Invertebrate Paleontology and Evolution. Fourth Edition (Clarkson, 1998)

Ill 11
9 780632 052387 >

Contents
XV
Preface
Macrofossils on CD- ROM xvi
Part One General Palaeontological Concepts 1
Principles of palaeontology 3
1.1 Introduction 3
1.2 Occurrence o( invertebrate fossils in Phanerozoic rocks 3
Hard-part pre ervation 3
Soft-part pre ervation 6
1.3 Divisions of invertebrate palaeontology 6
Taxonomy 8
The pecies concept 8
omenclanue and identification of tossil species 9
Ta.xonomic hierarchy 10
Use ofstatisticaJ method 12
Palaeobiol gy 13
Palaeoecology 13
Functional morphology, growth and fonn 19
tratigraphy 20
Lithostratigraphy 20
B io~tratigraph
20
hronostratigraph y 22
Bibltography 23
Books, treatises and symposia 23
Individual papers and other reference 25
2 Evolution and the fossil r ecord 26

2.1 Introduction 26
2.2 Darwin, the !>pecie and natural selection 26
lnheritan e and the o urce of vanation 28
Where doe. variation come from? 30
ignificance f allele 32
Murati n 33
pread of mutation through populations 34
I. elation and specie formation 35
Genetic drift: gene p ol 35
Molecular genetics and evolution 37
Ge ne regulation during development 38
2.3 Fossil record and m des of evolution 38
viii Contents
Microev lution 39 Part

All patnc speci.ltlon 4ll
Heterochrony ~I 4
Testing miL"rocvolutionary patterns 41 4
Analy~i~ of ea c ht tones 42 4
o-t·volution 43 4
Macrocvolu tion 4~
p ·cic~ selection 4~
rigins of lughcr taxa 45
R..ote~ of evolution, adaptive racliation. and extinction ~i
2.4 omperition and it eflects ~9

2.5 Utnl ary of palaeontological evolution theory su 4
Bibli graphy 51 4
Book:., treatise and sympmia SI 4
Individual papers and other rdcrcnccs 53
3 Major events in the history of life 55 4

3.1 Inrroduction ss 4
3.2 Pr karyotes and eubryotes 55
3 ..3 Earlie. t metazoan. Si
dia ara fauna: two viewpoint 5
The traditional view 5~
Mcdu. oid~ 59
Pennawla ·cans 5~
Annc.:Jjcb 6fl 5 <:;
Fossils of unknown affinities 61\ 5
V endozoan hypothesis 611 5
mall shelly fossih 61 s
Prccambrian trace fossils 63
Camcs of the Cambrian 'e:\1'losion of life' 64
Phy ico hemical factor · 64 5
B10logical f:'lctors 64 5
Biological evidence on metazoan rdation hip 65
3.4 MaJOT features of the Phanerozoic record 6
D1versification of invertebrate life 6~
Changes i11 pccics diversity and habitat 69
Problematic early Palacozoic fossil m
Marine evolutionary faunas 71
linutic ,md sea-level changl:s n
Extinctions 73
Pm~iblc L"att~es of mass extinctions 73
Earthbound mechanisms 74
Exrratcrn: trial mechani ms 74 B
ate rd vician (Ashbrillian) extinction event T
ate Dcvonian (Frasnian-Famennian) extmction event 75
Late P nnian extinction event 7
Late Tri:~ ic (Camian- Norian) extinction event 7(, 6 B
Cretaceous-Tertiary boundary extinction 77 (,
Bibliography 7
Book , trearises ::md symposia 7
Individual papers and other references 79
Contents ix
Part Two Invertebrate Phyla 83
4 Sponges 85
tl Phylum Porifcra: sponges 85
+.2 Clas'>Jficarion 87
+.3 Class Dcmospongea 88
Spicular demospo ngcs 88
'clcro pongcs 89
Chaetctids 90
rromatoporoids 90
phinctozoan 92
-1.-l Cla s Calcarea 93
-1.5 lass Hexactinellida 94
-1.6 !ncertae seru : Archaeocyatha 95
Soft part , organization and ecology 96
Distribution and stratigraphic use 97
n Geological importance of pongcs 98
+.8 ponge reefs 98
Spicular ~pongc rcc s 98
Calcareous sponge reefs 99
Bibliography 100
Books, trcati e and symposia !00
lndnidual papers and other rctcrcnces 100
5 Cnidarians 102
5. ! Introduction 102
- 1 MaJor characteristics and classes of Phylum Cnidaria
'·-
5.3 la s Hydrozoa
104
104
Order Hydroida 104
Order Hydrocorallina 105
i+ lass Scyphoz a 107
5.5 lass Anthozoa 107
ubclas Ceriantipatharia 107
ubcla. Octocorallia 108
ubclass Zoantharia: coraL !08
Order Rugosa 109
Order T abulata !24
Order Sderactinia 128
Coral reefs 132
Geological use of corals 135
ral as c lonies: the limits of zoantharian evolu tion 137
Minor orders 138
Bibliogr.1phy 139
Book , treatises and symposia 139
lndi\'idual papers and other references 140
6 Bryozoans 143
7 6.1 Introduction 143
3 6.2 Two examples ofliving bryozoans 143
~ B011'erba11kia 143
~ mifliua 145
X Contents
6. Classttic,m 11 1-1-7
6.4 Morpholo~:,ry ,md evolution ISO
6.5 col gy and dt'>tribution 154
~hallow-water bryozo.ltls IS..J.
Reef-dwelling bryoiOJns 155
Deep-water bryozoan~ 156
6.6 Str.ltlgraphical me 156
Bibliography 156
tlook. . tn:atm:s and . ymposia 156
Individual papns and other refere nces 157
7 Brachiop d 158
7.1 Introduction 158
7.2 Morphology 158
Subphylum Rhynclwnellitormca 159
Morphology ofthrt:'e genera 159
Preservation, study ,md classification of articulated brac hiopods 164
Major features ofbrachiopod morphology 167
Endopunctati n and pscudop unctat10n in shells 171 H
Subphylum Linh•uliformea 175 I!
Lin.~ula 176
Orhcr Lint,ruliformca 177
ubphylum Cr.miiformca 178
7.3 ntogcny 179 9
179 l)
7.4 Classttlcation !)
7.5 Evolutionary h tstory 183
18-1 l)
7.6 Ecology ;lnd dismbunon
Ecology of indiv1dual -;peC!cs 18-1
Eptf:mnal brachiopod, 185
:.nJof.mn.JI br,\chiopods IH5
B•·achiopod ass •mblagc and ·colllJlltulity' ecology 18!:1
rdovician pai.leocommunitie 18
ilu r:ian p::tlacocommunitics 18
Devoman brachiopod assemblag-es 191
Pernua n reef ,tssociJtions 191
Me <.woic brachiopod as~ociations 192
7.7 Fauna.) province~ 192 9
7.8 trarigraphic::tl use 193
Bibliography 19-t
Books. treatise<; and symposia 19-t
lnd1V1dual pJ~x·rs and m hcr rcft::rc tH:es 19-t
'I
8 Mollu c 197
8.1 r und.1mcntal org;.mization 197
H.2 la si ficarion 199
tU s
me .1spcct\ of ·hell morphology and growth 201
C01led shell morphology 101
Septation of the <;hell 203
~()J I)
8.4 Princjpal fossil groups
Class l3ivalvi.l 103
Contents xi
Cems/!lderma 203
Range of form and structure in bivalvcs 206
lassificanon 209
Evolutionary history 210
Funct10nal morphology and ecology 213
Ecology a11d palaeoecology 219
Stratigraphical ctse 220
la~. Rostroconchia 221
Cla~s Ga tropoda 222
Introduction ami anatomy 222
Classification 224
hell tn.tcture and morphology 224
Shell composition 226
[volution 226
Cla s ephalopoda 229
ubclass Nautiloidca 230
Subclass Anuuonoidea 23H
Subclass Colcoidea: dibranchiatc ccphalopods 251
'.5 Predation and the cvolmion of molluscs 255
Bibhography 256
9 Ech.inodenns 262
').1 Introduction 262
9.2 lamflcation 262
9.3 Subphylum chinozoa 263
Clas~ Eclun01dea 263
Morphology and life habits of three genera 263
Classification 269
Subclass Peti choechinoidea 270
ubclass tdaroidea 273
ubdas uech.inoidca and the morphological characters of euech.inoids 276
Evolution in echinoids 282
lass Holothuroidca 285
Cla Edrioasteroidea 286
9..t ubphylum futerozoa 288
ubclass A teroidea 288
ubdas omastcroidca 289
ubclass pluuroidca 290
Startl b beds 290
9.5 Subphylum Crinowa 291
Chtss rinoidca 291
Mam grmtps of crinoids 293
Palaeozoic crinoids 293
Mesoz01c to recent crinoid'l: articulatt.:s 297
Ecology of ctin id' 298
Fonnation of crinoiJallimcsconc' 300
<J.6 Subphylum Bla tozoa 301
Cla se Diplop\?rita and Rhornbifcra: cy~toids 301
xii Contents
tn1ctural characteristics 301 11

Pore ~tmcturc 301
lassification 3<!3
Ecology 3(!4
Class Bla tOtdca 304
D1ver. ity and function of hydro pire · 305
lassit1cation and evolution ofbla t ids 306
Ecology and di tribution ofbla toids 307
9.7 ubphylum Homalozoa, otherwise calcichordatcs 307
9.8 Evolution 311
Earliest echin denn and their radiations 311
Evolution of the tube [l et 313
Why pentamery? 313
onvergcnt evolution .and intermediacc forms 31-1
Bibliography 314
Bo k , treatises and symposia 314
10 G raptolites 31
10.1 rmcture 318
rdcr raptol idea 318
aetograptus chimaera 31.
D1plo.~raptus leptotheca 3211
rder Dcndroidca 320
De11drogmpt11s 320
Pre ervat:ion and rudy of graptolite 322
Ultrastructure and chemistry of graptolite periderm 324
10.2 Cla siftcation 326
I 0.3 Bioi gicaJ affimties 329
10.4 -voluti n 329
hape of graptolite rhabdosomes 329
Prmomal end in graptoloid 333
Thecal rructurc 335
ladia 337
tructurc of reti litids 33
10.5 How did graptoutcs live? 33 2
Passive drifting 339
A uta mobility 3~1
U c of model in interpreting the mode of life of grapt loids 34il

10.6 Fauna! province 343
10.7 StratigraphicaJ use 344
Bibliography 345
Individual paper and other references 346
11 Arthropod 348
11.1 lntroduction 3411
11._ la sification and general morphologr 34
01vcrsity of arthropod types 34M
Features of arthropod organization 349
Contents xiii
11.3 T riJobita 351
General morpholo~o,ry 351
Acaste dotlluiugiac 352
Detailed morphology of tJ.ilobites 354
Cuticle 355
Ccphalon 357
Glabella 357
Cephalic utures 357
Hypostomc 359
Eyes 360
Cephalic fringes 363
Enrollment and oaptarive stmcturc 365
Thorax 366
Pygidium 367
Appendage 367
Trilobite trac and trails 370
Walking movements in arthropods 372
Different kinds of trilobite trails 372
L1fe attitudes, habits and ecology 374
Ecdysi and ontogeny 377
lassification 3!)0
Evolution 382
General pattern of cvoluti n 3!;2
Microcvolution 383
Faunal provinces 386
tratigraphical use 3!:!7
11.4 Phylum hcliceraca 388
Clas Mero tor ata 388
Subclas Xipho ura 388
Subclas Eul)'pterida 392
11.5 Phylum m tacea 397
Bibliography 400
Books, treati es and symposia 400
Individual papers and other reference 400
12 Exceptional faunas; ichnology 406

12.1 Introduction 406
12.2 Burges Shale fauna .J-09
Arthropods 410
Lobopods 412
Other invertebrates 413
ignif1cance of the Burgess Shale faunas 415
Ecology 415
Geographical di ttibut1on 416
Diversity 416
Persistence 416
12.3 Upper ambrian of southern Sweden 418
12.4 Hunsrtickschicfer fauna 420
12.5 Mazon Creek fauna 422
12.6 Solnh fen lithographic limestone, Bavaria 424
xiv Contents
12.7 Ichnol gy 426

lassification of trace fossLI 426
Morphological and preservarional classification 426
Behavioural classification -1-26
Phylogenetic classification -1-29
Uses of ichnology 429
Sedimentary cnvironmen t 429
Srratigraphy -1-30
Fo sil behaviour -1-30
Bibliography 431
Books, treatise and sympo ia (exceptional faunas) 431
Individual paper\ and other references (exceptional fauna~) 431 The tir
Books, trc.ttises and sympo ia (ichnology) 434 li hed i
rndi vidual papers (ichnology) 434 new or
fact an<
Systemati c index 435 text for
year ot
General index 443 hapte
cion ha
chapter
a great(
of the i
for lonl
prove c
ana tom
Key
the tex
they ap
As\:
leagues
ance u
'olm
, dvice
should
taken c
thee c
Sheldo
J hn (
hdpcd
11llll1Cr'
abroad
I Wl
thank
for all
Preface
fhc tiN three cdit10n of tlus textbook were pub- given me at every stage. Had ecili nor undertaken
li'hl·J 111 1979, 19,6 and ll)92, and l trust that dill the vast job of rebuilding our teaching coiJectiom
n~w one. ncces. nated by so many advances both in and preparing new c urse matenals, I would never
fJct .md theory, \ ill retam lls fimctton as a course have had the tune to complete this book before the
te. t tor rudent of palaeontology from their second deadline.
\Car onward . 1 have made . ubstanual changes to To Raisin Morau of University allege, Galway
Chaptm 1-.~. 7, 8 and 1~. anJ the Trace Fmsils sec- I extend my grateful thank for her rwo beautiful
non has beln rramfem:d from haptcr I to the last paintings for the Part Title pages.
baptn. All rh other eh pters have been re vi. ed to I wish to thank also my editorial colleague ,
J grcJtcr or lesser extent. I have redrawn ab ut half Dr lan Francis and Jane Plowman, who have given
ufthc illusrranons: a singularly ongcnial occupation all pos ible assistance. From bn came the sugge non.
tor long \\1nter evenings, and I hope that these will wluch we have followed up, that the new edit1on
prove of Yaluc in helping student · to under. tand the should be coupled with a CD-ROM f Fossil
mJtomy .md tcnumologv of fo ~il invertebrates. Image . Tlus has been undertaken in conjunction
K y word~ appear in bold at their first mention in with the Natural Hist ry Museum, London, and to
tll( text. There is no sp<:cific index for them, but Nom1an Mcleod and Paul Taylor of that institution
dlC) Jppcar m the Gcm;ral Indc. ·. I owe a great deal. l am likewise grateful to David
1\s bctorc. I \ ish to thank all my friends, col- Hick and Eve Dainrith for theu meticulous copy-
lugut' ,Jnd f:UniJy who have given me such assi t- editing and proofreadmg, respectively .
.ttll't' m prLpanng this new cdnion. Derek Briggs, On a wet November day, over 50 years ago, I WJS
Ctlllll rutton Jnd Rachel Wood gave excellent taken into a museum in my native Newcastle-upon-
JUVKI.' at the begtnning of the project as to how I Tync, to escape fr m the rain. here, in a dusty glass
~twuld proceed w1th the ourth edit1o11, and I have case were two giant ammonites (probably Titanites
t.ikt'll on board m t of th ir suggestions. As alw;~y~. gigallletts), and 1 was given t under rand that they
thi.'\C collt·agucs, together with Oav1d Harper, Peter had lived million of years ag . hey excited J fasct-
hddllll, usan Rigb) . Liz Harper, I ick Jdferies, nation which has continued until now. and 1 am snll
John Ccpc, Alan Owcn and vanou others have glad to escape to tht: hills on a fi-esh summer mom-
lwl~wd llll' Jt all stages, dUd so have many other , too ing, to search for fossils, and then t bring them back
mlmernm to mcntJon, m my wn country and to the laboratory for study. 1 hope that thts book
1broad ..111d my thanks are due to all. will be f value to any student wishing to explore
I 1\0uld hke, above all, to record my heartfelt something of the richness and diversity of ancient
1h.111l' Ill my m1c and ~talwan friend ccih.1 Taylor, life, and of the methods available for its scientific
li.lt .Ill tlw 1upport and helpful suggestions she has study. ff o, I wiJJ have achieved what I set out t do .
Euan larks n
Edmburgh
Macrofossi Is on
CD-ROM
The Natural History Museum London and Or E.N .K. larkson have collaborated on the development of
rhi important new initiative in palaeontology teaching.
t\lacrofossils 011 CD-Ro111 is ,1 combined image lib rary Jnd database, containing rec rds for 1000 key fossil
genera. Each record contains a set of images, and information on stratigraphic range, hard-part mineralogy.
palaeoecology, palaeobiogeograph , etc. Tbe images have been captured using the Natural History
Mu cum's high resolution PALAEOVI ION digital imaging system, and the data and image reside in the
CompuStrat database mJnoger.
Thi ~y~tcm allows the user great Aexibility in displaying data. For example, users can ~imply bro\ se from
record to record; pull up fossil IT m a taxonomic index; or select and sort records by geographic range, life
h:Jbit, stratigrapluc range, or by a variety of other criteria. Range charts and palaeobiogeographic maps are
given where appropriate. and the user can print records, or export them into other applications. Within
each f the major groups there • re a number of labelled images and diagrams to allow the user to become
£1nliliar with key morphological tenl1S .
.\.laaoj(JSSils 011 CD-R M may be used a~ an adJunct to ll/llertebratc Palacoutolllgy, or a a tandalone prod-
uct. Taxa have been selected for their relevance to Earth ciencc teaching worldwide.
PC or MA , minuuum 6Mb RAM, D-R M dnve, 640 X 480 colour monitor.
For furrher infomlation and ordering details, please contact: ian.francis@blacbci .co.uk
PART ONE
General
Palaeontologica
Concepts
Once
Son
bl ing
p.trticj
the S
from
.tccrctl
Of
E.1rth,
thou •
been!
Oil j UJ
ing tt!
withit
.1n.: ret
green
old,
These
the se
.ltth
cyano
instnn
gave
ously
bee am
Ot~
~arth
rpro
when
of CV
m ann(
·volvi
in gee
Arnioceros cf. hortmanni (Oppel), on assemblage of immature ammonites from the lower Jurossic of Block Yen, Chormouth, England know1
(lower Sinemurion). These specimens were probably catastrophically buried, since the soft parts must hove been in place when they
died preventing sediment penetrating the chambers. Pointing by Raisin Moron; original specimen in the Jomes Mitchell Museum, Ncedl
University College, Galway, Ireland. bckint
Principles of
palaeontology
no record of terrestrial ,mirnal until mu h later. In
tem1s of our under tanding of the hist ry of ]jfi·,
perhaps the most significant of all events wok place
Onu: uplm .t cimc .. abOtl t 543 Ma ago at about the beginning the
~mm: ~600 million ye:m Jgo the Earth came into ambria n Period, for at this stage there \ as a sud-
b~mg, probably omtmg fi·om a condensmg disc of den proliferation of different kinds of marine inver-
pmid~ . dust and gas, \\ luch slowly rotated round tebrate . During thi cntical period the princip:tl
tht Sun l trger partKk,, or pl.metismals, formed invertebrate groups were est bli hed , and they then
tn101 d11 nebular disc, dnd a~ rhese coll.Jded they dtverslfied and expanded. S me of these organisms
J tTdl'U, en:ntually f(mmng the planers. acquired hard shells and were capable of being fo -
Of .Ul thl· nine planets tn the alar ystem only silized, a11d only b ecause of thi. can there be any
Earth, ,11 tar a 1 known, upporrs advanced ltte, chance of understanding the history of invertebrate
though at the timt· nf wnctng much tntert:St ha~ life .
bcw generated by the discowry f organic material he straufied sedimentary rock laid down ~ince
onJupirer's satellite uropa. it 1 , however. a tnk- the early ambrian, and built up through m the
mg !Jet rlut life on Earth beg,ln very early indeed, whole of Phanerozoic time, arc d1 tinguished by a
111rlun the tirst 30% of th plJnet's history . There t;ch heritage of the fo il remains of the inverte-
ar~ rt·rnatm of simple organim1s (bacteria and 'blue- brate that evolved thr ugh successive historical
~rr~m J]gae , or cyanoh. crena) in rocks 34UO M a penod ; their study i the d main of invertebrate
tM. 10 lifr presumably origmated before then. pabeontology and rhe subject of this book.
The->e 1imple torm. of life 1Le111 to ha\'c dominated
the llTill' fur the next 2000 Ma, and evolution
Jt th.u ttnte was very slow. Neverthele s, the
l\'.mob.tctcna and photmynthetic bacteria were
m1tnmltrlt.1l in changmg the environm ent, for they
~.trc off O\)gen inco an atmosphere that \ as previ-
Ollll) dl.'voiJ of it, ~o thar animal ltte eventually
b,umc possible. Hard-part preservation
On!) when s me of the early hvmg bemgs of thi
bnh had reached a htgh level of physiological and Fo si) invertebrates occur in many kinds of ed.imen-
rl'producnv~ organizabon (and most parbcularly tary rock deposited in the seas during rhe
\\hut IC\ual reproduction onginatcd) wa~ the rate Phanerozoic. They m ~1y be very abundant in
11f crolutwnary change aceeler:tted, :1nd with it all lim estones, ~bales, siltstone and mudstones but 011
lllJillll'r of new pos~ibilit1cs were opened up to the wlwlc are not common m sandsrone .
nolving lite. This wa. not until comparatively late Sedimcnt. ry ironsrones 111Jy have rich fossil
111 geological history. ;Jnd there ::tre no fossil animals remain . Occasionally they are found in some coarse
nown from sed.imcnts 0ldcr than about 700 Ma. rocks mch as greywacke and even conglomerates.
N~cdl s to say. these an: ,ill invertebrate anilnaJs The state of preservation of fmsiJs varies greatly,
I l1n~ bad.b nes. AU of them are ntJrine; there is depending n the tmcture aJ1d composition of rhc
4 Principles of palaeontology
original shdl. the nature .mJ grJm stzc of the cnc1m- Hoor during these ero">ion periods would probably
mg sediment, the clwnuc:tl conditions .lt the time of be trJ.nsport d or de. ttoyed- another linutatlon on
~t:dimentarion, .md the subsequent proce\ses of dia- the adequacy f the fos. il record .
genesis (chenuc,1l and physical Lh:mges) tJbng On the other hand, some manne invertebrate\
plan· 111 the ro k after deposition. found in cerrain rock types have been prc~erved
The study of tlH" proces~es le.tdmg to fi.1ssilization abundantly :md in t:Xt.]Ui~ire det,til, so th.lt it 1s possi-
is known .1s taphonomy. In mo. t ea es only the ble ro infer mu h , bout their biolo!:.'Y from their
ll.lrd pam of ro~ 11 .t llli11Jl~ Jre pre<;ervt:d, Jnd for remain . . Many the best-prc~crvcd fo.,sils .:ome
these eo be toss1hzed. r.lptd bmi I is nonn;l]ly J pre- from limc~ton c' nr fi· m ilty ediment' with a lugh
requt tte . rhe sofr-b died element Ill the faunJ. . calcareous content. ln rhe~e (Fig. 1.1) the ongmal
Jnd thme Conm Wtth thin org.uuc sheJl , did !lOt c.1lc,treom <>hclls may be retained in rhe filSstl state
nonnally survive dtagrncst~ ,md hence have left little \ ith rebtivcly little aheration, depending upon the
or no evidettce ofchetr (>.Jstence other rh.m records chemic:1l conditions within the ·eJiment .1t the tum
of thetr activity tn the tonn of trace fm,ik Wh t \ c..· of do:po ttion and <tfi:cr.
Gill see in the rocks 1s thcrdore only .1 narrow band A sediment often co n i'ts of c mponent~ derived
in J whole <ipcccnnn of the organisms that were f1·om v:1riou~ environments, and \ hen all of these .
once ltving; only vety r.udy lMve there been tound including dcc~ying organism~ , dead . hell and scdt-
beds contammg some or .dl of the ~ott-bodted ele- mentaty p:trti le , are rl1rown together tht: chemtca l
ment~ lll the tauna .1~ well. ' rhe~e JT 111ll11C11SCI)' sig- balance i un t. blc. The ,cdiment vvill b~: in cht:mt-
mticant for palaeontology. cal equilibrium only after di:Jgcneric physicochemi-
Tht oldc~t such f;:mna 1\ of 1.1te Precambnan age, cal J!terati ns h:wc taken place. Tlwsc may involw
some ul5 Ma old. and ts om onlv record of animJI recty~t.'lllizati on and the growth of new tnin~:ral
life before the Cambri.tn. An cher ~uch 'wmd w' is (authigene i ) a~ wd l as cementation and com-
known u1 Middle 'ambn,111 rock from Briri h p.Ktion of the r ck (lithmcation), and dunng ,my
lumbia. where tn additt n to the normally one of these proce es the ossil~ may be altered or
expected tttl bttes and brJch1opods tbcre ts a brreat de~troyed. hdls that arc originally of cJJcite pre-
range of oft-bodied and th.in-she!Jed animal serve best: arJgonite i~ ,1 lc~s st:1blc fonn of calcium Figure I
sponges, wonm, jdlytisb, 'mail shnmp-ltke creJ- ca rbonarc ~ccrcred by certain living organisms (e.g. shell wo5
inollyof
tun~s .md .munals of quite unknow11 nffimtie~ - cor;~ls) .md io; often n::cryst;:tllizcd to c:Ucite during genetic:. c
which .tre the only trace of a diverw fauna which diagcncsis or di,solvcd away completely. shell pret
\ oultl otherwise be quite unknown (Ciupter 12). Calcarcom skeletons preserved in more sandy or core; (h)
(j) irons!<
Then: are o;imibr 'windows' at other kYds in the silty sediment~ nuy dissolve 3fter the sediment h<t!'!
geologicJI culllmn. likewise ilhuninaring. hardened or during weathering of the rock long
Th e fcmil rerord ts. J .1 guide to the evolution of Jfter its induration. Moulds (often tnt~called casts)
,mctcllt litl:. tmql.leStl n,tblv limttcd, patchy ;md of the external .llld internal surfKe~ oftht· fossilmav rh ori!
incomplete . Usuall nly the hard-~helled clemem:s be left, and if the ,ediment i ttne enough the details t:Xtra r
m the b10t.1 (.tpart fn m trace tossLis) Jre preserved, the~e show may be very good. Some methods for while
and the foml .1~ embl.lge present in the rock may the study of ~uch mould~ arc described in 'icction t•xpen
have bet:n trJnsportetl ~ome Liistanee, Jbraded, dam- 7.2, with reference to br;Khiopod~. If a tos,J 0111Ctli
aged and nlLxed with elements of other fauna.~. Even t·nclmcs an oribrinally hollow pan\ JS tor instance served
if thick-. helled anim:tl- w~:re onginally present in a between the pair of ~ hell of a bivalve or brachi - h . dr g
fauna , rhey may not be preserveJ. m ~andy ~edi pod, this space m ay either be left empty or become (h '+)
ment\ in \ •hteh the orculanng waters ,tre ..tcidie, for filled wirh sediment. In the latter c ~c d ~edunen t core t<
instance, calcareous shells n1.1y dts~olve wtthin a few core is pre5crvcd, which ome~ out mtact when the withi.n
years before the sedunent 1S c mp.u:ted inro rock. rock is cracked open . hi, b~:an upon its 5uriace .m \ th se
ince the sea floor 1s not .1lway~ ..t reg1 n of contmu- internal mouJd f tht: fi ;~il sheU, whcrca5 exter- ·ngla n
ous sedunent depoo;mon, many , pparently continu- nal moulds arc left in the cavtty !Tom which it ou Ill\
ous edimentary sequence~ concam numerom came. In 1-:trc circumsr:mces the core or the ~hell, or ~picule
tn<ill-scale breaks (diastems) representmg penods both . may be rephccd by an entirely different nun- co nditi
of wumo\ving and nmi n . Any shells on the seJ era!, as happem in fmsilo; pre,crved in iroll\tone~. If genera
Occurrence of invertebrate fossils in Phanerozoic rocks 5
lil
/ (f)
Figvre l 1 Possible processes of fossi lization of a bivalve shell: (o) original shell, buried in mud (left) or carbonate (ri~ht); (b) fu e
sllell was calcite, wos buried in c carbonate sediment and was preserved intact other than as o small cry~tollized patch; (c) shell orig-
molly ol orogooite, now recrystallized to calcite which destroys the fine stnucture; (d) original calcite shell retained surrounding o die-
genetic core of silica; (e) o silica rim growing on the outside of the shell; (f) tectonic distortion of o shell preserved in mudstone; (g)
shell preserved in mud with original shell material leached away, leaving on external and on internal mould, surrounding a mud stone
core, (h) a calcareous concretion growing round the shell and inside Iif the original cavity was empty), with patches of pyrite in places;
!H lfOOStone replacement of COre a nd port of shell.
th~ nn"rinJl 'paces m the shell arc impregnated with the si lica incre:.r es markedly. and the ilica o
exm minerals, it 1s SJid to be permineralized , released will travel thro ugh the rock and precipitate
\\ltik th 1-'fOWth of \t'CUnd.U)' minerals JC the wherever the pH i lower. The imide of a sea urchin
expmsc of the shell 1s r.ep1acerncnt. ores may decaying under cli£fercnt conditiom would trap ju t
10nwtm1c1 be of pyrite. Graptolitc..tre often pre- ~uch an internal microcnviionment, within which
lcrml like tht, anaeorbic decay having relea5ed the ilica cou ld precipitate a~ a gel. Such siliceous
11dmg~n sulph.tde, wluc:h reacred with ferrou~ cores retain excellent tcaturc., prescrv111g the internal
I~· ) Jl\m in rh water to allow Jn mtcmal pyrite m rp hology of the ~ h ell. n the o ther hand, ilica
lnT<.' m funn . ometime~ a core of 1lica is found llhl)' re place calcite as a very thin shell over the sur-
within ,IJl unaltered calnte shell. Th1~ has happened face of a fossil as a result of , omc complex surface
Wtth \OI1lt' ut the retaceous sea urchms of ~ourhem reactions. T he e siliceous cru ts may retain a very
En~and . fh cy !wed m or n a .,ed!mt·nt of calcare- detailed expression of the surface of the fossil and,
nu mud along with many sponge,, wh1ch ecreted since they can be freed from chc rock by dis olving
'PIL"Ub of b10geruc s1hca as a skeleton. In alkaline the limestone with hydrochloric acid, individual
nmhtiom (above pH 9), which may sometimes be . mall to sils preserved in this w::ty can be studJ~d in
~·nmtrd dtmng ba tt"rial decay, the solubility of three dimension . Some f the most exquisite of all
trilobite~ and brachwpnd~ are known from material n:ma111s of omc part, usual!) a shell of ~keleton, of a
such as thts. oncc-Li ing org:mism; (2} trace fossils , -.: hi ·h ar..:
A rdativel) uncommon bm L ·q m~nc mode of trac~. trails, burrows or other cv tdencc of the activ-
prcscrvauon ts pho phatization . Sometimes the ity of an animal f former tmlt"S - omcttmes thest:
cxtcma.l skeleton, speoally 0f thm organtc-shdlcd arc the only guide to the former presence of sofi:-
a.nimaJs, may be replaced or ovn~;,rrown b> a thin bodicd animals in a particular environment; (3)
sheet of pho ·ph.He, or tht• bttcr m,ty reinforce :m chemical fossils . relics of biogenic orgamc com-
onginally phmpharK shell. In tht• former . ttuanon pounds which may be detected geochcmically in
the external fom1 of the body j, prcci,dy rcplic<ttcd. the r ck~.
Altemauvely .1 phosphat!L tilling nf thl interior of At the beart of invertebrate palaeomolob'Y ~tanru
the ~hell may form a corL', ptckmg out imcm.1l t~U · onomy; the cla sificatJOn of fo~sil and modern
stntctures in rem:ukabk detail. Such prc~ervation is .tlumals into orden:d and natural gr upmgs. Thc'c
probably .lSsociated \\1th b.ICtcn.ll activity directly groupings, known a taxa , mu t be n.:amcd anc
after the death of the am mal. M.my ~mall ambrian arr,mgcd in a hierarch;al system in which their rda-
fossils have been pn::~erved by pbmphatiution tion hips are made cle:tr, and as tjr ;t~ po siblc mmt
( 'hapters 3 J.nd 12), but much larger fossils may be be ccn in evoluti nary pcrspectJvc.
pn:served also, tor t·xample LTU~tal·eans \Vith a Evolution theory is compounded of various clis-
ftuoupatite tntillin~ and 'vvith Jll their delicate ciplme - pure biology, comp:tranve .111atomy,
appendages jman embryology, genetics <tnd population biology - bur
Fossils :~re often found in oncretions: cakan.'- it i only the palaeontological aspect th.lt allow~ the
ous or sili eou~ nun~s tomll'd <~round the fmsil predicti ns of evoluci nary <cicnce to be te ted
shorth after Jts death Jnd hnnal. Concn:riom fonn .tgain t the background of geo logical nmc, pcnmrs
under certam condition onlv, where J deltc,Jtc the tracmg of evolving lineages and illustrates ~ome
chenuc.1l b.thnc~: L xtsts between rh~: \ Jtl'r and ~edi of the patterns of evolution that ,lCtu,tlly have
ment, by proc es~c~ as ver nor fully understood. OCCUITed.
The rates at which aJllmals have e\'olvcd have
varied through time. bur mmt .mimal type'
Soft--part preservation (species) h;lve had a geologicJI life of only a few
million year~ .. ome of these t'Vl1lvcd rJpidly, ~uch a~ Figure 1.2
In very rare circumstances ·oft-bodied organisms
the anunomtes, others very slowly ( h, ptcr ~). A
can be pre. cn•ed J.~ fo sils, :Jnd the-.e provide other-
rock succe~ ion of marine ~edimenr~ bmlt up o L'T
\V1 e unobtainable evidence ot the divcr~ity of met:l-
many millions of ye.m; may therefore have ~cvcral ( t .~·. MlU
zo:uls living at patticular pt:nod~; thi~ is dtscus~cd m 1 ju t a
ossil species occurring in a particular eyucncc. each
Chapter ~~ - ~.:e ruin 1
. pt:ctes confi.u.cd to one part of the uc e sion onl)
and representing the time when rh:1r pecie wJs p 1. 1:!, thl
living. Heremlies the oldest and mosr general appli- ph IS lll
ca tion of invertebrate paJacomolot,ry: biostratig- Alcbo1
raphy. Using the sequence of fossil finlll ,JS, the dm tplin,
gt:ological column has been divided up into a 'erics ~ Jl. cnnt
lnvenebratc pJ.bcontolo~;.ry is normally scudjed as .1 of major geological time units (penod ). cJch of ll.:n11cd
ubdivmon of !,':l'ology, .ts ir i~ withm Earth ~cience which is further divided into a hH'r;Jrch ' of small growth 1
tlut m gn.·ate\t apphcatitm~ lie. It cm .iliu b~: seen as unit . The whole basis for tl1.i histonol chronology (H lll h,lS
a bioi gical \Ubjcct. but one Llut h,1s rhc unique per- i~ the documented sequence of fo~sils in the rocks. JJy~ 111 I
spective of geolo!.,rical time. W1tlun tht: domain of But different kinds of fossils llJve ditTerenr mati- lm h.t~
invertebrate palac ntology there an.· a number of graphicaJ value~, and certain parrs of the geologiCJI th~ ,)0\
intendated topic (Fig. 1.2), all of which have a record are more closely subdivtded than otheN Snln'
bearing on the olhers ,md wh1ch also link up with ome 'ab~olute · ages based on r:Jdwmerric d:1ting IJJVl .ll'vl
other . cienccs. h:1vc been fixed .1t particular pomts to thi~ rcbtivc I pu:ts I
Three main c.ncgone~ of t\.1,sih may be diStin- ralc, anti these provide a framework for under- tlllcil
guished: ( 1) body fossils , in Pthcr word-. the acrual standing th geological recurd in tcnns of real time \.Vhilll I
Divisions of invertebrate palaeontology 7
growth
and form
Ftgurc 1.2 The various subdisciplines of palaeontology.
(1.e. known pcnod! o million of year~) rather than 1 YSOs, is concerned with tht' relationshi ps of fo si!
J JUst a purely relative scJ.!e. This i · only po ible at animals to tht'ir environme t, both as ind1vidu I
t~nJill honzon , however, and for practical pur- (autecolog y) and in the fauna! couununiti cs in
f u' •, tlw g~·ologid time cale based on biostratigra - which they occur naturalJy; the httcr is sometime~
ph 1 unsurpa.scd for Pha.nerozmc ~edimcnc known as synecolog y.
Although tratigraph 1s the basts of the pnmary in cc the oft parts of£; ssil animals are not nor-
d1 npluw of geochrono logy. a small facer of mally prc crved, but only their hard shells there are
Jhwntolo!,'lCal stud\ ha~ a bearing on what may be relatively few ways in which their biology and life
tcnn~d gcochrono metry'. By counting daily habits ca n be understood . tudies in functiona l
;r1wth nng-. in cxtinc:t coral and bivalvcs, infomla- morpholo gy , however, which deal with the imcr-
rum Ju, h~t:n obtained bearing upon the number of pretation of the biol ogy of fo silized skeletons or
hv~ 111 tlw lunar month and year in ancient times. tmctures in tetms of their original function, have
Tlm hJs hdpcd to cnnfim1 geophysica l estimates on been ~ucccssfi..JlJy attempted with many kind~ of fo~
thL· 1l\l\\1Hg llfthc Earth's r ration (Chapter 5). sil , rcstncted in scope though these endeavour s may
Smcc .. rraographical application . of palaeontol ogy necessarily be. Ichnology is the study of trace fos-
hII'C .U\IJYS been <;o important, the more biological sils: the tracks, surface trails, burrows and bori11~
l'PCd> of p;ll contology were relatively neglected made by once-livin g animal and preserved iu cdi-
uunl rompJrativcly recc11tly. PaJaeoeco logy, ments. This topic has proved valuab le both in
wlud1 h dt:vclopcd parttcularly s1ncc tbe early under ·tanding the bch viour of the animals that
lived when tht· ~e hmcnr wa~ bemg deposited and m These word~ ~hould make entirely clear the fun- ti.Jome(
interpreting Lh<.: contempt rancom enviroruncnt. damentaJ n:nurc of ra.x:onomy. For. as lu~ often beer
Finally, ir is only by the intq,.-ration of ra,xonomic s~1id, to identify a ~ossil coJTectly t~ the fir-;t step . .tnc
data on locaJ faunas that the global di.~tribution of indeed the key. to findmg out further mlonnJtlon
marine inverrcbr:J.te~ through time CJJ1 be dun- abot1t it. Sound l1sS1tl :Jtion and nomenclature hr
dated uch '>tudles of palaeobiogeograpby (or at th e root of Jll biolof:,rical and pa.laeontolob'1C.1
palaeozoogeograpby in the chc of animals) can work; withom d1cm no cnherem :llld ordered
be u cd in onjunccion with g<.:ophy~icaJ daca in system of dJta ~roragc and rctricv~ll L~ possible
undcr.;unding the: tunner relative pos1tiom : m d Ta.xonomy. or systematics as it i~ <.omehmc
movements of conrincnr.li masse>. known, is the ~cience of cb sific:Hion or organi1m1
All of these a'p 'cts of palaconrology :~re intcJTe- · It i~ the oldest of all the blOlogtcal discipline~ ..u1d
lated. and an ad nee in one 111:1) h:tvc ,1 beanng the prinnpl s outlined by Carl Gustav Lmn, eu
upon any orhcr. hu5 J particular study in func- ( 1707- 1778) m hi pioneenng y~lt'HIIJ naturae ar<
rionaJ morpholo!:,ry nlJy b'1VC infonnation on st11l m us t d:Jy, thongh greatly modified .tnd
palacoccology ,md pmsihly ~ome t(:edback to taxon- cxtcndt:d.
omy a<. well . Liknvise, recent refinements in t:txo- ~ rt.ltll \\
nonuc prJrtKC have en ,tbkd the development of a The species concept piu lngy
much more preci (' '>trJtlgraphy. The fi.md<mlCntaJ unit of taxonomy IS the ~pcCies rh dduu
hemical c mpound' or bmlogic.ll origin can Animal species (e.g. Sylvester-Bradley, li.J56) lrC
now be recovered from ;lncu.' nt rocks ~md fom1 rhc group. of mdiVlduais that generally look hke ~:ach
b.tsl of biom lecular palaeontology Such to tl other and can imcrbrecd to produce otf~pt-ing ofthc
mok ule~ may hdp to di.tgnc '>c whtch org:tmsm~ ~amc kmd. They cannot mtcrbrced wtth othc
they come fron1 .md tht·tr brcJkdm ·n p,tthw.tys may ~pecic~. ince ir i~ reproductive isolation alone rh~
say nmcthmg ,l bllur thl· t'll\'1 ronmcnr. Molecular defines species it i. 011ly really po sible to dmmgw1l1
phylogcnctirs b.t.;ed upnn protcm scquen mg may clmdy related -;pecies if their breeding h,1bH' art
'how how tar two or more related organtsms have known. Of :~11 the described 'species' of livtng am-
d1verged from ,1 comm m .mcc~tor, and to ~omc mals. however, only about a ~ixlh arc 'g od', o
extent rhe av;nl,lble rcchmqm'' on be applied to the properly ddined, species. lnfo1111Jtion 11pon rh
rcu~nt t(1~stl re ord. lmnlllnological-detcmlinant reproductive preferences of the other tive-srxths (11
tcchmqul'~ c:1n be used to detect proteins and poly- allnarur.llly occurring animal populations is JUSt m
ard1Jndcs m fim!l shell' bur, fiJr the moment, only documented .
~hell younger thJn 2 Ma llJYe proved amenable to The ditlcrcntianon of mosr liVIng and .!11 fo 1 .lll d ~,[f;.ltJ
analy'li . here ,m· also promising developments also specie~ th crdore has ro be ba ed upon other anc I t:l 1ll
in p.1laeobt chl'lnt,try .md r~,mic gc ·hemjsrry technically le. vaud criteria.
which arc apphcabl ' tn the fossil record, though Of these by fur the most important, especially u
tht:se are beyond tht' ~mr( of tlm hook . pabconrology, i morphology , the ~cicncc o
f1 rm, since most natural >pecie~ tend ro be com- !-\lllSh th'
pmcd of indiv:idual of similar enough cxterm 0\\IJ ht,H:
Taxonomy appearance to be identifiable a~ of the same k.md \'tdttal to
ni .. tinguishing species· of living anima.ls by morpho \llllil.n in
Taxonomy is o "lcn undavalued .h .1 glorified logJcJI criteria alone is not w1th ut hazards, espc· erh. psI
turm nffiling - with eJch ~peci~s in it~ tulder, like cially where the ·pecies in que~non are similar atl1 .1 p.trticul
.1 stamp 111 m pn:,.:ribed place m .m Jlbum: but do el related. upplementary mtonmtion, uch .r trl.l b. l"
taxonomy i' .1 fund.tmenwl Jnd dyn.ulllc 'i ·io:nct•, the an:~Jy is of species-speCltlc protems, may be o tlw di tin
cll·dtcatcd to c.· plnnng tht• r,tU\l'~ llf rclatiumhip' help in ~ome ea es where there i good reason fo tit· ri k '
and simil,mrie~ among orgJnt m,. Ja,~tflc<!tium it to be ought (e.g. for disea~e- arrying 111~e ts). FO' ('1'1 VISIOO
Jl'l theoncs ;tb~1ur the ba'i' of 1utural order, not the rc~t some degree of subjectivity in taxonomy h. be tduHif
dull catalogue~ Cl)ltlpikd ~.mly to avoid cha ~- to be .lcccptcd, though this can be minimized log~ :1! nl
"17n· be· t 111011c~~mph:. arc ll'mks l![gf'llitH ... enough morphological criteria arc u~ed. ll'lho t.;,JI
(SJ. Could, 1990) .t ~l rtbl·d
Nomenclature and identification of fossil species comp:.tring th specie~ point by point. Some f the
In the forma] nomenclature t any spcncs. living or species may pr vc to be identic, 1 with already
·O\\U, taxononusts tallow the b10logtcal system of described species, or show only minor wtri,ttion of a
Lumacus, whereby each spec1es IS defined by two kmd that would be expected in .1 I al v,triant
u;uncs: the generic aud pe ifi (or trivial) tl.lllles. within the same species. Other specie~ in the fauna
For example, all cats. l::Jrge ~md mall ..1re related, may be new, and ifs a full technical dc~cription
and on~? parrirular group lu been pl.lced in the wtth illustrations mmt be prepared for ea h nev
genus Felis . Of the v.mou. pee1cs of h'lis the spe- pecies, which ~hould be.: published in a pal:Jconco-
cific names F. Ct1friS, r. ,11'(1 and F. pnrdw tomul!y logtcal journ:~l or monogr.tph. Thic description is
refer to the d mesne cat, ilon and le p.ud, respec- based up n type specimens, wh1 h are :~!ways
~vdy ln full c:t.xonorruc nom~?nclature the author' thereafter kept in a museum or research imtitutc.
name ,md the d.1t..: of publtcation arc given after the Usually one of the e, the holotype, is clectc.:cl as
.pcctes, e.g. Fells mills (LiJlnaem t 77~) (see below the reference specimen :md fully illustrated: com-
tOt further dJscu~sion). p.trative detail may be added &om other specnncns
In pabeontology it cm never be known for called the paratypes. There arc various other kinds
certain whether a populacion \VHh a parricular mor- of type specimens; for example, a neotype may be
phology w~s reproductivdy isobted or not. Hence erected when :1 holotype has been lost, or whl'n a
the definition of spene~ 111 pJ.Iaeontology. J'i in most species is being redemibed in fi.Illcr and more up-
h1~ug specuuen , mu t be ba ·ed :tlmmt entirely on to-date term · when n type pecimen has previously
morpholog~c.tl criten.1 Moreover, only the hard been designated.
pJm of the fosstl .umnal are pre ervcd, and much A new genus will be designated a gen. m v. by it~
11sdttl data h ~~ vam. hed A carefUl examination and author, this following the generic name, a ne
d (Umentation of .11l the Jn:ltmnical feature~ of the species a sp. nov. a.nd a new sub pecics a~ ssp. no\',
itl)Sll has to be the matn gt11de 111 cstabli~hmg that The new specie must be named and allocated to
onnp~ries is different from a rebted specie~. Tn rare an existing genus, or if there is no de cribed genus
ca1cs tht can be supplemented by ,t comparison of to which it pertains then a uew g nus must al o be
the chenustry of the '>hcU. as has proved especi,tlly erected. To appreciate the method let u con. ider
lh~tltl m rhe erectmn of rugher t;vonomic c. te- the tollowing historical ulc. Tn the early niJleteenth
f gones. Wirhm any interbreeding popubt1on there is century brachiopods were poorly known :md few
U1U31ly quite a spread of morphologiCJI variation. di~tinct genera had been erected. ne of these was
On a broader scall" there m.ly be b rh ge graphicaJ the l.iving Terebrawla, named by l .F. Mi.illl'r in
l .md strat1graphic vamttons, and all these mu t be 1776. When E.F. von chlotheim fim studied
i mefully documented 1f the specie LS to be tdeally Devonian fo '>ils !Tom orth Germany in 11LO he
·ublished. uch studte may be very igmficant m recognized that ~ome of the shell! were bra-
1 rvolution,try palaeontology. chiopods, and he described one of d1c most abun-
f When a pal.u~ontolog~.St 1 ancmpcing to dtstin- d:~nt fonn a the new . peci Te,-ehratula sMrimtlaws.
gut h the ~pecies 111 a nn ly d1scovered t:mn:~. ay of By I H30, however, much more was kno\\'ll about
J ioml brachmpods, she or he has to separate rhe mdi- br.chiopods, and G. Fischer de Waldheim proposed
\1du.ll fo sils our into groups of morpholobricaUy a new genus for this species. so that it became cor-
1\llltbt inruvidu:tl~. There mJy, to cake an example, rectly de ignated Clto11etes sarciuulatus (Schlorhcim).
perhaps be e1ght uch gr ups, eJch distinguished by This is the 'type "pecics' of lwnetes, :1 >veil-known
d parncular set of chant ters. ome of the. e groups iluro-1 evonian genus of the Cla s trophomenat,t.
nuv br dearly distinct from one another; in others Nme that where a species was origmally de cribed
rh diShnctlon may be con tde-rably le s, mere sing under a different ge neric name. the oribrinal uthor\
tht mk of greater ubJeCtJviry. These group. arc name is quoted in p:~renthc , e". In 1917 F.R.
provt tonally con idered as . pecte~. whtch must then owper Reed, then working on Ordov1 'J:t.n and
LS b~ identified. fhis i · don~: by consulting pabeont - ilutian brachiopod~ ofthe Girvan distt-ict, cotland,
if lobi'JLJl monographs or paper'l contam.ing detailed recognized many new species. One of these had
rcLhmnl dcscnptions and illustrations of prev10usly similaritie. in morphology to lulllefes, hut it was
dlscnbcd brachiopod faunas of sumlar age' md sullinently different to be regarded as a ~pccies of 3
new subgenu ; thts is written C/wrzctes (Eodw11etes) There may be various subdivisiom c f these cate- logicetl ( ,
advcna Reed \917 . When m 1':1:2!:1 the ta.xonomic goric~. e.g. superf::unili ' . , suborders, etc., and in ccr- '' hach the
problems of Clwm•tcs ::utd similar forms wen.! tam groups there is evcn a case for erecting phylo en,
addres ed by O .T . Joncs, the ne\V Supertanuly 'superphyla' . There ar only <lbout 30 phyla in the lll'til e\l'l:
Pie t•nnbonjtacea was erected to Jccomrnod.atc animal kingdom, . nd only about J dozen of the>e, 111L:JI1t thl:
11dJ 1l'l!c1 ,md m:lny otbcr related bra hi pods. At a e.g. Mollu ea and Bra hiopoda, leJve any ossil whit h nil'
later stage Eocho11e1es wJs cle\"<lted to the rank of a rcmau1s. an Lcn11'
full genu~. In the most recent rrearmcnt, I .A .T. In ta.-xonolll higher taxa an.' u ually di. tinguishcd I'OIIr.e grJ
Harper described a large fauna fi-om the irvan by their suffLx (i.e. -ea, -a, etc.). A an example , the oakr f ~
di rri -r f ' c tland, and wnlun thts he recogmzed following documents the: das tfication f the . rrJplltcal d
1'-.vo subspecies of f . adi'CI"'· of onH.:what ditrcrcnt Ord vician br:~chiopod EariWIIe/I!S adiiC/1(1 xracrlt • deu ' ng r
ages Jnd distinguished by nun r differences in re erred to earlier, according to a ta.xonomic schem~ t;u·onorny,
morph logy. The e. in Harper\ (1 YXIJ) monograph, in which the author of the taxon and the year of hJ~ fc a lw
arc wntren EachDitctr_~ r1dvc'1111 mft~ella Reed LI) 17 (de'- p u blicati u are quoted. l gJc:il cl.ts•
ignaring Reed\ anginal m.lterial). and Eoclumetc.s best nteth
adall!llrl Reed 1tJ 17 .~rrw/rs s p. IIOV. ubscqucnt Phylum Brachiopoda Dumcril LRU6 pl.ln: rnucl
auth r<1 wiiJ n:fer to the btter, m full. ,1s Eochom:tes Subphylum Rhynchoncllif< nnea Willi;Jms cl al . dcSCL' Ild,IJlt
ada>ell<IJ!'''cills HJrper 19~Y, or m abbrevtated fonn a~ 1996 I ')90).
E. ad1 t'lltl .~racilt\ Clas ttop homenata WiJliams 1!1 nl. 1996 I· l t ' 1111
Where. due t tndllti!rent prcservauon. or lack of rder Strophomenida Opik 1034 .111d subje t
m up-to-date m nognph1e ba c, a specie cannot 'ubordcr Strophomenidina pik 1934 kmd of clas
be 1denrtfied w1th ert;untv. it may be des1gnared a~ Supcrtamily Plcctambortitacea Jones 19~X 111 da,q al
:UT. (related to) or cf (ma be comp:m:d wirh), an Family owerbyellidac Opik 1930 h ~,11 r~cnrc
existing ~pe tes (e.g. Alt111ogmptlls cf. l'OIIIt'riiiiiS). Subfamil ·awerbycllinac pik 11)30 1.1 on m.ist:
Where the tl.), ~11 can be Identified as bel ngtng t a Genus Eud1011etes Reed 1917 opung tor
kn wn genu . . but c.lnnot he 3SlTtbed to ,1 species (as Species advl!lla Reed 1917 ml. rc:Jlisn
may be the C3St" where preserv. ti n is poor or if ub pecie~ gracilis Harper 19R9 \ 1nthat
only .1 fragment t , pre~crved, the suffix sp. (plural C]\I.Ultlfied a
~pp .) i. used (e.g. Cl/orcl'r/5 <;p.). If the specimen, in a In the above ~ection we have seen the divt ion oi ' I 'lu~ter
more cxtrc-m • case, -an only tentatively be referred the ta.xonomi hierarchy. but in defining the~l ' uld be l
to 1 gcm1s, one w uld wnre e.g. ?KIII<'~{!illa sp. If groupings how d t xon mist actually go about it? unteric ai
only the spenes 1 dubtous ~uch an a riprion as The basic principle i that morphological and thcr HI ~ome ins
Eoplectodonta ?pct~ki/1('11515 might be used. simi laritie reflect phylogenctic affinity (homol- nJri:'J ince
All t.1xonom1c w rk such as tlli~ must foUow a ogy). his is always so unlcs~. fc r other reasons, h best r•
p:trncubr ser of rule;;, hKh have been worked out simjbrity re ults from convergent evolution. But in
by a cries of lnreman naJ Commissions and are a.~sessing 'similarities' how does one decide upon Ill re impo
documented m full m the opening pages of each w lu h characters arc important? Hm: ~hould the~ l1brau1 111 ·a a
volume of the Treatise 1111 lllr'!'rtebratc Palcomology (a be chosen to mirumize subjectivity and produce llllflH:ncaJ t;
continumg serie~ of volumes pubhshcd by the natu ral order groups? There i no universally The third
Gcol Jgtcal ciery f America). accepted meth d of facilitating the e ends and I nt:, empl·
taXonomists have used different methods. [n recent lllJnitht, in
Taxonomic hierarchy years three sharply contrasting cho I~ have lh · distribut
Although aU taxonomic categoric~ above the species emerged: these are the schools of (1) evolutionary tms. It is
level are t some extent artificial and subjective, taxonomy, (2) numerical taxonomy and (3) cladism, · tematic!
ideal! . they should as far as possible reflect evolu- Until the 1970s m t palaeontologist , e~penally tu~omologis
tionary rdation h1ps. those working with fa . il m terial > hich they hJvc .lppltcd vigc
imil.1r pe i~s arc grouped in gen el'a (singular collected in rhe field, were evolutionary taxono- md racrJii
genu s) , genera iJl fam m es, f nulies in o r ders, mi rs. In erecti ng a hjer;1rcrucaJ clas~ification . . uch there were 1
orders in class sand cla~. es m the largest division of classical ta "onomists used a traditional :md very t1ex- . dmJt, the p
the ammal kingdo n : phyla (singu!dr phylum). iblc ombim1cion of criteria. First there i' m o rpho- evolved, da<
logical (or pbenetic) resemblance, the extent to far the most eflcctivc method for rccon tructmg
which th JnimaJs resemble one another. Sec nd, phylogc-ny. Smith's ( 1994) e::o.:planacion of cladistic
pbylogenetic relationship arc along with phc- methodology is so comprehensive that only brief
netic rr~cmbbnce considered m1portant. By this is comments arc given hen:.
meant rhc way (as far as c:m be dL'tennined) in Hennig was of the opinion that recency of com-
which anim.lls are acn1all related to L"ach other, i.e. mou origin could best be shown by the shared po -
m tcnru of recency of common origin, which of session of evolutionary novelrics or 'denvcd
(ourse gr~dcs into ev lutionary tJ...xonomy. The characters' . h us u1 clo~e l y related gr up~ we would
order of succcs~ion 111 the rock record and gco- see 's.ha.rcd det;vcd characters' (synapomorphies)
\it.!phical distributi n may play an important part in which would distinguish this gr up from others.
drriding relanomhips . hi5 practical approach to Hcr01ig' central concept was that in any group
tl.'<Onomy, vltich took all [actors into consideration, characters arc ~·ithcr 'primitive' ( ymplesiomor-
h11 for :1 long time been the bac bone f palaeonto- phic) r 'derived·. Thus all vcrtcbratt!S have back-
logiml chs ifi :ttion, Jnd is ~till considered to be the bones; the possession of a backbone is a prurutive
bN method by stratophenetic palaeontol gists. who character for all vertebrntcs and is not, of course,
pl.tcc much emphasis on time in seeking ancestor- indicative of any cl se relatiomhip between .my
descendlnt relatinnships (Henry, 1984; Gingeri h, group f vertcbratelt. What is a primitive character
1990). for all vertebrates is of cour'lc :1 derived character as
Fonome taxonorni ts, ho ever, the uncertajmies comp.trcd to invcrtcbr«tes - synapomorphy and
md ubjectivity wh1ch are almost inescap:~ble in any symplesiornorphy thus delineate the rebtive status
kmd of dassttlcation eerncd to be particularly acute of particular characters with re pect to a pec1£ic
mcl".l!l\lCal taxonomy. as d sd the limitations of the problem.
t0\1~ record m tem1s of preserv:~tion. The numerical Hennig endeavou red to provide an objective
uxonomi~ts tried to esc::~pc from this problem by methodology for determining recency of ommon
opnng for quJnrified phcnctic resemblance a> the origin in related taxa, bas d upon primitive and
~nly reallstic gtnde ro 11aturJJ groupinb"· lt wa their det;ved characters. uch relationships are exprc ~ed
11ew thnt 1f enough characters were nu:amrcd, in a cladogram (Fig. 1.3), in which dichotomou~
quantified and computed nnd reprc'>entcd by the me branclung points arc arranged in ne~red hierarchies.
tf \11 duster stati ti ~. the distan e between dusters Here taxa A and B share a unique common
.,
e wuld be med as a measure of their differences. ancestor. They arc said to be sister groups. They
umerical taxonomy has been found very useful both share an evolutionary novdry or synapomor-
·r 1n lome inst.mce , bm subjectivity ann t be elimj- phy, not posses ed by taxon C. Now is the sister
naiCd .ince the ope-rator h.as to choo c (subjectively) group of the combi ned tal a A and B, and D is the
>, now best ro analyse the measurements made, and ~isrer group of cornbjncd tax:a A. B and C. [n per-
n m1y n~ed t 'weight' them, giving certain char;Ktcrs formtng a clactistic analysis, ~:hcrcfore, a taxonomist
n more importance (ag:lln subJcctivel ) in order to assumes that dichotomou~ splitting had occurred
y obtain meanmgful results. Hc:m:e the objectivity of m each lineage and compiles an (unweighted)
:e numerical ta..xonomy is less thJJ1 1t m.jght appt:ar.
'Y The third school reues upou phylogc:netic critet;a A B c D
d i1l11e. emph.1sizing that features . hared by organisms
1t mift t. in nature, • hicrarchial pattcm, evident in
re ~distribution of character shart:d amonb>st Orbran-
'Y un. lt is known a cladi m or phylogenetic
n.. systematics - a . chool tound~:d by the Germ n
ly .ntomologisr Willi Hennig (I %6) and was soon
/C ~plit!d vigorously to palaeont logy (e.g. Eldrcdgc
)- !JJJ Cracrafi:. 1980). In the early days of cbdi~tics
:h .~re were m:my doubters (including, a~ I have to
1{- .dmit, the present author). tlut, as the method it elf
)- mlvcd, cbdistics ha c me to be re ogntzl.!d as by Figure 1.3 A dodogrom (for explanation t>ee !ext).
character data mat:rU . The more characters ami opposed by Ridley, I IJKS). the successive appearanct
character tatc.-s there are availabk- the larger the of ta.xa in stmtigraphy cannot be denied a an essen-
database. and brge d,ttabases arc often routinely tial ource of cbt<'l, however imperfect the fossil
processed and the construction of a cbtdogram record actually is . Thus as Gingetich (J 990) com-
speeded up by using one of several computer pro- ments, ' im e is a fundamental dimension in evolu-
grams. The PAUP (Phylogenetic Analysis Using tion:uy studies, , nd a major goal of palaeontology
Parsimony) program. for example, is a te hnique should continu e to be the study of the diversification
which lilll.k.es the fcwest assumpt1 ns in rdering a of major groups in relati nslllp to geological time·.
set of obscrvanoru. So, h:lVing constructed an appropriate cladogram.
How can we ciL tinguish symplesiomorpruc . the next t:~ge in exploring relationships is to com-
(shared primitive) from synaporuorphic (shared bine this information with biostratigraphic data . For
derived) character tatcs? The mu c useful way is thi. , mith (1994) discusses methodology in detatl
'outgroup comparison'. Hete an 'ingroup' (of which The result is a phylogenetic tn:e, wruch shows the
the relationships are under investigation) is specifi- ~plitting of linenges through time and is effectively
cally designated. and compared with a clmely 'a"best e·timate" of the tree oflife'. Very commonly
rebted 'outgroup'. Al1y haracter pre cnt in a vari- there i · an cxc Uent correspondence betwecn thr
able tate in the 'ingroup' must be plesiomorphic if cl::tdogram and the rock record; on the other band
it is also found in the outgr up. Likewise apomor- the combmed cladistic nd biostr.Jtigraphic approach tnCI~~.:pt '
plllc characters are only present in the ingroup. m y throw up unexpec ted patterns. LHl< 11 I.
Hennig distinguished three kind~ of cladisti The ultimate problem, not only tor cladi 111, but Lh · ~;Uti'
groupmgs. Monophyletic group contain the com- for :ill taxonomic methodology, results from con- h Ul ll b
mon ancestor and all fits descendants (D, , B, A vergent evolution. Resemblances in characters or lh · tW
in Fig. 1.3); paraphyletic groups are descended character stares may have nothing to do \Nttlt tl1 ·~~. In: ,
from a common ance tor (usually now cxtin t and recency of common rigin, but from convergenrl!, .lll th '11 1
known as the stem gr up) but do not include all and it may n t always be possible to di entangle the IUhtJ' ll
descendants (B and C, for example, in Fig. 1.3); results f the two. Thus W illmer ( 11J9U) and Moorc Tlti~ i\
polyphyletic gtoups on the other hand, arc the and Willmer (1997) in cousidering the relation hip' tm.l mor
result of convergent evolution. Their representa- between major invertcbrare gr ups argue tlut t1.tlv~i. :u
tives are descended from different ancc tors and 'cladistic analy i b::lsed on parsimony will tend to liiCHlltt '
hence, although the e may look superticially similar, minimize and thus conceal convergence·. and the\ Wtt1t tl
any polyphyletic group comprising them is artifioal. contend that c nvergence at all leveL is tar mor i. 11.111 I. i
A cladogram is not an evolmionary tree; it is an import nt chan has generally been believed. There 1 ~p · ilt ·all
analySJ of relationships. As such it is a valuable and certainly a problem here . Even so, cladistic method (PAL ' TA
rigorous way of working out :md showmg graphi- ology, oupled W1th biostl.ltigraphy seems ro be the <.J!.' 19 5)
cally how organmus are related, and it force ta.xon- best way forward, and an essential prerequisite for l.llld.Ird.
omis to be exphclt about patt rn · and groups. The drawing up meaning±ul phylogenetic trees. 111 d · mi1
method logy of dadist.ic · e peciaUy good when p~1pubrio·
dealing with discrete groups with large morphologi- Use of statistical methods tu unalter
cal and scratigrapluc gap· and to these it brings the Inevitably palaeontologtcal taxonomy c:mies a cer-
potential for real objectivity. A da.dogram show tain element of subjectivity since the information
how sister groups are hierarchically related on the coded in fossilized shells does not gi e a c mplerc Paloe.
basis of shared-derived h m logics, but alth ut;h it record of the stnJcture and life f tl1e animals that
portrays taxonomic relationship in terms of recency bore them. here are particular complications that
of common origin, lhe order of ucccssion in the cause trouble. or instance, pabcontological xou-
rock record lS n t taken into account (though omy c:m do little to distinguish sibling species,
implicitly dadogram haw a time axis). which look alike and live in the same an:~a but can-
o where doe_ that leave tl1c potential contribu- not interbreed. Polymorphic species, in which
tion of stratigraphy in rcconstmccing phylogcny? many fom1S arc present within one biological Polo~
Whil.~t a few 'tran.~fonned cladists' negate the value peci~s, may likewise be hard to speciate correctly . llll eco
of the fossil record .Urogcch~·t ( view vigorously rn p.irticular, where sexual dimorphism i~ str ng th 'If CllV
I
:e rh~ m;des ami females of the one pecies may be so ancient organism in their environmental context.
1- dJ\\tJJiilar m appearance that they have sometime AU animals are adapted to c:heir environment in all
;il hc,n dcscnbcd a~ diffcrenr species, and the true situ- of its physical, ch em.ic:tl and biological aspcc . Each
1- anon m.l} be h.~rJ to di~entangle (as with species is precisely adapted to a particul r ecol gical
1- .umnnnJte\; Ch:1prer R). niche in which it feeds and breeds. It is the t:ISk f
w Whcu 1t comes to the di~tinction of closely palae ecology to fmd ou t about the nature of the c
111 rcbrrd species, however, there arc a number of st - adaptations in fossil organism and about the rcb-
,• Oltlc.ll tcs · tlJJt may help t giVe a higher degree of ti n hip of the animals with each other aud their
n, ob ccrivttv. ne sirnpl l·n variate test in comm n environments; it involves the exploration of both
u1r, tor x.unplc, can be used when .l series of present and past ecosystems ( chafer. I Y72)
r p-owth stages arc found t gether. If a collection of AJ.t hough palaeoecology is obviously related to
il. br.1chinpods is made from loc.1lity A, the length/ ecosystem ecology, it is n t and catmot be the same.
1e w1Jth ratit'~
or 'ome mher appropriate parameters In modem community ecology much empha is i
·I. m1v be plotted on a graph a~ a scatter d1agram. A placed on energy flow through the system, but thi
ly lmc of best fit (e.g. a reduced major axi ) may then kind of determination is just not p ssible when deal-
'le lx drawn through the ,c tter. This gives a . imp le ing wi th dead communities_ Instead palaeontologist~
1d )' = 11X + b graph, here a is the gradient and b the perforce mu t concentrate on e tablishing the com-
.. b mtt'f(ept on they, xi&. A similar scatter from a pop- position, ~tructure and organization of palaeocom-
ultti~lll mUectcd fr m I cality B may be plotted o n munitie , in attempting from here to work out
ut tht 1,1mc graph. and tht" reduced major axis drawn linkage patterns m food w b and in inve tigating
n- fr llll tlm too. The relati vc slopes and separations of the autecology of individual species.
r th~ two axe~ may then be compared statistically. If Ma ny attempts have been made to summarize
th th~~ he Within a cerwin threshold the population~ categories of fossil residue, to provide the back-
:ef tan thm bl! regarded as being of the ame pccies; if ground for interpretmg original community ttuc-
hL outstde 1t then tht• specie are different. ture. The cheme proposed by Pickerill and
If(' Thh i' unlv one of wh le series of pas ible tests, Brenchley (1975) and amended by Lockley {1 <:l83) is
ps mJ more elaborate techniques of multivari. te used here:
I t .nuly,•s are becoming in rcasmgly important in tax-
tu <lll\llntC l'volutionary tudics. 1. An a semblage refers to a single sample from a
cy Wuh the adwnt of microc mputers and the pro- particular ho rizon.
Ire! ~~ion of pc iahst soft\ . re pncbges designed 2. An a o ciation refers to a group of assemblages,
. ~~ •pt itically to meet the needs of palacontologi~ts all hawing similar recurrent patterns of spcctes
cl- (PALSTAT; Bruton and Harper, 1990: R.yan et composi tion.
he u/, 11)95), the u e of numerical amlysis 1s becoming 3. A palaeocommunity (or fossil c o1Tll11unity)
~ ( \c1ndani. tati tical mec hods .1re likewt e es ential refers to an as emblage, association or gr up of
111 d fining palaeocommun.iries, in 'undefom1ing' association inferred to represent a once-distinc-
populations of defom1cd fm~ils md comparing them tive biological entity. Normally tins only repre-
to unaltered material. sent.~ the preservable parts of an original biological
!r- community, since the soft-bodied animals are n t
on preserved. This definition corresponds more or
~tL' Palaeobiology less exactly to that of Ka ffman and cott (1976).
lal
:J.al \anous categones may be included in palaeobiol- Palaeoe ology must alway remain a partial and
m- ogy: pabel1ecology (here dis ·u sed with palacobio- incomplete sc ience, for so mu h of the information
e, g<·ography). ftmctional morphology and idmology, available for the study of modem ecosystems is sim-
lll.- eJt'b o whtch requires s me discuSSion . ply not preserved in ancient ones. The animals
!Ch them elves are all dead and their soft parts have
c.u Paloeoecology gone; the original physic and chemistry of the envi-
·ly. IlK~ ecology is the study of animal in rclati n to ronme nt is not directly observable and can only
10g th 11 cnviro11ment, p:1laeoecology i. the tudy of be inferred from such secondary evidence as is
available; rhc bells may hJ e been transported ,1w::~y Modern m arint: t:nvironrnents arc graded accord- ( )llly tl
!Tom thctr origm,tl envtronmcnts by ntrrcn~. and ingly to depth . The liltoral environments of the mally pres
the fc s t! assemblages that are found nuy well be shore grade into the subtidal helf, and at lhe edge t'(liments
mixed or incomplere: p st-depo~1t10n.tl diagenetic f the ·hclf the continental slope goes down ro by b.t~.llttc
processe. may h.we dltered the evidence stil1 further. depth· th is i the bathyal zone. Belo" this lie Lhc ,n lowl
De pi re this pabcoecology r ·m.tins .1 valid , if partial flat abyssal plains and t e badal z ne of the deep- tlnally co
cience. Much ~~ now known about the po~t- a can trenches . There i often a pron unced zona- below the
morrcm history of organic rcma1ru (taphon 111); tion oflifc fonns in depth zones more or less parallel he o eat
ha peer 12). of \\hi h an additional dimensi n with the hore. ln ddition there IS a general olde t . ed
involve' buml pruccssc~ (bio tratinomy) . hi~ decrease in abundance (number f mdtv1du.1ls) bur Jgc . hest
help to di entangle the various factor rc'>ponsible riot nece sarily diversity (number of ~pecie ) on ,IJld are s
for dcpos1ti n of a p;lmcular to stl assemblage, . o de cent into deeper water from the t:dge of the shelf. tlwre arc
char assembl gcs pre. erved in si/11, which can yield The faunas of the abyssal and hadal region were now uplifi
valuabl pabe e t lobric.1l mfom1 non , can be dis- originaU derived from those of shall w waters but W11at i
tingui hed from .1 · emb!,tges thJt have been trans- 01re highly adapted for catching the lim1ted food therefore ,
ported. available at great depths. Tht:se region . are, ho - p rt, fth
Bio tratinomy or prcscrvatiun lmtory has both ev~r. impoverished relative to the shallow-water m nts oft
pre-bunal and pose-bun, I elements. The former rcgwm. litt ml, la£
mclude tramponation. phy teal, chemJCal or biolog- Animals and plants t at Live on rhe st:a floor are h ]f real!
ic;ll damage to the shell, and the Jtt.lchmcnt of epi- benthic; those thJt drift pa sivcly or v1m feebly m t wards t~
fauna . Pmt-buri;u processe\ may involve the water column are planktic (= planktonic) lying tfsh
di~turbance by burr wers and sedinH:nr eater~ (bio- ·ince they are rhe plankton. Nekton (nektic r r whcr t
turbation), cun-ent rew rking, .,u)ution and other nektonic fauna) on the other hand compri es a hve
diagenNic preservation change . swinuncrs. Neritic animals belong to the hall v, Horizor
waters near la nd and include demersaJ element\ The main
Modern environments and vertical distribution of which li ve above the continental helve~ and tt-ed t1 11 of re
animals on benrh.ic animals thereon. Pelagic or oceanic nature uf
Figure 1.4 ·hmvs the main environment~ within che fau nas inhabit the su r£1ce waters or middle depths of lcn e. Tht
Earth '~ oceam , t the present d.1y and the nomen- the open ocean~: bathypelagic and the usually cc.:ans is I
clature tor the distribution of marine animab within benthic. abyssal and hadJI organisms inhabit th~ th<: the r
the oce:tm. great depths. \ ·ale . T ro·
tauna.,, an,
but thC" lll
mean sea level relatively I
the specie:
photic individual!
200
aphOtiC
alinity
OCEANIC thousand.
i.e. confin
Ill few are e1
4000
a;~ alinity. Tl
E logically
5000
waters are
especially I
cializcd :111
gene~ can
the Jurassic
CONTINENTAL SHELF SLOPE ments whi
pointer to
Figure 1.4 Modern marine environments. A, B and C in the inset refer to suprolittarol, littoral and sublittoral environments. (Based
on a drawing in Loporte, 1966.) which the~
Divisions of invertebrate palaeontology 1S
1- Only the shelf a11d slope environments an: nor- Water turbulence may exercise a substantial con-
nuUy pre\crvcd in the gcolog~cal record, the trench trol over distribution. and the characteL of faunJS in
1rduncnr mely o. The ctbyssal plains are underlain high- :md low-energy environments an: often very
by b.Ldrtt rock. furmc:d at the mid-oreamc ridges disparate. Robu t, thick-shelled and rounJcd spcctcs
Jnd slowly movmg awclY fr m them to become an: normally adapted for high-energy conditiom,
,_ tinally consumt>d at the subducti n zones Jying wh reas thin-shelled nd fragile fonns pomt to J.
b low rhr o c3rlic trenches; it move a rigid plates. much quiete r w:ner enviromncnt, and it may be
Thr ocean basms Jrc \'Cry young geologically, the possible eo infer much Jbout relative turbuJence in .l
oldt-..,t 1cduncnts known therem being of Triassic fossil environment merely from the type of shells
agt'. ThcsL arc now appr achmg a ubduction zone that occur.
.md arc ~oon to be consumed without trJcc. Hence
thL•re Jl'C very fe~ mdications of abyssal scdim nt MODERN AND ANCIENT CO.MMUNITIES
now uplrlted and on rhc continent . In shallow co ld-tempcrantre se:t marine inverte-
Wltat r~ preserve-d in the geological record is brates arc n01mally found in recurrent c::colog~caJ
tht•tTforc only a fr:lct1 n. albeit the most popuJow; communities or associations. which ~1re usually sub-
·- pJrt, of thr b10t1C reJ]m of :m l nt tintes. The sedi- strate related. In these J particular set of species
tlJ(:nts of the continental shelf mclude those of the are usually found together since they have the same
lator.l!.lagoonal, shallow subodal, median and outer habitat preferen es. Withi11 these communines the
1h ·If r a.lrns. Gencrall ' edin1ents become tincr animals either do not compete directly, being
t•lWJrd~ the:: edges of the shelf, the muddier regiom adapted to microniches within the same habitat. or
i)l!Pllffihm·e. There ma be reefs dose to the shore have a stable predator-prey rebtionship.
tlr where there ts J pron unced break i.n slope. Communjty structure is normally well defined in
cold-temperature areas, but in warmer seas whc::re
Horizontal distribution of marine animals diversity i higher it is generally less dear.
The main control\ ffccting the horizontal di tribu- Pctcrsr.:n (19 18), working on the faunas of the
o n of recent .md to .Ll an..i uuls are tempera[Un.:, the K:megat. first studied :1nd defined some of these nat-
c nanrrr of the sub trate, sahniry and water turbu- mnlly occurring communities. He also recognized
)( lenn•. The larg -. cale d:i tribmion of animals in the two categories of bottom-dwelling animals~ infau-
y on·ans \. largely a fi.mc-t1 n of tcmper-.tture, whereas nal (buried and living withm the sediment) ;md
e rhe other factors generally operate on a more local epifaunal (livi ng on the sea floor or on rocks or
ll'Jle. Troprcal shelf regi os carry the m st diverse seaweed). It w;1s soon found that parallel communi-
!~tunas, and in th ~c rhe peaes arc very num rous ties oc llr, with the ame genera bur not the ·;une
bm the number of individuals of any one pccies is specie>, on the oppo ite sides of the Atlantic. Since
rL•iativeiy low. [n tempeldtC through boreal region this pioneer work a whole science of community
the )pecics diverstty is lcs~ . though the number of ecology has grown up, having it co unterpart in
indiViduals per species c:m be ery large. palacoccol gy. Mu ch effort has been expended in
5Jlinity in the sea 1s of rhe order of 35 parts per trying to understand the comp sition of fossil
thousand. Most m ::mne animals , re tenohaline , conununitics, the h:tbits of rht> ar1imals composing
t.~. confined to waters of near-normal salinity. A them, community evolu tion through time and, a
fc1~ arc euryhaline, i.e. very tolerant of reduced far as possible, the controls acting upon them (e.g.
illinity. The brackish w:Jter environment is physio- Thorson, 19 57, 1971) _ Tl:Us is perlups the most
lng~WJily 'difficult', and faunas living in bracklsh active field of paJaeoecology at present, as J host of
wJtt:rs are normJlJy composed of very few species, recent original work testifies (e.g. Crajg, J 954;
eipt:cially blValvcs and gastropods belonging to spc- Zieg!er et al. , 1968; Boucot, 1975, 1981 ; Scott and
ualizcd md ofi:en long-ranged genera. These same West, 1976; M cKcrrow, 1Y7H; Skinner et al., lSIHI·
genera ran be found in sedimentary rocks as old as Dodd and Stanron, 19YO; Bosence ami Alli>on,
rhe J.rrassic, a11d their ccurrence in particular scdi- l 995; Brenchlcy and Harper, U97).
ments which lack nomul marine fo sils is a valuable As HiNich (1977) makes ir clear, however, mo~t
d rointer [0 reduced saliniry in the enviroruncnt in fossil assemblages 'lack soft-bodied animah. Where
which they lived possible tr;lce fossils c:m be used to compensate for
thl~ but they an. nu real \ubsrirmc'. llence a~~em cnvtronment. Living conuuunJtles arc thadore gcn-
bbgcs, .tSsociatHm' ,1nd ~ven p;llacol"ommuniries cr:~lly well balanced, the number of speue\ and md1-
must not be comidc..'rcd ,1s din:ctly l'qu1vaknt to the v1du,1l~ of particul:lr ~pec1c~ being controlled by th{
\Ca-Aoor communiti..:' of the pre~cnt day. Us1ng nature .1nd availability offood resource~ ro,~il ~sem·
b10strannom1t .md 'cduncntolog~tal datJ the btlges 111JY be tC\tCd a Cl ordmg to this conu:pt. Ir thq
'dt•gree of distortion' trom the ongunl commumt\ arc 'unbalanced' then l'Jther (l) there n1.1y h.1w been
can m \ome tJ\C\ bt· c'timated. ~ot£-bod1cd unprc\ervabk organisms wh1ch on~nJil)
completed the balalKt' or (2) the as~cmblag~· h,t, het~
FEEDING RElATIONSHIPS AND COMMUNITIES mixed through tra1hportat1on and thm doe\ not
All modern anilll,tl\ tc..·cd on plants, other :m.imals,
organ1c dcuitu' or dc~rJdanou products. The tiny
plants of the pl.mkton .tre the primary producers
(autotrophs), a' 1rt \CJwccd~. Small planktic ani-
r~flcct the true onginal tommumty.
FAUNAL PROVINCES
Mannc zoogcography (Ekman. 1953: Bng~. Ill7~
-
Hall.un, 1996) i\ pnlliJ.rily concerned With the
-
mal~ art the primary consumers (ht rb1vores and
dctntm eater\); thcrc arc secondary (canuvores) global J.1,rribution of lll<lrme tauna~ Jnd wnh the
and tertiary consumers (top carn1vores) m turn. dcfimtmn of faunal provinces. Thes~ ar<.· l,tr!(c
Each animal '>Pt'Cie\ i' therefore part ot :1 food web geographical rq,rions ()f the sea (and mo~r p,lrtJC:u·
of trophic (i c. t\.·cdmg) relationships wherein there brly the continental \hdves) within wh1ch the fau-
.:tre a number of trophic levels. In palaeoecology it na~ at the specific, generic and sometunes f.muh21
1\ rareh pombk· to draw up a rc.1hmc food web level have a dt~rinct Identity. ln faun.:tl pnl\'inct') f,gvrc I 5 (
nun) of thc animal' arc endemic. I.e. not iound dmwing in Bri
(though thi' " om: of the more tmportant .l~pect~ of
modem ccolog)). but most fosstl ammJh un usually outs1dc .1 particular provmce. Such pro\ mct·s arr
be ass1gncd to thctr lOrre.:t teedmg r-, pe .llld so the often ;;cpa.ratcd fron1 neighbouring provmtt\ b) I •H ex.unpl
trophic levclm,Jy be c~tunated rcaso1ubly. f.1irly sharp boundaries, though in other CJ'\l'S tl11 IJitii.L, 1\nl.l
Of primary tOil~tHHef\ the following types are bound,trics may be mo1t: gradational. l\hlkr w,1t1
Important: Ftgurt· 1.5 ~how' the mam zoogcogr.1phial \Vc~t Atl:!nl
regtom of the prcscnc continental 'helve, a~ dctined !.lied tor ' '
ftltercro; or \U,pcn'ton feedel"'i wluch are mtaunal by Bn~" (197-l). d1wrgcd "'
or cp1taunal ,\IUil\,th 'uckmg 111 ~mpeuded organic Trop1cal ~helf,w:mn temperate. cold temper:Jte !ihl h. I ikc
m.1terial from tlw water: and polar regiom can be di~tinglllshed, who~c luml\ ( ri(Jp.q~m)
epitaunal 'colkltl)r,· or detntm fet•ders, which ::~re controlled by latitude but also by the ~prc.td ot 711\I!;CO~•.I,lp
sweep up orgalll<. flt.ltcn.JI from thc ~~a floor; warmu or coldcr watc1 through maJOr m.mnc cur- shl'lvc~ h.t\
some infaunal b1valve~ and wonm .m~ .1lso collec- rents. Tropical shelf taun.1s occur 111 folll s~par.lt. n11granh th
tor\; provmce\. Of these the large lndo- We,t Paotic m..:nt' v. h1c
swallower\ or dcpo,it feeders. which are mfaunal provmcc, extendmg from 'outhem Afnca to ca,tem nt these l'lll
Jnimals unsdntivcly 'coopmg up mud nch m Amtrah.l, i~ the riclte't aud most thverse and hl! 1111\\ long tl
organic nutcnal. been ,1 m~uor centre of dispersal throughout th( The dt•vt
Tcrt1.1ry. Smaller and gcucrJlly poorer provmces ol ht ~~~ c.'h.lrl(
Scconrury and tcrti.u-v consumer~. the c.Jn11vorcs. may tropic.ll fauna~ arc found 111 the East Pal·ifit, Wt·~r pre,t•nt Me
prey on :my ofthc\t', but H 1s the c.:onumm1t1cs of the Atl:lntJC and (lea-;t diverse oLill) East Adant1c. I he' \\ 1'~ hcmg
pnmJ.f} trophit level that are mo!>t conunonly pre- arc \Cparated both by land bamef\ (e.g. the Pa1unu l!Jll.un . 19
~ervt•d became ol thcir 'heer numbt•r of md1\. iduak lsrhmm) and by regtom of cooler warcr (e.g. wheff 1971J: Mc.:K
In many livin~ l'Ommuniries most of the 'bio- the cold Humboldt Cun-cnt 'weeping up the west· COn)\lflCtHH
mass' IS Jctually tUlltributed by VC£) !C.•,, t.u:a, llm- ern ~ide ofSouth Amenr.t restrict.~ the rwp1cal taun p<tss1hll' to
,tlly not more dun livt· (the trophic nucleus), but to within a few degrees south of the equ.1tor). 31lt"ll'llt t:nu
there may be reprc,cntativc:s of a number of other The ~hclf faunas of cooler regiom of rhe world tllll'Jlt.ll \ll;l•
speCie~ m snull mtmbt·r;. ln this sy<>tcm competition arc likewise restricted by temperature. .111d ,tgatn ZCI("Igt'Clgl'.lp
between the \pe<.'le' lOJKemcd seem.' to bt: 011111- wann or cold currt•nb cxercise a strong rontrol of a l-lcmng ll
nmcd . It ts thus mutualh beneficial \IIJCt the dttfer- thc1r d1smbunon. Somc zoogeograpl11c ·,.,lands' can 'lOOH' 1nt\.•rc
cnt taxa arc cxploitmg J.1tl\.·rcnt n.>sllurcc~ \vttlun the be 1\obrcd by rcg~om of w.:tm1er or cooler watc~ Cllfll'llt 'YSt
Key
re}'1.{{fJlrop~l shelf
~ warm 1emperaa~
F~~ cold tempera te
- . cold curroots
Figure 1.5 Dislriburion of modern marine shelf-living animals in founo l provinces, using Winkel's 'Tripe!' projection. (Based ono
drawing in Briggs, 1974.)
F r example, the tip of Florida carrie a tropical shelf Modem and ancient reefs
fluna, i alated to the northeast and north we. t by Throu ghout geologica l time animal have not only
cooler water arc:as, but though this is part of the become adapted to particular environments but als
We-.r Atlantic tropiCal shelf pr vince it ha been i a- themselve created new habitats and environments.
lated or some time and therefore its f: una has Within the e th ere has been cope for almost unlim-
d1wrg d somewhat from that of the Caribbean ited ecological differentiation.
1helf Likc\-..i.sc some ce:ulll islands (e.g. the Perhaps the m st striking examples of such bio-
GJ!apago } may be considereu as part of a general genic envi ronments are the reefs of the past and
70LlgL'Ographic rcgi n or pr vincc, but as their present. Reefs (Fig. 1.6) are ma sive accumulations
shelves bave been initi.all colonized by chance of limestone built up by lime-secreting algae a11d by
migr,mts rhey ma have very many endemic de- variou kinds of invertebrates.
ments which have evolved in 1solation . How many T hrough the activities of these frame-builders great
of these endemics there are may depend largely on mounds may be built up to sea level, with cave.
how long the islands ha,·e been isolated. and channels within them pr VIding a residence for
Thr development of coday's fauna! provinces bas
hern charted throughout most of the Tertiary. At
6ASIN SLOPE AIM LAGOON OR SHELF
present Me~ozoic and Pala.eoz01c province an: hkc-
\\1sr being documented (Middlemiss et al.. 1971 ; reef reef faclflS
fl.illam, 1973; Hughes, 1975; Gray and Boucot,
19 '1; McKerrow and cotese, 1990) . Whl'n u~ed in
conJunctlon with palaeocontinental m<~ps it may be
po ,jb[o.> to see how the di, tributiou pattern of
netent taunas related to t h~ p sirion of ancient con-
nnrntal masses and their shelves. omctimcs palaco-
zoogeographical and geological evidence may have
a be.mng upon palaeotemperature and even allow
10me mterence w be made regard.mg ancient ocean
cmrmt system . Figure I . 6 Generalized section through o reef (algol or coral).
innwnerablc kmds of Jntm,tl., .tll ect log~cally diffcr- ing tn clump~ and thickLtS on the reef \uri:JCc lr.tVt: cxp.
entl.1ted tor thcu p;.rmular mche~. When these carlic't of reef invertcbrJtes bcc::um du11ng d 1
In tlte bJirier and parch reef\ of the tropic. I se.1~ e. ·tinct in Llte Middle .unbriJn time there were no I rom \ hi,
of tod.1., which grow up to the ~urt"Jce. the wam1, more reef animals untd ~omc (1{J M,t l.ttcr; the only lllll t h,tV
oxy!Se11-nch. turbulent waters allow rap td calcium reef., were strom.ttolinc. In Middle OrdoviciJn ltlm When th
metJbultsn 1 Jnd ILcnce contmuous growth. The these algae were JOined by cor.tl., and strnm,Jlo- lll.JIJ~ piJ
principal frame-builders arc ,llg.Jt' .md corals, buc porotds 0imc-secrcnng ~p mgc"; -:h.1ptcr 4) a~ well fl'C r 1Ufllll
there .tre many ot er lands of mvertebr.Hc~ m the a~ by red .tlgae, which together turmd a reef cnvi- ~ lt:ll~ .1~ t
n:ef community: sponges, bryotoam .md mo11mn ronnwnt attracting a ho'it or
other invcrtebr;m:~ ll)l)dtl..'d
among t others. ~ome of rhe~e ::tdd m minor way~ to including brachiopods .md tnlobitcr; , For unknowr; t/:c UJJdJS[
the red· framew< rk; othe~ break it down by bonng rea,om tlm type of reef complex diJ not continu btate,.
and gr.tzing. The growth of the reef ro sea level beyond l.lte Devoman time. The red;, that arose 10 l hne
continually keeps p.Ke Wtth mbstdencc, bur it i also the C.lrbomferom were mainly algJl. ~trumato cot.tl reel
be111g continu..UI ' eroded In a typiCJl coral reef poroid .111d corah no longer playing ~uch .111 tmpor- the bcgm
complex the reef mclf is a hard core of cemented rant part in their cPnstruction. tmc.:d to 1
.tlgal and lOr,ll skeletons [King seawards, .md a the The s.1me kind uf reef continued into the Jll tu tl
reef sub tJe tl grow. outward~ over ::t foren:ef slope Pcm1ian, ::tnd many of the~c fringed th~.: siHinki11g U)lltHllUJ1
of rumbled boulder\ broken from the reef from. inland ~eas of thnt time. They rme ar the edge nl ,J!lt'LkJ b
Behmd Jt IS a L goon w1th a cor.1l ~and sediment and deeper ba~ins in which rhc water pcrindicalh \\Or d ft:'l
tidal flats along the shore colonm~d b c:yanobarreria became more saljne ,1s it ev.:rporatcd; there wa~ ltk.l· lltJrlllL cc
('blue-grcem'). Green .tl~ae .tre commonest in rhe wise evap r.ttH n m briat1t ~alt pam hchind the recf.11
back-tccffane': red .tlg.H~ Jre the lll.lll1 lime ~eerercrs w::tttT drawn through the n.·cf dried out in tht Functio
of the rt>ef 1tsdf lagoons behind. In the Pcnni.m red~ uf Texa~ .u1u Ih~ tunc
Largt• reefs such .Is the ab l\'C J.re lawwn as bio- northern England, w ich arc very l.1rgc, the reel he I.'~ I ;tbltl
hcnn · they tonn di~cretc mound~ rising from the front rose .L., a vertical wall of laminated algal 'het•t . :wologi ts
ea tl or. Biostromcs . on the other hand. are Aat turning over at the reef cre~t whenc- it re.Khcd '>CJ t 1w~rds c
lamlllar cunilltutune~ of rcet:.type anutul~ :md barely level (Fig. 1.7). ·h •n the
rise: above the sea floor. The upper ~urfi'tcc w. intemely coloni zed b)' lund1c na'
Throughout ge::~logteal htstnry there hJvc been stromatolitcs, which died out toward-. the ]; gounal 111.111 ' WJ~
various kmd~ of reef l'Ommunttles. whJCh h:JVe back-reef fa ie~. Thjs kind of reef in general mor- ' ~ l1 Vv'hl
arisen, fiounshed and become extinct. In all of these phol gy thnd(m: doe'> not clmcly approximatl' th~ knn n (~•
the frame-bmlder- h.tve includ d ;~Jgae, but the standard p<tttem of Fig. 1.6. Whl'n Lhe Permi,m ~bdl l',tl.t\.:01
invertebrate !Tame-bull :krs h.wc been of different seas h::~d dried out completely the Perm.iJn ret:f- :~r~,m \ I
kinds; the pre ent corals arc only the most recent in complcx type vanished, and then· were uo mon 11 rt:JI c:c
a senes of reef-building ,tmnuls {Ne\ •ell, 1972). red~ unt1l the slow bcginninbrs of the coral-al~al n:~1 l f bJ.zan·
The oldest known rects arc over 2000 M:~ old. sy. tern thJt arose in rhc latl..' Trias.. ic. Coral ret'~ th~ 11 nn
made up entirely of sedtmcnt-tr;lppmg and po ,,bly l1 Cllll.: ,
lime-secreting cy<mClb,1ctcna: the prokaryotic tro- t••WJtds i1

reef flat
matohtes . 'ome of these red~ re.tched con·iderable .ld,lptat!Ot
dimensions. l ne l reported tram the Great bvc t • .1 clean
Lake region m 'anadJ as be111g over ..J-50 m thick , lt rhe'e p
and scparatmg a ~hallow-\ ater c, rbonate pbrforrn till' I hodul
from .1 decp-\vater mrbJdtte-tilled basin . her arc 1 m o
no prc~crved metazoans 111 thc\e reef~. however. and I udwtc.·k
their ccolog1eal structme nuy hJve been very <;im- lu.·' l1r~~:
ple. o only
0 lOOm 11\ L']l[] ll c
With the rise of frame-buildmg metazoans m the
Lower ',u nbna.tt J new lund of n:ef ·o mmunity scc;lo l\leflt I tmd hou1ontall I wo n
made it :1ppeJr.tnce. SrronurolitK reels •.vere Figure 1.7 Crest of a Permion algal reef . (Redrawn from Smith. b dl."c.lllet
invaded by the ~pongc-likc .l rch.teocyat!uds, grow- 1981.) .liHI fomt
h,r.,c ~.).l1,l!ldcd and tloumhed smce then, oLhtT than Thompo;on ( 1917) it h.1s bccn understood that the
I' d1mng J c.ttasrrorhic pcnod m the earlv Cretaceous conformaticm of the parts f any organism ts the
0 tram which there .lrl' no reefs known (though coral~ re. ult of interacting forces, dictated by physicotmtb-
y mtllt h.tvr been hvmg somewhere at tb<it ome) . cmari a.l laws, which have op~:ra.tcd rhr ugh
IC When rhr corals recovered they were joined in ~rowth . The cenrr.1! 1ssue here (Thomas a11J l~l:'if.
,_ ffiJ!l places by the peculiar rudtstid bivalves, also I 'JIH) is the balance perceived between ·accidents or
u tl't'f on11rN, wh1ch at one pcnod Jlmost ~upplantcd lll';tory and the prL'SCnptwn of physical law~. as
i- l'UfJI' Js rhc dommant reef framc-bmlders . Yet these c:1uscs of org:111ic dc~ign ·.
-~ toll dird out in rhL· 1Jte rctaceoU'i, leavmg corals Ditlerent marine inwrtcbr:~te skeletons may be
n th~ undisputed and cl mmant rcd:.buildiug invcrt~: functionJlly convergent 111 the way they g1·ow.
ll' lmtc. and the ~amc kinds of growth pattems turn up
n Th~rc h.1s been some declme in the spread of frequently in represcnt:~tives of m:tny phyl.1. This
,_ mr;1l reefs .md m the munber of coral genera ~incc i~ became there <lre relatively tcw way~ in which
th · hq~inmng of the T C'rtiJ!). They are now con- an animal can grm and yet can produce ::1 h.trd
tinld to rhc Ind -Pactti c regton Jnd on a smaller covt·ring. 1nvertcbrare keletons, by contrnst wtth
iC cak m the Wc't Atlannc. rlm dedine may still be those of vencbratc~. arc generally cxtemal, :md this
lOIItinuing. though the rcef'i were not significantly narrow~ down the ~pectrum of growth possibilities
!Twc.d by the Plmrocene glanation. The future of -;till further. Some of these- rypes of keletons arc as
y wnrld rcrf rommtunrie~. tht: most complex of .tU to !lows.
·- mtnne ermysrems, rem~m~ ro be seen.
External hells growing Jt the edges only by
IS
IC Functional morphology, growth and form accretion of new material along :I p:trticular mar-
d rh• fum·t1on, of part!Cttbr organs m fossils cannot ginal zone of b<rowth. Very often such growth
~f bl' l"it.lhh\ht•d by m.my of the methods available to results in a logarithnuc sp1r:1l shell as in brachia-
s. wologi~r,, but tt is still pm ible to go mme W~l)' pods. cephalopods and in coraJh of cert.tin simple
tow.1rd~ cxp1all1ing how part1cuhr org::tm workeJ corals.
wht'll tht , n1mal th,lt bore them was ,1livc. Su h Extemal kelctom of pbtt'~- di~unct, conttguou~
funcoonJ!mtt•rprerarwn lS, however, limjted .uu.l in or overlapping- normally ~ecreted .tlong a smgle
al ntany w y. it 1s h.1rd m Jo{O beyond a certai11 point, zone whi h m:~y be but j, not ~11\.v~tys marginal. A
·- town wlwn the function of a particubr organ is good cx:~mple is the echmodenn skeleton 111
iC kJHlWl1 (~ertion 11..3). \>V)lich the pbtcs once fom1ed ,tre p nnanently
If PJheontologim .1rc often presented either with locked into place, thougl1 they m.ty thereafter
f- or~.lll\ whose function i• not clear ,111d which have gruw individually hy accretion ofnutcnal in con-
no rc.tl counterparts in b'l:mg ::mirnals. or with fossil> et:ntric zone .
nfb12aJTc appearance wJm·h .1re so mod1ticd from Extcmal skele-tons all tanned at the one t1111e.
rlw nonnal type tor rhc r:Lxon that they testify to The arthropod c. ·oskelcton is mmr typicaL
cxtn•me <ldaptations. Some J tcempts c:m be made Grnwth here i' difficult for the skeleton ha~ to be
111>'.Jrd1 inr ·rpreting these morphologJes in terms of periodically moulted, a process known as ecdy-
.tdApl.ltton .md mode of b fc, \-\< hich m rum may lead sis. When the old exoskeleton is casr the <1rthro-
w ,J dc.trcr understandmf!; of evolutionJry processes. pod take~ up water or Jir ,md swell~ to tl1e next
If thm problem~ arc ro be t.1ckJcd, then a cobcrcnt brgcr ~ize. :md the new cuticle which underlies
m~rhodologicaJ ~cheme IS needed. One particuhr tht' old one then hardens. GrO\vth is thus raptd
·~>tem o approach, the paradigm method of and epi odic and is only pos ible during moulting.
Rudw1ck (1%1) , has been much disru sed, but it The di~adv:mtagc of tlus sy;;tem is the vulnerabiJ-
lie, largely outside the ~t:ope of tbts work ;md ity m <.bmJgc and predanon during mouJung.
Ill only some examples of m :1pplication arc
ntt·nrinncd. Es~cnttally, any kind of skeleton, intemaJ or
Two related aspects of palacozoolo~ \v luch eau cxtcmal is 'highly constrained by geometric rules.
h, be deduced !Tom tossihZL·d remains alone Jic growth gwwth processes, and the properties of matcnals.
3nd fom1 . Followmg the dN>ic wor\... of d'Arcy Thi'\ suggests thi'tt, given enough time and an
extremely large number of evolutionary experi- sttatigtaphy :md chrono tra tigraphy, diJ of which 1\ 'pak
ment , the discovery by organism. of "g od" are ways of ordering rock strata inro meaningful "IH:h1~in~
designs - tho e that are viable and can be con- units (Hedberg, 197fi; Holland cl al., 1978) . th arly P
structed \vith .wailable materials - W:l' inevit ble and l1 mn tn
in principle predictable ... the recurring desigm we Lithostratigrophy ,Jhlll!d,mt
ob erve are attractors, rdcrly .md stable confi~ura Litho trarigraphy is concerned with the erection of .11 11 11 1ml.i tc
tions of rn;ltter that mmt necessarily merge in the units ba. ed upon the characters of the rocks and dif- 01\.l cs an
course of cvolunon' ~ homa' and RciC 1993). ferentiated on types of rock, e.g. siltstone, hmcsronc
1n pomt of £.1et the potential avail ble has bet""n or clay. lt is u~cful in local areas and es ~:nti:tl in
very well expl01tcd by living creature , though pe- geological mapping, but d1ere is always the dJng.:t
citic constraint seldom .ul w the production of tl~at even in a small area rock unit~ cut .1cros timt
ideal organism . Nature \ rks a~ a rinkerer, r-ather pbnes. For in. tan cc, if a 'horcline ha been advanc- v ry prcci
than · an engineer Qacob. 1977: '>CC aho Chapter 2) mg in one direction a particular suite of sediment~ I n any l
- a point to be bomc in mind in comidering all liv- probably of the a me general kind. will be left in Jb r1.1 lta.· bcj
ing and fossil organisms. wake. Though thi · bed will appear in the r d 'nlk tion
record as a ~inglc uni.fc rm layer, it will not all havr n Jll'\. this'
been deposited at the one time; since it cuts acro! nm w1th
Stratigraphy t:Hne planes it i said to be di achronous. Sud ht·t·n toun
diachronism i common in the geological record m;1rkcrs. ·
edimentary rock-; h:wc been built up by layer upon Furthermore, many suite~ of sediments are later:ill\ g I gtcal
layer of sediment, which sometimes ha been much m1perSJStem; different sedimentary facies may have l: .ltnpk.
the same for long periods of time but at other times e. · red at tl11:: amc time within a smaU space - J ~tls fnr M
has ch;wgcd it' haractcr rapidly. The indJVidual sandstone layer, for instance, passmg into a lulc Ja,t< d s m
layers within seduncntary rocks are separated by some distmce away. Lithostratigraphy IS thus only t ·'I n eh
bedding planes. Thco;e bcddmg plane~ are rime hori- real value within a relatively rnaJJ region. it 1' ubdi•
zons, and the histors fa rock ~equcnce 1 reflected The divi,ions erected in litho tratigraphy ar• [tnod~ wl
in its bye ring. E,tc h bycr in the rock "equence m11st arranged m :1 hier:Irchial system: gro up. forma· .1 t lillJ n )
hdvc been laid down on a pre-existing layer, so thJt tion, member and bed. A bed 1s a distinct layer 11 Th pr
the olde t rocks ar~ at the bott m of .m exposed a rock <;equence. A 1 ember is a ~roup of bed h \V ver,
equen e, the ymmge ·r Jt the top, unles~ the succe - united by certain c mm n characters. A form riot logtt al sue
sion has been tccconically inverted. This is the p rin - is a group of members. again united by characre < tlwr.. Tb
ciple of uperp sition, recognized as long ago as with features in common. lt i the primary unit nl
1669 by the D.tni~h <;cicnttst NicolJtlS tceruen lichostratigraphy and is most usefi.rl in geol gical Many k
( teno). mapping. Hence it i fonn;~tions that a.r nonmll) d\ cllin;
Strattgraphy i. cone mt:d with the study of strati- represented by di!Terent coloun; on geologtcal mal lived in
fied ro ks, their cla~ ification into order d units and and cross-sections. and a fom1ation is normalh mud 'I!:
their histori al intcrpretati n. lt bears not only upon defined for its mapping applications. FinaJly, a grouf ere o l
p t geological cvcntl- but aJ o upon the history of ranks above .1 form:Ition; it is composed of two 01 uon. of
life and is perhaps the most basic part of all geology m re form::ttions and i ften used for sunpliiy:m~ not fo
(Harlmd and Arm trong, 1990; Bentou, 1995). strarigraphy on J "mall-scale map. 'lhh ml
Much f stratigraphy is c ncemcd \vith chronol- Lt tin • F
ogy; the geological record ha~ to be divided up into Biostrotigraphy \tmvers;
time peri ds wruch arc standardized, as far as po si- In bio rratigraphy the fossil contents of the beds an 2. omc k
ble, all over the v ·orld. ne of the pnmary aim of used in interpreting the hi t rical sequence. h u 1
stratJgraphy has been t pr duce an a curate based upon the principle of the irrevembility
chronology in which not onl the order of ev nrs evolution. This means that at any one moment u· l'll$C fi
but als their dat , .uc known. "tratigraphical cl:mi- the Earth's hi. tory there w<~~ living a umque anri little U~t
ticatwn i basic to all of thi~. pecial assemblage of :mima!,, characteristic of that ood 1;
There are three pnncipal categories of strati- period and of no other. As time went on these wet~ ate an
graphical dassificarion. li tho tratigraphy, bio- replaced by others; eac su n:s~ive fo sil assemblagr h wg d
Divisions of invertebrate pa laeontology 21
1 is J pJle rt:Aecri n f the life :lt the time that the -t. ince fa sil pecies could migrate toll wing their
1 nclo ing sedunent!. \ c:re depo ited . Thus during own environ ment through time, there 1s always a
th~ early Pal,teozoic trilobites a_nd brachiopod were po sibility of diachronou faunas. The zom: as
the mmr common fossils. By the Me oz ic the mo t defmed in one arc may not therefore be exact! ·
.1bundant preservable invertebrate~ were the time- equivalent to that in another region .
f .unmonite~; they too became extinct, and nail Jnd
bivalves are the commonest relic. o cnozoic time. In the example of a graptoltte, thcretore, tor the
rlu is how it appears on a broad ~calc . However, rea ons outlined in (3) and (4), the total range or
\\hen the time range~ of indtvidual fmsil species arc hi ozone of a species is not likely to be pre~erved in
r c an1ined 1t is t:vid m that ~ome of the~e hsted tor any ne area, an d it is therefore hard to draw tdeal
onl>· a fraction of gc I giol time, char,lctenz.u~ isoch ronou boundaries or time lines.
\'cry prrc~rly a particular brief historical period. ideally, zone foss1 1s . hould haw a particular com-
In my local area, once the sequence of fossil tau- buutwn of characters to tnake them fully suitable
n.l' has bcrn precisely c. tablishcd through as tduous for biostratigraphy. hese ' ould be:
mUection and documentation from e::\-posed ec-
llon\, tlll1 known ucces ion can be med for correla- a wide horizontal clistribution, preferably inrer-
tmn witb orhcr areas. Ccrrain fossil pecies have continental;
been found to be p.uticul.trly good ·tratigraphical a ~horr vertical range so that they could be used to
mJrkcr , They chJrJctcrize sh rt secoons of rhe define a very precise part of the geological col-
g~olugical ~ucce · ion known :t~ zones. To r:~ke an umn;
example, amm nite an.: p.trticubrly good zone fos- enough morphological characters to enable them
ih for M e~ zoi · ~tracigrJphy . The Jur,l tc period to be 1dentified and di tinguished e.Jsily·
!.1 red omc 55 Ma, .1nd m th tandard Brinsh sue- strong, hard shells eo enable them to be com-
m ton there are ver 60 ammonite zone by whjch monly preserved;
1r 1~ ubdindcd, so the zone~ arc ddined historical indepe ndence of facie , a w uld be expected
p~nods whtch h;wc an average duranon of less rh an from a free-swimrning animal.
J mtllion ea~ each .
The prJctical probkms in btostrarigraphy are, All of the e conditions are seldom fultiJlcd in fos-
hO\wver, very comple:-.., .md some partS of the ge - ~ils u ed for zonation; perhaps the ncritk anm10nitcs
logtcal succe SIOn arc much more closely zoned than come close t to it, and it ts nor surprising that rhe
otlwrs. The main pr bl ms arc J.S follows. principle of really precjse stratigraphical correlation
were first worked out fully with these fos~ils,
l. 1any kmd of fc ssih, e pccially rho e of bottom- notably by d1e German palaeontologi t A. ppel in
dwelling invcrtcbrat , are f:lcies on trolled. They the 1 50s.
lived in particular environment\ only, e.g. lime- It was ppel too who first recognized that there
mud 'ea A or. reef. sand or ilry sea Roar. They are variou ways of using fo il in stratigraphy
were ficn htghly a aptcd for parti ular co_ndi- which partially circumvent the diO:lcuJties men-
ttom of temperature . •aliniry or ubstr.:He a_nd are tioned, and hence different types ofbioz nes. here
not found pre erved outside thi. environment. arc tour rnai11 kinds of biozones generally used
Thi means that they c n only be used for corre- (Hcdberg, 1976). Assemblage z ones are beds or
lating particular environments and tlm arc not gratip of beds with a natnral as emblagc f foSSlls .
universally applicable . They m y be ba ed on all the fossils prc erved
2. omc kmd of fossils arc very long-ranged . heir th rein or on only certain kinds. They are usually
rate. of cvolnti nary change were very slo\l . very much environmentally controlled and there-
They can only be u cd in a br ,td and general fore of use only in local correlation. R an ge z o nes
It'll e for long-period correlation and arc of very Jre perh p of more general application. A range
little u e for e rablishmg cl se ubdivi~ions . zone u ually represents the total range of a parttcu-
l Good zone fo~sils such as the grapt lites are dclt- l::trly useful selected element 111 the fauna. One may
catc and only preserved 111 quiet environments, therefore refer to the Psi/{lccras plt11wrbi.1 z m:, ba>(:d
being destroyed in tnore turbulent conditions. upon the ep nymous amm nite that dcfu1cs the
2 Principle~ of palaeontology
divcf'\ification tol- tr.lttg r. ph
c wbjc h mere is a' befor e: expans10n ~md
lowc sr zone of the - ur •pe.u1 Jura.,sic, abnv ction . 'ever al snch bwmere~ h.:~vc of' \llC C\Si1
i~ the rhiMIH'Imia 'm.!JIIf,lf,, zone . ·a eh

rangL' zone is lowe d by extin pper U11ly
studi ed \111('
h occu rs been defin ed in the inten JVdy
:Uways name J after J p.lrtl cul.u spent'~ whic .·rn uwan - 11111! , l h
lly usl.'- C.1mbrian of orth Ameri.c.l. The p.ttt~o
is
withi n ir. Whe re there arc a num ber of ZOlu be disce rned m ·tu.JII be
.1
specie~ ably s1milar and could prob Jbly
fitl species, or wher e tht' r:mgcs of mdiv iduJl l t 1s u~u ;
lll.l}' be othe r pms of the eological colu mn .1s \\'cU.
are- long, a more prt·u~c tnltl: deJin 1tion dtf lren r 6
~'Tdphical
given by the USl' of ovnb ppm g strJti tltcn: ,!re s
con curr en t Chronostr atigraphy
ranges. Such zone~ ;Jrc thcre fon· called r [ orreb tton
s .m.' u~efu l hron mtra tigra phy is more fw rc,1 hing than eithe
ran ge zone s. Acm e ur peak zone graph y bur has ir~ root iu both of pod f:m n.t'
in whic h bio- . or litho stran
locall)'· An JCme zone is a b dy of qraw ize the sequ ence of ht1.''· l,rap t
parti cular speciL'S is them . l t~ purp o c is to organ
the maxi mum abum iJnce of a o~tr:wgraphical .111d lnnc sn
zone s rocks n J global scale mto chron
found, thou gh not it\ total ran~ . uch <JCme worl dwid e event~ lJrtc•r , bclll
er hori- untts , o:o that all local a well a~
lllJ.} be n.1rrow hut Jn: nftcn u~eful as mark .ud scale. Henc e 1t ts Jll t1 d\ 111 \\
int rval can be- rdate d to a s111gle stand
zorn in geological mapp mg. final ly. an <lnd their nme rela- In MllltC .u
bio~traci conc erne d with the age of \trJra
z one is Jll mter val bcn: ccn t\VO di,tin ct rch1c al classi ftcat ion of ume - v~.:rtte,ll ~c
distJn crive tions . To do this a hier;1
graphical hori mn. lt may nut have any must be empl oyed . The rht.:sl twl
simp ly a equi vakn r unit~
fossils, or mdl.'cd an fi>sslls :~t all, being rcluc Jl sy~re m u,eJ is a.s ~how n 111 lm ·s. hur 1
of str:Jta cmw cnn n:tl hicra
onve uiem \v.ty of refcm ng to a grou p ,Jf'L ~ ms1bl'
aphic aUy Tabl e 1. 1.
bracketed be(\,vecn two nam~.:d bw,tr atib'T OtdO \.t'iJ i
defined zt ne~. ~l.lgt'\ j ·fil
- and
Biosr rat1g raphi ol umr~. unlik e litho Table 1. 1 Conventional hierarchical correl
ation between Fos" h t
unm , arc not hJeLJ rclua ll y gical unit~ tl
hron o<trangr.1phie:U chronostratigraphical and geoch ronolo Wo<:d L\
es, whic h
arranged, ap.lrt from in thl case ofsu bzon Geochrooological units NL·vcrtht•lt
divid ed Chron ostrat igrap hical units
are local d1vi~IOI1~ \.vhn c a zone can be bed~ of t:•
ewhe re.
more finely 1n a parti cuhr regw n than el Eonothem Aeon ht: f ~il
rent kind uf ~rrati ~'Taph ic conc ept, the b io- Erothem Era"
A tliffe an one ~
cJ a~ a Sy~tem 0 Periodo
mere (Palm cr, 19115. !9H-+) w.1s detln Epocha l'lll11 !l1011 ,
boun ded by abrup t. Series 0
re ci nal biostratib'T:lphic unit Stoge• Age htl-C:renr F.

demcnt~
non- volut i01ur y ch::wges in the dom irunt Chronozone Chron l.1tt. . "' m1
nece ssar-
of a 'ingle phyl um. The~c clung es ;~re not province~
en- •These terms are in most common use.
il · relate d m physKJ.l disrontlllllltH.·s m the sedim thnc fore
The
tary recor d and they lll<lY be dt.lch rono us. lllC'CS Ill V
in ~tudies
biom ere conc ept h;ts prov ed most u~di.Il relate qu1te stmp ly to gnph tcal c
a repea ted ChronoHratigraph.ica] unit~
of lat~ 'amb rian trilob ite taunas , wher e ical unm : thus the r cks of the there Jre 1
recor d. In geoc hron olog
patte m of event~ i~ l•v1Jeut from the fo,o;il were all depo sited durin g tht so rh.H i.ntc
in an initia l Cam brian Syst em
each biom ere rlw ~helf scdu nents conta of these term s are \elf- Jn chro
hical range Cam brian P eri o d . Most
tauna of low d1vet ·1t', ,md short srr.lligrap th.u the) b the f
fauna~ expla nator y, but it shou ld be recog nized
(one or two \pecic~ only}. I-I weve r, later dwid e m exten t. hs lure d
diwr~ific arc all, at least in theor y, worl
withi n the hiom crc beco me muc h more T ht· Psifaceras plmwl'bis chro nozo ne is a tim e Utllt who ever
c t, by this
and oflo nga ~rratigraphic range :~nd sugg whic h the s;nd anun on.itl' l.w.t~ bra 1
h of the equiv alt'nt to the nme in
stage, 'soun d adap tive ptm~· Jnd che zenit if it wa~ con tined to cen<~ in th ,,. 0('(.. 1
there i. wa in exist ence , even
trilob1te t:JUna . At thl t(JP of the h10merc d only. It hard indee d, howe ver. unhk.dy,
d trilo- p :ut~ of the worl
IS
often a r.1ther sp ci.IIJZcJ t:lUna of short -live , since SUI t•r,ed e
e .rti net .1bru pd . to be able to delim it chronozone~ accu rately
bites. then :ill the gro ps beco me to terta in geog raphi ·al ~ n trdy c
way most fossil were cont lncd
The succ eedin g biom ere bcgim in rhc ~amc of the ammJ J~ hv- tln n tr.
f ~inular regio ns or provu1ees , as .ne most
as its prede cesso r, uft~·n w1th trilobite~ today . Ther e are relan vely few \veU-est:~h[i,Ju:d mu'h to 1:
one. Thev ing
appe aram c to thme at the ba~c of the tim nts' :u; they luvc lwcn .tllJ rL'tric
111 trulll a stock of more slowly chro nozo nc ·, or ' worl d insta
may have migr :m·J ne', thou gb it Ius a real jC)HC)) COl
deep er- called. and ~o 'chro nozo
evolv ing trilob ites in ,m outly ing, possibly .1hlc to most pr.1cttc:U pre~cJlt str
new bio- mean ing, i. not :1 term apphc
wate r area. The latl'f devd opm em of the
Bibliography 23
1tr.1t1graphy A stage, on the other hand, b a group

ot succc~sivc zones ha\•iug gre.lt pracbcal use, c~pe
,a.lllv ~ince it i' norm.11ly the basK workmg time
umc of chroumtran~r,tphy, the n:urowe~t tlut .m Books, treatis es and sympo sia
rrually be used on .1 rq.,riuml sole. gcr, D .V . (1%}) Pri11dples 1![ PalactJeco lo,J!)' McGra•' -
lt J\ usu.llly at the st,\gt: kvd rh.1t rock\ of widely HilJ. New York. (Useful ba>ic tc:-.-c)
J1tTarnt fae1cs can be COJTdarcd. A s an ex.tmple liJrrinb'to n, E.j. ~ . ( 1967) Jn,frrelmrrl' Stntrlmt· tllld
then~ m~ some ditJILulttc'> in makmg preci~c zonal FunCiitlll . Nc lso11, London . (Invaluab le zoology text)
wrrd.1tiom between Ordl VJn:Jn tnlobtte --brachi o- Benton, M.J. (cd.) (J 9'-!S) Tltr Fossil Raord 2. <. lupnun &
poJ tauna1 and rime-cq uivaknt t3unas wtth grapto- Hall, London. (Jnv.tluable sun1111Jry of strangrJp luc.tl
lit~l. Graptolite ,tre rardy preserve d in the siltstonc d1sri'ibution of all known fmsil generd)
Boardma n, R..S.. Chectha m, AH. and Powcll , AJ. (eds)
.nd llm •stone'\ f.1voured hy the ~hdly fmsih, JJJd the
l~•ssil l1wrrrcbratc.,. Bbcl-..·wcll, Oxturd.
l.ma, bemg bentluc, could not mhabtt the ~ogn.mt
(I'J!:l7)
(hxcepn nally usetul rnulti-Jm hor rexL)
mud1 in wh1ch the graprolire~ were best preserved. Hmencc , D.W J .md Allison, P.A. (1995) (cds) !l1r!ri111:
In IOme areas, or CUllf\C. the f.mna~ do alternat . in
palm'ocru,irolllllenral ar111l y.1ts from Ji•sstls. Geologtc al
wual . egncnLe sitJLL' the ~lte~ of depos.irion of ·ociery ofLondo n Spenal P ublicatio n No. HJ . (12 use-
!ht'>C two tJ.cie fluctu.ltt:d with oscallati ng ~bore ful papns)
lilll'l, bur cl10ugh prcci~e znnc-ro -zone corrcl,ttions lloucot, A. ( 1975) EP,tllltioll a11rl E.xriurtlnH R11rr Commis.
.lfl' po·sibJ ;lt SOllll: level~ I( lS found in pr.1cticc.: th;H Elsev1cr, Amsterd.un. (Adv,mced tc.·t, nt.JinJy JL·JJing
Ordo•irian grapLOLtt' 70JH;S correLuc best wirh w1th brachiopt)d distnbm1on)
~tJ!:\tl defined on sheUy fu · d~ . Uoucot, A.J. (11JH I) Priwiplcs of Bmrltic Mariw·
Fos1ils give a. rd.ltive chrnnul ogy wluch can be ralacocmft,'?Y· Academi c Press, New York . (VJILublc tf
H:ilO~)'ll ratir)
u ~d J~ the prunary ba i of t hronostr atigraph y.
Brwchle y. P .•md l l.l rper, D.A.T. (l'N7) brosyuwts,
' vmhcbs , it is ufiL·n lurd tn correlate precisd
l::(o/ogy am/ E•'oluthlll . hJpnun & Hall. London .
bed o[ equiValent age in \\ 1ddy sep;lrate d areas, (New, VJgorous and fl!<t(bble tc ·t)
rh~ fo~ il sequence~. thot<gh wdl docume nted \\'tth Bngg~ . D .E. G. Jnd 'rowtl1er , D. ( 191JU) Pc1l.u·,,f,to h!l!)': a
my one .lrt•J, nt.ty cont,1m v~ f} few element~ in S}'ulhl'sis. BI.Kkwd l , Oxford, 583 pp . (The best md
ommon, tf mdt"ed a11y .H .tll. o;mce they bdong to mo~r useti.al p.llaeonw logical compend ium uf all)
ditfcrcnt fauna! provinc~e~ which <He hard to corrc- Bngg;<. J.C. ( llJ74) Marim: ~oogw,gmphy. M..:Graw-HiiJ,
atc. Som times, howevn . tht• boundar ies uf such New York. (Deltnut: ttion of GlmJl provmce'>)
•rtlllncr mJy luve mcilbtc: d m .111d fro. here may Bn1ton, D.L. ~ncl Harper, O.A.T. (1':190) Micro-
hm-for .tppcar demcnt of adJacent t~1unal pro - computer > in palaeontology. ColltrilmtiorJJ o.f tltr
Ptlllll'tlHto/o,~ <c<~l i\Jusrum, Oslo l 70. 1-105. (8 p~pers)
nr,"' 111 verttc:JI succe,,io n. th u facili tatmg strati-
Cowen, R . l <.J<)4. History of Lift, 2nd edn . UladcwciJ ,
rraphKal corret1tio n. At ntO\t •trJtlgrap hic.tl horizons Oxford . (Eminent ly rc:~d.1 ble)
thn~ Jre u u.tUy some ub Lqu irou~ worldwi de fossiJ.., Dodd, j.R. and Stanton, I J. (l9 0) Palncoccolngy.
mum mter(l ntmenta l corrcl.atinn J~ 11 Jt impo~siblc. Cv11rrpts .wd .;tpplirrtfions, 2nd edn. Wi ley, cw York ,
l11 chronostra tigraphy tbe rcbtivc ~e ~uence given 502 PP- (lnterestiug appro.1ch)
I\' the fomh is supplc:mc:nted and enhance d by Ekman, . ( 1953) Zoo,~co.l!mplry <!f tire s~a. idj..,'WJ k ,lJld
.h,olute dJ.te' which cJn be .1ffixcd .at certam pui.nts jJckson , London . (An old..:r but still usefu l text ou
1hrr '\'cr appropriate rock~ ccur. The e are usually m.mnc life zone~ .111d tjunal provinces )
brJs br.tcketcd between (omlifcr ou edirnent s, and Eldredge. N .•wd J':lcratt, j. ( 19HO) Pllylo,~encttc Patll'lll(
tnelr occurren ce 1s not t o comulon . It i' mo~t aud r/Je Evolurio11<1ry Process. olumbJJ mvcrsity Press ,
unlikdy, therefore. rh.lt radiome tnc dating will New York. (Expl.ti.m cl.tdistic n1cthous)
F.1irbndgc. R.. W. and Jablonski, D . (t·ds) (197'.1) 'n1e
11pcrscde palaeonto logJLJI correbti on: the two are
Et~rydtlprtlin ~r Eart/1 Srit•11ccs, Vol. VII, Tlrt' lincyclopedw
Llttlrci} complea ncntary, J.nd the brre.tt success of
4 J>alealllolo,~y. Dm den, Hutchms on and R.o ~.
rhmnostrarigraphy, in ~pit· of it hmit.1tions, owes Stroudsb erg, Penn. (JnvJJuJblc rcterenrc )
mudt to both . The u~e of ..turonutic d.lt< -proccs~ing GoldriJlg , R. ( 1991 ) Fassils i11 tltc Field: lujimllnfrc,,
nd rdrie al sy~tcrns i~ growing and m.l}' (Hughes , PvtCIJir.JI r111d <111 1/ysis. Lon~rnw. 1-brlm' . (How
%9) compms atc fur \Ome of the constrai nts in palaeontoloboical infonnan on i> garhcn:-d m the field)
)'r·,ent ~tratigraphJCal prarncc . (;ould, S.J (199()) /l ~llldcifiJ/ Life: t/lt' Bli':J!W Sh,l[c .uu{ rhc
ll<llllr!' tJ( hi.story. Hurdumon R.td1us, London. Moore , R.. . e1chen, ., Robi~on 1-t.A .111d KJe ·ler gT n ·th: .UI J
( nmmcm' on w..xunumy) R . . (succe ivc editors fr m 1953). TYI'tllise 011 J I. Bonn~.:r
Gray. J. and Ooucot, A. (ed') ( ll)7':J) Hlsr.mcal BiO,!(f<l- fttverrebrare Paft·otllolo,gy. Gcolo~o.oic<ll ocicry of UIIIVt:t lry p
~r<tphy: ?/,m· Tt•ctuuir.s rmd the Clwulli".t: E111•iHmmeut. American .md the Univen;ity K:an .1 Press, awrenc~. IIHll 11 , • • I
Oregon . t.uc Univcrstty Pre", orvalfu, Ore . (Several Kan. (The standard reference work on invcrtt•brate f. S· l ibt .11') • Lo•
papers on l~wn.tl provull:'cs) sils - each phylum treated in scparJte volumes). \ ith ·nntm t
H.tl1uu, A. (lQ73) .1rl,ts •1 p,,/w:,,bit!~C'ograf'lly . El•eVJer, Murray, J.W . (ed.) (19H11) Arias of l111•ertebmte M.l~(o.<5il.l Vtl ·n~me, ].\\
Anmcrd,un. (-l8 origm.1l p.1pe 11 disrnbunon of fossil Lon~rnan. Harlow. fi r rhe 1-'abcomological Asm JatJon Bicl>1
\ltlflllf
faunJs) (E. cdlent photographic cover:1ge of m:un brroups) {V.,luJblc tc
H.illam, A. ( IIJ9fl) ·tu o,u/im· ~~f flhau cro::t>ic B"!liCtll/r<lJllty. P3ul. .R.. . ( 1980) Tlte Natural History .if Fcmils. Wtliln<'r , P (
Oxford UnivcT'Iic. Pres~. Oxford. 'J~Zeidenfeld and i olson , London . (Simple but inter- Ill cNtty
I !J.rbnd, W .B. (ed.) (I 'J76) 'nu· p,,sil Rm1rd - t1 esring) 1n l.thl .me
~)•mposium H•itlt D~.•wmrlllntimt . GcologicJ I SoCJery of P1veteau. J. (ed.) (1 'J52-1966) Traite dt• palcowvlogtr
London. London. ( omains umc-range d1agr<.Um of all M..ts on, Paris. (The French 'Treaci~e', sbghtly older
fo il orders nd ch.ms showmg lhvcrsicy fluctu~tions than the Amcri an Treatise lltJ lrrvcrtcbmft• Pt~lroutology lndividu1
through rune) bur of very high quality)
H.1rland. W .O ..utd Ammrong, 1-t.L. (11J9!1) A Geolo,!(ical Raup, D.M . and t:u1ley, S.M . (1978) Pti11ciples ~J t o.ti~, G.Y. (
Time ,\'cafe. .unbnd~e Un.ivc.:T'Iiry Pre s. Cambridge, r,!lcomolo,S{y. 2nd edn . freeman, s~n FranCISCO. Slut (t n\
253 pp . (Es enoal :md up-m-dare) (Excellent textbook emphasizing approaches and cnn- ( >uartrrly jtJ
1-ledberg, H .D ( IY76) Iurmwtiorhzl lrati~mphi( Cmdc. ccpr~. but not morphological or srracigraph.ical detail ) I t1 "-1'1 . ( ,
Wilcy. New York. ( fftcial guide to ~trarigrnph.ical R..idlcy, M . (1985) Er•olutitJn mrd Class!firation. "nlr h :lltl.'). \t .A.
procedure) R..;_fomratitlll cif Clndism. Longman, Harlow , 20 I pp. tnno~nc~
lledgpeth, J W ..md Ltdd, J. . (llJ57) 'ltt~Ui\ C 011 Mari11c ( ' !car trcatmcnc of dJffcrcm ~pproachcs to classifica- •.
Ph,.~. Clll'lt
Ewh'~)' <11111 Pnlarocroh~t:l'· V Is I .md 2. M t' rnoir~ of the tion) I nd.ty). A
Geolot,>icJ.l ocicty of Amcri ·a No. 67. Geological Ry3n, P., Harper, D .A.T. and Whallcy,J. . (191J5) PAL· ( here tla
Society nf Amcn ca. lawr nee. Kan. (Sr.andJ.rd text, TAT: ser's mamtal mu! case ltistm-ies . 'tatistics jtll llil'\tch, F . .
with pJpns .md anno ted btbli graphy on ecology of pnlaeontologists. 71 pp. and dis . Chapman & Hall, bttHhK ,1.,
J.llliving dnd os~il pbyl.l} London and Palaeontological Assoctation. fl.tf<lt'OIIIO/o(
Henntg, W (1lJo6) / 1hy!ttQ<'Itcti, .'lysrrmarics. Umvet"iry f Schafer, W. (1972) Hcology allll Pala occolngy <if Marim l:uv.1lvc ,ur
IUinois Pre~>. Urb.tn,l, 111 ( )riginal d.tdi~m; second E1111iro11111£'11ls (translated edr.) (ed. G. Y. Cra1g) . liv~r l,mgcnl'b, P .
edition pubh~hcd in I G74) and Boyd , Edinburgh . (Standard work on Recc11t .'i)'lllill'.l·is (
Hughc , N.F. (ed.) (I Y75) O~~tauis ms 1111d Conrineuts Nort:h Sea environments with applicacious for p.tlaeo- Bl.t bwU,
17trt111,1ilt l'itHC peciJI Papers 1n Palaeontology No. 17. ccology) l l.lrf'l'l. I) .l
Acad.:mk Prc~-;. lundun . (23 origin.l.l paper.. on dj~tri chopf, T.J .M. (ed.) ( 1Y72) Models in Paleobrology. lrtllllr/111111
bunon of f.mn.lS; many cllJncd on global map~) Freeman, Co per, an Francisco . (10 original papel'l, ,or/t~trd, I
Hugh<'. N .F. (IIJHY) h 155ils fl( lt!f<m11nttotr: ew Rerordhtg many ~igmficant) Monngrap
allll ...,tmtal Condathm Tcrhmqu<'>, ambridge University core, R .H. and West, R..R . (ed) (1976) tntctme illld H~ per, I ,A
Pres. , .tmbrid~e, l..j..j.pp. Clnss!fi trlimt <if Paleocotw11u11ities. 0 wden, Hurchinsm1 ~)'\tl:lll lt)
Jo>•sc , K.A. and FndJy, A.E. (11JH2) Problems of Phylo- and Ro. , troudsberg, Penn. (11 original papers) St'III'IY tl..f 1
,I!CIICiir Rt•fuiiStntction. Syqematics AssoCiation Special Skinner. B.J. (ed.) (I Y81) Paleoutolog)' and paleoenfliron- 1-1 'IIC), j -L.
Volume N . 21, Academic Pn:ss, ew York . (I l nrcnrs: Rradirrgs from Amen'carr cie11tist. K3u mann. Lo p•11m de "
p3pcrs) Altos, a1if. (21 collected papers) dad.u.m)
Laporte, (1968) ...Jmimr bwirtl!rmrrrts. Prent!Cc-11~11. Smith, A.B. (1994) Systematics mrd tlte fossil rerord. I h•ll.md,
nglewood Cliffi, N.J. Domtlll'lltillg Pt/0/utilllrllry [•atlem$. Blackwell, Oxford '''· (ll.m
M Kerr w , W .S. ( 1 78) Tire Ecolo,~)' ,,f Fossils. (Lucid explanation of dadistic methodology) ;.:olngic
Duckworth, London . ( onununiry reconstructiOns) ylvestcr-Bradlcy, P. . (cd.) ( 1956) ne pecies Conet·pt i11 pp 1-·D.
McKcrrow , S. and Scotc ~. C R . (cds) ( l Q90) Pa.laeozoic Pt~laeO/IIIIIIIgy. Sy rcmacic J\ss ci~tion, London . (Basic lhoh, (1
pJlaeogcot,rr phv .Uld bJO~euw.1phy. Gcolog~cal work of several papers on palaeomological taxonomy) lltd-7
Society Mcmo1r No . 12. pp . 1--·W . ( bn y valuable Tcichert, . (1lJ7 ) Treatise 011 ltwerrebrare Pnlco/ltology,
papers iucludmg Pa!Jcuzo1c we rid nups) Part W (Suppl. I). Tmcc FomL,· at1d Problem alien.
Middlem.i ~. F. . , RJwson, J>.F . .llld Newall, G. (eds) Geological Society of America, L1wrcn e, Kan .
( 1971) hmtl<tl Ptt'l'itta•.r ;, Sp<~a· mrd Ti111r. cc I House Thompson, d' Arcy W. ( 1917) Q,, Grot/llh awl Ftmu.
Pr~ . Livt."rpuol. (13 ~)[igin.tlfHpc·rs on t:1unal distribu- c~mbridge University Press, Cambridge. (Ind1vid-
tiOn ) uali tic cla ic work on physical laws derenmning
Bibliography 25
~owth : an abndgcd cd1t10n of thts ~ook " J edm:d by K.tuffimnn, E .G ..md Scott, R . W (1976) l:h.~1C conct"prs
j.T. Bonncr 111 I W!l and al, J published bv '.1mbndge of community ecology .md pabeoecology. in St/llctiiYO:
Umvtr~Hy Press) and Chmi/icatiOII vr w.
Jlaii1Cilftllll11lllllifies (cd' I - ">corr
Th!1~>on, G. (llJ7\ ) lJ_/c 111 the :c.1. World Univn. try .md R.R. We,t): I owden, llutchinmn .md Ro~s.
Library London (S1mple, \ dl illu tr.1ted text: deals Stroudsberg, Penu., pp. J- 2R.
\Vith COllllllliiUt)' >tntltllre) lockk•y, M. (1983) A rcvi..:w of brachwpod domtnJted
;,
V.lien tine: . J. W. ( 1<J7 3; EFululwlltlT)' H1lt1Ct>aolog)' •>f tiJc p;~ l acocommun1ttes from the type Or oVJcian .
.\Iari11r Biosp!lm·. Prcnnct·-1 bll, Lngkwood Clitf>. J H.!lcreo•IH1111lg}' ZCl , 111-45. (T 'nnin 1 gy for pJlaeo-
(VJiuablc te ·ton evolution <tnd ecolu)!;y) community structure)
i. Willmcr, P. (I 1N!I) llwtTtl'l>r<llc Rdallc>n:;/up:.. Cambridge Moore, J, and W11lmer, P. \ 1997). Convcrgcm evolution
Univcr.icy Pre~~- CLmbridge. -itiO pp . (Modem, very .in invertebrates. Hio•h!f!im,' RL'VI l'li'S 72. 1-60.
readable and up-to-d:ne tre.ll:meut) Ncwcll. N.D. (1972) Tbe evolution of red!·. Scit'llltjic
.~mcrim u 226, 5+-65. (lnf(mn,tbvc short pap r)
r Palmer, A.R. (IIJ65) l:3iomere - a new kind of ~trau
'· Individual papers and other references graphtc uniLjoumn/ q( Pc~luclllllc>l,~~y 39, J.J<J-53 .
Palmcr, A.R. (1984) The biomcrc problem: evolutiOn of
if Craig. G. Y. ( 1954) The p;liJc:oecolob') of the Top Hostc .m idea. }twmal c?t' Palconrology 39, 14Y-53.
Shale (Lower Jrb0111ft:r u) Jt .1 locality near KIIsyth. Pcter;cn , . '.J. (191 R) The sea bottom .1nd it produc-
Qt~arterly Jollmal.if rht• Crolo_~Lml SoCief)' •?f l...tmdou 110 , tion of fishfood. Rcpom of rhc Dani h B10lo~rical
103-19. ( bmc pab~l ecolugica).,rud •) tatJOn 25 , 1-(!2. (The first ciefimtive ~umtmry ofben-
e Fort~y. R.A. and JdTcnc: ... R... (19Kl) Phylogcny and tllic commurliaes)
1y tem.Hin - .1 compr nme .lppro.Kh, m Pnll>/em.s (~( Pickerill, R.K. anci Brenchley, P. (JY75) The appltcanon
Plr)•lc~~CIIL'fi( Ruv11stnlctio11 (t'ds K.A. Joysey ;md A.E. f the community concept in palaeontology. ,\I,mtime
Fnday) Acadentit Prt:~!>. t:\\ York, pp. 112-47 erlim nfs 11, 5-H. (Tt:mlinology and apphcation)
(Where cLtdisnn 1s .1 d 1~ nm u.cful) Rudwick. MJ.S. (1961) Tl1e feeding mt:chanism of the
FUI'ilCh, r.T. ( 1')77) oralh311 (Upper jur.'L~Sic) nurine Pcnnian brachiopod Pr~>ndl lllt~(cllin . Palat'<>lltolo,l/}' 3,
bcnthlt $>OCJ,Itluns from England md Numundy. 450--71. (paradigm approach)
PaiMollt<l/o_l!)' 20 , 337-RS. (P:tLH_m·colob')', n1.11nly of ulith, D.B. (1 98 1) The Magnesi,m L!me'l ne (Uppc:r
btv:tlvc conunumne, _l Pcnnian) reef complt:xes of north-east England, m
Gmgcrich, P 11 . I <J90. StrJtopheneuo, 111 Pn/,u•vbi,,ft\rJ)'; a Europe1111 Re~( ,\lodels (c . D.F. Toomey), ' ociety of
S)"llilcsi.< (eds D.E.G . Bri!!b>"i and P.R. rowther). E onomic Palc:ontolngiHs and Mineralogists pecial
Bldckwdl, 0 ..-ford, ·L\7-42. PubliLariun No. 0, Tulsa, Oklahoma, EPM, pp.
J-!Jrpeo, IJ.A T (14!:(<1) Hrm/'"'1-'''d,· jr''"' tile Uppu 161 - S.
:J.r,/mdlic~u >IIM'-'>""' (()rtl01•1da11) 4 t/1~ Cin•<lll Disrrict, Thoma~. R .D.K. and Reif. W.-E. (I j'JJ) Th~ skdeton
'• ~1>tlaml. P.m 2. 7'l-l :!H. P.llaeontographJCal :">onecy space: a fmitc ~et of organic Jesigm. Et•oltHiou 47,
Vlonographs (caxono1111 pro cdur referred to Ul text) 34 1 0.
d Harper, D.A T . ami Ry.an, P ( 1990) Toward> .1 statistical Tho~ n. ,_ ( 1957) Bottom communities, in Trl'alrsc ""
n system for palJconmlogist~- _la11mal <!!" tl1c Geolo_~i,al i\lan'm• Ecolo_f!y and Pa/acoccolo,!!)'. Vol. 1 (c:d. J.W.
Ot1o'l)' qf u•llril>ll147. lJ3S-4H. (1~ . ..::nual re.tding) Hcdgpcth), Mcmou-., of chc .eological ociety of
Henry, J.-L (llJH.J) Au,dy~c cl.HJistiyuc: et Trilobites: n America N o. o7. Geological nety of America,
pomt de vue. Lcrlwi11 17, 61-6. (Use and lunitation of Lawrence, Kan ., pp. 461-534-. (. tand.1rd \ ork. invalu-
rl.Jd1~m) able, well illustrated)
llollond, Cl!.. Audley- h.trb. M . ;,, Bas\ctt, M.G. L't Ztcglcr, A.M., .ocks, LR..M. and Bambach, R..K
j ,1/. (I'J78) A !-.'lliJe m .. trangrapluc procedure. ( 1'16H) The compos1tion and strucntre of Lower
Geologtcal 'onety of London ~p ' nal Rcporr No. Fl, Silunan m. rine commurut.ics. urltma 1, t-27 . (The
pp. 1-U. (!Jwalu.Jblt' modern tr'.ltlllcnt) first major work on Lo\1 er l>ala zoi communitu:s;
llcub. F. (t 977) Evnlut1on .md unkt·nng. uellce 196. very well illustrarcd)
1161-7.
/,
I.
I.
g
wer which
Evo ufon and the fossil 1! e~ not pc

h 1wevcr, n•
rnuch and at
11;1[0111 • bi
record . nJ
Org;HH OlS
evHkncc in
f
cytolo~
~hen brou~
to .1 them;
[)obzhan k)
molecular biology arc transfornung :tnd adding tu wns e.· cp l
this ~ymhcsis, and new vie' on the nature of the The rcn."
gene seem to be changing our whole onception of b ~elf-~ le
Amongst .tll the ~ tence concerned with organic how organism evolve. quently disf
evolution it i nly p lae neology that has the In the following tex1:, tbercfc re, l pre ent fi1 . tl) a and t:mh-:)
umque perspet·tive of geologic• I time . 1t i~ the rock simplified account of das ic neo-Darwinian evolu- needed hen
record alone th:H prov1d s an hi torical perspective tion:try theory, ;1.imed particularly at students of Although
for rhe study of evohmonary vent , ,md thi time E:mh science who may only have a lim.iccd back- eh· father 1
dimension could never have come from any other grou nd in biology. This includes some information Tre m ny
omcc. Accordingly, the input of palaeontol gy t on the impact of new information from molecular cllJ t ammals
cvoluti n theory has be n in under tanding the hi - genetics on e olution theory. Thjs section i~ not o is of time
tory of hfe, in mterpreting pJttcrn of ev Jution intended to be comprdJen ive, nor due it pretend tTnm pre-c
(e.g. adaptive radi.1tiom) Jnd line~ of descent, and. to be a guide to all rcccm developments and dis- L arwin hi1
unp rtantly, in .1s. es~mg r,ttc~ of evolution. Now it coveries. Fuller treatments arc readily available else- wh~eh co ul,
has t be ;Jdmirted that the fos. il record is incom- when: viz., impson (1953), Mayn.1rd Smith (1975, f modem
plete, and to interpret it can in some imtanccs b' 19fl2), Mayr (1963, 1976), Dawkins (1986), prinutive ,1
'like trying to re d a diary with half the pages Dobzhansky cl al. ( 1977) Gamlin and Vines ( 1987), though rcfi ,
missing' ( hcldon, IIJSR). Yet it ~till rem ins an Banner (1988}, Endlcr and McLclbn (1988), he~ at rhe h
immensely rich source of primary dat..1, :md an Hoffinann ( 1989), CampbeU and eh pf ( 1994), wrcm ined
give, if re. olved finely enough. a f:1ir picture of evo- Maynard mith and Szathmary ( 1995), Futuyma 1V ·-fingcrt.:
lutionary events that ctually took pl. cc, however ( 1996), Strickberger (1996), Ridlcy ( 19%) and bratc , for 1
long ago. others listed in the bibliography. whil~ Valentine ofways. Tl
The smdent of pJlac ntology needs t have some (!973), Hallam (1977), Stanley (1979), Cope and of marine
background in bioi gtca l evolutionary theory, other- kclton ( 1985), Levin ton ( 19R8) and Skdton ( 1993) the hoof~ <
·wise he will be m the position of the man standing arc more directly concerned with evoluti n and .1r' .111 vari.
by the roadside :md " atching the car. go past but the fossil record. ln the econd part of tl1i chapter d1 er. ity 111
without any 1dea ofhm their engine. work', to use there i a more extended a count of what the fi ssil the Ga.l pn
Brouwer's ( 1973) ~:.rraphic simile. For the fos il record can tell us about the natur • of evolutionary en ounterc
record cannot give much infi rmarion on che mech- change. l~33, led
;mism of evo lution:try change; this has to be pro- dtvef'\ity in
vided b) genetic. , cytology, molecular biology and group of bi
p pul cion dynamics, building upon rhe origmal riginal pa
conceptiOns of baric Darwin. Many years ago the ~pccics had
tirst real multidi~ciplimry amalgam of data was pub- ends'. Geo!
lished as Erto/rttl!lll: tlu: !lfodem yn//l('sis (Huxley, t.111t tO CV•
1942). rom this highly uccessful attempt at ~ eld- The theory of evolution links together a multiplicity hm the 1
ing together infom1ation fr m various sources, the of biological phenomena md is underpinned by all ( . g. lungfi
neo-Darwmian synthe is t.tkes its name -and this is the evidence of the geological record. lt r ·main a mdicated Cl
a usdi.tl starting pomt. But re ent developments in theory, not a proven fact, for tbc inunense runt.'!>cale subsequent
Darwin, the species and natural selection 27
over whtch evolution.uy cha ng~s have token pbce few species to small ;lrca.-. only where each had
doe1 not pennit then wren observation. There is, become adapted to its own environment.
flowevcr, no other Lheury which encompa ses so Darwin was particularly mteresred in ~elective
much and Jccords with the cvic..lcn cc of comparative breeding of animals and plant~ unckr dome ·ricarion.
anatomy, biochenmny a11d phy.;iolot.ry, of genetiC"> He reabzed that the present great variery of dogs and
md cytology, and ut the relation hip~ between c:tttle had been produced in only .1 ft:w thousand
orgamsms percctvcd by ta..xonomy. Wherc.ts the ye~rs from only one or at must a few ancestral rypcs.
cl'idcn e m some of rhe~e fields is Circumstantial, He concluded that a great potenlial for 'descent
whrn brought together and mterprcccd it builds up wirh modification' must exist in all animals and
ro a tht!ury of formidable conscyucnce, and ,1~ plants wluch could be speeded up by such :mificial
D bzhan Icy has said, 'Nothn1g in biology make" br~·eding. This led LHl to the belief th:n similar
1ense except in the bght of cv Iuti m.' processes, tJ10ugh probably on a ~lowt:r tiJnc scale ,
The r cent facile artcmpl~ t discredit evolution had opcrJted in the wild state; in other words 'lnt-
f hv 1elf- ryled 'cre.11:1 0n ~cicnt1sts' have been so cl - ural selection'. Thu~ breeding experiments arc the
quently dtspatched b Dott ( 1%2), ]{jtcher (I 982) foundat:ion of classical genetics, where the mcch:~
and t.1nley (19H2) that no urthcr comml·nt is ni.;m of heredity is understood in terms uf 1tS effects.
needed here. Darwin was also concemed with palaeontology
f Although harles Darwin is gcner.1lly regarded ,1 bur he found that the fo. sil record w ;1· ~omcwhJt of
rhefather of evolution thcury (D<~rwin, 1RSCJ), there a ills:~ppomtment in supporting the case for cvolu-
1 1\WC many pre-Darw1111:111 ~Lientists who pmtulated ttonary change. He had hoped eo find cv1dencc
r th.\t .111un.1h ;md plants l1Jd ch,tngcd over long peri- of gradational change bc:twet:n :mi.mal specie~. of
t odl nftimc and that new type of\pccies' had ~1risen 'infinitely numerous transitional links' connt"ctmg
:1. (rom prc-cx1stin~ ou .s. The~c c;1rly workers ~'nd ancestor. to descendant~ .md of str:1tigraphica11y
OJr\vm h1mself 1dentificJ many dilferenr point arnnged sedes shO\ :ing 'descent with moditication'.
wh1 h could be regarded a evidence for th~: origin In 6ct. he d1d nm .tind " hal he had expc tcd.
of modrrn specie from pre-exi,ting and more Darwin assumed th t the imperfection. of the r ck
rrinmive ancestors All these dfC a cepted today, rec rd and the lunitations of k11owlcdgc at rh at rime
th ugh rdl.nrd and , dded to. Ta nnorny, ,1s always, \ ere the factors resp n ible. He WJS indeed partially
he~ ,Jt the heart of cvolut1on,1ry rhouglu, and losdy correct.. but to this p int we shall return btcr.
mtcmnncd w1th H i<> comp.trat•ve anatomy. The f the prt>-Darwinian evolutionists the mmr
a ftv·-fingered, or pentJtl,tLtVL lnnb f higher vcrtc- pr minent WJS the French naturali];t Jcan-1:3apti~te
i brJte\, for e ·ample, h~ts been mud1fled in ;J variety Lamarck ( 177 4--1 H29), who proposed I ng before
uf way . fhe grasping h.mJ of primate~, the Ai pp er<~ arwin rh:~t all living organisms had oribrinatctl !Tom
of marine mammals, the wings of birds and lxtts; primitive ;mcestors and rl1at in the slow procL·~s f
rhe hootS of hor e - the Jll look dissimilar. but such change had become ad.'lpted tor liv:ing in p.lrtic-
1r~ all vari:mts on ;J common theme. Likewise thl" ular environments. The concept of such adaptation
r dive~ity in strucmre and funct!Un f rhc beaks of 01iginated with Lamarck. He apprcci:~red rhat in
rh~ alapagos tind1es (CrMpi:::a), which D~rwin order to live, an.imab have to be effincntly acLlptcd to
t'ncountcrcd dunng the voyagc of the Beagle in all the physic, 1 and biological parameters of the ' nv:i-
1833, led him to say • cdn~ thi gradation and romllents they inhabit. fn a seJISe, of l'Ourse, an ani-
diversity 111 stntcturc 111 onc ,mall, i11timatcly related mal i 'all adaptati n'; it ha~ to be anatomically,
~up of birds, one might rca.Lly f:mcy that trom an physiolog1cally and trophically .tdaptcd to it environ-
ong~nal paunty of bm1 in tlli~ arch1pebgo. ont> ment, and it is critical to evolution theory to undcr-
>pccirs h:1d been taken and moditicd for dilfcrem sraJ1d how such adapL.won~ came to be.
cnM . Geographical distribution was sccn as tmpm- 'While appreciating the importance of ad.lpt::~r-ion,
onr eo evoluti nary th ught m other w .l'i~ too . howevc::r. Lamarck linked llis insight with ·ome
{ Thus the ex1stence of 'relict' and isolated ~pccies concepts no longer believed to be tenable. He
l (e.g. lungflsh) in d.iffe rem pam of the world ~urely believed that adaptation had come about through
mdicJtcd an original ..vidcspread JI1LCStrJI type, a some kind of internal driving torce, ::t ·vit:U spark'
1ubequcnt poplllatJ.Otl coll.1p~c .•md re~tnctmn of a which made ammah become more complex. He felt
.28 Evolution and the fossil record
th.tt new rg, m must ari!>e from new neL·ds and that the nao1re of variation and heredity W,l\ l.lrgel 1h I P.
the e 'acquired character~' were mhented, a in his unknown, s that hi~ icw on thi~ wen~ spcculao1c ltddcr n I
cLmic pmtulatc tlJJr the 11L'll- nf tht· 6'1rafle had. and imufficicnt f r the theory to be . cen ro work. pi
become longa m re. ponse tu a 'need' to reach The pioneeting work of the Au. tnan 111~111\;
leaws up on thl· trle. he the1 ry nf inheritance of .regnr Mcndel in 1865. and of the later ~chool tll
Jcyuired characccr' 1~ not htghly rl.'g;micd nuwaday. T. H. M organ which began in 1910, h1id the foun·
.md is generaily umc~t<lblc (Jithough there i~ datioll for genetic experiment and rheury. It wa'> thil
ome evidence of a kind uC gcnCtlC li!cdback from UMt ~upplied the necessary undc~tanding f hered-
the environment operating tu produce apparent ity esscnnal to the amplification of D · rwm \ theory. 1.11r,,
'Lamarcloan ch.tngcs'). Danvm, on the other hand, 1 (\l\111~ W1t
pn: VJded .t logtcal and teMabk theory f r eve lution- lti tbl.."
ary eh nge: one th .lt has 'to d the te-.t of time and Inheritance and the source of variation 1 ,mgcd 111
pr vided J ~tarting point r r later developments. \trun~ H gc

The full title ofDanvin\ lllaJor v ork of 1859 w~ In the cells of all eubryotes (all organisms except for sOilll', ,IJH.J •
011 t/1c nngi11 q_j speciL's by 11/t'tuts l,J 11111111'111 srlario11, or Vlruses, bactena and cyan bactena) there i~ J p~r d1r rTu
rile J11L'.I'CI'I'uth,u tl})at,orrred races in thl' ~tm.~~/e ji.1r /ffi•. n ucle us (Fig. 2.1) containing elongated thrcad -hk~ J~.:~' rih rh
The main pomts of the theory arc straightforwa rd. bodies, the c hro mosom es , which are made of pro- ;m th · pr
tein and deox-yribonucleic acid (D N A) . g u e tic eo
I. Animal ~peci ~ reproduce more rapidly than i:; tlms and i
needed to maintain their numbers. Animal popu- smooth endoplasmic reticulum twmng f
lations, however, though Aucruating, tend to I mf nn li 1
remain stable. (Here he w.t~ mflucnced by the ,d hat J
Engli~hman Malthu , who held written on thi ntH: k ut1de
ubjcct some years ·ar!Jer) vacuo le mitochondrion lt) l Ill ,1 ki·
//'
2. There must therefore be compenrion \.Vith.in and kttn:s, th~
between spcc1e' 111 the 'struggle for cxi~tcnce ·, for wh•Lh the~
food, to r living sp.J.le J.nd (within n1embcr~ ofthe- \:Jlnnu us.
~ame specie~) li1r lllJ.tC'> , if the character~ that indi- dltl·rminc:
vidu,t1s bear Jrc to be tr.ln ·m ittcd to the next gen- ,nnuo ao
eration. tngl'thcr t
3. Within ~pecies all J.11imal vary, and this variation ~omb111ati
i~ inherited. p trtJ ular
4. fn the ~truggle tor hfe tho e ind1vidu.us best tittcd en ugh p
to ~un ivc in .t pJrttcular environment arc the s ·,tl·m tu
nes to live and w rcprodu c. The other arc lmk them
weeded out m the mtcmc competition. The pr rem.
favourable h.n-.Ktcristics th,tt nuke uch survival !'rote in
po sible are inherited by future gcncr.ttiom, and 1.t the rib
the an:umulati n of diHercnt favourable chJrac- m . ns th .
ters leads to the separation of ~pt•cie v ell adapted lllt of tbt
to partiCular environment~. Thi~ i~ what Darwin I· 1r this
called ' narurJJ sclccnon'. du 't' ' ,1 !>i
but with
All tlm seem\ logical enough. though Oar.: ·in's early fat d oplet r placmg
critics, Mivart for instance. argued th.u Darwin lud 'transcri
not reall · shown how favourable characters were I NA) i~
actually accumulated, only ho tho~e animals le s nucleolus ribosomes pre~umal
fitted to their environment Failcd to urvivc. In this and att:H
rough endoplasmic reticulum
rhey wen.: not unsound, for the mo~t . criom weak- ~1mplc p
ness of the theory. as prescnrcd b Darwm, wa thar Figure 2. 1 Eukoryote cell structure, with orgonelles. them$d"
Th~ DNA molecule forms J long. twisted. ~piral thar gene\ themselves are made o[ two kinds of
bddcr of which the 't pright~' consist of alternate component :1rranged in cries. These are exons.
blocks of pho\phonc ,lCJd and the pcnto. e ugar wluch code for proteins, and introns. wh•cb do
~1eo\ynho~e. when.:a' the 'mngs' :~re m.uchmg pairs not. When the nuclear DNA ts tramcnbed to
of rclanwly sm.lil unm, rhc nucleotide bas~.:s. The\e nuclear RNA it retains the org;mizJtion of the orig-
b1e~. in D A, arc ;1c:kmne, gu.tninc. thymine and inal D A, bm when nuclear RNA 1s rcproces ed to
cvtosmc. They .m' .machcd to the pemosc ugar messenger KNA the i.ntrons arc lost and the exam
umts and project in'i ard~. linking up together m are spliced togetht~r. Tlm does not ahvays take place
pair;. Adenmc c.m onlv pair with thymme, .md in the original order, however. and uch cxon shuf-
ntosine Wlth guanine . fling may be the basis of r:J id c\·olunon of protems
In rhr chrommnmc the DNA ~rnnds are thcn1sclvcs in new combin:~tions. Provided that
lmngcd m dl\crete umts, the genes, which arc these are ti.nlcttonal, new gene y tt'ms 111~1y anse
,trung together along the len~:.>th of rhc chr mo- through comp.trativdy few uch shut11mg events.
mc, ;md of" hich rh 're may be everal hundred and i.n short periods of time. Tlus IS a newly under-
pt·r chr mo~omc . The genome 1s a tem1 used to toad phenomenon. but may hJve tmportant const'-
clt~mbc the \ hole genetic complex. The e gene quenccs tor evolution theory.)
.ll't' the pnmary unit' of heredity, carrying the When messenger RN A arrives at the surface of a
genetic rode, whi ·h is mvolvcd in produnng pro- 1ibo~ome it does not torm protein rhrectly , but
tdm and m dm:ning rhe devclopnK·m and fi.uK- through yet another mtem1ediate molecule, transf..:r
uonmg uf the whole organism. All the nccc~~ary RNA .•md when this complex process IS complete,
mfomu110n tor thc~e end~ is earned in the D A. the re ult is a protem seq~tence, coded for by the
Jlld what t important i the 'equcncc of the four nucleJr D A, the tramter RNA me.mwlule rerum-
mtckotide ba\e~ J.long m length . Though these ing to tht' cytopl:tsm.
hm1t .1 lind of alphJbet comi,ting only of four The understandmg of protem syntl1csis has been a
kttm. the number of possible combinatiom in major mumph tor molecular biology. but the mole-
which tht') c:m c:J<.-ist, codlng for specific proteins. is cular p:tthways that lead from genes to actual organ
cnormou~. Tlus sequence of nucleonde bJses nd characters arc ve1y complex, and at present
dt>rcm1tn ·~ the ~cquencc 111 whtch any of the 20 largely unknown. How the protem~ and other com-
jnltnu acid found iu living orgamsms :m: strung pound wh1ch arc produced arc orgamzed and built
t~gctht:r to make proteins. It has been shown tb:Jt up into fi.mctJonal organs and whole bodJe, remain
wmbinatinns of three b:J$eS acting together code £; r one of the mam tasks for molecular biology for the
prncular amino ;Kids . :md there arc more than future. Some prot,•Tcss has :liready been made; tt t
'llllll!lh pmsiblc combimciom in thts 'base-triplet' now known that some genes are 'structural', and
wnn to m:tkc all the btogemc .muno .lCtds, and to concerned only with the synr.hcsis or materials,
~nk thrm up m the right order to m:~ke J speCific others are 'regulator' genes, which control and
prutru1. orga1uzc the compound and dirt'ct their building.
Proteim Jrc 'Ynthesin·J. nnt in the nucleus, but uch genes relea~e · hemical pr ducts wluch ~tart a
,,, thl ahosomcs in the cclluLlr cyt pl,tsm ..md tlu whole host of complex reactions. Ln some kinds
mr.m\ rhlt .tl! the infi.mmtion h.ts to be tramfctTcd of development the genes are switched on and ofi·
IS uut 111 the nucleus to these ~tte~ of protein synthesis. in particular sequence, releasing products wl:uch
Fm thi~ to take place the nuclear DNA first pro- react together in rynthcstzing complex molecules.
J,~.·c,a '' gle-~tranded copy ofitseH: nuclear RNA, Structural genes can be accivatcd and deactivated
hut wtth ur:~nl repi.lcmg thynune and tibose sugar when needed; evidently the i1utial stimulu. tor the
n:plmng deoxynbme . Following thl~ proce ~ of ~witching on of sttl.1 cmral genes is given when a
tran~rription'. .1 ti.mha molecule (messenger sensor gene re eives an appropnatc stm1ulus.
Rt'.A) IS tanned, wh• ·h move~ our of the nucleus, imple organisms such as bacteria and fungi have
ftt\lllll'lhlv through pores m the nuclear membrane. sets of adjJcem genes kuown as opcrons, coding tor
<n I .lttJchrs 1tsdf to the nbosome. (This is not a a panicular metabolic pathwJy. These c.m be
unpl~ proces , however. for at this stage rhe genes sw1tchcd ofF and on together. Ln lugher organisms,
·h\'111\~lve~ arc modtfied. lt has been shov.m recently hm ever, genes which control difrerent parts of a
)0 Evolution and the fossil record
coordjnar ·d progr.~mmc may be ~cJtrcred :tround in number being conventionally defined a~ 2n . /'7'_.rr,._.-.... ~......__
mall groups. bm on d11ferent chromosome~. When rhe body (somatic) cells of .1n .1nin". -~----
Genetic:tlly identJc.Jl org;m1sms reared in different dt ide JS th e org;mism gr ws, the chromo on··· .. ~.:;:-:::.··:··'. ·· ·
environments will noc develop to exactly the . c1111e
fom1, as 1s w1tne~~ed by the different -lf pe.u:111ce of
divide by longJtudin::tl tission, and each ' daughtc · .// -\ -:f"S~·;('
. . .;,,
cellmhenrs ::tn exact copy fthe .. hromosomes J.ti "F \\--f.• : '· <..
-
"\ \
vegetable~ gro\ n 111 neh and poor s01lo;, respec- genes in the parent celL Thi~ proces~. knmvn ( \ ) . ) '
1
T:'-h
tively. The inherited genetiC ll1J.teJi:u in .my org:m- mitosis ( i~. -.2), ts effectively the same for JLI t i · ; ; ....... ._ ·~ ·~"'( ·
\ • ; • 1r \
i~m. known :Js the genotype. i~ reacted upon by d1e somJtic cell~ in an one body. \ \ -.( 1 .··~{-~ . . ·
environment (prob:tbly through semor genes) to When :l cell i~ not divirung, the chromosomes a . \ ...: i:J··.~:·"::>~/.
create a devdoped indi"'tdu.d. This individu:1l, the L1.r~c~y invisi_ble; this is_ k1 w~ :t~ interphJse A~ r"'"'( ·. ·--:--.-·-< .
product of both her •dity and cnvuonment, IS the d1VJSIOll begms, bwlogtcal s~nnmg reveals the chr '··•. · . ~
p h enotype. mosomcs, each aln:ady divid.:d into two chronutt -......_
. omeomc single g ·ne\ control smgle rhJracters. hdd together by a mall sphencal centromere, wh1• interphase
More often character' arc polygenk; th,lt 1 , m:my does not stain. The fir t ~tage of cell divi~ion
genes. e.1ch of ~null 'ffect, contribute to the bio- prophasl.', where two entriolcs move to opposi
chemical pathways that rc~ult in the fonnation of 3 poles and J 'spmdle' of protein thre.1d .tppe;
p::~rticubr chJr.Jctcr. Ag:Jin, some g nes ,trc between them. At metaphase the nu clcJ r memhrJl /""~-:---:~--"'"'
pleiotropic; th.1t ts, they affect sevcr.u characters v::~uishes and rhc chromosomes line up along d/ :'~· ~·.
smcc the s:une gene products may be mcd in differ- medi:m pbne, each ~plirtmg in t>vo o thnt eJ1 ·· ··· · · ·
ent biochcmic.1l pathways. A phenomenon which at chrom.1ttd becomes a daughter chrom ~ me wnh ;· ;
first sight o;cems puzzlmg '' th.u much f the 0\\'11 centromere. Then. during an;1ph.1~C they p1 : t ~j ·,/.'
I. .JI h '~ \ iJ' ,' ._ .. • _,.,.
genome const.;rs of rt•pe.lted DNA segments (multi- apart '0 that eac I1 ounLUe of d:mghtcr c romosom ,;_~ ···"':-.:/-"'
gene ~ll1lihe") with anythmg tl·nm r, o w 'evernl move~ to the oppo~itc pole. Fin.uly, at tclnphJ.~c.
thouo;and copies nf th • <;.lme gene. Though 1t may new nuclear membrane ap pear~ round each q~r
appear th,\t mm h chrom s mal nutcrial '' redun- chromosomes, the spindle vani~hes .md the c1
dant, there ould wdl be 1mporr.1nt evolutionary divides into two tdentical daughter celk ... ..
1mphcatiom here, as 1s discu. cd lJtcr. n the other h:md, rhe fonnation of eggs ,..,
spenn (gametes) in the testes and ovaries respc ·
tively involves .1 very dtfTcrent process: meioi · ·
Where does variation corne frorn? (F ig. 2.3). /'J i'1
here ::tre t\vo cell division . he first of these 1 · · ' : i '
W1thin the nucleus of any n·LI in the body of rnmt reduction division in wluch each d.mghtcr c(· .
living org:111i m there'' a 'pcciflc number of chrom - inherit, only ne chrom s n1e fi-on1 3n hon1olog . ·· ·
orne. with their gene,. h~.: chrom )'-Ome number is pair. Here ::~g::tin rhere ::tre fom phases but ~ome ~ · j
a! way' con,tant ti..Jr the <;pecH.•s. I 11 hum.1ns ther~.: arc the~e are significantly different fr m their mitot · ·-
JJ pajr. f hroJIIosom s, '"'hi le in the fruit Ay counterp::~m. During pr ph:~se of the first met I telophase
Drosophila there arc tour p.urs. The members of c. eh division, when the chromosomes p::m up, tht
pair <~re all homologou ·, i.e. sinuLtr in ,tppearanc dJVide longit1.Jdinally, fom1ing t:,'Toups of four eh re
md length, c -ccpt fi.1r one p.1ir, the ~ex drromu,omc~. matids (terr:tds) but with only one centromere fi
upon which the genes cgulating <;exual characters eac h pajr of chromatids. These four chr nutid ,1
Jre locatnl. [Tiw -;ex of an indiv1dual depend upon often tangled at certain points known as chiasmat
this pair f ex chromo~ me,. In one sex the chro- and a~ they pull :~part they break and re-fom1, ea
mosomes arc identical (XX) m the other ~ex they arc exchanging parts of the Jmc length with its homo
Ji imi!Jr (XY). The ~ex chromosome carry ocher ogous c hromosome . The resnlr of such 'crossU,
genes lhan those 'peciticall · responsible for ~t·xu.tl over' i that the unp;ured chromosomes p;~ sf.2 The stages of mit
char.:tctcrs and rhm cert..1m phy~lC<ll characrcrs arc down to the d,ltlghter eU arc not tdentic:tl wli _
scx-link~.:d. In mammals the fcmak 1. CX: and the thme of the parent cell, as they . re composed ofbjf gametes trom
m,1lc XY. bur 111 birds it is the other w y round]. of e::~ch of the homo! gatiS p 1red chro111os 111~ has h,1lf the num
When chr mo omc:s are paire likl" thi~ the orga- The second meiotic division 1s Like a somJtic rfOmosomcs are un
nization i~ s.1iJ t b · diploid , the chromosome mito IS, so that the end product o the two divistofl 11Y of the parent
·.·.
, ..
\
I ..··
.~;(...:
\
.:
. I .;
..~ '
.•
.· ·. .
.. .. .
telophase
fi!JK! 2.2 The stages of mitotic cell division in a eukoryotic cell. For explanation see text. (Modified from Kershaw, 1983.)
Ll'JJr gametes from a mglc pare m ceU. Each material was exchanged during crossing over.
liJ.te ha ha! the number of parent chromosomes; These gametes containing unpaired chromo-
h, Liromosomes arc unp;urcd, and none arc Jdenti- som s arc said to be haploid, and rhe chrom some
1 to any of the parent chromosome m cc genetic number i 11.
32 Evolution and the fossil record
k.ind: RR
Lhey ar of
possessmg
pose or c
tlons at Rr
.1llde R IS
tht~case r 1
SomctiJ1
klh>wn to
types, or
gtnc whi
.md r. Allel
r is rcc css1
has the re ·
pp or pr ar
has th~.· do1
The ah
~mglc ha•
of va
tl'IIls
smglc char
.Jrt under
ent ombi
or wlute 6
th.m wouJ
.1Uele'i.
hcse s
second \."11 , ·
/
------·:
·""-
'
\
lllllbinati~
anaphase .illele . et
((
' '
y \\
} .J
addttion, 1
I . I I ab1hty by
\ ... \'-..! -" ,I
./ ~ llle . all
Polyploid)
telophase gametes
R.ec ssi•
Figure 2.3 The stages of meiotic cell divis1on in a eukoryotic cell. For explanation see text. (Modified from Kershaw, 1983.) dunng ev
lcs 1tlun
unJt s. ther
When cg__gs an: fertilized by spcn n the homologou however, are nor necessarily tdenncal, for they ll1l mg 011e p<
chromo. omcs trom different parems come together exist in a number of'expressiom' , known as allele~ advantage
and pair up. In thi proccs-; of recombination a new each of which IS slightly dJffcrem chemicilly aru, tcm1~ oftl~
genotype resulb from already cXl~ting gene~. deriving will lead to the development of dillerences in tb.
from different ~ourccs. The efft·n of both cros\ing chara tcr. it controls. Such differences may ran •
over and recombmattou t~ that vari<1rion " ithin the from the minor lo the very sub;tantial.
products of reproducoon is high . It is mainly upon Sometime , as in the genes ontrolling hunt.' ,. here p 2 a
this v:m:uion that natural ~ele c tion operates. eye colour, there m y be only two allcles. desig dnnunant
nan::d R and r io convenlional notation, R control zvgotes (rr)
liug brown eye colour and r c nrrol!ing blue C\ \ tll only b
Significance of alleles colour. Allelt:s for eye colour may exist on homolo- advantage
gotL~ chromosomes in four possible combinalion• p.1rticular
Homologous duom ;ome lie 1dc by side, RR , Rr, rR or rr. In the homozygous conclitiw tmu, the b
matchmg gene b} gene . The homolo!:{OliS genes. ,11leles on homologous chromosomes arc oftbc sam. lllde witru
Darwin, the species a nd natural selection 33
1-Jml. RR or rr. In the heterozygous condition Evolution can thus be onsidered as based upon
thn ;lre of different kmds: Rr .llld rR. An individuaJ change · 111 gene frequencies through natural
p11. ~~- si11g RR will have brown eye,, \ here as the ~ck ·tion.
ro.wssor of rr will be blue cyed. Bm th~ cumbjna-
lt'n tl Rr or rR w1ll ::~h lead to brm n eyes, for the
,1lkk R is dominant o er r ;md nusks its etl"'ect. In Mutation
dm ~·a ( r i. said to be the re e sive gene.
Somt:timc~ allcles .1re multtple; ~ome genes are Natural clecti n operates upon the b'Tcat stock of
. uown to exist in up to 11 allele\. Human blood variation inherem in the gene p ol of a population .
!Vp~~. for exampk. arc controlled by only a smgle Th genes are continually res uffied by the meclu-
~'tll~ which urcun. as th rce a!Jdf's dcs1gnared p, q nism discussed above in new cornbinnrion · or geno-
o~uJ 1. Allck~ p and q have L·qu.1J donunance when:a types. Bm if ev lving organi tns re ever to give rise
r W\ivc f the four human blood group 0 to anyrhmg radically new there mu t be a ~ourc~ of
.h dll' n:rc ~ivc homozygote IT. ~'T up A may carry new variation. Where does this variation come
1• or p1 and group ll qq or qr, wh1k the group AB from?
, the dorninam heterozygote pq. lt sometJmes happem. that a gene \1 ill uddenly
The nbove exampk~ deal with genes affectmg and spontaneously change to a new allele. Tb..i5 kind
111~~ (haractcrs, but there ::~re more complex par- of random change i known as a g ene mutation. lt
Wll of variati n wbcn t\ o r more genes control i the result of ,1 chemical change in the DNA helix:
n~e characters. Thm plum;lge colours tn parakeets a ~ubstirution or rearrangement of nucleotide base~.
.tl under the coutrol of two g~:nes wluch in dtffcr- r <1 duplication or loss of a small part of thl.'
nt combinatiOns rua} produ c blue, green. yellow sequence. The simple t kinds of mm tion :tri e by
r wlute featl1e - a ' rder combinJtion ,f colours one nu leotidc base, or a pair of b es, being
dun would he pos~ible \vith only J inglc pair of replaced by another base or base pllir. his is ub ti-
.lldc. tution, but mutations can I o arise by additwn or
Th c s urce or
vanation cro \ll1g over. rc- deletion of ba.~c pairs and this may disrupt the func-
(\)ffibination, homo- and hctero1ygosity , multiple tion of the gene. Mut.-l rions on a larger cale can
illdcs, etc. - are 11 consequence~ of u1ploidy. In a.ffL·ct from tens to thousands of base pairs, and
Jdmon, many plant' increase their potential vari- chromosomal mutation , which arc the most
btbty by polyploidy, i.e. duplication of chromo- extreme kind, may invol e inversion, rearrange-
"'"lCI, Jllowing a greater number of loci for allcles. ment, addition or ddeti n f long sequences of
p,,JyplOJdy, however, is rarely f und in ammal . DNA. These tend to have larger-scale phenotype
Rccc s1ve genes might be expected to get lo t dfects and arc more ften disadvantageous.
lunng cvolunon. bm in tact the prop rtion of alle- Although it was formerly h ld that mutations are
1~ Wldlm p pulations do uot norm.Jly eh ngc, comparatively rare events, it is now apparent (Drake
unl s there is a speCial survival advantage in posse~s cl al., 1983) that . pontanc us '1esions' are continu-
mg one pamcular type of allele. lf there i no such ally occurring in chromosome. during replication;
d1·:mt;~gc a genetic equilibrium is lllaintaincd m tbere ma.y be many per cell each day. But such mis-
tcm11 of rhe Hardy-W ('inberg equanon: pairings are continually being n.: paired by a complex
enzyme system (DNA polymer se acting in con-
p.! + 2pq + q1 ::: equilibrium jwlction with other enzyme ). This acts a a 'proot:.
reader' which picks up wrongly paired ba~es 3nd
11hcre p~ .md q~ represent the relative proportions of puts them back in the right order. Even this pr ot:.
Jorrun,IJlt h mozygot·~ (RR) and recessive heter - Kading y tem sometimes lip. up. and mutation ·
r spectivcly. The equilibrium so defined
''O!L'S ("), then are fixed in the chromosome, to have their
1\ill only be hiftcd when rhere i J decided sdeccive effects upon the descendant.
,dvmragc in po~ c. ~tng a particular allele. Thus if a While mutations have been traditionally under-
n.uticular phenotype is preserved by natural elec- stood a~ being due to DNA damage or mispro ess-
~un, the balance wi.ll altl.'r ~nd the proportion of the mg, it has rt:cently been hown that many
rllclc within the populatiOn \vill increase. spontaneous mutations, cspec1ally rho c on a larger
34 Evolution and the fos.sil record
scale, may be due to mobile genetic elements (v riis not likely that mutations will pread very ta t 111 a lsolatio1
ously named 'ju.mping genes', 'transposon ', or large population, even if they arc dominant .md
even 'molecular parasites'), wluch are DNA mutation rate are hi~h. But most natural popula- lt sh uld ne
sequences abk to lUlWC .1long chromo omC'~ or liom tions J re n t evenly dimibuted throughout their nonnJily cs:
one chr mosnmc to <111othcr. They can alter the species range. They tend to be ubdiv1ded mto gt•ogr:~pht cl
whole genetic ~tructurc or an org;uusm through relatively small unib. of p pu.latton, known as being ut r
rc:11Tangemcnt or LXpansion of it genettc m,1teriJI. ga1nodemes. Examples of such llJturaJ populanan 1bnve. But
ome tend to attach themselve to particular site-;; groupings arc r..~bbit warren , schooh of fi hes, pond may b{. eq1
others seem able ro fit in almost anywhere. The commUJuties , the faunas of endosed lagoons and, 10 g~cal i1
l.'l"OI
capacity tor 1110 ement of the e mobile or qua~i th~ human sph ere. the inhabitanrs of towm and vil- r mjcroh
stable genetic ckmcnts allow a radical and rapid lages. They tend to be partially isolated from other I gtc:tl orb
alteration of the pattc n of gene expression by J ~uch gamodemes, bur not completely so, forth ugh tion' ~o tha1
single genetic event - a phenomenon which has each gamodeme carries Its own ge11c pool there is (the brecc
clear implicatiom to r rJpid evolution :md tor the some p ssibi l.ity of cxch nge and hence gene flow eluding COJ
origin of new type'> of organjzarion. between the pools. genetic i11o
A new allele, tormed by mutation. is h erit:~blc lt is within such small natural populations that but, bccaw
and will produce changes in one or more characters favourable mutations. especially those allowing bet- location~ oj
in the organism. These change are usually of sm aU ter adaptations, , re able to spreJd. A mut:mt condi- th hybrids
., ole and of limited effect; 1f they are larger they arc tion established even m one f.1111ily may, if 11 Th re a.r
vety ftl.."11 lcthll. Mo~t mutations are neutral, s me possessc de tive advanwge, oon spread through achtcv~.;:d. i'
arc disadvant.:lgcotLS, but others 111 y le-ad to new or the populari n in the gamodemc within r btivel)· trom . tot
omewlL.Jt d.iflerent characte-r; whJCh may carry few generations. It may then spread to other mt 1 m.tny
ome 'deceive advamage. An advantageous muta- gamodemes in the vicinity, increasing the dfccuvc d,llt 0 IS 11
tion, wluch. may only have a margm:tlly bencti i-al range of the usetw mutant condition. Such J mutant may c n1e
effect, will only be a~ le to spread 1f (1) it is recurrent conilition, especially if occurring in J peripheral only a few
wichjn rhe popuhtion ·o that new recmits are isolate (i.e. a single garnodeme. or mall clu tcr of tcrcd and e
continually added eo the mutant stock, (1) it gamodcmes on the fringes of :t larger widespread pooL Thes
catrie some sdect1ve advantage so that the population), will be the first tep toward. the fonno- pecie , a t I
Hardy-Wcinberg equilibrium 1S shifted in its fav ur rion of a new and distinct species. The accumulati01 an tnitJally
and (3) the population ts small enough for it to be of a fevv more mutations m:.~y then i. olate it com· nut by 0 11
carried and spread relatively qlllckJy. plctcly, as in the das ic D::u-wiru311 mode. The origin ,ltld C"-']JJ11C
All alldes have resulted from mut<ltion; multiple of species from uch peripheral isolate~ is known as 'genenc bo
aUeles h,wc ansen at different time~. Some mutant allopatric speciation (section 2.3). When the new n hi
V<l ria t1
aUde may sponr-aneou~Jy mutarc back again to the type is sterile with the parent tock, h::tving ruverged to he m Z)
original gene e.·pre,sioll from which they aro~e. enough genetically, the 'parent' :md 'daughter' ultun.ltely 1
The rates of mut tion vary with the gene. Some are ~pecies are then ennrely distinct.
mu h lugher thJn others, but they .Jrc gcnerJlly Recessive muranol15 spread compa ratively slowly
low. one mut.1t10n tn IIJ llUO tor a 1-'anicul..u gene and only in m:~ll population~ with dme inbreeding Geneti<
being considen:d a~ high. The nte of mutation can is there any likelihood of more rapid ~preading.
be increased by rJd tarion, mcluding exposure to However, recessive mut.ttjons can become domi· If a populOJ
ultraviolet light, or by chemical mcm~. n:mt under certain circumstances. E:-.:perimeut1 limited an
have shown th:Jt changes in temperature alone ab1hty \ VJtl
nMy be enough to modify the dominance of J or the ov
Spread of mutations through geue. ln natural populations there arc some gene1 ~m.11lcr tht:
populations which exist only a~ modiftcr<; , changes in which ma~ h.IIIU d do•
:llter domjnance ro rcccssivcne~s in ~ gene. If a At. each
Mut:~tion ~~ tht: primary >Ourcc of variation. but it is recessive allele is favourable i.t will be selected for v.1th its nu
only through the dli:cts of ruplmdy in higher organ- domj nance. thL' ~amet<
isms that the rnutJtions can spread. & mentioned. it lr< nor re
""' and a
Isolation and species f o rmation genetic drift. becomes irnportanr in tiJese mall
population . his i simply an evolutionaty dJJttgt:
'hould now bt: rl ar that ome kind of isolJtion is in gene frequenoes through random t:baJKe asson-
mnall; es~cntial tor specie fom1ation. lr nuy be J mcnt, without sdecoon being involved . Henc ~: Lt'r-
. 1graphic<1l l)olation, J.e. ~ome parr of a popubtiou tam localized popularious descended fruw an iuin ..d
n ng cut off from the rest by a barrier, as norcd p~11r or few pairs may become qmtL di>tinctivc or
lxlvc. But there are orhcr kinJs of isolation which may even differentiate as new sp1. nes mere!)
m1 be equally dfcctivc in certain cases. There is because the rounders of the population h. ppeucd [0
,:olog~e.lll~ohtion, in wh~eh ~pecies bccmm: adapted carry a particular et of alieles.
11 rrucrohab1tats in the ~:tme general area. Physio- In :my gene pool the etfecD. of all thl' f1ctor con-
r ~Lal or b1 logical fa r ~ ma ' aJso isohrc popula- sidered' ·o far are conrinually inrera -nng, thougb
ulh o that they . pc iate, c .g. changes in the riming mJn · biologists beheve that most mut;ltion .ln:
t'th~ brceding cydc or merely izc dificrcn cs prc- neutral and o cur ar a more or le s constJJH rate.
drn copulation. AgJin, there ma be a simple cne fi-equcncic~ w1thm the gene pool arc relatL' d at
tn~uc incompatibility; ferti)jzari n may take place the chromosome level to m tatlon. crossmg over
ut, because of different cl romo ·omc lengths or and recornbimtion. They m::~y be 'stabthzcd ' tn a
mons of alldes, recombination is impossible and particular kind of selection that actuJlly prevents
d1e hybnru abort 111 embryo or are terile. evolution in a large population, for 1f the spcc1cs ts
l Th~re are other ways in wluch isolation might be succcs ful, wid ·preJd and well adapted to its envi-
h 1evcd. A population ·explosion· or flmh, derived ronment. the undesirables are weeded out and the
m a founder population, ma initially expand only opcr.ltion.tl selectiOn is stab ilizing selection.
rr 'l many new an:a in wluch compl'tinon or pre- Thi~ 1 amlogou to what Mtvarr proposed in hi.
1ltlll l\ minimal. After the first flush, however, critique f Darwin. n the other hand, gene frc-
may come an equally udden cr.~~h. which leaves quencie begin to alter when the cnv1ronmcnt
®lv a few small isolated populations. widely cat- changes to be omc easier or harsher or when a pop-
md md each with its m n sample of the total gene ulation migrate into :1 new area and. finding lJCtl
pool. Th~ e in tum may act a~ founder; of new competition, bq,rins to differentiate cc Job'lcally.
peae~, at it-a t orne of which may urvive. Where The intcn iry of selection under these la tcr circum-
111 numUy Jarg~: population has been all but wiped . rJ.nces is much greater. At su h t1me~ there may
• 111 by\ me cara tr phe, rh~: speci~:s may recover occur directional selecri n towards one cxtn·mc in
Jl':qJ.llld af;;}in. H wewr. it ha P<JSsed through a re. pollse to a 1 ng-c ntinucd ~election pressure. and
·~~n~nr b1ttlencck' m whteh much of tbt: potential rlus Is seen in the fo sil rcc rd <b an evolutionary
, 1tl<l11 has been Jo~t. Tlu: loss of thest• allclcs leads trend (though ofi:cn it is hard to sec th~.· rL,t on for
homozyg Slty, and S1l.:h l.1ck of variation may the trend). Altematively, d.i ruptive kction 111ay
•nnJtd) pro ~: demmental. occur; the commonest phenotype in a population
can be selected a"rainst, and the two extreme~ arc
p itively sdected, fonniug t\v ditlerL'nt species
Genetic d rift: g ene pools fi:-om an original nonnally distributed popul. tion.
One would there£ re expect to find in the fim1l
111 pupubnon i, very small it will contain only :1 record reli ~ o[ long pe1ioch of species st1bility, . ep-
llltd and random s.1mpk of the total genet1c van- Jrated by breaks representing times wlwn spcciation
' within the whole ~pccies. i.e. only a fTacuon was ve1 r<1pid, and then again table 1 c1iods. Tlm is
•b.: nvwll 'gmc pool' of that species. The very often what i tcmnd, as will be cxpt11ned .lt
ull.r the popul.mon, the fewer will be the allele ~ 1me length in the next section.
•ILkd dow 1 to future generation-.. In any evol ing sy~tem the more nucrocnvm n-
At tJdl m~•o~is only one chromo ome of a patr, ments there are. the ~rreatcr will be the possibility at
.th Ill rmdom ' mple of alldes, is passed down to differentiation. pecies thus can be :ud to arise in
tbr .:,.~mete : my allelcs 111 a riny populati n that responsc to the cnviroru11cnts actually availabll'.
nut n'l'ombined in lerrilization arl' therefore It is well known that nan1ral scl ction i · operacing
I ~ md 1 1pecial evolutwnary process, kno n a at the prl' cnt time (Endler, 19R6). Evidence for thi~
comes fi-om anou~ ~ourn:\. TopKJll·xamplcs could tinkerer'. The whole or~nism is a 111gle ~y~t'm Mol a
includt: th~: 1i c of rnyxomatom-resi.~tant ·tnim of and the morphology and function of its componcn
rabbits r antibiotk-rc~i.,tant bactena. The case of parts have to be compronuses amongst c mpt:rin~ .I" i al g
'indmtnal mdarmm· in moth~ has been cxtcrlS!Vely demands and based upon genetic cllld physi al nt;l· IIH'Ill~ ~0tl
studtcd JnJ 4U~'tcJ. Before the industrial terial which may Jtsclfbe li.minng. sihl1·, throt
Re olurion 111 the north of En~.u1d, the moths For example, the lt'JlSes of a trilobite's eye J liit' othcr I
mhabltlng the lichen-covered trunks nf trees had a made of calcite (Ch:~pter 11). as ts the re'it of the ant·
pale spcdu~:d colour h.tmwnizing WILh rhc environ- mal's cuticle. Presum:1 ly trilobite~ werl unable r~
ment mJ r~:ndL·nng them inconspicuous. When secrete mJterial other than calcite for any part c ftbr
record, were first kept m the early 1XSUs these exo keleton due to inherent genenc limitations, an
moths m de up 91)'},, of the popubt1 n : the other 1% ~though calcite has the virtue of transp~1rency, 1
were dark-coloured types '' luch we;-re more easily disadvantage f r ;m optic, I s. rem i. that i\ :1 high!
pit ked out by pn:d_1tors. But by the end of the cen- birefringent mineral. producmg double image'
tury thl· wuod!Jnds ne. r the 1ties bad become pol- different depths, for ligh t travelling through it in ~n .11tJ nmi.l '
luted and the l11:hen . wh1ch arc very sensitivr to oth~r direction~ than the miner,llogica.l e-a·. ~btp l t ge
unclean ~1ir. h,tJ beell killed. c~'Posing the blackened Trilobite have overcome this disadvantage tiN b1 H 111 liUdt
tree tnmks below_ The vast majoriry of the mod1s in orientating the r-a.· i~ normal to the visual sUt{acc tJJnk·c tlar
these area were then the d.Irk-colourcd fonm, the and second by many ing;eniou~ moJificatinn~ o the [· en s
pale varieties .ll'Countmg for no more rhan 1%. crystal btrlce; they have al\o evolved lcm doubk 1111 purta 11 t:
incc 1t had become ::IdvantJgeom to be dark- which bring light to • harp torus. In ~pHc, thcr,. nt L: •olutic
coloured 1 th t prt>1.btor~ would less rasily see th ' fore. of an intmmc limitation, ~cleni n pressun Ule ~en t
m th on the d.trk tnmk~ . natural selection had bet:n has 'for ed' the best functic nal ~ystem that \\l d. tlJI11tc s•
mtensc m the changmg crwtronmcnt_ omc 70 p ssible. t IUH (e,
specie Jf morh .u-c known to havt· been affected by , uch anatomital aml physiological adaptatiom .w llunkJp1llt
such mdmrnal melanism witlun the very short one re ult of natural selection, but ammal populi I' ·rlup
timesp.m of 'iO yeJr~ Bur only a few gene an.: tions arc also adapted to their cnv1ronment rcp!'l)· 1 r hat nu
atfect.:-d here; It re:oqmres more changes to initiate ductivcly and trophically. In this re pcet, 3l1d 111 Ih.1 1 lht"'
spcc1,1tton, involvil'lg a grc:lter variery of ~enc. con- br ad and general tenns, two kinds of adapti~c tll u~ands
tr lling different character~ . strategic have been chstinguished. They arc differ H I.H d gc1
A more recent cx:unpk conccm.'i the amc cnt ways of . urviving and reproducing on a limitt mc::uungle•
G:1bpagos finchc~ ( t'1upi;::t1) ~tudied by Daf\vin. energy budget. What are kn wn as 'r-strategJsn Ik I Jt:b
ne ~pcCics is a spc ·iJhst feeder on small seeds. have very high fe undity so that great numbers I NA nd
ollowi11g a severe drought in 1 77, the av.1ilability offipring arc produced raprdly . lndividtwls may "tJnJer~ a
of small . eed~ bec:unc much reduced. Bigger seeds short-lived. rclymg upon their superior repr due 1JNA than
howc\cr, were more .Ibunda.nt. and thuse larger tive abiltties tor species survival. 'K-strategists', 01 lllJl ·nal h~
bird~ wlth ~rronger beaks that could ct>pc with these the other hand, produce fewer offi pring, but arr Swlll' o 1t
were abk to survtvc. It ~<.>llow that the pressures of u.mally slow growing and live a much longer time gcn , l ut
namral selection vary frorn pLtce to place and from they are better at competing for food and livin1 g "ll fa mi1
year w year and tb;It there is no reason why small- sp cc. Tll.is is an attractive concept and in sonic lllUlllpJe C
scale evolution ~buuJd not b~ reversible. respects still useful. Jt has. hm ever, been heavih •>plc' of r
In the <·xamplc ]Uotcd above lhc sdect1ve advan- criticized, since ome pecies have both high fecun· 11\ ll' lOpl
tage of the gene is dear, but in many ca es it is le diry :~nd superior competitiveness, and :~IJ pm~ibl 11lenricJl u
c,Isy to w1dcrstand the sclecti e advantages of parnc- combination of these end-p ints may occu: "lC\\ I I'
ula.r characters. ( teanlS, 1977). Furthermore, the concept of t:h redundant,

election, then, 1 seen JS a primary agent m evo- 'limited energy budget' is not invariably valid . ft tbh lear.
luaon . It Joes nor, however, produce ideal geno- therctore not very ea y t ategorizc adaptive str:l ,.,, ode m ~
type~, bul only the best avallable given the gies very simply, :md ir seems rather that population ,,tutt: rim'
JIUmerous constraints wftich operate at any tLrnc. will tend to maximize whatever reproductive and ~IT~.:ns of a
Thus, as Jacob ( l Y77) h:1s well said. 'NaturJI elec- survival potenti.1l they have, in the circumstance~ i dl'lcrcnou1
ti n does not work. like an engineer. lt work like a wh1ch they find themselves. , t[lic~s; it <
Darwin , the species and natural selection 37
Molecula r genetics and evolution \hiddcd' component. ft may even. afi:er much evo-
lutionary wvcrgencc, tum our to be usefitl in ~ome
Cl 111Cal genetic~ make\ me of bn~edjng cxpcri- category, to an eventual descenrlanr nrganism.
mrnt \O that rhe gene~ .1re undef\to d, as far as pas- The ':1mplifi •d' amount of information carried in
le, through the tr effects. Molecubr gt:neti~.:~. on multige11 e t'jmilics may, in way~ not .1s et fullv
dJ~ Ofh T hand, COilCentrat ~ llp 11 the nature of the under~tood, be n ccs~:~ry for the maintenance of the
.~nt , Jnd upon how genetic m.ttenal 1 · stmcturcd complex functions Jlld inter:lction' of rhe cell. of
JIIJ orgJnized Now that' e b:l\'l' covered the basic metazoam. lt had indeed been 'ugge~ted that the
' dJ~>k,tl evolution theory. ll \ ould -.eem appr - ambrian 'explosion' of metazoan life may be
rn U' to Cl m1der the impacr of ne\ di~coverie~ m related to the inioal emergence of uch mulrigene
•Jtorular genetic . When::-t~ these are rcvolunon- £uni1Jes'. When more is kno\ n of their funct1ons. 1t
\, th~ rhemical p:1thw.1ys thJt lead from genes to may perhaps allow .J furth er de clopnwnt l ) f thl\
-~1' Jcvelupcd org:mt ms arc 'till largely unknown, concept. For the mornent, mnltigcne t:tmilics '-l.'em
.IJ,d until we have a dorcr p~eturc of the relauon- to be primary organizattonal units withm the
,lnr of gem:~ w tkvclopment we .,h.11l \till be far genome, aud an inrnnsic parr of a Jyn;lmic an I
11 111 umlc[);tancling how evolution work~ at the in terKtive y tern of genetic control and regulatiou.
UlO!elul,tr (,111d hen Le the mo~t b.tMc) level. Thc~e new concepts have led to two pmsiblc
E1cn m, what hJ~ Jlrcady been e~tabli~hed is hallcnge ro neo-DarwillJsm. he titst is Kinlllr,1's
11 m1am fur undeNtalldJPg hitht·rw clu tve aspects (l97Y) 'neutral rheory evo lution'. Thi~ propmt·d
nulution the ut • ( lr 11 .,how' rh,tt rhe genome that populat1011S arc much more diverse than vould
• l' ~~nt!tic. appaJ-atu ) is ~ mu h mnre fluid and be expected from natural sele ·tion. The levd of
s ~ n.m1c yslt'JU than fonncrly ~upp 1scd. with dJf- variation h uld be much lower becJuse 'useful' dit:.
'lt levd~ of hicrarchial cmnplt•x1ty (e.g. tcrence shou ld spread through the pupubtion. and
!1111· pllkrrt al., i9H2) . 'hannful' v 1i.ants ~hould be climi11ated. Smcc then:
P(rltap the most unportaJH oC rht.'\C t b~crvati m i~ actually a high degree of cvtdenr vari.1bility, most
t tlt mall) ntkaryotc' cmy much more D A of it is nodilng to do with natural sdectiotL lt ,1rise~
tlun they need tor coding protcm~. Tlm eXl'\tS a~ from chance differences at the molecul,Jr level. <lnd
•h, u~;u1d of cop1c' of tltc ~me gene or Sl'tS of mo t of the va riation neither advances nor retard~ :m
rd.1tcd !(ene~. Such multiple n.:pcotion~o ' ccm to be injividual's survival. lr ts purely neutral, ami
Ill anmgles , especially sinLe there seem' to be no whether gen s for particular chJractet s persi tor d.i~
•r rdJtions!up between the Mn )lint of l'Xces'i appear is by chance alone. Trus is a debJtable point.
f D ~ und the ize or con1.plc. icy f the anim,tl; sJh- but conversely it could be argued that even if very
mmdm ilnd primitive h hes carry much more many diffe rences are ne utral, those few tlldt do con-
D~r\ than mo t mammals. uch uncs entia I genetic fer a s~:lective adva.~nage m~ available to be moulded
t ri.tl has been called 'redundJnt' or 'junk' DNA. by the redoubtable po'vver of n.1tural sekcb.on.
ne of it consJSts of many n:pctitiom of the .1me Anoth r challenge. deriving from the concept of
ne. hut mu t of the exccs' DNA cx.i'it:. ao; multi- mulci gcne fantili es is Dover's ( L<:lH3) theory of mole-
J 1 famtlies, i.e. groups of repeated !>ClJUCnces or cular drive. re is ba ed 011 the understanding tlut
mlnple copies o the s,1me gene. _ uch multiple individual memben of :1 multigcnc f1mily show great
• \\' of non-coding sequence exi t coo. and mul- similarity w ithin a species. The individual members
lrk copies of all types of gene ~equc.:ncc~ can be of a species do not evolve independently but <Ire
e Jd~IJ[l< .ll or qmte wid ly \'"Jriable. Wh,u Joe~ thi largely homogenized in a phenomenon known as
r ·r1 DNA, wluch at tirst sjght appears w be 'concerted evolution'. According to Dover tlus LS
e r!'<lund.mt, actually do? Some tim..:tiom .ll'l' rea$\ n- due to fixation of mutatiOns withm a populanon
l) •br. A mulngene fumil • for ex:lmple. can bc~.:auSL' of molecular mechamsms w1thin the
,n vdl tllJny closely related protem~ ,tt one and the genomL'. Thi he calls molecular drive . A number
mt nme, mher than just one. Moreover, the of uch molecular mcchanisn15 (e.g. gJm or loss of
ctln of a mutaur genl, ~vhich othenvt~c might be ecti ns of genome by tr:ln positwn) can operate to
ld ·tcriou1, will be butTered by the other gene allow genes to multiply o ut of ph:m~ with the organ-
t•ptn; lt doe\ no harm but ~lays 111 the ~y\lt:ITl as a i~m that bears them. The genes controlling ditfcrent
38 Evolution ond the Fossil record
characters tl11 thus incrca 'e m frequency due to just perate in sequence. First. maternal gene estab- ~pen.1l ins
'uch horiwnt.tl mn!ciphcanon, and not to any effects lish the antero-po tenor • nd dorso-ventral <Lxc~ of procc se . I
of natur,tl s~.:kction M lecubr drive can thus tl: a the egg, before tertilization. ccnnd, segmentation can be estal
variant in .1 mult1gcne 6mtl , and such tlxed variants, gene come into operation after the egg i fertilized. uncle t
constantly tttn1Ld over in a breeding populauon, give and they define the number .md position of seg- ogy, and at
uch homogeneity. uch ,1 c ncerred patrern of &..xa- ments in the b dy. The tlurd group conmt ot mumty eve
tion dcfin"' hiolog~ ·.11 di. conrmuine~ from one homeotic genes; the'e control the uniyuc.: identity Of equ<
specie' to thL m: ·t and may in Iced be one mean f of a segment, i.e . how it 1~ constructed Jnd 1; •11Jt lt from the~
rigin t bioi t>ital n vdty. Molecular drive in n- I oks like. These seem to work on :md Integrate the have :triet
~cqucncc may hL .mother wa in which ~pecie can ignal pr duced by the segmentation genes. A ha~ been '
ari~e, .Kridt·nt:tll), ,md by non-Mendeli:m causes. 'mur:~tion in ::1 home tic gene complex nuy produn hccn very
Thus whik n..tt~~ral selection is rradmon.lly under- a duplicate segment. re ·ultmg, f, r example. in J a, even over
~tooJ s op~..ratin~ ,rt the lcvd of the org< tnism .:md ts with four perfectly formed wings inste. d of two. ra~e that
fum.lamcntally tdaptive, mol cular drive 1 n n- hese three groups of genes act n eJch other Ill doe~ not F
adapti\ L and op~.:ut s at the level of the genom . It sequence and prcsmnably code for proteins that u 'tart'i. 'om
may aJ o bL contrasted with gl·netlc drift, wh1ch t\Jm h:we th eir effect on other genes. incluthn~t lution in t
operates at th level of the organi~m but i'i non-. dap- those c ntrolling stmrntre. ome genes (maternal denly dive
tive. Molecui.H dn ·c m.1 well prove tn be the thjrd segmental and c~penally horneotic) contain .1 short. adJptl·d fo
eau e of \pl.'ciatit n; but it JS hard t ee hm 'ii.gnifi- highly conservative DNA ~equence (or mottf radiation
cant aclapt.1tion might .tri't: rhert·fr m . known :ts th e homeobox. lt IS 1~0 b.1 e pairs long evolurion
Neither of tho.: 'm•urral' or tlw 'molecubr drive' and constant in size and sh::tpe. The homeobo wa1 ~tr. tigraphi
theories scnou,J undenn.incs nco-Darwinism. first discovered in homcotic genes 111 lJrosoplri/<1 Ill sound. tho
They do, howcv~.:•·. aise the question of how much 1984, but has now been ~hown to be homologJUI t the LO U :
of the variat1 n we see is there because of chanet.: in various invertebrate group~, including primmve n::tture of
alone and ho\ much has .tccumulatcd bl'cause it metazoans, but also in verrcbnte~. 111 ludin dieted, tlH
actual! LOnlers 1 sdccttve advantage. hum::uK Homeobox-containing genes must be Vtl) from ob\er
Furthe• dJsctwcries ,lt the molecular level include ancient, ~t lc:JSt 500 Ma old. ince they . re round St These v
transpmons, and cull et tivdy this new mfonnatio11 djver~ely, and they always fi.t nction in segmenttc detail in th
must have a bcJring upon rap1d but eo rdinated animals i.n conrrolling the spatial expression of th( the outsel
e\·olutionary ch::m~e, Jntl the origm of new rgani- body a.xi~. They are, h owr.: cr. present in non-seg- genetici ts
zational pattem.s. mented phyla .md evidently they do not alWJ\ he chang
operate in the same way. lt seem~ thJt these highh elective b
con ervativc homeobox genes c::tn coopt othrt take m:my
Gene regulation d uring development fun ctions, and so, in the evolution of development~ represente(
proet:sse 'the sJJnc rcgubtory clement<> are tl'C001· bedding pi
When .1dulr )rt,ram m develops fr m a ferri lizcd
111 bined into new developmental machines' (R.1fl ceed to ex
~:gg, it Jo~.:s
su .Kcmdmg t a prederennined path- 1996). Clearly the bomeobox concept h:t! an evi- evolution:
way, controlled by ~pedtic genes. These genes arc dent b a ring n one of the major areas of cvolutim· Mic roe\
switcht·d on and ff in regular sequence, and :my ary palaeontulogy; how new body pbns originate within Sf t
err r in thC. pr et could be lethal. The product of and how they can be mudified. In rhis context !Tom ones
on~. gl'nC 1' likdy to have a equential effect on palaeontology supp lies the time dimensiOn. whtrh the other
oth~:r gnti.'S, Jnd in long-rem1 developmem we ca n can be integrated with genetic and mokcubr d::tt.l. species lev
cnvisagt· a 'case 1dc. r g ·ne tic controls . Thm a gene wttb adapl
switcht•d on, or turned off, ::.tan one developmen- modes will
tal stagl.' controls the ~:'\pres. 1on f other genes at a
sub tqt.Jent ~tag~·. The cnJ result is a fully developed
organism in whio..h "there i. a place tor everything, Mic roe
and cvr:rytlun~:; 111 ir place' (Gee, 1~J%).
Tht>rt.' an: thn:c gn>up~ or regularory genes that We ha e already seen that palac01 tology, \ ith 10 The origi1
have di{fa llt Cunrtiom dunng embryogenesis ~md unique perspective of geological time, can ~m t1rgely thn
Fossil record and modes of evolution 39
•P oal insight!. into the nature of ~.:v lutionary an.i.Jn:1l and its environment. are generaUy .tttributed
nro nes. [n partJcul.tr, tk~ccnt of ev lvmg lineages to microcvolution. Although the generic and tsolat-
n be established. nd the hi~tory oflife through tJme i.ng processes by whi h ~pecies differentiate are ' ell
udcrstood. Thts i~ an important t:t k fi.1r pJJacontol- Lmdt·rstood, it i · only comp:.~ratively recently thJt
,- ' ~·.and at present both phyletK evolution and com- full confinna riou of hO\v they hJ e actually oper-
tf ntUllty l'volution are recc.:wing much Jttenti n. tcd through time ha~ been clearly seen. Simp on
V 0 equal unportance i-. the perception. glc:med (1 944, 195.3). working mamly with vertebrate~.
lt 101 the to~sil record alone . that rale of evolution linked gL'nctic the ry and palaeonto logical dat,\ in a
\J.Jw varied th rough time. In ~t me tmtances ·hange tund , manner. as d1d C rter (195-1-) . lt came to be
1u.s b.:cn very raptd, but rn other cases there h:IS undersrond at thi time that pccies could anse
~'ll vt:ry little evolution.1ry moditicatmn . t Jll , e-ither by lineage splitting (spcciation) or by tr.1nsfor-
"n uver long periods of rime. lt ~eem~ to be the m:Jtion of the "vhole poplllation (an, g(ne 1s) .
' 'e that evolutionary change m m.my m~tan es Unfortunately. however, there were very few
J1 ,\ not proceed at a regul.tr pace but tn fit~ and imt::mces where o ·j] popubtiom were actually seen
11 t ~. ometimc~ we can Jisccm r:lpid bu!"'its of cvo- to show a gradual modific:1tion through time:.
g 1 non 111 tbc ti.1s tl record, a ,ingle ~tock ma . sud- Indeed, many of the da.s ic ca_~e which were sup-
.nly diversify and irs dcscenJJnts qutckly becn111e posed t exhibit uch unbroken series have not
lJpt,·d for ltfe in v.mou habrtJts. ~uch adaptive tood up well to modem an. lysis. imp on md
r.diations :m.• one of the 111mt evrdent pattcms of others ,trgued, as had Darwin. that the rarity of fossil
olucion revealed by p.1beontnlogy. Although the specinwn mtennediate in morphology between
mgraphic . equcn t: i_~ IlK rnplctc rt still give~ a succc~sivc forms :ts due to the rncompletencss of
und, rhough in some HlStJIICl'~ tragrnentary, guide rht: rock record. here i~, however, : m alternatlve
the course of volution:1ry ·h;mgc. Although the view. enunciated by Eldredgc and , uld in 1972,
rure of the p;~ttcms seen nught have been pn:- that the reco rd can be taken at bee v.1luc, and the
ed, they would bt: nnlikdy tn be conhrmed very bet that there are so few rransitional ~cries
mob ervaoom on modern anrmal~ , lone . known ha~ something fundamcmal ro cell us about
Thc1e variou~ is~ues will be explored m greatc1· the nature of microevol utiooary proce ses.
um the following text. lt ~~ irnporrJnr to noLc at Accorcliug to Eldredge and G uld, !llosr palacon-
LJut\et, however. that pal.Jt'O!Holog]st. .md tol gtcaJ thinking h:1d been dominated by the Jdea
'ti\:1St5 are workmg .1t quite diflt.~rent timescales. that evo lu tionary change must inevitably have been
(hange~ that .1 gen ncist c:tn produ e rh.r ugh slow anJ tcady. This m del of gradu:ll de cent
1
,·ri\·e breeding owr 1UD-300 gencr:ltlons may wirh modificarion c med to be in rune with the
c many year~. Y er th1s s.nne timcsp.m m.ty be majestic:11ly unfolding. long-term operation of the
rmntcd gcologicallv by no more th :m ,1 <,inglc processes of naturaJ election as originally envisaged
tng plane' Bcanng d11~ in mind. we can pro- by Darwin. Was it not, :Jfter all. cla~sic Darwnusm?
J to examiue the two m:un g;raclc m· modes of The idea f pbyletic gradoalism implies an even,
luaon: microcvolution and macroevolution. sl w. rnerhod..i al change of the whole gene pool
~\tcroevolutton refers m small-scllc changes and hence of the charJcters fthe entire populatton.
hm species. and cspecjaJiy the tramfonn.ltion This transtormatiort wou ld invol e lar~;c numbc:rs,
mr one speoes t a m:w one. Manocvoluri n, on mual\y the entire ancesrnl population, and ould
ther hand, concems evoluoon :tbove the occur over ll as J large part of tht: ancestr.tl specie
o level, With the origms of major groups and geographical range. lr ccrns a rca~onabte con epc,
~ Jdaprive radiJtiom. These two cvoluti nary and certainly omc popularions may h, '"' ch,mged
~ will be c o 11~i krcd in turn. in jmt this way. Yet, as previously norcd. continu-
ou series of graded intcnned.iate fonm 3rc eldom
· c~..·n in stratigr:.tphic:U cqucn 'e - h.lt we normally
Microevolutio n ~ce i~ a sequence of stable fossi l populatiom cpa-
r.Jted by abrupt morphological break . To some
I ~ origin~ of new pene" from extstmg ones, dq~Tcc tlllS may indeed be due to incompleteness
I· through changr ng mt -rJcttOLlS bt:twccn the <111d bias in the fossil record, but ru Eldredge and
Gould 1t ~uggestcJ thJt evoluttonary ch.mgc occur-. c:i.tlly if the popul.ltton i' adapting ln a m·w lo<ll v. tthout tral
only in ,hort-livcd buM~ (punctuarwn s) in which a l:llVlrQlllJJent. fo begm with, JJl 1\0J.Hcd populJIIO next •dapt1v
new spt'CIC' :a-i~t'' ,tbruptly from ,1 parenr ~recies. \UC h as thl\ m.1y de\·dop .ts a 'r.t<. e · wtth d1lferct tOt:ll gene p
often w1th rdativdy large murphologic al rh.mges. ecological requirements , but (it put tu the test) 't' m·wl-. tsola
and thereafter n:m.llll\ more' or le~~ stable until it~ <. .lp.tble of Interbreeding with rhc {)( i~rinal populJ· hJ,J\t•ci \JI11f
extinctiOn The,c authors h.1d mdepcndent l) docu- uo n. Wirhm a rdanvdy few gcner.nmm. hownc• w holl gene
mented preci~e patterns of microcvnlun onary .md wuh contmued 1\0l.lnon. th1~ popubnon m;fl' dm m tt,el(
change in Devolll.ln tnlobltl.'' from t:J\teru North become reproducnvd y ..1' well 1s geogr.lphtc:!ll} 1 • k~, m.ty be .
Amcric.t (~ection 11 3) and m Pki,tocen~o· lanJ snail~ bred !Tom thl' parent pupularion. 'I he min:~l ntl'Jn· tht· lllhcr h
!Tom Bermuda (set nun 8.-l). All their ev1dc•nce indi- ber. of the pt•nphcral tsulate may well be fi:" 11 'g• lll'tlc bot
cated rht Jbrupt on~o-'1m of ne'\\ 'pec1c~. followed by number.. .md .trt ,t,ltl\tll ally likely to CJIT) :1 'et h IVC Jtttlc cJ
~rasi,, ,md to tllis n~t,dd of cvolunonary ch:mge they genes somnvhJt dtlferent to the parent popul.umn
gave the name ·pun ctuated equilibria' . In thest: ~nu ll populattons evoluuonJry ch.tngt c Heterochr
Thq· felt that th1' model 'cen1ed m .1t.cord with ta'it. Thest· lllJY be the founders uf J new :~nd flou<· Wh1.11 Jn o
what I' wen in the fo~'il rt•tord ..md 111 Jddition 1\hmg spcn<.''· "hieh ,lftc:r the op<. rJtiOn of fimh the dilTeren•
would pn:d1ct that mtcm1cdt.lte ' \\ ould nor be 'clecnon prc'>'>Ure wtU be ,ufficientl} moditied ttl t>:: SC\jlltllCC, aJ
found. lt wJ~. moreover, nor nul of keepmg w1th rt·productivc ly distinct from the ongm~1l specit·'· c 11ly t·omrol
Darmm.1n thought. for D.1rwm himself had envis- The ~tepwi\c p.ltt<.·m of 'pectcs chJngc normaU 111 ~~J1u~n1 ar'
aged thJt not all t:volution;~ry rh:mgc need be grad- 'ten m tlw fo,.,JJ rcrord may nO\\ rt•adJly bl' llltlr rate of devc
ua1. Indeed 111 rh~· lourth editmn of the ()ri~in o( pretcd. Thc· .. harp murpholog~tal dic;continuth lrom 1. .tch 1
Species there is the tnllo"' mg conuuc·nt: 'M.my .1ctuaUy 'ieen 1n \O many 'equenccs ltldiCalc tlut tb chrony. de
~pecil'\ when ann tnnm.·d tH.'ver undc:r~o fi.1rther de~cendant \pecies, wh1ch origmarcd .1~ a periphcnl npp1. 1rance '
changt: hut becomt exunct WJthmn lea\ mg modi- i'oiJte. h.1' cnonnomh. expanded .111d tu .. movt .cter\ (GOI
fied de,t·cndanb .•md the pcnod' during \\ luch the b.1ck tnto thc area formcrly occup1ed b, the p.~rc Ml Kmney .
spene~ h.1vt: under~;tonc modtht.ltlon , thou~h long 'pecies. lly tlus time the parcm 'Pl'CICS may ha1 l•\\" '' ;;:c Jlo
as measured in ye.1r., have prnh.1bly been short in become exttnct. or uuy have broken down 111t1 ~ ~olllple, of
comparN.lll \.Vith the penoJ, during \dllc h they 'miller populations (and perhap~ eventually to wp. of •
ll)'llll'lll
retain the ~ame fonn' . rate species). or m.1y have moved .1wav followmg 'I:Hlu.1l of no
If tin. punctuated cqu1libnutll modd 1~ .1 reason- 'htfting hab1tat, or (though this" k" hkely) tht• n lonn. howe
ably t~1ithful reAcccwn of wh.ll actually happened. .\pccies may h.we competltivel y d1o,plJc.:ed it. 111~ rile of d
we too have ro rcnmder explJtl.ltions of ho\\ r.1p1d From the bJOiogic:ll pomt nf vtcw. howcv bttt' nny go
change' occurred. l ,,.o of thc\e will be pursued .tllopatric .,pcciation " only one '"·" m wlu thl' .lllCl'\tral
here. reproductive 1solauon can occur. and m reCCIIl } tlt'm v. h ich
th<.·re ha~ perhaps been less c•mpha~is on th been rdati•
Allopotric speciotion .1llopatric model. Geogr:~phjc i\ol.ttion is not Jl gn'lwth tem
Mayr ( 1tJoJ) and othc~ propo~ed that new ~pccic' cs\enrial factor for all modes of 'Pt't:iJtion tho1. th ancestra
are onl} hkcly ro .m,e hy <;omc kmd of reproducove Lkarl) it t' an unporrant tactor from the- ea
isolaung mcchamsm~. for the process of ~penation The ongm of r;pcnes nuy bl· 'it'<.'n a~ a <..1\t' 1 gra!lliiiC.
must treJte some kind ofb.mier to gt:ne flow. On(• quantum evolution :tt the spcctt\ lcvel. The L\l
such kind of 1~ulanng mechallism mvolvc~ lated population has to P·•'~ out of the ~et of ctol"r- Ch 111gt·~ in
tcally prcu~e paramt'ter. to which 1t and the pare {Tilt tu re~ d
allopatric, or gcogrJphic -;pen,HJon. Let m 'uppost:
that a rebovcl} homogeneou s popubt10n eXJsts popuJarion are adapted tnto another ad.1pt1ve wt two posstbil
within 1 parricul.1r .tre.l. Gent' flow throughout the The intervcnmg conditlOlls Jre umtablc ami inadAI" .amorphosi
popul.ltmn is achJe\'t•d b} frequent um:rbrecdin g tivc: mml and indeed ur;ually all populatiom und • har \{ re-rs o •
and sr:~bll1ty " thm maintained At mmc pouu a gomg sud1 a tranSitiOn wtll not \UrYJve to g<llll t •f the desc
small p.lrt of th" population mt~ratcs aw;~y ti-n m th<.· next adapnv<. peak. Ir ~~ convemt•nt to com1dct t aiTc:l tl'l'i. so
parent population .md becomt·~ geographtc.ll ly im- problem in terms oLm 'adaptive 1Jnd~cape' (Wngh Jtlvr.nde of
lared. Such a popul Hion camcs tt~ own gene pool 1932) of high bills of .ldaprive filnc~s ~ep.tr.tttd 1'1 • ~.:rt~111 char.
and, 1f thl: popubnon i' \'Cl) <;111,111. rutur:~l \election km•-lying 111adaprive \\\.Unp,·. wiHch have to h1n tJ1led 1
or gcnenc dritt will mod1t)' it quite rapidly. espc- tr.werscd .utd in wh1ch mmt .~th:enturer., will '' eh~' C'lopmcn
uhout tr.ICl'. Thme th:u do ~Ur\'ivt• .md gain the 'frozen' in a more JUVenile form, yet they are \till
l}(xt adJpthc peak w1ll carry only ,1 fr.KtiOn of the functional.
IO!ll gene pool of th~ p.trent popu!Jtion . Thu~ this McNamara (J 990a,b, 1995) has sununanzcd three
~11h t,oiJred popui.H1on already ~tart' with a separate paedomorph.ic processes. Progcnem ou:urs
b.t,ed \dlllpk- of genes by compari,on With the by precono u ~ ~c:x'"lt al maturation (and thu~ .1tfl'L ts all
11hole ~cnc puol (Mayr'~ founde r princ iple), :md structures leading to a small and juvenile veNIOn of
th11 111 itself, 5Ub~cquently reinforced by mut,mt ,tllc- th e ance~tor). Neoteny involves a reduct:d r.ltt.: of
lcs.lna} be a 'tgnificant step towards 'pcci.1tmn. On morphological development (and may affect all or
t ~ other h.1nJ, if 1t Ius lost ton many gl·ue~ by only 'ome morpholog~c:~l structures; th e de~ccndant
~eneuc bonlencl kmg' referred to e.lrlllT, it 1113} •~ U\ually the same stze as the ancestor) Po~t-d•~
\'e htde ~:h.llKt' of renewal placement occurs by delayed onset of gro\\ th of p.tr-
ticular morphologJral structures relative to othel'\
Heteroch rony (only smgle \tructu re~ are involved and the\· u\uall}
~ hen .111 orgJmsm develop~ from J lcrnlized egg resemble ~mall and JUVemle versions of the ancestral
the dtiT~rcnt nr~m appear and grow 111 a pn·ci~e fonn).
Jlitnce. md the tllning of these cverm I\ gcneti- Pcramorphosis ~~ the converse process: the dc~cen
lly controlled. The evc:nn1al s1ze .md shape of the dant adult i~ morphologically fi.mher advanced than
Jl · tm~m are nonn.11ly lmked to the ttmtng of the the .mrestor. T hi' process can likewise .11Tect whole
t• nt dcvclt,pment, but they l";IIJ be Ul' l oupled organism~ or \ingle characters only. Ag:un there are
&unt e tch orher, .1nd tlus IS the bJSI\ of he t ero- three kinds of peramorphosis: hypennorpho~l\
chrony, delined a' 'dungN through timl 111 thl (delayed ~c'\u,ll man1rity}, acceleration (increa,ed rate
reJr,lnce or rate of development of anCl''tra} C: har- of morpholog~c.tl development} and prc-dlsplace-
:tm C:ould, llJT; McNamarJ, Jl)lJtl,l,b, 191.J5; ment (earher omet of growth). The morphology of a
M1Kmnq .md MLNamJra, lYlJ I) In the \1111plc~t descendant fonn may therefore be a mosa1c of nor-
stu alonl can be affected. We m a~ thmk, tC..1r mal, paedomorpl11c and peramorpluc charaCtl'r\.
:ample, of a mlob1te m wh1ch coordmatcd devd- Where,\~ hctcrochrony is more fully tre.ucd by
pmcnt nf 112c, ~lupc .md rimmg leJt:.h to .ut indt- McNamara ( 1990a,b, 1995) and McKinncy .md
11du.tl nl nonml \ize tor the ~pecie~. In dl'Sl endant McNamJra ( 199 1), two important point~ c:m be
nn. hnwcvt r, there may occur J mutatton .lfTcct- summarizc:d here. First, heterochron.ic change' arc by
11~ rJtt' of dt'\ ck1pmcnt. If ~o. the dcsct:.·mbnt trilo- nature 'mst.mt:a.n~.:ou,·, <lnd many of the mJcrocvolu-
te n1.1y ~o on grm.vmg untiltt is mmh IJrger than tionary 'stcpw1sc' changes seen in the fm'>il record
.mct:.~rul fi>m1. Then the genenc timing mecha- may have ongmatcd m this way. S~.:cond. hete-
n '' lurh Pll£' an end to growth (.md '' hich has rochrony ofi~I'o .1mmal~ .111 t.~cape from \pcciah7.ltlon .
en rtlmvdy slowed do\vn) catche, up and An ammal ~pec1e' nu~ be well acbpted to J pamcular
1\th tcnnin.lte~. The remit 1s J g~ant vcf'\ion of environment. but 1f rhc environment Itself bq~m\ to
4ncNr.JI tnlobite; equally a dw.trf l'tmld result alter, the only W<lY out for the specie\ 1\ to change
rom thr t•arh omcr of the growth-stoppmg pro- mto something eh~:, and heterochrony may cn.tbk 1t
~ lllllll.'. raptd.ly to do ~o.
Whtlc heterot h rony is probably a prilllary control
ngt:' 111 ~hJpe or morphology result tn d!llerent of microevolutionary change, it i\ pcrhap~ L"ven
"" rurc' dc,·eloping Jt different rate,. l hen: are more important in understanding tht• ongm of
0 r 1\\Jb!lme, here, paedomorpbosis .md per- macroevolutionary novelties; it is indeed one of the
amorphosis In paedomorphom tht:. 'Juvemle' most t.ntical conct•pr' in the whole of evolunonaf)
1 tc" ol tht• ance~tor are retamed uno the .1dult palJeonrolo!-'Y.
• tht dc,ccnd.mt. The whole or~lll\111 m.1y be
rClul, 'u th.11 rhc de~cendant adult re~cmbles a Testing microevolutionary patterns
wnil~ of tht· .mccmal fom1 . Altenl.ltl\'ely. only The fossil record i~ undeniably incomplete, Jnd 1t
rrt~m 1 h.tr.ttteN ch:mge -the ~tmctUfl'~ concerned has been argued that there are so many gap~ 111 1t tlut
hl11' [nkd to pa\~ through all the nom1,1l ~tages of 1t cannot really be m ed to interpret ratt•s of evolu-
,,(\tluptlll'llt thc1r morphology h.l\ become tionary change. T here is truth in th1~ ,tnd we t".ln
J2 Evolution ond the fossi l re<:ord
tor-descendant rcbtiomhips be pmtul.1ted wtrh t·cr- r.~11,1l
v.tnal
only be f.urly ~ure about pt~ll"t:~~e~ in m,mm· inver-
tatnty. In such ca~c~ dadistil an.tlysis is Jjkd} w llHIIt•than!
tebrates if we ,,unple on J !"me enough ~trJllgrapbic
'cale. Ide11ly, lo record r.ltt:\ and proll'''cs of pnwe uscfitl. Ml>lc llut 'o
gcuclil, \Ot
change. we need w unden.1J..l· very dctailetltmcro-
'trattgraphll lOilel.lion throu~h an unbroken ,edi- Analysis of case histories purel) den
mcntary ~c4ucnn:, using l.1rgc volumt:s o( m.ttt:rial l'hcre are now .1 f.tir number of well-documentt' ,l L.h.lllglllg L'l
C;lse~ tllu~tr:~tin g various pattern' of microevnllltiCln- I hcrt• an

tor ~lati~til.tl proces,mg, with ptl'ct'ic, independent
ary change (summ,trized in Cope .111d Skelton, J 1)K=i t 1 un·htn
scrau~phtcal control .llld m lr .1 broad geo~:.'T.tphi
lJI area Uarl-,on, 198H: ~hddun. 19lJO.t,b). There arc ~m111 SlllilJI-q .lft·
cx:~mples of long-tcml sta'h and of the Otlgll1 ol tllu~ \VCtc i

A1l the'c ob.Jel"rl\'eS ltl 'ddom obt.Jin.1ble. and
even 1f the) ,m:. 1t has to be rcmembcrt:d tlul sedi- new species by .1hn1pt morphok1gJcall eaps Fqualh I hC\l' l.Hl b
thcrl.' are sevc r:~l instances o t an .tpparently ~r:~du~ 1 'lW- whet
mentation in mmt deposnion.JI ~cttin~" i~ either too
\low or wo mtcrnuttenr tn P'-'nnit re\oluuon of phylcttc change. Both modes of l'volutJOn .1re p0\\1· '' J.s higher.
blc. and there " no reJson why they should I C\ ohmon.li"'
~hon-tenn, lOntmuous btOlll!,.'lc,ll pron'''l'' m the
fossil recl1rd Fvt.'n the most c,lrcfull) \Jmpled mutually cxclmtve. Illl'ntcd ot t
llUCro\trattgr.lph K \Cquence\ rl'prcsent \p,lm of time Of the ch~sit .md often nted c.1rher studJc~ whtlh (I'IX7. 198!
purported to show ~radual ch.1nge, many do 11111 ll'ntr.tl WaJ
many time' longn than hunw1 litetin1cs (Schindel
1980, 1982). The det.lil, of lm:al short-tem1 stand up to dct;likd analy~i~. They were nor earned .f( "'Ifs ne cur
out on .1 ,uftlcicntl) detailed st.tk and thetr .nnhot scn•,·d ill a
proccsse5 (t.:.g. l·olomzanon 01 Auctuatin~ popula-
were not umnAucnccd by prcconcepnom Th~ hale~. fhc
llon dyn.m11c') c.mnot nnnnally be re,l>lvcd by
p.1laeomolo~c .11 tcchnttJUC\. found what they expL·cted to tind; gradual mnrphCl •ud wnhin i
log1cal ~hi tts throu~;h time. Thu, .111 :maly't~ of th nl \\ hkh sh
This i., one problem; Jnothl·r rntlrl' M.'num one is
that rate~ o( ~c..·dirnentarinn m.1y ch.1ngc, often quite jura~sK oyster CrypfltU'fl by Trth.:'lll.lll in rht· 192H. 11h., m the
markedly durn1~ the depmH1on of a tch~ibterou~ ~ecmcd to \how .1 b'TaduaJ trend tow:~rd~ me ru,t'll gr.tdu,,J cvol
'equence - \\ htch will imposl· a b1a-s on intl·rpretJ- collmg. Hallam \ ( 1982) more finely resolved <.!Ud· gerwr:.1 .md 1
cwn. ~uppo~t thJt :t popul.nwn of brach10poth hv- te~. however, ~hO\\.t:d a pattern ot stepw1se th.ltl~ 1cr reveT\als,
mg in the.. ~.l!lle cnvlronrnl'IH h.1d slowly l h.mged in wtth paedomorphoM s as a control of spl'CJattOn. The ti om tllnc re
response to ,1 rontmuous WeJJ.. ~electiOn prc..·s~ure. If only indication of gradual evolution here i' j C\\,1rily ml·a
phyk·nc size ch.1n~;e through tunc. \llld} sp.111n
the cncJo.,n1g wdiments h.1d Jtcumubtcd .1t a con-
<.tJnt ntc rlwn ~ ~.:radual pJttern of cvoluuon;uy Of c..oursc, the researches of rldredgc and Coul hcldon\ (c
change '\\ ould be recorded 11: on the other hand. .1re not inu11um.· to t nncism e1tht.·r. Eldrcdg'-\ '<tlill Uppcr D ev<
~edm1cnt.mon h.td been llltt•ruutrent and thc:re had uf f>/1t1wps (sectton 11.3) covered .1 wide gco~;r.lpht (l·cl\t .ltld <...
been ero\ion.ll breaks, \VC ~hould ~ee :111 :~pp.trcntly cal area. !t demonstrated that ,r,1~1~ wa~ thl· nn1111 llllll W,l\ liS\
::111d that speciation w3s rare and probably raptd. Uu ll~tigr.tphy.

~tcpwisc pJttl'rn of cvulutitnl. An arhtinal .md pos-
\lbly undctl.'ct,lble bta~ ;., thudorc unposed, m.1king thl.' stratibrraplw w.ts not tightly comtnincd enou0, lound :t'> rwc
gr.l<iaoonJI l'\'ent~ louk ,1, 1f tlu.· y .1re punrtllJtJOnal. ro aUo" the prtKe\\ of spcct.mon to be pinpointt h •th dw cy
Our knowlt:dge of o,mall-~cJic t:\ olunon.H") l'\'ent<., accur,ltely. It could also be argued that the dwJ~ n•lll t'imco01
tl'r\ used, i.e. the number of tilt·' of lcmc\ 111 th Jdopunu of
therefore, is only ,\, good ,1, the sed.iment.1ry record
allow~ it to be..·. eyes of these trilobites, is itself di,conrinuou' .111 'h.tngl'~ are
would inevitably show stcpw1St: ch.mgc~. Likt·\\w ( l~t! \[r,lt1b'TJ
Yet a furcht·r problem ts how to rccogniZl' .mces-
tor-dc.;n nJ.mt rdaoomlup,. If we h,l\'C :1 t ononu- ,llthl)Ugh Gould's ~tudies of evolunon (sccnon X. odonh whicl
ous scJmlenl.lt) ~cquull.: c with t-.>T.1d,ltlon.tl cmert:d only half ,1 million ye.m of gcologtcal tunc Perh.1p\ tl
transfom1attons, theu th~ nkkr fos,tls would \Ct:m to Mlatl shells are rathl·r simple rhmbrs with few 'h.lrJ•· aud punrtu,
b<.· the ancestor.. <md the yclllll~er ones the dt"Sccn- tt·~ that can be mt'asured. hlJtCy\ (19~
Williamson (I I.JH I) made .1 vay der:Ulcd study~ bHl'~ ill <;pit
d.lllts. lf, on thl· other hand ..1 ~t>qucncc displ.1ys sed-
lllll'ntal) dun~e' coupled wtth sudden shifu. in Lltc Tertiary mollusc<. in a bh bNn in East Afn1 lt dol)l[t:' Tl
morph1.1l0b'), the ldanon.;hl~" .ue tJr le~., de,1r; the In all his 13 lme.l~t's the p.Htl'm which cmetg thev 'hm.\ a c
younger dellll'llt~ may hJw becn mdl•pcndcntly 3lcorded with the pattem of punctuated cqlllhbn abt upr ongi1
ami with lo ng pcnod~ of stam. AppJrent <.pec1.1t10. lftt• ~.IIIH' be
dcnVl'd ;mJ have migratt·d in from elsewhere. The
b'Teater the morpbolog1t.ll ~h1ft. the Jc,., c:1n .mcc~- events \Vere ,JCcompanied by .1 much highc..·r th C<ll(l,/ll(l'S, V.
·- llltul \'anJbility. and thl'\l' \l't.'lll tn have taken no sea (section 11.3) Tlm pdagtc trilobue. however.
mar~ th.m 50110- 5011 0011 ye.tl'\. l3ut it i' aJwJys pos- 'howed .1 connnuous and ~adual dunge through
h! that smm· of rhc \ hJngt.'' rl·conkd .ue not truly the same succemon. and the ftct that it due\ ~o
~Ctlt': 'omc mtght hJ\'c ht.·cn Cl npht.·notypK - a demonstrate~ that the JUmp~ be m ecn the olenid
purd) dc\'dnpmenr,tl n·,pomc of mdiv1du.1b to a spene' arc real, and nor rhe result of hiddc:n ~cd•
d .~nt'lllt;
.. Cll\'lr<lllllll'llt. mentary breaks. Thi' 1s the \ame succco;,ion in
There art· orhl·r t.·x.unpk,, e.g. iu rhe C'retaceom which we '~e both ~adu,tli'\tlc and punctuauonal
!Cl ur,hin Di.,C<ltc/t .1 (~1111th and P.lliL 1985) where change. The two modes of evolunon :m: n:al. and
mull-~~1c chJngc' th.tt t..lll bc t<>llnwed through the apparently competmg theones ntrn out. after ill.
aruc ww.• impo,ed h} cnvironment.t.l c<tme' alone. to be complcmcntuy.
The1e t':m he corn:latcd with .tltt•ratiom in ~ediment Phylenc change\ seem to be Important 111 rhe tine
al pc-'' hcm·ver the ~cdt~m·nt w,l\ rnudd.Jer, the te~t tuning of spec1es to the1r hab1tat~. We should con-
1- .n higher. Sut.h d1,111gc' .m· ct. ophenotypic. not sider howev<.·r. the followmg quesuon. What sort of
IC , )lutmnan. Yet otht.•r n .. unplc' h.tvc been docu- environments ,\re likely to fJvour punctuauonal
ucnted of tml} gencuc t. h.utgc. of whtch Shcldon·~ chJngc and stam and whteh would tavour gradual-
h ,!9S7, l9H8) study ol OrdnvtCJ,m trilobites from ism? lt h.1s been mgge\ted Unhnson, 1982) that
)[ etmral Wab (sedion 11.3) is cornpdling. These punccuational change is mo~t likely to ocet1r in ben-
·d fo"tl'onut in grc,tt nutnbns .md <trt.' 1uperbly pre- thic, and grJdual ch.1ngc 111 pelagic, environments.
'mcd m .1 virn.tal!y <.t>tttinuous ~equcnce of black This would seem to bt• tnw of Fortey's trilobites. If
1hah Tht.· scqut.•nce 'tmlit.·d 'PJnm·d \Om(• 3 Ma evolution 1s largely a marter of :td.1pting to chanhring
~~d 111tlun H ~n· e1gh1 Clllltlllon trilobite linca.gco;, c~ll envtronments, mtght we not expect to tind abrupt
I \dmh 'hO\\l.'d a tK't innt:J\t.' in the number of morpholog~c a) ci1JngL'S m environments where
nbi m tht• rygiJium. lt io; .l \lriking mdtcJtion of physical ch,tngcs were hkt'WI'\C abntpt? Conversely,
gndull c\ olurion mwrring in p.lr<tllel m the various pclagic environment!. and those benth1c reg10ns less
er.t and madt: all rht• mort· rcmarkJbk by charac- disturbed by v10lcnr changes would be expected ro
e. ltr rt\rerul~. i.e. tt.•mporJry derrea~t· in rib number be more the locus for gradual :tdapr1ve change.
1C t()l!l rune to tirnt•. Gr:~du.t.l e\'olurion does not nec- Shddon (1990a,b. 1996), \\ 1th the dt•ep-water enVl-
tsllrl!)· httlll unidirt.•ctwnal dtangt.''· In another ronmcnr of h1~ Welsh cnlobac~ in mmd, would
IIUd) spaumng Jbout thl' ,,,me length of time ao; concur wtth the latter pomt, commenting th,lt 'per-
ld Shedon\ (c. 3 5 Ma 1 trilobitt.·, from a Middle and sistent phylcnc cvolunon " more charactcrisnc of
iy L'ppcr D~\oman \l'dton in France wc:re examined narrowly fluctuatmg, 'lowly changmg enVlron-
1- Fast 4lld Cla1k,on, 14~tJ) . Hnt· wnodout evolu- mem,'. He sugge~t,, howcver. that \ta\IS i\ more
111 o .1 \\J\ used a\ an tndt.·pcndt·nt control of bios- likdy to prevJil 111 morL w1ddy Alll:tuanng and
Ut b'J';lph¥. ThC\l' tropldllt..llrvphine tnlobttes .tre r.tpidly cllJnging tnvlronmcncs Quttt.' possibly
~h uml as t\\\1 'l'p.tr,ttl' .111d \Utll'Ssivt• hnc,lge!-, and in some morpholog!L's ,trc st~ltlt: and comparanvely
·d Mth the t'Yt.'s be(;,\llll' rt.·duu:d m '1ze and tin.tlly i11lrt within qlllte wide l.'nvironmt.·ntal bound;Jrics.
non·htmtlonJ.I. Thts tlloty bt· coupled with the They may be comp,tr,HJvdy unaffi·ctcd by 'nonnal'
d,•phon oC an mlaun.tl, burrowing habitat. Such cnvironment.d c h <tngt·~ and it '" only when the
.hm~ts art uJtcrpretl.'d as 1-.rr.tdual ,md it is the pre- broad bound~ of tht'St' arc CXCl't'ticd that we might
,('. \lt' ll!Jtigr.tphtc,ll tOntml prov1ded by thl:' con- expect ~pcciation events to h.1ppen.
4) •'11~' wluch cndble tlm to be \O
.e. l'nlups cht• clc.trc:'t indit.1tton of bl1th gradual Cc· evolution
c- d punctuated e\'uluuon occurring together IS Many species arc eo-adapted to life in .111 mtmlate
For1 y\ ( l!.IHS) work upon Lower OrdoviciJil tnlo- as,ociatmn wnh ,mothl'r spctu:s, mually nf .1 totally
of t' m ')pu7.bcr~cn. • kre tht.re are many olcnid different kmd. Such as~OI.tatwm have amen by a
·a. tn: c tn. I he\ .m· .11111mt certa111Jv benthtc. and kind of complementary protn~ known a' eo-evolu-
ed the) shm\ a cl~.1r pane m l)f 'tqm l~t. ~volution- the tion and the ,ldaptJnom \)t the t\\O speoe' con-
rta btupt lmgm of new ~Pl'lle' tollo'' L""d by \ta~t~. In cerned are e"enually l1nked . A predator- prey
")tl s;tf11C bc:d~ one lint!, the: largl.'-l'"}'ed tnlobite system I) lme sut: h t'x,unpk, <I\ .m: sy,nbmnc and
.111 r:ua, "h1eh \\\am 111 the upper "atcf'l of the commemal rd.momhtp,, and of Clllll'it: there .m.· the
Evolution and the fossil record
mutually n en~~san n:latinmh1p' lH:twl·cn flowering cake place at any lt:vd: the genome. the individual. when tollowc
pla.ntpecte~. or above. Moreover, pt oct:,~e~ ~uch J uftctt a net 1
In \'lCW of the t lO\l' <.'thtdapt:u1on between selection and drift can opcr.ltc al all thc\e kvch. We may be sever
~pene~ a motMitarion of one will promme changes have already comidered \Ollle ol the mccham~Dl.l be reversed. ~
in other;. Tlm Js de.1r, fl)r e.x :unplc. in the evolution operating at the genome kvc:l, and we have \ceu plex:ity of the
of mollusc~. perhap' the Olll\l cnum·ntly edible of all that rnicroevolurion 1\ LOntcnH:d with changes ~.;hanges l vidt.:
mvcnebratC\, tn "h1rh 'evolutwnarv t-scalanou·. an \vlth.iu specie' o r w1th mmor taxonom.ll' ttallSiriont kno~Vtl for a I
'anm race' between prcdawr Jnd prey ha~ been a Macroevoluuon, on the other hand, refef'i to cvolu· vanous way\.
dominam control of their evolutionary history cion .n higher level~ or, to me Levinton's ( l 983 these trends
(Venneij, 19~7) V.m \ .1lcn (1973) developed this definition, With 'the l haractensnL tramitJOm th.u ~pecie) \qth
theme in a d1ffcrem d1rertmn. He drew up plots of diagnose difference~ of m,\JOr ta.xonomJC rank'. lt 11 lm survival
species surVIVOT"\Iup, 'hO\'-IIlg how loug the compo- in the study of 111<1lroevoluuon that palaeontolot,'\ J aptdl} and I
nents of an original \llllplc of orgamsms survived has, perhaps, its mo~t dircct lOntnbution to make. upon many fJ
ilirough time. Rather surpmmgly, he found that the Bearing in mind that \everal thous:tnd generation.~ of generation
likelihood of extincnon of any specte~ remained may have hved and d1ed during the time of dcpos1· thc~c ch.lr.lcte
constant; a ~pecics nught bClQllle extinct at any rion of onl y 1-2 nun of sediment, there ts no way Ill periods. Vaci.
time, however long ago tt h;\d originated. In other which a rruly fi ne locus can be obt:.uncd on eveut1 tton rate alot
words species do not neccs~ilrtly have better chances which took place during rh1s wne unless scdimentJ· .1tfect the dev1
of survival because t11l'y have eXl!itcd tor a long time. tion was very rapid. Detailed palaeontological srud· or dade.
Consequently spec1es must conunuallv be m a state ie) are better at re~olving the more maJor features o An impom
of evolutionary Aux s1mply ro kt>ep up with each evolution : species selecnon, the ongm of new struc· m.teroevolu 01
other to achieve ,, constallt bal.mce. Evolution is tural plans and higher tJxa, adaptive rad1at10m and Thi\ is becaw
thus forced mamly by htologtc Jl factor;. This is the cxtinctions, and r.1tes of evolunon. at thl: microe
Red Queen hypothe\1~. b.N?d upon Lewts Carroll's ~·qu r v.tlcnt sel
fomlidable per'lonage m llrrl' t/lftll~(!fl the Llllkm,\! Species selection large-scale p~
GltlSs who told AJ1ce 'NO\\ here. you see. u rakes all We begin with speCies sdecnon. \liKe tt is here tlut \nth phylet1c
the nnuung you cJn do tO \t,IV in the qme place·. the long-renn dttference~ berwecn mtcro- and Though spec
The R ed Queen hvpothc'i~ m:tkes the .lssump- macroevolurion are mo~t dc.tr. In nuuocvolutJ( 1 lllJCI'OC\'O)Utl<
non of progress, rhat !Jrer membcr; of a hne..1ge will the umt of selection i' the md1Vtdual, v.tn;ltlon~ an there j, aim 1
be better :tt \UrvlVIIlg than earlier one,. For molluscs from mutation and rccombm.JOon. and naturol where ba'>ic.ll
dus seems reasonable. For other animal' this selection operate\ by tavounng md1Vtduals w1~ gcnc..·ral direcr
assumpnon 1~ more ditl~rult to tcst, e\ en at the character com binatJom ' u1tablc for ~urvival. At th( dnnp:trable tl
macroevolunonary level. where mmt of the current macroevolutionary level, howt'Vl'r, a~ Stanlc:y ( 1970J level. These :u
debate hinges (Bcnton, 1990). An alrcmativc model has shown, the umt of \clectJon •~ the spccu:s, th sdcnion rem.
IS the StatiOnary hypothl''l~. Tlm propmcs that evo- source of variation IS in (rap1d) speciation event' pnrnary mod•
lution is forced mainly b) non-bJOlob<ical, cnv1ron- and selection acts upon o;pccJl''i with t:wourabl lonl{t:r time p•
mental t'ICtOI"\. In an unch.ll1b'lng environment there character combinalions. Th is is species selection. h
will be no change. Spet iation, evolutionary burst~ is, of course, analogous to natur .1l selection. Uu. Origins of I
and cxnncrions only h.tppen when the physical while rate of reproduction and potential for individ New types of
environmcnr change~ Tcst.lblc data remain fairly ual survival operate at the individual level, rate (/ of ammal o rg.
equivocal, bur as Uenton (J IJIJO) shows. continuous speciation (i.e. birth of new species) anJ spct.:lc C.1\\tl record,
'constant naming' m.1y Jmr not be pmsible for many longcv1ry are detem1inant~ m 'pcuc' selection. pcnod of evol
lineages in v1ew of tht• rumtrunional and develop- So having defined spc{ic~ ~election, how does 1 smalL dctinint
mental limits of the organi'm' them~elw,. operate? On the pumtuatlon model, spcciatJon ds. Much lJrg
events produce JJscret\: sp~'Ctes which, alter a bn( m the ongmal
and geologically instantaneous pcnod of flux. haYc phyla. Even
Macroevolution their characren. defined from the !~tart. Thcrcafr group~. all th
there is usually a long penod of httk· change. \'lnwl <Jnatornical, m
Evolunon 1~ :t lucr:trt hila I phenomenon. m the stasiS except, perhaps, for a lmlc: ph) lctll tine-~ lC'.Il elements 1
~cmc thar l hangc~ of cvolurion,try importance may ing. Widun .1 plcxu~ of related ~pent:~. howcH7., fllUillOU\Iy eo
1hen lollowed through long nme lntcrv;~l~. there j, The originatio n ofh1gher tax,J, in tlm 'cnse, n·m;um
olt~n ,1 net cvolunon ary change or trend. There the least understoo d of palacontolog.cal pht·nom-
IIU) be ~cveral couplc:d trends. \Ome ot which c,m ena. It is sometime~ referred to as mcga-ev olution ,
ix' fl vcr;cd. ~uch trend' (e.g. the mcrcds~ m com- though the term t\ seldom used nowaday~. "s 1t
pleXIty of the anunonll t' suture. or the numerou s ~eems to be another form of macrocvo lucion opcr:tt-
ltu1~,:~' n1dcnt 111 graptohtc phylogen y) have been ing .1t a higher level, and it ts hard to !..now where to
lnown for a long nme and h:tvl been mtcrprete d m drav. the boundary . When more is b..nown about the
unom WJp. In modc1n nucnwvo lutionary theory fun cri on~ of the gl·nome, and how the inlormati on
hc1t trends rt·~ult from speClt'S selection . tho'c earned in the gencti(. code results m J tulh dc.:\·el-
~ll'll w1tb cluracter lombina uom .uivamag eom opcd !ndtvldua l, we 'hall h.we more of an m~1ght
for 'urv1val bt•mg sele(.ted through ome. How into this most critic.d yet most dusivt: of all a'pects
nptdly Jnd how cfiel.tJvcly tht·~ spread depends of evolution
IIJIOI 11.1ny .3ctOr.. not the lca<a of whtch 1s the rate The links bet\wcn h1gher taxa arc obscure, Jnd arc
of g\'lll'r.lrion of new ~pel ies whKh happen to have but poorly rcpresentc.:d in the fossil record, 3!1 excmpli-
th~r ch.lractcN and the tcndem y tn surv1 vc tor long til.'d by the divemfication of hfc m thl' earl)
pq~ods. Vamnon m spcu.mo n r:ltc and m c:x~nc Cambrian , where tran.sitional or hnkmg fonns an:
r.un rat~ alone, or w1th selecuon opcr.ltmg . w1U absent. The geologtcal record give~ no indic.ltiOll ot
,tfm the developm ent of ;l ne"" l'voluuon .uy ple,us such relationsh ips, .md our knowlcd~ thcrl'of rl'St\
.xcudc:. cnttrdy upon the tr.ttlloonaJ dt~liplinC'i of comparatiVI.'
AnunportJnt roncept, srresstd by St.mky, IS tlur an:~rorny and embryolo gy, \Uppleme
nred, to ~omc
m.uo~volunon 1~ decouple d from minocvo lution. extent, by chemical t:t.xonomy. But what the fosill
Thu I\ bclau'l mdtvtdua h are the um~ of ~dectJon ren>rd dorc: give is many example' of
the 'mst.lnta-
~~the ll\lll'Ot'volutlonary level, .md sprucs are the ncou~· origin of new structural plan.~. The dcnvation
1qutnk·nt sdct tion umt' .lt lup;her lcvck H tnce of monogr.tptid~ from diplograp
tich, the bcgtmung s of
~r..·wole phylogen ctll trend' are nothmg to do LITegularity 111 cchinmds by the abntpt d1~pla<"cmem of
1 th phyletll trends w1trun .1 particul.1r spene~.
tht: periproct , the fi1sion of the mantle in b1valves to
lhnugh spcucs \dect1on is the ba."' of much fonn siphtms: all these arc importan
t, sudden key
nunocvolunon.try change, tt 1, not the only .1gcnt; mnovaoo ns which allowed
great evolutJOn:try dtven.i-
ltr~ 11 al\o what 1s known as dtrccted .,pea:ttio n, ficauon thereafter.
llhl·r~ bJ\icall}' r.111dom changc~ move m the s,tme Such changes a\ these art now understoo d 111
grnenl Jm1tion. and phylogen cnc dntt, wluch ~~ tenm of changes of rcguJat<>ry gene~ which
control
complrJble to gcncot dntr ar a m:1croev olucionJr y the operatiOn of structural
genes. Mori.' importan tly.
kvd. lhrst· Jrc less impurtan t, however, Jnd spl'cie~ evolution ary novcltJts may arise
by relatively minor
lUOn rtnt.um as 'mtl roevolun on m aaion · the changes in the regulator
y gene compkx. Tlm might
rrunm mode of m<~crocvoluoonary change. over affect a character or group
of character s, hut such
longer tune period~. regtllator y gene changes nught. on the other hand.
affect the whole organism .
Origins of higher taxa A well-kno wn process th:tt is pn:sumah ly con-
Nr11. l)'pc~ of mucmre, .tncl new gr:tdes and system\ trolled by regulator y genes :tnd
seems to have been
r Jntm.1l o~an1urion. appear very suddenly in the importan t m evoluuon IS change
in rdatl\ c growth
.mil record, teH1!)ring eo an inmal and very rapid of d1tferent parts of the body
givmg the dcnvative
r nnJ nf cvohtr1on. St11.h changes may be relatively stock different proportio m. Sulh allo m etric
llil, Jt:fming new group' l t lower taxonom ic lcv- changes nught be sudden or gradual, and nu~ht wdl
Mud1 br~cr changl!',, of great unportan ce. rt'sult account for much evolution :try chan~c.
ut tlu ongmJtJl)l1 of hi~her tax.1: orders, classe~ and Abn1pt and major change~ are less easy to under-
. 'i' hen m the tiN rcprncmatwe~ of the~c ~t~md. and have been the subject of much
di\CU,~ion .
up!, all the ~yscems of the body, and all the In 1940 Goldscbm idt proposed
a novel and cert.1inly
allallltlltc:Jl, mcchmica l, phy~iologicaJ and biOchem - somewha t £1nciful concept to ,1ccou11l
for the \ecm-
tkmcnt' therein. ha\ c to bl' prcn~ely and har- ingly mstantan eom origin of new speues
and mdeed
mou'l) l'\lMdinated from the VCT)' bcgiruung . h1gher tax:t. He held that
major chrornmo mal
mutarim" or rcarr.lll!,;l'llll'lll\ c. ould in rJrc.· cases pro- the aduJr tonn. A tl'rtain ,11\ti.'Unt of n:ordcnn•
duce .1 'hopdul nHm~tc.•r', J viJhlc new kmd of the lcm-p.tcJ..mg 'Y'll:lll, .1 lmle tine tunm~.
organi"n wllll·h nmld be the progenitor of an there we have J trm. ph.Kupid l'\e Wlut we c.mn
cntireh ne\\ t.l'l.llll with tt\ n\vn chJractcr. kno\\ i~ the 11.1tun: of thl· 'dnuon pn.·"urn 'du
Gold~chmtdt \ view~ wac m·vcr popular, largely led to thi~ lair!) dramatit rc.·mhranuatwn 'We t:~ll
became ofthe \Cl') lu\\' probJbilitv ufsuch a 'freak', sure, however, due ~imc du~ kmd of e)e \VJ~ tunc
even if H luppcncd to havc ne\\' JliJpuve characters. oonal m the JUVeniiL· ~t.tge. 1t mmt h.tve he
being able ro hnd .1 mate and brcc.d Yet there is still equally funcnunal "hen it w J\ adoptt•d eh m~
some support for a mo~.htlcd <md toned-down ver- paedomorpho~ts, as the Jdult t) pe.
sJon of the 'hupcfitl monster' concept. and it IS not T lus j~ JU't one t''-.!111ple. There .tre n1.1ny otheo
impos~ibk eo envi~age .1 vi.tblc .md 'not-too-fiTak- Pacdomorpho~JS ha' oti:l:n been 1.0midered as bea:
i~h · freak, breeding inmally wtth ~umhr novel otf- responsible lor the onwn of carl}- hsh-likc ame)\[lt
~pring or the \Jllle pan·nt Ill J. \lll.l.li population, as of vcnebrate~ trom the tadpole-lilo.e, tree-~w~
the founder of :l new l.l.'XOll. fC\\ would accept larvae of sea ~qutrts. L1kewt~c the:: \mall pbnktullll
Goldsclunidt\ tdca llt tt~ ungtnal and extreme form. copcpods may be products of the early omt:t Rotes of E
bur tt i~ qwtc pos~tblc th.ll a comparatively few matunry in plnnktontr larv::te ofbenth1c crustalt:.lltt extinction
quantum events over ~everal generations could have The importance of heterochrony (Could, JCJ71 l•vol ut 11 Hl.lr
led to an emirdy m:w type or organization. M cK.iJmey. 1YHH; Mll{inney and McNam.tr.!, I()() fi ·•m t 111 1 ~ 1 .1
Moreover, the modern vat·w of the genome, as a cJJulot be ovcrsrrc~sed, '" it can account for ahru1 nd D.ey. II.J
hierarclllCally orgamzed .md dynJnucally imeractivc evolutionary change5 ,H every Jevd of org.uuzatt\ll
~ystem, allow~ for llllll h tllOH' coordmatcd flexibil- Paedomorphom may be seen a' a spenJI c.e\t 1
Ity than would havc scc1m:d po~stblc a few years pre-adaptarion. The COIH:t:pt of pre-o~daptarion,
ago. lts simplesr fonn, mc.'.ms unly th,et .ll11111.1l~ utll'!
We llJve ,tlrcad\ com1dned tbe importance of already pmses~ character~ capabk ofbeing modtli hang' 1~ 111
heterochrony 111 the on~n of nev.: spenes. It seem.s 1f they are gomg ro be able to .tdapt to new envtroc su(• c~,fu l
to be even more so m com1denng the origins of ments. Somenmcs d1e'c characte~ may be m th: tnnc.: (11 u.tll
higher taxa and ot nev. wstems or organization. ln tom1 of organs rhar have lost tlleJr ongmal fimcuonr Jd<lptl\ c Ill
many ca~es it pro\,des an amwer to the emgma of they arc therefore.- ·~pare pam' and c.1n re.aJi. t111111) 'hu
how a hVlng or~ani~m c.m abmptlY produce a become adapted wHhout mh1binng an eXJ,u rxp;\lld the
desccndmr fonn, which nuy be qutte d1fferenr m funcnon. More oftc.n, howc.'Vt'l, they arc 1lrc2~ n " glt1llp!
some re,perrs and \l't 'nil j, a fimcttonal. coordi- funcnonal. bur by bt·commg mod1tied they Ill 10 11):111) \.'1
nated orgam,m. ~ krc " one e'Xamplc. The eyes of then become more flexible tn tlw v,mery of tun- :ulaJHivc t
most mlob1tes (st'rtlnn ll.j) are holochroal; that 1s. ttons they pertonn. 1 here ,m: mJny good ex.unpl \Oiuthm.tr
they h::l\'e many connguous lemes closely packed dra\\11 from the vencbrate~. ~uch ,1\ the use of .1 t, 1;a eh~ telic
together upon the \'ISU,ll ~urtJce. flm ts the primi- dtverticulum :1~ Jn .ur-brl'Jthmg lung in Dcvwu r nt ~p~·, ie,,
tive type of eye whtt h on be traced back to the and later lungtish hvmg 111 nvcl"\ .md pond., th;tt .er I] Ilk f ...
early Cambrian. Tht·rc .1ppt·ared. however. in the liable ro dry up; the ,,,me strunun· became tht: ~wtr.1 '\d tptl VC
early Ordov1cian .1 lll'W l-md of eye. confined to bladder in r.1y-timwd ti~b. •l,ll'llll .tlly
phacopid rrilobtte~ :tlonc Thc.·sc 'l'htzochroal eyes Numerous ex:unplc~ could aJ~o be nted in tht J! ahllo'l .1
have relatively much largt•r knse~. not very numer- invertebrates. the rcmarl-,1bk ad.apt.mon of olreaJ l ll g ly tllll\t
ous, and separated from c.tch other by cmicuhr exi~ting '>pine~. ci lia and tube feet in echinmd, for d1\ ,·rsiltL otl
material Thev loul- vc.·n: d1tlcrent trom holorhroal deep burrowing hcmg an 1dcal CJ\e :unhrurt,
eyes. lt 1s at ti~t ~1ght very lurd ro ~cc how the H owever, nm only may 'rructllrl'S be prc-Jdaprt km1ng the
schi1ochroal eye <ould have been dcnved from any before moditicatton: thl·y mmt .1ho be,·omc post clc11 , ,ri~u
holochro.tl prccuN>r The amwn lie~ m a 'rudy of a dapted while the1r timttton ,md probably forr Crll\1 ILC.IIU
the onrogcny For thl· Jun·nilc eye of an aduJt a~ well - are 111 the prnn''' of bemg tramfonncd t x;~mplc. d
holochroal-eyed tnlob1tc ''· m dlcct a miniarurc Prc-adapution h '' be, omc rcrognitcd a\ a vr~ the J11r.I''Jc
schizochroal eye lr h,,, relattvcly l.uge . .sl:paratc and crirical concept. indeed Cam·r ( 1954) hJs \\ nttro \\ lltl h 111.11
few lcmc~. Arre't it~ development by paedomor- 'No adapt.tnon IS pos\Jblc without mme grJdc of ,11\ t'rsttiCd
pho~IS, and the JUVenile ·~rhl7orhro.tl' eye becomes pre-adapunon m the 'tnll'turc to be adapted' from t lw
.~n earlier lOnccpr rcl.trmg to ontogl'llY .1\ ,1 tool .tdaptive radiation " commonly J new \tnJCturnl
nntlt l'lt.mdut~ cvolurionJI')' rl•l.ttiothlup~ ,,..1~ plan. ln mannc orgamsl115 adapove radiations .1re
.ofHacckcl, who bditvcd th.n thL' onro~l·ny of very ofi:cn al'o lmkcd wtth marine trJn,grc~\IOm, tor
anm11l \\,1\ ,1 more or k'>~ d1rcct rcL.lpltul.mon if new space IS created and new environment' an:
111 phvlugcn} [n other word~. the yl"nmg \t;tges of generated then c.1pportunmic group~ will anse ro
J.vdop111cnt ot .my :minul Jre very ~tm1l,1r to wh::tt colom7e them.
~e Jnce~lur~ of that species were hk.e. In emhryol- Once the initial burst of diversification j, over
11 and 111 p.tiJeulltology there is verv httk cvtdL'ntT and rhc main groups arc differentiated. thL· r.ttc of
tht\ (rhou~h one or t\vo pm~tblc Cl~l'' .lfl' dl'- species production slows down, ::tnd the spenes tend
~~ed \\ nh rt:fc:renu: to .muno111tc\ 111 \ll tton H.4), to survtve longer. Whereas the new tax<~ ongtnJted
:d illtte the widespread acccpt.l!Kl' of hctcro- by quant~un cvolurron dunng the early stage:~ ol the
1), HJcckel's 'biogenetic law'. '' 1t " ' ' ' c.tlled. rJdiation, it IS dunng the later stages that ph)lcttl
bw1 Vt'l)' largcl~ di~carded. mteroevolutton:uy change can pl..ly a part, for 1t
allows :~dapt1vc tinc-tumng, such as slow ch.1ngc: 111
Rates of evolution, adaptive radiations and size through time or alteranon 111 the tmunt; of
extinction breeding cycle~. I f a parocular spec 1 e~ beconu:s
·nlunon.ll) r.Hcs within evolving group~ .trL' fi1r extinct, it wil l probably be replaced by a new one
111 wnstant ..1~ has long been kuown (CJmpbell from a related stock and w iU become tuned to its
Wl\, I9X7) lt we comtder evoluttotury r:HC'> in environment 111 the same w::ty. lf the ongm of new
1 uf produ((ton ot new ~peLll''· ,tnd hm11 long speocs more o r le'' equates with the exunct10n of
U\a pt:r\1\t, a conunon Jnd genl·r,tl p.tttcrn older onel> (horotelic evolution). the lmeage will
~ 111 most t"U,,il group,. Tlm I' th:n 111 the ini- survtvt· Jnd rt'lllJIIl v1gorous. If spectes .1bundance IS
Vlgc' {ll thL· hfe of a higher uxon, e\'olutionary plotted aga1mt nmc, rhcn their dttferenttJ) survtv.U
~e ~much more rap1d rh.u1 tt " later on. Whl'n or extinction w11l gtvc rise to clades (i.e. clmtt"rs of
ue~ c"ful ne\\ .mnnal group Jppc.lr~ ltlr the fir.t branchmg linL'ol~l'') of particular shapes. For ex,tm-
nr (U\U,,Jiy wtth .1 new body pl.tn 01 wllh J key pk, when initial evolutio n has been raptd, and there:
1 pnvl' mnov.mon) rh ere
~~ ;1 gt c:at uutt.tl 1. vulu- ts thereafter a slow and gr:1dual decline, the rcsulrnnt
h)ll.ll'\ 'hnrst'. New taxa proliferate l,lptdly ,md pattern w ill rc,cmblc 3 fir tree in outlutc.
rJrllt thl'lf t!l'O!!fJphicaJ ran~c qutl kJy, ,11\d the Conversely, thouf!;h rarely, a more gr.tdtl,tl tncrcJ~c
1 1 grmtp' 1mm~ from tlus burst bet ome ad,tpted m number. of ~pcncs, tollowcd by .1 re l.ltlvdy
m.tll) 'llVIr<llltnl'llt,. 1Jus prOll 'i~ IS klHl\\ l\ ,1S abmpt extinction, produces a pattern of p.nntbn1sh
WptiH' radiation and I~ always J time ol r.lpid shape. More commonly. cl.tdc shape resulnng !Tom
llOlllf} , hangc (sometimes known as a rapid inittal bur.t, .md c>."Pamion. ft)llowed by .1
tKhrtelic evolution) and oflugh pn.ldunmn rates fairly rap1d dt:t linl', though wirh some members
pcm·,, gcnaa and pos.s1bly Jl\n t.muhe~ and persistin~ tht:reaftcr, over long time penods, t.1kes
I\ the fomt of a sprouting mowdrop bulb. In such .1
A,l,pttvt: r.lth.ltH)n may be vtrnl;lll) glnb.1i or geo- case the :.urvivor~ continue as rclicts long .tftcr the
phKJI1) t]Uitl' lot·ali7cd Moreover, tt may occur parent group which gave rise to them h.1s dtsap-
lllllo\l .1ny t,r.,.onomir level, but it ~ ~ IIIO'>l mik- peared. These survivors are living fossils, continuing
;1\ tllmtr.ltrd by -.uch phenomena .1~ tht llllttJl in the virtual abse nce of evolutionary ch.111g<.'
tTilhl 1tton of tnvcrtcbrarcs 111 tht" early (bradytelic evo1ution). Such genera .1s the br:t-
bnau, rlll' r.tdiJcion of Ordo\'ICJ,1ll trilobites tol- chiopod Lin,l/ulcJ nuy bl· comidcred here Tlm genm
mg !lu.• l.ttl'l C.1mbn.u1 cxunctiom. or the md- was already tn ex.i\tcncc m the Upper C.1mbnan .tnd
(lngm .mJ r.1p1d prohfCr.Jtmn of lugher smce then ha' dungcd relatively little. It IS Jblt: to
tcan' m the t:arl) CJrbonifl·rm" A further mrv1vc because early on in history 1t colomzed J
plc:. though on .~ more 'pread-out ttmc,c:J le, 1s 'dtfricult' but per.1~tcnt envirownent. that of mud-
lurJ"Il ~:vnlunonary c:xplo,lon ot ~t·:t urchm~. 111 fl,JtS and shoals expmcd at low tide (though o,pct ,_
u,h lliJny h11!hly 'ucn:~~fi.1l :tnd pt:r\l~tent orders mens are known from deeper waters). There wa<.
f\'1'\llicd fi·nm .m 1mti.t.l stock which had \lltvlved relatively little l.Ompcntio n tor this since the physw-
llll dtl' I'.ILtL'OtoiL. The initial 1mpetus for an logical problems impmed by Auctuatmg salinity .111d
period1c ~o:xpmurt: arc 'c:H·r~ But b~cllbc Lill,l!lll<l Evolvmg lm~ages in the fmsil record 'ometlllta'
wa' ahk to ems~ till\ .1d.1plln thr~shold, which show pattem~ of direcr:ionaltty collcwvely known
other mvcrtt·br.tte' h,l\ t.' r.1rd} been able w do, 'evolunonary trends' {McNamar.l, l991l.l,h). Tht
there 1' no rc.tson whv it should not go on living in may be ,tn,tgenet:Jc, occurring 111 ,1 ~mglc lll'l\0 1>.11"\ Ill C
\uch an environment imktimtdy. lt IS unlikely to be branching lineage, or cladogem·nc wh1ch mvol\ 1111J1t ort !Ill
displ.~t~·d by competitor... ~pcci.Hinn events. In some ca~c~ the.: rhJnge~ Lll!llt C\Pillp• !Ill
~ontt' 'br.H.iytelil' .\llillhlh now live tn the quiet about through mccessive paedomorphil ewnts <.0 Ill,\!)' ' Ill
and stable w:tters of tht• dc~p 'ea. I le-xactmeUid we h:1vc 'pacdomorphocline~· Some t•xamplc.:s .m T!m ~~ 1c
spongl's ,and the mrmoplalophoran mollu~c o;ummarizccl in Chapters 6, H. 10 and 11. thouw Ulll11' tit
\'c•clpr/111<1 (who~e most rl."cent los~1l relative as Mtddlc more fully treated in McNamara ( IIJIJOa,b). Man tnd thl' 4:
))~ vunian) arc .ml~.>ng-.t thc't. But the deep sea. trend~ ~cem to be adapnvc, .1m111g !Tom coutmul'd efl,llll k
thou~h J sublc t>nough env1ronmcnt and a haven natural ,election m a partitular dircltiUn. Yet tlu ther 11.11
tor rdJLt stodcs, is not .ln 1d~.1l home because of the role of ad.lptanon ha~ been called lnlU que,rmn. JJ tndl\'ldll,
rebnvdy limited food supply. rh o~~ Jnllllah which at le.1~t ~omc may be the result of rJudom \Orttn~ ~tlppl)
t'::tn copt: with mch unpopular t'IWironment~ have a alone. ht.:t\\ I.'(' I!
t~ur chance ofbecommg br.Hiytdtl unJl·r the inAu- Throughout the history of lif~ u1dtviduJl t.m l\ll) uf ,l
etKt' or \ t,tbli iZln~ sdcction. have originated, floutished for a longer or &horw Jh'{ ll'~ 0
ltdl ct) of once Important group~ livmg thus may ti111c. and then become cxtincL. The demtse of md1 111)11 I\ Ill
be re~tnc ted to certam h:tbttats. bll( this does not m·c- v1dual specie~ has lcfi little trace upon rhc bru.1d pat· Sllll(lhU
l'ssanh mean that such 'hvmg I0\\11•;' .1rc dcdinmg: tcm of evolution. Much more imporLant, howeVt'l .,f rnl
~ven rht> nghr condtnom thC''e r.1xa could go on for ha\'C been periods of mass exum:tion wluch arr Alll~.:lllU
ever Wht.>rea<>. as mennoned .1hm•c..', the..·} may mh.1bit dealt with from the geolog~tal and mattgraplu r here 1t
t.'nVJronment~ tor wluch thl·rc..· 1' htde competition. it viewpomt ~eparatdy in Chapter 3 Uut at tlm poul! lllt.:r onl
mtght he.> aJ,o th.lt the} h.wc..· ,1 "" tdcr 'nv1ronmental ir i~ worth stres~ing tbat when, through maJOr em,. An 111
tolt.>r.mce th.1n do other J111111.11, Ne..•\\ v1cws on the ronmcnr,tl cat.1strophes of one k.md and another tlhpl.al e
nature of hvm~ K1'''h .In: ~Lilnm.mzcd 1n Eldredge many k111d~ of organjsms arc lulled o~r sunultant'- 11~ 1 mge
and Stanley (I C)~H). Wtrh the.. c..·xcept!On of living ously, the whole evolutionary p.ltlern ~~ disrupted .md takt
tosstl~. it seems m he the lot of .111 t.tx..t to become When ~uch Cltastrophe!> happen. a~ they have mam ~ ltt11111<1l
extinct. In \t>tllc..' cases there 1s a kmd of relay of ti111c~ within the Phanerozoit., whole ct.my~tem• I.J[tl I h ~
spc..'c..le\, a ne..'\\ on~: con11ng m to occupy the vacanr collapsc along with their component o rgam<>ms. A., ah1 upt t
me he left hy the exu tll uon of.1 prccuf\or. In particu- the aJaptauom of individual speoe~. the result ot allnp.un
l.lr hrrnups then• 'eem' to he..• 1 dc..•hnitc correlation succe~sful mltlal evolutionary bur;ts and of subse· 1l IUVC \
bc..·rween .1 high rate of \pcd.mnn md :t high extinc- quent fine tumng over ulllliom of yc.>a~. are \imph llll'llf.ll
non r ne. An ordmg to <\t.llllc) ( 1979) there are two wiped out, togetl1cr Wlth all the dehcate ecolo~cal ~reocs
rc:tsom for th1s. One I\ th.ll spcltatton .111d extinction babncc\ between species. ltOII bt•J
are very high m \11\..tll, loc:tl popul..tuom. If J popula- When normal conditions are restored agam afret 111 the •
non become~ w1despread, tt limtt' further )peciation <>uch a mass extinction period. and especially if there ,Hhpt1VI
111 tht: ,1re:t tt inhabit.,, ,md 1ts broad geographical is a subsequent marine transgreSsiOn opening up the .mu
'prc..·ad is ;J bulwark agamst extlllltion. Though parts new sp.tcc and habitats, then evolution resumc1 ~tnt tly
of a wtdc~pre.Jd popuiJtion hvmg 111 local areas may based upon new adaptive r.1diaoot1s and nn the ll'~ttin<
d1e out, mamly due tn .Kcidcntal f1ctor;, the whole development of new ecosystems. Ma~s ex'tlncrions ,lfl 110[
popul.!tion 1\ le" hkd~ enher to bccOllll' c..·xtinct or to more than any otl1er factor, control lhe whole pat- Anot
.1llow the production ut new 'PeOl"> trom the same rem of the evoluoon of hlc, bv resetting 1t from 111 '>YlllJ
~tock A \ccond bctor c..lonunc..·nced by Swnlcy is that tnne to time u1 new and d1fferem combmatiom. 1: bnllflll~
rate~ of ~pnoation and cxrinc..non vary w1th behav- ~eem<> mdecd to be the case th;lt lll.lJOr phy~1cll l l\l' \()1
IOUr. \penalized bc..·ho.~\ mur pancm' Jrc dl.lracteristic or cnvtronmemal catastrophe\ have rau\ed m ore ~rc:tlc~
of p.trtll ular ~pedes. and tf Jll) 'PI.'lH..'\ bcc..omes too large-sc.de change than ha~ competltton between .1re ~er
~ren,Jii7~:d (~:.~;. in food prl-fen·nt:cs) tt may increase orgJtusms. may n>

rhc vulnl'mbility of th.lt 'pecie\ tL1 exLmction through So we ~hould now cons1der wlut rhe role ot pt:ting
evt·n minor cnvtronmcnt~11 ch.1nge. competition has been. t h ,\1\l< I
Competition and its effects 49
Evolutulnary COIIVl'rgcnce m.1y bt• mtcrpreted .1~ .1

stratt'b') for cocxl'tencc which would mmmliZt' tht•
;e dfeu' of ,cJecrion. \O rh.H 1t would upt:rate upon .tll
l- lhrv.1n empha,ill·J the rok- nf compctitu1n ,1, an the imliv1dual~ m th(' r.m~e of uwrlap ;b if thl')
e tmpOrt3nt Clctllr 111 n.1tural \t'k' rion . le j, de.u chat \\'t n· ju't one ~pet 11. ~ .llld not two competing one,,
IC mprnuon for food, hvin~ 'P·'' t ' .tnd nutc\ 1\ .1 pn- Diwrgt•nrc of t har.Kter~, 011 thc nrha hand, 'ug-
:o W\ comrol w1thlll a singk ~pt-cic' or popul.tuon. gc~l~ ~eknion witlun the two ~pct te' for ~lighlly dif-
:t.: ll j, rompcuuon 111 ~impk t'Ulhl~riral tcnH\. ~uch ferent lubitat,, 'll th.1t compemmn I' a\'Oidt·d l~y
;h 1petiuon 111.1) bt: JJrt•tt. in tt·m I' of .I~'Tl "'on h.1bitat prolifer.ttion.
I)' 'lllJ theN 1hh,hme111 of .1 hil·1.1n hil,1l onh:r \\ 1chin CJ't'' illu,tratmg the'e art• J...nm' 11 from tht· tC.l,,tl
·d :IWIJ kmds ot h1~I1L'r .min1.ll \OCJt'tll'\. On the rnord. In C:tmbn;lll .tgnosrid tnlohl ll'\ then~ IS l'\"1-
tc •ha hJJ1d 1t n1.1y be mdin..·ct, in tt·m1s of too m.my UCIICl' ut depth segrt•g.mon. :1' wdl a~ geogr:1plm .11
td i1\'1dual~ p.IS\JVt'l> exploiting the ,1\ ail.thk food 'epar.1t1on in dosdy rtl.lted spt:l'lt''· md at the ~.llllt'
tg pph. llut wh.lt art: tht· dlct'h of rompetttion tinu: \JZC di.,placcment 'ecm' w oper.'lte in ~e' cr.1l
~t"en 'recie\, e'pt:cially 1f \lilt' im adc, the rt·m- 'Pt·cie\. l t has been predtt"ted ch.u. in ordt'r to b1.· .In
ea 1) of another ur tf the gt'tl~rr.tphit·al range' of rwo ctli:rtJ ve '>trare~ry for ,1\'0Jtling cnmpcnt1ve ehnmu-
::r pr.le> overlap? Undoubted)) llltn~pt:tifit LOill~Kti- non, si7e displarelllcnt would m·~.·d to he in the r.1rio
i- '"'' highly tmport.lllt. a~ 1~ \\ illw~scd b) tht· rat.1- of ,tbout I: 1.::!!:! tor 'ympatric spt•ne,. The fos..1l~
l- ' ph1( redtlltion of the IIJtl H Bnnsh popul.1tton inve,ri~.Hcd, mdudin~ these .l~"llll\tllk ~~.·cm to
ud 'lJUirrd' .1!i:t:r the i11lloductton ot tht· come remarkably dolt' to thi., thcort'fKal tibrtln'
.11.re'· A.nJtn,Jn WC:Y \4Uirrd ltl the nineteenth ll' lltUr)'.
Tlt111' m no\\ vel) tew red ,quirn:ls left 111 Bnr.un .
Eld1t:dgc (I IJ74a) ha\ shown how dunng
Dcvonnn times iu North Ament.l there liYcd two
~t •ltrr only ;1bnut IOil y~.·ar~. spcot'\ of the large trilobite Pllrl!"llJI.' . P. itlii'~'IIW
1- \n llt\"Jdmg ~peuu may lw t'\pe<:tcd to t:nl to 111h.1b1tl'd the western area. \\ hcrt'.l' sucn·s\1\'l'
r, pl.i c an c,ubh,hcd \Pt't:lt'' • t11 t.tke m. Cl p.trt of allnp.nncally denvcJ \llb~pecil' of r . r.ma (prob,thl)
nngc and \Ct up a ternton.1l h.tlance. to intillr.Jtc origlluting as a Europe.m lllV.tdt·r) ocutp1cd till' t.:'r-
:i. d nkr up .1 prt·vwusly llll!lllllpic:d nitl11.', or to rain m the ca.sr. Tht· 1pec1e~ range of the two d1d not
l)' dumn.Jtc till' n.ltlVt' 'Pt'CJe~ t'ntlldy. How olit:n the nomlillly ovcrlJp - .1 t.l~e of llllll-tmnpetitivl .' gcu-
iiS ~ t<r happcm I\ not cle.lr, ft>l 111 the fossil recortl gr.1ph1cal separation- but whert: tlu: tv.o 'pccic., art
1 rupt fJunal brt-;tk, 111,1y \1gn1t) the mi!-,'TJtloll of an found together, fJ rrllhl aud 11• itlll't wis mutu.lll)
~f plmcally tknnd 'penes 111w ,lll area .1tler d1c rcartt·d: P. tllii'<'IIW dtvcrged iu morphology fTo111 P .
:- me 1tock ha\ become cxtmct th1 ough cll\'11 on- rrwll whtTc:as P rtllltl converged, thvergcd or \tayt•d
ly n,rn!.ll or other c.IUSt'S. Qtlltt t OlHDlOillY lll.'tlhcr 'neutral' 111 different dlJractcro;, exhibning a 'mixt•tl'
al rri~' become:\ exnnct and ttlmpctltJ.ve dimma- re,1ct1on. On the b:.Ns of these -.tudic~ Eldredge h ..s
n bctwc ·n sympatric spct lt'\ (1.e spent:\ hvmg suggested rhar ch.1r.1t:ter d1splau:ment nuv bt rHl
thC' \.11JJ • an:a) ma\ be .l\'CltdeJ by a vanery of mort· rh.m a 'magnif1ed trucrot·mm of the gena;~!
re ?U\'C str.ltqpls One \\'.1)' IS spatial sewq~-atlon; p.1ttern of mteraltJtlll between l\\"O 'petic' n t'll
1p he ~nmwh come to mh.1h1t ,1nd feed only w1th111 whcn ,11lopatrir'. prov1ded that .illopJtf)' is ~mlJinc<l
·<;, nnly loc.Jlizt·d ~eogr;lphtcall) or venic:tlly (depth) through rompeti tt ve exclu~ion.
lC '· tri<tt'd lubit.tl\. Hcnre tht.: mo ~ymp.1tnt 'penes Even though mch mechamsm~ cxi,t. compt·t1t1on
s. nor direct!}· rornpt tmg for ti.1od m linng 'PJCe. doe' operate eo tinUl ~penes Jnd perhap\ to lJU'e
(- :\nmhc:r 'tr.1teh'Y '' char.lt tt•r di~pl.tcement , ttthcr t:'\tlm non, at the mterspcofic. ccolog~cal k vd m
,11 Ill wmpatr11' \peric' or whcrt· the ranges of neigh- wh1c h we have referred. But doe\ H also opcrJtt' .ll
lt ' 111ring ~ptTICS ovcrl01p geogr.1pl111. ally. Jn the latt~r maJor extinction penods, and 1f so, how?
~I J e 'unJe r.1tlll'r t"uriou\ rt'.ll'ttnn~ between th~ two Sinn· competition has been tdentlfted .1s an evi-
re "' wuh tht•tr ~mubr t'l'\llllgJC al requarements dcnt .lgt·nr 111 evnlutmn, there w.J\ tor a long timt· .1
•n ntt up. Eatht r md1vidu.11, 111 the range of" O\'Crlap g~.·m·rall)', iflighrly held \'Jew that competition almw
com~ to look \ t'f)' ~imai.Jr to those of tht tOm- WJs perhaps the most Important t,HI~c Ji>r the
(Ull~ ~pcne,, or .Jlccmattvdy they may davtr~e in e\llnl cion of somt' ~rroups and th~·ir repl.tccnlt'nt
hmcters to ,1 much grt•atcl dq.,'Tee than nonnal. by others. Although competittOil undonbtl·Jl) l'i
.l"Cl'\~IVC al
impon.m t. the nlder view th.n pJrtlrula r t.1x.1 'omc- dectdcdl y higher than in a brac..:hiopoJ ftltenn~
rnut,1t1on .
how be< Jme nutdarc.·d .md "ere ~.1dualh rt•placcd app.lratm of the ,ame kngth. Moreow r, proct'\\1.'1
of p.trtidc tr.tppm~. sorttn~ and dtge,uo n 111 btv.livl, :tlh:lc•, '.111
by more. ctJlnc.·m l"OiliJll'tLtor~ ts no lnngC'r \t'cn as .m
arc highly dTcctive by compari ... on w1th tho~c 111 tlOJlS 11\liC
o~dcqu.ltc expl.uut ion for Jll typ~:~ of apparen t bc-cnt'l of
replace• 1ent 1 hough in so m~ respct"t\, tor cxo~mpk. bradnop oth, and bl\•alvc' have m any l.l~e a gn·at
enVJronmt·nrdl toleranc e. Thc,e .•md other f.\ctc•l"'. popultti Ott
tnlobitcs are bmlngn. all; lir111rcd, l.1ckmg stn.mgj. tws ~( nt 111 the
Jnd gtJ,pmg c.·qtnpm<nt. rhe'c: ,tnu1ul' \\Crt nmjmt coll.uc:d by Stl"de-P etrovic (1979), ,111 indicate thr
rel.lnve php10lo~cal ~upt'nont\o of b1valve~ 01
pnnun ((
fo r tht·l own wm:. but surd) tor .111 timL. n:m:lrk- dnvc 111ay
Jbly \>\'cll org:111iLt.'d .111d htghly f'untlumaltrcature~. brachiop ods and mgge~t rhat compctt tton cannot l'<
di,mi,\c d a~ a factor in f'aunal repbcem t•nt.
, lunge bu
Recenr 'mdil'' upon thetr function.t.l m01vho logv.
for ex.1mple . thor complex mcchani'm~ fur enroll E-.vcn \O. tbl" rl"pl;~ ccment of br.tl htopod' ~ Nn' '>P
biv.1lves wa~ not directly due to :1 compt·ritll ~~"" ( ont'
ment ::1' .t dd~nce ag.11nst pred.ttOT'\, 11Jrdly ~upport
t.tkeovc r. but the n·sult of one major eptsod~:. th tlten tn: '
the VtC\\ that tnlobttl"~ d1ed out 'imph 'bccau~r I .qu.tll\. J
they weren't much good', .1s one of tlte aurhor's snt- Pem1i.m L'XtlnCtton crim (Gould and G.llJO\o\J)'
,Jil\e .
19HO). Thb atTected brachiopod~ more ~~:verely thJn
111.1}
dent<> memor. tbl; phr.1sed tt 'omt· vcar' a~o. The croc hrony
htstol) of tnlobues , hke rh.tt ot 'o m.m} othc.·r bivalves - th\' lattt•r 'weathe red the srnrm' b~n~
due no doubt to their phystological ~upenurit\ lll'W ~pl'CI
group,, t~ pum tuated bv .1 ~en~:' of mJjor .1nd minor nfthl' pan
ex rim nons, \Oilll.' of w hKh can be dHelth rdated ro Compet itmn w,1s undoubt edly mtemtfu~d dunn
I V.H ,lilt I
phy~tc.llevent\ mch ,1~ mannc rt.'gTl"~\lom and not to thl Pcnm,H l .md rhc br.1dnopoJ-dom1n.tred cor
munitie~ were disrupte d. When the cn~1s wa' over
pnpui.IIJOI
compem ion. At <."ach of the m.1jor t'vent~ m:1ny dcctmn
tnlobtrt group' bcc.mte e:\\IIH t mon: or less sunuln- tht btvalvt·s. movmg from thetr origm:~l ne.lr.lw
~h.mg(' w
neomly. Tho~t' that were ldt continu ed to floun~h hab1t,tts, were abk to colomze rhe now vJC;\11t n~>
'P·Il'c. prcv1ously occupic u by bradllopod~. It w3.1 ~t.l'l~ IS eh
tor Ion<! pcnnu~ of tmw. Some.· of these were then pi:C
removn l dunng d1c: nex:t < nncal l'vent. and tinall) hac that the phy\iolob •l{al supc:nor irv nf bnJh Ot::W
w.1s so 1111portant, for rhcy respond ed d11Tercntl> tt cvl"nl. Cc

the ),J,t tnlobites dt~appc..trc.·J dunng thl· ~re.tt flllltlll'llt
Pemu,tn cat.htro pbr, dunng \'- htch over YO"n of the the lllJSS cxtinctt on p~:riod than did brachiop oJ
they were nor too ~criomly affet.tc:d by rhe PernnJ bal.lllll is
total 'hallo\\ -w.ltt't tn<lnlle in-. crtebrat c bim.t 11111< ut lo
bccam.: extinct cmts. Sub~cqurntly they rccoven:d :md untlerwe ntJ
lll:l.JOr radurion . whl"rca\ the br.1t h10pod' lo~t thro
nt.unt.un e
A t.t\t: .,d11d1 nu~ \t'em. p~:rhap,, mon: directllo dl~pl
related tu thl' effecr' ot tompl'lt tlon. w.Js the former dominan ce :Jlld remame d of low divef'l ll((\:1
tht<rc.lfter. Hl're, of courc;e, dinc·ct competl tion fi • tl ll' tllech

rcplarc.m ent ot br.H hmpod' Jt the end of the rcmarb b
P.uaenzm c b) btvah~.·, .1s th~. douun.m t manne 'c'- sea-tloo r \pace and food may h.l\' l' had a 'tunn .
1h<:~e lilt!
'ik filtt·r-fee dl!lg bcmho~. Brachiu pods dnmi1un :d cflcct on the br:~ch1opod btvalve bias, but the 1:1un.t
replacem ent W;t~ in the tir~t place \Ct in motion b\ MllfOI
the )c, cl-botto m ne: 1r.horc: .md shelf lubtr.tt' dunn~ pn tc
P:tla~.:nzmc ttnlt, but rht') wen: able: to share.• .tnum- m.uor physttal c.na~trophe.
A~ Ben ton ( 19H3) has ~hown , competi tion 011 1
SJ't: ll,\t101
hl"r ol lubir;m (espt•c.t.lll) 111 the ne.1rshorc region) popttl.lt lr
with btvJI\e\ .md .1 long-ter m t<qtuhbr ium wa!l wt own may mcrcasc the ltkeW1ood of extinctio n m
~hi Oil) ,
up. from rhe Mcsuzo ic ouward~. howeve r, rh1s hnc.,tge, hut it ts unlikely ro h.tve been tht' 01
c.tu<;c. Major ldtJ.\trophe~ or phy<;ic.tl change~. lu•wevc r
domiuau ce W:l\ Jo,t w th1. bi.. ah·e, .llld brat hiopod~ du:il pn
ht·came restncrl' d to rerum habJtJt\ alone Was tltts demonst ratt•d ~o oltcn tn the ti.'"tl record, han 1111J.
10 the pn
a dirt'L l comcqu enn· of t omp~tttton? Or wcr~: ally by themselv es been ~uffincnt.
\11 lllolf\11
phy.,ical tactorc; mvolwd hl"rL too' MaCHl
Therl' ts no doubt that bi,·dlve' ,\re much more cvnllltlu J
dfecnve ft.lrer fecd~:f\ tllJn Jre br.Khio pmb. The these." an·
pumpm~ ..tnd tittering 'Y~tl.. tn of rhc: btva1vc, h mm~ Ad.tptt\'C:
etlint·nt th~n the luphoph ore nf br.tduopml~. pltl lltltlli
thou~h both W\tcm' Jrl' compos ed oftilame uts ""ith
be;Hiug C1h:1 . HoWl'VtT, the.: org.llHz aoon of the EH)Iutio n can be ddin~.d a_<; a changt m gem~ 1ft. mgm.1l1
biv:1h·e feeding gill prodmc ' a rcl:ltlvcly stronger qucncics through t1mc. In any gene pool mmr van· tllg a brrl.'
J(lr p( ri
current, and the net cn~rh')' gatn for btnJvl's I'>
ation rc~ult, from recomb111:ttiou of dominJn t .1
Bibliography S1
nng (1)1\'e .1lldc\, but 11l''-" vJnJtlon ~~ mtroduced by only pm~ible where rhL•rc •~ v.tcant exmp.Kt'. They
:'iSl'~ unon and undl•r n~rt.\111 circum~tance~ mutant mually take place where a fom1crly ~table ccmy\tcm
lvc~ e on bcconH..' domtn.mt. In very small popula- ha~ collapsed, Jnd ""hen m a~' e1<-'tinct1om have
: in Oib much c\·olunon.uy change 1~ non-adaptive 'cleared the field' tor new kind~ of .mmuls and m'\\
,ttcr us.: of dw ctlel·c, of ~en~tll" dnft, but m larger ecosy\tcn~ to .m~e )uch ma\s exnnctlom, based
'Of', u!Juons sdecnon .1ctm~ upon thL· "an;ltion pre- upon ma..~or phy-.;ical changes, are probably the rnmt
the t 111 thl· gl·ne pool .md k.tding to .tdaptarion is the import.mt factor'i underlymg large-\C 1lc evolution-
)V er p:mury force m c\·oluuon.tl) dlJnge. Molecular Jty changes. and arc explored more fully in the next
t be inve nuy be .1 third tm·dum\lll of evolunonary chapter.
gc> but, likl· gcncnc dritt. ;, non-adaptive. Neo-Darwiman theory rem.um .1 viral bacJ...-
b: ~ew spcc1c~ nuy .tn~c by thl· 'd.tS\tL · pathway of gr~und tor the undcrsrJnding of ~ui.Jeonrolog~cal
tin· ,, connnuous dun~' 111 .1 l.u-gt· population. and evolution. tly respondmg Aex1bly to Jmtitit·d rriri-
th~ ere Jre some mdtr 1t1ons of this 111 the fossil record. nsm and as~1m1bting new dat<l and conccpto;, it ha'
vay, Equdly. Jnd pcrhap' rnnrc commonly. new species grown and developed wh1lc ~nil n:taming ito;
hail uVJnsr tbntptly Sonu.: m.ty originate through het- integnty as a theory. It io; bchevcd by ~omc
:tcr, w•hrony, l'~pcu.tlly 111 sm.tll populations. Other (Dawkins, 19H6) that the creative force or natural
:ity. 11'\1' 1pl·m·~ .m~c :1~ pcripht'Lil i_,obu:-~ on the fringes selection is adcqu.ttc to cxplai n all of th1..• diversity of
ring rhe 1wcnt popul.ttmn, tiii!,'T.1ting whnt>ver there is life and it~ evolution through time. Yet there are
)fll- n.ant ntcht• Onre l'\tabli~lwd over :1 w1de area. others who do not fed th:H this redurrionist
vcr .. pubuom tend to rt•m.nn M.ttil, bemg \tabilized by approach IS the only po~Slblc one, and who prefer eo
Lore hnon unks~ tht•rt " J m.JrJ...ctl envirmuucntal :>tudy organisms as mrcgr.1l wholcs . A new evolu-
·eo- ngt \\ lm·h mcn.'.l~t·~ tht• sekctinn pre~sure. Such tionary par::tdtgm may hl· emeqp.ng (Ho and
was \t2SlS u the nnm1 tl.1r popul.1tiom, .md the ongin of Saundc~. 1984). Tht·re h, ror
eX<llllple, SOil1l evi-
lvc~ 11(',\' lpn ll'' j, .1 r.tthcr spl'l i.tl and relanvcly rare dence. in ccrt.tin Lht'S for the mhentance of
{to nt Cmnpl'litilln hl't\\'l't'n 'l't't ies in a new envi- acquired charJctt•r. (Po!lard, 111 Ho and :)Jtmders,
:>d,: romnrnt nuy lw dm.· ct m the initi.U stage~ before a 19~4) and a pbmibk genl'tll med1amsm has been
uan l\t' h achll'n·d, but 111 J nmn: balanced conuuu- propo~ed. There i~ 'omc 'upport for a ~emantic the-
nta llll) oilongl·r ,t.mdmg .1 ,t,tblc pn:d.Jtor/prey rano is ory of evolution (Harbicn, I 9H5) 'uggt'sting a pJu-
fU.'lC 11l3111t.ltncd .md t·ompctitlo n is ll\ll,\11) avmded, char- rahry of evolunonary l.m., rc\emblmg grammanca.l
·sity :trrdbpbrcmtnt hl'tWl'Cn \llllil.1r types bcmg one of rules in (anb"l.lage. fhcre 1\ J llC\\ <!pprcCIJtlOil of
.
mg
for
. med t.mi~m~. ~ome t)'pt·s of biotic assooation are
-:m.trbhly pcN~tent nvcr Inn~ penods of time. and
genes having thetr own dynanuc ltfe .•md the possi-
bility th.lt genes can orgaruze thcm\elvcs mdepen-
dently of the env1ronment and tlut urgam~ms c:m
mal ~,. mdudc co-c\ olution.u: rdat10mhtp~.
)y .I \1t,mt volutwn rdi.:r' lt1 the ongms of new evolve \vithom any 'clcltmn prcs~ure (Campbell,
lp(nt' Tlm ,~,;rm to t.IJ...t• pl.lcc either by allopatric 1987). The new par.1digm nuy b.1lanre exogenous
1 tts fWJtion 111 accurd.Jnu• w1th the prediction~ of (nco-Daf\vinian) .1gainst enJogenous {intt:rnal)
tnJ pt1pubt11111 dyrumit tht·orit'~, or through hetero- cau$CS of evolutionary ch.u1ge. Perh.1ps, in :w or 30
mly hwny. fhc overlay or the seduncntary record, year., we rn.-ty be reaJing the rossll record in a very
, as htlWt'Vcr, obscures to a greater or lesser extent the diffcrem light.
ISU- nu.tl pmcc~scs ot ~pect.Hiort. Macroevolurion refers
to tht• pnKcs,cs ol t. volution .1t h1gher levels, ]e,tding
IOIIl.lJOT t<l'<OilllllllC 1.hst Ollllllllltles
MJln'C"olutwn.lry speues o;electlon lead~ to the
rl'olunonary trl.'nd~ ob~<.·rved tn rhe fmsil record;
Books, treatises and syrnpos ia
th~r are Je<.:<ntpkd from micml:'voluuonary trend~.
Ad.tpnw radiannn t\ .1 ,t.md.trd malToevolunonary Ayala, R.J. and V.tkn:me, J W (I'J7Y) l:t•tl/t·ml!· tlrt
pbrntHnenon refernng to thl· desct·nda.nts of an nrcoryllllll Jln>US50 "' Or!flllllt Ft•o/u/lt>rr 13t·J~J.IlUn.
fre- ongnul type (often wttll ·' key .1d.1ptation) undergo- Cunu1ung., Menlo 1'.1rk. Caht. (Vl'f) ~ood}
an- ~ a grc.-at bu"'t of l'\'t1lution and becommg adJpreu Barbicn. M. (ed.) ( Jl)g:;) '/lu '\rmiliiiiC 17Jtt>ry t!f I..It'tl/rttiOtl.
and \rc:otic lublt,lt'> .md nrche., \uch r.1dtations arc Han.vood London. IHH pp (Vl'~ de.tr prl·~cnt.Hion)
Bonnt:r, J 1 ( I9KH) '/71c h•t•/utiPII cif Ct>mpkury lty \leans '"' NcllurcJ/ HistcH)' Nonon, New Y or~: (V crv r~atLble Prl'ss. Chic.•go.
d/ Ncllllrcl/ Sdcwc•ll. Pnnt t'llltl UnJvt•r,ttV Prt:,~. N~w t"~>ays) biOIO!-.')')
Jt:r~t:y. Hall.m1, A. ( IY77) Paucms v( /·r•,•/ultCIII " U/mtrated hy 1 R.;~O-. R.A :md K.1
13rouwt:r, A ( IY7.1) I ;,li'lcll 11.JI,u·c•ut,,J,'SY· Ohv~r and fM.<il Rr(llra I )evdopmt•nrs m P~laeontology ~nd Et•o/uticlll - tilt' L
ll0>d· E,lmbtugh. StratJgraphy No. 3 El,c\ it•r. Amstcrd.un. Clla11!/L M :Krml
Campbdl. J.H . .md Sd11•pt, J.W (I'I'J-t) Cwlfi1•r Ho, M.-W ..mJ S.uuulcr., P,'l . (l'd') (19H4) &)'tlllci .\~ cext)
/;t•,,[,,..,;•,JtHlC\ .111.! lttrclctt, Bn,rnn, M.l''· (fi p.1pcr.) Dttnl'iui.<m. Ill flltr.•dll.-rlclll le• the ,\,,,,. f.c•c>lllflttlltlrf R.1dley. M. (19%)
<..ampbdl, K SW. and l>ty, 1\1.11. (t'd,) (19K7) Rcl/1'5 •if P.m1di~m Acadcnu,· Prn,, Nt'\\ YorJ.., .'\7(, pp Clear and up-r•
Et•,t/urr.•u AIJ,·n md Utmm, Ne" Y<•rk, Jl-t pp. (16 Hoffinan A (I IJ!W) ...lll,'lllllt'llh ,,, Ec•t>flllwu. Oxto:d Rw;c, M. (IYH2)
papt'r') Univer.ity Prt:~'· O!>.ford. £,•,>/rllillll Cc>IIIT•'
C)rta, (' S ( 1•1:;~) '"""''' h~•/cll"'" ,, ,\tudy <!f Rt·cmt Hmdey, J. ( IY42) l t•c•/uthlll tlu .\lodu11 -')'lllltt•sc• ·\lien (Excellent Jnd 1
1·,,,., ,,( "' C.rmn. Snl~\\1tl .md .J.t< k\t)ll, London. and U;1win, London (<. 'l,ls\ll lnt) \unpsou. G.G {
(CI.J"It tl' ,tbuok) Jepsen, G.L.. Mayr, t . .md Strnp~on, G.C •. (llJ~~ Columb1a Urm
Cherf.t\, J (l'd ) ( 1'IH2) I ),ln/'111 l 1t• '" D.111. l\rw Gmf/ic<, Palat'lllll<•ft•l,'y •lltd h•,•futHIII. l'nncetor. S1mpson, G.G. (
Sflfllli•IITI'C, I •llhlnn (2-1 P••P~ c' - lllll\t ll\t'hcl) University Prc"• Nt'\' Jer.t·y (M,m) Jndiv1du.cl papeo Columbia Unr
Copl', J W .111d Skdtnll, P R ( 1!JH'i) Ft•c•/urit•llllry Ctuc cla~~ic, if a lirrft, dated) combinmg m~r
Hisrc•rit·.~ F••111 thc f.mHI l~l'nnd, 'ipl'ti.ll l'.1per' m Kcr\haw, D.R. ( I'JHJ) <jlli/llcl/ /)wa~ity Unwm 11vm.m Stanley, S.M. (I'
l'.1heontoloh'Y No. U, Al'.ltk1111C' l're,s. London. London. (Very clc.u, t'Xt:dk•nr l1Ju,tr.lnom) Freeman, S.m I
(M.tn}' rt't l'llr d"~it P•'P' r\) Kitcht"r, D. ( 19H2) ·1/tulill,\1 Sdc'llc't': tlw Ct1Sl' A.~<IIIIS Skclton, P.W. ( I'
Co\\t'n, R, ( 111111) '17tr lfillclYJ' •'/ L!/1' 13l.it kwl'll, Oxford. Creatio11i5111. Open UnivtNlY Prt'~'· Milton Kt•ynt!l J>alacOtltdl.~~~ic·''
(limmt•ntly rt'.lcbblc) (Excelll'nt) Umvl'T\ity. (A1
D~m·tn, C ( IH.l\) "!1tc I '•')'•'1.''' ,,f the Ht'<llih Jnhn Murray. La pone. I. ( II;IH2) '/ hr /:,,Hi/ Rcnml .md El't>lrlll•>ll. Rt•tJcill's Sranky, S.M. (I
London. ./;VIII Scientific !11nfric.111 . Fret·m.lll. S.m fr.UJU\ttl. (I" FtlSSiis, GI'III'S
D.arwm. C. (I S:-~'1) < >u tlw I Jn~111 c•/ SJ't'fH' b)' \ lt..ms v.f cnllenl•d p.1per.) Nt:w Yorl... (S
,'\.mcrr/ ,<;,·ft c11<111' ,., rlu l'tr•rn•,l/1,>1/ ··f f<cll'tllllt'cl Rci(CS ,., I evmron, J. (1tJHH) C:c'llrtlc), / 1,1/ctc'cllllcl/t~tl}' <lltcf .\/.JtTJl. \tcickbergt•r, M .'
the '\t111~~/ /c•r /.1{• J•hn .'vlun.l\. I ondon '"'''"''"" C.m1hrrd~,. Umn·rm\ l'rc,~. C.1mbnd~c. (A. Hartlccr. Bo'r
Oawkms, R (I 'ISh) flu· Him cl 11 ./1,/cm.tko. Pl'IKan. cnmt maponJill 'Yillhc~") rreatmt:nt)
lim1111nd,\\onh. (N.ttllrll ~cluiHIIl c\ l'rL'.lriVl' force) '\.byn.1rd ~nmh, J. (1'175) Tl11 /hrcll')' f!{ / •c•t•futr.•ll. 3r V.ucntme. J.W
Dot>zhJn'l:' f, A~,1L1, t J, ~tchhm,, C l. .mJ cdn. Pc.:ngum, ll.amlt•ml,wonh (lnv1huble) \lmiur BIMJll
\'.llennnt·, _1.\V. (1'177) h·,>lullt•n. frl'l'lliJn, San Maynart.l ~lllith, J. (nl.) (I'IH2) E'Irv/111w11 '\,•u •1 Cn11m7 Ne\\ Jcr.cy. (~
fnnct en (( umpn:hcn "c .tdv.ann·d text) <!fra Dctnt'lll. M~emill.ln, London (MJn'r kev papcn Vennqj. G J. (I'
l:ldredgt•, N (I '1'1 I) I •'SH!.s, Ill• ft•,•/utlc•ll md h\timl••'" of \\1th mtroducnon) Hc>r.•ry ~( Lifr.
Spwr.s. Aunun, Nt'\\ York (CI~.uly \\Tltt~n. 'uperb Maynard Snuth. J..ultl S1.1thrn.lr). F (I <J<JS). 771£ "''''"'
olllur pholli!,'Taph~) cramctl<>ll> ill n•••/u/11'11 Frn'lll.ln, Nt'\\ York (Gcn.:IJ
[ ldrcd~oc. N .lrld St.lllll'\, s 1\\ ! I Cll<4) /.11'11(1? h•ss•ls. approach)
Spnngcr \'crl.1~. Nt'\\ Yllt~ (J-1 p.tpcn~) Mayr, E (I Yf,J) -lnillht! '\f't .,,., t111d li•·••lrilit'" Individual
Endler. J.A. ( 111Kh) Xcllllltl! .\dc~fl.•ll Ill '"' rl'dd. Prinu:ton Un.ivc~ity Pre\,, C.unbml~e. M,l\\.
UntvtNt\ Prt'"• Nl'\~ !l'r-~:~, (M,IJtlr rt'Yll'\\, hut tcch- Mayr, C. (I <J7f>) f:po/lllr.lllllllllthr 011'mrty ~~ U)t'- .\dtclt lknton, M.J. (1
mcalh· dllhrulr) bscl)'·' Udkrup Prc~\. C.unhlldgt•. M.1ss. wry ofbt'i:. l\
Furuym.1 , D.J (lll11t.) lc•cl/lltiml•ll)' flic•h•l,')'. 2nd edn. MJyr, E. (cd.) (I 97H) Fl'c~lmioll R£'ad111g5 .fmm Snwr!fJ< Uenton, M.J. (I
)m.lller. Sunt.lt rl.1nd M.1" Amt:n.-an. Freeman, S.1n Fr.lllCI\Co. (Nme .article..) Mvlogy: 11 S
Furuyrm, I>.J. .md "il,nktn, M ( I•IH4) Coc'l'cl/lltidll. M cKmney, M .l. (t:d.) (I YHH) lil'll'r<lclmmy i11 J;volultmr. Crowrhcr), Ll
Srnaut•r, Sundt•rl.md, M.1" (V.tlu.1blc text) AlultidisdplintJry llppmarh Plenum, N~:w Yotk. pp (,;lmpbcii,J.H.
Garnlm. L. .md VIm''· c; ( l 9H7) T1tc E1•cl/11111111 •!f Lift·. 1-348. (l6 paper.) Rates '?/ F11olc
Colhns, I ondon (h1w tn'.ltllll'nt, colour rllustr.1t1om) McKmrwy, M . l nnd McN.1nw·a, K.J. (1991 M.ll. Day), I
(;et", 11. (I '1'1(,) Hr:fc•lr thc• ll.ukbclllt t ic·u•, ell I tlu Or(~till ~f J-lrtrmclmmy - tlu· Fl'tllllticlll ll( ( )lltcl,l/fiiY Plenum. Ne\1 ( 'Jarkson, E.N.
the Vutrbr,Jin ChJprH.IIl &. 11.111. London York . pp. 1-137. ( fhe llltl\t mcful 'Ynlhest~ .iv.UI.ible brate ~pecics
Could, SJ. ( 11177) Clutc•t:<ll)' tfllcl / 1/iy/rll!fiiY BdLup/ McN.1mJJ"J, K.J. (cd.) (I 1)'10a) ct'••/uti.•llcll')' lit~~<l trJnsfom1atir
lhrv~nl, ( lmhndgt·, M,l\\. (I•"' <'llrm lmtonc:.1l .llld Bdhaven, Cambncl~e, Mm., pp. I-16K. (14 papel'l. 99, ISJ-71
phalosnphrc 11 tll'JliiH'Ill of tdt'l\ nu rcc.1p1tul.mon, di~CU\\1011 Jnd lJS~ lmttlrll'\) Dott, R.l I. (I'
nt'lltt'll) and p lt·doul<~fl•ltt"") 'VtlN:unaro~, K.J. (t·J.) (I'J'J5) /;1'•>/llllc'llcll)' C:/ccll~l/C 11nJ ,,m. JtlltmcJI
C:0uld. SJ. (I 1J7l:\) /r•o 1 'ill tot /I,Jnt'lll - Rrtftmt•ll~ cm Hcrcrc>dm>n)'· W1lcy Ne\\ Vork 11 \,llmblt• pJpt:T'i) .1ddr~")
,\,,wmd J-l,,,.,,y. I kut'< h I tu d,m (Vt'l') n·ad.ahlc) lt:Jtr. R.. A. (IY'J6) nu "''''J'I' .., 1..1/t' Gwu, /)c'J't'"'PIII<'III, IJtwcr. G (Jfl:
Gould. \ J (I'JXll) 1111· f',md,,'s 11wml>- Mc•rt• Rt'tfrrtic>IU .wd rlct' El'vlutwll <'{ lmm.t/ F.•mt UurvcNty ofCluCdi!O ~pecit"~ cvoh
Bibliography 53
Pro, ChK·:agc>. (lmls dc\'clopmcnr.tl .md cvoluuonary Drake. J W .. (,Jrckm.m, B.W. tnd Rtpln·, I (14X3)
br•>ln!-')) Updating the tlrelll)' ofmut.ttion. AmWI•III ,\,imtllt 71.
Ra[ R.A md KJufimn. T.C. ( lliKJ) Eml>ryc•>, Ct·11c> mrd 6::!1-311.
En•luth•ll -lltc' Dn•rlc•['mt 111•11 Cclll'tir B.bis t') l :'r•t1lutiotwry Eldredgc. N (I'J71) ' 1 he illopamc modl'l .md phyl1>gcny
c.lwr~c . M tcm rll.l/1, Nc:w York. (Adv.tnn·d llltt'gr:mve in P.tl,u.>n7oar urvcrtcbratcs. Et•c>lutwll 25 1511-Ft7
l~\i} (Very tmport.llll work)
lthll~y. M ( 11)\)(l) f:po/utwu. 2nd c:dn. 1313< J.. wdl, Oxford. Eldrcd~c. N. (I'J7·b) Clur.tcrcr daspl.tccnH'nt 111 lvnlu-
IU~.H JIIJ UIHO-tl.ttc:) tinn.rry hntt'. lmrrir1111 Zcwlt~({i.(/ 14 , lmr\-47
Ru,~. M. (1''~2) D11nl'i11ism D~ti·~tdcd -- ,, C:llldt to rltr (Advanced)
h,,/urt<'ll C.lllll<ll'fr~ii'J Adru~on We,lt')', Rc:.rdin~. M.tss. Eldred~c. N. ( 1117·1b) St.rbilitv. d.iVl'rmy Jnd 'PCl't.tnnn 111
bct-Ucnt Jntl rl'.td.tblc:) P..tlreo70IC epo:mc ~c.r~. jollm.r/ ••f fJ.rlrt>lllt>lc'I!Y 48,
\impwn, C G (1')44) ltlllf't' ,wtf .\fc>clt• 111 b·dlllhlll. 54<~ (lmpon.mt \\l•rk \trt's\111~ .illop.llnl' c:vnlunon)
Columbta lJfli\'C:"IIY Prc:~~. Nc:s\ York Eldrc:dgo:, N tml Gould. S.J. (1971) Punctuatl·d O:ljUthb-
::npson, G G. (14='>3) 77u· \/,,,,, r,·,rtwn ,,f £,.,,futwll. na an ,Jltcnuuvc to phvkuc grddu.tJi,m, m \lc1dds 111
Columbt~ Umvl'Nty Prc:,_, i'.l'\\ York. (( l.t\\IC work P.rwro/rit•lt>~)' (<:d. T.J.M . Schopl). Frcednun, c,,.,pt•r,
ct•mbuung m.ury .hpccts of cvolutwn tht'<'ry) !>.m fr,lf1li\CO, pp. H:!-115 (Allopauit cvolutwn)
~nlev, -;,M ( 197':1) \ lmTciCI'c>lmitJII , !J,I/Ic'lll .md P1Nt'SS. Endlt:r,J A .md MtLdlan. T. (19HX) The pnKc'' ofcvo-
hwu.111, S.m Frannsco. (Ain:,tdy ,1 d.ts,tc) lutton tQW,Jrd' a newer 'ynthe~ts. .·~u""''' /~c·t>icw ~f
~~dlt>n, !' W. (19'13). l:l'aluticm. ,-1 C.•milim•,/ Hioh,l{i.-alllnrl Emlcwy .wd Sy(trmmits 19, 395-421. (!::.Mdkrn sum-
P,lf,JtWIIcl/t~~··"l .-tppn>11rh Addt~on We,lev/Open m,ll) of evuJulillllary proce~st's)
LniiCf\l ty (>\tlnm~hlc fusron or .!ppro.tc.:hl'\) Fei~t. R. and Cl.trko.on. E.N.K. (19H9) Envrronnr~tlt.lll\'
Sunk}. S.M (1'1!11) 11tc .'\'rw Et•c•/lllrc>u.try ' fllllt'tahlt• - controlled ph> ktlt rvolutton blind ne" .md t'XtllldiOII
fDSStls, C..·r ,., ,,tf thr ( )ru~m of Srr,·!r• B.t\ll' Hooks. m IJtt' Dt"vnrH.ln trop•docol)-phinc trilubill''· L:t/tm,,
~r\1 Yo l ">unple hm wry gm,d) 22. 359 37-1.
Stnckl-oergt•r f.\ W (I ~'lb) Er c•lutwu, 2nd .:t.ln. JnliC\ and Fortcy. R.A (I 'JXS) Gradu.Ut\m and punctuated cqUthb-
llmlcu, Ut"mn , M."h\. (E.,n.:lknt, nltnprchcn~rve na J\ conrpl"trn~ and complemental) theonc\, m
trrltmenr) b•c>/11111>11<1ry C.r<c Hi.ilc>rics from tilt' h•~lil R.t•cor.l (cd,
\ cntm~. J W (1973) r,.,,fllltclttal)' P.tl•ll'<llll<>lc~I/Y •!f tire J.W. Cope .rnd P.R. Skelton), Spcct.u P:rpc,..., 111
1/.tnttr BtchJl/lrrr Pn:ncicc-H.lll, f:.n~kwood Cltffs, Palaeontology No. 33, Academic Press, I ondnn, pp
~~"' jl'N'Y (~cc Chaprer I) 17-2H.
'il'ntli'IJ, (:.] ( llJH7) J!po/utioll mtd E.mrlatiow ,,, Elclh~~ical Gould, S..J. ( I%()) An ('volutiott.lry mirrtKOM11
1/rsl.•ryof l.t{t•. Prinu.:ton Uruversrty Pn.'\\, New Jersey. PlcJ\toccnc and Recent lmwry of the l.md .~n.ul P.
(l'c•ecili::ollltt's) 111 Ucnnucb. Bullrtim •!I t/u• l\ltN'IIIII
of C'""Jl•!rollil'r z.wi"::!Y 138, 407-532. (Allop.ttnc
e\·olution)
Individual papers and other references Could, S.J .md Glllow.ry. C.B (I'JHO) Bro,:alve~ .llld
clams· slup' that pa~> m che night. p,,/wl•wl.~r 6 ,
too, \\j 19XJ) L.uge-~cak repl.lcenll:ms m the his- 3H3 ·407 (Why did bivalves cake over?)
roryc•l hie ,\,rlrtr< 302, 16-17 (Compl·tlnon ''not all) H.tllam, A. (I <JH2) Patterns of spcciJtJon 111 Jur.t\SK
Ilvn, M.J. (I 'I')!)) Red Queen Hypothe,l\, m PllltlPcJ- C.ryplmra. f><~lrc•lrialclf!Y H, J3~n.
~ /,•~)·: a .;;yllfhc.Hs (cd" D.l:: (; Unw .md P.R. Hunkaptller, r .. ll umg, H ., Hood, L and l mtpbell,
Crnwthcr), 131~ckwcll, Oxford, pp. 11 Y 2.3. J.H. (I IJ!l2) The unpact of modl'rn gcnctin on ~·volu
npbdi,J 11 (19H7) The m:w gem: ;Uld its cvulurion, m rionary theory, in Pmpectit,es in I::volmic>n (l'd R
R.IIc'l t>( Ea•cl/tJiitJtl, (cd~ K.S.W Campbcll, KS.W. and MJikm.m). Sm.mer, Sunderland, Mass.. pp I (,4--H')
MH. Dlv) Alien and Unsvm, New York, pp 2Hl-109. J1rob, F (1 '177) Evoluuon and onkcnng liw·11a' 196.
kton, f•.N K ( IIJR8) The on~n of m.maw invcrte- 1161-7.
mtc \pi:Clt'S .1 lrlOC:ll review of 0\ll rocvolunon:try Johmon, J c; (I'JH2) Ot'tUrrence ofphylettc gradu.th~m
tt:ln\filllliJ!IOn' PrCil<'< dit!i/S •?I tilt Gt\1/c•gr•IJ' .dsJcVIcliiOil .1nd punctuJll'd t·quilabna through [llm:. jcumltt! of
99. h3-71 Palcc•rrt"IO.t:Y 56, U11J-Jt. (Scdm1cnt.try nwrlay
R tl (I 'IH2) The cballeugl' of 'ocnutic c rl•anon- ob,curl'\ cv.:m')
u J.•um.rl 4 P.J/t'<>lltc>lo,s:y 56, ::!67-71l (Prc"dcnnal Kimur.1, M ( 1979) fhc neutral rheory of moll'cul.rr t'Vo-
dJn..,,) lulion. ScmllifJt Allrt'nrall 241, 94--104.
Dvm, G (l'IH1) Molt:cul.tr dnvc: .1 lllhe~tvc modd of K.itts, D.B. (1 '>74) P.1lcontology and cvolunon,rry them>
fli:CieHvuluunn . Mlllm•, L>11d011 299, 1I I 17, Er,o/11/ion 28. 45X 72.
Evolution and the fossil record
Levmton, J.S. (I C)tlJ} St.hl' an pro~-trt:\\ . the empmal ba~as and P.R. Crowrher}, Blackwdl, o,furd. pp 106- IO.
of nucroe\ olunon 111111~<11 Rn·rw• 4 Ec.•ltwy mrd Shddon, P.R. (llJ90b) '-lhakm~ up noluuon.1ry pattems
S)'lll'lll<llio 14, 103 17. .\!,uurc 345 , 772.
MJ\T, E. (1 1h4} Ch.mgt• Ill gt'nl' tiC cnvarnmm.·nt and Shddon, P . R. (19<)(,) Ph" lJ dunge: J model for 't..J/lltr<'ll .a;,, l'r.~o> (nl\J . lluxlcy, A.C. and evolution m dafferent en\'lronmcnl\ Palur(l.
Hardy and fLU l·ord), Alien .md Unwm, London. pp. ge<~l!rapll)', Palmwlm•·Jtolol!)'• Pttlanlt'Ctllt')!) 127, 209-27
157-811. (Allop.1mc 'J'l'( a.mnn) Samth, A. and Paul. C R C ( llJXS) V ana non 111 thl irreh'll·
McNamara, K.J. ( I'JXl) Ht·taoduony .mJ phylogenetic lac echinoad DL<ttlide.t during du: early Cl"nonuruan. u
trend,. PtJ/,,>/rr,,/,>:~:y 8. I \11-42. (Pacdomorphom ,md Et'Oiutwunry Case J-/wanl'.< fiam tilt F11.<>1l Rr(llrd (ed' J. \\
p<'ramorphom) Cope and P.R. Skelton), Spcn.1l Papcl'\ in Palaeomolo~
Mcf'amara, K .J. (J<J'Iob) 1 ktcrochroaw. tn P.rlnrol,olo'(y: 33. 29-37.
a Sy11tlrw< (cd~ D .E.G. On~' .mJ P R Crowthcr). Steams, S.C. ( 1977) 1 he cvolutaon of hfc-lu~rory tram
Blackwcll, Oxford, pp. III - I H. Am111al Rwlfll' tf ~:;c.,/,\1/Y aud Systwwttc.s 8, 148- 7l. (r f-----~-
Pollard. J.W. ( 19H4) I\ Wt'l\lll:lnn \ b.amer .lb\olute?, in aud K-~elccuon too si.tnpli~tic)
BcyiJnd 1\r,,·OcJn•~ll~>lll. (l·d~ M-W. llo and P.T. Stecle-Petrovic, M. (1'>79) The phy,aologit.ll ditfercncc'
Saunders), Acadeum Press, New York, pp. :!91-314. betwt:en artaculatc br.1cluopods and filter-feedmg ife began on E;Jrt
Schindel, D.C. (1 lJHO) Macrosrr.atagraphic ~.mapling and bivalvcs as a &nor in the cvoluuon of m,mne lt:vcl b.ot· ~thin the first 30%
the linut\ of p.lleontolob'll' reso lu non. P11lacobiolo,'()' 6, tom communn:ies. Pa/ncollwlo,l!)' 22, 101-34. (I' cum· .me the E th ' ()
.. . ') ar s a
408-26. (Sec text) petJtJ.on amport~nt. ·d 0 f fr
Sclundcl, D E. (I <.>H2) Rl'\Oiutwn analysis a new Van Valcn, LM. (ltJ7'i) A new evolutionary law VOI cc oxygc
approach m the ~·'P' 111 thl" ti)\\ll record. Palcohiolo.~tr 8.
E~·oltttiOIIllry Theory 1. 1- .30. (Red Queen hypothesi,) arter for_debatc.:, as
340-53. (~cc text) Williamson, P.G. (l':.IHl) Palacolltologacal documentation Jve c~nsJSted of C l
Shddun. P.R (l<JX7) Par.11ld ~radu.ili~tK evolutJon of of spcciation rn Cenozoic nwllusc' from the Turkan;, rcn1a1Jvc modds Sl
OrdOVICIJII trilobit('«. \11111111' 330, 561 n3. Basin. Nnwrr 293, 437-tJ. various other mi
Shddon, PR (198H) M.1kmg rill" mo~t of the evolution Wright, S. (1<J32) The rub of mutation. mbret:dm!, own that many ki1
dJanes. St'l , &wuist 2 1, Jo~nuJI)' . 52-54 cross-breeding and ~clccmm in evoluuon PrMecduw • '>Wd have been' syn
Sheldon, P.R (IIJ90a) Mlcrocvoluuon and the fos~il rite Sistlt lmmtnltiJIIill C.mt(ri'SS ,,( Cnrctics. 356-61 esc gaseous nuxn
record, m Pal.utlbialt>!!Y a 'irutlrt·~rs. (cd~ D.E.G. Briggs (Adaptive lanchcapc~) ~ergy source wa\ pr
'articular amino acu
cnrally. as have at .
.rbohvdrares. aJI vir
. Some sunple o r:
ctroscopicall} Jet
mplex ones have ~
The energy requ
olccules was c.:onti
ctncal discharges,
i!liations, hot spnn
·at. After several hL
the Earth was eo
ocean basim, co1
~
anic material (a
ldanc). The next
clear, since the a
'lught together a
1mtually to rcplu
cnc} of DNA. At 1
eit in a prirruove
lution, for the eo
~ecn the living n
nt, and allows an
~ O\\m.
lS
Major events in the
1-
n
l.
history of life
;,·
"·,_ Some of the earlv protem-hkc sub~tances proba-

bly polymcnzed 111 sphcncal boches, mch as have
been produced expenmemaUy tn comhuons simu-
tg life bcg.1n on E.uth over ]:;oo M.1 ago; th;tt is. lating those of the early Earth. These may be
t- 1thm the fir~t 3() 0/r, of the pl.tnl·t\ history. At this
regarded as cell prccu~ors. From these, by proet:~ses
1-
tutll' the EJrth\ .ltmmphl•re was rcdming <md still poorly understood, came the e.trliest true organ-
dvoid offrce oxyg~..·n. Jmt how reducing it was is a lsms. Smcl' the present st.tte of knowlcd~c of the
m.mcrfordebare, a\ i' it, ,\ctual compoMtion. lt may origins of We and th~.· proccssl's ot chcnucal cvolu-
)11 !t1\: 1·on'i~ted of CH~, Ni l,. 1-1 20 .md H ~. though aon have been sununanzl·d dsewherc (e.g.
n.l cmatii'C modeh ~uggc~t N , md CO,. wHh H ,O. Awram.ik, 1YH2; Wocse and Wachtl·rshauscr. 1990).
\'mUll' othl·r mtxturl'\. F11.pcnmcnt1l work -has these will not be followed further hert·, nor will a
111n thlt many 1-..md' of \lrnpk or~anic molecule, very dc[J}.kd p1cturl' be ~1ven of the later sr.1ges of
~·
of uld hm.• been s~nthe,lzl'd .1h10tically !Tom any of pre-b10ttc and early bwnc evolution which Cloud
,(, , thtsc g~eou' mtxtun:,, prO\ ided th:n a ~unable (1976), Schopf (19H3, 1992a.b), L1pps and ~Hgnor
nttg) ourcc \\"a\ pn·scnt to p ')Wer the rcacttom.. In( 1992) and others have treated.
mculu :~milll) acids have bc:en produced experi-
~ . a, have at leaq ..onw nucleotide basco; and
arlloh)'drltC\, all \'lt<tl tO the ongtm of hving mate-
Some \tmple organic molecule' ha\'t: abo been
~troscoptc.tll} dctll tl'J Ill inte!"itdlar -;pace and The earhest hvmg orgam~m<> were undoubtedly
mpll'X ones haw bt:l'll f~1L1nJ in mctc:onte\. prokaryotes: cells devmd of nucle1 and with the1r
Ihc ~na~>y requ1rcd f(Jr the '} nrhe~1s of such DNA not arranged 111 chromo~omc~. fhere are two
mohult... \~J' ~.:ontmually J\.lll.1bk m the fonn of main groups, th~.· Archaebactcna (h,1lophlles.
U'llal disdl.lr!tcs, ulu.tvlolc:t light and wnizing mcthanogens :md rhcrmophiks, abk to hvc 111 sm-
lutllllh, hm 'Prlllg'i. J.wa lllltpourm~ and solar gularly hmnlc anox1c cond1t1ons) :111d Eubacteria
w. Arr(r Sl'\Ual hundred nul lion years the.: surface (which include. amonbtf\t otht•r grm1p~. purple and
tth1· E.lrth was lOVl'rt:d by an OlC:ln. or a series green bactem, and the hving .tnd ros~tl cy.mobacte-
I tll(Jil ha~m,, ront.llllln~ a 'hot tblute ~oup' of ria, colloqui:llly if incorrectly known a~ 'blue-brrecn
',;liiiC material (a d,l\\Jl phraw due to J.ll.S. algae' (though 1t 1s kgttlm,tte to call them 'blue-
HJme). The next st.1gc~ 111 or~alllc evolution Me greens'). These Eub.1crcria were the tirst tmc photo-
ck~1. >irH.e the .UiliiiO Jc td molen1lcs had to be synrhesizers. Probryonc tommtmitH.'S c:.~n be traced
ought togt:dtcr <md polvmenzed a~ proteins, bKk to about 3500 Ma, and dommatc:d the scene
IIUllh to replicate themselves through the for a long nme tht•re.1ftcr
nq ot DNA. At thl' p1)11lt the on~in at the cell, By about 2500 Ma then· e"sted tel'ming com-
!Mt m a primmw lonn, "·•' e''enual for further munities of b.1cr~.·n.1. mcludinp; C}anobacrcria: thl'\e
uuon, for the cell membrane .1ch a' a butTer werl' of many ktmh and mc:tabohzt•d in different
\'tCII tht' ilvmg matenal and the outer em·iron- way~. Moreover, they Wl'rc <1blc to exchange
11. md allow' an l'ndosed incern.1l environment genenc rnaren.1l in rlw w.1y that thl'lr modem l'OUil-
0\\11. terp.lrts do Indeed 'the nnly lll.lJOr fonn of gcm·nc
56 Major events in the history of life
cxchJngt.· Yl't to he cvoln·d wa' the complic:ttcd e<1n h~: tested ag.1111st the m h to~\il n:cord of tlw !Jt. :t'i.SOlt~ted
kind invc.:lllt'd b) l'llklryotc~· (Dyer .wd Obar, Prccambrian. \ud1 to"1h ou.:ur in tlnTt: kinds < d•XC\SOT:S,
II.JlJ4). environment (Scrodu:r, I ')H'J) (2) Str•
All hvmg nq~.ll11\lll\ , otha th 111 prok.1ryotcs Jnd ( I) Chnt-tarbon;Hc l.1ue' Prokaryotic life-foro OIICll tour
vinht''• .lrt' euk•ryotlt . Eukaryote cell' :uc apprc- were hrst t{mnJ 111 1954 h} studymg tlun wctiOU\ IJ miiL'l cc cl
nably hrgcr tlun t),,,,l." ol pmk.1rymc' and of 3 oi the 2000 Ma old Cuntlint d11.~rt from eh!, llnlly tlm
much higher gralk ,,, org.tnl:t.ltlOII (Fig. 2.1 ). Each Prcc3mbn,m Iron Fonn.ttJOll of southern Ontan ~c· rt•tmg
cell h.1s a llll.'lllbr.tm·-buuml uudeus, within wluch Tlm \\oork n:vc.JicJ a nrh Jnd dive!')e a'sembbge fnnncd n
n•,idc the chromommc,, and the t vtopl.t~m contain~ prokaryote,, prt'Hinldbl} photl)svnthetic and p Ot.ltO)I[('~·
variom ·org.mdln · 'peuJlucd tot ~uudry functions, ~ervt'd as thrct•-dimcmJOnJI mould.s tn the chert; They m.l'
including ,e,ull rqmlULJdiOil. The mitochondna, Some of the~L' art· ven. \Ulu!Jr to living C\'o:lnobact(· ht•mi,phc
tor ill\t lllCI.", .trc th;;o 'ttt·s ol rdl re~pir.1t10n, and they ria. Further work r~:ve,1kd thJt prokaryote~ :~r be tracL'd
h.tn• their U\\ 11 I >NA, distulct frnm that of the conunon in Prt'lambn,ut cheit-carbon,lte honzom ('J7h) Wt
chromosomn. !:lo dtl th~: dlloroplas~ m the celb of The oldc>t 1-..m)Wtt mu:rotossils in chcrt come fro1: badl} .ttfc
grl.·cn pl.mts; thl.''l:' arc ~tllk.inglv silnil.u to certain the Wanawoona Group ot Western Austr.tha ~how c.1rly Ca1
kinds of tn·e-hv111g t y.uwb.n tetiJ. The cilia and Aa- ing filamencous and cocco1d stmcturcs 3sson~tn coiUJnnar
geUa (rollel t1vely undullpmlt.t) h.we .1 consc.tnt wich probable strom.ttolltcs m chcrts some 3400 M uluJt· wa1
\tntCtlJre, JdemH.al 111 nlll:'uk.tryottL t·dls. They have old, and .111 ,JCLi ve ccmystem must have been prescn ( ') Sha
a ccntrJI p.ur ot tontl.lttik nmrotlbrils. surrounded then. From .1bma 20()() Ma ago onwards. mh stl~occur
by 3 ring o( IHnt orhl'r p.1ir' !:lmh stnkmg similari- microbiot.ts are oftt·n found in a cypical chert- ncritarcb
ties, J!on~ with tllllth other t'vJdence, "-trongly sup- carbon.lle t:1nes. Very nch .t,~cmhl.lgcs occur m th t'ukaryott
parr rill' theory of ~:ndn'~ mbtmts, whit h postulate~ later Proterozo1c (e.g. the HSO Ma Btttn Spnn~ lw Timo
that dw organdk' w~·rt nnce tndepcndcnt, ITec- Fom1ation in centr.1l AmtrJh.l). These arc donu· tound Js
livmg b.1ctcn.1. Thq bcc.Jmt· mcorpontcd sun:cs- nated by prok.1ryote5, and m gcnaa] tenns 'h mmt oft
SIVdy in orhcr t•uk.u·ynuc cdl<.. Jnd lmt their comp.lrJtivcly httlc hght upon the time of origtn ,:an be rr
amonomv. I ht• euk.lrvotic l ell ts rheretore a probryotcs. One particular coccoid gcn~ time dive
'community ol ntll rn-org.tnl'nt''· workmg together Glclltlborryditlll. ha~ arrractcd much attention m thl: ~go ..md
as a ' marri.1gc of conventencc' (M.1r~,;uli<> and rcspecr, for Wlthlll each cell therL' 1s a dark spc .md uf m
McMcnamin, I'NII). hwn the acquisition of originJily mterpn·tcd ,h J nudcu' (Schopf. 196ti Jll l'OJst.ll
mot1hty. and the compln '}''tt·m of mitotic and Tftlm were ,o, Glnwblllrydu111 would bl:' J cukaryot !vb lll', s
mcwn n:ll dJVNOn proh.1hly lWSt' by the inva,ion Although the cell w.11l" ulrr.J,trut.turally compattblt 1.: xtmct. a
of sptrnth.lCtt' b.ll'tt•n.l, wlud1 h.JVL' nm\o undergone.: with a eu~1ryotc .tllimty, other evidtncc 11\a} mJ. PrL'c.unbt
such mothtir.HHltl that only tht•Jr microtubule' cate that tht\ org.umm 1\ ,lctu.tlly .t prok.lryotc TIK \ \1hen
rem3m 'nudc1' may be nn mort• than dq,,rrad.ttion prodmt d1 vc• '1tic
The ti~c cndosymhtllnts \H'tl' .tnn:\tral bacrcria of ccUular cyropla.,m. For Js Knoll and B..trghoon: divet,Jty
whirh llll orpm.ltnl utlll.·r b.tl'ten,t. were .1blc to ( 11)74) showl.'d, thL' t.ell contents in bvmg cyanobac mt~1 rht: 1
rcspitl' aerohlt.llly, .md wlm h \Ub~t·qucntly became tcria cm contract to the centre of the cell afit 1\ COil•
mitochondna. At tlm tlllH.' fil.·c oxygen was an dcJth, giving .1 spurious unprcssion of bcmg • Pt oterozc
active pmmn I~H mo~t b.tctena, and not 01lly did the nucleus. Then· arc some 700 known examples n s~·~lliCncc
~ymbiotic union nt mnothondna unp.trt an effective later Pn:c,mtbnan ~.hcrt carbonate Jsscmblages, ;m I ll rt. the
way of gctttng nd of lt, but tt .tho gave them a new Jltbough po,siblc cuk.tryote~ h,tvc been reported t" I \ITO!ll;l
way of .1c.quiring cm·r~. The butld-up of fret• oxy- other diver\c Jnd wcll-prc~crvcd .ts~emblJge~ je.g pruk.Jryo
gul in the.• otc.•am from photmynthesis may thus the 1.:2-1.4 Ma Ucck. Spnng Dolomite 111 Calilorm pl.•nktit
haVl' been the rnggc.·• !o1 thL' orig~u of cukaryotcs, (Lican, !97H) 1. lt ha' recently ben> me cle-ar dlll hvmg in
the tmung of wllllh tJil be linked to thi' event. It JS chert-t:arbonJtc fjuc~ an: not the nght place to lam tlll'llb . 'T
probable th,H the nudt•.tr nH:mbranc cante u1to tor eukar}otcs. Thq .1rc. l'ljUtvalcnt, m~tead, to tht <.,it 11olwtr
bemg at .1bout the ,,mll." tune. and plastids were nucrobtal mat' of todJ}, which arc almo~r cxclu- blagc uf
acqum·d m turn. lt •~ wry hk.cl~ tllJt all thi' took sivdy prok.1ryotk 1 hl''t' liw 111 narrO\\ I} ddin pCI.I!--rlC
p!Jcc radll."r quu.:kly (Dyc.·r mJ ObJr, 199-l), ccolog~cal condittom whtl h .trc oftt>n har~b enou wrncd u
bet\H't'll about 2.H .111d 2. 1 Ga. an h} pothl'sJs which to e.\.dmlc cukaryotK orgatmnh ,tltogl."ther. ott While d1
Earliest metazoans 57
r !!Omted. tor cx.lmpk·. likt· thc1r Prcc,lmbrian pre- of late Riphcan life, thc~c microbiotas 'how ~omc
f dcr~mn. \\1th evapontc\. tlling of the diversity and evolu tionary complcx,ty
2 .Strom.Hobtt•s. Prc~ambnan prokaryote~ arc of both prokaryoces and cukaryotes JUSt before the
:rm iound a~'Ol"IJted \\ 1th ~tromatohte\. These are rise of the fim mcta:wam All t•ubryotc~ reproduce
unJt~:d 'trurrurt'' bllllt up layer bv layer. nor- scx..uaUy, and bear the potenual tor raptd evolution,
\ through tht• ~t·titment-bmdmg and tarbonare- since thetr pmenrial for vanauon, upon whtch nat-
l't'nng :KtlvttiC\ oi thl' t yanobattcn.1 wh1ch ur,Jl ~election can ate, I\ muth h igher than in
nn~d m1crobi.1l m,lt, un rht· 'urt:1n: (\omc 'stro- prokal)'Otes. Y cc tlm porcnual doe\ not seem eo
lltltobtc,· m:ty ,Jctually be rhcnucaJ prcnpttatcs). have been explotted tmmtdtatcly. The oldest
Tht'. m.ty be li.lt, bur .m: ut(cn cuJumn,lr or tonn known eubryotc, Grypmua, IS a cot.led unbranched
hcrn~~phcrical mounds flw ~tromatoLtc record may filament up to 30 mm long, and it comes from rocks
e be rnrcd bJck to JOt HI M:~ 'itromatoutc-. (W.llter. over 2.1 Ga o ld. There arc likew1~c spherical micro-
q~b) were tht• tlr'>t tnlt' rcd-fonnel"i, but were fossils from China, somev.:hat younger, and the
bld!) .ttfcrtcd b., the ml uf gr.tzmg org:umms m the earliest eukaryocic cells k nown tu belong to any
1rh Cambn,\11 Thc lK'\1 tnotkrn l'Xamplc~ of modem taxon are red algae, around I Ga old.
11lunma1 ~tromamlitc.; .tn• tound IJVing 111 hypcr- The Late Precambnan ocean can bc understood
:1 hnl· \\Jtl'r\ iu Sh.1rk D.1y. Wc~tcm Austr,1lia. as harb o uring a dynamic and vital ecmystcm of
1t t3) Sluk-.....iJt,tonc folcit·s. Prcr,unbrian m•crofos- autotrop hs (organisms that m:1kc their own food)
h ·.1• r~rrur al,o in organic nch ~h.tk•s. These arc :~ll and futer teeders, b ut wJLb very short food
&l·ritarchs n:productiVl' or l'lKY'tmcnt stage~ of chams. Its collapse was abmpt and the mb~equent
1e uk ~noti( Jlbr:~l pl.tnkton). rhq wcrt• fmt reported Cambrian 'explosion' of ail lmaJ phyla, in a new,
b Timofn·v in 19l)•). Tht· .tcnt.lrch' .trc mually competitive and multi trophic ccosy~tcm was per-
uuJ ~~ flau~ncd and !olded con•pre\stons, and haps (McMcnamin, 1989) the 1110\t decisive evem
Dlt ut rh~ earlier Ollt'' .tn.· 'implc 'PhtrOids. They in the history of ltfe.
m I'C tr3ud b.tck to ~U111e I·WO \11.1, from whJCh
h. e di\'l'l\lt) CO\t' to 'ome 311 tn.a, about 750 Ma
us , Jnd some of tlte,t• l.1tu Jt ntar~h' were 'piny
ot f mort wmph:x torm. The't' wcre dommant
~). 01\t.ll \\,Iter;. U\ tht• ldtt·r Vcnd1J11. \Ome 630 The divef\ificanon of hfc w h ich began m the later
M.! Ill', some 70°u of tlm m h bmta h:.d become Proteroz01c rt·sultcd, by rhc early P.llaeozotc, m the
IWld. a m tJOr event ''hi eh tome 1dcd with the late c~t.1bh,hmem of the m.ml group~ ofhving orgarmms
cambna1 (V.trangun) ~l,tt i.ltton. which persist today. Of thl' many attl'mpts to classifY
\\ h~u tht dun.He amdiUJ ,\led there wa.~ .1 further the esseno,Ll major !:,'TOUps or kmgdoms, th,tt of
l l\ • ICf\llltattun ol cuk.Jrvotlt pl.lllktic .tlgac, but high Whittaker ( 1969) and Whtttaker and M argulis
rn •I'C111ty l~1d' \H~rt not rcJdtcd ag.un until well (1978) has been generally adopted and JS tollowcd
~c- llt' the Earl} C.unbri.m. hen:. These authonties ~uggcsr five major kingdoms
tcr ACt>nccptJOn of t't olo~cal dlVen.ity 111 the IJtcr as follows: ( I) Monera (all prokaryote~. i.e. bacteria,
a l'rnttrCl20IC ~~ g1w11 by .m tiUO Ma (R 1phcan ) cyanobactcria and green p ro karyotic .1lgae), (2)
ol IJIIrntc m Svalh.ml (Knoll .md CaJdcr. 1983). Protoctista (p rotozoans, nuckottcd algae and slime
nd krc rht•rt •re three dt\lltlct .I'>Sclllblages. The first is moulds), (3) Fungi [which cannot make their own
ll1 1 •tronlltohtc-out .~~~l'mblagc, wllh \IX or ~even tood bu t w hich must live as parasites. saprophytes or
.g. r )bl'\ utc ~l.'nt:r.t prc\t:rn·d in ,.bta; the second is a symbion ts (lichcm) I, (4) Plantae (all cukaryoric
fll.l nkuc J\Scmhbgc domu1.tred b) IMge acritarchs plants) and (5) Animalia, includ111g rhc sponges
113[ g 111 th w.uer column ,above the coastal sedi- (parazoans) and multkdlular heterntroph~ (meta-
uk tl Thac .1l~o octllf\ ht:lt' the much-dtscmsed zoam).
he blrrrd•t>ll. 1 lmd, thac ~~ qUttt: dill'erent asscm- All of thcst: were certamly csrablhhcd by the later
Ju- tli >N·-,haped mtt rofos,tls (V~Ms). probably Proterozoic. P r otozoans, whtdt ma\ be photosyn-
ted c h~tcrorrophtt protozoans. wlurh have thetic or prcdaron,1l, rcma mcd 'm:tll and evolved,
lhh td up mdcpcndl'lttl) Ill m,my other locahttcs. ot(en to elaborate fonn, wirh111 rhe tonfme' of a
lt:ll JuTercntul prc,en. .m on mmt limit our \'te\\ siuglc cell. They are rcprc,entcd 111 the to"il rt·cord
58 Mo1or events in the history of life
b} v.tnom kind' 0f mtcrot(w;tl~. mch a~ the ~1bcna. The lollo,ving M iddk .111d L11c C' 1111bn
for mmufendJ, Radtol.ma .md ( httmozoa la~teda remarkably short time (5.20- 505 1\.h). Wt
Anmul~. all of whid1 .m: multtccllul:tr. may have tlm ume~cale in mind we cm now l'IIINclcr the v
bcgtlll J\ clu~tcred agp-qptl'' of protozo:lm whtch tous buna~ 111 turn.
r~ muncd co~cther .tficr reil d1 Vt\1011. One line led to
the -;pongc~ (Chapter 4), rct:tinm~; 111 general ten11~
~uch .111 'aggregatt•' org~mizttion, anorher to the Ediacara fauna : two viewpoints
meta z oans. m which difTc.:tt'IH p.lrh of tht· body
bt'Llllle ~pec1alizcd tor dttfacnt functions :md tn Bdorc 1947 almost nothing w ,1s known
wh~~:h u~sue5 (mu~de, gut. nen-c. ctt.) developed. . natun: of Precambrian met:wo.tn life. The re
Onte the earltc.:st multlcelluhr .uum:tls were 'Protcr07otc' coined for the later Precamb
t'\t,tblt,hed. meuzoan life underwem succcs~tve (2500- 543 M a) wJ.~ ba~ed on the undcr<.tandmg t
raduuom. Of the,t• the tir-r "'·" rhe sofi:-bodied there wcrt organi~m'> (c g. strom.ltolite~) IJVmg rh · (a) Cyt
Edt.lt.lr,l t:lUna of the !Jte Proti..'WIUJt'. then CJJllt' but whether or not rherc w,l\ .mim.1l hti..· was rh
a r:tdtJtron of ~mall 'hell> tn\\tb at the ba~e of unknown.
the <. 'ambnan. and there tollowc.:J a dramatic Thl.' di~~:overy in that ye<~r of .1 rit h and we
'explosiOn of hfe' m whll'h calcareous alb>ac. prc~c.:rvcJ hluna of \Ott-bodicd :mtmJh m th
\hdlec.f protozo.tm, sponge.:' and arrhal'ocyathid~. EdlaLara Hills of South AmtraliJ (h g. 3. I) was th
(d) Pra
conodontophond~ .•lrthropoJ~ (c,pt·nally trilobite<>), ft!"llt .Kccptabic faun:tJ record rrom the J>reC:llnbn
pog.mophorJil\, mouu~c' ami ethmoJenns fi~t (Ciac.:~\nc.:r. 1961, 1962, I'JlN)
m.1dc the-ir .tppeJrJIKc.: .•1~ dttl IIIJilY ~hort-lived The li.ls\ih came from .1 \hallow 111.mm· to lm
'problematic' groups of dubrnu' atlintt} Difl'ercnt wqucntc \nth flag_gy and crmo;-beddeJ '\,llld,ron
kind\ of "kdemm appc.trcd, nude of both aggluti- the occ:hJoual emergence of shtUI\ " ,ho\\ n
n.m·d .md ,ecreted nutenal. of v.mous kinds of polygonal mudcracks at cc.:rtam honzom. The fil\
cht·mtc.ll compmtnon. Although most ot these early an: pre..erved along the intcrtace ot tine .trgtllact•
group'> were suspemton or pos,thly deposit fecdcn, lammae and hard sandstones. App.1rcntly the de (j) soft bod1
then• were some predator~ <lnd .111 c:colcJgicaJ balance an1m:t ls c,Hllt: to be stranded upon thl.' tidal mudfl
wa' 'oon ,tchil·vcd. Thi~ 'C.:unbn.lll c11.plo~ion' wa~ a nr m the bottom of tidal pool\ anti wl'rC prcsc.:rvt
n·.tll·vent, .md not .m .1rt~.·bn ot rhc record; it prob- when covered by sand. The saml is ot unu~uJI tc
,thl> Lt\ted nu more than ,1 t'l \\' tml11on years. ture. rather hke a foundry saud, whJCh j., a factor
The Proterozotc-l1mbn,Hl transition JS now tht\ uncommon mode of preservation.
Tl'llltrkably \\ell cahbr.lted (k.noll. I <J9h). due in part Sl·veral thousand ~pecimen~ of Ed1.1can tos'!l'.
to the n·cognirion th.u orbon 1sorope vanations of now known, belongmg to sollll' 30 gen
th.H nme were globally '>ynchronom. There is now a T radtnonally they have been comtdered a_s JDnclids) ..tlh
rc·tined 'tr:ttigraphy wh1eh alon~ \\'lth new fossil dis- reprcsem:mves of modem phyla. <.·speCially '"'"""""'•'llltic' ~mn
coveries have given .1 fresh pe~pccttve on rhe (GI:ll.'smer and Wade. 19c>6; Wade, 11)72; G ~ervcd, but
'C.unbri:m explm10n' .md the evolutton ofthe earlt- 19H..J.). A contrary. and adnuttcdly hcreucal v rh surlon.·s,
c~t metazoans. however, is that they :lre reJjcs of ,1 'fatled' cvolt
The base of the C.tmbrian (wh1ch in my scientific tionary experimen t, qu ite unrelated ro any modf~
litcttme has hovered bl·twlen 500 and 590 M:~) orgJmsms ,md ':1lmost like bfe on another plan
nm\ ~eems tirmly fixed .u 54'\ M 1 New records (Setl.lchcr. llJR9). ll'.\1)//1 lt'5 lS ll
of pm" hie mtt.uoans. thl· nkk·~t kno\\·n. Jre dated ha' .1 stronl
fro111 .1bout o15 :vta. .Jmt pnor to the Varangian The traditional view arrangcd In
In· Ag~: (c. o I11-590 'v\..1 . md the oldest Ediacaran Accordmg to the tr:tdittOJI.ll rmerprerauon a u.·nrnl <.t,
laun.ts from about S7S \t\.1 Tlw E.1rly Cambnan, Ed1ac.1r.l fo~sils fall 1nto four 111.1111 c:ategone,, diSC. Killlbt•
111 ,!\rending 'cqucrKc: lnmpn\C\ the Nemakit- hdore COINdenng altemative hypotheses, the) ... J.ler, bcll-,h.l
D.tltlyman (543-530 M.t), the Tommonan, tol- fim dl·scnhcd under these hc,1d111~. t'S. T hl' tl'Jll
lowed by the AJrabaniJn (together 51U-525 Ma), 1 he m3jo1ity are jellyfishes (medusoids) .md Kulptmc of
then the Botomian and tulJI!y the Toyonian cor.tls (pennatulaceans) bclong111g to Phyh:
(together "i25-5.21l M<l), .Jlln.llllccl ;Jfter type areas in C111d.ma; there are also segmented wor.
Earliest metazoan s 59
11
h
:-
(b) Tribrachidiu m
c
)1 (c) Spriggina
11
lt
(al Cyclomedusa
11
l-
c
ld) Praecambridium (g) Parvancorina
11
ill
s,
(e) Dickinsonia
1~
1:3 soh-bodied tnloblte

ts
d lipeJ.I Elements of the Precombrian Ediacara fauna of Australia. (a) Cyclomedusa \x 0.5); (b) Tribrachidium (xl);
(c) Spriggina
075), (d) Proecambridium (x 1), (e) Dickinsonia (x 0.75); (f) same, three-dimensiona reconstruction; (g) Parvancorino
0. 75);
11
CbomiociOOJs (· 0. 25) . (j) 'soft bodied trilobite' (xl). [(oHd), (g) redrawn from Gloessner and Wade, 1966; (e), (f) (>
based on
bcher, 1989; (j) from Jenkins, 1992.)
mnelids and some pccuhar orgJnisms of unkno\\ n Pennotuloceons

t.l uur,. Sometimes one surface of thc fos~ll alone ts Clurr11iodiset1s and Pteridi11i11111 are dongatcd feather-
r,. "\t'J, bm there ,Jrl' al~ll compmirc moulds Wllh shaped bodic,. In ClwrnhJtiisms there ts J ba,al dtsc
,. ,Urf.KC\ compres~~·d to~t·th~·r. With .1 central ~hafi: about ISO nun long from which
I-
spnngs primary branchc~ at abom 45° to the shaft,
ll
and ~lH11l' 20 or ~o secondary branches arc produced
t' ·n pwcs of jdlyfish h.avc been dcscnbed. from each of thC$C. Prcntlini11111 1~ \O!l1e\\ hat 'mular,
'511mr 1\ 1p to 125 mm .terms, i\ dome-~hapcd but poorly known, bt>c.mse of cla,nc dcfonnatio n
h,l\ d mongly st ulptureJ \lll f.JCc wtth brge radt- during preservatio n. The affinitic~ of tla:~c have
amngcd lobes. C)'duwt'llmtl j, much ~mailer and been much debated, but the morpholog ical compar-
c l mnl cone With conn:ntric nnt-." and a wtdc isom they bear with sea pens. t.e. alcyonan,lll coral<>
d tll·t 1\rml•rrcllrl (hg. 5.3) probably h,td a much (Chapter 5) wnh polyp~ ranged along the ~ccond
~r. bciJ-,hapcd dome With distinct lonb>itudinal ary branches, arc so clme tlut th~:v arc general!)'
s Tht· lcntJcl~:~ of EL'JlMT'"'I arc prL wrvcd, but bchcv~:J to be related to tlus group.
Siulprurc: of the other njnc jdlytish s~·ncra j, less
11
centre, Jnd as they neat the cdge of tht· pl.nt.. auwtwph
Annelids d mun.tte
Three..· spec1t'S of bro.Kl. A,tt blbteraJiy symmetrie<tl they are ~harply bent, cat.h giving me to .1 ~rm1
1· .1llll,l\
'worms' an: l.:nown, n:ll-rred to as DrckiiiS(lll ltl. They marginal ~triae. In nun) ways J tibmd11d111111 h
n:~.oc..ud~,.d
have .1 tlun central JXI~ and n.trrow segments nonnal ~uperfinal 'iinulanry to tht.' edno,l\te roid' (Char
Ne..'\\ loun
to it, arrangni Ill a 'quilreJ' p.tttem radially around Y), espeCIJily m a few gcner:J. which h.1vc trirau
'yuunetry . Some 'pecmten s seem to h,l\'l' a Sc::\lldina'
the from and n:.u. ~omc spc..omen s bear gut lobes. ,J
shaped Cl'ntral mouth .Uld bnstle-hk c apputdag~
lll.lflllC.'
but no mouth or e~c..·~ h.we been pre,Lrvl·d. A dos<-" fnma (G
rescmhhn cc h> the modern segnH.'nted wom1 the surf.1ce, cmKeiv.l bly precur<>ors of tube teet. T
thout pc:
:,pmthrr whll h 11'1:1.'~ c..\11 and l'Jt' ~pongcs, has been po~~ibiliry that dus am 1ent fos-.al wa' a kind
nf c~hout
noted. ~o Dul.:rmtlllltl lu~ been com1dcrcd to b~: a~ 'proto-e~:hinodeml· IS rherefor~o• nut to be dr
garded. Some ~pc:cimem of r;Hhcr trilobtte-1, Ntwfmtt
annelid. It dllll·r~ fmm Spi11tirrr only 111 that the..· di~tal
appearan ce have recently been n•torded Oenl1 de•·pc:r-Y
cl.m·<. (par.lp~>daa) suu.tted on the end' of the scg- pceunatc
mcnc; .md uwd tor c. r.nvhng :tn: lacking. Drd..:11rs.•nia 1992; Fig J.lj) from the EdJ.lC,lr.t Htlls, .n van(
~tJr-~h.tp
spclum·n s mu.dly lie tl.u on rhl· bcddm~ plane,, but ~tagc~ in growth. Mtght rhesc..· be some kmd
~oft-bodied rnJob1te prenmor ? We av.. ait furtf
on the 1
some. of rhl· l.1rger on<'~ arc ddonned by traltiOn, referred
allowmg det.uh of tlw quilted ~tnKturc..· to be dan- ev1dcncc .
the .tgc
fied . In thc,c..·, ~harp crcascs cro~sing the qtulhngs nu know
shm\ that the.. cundc.. nnm h.tvc bt.•cn leathery .111d Vendozoan hypothesis
~eti.Khcr ( 1989) reJeCt~ the view th.lt the Edtat
hue lakt·
ftextbk. Evi,!t:ndy tht• uppa .md lower cuntular roc:b nf
smi:tl't'~ wen· not knm\:d togc.·ther at thc..•ir outer cir- organism s are ancesto r~ of modern metazoan ph1
nmlar J
cuntll-n:a Kc but hdd ap.m h) -;ttH: btradially lie categorize~ all these orgaamam a' Vendo.
arguing from the srandpmn r of l'Oll\trurt iQnal n • ont.1in
an.lngl.'d ~q~mc:nt.ll 'L'Jlt.t gc:nu' (.
Th~o· elegant Sprrl!_t,!lllc1 ha~ ,\ hor'e~hoc-sh.tpcd phology. He comment~ that in 'Ptlc of ,Jn appa~
Umil
'he .1J' with t~>ll)!. prt~ll <tmg ~et.tt'. Some 40 sq~ments dtversity in the t-:tunJ.. nearly all the gencr.1 havt
~Jp bc!t\
tollo''. eat h. tt.cordan~ ro \c)lllC authontie s. tenm- striking ba~K umt()rmit y: the~ arc thm and tlatt(
round or leat:.like. and pos~cs\ ,1 ndgcd nr llllllt tht.' C.tn
nanng i 11 a slaort parJpmlj um, .md hence it ha\ been 1\ollll'd
con\IJcrc J tro be Jn .mneliJ. '' ith 'mnc pmnt~ 111 upper surfJce. Moreove r, ~uch tonns as /)itkiu,
hctore t
common wtth modem planl...wmc wonm. (fag. 3. l,e ,t) arc l·onstructc.·d of btrad1aJI} arn1
l~111pU1<
hollow tube~ of Acx;blc matcn.tl. open m the o
~ide. H ere, as wHh aJI Edi,1c.1ran oq~anism,. thne
ami the
Fossils of unknown affinities prc ·t·rv.
Pr<tt'!lllrrfmdrrutt h .lll n\ otd dtKotd.d fonn 'omc no trace of a gut <iuc h fcJtures 'ug_gesr eh \t all t
<~1 •• I'JlJ
5 mm long. 'I he 13 known 'pc:c1mens show three animals fed by p.1ssave abmrptio n of das,olved fi
I IUS \et
01 ftmr p.m, l>f r:J.i~cd lobe<>, ~cc:mjngl) arrangt'cl in .111d mctaboljt cs over thl.'ir whok ~urfJrc. If t
~peLt..)
segmc..·nts, bdund .1 ho~c~lm~-,lupcd antenor pare. were the case .•md 1f intcmal digc.>\tl\'C . v.Nul.!r
transport Jtlon sy~tcms were laclung, then the'e • cqm:nc
Some: authontac~ (C;Lic..·~sner and W.tdc:. 1971) h.wc
mals had no rcl.1tiomhip with any modctn gn ) utli1J4e
interprete d tlm .1\ an arthroptt d with a soft. non- cxt~ ud•
nuncr ahzed mtt:'~tmult. lt h.1~ ~ome re\c:mbbn c. c to (except. 111 fl·,·dmg mode. wtth pogonoph or
Ma)
and even Spriggi1111 may be cnvu.t~ed, not J\ a er~
,If
a tnlobtte prnt,l\pis or larval stage, hur the an.tlogy is ~!Idly
not J dme 1me. A ,i.Jmlar though larger fonn. ing wonn. but .t~ an upright frond , hdd 111 the
by the 'head'.
·11uwm
r 'rlllihl. ha' hl U1 found in Stben.l. rcscmb
[>,,,,,,corllltt 1~ .1 ~hjcJJ-!>hapcd .mimal up to If tlm tnterpret. Hion 1s tollowc:d. it " pc)\.\lbl
11 uly 1
:?.6 nun Ion~. wtth .1 hroad, am.hor-h ke ridge on the envisage the Vcndozo a J\ lying on the surf.111
bacterial or cy.mobac tcnal mats, and manv llldV b bout
uppn ~urf1n .md .1 nJrrO\\. mn~<:d nm \\ ithin I ukut·
the m.1rgin . F111e strt.ll' extend obhqudy bat kw.trds had endosym bmnt\ \<\'lthan rhear bodies. The w
tu!( tin
from the m.nn ndgt•. Somt· .luthorc; havt· suggested of an cndks~ ~prc.·ad of p.tssivc, AJt grt•t·n quilt!
quire appealing . Yet the 'fall trom tht' \..udcn nd c.
arthropod an 1ffiniue' 111 vie\\ of a suppo~ed re,cm- LllllCd
biJntt' to dw \hell' of the ltttk" crust.H.:ean fntJps, Ediacar.l' (MeMcn amin, 1989; McMctl.ll lllll
1lls be
but thJ!> is spcllli.ltlV l' McMena min, 1990) came abruptly. With tht
becu t
Tnbr.1rftidrum t\ dJs~:oidal wtth a n·ntral raa~ed of the tiN predarorc; in the early Cambn,111 the
tem of global c..Toloro- r hangni from pnn 'kt'lctt
pbtfonn. Upon thi~ thrc:e ',tnm· r,tdaatc from the
Earlie5t metazoans 61
11 roplur to tlw lwr~rOtlllphtL '"'tt~m that was to gasrnc cavity and lodged \\ tthin tht· me~o~loe.1
111 tc: unul tht· pre~ent tllnc. (Chapter 5). Thi' would givt• \Oilll' ng1dt!) .md
Ullli of l~dtac.1ran ag~: .mJ t) pt· havl.' been would anchor the coral 111 tht• sub,tr.lte Ncverthe-
rdcd trom vanom pam of tht· \\Orld: Amtralia, Jcr,s the body piJm of mmr Eds.lt.1r.ln org.11mm~
1foundbnt.1 ~tbnu , Englaud and Wall's, arc V~f) different from rho\l' of other met.l:loam.
nma and N.mlJbu . I hq .m~ gem·rclllY of and d1cr~: rernam~ lllllt h 'upport t(lf "i~JI.Kher\
ne hAllow-water ongm . I ht· diVerse Amtralian V endozoan h} pothe'1'
(C.Iu~''ner, I9H4: jt•nkm~. 191J:!) d.lte' from The Ed1acaran f.1l111.1 range' throughout much nt
perh.tp~ SilO M.t. A som~:what older tauna the Vendtan, a time 'PJII of '><Hilt' ~~~ Ma . .md tht•
llxlut 5115 'v1.1 trorn tht: A\ .1lon Pcnimula in divef!>ltY of t)'Pe~ incrc."e' through lllnc Thcrt' ''
rtoundland occur, 111 scdim~nt\ of reLltively only one undoubted ,\...ek talin·d f(,,,JI of mnilar .1gc.
00'-llltcr ongm. ar vanom hon7on\, Tht:y are the pipe-like an n ulated c.tltart'Olts tube knnwn
tute (romb-ltk~:), bushy. spmdle-,hapt:J c1nd as Clottdiua, which t<ln b~: up tn 4 u11 long; in
u-,hlpcd t(mm hdon1-,rtng to .1bout .'\0 genera, .and Nam.ibia th~:sc wbes Jrc so .1bund.mt, Jlon~ \\ tth
the tr.ldJIIIlrlJ! mtl'rprer.mon m1ght be largely undescribed goblcH h ,1p~d lllJLrofo~'ils, ·'' tu funn
t'l·d to tht Cmdan.1, as nughr be ~:xpected from biodastic sheers. lly the cud of the Vendi.m, .1 fcv,
lb'l' of the f.ll111 .1~. but I0 of the genera arc of small shelly fossils arc prcscnl (AIIrlbmitcs .md
~IIOI\11 ,tffimty. There .m• no wonn-ltke or trilo- Cam/lrowbulus), pre~a~ ng Lltc1r dr;un.llil cxpamion
11 ·-lilr i(mm pr~~enr. The to\~1hfcrom Ch;~rnian in the early Cambn an.
l. kl of rentrJI England .ut• now known to be of
Jr Jg<: to tht• lower Av.iltln '>llltt· and lake them
••'- tAJO .1bund:mt '>pt•rum·m of the long-r.mged Srnall shelly fossil s
1L Ul Clr.lllfl!ldl.\111~.
ntd rnently ther~ \et·nwd to bt• .1 subq.mtial In recent years nch .md van~:d tauna\ of small ,belly
bet\\ rut tht· l.tst EdJJc .tr.m fnm.1 and the base of fo,,j], (SSF,; Fib"' 3.2. L~) h;~ve been dm umented
C'.arnbmn. rrint(uong \olacht•r\ concept of an from the base of th1. C.1mhn.1n .md Jmt into the
ted tirst t•xpcntnt'IH', \\IHdl died out long Prclambrian (e.g. Matthew' .md Ml'-\.117hev,ky. 1975;
·Use C:unbrian b~:~.111 . But tht•re •~ no longer a R.aJbcn. 1981; Roz;mov and Lhuravkv, llJlJ2)
raJ g;tp bct\wen tht• voungl'~t I:dtat aran tJuna The hl''>t- known \l'l]llt11Cl'\ romt• from the
tht first small ,hcJh IO\\tk rht\ I\ purely a Sibenan Platf(mn. \\hen· late Pr~l·ambnan rak.lre-
k
~
~t~en•nonJI .trte(llt. In N.1mib1,1 (Grotzmg~:r et
19<15) thcrt ha' b~:~:n tk'l nbt•d .1 nthl~ tos,tlifcr-
ous bed~ with dolonunc. ouhm and \ tronutolitic
horizons are ovcrlam by t•arly ( ambrian '>edum=nt'>.
~s stqucnce with .111 .tbund.mt Edtalar.t f.llma, The Tommorian stag~: dot''> not comatn tnlob1te~
d 1.UI) Ptmd111i11m. ~t\l'ral tutl\ Within the but h<l!> Jrchat:ocyathid~ and .1bund.1nt <.;Sr:~. many
1- a.u1e pro\ 1d~; rdt.1blt· U-Pb J.m·~. md the with phosphatic shells. Somt• oftht•se (e.g. rlldrml'lla)
p .. 111~r\t N,mubt.ln Ldi.IL,Ita-typt' a\semblagc are hdically c01led gastropod~; thl't'l' .lrl' also
}. uJ, right up tu tht• b.t~e of tht• Cambrian (543 h yolithids (an cxtim t m o llu,can t,rroup) .md com-
I- \11 md •~ conternpor<IIH.'Oll\ \\11 h the ti~t \mall m on ~po ngc ~piculcs. Tommot1.s n fJunJs .1re rt'plctc
d ll1 tos~il ass~mbl.lgl'~. ff the Middk Cunbri.m With short-JiVL'd problcnutic groups of w hich two
111/IJJ'Ir/on from the 13urgt''' Sh.tk·. whid1 dosdy are of most interest: the hyolithelminthids Qami-
na ted phosphatic tubes open ;lt both t:nd~) .md tom-
,,
() mbb C/rami!llfrstll5 (Cormay M orm, 19CJ3) is
~n Edsac.tran surv1vor, .md tf more " known motiids (phosphatic conoi<hl 'hl'lh mually
c.: ut the \oft-bodi~:J tnlobttL''' lllL'ntloned by occurring in 'Yillllll'trital pai~) Thc~c !.mer have
11 u ' (1491) from thl' Ediat.lr.l I hll~. tht'n the dis- been compJrcd wnh pbtcs of banude., (nmpede
IS tllfl h·twecn .lt k.1c;t <;Oim: of tht· Ediacar.m taxa crustaceans), though they arc phmphattt rather than
>f .:J l.unbmn faunas bt•comt'' Jt..,~ dear. Thi' i~ sus- calcareous. If so, 1t would su~gcst a Vl.'J! early ongin
d ."; ')l·ilarher\ (I YlJ2) tom me-n~ on th~: curi- for these crustaceam . fh ~: rcsemblan<:l's, however,
WI. ar.m p,:tmmocor.tllt,t which '>t'em to have Jre only ~upertinal, and rh~: tommotiid, probably
.- tme .mrhozoan polyp,, t.l'. ·,and corah' wnh belong to an cxrinct ph) lum (B~:nh't~on, 1977).
tons fonned by ,,md gr.um wluc.:h t•ntered the Some of thc,e mltrolo\\ih may .lltu.tlly bt·
62 Motor events in the history of life
(b) Aldanella
(c) Tommotia (d) Anabantes
(a) Fomitchella
(e) Protoherzr
(g) Coreospira
(h) Chancel/aria (j) Yochelcionella

(f) Lapworthel/a
(k) Chancello
(ml Mobergel/a
(I) Cambroc/avus (n) Canopoconus (o) orthtd brachiopod
Figure 3.2 Latest Precambrian (Anobariles) and earliest Cambrian (others) small, shelly fossils. (o) Fomitchelfo (x 40); (b) Aldont
(x 20); (c)Tommolio in obverse and reverse views (x15); (d) Anoborites (x15); (e) Protoherzina (xl5); tn Lopworthella (x 20),
Coreospira ('> 20); (h) Cllancelloria (x 30), (j) Yochelcionella daleki (x 8); (k) Choncellorio romifundus (x 30); (I) Cambrockn<
(x 50); (m) Mobergella (~< 5); (n) Canopoconus (x 50); (o) orthid brochiopod (xl 0). (Mainly redrawn from Rozonov ond Zhurcrk
1992; others from Oion Yi and Bengtson, 1989; Bengtson et of, 1990 l
fi9ure J.~
foscicufoto
Dreponocl
tomponents of larger orgalli'm' Thus ~mall plate\ )JmJlar foss•ls ofTommottan age (llcngt~on tt
(Photogro~
Jnd sptnes w1rh a nct-hkc ornament (•\ficrodrayon) I990) arc known from Au~rr,tha, England
h,l\ c now be: en ~hown to be pJrt of the: armour of .1 Sc.tndln.w•a. Many of the ~penes connnuc weU
br~(' ,]ug-lil..e fom1 (Ch,lptcr 12). into the Lower Cambrian, when· they arc toun;
S"'h ap~
An cxcepuonally cumplc:tc and important section with tri l obite~. By contrJst, there <lrl' rcbtivcly ie
then ont
through the btest Pret·,unbnan and early Cambrian microfmsih in the latest Prccamhrian stage: tl
llnrthe.lSI
ha~ been <icKribed fwm Mcishmun in Yunnam, YuJomian. There arc only a few ~null tubul
K.1u fin,111
Chma (Q1an Y1 ,md lkngtsuu, 19~9). Within the organ1~1ll\, e.g. Atwbarites, a phosphanc tub-
•nn wic
Mcl\huum1an stJgc:. d1rcttly cqtuvaknt to the Clollllilla, an annulated cakarcom ttJbe, a
st.l~t: rest
Tommot1.1n. three thronozoue' and c1ght subzont"~ S,lbclliditrs. an organic-w,11lcd tube, thought to ary llcr
hav~ been erected on the ba\1~ of ~~Fs alone. The the skeleton of a pogonophor,m (a simple un~
di\.CI'\lfY!
tiN trilobite<; do not appe.1r unnl the overlying m~tntt:d 'wonn' that has no gut but absorbs d1s~oh
tlll' h'fl:Jt
Qumzhu\t::tn 'tagc. The ,1buml.!nce of trace foss1ls orgatuc matter over its surface).
th<' tulm
and .tcnt,lrchs m.1ke th1' .m 1dr.ll \ecrion for srudymg In the: type area in southea\tcm S1bcna and d
ongm of
the ~equenn· of e.uly C.1mbn.m l'vents, and a candi- where, the Tommotian rem unnmtonnably on 1~
IU111.1ry b
datt• t()r the PrccJmbri,111 C1mbnnn global bound- l.1te Pn:cambrian Yudomian, and thc Nemakl'
11 tonn<:r
ary ~tr:ttotypc. l).tJdyman stage is missing. Since, in the Tommoru1
Precarnbrian trace fossi l s
The e.1rlicst appearance of rr.1ce fossils may help to

date the origins and init1al r.td1arion of the mc•t.1-
zoans more readily than do the soft-bod1ed org,m-
isms themselves, \\;th rhe1r hmited prospect' of
preserv.mon.
Tr.Kc' fosstls, however, do not extend f.1r back
mro the Prorerozoic. The oldest so far dc'>l nhed,
which at first seemed to be the burrow of .1 woml-
zina like org::umm in rocks I000 M a old, is now bd1eved
to be a renntte burrow. Trace fossils ach1evcd \Omc
def,.rrec· of d1versity in the Vendian, and the1r nukas
Aoumhed 111 shallow ~cas With ~andy floaT\ (Crime\,
1992a,b). There are scattered records oftraet: fm\lls
in rock~ of over 750 Ma, bur trace fo~~ib do not
become· abundant or diverse until ne.1r rhe
Precambn:m-Cambn an boundary.
k:ma Mo,r of the early trace fossils probably n:mlted
from the burrowing acuv1ties of ~oft-bodied mf.w-
nal wonn\, but the number of type~ represented
tncrca\ed !,'T<.'atly in the uppennost Vend1an: hon-
zontal ~inuous feeding burrows, surface trails, rcsnng
marks ami vertical pipes ;1re not uncommon ~omc
little distance below the Precambrian-Cambri ,ln
bound,try. It is not alw,tys easy, however, to date
these 111 .tbsolute tenns, nor 1s 1t clear wh.H kmds of
me/la
I); (g) anim.tls nude them.
:lovus However, m many sequences traces ot .1 kind
::~vlev, norma.lly attributable to trilobite resting traces
(Rusoph yms) appear some cons1derable stratigr,Jplu-
r~L~ 3.3 (a) Archaeospira ornate (x 35); (b) Lopworthella
i:ic(IJ/o~ (x 70); (c) Maikhanella prislinis (x 35); (d) cal distance below the trilobites themselves. They
~ile$ dilatotvs (x 35), (e) Scoponodvs renvstvs (x 35). an: mainly sunpk fonns with paired chevron-uke
:t al .. ~s reproduced by courtesy of Or Stefon Bengtson.) marl..mg.., evidently the ~crarch marks ot the
and
appendage\. They may have been made by late
·U.ur Prt'cambnan trilobite pn:cursors with thm, non-
ound lppcar en m,h\e, tiH.· 1mpression 1\ g.vcn (hat minerahzed ~hells, propelled by 'a siJnple muscula-
1 fc\\ 11 ongm was very .lbmpt. rtecent d1scoverics m ture of low efficiency' (Crimes, 1976).
: the xthcJ~tcm Sibe1ia, however (Knoll, I996; In general, trace fossils arc more common than
b ular ·fuunn et 11/., 1996) n:cord a contmuous succes- body fossil~ in the later Precambrian, and whde they
tube, \\1th the b,ts,tl C.unbnan Ncm.tklt-Daldyni,m go further back in time they do nor rake the pre-
and mtm~ confonn,tbh on the Yudomun bouud- sumed on gin of motile metazoam very far back m to
to be Hm we see SSF, appe.uing sequennally aud the Prccambrian, certainly no more than 10011 Ma
nscg- ~mtymg through .1 pcnod ~ome 3-6 Ma long, and before the present time .md probably much less. 1 he
o lved gtllt dJVemty or SSFs m the Tommotian is only first good trace fossils postdate the widespread late
nrlnnrlolt:Ion of a loug evolution. Accordingly. the Precambrian glaciation. Following the fir~t V cndt:tn
else-
ot SSFs was a \tep\\1\t' process, ,tnd the evolu- divemficarion, a second raJiarion of trace fo'is1ls
m the 1\ burst thereof a mu1.h les.~ dramatiC: I.'Vt'm than took place in the Tommotian/ Adubaman, and
nakit- ::mer~· ~eemed to be. three trace fossil biozones have been estabh,hed tor
Jotun,
this mterv.tl. Cunou'l). there: w.l, \'ery ]jttle change scale. clustered .truund the Prcombrian-Cambnan virtually cxn
in rhc rrac.:c fo,\11 baot.l for tht' rc~t of the l ower boundal)• (Cook and Shergold. I 9H4; Cuok, 1992 llnn any otl
Palaeozoic.:. other th.111 ,1 po,~tbk nugr.ttwn of some It is likely that dunn~ the l.ltcr Prcombrian, phor me and decla
types mto deeper water. phoms-rich ~edunent\ accumulated on the dt.'e~ ~on temporal
occ.m floor as tlm clement w,\., d~plcted from atcd ,lnd di•
mrfacc ware~. Dunng m ext<:nded period 1 ommouan,
enhanced occamc overtum, m thC' e.1rlie't Cambrun brachmpod'
Couses of the Cornbrion 'explosion
(and po~sibly J~~oc;aated With the marine transgr~ Thae uc.::cur
of life'
sion) phosphonl'> wa\ brought up to surface watel' problcm.ttK
The prohferauon of 'kdctaliLed lo~,,Js 111 the Earl) to ,giw uKreased nutnent levd' jn the phone ZOIIC' tzcd. All oft
Cambn.111 wa\ ,1 !'\.',,) phenumenon and not an :rrtc- Phosphoru~ IS esscnual to life and tlm event nw organiz<:d ar·
tact of thl" geologH.IlrcrorJ. The .1ppear.mce of the have provided the trigger for the sudd«.>n diverslfic:a· ant:c, wh~eh
mam Jnvcrtehratc phyl.1 \hl' plldsed through some tion of living org.umms (,1s well a' the atcumulan understand.
31HU Ma, but tt \'v.l' .t c.:ntital tummg pomt in tbc ofphmphorites on the \C.l floor). It" ~;igmficanr tt There Jrc
hiscory of life. But l.lll we e~tabltsh why it hap- many early Cambnan org;.umrm (e.g. elements f .1ppcaunce c
penl"d? Spccifiolly, llad duugc\ in cxtcmal factors rhc Tonunottan faun.t, amllingulitorm brach10pod· One is that
dnve ir? Or wa~ 1t mcn.:ly th.1l hfe had reached a par- luve shells of c:tkium phosphate, which w::ts w~ perhaps due
ticubr evolutimtJry thr!.!~huld? Quite ('vidcntly, abundanl at the time. H igh phosphorus lcv~ oper.tting at
borb plwsicochcmJc.:,tl ,tnd biologiCal factor\ were inhibit calcificatiOn in living organisms, which nu :~t:qlll~ition o
involved, ro~c.:·thcr turc.:mg the.:· lliOSl dramatJc event explain why l.llcJreou~ skcletons cLd not appe unmincralizc
in the hi,rory ofhfc. commonly 111 the fos~il record until later. when thi bearer'> there•
c.trly Cambnan pho~phogemc event was largcl tnl not avail:
Physicochemical foctors over. lhough the
During rhe V cnJtan anJ L1rly Cambnan there were The spread of 1\llrTient-ennched waters over lu\1' non possible
maJOr change' in ,Hmmpla('nc composition, oce.m lanrude shelves, as recorded m C-tsotope ftucr aboo,;e group•
chenmtrv. pi.Jte teltomc.: altl\'lt) .md dunate. To tions (Braster, 19lJ2:t-c). ma} \\ell be lmked to tt was 'pre ad
be gm \\,th there ".1~ a ,Jo\\ build-up of free armos- global me m temper,lrure ,\\ lhe Earth emerged tr the Cambria
phenc o:\-y~cn JS .1 rc,ult ot phmo5ynthesi~ wluch a la re Prec:unbnan 'icehuu ..e' to the 'greenhou~t dJd .1 nuneral
evrdently anamed a Ien: I ~ufficic.:·nt to ~upport phase of the latt:r Camhrun. first mstancc
animal life (Rhu.td~ and Mor\e, 1971 ). and likewise Finally, there were t:ertamly four, probabl) \\ that the calc~
a decre.l\c rn the concentration of CO, in the chronous, cvt.>nt\ of global rt:gre~~IOn and erna 11 " compos1
.1tmmphcrc (Tuckt•r, I'J92) fhcre 1s still much gence dunng the time 111 question. The tiN t' pm\lbl) ,\CCU
doubt, however, ,\S to how much frt•e 0 was actu- Kodituan cnm, marked the end of the Protcrozo 11 Once it v.
ally present 111 the J:>rotcrozoac - there may in fact and the virtual demi~c.:· of the l:.dr.tc.tra fauna. 1 \eS\1011 of ::t 5
have been ,1 llur amount (McMcnamin and la~t. the Toyonian cri., is of the latest early Carnbn JHU\de attac
M cMcnamin, lY9U). Other f.1ctors anclude a world- is synchronous with the end of the SSP radiatu prmpects) an
wide ~cncs of man ne tr.1mgrcssions, the fif\t tollow- The effect~ of thc~e scvcralc.:nses ,1re currently a \UI 'elective adv<
ing rhc Varan~an glananon, ~uccl!ssivdy flooding jcct of detailed study. IIIVCrtebrates
the corltincnlal sheh·es <ll1d opening up new \\ :ty would tl
shallow-water ecological nirhes. Most of the conti- Biological factors •ry dcvc-lopn
nental ma~~es at th.1t time IJy in low latitudes, and The most faamliar rcpre~t:ntatives h.1d originate
thc wamung, shallow ~hclf seas were npe for colo- Cambrian faun::t arc trilobilt:s, archaeocyathid,, t to Jevdop h
niz.ltion. It Ju~ been suggested (Brasjcr, 1Y79, tain primitive molluscs, 'mall brach10pod-; (t:htc ~lb'IHficantly 1
1991a-t) rh at cvolut!OnJI)' p.mems of diversifica- with pho~phanc ~helh.) .111d the earhe~t c:chmodc• Eukaryouc
tion may be lrnked w1th \UccessJ\'t' stages of flooding (cdnoastc rmd~. hchc.:oplac.:olds and others). Somt nd produce
of the shelves. L1kew1se. the Jvailability of extra these are found in a few t,olated localme~ alom 111g. The; an
food, e~pec.:1ally phvropi.lnkton. nuy be related to a ilid not spread unnl l,1ter; othel"\ arc wide~p~ b10minera
major penud, of upwdhng Thi' latter hypothesiS from the start. The arc.:haeoq ath1d,, for in\U 1101 realize t
seems to be bnmt' out b} the remarkable syn- appeared fir.t and ~pread from rhc1r Stberian t:ct meuzoan eel
chronelty of large pho,phorite deposits. on a global of origin to or her part~ of the world only to bet P<'CJJ!Jzed or
11 ::uJ!h cxunct b} the end of the Toyonun. More tacc~ for cry,ral growth had become av~ubble I hc
~). ~ny othL·r group. they dram,ltlt-.lUy chart the incorporation of the citric acid cycle n1to the 'ellu-
,_s- nd dcdm<.· of the Early C.tmbnaJJ h1ota. Whtle
empor.mcom phosphanc-shelkd ')~F~ prolifer-
lar mctaboli~m of many organim1.s provtded .1 v.1st
increa~c 111 avatlable energy for biominerahnt10n.
Ill d Jnd divef\thecl. reachmg thetr .u:me in tht: Somethm~ of the vast divcrmy th.lt hfe had
of 'mnoti,lll, the fir~t trilobite~, echmoderms and reachcd by Atdabanian timt:s is shown by the
lll t ),btupod~ did nnt .1ppcar unnl the- Adt:tba1uan. Chengjiang 'window' (Chapter l2). So divcr;c ;lnd
S- rrc occur too ••1 good number of peculiar and 'advanced' is dus fauna that ic would sl'em to ~ignif}
ri okmJtll llh,j(,, often geograplm.tlly very local- either (I) e.xtremdy rapid evolution of mo~t rypc~ of
c. (J. i\11 of thc\c appear very .thmptly .md are fi1lly animal~ 111 the e.uly Cambrian or (2) .111 origm for
1\' uuzeJ anJ tMTcrt•nnated on their fir<>t appear- ceolomates (.muua(, with an intcm.ll cJvlty 111
a- \\htc:h ar fiN stght may \eem d1t}lculr to "hteh the gut IS ~u~pcnded) some ttme before the
m u.nd. Prec:unbnan- Cambrian bound.1ry. lt j, h1ghly prob-
:ll There .trc t\\ o factor; by whtch the very abmpt able that both laccors were involvt:d.
of ppurancc of the~e earl} fos,ih r.lll be explamed.
1s) l~e 11 that evolution .tt thi~ timt' \\'.1\ \cry rapid,
0 1rh~~~~ due 111 part to intense ~dt·ction pressures Biological evidence on metazoan
I~ • rjting .lt that time. The ~econd point '' thnt rhe relationships
Jllttlun of ltJrd shells a' nppo~ed to merely an
r unmlm:d Integument must have brought the There are sevt"ral level~ or grades of or~uzanon 111
1m thereof to .1 'fos-.ilizanon thre\hold' ..1 paten- mulncellul::tr a111mals hving toda), the compar:mve
not a\,lllablc even to thetr tmmedi.ltc .mce~rors. swdy of wh1ch ~heds hght upon the prob.1ble tourse
ugh tht• nuncraliled 'hell wluch made fmsthza- of thetr evolution (e.g. Clark, 1979: Wtllmcr. IIJ\)0:
J10"1blc mu~t have bet·n Kqum·d m all rhe Nielsen, I <J<JS; Get·. 1996).
vr ~:roup~ over ,, penod of .1 tcw mtllmn year; - Undoubledly the earliest multtcellular .mrmals
1111S '{W:td out 111 otht•r taxa throughout the re~r of were aggregate\ of protozoans. looscly mtegratcd
Ill l :unbmn .md the t•,uly Ordovkian Bur how but with cells :tll of the one kind. Some of today's
a nnneralizcd external ruvcring ori~11.1te in the simplest sponges would serve a~ models for elm
t 111\tance> One mggesnon (Giac~,m.·r. I 962) is grade. Sponges (q.v.). however. developed along
1- the lalcarcous or phosphatic m.ttenal of which thetr own evolutionary pathway and :tre not rc.tlly
r- ! romposcd ongmated a' an excretory· product, metazoam.
le bh Jl"CUmulating over the 'km and hardening All other multKdlul:tr organism~ ha\'C the1r cdh
c, On e it \\a~ formed qllltc Jcudentall}, the pm- orgamzed into tl~\UC\, i.e. cell layers or mJS\t'\ \pc-
'lt" n of 3 shell which could be med .•, .1 base for ctahzed for different fimctiom. and tt 1~ tlm char
n. k ltt.lchment (gtvmg enhan1xd lncomorory defin~ tht: mctazoam proper.
n- pcrts) and as a protection would ~vc a smbTUlar In the simplc~t metazoan grade, rcpre\cnted b}
nve .ldV.11ltage to the ammal po,\L'\Sin~ it. Any Phylum Cnitlaria. the organization i~ diploblastic;
Hnrhr.ttc~ with h.1rd coverings fonncd in such a that is, the body wall consists of two layc:r>: an ecto-
would then have great potcnual for cvolution- derm and an endoderm (Fig. 3.4a).
dndopmcnt. Moreover. once h.1rd mouthparts These together fom1 the bag-like: body wluch
on~;~nared, the selection pressure on other phyla endmes a \ingk cavity. ~erving as a gut, wtth a sm-
1dop h.trd covenngs for protection would be gle ope-ning which Jets as both mouth and anu'
firJndy mcreased. The next level, which includes all higher types of
E bf\·otic cells m general are able lO pump 10m animal-;, I) the triploblastic grade. In tht~ condttlon
produce prorem mamces capable of mmerauz- there is a thtrd layer, the mesoderm , ,,mdwtched
TIJn Jrc 1lready pre-adapted for the proce\scs betvveen the endodcnn and the ectoderm In tt'>
bln!llincrah7,1tion (Simkm. lYH<J), but the) did most mdimenlary fonu, as repre~entcd by the
l', r~aiJZt' thctr fi11l nuneraltzmg potennal until unsegmented flarwom1s (the Platyhclmmthc~).
c U7oJn ccllul:tr ~ystem~ were well LlliTct cmiated, the body is bilaterally symmetrical .tnd the nwso-
··l.tlitl'd organs had originated and exLcn~ive sur- dem1 fonns a more or less ~olid m:t~\ of tissue
(e) not have ro follm\ the contour<. of the ecrodc .m: le~
The coelom nuy h.1vc fum tioned pnmmvel)· a1 Pro to
ectoderm hydrmtattl skeleton (the conlamed fluid gt\ llld Ll
some n~d1tv) .•md tlm t1mcrion is retained in t nd ol
amtclid wonns r hcsc move by waves of contr.l '\C<fllCI
nom nmning fon\ ards !Tom the rear. Each eo rout c
tracoon force\ the lilt ompre''>lhlc Amd forwards earl)
the d1rCt non of movement, and 1t 1~ against the fl 'hbstu
endoderm mclf th.u thl· mmde' l·ontr.llt. Many other u Ill Va~
h.1vc been found for the coelom m htgher aninu hole .1
especlall:- a~ a ~p.1ce tor stor.lgl' of mtl'mal org the m
and m.lten.Jk The devdopment of the vanll 'luzo
orgam m .lmm.\1~ l.lll be fc)llowcd in embryolob~ dm c •
Organs of ectodeml.\l nngm mclude the .;km Jn .~red 1.1
nervous ~ystcm, mosr of thL' gut .md ~ome a~sociJtc the. ot
organs ,1rc cndodcrm.tl, wh1lc rcspir.1tory aml exLr .md tl
tory organs Jnd .1 1lmuscle~ .1re mc~odermal. tllll t:;
What t.ln be csl:lblished about the rclatiOJl,}H bC\011
between the mam invertebrate phyla? T he ftl\ (d~lltl'
n:cord 1~ of little value here \!nee the phyla, l.l', .lS
which at le.ht 32 an~ already defined on their f, th ·l.u
.1ppe.mmtl'. lt ,bould bt: po~~ibk to e\t.tbli\h hO'i lA
the phyl.1 the1melve' are hrrouped m terms of eo atnr~
(c) mon ancestry m 'mega-~roup~· or \uperphyla'. "1rd

Those schem~ th.lt h.1ve been drawn up ,jJ1 cc hmc
highly vanablc. Some author\ favour a more-or-le V.J\ • '
orderly progression from ~unplc to .Id\ anced or het~\ l'
ism\. R .uher more cnvi,agc some kmd of rree-b ''1wn

pattcm, wuh d1chotomous branchmgs, others ag (~ea '<
envtsage a polyphyletiC on~lll tor mmr inven very~
brate~. the vanoll\ phyla or ~uperphyb bem~ deri\ '" be

independently from promtan ance~rors. The p (plutc
lem, .1~ W1llmt>r (ltJ~O) puts tt, IS 'attempting Such
reconstruct .1 trt>e w1th evidence only from the m '.1 l.1rv
recent le.tves .tnd .t ft>w twJgs". J VC')
The hkely ,tncestor of later groups (itself denn lt I\ I
from a prot070,l n precursor) Js comidered to hav bvbri\

in melozoons: (o)
been a ' pbn11la '. a small, ciliated, ovoid mass of soh In 'h1
Figure 3.4 Grades of organization
diploblastic, (b) acoelomate triploblastic and (c) coelomate cells, flo:lting in the sea. r rom this were mdepen tlw :u
triploblastic. dently denved cnidanans, t'tc:nophore~ and pos~ibl ll'tual
sponges. The next ~ta~e was the origin of acocl(' ,,f la
mate A.1twonns lil,.ewi~c ongmaung, probably 'ri Junin
(mc:!>ench}me) bt>t\."een the endoderm and the ecro- era! tJme~ over, frolll .1 piJuula-type ancestor. Th Wi
deml (ftg. 3 ~b) I he endodenn forms the gut. the probably had ~obd rnc~enchymc and may well h~1 Jfl..' s
ecrodenn tht: outer \kin In all other invertebrates, resembled ltvmg platyhelminth flatworm!>. Th lhc I
a' well a' 111 the \.Crtd1rate~, the mesodem1 form~ a gave ri~e. agam pcrhap~ polyphyletically, to 11! 'W!lfll
hmng to rhc cctodcnn .md overltes the cndodcm1al coclom:uc~. UJIIC
gut aho, encJo,m~ a fluid-tilled cavity known as the Two coclonutc 'mpcrphyla' arc tradJOon thl' lt
coelom (hg ·' ·k) fh c gut ltes freely within the agreed as dl<;tmcr. on cmbrvolog.c.tl b'TOtmds, tbout, fill Ill\
coelom, and '\lllCC lt mav be looped or coiled it does <;ome aurhonoe<; chum that the apparcm differcn .lllima
lll b, dear dun tht·> might .lppcar Thc'it' are the Many .mthonncs bd1evc that the proto- and
- .I Jlrotostomia (atml'llth, mollmn .md Jrthropods) dcutcrol\tomcs wcrt' dcnvcd tndcp(•ndcntly ITom
Id dlC' Ocuterostomia (t hordatc~. cdunoderms an acoclomate 'pl.ltyhelmmth' Hergstrom (1989.
Jfm nmor phvl:l) In P1 otml01111.1 ( I) the early 1997), however, com1ders that dcutt·rmtomes were
tll e of cdl diVI\lllll (de a\ age), ti>llow, .1 ~piral derived from protosromes by loo,cnmg devclop-
and the 1.1te of t".lt h tell I' dctt:rnunt'd lrom an mcnul comtramr'; rhq: .m: twt really vt•ry fu dltler-
ugc (2) \\hen tht· t•tnbrvo h.ts reached chc ent. H e env1s..tgcs .1 \tt·m ~>roup ot mct.12:0.lll\, about
uiJ Sta~. rt' hl"n'lllt' ~ hollow ball of cells, 1t the platyhelminth brradc, whtlh W:l\ P'l'Udo-
ut~ to li)ttn J tloublt>-wallcd ~phcn: \nth a ~cgmenrcd .111d w1th a true mouth .md JllU\, creep-
ton rnd. llm pu1nt of inv.tgmJUon becomes ing along on a nltated lowmotory \Olc. These,
nouth (prottl\tomv): (J) the tOclom ames by the Phylum Procodom.tta, 111hJb1tcd the IJtcr
• or I~ , u:. b~ tlw mt'sodcnu splitting to pro- Proterozo1c shJUow \l'a', thl'lr ttrval \tagc bc:ing a
~ u domlt ea\ lt\ ••111d ( 4) the swimming, cih- trochophorc.
llnv,l h l rrod1ophme. In the Dt'utt'rostomia on They gave ril\e din:ctly to Mollusc,1 whiCh retain
'her lund {I) t bv.:~ge follows J r.tdtal pattem many prorostom1an character; .u1d v.triously to the
Ill,' rm• of t',\th t:dl l'l:lll.Hil\ flt'xiblt' for a long many phyla that djfft'rcntiated during the C.unbri.m
(~ tlK pmnr nt lllV:IJ!;lllatlon 111 the blastula radiation. T o mmoriids and machaeridians could
lShlp •'Ill 'tbt• ,111\1\ .1nJ the mouth devdop~ separately be prococlomate~ th:Jt dcvclop~J cxtc:rn~ll ·,cak~'
to~~~~ 1htomv); (3) tht• rnelom Jrist·s bv euterocody, and survived Ill to the PJiacozoic. WiUmer ( 1990)
• ot HJV;ijllllJtt•d poud1c~ lrum the uJtestuu:; and (4) presents a ,hghtJy dtiTcrcnL view 111 her exhaustive
tit t r;a L a 'tonun.1 review of in vertebrate relatiomh1p~. t'nvt~.l~oing a
hn\~ \ 111 et\ 'f\1.11 Vlt'\\ nf the use ot larvae Js indi- great prolilcrat ion ot" coclomatcs 111 the early
t'Olll 1 r IJnnmlup. hClwc:vcr, ha' been put tor- Cambnan, mdcpeudently dc: n vcd from acoelomate
'\\ 1lham,on (1'192). R..Jdtall~ symmetrically 'platyhelminths' and. although ~uperphyl,t such .lS
p J't' nu adult\ luw bt!Jtc:r.lll) \) mml·trical lar- the protostome lophophorates tan be d!\Unguished,
r-kss ~1 should thl' be: \IJ? Other m congrult1cs the overall impres~1011 IS ant of dramatic polyphyly.
• Tgllll ~~ and Jdult\ 1bound Wtlliamsou the majonry of metJlO.tn phyb bt•mg dcn\·cd
c:-hkc ully f'(omhzcd c~"' hclon~ng to asodJans mdependemly from an JIKC,tral 'tot k. The quite
• :ts:un 11 \\1th rhl• 'pcnn c,f ec:hmoids: the\e are remarkable c:bvcruty of the M1ddlc Cambrian
'erte- 1l} rdm·J j,'TOIIP' )mm· of the: se matured Burgess Shale. Unmh Columb1a (Chapter 12).
1 nvcd e appart·ntly tlllnn.tl \l'.l-urc:hin l.trvat: whtch includes m any arum .lb unposs1blc to p!Jcc in
prob- nd a fe.\\ 'urv1wd ,1, .tdult se.t urchins. modem groups, lends fi.1rrhcr ~upporr to a poly-
1ng tl 1 nrnenl\ kd Wtllt.Hll\on to conclude that p hyletic ongiu for the coclomJtC'i.
C lllCISt I trm< 3n bl· rnn,fi:rrt·d 11-om one aruma1 to The adYcnr of molecular gcncucs on the one
huntl~ rdm·d tll\t' hy cro~\-fertihzanon'. l f hand and dad1mc methodology on the other h,\5
~t'fl' et! nJ ~,J l'll''1blt·. through narur.11ly occurnng revivified the :~ge-oid debate on metazoan relatton-
n h.nc ht1n, tor ont· J...md nf manne mvertebrate shtps. While much of ch1s IS beyond the scope of the
( 11 ~\)hd l~· th~ i.ln .11 tiu tn of .111othc:r, .md thar present text, the to llowing works giVC somcthmg of
tdcpcu- tHll~~~~~ 1111111 ot dttli.•rt'nt 1-imh of o rganisms the flavour of the m odern debate. R.tlfs (1496)
PL"'Jbh I ><UIHld in tht• pJ~L. then the importance synthesis of pa laeontol ogic:~ l dam with developmen-
.K0c(,,. J1 pl1ylo~lllt'tlc: 111thcators 1s much tal genetics puts special ~..·mph.1s is on the role of
,) I ~('\ .. I I.J regularory genes. Likew1sc Davtdson er 11/. (1995)
·• { flll 11cf!lu rht• Jcutc:rmromt'\ a \t'p.lrate 'lophophor- explore the o n gin of b1laccral body plans in renns of
I IU\C pc•ph\ lum t'.lll he dhtm~;tmhed, mcludmg developmental rcgubcory mechantstm Gee's ( 1496)
Th~e lltath10p11dJ 1nJ Bryo:m.t, and the small analysis of the origm<> of dcutcrostomes (echino-
:o thl ' ln11\\ n .h l'homn 1d.1. The~e ~] h.we the dcnm and chordate\) rchc'o ~rrcatl) on cbdtstic
bnd ut' fi,od-br.lllll'nng .lppJrJtlt\, known .15 mechod'i. as does Nielo;cn·~ (19lJ1) \urvey of rcla-
tonllly lophophore. Nont· of thc\l' phyla arc ~eg nonshtp~ w1tlun the ammal kmgdom Moore and
:hOltgh b t tltt• roultl h.l\ c hl·cn denwd from an Willrncr ( 1997). on the other h.tnd. bellevl.' conver-
~re ne gent evolunon to be so w1de~prcad th.lt dJdJStK
Tnlobite' i, for example, whteh have been t•xn In

analysts ba~ed on pan.unon y (the choiCe of the sim-
for some 250 Ma, were undoubte dly htghly fun gnHt
plest cladogram ) will ntinimizc: and :~ctually conceal tl
converge m·e, kadmg to false results. They believe tiona) inverccbrateS, as is witnCS~l·d by the comp 111
morpholo gy of their eyes and other envirorun c Tht·

that the evaluano n of morpholo gtcal character .,
sensor;. ln a \Ct1Se they could be described as 'pn1 llld
roupled w1th mokt ular data, is the best way for-
w.ud. The debate w1ll C'-'ldcntly continue for a very itive', ince their appendag es arc undiffcrentlat ubj!
long time. and they lack some of the claborau om of o I liVe
arthropo d groups. The term 'prim.inv c', bO\.WH le-. c.

lt ts pmsible th,H the carlte~t mctazom s ltlhabited
should not be constmed as meaning biOlogic thct
the meioben thos, that 1s the poorly m:ygenat ed
infenor or ill-.tdapte d. for though m an t·voluoor rrcd
~ulphide-nLh la) er JUSt below the surtace of rn;my
~eme they were unable to c~cape from the lmuu prul
marine ~ediments. Whereas the meiobcn thos today
tiom of being tnlobite~ they '"'ere able to J.dJpt llld
h,ts been tll\ aded b\ some ~econdanly adapted ben-
rhos. at lca't some of 1ts present-d a) ~mall organi~m~ many 'peciahze d niche~ and .,.. ere amongst the r IOl:l.
succc~<;fu llife forms of their time. Ill cl

(protistam and mct,tzoans) might well be the relic~
of original Precambr ian mt·tazoan fauna, which 1n trilobite~. as in all of the invertebr ate f, Clltl
ongmate d 111 the mlphide- nch layer., and some of groups studied here. once the themanc pancn.
which invaded the oxygcn,lt ed layc:n. above as the orgamzat ion of each fctxon wa~ estabhshe d it tem'
progenito rs of the later rmc bentho~. l f this were the to remam very couserv.1tive. Thus, Wlthm any ph c
lum each hierarchic al level of orgamzan on can
C<ht', then the earl) e,•olutton of \tnall metazoan s
tmght wdl have bct,'Un long before the beginnm g of rclatc:d to an archetypa l pattern of comtrucn on
heritage which though highly t'imctiona l wa\ Jll I )u
"
rht· Cambrun . Unfomm atc:h such meiobcntho~ i~ hh
mmt unlll·..dy to be fossilized and elm we shall never same ume confining and restncrive . Every ar
, on•
know. typal pattem, however much cvolunon ary phl\tl(
was possible within it, remained an archl•typc for lwcJ
the phylum. At t<L\:ononuc levels lowlr rhJJ1 fccu
phylum, new kind\ of aninuh could wmcur ltllll
originate , brc:tking away from chc fimcttonaJ \}'t I 1tt

of che1r ancestol"\ but unmed1a tely 'il'ttmg up 'iOUI
archetype s, which were likt·wi~c continin~ p.llll

!'ill y
which allowed, within certaiJt !tmits, aU rn<Umrr
Diversif ication of inverte brate l ife b:t~~
new cvolution :lry po'isibilities
Once the ma.Jor ~rr0t1p~ of Invertebr ates were cstab- Th<.• molluKs, one of the most drvcrse uf spcl
very well how the potc llllrt
li,hcd t!H.') were .tble to expand .111d differenti ate, invertebr ate phyla, show
single archetypa l plan \\as reah IIIUI
though chctr rclanve dommanu~ has fluctuated uaJ mhercnt 111 a
lu~c· (see Ftg. R.l) h lJIII(
stgnilic:mtly though rime. The great CJ mbri.m A hypothen cal 'archimol
covering the mJs~ of vim:ra. a rJ\P
'explosio n' gave n~c ro many groups which were single shell
and ,1 postertor C:lVtty 111 wlu~.h rhr
'hort-hvt :d. but chmc phy!J wh1ch survwcd the mouth
located. From th1s Jt lea~t 'IX ne\\
Cambnan h.wc continue d to the present d.ty. Ta.xa were
time ranges. independ ent brroups emerged. Some, \llch a~ 1~
of lesser rank havl' gencr,illv shorter
to work out monopla cophoran s and polypl.1c ophoram. rem ellet
From most group' 1t has been pomblc Pc r1
thci1 phylogen y. dq>cndin g quite dose to the ancestral type; other.
~ome of the dct.u), of
h and mo..,l of Jll t lo\
on how l omplcrt• thc:ir l(,s,iJ record is. To some.: bivalves, gastmpod s, scaphopm
away m vmum dq· I Ill t
cxcem, through fimniona l 'tudies, lt I'> aho pm~ible cephalop ods - diverged
adapted to va.nous habitat\ .mJ mv tnd~
to Ulldt'r\l.llld SOillt'tlting of the bwJogica) quality Ol and became
life. Each of these classes had an archei\ rod:
the fossil organi~m~ even tl these arc of great .mtiq- of I' !I
'masrer plan' in whtch all the ft•arurc' of thl' u
urty, and 1t 1s vc 1'\ dear that even from the begin-
molluK were present. though mod11ird , addld f\
ning of tlw fos,il record mvertebr. lte hte WJs highly
ol the later and altered in a parucular functiona l combiuatH
orga1tized Jnd wdl ,1dapwJ. even if some
ca~c a form fi)r new cvoluoo
represent atives ol ,1 'rock apparentl y 'tmprovc d' on allowmg in each
the earlier ones. potcntul.
Major features of the Phanerozoic record 69
In the mollusc~ and 111 most other mvertebratc trilobites m the OrdovlCJ,ll1 and cnnmd' m the
lUJll the mitial djfft·rcnnauon took place very late Carbon iferous). Conversely, however. the number
·~ tht• Prcrambri:m or 111 the earliest Phanerozoic. of body plans or defined groups in the P::Jiaeozoil
.tl n, mhcst evolutionary ~tages are not preserved w:u. much higher. There arc fewer kinds of alllmals.
l- 1ldc:1n nnl) be rcconmucted by mferem:e and with i.e. groups of high taxononuc rank. today than there
·d C<:U\'It) Likewtse. though the fonmuon of ne\v were fom1crly, and this net reduction m typc<i of
;:r rnebme archervpe~ went on at lower taxononuc o rgaruc dc~ign is a mamfcst trend m the fo~'il
ds throughout the whole of tht- Phanerozoic, record. The highly succe~sfi.1l groups of today,
l ere b nomtally litde n·cord of the types that car- which include so m.my \pccies, seem to be
d the potential for funhcr development. This 1s 'winnowed survivors' of w hat Gould h.1s termed
pro~Jbl} because most of them lived in very small 'early' experimentation and later ~t:mdardizarion'
l' J r.1p1dl) evolving popul.1t10m contined to very The early experimental woups, ~uch ::1!. so manv
·~t :-alizcd aa\IS. The t h.ull"C~ of the1r preservation Cambrian Problematica, the Burgess Shale f.1unas
diX"O\'el) arc thndorc \'cry small. and such and some of the more b1zarrc lower P.ll.it·o7mc
il uall) m1portant 111temled1ates are rardy found. echinoderms, did not b~t long, but to \.\'hat extend
of it was comperirion from other groups that weeded
:J them out, or purely randon1 phy~ical processe~. mur.t
Changes in species dive r s ity and remain conjectural.
c habitat Not only has there bc1.'ll a 'hift in diwNI) and
a biomJ.)S through time. but there hc1.~ al~o bt•cn .m
1<! m1'rc ~pccie~ hve 110\V than in fomlt:r t1mcs? Or overall d1splaccmcnt in habitat. In the earliest
e- the number of speoes remained more or lcs\ Palaeozmc the immobile su~pcnsion feeder; (e.g.
n ,n,t.Jm \Jncc, say, the late Cambnan? Thi!> has not brachiopods, dendroids, bryozoans, corals, ,trthaco-
311 l~n Jll CJS)' problem to solve because of the impcr- cyathids and some kinds of cchinodenm) occupi~.·d
ne ntinm and bias inherent m the fossil record, and soft sediment surfaces. Throughout time, however,
onl now there has been no clear agreement. the soft sediment environment came to be occupied
n1 ttcrly. however, much d,ua deriving from various by depom and infaunal o;uspemion feeder' o;tu: h a~
w uM' has been pooled and show~ a conSIStent protobranch bivalvcs, Irregular echin md,, ,ome
llt mem of change - a real mcrease in specie' <.hver- crustaceans, holothunam and wonns. as wdl a~ a
of Lhmugh time (Sepkoslu et al., 19R 1). Thts is number of~ovengers and predators (Thaycr, 1979).
t ·J upon aua.lysis of (I) trace fossil diver~lty. (2) The great increase of such ~ediment-dmurbing
~·cdc~ per million yc·.lr,, (3) species nchncss, i.e. ('bulldozing') animals rendered the soft scduncnt
mm1bc~ of invertebrate ~pt·cic'S withm to~sil Wlll- surface less habitable to 1ts former 1mnwbilc
umtlc\ through time, (-I) generic divemt) through dcnizcm, and those that do survive (endoby\!>atc
me 2nd '5 diveNt) of t:lmtbes through time. bivalve\ ,\nd stalked cnnmds) tend to tavour rcla-
~11 five C\tinlJtc~. \t,lti\Ocally tested, showed a tivdy unb10turbated sedtmcnt There wa~. through
~{underlying pattt·m l nw diversity prevailed in time, a corresponding mcrca~c in tht' dtver.Jty ot
c C.Hnbrian, and although diversity was apprec1a- mobile taxa in sofi: sediments, and of mm1obue su~
~ luf\hcr Juring the n·~t of the Palaeozoic it lev- pcnsion feeders on hard st1rtaces. Some uutlally
.lltd off and did not go nn mcreJsing. After the late inuuobilc taxa, such as the cnnoicls, have mob1lc
mmn cxtmctiom thl·re was an imttal penod of rcprc~cntatives today. It~~ qune possible that the me
1 d1~c:t\ity, follmH·d br a real and connnuous of rt·ef-fomnng and dwcllmg animals 1s Jl lca~t
t"l)C in diveNity unul the present day fhcre are, panly due to the d~trucnnn of the origmal habit,lt
cJ. at leJst twice thl· number of speCie~ hvmg m by bioturbation.
JJ) \ocean~ as tht·rl' were at any tJme dunng the
al•rozoic.
Modem seas abound 111 gastropods. bwo:l!ves. Proble matic early Palaeozoic fossils
ll, ~>, ~dunmds and fishc,. In each of the~c modem
ry !llp\ the di\·ersiry of spl'l'le' cxcccd' that attamed The c,1rl y Cambrian tauna' represent a nuJnr meta-
an' Pa.lacozo1c phylum (other. perhaps. than zoan rad1at1on. Along with the 'standard' tnlobltc<.,
mollu~cs, marttcul.lte brachwpod~ and others that

were the progemtol"\ of long-hwd stocks, there
were many short-hved. oftc..•n gcogr:tphteally local-
ized and frcqucnd) 011arre fonm which cannot be
related to an} other group,, but \\ hteh mmt be clas-
Sified as exnnct phyl.1. lt \\.I' .1 wne dunng which
many mdcpendent hneagc..·, on~natcd. but only
some of\\ hi eh ~un.·lved (St.mk}. 1976), for Js prc-
viomly notc..·d there \\c..· re m.ln) more body pl.m~ in
the earl) P.d.le02'0lc.. than there.lfter.
The rapid d!veNtir.ltmn of mer.1zoans around the
beginmng of the C .unbn.111 wa' a real evolutionary
phenomc..·non, .md the .1ppc..•arancc of mineralized
exoskdetons w.1s onlv one ,1\pen of thts diversifica-
tion (Stanlcy, I 'J7(J; Beugtson, 1977). There was
surely an equivaknt rac..ll::uion of \oft-bodied animals lbl
at phylu111 level am.!. .tlthough mmt traces of these
have been lost, there..· .m: indeed a few isolated
occurrenn:s. For c..·x.uuplc, \Oil le curiou~ soft-bodied
pent:lmeral orgam'm' up to I 0 cm 111 diameter and
known on!-. trom .1 \tnglc bedding plane in the
Scottish Htghland~ (CJmpbdl and Paul. l983) may
have echmodenn alluuuc..-s. but .m: more likely ro V
belong to an unt...no\\ n ph)lum. Sun.ilarly. organ-
ISms which formed tl1c1r \hells by Jg,_glunnatmg par- Figure 3. 6 Tl
rides together arc found tor the tim time in the Paloeozoic e.f.,
Lower Cambnan.
The genu\ ,\altne/1,,, for example, of Lower
Cambnan .1gc. " .1 'mall. comcal fo~s1l w1th 1ts wall~ Marine
divergmg ar ~01111.: 211' and with ~ centr,ll tapering
bore. The walls ronst~t ot ~mall .tgglutinated grains Tht• Ed1.1C<
arranged m nc\tt•d contcal layers. each of which T lmHltonan
makes an angle of about 45" to the a.Xls. Salterella L'¥ .nh1ds rec<
bears a supertic 1.11 resc..·mbl.lllce eo cephalopods Figure 3.5 Mobergello, on enigmatic Lower Combrion foJ ctf marine h
with wh1ch tt was ongmally confl1sed, but it is From Sweden: (a) young stage with convex inner surface, l .tmbnan o
older example with concave inner surfoce (x 15). (Photogr~
now referred eo the cxtmct, \hort-lived phylum reproduced courtesy of Or Stefan Bengtson.)
tlf \UC'C..CsSiVt
Agmata (Yochclson, llJ77, l9H3). Salterclla is wide- 'ncs ofcxti

spread. ranging thrnugh much of North America, Tlm Phancr
and is found .1lso m the Northwest I lighlands of ~lll'C eSSIVC '
Scotland. cure, protected by the..· ~hdl. Suull'ipecimens of\uc l9'ltl) The
Many of the 'srmll ,belly fomh' (F1g. .3.2) from a shell could nor h.wc .1fforded .my protection to 1 Modern CVl
the Tommouan are of problcmauc origin. Some bearer unles-. it w,,, :1 burrower. ltkewise, the..· \Ill IIIC:nts ~een
may well h.we been pam of larger organisms. button-\haped 1.-~·,a~l?)lrian, ~otm· 0.41 111111 111 J1.111 ~l••v.ly togct
Bengrson (196H. I 977; Q1.ln and Bengtson, 1989). eter and consio;ung of a dc..·mt· tuberculated !Jpp loo F .\l h c'
For example, tlw round, di,coJd.tJ 'hell .\lobc~~ella, over .1 porou' con:, " of unknown afii.mt}. Mig srmplv com
known trom Scandtn.lVIJ, came\ mme seven pan; the\e have bel.'n dermal scleri tcs- J kind ofbodl :!JC cflt.•diVC
of marking; (wh1ch m.ty be either elevaoom or annour of an exunl£ ~rroup? For the moment. 11 I he Can
depreSSIOnS) in lt\ COilt'JVe 11111er ~urface (fig. 3.5). pcrhap~ inddinncly, our I...nowlcd~e of man} by rnJ,)bltc~
The,c.: mav h.wc bt•t·n mmck attachments, and thc\e b1zarre, e.1rly ( ambn.m 'Problematica' m h) <thth llh,
one nught envisage tltt• anmul as J limpet-like crea- rcmam ramah71n~l> tmompletc. denm. and
ligr.te 3.6 The three successive, but overlapping morine evolutionary faunas (e.f.) ol the Poloeozoic. Combrion e.l., block;
llbozOK ef., crosses, Modern e.f., cleor. (Based on Sepkoski, 1990.)
Marine evolutionary faunas (Fig. 3.6) Burgess Shale animals belong here too. This evolu-
tionary fauna divc~ified rapidly .md its members
~ Educara !tuna of 'v~:ndo7oJm' .1nd rhe were generalized depom feeders and grazers. le
mmotJan tJuna of \mall ~hdly fc.mil~ and .uchaeo- began to decline m the Ordovician and became pro-
lhids record tltstmct early q11~odes in the h1story gressively restricted to deep-water environments.
il Mmne li1e F·olluwmg thts, and from the early By the Permian it had aU but vanished, apart from
l) umbn.m onw.uds. there ha' bn·n a contmual relay inartH.ubted brachiopods and monoplacophor am
wcc~sl\'t' invertebrate faun,\\, punctuated by a which ~urvive until today.
ofe:o:ttncoon penods of grc.tter or lesser etli:-ct. Meanwhile the Palaeozoic cvolunonary fauna
'!1111 PhaneroZOIC record can be rc,olvcd m to three had arisen with an 'explosion' of calcareous-shell ed
Ct'\11\'C 'evolutionary f<llln,Js' (Scpkosk1, 1984, invertebrates in the early Ordovician. Rhyncho-
b ). Thes~ are the Cambn.m, PJ.lacozmc .md nelhfonn brachiopods, corals, moUu~cs, bryozoans
lcdtm evolutional) fauna': their componenr cle- and ednnoderm~ rap1dly djvcrsified, new trilobttc
t> ~cem to divemty rap1dl~ and to dec.hne families arose and graptolites with coll.lgen skcletom
Ill) 10gethcr, and they share orhcr charactensncs dommated the plankton. Much of the bcnthos
l •ol. Exh evoluuonary tauna is not seen tlll'reforc as consisted of epif:ltlnal suspension feeders, with
1ph composed of randomly .tssociated tax.t, but more complex levd'> of ecological tiering than the
.-ctively linked m some \\ .1y. Cambnan faunas had -.hown. The Ordovician radia-
The Cambnan evolutionary 1:1una is donunated tion of tlus evolutionary fauna established the pat-
tnlohaes, along \vith martJCulated brachwpods, rem of mvertebratc hfe on the continental shelves at
.uhthJd), monopbcophora ns .lJH.l early echino- least until the end of the Pcm1ian (despite sharp
ll. and tht· ~oft-bod1ed or organic \lH~llcd changes at the end of the O rdovician and in the late
De'>'Otuan) . Ahhou~h th~ V.Jb~ozoic faw1a was number' and dtver\tty and con.tined to only ccrtJu1 t:frect' 01
~everdy ,Jilec:tc:d by the end-Penman extmction parts of the world, finally dymg out at the end oftht \\ <llllllllg
en·nt. 111.111\ reprc,c.-ntattve tiumlies surv1ved Cretat.eou~. m:w m.u
throu~h tht: M~'o7mt to the present day. When, followmg the tinal denuse of t!Y. OIZ ltJnn.
Th~: Modt:rn or Me,ozoll-Cenozoic evolutlon- ammomtes, the carllc\t T ertl.ll) i:1t1nas were estab- ph.l\l" tv
ary t.1una of bt\>Jhes. arthropods. echmoids. lished rhey were not unhkc those of today. Tht calli\ wh
foramimtcritl~. cnt\t,tc.:eam. fiSh and other groups bard-shelled faunal assemblagcl> m the Tertiary con- en, is
had \Ome rc:prl'Wnt.tttvc' ~omg back to the SISt of b1valves, gastropods. echmodenm, bryozoaru. Upon
Cambnan 1nd h.1d 'lowly dn·eNtied through the sclcracantan corals and other fom1s. \Vlth br.. h<nt~e' 111
Palaeozotc Wlwn. tollowmg the Pernuan evenr, cluopods becommg a very nunor component progres~l'
\ acant eco,pan.· b~ tame ,watbble again on the con- · Indeed sheiJ debm on an early Ternary shordm order cy
tment.ll \hdn·~ tt WJ' the Modern evolutionary would not, m terms of groups represented, be veq 1'~'12) . T
taunJ whtch took lWt'r !'here are many evident ditferent from that on a modem beach. 1t would rh~·,e va~
change~ at rht' rtlllt', tor c:x.unplc, the braduopods contrast s1ngularly w1th a Mc..·~ozotc beach, whtrh p.1rt of tl
losing thctr domuunct• to btvJ.Ives, ;md the com- would be covered by ammonite shell,. ht(· hJ~ b(
plete replacemt•m l"lf corJI and edunoderm fuunas.
Very divef\t' ecolngtc.al types :m~ presenr amongst
repre\entanve~ l)f tht· Modem tauna. Climatic and sea- level changes Ex tinc
The: rhree p;rT.tt L'voluttonary fJunas overlapped
but they .tre di,ttnct .111d they provtde a meful way The Earth's climate ha' drarnJtiLally fluctuated dur· 11 \Ct'lll~
of unde~randtn~ rlw c:xp.1m1on of the biosphere. ing the Phanerozoic, n\utl.t.ting between altcmat evc:nluall •
One duracrt·mnc ft'.ltltrt· that c:mc:rge~ clearly is that '1cc:housc' and 'grcenhou,e' conditions (e.g. Frak(1 IWW t.l>.;l
each ~ucct·s~ivt.' evolutionary tJuna dtverstfied more et al., 1992). The late Precambn.1n (Varangtan) t('( llld dl\ap
\lowly rlun rh at \\ hu:h prt•ct·ded 1t. but has a higher age gave way to a warmer. though still cool ciJmatt ltcd gcol(
kvd of nuximum dl\ t'f\ity. and .1 grl'ater range of in the early Cambnan. By the late C.1mbnan lht lc,·d o(b,
ecologtcal \tr.uegte\ too The- <.tepwise pattern of Earth had entered a hot 'b'T'Cenhome' phase. bstu~ lllllUllll.'l,t
incre.he in global diWT\It)' \een m Fig. 3.6 has led to until the end of the Oevoman and punctuare-J on!] epl\Ude\,
T\vice a\ m.my ftmtl1c' living today than there were by the \hart. sharp. late Ordovician gl..tt:iaoon. Ther. exnncl n
ac an> nmc dunug the P.1heozmc. Parillcl~ may also from the Carbon.iferom to the late Triassic there" ,1Je CXtll
be \Ought in the cxp.tmion of tcrre~tnal .md fresh- another 'icehouse' wtth extc.-nsi ve tee sheets covet· lng~c.tl p
water pbnh .1nd .mim.ll' (C.r:1y. 19R7). mg rhe sou them cononent of Gondwana during th. tf, tined o
Oe,pitt' thl.' rnht•rt•nn· of t'ach of the three evolu- late Carbomferous and Penman. A warnung ep1~od. example.
tionary t:mn.,,, t'.lch h.l\ umkrgonc ~ub~tanrial inter- in the late TnassK led to a ~econd 'greenhow f:luna., tlu
naJ mmowr. Thi' ts \\'l'll 1llustrJtcd by the Modern phase, pealdng 111 the Cretaceous, .111d therea!tl'! tuH.Icrwc1
f.uma. ,mcl 1.''-pl.'tt.llly by thl' me .md tall of the there was a general dcclme m temperature, leadmt t.umhc' ~
ammonoid\. Wlwrea\ thL· rise of the ammonoids in to the present 'ice house'. ··nt1rcly n
the Dt·vontarl \\:t~ \lnw, by the: Triassic they had Such large-scale c1Jmat1c fluctuations (\Ill !J, 1'1X<:I). Tl
become amongst the mmt common and characteris- related to changes 111 the mtenul workings nf th g.r.tplucaJ
tic of all prcst•rvablt: lltVL'rtebratt'~ and very diverse. Earth. ' lcchouse' phases occur when plate tectom fi1r 1t. Son
Y t:t they too Ill.'.HI) bl'Ldme extinct again at the end movements arc nunwlJJ. At such times ocean ndw pl.llt!, tho
of the Tri;mtt. ltwing ,1lrc.1dy suffered a late subside, thereby ac.:commodatmg v.1st amounts o Such e
Penm.1u ~ethJl k. All thl· .tmrnonites of the Jurassic water in the ocean basins, so that the c.:ont1 dtt1~rcnt ~
and Crcratt'Ull' wen: tkn.,ed tTom a single small masses arc left high and dry. Lowered vulcaninr e~:osystem
group of Lttl.' T riassl( <UIIIIIOnmd.' which alone sur- tn.inimizes atmosphenc CO,. and heat loss mcreaso ng1d m a
vived rhc: n~OUT\ of chc end of tht· Tri,mic. With rhe onset of renewed plate tectomc acnvtl) prospect '
Ammonitl'S liourishcd 111 the Mesozoic, along contmcnrs move, spilt and coUtdc, ocean ridges 'u~h an e~
\\1th b1valvcs. gJ~tropmk cchmodem1s, corals and contments are Hooded, and mcreascd vu) rotl•ntial
braduopod~. '' hich were o;till locally common. In throws dusr and CO, m to the atmosphere so f{'placcme
late Crctat·eous nme the Jmmomtc' declinecl Over hear 1s kepr 1n. I hus begm~ a ne\v 'greenh rronusing
a pcrtod of \OIIll' Ill Ma the> hn.11m• reduced 111 phase. Such megacyciJot)· mev1tably has I hus ace
cl\ on living organi,Jil\, \llltC wht·n dm· to global Phancrolou.: lo~sil recorJ c.an be dtvtdcd into 12
u1mn~ the contiiH.'tll.ll 'hdvc' .m: flnodcd, v.1~1 levd-botrom 'ecologicll-c.·volutmn .lry. Ulllt\ rum-
\\ mannt habitat .lrl'.l~ ,1ft' •lpcncd up rm { olo- tuatcd by extinction evcnh of" a t,.rrcater or le~~cr
lllllll. These dl~appt•.tr dunng a later 'ltt•house' m..1~nitude. Within each of the'c units. commumne'
1c. Moreon:r. clmutll' ch.mgc hy tt,dt. l'spe- wen:- n:m.1rkably \t,tbk and thctr 'tmcturc.· w,l\
U) \lht'n rapid, i' hkdy m gt'tllT.ltt' 1 lm-.lo~cal n~hth comtrained \litthm t'c.ulo!Pt<lll)- dict.ttt·d hm-
lts. Dunn~ rhe~e long pcnOJ\ of stability thcrt• \\ c.•rt•
Uron tht:'e two 'grct•nhouse • and thrt't' 'tce- fev. cvulunonary mnovatiOih and stabililtll~ ,del-
u,c' mqtJtvdes ut" the Phancrozott ,lrt• unpmcd non operated both at the populanon and the c. om-
••gJC~'ivdy sbortet ~ewml-, third- ,tml fi.>urth- muniry level. When, however, major cxtimtiun
k rycb gl.'tlcr.lll'd h} other c.Ht\t'S (liallam, event~ dl\1llpted the \tnJLturc uf cco'y'tcm~. not
2). Tbt'se too h.1d tht·lr bH.>logiol dft•us .md onl) were new structural and phystological mnova-
e 1 Jf)111g cycles, op~.or.ltmg at dilft·n·nt \t'.lk~ • .1rc tiom C\tabl!shed during the ~nb\equem .•J.lptl\'l"
ut dtt' r~dess stJgc upon which tht• lmrory of r.1d1at10m. bur commumty t) pes were complctd)
lu\ hccn acted oul. recomtltuted m new and d11lcrent panem,.
Extinctio n s Possible c auses o f m ass extinctions
1Wns to be the f.ltt' of VlrtuJU\ aU urga•11sms Of all the. t•xtmction pcnods wh~eh haYc heen Jot-
~:ntually ro become cxnncr. In the ,,tme \\,1} char umcnted, by f:lr the mmt 'evt•re wJs thl" l.ltt'
w t:txa h 1·e a me.: n thrnugh nmc 'o they decline Pcnm.m cn\t\, which reduced the numl:-tt' l of
Jd ,ii,lppear and mmr gt'llt'r,l .md specie, h.tvc hm- mannc lllVl'rtcbratc famthcs by 57%, (with perh.1p'
. d gculogJcal rangt•s. Thcrt' ,, thm ~ continuous IJ5'(n of .tJJ ~pccJeS disappe,mng).
vd ufbac4,tround taun.1l rhan~eover, t omistmg of Thcrt· were other tmcs of intcm1edjatc: severity.
uncrablr· minor t',tlncnon and n.:platemcnt Up to 22°111 of all r:~mihe\ dit•d out in the. l.lte
node,. Somt:tilllt''· however. nuny tJX.l become Ordovtu.m (end-A,hgilhan) .•md cri~es of comp.lr.t-
t11cr more or k"' 'tmult.tncousl). Such l.trger- ble magnitude took plan· in the late Devnnun
ntmc 1011 cpt,odt•, tenninate L'.\Ch of the gco- (Fr.NH.m-F.unennian : 21° o) .md late f'na~st c
:11 penods and the 'Y'tt•ms tht·m,clve' are (Non.m, 20'Y.1), and fin.11ly 1n the late Crctat:t'otl\
d1111 J on the b:~s" on tlw taun,t~ they cont.un. For (M:~.tstnchti,m; 15%). The la~Hncntionl."d ofthc.'\l' t'
or
•lllple, at the elld the C.unbri.m tht• ttilobite of particular interest .1s it ,1tTccted land anilll;ll\ .1\
•u1> tlut had dormn.Jtt•d the scene fiu .1 long tlme well "~ manue fauna'>. None of the\c cpt,odt''· nor
ulmvcnt a worldwide change. rhe m.lJnnry of the le'' unportant cxtmLt1on cvcnts down to tht•
tilit'\ belOming cxtinet .mt.l being rt·pbccd by b1omere level, were part of a regular cycle of elit'tlt..,
nn'l~ ne\\ scod..s (Wiutungton. 19oh, Wt'\trop, but were Irregularly spaccd throughout geologtc.tl
%'9) Tht' abrupt t h.mgt• j, a cnnvemt·nt \trJti - nme
11ph1lll marker, but therc must have bl'l'll reasom The v1cw that extmctlOII events are cuupkJ wirh
'Ill ~~)lllC kind of t.lt.l\trophc \l'Cill~ to h.tVt' takt· n a 26 M .1 pcnodKity (whtlh would suggest a s111gle
r, though not netl'\\,lril} .1 sudden l>lll'. underlying causal mechani\lll, and pmb.1bly
~ uch events are oln wusly dc,tructlve, but 111 a cxtr.tterresmal) ha5 now lather fallen !Tom I.IVIlllr
tTercnt ,ct•,e they art: ,1l'o ue,mve For once new (Pauerson and Smith. I <)~7, McGhee, 19R9a,b).
r ) 'tenb are ~et up they tend w bcconH: r.uher What 1s dtfricult, and rathcr more spec.ulatt\ c, ,, ro
·!i!d 111 an evolutionary 'emc. Thert· i' not much find c,m,cs. Smce Alvarcz et al. (lt)tW) propo,ed rh:lt
101r~cr wichi11 them I or umm ativc change. Once the.· renmnal Cretaceous cxtmction evt:nt n:,ulr\:d
·. ,,,h .m ecosystem h,l\ broken down, howeVl'r, tht: !Tom an asteroid colliding with the Earth . III,ISs-
) !'l~tenn.tl then become:-' .1v.ulablc for upponunmjc e~:tinction analysis ha~ become ~omething of .1
cpl.ltr~llent and 1t ,, at \uch nme' that n~.o \\ and grm\th mdustry in palaeoumlogy (Silver and 5thul?,
m:,mg evolutionary development' c.111 prmpcr. 19R2; Llf\vood. 1988; Albntton, 19~9; Chalonc;r and
u1 Kcording to Uoucor (1983) the marine H.11lam. 19H9; Donovan, ll)84, Alvarez, li.J97).
74 Motor events in the history of life
Whcrc.l\ tlwrt• lw, been .1 nJtural. if mis~idcd Volcamsm Thl.' produltton ot mas~cs of vokanK l'llJllll
tcnJcnC) to look t(,r a common cau~e tor all extinc- du~t mto the .ltlllO\phcrc would undoubtedly rt....ul: rclurd
non t vtnt,, m.mv aurhontte' would prcti::r to seck m glob,u coolmg .md rc,ull 111 cnvumunental ~trt ~h.: mt
l'Xplanauom 111 nndnm combmatiOllS of more ordJ- whtch could bt.• long t·ontmucd. e nnt: t
IlJf) tJctor' (T1.•1d1l'rt, JliKI:!). Indeed. a~ Holland Oc~.·JnogrJphic ctfech. P.mcrn\ of oceamc nrctr rcsuh\.'<
( Jl)Ht}) h,l\ n~htl y pu111te,! nut. the u'e of the term !anon m.1y d1an~c throu~h ttt1le. ne.:\-\" system\
'ma'" exum ttc.>n ' c.· an nvadr.unanze the pattent" that up\\ellmg c.:ould bc \Ct up .md pO\\Ibly l.ugl·-sc.~ late
rhe t"O ..,il rcnmi .ll:tu.1lh' hJ' t0 'h0\\ . overturn of stagnant buttom \\.Jtcr could Olct: I f•\hlr
Putt.' tHIJI ~.: .llt~t'' c.1f the dc.·tcriorJtion of glob.u wnh lethal c.:tlen' on the pd.th'1c.: and 'helf tJuna <. high, [
ccmv~rcnh Wc.'rl.' dctcrnum:d by McGhec (l «:J~9a,b) the other hand, a dmunution of upwelling ~y)ter \HrC f
as tollo\\' could, over an extended timc penod. dt•plete clx, bl:ltk g
plankton of essential nutnents. 11h n.
Earthbound mechanisms lllO\t st
Glob:~! remper.1tur1.' dcchm· Tlm would particubrly Extra terrestrial mechanisms r;Hcd
afl"l.-rt ,h.lllow-w.lter trnptol (and rce~ ecosystems. Supernova rJdiattom. I eth.tl bbst' of radJatton fro exp.ltld
Manne org.tni~tm llvin~ ;H h1gher latitudes would a nearby supcrnov:. could wdl have advcm~ Jnd Jr
rnrk mit,rr.um~; tcmpcr.lturc belts and would be kss atfected the E:-~rth. lt i~ not poss1ble to tic any sm' <.'wlogi
severely atTl·cted, ;md hkc:wi'c inhabicanrs of deep cvcnr in with the.· fo~~il record, however, :md m<~ \\tth ••
water. mannc ort,r.mi~m' would have been protected bnr (Bn Ill'
M.mne rc·brres~ion . Hucttut1om in sea level have water. ,jtll'l tl."l
t:tkt·n pl.tc.c.· hundreds of tlme' during rhe Bolide imp:~n. R.trc but dcv;l\tating meteorite' ndont\
Phanero7otc \nnw h.wc.· bn·n no more than mmor :mermd collismm wHh the F.,trth could have toull rh.:re '
loc:~li:zed IIKUT\Iom m rt.'):,'l'C\~IOm of the sea; other~ dJ\rupted tht.· p.Htt.Tn of ltfe. If a bohdc some lO k lu1kcd
on .1 murh lllort.· \\tdc,pn:.1d 'cale. Regre~~iom in diameter htt rlw F:trth\ surt:\Ce, the first dfct.: W.1tcr (
would reduc.·c lubn.1hle art.',\ on continental shdves would be a sh<)< k wavt.· wlth 1mriJI dcva,t.1ttu hr.tchic
{though h.thir.u 'PJCt.' around oceanic 1sbnds would St·cond, there.· would be a pt•nod (lasring ~evr tht re :
IIKICJ\e) . A flnoded l'nntinent:tl ,hdf or epicontl- months) of d:trkcmng b} Jtl1lO\phcnc dmt. Follo\1 worldv
m:nt.ll ~c.\ pr<\\'tdt.•, .1 b'Tt'.lt are.1 of living 'pace <lnd ing the in1ttal hi.N the t.c",Hion of photo,ynthct < hiopo
very many h.thltat,. Thnt: \\ill be especially nun)' :tC!d r.llm .tnd lowered tempcr.1ture' due to \unli§ ( )rdOVI
habttat~ 1! the tcmperatur~.· e,rradielll <Kross the sea is bem~ un.1blc to penetrate the dmt would h.lVe h.;: dl\ t'T\It
h1gh. If the ,hell' St.'J' arc contracted, then the living lctlul efl"l.-ct' on 'ens1tn dv balanced ecosy,tems a tu 25 I
spat.e \"11 dJ\appt.•,tr. uwt~ of the bab1tats will vamsh could h.l\'e prct~pitated J llldJOr extmctton. Then ~nd th
and the ecolob'1c.·al d1sturb.mt.e w1ll be profound. A no doubt that the Earth ha~ been bombarded t \\'Jthm
tal11n sea le\ c.:I of L'lll} a t"l.-'' metre' could do untold cxtraterre~trial bodie<; rhrough time, for there are L I 1 >H~).
dama~c. Wh.u h.wc been termed ·perched faunas' to 100 known impattLraters ofv,triow. sizes on tl I he
. Uohnson. I 'J74), t.e . launa.s tlut have evolved Earth's surfatc whtc.:h have been made during tl:r lll.l:>\1111
raptdly and h.we c.· ulolllzt:d a shcli-or continental sea P hanerozoiC. The 26 km Rte~ crater in ll,, h.lhitJb
dunng a nme of m.1XJilllun floodmg, are especially Schw;iblsche Alb. 1n ~outhern Gennany, one oftb l~<m ex
vulnerablt" when thl." 'C:<1 rctn.:,lt:.. best studied, rs an 1mpact structure some 15 Ma old f hc
furthermore. 1f any 'key ~pectes ·, plant or .tJumal, the meteontc must have been l - 2 km m diam~tc hundre
occupymp. J trlllt.al pos1non 111 a tood web becomes A 70 km drametcr Trias~iL impat:t cr.lter is know I hm,m
exnnct, the whole net\\ ork of fcedmg relationships from Ontario, and m,llly of the lunar impact crate Bv ch1~
dependent upon lt wtU be munediatcly dismpted, are much larger sttll. It has been estimated d1 lcwl re
and those .tmm,tls .u hr~her trophic levels which meteontes of more than l 0 km m dJJmeter, produ&· \',hit h l
depend on n ti.)r food m,l\ well become extmct too. ing enters of 200 km diameter. should occur abolt the Hir
Clmuric hutf~ring effects m1ght lrkew1se vamsh. once every 40 Ma. ,tnd the ltkelthood of 111.11 JUs re-
M:mne regrt:\\tOn would. of course. occur during extmcnon rcsultmg fTom the tmp.Kt of celestial pr the Srh
gl.lu.lttOn, w1th w.lta bemg locked up m polar tee. jecnles ts actu.11ly quite h1gh. ( 11 1'\
The combin.ltlon of ~hrinking hvmg space and In analy~mg p:tmcular extinction events. s~ pod fat
cooler clim.ttt.• could be \CUte. mentolog~cal and geochenucal data are used C l \ TO\
Major features o f the Phanerozoic record 75
tli\IU~nun w11h 111fonnat10n from the t(mil Europc.:.m invnders colon r7ed the North Americ.111
,,1rd. It ha~ betollll' ev1dent ti·om this that no sin- shelf .~rcas (Sheehan, 1973). The ongin,tl North
mcth.tru~m lJil be m\oked to .lc.xount tor all Amencan communme~ had enJOyed a long
anucnon e\·c:nt,, .tnd indl'l'd some may h:ave period of \tJbility, but were narrO\\ ly adapted tu
tcJ from 'cvcral cmnodt•nt l-.1\l~t:, . parncular mches jn defined conununJC) structure:,.
The European specie~, however, had a gre.lter flexi-
late Ordovicion (Ashgillion) extinction event bility and were less dml'ly adapted. ' I 'hcse were able
.<,trd~ the end of the Ordo\ ll'l.lll, \C,l kvcl \tood to cope With adver~e environmental nmdmom, but
.:h productJVJt\' WJ\ v1~orou' .md ~.nvironmt'nl\ lt wa~ \OOH.' 3-5 Ma bdore pcf'\i~tt'nt communit)·
fa1rly st.lhle \helf &una' were divcr<>e .md strucntre!> were re-est.lbll\hed.
'l ~aptolmc \h,lk ,\ccumul,lted 111 deeper water.
l1cn, .1t the beginmng of the I hrn.mti:m (the top- Late Devonian (Frosnian-Fomennian) extinction
'' 'tJge m the Ordov1cian). the dim.lte detenn- event
d r.1p1dly, and the GonJwan.m ice 'heet~ Tlus event ts well atte\ted (with \Omc 21 "'o of
rrarlkd greatly. extending (0 qurte low latitude\, marmc f.1m1hes lost) but le~ well understood. lt
tir.111lC31ly Iowe.: nn~ St'a le: vel. The resulnng seems to have been 'paced out over ~ome 3 Ma
lnbrtcal Jismpt10n came m two abrupt ph.tsc~ (McGhec, 11J89a,b), but a~ a sL·ries of separate
1th ,1 more d1truse timt' of ~.:xt1nctions bet\H'Cn event~. and w1th a particularly sharp drop at the end
Arcnchley, 19'!9). fhe first pha'e (late R.twthL·y.m) of the h.l\111,\11. The wor't effects were concentrated
tl·J the pl.mkton (notablv ~raptuhtcs and t·on- on the troptcal reef cco\ystems and w:mn-water
idllnll) and al'o the deepcr-~hell benthm. where shallow m:mne conununnics. The vanou~ compo-
m w.l, a j.,Tfl'Jt drop in d1Vcrs1ry This may be nents. however, were ::tiTected in different ways.
nltd to ocean~<: overturn nl' JllOXll or toxic deep T abulatc cor<lh never recovered, though mgose
.Utr (Owcn .md ltobcmon, JlN5). ~hallow-<;hdf corals chd better. •md although 75% of brachiopod
nduopod~ w.:re .ll\o ttfected but le~~ \everely, .md genera became extJJKt, they staged a remarkable
c .trme tht: umquc, ~hort-iln•d, but vlftu.tll} recovery. a~ did the ammonmru. High-latltude
rldw1de H11111111licl C1una. dornrn.tteJ by bra- faun:ls .md deeper-wJter associations (as well <1'1
luopo<h, winch pcl"\istcd until rhe end oC the terrestrial biotas) were much less badly :-~ffectcd.
)rd,>ncian. \J..'hllt• tlm f.mn;t i~ nonnally of k,w These dTet ts arc commcnr with a 111odcl of dr.t,tic
rut\ (<10 genera), \Umc vanmt' may have up global cooling, but 111 th1s case sea level stood lugh
li ux-a, mcluding l<lcal re he b <lf earlier btttl.ls, throughout the whole h10ric cnm, anJ glaciaoon
d d1en: ti l'VIdeute Of b.lthymctnC ZOnJliOn was u:rt::nnly not mvolved. Another f.1ctor, how-
rthm the Hunauti.m t:nma too ( Rong and H .uper. ever, is the global spread of cuxinit bottom coudi-
l~N). tlons into -.hallow se.1s during the later Devonian,
Th~ \econd phase tKcurred dunng the glaci:U and thi, might itself h.we been promoted by a 'ea-
unum. Sea level \\a\ dr.l\llr.llly lowered and level lugh ~tand. One or more 111aJOr o..:earuc on:r-
b1u~k ~hell" arlJ redun:d, c.lll\lllg further dt\rup- runls of tlm ~tagnant bottom water would luve
n ~xtmcrion oC th<.· ~hallow-,hcl( b10ta. poisoned the upper warcr-;, and have led to severe:
The £hird and final phast• of extmcoon. ~ome fauna! extinctions. Anoxic overtums of tlus kind
h=ndreds of thum.111ds of) cars later. occurred in bee have been considered a\ 'idc effect<; of global cool-
muntllo rime\, at the 'Cl) end nf the Ordovinan. mg. At present (McGhee. 1996) a search cononues
1..11~ nme the gl.IC. I,ltmn \\a\ lar~d) over. and \CJ for evtdence of metconte bombardment Some evi-
I rose qlllckly bnngmg w1th 1t .lnox.tc waters dence: doe~ exist in the forrn of sever:~I mJCrotektHe
lud1 ~pread onto rhe ~hallow ~hdves .md killt·d off layers (melted glass spht·rulcs), soot honzon.!> ;md J
Himantia fauna. Only later dtd wann-water f.1u- poss1bk 1ridium anomaly, but these .m.• ~cattered ,lt
rc-c\rablish them\elves. trom the begmnm~ of various wncs in late Devonian htqory More dra-
~1lunan. matically, several crate!"\ attributed to bolide impact
In North Amenca che Late Orduvtdan brachto- are of late Devonian age. One of these, the 52 km
)(1 fJunas bec.m1e abmptly extinct, and when the diameter Siijan crater in central Sweden, may h.we
• .11 ro\e again tollowi ng the 1ndt1n~ of the: ~lant·rs. formed in the late Frasni<m, others at ditfcrent
tllllt'\ fhe touc~pr of muluple unp.tt t\, ,p,tn:d out bl•cn rcrogmzcd. While the hypers.1lme w.tters Crete
ova ,\11 a:xr~nded tlllll" r~ nod, ·~ beginmng to look these \t:Js \H re generally hosuk to htc, except Ior rht K
.1ppealmg the .1lgal rectl; themselves and a t'i:\\ \pecialiu unult.Il
111VI.'rtl'l1rJtl'), the ;lmount of ~air taken out of th :~ nd piJ
Late Permian extinction event upp1.•r water of the sea and pem1am·nrly locked ,l\H\ thnn.t~l
Tht' .ltme cm1~ seem~ to h.tve ~p.mned .tt Jca,t 10 in l v.tponte 'edimcnts must havt:· bl'L'n <on~tder.1bk UIJI11lC
M.t, ,111d w.ts .1bo coupled wiLh extuH:non of rcrrcs- Whctht·r this w.l~ l'nough to rt•duct' tht• ~ahn1ty fit rlwugh
tn.tl t:nlll.t\. 1 hose m.mne t:HIIl.t\ \\ hH. h 'lutfercd the wrlau· w:~tcr' 'n a~ to .1fti:ct advt·rsdy tht• life 01 l'a:nni.n
gre.tte'r rcdlltrlllll wert: tloptt':.ai 'lL'llohaline groups, \tcuoh.tlim: mannc organim1s i\ d~b.ttablc. m.trllll.'
hnth bt:nthK .md pd.t!;lt. and the umoid-bracluo- It \\,1\ ftmncrl} thuught th.n tlm Jrop 111 \C,l lcvd ~penes
pod- .md bryozoan-donun,ttnl et:O\)''Itt:In~ of tht: · v.. a, the rc,ult of global t oolmg, hut the PennQ- I ht:r
I..ue I'JlaeozuiC were tottll) dc:,troved . Y ~:r bivalves, C.trbomferou' ~anation w.t, fi1mhl•d by tht· J.ue C\ Cllh.
n.mnlntd,. ~asrropods .111d LUtHldt:>lll' ,uffered com- Pcrmian; smkmg oceanic ridgt>' •.., .1 more hke~ dunng
p.tr.ltl\ l'ly Im h.·. and Pumt.lll ~J,trupod' llldecd LJU\C. •1nlv :1
h.tvt: provt:J .m excellent group t()r 'tlldying con- A ILtrther factor seem\ w h,l\ t: lx·cn ma,,t\l t'\:tJtll tl
troh on t•xtmctton and ~urvn .11. vukanit H), centred in SJbcna, caus111g a dr.1m.1t1. th.mge
F:um.1l .mJ geochcnmal d.1ta \tlpport :1 gradual, intrt·a~c in atmospheric CO, level\. wh"h m.1y lm·, rt•mton
,md 11\lt .1 cata,trophic a:nd-Penn1.111 event, though led to oce.mic .tnoxi,L Tl~cse mulutanou~ t:<lliiL'\ < .thbr,tt
\Vtth \llCl"t''l'ivc sharp pulse' wuhin IL. M.my fucrors collcuivcly lcd to previomly unp.!rJlklcd env1ron· lll.\Tlllt'
\\'t.'Tt' mvolved, ,l!ld no one \lllgk t'lll\e 1~ envisaged mcnt.tl 111\tJbilit) and ~o to roliJp\tn~ t'lmy~tc1m . hryozo.
(En\lll. I<JlJJ). On~: unport.lnt t:tctor was the 13y thl end of the Pcmllall th~ 1,1\t reef enviwn· alsll det
p.ll.tt·o~t: U!,'T.lphy nf tht• tmll'. 'mce Ill the continene> ment\ had \'Jill\hed, and aU J\SOll.ltCd e<.:U\)'\ll.'llll llOlllll't'
wen· tht•n h'emblcd mw thl' ~tant 'upen:ontmc:nt lud coUap,ed. Nonnal manne tondiuom Jo not l rctJt'l
of P.mg.1c.1, w1th dw tnJlll.,'l.llJJ l'ethv.tn occ:an ~ecm to have been restored for 'l'veral mtlhon )'t'Jrs bt'( 11 gt
mtlenung ib CJ'Itcm margm \m h .1 configuration The Permiau extinction therdorc (MaX\wll. 1%9 md or!
would n•n:aml) h:~ve ,1tfct red d1m.uc and ocean seem~ to have mvolved glub,1l lOohng. manne nt.llll c
nrntl.mon ..md knock-on bioloh'1< .11 <.:tTcns would rcgre~m)ll, knock-on effects of thc\e and pcrhapj \'llk.tlll
h,IVa: bct·n mcvit,1bk. St•cnnd, tlwrt• wa~ the mmt \t'vcrtJl other factors. nwcry
cxl\.'11\IVC known m.truw rq~rl'\\1011 l>f .my time Ill lt.tly
during thL· Ph.mL•rozoic Tlm rL·grt'\\ion beg.lll m late Triassic !Carnian-Norian) extinction event ltyt·r I
the c.lrl} Pcrmian <md h\ till' t•nd 0f the Penman the: In rhc brc Trias~ic there were heJV) lo~se~ 111 num houmb
overall drop in 'ea lcvd h '' been cakulared ~ uwertcbratc groups and the ammonotd~ and bt\•alve~ " dcpk
210-2:iO m. The lo" of mannc h.1bH.lt \\.1~ very only JUSt ~urviVed Conodunrs. ~:on ul amds and mon: .1
great.•mJ dm, rogetht:r with mtrh·d 'ea,onal eh- strophomcmdc brachiopods beGune exnnn. Lan th:lt lt \
mat\.' '"'111h"'· led to .1 'tngul.tr L't nlogit Jl mstabiliry f..1una~ al~o ~uffered dramamally at rhts timl some 1
and l'xpbim aho \'vhy tlwrt' ~rL' \ll fi:\\ good bound- Exnnctions of marine tauna~ were spread throu~ 1-.trth ,,
.try >l'diom .tg.unst whit h thL \L t:!llnal changes can th e Scyth1:111, Ca.rnian and subsequent Nomn In \t
be I.Jhbr.llcd. W ann ~hallu\\ ~L',l\ lw1.Jille reduced to (Hcnton, 1l)90), and an extraterrestn.tl cause thu1 utdiunl
cerl.lln pam of the world only .tnd much of the sccms re more . Although strangr.tph lc resolution tl to 75 I
Penni.111 manne sedimt•ntatwn Wil'> depmitcd in less good 111 the T riassic than tt IS, for ex.unple, tn Ill 11 011-
inl.mJ ~ea'> nmne<.tcd to Lht· ou::tn by narrow the Jurassic, d.ltJ from .European pcctuud bivake~ ,,rlll!-,'1.
chJnneh .md bordcrt·d hy t'nonnml' algal reefS. and crmo1ds espectally Oohmon and ~tnHHS, 19:-19 tl•l.ttton
Ot tht·'>e the be~t kno\\ n .uc the 'ea' of westem shO\\ substantial reductiom m the: l.JnHan. conrem· d.tt.t tTo
I ex.l\ .md nurthwc,tcnl Europt'. Evaporitc:~ were poraneomly wuh Wldespread t~lues changes. ~unrm 1(1 ,1 dtJ
dcpo,ited both behind the r~d\ J\ ,,lit water washed and Ruffell (1990) couple the\l emmanom \\1tb l'( l1011
ovn them and m the b.mns thetmclve' l\ they peri- '}m hronous climatic changes, parricularly ro tht lJptlm
odlc::tlly dncd out. The ll1w-l~ mg rt•gJOns formcrl) oml'l of mcreased ra.inf.111 dunng .111 othenv1se ana I' JHf>) ,,
occup1ed by the~e sea' \\erl' 'omdtlllc\ fioodt:d, time. This may h ave led to a change 111 ocean ~ur ,, htg h -
on I) to tlry l)Ut again; m the Zel h~win ~ea of north- fucc rempcrantrc, pH or ~al nutv ch.111~es or loss o! tlW of
we~tern Europe tivc ~lllh dc'>iLtalion cycle' have tarbon.ltc habitae>. tht• md
Cretaceous- Tertiary boundary extinction lie: ;1 tcw millimetre~ of rock dcvmd of '>pore~ or pl.mt
fh,, K-T bound.ll) is nurlwd by the morl' or le~s cutldcs, but otten w1rl1 ~apropd or filsain, md11 .ltlng
lUltJoeom l'Xtmcuon uf tll.lll\' group' of 1n1m.1h rottJng or bumed vegctauon {.tnd mot layel'\ .lt tlm
xl pbn~. rangmg fi-om ph) to- to zoopl:mkton. level mdicatmg huge wildfirl'' h.1ve been t()und
1mu~h dnunonnc~ and beknlnHC\ to the tl rrc<,rn.ll, world\\1de). Thert• follow~ a few ccnnmerre., of ccul
ntmnr and tl; mg repute~ ,md land pl.mt,. Even or nmdstonc with onl~ ti.>m spore-, and mq~JI(l~stls ,
though tl11S cm1s was le~~ \l'Vl'te than thal o( the late and thereafter .Ulgio~penn rem.um mcrease wu h l'Vi-
Pmnlln. \OIIIC: cstmutc'> 'ug~t:\t that 15"'• of all dt·nce of greater pn:cip1ratton. ~howmg a ph.l'l'd .md
nnr !Jnullc\ .llld up to 75" .. of ill Crl't.Keous b'TJdual recovery unnl a tropic.tl ratntore~t \\a' l''>t.th-
el pt• c' \\l'H' clmun.Hc:d. h,ht•d The tu11t"\Calc here IS about 1.5 Ma.
;)- There n1.1y h.l\'c been no \ingk came t(lr these hv1dence oftlu~ k.md is compelling, .md j., roml\-
tc \tlltl, indeed tht: slow dellim· of the .lllllllOilitcs tent with a dra~tH: .1nd '>uddt•n environmnu.al
ly ·mng th,· l.m: Crl'taccom, .md thctr restncllon to tr.nrnu. followed by a ~low rctovery. The ht...dihood
J tl:\\ p.lrt' of the world pnor to the1r final rhar .111 astermd tmp.tct wa\ n.•,pomiblc h.t, been
xtuJCilon. m.t} 'ugge~t that ~r,tdual envinmmental much ,tcccntu.lted by the ruscovcry of a 200-JOO km
h.m~ \\':1~ .1 pnnury control Th1s vil·w h.1~ been wrdc crater of t•n<i-Crctaceom .l~t· .1t Clm:xulub 111
re.nfllm·d rl'cl•ntly (McLcod l't '''·· 19CJ7) by refined the Yucat:.m Penimub. lr is buncd by l.Jt~r 5Cthmem~
lihratiot1 of thl' tnne range:' of ~evt:ral Cret.ICeous but h.t~ now been ~cophysically nupped, and \how~
n- nn1' orgam\111\, wh1eh •hCI\\ that O\tracod'i. ,lt lca\t three vJst conccntnc nn~" of the kmd to be
ozom•. md bl\'.llvc~ ..Is \\ell .Is IJud re pules were expected from the tmpact of .1 bohdc over 10 km in
n- ~ dcdimng dunng thi, tnm Yet there w,t, ,t pro- dumercr. (AJv:1rc7, 1997). Onlhnt-." m the Canbbean
11~ lliiJCcd ~harp t xtmction .H rbe ven. l'nd of the m.tdt• 111 1997 grve t'VIdencc ot" gr·t•:-tt di~turb.mu: m
ot uwn·om, Jnd in recent y~'Jl> much ~upporr has ~t·dulll'nt~ of the K T bound.•ry, which :lfl' ,urcJy
rs m giwn to \.ll;l\trophlr' theones to exph1n this tlw n•,ult of shm!... w.wc~ gcncr.ttcd by this colli~lt111 .
1)) md othet wwr~ t:'\tincuon penods. Thetc .1re rwo 'lome gL·ologi~t\, however, bd1eve that intcmc vol-
llC :n.lm contcndt'r\ hen·; bohdc tmpact and tntense lallll activity around tht.S tlmt• (ould equally be
•P' vuhrucit\. The pre~ent dcb.ne d;ues from the dis- re~pono;iblc for lhe iridium hyn (or layer...). ~lub.1l
>my of Jn tridtum-rirh l.lVCI' in manne cbys wtldtires, and thl' drJmatic Aor.1l .md t~umal Lh.Hl~e~
JlltJiy and Dcnn1ark by Alv.trt•z t'f al. ( llJHO). Tbis duCUilll'nted, and tmkcd, the n'i M.l Decc,m Trap
he prt:osclv at the CretaCCOU\- r ertial)' Rood bJsalts m lnJ1.1 t·omcidc e;o.;.JCcly \\.'lth the K- 1
ny ~~d.i~. J~ fl'CO~'l11Zed by IJurul ch.mgc,. lndium boundary. The<ot tluod ba,aJt... entpted for a nulhon
{'~ 1epletcd m the Earth\ lnl\t bm i\ J() l 11! 1 mncs yean or so. Poss1blv India wa' nugranng over a hor
11d 1~1t Jbuudalll in n•cteontt''· .1nd H W<1S posmlated m.mtlc plume at rht· time. AlrlTil.ll!Vcly the vokan-
nJ lw1r wa\ dcnvtd from .1 l.trgc ,l'ttcrOJd or mt•rcorite Ism nught have been triggered otT by anothct .tstcr-
lC IOlllc Hl km in Jwnctcr wlurh colhded wtth the Oid unpact. There .tre iridium b\'l r... 'pannmg up rn
gb E.lnh al the end of the Crct.ltl'llll\. 20 Ma on either side of tht· K T bound:tl) . .md
l;ltl In support of the .1!>tcro1d nnp.Kr hyporhc\t~ rhe more needs to be known about the chstributron of
1US dJum l.1yer h,ts now bet•n t~mnJ wnrldw1de (up mdium Ul SedllllC!lt\ generally. f Ikl'\vi~C. ,) !110ft'
I~ 75 localines), often w 1th 'hocked quartz grains retincd corrclatiOll of marine and tc..•rrcstrial CVL'Ilts i~
lll non-nlJnne a' well a_, nunnl \l'qttence' Whereas neeJed as a test tor synchronctt)' Even though the
\'C'I grJphic n·,olution Joc' not pcnntt a prcn\c correl.ltlve effects of ,1\teroid imp.tn and vukamnty
~9) non berwcen marint• .md rerresmal 'equcnces, rernarn unclea•, .tml although this ilJ'' to be ~er
111- ·,;,,from plant to~~1ls over till' K-T boundary pain~ :tgarnst a backf.rround of other rh.mge,, the JVJII.lblc
111~ 1' J dramatic and sudden t'llVITOIH llental dist urb:mcc. !.'V1dcnce shows uncyt1ivocallv that .1 grc.tt ,md t:lr-
·ith SI tlM~ tn \\l''>tcm North America (Wolti.· and re:~chmg catastrophe, or indt-cd multiple c,ttastro-
the. Upchurch. 19H6, lloultcr c·t ,1/ • 19RH: Upt hurch, pht'\, dt·vastated the Earth 65 MJ .1go
.rid ~)are pamwlarly 111Structl\ 1: m tlwo. context. Here ~nt.lller. but neverthelc,~ 1111portanr sttp\\1\C
ur- htf,.ft-dlvermy cver~'Teeu brn.tdleaved tlor,t, mdJCJ- extmcnons took pl,tcc in tbc E<Kene and Ohg<KLne
uf nl t of rather dry conthtiom, .tbruptly tcnnuures at (Prothero, 198lJ) and dom.inautly .11li:cting: l:.tnd .1111-
emdium-rich K-T buundal) t l.1y layer. Above this mals in the late l>ki~locenc.
UniverstC) Prl.'~'• NcwhJvt•n. (S) mbtotK ong;m Awr,uml.., ').11

eukaryotes; biology. h1ot. hl"ml\try. palaeontology) hti.•, Ill fire
Margull~. L. ( J 9H I) <;)'llllrit•.•r., 111 Cdl Et•c•lutwu - LJ(t• auJ D G . Smid
Clll'iroummt cm thc hili)' Emit. freeman. ).ln Franruco. 1'1'· 3-1'1-6::!
Books, treati ses and symposia BarghCiom.
(Up-to-dltl" ~r:llemcnt l)l enJo,ymhm''')
McChl'e. G .R . ( l4lJ6) '17tr Lllc Dt'n•11wr .\l,u) E.\tiiiLitt~r. lme1ic.m 2
Albntton. L.L. \I 'IX9) Lrt.t-lr••rl"' I.prs.•dt•s ,, E.mll
tlrt· Frasman/hrmc·mum Cmu. l.olumbiJ Umwrol!J flrec,uubnd
1-li~t••ry T'optD 111 tht· E.mh Sueme\ No. 2. Chapman
& H:tll I ondun, 221 PI' (Cie.tr 'impll' prc:'l:ntation) Prc~s. New Y orl... ( l'he hl'\1 trt.',ltnll'nt .1\',Hl.thk) ne ngt\llll. ~
McMenamm. M A\ and McMen.mun. D.l .S (I'JW P;~laeozmc
Alvan·z. W . (14<17) 1' '~'·' ,m,/tlu Cr.rtu ••I Doom . Princeton
UmveNt\ Prl'' , l'nnl'l'Wn, 1\ot•v. . Jer.t•y, 1X5 pp. TI1r Emcrl!wu ..f -!t/1111<1/s - t/11· C.uulmmt Bmrkrlm•11~ (Lower C.u
. Columb1.1 Untvcr-.lt)' Prc", 217 pp. (Stra1ghrforwar.l. Hcn~tson. S
(Popubr .tccuunr ot lllt'tt·ome tmpa ct hn1othem)
readable text) mKrofo~'JI<
Ber~>ren , W A .wd V.m Couwrin~. J. (etb) (1984)
Nielsen. C. (1995) lrrmra/ Er•t•lwwu. <.h.torJ U111vcmt'l 20, 751-62
Catn>ll••rlrn .wJ f..rrrlt flr st••IJ The New UnJtormi-
Press. (C!Jthscic, cmphJ~tlcd) llcn!o:C~t>n, S,
ranam\m . Pnn nmn UntV<'~">lf\ Prt'\'• Princeton, New
Ra3ben, M.E. (cd.) (I'JHI) lht' /,>mmotimr ,'1/tr(/1' ,,J Il ( r•J•Jo) Ear
Jersey. ( IH p.tpt r' dt•.tl' wtth lll.J\S l'Xtmrnons)
Cam/mall l...t1Wt'Y HtHIIIdtlr)' Jlr,•blrm (tr.ln,Ltted frnm \lm~t>Hs t1/
Chaloner W.G .mJ I l.tll.Hn, A (1 '>8CJ) Evolution aud
ltussi.lrl). Amcrind Publi,hing. Nt.·w Ddhi. (Illustrated 9. 1-JII.J.-
excu•euo11. Pllil<•>•'J'IIIull 'Jhm_,,1rlitlll.•· 4 tlw Royal Society
R:ill', R.A. (I CJ%) 7'lte Sltllf'l' of Lrfi-': Gc11es, Dt•JtdopmCII Ueuton, M.
~~ L,mdoll U 325, 2-1 J ~·m. (1(, syn•post:t! p.tpers; most
tllld tire lil!ollltion <1/ t lnilllllf Farm U uiveNity n( ChiCJgt' Syt1111csis (c
imprunut)
Pre~~. Oinks p.llaenlllnlugy .111d mob.. ular gem·tic\) Lilackwell, '
Clark. R.D. (I 'Ill-~) IJ)'II<Imio i11 \lt•t.r ::•••lll E11ol111io11.
Schopf. J W ( IIJHJ) Fmt/r 's r:,,,[i,•st BitlSphcrt - ll{ ()n~,, Ucrgstrom, J.
Clarendon l're,s, ( >xtnrd (< ' l.mtc text)
arrd Ul'tllllli<llt Princemn Umversity Press. Pnncetnt new phylw
Donovan s K (ed) ( IYH'I) \la>S r\tlll<li<IHS: Pr.lt'C.~SI!S tllld
New jl'r.ey. (I "i \l:'CilOII~ <111 on~'ln nf hfe aud fu· Ui.'rg,trom. J.
Er•it!m•c Bdh.l\eu. loudt•n, 2(>(> pp. (Exrepnonally
et·osystl'lm) orgart.i,ntS
usctul lolllp11Jnon uf 'ewul p.1p0.:I">)
Se hop f. J. \V (o.:c.l ) (19'J2.1) .\ltlfM Fr•wts irr tire· History Boucot, A J. (
Dyer, 13 I' .md Olur, R .A . (I '''J-t) I Mt1!1{! tire History of
f..!{r.Jon~ .1nd 13artlcrr, Bmton, Mm. ((I wnthcttc pJpeo IOJ,.,'lCJ.I VJCl
bik.JI)'<'Iir Cri/, tlrr f:rll\'111•1111 -:r111/r Columbi~
Schopt~ J. W lnd Kll"lll, ( . (t•d,) ( ttJY::!) "I7n· J>rollcr. z \CIIlluLmng)
Um\iel'it(~ l're\\, f'..e\\ Yorl.. (Linr c.Nl) understood)
Bro>plrcrc Cambridge Umn:r..ity Pre,s, C.unbml~c . (11 Houlter. M.C
f:n,;n, D (I ')'J.' /111 <.rr.ll fJJ/..•;::,1/, Cnsrs Lifi• .md Dr,Ith
papers) P1ttem~ of
in thl' J>rnrumt Cohunhi.l UniveNf\ Pn:-.,, New York
Sliver, L. .md <.,l'hult1. (t'ds) (I'JH2) C:r,•l·~
P. cv•dencc, u
(Vay ~ood .md reJd.lblt•)
Implir.wom <iftltt· lmp.lct 4 Ll~t .·htc·r,•rc/s .md Ct•mrb Cl1rendon,
Frakc,, l A rune•'· I E 1nd 'wtl..ll,, J I (I 'J'J2) Climntr
rlre Enrt/1, Gt·ulo~c.•l Socltt} llf futKOl.J Spl·o.U Papc! Hmrcr. M D.
Af,>dt•.t o{ tltt l'h.m, r.•:z,•i ( .1rnbndge Umvl'mty Pre~'
No ICJO, (,cologJc.ll \ool'lY ot Amem..•. LWI.Tl'n 17n On\!111
(Excellent synthe''')
K.1n (M.111y p.1pcr.) I lomt•), ~y·
Gel', H. ttJ4h) &.·/;,,, tlrr /3.r.J.:b,•ut•: Vtnt•> l'" tlrt Onl/111 "f
W.uter. M.R. (eel) (I \J7h) Strom.ltt•lrtrs . 12, Ac,1dem
I 'trtrbf<ltt:> ( :h.tplliJll & I 1111. London. (Dcutero>romc
AnNcrd.1m. (M.1ny on~111.1l p.1pcr;) M.D. 1
Uro~,ic.:r,
reboonsh'P')
Ward, P.D ( l lJ92) 011 \lrt/tuq•ft~lr'< Tr.ul: u11in.~ ft•s ~•c•n./ormt.ll
Glaessner, M F (I 'IX-!) 17u· IJ.m•u t!f .i111111<~l L1{e - ,,
a11d the Great f!.\tirrtiii!IH. Fn:t•man. Ne\~ Yt•rk l!rJ~ler. M.D
Bwlwttmntl '\trrd)'· ( .uuhrul~c Unwcrs1ty . Press,
(Exn~Uent li~lu rl·JJmg) CJ.r(y ~kdt:ta
Ctmbndge. 2-1-1 pp (Nt•w, vt·rv de1r 'unun.rry)
WiU1amson, I) I. (I '>92) L.an•<ll' .md l!Pc•lution: I""''" is <J( Lo11dnn 1 4
H,lll.1m, A. ( 1977) P.utm'-' q/ l :r•o>lllt/1111 .rs lllwtrllred hy the
Nru• ?.aoli~flY· Clupm.111 &. f IJJJ, London. (Unonho· Br.•ster, M.n.
Fvssil Ream/. Elst·vit·r. fumtcnl:un. ( l7 oriwnal papers)
dox, but intcrcstin~-t) tlte biolog-i
H:tll:un, A. (I 'J'J2) 11/r•llll'lll ::'<llt' SctJ-Icl'rl Clw11ses. Columbia
WiUrncr, P. ( I'J'JO) lm•t•rtl'bmtc /{cl.rtrciiiJlriv{. C.tmbnd,, C',unbrian-1
Univcr,ity Prc"· New Ynrl... (Re.1d.1hle synthesis)
Umvcrmy l'ress, C.11nbndgc. -100 pp. (Modl'm. ver ft~olrllloll t!/
House, M.R . (t·d.) (JC>7<>) T/rr Ot!~i" <!f J\J.!f••r luvertcbratc
readable Jnd up-tu-dJtc trCJtlllc'IH) "'b'lmr), Pie
GrOIIJil, Sy,tt•nunc' A'~l>UJllon ~prCI:U Volume No.
4!'13-524.
12. A(.1dcmrl Pn·"· l nndon (M.111y e"ential papers)
Brcnchlcy, P.
L:trwoud G P (l·d) (I •IKH) f7\llllolltlll .nul Srm•wal 111 the
\!,w F.\.1111
ro.,sil Rt'ttlrd ,,,.... lli.\Ut.' A\\t.lU.I[JOJJ Speoal Volume
Donovan), I
No. 34. Cbrcndnn. Oxtnrd 36:l pp (1 '; p.1pl.'rs, cxcci- Individual papers and other references wmpbcll. N ..
Jenr qu.tllry)
from the Le
ltpp\, J.H . 1~1d Stf!llnr, P. W ( 1'192) On~111 nud Early Alvarez, L.W .. Al-.are7, W, A,,tro, f· .md M1c.:hcl. 11\
Wt:.'it Hi~hJ.
E•·olrm,>u of tlrt' ,\ lttll l•'·' · l'lt·num Pre<>~ . New York and ( llJHII) Extratc:rrc\tn.ll c.:.lllse for the Creuctcotn-
\•1it-bod!ec
London. (I(, p.lpt•rs) Tcrti.lry cxw1tuon ..).mru 208, lll'JS-IOX. ( fhe me«
CLuk. R.B. (
MJrgu.IJ\ L ( 1470) ( >rit:m tl( I.'u~wyc•ti, CelL<. Yale omc tmp.lct hypotht•,is)
Bibliography 79
n ntk. \. \1. ( 19H2) 1 he nn~m .111d c,trly rvolutlon of On,qm <1{ ,\ f11jor /ml£'ficbmtr Groups (ed. M.R. llou~e),
• in Tht C.u11/md~t' hr.-ydc•J•<~ditt c•{Earth S.iwces (cJ. 'w,tem.m cs A\Suciation 'lpen.tl Volume No 12.
G. Snurh Ltmhnd~.: Umn-l'tf ! l're-~s. C.unbrid gc. Aodemt t Prcs\, London, pp. 55 I 0 I.
r1 Wl-r,z. (U~eful wmhe,") Cloud, P. (1976) lkgiruun gs ofbtnsph eric evolutilll l and
tccm. E <;;. (I 'J71) Tlat• oldt•,t to~~• I' 'lomt•/it their b1ogeocb emical consequt·tKt:s. P.rlcobwlc>l?y 2
nUllt 224(5), JC~4:! (Fullv tllu,rr3re d work un
1'i 1-R7 (Recent work on late dll'ntic.tl early btolugi-
l't.<llllhn.lll t'Vnluttllll)
c.ll evolution )
:ngt<M, ~ ( I%M) Tht• probh.·nu ttt 1\mib m the l.'.lriV
Cl>m\JY Morn<, S. (1991) Ed1ac.ua n-ltke fo"tls m
p OZOI< {ctol ( llll't'YSit Ill\ [. j•,clflt'IIW 415, 1-71 ( lmbn.ln Bur~C\S Slulc-rype- fauu.ts of Nonh
L,>wtr C.tmbn.m pt nhlem.llt ta) Amem.1. PalacoJIIOiogy. 36, 5()3-tlJS.
~·t10n \ ( 1977) EJrly <. :ambn.ut butttln-,h apcd ( nok.. P .J. (l'JIJ2) Phospho gcncm .uound the
r ru"il from dtt ~Jben.tn pi.Jtlllrlll. fi,Ji,li'Nitc•/,!1/Y Protcro7o tc-Ph.lne roZJc rran<ttmn. J••ttmal c>{ t/11
ZO. ~SHi.:!. ( .eoh~~iwl Society ()( Lmdm1 149, 637-646 .
~'!On, S, (.onwa' Morn' S., <.'tloper, B.J. et ,,[,
( 'ool..., P .md Shergold . J (19l:l.J) Phospho rm, pho,pho -
'/'!( 1 Edr1y Cambn.tn fo,,tl' !Tmu South Au\tr.Ilt.1
ntl'~ .md skeln.tl evoluuon .1r thl C .unbrian-
lc /llc'lll 41/tr Assc)(lclfl<lll C!f • lrWrcllrl.llrl/1 Pctfctrc)JIIcli.>I/1.~/J Prrcambn ,lll bound.rry . Naturt 308, 231-<>.
~ l 3114. CnmC\, 1 P (1976) The maogr.1p hical \t~mficanct of
, M (1990) EnJ-1 rt,I\Slt , Ill P.t/rlt'clblc•io,l!)'· cl rr.1ce fth\ils. m lhr S1udy C!f Trace Fo.<SJI( (ed R W.
\ nrlrtsis (t•d\ D.E.G. Bngj;S .md P.R. Crowtha ). Frcy). !'>pnngcr-Vcrlag-. Dcrlin, pp JCll)-.30. (Trace to~
~well. <. htord. pp. I'H -H
'11 evidrnce on bc:ginnm~ of metnoa.n hfc)
Jo m. ' (llJX9) Th.: OTI!-''tn <>f Jntnul phyl.! d.nd thl'
Cnme,, T.P. (l'J92a) Ch.mgc~ m the tr.trc fos,il biota
n~w ph)lum ProcoLlomat.t l ..l'tlwia 22. 25 11 h'l .
.tt ross the Protcroz ic-Phanc rozoic transition . }t'llllllll t!l
om. J (l <J97) <)ngw of lu~h-r.mk ..,rroup' I l l 1/u Geoh>l/lf•tl.'>t•.icly 149. 637-1(>.
IUmt\. 11.1/ar,•mcllc>t:ICct/ Rl'>fllT<h 1, 1-14 C rtll1C\, T P (1992b) The rccord oftracc: fowls acrn'' thc
I U<Ot, A.J (1 1JH3) Dnc' l'vvluuon t,1kc pia'" 111 au t.:ttl-
Pt oterozoic -c.u11bri an hound;,uy. in Ori~itr dill/ Early
) ~1 \anmm? ./t'''""'' <?.1 p,,J,,,,,,,f.~c,:y 57, I '«I (V c rv [t•olrttlt'll of the Mrta:;w,, (cd~ J.l I. lt)lP' and I'.W
t n• 1ung} )rgJlor). Plemtnt Prcss. New York ami London, pp.
\ 8uult•·r, M.C. Sptccr, lt.A .•tnd Thnm.l5, ll A. ( I9HH) 177-204
m> of plJnt exumno n !rum \tune p.UJttlbo tantol Do~\ td,on, 1:. H., Pcre-l:"on, K.J. .md ( ~mc:ron, R.A.
., 1
·nee. 111 £""'""'" .md ·"""'ll'cll 111 1/tr F''·'sr/ Rcr.•rd,
( crcndon, nxford. pp. 1- 11> .
.\I 0 '1979) Th,· (. .unbn.ln r tdiatmn ~ Vl'nt, 111

(I 'J':l5) ( )ngin llf bilatl'r.than bndy plms: l'VOlunon or
dcveloptnent.U n·gulator y rnrchan.isnl5. &irrut• 270 .
I' U19-:!5
Tlrt On.~rll c!{ M.tjtlr Irwmt!mn r Cr.•up> (t'd. M.R. Gehh. J.C.
!WO l.:.Jrlie\t known c:chmnd cnn - ,1 new
!cou,e). 'y~tl'tlLlllC~ A~son.tttnn Spt•tial Volume No cdJac.lran fossil from the Pound sub~rroup uf ~ourh
1 4, Jdenuc Prc''· l ondon pp. 10.\ 59 Amrrah.t ,·Hdtrn~t.t:a 11 . B7-13.
I,M.D (11.)92.1) B.1dq,,rround to thc: Cm1bnJn CXlllo- Gl.tcs)llcr,
M. F. ( 1%1) f>rccambnan anmuls. Stimt!ft•
on J'llt'lllll <'/lire Gnlhlgico~l Sc'firl}' of L.111dou 149, SR'i-7 • lmericau 204, 72-H. (Edr.rcar.t f.1un.l)
· M.D. 1992bl Nutm:m -umt.hl· d Wltl'r\ and the Gl.tl'~sner, M.F. (
l'J62) Prccamb nan lil\stls. H~crh>~tcd/
urlv ,kcJetal fnm.I fl'lord. Jc•~tma/ cif tltt r.c,,f,~t:"t~l Sool'ty Rt•l'iew.' 37, 467-1)4. (Edt.lcarJ Clun,t)
L.111don l49. 621 ·9 Glw~sncr, M.F Jnd Wade, M (IW.l•) Th~ !at<'
MJ) (I '19:!c) l'.tbc:mc:.mograph)- and dung!:\ 111 Prccamb nan fo\\ils from Edt.tr.trJ, South Amtr.ilia
btoln!-'1'·'' cyrlmg nl phmphoru~ .ll rm~ thl' Palaet'lllohlf!Y 9. 599-62H . (Edt:H.:ara fuuna; technical
mbnan-l'recambnan boundlry , 111 Ori.l!'" dllcl b1rly de~nt ptm ns)
11 11 ••I thr .\/rt.t::oa (l'd' J H Ltpp' utd P 'W GIJl\~oer, M.F. and W.1dc, M. (ll/71) Pt.u·..ambndlllm: .1
t:;nor), Plc:num Prn\, New York .tnd lnndon. pp. pnnuttw .trthropo d. Lctlwia 4, 71 - fl. (Edtacar~ fauna)
J-524 ( ;rt}. J (llJH7) Evolutio n of the frcshw.ttl'r ecu') ,tenr
hln. P (1989) fhl· l.ill' Urdn\tli .m cxtincno n , 111 the fm,tl rel·ord. Prllclt'tl,I!CI>glclplry, p,,/,u·ocllllll11<11t~I!Y·
ft>, E:~tlltdnms: Pma·.un mu/ Er•rdmcr (t·d. S.K fJa/cli'C)I'tcllt>I!Y 62. ::!· 214.
~nn), Udhaven. I ondtlll, pp . 10+-32
(,rotnng lr, J. P. l3ownng, S.A, Sa)'lllf, tl./ .md
_::rbt I, N . 1 td P ~ul, C .R.<' ( llJH' Pt'U(Jfll r.U t(l,stl, KaufiJJ.IIl, A.J. ( 1':1'!5) lliO,tratt!-'l'·lPhic .md geoLhron o-
111 thr Lowl'r C.unbn.m ptpc nKk of lht· Nnrth-
lo~-,ric comtrain h on early animal cvolutJort. Scietra· 270,
~N Highlmds. S!llttisil )•'""'''' ,,f (,c,,f,,~}' 19 347- 54. =;<;H-604
•-txxhed problenu nc,t) I Jnlland, C. H. (I IJH':I) Synchron(}logy, r.1xonomy ;md
1. ltB. ( llJ79) RJd!.ltlo n of tht• Metazo.t, in 'I7w r<:<lliry, 111 a,.,,fruhlll mu/ Extinctil•rl (cds \V C. Clulnne r
.md A I l.ill.uu) P/rr/'"''J'IHw/ /i,,,,,,ffiellr.> •!l the Rt•ynl McMcn:unm, M.A '· (I')~'J.) 1 he ongm~ and r,ltiJ.l ... Jl .. ~ .... ' !
St•ncty R 325. ,h.\-77 of thl' e-Jrl) Met,\70,1, 111 I ,.,,Jraicm 1111d tlrr h15stl Re; IJ(\IIUl diV
Jenkm~ R .J f l'l<l..!) FundwnJl mJ ~·colug,K.U <ll>pCC!~ (ll"h K.C. Alien .1nd lJ f G Bn~"). Bdh2 ~ tutctilm
of E.li.tt.Han ,,~cmbl.l):t''· m CJ11~111 mul /;.uly r;.,,,/utiOil London, pp 7J-IJH. ~ pki>Ski _I. (
·~/ tftt \1, ,,,~,,, (l·d, .J I I l1pp' .md P \V. Signor). Moore-. J .tnd \\lillmt"T P ( llJY7) Convergent evol syntl!r.sis
Plenum Pn.'"• 1\:,.,, Yn1k .m.! lon..Jun, pp. 131-1 - 6. in in\l'rrebr.Hl''- Br,•/~r,,r/ Rrl'lt'll'> 72. 1-<•0. BIJLk\\dl
JnhiNln, A LA .md Sumtt-, M). (19R'J) Thl.' timin~ Jnd Ncwell. N.D. ( 1963) C.. me~ 111 the [u,tol') of lite..'i.: SC'pkmkt,J ..
C'JU'< of rlw lltt' I n.l\\ll m.~nn.: mvcrtt·brJtc cxonc- \mnmm 208, I I o. ('Ctt.Htmphl'm · " a dctcnn I \V (l'lH
Ulllh t•ndl'll< l' from ~ •.,IJnp' .md cnnmds. in \/,us for nUJOr cxrmcnon penod') rr.. ord \.',
£,-,,,w,•m: Pr''"'JSI'S .md /i'i•/r/111' (cd S.K Donov:lll), Owt'n, A W. and Rnbc1t,on, D.D.R. (1995) Ecol~ ll\ y throu1
lklll.lvcn. Londun. pp. 17+-'J.f. th.m~c' dunn~ the: cnd-OrJo\ ioan c:xtmctmn ..H ~hcch.m. P.I
Johnsun. J.C. (1'174) Fxnnltmn of pcrched tauru~. C.~t't'l••.~y 20, 21-.W gl1n.tt1on
Gn•/c•~r 2. ~7<J-82 (r.IUII" lt1lnn1zmg flooded ~helve\ Pancf\Oll, C'. .md ~uuth. A.D (I <Jl-17) h the penucful. 1\:orth An
lr~ mmr vulnl'r.lbk to t':>.Unnton whl'n thl' 'ea rcrreats) of I.'Xtlllltl011\ .1 t,IXOillliiUlartdJct? [\'•111111' J ::>uubs~. K.
KJufinan. A.J .• Knoll. AJ . ..,cnuiJ1.1tnv. M.A t'C .rl. (19lJ6) '24H-5 I gc:nlo~c.tl
lntt'b'1 .Ht'd tlll<liHhtr.ltlgr-.•phy nt the ProtcrozoJc- Prothero, I) R . (I t)~lJ) Stl'(lWI~C t'xtincriom .md elm hfml>~ll)?ir ·
Cambri.m bnund.1rv 111 tlw "e'tl'nl An.1bar region. dc:dinc during the: brer fou.·ne Jnd Oligoccnt", it1.ll Sulllm M J
Northcrn ~ihcn.l <;,.,,,,,!!,.-,,/ .Htl!!•'::orc, 133, 509-33. Extirwit~m: flr.•a•sw~ nnd EPidt•nc•' (ed. S.K. Dono1 tlnn.Hk c
Knoll , A .I I ( I'J%) I hughtel ot ume. /),,/rc•hi,•logy 22, 1-7 Uclh~ven, London. pp, '217 .H. ( ;...,,)\? y 1 ~
(updJt.: on Ptl'l.llllbri.m C.uuhri.m bound.uy; excel- Qian Yi .tnd Uengtson, S ( 19R9) P.llaeontology md b1• 1\t <lilt'\ , S.N
lent) trJtignphy of thl• E.-trly CJmbnJn Ml"t~hucuman '~ .unhri.m
Knoll, A I I .lnd l l.tr~honm L ~ (llJ74) Precambnan 111 Yunnan Provmce , Sourh Chin.!. J.',usrls rmd Stmtal' "<'lt'lltt' 27(
euk.tryOIIl ur~.1111'111'' .1 rt".J\\e,,menr of rhe evideuce. 1- 156. Suu 1ht:! , P .
~flt'll<l' 190. 'l2 4. (C.t'h dLIUbt upuu fL~>>il euk.Jryonc RlmJd-'>. D.C .md Mof\e, J.W ( I'J71 ) Evnlution.ll)' f't•ssil Rc1
.llj;Jt) ccologJ1. "J..'Tlltilanu of oxygc.:n-dc:tic:iem 11 Bdh.lven •
Knoll, ·\.11. .md C.tld,·r . .., ( 1'IHJ) M•• rflhiOtJ~ of the bte bJsms. Lt:clrt~rtl 4, 41."\ 2H. (W J.' early evoluoon lirn!tC \-.1\'r.stcr-Ur.
Prt'l m1hn.1n lt \"I.. I'• f-onu.lllon, Nord,m~tlandet, b) 0, detiocn• y ?) p,,.,' t'dlll.~l
"'valb.ml. P<1/,, ..,,,,,,,,~)' 26, 4h7-1Jh. (01\'Cf\t' prokary- Rong, J.-Y and H.~rpl'f. f) A. T. (19K9) A global w:u~ l hemica
om Aon) m of the I.JCC\t OrdonuJn I hm annan brach10poJ :r :app.m. H .
Knoll. .'\.K ..111d s,H'It, K (19!(7) Mu:ropabeontolo~ 11;'1~. fftiiiJ<I< li<ll" •!/lift• /{,l)'clf "''01'1)' .,( f<flll/111~~/1. CIU\C:'i of
.ll ro·' llll' Prt·, uuhn.111-C.uubn.Jn boundary in Smut·.• 79. 3!n -W2 lmrm~> •!I
SpitZhL·r~.:n . .J.•Ilnrcl/•!1 1'.•/"'"''''•'l!l' 61. :i'l~-IJ26. Rounov. A. Yu and Zhur.wll'\' A Yu 1'19~ S1hc:r .md
LIC',m, C.!{ ( 1'17~) Bll'g•·olog) of the late prc- lower C.l!nbn.ln fo"tl rt•n,rd of the SC)VIl't Umo spcn.ll Pa
Ph.ulerntol< B(ck Spnn~ Dolomite of eastern Onl?m .wd I:arly F"••lmro11 ••f the ,\ lrltl::tl•l (ed~ J.l I l I "'re nee
C.tlil~mti 1 f<•wn.r/ N fl.rltwrc.•l.•~!~ 52, 767-92. and P.W. Srgnor), l'lenum Pn•s,, New Yorl I C'tt ht·rt. <
M.1rguh,. L .md '\ IL Mcrwmn (I 'I'll I) M,1magc of com·c- London, pp. 205 2H'2. 1'.1/,rc cibio/.
ntl'ntl' !'ltt "'"'"t'' 30, 31 h (Luk.tryote ongins). ~unucgar, U. ( 1'192) l:.volunnn of rhe e.ulll'~t .1mnu. 1h~y.·r. C.\l
M.miH:w,, S.<... .md V Ml\\.trzhev,ky ( I'J75) Small shdl} ,\/,!jcJr EPwts 111 t/11• l/i.111>ry ••( Lrfi' (ed J W . Scb· luntm of
fos"l' of l.ttc l'rCL.Ullbn.ul .md l',lrlY C.unbri.m age: .1 Jonl'\ &. Bartktt, U,l~ton, M.m., pp. +l 62. -l~H-6 1. (•
rCVIl'W of rt·• ent wnrk l)ll.u-t.'li)' }<liiYI/<11 of tltr Gr,J/O,'o!iml Srhop t~ J. W. ( l 'lhH) MKmAor.t nf the tltttcr Spn r n<' kt·r, IV
Sorh·t)' il( L!incl.m 13 1, 21'!11 JO-t (MknJI(mJJ evidenn• Form.ltion, l.1te Prc••1mhn.m , Central A ustr,11i.1. Jmrr ht.,lllld.Jt')"
un L'Xp lo~IOII of lik 111 l'.ulv C.unbn.m) <!{ Palt•tml••lv,~)' 42, 65 1- H. ( Gl!'m•/)(>tryt/i(llr - po<11 nutncnt ~
M.1xwtll, W.J) (1 1JH'l) Th,· mHI PrnniJn m.1~s cxrinc- cu k.i ryorc ?)
tlon m ,\/an I "irrdic•rrc l'rt•<•'N's '""' h•irll'llft' (ed. 1\.K. Schopf, J.W. (I'JIJ2b) ' I ht· nldc~t t(mils and wh.1t rh
Donov.m). lklh.tH'II, London. pp. 15.2-73. me: .m, 111 AlnJ••r Er•t'llts 111 t/r(' 1list11ry of Liji· (rd J\i
McC,hl·c. c; (I'I!W.1) C.11.1~1ruphc' m the lusron ofhtc, in Sclmpf), Jone~ & B.irtk·u. Bo~ton , Ma,!>.. pp. 29-6~
l:r•lllrtti•'" '""' t/r,• h•wl Um•rd (rd' K.C. Alien and ~chop( J.W ..md (khler, D Z. (J'J76) How old .let
D.E.(, Bn~">), lklh.lvl'n. Londt>n, pp 2<.-50. eukaryot~">? Sticmc 193. 47 9 (Defend!' PrecJ.m
Ml Gh..-,, (; (I 'IH'Jh) I he h.t'lll.ln FJmenmm ,·xtinc- algae J\ l'Ukaryuuc)
non l'Venr 111 ,\/.1.•.• 1:.\llmlh•lls: l'r••cc<.•o .md fl'idnru Sc:J!Jch.:r. A ( IIJH'J) VL·ndowa Orgamsnuc lunstruc
(cd .... K l>nnov;~n). lklha\l'll, 1 nndon. pp. I:U-51. 111 thC' I.Hl' ProtL'flllOIC h•o,pherc LHhtlltl 22. 22'J-3
Mtlct•d. N 1'1 er/ (22 .llltlw~'>) (1'1'17) The Creuceou,- St:ti.ICh.:r. A. (1992) Vc:ndobtt\lll.J and Ps.unmocvr
TCrtl.ll') hllmt lf,lll\ltlllll.)<•rtrll.r/ <1(//r, Gt,•l~lml S,>crrt} Jo,t comtntlllon' ol Prcc,unbriJn cvolunon. ]t>rcm:l
IJ/l..c•lldl'll 154 ~h!'-'l..! rlu Cr.•!.t:erMI.'\,l(wty ·~fi.Nrd,,, 149. (,07-U.
Bibliography 81
P~'-"kl,j. (l'!K4) A kmenc nwdd <>I Ph.uwro7oi, raxo- biominl'r,ilisauon. }oun1t1/ <1f tlzr Gt·ol~ttlfcll St>nl'ly of
ncnuc dswr H) Ill. Pmr-Pal.1eo2'm, t.muhc, .md lliJ'>> Lmdorr 149, 65~.
(:tttn•tiom Pa/ut/•r••lt>gy 10. 24fl-f>7. Upchurch, G R. (1989) Terrestrial environmenrJI ch<~ngc'
Srpko•ki, J. (I'''JO) Evolunon:uy faun.1s, Ill p,,f,,,.,,,l<lf,,\1}': a and exuncuon pattems ar the Crcuccou\-Ternary
'f'Hirt••is (rd' D.C.<.;. Ungg> .md P R. Cruwther). bounciuy, North America, in ,\1ass Fxtmtlit>IH. P~t~rrm:s
BIJ,kwdl. Oxford. pp. 37-41. and Evid111cr (cd.~.K. Donovan). Belhaveu, Lt'ndnn. pp.
Scrku1k1,J , Uambach. R..K .. Raup, D.M .•llld V.dcnunc, I'JS-21 C1.
J ~·. (19HI) Pluncrozoic m.mnc dJVcr,ity .md the fm~il Vid:~l. G. ami A.H. K.noU (1982) Radiatit)llS .md t·xanc-
rc..1rd \'.llurl', London 293, .u;-7. (Ch.u1~e in divcr- uom of plmktou in the late Protcrozn1c .md c.uly
•uy thmu~h tulll'>) Cambnan. Na111rr 297. 57-60. (Early lmtory <)f JlgJe)
han, P.M . (I 97 3) The rdattnn of l ,u,• Ordovtn.m Wade. M. (I 968) Preservation of soft-bodlt'd .uumah in
£l.!cunon 111 rhc Ordo\'lcian-SLiun.m changeover m Pr.·cambnJn \Jnthtone ar Ed.iacara, )outh Amrrali.1.
i'lorthAillt'rlt.lll braduopod i::IUilJ,. u;t/rm.l6. 147-84. Lttflaz,t I. 2JH-67. (Why the Ediacar.1 f..mru w." pre-
:nkm, K (19H<J) Btonuna.l!J,Jtmn 111 the contexr of served)
~~~~~~,tl nmc. Tr<111S·Irtrom <I{ rlrt• N_,,y.zl 'iNirry ~/ Wade, M. (1972) llydrozo.r and Scyphozo.1 Jnd other
c E.lrrrl·u~ll l..rrr/r .v,·zut•s 80. I9,"\-9 medus01d' from tht• Prccambrian Ed1acarJ f.Hm.l. ~ourh
1111, MJ 111d Rutfcll, A.H . (19')0) Synduoncay of Austraha. Pn/nccllllrtl<l~Y 15, 197-22'). (fethnu:.ll
I, ,hmalll dtJugc Jnd cxuncnons 111 the l atl' Tn.1s~ic. descriptions)
(;.oilt'J!Y 17 2h5-H. Westrop, <;. ( 1989) Trilobite m.1s; exnncuons nC;lr the
lunlcy. S.M (I 1)7(,) Fms1l d.lt.t .tnd thl· Pn:t.unlm.m- Cambrian-Ordov1ci.m boundary in North A111crira, 111
( lmbnJll •·volurion.try rrans1t1on. t lmmcmr J••rmr.tl of i\ltlu E\tillrti<lrrs: Profrssrs a11d El'rdwrt• (cd S.K.
:ltrnr 276. 'ltl-76. Donovan). Uclh,tvcn. London. pp. 89-103.
th,·r. P ( 9X9) Prc-mt'tJzoJn lilt-. 111 I :..,,flllwll !llllitlzc Whmaker. R H ( llJ69) New concepts of k.mgdorm ol
1.1 1 I Rw•r.f (c:d' K.C. All.:n .111d D.E.G. Bnggs). org.11mm,. Sczmct 163. 150-60. (Five 1Jngd01m)
e Bdhl\CO, London. pp. 51-72 Whittakcr. R 11 and Ma.rguhs. L. ( 1978) Prntl\t cl.l"lfr-
i .:ntcr-Bmlky, P.C. (1975) The w.trch for prorolifi· cation .1nd tht kingdoms of orgarusrm. Biet>)'>l<ms 10.
lmttdr11~< ••} t/r. RoJ••tl S,>urt)' B 189, 213-33. 3-18
!- t hcnur.1l and eJrly bJOiugJcal evolunon) Whlmngton. II.B. (1966) Phylogcny and dmnbution ot
.1- "~" ·H. (1 1182) Extinction or \UfVIV,tl: sdect1v1ry .md Ordovici.mtrilobites.Jauma/ <!fPa/eo11tcJI11,11y 40, 69h-737 .
111 ollllr' of Pl1.1nerozoic crisis. in C.co/cJ~Ifd/ IIIIJtllfcllit•zts of Woe.,t:, C.F..md W:ichtcrshaliscr, G. (1990) Ongm of
lmr,w •!!" Lrw· Astc1oids 1111d C.tr/ll'IS VII lilt' b~~rllr (eds L. life, in f>a!.tt'ctlri,JIO,'ll': a Synthesis (ed~ D.E.G. Uriggs aud
~1lm ~nu I' Schult:z). Gcologttal ~odery of America P.R. Crowther). 131ackweU, Oxford. pp 3 9.
~pt:Lilll'apa No. 190, Gcolot.'lcll Soc1ery ol A men ea. (Valuable, up-to-date symhesis)
1\l'flll'e. KJn . pp .:?.65-7(, Wolfe,J.A .•md Upchurch, G.R. (19H6) Vcgetatton. cli-
rtt:hcrt, C ( 14!-18) Exnm:uou' .111d exum:nons mate .md floral ch.mges at rhe Cretaceous-Tt.>rriary
l'ill.u ·h~/,~y 2 . .j 11. bound;~ry Nature 324. 148-52.
Ill rr, C.\\' ( 197'J) 13tologtcll buUdozcr. md rhe t'\'o- Y ochelson. E L. ( 1977) Agmata, a propo)cd cxuncr phy-
Q. '"11 nf nunne benrh1c cnnuuurutil'' "-<rma· 203 , lum of c.uly Cambnan age. Jmmzal •!I P.1lt·mztol<l\1)' S 1.
'1'-61 (Ciwt~·· in h.!bitat through ume) 437-54.
• lrr, M.f. ( 199::!) The Prccamhnan-l.ambna.n Yochl'lson, E. L. (19!:13) Sallt'rclla (E.ll'ly Cambnan;
~1und.uy . \C,l\\.,UCr chenu~trv. oce.m t rrl ul.mun ,md Ah'luata) from the Scotti'h Highlands. Palal'cJIItt>lc1,1/Y 26,
nutm•nt supply in met.lZO.tn evoluuon, exuncuon md 253-{)().
ICY
w
the
]~ll
ion
I)
,b.l
•I •'/
PART TWO
Invertebrate Phyla
!Ill 'PC
rcg.trtk<
of ( dl,
I here 11
o( .1 gd
.lllfl llll'l
p arazol
proto7t~
tllcy, n
1 he ~
fi>r nl
morpho
d p nch
r marka
pongc
m o
r move<
JU t hkc
1rc only
Spon!
1re all t
pongc I
'' uh ,1 <
top vi.1
~pong( 1
\\htch I
chamb t
The-se
c h oan o1
the ch:m
1 xh.1l,
d\lt)'. 1
mnt
I urthcm
Orthocone cepholopods cut tongentiolly, showing intemol septo ond siphunculor structures, in red Ordovicion limeslonel lup bet
Sweden Pointing by Roisin Moron, originol specimen in the Jomes Mitchell Museum, University College, Golwoy, lrelond I die
Sponges
J'i tht• Choanoftagel11da art· m form \'Jrtu,tUy Jdt'IHI-

cal w1th 'ponge collar cells. ~ome pbnkconil· m.mnc
choanoA:~gclhds. like sponges, constmct b.l'h·t-like
r sp110ge~ are mulncdlular orgam'm' but are not cap\UJc\ Of geometrically 0Tg<ll11Zed rods Of ,jlic,J -.1
ll,'1Tdrd a' metazo.1n~. 1 hey have only .1 few rypes further mdic.ttion ofbtologJcal afrl.nity.
•I ,dJ, Jnd rhe~e are nm really organized 111 tissues. £;u: h colbr cell is a small globular cdl With ,t
T rl' 11 Oil nervou~ ,ystem. ln many way~ lltt'Y are cylimlncal colbr projecting from it, wmpo~eu of
a brrJde of org<llllZltlOll 111 between protozoans fim: pwudopodia. This collar encirdt·~ J 'oht.lrV
m~uznans: ht'IKt' the\ .1re somenme~ known a~ n:ntr.1l flagellum. In hie .111 the tlagella be.tt contlllll-
ptmoan~. App.1rently rhey were derh·ed lrom a ally, the tip of each wlmlmg in a sptral motion. The
ozo.t. anet·~tor hut wen.· ,m eYoluuonary blind currt·nt generated by 1ll tht· AageUae draw' '' .ltl'f m
. , not ance~tr.tl to 1ny metazoan. through the ostia to the chambers and out to the
lh~ tint' structure: ot sponges can often be used n·ntral ctvtty and the oscu lum. Particulate org.1n1c
• •J~nt1i)·ing tht· sp1:cies. but the external nMttcr then .tdhercs to the ~ncky outside of thc coi-
rpholn~:,'\ may vary w1thm Eurly bro,td limit~ !Jr, wh1ch 1ngcsts it. Though the whirling ut l'.tth
niling upon tht· cn\'m)nment. Sponge~ have fiagellum 1\ comparatively \low, the combined l'IK•ct
ubi~ pO\\ t'f\ of rq~eneration. JnJ a living of all the flagellac operating rogether prodm:e\ ,m
msr lqUeezeJ through ,l Silk net \\Ill re-form dlicient though low-pre~~ure pump wh1ch t.ln pa~~
\'11 the other s1de But md1V1dual celb through the ~pongc every m mute a voltune of w.Her
IOI'~d from sponge~. even though they may look equal to th~.: sponge's own volume. Sponge~ gcncr-
t l1te some prowzo.lll\, will not hvc long: they alJy have other kinds of cell~ than d1c collar cell~. In
1)11\y vuble a~ part ot the 'ponge. one major b'Toup, Subphylum Gelatinosa (fig. -lA),
Sponges are genn.tlly ,t•ss•le benth1c .lllllll.lis and d1e outer surface is covered With flattened epithelial
ill idter fecdm (Bt·rgqul'•t. 197H). A typical cdh (pina cocy tcs) of ~ h1ch those in the pore
~ tF1g. -l.la) h.1s .111 upnght bag-•duped body rcg.om (porocytes) arc perforated and can tlo~e oO·
J u:ntra.l c.Jvity (paragaster) opcn111g at the the pore\ by contraction tf necessary. Each pororyte
vta .m osculum . The outer sur6ce of the is stimulateu to dose indivtdu:llly by the pre~en.:e of
'{:.e I> pcrforat~:d by numerous tiny hole~ (o s tia), noxious substance~ in the watt·r; the condttl twn of
h IL·ad to inc u r re nt canals and thence to such ~tunulation to nc1ghbouring cells may take
cbmbcrs \\ itlun the ~ponge body. place, but in the absence of a nerve net ~~ VC!] ltm-
These dt;~mbef\ .m: lmed by colla r cells or ited. Then: are also amoebo1d cells (amoeb ocyt cs)
dloanorytes (Fig. 4.2). c.Kh of \VI11Ch faces into wh1ch \\andcr through the <,ponge body .md tr,lllskr
dumber. noun\hmcnt from the choanocyres to other p.ui~ of
h.1lant passo~g~:s k.1d trom these to the central th~.: sponge.
1 • fhc col1,1r et· Us .lrt' the mmt Important ele- Mmt \pongcs have a ~kcleton; this may stmply be .1
1:1 m the or~Jnit.ttion of the ~pongc. colloidal Jelly, but in mo~t living kinds it COilSISts of a
crmore. tbe~t· cdJ, dcmomtrate the rclanon- homy material (spongin) or of nlcareou~ or stltccou'
'es from tv.ecn ~ponge' and protozo.Ul\, "tnce the sin- sp icules (Figs 4.6. -t 10, 4.11) or of both Some h\-
C'd or sometime\ colomal protozo.ms known ing ponft'ratlS (<>clerosponges) and several fos~il group\
86 Sponges
F1gure 4.
arrows
(choon<K.i
la.tl ope
lcucon
JIUillbcr
ccncr.J
ponge~
(a) anunals
uons m .
uvely ~·
ycon a
bcrs ex
and gro
rh en
I \
'
ot whtcl
tcr) 1 h
tolcranc
\.,.,/) m the
siltceou•
sull wat
the cxtt
(b) Asc:on (c) Sycon (d) Leucon (rhagon) non.1l n
t'};t'urret
Figure 4.1 Elements of sponge morphology· (o)leucon (rhagon) type showing passage of woter currents; (bHdl ascon, sycona
ltl"tswip
leucon grodes of organization [(a) based on drawing in Marshall, 1978.] ~talked
~t:tlk .ltt;
111~ thro
h.l\'1.' J c.dt'.lll'llU\ 'kdctun 111 ,addition to the 'Ptcuk~. ~omctimc' 'Ptlllgt'' Ll\e 'hdl ucbn,, und or even Sronl
,md the c'I.Unn .Ill h.lt'ocy.uh.lm h,1ve ,1 ~kdeton con- 'PKUIL''> of dc.td 'Ptlll~l'\ w strt'llt,rthen their ~kclcr dtfl'CtiOI
~JstJn~ of l.lluum cuh•n.He alone ~picules C<ln be: Tlu:rc art thn:l' grade' ot orgJnization in ti1 one ~tt.lt
to~'thn·d, t''Pt'n.lll) wlll'n they are united ~o a~ to 'ponge,. In tht· 'implt·,t (ascon) grade (Fig. 4 11110 the
hold rlw both n! thl" 'Pllll£l' to!tether ,tfter death. the ,,K-hke budy i' llll'reh ,, ...inglc chJmber 1t (Ftg.
1 hm c:ompktt' or p.trtial 'pon~e bodic:s. m.m of "11h cho.tnut-.ll''· with .tttl'lld.mt l"pithdial cdh Villllllg (
'Pit uk' <W ''ol.ned 'Pilule, .m: tound in rocks cxtcnd- amoebocyte' Sycon 'pongc' (Ftg. 4 .1c) lm \\:tter. A
mg b.Kk tn tlw <..·.unbn.m .md po,sibly fi.tnher. numbt•r of groupt•d ,t,nm-hke chamber... \\ ith a c.nn(1ids
Classification 87
la) (b)
F~ 4.2 Wall of chomber, showing incurrent pores with Figure 4.3 (o) Life form of o stal ked, still·woter sponge and
arrows mdicohng current directions ond collar cells (b) life form of o 'rheophilic' sponge with a wide lateral osculum
W.00nocyte!) directed owoy from the current. (Redrawn from on illustration in
Carter, 1952.)
~I uprmng, but thl.' v.t~l lllJJority of ~ponges are of

leucou (rhagon ) typ~: (hg. 4.1 J,d), in which a
SPICUle ;/riR ollllOI'bOCVllc!
nbrr of ~ylOn-hkc: dcm~:nts opc:n into the large

Mttral CJV1£y (p.ua~.l\tcr). In the larger modem
ngcs, ,hnmps, ophtunmh or other com.mensal - rncurrent JJQre
JllllUh m~y hvt• 111 the p.u.tg.tstc:r. Jnd ~uch assona-

nll) bt• found 111 lO\\tl lonn'>. A.,cons are rela- prnJCOCVIO Lwers
dv mall and not nnu:h l.1rgcr than lO cm high.
'COO and lcucon spon~t·s wath their folded cham-
:m ~xhabu obvtou'l) grt•.tter filtering efficiency
ni gro" ro .1l.lr~cr •azc.:.
There art• ahout I500 gcncr.t of modem sponge~.
\lhtch ~ome HO% .ut• nunnt• (the re't arc frc<>hwa-
Figure 4.4 Wall strv<:tvre of on odvonced gelohnoson sponge.
The 'hallow m.tnnt• gt·m·ra 'how a remarkable (Redrawn from Reid, 1958-1964 )
.rJnce of mtertad.tl o.:ondattons Sponges abound
the dn p \t'.t, cspt•uall} the I kxactinellida or
ceon~ gl.l's ~ponges, whtdl .trt' confmed to qutet,
\l,ltt'r<;. Spongt' morphology as well .tdapted to
th cxtcm.11 cawtronnh.'nt, t'spcnally m such fimc-
~onalnt't't'S\ltlt'S .ts the scp.trauon of tncurrcnt and Sponge taxonomy at the highc~t level ts b.tscd upon
turrt•nt w.tter (Htdckr. 1937; l lartman and the soft rissues, though only the skeleton 1s pre-
, ond Rct,wig. I971). 'I hm m.1ny dl'l'p-w.ttcr sponges are served in rhe fosstls. Two subphvla have been dtstm-
1talked, the inntrrl·nt w,ttt•r commg in near the guishcd (R.cid, 195H-l904) on the stntcture of the
lk attJdmtcnt .md the l''<t'urrcnt wastewater pass- ~ponge wall. In Subphylum GdJtmosa (Fag. 4 A) the
through the o~ntlum ( l t~. 4.Ja). omer epthelial layer of flattened panacocyte~ overlie~
L the Sponge\ hvmg m ,h;~llow \\otter 111 wh1ch current a gel.ltinou~ mtddlc layer (mese n chyme) in which
·ons. ~lfJOm .tre ron\t.l!H mJy havl' the osculum down the sptcules arc 'ccrctcd by scleroblasts and
ving c ,,Je. 'o tlut tht· bnudcr t:m-likc surface taccs wherein the .1moebocvtc~ wander.
. I b) o thl· rurrem .md the l.ttt•r,\1 osculum Jway fi-om The tnnenno~t layer. 111 'pongc~ of J\COn grade. t<;
inc:d Ftg. 4.3b). Tlm not on!) make' u'e of the prc- that bcJnng the cho,HH)l ytc~; 111 more advanced
and ' ,g (Urrcm but .tl'o pn n·nts reme of excurrenr grade<> the~e arc lll\.tglll.ltt'd wathan ch.tmber;.
IC I li r A .,,trJllel wtth tht' " fC.mnd in brvozoam and The other ~ubphylum. Nuda (Fig. 4.5). has
_en- T.lOitk nettber ptnacocyte !Jycr nor mt:~enchyme: rhe
88 Sponges
PHYlUM PORIFERA (l .1111.-Rn.) MultllellulJ r anunal~ ''

choJno.:vtc' .mu .1 ,lo..d.:tou of '1'011!(111 .md/m tJk,treout
~ahceom spanal.:~ anu/m 111,1\,1\'l' ,,uhnn.ltt.
SUBPHYlUM I. GELATINOSA Porift·r.t ,-.Hh pin.KO")
and mcwn< hymc: dttl.tno.-vr•' on tht• umer llll"'ench~
,urf1cl'.
CLASS l DEMOSPONGEA l..un.-Rn ): Gdatmo;~
leunm gr.u.l.- "1th ,,h, <'<)11' ~p~<uk' md/or 'pongm
\Omt•umc' "•rh torngu mdm1011~ \pit uk r.ty' U>
dl\'l'rgc .u hll or I.:!W ll\ 111~ >ekro,pon~t'), clut·tc
mo~t ,phuu to7oJn' .md mo\t 'trl'lll.U•'POrOld>
mcluJcd here
CLASS 2 CAlCAREA (CAlCISPONGEA) (C.un -Rr
Figure 4.5 Wall strvctvre of o hexoctinellid (Nudo) of leucon Aseam. \yrnm or kunms \\ 1th a 'l.ckwn uf .akuc
grade. (Redrawn from Reid, 1958-1964.) spicules A tcw 'Ph11llt\lZO.Ilh ,1nd nwnutopor
bclon~ fO tlm llJ~,
choanocyres ,ue borlll 111 ,1 net\\ork of \yncitial fila- SUBPHYLUM 2 NUDA: Ponll-u wathom pm.tuKytc ccU,
ment\ (t ra b eculae) but .1re lJkc.:wi'c organ ized in mcscnchynu:; dH,)J illll ytc' '<'I 111 .1 netwnrJ.. nf J'rowpli!
mic tlu·t•.Jds.
chambe.-.;. CLASS HEXACTINELLIDA (HYAlOSPONGEA) 1
The: lll.1JOnty ot hvlllg Pori fer..1 luve :l skeleton or C.un. Rt' L ): NuJ.1. mu.ally lc: ucnn'. Wlth ~plntbr n
spicules .•llld unti l rt'tl'JHly it \\..1~ imagined that the d•vc:r~'lll~ .11 'Jil" Spatnl.u \lo..t' lc:tnn nnlv. (a)
only ~pon~c~ in the f<w•il n·conl were likcwi~c spic- INCERTAE SEDIS. ARCHAEOCYATHA (1 - M CJm (d e
ular. Smcc thl dm OH'!'\ of Rl'Cl'nt ~dcrosponges, C.1lc;1rcou\ 1wrlnr.1tl·d ~kekmn nf two 'nested' mvrr
with both .1 ~pit ubr .md .1 l'JicJrl'Oll\ ~keleton, the nmtos JUmt·J b)< r.td•~l p.1rtH1<1lH. 1\ few \phmetozw::
.md ' tmm.Hnpnmtd, .lrl· mdudcd here
claSSlflnnon .md n:l,ttHm~hip' of fo~sil and livmg
\ponge' h.t<; bet•n drJmJt1l'.lll)' n·' 1sed. more ~o. per-
haps. than Ill any othl•r fm,il group. It lu~ nO'\\ been
rerogmZl·d rh.tt tlw 'kdl•ton o( \tromatoporoids,
rhaetend<; lnd sphllll tozo.tns hkewi'e possesses both
sp1cuks and cak.tn:ous m.m.·rul. Though the arago-
mte skekron " often rl'l'f)'t.tllizcd, obscuring the Spicular d e mosponges
pnmJl)' \tructun:, .lttd l'tching and SEM microgra-
phy ha<; <.:n.lbled 'P" uJe., to be detected. These ,"vtost t(w,tl dcm mpongc' are repn:~enred
group~. togl•ther wtth the enurdy calcareous s1hceous .,piculc:s on I}, the ,kdcton havmg c
arcll..leoc>.Hhtd,, .m: 1\tl\\ .111 regarded as tnte lapsed. Such ~picu k<. !ll.l) luvt: etther one single
spon~e\. lt h.ls .tl\o bcnm1e dc.1r tlut solid, calcare- (m o naxon) or four rJy<, (tetr axon) drverging at
ous ~kdl'ton> dcvelopl'd independently within at or 120° (F1g. 4.6r).
least two ot rill' mam poriteran classes (Demo- Anctent ~pt culc~, l'VCI1 ur
Cambri.tn age. CJJI 110.: F1gurf' 4.6
cunol ~ and
sponge:. <1nd (\tlc.t rca), and that 'stromatoporoids', tmlly be rehtcd ro mo Jcrn t:unilic~. Only in Ore~
\ponge (I( C
for example, .lfl' tbm bc~t regarded as 'grades of Litlmtida (Cam .-Rcc.) art: the spicules of dttferc ~ponge (~. C
orgamzation' r,ltht•r thJn .1s of 'eparatc taxonomic form. Th e~c knobbly or tubt:rcular desm as (f
rank, and the) un be .Kcommodatcd within mod- 4.o::~,b) .tr<.: oircn 'o int~.:rlockcd that the skekt
cm sponge d.ls'c' (Zhur.tvlcv t'l fll., 1990). Because holds together .lftcr dl..'alh. Lithisrid genera an.' (0
of such convergent evolutiOn. many problems mon fro m the jurJ.,,ic onwards. and such types Sclerc
remam m rrymg to produce ,, ~.ltisfactory classifica- the Crctan·om S1plumi11 'ho'" det.1ils o f the c.m
tion of the Porifer.l A provi,ionaJ dassit1cation ., chambe r~ . parag:tstcr .tlld mculum when sett1on
given bclnw: thl\ no doubt lllJ} well be modified m Cliomd \ponge' <.an pt:nt:tt,lte mto hard shell
the future limc\tolll' nutcnal. 'pn:,,ding out below the surfr
and sccrering a 'PICubr ... kdeton wtthm the lh
Such \p ongt'\ contribute b'~'t:.itly to the destrul l g•>llltll
of hard ..,hdl tt\\lll'\ ,md thus arc ot ~ome gcolo gm and 'I
imporrancl. ~~~~ •>fbut
Class Demospongea 89
[
0
(c) Demosponge spicules
)f
;-
(a) S/phon/11 (b) Uthlatld desmas

(demospongel
(e) Peronk»ll11, with spicules

(f) Leucendr• (demosponge)
or 4
r9Jre 4.6 Demospongeo and Colcoreo: (o) Siphonio (Cret.), o stalked example (leh) cut in section (right), showing the internal
fer O.'lOI1ond porogoster (x 0.5), (b) lithistid desmos (x 25); (c) demosponge spicules (x 30); (d) Rophidonemo, o Cretaceous calcareous
~nge (• 0.25); (e) Peronidella, a Cretaceous calcareous sponge (x 0.5), with 'tuning fork' spicules (x 30); (f) Leucandro, o demo·
Ill
~ (< 0.5), with tetraxon spicules (• 30). (Mainly redrawn From Hinde, 1887-1893.)
ig.
Oil
11-
Sclerosponges engulfs the spicular \kdeton dunng growth. The
spiculcs then bccomt' mcorporatcd 111 the calcareous
me 13 ll'v1ng 'Pl'lll'' of 'umtllinc \pongc~· have matenal.
or .n de~cnbcd 'intt' I 1)70. inhabinng 'ubmannc The outer ~urface of the hvmg msut' mvestmg the
lCC es JnJ other d.1rk t'l) plk rct"c<.'c' (H.1ronan and skeleton ~~ p1erced w1th two stzcs of pores: small
dl JI)R'~u. l<J70). Tht:\l' '<..'nctc .1 m,l\,ivc, encrustmg, incurrent ostia, Jnd l.1rger. w1dcly ~paced excurrent
gruuu, skclcwn in add1tinn to a 'kdcton of ~pon oscula. These J0111 With a system of radiating
md 'Pil ulc~ . The t:akarcou' ~kekton may con- star-shaped canals, often nmnmg 111 grooves wuhin
ot both .uagonitc .111J c.1kill' ,u1d it progrc~sivcly the outer layer of the ~keleton. Water 1s sucked in
90 Sponges
(a) (b)
Figure 4.7 Chaetero5 (o) woll structure and (b) drawing showing appearance in thin section, w1th sprcule pseudomorphs (Based
0 crowing In Groy, 1980.)
Figure4.8
lured port
Stroma~po1
through the mCI.t h} dw.uwcytc~ lying in linked Stnllnatoll.\tttclfa thcrl' ,tre .11\o ~picul~.·, (Wood .\l ( 1) (BasE
dumbe-r' ,tnd ~·xptlkd thmu~h tltL' cxcurrent O\cub. Rcitncr, I<JHH), but lhL'St' .trt' nf quitl a ditf\'t( Invertebrate
Fmstl sdcnhplllt~t' .m: !...no\\ 11 lrllnt rh~ Tcrti:111 kmd, <llld of ol Utffert•nt t.'T'OUp
l h,tl',ll [l'l'l\lll
of Ca!ttorn1.1 . dcmospongc~. lntkcd, 111 lll,my wap tlm gl:lllll
intcnncJt,lt\.' between duL'tt•tid' .md \trunm h:~pcJ l'X
porord~. Frm n \Ul h 1.:\ tdcm L' . it 1' no\\ clcJr wspnngcs
Chaetetids 't:luctctld'' .1r1.· polyph)k•nt:, .111d that tlu, 5'Tadr The str
orgamz.•t•ou •~ lOllVI.'rgt•nt. h,l\'ing ari'>en more 111 't:rucal
The.:,~.· .m~ l.umn.tr, hcmt\phcnl".ll or em·rmting once in geolngll,ll hhlOf)- . stght tu bt
org.tlll\lll\, nt{en ,,f qmtl' l.trgt• ''7t'. Thq were long ullll · tntl
bdinTd m b~.· uhul.ttt' ,·nnJ,, bm rlwy ;~re now rec- l\'PC'o \\.'1'\' C
ogmnd ,,, 1. hhd} rd.1t1.'d tn ,cJcru~ponge'. Strornatoporoids

f/cll/lcl hlpcl
Sp~tuk·, "t'rl' ll!"t dhl'lWen:d m tht• Carboni- lOCnostet
tcrou' Clr.tt I I'll'.$ (hg. 4. 7) (Cn). 19Xfl). Srrom.ltopormth .trL' l\\k.tr~.·nus m,1sses of ho\\' sto1
l11 thi' g~.·nm tiH.' tnlmlll.'' dt,pl.l) rhythmic .md \trucrur~.·d m.u,•n.ll t(·mnd in tr.wcr..cd I
~rrowth batllk .tnd on 'ettitiiHng the)' 'how an ~cqucnccs of C.lmhnan tn Ohgoccnc age and dtltcrt:ntu
trrq,'ltl.tr to <oubp,,J~~tm.tl n·llul.tr 'tmnun.>, the celh n.1nt 111 the l.itlunan .md Dt'\'0111,111. Pal.1eo7oic ul.ttl' pattc
bcmg up tu 11.5 nun '''rt"' md known .1s c,1hclcs. nutoporoids, espcu.tlly rhm;: of the OrdovKIJn ·r here 1
Thl· ca!Jdc '"·'"' \.try 111 thtl"l...n~. ''· tlften extending 11cvoman, were tmport.mt r{'t?f ton11ers. between c
mto ridgl''>, wlu~. h 111,1} 111 tmn dl·velop imo a pscu- common enough .111d h'Tt?ntly \hldicd, thcrr llL'W Jl1atet
dmcptal 'lrmtun• .111d Plt",l\hlll,lll} mto j,oJ.1tcJ c:~l nature ha~ only rt'< cntly bt.>en rlariticd; tb~) "nh the: tl
columm. l'hl'Sl' \tl'llrttll·l'\ tnmmonly t,rive rhc likcwi~e dcmmponge~ wirh bnrh ~p1cuks ,md a,. thl' SllCCe\•
lalidl.' w.tlh .1 ,L,dlop~.·d .tpp~.·ar<lllt l' The spiculcs of carcous ~kckton. L.tlcarcom
Clwctefn arc hnmd wtthm the t.llick walk In thi~ They lw.tr st1mt· rcscmblanct· to compound tJhu luc~ in con
lllJtcn.ll trulll W.lk\ tht'\l' \\ .n, .ll'l' often diai!Cilt'ri- late coral<., siiKC rhc r.lk:trcnus •kdcton is of i Aflinostt
cally ,.hotled .. 1 pror1.'" "hil h h.t, prc,crved the cal- layaed fonn .md formmg rounded m.t\ses or thin 'umc:what
ctt;: 'Pil"Uie, 111 p.Hdlcs. The 'Ptttdes thcm~dvt:~ arc \hct't'· Ol'C:l\1011.\lly cvlmdncll or dt~Cotd.tl (Fig. ~ JllJ more
\tr.ltght or sh~htl) llll'\in~-t ~lmt) Je, up to O.J mm Where the uppt·r \llrl.ttC' 1s pn:scnTd 1t normJ prOCC\SCS l
long. ,Jurpl) pmntcd dl\ully, .md wirh J di~tinct show~ J p.1rtcrn of poh gorul m.trkmb"'· lt .1!-o ~re not cot
ho" ut th1.· pnn.im,tl cnd. The prt'\CIKe of ~uch cal- have small swdhng-. (m amelons) at tntcrvah pt.'l'll'\.
l" Jreom ~picule, ~Cl' Ill' ro 'ctdc thc .ttlinttH.'~ of the trcqucntly .1 mmt dur.tdcnmc t(aturc of L1balud
Ch::teretid.t .md thl'\ .trc 110\\ recognized as being poro1d organi7.\tion, \tdl.ltl' ~-orroo\'e' l..:nown horizontal
rebtcd m \t lnoo.,pnngc.... In the CrctJccons astrorhizae r hc,c .lrt.' \tnkingly sitmlar to the 11hcrca~ L
Class Demospongea 91
(a)
fb)
on
f-gwe 4.8 (o) Stromoloporo (Dev.), showmg lominoe in o froc-

llred port of o colony ( · 0.75); (b) upper surface of Figure 4. 9 Wedge diagrams showing hard tissue in (a)
Stromotoporo, showing momelons with openings of ostrorhizoe Slromolopora and (b) Aclinostromo. (Redrawn from Steorn,
nd I 1) (Based on a drawing by Lecompte in Treatise on 1966.)
!lit l!l•erlebrote Paleontalogy, Port F )
of
IS
ruped t'xh,lknt c.m.ll ~Y'l\.'111~ <)ll the ~urface of ~clc / .....- .....
hat Oc\pongt''
12 EH-u:!H
of Thl 'tructurc of 'trom.unrormd\ may be smdicd
~rrtte<tl and t.lngulttal \\.'dtom. It appeal"\ at tlrsr ~ ~ t
1.111
light to be b, dctim·d th.m m t' nrak but the donu-
JH
nmr stntcturl:\ tfl' \.·kar cnou~h J\ fl.'w ~elected
~
[]
~
j1C'\ ;cn·c to illtl\tr.ttl' the r.mgc m morphology. In
(a) Actinostrom•
111!al.lJ"''•' (Ord.-l't nn.: ltb"' ·V>. 4. JO) the
roc nosteum (rolnnt.tl 'kclcton) in vanr.tl ~cction
·red bows \(our, upright pillars, JOmed .Jt tntcrY.tl~ Jnd
1.\tt: trl\'CI'\Cd by thm hon7Dnt.ll laminae, though the
tnl- diffm~nt!ltcd \tntCflll'l' i' often lmt and only J rctic-

tro- te p.tttcrn of .HI.t,tommiug l'lemcms rcrnJms.
1 tO fhl'rl m.ty .tl,o IK ta bu lae makmg partitions
ugh twecn rh.tmbcr'i. With upw.trd f_rrowth (Fig. 4.c))
t)gt - r11 m.ttcn.tl\ '-\'~ n . ~Ll'rl'tl·d by tht.: cclh in contact
. an· 111th thl up(K'r t m·nmtc·.d ~urClct·, tit m roofing over (c) Strom•topo,.. (d) Strom•topor•
cat- he 'li<H'~'ivc :htrorhtL.tl t an.tk In Srro111atopora the
:Jl m~ou~ nl.ltcri.tlt~ C< llt~truncJ of c.trbonatc sphcru- Figure 4.10 Structure of different stromotoporoids in section
(x 10 approx.). (a He) vertical sections; (d) horizontal section.
abu- [,1111 contact wtth one .motbt·r. but thi~ 1s unusual. (Based on a drawing by Lecompte in Treatise on lnverlebrate
~br AriHic"lltlfll<l (C.un L. C.ub.: Fig-; 4.9-4.11) 1~ Poleontology, Port F.)
, fbt mcwhat \tmtlar, thou~h tht p!llar~ arc c;trJJghter
4.H). l mo~<.· ~lcnda and .JOII1l'd b) horizont.tl radi,tl
n.ul :t\(\'"l \ 111 .1 kind nf l.1nunar m twork. A~trorhi:uc pronuncnt whulat• bt'f\\'t'cn the piiJaf\, com·ex
Ill :I} not common .md Jr\.' .lb'l'llt ~ltogt•thcr m -;omc upwards. In other gen\.·r.t honzont.ll l.tminae, ottcn
.wt! ~IC • \\'Jth cdlul.tr pJrttttom, ,lfl' weU dctincd. These
1,1t0- Llbalud/,r (Stl. Dt•v.) h.h mong p11l.1~ and thm strurtur.ll ckmcnt'> piiJ.u,, l.umn.lte. tJbube.
n as nzont.1l tJbulw .mJ pu"l'"e' r JR' ,1\trorhiz.lc, amorhiz.tc, ctt.:. - .m: t(>Und variously developed in
,t:u- crca' L.r/trc/u.r ( Ft~ . 4.9. Ord.-c.ub.) h.t' \'Cl)' diifcrcm stronuwporOtlh: 't)lllCtimes one kmd
92 Sponges
(a I
osculum astrorhozal canal
flagellated chambers
basal plnacoderm
connective !Issue
(bl
Figvre 4.12 A generalized sphinctozoon sponge, shoW1"'8

structure. (Redrawn From Wood, 1990.)
(cl osculum
~~~s
~ ~
- · J--spicule
.1ragomtc. Some authorities (e.g. ()rc,Jm. lllll{J
\uggested an originally MJgUtlltK -;kdeton "b
u~ually much altered dtfl~renn,tlly b} Jugm-
: \ .· I '
)team attempted to di,tmgui~h prima[} from
nndary nunosmJCtun:s and by 'll Joing he
remove the source of much t.Lxononuc wufu
Ftgurc 4. 11 ~bows the rccomtnlCt~J 'lltl-p.m n
phology' modelled upon that or ltving ~tlcru,rm
~tromaroporoids grow as l.tnmur ~beet~ .Kt
one alter the other - " pattern which lm r
.uncnablc to computer 'illllUiallon, as pix~b
rastcr ,1rray (Swan and Kcro;h.n", l1JIJ4).
Figvre 4. 11 Restored morphology of stromotoporoids mod· ~rrowth fonn~ corrC\pondmg to attu.1J 'trOIIt.
el red upon sderosponges (o) Actinoslromo (Com.-L Corb.); (b)
por01d morphologies wen: produced, '
Stromotoporo (Ord.-Perm.) in which the hard tissue is composed
of carbonate sllherulites., (c) Adinostromorinino (U. Jur.) show· according to simulated seduncntauon pl
ing how spirules ore trapped by sponges which then becomes domal tcnm\, merging and branclung (}1'C)
mineroliz.ed to form o primary calcareous skeleton. This skeleton columnar stacked sbeets wtth ragged margtm
then grows a ~dory ~brovs rom ((b) based on Steorn, 1972;
gou~ m real type~. Cunomly. 'omcal-ba,t•d
(c) based on Wood, 1987 )
phologu:s, easily gc11<:rated by the con
program, are rarely found 111 nature. The~c cxr
of structure develops much .1t the expense of others. mt:nt~ suggest that c:~r h ~tronu ropm01d 'gn•
Sptculc~ haw been found in the skeletons both of unit' wa\ largely autonomom. like each ptxd 111
l.1te M esozoic (Wood, l t>H7) and Carboniferous model, and there i~ no dear rvtdenn.· of centr
~trom.uoporoids. Mmt ol the sp1cules are siliccom, wntrol of growth.
.md thm the maJOnt) ol thew ~tromatopor01ds arc
regarded as demo,ponges. Apparently the primary
'kclcton was of 'P'l uks. Thc\c then became trapplc"d Sphinctozoans
h)' a coating of c;pongm ..md Jt was upon this tf:unc-
work that the cakareou\ ~kclcton (usuall} Jr.lgo- ~pongl'S of 'Phinctozo:m grade have \egm Figure 4
nitic) became secondanly prectpltated. trrcgularly proliferating d1:tlllbl'l"\ arranged rou1 ~urfccn
The original compo~1rion of stromatoporoid Lentral cavity (Fig. 4.12). rovndm
upp~rmG
coc no~tea is unknown: it m.1y h:.we been c.ucitc or fhc walls arc perforated .md m.1y be of COillt' r produc
Class Calcarea 93
li'gure 4.13 External appearance of on unusual twinned

IPOilge. Raphidonemo, from the lower Cretaceous of Foringdon,
[ng~nd 1~ 0.75). (Photograph reproduced by courtesy of C.
C!loplin.)
'C' Figure 4. 15 (o) Port of o skeleton of o dictyonine hexoctinellid,
lS showing overgrowth of contiguous hexoct spicules by o siliceous
s. envelope and (b) construction of the hexoctmellid skeleton with
outer, middle and inner megosderes and scattered microsderes
:- (Redrawn from Reid, 1958-1964.)
1.
r- c;rructun.·. They Roumlw.! rna111ly in tlw late
s. Palacozoic and early Mesozoit. but sevl'r.tl
td Cambrian genera arc knO\\ n, .md .1 Rect·nt fomt.
•d Vacclwa, h.1s ,1Jso bet·n dtSCO\.Cn:d, hvmg m nypnL"
3 habitats. There .1re mml.lnncs bet\\ ec:n 'PhllKto-
l) zoam and \OillC archacoecyaclud,, suggec;ting ,1 pos-
siblt: biological reLniomhtp between the two gr.tdes.
r-. The skeleton of calc:~reous sponges norm:~lly con-

a sists entin:ly of C<lkiw sptculc~; there is neither ~pon
';J- gin nor ~ilica. The spiculcs arc often of rtllllng-tork
th shape (Fig. 4.oc). In the large Order Pharctronida
he the spiculcs form a closdy p::trkcd mc~h of difFerent-
r:J sized 'tuning forks', givmg ,1 ngtd and eastly fos-
sihzt:d ~kcleton. Rich jur:m1c calcareo u~ ~ponge
faunas arc known, \omcnmt'~ ::t~\OC'Iated wtth reefS.
Well-kno\vn Crctaceou' gem·r.1 mclude the v.lse-
likc Rapludvllcma (Ftg\ 4.6d, 4. 1 '\) .md the dj~tJte
Pcro11idel/a (fig. 4.6c) whtrh togl·th1.·r tcmn extemtve
:d. lipe4 14 Volospongeo gigonleo (M Cam.) from Utah The sponge bed~ in the Lower Crl't,ICt'OU\ of \ourhcm
I3 Ub:e of this keg-shopoo M.lloctinellid is shown, with low, England.
~mounds covering the entire surfoce The osculor margin is
~stand spicular impress1oos ore deor I 5) (Photograph A fe\\. ~phmctuzoam 'et·m to bdong eo the
ex ~ed by courtesy of Or JK Rigby.) Calcan:a.
94 Sponges
bcll-slupl.'d mltrmdt•t L'\, not oftt..·n found los,lhz
Often the mt·g.t,deu:' umte J\ they grow togcr
Ht•:<.:,tdindhd' .1ft' of the normal sponge ~hape ;md b} t..ompkte fll\ion or b) the fom1atton of ~iltt
lllJ) bt..· .mchllH'd b) ,111 t•xpandl.'d lbngt' or b~ a hnk' between .H~.tt.. L'lll 'Pttult"'> In ~ut.. h dn·p-
wft ol gl;"'' fibn.''• c'pcu.tll} m th<:" deep-sea fonm gl,"s ~ponge~ ·'' l~uplradlc~ the whok ,!-det
(f tg. 4.1 (l). betomt:' ng1dly UIIHL'd, usu.tU) by the grov.th
UJg-,Jupt•d. VN:-ltkl· .md dendritic form' are C0111111011 \ht:,Hh uf,thl,l t'ttvdopmg .ldjJcent pJ
!-no\\ n, \llllk' ' ' ith Lttct.tluscul.t fhc ,kele£On 1~ of rays. Hcrt..· the 'kdctoll fonm .1 n!:>rtd l.1tuce p conttne
np.1hm ,tht 1, ,·omt,tin~ ol large (Jnegasclere) and rectcd :lg;tm't rorston by 'ptr.tl ,iJil'cous g~rdl tl d.i) 1
.,111.111 (micro!>clcrc) 'ptutb {ltbr<; -1- 15, -1- 16). . runnmg round the oubtdc ,lt 45° to the axh. there h
I ht..·~t .lrt \\ 1.'11 ordt rt·d. t"xpn:~'mg cubic'' nun.:- Hcx:~cttnt:llit..h .tre dJ~~tlicd nwnly on the bJs !IJ>; C ht
try I he mq~.\\t k·rl'' ltm· the outl"r .md inner \\ails tht'tr ~kclct.ll ch.tr.ltll'l ~
the stntcturc of llw 111 llt<.:.r det
and arc ht rt• ftn--r.t) cd. wtth the ndd ray {axon ) .md nmTmdcrt'\, whl.'thl"t the\ unite ;~nd ho\\ llr
pomttn~ nm .mk \tx-t.tyed mcg.1sclercs are found link, hut ~uKc IIHCIO~dcrcs ,tre ~cldom fo~sihu
tn tht rt..·ntr,tl tr.tlll·t..ul.u and cho:mocytic l.tyer~. there arc snml' di llicult1e~ with the ta.xonomy ol t
together with lht.. tll\lt h 'nulla stdlatt' or dumb- stJ fom1s. I k x:.ttttncUids CJll b~ traced b.tck to dJi
I Ill" .111
•:11com
111\CI tl'l
1bun,
( unhtl
(.;.unb
r. ra
\ t
thcsr tJ
llu:y li
'\(fOill,lt
h rm ~ t
h 1\C bt
uurna h
recti tl
< llt<"l c
Arch
th ht
Amtt .tl
tlhcnt
Wht..'tl'
ll..mt.t,
\lt,l[i~·1
\td
n1 g:.m~
I \1 SJll)l
I he
I• lf)'t
f 111\:t
uutcr •
tl 11 r:l
1•1 Aulorossella
F1gure 4 16 (a) A deep·seo hexochnellid gloss sponge, Aulorossello (x 1) and (b) spicvles of Tholiosterello (Hexochnellida)
micro· and megascleres (x 10) ((a) redrawn From Schulze and Kirlcpotrick, 1904, (b) redrawn from Hinde, 1887-1893.)
lncertoe sedis: Archoeocyotho 95
m~nan, .111d nuny nnW-t'Xtll1lt ~roups tioumhcd

the Upp~r l'.tl.lt·n.-oH . l'ht'st' 'pou~c:~ were
cttd '~vtrt'l) by tlw Pt·mu.m t'Xlllllllom bm had
\t.'red b} rht• Jur,,,,ll, '"hen IIIJil\i ne'' ta:xa
• Ordt•r I kx.ll'tlno\ld.a \\'Js 'H'r\' unportant in
tn:tJCcous hut " no\\.' nt'arly exrincr.
exlcundh<h wcrt' .1hund.1nt 111 the l'ertl,lf) on the
nncnttl \hcln·,. bur s1nt c mmt hn~.tctmelli{i
\ hvc m rh( b.ulw.tl and .thv''·'' ?Ones either
hJ' ~ccn 1 'h1tt <'i lnhn.u or the 'helf fonm
\t become t'Xtlm r .1nd only dl.'sct•nd.mts of tor-
Figure 4. 17 Archaeocyafhid morphology modelled on

lt .mh.tcory~thld' .lrt• .1 vl'ry t·.trly group nf ol- AiocicyatllUS (Cam., )C 1.5). (Modified from Hill in Treatise on
Invertebrate Paleontology, Part E.)
•rr"u' fmsik mu.11ly in tht• torm nf ,1 porous
cncd cmw, t"lmnd m.unly in lmvt·r C.nnbrian
onJtc f.1ett'\ and pt'Nstnlg only until thl' upper The ourer :~nd inner \\,tlb .trt pertor,tted b)
nn~n.m . I ht'lr .u llll \\,Is during the lmver numerom hole~ .unngt•d tn lnn~'1tudin 1l row\; till'
1hn 111, hut rht') dt'< lnwd tlwn-.lttn, bt·commg scpta arc '>p.1r..cly perfor,ltt.:. In lllt.l\l trch.ll'lll y 1th1d'
rut· .md tonlllll d to trYptic h.Jblt.Jt' bdt>n· the uutt.:r w.tll h.1s ,m,tll pun:' wht.:n.'J' thow nf the
r finJI cxtmttion 11 thl• end ot" tht· C.1mbri.m. mner wall .tn mut.:h Llrgt'r. Tlw llllt rt.l\tructure of
~~ hvcd 111 'l1.11low w.att·r... tlftcn .1"m uted I\ ith Ull' c.:up 1s of pol~,smtruu)·\t.tllint· t uutt, mtt.lllv
lUtoht(' md iimmng thll ker... loc.tlized bm- altcrt d by diagt.:new•.
ur bmstmnH·s. or·" i'tll.Jtt·d indindu:1k They \l;luJe _-fj.Jtity.ulm~ "ofrcl.ltl\d) sunplt.•um,tnt~
, been COINlkrt•d to lw tht· tiN 'rcd:-tom1in~' tion. other .trchacuq .nhid' 1111\ h,l\'t' patnr.Hed
uls, but tht'} .trt' mu.lll} 'uhorJin.1te in thc~e walls and scp[J, and ,,(,o .ll'tl'"m) \tru~ tu re' uo"mg
to strmn.tt<'htsc.: .Jig.H.'. gn'" 111g in dump~ or the 1nten .tUum (Fig 4.1 ~).
tmd on the 'urt:ltt'. Tht:5t: mdude r.td1.1l roth. pt·rfm ttl.· tt.mwcf't'
~rrhscoC\,tthtd' .1re timnd 111 .111 tOIHtllt'llt'. hur pl.Hes (tabulae) .md mtperfor,ttt.: .1rd1t.:d plJtt.:s (dis-
bt~t-known tilun" .m: 111 R.u~~IJ, South sepin1ents). These dt•ment' tend to be concen-
tultJ .mJ \\t..'~tcm North Amcrtt,l. Thl'} an: trated tow.ud\ the ba\e ot the tllp. A kw genera do
ellt fi·t)l11 the Bnll~h 1\k, .md norrhnn Europe. not h.wc an umer wall; sollll' ot these are very flat-
.re the) Ott..ur 111 u>ntinunu' 'cqucut..c. J' in tened cones, nt·arly dtscoJd,tl 111 shapc. Ot1tgrowths
u tJ, they luve prm cd to In: of con~idcrablc trom the outer w.tll arc qlllte ct.1mlllon .md include
• tl~l'lph~~:.1l value. tubul,lr rods, tubt.:rdt•s or s1mply dcnst.: Jrrt'gular
Archacocyathu.h (h~. 4. 17) .m: nolnl.llly solitary masses; the l.ttter .tpp.trently grow .1s .1 defensive
be.mng ,1 supc1 hu.tlrc~t·mbl.tncc to corals
I!'Jllmm mech.m1sm where nt•tghbourmg spenmem tontact
j'~)ll~t:~. the mdiv1dual m overcrowded t·ondlttons. I hough
flw skdl'l\111 or utp 111 ,, typ1t.1l cx:.unpk 'uch a~ mosr archacocyath1ds are lOillt<ll and solitary some
)~thll) (Cl."' Rq~lll.llt'') " Ill.> more th,m a pair rare colomal fonn~ arc known, w1th mdi\•Jduals
~t·rtcd t onc~. one inS~dc the other. tomling bemg e1thcr arranged Ill chams or bl.mched .1nd
uter nd inner walls \\hit h .ue numectcJ b} vcr- shrub-hke.
od,ll parntlllll\ (scpta) JnJ 'ep.tratcd by <tn The mu.ll "1ze of Jrclueocy.nlud" " 10-25 nun 111
ul.lr ca\'11')' (the intcrvallum) . A l,tr~e tentral chamt'ter and up to 50 nun h1gh, though ~pecunem
I) ~('Ill\ an.llo~ous to tht p.tr,t~.l,ter ut '>ponges. can reach a he1ghr of I 50 mm .md ,1 few gtam-;
.vith km er part ot tht.· tup " mu,tll~ expanded ulto ,\ exceed rht~ r:~nge.
fLtn~c \\ tth root-hkl' ht.,ldt.l\t~. Mmr arch:~eocptluds. likt· AJtii'IC}'•IIIuls. belong to
96 Sponges
were t()und to b~.· better .1li1pted to rap1tl t
vdonty th:m tall, ,\emkr on~.·~ Appan:ntl} the
ical ~hape of ;lrch.wocy.tthid,, and in parucullr ' I fl
large t'Xt urrcnt npt:ning. "as wdl .1daptrd for

gencr.mon ot '"rh pa"ivt: tkm _ Thou~h the p
gcnerauon of turrents m.1y ''ell luve been 1111'
tant, 1t is unh~dy that thl• .lrclucoq·athtd' a
doncd the \1\dul tlJgdl.1r pumpmg '}~tem de\
by thL·ir pre\umcd protozoan aJKestof\ ..tnd I(
m.l) h.1Vl' bel'n a donun,Hltl)' .1c ttve pnKc"
nomJc.lll) boosrt•d by the pas,ive Auw componcn:
Archacocy.Hlml.. ha\c: bel'n reg.trdcd b} m.ln\
'n:mm: ·~ tir~t attempt to m.tke ,1 rnulucellubnk
r
lrr {l•Jiar
tal org:mi~m'. Bul tht.• 'ucce~\ of other ktnth

sponge~ m;1y well h.wt· been .m unport.ml f.1ttllr
thc:i1· early extinction.
Nc:-~rly .Ill arducot:yathtd~ comc from c.1r!'lon
Figure 4 18 Structure of the orchaeocyathid Metaldetes shown ~hdf <;t·dimcnrs Jqw~itcd in wann sca~. They uin
in cross~ section (left) and oblique longitudinal section (right) with
rod,ol sepia (Drawn by E Bull from photographs in Debrenne
monly adhered lo the sub~lr.He hy holdfa,t\ or ·.un f•gure 4.
br dcvic"''• though in ~o!llL' of thL cup-ltkt· lt Cl wmblog
and Jomcs, 1981 ) liOns) (Aftl
found in o;oti ,edimeuts the holdiJ\t\ \>.ere wry~
l 'I '' R~.·,~ul.m:, , I h~ t.Hhcr eh,~. lrregulare~. ha.' or .1bscnt. Th<.:v arc 11\0)t t ommonly ;w.ort.ltcd 1
lnH:r llll'tllb~t'>. dt,tm~tmhcd hy IITl'b'lJlar pore al~:,ral m om,llnlttcs; howc,·er. they .1ho o, l lwrc
'tm~.-wrc' u11 bnth '' •• 11, ,1 nJ nftcn ;111 irrq~ular out- bctw~:en .md Jround the .1lgal red\ .md m heddtl.
lim: o( tlw l up ·'' wdl limc\tolll''> wnh trtlnbttc\, brat h10pmh
hyolithid,, but vcrv r.trc:ly \l.'ith other ~rong~.
Thc:re t' cv1dcme from rhetr occurn:nCl' 111 \U
snu<;hcd. tumblct.l block~ .md their common lh
Soft ports, organiz ation and ecology
.1t1on wnh .1\gae that the\ wac dt:pth liuuh:d.
1 ht.• ,oti lt\\lll'\ of .trch.tcun-.tthid' arc quite optunum being 20-3!1 m. mdlvJduaJ, bcco
unkn~.>\\ n, thouf;h the~ pn:-,unt.tbh posse<-~ed -;malkr .md r~.·wcr dO\\ n to I ()(I m. Thetr modr
~.ho.ullK)'te,, hk1.· dm,t· of ~nhcr 'pon~e'> . One usdul growth 1s !.hown trom .1 hfc ·'"emblage of OCLlll 1.'1
po111te1 t\) dll'll' hiolog~l .tl '!-.rr.ldt.•'. however. is their Low~r Cunbri.m .1ge trorn Amtrali.1 (Ftg 4 r lt,h re•
0
LJp.ltlt) to l'l'!!,l'lll'r.ttl' th1.· t up '"here d..m1.1g;ed. wluch Br.mc:r. Jl)7(J). lll Sll "
pmnt\ t<l .1 qtuhty of organi:t.ltton and 'individual In tlm t:lUna m.my sp~Cit'\ of Rq~ulan: v. I ''"I' 11
111tq~nty' .tt l1 l\1 ,\, good ,1, that oC the living Poriter:l. found dmtcrcd togcth~.·r. grnwmg in the ,,um g \\Hh et l
Mo't .nathnnttt'' .t~rn· thJ.t .trdtaeocyathids were cral dirc:rtion and rc:tchmg hetghts of up to 901n l ltl ~elll
ltltcr fccdt r,, .ttlt\'Ciy pu111pang water through the l rrcgularcs arc prc~cnt but tewcr. The larv;H? .tl\1 (•r ~up-1
cup :tnd \tr.tinmg ntrthe ruud p.uudes, perhap~ with ~cttlc:d on de.1d .trchac:ocy.lthid~ :md usuall) on 1\ l'ldcJ
cho.HlOl ytc-hkt· rdls, but the filtering function is outc:r wall ne.tr lhc top. WhL·re the juvcu tt l h,H
not ol)\ lOll'- 111 hTegul.irc~. Fill me t<trt.k experiments attached they fixed thcmsclvc_·' by exothecal ou !h.tt ofc
11'-111~ anodd .lfd1,1l'<lt y;Jthids (U.Ilsam and Vogcl, growths lrrcg·ular masses of l'Xothccal mJte ltmh .tre
J<J7 J) 'll!:\~l''t th.tt 'l'·'"ivt· How' w1thom flagellar tormcd whC'rc andavtdu.1ls of tht· \;tml' .md difle 1\-l.lll)
pumptn~ nu~ h ,l\'l' bc:c:n 1mportant The model~ 'pectl'' come Into ront.ld 'ugg~.·,t cnmpctitiv~ 11 ~'l'"' i .\11
\\ e r~.· tix~.·d 111 1 tlunte t,mk in a lammar current .tcttom - pcrh.1ps the oldt•,t knO\\ n ca~e in the fi l • n tbri.
\\h(hl' 'nt·n~h lould b~.· v,ml.'d. ln th" current a record. In 1h1~ f.tun.l .1 mtcrttl' l.!ycr ~urround5
n · lm lt\' !-!~' tdtl'nt ''·'' e'tabb~hed. gcneratmg a ~cc cup, the work of the probkm.ltic l'ncmstmg ur
ond 11"\ At)\\ ti-\llll th~ outer wall through the mtcr- l'>lll Rl'll<1lm. wh1c h bound tht• .m: lucocy.ltlud eo mdl\ rdt
\'J llum .md llllll' l \\-,Ill w till' ~.entr.tl ca\·ity and out mumry together .md \O cn:tbled H to be prcscn·td 1992). ~
thruu~h till' 1 " \ ulum \hurt. stout archacocyathtd~ a lite a~sembl.1ge the whc
lncertoe sedis: Archoeocyotho 97
vamJu~ directions. Modnbrity has pwvcd c~pcci.tlly

,tppropn.ltt' for reef-building arch.!t:OL)- atham; Lltl'}
<.:an grm\ larger, the} h,l\'e improved rq!;ener.ltl\'l
po\\ er- .md they l an enaust sub,ll.lll'S. lndt:ed,
ard1ae<Ky.uhans wer~· the tirst ol all ,J..cJetolllzed
llll't:llo.1m to havt• developed modulanty. It wa~.
ho\H'vt•r. only the forms w 1th perforated ~ept.l
whit h bt:camc modul.tr, pre.sum,1bly beou\t' sott-
tts,ue connecriom an: c.'\\enrial for \Uc.h ro de,·clop.
The lkgrel' ufintq,,rr.ltlon varies m .uch.1eocyatham ,
.1~ wlth t.m.tls. Tht: k·~~ wdl-mteg!Ollt'd form' .uc
'pst.·udut.ohmial ', i.e. rhc.·y consist ol m.111y rdativdy
similar l111kcd clemt:tlt,, and tht· bra m hmg fonm of
the,t' were the mmt ~un c,sful and long-la,ting of all
archaeocyathans . They wcrt: common m earl)
C.unbn.ln reefS, whl'rC mrbulencc, ~cdnnc::ntatlUII
rate~ ,111d nurrient ~ upp l y were f.11rly h1gh. Otht:t
arch.lt'Ocyatham . however. became tnl'reasmgl~
well mtq..,'lateu. c'~x·c.·tally the nus~l\'C l;muua.r l)r
columna! type~. l>O the whole org.mt~m m.1y be cun-
siden:d as a ~ingle indl\'ldu.tl, ratlwr than .111 aggrt:-
g::tte of ~ep.uate m udlt k·s. Thi~ i~ partJClll.uly the l'.!Sl'
fhm.: ,1rc v,ull)uc; exa1npk' ul rcd-.J\\oriJtcd in the Irregulares. some of ·which be.1r ,1 srrik111~
~<IC}"JthH.k One mterestm~ tum.1 dc,crilwd by resembl.mce to strom.ttuporoids .
hrrnne and J.lmc' ( Jl)~ 1) liom l abntlur .md tht•
~g lO.t\t of Nt·" toundland ~~ of L1te l mn-r
1b11.111 agt. lt tonsJsts onh l>f !rrcgulJrc,, .md
Distribution and stratigraphic use
1u~h only a very few speoe~ ,Ill' prc~enr thc'c .1re
~h packui. 1 he paunty ol 'pcne~ 111.1) be Archacory:.rhid s Jre tound in .111 <-'<.'lltHicnt~. though
Ult' by rh1' ,r.1ge the arrh.teoryarham wac thc1r dt\tnbution is patrhy; none are reported !Tom
nung ie\H'r 111 an) case. I h~: arrhaeoq.uhan ' the Bntl\h Isles. Thetr abundance in the Lower
r l"tthcr with t.lbular btmtromes or \Hth ~mall Cambn.m of the USSR. has enabled very prcnst•
~~reds (b1ohenm). where they m.ty compri~e up time r.mges to he worked our for ovl-r lOO gent:r;l
51) of the rot,ll volume of lh~ roLk. Sub~idiary and uwd stratigr.lphKally most sm·cc~~full y. Thl·
l mdudc tnlobm·s ::llld bt.ll h10pod~ together oldc~t known ~penes come !Tom the Tommoti.1n
~clnnodcnn .md hyolith1tl debm. In thiS rock stage in R.us~1.1 (R::taben, L9~H). Tht:v Jm~r..ilicd in
nu' Mu.zldrtt·~ (rtg. -l.lH) 111,1.,. be of ~tick-hkc the subst"quent Adt:.b.mt:Jn, reached ;m acme in thl•
1p-hke form and mch.,iduals .m:' 'urrounJed .md 13o tomi.m , ;md dcchnl'd greatly 111 the Toyoni;m
ll:d together by .111 exteJISI ve · cocncn ch y m c ·, stage w hich tcrrnin.lt<.·d rhc I ower Cambri.w.
11 th, ntp~ h.wc a 111a.sst\'C lubtt, resembling Specics ,lrt: fl'w 111 the Middle Cambrian, and onl>
fwra.h, nd from the coell\:ndwme nunwroth one Upper Cambnan survivor has bct:n recorded
ve found to 'wam1 from Am.uctica. There ts some n1dencc of pro\-
'llln) archaency.uh1ds are sohr.11-y. and thl'~l' .m: inciahty, with three or more provtncc' in the
11Jllv common 111 the e.1rl} part uf the Botom ian, limiting intercontincnt,l l corrd.1tiun
m~ndll. Sub,equcntly the modular tonns bct.lllle using ,1rd1aeocyathids. Fven so, the R.u~sian work
commot• Jlld surce~sful. In tht:'e the org.umm ha' shtm n the potenn~l ~tr.ltigrapha .1l value of tht:'l.'
1hole rom1't' of repeated funt.nonal Utllb or renurk.1blc exrinct poritcrans, whos~ zon.t.l u~c w1ll
,Juah wnh ,ofr-parr contmutt) (Wood rl ,,/ .. ll11UOllb[L'Jiy JnCfl'J~t' ,)S the flllllJ~ become bettl'r
). StiLh muduk~ lllJY be .ldt~kd to any part or
1
known.
hole organism, which i~ then ,1ble to gmw 111
98 Sponges
modem ~ponges. ~.g. Cfl,ltld (Col:>b. I 1h J). ho
1 1 alw.•~
mto hard sub~tratt'\ .md may cuntnb ute qt~~te \lt{llll-
tt:ph.
Jtcd
•ramly to tht• destru ction of ,hdls.
:lnthc
.lrt• r.
Spm1~;c' arc of link ~tran~raphtc.ll v.1lue. though
bet\\•
rht•y have been u~cd .1~ marker s .lt ~pecitic honzo m.
M<
Tltur mam tmport .mu.: gcolog ll.tlly hcs in thJt tht·v lot.lll
haw bt:en a smmc ol bwgcn it Mlica and th.tt rhcy Jd.llt'
an.: an ekmen t though n(lt otten .111 import .mt am·. Spicu lar spon ge reefs
'cor.tl
in reef fabncs . .11
Rtch f.tunas of 1. mcd mtddk• Cambr ian derno- In the Jura~\lC thnc dcvdo ped v.trinu s .111d mud•· Jn.'
\tudied kind~ of ~ponge reel;. Tn Norm. tnd) fo

'lponge s .md lwx,tttmdlld~ are known from the
Burgcs~ "ihale .md trom Utah. In the l.mer regton
examp le, sm.tll Middle Jurassit: biohcr ms were bu~
tht·y .lre often prt''ICr wd lhttenc d but ll)mpl ete (Ftg.
Ltp by the ltrlmti J Pl,ll)'f"cmi<T (Palme r and Fill"\1 Ca
I <>!H), mdtVid ua], of wh tch c~:mentcd to t'.Kh orl tl
-t 14). and the\ haw recentl y bct'n dc~tribed in dc- I hrot
to fonn J ngtd framew ork. The,c probah h gre''
gant det.1tl by Rtgb) (li.JHJ. llJHfl). (srron
the lower photic ..:one, below nomlJ I w<~vc b.m· b1
1-le.xacunellith throug h tmw 'tTm ro luvc bct•n hn·t· t
.1bove \tom1 b.!!>e \!nee some of thtm )how stgn
m.unly ~:onfim·J tu the 'ntT-rc cf t3cic' 111 deepa Sta
\Lorm dam.tg c. A low-dt vcrmy comm unity d(lrru-
w 1tcrs, where a' dcmos pongt' ' and c.1lt,1reom to ha
~ponges pn:t\:rr t·d ,h,illo wtr \\,Het"' . Ordm·
in.m nated by the oyster .Am 111 grew on the upper sm
,,dtnt
-;pongt ''• c:~pt•tia ll)· tht· carbon ate bctc~ t,tce, while the qutcte r-v..ner lower surl.l
of
Jitlmtt d 111
str0111
North A.mcn c 1. \H're qmte unport ant :t\ frarnt· suppOrtl'd .1 \l'f) nth .1~sembl.lgc of tncnt, ung t'21 •
sponge s, cydmt ome bryozu ,lll' .111d o;m,tll th: r hey
bmlder s 1n ~tmm.ttopurotd, brynz.o an (Rigby , II.J71)
ctdeace .ll) brat hiopod s, :tll of which were 'u~pen,i \\ lthil
and 1lg.1l reef,, 'umett rne' bt•mg ''muc h .h half Llw \\llhu
lcedel"'. Sm.11l nemng br,JCh iopud,, "agile t'L"hlllo
totJI reef volum e. from time to tune •n rhc h ltwk~
P JI.H.·ozotc tl11.:rc wt·rc qmtc: t'Xtcm t\ c lncali7l~d .111d 5t.lrfish and boring wonm and bivalve ' .all
bur
~tlun.ln ..:olon t71.'d the red, whcrc:a~ erect bryo70.UI\ I
devdopmt:nt~ ol sponge~. stKh .1s the nch .tltr.:trt
1 l'l' and the Lkvom .m .md un the mtcr-r cd mud surf.'lct•. Thm a chvel"\c I
<,pon~e t:luna' ot c:tme'' ,, 1101{
IS\t~stp pt .tn ~;Lm ~ponge ' ut New Y nrk ,\nd b.tl.lne t•d CtO~)-~tcm bJwd upon tht· tT.1tlll bu
M 1r( h:w
fJ/,uyclrt•nia was abk to dt•vdo p.
Pcnmy lv.tm.l , but rht·<,c \\I.'TL' nnt J<,snci.m:d with horL'I"\
red~.
Tht'\l ~m.tll red\ contra: .t \\.tth rhe much I
Jur<t%te sponge al~al reds of south11t Much
In the Penm,lll tht· r,pon~t· tJunas J'~oci:ttt·d wtlh Upper fcmn l
m). \\ hich ~cw to \Oillc: cotlSldenble !11
the Tcx.tn red tl..lmplex 1rc n:ry weiJ known . Here l;enn,
above the sc<t floor. In tht•se (e.g. Fliigd :tnd StClft
Ill h;)l'
tlw (Jlcisp ongt'' dcrivl' d trom CJrbo ntfcro u' 'helf:.. )cp 1r,11
hvmg 6una' bt·t.llll l men::1\mgh import ant on thl'
11JR I) tht• domin :tnt !Tame bmlder s .lre ht xactm
.md lithisti d siliceo us \pongc~. but t-llc,,rt'OLls AI \llrt:tL'l'
p:ttch n·cts and hn:tlly on the barrier reef<>. wherca~ ' may \:lie h.
hex.tctindhd~ '' mn.1l prdi:rr cd the olf-rt·c f
mar- al\0 pl.tyed .} domu unt rok. rhe ~pongc
<.llp-sh ,lped or dt~coid al, depend tng on wheti tl 't)r.
gm.tl t'Jtit·'· ~L)I11C well-J l'vclop ed ' I na'>Sll sponge -.urfalt ''· rL'\ rill'
on hardgr ounds or soft
red:. Ollllr 111 the AJp,, 111 Cl)lltrl \t, the Jura\\IC they grt•w lt·s gmwth ~nd N
nvely. l n rhe be5t-m tdied l.'x.tmp
sponge red; ot '\t.lttrhem Cemu ny, wlurh .trc very Ud.tw.
pnmal'\ o trJ.Jnc- c vclical .md the reef built up in ~tage~. An nu
well kno\\ n. tOnt.ltn '1'<>ngcs as norrno
butldc r,. I !ere thL· 'pnngc s ,\re m.11nly 'lliCco tl' thtn-
'rongc !.lye• was t"ovcrt·d by marl and l..tlkJ, t
more ~ponge~ grew on top. fhesc died 111 w 1 lllmg
w.lllcd tl.mm. oftl'n pre,cr\ 'ed 111 hfc pmitio n, lfl ludt
fomun~ 'mall 'POllf!;l ' mound s, whit h on.tsJO
nally .md tints growrh of lurthct 'pon~e' tontm
llhcr
'ponge J.Jld marl layer' alt<.·nutmg. P<.•rwdtc
l
grt'\\' ro ~reat '\lZt' md united to proJm c l<trgc.> dc,;aJ 'r
<. }.lnup hycean algat• !:,'TCW on t''P ot rhe
OlllnH
m.l,\Cs. Hut tht'\C spnngc~ dtd nnt rat'l' thcm)e lvt'\
a11d tow,lrd~ the lop ofthL' red~' the-.c brc"3! ( •lllltlt•h
t:•r abo\'t' rht· o;ub,tr.Hum, tomung thin A.tt l.tyer; de.tl ot pu,urnr ;
l:nmtrome~ r:tther rh.1n tntt' reek Other orgJm
\lll) domm ant. Thert• \\:l$ a gre.H
tl()n red\ pre1
'tl tl ''··
bw,tro me"· n:-crystallization .md di~solu
111 tht
.lTL' t.u-d~ J\\oll:l ted '' 1th thc'e Imas'K i) Ch:H
bt·in~ un.tblc to prm tdt· '\lltJbk · .till horagt . Many that rhc angina l torm of the spong<.
Sponge reefs 99
\lV\ pr~servt:d. )omt nmrtHIILHpluc t:llln.l~ of with red\, but tht' llla.JOr :.punge reef ftm11er'
plulnpod~ Jnd br.tc.h10pod~ lfl' sonlcttmc.'\ .l~'oti were the..· ~tromatoporotd~. Many 1-':JI.teozoic stro-
, l '' tth the: 'pnn~e reels. but oti:c.·n no ~utt.tblc nutnpormd~ were reel bUilder.. .mJ. \\Ill L thev h.td
~ .1~e could bt· prm; tded .llld nrher or~.llll\1\l\ geolo~cal rt:qmremc:IH~ ''milar to rhme of rt•tf-
rut \ne\.. bn o7o.ul~ m pl.tt L'' ti.Jnnt.>d mcadm,,·, buikhn~ tabulate LOr.ll,, the two .lrl' often found
rwccn rhe \pongt• beds. togc:tht•r as fT:nne bwlders. Strum.ttoporoid~. 111
~hl\t Crt't.lcc:nm .md Tcrti.tn spongt·~. thoug,h gcner.tl terms. tt•nd to displace tabul.ltc:. in lugh-
11\' ~bund.l!lt. do llOt SC'l'lll tll luve bt'l'll rcl.'f cnerg) l'I1Vrronmcnr,.
l.ucd j.Kk,on et ,If ( 11.)71) ha,·e dc,rnbed Rec.elll Four lu~ic growth fonns are found JlllOII~"-t
rJline· ~pon~c br.tt h10pod ultnmunint:,, whtdl ~trom.n.oporoid~: l.lmm.1r. donuc.1l. bulbom. and
ah(l of p.1l lt'onrolot,'lc:tl •ntt:rt'\t. dc.·ndrmd (Kersha\.\, 19H4. 1990). In Dcvonian red\
the l01r~c domical kinds ('ballstont:s') and lJnun.tr
torm' .trc a~<;ociated wrth htgh-ener!-,') l ond1riom on
Calcareous sponge reefs the reef crust and .uc maJOr fr.tme builJcT\
Dcndro\{1 .md ~m.1ll bulbous fonns prt:ferred qwel
ronghom ~L'olo~rtc.tl tlmc.. .trchaeocy·.Jrhtd'\ water nmditiom 111 b.~ek-rcef lagoon\. There 1:. ,,
trumatoponlllh. c. l!.tt·tt:tid' .111d sphinct01o:ms) dirclt pat\lllcl here between these and living cor.tl,.
1t contnbutetl m v.uwu' "":l')~ ro red" torm.HH)n Vnv large rect:.bmldmg strom.ltoporoid m,l\,e~
~lime Jrtltaeoty.ttlud' Wt'n .1hk ro tlx thc1mdn·' Jre ofll'Tl found to have m.ugmJI tnv1gmattom llf
h;ud ,ub,tratt''· Othl'r. tt'lllkd ro root Ill ,oft sediment and mdm1om of 5ediment wtdun the
11cnt\. 1he) .lrt• tlli:cn t\mnd .!''>ocutcd w1rh cocno~t~um, indic.ltiV{ of ce~<>anom of ~rowd1 for ,1
Hnarolttc red\ but thc1r u111tnhutmn •~ mmor time. Tongues nf m.lr~1ul 'cdimcnl m:1y be found
he\ flounshcJ, hm,·~.:,·c.r. 111 'cryptic' h.tbit.lt'\ on om 'idc uf the ~tromatoporo1d m.t~s. SL1ggestu1g
thm rhe eJrh l .unbn m rt·ct:.. 111 1:-'Tottoc~. c..·:witie~ banktng up of srdirm•nt on the Ice ~1dc. The growth
thm the tim1t '' ork. .1nd on the..· under.tdt:~ of of \Uth ma\\c' i<> ofkn ,l,ymmctncal, ,llld they ot'i:en
h of debm .md 'prc.~dm~ ,J..ekr.tl org.m"m' scc·m to k.m over mtn rhe current, from wluch, ut
huralr\' and Wtllld, 11>95). \ud1 ll)pts nttert•d .m cuur~c. their food Lime·. The llH)\l ~pectattll,lr
llr:lrnve habtt.ll, w1lh redun·d ~.·xpmun~ and cnvt- Pal.1~:owil reefs built l.rrgdy by 'tromatoporoid s arc
unenwl stre~~ .lllJ hou,cd 'olit,lry chambaed thn\t'ur the Stlun::LII of C:otland. where tht.>y furm
!l.lcocpdum, l:alnltl!d cy.mllb,lttcna amlmicru- huge 1 ed nussc' up to 211 m h1gh ,111d 200 m in
r.;., 'PKulatc spon~e~ .mJ lHher org.mi\111\. diamett.>r In the\e biuhenm strom.ttoporoid ' were
!ud1 of rhe bwm.1s' ot' the n.•t•t:. \\ ,1, dCCUJIIy 111 cht: domuunt and corah and other dt:nll'nt\ subs1d11ry.
:-~n of crypt-dweller,. 1-rom thl· c..·.~rhc\t C.unbnan, The slromatoporoa.ls m thc~e reef~ developed
1 rocyatham .!ppc.m:tl ~unull.lnCml,lv. b11t in variou~ growth ~tratc..·~ie\ Jnd a v;mcty of respomc.•s
rJt~ assonattom both 111 1.1) pt' md on thc open to t:nvtruJunemal pre'<,ures. Abntpl lll.lnge:. m the
:Ut, Jnd 1t 1\ unltkeh,. that tht: ll) ph \ervcd .1\ direl tton uf growdt .tre due to the 'h1fnng or ovcr-
ha,·em' tor former Jl!mll"n' ofrhe expo,l·tl st·.t run1tng of the coelHJ\tcum. hentc: rht• ~trom.tto
pormd5 n.'cord tht· vanou\ event~ lh.lt h.tvc taken
The inunense Pelllll.llt C.1p1l.m Ked nf Tcx.J\ place while they lived (Kcrshaw, 19H-t-). Orher surh
Nrw Me'XIl o nm~ the m.trgm of the .llllil·m Silunan rects occur 111 the Great L.1kc, Rcg~on
1ure Ua\in \omt' pJrts Jl'l' charactnizt•d by of North Amenca (Hc·c).;cl and O'Bm·n. 1976). In
:nnous piJf) 'pongcs, Gi~dll/tl)t'tll(\?10 tannin~ the tht.> Wenlock of the Wcl'h Bordc.•t, J giant cor.1l
Ill'\ of gre.Jt Cl\ Hll'\ hOll\lll~ .1 I'll h l rypttt 1,\llll.l. strom.troporoid reef, with \uh\1d1ary bryozoam,
lu,1mg pcnd.1nt ~phmnuzo.m' .111d brvuzo,uK fonned a margiual b.u-ricr between ,ltl outer deeper-
ki rcgiom have.. :t foundt:r bl)·ozoan--,pong c w.lter 70ne m which reef~ werl! .lb'>~.:nt and an inm·r
UllUIU[)' \\'lth th~.:· Slll.llkr piJt\ \pUil!J;l' shallo\\ water regton \\here ~malkr rel'f' aboundc:d
) lliJ''•l, but hkt \\.1\l: "' tth .tl'l'~pnt t:mna (Wood (Swffin. llJ7 t). The. .u.:me of coral 'rromatopormd
. IC)<.JO). A, "nh thl.' l..111tbnau ret.>fs. lllO\l of reef development w,1, m the Dcvoman. and
prt~crvablc: lwnthos w,,, homed in the tryprs. immeliSC reefs of thiS .1ge arc tt)UJ1d Ill SOUthWt:\tt•rn
l h.u~n:nds .md sphmctozo.\11' .m.• oftt.>n ,l\\l1t 1.1ted England .111d in Albc..·rt.l. Mcsozoic 'tromatoporoid ,,
100 Sponges
R.taben. M E. (cd.) ( I'JH I) '17tf Tc>/1/IIIOitcl/1 \ftli/C cllld 11 u .• lity o
\\ l11ch Wt'rl' in 111.111} '"·')' dl'\tm.Iar to thml' of tht>
Cmlll'rillll L'll'ff Hcuwd.ll)' l'~t•blnll Amumd Pubh~hut;:. UoJrdm~
P.li.ll."o7oic. \H'rl' IHll ~t·ncr.tlly n:d fonnCT'i. J)nwd<'ll
it \CCllh rh.tt '\'llll~t'S tlt \'Jriom kmd' wcrt able New I >cllu.
Rctd. R l:.ll (I'J5l'l- l%-l) lite I 1'1'<'~ pp 5h7-
to sccrctt' ,rron~ t .tk .trCtHI\ 'kd~:wm in r~:sponsc to Hn.trtmtlltclcl P.ll.tconmgr.~plm.•tl ~Ol'll'ty. l Jmdl', C.J
envJronmcnr,.t 11ppmtumtlc,, .1t dtfli:rcnt tin1e~ (C:Ja,~iliLatiOII, wowth. llltli'}'Jl\1)0"'')') sron~C\.
throughout gcultlgtt ,tl lmlllf\ \ud1 l.tq;c colonial Renncr, J. .md Kcupp. H ( llJII I) 1·•'"'1 .md Rt'<l'lll >J'<>nftl Jlck,,lll, J.l
fom1s .trl' n:f) ~uud .tt llllomzmb h.trd .. ub~trates. Spnnger- Verl.tg l:krltn. 5'J5 pp (MJny 'alua Rt· .. cm I
Yet hvmg 'Pl'll~c~ "tth l ak.1fl'OU' ~kdctons arc paper.) thctr p.tl.
ti:\\. and of lo\\ dt\'e Nt\ , .llld ,trc rontmcd to cryp- TcJchcrt, C (ed.) ( 197 .:!) TruJit(( ,,, fm,•·rrtl., Kcr>haw, ~
tiC lubtt.th. ·r he dctltnc ot· l nr.tlhnt: 'pnngcs. <~rtd Pc1l.tcci/IIOlt•~y. fJ.m I! (rn•rsnl ,,,/rllllt), ArdJctt'c'•'fJtk: levcl-bo
Geolobrical \oncty of Anu:rit.l .md UnivcNt'. (11 Kershaw, •
e~pccully thc \lWnl.ltopmoilh, ath.:r the early
Kans:l!> Prc'> l.1\'> r<'IKt:. K.tn nphonu
Meso/utc " prub.tbl) tnrrd.ncd \\ ith the rise of
Swcdt•n.
rccf-bllllding ~l kr.H:llfll,lll l or.tk WhcrcJ\ living Palmer, T .
caluticd dc tt H!spongl:'' h.wt· much tuughcr ~keletom from th<
th.m do sdcr.tdllll.tm, the l.ttrlr grow very much Individual papers and other refere n ces 24, 1-2.
more quid.ly Slll h r.1pid growth l'l .1 consequence RJghy. J.l<
Ualsan1, W.l. and Vogcl, ~. (J117J) W.tlt:r movement tb
of tht:ir 'YIIlbtOlll rd.tnomhtp with woxanthellac. thrtmgh
~rch,t<•ocy.trhitb. cvtdcnn• .111rl 1111plications of prt'!osurr
Thc~L' cor.ll\ t.ltl thu-; l olontzc .lv;ulabh: 'pace very (',,/t•tl/11<11
Aow in models. Pal.tt't'lllc'/•\1!)' 4 7 , 1)79-H.t. (FuultlOn
r.lpHily· thl') .1rc tdt:.tlly romtttuted tor building 1174-R!
morphology)
red\. l:hddcr, G. P. (1937) 1lw pcrfi:rtlnn of sponv;es. p,,,,.a.f•"t;
lti~by. J.f
Cor;tlhnc ~pllll~t:,,hm\' l'\Tr. l.tck zooxanthellae. 4 rile Lmm·tw .SNit'l)' <!f 1..,,tftiJI 149, 1 14-ln M;U]lllll
Mounta
Since tbc 11\1.' ut" ,cJcr,tl nni.m' the~ havl.' been DrNer. M. (I'J7o) F..trl) Cambti.m mrcrgrowth'
(cxccllc
muhlc to compt'tl' .1~ n-cf timncr,_ and have lmt arclueOt yarhid\, Rma/ds .md pwudo~trt1marohtc' froo
Rtgb).J.I<
thcir t'imna dommanCl'. Thq nu'>\ l'XlSt only a~ ~ourh Au,tr.lli.t . p,,/,rniJrtolt'.~)' 19, .::!23-t5. (Suct~,n
C.1mbri,
rclid' liv111g d.trk md t:f)p!lt habitat<; where hcr-
111 generanon~ of.lrch.Jl'lKY·llhut- 111 htc posttton: btbli
CmJ,Jdic~
m.uvpu: 'ur.1l' c:tnmn p,·nctt.ttc. Yet it is these few raphy) Scotlin , I
Cohb, W .R. ( 1969) Pem.• trtttnn o( l.tktum cJrbonr.
\urviHlf\, thl' lllOlkrn ,~,·Jao,pongl''· undiscovered Wenlot:
mbsrr.ttc~ h) thl· bunng '('OIIl!C C/ior~or .-ltutra
unul rcfcntl). \\ htl h h,l\'L' t'u.tblcd .t mbstannal 17, 173
Z.•vlt-.x•sr 9, 7HJ-'IO. s hulzc,
rcimcrpt et.ttiOil tll the Phylum Ponfcra to be Dcbrcunc. r tnd J lllll'\, 1\1 p ( 111H 1) Rcct:..J,,OCU'
ctfccr~.·J.
I kx.trn
Jrch~coq .llhJJs frum thC' ltl\\'Cr CambnJn of L.1lm
19111-0
.md Nn\ltllmdbnd fl.t/cJr<tlll•'/•li!Y 24, J43-7h. \
1-6:!. n
text) StCilll, C
Dcbrcnn~:, F .111d Wtlod, R (I«J[JII) A ne'' C.unbru:
sphtncto7ll.ln sponge front Nmth Amcnca. 1t' rdall
to Arch.lctKyath~ .•md the n.Hurc llf e.uly sphtn(U)o
zn,m~. Ct'tl/t\~it',rl .\/,r\!11 ~,,,. 127, 435 43.
Books, treatises and sympos ia
Flligel, t ..tnd Stl·iger, T . (llJHt) An Upper Jun"'
Elcrgqui,t. I) IZ. ( I 1J7t!) "~/'<'11.1!1'·'· Untvcr,tty of Califomta ~pongc-Jigal bUild-up front th~: northern Franhn~l
Prc"· Ht•rl..dl·y, l .tltl. (l1vmg l.mn,t\, 1llmtr.ttl:'d) West Gcnn.my. in Uuropt'tlll Rrd ,\ foJc/., (cd. IH
C.trtcr, <, ~ (I 'h2) t .nln,ll /.,,,,f•'~l' •!I rlu· ltwcrtebratl'.<. l'noney), Socicty of Pnmomte PaiJcontologisrs .11
Sidgwit k .md 1.1t k\nn, lnndon Mmcralogtsts Sped.u Publitallon No. JO, ~ociet) •
I teckel. P. ,lJld 0 Bnl'll, (; I) (I 'j7h) Sr/un.m Rnis of tire Ecouornic PaLH."Onroklg-t't' .md Mincralogtm. Tu
Crrtrl L11kr< /{,•1/i•'" ,,f '\.mil .'lmrnc.r. AAPG i:teprint Okl., pp. 37-9H (T) ptc.ll sponge red)
Scn••, No. 1 I, A nil n< 111 A"u,utmn of Petroleum C.r:~y. D.l. (19HII) SpKult l''~udomnrph~ in a 11
Cot:tiO~\h , Tukt, ()I.;\ (Nunll'l'llll'l pJpc~) P.J!Jeozmc ch.lctcnd Jnd 1l, ,.:Jcrospongc JJ:fuUu
M.1~fnll . N.B. ( I'17H) -l •t'"''' ,,f Dnp St,l Bi,>lt\11l'· f'cl/t1('0111il/t~)' 23, ~()}-J(l
Bbndtonl. I ondun. f l.mnun, W F .md C:nrt 1u. 1 . E. ( 1970) Jarn.l

Moor~. R .D . (cd ) ( l'l::i:i) lr<•lltH' ,., flwatrbmtt' tnnlhnc ~pOllJ,!;l'\: thur morpho!OJ,."\', ecology. anJ ,
l'.llllt'cii/I<>I.~!!Y· P.u t I Ir, ltcl<l><1st,\ <'( 1/tt Z.w/,1~11<1/ \,l(tt'
Geolt,gtc.tl Soc11:t\ ''' Arn~n,.l and Unt,·e~ity of l..Nrdo11 25, .::!05 -4 3 (Sdew,pong~de.Ul demo,po11gr
H drtnun, \\ !) .md Rei,wi~. II.M . (I'J7J) The ind1
K.uh.h Pr<'"• 1...1\\ I< tll'l'. K.lll.
Bibliography 101
U) of 'pongt''· m • l11111t,r/ ( .'oh•mrJ (cd~ R..l.o. por01d~. Pal<ll'OIIflllcli!Y 9 , 7 ~- 124 (J)i,tincuon of pri-
llo.udnJJn, A.ll. ( hc,·ch.tm .md \V A. Oliver). mary from secondary nutrmtructure)
llo\\drn. llutdumon md Ro". StroUlbburg. Pcnn., Steam, C. W. (1 972) The rclanon,hlp of the monuto-
rP Sli7-ll.J . poroJd~ to the ~cll•ro,pongl·~ . Lt·rh.1i,t 5, ,'\(,lJ-HX. (~ec
, (;J. (I XX7-'JJ) A morlll~'T.tph nf the Bnttsh fos~il text)
lf<'ngt". f>,Jfa,,ltlr.~r.IJ>lll• 1/ •.,.. tfl)' ,\1,•11•'\!rtlphs.l-25 -t. Steam. C.W I '>75) The 'tronutoporotd anunal. Lcth.lw
wn J BC.. Gort'.IU. f I ami H .mman. W.B. (1971) 8, R<J-100. (Sponge .1ffinme~ of \frolll.Hnpormd~)
R('(cnt bradtiopod coraUmc 'pon~c commumncs and $\\,lll. A.R. Il . .md Ke~h.1w 1., (I'J'J-l) A cmnputt'r
cbftr pllacoccnlogJC·.ll ,j~,''llllk.mn~ '-•mu· 173. (,2.'\-5. model for ,lo.dt•ul ~'TO\\ th of 'tromatopormd,.
l\t. S 19X-t) P.lttnrh of 'twm 1topor01d growth m Pa/a.wrt<•lt';{?y 37, -+09 23.
lc1c.-bottorn t'll" imnlllt'IH'· r.,/,rc'<IIIIL•h~} 27. I U 30. Wood, R. ( 19H7) B1ology .llld rt•n,ed W\tt mJucs of 'ome
hJII, S ( 19'JO) l.,rmm.\lnporotd p.tbeobmlogy and late Me~ozott \tronutuportud,. .\p'''"'' Hrper' 111
honomy in a l.otlunm bimtnllllt' m Gotland. Pa/ac<llllof<!I/Y 37, 1-89
s~~~~n . P,r/m·orll•'"!l!l' 33, MH 706 Wood, R. (1990) Rcct:..building 'P<11l~l'\. .lm<'lltclll
J:ll1~r. T. and Filr,ich. r ( l'JH I) l·colo~oy of 'pongc-rcef~ Sciemrst 78 , 224· 235.
!rom the Upprr fhthom.tn of Nnrm.llldy H!l<leclllfcliO,'?Y Wood. R. and Rei mer, J. ( l98fl) J>onfi:t.ul .11linll 1 e~ for
H. l-13. Me~ozoic ~tromampomids. J>ai<IC<IIII<rlc'l!l' 29 . .fo'J-75.
. il([>) J.K. ( lY71) Spongt'' .md red .tml rcl.1ted facie' Wood, R. and Rc1mer, J. ( I1J8H) "I ht• Upper CretJceous
thmu!(h tllltc. flrc1Ct'l'di11_s:s 4 tilt' Nm'llr Amerim11 'chaetetid' dcmospongc s,,,,,,,uaxlln'lla irn;l!lllaris 11.g.
I'.Jr,,llr.>lcr.~ir,.! C'.ltrflt'flfi<lll, Chtt.lgo, I <)690), pp. (MichcJjn) .md it> ~yslt'tll,ttil uupltt.tllom. i\'c11n
r-1-88. (A rt·cc nt treatntt'llt t1f spon~e n.:ds) Jaltrlrllt'h fiir Pa/il!l/llt>lt!liil' 'lblrcllldiiiii~CII 177, 21.'\-22-t.
~by. J.K ( l 4HJ) Spml~t'' nf rht· M1ddlc C.unbnan Wood, R., Dic lo.son, J.A D ..md K1rlo.l.md. U L. ( 19%)
~llrjum ltmc· ronc from rill' llollsl' R..mgt• ,tnd Drum Observatiom o n the et·nlot.'Y of tht• Pnnu~n Capll.m
~buntams of Urlh . ./•Hrm,r/ •!I l'.lfc,•III<•I••I!Y 51 , 2-t0-70. Reef, T ex.ts .111d New M,·xico. !'.1/,u•tlllf!lh'.l!y. 39,
hccUcnt illu .trauum) 733-62.
v J K. (19Hh) Sponge' of rht• IJurgl''' Sh.1lc (Middle Wood. R., Rcnner, J..111d Wt'\t, R . (19X9) Sy\lt'llldtlcs
mhrun'. Bnct'h Cnlumbt.l p,,J,,t<,llft!I!Y<IJllll«l and phylogenenc impliutinm <1f the h.lp)o,cknd stro-
C4tt.WIJI'•1 2, I 05. mamporold Stwd/i,1mtra nov. gen l.£rlwa 22. 85-91,
1, TP (! 971) fht• wndttlllm ni ~'Towth of the Wood. R., 7hur.llev. Vu A, .111d D ebrt·nn<. r (llJ92)
Umlock reef• of \hrop,Jmt• (Engltnd) '\t•dimmwlt•i!Y Funcrion:tl hJOIOg) .111!1 en1ln~n, of the Arc h.1eoc) Jth.L
17 17J-21lJ ({'1.1\\ll) Pal.tr<l> 7. 131 56.
c, P.E. and Ktrkp.llnd... R . (I 'Jll-t) Die Zhuralcv. Yu A. and Wnod, R. (JIN5) Lower CambnJn
He.'UctmdiJdcn dcr Dl'llhclwn ')odpobr-Expcdition reefa! crypnc commumm·~. P.r/,uwlfc>h•..:y 38. -t-t '\-70.
• ..J)J. o,,,, h \ildp,•lm J \f>nllfl<'ll 12 . Znol. -l. Zhura\'lev. Yu. A. Dcbn·nnl'. F and \Vuml. RA (I<JlJO)
2 ltenncr Bcrhn A synonymi,ed nomt•ru.:l.mtre ft>r l 1lnticd 'pongcs.
1, C W ( 1966) The nuau,tntuurt· of 'crornaro- Gc,,fo.l/rm/ .\la11a::mt• 127. SX7 •J.
Cnidarians
"' r
• nl r
me.m\ of .m10ebotd p'eudopodi.1 ; .lit~·matl\
the\ ~.-an extend H.1gdl.1 mto the Llltt'ron to '!'!
coments.
Corals, sea .lllL'lllOile\, Jell yh~h .md the small colomal fhe cells of the l'Ctmknu ,Jre more )ughJy 1hlfi
Id I
hydroids arL ,11l n:pre~enrat1ve~ of Phylum Cnidari.1. cntiated. Of the\e thL prinnp.1l types at~
They were tonnerlv grouped with the cten ophores, musculoepithelial celb- w lumna1 n:ll~ with\
'sea goo~cbcrries' m 'comb jelhcs' (Fig. 5.3a) in tr.Krilc fibres- and there .1re .1l~o ~eme cells lrJI
Phylum Cm:knlt'l ,H,I, btlt <.. n1dariam and cteno- to the nerve net below. In .1ddition then •
phores .m: mm rq~.Jrdt·d .1~ ~eparate though related ectodennal st111gmg cdl,, the ncmatocym f
phyl.l. Ctem.>plwrn .tn: ~;lobul.u or elongated gelatl- 5.1c,e). These ;m· l.1rge cells \\ith a ~calcd C1
nou\ org.1msms ~\\ unnung by 5Cn;cd combs of fu,cd cavity conr.lullng pm~onom fluid. within \dmh m1nut
Cilia arrangt·d 111 rmH fhe) catch food w1th rhe1r a t1ghtly lOtkd dong.tted tubular thread canvr
long tenracks. \\ lm h generally arc anned \\;th barbs and srykt\ 111\lde 11. A mJ.tll \ensory hair on
adlws10n n:lls fm l atrhmg prey; one species Jlonc mw;tde of the nem.ttory,t, till· cnidocil, '' 5Cilil
h,J\ \tln~ng n:lk Unt1l recently they were believed eo VJbranom 111 the w.ltt'r, .md \\hen .1 ,mall •
to be: the onh phylum With no fosstl record. but s~m passes cICI\L' to the nml.trian it trij!gcrs then
two 'Pl'cimem ti.Hmd 111 the I ower Devoman of mcysr ro d"ch.1rge. When thi' happen' the ~e
Humrild.:<.dlll·kr (Ch.1pter 12) .•llld rewaled b\ X- c.1V1ry 1\ lnt\: nsdy ulmpn:,scu by ~trou~ n
radiography (St.lllk) .md Stunner. 1983). c.\.tcnds tibrcs and thl' sl al bre.tb: tht· thre.ld c. shot out
the known r.mgc l.1r b.1rt... 111 nmt·. great force, tuming imide out as it extend' o
C n1dan.1m .lrl' the \lmplest of all true metazoans, cxpme the spir.1l b.trbs, '" luch pcnetr.lte the prC'f
but they art· of an t•vnlunon.Jry gr.1dc higher than rhat ir " lllJL't ted ,1nd p.traly,ed b) the po
spllngcs \Int't' che1r rdb ,ue properly organized 111 Bactent'' of nem.nocy'h .1re found on the tenu
m~ues whit h .uc nornully romtmcred on a radi.tl which all dl\tharge rogcthcr. The captured
p\.111. fhe t dl w,11l ts diploblastic, having rclh held f:1st by thl.' thre.tds, is <.onveycd to the mou
organ1zed 111 mo l.tyen only (Fig. 5. L1-c). the tcnt.1dc thcn bend~ iuto it. Further tlampo~
fhesr ,uc the nuter ectoderm and inm:r endo- food i~ by mucm slrillh""·
denn , wh1ch havt: no body cavity bt:tween them C nidan.1 .trc ch,mnerized by a life cydc m 11h hgure 5 .
but only .1 JCIIy-ht...e \tructurdrss laye r. In thi' successive gencr.ltions are or dt!Tert'lll kind~. r rnnsogl001
and life C)
mesogloea runs .1 Simple ncrvc net. The mcsogloea alternation of generations or temporal po~ lMeoftfi
ts somctJOK's inv.1ded by cdh from the t\vo primary morphism i~ typical of tlw Jc,, spl;'cJalizcd Cru I"'!Ccntly b
layer\. r hc ~lngk body t'.Jvity (enteron) ha, only but m ay he ~uppre\scd entirdy 111 the Boschmo i
one opcmng, tht· mouth, \Vhich also ~crves a:. an ·advanced' kinds. Two typt·, of 1ndividual. rbe
.mus .md i' normalh surrounded by a ring of rema- nully tixed polyp (Ftg. 5.1 ,1) and the free med
de~. lt i' lim·d by t•ndodeml whiCh is somenmc' (Fig. 5.1 b), .1lrcrn.nc \Ul'CCs\ivdy so that che b ld) lH
mfoldcd to form radl.ll partitiom or mesenteries, give~ me a\cxually to mt·dusae. whtch rep m g of
mcreasing the are.1 over which digesnon may take scxualh 'o th 1t the1r zygotl.'\ prodtll'C poh-p'>, :md dun Mt
place. tor eh~· pnnurv funcuon of the endoderm i~ on (f-1g. S.ld). plankl•lt
dtgesnvl' I hl' o:rll' of tht· endoderm ingest food b} A polyp 1s a \cdentary .mimal '' tth a cylm UIVCrtcd
Introduction 103
(e)
Jhre<ld_.
I
endoderm
cmdoc1l
pQISOn ,;,JC
(dl
mesogloea
manubnum (f)
QCISirOZOOid
Q-medusa
1
t
coenosar<
0
b
} .
...
planuld larva
·h ·~·re 5.1 la) Schematic diagram show1ng cmdorion hydroid cut longitudinally; (b) cnidorion medusoid with a thickened
""'IQioeo above the manubrium; (c) body wall of a cnidorian, showing details of the diploblastic structure; (d) Obelia- morphology
~~fife cycle lx 25): hydroid phose (leFt), medusoid (right) from below; (e) discharged nematocyst, showing spiral barbs near the
~ :lleof!he thread; (f) Milleporo (Cret.-Rec.) -longitudinal sedion (above) and surface view (below) (x 10): the small meduso has
'IJ :e-Jiy been budded off from the ampulla. (Id) MOdi~ed from on illustration in Borrodoile et al., 1956; (f) based on a drawing by
re ~in Treatise on Invertebrate Paleonlology, Port F.)
·r-
sa
'11 and an upward-l.tl.'ing mouth \Urrounded by a body is di~coidal, with the mouth prolonged
ce o( rcnraclc-..I hL 111L'sogllK'J j., comp.tratlVel} cenrrally into a downward-faung funnel or
.\ledusae arc tree-,\\'inumng inh,tbitant' of the manubrium. The mesogloca i) ven thick, and
ton. often very 'mall ; their oric..•ntJtion is bclo\v 1t the enteron cxtCJH.ls through four or more
cal ed rdati\'c ro th.tt of polyp,, Tht· medu~oid radial canals which JOIIl ,\ pcnpheral urLular canal.
104 Cnidorions
tomung a ktnd uf <.'Jrl"ul.nory sy~tem tor dis~olved ~cap~.· tl

tood m.ltat.ll. I he t<.•nt.Kk!> hang dO\" n free!)' ficd} .m
bcm·.1th the oLH<.' I nm, t'Jch muallv c.tnying at its The H ydruzoa arc: mu.lll) pol) morpluc, wtth r• dun· \C
ba~c an or~an llt b.1lanu: or \t.ltot pt. and often tctnmcral wmml.'tt'). ~cveral hvdroz C3l>ptul
~IIKC nwdm.1c art• the ~~.·xual .1s well as the dispc:r- order\ embr.KI.' .l \\ 1dc rangc of tonus: fonn dir
s.ll ph.1sc of tht· q:dr. tht'\ c.my tht' tour gonad,. eJtclo~cJ
ont' of wh1ch ~~ loc,Ht·d ,tlnng eat h of the pnm:tf) ORDER I HYDROIDA (C.un.- Rn.): H.tve cnlonul t lhSl]
rad1,1l c.tn.1k Wht·n till g.1mete\ arc ripe they arc pol\P' ami plJnktumt IIH:Ju,.ll' e .g. Ol•d1.1 Ordm:it'J
'hcd b~ rupture nf tht· gon.td w.11l and tcrciliZJnon ORDER 2 TRACHYLINA (?Jur - Rn ): Frn· mcJu,ac ouh, lo .Uitics.
polyp \r.tg<' !1.1\ tng lwcn \upprt•,,cd en tu cl} . c .g. Pc1.1•11 d.lltl rcrt
t.tkc' phn 111 tht· 't'a \\Jttr. The zvgotc then devel-
ORDER 3. HYDROCORALUNA (Tnt.-Ret ): Th~ setrttl
ops mto .1 ,m.tll tdwed planula larva . At tirst thi' arc oti:et
Lalc.ul'om ,k,·ktun .md Jrt• utio.:n rt·ct tumw~ . l'.g ,\1,1
has .1 \Dhd con· of t•ndodt·ml, \Vh1ch eventually StylcJ.\(t'T. {l>rvtHiopl
devdop' .1 n·mr.tl c.tvlt)', and ~ettlcs down to ORDER 4 CHONDROPHORA (l'rt't.tm.-Ri!l.) Thr hrdtt hydrt)td
bccumc tl1t m·xt polyp gcner.ltlon ph.t'l' ., .1 'inflt· l.trg<· poly )'I. tln.ltlll!-; .1l tlw ,urfJcc b\ l ~ tllh~: ( ~\..(
Mt·chi\Jc IIHWl' by rhyth111ic puhanon of the bdJ- ftllcd Rn.n. Reprodut ltvc ph.t't' ts .t 1t er mcdusa. r .g. I l }'OllClltS:
Iike body through l11ghh 111odificd mmculo- cpithc- l'llf]llfll. \\lilt h cc
ORDER 5. SIPHONOPHORA (l't'c.)· LJrgc: .tnd cornpkx ft,, I ltc poly
lial cdls. Tht·y nonn.1ll) die .tfkr reproducing.
ing colo ni c,, nor known .1~ los'il': c:xtn·mt· polymorphtsm
Though .tltt•rnatmll of genl'r.ttiom ~ccms to have mr orpot.
zoo td~. no free ~C\'ltJI ntt'dus.ll'. c .g. fJI1y.<alra.
bt•cn b,I,IC to the IIHifl' gcner.Jhzed Cnidaria, ~uch J\ ORDER 6. SPONGIOMORPHIDA ('1 m,. Jur.): Form m:u lorlll111g
mmt of Cl.1" ll ydrow.1, 11 h,t, heen modified or colonu:s wnh rath.tl ptll.lr.. tltlltl'd by ht.mzont.1l bJn, ~11br apt
'upprcs~oed tn others In Cl.t\\ )cyphozoa the mcdu- \trut'ttlfC\ Tl'~l'lllhhn~ tltl' ,lSlTOffllZ,ll.' of \lrt!OlJtOpOrotJs prob
\\,IS
'>Old ph.ht' ., J~lllllll.llll a11d th~: polypoid phase very w.ty th:u
redutL·d. '\ hl'fL".ts 111 Cl.1" Anthozoa the mc:du~oid Prt•brHcith
h.-1s bel.'n ehlllltl.lll.'d L'llllrd) .111d th1.· polvpoid pha~e Order Hydroido \\nnn,.
has bccomt· the 'c:xu.tl g~.·m·l.tnon. Jlr,1t11la
1\ dc'1. nptwn of tltl' nujnr divi,ions of the H ydr01ds arc rath1.·r gt•ncrahzcd hydrozoan~ Olllt:: o(t
Cmd,1ri.t ('•mpltfied) lollow,. wh1ch thcrc •~ ·' t httmom external \kd I cdmg ~
(perisar c). The order mcludes thrcc subor<b rum pro
Elcuthcrobl.mma (free \nhtarv naked foruh, nCIIl.llOC.:
pond Hydm b~1ng one). Calyptoblastina been .1 'u
Gymnobla,nn.t. R t·pre~t·nt.ttJve~ of thc lau~:r t.ttJon is
~ubordc~ .1rc on ,1\Jtm.tlly found in fO\s.J iort puhd~ .tr>
Obelia (Suborder Calyptoblastin.l, F1g. S.JJ), Jur.t\~ic tc
The.: Cmd.tn.J (Prt't',lllt.-Rec.) Jre metazoan~ with
st.lllcLird example of a colnlllal hydr01d, ha
rJdt 1J or bJr.tdi.Jl wmtm•try. They h.tve body walls of
hollow root-like structure for .tttachmcnt fr,
l'Ctodc:mt .tnd l.'ndoJ~.·ml \cpar:~tcd only by
which branching tttbl·~ .1ri~c, g•vmg ris~ Orde
mcmgloe.t; ,1 ~•ngk body 1.avit) (enteron) with a
polyp' (hydranths) alternately on each ~ide. n
mouth but no \l'p.Jratc ;tnm; a 'lmpk nerve net; and
pcris;lrl. i ~ .mn ular in place~. especially below cl I he hyd
no sep;tr,tlt' I.'Xt'rl'tmy or circulatory systems.
cups (hydro thccac) whi<.h ~ urround each puh .m hydn
Ncmatocyst~ .trc prc~1.·nt. Th~:y may be solitary or
Though each polyp is an inJividuJI, the pnh ,utd m.1y
coloni.tl, .tnd thcv arc nflL' tJ polymorphic with alrcr-
arc aU connected Logcthcr by a tubular sy~tr ..\ ft/lt'J'Il/t/
Jl.lt<. polyp' .111d na:Jm.tL' Mall)' have calcareous o r
(coenosarc) through wlud1 the enteron contlnu ll\IS 'kdc
org.mtt' \kektom and they .lrl' frcqm·ndy coloniJl.
E ach polyp h,J\ many lenwdt·s and a prom.inend pIll HI Oil!
Thnc Mt' thrt•t• d:t~s~.·,:
bow. funnel or manubrium upon which the mo an.tls.
ClASS 1 HYDROZOA (l'rt't'.lm -Re• ) \n· below i~ ~et. Toward~ the base of the colon) are cyhndr J he Sl
ClASS 2. SCYPHOZOA (Prt•,·Jm - Rn ) Mo,t brgc Jcllvfi,h. gonothecae: Aa,k-~haped 'trutture~ wtth J c kdeton,
CLASS 3 ANTHOZOA (l'n-c:Jm.-Rcc ) l.orak ~c.1 .111cmonc,, stncted aperture. Wtthm eadt of these ts a a the skclcl
~llr¥Omlns, ,~,·a p( n, , ~tcm Jn\mg !rom the coenosarc, and from th11 UC'JI tube
mc:dusae £om1, bemg budded off continuaUy :6
the upper end of rhc stem ~ they nlJture. Th
Class Hydrozoa 1OS
re through th<.• ap<.'rtllfl: .md thereafter \Wim
eh and grow uno! thl.!y are old enough to repro-
e 'exually Gorll)thec.le ,lr<.' ryptcal of the
:ptoblastinJ; m the Gymnobl.t~nn.1 the medm.tc
directly from the coeno\,lr<. wtthour b<.·mg
d,,1ed, and then: t\ no hydrorhccl.
!o,\Uchionom hyJrotds ;lre well known in d1c
.lomian anJ h.l\'l' been rl'l orlkd fi·om ~ever.1 l
u~. The cak,trtous tube' ot 'l'\ lTJI Mc\OZotc
Tertiary serpuhd tubc-wonm (e.g. ParSIIIIOIIi,r)
otten tound mfc:steJ by .1 wlomal org-Jnt~m
tu/,Jp!lila) wluch h.1s been mtc:rprctcd .1s .1
.lrotd buried in the outer part of tht: calc.lrcou'
~crutton. 1975; ftg. 5.3b). Thi' has two t"Ont-
!rnts polyp dlJinbt:rs, and \tolon.ll net\\ orh
h connect them m a regular sc.:ale-hke pattern.
, polyps grew .1round the run .111J were probably Figure 5.2 Mi/lepora, a Recent specimen from Barbados
(x 0 .35). (Photograph reproduced by courtesy of C. Choplin.)
nrporJred into tht' tube as the scrpulid t,.•n·w,
ung polyp chambers openmg through ~cnlllir
lperture5. rhts hydrotd-\CflHtii J d\~OCl,lt\llll SUrUCe Jlld capablc of retracting IntO the tube:
rrohably svmbtotlc or commemal, Ill tbt: ,,unc the gastrozooid s, whtch arc purdy manubrium-
dtar several spl·ues of the modem hydroid like mouths; the dac tylozooids. which are elon-
hid,Trtyla :trc symbiotic \\ tth tubular sabc:llid gated ten tacles w ith batteries of m:matocysts; and
~ 11 \. the ampullae, which produce mcdu,ac. All these
ru!opltila would have been ,1blc to teed on are com1ccted by soft ussues nmnmg through the
of rhc supply offood parttdcs drawn in b} the honzomal canals. Thi\ kmd of colon) illmtr.ttes an
lmt anns of tht• wom1, whcrl'·'~ the serpul.J<.l in effecttvc 'dtvision of labour' and comcquent modi-
·11 probably derived ~ome protccnon from the fi cation of the zooids.
lllltorysr battencs of the hydrotd. It must holve Millepllrtl in its natural reef habitat is very variabk·
~ \Uccessful associ.1tion. for the degree of mfe,- in fonn. The ntillcporcs of Barbadm (Steam and
,, 'omcumc~ as htgh as 95% :md mfestcd \c:r- Rid m g. 197 3) etther encrust dead coral~ or gorgoni-
dl lre found in bed~ ranging from the Mtddle ms m th m sh eets le~s than I cm thick, or fom1 'box-
,,ll to the Pliocene, 170 Ma m :1ll. work', t.<.'. thick near-vertical blade~ joined along
their edges to fom1 box-ltke, mbanbrttlar, upward-
opening cavincs up to 30 cm luglt. Alternatively
they m.1y be bladcd, 1.e. wtth erect blade~ buttressed
by other blades, m akmg colonies up to 50 cm high,
hvdrocoralhnes (nulleponncs and sryla~tcrincs) or branch ing, up to 20 cm high, w1th delicatt: or
hvdrozoans which superficially resemble coral~ stubby branches.
~' J} be qum: unporrant in ~umc: modt·m reef,. lt t\ pm\ible that the boxworl.., bladed and
p.w (Figs 5.1 f. 5 2) has J th1c J... IJntinar c.llr,Jrc- brmdung forms ,ue dtstina notl-lntcrgradm g
skdeton of lll.lll) vcrtic.tl tube' with cro"- bio~peoe~. bm the encntstmg fonn wade~ into the
nons connected by thin honzontal ramtfymg others and i~ probably Jn environmental growth
h h fom1 of the o ther species, being most common m
ll !he soft tmues uwcst only the L1pper part of the ~hallow w.tter or cncru~ting gorgomans. The habitaL
leton, degeneraung m the older lower layer\ as preference~ of the other ~pecics seem to be governed
~clcron bUJid' up There are three type' of vcr- by water rurbulencc. bm:work fonm being the
c tube. each wtth tts own zootd .1ri~mg from Lhe strongest ,1nd b laded the most delicate.
ll
(c
106 Cnid arians
(c)
(a) Pleuro brllcht ll (b) Parstm onia
lh
UltJn: l )
[I 1\Cd I'
'" g I v<lr oo<J F1g. 5.

~trnhil
l 'lHs
uuuall)
•t t)(l~l
ntop
lt cc :IS
jdlyfi~l
(e) Gorgo ni• (g) Fe\\
tJHSl' , •
(k )
(h ) P#Hin lltul•
11) Peraco nularia
like, fused plates of cilia entero n and tentacles, which have jull
Figure 5.3 (a) A ctenophore, showmg four of the dcomb· - to the right 1s on illuslrotiotl
infeste by the symbiotic hydroid Protulophila (x 1)
copepod ("' 1); (b) Cretaceous serpulid worm arran9ement of l~
ds. The warm tube below is exfoliated to show the
ing detail of the rim of the serpulid w1th living hydroi scyphistoma releasi ng ephyr a larva; (d) internal
ating scypho2:oon
chambers linked by connecting stolons; (c) strobil (e) a Recent Gorgo nia with visible polyp sJsee (f), (g)]; (0. (g)
Kimberella, o probable Precambnan scypho2:oon (x 075); into orga nic coenen chyme with centra l axis on spicules ( · 2001
set
through gorgonian stem with eight·armed polyps affinities, (k) Tub/F.
Pennatu/a, a Recent sea pen (x 0 25); (j) Poroconulor ia, a conulariid (Cam.-Perm.) - possib!Y of cnidarian illustrations by
1975; (d) redraw n from Wade , 1972; (1), (g) hosed on
Recent red octocoral (x 4). [(b) Bosed on Scrutton,
Treahse on Invertebrate Paleontology, Port F;
(j) redrawn from Slater, 1907.)
Class Anthozoa 1 07
~q phozoam. mdt•pl·ndentlv .11 qumng .1 phosphaac

t'Xo,kcleton .
Conulariid, .trc not oti:l'n prl'~l·n·cd hcnme they
\nphuLllJ .lfl' tn~ l:-s\\ tmmmg ml:dm.u.:, broke: up t'J'il) atll'r tk.Hh, bm the: pn·seiKl' of \'cry
rm m.tnll\.' .md u'tully \\ tth raJul \)m me try numcrou' fr.lgment' dl\,ol\'t'd our of lunt''itonc'i in
on a tetr.uuc:rm" pl.tn rlw s~:s'1lc: polyp ,tag~ Godand rc:,ritil's to thc:tr tonner import.ltKC .h J
i3c) 1\ c.1lkd .1 scyphiMorna. \\ hKh buds by conuuon componc:tll of Pal.tl•nzorc fnm.\\, When
ruobilation . they arc found "hole, thl')' nuy he ,oiJt.try. m
Tht1 mca1h rh.11 to\\ ,1rds th uprt•r t•ud 11 1, coo- occur 111 mono,pt•otit· (pm,ibly don.tl) ·'"cmbl.tgc'
lly grm\ tng and bt'lll~ dt\'IJed bv tran.svcrsc att,tdlc:d to tube, or n.lutiloid 'hdk
:tOO\c,, 'o th.ll .1 ptlc: ol ,m,tll mt•dw••tc: tonm, one
top l•l the otht r .•md .lfl' rdl'.tsl:d 111 turn to bn.·ak
a) cphyra larvae . the JliH'mh.· , of thl' adult
:m h.
f~w Jrllyttsh art tnund m the tos,tl retord. Of The Anthozoa .tre pol} P' wnh no lrau: of .L medu-
&, somr .lrt tou 11d ·'' l' omprt' \\JUils in very tine: 'oiJ sta~t·. Anthozoam (torah, Sl'a .mcrnonc~.
,hmt•nt; orher<i, tnl llldmg .11 le.l\t sonH~ of the gorgouiam ,md sea pens) .1rc soiJtary or tolonial,
Jrar.~n fin111J, ~re prt''t'l'\ t•d .ts ,,md mtillings of eutirdy manm· cnidanans. Thl' polyps produce
~~ CJ\ tt\ (bur \t'' (. 'h.tptt'J 1). The Pr~:cambnan gametes whtch develop Jirctdy tnto planulat larval'
•wm·/1,, lron1 hil.1<.1r.1 1' prc:servt:d .ts pmmvc .1ft.:r fcrtihzanon and tlwmc to .1 new ~enc:ratJon of
upositc moulds. lt ., .1 pnmllr\'t' styph<lZOJn of polyps; the mcdus01d v;u~t•rauon h.ts been c:limi-
Jm<' fonn (\V.1dr. 1'172). po"thly rd.ttc:d to the n,tted enttrdy. They rcsembk hydroLo.ln polyps rn
grubomedm.1m ..111d thl· gon.tds .tdhered to the many re,pe<:t~. though the) .Ill' otlt·n Vl rv brgc.
ltanJl Jnd pn~kttt•d tnto th~ Cl'lltrJl l'.lVtty of They alw,tys h,l\'l' ,1 tubul.tr gullet or stontodaeurn
bell. leadmg down m to tlw l'Htc:ron. \V htt h hydrozoau
'"~n ~oup of prnbkm.mc t(,"!ls. tht• ConuJat.l polyp~ do not h.wr. 1 ht• m tenor Jtwlf ts dJVid~:d b)
ubnids) (C:un .-·r n.ts) .tppl' 1r-- robe of ~cypho- ra~.hal parnnom (mc,l'IHen~.·,) whmc number and
3rtlnmt-s. ' J ht'\1..' .lH' \tl'l ply pyr.tmtd.tl pho~ morpholog) h unporr.mt m the ,ubdt\'tslon ut the
to"tl' (ftg. ;) .J,), \\'h1d1 h.1v~.: .1 qu.1dr.He class.
-sccnon .md otrt·n nurkt·d hl.'mnghone ndge~ Those rh.1t 'ecrt'tl' h.trd p.ut~.•md L''pcnally the
nth~ 'ides. \~'hcre.1' rht·y h t\'l' bt•t•n rdic·rrl'd to corals. ,trc of grl',lt gt•nlog!c,tl unport,mcc. oti:en
us h'Tuups (t hord.rtl'\, .uwdtds or .m cxrmct tormmg rhtck hc:d' m tltl.' i'.li.JL'07ntC .tnd somc:umes.
.mn, thbnwk. 11J') I). rt'< nmrrurtcd crms-sec- in asmo,mon wtrh ~tronutoporotd\, n:ct:hke
through f,l<'clllll/•111•1 ftlc ,/,,,, lrom rht• ~tlunan of m,ISSC\. rrom fertury tlllll' Oll\\".lrds true LOr,tl red;,
t!Jud. \how ,1 tin•r-ln.IIH ht•d llltt'rll.tl p.lrtltlonmg of v,tst thicknesses h.wc tonned, .llld n11 als lt\ tng ~~'
maJll~ iJentic.d to t h.1t ot lt\ ing comnatt•d reef tom1l'T"> ,m.: proh.1bly rnorr nnport,mt nov.. than
hozoam Ot·rre, !CJ<JJ, I'J'J.f) ltkt'\'.t~l' rn thrce- at .my oth<.:r tlt ne
m,·mton.tl <onulJriHh front tht· USA (v.m Iten. Anrhozoan~ .1rc t,rrOUJX'd tn thrt'l' subclasses:
ll; \'an ltt•n t'l cl/., I 1) 1)(1) intt·m.tl 'trueturc\ .m~ Ceri,mtip.lth,m.l, Ot tocor.l llt,l .111d Zo.mthan.l. Only
Mmilar, .md ntort'O\'L'r tlK· ultr.!Structure reve;ll~ the \,1\t of rht·sc, whtclt mrlu de~ tht· LOrah. wtll be
;mkmg rcsembl.uKt' 10 t hl' tht•t.tc..' of coronatt•d constdrrcd in der;ttl
rbilltldll\, ,lllJ ,\11 L'VHknt \llllll.lnty Ill modt• of
,ht 0
lOW· IUlllld tnjury rql.lll. rhcrt• )\ \lllllt' t'\'tdc:nct' of
.olyp 11ltion .llld, 111 thn:c:-duut' thton.ll spcnmCn\ Subclass Ceriantipotharia
d of t~pam, oflon~illldtn,tl fi"ton Pm,rhk tt•ntade'
dion
t; (h) reen rl·portt•d bdov. d!l' pyr.llllldal \tnJCturl' Cerianripathana ,trt' soltt.u-y nr coln111al pol) P'
ro,o rue umancc\ ,111d Ollt' r.uher tlutened gl.'nu' w hi eh tor rnorphulogiL.tl re,l\nns an· pl.Kl'd .tp.trt
·er in 'l'dlt~) r~:st·mbk, .t lllt'dtl\.1. Conubnid~. from other group\. Tht·y .trc: \ trlu.tlly unknown J'
ton: now 'l'l'lll to lk· rcl.nt•d to l oron.ne fos,ik
108 Cnidarians
ORDER
Sub<:lass O c tocorallia Subclass Zoanthari a: corals ROER:
ORDER:
Octocor.~llla (?Prec.nn., Ord.-Rec.), poorly known The ~trucrure of rhe Zoanthana " ewmplified
ORDER •
Js fossils, .tre c~pccially represented by the gorgoni- the Upper .J ur.~ssic to Recenc sohtary ORDER:
ans (Order Corgonacca) ronunon in many modem Cnryvphyllia (Order Scler.lctini<~). Rect•nt ~peoc1 ORDER 1
t.Or.ll red-; (F1g. 5.Je,f,h). The colonv forms a flat fan of co~mopohr:m dtstribuuon, living on \,tnou ORDER i
of anascommm g bramhc~. The ~kcleron conmt' of ~trates and ar depth~ of o-17C)O m. C ~1/tllhii In ORDER f
horny br.mdung tubes, each of wh1ch may h.we a cool, temperate water~ otT western Sco''
central calc i6ed core. From the outer surfuce of the (WiJson, 1975, 1976). In regions of strong ndal•
branching tubes proJCrl the many small polyp~ rent!> 1t is found attached to pebbles or boulder
(autozooids). each of which ha~ e1ght renr,ldc'. in weaker currents on the outer ~helf H can lt\· Ord
char.~ctcmtit of the '>llbdass. Thc~e autozoo1d' arc variety of subsrrates; the rnost conunon too
sct in .1 rhirk gelatinous outer coating owr the attachment in sand parchc~ is to calcJreom mbl Mor.
branching cubes, wltlun which are ~er innumerable the wonn Ditmpa (Fig. 5.4a). rht R
ca!careom spiculcs wiUl h help ro support tht. .tuto- fhc polyp of Caryophyllla, whith ts 'ome 2-J <•lna
zoolds. These are the on!) p.1rt to preserve as fos~tls, acros~ (Fig. 5.4b), i~ many-tentac led .md ha~ a \ lnl1ll'
and only trom them c.:an the fornler presence ur gor- modacum le,1ding down to rhe enteron, whi.~ tlllliH'r<
gomam be interred. divided biradi.11ly by numerom mesentencs. EJ, S1 1h f
The fm,tl rerord of occocoral~ 1s spar;c, bur these has fiillcd &cc edge,. The sofi: lma.l thc•ngh
recently organiC skdcrom believed to be of gorgoni- secrete an aragoniric cup or corallum wh; 'f ~epr.
am have bc:cn found 111 rocks as early as the Lower short .md hom-~hapcd. lt has ,111 outer ~~Ill .lt
Ordovician (Lindstre,m, 1978). Furthermore , abun- (epithcca) within which arc numcrou~ ra show tl
dant spJCules of Lower Stlurian age, u~ually ISolated, arr.lnged septa which IJc between the r SJ'I'l'ld
hut someumc' found 111 nc'it-likt: a~rregatiom, have mesenteries. 1 'he firsr-tonned protosepta 11 psut
bcm shown to bdong to o~tocorals of Order SAc) arc larger and more pronounrl·d than "t 1rnl ', •
Akyonacc.1 (Uengtson, 19Hl). These: warty, spindle- n1ctasepta intercalated between them. fh c
shaped sp1c ult·s of the genu\ Acrartosel/,1, once thought When preserved as a fm,•l. tile aragomtc b, oVert'C
to pertain to sponges, arc Vlrtually mclistingui~hablc altered to calCite or cli~solved compkrcly b1, 'h1ch,
trom thmc of modem .1lryonaceam, hence tlm group negative mould 111 the matn.x. Ar.~go111nc ,keJ LJL Ul
ha.~ much more anncnt origins than was formerly arc known as tar back as the Tnassic, :111d almCI thl 111\l
believed. -1tmrt,>sella prob;1bly lived on a san~Yravcl Scleractinia arc aragonitic In Cary.,plryllra, a~ 11ll del
Sclcractinia, rhc septa arc l:ud down cvch~ Rug,,,,
'ea floor, lt was not J rccftormer
HeliopMII (Order Cm:nothecal ia) has a mass1ve muloplcs of \IX. When the cor.1l1~ very ) our. mcnh (
comtlum is simply a basal plate w1th \IX '1mll t il
.tragonitic skeleton, brip;ht blue: in colour. Tt is an dl:l
nnporranr reef fonm.·r in the Indo-Pacific proVlnce. epta inserted between the mc~entcrie,, Latt C'f'.WJtl
lt\ sUJUlant\. to tile tabulate Hclwlitcs has suggested mcuscpta appear, and there may be t\\o N honzo1
that the !.mer 1s a Pabcozoil o..:cocoral, but tlus gc ncr;~tions of thc~c so that the whole hero me' 7..aplu, r
rL•semblance 1~ far more likely to result from conver- riscptatc. A shght shift of thctr sp.King thwu~;! I he
growth as the roralJum beLomc~ cltipllral im ~;.!lau,
t•
gent evoluuon.
The red octocoral TulllporcJ, the ·organ p1pc' coral the charactemul biradMI symmetry Thw hl3turit
(Order ~tolomtcra). lw polyp' mhabtting long growth new material is added to the t·xpo,t-d I hcu 11
homy tubes (Fig. S.Jk) .tnd often fonrung large of the sep ta .md to the cp1thcca \O tl1.1t tbc
rolmualnu1s~es. The ~~.-a pens (Order Penn,ltu lacea; expand\ a.~ it ~rrows.
Fig. 5.3h) .m' another k.md of ottocoral. Here the In broad and general renm the snnpk mud
lolon)' has the fom1 of a leather, the base of,, htch 1s Cllryoplryllw applies to ll\0\t IJ\ ing and fo,"l
\Ct in mud wh1le the feathery upper branrhe' :uc but there .m: maJor difference'\ m ~kckt.tl \trU
hneJ wllh .1utozooids. The Ediacaran Cllnnlhnlisms septal arrangem ent and mode of colon) fonll.1t
Eight o rder. arc now defined (Scmtton, 11)4"1[ IC'~(If bl'C
Ius usuall) bcu1 mterpreted a~ J 'ea pen (though sec
hte .,f t1
Chapter I tor .UtcmauH· view~) .tnd tf so mdtcates
e.1rly ~Utlt:~' fm rills group. Will·
Class Anthozoa 1 09
Efi 1. RUGOSA (M Ord.-U. Perm.) rum dlVtdcs tt (Fig. 5.-tt). soon becoming ,c.·p.lr.nt:d
~~ 2 TABUlATA (l Urd.-U. Pcnn ) mto cardinal (C) and counter (K) pro,t·pt.l. fwo
roEn SCLERACTINIA (Tn.K-Rer) other pam of pro~cpta follow: the alar (A) ld.pcem
DEl 4 HETEROCORAlliA (U Dcv -l C uh.)
to the cardmal \Cptum, and the counterlateral
DER 5. COTHONIIDA (M. <. .tm )
:P.DER 6. KllBUCHOPHYlLIDA {U. Ord.) (KL). Even at thh &tage some bilaterality t~ ev1dem,
~DER 7. NUMIDIAPHYLLIDA (l'cnn.) which becomes more pronounced when the
OlOER B. TABULACONIDA (L CJm.) metasepta an: in~erted serially in four quadranb
only, on the 'c,1rdinal' ~ ide of the al1r ,111d LOUntcr-
lateral septa. Through differential gro\Hh the wun-
t terlaterJ.I and alar septa move towJrJ, the
Order Rugoso counterseptum to make room tor new met.t,epr.l.
Short mmor sepra are laid down bet\\ecn the
Ncrphology metascpta. Around the cardinal septum thert: 1' a
. Ru~o'a .1rc an exclusive!\ Palaeozo•c group of cardinal fossula where metasepta arc not JllseneJ,
ll llfY and tUioiUal corals. They \how bil.ncral and especially dnnng the 111termediate growth stages
~ mmetry 111 a prinun\'e m,111ner, arim1g bl'Cause the there are la.rer.1J (a lar) fossulae as well. The~e JfC vis-
IS uncrou' met,m:pta arc inserted in l(n•r loci alone. ible in the adult Znpl!rcnticcs. but m many large
f eh bliatenllicy ts clear in .;omc ot lhe Rugosa. Rugosa, wherl' the major and minor septa have pro-
es u~h 111 other~ it i~ ob~curcJ b) the prohfL•ration liferated greatly, the fmsulae are so compre~~ed as to
IS lti'Ll. ,mJ 111 genera ~uch ;:1' Hc.,-,,~'''""1<1 110 pat- be hard to d1stingu•sh on account of the mcn:.t-.ing
~~ tll •~ \'hiblc T\\0 examples ,lrl' d10\Cl1 to radial dimibutton of ~epta. Many Rugo,a, e.g.
,, the hNl ~rrucmre of rugo~c cor.1k The widc- Pllillrpsnstmctl, have no rrace at .t.ll of a fo~~ul,l.
J Carhomferous Zaphrwtitr.1 (Supcrfanuly In Zaplrrmtrtt•s, as in mosr other Rugm.1. two
!:11ophylhJae; F1g. 5.-ld-t) is ;:1 \mall. \Olitary 'horn other skelcnl elemenrs are (1) the epJthl'ca. Jnd (2)
, \0 c.tl!t·d became of it' ~hape tabulae. whtrh are lhr horizontal plates fornung a
fhc outer hom-~haped part of tht• lorallum is floor to the c,wity in which the polyp restdcd. The
rred by a rhjn cakareom skm. the cpitheca, maJOr ~c:pt:l join together in <1 centraJ boss forrncJ uf
'' h t'XIt:nd\ from the tip to the uppa (distal) sur- updomed tabulae .
. or ea lice, wherc the skeletal elements filling Heliophyllum (Subt3mily Zaphn:nttd,lc: I ig.
1111ide of the: coraJJum are exposed. These: inter- 5.4n-q), ,1 l,trgc North Amcncan mgos,m. 1s ~ome
clement' fom1 the basi~ of da~sttl\.atton of the time~ found .1s poorly developed colo111es. In tlm
.igf•l.l 1nd arc of two main types: tht. vertiCal de- coral there ,tre \O many scpta that it IS not ea~y to see
lsepta md axial structur e) and the honzon- the four m,tin line' of emplacement. The ~p.Kmg of
mcn~ (tabulae), whteh will be c.Onsidered the septa j, mort• or less equidistant, and the 'YJllme-
rhe rdy . tDissepiments. "luch arc m1porranr try is virtu.tlly radio~!. There are some mucrural ele-
)Ii'C zonul demems in mme Rugo)J, Jrl' lacking in ment~ not found in Znphretrtites, e\pCcl.llly the
1Ul- lrllflft'.l.} dissepiment~. which are concentnted 111 a broad
DOtH fhc \Cpt.'l Jrc thin vertical piJtcs ;~rranged m a marginal zone or dissepi.Jnentariu1n . fhc~c .1rc
}arts ;ntttcri,ttc btradial pattern, v,hich develops to small curving pl:Jtes located between the septa and
10\ll Jnt} throughout the ontogeny of the coraL set nonnal to them, inclined downward~ ,lt abour
0~1.:' r rnmnn of insernon can be stuthed by mvesri- 45° to the: c:pithcca. The major septa may reach the
:or,tl •• the ontogeny of the cor.tl from the early centre, m which updomed tabulae are the most
: thl\ '' mually done by makmg 'enal secnons pronunent component.,. Another charactenmc fea-
re ot tu tht· .1x.i\, from the tip to the l'ahce, and ture ~ the presence of carinae: short 'yard-.11m ' ba~
ilrak gJilK thl'm in a succemonal ~enc~s. Tlm pro- proJecting laterally from the septa.
nnc. ~ iull record of how rhe coral grows. smce Though bilateral symmetry has been ma.sked by
tic.'n 1urr1 once formed by accretton .tre not radial ~tructure, the cardinal fossula can mll be seen
): ••hd hut retain their origin.1l fonn through the where the di~scpimcntarium narrows and the axtal
d the coral. ends of neighbouring ~epta arc shortened and curve
I); h~n 7,aplmmitrs is very young ,1 smgle prosep- round it.
110 Cnidarians
lql
(b)
(a)
en1ocoet
1•-cl Caryophyllill
Id-f) Z•phrentites
ea lice
(d)
(e)
c Ml
Cllrdrronl
fossula
IQ
Kl
~I
~\LJ1,
ID CD CD \iDY c ~:...
C A
Class Anthozoa 111
(p) H11l1ophy/lum
cut sl!ptal edofls
'--v---' ......_,......
" " tabularium
dissepomentarium
(n)
l
M2
''¥' 5 J (a) Young Caryophyllia, o Recent scleroctinion coral attached to the tube of the worm Ditrupa; (b) Caryophyllia vertical
~
Ml
ml l!dlon, (c) tron~verse section through young (centre) and older (right) sderoclinian coral I 1, protoseptvm; 2, 3, metoseeto); (d)
MJ ZDpllrentiles (Corb) with port of ttle epttheco removed, showing bilateral symmetry, (e) some, in transverse section, (~) septal
1\ ~I \hown by serial sections from lip (C, cardinal septvm; KL, counter·loterol, K, counter·seplvm, A, alar); (g)-(m) examples
J~ Zophrenlltes delonouei group (Corb.). including (g) Z delanouei, (h) Z. paro/lcla, (j) Z. conslricto, (k) Z. disjuncto (early) ond
MI Z disjuncto (late) (not to scale), (n) Heliophyllum (Dev ), transverse section showing neor·rodiol symmetry (x 1.5); (p) vertical sec·
mI• 1.5), (q) enlargement of peripheral zone showing relationships of sepia and yardarm carinae. ((a) Redrawn from Wilson,
976, (d), le) based on Milne·Edwords and Hotme, 1850, (g)-(m) from Corruthers, 191 0 ; (n)-(q) redrawn from Hill in Treot1se on
l!ltrlebrote Paloontology, Port F.)
0 0
Dendroid • Phaceloid Cerioid
Amural astraeoid Amural aphroid Amural indivisoid
~"55 Transverse sections through compound rugose coral colonies, with terminology of different types.
112 Cnidorions
(b)
(a) S
Lithostrotion
(d) (e)
(c:)
S. JUnceum S. arundineum S. martini
Figure 5.6 Lithostrohon, a Carboniferous compound rugose coral: (o) transverse; (b)long,tudinol section; (c)-(e)laterol inc!OO'.l
three Siphonoclenclron spec1es, S juncei.lm, S. orunclineum and S. martini. (Redrawn from Jull, 1965 I
Range of form and structure in Rugosa which a \lngle ~l.;clcton (cor.tllum) is produced
FORM, TYPE AND HABIT OF CORAUUM the lit"l· activine~ of numerml\ adjacent polvp'
Rugo'>J .m• ctthcr solltarv or colonial (compound), conrributmg a corallite ro the whole.
and ctthn form may eXIst even within closely The h.1bit of the colony may be variablr tru
related gt·ncr.t or cvt:n ~pcctc~ of the same family. l n enccd by the environment. but colony type, \1
'olitary Rugo~.1 (Fig. 5.4d) the usual shape is a includes the morpholob'Y and rcl.ttions of the <UJ
curved horn: the ccratOid shape of Zt~plireutites. If lites, is more rigidly defined genetically. In thd
the 'cone' ha~ l.'XP<Indcd very fa..~t. a flat discoidal ciculatc type the corallite~ :m: cylindric.1l but mlG
~hapc ma} rc,ttlt (Ptg. 5.7c); with progressively less contact. There are two kinds of tJ.snculJtc 11
abmpt l.'Xp.lmton patellate, turbinate (fig. 5.7d). phologies: dendroid , with irn:gular branchr~
troch oid .md cer atoid forms will be the product. phaceloid, in which the ror.1llitcs Jrc more ot
Cylindrical corals .1re vmually ~rrajght-sided, parallel and someumes jotncd by conne
except lt1r the liN-fom1cd part; scolecoid tonm proces\C\. Massive cor.ll' .trc tho)e 111 wh1th F"l()utii ,
corallttcs arc so closely p.ICkcd a\ to be polygo PaiOOOC)
.trc Jrrc~rttl.trl} nvMed cylinder;; pyramidal types lunon
haw \harph angled s1dcs; where~ calceoloid ~cction. Several kmd~ of nu,sive corals arc
gencr.1, oddt·\t of all (Fig. 5.1 Oq), pos~ess a curved brui~hcd (Fig. 5.5).
c..orallum 'Aith one A.mcned ,jde and a lid or
oper culum cerioid (e.g. LuluutMiml). in \\"luch each oo
Colomal Rugo~.~ (Ftg-; 5.5- 5.8) are those m ret<.lllh tt) wJ.ll:
Class Anthozoa 113
'~ S1phonodendron (b) Siphonodendron (f) Kodonophyllum
((e) Acervularia (Sil.)
(g) Entelophyllum
u- Q~abdocyc/us (turbJnate)
~
:IJ.
tl-
IS-
in
Jr-
)§'
nd
n 5.7 Fonn of corollum in Rugose· (o) Siphonodendron, fosciculote dendroid, (b) Siphonodendron, phoceloid; (c)
Ill 'T.eocyc/us (Sil ), discoidal; (d) Rhabdocyclus, turbinate Form; (e) axial increcse in ceriod Silurian Acervularia; (ij rejuvenescence in
WJ~ Kodonophyllum, (g) ofhets in Silurian Entelophyllum. (Mainly redrawn from Milne·Edwords ond Hoime, 1850.)
111-
lite
114 Cnidarians
astl'acoid (c.~. J>hi/lip~.wwca}, m whteh the mg cor:~llircs an: tlnltl'd hy .1 7nnc of d!'.,cpmt om
cpHhct.l h wholly or paru.illy lmt but the septa alone; llll
\(,IV unn·dutcd; indivisoid (t'.g. Oricllw~rr.rer~), 111 \\ hu:h 'CPtl Su
thamnastcrioid (e.~. H.rpiMircci,z). m which the ,1b~cnr .111d dis\t•piment.ll material '' don ft.l
scpu of ,lt~.ltt'IH tor.tllitc~ arc confluent and often rlw, type 1<1 \try uncommon . dot
s1nuom or t\\ ''ted: thctc
aphr oid (t·.~. Ar.ulnwphyllum), where the septa FINE SKELETAL STRUCTURE OF SEPTA ( bJ
.Ire n:duced .H tht·lr outer end~ ~o that nc1ghbour- The \epta .1re cmnpmed of \enally arranged ra A!the
ing whorl' of tiny fibre' (sckrodcmlitc,). grol!j 11'111)1
111 C) limlric.tl bundle, known ,,, tr.tbetulae the: SI
5.9). lllll
Tn \OI11l' cor.ll\ the J.Xl'\ of thc't" bundks are
pier and p.1ralld (mon.tt.tnthtnc trabccubc), \\hilt fYP
others then: arc 111 .tdwuon 'ctond-order trabeL'U l u th
radiating upward~ and out"' ards (rh.1btlacanrh a lllt'lll
rhipidacmths). ThL· sderodennne~ are ~ecrcted I tgh
linear series from .1 contmu.1Uy producmg centrt lut p
calciflcat ion: the tr.tbccula t,rrows a5 long ,\'i calnti tile t1
tlon got:\ Oil. 'l'rabeLUJar tlSSllt.' is pn111.11')' Jrr• st
Rugma, and uthl·r l.ind~ of tissue that h.we b 11 >l f)
described (tlbru-nonn.1l .111d lamellar) rmy b~ Ntdrtl

re\ult of ~:.rrowth llltrl'llll'nts or d1,1gene~is .md tru r rts
fore \CLOnU,lf) (SorauC 1971) Jr(' , c
The ,J..elcton of Rugo\J prob.1bly origm.l!h l lu
ststed of htgh-Mg olote Th" generally nunt\ 1 uaU
original minmtructurc\ .tli:cr death, though rugo
htgh-Mg c.tkite li\U,tlly convc:rt~ to low-Mg cah vtdcd
Ftgure 5.8 Actmocyathus florrfonnis (L Corb.) from Bothgote, at the e.trlie,t 'tage of di.1gcne'i' ..1fier \\h1ch trab~:~:
Scodond· o smoll colony showing loterol increase with o few
small encrusting Syringoporo (x 0.75) (Photograph reproduced ongmal mtm'r,liOg) may ,llt~c"r . Ar.tgonite. ho\\ nd ~
by courtesy of Jeremy Jomeson.) seenh £O luvc bt•t•n tlw pnm.uy skclcr.ll mml"l21 the
td)ICC:
fluent
'"lucl1
prop
~UII1u
AXIl
\ ·ry
:; It 1.1·
llCJ1
broad
111 om
l 11 tf[j
Figure 5.9 Fine structure of septo in Rugoso. (a) monoconthine trobecvloe, (b) rhobdoconlhme lrobeculoe (holoconlhs ore~ olurn
bly reaystollized rhobdoconths or monoconths); (c) rhipidaconthine trobeculoe. (Redrawn from Scrutton 1986.) Olllp<
Class Anthozoa 11 5
s n~ Penman Rugo'a 'hordy hl'lort' their c:xtinc- the lancr genu\ it is divided b'y ,1 c~o.·ntr.ll \t'pt.tl plate
n (\V llldt. I 9911) tonm:d by thl· counter-septum. -iu/11hl h.t\ .1 Vt..'l)'
Smc~ the- gro's morpholo~ ol t'tlr,lls I\ so lughly sjmpk axul ~o.olumn. the aulos. wluch '' .1 't..•rticJI
r·
• nJblt• and h.l\ been 'l1bject, m 'o m.my mdeprn- rube trurH..ltlng the innt:r c:nJ~ ut the ~~.·pt.t ,llld tr.l-
111 ltod;,, fl' rcpc.1ted conver~enr cvohmon, vcrscd by honzuntal t:tbulac:.
1~r~ ~~ .1 llt:.lr nt•t..•d in taxonomy w tmd more:
·u.1bk·' rh.1r.tncrs to ml' ·" tlw h.1~1s ot d.l,~ltlc.ltlOn. TABUlAE AND DISSEPIMENT$ (FIG. 5.1 OF-H)
r- ~ I111Ugh nurn,~trutturt..• h,,, prm•td meful here T.tbu l:le .lfl' transverse plate~ "' hrch 111.1} lw tl,1t,
d ng '' uh ntht·r durat tcrs, it' mtnprt•t.mon 1~ 'rill convex or roncave. They mually t..)l'cupy a tc:lltr.ll
!;· ~~ 'ubjtct nt m ut h dt\l'll\Sion 1ud lt 1s b} no me.lll'i ~p.Ke or t.lhul:tnum. and 1.f there 1\ an .1\llltornplcx
unt\·crul kl'\ . rhey JOin with 1t. Tabulae rn.1y bt• rt•pl.ll..~o.'J b~ .1
l- number of ~nuller pbre' c.lilt>d tabdl.1c. I .tbul.1e m
lll rhe C)athaxonircae, the most pnnurivt> \Upt•rtJnuly.
e. rhr Rugo'·' \t'pt.tl micro,tmt turt' .md lrrJnge- gener.lll) exrt·nd nghr across the cor.lllunl, but 111
1d '.11) ~rrc.ltl) ••md thmt..· ditli.·n.·nce' .ue of
tllt lllJ} mmt Rugm.t they rennjnare extcrnalh 1n J margtn-
111 gh l.IXOIHHlllt v.th.lt'. St•pt,J lrl' nnrnull) lamuur, anum. whrrh 1s erd1er J septal stcreozone or a dJs-
uf 11 pt•rfnr.ltld 'ept.1 do occur 111 \DiliL' gt..·m·r.l where sepimentam lm, and hence do not rt·ach rhe edge.
.1- u 1r.1lwwi.H.· .m· nut dmdy jmnl'd U\ll.llly they Dmeprmt..·nts. the sm.ill pbtes mu.tll] found
111 1tl.11~ht, 'lH11CtllllL'' ztgz,\g Commonly -.epta arc row.1nh tht· edge of the cora.llum, he p~..·npht•ral to
en •olllllltnmt thickm·•" thrnughnut tht'lr kngth. In the tabul.mum .md hkc the t:Jhul.re .lrt ( omtrultcd
he thplrylluw (Fi~ S.lllk). t(lr lmt.lllte. the outer of tlbro-nnnnal nssue. They are of \,tnom kmds.
ut th.: 'ept.t .1re \U b'TC.ltl) dm kt·m·d th.lt thq. They m.l) ht• \Imply mclincd or ,Jightly 1\\\·olkn
onugutlll,, tomung a \\C:II-niJrked 't~o.•reozone. pl.ttc-. dipping row:rrd\ the .L-...,al rq~1011, but m
lll- In Auhlpfl)'llrmr (Fig. 5. 111.1} onh- lulf of the cor.tl Plullip.\.II(TI'<I. nr.rmrwphyllrmr and n·l.mve\. ti.lr
llS mJI!y ha.' mt..•thally duck~ncd \ept.l . Th~o.· ,jJc, of nl\tJlKt', thne evolved an extraordmary r.lllgc of
th~ < \,lll 'q't..l .ITL' li\U,lll)' \11\IHlt!J but .liT oftl'll pro- di,\eplllll'ntal tvpe~. includm~ horse~hoe-~haped
~itc· 1,tl \\ rth fbnge~ 1.1r ~m.tll dt•ntH ul.uron~ wht..•rt• the one~. wh1ch h.tve proved useful m cl.J~stfit.ltiOn.
'tlw tr.1hl'tt1br t1p' ~how through. In Unc•II•ISimca Di~~epiments are not found in the t•arhc't rugo~l'
~ r, J lllfllt' other .1plu·oid gem:rJ th~o.· mn~o.•r p.lrt' uf gc:ncr.t. On their first :rppe.1rance m l.1tc Ordov1t 1;111
lm scpt.t .m: rL·plan·d by d"'epurlelll,, whibt strt..·ptc:h:-.nuunids and column.triids the} ,Ill' con-
lent wralhtc~ ut thJmru,tc:rorJ t't}'e' have confmed to .1 thin pt..·npheral dissep1ment.mum. In btcr
nt ~rpta. lmpl!'.\115 illmtr.tte' 1 t'. p~o.· t)f toral in genera. ho\H'\'L'r. they berome mort unport.lnt
h 'epta .1re unperti.·ctly dt·vclop~o.·d. unh bc:ing stmcntr.tl component\, most noun· thl) m type'
ClpC'fl) timm·d rmmcdratd\ .tho\l' l".ll'h tJbular 'urh J' l.11u~d,,h 1t1 Rarely. a' 111 \.ywplr)'llum. the.·
dl\SL'punt:nt;tnum together wnh t.rbdl.tt' With \\h11..h
disseplmt:Jlt\ nuy intergrade. nuke up virtu.lll) thl'
whok skdeton. \inn: th e septa .lrL' reduced to \ep.l-
.n 11\Jnv Rugo~.t haw .1 n:nual \lrlltturc (Fig. r.Hc: trabt..·tttl,le devdopcd only on thl' uppc.·r \Urt:1ce
I .1 t), 1durh m.1y be one of th l'L't' b."lt k rnds. An of tht· tabell,tc ,111d di,sepimcnts.
mrtt x 1\ tC.mncd by the JOilll'd .rxial c:nd.'> of the
or '"Pt~. 'lightly tWI~ted ,l\ Ul J>t yd"'Pil yllum. A CAUCE
mdla 1s ''mply a d1l.ltt:'d end of tht• counter- The calicc " th.lt p.1rt of the coral "hkh. m hti.·. ",1,
m and ~~ reprt·~ent.lll\ t' ol ut/hh/rtl/ltlll, in cont.ltt with tht..• ba~.ll ectodt·mt of tht• pnlyp lt "
:t.ltll<irllfl, L.lp/wplryUrdium .md othc:r ~imil.rr in c:fJi..rt a lllOU)d of tht' '\CCTetol'\ \llrt:ll'l'. Jt I\ not
m. An J:\:ul colunm. a' m !uh,plly/lum, i~ a LonunonJy \CL'll 111 'pecimem prt,trved in hnle,tonL''
a.xu zone fonned oC ta b ellae. i.c ~mall unk" thl·'e h,t\ e been Wl".tthued. but in mJIV1duah
nplrt{' ubnl.te wHhm th~..· .1\.1.11 t omplex. l u colleued from ,hJ le~ calicular surt:lrc.·, cm oltcn bt..•
l1/rr,1 'lcritl!'f)'•ltlws <llld Dibtu~t•plt)'llrtor the axial cle.trly st'l'll. u,trally there j, an annul.1r t.tiKul.tr pl.lt-
,unn appc.l r"' web-like rn 'c:ctron ,111d •~ Ukcwi~e fonn ~urround111g a deep central deprt'\\I0\1 (t.llicular
ll\1\\t'd l.lrgd] of t.tbul,lt' v. ith ~t..·pt.l l russing it; in pit}. So111e genera With ,111 .tx.J.II complex h.IVL :1
116 Cnidarians
(c) Actinocylfthu~
nwk'
(a) Auloph yllum [)U( tl
u qui
5 l!lt
Spou
~h tpe
sur
(FK
Whet
(f) Oibuno phyl/um
(g) Phllcttl lophyllu m prnt<
IIl.l) f
(h) Cystiph yllum
1 use:
() CJH
lHho
lhR·c
rt cv
p r 1\
tt 1 n
(k) Kodono phylfum Ax

(m) Clllceo l•
iis"
the
Ih
111((
.lXrJ
the
I h
\l7t'
llltJ
S,l
(q) Metrio phyllum
2 Pu-:
th~.:
(n) Lophophy/1/dium (rJ
p.m
!;l,l
L-.~rc
' O i l' I
n 11
gat
C(
he!
hcC"J
ed sepia I• 1
axial structure and semic1rculor z.one of thicken
Figure 5. 10 Morphology of Rugoso: (o) Aulophyllum (Corb.) with (Corb.) with lonsdoleoid dissepiments or presepimentsJ· I 2.5:
Ptychophyllum (Sil.) with axial vortex( >< 0.75); (c) Actinoc yothus
odendron (Corb.) , vertical section showing dissepimentorium on tolnu
Aulino (Corb.) with median oulos (x 1 6); (e) Siphon llrill
e and location of central tobulor ium and dissepimenton
(x 1.5); (f) Dibunophyllum (Carb.), showing axial structur horseshoe-shaped dissepiments (..: 4); (h) Cystiphyllum (S•I.) ne
Phocellophyllum (Dev ). vertical section of dissepimentorium with
vertical and transverse sections (x 1 5); (k) KodonoPhyllum (SO tho,
ing brood dissepimentorium (:. 0.75), (j) Stereolasma (Dev.), coral with operculum (.., 0.5); (n) Lophoehy/lidium (Corb
septal stereozone (x 1.5); (m) Co/ceo/o, a Devon ion solitary rugose . showing downturned outer edses of tabulae formm9 o ' pu
vertical sections; (p) Diphyp hyllum (Corb.)
two transverse and one e and epitheca; (r) d1verse colycolshapes in Spongophyl doi
(x 1.5), (q) Melriophyllum (Dev.), with very thickened axial structuron Invertebrate Paleontology, Part F.)
gramm atic). (Mainly redrawn from illustrations in Hill in Treatise
Class Anthozoa 11 7
m.trkcd calicular ho's where it l'nwrgcs to the \Urf.1cc. stH:h .1s S. 1/lclrfi11i with a wide di~~-.:piment.mum,
rut thii I\ not :JIW.lYS present. I he foml of the ea lice the daughter ,mw~ from well within thl· pMent
qu1tl' van.tble even ,,;thul tauuhcs. a.' ~hov.. n 111 Fig. cahce and inhcrib ir~ in.itiJI st..pt.l, e'pt·t..-l.tlh thmt'
'lthn, whu:h 1llmrratcs vanom tom1' found m the of the outer \">all, from the end-; of the p.Irt•nt
pimguphylhd.lc 111 wh1ch thnc .trt' conic.tl. bcU- ~epta. Cenoid ~peues of Srplt,lllcldt'lldflm mtrc he
pt•d, \Juccr--;h.lpcd and ewncd type\. 111 ,, manner Jn.1Jogous to the third typ..:, blH the
d;wghtcr rcmams .ttlached to the parent .111d ts
iiJ0()4NG AND THE PRODUCTION OF NEW CORAWTES polygonal from the start. Nudd' ( I'JX I) h,ts extt..'ll-
(PG. 56) '>IVcl) reviewed colony tonu 111 Srpltcllltldttllllllll .
. n n a .:ompuund coral ~ow~ thl· tif\t-formcd
protocorallite grvcs nsc N~xu.tlly to offict\, which Th(•re arc some ru~mt.: coral' (not,tbly AtlllhlC)'Il/lttls)
·.w t(mn m .1 number of w.1ys. The tenn budding whtch show a cunous feature a"ou.ttcd w1th otf,t·r
"''·d tor rhc ~oft pJ.m, increase tor the ~k.ch:ton. production. A p.m of narrow tubular ducts m.l) be
Deuiled \tudtc' h.lVe bn·n mad..:. 111 the ~een cxtend.ing ti-l'>lll the base ot the otF.ct obhqlll..'l)
o'trotlollld.lt. .md mJJn other groups, of the upw.lrd" through the d1s~epimcnranum of the p.tr-
~e mam method' of mcrc.m• :md buddmg. There cnt. The ducts m a} then merge .11 h.ighcr k vd' 111
' rwo ~ind\ llf incre<l~e. p.lrrtcidal. m whn.h the the parent coralhte. They can only have \erveJ
{Uhllt is dl.!\troyeJ (I, 2) ..lJld non-pamnll.ll, wbcrc to prolong ga'itnc and ncrvou' commuull.Hlnn
not. bc..·twec.:n the parent corJllite and tht.: d:tughtl'r umil
tht l.lttcr was well e~tabhsh..:d (Sc..ruttou. I CJl{\) .
Axllllnlrt'.l\l'. In tlm mndc rhe corallm: ~phb by Oltvcr ( 1976a) ha~ rev1ewed many ,l,pec~ of llllon'i
ll\\1011 llllll two or more d.1ughter polyps within forlll,Ition in Rugo~a .111d other corals.
the catin~. ~uch budding is incVJt:~bly parrictdal.
The upper ~urt.K..:s of many compound corals can REJUVENESCENCE (FIG. 5.7F)
ufrcn be ~l'l'n with 'ever.tl cor:tllttes undcrgomg In mhtary cylindm.1l funm thl' c..orallite '' ottcn
a.Wl mm: J\e .H approxmnrl!ly the same rune. As found to be comtnctcd at trrcgul.lr mtervals. kJvin~
'ht colnny cnntmuc~ to b'TO\.\ the p:~rcnt cor;tllite a bro.td o r narrow shelf where the \C(>tJtt.: olde1 ~,.·,,l
bunt'd .md the: lbnghtcr coralhte> grow to fuU ice is exposed. Above this it may exp.tnd J!,(;tlll,
uc. Tlm 1s the ka\t n>nunon of modes of bdore once morl bctng c:omrnctcd. 1 he~e c..un-
reJ.Ir 111 thl· Rugo,a, though nom1al 111 the .,tnctctl bands represent times of ~en·.,,. Thm tn peri-
umn Arcn•uf,ma (Fig. ::; "'l') od' of f.lmiuc tllt! poi}'P rcsorbcd its O\nl msut'' in
PmphcrJI lll(fl'.hC. Hc:rc mtall offSets ame round order to stay aJive .111d shrank ,1\\:lY from the edge,,
lu· parent (Orallite (Fig. =i.7g). This is Likewise becoming sm alkr while retaining 1ts fonn. Till" next
~.!ITiod tl; 1l ts quite cotnmon m ta~cicul.ttc gcn- penod of increas..:J luod mpply pennmed gro\\ th eo
n. mmdt .h L:'ntdtlp/ryllum bcgm once more; this i\ r cjuven esccocc
Lltml mcrt·a~e. Thts " rhc mo~t lOlllmon Starv:~tion and regrmHh, hO\\l'Vl'r, are onh one.
~od ot maclw in the Rugma and 1~ not par-
i7 cxplananon; pa.rual 'morhcring of rhc coral by sc..·dt-
hid. In Sip/Wt1(1c/clldrmr 11 h.ts been wdJ mvesti- ment is another altcrn.ttivc. ln some specie~ rt.:JUVl'-
f1 r ,d 0ull. IYCIS) but h.ts proved to be so variable ncKcncc may be ..:~pcci.II Iy strong. so that sh:~rp rum
n to be ol ~nmed ta:xonmni( value at the generic arc t'fm11ed with deep .md almo\t sht-likc cnntrac-
-d. Three dl,lltll't t~ pl'' of lateral iucre.t\e h.we tiom berwecn. In \Omc tmtanc:cs mull bud' .1rc
n dt~ttn~lll\hed ut ,...,rpllc'IIOdmdmn usmg ,l·rial found on the upper surface of.1 l'Jltlc. Tbes..: prob-
('(t!Oll\ (I 1g. 5.6c-L'). ln one type (e.g. S.jiiiiCt'lllll) ably result from J !arc n.juvenesc cnce of ,t uH.Jl
· b{cr.tl bud ames fu lly l xrernal to the parent whic h almost d1c:d, fi·om ~rnall ai'l'as of stJll-ltving
1); (b) t and IS umi<lUy a'cpt.He; scpta onJy fonn ti\\Ul' .
5).(dl ~ In .1 \t'cond type. t\ p1l .tl nf ~pene' '" 1th a
Jlan vfll
m J,,,eptmentarium (e.g. S .•mwdurmm), the Evolution in the Rugosa
m; lsl
show curJiht~ anscs from ne.1r rhc penphl'l)' .md, Tht' Rugos:J have relatively ii: \\ phylogcncuc.. .llly
J), W1\h lll•u~h 'eptatc from the st,u't, does not tnherit usdi.d ch:~ractcr~. Furthermore, they were Vl'f\ plJ\-
·Perm),
a tvbe
u !Torn the parent. lu ,1 thtrd type, 111 ~pct..·ics tic in thetr cvolution.uy potenti.ll .md notonomlv
a o (dta
1 18 Cnidorions
~ul~jel' t to ht'lllennwrphll' trc:mh m ~q.1arare stocks. dcdine llllo the Pcrrm.m. ThL· la\t rugmam Ill
Y..:t ho\\l'\'l'f c.:ummnn 'uch Jtc:rattve trend~ may gcncra stu.: h ,1, Pit ftlJih}'/111111 .md IJ~,,~I!IIh'J'"}'//um
llJvc b~l·n, tlw gu1ding pnnciplc: controlling thc1r rugmam .trc known .1th·r thL· end of the Pennun
cvuluuon \\ ·'' .th\ .n:, to\\ .Jrdo; .1 ~rrong and firm thl' whole tht: r.lttt-' of ,o!Jrary to mlcmial
skc:kton . 1 hu~ num of thl· ob,crvcd 'rrucwn.•, cm remain' relatively t'Oil~t.lnt, the o;olnarv fonn~
bc mtcrprt•tc:d fum tinn.Jll\': t'l)r imt.mce. the compli- domin.lllt. but chnt• JS .1 \\eak tn·nd toward\ a
c.ltiOn ,11 the tr.lhnubt• inCTl'Jo;cd ,kdetal 'trength, pcr.:em:~gc of .mmr.1l fonm.
the cmll.ll' hdd tht• polyp mort' tinnl} and the ,JXJal
column g.l\\.' grl'.Hl'r mppnrt m tht• central region. EVOlUTlONARY TRENDS
Tlwrl' '' rhu' 'l'llll' ltllll ttnn.tl 111e.ming in all. the Rugose cor.1l' h.1d onh .1 lin11tni cvolutlllllJI)
trend~ th.lt h.l\'t' bt'l'll {,b,cnTd. though it j, not u.tl, hence it w.l\ mhl·ruHly bkdy that the 'am
,,h,ay' ck.tr wh.lt tiK· ltllll t1un v.. a, lli \trucntrc v.rould bl· c volved st·vcr.ll utn~
ActOrdmg to l.1ng ( llJ2j), who fmt tried to C'i
OVERAll PATTERN OF EVOLUTION such repennve 'tn:nJs', 111 Carbun if~rou' t"orJI
fhe c·.nlit·,t 1-110\\ n nt all lllr.th arc the Lower clur:lcter t(.)llow' in v.Jrvmg dq.,rrees and ,\1 1
C.1mbn.1n 1'.1httl.trmud.1 and the Middle Cambrian rates one of .1 'ontp.1r.1tivdy feY\ poss1bll• Je\
CothontonHI.ll'. Undoubted Rugosa do not appear men to; pomtmg to Lhc l.'xistem:c ... of lmuteJ ~
unnl the M1ddk Ordov1cian (Blacknvcran) in dencieo; ... rL·pc.Jtcd 111 each lineage'. In l~xot
North Anwr11 .1. Thl'y '>prl'.td rapidl) ,md evoh-cd in whiCh bl'conw~ confii'L'd h) ~lllh trl'lld.,, the
1 ~l ne<; ot \IICl'l'\\IVl t'pt,otk~. Dunng each of these w;ly ro Sl'pJr:ltl' .tnd 1du1tlf'v tbL· g~:ncr.t ~~ b\
one partic u\.11 f.tun.ll t,'fnup \\ J~ dominant. ;llld sub- dcr.Hicd sn1dy nf IIIOI1lhology. fine 'trucrurc:
,jdi.lr)' d t' llll'llb tl the tlllll' wt..' re ( 1) <.mall persi,tent ontogt•ny.
br.1d~ tcht 'tm k~ (2) t'Jrh 111entber' of a later domi- In gcncrJl ten m the Jevdopment of a rnJ!l,"
n.Jnt \fll( J..: .mJ (.'\) rt•nm.ml\ of a lt1rmcrly dominant 111111 (<.tert'070JH.' or dl''l ptment.lnum) lS p~rlu
'rork. mmr pt'f\1\tt•nt :111d unive~al o( all trends
·rh,· r.Jrlir,t rugo t ' tnral' .tppear ju't .1fter d1e fmr Rugo~J. ,md o;inn· tlm h.1ppen~:d m several
T.lhul.1t 1 .llld thl' m cr.tll p.lttt'rm of evolution in the mdept•nckntly m the Upper Ordov1uan and L
two b'lnlll'' .m.· rl'lll.lrkJbl) sunil.tr (')rrurron. llJgX) )ilun.ln jll\1 pnor to the fir..t \llll"l'" of the R:l;
Jn dw M id-OrJm tn.m to Lo\\ er SiluriJn small a' rt·ef builder; 1t wa' dearly 1mport.mt 11 ~n
\OliLII'\' I !ll.d\ \\ l.'fl' dunun.lllt. lllJUUY \VlthOUt dJ~ incre.l'l' in l'ulony ,jzt• whtlc hnnly 'u
't•pillll.'llb nml '' tth o..llll) t!:eblt• d~:vdopmcnt of tra- tht: polyp .lt tht· 111.1r~>in A number of othtr
bt·nd.u ti"m·. ·r hert·.ttter tht•re was a stead) im:-n.•,Jsc fundamt·nt.ll to ,1U the Rugo<;a haw als
in lllllllbt•t, oi Ru~o'J up until the M1ddlc di,nnb"\ll<.hcd.
De\'oni.ln .1ml \\ tth lllll1tu,u corals, tasciculate cen- Thl' phylogL'IIY \If thl' Rugos.t, howc\'er
md ;Jnd .llmtr.d bt·l.'onun~ incre.tsmgly important. tOt) poorly 1-nown lll allmv trend~ to bt• p1
The rtt•t" lublt.ll "' ,1, tolomzed, though in mo~t reef documL'Illt:d c.xn·pt 111 .1 ti.-w ~pentic case'
h.1b1tJh the Rugos.l wert' ,ubord.inate ro the stro- Evolutmnary ~tltliJe, L1f trend~ wtthtn f
m.ltoporutd~ .111d t.tlnd.ltt· t nr.tk The succcs~ion i~ Litho~troltlHIJd,tl' h:~vt• shnwn that tiJ( ~w.tll
de;u fmnt h~. 5.11 .•md p.111Kul.:u orders come into ~0 clnst• Ill ~CVL'J.II dL'SCl'JH.ienl ~tocks dut
dOJuin.llll't' in turn. Tht'n 111 the Frasnian (L;HC' ongm.llly lu1uped 111 the s,tme ~ent:r.l Wt'rt' u
Oevoni.111) Lllllt' .111 .lllllll't tot.1l collapse of mgost• (\t-riwd pnlyphylctlcally. rhus 'Urio11t1SIIt1t'~
t'Or.1l LHtn.l\, p.11.tllded h ,111 t'lJU.tll) dr:unJtic drop h.l\ .1 Wt',ll- Ill ah,cnl columelb ,md 1 tend
in tht t 1hu Lttl.',. \Cptl to bl witbdJ.lwn lrom both the axu
R.t·ul\'t'l') hnm dw. lllJjlll' cxtmrt1on episode w.t, pt•ripht'l'\. i' c\ 1dlndy polvphylctJc .tnd h.u
r.tpHI Jnd tht• nwallml' l.'cmy,rcms were \O ttllickJy be ,pJtr stllt'l' 11 mdudL·d morph-; that art
rc,tot keJ 'o th.H hy rht• l.tr.·r l owl'r Carbonjferom funn bur dt•rin·d lro111 diHi.-rent Jnce,tral spr
tilt I~Ltgi''·J lud nc.uly .JchJe\-cd their LowtT Ap.trt trom the'~: tr~:nd~. therc " .1 anmd
Dt nmt.lll nuxunum. 13) tlu\ time the Lithmtrotio- 'ome Litllo,trottomtbc .md m .1dd1t1on a pun
md.Jo.:, Auluph\ lltd,Jt' .md otlwr \: la'>~IC' C.lfboniferom mtere,ting rrt•nJ wllldl 1m·olw' the lo~s of
ni!;OS.Ill' \\ c1 ~· d1 •nunant. but rh ere was then a ,}o\\ ment' throu~h time. In the e.Hiy tom
Class Anthozoa 119
\! lt~no.ll'lhlr,,, pr.ll'lllllltlm ) thc:rt• lrt• \I'\ Cln let' of tht cur.tl' lll'l'Ur onl\' ~t ccrt.1111 hon7on' \\1thm .1
0 punc:1t,, a·dut ing m tht· dc:,n:ntkn t <!. ,,,,,; h1ghly vari.thk sedimental '\ \CtJUencc, 11 WJ'\ nnt
11 ull!ur .md tlwn ro rwo. In .H.idltHJt J tlw ~cpta ~c: n c:r pm\Jblc to t·~t.lhlish cont tnuJty of vari.Jtion Second,
<; .1! ~ ~wnnt• t'l·\H'r and tht tabul.w more ""tdcly the \tudy \\,h done wrthin ,1 lumtc:d ,\Tl'l .tlone .•md
g d. The e\TI)unona r\ prmnple !:(0\'l.'mmg the tt 1~ nut dc.:.tr ho\\ t:u rhe >e4t1tntt can lw ,nbst.mn-
~I n1,111 nf th~.: nc:w t\ pes 1\ probabl) p.a:domorp h1c ,\tnl in other regiom. Also, ~llll'C: diffc:rt•nt \pcries'
m1n' Lh t• de~rt•nck·nt .1du lt, tend w rc:,c:mblc JIKC\- t'Ul''\iHed in tnnc (.Z. dcf,moun .llld Z. p.u.llll'ia) .111d
!al Jm t'lllb. "iut h mOJvhnlo!-.") '' uuiJ 'c.: cm ro be there i~ \OIIIt. evidence of the '<I me 'PC:I. ic:~ brin~
~<~ilh Jppmpn ne tor .111 t'n\ 1ronnu.:nt in '"Inch found Jt thtT~.rcnt honzons 111 Jlfl-ert'lll .lrt:a\, tlw
l- np,,t ~10\\th w1rh mlllim.d mewboht· expenditu re p.ltt~m of ~.·vol uti on w."
probJbly very complex .11HI
Jd 1 'kdt·ul tkmt•m, w.l\ dc:~1rabk l ikewi~t'. the 1' perhap' bc't regankd as the c'\prc:s,wn of .1 trt·nd
~uOJgt• oft'lmd ,Jurin~ bl.'t\\Cen the hnkt·d pnlyp' w1tlun a 'pc~.;ic' group rarhcr than J~ .1 m.Jttl'r oi
th~ 'onona,tr ll'tnd' .1phro1J murpholog~es, .md l111car dl.'o;u·m involvmg sepJr.lte speuc,. In t'imc-
d11' f.tn th.lt tn tht'\t' ,Ill tht• .IVJi J,Jbk \Urf:lce \p,lCl' is tronal tenm the mnrpholob 'Y nt X. c'I111Sirilla m.rlt,
v~r~tl by polyp,, would k:td to gn. uc:r dtielt'IICY ti.)t .1 ~trong1.1 axial rq~1on than rh.lt ot %. ddml(luri.
tht' lOIIUtun J\ .111 111tC~f.lrt'd Ulllt but that ot L. cli41111!141 " de.Jrl) weake1 Pt·rhap' tht·
bch of rhe't trend~ thcrdore. t·.xpres~c:d poly- need ~or r.1p1d growth, .1s "''lluld be allowt·d by ~cp
phi l~tJtaU\. h.l\ lt\ O\\ 11 ,ldvant.lgl.' ~uitahk !t)r t,ll reduttmn, may 111 thi' ca~c h.we othet rhc ax1.1l
.llt1(1JI.Ir rcolo!Pt'.ll rond1nom wt.1kness.
\o many f.Ktor'i \l'CJll ro h.1ve been uwolvc:d in
the evolution of the Z. delallclll('l ~roup mcludin~
1re rdati\ t..h k\\ w ell-docume mt·d ,\Ccount'i of trt ndmg. rc\pOn~t: to loc~l ~·m·lronmt"m.ll cond1-
- lie: evoluuon.tf) rh.mgc m rlw Rugu,,t One nons, p.1edomor pho'"• allop.1try and polypln+~
th~ hN known 1\ thilt nf C:~rrutht•f\ ( J t) 11)), \\ ho th,lt the ~:vol ution Jr~ pattern now ~CL'lllS to be 'll
ked nn the /.tp/mllli/n drl.llltlllfi ~rroup in \nnl.md complex ,l\ to eludl.' .ma)v,i~ until bt·ttl'f dat.1 an:
:~ p~ r.d Engbnd Tht 'Pt'tit:~ b'TOllp Jctullh r.ln~t" ,,, t:u J\':nhble
er !klb'llln H1' cullectmm wen· m:~dc fiom thin
tnfltolll'~ 111 .lll .tltt'nl.lllng lime,tnnt.~h:tk'-'I.Jild Classification of the Rugosa (Fig. 5. 11)
c-w.Jl \cquenn:, all \\ell lncJted ~tnngraphilaUy. ' I ht: follow1ng cl,t\\lhtatm n. atncd upon the vm-
1Ki\1ng the pupubtillll \ Ill ,L,ccndmg ~equenn he Olh ~tructural feature' alrcad\ de,cnbt•d h ba,t·d
I dw most ch.tr.lt'tt'f\ ('lupe, ~17t. t.tbular ,p.~t.utg, upon that 111 the "Jit·ati.1c: 011 llll'l'rlrbrarr g,/n>ll/11/t~~y.
t I tl•nded to rclll.tin 'oml.llll but th.ll there ~ecmed lt j, brgcly founded 011 mcga\t opit t\:.1tlll"t:\, knowl-
•llll't">~iw ch.mgc' 111 the ~lupt <lf the t.lrdmaJ t•dge of mJCrmtmc tun: ~~ n<lt ver :1dvann·d enough
.md rhe k n~rth ol tlw tn.~Jt)r sept.l (Fig. t<l ht• of much use. 'I he ine\ 1Lib le elfec t of parallel
olg·m). The oldest bc:d' tomam .Z dclamw11i ssp., trend~ ;lnd con vergen t evolution . howt'Vt'l', cn:.ltt''
lu<h h.t, 'cpt t lllll'OII~ 111 the Ct'ntrt: .md a lar~~ tar- t.1xonormc problems \\ luch 1f rc,nked 111.1) lead to .1
ml) fi\\,ub c:xp.1ndcJ .Lxulh . A v:tn.mr nf tlu~. /.pm- r.llhcal re.1rrangement of the group. Cl.t"Jtlc.ltlon oi
ar . wnh 1 p.lralld-,Jdt ·J fo,o;ub, on u~ m tht· ,,mic the Rugosa "in a sr.1tc: of some Aux; nt.lll)i authon-
t'IC~ J1 lt \\.1\ prob:~hl~ Jcnwd pacdon1nrphicaUy. lit•s (e.g. Scmtton. I l)l)7 a) no longer mL Cy,t1phylhd.1
ltlh A\lll\rt-hvcd \Ilk bram h lc:d ro rh~ \111.111 / /,w•- .md Staumd.1.
11\h l)b In youn~~.·r bed, 7 dclm1nlln .md /.. Jl•lltli-
t(,r ,A tfl' 1.1n· and replacl.'d l.lrgely by 7.. ronstllcttl in
ORDER RUGOSA (?C.tm./Urd .-U. Pt•rm.): Sulit..uy or tolo-
th~ llh th~ Jxi.1l end of tht· t Jrdin.ll fil~'ub I\ con- lt.tl cor.J, wnh the llUJM ~cpt:.l tll\t'rteJ 111 lour qu.tJrmt'
lfl tcd w .1 lcy hok ~hapc. In rl11. 'uunge\t hmc- ml\ rei.J.ttvc hl the pwto .ept<t. [.puheca nom1.11l) pre"'tH.
ml1r c; there otl'\11'\ /, dl.'fllllda; tlm ft)nll p.1,ses .mJ up to tlut'c order. nf wpta. DI"<'Jllmcnt' lll'tur 111 111<\f<'
nugh a ccJIISIMI41·type morpholo~y 111 it~ juwnilc ,\dV,111CCd groups; tabuf,Jt' 110m1ally (11C'iCIH. rnsl- Jl]d \1'1-
dIll ·dt1pmcnr. bur rhere.1ttt'r 'epta rl·tn.:.lt a\\ .1y trom ond-ordcr 'cpr.1 \\ell 1.kvdoped: thml-ordcr 'l pt.l lcs' '• •111-
lll<ln
ltrl~ tUITc
SUPERSUBORDER CYSTIPHYlliDA (M. \ )rd 'U. D<'\ J
.l pl Th,1ugh th1~ h.1~ been rc~arded ,1~ a d.1~~ic study H ~olitarv tlr somcttnw~ !:1s< it ul.11<
or ma."JVI' Srpta u~u.11ly
(q~ vcrd; lmutt·d 111 two rc,pects Om· ., that ~•nee 'PlOY Jnd weU-dcvdttp cd btll nt.t\ bt mu< h rcdur• d.
120 Cnidarians
I ahuiJt· .md d"'l f'llllC:Ilt'> v.m.1blv dt•\'dupc:d and of vesic- cement themsdve~ to the ,ub~rranun, though ~orm "Pl'll I
ular lonn, not dt•.•rh· J•,nnu from t\ICh mhcr. floor of have Clcamce~ of JttJ( hment or roor-likc 'talo1 \:lllcrg
tlhct· ionm 1 ,JulJm, bowL c g. l'.J/,Itwydus. Rhabd<>rydus, which may have helped to fLx them 1 hey seem 1 C01
l'rypl.J.<III.I, //,>lml'tlllyllum. Ct>~thiJ'hrlllllll, c.IICC<IIa, Cysti-
have preferred soft subsrratc:~. but smce relanvdl 'l'h'<.'\
pll)'llllm, \ltS•'I'hyllum,
few 'hard~'Tound~' are found pre\erved 1r 1~ nOl o( hare
SUPERSUBORDER STAURIIDA (M. OrJ -U. Pc:nn.): Solirary
or colmn•l "Hh l.immJI '' plJ u'ualh .:umpo>ed of trabec- known how far they could coloni7e ,uch hard sub- ofl<.'r •
uiJe I ht wptJ ~re muJII\' \\ c:U devdoped bm manv strares. The horn-hke ~hape '"ould gtvc.: a reasonJb \\Ould
retH'Jl hum thl" penpher.. A IIIJrgtn.JI \tc:reozone may degree of support 1f tht• l or.1l wert to be half -.unk lliiCl'll
Jevdup. 111 whtth '"' dt"cpmtcnb mJV be absenr. mud, com•c-.;: sldt• do\\'n. Tlm would allow tbt ,llld CO
T 1bul.•e mu•lh prcwm, though llt.l)< be m complete. polyp to be exposed ,1bove tht: ,e,1 floor and wou been tl
Then· .ut· H k•.t,t If• 'ubnrtlt•r> (f1g. S.ll). common gen- ·
enable 1t to my 111 the s.1me gener.1l ,ltmude \\h lit \\hich.
er.1 lilt ludt• 'il,lllllol, ,'\111 J<f<i<hlll./, Calt>styil.<, Cy.uha..\'1'111<1,
the coral grew. ' I here IS some ev1dence th:tt RU!,'\. , hr:r ,,
F.nt<f,>pllyllulll, 1r.uhth>t'hyllum. 4n·n•u/.m,,, Phillipsasrraea,
such as A~tlophy/111111 could grow like rh1s, orient~tt r~rdy I
H. lh'J'h)·llum, %.1['/llmlit•·•. ( '""";,, lult>phy/lum, P"l"e,Jsmil'"·
l il"•"''"'"'"· -;,/,,,,,,,,.~,.,,.ft,,/1, l..tlll<tllllfl•l. wHh respect ro current~. Commonly, howc\'er. 1 .llld su
:tny population some or all oftht: corall,\ art: tWI'tcd: m the
they must h.wt: toppled over .llld then grown up Pabco:
Ecology again. It IS
Solit.Hy ltu ~m,1 h.KI 1w really t:H"t!ctive means of The sohrary Cc1lr1'nla (Fig. 5. IOm) l.ty on th flO\WSS
.m~.:hora~t: 011 thl· \l',l floor !'hey d1d not nom1ally seo~ Aoor, convex side down with m operculum 1hr:ir ti
pholog
.111d j.v..
did llO
.lrl' hkt
IOgenera IOgenera Rug
10Jlll1el
C'llld l'(
t'\:nlog.
g~es h.
.,
.. .,.. .....
c:
.;: .,
.., ~"'
,.., .. .,c: ., ..~ :::.. "'.. .. §.. ..
.% !; .,.. ..., ~ ~
t:
..,.., ..~ .. :::::... ~ .. :t . g .

c:
..
I,.
c: c:
c: € c: ~
l1i :::: :.. ~
"i. ::: ....t: ~ ....
.. ~..
<: ::... ~ :.. .!!> e ~
l
!! <:
~ "' ~ ~
ID
c; <: c:
.. '---v--1
~
¥"' l
~ ~ Cl. ~
~ ... Q. "'
{"' .. E
.,~ ~
Cl. "0
Q.
" sCl. 0
.,c:
0 Cl. !;;
"'
...,.., ...8 ~ii"~12
.... 'ii ~ ~ c:
"' ~ Flt\IOSitrna ~
~"
SI ~
~ ::... ~ ~ ~
a;;; u cii ~ ~ tt u 8 it 1.1)
"
<( u ..:;
.._____...., ~
<( 1.1) - ~
~ ~ Q
-.;
>-
u Stau11 da -' Auloponda ~ "'
X X ~ FaYOSitlda F1gure 5
mentol~
R u G 0 s A T A B u L A T A (dl Do q
Polo~
Figure 5 11 Time ranges of supersuborders (and some suborders and fomilies) of Rugoso ond Tobuloto (Redrawn from Scrulloo, 198e! bent lifest
Class Anthozoa 121
p(n to lJ0°. Prc,unuhh it lotdd ~hLtt it in ca~c of PaL1eozoic Rugm.a th.lt lavl.'d un J \Oft \Cl floor
tlll~~l!llt) (Neuman . llJHH; F1g. =>.12).
Compound R.u~th,t drd not normJJly tl.x them- Mmt wcrl.' hbcroo,c"1k. 1.<:. lllllt.tlly attJchcd for .1
\'~ to the \l'<l flour l·itl1cr, thou~h rJrl' example<. short tinll m .1 'eduncm gra111 .md thcrLali.cr lying
lurdground~ ctKnt,tcd by Rugm.t Jn.' known. In rccumbem (e.g. l h•lt•phrt1f!111,1). Thl\ j, n 1dcnt from
'liJit(r\ub,tr.m."\ the \\l'tght ol the tolon\ in the mud the ~hapc of the c01.1l ~11d it' .ltt.tdunem \Car.
would no doubt ~vc ~Oillt' ,t.Jbrhty. ::,mte they \V ere Ambttoptl lonns sut h .1s Ph,utf,uus were JttJched t"l>r
lillcemented the~ w~..·rc l"ll\'lrnnment.lll) rc~trictcd a longer penod, to .1 'hell, for example. bdorc
mJ could nor butld proper reel\. llm ~eems to h.we falling m·er. RltL"I!IIIt1J1Iryllum 'ecn1:. to h.1ve gruwn
bttu thetr mo~r ,e,·ea· l'voluuolury limitation. ~rom uplij;ht, 1ts base cu1beddcd in ~olt 't'dunent.
~:~luch. m 'Pite of lhcir 230 million year history, RhiZmemlc ~em:ra have ruut-likc holdta't' emanJt-
the) \\t:re nevt'r .1ble to t'\l,tpe fhu~ they are very ing from the corallitc at st'Vt'rJl kvl·b. ,1nd fastenl'd
urel) found \\ llhiu tht> Iunit\ of ~trong WJve action to sediment gT<llm (e.g. Dotwphyl/11111). Thl'sl' form~
111d 1u wen: lll'Vl'f rl'.1lly itl\'olvt'd '" fi.tmt" builder~ grew upnght. Tlte ~mall di:.lotd,\1 PtJfrTCO(ydm,
tr d1c algal and cot.11....,trom.1toporoid red~ of the which inridcnrally forms a glob,llly wide5pread
l11lwnoit (Well~. I 1JS7). marker ho1izon in the Upper ll.llldtwerv. m.1y have
1t •~ not known for Cl'rtJIIl whether mgose corals been vagile, able to move <\round by its peripher.1l
f01Se,scd CndmymblOtiC ,1lg.H~ (700X.ll1thellae) in tentacles on the sediment ~urf:Ke. Fin.1Uy. several
dvir tissue' or not. Judging by the low level of mor- species of Cmllillkit~ .md other genera had a fLxmes-
phologic~! uncwation 111 nL~ost• lor.allite~ (Coates siJe mode of life, firmly .utach<:d through life to
djack~ou, llJH7), howewr, 1t j, pmb.1ble tlut they p;Hches ofh.ml botrom .md to mht'r tos~ils.
did not, smcl' modan t:or.1l~ l.tcktng zoox,mthellae In large, strongl> t urved rugmam 'lllh as
r: hke\\ise poorly inrq~r.1tcd. A 11/r•pfty//um, the hom-ltkl· foml w 1<. ,tL·IJpred for
Rugose coni bun.1s "l'rt· \ l'l) senqtJvt to cnvl- resting 111 the mud, conwx downw.1rd<. .md partiJIIy
ronmemal condtttol1\, .tnd the t'Clnn nf both sohrary embedded. If it toppkd over 11 regrew upw.1rds
md colonul ~pene' ".1, 'tmnglv mAuenced b) .1gain. which Jet:Ollnt\ to1 the convoluted shapes
rcologt1.'.ll fartor; (Wdk 193'"') V.mnm life st:rare- often ~t"cn in \UCh l orJI'
i:X"i h.l\'e been dl,£11l!."lll,hed 111 \oht.lf\' I ower <:iep.lrate f:lcic' f.wn;l\ c.m be dt\tmgtmhcd,
~~\~Nvu:~~,
(~:r;i'W.~~:·N,.', r
~'----~.&:;T.
-li -
(b) Laccophyllum 1 (d) Dokophylltlm

(f) Rhabdocyc/us
(g) Ho/ophrsgma
(c) Phaulactts
(a) Rhegmaphyllum
(e) Pa/eocyclus
~ 5 12 Life strategres of some lower Poloeozoic solitary Rugoso: (o) Rheamophyllvm growmg on on onginally unstable sedi-
euy gram; (b) Loccophy/lvm, aHoched to a vertical shell fragment: (c) PhavTOcfls, small specimen adopting a recumbent life style;
~ Doloplryllvm, a rhizosess1le form with tentacle-like appendages producing tubular holdfosts oHoching to sand groins, (e)
~Ius adult form possibly copoble of ~ile movement by lateral tentacles, (f) Rhobc:locyclus, odvlt turbinate form with recum-
86) ~ lilestyle; (g) Holophragmo , recumbent lifestyle (All redrawn from Neuman, 1988.)
122 Cnid a rians
a<LptcJ for p.micular c.:irnumwnccs and nonn;llly the earhesr of .111 rugose coral f.1una~ '>eem to hN h
found with dur.Ktemllc .t~~ouattom ofbrachiopoc:b been of tlm type. though occurnng m mudstom lJ
and tabul.tte,, Three such t:lnt' faunas were noted It i<. mu.tlly found wtth t:lbulatcs, small brr 3 (
by Hill (I 1U7) \\'orl..in~ main I} , ..;rh Scomsh ch10pods .md tnlobttes. As repn:sentcd rypicill <:(
Carbonitcrom \llll't'''iom. The'e are ao; follow.;. in the C.trbo111ferom the sea floor wa' gcncr.ill se
muddy. wnh detaymg ort-,r:~nic matter, but mUll ne
I. Cy.u/,,,,,,,,,,_l.tn:ophyllid fauna. Usually found have been f:urly well oxygcnated.
111 bl.tck or grt't'lmh c.tkareous ~halcs, though 2. C.mima-chstophyllid f.mna . Thi~ fauna, tound
~ometime' 111 brvo;o.ln ret>f. .tbo. these corals arc fil'lt 111 the Tourna1si<lll but prohabl) older, t"'Oil-
all ~m.tll .tml 'olitary w1th dmepiments poorly ~i-.r~ of large 'ohtary genera with dis~epime1
Jevdoped Thi' tam.·s t:mna ts very long ranged Clisiophylhth gradu.tlly tuok O\er from c.mmi
and n:rt.unly .mted.ltt'S the Carboniferous; indeed with time. These t:tunas were ad1pted to shJiill
Back reef
Redwater Reef. western Canada (Kiovan 1964)
Amph1pora
Stachyodes
4----.. .
........ strom-algal
- consort1um
~ rugose corals
tabular corals
mass1ve corals
Central European Reefs (Krebs 1974)
Amph1pora type
stroms
Stachyodes type
stroms
rugose corals
tabulate corals
tabular stroms
(e)
mass1ve stroms
F1g 5 13 cont. on p. 123
Figure 5. 13 (aHd) Distribution of NgO$E! corals, stromatoporoids and other reef-building organisms in various Devonion reJ;
Redwoter reef, Western Canada; (b) Central European reefs; (c), (d) Frasnian (c) and Famenman (d) reefs in the Conmng Sa::
Western Australia; (e) polaeoecolog1cal succession in a Devonian reef from pioneer to climax stages [(a) Redrawn from 1(1
1964, (b) redrawn from Krebs, 1974 · (c), (d) redrawn from Playford, 1980; (e) redrawn from Copper, 1974.]
Class Anthozaa 123
limNonc \1.'3\, \\'dl hr .tnd O'-)l?;Ctutcd but with ered to be ret·f faunas .m.llogous to modem reef
httlc tcrngcnnus lll.Ht'n,ll. corals, but smce the R.ugo'>a wt·re not really reef
3 Colonial or 'red~ cor.tl t:nm.l\. Compound rugose fom1ers they arc best comtdcrcd a~ JUSt J group of
cor1h "ith di"t'plllH.'Ilt' m.ty occur in ~m.11l and colonial corals. ·1 hcsc colomal corals evolved
CJttt•rcd 'h10henm': they .trt· reprt·sentcd m bed- from the Cmtmit~-chsJOphvlhd fauna.
ded limcstonc' ·'' well Tht'\l' \H.'rt• first cotmd-
- fish, crustaceans, rad1otanans, tentaculillds

nauttloids
-: - ammono1ds
: - conodonts
cnno1ds
<; sponges : :
braclllopods
V gastropods
~~b~iv~a~lv~e~s~:~~~~~~~~~~~~~
corals •
I Amphipora:
Stachyodes
bulbous stromatoporoids:
massive stromatoporoids ~-
stromatoporoi~s~:~~~E~;;~~~~~~~~
flat algal mat :
tabuldr lam1nar
I oncolites
columnar stromatolites
-
Renafcls-- ..;..11111.-..;...- - , - - - - - - -
Sphaerocodwm _ • ~ reef margm
1 reef~ l slo e: · /."). reef flat platform
margmal : )
bas1n t slope -· •.
(c)
: .. .. fish
.,..~
. .. - - nau1tlo1ds
-:--:--- -ammono1ds
- +-=.
- conodonts
cnno1ds
sponges
brachiopods
..
.-.
gastropods
I
corals
b1valves
. .. --
stromatoporo1ds -
flat algal niat
oncolites
. -
non-fenestra! columnar stromatoht es
fenestra! columnar stromatolites
Renalcis • •
--
Sphaerocod1um- ~ . ..;i:-feef margin
I · 1 : / . reef flat
basm
.f s"'''~
marg1 nal
Iope
j_
•
--. __
... --
:::?
-z~~
~
•''""m
- - .... ___
-::;:._
~__,.._,_...
(d)
STAGNANT WATER QUIET WATER HIGH ENERGY
123
!!efs. (o)
~ Sostn, mtermed1ate
Klovon,
124 Cnidarians
Htll\ thrt·c t:tttc' ,1\\0tt.lttom ban· ~cncrally been with rcspcu to ,ancient reef, h,l\ bt·u> gmcr.lll)
well \ttb\t.ullt.llnl, tlwugh tht· Cy.lll"r.\lmia fauna is firmed (e.g. J.lllllt.'\011, JCJ71 ), a mme detail<'<.!
\et> mud1 \Cp.n.ttt' '" hc.:rc.t' t!Jt.' other two arc: rurc of the '.uiou~ ror.Jl--.trom.JtoporOJd t
\omctitm'' nllt"rgLading .md et ologicall)' less wdl J\sociariom " emergmg. <)crutton (I 977). tor c
dctmed . Thu' at '' not utll'ununon to \C.:C rapid altcr- pie, who workt•d upon au evolving rcc:f compl
attom uf f.aun.l\ (2) .ami (3) in 'c:rtJt\U ~t·qucnce, as in \Outhwcsrcm EngLmd. hJ' 'lmwn ho\\ the
the C.trbuntkwu' of the Wl' 't ot lrd,md where. 111 m stromatopor01d rt•~:f ltme\tonc' dtfft:r from
,lddntun. tlud.. t·r, nf IN.iua!Jtc litho~trorionicb an~ in b1ohcnnal, btodJ,ttc .111d bt'dded do~rk lim~
found \\ tda rhc: l.irgt.· \\Jhtarv .1ulnphyllum growing Coppet ( 1974, r 1g. 5. I I le) h.1~ de,wbed .t g
w ithut the £1Hr kt·t, . ahzed pal:~eoecologtt.al '>Uue\~1011 of .1 Devo
More ret.ent \\ork h.l\ cumt·ntraH:d upon the das- · reef from p1once1 through intcrmedi.ttt' to J n
rnbunon of ru~O\L' .md otht:r torah tn PaJaeozotc reef sysren1. Dt•vonl<Jll rt·cf 'Y'tems were endc
reeE Klov.an (IWd) .and Krl.'b' (1974) havt> charted more cxrcns1vc and elJborate th.m thme of
(d)
thl.' lanc\ ,1\\0tl.tttum 111 ror,tb .md other reef organ- Stluri,lll. but by thl· end ut the DevomJn ho~d
i~rns Ill Dcvont.lll l l'd\ nr we,tern Canada and cen- l.lpsed. A fuller lrc,ltnlelll or the theme hi\ be
tral bmnp~: rt·spt'tliVd) (hg. '1.13). given by W.dkcr and Alben.L,Idt (1975)
In the~e re eh wlnni tl Rugns.l tend to be confined On a globa I scale. evident e ~~ .1bo accumuiJ
to the qmctct tnre1 el'f I.1ue~ around the flanks of rigid on tht' biogeogr,1plucal di:.tribution of mgose ut
trJmc n·e~. the\ do IWI no11Hally occur within the (Oliver, I976b).
ret:f tbclf nor, t.·xupt in localized patches. witbm the
bad,-rl'cl fltll'\ \o!Jt,ll'\' RugL"·• 1re likt>\vjSt' con- Order Tabulata (Fig. 5. 14)
tined but ,Jrt: mtmt:m .dh im1gmficant. Probably one The tabulate coral, .lrt• ent1rdy Palaeozo1c
of tlw hC\t t'\.HHpJ~:, tn ~hrm .1 dunging reef-rel.lted though they appear a ltttle earher than the R.u
cor.ll ecoi0)-."1 1~ 111 tht· I kvom.m Canning l3a~m m tht'y have an orhcn\'l~t' '11111lar tlme range. Th~
Wcqcm Au~tr.1h.1 . I krt· there 1\ .m exhumed Middle alw.1y~ colom.11. ncver ,ohtary. ,md muallv
to Upper I k\'om.m b.1rne1 rt>ef belt ~ome J51l km coralhtcs are mull. lnvan.1bl} they haYe promm
long (1'1.1}1nrd. I 'J~II); tht' ret'f romplexe!> fonned tabulae. but other 'kclet.al dement\, in parricul.tr
lmlc,tOill' pl.nfimm ,t,mdmg rem to hundreds of septa, are red ut. cd or ,1b~em
metre' ahcwe tht: l',J floor .md were flanked by mar- H .wmg n:lauvdy fe\\ \tntcrural elements.
gm.ll ,Jopl.' .md h.l\111 lkpmll\. In the f-r.lsnian. mgo'e T abul.lta are of compararivdy \lmple romtru.
cor.l". togt'rlll·r "ith \tf<llll.ltopnroid~ .md .1lgae .1rc FrWMIII'\ (SII.-Dc\.; rig. 5.14.1), fi1r example.
ch.Jr.Ktt.•mtll of the rn·f m.1r~n. reef A.u and plat- elongated, chm-wallcd, pnsmattc. polygonal r
timn. then number.. dtmtm,hing shorewards .md ,1lso lites with hon70ntal t.lbul,ae exrendmg right ;r.
do\\11 the rt.•t•l ,tnpl' Thl· rnn), \Vt:re abundant at the coraJium (Stcl, I97lJ). The :.epta arc redutrd
depth~ of down to I() m < .llld r,lre at greater depths. short and ~omewh;1t 1rregul.tr 'pmes. All th~ \\
Followmg tht.• b'1T.ll glnh.d l'Xtinctiom ,lt the end of are pcrfor.1tcd by numerous mural pores. ronnnn
the Fr.JSm:ln .md the tcmpor.Hy drowning of the the cora llitc~. In FaPosilt'S, ,1\ in t,lbubte' gcncr
n:d-;., tlll· char.lt'l~:r of thl' reef tomplc.:x changed dra- clement~ of \ l..dctal micrmtrucrure' (t1buiJe
m.lhcally .•111d rill' fc\v cor;!I\ that per-;i~ted were con- wal\<;) ::trc fibro-nomul. Tr:a bcculae are scattrrrd
fmeu to the reef.\) .1ml 111.\rginal ~lope, the reef margin t::tbulates. l n I;atJosilt>s they are nonna) to the 'urt
and tlw n·ef A.H. of the tabul.1e bm inclined upward~ and outw
Tabui.Hl' u>r.tk npt.·c i.11ly thme w1th a creeping along the a xi' of tht.• ~ept,l. Jn other gencrJ
or t(,Ji.H nHI\ l1Jb1t. wac \OI11C\\ hat le~s restricted spines an.· nor ne( c~sanly tr.tbt'culatc. Some gtr
but t.nuld not t umpctt.' '' ith \trom.uoporoids and (e.g. Tmlwrrditl'~) h.IVe trJheculate w:lll\.
algae Ill tht• wrbult·nt tolll' of mmt active reef Halysite.• ha'> tr.lhl'l lll.lt' 111 the walk F,gvro5 I•
growth . The lunl ,kt.·ktom ufboth rugn~c and ub- mural pot
ulJtl' n1r.1h, ho\\T\' ~o:r. \l'l.'lll to h,we contributed to Range in form and structure ~tons sht
c,tab\i,hm~; .1 farm lmmd.1ttnn tor btcr reef growth FORM Of CORA\lUM ( 0 75); I•
of .1lg.1t: .md \tronurnpon"lk Although tht' broJd Tht: Loral\um (culoma\ skeleton) t\ btult up by
I 3 SI. (fl
ttJlony clco
panem of di,tnhutlon of mgow and tabulate coral' vjdual polyps whKh may or may not be lm.t~rtcbrot~
Class Anthozoa 1 25
(a) Favosites (b ) Pleurodictyum
(d) Ha/y.ites
(e) Syringopora
connecting tubules
tnfundtbut.forrn labulaP.
ll Hl/io/itn (g )
<l'lncnchymc
Jl-
. >"''
ro
!llh
ng
)1).
1nd
I U1
:,~~
lrtb
pial
lClll
and
514 Morphology of Tobuloto (a) Favosile5 (Sil.-Dev.), vertical and transverse sections showing tabulae, reduced spines and
!3pom (• 4); (b) PlevrocJ,ctyum (Dev , internal mould) with commensol worm (x 1.5), (c) Colioporo (Dev). vertical and transverse
~ show,ng shelf-like squomuloe (" 4); (d) Ho/ysiles (Ord.-Sil). on external view of o colony showmg coten1form growth
SI; !e) Syrif190f>0!0 (Sil.-(orb.), vertical and transverse sections showing infundibuliform tabulae and connechng tubules
::.5I 1n Helloli~& (Sii-Dev ), transverse and vertical sections with corollites embedded in coenenchyme, (g) Holysifes, o Silurion
cleared of matrix and showing colentform growth (x 1). [(oHe) Redrawn from illustrations by Hill and Stumm in Treofise on
Poleonlology, Port F; In redrawn from on illustration in Woods, 1896; (g) reproduced by courtesy of C. Chaplin)
126 Cnidorions
ronm·ctl·d to l'.ICh othu Ccrioid form.\ (e.g. rh<: cocncnchymc-. Thl\ L.lll lw comtnKtl'd in van 11\ 1}\f
F.lVmltliM, h~. 5.14.t) l1<1w pulvgonal corallites all in way' and l'VIdcntly L'\'olvcd tndcpcndcntly m ~t'\ tlc.')
coiH,Itl Catcn ifonn rolnnil'' (q;. the Halysirina: \tOI.k,, tabul.lc, tnbLI.liiJl' borne on di,\cpim tiH I
Fig. 5 1-td.v h.J\ l' dongatcd LOralhtc' joined l..'nd to .111d £rahcc.:ul.lt' dmch p.lL ked or ~ometuncs o lrnm
end 111 \\,tlllkring p.•li'alk' Fasciculate rabuLm:<> mznl ullo tube-' 111.1~ .111 hJ\'l' J part in it. The pr Ordo·
(c.~ S}·nu.:•'/W.I) (fig. 5.1-k) have c\'lindrical coral- dulfion of 1 c: Ol'lll'lll hvme 1' um: of the 1 llldS I\
lltl'' \\ llll h 111.1) lw lk·ndnnd m phan:l01d and m.1y be unporrJnr tvolurwn.ll)' trt·mh in the Tab tnlrtl
pwviJt.•J \\tdtt.OIIIIn tmg tubule,. Auloporoid ~t.'n .motht:r bl.'ing the de\ t'lopment of nturJI l lfj
er.• h,l\'1.' .1 hr.mdtlll~ (ramose) tubular stn1cture and or ronnet'Lin~ tubule:' f'l)r lnter(onnert•on ot l.t~JIII,
l)frt:n .111 Clllnl,tlllg crccpmg (rcptant) habit. They t.'Orallitc\. Lhougl

u'uall~ em n"r lurd 'ub,nate~ but art' ~ometimes free they I
livm~ One ol thl· l.llt<:r Ault1por,, ~pcot.-s ha~ been BUDDING AND GROWTH OF THE CORALLITES t.thul.lt
r<:lUil\truttnl. ll\lllg ~ cmnptltt"f progr.un for st:rial Tabula£c lOr.d LtlloJUt:~ h,t\e n,..,o cnd-m~m l.t , 1< 11
\t.'lllOII' (Suutton, l'!~N) finally , coenenchymal adapt1vc Kl'OWtll 'tr.1Lt'g1e~ ("-,L nltton, l 1llJ7,t).
(hJ:?; 5. 1-H) rvpl',, duratt<:mtil nf the more advanced periphcr.1l growth, thl' ne-w tor.lllitcs an~e J!
t:tbui.Hc<; (e.g. dw I khohtina), have no dividing walls ed~t· of th<: t oh)ny; 111 medl.1l growLh thev emer
bctwec11 rhc lOr.tlhtl'~ but mste.1d .1 common mass of between tor,tllitcs which lormed earlier. S0111L
tompkx n'~ue, till· cornelll"h\111e, deposited by a ulates could on ly grow in nn1.· of these two IIJI
common lOionl.tl t1"ue (~oeuosarc) in berwc:en the: other~ lould lh.lll~e lheit !!rowth str.HI.')o.'lC\ l~ th
polyp' .md t(mnin~ a dt:tl\C~ cJk,lrt:ous mJSs in wluch dcvt:lopt:d
till cnr.llhrl'' .1n· t·mhcdded A, 111 the Rugosa,
colony tonn, wh~rh H.·fi.·f' ro the relatiom of the Evolution ond ecology
cor:tlhtt:,, IIHN ht• dJstm!,;llt'ht·d from colony h.1b1t; T.tbul.ltr-hh l'Or.tls aa· n·curded trom
rim~ '/7MIIIII<lJI<llrl J\ .1 r.1111mr bur cenmd favm1tid ( ',tmhri.m, hut 1hl· rt''l'lllhlancl'~ arc ~upcrtim1.1
Other rabul.ut·~ 11\.l\ he massive, foliaceous (m e.uhe~t undouhtt:d T.thul.lt.l .1rl' known 1n
whu:h luhu rht lor.ll tonm thm overbppmg lanunar C.madi.m (l ower OrdtWIC1.111) of North Am
'hct.'h) or c reeping ~omcwh.n .1nrcd.Hillg tlw tmval of the tir:r Ru 1\ t.:
.md hy thl M1ddlc t )rdonnan rhe fi1ll r.lngl.' of I s th;u

SKELETAl ElEMENTS ul.•rr colomal f) p<:' h.td .1men . Thr1r nver.•ll p pcnphl·
T.1bul,tt'. thl· Jllll\t unportant 'kdetal demcmo;, arc nf e\·oluunn 1s rem .• rk.tbly \ltnlbr tn th.1t of form. A
.lh\ .1y' prc\t nt .md tllllllllnnly tr.\VCf\C the coralhtc Rugo'><1 (l:,cnmon, I'JHH: we .tl~o f-1g. 5.11 ). T.tbtili:. hal}~llt
horiwnt.tll) Somctiml'' thl·y ,1rc replaced by were .tctually more muncmus than Rugo\,1 pnt!r ncdtJI 1
'n•.:~ller t.1bdl.1t' In the Cl,Ucubte genu\ Syringop,,m thl' St1un.u1, bur thl'}' w~: rl' more .~rutd} :1tlet1cd w dCim.
(F1g. 5.1-k) tht.· 1.1bul.ll· .m· funncJ-,h,lpcd (infundi- rhc- L ltl' On.lovll'l.lll cxtinctlllll L'Vcnt tlun \\trr h< \\C\C
buliform) .md run through thl· tubule~ th:tt connect latter grnup. f ht'fl'Jftl'r they rlunbed to,, Ill'\\ 1 (Ofullll'S
the- br.lndlt'S ~q1t.1 111 t.lbul.ttc) .trl rarely more than of UIVeNty in thl· M1ddk Devom.m. hut "1(1 (,(
shun spllll'\, oftl'll 12 111 number, but iu some c.1sc~ undcrwcn t ,1 l.lt.!~tmph 1r lkdml', contL'mpot b 11~1: I ,
rhq do rc.1d1 tltl' ct·ntrt• of the corallite. In Caliapom l'Ou,ly with thl• l ~u~m.l, 111 tl1l' Fr.lM11an Tl ll<)f J 11
(rig. 5 I 4c) .111d othl·r gcnaa of the Favositina they never rerovcrl•d ti dl y ti·mn llm. though they c01u I hl'rC
nuy bl' n:pl.ill'd b~ ~mall 'helves (squamulac) m·d until till' l'lld of thl' Pl·nnian whl'n they fie
'' h1t h JUt 111\hlnb .1t mtcrv.1h from the corallitc became t'Xlll1d. lt •~ con\ltkrcd hl-..dy rh.1t tlwu
" ,1ll. '-,cptalmst:rtlllll 111 ,1 SC4Ul'lll'l' mnilar to tlut of ltl' U \Ul't.l'S\ m tht· l ltL'r P.1lacnzmc nuy h,,,,.
dtc R.u~m.1 •~ 'l'lll Ill Jl,d,,c,tci~ ,md a ti.:w orhcr wbu- p.1rt1ally dui..' 10 the late Dt·vom 111 l'Xtincnon o
l.Ht' \ bur ih \I gill ht .\lltl' 1\ U1Kl'rt,lll1. \trom.lrupomllh, \\ hll h I1Jd 'rcarl·d the t
Whrrt' 1hl'rt' " .1 nl.lr!--"'nal zone. a~ in the more .md other mdl<:\ 111 \\ ludt t.1bui.Jtes had
Ad\ .tnl ed t.lbttiJtl..',, tt c Jn be of two kmds. It ma} be flourhlwd.
onlv .1 rhil h·m:d zo11c- of annul.tr lamellae a~ in Throu~h ttnll' ditli:rcnt
~,,,.,,,,,,,,,,,, cnmtncung the Jp<:rturc. ln the other 'tnll'turc' Jumin.ltl·d in rum
1-..md the !--'TO\\th of the 111,1rpnanum is Jccompanird \\ere comnHHll'\l up ro tht: c-Jrly <;i]urian (n
b\ the lo" lit' tlw l t>I,llhtc walh.. re~ulnnp; in J l ocnl'lll'hynul imperii 1r.Jtc 'lnKt url ). folio"
Class Anthozoa 1 27
tiVCl\tnth m the Dcn11u.m (ma,stn· n:noid pt•rto- ,md tht• cumul.uive evJdem·e IS heavily in 1:1vour of
wc) and finally the f.J~tltui.Ht s\>nngoporolth .1ftcr Cwmmds bcmg corals.
lh,· 11t·vont.lll. Tlm n·prcst·nts an oVl'r.tll tkdmc Polm't?f;wositcs colomL'S from the Siluna11 of
l~•m the range of l~pt·~ L'~t.lbli,hed in tht• cJrly Qucbt·e L·an be produced botb sexually .111J. more
lrJnvician. but al'io .1 ~l'ner.tl trend ,1\\,\} from commonly. by ti-agnwntatlon (lee and Noble,
U\\t\'e type\ and to\\ ,1rJ, .1 IUI.)rc open. ere.~ et 191)()). ",t·,.:u.lll)' produced culome!> alwa)S havt con-
I!JIICntrl.'. iLal h.l~C,, ~hawing ''-'hL'rt the colony ha\ been pro-
l.;~rgt'r tabulate' trt' found 111 coral- duced from .1 ~ingle protmnrabtc ~uch colomL's .11c:
lltonwoporoid reds .111d were rclativd} 1111p0rt.mr, fi1und r.111domly disper'icd. l'opulatiom produLt'd b)
hough they were nor re,11ly fran1e butldcr~ ~ino: fra~:,'lm:ntation, however..Ire f()ttnd in dump\, L un-
ht•y hat! no propn m c.lm of ,lttachiJlcnt Sn1.1lkr ~isting of many colonic~ wtth densities of up to
ububtc' tcndt·d to m rur 111 Jec:pcr '' .ttcr... and 50 m ~ These never ~how ,, prmocorallite. but h.wc
tlsctntl.ltc genera usualh liwd m gutett•r t•nviron- (ircul.lr .tml conca\'c b.t\l'' .md limited van.1t1on 111
rnt,. Ordovicun .md l mn:r ~tlun.m t.lbul:ue~ corallllt' ,Jupe. They prnb.tbl) originated by fi .lg-
~mJII \IZt' arc oftt•n found "'1th t.·-.1rl> ~ohrary menlatton of .1 parent colon), .l'- .m ad.tptatJOn to a
R~~mJ, lonnm~,: .1 r h.lr.lcteri~tiC JS\Ott.ltmn in mudd> sub\lrJte. Sexual reproductiOn here lll:l)
1kart·uus mud~tone~. In l.tter rocks .tlso the two .ue have been comparatively r.trc lor the purpo,cs of
'lut•ntly .1ssoci.1ted, ,111d evidently the Rugo~.l and dt~pcr~al .md gene flow onl>
Ahul.lta l1JJ \llllJhr h.tbir.tt prcferenc:es. Ct•ncraUy T abul,w:~ are not uf gre.ll 'itr.ltigraphtcal v.1luc,
t) ltvcJ 11 conditton~ 'umbr eo tho~c 0t modern but they do \Omerime' occur Ill useful m1rkc:r
J!,, but they .1rc not nonnJ.lly found 111 very b.md,. Tht• Lower Oevom,tn Plwrodicryum ( I i~.
~-t'llC~ CllVHOillllliH~ 5.1-lb). J \null domed t.thul.ltc .th\ ·'Y~ tound with .1
A 'tuth of S1lun.m p.Hrh reds 111 tht• Welsh commcmal tubiculous worm 111 the centre of the
>tdalanJ, whit h .1rc do m mated by t.1bubtcs colony. j, L h.tracrcristic of ~andy tacics, by ronu,m
\mtttou, I 997a) ~how~ how dilTcrt•nt ~rowth with other L orals. Alun~ with oth~.:r fus,tb of tht•
Jtcgtc~ .llld colonv f(mm rn: vailed through suc- same t:tttt's it ha\ been ust•d ~ucccs~fullv to dl'tcr-
1\'C \l.lgc~ of reef dcvdt)pmenr. W Jter dt·prh was minc Jiret ttom of fimnal llllf..'T.ttmn ac:ro~' Europe.
IIIJil JO m (Scoflin. 1971 ). In rhe 1111tt.11 mges,
pltcral !,'fll\\ th lcJ m .1 dommann of t.1bular
Classification
::m As tht reef~~ e\\, ,1 tore dc,•dopcd Ill wh1ch ORDER TABULATA b dtvtded tntn wvcn or ei~ht suhnrdt·"
v md~ were ,Jbund.mt, .tnJ both pcnpht•r,tl .md whith .ue 'iUHc clearly cJnumscnh.:d and eastlv d1\1in
dtJI growth strategies in other tabulate~ gavl' ri~L' guishcd. The (ullowmg dJs>tliCltiOu is sunplJhnl md ~
uomal and bulbom n•lonie~. Jn the reef nurb'1n, sli~hllv lll<lrt· ..:lunplex arrangcuJl'IH " shown in hg. 5 I I
rt\er. mec:lial brrowth '""' prcvJ.lent md nodular ORDER TABULATA (Ord. l't:nn.). luvambly rul,mul.
m~ were common. In tkeper watl'~'> ehcwherc (;eu<'r.lll~cfuupo'e'l of 'mall tnr.llhtl'> With ubuiJc, 'cpu
"I 60 m) periphlr.ll b'Towth donun.ttcd and (u,tt.IIIV 1., in number) n·dun·d 111 ,·ertit~l TO\\~ of 'PIIK' or
·-
! )'
:1-
I
~ tJbular and hm -do mal colomt'' wae the
Thtrt• has been much & .. putc abom the aiTimtics

ab~en1
SUBORDER 1 liCHENARIINA (I Ord.-L. "iil.) M,,,,1w
colon1es. cor:Uiites pri,m.tti, with I(, or m on• H'pl.l .tud
wtth mural pores l.lcl.ing m l.tr<'; <'.g. Udtt'llllll•l
ly F~l~>,ilt'S and rdatcd ~cncra. Some org.tnisms tor-
SUBORDER 2 SARCINULINA (M Otd.-M. Dl v) M.l\\1\ ,.
ll- 1) behcved to bt• t.tbui.He tora.ls (l'.g. C/wctctt's)
c:ocncndn rn~l colnnte'. "irh cuctu.·nchymt l'nl h"lll~
_n no\\ bccn 'ho\\ n tu bt. ,dernspongt·'· .md a honzonul ~p.1cc~ and lomwd hrg<"lv of Ybui.Jr .1ml "pt.tl
he hJs been nude: tor tr.ul'.. tcrring fwo\lthh t(l the ntt·n,lom Up m 24 \lf't.l L I' .'\.wi11111J.
ta l 'fOipougt.~ too. ~lt uuures c!Jimed J\ 'Pltllk' m SUBORDER 3. FAVOSITINA (M Ord.-Perm.). V mlum•
-h Jl/o, however, sct:lll to be endolttlm lmnngs Lulonll'~ wtdl ,Jcndcr corJlhtc~ luvmg mur.tl (HII<'s .mtl
1\ltton, 19R7) and thL· mural pores .md La kite ~hon spmosc \t'ptJ; L'.g. hll'cl.wcs, Ah,colitr.<, t\tuhrlllli•l.
P/nm>t/ul)llll/1.
eton o( fibro-norn1.1l ti~sue of fm•t>SIIr' h,l\ uo
SUBORDER 4 AULOPORINA (0rJ.- l'cmt.): Cor.th v. tth
tcrpm among-..t the \pon~;es. rn ,ltldltl011 t<.)~ <r<'epm~ <lf ... rcct h<1b1t, t:.,uwl.uc, Wlth tJbu!Jc \\ 1dch
ltd polvp~. e:tth "" ith 12 radi.uing tl'nt.ldt·' h.wc 'J'·Ilt'd L'r lb,em; c .g. ~ llic•p<>r·• ·
found in CanaJ1an Fui'MIIl's (Coppt•r. L9H5), SUBORDER 5 SYRINGOPORINA (M. 0rd.- Pt·nn.): Ltr~L
128 Cnidorions
er•·•• ct~r.tll., d,•ndnnd <lr plucclotd. wtth cylindri.JJ straight (Ftg. S.JS.1), though horn-~h.tped and C)lm-
n•nlhct•• ''''"lnlnl h\ honznlltJl tubules. Srpra ofi:en dncal rypc~ .tn· l mumtHl.
•b,~:nr. ubul.••· hnnwnt.1l or funnd-,hapt:d, ~·.t;. In colonnl "icler..lt t1111.1, a~ wtth other colonm.
Srn"~·'l'•"ll. the coralhre' .m~ 111ten omlccted a:. a result
SUBORDER 6 HALYSITINA (M Ord U Sil.): C.u~:rutom1
repeated ;I\eXtl.ll d1v1,mn b~ the polyp,. CoiOI!f
"''''"'c' ni unpcrtnrJtl' n>rJII.t h.1nng m ~omc Cil\e'\ 'mall
wmcJl nbuiJtc tubule' hl't\H't'll tbe largt> tubt..,: c .g. type t' g~..·nettl <111) defined .md re fee> rnamJy to tbt
f/,J/}'$1(C • rebrioshhlp\ ot the roralhtL''· A~ \Vtth the Ruga!i
SUBORDER 7 HELIOLITINA \.l nrd -M Dev.)· Ma"l\'t' there .trl dendrmd. pluccloid, thanmasterOld IUld.
rolomo:' wuh 11 't·pt.uc cnr Jlhtt"> embedded m r.lTcly ••tphrmd type,. ln 1..eno1d sderactinian~ eh;
COCIIl'lll h) lilt'. (fht'\t' .tr .. "'llll'tllllCS cla\Slficd .!p.lrt Jml thecaL' (i.L·. "" .tlh) nf ad_pccnt polygonal coralhtcs ar
h.1\ c '''Ill'' '"'ul trtt) Ill ll.'tt.mr llctocor.tls) e.g. Htlr.•litr• cJo,cly umtcd Jud of d1"epimental or ~cpwl on~
by contr.1~t with the \\ails ot terioid mgosans, whl
Order Scleroctinio (Fig. 5.151 arc cptthcLtl. Plocoid types of coralhtes havr dl
All pmt-LO\\Ct ' I n.l\~ll cor:Jl~ an: included in Order tincr w.tll~ but the corallites are untted to each oth
Sdet'<Kl1111:t L1kc OIY}'•>Jlh)'/lla (q. v .) thL·y secrete an by ths~cpmlent,, 1-t,unose types of crecpmg luln
.tragomllL l'\:thkdett111 111 winch thL· \epta <1re inserted paralleling the tablJI,ttt> Alllnt~t>rn, also eXIst.
between thl· 111L'~l'ntt·nes 111 multiples of six (Fig. In addition there ,lre two other typ<..·s which''
5.4.t~) Atter the til"'t ~1'\ prntmcpw grow, successive not corre~pond to ally wgman type: (1) the mean
cydcs ot ~ix, 12 .llld 24 mct.t~cpt.t .trc inserted in all sLx droid type (Figs S.JSb, S.J6b) 111 which coralln
quadrants. ' I hrcm~h thL·tr pJttL'm of~eptal insertion the arc arranged in hul'ar ~cries with the crmH\. ,
SderaLU111<1 ,trl' lmmedtatd) d.Nillf:,'\li,hed from rhe ab~ent, and LOnfincd wrthin the lateral wall~ wh·
Rugo,,t. L.tt h of thL· l'Jrltn cyrks i' complete bctore run irregularly owr the ~urtJce hkc the convdr
lhe ne~• cyrk !P'ow~. though in 'ome ca.~es tlus system tions of a human bra1n; (2) th~o.· bydno phoroid t
bn.-.1k~ down 111 th~.. h1~hn cyd~o.-... (f1g. S.ISc) m wlm h thL· t entre~ of the coralhto
The 'kdctnn origmm·, J.s a thin basal plate from arranged around ltttle lullock~ or m o nticulcs.
\\hi eh rhe '<'Ptl .tri,l \'l'ntl.. illy. As dllS skeleton In hab1t 'dcracnruan colomes may he branch
grow' upw 1rd' th1.. IO\wr lliJrb'lll of the polyp hangs (ramO\l'). lll.mtve. cn<..nt,ting and crCL'pmg (rtpUr
U\er 1t ,l\ .lll edge zon e, \Vnhm tht> cup there may or fohaccom Sometlmt·, adjJCL'nt colonic' oi
b~..· d1s~cp1menh ·" wdl J.\ wpta. In general. the sJme 'pectes h.s~e to fonn .1 ~m~k colony.
srrucrnre of both \1111ple .md compound scleractmi- lol
.Hts I\ hght .md porou\. r<1ther than solid a~ m the SEPTA AND ASSOCIATED STRUCTURES
R.ugo,,t. In ,~., \'<;r.tl n:\pl'lt' other than the pnm;uy !:lepra arc tom1cd of .1ra~onmc tr.tbc.:culac ('implt
~eptal plm thL "-,der..tttinu diiier from Palaeozoic
compound) nomtally arr.mged in ,1 t:u1-likc s~'<t
group,, ,1\ tlll'lllllllll..d bndiy bdow: independent and otkn With .1 dcnnrul.ue upper 'ur6ce, but Le
C\Jdcm·t• (Ohwr, 19HO , 191)6; Scmcton and uur t1~sue 1s unknown (f-1g. 5.15). Usually the t
Cl.trkson, I 1) 1JO; !'ltrutton, 191J7b) now shows that beculac arc unttcd, though ~omctimcs, paralldr
St leractliH.t .trml' lmm ,, group of sea anemones the Palaeozotc Family Calo~tyhd.tc, thl' trabm
r.tthL•r than from any PJI.teozoic cor<1l. fr;tmcwork i~ lomcly tmttcd .tntl may be pt·rforlt
Such fene strate septa ;rre more impona.ut iu ~1
Range in form ond structure .tctiuiam rhan nrgosam. Th,· t,rrouping of trabee1rl
TYPE AND HABIT and thetr srnsctnre ,tre important stable charart
Sder.ac tlnl.lll\ may be \Oht.tr> or compound. In soli- for taxonomy. They form initially from aragor
tJry ~dcncnnuns tlw tt.mn ot" thl' corallum depends spheruhte~ 111 .1n org;ma m.Jtnx (SorJ.uf, 1':172).
on the rel.mn· r.ltL'' of \'Crttcal. Jnd peripheral The <>epr.l ori~., n.llt' between the mescnten;
F1gure 5
growth OJKL' rhc h.t,.tl pl.!tL' has been secreted. wh1ch .1re .tl,o the.: 'He of d1ge~tion, excretion Recent m
Where pn1plwr.tl 1-.rrmvth Ius been donuna.nt a dts- gonad devclopml'nt. The,e ntt.~L·ntenes have ad 0.0 hydt
cmdal fonn r~..·,uJt, (Fig. 5.1€1J). often with everted blc layer of endoderm 'eparatcd b) a tlun iliecr M,cranx
(1 e divis
\cpu m C.hc' wh~..·rt• tht''<' h:t\ l' grown rapid!}. mesogloca. MtN·Je, ,\rt' arranged on one :.ide of
sderoctin
Perh.t~" rill' t ommom·,t kmd~ arc thme wtth mesentel) onl~ (F1g 5.4c), and the mesentcn~ (Of,so .- ~
rurbmah: lll' t tlnll'.tl t"oralb m which the aXIs ts group1..·d m p.1m \\olCh thL pleated mu,cle bl f 'Wtth th
Class Anthozoa 129
t•l Montl1111tlti11 (bl M e11ndrfn11
(fl Goni.stn•
(el Gt~IIIKIIII
(dl Mic;rltrbllcill
(n)~ent
ex
8 nl
"--- - - - - - (p) Lophelill
7e 5.15 Sderodmion morphology (a) Montl~t~oltio (Jur.). o well-preserved specimen (x 1); (b) Meondrino, upper surface of o
a; meondroid coral, showing o growth pattern resulting from introtentoculor budding (x 1); (c) Hydnophoro (Rec.). upper surface
'I- ol,dnophoroid coral, showing o growth pattern resulting from extrotentoculor (circum-mural) bUdding (x 2); (d) synopticuloe in
f wcio (Cret.-Rec ); (e) trabecular structure in Go/axeo (Mio.-Rec.); ln Goniaslreo (Eoc.-Rec.). showing distomodoeal budding
~ d'vi!IOO of 0 smgle stomodaeum into two) on the upper surface (x 2); (gHnl the origin of poli, showing (g) 0 sextant of solitary
~mon, (h) the some divided into layers and (j)-(n) transverse sections of po yp and corollum at each of these levels; (p) Lophelio
qgo.-Rec.), o dendroid deep-water caryophilline (x 1). (Redrawn from Wells, 1956 in Treatise on Invertebrate Paleontology, Part
.,~the exceptions of (a) and (p); (p) is original, (a) is redrawn from British Mesozoic Fossils, British Museum, London, 1962.)
130 Cnidarians
(a) (b ) p11013ry
b\ Ill
Ill cc
31 Ullll~
th• C Ot
xmltN~
but lt
lmlang
lJVIIlg li
tll.l s.
SUDDII'
I hl Ion
dc t cnn~r
rdatlvt·
hccll d i\1
Figure 5.16 (a) Fungia, a large Recent solitary sderactinian, showing several orders of sepia (x 0.5)i (b) Meandrino, a Recenl~ lnt 1.1t1
niol coral from Baroados with typically meondroid structure resulting from introtentocular budding (x 0.5; cf. Fig. 5.1~ J, \i l
(Photographs reproduced by courtesy of C. Choplin.) d;utgh
or m e
f:1ung c.ll h otha. Thl' space within ' u ch a pair is Di,~cp un e nb, like tho'c of the Rugosa, arc ~elr wod.1~
thl' c n tocoel. \\ hik in bctwt:cn (the muscles facing by the base of rhe polyp. As the latter gro\\ uch
;n.\a) trom c.1rh otht:r) 1s the exococl. llut two pairs moves upward,, dctadung a small part of the llllh·d
of lllt'\l nrcnt' on oppo\Jtl" ,Jdcs, the di rective from the l'ahcc at a tunc and fornung a disscp (I Jg.
m esen ter ies, h,l\'e the mmde pleating reversed. to cnclo\c cad1 space. Where the fute skeletal 1ntr:1tc
gJ\'Ing .1 b1l.llt r 11 ')'lllll1l'trv to an otherwise rad1al turc of d1s~epuncnrs have been studJed \SO ~Xt~
coral. lfthl· n>r.tll\ dhplll'Jl thc long .1xis extends in 1972). d1sscpuncnt~ have been shown to have type n
the dirt'nJvt· pl.utt·. layers. a hest-formed pnmary laycr. overlam b) [CIIllt: l
SLpta .Ill' thu~ CntlKOt'Ul' (entosepta) or cxo- neatly ahgncd clusters or aragomuc spherul::. la1 bu.:
col'ill (cxoscpta) 111 ongin. usu.1lly rhc first two Endothel'al d1ssepuncnts arc contined \\ I ht• t:
cycles an.· olt·ntost•pt.l, the rcst of exosepta. In some the coralltre, but 111 SOI11l' colomal scleraonu11.1 (wheth
sder.KtmJ.Hls thl'fl' Ill.!~ bl· a peculiar grO\\ th system cor,1lhres arc un1tcd by a conunon spongy ussue pi ol
produung VI.'Llll.il pill.m (pali) along the inner coen osreum, wh1ch may be formed p.uti.t!h 111tegr,1
edges or SOilll' l)[ thl· mtosept.l (Fig. S.!Sg-n). exothecal d•~sepunent<; and partially by rods and pruduc
If a rolumdl.1 tunns it ts .ilways of septal origin lars (co s tae) , as 111 Gaf(/xea (Fig. 5. lSr). hnwcv•
and may bqpn .Is p.d1; 1t I \ usually a central ro d or a Tabular dissepimc nts arc flattish plates cxtc rill' ~oti
div1ding pl.lll' ti.m~~t•d from .1 proseptum . ing across the whole width of the coralhte or llc not
The septa arL· ollm l'onnL·rted by noss-bars called fined only to its ax1al part (thm resembling tJbt
syn a pticulac (f 1 ~. 5. ISd) whid1 grow towards in rugmam) . Vesic ula r disscpiments arc 1 Evoluti<
each other from tht· w.tlls of adjal'ent scpta and arched phltes, convex upwards :md overlappmg. The okk~l
C'-t'lltuall) fll\l', pL·rfor.Jting the mesenteries in the usually inclined downw.ml~ .md inwa rd~ !Torn 1 hey
prOCC\\. edge of the corallite. thetr tirst
OTHER PRIMARY STRUCTURES SECONDARY STRUCTURES

The thm b.l,.d platt' '' ~cnutransparem and tlmlly S te reom e ~~ ·m .1dhcn·nt l.1yer of secondaf)
,tdhcrcnt ro thl· 'ubstrarum: 1t nta\' later be thick- whtch m.l} cover the septal surface. lt 1~ con
ened by 'l'l'Oillbrv dcpmm. The ep1theca is not of transverse bnndlc\ of ar.1gomnc nccdle' Stc
developed 111 lll.ll1} \l'lcr.JCUnians. bur if present may nonnally rhtckcns the cpitheca 1ntemall} !'
(omi't of dll'\TOn-hke av,t.lUitcs of aragomte. bur 111 some C.l\t'\ lt\ fimc non 1s taken met tlu tlmc.
Class Anthozoo 131
rnmuv thickening of rhl' \l'ptJ, by di"epimen~ or Jurassic and the fi.r;c tmt: decper-\V:Jter ahermatypic
i)11.1pticul.u:. corals (Subo rder C:~ryophylhm,t) emerged at about
In compound ~dt.-r.ldimam ind1vidual corallitcs that omc. though these were of lumted importance.
u,ulll} 'cparatc:d by .1 l'omplcx perforated tis\ue: By the MJddk Jura'\\IC \deracnman~ were on the
. cocnostcunt. llm n1.1y cOIN\t cntirdv of incrca~e evcrywhert.', .md p.ltl·h red\ cntpted wher-
11h~ol d•"cpmleliC' (t.1bul.ir) or other material, ever condmom \\t'rc ~utt.thk. In the Tt·thyan Oce.m
but lt provide' a 'uppnrt ltlr numLrom canal~ the tl.r;t large rec~ developed, pl'N\nng untll a general
linhng the mdi\'1du.1l t ur.tlhtc~ and bmdmg the '\etback in the early C rctaceuu,. Lltcr th1. largl reefS
mgccher 111 a tuncumully cohc~ive and cot'.U b.mks devclopt•d, .md ,1lthoub>h \Oil1e of thL
older Cunihc~ d1cd out other~ .tro~l' _ Th~o· \Uccc" of
reef coral\ fluctuated in the C'retJceom, .tnd the aher-
matypcs became really 1mport.mt for th1.· firo;t ntm•. By
lom1 of dw colon> lll tht• ~clcr.lLtinia j, largely the late Cret.1ceou' both hennaryptc .md .1hemutypk
tkR"mlllll'd b} tht: tuoJL· of budding combined with cut'Jls were becoming <.1f dJ~bnnly m odt•m type.
I UOVC ~'TOWth rate\. 1 he folk1Wing type~ have though like many other ~:,rroup~ they sutft:rcd a major
'U ru\tln~Li\hcd: extinction event at the t:nd of the Crl•tac~:ou~. By tht:
end of the Eocent: the do n1in:mt b'Toups 1.vcrc much
lmr.ucntacular buddmg H en· the polyp divides like those of the present ~in cc the formerly important
olo-
2b). b1 'imple ri~swn .ICI'll''
the '>tomodaeum, each Mcsozoic 6milics (e.g. the M o ntlinlittidac) had dis-
dJ.ughter (of wlm h th1.•re may be two, three, four appeared. During the late Tctti:try the two main coral
•r more) rct.11mng part of the ongmal '>to- Etunal provinces of the prt.><;Cnt the lndn-P.ll:ific and
~red ltlud1l'Um regt·m ratmg the rc't Tht productl> of We\tcm North Atlantic provtncc\ lud become dj,_
''> lt llllh polystomodacal budding may remain tinct, and reef- but.lding corab floumhcJ on an
base dllked or bet ome 'cp.•r.llc- Mcandro1d corals unpreccde ntt:d '\calt•. Many reef'> were killed by the
1t"J1( :f1g 5.1 'lb) .1re dear!) the r1. ,uJt\ of mcomplete lowering of ~ea levd m tht' Plcmocenc, but their
ntc- !utentacul.tr buddmg. remains have proV1deJ \table \UriJu:~ for the rcb'T'Owth
rauf. btratc:marular bu,!Jtn~ ( F•~· 5.151). In tlus of cordl ~inre thc1r submergence Compat'auvdy few
cv.-o 11-pe OC\\ \COIIIOd.1e.1 .m: prodUCt"d Olltslde the genera were .tlfel'ted by the condmom of the
ver- lrntlCIIIar nng of rh~ pan•nt, fhe~e extratentacu- Pleistorene, ~o that PltOL·cne c.: oral tJun~ts arc etli:c-
lhtco;. bud\ ,oon ~cp.lr.:lt\: md do not remam linked. tively the sam e as tho'e of the.> prt'~ent.
;chin The colony ch:lt n:sults from ~uch budding Most tanulies of \der.lllJtiiJm (Yeron. 1995)
iS t\H; "hethcr branrhtn~ nr ma~sl\'l) thuo; comt'\ts 'im- show evolutio nary tremh, espenally from sout.lf) to
~.rh~ ~ of numcrou~ \Cpar.He tndr" 1duals ,md ts not compound roralla anJ, in colom.1l t}pe~. from
ly of ·egrated fum:tton:tll> 111 the ~:~me- \\ay as the phaceloid through plocoid .md cen01d to mean-
d ptl- •mduct' of mrratent.1ru!Jr budding. There may, droid tom1. Tlus probably uH.reased the degree of
owtver, be some m·rvous or chc-mtol lmbge if coloni.1l incegr<•non .
rtend- thnoti tl~~ucs an.' 111 cott t.1ct, even tf the enterons
r con- , nor ntttrcd. Ecology
ablll.ll' Scleractinians E1ll into two m .nn phys10log~cal cate-
small gones: zooxan thcllatc .111d noo-zooxanthellate
tg, and 11IJN known sch:r.lt ttni.ms .ue Middle Triasstc in (approximately cqUJvakut to the ecolo~'lca l tenns
1111 the llwy denved ti-om .1 !-'TOllp of ~oft .uu:moncs. On hcnnatypic and ahennatyp1c). In zooxanthcllatc
fi."'t .lpp<.'.lratKe th~.·y .m.> .urc:tdy differentiated corals the cndodennal cells arc replete w tth <;ym bl-
a number of unport.mt t:1nultc'>. Hennatypic in otic algae (zooxanthcllae, wh1ch are J kmd of
tht~ formed \111JII p.1td1 red\, m.•y haw pos- dmoAagellate). These symbiOnt\ benefit the corals in
..; (1~\U• zoo...,lnthellaL' .md w~.·rt' confmt•d eo wam1. two respects. Ftr<.dy. up to 95"il of Lhe org.uuc car-
~lpo,~.·d ~ well-hr W.:ltl'f'i. By tht• Lite 1 na,,ic they were bon produced by the zooxanthcllae ~~ t.1ken up hy
:erCt'IIH ::JiJ ~£1st m m.my p.1m of tht.• world. Jnd many the polyp and used a~ food. l.ie~.ondly. zooxanrhellate
a" wdl, Jrtht'. I he tiN 'null coral ree& date from cor:~ls :~re able to grow up to three tunes more
:>Vcr b 'l:llt' l'ht\ p.mem pt'T'il\tt•d chrou~ the Lower qu1ckly than tf they had pOS'\C'i~ed no symhtonts due
132 Cnidorians
ro the , xtn () l'rodu. 111111 b th~ tlg.w l he''- .dga,· tuund 111 nth~t or~.l!ll\111' '"" Jlw ctoln!-,ry ot nwa
,utd tiJl )',In'~~~ I.'S,~llll:JJ
l Xl~t Ill cll•'rtiiOil~ lllllllll('f'i, t'tll tlll:lb h ~~ bl't'll m ut h ''"d1ed: U\t'lul rder 111
''' rh,• tllc.:t~holt,m nt thl· l or.1l uppl) '"!; 11 \\lth .He YollL-:1.' (I•Jt,S), ,\t1u~. umc Jnd l~nlwiT I lu
tllllncnL' ntd t.n.,gcu th.lt ~\ll h lOrJl~ c 1n ''Ill) flour- L:\\1~ .111J Pn.e ( JtJ7!=i) l.all .ntci { •1.11CS I OlC
hh 111 the phon Z•llt~. Jt depths nonn~lh k" dun ( ;on:au ct 1/ (I <J7lJ) .md Vo\hmgh (1 'J~2)
50 111 r.ud) 'Ill m) S111, l' the cnr.1l~ hJ'.: firm
.mchorJ~~.: th ' gro\\ \ih, 1.'~\lulh Jnd nu"t Jbuu- Classification
dantl) \\ltllln the zone of\\3Vc JdlOll .•ll depths t.:,, In d.mtt) 1111:) rhe Stll'r.H ttlll.l, 1S \\ Hh the R:
than 2•J 111 and I .tbul.lt,l, ll h.1, h<.·c·n nt'< c".u) to hnd
Reet-bmld1n~ t•r hcrtlllt\ ptc: c••r.tl' :Jrc n.:.lfl) Jl) [:\XIIllUil\1~ l h tr.ll [CT\ ~l pt.iJ \(nlltllfl' tall bt
zuux liHhdl .111d ~cucr.tllv m cd ,, ullntruum '' Jtcr
tl tor dctimng \tthmd111.11 t.llq?;ories \\ luht 110
tcmpcr.Hur~ PI I~ l .md floun,IJ bc,t hctwccn 2;:i u•lnll) to1111. t1n11 " nt' v.dm• onl) .tt the fl
md :!'I'(' .md 11 nnrnul ,,Jinllt\. I Jcn,c mmt r.:ct· k·vcl. ll.thlt.tt 111.! dnmn,lnll\ .Jt .Ill)
l.lWJI' .u <" n·~n 11 red tu ,h dln\\ . '' dl-ht. '' Jrm \\-.Her. c.ltlull .Ut' nflurlc v.tltll'.
In .tddmCIII, rhq llCl'd 1 );OPd 'upph <>f oxygen and
gl.'tll'r.lll) pn·kr 1 lt1111, ll• •11-!llUddy substrJtc fur ORDER SCLERACTINIA (M . I n,1, !l..n.): St•Jtury t>t"
, lll.ll, \\ 11h llh'~<'l11<'11l'' p.mt·d Jnd lht· wp1.1 .uull~l'tiu
111 hcr\\'t\1' tlw pl.lllld.w \\ 11l not Sl'ttk. There ..re.
t1pl,., o1l , ,, l1.1S.1l pl.1!t' !-:IVC'I "'< to 'tp1.1 .111d l'('tlltr~
hn\\ I.'\'~ 1, hl'I111.11VJ'<'' ltv111g .111d building; reel~ in SUBORDER I ASTROCOENIINA (M TIIJi
nm,ldv 1cgu•n' l1~h1 " pc.:rh<1ps the IIIOSt unpon.mt Not 111.dl\ cnh llll.tl h1 n11.1t~ P'-'' \VIth Slll.lll , ,,,,Jillf
llt c1ll eh, luliHIIt ~ t lttnr~. rht•ugh .t red c:u1 be kdkd nl 11(' 1n Cl~thr rr.1bn ul.tl. 'PIIIII'<' tn l.untu.l1, e ~· 1 In nu1
h\' ud.11Lll1LI •ciHt' dunng III.'.IV\' r.uu, dunng ,1 hur- J .,,,1,1/lol ,, ,,, jlt 11) ' (
11• ;me tor 111~1 tllll • •1nd 1111) t.tkL· 2fJ )'t'JI"S to re- SUBORDER 2 FUNGIINA (1\\ . T11.ts -Rcr ). "''b V U•gct
' ,t,1hhsh 11~, If .. ,lunul wnh p•tlnt.IIC: SC'f'll hnk"l h, '\n.•r
( <•r~ll•r~, ll\Udll} lu~c· M.nnh lwrm.ll\1'1<, c g I
N.m-zuox 111thdl.1tC l oral~ JTL' lJr~dy .1ha- (:,·.1.•1•11>, f,,,,,,,.1.
m lt} ptr (t.•' 11~>11-lc.:d h'llllcr~) ~'1lllt' do hvL' m SUBORDER 3 fAVIINA (~\ I n.h.-1{,,·,.)· L;unuur
,or.tl rcrt~. though othl .-.. .uc lnu11J at depth' nf up "uh l.m-hl..c 11 1b ul1r I >l~'l'l'llll<'llt~ pr~<nt btn
tO nOOO Ill I hn ITC. ho\\e\CT, lllO~t abund •• nr 111 ul u· r I< \.\ unh ht'rlll.lf) pt<. < ~· lur·ci<'S, \1
down Ill :.1111 m ••llld though the' c.ut 'urvi\'c rem- '171niJ.</IIt/iol
per tturc' hdo\\ I) { • th,., t1oun'h hc,t hL'f'\\ Cl'l1 SUBORDER 4 CARYOPHYlliNA tlur Rn ) Non
l.lr\' Jnd .1ht nn ot) pll. :-.epr o l<mlnar "nh ~unplr tn
=i md 10 C. hmh\:tllll•I'C. thc'i can hw m rota!
I.Jc Ill t.1n ,Htllll; r.u< '} n.<pttcul.ll•. e.g [>
,l:u km:,~. <het t \\ t•-tllltd\ of these 'decp-~ca corab'
( '"l'•'f'h}'llhl
.m ,oht.lr} 111d dtl 11111 hnm rcct\. thnu~h rhc u•lo- SUBORDER 5 DENDROPHYWNA (U <'n:I.-Rt'C..
llt:tl dl'lltlrurd c.u) ••ph} lhll L.t•phd~tl (!-=t~. =i.ISp) nr ,,,Jnnul '>l·pt.l ltnllll.ll bu1 trrcgul<ul) pl·tfi•orr
lot m' Jt•cp-\\ llt'l b.tnk~ or thtch·r, Jt the cdg, uf nt •mollcu S) u.1pttutl,,t' M.Juol~ ,Jhcrnlllypt
rhc "'''l~tll l·umpt.llt ~.otHIIIU11.1l ~hdf / ), 11./r,lJ'h)'//lol
fill' 1.' lrliL''l ( 111,1\~ll) \l kt,h tllll.lll\ Wt'TI.' I.'Vi- th 'llllllll
den tly ht'tlll.tt \ P'-''· l hnut;h pcd1.1p' not 'll adapted Coral reefs tf1c l'<lgl'' t
lot tht· tl'd lublt.ll ·'' tho~;e ot the Pfl'\l'lll -;int:l' they Rl'lc lH uu.il r~ds (Ymlgt•, 1'J6.'\; Stoddanl, I 11 " 1101
lotlltl'd llllh 'n1,1ll lt'cl' Ahnm.lt\111t' corab dl.'vcl- Ladd, 1971: l .11lll'tnn t'/ ,If., I 1J74: Shepp.trd.'
op.:J twill u\.111\ ht11li.JtypH \toe b h\ tlw ,Jo\\ Verun, llJX(, W db, I'JXH) .tn• confint·d lll ln
'prt'.HI 11l 'l'rl,llll l\ pt'\ tlltCI CP)d,l, dcepcr \\',ll\.'l"i. \l'•" .md .!Cl' bt:\1 devdopl·d 111 the lndo-PJnh
lltl· h"t .knu\\ 11 .!lll'tlll.lt\ pr' wcrl' the JurJ\\ll. .111d 111 dw \\L'\tcrn Nnrrh Arl.mnr. Th~ lndo·l'
t ,\I\ up!!\ llni' "ltu~c dt'\l'l.'lld.mt-. tl't,l111Cd thl' hJblt. ru:l,, wht·r,· IIUXIIII.lll\' dn·dopt:d tMcbn
but tt p1c,t'lltJII\ ,., ot m.lll) nthl't f::.'1'0l1P' ,1lso '-.uuthe.l\1 Ast.l), h.l\'l' ')2 rnnrded genera and
b~ u•lllt' .1lwn Ill!\ ptc. 'o th.1t the hcnnal \ pt~Jhcr gcnaa .md O\"l'l" 71lll 'J'l'l·i.:, (~c,· Vewn, 19
llllt\ pe dt, 1~11111 tnd" 111h ·" 1o" tnan\ tam!lteo, and tllmcr.mom .1nd dl\tnblltwn m.tp,). \\ luk the
l:\ L'll gcllCI.I t:um.t ol 1h, urth Atl.mtir '' \lllJ.Ikr \\1th 01
lt rs ltkeh th.H rhc gr~ 1t t'Xp.lll'lllll uf tht· her- gl'ncr.J .md C.2 'Pl'lll' 1 ht• ~nrth AtbntJ
rl'ef htnlckr' \\ ::&\ chrltth huk,·d '' llh dtl.'lr
lllll'\ pll ,eem' w h.l\, bt·~.:n tknH·tl uut•.•lly trom " Jl
o~dopuon ot z.onx.mrltdlac Sm h ') 111bwtlt algal.' <1rc \uunc, Sllll { )h ~oCellt' rt't'h nf thl.' '1/..'c~t I
Class Anthozoo 133
h" ~itH c dnci-

llll' 1 otal,, h111 11 utc 111 tiow. I ht• III.IXIIlllllll r.ltt' ol 'ub~1dcnce h.1~
PJI!h ul.trly \lilt e the Amen< .I.~ bct:ll 1bout So cm 1wr Y'' 11 ( 'unltml.ltor) ,,l,llllt
rh~ l'.uJ.IIn.• 1\thlnu, 111 the .llld dnlh11g evllkllll' fnup oth(r ttoiJ, mggc~t\ rhtt
()('l'llt thi' •~ the llllllllhlll I' ttlt'nt I he now sunk~:n \·ol-
struc tura l r eefs d1( st k·1 •~.tuuJn' hJ\'<' butlt unil ,,lmd~ olfO\l 11 'hm <.pot~' 011 1 ~uh111.tnn~·
nusm ( \\a\ c-rl'\1\tJnt \ttw tlltl''· l>ltcn to L'Tl"at ndgc Lft:~t Jn,i hJ\ c h~ en c.1rned "' J) 111to dccpl'f
:rkn('"l', ''here the hulk nl tlw nuten.tl ~~ con- \\Jtt:r~ (:1' on .1 uul\l'\•lr belt) " the oce<111 tlnnr
red l->\ C<>r.ll\, C\ en rhnugh <.!I< trl'Oll~ ,II);.IC 111 '} ~pre.1d 1\\.1\ fn>m the- n.lgc (Sloflm .md I >Lxon,
n unpon tm part 111 reel Mrucrure. ~~~~ h .t l.n~~: l'lSJ).
tntcmre htnlt up b) ~~o:lll'r.HtOIIS nt torJI\ 1' the I hr\\ m\ tlll'OI} ut <Hull ltmuatJOII, though 110\\
ltJt ot a hno,r of dJ\'l'r\t' t>tg:llll\111~. tlu: \\ht>k 'tndiL.. Ited, did IItH t4kc uno I• unutt Pl<-1\to( tm
u ng one lit the lll•J t < Plttpk·x Cl my,tem~ th.ut~c' t•l ~c.1 lenl, '' l11ch ''''re C:1r r adung \llllt:
1- m. In co1.11 red' ptPdtll t" tt\. t.dlllllll met.tbo- the reef, \\l'll' expt"' d .111d thl· u>lill' kdkd nil I h~
'11 Jnd r.1rhnnatt' hxHu•n (J'itlll g '·• ) .m· vt'rY
1 t'XpO\l'd \\lll,\lC\ \\l'IL Clntkd tO \l}llll' l'XtCIIt 111J
1 l"ht• r.tngc ut ll.lhtt.tt\ 1\ \Ill h th.ll !here " .1 ut ten trt'ttt'J llltu .t k.mt tupogr.tl'll\. llcnu: the
~l tle).,'lH' (lf,pet J.d17ltlnn 111 tht• ·'' <>< t.tlni t:111n.1. vent't'l ut· l or.1b whtt h h.1~ g1o\\ 11 'Ill Cl' tht· end ut
I'• 1.111y tlo,hc~. mnlltht ~ .111d .lrthmpoth; the'c ,Jrc rht l'kl\tolt'llt' h." lt'•l'\t,tbll,hed ttselt on .111 .mom
.1 m znned .1nd lot'.Jh/t'd llh Ill'' 111 v.trmtt\ p.11"ts .t!um tnpugr.tphiL sud.JLe, .111d 11 is I'L'ltUs•· ut tht~
ht•rtt{ th.H tilt' ll11111 uf .I llllldt•lll I cl'f 1\ lllll \llllf'h .1 rdJe(-
1 non- reef con1 n tunities, IHn\ t'\TI, rhnu1;h non ut cunstntttiun.d atll\ 1ty
:ill\ rontmcd ttJ '"'P" Jl ~c.t,, th•· 1 t>rJI, n•erdv
t ·~crht'l \\'lthout bmldu1g Ill' .1 n~1d \lrll• tu re. Growth and zonation of corals
nr:-tl\ ••n ret'l' h.I\C ll'\\ Jhlt\11 d l'lltnlleS othl'l tiJJn
1
ypes and genesis t:mal-c.ltlllg hsht', 111d 1 he ~t.lrllsh "I mrtlu~lt'T 1 he
tr.l! coni ret'l\ to,! I\ lwlnug to three JTI.lj•Jr IJttcr ha' fl( t'ntl} "lt·.•kt·d IJJ\Ilt "" \llllll' I'J<IIll
fringin g r<'efo,, barrier r eef\ md atolls. n:eh. for atte1 rlw depn: d.tuum tJI the ~tarta~h the
re ~ de\ t•lop .tlong shnrcltrw,, e~pt•n.•ll\ <k 11i cnr.1l be1t>llll'\ t:o\ l'n·d \\ nh ,111 ,tlgal ftlm
of volt.lllll hl.llhk lhnwr rcch .1re tnnlll'd "l11ch pn:' ems t tll,ll pl:mulu· from ~etthn~.
di\Wlle frnrn the 'IHlll' Atnll' arc nn-,llar <1r H..epC.ItCd .4r.711/fr,l IIf 'tll\'1 lliiiS•, Jm\\ C\ ~I, .1ppe.11
be u hoe ~h.lpt·d .lllll li'UIII~ 11ncnt.ttt:d w1th to folln\\ ::1 lVlhnl p.•ttcru \\,lXIII~ .md \\,lltmg U\'t'r
IIIIIUJ to the prt v.uhng \\ 111d. wuh n ccntr.tl ,lnl- nUll) rhnusHah 111 \l'.lr\, .md thl d.lllllgc to tht• red
'liOII 111 \\}Ill h \lll.tll p.ltl h fl'l f, llt:J\ <fl·\·dop. " evcntuall} rq1.Hred.
.rut B.•m<'r Re<'l (r-..1,1\\\ ell. IW1X, lknnerr, fkcaml' of the \tl't'p l'IIVJrunntt nt~l ~r.1rllems
tT caMcn1 Athtr.tlt,l " 1 'Jll'tl.il r.N'. hem~ .1 \\ Hh111 thl' rt'l'l tnmplcx. rhc tt>r,tl' them,cln·s Jfl'
f(~tiiTt' p tr.11ld to tht• t nllt-houndcd m.trgm of ccologic.llly 70nt·d w1d1 'pentK rt·kn·nre ro thC'
llllltt 111, \\ lll'rt' t or.tl ~n >\\1 lt \\ ·" 111111.HI'd nn 'horelim· (r:ig. 5.17h.r) I·.H h 'JWl 1c' ut tuptl'' .1 p.u-
gt~ uf .111 upr;II\L'd fttd!. ttrul.tr h.1lm.1t .tnd 1' nm tou11d d't'Whl'l'l'. Wnh111
(l)(l') I. I 110\\' WL'II ktHl\\ 11 th.tt t ill' thrt'l' 111.1JOI" type~ p.1rtHIII.1r h.1h1t.1t' tlwrt· j, .1 l\111d ut 'pn king nrdl·•'
1'18\ ~ l ll tll'S\IVl' \l,tg<'' 111 the growth of coni 111 cor.th; donn11.1111 ' IWCil'\ m.t\ ,H t\l,llly l'.lt other
·op1,tl 1d J 1111kmg volt.lllll hl.n1d (l;tg:. =).17.1) ~pcru:~ ot- till' ~anw ~l'llll\ h\ llH'\L'nler: l xtrusmn
i< .1r~ ~-orro\1 th ,,r <or.tl' h.\\ kept p.Kt' ,.,,th \ubsl- .md digeo,non Ditli.·n·nr 'Pl't lt'' mh.1b11 till Wind-
P.tlltl ll hllc hrokt'll rn·f t tltl\ h." torml'd , ,1\t.Hlm~ ward .md lec\\'.lni \ldL'' nf tht· 1\l.md. 'I hnl' ''· fur-
ug du\\n to dt'pth 011 the Ollt,Hk. I ht' tht·- thermore, .1 gre.H rh.mgt· hnrh 111 'PL'Cil'' .md 111 thr1r
gllllll\ \l1ggt'SIL'd b) I >.11 \\111, h.1, ht:l'll 1 on- !,.'TO\\ th t(mn~. from rlw uppt'r p 11 t nf the- filr,·reef
b} lcl'p btmnb"' Ill !I> ,lttllk At En I\\ L'tok 'lope 1Cro~' the red· fl.tt, to dtc 1 cntJ,tl hgoon \nth
rro Ill 1'151, b.1\.llt \\.1~ ~lltllllliiCH'd .11 ,Jl'Olll ih 'hdl ,,md flnor .md p.ttl h rt•d\. In '' 1n1e Cl'C'
bdo\\ the ~urf.1u:. I he ltmL''tolll' o\'crlyuq.! lliJ\\1\'e rounded cor.Jh ~111 h .Is D1pl.m11 tend to
1 \\al E(llL'Ill 111 .tgt·. fhus hll\\ t:tnk ~~ ,1 inlubtt the 'urf 7om•, \\ htlt• rlw 'tnllt hratH h111g
~ pul:lr 1.25 kn1 tlm k ''·'"'hn~ ou .1 dt·cply 4•'•'1'•'''1 (rhc \r.:tg\ horn' {or,tl) ll\c' .u ~reltcr
\Oic.ttllt ,,l.md 't>lllt' 3.2 km lug:h nu the (kptlh out of rhe r.mgl' ot ,urt .11 t1on. In other lrt'J'
134 Cnidarians
(c, 111.1\"i
reel llat Am1p
0
19H2)
th en~ I
cculo
lll Cl
UllpOI
m Zu
\ '.tnO\
fi>r 111
tion.
go mat
thc lr
60 \0 .lrt.'
lr1n11 •
h.1bita
The
t.'I1Vlft
(b) massive soh reef upper low er
roelgrass dead corals llvrng corals octocorals edge reel slope pl'ryc
more
rt.'.H:h
0
1t ha.,
W11
t10n b
m
also a
equall•
and c.:c
my of
tht.' lajl
(d)
Dee
I>t·l·pe
.md L
trom t1
of the:
toloni,
'-'
l m
............. ftMrh
The
u
l m
I <J64) I
thl'rc i:
u by oth
lm
whir
111
or of d
rnnme1
hms ;l
l haract
figure 5. 17 Coral reefs. (a) growth stages in the development of on atoll by volcanic subsidence according to Darwin, showi~
fringing reef (top) a barrier reef (middle) and on atoll(battom); (b) section through the edge of a growing reef, showing zonaltoo t.1l dcb1
based on Monouli reef off the northwest of Sri lonko (coral genera ore represented diogrommoticolly: 1, Symphy//io recl:T. prm,di
Acroporo formoso; 3, A hyocinthus; 4, A. humi/is; 5, Nlontipora folioso; 6 , the soft corals A/Cyonorio, Gorgonorio and Anti~ otht.'r a
7, Glovio, Pocil/owro, Gonioslreo and Poriles); (c) coral zonation on the growing edge of Amo Island (Morsholllslonds; I, Pori
dominant, 2, Pochyserio dominant); (d) stages in the development of a deeP-water coral bank. [(b), (c) Simpli~ed from Mergne< bt: 'eve
Scheer, 1974 and Dohl et of , 197.4, respectively; (d) is redrawn from Squires, 1964.) mubric
Class Anthozoa 135
1\~ cor.tb arc commoner in quiet waH.'I"\, .md b ank devd<'p~· a topographtc enttty \\1th .1 core of
:rap.>r,, occur. in hi~h-ent."rgy lOIIl'' (Vo\burgh, solid ~kdct;.ll dcbn,, .1 lO\·enng mat of more open
~~). The appan.·m ~tructuraJ ,trength uf cor.tl' '' dchm and a capptng of live coral. The proportion of
rcfore only onl" factor an1ong-t m.111y th.u t ontrol living to dc:~d coral dccre.tses wnh rune.
,,]O)!llol} ~Oil.lllllll; ability to COlltpell fi1r 'P•Kl" .tnd The cxpmed ~cqucnccs studied by Sqn1rt'\ wae
I• cope w1th 'cdimcnt uiliux may lik.ewisc be Tcrt•ary in agl', but 'imil.lr b.mk~ arc fonmn~ tl'!thy.
nport.utl. The study of fo~·al thickets, copptCl'S and hlnk~ (';111
/nnmon v.u1cs ti·om reef to rt."t."f lk·pending on yield valu.tblc pal,tcoecological data.
~,,.,,
facto!'\ I' he pre~ence of re'i~t.wt albral 1idgc~.
tmtJnce, l.lll grcarlv affect the p.tttcrn of zona- Geological uses of corals
. Not onl) the ~dcracti.JuJJI\ but al'o the gor- Corals as stratigraphical indicators
ru.~m (111 the C.tnbbe.m reefS) a.nd m:wcor,tl\ (in Corah .trc gerwr.lll) too long-rangcd for U\l' '' lone
r lndo-Paettu: reef~) arc ecolo~'lcall) zoru:d, and fo~'ih, though the widely di,tribmed !'tr/,uwrrlw
MC the vam.1u' b1valves and otlu:r nwlhi\C\ that provtde' ;r tinl" ex.tmple of .1 useful :JCtnt' bto:wn,ll
~n 'Peetahzed communitie~ in the verv vaned reef indJcaLor fl>r the l.1tc Llandovery-e.trly Wt•nllKk.
~1t.1~(Yon~c, 1974) Ncvcrlhck\\, cor:1l' h.tvc been mt·d. whc..•tl' no
l hl' growth r,1tc of corals varies gre.ttly With tbc shorter-ranged f(mlb arc available. c~pt•c..i.rlly 111
nronmcnt md the ~ea~un but m.ty n:ac h :2(, cm the Carbonit(:rous. Thi~o work bcg<lll 111 lkl~um
rv~.1r 111 branching corah and 1 nn per~ l'.tr 111 the and in 1905 Vaughan, working on tht: l owcr
ore nus~ive rolo111es. Reef growth .ts J whole ma) Carbonifcrou~ ut Uri,tol. England. l'rt:l tcd .1 /Otl.ll
h.1 maxuHum of I 2 cm per ve.tr, but 111 the pa.~t scheme b,1,ed upon the fir..t appc.tr:tncc of cor.1h .1nd
Ius gt<n~:rally been lcs\. brachwpmh. Th1' 'l hemc '"a' later extl·mkd to the
\\1thin rl·ct\ rherc •~ not onh commu.tl romtnrc- North of England .mJ proved, though l'ntdl• Jnd
by tlw ,cJeracumam and enrnl\tttl~ .I I!(.le, bur inexact, to bl' apph(ablc, with minor moditllartnm,
o ~ ron,t,tnr armnon through w.tve .H non .md, to much of Europe.
l'llly unportJnt, dcstntetton b, bnnng l)rg.Hmms Vaugh<tn bd1cVl'd tl1.1t the t;mnal wqm·tll'l' he
J rnral-e.ning fi~hc~. These produl c 1 grc..':tt quan- erected rcAl't ted evolutionary lin~:agcs, but 111 f.Kl
1\ nf filll' lOtnlllltllHcd coral ~.md, which till\ up the corah re.tlly Olcur as .1sscmblagc biozmlt'~ Wtdl
hgum1 .1nd i' 1ransported to deepl.'r w.HeN. each of the lllJIIl cydc..·~ bringing m ,\ new tn1gr.111C
fuuna.
Deep·woter corol bonks Rcccntlv Mtt~ hell ( llJH9) has plotted thl· tune
t> r-watl r rorah (ahennar\ pc~ ~uch 1\ Ltphc/w range~ tor 6H rugo'e corJl speCie~ ahram~t ,t,mdard
• Dcu.lnlphy/li,1) mJ} tonn ,trucn1re' drflcnng Viscan ~t,tges. -;howmg that J. now-retinl'd l·or.tl
!Ill dlll\l' of thl•tr '>h.tUow-water coumerp.m~. Some stnt1graph) rs pm,tbk for the Lmn~r C.1rbomlerous.
the-e: Jl•ep-watcr c..·oral~ ,tre ,oiJtM)': orheN arc supplemcntmg other stratigrapluc data ba,ed on
'lll.tland dendrifonn. goniatttc~. c<.modon~ .md toranumtcrids. Ne\\
The.. dl·velopment of thcst· \trucrurc\ (Squires. ~cages b.tscd upon transgressive cyde~ 111 the Lower
''fl'l\ tJkc~ pl.!Cc in four .. tagl·~ (Ftg. 1.17d). At first Carboniferous are now standard. but nn1d1 rehnc-
re 1~ J slllgk colony. Wlwn tlm colony j, JOined mcnt is sttll needed.
dhm rt bt•come\ J thic ket .1 few mc..·trl'' al ross, Coral zon.1l1on 111 the European Devunun. once
wlud1the member.. ma.., be all of the ,,lllll' ~pccics elaborated 111 Wl'.tt dct.ul by W edekmd, ha-. prO\ cd
lid1tli:n.·m 'pecic~. Th1~ newly dl'\'l'lnpc..·d envi- to be overoptrmlsnc
IWt Jttr.llt'> other urgani~m... e.g. ti\hc,, moi-
J and t.nl\tacc.tn~. givmg tt a m·w l'tologtcal Corols os geochronometers
ICtt'r As the. duckt t ~prcad' mJ m.Hurc..'\, ~kell' In wcll-pre'>l'rvl·d '>penmens of m.my ru~o'c Jnd
go
pnd hm JCcumttlate\ from broken .tnd bored coral, sclcranlnt.m c. orals tht· cp1thecal surt~Kc -.Ill!\\\ hne
·2 11Jing a new mbstrattun wh ll'h .tg;un .urr.u:ts growth ndgcs, some 200 per cm. Eat h of the\e rep-
Jrio 1'1' mimak Thi~ ~tage i~ now ,1 coppice and m:1y resent~ a growth mcrcmcm: the fonncr pO\Illl\11 ot
" ites
~~·wralmclrcs .1cross. Eventu.tlly. wtlh 1he accu- d1c rim of the caltcc. These fine growlh ml~c' .He
and
l.tUon of more dcbm and furthn lnr.t! t,rrnwth, ::t often groupl•d 111 pronuncnt bJJ1(-l\ or .lllnul.ttion~
136 Cnidorions
between v.hlth the c.:pHheca I'> consmned. It is nor- Core

mally recugnm:d that the tine growth ndges repre- evolt
'>t'nt daily gnl\\ th I Ill rcnKnh, where:c. the banding ( l>lnni
is monthly ,md thl· broatkr and more widely spaced 'lrurnr
annulauom .m· \"l'Jrh. Wdh (19<>3) first established 'kcleral
111 the ,cJeraetinian ,\Jcml<lllll that the growth ridge'> lml.cd
were Ulllrnal. and \liKe the Recent Lopllelra pcrwsa. CO.Itl''
!Tom Nomq~i.m ljord~. h,J., .'!S growth ridges {pre- :Suc.:h
sumabl~ dlllnl.ll) per h.md. each band corre,ponding gr•ptol
to a lunar tyclt:. the month!} banrlmg hypothe~~' 11d IIH
seom at k.M pm,ible. otl fron
W cJ(, ( 11)63), \\'tukmg on Devoman corals wJth \\ h ert'.l
.uwual r.tthl•r thJII monthly groupmgs, counted the ot tlw 4
number ol wowth ndKe~ per annulation in a num- Se• th.lt
ber of ~penes l'hough the results were limited by urur In
the pre~ervanon or thl' cor.) I~. he concluded, mainly ll.dit~d
from observatiom tHt llcliopllyllullr, Eridoplryllum and 1l1Litlbt•
Fa11osrtcs, thJt thert• were an average of 400 days in Adv
the Oevoni,m ye.u. Thi, ~igure coJTesponded well to ll.llll ly
astronomical estimate~ that the l:.arth ·~ rotation h:t~ 111111 .1111

been slowmg do\\ 11 through odal friction by about 111d le'i
2 ~ per I 00 000 ye.m. Assunung chat the E.1rth 's dutmg

armu;~) nrc un round the ~un was the 'ame length, dlet:tivc
wluch seem' to be f11rl~ well sub,wntiated, there 111 \\ IHc
must haVl' bl·en mon· day-s m the Devonian year, ooperJ

but thev wcr~· ,horrer (~crurron and H1pkin, I 973). "' th
Annual band1ng ~~ nor often found in corals, hav- dt\lmc.:t
mg only been recorded in speclmt•ns that ongin.tlly ~1\'e
Lived 111 ware~ where therl' were seasonal flucrua- llltc !!Tat
nons. lt ts nonnall) e.Ner to work with the Rugosa Figure 5. 18 Bonding on the epitheco of the Middle ~ mg pem
that show the growth ndge' t-.'Touped in monthly rugose coral ?Heliophyllvm sp (x 3) (Copyright pholog\ll the ver
reproduced by courtesy of Or C T Scrutton.)
bands. The'e are more .1bundant .1nd do not have to <r.,gcs. t
have a complt:tt• epltht·ca. Many Recent corals have :ZOIIIUS !
monthly brcl'ding cyt le' dunng which carbonate upon a biologic.1l rather than a physic.1l sy~rem. mosr cx
deposition 1s mh1b1ted, Jnd this may have been the has. however, become mcre,mugly cl~ar that Cl) lwt.. mg
case w 1th the R.ugosa. are not idc,1l tools lt>r tlus, lor wh1lc diumal ban hv ,, fc~
The monthly handed M1ddlc Dcvonian corals is clear enough on all unworn corals, cleJt montl c..luuy,
studicd by Scnmon (I 905; F1g. 5.18) had an average and annual eye.: le., arc harder to detect and lllJ\ r 1tllllds'
of JO.f1 growth ridges per monthly b;~nd, and if be present in many ter.tls (Scn•tton, 197R). T Ci"ICI ICII C
the tigure of 199 (th~· ,1\tronomical estimate tor rhythmitcs (L·bb-tidal <.kpo'>it.'> whme v,1riable lam \11pphes
length of the DevoniJn year) is divided by the thicknes~ encodes .t full spectrum of sernidium lllt'l11bcr
average growth-ridge rounr per band, the result i~ lunar mod.1l p.1l,1eotidal events; Williams, I~ 21101d~ ft
comistentl) 13 offer excellenr potential for geochronometry. e 111d the
Hence it seem' that 13 bands each of about 30.6 cially for the unfo,-.Jhferous Precambnan. R.m.~ tlu: tan
growth ridg~·' were la1d down by these corals in the events may m any case have left a masking recor r-Jis cvid
M1ddlc Dcvoman year. the b'TOWth of the .,kdeton. P enod1c growth fe {tl rh c
The 'coni dOlk' prov1des a corNstent check on in bivalve\ are now better unde['jtood and I're< :unt
estnn.lte' of tht• ,Jowmg of the Earth's rotation based prove to be mon· :.uttable for geochronometr} th:ln<~m
ulmH:tl
filii} mor
Class Anthozoo 137
Corals as colonies: the limits of zoantharian During the Palac:ozoic the ratio of \Ohtary to
evolution colomal Rugma remamed relatively constant, there
domes haw been cktined ,1\ ·~roups ofindJVtdnals being a somewhat higher perccntagt• of -.olitary
'l\lcturally bound ru~etha in v.uymg degrees of corals. FoUowmg the Devonl.ln extinctiOn there
cul.111d phy\lolo~c.tl mtq~r.ttion .•tU genetically was a slightly enhann:d trend toward' t:oloma!tty.
ed by de,cent trom .1 ~mgk· founding indJvtdual' and a fuller mcegrataon of the colon)'. Tabulate
i.lJtc\Jnd Ohver. 1')7.\). corals were all colomll. and m Fnl'c'SIIt'S the mural
\uch colmm.'' (t•.g. tmd.mam, bryozoam and pores suggest enteron connecttom. mdte.ltmg at
tohtc') cm c \.htbH .l w1de r.mge of organization least a degree of colomal mteb't'.ltlon. They were
.111 mtegr:~t1on. At om· extrt·me tht· zoo1d\ budded severely afferced by ~uccc~~1ve extmct1on eptsodes,
Htrorn the pm:nt may bt• lOmpleteh mdependent, however, and the dtve~tty establlshed m the
m•as at the other end of the 'ea le the mdividuals Ordovician was lCtually reduced through tune
rthl' colony lll.l}' bt•nHHC lmked and coordmated (Scrutton, 19!l8).
• th:tt the whole coll)llY tan ti.mcnon as J single Although ccnoad ;tnd phaceloid forms are known
ll !11 some ca'c~ zoo1d-. m.ty become highly ~pe amongst scleracrimam, a generally h1gher levd of
IZed, promming dJVI\ion of labour amongst the integration i~ apparent m the preponderance of
<1Jtbers of the colony. meandroid and coeno~tcoid types. Me.1ndroid gen-
Advantage~ of the wlonial st,1tc pre~umably are en. have a polystome ~yste m in which hundreds of
unly connected with h"~'t.,ltt•r dlictency 111 protec- mouths with their :~dpcent tentacle~ connect m each
n.md \tJL'lllity ,1\ well .1~ 111 n:production, feeding ·valley' with a single elongated enteron. In coenos-
Jd r~\plrati o n. Thm .1 tolon~ of a,t·xu.llly repro- teal Scleractini.t the enterons of adJacent polyps are
~ng mJtvidu,tl' tan \OOn dcvl'lot' .1 l.lrge .md aho all connecred, and 'ikcleton buildmg hJs become
ett1ve b1omJ\\, wHh .1 'trong .1nd \t,lble ~kekton a cooperative venmre. Bur even the mmt Jdvanced
wluch rhere ts the polt•nu.ll fi>r inteb'Tation .md sderactinians, with the pm~1ble exception of a very
•peranon h~·t\H't'n ztlotth fe·w rare genen, do not exh1b1t polymorphJ,m.
In the nttocur,tl' ( B.l}'l'r. I <>73) there are two Despite the fact that rugose and tabul.lte corals
et tuntrional typt•,: (I) thme wh1~ h 'ecrctc a were able ro make 'orne he.1dway tow.lrd, .111 mte-
t\'e 'der.lttllll.ln-hke 'kdetun wtth \ef\ little gratcd colonial sy~tem, rhc> \\ere generally les~ suc-
e~:uuon, .1nd (.2) thow (the\ J\t majonty). includ- n·~,ful in this than the Sclcracttma. But thetr
Oil pcnnatulateam, m whu:h mtcgration vane-. from comparatJve failure as reefbuJldcn was probably due
ph \ery· ~unple, chrou~h numerou~ intem1ediare uot so much to this a\ eo cert:un other features of
~ to ver; complex tulonte' with the variom their orgamzation. The Rugo'<l and to ~ornc extent
1ds spen.1hzcd tor dttlen:nt fum:tton~. ln the the TJbulata had slow-growing and rJther ~olid and
t extreme CJ'Ie~ not only arc 1nterzooidal canals heavy skcll'tom in which :1 great deal of calcium car-
~mg the t ntcrum ret.uncd, \o that food captured bonate was deposited in internal stmcnues. They
a few zuotd~ t...1Il bt· shart:cl by the rest of the lacked an edge zone and could not :~ttach themselves
lony, but also there .uc 'penalized 'siphono- firmly to the substratum. Tt i'> not known whether
"uls' org.1m1ed for pumpwg water through the Palaeuzoic corals had zoox:.nthcllae in their tissue; if
n(nchymal caJl.lls. to cmurc that adequate they did not, one of the b'TC:lt adv:mtages possessed
lll:l phe~ of oxygen.Ht'd \\ .lter tor rc~ptration reach all by reef-building ~clcractiniam was denied to them,
to lbt·rs of the tolony. Suc.:h polymorphic and this would have affected thc1r metabolic effi-
8~) 1d> funcnon together but c:annot function apart, ciency, their ability to get rid of wa~te mJtenal and
pr- th~ colony act\ Js ,\ \upenndividual'. Despite other factors.
om ~a that dm level uf unegrauon 111 the octoco- However, the \trong, yet light .111d porous. stable,
ii.n r\1dently ongm:ncd vel) early 111 their lustory fast-growing and ar rht: ~amc time mcrcasmgly inte-
Jre' !he mtcrpretanon of ClumlltldlsCIIs as a grated skeletons of the sclcractimam permitted the
mmbrian pennatulacean ,., correct), the zoan- development of the coral reef envtronment, so lay-
I.llll haw never 6.11ly reached the ~ame le,•ds of mg down the framework for \Omt' of the most com-
~mce thetr £aoids are nor plex and productive of all ccosy,ccms that h.1vc ever
evolved.
138 Cnidarians
Minor orders K
In n·tcnt \'LJr' Jll,lll\ \:or.1llim: · taxa have bt·en
ckscnbcd trom tht: <. .unhn.1n and intomull} placed
in tht ( or.llolllorph.J <.)f tht•,c. two orders ,1re tme
Ztl.lllth:man,, 'l ,tbul.lromd.l :md Cothonionuda. 3
Neither uf rhc~c: .1ppt•,tr' to be: .mct:\tral to .my post-
( ambn.111 p-oup. 1 hc:'c .md othn invatebr.ltc~ and
uther mmor onk·f' .trt• d1scu"ed in full by Scnmon
(I 'J'J7b).
c 2
Order Tabuloconida
'],,J,u/,ln•llll.' (I <. .1111 ) 1\ .1 \llht.try ur dendro1d coral. l•l Heterophylli•
\\ 1th largt· (>I(, mm) cor.llhtc:s w·hich arc u'u..1lly
.ht:pt.ltc bur With t'\ ttknt ubul.1t:. Alt>t>rt>II'IJWm from
the Lm\'l'r C.11nhri.m nt' Amtnha (~or.mf' and
S,\V,lft'\C, l'l')'i) 1\ .1 ccnoid torm Wtth ~epta and tab-
u be, ,lJH.l 4 l'r!111'iJltlfc1 111.1y ,1lm bdong here. None of
thc~c have mur.tl pnrc,, the1r mode of growth is
unhh· th.tt nl r.thul.Ht''· .tnd they are cbssificd scpa-
r.ttcl}.
Order Cothonioniido (Fig. 5.19b)

flu-; monmpt't lfll \UbdJ" ts known nnl} from the
F'I(Jvre 5.
<':l.rly Muldk l 'unbri.m o( Ne\\ ">outh W Jles where W(;redel
mdJvJdu.l), lft' 1hund.mt
I he t t)r.lllitl.'' o( G>t/11•11i••11 ;u-e ~null and conical lb) Cothonion
.md up w I 0 mm m dt.uncter. Tht·y ha\'c "' eak
epn mtenull) hut no hunzontJI ckmcnt\. There •1ul ( I
ptll
•~. ho\\'t'Vcr.•111 operculum or lid tor each corallitc
1\,1/m.ll
upon tht undt·Ndc ol wlm-h .1re \trong btradial
l11lll·ll
sept.l Jdl .1nd Jt·U ( I CJ7(•). who fiN dcscribl'd these
c:or.1h, 'illggc,tcd th.tt till~)' •mght bt• allied to the " lOill
lt I\
Rugm.1, but thev .tre mmt prob.1bly a quite inde-
lr.JI to I
pcndcm group. 11nw ckv,1tcd to the rank of order.
( lrclm•t
tll;lt /\.11
Order Heterocorollia (Fig. 5. 19o)
I hl' hctt'Tllll>l".lls .tre .1 snull group of elongated,
'kdliOI
p1pe-likc lOt,d~. umtined to the Carboniferous of
( '>r.tlli
111~ 1 S\11
r.uropt' .111d A,i,1 .md very ~hort-livcd. They have
llm
'iCpta .md t.lbuiJe but often lack .1n ep1theca. Thc1r
IH rt-h
pattern of \l.'pl.tl 111\l'rllOn ts unusuJl and charaC[eris-
nc. l.,omt ,lut l mtlllt'~ behcve that they led an epi-
md 1'1
n1gos 111
pbnkwmt hti:, .Ht.tched tn tloating st'Jwceds.
\H:n.: (
lnCIIllll
rh~ dur
Orders Kilbuchophyllida (Fig. 5. 19c) ond
ph m m
Numidiophyllida F1gvre 5. 19 Representatives of mmor orders: (a) Heletc#. hort-h•
A 'mgle l!tllll' ol ,dcr.JCtt 111.111-hke coral h:1s recently (" 5), (b) Cothomon (M. Cam, 2) (c) odoleKenl Ki/b,drf.
y
(>< 3) ((a) Redrawn from Hill, 1962, (b) redrawn from Scr. pcndt'lll
hcl.'ll dc,ntbt•d from 111.1"-tlm" Jepo,tb 111 the
Uppt·r Ordu\'it"i.m ot ':>nutht·m Scotland (~cru tton 1979, (c) redrawn from ScruHon and Clarkson, 1990.) gJlll \11
Bibliography 139
Prec.
V
E 1.m:pho T" [ y-•
Palaeozoic
P
Tr
-
:
•ne. Cothomonnda
Tabulaconida I
? - - Heterocorallia
I
(50 genera
Zoanthiniari a
Actiniaria/Corallimorph~r~__ _
r
uc~ophyll ··-:
~5 20 Relationships of the various groups of zoantharion corals. The ranges of the uncolciRed sea anemones from which they
O!rederived (Zoanthorinario. Actinorio, Corollimorphorio) ore shown by dashed lines. (Simplified from Scrutton, 1997b.)
dC!Jrkc;on. I 'l'Jl ). In H' .tppt'Jr.mcc, p.urem of

put 1mcrtion .tnd \epr.tl microc;m1cturc,
h,,phyl/i,, " rcnurk.tbl~ mnil.tr to recent sclcr- Books, treatises and syrnposia
uruan.,, :mJ 1' 110\\ known to h,l\'t' been .1ragoniric
compo,ltllln. Oennen, I ( 1971) /7u· ( ;,,.,u IJ.m11 1 Rn:f. l .tmdownc. Dl'e
h11 not hkd~ th.tt tlm ~t'JHI\ \\,1\ .tctu.tlly anccc;- Why Wc~t.
Borradatlc, L., Pom, f-.A., f~a,th.un. L I .~ .•1nd S.nmde!"'i,
!O l1vmg sc:ler.tt tltll.lm on .tt count of the
J.T. (1956) '11u· lm•t•rtcbrtlltl C.u11bndge Umvers1ty
.lol'ici.m Tn.t,,ll tum· g.tp. Wh.tt 1\ probable 1s
Prc~s. C,unbndgc.
11 Klibufiii'JIItylha repre'L'nt' .m t'.trhcr '.tt£cmpt at
Camcron , A. M. et nl. (ld~) (1'J74) J>mcl't'drn.~~ tij'thc Sm111d
dctonmtlon' by thL' \;lmt• !:,'TO Up of sc.1 .m cm ones Intematioual SympMIIIIII ••11 Cmc1/ l~afs, Vols I .llld :2.
Coralhmorpl1.1ri.t) ch.tt g.tve nw to true sckrJctini- Grea t Uamcr R eef Comn1Hrcc, Urisb.111e.
' \!lmr .2.20 M,t l.ltl'f (Ftg. :1.20). Hill, D. ( 1937) A Mtlllt~I/Ytlph of t/11 Carlmuifi·rciiiS Rr(I/11Se
Th11 cJrly exampk of skekt.tl .Jtqui~ition was Ctlmls (![Swtlmrd. t>.tlilt'Ontngr.iplmo~l Soucry, I ondon.
)it-ltwd .md very Im .tll7t·d. 111 Southern Scotland (T.1xonomy .llld Lnr.d LOillllllllllt~ \lrtll"tllrt•)
...J North em Ird.md Thl' nmtcmporaneous MaxwdJ, W.C.II. (I%H) 1ltlas ·~I tire (,real Bc~mcr Rct:l
Jt,'Wm, wh1ch ongm.lt<.•d .H .1bour thL' same time, Elsc\ncr, Amstcrd.lm. (lllustrarcd .tti.Js)
e deriH·d from .1 ditTncnt group of sea Moon·. R. C (eel.) ( Jl)'j(,) 'linlli.or ''" lm·crtd>mtr
P,l/m·orlloft•~y. Part F. C.>t·lt 1/tcr.rf<l Gcolohri.c.tl Sool'ty of
!One' (Zoo.mrhin1.tri.1), bt·c.tmt' dominant for
America and Umvt'Nt)' of K.lll'-l' Prc". LJwrl'ncc. K.m.
duranon of thl' P.tl.ll'otoit•, .tnd tht· corallimor- Musc,mne, L. .llld ll•nhofl. 11 M (I'J7 ~) Ct>dmttmlc
'c:a anemom·, thd not, other than the Br,,lt~I!Y· Ac,1tlcmil Prl'\\, Nl'W Yl)rk. (E:\ccllcnt compi-
Pennian \'ut~uc/i,lpilyl/um, which inde- lation)
nt!} dL·\'doped 111 tr.1gonirir ~kdeton, calcify Shcppard. <.R .C ( I'JIB) t .\llfm,,J I /isttll')' t1{ the C ••rt~/
OlD unnl altc:r dll' rugo,.lm lud becotnl' cxrinct. Re~/. UIJndtord. Pnnlc.
140 Cnidarians
Stel, .J.II. ( I 979) Studrn ''" tire Palacc•bi<JhWJ' cf Frwositids ~ion 111 the northwest of England. Quarrn/)'./••llrtt.J/(Ij I 1\\
Stabo. Groningcn. Gcolo.!flWI SvciCt)' of L.mdo11 68, -1-l'J-S:,Y(, (lnru
Ul~
Tt:ichcn. C. (ed.) (1 CJH I) Trt'Jttsc• 1111 bwertdnc1U' using cor.ll .1nd brachtopod f.lUna) tlu
P.r/ewllc•/,~1{)'. P.trt (Supplt:mcnt I), RII,I!•'S•l .uul Gill. G.A .•1nd Co.!tl!'>, A.G ( 1477) Mobthty, gmwih mdl
T.r/tu/o:~tcr Geolu~•c ll Socicl) of America and tems and subsrrate m 'omc fu~'\JJ and rccl'nt c \ ))
Uruvt:l"'ilt> of K.lmd~ Prc''· Lawrence:. K.m .. Lctlwia 10, 119-34. ·11
Vcron, J .LN. (19Rft) Cc1ra/s of Ar~sttolra 1111d the Jndo- Gorcau, T.F. ~nd Yongc, C.M. (1968) Cor.tl comruu M a~
Pc1{Jfh Angus .md ll..nbcrtson, Nurth Ryde, NSW, on muddy ~.'lnd. Nature', 217, 421 .3. (Soft-l,.;,n .7m
Aumaha, 044 pp. (Superbly illumatcd \\.'ith colour hab1t.1t) In(
photo' .md dmnbuuon m:.ps of all ~pc:ete-~) Goreau, T F. Gore:1u, N.l. and Gorl'.lU, T.J lli !Jrll
Vcron. J.E.N. (II.JIJ'i) C.>rals ill 'ip,ue anti Ttmc CorJls ;md coral rct:h Ylcmific .1mmt.tll 241 l I
Bicw·o~rnplry artd c.,,,[,uou of tlu Sdrmrt111ia. Comell 110-21 Uri
Univcr.ity Prl'ss, lthJc.L Jamicson, .I::.R. (1971) 1'alcoccology of Dl'Vlllll.ln rrr~ /\lciiH'
Wclb, S. (cd.) (ltJHH) c,w/ Re~(s t!f tlrr H'orld (3 vuh). western Canada. p,,,n•rr/iu,l!s 4 tlu· l\'cnrlr l1 ut
UNI::P IUCN (Urmed N,ltlom Em'lronmcnt P.tlcomoh~ll<ll Corwwtfllll, Cluca~o. 1<1(1') U \ID;
Progrmmu:. lntcnution.U Uruon t'l)r Conservaoon of I 300-40 (Reef environment->) MHch
Nature and Natural R esources). Jell, J .S. ( I %tJ) Septal mtcromucture and cla"lticAtlon n1~
the Phllhp~~straeidal'. in Stlati,ilraplt)' .md l'ct/cm'lf' ( rf'(
(ed. K.S. W . Campbdl), Au~trilian Nanonal Untw Neunl
Individual pape rs and othe r references Pre~s. Canberra. pp 50 73 (Com po~ttll•u of 1k, Cflr.
elements a.nd c hcmmry) Ncum
Babcock, l ( 1991) The emgma of conul.lmd affirutu:,, 111 Jell, P A. .md Jell, J.S (1976) Early M1ddlc Cam P1l:.
Tlrr Far!)' Er•,!lrlllc•ll I'{ .\lrta:oa ami tlrr Sig~tU'irartu• •!f corals from westcm New South W.1le~. Althrn Nudds
Prctltlc•mcwr Taxa (cd~ A.M. Simonett:l .lnd S. Conw<IY 181-95. (The oldc't known cor.u~) luh\
Morn,), CUP, Cambridge, pp. 133-43. ]ern·. F. (1')IJJ) Conulari1d n1icrofossih from lht S1l •n Jlll-
Baycr, f- M ( 1973) Cololllal o rg;~ru~Jtion in octocorah. 111 Lower v.~by bccb of Gocl.md, Swcdl'n l'.tlo!oL ( lhver,
4111111<11 (.,,fouJrs (eds R. Boardm.m, A. Cheetham .md 36, 403 24. men
W.J Ol.tver). Oo\\den. Hutchul\on and Ro~~. Jcrrc. F. (ICJ94) Anatomy Jnd phylogcm·Ul '•gn.IIOOICt (lom
5rroud,burg. Per111., pp. 69-q3. Et>nmu/,zn,, lt1mlata {Wun.m I R95), cl conul.lntd 1.. hvn,
Bengt,on, ~- (liJH 1) ..trrmtosrlla, a SrJunJn alcyon:tcenn the Silun.m of Gutland. Letlwia 27. ':17-1111 o( D
octoror.1l Joutll<ll (){ Paleolltology 55, 2H 1-9-l. Jull, R. K. (1965) Corallum increase 111 I.JI/1,,,,, \c,:=;-
Carrudu:r., R.G. (l lJ Ill) On the cvolutmn of Zaphrt'lllls Palatwttoh•f!)' 8 , 20-l-2'i. (Vannm method' ofgro11 l lh\CI,
dd,tJfllllt:t rn Lower Cuboniit:rou~ mncs. Qrwrtaly Kato. M (1963) Fine ,kcJctal structure~ m Rl:j; or.1l•
Jourucr/ .if rhr Grt~lc~i!tl·11 S<~<iet) ·~f Llt~tdo" 66, 52.} '\6 Jcumal <!f the Faculty of C.:..ttl/(1' <!f I l.>kk,n.f,, U, (fan
(CI:h,IC. though now dub10u~. evoluoonary ~mdy) Series 4 11 , 571-630. (Mtrrmrrucmre) lllrH·t,
Cuatc,, A.C. and Ja ck~on, J B.C. (ltJ~7) Clonal growth, Klovan, J E. (19M) Fanes Jll.l.ly~i_~ of the Rnh\.tkr Jl P.,IJl'<
.>lgJl 'ymbimis. .111d reef fornlJtion by n1ml~. Complex, Alberta, C.lllada. Bullemt ••f C• Ill,,,,
P.rlat'tlluoft•l[y 13 , '\(,'\ 7H. Petrolt:um Gcolol/y 1 2 , 1- 1Otl. (Distribution uf,
Coate,, A G and <Jhver. W.A. (l'J7J) Coloniahty in
r he r
within ,\ n·cl} pp. I(
zo.111thanan corah, m .1uima/ Ctllomc..< (ed\ R. Krebs, W ( 1974) Devon1an t.arbonatc compk·xc( of I'Liyl<•rd
Hoardmm, A. Chet"th.un and W.J. Ohver), Dowden, tral Eumpe. in Rtr6 '" Timr a11d .\pll•< (t J ( :llllll
llmchmson and Rm\, Stroudsburg, Pcnn ., pp. 3-217. Laporu:). Society of Economic Mm!!r~lngilt'> ,,J Jl.•fr
(lntcgrJtion of col omc~). Pakontolobrist'i, Tub.t. Okl.. pp IS~ utdlwr!
C0pper, P. ( 1974) Structure Jnd development of early (Di~tnbunon of cor.ili wnh111 ree~) lilt' n:
Palaeo1oic ree&. 111 l'roradtiiJl> tl{ tht ')erclllti flllenrattclllll ladd. H .S ( 1971) Ex:1\0ng rcc~: geologwal ali J~.umal
5)''"1'05111111 cn C1r.1/ Ht•t·{s. (ed~ A M C:uneron re a/.), Proa:etllll.i!S of rite ,~>,:crtlt 'imrnc.w Pah·olltcllc!l!)' C~t:l ITII<fll
GrcJt B.1mcr Reef CL1111111lttee, Umbane, pp. 365-l-16. Chtcagn, 1469 U), pp. 1273-1300. (\.o()d ~Uit \1 offin 1
(Ch.mgc~ in reef 'lniCtltre from mCiplcnt to maturity) and b1bliobrraphy) Wc-ulo
Cnpper. P. ( l ll85) r~l)\ili~cd pol)'P' m 430-Myr-old Lang, W.D. (1923) Trend' in Bnt~Sh ( Jrhonif 17.3-2
Fat•••~rtt·:; corab..\.,1/llrl' 316, 142-4 coral~. Prwcedill,il> of tltt• Gcofo~rc.tl 4l>tlolJtl cut1in 1
DahJ. A .. Macmryrc. I G. md Antoruu,, A. (1974) A 120-36. (An early analy'•' of cvolution.uv tren;h; \tnll:tu
comp.tr.mve 'urvcy of (Oral reef rc~earch sJtc:s . ICcl/1 ally ofl.tnmcd v.Uuc) IJJrwu
R<'Wctrdr Bllllcmr 172, 37 I 20. Lc:e, J. and Noble, l.P.A (I <JlJO) Colony d~:velo 11-JR.
Garwood,. E.J. ( 1C} l.\) rhe lowe r C.lrbonifcrous SllCCl'~- and form.1tion in halysltld cor:1b. L:lhma 23, 17'l-' s~, \tttun ,
Bibliography 141
le1m. JB. md Pritc, \\ S ( I'J75) fl'ed111g mechanisms wowth. l'alat'ollmf,~'IY 7 , 552 ·H. (.\1onthly b.mdmg Jnd
md ftcdm~ ~rratq:1c, of Atl.mtJ< reef t·orJ.h. jo>llmcJ/ of the length of the Dcvoman yt·M)
~ 7A~,,f,>cu,lf ,s,,,my ,,jJ ..,,.,ft,, 176. 'i~7-H Scmrron, C .T ( 1975) ll ydrcud-.t·rpulitl Wlllluo'" m rh~:
~J,trom, M. (197~) An tKlOtor,J fmm thc l ower Me~ozoK and Tertury. Pa/<tt<>lftt•/c•l!)' 18, :!25-7-1 .
OrdO\KJn ol '" t•den Cn•lt•t~rul t't P.tl.lro>lltt>l•~l!'ca 12, (P.1rsJmo11icl and Protrrlc>p/u/.1)
~l-52 (l )\dc:q knm1 n) Scrutton, C T. (I Y77) Reef fll'll'' 111 the Devon1an 0f
lcrgnc-r. H .111d Sd1c-cr, G (I 1174) Tht· ph)·,iographic eastt·m South Devon. Enghnd \lwrorrc·s Ju Bmt·mr !le
11 tonlnun .md "'''lngtt.ll tomliuun~ tlf 1vme ~l1Uth Rt.'CIIrrclrrs Gt'oJ.~I/''1"'~ d \lrmbt 89, 12'i-JS (F.mn.J
lnd13n and Ccvlon tor.ll rl'cf... Pt.~(t-nlill.l!> of tilt Sec<1mf \',m.mon with f.Kil•,).
l"t.-m,Jtltlf<.JI "Y"'I'•'HWII ''" Ct1r.JI R.-r{l, (ed~ A .M . Scrunon, C T ( IIJ7H) PenodJC growth tc.uure~ m fi>ml
Carneron r'l ,,/./ Grl'.ll Bamcr Recf ComtnJttee, organism.\ and the len~:.rrh of thl' dJy ~nd mnnrh, 111
Bmb.me, pp. 3-.311. TiJa/J 'riaion allclrltr Eo1rtlt '.; Ro•l·lll•'" klh P. 11mt.ht and
lll \l;:n~-!:dw~rd,, 11. .md lla1mc. J. (I HSO) A monograph J. Sundcm1ann). Spnnger-Verl.1g Bcrlm. pp 15-1-96.
ot the Bnmh to"1l , or.1b. P.rlal't•III•!S{'<1plutal Society (Corals les\ v.1huble dun biv.1lves)
""l'· .\f,.,,,~raplr. pp. 1 15f>. Scrmton. C. T. (I 979) J::,lrl) fos~t.l cmdJ.tlam, 111 Tlu:
\htthdl, M. ( llJH'J) i3H1\tr.lll~rtphy of V1sean (Dmantian) Ori._~i11s <![ flfajt>r flll'frtl'lrr.llr Crl!IIJ1> (ed. M.R. H ouse).
ni[:O'<:' coral fllln~\ 111 Bm.nn . l'rc•m·dil{(/> o.f tire ~ ·orkslriTT! Systcm~nn AN)CJ.ltlon Spt'CI~l Volume No. 12.
f..et>/t~~ICol/
So.ifl)' 47 . 21'\· 17. Academic Prc~~. Londo n, pp. I (d -207.
l-J~unLill , D (1984) Ori!-(111 .Uid ~:a rlv cvolutmn of ru~ose Scrurron, C.T. ( IY83) N<:w oiT,cHls\oci:lled srrucrure~ in
orah P,rfcrllf!Wrc1JIIIiCt! lm cticclll<l 54, I lll-26. ~ome Carbonrfcrous rugo~l' cor.tk J..ctlt11ia 16, 12l)-44.
'lurmn, 13. (1 tJKH) Somt· .1~p~:cts of life 'rrateg~es of Early (Remarkable growth \truu urc~)
'\.Ill Pa!Jrozort ru~mc: COtlb. L:tltctia 21 , 97 114. Scrurron, C.T . (ll)84) Onw11 Jlld e.trly evoluuon of rab-
I l, udJ,, J.R. (I 1JX I) An lllU\ll .n~:d key to the Briti~h ulate corah. Pa/m•tllllc~l!rctfllllr.J 1111wrim11a 54, 1111-HI.
thomorud ro~ l~. 1, ttl llalcll't>llfttltll/l(cl Ptt!mtJ(cl 22. ~rmtton, C.T. ( llJKfl) Cmd.1n.1. 111 tt/"' ~~r ltwmehmtt'
r un • l-l04 Marr<!f<>ssil.~ (cd J W Murr.1y). lnnt-.'1 11.111, I IJ rlm~. pp .
,Jog)' Ytr, W.ll. (197hJ) Sonw ~'pe(t\ llf ~olony dcvelup- 11-46.
mrnt 111 .-orak ,\ /,m,•m <'{ t/r, J>Jlrot~tolt~I/'C·II .\.-•aety 2 Scmtron C.l . ( I 1JH7) A rcncv,; of favo~1t1d JfflllltJes.
•t• of lfcl•r•••II•!J PJft'cllll•''•l'S}' 42 "uppl.). IC>-H. P.11111'!llltology 30 . 4H'i-'J2
rom rrr, W.A . ( 197hb) Pr~'ldc:ml.ll .tddre" biogeography St.·mtton, C T . ( 19HH) Pattern' uf o.ttnttllln •md \Url'lval
-:l' Oe\om.m ni!!;O\t' t:or.\k /•'11nr.1l ,,( Pail'.:wtolol!)' 52, m Palaeozmc coral,, m l::.\llltfflt>/1 .wd tin• 1-o.<.<JI Rt'.-,>rd
~73 . (Cor.ll fnm.1l provmet·•) (ed. G.P. larwood). \y,tem.ltrc~ A~\OCJJtlon Speoal
''"'"
lth) :tr, \V, A. ( I1JXII) The rd.ltll>ll\hlp of rh~ ,cJcnctiman Volume No. 3~. pp. C.'i-HfJ
i;\'~ I ;;nil tll the m~o\e corJh. J'.J/e,>lllt>l,~y 6 , 14C:r60. Scmnon, C.T. ( I'JH1J) Ontog..·nv .tnd J'tngcny m
•rmry f.!~ our\ on~n of 'dcr.ICtmiJn' from \eJ anemones) .-111/t>pora .md Its ~lgrllficance Illu,tr.ned by .1 new non-
.:r, \I, A. ( I'No) <. )ngim and relanomh1ps of encrumng ~pl'Cle~ fmm the I )evom.m of southwest
Rc.:f P.d.il'OZUlt coral gruup\ .md th~: on gm' of Sderaccinia, England. L.ctlwrn 23 , (, 1 75.
I<Jt/rdll 10 Plllt\>l•r(l/t>i/)1 t1111f lJHJI.>!!y <?( C,•rals (ed. C.D. Stanley). Scn1rton, C.T. (1997a) Growth \lrJtegi~:s .md colmual
.::or.1h fhr PJleontologtcal Son cry P.lpt:f\ 1. L.1wrence, Kan., tom1 m tabulate cor~!~ &•ltt111 I<<'•'' 'l•••u·c/,uf f:'sr•iiiC•Ia de
1 1117-34 Historic1 Naturnl (.Srcrion Grt•lc~l!id) 9 1. 177-8\1
f ccn- ~ord. P.E. (l'lHO) Dev1.111iaJr 'Gre.lt Barrier Reef of Scrutton, C. T. ( 19'J7b) The Pdl.tcowi~ cvr.1ls. J. Origim
. L.f Clnnlng U.Nn, Wc:,wrn Au\trJIJ.I. Amt•nrau .1ssociatio11 and rclatiOil\hlp~. rr.•rctdiii,~S v( the ' ,,bltm: Geoltll/1(•11
.s :wd (Prtrolm111 l.cr/rl,l/itts l!ullrtill 64, H14- 10. Soriet)l 5 l , 177-20H .
'l-20K 1 dbc1g. P A (I Y7'i) 13rymo.Jn di.1g~: nesi\: be.1ring on Scmrton, C.T. and C larbon. E.N K . (l 1l 1JI) Ant'\\ sclc-
th~ n.llurc of lhc on~ n.ll , kclcton of nibrose co raJs. ractiJJim- like coral from tht· l)rdoviciJn t~f the
tspe~l~ ~miJ/ of Ptllemrto>lc>I!Y 49, 5H7-606 (Detailed Southern Upbnd~ . Scotland. p,,/.,c•c>l/tolt~I!Y 34. 179- 'J4.
llt lllft11 •rmnucrur.ll ~ludy wnh good bihliogtoJphy) ScrUlton, C.T . ,1nd Hipkin, R .G (1971) Loug-tcml
m nul) ll'm T. P. (1971) lht: tOndmon' of ~owlh of the change~ in rhc rotmon ratt• of thl' E.uth. Earth S(lm.-e
WcnloLk red\ n( ~hrup~hl!'e St•drmeutoltli/Y 17, Revrn1• 9 , 259--74. (P.1roallv ha,cd on cor.ll b.mdmg)
,ifcrtHI~ J-!1':1 S!Jter, I.L. (\Yl\7) A monograp\1 ol Bntl\h Conul.m.u:.
··111 34 n.TP.and Dl';nu,.J .E. ( l 'lH.l) The thsmbution and Palartmt•!I/T<Ifllllrlll '\ont't)' ·""'''~1/Ttlplr, pp. 1-411.
Is; ;H m- ::ucturt of cord) n:ct\; nut· hundred year.. '111ce Soro~uf, J.E. (1971) MIUO\trunurt m tht· t•xmkdcton of
:M\1n. Br11lt~~"''' )<•unr.t/ of tlrt• Lllmtall .S.>cirry 20, some Rugm.l (Coelentcrara) . .Joum,J/ <!( Pctft't>lllttlt!~l' 45 .
1opmem 1-38. Red\ ~nJ pl.1te tel'tomn) 2}--32. (St:an nm~ elt·llrun nucrn~raph')
'9-94 r.nron. C T. (I'}(,::;) Pl·nudltlt\ m Devonian coral Sorauf. J.E. ( 1972) Skelt•tal nucro,tnl~ turt• and truo:ro-
142 Cnidarians
~rchttccrun· m \da.tCnnt,t (C.ul'lcnta.ILt). Palat'OIIIt•I·~!?Y and ecology \\ lth nKrt•asm~ nurunry)

15, HH-107 ('il mntng t•krtrnn mterot,rraphs) Wang, H.C. (1950) A revt\1011 uf tht L.o.mtham H.ub'Oil
Soraut . .J.I:: ..tnJ (\.w.trt•,e, M (I '145) A Lowc:r Cambrian m rhc h~ht of th\otr minutt' sl...dttal ~mtwuc
coral trom ~omh AtNrtlt.1 PJI.ti'otrttll,~y 38, 757-70. Plliltl.ltlJIIII(tll rr.m.<tlmt•IIS <?f tilt Rti)•J/ '>t>(lrly B 234
Squtre<; () T ( I'JI•4) h>"il cor.1l thicket' m Warr.~npa. 175 246 (CI.t'>~ll' -.tudy, now r.1tht;"r SUJWT\Cded)
Ne\\ L~·.tl.ind )•'"""'/ o/ f>J{r,>ul<>l<~llY 38, lJII+-15. Well\, J.W (1'1'\7) 1ndn.idu.tl \",m ttam m rh<" n•!l
(Devdopmcm ol tlud.. t•£\. ~opptt:C) .md bank\) coral 'pecac' I /t'llolpl•ytlwu /ll'//1. Pt~/.u·o•lllt•lt,t!i.J Amrr
Scank·y, )),(, and \rOnncr, W. (l'JHJ) The first fo)sil 2. 1-22 (f nvirollllll'Ot.ll f.IC[()I"' tn Vln.ltlOII)
cccnophorc lrnm rill' Lower J>cvom.m of Wc't Wclh. J \V ( 1957) Annm.tted htbltC'I~Jphv: ror.lls
Gtmwn\ '\',umt 303. <;I H-20. Trc.wsr <111 .\lanm E ,,f.~} am/ J>.,/,Jt'<><'u•l~~>·· Vol ~
Steam. ( W .tnd Rtdmg. R (1'>75) Flmm ofche hydro- J. l ledgperh). Munmr\ of the (,colog~cal Snt1~
zoan Milt.po•r,, o)ll .t Rcn•nt l'Or.J.l rcc[ Lrrhma 6. America No (,7, Grolu~c.tl ~onerv of An :n
!87-200 (liVIIItl, milkpMIIll'\) Ln'llrence, K:lll . pp. 77 3 K2. (I m .1lu.thle \Ourcr 11 rl:
Stnddud, D C (I %9) holu~y .tm! morphology of Well~. J W ( 1%j) Cor.tl b'l'O\\th .mJ ~cochronumctn
Rct cnr cor.1l rt•ds u,,,[,~~'"" Rn•itw.\ 44, 433-98. ?'olclfllr<' 197, 94H-50. (Annu.1l h.mdmg Jnd the '1011
(V.1Iu.Jble rdnt'nrt• worl. wtth btbhof,rraphy) of the E.1rch 's wt.tttou) Bn:o7oa
Taylor, j.D. ( liJ<tH) C:or.tl rt•cf .md ,l)~oci.ttc:d invertebrate W cndt, J. ( I'J'JO) The liNt .lragonitil ru~o~e corJJ }twm
l aHI to
communttws, tllJ.tnly molhtstan, around MJJ1e, •!/ Palrc•ut;•/tJf!Y 64 ..BS 40.
t,>rottps f.
Seychellr~. IJ/,i/,t.(llflhiml 'I i·,,s,mrotts ~f r/lc Royal Society Williauts, G.E. ( llJH9) Late: Prct'<llllbri.m tid.ll rhythnu
!ll(lll Ill
B 293, 129-2011. (Cl>ral/nmllmtan ecology) 111 South Amtraha .tnd the hi\tory of the Earth\ rotl-
van lren, 11 ( I'NI) An.tromy, p.1tterm ofoccurrency and tion. jmm1111 4 tlw Crttltwiml Soncty of Loudt•ll Uh ltrd.ty; .11
nantrt• ol thl" , nnnul.mtd s< hott. PalacOIICClft1~Y 3 4 , 97 Ill. olfl' kll\.11
439-54 W1hon, J. (I '175) !'he dmriburinn nl tht· LollllllliTil

van hen, 11. , FH£1.c, J A .md Cux. R.S (19%) Cll1J'<lp/ly/lr,1 '""''"' \ .md B un th~; ~cuttt\h lOIItlJ ~h lct0111
Problcm;mcal to"tl tllldJnan' from tht.' uppc:r 'heir Jcmnwl 4 rlac \ltll'illt' Httl/cJ~tml oU.I•'"·'"'''' n•lc•ny h
OrdtWtCllll ,,f tht• north-< entr.tl U'IA. Pa{,,,.,,,tol~y 39. L'mtfll Kill(!dt>rlr 55 , () 11-::!'1 (Pren~e documcnr~tl All h'l
1037 64. Wtl~on, Jl3. (I <J76) Att.ll hmull of tlw corJI C.IT)'<' Ul Lht: Ill
Vauglun, A ( l'ltl5) T ht· p tl.lt'ontolu~(:KJI 'equence in the >mitllli ~ llld B m tubt'\ of thl' polvch:tt:tc V.
CarhumfC.•rC'Iu' ltlll<'~hm<' ot' the Hnstol are.l. Qunrraly •lnffllhl (Mulll·r) .111d othl'r 'uhsrratt">. }o•llmJI o(
J•••mw/,•Jr/,, C';,,,J,~'"'/ ,,,,itt)' ,,f L,J,,, 61. 181-J07. .\lanm Bit•lt•!!U<II Am'<l.lllo>ll ·~I tlu Uuir.·cl l'm~.lm ·
(t'lt\\ll \tlld} U\1111,>; , nral br.t,luupoJ zun.ttton) 291-Jt)J (l>ocumcnr.tnon)
Vo,hurgh. F. (I '11\2) .-lt''•'P•''·' ,,-r,,u/,,,.,: ~tntcture. Yon~c. C.M (I%J) I hl' bmln~·y of coral
nu•, h.mtro., .llld ,·,·nln!.'\' nt .1 rrt•t c,1r.1l. p,,,,,ycfm~?s '!f rht· .-J.,J,,mu.• 111 .$,mm· Hic•ft~)'· Vnl I. (ed F ~
Rt•y.t/ S<Vttl)' H 2 14. -'H 1-94 G<.'orgt: AJk•n Jnd Unwtn, London. pp 2 llh!•~ ncit.•c
W Jdt., M ( I '172) llydwnt,t .md ~~ yphozoa and other (Valt1.1blc -.umnun .tnd htbltn~rrJplw) llllCjll('\tll1
medu,,,,tJ, from thl• l'n:r tmbn~n Edt.lc.tra tJunJ. South Yongt', C.M (JW,H) 1t\img coral, / 1rtlfft'dllli!S ••/ lilt I hq wt•r
Au,tr,th.l p.,{,,,.,,,,o/•!1/Y 15 I'J7-22S. Stlriel)• H 169. 32lJ H. (lnv,tlu.lhlc: relerencr) LlllS Ill IK
Wall.cr, K.lt..tnd Albct'lt.tdt. L.P. (IY75) Ecological suc- Yonge. CM. (1')74) Cor.1l red; and moU Ill !),; tO I
ccssaou as .lll .1spen uf ~tru,rurc 111 fmsil communities. Tr,w>llCIItws t!/ till' R.•yal StJm•ty o(/:dwbur~lt 69, 14- the n,JJllt:
p,,{nrttbicJI<\'<Y 1, 2]K-S7 (Reef srntctur.ll development (Environmt•ntdl nilhl's in n·et$
lm l• r1:1tu
lllt:t' botl
the phylu
lll\ c1 tehr.1
tm•c' (.me
1'11\ltatll r
IJlOtc Cnlll
l \\ o gc•
mo• ~ cmn
morpholol
Ill ~· \11(( l'
)S.I
re
14
Bryozoans
Ill
rd
of
ea.
rk
In
in~
l\"0103, became they are ~mall ,md oft~n d.:Ilcate,
nal
1tld to be less famtliar th.1n mo~l Invertebrate
Bowerbonkia (Fig. 6. 1)
it~:.,
•ups prl·~aved 111 fm~il form. Yet they are com-
ta-
m 111 S{'dlmcntary rocks aml abund.mt 111 the sea Bou•er/)(111kio (Order Ctenostornat:t: Fig. 6. I;\ e) ha' a
46. h: Jt I ·.1~t 3500 living and lS 011!1 ti.mil speoes creeping cylindnC<ll stolon from whiCh numerous
kno\\ 11. The fronds of fl.n seJ\H'Cd~ such as bottle-sh.lped zooids arise m clusten.
,nJ rwn.t arc ofren covered \\'lth the l.tt:y calcareous The body w.11l~ of each living zomd (membra-
tal lth'm of the brvozo.1n \lcm/,rmliptlrd, each nom .md unpre.,ervable in this ca.~e) Jre c.tlleJ the
tilt 11) !wtn~ composed ofhundrt•th of Individuals. cystid. (The tenn z ooecium, wh1ch hJ' \Oilletlmes
) All bi')'<'ZO,Hls are colomal. mosr an: rnanne, and been used a\ ,tn altemative to cysttd, " now res-
I/ lit Ull' nt:tjonry of case~ the tlllHs or zoo1ds of the tricted to the calotied zooidal ~kdeton.) The laiCI-
ny scncrc ntbes or boxes of lime p.un.11ly encas- fied part\ of .1 whole colony may be called .1 zoarium.
~thc sol[ pam (e.g Fig. 6.~). Lath zoo1d is basi- When feeding, the zooid extend~ its lophopho re, a
l y rylmdnc.1l, h:ts .1 ring of tentadcs :md at first ring of' I 0 cdiated tentacles, into thc su rrounding
ht '~cn•s w resemble J sm.11l t nidan.tn polyp. water. These tentacles converge on J u.:ntr.d mouth
lll'l'Va, dw zooid~ are coclom.tte, h.winp; a freely from which the gut descends m to tht· body of the
-:;pcndc:d b'llt wHh both mouth .md .mm, and are zooid. Tlm gut rs U-shaped, hangmg down 111 the
UNton.tbly of a lughcr brr.uk of orbramzanon. codom .•md h,ts an oesophagus, stomach and imes-
\\Cn rccogntzcd as bemg dl\ttnct from cmdar- tme \\hrch opem bv an anus close to the mouth but
m IS20, .md the name Bryom.1 wa\ gtven for- outside the nng of tentacles. The tent.tde~ .m.1c:h to
y ro tht•rn m I ~31 bv Ehrt•nhng. 1 ye.u after the body b}- an eversible tt::ntade \hcJth. In
nunc Polyzoa had .1lready het·n b"~''en by the Bvu•erbn11kia and tts relatives the sheath t\ protened
1 IWUr,1hst J. V.lllghan Thompson. llowever, by a collar, but this is not reprcsentcd 111 other
1 r both Thomp~on and Ehrcnbcrg included in Bryozoa. l3crwecn the mouth and anlts ts a maJor
phylum. ,1\ understood by them, ('L'rt.tin other ganglion from which nerves run to all parts ol the
trtchr,tlt' group\, the true bryo1u.ms arc some- body. From the base of the stomach a thread, the
e' (.tnd e'pecrally rn Amencan U\.lgt·) known as funic ulus, connects to a main funic ular tube run-
lum htuprocta The tenn Ul)n7o.t, being in rung along the ~tolon and connecnng .111 the zoo1ds.
re common ll\e. IS ret:uned hen.· Tht· tcntacles can be either extruded for fCedmg
[~~oo gt ncr.1, one "m pie 111 con,tntt t1on Jnd one (F1g. 6.ll) or retracted entirely wtthm the body for
cm11pkx. Jre de,crib~d here to 'ho\\ the ba~u: protection (ftg. 6.ld). In the restmg zooid the ten-
!phtllogy. Both of rhe~e bdon!t m tht· Llrge~t and tacle slw.nh becomes mvaginatcd, wtth rh mrface
,, IUCtl'\\hll marine ctw,: 1he C) mnol.lemata. facing mwards and surrounding the tentadcs rather
144 Bryozoons
rh:an fucing outwards below them. Eversion of thr

tentacles is accomplished quitt• ~imply by cmnpw-
\IOn of the bod} wall b) tranwer,e mu~cb; thil
ra1ses the hydrostatic pn.·s~ure of the coelmmc thud
\O that the tentacles have to emerge. A lJrgc n:trac-
tor muscle. attached to the tentacular ba~e Jnd
accompanit:d by longitudinal lllusclcs. pulh the ten-
tacles wahin the body when d.mger thrcatcm, and
the zooid I'> finally dosed otr b) a circular 'Phim:trr
mmclc jll'lt bdow rhc collar. T he colbr then tiJld1
inwards in ,1 series of pleats and completes the
closure of the 1001d.
(d) When the 1ooid feeds. thc t~ntades Jrc extended
in an erect ti.mncl by hydrmt,ltlc prc\\Ure of th~
(cl
coelom. Thq ar~ quadrate 111 cross secuon and haw
ciliary tracts on each stde, those of adplcnt tcnt.l-
Hmtacle she~th dcs nearly touching. Another trrtct of 'frontal cih~
on each tentacle faces the mouth, bciu~ mor('
strongly developed tow:~rd' th~ ba\e. When th~
lateral ctha beat dO\\ 11\\ ards m a courdirutcd F1gvrE
merachronal rhythm, current~ arc gencr,ucd whl\.h (Anas
pa~s ~rraight down the funnel towards the mouth
( 16)
Toylor
and out between the tcnt,tcles ( l~ig. 6.lc). The food
particles (mainly phyropl:mkton) in the mtomm~
stream are in~cstcd by the mouth, but prt'CI'ely how dt•vc
thts is done 1s not entirely clear. The Atm rJ!c~ of ll.lO:U
l urrents leading directly to the mouth an: mud1 pore•
higher than peripherally, and posstbly only the pmi-
dcs in central srreanu are- c.1ptured. The operation
of rhe lophophore and lt\ ~tructure .1rc deCided] Sr.
smular to thmc of brachmpnd~. hence thc 13ryozo1
and B rachiopoda are ammtcd to be related and arc ,..,,;rt
(e) t,ueral Cilia
grouped together with phoromds in 'Superphylum' ph or;
~~l~~ Lophophor-ara. bryo2
~5...
There are no sepJrate excretory. Clrlulatory or ndua
~~~
respiratory org.ms. The colony grows from an 1rutul r.ldlat
zooid, the ancestrula, by h'Towth of the \tolon 7001d
(itself a seric~ of moditied toOids) and by asex'11~! .ltc. I
fromal coha
budding of new zooids (Fig. 6.3). New wl01u~. swolJ,
however, arc produced wxu,tlly. Thc 700tds arc l'g£'1 •
hermaphrodite. but the ov:try and tesns IDl~ Cl VICe
develop at d11Terent time\. The ovary IS a duster ol next
several egg cells, wruch .1re released one .n a tune \\',111;
Figure 6. I Bowerbonkia, a Recent ctenostome: (a) appearance into the tenl<lclc sheath where rhey nrc ferriJJZed ~l'para
of colony ( 1), (b) section of zoorium with extended and

retracted zooids (x 7); (c) zooid with tentacles extended (x 35
Each then develops into .1 trochophore Larva nr or
while still in thc tentacle ~heath. When fully nururt (thOU!
oppro)(.), (d) zoo1d with tentacles retracted (lC 35 opprOJt..); (e)
section through tentocle crown, with arrows showing current the larva ~wum away; meanwhile the zomd degen- pnma
directions. ((a) Redrawn from Bossier in Treatise on Invertebrate erates. A new egg i~ relc'l~cd only when the tan mdcl
Poleontology, Port A, (b)-(e) redrawn from Rylond, 1970.1 fertilized egg has developed into a larv:1. Spenn t·~t po
Living bryozoans 145
(b)
(e)
rg.Jre 6 2 Bryozoon structure· (a) Smiltino sp. (oscophoron cheilostome), Holocene, Antordico (x 16); (b) Coslonoporo magnifica
':'IO!Con cheilostome), Cretaceous, England (.: 1.4), (c) Orbignyopora (cheilostome borings on brochiopod), Silurion, Pennsylvania
61. (d) Crep,dacantha (oscophoron cheilostome), Recent, New Zealand (All SEM photographs reproduced by courtesy of Or P.D.
cylat)
doped '' 1thm the re~tt\ 1' wc:n 111 man) br.-
r zom' to make m way out mto tht· st'.l \'ia tmy
fl'l Ill thl' tl'llt,Kic:s.
Smittina (Figs 6 .2a, 6.4, 6.5)
ltm,, (Order Chetlo~cum.HJ, "iuborder A~co

oul h an example of 0111: of thl mort· complex
uzoam. The colony is encrmtmg..md the indt-
wl zootJ, .m: arr,mgcd likt· fl.tt dlipric tl hoxc~
1diJting a'' a} lrom the ;mccstrul.l or tlr,t-fonnc:d
oo1d. The\ .m: all comtructcd of t.llnum carbon-
~ Mature zooids may dcvdop dt\l,ll ovicclls,
ollen spherical structures in which the ferti lized
.., develop mto larvae. ln dttli.•n:nt tlll'tlo~tomc~
(b )
1cell1 111.1) lte \\ithin the zooid. n'verl.lp onto the
f xt one dtstally or be cmbeddc:d in ih posterior
e : altt:mati\·cl} tht·y nu) bt· whnll) or p.1rt1ally
~epmtt:d. Din:ctl} bdund tht· 0\tt:c:ll " tht: o rific e
a opcmng to the zooid. Tlm j, kc) holc-,haped
ugh tlm ts not munedJatdy t'\'tdent, 'incc the
:>nJ!Ur)' onhcc ~~ h1dden beneath tht• uppt•r 'urface) Figure 6.3 (a) Cupulodria (Mio.-Rec.), a young zoorium show-
ing growth from central oncestrulo - blank zooecio ore the lost
:t J clo~ctl by .m oper c ulum, lun~;t•d .lt tt' n.lrrO\\- formed (xl 0); (b) probable life position of zoorium. (Modified
n tl pomt~ on cardelles, so that when the opL'rLulum from Logoiij, 1963.)
146 Bryozoans
OPf•rculurn
rfmacted
ZOOid
reu.u;tor OVJfY S11ptuhm1

mu~cle
frgure ,
(b)
muscles
loshrng
Smlttoldtn mermor1111
Figure 6. 4 Structure of the oscophoron cheilostome Smittoideo

mormoro, show1ng mature 1ndividuols with OV1cells (right). Figure 6.5 (ol longrtudinol section through o generth
oscophoron with zooids retracted, (b) Smittordeo /!lCliiQI:
showing progress1ve development of the penstomrol ,.-
young zooids. (Redrawn from Hayward and Ryfond, 1979
ope m .1lmut tlm fuk nmt th~ d1,tal part (anter) ri~c'
:tnd the proxlln,tl p.trt (p oster) ~mb. When the
p olypide i\ retr.td~d. 1t be\ ~:nurely withm the The frontal wall (i.1.·. upper ,urface) i~ eo
1. }'tu.! .111J tlw onfiu~ 1\ doo;cd. When 1t emerge~. it Ill struuurc With a regular ~culpture, \crreu
comes through tht dt~tal p.m of the orifice :t~ the ~\.'vera! Ltycl"\. Chcilmtomc' (h g. h.2) ;ue 'tnm
Jnrer lift, up Conuet:tcd to the proximal part of polymorphic. a\ i~ ~hown by the 'prw
rh<. onhn· bl'IO\\ till' po~tcr 1s a o;ac, the ascus. stmcw rcs avicularia ,mJ vibracula, which
somcnme' I-.1HJ\\dl .1~ .1 con1peusatrix, which is both m o dified wuitk The aviutl.tria (Fig. 6.6~
wspendcd from the bod} wall by many parietal attached to Lhe uppt•r su rf~I Cl' at <;pccicHp
muscles. ·r ht'i tompel1\,ltrL'X 1s conccmed with location~.
polvpu.ie cxtrm1otl. When the radi.1l muscl~ con- One ku1d, fou nd in B~tg~tla.
tr.llf. tlw t ontp1.·m.1tnx t:xp.llld~ and the po~tc:r is head in \hape and conlaim a -;inglr mt
deprt'~'ul ~o th,ll w.\lcr cntel"\ thc \JC. A~ the com- polypidc. h h.t\ .1 h1ngcd d11tlltous mandible \1
pem.ltnx 'iwt 1". tlw polvptdc ~~ d1spbced and ha~ to opens .1nd \tups ~hut in ,, to mt.mt motion, t'1
emerge from llll .um r bcousc of the hydrostanc to d1scouragc both ptl'datol"\ and ~etthng l.m-a
pre'"tre. 'I ht• poh. p1dt• t.lll be pulkd back 111 ag.1in The polyp1dc 1' tt:duu:d to ,, rudunent, l
whl'n rlw rt·rr.wwr mu,cJ\.' rmltr.lct\. This cause-; the mam mtemal \Oft-pall 'itrunurc\ are the
compcm.ltnx tn b~· C\'.ll'u.th.·d hydrmtattcall} {~incc: ant.lgotmttc mmdt· ~l't'i w1th which the n
d1c r:tJi.tl mthdt'' .lrl' by tht\ tune relaxed) and the snaps. The m.1nd1bk j, mfn:yuently prN:n·cd Figvro o.7
npcrculum ,fHth. but -.mn: m t•dgc: gt•nt·r ally lib the avicubn ozoan grO\J!
Classification 147
fonn is quite weU rdkctcJ by the pre~ervable

skeleton. Aviculana may bt• \Cssllc or, in the most
extreme fom1, peJunul!,ttc, t.e. mounted on a 'ho1t
stalk. The least-modthed se~stle .IVtcularia are c;aid tO
be vicarious; these ,trc ,IJghtl} muller than nonnal
zooccia and rcpbce them 111 the t"olony at regular
intervals. Interzooidal ;wtcul.tna Ol"l"Ur between
zootds and are rt·duccd 111 ~ize, w h ereas the much
smaller adventitiou s .md ~omenme-. peduncul.tte
avtculana .lre found m,unl} 1n chetlo~tomcs in
whic.h they may occur anywhere on the frontal wall.
There 1s a commm1m between the t\.vo kmdc; of
avtcularium. R yland ( 1970) has dt~cus~cd their
possible evolution in some det.ttl.
F.gure 6.6 (a) Avicularium in lateral view with different sets of Vibracula (Fig. 6.6b) po~Sl'S~ a long wh1p-uke
11\Jsdes to open ond close, (b) vibraculum, with musculature for bnstle (seta) projecting from a sessile basal chamber
1oshiog shown (Redrawn from Ryland, 1970.)
which contains only the musclec;. The seta swings
on a pair of opposing pivots jmt above m lower end.
The mmclcs arc actacht•d hdow the pivots. When
triggered into acnon the conrracnon a~· the muscle~
cause~ the sera to la~h vJOicntly. Th1~ mmulatcs
neighbouring vibracula, .md thl· whole ensemble
\vill strike hard ag:llmt .my .then object, diKouragc
~ettling larvae or wmnow .1way \edunent. In lunuh-
tifonn bryozoam, vtbracula mw be modified a~
'legs' on the under\ltk of rhe colony, t'n,1blmg H to
move.
PHYLUM BRYOZOA {Pt1lyzo.1. Ord Rt:c} SNLit: wlotual

coelomat~'S. uomlJIJy man m·. r.1rd} frt:~hwatrr, lOnmtiug of
>mall linkt'd zooid,. li\U.tlly \\ tlh ,, lak.lrt'ou~ or mort: rarely
an organil skeleton. Zootd\ have J lt:ntadt:-bearmg retrJcolt'
lophophore and a U-~lupt•d gm \\ ith the Jm" ourstde the
tentacular ring. ColoniC\ .mw fiont .111 .uKntrul.t (nr r.trelv a
statoblast}, .tnd .trc cncru~ttng, rrt•cping, ca•t t m m ch.tms,
polymorphic in some groups.
The cl.tssiticauon .tdoptcd hcrt· t\ th.tt of Bo.trdm.m rr <tl.
(19t!3) in Robt~on, l'fl'tuiu ,,, ltll'rrtt•lmllt• Pr~/rolllo/()1!)' .md
McKinncy and Jackson ( 19H'J: 1-lg. (>.7).
s CLASS 1. PHYLACTOLAEMATA (' I crt.· Rct ): Non-CJ!carc-
ous freshwater bryozoam. /..omd~ wtth horsc~hoe-•hapt·d
, lophophon><;; \tJtobiJ\l\ produn·d .,, rntmg buds Twdw
c gencr.l only. J>/Jmllllc/1,,, Cmrmd/,t
CLASS 2. STENOLAEMATA (Ord -Rn.) CakUit·d manue
bryozoam, usuallv nlm-opt·rutiJt~. wJth JJt o:xtcm1vo: fo~
e
~11 ro:lord 7oOJd> .tre t ylmdnlJI aud clongo~to:d lOOl'llJ
:I conmmmg to grow throughout tht• ht~ of the .:oluny and
e ~et at an .mglc w lhl· dtrt'lllou uf n1louy h'T<l\\th Each
I, figxe 6.7 Time ranges and relahve abundance of main bry- polypidt: i~ 'urroundo:d hr ,, llll'lllbrJuou' '-it. T~·nudo:
czoon groups (Redrawn from McKinney and Jockson, 1989.) extru,Jon m hvmg timu' i' hrnught ,tbnul by mmcuiJr
148 Bryozoans
(a)
"
Z•
Figure 6.8 Trepostomes. (a) Anisotrypo (Carb), Alabama (x 6); (b) Dekoyio (Ord.) showing o longitudmol section (left), a tangen
tiol section w1th aconthostylos (above right) and a transverse section (below right). (Drown from o photograph by J.P. Ross in Joomd
of Paleontology, 1962 )
Figure
Englan.
(I) (b) (d)

Chllonous
nembrane
costa
(o)
{b)
Figure 6 9 (a), (b) Pouible ongin of Cretaceous cribrimorphs From membraniporoids, showing (a) ancestral membraniporood t1fX
with radiating apertural spines and (b) spines to form costae covering frontal chitinous membrane, (c) Pelmalopora gregoty~
Cretaceous cribrimorph (frontal voew) (x 25); (d) Tricepholopora pustulosa, a Cretaceous cnbnmorph (x 25 approx.) [(o}-{4 Figure 6.
Redrawn from lorwood, 1962, (d) photograph reproduced by courtesy of Dr G P. larwood ) (Redrawn
Classification 149
un fi>r 11~ ''"loll Ill thu,luno th,•pn>xun.ll p.1rt of the ,,•par.lted b\ ~".:udopornu' lronl.ll ".tlh. or rontJg\JOU\
..-t dt,pi.Jclll~ IIH t<mMI<, dl\t •lh. :;:;o gl'ncr.t pol) gonal Jpertlll o. lntt'll<HIC, 1.11 \\ .J], mtulh pu.:rccd
011DER 1 CYCLOSTOMATA (h< 11<1 k11111\n ,, Tubuh- b, mural pores, <'.)! Cnu.r, HtHIII(c,r (Fl;! 11.11),
por.tu, th< 11"-lg<' adopted 111 the I'' ntl<l' ''" ilwn1•·/lr.111' St<>lllollt>J'<'I•I (fl..: (I 11 ), ,\J,HIIItiiJI<>f I (h". h.J:!l
/'w/t(l/11(1/ !1!1~ l lrd - Rl"C ) I Ill t 01 <.'Ill nl\llllg Zll.ln.l ORDER 2 CYSTOPORATA (L l )rd U Perm .) ~mu!Jr to
ol wbubr zuo<l"l.l po,\C:,~Iug t"lthcr nrcul1r .tpertun" C\, hhh>me' hm '' 11h Cl'!;'""' of <lln ,) cy~tiphrag~ns
'eparatm~ /OIIC'<II 111d/ur hi\'< <IC•tlltl< pr<:Jt'<tlo'
(l unaria) .uouud zoo<'< 1.1l .tp< rtun•,; <' ~ h<tufll'•''·'·
( .:ff,lllhlJl,lf,l
ORDER 3. TREPOSTOMATA (Ord.- frm.) ~ronv hrv-
ozoam' fonmng m.l\\1\ < 111 m 1 "1th don~.lt< auto-
zooecia wlndt .m· mm.tll\' rhm w.•llcd but beromc
rill d. w.tllcd rlow to rlw 7o.Jn.ll 'url.lt c: whert' 'mJIJ
mesozooecia. tdkd by dmt·h· 'P·•rnl diaphrag ms.
Ol.l\ utter\'enc bct\H'CII .HH07oot·n.• Mur.1l pore'
;~b,cnt. e.g. \font~culipc>r<l, . ltm••tr)'Jl•' (ftg. fl.HJ). V<k.t)'t•t
(Fig. 6.Rh). f/nl/c>JW•t.
ORDER 4. CRYPTOSTOMATA (l . Ord, U. P.:nn.): Zo.tm
erect .llld !lt't' 111,,. 01 li mu ing hil.tllllll.lr sht•t•ts. auto-
zooid' sho t t, wnh h.lS.Il di.lphr.lgll" or llll0111pktt· p.trti-
tiom (hcmisept.t), t·.g . .Jrtlm>pllr<t.l!lll<l.
ORDER 5 FENESTRATA (J Ord U Pmu.) l:o.m.1 nnt
,111d llt't-Ji kt• \\ lth llllll.llnllll(t· lH,\1\.Jlt'\, Jllll>LllOith
f.gvre 6.10 Theonoa diploporo (Cyclostomata), Jurossic, ' hort .md rommonl\ wuh hemt\l')'t.t: t'.g. Tut~•,ftlloJ,
England, showmg radial ndges (xlO) Rlr.ll>dc>m•'''"'• .tl tcltrmcdt•!
ti}Xe 6.11 (a), (b) Bereniceo and (c), (d) Stomotopora, two Jurassic cydoslomes with contrasting adaptive morphologies.
;;Jdrown from Taylor, 19791
150 Bryozoans
m groups ,tt the \Uilllllll of small mound~ (n1ontir·

ules). The zooecial walls arc comtructed of rh1
laminae, With the mdtvJclual l,munae ,lt intervili
fomting melll\tlh-hke duphragrm or cro~)-paltl
tions There 1s no cummumcatwn through aUto·
zooccial wall~. Ucrwct•n the l.lrge zoot•nal apenur~
(autozooecia) tlwre arc oftt•n 'nuller operung~
(mesozoocna): thc\e 'uggt:st the lonner presence
~ome kmd of smaller pol}'morph1c zood
Fi!Jure 6. I 2 Mesenter~poro, showing lunate structure by means
of which overgrowth of old zooecoo has taken place. (Redrawn Somettmcs acanthostylcs (rod-hb..e spme' of con
from Toylor, 1976.) in-cone ak tte) arc vmble :.lt the mter~ecuom t
zooccial w:tll~.
The Trepo~tom.lta, wludt after thetr mittal.:vo-
CLASS 3. GYMNOLAEMATA (llrJ.-Rn.): Mannc. occa- luttonary burst bet.:.lllle \O ,tbund,mt m t
\tuoully brJdmh 111 fre-.1\\\.lt<r brH1ZOJn> whKh ma; be
Ordovtctan and Silun.1n. dechned thereafter •
c.llnfi,·d /.ouoJ, ,,.,. bm.-hk,· lll uo.oy form ~hon (·ylinder:s.
Tht:u \llC: "ltwJ ,•orlv 111 dewlupoHc:nt .1uJ tlwtr long ,JXIS
cryptostomes became domw.mt. and they timlh
uoim idl'' into thl· lu«ll tlut·rtion of growth of the: :zo:lr- dted out m the Triasstc.
IUill 7.omd, (lllllll'lll'd by .1 tionkul.tr network. The Cryptostomato appeared in
Lnphopht11C\ l'\'l'llt.:d lov lllll!ollll,n ddilmUtllln or pan or Ordovici:1n and, dcdining somewhat thereafttl
tlw body \\.111 St10ngl~ pltlynlorphu 650 geuerJ. held their own unul lhe end of the Pennian The
ORDER 1 CTENOSTOMATA (Ord.-Re,·.) Zuuid., unt.<l- Fencsrrata however, to wluch they arc related
uh,·d \\.111, memht llh HIS ur ~,·Louuou\, IJcking ovirl'ih,
reached rhc1r acme 111 till' later Palacozmc.
trnJn.-ntl\ pc net ran t l'x.unpJ ..., indude &,,,.,,-/.,mkia
(lot:;. (o.lo-.- Ho)'••mdtllllt, Oo/•r~ll}'•'l''"'' (F1g. 6.2c).
Fencstcllid colome-. (l;1g. 6.13a-d) grew from an
ORDER 2 CHEILOSTOMATA (Jur -Rec.): Zomd.~ ancestmla fomung tirsr a nng of zoocoa and then
cak.ncnu,, "tlh 'hnrt hnx-hkt· zonccoJ havmg .1 do~l Circlet of upnght branches: tht.•se btfurcated at uuer·
onftn• dthcd hv 1 hm~cd opnrulum. Aviculan.1 vals gtvmg the colonv a cup-~haped form or m ~m
mJ Vlbr ~rul t ' t unnwn l•mhn n' hroodcd m oviccll• spcncs the form of a haH-cup or fan
Tht· four 'uborJc:n Jrc d.:ftncJ on calnJicauon of Whatever the tonn of the colony, the br.an.:h (I)
the frt,nt.ll \\ .11J .tnd Juordmv; to how the lophophorc
are submangubr m cro~\-\I.'CUon and have row~
1~ prutmdnl [ \.unph:' mdudt Srmtlftr.l (Ftg> 6.2. 6 4,
zooect::t opemng omo onl} une t:-tce of the rolon
1>.'1) • •\loml•r•IIIIJ'•''''• <.n/Jnllllol, l'tfm,li<'J'<'1•1 (F1g. 6 9c).
Curm/,,./"'' {f-1~ t. 't,h), C"'l'"ia,u,r/w (Fig. 6.~d).
C w'""'l'''''' (Ftg. h .:!b).
The branches 1rc connected by rrus~-b::tn \\1th rn:-
tangular ~paCe\ or fe n estru]es between (Fig. l).\3e
The skeleton w:~s probably secreted b.,. a thm
-·
eptthdmm whJt.:h extended over tht" whole \Ur!J,
and, as tn some hvmg cyclostome~. the zootds pr11b-
ably sh:~red a common Ct)clom. '
~hortly <lttel their fiN appe.1r-ance in the early Since the growth of the bryo7oan skclet<ln 11
Ordov1rt.'ln the thyozo.l underwent a great burst ot 'coloni,tl effort', the colony may become lngh
evolutton, re~ultin~ in thl' est.1bli~hment of the early integrated r.1ther than n•m::tin .1s no more than
~rcnolacmat:1, wh1ch by the l.1te Ordoviri:1n were aggregate of md1vtdu.ll~. The fum·t10naJ morph,,!
vt"ry ,Jbund<lnl .md c..ltwrst" Tht" !:ttenolaemata are all ogy of bryozo,111 coloniC\ IS potentially .1 frUit'
extinct nnw .lp.lrt twm the Cvclmtonuta. but were field for research. l"cncsrclltd colomcs, to t.1h
the dom111.1nt d;w, ol P tlaeozo1c bryozoans. Of all example, h:1vc been .1nalysed functiOnally by Co\1
the ~tt·nohcm.lta thl' mmt 1mport1nt and abundant and R.idcr (1972). I he arr:mgcm1.'nt of the zoooo.. F1gure 6 13
fenestellids:
in Lower P.ll.wozoic rime\ were the 'stony bry- and the fom1 of the zoanum are clt'.1rly 1mpon.mt zooecio (b)
07o,lm', the ·r repmtom.H.l (Fig. 6.Ra,b) The}' under~randing how the colony operated as J wh Archimecles
tomlt'd 'nck-lih· or glohul.tr c,llc.ucom zoana up to It has been pmmlared rh.lt dw 'opt•t-ation.tl subu · estcllid, u
of the fcnc~tclltd colony 1rc the tcncsm1b. In (Carb.-Perrr
50 cm .H ro''· I he mo<.'<.' i.l .ue tubular and closely
os ext?"dill!J
p.td.. t'd With crm~-p.lrtltiOn' and thickened dJStaJ mam branches. the lophophon:s of equally 'P.K pre!ohon of
p.lrt\. ')omt'tlllll'\ tlw 1om·n.tl aperntres are clustered zomd~ on oppm1rc <;Jdc\ of .1 mcdi.1n ndge Coweo and I
Morphology and evolution 151
(cl (d) considered to h.•ve L'Xtc.;·ndcd into the fenc~rmk

fonmng a tiltenng net, rhc u·o,~-bar. gl\ mg \llppon
and ilividmg up rhe lcnc.;•,rmlc.;·, A combined fct·ding
cum:nt set up by the 7mmh '' ould drt\\ water in
through the tcne~tmk,. c..'tl.lbhng tht•m to 'train off
all tht> food matenal \\'Hh thc.;·ir lophophon:,_ Jnd
would thm g~vc lugh tilcering c.:Otucm· ~ .
The growth fonm of knc\tdhd c:olomc' m.ty
have been adapted for nuxtnllztng tiltcring dti-
C1CtK} Jt tht> tolumal kvcl ' I wo kimb of ntp-
.shaped colontcs .1n: kno\>vn: thPsc.;• With zooeci.1
tacmg outward, (c.;ummon 111 till ~ilun.ltl), and
thost> with inward-l.tLIIlg :wocn.tl .tpc.;•rturc.;•, (mor~
cotmnon in the Carbotukrou,). Thl•n.• .1re .1l'o tan-
shaped (half-cup) and 'PII'll colonic'. The cup-
shaped colomcs would probably be L'I.Jllally ctliricnt
whether they drl·w ill W<llCr lt om the top .md
exhaled it laterally through the tt:ne~tntlc~. or
whether they ~ucked it in at the ~itb (like a ~pongc)
and sent out an cxcurn.·m strl•am trom the central
caVlty. Fan-~hapcd colonic, (agam ,1, wtth cc.;·rt.lin
sponges) would haw the bc~t ,h,tpL' for takmg 111
w.Her 111 .1 regt me wheu: .1 \\ cak l urn:nt w:h con-
~tantly Aowmg nonnJlto rhc tan 'urlac:e.
The pecuhar Ardumcdc.1 ha~ J typtctlly tl-nc\tdhd
network ~tmcmre. but tlm describe~ a hdtGll 'PlrJI
around a duck ccnrr.1l cakarC'(lll\ LolunUl. ~ud1 a
tj bf}'070.1!1 ~tandmg upnght ort th~: 5e,t floor" ould be
r fi.mcnon.tll) effinem tf the rombtned anion of the
lophophore~ generated a current 'cream coming in
.lt the top and nmmng 'ptrall} .tlon~ the 'dec.;k' to
the b,t"'· J~ do\\ n .1 f.ur~ound hcltcr-skclter (I ig.
o. 13~. A~ the m:un srrcam travelled ~ptr,tlly down-
wards, some of 1t would be ~m ked tltwugh the
innumerable fenestru lcs .1 nd pa~\ed ,\\.\ Jy 111 .1 c.;cn-
trifugal stream jmt bdow e:1c h deck. !:!inn.• the fen-
tl estdlid feeding current rN·omtructiLms were
y conceived, based upon fo~~i l materi,tl .tlone. some
n very ~imilar fu nctJOn:JI :1n.1loguc~ h.tw been found
in the modem chcilostome Hcrcpttl,l, wh1ch n:scm-
11 blc~ and feed~ in prcci~cl) tht· m.ltUH:·r whJCh had
ll been predicted for fi:-tw~tdltth.
ll Nor uncommonly, new sp1ral .-ilf!Jimrdn colome~
~ ~ 6 13 Morphology and d1reclion of cvrrents mferred in
.a originated from prC-C'\1\1111~. ,lgl'lllg branches.
'II!Siellids: (o) Carboniferous fenestellid with inward-facing
11 :«10, (b) Silurion fenestellid w1th outward-facing zooecio; (c) Possibl} hundreds or even thou-.mds c.;lf genetJLalJy
11ttnedes (Corb -Perm ) a sp•rol colony (d) fan-shaped fen- idcnnoJ colome~ were pmdltlcd m thl\ \\ J) ti:om
using unidirechonol current, (e) Fenestella an original t(mndcr (Me Kmncy. JliHJ). L1kc\\ tsc..· the
~...Perm.), port of the zoorium showing zoo1ds reconstructed
•C edeoding into fenestrules (K 8 approx.); (f) Archimedes, inter- living cheilosmmc Hut:u/,, llllfif,l ha~ crc.;•c.; t sptral
of Feeding by spiralling currents. (Redrawn from rolomc\, the result of cqmv.tknt ~owth parameter.,
>'!11 and Rider, 1972 ) as computer modclltng h,1, ,!Jown
1 52 Bryozoans
\unll' knc~tdhd' \\'nl' h rc-,h.tpcd. c .g. Lyrtlj>Nfl. hl.'rc, both from d1l.' dttli:n:ntul spanng of autozo I he
ThL'} h.t\'l .1 tha knll'd m.trs>in.ll rim 'upporting .1 l i.tl .tpnturc' and from thctr onentatton (Ta)1 ICH!I
I~ nc,tr.HL' ne m ork m tlw ~h.tpc of ,, bowed tan. I979). '-.omt• bryowans hJ\ l.' regu!Jrl) spaced 3\ L(lfO
I hc'c hryuz, l,llh (I\\< K mnt·~. I 977) t'Vtdcmlv l.t) zonen.tl .tpt·nure~. often .uT.mged m a sy,rem Mt:
thr upon 1ht' 'L'.t floor \1\ th.n there WJ~ .l n open he\..tgon.tl do,t· p.Kking. Uut 111 many kind3 ot b l) lh
v.llllr lwhl\\ tlw l.tn. the zooid' being on the upper ozo,m, the: .IUlO/Olll'Ua arc UIH.'Vc.'IUY dJ\tnbUt Cl<!,
\Urf.ICt'. The fm '' '' pn,h.tbl} orientated with tht· o\er the colonv ~urt:.c:e. l·or example. the Jur$ .md
proxim.tl end nn thL· up,tn:am ''de. ~o that '',Iter cydosmmc ''""'f1'1''', an crec t br.lllching genus. uc.ns
\\ ·'' 'urh·J throu~h tht· tcne,tmk'i. tiltered .md dis- .tutozoocctal aperture\ .lrranged m b.mds .ur.ull# pu1c
charged thwu~h the 11pen cud of the \'Jlllt. cros~\Yl'\c.' ro tht· .l'\1\ of rhe br.mch In 'nu·ctn.>a I over]
Cf) ptmtomc' ,md fcne,tr.ne' outcbsscd the trc- 6.10) the dl\rotdal .lpcrturc\ are arrJnged m t2dL lugt•
pmromc' 111 the l.ut· P.tl.tcozoic, but by the end of bLuH.ile, (l?t,ttck~) wluch can be rcg.udc:d .1~ u.tllv
the Pl·nni.tn they tm1 wt·re LXtmlt. colontc,. In the l.trgc. mbsphcncal colome' o• h .lvt•
M.my l..tn<h 111 !JJI,tcozotl bryozo.m~ exhibit Pltoccne .\lt'<llldr<tpora the 'ur6ce dhplay, arcu lllllll(
humeonwrphy. smce there .1re unly a lmmcd f:.J~nclcs. ead1 bounded by .In exterior wall. Du goll.'r
numbcr 1>l 'w.tv' ol bL·mg a bryozo.1n · (B iake, 1980; rhc growth of tht· colony rhe'c fa~ciclcs altcm.ltt tlowc
l.1ylm .md U.tdVI.'. llJ95). Such cvolmionary dif:lcrenn:trc .md anasrommt· (B.tlson .md Tay;tl! Jb(J\'(
l UliVl.'l'genu: tall npl.'t.ltl' l'Hht•r tlt the level of the 19H2). 'i(tl
colony m tht· Zlll'td. I hm there .uc lyre-shaped In .111 rhc~c r;~~c,, thl.' rq;iom of du,tered am"' tuni.
fcne,tr.tll'' .utd L' dost1Hm:~ of virtually ident1C;t] zoocn:t - whatCVl'r rht·tr arr:mgemcnt - muq •.Ht:aS.
form. upt•t,tung l'lmltJon.tlly m the ~.tme w:~y. In s1tc~ of 111h.tknr How. The do,cl~ packed zo, grt> \\t
rara 111st.Uitl'' both zut,Jd .1nd colony fom1 arc coopcr:ued ro produ{ e a ~tronger current, file I~ po
\Uongly humt•nmnrphtt, .ls 111 the Crcnct·ous more \\ater 111 11111t rtml'. th.m \\ ould h.tVL' bnnp ~ol,ln
CltipJ.,nk,uitltl. ,, cht·tlu,ttllllt' "nh long rububr stblc 1f rhev lucl npl·r.ltecl mdivtduJIJy. The l"XI:ll rdugc
zoocn,l .JU't hkt• tJw,l.!' c,f c~ do,tonu·'· :~nd a colony CLIITL'llt\ \\.ere dt.lllnl'lil'd ,l\\'.1)' through rl'l:l \ H ill Id
type of rcnurbbl) 'uni!Jr tonn. dev01d of auto7omd\. Another '>uch ~y,tem h cxl:t .uJ.\. I
I he Cydo,tnm.n.t (f tg 6 . 1 I) \\'Crc rhc onlv itcd by tht· m.m\ hrvmo.m colonic' in \\luch J, Wf}'
~rroup of , .tk.ll't'Oil' bryn70.111\ ro cro~o; thl' 'urftcc t\ nt,c'd mm mounili or mooticules Wd~ rn
P.ll.leozot c-!\1t·~o7mr hound\!")' (though the uncJI- amozootd, an: lul.ltl'd 111 the deprl'\stom be~WC~:. rcplat ~
nticd ctt'nn,tnmt·~ dtd .tJ,o and survl\ ed into the,e monncuJc,. md betau\c of thetr settm~ on ,ompt•
tht· Jur.l~'tc) \IIKt' there \\'1\ little competition conc.tvc surt~KC thc:ir tl:nt,lde crowm ' rl"gardc
dwy Wt'rt' .thk to dommatl' the Mesozoic srenc. together. Thc'e dcpn.·s~Jom must have been m lcrm,,
1 ht·y h.td bL'l'011lt' Vl'ry dtvt'f\e and important by the ~ite\ of COOperclllVL' lllhJlJnt CUrrentS, Wht(e ''herea
lnwt·r ( rct.ll t•ou'; tmkni .tccording ro R ybnd momicuk \Ulllllltb. "'here autuzoocLiJ are 'PlN' I he
(I CJ70) 'tiHtr 7t'l1Hh durmg tlm period comtitutcs absent, at tcJ Js c~luiJnt duumcy~ (Dantl r1 t lllll tl
one oftlw htghhglm 111 tltl' lmtory of the Bryozoa'. 1974; T.tylm, l <J7H). fhe powerful exlwlant th e gt
They h,tvt' dcclmcd (rcl.nive to the chcilostomcs) rents, in their turn. were ,1blc to clear \edimem < h. dos
smct·, chouf:{h Mlllll' ~urvivt· to the prt·,ent, and since to di~pen.e l,trv.w or t'Xl'rct.l. tlt l CS\ f
they .lrt' dtL' only living repn.:\ctltativc3 of an ancient 11 omeomorphy, .1~ previously mcntionetl. pnth;-.bl
~tock, tnmt of onr t nnrcptitm' of soft-part mor- common. ~uch l'volutton.tr)' convergence canv 'tcnnst<
phology m P.tl.tt'n7mc brynzoan' arc based upon ate cithc1 .tt the level of the colony or tht' w wit l\t'
them The r¥clmtnute zootd i' quite like tlut of Thu~ there .lrc lytt·-shJpcd fcne~trJtt>< Utt tl tt(
Ji.,wcrbmtkw, though \\ 1lh .1 tL w dlJ.ractcri~tic diJfer- cyclmtomc\ of vtrtuaUy tdenticaJ fonn. In t ( >rJt•
llll l'\, .md tltt rt' '' \OIIIC dq~rt•t of polynltlrph.i3m. imtJntes both zou1d .md colony form are m lour ~u l
,'v1.my n·dmtnmt• tnlonic~ 'ccm to have been homeomorph1c, ,,, m thl.' CrLtJccous Clt~pli!rtb llld "",,
qun~: \\l'llmtq.,rr.ttnl ..md the tolonv rather than the a chctlostomt• whete thl' ?omds, long Jnd cl ~ n bc·
indi,•idu.11. ·'' with km·~tdlids. i' the funcuonal unit arc vmu.tlh tdt'ntii.JI wnh thml' of c\·cJo,tom \\tthm t
in tt·nm of ctTn tl\'t' ll'edmg. the t) pt· ol wlon\ (1 avlor .md Badve. IIJ95 ll(ln Ill tl
lt i' po"tbk to tnlt r the tonncr pre~t·nce and During Jur,\\\lt lllnt dll.' genera '\~cwwtc'~ 111 the ~c
dtrccttnn of t'Xlr.tltHltdJI tceumg current sy~tcrm Bcrnllc"C<I (Ftg. 6.11) were the donun.lnt cvclos I hl' ,
Morphology and evolution 153
The) m: both cucrmtc.-.; .mJ whereas thq .lrt· ottcn extrude~ through the acuon of mtemal mu~ck~ on
found together the) .m: qutte unlike each otht-r 111 rhe flexible frontal membrane. Uoch the Anasca and
''''ll) fi)ml. and theit .1d,1pt1vc strateg1es wen· very the Ascophora are very important today (Ryland
ITacm (T.1ylor, I f.J79). o;,,,lll,lfllpom colonic~ tonsi~t and Hayward, 1977; Hayward and Rybnd, 1979).
f JKhotomously bra m hm~ tumcnal rmn of zoo e- Cribrimorpha (Fig. 6.9). however, after J. bncf
h\ meam of whKh the colony spread~ rap1d.ly though \Ub~tanrial expansion 111 the late Creraceou'
JDd r:~dully The angk bt·twl'en ~ucces~1vc b1turca- (e.g. Larwood, 1962) have now greatly dechned.
~ dmumshe\ as thl• colony grmvs and. ,h com- These arl' U\ually unilaminar encrusting form~ wllh
rut~r sunulatiom haH' \hown, thi~ dday~ the calcilied ~idc wails, though the primary frontal wall
~riJppmg of brandtc~ unril the colony IS very in whith the aperture lies is of chitin. Tlus membr,l-
r~~. lkrellift'<1 colonies, on the other hand . .uc ini- nom vial! is overarchcd by calcareous costae or rib~
Jll, I~1Hhaped, lat<:r lK·cormng dJsc01dal. lhey which fom1 ,1 ~econdary frontal wall and meet 10 the
it mu~h more clo,eh p.Kkcd zooec1a. hnkcd by nudline. These fuse, making a kind of porous cage
:llll(rous pores. Tht: lolon\ w:~~ weU mtcgnred, over the pnmary wall - perhaps the most elaborate
~mmg cxrrazoOJdal kcd1ng current'. which wail structure<, ever evolved 1n the bryozoans. The
wrd rad.Jally mw.mh .llld discharged upwards apertur.tl region is usually protected by a scmJCtrcle
,,,~the colony centre. of oral ~pines.
\1<111/trlopom has been intcrpracd J-" .111 oppor- Cheilo,tomcs, by their marked polymorphism of
·nistic ~peoes capabk of r.tpidly colomzmg new rhe avicularia and vibracul.l <lnd by the connect1ons
lhc advantage 1.•l lt' urmcrial dichotomom between zoo1ds, e><prc~s a h1gh degree of mtegr.t-
111h i~ that the colon\- lllllld spread l.)tllckl)- from tion. \duch may have involved some mod1ficanon
romt of ong:m. Somt: pan' of the exp.mdmg of the ~lnll cure of the colony to make max1mum
~~~~~ mtght there fort: lx· Jblc to lOt Jtc o,Jfc ctfecnve use of the feeding currents.
hl!:tc~·. e.g. recessl'~ m th~: sub~tratl', where they The geological history of bryozoans is incom-
uld bt' protected fi:om pn:da6on or other h.tz- pletely known because of tht: poor record of the
1 fkmucea, on tht: othl't h.tnd, although more non-calcified forms. Ncvcrthcles!>, it is clear th.lt
:h· ~owmg, used ~p.llt morl' econonuc.lll\' and pamcular group~ were donun.lnt at certam nmc\
much more fully mtCh'T.Ited. Bcru11m1 nonnally (Fig. 6.7). In the early PalaeozOJc the trepostomc~
lan·d i)r<llnoltopora m ccologtcal sucn'''IO!l .md. and tryptosromes were especially unport.Jnt,
~ung more etfel.tl'vd} tor ~ubstrat1. \fldl'l'. i' wherc.1s the acme of the fenestrates wa~ in thl' late
tied,,~ an equilibrium specie3. In cvolullon.try P:tlaeozoic. After the Pemu.m extinctiom the
lC n,, thndorc, SttllllclftltJtlY<l is an r-~tratcgist, cyclostome~. wltich had been present throughout
c rrm Brrewcea i~ .1 K-str:ltl.'~t5t. the Pal.1cozoic, expanded vastly in the Jurassic and
I Tile rcbriw numcnc.1l dcdme of tl1l' Cydoo;to- late Cretacl'ous, decliJUng m relative abundance
r. m the Luer Cretaccuu' Jnd Tern an relate' to only whl'n rhe cheilostomes arose to become the
.:rcat conccmpor.Hli.'Oll\ e:\.-paoston of tht: dommant bryozoans of the latest Cretaccou' and
!hlltomata, the b~t 1nd perhaps the most Tl·rtiary. In Rccem seas they arc perhap~ the mmr
~''tul of tl1c bryouun orders to Jnsc. They numcrou~ lophophor::ttes. In each of the~e group~
•bJhly originated nllt from cyclostome·~ but lrom tht• degree of colonial mtegratJon varie~. lr 1s
IM>~tomcs, which wac rhen 111 eXJ~tCIIl.l' but claimed indeed that the colony rather than the zooid
~ fo,~il record io; pc10r ~1nce they 01rl' ah\,\}~ ha~ been the unit of natural selection. Thm in
lld :.llntlcd. R ccl'nr sea\ the specializanon of .lvicularia and othn
:lrdcr Chetlostomat.l 1s d1v1ded bv RylanJ mro polymorph~ can only be ~een 111 terms of bl.'nclil to
IUbordc.-.;: A~coph<lr,J, An.l\C,l, GynmocptJdca thl' colony "' a whole_ In addHion, polymorphism
Cribnmorph::t. A~t-nphor,lll ~tructure h;ts been occurs i u 75% of all living che1lostomes, e~pct i.tlly
1nb~u above 111 rd.tnon to Smilfirw, though in those ~pecics living in pn·d1ct.:Jble environutl'nt~
m tlm suborder then· IS much stntctural V<lrJ.l- where ~ufficicnt food resource~ would aUO\\ the
ui the pmition of thl.' ovtcdl <lnd the on fin: ,md 'luxury· of non-feeding zoo1ds.
Slulprurc of the uppc•r \UrflCc. Several trends, indicanng progressive adapnve
ht Ana\O l,tck thl• 1\Cll\, .md thl.' polyptdc evolution m bryozoans, have been dcf111t:d
1 54 Bryozoans
(McKmtwv tnd J.tL·k,on, I CJR')) Example~ arc The depth rangL'~ of many brymo.1n 'pecJes fr lnlr,J:t
c.tloliL.ltWn of ZOOJLk m creased integra-
I Ill rea sed 'hallower water> arc well knnwn .wd they r~nd t Q tlc•t
tHlll ot zntltth 111 l hdlmwme<., .md " ~hift of rig~dly fonn t·har.lctt.'rt\llr '"'nci1ttnn' with otht:r on: rtm 1
actt 'PL'CIC' Into JeL'PL'r water. Such long-tcnn. t\lm, som~: of whll h have bn~n rem,trkably pt 6 1111<
ottcn pul) phylcuc !d.1pt.1uum c.uJ mu ally be rd.ned rent through ttme (Hayw.ud 1'1 al., 1994: SITI! of 1 1\'
to thL· 1rnprm L'llll'llt of the colony .1~ .1 living mech- 1995). ptc\ 1C
.ml,lll, or w dd~nd a~df ag.umt predator:.. Bryozoan' may be cncruslcl">. oi erect culo 11\ lllg
form. free hvmg. or lllJ)' live rooted 111 soft Em
mcnt. Such growth ton m an.· a rcfkcuon of ad.p- ht'Cl<;
uw 'trateg~es, and J<; McKinne)' and jaLkson (19 op of
comment, 'growth p.:mern .md tom1 of a bryoz tllll ( 0
~
Uryozo.1n' .ut abund.mt 111 all oceans with a maxi- colony i-. .1n expn.·s~10n of Jt\ ccolot,.'lcal ruche m lltcn·
mum m the W\.'\tcrn P.tufiL . ThL·y .tre found at ail than 1t~ phylogenetlt h1story' . mdud
dL·pths tn1m till' ~horehne down to the abyssal zone Encrustmg tonm tl.'nd to be I.'Onunon m sh.illo ~ul"tr.J
- the dcepc\t rt'l urd " over l:l5tlll m - but they w:~ters, owmg to t ht> ,lv.ul.tbihty of suitable 1ub fl,,,,r I
detrt',l\l' 111 number' .md nnport.mce in the tauna strate~, but arl' more common on ephemernl tl JhOVl'
w1th depth. long-lasting ~ub~tr.ltcs, e.g. dead shells. and \( and '"
'J he control\or dlstnbution .m: •• ~ follows. weeds. Although thl.:'y do occm also on stablr \UI' lnllmJ
\trates, they .m: mually Otltl:tsted by domuu IIIISII<I
-lempe1 .llule, ftll though ,, fc'" 'pccics arc cury- competitors. Some Mesozou: spt:nes were able wttbl
thcrm.llnttl\t h.t\l' rc~tnttcd tt·mpaatLJre ranges. produce frontal or penpher.tl o;ubcolonies. and ntM gcnc1;.a
Wave .tdH\11, \lllu: tilL Loloull''i are liable to though not all, \CL'Ill to h.we reproduced onct: "' •n,l .11 c
J.unage. :llld d1ed ~oon ati:erwJrd~ (McKmncy and T~Jl ddJliill
The J\';lll.ibJilt\ of .1 hard \ulmrate upon wh1ch 1997) They may be nmnero; (cf the Jum 1t Ku1
dw l.uvae t',JII 'ettk·. Tlm 1~ e\pcciill} important Scomflfn{'ofll), or t(mn ,beets com1mng of a s1 tJVC ~11
111 limiting Lkpth range. tiJr deeper >ea sediments l.1yer or ~evcr.1l layer<;; the latter kmd may i proble1
1re mm h liner tl1.1n tho<;e of the contmental mound' and th1ck sheet~. Mulolayered 'hect'l A lr
shc:lvn and dt.•t•p-~t'.l ooze, otTer llttle prospect of common on sub,trates of lumted dnnemion~ s 2•':111,
:1 hrm .tilt hm.Jgt·. I here .1re. however, some jur.1~~~c enrru~tmg bryozoam are known to 114 \[011 •
gymnola<"m.IIL'\ wh1ch r.1n \ctcle on fine panicles. made the be,r u'>c of l11mtcd mbstr.ltes hy rlpa •HI tlw
.ltld ,111chur tht·lmdn·, bv root~ which p<"netrate overgrowth of old :roocc1.1 b)' young autmooeat Smcc tl
the ~u b~tr aru 111. orientated 111 the same direction. During \uch 'mJII d
SaiiiHt). whtl h I~ .1 r:url) unportam control. For nl:uncll.lr' gmwrh (T.1ylor. 1976). nnv zo h1vc .11
exampk. ~inn~ \\,Hero; olflargc nver mouths have develop :.long a charactL'mtlc C-,hapcd gruv. fonm tc
redun·d \,thn•tv ,1\ wt.>Jl as much suspended sedj- m:~rgm (F1g. 6. 12). The two end~ of the npplt·d
, ment. ft•w bryo:zo.tm .1re found there. Even ~o. remained stationary whik \Ucces\ivc increm <>ne
rher(' ,1r(' ,1 fnv L'uryh.llmc bryozoans which can expanded ,\way from them, rct:.uning the 'C' 'h I dll' \
wirhst.111d ,,\ltnitte~ of only :w per mille, including As the growth margins cxp:ltlded, they usual!)' t 1 m. 11 it·u s
rhe llhiCJUIIOU\ l1nwcr!ltJIIkh1. in contJCt with other C-shapcd growth margin~ IHlll 111 I
which tht•r~: wne 111.111Y per colony, and till rt h ~~ hl'l'l
tant inter.tctiom were nfren nf complex fom1. lr hvcs .1
Shallow- water bryozoans gn·.1t .Jdv.mtagc of tlm 'Y'tcm '" that the colnn\ 1~ lilust
whole thm has a longer hfe'lp.m than normal. lthur.Hc
Rd.1nvclv lt'\\ Rl.'u m brvozn.ms are intertidal ~llKC repeated inrrustJtton of the same area. 'v\hcrc.h, rate ten
rh~: high L n\lrotlllll'tlt.tl ent·rb')' and the problem~ of zooid is allowed an t.qu.tl k·ngth ofltfe bcfi)rt' I 1\ llt lrtll<l
d~.:,icc.HIOII ht'l\H't'll udes arc too great tor ,uch dcl- covered by the next layer. to saltnit
icatL' org.umms ThL· sublittoral zone. however. ha~ Another kind of mulnlamdlar growth ha, l he I.
.1 wealth ofbrvoz~l,m cnlome'. Thq feed largd) on 'hown in the Mcsozoic tubular bf)Ol tlw colc
thL· .1bund.uu ph~ ltlplanktun of tlu~ zone and ,1rc Rcpt!lmllltwwa (Nye and Lemone, 1978). He (c g. qu:t
e'pcn.tll} c umnmn .lt dL·pths bct\YI.'l n 20 and 80 m. parcnt.tl zom·num g;.wt• rise to two or more .tnd ~hell
Ecology and distribution 155
crazno~l iJJ bud.' (fouuJmg zout:nl). The'>e then dcmem, but nor too brgc. Cupulndnn (Ftg. 6 3) cJn
llesccd ,1\ tht:y grt:\\ up Jnd over the .1pertural tolerate .t certam amount of mud l>incc the vtbracul.l
nms of th~ parent zoocciJ and merged \\ ith .1Jjacent whip constantly Jnd prevent the ~cctlcmcnt nf day
undmg zoot..:ria. Thu' there ongm.neJ .1 new layer particle>.
i nvcrhrrowth~ wluch, a> u1 the gcrH.'I.l mentioned Colollle~ Jre .llwJys lunulitifom1. i.e. h.wc .1 con-
~tvtousl y, rcmed eXISting space wnh a new l.1yer of cavo-convex lc nsoidal form, :md re~t on tht..: botwm
nng ZUUCCIJ. r:~ised o n vihr:~cu lar setae. M an y lunulitiform types
En et hryow.lm may ton11 unihmtn.u or btlamm,lr (such a~ the rcbtL'd Sele11arin, though not Cupulndria)
eth, tlr arborcst..:ent colonies where the growmg c:~n 'w:~lk' on thc~e setae acros~ coaf"'e uncoiiSob-
of the brmchcs h.we repeatedly d1v1dt.d to pro- dated ~cdimcnt-.. Sinre the temperature lmutl> of
~ trc~.·-likc.. form (adeonifom1 when btl.munar). Recent colonic\ arc well kn0\\11, and asmnung no
ben· .Ire \(.vera! ad\~Jntag.:' 111 erect growth, change of hah1tJt preference through tunc. H ha'
1
udmg tht. uKreJ!>e of tis!>Ul' Jrea rd.1uvc to the been pm~1blc to use C. cmwnwsis a\ a good
tntJ, the r;ming of the colony .1bovc the ~ea pJbeorcmpcr:~ture gauge. Since rt i~ conunon 111
r b~\'lllg unproved acce~s ro feedlllg b'TOunds Miocenc and Plioccne sediments of the North ~e.1
,)\, tht.· ~c.1 Hoor . .1nd prorectton ti·om predarors Basin, the temperature of the water dunng dcpo~i
utd ~\\~rnpiug by 5cdiment. Of tht.• t.'rt.•ct fonm tion of th e~L' sediments m ust have been Jt least 9°C
unJ m ~hJllow waters, many, e.g. the bJI,lminar higher tlun 1t i~ today. The firsr appearance of tillS
u~~~ h.wc tlcXJble zoari:1 whtch ,uc better .1ble to species i~ a good stratigraphical marker to• the
th.'tdnd lUrrcnt act10m. Stout!) comtmctt:d crt'ct Lower Mioccnc. and on this criterion !>everal su1te~
r. nm '~n w1th~tand modcntc current 'trengths of Tertiary \cdimcnt' have been amgned to thetr
~rr dblt• to .tb~orb the ~trc'"~'· whtlt.• th~· more correct sysrcm
r, .!clic:ltely br.mched fum1' prd~r quil-t w atcrs. Rtwtcd bryo:.:oans (most of winch arc
IC 1 1\umcy and j.Kk~on ( 19H )) di,cu'' \'.lnOU\ .ldJp-
1 chetlostomt.''i) are erect colorues anachcd bv long
le • 1trate~c:' 111 erect bryozn.m'i Wtth re'ip~·ct to tube~ from the prox.tmaJ end to gr:um of sedmtent.
m 1' blwts nfbreakage and Ao\\. Some of the~t.· (e.g. Splltlcropt>ra) proJect .1bove the
re \ lice lrvmg h.1bit has hecn commnn 111 bry- sediment surf..rct.· like small fu ngi, though some may
Ill' o.mnince the Ordoviet.ln, :md whik rn.my ufthe reach IS t·m in l ~t:i~ht. R ooted bryozo::~m are com-
'\:e m1' trcpn\tome\, for example. may h.tw rolled mon in soft \t:diment~ in deep water; (> 1000 m).
~d the '~J floor, other. l.lrgc colome' l.ty p.l,,ively. lt may bt• possible to infer depth rclaoonshtp~ in
13 , the btl Crct;tn·ou~. howt·vcr, man)' kinds of fossil bryozoan-beanng assembl.1gcs from colony
U,ltiCl1td.JI or cap-o;luped fret.•-ltving colonic~ fonn, but only rn the broadest ,md 1110\t general
~,. ia • an«-n. The'c an· the domm.mt frce-hvmg sense and 111 conjunctton with other criwriJ, \li1CL'
C'th od ry, \l!CCC\sfi.tlly colonmng lnmt.·. movmg by it~elf the shape of a colony is not an um·qmvoc.u
c· rleJ 1111tk palaeoecolog"~cal inciJcator.
·n1s ne sudt \penes of p.trticul.tr etologtt.tl tntcrest
tpe. tht w•d~,prc.1d an.t~c:~n cheilostonw Cupulc~drw
Ime ncmi.r (rtg. h.Ja,b). for it' l1.1bJtat .tnd c.h~tribu- Reef- dwelling bryozoans
• ot 111 Recent sc.1~ and !Tom the Miot.·cm• onward~
~ul bcm wry thorou~;hly re\carch~·d (Lag.tii.J, l %3). Bryozo:.tns h.tvc been quite signifiLJill .1~ frame
The c-. dt de!pdl, bct\\'cen .tbnut 5 ,md 5110 m, but it builders or a~ 'edm1ent binders in VJnou'> ktnd~ of
4\ J t cummon nn contincnt.ll 'helve' on a sand reefs through gcolog~cal time. In Ordovician to
. h-. tl 111 the Atlantic and E.N P.H:tlic. It can tol- Devonian cor:ll.-.;tromatoporoid recfc; they formed a
C,tc:h tOI!.pCrature' between 11 .md 31 )C. though it subordm.uc p:trr of the reef fabric .md a\st\ted 111
t.·in~ 11.1mulh contined wtthm tht· l-l C. 1\0I..f)'lllC ~nd bmding the sediment. CommonJy thq bndged gaps
muc, between l7 anJ 37 per null~. and allO\wd cavittes to form bclo\\ them. these
been "he btv;ll: h;tve \trong ~cduncnt prl'lerences, so often bccommg tilled with fi ne sediment. There is
tZOJtl coloatl'' ,m: alway5 tound whert.• the p.1rticles some evidence of vertical zonation of bryozoans in
~n: .I · qulrtl t,rr:~in~. foraminili:riJs, glaut.nmtc pdkts some Silumn reef:~.
llC" hlr 11 frnhrrnent') .m.· large enough to pemut set- In the large Pcrmo-Carboniferous algal reds they
1 56 Bryozoons
contnb\lfl' 111 .1 mmor \\',1)'. or on .1 localized scak

lnd
smm·tmll'' mon.• •mport.mth. to the reef frantl'-
work. J·cn<."<.tdlah h.W<.' bc<.·n t~Hmd m life pminun. U !son
)) ~tl'
pro.wrnn~ nul\\ .mh ti-om .1 't<.'cply dtppmg reef t"Jce. Books, treatises and symposia
1'110(
rhey .IJ,o nrt llr Ill p.H<. h J"<.'<.'t\ ,I\ ti-.tme builde~.
L3i~~y. 1·. 1'. (<.·d.) (I'NI) Bt)"Cl7oatre' .Ktud~ et 10$ S2'J-
Modnn cor1l r<.•ei\ m.l} C.IIT) abundanr tauna' lhnta,
of cn~.·nhtmg bl') 07o.ll1,, '' hll"h arc in places ~i~lifi Hulft 1111 clt• 1.1 S,o,u·tc: ,-;, rt·uufi•(llt' .I '/ulll"'' \ atu
c:mr ti-.unc builder' Stronglv built. thjck-walled /',.,,.<t .le /.r I"'"''·Mcm01r~· I. (55~) n•p•Nwu f!Jr- Uryo
lHS ,
Bu,k , C. ( tx=.•J) A i\lonugnph of the h1"tl Polyz
cncru,ter\ ,Jr<.' fi1und 111 rq~mm of turbulent water. Bl~kc,
dw Cr.tg. P.tl.tl'lllltogt .tplunl Soctl't)', L•mdon ~
wlwr<.'a' the more ~.khc.lll' cnbrunorph~ Jj, e in shcl- dL·scnpuun uf ,t hi) ozu,lll-nl h bed) 071111
tcn~d nvltll'\. Tlw l.lrV.IC of thl''e type' have stron"g ll.tyw.mi. PJ . .tmi Ryl.u1d. J.~ . (f'J7'l) Hrtll.<lt -lsu•p 451-
h.1h1t.lt prdi:rcn<.<.'' .1ml will only \<.'td<.· on particular Bry,>~o.J<IIIl,
( 0\\ C'll
'uh,rr.ltes. usu.lll) <."or.dlut<. al~ae or d<.·ad skdeul ~vnopw' of tbt• llrili,h r:.n111.1. Vul. I I. A..-.ult'mt l'lol<.'lil
m.1ten.tl . W .lt<.'r tut buh.:nu~ \<.'<.'ms to be the primary London. ( I~x·· dk•m tllu~;tr.ttwn~;) 1 I '•ll!f,
I bl)·
control nl dl\tnbutHHl "Jtbm the multit~1rious habt- ll.!yw.Jrd, P.J .. Ryl.md . .J S .lnd 'l.•vlur. I'.D. (l·d~)
dl\trJl
tlt\ prov1d<.•d bv th<. coral n:cl and bryozo:ms of var- Bioh!I!Y cllld l'trlao ·o/lttl!t~l/ )' •'./ Hry••~• '<IIIS. Olsl'll & (l
lt \\I h)(
wm type,, though .1l1umt all encn1sters will Aourish Frcdensbm~-t (M.111~ v,du.1hk p.•pet')
f.>f M)
L.lrvo~nod (; Jl. (t•d.) ( I'J7 ~) Ur•r111! tlltd h•>>il B')"<'
tlmmt .1 11 ~ will' I"<.' th.1t 1s lr<.'e of ~uspended sediment. }lllllll
Ac.Kkrnit PtL'"• Nt•vv Ymk (M.mv ong:malpJp
Mo't speci~.·, se~· m tn ha\ c parntular funct:10n:tl () , r-;
,t,111d.1rd n:ti.-rl'ilU; Wtlt-k)
td.tpt.ttwm tl1r o,ud1 lubll.ll~. thougb many of the~e i\1•Kn111
llf\Vnnd, ( ,I' md Abbon. M.L3. (I'J71J) 1d•••ll~
hav<.' not been inn·,u~.Jtl·d 111 det.lil. h 1(>t'(
HI')'•'· ''''''''!>'· Sy,ft'111.1tK' A"ou,1tmn Spt·nJI \ ol
No IJ. A, .tdllllt< I'll'"· I nndon. (M.111y papt•n
L1 m·ood. (;.!' . .tud NL·d,.:n. C (<.·d,) (14HI} R
Deep-water brya:z.oans F'"'" H•T•'.;:''''"'· <. )J,<.:n & ()J,cn. fn:dt•n,borg.
v.1lt1.1hlc p.tpt'f\)
Mmt of dw .thy,s.ll .llld bathyal Bryozoa oi Re<. cm L.mvood, GP • •utd Rt>Wn, ll.R (e<h) (1()7'J) Hu•l;:
\<.'1\ 1re dl~ thhtome~. :\I any 'PeCII."i taken from -;y.<li'IJI,IIh~ <>{ (.(>/,11/1111 <Jrt:cllll.11115 At.J,It'I!Ul
d<.·pth~ of over )I HIll m were att.tched to shells, London (lll)'o20.IIl\, ror.tk 1:-"·'Ptohtc. ctL)
pt·bbh:' .md ntha h.trd ~ur6cc,, .md the} have bt·cn Mt Km ne), f..K . .m.t J.1t k"m, .B C. ( l'l!lfJ) J Br[:.
b•,o/utr.•n (cd, C.T Struttnu Jnd C.P Hughc:s)
found .lt 'llllll' 25% nt' d<.·~·p-\cJ qanom. Rooted
'I ''l''f' 111 fi.J{,I! •lllltlf,~t:)' 2. Unwm H) tllJII , Lt>ndo:L
speCie'>.\\ 1th ltmg rt~llt-lik~.· thrc.1ds .:apablc ofhold-
b<.:'t ')nthL'"' .a-.atl.thlc: l'Xl"l'lknt \.l]uc)
mg thl'lll 'l'l'urdy 111 'l>lt ,~.·duncm. arc now known Muur~·. R .C. (t·d) (l'l'i.') '/'rt',I/J<t ''" /m't!!
to be Vl'l')' .tbund.tnt llld dt\'<.'r'l' 111 decp waters. and Hllntiii•J/,11!)', P.lrt G. Ut}'tiZ•'•I. C<.·ulo~tul S1.1Ct
.lrc dnmut.llll hl.'l11\\ 111110 111. AmLTil' ,t .1nd Un1Vtl'\tt}' ,)f K311s.1~ Pn·s,, b
K.w
Rob1,on. lt. A. (<.•J.) (19H.1) 7n•crti.H ''" lm
l'rlh'ollllll/.~1/)', l1.1rt (; (r<.'\'lsc:d) Bry,a.::v.T, I il/
llltmrlllrti<lll, ( )n/ct <:r''''{ltll"ata, <>rdrr C}'P'•"I
Smcc P a l,ll'IWilil bryozoan gcner.1 tend to be long Geolowral Smt<.'l)' ot A ...enc.l ,md UntVCt\lfl
1<..111\,IS l'rcss, l .twr<.:tKc:, K.m.
r.ll1gl·d .md latt<.'S (.(ltttrolkJ, thctr srratigraphical
Ryl.md J S ( I'J711) Brymn.m~;. Hut..-hilhon, l•
applicatiom m• u,u,11l~ pom Spcncs assemblages
(ln valtt.1ble .1n out !I of Rl·u·rlt .111d fcw.tl hrvnw
w1thlll giv<.'ll (ll'Jl'' H"l' rq~wtt.lll) very useful tor Rvl.tnd, j .S ..111d ll.1ywml. l'.J. (l<J77) Bntllft i
zon.tl plll Pll'l''· lwwl'V<. t. co,pec i.tlly 111 w1dcspread Hr)'•' ,,,,,, l.ovm1p~<.'' nf the Bnu'h Faun.t, V
c.trbon.lt<.' ,Jwll '<.'lhlllt'JH\. 'wmc of the Cretaceous At ,Jlknut Pr,-~, I ondnn (L·xn·llcnl tiJu,trJUl)lll
and Tcrll.ll)' brvozo.ttts 'eun to have !muted vertical Wooll.tnm. R.M ..111d Zint11wr. P.l (<·rh) (I'IT
dhtributmn, .md ·'' tlll'lr ttme r.mgcs become bette( 4 Bl")•••:::oaus. At.tdt'llllt Prt'"• Nt'\\' York. lNu
knm\ n ,11 tlll'tr 'tiJtt~r.tphllal pntcnnalmcreases. p~per;)
Bibliography 157
Individual papers and other references

,'\1.:Kmney. I.K ..md I .1ylm, P.D. (IIJIJ7) I tit· h"m•tc' "r
~l.lllll' J\il''>OLOII C:lltrll\tlllg hr\'OLO.tll\, J>,r/,l('<>llh>f,J.I!)' 40,
>11 PS nd T.1-.lor, P.l) (l'IH2) l'.ll.tcobtoloh'\ md 515-56
'11110< >f lng<' < \'< lll\h>llh hi) n7o llh tnun the Nyc. O.B. .111d L,·mon<', 1>.V ( I97H) ;\1ululunintr
P ocene CM1lhnc: t 'ng nt Snffi>lk P.t/lll'•'"'''/•!..: 25. ~orowth 111 Rtplt>lllllilll•ll'·' h ''"'''• a lit:\\ 'Pt:tie' of
i!tf-54 (\'\'c.-11 "orth r<•.tdmg) cvo:lo,tomc: hl')oto.t. f,,,,,,[,>j f>.lh(>tllt>l.•..:y 52. ~J0-45 .
a \\.C.. ~h Kutlll'), r.K md Ztnutttr, I'.L. (I 'J7~) \chopf. l ,J.:\11. (1977) P.ltll m~ .1nd tht'OOl'' of t:\·nluuon
Br.ozom monn, u),·~: •·srurr,·nt \\ .trcr outkt '>t·•wn· among the Bl)'07o.l, 111 1'.1111 ms o!f l't•,>lutioll (l·d A
185, "'S)-4 (lmpunuH C1lllt cpt) H.tU.un). Fl\t'\'lcr. Am,cc:rJ.un, pp. IS'J 207.
e D {llJXtl) Houll'OIIIIlll'h\ 111 l'.ll.!,•o/lliC bry- ~muth. A.M (JIJ'J5) l'.tl.leol'll\'IWilllll'nt.ll .uulyqs U\lng
.no.tns: !I se m:h fl11 cxpl.tn.tllom. 1'.1/,,, ,,f,,,,f.>xr 6. brvozo.ms: ,, r<'\'lt'\\, 111 \ l.trilll' Hr/,, ,,,,,,,,,c,,,[
~i).-{,5 A;,,;[ym .Jr,>m tlu Ft>ml R. .. ,ml (ed' I) W J. Bownrc.
en, R.. lllll Rtdu,J (I '17.:!) hlllllllllt.tl .ut.ly,,, ofti:n- D W J Jnd 1'. Alhson). ( oC\>Iogt\ .11 '-lottl't) ot I on don
t ll'ilid bl)ULO,Ill ltlll>llle\ [.J·tll<ll•l 5, I ~7 -fl-1 (Sl'l: tc:~a) Spcn1l Pubht.mon No. HJ. pp :!.)I H
J.nu. R. (l'Jfl.\) (.'"1'"[,,./rr,, """"'~'""-'(Busk) porrr.11t of Taylor, P.D. ( 1')7(,) Multil.uncii.Jr gr.m th 111 two Jur.l%tr
:1-) brymo.1n. Jl,,[,,..,,lll,•l<>l!)' 6, 172-217 (Morphology, ryclmtolll.ltou\ bryo1o,t P.r/at't>lllt>lt!I/Y 19. 29H J( 1(,
n. dNributinu ,md ,., o I ''14)) Taylor, P.D. ( I <J7H) FutKttOJl.ll "V:"IIir.uKl' of rontr.ming
1rwood. G.P. (I 1J112) I he lltorpho lot,.,•y .md 'YStl·nucics colony fim11 in two M~:sn70IC l'IHTmttng hl)·n7n.ltK
15
nf \Oillt' ( rct.lllOll'\ 'rtbrunoq>h l'o lyzo,l (Pdm.tto- PaitiC<'.I/CO'(I'ttphy, Pt~lam limato>lo!'i)', Palllt't><'<tli<!I!Y 26. 15 1-8.
J
ronnle) Hull!'ti11 t>/tht Hnri.•li \ l11.•mw t>/ .'\l,rtllr,,/ HistMy Taylor. P.D. (1 '>7<>) !'he lllfl·r<'II<T of extr.tzooid.1l feeding
6, I-2H I. (l:.x~:cllcm lllusrr.uioll\) currents m fmstl hryo1n.w rnlontl'~. l..cth.litl 12, 47-56.
Ill
Kmnry, r K ( IIJ77) htnli1011.1111Hli)'Tl't.I!IO!l l>flyre- Taylor, P.D. and lbd w, R.M (JIJ<J'l) A m·w ,h<·ilmtom<'
Jped bl) nm.ms. p,,/t·,>/or,•f,,l!)' 3. lJO .., bryozoan front thl' Cr~:t.llt:llll' of ludt.l Jnd Eump~:. a
IIC
1\Jnne\, •·.K. (llJH.l) A'<'XU.ll n> nil\' multtphcltton by cydO\Wll1C hollll'OIIlllrph. Jirl{,JI!lllltlf.>I!J' 38, (J27 'iH.
wl fupn~lllJtlltn: .m unport.tm ""'d'• of g.: m: ne longevity
h)
the Ctrhomti.·nlll' hr-.o7o.m !r,/,,, dr< p,,/r,•bit~l.>l/}'
9.35-•H
r,nd
':'OS
f.
alol
ot
o n.
5Ct111
10.
Bra eh iopods
I
Ih
I ,.
tal
Brach10pods arc bentluc m.mnc animals whose sofi. Ccrrain fundamental ch::trJctl'rs .ue common 1\)
parr\ are enclosed withtn .1 two-vaJved shell. They brac:hiopods. They all h.lvt• .1 -.hell of two \,11
have some resemblance to bwalves in that they po~ u~ually fixed to the sea Aoor by means of a 't.tll
~css a hinged patr of valves and feed by drawing pedicle (though some arc n.'tnl'nted, :1ttached
w:tter mto the ~hell Jnd tittering off the food p;lrti- spint•s or !Tee-lying), and ,1 complex food-g~thc1
clcs, but zoologtcally they .tre quHe separate. The organ called the lopbophore. Until recent!).
two valves are ot dtfferent \tZes bm symmetrical .1s in the third edition of thl\ book), brachtOf
1hout .1 m edian p lane, by contrast with the were brrouped in three cla~'>t''>: L mgulata, lnarttl'U1
cqual-~zed but mequtlater.tl \'ah·es of the bivalves. .mJ Arttculara. The-;c arc broadly equivalent to
Brachtopod~ arc tiN t(>Und m the Cambnan and are present Subphyla linb'l.lltfomtea, Cranntonnc-J ,,.
very abund.mt m the to<o~tl rc<.ord, often bemg the R.hynchondlifomtea, rcspe<.ttvcly, in the new
commonest and most ubtquttous fosst.!s in any shal- ~tficanon of Williams et di. (1 1J96). Tlm b.11 lx
low-water dcpostt. In the Pabcozoic they were .1 c~tablished on a cladtstic bam .md tested .1gaimt U.
very important phylum .md, though they were Jec- stratigraphic record. While the onginJl tt·nm
imatt:d in the Penm.m, 'ome genera conttnued a~ now superseded, it is still proper to refer colloqw.d.
the dominant benthm 111 loc.tla7ed areas during the lO 'inarliculated braduopod~' ft)r the 61"\t two IU
Mc~ozotc where tht·y m.1y be v,t,tly abundant. and phyla. and 'articulated br.1d11opmls' for the th1rJ
by tar the cummm1c't fm,tk Although thetr tmpor- detailed classificanon i~ set out 111 sct.tton 7A.
tance h.ts ~mcc dedmed. they .trc, ho\vever, com-
moner toda} tllJn f(,mtl'fly thought. especi.tlJy 111 SUBPHYLUM 1. UNGUUFORMEA (L. C.1m.- Rec.): Br~ tel
deep and cold watL'I'\ Tlm nuy represent a real shtfl pnth on which the valve'> Jn nut hmgt·,l b~ teeth and sock
111 en vironment through ttnH:. Jnd on whorh the shell os duunuphc"ph.tll< 111 comp011
The shallow water\ otfNew /..ealand cont:lln abun- The ~wdodc ts fleshy and mustulJI, .111J t·noc:rgcs bcmefl
d::mt brachiopod\ ( 12 ~prc1c~ of tivc genera), and thts vJivc' or from the apex of one- of thl· v.olvc'
SUBPHYLUM 2. CRANIIFORMEA (L C.tm.-Rer.)
~~ the classic area for rc~earch on living brachiopods.
ltkcw1 ~c lackmg teeth and ~oda~r\, hut wuh
There are. however, m on: divL•rse f.1unas elsewhere: shell. Pedicle reduced or abscm.
the Untish Isle!. have 21 ~Pt'lll'' of 17 genera (Brunton SUBPHYLUM 3. RHYNCHONELLIFORMEA (l. Cam -Rrr
and Curry. 1979), though the'c .trc normally found Calcart·ous-sheUcd brach1opocb tn whoch the nlvcs
111 deeper water.> and arc hcnc:c rarely ~een. hongcd b)' teeth in one valve .md \ockrt\ 111 the other 1i
Over 4500 fcmtl gent•r a .trt• new. known. though pnhcll' ·~ made of~ dead homy m.ucn.tl .md on ..omr
g<'lll'l'4 appears to hAve .otroplued Tht· rhynchondlu
no more tlun lOll an: li\ ing today, albeit these 1rc
.trc much more diverse and JbundJm dun the ltngulu
w1dely distnbmed and tound ar all depths in the \t'::t. ur uannfonncs. They appc:JI at the SJim.• level m the L d
M.my of the fmsll genera I1Jvc been found to be Cambnan; however. d1e main cJ.rly rJdiaoon of th~ rbr.
~trattgraphically useful .1t vanous horizom. t honcllofonnes was not untJI Llw c.uly OrdovtCJan.
FiguN.
view (
ofbro
Morphology 159
7.2) ha~ beeu well chowu by ttW1)' ,tuthof\ a~ a

Subphy lum Rhynch o n e llifo rmeo
typical example of a modem rhynchoudhfonn
Morphology of three genera brachiopod.
It shows ck.ulv ho"" thl· hard pam relate ro the
Mogellonio living anatOlll)', whiCh •~ mdi.tl tn tntcrprrtmg tilt'
Tb rcccm brat h10pod \ ft1~cl11111i,t (Order ubiquiwus tcrcbr.uulu.le' uf the Mc,o70tl·.
Tmbratuhda. ~ubon.kr T l'n·bratelltdtn.l; Ftg; 7.1. .\la,l?cllcmltt ltvc\ cptlaunall'. .•tttac bed b; tt\ pedicle
\11 (b) (d)
e~. rnllo
01
bv
11~ comm1ssut~
nd
)lh
at.l
;he
md ....1h '"
are
ally
\lb-
I.A
ltl (I)
hio-
.:kc:b
tion
aXIS
n dw
~wSIOI
111usde sca1s
!iCliS (g)
l..t'C ): iJnH.mor lobe

.. <11 t'
, ThC' mantle
fo,\11 rudiment
::>mH~s ped1cle
omll'< rUdiment
_QWC'T
rlwn-
:re 7.1 Mogellomo Rovescens (a) upper surface with brachial valve (x 2 approx.), (b) lateral view (x 2 approx., (c) antet"ior
lt<'(\ 2 oppro.x.),(d) tn life position, showing pedicle allochment; (e) internal view of pedicle valve (x 2 approx.); (f) internal view
bochrol valve(>< 2 approx.); (g) larva [(aHfl Bosed on Davidson, 1851; (g) based on illustration rn Percival. 1944.)
160 Brachiopods
adductor muse lP lhnu

CXoll tly
gut dlvenJCUI 1m malle-r I
the pcc
1 hrnugl
opcratm
er~(br:1c
INTERN
lo hg. 7
uon cut
'T\ilntle cavny
tJrbr.Jns n
:ill' "hie
'hdl i'l .
dJductor muscle
\t t rl'LC\
.1ntenor body w~l l lll.lr6'111.
body cavny
,h~ U It i
Figure 7.2 Magellania Ravescens: median section, somewhat stylized. ln.tt hiop
tic tucl<
Jll l\tcnnt
to and 'tone~ .1t dt·pths of 12-600 m in
1 ocks line of JUnction or co•nmissure. rn convcntw' v.alve~ at
Austrahan and AntJrcnc wart·rs. orientation the umbonc\ an: Jocned at the postt· )11111 , for
The rwo ovaleJk.tn.'ou~ valve~ dtffer in 'ize but arc rior end of the brach1opod and. <,mcc the hing. body w
eqUilateral and d1v1ded b, a single plane of symmetry. line j, in front of them, the valves gape at the antt- 'rp.uatcs
In ~tandard oncntatwn the 'upper' valve is ~mallcr rior end. l:.tVity.
tlun the 'lo\\~r· Thae has been \Ome controversy as Each v:~lve •~ extemally ~culptured with faint ribl mantle 1
to ""hat the~c t\-VO valves 'hould be called; the 'upper' radiatmg from the umbo The~e become mo m1c ted
valve IS vanoml) known J\ the bracbjal or dorsal amcnorly ~o that the antenor parr of the Cll!TIJE: hum. T~
valve. the 'lower' ,ls the pedicle or ventral valve. sure ts cn:nulated "'hen: the nb\ mt·et u. Thr {Of} OlC1
The rcml\ 'br.1chtJI' Jnd 'pt•ciick' refer to structure' growth lines arc subconcenmc round the umk tlw lop!
conr.1med wHhm the valves and rhus are more tech- and LkeWlse become rather crcnul.ltcd .mtt'no fhud-fillt
mc.llly correct Furthcnnore. the conventional dorsal where the nb~ are stronger. E.lCh of tht•sc gro 1 his is <.l
or ventr.1l oncntanon " not ncce~arily a life orienta- lines represent<; a record of the former pminon nbbon ~
non; m t3ct the brachiopod, when ftxed by its pedi- the edge of the ~he ll , and ~i n ce they .1re cspt:c ill rht• 111cdi
cle, ha~ both valvt'' verticaJI> hdd or may have the prominent where growth ha~ ce:l~ed for a short lln:t lll\it!L' ur
ped•cle v::~lve uppennmt. H ence the terms 'brachial' the shell retams .1 pcnn:lllent record of it~ O\\ lJ\Jty. 1
and 'pcdtcle' ,lrt· adopted here J~ standard practice to ontogeny. tht· 'hell
de~ib'llatc the valve~ though tht· other system is admit- Th~ pedtclc valve ·~ th~: l::trgcr of the two Jod 'l•ttcd a\
tedly simpler. intumed posterio rly ~o th:~t the m m t convex part rrom th<
it can be ~een ti·o m the upper ~ urf.1cc. This part lophophc
EXTERNAl MORPHOlOGY the shell is the palintrope. The umbo is perforo11 p.ualld £
Tht· ~uperticial re~embl.1ncc~ of such brachiopods as by a round pedkle forameo through whtch uh.1. Th
Atagl.'ilcmrn ro Rom.m oil l.1mpo; ha!> long been drawn. pedicle emerge~. The httcr i~ a ~talk of homy tt~lllt rntb th.
hen et• the vc:rn.tt ulJr tt.·rm 'lamp shells·. The two cemented Jt 1ts d"t.1l end to rock\ or pebbles, .1nd (IlTH nt~
valve~ arc unequall) bKonvex. Each grows from the lt the brach10pod i~ raised above the bott ;~rr.mgcd
firsr-formed p.m. thc protegulum . which later Between the umbo .111d the inner edge of tht' 1 p111y too,
become' p.lrt of the umbo. lt grows Ul a loganrhnuc ts a tnangul.u cavil), the delthyrium , do,eJ bj hltt red"'
~p•ral, as doe' the 'hell ot a bl\:aJve, .md at each patr of delridial plates. They arc markt'd The fooc
growth tnt. rem em new ,hell m.ttcnal is accreted straight. p:~ralld growth lmc~. tranwcr<.e to the h ment in
round the b'H)\\·mg e-dge. The vah·cs meet along a chiopod'~ plane of 'ynunetl) dlon~ the
Morphology 161
Thoujo;h the commmutc:~ of the two valve~ fit The mouth is situated m the .mtenor body wall; 1t
ttlY nearly all the w.l)' round the shell, the lead~ to a small gur wich an ot..,.oph,lgtt~. \tomach
'Jlill~r bradual valve has th umbo tucked JUSt mto and blind-ended intestrne, but no anus. There arc
mt pedtrle vJIVl' Jmenor to the delthyrium. abo digestive gland~ or dJVcrncuiJ.e .t~~octatcd w1th
ouj!h dm reg~on ntm thl· tranwer.e hinge axis the gur (cf. Fig. 7.3c).
per~nnt-: on mtern.1l teeth (pedtde \'.live) and sod,- Excreta arc voided through the mouth into the
(brJchul \~lh·e) mantle caVJty and di~po,cd of by the exhalant cur-
rent. There are no other pc..•rforauonl. in the Jntenor
NTERNAI. ANATOMY body wall other than the patred funnel-~hapeJ
fJ);. 7.2 the: 1ntcrn.1lor~Jns Jre dt~played m a sec- nephridia ('k.1dney!.'), wlu<.h remove 111trogenou'
ll cut through tht pl.me of 'ymmetry. These waste taken up by wanJenng cell\ (coelomocytes)
llglll~ rdJte to structures made of calcium carbon- and also act a~ passage~ f(Jr the escape of gametes
" which lJil bt.> ~een 111 the: wparJtcd valves. The from the gonads. Sometime~ the latter become ~o
tilt\ 'ecn:teJ by .1 cellular epithelium which swollen with eggs or spenn a~ the brecdmg sea~on
.rett:~ ca.lc 1rcous tn.ltcn.d m.unJy ,lt the valve approaches that they can expand into the mantle
ITI.,~n Tlm cpithdium underlies every part of the canals. These canals arc tubu lar branching exten-
h].lt i' rwo l,•yc:r. th1ck 111 the .m tenor part of the sions of the body cavity (codo111) which run
,hiopod. where the mncr l.lycr form~ the man- between the mantle aud the inner epithelium.
,, nclosmg ,1 l,trgi.' mantle cavity. Towards the Within them coelomic ftuid circulates, being used
oomor end of the o;hcll the mJ.ntle~ of the two ma.in.ly for respiration. In dead ~peciml.'m of
•:eo, .Jbntptly leaw the tntenul valve ~urface and Af,l~c'flallia, divested of sof'r tiSsue. the \car; of these
, fom11ng J 'mgk• ,hec..•t of tissuL', the anterior canals are clearly seen on the Jm1Jc of the ~hell
kldy wall, \\ htc h cnmc:s bet\H'Cn the valves and The only other organs tound 111 the body cavity
·.-nte' the mantk l.tVIt\ trom the..• pmtenor body are the ped1cle and related stn1crure~. and the mus-
rity. The only organ t:OntJmed wtthm rhe cles for opening and clo\lng the ~hell. The ped.Jcle is
tk ravitv j, the lophophorc, which Itself i~ a cylindrical stalk having a duck extemJJ cuticle,
~tcd ''ith a ronnnu.ninn of till· mantle epithe- within wh1ch is a thin ep1thehum and a central core
Tht, i, the rn.un ti.)od-brathcnng and re,pira- of connective ussue Though the sheU I\ tixed to the
nu:(h,umm of the br.td11opod. In ,1/age//ania pedicle, tr can be tuml·d in any d1rl'ct1on by two o;ers
lophophore h.t~ .1 hydrmtatically supported of adjustor muscles near m ba~e 1 hese allow the
-!illed c.tndl (the brachial canal) a' i~ axi'>. upright brach10pod to swmg 1nro or away from the
L\ ~upported b, .1 long, loop-~haped t:alcareous current. Frequendy the ped1cle split<; into several
n (brachjdium) '' hll h '' 'Ynlllll'tncal about srrands for attachment (cf. F1g. 7.3e)
median planl' .mJ ,Jtt,tL hed .It m·o pomt' to the Brachiopods open and close the shell mmg two
:k of thl' bradual \ alvt• ,lt thl! re.tr of the mantle sets of paired muscles. The adductor muscles
VI\ ty The bradmlium !II,Iy rem.tin ,lttached to which close che shell are ana logous, though not
,hell dlter death \\-lwn thl' mvc~tmg tissue has homologous, to the ndductor~ ofb1v:1lvcs. They join
I~ UW 3\\':ty, but lliOrC ottell It is not pn:served. the two valves somewhnt obliquely, and their points
of .1111 the strong br.ll ludtum-supportcd .txi\ of the of attachment arc both anterior to the hinge axis, so
of 1ophore there ~pring .1 l.trgt• munber of slender chat when they contract the shell must close. By
:ed alld ftl3menL~. Tht>sc ,1re \tttky and lined with contrast, the diductor muscles which open the
:he: The beating of tht·se c.ilia ~cnerate~ the cur- ~hell have a quite different line of action, with
lll". l\ that bnng 111 •.md cxh.tk• water Inhalant thejr bases on oppmitc <;ides of the hinge. They
by Tltlm come in latc1all>, .mJ ,ill tht: tilamerm are are attached to the pedicle v:tlvc JUSt outside the
10\ gcd so as to prO\ tde .tn effective net for trap- adductors, but they arc fixed tu the brachia] valve
.lvc "' iood parncle'; tht:y ,tre 'tramed ofT wluk the by a calcareous bo\\, the cardinal process. on the
y l d water IS cxh.1leJ .mtenorlv m a smglc stream. posterior side of the hinge a•m If the d.Jducror
bv food parudc~ t.tught are pa<.,cd down the fila- musdes contract while the adductor; correspond-
>r:a- ! m a mucu~ bdt to a food groove running ingly relax, the cardmal prorc\~ ""lllb"i downwards,
, the brachial axt\ and \O m the mouth. and as the rc\t of the shell lies ,m tenor to the hmge
162 Brachiopods
1•1 (b) (cl Pet

ftH
:~dd
and
The d
the pc
more·
throug
ontogc
n npt
let 1111gr.ttc
gtllW\,
the gro
ltatcJ •
Ill IJI,III
brat hid
r.ardina
delthyri~
(f)
notothvriu
(Ill
nccesso
Figvre 7.3 Terebralvlina rehlsa (Rec.), Firth of lorne, Sco~and: (a) interior of adult ( 18 mm long) brachial valve (x 3); (b) enbf
ment of lophophore (x 6): (c) digestive diverticulae (x 100); (d) juvenile brachial valve showing alignment of punctae (x 25): (e)fd
de showing separate attachment strands (x 75); (f) enlargement of growing edge (x 50); (g) lateral view of pedicle foromet
complete juvenile shell (x 70). (Photographs reproduced by courtesy of Dr Gordon B. Curry.)
the valve' g.tpc opt·n by a few dq,'Tces. Feeding may the brachial valve, but thc~c .tU bdong to the Ja
then beglll. tor mmcle' since the lattt·r div1de m two. adductor r
These two 'l'ts of musdes. and also the adjustors, Separated valves from wluch the soft nssues scars
leave well-ddincd m uscle scars imide the valves, been removed thm <.how the following charact
vt~tble m J\OI.Hed valves of dead individuals and
someume\ m fos,tl v.tlves also. Of the scars m the Braclml (dor<.,ll) valw. Sockets. cardmal pr diduc
pedtcle valve. the larger outer patr are diductor adductor muscle ~car<., bradttdwm, mantle C)
sc.u-s, .md tht· smaller inner p.tir the adductor scars. there may al~o be a median septu m or
Firwre 7.4
There arc ltkt•wtsc rwo main pairs of muscle scars in 111 the plane of wmmc:tn ncil view of
Morphology 163
l'td1dc (wnrr.1l) v.1lve. P .llmtwpl". tl'Cth. pcdidl" ;\.la,l!rllallt<l rs an example of a br.1cluopod wnh .1
1 rJmcn. drlthyrium tlmt"d by ddnd•.1l plates, non-strophic sh eU. The hinge J..'\.1S paN's through
Jddurtor .md dldUCtOr mmck \Cal'\, aJjU\tOr ~Cars the teeth .md sockets, v..·hich arc 1t~ only fukr.1. The
and nuntk c.uub. sheU edgt·s at\pcent to this arc curved and not COIIlLI-
dl"nt with the hinge line. Mmt livmg br.1ch•opods
1' tklthynum, InCidentally. i~the ,lncc,tral site of (e.g. the Terebr.lLulida, TcrcbrJtdlida and Rhyncho-
pnlh k toramen. In hr.1Ch10pod~ thar retain the ncllida) art' of tlu~ type. Many groups of Pabco:zoic
111 'pnmJtiVl' '}'Stem' the dclthynum 1s open and braduopod:. (e.g. the Orthida and Sp•rifaida), how-
ough tlm the pedicle cmt"r~t''· In rhe early ewr. h,tVl' strophic sh ells. ln these the hinge hnc is
tog1n} of \I,J,~cllania the pedldt•" til'\r located m straight. oftt'll l'Xtcnds the full w1dth of tlw \hell .md io;
'fltll ddthynum. but the ti1r.nncn ~raduall} cou~'odcnt \\ 1th the hinge ax.i\.
tt to lt\ .1dulr po.,•uon ·" tht• br:1ch•opod
\\ ..md the ddrhynum do~t·, utr bchmd 1t b} Visbyeflo
!10\\lh of tlw ddt1<h.1l pl.lte\. Tht• ,hdl Jnd a'\so- I'rsbyt•lftl (Order Orrluda, Suborder Orrl11diru) io; a
tl'<l \ITUthlrl'S 111 the Recent '/ ndllrlfulma. smular Silurian bracluopod with a strophic shell (Fig. 7.4).
111111} \\;IV\ t0 J\fcl_~rl/,wia, rh011~h With .l shorter It belongs to one of the earliest o( all rhynchonel-
d11dium, .m: shov. n in F1g;. 7 .J. liton11 br.tchtopod stocks to have evolved. the
mterarea
/
umbo
- hinge I
commissure
pedicle valve
(b)
teeth
cardinal process
large-
] pedt
nen tn
lduc-
h:l\'\:
:CT"\.
-
llductor muscle
dental plates
(below)
::>CC\\,
anak (d)
tition •111e 7.4 Visbyello visbyensis (Sil.): (a) complete shell, upper surface; (b) lateral view; (c) internal view of pedicle valve; (d) inter·
view of brachial valve (all x. 5)
164 Brachiopods
OrthtdJ, fir.t lutown from the Cambrian. The two deeper and convex cxtcmally The 'hell j, \troph
valves arL' d1~sumlar m ,tze and convexity. The large and the hinge extends the full w1dth of the 'cnucu·
ped1de valve IS very deep, the brachial only slightly cubr shell, being bordered by very narrow intrnr·
plano-conwx. but both have the ~ame surface ea~. There i~ a large cardmaJ procc~' which bloch hang a
sculpture of tine nb~ or costcllae radiating from the rhc ped1clc opemng whc..·n the valves arc cl<M
umbo ,md n.trvlllR ottt\\Jrds laterally ro the com- Externally the valve~ have .1 \C ulpturc of Wl) t~
ml~~urc A pronounced umbo characterizes the rib~. with somewhat more pronounced ~m~c nb
pedicle valve .•mtl'nor to wluch ts the large open spaced at interval!> and div1dtng the shell into rJ
delthvrium Without ddtidtal plates. Tills ts flanked segment~. There arc no teeth. The\c have been I
by a tn:1ngular interarea, wluch fonns a flat, slightly in the evolutionary hmory of \trophomcnidc..~ ~
slopmg shdf whosL' antrnor t>dge is the straight their function ha\ been taken 0\·Cf by 11lJ11} ~rm parv
hmge lmc The tmerJrea ha~ doscly spaced growth denticles whith run aJong the hingt· line, tho~e 1
line~ p.lr.lllcl wtth the hmge. If the valves are sepa- the rwo valve\ interloc:.king. The ventr.tl mu,,
rated, the l,1rgL· rrcnul.m:• teeth are vtsible, situated field has a bilobed diductor \Cat Within which tl (a)
anterior to the hinge lmc where the dclthyrium smaller adduttor scars arc located. Jn the braclv
reaches its grcJtl·~t wtdth. joining the teeth to the valve there is a large V -~hapcd cardinal proct
floor of thl' pcdick• valve arc the vertical dental which prqjects posteriorly carrying the point "
p lates, which cnclo~c a deep umbonal cavity con- attachment of the dtductor mu~cks well to the re:.
tinuous with the delthymtm. Flooring this cavity of the hinge . This is necessary in view of the ve~
and ext('ndmg somL' way Jntt:norly arc the recessed narrow body space instdc the shell and the rc~tnCI<
muscle '>car<.; the d1ducror \Car\ arc outside those of line of action of the mu\clcs. A scncs of vcrtic
the adductor' .md tht' \.\hole muscle field is plates supported on ::1 raised pladonn (bema) h~
hcart-\hJpcd. Tht. Jdju,tor \C.lr<. arc not recessed and down from the roof of the braclual valve, Jlrn
arc b.trcly Vl\lblc rouch1ng the floor of the pedtcle valve when d:
In the brachtJI valve there j, an mterarea, shellts closed (F1g. 7.5b,d,e). Pre~umably tlm MI\11.
though ir IS narrO\\l'r than that of the dorsal valve. cure was m some way connected wuh (el
Bem:cen them thcrt• " J 'mall tnangular openmg, lophophore whteh m:ay have adhered to 1t. lJr
the notothyrium. wl11ch he' dtreccly opposite ro elm lie the adductor sr.trs. An mterestmg feanl:.! Figum 7.~
the delthynurn \O thJt the pedicle opening is of the instde of the 'hell are the numerous small rr. ofbrochiC
diamond-~hapcd and i' fom1ed b\ the ddthyrium jecnons (taleolae) wh1ch .tre t.alcite rods obl.tqcd-
and nototh} nu m together. A btlobcd or trilobed sct in the shell and rad1alh arranged p.aralld wtth
cardinal process proJClt'> pmteriorly and almost costcllac. These arc rod-ltke umt~ lodged \\;tllln
filb the nototln num ''hen the valves an.: closed. shell irsclf and projecting tntcmilly, and the) a ot form
The adductOr \t.,m .1rc VC!) large. Supports for the found only in the Strophomemda. ht,•m1L'a
lophophore arc n.:prcscnted only by .1 pair of diver- The dclthyrium and notothyrium arc • tlll.ll ran
gent brachiophore bases . which arc simply a pair blocked by the large cardinal proces~. and the 1m Jcpart t<
of oval knob' inro wllt(:h the ~ockcts Jre recc~scd; supra-apical for;lmcn is often ~ealed. If there wa 'hown 11
nothing elst.> IS lu10wu .1bout the lophophore. In pcdklc for this strange, thin shell, it cannot hJvr h lt.\1 geno
both valves rherc arc mande canals of distinctive a supporting fun ction and the apparent absencr ~OIIll'Pc
fom1 The costcll,at.· .m.· lltOrc pronounced toward' adjustor mu~cle ~c1r~ \uggcst th.at it wJ~ .tb~, l k ht CXt<
the t.O mmissurc '" th,tt they interlock along a Individuals mu'l thcrcton: h<Jve hin flat upon bt.tchiop
crcnulated edgt. se.1 floor (ambitopic h;lbit), .1s ~~ generally comm T lw n
in srrophomenidcs, which lmt thl'ir pt·dicle~ du the1r con
Eoplectodonta ontogeny. th on th
Etlplurodcml<l (Order ~trophomemda. Suborder <lthcr fc:..
Strophomemduu, ftg.. 7 .5, 7.6) is a Silurian genm Preservation, study and classification of h111gc, de
in which, as 111 m,my brachtopod~ of tlm order, the articulated brachiopods to bt: me
t\VO valVe\ are COtllJVO-l'On\'eX The brachtopod 'pccit'\ c..ksrnbcd above, one ~P(' ic, },
They fir umdt. om· Jnother. the bracrual valve and rwo fo"1l, belong to \ttccc\\ful and u· nnr•~t;-tl f(lund 01
havtng a concavt• L'xtenor. the pedidc valve being groups <1nd t0''e some..· md1canon of the vart~ A111ong;r
Morphology 165
cardinal interarea of brachial valve

lnterarea of process cardinal process
pedicle valve denticles
brachial
valve
parvicostellae adductor
muscle scars
be ma
dethyrium (b)
dentlcles
of ~~~~~~~f.i·•:P'l,:P~~·~
!lf ~r~
rr
ed
(c) ~
\
bema
at vertical plate
gs
se brachial valve
~
te
ldddidtuctor} muscle scars
a uc or
(d)
pedicle valve
~ 7.5 Eoplectoclonto penkillenSIS (Si I ) (a) complete shell upper surface; (b) internal view of brachtal valve, (c) poslertor view
{brachial valve; (d) verttcal median section, showing disposition of vertical plates; (e) internal view of pedtde valve (all ' 6)
fonn and functton 111 Subphylum Rhynchond-

•nnca. Yet tht ) h} no mea m show the full struc-
ly :ur:tl range, eve 1 though mmt brach10pods do not
ill• d.:p.Jrt too rad :all} from thl' kmd of morphology
;1 uwn in thc\C. Ncvcrthde~~. rhere lrt: some pecu-
d iJr glmcra ,ltn011b'llt the Br.tchiopnda , especially
)[ o()Jlll' Pcrnu.m Procluctida wh~eh arc ~o modified, at
(. 111 cxtcrrully, that they arc h.trdly recognizable as
LC rJchiopods.
n The majon[) of bradunpods c.m be ass1gncd to
hctr correct order and U\tl.llly rd"t·rred to their fam-
~ on the bast' of l'Xlernal morphok1gy alone, but
thl'r featu n.·, chc luphophort.'. muscle scars,
hinge, Jenhnon. cardinal prnce\\e\, etc. - may have
ID b( used 111 tdcnnt).Jn~ a brat h10pod at genus and
g '!"C'nr~ levd,. Unfimun,ndy the bradudium is
Figvre 7.6 Eoplectodonla penk,flenm (Sil.) from the Pentland
lt lllund onlv in \Oilll of tht· many fo,,tl groups. Hills near Edinburgh: an internal mould showing the pedicle
:y ~ong.r Palaeozutl ,mfel'\ the brachidium is valve with mantle canals (x 8)
166 Brachiopods
still retJms it~ shdl. the mtemal \ITUltures ma\' br rublwr
present onl} 111 the \pmfcrid.1. Atrvpld.t, Athynd.l
and Terebratul iJa, and poSSibly \omc Pcntamem b studted afi:er removal of the ~hell by and or b) nl- ,IIIOWt'
,,(this
(Fig. 7 7). cining, t.c. by burning off the Olllcr ~hell so th.url11:
mould m matnx 1s vt~iblc. dl) lll,l
In the'e th~ braclmlium nuy be pn:,crvcd h'r
M.ul) Pal.leozmc br:ll'htopod~ .tnd son~t· morr b'T<iphlt
mmn.U im.rust,nio n, tl the ~hell 1~ nm mfillcd, or mally j
buned in nl.ltrix wht:rc tt 1s. In the latter case tht· recent ones may be pre~t·rved m stltstoue or lfllh-
structure t.m be revealed by dt'"ctiuuo r sectioning. \tOill' from w lurh all the c.1karcou~ matcri.tl, llldud·
ing the shell, ha~ been lcachcd awav hy Jnd! Majc
Many rhynchond ltfonn bnduopod s are pn·-
\erved with thetr shcllm.ttcr ialtntact. ~lllCI.' the <;hell b>roundw.tter. 1 he result.mt mtcrnaJ moulth nf!Cll
.appear very different fi·o111 mould~ of the cxtnn Forn
is calcitic it ts not sul~lt'l't to tht• potcnti.1l dtagcncric Br.~chic
rr.msfonna tton of an .u.1gonit1t shell. 1 he cxtern:tl surface. Hence 111 the E,,p/cctodonttl ~pccinll'll\ illu-
morpholog y of ~uch ,hdh j, oltLn \'Cl) well pre- rratcd lll Fig. 7 6 the umhon.U cavlt)' of tht• petit lllh Ill''.
valve seems very pronounce d, the nuntk t.Jttl I he ul

scrvt:d and, provided rh.lt rhe) Ltn bl." t1·t•cd cleanly ,ftdJ j,
from lhc m.ttrix. may rt'<ldily be l'xamincd. Since ir grooves ,Jrc represente d by ridge~ and tht• lliU'rlt
tield~ are in neg.mvc rclieC ln the brachial -.:ah~
dJtlcrt'l
i\ very oti:en the mtenul morpholog y that is really At any
diagnostic, 1\0JateJ V.liH'\ arc uf parricubr value. If Visbycll.t (Fig. 7 Ad) deep mdent.mo m would n:rrt-
\t:parated v.1lvc~ do not occur m the fauna, 1t may he sent the brachiopho re ba~c~ .md the cardimlJUOll."
pedicle
llt:lcs,ary to undt·rt:~kt· \t•rial gnnding tn order to Much of the relief reprcsems, of lOurse, \'Jn.ltHllll valve
rcwal mrernal smlt tun:. A parallel gnnding in tludmes~ of the ,hell. The tntcmal 'trut tm
nud1ine th.lt can gnnd to ti.\.cd d1stancc' IS med for ~uch ,l\ the mu,de scal'li, .trc ottcn ~trikmgh 1\
rhi~. but c.H h mccc~'iw ~:o>round lace mmr be pho- preserved and c:tn be ~rud t ed 'vv:ith tmnimal pn·pJr
tographed or drawn hl'lorc rq,>rmding. 1 hough rim tion. lt is in f.lct very often easier to work wuh
1~ tunc con~unung, \lll h scnal 'ections .ut: often the idcnttl} brachmpod s from mtcrn.tl and Oll
only ~ourcc nf infonn.llto n on tntemal morpholOb'Y· moulds than tt) ll\t' speumcns with the calnllt jh
A complete uHenul rt·construt uon C.Hl be prepared preserved.
!Tu m them by bmldmg a ~\ ·'" model or. more Furthenno rt'. latex r~.·phcauon alloW'; 'po'tU'
rJ~miJy, by l"nmpmcr 11mdcllmg. to be t.1ken whtrh show .tU the fe.ttun·, of,h\:llm
If the br.1rh10pod ts prewrved in a silry lltatrix bur phology just ,,., 1hl'y were 111 lite. Ill' re .1 th111 f1lm
(a) Atrypll (b)
(c) Spirif~r
(d) ~
Cor
~ . ~)i~
'-- ~,,!_V
~
below ( 1 251
figure 7.7 Sp1rol broch1d1a - their form and fvnclion: (o) Alrypo reticuloris, intenor of broch1al valve from with shtl r
fgure 7.8
some, vertical seclion showing sp1rol brochidio with unRitered and Rltered water; (c) Spirifer striata, stliciRed specimen expose spa
~leering system
tially removed, showtng spiral brachidia (x 1 ); (d) same, vertical section showing laterally directed spires w1th (Sil) in fro
reverse of that in Atrypo.l(a), (c) After David~n. 1851; (b), (d) after Rudwick, 1970.) Composito
rec:t•morgin
growth ved
Mo rpho logy 167
rubber ~olut1on 1~ poured on w a '>hell surt:lce and rcsol\'ed inro radial, ant('nOr, btcr,tl and vertical
JUnwed to dry; rhen a !.Hex block 1~ built up on top component.~. If all of these keep pace throughout
t dm from ~ucccssivt.' d11ch r l.Jyt·~. ami when fully development the result Will bt' a rectimarginate
dn may be 'rnpped otT. lt 1\ hkt· producmg a photo- shell. i.e. one with a planar comm1ssure (hg. 7.8f.g).
nphJC print from a ncgatin·, .md tht result IS nor- On the other h.md, 1f the vcmcal component
~mUy Jmt a' cl.mt)'1ng. grows more rapidly than the other\ there w11l be a
localized growth anomaly. resultmg m a vertical
Major featu res of brochiopod morphology deflection of the comml'i\UfC.'. Wht'thcr the deflec-
tions arc median, p.ured or \t·nally arranged (Fig.
Form of shells 7.8h) depend. on their lot.Jhcy, wherea' their ampli-
Ur.JchJupod ,hdls, ·" nott•d bd(m:, grow by accret- tude 'and whether they arc -;harp or gradual is a
ml: ne\\ nutcn.1l !Tom rht· m.nltk at the valve edges. producr of rhe rapidity of changt' of rcl.1tivc growth.
Th~ ultinurt· l(mn of .mv dcvt·loping brachiopod Relatively few shell ~hapc' have actually been
'hdJ i' a produc r of rht• rt'l,ltlvt• growth rate~ of the adopted by brachiopmh, prc~umably for good func-
Jitfcn:nt part' urthe V.llve edge\ (Ruclwick, 1959). tional rca~om. Biconvex brathiopod ~hell shapes tall
Al any pomt of tht· v.tlve margm, growth may bt: within a t1i rl y narrow actual ran~c, ouL of a much
pedicle
valve
ped icle
fora men
(b) (d) (el
Kirktdium Cyrrta
(h )
med1an plane of symmetry
••· antenor
··......
vertical
rad1al
Composits
b) fY;..-e 7.8 Srochiopod growlh and form; (a) Kirkidium knightii (Sil.), lateral view (x 0.5); (b) some, with external shell removed to
1r· !JOse 5POO<Mium and cruralium, (c) some, in section, with body cavity enclosed by spondylium and crurolium; (d) Cyrtio exporrecto
,e \Sll in lrontcil view and (e) C. eJ<POfTecla (Sill in lolel'ol view, sh0w1ng g reatly expanded pedicle valve interoreo (x 1.25); (f)
~ilo (Corb.) on othyrid resembling o lel'ebrotulide but with spiral bra<hidium; (g) radial and concentric growth elements of a
'!Cimorginote shell, showing radial vectors, concentric zones, median plane of symmetry, commissure plane and hinge axis; (h)
pd. vecto~ operating at any poonl on the odge of a shell. [(fHh) Modified from Rudwick, 1959 I
168 Brochiopods
broader r.mgc: of po~sible ~h.lpc,. Many theoretical longed m to hollow or soLid spmes. One funwou ol
shl·U ~hape~ thJt rmght hJve heen med were never, such \pine') appear; to have been for anchorage Tht
in t.h.t. adopted .•md the shell type~ that can be IJrge PrtJductus has many \pmes on the base of the
obwrvt·d m nature ~eem to reprc\ent an optimiza- pedtcle valve which seem to have had thi' funcnon.
tron of are.tl ;md volume c nmtr.unts (McGhcc, and the thin-shelled strophomcnide Clto11rtrs, \O,.hJCh
JliHO). Furthermore, m.1ny of the shell shapes th.tt Iw. ~p in e~ along the h.ingt: mar~n , could lOllWV·
art• timnd m nature. even rf they depart markedly ably have used them for fLxing melf upnght on th<
from the bJConvcx norm. tend to appl',tr over and sea Aoor wit.h the spines held vertically. llut m th
over ag.11n 111 unn:Lltecl .. tock- through time. In rhynchondlide Acarrtltothiris the spmes of the ped1clt
such h omeomorphy. the n· .. ull of convergent and brachial valve~ when tlrsr fomted at the edge
e\'olunon. the descend.mcs of ,J ddim·d stock come the m,mtle were hollow and luve been mrerprett(
ro h.n:e <.hell 'hJpc<> (or mdeed other character>) as <.emory tubes for exteudmg the \emtttve np oftht
dmd\ reminr~cent of rho\c of other and separate mantle well away from the body. another kmd o·
~'TOilp\. eJrl)' warning sensor (Rudw1ck. !965b). Th~
C ,mrpomn (Order Athyndtd.t; Fig. 7.Rt~ 7.16) is in spines, uuually holJow, became tilled up With cak t
l''tem.tl appearance strikingly hkc mJ.ny terebrar- a~ the mantle edges migrated away and ewntu•l
ttlide~ nf the ~Jmc and ditli:rl'nt ages. Both are took up an anchoring functron. External omamen-
non-strophic, though the m.1jonry ofspiriferide& arc tation in mme silicified br.lchJOpods LS ,Jwwn 1,,
strophll. The only real c'tt.·rnal d1flcrt·nce is in the Fig. 7.Y.
~t) ll' nf perforation of the umbo, though the internal
mnrpholo~') I'\ r.ldteJII) di,,tmll.!r of cour~e. This i~ Microstructure of shells (Fig. 7. 10)
hut nnt: l'X.unplc of 1 ub1quitou' phenomenon. In all rhynchonclliform brach10poch the 'h~U
rhl re an: isochronous mJ beterochronous multilayered, and the vanous layers c.tn l1\U.1lh be
h omcomorphs (t.e. pre,em .u the \Jme ttmc or at d1mnglllshed m fossil brach10pods :1' well a' m lm~
di!Tl rem urnc~. rc,pel nn ly). anJ they may occur one\. Shell structure and development Jra, b:t>
eitha \.\ ithin the ,,tmt• taxon or m d1tli:rcnt t.1xa. extemtvely studied in the R.eccnt rhynchot~dlt,
Sntnl· br.lclHopod,, wd1 .h the rhynchondlidcs Nt~tasaria 11(1lricans (Fig. 7. 1Oa), which st•rves Jl
.u1d 'pirifcndc\, luw pronoutKl'd serial zigzJg ~tanda rd model (Williams, 1tJ6A). There an: tlu;
deflecl1om all along the .lnteriur part of the comrni~ sbeU layers: an outer non-calcareotl ~ periostracum
~urc. "ith a concave c~.:1 tral s ulcus on the brachial a middle calcareous primary layer and an tnnCJ
v.tl\il· fitlmg mw a pron lllll cJ fold n1 the pechcle secondary layer of calcarcom and inorgantc mJt~ F•gure
vJIH'. Such \tntctmc' prubabh re,ulted from differ- nal. morph<
l'ntl.lllon of !,.'fO\\'th rate' .tt loc.tlizcd site;, not only Wtthm the periostracum there arc three rrum OfOO W
sptre b
111 the \ erncal component\ but m the radial ones proccmaceous layers which underlie an outu gcbb- o 1uver
~~~~' Thl",e 'h.1rp .md ~cnall) arr.mged ztgz.tgs gave ,\ nou~ sheath. This sheath protetts th1.· growmg (l!£t $howtn
rclo~m cly much b'Teater eflelttve length of commis- of the ~hell and is the fint elcmcm to b~ tonned, bu: eh a roe
betng gc!Jtinous it is soon rubbed otf and does nn~ tnt ron
'urc over which food p.1rudes could be taken in. for
no cxtrJ gape. When the shell g.1ped slightly, small ClCtend far beyond the edge of the ~hdJ.
parttde~ could be drawn 111 while large and harmful T he pnmary sh ell layer beneath the pcriostmm
\Jnd grams were excluded. Only Jt the sharp angle is of rather structurele~s crystalline c:llcite. where Jll.lllll
or the llgzag could larger p.mtcles enter. and these the very distinctive secondary layer comtst\ of clou- grnnv
Jngles are nonn.lll) prmectcd b) setae (which .1lso g.ltcd calcine rods (fibre~) \'.lth rounded tn tiv~ ;
Jl t .l\ early \\ amtng ~emors) or occasiOnally by mclmcd at about I 0° to the ,bdl ,urface and lmw re pt
~ptue,. The told and 'uku' '}~rem ot rhynchond- a regul.tr sy~tem of scackmg Jnd J charJctcn~ h )\\,lf
hdc:~ 'c:etm to be lnsmmJc:nt.tl m separating lateral tr,lpezoid.ll cros~-~ection. Although the pnnu.; la} l•r;
tnlulant currents from a median exh.llant scream. layer ts of constant thicknc~~. the ~econdary U\tr ~l'CII.!t
~ptnl~nde~ ,1lso have .1 ll1l'dt.lll fold and sulcus, but keep~ on growing throughout the hi~ of the br oft lt ~
thcv are rebtwely smaller and rarely ornamented by chiopod and so is thickest neJn.:st the umbone' tdl h
ztgz,lgs ,md prob.1bly mark the exhabnt system. A ~cction th rough the growing edge of the shcU .1ln:.td
T be co~tell.1e ur nb~ m.ty sometimes be pro- shows how these several layer.. arc secrt•tcd Th lit' t.••
Morphology 169
r
y
l-
n
IS
>e
lg
·n
Je
a
.e
n.
er
e- hgvre 7. 9 SiliciRed broch1opods frc n the Viseon (L. Corb.) of County Fermanagh, Ireland, showing details of surface sculpture and
'IOI"phology: (a), (b) on adult shell of the spiriferoceon Tylothyris laminosus (McCoy) viewed posteriorly (showing high ventral inter·
lO :reo with clelthyrium almost completely covered by deltidium) and dorsolly (x 1.5); (c) Cleothyridino Fimbriata (Phillips), on othyrid
101re bearer, showing port of the spire and hovmg typical spinose lamellae externally (x 2); (d) Productidina margarilaceo (Phillips),
:I-
c llf'enile specimen of o productoid, showing the pedicle valve posteriorly (x 7); (e) Dosyo/osio ponicula (Brunton). on adult shell
ge !-cwing two·directionol spines on the pedicle valve (x 1); (f) Overlonia fimbrioto ( J. de C. Sowerby), o young pedicle valve with
ut dicroderistic interdigitating ridged and spine poHern (>< 2); (g) Dasyo/osio ponicula (Bruntonl. on adult shell with o well-developed
Ot ~reo. (Photographs reproduced by courtesy of Or Howord Brunton.)
m
:as n.mcle, which setrete~ the shell, i' tnfi1lded inro a from tts upper surface, embedded in .1 protctn
n- 11.0ve under the -;hell edge; here there '' .1 gen e ra- cement; the~e then grO\'- together and amal!-f.~m.ltc
:h, ti\·e zone from \\ hich ,111 the ~hell-\t'crcnng cells as the aystalline prim.try l.1ycr. The cell then ~wivels
ng ·t produced. As e.tth ne\\ cell ·~ tom1cd tt moves round an a.xis of rotatton. ~o that its dtstal surf.Ke
tlC \nrds the edge of the ,hell, producing rhc v:mous now faces outward~. <tnd becomes pt·nnanently
Lry w}'c" 111 turn from m outer surtace. r:mt the cell fixed Ill its final pl.1cc a~ the growing edgt• of the
rer ~teretes the gclatinom ~ ht'.tth, then the three layers shell moves away from it, all the while ,tecrcting
ra- ~f the periostracum 111 tu m. Thu'i. by the time the new material in conveyor belt fashion. When the
e has reached the outer edge of the ,hdl. it has secretton of primary cdl material j, complete, the
ICli a!rtJd}' produced four dttl"erem kimh of m.nerial m secretory function of the cdl changes for the sixth
'h e tocCl~~ton. h then begim eo gro'' c.1lcitt' rhombs and la~t time and 1t produte' a -.ingk long C'.tkitic
1 70 Brachiopods
penostracum J'l Oh'T.lll
I•) e-.temnl penostracal layer 'Jd\ lllCt
pr•marv shell layer
below tl
1nne1 bOU1101ny rntlrnbr<~ne Sf'Condary shell laye. JIIJr l.':lk
lac.') Bt
c~:~llular eprtheloum Supert"Jn
mcntdin.
protem c eme'lt
Sl'COIId.tr
t\otu.1o1rll1
Rcn·n
vacuol~>
cndu,kd
the ltvin 1
mucopolySi:tCCho;tnde spt< uk~
uhdc~.
secret•on droplet
(b) (d) Endop1

In rhvncl
endosprnes
rh~ ,J~t·ll
C <1\'l(tC\ \V
k11Hh of
lt.I\'C endo
~hdl ~true
rl'gnl.trly
puncta e.
taeolae
q oneratove 1one lUbular o
t .:aecal.' .tr
(c:) le I
tht: stand.J
\\luch bel
de A~ the
~ecum
.lrc
L,&(.'c Clt'
Cleft <~.tel·
through nu
ond.1 ry lay
1Jw C.IC
Jlld l\ COil
Figure 7. 10 Shell $lructvre in brochiopods: (o) standard secretory regime in Notosorio nigricans (for ellplonation see te>tf.t 1111 rnbc,
impundote shell (e.g rhynchonellide); (c) endopunctote shell (e.g. terebrotvlide) with coecoe possessing core cells freely w~. thl' hr1t~h
the cavity- the secondary shell fibres are shown os cut in section; (d) pseudopunclate shell (e.g. strophomenide) with no pnrnoryla,i
but with taleoloe prolonged internally os endospines; (e) shell of Lingula, with primary shell constructed of alternating loye11 ol ft paotnm; tl
photic and organic material - o marginal seta is shown. (Mainly based on illustrations by Williams and Rowell in Trealr>4 JtV hllow,
Invertebrate Poleontology, Port H ) 11111 \\ Jth
11, that 11
c;:t
fibre. one of m,lll)' tdt'lltK,ll dements m the scc- The modern terebratulide ~1/a/ttlllltl produr f' 1111.1nly SI
ond.lry l.lycr In dm l.t)'L' r the regular pattern of the shell according to the ~amc ·~randard \ec and tlwy 11
man)' long tnchncd tibrc~ n.tturally reflects the regime' as .\/MtJ.Iaria, but the prnmtracum 1111 pcnosu a ea]
ordered ,lrr.mgcnwnt of the indtv1du.U cells that pro- thtckcr and of J mon: compkx labynnt the n·pJtr <
duce them \mcc new nutc.:rJJI is contmually .added srrucntre. UIJ} ruptur
ro the Iowa t•nd of thl· film: by the ~urfuce of each Most rhynchonclltfonn braduopods, hvlll£ lflll' llll
cell, tr ~~ nor 'urpmmg to find that the secondary fossil. probably secreted their \hclh m much chaopuds d
layer " thlt'kt·,r .tr the umbones, whJCh are the old- same way as 'ofMcmrl. Otffercnce' m ~hell s~ u a;cstion 1
est p.1rr of tht• ,Jwll . of some groups, however. mdtcatc that the~ a 111~ wou!J
Morphology 171
progmum~ must h.l\'~ bc~·n rnodlfi~d. Io the Impunctatt' .md t•ndopulll't,lle ..bdh m rhyn-
~ivJn(ed' ~trophllln~md.t (rtg. 7.1 Ud) the shell choncllifomJ braduopod' -.et:m to cut ,Jtruss e'tab-
bdo\\ the penmtr..ll'Uill '' only a 'inglc layer oflam- li,hcd "Y'temauc bound.me,, .md tt ~~ probabk that
uur falntc trawr.t.·d b\ till hm·d cakmc rod, (taleo- endopunltaoon t•volvl.'d more tllJn once. T hu'
bej But m tltt.' ,\lll'l''tral 'twphomcmdc group. mo)t onludc, .ul.' unpum.tatt'. apart from one ~upcr
aptrf.uml) Ph!l t.unhllllltJt ~.1 (~ubordcr ~tropho f.mul}. the Emdet,Kt:,l. All Jtf) p1dc' and pen-
~mdmJ), tu wllll h l:ttpft·ctctdttll/d bt'longs. there '' a tameridc~ ,lrc 1111pum t.lte . 1\\o,t 'Pinleridc\ JnJ
ond.uv bycr of mclmcd l'.lkltlc tibr~ like that of rhynchonelhde' are al~o unpu11ctate, hut there i~
'IIHJn.J one mamly punctate 'pmtende \Upe1tamily. the
l{~t'rnt Tercbr.JruhdJ .llld I hcndctdina have an Sp•rifennace.1, aud one punctate rhynchonclhde
nJ,1,keletun ot l.lk.lrl·uu' ~r•rule~. o;ecr~ted wtthin supertam1ly of thrt•e gent>r,l, the Rh}nchopor,Kc;l.
e ll\ mg tiSSliC .1nd ,l\shtlllg 111 1t' support. ~uch All terebratuhJcs .1re pumt.lte.
!F''ult·, .1rc tmmd m \Oill~ Creuceou~ terebrat- In the shd l' oC mo~t !)trophomen,lt,l (F1g. 7.1 Od)
uhJcs the thm and often 1rrcgubrly formed t.tleolae gtve a
spunous unpre'\~ion o f punct.ltlon, espectally to
Endopunctation and pseudopunctation in shells weathered spccimcm. l-lcnce the tenn pseudo-
In rhym hondlifim11 hr;Khwpod~ such as Notosaria punctate is m cd ro distin ~lll 'h thl'm, perhaps
h~ ,hrll t~ impunctate. u:. ha~ no pcrtorations or unfortu n:uely, for in or her re,pcct\ their nucro-
mt•r~ w1rhin rhc ~he ll ~tn•cturc (f1p;. 7 .lOb). Other strncn•re 1' clc:~r l'IH)Ugh .
mds ot rhynch,)ndhtonlll''\ (e.g. tcrebrarulides)
,.e endopunetatc \hl·ll, (l 1g. 7 .We) 10 wluch rhe Hinge and articulation
Jl'·ll sm11 run· " pt·m·tr.m·d trom the i mtde by large, Strophic :md non-,trophlc \hdl~ .tre d 1\t111gmshed
gubrly Jrr,lngcd, dnn~.1tcd c 1\.1tH.'~ known a'\ by their hinge ~mH rure. Then· arc no ,tppar~:nt
puuctae nomul to rht· ,hl'll surt:te~: Thc~t· rontam 1ntt>m1 cd iarc~ bct\\'l'l'l1 rhem; mdecd 1t would be
hbr ouq.,rro\\ ths o( rht· m.mtk. the caecae. hard to unagme a bra< hmpod of 111tt'nl1edtate km d.
(mat' are fomwd .11 till' t•dge of tht· ~hdl during The variom \tnlttllrt·s a~~on.ncd wHh the hinge
dle ~und.mi \Cl n·tory rq.,'lmc by 'mJ.II knot'> of cells region arc de:~lr wirh 111 rum
-hi, h hch.l\'e independent!) of tht· rest of the man-
Ut ,A,, thl' ~'TO\\ ing t·d~t· of thl· ,hell move<; """ay the PEDICLE OPfNING
lleCW .m: pcnnancntl) lm l...nl in pos1tion but. since The ddth yn um ami nutothyrium ,1rc more dmely
:oc
uch cactum rct.1im ih umt,lt t with the mantlt· ~een on \trophic \hell, slmt· the hinge j, stratght.
throughout dll' dt•po\ltion of tht• pnmary and sec- Pri micivcly. as in ord11dcs, cnll'lc[ldt·~ and pen-
,ndJry layers, till' lower end nlJy become very long. tameridcs, they LOgcrhcr tC.m11 o.1 d1amond-<,haped
Tlw tat•cum llt:.u h rt',ll hl'' tht: outt·r ~hdl surface opening for the emergence of the pedicle.
nd is n>llllctted to it by ,1 'pcnostraraJ brush' of However, this IS not thl'ir ~olc function. for in br.l-
(b) on~ rubcs filled wtth mutopoly~.Kch.lride. Below chiopods with high intcrarcas they .1lso .11Jow a space
din th~ hn1sb ,\re ton.• n·lb fillni with glycogen :md for the diductm mmd c~ when tht:)t: are Lontracrin~.
l)'er
105-
protein~; these hang down frt:cly uno an empty c.w- Since the didutto r mu~d c~ arc atl<tchcd tu the cardi-
t on IIY bcluw, so th,ll the only u>nnclllon of the cae- nal process, their line of tJCllon ts brought dose to
.um with the m antle •~ by the tlJttcncd peripheral the hinge, and il there wert· no o pcnmg the muscles
dl> that line lilt' w.lll of the puncta~:. C.1ecae are would catch on the hmge line and fr,ty.
its "'1maril} storagt· lh,lmbel'\; they are .1bo respi rato 1y, Where thert· i) no pcdidc the delthyrium ;md
ory 11d they 111h1bit bonng or~.llll\llls. In addnion, the notothyrium may be protected by v.mous kmds of
.lCh enostratal brush on rdeJsc .111 orgamc 'glue' (for plates. In the pedtdc valve there 111.1)' be J smgle
like thr rcpa1r of llljury) 1f the pt•no\tracum ts acriden- plate (the dcltidium; ftg. 7 .llj) or. as m ,\la~cllanw,
clv ruptured. a pair of plate~ (dcltidial plates). umtcd by a
and Smce nucrobonng. 1n thl• 'hell~ of Recent bra- median suture ,111d arched <,o th.u they do not Inter-
the ilopodo; de.uly .wo1d punctJc (l.urry. 1983a). the fere wuh rhe hne of action of the mmdes.
:ure IUgge'\tlon th.H t.lt'lJicontenr~ mh1btt bonng organ- The notothynum n1..1y hkewm· be plated by a p.ur
:ory !SmS would appe.u ttl bt• ,ub,t.mttated. o f chilidial plates or by a smglc cbilidium. Rarel)
172 Brachiopods
d.t
~"' ·~
(a)
pn
~ 4r 11H
b~c
Ill\
'"6
(g) (h)
led
tP
cp \\I
i(~~·"~'" ~
Ill<I
c.;o:
·e (j) (k)
eh
Ill\
J'e
·~
""'"''"*-""'''m
dellhynum · dPit•diJI pllte~
~ Ill
l)IJ
~0
ll'(
Figure 7 11 (aI (dl Inclination of interareos: (al brachial valve hyperdme, pedicle valve anacline, (bl brachial valve anadine, ped>- be
de valve opsodtne, \c) brachial valve apsodine, pedide valve apsocline; (d) brachial valve opsadine, pedicle valve procline; (el-l 1 10i
cardtnal processes (el Strophomena, (f) Pustvla; (gl Hesperorlhis; Ihi Leptel/ina, fil pedicle Foramen with dosing structures; (k) trn
gular stegidial plates dosing delthyrium (Redrawn from Williams and Rowell in Treatise on Invertebrate Paleontology, Port H )
(c.
nl·
)m
111 'Pintc:ridc' then· 111.1}" bl· a stegidium (Ftg. evcncu.lll} JOin in tl11.: nudline, clmmg oil' the J'(d!- Ill'
7.11 1\). r lm 'Pl'n.ll tvpe ot delthynaJ coverin!! is a clc opentng t'llltrd) from dw ddthynum: tlm nu
'cnc' of l Ollll'ntnl tnangular tccrenons tom1ed then bt• dt~tm~n,ht·d .1~ .1 pl•dide ti1r.lmcn. br
dunng !--'TO\\ rh "htch 'ucn•,,•,·dy dose che ddth)T- tlu
i,ll t:.l\'tt)' IT1llll the nuNdc 111 . tNTERAREAS oc
Ccrt.un 'trnphomentdc' have onl) a very tiny I nrcr.Jrc.H pmpt'r .lrt' ti1und only in strophic shdh d<
hok 111 the umbo tor tlw cmt·rgence of the pedicle. Their artttlldl• rcl,tttvt• tn the hinge line ., VCC) l.ll
Likl•wtw 111 thl· 'ptntendt• Cyrtttl (Fig. 7 .8d,e), tmport.lllt tn taxonomy, and tht nomenclature o \\'I
which h.1s .1 tnurh exp.1ndc:d pedicle v.uve imcrarea, some ot the v.trmm .1tt1tudes i~ t!lustrated m F.,. b.t:
the pt•dtcle opentng rc:stdes 111 the ccnrre of the 7. 11a-d. Ot the~C' apsacline .lllJ anacline, resptl·
tnreran·.1. Any pedtck emergmg from such an open- tively, .uc thl· mmt common ~ome shell~ lu\'e thL shc
Jng cou lJ 11\ll h.1ve L'tubled rhe brach10pod to be interart'.lS of rhc pedick v.t!vc: .Htcnuated 111to J pro- thc
held upright 011 the ~L',I Aoor, hut could certain ly nonm-ed bctk-like rostrun1, whtch in l 'llole.( t< 'P·
haw .1ned ".1 tt·ther to hnld n 'o that the shell was curiously rwt~tl·d en nnt 'ick. lll
1\0l \Wepl .IW.I\ by CtiiTl'IH\. f\11
Non-,tmphH 'ht·ll~ \uch .lS the large Silurian CARDINAUA
l\nkidir1111 (Ftg. 7 H.l-t') h.J\'t' a large. long pedicle The stntcrun.'' .1r the pmtenm end of the bncht
v.llvt• wllh .1 \lrongl) L un·~·d umho but an open \";llvc. cnlkcnvcly knmvn 1' cardinalia ..1re l11ghl) At
ddd1\ nu m ~orue Ji,t.tlll't' .Interior ro the umbo. In differentiated .tnd 'l'rw varinm timctmns Of the~t 111.
JtiVelllk 'hdh of Mn,r,:t //,1/11<1 .md other terebrJmlidc\ rht' medi.tlly pl.tt nl r.trdu1.1l prnce's 1-; ofren th~ \).~
the pcdicll' opt·ning •~ lnc.ued in a nom1al mmt prominent (Ftg. 7 . llt'-h). In if'i simplest form.
ddthynum. A, tht• 'hell grows rhi' opening migr.uc' J\ found in rht• mure prinurive rhynchoncllifonnes.
po,tcnorl\ tm\.mh tht· umbo lw re,orption of the ir ., no more dun .1 p.ur of undttTerennated mu,ck r,u
'hdl Ml-.111\\ htll· ddttdi.ll pl.nc~ tom1 anreriorly .md base, behmcl the nntnthynum (fotg. 7 . I I h). A mort th·
Morphology 173
bor.uc kmd nf ~rmcmr~.· " J median ridge: tlm 7.Ha4.·). In thc,e dw pedicle valvt' h,t, ,, wlttul \\all
nmthhl)' s~.·p.tr.trcd the t\\ 0 Jre.J~ of mu,d~.· .lttach- Jivtded m two dhtally so char ir appe.m Y-sluped in
tllll, h111 in more .ld\'JilLl'd timll\ tr .ll'rtJJlly cro~s-st.>ctton. !'hts ~Lnteture. the spondylium. lit•,
till' rhe ,ire of thducror .1tt.1chmu1t through the duct d)- opposttt' a cruraliurn a p.111 ol 11\.',lr-vt·ni-
1\\ard mtgr.trion of rhc mu~cl~.· h.t,cs (I tg. 7 . I Ig). cal pl.lte~ wludt .1re out~rrowlh' u[ Llw br.lt h1.tl
A titrthcr dcwlopmcnt of rim kmd ot stt'lll tu re pron:'~L''>, h.lllgtng down ti-om the H•of ol tht•
ktl m .1 ,Jutr with ,1 hc.1d (myophorc· hg. 7.11 1). pedtck V<liVl' 'J ngethcr these stntllure' t•ntltl~t· .1
htth llllfl,t\1.'~ tl11.' ,Jrc,l of ntU\t·k ,lll,tdHlllllf .111d 11.1rro\\ verttt.llc.lvlty, open antenorlv abtlll( 1 thtrd
1 be \',111\lll\h ucnul.ttt·d. forkt·ti. tnlllrilnb.tre or of the w.t; t'rom thl.' umbo. in wluLh the lllll\t.k~
mb-hk .tnd prob.tbly .tll tht· otht·r org.ms of the body c.1viry
Othl·t rardin.tlt.J lllllmk socket walls .mJ bra- \\ere· cndo,ed ·" wdl.
chiophorcs, t'7\.p.mdcd knob, "l11ch lilt} h.tn been Mmck pl:trtonns such as these .1rt· not unu>m-
tm'Oht·d 111 'III'POrt of the lophophmt' mon 111 rhynchonelhti)nn bracluopods, ;md .11\o in
L"r:ltmfomlt'' 'mh as T1imcrd/a (hg . 7 . 14~) and
'ltEIH, SOCKETS AND ACCESSORY STRUCTURES \l.t'm eo h.IVL' ,tn\en several rime~ dunng t·vohtttott.
Twh .trt• alw.t\ s litund in till' pnhdt v.th t'. ,m-kl't'i RudwJCk (I 'J70) h.1s not1.·d that tt IS not llt'CI.''';,ty to
111 tltL' bradH.1l. Tlm wslt'lll .lpp.m·ntl) 1. vulvnl only havr a contrattile rnu\de mnning tht· ti.1ll ht•tght of
tile .111d tbt·n remained n:l.tti\TI} 't.tbk Thc n:cth tlw \lwll in orckr to open .md dos~.· tt; .1 shorter
uli't~lLJf knnlt, ot· Sl'nmdJry ulutc "1l h 'month or rnu,ck could dtt the JOb more economtcallv .llld
crenu!Jrt·J .,urf:tt' L's. In lllJII\' bralhtopod, the) nuy \\'tth brre.Hct control. Muscle plarfonm tend ro oaur
~uppurtcd b) .t patr of' ntic.d d~.·tH.tl pl.tre, \\ hich m rhmt' ''It'll' \\ lmh Jrc highly b•convn: rhcy
111 them tt• thl.' tloor of tht· pt><hl k v.tlvc. .1llo" ii>r the :tttJchml'nt of a 'hon but opnnully
lhe tc~.·rh may be ~upplcml.'ntnlut 1.' \ en tepl.llcd dl'"cctiw mll\dc '}\Cl.' m. Nor Jll 'tron~l; btl'onvcx
t g. E.•plroo.f,.,rt.l) Ill 111.111\ 'troplwnlt' llllk•, by den- shdh h.tvt' this 'Y'rem. the \UrvJvmg rhynrhnnd-
dl~ wluch ~ru\\ .1lon~ rhc nt.lr~lll' ut !111.' h.inge lide' and tt rd)r.ttulidt.'\ have p.trt of the mmck
nr. M~ m her' of Order Prmllll tid.t g~ na.tll} lu vc ,rr.md rcpl.tr~.·d by .1 non-contractik tendon, \\hi eh
~ttht·r tt'L'th uor dcnttde,. j, Jm>ther "ay n l nper;lting vvuh t'qu.ll dltut.·ncy.
So~kcts, ti.mnmg pMt of thl· L.trdtll,tli.t, Lll'tL'll have
.tr~· 1Jld thtl k m tenor w.1lk frt'l)lll.'lllh " l.1r~e as Lophophore (Fig. 7. 12)
rh~ t('dh thcmsdvt·,. In ~m h t.l\l'\ tht mtt"rior The bt.IL hJOpod luphophore, whme \lrm·lurc •n
iOckcr wall nuy even mdcnt • secondary socket ,\lt!l!dl"''w h." b~.·~.·n dt',cnbcd. 1s tunruon.1ll~ 'imtbr
lose t•' dw COL, rh on the pt•dttlt '.1ln \ot kt t \\ Jlls in .tll br.H:hiopod,.
~n 1:-e \\\'ollcn .1nd. 1' 111 I 'i.,fl)'• 1/,1, tloseh ,1\,0l i.ttc:d lu the br.ll.htupod\, though thl.' fil.ultl'tttou' 'truc.:-
l\1th tht• brachtophorc' m .uktng tlw lophophon: ture of the lophophure is comrant, there :~re 'e' era I
cs (I i~. 7.4d). distmlt WJY' in wluch thL lnphophore thdf ''
I reth and sockt·r~ .lte comrn11. wd nt scLnndary arrc~nged wtthm the mantle cavil~. ftu th~.nnme,
hdl. A~ the \hell !1:1'0\\'S, rhe tel'th move·'''·'~ from though ~Ollll' br,ll'h.tOpods have a t.tk,trt'Ull\ suppon
thl' hmge luw, r.tking up ~uccc~'IVt' though clo~dy for the lophophurc (~pirileridc~. atryp1tlc~ .111d rerc-
ipJC<~d postttOm .111d lcavm~ .1 rr.1ck ·. whtt h ts the bratttlidcs), others do nor, and in tht· onhides, a~ in
nunul.mn• dl'"cct of the posttJnn due tllc\1.' \ln.tc- mmt stmphomcnides, pentamendn, .md t"lJ"il hn-
tur~' h.1d throughout onto~en\ guhforml'' .111d crannformc~. 1t 1\ h.1rd to tdl what
the ongm.tl lophophorc ~trucmre ntll\l h,t\'l' bet·n.
Muscle attachments Linng rhvm.honcllidc'i likcwi<;c haH' 110 r.tktlil.'d
\Jductor. dtducror lnd ,tctJu'tl11 mu,tlc' Jrl' nor- lophophorc 'upporr, and m thc\C, J\ 111 li\ing
:mll) :Jcrachl'd dtrt·ctly ro tht' lmtdc, 11f the rwo hn!--'111JfOnHt'\ ,ltld CfaiUllOnllCS, rJw Jophophon.• IS
live,, fnrmmg Kar pancrm which .la' ~L·nm or held up purdy by the hydrostatiC prL'"ure of thud
1\X'( I~\ \Pl'l'ltit . in the gre.Jt bradnal canal (which l'.lfly tn lttc
Somt'tllllt"\, however. 111 th1. nHI\l ks nuy be becomes dt)sed off from the coelom). In man}
tmcd otr the floor by muscle platfomlS. ~uch .1s Srrophoml·n.H,l there arc tr,Jte'> t)f po,siblc
tho~t· t k.trlv di\pbyed in the pt·nt.tmt•r,ltt".lllS (Fig. lophophme structures impressed upon tht' imide o(
174 Brochiopods
(d )
hr.ll'h
l•l orrgrnal trocholoph!'
fc"" 1
Ill lh
d~vcl
(e)
great OIIQIOiil trocholophe

brach~<~l
can a
(lJlj
tou
(b) ...... Ill ~
nnt
termtnal termrnal spi
generatrve zone generatrve zo~e
WIC
rhy
(f) Amphigenflt (g) Gryphus
(c) ~pit
trocho nth
O 'mouth
In bo
l ~p1rolt
- uu--
lYQC>- <;(.hllO early spno· J0111CC
(fl) 6f end.
Tht
!
plecto
! ptycho sprro-
know
Ch10p1
how
~ UiJUD (0@) the.: ci
IIIITCI'
1
Figure 7.12 Brochropod lophophores: (o) sectional view of lophophore with Aurd·~lled great brachial conol, food groove ond ~ Ftg. :
ments - arrows show the drrection of movement of food particles; (b) arrangement of filaments in section, (c) the main lypes cl tiN of
lophophores showing various developmental pathways; (d) Terebrotv/o plectoiOphe, showmg mode of growth from o tenninol geroer eham
olive zone, (e) Rhynchonello spirolophe, some, (f) Amphigenio (Dev.), primitive loop slrvctvre of early terebrotulide (Centronell,c/ioat Ull OL
(g) G'Yf?hus (Rec.) short loop of Terebrotulidino. [(o)-(c) Redrawn from Rudwick, 1970, (d). (e) redrawn from Williams and Wri-Jt
1961 ; (f). (g) redrawn from Treofise on Invertebrate Paleontology, Port H.) 1 .~:. int
pmicl
th1.. br.whial v.tlvl', wh1l~ the: .ur.mgement of the ver- C hange!> 111 the form of the br.lchJdmm dunn rent~
ontoge ny provtdL· ·' ba\is tor the das~lficaoon ~:xhal,J

tiCal pl.HL'~ tn litlp/crttldtl/1111 .md rdatcd genera sug-
[o!;l..'~t~ the f(mu th,lt the lophophon: mu~t have had: lophophores tnto d1fft·n·m type~ (f-ig. 7.12c) All mocin~
lopho ph o rc~ pJ~~ through tht: same l'arly ~t.1gr~ m the \h
other posMbk lophophorc ~trunures give little
information on the ongu1.1l form. developme nt. The firo;t ~tagc, whether support~d or •"OilleS
The lophophore support j, represented in not, i~ :1 trocholophc. li ne the lophophore 1 the ant
m erely J p.1ir of curvmg ' hnrll\' proJecting horiZl'll ~ep.trat
rh\ ru:hnndlid~..·, .trld ,~,.)me pcntaml..'ridcs o nly by ~~
p.nr of ,hml \lrttls (crura) proJecring from the car- taUy on ellher ~idt' of th e 1nouth .md fomunji; 11: NCltc
dm.tha. ln group~ where the br:1chidwm b a cakare- lllCOmpll'tc nng. Only one genus of m1cromorprJ wnica
ou' ribbon (t~rcbrawhdl..'o;, .11l)'J11dc' and spirifendc5), adult brad110pod~ ha' tlm kind of ~tntcture a~ tlw in 1
1t .Llway' ~row' (or grew) by acnctlOn on the (ame- since the trocholophL fonm the inm.U developmen- "lm h
nor) gru\\ m~ edge .md by rc,orptlOn on the (po~te tal stage of all brat hwpod,, llu:. parucular C<L\t' ch.unb
nor) tr.uhn~ edge ,,, tht· \hdl l..'t1large~. ln'v<triably probabl'r .1 ncott•nom development. fhe next ~~~r. 7 ~a,b)
111.~" tiLnnl'nt' .m:- .tdded ,H the growing end\ of the m dcvdopmem 1\ the schizolophe . where the r., mgly ~~
lophophorL' alone. nt.'vcr mtercal.ttcd withm the origtnal hom\ h:\vl' bL·tom~ larger but are bent bJd pomt u
fil.unent \l'rll''· and nm par.l!lcl, dtrcct~J tO\\Jnls the momh. ~
Morphology 175
Ulhiopod lophnphorl'\ go rhrough this stJgc too; a <.:alntied bradudtum. lr \cem' rea~orublc to a!.stiiTil'
fell nurromorplm ten·br,ttuhdc' h.wl' srhtzolophcs that currenr sy~tems fi.uKuoned 111 a ~unilar mannc:r
n the adult. hnm herl· then· are three possible and that the unilltt:rcd water lay 111~1de the 'pin:~
lb'l'lopmcnul p.ulm .t\'s: wh.tle the exhalant ch.tmbcr w,t, nut!.tde them. In
Spirij£·r. however. thl· 'Pile\ poutt later,tUy (Fig.
one: 'Llrung with J zygolo phe, \\'hirh lcJds to a 7.7c.d). and wacl'r must have hllcrcd from the out-
plectolophe: the long or 'hort loop oi mJny ter- ~tde to the ms1de of thl· sptrc~.
~bratuhJc,: Ple crolophc~. as dewloped m modem tcrl'brat-
ont• leading to the multtlohed ptycholophe, :ts ulides. arc htghly ctlit tent \tniCtures of complex
found 111 the nwdcrn Lut~:clla, ,t theodeJn, and three-dtmcnston.ll tomt [ .Kh .Inn dl·vclops as a
111 \Omt fi),,s) prodUl ttdn. cur-Ving horn (the trocholophc ~ta~c). then recurve!>
on~ kadmg to rh;.: twin 'Pir,u 'Y~tem of the sharp!) ,1, .1 h.1r runrung p.tralkl w1th the original
spirolopbe. pt•rhap~ tht nmunom:\t Jnd most arm and lymg .tbove tt (the zygolophc ~rage). In rhe
111dc,prcad tumtional 'Y'lt'111, po~'l'"cd by Ren·nt titlJy developed pknoloph~o.· thts b,1r lllms mward\ to
rh}tKhonclbde' .md ltnguhde,, .ts \H'U "s by fossil tcm1inarc in J vertical sptr,tl cn1l. The two sptral
1pmfendcs ,tthyndc' .md .nrypide, and po<>sibly median coi l~ developed trom rhc p.med anm lie stde
llther foss.U groups. by side ami with their til.tmcnto; occupy much of the
spacl' in the centre of the ~hdl; between them is
In both the pkuolopl!~..· \)r tcrebratulide' and rhe inh<Jl:tnt water.
ptrolophe of m.lllj h'"H)ups tht: twm ribbons may be Members of Supcrf..1mily 'Tercbratcllacl'a
JOined by 1 stout rod. the .JUgtun. rH:.u the posterior {Subordn Tcrcbr.ltl·llidm.l) h.1vc plcrtolophcs sup-
nd. ported aJong mo~c of tht•tr length hy ,1 long calcarc-
The mode oi opl ratwn ot sp1rolophe~ i~ wdl ou~ loop in \\hi<.: h the two cmls h,JVt.' JOlllcd up,
mo\\11 Ill S0ll1l" llH1dCrn rhyn~o.honellide bra- though the median wtl J' 'upportcd by hydrmtanc
btopods, and \Ollll mfi:n·mc~ can be nude about prc~~urc alone. ll owt ver. m tcrl'br.Hulacc.ms onl)'
!w11 It funcnoned 111 fo"1l' A\ 111 all brarh.iopods, the ba\al pam are 'uppmt<.·d b; ,1 muth 'hon~o.·r lnop;
!he ciliated fi!Jmcnt' borne on the brachia cn:.1tc chc rest rem aim erct t through hydro\tJtte prl'''urc.
rurren~ ~h~rh nonn,11l~ Cntl'r the ~hell laterally though some addmorul 'uppon t' ~rtvcn b~ the l"aJ-
Fig 7.-). When th~ \\ .uer pa~sc~ into the ~hdl. it careous sptcule' wirhm the lophophor~o.' tt,l•lf
nfalllrc' ouhrdc thl· lophophore m an inhalant Ptycholophc, arc gent:r.tll)' rather flat muctures
); hamber lt then ha\ w P·''~ through the nlia from and do nor form the complc~ three-dimen,ional
t, outside to thl· 'P·""l' l'ndo,ed wJthm the spires, cotls found m the other t\.\.O lophophore typl.'~.
r mto an exhalant chamber. As Jt doe~ so. food Pn:sumably they arc le" dliuent filtenng ')''tcm,.
p;~rncks .1re ~rr.um·d of[ Urachtopod feeding cur- They arc uncommon toJay, ot.:curring only in
renh arc very prl'CJ~cl) controlled lnhal:mt and Family Meg,tth yrididae (!->upcrf:umly TLrcbratcl-
xhJ.bnt curn:nts n.: m.un tntact and there 1s neither lact::a) and thccidc.tns, but they may h,tVl' bl.'<.:n thl.'
unng nor eddying oncl' rht• current~ have entered uonual lophophore type 111 stmphl>menids ..1~ i~ wit-
11 rhc shell. The w.ttt•r from thl· cxlulant chamber nessed by the brachial ridge systclll ot" Evplectodonta.
)1" .umc' out mcdwlly, and the marked deflections of Most forrunarely, int:lct calci fied ptyrholophc~ hav<.:
h he .mtcrior margin of rill' rhynchondhde~ appear to been tound (Gram, I'172) 111 the ~mall Pl'nnian pro-
1- pmtc inhabm from c'<h.ll,\nt ClllTents. cluctacean Faltt{er (Ftg-. 7.17.1 ·t).
111 \',,r,wria t' J R~o.•ct'IH rhyrH honclhde whose t\vin
jc onical 'Pire\ pomt downward~. 'I he~e spire~ form
d. e mhal.mt chamha .1nd cont,un unfiltcrl'd water Subphylurn linguliforrnea
I- hllh it p.h\e\ through thl• tibmt·nts t0 an cxh.llant
I~ .lumber out\!Je the 'Pin.'' In Ordl·r Atryptda (Ftg. Th1' -;ubphylum of orgJnopho,phattc-shelled bra-
·-J,b) tht tnorpholot-') of the lophophore 1s ~tnk- chiopocb 1' con,Jden·d to be more closd; retned to
.'0 ~~ ''ffillar. with tlw 'ok• dttfen·me that the spir~~ bryozoam and phoromd~ than to other braduopods.
ck nt upward' rather tl1.1n downw.1rds. and there is a The be\t-known repre\Cnt:tttve 1' /.,u~t!ula
\.U
176 Brachiopods
Lingula tem i' unhke th.tt of a reil ul.lle,. The two v:U1· \(1 ~
u11~u/a {Ordt'l Lmgulid.1) 1\ J plllhpil.lttc- ,hdkd clo~e b) adductor mmcJe,: a '111gle po,tl nor mmdt fJul t
lll.lrnculat~ br.lt l11opod \\ h1 t h ha' per.~\tt·d ~iiKl tltt and .1 p;m of antt nor one!.. When the,e relax. r~ :lithe
C.unbrian wtth rdarivc:l) httlc chJn~c:: .1 type exam- valves mow ap.1rt 'lightly rh rough rhe el.l~tlClty ol tht•y
plc: uC bt-,tdvtdlc evolutu)n. Modern Lill_l~rtla hve~ the muscle~. but otherwi~t· the valve~ can onh m (E
suuc:.,,fl.tlh Ill a hr.tck1~h ro mt~rud.1l t'llVironmen t. rot:Jtt'. 'hear or ,Jtdc ag:limt one another through th
norm.tll) 111 tim· ~Jnd (t rug. 1951) Tlw earl} 111\,1- acuon of ' 'anou' o blique muscles \\ h1C h .uc lartt Ot
slun nf Lll/~11/,1 Into >Ill h .1 'dttlicult' envtronmt·n t and wd1 dcvt:lopr.:d. The ped1cle h,1s a leath,r) I lwr
lllt wluch dwrt' were t\.·w cmnpctltors i~ prob.tbly extern,\1 cuticle w1thin wluch is an cpithl.'bum I 111h'1
Oil\.' of the pnnury n.'.l\011' fnr m \'lrtliJI evolutmn.- emhi)'Ological1) part of the mantle. Imide it th tour
an 'tJw;. Nt•vc:rthcJe,~. thuc j, rCJ\Oil to bellt'Vt' are mu,cJe, en.tblmg the ped1ck to contr.Jlt. ~o d wh•cl
th.n some t"o~\11 l..JI~i!tt/,, 'P'xie~ ltH·d m envtron- when the end ~~ timtly t:lxed 111 the ~.md the b' • <l\\1
llll'nt~ orher th.ll1 the intt·md.tl, indudmg shell :1nd chit)poJ c.m be withdrawn ulSide Jts burrow. A1, Urllbc
b,,,ll\,11 region (Chern,, 11>74), m tht• u\e of Lllll!llln arricul.ttc'. the shell i~ secreted by cdh produced 111 n 1nu
,J\ m em~runmental Jmilc .Jtor I' linuted. Some dtt: gcncr.1rive zone 111 the mantk. under the '.uve n. 'lt'<lllt
kr• nce-. 111 I .IIIJ!Iti•T-bc:.trtllg · pJl.H·oconm1umnc:\· In Lil~l!uf,, (Fig. 7. J Oe) the penostr,JCum "under ud,, r
through time luve b~en documented by We,t by J primary shel l wnh m.tny ,Jitcrnating ~tranhc lllllrtl
( 11176). layer' t1f organic ,md phosphatic material. The) Jl I'Jiwc

lodem li111!11la (Fig:. 7. 1-tl-c) h." .1 p.1ir of :1lmmt all 'ubp.u-allel wnh the shell margm, and rJthn tJ wherh
tdenuc.ll, h1bter.1llr \\ mmetnc,tl v.1h c:~ Thq Jre being rontinuou' layers rh~:} arc elongated J(lllO I nul)
gcntly conVt''l:, .111d from hcrwet·n them projen' ,111 whose mode of fomt.uion 1\ tncomplcteh unJ ltiiVn
clongJtcd pt·diclc. The Juunal hv..:~ in a VL'rlJC.II stood. The phosphatic layer~ .trl' thicker '"here th '' luch
p<Ntlon in .1 bm ruw .•md it~ pedJCk extend' deep overli~: the body CIVlty. Thcrl 1~ no 'econdlry lJ d1v1Jt·c
imo the ,.md. w1th tlw upper edgt'' of the t\\O nor Jre there punrt.le. Jnh: r .w
VJIVt''JU~t .H rhe kvd ot the ~ur£tce In htt- the di'tal It ~~ imerestmg to notl. how L111gu/,, bui'TOI!! tllb'ltlu
part of rhe pedicle ha~ litth: roots which :1nchnr the (Th,1yer aud Srcde-PetroV lc. 1975). Tht> bngt 111mu,f
.mtmal tlrml} ..md t.f Ltl(!rd,l IS dtsturht·d the pedH:k Cf~~ttidi,,, which " the bc~t 'tudil·d. entl'T\ the scdr
i1N.mtly cuntrJl h. wnhd1Jwmg tht ~hell dt·ep mcnt .mterior end first w1th tht· pedicle uuu.illv
Wlthm thc btu row. U) .malogy With other bra- as J ~utr prop ;md l.uer rr:nhng behind. Tht' bra~.
chwpod~. the ped1de end ts descnbcd a~ po~tl·rim; pod then burrows downw.1rJ\ through the >cdum
tht ,Jtghtl) l.ug:L-r valve 1s c.1llcd rht" pedicle valve mmg ~cissor-hke .llld rot,lT)' 'awing movemt'ntl
.md the \lll.lllcr the br.1chul v:JI\'l'. Honn 'CtJc the \al\'t'S. In addmon. penod1c rapid valve dos:<:
round the expmt·d edge' ot the ~hell nurk otr chn:c ejel t~ water from the mantk c,lVlt)' and the
.1pcnure~ k.1dmg to the HNde. the outer twn tor setae pa'i.~ mucus-boun d \.tnd upward~. Whm
llltllrrent \\,Jlt·r. the cenrnl for excurn·nt water. brach10pod has thm burro\\l'd, anterior end doll
The shtU " lmed by .1 mantle .. md the antt:nor \\Jrd~. tor some d1stam·c, it then rum~ upw
body \\all, rhe lophophort Jnd othl'r mtcrnal organs descnbmg a U->ll.lpt·d trad... W~thin a le\\ houn
.lrt .1, in the 1 hynchondlltom1e~. Tht· lophophure :s amm:tl comes to the surf,Kc Ill ~tand.lrd tee
.1 ~piroloplll' with the two ~pirc~ directed inwards, pos1tion, the correct way up This remarkJble l" ( IJ)
t'.tL h bein~ .tn 'tnlubnt 'P're' c.JrT) ing untilrt•r.:d rowu1g sy,tem dOt's not .Jt any wne 111\0ivt rh<
\\ lttr wh1ch " \tr.tmed Jnd exhaled mcdiall) 1 he of the pedtdl' ·'' .1 d1gging 'foot' (as m b1vah•cs.
gut, however, h." an Hlll\ d1sch.1rgtng po~tcnorly. ,1 method unique in the an1111.1l kmgdom.
;lnd there arc lour nephndhl. The circuiJtory ~y~tcm Un,{/lfla is widdy rcputcJ to be .1 cury
'' '1mple, and there IS .1 tll'T\ ou'\ 'Y'tl'IH with 1 gan- genm. but the curvhalinity 1~ in t~ct only mo<l Chile
1
~1011 ne,1r th.: mourh ..h tn the ar!ll ul.Jtes. Ont' of 1 hm the Ausrr.than L. cm,l/ill<l can toler,ltt'
tlH.' greare~t u1lterenccs between Llll.l!rt/,1 and arutu- level~ from 20 w SO per nu lie. but on I) for Jbct': F!EJUTO 7. I.
l.lWd brachwpods lie' 111 the mmculatme for opt>n- weeks. :md such fi:,1tures '" pcd1ele regencr.m~ pedorotio11
,1void:mcc of prl'JJturs by rapid withdraw J I volve (>s
tng .md clm1n~ the \hell (F1g 7 1-k). Since there IS a~mple te s~
neither a hmgt· line nor teeth and 'Ul ket~. the ,Jwll imp;urcd (HammonJ. 19H3) m a much ,hort' of ped.de,
<".Ill IItH open bv lev~r.lgt• ..111d ~t) tht• mu~cul .t r ~y~- Othl'T linguhlh \UT\IIVC Jt ~.1lumies ranging lr from lreoti!
Morphology 177
t 42 p~r nnlk, .md arc \H'll atbptt·d lo \\3h.'T\ of rl'f-;;n:nCt' to tht• mu,de an.Khmcnts .llld. though
uo.n1.1un~ salnuty. u'uall~ in '' .rrc'" nl X Ifi°C, and thcrc arc rk.rrly ddined Jdducror .tnd obhque mus-
althmtgh thn 'hm' .1 prcfcrt'lltl' li1r ,h~llnw '" rti.'T\ dc,, ~Olllt' t'Vtt'knce 'ugge~rs that the lc~,·eragl' sy,rcm
tbl."i hJH' bt'Cil Cl' corded Jl tiepdl\ llf dm\ ll In SilO could h.IVL' fimt noned more hke th.ll lll .1 rhyn-
I EIIIIH t I rr/., 197X). chondhform rh.w rhJr of Lill.l!llla.
Order Acrotret1da (L Cam.-R.ec ) o~tcn Ius emu-
Other linguliformeo (Fig. 7.14) Jar v:tlvc~ of dtffcnng convextty, ~omt' pn\~L·s~mg
rhtrc art• rwo classes \\tthm llm ~ubphylum, high. conic.rl pcdick v.1lve' .md AJt, dt\C-\h,Jped
lmgulat.l .md P.1tenn,1ta. CIJ,, Li u~ul.rt.l t un'i'ts of bracht.tl valve\ (cg Gmvlr<'ta, Nwumw111r1) and have
fimr ord~r'i (Hnlmcr. I 1JHlJ). Order lmgullda, iu mcdi.m wpt.t 111 the pcdKk v,tlvL, oftL'Il tl! cnmplex
h1.. h thl' ~hell~ .m: num.h dungllt'd o\,tl, \\ ith lonn. ·orJa DhumJ.t h.l\ rather Att. round or ov,1l
\\th lmt•s, roncl'ntnc round .1 ucar-marguul 'hdh. with 1 'ubm.trguul umbo and us11.1ll\ \rn)ng
IDtlbo. \l'l'lll\ ro repn:sent .1 n.mnal gruupmg. ~ubcotKL'ntnc g£0\Vth hne': the living Dl~riiii'Crl h.l\ .1
l hon~h I.w~u/,1 .111d ItS rd.Hive' \t't'lll Ill bt• typ1cal :.ubu:mral pL'lhde ti..>r,lmt·n from "hlt'h cmngc~ .1
\mg 1(1,,,,,·.
h,,,·mg pt•rststcd for long uml' pcri- short stout pL·du. k, the foramcn of tht (. >rdovttt.lll
.1 h. tht•rt .\n'. however. deJr diiTcrt'llt t'~ in 'hell Orlnmloulc.> " 111 J ~imilar p<hitton. Order
d I!Urrmtrurtun· het\H'en li\ tng .llld lower ~iphonotrctrda (~.g. N11shbidla; Fig. 7.15) dtlkr.. from
c P.tLt·,,zok l ill.\!11/,, -.pecte~. v.. luch c.t'l' doubt upon the acrolrcmh utJinly in 'hdl stmcture. onto~eny .u1d
'1 1 h~lhcr tlu.• lJtta do tn tjcr pert.ltll ttl dm genus.
Fauuly Oholld.Jt' bdont--..; tn rim order, 11> rt·prescn-
the Lkvdopment of thl' pedicle tor:mten.
Cl.t~' P Jtt:nnJt.l (L. Cam.-l.lte Ord .) . unhkt•
!1\e\ hJvt• ~ubov.1l nr rounded ,hc(l, (I rg. 7 .I Je) in LingulatJ. ha.., \ttophic shello; and \Oillc otht'r 1~:.1turl''
hkh tht• prdtdc '.1h c nftt•n h,,, .Ill tilt~ rn.tl fl.mge suggestive of mctpicnt rh> nchondhti..m11 'tntcture~.
di tck>tl mto C\\ o p ropareas \\ hrt h rl'\Cmblc the including .t rudunentary nototh~ num, ddthynum
t rJrl'.l\ of 1111llllact·tl brac hmpod, .md lun· .1 tri- and pain:d Jdductor muscle~..\lioouutrrl '' typtc.tl: m
ll b'Ultr pcdtclc g r oove in bet\H't'Jl tht'm. l he the gcnus VICI}'llllllcs the brachtal valve h,ts a rt·nurk-
usculmm nf Ob,,/,s un he rL·n~mtntlted '' tch able sur£1ee \CU!pture of open pores.
o-
~
mt
of
Jrc
•r.tl r
the (dl (f)
0
~
r(b.
the
~
(g)
~
(h) ~
~ (m)
le)
iuc
rate Clule Bicia Kutorgina B1llingsella Glyptona
lOll)
,ut 4 ~713 Cambrian brochiopock (a) Chile (Chileida), complete shell; (b) same, interior view of pedicle valve, showing central
,md nro'1011; (c) some, lateral view of pedicle valve ( ~ 15): (d) Bicia (Obolell1do), in tenor of brachial valve; (el some, Inferior of pedi-
arc volve (>< 5): lfl Kulargino (Kutorginidoe), interior of pedicle valve; (g) some, interior of brachial valve, (h) some, postenor view of
~shell showing Pedicle opening (>< 8). (j) Bil/ingsella (Billingselloideo), exterior view of brachial valve; (k) some, interior view
;111\l',
ped1de valve ( · 3); (I) Glyplario IProlorthido), complete shell; (m( some, interior of pedicle valve (x 3). ((d). (e). (j). (k) Redrawn
Tl 13 Tre<J~se on Invertebrate Poleontalogy, Port H, others from Holmer, 1996, Geologiskt Forum 9, 1Q-14 ]
178 Brachiopods
Tnm~.:r
antenor lnge b1
posterior
(c)
0
<--~oblique
adductors
~ (dl.
muscles Micromitra
, .-'
homeodel
tidium 0 propareas
bl) .1ra
pseudc
\\ lth ,I
f~.: Hurt•
mppnrt
down~
dcvelo1
indcpc1
pedicle
(e) (f) (g) Trimerella (I) (m

(al (b)
Lmgula Numericoma Orbiculo1de1
Tlw se;
ut the
Figure 1.14 Lmguliformes [(a)-(f), (I), (m)] and Craniiformes [(gHk)]. (a), (b) Lin9u/a (Rec.) in its burrow, frontal and lateral Ylev.\ uptlllll
showing inhalant ond exhalant currents; (c) some, showing internal muscles in the brachial valve (x 1); (d) Micromilra (Poterinidol~
l
In ncphn~
Cam), complete shell (x 5). (e) Numericoma (Acrotretida; Ord.), interior of brachial valve; same, exterior of pedicle valve(!( Al. I
Trimerella (Ord.), interior of pedicle valve (x 4); (h) Crania (Rec.), exterior and (j) interior of brachial valve; (k) same, inletio. I ltt· \CJ
cemented pedicle valve (x 2); (I) Orbiculoidea (Acrotretido; Sil.) pedicle valve; (m) same, complete shell in lateral view (x 2).(b only w
mainly on photographs by Rowell in Treatise on Invertebrate Paleantology, Part H, except for (e), (0 from Holmer, 1989, fosl:ls Appare
Strato26, 1-176.]
tng rim
\C.Ir.
A \p
\\ dl tl(
rum v,i
Subphylurn Craniiforrnea
(< urry,
In tlm ~ubpll\ lum. wnh 1ts 'lnglc CLl\s Craniata, the utachc(
,hdb dre of takium c.lrnonatc, aruculation 1s absenr m late •
and thl· pcd1lk IS .lbst•nt altogether. The alimentary pcrarurc
can.ll has .1 true .1nus. ' J h1s d .lS\ i' regarded as closer hi') p
m Rh}ndlOndhfonne.l tlun to Lmgulata. Three dnrcas1
~pJn. 1
orders h,lVt' lwcn tktincd. Members of Order
Craniida (t• .g. Crcllllct) .m: unmu.tl crarmfonnes in growth
h.wmg a ~hdl of protcm ,lnd t.llnum carbonate. They Stzc di
~ur\tVC tod.l) ,1, the most \tu: n.:s~fu l of calcareous- 'how n
shclkd cr.lmiti.lrntes. In tht• br.lchial valve the shell is b1::mnua
Lnplc l.tyen:d Extt•rnally there 1S an impersistent lllmed'
mucopolysan h.lridt• ~ht'.llh with the periosrracum flll·orcti
below. Udow lht' is the prim.1ry shell made up of In a I
Figure 7. 15 Nushbiella, (lingulato, Siphonotretido), bro&
nct•tUe-hkc calutt• cry~tallitt·~. ,md fmally there is a valve (x 10). (Photograph by courtesy of Dr Lars Holmer.) d1L tcrt
sewnd,lry byt'l of t·:tkiuc laminae ,Jteathed by pro- brood
\CClllS t
rein ~hcct~. 1 he ~ewnd:u·y layer 1s found only 10 the
bradual v.1lw 1t ts abst•nt in the pedicle valve. • hwpoc
wh1th ccmcnb mclf to the 'urtacc by its outer de and the pcd1t le vah c cc memo; it,clf ro an appr I he
mut.opoly,acch.mde ti.lm. Craniaccam are punctate pnate sub,rrarum. I\ ljUI(C
with branchmg c.ICC.It' m both \alvcs, used for stor- The ~mall Order Cramop,ida (e.g. CrJu rhynchc
ing pol}'archandc' .md protcm. Where the muscles includt·, fonm wnh an impunctate ~hell ha\'ingcn embryo
arc mscrted, tht• c.llcitc of the pnmary layer fomlS tallme pnm.uy and l:uninar secondary layers rutural (
platfonm 111 the central part of each valve. ),.,... a ha
blades nonnal ro the mrf.tcc Cr.miid' have no pech-
Classification 179
Tnmerdl.!t l',l (Ord -Su.: Ft~;. 7. 14g) consi~t' of hotly, the othl·r the mantle. From the numlc ~eg
Oif!!' bracluopods wnh thick bttonvex shdb, proba- ment there dcvt:lops :1 mound-hkc nd~c. which
' ua~on1t1c, havmg m the pcdtdl' valve a large stam to grow lophophore fibmcnts wh1k the lar\'a
r)rudointcrarea and strongly ddua.•d proparcas is \till free-\Wimming. At the same tune the l.trval
·1th J ho m eodeltidium in between. A pcculiar shell (protcgulum) develops ,l\ .1 ~mgle plate
I tun· IS the rnmmon presence of lllli\L le pladonns (though tlm IS unmual in any br.lchmpod), Lm:r
'upported by .1 central buttress and often tumed splitting into two separate pbtes for the hr<lt h1.1l Jnd
lo\\11 .lt the edge~ to endosc p.11rcd ch.1mber~. Tllis pedicle valv~.:~. respectively. The pedick too grows
1dopml·m of muscle plattonm ts, of LOursc, quite from the mantle ~cgment, and wht•n 1t IS brge
Jcpcmknr of chat in rhynchondhfonncs. enough the brachiopod settles upon it, mually b>
the r'irne some 10 or so mantle fibmcnts h.we been
product·d All other \tnlctures (e.g. mmdes, gur and
nephridia) onguutc· from the body segment.
T he rhym;honelltde Xowsnria (cf. Ftg. 7.1 g)
TbC' 'L'XC\ arc~cpar:nc m brach10pod~ l·cmhzatton shows the very ditTerent character of rhynchonclli-
t th~ l'gg'- r.1kc' pl.u:e m the sc.1 water after the fonn dL·vdopment. The larva h:t~ thrCL' segments;
IS,
L. "rwm gcn11.:11 products have been shL·d vu the the upper is the globular body segment or anterior
phritlla through the m:mtle cav1ty .md out imo lobe, the middle th e presumptive m:mrlc, and the
~~ ~ ICJ Breedmg habits arc poorly known, being lower the pedicle segment. Initially the m:~ntlc seg-
Sd nh well documented m LJn.~ula ,\lld li·rclmrwlina. ment hangs down freely over the pcd1ck segmcnt a_s
nd
~pJrcntly \Ome ~pecies have 1 single annual breed- a cylindrical 'skm'. This then inverts Itself w cover
+time: other. produce muluple brood' dunng the the body segm ent, leaving tht· pcdJclc expmed.
Shell secretion then begms on thc now reversed
~ 'PC:l'IL'' in whtch spawmng p.ntems are now outer surface of the mantle, but the brachial and
rll ducumentcd 1\ Trrrbmtllllllc! rt'1Wcl (ftg. 7.3) pedJcle vJlve protegula .1re separate from the bcgm-
1111 '' Jtt'J'\ 100-JOO m deep otr we~tern ~codand ning. Nter a very brief larval life (~hortcr rh.m m
Curry, IIJH2). The~e brachwpods nonmlly live most in.~rtlcul,lted brach iopods) the yo ung mdivld-
ltllChL·d to biv.llve~. The1r bLmnual sp,l\vning cycles uaJ settles on the pedicle and compl ete~ its dcvclop-
1IJtt: ~pring and late autumn arc 111itbtcd at tem- nK·nt.
'IJturc' ol 10-11 °C, .md the n:prodtKtiVC cycle is
tl prcu'cl} tllncd. Growth I\ 11 tir<it rapid,
.nrea,ing throughout tht• (maxumnn) 7-ycar life-
m. There " .1 ,Jm\ mg down or ce,~anon of
wth m '' intcr, producmg dear J:,'TO\\th b.1ndmg. Bracluopods h,\VL never proved very cJ\}' to da'~1t)·.
:ze dMribution pattcm' rccortkd .h hmogr:uns and to some cxtcnr the dtflicultlc~ ha\ c bccn com-
1\\ rq.,''l.ll.!rly spaced p~:ak\ t nrrl·~pnnding 10 the poUilded hy homeomorphy. Some of the earl)
~nnual SL'ltkml·nt ·cohons·: the,t·. mndcmally, schemes \NCrc ret.uncd for fjr too Ion~. s1mply for
umcd out to be idcntical with complltcr-lu~ed and want of :mythi ng better.
lil'Ort'tlt.llly predicted sunulatJom. T he basis of a new damficatton of brnc h iopod~
In J tl:w RccL·nt brach!OpOlh the tkvclopment of was laid by Muir-Wood (1955), who made 1t cb1r
1ial
lertil1zeJ q~g.' takes p!Jcc 111 spt·c 1.11 inv:~gmared th,\t the ~:onccpt of working down from 'orde~·
rood pouc hes 111 the shell of the ti:malc, .md this predctemuncd by a few key characters h.td to be
m' to h.l\o c been thc case 111 cnuin t(,,sil bra- abandoned m favour of a scheme of bmlding up
~pod, .l\ well. from th e gcm·nc lcvd to higher umt~. thu~ rccog-
ro- The development of the zyg1'tc and l.!rval stages nizmg but avo1dmg the problems ari~ing from
quite d1tfcn.' nt in linguhfonn, t ranntorn1 and homeomorphy. rim defined the m.un ordt:r..,
tps) mchonclhlonn brachiopods. pmv1dmg good which arc clearly ~et out m the TrcMtsc tilt Jm,atl'lmllc
i'l'- JbryologJl.tl evidence that the thrt·c cl.l~sc·\ arc Paleomo!t~I/Y of l lJ65. However, there re111amcd a
md rural UJVI\JOm. In LJII,I!IIIa the f'rcc-s-wimming cJrly fundonnent.1l problem, which wa~ how the defined
d cr il\'~ h,1s two ciliated ~egmenb: one become~ the groups were actu.1lly related to eJch other, and a
180 Brochiopods
< n :aenozoic I
m.. J
I ~ '- :")
~-
"l) Cll llt
.~ 0
Lingulida I dat.!
all b
I Discinida
le tdcr
be m
I s.,....uuu 'ouua
lCf . . . .
T
,! LINGU LIFORMI A
Ca111
IPaterinida ha~ r
...
--+.~
ou'l)
t- - -- ·- - - - - - -- , - - r---- -- he!le
~
,(~ Cr
niida C\olt
f.llln:
Q
1\ 1erellid< CF A~"'"" , .."" - --
Ant.t
- - Crulclda
- c--
T'
- - - -- -- -- 1- --- bcc:n
snb,l.
Die yonellid na chJOr
- Ob~lellida ··~ llm
gr.1ph
,1dopt
-KT"'"'"' bdo~
~
>rthot 'tidina dun'1
~
(J1
dw ,j
lilX ll
II~OR I ~EA
110\\'
~\.ll'l1t
~~
!!I Cl it 1mbonitidin~
I PHYlUA
"Or I I
~~
StroP,homeni(!a
honetic
P•od~ '::~: (D
b1valn·
liOOlllJ
lcarcc
fonnC'd
- ~ -~
...... W.·
Protc rthida
Orthid
litter \"
SUBPH\'
hmt;cd
1111 lily
I' ~ v.ohc, t
\V~
CLA~
Pe rtamerida
)_~
urga1
HtJC
Rhynct onellida timn

Atry1 ida Cut
~
...
(J1 ~ ~~ ~~
_i~j
OR
~
~ Spirifer da
-~ '
.
Frgure 7
~
I
I ThE 1deidina groups,
mdude l
~f)
~ Obolella
Athyridid Rhynchoo
_.. Mrcromil
Terebratulid
7, Vel/on
:I)
:z Hcsperor
< ~et> 0 (J) 0 n , I ~ '- n Caenozooc Cyrtospit
Classification 181
tinuin~ pmbll'm rcnuim 111 lW\\ w rrconcilc r1ung.tfl", ll".lrdrop ~h •pt·d 01 hlllth kd. unll>onc.:' u,t,JU~
;U froml11mg .md lo"ll br ,tt hwpod' Ovl,.'r 9'i% of ''" nu! rcrmin.rllv. JlC'tflc k gw" 111g <~lit po,ttTHHI)
brachwpod !-(llll'r,t ,trt' .:xunct uHI th.:n· ;~rl' l'Oll- bt"tWCt'll \'alvc'; l ' g. I.JIIgu/,,, ( 1/1,>111
nblr ditferentt'' 111 m.ttolll) .nul nntt lgt:ny ORDER 2 ACROTRETIDA (I <"uu-Rcl) Sh,IJ Llllttl.u
nr ,uburcular. ,,lf,·n ,.J.,n~tul puhdt ••l'l'lllllg whac
~cell thr VJnom hrJdllopod group,
pn.'>l'lll tonhlll'<l r,, pr.lic k \"aht c g 4auut.1
fhr IIUIII hr.tdtlllpod lJX.t ongut.ltcd 111 rhl' (_;._,,wlrt•t.l. ,,·,,,, tl(t'"'·'
nhro-Urdm·tct.ut, :.md hc:rt· "~ufic:.mt ne\\ dua ORDER 3 DISCINIDA (M . <lrd -Re .)· Shdl n1wabl,
tl'C~nU} bC:lOIIlt' 1\ atiJblt· 111 the torm ot prc\'i- p< Jto:k ope 11111g (.lt lc.m 111 vnuub hdh) lllnn\ 3 tn.ln-
unkm•wu t:nltl.l~ of \ t'n c:,trly ulcar.:om- ~"r nutd1 m p<)\(t·nw tn.orgtn ut pnhd, \ aln·. c g.
ed brlduopods tt'prt• t'tlttn~ J tritical stage 111 Trnu,lli>. u,.,,,,, cJr/.tcu!.•lcl.. 1.
ORDER 4 SIPHONOTRETIDA (U < 1111 <)),I.) :-.hell

oluuon (~ g CJ,,,, Cluk.tl.l). Tht· f.u t th.ll '>ut·h
b!cOH\ L'X, rnuml,·d 111 t< Lrdr,>p-•h•r• d '' rth hc•lln"
wu1 were rctll\'t'tl'd fi nn1 Ktrghizt\t.lll .md >pmes, pc•d1tll· hlr.lllll'll tlllltlu Jlld tpll tl, t' g
trclll~ ~\pl.tll" why tht:v h.1d 11ol prl'viomly
I en rrrurtkd WtiiMtll' rl ,zl ( 11)1)(,) prtfimnl'<l J
"'J'"<'Ilfllft'l,/, '"'"''lt 1/11.
CLASS 2. PATERINATA, ''irh r1H ''"~I· < )rJt·r I'Jtcn111J.l
.a~tJtlttal compulcr-b.t,nl d,tdt\llt JnJly'i' of bra- (I . C.mt -U l )rd) Shrlllllllll<h·d tu t:lliptll.tl hut \1 llh .1
luopnd r~lltron'h'P'· tm ot pm ,ttmg thi' new data. lletr-,rruglu cdg< hl1'1111ll!,l .1 Jl't'Uillllllttl.lll'.l \\ith .1 (uti,.,,
Jl11, proved to be tullr < umt~tl'ltl with the strati- clost"d) (lt•lrhvlllllll l•nlult• proh.,hl) ·''"''Ill,,. g. P.llcnll•'·
:r.•plm rt't:ord of .111 br.u hiopod group~. lt ""tll

bt:
,, fic'Ytliiii/Ttl Diff)'<'lllfl',.
SUBPHYLUM 2 CRANIIfORMEA (I 1 ".nn.-R c: c .). llm·hiopod'
ul'tcd gt:nt'r.!lly .1nd .1 'imphltl'd vcNon '' gtv.:n with cllnrl'll lh slwll' !.1,1..11,~ l<<th .md """' 1,, .md \\llh th<'
low. Jnd tlltl\tr:m.•d 111 hg. 7. lh, hotu the ~tu pedrclc n:dun·d or .1\''~"1 p,.,IJ, it- \.1lv••, "" ntul, ._"'tl "11h
tt\ point ol \lt'\\', 11 '' pnh.1p' llltlottllll;llL that .mu' Smglt· Cl.!" CRAI\: IA I l\.
• e 11111pk, and c.t\lh ltlllctn bncd d.1mtication of ORDER 1. CRANIIDA (?M. l...tlll -H.u ): :-hell rnundcd.
pUUCI:l[C, U>U.Ill~ .f the pC'dtdl"
t OllJOr ordt'f\ ol ·.lltll ul.ltt>d' br.rd11opod'> has
\aln:, brJ1hi.1l '1lVl' tl\ll,ill) ulllltll, 11111sc le ~c~r, pn)-
11 pJi\t:d mw hr~tor>, hut \Ill.: h t\ the nature of
nouuted. c.~. c;,,mr.l, ,\'«li'·"''''• I .d.lu•lntl')''b
'nofic pro!.rr"''· ORDER 2 CRANIOPSIDA (~ . <)11! l. <. rrb ). Shl'l1llltp-
llcal tu !JJtt,'ltlmJ, With m .• rkc.lt un C'nmt ~'T<J\\ th hot"l>.
C.uu Rt•t ) s,lht.lr) mannt· uuputH t.ue In\\ nnl\clt• p!Jrfi•rm pn <'Ill, C'CIIlLIIt.:J O\
ll1't'J ••~elom lit~, IIIC•JUI\ Jh cJ !>ut biiJtcralh S} uunctrical
.lplt<JI '"b""n nl pe,Jt, le \ "'" l' or frn" lyntg, ~ v; CroJtlt(lp.>
mult•> the c-mmm,mrt· pl.u1c Sht·ll cluuuophmphauc ur ORDER 3 TRIMERELLIDA (M l )nl LT S1l.). Sht-11- !Jr~"'
iDfnlU\, pe.itciC' Jftll hc-d, (C IIll'lltCd Ill lrtC V.lh't'' ;md th1rk "1th lllli>Cit: ''dn 111 hmh Hl~c~ •.• m, ubuou
ted l>y nunrlr, t•nclt"'"S hnth .md m.urtk , .t\'lltc,. th.: by hmge-phr,• 111 bradH.,J \ '"'' m .I uuduul '"c kt"t 111
mtr mrh J tilmtcntou' lophuphnrc pcdJde \';tl\·,·. All wcrt• lrce lylllg. ' g J'wm rr/1,,,
::.!PHYLUM UNGUUFORMEA (I . l .un - Rec )· VaJw<; not VttiL'bt•/u;,
J b~ teeth .mu 'nrl..t't§ 'ohdl org.utophmph.ltlt. Pt'd1clc SUBPHYLUM 3 RHYNCHONELLIFORMEA (I l \m1.-RI."c.).
lilY pn:\cnt ftc,h y md nm,<'ul.tr. t"nwr~sng betWt"CII Shell ukJrt•nu~, ,·nclnpnnrnrt·, lllljlllll<'l.lll' n1 p"·udnpu1H-
e> t>r tmrn the .rpcx ut .1 \.llvt·. l'l.mkm hr\'1 . t.ltt· ' I cnh and mrket' prr~t·nt. rhou~h ~· lllll'tllllL"' lmt , ... , -
ClASS 1. UNGULATA (L. ~ . uu.- Rt·,·) llr.Kh•optld~ With oudJ.nly. Lnrr.1 usuallv prt·st·nt, somt·run~s pn1lnngcd ,ts J
ul)\Jnopho,plunt \'.tlvn l.ttl..lll~ ttt·th nr sorkc.:ts. v~lvr~ brKhJdium. VJivcs npcncJ bv dtdunm~. rlnwd by .Jdduc-
~rtJthl"d b~ 111usrk~ .tnd btldV w.tll 1m ly Must ul.1r pcdrck ton. Pethck .tnsc> lrom 1.n·vJ1 rudnm·m. Uut Without ,mm.
fonncJ ,1\ ;,ut.gn•wth Cllnlullil•. lnphoplwn unsupported. CLASS 1. CHILEATA (L. C.un.) Shc.:ll stmph1c. m.trtlntlJtnl.
Gut with tn1c <Ill Us. pedtdc: V.IIVt' Wtth cJrthn.11 llllt'f;)l'l'J, Vtlltl.ll V.livt: pcrf.>-
ORDER 1 LINGUUDA (I t ·.1111- Rt•t ): Shdl bKunvcx, ratl.'d. Siu~k OR.lWR CHII Ell>A: q; Ut~lt .
CLASS 2. OBOLELLATA (L . <. J111.- M. t .nu) '>hell cdk.lrt-
ou>. br.:onvt•x, mbt lrt uiJr to uv.tl . l'wuduHitUJre:l 111
pedidc \<lhl', p<NUnn of pt•dtdt· l>pcmn~ v.lrtJbll'. bt•ak of
'VJre 7.16 Time ranges and relotionshtps of motor brochiopod br,u;h1.11 v.tlv< nurguul S1n~ll' l lRlll:R l )llOLELLIUA.
~· with represenlohve genera illustrated The n~ne dosses e g. 0/~tJ/d/,,, T,. ,,,,,.,,,,,,,.
lluCie ungulate (Li), Poterinoto (Po), Craniata (Cr), Chtleoto (Chi. CLASS 3 KUTORGINIDA (I. C~u1.-J\1 C.11n.) Sht 11 t rl-
lolelloto (Ob), Kutorg•noto (Ku), Strophomenolo (SI) and
S:yochooelloto (Rh) Genera illustrated ore 1, Lingula; 2, oreous, Oll'Oll\'<'X, <.Utl111.11 .rrc.c iu lmth \ Jh "'·
l'.iaoolllro, 3, Crania, 4, Dictyonella, 5. Orthoteles, 6, Triplesia; ddthynum and nmmhynum pr~l'lll hut !Ill tcl'th, "lCkt·~
Vel/cm10; 8, Eoplectodonta; 9, Chonetes, 10, Dictyoclostus; 11, or c.trdm.ll prore,;,, ,. g. Kill•''~"'•'
~is; 12 Pen/omerus; 13, Plicirhynchia. 14, Alrypa, 15, ClASS 4 STROPHOMENATA (M . C'un.-Tn.Js.) Nont111ly
yrlospirifer; 16, Thecideo, 17, Camposilo, 18 StiphrotfrY,is. t·cmcJvn-cmwe'\ nr phnn-l on\ C'\ shdl\ "llh ~ln>phu
182 Brochiopods
hm!'(c. \1111pk· tn:dt, nlll'll l<"l· ~hdl u~uaUy pscudopunc- hous. !:>ubord.:r. mtludt
tJtt·. 'uml' umhnn.tlly <C'IIIl'lltl'd, rubul.tr ~pmc~ may be SUBORDER 1 PROTORTHIDINA (L. C1m.): l'nrntm
pn:)l'lll , luphophurt· \Upport\ r.nt>h found. Cardm;~l ortludc,, l.td,.tng .1 ~,,1rdm.tl proce'i.\; e g. f>r,•t. rrhu
pru<<'" oti.cn htlnhcJ. pnhch." fitr.unen often do~ed. SUBORDER 2. ORTHIDINA (l CJm -Pemt.): :-.lomu!Jr
ORDEI!l STROPHOMENIDA (Ord -1 n:t~.). Umallv con- tmpumt.Ht', find:,. nbl>nl 'hell' (other d·-.q
cwo-«llt\Tx. 'lllllllth or ",th hne nbs. srmplc or <;UPERFAMII Y f!Nl rLETACEA m whtch
redlltni renh . ,hdh 1ft:' pumtJtc). ~ g. Orrlu.<, flrtm•rtlu<. I 4/,
SUBORDER STROPHOMENIDINA (Ord.-Tna,.). ~ JJI>y.·ll.t 'W-ili •'1'/""M, D.rllllltufll,t.
Pl.mu-rnnn·x tu contJn>-u>nve:-... ulicn small \\,th ORDER 2 PENTAMERIDA (M. C:un.-U.
tine rJdr1l nl\\, ~unplc tnth uttcn rcp!Jccd by denu- l3Konv< "· tmpunct.nc, mu.tll)' non-,trophtt 'htll•
de,. c.~. b>JtftH,.,l,•lll•'· ::;,•uHbytiJ,,, Pltctmnb,•mlf-'. 'pnndyhum m pcdadc v.Un Jnd "'mcttntt.., J p.u
SIT<'I'h<•<l,•lll.l vcmc.1l pl.ltt' (crunhum) oppo'm th~ \pund,lium
SUBORDER 2 ORTHOTETIDINA (Ord.-Trias.): Pedicle whole cndo,mg the mu,dc ovar-, Suborder; m,lu,Jc
nun-fUll< th.m.tl 111 .tJult, llll' pedtde v.tlvc IS ofh·n SUBORDER I . SYNTROPHIIDINA (M C.tm.-l Ot'\
ccrm:ntnl, l<Ttlt \tlllplc, J '~nr.U lophophnrc IS Pencamendc, wtth pronlJlll'nt fold mJ <uk111 11111
mfcrrl'd lOt m.ltly ~t'llt'r,t, e.g. hmluua, OrtlliJittes, open dclthyrllltll; , ntr~ltlllll rJre; c !;· PN<Jiil/~•nim,
,\lrrk·t•/1.,, St't'J'Itt'h y111f11a .. SUBORDER 2. PENTAMERIDINA (M Ord.-U D '
SUBORDER 3 TRIPlESIIDINA (L. Ord.-U. Sil.): Llr!l;t' Mrougly bttonwx ~helb, usu,,lly l.t,km~ f.,IJ
l3t<'llt tVl'X tmpum t.tl<' .\hell~. pedide valw intcrarc.t and sui< us: rrur;1liuru wdl developed, c ~· Pem.tml'n•
wllh .Htht:'d pwud<•dcltidultll, br:tcht.tl valve mtcnre:t Stntklmrdi,l, c">idwwl/a
l.tdcin~ wtth Inn!' fiHkcd L'J.rdmal procc~s; e.g. ORDER 3. RHYNCHONELLIDA (M Ord.-Rec.): ~h
Tn[tft•\ltl nonually nnn-,lmphi<, mu.tlly wirh heal.. Jnd htt""
SUBORDER 4 BllLINGSEllOIDEA (L -M. CJm.). tioaul pcdtclt·: dclthyatum p.1rti.tlly c:lo~ed; crun 1
Btcnnvl x "cll-d<'veloped hmgc, mtcrJ.tras and br,tchi.ll Vit·W U~u.tlly Wtth ~o.tr.e nb~ 11\eCOOI-: alo~
cceth. \tlllplc.- , m.Jut,tl pro~.,"· \\ tdclv illvcrgenr 11gz;1g commt~\urt· ..111d often wtth .1 prunoumcJ IOIJ
mu~ck '~ .1r.; <' .~ Hil/ut~.'!·/1.1. .md mkm. Sptroloplwu~. Muscle: p!Jtt(mm d~•ckpl'll
SUBORDER 5 CLITAMBONITIDINA (Ord.): Wu.lt·- only m ~UplrfJnulv St,·non,mJWl'C<~ 1mpun ~ T ht• C.l
hmg~d unpun.wtc pscudupun~cne 'helk havtng except for ITil)llO~<·ncnc Supc:r1.umly Rh\ mh0pon;:
;~gc, 1nc
pronomKcd pwudvddutiium, e.g. Clit<IIIJ/•••'111<'>. e.g. Rll)'"'"'lnt'llll, fJu~uJ.\, Cam•lr•''•'t'clull, Grb/•,rlr)
I 'mmulrh, '/(tmrllyu,lua. ll'rl<<l/11<1.
drtined
SUBORDER 6 . CHONETIDINA (l. Sil.-Pem1.): Small ro
IJr~t· piJrw-tunvt:x shdk Wtth prommenr ~pin~ ORDER 4 ATRYPIDA (M OrJ - U . De>. .): lmru r here J
.tlon~ the pn!tdc vo~lve mter.tre.l; c .~;:. '>tlt>plu•diC>ttt'tt'5. and btconvc:x ~hdt, \\ uh 'hort lunge lme Jnd ruml\1 hnguhtc
CII<>IIIU• . Jb~cm tmcr.Jr~•h. l)pcn Jdtlt} m1m nr umbonJI pcJtdt th:u '011
ORDER 2 PRODUCTIDA tL Dev.-?Tn:ts.): Shell usuilly foramcn Sparalr.t m.tv be dor-.tlly ~·r IJteralh dt \\ h<lk· I
pbni>-H>IlVt"X, li\UJtl)' \\lth deep OlJ/ltlc body C.lVJty. e.g . . ltryp.r, c,,,,.Y.I/•'· Cl.l•\111, D.tylll, Rct::tol, i)J I~ dnopod
ttn.t,ion.llh <'<11\t<.ll .1nd bi1arrr m fonn. V.Uves ottcn ORDER 5 SPIRIFERIDA (M. Ord.-Pem1.): Shells b
li t< ludcs
gent,ul.lte, mu.1lly prulnnged uno J 'tr.ul'; delthynum vt•x. U)UJlly 'truphll, nltt·n wtth J long hm~;~.: line. t:\}'1'
li20.lll~).
~nd nllloth~ t aum trequcntly dosed. Shell usually \\,th ally ribbed. imer.1re." wt'll developed, e~pcc1all\ m
pcthdt· v.tiYt'. l.1tt:r.1lly d1re.tcd ~rinl br.tchtdJUm, fh~..· (
tubul.tr 'f'IIIC''.
SUBORDER 1 PRODUCTIDINA. Usually ccmentc:d m or wtthom Jugum, pumt.ue, dclthvnum open urci<J! J (Will w r.
early \t.tge~. free lavmg d' .tdltlts, cardinJI pro.:t.-ss bilo- e.g. Spmjrr, Br,Jdt)lf/ryril, ,\lw "'·'l"'!fa, Cyr1111. dw cr<~n
b.ut• nr rnlnb.ttc; e.g. P~t>tltultts, Gi!!atttllprcttluatts, ORDER 6 SPIRIFERINIDA (L. CJ.rb - L. Jur.): U~uJUv ;~nil dosdy r
11 '•ltl~"lllftllttlhl, L>lltyctdt•.\111.•. ~he ll~. 5111llbr in most n:spects to Sptnfc.:ndl, buc nt.
llllh'1llit(.)
SUBORDER 2 STROPHALOSIIDINA. E.trlv st:Jges usually punctate: e.g. Spiriji•ri1ta, Uriplica. Puoct<•.•pil!fc'l'.
hC"forc
t:C'tncntt•d. !Jtcr >rJg<'> .mdmrul by root-like spine\. ORDER 7. ATHYRIDIDA (U. Ord.-Jur.): Smooth impu•
rate shells wHit turrow hut~t· line; bt•.Jk ofle11 mmrJt fill· ea m b
lndudc:s nt,my biL.trt<' fumts; e.g. StmplllliMra,
CltcttW.IIt:!!"· Cytla.-.wshan.l, R~tlttll•!fruw, l..<•ptttdus. by umbotul tor:mwn; l.tta.11Jy dm:ued \piralia J<>ll u111t en tr
CLASS 5 RHYNCHONELlATA (L. C~m.-R cc.). by JUgum of pccuh.1r Jtld lumplcx fi,nn; c g. 11/t tfll'SL' lUI
Rhvmhunl'lhltmm·, wHit hu.:onvex, cJ.kitc ~hell~. Cotltpost/a, .\ fc,.istcl/,,, J ctr.utmcl/,1 ~ll i.tll -\C"J
unpumt.lll: ur <'tttlopumtJlc:, J taldfied br;~ch1dium may ORDER 8 TEREBRATULIDA (l Dev. Rec.): ll•,onv Jt(H Ot'Cl
bt pre~cnt . .trtKul.lll') wnh \hon n!ln-qrnpluc lunge. mJ \1
b.N' of tl
ORDER I ORTHIDA (l C.1m.-U. J>em1.): Amculata \nth fum nonal pnltdc t'lllt·rgmg through umbon:tl fo~
utu:yuJII) ht<.t,nv('' ,nuplm 'he lis . Delthyrium and Dcld1~ nu m g<·twrally tlll,ed by ddthvn.tl pll1e1.
m the li~
uur,,rhynum mu.lll) op.:n: lardmal protcss nonn.tllv puntt.lte Brachitlnun lunp-likc:, u,uJ.lly plccrolof ablt:. An
pre,eut Shelllmpun.wtt·, r.uel) punctate. Vemr.tlmm- Subotder. mdudc ,h(lncJhfi
clc: field 'null: mu,cle pl.ldonn rarc . Uracludta ab,em, SUBORDER I CENTRONElliDtNA (L. o ..~ -Per." KtJtorgm
hut lophophorc pmb.1blv ~thtzolophou!> or sp1rolop- An:h.li< tcr,·bro~tultd<"~ w1th pnnutt\'<' loop of llt~tl:)L'IlllC
Evolutionary history 183
fonu l'.~ Cnllf•'llt/1,,, lmplu~:rm<~. ·''""l?•lul'lt.t/us the tir't orthtdes and penramertdes, as well .ts the
SUBORDER 2. TEREBRATUUDINA (L. lk~.-Rt'l ). Short first rhynrhondhformes. The earhest cr.miitomt
IOt>p, mc.-dJJn 'eptum um.1llv Jh't nt, mtcm.ll ;pu:-ules Phill~t•dra ts doubrntlly recorded tfom tht· M tddle
J~vdnpcd: <' g Dtrlmmu, IC rtl•r.uuf,,, Prl!•'Jif.
Cambn.m, but tlm <>ttbphylum dtd not really ger
1'/urc•rlsrm
SUBORDER 3. TEREBRATElliDINA (I nd' Rn .): Long under way until the Middle Ordovin:lll
loop w1rh c.u-Jinalia .111d tllt'th.w wpttnn. e.g. Brarh10pod~ .1 ~ a whole had beconH.' quttt' com-
Ha~l'llcllllll, Zt>tllma. D(~<•lft•l/,,, T<•rdrrt~lc/l.t. mon !orally m the l:tte C,1mbnan; tor ex.tmple,
ORDER 9 THECIDEIDINA (Tn.l,.-Rl·c- ) Sm,11l d11Ck- swarms oC the orth1de Orusia in Sc.tndtnavt,1 cover
~hdl~d trtlcul:ltc\, U\uaUv cen•emt'd .mJ w1thnut pedi-
endless bcdd111g planes. It wa' only Ill d1e l'.lriV
dc. <JpJble of opemng Vt'l'\ wu.k Pry, hulophe
Ordovician, however, that there waoo; .1 n.'.ll evolu-
rtct,\td deep \\ nhm gnnular 111tc:nor. Thq h.wc been
rtg,Jrd~ll v.mou,Jy J.~ \trophomcllldc(, !nd>r.•n•lide~. or
tionary bur\t, .md it waoo; mamly confim·d ro the
rtnfcntlc,; l:i.llcr ( I'-NO) on the ham ot \hell rhynchondhfc.mnc,. The linguhforme' and aam-
i:) •w,co,trucrurc refer- them to the ( )nkr ..,pmtcndl; e.g. ifomte) never thereafter played more tlun :1 \uhor-
11ttil•l. /.<Jm::t'//,1, ,\1,,,,,~1/mo. dmatc role, except locally.
l)rdt·r llllll'rt<llll lly early Ordovician times mo~t of the rhyn-
SUBORDER DICTYONELUDINA (M Ord.-Pcm1.): choncl!Jform groups which canw to domin.ne the
Snull group of unu~uJI hradm..1pvtb m wluch umbo
Pal.Jeozot<. wen: becoming fully ~~tabli~hed, includ-
ol pn.lidc valve j, 111~rkcd wtth pl•wh.lr rnangular
umb<>t1.l! pi.Hc, c g. F.idllt'•tldta. ing the e.1rly ~lropbomcnide~ and rhym honcllidc~.
;ll
while th~ orthide\ ,md pentamcride~ cmmnued to
r-
tn
diversif) Then came the late Ordovtnan tee age.
;a which h..1d J significant!) adverse dll'tt on br.l-
Id chJOpods. though none of the maJOr t.L\J lwc;lme
~d extmcr. Dunng the Silurian there wa' 'ome rt•duc-
Ill' braduopod~ .m: of lower Carnbn:lll tion m the dJvCT"'lty of inamculatcd bradllopnds,
.l;
-t, ~nJ tht.'} arc readily diffc:n.· nnJted uno the three and of ortludcs and mophomt>mdes. though rhyn-
fincJ 'ubph} la wh1ch pcl"'ISt unnl the prc~cnt day. chonelhde' and ~piriferides continued to cxp.md. In
..JtC lhcr.: .m~ 'ud1 evident difference' between llecent the uppcrntO\l Sllunan to Lower Devolll<ln the last
,.,r ~•guhfurmcs, cr:miifonnes .md rhynchonclhformcs of the impo rtant o rders, the TerebratuhJ.l, was
ell' Jl )OOlc ,Juthoritics believe thl· b1 .ll hiopods :tS a added, probably derived trom an .tthyndint.• ances-
cd. holt- to be polyphylccric. In thl' VIe\\ the bra- tor. During the Dc,·onian the sp1rif~nde~ reathed
llicl. opmh Jn.' J 'dadc nf org.Jmtatton' wh1ch thei r pe.1k. and there seem~ to h..1ve been .1 gcncr..1l
on-
luJe-. ut her lophophorate\ (plwromd' .md bT)- brach10pod expansion. By the bte Devoman there
1-
thl'
.»n,) . was a general decline and \.\ idespread cxtmcnon,
\ llh The t.h\'ISIOn of llrach10pocb mt~) thn•t.• ~ubphyla and the pentamendes died out. In the later
, ...d illi~nh ·t ul., I YY6) implic' lllOllllphyly, though Palaeozmc there was a new bunt of cvoluuon. par-
lfJilllfnnnes and rhynchonclht'lmnL'' .m: more ticularly 111 rhe Order Producttda. wlm h mdude
null , 1dy rd.Hl'd ro each other th.m tht.•} .1re to the many large .md pecuhar, though htghly fi.mdtonal
&inly n~'llliformes. There is uo record ut" br.IChiopods genera. These first ::tppcarcd in the '-,llun,lll and
llttorc rhe Cambrian, ,utd .tllhough some extended through the Permian. Thetr grl'.lt success
._l llt'-
·,lt.:d (ambnJn 'braduopods' or
teardrop slupe with was partially became they ~ucces~fully tolontzed .1
•tncd Kentrit nngs werl· descnbcd 'ome yeal"' ago. habitat previou~ly closed to braduopods: the
'')'fiJ. e tumnl our to be inorg.lllll'. n·,ulting from gu;ISJ-mt:wn.u or :~lmost buned (and heme pro-
'llll-~calt mud flows. The earlie\t brathiopod, do tertt•d) mode of life. Amongst these art mcluded
nvcx occur unnJ 'ome dt~uncc ,lbo-.e the defined Ct.Jlrllltopr.ldurrui, the largest of all brat h1opod\. The
With
of the Cambnan. llm thl'ir divcl"'lt) ....-..penally Pem1ia11 \.\," .1 rime of great evolutton.H) plastictty
Jlll'U,
$hell
:he hght of new dtscm•l'nl'\, t\ alrl·.tdy rl'markin producndc\, culminating m tht> ongm of
hou•. Among.r the earltl'St arl' ~hon-ltvcJ rhyn- o;ome very p<:culiar Pennj;m group' - t>Spt•ually in
•ncllifoml orders (Chdetda. Oboldhda and Subf.·ltllily Srrophalmiidina (e.g. Rirltt/Jtl{i:lli<t,
:rm): Jlorginid.l), the Billingselloldc.t \\ lurh were tht• Cydacn11tlwria, Gemellorvia) - wluch cololllzcd reet·
- ov.tl 'genitor\ of the Strophumcnat.t (11g. 7.13), and environments and wh1ch, in spite of then ,1berrant
184 Brochiopods
.md q111re unhr.u·hu•pod-hkt• .tppc.trance. were po,t-Jl:tl.lt•07<'ic t'Xp.tmion of the bt.H.:hinpods. h

ht~hly Jd.tpted to p.trurul.tr t'Jl\'ironment~. ~ome of \\,h nor, howcva, a dirett dl\.·u.J(,r onl~ aiiwtt
tlu:'c, inrerpn.•tt•d hnlrtinn.tll) by Rud\\ id... ami Penman t'Xtincnnn' ttid th1.· bivalve\ come tn donu- Fi.
Cowen ( ll)(,~). involved Yl'f) proummg adapta- nance. 'I ht·y 't't'lll tn h,t\l' 'weathered thl' srorm 1 hcs
tinm, such .1~ rh) thnuc tl1•\\ tC..·c:dmg ~uppbnung cil- bcttn (Ciuptl·r 1) .mJ, being the lint m the 'ro~ce t \\ lttt
i:tl] pumpmg. lOioniz~.· tht• 'c 1 tloor .tficnvJrlk 'tayed rher(' as l:nm
r [O\\'l'\'l'l', \\ ith th~.· ~1.'\ ere l'Xtllll'tlom of Ltte donun.mt l nmponl'nt of Ill'\\' but b.1bnced t'C'tn) dllU(
Pemn.m time. 111 ' ' llll h .tll ti)NI ~roup~ wen· tt'llh, whKh gl'ller.tll~ exduJed br.Khwpoo torn.
aflcLt~.·d. t'\ eu tht·~~.· uc\\ device' did nor hdp rhe Mnn·owr, ,1, R udwtl k ( llJ/ll) hJ~ sho\\11, thr the p
prod ut tide' tu ~un t\'l'. Unly .1 vel')· tcw (~uch as tlw mt:mn.tl h.tbit.lt' 'o \lltt.e"t'llll) exploited b)· 11-t SI)Jll('
t:uml~ Tht'l mpmJ.tt•) nw..h: the lt ,tmition to the siphon.Ht' ht\'Lllvc' .m: not thme 111 wluch brJ-o l11
Mc\07llH'. Tht• re'l bet ,\lllt' c:xtmct. a~ d1d the chtopod' \\'t:fl' n·t.•r \'cry \UlCe\\ful. Nher th.mlk' tntn 1
lll:lJoriry ot"lht· P.tl.teolllll gruu~h Uy the end of the qu.t.,i-int:tl.tl,tl producude,. Pu'"bly Jt wa~ c ltkt• r<
M idJ!t: Jur.t"tt tht l.t\l 'PIItf~·raJe~ hJd gone. leavmg when tlw l.ttter were cxt11tt1 thar it was pos~Jbldlf '" dt•.
only rhy udwudhdt•, .111d lerebratulide~. which tht• hlv.llvt.'' tn cxpluit the 111t:mn.1l habitJt ro la te
rll~l'l her \\ ith thl ,lll,tll 'uhorder rh~.·ndcidin ,l Tl);lke higlwr dq~n:c th.m bdim.·. ( .h!tdt1
up lht gmup' th.llll.IYI.' 'illf\Wed until Recent time~. Annthct t:tllnr 1nay h.tvc bct'll tbt· mt• of prtd.i· th\1\ ,If
r. i\ ing rh')' udwndhtk~ ,tre ,tblt• to t'Xtmde their tory ~t.trft ,h in the cady Me~o7nic, the cormnm rhts rn
'ptrolopht·, out from tht• ~hell 111to the w.tter for ent.lll'l of both br.tduopod' .md epif,nm,d b1vah brrc h1r
more dl'ittt'llt fond ~.ttht'l'lll~ ..md pm~ibly also for Although tbt tiN ~t.trli\h wrtb mu~rular .mm in h.w • r
right in!-( ,111 o\'t'rtll rn~.·d ,h,•ll by leverage Agcr kmm 11 fitllll the C.trbomll-rou~. tulw ti:.:r 11 JtJ hv
(II:Jfl7) \ll!-\!!,l''t' th.n tlm 111.1\' be one reJ\011 why \Ut"kt•r,;, ;1 llcxib\l• lliOlllh fj ,)Jlll .md ,111 l'\'CI'\1 2 Jlt ' •
rh) ndwndltde, \Ill'\ I\ l'd hut ~pintl:ridcs f.ulcd w ~tonuch did not .tppt·.tr until the Me\tl70lt . tlt\-ali'Cl CXlllldt
do so. being priuurily ml:nlll.ll, tn my tJ\t' h.tn· lx:l \\ c aknt
Thct·llktdmt·:. (l.'.g 1~~~:.1.::.•·n1) .ut· .1 ,m,all ~'foup of ddcnn:s .tg.1111~t predator,, wht•a·.,, li\'lng t:ptla bc~·n l
t:alc.trl'oll\-\hdh:d ptycht•lophou' Jrticu!Jrl'' whmt· 'e"ile braduopod-. h.l\'l' none (other dun a rt'pt' Jncf 1311
.tffinittt·~ han· bet'll dt ... putt•d, bur on rhe basi' of t.J\te ami prdi:rl'IICt' tOr cryptic h.thn.w;) Thus the ub
'hell 'tn.tlturc .trr llll\\ rq~arded a' \piriferide~. Tht'\ Me,uzoit br.achtopod~ ..th·eady n.•tentl\' dt'p1 n~c 111 t
rem.11n an llltt'rcsttn~ t'ntgnl.l Derailed ~tudie' of m.r\' h.t\'1.' been pn.•vented from rc-r.d1atmg m the
these n:m.trktbk br.t..:htnpmh h.t\'l' bt·en mJde by predatory ~t:trfi,h. w whtrh the) \O e.l\lly ldl pw 'el11ut~
U.tkcr ( llJ70. I'JX,\, Ill' lll) re\Crst'tl
Lmt,"'.tltfunnt·~ sel'lll to h.t\'t t!lJng~:d little ,jnn: the br.1.
their Jl·dlltt' .H till' t•nd t)f rh~: Ordovician, though \\Ould I
their habiut' h.l\'C.' b1.'t'Utnc le" d1n N.' 1f rhe kno\\'n llllllll 11.11
prescntt' ol dt'<'p-w.ttcr lmguhde' 111 the (;)ilunan i~ a lll.aJc lho
rcltabk gmdt•. Ecology of individual species EnC'n
Lr.tnit!Clrmt'' wt•rt' 11l'Vl'r more than ,1 minor b> till' J'l
cornpom•nt 11frht• br.tthtopud fllln;t a' a whole, anJ Nc:arly .tll br.tchtOpods through t11ne hJ.Vl. been I rh ~ lr1·it
arc n:prcscnted rod.1y only by three ~ener:l, nf thtc. A tcw lm~ulifnnnes t(,und in t-,rraptohtl'\11.11 cocrmte<
which on I\ ( 'mw,t '' t"mmnpoltt.m. and rare tmy rhynchom·llifomtes (e.g. chon'~ IUJIJ . t\ t(
!'he t h.u.ttlt'r nf hr.tchwpod f!Un.ts thm '>cem~ to 13urttm ,md Curry, 1'JXS) 1my h,tve bet·u t·rrrl
hJve ch.tnged ~rt·.nly throughout the Plnnerowic. w m c tf.l t"om·emull h.tbtt.lt wcrc ,tv.Ji!Jblt· U
and even .1 ~upt•rtict.tl .malyw. o( anv fauna will nor- ( llJH4) ltti.·-\trat~~ groupinb" propo\cU tor
m.allv en.tblt· 11 ro lw rl'li:rn d to :1 particular ~y~tc:m. ~tlurian c:m bt I.'Xtt•mkd to covc1 \'lrtually aU
V tnou' theont·~ h.l\'t' h~.·.:n propmcd to account tor thte br:ttiHopod~; thl'\e caregorit:' .1rc not mu 'lr or l't<
rht rt'lllo\rkahl~.· romt'r\',ltt\ITI of po,t-PaJaeozoic br,t- exclustvt but oltl.'ll imergr<tdc. Whcrl'.t\ nem. \ he 111d
chtopmk Recent hrarhwpnds remain ,ltt.ll.hed to h.m! \\hen th•
Compt'tllton \\ 1th htvalw,, e'pccially thmc with 'tr.lle~. nnny l'.tl.rt'o7mt" faun.t' WtTt' o~hlt• to~ md~pcndt
stphons. nu\ h.we been J factor m limiong the ,otr ,edim~.·nt ,ub,tr.ltt'\ " •lt!('< lltlll
rhc 1'~· 1
Epifounal brachiopods Fig. 7.17.1-t:) in which. incidentally, tht• lnphnphorc

support j, p1 e\erved, the mbular 'pme' Wt'H' ll'cd
fixosessi/e brachiopods t(n attathmc:m Ill .1 h:~rd 'ubstratc.
cs n·m1111 .lrtJched throughout Ilk. Thost' Man-. \tll'h Pennian productide' ll't'd tht•tr 'pim·,
Inch h.1w \tom, 'm~k. unhrJm hnl pt'(ildc~ .lrt' in thi~ '"·L)'. The Lower Pennian C/i<11/MT((!I'S (F1g.
no11n ,,, plcnipcdunculatc . Ml>~t modern bra- 7 .17J-n) I~ ll\ll,llly lcllllld CCilH.'IItl'd [(l J.trgt r br,l-
hJOpod~ ~n· ot this kmd .tnd .1rc nonn.dl; Jtt.lch~.:d chiOpmJ ~ nr coral ~. Here the ccmentt·d b.tw of thl
ro rurk~ md dl'.1J shdJ, on the \t':l tloo1 b; lllt:Jil\ of ~hc:l11~ tht, thl' atltcrior p.lrt is raised up .md !Jxcd hy
h' reJilk. f:n,\il pkmpeduncul.Jtt' br.ltiHopuJ, are- tttbu!Jr ~pine\ trorn tht' pedide valvt·. .111d tht
111Ctlllll'~ t"lnmd in 'nc't~' or dmtt'f'. bradua.l vain· bcJr. littlt' funnels ut uurert,\111 ltllll'-
In ~onll' br.llhiopnd, the pc:didl· •~ dtvtded l1on arranged Jlong the edge of the \hdl.
thin thre.llh likt· by,,tl\, whtdl l.lll pt•nt:rr.ue
e r01•t' iuto tkep-,eJ ooze, tnd .ltt.tCh rhetmdve~ Liberosessile brachiopoc/s
dtAd t\1r.uuiniti:tid '>hdk ~ttl h rhizopeduncu- Mmt brath10pods \\,th a dosed pcdick opt:nmg
bte ~tJdnopmh as the: livmg terchr 1tul.~rc.m (e.g. m.my su ophomemdes, 'ome 'pm le; ndt•' m
dtli,•II<IJ!lh•t•l .111J tht: rhynchnndltd~ ( .ryproptlm Jrt' orth1de~) IlllJSt havt: l.nn upon the ~ea Aoor. Such ,1
1 m.tbk tn tllthnr themsdvt·s nn \oft ~uh,lr.lte' and hab1t i~ r.ue 111 R.et enl brachiopod,, and d11~ ntJV be
·lt111lludt: uJ lite: nu~ have: bet'll quite tOilllllon 1n linked With the llltrc·ase in bioturb.HWII ~IIH c the
luopud, gennallv. The pcdide' of hr.tdnopod~ PalacozoJ( Brat h10pod~ that lie freely on thl. o,e,1
ve rrm.lr).;.lbk· .lrtJchment \trength' 1nd m.my floor J~ .tdult' Jre tc:rmed ambitopic M.m~
1 lin· .1humiJml~ in ,ubtidJI currt'llt 'trt'llhrth~ nf 'itrophomenide, h.1ve a dish- like rnumbent lulm
• n1) . 13rachiopmh do not. thc:rct\.lrt•, 'l't'lll to he that ~prt·ad' the we1ght evenly. Chont:tJdint'' h,IVl'
ludcd from tttrbukut cnvtromm nh b) ;m} 'pine\ along lhl' ro,tcnor margm of the ~hdl \\ hidl
:<C'Jknc\~C' 111 the p~d1de (T ha~n. 1'175). lt ha' m.1y h.lVl' 'upporteJ tht: recumbent \hell hk.e a
bfen rakul.nt·d .md ,hu~\ u t•xpetlllll.: nt:dl~ (Blight ~now~hot·. Altt·marivel) the shell nught h .1\'l been
d Bltght, 19!JO) th.tt ~ -lmtrtJ.'f"'1/l'"l, .1mltmed to hdd upright wtth thl' spmcs plunged vertlt.lll~ in
'ubwat,\ bv a lung U.lin pedidl', \\'lHtld .tnually the: o,edimenl I he Si lu rian spiritcrid CyHrtt l.ty R.n
lt Ill the w·1ter in much the ~.um.· wa} .1\ .1 kttt: flies upon 1t~ expanded inrerarea, and lhl. ~lcndt:r
~~the .m. Tht\ v.ould only happen .tt .1 current conc-likl· pl'dtdt· emt:rgmg from it could h.lvt• had ,1
lonty of >11.5 m , '. I f thi' t.tl.:e' piJtt.: .1 rt gwn of 'tethering' ti.1nctton but would not llJvc: been sup-
n-erscd Ao\\ t.tk.t', ploKt' round the ,11\tl'nor t•tlgc of portwe. Pl.afoml\ md ·wmgs' were \Omct unes
e br:~chtOpod. The ,tdv.mtagt· nf 'ud1 'flymg' dt:wloped, givmg maxtmum support on the ~ur6ce
uld be enh.mred tllter tl:t·Jing Jue w 111 ' ' 1th of the: ~cdunent, \\hi le Atrypa had the 'hell extended
nm1al clli.m. 'mcc the ·reYer,c:d thm · could be ·" a bro.td fl.m~e. Co-,uppomve J\SOCIJtiOil\, wtth
••~m~ ul Uuna.l would ahu be ,woilk·d . br.KhJOpnd, nghtly packed in .111 umbo-down
Encrusting hr.1chiopod~ attach to hard \Ub,tr,ttt'' postun·. o~rt' \OtiJetinws found. In tht''>l' the t lmt'
.Ill' peditlc v.tlvc direttly Ulc: !Jrva h,l, settled. In trowding would gJ\'e mumJ.l support Tlllc; kmd
tnt' l1V111g Ct·,mia,' <llld ~ts tm~il rel.lliVt.:\, tht: of rdatiomh1p h lllO'>t common in pent.llller:~ce.ms
·ncmsrctl ~hel l follows the contour\ of tht• 'ubstrJ- and other thick-shelled brachiopods of t,ml>
ntOI. A ti.·w ~duri.m rhynchondlitolllll'' .tn~ likewise high-ent•rgy, opL'Il '>hclf environments
ru~tcrs. ,1\ .1rc some Lngul,ttl'' (l·.g. Orbrwlotdm).
eh .tn encm~nng srratcgy wa' often umtined m Endofounal brachiopods
wly ontogt·ny in umbonally cementing bra- ull.l!rt/a (q \ ) I' .1 htghly adapted burr owing bra-
ilopod<,, .md j, be~r known m produtttdt:,, "here .1 chiopnd ""hich j, .1bk· ro live ctTccnvch 111 '>Oft ..ub-
Jr or oc.Hnx often nJJrks the umbo l,f the: ped1de ~trJta A f.ur numbtT of 111\tance'i of fo,"l 1ndt\ tdua.ls
Jln· indtc.Hin~ attJdunent of tlw jli\'C:nilt: sht·U. in ltfc pmltlon h.1w been de~cribcd. ~onw ~llurian
~'h~n the \hell h.1d grown enou~h to bt• ,t,1bk ~penc~ of l.ltl.(!llla, ho\\ t'vcr, seem ro h.ln' Otlllptcd
l. Jrpendently, ir broke ofT .tnJ bcl,lllll' lrl.'c-lymg boring,, m~n.k .md 1b.mdoned hy othl'r or~.umrm m
ll;r<l_~t'llnwllfhtt, q. v.). But u1 \ll( h produttJde gener,1 large cm.1l rolonk\.
tht' Pt:mH.lll hrhrfcr (Subtmkt "itruph.th.Nidma; Though rhL· ini:m1ul niche so \Veil t>xplattt·d b~
186 Brachiopods
/ .J/1(!11
lormc
stro p!
00
(a! (b )
p.art1al
cover
~Jid t•
7 17)
J CCtcd
.11 the
late r I
wirhd1
,;
·~~~
(cl
CJt Falafer
(d)
Waagenoconcha
lwpl:
<"rt'\C C
'' ould
lllt'tll ,
l.,llllOl
dutlid
(f ) which
IIL'Jllg
fl tl<lgt•
l) lind1
,, hi eh
mg <.' :
.Jtt~c h ~
l:~rg<.' .1
floor.
anchor
\tar a
Kochiproductus
Figure 7
(o) Falal
lophophl
hurd su~
product le
,. hon ofvc
l>ryoz001
Cyclacantharia Fr.rmion
of how s
IJroch,ol
(k) Kochiprc
infounol
ping voh
Kochiprc
uxcludin~
mtermedi
parttcles;
{Perm), (
nelson~
1howing
by tubule
---
(m) Chonosteges
1966o, I
Poleontol
Sheills, 11
Ull~11l,1 wa' nev~:r tull~ '''lonaJed b\ rhym:honclli- [n the ca~c of Kadup~tltllldll\ (F1g. 7.17j,k) the tr.ul
funues, many prod m nde~ and perlup~ \Omc:: other fonns a wtdc, R:n, horizontal R,mgc whiCh must
s•rophomc111des 'ecna to hJ\t' 'pent tht:tr ~duJt lives have lain fiar on the ~ea floor. Experiment' on mod-
pun.illy buned in wdimt>nt .and '' ith a \edimcnt ds \\ith a flow rani.. (Sheilh. 19nR) 'htm cd rhat the
cover over tht: com'.I\C: ba.adu.•J \'aJvc. Thc:sc arc \>.ide Range acted .1~ J \Cttlmg table for cxrram.ou'
wd w be qua~a-int:•un.ll. Adult productide~ (F1g. and incdjble parodc\, while: 'rnallc:r pArtldc~ th,lt
- 7) were .mchcnl'd b~ tht: \lrong 'pines that pro- passed through the tirc;t bafHe \\ere tr..tpped b; talc-
e<tcd from the pedKll' vah-e. J'hese 'PUles. f<>Jmed olat: extended tikt' a rO\\ ot ~take\ at the entrance to
&! the margm of tht• valve, were ori~nall} hollow. the mantle cavuy.
..uer beconung tilkd with t.llutc as the mantle The quast-m£1unal mthc doe~ not \ccm to have
l\1thdrew ~1ncc the marg1n' of both valves are been· adopted by any RccctH brad11opod~ .
ilurply flexed upward' Jnd dt ,l\\ n uut mto a narrow A rather unusual Jdaptatmn of the pcdide h..t'
JCsccntK flange or trail, only tlli~ dun creKent been de~cribed in the Recent tercbratulide ,\la.~(ldtlla
would shO\\ ,lbO\c the gl'neml level of the sedi- crrllriiiJZi. Tllis tives in .1 h1gh-cncrgy c:nvironment m
m,·m, and the: brachtopod Wllllld Lhus be well drifting, mobile :.hell sand. Mo\t !>hclls arc partially
tlllJOUflaged. f t tltl,(!l'II!WIIc/111 (f"tg. 7 .J7d-f) is a pro- buried with only rhe1r anterior edges promading; if
,lucnde w1th dd1cJte 'PIIlt'S on the: br.lclual valve their shells are butied they push thctmdves to the
wh~eh probably prcwmcd ~etlled sediment from surfuce using the finger-like pedicle ,., an elevator.
h~mg wmnowed JW.l)'. Dunng the ontogeny of These observations (Rich~nlson and Watson, 1Y75)
tl;l~~t'II!WIIc'/lrl the ~ceding larva Jttached itself to extend quite unexpectedly the range ofkn0\\'11 ~ub
{Vbndncal bryozo.lm 111 lhc ~Jlllt' envirotmlent, strate rdatlom in br;~chiopodo;.
llhJCh 1r embraced Wl{h nne 01 mon: pam of curv- A very few living mtnomorph1c brach10pods live
g cJrdm,Jl ~p1nc' .wd to '" htl h 1t remained as interstitial fauna 111 mana1c \anJ'> and tt 1s possible
tuchcd for mmc umc When the shell became roo that some nny fo,o;tl hnguhformc\ (e.g. acrotretide~)
hrt,'t' :md hcJ\)' 1r broJ,..c ntT .md, landing on the sea could have done the same:. Some brach10pods lived
toor. gre\\ long ~tout 'pine' wluch it used for tn nc~ts or clusrerc; whme early members anchored
411Chorage m the mud, e\'cntu.•ll-y becoming buned themselves ro a chance dead sht>ll and later the1r
tir a' tlw tr,lll ((;r.uH, I%Cu,b) own dead shello; prov1ded a ~etclmg pbce for later
generations. 1n o;uch ne~ts at lca~t three :mnual
broods have been found (I lallam, Jl)62). conraming
brachiopods of rhc same or dttfcrent spene,. The
nswe 717 Adaptations in Permo·Corbomferovs Productido: nests came to .1 sudden t•nd, probably by bc:mg over-
!Ill folofer (Perm.), interior of pedicle valve, (b) the ptycholophe whelmed bv the ~cdunc:nt wh1ch prc\en·cd them
~e, (c) reconstruction of Folafer, attached by sptnes too
'Dd substrate (all "' 2): (d) Woogenoconcho, a Permion spiny Spiriferides, which ~omct1mcs grc:w in ~anul.u
·~e. reconstructed, (e) median section showing disposi· clusters w1th only a smJ 11 b.J~.tl att:tch ment area, are
r.-1ofvolves and spmes, (f) juvenile of same attached to on erect sometim es found to bl· ;~~ynlllu:tnc::tl Jnd distorted.
:ryozoon (all ~ 0.8); (g) Cyclocantharia, o highly modi~ed probably through crowding b> adpcent ind1v1duals
=m•on reef-dwelling prOductide (' 2), (h) Rudwick's conception
~how svch o brachiopod fed by Flopping the valves: o rising (Ager and Riggs, 1964).
11ochiol valve would Rush out the interior; (j), (k) Recent plen.ipcduncubrc brachiopods have the
txhiproduc/vs, o Corbontferous spiny productide of quosi- ability acrively ro onenr.tte thcmsdvcs w1th respect
·louool habit, shown (j) cleaning sediment from Range by dop-
"i!l YOlves together (>< 0 4), (k) cross-section through Range of to current~. and the pOSitiOns taken up by livmg
~iproc}vclvs, showing various sediment boffles (1, spines spccimem 111 area~ of current flow coHfOml to
u:Wiog large particles, 2, Range acting os o settling table for hydrodynamic prcdicnom
~iole particles, 3, toleoloe (endospines) trapping smolt
lf rhe anrenor-pmrcnor ax1~ of the -;hell "'as par-
;cr.odes; 4, only food particles get through), (IHn) Chonosteges
111!1 ), (I) in life position, dorsol vaew ('< 5) (m) side view of fun· allel with the current (and cspcc1.1lly 1f the excurrcnt
illonbroch•ol volve (><. 20): (n) medtan section of Chonosteges, rcg10n faced inro it), nhary pumpmg would actually
~ posterior cemented port, rotsed anterior port ottoched be oppO\cd by pre-;surc d1srnbunons. If. on the
t'Y ~llulor spmes, and funnels (x 5) [Mainly based on Grant,
other hand. the liVing annnal lud the JXlS perpen-
~ 1972, ond illustrations in the Treatise on Invertebrate
ei!cntllogy, (h) bosed on Rudwick, 1961, (j). (k) redrawn !Tom dicular ro the current (.md <:'penal!} 1f one of the
s 1968 I incurrenr reg.ons taccd mro 1t), pumpmg would be
188 Brach iopods
,,,~1\tc:d.Tlm 1s, 111 tan, t :-..•t tly "h.H h:tppt·n\ m many mJrticul. lttd braduopo ds .md \lll.lll plccun
'pedes studini, and brJcbiopo ds may \Wing through bon.itat cans (SI'nwidcl l, Clwnl'l<llil l'o). Till' ~1lty nud
.1 \Ub,tantt:ll .1r~· to t.lkc up tlm prc:te-rrt>d pmition
dle p.lrl of rht 'rectnun ~~ of\\ 1der diwr.ity arnl
rd.1tivc ro prn·a1llll~ t'urrent~ (Lt ll.ubera, I'J77). donun.lte d by tblmencl hds. lll'tnorth ld' and ~
1\nlCtH)n.ll p~:dtdt•' '\l'l'lll to have b~:cn present in plcctamb onit.Kl'.t m suth .v; Soll'l'rf,yello, wlud1 h11
mmt rll\. nd10ndl uk\ and rerc:br.ttulidc,, most rdJllvt·ly widt lanes r angco;. [· xtrapolatm g l\n1n
orthtdes and ~pmfi.·ndcs, some penr.m1cridc:\ Jnd .1 Lockky was abk• to show th.u rh c.:· 1 angt nt br 1rh
few stroph11m emdt, pod ,l\\ociatJOm could be djvulc:c.l mto t'lght 111
tommlnu ty types, whu.. h pef\1\tl'd throughout!
whole t>f thL· OrdoviC1.111. E\'1.'11 though thr ).,rt
eha.ngc.:·d through time, the b.1lancc.:·, tauc~ JM<
Brachio pod a ssembl ages and
uons Jnd gen1..•raJ aspect of the~c.:· c:1ght u>mmu•nt
'commu nity' ecology
remained relatively suml.u.
fo,\11 brarh10p od' tend to he found 111 recurrent
a'semhbg... '· often c. ompc.N~d of se\ cral spenc:,. ln Silurian palaeoco mmunitie s
these ,\\\Cillbbges other mn rtebratcs .1bo occur. bm Early dJs\lC ~tudJCo; of br.1duopod-donun
tht· br tcl11upnds an thu.1lly dominan t. Thc~e assc.:m- palacot ommumt ies were thme of Z1eglcr (I
bb~t·s h.tv1.. been much med in n:n:nt \"CJ!'\ in and ZJc.:•g;lcr et .1/. ( 11Jf,H). updated by Cocb
:mempb tu undcl'\t.llld the 'rrutturc of .mnc:nt McKerro w (JIIH~). whiCh \\'Crt based on rcrurrrro
(m.unly cpil:lllnal) p.ll.lctKnmmtmiuc~ .md the con- Jssembt1gc:~ of Upper I l.lndovl'r y age 111 the \\
trol' opcr.ltmg upnn them. Uordcrl,m d (Ft~. 7 llJ)

The recurrent ,1\,embl• gc' Lhar.tucr ize p.1mcul.tr Du nn~ tht• l:.Uher l!Jndove n a 'hon·hnt
t'llVII'Ol1l llc:nt\ and 111 hfc lllll\t h.lVI.' rdared to such eastward ' from H :~vert(>rdwt:~t Jnd, rurv111g nortl
par.unc:tef\ ,,, depth. tl'mpc.:·r.llun'. s.1lmirv .md \Ub- ward, nc:Jr rhe rown of Lbndnve ry. extc.:·nd(j
'trHc:. Mmt p.iiJCtll't c.>lngl\b worlun~ With rc:rur- north\\ ,1rds. On the 'hdf 'ea bem ee-n tlw ~horc
rcnt •l~~c.:·mbl.iges h.we tnc.:·d not only to detlnc them rhc graptohrc ,Juk arc.:\1 to the west ltvrd o1
in rc.:nm ot thc:ir composiu un Jnd rlw rebrive abun- f.mna. lh the nme the -.horl'lim· had rt•trc.Jtcd to
.l.lmt' of thc1r t:1lulll dt:IIIUlb. but ,1l~o m deter- La~r ut the Malvern Htll'> LD the Uppt·r Il.tndtr.
nunc:. JS 6r .1~ po<:~1blc, the nature ot the controls. the ,hl'l( was much broader .111d shclly taun 1\ ¥.
')o rnud1 rc:cenr \\l..lrk h.•~ bc:en done on thl\ mmt ditlcrt:ntt atcd nwr it. Five Ji~tmcr though £1
vttJI nf p.Il.!l·neculll~'lc.ll fa~lth thJt the bulk of lirer- uonal pJJaeocN nmun•ue ' donunace d h)' b
Hurc: ts comtder.1ble. Only ,, lew ex.1mpks .m· hert: pod, (though w1th '>ubsidjary bJvalve,, g:11tro
\l..kued tor d1scu~,ion coral!> .md trilohHc<;) 1.1\ 111 dl\tinct COliC.l'lltnC
p.1ralld wuh tlu ,horc.:·
Ordovic1on polaeoco mmunitie s Nc:.m~st the 'horc l.1y J Lill,(/11/a pal.worommu
In recenr ve;u' volum111ous knowledgL' of with .tbundan t .,null rhynchonellidc.:·' (pn•l
OrdovJn an p;llJltltlliiU11Ulllt\ stnll·rure h.1, accu- Stt'(!Crll)'llclms or Rvslrlcdt.tla) and Li"i:'t/,,, The
mui.Jted. Ail hough tlm o,tannlla rer tlun comp.uab lc from-'>hore E11C••t·lra p:~IJeocommuniry h.1J
studtes on thL· Silun.m. over c;o difl(~rcnt J\~uuation~ rhym hondlidc.:·s (Et•wdia) bur ~ more diwr'e
h.l\e by no\\ bc:c:n de,cnbed . lt is h.1rd ro stunrnari ze of other brach1opod.,. In the 1-'tlllllmm•l p.11l
.llld mtegr.He all dll' infunn.Hl Oll 1nto <l coherent mumty whtch by tJrthcr out thl' eponymnu1
p1tture. but 'nmc: kq work' h we ht·c.:•n ~llltL'"lul m domm.1ted .111 othcr1, often rc:Jching ,, l.u~~
dom~ so 1 m kk) (19H3). ming du,t...r anJ)y,•~. and 1pcnmcn s an· \OIIIl t11111..' found m hti. J'lNtJOil.
workmg on Anglu-W ehh t:llln.h !Tom the t}'pl' ar.:,1 large: C..•stistrirkf,llldirl W.lS the ll\O'it ahunJ.tllt
of the Ordovici an. has bc:cn .tbk to discun some ch10pod in the penultm1 are p.1bc:oumm
rk·.1r pattnm of d•,tnbun on of fll\,JJ bratluop oth rhou~h <;maJkr brach10pod~ wc.Tt. JJ,o unpoltllt Figuro7
lub~troht
rd.1t1ve ro nngm.1l \cdimcnt type' (Fig. 7. I H). the Outl·r--.h df Clonrufct pJI.tl'OCOTllllllllllt)
Cn.lf\C ,t•dinwm~. JS h,,, long been known. :~re were .11 lca~t .l dozc:n genc:rJ of rc!JtiVdy ,m.alJ
donun:ue d b\ coar,l'-nb bt·d orthacc.m ' (Ht pt n•rtlus, ch10pmh . \O dut tt wa' the mo't dtvef\t oum
01/t!ll'this) , \\ht:rL'·" \"l"l)' tine scdunem~ contJII1 of all Beyond thi~ l.ty .111 ~n·a of gr:~ptobn~
J
l
)
l
llandello
'l
e 1
d
h
e
y Uanvlrn
·e
l-
1-
s.
ts
J
y.
ly.
(-
ar
~c:
he
·a-
J
ty.
In fgure 1 I8 Generalized poltem of focies relationships and substrote preferences of Ordovicion broch1opods through time: coarse
:re dlstrotes ore on the left, with a predominonce~f ribbed brochiopods; fine substrotes are on the right (Redrawn from Lockley, 1983.)
190 Brachiopods
[dr/y Upper llanrlo• ory I mgui.J sp De•
0
N fJ!oiJt.>ogrogrJpny A pn
Q outt.I\)J;)i•tfb
grJlC
b!Jgt'!
p PG/ tlfflel{l~ C()t'1'\Uh..,"l'to
S lOS~!uc. • ~MC'OI'J\fff\11\11\
Oucnt
C C~J tomrnurl•IY tolllw
mU<~>
G
T
(IJilf loll"'
lJrl't\llc
0 priJm
_ / S!lell rnm 1 11 Ill dC'
..._. c: tt-,nt -t.tVC.trOn ch~ml
~ ln<l.or~a tunc
In()ll ,
CAR!liGN'i lh
B·H
hght '
c.r:-iil
111.11 i I)
I CIIIH
111111l'
Ill rnl
Co:;tl.stflcAidncf'b /!rata
lmn t
(j
CVt11U I
jl(htul
I C llO
I .
In
"t1rk'
Ordo•
1en1l\
Ctoflnda globosa
.u;,enll
0 rcpc::ll
Ime'.
'Olltr<
depth
Figure 7. 19 Poloeogeogrophy ond brochiopod polaeocommunilies in Wales m lhe early Upper Uondovery, showing illuslrat!O!U Cl.mn
lhe brochiopod genera de~ning lhe poloeocommunities. (Lorgely redrawn from Ziegler et of., 1968)
p.1l.tc r.
·"core
ddine
:lCtumul.l{ln~ 111 .1 dccper-\VJtcr environment. inclmlmg pres~urc:, watl'r turbulence. saluuty. 'ub- .lhk t
Sin11Lir p.ll.lc:ocommumtJc~ hJvc been found in strate and tooJ ~upply. Tc:mp..·raturc:, however ,I\Sf!lllJ
roch of the ~;mw ,lgt• m othn parts of the world, which den ca~e~ 111 dcepcr \.Vater, .1ccording t punt'll
and in ~Ollll' c~sc:~ thl'Y seem to be equivalent ro Boucot ( 1975). 111:1)' be the most 1111portant of'811 turbuf
thmc Jt'fincd h> Z1t•gkr et ,r/. (1968). these variable~. bi.J~C'
The l'Xampks of Ztcgkr t't t~/. ( 1968) show a reb- Where other ~llunJtl br<1chiopod paJacoconunull!- AIN>h
nvcly st-r:t1ghtf0!'\v.1rd pattern of d1stribution in nar- tics have been Ul''l ribcd, howt·ver, ~uch a\ tk g.ucd
row b.md' '':h1ch were lar~dy Independent of Ludlm;an a~scmblagc~ of L..tw~on (1975), 1t l> k"'' t olrt: 11(
sub\tratl. and p.lr:Jikl w1th the shore. This seem~ to to assess the comroh ol Jj,tnbutJOn. Lawson, ,uggn;.. (0 60
suggc\t th.lt the di,rribmtnn pJttem ~~ in some way mg th.lt substratl' ma> bc more unportant thm dqu t/ ,./.'
related to Jlpth nf w.Jter. Depth alone. however, in some coc.c~ and noung that bracluopods bclonpit TIH
ha\ been gt·ncr:tll} rcg.lrdcd .h a rather elusive tactor tO different assonatlom nuy have varymg tokrana ofien
whose influt•ncc on <ot•dmlcnt~ or ammJI~ is indireCt. rangt-s, reJected artempr\ to unpo~e the s1mple pancm brgd)
It '' not tkpth ·'' 'uch th.u i\ .1 primary control. but exlubucd by the Upper Llandovery pabeoconunuru- they
cc1tam 1mporwu tactor\ th.u VJI)' "vith depth, ries on orhcr ktnd'i ofa\\cmbl.tge. t\\'ign
Ecology a nd d istribution 191
Oevonion brachiopod assemblages nlar BA may be difficulr where there ha<; been an
~ pnnury t'l~k for p.li.H~Ol'rolo~ i\ to try eo lnte- abmpt t'Jun:~l replacement 'o that there 1s no conti-
tc the e·mtmg bulk uf d.ltJ on l'.lheo70IC a~<;em- nmty ,.,,th tom1er a~~onltlons However, where
~·' .md to t:\t.thh~h how tlwv luvc evolved in neighbouring a~~onattom h.1ve not changed. then
ifu<ru.mnp; cnnronmcnt' through nme Tht~ \\'Ork, the :membl.tge m que~uon tnJy hl· 'bracketed'
iJIIo1\ing th:lt of Bn·t~k y ( Jl)(,l)) .md Boucm ( 1971), between them and \O referred to it~ correct BA. If
p 11111~ well, bur tlw lll.'l.)nr h.ward enlmmtercd is an unknown J\\ociarion he' berwccn BA 3 and BA
mdctwnining thl· l'tl\'irnnllll'nt.ll rquivaknce of 5 it j, probJbly BA -+. l'Wn though tt " dl'.)Ulll' t.
.!:<Juncr .1~mci:H1om \\hi eh rl·pl.tn·d l'Jch other m Siluro-Dcvonian chom•Ltccan\, ·" dt\tributed
11mr but \\hit h m.•y h.1w tcw ch.n:\Cters in com- wuhn~ bcmhic .l\'\embbge\, ~how a dear n:latJomlup
mon'' ith the1r prcu•r;ors between morphology :md l'llVIrOnmt:nt (Rache-
l he m.~or work ot Bum or ( JlJ71)) hJ<; <;hed much boeut~ 191)0). Choneran·.1m \H'rt• nor unt:ommon in
l.!hr on tlm top•c Ill umkrtonk .1 IJrgL'-\C,llc study the Silurian, but with diveNiic.Hmn of ctwtronmcnts
Jf Siluro-Ocvom:m br.H·hwpod dJ,tributwn, pri- in the early Devoman the> llndt.•rwcnt J lllJJor r.Jtlia-
rrunly to .1'>\C\\ the \17t' ol' brn~ding popnlJtions (in rion. Variom ad.1ptations in ~hcll shape and the devel-
rmm of rht Jrc\1\ thl'Y orrllpinl) .mJ ~o to dcrer- opment and di~tributions of hinge spine~ h:tvc been
minr. how utes of evolution .mu t'Xtmrtion v:1ricd directly related to bathymetric gradient.
mflllpnl.ttiom nf dilll•retll Mll'~. I k "'"'able to con- Boucor\ model of hl' tlt h it· associ;1tions is ~o far
lrnn the prrdittiom ot poptd.ttlnn dymmici,ts that only a provisional ~rep tnw.mis the intcgr;~.tion of
'"lunon would he r.1p1d in ~nulkr popubt10m .md 'palaeocommunity' d.1t:1 .111d the interpretation of
fl!l'lulared .1 complex 'LTil'\ uf hmlogKaJ and physi- thi~ in environment.1l and cvuluumury terms, but it
~ rontrols over exlinn•un evenb provide' a nccc~sJry framework lor future dcvdup-
In csubhshing .1 \r,llllltrd c.ommumt) frame- ment~ of community \tudic' in Palaeozoic bra-
tick' for thi, \tu d), UntKllt rnktu u:d all kno\\11 chiopocb.
OrdoVJnan w De\ Dlll.lll p.d.Jc.'lll.O IIlmunitie~ in
;an, of b enthic a ~semblagcs (BA<.) A benrhic Permia n reef associations
mbbg.: ' ',1 group ut c.unununitie' ou·urring In western Tt·xa~. Somht:.l\t A\1;1 and ~ll ily there arc
~redly 111 thl' '1n1c po,ition rebtive to .1 \horc- inuncnse reds of Pemuan a~c tomposed maml~ of
lK''. AccorLhng tu Uuucot, the\l' \\'l."rc probably algae. bryozo.tm .md sponge\. Tht: Cap1can reef
nrrolkJ l )' water temper.1turc.•. denea<.in~ ,.,.ith complex of we\tern fcxa~ ~~ the best known of
rrpta Holll or tl1us rc.•g.1tdcJ tht: L."~t:"la through these. le marks a late ,t,lgc 111 reel development,
nm/,t 'rommumttes' ut /.1e~lt't t'/ ,,[ (st:t' Siluri.111 fom1ing a g..111t bamer rrct round the margtm of
?WcocommumrH.'\ .1bm·c.·) .1~ l3A' l -5. zoned deep-water basms whlC h 1t enordcd; the~e had only
LOrdtng w di\t.lmr tron1 ,Jtordinc. He ,1lso a narrow outlet to thr <;C,l on the west. 13ehmd the
.ktincd a sixrh l3A lytng st\J\\ard~ of l3A 5. He was reef l.ly large salt flats : ev.1poraung p.m~ 111 wh1ch
,_. k to reLltc :~11 knm\ 11 Ordovic i,m to Dcvonian dolomite, gypsum ;tnd .111hydntc were deposned m
r. •rmbl.1gcs to .1 ~y~tc.·m ot SI" UA~, though the com- sequence away trnm the reef JS the w.:~ter became
eo ~lll!'nts mi~ht v.lry laler.tlly lkpcnding on w~uer progressively more hype rs.1linc. T ht'Sl' were rcpkn-
ill trbulence. ~cp<lrate quit't .md rough W<lter assem- ished by sea w.lter wnslnng over the top of the reef
!,'CS could be disungtmhnl 111 111ll~t nf the\e !:JAs. The reefs .ll'c nor espcn:dly fo~~ilttcrou~ .md contain
ll- IMimc depth, for BA, luvl' now bt·en mvesti- only locaJized devclopmrms of br.lciHnpods, 111
h~ . ~d u'ing st•veral hm·s ot t.'VIdt'lll'l', .md BAs 1- -+ pocke~ of low divcr<;ity. Pnor to the development
ISY ~re now beheved to he wtthtn the photic zone of the great :tlgal reef~. numerou~ smaller reef~ or
st- l.f,l) m), :md pos,thly .111 nt p.m of BA ') too {Brett b10hem1' grew m \utnewh~u deeper water. fhese
tth .J.• I 99J) patch reefs, best expo,cd 111 the Gh1" Mountaim .
ng The compmmon ot BA, dunged through rime. gr<'w up ro XO m h1~h. though they wt·rc muJlly
LCC ;fttn ahruprly. Where.· thl·rt· wae reef,, ...,,th their smJiler and .1re .1bundamh tossJhtl• rous, contJming
~d} endc.·mtc md r.lptdly cvolnng populations, 'ome of the most h1ghly moti!tit·d hrach10podo; ever
ru- were mu.lll}' a\\O{'I.Hed w1th UA~ 1-3. to have ltved (Grant, 1971 ).
~ing .1 given hr.ll hmpod ,1\\l~l'l.ltion to J pamc- Mmr of the Penman h•nhl·rmal brach10pod~ .m.·
192 Brachiopods
of rehuvcly nnn11.1l form: the majority were pedi- example, that Lower Po~l aeozoit nrthides \\lth 1\ le
de-.lttH hcd md .tbh. to cxplmt o;cvcral different coal"\c ribs pn.·fcrrcd .lrenat.eous cnvsronmcnt'i, ~"d C.c
h.lhttat\ withm .md penphcr.tl to the biohcnn. there ts a gcncr.1l tend cm" in the Mesozoic fi1r sun- M,
Otht·r k1mh Wl'rl.' dl'limtdy 'mti-rl'cfal' and lived in ilar anarom.ic.tl feature~ to ou:ur in unrelated ~totkr l:)(l
tht flat 'ednnent ht't\\·ccn thc red\ Orhcro; again in 'imilar cnvuonmt·nt). A~er ( IIJC1S, 19ll7) sho\\ cd Bl
well' rnntinc.:d eo the red. often being of conical how Me\0701l bradliQpod, lJn be related to \C\'CD bo.
tC.mn w1th long tubular 'PiliC' fomung an interlacing m am b1ofaue\, .md the nawre of the mbstratc seems (l.r
mc\hwork w ludl ctmtnbutcd to the reef fabric. to have cxertl\ed a donunant control over dtsmbu- Ne
Thco;e r.nhcr ttlr.tl-likc braduopod~ wen:: all modi- tion. Thus there arc charactcnsnc as~embbgt~ from (Ne
tied \trnphJio,nd produnide) in which the conical ( 1) vel) shallow water. coarse ~edunenrs. (11 Stt:
pelhdc v 1lvc w,t, t'l'lllt'ntcd in the early stages, and sand-grade ~hdf \edunent,, (3) pen-rectal sedtmenu Jt't
luJ tubul.1r 'PIIlC\, ,,, in Clu111ostcges (q.v.), wluch (4) rocky bottom~. (5) mud-gradt.> shelf sedtmenu, J
held them llp )m h 'Pille' wcn1.., to have provided (6) fine-grade, probably batbyal ~edm1cnrs and (7 trnl
.ill 'evolutllHIJI)' 'pnngbo.1rd' for their posse)sOn.. epiplanktonic bracluopods. Consequently. an} ten
~omt• biOhl.'nm were composed largely of assemblage from wry ~hallow watt'r'i genml I' <011
R.Jththoll:ntid.lc (e.g. CC'~drwria and Cyc/acanrlwrit1; includes rhynchoneUJdl'S w 1th co.me nb) .md lar~ott: I ht
Fig. 7 .17g,h). The't: have long. poinred pedtcle pedicle opening-; supportsng stout pedicles. a~ wdl J.l kllll
valves suppmtt·d by tllbular spines. The brachial terebrarulidcs w rth sharp commissure fold'i. A qune 11,1\
v:.lve ~~ red m cd to ,, thm, flat, horizontal plate d itre rent assemblage wa~ to bc found m 'pen-n.'t'fal t.rtl
recc\st:d \VCII wttllln the pt·dH..le valve; ir could open environ m ents, w1th large asymmetncal rhynchon \. 111.1 r
to Xll 0 ur ~0 .md h.td " prott:cllve grille of spines or !ides and tcrcbr.HuiJdc~ w1th elongated beaks. Sl,mt f11lll
mc,hwOJk ,lbm·t· the opt."neJ valve. R.ud\.vlCk peculiar brach1opods, t'.g. the pl•rfi1rarc Pyi,!,>pt, Wtl w.rtl
( JlJ(, I) m.ule .1 ' ' ork mg m odd of one of rhese and confined to c.1lm, deep ~ea,. wherea~ 'm Ill al l
,bowed th.lt tht.• b1.tchiQpod could have generated thm-~hclled rhynchondltdc~ were cvJdentlv t'pt· lmt1
strong rurrt:nt,, hrin~ng m fc)od Jnd flmhing our pbnktomc. cudc
the mantle c.wirv \tmply by raptdly flapping the One \\Cll-mldJed \ection in tht· Frt·nch JurJ, £r lh('
bral hill v.ll\'l· ming tlw .tdducror and d1ductor mus- example, ha' fimr sucCC\'\IYC rhvm hondltdl l!<lll·
cles alrern.1rdy. Rlractorlr)•lldri,t at tht• ba\c, J.\\onated With ret£. ern ,
Though rim intt:rpn:t.trion h,,, been cririCJzed by Acmrtlwtl11ris of nonnal hcmhomc h.1h1t and found Con
Gram ( 1972) - on rhe grounds that the known ~ands, LurmMdla. a~\onated with 'pongl.'\. and 1 lllOJI
o;rrucmre ot tht• l.1rge ptyt holophe in the Pennian the top the tsolart•d Acmrclwrlrymlrra, \\ hich orcun 111tf
producw:lc, (Hg. 7 17.1- c) would make rht~ clays and wa' probably epsplankwnic Likn\ll such
unlikely, :md th,lt the brat hmpod could have fi.mc- other rh ynchoncl!Jtl, from ditrcrl'nt \tocb sllu,tm Sl ll ,ll
noned pertc:nlv \\ l'll w1thot1t such a valve-flapping homeomorphic ton vergence m .1daptmg tn rl·rf.tl or tll.Jr~
mech.1111~1ll - the po"1bihry 'till remains that the peri-reeful hahltat,. B'
nchtokniuko; could have U\ed Rapping valves to bct'.r
o;upplemenr m rcpl.lt't' cthary pumping as a m eans of clu\C'
collt•cring fnod . ,uiJ I
Some othl'r Penn1.1n produnidl's (Gemellnroia :md At
Lcpwtl11s ( l.ytll>tlltt) .m~ even more peculiar and The.: globJ l di~tnbution ofbracl11npod~ is mmt full ~l Clll
alrhough t!1( funroon.1 l morphology of some of documented in the Lower Palacozmc, wlndt 111 ( ;1)11(
the,e pt•tnli.lr ic1nm Jllay be open to que~rion, "llrely any case tlur p.m of the ~cnlo~ral column "hen lllllt.
Rudwick 1s corrt·n 111 \:lVmg that these adaptations they arc of greatest ~tr.llib'l"aphical value and \\lrer (tli\ct
'bc.u Wl(ltl'S\ to .1 degree of pb,ticiry unparalleled at they show an mstrtsttlVC parallel with conrcmpozr f\ ptc
.m~ nth er nmt' 111 br.Jt h10pod evolution'. ncous triloblle,. fc\\ ~
The mmr reLcnL sumnunes arc b) Boucot (19- b'l"Jdi(
Mesozoic brachiopod associations and Cocks and Foncv ( 11)90). Data on brachsllf<l( t
J.lld
In man\ lll\fll1ll'' thl·re \Cons to be J broaJ and dt\rribunon arc coml,tcnt with tho't: from trilob lu~hc
geuer:tl corrd.ttwn of hral hmpod '>hell fom1 with and graptnhte\, .mJ fit m well with model!. dJd 11
cnvlronmt·nt,, lt w.1s rccngmzed long ago, for palacogeographital rt.'<..Onstn.ICtlOm (Srotne II.IS, .1
Strotigraphical use 193
~ld\mow, JlNO) . In th~ Ltrl) Ordo\11 i.m rhe large Silurian brac hiopod fJunas rnovcnng from th"
Gond\\,m.m conttm•nt ('Hllltb AmericJ, Afuc::t, decimauon became co,mopoht.ln, thou~h two 11C\\
M.!dl!-}hl'Jr. I nd1.1. Ant.tr• ttCJ .md Au,rraha) pronnct·s arme Oil\.' t h.u.tt tenzed bv Clmkchl
tendni mm the Smuh Poll.' to \11111\.' ~() N. "hich beeam\.' C\tabli~hed 111 "louth Aml:nc.l, .1nd
BIn ~ (Snndm.1\'t.t tud w~ ,r rn Ru"t.t) by mu- the orher w1th Atrypt:/1,, \\ htth aro'e m the fonner
!lo.lrd bt'l\\ ~·en '\0 .md hll ~ '' llid A ,·aJonia USSR. Greenland and wt•,rem "orth AmcncJ.
Enghnd, 'outhan ln.l.llld and ea,tcm Dunng thl' Devo111an there was htrthc1 provinnal
Nmfoundl md) . I he l.lr~l' Lwrentun lUUrinc::nt diAerenrianon. rl·sultmg t'\'Cntu.tlly in tive fauna!
'-:(lrth Americ.t. \\ e'tem :-Je\\ tmmdbnd and pronnces wluch h,wc bcl.'n full) documcntt·d by
'i.:orbnd) 'tt .t,lridt• tht• ~'llll.ttur.•1~ du.i the mdcpcn- Boucot (1975).
:nt landlll.t\\1.'' of E.l\t A,i.t .111d S1bcnJ. ·n1e stnkmg parallels bcrv>t'cn tnlobne ,llld bra-
A' di\CU,,t·d c trlu:r (Ciuptcr 1). the primarv con- chiopod d1~tribut10n 111 rlw l OWl'r P,ti.1COIOIC show~
'rtll of dt,trihutlllll llf t~1J.1y'o; \hdf-ltving fauna~ lS how pabeob•ogcognphy c.tn be J powerful tool in
tcmptr.Hurc, though the tclaun· i~olanon of the timetabling maJor globJI t•vcnr' .md comrraining
t\IJIIIII.'Iltll l.llldlll,\\WS ,ti\U hJs a ~Jgnitit,lllt role. plate tectonic models.
Th~ same control' upt'l .ttt·d 111 the p.1st, .llld the
kn\lwn discnbution ut Ordo>JuJn bt<Khiopod f.m-
111 h tull) torl\l~tt'lll \VI th them. Tb m there w.ts ,In
,trh· OrdtiVIU.IIl I.HIII.I ol t·ndcnul lingulates and
tJrticul.lti.' ottupymg cold pobr \\',H\.'rs. later M.tny br.lChJOpod gener.1 ,IJld ~penes .1re relatively
Jlllll~d by d.tlm.Hlcllids (TillS llllllptt>d the :.ame long-ranged .md thus ofhnmed v.tlue 111 corrd.ltion.
~Jtt:rs ,1\ the contcmpur.llll'llll' ctlymen.ll c::.tn-chl- This is p:~rncnl.trl) 'o wnh Mt'\OLntl' brJduopods,
ruamt.ln'Jn tnlobitt· prm int t·.) Lik~"" 1\e the low- which though \OmetllllC\ ll'<t:ful on .1 !mal \tak Jre
utudl.' arc." ot' Cond'' .m.1 c.uri~d thetr o,. ., ofles~ ~tratlgr.lphtt.tl .lppltcJbilit) over wider an.'J\.
~nJenuc b.:1chiopod fnut,l\, md \O did l:hltica .md In rhe P.ll.IC070il, howcvn. braduopods have
d1e ired\ floating l onciuent\ 'tnmg out along the proved of much grc.tter v.tllll.'. togt·thl r \\.tth trilo-
rquator. 1 he ·,ubpol.tr' <. )rdt\\'KI.lll f.llln.l' of we~r bite~ they .ue the pnmal) 'tr.ttigr.tplucally usd\.u
em South Amcnc.1, North Afnca and Europe (we\t fossils in the \h.1llo\\ -\\,Iter t'Jcies of tht: Ordov1cian.
Wlld\\,ll\3) C,lll be dlsrlllglll\llt·d from tht'SI.' of the Brachiopod J\~l·mblagt'' .m:. of cour.e, vel) much
-.>r~ tropic.tl ·e.tsr Gondw.uu', n.11ndy Ausrralia controlled by t:KtC\ and on a global -.call.'. by realnu.
111 ~outlwasr A\1.1 . Spt'll.th7t>d deep-\v.Her fnmao;, and provmcc,, but corrclatwm at the \tage level
~h oh thl' !.Her (. )rd0\'11."1.111 Ftliltllllrlltl fauna, of bast·d upon concurrent braduopud range~ wtthjn
1111al.l thm-,hcllcd hr;whioptlth coloni:red outer-shelf province~ cm be Vt'l) good Tin" ~traugraphy and
uglll \ttt '· stn~cmre m rhe C.1r.1doc rock~ of tht· G1rvan district
13) the l.ltt·r <.>rdlWJn.nt br.lt h10pod f.mnJS of Ayf\hirc, Scotl,md, h.1vc bcl.'ll correlated by
bctJme \l>mcwlur I"' prnvmr 1.tlt1ed due to the refert'nct• to the st.llld,trd t ominuum brachiopod/
do,er .1pproach of the t'OiltiiH'Ilts, rhou!!h there was trilobite-bearing scries uf Virginia, Alabama and
I mlll11gh endcm1cll) 111 place~.

At the end of tlw OrdoviCI.In .111 1ce ~beet. which
~~·cm~ ro have t·xt~rl'd 111 polJr n:gions (west
Tennc~~cc, both .m:a~ being part of tht> Scoto-
AppJbchian proviiKc of the Antt•ric,tn realm.
So ~ucn·s~ful h.t\ tillS str.ttigraphic.tl work been
lr\mdw.1na) from rlw t',lrly <. '.1radoc, expanded sig- that ,l ca\e can bt: pre~cntcd fi.>r thing 'hybrid' )trat1-
uicantly. As~OCl.ltlll With tlm IS tb(,' llillh1111ia fauna graphical rabies in .1re.1s Slll.h ,1, the l3ntJSh Isles,
:: discus,ed 111 lull h) Rong and H.trper, 1988). where the Incur.~ on of· Amen can· fonm at a p::trtic-
Typt~:ally thts rOml\1\, ,1, 111 l~urnpe. of onJv ,1 very ular period m time .1llows the rccogn1tiou of zones
rew gcm·r:1, thou~h rhc1 t' " .1 \tccp bmgeograplHc.ll and swgcs ongmaUy defined 111 the Umted Statt'~.
) £fJdient toward~ ( hm.1. \\ hll h W,l\ more tropical, There i' now .1 good torrclauon between the
:i 1nd the dtVCf\if) of I fm111111r 1 f.mn.1~ j, .1ppredably qage~ of the Ordovll.IJil .me.! S1lun.ln 'hcUy tJnes
jthl.'r rh.~ late-( )rdovil'l.\11 <•timanti.m) glaciation .1nd conturrt'nt grapwlite zone,, mdecd there arc
Id much h.mn t<) hr:lt hmpolk ,,, to the other fJu- !>even Caradoc stage' Ill Engl.md and Wale\ for only
lS. and cnllcl uvdy the hmt.l\ rccovaed slowh. m o graprolite zone,, mdilJtivt' of .1 tine rdinemcnt
194 Brachiopods
of corrdauon. J'h~· Am~·rH:.m shelly facie~ st.1ge~ :1re B.1kcr, I' .G. ( I'J70) fhc 1-o'Towth and \hell nucrosm~ecurc Cum.•
less findy drd\\ n but 'till good . However, such cor- of rhe thendcKC,\11 hmch1opnd ,\./omelllll" ~wrlli04 ch~m
rcbtHlm an: ~ull mor~· tinnJy ba~c:-d 111 some parrs of from the Mllkllt• JurJ\\il of l ngl.md. Pa/,wout,•l•'g)' ll rmup
7h-9iJ. I.)}- I
the \\ orld than m othe~ .
Baker. P (., ( I<JH3) fhc dinunuuvc thcodcJJme hnduud Euug, (
In th~: Co~rbumtl-rom. brad110pods h:1ve been of
Erwllt~~flt••""'a /'):I!""Jra (Monr..-) from th..- .\1ldJir lml.'tl
~omc ~tr.ltl~raplu~·.ll v:~lue when combined \\'lth
Jur.t"ic ofEnghnd. P.r/,,,,,,,,,,,~1!}' 26. M•.\--70. colog
~oral' 111 the 7nnt•s crert~:d for the Lower Baker, I' C ( I'J'JO) £he ciJ,,tfilJUuu, orib.-in. and pin Enug. <
C....1rbomterom m Engl.md h\ V aughan ( 1905) and logeny ol .lrticul.l!t' hr.Khmpo<h. f'al.rftllllc•I<'K) 3J H..dlt
G.ln.\ood (191J) . I hough 'till valid, these zom·, are 175-Y~ de-~li
nO\\ suppkme11t~·d by 'tr.Jti~'TJphical work usin~ Ha,~crt. M.C.. (1')~-n L1fi: 'trategle' ot Stlun.u1 b!l- l't'rgu\o
llll<rOil1\\ll\. wh1ch gl\"l'S ,\ mor~· pr~·cise correlation. ..:htOpods, m llw.·r.•l·•~y tl/ Stlun.m Orl!aiii.IIIIS (cd, MG C.arht
Ua,\crt ..md J D. L1wson). 'lpecaJJ Paper' 111 PaiJtO.n- /'tJ/1'(11
tolo~:.ry No. 32, At.tdenuc Prc\\, London. pp. 237-fiJ Mudy
Bhght, F.G ..111d BhKill, D.l·. (1990) Flyiug ~pirilenJ: '-'·'"' 00
,mnc thought~ on rhc lift- ~rylc nf ,, Devuruan spurl~nJ 'lurl i
brJChwpod. Jla/tlfi~!/1'1~1/''IJIII y. flttlll<'tlrlillwtcllcJ.t!Y, P.rldfll·
Books, tre atises and symposia I'COic~f!Y 8 1, 127-39.
( ''''"'
11\lllg
Boucor, A.J. (I 971) Prauical taxonomy, zoogt"ography. \• r.mt, I
Houcnc, A J. (1 117'i) Hl•c•illlh'll ami i;:l.'littctivn Rate Cottlwls. palt·occolo!-.'Y· p.llcng<'llgraphy. .1nd ~trlhbrr.tphv r,,, lt tl' h t
fht'Vlt:r, Am~tt:1.!.m1. (I >evel<1ps concept of bcntlur S1lunan Devuui<m br,tduopl'd' Procct'dilr.l!s <?{lire,'\ durinj
,h\l:mbl.lge') imrrirolll flalrolll<'h~~i(tl/ C<lllt'CIIIitm, Cb.ic.t!!;U. 1'16~ (r Gnnt. l•
Bmnmn, C.II.C md Cu1Ty. G.B (1979) British pp. Cj(,{,-(,] I. (L~rl~ qudy; base for Boucot, l '1751 thc p
l3r.uh111Jic'd'. lm11<.',111 \ntKtv Synup~e., of the Driu.b UretsJ...y. P W (1%4) Cenrr.al Appalaclllln Utt l'r~ft·cm
F.11111.1 N<l 17. 1\t ldt·mu l'rc,~. London (All Brimh OrdovJca.m rvnummm~·\. Bullt•lm tl/' tltc Gr<•t'rg/ld <:ram, ll
'>Pl'l ic'i) ,\t>Clt'ly t!/ Im men 80 . I 93 212. (Commumt\ ~tructurr J\1111111
D.l\ ,d,on. I . (I x:; I) I Mtllllii!I•IJ•h ,,, lilt• Brill.ill FMsil Br<Jdt- Brctt, C. Boucot, AJ .m.! Jone,, R. (l'l'IJ) Absd 1'.1/('c.l/1
"'1","'' · l'.lllet•ntll!--'1'.1plu~.ll <.,oncty. London (Standard depth r.m~c' of Stlun.ln bcnthtc 1\\emblage\ Lt:l 14H-
,urcrbly alllhtr.ltt·d u:fi.·r.·nc ~ 111 II\<' volumes. raxonomy) 26,25-tll Gr~m. ll
McKannon. 0, lt·t·. fl f: ••111J C.unpbdl.J.D. (eds) ( 194 I I Burron. C.J and Curry, (, H. (I 'JH5) l'clagic buduo nmns
Hr, -/u•'I'•'.Jr l7mmi!IJ /imt". Ballcm.1. Rotu:n.Lun (scvcrJ.! from the Upper Dcvom.tn of E.N Cornwall. 1'roJtrdl 1'.!/c-tlll.
up-to-d.ltt' p.apt'!'>) t~( tlrr l'sslrn Stl(H'IY 6. 141-5. lophor
Moort', R.n (c<l ) ( l4to5) Trf Ill it ' 11 ltwertehr,J/1' Cherm, l. (197')) ' ) he cnVIronment,ll \igtltficmc~ l bU.uu,
Palt.Jtl/tl/,,1!)'. Part It. Br.atluvl'•'"·'• 2 vols Geolog~cal Lmgu/t1 m rhe 1 udlow <.,crit" of !he Marl~t
SocH:ry ol Am<'nl'.l .111<1 UmvcNty of Kama!> Pn:~s. Welsh OorderlJnd .wd W.1le\ /..er/will 12. Vi- ti5~9
L1wrenn:. K.111 (K.u:,ll r. lt.l (ed.) ( 1477) Vol. I of (Dccp-watcr lin~-o'\ilitb) l l:llllllli)J
th<. rcvN:J \ l r'Km ts 1111\\ pubh\hed) Cwg, G. Y ( llJ'i 1) A cornp.lr,mve 'itudv of the ccol~ ttll('r<lr
Ruthvick, M J <., ( l 'J70) /.11'11~~ arttl Foml Bmchwpods. ami p.tl.leoel.olnh') of Llt(~lll.t. i'PdiiS<Ictiotl< •!I !lOll 1
Hutt himon, l nndtm. (lnvalu.1hle m:.ltmcnr. Wl[h a s,,,,cry
Edi11bu~~~~ Gn'/tJ.I!Imi 15. 110-:20. (Fosstl Uttf jt•unwl
funuiotul ln.1~) m life po$ltlOn) 1 loln11.•r,
Cocks. l.J{..M. ,lnd r ortcy. R.A. (1'.)90) BIOgcogrJpln· 01.11latt
Ordovician and Silnn.1n f.utn.l'. 111 Palae~: Swede
Individual papers and other reference s P,1/acot:co.~rnphy and Uh~l/i'O,I!tllplr}' (cds W.S. McKerro; I a. B.~rbl'
and C.R. Scnte,c), (;eolog-~r.l l Sncu:ry Memou No IJl()V('IJ
Ager D. V. (1%5) 1 he .1d.1plmonufMe~ozoic brachiopod~ 12. pp. 47- 104 <'rlC'nta

tu d1Hcrcnt <'11\'lrtHllll<:nts. fl.li<lt'<~Co~mphy. Pa/,Jcodima- Cork~. LR.M .111d M~.KciTOW, S (I YH4) R.ev1ew ,,f I .>~WSOI I,
t.•/•!1/Y· Polocclin'l''.iiY l, l·t'>-72. .-ornmoncr anml.ll'i 111 Ln\H'r ~dttndn nunn~ b~ntht / 1,1/,u•vt
A~er, D.V (l'lt.7) l3t.Khi~o>pl•d p;dacoccology. E,;rth <.OlllmUnlfle~ PrJ/,u•cmltl/cl~Y 27, hh.\--70. l'Od-d<
'iOt'll<t' f{fi'IL'II'I J. 157- 7'1 Curry. G.B. (19H2) Ftoltlf.'Y .tnd popul.it10n structurt todd~.·y.
-\gcr J) md lttt:t-."• E.A. (1%4) 1 he mtcmal amtomy. th~.· lle~.cnt braduopud fnrbroltltlillol from Scot.. raatt•J 1
~-o•wwth Jml ,t,~ltllllt'try ol .1 l>t>\'om.m \pm fend. )c•unwl Hl/olt'tl/llcl/O,t:}' 25, 227-46. f",Jitlt't>/1
4 Hllt'<''"''h'X) 33. 749-(ltl (Cn.>wJmg results in CulT), G (I 1lH'h) M tnolmnnt-."> an Recent br.tchn~ Mc.c:hce,
.1srmmetry) :tnd tht• ftlllCIIOll\ of l '.II:GI , U'lil<lltl 16. 11 'J-2" IJte tlr
S7-7h.
Bibliography 195
~·. G (I tJS3b) Pt·olob')' ot tht· Ren:nt dt"ep-warer rhyn- Muir-Wood, 11. ( I'J55) .-! Histtll''}' ·~I rlu: Clcm!fic.rfttlll <!(rite
,hwellid br.t< h10pod CryJ•fc'J"''·' trom the Rock.ill Phylrm1 Br<ulrwptldcr Urimh Museu111 of Natural
trough fl.l/iJnl:t!<·,~mp/1)'. fl.J(,~e,l(fllll<llc>h~)'· Palan•t'U•Ic':'l)' 44, H1story, pp. l-12·t. (fiNt .mt·mpt tn ercrr a modem
9HII~. (l echering pnhck·) classtficauon)
t1111g. CC. (I'J~l) lmplit.ttlnu' d<· donn~ rr:ct"nt~ sur les Pernval, E. ( 1944) A umtnbuuon to the ltti:-hmory of
lll'"'tdt") acrudk... d.tu' lt... lntr:rprr:ranom p.llt"oc!- rhe brach10pod 1crrlrr.urf{,, tii(IJIIJ}'''"'' !:>owcrby.
~·111<''· L:thrr,r 14 , 151 h (ll.lhltal:\ ot'hngulrd~) Trmr.~artrtltt~ l!l tlrt· Rt~ycrl ~<lfll'f)'<!( \'t'U' h:a/,wd 74 . 1-24 .
:tg. C .. G.11l, J <. P.IJ nrd D md Pl.vw. J.C. ( 1978) Rachebocuf. P ( ll)CJO) le' br.ll h1opode' ChonetJce' dim
Rrae\tOm cntll]llt'' 'ur l'l't<11ugu: t"t la ,nci:marique Je, .IS'-embLtge~ be111hi4llt'' \llurrt'll' et dt·\·oniem.
d..>s hngult'\ Jttudk·s t't '''"'lt·~ Ct'clbit•S 11. 573 o09. Pn/aco.l!c~r.1pl1y, f>Jic1t ,,dmlcll<•lc!~!)', £>.1/ctt'<'U<li.~!!Y 81.
MglhllO, L. (I')():!) r hl' p.d.IC:lll'COlllb'Y of .1 Lower 141-61.
t:arbonrtl-rnm m.mllt' rr.ln,~ri."''\IOn. _/ounw/ '!f Richa~dson, J .R. and W.mon. J.E. (1975) Fom1 and
P,rkc!lllc•l•~li) 36, IlNO- I I 0 I (CimK p.U.tt:OC(I)lO!;.'lC.ll function m J Recent free-hvmg br.•ch1opod.
\tudy) f>al~ol11olo}!y 1. 37Y-H7. (U nu~ual l1k h,1hit')
l;ffi\OOd, j b. ( 191 J) I he I OWt'l ( :.lrbo111ti.•rou' SUCCe\- Rong, J.- Y ..111d l l.uper. I) AT ( I 9HH) A gk1b.d ,ynthe-
,j()IJ tn rhe north\W\t ''' [ul'\l.mtl Qll•lflt•dy jtHmra/ of tlw sis of the latest Ortloviuan Hlrnanuan brach10pod
C.v/o~llcl/ ."'c•ot'fy tl/ L<•11dc•11 61:1, 441J-S'Jn. (Zonation fuunas. T'ratHtUticiiii 4 tlw N.cly<rl Stldt•ty 4 Udmlu11gh 79,
Ulmg coral .111d br.Khmpnd t:nm.1) 383-402.
••nlll. R.C. ( lW16.1) A Pr:mu.m pmductnid brachinpod: Rudwick, MJ.S. ( 195Y) The h'TOWth and form of brachiu-
litehistory Sm·lltt' 152, I•W 2. (C hJu~e m life hab1ts pod shdls. Gl'olc!~im/ Jl.la.l!•1.~illt' 96, 1-.24. (Growth vector.;
during on11 l~t·ny) in recti-m~rginat~ and phc.lted 'hdls)
r.tnt, R.E. ( Jll(,(,h) !'-.pull' m.lltf.t>mr:m .md lite habits of Rudwick, M .J.S. ( I<)(, I) Tlw fl•edmg mech,uusm of tbc::
tb~ produttoid bra<hlllpod rJ ,,,N('II<'"'"dla. Jmm111l <!( Pcm1ian brachtOp\)J f>rc•ridtihoJi·llicr. f>alc1Clllllllft>f!Y 3,
PJ!umrc'/••1!) 40 I 061 'J (A, .1bon~) 45ll-71. (Worktng model shows oper.ltlon ol Aappmg
run, R.E (I '171) UrJc:h10pods 111 the Penm.m reef envi- valves)
'11llmt'nt of we't I c:xa,. />,-,," .-drill!.• •!f the "••rth Arm:rica11 Rudwick, MJ.S. (1964) Thc: lulllllon of Lig-ug deflec-
c.,,.,"'"'"
I'J/conr,•/,~~i.. r/ Clut.l~o 1464 (J). pp. tions in brach.iopod~ . Pnlmcllltolc'i!)' 7, 135-71. (Z1g:zags
1-14-t-81 . (Broheml\ With unu,tl.ll spt:<'r.llrzcd f.uuus) interpreted .h prorccnve)
.unt. R E. (I '172) ·r ht• lnphophnre .111d feeding mecha- Rud\\;ck, l\.l.j S ( !965a) AdJpuve homc:umorphy in the
!1Wll5 of thc Pmc.lut 11<1111.1 (Dnduupuda). _lounwl '!f brachiopod' Trttmtmd/,1 Bmner .md Cltrin•tlrym
P.dt.Jm,>/c\') 46, 213-4t'l. (hN Jt"'t:npoon of producrid Rollicr f>clllicllllolc>i!i~drr .Znr.q-/mJi 39. 13+-4h. (An
lophophor< .uu.llunt'O< l1Uitnrt·rprt·t1nun) extreme case of homcomorphy mterprcted fimnion-
fWJ.un, l\ (I W•2) Br.Kiuopod hti.· .mc:mhlagl'l> trom the .illy)
~lul;tone Rod: bt•J ot Lt'IH'\tt:l'\lurc:. f>nf,l<wrt,,f,~~y 4 , Rudwick. MJ.S. (19o5b) St•mol)' ~pmc~ m tl1l JurJ..s.,Jc
6i3-9. (Suue~\1\l' uulu.tl bromh) brachtopod ."lwrtlwrlum. Hrl.rl.'cllltoltli(y 8. 604-17. (!'o.pme'
ilmnnond. I .~ ( I'Jln) L· xpt·nment,ll mrdtes of ~.llmity a~ early w.1mmg \ell\01'\)
toltrJ.me, burrow1ng hdl.l\ l <l llt .utJ petit de regeuera- Rudwick. M.J.S. and Cow.:n, R. ( 1%R) The functionAl
cion in Ltl(~u/o •lllolllltcr (UrJthtopod.l lnarttcul.u:.a). morp hology of smm· .lherrJnt srrophomenid bra-
}<•ttma/ <!f fJ,lft'oll/tlft~I(Y 57, I ' I I I f1. chiopods h-orn the Pem11.111 of Stuly Rullrri11 SNict<1
lnlrner. L.E. (I'IH9) Midt!lt· 01dovtd.m pho~ph.1ric inar- Pafcollwlo,~rctt lta{i.wl' 6, 11 3 76. (R iththofelllids .md
rit1llate brachiupod' ti·urn V:istn~Otl.md .md Dalama. other\ replace duary pump111g by valvt· ntovcmcttr to
Sweden. F,'.~si/s ami Srw,, 26. I 172. gcneratt' currents)
J BuberJ, M ( I<.'J77) UrKhiopnd orientJtton to water Scotese, C.R. and M cKerrow. W .S. ( 1490) Revrsed world
rnovt>rnent I "[ heory. l.lbor.ll<>l) lwh,wiuur <tnd field maps and in trodu ction, in J>altrnr;:,rit Pa/aco,~c<~~rnpliy
oncntation Palt•ohio/cll!}' 3, 270-l-!7 ami Bio.~~'tW1'11flh}' (cd~ W.S. McKerrow Jnd C.R
1Jw1on, J.D ( 197'i) I udlm~ benthoniC .1\scmbl.lge~. Scott"se) Geolob'lc~l Sncu:ry Mt"lll<>m No. 12. pp. 1-21.
P.rla~tlltrlll~!!)' 18 . 'iO~ ')::; (I )1\tllll tmn ol four brachio- Sheil.l>, K.A.C. ( 1968) 1--.'<,,friJIItldrulw wwrrttl n. sp. fTom
potl-donun.ltt d .t\\l'lllbl.t~t'' and thetr ullltrob) the Scocri~h V1~C.lll, ,md a pt1\\tbk mcthamcal advan-
f W.Ucv, M.(;. (1 1>H3) '\ rt:\'1<:\\ tlf hr.IChJOpod-dorm- tage of it~ R:~ngt' '\CnJct\lrc. I r.mmclit'"' <!I tilt' Rl')'•tl
lllrcd palaeocommumtll~ from rhl.' cvpe OrdovJCtan. Sanety '!I Etliullllr~h 67. 477-507. (Uw ot'liow t.1ble in
/'.dott<''"''h'~)' 26, Ill l'i (Ex,dk·nt trt'Jtnumt) dc::temuning partidc 'cttlrn~ on fbngc:)
!.-Ghee. ~.R. ( 19tiO) 'iltdl fimn Ill d11.' btt'onvex arncu- Tluyer. C. W (I 975) 'mength of pedKk· .1[(.1( hmcnt in
hre Brad1111pod.1 J gt:<lllll'lnt· .m.tly,rs HtlftlbicliO.I!l' 6 , arricul.ue brach1opod,. ccnlogtt' .md p~leoecologic
57-7£., stgmficanc:c Pt~ltobit,fc•l!)' 1. .\KK-99.
196 Brachiopods
Th.t\er, C. W . .md Stl"l·k-l't·trovic, H.M. (1975) stacimcs; development ,llld dunge\ in flllnal provmcc
Burrowm~ of the hnguhd hr.Jcluopod Clt>l!ldia pyra- Williams, A ( 1976) Pl.m· tct tomts and biofatics evolu-
mld.t: it\ t'l'nlogu: .md p.lleoel'olo~-,ric sigruficanc.:. tion as ranors m OrdOVICI:ln correlation. Ill 7lrt
Lt·til<~ia 8 . .20IJ-21 . (llu\\ thl' fll'dldt• is used) Ord.wicinll Sy.(t£'111 (t·d M .G lhm·rt). UmvcNl)' d
VJugh.m. A . (I '105) I'he p.li.JennrnlngJc.11 sequence m the Wale-~ Press, CJrthfT. pp 29- (,6. (F.tunJl pro\·incc:~311d
( ubomti:rnus hmt·stone nt tht• Bri,rol area. Quarterly srrangraph1cal probktm r.li,ed by plate closure md bu-
)mm~<tl •!I tilt• Crolt•l!•r,,/ s,.,,ft)' 4 Loud,•n 61, 181-307. nal nu..,rrauon)
(CIJ"k ,Wd\ us111~ tOr.JIIbr.u.:hiopod zouanon) Wtlhams, A. and Wnght. A.D. (1961 ) The ongm oftbr
We,r, R R (If (•) Cnmp.m,nn of '\C\'en lmgulid com- loop in art1culatt' br.ad11opod,. f>ul.lt'Otlt.>loJ.!lY 4. t~•J-76
munml'\, 111 .'>rmam, tmd Cl..zn!Jit·,uj,,, c>f Palart>cmn- Williams, A , Carl~on, SJ. Bnmmn. H.C. a al. ( ; •I•Jfl) A
mwlltln (ed R W Snm .md R.R . Wt:st), Dowdcn, supra-ordmal cl~ssltit".lt1011 of the Ur.JChiopod.l
Hmdumon .md Rn". "itroud,burg, Penn., pp. Phi/Mop/11(11/ Tr,mstlaitm~ •if tilL' Roy.!l SoXtrty '!f L.>t~Jon R
171 - '12 (I 111!-,'l.lhds wcTe .tti.lprcd to ditTert:nt environ- 351, 11 71-93. Thl·
lllt:lm) Zicglcr, A. M. ( 1965) !:>alunm m. m ne commumoes llld tcbra
Willt.~~us. A (I 'lhH) A history of skclet.ll secrcrion in bra- their environmental Slgluficllllt'. Ntltllr<' 207, 2711-2
livint
duopoth. Lt'tfl,llll 1. 2hH-H7. (l::xcdlem general review) (Asscmbi.Jge disttlbutloa, w1th m.tps)
Wilhams. A ( 11J7)) DlstrlbUtiOil or brach!Opod a.ssem-
feren
Zicglcr, A.M., Cocks, L.R.M. and l3ambJCh. IUI
blagt'S an rdmou lo Ordov1u.111 p,llat'ogcobrraphy, in ( 1968) The composition ,md stmcrure of I 01wr
mduo
Or~mmms ,,,d C>11/i11o'tll.> 'f7tnlll.~h Time (ed. N.F. Sdurian marine U111lll1LIIlitic\. LI'tl111in l. 1- 27. (Cia111c, copho
llught•s), ">pt'CI.Il P.tpers 111 (l;JI.Ieontology No. 12, well-illustrated work) shells
Ac.ldt'tllll' Prt·~s. London, pp. 241 - 119 (M.:thodology, tors (
romp
ccpha
tlcfi~l·
amnH
m ann
rropo•
~rroup
nukc
JI1fcrt
unlccc
StnJC(I
much
lt IS p
L•rganj
rhettc;;
ofmo
Clcular
Wh
he OUI
~lllCC I
fhl' CO
them·a
the eh
mod er
(rig. 8
many ·
dunng
Molluscs
a.
B
The Mollu,t:,l .m: l.llll.' l.)f thl.· most dlvcDc ofall inver- 1\fcopifiua IS a member ur the: mollusran Class
~rate ph\ I.• anJ mdude a whok rangl.' of animals, Monoplacophora, wlud1 h,,, toss1l represetHativcs
bvmg ancll"l1~~il, whu:h .1t llr;t 'lt.~ht \et>m to be so dif- known from the Cambn.111 to the Devom.m. The
lfrcnr as to be unrebtl·d. The unportant groups discovery of tllis 'hvmg t\mil' ha~ been of the great-
K
lllcludc the cunnus pbtl·d chitons (polypla- est mterc:st. c:spen.11ly Slltel.', contr,\1)' to l'Xpect.l-
lC,
:ophorans). the slu~ .md ,n,uls (~.1stropods). tooth tions. 1t shows tr.1Cl.'S of 'egmt·nt.ltion.
~ell' (\c~phopods), hiv.1lws, their plmible progeni- T he structure of the ,uchunoUusc ts l[Ultc simple.
'Cll'l (the ext1nct rmtroconchs) .md tin,1Uy the most It has a cap-like s hell setrdl'd by a layer of tissue
omplcx .md h1Ahly org.umed uf ,11l molluscs. the known as the tnantle. Uclow tlus lS tiK· body wtth a
tphJlopods, whtch 111dudl· thl• modern squid~. cut- mouth at one cnd and .lllm .lt the other, the latter
tldilh, Ol ropmc-. .1nd pl'.lrh naunlu'>, and the fo"-~il d1~chargmg mtt) .1 po,tcnor '>pJce, the- n1antle cav-
.1111monol(h and bekmllltl''> Mollmc-. arc nuinly a ity, m wh1ch al'o re~1de the gtlk
mrinl' phylum ..md onl) .1 tc\\ hivJlves and gas- In other rnollu~c' tlm \tntcture ts mod1fie-d m dif-
~pod, h.lvt· hel.·n \IIC<.' l'''tul in trl·,h water. Only one ferent ways. Most mollusc' h.Wl' .m extemJl o;hell of
~p of ~l\tropod,, tht• pnlmonates, w.1s ever able to CJioum carbonate. though tlm has been lost m var-
ke d1c rr.m,ition trmn w.• to Llnd . The divel"'\11)' of IOus gastropod'>, partrl ul.1rly tht• m.mne nudtbranchs
.fui~rrm kinds t)f molhN·, i' extrt•me. yet they are all and rhe terrc~tnal .,Ju~ In squid.-. and curtletish, as
llllltcd h) .1 tommun ~rround pi.Ul ••md the -;m1e ba_sic well ao; m the l.'Xtmt t bdnnnite.-., the o;hcll h.l.<>
JIIU(turc' .trc to hl· t(mnd Ill .tll molluscs. however become moditicd :l\ :m llltt·mal \kekton. and in the
ch they h~n· hL'nmll' <hfli:rt•ntl,ltl:d. octopuses 1t ha\ been cntJrdy lmt Where the o;hell1s
present ir 1s alw.ty'> \l't rt'tl·d by thl· m.mtle, "hi eh
directly underlie' rht· \hdl. Throughout growth ca.l-
cium carbon:lte 1' added to the edge of the shell
!Tom the m.mtle. In mmt rnollt1scs th1s mantle
lt 11 probably simpkst to mmider the fimdJmental comim. of only a few cl'll thickneS\l'S, but in the
III),'Jilization ot mollml~ w1th rdi:rcnct· to a hypo- cephalopod-. it is t,'TL'atly thich-ned .md \Upplied
t'l~ucal archimollu~l m wl1ll·h Jll the basic feature~ with powerfi.ll nntsck... Thl· lll.llltle tavity j, a con-
,,f mollu,tan strurturl' ,lrl' pn:sl·nt, though not par- stant feature of .lll mol l u~cs and (in different mollus-
4Uiarly specialized 111 any one dirl·ction (Fig. 8. J). can classe..) has been put to variou' U\l'S. T l1l' gill,,
\Vhik thi-. w.1y of prescnt1ng ~uch concept~ may w hme primary fimctu)n " n·,ptr.ttory exch.mgc, arc
:c out of fashion at p1 e~cnt, I rontmuc to use it present in ,111 molluscs except the \;md gastropods
lt ffiO\t student~ find 1t hdpful m undcrstmding (pulmonares), ~here thL· inlem.ll \urt:tce of the
umplcxlllL'~ of mollusc.1n anatomy. Tlus hypo- mantle is highJy v:lsc;td~r and i\ modif1ed .t\ a lung.
Jell mollu~l..lll lllCL'stor ha\ dusc sinularjtics to In the bivalve\ the 1-rills arc not only re,piratory but
dmom and an c\'en timer resembbnce to the also adapted for thl' ~.Hhl'rlll~ of ~uspended food
~cm lunpct-hkc pnrnmve mollmc .\'copifi11a panicle~ from the water. Tht· m.mtlc l.tvJty ha.' both
ne 8.2). whu:h \\,1\ unkn0\\11 until }t)53 when an inlet and ,m outlet to thl wa through which
)' ~pecun~:ns wcrt· dn:dgcd from the deep sea waste matenal !Tom the anu' dl\thargc,, a' \\'dl a~
tmng the Oanish 'C.tlathea' Expedition. respired water. Tlm oudet ha.' alo;o undergone a
198 Molluscs
Po lyplaeophora
(a)
Figure
$howin~
dorsal~
lion by
M onoplacophora
1\IOllOI
\ shell .111 slo,
7 gonad l l lliSCU
"'---
r:ltlula
a belt
\\ h1ch
cngcr,.
hypothetical archomollusc have a
J which
are mo
deposit
normall
ccphalo
hunters
mg. but
Cephalopoda (nautiloid) erfi1l j.r
cil\e or
r•)\\ er l
t~.:nr.tnd,
Gastropod a
rim hab1
l ·cdtugl
Cephal opod a (squid) •ntr.tnor
eran~ rh
Figure 8. 1 Morphology ond relationships of molluscan classes with reference too 'hypothetical arch imollusc'. groups.
dtffcrennal modification, mmr markedly in the and phy~10logy. have most features in common
ccplulopod\ where w.ucr, taken uHo the mantle Such then is the fund.mK'ntJI plan of moD
Cl\ tty .mtl Sljllirted out as a Jet through a funnel. is organization. but "'hy ~hould mollmcs An Ol
tht: pnm.IIJ mc,ms of proplllsmn. diverged from lt 1n so m:Jn\ d1rections? lr ~l'ntll \\ IUllll PI
Abo\'l' tlw mantle c l\'H) Ill .tll mollmcs IS the vis- much of the evolutionary dttTcrcntiation of m
ce ral m ass, which cont.um the gut, digestive glands is bound up wtth thctr mode of nutrition, .and ClASS 1. I
and ktdncy' .md the ncrvom, circulatory and mm- inrcrcsnng to sec how thetr <;tmctural diffrrt'n l'l1p ol p
tke ~he%
cul.tr ~ystl'll1s These too, in srmcturc, arrangement is closely connected with how tht>\" t~
Classi~cation 199
-foot
(c)
~e 8.2 Neopilina !Monoplocophoro) la) ventral view,

lhowing central foot and serially arranged gills I'- 2 approx.): !b)
doool view !x 1); (c) lateral view (x 1). [(a) based on on illustro-
Cllby l.emche in Treatise on Invertebrate Paleontology, Port 1.) Figure 8.3 Chiton !Polyplocophoro): la) dorsal view; (b) ven-
tral view (x 21.
nonopLlcophoram, dlllom and mu't ga,tropod~ arc g.tstropods Jnd di~unKUt\h.tbl!! from tb~m on the basis of
Uslow-movmg nml!Lt\l' wh1rh rn:cp .tlong on a muscle scars) Thc:y h:tvc: JXtirl·d. wnally repeated mmdcs,
'!lu>cular 'lt•ot'. The> arc .111 providnl with a radula, gill~. nephridiJ (.:xuc:tory org~ns) .l\ wdl .l, other uttem:U
bdt of senally .lrrangcd tt•cth wtthin the mouth, organs. The fi)ot J\ cir<uLn .llld u:rur,tl .md nnged bv the
mJnde cavtty in whu:h he 1h~ !{ill\ Mouopl.tcophur.tm uc
hich firs them to be het btvort''• t'Jrmvorcs or \CJV-
th(· onlv molluscJn d.h~ wttl1 J rruc mtem.1l '~~n~nw
:nger.;. On.ly the ga,trupod~. of tht·,c thn-c cla,ses, non, ~ugge,ung ,1 zoologil.tl rt"l.ttmmhtp wtth .1 \t'g;mc•med
I'<' a wtU-dcvdupcd ht·ad \\'ith scmc organs anet•<;tor (but ~cc C'haprcor J). Fo~~th \Udl J~ Pllm.1 and
I
clnrh enable them Jcuwl~ tu hunt. The btva.lves Tryblidmm .ue !mown only tT<Illl rhe P.1lacozotc. •\'n>J'rliii<I,
rr mosdy suspcmton lct•Jn.., and lcs\ commonly I 'ema 1nd a tcw mhcr g<·na., or cur rodav mamly m the d~<"J'
~m feeders. The) h.t\'c no lw.td, no jaws and sea
CLASS 2. POlYPLACOPHORA (AM PI II NEURA) U. C.unb.-
muUy on.ly limtteJ clp.tnty tor movcmcm. The
RL·c : ng. 1-i.J):Mannc mouu,C1, IJJnn~ .• bll.ltcr.IIJV wmmet-
~hllopod,, on the t•thcr lund, an: fa~t-moving
ric::U >bell \~ tth ~cn·n or ct~ht lJk trcou' p!Jte,, hue othcrwtsc
tt'l'l; here rhc radula j, mt•d main.lv in S\\ a.l.low- re,embhn!t the an:h~tH•al mollu,on phn m havmg an
' but tht• curttng up o! fm)d 1' done by the pow- amenor mouth \\ tth rJJul,t. ,1 ptl\h.:nor mantle canty wuh
i!1UI jJW':i. The he.1d, '' tth tt' htghly developed .mu' and gills. and a \'cntrJJ t'l)()t. Polypi.Kophnram .rrc r.uc
.erne organs and wcll-m~atlltt·d br.tin, the animal's foo;sil, but ou:ur \Can~n·J throughout tlu· l'lwncrozoK, e.g.
wer of movcmcnr. tt\ huoy.Hlc > dl..'vitc~ and the Ch,tc>ll.
CLASS 3. SCAPHOPODA (Ord RcL , Ft!l R.4a): M.mm: mul-
ntJCitlar apparatu' .111 l'\'olvcd m anord;mce with
lmn wuh sm,tll tape•ing <Urvlllg 'hclh opl'll ,tl both emk
b hJbtt oi cJtC'htn~ ,\lttVl' prey. lt is the mode of
The .1mcnot wider eud wrth rhe mouth i' pcnn.111cmtly
· ~mg th3t h.1s been aL tltc ront of tuoUusc.m dtfier- embedded m scdunc:nt, 1hc: .Hi rm,,), ft·ed 011 ~tnall orgJut~nts
ttJhon; thts should be rclltl'lllbt'rt·d wht·n constd- us1n.g 'pecially arl.lpted tt•ut.ldcs. The anu~ ., .tt the uppa cud
mg the srrucrur:~l pl.m of the VJnom molluscan Jnd the gills ~re much redurccl Rct:c:nt 'Jll'l'll'' arc mort'
;roups. abundant than fos.,JI one~ ,1nd ocrur domw.1ntly nn the: con-
tmemaJ ,]ope or 'helf Tht• c>rll' dt·t..ulnl 'tud\ \O tar mJde
of the ecolofzy ot ,1 fm~tl '<.tph<'P"d, the Ordm'KiJJl
Pf,wro.f!lypla, sugge,rs ~uml.1r luhth to modern lonns. e.g.
Drllfalium.
5(31\
CLASS 4. BIVAlVtA (LAMELlll.W..ANCJ I!A or I'ELECY-
\J\ c An om!tm· cJ.,,~thonon nt" tht' n1.1in divistons PODA) (Cam -Rec) Btvalvcd mollu'<' or 'dam,' wtth no
th!lt dun Phylum Molht~t·.t 1~ ·" f(,IJow,. defimte head. bur havmg the \Uft p.1n' o:ncln-.cd bcrwc:en
!u,~·s p;ured but unequil.ltcral c:.tk trl·nu' ,hell~ united bv J dol""al
it )\ .JSS 1 MONOPLACOPHORA (C mtb.-Re<· : F•g. lC!) A hinge. which may be wothcd. I h~ vAlve' cJn bt: \hut by
oi pnmmn• m.mnc molllh(' wtth um~ .th-ed lunpu- 'rrong intem~J mmcul.11un•, and upc:ned b} the outward
ttion
ilirU, (though unrcl.lt<·d m lnnpef'o proper, wh1ch .Ut' pn•"ure of ,t ~pnn~•y h~.llllt'nt alun~ot the lungl' \\hen the
The
200 Molluscs
ClASS 6 GASTROPODA (C.un- Rl·c. ): Sn.1u' of all kmds - (k..\lllll
mJnnc,l.111d Jud ht·,hw.lll'r - \\luch u'uJII\' trl'CP ;iluu;!lolll Cambt
tl.tttcncd 'timr' A. !TU•' hc.td l\ pr<''<'nt \\lth t:\'l..., and tltbtt l>ut .tn
helen scmt: or~m ..md the .m.;<.' 11111\ .1lwd 'ht:U '' rmlt:J, pla.nispi.
anord
r ally or mor,· otren h e lic ally 1 he mtcnul or~m arc twiSted
operculum by a I HO" torsion \0 ti1Jt tht m.mtl,• C'J\Jt\ fJu......lllh!noolr hi\'J.!vc
Som. '~cond.mly .hcU-k"' W\.IU~" tlut luw lmt th<"ir tOnQ hack r
.1rl' k.no\vn. Ptaopotl, .tr<' m.mne g t-trnptkl,, atbpteJ scapho1
swtmnung. 111 \dlll h the." ti>nt '' prnlon~cd m to btt'ral \\1 sent ev
for lite in th.: ot<'.tn. Th.: t.apenng ··munl ~lu·ll\ nl rh("S(' 1-.lmk· Cambr
tolll< g.t~lmpud, .ar,• ~ntllt:llllll'' fi11111d m ii1"!ltom1
l'VtJenc
CLASS 7 CEPHALOPODA (U Cun.-Rt'c.): I he m
o~dvanced ut .tit mollu"·,, h.t\'111~ cxtcrn.u or mt<·nul dum·
rhe\c e.
bucd ,ht:Jis wtth thl· , h.ltllb~t' hukni by ,1 'tphundc JJI.l · md of
(a) ing htHWlnq The\ h.wt• .1 propcrlv dl'fam:d he.td \<rm .1bout r
eiJbor.uc ~cn~c nr~.Hl'..md move by .rct propuhwn ol \\lie \l't'lllS t•
from the m.lntk rav•ty. 'I he nwdlTII ;\i,mri/11), "lu'd' ~o~ orif,'111 c
ocwpuws :u~ here tncludc:d .•~ wdl .1s Jnunollttl'S, h.:len:· l•mg-dc
mtc:s and ~omt: d~<~mbl·rnl C;uubt i.111 mollu~n (Fi!!. R.5
Figure 8.4 (o) o Aattened hyolith, show•ng operculum and till reasc
exrmet rc!Jrivt:s of 1\·,mlrlrl.l.
paired helem (M. Cam.; x 4), (b) Dentalium, a scaphof>O<l, ~·subl bl
!)(
reconstructed in life position 3). [(a) redrawn from Butterfield whtch I
and Nicholas. Journal of Po eontalogy 70, 1996; (b) redrawn The Jbow clJ~~ification ~ comerv;lti vc and some
from Kershaw, Animo/ Diversity, 1983 ] -1!-''l"CCd c
authoN would aUow 11hlllY more cla~scs 111 the
OrJovic
Cambnan. Anordmg to Yol hdson ( 1979) , for <:Xlilll·
mu~ck' rciJ..\ I ht• ~IJ, art· l.lr~<' .md mnditicd for fiJr,·r fccJ- plc, an earl) mollmt.m radt.Jtton 111 C.unbmn nm
tlu: mol
mg, ,\lh~tlw nun de c3VItV 111.1y be connn tcd to the outer cJJSSCS <
gave rise to .1 number of ~cpar,Jtl.' ~hort-hwd cl~
<.'11\,rr.nmcnt b\ sip ho ns indudmg nuny kind~ of ''mpk t aJ·Hhapt·d ftmm (L"
ltllCJgL'S
ClASS 5. ROSTROCONCHIA (L. C.un.-Pcnn.: f1~ 8.16): g.l.~tropc
Hdcicmclla: hg. H.l6c). l'hc'c \Vl're subsequently dh-
Exnncr molltN ot bl\ .tln--hkt· .lppl'JCJIKl' but with one or tant fiw
more of thl• ,ht:Jl bnT' Hlntmuou' Ju"'' t),,. durs.ll margin placcd b\· more effective c.1rly rcprc,cnt.IO\N
.lttenttot
'o thJt J dof\.1! , umm•"urc •~ I.Kkm~ TIH: JUH.'nue shells are ·modem· grm'l'" 111 a \Ccond. btL CJmhnm-
the othc
Ulll\'alvcd and tmlnl early OrdovKun r.1di.1tion. The .lltcmati\C \1C\I
dl~l.-U'>\CI
septum
,.....""'~~·~A'.t,~~~'
.,~:"' .....
. ···.::··~ ... · \,;.
muscle :· ~i
band
R-::1-ll*-- - paired
gills
(a) Plectronoceras
tentacle
(b)
F1gure 8.6
Figure 8 5 Plectronoceros. on early cepholopod· (o) specimens from the late Combrion of Shantung China (x 3): (b) inn puter-simul
resloroflon of soft pori$ with shell partially removed to show man~e cavity with gills (Based on Yochelson et ol, 1973.) shope note
Shell morphology and growth 201
Runncbrar. llJ9n) 'uggt·,t~. however. th.lt the small

C'.ambrian mollu~t:' do not pcrt.1in to cxonct c!Jssc~.
but lre earlv rcprcscnLlttvcs of ext.mt !,.'TOups. Thm,
urording to elm pcf'pl·rnw. monopl.lcophorans.
ln\alvC'i, gJstropoJs and rmcrorond1' c.m be traced Smcc the ~hell t'i the only pJrt of the mollusc nor-
luck ro the l'arh C .u11bn.m ''lull: ccphalopocb, mally tO be fos\lltz~:d. <:crtain ..t\pl·ct~ of 'hell shape
10phopod~ and poh pi.Kophorans. 111 the light of pre- ~hould be fi~t wmtdcn·d.
1(11! e\1dencl'. do not 'l'l'lll to .1ppear until chc latest
C.unbrian or l.'.lriy Ordovician. The morphologtcal
.:\1dc:ncl' ~ectm to f.tvout the Lttt•r view but, since C~iled shell morphology (Fig. 8.6)
:h~ earl} Cambn.u1 mollusn arc till\. HKonspicuoLL~
md of low dtvcf\tt\. \tlll n:lauvd} latde '' yet known Although some molluscJn ,bells an: 'unply ~tra.ight
lhom them. TO\\ard' tht· cud of the Cambnan there tubes or cones, very m.wy uthtT\, whl·thcr b1valvcd
~m~ to have been ,a 111~1or (,md tftht· t•arl}' Cambrian or ltnivalvcd, arc coikd. Thi~ lOJhng ~~ most eVIdent
,,ng~n of at least some moUuscan das~es is aJJowed) in gastropod~ and ccphalopods. but even tbe indi-
lung-delayed r.!di.ltlon. cmapkd with a dramatic vidual va Ivcs of a bivalve shell are coiled; the 'open·
mcrease in size. ll \vas rim th,tt led to the forceful side of the valve, which 111 life enclo~es the viscer:1, is
rublishmcnt of tht· dtHHU1<\nt lllOiluscan groups analogous to the aperture of other coiled shells.
11bicb became then so impmt.mt. fhc two models are Leaving aside for the moment the highly modified
c
l~ed on the tmport.lllCl' ot the late Cunbrian-carly shells of belcmmtcs and ~qlllds. it 1s not hard to see
e
Ordo\~CIJil radi.mon d1l' daffcrcncc IS in whether that mosc coiled molluscan shells :~re ~imply hollow
tbr mollusC\ of Llm tune bdong to newly established cones rolled up on themselves to a gre,lter or lesser
:!.m~ or an.• ''m ply ne\\. l.lrgl membe~ of ancient ex'tcnt. In such rolled-up cone~. whtch grow at the
rages. Of thc'c v.mom dome' onlv the bivaJves. apertural end only. there :~re very lntcrestmg math-
~·
a~tmpod~ md Cl'plulopmh art common .md impor- emarical propenie,, for chc cmhng. reprcscnred by a
s-
unt fo~\il~ .md w11l be cre.ued m 'ome det:ul. Some line traced along the edge of the shell from the
()f
Mltnrion t\ aho given w the R.mcroconchia; hoYvcver. fi~t-fonned part (protoc onch ) to the aperture.
)-
other three da'"'~ .lrt• rarl' fo,,il~ and wtll not be mvanably has the fom1 of .1 logarithmic o r
\\
.:;srus~d further. equiangular spiraJ d'Arc\ Thomp\on ( llJ 17) nuy
(a) axts of coolong (b )
IOIIhll planosp•ral
qeneralong expanded
apenure
/ '
/
lo)\
/ T mcrE!asmg
qeneratong curvt!
.1hor ,J c;onqte
complete ruvoluloon ~~I
"f!8 8.6 Theoretical rT101'phology of the coiled shelL (a) schematic diagram of port of a helicolly coiled gastropod shell, (b) com-
!!Ted •·simulated shell shapes derived by varying vonous geometrical parameters D, T and W; (c) a theoretical but non·fvndionol shell
• no/ adopted by any molluscon group. (Based on Roup, 1966.)
202 Molluscs
be quoted hLTl': 'it j, pl·rul1.1rly duracteri~nc of th<. The r:ue of whorl t..•xp.ln~ton ( l f) .tfter onl' rev .llllllllllt
\p1r.1l ,hell 1h.tt it dm·' nnt .1ltcr a' it grow;: each lution. In the ,h,1gr.1m 11' = 2, 'mcc the dtlmdc po~IIIOI
incrcnll'IH j, 'imtl.tr to 1t' prcdccc~\Or, and th<. of the ntbt• :tfrcr a ~m~k rcvolunon 1~ t\\1l'l' \\1-iJl crcnt..:l'
whole .lftlr l'.ll h 'Pllrt nt" !-'Ttl\\ th. 1\ JU~t hkc what it it was on~.· whorl bl'fort•. (;Cilei',H<
wa~ bdon:'. I h:nn· thl· 'Pir.tl \hdl cm .ICactc new The po,lt10n :tnd onenr.uion of the gcncm ulue,
mart n.1l .H rlu: om· t•nJ onh \VHhnut chJnging 1t~ curve wtth rt''Pt'ct to the .v.;,, (D) In our exmljlk 111\'0ivc<
shapL In .tn idt·.tl ,hcJI ('uch a' th.tt of S.wtilus) of the Clrt"ubr tubt..• 1' \t'p.tr:ttt..•d b} .1 t'OI1\t.11lt JJ~
r.tdiU\ I, till' eqU.Illllll ol thi\ spir,U \hJpC j~ I = u9 ranee from tht..· uulm~ .1xi, l'qu.tl w lulf tt~ 0\1
\\here <1 j\ .I lOil\tJ!lf and 9 1\ the \\ hok angk diameter; dm I\ D. Septt
through wllll h dtt' 'pir,tl h.t~ bet.:n tr.~eed. The rJtt' of w hurl tramtttmn .tlong the :.Jxl~ (1
Tht· U\ctltlm·" uf rh" "" J\ of ~:.rrowing i\ undoulH- i.e. till' reLHIVt..' dt\t,IIKl' bt..'t\\ een \UCn~ssivc m·~ I here ,,
cdly the rt'.l~on tor it' aJopuon b) so tnatl) livmg lutwn' .tlong tiH' .t\.1' ,1, compared with ~\1 sh,. 11, nl
orgam\lm 111 the 'hdh ot braduopods and mollttSC'i, trom tl1c .txJ,. pan 1t 10 n
m the foramm1il'nd,, 111 tht· hom' oi lllJJJUJtals, attd ( t·ph.tln
even m the cyt:' nl ttlklblle' - wherT the ability to In <oomc of R:~up·~ tnotlch thl' p.tr~uucter~ wcr~ kef' of t.Jv;tl·
grov• •lt ont· t:nd onl)' Without changing 1orm has consont; in othel'\ ont..· or IIIOTL' o( thl'lll wen· m, '~Jll.l, ,,,
been unport.1nt. ro t..hange a~ thl· model was being gencratl'd If · c.;,,,phopc

Nevenhclc~s. 111 n,tlurt· l oilt:d shells often only 8.6b). Thu~ .1 ~.lstropoJ with an im:rc.tsln~ I .11111 itll>~l l
approxim.tlt' to tht• lt.lt:.tl lllJthcmJtical fomt, J.nd \.Vould have a t..OllC.l\ll' ]Jtcr.tl surt:-.re. In gcm·ral tb th,· 'hell
otten n is the dep..rtun.-, lrum the hypothetical that coiled ~hell~ of any one nmUu~can or odtr. ho\\CVl'f
Me ol tht· ~'Tl',ltl'\t htnutun.tl mtert·st. ln anunon.itcs ~hell-secreting group t..,umot re.td1ly be conftB <cph.tlop'
(fo~stl ccphalnpod~) then~ n1.1y be several m.inor with rh me of othe~. 'I hey evolwJ to d1tTc~ ruhul.tr ,j
changl·~ 111 rlw ~pir.tl .ln~k thruughout growth, and fimcnonal ~.:nds Thus g.lstropods .tre U\Ua.ll) COII11CC(j l
nor all pam of the ammonite' nccessanly obey the calJy coued and rend to have J low 11', but J .1\ C<ll lllel'tcc
mic, oi the log.uithnuc 'Pir.tl iomt. fhe ~pacing of vanablc:: and may bt· c::xtrt·mcly h1gh. g~ving \ rc.•ccnt m
the nbs on tht• ~hdl j, nothm~ to do with the long, thm, h1gh-sp1rcd shclk In b1valve\ T IS tnt< turc.
gro\\ rh of the 'pir.tl .md Ill I) alter \lgruficantly ar and 11 'vcry h1gh. whereas 111 brachiopods 11'1sa. whole of
vanou' pcnod' rhrm1ghnut growth. whereas the high but r 0. Umh thc\e shells haw form~
:.perrurt' 1s c,fwn pc~.· uharly contr.lctcd c::xpanded apl·m1rcs <..ephalopod~ MC nomully pbm. i'lllll'
Follu\\1ng d'AH') '( hnmp,on ·, e:.rlv stud1es. the spiral; 11 ' I\ never nonnally h1gh and T = 0. Ut· 5hdl~ h.J'
tht•urt nc.1l nwrphnlo~n t.li the co1led ~hell has of the dtffen:nn· 111 W betwel..'n ceph.tlopod' (Ynchd~t1
attraued nm,tdt•r.lhk \ttt.·nnon. The he\t-known brachiopod~ there " no overlap m sheU sfur hell~ of
recent studte' arc tho~t· ot Ita up ( 19n6), who u'ed a between them, thongh some plarusprr.U gotltrllj:«: CIIU~J>ft•c
compmer-h.l\t·d brr.1ph1r.tl rnt•thod to produce vari- are approxmtatcly t:qu1v:dent 111 sht~pe to m1 lhc Uppl'l
ous kmd' ol hypotht' tlt.ll roiled ~hape~ and was then cephJJopod~. What makes such dtffcrcnccs w t 1.'\ ~·r • ~~ •,j
able eo 'cl' how lllJI1)' n( the'e pmsible types had in fom1 of many pl:tmspiral .md hclic.U shells L~. ru .1re 1
r..~n,
tact bt·en adopted in n.tturl' Hl' was J.hk to generate ever, the shape of the generarmg curve, i.e. the n• ccplulnpn
a largt numbt•r of ttkal ~h.1pe' lhlllg only four para- in cro~s-scction. In ammomtes, for 1mr::mce, the \h. ~ophnrJnS
meten. in the prognmmt': tht· shell ~hJpes were pro- of cadicone shclh (Fig. ~.25) contrasts markc.:: 111thur a
jetted on an o,cil!u,tnpe \crt·en. T.1king the t:xample with that of a compressed platycone, yet it I' c t volvcd m
ofa hdit.tlly t'Oilcd g.\Slropod (Fig. R.na). and allowing really chi~ parameter wh1ch IS rad1c.tlly diffcrrm de) 111d pt
rhat il\ 'hell '' 1m mnrc rh.tn .1 hollow cone growing Of all the possible ~ht..·U shapes ri1Jt Raup \U~ Ill'"'
j, rut!
.lt the .tpl'rtural ~.·nd .md t o1lmg about a vertical axis eo gencrJte, only rcl:trively few hJvc been faun UntOrtllll.ll
a~ 1t grow,, tht''l' p.lr.Jmett'f\ .Jr\.' .1\ t()lJows. be mcfi1l biologlcally, and ~unilar ~hell shape.' Jen'' "~f h
been adopted time and ttmt: again wnhm rhr gr t;d..'l'JI pla~.·c
Tht• ,hapt' of th1.· tube in \ecrion. otherwi'e th.tt bear tht·m. The other type~ (e.g. F1g. ll.t>C) ll IICilll)' ,1
kilt. I\\ 11 J\ till' 'hape nr the gencradng curve not been able to be put to mdi.JJ tuncnonal pu~Jl<r tru.~lt:1fll1~)1
(ctTct lively equiv.th:nt m th1.· o;hape of the apt:r- and so are rarely, if ever. found in any lmngori logt n•e~. · ti
tun.:) . In tlll:' l'X,unplt· iiJu,tr.ltt·d here 1t ~~circular. group. up1dh• sup
bur the apertur.Jl 'hapt:' of g-ao;tropods usually A ne\\ .tppro.Kh 1\ Obmoto ·~ C'XI~IC 11Cl'.
dcp.lrt trom dm tim11. method (Sav.l77i, I 'JCJO) I le re the

Principal fossil groups 203
mount of ~'TO\\ th .m : ddtm•J rl'l.ttiv~ to tlw current

f'l"ltlon ol the ~hell .tpertun.·, r.ltht·r than a tixed ref-
CJ(UCL' framt· as mt·d by R.tup. flm method c.m
Class Bivalvia
!:t'llcratc \'arious kmds ol surt,lt~ ~culprure. and sun-
ubtc:-. more d1relll y the bwlogttal proce~'es All bivalve~ have J \hell tonsa,ung of a pair of cal-
ln\'Oived m shdl h'Towth and morphogenesiS. careous v.1lves between whtch the ~oft parts of rhe
body ne enclosed. Unlike rht• br,Khiopods, whJCh
arc aho bivalved, tht·y ,m· very abund.1nt ,10d diVerse
Septation of the sh e ll today. and most shordull'~ art• littered with thetr
dead shells. The 11l:lJOncy of btval\·es arc marine:
There ~~ a fund.Hlh: nr.1l dt~nm· tion between thme mmt arc bcnthtc .111d ltve mt:nm.tUy or ep1taunally.
mdl> of mollusrs .... htt h .m· divldt•d by int~mal Some gener<l have mcces~full} colomzed tTe<ihwater
pll'ttnom (scpta) .tnd thn~t· v.lm·h ar~ not. habitats. The earliest known gener.1. 'urh ,1, Forddk
l'rph.tlopod o;hdls .tn.• ,1lway~ septate, wh~n.·.tS those are Cambrian, bttt bavnlve' only bec.1me divel"e in
,,f bivalvt·~ .md mmt g.l'itropnd' .trt' devoid of the early Ordovician. They nrc. howt'Vl'r, gencrnlly
~tpt.1, Js arc thc 'ihdl' nf till Polypi.Kophora and of relatively limited abundanct· in the Palacozmc
\aphopod.t. But tht·rl' .11-c sonH:' ga,tropods, living except locally. l t wns not unul the w.1ke of the
md fo~sil, which do h.tvt• wpt.t in thc uppt•r part of Pennian cxtinctions that btvalVL's moved from
th~ shrll. .md they .m• IHH tttKommnn. There is, their origmal near~hnre hablt.u to the ot~horc ~helf
'IOI\t'Vcr, .1 ~'Te.lt dlflcrt'lltl' betwetn ga!.tropod and rcgiom fonnerly inhabited hy hr:Khiopocb.
, phJlopoll shells, 111 th.llt eph.1lopod~ always have a Thereafter, in the Meso70ic, they hl'c.lmc much
Ill tll~ular '1phunde running through .111 the septa and more common, .1s d1d the g.htropods, and trom the
1- nnccttng the th.tmbers: dm strutturt' t~ mtJmatcly early Tertiary 011\\,lfds they h.we come to dominate
r; :ron~ctcd wtth the buoyancy of ccphalopod~ both the hard-~hdlcd ,h,tllow-m.mne fauna. The name
ry rnt and fossil. Ga,tropoth do nor have such a Bivalvia was ori~'lnall) g~ven by Lmna~u~ 111 175H.
l\\' wctur~. and tht•Jr vi,ct•ra nonnaJJy ftU up the adopted from the m.agt' of Bonnam tn 168 I. but for
un :OOic of the tmidt· of till' ,Jwll. t•xn·pt in the c;eptate r:nher complex lmtonoJ rea\Oil\ the later tenus
~ry
'!!m\, Pclecypoda and Lamclhbr.lnt hta hJve bt.·en more
JU- Sclmr lu~h-~pirt•d C.unhn.m monoplarophoran common!} 1n use. fhere 1s much to bt· \J.lJ. how-
US~ hili h.l\c been dt•scnhcd wlurh have septa ever, fi)r 'uppre'"ng the two l.trtt'r names both tor
md Yochel,on er ,,/,, llJ7J): tlll'Se dmdy resemble the taxonomtc corn.•crnes' and to .wo1d confusion.
OlpC a~ of the earltesr known ccphalopods of the
od~ nu1 P/e,lnllltJ(Cr.l~ (I ig H 'i.1), whtch conK'S from Cerastoderma IFig. 8.7)
ilcd it Upper c,unbn.lll ot ( 'llln.t. Plcartllltl(('ftiS, how- The modern cockk Ct>mstodemr., n/ulc (Subdass
the tr, 1\ ·~tpbuncui.Hl '. \\ here.ts the monopbcopho- Heterodonta. Order Venerotda) ".111 mt:lun.u blValvc:-
;)\V- rtl .tre not. lt " quttt' pm,Jblc that the earliest which lives 1n the interudal zone m European .llld
:ubc phalopods were denvt•d trom scptJtc monopla- other waters. It ts still sometime~ rdetred to m the
1ape phorJns, the tiNt fonm being bottom-crawling as literature as Ccmfitlllt cdrtfe though, su1ce Its mor-
edly .,, tllClr ancestol"': rhl' latt·r ones, whKh rapidly phology departs too greatly from th.lt of the type
onl~ olvcd means ofhuoy.111ry (dfectl·d by the ~iphun- speCies of Cardium originally descnbed by Lmnaeus,
~nd propulston, were .tbk to explmt the vacant the genus Ccmstoden11n has later had to be erected for
abk rtUt mchc, whtchw,,, h!lhcrto clmed to moUuscc;. it.
1d tn !ortunltdy, the t(w.tl record ~VC\ no further evi- C. cd11/c mhab1ts a burrow a few ccntunctres
have ocr of how tl11s rt.:tll.lt k.1hlt tramttlon could have deep, in whtch at IJvcs with 1ts two valves JOmed
·oup~ en pbcc: nor '' rlwre t'Vllknct.: from comparative dorsatiy by a hmgc, w1th the lme of dosure (the
h.l\'e •omy .tnd cmhryl)lllh') A' 1s commonly and conumssure) being verncal. fhe two valves arc nor-
po<.C:'i lill'Jttngly rhl c1'e Ill .lttl'lllptmg to unravel phy- mally sbghtly open, allowmg the mmcular axe-
fos~a nl(,, the imn.,ung intt mll'diatc fom1s were shaped foot to protrude bet~\ een them antenorly
~ ~upcrscded .1nd h,l\'l' left no trace of their and the stphons con net ttng the an una! w1th the ~ur
fra.rm• "llllcnce. &ce to project backwards and upwards.
.1 and
204 Molluscs
th\.· (
(cl
whirl
u111bo
am cri
b1valv
PIC\CJ
Vl!rric:
d•rccrc
valve!
!:ttJrm
togc:th
llCdfl'SI
If ei
8.7b) t
A A;~
tect
tu rl
c:nsu
(d)
IIQ11llent place
I I C. J
Cnn
.md
rnaPIIe Ci!VIIV
q1tl (hhbranchl
due<:
card
tht.> r
m11r \Ct b
dc11n
1dem
The
1:\VU '
dong
ment
dl•,td
1\vo
of t1H
'ltt·~
Figure 8.7 Ceroslodermo eclule, a Recent infaunal heterodont bivalve: (a) posterior view of intact shell; (b) interior of leh vok h~:twe
right lateral ViOW of living animal in l1fe position with shell portially removed to show soh ports; (d) vertical section of living anilll' ,hell t
(e), (f) cardinal region of leh and right valves, respectively. fhc 1
' l ar~ a
.md ~a
Shell morphology and orientation chiopod~. an: the l'.lfl}-fonnc:d part of the ~ nt 111,
1 IH. t"' o v.1lvc' on t•nhcr ~idc of the comm1ssurc are From the ,ide the v.1lvcs do nor appear ~) UllllCtni hJV:lJVI
v1rtu.•llv rmrror unagt'' of each other; thev arc ,,,id but arc ineqmlarer.1l and <;omewhat lopsJdell. ft the tra
to OL cquivalvcs An intact Ccmstodermn when the umbones radrate 22-2H ~rrong nb~. cw~~cd Thouf!
cx.1mrnc:d trom I.'Jthcr t•nJ appears heart-~hapcd. concentnc growth lines w hi eh arc rc:corch of many I
w1ch the t\\ o ,.,,IV\.'' ~)' nulwtnca1 about the vcrtKal fonner pos1UOIIS of the edge of the ~ht·U lial ~ir
umbones are ser -;hghtly tow.ud\ one end o{ pr1.:~t·n1
commi"urt' .md their umbones dose together .md
t:teing c.1ch ,lChcr. ·r hc'c umbones. as wtth bra- ,hell, md IllS dus whKh 1\ 1mportant in detemu:::; and aiJ<
l!!t onentauon of th~ ,hell and whit h v.1lvc: j, awa} 111 ose of dangc:r (hg. l:ll1). Tht palli.11 ltnc
· ~1<h In standard oncntation for Crt<llfcldrrma the hJ<o nothmg to do w1th the adductor Jllll\dt' ;\\
lllllhones face antenorly and an: do\l'\1 to the such : tlts the line along w llllh the inner llltl\tlll.:ll
tcrior eud; thi~ i~ typical of mo~t thoug;h not all p:~rt of the man de 1s att.ldtt'd.
~111\'e~. lf an im.tLt Ct r.tSftldtmw with both v.tlves
~nt is held tn th.: h.md with the tumm1"urc Internal anatomy
ltltlal. the lunge honzontal and the: umbones The ,hell (F1g. 8.7c,d) ts tormc:d b~ rht• m.uttlc.
·red may from the ob,erver, tlll' right and le ft wh1th 1s Jn:~logom though not homologmh \\ trh
11l\"es m.· tnunedtatdy d1~tmguished. In thi., oric:n- tbc mantle ofbr:1ch10pod~. f herc are three Jayt'~'> u1
mon the ligament, "vhich holds tiK· valws the ~hdl, :Jll of whJCb are st·cretcd by di!Tctt'nl p,ms
~~:thcr and is untrument.U in opcnin~ thc: .,hcll, is of rltc m.mtle. At Jt~ periphery tht' mantk ti.mns
rl't to the ob~erver .111d ts thus posterior. three fold~. only the outcnnu't ofwhit:h i' 'n1etorv
If dther valve t\ cxanuned from the tmtdc (Fig. and ~hcll-formmg. The tlllddle fold is scmory, the
Th) the followmg mtcnul features arc ub\cn;c.::d. inner nnc muscular. '1 he omer part of Lltl' ,hdl
•~ a thtn Llyer (the p eriostracurn) m.tdl' of
~ ~atrened vemc.1l .m:a (the hinge pla te) bearing d~lrk-t·olourcd tanned protcm. In somt' modern
teeth between wlm h Jre socke ts ('Otrcsponding shdb thi~ is thick ,\11d dtstinrt and protects tht' ul-
ttl!he cn:th on tht• othC'r v.llve: thc~c atlas guides carl'Oll\ <ohell ag.:~inst damage and dissolutio11. Wher·c
ClhUnng that the two v.llves go exa,dy bad. into it i' tine .md ftcXJblc 1t allow~ the m.anufatlllrc nf
~be~ when they do\c, nl.lktng a ~c' ure md nght intncatdy sculpturt·d ,hell morphology and uma-
fit. Dentmon 111 b1v.1h·es is of vanou' k1mh; in rncmanon (H arper, I 997) In pte1iomorph\ 1r j,
Ctrir,;tocf,nna the tcc rh are of h etero d o nt type vani,hmgly thin and t' r.1rdy prc~cn·e d 111 tos,ih. lr ts
and fall mto thn·c groups. In C.:nmrtldrmra, formed b) rhe outemw't m.mtlc fold Jt the \hl ll
directly below the umbo (F1g. 1:1.7c .t) arc the large edge and is th e only shdl l;~ycr to cross rhc: dorul
cardinal teeth , two in the left valvc .md rv:o in margin. Udow this arc two much thicker I,Jyl'r\ of
the nght; the two sC't'\ of elongated and obliquely cry,talline calcium carbonate, fom1ed by dcpmmon
·t lateral teeth ltc ~omc d1scancc: a\uy. Uivalvc: in a protcinaccou' (conchiolin) rn.ttrix 'I h( tnncr
1
'lltltlon 1:. veT) 1mporrJnt in ci.l~\lticanon and layer 'tops short at the p.1lh.ll line.
1dcntitic;atton and i' cons1dcred 111 dc:taiiiJter. WHhtn rhe shell the upper regton " on uplt'd b}
The ligamcnL a rubb,·ry matcnal connccttnp; the the vlstcra.lmass and the lower by the m;~mk t.1v1ty.
t\Hl valve., and hold1ng rhem open lt l1es in an The mouth is set antcrmrly .md the anus :1t the pm-
dongattd pit puslt'tior to the umbo rhe liga- terior end of the rnantlc clvlty. The l.1rgc < c...·ntr;tl
•ncnt 1\ rarely preserved when the shell h,1s been toot is ·• mmcular org.m uscd m digging. The .m1m.11
d.:Jd 'orne tunt. c,m altematcly contract .md ~ xpand it .md thu' c.m
fwo large: ovoid \llllloth areas tO\\ .ud' rhc encb u!>c 1t both to dig it' w.1y 1ntv the ~ubstntt•m .tnd m
the ~hell. Tht'\l are d1L' scars or machment move honzontall} Wltlun 1t. On cithcr \Ilk of the
tn for the adduc tor muscles, wh1ch nm foot the long ftla.mcntou' gtlb h.mg down mm the
1rtwcen
e; (c) the valve\ .1nd when contrarred h·cp the man tle c....tV11)'. Near tlw mm•th two cxtr.l t,'111-ltke
1imol. 1h~U dosed. structure~ arc found: tht· labial palps. Thc~c
Ihl· pallial line whit h joins the t\Vt) .1dductor together wi th the gills .ue U\~·d in feeding
111 and rum paralkl wtth the edge of the shell Su1<.e Cemstorlemra liw~ a~ an infaunal \ll\pl'mion
wJ~ome ciJstancl \\llh111 it. Th1' m.1rks the point teeder protected ''ithin the 'cdtmem, 1t ha' to be tblc
1hcll. f mantle artachmem to the shell. In some to m,umam connection With the: surtacc watt'T\ for
nC"al 11·.Uvcs. though not Cmutodmrl<l, Jt ,l),o marks teedu1g ,md respirc~tory c...xchJnge. Thi, ~~ lanlnatcd
~ rum tbnrammon between microstrut:tur.Jl shdl types. by two large siphons wllldl projcn upw.mh from
d h) Though the pa!li,1l line of Cerastodmllrl is entire, the posterior end of the ,111imal. Water i~ drawn inro
ftlw n.il1y btvalves luvt" .1 del'p indcntatwn (rhc pal- rhc shell thro ugh the inhabnt <iiphon by the r diary
The lilt sinus) tow,trds the rear of th,· ~hdl. Th1s is pumpmg action of the giU filame nts. The,c wm-
f the f t>Wt only Ill genera with rctr.Kt,\blc siphons cal 61.\mcnrs form a comb-hke \trucrur, lined wtth
uung ;nJ illows a pocket in \\ hich th~o·\c c.m be rucked 11mumcrable c1lia, wlu~ h beat in \Ut:cc"ivt• .md
2 06 Molluscs
coordmarc..d \h1VC' of movcment and collectively ~et -;how J renurk.tble clegret> of mherent or actuJit7rd sir. I
up one-w.1y prt'\'lln:\, rhm generating quire strong evoluttona ry p1.1~ttc1ty. Then arc '>ome features 1r Of J1
mw:trd currc..·nto- '1 he ~~~~ and bbtal palps not only the h:trd and \Oft parts th.lt h,1ve rem.uned fairly 'tJ- Clll i
generate the.• current' bur al~o trap the food pamdes. ble smce tht'lr ongm. rhey IJJve not .1ltered mud1
by 'c:uc..·tmg .1 'ttc.. I.) murm Thc: food pamcles or ha ve undt>rgont.' only mmor evoluuonm
adhne to tlm .md .m· nmvq cd ro the ventral edge changes. Suth char.Jch:N art very mcful in claNtll'll·
oi the gtll or p.1lp bv tht' nha. Tht'y then move m a tion tor dctinmg the h1ghcr taxonomrc catcgor.
lmc along tlm cJgc..· unril they come co rhc mouth. they tnclude 'hell tllllmstrudurc, gtll morpholo~ .11
Wa\tt' w:m:r c..onc.nnmg t.trbon dJOxidc. exchanged and dennnon, thou~h the latter ha' on rhc whol N
b~ the gill\, " takc..·n J\\ a) through rhe exhalant been mort' variable rh.111 tht.: other.. Other ch.tr•• ,Jr.
~iphon rogcrlwr with l"XC ret.1 from the anus. which· tcr<i. however, includmg the ovenll form of dt Ul
i~ pbccd vc..r) c..low m the.· l'Xh:tbnt stphnn. <;hell, the rnu<;cul:tturc..• .md the pre<;cncc .md rehm olll
The onl> otiH:r mtpnrtam ~trucmre~ within the development or \lphom .111d .l"OCiated StntCtltrC• Ft
;,hell arc thmc..· n>llm·c..·rcd -.vith it~ opening and clo- Jre more direttly rd.m:d to the btv.tlvc,· spcctti .ur
~un:. Crrmt,•tlmua, hkl' other bivalve~. Ius many nat- adapt<~tiom to p.1rticular lllude~ of lite and arc thU' Cr
ural cne1t11t:\, C\pt:t i.dly bird~. gastropods and of more value at lower taxonomiC lcvck ara
starfi~h. :md tb u11ly ddt:JKe .1~aimt them i~ to with- '>P~
draw the foot in~itk the.· sltcll and to keep the valvt"s Shell microstructure and mineralogy (Fig. 8.8} tlli
tightly ~hut fix ·" long as pm~ibh:. This closure i\ Thc cakan:OU\ ~hell~ of bivalves arc mulrilaym l.lll
cflet red by till· 'tmng honzontal .1Jductor muscles, :llld comi'r of two imcm1ixl'd pha~e~ (Wilbur 0111
whit h arc.· both about the ~am c.: ~izt: (the dimyarian 191)1): ( I) an or~anic matnx JJIJ (2) rry~talliuc: l:i!· lll
lOndition) .ual .madH:d to t:run~ points OD the cium carbonall' in the limn or calcite or Jragonitt lar
mtcmal \\ alh uf tht· oppo~ing vllvcs. When the Oyster shell~. (i,,r eumple, are m;u.le of cak1tt Ho
adductor. rcla:\ thc v.Jin.•, open automatically to let though they have ,1 Lhin .Jr.l~un iti c p.!d \\ h~Tt' the .11lh
the tool out. ltlrc..cd apart b.,. the ligament which acts mu~des arc auac hcJ. Other bivalve ~he lis m
like a <.omprc"ed ~rnng and whmc clfect. acting entirely aragomtK. but perlup~ the majority hm ()t all
alone, would keep thc: vo.~Jve, permanendy open. d11ferenr layer. compo,ed of talc..ite and .tra~orutc. ll('tJc:.tl
Rcc..e mh tkad ,hdh in which the muscles have Vaterite, another fonn of calcium cArbonate. ht\1'\ ,;
decayed c..1r bel·n eJtl'n rll.l\ still retam the ligament, reported from lllJUred ~hell~ w ht<:h the nt.ulllc IUI 1111\nl\1
and the f\\ o ',1h•c' 'ull JOIItcd together are always been .1blc to regcncr.ltc. they a
found m the open posttton. The f\vo phJses tend to b<: found m bl\'.1.1\c.., Ill I hJ
:111.1<
B 1valve' thu~ opcn .md clost: the1r valves by an number of recurrent p.ltterm, occumng tn d1scrcte pti\lll.l
amagomsnc mu,dc..~hgament S)"tem, which is very tltc- cal
effecnve but rcqum:~ contutual expenditure of ontt·r
energy when the ~hdl ts do~ed. The shell closure llllll·r
-
system nuy bl" c..OtllrJ\ted w1th that ofbrachiopods, biv.1lvc
which oper.1tes l)n t\vo set~ of antagonisnc muscles. (S11hcl.l
Though then.• " tW hc.1d 111 biv:tlves, there are fhl rt• i
wcll-devclopt•d nrcul.ttory, excretory and nervous ( a) (b) (c) (d) lllll;lll}
system~ ••tdequ.ttt' for .lll thc..•tr physiologic:tl needs. IIIUdt•
Cmwadcmtrr rs very wcll .1d:1ptcd for lite as an intau- I tll\~Cd·

nJI suspcmron feeLkr. lt h.1s a lugh 61tcring rate and, 11.1\.'t' tl
havmg colom7c..'d the r:tther 'dtfficult' intertidal p.mirul
CJ1Vlr011ll1Cnt, 1\ htghJy \UCCe<;sfuJ. fhcy 3
wluc ha
Range of form and structure in bivalves tly ll ,j,t
Even though c..crt.un fc..·.ltun.·~ of bivalve orbramZ3- Figure 8.8 Bivalve shell layer morphology os seen 1n thm se rh ~· 1
lion: (a) simple prismolic; (b) compound prismatic, (c) ~
tton, \uch a' tht· .tb,c..·rKc of ,1 head, have limited U111hrc
nacreous; (d) foliated, (e) crossed IOmellor with inset !k.,
the1r evolutton.ll) potc..·ntial, the- range m their fonn disposition of stocked orogon•te lamellae; (f] ~ ~ru Sup
and the adapt.ltinm rh.lt bt\.,llvt' 11Jvc undergone (Based on Taylor and layman, 1972.) tropical
bdJ IJycT\ These hJvt hcl·n u~ed m the unr.wellmg whereas the pcrcent,tge of c.tlcire mcre,tse~ pro~'T'l'\
~phytogenic~. Srx pn11ury t)pes haw been dttler- ~ivdy towards cooler w:~tcn.
ltl.lt~J (T.1ylor ('/al. , 1t;(,l/; l:hthunt, l 975). AJthough shell strunurc ,wd mineralob'Y ;~re very
u~efu l in t.txonomy, they ,1re rarely pn:~crved
Simple prismatic \trunure '' tth colulllnar unchanged m older foml~. fcrtnry fo~s1ls may rer.Hn
['IJI\~otul calntt' or .1r.1gumte pn,ms. the::rr ongmal aragonite. bm there are no kllll\\11
.; Composite prismatic 'mtcture wtth rmy radi- mollu~l' With aragontte whiCh arc older than the
mg ant ular cry,t.lh. C.trbomli:rous.
Nacreous stntctun: 111 which tabul.!r sheet~ ot
n,tgomte are found rt·,cmbling a brick w.tll when Gill morphology (Fig. 8. 9)
ut m section. These .1rc usually found 111 1mddle The 'structure of the gill~ ~~ ~ddom preserved in tos-
1d mncr shell layt'r\. ~ils. but it IS mos( vJiuablc tn taxonomy. The gtll~
Foliated stmctun: of lath-hke c.tkitic cry~taUitc.s hang down IntO the nuntlc cavity on etthcr \ilk of
' n11;ed m sheets the foot. In Order Nucul01d.t (Ord.-Ret.) tht· g11ls
Crossed-lamellar 'nul tun.· which '' mmnally are protobran ch , bemg ,m,lll and leaf-likl r.Hher
-:~gomtlc. Here tlw ,hdl is m.tdl· of dmely hke thme of amphiucuram and ccphalopod,. fherr
'l'·llcd lamellae, wHhm e.Kh of which :trc found unmoditied appearance and the iact that thl·y occur
rlun stacked plate\ of .1r.1~unite, thme nf .t(lj,lcent in :1 primttive group suggc~t that they are nor f.tr
·d lunciiJe being indut~:d in opposite dirt•tttom to removed from the ancem.tl type. Filibra n ch grlls
on( Jnother. In some l.I\C~ mtergro\\ ths of blocks tom1 lamellar sheets of tndtvtdual tilametm m a
'·
1- ,j CT)·,tab are found (cosnplex crololocd- la tnel- W---hape, a-. do eulamcllibra n ch gilh (occumng in
e. lar mucrure) With lolll pnnnp.tl om:nt.ltiOJI\. Cl'rclst,ldt'/11111). but in the latter there:: are l TO\\-p,mi-
e. Homogeneous 'tnh.:ture with ~m.tll b'T'Jnular tlom jommg the filament~ and tnaking water-tilled
1l' mhtdral crystals cavmes between them. The vast majority of b1v,1lvcs
fl' b.tw gills belonging to rhr latter two types.
VC li all thc<ic, nacreou\ o;tnKture ~eem~ w be ph) loge- Septibra n c h gills, wluch are contined to ,1 ~tngle
l'. ooll) tht: oldest. not only m b1valve' but m mol- 'upert3mtly of rock borer<~. the Ponml\ .tcc.l
I~ genrrally; it 1\ .tbll the \tronge~t. \\ lmlr r.ti't" che (Subcla\~ Anomalodesmata). nm transn·~d) Jlro...~
),1~ iOl\'rd que'\tion of why the other typL~ evolved if tht: mantle cavtty, almo\t enclosmg an inner dum-
, Jrc le" ~trong? Underlpng the art'.h ol muscle bL·r whiCh m.1unains only a small connection w1th
11lnm:nr rhcre ts .1 'pcctahzed region of irregularly the out~·r cJvity.
• tc.: ~· ~~mJtir crystals: the m yostracum. In CeraMadcn11n
' tJkarcous part of the \hcU is two-IJwred, the
layer being of crosscd-lamelJJr form, the
-:J(f being compl~ x cro~'ed-lamellar A tew
llvl-., such .t\ rhtht' of Super£umly Lunnacca
al>dJ." Hcterodont.1), h.tw three c.tlcu cum layers.
ere IS snme evidcnn· th.tt tht· type ofshell stnu:turc
cru.illy present in bivalws rn.1y partially rdlt:c t their Protobranch Allbranch
k of liJc. Thu~ tro,,t•d-lamdlar. complex
·d-lamcUar and cumpostte pri,matlc ~hell layers
d1e htghcst llJ.rdm·" values but do not have
l!liCULrl) high nlmpre\\tVC or tcns1le \trengrhs.
, Jfl.' conllllOIIL''l 111 hurrowm~ hlvalvt·s. .in
lu;h a good re~ist.HKt' to .lbr:t,llln, bt;t not ncle~sar
rrslstancr to betulin~ \trl'\~cs, would. be u~eful.
sec- Ihe mincralogJc.tl dtffcn:ncc' in thl• ~hell may be Eulamellibranch Septlbranch
heel tr ecological control; for instJlll'l', 111 the mod-
.,jng figure 8.9 Bivalve gill morphology, four basic types shown by
iOU~.
Superbm.il} M" ttlat c.1 (Subcla"" Ptcriomorph:t) transverse sections: shells ore shown in block with the foot pro·
~ea! specie:, h.n'l' l'lltin:l) aragoniuc ,hells jecting centrally. (Redrawn from Moore et al., 1953 )
208 Molluscs
h
t~
d
...
,,---- .
I
t<'
\
"........ }>
----- hJ
(c) G/ycim erts
th
\',
lu
~~
lit
D
(f) Neotrigonis In
fht•
t i :!St~
whet
~
. vre 8.10 Bivalve hinge lines and dentihon (not to scale): (a) Nvcula (Ter1 -Rec.), right valve (note that the umbones lace po*r:
). toxodont hinge; (b) Proeleclo (Ord ), right valve, modiRed taxodont h1nge, foot position inferred, (c) G/ycymeris (Tert-1.« Mt
g I volve, toxodont hinge; (d) Mytilus (Rec.). right valve, dysodont hinge; (e) Sponc/ylus (Cret.), right voiVe, isodont hmgq J Ill·
Neotrigonio (Rec.), schizOclont hinge; (g) Myo lrvncolo (Rec.), desmodonl hinge with chondrophore. [(b) Redrawn from Brodsi-.ao
nJ,I}
1970: (c) redrawn from Woods, 1946.1
m ten
rccth
Fm~tliz~:d ~111\ b.tvc lK'I:Il de~cribcd in Upper l.1rgcr nd~e-ltl.c tet.•th. pr<..'\umably allowing dt .1 chq
Jur.tmc /'n~c,tll<l. Unsurpmmgly, they are very like protrll\1011 of a large foot below tht·m which tht\' ht• .tl.>
the g•ll~ ot \'t•c•/r~~cllllcl. hvmg today. protected from .1bovc while the shell w.lS open.
Dysodont dentitiOn comi~l'i of small ~m
tt.·cth near the edge of the valve, as m Mytil11s u ton.•
Dentition (Fig. 8. 10}
There art· ~evcr.d k tnd' of tooth and lunge srructurc Isodont teeth :m· very large .1nd located on rtth H. l 2h
in biv.llvc~ of whtch the following can be clearly side of a central lig.nncnt pit, as in Spotuljl t \'~ I ,
n:cogmzcd . Such teeth arc ch,lr;tctcri,tit of one superlam ~Otll)
In taxodo nt dentmon the tn:th are numerom and only. the Anomi,1ce.1 (Subcla\S Pteriomorpha: se tllo
'ubp~lr.llld t'r rad1ally ,trr.111gcd, .md in modem ta..x- S chizodont teeth arc co•t.find to Suprrf.m1 \'IIJllS)
odonr bivalves 'mh ·" Clycymrris and Arru they arc T rigolll.llC:t (Subd.m Palaeuheterodonta Utd:l Ill On~
all rarhcr <.itmlar ,'\'uc rt/.1 and i~ relatives have ta.x- Trigout•Jda); they an· very lar~e and have 11lDI
odonr t<.'cth. a~ do n1.1ny of the Ordovician p.trallcl groove~ nomul to the JXlS of the toro: Orl
bJv:~lvc,, though m \Oilll' of the~e there is quite In Tn.~o11ia the kft valve Ius three teeth, rhc n#' M lht
<.mmdt·r.lhk d!tfcn:nuauon of the teeth. In the has rwo. m•dlu

Ordo\1CI.lll palaeotaxod ont genus Pratfetla, for Mmt Tcntarv and R.cccnt bivalves hJve hfl· thOg)
m~t.mre. th( n· I' .1 po,tt.·rior secnon ,,,th lo'" crodont tec~h, a' cxcmpli.tied by CcrJJt,,J b&\ 1Jv
rid~c-hkc t<.'cth .md .111 antenor pan with much Nonnally there are two or three c.udin:tl t clc:pn::
helm\ th~ umh<), ·" wdl ,l\ th1.· dont;.Jtcd lateral in fTont of.md behmd rh<: hm~e. rl.'~pt'ctwd;. Tht·ir
tC<'th ~nrcnor and pmt~:nor to tht:sc. Heterodont function may h<1ve somethmg to do With burrow-
dtntttror <.111 be tr.l< <.'d b.tck to thl' Ordm;cian, mg. bur rhr1r pn·~ence .tnd ~h.lpt' depend on ho\\
tki 11 ha, been \ll~~l'\ll'd th,ll both the p.Uaeota:.,;-the ~hdh grow. Nt·vcrrhdc'>s the.: Tna,~Jt Li111<1 lin-
odonr .md palacoheterodont teeth in the cata (Subcl.m Prenomorph.1. Supcrtanul; Lunan·a)
b1\'llH' of th.1t 11111<.' \H'r<. dc:nn·d ti-om an aiKC~ .1ppan:ntl} tound a uw for Its lunuk. ,1\ h:l) been
,, .. \\1th multiplc-rid~t·d tn·th. ~hown m some mtere,ung ecologic.11 ... mdies Th1s
Pachydont d~.·mitwn on llf\ only in rhc: pet uliar Lima. like it' modem 'pecre~•.tpp.uentl) re,tc:d in
luppunt.d (rudist1J) lm .tl\'t:s. \\hi eh ct:ment life "ith 1t' lunule on the ~c:.1 floor, .1s j, .1ppan:nt
tbwhd\ t:\ w tht· ~ull\lratum by .1 \'cry largt· right from the..· orientauon of t'p1LOtc pccttnan·am and
1 I\~ ot l or.tl-hh· ti.Hitt. Thl' tn·th .tre vt'f} large. othe'r small organl\lll' on the 'hell, and from the
1\:l\~ ~mJ blunt. F\ Hienth' this pn:uliar dentitlon pmition of the shc:lh themsdve' when fo,silized.
I' dirt't tly t unm'<.l<.'d '' Jth the unmual mode of Juvenik \pecimcm of Lillld ,m: otlt:n round clmtcred
life at the antcnor end of the lunule. They have been
Desmodont <.kn llliOtl I~ .I l Ullllllun to011 or interpn:rcd Qdferic\, 1W10) .ls photonegativt' young
Inn):~ structure 111 whKh thl' teeth :~re very which enten:d tht: c..tuk ~pact: between the umbones
rt•duct·d or ab~t·nt. hut .tucs~Ot)' ridge~ lying of the adult and lived there ror a while. T here are no
1lun~ the hitt~e 111arg-in l.lkt: tht·ir plan:, and large juvenile, in the luttuk:\, however, and pre~um
1ttt:n, J\ in My.r, .1 l.1rgt' projt:cling intnn.ll ably thc:y mu\t hav~.· rnuvcJ out when they IMd out-
prou:ss (the chondrophorc) <..1rrie~ the liga- grown their dark ... hc:lter. Many mudt:m Lima
ulcnt. All lm .llvt:s po"l~\lng dc:smudunt binges \pt:cies buiJJ llC~t\ or by\\US in 0.11k l<lVities, d.nd it
Jet' int:tun.tl suspen,ion f~c:dc:1 s ;ec:ms th.lt they ll1U\t have rnhet itcd thi~ prderence
for the dark from theu ame,tors. at lea~t as far back
'!!ft'Jt \,uict) tll\knt.tl \trutture in bivalve~ con- as the T nassit.
11 'rron~h \\ rth th~.· sltuJtiun 111 brach10pods, The onentation of bivalves and the disunction of
erl' 11 1s ;~lmmt t omt.1nt. ldt from ngln vah-e, 1\ I.K1lttated 1f the following
!fi• comidcrJtiom Jft' bomc 111 nund.
;:.) Muscles and ligaments
~~ "he hgJrnt:nt 1~ a VJn.tblc \tJucture 111 bi\ .live~ and Mo~t b1vah-e~ .\rc prmogyral. though \'uw/,t.
,..,.,
y haw t\\0 parts: mw extettl.ll .111d rhc orher Umn and Dtwa.\· ,1n: exceptJOns to thl\ rule.
;:etnal. 1'hc latter m.l) rcs1dc in a pit bet\veen rhe The pallial smus 1'i ah\.,lY~ postcnor. and if the
::tth (e.g. 111 1~odonr ~hdJ,) or nuy be 'upportcd by external ligametH 1s nor central, 1t too 1\ muJlly
:he fhondrophore (c.~. ,\1}'<1). Either <.omponent may postenor.
IC:'Y • ab~ent. Though the adductor; may be the sam<." s1zc, 1f
CrrcllhlCit•tmcJ ;,, .111 c'\,unplt: of ,m isomyarian one i~ redult~d 01 .llheiH 11 1\ mually tht• antenor
ple alvc, in which the two .Hiductor muscle sc.us are one. though not, tor example, 111 L:'mis.
11C or lc~s the ~.\tnt' m•e. 1\l)'ti/us (l~igs 8.1 Od,
~er 12hJ) .md orhcr by~~ally att.1ched genera. how- Classification
lu.~. er, have a wcatly reduced anterior :Jdductor (ani- Biv::~lves have always been found h,1rd ro cbsstfy.
lily wmyarian condition), .1nd in the swimming The problems are not actltc at lower taxonomic
lllop Pcclt'll (f 1g. H I L1) the ,llltcrior ,tdductor has levels, for species, genera and !:tnnlics seem on the
ill) umhed .lltogcthcr .md there remJm' only the large whole to fall into clearly dcbncd natural grouping~.
der 1nonomyarian pmtt•nor .tddunor. These are based upon shell fonn and stniCturc, the
an\ presence or .1bsence l)f .l pallt.1l smllS, dent1t1on and
)th. Other shell structures other chuacter;. Some of the f.umly group-; ha\'e
ght st bl\'alvc~ have tht• umbone' antcnor to the rcmarncd very comervativc over long pcnod' of
dline (prosogyral), but then· .1rc some opis- time, and a number of gcner.l haw pt'J"\IStl·d with
1et- :rogyral gcner.l wHh pmtcrior umbones. Some re!Jtively little ch.mgt• smcc the Palacozo1c.
111a. lire shclh, whl'tht'J' rro\0- nr opl\thogyral, have Howe,·er. lt 1s nor \O c:~sy to ddim· l11ghcr taxo-
~eth re-sed .1reJ, (the lunule and escutcheon) phlcc:d nomic categone' of .1 kmd that c,1n bt· related m a
210 Molluscs
phylogcnctltJII) mcJmngful \V,ty. Then: Jrl' too tcw SUBCLASS 5 HETERODONTA IOrJ R..-..- ): I lctcmtl '
morpholug~t.ll dtu:, ..md mmt of the more mcfitl eulamclhbr.lllch' rn "h1ch mmt modem gcner~ I· do
neJrly .111 \\11rh Jr.tgumuc uo.,.,,.d-lamdlJt ,)1..-Jl,, .td~prnl
stable ch.tr.lt tt•rs .m: tn the \Oft pan.' or shell vmcJ mud'-"' ot hte .md cspt'C't.tlly eo mfJutul ,tph.,nrec.hng.
mJCrmrmnurt·. Tlm t.lll'e' difficuluc-; \\;th the Thc htng•· Hrulturcs nu~ Jq.:<'Ut'r.ue to ) de•mo& la I
fo,"l lnrm,, w l11.·r~.· the ,oft pam have v.mi!>hcd and condatwn
when.· rhc ,lf,I~OlliUl p.m of the -;hell is norn1ally ORDER 1 VENEROIDA: AlllH hl·tc:rodonrli ,.,~th lTIIC be:
rrcry,t:lllizt•d or da"olvcd. r:urthcnnorc. paralld erodom II.'Lth. ~ ·~ Llllfll·'· "nuti . c,,.j,(,J. Cr.uJJ!
evolution. \\ lu~. h ha' conli.t,t•d m<tny hne:. of Cmutt>clt·m•••· I t'lllll ••\lllllhl, T.l/m.1
ORDER 2 MYOIDA. ThuHhdlc-d t->urrowcn uul b..·r~
dc,ccnt, ,1~11n r.mc~ t.lxonomtt dtfficultics. .1nte
vcrv mcqua~Jl\ c:, luu~c degc:nc:rJtc. \lphom '"11 dnd-
A n•tcnt ~.t.t~\tht.ltlon. ~'lvcn below and ba~ed on. oped; q~..\/yc1, C.:miHrlll, Plrc1lo1~. l'tr~d''·
tlur an dw '/ f<'<tlht' ,,, ltwcrtd,r,ur Pu/e,,/ttology, use!> ORDER 3 HIPPURITOIDA: LJr~w nhcn (or.ilload. •cmrnt d
,hrJI mtlTO!>tructurc. dt·utatlon .md (to <;omc extent) exnt\Ll biVah<'' w1th p.Khvdom dcmmuw ~g. Ourr lb
hin~e struttutt', ~ill type, stomach anatomy :Uld the Hipplllllt'.\, R,lllrcrlit<'•
narun.: uf the l.tba.tl p•.Jp, .1~ tht• 'table characters, and SUBCLASS 6. ANOMALODESMATA (Ord -Rn) lhnmmr
in thi-. d.ts!>ilit Jtion \IX 'ubda,,c, arc defined. or boring Ion m, very mmlt lll.'d, wrth JrJgonttll ~heU1 . hI
clcsmoJont dc:nnnon. One ordt:t u11ly, Lhc Pl1l1l.tdomyord,
Ch.trattc:rs ~urh .1~ shell ,h,tpe arc \U closely relarcd
c:.g. Pltc>l,t<lrrw)'<l, Fdmmrdi.r, 11h•wcrllr)'tl.
w hfe h.tbat~ th.u they .trc 'umt:Jblc' and useful only
in cl.t~~•ti~..H1un .tt lower t:lxonnmit levels. Evolutionory history
Virrually <111 tht" m a.~ or btv:1lvc 'toL k.s were estJh· (C)
SUBCLASS 1 PALAEOlAXODONTA (OrJ.-~cc. ): includes lishcd by the: Mtddl<: OrdlwJuan (Pojl'ta, 1475,
only rhc om: l)JU H.R NUCUl Cl!DA. 'mull. procobranch,
1978), and from chc:n on lhl.' lmtory of brvalve~ 11
r.<xu.iom. miJun.1l. l.1bul p.1lp ll-c-dc" wHh Jr~gomtic shelb:
straighrtorw.ud. The scJrch for bi-..alve ,tncc:stors
c .g. ,\;r1w/.1, Ctt tllldc>rrt.r
SUBCLASS 2. CRYPTODONTA (Ord.-Rcc.): A l.rr~dy tooth- however, h.1~ proved le" \Imp le. Such a\ do e:1dst JUt
It~~ (d)' "'Jum). 111 htmJ I, .lrJgomo..- _,hcUed. mainly tmy, of mtllmH." rre ~l.lk. T hl.'y are vanouslv ennl-
PJ!Jcolllt<" group \\lth "''''"'I'••
(ORDER SOLEMYOTDA) agt:d as surface dwdlt:f\, l)'ing on tht>tr stdes, iecdtllj!
l\ the unh h\ ang rcprc,t·ut.ttwc. Cmli~>lo~ Jnd &1ptm.1 Jrc wnh the foot .md rc,pmn~ \\ ith thl.' gill~. or J~ sh~
rcprc,cnt.lll\l" .,,· rhc- ,,nJv other order, dw Pai.J.:ozoac low burrowt•r<;. rtw bc,t-knuwn genus. fwdtll.t,
ORDER PRAEC:\~Dil>DA
from the Lowc.:r Camhn.m. \Celll' to be a VL'f\ 'lllill
SUBCLASS 3 PTERIOMORPHIA (UrJ Rt'<.): A ratht•r hetero-
geneou' l-'TOUp ,,, nnmt.liiY by,,a\c btvJivc" wath VJnablc bivalve. though not .111 would :ll!fCe th.lt tl is 'uch
ntu\<~tbntr<' .md .tcnuuon 'ihdb tn.av lw cJlcmc. ar.t~oruuc (Yochdson, IIJXI) . However, mmt othn authon
<lr bnth. clam1 tlut tt h:1' .1 true lunge line, a smgle tomh .ll)d
ORDER 1. ARCOJDA homym.m tihbranchs wtth ~ocket 111 c:.teh valve, and J 11,1rrow postennr li~ (jl
no\st·d-l.uncllJr 'hell,, taxodont, c-.g !rea, Clycymms. menr trough . Anmhcr nu11Ute genu~ (I mm) 1i
ORDER 2. MYTlLOIDA: Am,omyanm tilibTJnchs .1od Pt!jcttlia, tTom thl' early C.unbmn of Au,mlu
t·ul.lmdhhr,uldh, pnqn.nadn.tcreou~ shdls. byssatt·. F~n
• dy,uJulll, c ~- \fytilrl!, Puua.1, I .it/r,,p/lw.'(•l.
ORDER 3. PTERIOIDA: Ant,oanyJmn or monomyarian fui-
( ~unncgar and Bentley, llJ~2) . l hts h,1s characten,.
tics rcmimsccnt of OrJovJCiau p:llaeotaxodmn~ JIIO
'
Myo
),
it ts qnttc likely lllJt these early, :md tiny, thou~~~ poste

br.wt h~ m cul.ruH:lllhr.tnch,, 'hc:U Hrttclllrc varied, byssate pallia
or cementl.'d Imlmk' .dl ~allop,, oystcn, <and p<"lrl clams; disnnct genera wultl have provided the roobtod sholk
t.J,;. /'l'd!'rl, Ptcrirr, ( .m•illt'<l. ltw<rrmruts. Umn, Ostn·a, E'.-'"!.~ym. for most known hi ~her btvalvc ta:\:a. (" O.i
SUBCLASS 4. PALAEOHETERODONTA (Ord.-R.I.'c.): A dom i- Cunou ~l y. n o biv.1lves ,1rc known fwm th1·
nantly P.al.tl'<U<ll< .lr.ll.:<lllllll-llwUnlt-..'WUp indudiatg the fi11- Mtddlc .md Upper Cambn.ln .md thme f{mntrl} (CoJ
lowm~ unJ.:r,. bclo
ORDER 1. MODIOMORPHOIDA The: PJl.1e0zoic pn:~o~
recorded as bivJlve\ .trt• now know11 not tu bt• ~l'
E.1rly Ordovil i,111 (T rem,1doc) f.1una~ .lie uf low dtvc
,,1 llltl\1 btt·t I:>".JlVl'' \'<tth h•·tt•rndont tl.'c-th (~conodonts);
dive~lty, and o nly 111 the Art·mg did bavah·es a!
\Ill
l' ~· \(,•.lflrf,•r•"'· R.r.l<''"''
ORDER 2. UNIONOIDIA: Hetero.lum uon-mJnuc genera become comm on, \\hen tlH.'y und~:rwent an imtlll by l:
w1th .1 Ion~ tHIIL' r.m~~. •·.g { ,;,, great burc;t of adJpnve radu[lon. Thi' r.1pid eYolu· T
ORDER 3. TRlGONOIDA ll•-.alv("'; w1th l.trgc trigonal sh._.U, tionary dunge tool.. pl.Kl' m the clme-mshore nsc
md "ell ,kvdiiiWd "·h11udont tet:th. Common Ill the dpc
habitat t:IVourcd b) early btval vc,, buc such environ·
MC"\OZlll• "ht·u the\ \\l'rc Jmong'it the mmt numr:rou'
mcnt' arc ~cldom pre\erved m the fov,JI record. One four
bav.Jlvn. """ n:ptc,entnl h) Sr.,tn,~ulct, thl' on<.' li,·mg
-;uch oct urrence from the early Arcnig of W.1ln p.tlp
~I."IIU\, e.g. 'frlS:olllhl
d
tal
ha
8 p
!,
'·
t-
d pallial sinus
lbl ha p
s.
(g)
dv
lg
"'a; pallial sinus
..,_
(d)
s. \/'P
Mya truncata
re
1'>-
ng
il-
Ia .
.all
eh
)f'<, (m)
nd (k) (I)
ij) V
1~
Venus Divaricella J
llia
~ 8.1 I Morphology and mode of life of infounol bivolves, showing the disposition of the hinge axis (ha), dorso-ventral axis
;tS-
6--v). ontero·postenor axis (o-p) and shell modi~cation; small arrows represent incurrents (plain) and excurrents (feathered) (oHdl
n<J 11)11 rrvncolo. (a) external view of leh valve; (b) 1nternol view of right valve, showing muscle scars, pallial line and pallial sinus; (c)
tgh po-.terior view showing gape (all x 0.5); (d) in life position {x 0.2); (e), 1n Mya arenorio in deep burrow with long siphons and deep
f(l•lolsinus (x 0.25); (g), (h) Phocoides, o lucinoid in its burrow, with mucus tube to surface (x 0.5); (j), (k) Venus, a venerid in its
xk ;~)low burrow (x 0.5); (I) Divarice//o, a bivalve which burrows using its surface sculpture as a saw, by rocking through the angle
075); (m), (n) Ensis, a razor shell in its long shallow burrow (x 0.3). (Mainly redrawn from Stonley, 1970.)
the
:rly upt• )')96) comprise' no k~~ than lH genera, flexible: extensions of the palp~ wht<:h project from
so. l.m);lng ro at least seven 'ubdas't''; .111other ts the the ~hdl anti collect food particle' dtrcctly from the
QV. me cnol-w:uer pen-Gond\\ an.1 t:wna, mamJy of sediment (Fig. 8.12d).
lve~ ~I ,(ultow-burrowm~ dt•pom t~cders described Second, there were shallow burrmvtng ryp~' such
inat B2hin .md Gunerrez-M:uco ( l'l'J 1) from Spain. as the Astartidac (Order Vencroida), \\tth no real
11U- Thi, t:.trl> Ordovinan cvolunonary burst gave Mphons (Fig. S. 12e) Then there were t•ptt:mn.ll,
t0Tl' , to ,cvcral supertamthe~ h.wm~ ta..xodont, byssally att.Jt hed bivalves, the pteriomorph Order
on- . udont or heterodont hmgc:~ .wd bclongtng ro MytiluicL.1, ,unonb"\t others (Fig. 8. 12f j), and finally
)nc ur kL·ding rypes. First there wt•rt• tht• c.ulicsr labial thert· armt: the infaunal mucu'>-tubc ft•edcrs, the
·ales lp ti:L'dL·~ of Order Nuculo1da. whtch have paired supcrGmily Lucin.tcca (Subcla~s I ktcrodonta),
212 Molluscs
_t l_ rhe1
not
P.1la
(C) Hiatella
•
(e) Astarte
srn1o
Slph
the •
dw
great
de,n
crod
aho
V (h) Myti/us lubit
prev1
burn:
(b) Aspergillum
k.cy c
mode
(a} Pholas Sm
a
(j) \Jn(l•
a
rowin
l ~~l·nr
Figure 8. 12 Boring, cavity-dwelling and other modified bivolves: (a) Pholas, a stone borer (x 0.5); (b) Aspergillum, o highly~
Red sand-dwelling oivolve with reduced valves (v) (x 0.5); (c) Hiotello, a 'squatter' which occupies old borings (x 0. 5); (d) Yoklio-a C'rc•t:tc
labial-polp feeder(~ 0 5); (e) Astorle, o non·siphonate infounal feeder (x 0.4); (f),(g) Modiolus, semHnfaunal and byssolly ~ S.JJJli.' (
(x 0.5), (h). (j) Myrilus, cpifaunol and byssolly attached (x 0.5). (Mainly redrawn from Stanley, 1970.) <'ally t
C.llly '
other 1
r.ally v
'"hieh wcrt· th1.· ouh P J.l.tcuzolc deep burrowers It 1s worth noting rlut the pre\cncl' ol the mu t>flow
(Ftg. B.ll g. h) . All d1c'1: lct·dmg tvpe' ~urvive ro the fom gave an inherent .tdvant..tgc to bivah:t.., m·cr IIJCt hJJ
prc\cnt. as do tht· \Upcrtanuhc~ m wluch they arose. rhynchondJifonn braduupods, wluth other than m. l homa
Apparently tht• ~.-.trlic't t' pcs \\ere infJunal. but epi- troconchs were d11.· riv.tllurd-\hell~:d suspcru.tou fetd. fum tio
faun.ll ~cnl.'rJ beg.1n to diVI:~ify later in the crs. Only b) me<ms of the toot lOlUd bival\'es colonG: shell dj
Ordm 1oan. in.fuun,IJ CllVIrOJUllt'llt\ SliClC\Stt.J.J.y, wJuch tbe rh tlexibili
In onl.' genu' ol <Jrdm-tnan bivalve~. Babinka, the chondli1om1l'~ \Wrc never n.·.ill) able to do lndet.: (Ofl\Cf\'
shell ha~ J r.tthl.'l n11tou~ couJunnation which ha~ the second great e:\.p.tmtou uf the btvalvcs dunn, 1-ami
attracted comldn.tblc attention (McAlester, 1962). early Mcsozuic Jnd contmuing throughout t'• 11HplcJ
Tlm ~en m 1\ .111 t'.HI~ h1l11101d known only from the Ccnozoil was d1rectly dw: tu the tact that the\'' \\hith li
Ll.mvtrm.m nt the Czech Repubhc. Babi11ka has an burrow. H oweVl'r. burruwmK J.bility alone \Hl oJUilllll<ll
isomy.:~n:tn shell of ~t.llld,mi form, but between the enough. for wh.H g,lVl' the unpcn1s for the 11: lubJtat'
large adductor sc,m .1re nvo chnins of smaller muscle e:-..-ploitation of' rhe tniaunal h.tbitar was the fusllln llom of
~c.1r~ lik.c ~m"h" of bc.1d~. The upper cluin has eight the pmtcnor cJ~~.·s of the mant.lc to tonu true ~tph tb t·m C
scar~. tht' lower VCI) lllJilV tiny one~. These have (Stanlcy, 196H). In the early Ml'\OZOic a large num !P d.)) T
been lnterprt·rt·d ·" toot (ped al) and gill (ctenidiaJ) of new hetcrodout or J~,.·,modont \Upcrbmilic; Jl th~: \
must le ~car\ .111d bc,u .1 rl·m,trkablc sunilanty to those of which by far the llh~lonty wt:n: \tphou.Jtt, hugl r _,
of the monopl.lulphorJn Yropi/111<1. Babmka, bemg a their siphon.~ tor !l:cdm~ wluk remainmg pro! lllort etfc
lucmntd, \\~1\ prnb.tbl) 1nf.1unJI .1nd a mucus-tube deep bdow rhc ~cdimcnt ~tu:f.1cc. ihm-tced
feeder, but lt dm·~ srrnt thJt it rct.1im the muscle Many of rhc~t· bw 111 the mterudal zonl'' tcature~ ,
p.1ttl·rn a' .111 mht•nt,ml't' from an ance,tral mollusc, they t'ltha occupy deep (rd.ltlvc to body ~tze), Jd;JptiVt',
somcthm~ quilt' like a monupiJcophoran. The e\'1- mancnt burrow~ (c g .•\1)',1, .U.mml•t) or, Lkc C:V< llt\ of
dcnce i' \kmkr hut is in .ICcord.mce ,,;th deductions more shaUow-burrow111g Ccr.uftldfnii.T, can b bl dtVei'\C
about mollmt•.tn phvlo~cn~ from other sources. 111 agam qwckl} 1t wa,hcd out and thus re-
thtmselw~. Quit~: pm,1bh the Intertidal zone was Functional morphology and ecology
1101 wcatly colom?ed by bivalves during the All b1valve shells have to be able to open w1thout
PJbt·ozolc 'mce they did not have the requisite the umbones commg together and so the umbones
lttUCmral potenu<tl . I hus the mantle fusion and have to be kept some clistaJlCe apart. At the same
11phon fomuuon. ongm.mng in the Heterodonta at time the elastic hgamem has to be prevented from
t.icvcry t•nd nf the I'.11..\:0ZOIC. appear to have given undue stretchmg and breakage as the bivalve grows.
d!t biV:tlvc groups dlJt inherited th1s \tructure a The~e two geometncal requirements :ue 111 confl1ct.
g:nt nn\ cvolurion:uy potcnua.l Probably the and the shape of any b1valvc shell must therefore be
&,mmlonr hingt·, \\hich \\a\ denvcd from a het- a compromise between th em. The range of shell
trodont pn.·dccc"nr. i' ,1 \Ub,id1ary modification morphologies ava•lable 1\ thm constt:uned by the
Jl,o d'~onatcd \Vith rhc mf.lUn,il ~iphon-feeding geometry of the co1led shell. Many d1fferent com-
1~n The cxp.1mmn of '1phon feeder' mto new. promise soluuom have been aclueved. and can be
1rrnou\IV uncxplOitt•d h.1b1tat' (mtertid,tl, deep simulated by computer moddlmg. Among those
t LtrO\\ing. bonng. c·cc .). ckpcndcnt upon one new extrapolated by Savaz1i ( 1987) arc. for example.
lr) rh.1r.1c ler alorw, ha' been dc·~cribc·d ~ an ideal evolving a whole range of incquivalvc adaptatiom.
m1Jdd of aclaplivc radiation. very wide ~p;1cing of the umbones and h.tving a lig-
Soml bivalve slOl b havc· bct:n remarkably stable ament that is continually replaced a~ it breaks during
mr~ thc1r fiN .tppl'Jt,lllCC. Thm the ~hallow-bur ontogeny. cessation of growth before the ~hell ha~
rowing t.txodont Family Glyc ymeridac ha~ retained coiled to half :l whorl, and decreasing whorl expan-
twntiall) the same fimn sincl' the lx·ginning of the sion rat~ throughout ontogeny. Thert.' are only a
.di-
,a Crt•tarcous .•md Its rt·pre'l'ntativcs luve lived in the limited number of ~olutillm to thc\e common baste
,ed ).lllll' <unclll-\wcpt habltal\ Thl·y arc morphologi- requirements, and these recur quite often m parallel,
ull} umpcllahzed, and although they arc ccologi- in unrclalcd lineages.
oll) spcualizeJ a... rq~anh ... ubstratc: prdercnce. in Shell fonn and mode of hfc in b1valve~ is illm-
other re... pt:t:ts they arc rcnurl,.,,bly tolerant of physi- trated in f ib'!-. !3. I I - H. I 4. Withm the bivalws the
nlh vigurOLh conJmum, and a'sonatcd fauna.~ arc structure of rhe hinge, dentition, rnmeralogy and
oflo\\ wvcrsity. The 'hell fonn 1' detincd by rigid shcU uructurc and compo,ition do not appear to be
mnhanic.tl and gcmnctrical constraint~ and, as characters of much adapuvc ~1gmticancc. They arc
Thoma... ( 1975) ha., \hown, the demands of the unportant in classuit:ation for ddirung major taxo-
runcnon.tl and gcomc:tric<~lnt"t"J~ of tlw animal in i~ nomic group~ but have llllk LO say <~bout the adapta-
tZ<.' !bell did not .1UO\\ the t~mily enough csotutiouary tion of the d11fcrcm !iOrts of b1valvc~ to their
{11- dexibiiH) to undcr~o tUI tht•r radiatlotb, hence its envrronntent.
ed. cumerv,lti'm rhrou~h tunc . On the other hand, the \hapc and gcm·raJ mor-
the Fmuly Tngoni1dac, rhar..Ktt:rlZt'd by enonnous, phology of bi vaJ ve shelb ilircul > rdlclt~ thcrr mode
th~ complrx hinge t c·t·th, 1\ another unportant group of hfc. lndct:d our currc:lll undcr\tamlmg of the
•uki ~hich livrd 111 n>.~rSL' 'bifbug s<mds. They were the ways in which modem bivaJvL·s an· adapted to par-
not domm,Jnl shallow- watc:r burrowers of nearshore ticular modes ofhtl: cnabks rc-a,onablc iulerenccs to
n..u hlbitah during tht.· t•ady Mesowic. but the extinc- be made as to how extinct bivalvcs liwd.
1 ot nons o[ the late Crt·tau:ous proved disastrous for Modern bivalves can bc grouped mto several
l(lll~ rhcm. One relict ~c:m1s only, N£'1'1rl.(lollia. survives morphoccolog~cal categoric' (Stanky, I Y7U):
1ber ,odJv. The: large tt·cth lllaintaim:d valve alignment
O,t.', Jt the \\ 1dc g.• pc required for extrusion of the infaunal ~ha.llow burrowing;
-;ing uge T -slupt•d toot, tor ""hich g.wc the tngoniids infaunal deep burrowing;
Ct\:J more ctfrcrive mob1hty than .mv P.uaeozoic suspen- cpllimnal. attached by byssus thread~ to rhc sub-
non-fecdmg h1 v 1hcs (St.mlcv. 1Y77). All other ~tratum:
1ere tarures of rhe1r unmual morphology are highly cpi.fuunal. cemented to the rock:
per- lJaptive. .1nd h.1d ll not been for the disastrous free-lying:
the.- m:nrs of the Late Cretarcou~. thq would no doubt swi.mnun~
TO\\ t>ed1ver.e and Aoumh111g today. boring and L.lViry-dwclling.
:>li-.h
214 Molluscs
Examuuuon o! the dc.1d shell\ of mmt bivalves and ends for the toot md me Stphom to \"OIIll' out. Pt'dll fl{IOT.
mdccd abo fo,~tl om:' \\'ill .illuw the correct care- and s1phonal gJpcs art· more durattcmtlt of lh lllg CV
go!) to be mti:rrcd . In the burn.m mg 'P':cies. shell dcept•r-burrowmg genera. <:;oml' ,h,tllow burrtl\\'fll rt ,, P•
fom1 •~ .tl~o rd.nt·d to ""'lwthn m uot tht· ammal was (e.~. Ii'llllla . Dwr1nccl/a) hJve a um~>ll' extcnul lt)r ht
.1 -.lm\ or npui burro\.\ er JtH.I w thl· n;1ture of the sculprurt• of ndgc~ on tht' mtlsllk t>f tht• ,Jtc!L ,, PC' j
scdimt•nt mto wi'ncb 1t burrm..,cd. Mo~t bivaJves, DiJiaricl'lla h.l\ .m unusual W -\hilpcJ p.utem 11f lnt( 8 l.?h ,
liJ...c Ct•rcUttldemr.J, arc su~pensio rt lct·dcr~. There arc a ndgcs (rig. R.lll). When it burrow~ tl~ rodcin~ IIJ'II~h
few gem ra. however, mt'mbt·r-. of tht' Nuculacca. movcn1cnt of some 45° i~ aided by tlw ~'"P ~twn '' tlm·.1d
\\ hich .1rc dt posit feeder... Nmul,tceans m eh J\ Lhc ~hell by rh~:M? ridge~ as tt 'saws' 1ts WJ} Jown 111m lilt .ttl:{
) ,,Jditl (F1g. H. l2d) t'Xtmdt· 'Pl'tl.tltzcd extensiOns of the tim· \,tnd m which il li ve~. The obhlJlll' ridges S~ I H'tl t
thctr labial p.tlp<. tnto rlw 'icdimt:nt and so collect · the mort· elongate Te/lir111 .l"i'L the burruwmg func· t.•on , I
or~.mtr p.trricle' trom tt Till' vcm•w1d ~uper&mily non 111 much the l>JJnc \\'3). P.lttl'ms ,udt JS these- :u \I.'IHr.l)
rdlinJ(CJ JTl' p.lrtly depOSit Jlld part!\ SU~pcnsJOn uml\u.tl. Jnd the majorit} of burrower' arc ~111001l ~hdl 1
fi.•ctkr... TdlmiJ.~ u~c thctr lon!t mhabnt stphons to ,hdlcJ .md mt·amlim:d. th~ dh
'ucJ... up Joml p.1taclc~ from the ~eduucnt surf.tcc Sh;tllnw burrowers do not nonn.tlly luv,· \'01 tuo 1h~
r.1thc1 li J...e a vacuum ckmt'r. Otlwr than these. the elongated shdls. There arc. howevt:r, .1 number C' md t ho
bivalve~ dtscussed 111 tht' tollowmg st•ctwns arc all well-known, Jcep-burrO\vmg ~encr.t, wluch hJ\ ~ll~' SI!
~u,pe1mon feeders. ~ hclls of very drawn-out form adJpted for life 1fl rln l11n
deep e)(e<tvatwns which are VJrlu.lllV prmuncur \\ lt ~ I h
lnfovnol burrowing biValves (Fig. 8. 11) R.c:prL'\t'nl.ltive~ of many biv3lvc fanuhc-; h.1v.: mdt ,,, Olfl}" l
B1\ ah c' thJt burrow 111 ,oft ~ubqrata. ~uch a~ our pcndcmly evolved to this mode of ltfc •.md rlJo ol hv"l
rvp~:• ~XJmpk CcrJstcldrmw, lnvc a well-definc..·d ~hdl' han: become mod1tied m wry 'umbr '' Af)'tll
'equclll:C of movements wh1d1 t'n,tbk• them to pcn- The lar~e Alyc1 tlft:lwria. for um.mcc (£ tg lU k rd.ttl\'l"
err.Hc the scdmu:nt. hr;r the foot probe" dowu- a !.lug~l\h bivalve which burrow\ 111 finn \:lnJ IS .t gl!t
\l.<lrd' and 'wells wtth hlood fmm the: circulatory mud. lt ha, a long clliptir.tl dt·,modont 'hdt \\lu 111luhit
syHem; then the s1phons do,c Th1' IS followed b} 3 " very thin, with a cunomly m odified hg.unl!tll •tt·• he\:
r.1p1d ,tdducuvc movt'mt'nt of the two valves which (ch ondrophort·) , much redm·~·d tet·th, and p~ "'rc.Hiy ,
thi.He~ the toot further and ~queen·~ w.ltcr out from noumcd Jlllc:rior and post~·nor g-.1pc~. The \iplH'il Wttlt Lh
between them. When tht· loot •~ ~Ub\cquen rlv are fu~c:d wgether, and thoug h they may ht tl' cxr.lu(k
rellaltcd the shell sinks down mto tht• ~cdimem. lapsed a~ an escape reartmn when tlw hiUt (Z(IuX.tll
Then the muscl~ rcl.lx pnor to the..· on,et of tlw next supportmg them is drained they c:mnot bt• "'t:· \\ lth thi
t yde The .mtenor add m ror mu.tlly rontran~ first. drawn ms1dc the shell. R azor ~hell' \Uth ,1\ I lhuu
tollowcd by the poqcnor llllt': ht•ncc c1 roclcing (f1g. l-3.1 I m.n) and So/r11 h:JVc \'Cl) long, .tbr thl' con
mo,·uncm 1s unp.lrtcd to the 'hell. whit"h c.m be up rube-lskt: shells. with reduced teeth .lt the .mtett '' lud. tl
ro -l5 in some of the more: di,w1d.tl ,bells bur ts cm! alone and pemunent amcnor .tnd pu,ten the t \\11
normally less. Whtle tlm proc..~:-., ( ) f 'dig,_ging in' ts g.1pe~. Th e~ occupy n1bul.1r burro,,., dm\lt \1 ht c ;t( mlin
goi ng on, the s1phom cxrcnd to keep contJct Will! tht·y tan m ove for protection when thrt'.ltcm.:d. h tbll of
the mrt:Kc. Not all deep burrower' ,u e of ~lll h mod•ll tCill\, tl
Not all burrowmg bJVJJve, h.wl' \lphom. Astllrre, lurm. Some lucinoid bivaivt·~ \Uth a' Pltac(lidn (I~ t..Xh.li.Hll
for Jll\t,mn: (Fig. 8. 12c..·), \\ btt h hw' jtt\l below the H.ll g. h) have conventional-looking 'hdl1 lht• llllt.ll
'urt:Hc ha.' open, <.ln-hJ..c m.mtk• l'dgcs. not fused ncar-circul,u fom1 and a nt:.trly hnrinmt.ll l111 'rJrcd' h
tnto ~1phom . Mosr <.lullow-bunm\ tng generc1 (e.g. aXIS J'hen: 1\ a long postcnor (t xJuJ.tnt) \lphon, 1fo•lll hc..·il
L11o11.r. OtJIIIL\·, I 'm us). howevt•r. an: 'lphonJtc. They the mhalant current 1s drawn 111 through • I~
han· cqmvaJved ~hell~ \\ irh tht• two adducror mu.~clc mutus tube connccnng with d1e 't·Juncnt 'llrtut Cemet
~t'lr. of ahout th~: 'amc \!Ze. md often h;n e pallial Tlm h;lblt 1s probably :tiKll'nt, ti)r luunmJ (Fig. 8
~tmt\C~. Tht: antenor-po~tt'nllr hu~:· in tht-se Goming found .1s t:u back as the OrJm m.m. fhc be\t
the dors.tl tlps of the addurttH ~(\tf'i) b .tpproximatcly llli:Jmdvt
par;tllcl with tht' lunge lint!. The: anterior .md poste- Byssally attached bivalves (Fig. B. 12f.-j) •lrsr~:r-. (C
nor 't'ctiom of the t"ommis~urc IIJ.IY be pl'rm.mcntly Many bivalws secrete threJd~ nf the pmll'tlt (t> •n•ltphnlc
parted; rhc shcllnuy tht:rl'll>ll' ~apt• at l'ithcr or both ~en. w1th which rhcy attach thl'lltsdw' to th IJI\tllt'd 11
Principal fossil groups 21 S
rltJor. The mannc ,\J,,Ihtl//1.' (f-ig. ~-I ~f.g) ha~ a taper- Srcnzel ( 1971 ). Thc~c .lrt' pL·rlup~ the mo~t ~ucce\s
Ill!( cylindnol 'hdl and lin·d with only the po~te ful of all bivalve~. h.wmg J very dfint·nt fcedmg
nor pan of the ~hell e:\pmed One 'Pt'cic:' i' adapted mechanism which mwgr.1tc~ dw .lcnvmes of p.tlps
lor hte m \.lit nuC\hc:\. hum mch endobyssate and gill~ in a way that no otha btvalv<.·~ have been
!)pes on~inated the LOilllllun mmwl .\Jyt,fus (Fig. able to do. Oy,te~ .lrL' Jbund.mr m .metcnt .md
S 12hJ). wluch hve' .mad1ed to the 'urt'Jn:, i11 an modem \cdmtem' .md arc normJIIy presuved m
ppnght posmon. wnh thl' l ommi<.\ure vertical. Tht' their nanm1l ltfe po,mon. Man) hardground~ m the
thr~a&•. known .h by"u'. arc \t'Lrcted from a gland Me)uzmc and Ccno7mc an: marked b~ the prc!.encc
locltc:d lt the:' b.l~t' of the toot. r he:~ form sticky of many oystcl'll wlut h ,1ttac hcd d1em\clves to wh.a
~ecrctiom '' luch lurlkn raptdly after che1r fortn•l- was then J harJ. rccenLI) l>Ubmergcd sub\tratwn.
non l'hr 'hell h don~.ued, ofi:en \\olth J flattened · When chey settle. uym:r l.trvac ,ltt<tL h them~elvc~
venml mrf.Kt' otlenn~ 'upport .and stability to the to che sea tloor by the1r kfl v;tlve, which become'
.\hell l here 1s a byssal notch m.1rlmg the base of cemented to the rock.. In the pmccs~ of Lemcntation
the ~hell, !Tom whit b the- by,m<. emerge~. Usually (Harpc::r, J Y91 a; F1g. IU Jl), .1 dun ~hcet o( pcno-
fY too the Jntenor p.lrt of the shell ~~ much reduced, stracum form~ Jt the 111,1ntle margm. Thi~ 1~ secreted
of m,J tht .mtenor .1ddunor 1~ t-,rre.1t1y dinun.ished in dllling cementa! ion and rhn.) ugh it km 'c-xtrapallial
ve ,izc so thJt rhe .mtenor- posterior line: 1s oblique to fluid' fi·om .1 spaLe between the shell ,md peno-
in tlu hinge ·'""· thnu~h tlwre ~~ no cle.1r a~rreemenr stracum and the mantle. f-rom tlm fluid a Lalcareous
1t. why th1~ should bt• ~L1. Almn~t ,tll JUVenile bivalves nuterial crystallizes. lt i~ wmpositionally ltke t11e
k·- .l2ny hrroup .trt• hy~\.11ly .ltl.IC'hcd, and the retention shell and forms J strong-bonding cement. Since
en~ lihy"u~ 111 .tdult\ I\ prnb.tbly paedomorphtc. the two v.1lves must dose ex,~tlly .tlong the ronunis-
y>. \fyti/11., '·' cpit:llln.tl, but m.my of m modern sure, any irregularity in thc ldl valve" also reAected
) h rtbU\'L'' liw p.1n1allv or nlmpktc.:ly huned. Tridam.t 111 rhe right valve. I hm 1fthe larva serrle' down on a
tl[ 112 g.:nu' of c-nonnom thH:k-,helkd clams which dead ammomtc 'hell, both the left and right valves
ich mlubit rropic.•l rL·d\ .md •~ hkt•wt\C byssally will have ;111 unpre\SIOn of the .tmmon1te Oy~ters
ent .aadu:d lt' '-lphum ~re: dm.·c.:rt·d \tratght up and are have a smgle large addunor to do~e the vllve-. and
ro- ~~ltly expJnded. \Vha~:.l\ the: by\\US t\ m1dventral are often of \Omewhat arcuate fonn, With the gtlls
ons 1m the hinge ju't Jllhtcnur to tt. Withtn the lymg honzontally .md the Jntenor-po~tenor .wis at
~ol xrmd.:d \lphon (1\\lll .\(1.' lllllllllll'rable algae <;ame 60° to the hmge axl\ Though 111 mo~t genera
Ooli ooxanthdbe) living 111 1 wmbioti<. rclanomhip the cormmssure 1s mon.· or le" fiat, some fossil and
1th- :nh th1~ dam, J' thev do '' 1th cor,tk Recent gc:nc:r,l (e.g. the Cretaceous Arct,l.>trc.r (Fig.
'!1151~ Though tJ1.111) by".tlly .tl!Jthed b1\ alve~ Live with 8.13b) and the modem o~trm .from] have ztgzag
110\t !le comm1s,urc pl.me vcrtH..tl, thac .trc othen. in commi~sures hke thmt• of nuny hrach1opods. and
·rior hid1 tlm i~ not \O. f>u·,;,, tor tmtancc. ha~ one of thc~e prc~umably fulfilled a 'uml.1r ti.mction (sc:cttun
~nor e two 'ears' Jlong tht· hntgt• hm: gre.ttly enlarged, 7.o). Oysters rnay hudJ up ~ubstantial biostromes,
loch xrending th<.: ltmgl.' hle .1 wing. In the spec1a.lizcd and as they often lie in bclb pJmlld with ~horclines
1bit or Prrrir~, whitlt live' .tttached to alcyonariJn they have beL·n u~ed succes~full y in dctcnnimng fiJr-
.1f1cJ t~rns, the ti.mruon ot the wing \eems to be that mcr shore po~itions .
(Fig. •hal.1m cmrems are rcmovcJ .1~ IJr :l~ possible from Cementation has orif:_rinated independently in
of r inhal,mt rq,"'on ,md Jrc not recycled. ln bentltic some 20 major groups, and always from J by~sate
1ingl' m~d' b1valvt'\ the e:\temion of the ~hell prevents it ancestor. Such tcmcmation i~ tlm~ polyphyletic. and
1,but •m bt·mg overturned by lllrren(.!). cherc: arc: even example' rrom fre'h water. Er/1cria,
long tor inst.'lnLc, fonm large oyster hank\ 111 fresh water
rt>tCL'. Cemented and secondarily free-lying forms in We,t Africa. MJny ~:orruup~ of cemcmmg bivalve.~
l~ an: IFig. 8.13) appeared mdependcntly 111 the early Mcsozmc,
{ bc~t known uf thO\C' b1v.Uvcs which attach cointiding w1th a gn·at lllL"rCJ\C m populauon prc~
~1\Cive' by l'elllt'nt,ttlon to the sub~tratum are the sure. H Jrper's (llJ91 b) c:xp<.•nmt·nul work -.how'
\lt'n Order Pu:nmda. ~ubordcr Ostrema) whose wby tlm should be 'o: prc:daton ((rabs and startish}
colla- i!phology .md eYolutton luvc been full} docu- were ofi't:red a dtoiC<.' of bvsl>Jle or temented prey.
he \l'J tcd m tht• '1rrull>f ,,, illl'crtcbrdtc P.lfC'tliiiOit~\l}' by and the latter ~urv1ved far more dfet.nvc:ly smce the
216 Molluscs
pr~d.Hol'\couiJ not readily nunipulare or pluck off rccogmzable <tl> bivalveli at all. Thl''l' inhabited the autcq
thl·ll'mcmcd bivalve~ from the substratc. shallow troptcal seas of the Tctl1\ an rc.1lm Juring \,th·e
An ~xrinct group of n·mcmcd b1valves, the tl1e Late Jura~s•c and Crctalcom. Ex;tmpks ure ltkl· ...
luppuritot&. or rud1stJds (Suhcla\~ Hetcrodonta), Hippllntc/1,, and Radiolites (Fig. R.13j,k) !Tnm the !Jyl'r!
bctarnc very hi~hly modtfied , \O that they are hardly Upper Crctac~:ous, in which the valve' ;Ire Wl) j;lg.ITl
l:Jr(;l'
~ur(tc
adductor
muscle tht'rl'
Sl10pl
r~rtlm
a cor 1l'
lty IU
ha ~l>k t
valve,
m.tde
p I 976)
adductor
(b) Arctostrea
muscle
flllletr
(cl Spondylus
ur to
lu lle\t•
f(lTIIIil
~ P"'"""""m
~mall
~
ullltril
' extrapallial
· · .. · : . fluid r he
t'llt HI\
had hl
shell cement umbot
(f)
~.I 1h)
R hy Ollt
(g) Gryphaea arcuate
tl.cy r
ne\\ .tt
unn,tlc
'uav.1gi
nc.ttlv
ff>nii•O
(h) Diceras
IIJI fig:
lJt IJ\ V
slwrten
j'fl-.td.J
11110rru
(j) tbk to
(kl Radiolites acic
111 v
(I) Torreites I he fi11
Otlt 11 t(
F1gure 8. I 3 Cemented or secondarily free bivalves: (a) Oslreo, cemented by its left (l) volve; right valve (R) overlying (~ 0 31. (t:
\H: 11 ,abe
Arcloslreo, o Cretoceou$ oy$ler, cemented by its left (l) valve, showing gill, spines, odductor muscle scar and orientation (~OS:
(c)-{e) Spo_ndy/us spinosus (Cret ), (c) left vol've, (d), (e) in lateral and anterior view, showing how spines ore used os snowslm ut Hntd
(>< 0.75); (f). (g) Gryphoeo arcuato (L Jur.), o secondarily free-lying oyster, with the heavier left vol..-e below (>< 0.5); (g)
through the edge of on oyster shell, showe~ processes of cementation (see text), (h) D1ceras (Jur.). probable ancestor of rvdeshcls. ~
*""' ,IH' IJrg
twisted umbones (x 0.3); (j), (k) Rodio/ites (Crel,), showing coralloid right valve (lower) and smaller left valve (upper); intemol s~
run cu t·
lure of latter shown in (j) (>< 0.3). (I) Torreiles, a Cretaceous rudist reconstructed os with thick extensions of mantle margin between~!! lugh &
valve rims. [Mainly based on drawings in Zittel, Textbook of Paleontology, 1913; (d). (e) redrawn from Carter, 1972; (I) redrllW'r lt'~ tittnl
from Skelton ond Wright, 1987 ] ,td.IJH.Itil
the lll'CJll.ll tn siz~:. ' I he right v.1h-c j, conicJI; rhe left The large rudi~t 1~m·eiws is n:con5tructe d (Slclton
nng niH• 1\ nt·arly 1'1.\t and sm on top of the n~ht vain· and Wnght, 19!17) "' hanng had rh1ck extemJOm of
<ltt: bl~ .t hd. The n~hr vJ.!vc h.1s n: I) thu.:k \\ .1lh of ~vo mantle ntJrgin prOJt'ctmg out between tht valvt:
tht• brcr'l, .1 rcl.lttvdy smill body tavity, .md a single mm (Ftg. l-!.131), a,\ 111 the livin~ gt.lnt clam fritlama.
v~:rv l(l!fllltlt' tooth \\ hich articul.ttcs with .111 equally This gt>nm is found in both Oman .111J the
:'g\' pm of tet·th hangin~ tkl\\ n from tht· lower Caribbean and was probably dt-;pcrsed b) l.uvae
rf.1cc of tht· ldi valve. In t'Xtemal morphology along shallow \tag111g po,ts' m tht• P.1cific .llld east-
ll'IC .1re ~ome '1mli.lnt1c~ to the b12:.trtc Permi.1n em Tecbys.
rnphomcnide br.lt hiopod gent'ra Ccmd/aroia and A geomccncal model for the b'Towth ,111d form of
Cy.fu•JIItlwria. and m a remote 'ense even to \Olitarv rudi~t b1v.1lves and its implication for their da~"tica
on!<, but tht•n: 1, tertainh littk funct10nal \tmJ!ar- tiOn ~~ gwen by Skclton (1978), and ;1 full treatment
~ to the l.ntcr \nme rudJ,tld, ~ccm to have be<.·n of rhc1r evolution, etology ,md role as reef formers
•h to \lick in water through the perfor.ltl:d left may be found in K:lllfinann ;md Soh I (19H l ).
aln, .md dt•uilc:d funct1on.1l analyses have been The very spiny isodont genus Sp(l11dylw. which
maJ ,hO\\ mg pwbabk currcm <.hrett10m (Skdton. hves tod.1y m coral reel~. is usu,tll) cemented. Some
976). Most md"tid' were 'null. on.ly .1 tew n:n- speoes. however, have become 'ccondanly tree.
lmctrL'~ h1gh, but then: .m~ \ome immense spec-it·~ The Cretaceous S. ~pi11osus (Fig. 8.13c-e) ha' spines
up t•' 50 cm hi~h. They arc found m Cretaccou~ Jrr:mged ar 1ight angles to the ~hell margms: these
111 tone~ in Fr.\nce .md che\\hcrc, ~omctinu:' apparently acted a' snowshoe,. pre,·enung th1~
m ng extctN\'t: duster; "l11ch 111.1\ t:vcn fiJrm frce-livm~ btvalve trom sinkmg mto the ,ofr ooze
IJll reefs. tltllugh only \Ome spt'LIC\ actually upon wh1ch ir lived.
untnbutc tow<H<h reef form:mon. Among<;t other secondarily free-lying gt•nera i~
I I ~:arlicst rudM' (F.11mly Diccr.H1dac) \\ocre rhe large oy,rcr Cryplwea (Fig. H.lJCg). whme evo-
ntomte~. attadtcd b) the umbo of one valve. The lurionJt}' developmen t has been much deb.Hed in a
(lJl h.1d hc:a\)' ~hdh With an e'\tt'rn:ll lig;lnH!'nt, and the 'cries of papers since the early I 9-Hh. r never med to
umbones wen: twl~tcd in .1 \pirogyr.1l' f.t~hion (fig;. have much enthusiasm for tht• unattractive- looking
RDhl. This comcrvative t Othtruction wa' retained GryJlhaea. I felt that 1t had too IC\'1' morphologtc al
\ onr: group mml the end of rhc Crcraccous, but characre~ to sust:un the endle'>s flow of p.1pers
<el :DC\ n:mained J\ cnmt\ter. :md d1d nor invade which purported ro describe its e-volution, .1nd some
rw .ld.tptive zone>. A senmd group, however, the of these works d1d m fact prove biollll'tncally
uucoJkd rud1st' (Skdcon. I ')78, 19H5), have .111 umound. H owever, the excellent recent 'tmlic~ of
llll.lgtnatcd' lt~,unenr- tl j, no longer marginal but Johmon and Lennon ( 1990) and Johmon ( 1994)
n ltl·, rucked ,1\\'.1) wnhin till' ~hell rhe growth have restored thi~ genus to emmenr re.,pect.tbJlity,
~1m1 of rhe shell 1~ thus lWt runsrr:1ined by .111 exter- as tllu~tratmg a complex, tiOil-lll11direction:JI and
Hll liA'lmenL 1t W.l\ only btcamc of th1s that the b roadly graduaJ ch.mge through tune. Cryplwm hJs
h··ll1 were able ll.l uncoil, and Skelton regards the a very thJCk convex left valve. nece~!klr)' fi:>r 'tJbJlity,
iliorwung and lmagmation of the h~mcnr a~ a for if Jn individual was ovcrrumed the comnm~ure
rr-acbptation tor rhe1r raJiJtion into hitherto wouJd be blocked, wHh fatal rc~ults. ln thi'> tt)ntext
,JIIOI'tllpied ad.1pt1ve zones. Since they were no\V selection seems to have favoured a more stable,
~bit m grow clon!tared tubul.1r valvt''• they could broader and flatter ~hell !>hapc ..1nd mcrca,t·d size as
1.1dt> ne" hablt.\t' as 'elevarnr.' and 'recurnbeuts '. an adaptJnon to a ~table food mpply.
The tonner art' tJll romcal or barrd-,h.1ped shelb. Amon~r.,t the strangest of all b1valves are the alaro-
1iren tanning th1ckets, and fet·tling in c.1lm ware~ conchids; the 'giant clams' of the Terhyall Pem1ian.
1.3); (b) 11'el1,1bove the scdunent ~urf.t<.l' and l1llt of the zone GcnerJ :,uch a~ Sllikamaia (Tachillftltl,l~ia) reJch
(x0.5),
wshoes If muddy wJter 1-tccumbent\ , on rh~: other hand, length~ of up to I m and the \hell was up to J cm
section lit' l.1rge ex'tendt·d 'hell~ lym~ freely bur \tably m duck. The~c ~hell~ had wide wm~-like Aan~c1 on
:h,with urrenHwcpt ~hore~ on the nest~ of build-ups. The each v,tl vc, extending postenorly and giving the
Jl ~true h11!h diversity of 'urh Cretatt•ous um·<)Iled rudim appear,1nrc of a very A:1ttened shell. The juvenile~
-een the
edrown ()(ttied ro the ,tdvantage g1ven by till imtial pre- were byss.1lly artathed but the large and heaxy
&bpt;mon. (I 0 kg) adults lay freely on the scdunent ~urtacc, the
218 Molluscs
wetght being p.u tl.lliV bornc bv the flanges. Tlm ts row hke o,1r~ whtl~t the .uum.1l '' sw1mmmg. wh•rh the ,
one of the tC:\\ btnlv~.·, that lav fi.1t on the sea floor ,tdds to the spet·d of mov1.ment ,hcJh
with J venil"JI lllllllllt"urc (Y .mccy and Boyd, Pccrt'll and L 111111 prob.1bly mhented thetr n.lt.W>t'\ very
Jl)~D). abihty from a common .IIKestor. and there 1' (\1• arl.' w
dencc that as t:u back .1\ the Carbomll:rou~ som Sl'Lrel
Free-swimmmg forms (Fig. 8. 14) bt,·alve., could S\\1111. Fm,ll Pet onacca .trc no; tr:H.: Ut
fhe large \Callop flt·at'll norm.JU\ ht>s free on the sea uncommon . Some: of them were hke Pcaen m mor- ~()I\\.'(
Roar. bm tt c.m '"''m by v•~orom and repe.ued pholO!-,')' :md h:tbu (e.g. the Carbonitcroll) 1\!0lll
clappmg of the ,-,,lvl.'~ w~~.·thcr 'o .1s to expel water Pteri11ap1xtrn): orhcr. \tKh ·"the Devoman to Juraqc wlnLI
m succi.'S\l\'1.' Jl.'h on bnth "d1.·' of the 'car-.;'. Such posidonian genu' Alwtru .111J m rel.ltlH'~ may ha\'r ahom
activtty •~ cxhatl\Ung tor the btvalve and catmot be been speciali7ed for .1 ncktoplJnktomc (cntud) Jtltll I
~ustamed tor VC!) Inn~. but tt IS nonnaUy used only free-,w1mmmg) mod~.· of hfe . Hmt!otli<t 1s characrcr- thlll\ .
to escape from prl'd.Hur. •sncally present m hbck ,Juk·' (l'colog~c:tlly equJVJ- Sl)llW
In J>crrl'l/ thL n\o V,\IVt'\ Jn: tlllt'I.JU.ll 111 size. thc lcnt to the earht·r gr.lptolitll' f.Kies) in which d1e Anou
lower hem~ the more convcx, hut they arc nearly only other !(,~si], .l rl· ,Jilltllonitc~ or gon.iatJtl'S, hu• ror th
cqml.1ter.ll. On l.'lthl'l' Silk of the umbo the hinge i~ it m:ty al~o occur in ,],;-allow-water limcstonn pn rmt
prolonged mto two 'l'.IT,· ofnc::~rly equal sizes. The depmin:d as lime mud~ fine t'IHJugh to preserve the wood-
ligament i' wt ~.·~.·ntr.llly .n1d 111tcm.tlly and cmphccd ~hclls (thr.; p.lr.dkl ~ ilh gr.tptnlitc prcservJtion 11 ,., olvt
m J mull tn.mgul.tr pH. A smgk large .1dductor ag:tin JIOtcworrhy) All pmidoni:ms arc thin-\hcUrJ l';ti.Jco
nmsck octupll'' much of the central ~pace. with only two \hell l.lyl.'rs. Thl.'y h.we gapt'S un trom c
Ev1dcntly tlm kmd of ,hl'll wa' derived from a either ~1dc of rht• hmge .1~ in thl.'ir living pcrtmaet'lll Un•
by'~" lly ,Jtt,ll hl'd .IIHl'\tor. d11. t\v o kmd' of \hell relarivl',, bm rht•y lll'VlT had .1 by:.,al notch Jt :Ui) 111 rhc1
ha' l' lllJil\ ll..•turl.'' 111 rommon, though that of time in their onlllgl.'tl} .•md they prob.1bly nci'Ct round
PfCrcn I\ t'\ll.'lllkJ h\ 111 lilt re.t\ed umbonal augle. went through .1n .ltl.ll hcd pluse at any Lime m rheu (Hl'Up,
,MI~t111g it\ t ,lp,llll) .1\ .1 hyJrott11J. life lmtory. Spcc1111en~ .m: .tlmo~t Jlways pr~~rYed S\lmet
Un111 j, 1 hy".11ly .ttt.tched ~t·nm. but it can abo with both valve\ open. thou~h m modem limJCt'JJ \\",1\ Ill(
\Wim ·" an t ,t.lpt• n:.lt'tlllll . I nthvidua], can rclca,e and other ptt·notd.' tlm 1~ rarcl" so. ExpcnmcntS by clfc:n.
che1r by'"" .md w.11n by \',tlvc-dappmg with the Jctfcne~ and Munvn ( 1965) showed that the vah-n Luc j m
commis,urc hdd vc:rtic.tllv. Tht· m.uuk 1s here pro- would be prc,crn·d 111 the open po,ition onh u mnpk
longed imu 'tt·ttt.ll'k'' .1muml the tomuus~ure; the~c their nomul opcnmg .111~k l'Xcecded 60°. 1\ lu,b bUL 1111
nu~ indeed have been thetr nonnal angle of opett- htv.Llvc
ing in bl't\vl.'en \Wimmm~ tontracttons. Funher thl'\L' tl·
resillum experimenD and L.lk ulatmm have mdtcatcd dui ••pproxi
~uch a swimmin~ b1v.1lvc would not haw 'u1:k Nnt
rapidly. c~peci.tlly 1f the dr.lg ell~ct\ were inucJv·l 11\'l. hi
by ,l fringe of Sti ff letllacles ,lround the commtSSUI~ :Somt: .1
(b) .1s in Lhe modern Li11111. l tl thl.' c:
adductor Posidonin therefore mny well have t•xploncd Jn
muscle ecological 111chc, that of the: perm,lllc:ntly swm1mmg Ecolo
nektopl::tnkton, wh tch no Lu~.·r b1v:tlvc h:ts bcc:n Jbk Wluk
to mv;lde ~intc tlw cxunction of this gcnm 111 tb~ shown t
Cret,lccous. ut extin
.~r.· 11101
(a) Pecten Boring and nestling bivalves (Fig. 8. 12o-c) l'IIVIIO!ll
Ccrt.llll btv.1lvcs.m~ .l<ilptcd for hti.- 111 h.ml 111~ ht>w he
Figure 8. 14 Free sw•mmtng b1valves (a), (b) Peclen, a swim·
ming bivalve in which the anterior adductor is reduced com- stratc~. The ~tOill'- .md wood-boring ~:t<ncn U\c.:d tU)!
pletely ( · O..i) - brood arrow shows the direction of shell l.-~tlwpllt~~tl (Order Myttl01da) .U1d Tcn·dtl (Ord<r ro111uent
movement when woter IS e1ected (small OITOW$) neor the hinge; Myoida, Suborder Pholadina) haw elonptrd A ,tu(
(c) Bos1tra buchi, a Jvrass1c bivalve, possibly nektoplonktic, with
valves open in inferred life posillon [(a), (b) Based on Stonley, ~hells of cyhndnc.ll fom1, and like modlfil.'d d<'q Ill God,
1968, (c) based on Jefferies ond Minion. 1965 J burrower\ they ltve With the1r long "''' nonnJ! sym ( nlo
thr ~urftce .md h:1w Vt'l) l'xtendcd stphom. The Most Palaeozoic b1valve' po~se~~ few d1agnosnc fea-
lhdL, of tht:,c btv,tlvc' .m: nn thm Jnd must be tures and are rhus h.trd to cla~~~fy and tdennfY. Only
m n.•,i,tant to .tbr.l'>lon. for the edges of the shdls when the mtenor IS reJlJ~ well pre-.ervl·d tan the
lf( u,cd m cxc.w.ltm~ tht·lr burro\\\, atded by aods anatomy of the ~oft parts be reconstruncd w1th con-
l."i:rrtcd by cell, Ill thl• tn.111tle. [he 111\0JubJe perios- fidence .md thus allow the ltk posmon and habtts of
tracum prcvcnh thl'lr <m·n ,hl']l, from being dis- the vanous t.1xa to he mtcrred I he Mollbos f.tun.t i'
hed. Frcqucnrly the 'hell t•dgc' Jrc prov1dcd with stltcified (through the 1111ttal .lgl'ncy of endoltthrc
rout 'pine,. mt•d ·" ' l r.tpmg tool' 111 excavauon. m1croorbram~ms). and WJ\ prt'\crvcd 111 51lll. The
which i\ ctli.-ned h> rorking rnovemcn£<; of the shell bl\·alves, tsobted from the mudd> m.urtx by ,tcid
.1\•ut the long .txi,. l.,oml spCliC\ of I..ulwph.t,l!d bore etching. arc prc~l'rvl·d 111 such dctJII that the1r adap-
nro he coral, .md hJw l11ghh \pt:et,lii7ed ad,tpta- tive i11orphology cJn be t>~t.tbltshed.
:Jillh Nor all htghl} .lttcnu.ltcd b1vJive' .uc borers. u~ing muscle ~c.tl'\, ~hc:ll sh,tpe'>, ~17C and muscu-
~>llll' hve 111 tinn \Cdlllll'llh, c g. ISf'C~I!tllrull (Order lature of the foot .•md by compJnson With hvmg
\norn.alodc\nur.t; F1g. K. l2b), .Uld Jrt: well .1cbptcd biv:tlves, LiUed.thl wa' .1bk· to J\~t'S' tht· burrowing
lt>r tlm mode of liti:, t•vidt·ntly tlm represent\ the potenrial of the v:tnou~ spt:cit·'· St.ttl~tinl .m.tly~" of
l'nmitiVl' Londinon frn111 whtt h the rock- :md the trophic ~tru crure of tht· COIIIIllunity ~howcd that
wnod-boren. late! evolved. The bonng habit hJS it wa~ dominated (>901,l1.) by deposit feedcrs, which
tlolvcd nu•c t imt·s i ndq>cndcndy since the early unlike suspension ft•etkr~ Jrc .td,tpted to a rby-rich
PJhcozoic and h." hl'lOilll' c\pccially importanr mud environment. There Wl'rt' three differt:nt f(·ed-
r·1m the Me,u70H. onwJrtls. ing levels within the ~eduncnt (Ftg. H. IS): (1) 'ur-
Boring bivalve~ .\rl· roml1Hmly found pre-;ervcd fact:/scmi-intjun.LI, (2) iniJunJI .llld (3) deeper
tht'ir burrow~ ..md \\ hl'fl.' the bunn!P' tit tightly infaunaL
ntund th~ .,hdl' 11 would \eem probable that the OnJy rhe shclh of rhc tiN category werc often
up.mt wa.' the t rt·atw of the domicile. fragmemcd and nomully cncru,ted by cp1bionrs,
5->metune~. howcvc:r, the los~1l widtin the bonng and likewl\e rhc int3unal 'Pl'Ue' wen: more com-
lb mcrd} OlrUp) 1n~ .1 '.llJted r~:,id~:nr~:; it w<tS, in monly Jrticulared. Tht: ccologKal .mumptiom made
dt((, a \quJtter '. Kellv ( 11 >130) has sho\\ n that rhe abom dm tauna of truphtt:ally tiered dcpos1t feeders,
~ Jurassu; H11ud/c~ ou urs in two habJtab. tint as a mhabinng a muddy 'ol(-bottomcd cnvtronment,
f :nnple by,.,.lJ llt'\tlcr on IJJrt.l ~ub~trates. and ~econd, fully accord \virh tondu\1011\ reached through tunc-
h tilt more common I). 111 honn~ nude by other tlOn.ll morphology.
~lrcs ptnetr.mng hard sub\tr.tte' (rig. 8 12c). In There IS a dear c.omr."t between the life habit~ of
·r ~e the contours of the I liclldla ~hell do not closely the Mollbos a~~emblage and the ~omewhat younger
H proXJm,\te the shape ot the burrow. Grogarnshuvud assemblage, .1bo from the Siluri<m of
k Nrsclmg btvalvcs l.lnnot bort' but .1re photonega- Gotland (Liljedahl. 1991). Hen: the lul inmd 1/rowd
·d vr, like Lilllcl, .md tlllllpy pre-existing cavitie~. prism is touud, u~ually in Lit(:- position 111 sh,illow sub-
t' .• ;.,mr Jre byss:dly attJched, .llld the1r shells grow to tidal lime mud. lt 1~ very )tlllllar to livmg, lucinoids
tthl' c.wity even 1f tlm 1\ of Jrregular shape. ( l~ig. 8.1 I a) and wa~ likewi~e a rllucus-tube fet'der,
living at ~ome depth within the sediment. llia11ia
Ecology and palaeoecology oriented itself obliquely to the direcuon of wave
~lult' ~hell ~hapc .md other factors have been action, thereby maximi:zing the intake of suspended
~own to be useful 111 llltcrprettng the mode oflife food particles through the mutm tube and avoiding
If cxnnrt b1valves, other k111d~ of ecological study touting by its own wa\tt' matenal. lt may also have
more dtrectly COIKt'rnt•d wtth the orgamsm- lived in symbtosts wnh ~ulphur-ox.id1zing bacteria.
monmcnt rdanomh1p. l'he followmg examples The following Mcsozoic example ~~ on a larger
lb- ·11 hm\ hiolo~'lcal :1nd gt·olo~lal tnt<:ria can be scale bm equally of mtcrest I he Great EstuJrinc
~ra ;N together m g~vc: mfoml.ltlon about pa't envi- Series (M. Jur.) of the Inner I kbnde' of Scotland
.kr ncnt' wrth f.tnl ,_t nntn,Jkd f.nma\. consist~ of a 'hale and sandsronc ~cquente Ill whiCh
ted \ ,tudy of .1 <;tlun,m btvalw fauna from Mollbos hivalvcs, along w1th some gastropods and ostra-
~ep Gocl.md (Swedl·n) romhmt·, both auto- and codes, arc the most dommant fmstls (Hudson.
to rcologi~·.tl dat.l (L1Ued.1hl. 19R4. 1985 , 1994). 1963). fndividu.ll b1valvcs are very common but
220 Molluscs
1 Nucutoidea lens 6 Freja fecunda 1!1t'01

2 Nuculodonta gottandica 7 Molinicola go!landica
IIlO lT
3 Pataeostraba batuca 8 Goniophora onyx
4 Caesarietta llndensrs 9 Maminka sp. w th:
5 Janeia situnca 10 My!llarca? sp. ~ll, ,,
currc
l,IV,lh
rnhnt
lu ICl
lm b
tH·ttc
mnlh1
Runr
Ro
\\(.'fl'
t"\Jil11
mllcd
llll y (
fi1r tb
Figure 8. 15 Suggested life positions of Silurion bivolves from Mollbos, Sweden (redrawn From Liljedohl, 1985.)
I lwsc
l'lli.IJI,
belong lll onh a le\\ \pcoe~. and within the Unit, and the small I 'i••ipllr/t()· through \-lr.Kktd (IHllh
~cqucnn: then m: h.mllv .Ill} nom1al manne fossi.b water (the mo~r 'dtflilult' env1romnenr to colomzc ) llllll
c'Cn pr .lt the V\.'1) top. ~uch J\\ociatiou i~ normally tor phy,iolo~cal reasons), where only tht• ettl"\iu nt v.1J.
md!c;,u:ivc of .1 'c.IJI1•rult' cm•tronment. which fc:w line .\'eom10dvn wa~ present; thence to hr~rh~
'pcnc.·, llJvc belli .1hlt: to colonize and in wluch the marine. where oysre~ and mynlrds thnvc:d· .md
'PClll' that h.tw Jonl' 'o are mccc:ssful. The beds finally to fully salme manne env1ronmen~
wcrl' JcpmitcJ 111 very ,hJ.lJow water a.s shown by More recent work on oxygen .1nd carbon 1\().
mud c:r.u.:b .1t u·rt.tm honzom ~uggesting periodic ropes has shown rhc e~sennal corrcctne~s of dm p•r·
de\ICl.ltiOn. ~Ulll'\\1\'L hthologies within the ture, though a few modlflcanons were nct:dttl,
\equc.:ncc nlllt.nn lm•J.Iw launas. Hudson tound notably that Unio and J\;romltldtl/1 of the Jun I•
rem,1rkdblc.: .lllJ log1c' \\ ith the present shallow apparently lived in nonnal manne condinom
lagoonal bJY' of till· Tcx,,, co.1st, from the point of
viL·w bc.Jth ot envirmlllll'llt Jnd of the bivalve gc:nera Stra tig raphical use
li v111g there. ~c;·vcr.u ot these modem genera l3ivalvcs :~re on the whole f.1r too long-ranged 111
(.Hyti/us, Osm.'tl ,md Ulllt•) have dtrecr coumerparrs rime to be of much zonal v;tluc. However, tht•
m the Great £,tuaru•c.: Serie~; furchem1ore, in Texas have been used 111 a broadt•r \trJtib>Taphical ~cmt, ~
thL· bivalve ''''l'lllbhgl'' 111 t"Junal content, which in Lyell's divisiOn of the TertiJry. which has ab!ln
can be dmcly tompJrl·d with the Jurassic ones. are dant bivalves and gastropods, mto four ~em·, boNd
~.~limty LOntrollcd Thl· Texan Cmssostrec1 specie~ upon the relat1ve percenk1ges of the mollmcan l~t:·
.md mytihd~ hvl in \\Jtcr of rJther reduced salinity. nas present therein now hvmg.
and H ud~on llllerrl·d thJt analogom Jurassic associa- The one nrcum~tancc under whiCh bivalvcs hl\
(b)
nom hkewi'e hvcd m hyposalrne water. been used successfillly as ~trangraplucal indicaton
By marchmg tht• analo~ue~ Hud~on showed m the Briosh Carbomfcrou' Coal Measures, whet:
dearly that the ovcrlappmg assemblages were con- non-manne biValve~ .1rc abundant at cert.lm hon-
FigureS
rrolled bv 'almm ''anat10m through geological zons. These genera (Cuiltllll(tlfa, Xmaditn m! [Dev ),o
nrm·: trnm lrt·'h w,uer (donunared bv the b1valve • 111thraco11ma among.t mhcrs) arc not unlikl' lh: x2 5). (I
m01km freshwater l '11i11, hut they differ 111 certain of the sheU byer; arc contmuou~ acros~ the dorsal
morpholog~ral char:lcter<;. The ~pecks are not easy margin. T he valve\ mu~t luve been held ng~dly
di~nngutsh and .m.• r:~rhcr long-ranged. But even together. the dorsal m.1r~n functionmg at best as a
~ 1ix or ~rvrn 7onc'> h.1ve bc:t:n defined u~ing con- poorly elastic structure.
(U!Ttnr r.~ngc' of ditfcn·nt 'Pl'Cil'\ of non-marine The earliest known genu,. Hcwu/tipe,(?ttra, ts
:.Uw , and the: zone' thu' dcftned have been cor- Lower Cambri,m. Later rmtroconchs rcad1ed the1r
, bowed by plant and ~pore lo"ih. maximum devdopmenr 111 d1c cJrly OrdovlCian,
almost nvallmg btvalvcs at that stage. but they
declined thereafter and on.ly one order. the
Class Rostroconchia Conocard10ida, continued unul the Permtan In the
early rostroconchs (e.g. Rtbm1ia) all the shell byers
recent yi.'Jr\ .1 ~mall group of J>:~lacozmc molluscs traverse the dot"ial nurgm. whereas m tbe ,1dvanced
IUs been recogmud ·" bemg of muquc phyloge- toruLS the outer l:tyer doe~ not cros~ Jt, suggestwe of
llr mtere~t .111d ha~ been ~cparatcd out from other an independen t ~tcp toward~ the condition already
mollu!iCS J~ Clas' RostroconchJa (PoJet;J and achieved by hiv:tlvcs. Colloc,rrdill ll l (Fig. R. l 6c,d),
RunnegJr, llJ7 6). one of these adv:tnced fom1s, has a gape at one end
Rostroconchs look superliCJally like bivalves and and a very pro no n nccd rostrum :lt the other.
11ere probably f.1irly ~umbr to them mtemaJly, for lt has been suggested that rostrocon chs occupy a
mmple 111 the possc~sion of a protrmible foot, stg- key p0~1 t1 on in mollmc:tn phylogeny. Pojct:l and
ii~d by,, marked .11lrl'nnr ~ape m the shdJ. Where Run negar ( 197()) have proposed that hclcionellans
:hev d•tfcr i~ in the morphology of the lunge line. are extmcr mon op l:~cophoram and that these gave
r they cio not pm,t·~' .1 functtonal hmge at all. n~e to the rmrroco n ch~. losing their ~egmentatton in
be,e molluscs beg.tn thL'tr ~owrh by producing a the process. The~e in tum produn·d the h1valvc~
mJJI, hmpct-lih. htl.ttl'r,llly \ytnmettical proto- on the o n e hand (by separanon of thl' valves and
:onch. From rim Llll' .tduh. ltkt·wi~e bilaterally development of :1 proper hinge), and po\sibl) the
tmcrrinl ~hell (dissoconc h) grL'W do\>\ n ao; a pair Scaphopoda on rhc orhcr The cL·ph.llopom and gas-
ri l'lk~. But then· " no trul' hm~;c tor ~ome or all tropods, accordmg ro chc,e authol'\, were probably
::l-
e-
d.
)1(
(c) Conocardium
lfi
ey (a) Salpingoscoma
a~
n-
cd
.u-
IVC
(b)
>lS
:!re (e) Helcionella
·n-
'11/te 8. (o), (b) So/pmgostomo (Ord.) o bellerophontid, in dorsol and lateral views(:< 0 5), (c), (d) Conocorc/ium pseudobellum
I6
.nd in lateral and dorsol view (x 1.8); (e) Hekionello, on early Cambrian mollusc (gastropod or monoplacophoran,
l;.e,. ), a rostrocanch,
the 25) (Redrawn from T~trse on Invertebrate Poleanlofogy, Mollusca I, 1960; Potela and Runnegar, 1976 )
222 M o lluscs
denved mdep'-·nJ..:mlv li nm tlw monoplacopborans. pteropods of tht' 111.mrH: pl.ml-ton. ThL ca.rlicsr gen- tor 1
When the ro~tfO\.OtKhs h.1d ~rtVl'n ri~e to the more era arc Lower C.m1bn.1n, and though they probably the
adaptable bl\ a(w,, comp..:tirwn may wcU have been are more abund.lrH now th.111 Jt .my other rime they pbtc
an tmport.lllt ftctm 111 th..:ir d..:misc. can be found m \edtllll'nt.lry ror b of all age,. ~J 1
Thc~e htn~de\' 'bt\'Jh ..:d' molluscs have helped The majorit) of prc,ent-dJ\ g.mropod~. and the it\ Jo
m an uncxpelled w.1~ lO brid~t: a large morpholog- only one-; pre~erved, have l'oilcd shdk Ga.mopod' <tll whic
tcal g.1p m the phylo~cn\ of early molluscs. Perhaps have a true hl·ad, usually c4.uippcd w1th tcntado o( tht
morl' dl\cov'-•nt'' of suutl.u lund will illuminate the eyes and other scnw organs, wluch ~ more or b, tubul
rel:ltlon,htp' of moUll\can classes even further, continuous wtdl the elongated body, tills tvpically lu whicl
allowmg J tc't t 1f tlm t•voluuonary model. a flat, ~ole-hke lower ~urtace upon v. hi eh the arnnul Like
Creep~ by smalJ-scaJe \\JVC~ of JllUSnll,lf l'Olltractious, male
lubncatcd by slime trom mucou s glands. The \1- How<
C la ss Gastropoda cera! part of the body l.1rgely rcsick·~ i1mdc the sbdl. mg P·
which in such ,\n example a~ /JII((illlllll (Fag. H.l7a,b), pc:n"
Introduction and anatomy the common whelk, tS hdic.llly coded. arc sh
Gastropoth in< lude all ~nail\ ;Jnd ~lugs living in the The h ead- fo o t, 1.c. the prolnmble part of tlw the: be
sea, in frc,IJ w.11cr~ and u11 land and also tht' body, c:1n be withdr:~w n in~idc the shell by retrac· .1~ a sti
Tht
COilllll
m eau~
from ,1
(a)
tion 111
howcv
.tlnng t
1ty he t
tzed or
to be tt
Jt IS
CJVlCy
im'-·mal
systt'lll\,
merry, I
pod me
'ome of
nnm1alo
This t
111 now;
of the: si
dXIS
suggc~tic
fusllotno
the mo~
(cl Eugltsh ;
suggc~tet
uve adva
and
Sr.llc
tlw an i rn.
There
.S)'fi'I'S{ftlSJ
number c
Figure 8.17 Gastropod morphology shown by Buccinum (Rec.): (a) living animal ( I approx.), (b) shell (fusiform); (c) trochopllcn nm so rh;
larva, (d) veliger larva of gastropod [(a) Redrawn from Cox in Treatise on Invertebrate Poleonlo/ogy, Port I) hum the
ror muscles (till' on h .m.tehmcnt of the ,oft pan to mass was able to rcrr.:lct deep mto the shdJ. Such
'! e ~hell) aud l lmnl ntl llmn the ouNdc by a monopl.Jcophoram ~' tlm would seem to be
~Jte-like trap donr (tht oper culum). prc-ad:~ptcd for tOr\1011, .:lnd 1t 1s probJble tlut the
The mmuh tont.llll' .1 r 1'p111g J;l\\ (tht• r:tdula) in first gastropods arme from these (Peel. 198U). The
t1IO\n'r p.1rt, compo,t•d of .1 IIIUltltude of tiny teeth great succe~s of g.tsrropods i' ,, tC\t.lment to the value
1\h~eh \crapc .1gaimr .1 hornv pi.Ht' in tht· upper part of tn~ion m .1llowmg full retracnon with m the shell.
1)1 the mouth to ~hrcd h>tld matt·ri.tl. Buu 11111111 has a '\uch a (Jnluy n1.1y havt: been acqmrcd mdepen-
rububr exrcm1un ''' the lllllllth (tht· p r o b oscis} dcntly b~ vJ.nous b'Toups ..md rhu' tht• g.l~tropods
v;luch 1~ pn:~cm 1n tht• more .sdv.mu:d g-a~tmpod~. might be polyphykttc.
lsle mo~t \tl.lll~. tht: "hclk i' hcnu.1phrudJtc, wuh When gastropod egg' h.Jtc h the)- turn m to pl.mk-
uulc .md lcnule org.sm 111 tht· tmt 111dividual. tic l.irv.tc known ,1, troch o pho r es (ftg. R 17c).
lluwevrr, llldlndu.J!, lupul.Jtt' \\ 1th :.1 ~:omplex mat- which do,cly reM:mblc tht: l.lrvat· of n·rram marine
mg pattern, uwolv111~ the l"e of J \er) l.lrgc male wonns. Trochophort'\ ,1re vt•ry ~mall .md more or
rl'lll' ,lnd the l'XpUJ\Ioll of t.lk.m.'lJll\ dJrts, which le" globubr, with a fringe of dli.1 round tht:lr widest
u-e >hot our ot ~peu.1l .lpt•rturt:~ t~nd t•mbeddt·d in part and another tufl .1t the .lpt:x. The mouth, situ-
t~c body of the l"llht'l llldtvJdu.ll. tht~ Jpparently acts ated Jt one sid<.: ju~t below the dli:try nng, leads to a
11 J 1tunulanng prot t:durt·. simple gut temtinatlng in .lll ,IIHI'i .lt the lower pole.
The mantle cavicy lie~ ;Jtltenorly and in H11m1111111 The next stage in Ltrval development i~ the velig er
<)llll\1lll1iC,lfe~ WHh the l'\tt:l n.tl <.:IIVIfOIIlllent by stage (Fig. 8.l7tl), during whiCh ;I thm \hell is
mns of an mhal.liH '1plmn ..1 wbul.u org.m fom1cd secreted over the upper pole .tntl tht• region under
'mm .1 fold 111 rhc m.ltl!lt· ,nd on upym~ an mdeuta- the gut become~ cxp.mded 111to ,\ large sail-like
()D m the ,hell nuq.,'11l. M.1ny g.•,n·opods do not, v elum: a Rat bilobt:d mg.m rovt•rcd wnh cilia
RliVcvcr, h.tv~ ~uch .1 siphon, .llld water 1~ drawn in which propel th e vehger through the WJtcr by then
ng the cdgt· of rht: ~hdl. 'W nh1n the mantle cav- coordjJJated anton. It ~~ .1t tlm \tage th,lt torsion
hc the g~lls, .mm. mucuus gl.mds Jnd .1l~o ~pecul- OCcurs: a rather sudden t\\ !Sling through UUI0 • a~ a
~ organ' rhe osph radia) \\ hme ti.tnction seems result of wh1ch the larg~ velum r.m be withdrawn
lx- to \.1mple tht: \\ stc:r t'lltenng tht· CJ\'ity . U1Side the shell m ca,t· of d.mgc:r (,,Jf\tJng has sug-
h ., channen'nl· of g:t,trnpuds that the mantle gested chat the advantage contl>rrcd 111 bemg abk to
lt) fJcc' .mtl·norly . A"ort.Jtt•d with rhi~ the tuck away the vclum "' :1~ ~o grt:at that tht> rt>~ultant
llml.ll org.m,, mdudm~ tht• nt•rvom and pallial displacement and a~ymmetf) of the nHenuJ org.ms
erm. arc pcnlli.1rly t\Vhtl•tl. This lntt·mal .1~ym wa~ :1 nu nor price to pav for the ~ate()- rendered. (In
.try. known as tof\mn. " ti.md.tmt•nc.tl to g.lstro- addition to hl\ senom paper\ on the subject,
od ruorpholog}. ,md thoul!h 1t hJs been lo~t in G.trstang al<>o wrore about hs<; concept' 111 the
11ne of tht ~bell-le" tt>nll\, tlm return to .1 more unusual medium of com1c verse, .111d l11s poe-m Htlll'
:oo~ul comliuon " de.lrl} ~econdary tile l'el((!cr got rts 'I 111isc cry<;t,lllizc<; the arguments m a
Thi, cunou' dhpl.itt'lllt'nt of tht• mtem.ll organs is memorable f::~shion). The 111.1111 problem th:u the
, no way connt•ctcd \\>lth tht: .twmmt•tnc.tl coiling g.1stropods ::~cquired through ror<;ton was th.lt ex-
I Ull' shdl but h.1s .1 tJllltt' diiK•n:nt M1gin. Many cret.1 from the anus (loc:1tcd in thc m::~ntlc cJvity)
J!!&CStions h.IVl' bet•n propmed to .tnount fi1r it, would be expelled jmt over the mouth. In many
·r most gencr.llly ,ll."lt'ptcd being th,lt nf the gastropods there .m· dcVlcc~ that cope w1th this by
1 ~.,b zoologl'>t Waltt•r Garst.mg (I <JS I), who has ~cparating the inh:~lant nnd exlulant water Exhalant
;gested tb.lt \Urh tnNnll g.tve .1 smgul1r prnt<.:c- <ciphons arc pre~enr in <>ome, and 111 othe~ there IS .m
'~ Jdv.mtagc to thl· ga,trnpod wht·n 111 the lJrval indcm::~tion (slit-band) in the <;hell, fomung a ch.m-
< and w.l\ rt:tamt•d 111 tlw adult wlH:rc it ,11low~ nel through w h ich the cxhakd w,1ter t~ nmed dor-
m1mal to n:trau uHu lt'l ,hdl. sa.ll} away from rhe 'hdl and ht·.1d region (F1g.
There are smnt• n nkd llllllll>plJt·ophor.m,, ~.g. R. \XI).
nrrt•lplltlt"r,l .md \iuuitcs. wh1t h hJve reduced the The vclum of brval g:mropod<> cvcntu,tUy
da of lllll\tk all.lt.hmelll 'ite' on the ,hell i.nrc- becomes the foot upon whKh the adult glides. by
:>hore so that the tiN mu,cJt• \t'af\ ht: hali a whorl bark tht· loss of ciha and rhc development of an JOtt'mal
;:n the ~pecturc . Tlm 'uggest\ that the vi'reral ~~tem of longtmdm.ll mmcks which .1llow vc:ry
224 Molluscs
small rhyd1m1t \\,1\'l"\ ot nmrractlon to pass back- ORDER 1 ARCHAEOGASTROPODA (I CJm.- R,<.). Th•
ward' along the foot; the luhncanon of the gastro- gills here arc aspidobranch (1.e. hJw theu til.unrnb
pod's passage 1\ .h~l\ted h> the mpply of c~pwus arr.lll!!l'd 111 a douhll- comb nn either ~ldl of tlw ~Xh ;M
shme from the muc.:ou' giJnd'. arc free .lt one ~nd) . rlu·rc mJy he t\H> g1lk or one If
be lmt The 'hell ~tnKtur< I< v.m.1ble. 'ome \}'11Unemc:a!
G.1srropod~ nuv Iced 111 a van et\ of wavs. Some.
fonn> c:xm llr Ut' found J.\ ,,,,,11,, bur the ,hc1J, .1re nor-
such .1~ the muriod' .~re acuvdy <:J~mvorot~~ and can lmllv heh,.ll <plrl"> Nearh JJJ nunne; c g . £uc>lllp/UtiKJ
rasp away the ~hdlof .lllother g.t,tropod or a bwalve P/cumlcllllcln•l P.lltflcl, l'lolly(tf.JS. rr.,.:/ms, .\I.Jdunt<'S
ro re.1ch the flc<.h fhc > drill a neat, round hole in so ORDER 2 MESOGASTROPODA (Ord.-Rcc.): ,\1\Jml) luvr
domg. \\ htdl may t.1kc 24-4H h. They then inject a pectinibranch gill\..1 mud1 mor~ l'LlborJI<" md cffiClCDl
muscle relaxant through the nC:\\1) drillcd hole, the \)'~tem rhan the ,1\f'IU<IbrJnch lOildltlOil. \\lu.:h perm
~hell then open\ and they .tre able to feed. The old- free: flow ot wJter through the lliJnt.lc uvtt\ Ln'UIE
mNog;mropoJ, .m: ci.I\Sihed on rJdu1Jr <ttul'turc ; er,
est known prc..·d.1W11al g.l~tropoJ drillholcs arc prob-
Stlc>m/lw, Cn>rar.l, N.uic,l. Ct•nt/1111111, 1\enncc~.
ably Dcvormn (South et ,d., 1985) and drillmg ORDER 3. NEOGASTROPODA (C'rct Rec.) The m~u,.
hab1t' have .1lso bet·n recorded 111 the Carboniferous and ncoga\tropod~ .He otientomlm1ed 111 tlw •mglt• Ordd
Plntyct'ms, a spec..1111cn of wluch was found located Cacno~J~tropod.1 fhcy luvc rectmibrJtKh gtlb. "n
directly .1bove .1 COillcal hole m the tegmen of a in.halmt :01phon leads m to the m.1r1tle c:tvlty , c:har;llteruu-
crinoid. Tn th1s case, however, the gastropod was cally with .1 ;hort or long j,."l'oove 111 the shell carrying,
probably p.u-.1~it1c r.)[hcr than pred,ttori;ll. lt was not, siphonal mbc antcm1rly from the ~hell: e.g. .lfurn.
Brmi1111111, I 'lllllltl. Cclllll.l.
howev<:r, until tlH· Lower Cretaceous that ~uch
SUBCLASS 2. OPISTHOBRANCHIATA (?Carb -Rec.). Mannc
drilling became Wlde~preac.l. and 'forced' evasive gastrl1pods which h.1vc br~cly tlr complcteh ln~t eh.- sheU
adapt.ltlom on tht: prey. Today ,\!ttrt'x and related They luvc umlc:rgom· c.ll'lOI"IOil Jnd str.lJghrcncd them,dvn
gcnerJ arc th~: most dlenivc..· of all the 'driller ou[. The~e indtiUl' the planktOill( ptcropod' and nud1hranch
killt:r\ ·, as thl'} h IVl' b ..•en memorably tenned. ~eJ 1lut;t'. wh1rh rury \l'nmd~l) gills ull the dorsal surf.1c~.
Turrirdlnw ~.1\tmpod~ have been extensivdy SUBCLASS 3 PUIMONATA (Ml'~. Re, ): In tbc~c l~nd
dwellmg {.tntl set· ond.lrtlv 11 c\11\\ .rter-d\\ c:ll.ing) 'IuS' w
drilled and 'pt•dt•tf (i.t•. have had rheir apertures
m;ul;, rhe wn~ .m· lost Jlld thl whole ) \ld:ll'l' nf the IJ1.IIIdc
broh n} br other gamopods from the Late canry IS mod1fied ·" .1 luu~. l1ber.tll~ supphcd w1th blood'
Crcra.:cou' until the pn•,cnt d.1v. Y er such prec.Ution VNscls and kept JX m1anently llllll\l The pulmonart·, ;m- rhc
ha~ nor app.1rently ii.)rCl'd .m} evoluoonary changes onh mollusn rh at h.tvt· m.1dc:o .1 rt·Jih sm rn,fi.JI rrans10un tu
in tht shell~ of tumtdlim·, dunng th1~ rime: there is Lllld. Mo~t of them h.IVl' rct.llned ~ht·1r shells. though tht
no cvidelllc of .m ·.1rn1' race· a~d predation seems land )Ju~ h.11.c: lo~t them altogether.
to have staved .1t much tht· '>.llllC level smce the later
MesozoK (AUmon t'l <1/ .. 1990}. Shell ~trtlcturc and morpholob'Y, which arc ~
Many g,L\tft)pod' fi.•cd on dt:tntus or decayed that 1s ldi to the palaeonrolngi~t, arc not generJ.li~
matcnal. Snll otlwf\ h.lVt' 'pcnahzed modes of teed- h1gh on the hst of cnrcria fur dctc:rmining com·c•
in g. such <1\ thl wpropluhc (facccs-mgc~ting) gas- sysrematic pl.~ecmcnt. Furthcm10rc, the common
tropods found clu\tcreJ round the exhalant spires of tendency tor shells of unrclalcd stock~ to acqmr~ Ul
CarborutcrOLI\ bla,to1ck similar fonm through ho111comorphic cvoluuon

Figure'
causes addit10n.ll problems. lar, lov.
Classification folded r
Shell structure ond morphology constric
The clamtication of gJ.,tropods is lar~dy based upon Glabra.
soft parrs. Gill .1nd mphrad1al morphology is most T he shell of .1 gastropod is ba~ic.tlly an elongated Symma
1mporcant, a' 1' the \trudun· nf the nervous system, cone, rare ly sept.lte, which 111.ty bl· coiled in anum- Bntish f.
heart, kidneys and tcproductJve ')'Stem. A con- ber of way~. Tht.• mo~t charac..tct istil ~hell shape 11
densed vers1on of th.ll m thl' Tre,ztl:il' 011 bwerrebrare helical sp1re, coded aboul an .1..-...i,, wluch i~ U\U.illj lll.ly b
PnleMttolt!I?Y I> g~vc..·n ha~. though more reccm smd- illu~rrated as vcmcal. 1 hcrc arc ~ome gastropod!
llllll' lll
ies (Ponder and W.ltlll, llJHH. Ha<;zprunar, 1988) with planisptral and hence 'Yllilllctrica.l shell' try of
pre~em a somewh.lt dttft:rent arrangement. u~ually the whorls aU touch one another, but m
of ga~t
some cases they do not embrace. Cmlmg i' charac· ,elf-ex
SUBCLASS 1 PROSOBRANCHIATA (L Cam -Rec): ShcUcd tcmtically dextral bur may be ~imstr.tl; in very rm iJJtJ\tTJ
g;l~tropml, m \\ hJC h h .> Nun " complete:. ca~cs equal numbt:rs of iudiviJuab in a population
lt lS
'11~ (c) Ampullonatica

'Ill
lit~
(d) Athleta
ng
.g.
(a) Acteonella (b) Tumtella
,.,.
l~r
11
(jJ Vermetus (k) Tibia

(e) Murchisonia
(f) Nerinea
(h) Gastrocopta
selenizone
.tll slit
111)' band
ect
(I) Glabrocingulum (m) Euompha/us (n) SymmetrocapultJs
";Jre818 Gastropod shell shapes: (o) Acfeone//a (Rec ), convolute, (b) Turrite/la (Eoc.), turreted; (c) Ampullonafica (Eoc.), globu-
:11", low spired; (d) Athleto (Eoc.), volutospine; (e) Murchisonia (Carb.), high spired; fn Nerineo (Jur.), with columella and internal
'Oided thickenings within the whorls; (g) Phyl/ocheilus (Jur.), turbinate, with digitate oper11.Jre; (h) Gostrocopto (Rec ), pupiform, with
~s1ricled aperture; (j) Vermefus (Rec.) irregular, partially uncailed, (k) Tibia (Eoc.), turbinate to high-spired, siphonote; (I)
~"Jobrocingulum (Corb ), showing slit bond for exhalant siphon, (m) Euompholus (Carb.), d1scoidol, near plonispirol; (n)
Symmetrocapulus (Jur.). potellote (Mainly redrawn from Treatise on Invertebrate Pafeontology, Part I, and BritJSh Cenozoic Fossils,
n1- hloh Museum, 1979, not to scale)
is a
llly euy be dextral ,md 'llmtr<!l. Ltmk} ( 1977) ha~ doc- of Cl'rt,un g.tstropod~ (the prmoconch) .trt' ofTen of
od~ um~ntcd some uHcrc~tmg comment\ on the gt•omc- pt·culiar form and do not netC\\arily cotl in the ~amc
~us. ay of g;tstropud shdl loru1. The b,1sic rcrnunology ,1)1.1~ <l\ rhc rest of the ,lwll: t"urthcrmorc, tht·y 11\llally
111 ~mropod shell\ '' gt vm iu Fig. R 17b It ts largely h.lVc ,, cittTcrcm surC~ee orn.nncnt.
'aC- lt:.t•xplanawry, ,1, .m: 'omc oC the b.t\ll" 'hdl fom1s fhc cll."temal whorb may be ornamt·ntt•d wirh
·are 'rrlred 111 Fig. H. IH. ,·anom characrensut fc.1turc~: nbs. spine' or vcrlical
100 lr h worth notut~ thac the carlic~tJll\"l'tllk \.\ horl, bal' (varices). The Jpaturc Ill.\)' bt• cnnrc .md
226 Molluscs
unnurkcd by any feature (holostomatous) or. a~ in Evolution g;.~tro
neo~.l~tropod genera. ma\ bt c.:qutppcd with a General features of evolution ol the.
groove which is the ourlt•t fm tht c.: xhalant s1phon G.l\tropod'\ mmr have ari\cn tTom .1 btl.trcr.ill} }) 111ullu
(siphonostomatous). The..> s1phon nuy be set in a mctnral mollmcan ance~ror. Jpproxnnatm~ 111 !!.Cil- So(
deep ~ht-b:md \VhKh wparatt·~ 1t t'Jr from tht· er,tl morphology ro the hypmhetJc.tl .trchunoll hi\{'·
mourh. The cllcitieJ ~tnp u:prt·~~.·nting the 'track' nf dt:~t·ribl·J t'otrlicr. The protection gtvcn by tnl'h (Jp-\1
the ~lit-band a~ it i~ calntied J~ tlw sclcnizonc (Fig. and the resultant ability ro 'vVithdraw instdc thcu tmidc.!
H.l HI). In ~uch genera a~ till' sJnd-dwdlmg Aporrltais ~hdh mu~l have given gastropod~ .1 ~rt•;tr
::t.dvJnt ~ gllltl' t
.md Plryllocltcdus rhc apl'rttlle m.ty bt· enlarged into a K)r from their fir~t .tppl'ar<HKl' in rhe Low radter
Range wh1ch helps ro ~ubilize thl:' 'hell while the CambriJ.n the} !>cem to havt' bel'll rcmJrbhl} u alh !I
~astropods arc feeding. CC\sful. lwlls
There ts :1 ven .. le.tr dt,tllld!Oll between tho.,c.: Thl' earliest known ptmtbk· ~unph:
sheU' in which the late-r whorl~ do not meet cen- (Cc.lW'.'Jiim. Hclcronclla) arc found m 111011 l
trally•.md rhm have .111 umbilic us. and those in Cambn.m. They have coiled 'hell' wl(h :1 \\"~ f';al.wo
whith rhey touch. In the lattet o'e they m:: welded expanded aperture and an: platmptr,Jl, hence bthtl!· In
to .t centr:ll rod (the columella). Likewise, ally symmetrical. Thest· very early gl·ncra an~ JW· ,t,yllln
high-\plrcd fonm, in which the .1pical angle is low, mally tla~sifit:d with the early o~rchacoga,truro !O tl
arc very di.~rinn from low-spm:d genera, which Suburder 13ellcrophontacca (L. Cnn.-Tria~; FJ& l'lt•uro
have :t higher apic.1l 01ngle. Tht• ~hapc of the whorl~. 8.16a.b) named after the characteri~tic genu> M exdm1
::my unusual coiling p.ltterns. the.: presence or la<Jplwn (which was the fi~t Palaco70lt mllllu'r mr thC\t' •
ab~encc of dttferent kind~ of t•xtcrnal ornament, to bt• UC'\l ribcd. by the concholog1st de Monttor. rorn.1n
apcnural ~hape. Jnd the pn:,ence or absence of in 1808). (The systemanc pmmon. howC'vcr, M~.uu:
sht-b:mds or of stphonal gTtlOVes .ue all used 111 fanuhcs SmumdJe and Bclkrophonud.1c lm bee pod g
dJ~~ttic:trion :t..nd idenntic.ltlon. much Jcb.m·J. Spccimem of &·llt'lllpholl \omen~ lntt·rn.
~how paired muscle scars and thu\ tlm genu' and \1111110(
Shell composition .illie~ arc con~idcred by ~ome authorit1c~ more bke org.lll\
The ~hell~ of g.1stropods h.1n· an outer homy layer. to bl· monoplacophoram than g.l\tmpods.) !lOll, S
the pcriostracum, below wluc b ltt·~ the shell proper, lklkrophontidid~ have been lll ll'rprctcJ (HJrp,r nn t rn
J ~tntcturc normally compost•d of layers of arago- ,1J1d R.ollim, I 985) as infauna l m senti-int:tunal Ill • tht wa
nitl', though many g.Htrop0tk t•spt·n.illy those hv- I uses with thl· apertural rq,rion enwrapped by tlh- lcrttlt7:
tng mtl'rtid.llly. luvc .1 c.tlottt outt'r byer. In tom! mantle and possibly al~u the toot. One lc1m1 Ilk ,,frhc I
\hl'll\ the aragonite either rco-y't.1ll1ze\ to c.tkitc or S1lunan PtcrMhccct trimcrc/loirles (Ciark.,on r/ ,,/., 1995 Br,
m.1y be..> dis,olvcd. c'ren.tlly whert' there is leJchmg ~how\ an extraordinary rangt· of form wltlnn \t'T)' ne
111 th~ rock . one spenc~. Possibly tlm reprl'scnt' J \UTYI\'al 'll'E- lllgly c:
M.tny ,JtcJl, hJvc .m inner n.tcrcom layer of egy, prc~encing to intendmg prt·datul"\, I.''Pt'CU:: OC'<llplo
very rhin Jra~omtc lean·, par.tlltl wtth the internal t.ephaJopods, a mosaic of dlf1"erent ·~c.lrfh lll1J~~ ot rht• 1
'hdl 'urCtcc. each 'eparated by t'qllally thm org.mic .1nd thcrt·b}' confusing 1t. { :.1rbn1
layt•r,, External to tlm 1~ the: cro~St'd-bmcllar layer The bdlcrophontaceans .1re ,, 1110\l iruport<ll I ll11l'~CI
in ·which thin lamellae (0.02-!UH mm thick) are group of fossil gastropods, and over 70 valid gcni'D r·nglan•
Jrr;tngcd nonn:tl to the.:- .;hell mrf.1cc. But each h.we been described. Howl'vt·r. \Olllt' ot t Hit ~pc
l.uml1.1 ttsdf ~.onmt~ of .;tnps of extremely thin Cambrian bdlerophontitorn1 ,!Jell\ luvc ptl'UIJJ!I- I O\\ ~r
.Jr.tgomtt•. ,1rranged obhqucly, the 'tnp' of :tdjacent ucs uncharacteristic of the bdlerophontan•Jns .ti (B.mcn
brndl.tc bc..>ing arrangt•J .tt nght .1ngle~ to one ''hole (Yochelson. 1967). Hc/1 tr11r//t1 and it' rdlm re haec
.1notlwr. I1Jn' a sunplc apenure wtthout .Ill) indcnuuont :~nd lu
There .ue otht·r kind, 1)f .;hdl stmcntre in ga~ but Jll other bcllcrophontactJIIS have .1 pronoun C.ll'llO~,
tmpmh, .ill of 'Y'tcuuric unport.tnce. bur••ls m the note h or enurgmacion 111 the plant of 'ymmem <.m be 1
C:l\e of the 1.1) ~rs dc'l nhl·d ,tboYc, they .1hnosr Tht•re .trc other reasons for believmg that Hrlac'N nf the I
alw.1y~ d1~.tppcar in fn,~JI\ .1nd no trace of them is not a normJI bellerophontJteJn, and ~on In rllltr•
rt•ntatn\. authorit1c' have suggested that 1t nMy not ~ 1 her
havtng
tNrnpoJ Jt all but •·.nhl'r 111.1~ bl' .1 n•pr~l'ntativl' Carbom£t:rous: '' first mv.1~ion of tht• habitat ~o
ri the monoplacophor.m~ or l'f ,111 exnnct das~ of successfully coloruzcd by the pulmon.Jtes much latl'r
ollu~c-~. on in the Juramc and Crctae<.'OU\.
~omc other P.tl:il'Ol\llt" bdlerophonufonn mol- The gastropods were atfectl'd, as wt•re mmt otl1l'r
), tht· u~htl} cmkd C)•if,,, )'rttllldftl and the more orgamstm, by the !,tTcat ~xtmcuon pt.•nod at the c:nd
r-<hJpl'd '/ rybfldww, h.IVl' p:urt'd mmde ~cars of th~ Penman, but they tontJnut•d to t."volve
r !de the ~hell .111J dt' not .tppl'ar to hJve under- throughout the Me,ozo•c Man) duraaeristic
.'lll e tOI"'JOn; rhc1r mmde ,c.1r p.Htem mdtcate~ that, groups ofMe.,OlOJc age bct.um· \Cl) 1mportaJlt for a
r :her tlun bl'ing bdlerophont.Ke.llls. the} ue Jctu- while. such as the Nenne1d.te (F1g. H.lHO. The~e are
~r rnonoplacophor.m~ . llencl' bdlerophonnfonn a 6mily of lugh-spired Mesozorc me~og.tstropods
dh .1re not necess.111ly .tU ~ra,tropods. but such fou~d in carbonate scdunents: 111 them th~ mstdt." of
1nnple morpholo~ r.tthl'r \l.'t'lllS to hJvt· bt'c:u com- the sptrc is duckcn~d by foldl'd caloum carbonat~.
mon m dive~e molhlstau ~rroups dutmg the earlier Spiral calnte rods ru1uun~ wtt!un the spire nuy bl'
l' P.U.ICOZOll the original duct systc:m wrtlun the digestive gland
In the Upper C.unbn:m ,1re t'ound the ftrst (Barker. 1990). lt Ius bet"n argued that many ner-
,yuunetnlal, hdtc.1lh cotled ~hell,, belonging ineids were int:,unal and th,lt ~•nee they lived in
to the 1mport.111t archacoga~tropod Suborder organic-nch carbon~lte mud, on which they ted.
g. fleurotonl.lnlll.t (C.1m - K..t"c.). ' l here is .tlso an they would not have needed the nutrient storage
,:1- rxd1Nvdy )J,,)aeozotc ~ubordt."r, the Macluritina: all units that other gastropods possessed in the spire.
'le\e earlv form~ arc low-spuc.'d. Rc:ccm pleuro- Hence the space was taken tlp instead by calcite
•omJnidcs are 'hvmg to"th · 111 whtch th~ grade of which followed the contours of th~ digestive gland
pruzanon present 111 '0llll' of the e.ulil'st gastro- and gonad. The calctte taken m wtth tht· food was
X~J gener.1 c.m bl' \een by d~rect homology. thus convemently d1sposed of Other groups. such as
lcmally the gill~ of modem pleurotom.uiids are the gtgantic neoga~tropod Famtly Srrombtdae and
.::uuodttied (.hpldohrJnch). The reproductive the cowne-shell Fatmly Cyprae1dae. appeared for
~m are Jjk~WN~ Ill a rd:ltlvdy pnnutlve conru- the tirst nme m the later Mesozote. But tht." real
~. smce the rap.Kll)" ti.)r mternJl fl'rnhzation ts acmt." of ga.stropod evoluuon was reached m the
per t presenr Jnd eggs .1nd 'penu are merely shed mto Ternary, conru1tung unnl the present. wtth the
ul- e \\,tter lt \\,"J' only tlw tkveh1pment of intern.1l great succ~ss of the long-~tphnned neogJsrropod'
thc m~hzJnon th.lt rentkrl·d pn,,tble tht." ):Iter invas1on (e.g. Fig. ~ . I ~k) wh1ch domuwc roda} ·~ gastropod
th\! the bnd .md frt·,h w.tter,, t3una.
95) lly c~rbomti..·rou' tlllll'\ ~.1\tropod tJunas were Gastropod~ ~ecm to h.1w bct•n a \table and con-
the >try ncb and dn:t'r.t·: rndt•t·d 1t 1' ht•conung increas- stant component of the m:rnnt> f.ltlll.l smce e.trly
rat- ,11 clear that l':tbet1ZOil" ~·''tropod' ~eem to have times, and may domuutc ~ome m.mne commum-
.allv L1tpred .1 r.mg<.' of h .tb trat~ .l pproxun.tnng to those ties. Fre~hwater Ternary hme~tone' may be
~e<~', :the prl'~l·nt day In one taunJ of tindy prcs~rved compmcd cnttrcly of g.l~tropods, and a particular
Jrbomfcrou~ ga,tropod\. the: Vts~.m Hotwdls long-ranged ::ts~ocr.ttton of gastropods and ostra-
tant Lunc~tont: fJl111:l of the: M~lllllp I hlls m Somcr<;ct, cod~s. mually .d~o with bivalve\, i~ often an indiCa-
11era Engl.md, no le'' tlun 4S gt•m·ra :tnd upwards of tor ofbracki~h-l::tgoonal facics.
th~ ·" ,pcnc' occur 111 ;l\\0Cl.'ltJOn w1th a typical Though g.1~tropods .1rc long-rnnged .md evolved
iari- lower Carbon&•rou~ coraJ-hr.lrhlOpod .1s~oc1ation ~lowly, new stmctural developments allow~d
as a Bant•n. 196(,). In tlm f.Ju11.1 tht:r~ arc many important Jdvance~ at dttterem nmcs in g~ological
tive~ hac:-ogNropod' (hdlt:rophonud,, pkurotomariids history. The modem vcnncnds, for m stance. arc
ton~. md lnnpN-hh· gl'm•r.l). hut thcrt.· arc some ga.~tropods with pentharly tllltOtkd shell~ which
nc~d nogNropod, ·" wdl; the .mcemy of the latter may 1ivc permanently att.lchl'd to branchmg corals
etn . :m be tral ( d b.JCk tn the t )rdonoan All the ~pent:s and fl'ed by strammg otT food p.ut1cles from water
nu·lltl IDl' Hotwdh Lmlt'\tollt' wen: .1pparently adapted passed between the edge of the shell and the oper-
,orne ourronrrh~, wttlun the1r t• nvirnnmt·nt. culum. In Vmucws (Ftg. 8. 18j) and Tcnnrwl<ma.
be a There " ~ome cndt•ncc of c.tcnog;tstropods wh1ch arc typical examples. two adv:lntage\ scl'm ro
nng nugr.ued mm .1 non-marine hJbHat by the be gwen by such uncoihng (Could, 1969b). One ts
228 Molluscs
raptd upgn''' rh tov.:;trd~ the..· 'ourcc of food p.lrticlc~ l'.l~
r.nnmg dn\\'11 ~' dctnru' front thL' 'urf3rc. The othL•r west zonatus east 1onau<. l \t
'" the cnmtclc..•r t!-1k· fkx1hthty rc..•ndt•rcd po~o;1blc to l.tst
mdJVJdu,il' \\ IH n gnm mg round ob,tadc..·'· Co
There arc sonh pccul1.1r Dc:voniJn ro Tna•;<;ic fw
< ountap.trl\ nf the, c..• , c'pcrially .1bundanr in the IIIJ
C.1rbn111fi: rnu : the..• \n-c.tlkd 'venniti1nn g.l\- oth
rropml,'. The..·} tc..:nd w t"lmn dcmdy packt-d bio- l'.lt'
henm, nftcn a c.. comp.minl bv calcareous alg<tc, ufc

e-.pcci,t!ly 111 n :,tnttnl \h,tllm\ -lagoonal habitat~. .md
The.. ,t•, hm' l'Ver. .tn• not .Ktu,tlly gao;tropod' role
(Wc..ednn , JIJ 110) . f h~o.') h,l\'l' .lll initial coiled buJ- I
hou' prnto< nnch .llld .1 wuque rhree-bycred shdl plm
~trudurc... It '' pl)\~Jblc on the basis of shd1 \V Ill
lllJLTO\lrm Lure.. th.lt tltc} JfL' rcbtt·d to the tenuculi- togt

tld' (q v.), .111d Wc..·cdou regards the two as a sister I p.trt
I quit
group nl .dl otllL't mollmn, h.tvmg branched off
l '
cJrlv in lllllc.. -','--~------ 1 pcri
In c..,lrl) lcrti.1ry time..·~ there .tppeared the ',', 1
ptempods. wllll·h .1re \111.111 (< . 2 cm) plankronic Figure 8. 19 Evolution of Poecilizonites bennudensis by tlero
wht·
opl\thnbrandl\, Thc..·\t' tiiJY h;wc dun shells wh1ch live development of poeclomorphic subspecies in two seporol! .Jr Lt
c;m he tolled or \IJ,Iight: .IItern.ltivdv they may have oreos. (Redrawn from Gould, 1969o ) be u
no shell\ .lt .111 . I ht') 'Jlt'nd all thetr hves afloat and \);
;ue unpon.mr complllll.'nts of the: plankton. It ~~ moll
ofic..·n \u~gc,trd th.u the ptcropod\ had a paedomor- small compa~~ of rhe tsol.ned l.:tke tlm may lu~ fur~
phtc origm twm fi(1,1tlll g \'ehgn LirVae, wluch seems been po~"blc wJrlun different local habitats. men
an cmtm•ntl) n'.l~ll11.1ble proposmon. Their primary A now-d.:t<;stc ~rud) " that of Gould (I <JoYa; F~r pr<>V
gcolo~l<\11 mlporr.Jmc..· 1' ,1, ont' of the main compo- ~.19), who eluctdarc..·d the cvoluttona11 histol} r1 strati
ncnb of prernpod nozt· 111 the dccp oceam. the Plct~toccnc pulmon.lte ga,tropod g~nlll gastn
Till' t·.uht•,t .luthentll'.Ht•d ptcropodo; came !Tom Pomlo::ollltl'.\ whtch lived 111 Bem1udJ dunng the lr.t lo~:orit
the Et"~l"t'lll' : g~·m·r.t de,rnbc..·d ,\, 'pteropod~· from 3tH) 000 years of the..· Plctmxenc: one ~ub,pecl~ 11 Will!
the C.unbri.m .md 1lthn '}"'tt•ms .1rc now known to hvmg rod;ly. Jut101
be 'hell' of hyohthid,, \\ h1ch .1re extmcr animals of Fms1l ~hell\ of rim br:t'tropod occur abund.mtly Ill lm v
unknown .tt1irmic..·s. unrd.ttcd to ~·1\tropod~. a st·qucnce of :~lrnn.ltlllg reddl\h smls and wmd- 'puJH
blown sand, whtch Gould carefully doe um~meJ. ol gl'
Microevolution in gastropods The \hordmes moll::tted through the l.!rcr With
C;mropc1th .m· of ltn111cd v.1luc 111 \tr.ltigraphy but Plci~rocene, ~o th~t i~l:md~ on which the gastropod see me
h.1vc.. pnwc..·d 111 bt· unport.un in microcvolutionJt)' lived were altcrn~tely p.1rti.tlly ~ubmcrged and 1111· rq,>Tc·
\tudlt''· ·r he..· t',lrhc't of the~c. b) Hilgendorf ( 1863: latcd, .lnd l'xpmccl and hence linked, though ottcL popul
SCl' Retf, I 1JH 1) llllll"l'nll'd popu!Jtinm of rhc rresh- in different cnmbm.nion~. In his study of the chan~· show,
\\',ltl'r pl.111mlnd \11.111 ( :yr,wht.\ klcmi which became ing popul.mons rhr()u~hout tlm time:. Gould founa cxtn.·•
1\ol.ncd 111 tlw J km th.lml'tt't metcor-impact crJtcr that tWO popul.Jtiom of Jl. ::o11nf11s bcrmlldnt.tiS, di~ ofsut
l.lkc..· of ~tc..·mlh' llll in 'ourhem Gcnn.my The popu- tinguishcd pnmanly by colour bandmg. liwJ tn .1ted
I.Hillll\ h~.·n: \\ c..Tt' .td.tprt•d to .1lkaline condiriom. ca\tcm .md Wl'\tt•rn n.·~:--.,om .md were prc~urml \ Altho
Thl're .Ire dc.tr l'\'ulutionJJ)' ch.m~t'\ here. e\pc- iso!Jrcd thmughoul mmt or .111 of the nme. Tho. .1~ an
l't.lll) ,1 g~o.'lll'J~il eh mgc..· ti·orn flattened to high-'>pireJ e.lstem and WC\tl'm J> ::<>llcllll.\ popubtiom formed docur
morphulo~') 111 dw nuddk part of the 'ucceNon, the parent srot ks (rom which other 'ub~pccic\ \W (Iones
and l.tter ,1 re\'L'r-.tl to .1 lm\ _,pi red fonn agam denvcd. Thcre .m t(mr dni\'cd sub\pccin: o c' ent\
t\1\\ ,trd' tlw wp. lt " not e.1w tn e't.tbh'h \vherher ont,'lnJting from thl· wc..·,tc..•m P. ::t~uatlls populabor., tonn'
or 1111t 'Jll'll.ltic 111 ",1, .tllopalrtl, tiiiJugh l'\'el1 in rhc rhc orhcr three from the c,l,tem brroup. The th~ 511 ()()(
tc·m fonm .tppt'.trt·d \Ut'Ct'\\lvt'ly. t•tch b~:coming \tability. and this mt·am a~ m.llly as ::!11 0110 gener.l-
tJIICI bdort• tl1c..' .lppt•,Jr:mrt• of thl' tH.'Xt, and the nom lwtwt·cn \t:Jhk '>J'l'lll''· Tn Jnnt·~ Wt' owe tlw
t-produu.•J \lllhiWlll'\ 1\ tht• 1111t' stt!l hvmg roday. comment th.1t 'Ocpt·ndin~ upon tlw tlmt• ~c.tle to
{j(luld \\',1, .1ble to ,ho\\' th.tr .111 fc.mr 'ub,pt•t ic' 1rose which the invcmg.nor is .Kcu,tomed. nnt• m.m \
om 1\ol.Jtcd ~m.tll group~ on tht• pt•ripht·n of the punctu.ued CtJUtlihrium nuy lw mother\ en"'lu-
m.un pnpul.llion .wd thcntt' 'prctd to colonize rionary graduah\m'. Tht, i' surdy a v1hd l'nmnK'lH
t,llhcr rcgwm: .1 d,t,,ic t' ,l\t' of.tllop.1tnc \PCCt,uion in m consideratiOn not only nf ~stropod C\ ,l[Uttlln
h ttNanu:. He..• .tho tlt.tdt• it dc.1r that the origin specifically. but of cvulution.1ry phenomcn1 111
e.uh popul.ttinn w.a, the..· rc,ult of .1 paedomorphic general.
tnd hl'llt:( imtanr.ult'Oll\ l'hangc, l.'mpha_,iztng th At th1~ point, \\C should mentum th <: tt'llt.llllli-
, J' Jn unport,lltt nmtml of t'vulutton. totd\ (Ord.-1kv.) ..111 t'nlb'111Jttl ~:;roup ,,f t nniul.
H.1d the \lr.tti~rr.tplncul. ~~o.'Oj.,'T'.lphtt,tl .md mor- dun cakite ,hdl-.. 1- 3 l m ltln~. with 'tnkingly
hol,Jgical dmun~<:nt.ltion bcc.:n le" co111plcte. it anmd.ttcd l'Xtenot \Urtltt '· Thl'\" 111.1y nr m.ty not
,,u)d have been vcrs c.tw .111d tl'mpting to fit bt· mollu\c~. T ht·se h.tvt· bnn l''-lt'n'i-.cly rcsc.lrcht•d
:tJb>tthrr the three e,l\ttTn P . .<:ulhlfiiS ~ub,pecie~ a~ by Lardeu' ( 1969) .mJ I ,lr"tln ( 197')) .md h,t\'c
~m of,, \IU~k evolvtng pkxm r<tthcr dun as three proved of indubit:tbll.' 'itr.ttigr,tplm 'v.lillt'. Thctr ,oft
~ane dJstllln I let anvc popul.tttnm whtdt started as parts and lif<.· habiLs ll.'lll.1in unktwwtL
.'.nphrr;~lt~ol.ue~.
fhcrdorc, in onk1 Ill undtr,t.tnd eh~: pnnnples
1hncb\ popul.mon ditl(:n:ntt.1tlon t,tke~ pl.u:c. it is Class Cephalopoda
D
le 1thi, level of dt·uikd ~maly''' th.n \tudie\ have to
undert:lktn , The cephaJopod,, wh1ch .Jre t•ntirely mmn~:. ,tre the
\\ tlltarmon ( I9H 1) 'tudic..·d .1 \l'rics of lacusrrine most highly evoiVt·d of itll mollu~c~ W •thin tlm cJ..,,
llU\C'all f:lllllJ\ in the l.ttl' [ crt1.1~ of the eastent are included the modt·rn ,\nu11d11 thl' :ugon.lllt\,
I • urbua UJsin in Kcny .t. Tht· 41lll 111 of lake ~edi sqmd' and ocropthC\, .llld till' t'Xtinrt mu11onmd"
nt are punctu.tteti br nt:lpp.tblc tutl\ "hi eh and bekmnHt~. All modem l t'ph.llopod' are dl'itm-
rol'ed to be: u,c..·ll.tl 111 llrclc..•rmK the flUn~ m gui,heJ h~ luving .1 prnperl) devdnped he.1d wtth a
tl~phtc ~cque nce. ' I bt• 'tudy of bt\·alve~ and good brain .llld cl,1hor.ltt' 'emory org.1m: the ~rrur
ttopoJ, pro\'ldcd chc fiN tine-,ealed palaeonto- tural and func..tinn.tl p.,r.dlcl' between the C\'t.'' <)f
:lCJI rc,olurion of \Pl'l iatll>n cverlts so that cephalopods and those..· of vcrtebrHl''· wluch trc so
~ illianhon wa~ able ro C\t.tbh,h tht• nature of evo- well known. \en·e m t!Ju,tr.ttc· thl' pn"thlhttes of
J:Ilon.tl') phcnoam:n.1 111 tlll k~s that 14 lineages. In evoluttotury attainment inht•rt·nt 111 the mnllmr.tn
11\ 1 n ew .111 the spcctJtton t'\'Cilt\ continned the archetyp•ll plan
d- un.nt.ltcd' etltuhbrium tnmld, 111 the pe~pectn.e lt m:ly seem n:m.1rk:thlt· th.lt the h1ghlv mob1le
•d. l geolog-.c.1l tune. lllll!); penod) of \CJ\IS alternated cephalopod~ .tre comrntned upon the 'ame ba,Jc
1th ,hort bursts of r.1p1d 'pcu.Hton. M.ljor changes pbn that is found .11\o 111 the hc.ldk-.<. .md mainly
"t'mt'd to occm 111 penpheral 1solates dunng bemhic b1vaiVl's. Yet as f1g. H. I shmv,, ,Ill rhe com-
.!!fC~'i on s of thl l.tkc Dunng these ttmes the ponents of 1he arthc..·ryp.ll nmllmc art prc~enl 111 the
~pularions. bcang 1\0I.Ited ,md under stress. ccphalopods .1s they .m· 111 other molltt~cs_ It i'
g- illllwed comtderahlc phennt) pie variat1on refl ecting largely because the teph.llopods wne .1bk. t·.1rly 111
nd ·xrrtme development.1l tnstabthty. I rom members their evoluoon.try history, Ln dt·velop an d1ecttve
1such popul.ltlom nt•w 'Petit'S .1rc..1se .md prolifer- meam of buoy.tnry ming the rh.11nhered \hell that
red dunng subseqm•nt l.1nmnne tr.111~gressious. they were able ro free thctmclve' trnm the sea Aoor.
i:though the TurkJn.1 ~cqul.'tKC ha~ been accepted and colonize tlw ncktit lub1ta1, wtth at' nch tcJOd
m excellent cx.1111ple of prt·n~e pal,1eontological rt:sourccs of actively moving l.trgl·-,ized prey. The tr
rorumenration. \01111..' doubt h.l\ been accorded e\'olutwnary history .md funcoon.1l morvhology
)one,, I91\ I) a' to Jll't lum ·,uddt•n' rh~: ~pecianon shows how well they \H'rt• .1bk to exploit tt. ti.1r
1\Ynt~ .JCtually \Wre. I hl' 'mtemtl'dtate' un!ttable they arc ne.trly all ,tni,·c..· t'.lrnivore' .md. other dun
:m. klnm pe~t\ted 111 tht• 'I urkan.1 ltnc.1ges for up to tl<;h, [he mO\( accomph,hcd \\\'llllmt· ~ in the 'ea.
rei! UUO yea!'\, bctWl't'n wrv llllll:h longer periods of There have bcc..•n cnmidl'J'.Ihk ditliculttcs iu
230 Molluscs
cl.l~sif)'u1g n:ph.1lopmk cspeciJUy 111 erecting large raced by a AuiJ <.usluon from the l:m ~eptum. Tl 1•1 N.
natural carcgont'\ soft patt~ can be cons1dered (cf F1g. ~.21b) J\ 1111
The scgn:g.mon of thl· Ct'phalopods mto two separ.ltC UIUt\: (I) the bod). \\ luch 1\ nllly l.'tldo
broad diVt.,tons, I ctrabr.mdlt:lt.l and Dibranchiata, by the mantle and contam~ the vt~cera and the ITllll
erected by Owl'n in 18J2 .111d based on gill mor- tic cavicy wtth 1t\ contt'nb. and (::!) tht hc.1d-l~10t
pholo{zy. h.h tlll\\ been .tb.mdoned. Wherea.~ many cartibgc-supported ~tntcturc With JX tenudc, sur r11arnt
aurhorinc' (e.g. Duno,·.m, I 964: Teichert, 1967) rounding the mouth With 1t~ horn). p.urot-1±:
prefc:r ro divide tl·pll.llopods uuo ~e\erJl mbcla,~es, jaws. The eye<> are pl.lcl·d later.tlly on the head-too
orhe1" (c.~. Hull.111d, 19H7) recognize only two, the The dorsal parr of the head-foot abo\e the trltD-
EctOl<Khli.t (u:pll.llopod' \\'tth external shelli) and. clc~. ha' the fonn of .1 h ood \\hich nom~
Cole01de.t (n•ph.1lupmb wtth tntem.1l shell.;). Dzik's extend., some way up tht· o;hdl. Th1' h.t' a toug
(llJH4) d,l\\lfllJtHlll 111 whKh three 'ubcla.,,es ;~re warty outer ~ku1, so that when the cenucb :u
propo,ed (NJulllotde.t. Ammonotdea and w irhdra\vn for protecnon the hood dos1..'\ the apet
CokotdcJ) " tolkl\\l'd here. cure, prc~cnting a Llrgely impenetrable and tuum·ll
ing surt:lCI~ to :my predator. The full ~.>xtent of tll(
SUBCLASS NAUTILOIDEA (U l'.u11 -Rt'c.): Shell (phragrno- unretracted hood is mark.ed by a black film on 1h
conc) exll'rn.tl (<'dllt·m hliatc), str.tJglll. curved or c:o1lcd, shell. only expost·d when the hood i~ closed dm1u
chJmbt•rcJ wllb \llliJlk 'utun·,, ~iphundt' central or ~ubccn over the tentaclt~. lklow the: t~:ntaclrs 15 th
tul. nft:cn of Cl)lllplt'x timu Fmu g1ll' pn:scnt h y ponome or funnl'l: a long tubular ~truuur
SUBCLASS AMMONOIDEA (l Ot•v -U. Cm.)· 'ihell extenul;
cotlc:d. ulll'l1 nblwd t h 1111l>cred v' 1th compll'x suture\, which can be t1.1rned 111 .my direction.
stphundc: ventral u1 11C'.1rlv dor-..ll, nf ~•m pie fomJ. Gill nwn- Nautilus ~w•m s by jet propul~ion; water enter. cl1 le)
ber unlulu\\ n mantle cav•ty through a ~ht-like mhalant p~
SUBClASS COLEOIDEA (?C~rb .-Rcc .) Shc:ll intcm.U. ,tr.1ight and, passmg over the tour b'~u~ withm tbt> m;~ut!
or ulllt"J, "phunclc: nuy he I.Jcl.m~. 1 wu jpll' presem. caVIty so that n.:.,plratory exdungt> 1s dfetted,
~qumed through thl· lwponmut> by the contracn
Subclass Nautiloideo of powerful mu\t Jc, of the h}pOm.ltnt:. (Unlik
Nautilus (Figs 8 1, 8 20, 8.21 a-e) 'qmds the m.mtlc ot' '\'auri/1b dot'S JK1t have mu
The only hvm~ u:phalopoJ genu~ w1th a coiled lar waUs, and the mu,rul.nurt " confined to a !pe-
external shdl '' , ,,tr/us, of ...,·Jlich there are ~ix cializcd 'ac· the branc hial c hamber ) A; u• 01L
h\1ng 'PCOl'' lnnlim:J to the (ndo-Wcst Paofic ccphalopod' th~..·rt· IS J coordmated '>)"'>tcm ot rh1
fauna! pro\ tnl"l! between the Pluhppines and Samoa mic flow by mcam of which water J\ pum
(Sa under\ wd [ andnun. 19~7, Ward. 1987). The through the mantle l.l\'lt) with t:·mly gt·ntle rcg;b
Nauttlus ~hell. ~omc.: :w cm m dJam~ter, is planispiral movement\. '\'mttilu~ Jl\o h,ts an e'\cape re.1lCJ
G
and omamented l'xtenull) w1th J radial colour
banding of irrcgubr .tnd btlaterally symmetrical
orange--brown 'tripl'S. Thi~ sh~U is divided into
mcemal g.1s-tllled dwnbers (or camerae) by septa, /
concavt• toward' the .tpcrture; the animal resides in Hr up
the b~t ch:tmbcr (body c hamber) and mm·e~ for-
wards each tinw a new sepwm ts secreted. A single
tube (the siphunclc) p.t~ses through the centre of
each scptum .md connects tht: chambers. Each scp-
tum ml'rt~ tht• mne1 w.tU of the external shell along
J ~hghtly curwd hue (suture line). Fossil I',:,wtifm
and m rcbtive' art> U\uallv pre,erved \'\,th the shell
di~snlved .md the 1.. hJmbers filled w1th spar or Ftgure 8
of ihe or
matnx. In 'ud1 l';t\e~ the pos1U0n of each scptum is (Jur ), th4
marked by a \Uture hnt: (irowth 1\ wry rap1d in liv- tube; (e)
mg Vautrlu~. ne\\· \~.'pt.l bemg emplaced .tbout every Figvre 8.20 ltving Novtilus pompi/ivs (x 0 25) from a~ tlcbOne c
1 \\eek., on .wer.lgl'. The liVing animal it~elf is ~ep.l- graph. boloncill
cuHiebon
of shell sJ
Brown, 1
on Dento
f•l' fNIItilu• pompi/ius

~nal tube !porous) (b)
sophuncular epnhel um
sophona1 tube
Ill
Slphuncular veon
11
4%
~re8.21 (o) Noulilus pompilius (Rec.) shell sectioned subcentrally (x 0.5); (b) ammonoid morphology reconstruded on the basis
ih! anatomy of Nouli/us showing inferred disposition of soft ports and marginal siphunde, (c) reconstructed 1aws of Hildoceros
\f.), the lower jaw having o poir of optychi on the ventral surface; (d) anatomy of Nouli/us siphunde- transverse section of siphonol
be; (e) longitudinal section showing junction of septal neck with siphonol tube; (f) buoyancy mechanism in the cuttlef1sh Sepia (cut·
to· !!bone of olaer chambers is ~lied with liquid ond younger chambers with gos at about 0.8 otm pressure; o lift of some 4%is imported
dancing the excess weight of the onimol for hydrostatic pressures of the sea up to 12 otm; x 0. 25); (g) structure of lower port of the
,llieboM with chambers supported by pillars and separated by lamellae; (h) Spirula, o small Recent cephalopod- median section
~!hell showmg ventral siphuncle; (j) onimol in normal life position, showing location of shell. [(a) Redrawn from Denton and Gilpin·
~!own, 1966; (b) based on Trouth in Treotise on lnverlebrote Poleontology, Port l; (c) redrawn from lehmonn, 1981 , (d), (e) based
Denton and Gilpin·Brown, 1966; (0, (g) based on Denton, 1961 .)
232 Molluscs
Hwolv111g a violt-nt comr.lctnm of the br:mchul \\;Jrd-puintin g septal n ecks p•L·rnng ead1 ..cptum lltl
ch.unber. ~o th.lt the .mim1l htcrJJiy JUlllp~ out ot ventrall). In hfe thn< cam the >lphundt·. a sm~_ok cpi
the way of an" prt·tbror m t'qmvalcnt mecll.llllSlll ~trand nf h\ing tl\sllL' rxtendmg from tht hodv t l.hi
m \l]llld~ 1s otten .accnmp.lml'd by tht• t'nm~ion of the protoLnnch .md c.lrryll1{!; .1 nch blood suppl} Cl' I)
ink 111to the \\ .lll'r rh rough the hyponnme from Jll
l.f)"
ink ~ac wnhin the mantle c 1\'lty. Thi' btolo~caJ j~:t BUOYANCY OF THE SHEll IN NAUTIWS AND OTHER COl
propuJ,•on 1~ the ,t,mdJrd me Ill' llf lnconH)tlOn 111 CEPHALOPODS Wll
crplulopods. ~mce the whule 'un e'~ of the n·pi1.1Jopod~ Ius been tim
1\,wtilus. b..:111~ ,, mobile t..:cJ..:r. h.1~ .1 highlv so llltlm.ltdy bound 11p ,,,th their po~\C'i\IL1H of Hot
org.ma1cd bram l ouplcd \\ ith a..:ooJ 'Cil'~· org.u1s .md buoyant 'hc:lls. 1t '' appropnatL' ro com1dL•r hull .md
a <:ompkx brh.t\'tour p.ltt;rn. \N11L'll hunting, · modem cephalnpnlh uf different kmd\ at:tlWlr
rh(
.'-:tWtt/11$ sprLalh lh Olltl'T tl nt.ldL'\ in .1 'COTlt" of achievt sULh bun)ancy.
tals
~ea re h', bur wht·n 1t c<~ptur~:~ til()d tt ll\l'\ rhc mnn The rL'St:an: he' of Dl'11 ton (I')(, I , 197 4), l lentull blot
tcnt.•rk' to h.mdlt: 1t. I hl fi.KHJ, "' h•t h com1sr' and G•lp•n-Hro\'> ll (1 1)Ml), \V.1rd .md Manm (Jl)/:o! able
numly oflar~t , ru •t.ltcan' .llld ti,IL j, L\lt up b) the and other.. han· ~hown clear!~ tlur rh~ buopncy o:
sile
bt 1k .md ~tnred 111 111 cxpamkd oc,oph,lb'll\ prior to ccph.alopmh work' on a qmte ditferult pnn.:tplt
~,JfC
beilll' p.IN.·d (l1 tht '>tolllJCh .111d digco,tl'tl. from tlut of fi~ht:~. In tishe~ the 'wim hbddrr con- inte
'\c~lltliu' ha' a r.lthcr con1pkx n:produa1vc: p.H- tJ.im g,t, .tt .m equal prc\'illfl.' to th.lt of the ~urround the
tcm The \t'Xt'\ o1re 'cp.lr<Jte: tC\te\ .md o\'Jrit·' ,1rc to mg \t".l" m deep-\\Jtcr fi~hc' tht• mtcmal prt·s~ure 01
T
bt• tnund at till' postcnor ~xtrcnut) of thl· body. rhe s\\'1111 bt1dder ~an bt: l.'nommus.
rum
Dunn~ copul.!rwn the mall' tr;m~fcr~ ,1 b 111 of sperm In contrast. thl.' ,J,dls of < cph.1lopod' \11Ch ~ rhe
(sp ermatopho re) to th~: nuntll' t,IVtt:y of the J\:'a11ti/u, ,md the 'lJuiJ~ StJI"' .md Spil11/,, (wh1lh scp.1
fcnulc u'mg .1 'Pt'et.lllv ,ltbptcd er\.'t ttlt group nl luvc •ntt:rnal shdh. l1g. 8.21f.h) .1U cont.1111 gas ~!a new
tcnt.l ~c~ (the spadix). rill· lcrtilizt•d l'~g'> lrl' ltr~t prcssurt.• ot lc~s th.m I atm, anJ neutral btalpllC) at mit1.
and tht• JUVCmlo ot '\,mllfll, lurch at 'omc 2'> mm diffrrl'nt d~pths 1~ .tdllt. vt:d nnt by prc\~urc equ.tlm·
\l:ptl
in 'hdl d1,nncrcr wtth Jbout ~cvcu dumbt•t-.; .llteJdy non but by dcmrt)' control. In both ,\:,lllri/us lnd
~t.llll
pte\L'nt Other tt•plulopod~ uonn.tlh h.t\ t' only ont 'icp111 the chamber' u1ntain !:!·" ,1t pre\\Ufl'' fJilt,"'n. pum
modaticJ tt·ntacll· (the h ectocotylus), but thc1r p.tt- from Jb•mt lJ.J arm in the mur~.: retcntlv fonned
111101
tenl\ of dt,pla). t ourt~htp .md tOpt~l.ltlOn .lrt· often clumhL·rs to about OX- O.I.J arm 111 thL oJJ,·r ch.un-
'•lJ V
qmtt· complex. All spct'll'' of \,wtilt~s h.wt· ,Jitc:rn,ltl' ber:<; of the shell. The living N.mril11s i\ ,\ightl> ncgJ·
but t
period, of reo;t 111d ,lt'tlvlt)' lndl\·iduJ J, .lr\.' active .1t tivcly huoy,mt. 1 c. heavier rha.n \c.l \\. tl'f. In tunc
mght .md dunng the d.l)' thcv ~mk to the 'ea tlnor immatur~ 'pennH:m the c h,unhl'rs contJm " tan ~(l th
where tht.•y rt·~t \\ nh the hood pulll'J down Jml Jmount nf lrqllld, the cauH.r,ll watl'r. hut m fully
\Cry
allno\t covcrin~ tht eve\ grown \bells the volume of t.unc raJ w.ttt:r " VC') cxpiJ
o,mall lt ha~ been removed by t•xtraction viJ the tom,,
THE SHEll OF NAUTILUS 'iphundc. Although 1t w,t\ iormerl)· bdre,·ed dut buor
The ,hdl of '\,uw/us j, nJJdc of aragomtt' m a con- rhe .111111111 could n~e or smJ... 111 till \\.Ill· I b)
~djll'>
chwlln matnx 1t tmm~t' of rwo m.llll l.1yers: tht: ~hort-ttTm 'ccrt'tJnll 01 extrat·uun of CJ.llh:r.tl h<.JUld,
remo
outt:l pt.lrccllanouo, mtrJcum .111d the mner nacreml\ it ha~ now been shown (Ward. l 40(J) that thio, nptr·
!Jquic
lan-r The outl·r 1.1\'er i, lnrm~.:d h!'\t .md grow' arion t.lKt'' place on I} V Cl)' ,Jow ly. In tht.• ll\ a,
long-
trom the nt.~ntlt· .H rhc cdgl' of the shc:ll, be!Pnuing '\'m wit" only tht.> more recent!)' fonneJ L h.11nhm wrtic
a' ,11.1gmut:e wl·tl, m a t.OnL h10lm matn x; thest cont.un hqmd, and th1s i~ pr~o:o,t•nt in dnnim,hmg
port •
benmte brger .md clmdy pad.ed to~etht:r ..1s they amount~ from the newest w the older ch.u11hm ,]l t~ J;
grll\\ mtu vertll.ll pri~m,. I ht mner n.ttreous IJ)t'l Thm the c h.1mber' b~:yond the tl'nth ti·om the body mal i
i\ lat~·r depoSltl·d b, the nunrlc m a 'enes of furm: chamb<:r ,1re empt) of liqu1d.
cxt~H
rathL'l like till' n.tl rt:om l.lycr of b1v:tlvc' lt com1st' It ~~ the \lphumle thar extrad~ tlm hyuiJ. Tht uid lS
of J bnck w.lll \trtllturc or hexagon<ll aragonite siphunrle {Ftg. H.::!l d,e) con~l\l'- nf two p.arts: th, buoy;
Cl)'' tal' wlth rom hiolin l.Jyt·rs bet\VCL'll them. The imperntl'.1bk sept1l nc:cks, ami thl pennt.lhiL• o,tr.lndl \\:.Iter
!>ept.l ha,·c fuml.lment.lll V tht -..une 'trut.turc. An brt\VeUI them (siphonal tube) ] hj, tulW h.u all
b, hy
elllp[\ ,\",wti/11~ ,hdl (F1g X.21.1) sho\\' \hOrt back- inner core of livm~ m:ucnal "' lth .lrteric' .111d \~Jrn
Wl
rl'nnmg tht: whok kngth of H. w1th .1 cylinder of hyponome. the ,hell does not go mto a 'PII1, 'inn:
pnhdi.1lcdk lh1Nd~· tl11' '' 1 horny tune of con- the centre of gr~wil) " loc.lted a f<..•w millimetre~
chll)lm fihr~\..md 'urroundmg tlm .1g.1in is .1 con- dirl'cdy below the Cl..'ntre ofhuoyancy (t.e. the n·n-
ennic tubl..' of 1rrc..•gubrly ,1rrangcd Jragonitc tre of gravity of the di,placed \\ .ltc..·r). Tlm imp:m' a
Cl)"itak A \·cry thm c..·~tc..·rn.ll pdhdc of conchiohn remark.lble stabiliry to the 'hell, .md 1t would need a
complete' the..· c..•mc..·mhk. When· the..· \lphunclc join' \'Cl')' strong coup!t• to n1rn tht• .lnmul on it' \ide or
111th the..· \c..'pt.ll twd'' tlw horny t11bc..· become' con- through 90°
nnuou, with tlw n.Jc.. renu' m.ltc..'n.tl of the neck. N.mttlus " unlike .my c.Jtha linn~ cephalopod
Both rhe .1ngon1t1..' .1nd htlrnr l.tyero; .1rc..· very porous \II1Cl' H c.m hvt .lt rc..·marbbly kw..· oxygen ten,iom
l!rJ pennit the P·"'·lgc..· of hqtud through them. (Well' t'l al.• I9'.)2). lt nunn.tlly inlubit' rd.1tivdy
nr
Thae I\ .liSt) ,\ thlll l.t} c..•r $,\ttl'rc..·d ,\ragonitc..· cry\- dec..;p watl:r, m $r do'e to the: m.:ygcn-mimmum
uh on dtc..• rotll.l\'c..' w.11lof till' ,c..•pt.l, Wlll(.:h .Kt\ like l.Jyt•r, growing ,Jmvly .111d mtng onh J lunitec..l
bh1ttmg p.tpn ,llld 1'\~nc..kr, tlw ".111 ".:ttabk .md so amnum of encrb'Y· Tl11., ~..·u.lhk' H to ,tvoid ~..ompeu
abk tc..• ret.un tlw c...llllc..'r.JI hqtnd. In tenns of it\ tcn- tion with 6~Hl10\'1llg h'h .llld sqllld 111 the upper
f IIIC ~trcn~oth, the \lphunck· h,1s bc..·c..·n likcncc..l to 'a \V,llCf\ Of thl' \e,J; il Jt,l\ .Ill optima] <.kptJt range of
!: g;trdeu ho\e ln.1dnl llltc..•rnally by water pressure and 150-300 m. \Wimn1mg .1bnvc..• the \ea Aoor as a
nJtemJlly supported .1g.11mt 'lJlllrmiog imt.lbility by rathc:r gc..:neralizt:d, ,Jow-lnOVIIt~ \l'<tvc..·ngcr or cami-
the \t'pt.J' (Hc..•w1tt .\lld W c..•stermann. I 1 )~6. 19~7). vorc. Nautilw only p.1y~ brief nocturnal ,.i,tts into
The bud~ t.ll r\!,mtilm I\ Ill ~ Ol\t,H t With the sep- the ti~h-dominated :ton I.' ,1bow, .md is \cry rardy
!\1111 dunug rhe tmt~· tlw lattl't " li.11 med. Eventually found abnw 75 111.
h the hod\ tllO\'t.'' .m ·'Y frl1tll the: (a,r 'eptum <llld i) A fin.tl point romcrm the..• dc..•pth to \Vh1ch the li\'-
h ,p.1wed from it bv .1 ~wduon ,,f h'-!uid. When a. ing Nautrlus c..an 'mk before the .. hdl unpludc\. The
.l ew 'cptum " f(mnt:J 1t .:nck"t:' a dtambcr which shell is very 'trong .md ngtd, Jnd \:Xpenmetm have
mttJlly retaim .ill tlm lttjuiJ. Howt:vcr. when the shown that an .tdult shc..ll would nnt implode until J
~eptum 1s fully t(mned .mJ '' 'tron~ enough to Wlth- temtle mength of U I MPa \\a~ applted, corre-
IUnd tht.' pre,.,urc of thl• 'l'.l. the..· ''phunck begnh to 'pondmg to a depth of 7110 l-no m Newly h.1tchcd
ump out tlw ltqtud, lc..•.wmg l'lll) a ,m.UJ rc,idual ;\'muiltts 'hell~. howcv\.·r, would tmplode at a depth
omount \\ lndt '' r~ducc..·d 111 ,,tit anJ hypotonic to of only 300 m (Hewm .md Westenn,um, IIJIP). the
sea \\Jter. Thl\ purnpm!! pwce'' works b) mmosis. last ~eptum bl·mg the Wl'akc..•\t p.1rt. Wtth age the
but the \\ .tter .Ktu,JIIy Ill the \tphunde .tt any one .uumal 1s .tble to go down to gTt"Jtcr depth, and
an1~ may be 'dccoupk•d' fr~..1m that 1n the chamber, althou~h there are rl'port\ of ltving individuals ven-
10 thJt the ,tctual wurk dont.• j, not \'t:ry gre.tt Gas turing to depths .1ppro.1~ lung tmplos10n limits. thi~
l'rl) ,[o\\ ly dtfli.I,C\ tllto the ~p.lll' ldi:, whid1 may 1s rare.
txpbm \\h) the.. lo:·" prt:\\lltl'\ 111 the mmt ren:ntly
1omu~d dum bel\ .trc..· luw. \,11111/us c.mnot adjust its
Evolutionary diversification
:\M ~lit>} Jilt. y quH.kl). It 111.1) tJkc..• wct.•ks to makc a full GENERAL CONSIDERATIONS
b) diu,tm~..·nt \IIKt" tlw m.lxtmum rate of ltquid The earl1est known cc..·phalopod~ .lrt" the sm.1U curv-
tic..!. r-:tnoval ~~ no more th.111 I .o nu d.ty . The cameral ing shdh of JJ/cmt>rwn•ras from the Upper Cambnan
~:r tqutd-phr.lgtnol'onc \)'\tc..•nt m.ty be used for of China (J~i~. 8.5). rhey are endog,l\tric, wtth mar-
ing mg-t~..·rm buoy.1nc.. v t. h.mgt.•s bur not .1s an aid to ginal, empty s1phunclt-s Where these occur they .1re
K'TS cmcal movement. ( ·,lmt·ral watt.·r 1s needed to sup- very rare, but at a slightly h1ghc[ honzon there IS
_in~ n each septum a., 1l 1\ bemg fanned, .md 1t Jlso ev1dence of J modest adapuve radiatiOn. r ow;~rds
crs. 1ls .1 r~·~ervon ofhc..]Uid halb~t which. as the .mi- the end of the Upper CambrJJu. how~ver. there
odv lllll f.,'l'OW\ and IJlc..'leJW~ 111 Wl'l£;ht, IS Steadily was a ph:~ se of C'\.1)los1vc rac..h.ltton ('' hKh mduded
mracted. In .1dult' rht• httlt• rcmammg c..'ameral liq- the tirst ellesmeroc..erttld~) .md \\ lm·h Holland
fhC' ;;..lt~ u~l'd onl) tc..'r m.umantmg ,t o;hght negati\'e ( 1987) regards a-; ·.1 Ltte ph.1~e of the uunal met.uoan
the..• ;10pnc). When \clllti/11~ nugt ate' upwards in the radiation, m which perlup~ J new combmanon of
mds ~ltrr column .H m~ln m sl..'.lrc..'h of tood. tt does so physwlogy and morphology took it~ mt:hcs in the
' 311 ~ hypononuc "" 1111111111g alone. ammal world'. From ~uch anei..'Stor<. caml' the vel')'
·cim \\cl1en the \,llltilu.' ~qu1rl\ w.Jter through its nuny Ordo~ 1c1 In ~enen \0 common 111 the fossil
234 Molluscs
(
ORC
Yt'lll
Orthoeeratoda
.. tnllo
ATI
,.--.-.. ~
.,:; Pllt'l
z cu
pn:v
'cun.
/'ilt.,-,
ORDI
gaccc
e.g.
ORD£
'hell,
<'.g. L
ORDE
\hdh,
\hort
l'nlfal
OR DEl
families \\ nh l
..____0
20 •t•Jy b
~•phun
ORTI
Oflt.llll
~ht:ll i
Ammonoodea Coleoodea
COnnet
r)rrlwa
Figure B.22 Time ranges and abundance of the three cephalopod subclasses. (Modi~ed From Dz1k, 1984, House, 1988.) rile IM
dncal s
CXCl ..>:\,~f
f.lllll(C$,
\l!O\Jdl'
record, for the t'.1rlv Urdo\.ioan was marked by a and chrect competition 1s not neCl"S.S.lnl} the onh Op.:J <
long 't
further cvolunon.lf) cxplm1on of ccphalopod gen- tssue. One tactor n1.1y be that the anunorutn wr~
hfllltltt\
era. The~c lud 111.1111ly straight (orthocooe) or exceptionally well adaptt·d tor reststm~ unplosJOn ORDER<
curving (cyrtoconc.") 'hdls and were generally of the depdiS they utl1abtted. Another 1s that th g;•ted ,]·
much brgcr ~•zc thJn the•r httlt> Cambrian ances- became ccolog~cal ~peoall\t~ mvadmg a grc.1t num llJ'I llt'IIC
to~. From the om· on~11.1l order. the Endoceratid.t, niches. unllkt.• the comparanvdy unspcnahzcd Sl';ll'- llndudt'
there .ume by rhe e:1rly Mrddle OrdovJctan many enger, Nauri111s. Yet the ,unmouoids became t•xtinct I'.•Ll•ozl
other order~. whu.:h pe~mcd through JIJ or part of the end of the Cretan~ous ,md Nautilrts did nlll. IINA (i
NAurr
rhe P.1bcozoic (f•~· H 22). Ammonoids were very much the prey of fi.~hes •n dll'\l' l'J
The l.m·,r of thc orde~ to appc:~r. in latest marine repttles ;md there are plenty of b1te-nurb o horn•'<) !I
Siluri.m or t'.lrly l>t·vonJan tJme, was the Nautilid.1, thetr shells to prove it (Marttll, 1990), but then thcrt
in \vh1ch rhc lt\ •ng N.lllti/1/_1 i~ cl.1~sified. no reason why naurilo1ds should not havc been 'iO t~ GeoJ,
Mosr of the\C Pabeo7ote ceph;tlopods became Holland's (llJ87) suggestion that hatchmg stzc IlL I hL po
c~:tincr by rhc end of the Penn1.1n. only the coiled- have becn important seem$ cnunently reasonJble. abund,u
~hcll N:mrihd.1 \liT\1Vmg through the Mc~ozoic to the young Na11tilus h:~rche-s :.s a 25 mm animal mm~ frt.>lll th
pn.....,enr. The dccllllc of P.tlaeozmc tcphalopods seems arely able to take up the deep-\\ Jter foragmg life Y} It'pre,cu
to bear \ome rd.ltiomhip to the mccc~' of the1r prog- of it!. parenr.>. Anu1101101ds. on the od1er lund. ln•1ulcls.
eny, n.uncly rhe .lmmonmd~. and e\pt>oally ro the tmy hatchlm~. much more vulnerable to the col ual .Jrc
ammomrt·, of rhe Mt',o7oic, though as m all such of the planktoruc ecosystem which they inhlhited, orhcr ta
Gt~es of.1pp.1rcnr 'tJkl·ovn'. mJn} !Jctors are involved the end of the Creuccom. ~achly b
Classification Only a few of the P.tlat.·o:zmt n·phalopods have

ORDE~ 1 ENDOCERATIDA: l'nmmw n.aut!lnhi, wHh a \VJ.de. pl.lnispiral shdb. Thc maJOnt:y an: of either cyno-
rrutnl n·hndri,o,al ''phunck .ami .1 ,tt.u._:ht to <·nJo!(.btncally cone or orthoconc sh.1pc. Thc'ic lllJ) be \'eJ0 don-
,oaled 'hell jlnduJc, Subord~r. ELLESMEREOCER- gate (longicon c) or ~hort and rath~r ~wollen
HINA (~h,,n -.·pulm·, k~ \\lth duck cnnnl'ctul~ nnhrs). ~:.g.
(brevicon e). lr ha~ bl.'t:n gt•ncrJIJy a'~tunc.:d that all
E!lomcoto>(ttt<, H.~tlamolt'oils, l'l.ur.>llt'<<M~. mJ ENUO-
CERA !IN'\ (H'r. k1n~ 'cpwl nn k, wh1(h lll.l\ mtruJe the tossiJ cephalopod~ w1th ,rr.tight or cur,·cd ~hells
prt\10u~ '(pm m: vcmul ''phundl· tilled .1p1cill~ with were ectocochlear . L t. that the 'hdl wa~ wholly
,an(-m-colll'' 'tnKtllrl'' - <·mluc••nc,). l'.!'· Eudt'fl'rou, extenul ro the hod). The X-ray photogr.tphs taken
1\l>ar.t< .] by Sti.im1er ( 1970) of ccrt:~in gen~r,t from the lower
Oi!DER 2 TARPHYCERATIDA. Exu~Nril.all) coalt·d 'heU. elon- Devonian Hunsri.ithducfer ,fW\\ very ckarl) that
~.red bo..ly d1mabcr. n hndm a! "rhund<·, l.1~c protoconch; m some ca~cs there wt•rt• hvmg us~uc\, including lat-
r.c. r.,ph)'•~'~•l>, !J,,.,,,.,,,, Tr,,,ll,•lrt<'<.
eral tl.ns, cxtcmal to rhl' \hell. Some of the ortho-
ORO€R 3. DISCOSORIDA FnJngNnr.allv <urved. compressed
1h.U. \\1th \'l'ntr.•l ''phuude mJ antl.atcd <nnnecnng nng;; conic P.llaeozo1c ccplulopod' were apparently
t.g. Drs"'"'"". Pllr,r!!m"".''" ancien t sqmds, preCUf\OI"'i of the I.Jtcr bdemnires.
ORDER 4 ONCOCERATIDA. Mmtl} cxng;t~tncally curved However. at present, w1th only the
lladh, v~unal ''phu11d<· wah anH.lt<·d conncctmg ring.. and Hunsriickschtefcr 'window' lo look through, it i~
•hort living dl.unbcr; <'.g. U11comru, Rrch.m/s(l/IOCI'fa>, not known how n1Jny ot' the Palacozoic orthocones
Ptllfcll/ll'l<ltt'I.IS .
\.Vere really endocochlear , for in Stlim1er·~ pho-
ORDER 5 ORTHOCERATIDA. Srr.11~hl to weakly curved shell
tographs both intcmal- and extemal-~hclled kinds
\\IUl ~ubcentr.!l''phunrk In some c:xtr•·mc fllmlS the sheU
111.1)' b( •hon or exog~'fm,Jll}' 'parJUy c01lcd and with the arc evident.
~phunde \·cnml m dorul. (Include\ ~UBORDERS Nautiloid~ arc classified on various cl1dracters, but
ORTHO( ERA11 NA (Ion Jot ,trJaght 'hdh \dth transvl'r.IC perhaps rhe 1110\t important arc the fonn and
~namcnr wncrAI \lphunrk Jnd mllJtc:d, embryoruc part of arrangement of the various Slphuncular structures.
lbrU mfi.H< d, h\111~ d1.1mbl'1 Ion!(. ~ubn·ntr.ll "phunrle. Modem , autilus (Order Naunltda) has only
(Unnecong nn~. • ~ lmdncal to 'hghtly m Rated), e.g.
septal necks with wh1ch to ~uppon the: s1phuncle. In
Orl1•¥tr•li, .\lnlnlwN< 1.11, -l.m'<t ''''· Cycl.vttt.< (Dzik mclud~
che BACTRITIDA here): Lll UITINA {'ubcentral, cylm- the ancient group'> there an: a variety of other stmc-
Jneal S!phuncl<• '' 1th hmH '''1''·'1 ne< k,, JPICJI plrt of ~heU run~s. c;omc of them of verv lOmplex fom1, and in
CXO~•IOC:I')) l'OIIeJ, hlllnc-1 \1111" ll.liTOW J.Jld deep). e.g. particular there are often stntc tu res w1clm1 the
Utuot >;,.,,, •r.t< ; ACTI~OCERA11NA ('aphunclc wtth c;iphuncle tt\df (en dosiphuncula r \tntcwres)
un,•Jrnbl) •ntl.Jte.l COIIIIC'< till!; nn~ .md wah well dcvcl- In Order Endoterauda the -;hell\ an: either ortho-
cpcd lJUll'r.ll dcp<l\lh 111 r ad1.1l block.,, l1rg•· protocone h. conc'i or cyrtm:om·'i and the "phuncle~ arc: usu,tlly
.
..
O!; maaglu 'hdl, '•phund•• ttl\\.ard, vcntrJI ''de), e.g.
trm''"""'· R·•Y•'IIIIt>tt'l<l•.l
broad and nornull) mar~n.1J. and mmt genera h.wc
~DER 6 NAUTILIDA I.:C.wg.l,ti'H\llly CIHird. modrrJtcly don- nested endO\iphuncular lOtucal she.Hhs or .tltema-
ttd shdl w1th 1\.lrrt>\\ 'ubn·mr.•l ''phunclc L;lfVJl dcvel- tivcly radial lamellae rumuug the length of the
•pmem '' Hhlfl C!-:1-t rlp~uk. nn pl.anl.romc IM\·al stagl!. 'iphunck.
llndudC<~ 'IUBORIH:.IlS <.. L:.N'TROLERATINA (Early Order Orthuterattda, wluch permted inro rhe
l'.a!Jeozl"c), <'.g. <·..,.,,,,, 1115, l'rt!clwccr.u; TALNOCERA- Trias\ic, arc mu<llly ~tr.lip;ht-shdJed ,111d have ~imple
lt
TINA (L.ltr l'.11.lcu7nl( .111J l'n J'>~K). e.g. I 'estinautilus: and
t. scpral necks with only thm connecting ring<;, .md
N:\UTILINA (pmt- fn.l\~ll') c.~ f\;,uttrl~<~•. Jrurit1, though
d th~~ cJnnor be S.ltl\f:Ktllllly dlJj.ttlO~l·d d11t t\l frequent
sometimes cndosiphuncular structures.
•0 h,,meomnrphyj . Rcprescmativcs of Suborder Actmocer;ttina all
IS have orthoconk shdl\. The septal necks arc short
o. Geological history (Fig. 8.21) but have inflated nn~l'i between them, withu1 \\'hich
l)' The pmt-Cambri.m nJuulollh ,m: wry diver;e and is a delicate and wmpkx system of radul canals.
lt" 1hundant ;md oftl.'n qlltll' J1ffcrent in shell ~hape Some orth occrarid gl.'ncra have shells of deodedly
&-om the living '\,lutr/rrs. A~ 111 their sole modem odd form, ~mh as the OrdovJCI.lll UlHtH'.s (Fig.
rrrrc~cnt1ti\'e, th1.: \Utllfl.' J.mc~. J~ seen 111 Ultcma) 8.13h) m wluch the early ch.lmber<; .ue coiled
mould~. arc m:arly .lh\ ·'Y' -.traight or ~hghcly curvmg and the r~t of the shell 1~ a raptdly expan<ling
lSt: JJd arc \TI) r.trd) more ~.·omplex. On rlus and orrhoconc.
at ~er £1etor; L v<.·n thl' pl.umpiral uautJloids can Even stranger arc the shells of .-lswct'ms and
1 Jdtly be dt\ttn~mhc.:d fmm the .1nunonotd'\ GJ.moccra.s (Fig. 8.23t). In these the \hell con~i't" of
236 Molluscs
(b) VaginocertJs (c) tWO (

...----;, body cavcty small,
C)'rtOI
scphon
so met
brevi<
septal neck
modi I
to rh1
connecting m19
fmm
fom1e
dorsal
cameral deposcts ;eptum
her. 1
and tt
tion v
mal \\
(d) CtJmpylocera:r (e) Michelinocerss (f) Glossocer11s W,IS it
w,t~ a
(I)
ceri<l,
nt·kto
The
cepha
gJve r
mere a
~oiled
\'le!\\" •
;111ce.
(g) Pentameroceras
(11
life, a
been ~
Occ
two la
(h ) Lifuites dorsal
cone~.
wtthu
the ~~
operct
(kl Lobobsctrlt. .
Figure
camera
(Endoc£
and lor
shell o
Campyl
inocero
and coc
Sil.; xl
(ii) trun
(iii) mo
(g) Pent.
hed opt
port, p
with 'c
Wissenl
noutiloic
dvced ~
two quite chstmct pJm I he hr<it-tormed part is a It I\ generally accepted that mo't orthoconic
11nall, rather \t,md.trd orthocone or ~bghtly curving cephalopods were free-\Wlmmmg forms carrying
cynoconc, With .1 thm \tr.ught stphuncle. This ts the1r shdb 111 a horizontal pm1tion. Some ev1dcncc of
sornctimt'\ found JOtncd to .1 much tlucker, swollen tins come~ from the rarely pre\erved colour markll1b"'
brc\icom· where the mtcmal stn~entre ts lughly tl1at occur on tht• dol'\al '1dc of the ~hell only and
rno<hfit·d. In tlm rht• ~1phunde 1s short and confined appear to h,tvc been a kJnd of <.amouflagc. But more
10 the apit"JI end. w1rh tnfl.ued connecting rings: importantly. the intern.1l ~trunurc' of tht· ,hell aho
from it the highly moditit·d 'ig~no1dal ~cpta are mdicatc a honzontal pmttion. Wtthin the ~hell, a~
fonncd to endo'c l.trge ch.tmber> located 111 the mentioned, there .tre endos1phuncular depo,it...
dof\al part of tht• 'hell 1mly. abon· the body cham- There are al~o regul:lrly 'haped m.tssc.:' of talcarcou~
ber. The t\\o p.lrt, .trc \W)' r.ucly found together, matenal known as cameral deposits (F1~. H.23c-e).
md it j, genc.:r.tlly .Kct•ptt•d that the longicomc por- These cameral depmtts were ~t:<.rcted prugrcssivdy
uon wa' tkuduou' .md rhrmvn away when the ani- from the ap1cal end a~ the shell grew: thq are con-
mal wa' maturt• Prob.1hly .1 th.tngt' m mode of hte cencrated near rhe :~pex and dcvdo~t·d le:.s ,md lc:.s in
was involved, 1t 1' qllltt' hl-..dy that the juvenile shell the chambers ne:1rer to the ,tperture. T ugcther with
was a nekrobentl11t .111im.ll \\ lu:rca\ tht• mature aKo- the endos1phuncul::tr depmit~. tltt'Y must have given
r~tid, whid1 h~J \hed 1t~ L'.trly ~t.tgt•. wa~ .ltl active extra weight to rhc apicnl end throughout the growth
n~kto1tic hunter of the shell, rhu~ :11lowing :l contmucd t•quilibrium in
The great dJver\llY Ill 'hdllonn 111 the Pal::trozoic a hori:zonra.l ~rructurc, with thl' ccntn·s of buoyancy
rcphalopoJ~. other th.m Ill thc't' hiZ;1rre ex:~mples, and gravity st:~ymg clmt." to one another (Flower.
~Jvr rise w .1 f(mnl'riV hdd conu:ptwn of a gradual 1957). Only orthocomc :lnd cyrtomc ~hl'lls have
incrtasr 111 cod111g from dll' ~tr-a1ght to the fully cameral depmit~. sm ce m thl" cmll'd ~belli of the
coilt=d (ondiuon Till,, lwwev~.·r. 1~ .m incorrect Nautilida and Ammonotdc.'J the two centre~ Ut' m thl"
Vlc\\, for each ~hell fi.mn. 1.'\'t'll if of unu'>ual appear- same vt·rrical plane m that the problem of st.lbility 1s
ann:-, i~ titlly ad.tptt•d to lh own parucula.r mode of ~olvcd another way. Pn."iunuhl}, .11- m the modem
!Jle, and buO)'JIKY .1dapt.llllln' Ill p.micular have 1'\'maiflls, all Pabeo701t n:ph.tlopod~ had some kJnd of
been .1 prinur, e\ oluuona~ comwl. 'lphuncular buoyancy control ..lJld a ~ltght change in
Occ.mon,tlly etrtul.tr 'tmcturc compmed of demit)• would have en:tbkd rht· livmg ammals to rise
two large, sym1 ·trit·al l.tt~nl plates and :1 triangular or !.ink. The ~·olkn, c.•gg-\hapcd ~hell~ of Order
Jor;.tl pl.ue .1rc tuunJ 111 .1~\lKI.ltJon w1th ortho- Oncoct•ratidae (Fig. H.23g) Jre qu1tc common m the
•One~. Tlm apryc-hopsid h.t~ been tound resting Ordovician and <;ilunan. They may have floated with
"1rhm rhe ~he-ll .tpertun. 111 a few \pt:cunens from the long ax1s vertical but with the aperture down-
the Sllun,111 of "l\Vl'dl'n. and IS ~urcly a protective W<lrch, 'earching tht· \L'J Roor with thetr tent,1clcs. In
''l'"rcuhun rhough pmb.1bly nm J .J.lW structure. these the aperture i' often comtncn:d mto J number
of sinu~e~ which t1kt· up their tin:1l ,Jupl' only 111 the
adult (Stndsbcrg, I CJH I , ICJHS). Thi~ nuy h:~ve been
F,gure 8.23 Morphology of fossil 'nautiloids': (a) Prolero- e%entia1Jy a protective devin•.
cameroceras (Endoceratoideo; L. Ord.; x 0.4); (b) Vaginoceras
IEnclocerotoideo, M Ord ), with cndocones within the siphuncle, By calculating the scptJJ ~lrcngth index tor
cll<l long septal necb (v 0.35); (c) actinoceratid morphology, Palaeozoic. nautiloid\, W e~tcnnnnn ( 1985) wa~ :~hie
1hell ond siphuncle partially dissected (x 1 approx.); (d) to show that v<lnom typ~:~ w~:rc adapted to ditferent
Compyloceros (Orthoceratotdeo; l. Corb.; x 1); (e) Michel-
•noceros (Orthocerotoideo, Ord. Trios.), with long septal necks depths. below which they wou ld tlllplode. Thm in
cll<l comerol deposits ( · 1), (f) Glossoceros (Orthoceratoideo; the later Silurian of Bohcnti.l there ue ( 1) hn•vt-
Stl., x1.8), showing three growth stages- (i) juvenile cyrtocone, cones wh.ith were probably l'p1pcl.tgic and gcm·r.tlly
1) truncated cyrtocone with 'ascocerid' portion growing and
restricted to depth\ of c. 35 150 m, (2) long~wnc'
u) mature a~cocend which has ~hed the cyrtocone port;
g) Penlomeroceros (Nautiloideo; Sil.), on ovotd form with modi- with thm, closely \p.lcc.·d ~epta. hvmg at Je,s rhan
:.ed aperture ( · 0.75), (h) Litutles {Nouhloidea, Ord.) proxtmal 300 m depth and (3) longKone' \\'lth ti1JCk, widdy
oort porttolly cot led (>< 0.35). (j) Hercog/ossa (Paloeocene), spaced .;epra wh1ch (Ould ha\'1: \\"ltll\tood d~ptlv;
WJ!IJ 'goniatitic' sutures ( · 0 35)· (k) Loboboctrites (Dev.)
Wissenbocher Schiefer - on X-rodtogroph of on endocochleor of < llt}O m. Likcw1sc . the CarbomferOU\
roohloid of toto! length 65 mm_ [The photograph in (k) is repro- .\lidtclitwwms hJ\ a '>trc.·ng;th~ncd 'hell which would
~by courtesy of W Sl\irmer.) nor llllplodt: unnl a depth of I I 25 m.
238 Molluscs
Several 'l hth0<.l'r.1~· lnnc,roll\.'' have been ttghtly coiled ammonmd~. All of thc~c haw. oth~r
Je~cnbed (<: g. m Morocco. ~t.mdmaVta and th.tn in their dch'TL'L' ot coiling, Vlrtu;tlJ~ tdrnnc.~
Chim). in wh1l'11 thc.•t·e .ue v.t~t ~·onc.t:ntrations of stnKturc, and it is in cephalopods ~uch ,,s thrse thlt
urthocone shdk Thc.·~e may bt: the.· r<:sult~ of ma~s the Jncrstry of :munonoids should bt· sou~ht
mortility aficr lll.ltmg. or ~.•lmlt\ dungcs in the (Lrben, 1966; Chlup:il, 1976).
v. .uct. A very ii:w orthncomc n.HI(UOJd\ '>llr\'tVcd
tnto the 1 n '"ll. hut llthcn.' i": onJ} the coiled How ammonoid shells differ From those of other
fonm of the N.lllnlid,t camL·d on. lhc elaborate cephalopods (Fig. 8.24}
stpbuncles of the. otll\.'r on.lcr.- .1rc.• uc.·vet found here; In the Jmmonoids, except m cert;un pc.·uth,lr genal
there is only ,1 thm. ~ubl'cntr.tlly ~•tu.w:d siphuncubr . known as heteromorpbs, the shell i' pl,umpinll)
'>trand. Most N.multd.1. cxtcpl tor ~01ne l.uc coded. Some Pabco7oic nautiloid shell' also hm
P.:tlaeozoic c.-yrtmonc,, have coiled 'hdh which may tht~ form. bur rhe.,c usually have .1 c. ( ntr.tl perfor:t-
be inv olute or evolute. Involute ,Jtdh have the Jaq non which is :tb,t·nL m all ammono1d' cxc<:pt 'ome
whorl cnnrch CO\'cnng <.~U the lot mer whorls: tn of rht• early OllC'i. The \Uturc une nornnlly tollowll
evolure ~hclh tlw tc.mncr '" horl~ arc all vtsiblc complex pattern; ca~.:h suture mark~ rhc JUncnon <'I
Somenmc~ thl'f<.' 1' cxtern.tl nbbin~ or even spines J scpmm with the inner surface of rhr ~htll wall, Jnd
in the po~t-P.1l.tcozmc genera, and occasional fonns, thc .Jrray of suture lines is visibk in ne,1rly dll
such as the Tri.l\\tc ClymeiWiftllllrlll) and the early .unmonitcs since the aragonitic shell rcadlly dt,sol\'l'l
Tertiary HtrlllJ?ft>~:;.r (Fig. 8.23j) .md Awria, have 111 dtagcnesi'i. If an indtvidual .unmomtc 'icptum nr 1
~uturcs rcmtm,ccnt of the ~oniatitcs of the mould of it is examined in face Vte\\' (i.e. as tflook·
Carboniferoll\. mg down the aperture}. tt appears lht or ~hghd}
curvmg in the centre but become' mcrlJ,ing~
Subclass Ammonoidea fnUcd toward~ it~ pomt of attachment to th<
Ceph:Uopod, of Subclass Ammonotdca (Dev.- shell, w here it bccome~ the suture. In the.· earh~r
Crct.) and l'\pC{ i.tlly thl' Ml'sO:wtc l~.mns known tn Jmtnonoids, of Dcvonian and Carboniferous ttmr
the \'ernacubr .1~ ',1mmonites' arc amongst the mo~t the \Utures are often simple zigzags, but from rh,
abund:mr and well known of all fo"ik Thctr beau- fnJ~~IC to the Cret;lceom the complexJt) of rht
ntitl planisptr.tl ,hell\, oti:en strikingly omamenced .1mmonoid 'iururc "lonsJderablc.
wtth Cl>."tcmal rihbin~. have been .tc,thetlc treasure\ ln most anunonotd~ the stphunclc I\ '-ltUJted nc3l
to innumnahlc. l ullectors. Yet to ~trJtigrapher; thl.'ir the outer margm (venter). One order. the.· Uppa
usefulnes\ lramc.l'lld~ th<:ir VI\LLll JttractiOn. for by Devonian C lymcnud.1, i~ ryptt'icd by ,J dor>JI
nature of thctr rapid evuhJtion, .tbundanee and ~iphuncle rwming along the mncr margin of thl
w1despn·ad d1,trihuuon they .m: the most v::tltt;tbk ~hell (dorsum); 111 tlm ~roup too the.· ~eptal nnl
of all fm~it' lclr /omng the rorko; in wluch thev .trl' rctrochoarutll (bar kw.1rdly pomtmg) a-. in mu-
occur. ThC\ have proved of ~petl.tl e tfccnvene'~ m ttlOJd\. and very long. This contr.1st' wHit the ou'l!l
the Trias'>H.", Jur.tS'il .md Cret.t,eum systems. where .ldv;.mccd anunonotd condmon. whc.·n.· the ~ql!Jl
their high turnovl't of spenc.·s h.1s nude 1t posSlblc to nec.ks are short and proch o aniric {cLm·rcd
erect zones equivakm to time penod' of less th;m a forwards).
million yc:m' dur.ttlon. Whereas the ~b ells of coiled nautiloid~ .tre oftcr
The anunmwtds nuy have been derived from thl.' unornamented or h.we only a feeble cxtcrn.tl \culp-
Bactritoidc1. a straight-shelled t<-·ph.1lopod subd:t\S ture, anm1onotd she Us Jrc frequentl)' nbbc.'d ..md the Figun
erol v1
wh1ch ranged through the Pa!Jeozoic; the shells of nb' may have knobs. tubercles or 'PIIll.'\: in Wl) OYel)
thcsl have .1 bulb-l1ke protoconch .md mar!,."lnal c.ompresscd form~ there may be a k eel In such gtn· erol VJ
Mphuncle. arc very .. inular to thml of Jnunonotd' in cr,l J\ the Jurassic fo.:o~moccms such development' .u Poleo1
.1JI respects othe1· than coiling, .md like them have c.trncd to an extreme, and in .tdditlOn I,Jteral lap-
no c;unt·ral depll,it~. pets arc present on l'it ht:r side of a compn.•s,cd Jp~r Non
A complete morphological \t·rie., fmm the Lower ture (Fig. ~US). Such lappet'i :m· rc\trtctcd 'tn I 011SI
Dcvonian Humri.irk\chietcr of the Gcrn1:tn nucroconchs. IIIoit

Rhindand allow' a ~t n1cmral sequence.: to be traccd Such primary ditferrnces in morphology cle.uf,. rnadt
from straight ti.,rnt' to loosely cmlt·d .tnd then eo dmtnb"llish anunonotds from thmc.· nf n:\lltiloid. !J
If I\
(a) Ma ntlcocere• (c)
dorsal
tmpressed
M + - - area ----1-...,....."
umbthcal seam
outstde umbtltcal- protoconch
dt<Jrnetcr
I
f
l
1
(d )
t umbtltcal
s
a
y
y
IC
;r
e.
(e) M/magoniatites
lC
1e
ar
sal
he
:ks
IU-
OSt
(I)
>tal
ted
ten
up-
the figvre 8.24 (oHdl Monticoceros (U. Dev.), a goniatite exemplifying ammonoid structure: (o) ventral view; (b) cross-section, (c) lot-
'Cry
mview with shell partially removed, showing internal mould with sutures (oil X 0.7); (d) sutvre line; (e), (f) Mimagonialites (L Dev.),
avery early goniotite With perforated umbilicus and exposed phragmocone - the shell is partially removed to show chambers; (e) lat-
;en- 1101 view (x 5); (f) sutvre line [(aHdl Redrawn from Miller and Furnish and (e) , (~ after Schindewolf, all in Treatise on Invertebrate
are Poleontology, Part L)
.ap-
per- Nonnally the ~ofi: p.1rts \.li .1mmono•ds h.we been piCture ~~ generally corn·n. th~: hiolog~cal affinities
lto COJISldered to bt• b.l\tc,,l)y mmbr to those of nau- of ammonoids arc clmer to cok01d~ (dibranchiate
lilotds, and ammon01d reconstructions have been cephal opod~ such as ~qmd,, c uttldi'>h and octopu~es)
~arly llllde very brgeh: on thts b.1sts (e.g. Fig. 8.21b). Yet than they arc to .'\·mwlus. R adul.tc have been
oids. Ill' generally hdd (l:.ngescr. 1996) that although thiS found m a few .lmmotutes (Ntxon, 1996), and the~c
240 Molluscs
(b) Dimorphinites (c) Longobardites (ovvc:onel

(~phaerocone)
(11
llt;l
{Cl)
wh
\t t•
\er
(d ) Ca/ocer.s tserpenucone (e) Spi nlkosmoceras (f ) Grossou11rla (g) Dlploc•r•s
planulatel
11\,\
.Ht'
.tb(
lllll
the
\\ h
.ll
bt•(l
llldl
Spil l
Figure 8 25 Shell shape and morphology in ammonites (aHd) nomenclature of dtfferenl shell shopes, (eHg) aperturol modillat ut 1
lions (lappets and spines) tn Jurassic ammonites. (Origina l drawing by E. Bull ) c1.1,:
11101
hJvc 'l'vcn teeth 111 l'.ll h trJnwersc row, as in anthnuc '>plral tram tht. protocondt, but in ,omc I.Oll
tht: Rct:ent dtbr.llldlt,tte l'l·phalopods, whereas PaJaeozo1c genera the ~hell 'hape may be runoml) the
J\-cmt1/us h,,, nmc. In .tddiuon, ammonoid upper quadnlatcral or tnangular. A\ the shell grow' from lCp)
J.''"· \dH.:re knm\ n. t:orre,pond closely with those the apcrrural end rhc stphunck grows with 1t. J'hc t1u11
of nxcnt octopu'e': titrthermorc. there are known siphuncle ~nm a~ a ~ma.LI bulb {the caecum) w1tlun rt''P
.tmmunotd ml 'll',, \\ hll'h the Recent l\:a11tilm does the protoconch; it ~~ lllitt..tlly th1ck rel,ttive to the llllll'
not luvc. ~ud1 C\'tdencc. though .tdmittedly slen- firsr-fom1ed chamber\ ::md 1my w.mder .tbout 111 1~
der. 'nggt'\t\ tltJt Jmmonoid' and cok01ds stand position. After the til"\t one or two whorls. hOI\·
r.ttlwr d<lSt' to~ether phylogenetically. whereas ever, it settles dow11 tnto it~ .1dult. nonnally ventr.~
f'{at~lifll, lllJY bt• liHllt' d1~t::unly related position, occupying relatively less space as the shdl
grows.
Morphology and growth of the ammonoid shell T he sept:1 were probably secreted lJl rapid growth
Ammonoid .. hdl~ ,1re characteri'ltically tightly coiled episodes, like thmc of Na11tilm. fol lowed b> rt'mnc
m a pbn1~pir,tl I:hhwn. Tht:y fomt rolled- up cones peri od~. an d the part of the shell where the anurul
of which the t•.Jrlll'~t-fimned pJrt i\ a ~maJJ bulbous :Jctuaily re~ided (body 'h.tmber) grew progress1vd~
dLp)otJal prutOCllllCh. rim WJ\ prob,tbly initially htrther from the centn· during the sptral grm\1h of
mh,tblted by a piJnktoml larva The protoconch is the phragmocone.
usually locJtt:J 111 till' rcmrc of the shell. but in T here arc ~ever:~l useful tem,, for descnbm~ tb<
occa\lonal L.lrl) t~mm (c.: .g. F1g. 8.2-te) the C"mling i~ fin::~l fo nn of the ~hell {fig. 8 25). Thm~ \hC'Us
looser so th.tt there I\ ,m umbtlical pcrforacion which arc mvolutc h.IVe the l.tst whorl covenng all
between the pwtmwll h .md the first whorl, like the preVtom whorls: 111 evolme shells most or .ul
Figure
that Jf thl• early cu1kd nJutJioiJs. The \eptate part the preVtou~ whorl~ are exposed. The umbilicus Trios)
uf clll' ~hL·U (phra gmocon e) u'ually grows in a log- which 1~ the conc.we \urt:\cc on each stde thruu• 'n Tre<
wb11h tlw 'ptral .lXI\ rum, 111.1\' ht· .ummt flat or in life (Strid,berg, 11)90); tlm prub.1bly will be done
ti<'cpk 1'xrav.ncd l'hl· ,!Jdlm.IY lw of normal rather in tururc.]
iUU<.'ned fonn (planulatc), n-ry rumpn:,,ed (oxy- The curious lOmprcs,wn of ~ome .tmmonoid
conl.'), \'t'l) t:tt .md mfl.ttt•d (cadiconc) or almost aperture~. and thetr ht'quc:nt ornamc:•Hation b\' bp-
globullr (sphaeroconc). pets or spmc,, have alteady been mentioned ~ome
ome ammn11md' I ,1\'e prunounu:d vennal ked' of the more pecuhar tonm .trt• llht,tr.Hed here (Fig.
fg '25:1), wlur h 111.1) be blunt or 'harp. The ked X.~Se) . bur rhe\t' .1n· umt,u.tl .md rhc lllJ.)Onl)' of
Dll) hl\'l' .1 .. in~;k, duublc or tnplc parallel •·idge sy~ .unmonmd apcrcur~'· .md t·~pt•ct.tll} thme of
trtll, nmning .1long thl· \l'ntcr. In t:W"-'ecuon the Palacozoic g~ner.a, .1re \llltpk hke thme of ruu-
whorh nuy t.tJ...l· 111.111} foun,, \\ hid1 ne often char- riloids.
ctcnsnc of p.trttlul.n .11nnmnmd t.unilie~ .mJ thus · In ammonotds of v.mom J..:imh the body chamber
1 'I)' lh fi.1l in t.tXononl\ m:l) he Ion~ or short. ,1\ dt\l.:ll\\ed in the <.cction on
Tht 1JLHt'r 'urf1t·L of .111 .Ulllllllnntd shdlt, mually buoyanq.
IJJrkt:J by l.llnt ~Ill\\ th lmt·'· but JJJ .tddttJUn there
t!t' u,u,tily nb~. r,tdr.dlv ,HtJu~cd and projecting Ammonoid suture (Fig. 8.26)
b<ll'l.' the surtaLt' I hne ,trt· ~ll m.my p.Hlt'OlS that Mo~t :mtmomHds ll"L' pn:~erved as intern;!] mould,,
,,nlv J few c.tn bt· iliuM 1 ,tlcd here ~omc nb~ go over ,md in these tlw Jtmcnons betwL'L'n wplJ and shdl
'hi \'t'lltL'r; other\ .ll'e IIILL'rrupted; 'ume are str.tight walls ~how up rk.1rly .1s \Utun· line~. which in
,,lnl~ltltl1l'r' rune m br.tnlh; th<:y m;ly be united ammonmds .tn• .tlway' more l"OIIIpkx than in
1r nodes; and \hortt•r rib~ n1.1y be intercJiated nautilotd~
IWt't' longer une~ .
1 In the eJrly .1mmonoid' (Dcvom.u1 .md C.trbon-
fuberdt•, m:~v t' rupt on point' on tht> nd~e or be tfcrom) rhe \tmm·s 'Wen nomully of .1 fa1rlv ~unplt:
~dt'pt·ndent of rhc:m: thev m.ty ht' prolonged imo tom1 Jnd l.irk .ttt.L'"ory Lrenulauom. but by
pmr' or united bv H.u node' . l ht> c:xrenul parrem Pcnni.m tllllC\ \nnlt' genera were ,howmg more
:o- <inb' -and ,1\\0l i.Ht'd \llllltlln.•, i' ot ~rrc.H value in complex \Uture' of .1 kind th.tt reathed the1r full
LINficauon .md idenntil.ltlon. but 'uJCc: homco- ftowt·nng in rlu: Mt·,ozoic. ''hen g~nera with
morph\ ~~ .tlw.t\ ~ .1 po~'lbtlny. rhetr use mu~r he cxtrt•mdy complex \\llUrl'' wt:re the nonn.
ne roupled \nth due ot ll[hcr cll.lr.lctel'\, particularly To dant)· .1mmono1d sutur<: morpbolog}, the
~ly !he n:uurc: of rhc ~urun: . IMo't tllmn .tnom of fos~il sumre' .1re mually rt:prc,cnted graphically.
)Ill crph.tlopod~. tndudtn~ thme hen:, follow tradi- Neglecting surtJn• t:onvcxity of the: JlliDJonotd
he on:u pncnn· .111d ,Jtow the ~hc:ll up~ide down rn ~hdl, the suture' .1re drawn from the venter to the
1111 ~peer to m li\'lng pmtnon. lt would .K£U.llly be umbilical seam (the extnnal \uture) and thence to
:ht: much more ~emthk· tn illmrr.ne them ·'' chey were the dorsum of th.tt \Cplurn. Involute auununoid~
lt.~
w-
rul
lcll (d) Oxynotlceras oxynotum
(c) Puzosill
vrh
ing
mal
>e1y
1 of
(b) MHitoc.ras
the
1ell<.
pll
ll of ·~ 8.26 Suture morphology in ommonoids. (a) goniatitic suture (Neog/yphioceros; L Carb.); (b) cerotitic suture (Meekoceros;
:::.u s. '<ios.); (cl ommonitic suture (Puzosto; L Crel.), (d) svlurol ontogeny in Oxynoticeros oxynolum (Jur l (Modified from various authors
ugh 11 Treoh·se on lnverlebrote Paleontology, Port L.)
242 Molluscs
havt> a d.tsnntt mrem.tl ~uture, bur 111 the evolmc \uturc' m all ammonouh are tar le~\ LOmphcated
fom1s the mtenul suture 1s small and umgmticam than the htter one~. and the whole lmrory of devel- ~pen
and i~ nonnaUy not drawn. Since the full suture i~ opnwnt of a mature suture can be traced ust ng senal pcrfc
symmetrical abolll the dorso-venrnl a xi~ only half :1 diagrJms of succcs~tve mtures. Such ontogL'ncm lllll\1
wptum IS nomulh drawn. Hccamc no account i~ \Cncs have sever.!l use~. They have been ll\ed. for llun
tal-en of the surf.lCl' l·onvex.iry, the \UttlrL· dtagram j, ul\t,mcc, to distmglllsh homeomo'l1h<; 111 whtch the IUIU..,
only a proj~c:tion, '' h~eh can never bt.' L'llrtrely accu- mature <;uture m two dmantly rellted ammonmd! de m •
rate, but thl\ i\ llllllllporrant t{lr purpmL'\ of compar- look' ~mular but has been amved .lt ontogcnclll'all) phu'
Ison of di!Terent \lltllre lines. Such th.t~rrams .uc not tn quite different w:ty,. tphl
only useful l{lr rnmp:~ring the \Uture> nf ditlim:nt Pt>rhaps the greatc't value of smur,1l ontogl'ny " bona
'Pt'cie' .md gcner.1, but Jlso e~senri.ll in tracing tht' in unravdling phylogl·nics. especially m Pal.teozoll ll'(.k
chang-es 111 '\Utur.ll morphology throughout tht' fonns, for in these .1mmonoids the c.lfhcr 'tagl\ u .\i,/11/
wowth of an indlvidtuJ. which hJ\ \'.lnOU\ uses. development of an 'ad\'anred' ~uture do'd} re~cm a mm
In drawmg .1 \lltllrl' the venrer j, ,Jmwn on the blt' the m:Jture <;ept:.t ot .l more primltlve .unmon01d (.lgJi
lcft-lund \tdc with .m liP'' ardly directed arrow type. Thus ancc<;to~desrendant relatiomhtp' t.w bl S.J), I
pomtmg tow;mi' tht· .tperturc; rhc umbiliLal seam i~ pmrul:~ted. and phylogenies have been drawn up on dcptl
'hown as ,t curvm~ ltne, and anothct vertical arrow th1s baSls which h:wc latn heen shown to hl)ld 1trn1 Whl't
nurb the dor~un1 on the right-hand side. 111 the light of more t'videnre. h,J\
lnlkcnom on the 'uture line pomrtng upwards (the Uuring ontogeny, in Palaeoz01c .1nd Me~OZI)!C" 01111111
apertural direcnon) .uc the saddles (l·.mlv rt'mem- genera. primary lobe\ .1nd saddles .1ppe.1r early, :mJ t\\ u
bered ~ince .1 hor...t.' \ ... addJc tacc' ton\ .mh). wherea~ nonn.1lly the tirst and ~econd l.neral \addle\ arc very Stren:
the b.Kkwardly pointing inftct:tl011\ {t:lllllg down- dt\tmct. Though rhl·~e per...i~t mto adult 'uturc, 10-1
w;lrd~ on the Jiaf:r.1111) arc the lobes. adventitious lobe' m <oaddle~ may appe.1r tn t.1vc
The ~uture ~~ ntmt important 111 taxonomy; between the primaries .md cvemua.lly ~:,rrow ,\\ l,•rg~ lll.tlit
p.lrticuJar kinth of ~lltllfCS C h,tr.lllt'nZe disnnct a~ the pnmaric\ rhetmdvcs. The only w.w to distin- l'hyll
ammonoid t:!mtht'' .mJ are vcrv meful in dassifica- ~'lliSh whtch tS which i' then to tr.l<..l' them bark to ~rout
tion and !dcnttfic.Jtion. There arc 'ome broad and rhe earhe\t omogcncnr \t,Jge~. low '
gencnl ccmt\ ddimng dffimonoiJ group~. based on In the ammonHl'\, wht•re the ~uturc readtes 1:. I ht:
-;urural morpholu~y. \duch arc ottcn med to char.K- 111JX1mum complexity. the lobes and \Jddle~ are ill Lye m
tcriz~: c:hfferc:nt gt alk~ of orgamzatton. Thus the c:renubtc, and the s.1ddlc all have acu.:s~Ot) lobes ~CIIllC
goniatites arc P:~l,ll'OlOK amrnon<Hd' with sharply The terminology of :11l the~e. as gwcn 111 Ptg. tL2~d. ''her
.mgular .md generally zigzag ~ururcs, without any become'\ rather totnplcx. Some rather pecuh:u Wtlh~
.tcce~'ory crL·nubtiom. Not all .lmmonotds wuh sutun.·~ :uc prc~cm in ,1 few Mesoz01c genera. In ct'r· whit
\UCh suture' ,UL gont.ttites: there .11 e ~ome unrelated t.nn cx:~mplcs each m,Jlure ~uture is apparently tn11 · 11)'111
Mc\ozoic gcner.1 wnh ~mular sutures In the Tna,stc C:lted :~gam'ir the nnc 111 front, though 111 fac1 tht \V,IJI
cer atites the lobe' an: fulled though the '-.1ddles arc 'mt\\mg' parrs contmue below the preceding )t'p· 'liKC
entire. Some cunous Cn:taceou' genera (pseudo- turn Septa m.ty be ;t<,yn1metrical, and cwn taxo- .tlld e
cer atites) ha\ e 'mular sutures. l'hc Mcsozmc nomit:.tlly unstable within the one genus. lh
ammoni t es Jll h.1vc fmdy subdivtdcd and complex •>I thl
lobe' and saddh:,, though there are also ~ome Siphuncle, suture and buoyancy hdi\.·1
Penlllan ammono1d~ wtth surure' of tht~ kind. Ammonoid ~hcll'i an: .dway~ thinner than the ~hclls •l"flC.d
Thou~h then: j, thu~ a gener.tl \tnt1graph1cal of 'ctutilus, their ~1phundes are \'emral (except m tlnl\
mncase in surur.ll rompkxny. the progre'>'!On from Order Clymcniida). the septal necks .uc nonn.tll) shell.,
gomatite through n:ratltt: ro ,lmmontte '" not. how- prorhoanitic 111 ammonoids and retrorhO,Itlltll 111 (.tJur.
ever, a direct phylogt:lletic line. n.tutiloids, and the ~cpt.l 111 ammon01ch are U\u.illy \lllllll
l 11 many Pal.lt•ntmc and Me~n70tl ~mmonoid~ convex, not concave, towards the aperture. hl'l-.1~
the ontogeny of tht: Slltltre h;l\ been worked Out lt i' gcncraUy hdd th.tt .umnonotd~o .H:httwd th.Ut
(Fig. l:>.16d). normally by breakmg ofT the chamber; buoy.mcy in mu<..h the ~ame way as doe~ \.,umlu) ,'\,llltr'
ont: bv one as t:lr ·" the proroconch .llld drawing the .md that the fluted \epl<! amongst other timcoom Slltllrl
\Uturcs as the 1nner whorls are cxpo,ed. The e.1rly m.l} have increased the ~trt'ngth of the sheU ~o thmt n J'lH
IU.Slbk• ro rc,t\t unpl<hion .H d~pth. But to be more Stanley (I 971) and b> Kcnnedy .md C..obb,m (1976).
lptciftc th.m rht,, and most particubrly to be able to One problem, howcwr, i' that beyond a certain
~rfonn mathc:maltt·aJ JnJ!ysts, variom other tactors lcvd of complcXJty. inc;rc:J\c:d septal Auting
mu~t be known, indudtng the nature and composi- decrt:a\cs the angle bt•twecn \t'ptum and ,bcU. Thm
non of the ,jphumll·, tn 'ee if the ,heU could have the shell of an ammontt<.' with d \'Cry complex
funcnoned hkt· th.H ot .\,mtilrts. 1 here i\ good evi- suture may be more prone to t.l.mugt' through
lknce from rare unaltered \pcomcm that calcmm bendmg strc\sC' than one with a less complex
pho~phate t\ the pnm.Hy constituent of anunonoid suture.
11phuncks. thou~h .fll\'t•m)e, havt• more: caknun car- ~everal altemativc functiom h.1ve been proposed,
b<matc .md proh.tbl> the pho\phanz.mon was pre- which mvolvc phy,iolog-tcal r.uhc1 than \tntctur.ll
ctJrd by a ca.lcmc \t.lgc: 1 he cakmc outer tube of ne'cessny. Ho"'. tor C\..unplc. couJt.l the ammonites
\1111i/us ~~ mtmng. N(lW in ccrtatn Mesozoic have rnaintatncd ,, \ecurc attadtnteJH of the body
•mmotutes the strt•ngth ut' the ~tphuncular rube muscles to that \hell during episodit forward
Jgatmt nplo~10n) In~ bct•n cJ.kulatcd. Needless to growth? Were the'c mu'otle~ ,ttt,lt hed to tht' sepwJ
\ly, tht• tube strength t\ .In tmport.mt hnutarion on ftutings, and if so nught the !.mer have increased the
.kpth. Tht• rebnve ~trengrh mdex (Ill r) X 100, sur£1ce area and thus providt'd a better •Ht.lchment?
11hert' h 1s thc: w.tll thtd..:ne~s ,tnd I' ts the tubt· r.1dius, Or w:ts the pnnctpal li.mnion of the seprum to
h.h been c.tlcul.ltc:d for st•vc:t-.11 Mesozoic sprend the ellhts of hyt.lrmt.Hic pres~ure evt'tUy
mmonotd~. Jnd tt 1~ mmt tntt:rcstmp; to ~ee tint acro~~ the shell? Wt'rc th~: ~eptal Autings consequent
rwo ord~:rs, Phyllocc:ratld,l .md Lytoceratida, h.ave a upon the form of dw org-;um template lrom wh1ch
rrcngrh mdex hke rlut ot i\',llltiiiiS (11/r X 100 = tt was calcified (intti.tlly fnn11ing a visrou~ tinger
111-19). whcrt'.l~ tht·tr dertv tttVt'\ the Ammonitida pattern), itself ju~t a by-procht<.t of geometric
n lul'<' a \tgmhcmtly lnwer \,tlm· (.1-<l.S). This infor- nccesmy? Or w.t~ rhe t"lmctton of the Auted seprum
e uttnn tccord, 1\ ah the: r<.'(Ognmon that the connected w1rh the removal of lJmeral liqutd?
PhlliOll'f.tnd.t .md l yton•r.md,\ ''en.: det•per-watcr Po,~1bly 'ome or all ofrhe,c .1re v.1hd 1f p.1rti.1l expl.l-
0 gtlUP'· whme cJenv.mw~ repe.ltedl~ invaded shal- narions. A recent re\IC\\1 of thh \\'hole mue, ba!>ed
\\ \Uta by .1 d.t"tt" prot c:" of Her.nwe evolurion. on phy~tcal and Cllb'1nccnng pnncipJe, (lle'\\;tt .1nd
Thr dcul.mom 'how th.a the Phylloceranda and W csterrn:mn, 19H6, 19H7) stresses the adaprarion of
L)'ttlC~rand:t umld \\ ttll\t,llld a w.\tt'r column of ammonotd sepra to rc~tst hydrostatic lo.1ding. whtch
!S. me 45• I m, .tbom the \.tmc a\ that of '\'autilus, could be applied both VIJ the: body t ham her of the
d. ner~J\ Onkr AtntnOJll(ld.t could probabl} not thin ,hdl wall The rc:l.ltt\C mAut•ncc of e.1ch of
1ar th,r;md t.lc:ptll' of more than lOO m. Other data these would depend upon (I) hallltat depth, affect-
:r- 'bKh have bt•t•n mt·J 111 .mc:mph to mfcr bathyme- ing hydrostatlt pre~mre, (2) whorl prnfik aflecung
n- 1) m ;tmmonmll\, 'ud1 ·" the rhtckncs' of the ~hell stresses in the ~hell wall, ,1nd (3) the artu3! ~tze of the
.he ~~~11 .md of the \t'pl.l, h.tvt not proved \O useful whorl, wluch would atTect bending \tres'e~ m the
:p- Hlle rhl rclJtmmhtp\ in.,olv~o:d arc wry complex shell wall and the fluted ~eptum. Thm a ctrcumfcr-
ea- tJ dudt: 'imp le an,tly'l~. ential ~ upport t"lmction is prinJJry lor the .tmmonoid
Tbcrc hJ\ bc:c:11 llllH. h dl\tU\Sion of the function suture, and it must havt• opt•ratcd in the e.trlic:~t
!the comp]c, amm<HIItc \Cpllllll. Many authorities growth ~tages too, as wottld be ncce,~ary in swim-
dievc th.tt the pnm.try ltllll.'tmn ut tht• convoluted ming animals adapll'd for o!Ehorc breeding.
tell-; ICj'tal pl•npht•ry WJ\ l\l gtH' till' .;hell extra strength. In ccphalopods generally. and undoubtedly in
t Ul 11 prcventmg implo~ion ,\t J<.pth. Ammonite ammonoid,, the main adapt.ltions were concerned
till) .be-lls ar<. tlunner th,111 tht)\e of ruutilo1d' and the wnh csublishing ,\11 dfl-ctlvc, neutrally buoyant
~ Ul .dor.illv wnvc') 'eptum ,.,uuld hdp to m~tain poi~c of the livtng anun.J) 111 the w.ller. Such 'truc-
lall) ::ong pn:"urc~ bcc.nl\c ol tt\ flutmg. Moreover, tur<.' .1s node,, nbs, tuben.lcs and 'Pllll'\, a' well as
.6UIC of dm mtem.tl buttrt ''mg 'heU shapes other sepraJ approximauon m mature mdtvtduals. may
~vcd Wl tht: st.mdartl ov.ltt' 'hell. 'uch as that of have been concerned ,.,ith tinc:-,clle tnmming of
uilrts ~li/1u, <~re po"tblt·. The Yit·w that anuuonite the pobe. Perhaps paradoxtl•.tlly, external ornamen-
nons ~ fum:tiom:d m.unJy <J\ prc,~urc re~mof\ seems tation can actuall} red ut c drag. by mtroducing tur-
1;\t it JlOnable .•md '' the model pn:tcrrcd by Raup and bulence 1nto the boundtl)' l.tyc:f\, ,1\ With dunple~
244 Molluscs
on a golf h.1ll . 1 he h Jrmt.ltJC .ldjU\tmcnts of thl that in ~ome .tmmomtc<> pairs of ·~pene~· cnuiJ bt H:IJ
.1mmono1u ~hell. 111 gt·ncr.Jl tcnll\, prob.1bly worked distmgm~hcd, dlfft•nng only m the character ofthell tllll
,1, 111 Nt~wrlrh, ·'' d1d by\tt•re'i' llll'l' h.nll'm' whiCh Outl'r whorl~: rhc nudt'l could nor be tOld ap.m hkc
.tllmo, c.:d t h.111~c' 111 hquid ,,.1th111 tht· <h.mtbers to W.tagen. however, d1d not bcheve th.1t \t'X bt•
tJkc pl.J~.c nunp.1r.1t1wly ,Jowly. Thl'> pcnnitted dimorphl\m could be invoked .1~ an cxplanaoor pof
,Jtorr-tcrm t'Xtlll'\llllh nllo dill~ rent k"l'l' of the 'cJ and 1t "·'' nt'Jrly IIlO }'C'Jrs later that Callom (Ill
and JlltolliJCit rc.:turu to till' \t,trtlng Je, d (W:1rd and (1963) Jnd M.tknw,ki (I %1) com·incmgly dcnhl!:· ~011
M.nnn, llJ7~. Hou,c, IIJHl). \tratcd tht tnten<l th.tt lOttld hl' u'cd in e<>tabhshu \\1tl
dimorplll\m unequJVOl,tlly. )mtc then nun Ill.\\
Sexual dimorphism (Fig. 8.27) Mesozoic dimorplm p.1m h.tVI." hl'l'll deJrly JJ\·nn- I eh
The qut'\tlon ol ''het her or not amnwnoic:b \\'Cre tlcd, ,1nd there .trt 'poradtc rltnni~ in P.t1Jt'i1ZOX. l':~fl·
\CXU.11ly dunorplllt h,l, bcl'll dchatul ,lt great length 311UllOIIOilh too (D.l\'1\ r1 <1/ •• 1996) tit tll
for J very kmg nmc. A~ l'.trly .1\ I ~69 Wa,tgcn, one In trying to work out dnnorph1c pJirs 1t t\ C\~ lt
of rhe c.1rl~ Gcnu.ln p.tlacontologt'r', pointed out ti,ll that all the .1mmmme ~hdb \tudied ,trt• nwur wlw
th.tt 111 ,lllllllllllltcs ut' F.unily Oppcli.idac (Order 'pccimcm, i.l'. ,\dull ,IJdls 111 which b'l'owth h:ld <(Ill'S
Ammonit111.1) thcrt" wcrt· .tppJicntly two parallel ceased. Such lllJlurity 111 ammonite 'hdls I\ m.uxcd h.tvc
phvlogt'lll'flt hllt',lgc' dcvdopmg throughout the by (1) crowding (.1pproxim.1ting) of the 1.1~1 fe~ lll,l!t•
Middlt· .111d Uppt:r Jm.l~sJc Furthermore. he found septa, ULie to dimimshing growth rate~; (2) J cban \ltnv
111 the cxtcmal sculpture near thl· .1pcrturt' -tht' nl>- IMgc
bing nuy be dttTen:nt, m the .lpl'rtllrl' ms~ bt ~<Hll(
constri~.tc.:tl or m.1rh•J by l.tppc:ts or honl\: ltld <"ll~r:

~omc ~lig;ht unc(lllin~ of the.: bod) chamber from tht s.,
rest of the shell. N~H diJ these k•,Jturc\ arc lound dt':.'\1 r
.:mv one .1mmonttc, and mdecd 'omc of them at '' irh
contincd to one wx only: nl"Vcrthelc,~. thc pr~ a~ d
of only onL or two \Uch c.:harJctc.:~ will ciUblc 1 b'T'OUJ
matUre shell eo bl' dl,tinf~ui,hed MO\t ammonoid logic.
shell~ found arc in t:\Ct maturt'. and 111 ~pcarn Qlllll'
collected from the same bed mature; \hdb of u.
same kind arc normally very mud1 alike 111 'lt<:, C'JII> '
When mJturc ~hell' from thc ~amc bed and loa:· 1 dl'a
iry are separated m to \pedes'. they often cornC' out IIOl I·
into t\\0 qu1te dt\ttnct ~'Toups, Without llltCmlcd:- grc.tt~
ates. fhe~C oll'C dmtngutshcd pnnunJy by SIZC, OIIC tn·di
group bemg two to tour tunc~ l.uger thJn th~ othc 11 Ul'),
so th,lt the two kmds .~re gcncrally referred 10
micr oconchs and •nacroconchs. lu nulrOCl1ndll Am
and macroconch~ from the" s.tmt' horizon. the mr.a ( lllllJ
who rl~ arc normally 111dtstmglllsh:~blc and only 1ht 111Ulll
outer whorl<\ dtiTer m numbe"t, ~lZI!' :~nd morpbol aptyc
ogy. If there ,trt' .111y peculiar li:atures of the lp:r (Hg. I
tu re. the~e .ue nornully found onl> 111 thr tllllg p
microconchs, whereas the lll:'IL rot onchs ho~w stmplt thl· .tr,
apem1re,, and the mor~ extr~me ca~es ot J~uch ·~ Ill
rnent of the body t hamber trom the prC"<cdJ rct.un
Figure 8.27 Sexool dimorphism in Jurossic ommoniles: (o) whorl are hkcwt\e char.tct~rhUC of the microconc \oiJh'l
Pensphinctes (Arisphmdes} ingens and (b) Perisphincles It t\ theoretically po~o;tblt: that the Jpparcnt se \\ h~:re
(Dichalomosphinctes} rotoides, both rrom the same horizon in the p.un .uc merely cJo,eJ) related taxa h\1ng 111
(Ofollion (-.: 0 15, these ore probably dimorphs); (c) plctt•l>•
Grophoceros cO\IOium and (d) Ludwigma cornu, o Bojocian <;,1111C plate Jt the S.lll\C tlllle, but \lllCC tn \Urn lm. ch.unb
d1morphic pair (x 0 65) (Redrawn from Collomon, 1963 ) cage' new ch:.trJcter; (~tll: h .1\ moditic,ttion' 1•1 CXU1 ud p.t
JUI sfulprunng) .1ppcar in bnth group~ at the ~amc There is no questtnn but that thl'w were part of
nmc. then sc:xtul dm1orph1'm app1.·a~ to be the only the ammonite ..and Trauth, working m thc 1930s,
hkely po,stbthty. llowcvcr, .:~11 thc'c cntcria have to was able to l'~tabh~h th:ll ditlcrl'nt type' or :.tptyclu
be med Wtth c.1utmn 111 IIHl'rprcting ammon01d bdong~.·d to dilft·rcnt .1mmomte group,. lt wa' Ius
population,, f(,r till')' d0 not alw,\y,, by any meam, VlC\\, .lt tlut time. th.H aptydti wcr1.' some kind of
occur m p.11~: th1.• ~.·xp~.·~.·r~.·d ."\11;511 ratio i' only
opercula.
romenrn ...., found, .md m 'nmt• c.t'c' the van.tnon There .1rl' .1l,o anaptycbi, wh1lh (Oil"'t of a <orn-
Wtthm .1 pl)pulatlon collected from .1 ~mgk localiry glc plate or clutin or other org.lllll nutcnal. nor-
m~y b~.· l'tmnnoth. Ht•nre we h.tVl' tu .\b'TCC \\'ith mally flattened and in J buttcrHy ~lupe. Aruptyclu
lchm.mn ( 1971 b) who .1djure' u' 'to be extremely arc confint·d m scdunelll' of Upper Dcvom.1n to
~retitl when it ~,·nml'' to m.Hnt;limng 'exual rela- Lower Jur;mtc .1ge •.n \\hich 111ne the e.1rliest aprv-
rionsh•P' betWl'l'tl h1th1.·rto blamde'' .unmonites'. du appeared. The IJst .lptydu .m: toumi with the
lt i' h.trd to I.. now wh11. h murph •~ male and last ammonites at the end ot the Cn.·tateous. Jmt
whiCh j, f\:nuk. ~Pilll' .lutllllf' havl.' romidercd the wh,tt the ;1ptychi were .111d hnw they fi.mlt1oned
rem::tined .1 martcr lor ~ome dr-.pute. though most
qu1.·stion ~o .11. ad~..·m11. ,1\ t\) be un.lllswerablc; others
h.we sugge,teJ po~lttvdy tlt.1t the 1111croconch is the palacontologi~t~ bdieved th,H they were opercula
male, and the lll.ll roconlh the t\.·malc. Nautilm and that anaprycht wetc ~ome kmd of.1aw ~tructure.
1hows little ~..lunorphi'>lll (thl' male Js ~omewhat Lchmam1 (J 967, 1971.1,b, Jl.)79) finally settled the
brga and bro.tdl.'t) Jnd i-. thu' no great help, but nature of these ~tnrt rttrl'S by ~ttld)'lllg exceptionally
llllllt.' Mesozotl n.Hillk)Jd' h.1ve beu1 described as well-preserved material .1nd 111 p.lrtlt tlbr by making
(kdrly dunorplm serial sectiom. I k ~hnwt>d eo m lus1vcly that th1.·
~mce .111 km m n dimorphic pair\ were origtnally anapytchus i& ,, lower jaw, nri~'ln.tlly V-shaped but
1Nnbcd as scp.tr,Hc 'Pcl'il'\, what should be done usually prr~erved Aancnrd and \pre.1d out. In addi-
with them t,lxononu~.ally when th~.·y arc recognized tion there 1~ .1 beak-ltke upp~.·r J<lW of tht· same
~ dimorpluc; ">h<'uld dimorphic 'spenes' be horn) materi:JI. very <;tmibr to thl' jaw' of modern
1 .:rouped tngerhcr under the one name a.s true bto- coleoads. lr is mually prt'\Crvl•d crushed on top uf
j ~'!c.U 'pecic~. or 'hould the l''>tJbh.shcd names con- the anaprychus .md l1.1d not prrv10mly hccn noticed
s tmu( m usl.' .1' 'm or pho'pl'l'll'~ '? as a sep.mue strucrurc. Aprycha. .lt'conling to
e Undoubtedly rhe tonm·r pralttce is more techni- Lehmann. .we C;llc.lrt'Oll' depu\it' v>hich lun·
grown on the cxrcm.1l (wmr.1l) ~urtJCl' of thl' lower
C'lll}· corrrn. bur thl· Ltrter 1s perfectly adnussible tf tt
.- llcll'ar rlur we :trc ulkin~ .tbnut ·morphospeCies' and J<lW, .1~ •l parr of thrclerwd pl.tte\, .md the,c did not
1t nor b1mpcnes, :tnd it do~.·, have the .tdv.muge of develop, cVldentl't, until dw Lower Juras,rc.
l- \;~'mer flcXIbult\'. for rht· moment. p.1laeootologim Lehmann \ mterprl'tatanm h.1Vl' been nmtim1cd
Ie ue div1ded on tlm lJilt'\tron. though the defimtion of and extended with rht• dhCuVl'ry of rnmpletc jaw'
tntC biO\peCJCS ::tppl',lf\ to b1.• ~Jllllllg ground. m other jur.l\<;IC ;lnd Cret.Kt'Om JllllllomtL'' In all of
as- these the j.tw ~trurnm.• ts rcl.lttwly hkl' th.lt nt octo-
hs Ammonite jaws puses, hut there IS on~.· unportant ditll-r..:nt e. n.u11cly
er Commonly found rn .1s~on.1tinn with Mesozoic thar the lower pw IS vcty largt•; in rehtion to body
.unn1onites .tn• pa1rcd l-.tkirir pl.rtcs known as size tt IS about four tlllll'S largl·r m the ammonites
aptychi . wlm h ~llpl-rtin.tlly re~emblc bivalvcs than in any modern cokmd. To wlut extent thrs
:r- f-ig. R.21c} Aptycht :~re mu:1llv \l'.lttered on bed- was related to the Jllll110mtcs' dJL't (probabl)
he ding pbncs where ollllll1ll11Hl'\ .In' abundant. Where slow-moving live anin1,1h .md urnou) i\ quitl'
)le me Jr.tgomm ~hell-; nf the Jll11110illtC\ are crushed, unknown. lt now seem\ ltkd) th.at aptycht ti.tnc-
·h- m so nuny Jur.l\\11 ,h,de~. the calcJtt: aptychi rioned both ,1\ klWL'r J.IW~ and ,1\ opt:rlltla (Lchmann
ng rwin tht·ir onh"ln.ll, sh~hth curvtng relief. .1nd Kulicki, I <JIJO). Tlm would l'L'rtJmly h.wc been
hs. Someomes no .llllllllllll!l'' .1re ti1und, only aptychi, po'>sible 1f the ammonltl•\ lower j,l\\' WJS rebnvdy
ual where the ar.a!-tonitl' ,hl'll' h IVL bcl'n dissolved com- mob!l~: \\hen fi:~:ding. with mo't of the biting work
the Fktrly. R.1rd'r .lptyl hi h.lVl' bn·n ti1und m'ide body done by the upper Ja''.
111- ch.unbeT\ or amnHlllltl'\, meeting like double doors Nor unl} do ammomte~ po,~~.·,s J.lw~ but .1ho a
:er- md parri;llly ntrin~ the shape of the aperture. radula, hl..c th.lt of llthcr nwUu'l' with semed row~
246 Molluscs
of teeth. lt '' \L't 111 between the jJ\1.~. Such radulae ORDER CLYMENIIDA (U. Drv) 1 ht· only munou01d 1.10 till
were fir,r dt,ulVI.:rl'd 111 Carhomfcrous gomames with a Jof\al. mar~n.u "phun(k: tl1cv mJ\ have "b>oruan~ I)T
from Braz1l Ill IIJC•7. but haH' 'mce been found m sururL~. OR
'L'H'ral g~o:tH:r.t ofjur.I\\IC ,unmomres. SUBORDER GONIOCLYMENIINA (U. Dcv ): The ~heU 11 otlt
often tnJilb'Uiar Ul latu.U VIew; q~. ll't'C k!umau, I t•t
c;,,,,.,./ymtlll<l ORI
Phylogeny and evolution in the Ammonoidea SUBORDER PLATYCLYMENIINA (U De\ )· '\utul'l'' usw!!) 11111
~unple, 111volutt" nr cvnlutl'; c:.14. Clpllilllil, PfilfJ ''J'':ttll'- .mu
{Fig. 8.28)
ORDER GONIATITIDA (M . Dev.-U . Perm.). Ammunmd'" e.g.
Tht· follmv mg order\ h.we bt•en di~tinguiilied m the \eh/
venrral marguul '1phu11cle. umJUv prodJO.lllltiC b"'ilUt:
Subd.h, Ammotl\)tdea. suture of c1~ht lobe,. ORC
SUBORDER TORNOCERATINA (U . Dn -cJrb., c urdt
ORDER ANARCESTIDA (L. Dn .- Pcnu.). BasiC anunonmd Timrt>ct'm.,, Clln/,,c, '''~· C.. mwd,•r/111) .t11d ')UflORDER Crc!
~tock, vcntr.ll r<•tn~<:hu.ltlUt. ''phunde bulb-like protonch, GON IAllTINA (m:unly C.ub.: e.g. G•tn«fl' '"'}
ofwn '~ 1th p<·rft.>r<~rcd umb1!1Cu,, "mplc sutlJrcs. Cwllorp11tl(l'r.1S, Ca.ftnoa-r.t.>, "''Jlollltl({'fcH) .ue ru~tlJlgw_dltd
SUBORDER BACTRITINA (U Stl. Perm.): Orthn- or cyno- pnm.1nly on the \Utufl'. The
cone, prob.1hly ·'''' c~trJI to .Ul other munonoids; e.g. ORDER PROLECANITIDA (L. C.trb.- U. l'cnn.): ShcU ,nuiOII! Jge.
Harrritt'\. with large umbtlicus. vcucral. m.trh'ln.U, rctrochu.mm An.1
SUBORDER AGONIATITINA (1 .-M. Dcv.) Loosdy to siphunclt•, ~uwrc' gon1atittt to ccratmc; e.g. Prvl.-tom•'B, ~lrai
t1ghtly mlkci, tl,lll\ltiOnJI between bactriods and t'arly Mcdlimflia.
In I
gomamcs; r.~. -lnt'ft•ft'trl.•. ORDER CERATITIDA (U. Perm. U. Tna,.): Shell oflw h1~~
\Ulll
SUBORDER ANARCESTINA (L.-U Dcv ): Serpenticone ro ornamcntcd, \·entral prochoanioc ~1phunck. reraoric ;utw~
globul.1r uwnlutc, s1mple \\ wy surure; e.g. Afrwriorrms, but some \Uturc\ att;lin grc;lt complcxny. Tlus order unde- wh11
titllli'CCSft'S. went .1 dr.unatic r.~d1o~tmn 111 tlw Tn.1mc: e.g. Cmn fnrar
SUBORDER GEPHUROCERATINA (M .-U. Dcv.): Serpenti- Xcnodisws, 1\:onres, PIIWft)(CT<I\. don
com• ro mvolut~:. mort: complex suture; e.g. ,\1anriroceros. ORDER PHYLLOCERATIDA (TriJ~.-cn·t.) 'imooth ,,, \Wkh r:tplC
luuo
sutut
early
but a
all 01
order
nuy
!-:OlliJ
<•oni:
P:tl.ll.'<
~tphtll
t) plC<\
Tht
the r
Pl'ntll.
tild so
lhL' gt
only tl
gn·.lt r
Upper
de, lmc
J'l'IIL'd
Wa\ the
to Ordt
bound:~
Shon
Figure 8 28 Evolutionary pattern of ommono1d orders, suborders and superfamilies (From House, 1988 ) I'emuat
om)m~mc:d ,fleD, \\ tth a .-h.u-acl~ mttctlh ph~ Uotd suture: 'cxplmivc:' ;llbptive radianon. Thc Tna~,ic
t)TIC"Jih mvulut< A ,-ery pem)tent •tud.;; e. ~. f>/ry/l(l(tr.ts . anunonotd~ nearl)' all belong to Orda C eranrida.
OI!DER LYTOCERATIOA Qur.-Crtct.) Ev<lltatc:. hlo\d\ ~:otle<l which h.1, ~cvcral ~upertamihe~. They arc bro:tdl}
o:w1 with 'tl.trc:,' Suture-- .:umplc:x, utH phyllmd, e.g.
known a\ \·c:r,ltJtc:.,·, and mdeed 111 mmr membel"'i
l.yr••«·r.rs.
ORDER AMMONITIDA (L ·rn.l'.- U. Crc:l ) The true thl' ~lllurc j, \Ollll' vari;mt on the t:~t.tbh,htd ccr:mnc
JrlU11\lntte-~ tlf the- Jura~qc .md Cnt.t«'<lll,, with cvpical ~utun: theme:, though the external ornament
itnmomt~t 'nrurt·. Normally courd .1ml utit•n dimorphic: became t]Uite complex in SOI11l' group' (e.g.
'·i-1· 1!lr ,,,,aas. Dcn>et'ntJ. r;,,,/it>• "'"'• l'ai•f'lrillct.-s. SuperKunily Tropitacea, whose mrmbt·r~ arc gener-
:\ilii,•!III>.Jdrr.J, I J.,pJifl·s. ally onwncntcd with strong rib' or nodes).
OI!DER ANC'YLOCERATINA (C ret.) Au cntlrl'ly Crctl<' rous H O\-vevcr, a ceramic suture is not t hararrcnsric
UI'J<:r, mdudan~ nonnalh cotkd tomt,, hut .llso all the
Cttuccou' hctcromorpln. e g. De il.rrr•rr.-.s. Tumlitt's.
of all Cl·ratitina. Indeed, as has ottcn been poinrcd
!n.-y/,~'Cr.l. , Vtli'/IIIU. out, the \Uture m Pmawccra> (Supt:rf.umly
Pmacm:cratacea) reaches a degrc:c: nt 'ammonitic'
Thr r.trlic\r .unmonoids art' of Lower Devoman wmplcxtty bardy rivalled h) that nf Jura\sic or
1gc. Thl·~ belong to an ancem.1l 'rock, Order Cretaceous genera. A fi:w peculiarly coiled 'hercro-
Anarcc~tida, wh1ch may have bl·en dcnved from the morphs' are known.
\lrJJghHhclled Suborder Bactritmil (hrben, 1966) ln the early Triassic there also aroSL' the: a1nmomtc
In both groups there arc similar, almost straight stock that was eo give rise to aU po~t-Triassic
'urures, <lnd there is a large bulbouo; protoconch ammonotds. Thts is Order Phylloccratida, an ;~\mmt
wh1ch 1s .tccommodared 111 the An.trcesttda by a per- smooth-shelled group with a charactcri~ttc 'phyl-
(or.tte umbthcus. Ncvcrthcle~s. other early genera lmd' suture (h g. 8.29a).
do not h:we the perforate umbJLH. us, .md there very Tht> lmtory of this order " intt:rc\ting. on
upidly armc more 'advanced' tcJrure~. 'uch as uwo- Jccount ot borh tts stratigraphtcal pcr-.i,tencc (it
lunon of the shell and incrca<,cd '<inuosit) of the contmued unttl the Cretaceous) Jnd its remarkable
1Uturc' lr.1ding to rhc gomannc condition. All the conservausm. Withju the group there was t:~'traor
e3rly fonm have ventral retrochoJllltll' stphuncles, dinarily liuk evolution, but its evolutional) off-
but ~ second Palaeozoic order ts dJstmgwshcd from shoots became lhc very d1versc amlllllnites of the
~I other .mm10noids by :t dnrs.tl stphunde. In this Mesozoic.
ordl·r, lht: Upper Drvon1:tn C lymrmidJ, the shells From the Phylloceratida there arm~· ,uJothrr per-
ntl\ be: tnvolutr or evolutc, .1nd the '-Utures are sistent order, Lytoceratida (f1g. 8.2%). which origi-
mum11: The thtrd of the PJ1Je070lt' orders is the nated around the T ria~sit·-Jur.tsm buumbrv.
Comatmda, which include' by 1:1r the lll.IJOnty of .ill Though the mam rootstock genera (such a~
Pabeuzoil fonm. All the'e hJw .1 prochoanitic Lyroccr<1s) 111 parallel \\-'lth th~: Ph\ollon'r<tttda became
l!phuncll·, apart from a fc" pnmttive ~enera, and little dJtfcrcntJated, they produced d greater number
tvptc.tl ~oniaritic suture line<,. of radtJllOns than did the parent Phylluccratida.
Thew PalacozoiL groups cnntmul·d throughout Both the Phyllocerarida and Lytocentida produced
da, Dcvoni.m and C.trhomfcrous .md mto the radiatmg ltneages, but the phylogeuy 1s ~o complex
P(rmian, buL rhcy underwent m.my t:me' as they and the difficultie~ in reconstructmg phylogeuy arc
did \O. During rhc bte~r Devonian, m p:uticular so great that in many cases it is not known for cer-
h~ goni.ttites wry nearly bl'C.tllll' c:xnnn and tain which of the parent stocb gave rise to which
11h the genus Tvrrwcem~· surv1Vl'd to gtvc rise to the descendants. H ence the taxonomy " unde.tr. and
gtJt wJi.1rion of Carbomferou' gn111.1t1te~. By the ammonite \ntdents have nom1ally retained a pol}-
Upper Pcnnian the goniatit~:' \\l'llt ulto .1 final phylenc (or 'r,1gb.1g') Order Ammomcida to acwm-
de<hnc. ThL·n. at tht end of tht Penman. J\ hap- modatc them. more as a matrer of convcmcnce than
l'tned "Jth so many groups, wldl·,prt. 1d {'XtJncuon for '\Ctenutic accuracy.
:11th~ n1lr, and only J VC!) ti:w ~lncr.1 belonging The pattcms of evolunon illustrated 111 Fig. H.2H
Order ProlecanitJda crm~cd the Pl'nllian-Tn.mic have been dcscnbed as iterative evolution, m
numiJry. whtch an :1ncestr::ll stock from ti1nc to tllnc ~Jves nse
Shortly .Jftcr the beginning nf the Tmssic rhe few to short-lived groups which replace cat h other suc-
P~mti,ut ~urvivol"'i gave rise to J v,t,tl) ~uccc,~ful Jnd cessively Thr~e new group~ expand ,\llJ dtversity
248 Molluscs
prl'~t
ut a
Card
larly
tunc·
almo
~ror.
nd ~
Up pc
genu
WJse
me m
rcm.t
.tnd ,
uthl·r
~rllUJ
Tl1
throu
.!ppn.
Mmt
that 1
lll,lkl'
,lllJ .
Figure 8 29 (o) Phrlloceros, on Upper Triossic phylloceralid from the red limestones of Hallstatt, Austria (x 0.7). The shell has been suc:h
removed ond the suifoce polished to show the body chamber (~lied with brecciated debris) and sutvrol patterns, picked out by oys· 11.11)'
tallization within chambers This group represents the rootstock of all post·Triassic ammon1tes1 but itself remained very con~ \ Jrtua
(b) Lytoceras fimbriatus (L. lios ) from lyme Regis, England (x 0 35), o representative of one ot the two deep·woter stocks !Tom ...+ucll CO!ll',
many Ammonitino arose ((o) Reproduced !Tom o photograph of o specimen in the Royal Scottish Museum.)
I hr01
SIZeS 1
for .1 whik. bm they arc geolog:tcally spcalang stmctioml pl.ln from which thl·re wne reltm·d., (corJc
ephemeral and 111 due cour..c become e~:tinct. The fe\.v devianons (l- g clymeniid\ and heteromorphS vtrtua
\hap~
me he thl·y v,tcatc i~ tlu:n occupied by de~ccndants of Withm spentlc hne.tges m.my evolutionary trends
the ~ame lon~-1ived ancestr.tl ~tock. Bayer and have been documented. in the ~hape of the ~hell. m CllCOl'
M cGhee ( \1)~ I) show how in southern Gcnnany omament, the ~uturl' line' and even m size and raoo arc re
'simiktr direuwnal changes 111 the physical environ- of sexual dm10rphs (Swan. l9H(>: Dommergu~. F.uml·
ment arc nurrored by ~umbr morphological changes 1990). Though the ad,tpti vc o;ignilicance of the!( ot th•
frequently dudes ;lll,\ly'i'· there is very often a dt·.u mode
in the ,u1unontt1: bunas'.
fhe Lwo ancc~tr.1l order~. Phylloceratida and relatiomhip bctWl'Cn trenJs and hetcrochron} Quite
Lytocer,md.t. contimted throughout tht: Mesozoic demonstrating here. ,1\ 111 so many oLher instanr~ just d
what
and gave nsc to supcrt:tmilial side branches (the the Importance of thi~ phenomenon ·'" Jn cwlu·
EoderoceraLacea, lltldocerat.Kea. etc.). Each of tionary control. Bcc.tu'ic of the rapidity of tmnowr morpl
thc~e dom111atl'd thl· ~ccne for .1 while and during m wide dt~tribunon. abund.tnce and ease of rccogru- C.tllot
'little hour ol ~;,rr.tn·' constituted a miniature adaptive non of the .1mmonires. thl'Y .m.· of outstanding zon.ll ciscly
radtauon. raptdl-) produnng .;hurt-lived families so value. why'.
that the phylo~em·nc crel' rl·~emblc:~ the prongs of a
toasnng fork . lteratt\'t' cvoluuon a~ shown by the Evolution in the Cordiocerotidoe Hell
Mesozoic ammomre' ha' .tlm been described under Ammomtcs prov1tk 'iOme of the best exampks of om1
the wcll-chml'll tl·nn 'thl· p.tl.teontological relay'. evolutional) line.lgl·s. 1 h e rap1dny at which th<1 lr ·~ a
On the whok the ammonoids torm a rather evolved. their abund.uKc and ul·uqlllty ..md th gcolog
homogeneom group b.l\CU upon a conunon con- diversiry of morphulogKal characters wh1ch th t"\.'Oive
prr<ient make them l.'~PC'I.J.Ill\' amenable to this kind are known a~ hererornorphs. ::,omt' appeared during
of analy,is. The lmt<>l) tlf th1.· Jur.t\\ie Famil] the late Tria\\ic. \Otl1C m the JurJS\Il'. and there
Cardsoccraudal.'. whid1 ~panm·d :w Ma. i\ particu- wa~ a more cxrcns1vc dcvelopmt"nt ofheteromorph~
urly ishtruCttVI.' (C.lllomuu. ICJHS) In l.arly Jurassic during rhc brer C"rct;~n.ou' llcccromorph~ may be
amt'5, the northcnl llHltllli.IH' clu,tcr~.·d round an loosely coiled wl(h thl.' \\ horl\ wholh or parnally
ilinmt landlocked Uort• tl S~o:J Thi' 'ea was on I) 'cparatcd. e.g. Cllttn~f«Jftr•h (Tna~ ). "'Jllrllftrds Our.)
'ror.~dJraUy t:olous:r~o:J b\ .munomte~ 111 the Lower .tnd Lyfllmorrras (C rcr). Other genera Jrl.' almost
~d earl) part of thr MtdtUc Jur.l,~tt'. and then m the ~traight, l.'.g. Bn.lntJmtt'l (Cr~..·t.). rc\emblmg bacuh-
Upper B.lJOCIJn, (k,t-cnd.mt' of the Pacific Ou.'an tids. Other<; have tht· bud) l hambcr hooked O\'Cr
gtnus SJ'hacrclfc ltl.• tou11d thur way intl..l the other- the top like a walking 'ucJ..., e.g. 1/amu/in.J (Crct.) . In
\\h'" unc olontzt•d Borea! !:ll',t 'I hey were the tlr~t such btzarre gt·ner.t 'urh '" the Crcraceom
member<; of tht• (. .trdllKl'<.tttdac. ,\ tamily which .\lacro~caphitn and Sf<~plutt'.'· though the earl} whorls
remamed bro.1dl} 111 the ~.irutmpolar llon:al habitat are of nomt.d fim11, the 'hell i\ then ~tr.ught and
•nJ cvolvt·d 111 ~~nl,ltlon. nftl'n 111 the abscnee of finally sharply rct urvcJ nc.tr 1ts tcnninatmn, ~o tlut
other .llllmomte\, unttl the abrupt cxtmcttou of ti:Us the aperture t:tu:s the fiN-f(mned whorl~. The
b'l'oup .lt tht· rnp of tht• LO\n'r Kunmcndgian. Tnassu: Ctlddoa•rclS, ami l ht: CrctJCt'om T~trrilices.
Tlw evolution ot tlm f.muly c.1n be traced Ostlit~i.!t>ccms and othcr ~wnna arc hclically coilt'd
rhrough 2H 70ilt'~ .llld (l2 ~ub:wne~. the latter like ga~tropod,, whik l ht: Crt'taccous Hrte1wcras
.tpproxlmatmg \Ol11t' 25() (}()() ye.1r~ c.tch 111 duration. has ib fir.t few whorh or hdic:ll ~lupt' .md the re~t
\ltN inrcrcstin~ly, .dl of tlw 100 <.H ~o .1ssemblages of the shell likt· Scaplutt•s. NIJIJIOIIItes. from tht·
that hav~ he en dt'\t nh~..·J .tre motlll,pccitic. which Crctac~om ofJ.tp.ln, '' pt•rhap' th~· mmt extreme of
make~ it c.t~')- w di,rlllgussh l'\ olutwn.u·y lmeages. all heteromorph,, h.tvmg a \cry long tububr \hell
md aJ,n dunorphll' p.1m. Within ~omc 'perie~. coiled m J sencs ot U-bcnth into .m unlikdy tangle.
uch a' Quc·mtt•dtclt"l"fcl.< /,uubuti, there j, an ext.lordi-
Dlf} rang~.. o( uHr.t\pn·itlt· v.1ri.uion, embr.King
~cnr§·
•e, Muall) the \\hole r.mgt· from caditonc to m ..")·-
eh
on~:. but mo't other 'Pl'Cit·, .trc of low variability.
Througl\llut dtt' evoluuon of thi' group. ~ha pes and
mrs dtangc. ami 1 h.tJ.Ittn' 'ueh J' rib' and keels • Hamulina (Cret;;coous)
•I} rord~.·d and UIH:orJeJ) 11\J)' <.' onlc and go; mdeed
!>) 11rtually tht• '' h11lc r.m~~<. of Kulptltrl' .1nd ~hell Choristocaras
IJs !lure~ f(ltllld 111 th1.. .llllmontrt' .1~ ,, whole arc (Tfi3SS<CI
ItS !llli1Un£1..rcd Ill tht· ( .11 dlornandat• <;omc morphs

tlO ue r<-markabl) suml.1r to tho,t· nf thl.' earlier Liassic
lt:,. mul) AmalthiJ.tc. Tint' tlu.: cvolution.try history
t:Si: f the C,lrdiOU.'I.Jtid.lc 1' n~.. hly documented in
t:;}f nudcm tenm. 1luwcvcr, thert· n: maim a problem.
)OV ~u1tc sunply, why dtd thc\t' <.han~t:' takc place? We
J;lll do not knn\\ . 'l ht'll' i~ no ckar t·vidcncc of
CC'~
)I li- bot kmds ot sdelt1un prc"mc~ forced di ffercnt Macroscaphites
{Cretaceous)
ver. •nphologte~. d lllJct·d they did so al all. As
~lll illomon tununC'Ilh 'wt• now know rathl.'r prc-
~nal ~ly how the C.miHlu:r.nsd,tc ~:volvcd, bur not
h'
Heteromorphs, extinction and habitat in Hyphanroceras Ostlingoctwas
)faH ammonites (Fig. 8.30) Cretaceoos) (Crer 100~)
tht~ 1< a wdl-kno\\ 11 Cll t th.lt at LCrt.un period~ m

Figure 8 30 Heteromorphic ammonoids, showing variety 10
the lo~~.-al hlstOf)' \Utllt: ~r{lliP' of .urunonites foim. (Redrawn from Treatise on Invertebrate Poleonlofogy,
they 'l'cd shells of lu~hly abl·n .mt turm. Such 'hell~ Port L)
250 Molluscs
1 here are man} lllhcr 'uch irrcgul.trl~ cmlrd gcnen buo}ancy calculation~ (Trueman, 11J41) th.tt hctcro- .lppt
111 the Cn::taccou~. ,\lld thetr cxJstenn: ha\ gwcn rise morph~ such as i\1am1~ft1JIIIiles and Lytnc rian•rm wen.· Wt\1
to very much evoluttlltUT)' ~peculation. well ,1daprcd tor flu,1ting in 1he water m p.trtH.:ular dfc<.
ror a long tlllll' the p.!laeontolo~ir.ll htcramrc on .1ttitudcs (Fig. 8.31). of a
hetcromorpln wa' dominated P} thl· ttica th;u Although the aperturc' do not necc"arily lace m bly :
~uch "hcU ton m wal' Jegcncratl·. retnl!,'TC,~tve and till' ~.mH: directions a\ tho'c of nornully 1·mlcd tn\'O-
bwlogit allv mad.tptt' c Furthcnnon·. because the
IThe
lure or evolute genera. there i~ no rca~on lO bdiev~ thJ, l
Triamc and Cret.ll cou" ep1"ode' of hl'tl·romorph} thar rhc forms were m any way unfuncttollal. :11 tho
rook place <.l10rU) bctore major cxtmllton petiods I khcally coiled .munotute ~hells have llll' "Piltln- tton
for the .1mmonotd,, tt M.'C111ed to m.my p.ll.lconrolo- cle in ~ dorsolatcr~tl r.lthL•r than ventral position ou hi \
g-im that there w.1~ J dctimte rdattomhip between thl' ~hell . These probably swam apex uppcnnmt mode.
hctcromorphy Jnd cxtmaion. IJunn~ the 1910~ and 111 thts posit:.ton the cameral llljlltd would rc..•gn••
Jnd 1930s the \ ic" \\J\ quttc \\ idc:,prc:Jd that inter- derouplc rapidly. Ullmcd•atcly a nc\d) Conned the rv
tu! rhythm~ m c,·olutton cventu.illy "ulmin.ucJ in a dumber wa~ emptted Such decoupling im reas~ the fi
kmd of ·rac1al ~cm:'' t'nce' during \\ hKh bizMrc and the dfictency of \'l'rttc al nugration 111 l{cu:nl tlwrc
OVI.'l'\pcciahzed tom1' were produu:d, .1s a la~t and ccph.tlnpods, .md thl• hclic:~lly coiled .1mmomte1 \t,tgc,
final extr.wagann bdore inevitable exttncuon over- were probably :tdaptcd to .1 ~imibr verttcally llllj.,'rJm l tuda
took the degencr,ltinp; srock. mmk ofl ife (W,trd. 19HO). ~ltturc
However. the rollt cpt ofheteromorplh a' degcn- Though there were \omc heteromorph, ar rht' Olll' t
l'r.lte and mad.lpttvc phylogcneuc cnd-f(mm, plau- cud of ammonnc hMory. rhe nl.lJOrtry of th~ 11111110
"bk though it \Cl'llll'd tn ~chmdewolf anJ tb otber ,unmonite' at that oml were nom1aUy rmlcd lu not m•
pmponcnc~. is now no longer getH:r.lll} held. To thl· l',trly Upper Crl'taccom the ammomrcs wcm 111111101
begin with, ir ha' beL'n ~hown (Wtl'limann, Jt)69) into ,, \low declinc ovcr .1 long pcnod of ntllt' , Jnd We
th,lt hctL·romorph~ appe.1rcd in ccrt.1m ltncagcs only towards their final end they h..1d bccomc re .. rnctcd ,nd in
and were not char.tt teri~ric of the ,tmmonoid~ as a to certain parts or till' wor!J only. The number oi Om· v1
whole .1t any om· tune. In the l n.mtc there were genera became fewer, .md finally no new characrrr ,,, 191)
only tour dowh rd.ucd hetcromorphtc genera.
111t'llt\
qum: lung-rang'-·d .md prob;1blv 'penalized for
more c
bottom livin~. wlllt h bL·nme exnnct .1t Jbom the
tlw P,1l.
same nme a\ etght normally cotkd 'upcrfamilics;
lt•vd, 11
clcJrl} there is tW !Jkdthood of a ram,tl connection
th.lll tht
bet wccn heternmorphy .md extinction. Likcwisc
thel'l' W•
the seven known !;l'lll'r.J of Jura\SK ht•teromorph"
l.ning tl
were probably monophyletic and WL're \pecializcd
''·Iter. 1t
bottom dv,dlel'\, where.t~ Crcnceou' hernomor- tAl-•• (evolutel th,lt of
phy. wluch n:,Jt hcd I!' maximum dl·vdopmcnr m
In\\ OX)
the early Crct,Jteom, \Cl'lllS to h:tVl' been J poly-
lOJnlllze
phylett.c phenomem>n. Curiou\ 'hdl fonm .1ppcarcd
lwt Wl'Cil
t11 several lincag~·~ l)liHL' ~uddcnly. often associated
llllllpt•tit
wtth reduction of the prinJJ.ry "uturc. M.my of the
ttk•\, \\ht
uncotled or partt.dl~ coiled hcteronwrphs that had
\\Jtl'l'\. l·
been present in till' Lo\\ er Cretaccou' produced
c•xygcnar
descendants \\llh nunnal or near-nomul cotling;
:uumonit•
tlm agam taJ~it]e, thl• Vtl'\\ that hl'tanmorphs were
ntll' the c
overspeCialized t'nu-fLmn~ from whJl h no further
Jmplodecl
t'volutlon was po~\lblc. PrcsumJbly thc1r ability to
thl W:lll'r
retum to normal l·uilmg when .1ppropri.m· erologt-
trap. In
cal mches wcrt' ,t\'Jthbk was a tJrtor tn the subse-
Figure 8.3 I Possible life attitudes of nonnol ond het~ alrt!,td\ 01
quent \ne cess ut' the n·-co1lcd genera ph1c ommonoids os calculated by Truemon (19 41 ) The bo:) of the Cn
h ha~ been 'hown, furthermore, by density- chamber is stippled. thL·ir l'nu.
Principal fossil group s 251
trpeared. fhrn: 1\ no l)Ut·~ticm of :m intcmally A ~onu.:what dttTerent vil'W t\ that of Westcmunn

\lie~kcned ~tllt:k: the Jerhnc w,\, rather tht· adverse ( 1990. l 'J96) who prc\cnt~ habtt.tt diJgram.~ for each
df~ct of long-c.:onunueJ t'll\'tmnmcnt.ll t:ondttions. geological penod. H e sug_~c'~ that Juras~tc and
fa kmd detrillll'nt,tl w the ammonite~ and proba- Cretaceous JJmnonite~ wcrl mamly pelagic (dnftcrs,
b!) l\\OOJtcd v. idt .1 'l'ne~ ot m.mnc rq,rressiom. swinlllll'~ and vcnic.ll nugr.um) W lulc ~ome may
ifherc 1' now 'onw doubt J' to how pronounced have colomzcd ncritic .md oc.:eamc habttats down ro a
dus dcchnt• acn1.1ll) w.t,: there: w.b indcc:J .1 decline depth of HOO m. mo~t hved between ~() and 120 m.
1 the l'nd of tlw C.unp.mt.m. but mtcnstve collec- !)ome of the larger strc:unlmed O\.')l'Ones nu} have
tiOn from rnttt".tl honzon' m the Ma.l!>tnchtl.m come up dme to the ~ur6ce. Accordmg to tlus ,,ew.
led Ward ( IIJY(J) w t:l\ our ,t more.> \teady-state the nerinc zone. above the c:onnnental 'hdf. mcludes
modd tor that nnw.l rlw l.m maJor Cretaceous (1) · pl:mk.tonic ammomtc larvae, (2) pa~stve pelag1c
~t>.;r~~ion (.md the: ChJU\ltlub mt·teur tmpact). at dritters such as the long-bodted Jura~~JC serpentlcone
die Mamnc:hoan Dam.m bounJ.u), comcided with D.utylh,rcms, some ~plucroconcs and some Cretaceous
th~ final md ~uddt•n dt:llll\C of thc:- b't .nnmonites; hetemmorph~. (3) pebgtc vcntc:JI migrant~ mchtdmg
there arc ceph.tlopmh 111 the.> ~ucceedmg Danian T11rrihres :md Sct~pflites, (4) sluggi\h nektobenrhos. such
)t.tgc. but they .m:- Clllled n.lutilotds of the genera as the hjghly om:m.- planulates (Arit•tites and
t ~/111ir1 .md l l£'r(t1,~h~~.~., (l tg. H .2~j) which have zigzag Aamtlwrenu). To th is habitat belonged the Jurassir
sururc lme~ The~!.' prcstun:tbly Wl"re occupants of Crnvesia and the C:ret.:tceou~ tH:oceratites. whtch were
\C t1flC of the ec0IO~IC,II mche~ V,ll'Jted by the among-;t the sh:tllowest- water :munonite.o; of all, <llld
he urunomtn, but I ),tnt.ln n.tuttlmd~ apparently did (5) mobile nektobentho~ . l'hese wen." stre:tmlined
ln not mvadc tlw mu lttph<tty ol mu:rom che' that their forms, such as rhe oxycone oppehid~ and some platy-
·nt unmomtl' prcntr<;or; h.td dnlll' concs (Flilda{era.s :md CrmmiiOfl'lil..<). Many oflhe<oe may
nd We h.we seen th.u \J,wrifll.\ ltn·~ at some depth, have preyed actively on other .unmonite~.
cd d m regions wlll're tltl oxyg,·n tem10n ts low. The oce:~nic regton had an c:ptpclagic zone mhah-
of ,)ne vtcw of .lllllll<1nite depth prctcrcnces (Wells e1 lted by phnktonic larvae, wh~t:h lived there for up
ers 1992) 1~ that they h.td ~inular habit.u require- to a month before moving cl~cwhere. I n deepc:r ofl:
m~nt( to \ ',mttlm (e\'cn though thcy Jre actu.illy shore waters (lower tpt- ;md upper mc~opelagic)
morr clo •tl) rd.ttl•d tl) the coll'OJd~). Throughout there lived mamly tT.lgtlc-\hdled ammonite~. \UCh .ls
mt P.~l.tC0701< md much nf thl l\1c~o7oi,·. OX')'gen some Crcnceom hl'tt•romorph~. and importantly
,e), m thl· deep ,e,t .md mtd-w.lten were lower the deeper mesopehgtc re;tlm wa~ mhabitt•d tn the
~n the) .lrl tud.l\. In thl· Jur.t~sir and Cretaceous Juras\tc by Pflyl/tltt'r<l.\ (>500 m ) and Lyt<ICH<LS (>800
tbm wert· no pol.u '.tp\ ..md thu' no \\lay of venti- m), both of whtch were vt·rucalmigr.um within thi~
,wng tlw deep mum by htgh-demtty melted ice- range .
Jtcr. lf.unmonttl'' h.td .1 comp.tr,tblc phy~10logy to Why did Nt~rlll/w survive \\hl·n the ammontte~
1t of .Va111dus.•md .1 \lllliLtr .tbtlity to w1thst:md dtd not? Pmsihly hl'CJU~e iL was an umpecialjzed
111 O\.'ygen kvds, tht·y could h.wc successfully feeder, especially well adapted to low o~.:ygen condi-
olnnucd thl! v.t•\t hypmoc rqpom of the ~ea tions, and l:tid large tggs. with high survival poten-
rwccn 50 ~tnd :mn m, where th<'rl' w:.~s littk direct tial, in deep waters. On the: o t her hand, perhaps it
rompctmou from fish, lx·lc:nmitl's or marine rep- was no thing more th:t n good luck.
lib, which mhabttt·d the high l) m;ygenated upper
1\'ltt'l'i. Pcrlup,, ,1, the on'.lltS became increasingly Subclass Coleoidea: dibranchiate cephalopods
~v~~:nJt<:d, the ')1l'n.tlin·d hypoxtr h:tbitat of the Modern coleoids
•momt<'' began (l) 'bnnk.. They t:ould not colo- The dibranchi:ltc ccphalnpmi' or lOicoids indude
7~ tltl dct•p -w:ttcl':\ bct:.tll~t tht•tr 'hells would have squids, currldish, Oltopu\c~. thl' p.•pl'r-nauttlus or
:!lplodcd .llld, < h.t,t•d dmvnw.1rd' b) pred1tors from Ar;gonawa, and variom t'xtinll groups, notably the
wateT\ ,1bm:l' till' .mHnnnitc~ were caught in a bclemnitcs. Thq. all h:1ve only a smglt: pair of gills
r In the l'.trh Uppt•r ( n.'t<lCCOU\ they were withtn the mamlc cavtty, and tt j, th.is fearure that
~d} on the 'kduH', .tnd thc c:.lt.l,trophic events 'eparacc' them !Tom all rhe other t·eph.tlopod\.
romor-
~ body th~ Creweeom- 1 \·rtiJrv boundJry only h~tc:ned Tht modern <i(jUJd' and cutddi~hes range from
r.:ru end tiny ammah on!) 4 c rn long to the gigantic
252 Molluscs
.tlrdutl'utlus. wlm.:h 1s (indudmg the tentacles) 18 m mro an intcrc~ting evolutionary perspective. He h~·
long Y l't their ~t=ologiol impurt.l!Ke ~~ limited; how by early Palaeozo1c tttne~ cephalopaih
~h0'-''-' 11
only the bclcnuutes .1re nomully pre~erYcd as fomh, were the mmt advanced and rnobtle of all rrunnc
.111J m am. case coh:01ds Jll mon. divt:rst' .md abun- ammak With their chambered shclli Jnd buov1nCl
d.lllt nuw dlJn cbt·r wen· fonnerly. wccb.mism along the Lines of Nauri/11s the) we~
One kmd nf dtbralll·hLue 'tmcwre is shown by highly ~utTes:.ful, though linmed tn how deep rh()
thl uJttldish St'fli<l (Order ~~.·puda), whtch is illus- couiJ go due to danger of tmplo~•on. However,
rr.ltt'd in Fip;. K.21 f.g. lt differs from Nautilus iu hav- when Ji'>h began to dtverstfy and mvadc the upper "'"
in~; cmly t>ight .mn' .md two tlnWdC\, which arc W.ltt:rs of the sea. in the later Pabt'07mc, one dfea
prov1dt>d with \Uckt•rs .md hook\. These hooks. like of \llth competinon was to put pn·~'ure on lint~Z
cht• j.tW\, are homy .md .lrl' not tml.lily preserved a' · space. fhe origin ofrhe 'anunonutal' buoy:mn )}·
to"•l~. thoup;h ~ome r .lrt' i(... "iliz~.·d l'XampJe, are tem was one answer to the problem. for tr tir
kno\\'n. In tht• mamlt· t.'.l\ity with it-. two gilh those cephalopods which posse" it from the hmiu-
then: 1\ Jn ink ~.lC. u~~.·d fur dtllldlllg the water as uom ot the ~heU. and it IS perhap:. not 'urpri\111~ thl
an l''l....1pc re.Ktion from pred.ltol"\. Otherwise, and the deeper parts of the sc.1 h.lVl: been cxtctl\1\dy
.1p.1rt ITom ~hdl ~rrucmre, the mtt'mJI org:.mizanon colomzed by anunoniac.:tl sqUid. Another ,oJutto
of the body i~ hro.1dly wmp;uablc. to LOtnpetmon from fish wa'i th;H lll.llly rephalopodi
ThL shell of St?pia i.l whully mt~.·mal. It is a large bet ame nwre ftsh-like, stmctur.1lly .md tuncttonall
ov.d hmly. known m th1. wm.ltlliJr Js the cuttle- ~o that the two were then competmg on more t>qtUI
bone, looted dorsall} llld <mnpQ\Cd of do~ely tenm, as Packard commellt\, 'ceplulopod,, tun·
<;p.tc ·d. oblique. C.lkar~.·om panitiQm 'upponed by ttonally are fish'.
pill.lf" . Tlw cutddi'h uuuroJ, lt' buovancy by secn·- Sptmla (F1g. !'!.21 hj) i~ a lmk 'lJuid 'omr 10(111
rinn .md extr.Ktion L,f hqtnd trum within the spaces long wluch h.1\ an open. 'Piral, dwubered sld
hl't\\ t'l"ll partitium. A 'hell 'uch ~s tlm 'eems at tirst enclmed l'nttrcly within thL· body. Thl· 'hdl Ius
'ight vel) dJ~'imilar w tlut of .1 uautiloid, but 1t JS vcntr:~l <\lphuncle whtch connc.·tb \\ rth the ch:nnbc
mmpholo~c.Jtly the dor'i,\1 h.1lt of the nautiloid only through a very ~mall porou' region in nclt
phr.lt,'lnmone which lu' becullll." Aattened and The lower chambe~ arc lar~dy fill ed w1lh li4u:d
exp.111ded. In the '4uiJ L1h.~,, the "hell ("ea pen) i" which is nomully inert within th..:m, hut thiHatl boo
horny. .1ddcd to or subtracted fi-om through the pvr1
~Hill' the ch.unb~.·r, wttlun thl partJtlOJlS are very rl·gion ~o that the ~hell\ buoyancy cu1 be coutnJll.
nlrr<m, thL change 111 buoyamy through the in a manner ~tmilar to th.H of ,\i<JIIfri••S. Th:
pumping out uf water c.ln b~.· .1 very "low process. at evcr-chJnging colour pattem' of ~qutds. in 1\bxi (
!
lt:.t,t ·" n:~ard' water deep wtthm chc chamber. rollmg colour waves, sudden darl.cning. of the '
"'t' erthde". buoy.uKy rq~ulauon b)' the alteranon or colour Aashc~ Jrc U\ua.l, r11ay IIIC\IJU:rizc ~
of WJter balance w1tlun thl chambered shell IS camouAagL' the predator or tJcilit.Hc loum1um
not the only system m modem \C(tlld<;. Quite a large non berwet·n individual,.
number of o;qlllds. e.g Cr.mdti<l. have in,tead The ~hell of the female Ar~tlll<l/1111 ('paper IIJI.-
adopted the use of ;umnnllllllll rhlonde within che tihh'; Fig. 8.32n) is a very thin ~piraJ ~heU, but a
tt~meo;, whJCh IS I~Otomc .1nd 1<\mmotic with sea formcJ by Sl'Crt.'tion of the ll'lllJdcs Jnd cam
w.lter and thus gJVes neutral buoyancy. In these cxrc.:rnal to the body.
tl.Jilchid~ the codonm cJvtty 1<\ \ol'tly d1stcnded and lt i~ med as a brood pouch. more r.1rcly fo1 'm)
lilleJ With llrunom.lc.ll lilJllld of relative demity mg ahout ,, captured male! The nu le. mndentJ!Jy
1.11111-1.1112. whKh j, Je~, th.1n due of ~eJ water \hdi-Jcs, and only about onc-t\\'Cntit·th the :illt
(rd.H1vc density 1.1126). Though ~uch anunonial.ll the ti!malc. le hJ\ a very long wxuJJ organ (the ht Figure B.
'I.Jtllds .1rc vel)· Jbumbnt .md the U\C of .munonia ~~ toc otylus) wluch c.tme:. tht= ~perm tlm can dew guord cui
roconstrvt
not contined to the n.lnllmk the \y,tcm c.mnm lt,eJf from the male .tnd sw•rn alt1vdy to\\Jnb ISOtope re
app.lrently be used for r:1pai huovarKy l·han~e\. female a kmd of 'guided m•s~tk L·opul.ttmn' "b1 (MIO,) m •
J> Jl k:Jrd ( 1972). 111 dJsCU\\lng the evolutioi1Jry \Ct'tlls tQ be unique m nature. '" Troite c
convergences .md intl·r.lCtiOll\ bet-..veen rephalopod~ I )c~pitl.' the fragility of the .lrgonJut \bell, tll:
.1ml ll\h, hJ~ put th..: ,lmmonl,IC.ll .md other <;quids tl)~~il record can be traced b.tl k to thl Olr~o<:
(d ) Hibolites
(I ) (c) ; .
(j) Duva//a
pro-os1r.1cum
l)hragmocone
guard
Te m pcr<lture (Cl
(h)
16 20 23 26 (k) Aulacoceras
(m) $piru/irostrfn11
( j) Ausseltes
pr!XIstracum
Conlilf':tan
San10nibn
guard
I'\
·d quard
·-
IS
of
c- :;we 8.32 Coleoid mol'phology (a) l~gitvdinolly sectioned belemnite in ventral view; (b) some, in lateral
:h ,-d cut transversely (all x 0.75l;" (d) Hibollfes (Cret., >. 1 approx.); (e) Actinocomox (Cret., x 0.75); (f) Duvo/ioview, (c) belernntle
(Cret., x 0.75),
·m1slruction of swimming belemn1te, with the guard wholly internal; (h) climatic Ruclvotions in the Cretaceous, (g)
hl' ~ratios in belemnite guards; (j) Ausseites phrogmocone (Trios.; x 0.75); (k) Aulococeros
plotted from oxygen
(Trios.; x 075); (m) SFJiru/irostrino
eh -10 I in section (x 0.75); (n) 'shell' of Recent poper-noutilus Argonoulo (x 0.35) . (Mainly
based on illustrations by Noef and others
Troifi de Poleontologie, 11 (h) after Bowen, 1961 .)
hr:
ne
254 Molluscs
(Holland, ilJ~H) Tht· ~.uhc-st .1rgonaut shells are ture :maly'>l\ of the J ur.t~~tc .md Cretaccom h' gl'lll
smooth and m deed wry hke those of nautil01ds. It '" ••0:"0 ISOtope r:ttiO\ (Fig. X.32h). Mcs
posstble that rhe Jllle\tral female argonauts firstly The th1rd compom:nt of the bclemnirc shell!~ the Cc..'p)
used d1scarded n.luttlOld or .1nunomce shells for pro- pro-ostracum: a long, flat. expanded tongut· pro- nf r~
tccnon and then comrnu:tc:d thetr own on the same JCCtmg forward, and pre,umabl> cm·enng the :mlC'- 111£0
model. rior part of the body It \l'Cill\ to be homologou\ to b'TI)\\
the 'pen' of the ~qtnd T~'''.11'' Tlm 1s rarely pre!>en•td. n
Extind coleoids and their evolution (Fig. 8.32) however, and 1ts fum tton is poorly undcNood. l US\C
In vel) m.my Jur.l\\IC and Cretaceous sedimenrs Sometime~ 'Pt'nmcm wtth eight radutmg \ets 01 thee
there may be .1n .1bund.1ncc: of fmsil bclenuures. hook!\ hdvc been found forward' of the: guard. lc,u- the
These .m: the mtc:m.tl ~hell'> of toss1l squid-ltke fying to the fonncr prcwuu: of arm\, md l'\Ct Aul.lt
cephalopod\, bm thq· dt) tH..H dmely re'>emble the to~sih1ed mk sacs h.we hceu located in place. l1ltcrr
shell~ ot .my modem 'lJllld or cuttldi,h. Bclcnmites \ccm thcrclc>rc m h:1ve been a kiod vf Bden
In the typtc.ll gl'lllt\ Hrft·lllllift'.l tht• shell Ius three fossil sqllld, with a c.litTcrcnt harJ-p.trt con~tructlon Au~rr
parts. The IJrgl!'l .tnd most pmtc:nor '\ccnon i'> the to other' .md tlwrcfore with .1 umque ~y~tcm ul Wtth
guard , ,1 lll.IS\lVl' btillc:t-~h.!ped cylinder of '\olid cal- buoyancy conrrol. cxten
cite. lt 11, parallcl-~idt·d. t.lpc~ po!>tcriorly to a point Ma~s accumulations of bdcmniLt: gu.mh a1c not othen
and is indcntt'd .11 ir' .111tcrior t>nd by a conical cavity tull'ommon in tht· Ml..'sot.oic, and such 'bclcmuit~ here t•
(the alveolus). fl rht• g;u.1rd i' cut transversely, rhc battlefields' may, :JS with modern squids, reprc,lth orth oc
struct\Jrc i'> ~t·cn ·'' rJdt,tlly oncnted needles of cal- mdss mortality :~.ftcr 'pawn ing;. Similar JnUiuub· 1 l'IJ \si
Cite, w1th lOIKL'ntnc !;.'T<>wth rin~. which arc :llso tion~ of orthocone nautiloids arc common iu the ~·xp.mc
appJrent m huntont.ll \l'ttton The a.x1s of growth IS Palacozoic. H owever, there an: other pos~lbtltUl'S, and (
not crntrJI. ho\\ rwr. but placed towJrds the ventral catastrophlC n1.1s~ mort.lhty, im~m~ pred.mou, and deciJn<
margin U\ually the 'urlate of the guard 1s smooth, concentration through Wlllll0\\'1llg of sedtmcnt contjm
bur it may be granul.1r or ptttl:d. Some genera. such (Doylc and Mo~tdon.tld, 19lJ3). it ts usu:tllv ptt"tbk extinct
as Hibolitc.•, have J Ion~ \'elltra.l groove cxtendmg to establish wb1ch ofthe~e Jgenet~~ operJted m par- A pc
rwo-th1rd' of the \hi\ lrmn .tlvc:olm ro point, and ticular ut.~tanccs. n:prcsc
rhe po,tcrior part of the 1,'1.1Jrd 1tsd.f 1s somewlut There 1~ evtdcnn: of 'l':wal dtmorphism m s•.lllle Jura~sic
~wolleu, Allllltltallhl.\ 1~ more par.illd-~ided but like- belenm1tes (e.g. Yollll.l(ibl'iw. Doyle, 19H5). a\ Mth hut is •
wise has a vt·ntul 'lit bdow the alveolus. Som~ <lnunomte!>. ~ htherco bdemmtc~ ha\·e onl> b« cone 15
spene.. or DrwcJlill hJve ,I lltnOU~ lat~rally flattened used for ~tratlgr,lph!C,ll purposes m the UpJXI O~tr::ICUI
guard. Cretaceous of northem Europe. Yet they are abun· known
Wtrlun the .tlveulu\ ,md fitting 1t exacdy is the dant, rapidly evolving and usu.1lly mdependcm of .11c ~om
phragtnocon e . 'T lm ts .t tolulal, tlun-walled arag- racies. Though at first Sight they appear to look hut oth
OllJtic ~trucwrc wlut h proJect~ outside the alveolus. rather sinlil.1r, tt is .Ktually 110t h.ud to tdenttf)r thtm type's is
lt is scptarc With tb ~epta nmc,we anteriorly and (Doyle, 1 <.J90; J)oylc and lknnett, 1995). ntict thcr. H <
separanng f.ur-sized <;,tmcrae. A slender ~iphunde they are found where the ml''lre fragile ammt1mtd J'emary
thread~ through the ~cptJ ,lt the ventr:tl margin. are abscnr, :~nd the1r blo~trat!gtaphic use can bur lie cont
These ~tn~e iures, together with a tiny bulbous pro- increa'>c. rl·duced
toconch, leave no doubt that the coleoid phragmo- rh~.,· pb1
coue 1s the diren homologue of the shell of a Geological history M10ce-nc
uaunlmd, anunonoid or Spinrfa. Bclemnites arc grouped mto two taxa: tbc ••phunclt
The guard, wll!lh ~~ ~uth J 'ubstantial part of the Aulacocerida (?Dev./Carb.-Jur.). which nJJy h 1 tht' phra
belemnm·. ha.' no dnect homologue in other retained the body chamber.•md tht' Bclcmnwd: ~trutrure
coleoids. lt 'eem' w havt· .\ded as a necessary coun- proper Qur.-Cret.), m \\ htch thl' latter \\'.ls n·duccd of the l
renve1ghr to m.lmtJin the bclemnitc body level to the pro-osrracum. Both b'TOup~ have b'UJido r('duced 1
when ~w1mmm~ t c. fi.tltilling 111 a difrerent \s,.·ay the which are nor ea"ly dl\tu1t-,'1.mhable. Other th2!l lltt· end ,:
~ame functiOn J\ tht• omer.1l dcposm of nauriloids. doubtful Dcvoman record, the l'.uhest aubcocend. ltdv.~qna
Smce the guard conmt~ oftluck .md unrccrystallized have been reported from erratiC boulder.- in ~o:tl. mdl't·d, t
calcne. tt h,,, proved 1110\t usefi1l 111 pabeotempcra- America behcved to be of Mts~tmpptan age. lt mure tha1
Predation and the evolution of molluscs 255
~'ncr;tll} .tgrced th.lt the~~. for~.>runner; of the Order Tcuth1d.1, to which Loi~~<J belongs. \Ctrn\
\l~ozo1t bclemnHe\ were denvc..•J from nrthoconc to have developed along .mother line. and tht· pro-
.rph~lopods about th1' rune through the c..·xp.msion O\tracunl h.t\ become an Important mtenul \tnlc-
1f dtc thm coverm~ of the t1p of tht.: phragmocon e tun: at the expense of the phragmoron c and t-,rt1.1rcl.
tnto dw m.1ssivc gu.mi by a simple process of relative Sepiids and tcuthoids .1rc found in the Jur:Mic
,~owth Solnhofen lunc:~tone. :md tcudud\ at le.N .tre
Tbc: sy,tcmatlc' md phylogeny of colemd~ ~~ di~ known from the Lower Oevoman J lull\rticbc..hi cfer
u~c:d by Doylc: er <11. (l9li..J.). l.tttle h known :~bout (Chapter 1::!). Phosphanzed soft pans, re~ulting ti·01n
th~ earlier st.tgcs ofhclemnite evohttJ.on, and only in very early di,lgcnesis, ,\re known in squids from tht.•
m~ Tnamc .Ire rhcrc..· t'llCOlltltercd well-known O?Cford Clay of the Enghsh ]urass1c (Aih~on. I ~HH).
~dicolt rid.1, whit h 111 'ome ways luvc c h.trJcters fhe anccstl) of tht· lmle sqUid Sprmla w1th it,
tennrdJJte bctwc..·c..·n those uf orthocl)lll'S .1nd coiled intcrn.1l shell i' unknown. h h,\s no guard m
llelemmttd,1 . The..• Tna~sic liusscift•s, a genus from pro-ostracuw , but only a loosely coiJcJ pluagmunm e
~u,tria, ha, a '>null gu.ud md .1 lung phragmocon e with a large prutoconrh and marpn.ll '1phunck
f 1\101 01 marginal \I ph uncle . ...tul.lutlcr<ls ha' ,1 l.trgcr, Smce the buoyancy rnech:nmmc; of modem
.\l~maU'r ribbed !,."\I,Ud, though the phragmocon e coleoids an: now beginning to be well undentuod,
•chmvise resemble' th.lt of Amseites. It i~ pm~iblc tt may be hoped that tlm will give n corresponJm~ly
b~re to assume an c.. volut10nary progression from :m broader conception of the meam whereby fossil
nhowne with extemal rtuckemng, through the rephalopod\ , mduding coleoids, \VCre able to c..on-
T:msu.: ~nera to the..· rmc belt>mmres. The latter trol their buoy.1ncy. lr ~~ unquesnonc: d that th1~
c .xp~nded anJ thvc..·~1tlcd throughout rh~:: Jurassic has been the most important ~ingk f:.1ctor 111 the1r
llld Cruaccou~ .tnd then, hl.;e the amlllunitcs, t•volution.
dined t>xcced.ingh·. though .1 very tew bclemnitt'S
ronunucd mm the e.trly T t'rt1Jry be tore beconung
~1111Cl.
AposSible side bran( h of Me~07mc 'belemnitc~' is

lpn:,ented by the.. friJ~SIC PJn,I~IIIMt'utlli.' and
c jurJ\\IC &•/cmuoteut/m In these the gu.ud 1\ presem
h ut b Vl"ry thin Jnd dchcaw, wh1le the phrJgmo- In mollmcs, a~ 111 .111 orher animal' w1th volum1nom
11 ronc 1\ short and bro.1d with .1 verv long pro- and edible At.:~h. the clanger of pred.1tion hJ~ luJ J
er !Jitrmun exrendmg from it. Tht· tirst octopme~ are decided imp.1ct upon the course of thc:1r cvnluttun.
mol\11 timn the Crt't.KCOU\ of Lebanon. and there ~uch prcdJtlon ha' increased throu~h tunc and d11:.
~ 10mc fossil argon.1ur' known tr-om the 1 crtiary. fcrent k.inch of moll me'\ h:wc responded to 11 111 dif-
~ut mhcrwisc the f'o,:.d record of these modem ferent ways. In fossils th1~ can be ,1ssesscd and it~
:l'flC~ i\ so poor tlut 1t 1s not worth rnentJOmng t'ur- eH'i!cts interpreted by cxJmining tr:JlCS of prcd:lllnn
rr. However. there .1re some cxtmct coko1ds of on the shells. V crmCIJ (1983, llJH7) argue~ dut
atUI) .1ge whme to'''l hi,tor. t\ mtercsun~ tf a lit- through omc the food-cnchiu g cJpJctty of preda-
le roufu,ing. Somt> genera luvc .1 ~omewhat rors (on mollusn and other mvertebrate~) ha~
.tlur~d guard and .1 '>~mal coil to the e.1rly part of Increased dramaticaiJy, bur has necessarily bl'cn
phn~'llloconc fhe Eorene Be/ttsrpicl and marched, in the prey, by increasing!) clfcctlvt· ,\lltl-
!ioccn~ IIJIIru/uu5111111 are of tlm kmd: m both, the predator dcviccs. Such adaptl\'e C\calJnon ha.\ kd to
uncle I\ much e'panded Jnd the ventral part of .1 'biologic.~! .mm race·. wh1ch is lllO\t evident fol-
' phr.l).\mOconc reduced. Though this kind of lowing period~ of mass extinction.
'JliUrt '' m some..· ways intcrnwdi,1te ben~een that Ccphalopodc ; depend upon escapt· or avoid,ltlce.
the bdemnite' and St'P'''· '' hirh ha' much The1r capanty for jet propulsiOn. coupled w1th the
.I ed j..'UJrd, the IO\~tl~ are \ el)- fe\\. Jnd to relate ejection of mk when disturbed, Ius bt:en .111
~nd members dm·ctly 111 .1 ~eyuencc Bdclllllites- undoubted l~1ctor in thetr surviv:JI and SLICl'l'S\.
1 lfJIIci-Scpitl is at pre~cnt unjustitied. It I\ pm,1blc. Ga'ltropods, e~pccially from the Juramc onward~. arc
~. that the\c..' c..•,trly TerllJT} colc01d' arc no rrorected and well served by theu th1ck shell. nJr-
t I thm ,malJ pJrt<; of large ordinal') c;epuds. row aperture, and retr.lct.lb1hty of soft ti~mes w1thin
256 Molluscs
the ,hell. The trc.:·~1uc:-rKy of gasrropoJ 'hdh which B.utcn, R L {IWl(J) l11r l..t•w;:r C~rltc>II!(<Wih r..hi10J'Od
h.we been rep.nrcd .1ti:er damage te,tifie' tu the /,,,,.,1 .fnmr rlr~ Htlllf'd/; Lnm:slo/lc 4 Ct•IIIJIIo•ll ,H,IJIIII,
etTn:tivene~s of the g.l~tropod shell .1~ .1 tonrcs~. for Stlf/lt'r:.d. Palaeonww.cphtc,tl SocJcty Mnno!-•Lrph' No,
though such ~hell~ h,lVC bc~·n danuged, their inhab- I 1C) .end J 211. P.rl.teontographlcal ~nu et}. l.omlon w.,'
it.mt~ have nnt bc~o·n krUcd . (Monogr.1phic 'tudy ut ,, drvaw taun.1) .I
G.IN.Ill~. w. (I 'JSI) [..m•.ll Forms ami Otlu·r z,.,,J,ogr.-nl Wu
13i\'.ll\'e 'hrlh ,rrc.:· much more ,·ulnerabk than arc
I ast·s. Blackwdl. l )xtord. (G,"tmpod tlH,rnn) l
g-a,tropod~. They luvc an rntnn\IC hrnitaoon
Kcnnctl\. W J and Cnhlun. W.A. (I <J7(•) ,-1>/IMS of Wu
unpmc.:d upon th"''" b, rhcrr shell c.:onsrnrction. Fur
' \mmc111itc Bwlo:,:y. B~<l\!ttl\!I<IJih}• mrJ Bit>.,tr.lfr,C!I•IJ'f')'. Sp.:~'UI l
\\ ht•n the 'hdl " broken at tht• margm, the seal P;rpcr. m P.Ucomoloh'Y No. 17. (Jnvaluo\bk trt•.mnrnr) [(
between the v.tlvc.:·, 1~ demoyeJ, body Rurds leak out Landnun, N.H., 1 .cn.rb<·. 1<. ;md D:rvr,, R.A. (I'Nt.)
am.! predator; c.:.lll 111.1ke short work. ot the shell. ' 'lmmmroitl Palei.IL1i,•lt~t.:Y· '£opic~ in Gt:t•biukJKY No. ll
While thi\ ",1, .riW.l)'' a problem for brvalves, it Plenum. Nt'\\ York .md London. (Jndi,pt•n,.lble rnm-
bc.:t a me acute 111 the earl) Mcsozmt' owmg to the pll.uwn. 2tl plplr') lr
proliferJtion of m·w kmds of prcd.Hors (g."rropods, lcbmn. P (l q'}fl, 1'197) lmmomtt·s I l..~ 2 ,\lrtlttiiiiA tl
trustatcam .llld ~t rrh~h) Th1s, the Me~ozorc Manne f-Msrft•, 2. Ed1tion' CEDJM. Pan>. (Fin.: Lolour pl.11rs Alh~·
Revolution (MMR) ofVermeij. put llluncnse pre~ I d11u~rm. U. (1Y81) '17rt· lmlll<lllitcs- thur l.r(to mrd rltm tit
\Urc on the bJV.tlvc.:·~. and they responded m a v.1riety I I nrld. Cambndge Umve~JC)' I' re''· <. .unhndg~ All111
(C1l'.1r, maightlorward tc:•;r) J),
ol wa)s. primanl) c~cape .md .woJdancc. to protect
l.d1111,11111, U. (1990) .Anw •l((lidt'l'll- Lcbm ::rr•rfdrc~r Slq/1• [ ,c
drem~ehc' (Harplr .md Sk<..·lton, 1991) those that
rmd Clrarybdis 1-'crdanand Enkc Vl rl.lJ.:. \tutq.'llt U.lbll
n:mamed eprt:turul wen· e'pecially .H m!.. cc:meuted (Sl'<:ond edttlon of .Jblwe) Or
furrm being le'' 'u than .mached bi,·.llve~ (though Moor.·. R C (nl.) ( IIJ57) Trul/IH' •'" Jm •rncbrlf1• p.tl
not d~dimt the.:: drrlkr kllief<'·. the munt"ld ~as Halt Cllllo>lol/y. P Jrr L, ,, J,,l/u<u1 -l. (,eolog~t .11 So, l~IY ai Uo~rkt
tropods). Epiby\\,lll" forms can tr.1p 'mallcr assailants A menu and l.J nJVt'Nf)' ul K.tma~ Press, I tWrt'nre jot.lS
\\tth byssal thrt\ttk Tightly ,c.1kd v.tlves, with K.rn (Ammonoid~) U.ryt•r
mtcrlocking tullllllimrrt·~ {Carter, l 1H1H), help to Muo.)f'l:', R. C. (et!.) (I 1J64) Trcu/rsc 1111 lm•nrl'f,r:tU Re
Jcter predator, ..rnd 'pinose brvaln·'· togt•tht:r wtth p,,Jt·cml<llq~y. PJrt K, \lcrllusc,r }. (~t'nlo~H.tl '>tli.ll't}' ,{ An.
bonng .1nd uc,tlrng type,, rncn.·a,L·d grc.uly afi:er the Aml"nC.l :md Umvt•r-~ty uf KJmd~ Press, L.mrcncc fJ.1It
K.m ("'aulilmill).
M MR. Spmc:~. hu'' e~ er, can only be generated m llOI
Muon, R.C. (ed.) ( I'Jh'l.l) Tro a/bt ,,, lrll'mtbr ll,tyt:r,
brvah l"S wtth a flc,rbk peno,tr.Kum. not rn others,
Jl.J/r,mt<ll•>gY, Part I. \ /,,flm.., I Gcoln~' .11 Sout'l) of ut 11
and 111 gener.tl potc.:·nu.1l a<Llpt.ltmm arc nor um- Anrcnca and Umvcr.lt\ of Karu.:t\ Pn·"· I d\Henc~. Unwt•r
formly sprc:ad but IL'\tm:tcd to \Oilll' groups only, K.lll. (Principles, g.IMTOpods) 11 1C,J
through stntc.:tm,\1 comtraint. Burrowing. which Moore, R.C. (cd) (1 1J6Yb) Trc'ati1r '"' llll'rl1tln.tl Run
became po~sibk through m.1ntle fit~lllll, .-one obvi- Polt•tllllt>l.lgy. P.1rt '1. Vol' l-3 Gl•ologiltl So~Jet\ o 1117:
ous re,pons<: tu prni.lnon 'inct thl· MMR: avOtd- Amt·nca and Unl\'cNty of K.UJs:l\ Prt:"'· I ''' rcnc Ur.rd'h
.mce by attrvt• '" unmmg. shl'itenng 111 \eagra~s Kan. (Bi\·alvcs) \t>lllc
cornmunitic~. the rrwa'iOn of the.:· phy\lologJcall) Moorc, R.C. LaiJCkcr, <. .<.. .md Fi\dter. A G (I'J5J hnr~
'dtfficult' mtcrud.tl zone. colom7ttinn of deeper llll'rrttbmtr r..,s.<rls Ml (.r.l\\ Hill, Nl'W York. (U,dUl C.1llum
dtslU~sion of mollu,c.m .macmny)
water and hihh ti:n111dlly arc other' Th.1r these have .unm
Morron. J.E. (I ':167) Altlllllm. llutchln\011, l.orulon J>!rilo
been successful ,,,,\lL'gw~. despite contim1.1l preda-
(V.1luable summ.1ry <>f ucorpholob'Y and rt•l.ltwmluJ>l) (M.1nv,
non pressure, j, evttknl from the ~n:;lt number of J.livc:ll'au. J. (ed.) ( I'J5::!) I'mitc d< Jl.llt\mtcllo>l[ir, 11 l rllo;n.
drverse bivalvt:' livn1g tod.iy. ,\1,,/lmqucs. Mas~on .111d C1c. P.tri\. (~hnrter chm tl .unm.
/'rr m<e, but an C"\enu.tl <ompil.umn) //i.\l(l)
\aunder.. W.B. and Lwdm.m, N H (1~1'7) ,\tMrl I' W.
Plenum, New York. ( 14 nmnly b10logJ<.ll p.rpcn) 'B, A
J'homp,on, d'Arcy W. (ll/17) On c,,,ll'rlt .11111 l•m l dlonH
Books~ treatises and symposia C.1mbridgc U niver..i1y Prc.:·~~. Cambndg<·. (I ndivtdu.J .md Jt
d.miL work on phy,rt.cl l.cws dt'tcnmnm!-( b'TO\\th: n ( .nrlv,
U.Jthur;t. R.<.r.C. (I '175) Lar/J,marr \nfmrcul~ ,,,d rftw .1bndgcd ed1tion l1r tlm book wa\ cdtlcd b) JT No I
D1<1.1!• nnk f:lwvrt•r, AmsterJ.rm. (hnt· 'tmcturc of Bl!nn.:r u1 I<)(, I Jnd .1l'u publish.:d by C nnl.m:!!;r C'o~nt•r,
mollu~cJn \hdl~) Umver-.i('.. Prc\\) C r~·t.l\
Bibliography 257
Vcml(lj, G.J. (1'1S7) b•o/ulro>/1 .md l !sr.al.llftlll . • lu f.i.·,>l<>;<?i•u/ Cmcr. R . M. ( 1972) Ad.tpt.mon~ ot Bnn~h Clulk
H1sh>l) o( U/< l'nnn:ton UIU\"C:Nt\ l'n:,s. tl h,• evolu-
Bivalv1J. }<lltmrll t1{ Pc~lcomolc>~)' 46. 31'i 4()
non.arv 'arm' a.llc:' bc:t\Vl'Cil predator .111d pr~:y) C IJrkson , E.N.K., ll.u·per, D .A. I . J lld Pl•cl, J.~. 1'1'15.
Ward. P D. (I 'IH7) Thr Scltlmal HNtll) ••I :\i,mtdiH Alien T .t.xonomy and p.ll.leoecnlnt.") ol tht' mniJu,c p,.,,,,t/11'"'
~nd Unw111. Ne\\ Ymk. 1 1,.., pp
trom the ( )rdo\'ll'l.lll and Sthm.an of Scorhnd . L.tll•ll·'
\"icdnunn. J. 111d Kullm.mn, J (cth) I'JH!t C:rph.llt•p•>ds 28, 1111 -4.
Plnt'llf cmd H1•/" ~chwcazt·rb~rrt(hdw. ~tuttg.lrt C hlupk, l. (1 ' l7fl) l'he oldest I{Oilt.lritc f:aun.L\ and their
Woulk I I. ( l 'HI>) J>alolt'lllll<•l.•'(y. lua•, att"br,lft. 7ch cdn 'trati~raplucal \tgmtlcancc Lctlwra 9. JOJ-1 5.
C.unbridg~· Unm.:rslC\ Prc"· C;unbridgc (...,tudcnt Cope.J ( .'W (llJ%) . E.ul} Ord~Jvician (An:mg) btv.tiH·s
text, 'rill U\di.al) from thl· I IJngynng lnhcr, 'iourh W.tk·'· Pcllil<'tllll••lt'.<?>'
. 39, •}7<). 1026.
Davi~. R.A., l.111dmJn N.H .. D"mmmcrguc~.J.-L. rt .1/.
( l9lJ6) Mature modific:atlt)ns and dunorphi,m m
Individua l papers and other reference s .tmmonotd ccphalopod s, tn . lm11wmnd Prllcollic1/tJ.t:)' (~:lh
N.H. l.111tlm an, K Ta.n;lbl· Jml R.A J>.w1\), Plenum
.".llmln, I' A. (I ')HH) Pht"ph.att\e d sotr-bodu:d ~quad, from Pn:o;s. New York Jnd London, pp. 4h4-4'
the jUrl\\1( o,tord ( 1,1\ I c'th.ai.l 1 1 tll3- J(l Demon. f.) ( 19h I) The buoyancy of fi~h .md
r\JlnH)II, W.D .. Nich, J ( md Nl>m,, R.l'. (1990) cc:phalopod s. Pro)!rr.;,; in 1Ji<>J1ltys1rs 1111rl ClrcmiHI)' 11 ,
Dnlling and peclang ol turnrdlmc g,)\trr-, c>ds \lllll' tl1L 1 78-~J4. (Esst'nti.l l rcfen.:nn· onltt.unmu n of m·~.mve
LJrc C·c:uccuu,. [>.,J,,..,,IIft•/~)' 33 . SQ')-(, 12 bUO'tJIK)")
B1bm, C. ~nd Gmnerrc1- M uco. I.C. (191JI\ M1ddJc: l>c:nton. EJ ( 197~) t )n buo) Ull y .md the Jiw~ of mnd-
Ordo\ tci.m h1v.1IH•, fw11t \p.un .md rh car phylcuc ,mJ crn .md t()~~il ccph.1lopod' Crooru:~n l cnurt• JlJ73.
't' p.li.Jl'<1~cogr-.tplm ~tgruti,,anc~·. Hr/mwllo/oJ[y 34 . l (l'l-48 Proceniill~\ ,~f tlw Royal Soctfly H 185. 21 "\ -9<}
Bukrr. M J. ([I)'Ill) Th.: p.il.tt"nbJt•lug} ~Jf nc·nne.lcc;~n I >cmon. CJ and (,alpm-llrm \.n. J.13 (IW;h) On the
ga-troputl, l11~tcm ·,t/IJ1t>/cl'()' 3. 24'1 <•4. buoy.llll} of the pc;uly n.mnlus. Jou""'' of t/11 \/cmne
,, U1y~r. U. (1977) Ccph.tlupoden-~cpten f~:1l
Rcgclmeclu ni,men 111 ( rdl.lu~c .111d Sc:ptl:'lllb.Ju dcr
.., Biolt[(!ir<tl -lss<)(t/111<'11 •{ lilt• Uw'rcd Kllt,l/d''"' 46, 7~.'-S'J.
(Sip huncular ~truuurc ami funttion)
t 1\mmunitcn '\curs j,JIIrl•u<h /iir Crdc>gi•· 1111J Domma~'I.IC\, J.-L. (I Y90) Amrnono1d s, 111 J::.po/rltlt>llolr')'
e. Pa/iJ,·•u,•lo-?11' 155, I 65-21:1 (Ainnlllmh. ~e-pt.al func- frmd.o (t·d. K.J MrN.un.IIJ) , Bclluwn. Boston, pp.
tion') lli2-H7.
Blyer U .mJ McGhcr. ( , lt (19H.J) ltcrativ~ lVnlurion Donov:an. l>.T. (1 1H>4) CcphJlnpod phykogcny and cl.t\\i-
,,fM.JJic Jur.t\\lt nnmonate t~un.l\ . Ltthma 17. 1-lll. tlcanun BIIIIO,I/icill ~n·irw< 39 259-87.
& ~cn. R. ( Jl)(, I) Oxygtn botnpt• p.1leorempcr arun.: \)oyle. P. (1985) ~L·,u.JI dJn11>111lusm tn the- bck·nmirc
mcJsuremcnh m• Cn:t.t,cous Bdt·mnmdc .t fi·om Yotlll,l/lbcltts trom the Lower jm.1~~tc ol YorkslurL
Europe:. lnd1.1 .md J 1p.111 f••um,,J ,,, P.de''"'''"'l!)' 35. Pt11acttlll,•h~I/Y 28, U Hll.
)1177-lH Doyle, P (1990) I'hc Bnnsh To.1rcun (I ower Jur.hslc)
Bn..hh.J\\ M 11970) Th~: Jcnunon .1ntl IUU\t ul.nure ol beh:mm te~ . P.1rt' I .md 1. Palaftliii<W/o1f>llinJI Soauy
IOill~: MidJh: OrdtWKl.lll (IIJndt•llo) bivalvl'' trom
M''"0.~'''J!h, 1-49. 50 7Y.
rini\lCrC. Fr.ltl\'l' Palclrc"JI!I/c>l[)' 13 . (,2 '-45 Doyle. P. and Bt:nnetr 1\1\ ( 1495) Bd~·mnttt:s 111 blll-
C.Uomun, J.ll . (!QhJ) ~ntul dtmnrphi~rn 111 Jura~>IL ~trangraphy. P.lf,tro>lll<ll~)·. 38. H15-2':1
ammor Hn 1 r,,,,,,clfolh c>/ rll<' l.t ru"Jrn Lrll'fcll)' .wd Doylc, P and Macdon.tld, D.l.M. (lq'H) Bc:lenuutc bat-
PluiMnJihit'lll SClrlt'l)' 57. 2 1-5(>. tlctickk l...c:thaic1 26, 65-811
) \boy ~:x.unpks CJtcJ, \\ith lllc:thodoloJ:,"V) Doyle. P .. Donovan, D .T .mJ Nt.xon, M. ( I'J9l)
11 Ullumon, J.H. ( IIJ85) 1 ht evolutH'In of thc Jurasslc Phylogcny .md sy,tematic' of rh!.' Colt nadeJ. l.llll't'rHI)'
th~: •llllll<1mtt: Lulllly C.udann·nnd~.:, 111 t:.r•olutr<•llotr)• Cnsr 4 k.a11111.> Palacttlllltllll/li•ll Cclllllll•uri,,IIS 5, I IS.
Hi<l<lri<.i fr•'"' tire Pv.<s1/ Rt•wrd (cds J.C. W Cope ami I hik. J (1 984) Phvloge ny o f thl Nautlloulca .
I' \V. ..,kdton) Speual P.lpc-r; 111 l'.tb(ontulo._,") No. Pahll'ollltlh~l/ica P,•lt•lll<"•' 45 , I 219
13 1\<Jdemt< Prc". London, pp. 44-411. Engc~cr. I ( 19')6) The po,mnn of da<' AnllllliiHIIUC:.I
omun. J.H ( I'J<I'i) T llllt' ii·om f(l,"k S. ~ Buckman
'""·
lu1l ;nd Jur.tssJC h agh re~oluttun ' LraCij,.'Tdphy, m Al i/nl<lllt'S 111
wi rhm the C~:ph.lhlpodJ, m ~1111awnmd PolmluoltJ,I/)' (l'd'
N.! I. I .mdnun, K ran.th~: and R..A. J),wis), Pi<·mmt
Jll c,.,,f,•~)' (ed M J le: Ua, . (,cologJ<.tl )one£\ Mcmmr Press. Nt'w York Jnd London. pp. 3-l<l
j.T No Ho, pp L!"'-'50. l rben, 1-I.K. (I 'Jh6) Lbt"r d1c: c,..,prung dt'f
oge ..11\rr. R.M. ( l'lfl>l) Functaonal srudte-' on the: Anunono1d ea. Hlt>lcll/li<l/ Rti'H'H'S 41 , 641 SH. (l:..1rlrest
C'rctaccom oy~ra Ara,>.Hn·.J. flt11at'OIIf<l/,>~y 11 . 45H-85. ;111U11 0110id~)
258 Molluscs
Fischl·r. A .G .•md Tetchert. C. ( l'lh'll ( uncrJI deposits in H ome. M R ( !9RR) E...-..rintrron .wd 'utYI\ al m the l tlw
ccphalopnd ,hc:Jh lmr•fmty , 1 1-...w.<IS Pait'tliii<•I•>J!iml Cephalupoda, m £wruc1it111 .m.J 'irtn'll'•r/ 111 rlrt I G
Ccllltri/•rUWIIS 37. )-}Cl (C.mtt:r,tl lltO'l-'hoJob')' .md U\C Record (l·d. G.P. LlTwood), ')y,tt·nunn A,,o,ratu D
L•lit'
.1s cuuntcrwcJghts) SpeC'I.ll Volume No. 34, pp. I 39-9~ bi
f-Jo\wr. R.ll (1lJ57) N.umlmd' nf chc 1'.11Jl·o7oic, in Hu <bon, J.D. (1463) The rcco~mtion o t qlrnll:y-.on- 5.3
7 rt'dlr <t ,,,. Alan"r r:.-oh~~y ,,d fJ,Iftlt"c>cwl.~!i)' 2 (cd. H.S. trolll·d rnollu\c ,rssembl.1ges 111 thl· Grl'.lt btulnn
l lbl'•
Ltdd). Geolu~o.l Society uC Aml·rk.l Mcmo1r No. 67, Srnes (Mrddlc JurassiC} of the lnno:r llcbnde1 ht
Gcologrl.ll \onety of Aml'nl.l, I ,\wrcncc, Kan .. pp. P.llat'OIIIt•IPI/Y 1 t , l63-R2. (Set' text) 21
H21J .,:,:!, (Fcologv .1nd .mnor:ltcd brhlrngr.lphy) JdTcnc,, R.P.~. (1%0) Photoncg.Jtrvc youn~ HI th lllje<
(,mrld, '-, J (I I}&Q.•) An evQ)unun.&f\ microcosm: Tn.1ss1c I.Jmdlibranch Lima /irrmlol (Schll)thcun). I'JI<lf lw
Pll'I\Ullenc and R.:ccm lmwl) uf th~ l.md ,n:ul P. onto/Oil)' 3, 3(12-Y. (U\e oflunulc ,r\ .1 brood rh.lmbrr) l 1ml
(f>•••·.-rf,,z,•mtl'>) m Bcm1ud.t Hu//t'trll 4 tile \luserw1 o.f Jeffl'ne,. R P ~- and Minron. R . P ( I'Jh;i) The 1110<!~ of 1'.7
(.OIIIJ••Ir•llll'<' 7..oclh~t:Y· Ht~mml l·mrwsitt 138. 407-532. lilt- of tWO jUI<bSIC 'peu~ of f'c>.'hft•lli.l (Bl\~hl~ :\1lA
(Allop.rcric \pcl"i.rtion) P.rl.lcoi!Wio.~)' 8, 156-HS. (Pianktomc mode of hie- pt»- h.l
(,ouiJ. S..J. ( l')o9b) Ecology .llld tim.nornl c;igmticancc tuiJtnl) hi•
tlf uncolling in I cnn<'tll/omcl 'l'"•lltl" .m C\~.1} on bra~tro Johmon. A I .A (1'}94) Evolurion of Europl'JJI lowrr Sllj
Pl'd ti.mn. Ur~llctm •!I .\l,mrrl' So~t·rrn· 19, -+32-45. Jur,~.,.,,, Grypluwa (Grypllaea) and contl'mpnrdn'''lll M cl<
(Unwilin!{ usdullor r;rprd upward h'HlWth) biV<IIW\. Hisrm'im/ B1ology 1, I 67-H6. ~p.
! larpcr. E.M. (19l} la) Tht· rok nt prcdannn 111 the evolu- J o hn~\111, A. L.A. and Lcn non, C.D ( Jl)C)(J) Fvulunnn ul
Zc:
tion ,,f rcrnt·ntanon in bJValvt·~ . l'.r/m·o>ll/cd<\~l' 34 . gryph.reatc oysters in the Mid-Jur.t\\lt of Wt'l~rn M.IJ...<
455 r.o. Europ~. P11lo11'c1111ofogy 33. 453-4K6.
.1111
I!Jrpl·r. E.M (lll<llb) Po\t-l.lrvJI tcmenht.mon m the Jonc~. J.~ (I YH I) An uncensored p.r~l· ut lu"il flllt<JI)
(C
(. ),rrcrd.1c md w. •mphc,ltlnn' tnr nth er remenring Nawn· 2 93 . 427-8. (lntl'rpretauon of W1ll11msvn C.I
br~Jh ~~- Jou,wl •>/ \lc•//umm ~tu.lrt 58 37-47. 11J81) :Vbru
Harper. E.M . ( 19<J7) Thc muJiu,l.lll pcno,tr.lrum. an Kautim.nn. E (., and ')ohl. N .F ( llJX 1) Rudt,t\, 111 Thr on
tmporum COibtr:Unt on bwa.lvc C\'c1lunon. Palart>lltolo,~)' h1rytloJ111t:dra <?f P.rlarour.>f,~I!Y (ed, R. W Fairbndb'l.' and uf
40.71-117. C. J.tblon~k.i). Dowden, Hutdnmon .md Roo Mulv
llarpcr, E. M. md Skclton, P W (I •N3) The: Mcsozoic Stroud\burg, Perm lk
M.mnc Rcvoluoon and rprt:lun.ll hlV~Ivc~. Smpttl Kelly, S.R.A. ( I<JHO) Hiarella- .tJur,rs\lt btv.rh-t' '<lll~tt~rl Ill\
Gcolor:ira Sprci.Jilssuc 2, 127-';J Palm·ttr~rc>ltJ.I!)' 23 , 769-R I .
~k
llarper, J A.•md R.ollin.s. H.U. (l9HS) !nLillnal or scnli- Kcnnl:Jy. W.J., Taylor, .J.D. aucl 11.111, A. (1'16!! Mul v
urf.~t~n.ll bcllrrophont g.1stropod' .uulvm of l:'uphcmitcs t.:.nvironmcnt.ll .md biological co11trol' on b1v,tlve 1hcll ~ht•
.md luncanonally related t.tXJ. 1..1'11111111 18. 21-JR. nuncralogy. Riolo.~iral Rfl'h-u·~ 44 , 499 5 30 (Shell mw P.rr
H.1wpnm.1r <~ (I '}XH) On the origm nt lllJJor !,r.lstropod t'r.tlogy ~Ubjcct to man>· controls) Per
gruup' "' nh ~rniJI rdercnl"e to the Strophoncura. Lmlo:ux. H . (1tJfl9) L"' rcntJcuhtc' d'Europt' ncodcmJle ( ;('(
JoutJIIII c!{.\!d/umw Swdrn 54. Jt>7-·H I. et d'Afnquc du nord. Ca/uw dt' P.rlt:o,r.•lt!~'r C:dull'IU ([),
Hcwm R.A. and We,tc:nnann, (;,{:,(; , (llJH6, IQ87) C.S.R.S.l 238. Ntxor
r uncunn ot complexl) Aucl'd stpc.r m Jrrunoumd L.tmon, K. (11J79) Silunan tentaculmd;, from Gorbnd and .11111
,ht'lls I Mechanical Pnnoplcs Jlld furKnonal model~. Sc.m1a. 1-oHils and Strata 11 , 1-1 HO l...t1
11. St:ptal evolutiOn .wd conclm11'm 1\'n1rs Jolfrrlmcll fiir Lchmann. U. ( 1967) Ammoniten mit KtcfcrJpp.tr.u und Ne
Gn>/!11/11" mrd Paliitmtoh>gu Ah!llltrdiiiH~c·rr 172, -17-69; R..tdula ,JUs Ti.lS Geschiebcn. PaliitliiiOhJ.I!Isdws ai!Sdmh l'Jl k.l
174 , 135-69. 41 , JH 45. (F1rst dcsc:nption ofJaws Jnd r~dub) CO l \
H o ll.md. C.H . (19R7) The n.luttlmd (l'phalopods: a Lchmann, U. (1'J7Lt) J aws, radulJ md crop of l mioo<w (h ·
~tr,m~c '>UCCC~~- jouttlfll <?f the Cnlltl.~IC•II -"••ric•ty of Lmdvn (Anu no no idca). Paltli'OIItolol!V 14 , .338 ·11. (UnUiu.ll t:t.•p
144. 1-15 prC\Cllt.ltlOT1)
l'~d.
llollJnd. C.. IT. ( 1988) The P.1pcr N.lUu]u,..Vtw Mexico Lehm.rnn, U. (I 'J71 b) New .1~pccts m .111111l<llllte biol•>to"
Ulf'
Burr.w of ,\lmc.s .md ,\l,rm.l! Rest.llcil 44. I 09-14. Pron•t·driii/S t!l tire Xortlr .-illlrnc<l/1 P.rlco>mol,'gll• tlu•
llou\c, M.R. (19XI) On the origrn , cl.t\\ificauon ,md C<'m•mtwu. Vol. I, 1::!51- n9. (A' 1bovc, \\ith rntcrprr-
l'<"!Jl't.l
c\·olunon of rhc e.rrlv Ammtmoid.:-.1, Ill 17Jt tauon)
l }P
.-lmrrlt•uoHit·tr. Tiu· I::.r•,•lutit•ll. Cl.t;sr/rralr•'" .\lode <?f LJ_(t., Lchmann, U. (1 'J7Y) The pw'> and radula of the Jumm ,3(,_~
•md Gt'cllt~~~rml Us~Jitlrrrss c>/ a ,\1,1Jtll !'.>m/ Gr,>up (ed; ammonHt: Dtlt tyilt)(cro.;. Polatwrlt!l!l,~l' 22. 265-71 • I'I~Jl't.l
M.R I lomc andJ.R. Scmnr), ~ystl'111.1llc\ A"!iuciacion Ldm1ann, U. Jnd Ku.licki, C. (191JO) Dllublc funcoon of
nf 1
Spcc>,rl Volume No. IR, Ac.Jdl:IIIIL Prcs>, London, pp. aptychi (Ammonoidca) a~ jaw elemcnrs and opercuh St1u.
3 3(, (Srill valuable) /...et/wirl 23, 325-32. P!.~Jt't.l
Bibliography 259
l1~edahl, L ( 19~4) S1lunJJl sihnti..:J h1V:1lvcs from Rt>Ht'llCt>mh .\1t>lfusk:> .md tlu Early Hislt>r)' •!fthr• Phylrmr
Gotl.md. SPrn.I/O <:c,>I~Jl'·lka ( 'ndm••klllll.i/ 78, 1-H2. ,\ fnllusca. US Geolng~c~l ''lllrVt')' Profes\Hltl.ll P.lpl'r No
l il_]l'dahJ, L. ( I'IH'i) Ecnlog~~:al ,l,p<.:Ll\ ol a ~ilicific.:d 1J6H. (Rostroronch~ as bJVJive
Jncestors)
biv.tlve taun.l from the Sllunan of Gotl.llld. LNiwitt 18. Ponder. W.F .•md Watcn, A. (1988) Pro,,,br.1nrh phy-
'i}- (,fJ. lo!(l'HY. !.la/.,cc>lo.l!•ca/ Rwit·u•.; <;;upplt'lllt!lll ~
L1l1ni.1hl. l. (I 'I'J 1\ ( ontr.Jsnng fn·din~;: str.ucgtL'S in R.aup. D.M. (l'Jnn) Gcomcrnc:1l an.1ly'•' ,,f,ht:llu,iJin!(
b1vJivc~ from thl '>1luruu ofGotbud . fltl/,1t'c>ntt>l~)' 3 4. gt'm"Tal probletm }•'""'''/ 4 Pttlt·olltt•lt>gy 40. 117&-'J(I
.219- 35. (:-.cl' text)
l•bt:tidhl, L. ( 11N4 ) \1lun.m nuculoJd md mod10morphid Raup, D.M. (lQ67) C.t.'nmctncal analr~" 0t shdl wtlm~:
lnvalve~ frum 'iwcdcn. h>ssils am/ '\tr.Jitl 33. 1 -89. th..: cephalopod ~hell )•lllnlllll!f Palet>tll,,f<~l!)' 4 1, -IJ-{15.
L1n~lcy, R .M ( 11177) Solllt' 'laws' of gJslropud ~hell fonu. (Sec text)
P.ll,•c>biolt!I!Y 3. I 'If,. }Oh. Raup, D.M. Jnd Sunky. S.M. (1971) rtw ccph.alopod
M,AJcstcr, A I ( l'lh:!) \y>tcnuncs ..1flinihc' .md lift: 'unm: probll'm, tn f>n11tiplt•.' ·~f p,,/c,,,,,,f.~I/Y· Frccnuu.
h.1bm of H,,/>,l.:,r. 1 tr.m~monal Ordli\'ICI.m lucmoid ~an Fr.mci~ro. pp 172 HI. (A ~ood trt'.Jtmcnt of tlm
l>w.tlve. P,,/,,,..,,,.,f~)' 8 ..2JJ-41. (Muluplc mmclc sea~ th~me)
T \U~L''t .1ffirultc' wnh w~o:mcmed ancc'w~) R.c1i, W.E. (I<JX.)) The Stcmhenn ~na1ls (M10cene
\ld<.UHIOn. D.l . (I 'IH2) Tu.mm~''' paf'lll<l, n. gl'n and u . Schwab1\chc Alh) fmm J neo-Darwmian pomt of
sp . .\ late M1ddlc ( '.1mbnan pel..:1..ypod from New view: J discussion. Pc~liimlh•lt!l:iscllt' Zl'itsdm{t 57.11..2-25
,r Zr:ahnd. Juurwd 4 Pt!leol/tolo(!}' 56, SH'J 'JH. (I lilgcmlorfs ~naih sec text)
1\ M1kow,ki, H. (I 'J6 )) l'roblem~ of 'nu.tl dimorphism in Rtlllncg-ar, B. (I <JH '\) Mt)llmcao phylogeny r,·visitc:d.
.umnomte\. 1, ''' JJ,tfarwrt.>lt>l!tta l'olt>lllt<t 12. 1-92. i\lt•moin 4tllc .-lu<>!IIIIWII tt[ .-iustrdlillll Pal,lrt>llhl/tl'(tsL' 1.
..... (Comes imlcpt·ndt·ntly to the 'Jilll' l'Onclu~tOn, as 121-44 (Most recem trcJtrncnt)
n, Callomon. I'lh \) Runncgar. B. (19')6) l::.uly evoluoon of Mollu'< !I ' the
Mutili, D.M. ()'I<)()) Prl'<ltnon on K,•.\llh>a·rrl> by semt- I0\\11 record, m ( )r~ll"' awl hv(l/IIIH>IIII'Y R,,lt,l/tt>ll <>(tin
"lit' oncnci fi~h in the 1\hddl..- Jur.b~l.: Lnwcr Oxt(>rd Clay Afr>//um1 (ed. J. T.1ylor), Ox±ord Um\'l'~lf) I' res\,
ud ofEngland. Palrtl'tlltlt>f,~l!)' 33, 731J-42. Oxford, pp. 77- H7.
1\S. Murwi. H. (1971) The s1phona.l lulw in Jura~;ic Ru1111cg.tr, B. and Hcntlcy. C. (141'!2) An.1to111y, ecology
llelcmnltld.J .md AuhrtK<'rid.t. Bu//t'llll 4tln Gt•olo,gmrl .111d ,1ftimtie~ ot the Australian Early Cambn.tn b1\.tlve
l11stiwtt t1[ tllr I 11111'"'''}' 4 l'pps.llo.~ 3, 27- \t>. (Dewilcd Pt•frl•ll<l nmm·l!an Jdl jmmwl •?f Paltwtt<>f,~~~l' 57, 7\-92.
tledron HUlfllb'T~ph\ ) Runncg.1r. B. and Pojet.l.J . (1'>7-l) MoUmcm ph,(ogt•ny:
8) \lutvcl, H. (1'>7..2) UlrnsrmcturJl 'tudit."'. un ccph,tlopod the pJlaconrologtol viewpoint. Sncn(<' 186, \11 - 17.
odl >hdk Part I Tht' "'f'£.1 .md siphono~l lllbl m 1\'tmtilus. Runnq..~r. ll., Pojl't.l, J, Morris, NJ. tl ,,/, (llJ75)
Ull- P.m 11 ClrthtllUJUC ccplulopnds !Tom the Bwlugy of the Hyolnha. Lnhaia 8, Hll-'J I.
l'cnmylv.miJn Buc:khnm AsphJit . Hulll'tiu of th~ S.avazzJ, 1:. (I 9X7) (;comcrric ;md funttinn.al LOmtramt~
Cr,,f,~,giml btWIII/t' t~lthr• L'nll'cl'sity .{ l'pfl.'"''' 3, 237-7J. on bivalve ~hell morpholo~. L:tltaia 20. 2'J3 106
(l>cuilcd UULW~'l.lphy) S.IVa:UI. E. ( 1990) U•olog.c;tl ;lspcct~ of thl'nrCtll",ll ~hell
'lltxon M. ( l 'N6) MMphology of the J-IW~ .md radula in morphology. u·tllcJI<l 23. llJ5-212.
.md 31'111 onoid\, 111 ,,.,,,,,,,;J p,,/cLJblc•lt•l!r (l'd-" N.H. 'it•II Khcr. A. (I 9S4) Okologte der tri,l\SI\ch..:n Mu\chd
l..lndm.m. K r.lll.the .111d R.A. D.lVi>). Plenum Pr~~. l..1m•1 lincar.1 (Se hloth) und ihrl'n Epi\kt•n .'\rues
:-le\\ York JnJ Ltnldon, pp 2.H3. j.J/rrlmch Jiir G!'c>loi!W 1md Pt~liirmrolo.'!. ll' ,\(,,,,lf.ll~t:{tc,
flltlurd. A. (1'172) ( tph.lloporl.~ .llld fish. the limits of 163 ~3.
cunver~ence. Bll>itl\!lr.tl Rt•l'it'IF.' 47. 241-307. Scil.lcher. A. ( 19611) Eprzo~ns ,1, ,1 key to Jnunono1d o:col-
I)Ctl<l) 1l'volunon.1ry llllt'r.aLnon~ berw..:en fi~h .Uld •lh'Y. }cm mal (!f' Palcvntololiy 34. 189-9.\. (Set• rcxr)
u~ual ClphalopoJ,; .1 \ ,llu.lhk· work) St.'dJchcr. A. (I%H) ~w11nmmg h.1b1t~ of belemnitcs -
Ptc • J S ( I 'IH!I) 1\ n"'"" Stlurian r..:tr.lctilc mouopi.I- rl'corded by bum1~ b.uuades. Hli,ll't>gc·•~Jlr.1p/Jy,
tklgy. C•lllhoran md th< ,,nh'111 of tl1c: ~.1\tropo'h Pnl(t'fclm.~; of p,,[,,ro.Jmwt;>/t'i{y, p,,/an>t'u>l.~y 4 , .:!79-85 (Suggc~tion
lt>.~:"al tht Ctoh'IIU<~I .-J~,,>o,lliml 91. 'J 1-7. tlut the guard WJ.\ in hfe exposed)
·rprt'- PU~l.l. J. (I'J75) h•td1llr1 rmynrm llarr.uadl .md ..:arly pelt.'- Skdtnn, P. W . (1'J7(>) la1uction,ll morphnlog}- nf the
, ypnd phylngell} . /Jul/,·tltli •i AlltWC•III /',J/t",>llltJlc~!?Y 67. I hppumidae. LL·tlt<IW 9 , H3-l 00. {Pt'lior.lted 'lrd'
3(,,HH. ,1llo"' cd inwes~ of w,ltcr wrrents)
l'oJ~t.a,J. (l 97H) ThL on~n ,md ta.xunonuc· divc11>tfication Skclton, P.W. (197H) The l'Volurion ofti.mLUon.ll dl.'si'b"ll
'f ~)ek·rypntk Jllti/,>.H>JIIaim/ Tralls.1111t'11-' 4 tilt• Royal m md1~t' (H ippunt.at..:.l) and Its wxonom1o.: unphcJ-
\Jtt<'IY t~{ L.>mf,,n l1 284 . ..225-46. tiuns Plult>SopiiUal Tr,m>.UIIt>n.< •if tllr R,')'•ll <;,>,ll'ty ·~I
ru. J. and Runnc~.tr, B (l'r"c. ·n,. (J,/,,c,>molot:>' 4 Lt•11drm B 2 84 . JoS-1~ . (Gt'ometry ofwowth)
260 Molluscs
Skdton, P.W. (1985) Pn:ad.Jpt.mon and cvolunonary Pn·"· Ltwrent.e. Kan.. pp. It> 1-2111. (ClamhCltlOO w
mnov.1nun m rudr\t brv.llvc,, m L·r-.'/lltHln<try C<l.>t' used 111 dm chapter)
1/w, nt'l fmm tllr /·ossrl Rt'ulfcf (nl> J.C.W. Cope and T honm, R.D.K (1975) Functional murpholob">· i.'<'OI-
P. W Skelronl. Specral P.1per-. m Pabconmlogy No. ogy. and evolutionary con\t'rvauon in rhe
ll. Ac.1dcnuc Pn-.,~. 1 ondon. pp I 'ill 171 Clytymcndac (Bivalvia). J>alac,,t<,fc,g)' 18, 217-5~
Skelton. P.W . .rnd Wnght. V P (1'11:!7) A C.1ribbc.m rud- Trucm.m, A.E. (1941) Tht· Jrnmomtc bndy th.tmber Wt
iM btv.tlvc 111 OmJn: i'hnd hopp111~ acro>~ the Pacrfic with speci~l reference to tlu: buoyarlly aud mode llflr((
m dw brt· Cn:raccous. P.Jiat't>llfllft)~y 30. 505-29. of rht: hvin1-1 a.nu uonitc. Quarterly }t>llllllll <>( the
S111ith, '-o.A., Tluycr, C. W. .md Urerr. C E. (191:15) Gt·vl.~l/il a/ SctCH!Iy '!{London 96, 33?-HJ (l'l.l\~lt m•dy uf Y.u
l'rt·,l.rnnn 111 rhe Pai.Jeo701t g."tropud-like drillholes funcnona l morpholOf,"Y of umJ~u1l 'hdl ,J~.~p~~ m I
111 Dn·nm.u1 br.tChiop"''h "W~tiiCt 230. IIJJ.'\-5. .unmonord') l:
StJnk·~ S M (I !JbK) Po~c-PJ.I.Il'Ol<lll .1d;~pcin! radiation Vernlt'll. C..J. (1983) Trace~ Jnd trend\ ofpn·dmon. \\1th Vot
of mt.nm.1l btvalve mollu,,y .1 "llt~nJUl'rlH.' of mandc 'Pl'UJ! reference tO hh·:1lved .minu)\, P oll.rn>llltlh>gy 26, ,,
flhl<)rl :tnd ~iphon f(.lnllJtiOil . /c 1 1f/ll,r/,~/ J'Jft'OIII<lfOflY 42, ~55 <.6. f
21 ~ 21J. (R.1dianon depcndctH on <lilt' kc\' ch.uacrC'r) Vogd, K. .md Gutmmn. W ( I1JKO) The dcnv.m•m of
StJnky, S.M (I 'J711) Rd,Jtll'll "(.,/idl/<>rrll re> life· lta/lits ;, tlrt· pdt'lJ'Pl'u': role of b10mcch.111lC\, phv,iolngy md
Bir•.r/l'i<l (,\1,,1/mr,l). Geoln~;i( 1! 'nKrl·t~ of Amenca env1ronmcnr. Lrth.1in 13 , 169-75.
Memn1r No 125. G~:olo~t<ll Suu~:cy of Amenca, W.1rd, P I) ( 1979a) Cameral hy111d 111 ,'\/mHt/11~ And
L~wrt·nce, Kan. (Extendni trc.nmcnt <>f Recent .mrr l lunite~. Palcobiolo,{/y 5, 40-'J.
b rv;rlvr functlon.llmutvholo~;Y) W.1rd, P. D . (1 979b) Funrtmnal nrmphnlogy nf hdic~ll)
St.mky ~.M ( 1972) Functiun.ll ulllrphulogy and cvolu- co1led :unnwmte ~heUs. Palenbwlt~l/)' 5. ·11 5 22. (MnJc
tmn ot by\S.tlh Jttac.:hcd bw.tlw molllt'k' }•rrmw/ <?f' ot' htc lllVolvc\ wrrical trub'T'anon)
f>cliC<•IIInlc!rtY 46, I h5-:! 12. W Jrd. P D (I ')80) Comparanve ~hell-,h.lpl..' rlr,tnbuoon
St.mln \.M. (1'1-7) CoJd •pt.ttlun 111 fngonrrdac; a 111 jur:J\\tC· (. 'rctaceou~ .unmonitt"> .md Jur.l\,tC-Tl'rtiuy
r~·m1rk tble f.rrnily of burro\\ in~ hi\ .tlvc,. P.rl,lt'i>trtolt"(y l\JIItlhtk Paft·c>l>icl/flgy 6. 3~-43.
20 l'lh'l-lJll, (Well '' onh n.·.tdm~ W.ml, P J) (1996) Ammonoid cxtJ.n..:tHm. 111 -~"""''"'''d
'-otcnzd. H.H. (1'171) Oy~ter' /n•otisc ,,, lr111crtcbr.u, P.rlcc>l>~t•lor:y (l'd' N.H. Landman, K. I ,111,1lw .u1d it A.
p,,{,"''""''·~~y. Part N, Vu! '\ Gt·olo~ttJ.I Society of D.rv!S), l'lcnum Pres~. Nt•w York .1nd London, pp.
Anu:nu .mu Umvt·mrv of k..ul\,1\ Prt'''· Lawrence, RIS-24.
K.m. (A full .1ccoum of morphnlngy, evolution and WJnl, P.•md M.trtm, A.W ('197H) On t he buoy.mC) "'
t'n1((,~"V· dcals sucrinnly wllh l'Volutmn in GY)•J>lracn) the pe.trly n.1ut1lu~. Joumnl '!f E.\f!Ctillll'll/<11 7.oi>lt~~y 205.
Strithhcrg, !), ( ll!X I) Apertur.tl comtnttlllnS rn ~ome 5 12. (Es~entr.1l rl·.tdinw.
Ulllt>.:t•nd tepJu)upod\ u·t/hll<l 14 , 2fl1)-7fl W ccdnu, M ( 19'lO) Shell strut:turl' and .1ffumy nf Vl'mn-
Stnd\bt Q~. S. ( 1')K4) A ph c hop,•d pl.llc' - j.l\\· dements or ti)ml 'gNropods'. Lrtltaia 23. 297-'\ Ill
prott'Ctl\'e opert ulum L:t/i,,,, 17, '1:\-!t Welt... M.J, Wclh.J and 0'11or. R. K (1992) Lifi:adu\\
Stml\lwrg, S (I IJKS) Silun.m onwn ml n·phalopod~ from O:\)'~l·n levek the btulogy of Sml/i/us pclllrpr/rr, and
(,od.md [7,,_,.si/s JIIJ ,\fr&Jt,J 18, 1-h:; extinct !'i.lnn\. Jounral if tire .\lanm Hrc>l•'~'"'l .r-1Sfc~MII,n
"irndsh,·rp S (I ~':Ill) OnentJtmn uf teplulopod shells tn c>f tilt l 1mtt•d Kmgc:lom 72, 31 3-2K. (llabtt,lt uf ,'\~ru11lrr
•llu,rnriorh Pnl.l<'<'llln/,,1/)' 33. 243-l!<. a.nd .1mmnnttes)
Stlirtlll..'t, W. ( 11170) ...,oft p.lrh Ill rnlob1tC'~ and We\tenuJnn. G. (1985) Pmt-morTt·m dl•,ccnr wiTh 'cpul
ceph tlopods. Sdnw 170 , 1300· '· (Snh ris~t• e' external 1111pl o~ro n in Silurian n:mnlmd\. Palilcm/11/,,~;,,/II'J
to shdlnf DevoniJn orthut ont.·s) Y..citJdrri{r 59, 79-97.
t;wan. R. ll. ( 198(>) l kterodunml ln:nth in Namurian W ~:\tenmnn, G. ( 1990) New t!cvclopllll'llts 111 c:colob'V <ll
JI\UliOIH)rd cvolunon. fl<~lmollt•>l•)l/)' 3 1. IO.H-5~. j ur,\,~1..:-Crl'taccous ammono1d~. , lrtr dei SI'Cmrd
f.1ylor, J.D. and Ltym.ln, M (1'172) The rncchJnic,ll Ccllll'(!!llt) lmrma::rollalc FM.1rl1, l'r•t>llllll>llc, .rlmlllt'llf
pmp•·rttc' of bt\ .lln· (MnUl"'·') ,heU \trucrurc~. Pa..:ol<~, 25- )0 October 1()87, pp. -l59-7X.
l'.r/•lt Mu,>J,,~y 15. 7 3-R''. W ntenn,mn. G. (1 1J96) Ammonmd hti: .lllct h.lbltJt, rn
l'a\'lor, J.D .. Kenncd). W J .llld ll1l1, A. (1969) The sheU tlmmtlllclrd !'.Jie,>hic>lt!j!y (cd' N 11 LJndrn.lll, K T;~n~bc
~mtcture .md mmcrak>):.') ot dw Ui,·ah 1.1. lntroducnon. .mu R.A D.1v1s). Plenum Pn.·"· Ne\\ York ~nd
Nut'lllJn:J-Tn~oman...J Brrlktm ·~f till Hntr<lr ,\/ummr o{ London, pp 608-707.
7-Mhl!)' \uppl. 3, 1-125. (:--.t1ntbrd work) Wt~:dlll.lllll, ]. ( J'J69) Th~: heteromorph' .md .1mmonord
t ,·ll h,·rt, (. •. (11JC>7) M~JOr ti:.1tun:' tll n:pl1.1lod evolution, t':o-tmrrmn. Bioltwiral Rn•irtl'.\ 44. ')(,'\-hll2 (hrll dotu-
111 f "'')'-' 111 P.tldcllllto/,~1/Y ''"cl Smrrr~r<~J'Iry, (t'd' C. n rt·rrt.rtmn of hetemmorphy .1' .r tuullwn,tl plwnom(-
Tc:1dll'rt .md E.L. Yochebon), Unrvc-r~rcy of Kans.ts non)
Bibliography 261
\\rdbur, K.M. ( 1901) Shell formation and regeneration, Kan., pp. I 41 61. {Bellerophonods may not have been
m P/Jysiolt~f!Y «?f .\.fol/usca, Vol. I (ed~ K.M. Wilbur gastropods)
and C.M. Yonge), Acadenuc Pre''• New York, Yochelson, C.L. (1979) Early rad1ation of molluscs and
pp. 24;\-81. (Sccreoon and development of bivalve moUu\c-like groups, in TI1e Ori.~m of Ma;or lrll'rrubrate
\hell) Gror1ps (ed. M.R. House), Acadenuc Press, London,
Wilhamson, P.G. (1981) Palaeontological documentation pp. 323-SH.
of spec1ation in Ccnozoic mollmcs from the Turkana Yochclson, E. (19H 1) Fordilla troyensis l3arrandt·: 'The old-
Da~in . Nature 293, 437-43. (AJJopatric spcciation) est known pelecypod' may not be a pclccypod. jounral
Yanccy , T.E. and Boyd, D.W (19HJ) Revision of the of Pa/eoutolo.f!Y 55, 113-25.
AlatoconchJdae: a remarkable f.unily of Pcrmian Yochclson, E.L., Flower, R.H. and Weber.., G.F. (1973)
bJVJive,. Palaeoutology 26, 497-520 }he beanng of the new later Cambnan monopla-
Yochcbon, E.L. (1967) Quo varus &//mlpho,J? in Essays cophoran genus Kniglrtocouus upon the ongm of the
m Pal.lt<llltolo.f!Y aud Srrat1.~mplry (elh C Teichen and Ccphalopoda. Lerlraia 6, 275-310 (Early drversification
L.l. Yochelson). UniverSity ofKan\3\ Press, Lawrence, of ccphalopods)
"
IS
al
..
es
of
do
te,
m
be
nd
;lid
:u-
1C-
ThiS 1
Echinoderms rct.unc
evt:r, 1
dcnvec
the ear
hcrwet
(clcurh
LOITI.'Ct
4liCilCC
(cnnoic
urchm~
Ol: WO(
d,Js~ific
rillStex
The f.umli.tr slarli~h and \C,l urchins wluch arc so A modem ~ysLe 111 of L'C hinoderm das,itllaOon, 21 non 9.f
common in \h,tllow watt·rs arc rcprc~cntati vc of dcfmcd in the Trcarrsc a11 flll'crlciJmtc Palcontology aM there a
an entirely mmn~: phylum. the Echinodermata modified by Sprinkk ( 11)76) i~ a~ follow~. marked
(Cam. Rcc), whtt h ~t.llld~ ap.lrt zoologically from tla~~es i
nearly .tll othn inwrtdn.lt\: group~. Echinodcmu; of tu corn
all kinds luvc inn:ru.tl mesodermal skeletons of SUBPHYLUM 1 ECHINOZOA: ~~~dim: t•chmodenm, usulU
globme or d1~tnid.1l arc hart
porou\ t.llCltt· pl.un. whith .1rt• nonnally spiny and CLASS 1. ECHINOIDEA (t )rd -Rc·c .): Sc.1 urchm,. .1 smglc
covered outS1lk .tnd 111 bv .1 thiu protopL!smic ~kin. CLASS 2 HOLOTHUROIDEA (Ord.-Rt•c.) \t:a curw11~ llut the
Nonuall:r the skdl'tom h.tvc ~ livt·-rayed or pen- CLASS 3. EDRIOASTEROIDEA (l <: un Carb.) r haractL
tameral symmetry . though in some fos)il groups known
tlu~ 1\ not so .md 111 somt· modt·rn and tos~il ~ea There arc ~cveral other cxtmcl classes. et: hmoz
urchms a btlart·r,tl symmetry j, supcrunpmcd upon
the: radial plan Anothn unponallt l~aturc of cchin- SUBPHYlUM 2. ASTEROZOA
odcnm is tht: water-vascular system : .1 complex CLASS STEUEROIDEA (Ord Re.-.).
intc:rnal appar.nu~ ut tubt:s tnd bladder~ cootauung SUBCLASS 1 ASTEROIDEIA (Ord Re.-.) St.uttih.
SUBCLASS 2 OPHIUROIDEA {Ord Rn.)~ Bnnle \lir~
fiutd fhis ha~ t'Xlt'll\1011\ whid1 t:mcrgc through
SUBCLASS 3 SOMASTEROIDEA (Ord.-R.t:t. ).
thc skdcton to tlte oul\tdt: .1s thc tube feet or SUBPHYlUM 3 CRINOZOA 'Pdm.ttozo.lll~·. i c. cdunoJm:.
podia. Tube th·t h.tvt· vanou~ tlHU:tlom, espeCially havll1g a ~m.tll pl.ucd body (oly') ILxcd by J ~tcm. and vo
locomotion. respiration .md ti:t:diug. They may be pinnul.m.· Jnm ,1l1Jp!l'd for f~>od ~adll'OII~
consJdt·red .ts .11l-purp1..hc orgam mt'd by ammals hv- CLASS CRINOIDEA (M CJm Rn.): SeJ-I.thes
mg tustdc a l akitc box tm mampulatmg the envi- SUBPHYLUM 4 BLASTOZOA: 'Pl'lnwozOJm'. often ''"lk
penpr
ronment. l,l('king free .mn~. hm w1th lm ... n.t1 brachioles for fond FJth
ering. ,md otrcn v.mllll\ n•,pir.ltory ~trudllrt'' 111 the rup.
Because of th1..·tr 1..,1luttt ~k.ektou edunodcm1s arc
CLASSES 1 and 2 DIPLOPORITA and RHOMBIFERA (?< m
very abundant 111 the fos~i l relord .•md often their Dev.): 'Cysr01d~' - cxrt nct group~ w1th perfnr.1trd pi:
rcmams have greatly l ontnbuted to carbonate sedi- in the calyx.
ments. rh m Lnlluld,JJ(imt'~tOilC~. I..Olllposed largely CLASS 3. BLASTOIDEA (~ 11. l'crm.): Extl tJCt 'pehmtozou
of the stem fr.1gi11L'nts of sea hltt·'· arc very common wnh complex rc~pir.llo1y \lnlt tu re,.
in some roL k'>, not.1bly the Carbotuft·rous. ln such CLASS 4 EOCRINOIDEA (L. (' un .-~1!.): f'nmmvc rrhm
gem
rocb the porom plates h.!Ve often been unpregnatcd dcm1s wnh pores Jlong the 'utun:s. platE
SUBPHYLUM 5 HOMAlOZOA (Ord ) RJrc pecubar ot~ll· I
with dJagenc:nr t.1k ium l .trbonate but arc otherwise
1~ms, c:alotc plated but wtth llll pl,ltln of ~ymmct:rY. Th
unchangt·d. Echmodt•rms .ue. however. stenoha- have been till' 'ul~crt of rnurh t'Ontrowrsy and
lme, and their rcm.um are onlv tound in scd11ncnrs be .1 scp.1ntc dwrd.ttl' \llhphvlum on their 11\\ n:
of fttll) man ne on~n. CAlC'ICH ORDATA
Ftgvre 9.1
frOm Dvrho
Subphylum Echinozoo 263
Tht~ j, J u~etltl ,1nd W<lrkable d,I\Slht.."ation and is and LepidocystoidC.l) have had to be establi,hed
mamcd for t..onvc:mcnc t here \n11th ( 1!.J~4b). how- to acconunodate them. Though there are ~omc
ever, proposcs .m .tlrcnuttve one \vhich ha~ been Cambrian cch111odenm, tht: epoth of their ma....:i-
dtnn·d dadisttt·aUy. I 11 hi~ c'tlnMtion, the view of mum proliteranon (.1t da$' level) wa~ the
the eJrl~ taxonnmi,h, thJt tiH.~n~ ts J n,ttural dl\ tsion Ordo\'il ian, a~ cJn be ~cen lrom thetr time~ of ori-
bmvren fixt·d (pclm.HozoJn) and trcc-ltvmg gin in the above h\t. At genenL level, howe\·er.
dcmhcrozoan) tThinodcnm j, phylogcnerically cchinodem1s ~were mo~t abundam in the
correct.•md rh.u the t\vn .uc ,j,tt:r group~. In conse- Carboniferous. A whole range of rem.ll'k.lble fonns
quence he propmcd two subphyla. Pelmarozoa arose at that rime in J greJr bur;t of ,1daprive radia-
crinotds, t·rc. ) .md f:kuthcro:wa ('tarl1sh. sea tion, bur only mme of these were ~ucce,sful; the
~rchim Jnd ~ea cut umber.) M.my dctaih remain to od1cr~, to wh1ch the Cambnan ci.1S~es belong. pro-
lx worked our, however. Jnd here I me the ~an1e duced no new lmes of descent .111d bec:1me extinct.
dmtfiranon ,1\ 111 tht· \ctnnd Jnd tlurd editions of
!hi~ rn't. Some furrlwr commcnh .ue mJde in sec-
cion LJ.~. Lk\ldt·, the: n hmmkm1 !-rroups nored here
there art· several others. In p:~mc uhr there was a
marked prolifcr.nion nf ,hort-lived echinodenn
b~~cs in the I own PJI.tc07mr. Some of these ~ecm C lass Ec hinoidea
m combmc ch.tr.tdt'r. typte<tl of m.my groups and
<re hud to cb~,tt)•. They ,tre often known only from Morphology and life habits of three genera
1 >mglr locahry and ,1 ,m,tll number of specimens. Echinus
Bur rhm1gh rht·'c .Ht' ckarly l'chmodenm, their The common sea urchin, T:chi11rts cswlflltrts (Order
haractcr.. .lrc 'o different trnm tho'e of other Echmoida), whtch hvcs m ,h,tllmv waters around
ln0\\11 cchinmlcrm' tlw wp.u.tte da,se~ (e.g. the the North Acbnrir, 'how' the fundamentals of
«hinoll).m dJ"c' Hdinlpl.tt..mdcJ, Cyclocy,toidea echinodem1 'tructun: ,1, orgnni7cd m an anmul well
ambutacn.HP
mcidreporne goll notcht~~
nu
1th
buccal plate
cd
th-
11.
~~t:'~
Jl(;-
~~ln
ht:''C
lll.l}
the tubercle penstomP
(b)
fisln 9 I
Echmus escu/entus (Rec.), lest deprived of spines: (a) aboral (odopical) surface (b) adoral surfoce (v 0.8) (Redrawn
m Dumom in Treotose on Invertebrate Paleontology Port U.)
264 Echinoderms
,ld..tpt~d for .1 hc:'nthtl ~n.·c-hvmg mode of life (Fig. tome rhc: test IS nmted bacl-. uno a perfonted Aang
9 IJ,b). which IS the perignathic girdle (F1g. <).2)
~ome I 0 cm in diame-
reil ill liS h,t\ .1 globuiJr test Thjs gudle form.. a 'upport for rhe ma-;tteatoif
u:r wluch ts ~lightly flart~ned .tt the poles. In life it is apparatu' of the echutotd: the Aristotle' s lanttm
covered w1rh 'hort (1-2 cm) spmes; tf these are (Figs 9.2, 9.25) . Tlm lantern has tive mong iJ\\~
remo\'ed dw pl.lting \0"\Kture is \,sible. On the each w1th a smgle calcnte tooth. The whole :t: ,('m
uppc.r (a b o r al or adapical) ,urface there is a central bl) ~~ \USpended by ltgament~ and muscle~ JttJchcd
apic al disc: ,1 double.: nng nf plates surrounding a to the pcngnath1c gtrdJe lt operates a~ a kind
central hole or periproct, \\hKh contains the anus. five-jawed grab .md, rhou~h eachja"v has only hm=
The. .lpic.J.J J1sl 1s fonneJ of two type~ of plates: rhe iced play, the teeth can r,tsp .tway at org.uuc detnCI
lJrgcr genital plates . and the \maller ocular plates or algal m aterial on the \ea floor and pa~~ lt im,anb
whtc h ,tr~ usuall) outside the ring of genitals. Each eo the gm. Within the tc~t (Ftg. 9.2) most of the lOt
i' pc.·rfi.lr,mu by ,1 pore. The genital pores are the parts arc rcLucd eo the stnlt'nl rc~ already dcscnht-J.
outkt' of the gonads, .md the ocular pores are Inside the rest i~ a thm layer of protopl.mn, ami ,1n
part ut" the \'>.ltl r· v,t,c.ulJr system. One genital the gut is only a \i mple tu be nmning spirally round
plate (the madrcporitc) IS IJrgcr than the others. lt the inner wall from mourh to anus, the body ofth,
h..ts numerous uny perCmatinm which lead into the test is largely empty. llnwcver, at breeding Ontl'
water- v.m:ular ~ym.•m bdow. The anu~ resides in which is no mtnlly in the ~u mmcr, the gomd~ \Wr
the centre of .1 numb~.·r of small plates <lttached to a eno nno usly before releasing their products tl1rou~
tlcxtbk-. r,1rdy fm,diZt·d m~.·mhrJnc extendin g acro~s the genital pores. Since echu101ds often h\t
thL' pcnpron. clump~ or congn·gatc to ~pawn, rhe chances
1 he lc\t '' dtv1Jed utto ten radial segments cross-fcrtilintion of e~TS and sperm from male~
extenJmg from th~.: ap1caJ dtsc to the peristome female indlViduals arc f.1ir. Tht· niJJtcd echinoplu·
(q.\ .) whil'h sunound, the mouth on the lower teus larvae whil h grow tl-om tht· zygotl" 1\
(adoral) 'urt:tcc. The fl\ ~.: narrower segments are actively in the plankton and undergo man) tr.ln,for
the ambulacra (ambs) whtch connect with the manom before finally \Crtltng down .
m:ul.u pbtt''· whl'Ct',h the broader interambulac ra The coelom h.1, vanom tubular clement-
(intcramb~) termmatc agaimt the genua! pJates. Both these the haemal and perihaemal sy~tcmJ. \t:em
ambubaa and imcrambuiJ~:ra cons1't of double be m vol\'ed 111 nuterial tramfer. .md the aJO~
column'> of don~ateJ pJJtcs \\ lm:h meet along a organ seem' m he <1\\llCI,Itcd wnh the n•plir
central surure m J zt~zag pattcm. In the ambu- injury, but thc.·1r spenfic funct1om .1re undc:rr
lacrum dm 1s tht' p er-radial suture. The interam- The w,ncr-vJ\tul.tr W\tem of the n:hinodtrtll!
bul.lcral pbtc.·s .1re l.trgL' Jnd mbercular, without F1gvre
pertor.ttton,, but the ambubcral plates each have foolwi
three set~ of p.tm.J pmes neJr the outer edge of the sltl like
pl.1te. I hest• pore pairs .uc the ~ites where the tube
teet emerge through the rest from the internal part \\ htcl
of the w.1 ter ·v,lsl ul.tr ~ystem. rlll· ,11'
The .mtbul.tc. ra anJ lntcr.tmbulacra are widest at .m· th
the ambitus, wlmh is the edge of the specimen tc ~t j,
when seen from Jbovc or bdow. The peristome i~ a (the s
large adoral Jre,t, <.overeJ in hfe by a flexi ble plated I.m tc 11
membram•, wh1d1 contaim the mouth centrally. In \\ luch
fos~tl specunem. however, the membrane has nor- rung 1
mally gone. leaHng .1 large CJrc ular or pentagonal the!'t'
CJVIt)'. ftve p.urs of gill notches are fou nd where only tc
the mterJmbul.tcra abut the edge of the penstome, pengna\nle ~
U\ ~ in
and from these projel't feJthen bunches of gills \el\ th(
Figure 9 2 fntemol morphology of Echmus himpli~edl pmq
wh1ch prov1dc \UrtJl'e\ tor re~piratory exchange through on ombub:l'\lm (left side) ond interombuloavm V\h leadmt
addttional to those uf the tube feet. lnstde the pens- side). {llg. l)
Subphylum Echinozoa 265
lb) A,.,.cia (c) Mel/ita
-rosettt!
- -cotklgen - -
s
t
l.
e
d
lC
• •
e.
:U
~h
1n
of
nd
u-
im
)r-
Of
1 to
tial
of
Figure 9 3 Structure of echinotd tvbe feet: (o) Echinus tvbe foot (with sucking disc) and pore pair; (b) Arbocro muscular-walled tvbe
~with large sucking disc and pore potr with expanded attachment area for collagen Fibres (c) adopical respiratory tvbe foot and
llit·l ke pore pair: (d) sub-anal tvbe foot of Brissopsis with 'Rue brush' and associate(! pore. (All redrawn from Smith, 1978, 1980.)
11hkh 1\ .ll\o cod~)lllll, n:st•mblcs nothing else in At the b,1se of C:<lch tubl: foot is an inAat::tblc sac
tnc ammal ktngdmu. lt~ prinwy function 1s to opcr- (the ampulla), and the.: tube: foot leads outwards
1~ thl tube lt•t•t. In cchin01ds m only e>--it from the from thi~. dividing as it p.1s'e~ through the pore pair
.~11" vta thc madn:poritc:. From this ::1 calcified tube and reforming on the: other side:. This device pre-
thr ston e can al) tk~rcnd'\ to near the top of the vents the tube foot from being withdrawn nghr
I 1t.m. I kn: 1t tmm the c irc umora l ring, from inside the test when rctr,u.:tt·d ,m d. since one of the
~lurh tiw rad ial water vessels extend, one nm- funcnons of the tube: foot i\ rc,piration, it also sepa-
)11~ up the u:ntrc of t'.tch ambulacrum. Each of rate~ incommg o:-.-ygt·n-rith water from the outgo-
nng ese pa,~cs tin.tlly through an ocular pore, but it ing Auid deplc.:tcd in oxygen. The tube foot
1) ti.mm a tiny dmt:d tubt· (.1pparently hghr semi- pmsc:s\es longitudinal mu~de' and has a sucuon cup
gtrdle ~ 111 some: t•clunoids). from the radial water ves- at the end, rendering it prehen~ile. Wirlun the
there ,\rise:, Jl imc:rv,ll\, p.tlrt·d l.ueral tubes, each watcr-va\cul.tr, haemal and pt·nhaemal sy:.tems are
passing many amoebOid cdh (coclom ocytes) which per-
11(right
Uding to .1 rubt· loot .tnd ~mociated appararus
•. 9..'\). fom1 numerou' functions.
266 Echinoderms
l:cl111ws mm·c~ by ming ih ntbc.: tcet, cspeoally ball Jo•nr of the tubercle below it; tht• tuberck r R.cJ
tho~c on the lo\\ er part of tht.· body. It can extend cry,tallographically continuou' with the platl' (Fl!:-" to •
the d 1mt tube tcct for a nm~1dc.·r.1blc dt~tancc. 9.4a-c, 4.23). Round the tubercle 1~ .1 ring ofmu'- I ht
appro:..1matcl} half of thme in .my one .1mbubcrum cles. attached m the spine ba~e ~o that the ~pine can not
bcmg extended at a gwen time, the other hJif be moved in any direction. nun
retr.Ktcd. A rubc foot will exrcnd when water pres- Amongst the spines are small org.u1s of babnrt> \\ISl
~llre vvllhlll it increase' dul· to Clmtr.lction of the (spheridia), on the adoral part of the pcr-rad1al
I\ ·"
ampull.1, the rad1al watl'r ves'd itsdf or,, neighbour- suture. and ophicephalous pediccllariae, whJ(h ocul
ing tube foot. As \.\'Jtcr come\ 111to 1t the lon1,ritudi- .m: tiny ~pines with their he.tds modified .1~ pincer. dod
11.11 musdes rel.1:..; wht.·n thc\C nmtr.Kt water i~ (mmc \.VIth pmson glands); the~e dt'an the surface, 'J he
torccd back mto the ampulla wlm h t:orn:spondingly 'dtscourage predators and prewnt larv.lt. from ~ct irre1
rdaxes. At the time of nux11m1m cxtcm1on, the suc- thng (Fig. 9Ad). Normall}' pedJcdlan.te he r~.·cum )'Ill I
tiOn tups on the end of tlu: extended tube ftet bent on the surface, bur they can be ert'Cll.'d and \nU thl· f
adhere.: ltl an adJ.acent part of thc 'cJ lloor. When the ~nap 'hut on any extraneous obJet t. 1\Jt'\\- pedicdlar- W\tCJ
tube ft'et contract, the eduuoid Ulovc' along the ~ea tat.' .m: fonncJ when any are d1slodged m ddeJlle. dc~n
floor. 'upported by spme~. L:c/111111\ moves ,]owly Ed111ws i~ a regular echinoid: one 111 wluch the nhir
over the sea Aoor in this way. feeding voraciously pcripro( t opens in the centre of the .lpKaJ dbt
With lt~ JIWS and dt·fendcd .1g.umt predJtor~ by its (endocyclic). Such regular cchinouh ,m: com.moll Eel
.1nnament of ~pmes. E.Kh spine. lil...c the individual today and 111 the fossil record. and they hve e1ther Ftlu11
pl.nc~ ot et.hmoid~. is a smgh: t.ry,t.li of c.1lcite. The on the ~c.1 Aoor or, like Struii~Yit'c''/ltrows, 111 CJ\1tli.'S ,J).JIIO
~pmc ba~e tonns a ~o~..ket wlmh .lfllcul,lte~ "vith the m rocb wh1ch they may have excavated thetmdn-s. \\ Hhu
rtied
(a)
lift.·. r
(cl crtmui.JIO
platform
agJim
lOIJnC
ptr,tto
\\ Jtb \
pruble
prubk
the le•
feet an
f..• rent
In f:.
pmt'd 1
\\'11 h .1
areoiJ fC:IIlOV(
areol.o tc.'nt'd c.:

.111 don
wtch .1ppar111Us .•hsttll;
pn~tc:ric
dt,\tmtJ.
po1son sac b'fOOVe
orhC'r ,u
thl·,e ''
mto tou
,Jh<lve tl
tlm !eve
Figure 9 4 Structvre of echino1d spines and tubercles: (a) surface view of standard tvbercles and (b) vertical section through sorrt rlw wrf.
type of tvberde and spine base with muscles; (c) asymmetrical tubercle with crenulate platform and wide area of muscle attochme<11 lllltl'f pt
at lcrt; (d) opened ophicepholous pedicellaria with sensory hairs ond poison sac. [(a)-(c) Based on Smith, 1980.) ~11d wid
Regular cchinoids arl· norn1.11ly illu:.trared according (a) ambulacrum Ill
10 a convenrion.1l Mll'llt.HIOn (Fl~"> 9.la, 9.19a,b).
The madrcpontc h .lh\JY' 'hown at the right ante-
nor and \\lth Its dept'ndent mtcrambulacrum is
pelalotd •mbutacta
umbcred 2. I he numbt.>nng proceeds anticlock-
i1\C (as ~ccn from the .Jd.lpiC.ll pole) so that gemtal 5
Jlway\ pmtcrior Rnm.m numt'r.lls dcstgnate the
1rulal"\ Jnd arnbuhc ra. hkc-.n~e numbered anu-
loch\i\1~. but srartmg to th~: nght of the genitals.
1c 'ame sy,tcrn 1' m<:d 111 numbering the pl..m.·s of
1rregular echino1d,· thmt' \\'Hh a dommant bilatct"'al
munerry marked pJrttcul.lrly by the posiuon of
h~ pcnproct \\hie h I\ no longt'r w1thm the apical
)\Item (cxocydic) I :chinvwrdi11111 and Me/lira,
.bcnbed below, are two very d1mrml.1r irregular
rhinoids, with ditli:rent hfe l1.1 bits.
(c)
Echinocardium (Fig. 9.5) sub-anal---.....-

fasctole
;: /wwwrdi11m (Ortkr Sp.lt.lll~oid,l) ts .1bund.mt in
billow \\.Iter, but unhkc Fch11111.( ltvc' 111 a burrow
linbin the \l'UIIll\.'nt. Its morpholng} ~~highly mod-
ud in ilt rortLtnt l' "'<~rh dm hurrowmg mode of
~ir. Livm~ below th\.' \urf:.n: b"~ Vl'\ 11 protection
~llll\t prLdaton., bur it h,1, to m.lint.llll an adequate plastronal spmes
OMection wnh the 'url:ltl' tl1r t'lmd 'upply and res-
torv e d1an~\.', .l!ld it aho ha' to be able to cope
lllh \.lllltatlon. le ha~. in t:•ct. e\\cntially the same (el
roblem~ ofllfc J\ a subsurfal'c-Ji~,ing bivake. These
problems have bl.'l'll solvnl nuinl} b" re,hapmg of
me tCSt J!ld b) CXlrl'lllt' lllodificatron of the tube
o:tt Jnd sp1ne~ iu dlfle1 em \\ av' tor pcrfomung d.if-
'tr(lltJOb~.
In Edrillomrd111111 .1 bilatt:ral synum:try j, \Uperim-
rostd upon the r.\til.ll Wllll11etn. The test ts covered
th l lllJt of shm t ~pmc~. but when the\e are
moved rr 1s wen .1s he.trt-sh.1ped 111 plan. a flat-
nrd dllpse in protik. The aboral surf.'lcc possesses
nelongate apical disc from whtch the periproct is
hsent; rhe latter 1s loc<1ted on the nearly vertical
t'(ll[~rior wall of the re~t. A s1ngle ambulacrum (lll),
11m1lar to the otht·r-s, IS located in .1 deep anterior
!JO<W~ :md got:s stra1glu tow.lrds the penstome. The
·n~r ambulacr.1 are p.mt:d (I I + IV. I + V). Each of
;,e 1\ in two part' J'he .1bor.1l p.lrt\ are expanded Figure 9.5 Echinocordium cordotum (Rec.) (o) oborol view
lDlOfour recc\~ed lcaf-hkt· 'petals', which ten111nate (x 0.8); (b) odorol view; (c) loterol v1ew with oil sptne5 present;
(d) oblique loterol view showing current directions; (e) in life
~e the ,Jmblt\1\. Tht• .unbulKr~ contmue below
position in burrow (Aher Nichol,, 1959 )
level but ilfl' of morl' 1Hm11:ll tom1. Aush with
surface .md Jcs, pronoumed. In the petals the
me
en! ::ter pore of c.1ch pore pair i\ elongated, sht-like
1\tdcly \Cpar.m•d frl'lll the round inner pore.
268 Echinoderms
From these cmcrgt.> Aancned re~piratury nrbe fel.'t, .tppear only m the embryo and are \OOII lt1't throu fht'S(
k·,lf-like and rc.ct.lllg'UI.u. Elscwht'rl.', rncluding bcmg re~orbed. Other than the modrlu:d ,J,.rl'< o sl dm
withtn the .mtenor ;unbuhcrum. thl.' pori.' pairs arl.' the test, it is marnly a divisiOn of labour bctwt'tlltll ltyl'r
more normal, thnu~h they m.ly c.ury tube feet ~pe ~pmes and ntbe feet that enables Ecltillfll<mllllm 1 dt'll\1.'
ciJlr7cd for othC'r tu m nom. AdorJlly (F1g. 9.5b) the live f:lr below the \urt:lce while feedm~. rrspm ~Ill, 1
pcristomc •~ lnl~,uni t:u ton\ ani' :tnd hJ\ a project- .1111.l e~creting dTccuvcly. Cllal •
mg lip (or labrum) below. 1 her\.· 1\ no Lantern. ~pccific btoturbanon ~trucrure~ m till· 'cdtmcm 111 outl
AdmJJly the pbtcs 1re l.'nbrged. and tiom their large ol(c...n of complex toml, result from the fon ud long
pore pairs emcr~c: 'ncky tood-gatherin~ tube feet. 111ovement of hc.lrt urchins such as Fclzuz.'""drum llllllttl
Behind rhe mouth " ,l flattened .m:.r {the plastron) Thc'c arl' clearly rcLngnizable and resufy ro the tor· nun 1
f'lmm·d from th~· mndttied pmterior mtcrambu- · rncr presence of ~ea urchins moving withtn the~~ lt)f\~;11
bcmm and d~mdy covered with HJt, paddle-shaped iment even if the animals themselves ha\'c not bet llll"liCl
'pme' fo,\lhzed. thro"l
In hfe &/ulltlfcUdium lives in .1 burro\\ up to 18 cm and it
Jeep. with J ~mgk· tunnd connecting tt with the Mellito {Fig. 9.6o,b) po~ter
,urfJce (Fig. 9.Sl·) I ht~ timnel ts both created a.ud Mdlita qllilltJIIicspe[!i)/',1ftl (Order Clypt'J\tCr01d.1 u I ht
maintained by euonnomly long tube feet with star- AattenC'd sand doll.H common in littorJl :md wbht ( ;hrol<
,haped ends n•,t·mbltng flue bmshe~. These emerge toral ~ands in southeastcm North Amcric:l lt lr\~ llll fl',\1
from the ad.lptl rl pJrt of the antenor ambulacnUJ1, either on the ~urfate nr buried horizont.lll~ \Hth gather.
rheir ba~es protl't'tl•d by 1 pyramid ol sptnes wluch it. It '' very flat wtth it' Jmbulacra petalo1d .rb..mll J(l()df
help in building the lower p.m of the t'lnmel. A set Jnd wHh five pertoranom (lunules) 111 the te~t !\\ lltJ)' al
of \unilar mbl.' l~et. emerging from non-petaiOJd pam in the pam:d amhulacra. the othrr br:cg tl·ndcn
regwm of dw two po,tcnor ambul:at-r.l below the unparred. Ador.1lly thl pcristome i~ Cl'ntral, wt h 1h <kll ( 11
v~:rtical rear \VJII, burld J smglc \,1tllt.lry tube' to .1mrs very dose behind 1t and just in from nfthe pCII- The
n:cci\·c excreta. terior lunule. On the adoral surface nre five dichor~ 11011, ;]
Two rcgiom 11f tht• te'>t known as fascio]es gen- mous chamtels {the food grooves) which are lmed dlJn 111
erate current' Tlw hrgl·r t~l'\ctole \llrrounds the with tube feet. They run between the lunul~ cl., the
.mtcrior ambul.lt'rum a' ,l nbbon-hke ~trip; tbe converge on the mouth. The whok ccq 1\ rm'Cfftl Jn uurl
smalla (sub-anal) fN iok j, ,m cll1prital nbbon \\.tth ,t dense mat of fine 'pines, of which the ad..'lfll h} cih:
lotated bdow Lltl ,\ltU'-. In both tht• ~urt:Ke is coY- ones .1re used for walking. tin\\. ol
ered with sm<tll vertil.tl 'Pine~ (tht· clavulae), each lr usually lives honzontally, burrowrngjmt bclu11 J '<Ubst,
of which i!> cmen~c.l by nmtuner:~bk nha, .1~ in the the surface and covt:red by a thin l,l)'lT of ~1nJ p,t\sCd I
intexvc:ning epithdlllm; 1t i\ tht: coordmated beat of I\ \ ftolfi
these cilia thJt produce' the nrrrt•nts who~e direr- .\f SL
food grooves
non '' shown in Hg. 9.5d. A ~trong current goes ~llllii.U
down rhe antcnur .unbulacrum. md food particles 'durh i
are caught b) dtl.' \lit k) tube fi:et and p.1~sed to the htgb .Ill
mouth. Other lllm.·nt' bathe.· e.\l h of the p::rired rrnr p.tM
:nnbulacra, tacilitatmg rl.'\piratory ex~·hange, and the Some
current from the ~ub.m.rl tasciuk propd' W<lSte mat- mg m tl
ter mro the >amury tub..:. When the ''Ulrt.lry rube i' .md dlc\
filled up Etllitt<lr,mJ""" move' torw.trd\. The funnel- hdt wh
butldmg spine' ~re withdra\\ n. tltt• anterior spmes ~.md
'l
.lrc erected and r.IJ'l' away at tht· front w.11l of the t• hmmc
burrow. wlubt the p<~ddlc-likc ~pmc' atLlched to the hyt.onri
pl:mron move the l'clllltoid forw.1rJ~. A new funnel l'lll' pl:t
~~created wlule the uld funnd, burmw and sanitary lunules
ontogcn
wbe collap~e bl hrml 1t (a) Ib)
llltt·rluck
The whole org.umm ha' the \Jill~· ba,ic clement\ Figure 9.6 Mel/ilo quinquie5perforalo (Rec.); (a) oborol C'll Sand <
l\ Fchiuus, apart from the lanteo·m :tnd gtrdle which (b) adoral views (x 1.5.) l';ul~ T C'l
The~c cchinoids i'i.-l·d as they mov~ through the a consequence of thetr mode of ft·edmg. fhe dJtTer-
scd11nent by ~tevmg nut tine derntu~ from the sand t.>nuauon of adapical ·~1eving' ~pine~ and ,Hinr:~l loco-
layer above. Tht• fnod particles fall between the motory spine~, the AJt-bascd sharp-cdp;l·d test, the
dcmc spine canopy•.tnd arc canit>d to the test mJr- rigtdity given by the sutm.ll interlocking ot the
gin. There food 1\ ptc ked up by the tube feet of the pl.ttes, the adoral br.mched food-gTOO\ l·~ and
oral ~urfacc and c.tmed in mucu~ \mngs to the the Junular pcrforatiom through the test make up a
moUih, a.~smcd hy both mbc fret .tnd nny spines. 'pcctahzed and lughly etfccrive innovation m fimc-
L1)ng oral spmc' f(mn a protcctiH' mesh over the uonal design (Seil,lther. 1979, J9C)O; Ktcr, 19H2).
mouth; they \cn:cn out ~and but help m the mges- llowcver. dill seems to have onginatt·d m the
uon of org:uuc Jctntu~. Mrllita nomtally travels Eocenc at least four time~ over - a te~r.11ncnt to
forward but it can .\l'o rot:ttc and right itself if over- 'the kind of adaptlvc breakthrough' pmsihk from
n tumcd. A~ it moves tt makes a cluraneri~tic trJ.iJ the c:chinoid 'llauplan', but it would unlv work tf
through the thin vutecr of ~and \\ hidt overlies it. the many component elements an: \UCl'l:\\tVcl)
md tt :.ometimc' pa"c' 'and ttp\\ards through the llllorporated and fitll} coordinated.
post en or (at1Jl) lunule as tt prob'TC:"t' ton\ ards.
a The ambuLtcral lunuks. accordmg to Smith a11d Classification
t- tihiold (19H2) h,tvt· one primary funt tion. They The raxonomi~: divlSion of the Cl.l~s El hinoidca
cs mw:a~e the perimeter nf the test ,wJil:lblc for food into Regulari:.1 .md ln-cgu1aria, originally proposed
in ~athenng. thm Jt tlng ,,, 'hort nit\ for tr.msferring 111 1925, was used until the 1950~. Neverthebs.
tood trom the .tboral 'urtace to tht· mouth. They authontJes such Js Moncnsen (l92R-1952). who
'
\'U m~~ aho act as hydrodyn.unic '\tJbtlizer.. linutmg the contmucd to employ lt, dtd not nece\\.trtl~ bdtcve it
n~ tendency for \JJld d{lliJrs to be tl1pp\:d lWCr by sud- to be :t truly phylenc das~ification. but simph a t.on-
be Jm current!.. vcnicnt and provJston.ll om·. The d.tssth<..atton
s- lhe aual luuuk ~l·ems to serve ,, Jitlcrent func- l'rt•ued by Ourham and Mclvillc tn th~o: 'J'rccrtiSt' Oil
~(1- tion, and forms in JUVcmles by re,urpcion rather flllwr·cclmue Paletllltolo.t:Y· later amended (Durham,
lt:l\ thJn uurialiy by tlllknt.ttion of tht ~hell margin, as 11)66), and used in the tirst edition ofllm buok, w~
Uld do the .uuhul,tcral lunulcs. It~ roh: ntJY be to .tet JS h,tsed upon a \ndc vantty of stable ch,lr.tCters ddin-
red u: outlet for the l'Xl'e'' \\,tter dr.l\\11 mto the mouth tng m.lJOr groupmg~. nuny of whu:h ,uc \till valid.
~ral by rtlwy teedm~ currt nts, wtthuut di,rupring the The'e character<; mduded the overall ~tru ... tun.' and
'ow of the fcedttt~ l urrcnts them~dve,. lt may serve ngidiry of the test, the number of .unbulacr.U and
k1\\ \Uhstdiary funttton .l'; ,1 pas~Jgew.ty for excreta intcrambulacral columns, .md the morphology of
tnd. pmd through to the .1bornJ mrfJce .111d left behind the plate\, lantcm and gtrdle.
i! Jlel/ita progn·~'c' torwnrds. A critical and dcb,tt.tble i~~uc in cdunmd d.lssttl-
.\1. st•xie~pc1;,,,1f<l h.t, ,jx lunuks bur live~ m a Vel) cation i~ whether Jrre~"ltl.mty had amen otttt' only
S!tlllbr manner. Dr/11/r.wt•r. another ~.md dollar or whether the trregul.u edunmds .tre poJn,hvlettc.
Ytluch i~ non-lunulate IS a filter feeder. h lives at a Snuth ·~ (19R4b) das\ltlcmon, based partJ,tlly on new
;h mglc to th~ 'ubstrate. w1th mo't of the poste- dat.t on lantem and tooth stmcmrc, and exp<tnding
1nr part showmg .tbnvc the surface th~o: ideas of MJ.rkcl, favours the VJCW that trregular
Some specie~ of i\IC'tlita and other sand dollars liv- ~o:thinoiu~ arc monophyknc, and in comcquence he
m~ m the ~urf 7nm: ingest m::tpletJte trom the s::tnJ, r~.=vivc~ the old b'Toup lrrcgulari,t. Smtth\ \theme.
mJ they dt\tnbutl' rht~ wtthin the bod) as a 'wt>tght whidt is partially cl.ldmtc, ts adopted here. lt num
bdt' \ htch J;JVe\ \t lbtltty be appreciated, however, rhar cl;mttic.ltlons gener-
S1nd dollm. like other highly modified Irregular Jll) arc 111 a st.1tc of flux, and thts m.tv well llunge
hmnids. do not b'T'O\\ m the ,,une way as Ecllinu.\ ovt·r the next few ~l'Jr\.
r wnrinucd ~ccrtrJOn of pl:ttc' trom rhc aptcal disc.
11w plates an: ,111 t'om1cd m tht: carly ~tages of SUBCLASS 1. PERISCHOECHINOIDEA (Ord. Rn.): Rq,ru!Jr
cnJorychc cchinmd• wtth mtcr,unbubcr.t 111 rn.utv l ulurnn,,
1,togcny ,md b~:{·ome very ri~'1dly muted through
.uubul.tcr;t m 2-211 cnlumm, 111.1 compound pl.!tC>.
terlodung suture' .111d mtemal ptllar 'iupport!.. l'l·rign.uhtc g~rdle ~m1pk· ur 1b\ent .md !Jnrau \\ 1£h mnplc
rol onJ s.tnJ doll,m ongin.ltl'd from clypt:.\Stl roids in the grooved teeth. Include' JJI P.tbeozoic ct·hnllllth C:Xlcpt dac
a:!) Tt:rtl;Jf). ami their \UCCeS\ and rapid spread w;c, Cld.lrl11lh. Tlus a< J datltu1lr gn1up to cb<"f) : 11 meludes fiVl'
270 Echinoderm s
group~ ( I) lep1do, l'lltrid,, thl <.'.lrhl'\t echmo1J, such .1~ OrJe1 llulcct)'POid.t h.wc a f11llv dt'.illlll't p<'llJ1r<1•t
Aulrdrmrh, wnh cwCl , nlumn' ol .unhuhtr.U pl.ttl'' .111d 0\Jll\' but mhcrwJSI.' rcrn.1m quire ~1mple (l/,>/nt}/'" )
1mbmmng ambulan 11 plue'; (~) t•dwtoq,tJJJd \, wtth .lt SUPERORDER 2 MICROSTOMATA The lamc:m IH>fi
l.:;tSt tilut column' nf piJtl'' 111 l'JCh .unlml.lcrum lmt 111 the: adult. but tf prcsrnt 1\ whnlh uor 1'11.1!
(Echllrt•tystitt•s, C:rm•..ru•,Jt~rrtr~); (.') k·p1destlmh. V.lth very hJw ~· titlly dtS,Jllllll prnproct, .111d a 'mallpt·mttllliL'
niJnV pl.trc' m cJch .unhubcrum (1'1'""'"11"·,), J dtJractcr Tuhc:r.Je, on the adm.u \ttrfact Jrl llTJngt•d ti•r unlo
,hared b\ (4} prm~rotldlnd,, but 111 tlw bttt:r the .unbu4cra dJreCIHuulloc uuwuon
arc mud1 l'Xpanded on the JdnrJI ~~~rt~c:e (Pt••lrr,,tcl.ms}. (5} Nt.>o~narho~tom.H.J h.lVt' a muml..:d. pent.l!-'<1n~l
pabedliiHHds wnh J'llly~onll amhul...-r.ll piJtl'' w1tb snlMI dhpt1c.1l omline. Tht·y retJlll the 1Jnreru, Jt k.1 1ID
~ecnnd.try tubercle~ {rtrl.u ·dlll/115, Mc/,•llllrs). thr JUVenile~. Thry include fn111 Mdt't>, pJrlltulorl)
SUBCLASS 2 CIDAROIDEA (Dt>\ Rt'< .} Regubr enJm·ycli. (1) ORDER. CASSIDULO IDA (Ciypm.<, "Ju,/u•/11
cchmo1d \\ 11h onh m n tolumm of plltc.; 111 the ,unbuldcra Cu.<qrfu/u.,). whrch uc subg(,)bui.Jr \\.ith ph)lloJo
and m rlw mteramhulll r.1 of!drer ~··ncn. f.Jch of the inter- aud bourrde~. and nonn.1lly lme the l.uu,·m m ~
.unbul.u:nl pl.1tN .lrt oiiUlllt-nted wnh J lon~er \lngle .:cntr~l adult; (.2) OR.DI::.R CL YPl ASTEROIDA "I"'
rubcrclt 'upport1n~ 1 111.1\SJVl' "1Lid 'pine The upnght compmes flatten eel or va1dt... d n lunoid' 1111ludm
l.uunn h.!\ nn forJ111l'll 111.1gmuu, .111d the pcn!-'ll.tthJc gtrdk the \.IIIU dolJ.tf\ (thcv are the mo't rt'cent otJ .,
has apuph)'l"' onlv. Lx.unplt:~ u1dud.- Cid.m~ .•-tr,I~<U•lfld.m<, edurll>IJ,, fi.N ;~ppcanng m rhe Palt:llf<'lle, t..
,,/itlftclun • Cln•m.•ta, ,\1,1/u,r. R<'lu/.1. &:uttllomr).
SUBCLASS 3. EUECHINOIDEA \U rn.1~ -Rt'l .): Tlm group Atl'lmtomJIJ Ml' h,·,m-,hapt•d urchuu wh1th ha1r
mcludt•, .1U pust-P.t!.tt'OZOIC cc:h11Hlld~. 1~:gul.u u1 mcgultr, lmr .d) trace of thl' bntrm. Thcr1• .1re tbrcr ord~f\ (I)
with bltulunlllar Jmhubcra and mtt·r.unbu!Ja.l . The pcnt;- Ord<'l Di...a,tcroJdJ. wid1 .1 ~pur lpK 11 "
nJthJc );lt<llt Ius buth .nmclc;; md .tpnph\·sc' .tnd th< Lmtcm f),,,,,trr. C:.•llyrrtt')): (:!) Ordrr I l o!Jsrero1d~. 11 11b ~
mav ht· •n onduilv lmr m '0111l' ~umps Till\ 'uhdJ,, is '11 clon~.11ed Jpical ""' (H ,,f,IS/cr, l'<•ltrllllma) Ut<lliUII)
large tlw ll h.t• b.-uJthVIdt·d mm extr 1 division' wluth ~.;oml 'suh-'JUtangot d' lcJtun:-s; (J} Order ~pJt,lll!-'l'ldl
betwt:en subcLt~s Jml u rJcr . Thci'C: ,ttt• two infr aclasscs: wtth J rompacc .1pif.1l 'Y'lem and clt,lr.JCtCI., <'Xl·mp],.
INFRACLASS 1 ECHINOTHURIOIDEA· Dccp-\e.t cchinOJth fird hy EdtiiWI'otrdllt/11 and ,\/irt,l\11 1
\\ tth 1 flex.~blc lt .c .mJ J Ul11•Jlll kmd of P't•udo-com-
poundmg of dll unhubtral pi.He' (Ecltmt>rlwra.r)
The nme r.mgcs of all taxa arc g~v..:n m Ftg. 1 }.~
INFRACLASS 2 ACROECHINOIDEA: Echmoltb w1th .111
upnAhl lantern h.lvlllg .1 dt'l'P V-sh.1pcd for.unen 111.1~
In the followin~ discussion of morpholn~.
mmJ. l'Ompouml .unhul.ltrJ. Jlld pt·nstontiJI tube feet ti.mcuon .md habit lhl' rhrce prtmary cchmow n
Th1' mfnciJ.~' mdudn thr.:e cohorts subclasses, the Pensd10echm mdea (pnnnmt
COHORT 1. OIADEMATACEA R .. r;um mJnY pnmmve C1d::tr01d~:t and Euechmmdc a (adv.mct•d), .1rt 11ktn
fcuurc' such a' rc~11briry .md !(Ttltlved tt•eth m .1 nd1.:r \cpa.ra tcl y.
dd.tmld-hkc l.mtern. Slullu\1. -w.tter 'Pl'tH:\ mcludc
cor 1l n·cf mh tblt.lnh wnh pn!'on ~pint·, There are
thrt•t• orde~ . Ex tmp(c, m.ludc: (),,,,/..,,,,
Subclass Perischoechi noidea
COHORT 2 . ECHINACEA All h.1vt keelt'd ct•l'th, 'ohd Pcri~choechmoid.s (On.l.-Rcc.) arc thL pnnuo
'PIIIt'S .1nd sutun:d pl~tcs, ,1l l ot wh1c:h ,rre .ldVJu ccd fe<~ cch.inoid ~tock
wluch mdudes all extinct Pa1Jn1Wt•
cute~. They .trt• tl')tllbr wuh ~dl ~~~t~. ,·ompound plates echinoid~. Only 37 genera and Jbout 125 spcci,.., n1
.tnd .1 t'ompl.:" pcngnuhll ~1rdlc. TheTe .ut' rwo ordt:l'i PJbcozotc t:chinoids art: currently knm\ n. Yl'l \IrK
{I) <;TJRODOt-.:'1 A e.g. -Jctt•.• r/t'rllol. Hnmo~rl.ms. s,,,,. their prc.,crvatton may be good. phylo~cny aod
,,,,,,, -lrh,,,l,t. {'/,l'""''''""') .mJ (~J CA1v'IAR01) 0NTA evolution have been studted m dct.ul. c'PL'nJih 1n
(e.!! Fr/unus. '''''".~)·INt'llll<lltt.,), Jr,tin~u~hcd tlll tlw
\tnu: rurc of rht· l.mrcm
the m:Jrathon work~ of Jackson ( II.J 12) and Kc~
COHORT 3 IRREGULARIA: Attordm~ eo ~nuth the irreg- (1965).
ular cchmo1,h \H'rc monuphylcnc. ~nd lrnm 'uch Pabcozotc echjnOJLh, mcluding odaro1(k th 1
'mt,·nncdute gt·n, rJ ·" PI. <~rtlurrus and Pyl!•l.</ft t·volved from thetr l.ucr counterparts 111 \t'Vcral rcspetL\
to rhc grc;,r d1vt·r.uy of mm• UJder.o grouped in rJ1ree
supaordcr<.. All lncgul.m.1 (n thl'l' thJn rhow rh.n haw M.my of them have tlcxiblc lL'\t,, often (11 lu~
now Jo,t the IJnran) haH dlllllond-shJp tJ nr wcdgc-
~ize and w1th the pl.ttes not ng1dly united. w d
!Jkr tt'Cth. Frgure 9.7
SUPERORDER 1 EOGNATHOSTOMATA Dt,ttngu~>hcd they are ofit.'n found 111 a collapsed statt.' ln ~011\C' represents c
on the structure <1tthl' l.ultC:JII (fig. 9.2'i), h.•ve ~JJ\1- case~ thl' pl.1tes arc thtck ::md have bcvelkd t'llt: (Ciypeosten
ple un~pc:o.dllcd 'Plll<'' .tnd 1ubc tccl. In Order 'o that they can ~lidc over tllll another .11 tb
P~gJstcnmkJ the pen pro• 1 ~~ hyhok-,Jup<' d md mar!:,'111\
lulf\\.ly uut ,,f the lfnc.rl dr,, ( PirHnlulltl>, f1'1!•UI11), Either tlw ambulacr:~l or mtcrambulac ral roln
0 2 ®
"""""'~'"'
5
Ech~nacea
f.~!::O """'
I
Ne<>yOldU >.V"
OcWooda
Ooade
metace-t
Holecty·
poodd Cassodutooda I Clypeasterooda l="er SpeWlgOI<II S••odof'la C..marodon...
~I Ill Il
I
~
I I
tS.
CS
he l
'
I I
\11
i
u
h I
11~
nf
g.
VC
5
iz
\ ~-
r-
lJJ r,
r=:==
l. J,..
.->-
[.)-
~- ·-
(1)
I~ la
cm
an
1fi)'
L_,_ -- ~-~--------- -- ------------------
-- --- •
1da,
ph-
I! EUECHINOIDEA
\~
..l
1- .
rgy, r-
3
oan
\'C),
s
~
ken
1- PEAISCHOECHINOIOEA
18 I
tin: I
J ,,
r-
zoic ' '!.-- J
!S of ~ rI ,
in.ce t
and 0
1? ~ ~
ly in
Kier
iiffcr E
I
"
large
> that ligure 9 7 Time ranges and Smith's (1984b) dossi~cotion of echinoids, illustrating some representative genera. Each vertical bar
SOlllC yesents on order 1, Au/echinus; 2, Cidoris (Cidoroida); 3, Pygaster (Eognothostomoto); 4, Cotopygus (Cossiduloida); 5 Mellita
ICiypeosteroida), 6, Echinocordium (Spotangoida); 7, Echinus (Comorodonto); 8, Acroso/enio (StirodOnto)
edges
1tthe
UllUl~
272 Echinoderms
or both nuy t'Clmt't of many rolumn~ of pbtc~. (b)

but cumplllltHiin~ of the ambulacnl pbtes ts
unknown: th.ll ''· thert· " never more than one
pore p.ur pet column.
Th~.· pengn.lthic ~tdle j, nonnally absent or of
stmple constr\lltiun, .md th~.· l.mtcm ~~ A.mish and
ks~ clabor.1te than dlJt ofl.uer cchmmd~.
The tl'St '' li\U,tlly glubul.tr and invariably regular,
with the penprolt 111 the l·cntn: of the apical disc (c) (d)
(endocyclic). Onl) J t'l·\\ flattened genera Jrl'
known.
Some of eh~.· e.trher g~.·ncra have the radial water
ve\scl l'ndo,~.·d tntnrully by tnwardly projecting
fl.mges trom th1.• .unbul.lcr.tl plate,.
Ech.moid\ ~eem to h.1ve been a t:urly dt:ar-cut group

from the hegmnm~. but the pn:sence of a lantern Figure 9. 9 Portions of the ombulacra of perischoechmotds la)
of rather cchmotd-ltke Jppearance in Cbs~ Aptilechinus (Sil.!t internal view; (b) Ectenechinus (Ord). wi1h
Ophtoct~tltmd~.·.t. and the ~umlanty in jaw structure:
pores near the per-radial suture; (c) Pholidechinus (L Corb.) Wl
pores remote from the per-radial suture; (d) HyoHechinus {l figure
between echmotds .1nd ophmrotd~. make-. the di'- Carb.!t internal view, with partial enclosure of the radial W(lfe oboral
tinctton le" compkte tiJJn was tonnerly thought. vessel. (Redrawn from Kier, 1973, 197.d, not to scale.) reprod1
P.tl.u:ozotl l'Chtnotds seem to have been a rda-
hvcl} \lll.lll .md ummpnrt.mt componcnr of the le)
Pabeo1oic bwt.l .md \\'t.'tc probably cnvironmentalJy atl'd 111 a deep g~oow wnh the ,i_ngk unp;urcd po1t
re~rncrcd. 1lw fll'xtble kptdocentrids mdude the on the aboral Ol.lThrin ore.tch pLtc dost" to
\Try c;trlil·~c m.a.· t•rhlnotd\: the Upper Ordovi~.i:u1 Furth~.·m10re, the r.1dial \\.Jter Vt"'d '"a~ cndo'lcdb)
-1ulr.-hinus .md hrwcd1inm. Aulcdm11ts (Fig. YJ~) has a tububr CO\'L nng anstng lrom the lowrr ~urt:trc:
only rwo rolumm ot pl.1t....... m the ambulacn. the ambulacnl plate~. evtdently the ampulla L1) bd
Th" ·~ ch.tr.ll tl'OStlt' of the pnminve Famil) dus. The rebred Lctmc,/rinw (Ftg. 9.%) has p;ur
Lcpidocenrrid;lt' to whll"h 1t belongs. The :unbubcral pores, though one pore L\ ~m.iller than the oth~.:r, Jll
place-. :m: curiou' 111 th.1r th1.· per-rad1al suture is smt- m later kptdocentnd ~enera, e.g. the )iluriau Apllk
,Jmrr" (fig. lJ. 9.t). the pore p.lm are tdentical thou.
mU closl.' to the per-r.td1al suture.
Th1.· .lptcal ~ystem of 1.11/c,/llnw anJ other lcptJo-
madreponte v ccntnd\ •~ pecultar 111 th.lt there ts onlv oue gc111u
pl.ttc with :1 ~rngk pore, thou~h there .trc five Ol\l- f'tgure S
lars. Jaw~ ;md other parts of the bncem have b~l'll ~hawing
found, hut the l.mtern 1~ much k~s complt:x thJn U1 lion ode
~hown ir
the Cttcc hin oid~. In these early echinoid~ the uucr
amhulacra art• ~mooth or have only small tuberdc
but .'-lpriferllinrts h:~~ large spines on the ambuLm rdatcd
(K ter. I 97J). b1
I.ICT.l
The kptdon·ntnds l.med until the Carbomfcrou llw
but g.we me ro .Hlntlwr fJmil}, the .Echtno- l':ich v
cyo;ttndte, 111 whtch th1. .tmbubcra h.we more th t1cprc'~
two columns of pl.ltl'~ and pnmary tubercle~ on th Ill list ul
11 Ill
intcr.1mbul.tnal pl.n~'· Thl· number of inrerarnbt; et." hi not
larral c olumm nngt'\ from JUSt one i.n CMI'CIIctlu In tl
Figure 9.8 Aulechinus (Ord ), lateral view ("2). (Redrawn fr-om
MacBnde and Spencer in Treatise on lnvertebrale Paleontofogy to 32 111 ~lync1siiCIJt·s. In the Leptde,thtdae rhcrc ~ \lclouc,
Part U) agam m.my colunu1s of .unhul.lrr;ll plate\, aud m thl- Ethmo
Figure 9. 12 Palaechinus memami (l. Corb.; Poloechinoido).

preserved fragment of test in lateral view (x 2 approx.). (Redrawn
from Kier, 1965.)
'~J~re 9.10 Proterocidoris belli, a Carboniferous echinoid, m.tlly thick h~:xa~onal plate\ arranged in more or le"
cilc:J..ol view ( 0 5 approx.) see also Fig. 9. 11. (Photograph
aproduced by courtesy of PM Kier.) \l rnc.tl columns (Fig. 9.1.2).
V;mous evolutiOnar} trend~ havt bt't"U uutl'd
within the ec hmotds of the Palacozotl ~ome, Hit h
,,, the Aattt·mng of the te•\l and ador.tl t: xp.lnSton ut
thl· .lmbulacr.t, .trc found m Order [chmocystOJ d.l
.tlom·. O ther trends are found in more tdv.mn·d
P.ll.teozoic ech inoids. sut h .1~ a gem'l"<ll increase 111
tht· ~ize and number of ambulacr.t; an mere l\e 111
tht wrnplcxil:) of the Lmtl.'m; the devdopmen r of
n·~ulanry m the mterambul,ll ral pl.Hc'; .md rhe lm\
of the cn do~u re of the radial water vessel Mn~t
c b,mge~ were allometnt, but some .tmtogenetll
change IS ~ct·n, 1.e m tht mcreased complrxiry of
tht· lantern and m the deYdopmcn t of tubt·rcle\ .tnd
spme~ for the tirst tunc.
~ 9 11 Prolerocidaris belli (L Carb.; Echin~toida) (a) Subclass Cidoroidea

fOtitog expansion of ambulacra adorally; (b) and then controc- fh~.: odaroid' ,lrt• the cml) echinotd group m
'-ol odop1colly h 0.5 approx.) (Drawings of lop of spec1men
lhown 1n Fig 9 10, from Kier, 1965.) ~urv1vc the P.ll.teozmc .md they stJII per~ist today
·r hey f01med the rootstock o~· all pmt-Paheoz oic
ethumids \\ hli'r thcm,dve~ 'hO\\ 111~ rdanv~h
~btedPrott·mcidarid.lt' (F1gs 9 . 10. 'J. II) th~ .unbu- lmlt· unportalll change. r hc modem cidaroitk
n bccatm 1-rrearly cxp.mded on the .tdor.tl ,urf.1ce. which hJve thanged very htdc at lc.1sr smce the
They W I ISI~t of gre.1t numht·r~ of ~mall pi.Hc\, C'rct.Keous, have ofi:cn lwen rcgardt·d as 'liv1ng
d t wtth ' single pnrt' pair 111 .1 broad. round tcmll~' All udaroub have rdatively n liTO\'. .md
:presston. Almost ccrt.unly the'e were rhe 'itc\ of frt·quenrlv smuous ambulat ra composed of small
l'tular sut kercd pml1.1 \\ luch could anchor the pl.trl'S, each wnh a smgk pore pair (ftg\ 'J. U, 9.1-1}
hm01d securely to rh~ ...e.l floor. The interambulac ral p i .J ll'~ arc vety l.:trge, w tth a
In the Pai.H.:dunid.ll' (e.g. \/cl((clyct, Pttfllt'lhinus. ~inglcl.trge tubercle in tht· ccntr<: of e.tch plate. tu
' Ollfchiuus), prob.tbly deriwd from the whKh a mon~ \ptne is attached ( Fi~ IJ. I4,l,b, IJ . l5).
c E:lunocystmd.le, thcrt• Wt'rt• man} t olumns oi nor- Tht' mamelon, or centr.JI bms, 1~ \Urrotmded by
274 Echinoderms
ocu ar plate
adradrill suture
.nrerradral suture -
rnlerambulacrum
Figure
Figure 9.13 Cidaroid morphology, odoprcol surface, with ter- ombul
minology (x I). The onus rs located withrn the periproctol mem
brone at the odoprcol pole. (Redrawn from Fell in Treatise on
Invertebrate Poleontology, Port U )
pl!Ll'~
(hg.
Figure 9 I 5 Tylocrdaris Rorigemma (Cret.), showrng d-~ Mtnc
shaped stabilizing spmes, some of which ore still rn posrhon
only
lHIIlJ
(el muscle
attachments .rnd 11
~-~"
l'lhin
1•1 Jtlll'SI
In
bl~·n
,, )ll)ll
Th,
(bl aurrcle the T r
~~" t) pc.
Me:,m
Jbund
r&doal water \Oesset 111 the
dll' Cr
Figure 9. I 4 Ambulocro and interombulocro of Archoeocidoris Figure 9. 16 (o) Perignothic girdle of cidoroid with opoph)'llll t tJ
(l Carb.): (a) reconstructed and (b) with spine preserved. (Based only showing muscle attachments (viewed from within the peris
on Jockson, 1912.) tome); (b) perignothic girdle of Recent Parocentrofus (Echincido)
dcvdo
with apophyses and ouricles (Simpli~ed from Cuenolrn Treol1~ .llld p
on Invertebrate Poleontology, Port U ) si1Jlll)\J
.1 wid.: smooth areola .uound whtrh "a nng ofuny till he;
scrobicular tubercles. Our~tdc th.:.,e a series of Ill rou
~mall 'ecundan tuhcrclc' is trrcgularl) dtspcrscd. The old~t knov. n ndaro1ds, other tl11n pm~1hl) lt\U.l} it
The apic.1J 'Y'tem t\ usually Ltrgl' and has five ocu- ~0111c Silurian iorms rc:prc~o:ntcd only by ~pm~
lar and five genttal plates, but in most C1d:1roids these whilh have been referred to thi~ group, belong to Both
.1re not rigid!) unlCcd to the te~t and nom1Jlly drop Family Archacocid.lrid.Jc (U. Dc\·.-Pcrm.), \\h!th lt seen
out on tossthzaunn. The lantern " rdatwdy simple; like mher Pahcuzo~e cchinoids h.wc Aex1blc- tNI rduno
the p.:ri~'Tlathir ~:irdlc h.1s apophyses only (Fi~. ami multiscrial intcrambui.Kr:t . .'\'ort.lllcdwlll_, (D..-, noids •
tJ.l6), 1.e. flJn~~' rdkrtcd from the tnrc:r.1mbulacra. .md L.prdt,cidari.~ (Carb.) h.wc six ro cr~;hr columm of gcncrat
Figure 9 18 Botlrrioc1doris (Ord.), reconstructed, showing

r<gure 9.17 Miocidoris (Perm ), collapsed so as to conceal the spines and tube feet. (Redrawn from Durham, 1966.)
o:nbulocra (l( 1.75). (Based on Kier, 1965.)
plu~~: rhe common Carboniferom / I trllllt'<,ridaris Botlm(uidans (Fig. 'J. H3), \-\-hiCh has been one of the
Fig 9.14a) ha~ tour However. ctd.uotd\ of Family mosr controversial of all fos<>il echinoderrns, comes
\lioCidaridae lincludm~ Miocrtiaris (Ftg. 9 17), the from the Ordovictan of Estonia, but recently speci-
. nh known genm to survive the P~:nm.m C\."tinc- men~ of a related genus have been found in Scmland
!lt111l have only two Lolumm o( plates 111 Jmbulacra (P;ml, l<J67) as weU as ,1 ~inglc tmc BMhrit>cidaris.
nd tmerambulacra, ~cttling rhe pattern for .tll later lr h.t~ tive single-columned interambul.tcra :tnd
hmotds for "'-' llll h ;\fHicidaris .tlotw wt~ the tive double-colunUlcd ,1mbulacra which do nor
·csror. extend as far as the pemtome, each amhulacral plate
In Palaeoz01c cid.trOtd~. adapttvc rn.·nd' have h.wmg two large spm<.>~ .md a single pore p.m. Tht·
bffll documented wlm h Jll relate w unproved pemrome has a prinuttve lamcm but no ~'lrdlc. and
~·omotton on a lurdct \ubstrJtc. 1~ surrmmded by a ~mgle or double nng nf pbte\
!'here were a few snuU-stzed cid.:trmJ genera in wtth pore pairs. On one mtcrprctation there an: no
rhc Tnamc, including the first cidaroid' o( modL·rn gcnitnl pl.ltes, md the madr<.>porite is on one of the
~pf, but the ctdJrotds rc.tched thur ,ttmc in the ocular~; altemarivcly, the .tptcJl plates rradirinnally
\lc,ozmc. dcclinm~ l.tter, though they art still regarded as ocular pl::ttes could actuall}- be genitah
lbund.mr todav m the I ndo-W c~t P.tcitic province. (Durham, 1966).
1 the Nonh Atlantic .md - as an endemic family. BMimllfidans has been v:mously regarded 111 the pa'it
nr Ctcnondand;te 111 the Anrarctll. . .1~ nn echmotd, a diploponte cystoid. or a holothumid.
!5 Cid.trmds arc nott·worthy tor tl11.· remarkable Its lantern plates are small a.nd poorly developed, ;llld
;-
lrvdopmem of rhcir ~pmes. Fusitorm, dub-\haped there is some evidence that 1t nught be morl' dmcly
l)
;e nd peculiarly sh.tpt·d 'pmcs are found in the related to holothuroid~ tlun to echinoids. lt~ ta.xo-
d llow-water gener.1 Mmt of these .trt• 'luggio;h and nomtc position is tar from dear, but it is nor bcltcved
~ .. heavy spines seem tO be used tor ~cab1hztng them that any later cchinoids were d1rectly de'\cendcd !Tom
rough water. !:>lender elongate sp!llC\ arc more it. Eotlwria, another Ordov1c1an oddity, was origmally
ly ttJitn the deeper-\\ .llcr or mud-living ~peocs. descnbed as a plated holothunan (sea cucumbt:r), but
c~ other than the presence of m.1ny ttny opemngs in each
to Bothriocidoris ambul:tcral plate ir ~<.>ems to have echinoid features,
:h 1ccms .1ppropriate here to mcntton .1 group of
!1 albeit deviating from the nom1. 11m again wa\ an
;t~ hmodenn generJ \\ htch may or may nor be cchi- earl>' off.\hoot of the greJt Ordovician radiation of
/.) tds and whiCh h.we been a singul.tr puzzle for cchinodcnm :md do~ not seem to have produu:d anv
of m1crarions of echmologtSrs. The noronous known descendant<>
276 l:chinoderms
Subclass Euechinoidea and the morphological Tube feet a nd pore pairs

characters of euechinoids In regular and 1rreguhr eclunotds alike, tubt I
TIK cul·dunoJd' (1 n.l~ Rl·t .) mclmk .Ill Me-.uzmc oth·n highly muddied and pl'rform 'anou• fun
ro Rt·• cnt Cl hllllltth Mht•r th.m od.troid~. I hey .111
luvc ttVl' hh "lumtur .1mhubcra and ti\'l· ba:olum-
tmm. The \tJnd.ud' ulullll> tubt:• foor {Fig
h.!\ J rdanvdy thm muscul.u lay~r. and 'lllllC' ~ •v t
nar intcr,ltnhlll.taa: thq nuy he rq.,rui.Jr m trrq.,rular:
the Lmtcrn nuy ht• pre\t'nt but in irregular ltmm LJil
nmd tube feet h:wc virtually nu muscle. lu th• 1\-c
ll'l'l of <>uch cc hinoids a~ live tn lugh-em:r10 ~~~ 1
4\
bt· \CconJanlv lmt. Till ttd.lrllJds Jnd •·uechmotds rtliUncnt,, e.g. ·l rii<!CIIt, the tubt• feet arc h1!!hl) tllU
~re now bd•t·ved to h.l\l d1\.~rged pnor to tlw tul.tr Jnd h,l\'l' .1 wcll-dcvdopl·d suckin~ J fi
.lppeJrJnce of ,\fit,,l.f,m, \nmh .md I h)llm~';orth. .tlinb'lng ro ruck ~urfaccs. I ubl' feet modtfic l
I 'J'W). 1 hough the f(l,,!l rnord 1s scant, at le 1st two rc:~pirJtion and tor funnel h111ldtng (Fig. lJ.3t,d) hl\
cchinoid lint'.lgl'\ tnu't h.lVl' pas~ed .Kross the ~pct:.ifJC modific.1tions and, .l~ o;;mith ( 197H, l'lkO
Permo-1 ri.t,,il' bound.JC)'. Otlt' gtving rl\C: to the hJ~ ~hown. thc n.nurc of tht:. tube foot can rc.1~ltl~ 11<:
moden1 ud.tnmh, tlw tHht•r to thl· cuechmo1d' 1ntcrred from the strucrurc of thl' pore patr bv rtfa
"hich beg.ttl w dt\'t'NI\ Ill rh~ bter 1 n.1'>SIC and \
l'llCl' to modern examples. f'h1o; In tllnl ~hcds ltgb
c.1rly Jur.IS\IL, under~mng 1 S'fl·.lt .1d.1pt1vr burst and upon the life habit~ even of c:xtmct cchm01ds.
cvolvmg Into many familit·'>. At this ttme there ·were
,Ill matull:r of" lltrlllHitl.ll lllllOVJrlOm (K1er. 197~). Apical disc (Fig. 9. 79)
\\hieh .lllowc:J l't:olog~e.Jl dtt1C.•rt·n~i.ttwn dcmed to In regular eucdm1oids tht apil'.tl di.;c i\ l'ntlrt:• and
the: comct\":tli\'C: pl'n,tlwc:..:hmo•d\ Jnd ndarotd, LOnt.uns the penproct. Tht latter bears tht': amn If)
J he elhmold, ol tlw P.tl.ln>znk Wl'l't: .111 rl'gular and located centrally Within a ~t·nc' of small plate~" luch
•:ndOl·) dil·. I hou)!:h tlw rt•gular cndnrydic sy~tem cover the periprm:tal m~:mbranc. The dm nuy b
Ius becn n.•t,ltnl'd by lll.ltl} llfthl' Mt·~ozoit: .md later cxsert. with the circlet orocular places out,ldl th
rn:a. numt•rmts other grtHIP' quttl' tntkpcndentl} .1p1cal d1sc, ur insert, with m·ular and gcmt.1l pllt\1
bt•camc tnegular, and .tll of the'e .td.lptt·d for :\ .1ltemating 111 a \lll~le ring around the pcnproct. llLil
\\ holh or parn1lh mt uuul exi .. tcnn·. There Jre, t'Vl'll witlun the sJme spcnl'' there may be m.ll\'ld
however. tiltlctcnt ·~r.Jdl',· t)i irrq~ul.lrll) in the u:lls w1th a tl.1lly m~ert, exst:rt or mtcmwdwe cnndt F1g ure 9
cchin01d~. ::-,ome art• t.ltht•r ~impk. wtth the tton. Genera stll.h as Hwo~nh·11ia (Fig. CJ. I'Jp) ~nd 1974 .)
penprnt: t \till 111 the ad.tph".il rqpon, though out~idc Pdt.tstes h.wc ~l'illly expamlcu .lpJCal di"' l'o~cnn
thl ap•c tl dm·, wherea, m others 1t h.t, llllt.'TJted to up to a thtrd of the total surf.Ke area of the cchinoill. py~.MC
tlw lfl\\ er ,urf1ce. The l.'XI refill'' uf fundllli1JI d.tffc:r- l'ht·~e haw .111 extra (suranal) plJtL h) the 111 3p1t:.
t'ntiatH•n. hm\'t:\'er. v.crl' not n:adlt•d until thc pcnproct, and the ocular and gcmtal pbtc' .trc ott n pmcly
Cret.Kl'll\1~ w1rh tbt nn~111 l'f Order Sp.ttangmd.l. highly onwnenLcd. pat pro
Thc early Mcsozolt .ld.tptlve radi.ttinn nf cucchi- In irregular echmoids thc pt'riproct nuy hwc by dor
nnid' g.1vt:· n'c to rt'pre~ent.ltlvt:.:s of .111 lt:'llr 'upn- wholly or paruaUv escaped from the apKJI dtsc ":lnd (.m eq
order-.. In \IH1lt' of d1e\1.' thl• .Jrchetyp.tl pl.m tended be located out,tdc 1t. One <.:an 'cl' this actu tlly hJp- \udr<ll,
to remain r.lthn coml'n'.lllvt .tftl'r Its l0und.ttion in pt'mng in the development of modem irrq,'lliJr stl- tlw g~..·n
tlll l'.trly Mt·w701C ..md Ihue \\..IS Vt't) lit de func- urchm larvae. The periprott in these t'> miti.tll p.lftiaU:
tional cvolut ion rhert·.tikr. In others cKh stage. .1picJl , but .1ti:er ,, short period of growth 1l movCj ~hapl·d
dwugh 111 tt,clflughly tunt t1onJJ. wa~ l.tp.Jblc of yet externally. dt~rupttng the cmbrvonic platt:.'' a' lllhla 'J. 1<)~.,·. (
tl.tnher fum t1onal modtlit·.mon. Tlw 1daptl\·c ditfer- o,(); these tht·n re-fom1. though 111 an J\ynun~tn fn tl-
l'ntiJtlnn of tht· 1norc extrl'lllO.: torm~ of burro\\ ing fl,lnon. and th~ genital V 1~ u~ually mi"mg T gtnital
cchinmd, ·" n'prescmcd b) I fhintlmrdium. ''a far cry trregular pattern~ are. of cour~c, retJllll'd m th found.
from thl· lllllll~holoh"Y nf lt'> rt·mcHt'. regular, adu lt. Variou~ kmds of aptl'<ll dtscs in towl .• nJ Otht
P.tl.teu.:olt ,tnu:,tor. Y l'l on re the on gm of irrt:gu- rcl·cnt cchinotd~ arc here illmtrated in J morpho!(l~ 1 c:Jucti•
l.mty l1.1d provided thl' lll1tl.tl \tunulu\, there were lt'al \cric' ~howmg .1ll ~nge' 111 rhe migration uf tht l00Ul1l
rhcu pknt~ of lit:\\ ntorphologJc.tl duracters for pcriproct. though this ~~ nor intended to dj,piJ} tlu: llllkcry
evohmon.\1) pnHT~\t:\ to work on. .ICtu.tJ COUr\l' Of cvoJution. merely grade~ of OT~<inl· thl·rc ar
7,ttion. T h t• Pygasteroid;~ ~how som~ H1lerl·~ung tinn g<
evolutionary trend~ in tJus resp~ct. rhc Larhtit ~i nglc: ];
(b) Echlnus
'" (k) Orblgny•n• (m) Nucl.olltH (n) N cfunicuf•,is

efongetus
'"
4
wtth conventiOI'Iat notatton ~

f
~
:·.
tcl Plulechinu• (d) Anorthopygu• (e) SpeQngus
if]!
.
. ·. . ~
.·
·.· . ~'
()) Br~•sop•ls
(tnbasal)
lt
!-
1- frgure 9.19 Apical dtscs (not to scale): periproct, where present, shown in block (Redrawn mainly from Howlun~. 1912, Kier,
td 1974)
1!!
d. PY~.bteroiJ, Plolt'cllillw IJ.111'kim1 from the Lias, ha\ A monob:hal 'Y'tem h,l\ been .1dopted h) .111
"lC :m aptc.tl dl\t: hkc th.tt of J regular echinoid and is clypc.HtcroiJ,, Mmt ,p,tt.m~otds .lrt• tctraba,.ll. 'iOllle
:n purely endocydit. In \lit.:hd) l.ttt.'r pygastcroid~ the tribasal .md .1 IC\\ monob.l\.11: 1t \eem\ th.lt these
~nprott " cli~pla~:ed oubidc, thou!;h still enclosed mo .trc umkrgom~ .1 gcncr.1l rl·duttton to a
ve by dongatn1 plate,, .utd tht.> gcnit<li V " still present monoba~.1l \y,tt m .
ud (m eqLHVJ.lcm Londnnm •~ wen in some early
r· ,\udrltlltt'S spenc,). ln mme 'ad-..mteJ' pygasteroid~.
:a- the gentt.tl V ts ,tb~c.:nt, ,1nd the pc.:riproct ltes at least
lly parn.tU~ out~tde the apttal Jist, lomnng a kcyholt:-
.res shapcd deprc~s10n un t il t• ad,•pil ,u ~urfitcc (Fig .
::>es 'I.IYc. 1l.20).
cal In the majonty ot' itrq,:-ttl,tr nlunmds only four
·he gcmtal plate~ rcmam, though occ.!'ionally a fifth 1s
the found.
md Orhtr th.ln the bt e.thntt of the pcriproct, a
::>g- rcducnon m the uumber 1.1t" genital pbtD has been
the conm10n m \-anuus tm:!o{ul.n nhinoid lineages. Tn
the hol~rtypmds ( l · t~. 9.21) .lllJ c.t~'iduloids (Fig. 9.24)
1111- th~rc are dctincd tr~nJs Irum ,t '"'tcm "tth ti1ur dJ,-
:ing onlt gcnnal pi.Hcs (lctrabasal) to a \}'stem with a Figure 9.20 Pygoster (Jur ); (o) odopicol ond (b) odorol sur·
liest llllgk large plate:: \\Hh lour purl'' (monobasal). faces (x 0.75).
278 Echinoderms
have smglc pore parrs, but arc very muB and hem rut"!
numcrou~. Alternanve ly, compound tng of plat1.'S u
'P·•t.
common in m~ny euechmoid s. Compound pbt~ st.1gl
have two, three or more demiplate s wnlun thr npc
confine<> of a single ambulacral plate L1ch deuupb
has it\ own pore pajr. Such morpholog y 1\ UIUlt PH
pronounce d in genera like HetacJWilT! Jlllf (hg Cert
9.22d) m which the large central ntbercle 1~ W· loid'
ver.;ed by numerous denuplates . Componnd1ug Jlllbt
results from the subjugatio n and incorporation of Thcs
embryonic plate~ into ,1 ~maller number of 'm. [Ct l.!cru
plate•; .md i~ important 111 .1Uowing J larger number plO\(
of tube feet per umt area. lt IS also commonly .w.o buLK
ciarcd with the developm ent of a large ccntrJl phvllo
tubercle. and another advantage of the system must
(C) be that a larger tubercle could be 'upponcd b l•l
larger plates fonned by the fusion of smaller nnc!.
Figure 9.21 Holectypus (Jur )· (a) adap1cal; (b) adoral $ur-
faces; (c) lateral view(> 1 approx.). Compoun ding .11\o separates the n>w' of tuhc tce1
withm one amhulacru m. ~o that the tc'\t bc.m It'll
widely spaced columns ot tube fel·t r.nhcr rh.111 lilt
(cl Conulu• closely spaced p.lirs of columm as in the cid,trollk
PETAtOID AND SUB PETALOID AMBUIACRA

Irregular ccbinnids such .1s Edmmmrd111111 hJYC
.1mbulacra fom1ing 'pet.1h•. J.c. w1th the adJptd
part~ expanded to fonn J Aower-likc rosette. Wnlun
the pct.lh the pore pairs arc alway<> dongated and 1 r
individual pore~ widely ~ep::~rated, one bemg ~Ill·
like; they arc al\\'3Y' assonated w11h AaneucJ fl •PI-
(d l Heteroc.ntro tus 1•1 Clypt~nt•r ratory tube feel. Some of the earher and 'pnmttt\t
irregular eclunoid, fc.g. f-lolectypm (Fig. l) 21, lbl
Pygastcr (Fig. 9 .20) I ha vc no petah but they do, lllCt·
dentally, have gilb. The next grade m orgattiiJilOil
ts a shghtly expanded o;ubpetalmd condition. JnJ
more advanced brrade still IS the 'Y~tcm of \nm
dypeasterO ids and caso;idulmds in whtch the pcuh
are large but flush with the ,urface. In the e.trl) tUt·
tcncd echin01ds, so many tube feet would havt bl·es
Figure 9.22 Ambulacral plating showing simple (cidaroid) and
located on the lower surface, and used for food·
compound plates (not to scale) (Redrawn From Durham el al. 1n gathering, that re~piratory l'xchJnge nught l1~
Trea,se on Invertebrate Paleantology. Part U.) been ttnp:u.red had tt not been for Lhe development
of pet:ll~ on the upper surftce. These pctalmd .lntbu-
lacra were then prc-adaptt: d for resptratol) m~ 111
Ambulocro deepcr-bu rrowmg ech.inmd~. Some holasteroad uld
COMPOUND PlATES !FIG. 9.22) spatangotd s have depn:,~cd petal\, espectJll) the
In C\Jechutotd ambulacr:1 there .11·e neveT more than geologic:~lly later ones (Edllllt!Cardiwll being .l pnme
Figure 9.2
two columns of plates, but tht•re ofien are a very example), and the decpctttng of the petal' s~ $ucb nitidvs (Re
large number of pore pat!"'\ .md hence tube feet. Tlm that they fom1 an cffectlw channellin g >trucntrc for !he spine r
has been ach1cved m different ways by variou~ the rcsptratory currencs .ts well as providing ,1 b~11 ( 100 OF
brroups of t:chinoids Thu~ in th1.• 'idaroids the place-. England, a
for a protective Lover of tl.lt-lying spines. fhl· Jjf. (SEM phon
lerenti.ued anterior .unbuLt<:mm of hobsteroids and

\patangmds aim ,trow through rd.mvdy gradual
1tagc' from an ont-,rin,tlly Amh and und1fferennated
type.
PHYllODES AND BOURRELETS (FIG. 9 2A.A-c)

Certam 1rregubr echinoiJ~. e'pecJJJly the cas~Jdu
loiJ,, and a lt'\\ rq,\'ul.tr gena;\ have ~pcctalizcd
ambulatr.tl srruuures 111 the ncinity of the mouth.
Thc,e an: the phyllodcs: are.ts in "hich the ambula)
bcnnn l'Xpands uno .t k·,tt:lik.t· sh.tpc clo~c to the
pcn~tome. The phyllmk' .1re separated by mtcram- bourrelet
bulacr:tl swelhnh~ knm\ 11 ·'' bourrelets. In the
phyllodcs the pl.lles .trl' Vl'l) nowdl·d, a much larger
(I)
e Figure 9.24 Calopygus (Cassidulo1do; Crel ). (a) adopical and

(b) adoral surface (" 2); (c) enlargement of phyllodes and bovr·
aJ relets (x8)
n
IC
t- number of tube t't.·t:r <Kt ur th.lll d'e" hen· in the
1- ambuJacra and. furthl·nnort·. the por~' tend to be
e' larger. In the 1rrq,rul.tr et hmoids thl·~e extra tube
l). feet arc med pnnl.lnl) for t"t:edmg. Some. 'uch a~
:1- the spatangmd ,\ ft•t)l/111, lcl·d on orbran~e-nch ~and,
::m and the tube feet .lrl' ml·d as ·~ncky 'hovels" to
la deposit sand m the rnouth. lll·guhtr echin01ds may
ne also have phyllodcs, p.lrntul.trly thme li,1ng m
:ab l1igh-encrgy envuonmcllt~ whc:rt• the tube feet are
at- used for clinging to the sub~tr.tte. Prcstun.tbly the
extra adoral tube fct·t in Prolt'Wtidari.l .wtl other flex-
Jble echjnotds m.1y h.1vc serwd .1 similar purpose. A
l.Vt' tendency ro develop phyllodc:s h.ts bt·t·n noted in
cnt many lineages. 1 .Iter nwmbe~ oft~n have broad
L)lJ- phyllodes \.vith l,trger but tcwer por~ pau"'i. In m1ny
. Jll species the tube feet emt•rgl.' from 'mgle pores. the
and respirarory fundwn wlud1 m·n.·~"tates ~eparate
the channeh tor oxygcn-nd1 and o:\ygcn-poor
ime '.gure 9.23 Structure of Recent echinoids (a) Cenfrostephanus codomjc thud havlllg been lo't.
uch ~s (Rec.), Indian Ocean showing the ambital spine base -
·for re spine muscles attach to the raised wedges on the upper port
lnterambulacra and spines (Fig. 9.23a,b)
lOO opprox ), (b) Psommechmus m1liaris (Rec.), Torquoy,
)J'\15 hglond, on aboral pore pair with neural groove (x 50 approx.). Cidaroids, as we ha\'l' ~ecn. Jl"l' dl.lr:tcterized by
dif- :EM photographs reproduced by courtesy of Or AB Smith.) mterambul.tcral pl.ates h.lVmg a '"'~.de l.nge n:ntral
280 Echinoderms
tuhl·n k . l'hi, 'Y'tem wa' rL"t.lllll'd 111 genera such _,, demtpyramlds foremen erntd
Hnmcidaris anJ 1\,r/ori.r, tlwu~h the tubercles arc magnum h 111Ctl
gcncr.tlly smalkr. Ho\\ L"Wr. 111 tbl.' m.uonty of other 110( 11
cucc. hmoid~ the \Ill" ~)( the primary tubercle~ 'VC 117('
dct:rt'.t,cd nurkcdh. .md lllO\t parucularh in the feed<;
irregul.1r. <. )n th~: mtcrambul.ll ral plate' of S.lnd- lrrcgtt
JollaT\ dll'rc IIIJY bt.• up w tl'll prmury tubercle-; per Et:hlnocy•-
fccde1
1' ~
~qu.1rc millunC'trc .md hundrclh per pbre.
Th" tendency, \\ hil h \O greatly altered the whole
appear.lllc.C oi the rntl·r.unbubcr.tl pl.1res 111 the evo-
Eucldaris
+
Diadem•
laulodonll
Strongyloa~ntrotU5
f ,,ry HoriQr.l
P\rfl
u;.
l'.ll
h J\C ,
I + hgar111
j Echinol~~mpa
luuon of cucchrnOllf.;. ,, ltmctton.1lly \Ound. since rnr. 1
I
w-- ~ ·~--c~
the nn>dem t ilhtllid~ c.u1 lll'ithl'r bur,; thcm~clve~ in ! ,,f t hl·
Jctrituo; tor c..unmtH.l~t· nor burrow ...,horr spinl'S .ue c)f, tdl
.1 prcrt'c.llll\tte fnr hurr11W111g t'c.hmmds: they Jclso .fit· "\VJ
provrde covn .1nd sh.1dc. l'crl1.1p' rhe most striking
moditicuion o t :Ill l'l hinord~ is m rhc modem recf-
dwellrng regul.1r l't:hmntd f'l>dt>plwr,, m which the
.1dap1L.1I \p n te' .tn tllmluoom-shaped, their rops
;;v~ • Arbacla Ho/er:typus
I.H run
:turicl
w lmh
thl· 1.1
(sttrodont)
bemg lhr poly~nn.1l pl.ltl'' l(mnlllg .1 connnuous 111 .td l
bJ\.IItt!orm tiHl\,Jtt , wl11l h ~'1Vt'\ protection m heavy Figure 9 25 Various types of lantern pyramid w ith (in bloclj ~prc.Hl
~u rf; thl:~l t'l htnlllds 1 .tn .1ho rlu1g to the rock~ on shapes of lceth, keeled or grooved, in section The arrows repre- po \ th
wnt possible relationshtps. (8osed on Smith, 1982.)
whtch they ltw h~ thl'H phyllnd.tl mbc feet.
Gill
Lantern and perignath1c girdle (Fig 9.25) Gill n
The I nran t~ 111<\\t unport.tnr in Cl htno•d ta...,.on- Jun~'IC Dr.uiciiJ<lJ'•O lJllltl' like tlt~t
" lur:l\si
omy at thl· 'upc r-ordm.1l lcvd Although tt 1s r.1rc to .-1rdr,7t'<l1Jd,m•, rhough wtth deeply deft for;uncn
.1 ular' a
tind it fil"ilin·d , t' lhlllg h ~pt'l' tm c rh h.l\'l' now been nu~>num :md rhe cct·th .1rc bkcwi~e Vel') ~mt r llolt'O,
di~cm·ercd rn .1llcm .1 phylogcny of cchtnoid\ to be From rhl\ c.1rly rom,tock genu~ \Ccmmgly 01rosr v:tr. w lud1
cnn~tmncd U\111 g the tt•t•th .md l.mtcm as llUJOr rax- tous ':llllodom' type<. (e.g. di.ldcmatoid,). m wluch I \11'\' tl
omonuc t ntcn.1 rlw lkt.Jil' of the lanrcm and it~ tlw epiphv'c' ~:-rrnw im\ .mk .md - b} other patlu
~trucrurc .Ire beyond tlw \COpe of thi\ work, but ~nrodont .tnd C:lnl.trmlont l.llltt'MI\, m winch th M a,
H j, impmt.ltH (1\ •pprlt i.ue the nl.ljor stages in it\ teeth, r.lther than being b'TilOVed. ha\'t' .~ rtn· Scxu.JI
evtlllHtllll.tf) dt•H lopmcnt . I'hl· bntcm consists nounced \[rc.•ngtlwnrng kl·d In l.murmlont IJnrrm !lOll".
uf fin• 1dcnt1t.ll pyr.tmHf, , c.tch compmcd of two the cptphysn JOtll at the top . '>mnt.· of d1c\c lllltcm rm nnr
dcrntp) r.111lllk Hl'tWl't'n these j, .1 dcpre,,ed tt)rJ- type' dctim• ut.IJOr t.I\OIIOI1111 brrouping' tn th~ fl'lll.llc
mcn, .md .1bnn• ,m: 'hnrt, rod-Ilk<. cp1phy\e'i. E.1ch irregubr cdllltOttk m m let
pyratntd con t.tms .t smgk c.tlntl tooth which grow~ From J pm n itivc Dwdt•mop.m-liJ..c \Wck c.une thr o pe-d '
down ,·ontlJHl,lll\' .t\ tt t\ .tbr;Jtkd. ti rst irregular eehino i<h , in \vhid t tlw 1Jntem is dJI· t o 111tt
I'hl· cl:~ssil wm~ of J.tl bon ( 1'1 12) tlisrinbru i~hed rint,rui, h eJ by the k)\~ nr
rhc l'p lph y~c\, but in whtch chn:c.:d
four lll<llrt t ypc' of 1.11 ttcrn , cic.broid, aulodonr. the rceth hc:C.Hlll' indepc:mkmly kcdcd. From thl N t'.I Tl )
soroJont and t ' 111 t.trodnnt. bul ,l, Smith ( 19R I , pyga\tCrOid TOOl~tock ,J TO~l the h okt:t)' poid~ Jlvl \\ ltC i l'
19X2) Jnd tlthn' h.l\'t' ,Jlll\\ll, rlw r.mgl of structure tinally llK c.mJtlulmd~ ;md t lypc.Jsreroid,, thl' lJHcr hruod
and ttmctton ,, r.nht.•r mort• complex. lsol.ltl'd pyra- havmg rcmarJ....tbly l'\.p.mdc:d wedge-like tecrh. gott.h,
mid' 11lu~tr.1tl'd lll'rl' ~IHm '-l1111t' of the nt.lm bntcrn All thc,c dungcs ,m.: ·'"oti.ttcd with impro\C· md T 1
type' tn .111 interred l'\'tllutionary rel.ttiomhip Thl rncnt~ m fumtion . The l.mtern oft idaroid' CHI onl s ~ Han
prinntt\'1! ,m h .tl'llllll.md p~ r.nmd wa' rdativl'ly mow up ami Jown .mJ opt'n and ~hut like 3 tiH·· e1n At
bro.lJ , wilh lbttt'h ~:-rrom·l·d t.:cth , whtlc latt:r jawed grab. The c.tnurmlont IJntcrn, however, pal tco
t'td.trollh h.l\'t' a lll!lrt' compre"-t'd pynmid With a 'l
cap.1bk l1f \ttkw.ty' r.1ping. "hich ts r11uch more cold at
,h.tllo\\' ltlramen m.lgtHtm and deeply ~rroowd cfii:cu\'l' in bouom fl·cdmg, .md it' \\ hok orgJn!Z4· Au~t ra
teeth. Tlw rl'll'lltl~ dt,c.:o\'cn.· d lantcm of the l'3Tl) tion 'eem~ w hl· rd.Jtl'd w thi, cud. The dyp{'dlt· 'hUll!
crmd bntcrn wllh th cxp.1mkJ wc:Jgc-like teeth i~ a

functton,,} tnt\hlllg .tpp.tratus, ,\lld although it can-
nor move ''lk•w,l)' 11 1\ vl.'ry puwc:rtul ~md can pul-
vcnzc: th~· bottom dt•tntm upon \\ lud1 the echinoid
feed,, Tite l.llltt·m ha' bt·~·n dt,pcmed wtth in t!H:
tttq:ul.tr Atclmtnnuta , wluch .ut• nncroph tgou'
t~Jm.
P.tl.teuzu•~ cdulll)ld, othn th,tll nd.trotd' do not
lu1·c 3 tnlt.' pt•ngii.Hlm g•rdlt·. Jnd tht• 1\HI\clc~ .md
~g-Jments \H'n: .ltt.tt ht·d dtn.• t tly to the tmide of the
tot Thr.: pn•,t•nu: nt .1 ~mllc r.u,e~ the: ;lttJchment
of tlw mu,dt·~ .md llll't' h.tlllt.tlly tmpro\'es thctr hnc
t,farno11. C1d.tru1d' (lt~ t) I(,) lun: .1 primtttvc brir-
dlr wtth pr~JC:rllom (apophyses) in the mter;tmbu-
Jmun l.:.ut'l h11Hmis h.tvr both .lpophy,es and
auricles \llpport\ .11 tht· h.l\l' of tht .1mbulacrum
wluch. m 11\0it: .tdv.IIH'l"li t't hinmd~. m:~y .uch over
rhc tad1al w.ller ve,wl. I ht· lkvdopmcnt of ;ntride~
Ill .1ddttH.Hl to .lpophy\t'' prnb,1hly mcrt'.hl"d tht
l'l~.td of mu,rk· .ltt.1chmcnt .md thm ,tllowed tht
pn~sibthty at L!tt·r.tlmm·csurnl of tht· IJntt·m.
Gills
Gill note he" .11 t' found tn Cl hmollh !Tom the earl)
~c Juu,s•c onw.mh I hc\' :~rc prc,t·nt in nc.trly .111 rcg-
n clm ml\ Ill 'onw e trly or pnmstt\'t' trregulal'> (e.g.
r. Hlii'CiytJus. fly.~n(fl r) hut not m the l.ttc:r trrcgula~. m
:-- wluch pculmd :nnhul.!cr.l h.\\'mg cffiut·nt rcspirt-
h mry tube ti:er .ue d evclopt·d
Morsupioe (Fig . 9.26)

\,!.~tal d1morphl'm 1" not nonntlh n ident in echi- Figure 9 26 Pentechinus from the Tertiory of SO\Jtneostem
:ll>ld\1 and wht•n tt dm·' nrt ur tt j, in the tonn of Austrol1a, showing morsupioe (.-: 1) (a) odopicol surface, (b) cut
specimen morsup1ae seen from 1ns1de (Based on Philip and
mmor d1ffacntt'\ 'ud1 ,,, l.tr~n gcmtal pore' in the Foster, 1971 )
knule I lmvt n r. rlwrt• .lrt· \llllle 2H 'pecic" of
modem cchmosd' 111 whwlt tht• lnnak' h.tvc devel- evidence o n the norrhw.trd clntt nf tht· Atl\tralian
ht' ~·ped ~pcn.1l hmod poucltt'' (marsupiae) in which contment .tway ~rom Anr:~rrnr.t 111 the l.Her T l.'rtiary.
JS- hllllCllb.ltc the ti.·rt1lill·d l'~b'S. Thc>c then develop The temnoplcuroid J>mtcrltillw h.l\ (111 dw fen1.1k)
ch tbrealy wlthnut dw fiTt'-\\\ imming l.trval stage. each of the fi ve gcnir:tl port•, npt·nntg ulto an don-
he \icarly 111 l11 tlll'm livt· 111 cold Anlarttic waters gatcd dcpre,~ion cxtrndtng \llllll' \'v.ty down the
nd 11here dt•vclopmtnt .111d gH)\\.th .trc ,Jo\\. Such in terambulacmm. -whereas 111 fJrmrd!lXI'( lti1111' a deep
ter rood pouches on ur 111 rt'< t'llt ud.1rmd' and spatan- annular deprc~\Hll\ ~u rround~ the .lpit:~l dl'r, .md the
' 11ds. but tht·y .1re ,,],o finmd in 'omc Cretaceou~ gemtal pores open d1rccrh tnto tht\. J>,mHpatallgHs
ve- and fern.1ry ccht1111Hh nf Onkr' T t'nmoplcuroidea. has a shallow dt'prec;~tnn 111 whtd1 th~: .tpical dht lie~.
nly ~.ltallgllldl•,, .md l. lypl'.l\tl'nls(ka trom 'outhe.ht- Nomully 111 ma~upt1tc cchtnoid' the brood
\'e- ml Aum.1h.1 (Ph1hp .md [·mtcr. I ~71 ). \\ h11.·h !Tom pouches .ue .td.1pil :11. hm .m ,Jdm .1! m,tf\upium i\
• IS Flaeoclim;1nc t'\ tdcnre ~cc m' to IJJ\'C bl.'cll very prc~cnt 111 tlw clypc.l,rcrmd p,,_,SllfciStn. Tht> mde-
on· ro!J .11 th.u ttiiW. Tht• prt''l'IH't' of ,ut h genera in th.: pendem ongm of m.tf\up•ac 111 'o m.my cold-water
lz.l- ~u>rnhan l'ertl.IT) .1nd t11etr ,,b,ente 111 modern off- tonm ccmtie' to .m c:vtdcnt .ld\·antJ).tl' of vtvip.mty
asr- hort' Au,tr.1h.1n fnm.t ,tlt ord well with geological in low-rcmpcr:tture nmd•twn,,
282 Echinoderms
Evolution in echinoids Abrupt changes of this kind .tre seen 111 lttlnl•tt
The tint retognizabk cchinoids Jrt' Ordovinan, molluscs and brachiopods, :l11d the whole pro~.'" ~f
appeanng at the ~ame lllnc ,l, a whole plethora of undentanding phylogeny and 1ts functional )l!;n.t
other echmodem1 t)pc,, 'ome .lt k,l,t of which. m cancc is bcde\11lcd by morphological dhtOnnnu• "'c.
certam tcatun:~ of thc1r org.tmzauon, .lrc renruu~ 1tie~. Even though there are gaps m ccluuoul
cent of cclunoids. Amongst these Cl""luld he mnn- phylogeny, 'uch a~ in the c~t.tbbshment ot l.'lttdn·
bcred holothuroids .llld the echll1n7oan CIJss no id orders, they .trc not :tll th.lt abrupt .•md \U~oh an
Opl11ol·y,t01dea, wluch h.1vc enonnm1' plated tube 1111p0rtant stage a~ the brcakout of the penpr<11.l C2l1 Le
feet .md .1 lantern strutturc (Apparent re~embbnces be seen as a rcbtlvdy gr.1dual process. Tht· ~volu- C<
bennen cclunotds and cdrioa.~teruid' or t·hplo- . nonary trends within both PJI.tcozoic and Mc-sozo1,
porite' an: ~uperficial.) The l~aturc' of true echinotd to Recent c<.hinoid,, 'vvhich K ier (1 965, I 974) h1
orgamzat10n became mort• clearly Jctined :1 little c,trefully documented, occur in ~cveral group~. l~11t·
later. and by the C.trbomfcn.)U\ tht' l.lrge regular fymg to cont1nuccl ~election kading to 'the ~'ladu~
Aextble-tc,tcd pemdwec.:hmo id' With many tmprovemcnt of the amm.tl .1~ .1 living mcchJnbn
columm of pi<Hes were becoming dtnrse. though The remarkable functional difft:rcnces bernwn the
never rcJlly abund.lnt. Tht only ftmll echmo1d early Mesozotc ctdaroids and, ~ay, the ~p.lt:~ngl'ld Of
genu~ knmvn to cro~s dH: Pcrmo- Tna<;sJc boundary clypeasteroid~ has been achit:ved only by mod1tlet·
is Mioci,lt~rts, though it ~~ ml\\ bclit•wd that the tion of ex:istmg organs or ol dcmcnts .llrc.ld~ prnet
c
euechmo1ds were dt·nved !Tom a d1tlcrent Late in the ancestor. Even the IJ\Ctolc' of the ~p.ll.m~otdi
Palaeozmc lmelge. in the Tria..stt <trt' t()lmd small .1re merely trach of modified 'pmcs, the ciJ\ ubc, 1
ndaro1d~ .•md then m tht" orl~ JurJ\\tl c.1me ;l gn:.1t wh1ch the surface IS covcrt"d with J c1ltatcd ~opuhe
burc;t of l'llC'Chllloid r.ldJ,ll!On. Of tht:·~t· t:tx:l SOme lium of a kmd n,1rive to echinodcrm orgJIIJl,IUOI
:ue 'tmproved regubr; · and the rc~t irrebrtlbrs, In tact the only new kind~ ol org:m m tlw cucd11 L
T
.tdaptl·d for m .1ny rnodt'\ t:)l hfe. roday elhinoids arc noid<; not posse~sed by the1r 'id.1r01d tor~; be m 3rt
a vHal p.lrt nfrhc unertebr.He marine n:.1lm md thev the auncle) of the girdle. ~umc \ll1Jil compoucnt
are proh.1bl~ l~ abundant nO\\ as .lt 111~ ume m the rhe lantcm and the oph1n·ph.tlom (pmu.·r-hlr
pa~t. In thl' ltccent 6llll,l, 25''o are ,p.n.u•goids, 16"•u pcd1celbnat·.
cidarOJth, 14"{, clype.l\tero•d~ and 14'h• tt'111110plcu- L
Furthermort•. the funcnorul significanet• ot lll\ T
rotd~. The fortunes of V.lrtous ord~.·r, h;tve vam:d \tructural change~ in exlimt genera may be inter
b'Tcatly Junng the Tertt.ll"Y. for m~tancc. CJ\\iduloid' pretcd because of cornparablt· function,tl ld.lptJtlun!
were \'cry unportant unnl the Plion:-ne. \\hen the\' m many modem rcpresent.ltlVC\, using \llllll.u mod
declined b'Te.Hiy ..1\ dtd the clypea'itcrOilh. and many 1ficttiom nf org;ms generally present m t•chlll\lld,, Figure!
old group' such .1s tht' holectypoids .trt• wry nearly e.g. spines, fa<;noles. t:tc. (for the same gcnaal pur Creloce
extinct. po,es). In thts context the rt·cord of cvnluttoll 111
Mod~..rn t•~:hinotd, .trc very ~ucce"tul feeders. ;\ ficr.uter, J Cretaceou~ 'Patanguid. I\ e~rn~ul
Some even devour other echmo•ch: the proce~s of good. lt h.1s been elucidated \trattgraphic.1ll), t\'!tcd
such fcedm~ h l\ heen ,hown rccenrly b) rune-lapse \t.uistically and Interpreted luncuonally wtth rei
photobrr.1phy, wh1ch ~hmwd three l,1rge cida roid~ cncc to Its modern rebtivc Ed~t~wcordut/11 .111d othm.
devouring .1 ~.md dulbr. t•.Hing ir trom the edges like
a btSCUil. Evolution in Microster (Figs 9.27, 9.28)
Tht· prctl\l' rl'l.llwmlup ben\'een stnl( tural and ,\ ltcr.J.Ster ~~ an Upper Crctateou' genus wtdc\ptr31!
functional dttfcrenn.ltlon ~~ perhaps more clt:arly over much ot E.urope. and \'Cry common 111 ltilnt(
Seen m the cdlln01dS th,lll 111 olll)· Other lllVertebratl: and Engbnd .
group tly contrast w1th the cvolutwn.1ry pattern ot ln 189Y H.owe, work1ng 111 ~outhcrn En~Jnd,
most fth~tl group~. l.'volunonary ~.lunges anJ rarcfwly collccred some 2000 specimen' of Mlfwta
repeated trclltls in tht· Ml'\n7oic and I,Ht'r echinoids in ngid ~trangr.lphtc:tl sequence through \IX )Ul~bo
have been nther ,]m\ .md \prcad out !::lo often in \I VC 7011C'> of the Upper Chall. He ob,crv~.·d ~.un
other taxa the vitallv tmport:mt intnmcdtate stage\ dear change~ m going trom the lower zon.:~ 10 tl e
are cumpn:~'cd in time. and there arc no dear link' upper, confirmlllg previous suggestiom; tht~ w~•
between o1w gradt· of mg.mizltion .1nd anothc:r. all the more \ tnkmg in vtt:\'v of the nWIIlttomtu Figure)
Upper M west/a(et
Campanoan
I
•
t '-~
Lower
Campamm
Santontan -1_-=-t
M rostratus
M corangumum
0
r
ConodCo<~n
.s
n M decrptens
1.
1-
Upper
Turonoan M~~"M--•• ~0
re
U1
~u---
e) M lesket
Lower
1)" Turonoan
·e- Eptaster
M corbovrs
ns
d-
ds, f1gure 9.27 Evolutionary relationships and time ranges of the Epiaster-Micraster stock in the Anglo·Parisian province during Upper
.If-
Cretoceous time. (Based on Ernst, 1970; Stokes, 1977; Smith, 1984b.)
in
illy
te<l
(a) M lt~skei (b) M. cor8nguinum
er-
ers.
'l ... '. ~;.
·ead
nee
md,
·aster
ceo;-
ome
, rhe
'-''aS
nou' figvre 9.28 Inferred life positions and burrowing depths of Cretoceous microsters. (Based on Nichols, 1959.)
284 Echinoderms
hthuloh'Y throughout 1 hl• nu m ch.111ge~ Ro\\ e enmg of the anterior, tooJ ~'Tot~H'. thl· fi1rward
nouccJ on going up till' Wljlll'Ol'L' \\c.' CL' ,1\ follow\. movement of the penstomc: .111d thl.' dewlopnwnt o:
thc !Jbrunt arc all advantagl.'ou~ in thrL'l ring feedtnj!
tlw test become' bro.tdc.•r, ,111d bnrh rhe raJieq and current~ to the mouth 111 a dc.'L'p-burrn\\ ing crht·
bnlJde~l p.lrh move; rn~tenc ,.-Jy: no1d, ,\., .m: the incn.. asc m lllllnher' ol re,pirJtol'}
till' .mterior groove dn~pl'll'i .md tncre.tsc~ m tubl' fi:ct. Other than dw ~ubJIIJI f.ISl.lcllc: then· i~ no
tu berLulation; other dearly Jcvelopt:d tasnok 011 the: adapJCJI
thl mouth move.·~ Jntenorly , anJ till l.tbrum or rc.•g10n, hut the whole highly tubLrLui.llL' \urfJt:c. ot
lip below 1t becomes very pronoutH:ed eventually the .mtmal may have acted ,,, .1 'dlfhtsl. t:"ciulc'.
l'o\ enn~ the mouth; bc.·ing ron·rcd \Yith the ba\eS of wh.1t \\ l'fL' probJhly
rhe p.urcd JJnbul.ter.l lengthen .md become cl.l\'ul.1e.
'tr.1i~ln, whcrea\ the interpnnt\.·rom are.1\ change Not :11l .uc agreed that the .\I. lr~kt i-M .•vr:u,·
lrmn being ~mooth through .1 v.mcry of intcmlt:- .1?"'""111 Ime nccc<;sanl\ h.l, to do \\'Jth .1d.1pt:~ti01b fot
di.UC' unnJ the tinal p.lttl.'Cil '' om· with 1 deep dccpa hurrowrng. ~nuth ( llJ)-;4a) mtcrprc.·t~ .\lia,uw
groove per-r,tdially and mll.ltl.·d l.ttc..'ral an.:a~. evoluttnn ·" .111 miriaJ brc.tkthrnu~h to .1 m·w hdbttat
(from s;md-hvmg ro clulk sl'.t-dwt<llmw .tnd that the
Though Rowe unagi.ned th.tl th~.' onginal lim:agL' change~ ~eL·n J.m:r arc bc~t 1ntcrpretcd to more
had \Ubdtvlded more t.l1.1n L>llce. 11\Ul h btcr work dl1ltl'nt lit(: (burrmving and ti:t•dmg) WJthlll tlw
wa~ ll:qtured before the umltl'c.·d ta.x,momy could h.lhlt,lt. Figt
be ~orted our. The early '' 01 k of Kenn.tc~ ( 195-J.) ,\frrr,uta ~ecm~ to have bt'en uut1.1Uy ~mce,slullll ado
WJ\ greatly updated by Ermt (IIJ70) and Stokt:' th~ d1.1lk \l',l~ bcc.1mc lt had .1 den.se .td.lpH:al unopy fosc
( 1977). Jnd it j, 110\\ undeNtlod th.tt 1.: \olutmn
wok pl.11:c in rwo ,c.·p.trarc Jrc.· 1s - .1 north em
of riny nulury ~pine' \\ ht~ h kt'pt the 'cdllncnt ro~
off thl· cc h moid mrfatc. .111d .tllo\\ l·d 1t to li\'r
pro\'lnc.c.• <Jild till. An~lo-J'.m,l ,lll pro\'lnC~ - m tht infaun.1lly.
latter ot' which the phvlo~Cn) 1' hc.·~t known. A rc.'lll.lrbbly ;;imibr c. volutton.try lrnc.• h.t, bm hol.
.\lttr.blcr lt·skci i~ a ~m.ul. t Hhc.·r ~lohuhr cchmmd reportni !Tom Australian sp.tt;JngtLh. where 111 th~ tl.lt
with .1 ~hJUow antenor grnovl .111d the mourh Terti.lr) U11thi.1 led through f>.lltiSfCI to S.-hizc~stn . .'\11 ....
mthl.·r f:n b.td.. without .1 l.tbrum. The earliest rnte rhrec gcncr;l :tre st1ll extJnt. f.illtllltl, supt.'rtin.tlly hlic lt et
,\Jiutt lt'l onguuted from rhe l".trltc.•r gull!~ Epi,wn EJ1t•l•tt·r. live~ 111 co.trscr ~l'dillllllt'\, HmiStt•r m tiner att.1c
found m.Hnly m sandy tJuc.·,. l·ro111 thl\ came the.· m.ltl.·rial anJ Srl1i::astct m very tine mud Th~ .1llm
lu1c.' of dc~cenc ,\I, bkl't \I dt'itpirm ,1nd .\/ Ctlf<lll- Lh.1ngc' ;Jg-JIIl arc.· intt'rprt·tc.·d ·" .td.tpr.mom to hfi.·u: tron
.~~~illlllll 111 which the ch.m~c.·, norc.·d by Rowc and lint· ~l.·dtllll.'IH. tt:ret
tontiuned by Kerm;Kk took: pl.lc.l', I hc 'penes \UC- Althuugh the high zon.t.l m•~ r.t,rL·~ ,m.? tntrr- rent•
c.l"c.'U c.ll'h other with \ Cf) lmlt· mdicanon of interprt<ted .1~ del'P bunowcr, .•\/..~111/111> \\,1\ probably \ {'!)'
nwdt.tte,, .md tht~ C\t.lhh'hL·d hnL·.tgc.• is no longer ,h,tllow burro\\c.T (F1g. <J.2lk). Tlw 'ub.m:tl fN·tolc nl\\ t
etliNUc.red tu lw b'T.ldn.th\tlt. I hc.•,c.· ,\1,, r.rJIL'r spcoe> h.h gone, ,tnd 111 addition the pyr.umd.tl tC\t "'''uld tn 1111
Jre found on I) iu rlulk, ,1\ " .\I ..~tlllms (formerly help to prevcm partidc~ from wtrlmg. rhcn· ar~ In
ktHI\\ ll ilS .\f .IC'JHIIIC'IISIS), whirh tfc..\Cl.'l1ded from /If. very m.mv respiratory fet't, ,t]] of ,rn.1!1 st~c· rht, '!Jilll'
lcskd by a Ji fTcrem line. dc:vtcc probably cn~ured ag.1imt ptnl.trion AI .~ibb1u ll'\1 ~
Ntdmh ( 191)9) ~hcd much lil-)ht on the b10logtcal '\Ct'lllS tll hJvc been dcnvc:d ltn111 the mid-7oml of bt
meantng of these ch.mgt'\, bv \mdying the prcctse nuctaster~. po~Mbly a' .1 pl·riphcr.ll 1\nl.ltc . •md Ill tll h.
n. lJtturh betwcl'n stmrture ,md t'itncnon rn the bv- h.1w lllJ~rtatcJ bal. k into the \tlll thnn En~la nd .11'0 (jfltfitt
ing Ethilltlulrlllllt/1 and otlwr l.';\t.mt sp.1tangoids: the toward~ thl.' end of the Uppl·r Cret.lcenu,, whtn 'li.ltlllj
rnuJern l'4lli\·alcnts t)f All.r.J•ft•r. lt \\',1\ h1~ VJe\\' th.tt interbred \\ ith M IClrtiJ(~I/11111111, produnng .1 number (IIJ cl
,\I, lt~kci ''as ad.tptL·d tu ,Ju llm\ burrowing JUSt c1fhybnd~. f.tll'l'/1
below the surtau. and th.l! thl' ch.m~l'\ m rhc main In rlw prohfcrauon of both 'P·•t.lllgoiJ, .md hola stabk
lmc ot' dc~cent from .\1. lohr tn \/. CtiY<IIIgrmwllt tc.rmd~ m the Lrctaccou,, nunlc.'nlll~ type' aro they ~
giw c\ iJeme of ad.Jpt.ltlon ll1 progrL'''ivcly deeper .td.lpted tor dt:c.·p or ~hallo\\' hurrow1ng, .md 1t lu WL•dg
burww111~ comhtium (F1~. ').'>H) been poss1blc ro intcrprL't sonll' ut t hc.·~c.· fuuniclll· itJh,tbi
T'he~e .1dapt.1ttoll~ sec.·m tlcar enough. The d~:ep- .tlly. L'dtillllft'Y)IS (Fi~. 9.29,1 J) 1~ .l high!} ntnJ1Iinl rot k111
(1) ctrcum"ral nng
buccal tube feet ambulatory

tube feel
Figure 9.30 Hololhuria, sectioned longttudinally. (Based on

Nichols, 1974.)
c
e
~~
Figure 9 29 Echinocorys sculolo (Cret )· (a)-(c) in adapical,
adoral and lateral views, showing marginal diffuse and perianal
lascioles (stippled) and current directions (x 0.6); (d) detached,
pocldle-like spines from the plastron (•d 5 approx.). (Redrawn
LJL
Figure 9.31 Holothurian sclerites (anchors, gratings and
wheels). (Based on Frizzell el al. m Trealtse on Invertebrate
lrom Stephenson, 1963 ) Paleontology, Part U.l
~olmcr01d \\ Hh .1 lu~;h ~tet•p-,Jded tl.'\t. an ahno\t forward uno tht• st.•thmcnl; the wcd~c:-likc shape h
ti.Jt hJ't and a ,h,trp amhitu... esscnttal for tlm. Domed 'patJn!;oid, \\'tth a tl..u H'n-
Short thorny 'PlOt'' wc:n: cndcntly tor the pro- rra] ~urf.1cc (e.g. Li11t>p11mstcs) do not burrow. but
ttcuun of dw .td.tpll'JI ,urfatc:; 'patulate spines hve eptfaunJII> on mud. ~lH. h modlficatiom of te~t
ltLidlcd tu tht· tubcrdc, of tht• ador,u ~urtace ,hapc \eem ro have bcc:u of kev unportance m
21lo\\ c:J forw.trJ pro~n:'''on .•mJ cili.uy lurrents enablmg ~p.ttJngutds to adapt to .1 \\ide ran~e of
Ul from the lll.lt~ll.ll Jttli.Jse t:hoolc: ,\ud the more dif- habttat~.
frrentuted 'ulwul t:t'l iole prodtKed ti:edmg cur-
er- rent\. Tlw ,unbul.!t r.t arc only subpct.tloid but are
\try long. EthilflltVty~ "'"' prub.tbl) ,, shallow bur- Class Halothuroidea
' .I
ok ruwcr but pre,um.1bly mh.tbttt·d ,, dilfcrent tut.he
uki from the htgh zun.tlmil't,l~tt·r~ (Stcphemou. 1963). 'There were nasty green v.arty thing.<'. like pil.'kled
are In spatango•J, ~t·m·r.tll>. the arrangement of gherkins. lymg ou the bt:ath · (I I. G. Wells. At·pyorlli.s
thl'> 'pmcs '' t:mly nmst.llH, but tht'n: i\ a wide variety of lsla11d, t 927). Thus has the t'Xtt'm;il appearance of
JbiiS ''"~t shapes. Thest .ut· ,Jtbpted to different methods holotlmroicls (holothuriam or se.t cucumbers) been
)1),11 ol bum>wmg (K,ul,\lJ\.,,1. l ';N~). which <~rt: rd,tted most gr.1phicaUy tksuibcd. llolothuroids (F1g. 4.30)
l to to habJt,lt. l'hm ~lubular eduno1ds (e.g. Echitw- :.tre fusifonn or cylindncal t'dunozoans which gcn-
,,n:: a .rrdirmr). wluch hH· dt.•ep 111 sand. burrow by exca- craUy ltc on the: sc<1 floor (or burrow mto it) wlth
·n tt utmg ~edmwnt trum tht.· front and an umulating it their long axis honzont,ll.
1.bcr on the back. 'I hu~t ,., •th fiat protiks. such as The mouth .md anus .trt' Jt nppoo;ttc encb. In the
~Awcm.r. livt· dme to the 'urt:1ce. though they are type example. H,,Jllrlwria. tht• mouth end l1.1s a teath-
1).1~ 1t.1blc agam~t current 'counng. ,1110 when burrov. ing ery mflorcsccntl' of ~ticky tentacle' (the b u ccal
.[0\e he\ pu'h ~l."dimt·nr to the pmtt. nor ~u:k~ of the rest. t ube fe et) which <1rC' usc:d m tccding. The ,km is
- has \\edgt.·-~ haped ,p.tr.tngo•ds ( ~-lu~<~stcr. Bnsastcr). warty and lt'Jthery. and wtthin it are embedded the
JOll- mlub1t 'oti- l nht',l\'t.' mud .md burrow by repe.ttedly calcmc demt:nt\ tlut are the only part~ to be tiJ~
itlcd rockmg tht' Jntt•rior t•nd up .1nd down .md sinkmg st.hzed_ These calcmt pl.nes .\re norn1ally rt:duced to
286 Echinoderms
!mic scleritcs. h.wmg thl· ti.mn of anchors. granngs h olorlm nans, whtch creep along on thrtr W11tral
or 'pok.l·d '' hl·d, (r:i~. 11.J I). tube fct't or on J mmcular, ~lug-hkc sole 111 \Vhtclt
Ven r.1n:ly they timn .1 cnntib•uom covl·r; more t he rubr fret arc reduced. llowe .. er. one of the ~rt
u~u.tll} the intq.;uml'nt " wry tkxJblc. 1£ is from doch irore suborder\, tltl' Fl a~rrodJ, h.1s S\\ltnmmg 1nter
\uc:h 'lkrite' th.tt ft)"tl gl'lll'r.l can bc Identified and reprcscnt.1nves whtch keep themselvc' afloat bY pul-
alloc.ncd w t:uniJtl, , \. )nJy 111 thl· mourh regton 1' ~arion , hke jdlyfi~h. and an.· pl.mktn·orom. Some at
there a ri~:,rid 'c:ric, of pl.ttl''· fomung a cakareous th l' bl'nthlc gl·ncr.1 ~Wll'p the ' l..'J floor .thod 11!
perioral ring. pcrh.1p' homologou' wHh the echi- them a' thq muv~: with thl"lr 'til·ky bucc.tl tubdC'(t
notd Llnrem. ~nd 'o pic k up org.1n1l dctntul>; otltl."!\ 'tm.:ad chm~
The ti\t· ambulalr.l .m.· ~rrangcd 111 two \et' paral- out in .t horizonta l collecting bowl, a' d,, the
ld '' ith the Ion~ .txi': thrl'l' VL'Iltr.llly ,\11d two dor- \luggt,h, burruw111~ Apoda.
'aJI>. The tube t(..et nf the do!\,tl 'et fonn small Holorhunan' rod.1y an: .1hundant in w.um, sh~llnl'.
sensory papillae, wluk liHN' of the Vllltral set arc w.m:r\, but they art ahu sttttes~ful m the deep so,
ambui.HoT) .111d h.t\ l \lll k.c!\. Thl' mtl·nul struc- where u ndc!\l'J photob•Taphy ha' shown dcndmcht-
tu res arc m uch 111mhliL·d. fhcn: .m~ ,1 p.ur of respi- rotcs 'congreg,ttmg tn hads like gr.tzing cow~·
ratory trees .tmtng lrou1 the 'wollcn cloaca at the
hind end of the lll[e\tlltc. Water Ltllll ing in thro ugh
the anus is Jlll111pl'J by rhythmic puls,ltions of the Class Edrioasteroidea
duJla l!lto thew trL'l'' filr re, ptr,ltOI) exchange w ith
the lOdomil fhud. In Subda,, A,p1dochirota, to Thc EdrioasteroideJ (L. C.1111. L C.ub.) are .t 'mall
which HoiMIIIII'i<l bdon~:-."i, thl·rc an: unusual defen- cxri.nct ~oup or et hinodenm with fiw di\tlflll
~t vc ~tructu rc~ lol lted \\ ttlun the body. These are am bulacral a nd tivc mterambul.tcral are.ll>. con.tined
the cuvierian o rgans thread-like masses w hKh to rhe upper (ador.J I) ~urt:lte. These run to J ccnt!1!l
can be ~hot llltt of tht .mm to cnt.111glc ~tick.il] any momh. M ost ednoa~tcrotd\ arc flattened anJ dt'·
predator rlur di~turb' thc holothurian. coidal though some ,1re ~l obular or dongatcd.
The earhl'\t lwlothun.m renuu1.' may bl.' of J.:d,iMstrr (M . Ord.; r 1g. lJ.32a,b) ha~ a lieXJb!c
Ordovt~.:ian .tgc: tlw c.trhl·~t undoubted holothurian theca m whKh tivc btscri.tl .unbuiJcra .1re re~rularl)
'>ptcuJc, ( Pc~/.tct•llt(lllll•m•l) .m:. howe\'cr. from the plated :md alternate w1ch irrq:~ularly plated lllteram-
Devoman. L:t•t/um.l (CJa~, Edunotdca. ~uperorder bulacral Jrcas.
MegaJnpod.tcea) rm~lu h.t\'t· been somewhere T he ambulacra are 'iuuous, lour haYing an ~lltl·
near the ance,rr.tl lml' .md mdced wa~ first described clockwlse rw1st whereas rhc tlti:h turns clocbm. lt g i
as a pbted holmhun.m. RcJucuon of the plates ;lnd curb round thl· cccentnc pcnproct. A •null
ro ~clcrite~. ~VIII~ tht• body tt\ present fl.eXJbility. b ydropore near the mourh presumably lcd to the
seems ro h.tH' been .tlnto\t li111H'!\..tl. A very w.ltcr-vascu lar syHem. ThL· ambubcra h:~ve two ;en
pecu liar 1 >evonran ophi1lt y~totd-hke echinodemt, o f p lates: a lower ser of flooring plates. arrJJ1ged
Rortbamrs, has .1 l.trge lantnn, but the skelewn i~ as in an erhin oid, the ma rgms of each pterrcd lw d
reduced orhen"1~' eo whecl-hke sdentes. T lus too single pan·: and an upper set of cover plates,
m.ty be .1 holorhun.1n. Slllcc the ~keletons o f the hinged o n top of the fl ooring pbtes. \.vluch could
maj oriry of ho lnrhtmam arc red uced to microscopic op en to let o ut the m bc tcct o r clme down to pro-
sclerite~ only, thc tomh1.1t1011 potentiJ I of intact tect them . The peristomial plates covcnng the
spcC!m em is vcrv low. fhere .1re, however, rare m outh were probably tixcd (Js in the cnnoll~ f•gvre
examples of complete ~pec11nens of th1s kmd. The tegmen) and could nor open. The whole ambu- remove
best-known taun.1 ~~ from the Middle Triassic of lacral plating sysrem I S strikingly remimscent of that Stroma
a prec1
northern Sp.1m (~rmth .md Gallenu. 1991), where o f the Crino1oa (q.v.). As 111 that ~ubph] lum. tht ombulo
supcrbly p res('n·ed pbred spenmem ~how ,t consid- radial wa ter veo,sel m:~y have mn in J medJJn groovt ((o) Bos
erable morphologic.tl .md ecolo~cal dl\'ersny. Only along the pcr-r:tdial sunm:-. though Bcll (1Q76). in a on lnve
a tew ph red gena.1 extst cod:ty. all belonging to the comprehcm 1vc work, has suggested an JJtcm.lO\'t
Dendrochirot.l, which '' ith the Asp1dochirota and pomion. There 1s some evtdcnce for thts, tor mco ,tbural
the Apoda tonn the thrt·e detined subclasses of Class of both the circumoral ring .md the radial water ,tmbul
Holorhuroide.1. The tiN two are mamly benth.ic vessel have been p reserved 111 E. b~tchi,mlls. Tht mg ,1 1
In ~p1 tc of tlw ~irnibnty of cdrioaster01d .unbu-

lacr:J to crino1d food grmwe' there wen~ no brachi-
oles ('hem fingef' ~.:xtemhng lrom the te,t}. T he
mterambulacrum floonng plates nature of the ltlbe feet " unknown. Edn'tlclsft•t may
.l'
havc been \e,\llc but \\ not hxed to the substrate
.md might havc moved ,tlon~ by pul~attoth of the
Aextbk theca
T he 'upc:rtiu.1l n;\embl.tmc: of edrio,meroid~ ro
the Prc:cambn;tn 7 rrlmJtlmlr'um ha' often been noted.
~hough tlm may be: no more: than comndental.
Though the ong111 of c:dnoa~tero1ds I' re,tlly
unknown, the .1 bm.d feature' th.1t they \hare with
early crinozoam wem to suggest a common nngin.
(a) Edrioaster
There arc tivc gcnc:rJ of C<1mbn.tn .1ge. of which
Stromalorystirc_, (L. Cam.) 1~ the earhcst, and indeed
amongst the first o f alJ eL hinoderms. it JS of pentag-
onal 'hape and h<ld tiw ~tra 1~h t ambul:tcr.1 w1th
cover .mJ flooring pLlles. lt was .1pparently a free-
living form. lt ~~probabl y best re~arded a~ belonging
to the '\tcm group that gave rise on the one hand
t to thc other four Cambrian genera, and all l.tter
~ cdrioa\tcr01ds, .md on the other hand to the
I deutherozuans (all !Tc:c:-livmg cchmodem1': F1~
9.32b, !/.51}. le thus ot.:u1p1e~ a central plal'c in
edunodenn phylogeny
c Edrioastero1d~ rc:1chcd the1r acme in rh~: Middle
(bl Stromatocyst1tes
Ordo~rc1an . ·1 hey are not uncommon in the late
r
Ordovtoan, but rhcrc 1fter are found only in ~mall
number\ until rhc larer ( .1rbomf( rous w 1th a slight
(- expansiOn toward~ the end. Vtrtually all known
;c ligament :.pe<. i ~-. come from northl·nt Europe and North
1!1 Amenca. Some genera wcrl' frel·-living. but mmt
IC wc:r~ pennanently fixed by thl' margtnal ring to the
ts substrate. Somcnmes hv111g .. twlh Wl'n.. app.m:ntly
:d used ;ts ,, ba~e. Jnd the h1gh \electivity nf p<lrtil·ubr
..r species for onr k111d of \hdl ~u ggt:\ts that these may
:s , rad1al water vessel hav~ been commen'ial.
Id The m:tjori ty of gcm·r.J were di~coidal and nor
n- (c) unlike Ednoastl'Y. In ry~l(Otys ris anti Rht'IIVCysti.s, how-
he ever, there •~ a long rower-like stem of imbricating
·id F.gure 9.32 (a) Edrioosler bigsbyi (Ord.) with cover plates plates below rhc ti:'lr .1dor:1l \urface; Cyatlwcy.(/ls is
\.1- <emoved from three of the ornbulocro (• 1.5 approx.); (b) also colunm:1r. The,t· U)ntl\ ntJy luvl' been penna-
Stromotocysliles (L. Cam.), a stem-group echinoderm, probably nently t-lxed 111 ~ofi mud. In general tem1' ednoast-
l.lt
o precursor to loter edrioostero1ds (x 1.5), (c) section through
he ambulocrum of edrioosteroid with inferred disposition of soft ports. crord~ seem to be mmt abundant whl'll c:ucrusrmg
oVl: !(a) Based on Kesling, (b) redrawn from Pompeckj, both in Treatise hardgrounds. Ar one loLalnv 111 the Devonian of
, a on Invertebrate Poleonlology, Par1 U; (c) bosed on Smith, 1985.) Iowa (Koch :md ~rrimple, I 1J6H}. lllJil} ednoast-
LVC eroids (Allc/,zrnwtn lr.wor•rn) along \\ rth the coral
ccs 1boral surt:tcl' cmht'b of plate~ like those of the Au/op(lftl .md che cy,rmd ldtlcfl!ltfSib are present
.ter unbul.Kr.t. but thl· rnargiual pl.ttt:~ are larger, fonn- in what mu~t h.wc bet'n thei r hfe habit, att.1ched to
"he mg a dl,tlllrt .uul ng1J nng. a dtsconttnmty \Urtlct'. Tht· 'ea Auor <tt that time
288 Echinoderms
comi~tcJ of w.tvt·-frt•ttl.'d limt'\lnm·, t•roded into anns ~

knobs cJno.l\lerottl\ ,tn: ftHttld fixt·J to the sloptng- Cln th.
surfaccs of tht:\t' knob': .tll ~tagts tn b'TOWth are JU:I:cd
found. Anotht·r eJnn.t\tl.'mtd spt·ue\ lived on the l:11.:r:•l r
,jJe~ ut the t ~stotd thc.·c.·.tc. The.· wholt: as:..emb!Jge pore.. t
was O\'erwhdtm·d by ..ilt .•mJ JU 'P'-'cimem are pre- VC~'>cl!
~crved in place. t.lu.• ad.
at~ All
ngtd p
Bet\
pr~JCC:l
Inc. w1
Jl UVt' ,
A\tcrozu.tm (Cl.t" Stdkwtdt·a) .trt.· grouped in three C\\t.'llti :
'>ubdasst:s and utdlllk tht: prinutt vc Subclass ,l\lt'I'Oil
Somaskruttk.t, \l.llli\h (Suht IJ" A~tcroidea), and { .ucn~t
brittle -.tar~ (Subd.t\s OphiuruidcJ). They arc rela- ht.'l.'ll p
tively rare in the.· hmil rnnnJ \incc they break up c.tr!) J'
very c.lsily .tftt•r Jcatlt ..tnd llu: pmcity of their timil .unpull:
rem,tim ., no ~lude tu tht·ir tonncr abund.mcc. I ltc
Ncverthdt·ss the.~ m: .unon~:-"'' thl' mmt abuncLnt of t'Lll ti-c
ammah living tud.n 011 wd.y ~hore' of continental C't"hlllO(
'ihdve!.\ .md 111 tilt deep w.t. 111<1 \'Ill!!
In aU .lslt'rtJZtl.ll\\ the <.t:ntr,tl p.trt of the body dctnt.JI
(c)
(tli~c ) c.-xtt"nd' l.nc.·r.tlly into t1w or more: a rms. rhc < >rdovi
momh fJcc\ do\\ nw.trth .md tht> .mm. where prc.·- ha.~ bet
sem. t'> .tbonl. ltt ,1\tt·Hml, thc .trm' arc not 'harp!} ;bter• 1id
marked oil' ii-0m tht· u:ntr tl dJ'c. \\ hcre.tll in ophi- •.1pac1t)
urOid~ tht' central di,c j, dc.trh ddinutcd and bear\ stmply I
tivc: Aex1bk 'nakc-hJ..c .mm (Gn:ck. <>$toe; = (Hhkc .
~nakc). Mo\t uf th(' \"1\t t't.tl clcmt:nts in a~terozoa.r1,. hte of e
mch as rhe llltc.·mnt' ,\IIJ ~onJJ,, .ne broadly homol- .!IJd W it
ogous Wtth thn,t• in othc:t c:thinodcnns. The rcl:at•d
W:ltcr-V.l'illthr 'Y'tcm n.:'t'lllbks that uf l rinoid' in IIIOdl'nl
that the r.tdt.l l WJtcr vc,,c:J, lie.· 111 deep grooves in mollusc:
the vcnrr.tl (.tdor.11) \llll.tt e. flm ts one of the many When ~
ti::aturcs th.lt ~li!!Mt·~t ,1 rd.ttintl'.. lup v.tth crinoid;; ·hmbs 1
rather tlun et httWilb.
'" ~t1Ck•
lcmgcr 1
8Vt'll nJ;1
Subclass A steroid e a (Fig. 9 .33a, b) till' b1va
Figure 9.33 (o), (b) Asterord anatomy· (a) external. with food sltghtly.
A,teroid' u,u,tlly h.l\'l' tivt• .mm, though the~e mav grooves and Aorescent gills; \b)cross-section of arm, showtrog
tt' mou
be furrhcr 'uhdi\'itkd. Thc.;e .mm nonnally extend plating structure and externa position of radial water ~Id
(cHe) Ophiuroid anatomy (c) adoral wrfoce, (d) crass·sectrcn opened
like digit' from tlw dt'c, but \Otnt• starfish h.we a of arm with ver1ebroe and rod1ol water vessel behind; (e) loler~ \\'tthlll t
very penr,tgmul outhnt• wtth 'hart anns. On the view. [(o), (b) Modrhed from Sloden, (cHe) porltolly redrw11 plt<rl' tht
upper 'un:llt' then: .m· .1 'cncs of pl.tte.;, whJCh !Tom from Woods, both in Treatise on Invertebrate Poleontology, Pert
U; (cHe) also partially redrawn from Nichols, 197.4) Afr~.·r tet
the rop down .trt• till carinals. dorso-la t er als and lllOilth~
marginals. Tht·,c.· 11\.1\ mn:rlock o;o that the tesr ts from d
rigid. but more oftt'll tltt')' .m· fkxtble ..tUowmg the hJVah·~.· •
Subphylum Asterozoo 289
~mh to curl, whtch is wry u~dul durin~ dimbing. tccdmg habtt is .ttll'lt'nt, e.xtcndmg ,l\ f:tr hJck a~ the
On the lower surt.H;c: th~· pl.ttt's .trc !Jr~er and org-a- Dcvoman and prob.1bly fi1rthcr. In J well-known
mzed 111 l'.tch tmt uno .1 d<1ubk rulumn u~ ambu- example (Clarkc, I 912) .1 large heddmg \urfarc of
bml plate,, thrnu~h whu.: h the tubc t(-ct connect by Devoman rod. rcvc,\lcd ~ome 4011 'pennwm of
j.X're~ w thc .nnpull.lt• I.'VI.'Il though the radi,tl water Del'<'llastt'l' e11dums in illlllll ttl' a\\lKt.ttlnn wtth the
\'C"selt~ nn th.: mmidc nf thl· tl.'\t. <. hmtde the'e are bi\·alvc.. CrammyJ~<l .llld f>tmm·cl, the environment
the adambulacral plates \\ htch .ll"!Jom tlw margin- having bl'cn 'inv.Hkd b~ \tJrfl\h which conhrrehr:ttcd
als All rh~: ,\dor.tl pi.Ht'' llllltl.' .trnund the mouth in a m v.tSt number; 111 order to ti:l'u on the dam,·.
n~d peristomal ring. There i\ now soull' doubt about thi, daim, and in
Bem·l·e•1 rlw pi.Hc' (us~kk·~) of the upper surftcc any case not ,1U J\ll-'rold\ feed m tlm way; ~ome feed
project extem1om ni thl· col'lom . known J~ papu- mtr.lOr.tUy. a~ do many ophmrmd'>, b}' taking thl'
lae. which .tn· rt•,ptr.ttnry; ,ince .htt·roid\ are very prey. whl-'thl-'r gastropod or bivalve. ultu the mouth.
lthve .lnml.lh, 'uch \·xrr.t' rt·~pitatory stntctures ,1re Even here. however, Jntrll\1011 of the .'>tom.1ch lobe'
~~t:nri.1l. ' J'hl· w.ttl'l" · V.I\t tli.tr syste111 ot" P.tbeozotc mto the b1v.tlve ,lppNr\ to au:omp.my Jige,tion.
Neroid' h,,, bet•n rnomtntcted (Bhke and Other ~tartl~h ,1re dt•postt teeders, and basket ~tars
Gut•mh~:rg, I1>RH). Ampull.w m.ty or m.ty not have :~.re smpemton feeders. Smu: some stadish mJy have
been prc~ent, .md if prl'~l·nt Wt:rl' l'Xtern.ll in the preyed actively on blValvcs tor at least ..JO(I nulhon
ml)· .t,knnd,, h) romr.l\1· with the large internal years, 1t 1' highly probable- (Cuter, llJ67) that some
lmpulbc 11f mmk·m types. of the rh.tr:~cknsttc physJt>log~tal .111d stn1ctural
Tlw fir,t \t.nh\h proh.thly dnivt·d from J pent:ml- ch.tractt:nstJCs of b1valve ~hell' (c.~. tht: interlocking
crnl tfec-bvmg \tl'nt ~roup, which included such commtssurt·~) can be comldl'rcd as dcfemive \true-
cchmodt•nn' ·'' Slrcllll.t/tlry~ttii'S, tht· e.trly members cures ag:unst ,tstao1ds. the C\Tr-present enemy.
movmg lrcd) ttn'~' thl· ,c,t tlnnr .md feeding on Indeed the morpholo~rtcal mnovanou of mterlock-
J~mt.tl partidc, , fht· t'.trht:~t tntc .t~termd~ were mg commt~sure\ m b1v.1lvc~ arose Jrolmd the same
Ordo,•ici.m, derived lrom .1 \lllll.l'tt·roid .mn·stor. Ir time ::1~ the gn:.lt cxp::tmton of extraorally feedmg
tu, bt·ell pmpmcd (C,tk, I'JX7) th.Jr rhe'c early asterOid!>, m the later Ordo>JCian and earlv Silurian.
~ternid, wen• rel.1tiwl} infkxible. ,,·ith limited lt" po\~1ble. too. tlut the failure of amcuJate brJ-
.ap~uty fi-,r .mu ntoH'ml'nt~. Jnd they mJy h'!!ve fed chiopods to radtate anew following the late Pem1ian
ltmply hy 'cdum·rH 'howlling. An alternative \'lCW exrincnom. 1s dtrcnly hnkcd to the early MesozoJC
r13bke anJ Guen,berg. l 1lXX) j, th.1t the modes of radianon of .&SterOids (Donovan and (de. 19911).
tit~ of earh• Jstewid, \Hrt: dtwN.'. Jnu bro.tdly par- Agamst the'e the eptt:nul.ll br.1rh1opods had linJe
:illd wuh rhow ofmd.1y. A,tnotd radiJtwn" clearly protecnon, <llld were C\pcetally vulnerable.
rel~tcd ll1 ,, dtvcr,itic.llwn 111 lt·edmg habih. Most
modem ,t.lrti,h .lrl 'maunu' pred.1to~ and feed on
molluo;c,, 111 panintl.u pn·ying .llttwly on biv:l.lves. Subclass Sornasteroidea
When a starli,h 'lllh .l, J.,trn<~.' loe:Jtl'' J hiv:l.lvc it
climb~ on cop ot it, en~<LKl'' the two valve~ with The carlic,t known \Larfi~h' (e.g. I 'illcbrrma~ter,
ttl suckt•red tube leet, .tnJ tnr \t'\'t:r:tl mmute' or Chinianant•r) are Trcm.u..luci.lll and come (rom
longer pull~ the two valvl'' 111 uppo~itc directions. southern Fr<lnrc and Bohemia. Thl''l' belong to the
Evcnruall) the ~ere'' \l't up bcromes too great f(.,r Somasteruidea, a short-livl'O gmup probably ances-
the bivalvt· \ ,Hldllltor lllll\tlt:~. aml thc valve'\ open tral to both thc A\tcroide.t and tht.• Ophiurotdea,
1bghtly. 1 he st.lrll\h tht·n cvtTts its \tornach out of though primitive: ophiuroid' occur m the ~amc
Its mouth and '>quccze'> ll bit by btt tmide the fuun.a.
upencd vain.-, Dl~t·stion ot' t ht· '>Oft part\ take' p!Jce The: Soma!>teroidc.l h:tVl' a pentagoml hod) ~hapt·
1•·nhm the biv.1lw ,ht·ll, ,wd \\'hl'Jl tceding i'> corn- and the: am1~. rather than cxtend111g from the hod>,
pier~ the: ~tonud1 rcturm imkk tht• st.trli'>h ag.Un. are jmt bq~i nmng to dlllt•rl'ntlatc. The ~kcleton
Mrer fcedmg. tlw surti,h l.m gu \vithout food for COnSiStS of tJvc bi\COal r()W\ uf piJtl'' fom1in~ the
1110nth\ \\'lthout ill l'lfnl\. Tht•rt• j., \Oilll' evidence ·anm', which haw latcr.ll rod-like 'virgali.t' project-
from characteri,tic ,1\\ul'iatiom of dt,JrtJculated in~ from them, falling up tht• 'P·In' bctwt:en the
biv.llw ,h.:Jh with ti.'"il surli'h that tlm extraoral arnn. The ,l..cJcton ha' ,l thin rim of rn.lrbrin.tl pbte,,
290 Ech inoderms
In some respt·tts structure uf the arm~ fl'~t·mbles that oceans, espect,<lly :n bathyal .1nd abyss.1l depth~. .llld,
of the pinn.1e of a bisen.d inverted crinoJd, .md some where they .1re often found crowded together 111 <oul
p.llaeontologtst' beltewJ th.u 'omJ\tcrOid~ were great number... They Jre mainly ~uspens10n tec:dt•n, n>ul.
derived from l rinmds. 1 hh vie\\ bl'CJO\t e~pec1.11ly but there art' tJmivorous ~peCJe' wh1ch feed on 1.1~: r.l
popular when rdl (1963) dc,cribed .l Rcl'Cilf deep- mull bivalve,, though the'e do not have the l'Xlrac... ~han
~ea genus, Pf,lftt~ft'llciS. ,l\ ~ hvi ng ~om,Jstt'rold, and r:1l teedjng hab1t tJf asteroid,. OphturoJd~ Sl·em to .l \~:
proposed v.1nom 'tagcs i11 the dcscl'nt of ~omas have retained llllll h the same org:mi:zat10n ~111n th~: norn
tcrOJds from t nnoid~. La tu \llllht:s mgg~:,t, how- Ordovicia.n ..111d thc1r genL'r,l, likl· rhmc of Ncro1d~. \\ clJ
ever, th.lt thi\ lll,l) be llllft.' lh\tiC and pf,lltlSICrias ha~ arc very lon~-ranged. They dn not ~eem t•~ h;m:
been re,tudll'd m great dl·ta!l by Blake ( llJH2) who l~ccn badly at1ectcd by any m.lJOr extinction pennJ,.
di,clairm Fdl\ vtew th.H tt " ,, living ~om.t,tcroid. In
his csrimatton tt 1s .111 unetJlllvocal astermd, though
primitiH in \Ome respect~. such as the .tb,ence of Starfish beds Cri ne
\uccon.U tubl· tect (wh1ch 1t does not need \UKc Jt 1' ('pdn
a microphagou' feeder). lr 1' now regarded J\ .1 \Ub- fhe rcmams of fms1l asterozoJns are all too 'cmr,, lOillp
genus of the rml' ,1ster01d L.lltdltl, and it~ morphol- smce rhe plate' nonnally dt~.,rnculate aft~:r dt'.lth, lTillOl
ogy, rather than being .1 u1nsequence of crinoid not being bound together ,1s ,lrL' those of elhtnoidl. \Ct 0 11
ancestr)-, i~ Cllntmgcnt on beh.wiour aml h.1b1tar. Sever.tl well-known starfish bL·ds are known, huv.-
The tmt tnw a;,tcrUJd 1' of early Aremg <~gc, and cver, in North Amenca. in Grc.lt Britain Jnd in th<"
b~ the M1ddk· OrdovJCl.m the som,t\tl"rtlld-; had Oevonian of G~:·nnany. In the latter c;c.e ~tarfi~h att Clc
largdy dl\.1pp~:•arcd. Mmt R.ecem spel tt.>\ of the almost alway~ .mociatcd \Vlth the srrang~: .lrthropoJ<
A~t~:ro.ideJ. ltke rho~e of tht• nrhwroJOC;I, belong to .ttimNaster and r ~!clwnisia (St(irmer and Bcrb!'\ITllln, Cri no
long-ranged g~:·nera wh1ch lllve persisted ft>r a long 11)76) (Fig. ll.H). P.llaec
period ot gcnlogtc:~l time Tht:rc arc three: or four Bnu~h starfish bt!d, wh1,h tnbntt
h.1v~: long merited ,mcnoon Of these, one 111 tf1e l 'nmp
L1.1~ of DoNet, 1\ fitl1 of ophmro1ds (two ~pc:(le<) r.1rl'ly
Subclass Ophiuroide a (Fig. 9 . 33c-e) hut there are no other fauna. ~t"duuentolo~tlaltn Cri1
teria (Goldring ~nd Stcvemo11, 1972) make it pl:un the pn
The mmt JJStmctive fc:.lturt: of ophmro1d~. namely that these ophiuroids were ~mothered and raptdh l'r,J anc
their thtn, snake-like anm. h,l\ been .111 unportant buned by a th1cJ... cloud of \llty ~cdi.mem. Modcr \\hich
tacror in the1r great 'uccc~'· 'mce they me ~muous opluuroich cannot escape from 'edJmcm more: th.n One o
movement' of the ann~ !ur locomotion. 5 cm thick, and the\~: Lia\\JC 'Pl'cimens probJbl> ~n IJ.J5) i
AJ] tht' VISCerJ ;trC lOllt<l111Cd in the l CIHI"<ll disc, died in the1r life position. uf trin,
and .tU thJt he witlun the an ns .~re: the greatly Th~: fumom Ashgilli:m st.1rti\h bed at Gtrv.m m . lnte
enlarged vertebrae, hu111ologou' with the fi.1sed Scotland ha\ an ~:xtrcmcly nch f:Juna of ,t.trli~h. c olum
ambulacral p!Jtcs of J~teroid\. The-se have speci.U tnlobttcs, brad11opods, mollmc' :111d the early l'dU· bd(lre
arricubtmg hmgcs so tilJt the ann C. very Acx1hle-. A nOldS .i rtlec//11111~. Ectt'llcrlllllll~ and rotllllriil. from rh, o-awl o
sheath of pl.nes <.urround~ the vertcbr.l~:', and the scd 1ment ins1de the echinoid tc't' Jnd other cntc:na The
\!Tong mu,dc~ that move the arm~ .tre found in the Jt has been conduded that the f.mna wa~ shifrrd theta) f
-.pace between. Thou~h tht: carhest oph1uroids are some distanct· and rapidly bum·d following mrbu- The rh
found m rhe sanH.' beds J..\ rhc tlr.-t somastcro1ds, the lence m a shallow-m:uinc env1ronmenr. Other ca~o ngul pi.
latter w~:re probably the1r lllCC:\tOr~. Thl .lmbulacra wluch have been coru.iderc:d c ontim1 rhar onh J~; Ae:o..•blc
enlarged to t<.)ml vertebrae ..1nJ the rad1al w.HtT ves- re~ult of 'catastrophic' cond1t10m .ue surti,h ltkd} l.'r.1l am
sel becanu: enclmed by the growth of .1 protective: to be preserved whole. Thu~ .1 thm lower Jura~~j, .llld :l
ventral shield. Then.· .uc: two order~: tht· bed .1t Gmlind 111 Gernuu~ ha~ ,1 rich f.uuu of maJrcpt
Oph.iurid.1 (Ord -R.ec ). 111 whtch the ,1m1~ can srarfi~h and also of other eclunodcmH (Ro,t'nkr.mz gonads
bend only m the hor11ont.ll pbne .•md the: Eurya!Jc I l)7J; Seilacher t'l al., 19XS) There is, hOWI.'\'tr ann~.
(Carb.-Rec), in vvhJCh the arms ~:an movl' in all very little other fauna apart trom oysteN killed bv Smce
planes; the l.lttcr arc oti~. n clu nbmg tonm. the same smoth~:ring event. The shelly borrom talyx) i!
Ophium1d~ arc inmu.·mdy \w.:ccssful 111 modern f.wna was overwheLmed by .111 mflux of scdumnt pl.ne wl
Subphylum Crinozoa 291
and, whereJ~ the mobile pectinid .md limid bivalves
LOuld \Wtm upwards to t•,capl', the fixed oysters
could mH and the cchinodenm, with the1r .tmbu-
lacral '>)'\tetm ~:huked b~r fine \cdiment, had no
chanu~ of \Urvival. but were pre,t•rvecl whole and as
a 'sdcctcd' fauna. Such taphomuJnc ~electivity is
normal ut other beds m which ethlllmlc:rm' arc very
wdl preserved (Bren and E1. J...LTl, 19R2).
Cnnowam are prumnvdy \talked cdunoderm~

('pelm.<rozoan~') \\,th long Jflll' Jnd nomully l.lck
comph:x rc~p1ratory ~tructure~ I ht· comatulid
mno1d~ have. however, lmt thetr \talks and become
;econd,mly tree.
Class Crinoidea
CnnmJ, ,1re very dl\'c~e

and irnportanr in
PJio~eozoJc 6unas. and thl·lrrem.tm' have con-
tnbuu:d mbsuntiall) to P.tl.teulotc: limestones.
Compktl' tnnotd~ (e.g. F1g. IJ••'\4), however, arc
rJrdy pre~crvcd.
Crinoid~ an: le\\ abundJnt th.m they were, bur at
the prcwnt thcy Jre rcpn:sentcd by 25 stalk ed gen-
m and by mme 9() genera of umt.tlkcd com,ltulids.
11 1\hKh arc the domin:mt group nf modem oceans.
n Ont' ot these. rhc modem frt•t·-hvtng lmedon (Fig.
ill 9.35, 1s t.tken as a type exampk \bowing the basics
,,f tnnOtd morpholog).
;n 1utrdon ha~ a stalk in lt'> e.1rly life, formed of
h, columnar p lates, but tt \OOll bre.lks free of it Figure 9.34 Features of crinoid morphology shown by
ai.:- ":lort 1t is fully grown ,md j, then free to ~w1m or Ordovicion Dictenocrinus. (o) os redrawn from Bother in Troite
rJwl over the sea Roar de Po/eontologie, 1953; (b) reconstru<:ted os a rheophile
hc
na f'he body cousists of J globuLn pi.lted cup (the
cd theca) from which long plated anliS (brachia) ari5e. stem . Above thts IS a ring of five basal plates with
lll- lhr tht"LJ has two parts: ,t lower region of thick ano ther ring of five radial plates abuvc tl. Attached
'>t"5
n~d plates, pcntamerali) \ymmctnc.ll. and a do m ed to the r.tdwl plates are the brachial plates of the
Le; a de~..ibk roof (tegmen) wJth .1 Lt'ntr,JI mouth and lar- lower p.lrt of the arms. These arms \ttbdtvulc almost
.dy trll anu,. lmtdc the theca is a 'p1rally cwmed gut lmmedt,ttdy so that there are ten .mm m all, each
!>SiC 111J J circumoral ring ,1~ in cchinoiJ,, but rhe armed \Vtth many pinnulcs comtruueu of pinnu-
of !:Udreporitt· 1s replaced by Lili.ttcd funnels. T he lar plates. In .1111edon and other l onutultds the
UlZ , •n.ld, .lrt· borne nor \.\. lthm tht• theca but on the ngid plate~ of the calyx arc very )mJil anJ contain
vcr. illh. o nly a chambered part of the coelom A number of
I by ~mre there 1s no stem tht· b.t~L' of the cup (or Aexible cini arucubte on the centro-dursal pbtc
l0111 .J)'x) ts m.1dc of a single, l.trgc (centro-dorsal) and c:~ n be used for temporJry fixation or locomo-
1Ctlt IJtt' whJrh is morphologically tile top plate of rhe tion of thc hvmg cnnoid.
292 Echinoderms
M
(c) tube foot tood groove I he
trom
later
coe10m1r: spaces lllllSC
rebt'
l·ocr-
Edmt
l:ttcr
there
porat
l!JH~
brachoal platP. \CC Ill
(o~~oclo)
pl·no
re.JCh
r:tdiat
ncula
(e) rare.
(b) rrtl>Ulh
(Jiso
I ... rem
~Ill CC
brachoal fa cat
... mens
Bnti~
\llg'gC
other
Tr
Aren
hdd
gut \\ hicl
Th
HeXJI
C Hfl onl)
\V her
Figure 9.35 Antedon, a Recent comotvlid crinoid: (a) living specimen in dorsal view (x 1); (b) vertical section through calyx. (c) llltO t
crou·section through arm; (d) larval specimen still ottoched to stalk; (e) lateral view of proximal port of a living specimen with Aexiblt lna
cirri below [(b) Redrawn from Claus, Textbook of Zoology, 1884; (c) redrawn from Treatise on Invertebrate Poleantology, Port UI
other
Rl'Cl'
H)IOcr
mere)
The arm~ .trc tkxtblc, havtn~ articulated brachial the mouth by ,t c iHa ry tnuc u s t r act along the food Ag
ossidt:\ .llld two 'cgmelltrd ~Lraml' of muscle with groove. The pinnular food grooves :~11 join wtth mno
w hid1 the arm un 1110vc in .my direction (Fig. thmc ofl11e br:Khi:J, which in tum unite '>O that ti\'t' Amc1
9.35c). rrom the liruunmaJ ring d radiJJ water primary groove\ cro<.' the tegmen Jnd enter thr mode
ve~sd tum Jov.n <:alh .trl11, whid1, ,1\ 111 \tarftsb bur mourh. When di,curlx·d. the tube feet can rermt :Kqtu
by comr.l~l wttlt crhtnmd~. hes uu~idc the plate~. A into the food brroovc. and J\ they do \O \mall ralcttl(' the ~t
double ro\\ of tube ft'l'l anw\ trmn tlm ve~sd, with cover plates (la ppcts). normally held erect. come de ten
a medtan tood groove m b~:tween. When the dmm like rrapdoor, to protect them. t:rinm
crin01d i\ t~eding. dw pinnulc' \\ith thc'e tube feet Anu:don ha\ only I 0 .mm, but other cotrunthds tet·dll
art: extended and, bl·ing !>ticky. catch organic dem- may have a\ man> a' 200 The dt>~pcr-\\Jter 'pectcs UllliSt
tus ami smJ.ll or~.u1i'm' which art.' then carried to ~eem on the whulc to h,l\l fc \\er ami\. lOOn
Main groups of crinoids
The oldest kn0\\11 cnnmu ,, L;illlll<lftlcnti/IS (M. Cam.)
from the llurge'~ SI1Jk. but 1t is not vet)' sinular to
l1tcr cnnmds .md onl)' tdt•nnf"it·d .tS ~uch because of its
uruscnal anus and tlw \\'c:ll-prC',ervcd tube teet. Other
rdatc.'d eclunodt·nm, mduding the blao;t070Jn Clas~
Eocnnoide.1, .m~ ,,J,<) C.tmbnan . ' rhe rdaoon~lup of
liilmll1f<>mmH to tr' t )rdovician de,ct·ndants of 40 Ma
Luer i~ not undcl"\tood. but m the c.1rly Ordovician
there was .1 wcat r.ldt.Htcm nt L'.trh crinmd~. contem-
por.meous wlth that of other L'dnnodcnm (Donovan.
J98Ha) pmbabl~ wnn:nrr.ttnl in tropiCal seaway~. lr
~eem~ th.n the Ordovtt 1.111 w.t' .m 'c'\-pcrimcntal
pcnod' 111 cnnotd mo'l•holngy .tnd Ordovtnan faunas
reached a h1gh kvd nf di\'L'J').tt) in \UCCl'~o;ivc: adaptive
rndiatJom. Sine<: rht· t rinnid ~k.dt·ton. however, di~ar
riculatcs nptdly .tlrl'r tk.tth, l'nmpktl' spccimem are
rare, The 'rudy ol the nng-,h<tpn1 columnar plates
(also known .ts rohnnn.tl"\ or o"idc') of which the
~tem i~ rompmc.:d i~ hdpful 111 .tpprcciJ.ting diversity, Figure 9,36 Cupulocrinus (Ord,), os preserved in the rock
'mce tht'} .trc 1:1r Jllllrt' t011llllnn tl1Jn compktc speci- (x 2). (Drown from o photograph by Romsbottom , 1961 )
mens or t up' (Don m .m. I'IHh. I 'JHlJ). Thm in the
Bnti~h OrdovKi.tn tnmml t:wn.l, <:olunmal diver:.ity
~~~~L'Srs .1 vancry nf \Pl'l"ll'' 't'veral tUllL~ greater than
vthel"\vi\c rccogniLt:d. Crinotd da~~lficatton •~ lUrrently m a ~tatc of flu:x,
Traditionally. rhe L'.trltL''t tml'' crinoids (e.g. the and Sunms and )cva,topulo ( I'N") .15cnbc: many
Arcnig Dnrdn•m11w .md Cupult•tnllm) have been Palacozmc group~ to the ArttCulJt.l; ~omc of these
held to belong w \ubd."' lr1.1dunat.1 (Fig. 9.36), likcw1se h,1d Acxtblc. 11\11\Clllar .mm. Tht·se authors
which havt· rigid c.tiH.tl pl.ttC\. recogmzc the C'amerat.l '" morpholo~;~call) dl\ru1Ct
Thl'rc arc t\\ o otht·r Pal,tl'ozoic ~ubcla~se~: the from the bcgmmng. hut rht•y rt·plan· the Inadunata
FleXJbtha (Fig. 9.4.\l), in '" luch Lalycal plate~ <lre b} the two subd.mcs D"pand.1 (P.tlacoz01c) ,md
only loo~dy umtt·d ..md the Camerata (hg. 9.37). Cladid.1 (Ord. Rct .) They comidc:r the FleXJbilia
where dll' proxirn.tl .mn m'tclc~ ,ue incorporated and the Arncubta to hl' infrad::t"c' of thl· l.tttcr, ,md
mto the tht·c.t. seck the a rlCC\try of thl· Arlicul,ltJ .tmongst the law
hudun.Jtc' JUSt t rmscd uno the TnassJC; the Pabt•ozoic dadid~.
others bcc.tmc t'Xlltlt't 111 the;· Pemti::tn. A possible
ltecc:nr surv1vor of lhc tnadunates n11ght be Palaeozoic crinoids
Hy(lcrimts , an Alll.lrlltc form, though thts could M ost, though n or ;11l, Pabeozoit crinoid~ have
merely be an aberrant arttcui,He. ~tems, w hich in the r:t~e of \Omt· Lower
Again. tr.tditton.llly . .tll Me~ozoiL to Recent Ca rbon i femu~ forms arl' uC great kngth. They gen-
mnmd~ ,uc pbn:d 111 a lourth subclass. the erally have l.lrgcr c.1lycal platt'\ than A11tcdo11, and a
Articulata. in wh1ch the .mm .are very flexible. Most third circlet of phtc~ (tht• infcabasals) may be pn:-
modem cnnotth .lrt' umt.tlked comatulids; the sent under the basak In dc~uibing cnnoids. the
ll'qUJ\Jtton of br.tdll.ll At·xthlltt) and liberation from morpholob'Y of rhc cal) x j, u'ually rcprC'~cntcd by an
the stem were undoubted!) of gt'l'Jt tmport.mce in 'exploded diagram' (Ftg. <J.37c). and the plate~
Jerennining thL' evoluuon.ll) potenual of this arc distinguished symbohcalJy as brachiab (BrBr),
mnotd group. allcw.tn~ an .1ctivc st·arch for good radials (RR), basal\ (UU) and mfrabasals (lBB)
s 'ixdmg h't"oumh. ThL' ltvin~ \talked ~penes are Where there arc only RR and BB prt~cnt the
,, unusually \mall. mhabiting dl'pths greater than crinoid is monocyclic, dicyclic cnnoub h.1ve lBB
Ill.! m and having \tem' r.trdy cxceedtng 0. 75 m. J\ well.
294 Echinoderms
I facets
Actmocrimtes
brachtal
(b)
facets
(e) Cactocrinus
(f)
Figun
Fig 9
tbrod
nol cc
Porl L
m;ldl
h.td
gl'IIC
Scyphocrinites er,lll)
the l.
gene I
Figure 9 37 (o)-(d) Actinocnnites (l Corb.), o monocydic comerole crinoid with brochiol plates incorporated into the large col)'l< Nt
(a! lateral interoreol view with brochial facets (x 1), (b) legminol (oral) view; (c) exploded plate diagram, (d) specimen with brocl11o dt~Uil
sti I attoched; (e) broken specimen ol the related Codocrinus, showing subtegminal enclosed food grooves leading from the brochiol
facets to the mouth; (f) Scx_phocrimtes (Dev.), o large comerote, with pinnulars incorporated into the calyx (;... 0.3). [(a)-(d) Largely kno\\
based on Woods, 1946; (f) redrawn from Moore ef ol., 1953.) in,u:l u
lh
~'I Lh
lnadunate crinoids (RA). wltluu the radial circlet, and there 1~ usu:tlly pl.m:,
The lnadunat.1 (L. Onl.-Tria'>.) j, a very large sub- an anal (X) plate in the brachi.1l nrdet (Ftg. 9.3Rc). uuprl
class wnh m ore than ::?OO 'pccit~. ln.tdunares are The va~t majomy of inadunates arc d1cyclic, .1 tcw tn:.c .1
probabh the mmr pnnutsw of all rnnoid groups are monocydic. Radial plate' ma) be 'com- fOod
(Fig\ 11 36, Y.3~). ha\'ing " ri~>id c.llyx \Vith the pounded', 1.e. djvided traruvcl"\cl)' ulto infraradials The
br:H.:htal~ free or loosely ronneLted above rhe radials. (lRR) and supr aradials (SSR), aud there .ue radl- '[ n/)(1/
The mouth is below thl tl'gntcn, the food anal eqUivalents (IRA and SR.A). In ~omc Jn anal have
grooves above it. The .mm m.1y ur may not have tu be emerge~ from above the r.tdl.lnals; thi~ is Paul,
pinnulcs. Commonly the pentamcr,tl ~ymmctry of greatly elongated in geuera such JS the Ordovimn In
the c:~lyx is disrupted by an extrJ plate, the radianal Dictetwcrhws (Fig. 9.3~) and Demlr~ICI'i/111.~. where irIS ridges
(11)
IBE Cd)
(b) (c) Cupulocrinus
mad repone
pores
(e) Cysrhocrln/res (0 umsenal
f"ogvre 9.38 Plate diagrams; (a) monocydic calyx; (b) dicydic calyx; (c) Cupulocrinus (Ord.), dicydic calyx with an anal tube (as in
f'9. 9.36); (d) lateral view of a dicydic calyx (188, infrabasals; BB, basals; RR, radials, in black; RA, radianal; X, X·plate; 1BrBr, prim·
aYochs; 28rBr, secundibrachs). (e) tegmen of crinoid Cyathocrinites; (f) evolution from a uniserial through an intermediate to a bise-
~condition tn the brachia. ((c) Redrawn from RamsbaHom, 1961; (d) redrawn from Bather in Treatise on Invertebrate Poleontology,
Pan U (f) modi Red from Moore el o/, 1953 )
nude of dun, folded pl.ltl''· Th~: t:.lrlil-st inadunate~ work. where rhe pl.ltc\ JOin there are thin cm:ulJr
lud an dong.ul·d 'rr.llght lllp, but m many later area' of the calyx beanng folded calouc membrann
gcncrJ the calyx bi.'C.lllll.' tiattn1t:J .md expanded lat- (goniospires). Each gomosp1re co1msts of three sets
tnll¥ FurthemlOrt', tlw .tlmO\L veruc.tl direction of of folds, o ne for each of the ad_1.1cent plates. The
the d1chotomom arnl\ Ill tht: earlv maJunatcs wa>
gmerally rq~IJct:d h'r a 111orc tlmwr-hh· crown.
Numerous orh~:r l.'volutllJil,lry trl·nds have been
d11tmgtmh~:d, .md nuny 'peca.Jlized torms are
known. Peltdl>CriiiW (h~. 1J.Jl)) is .1 tjny Silurian
madunatc rc~cmblmg .1 num1tc palm tree.
Here the stem IS short, the calr-. small, and each
of rhe five br.lcl11.l fused llllO :1 kaf-likc wvided
r!Jtc, w1th a sencs t>f d1t hotomom {ood grooves
·mpressed upon m >luiace. H)•/l(l(ysrrs has only three
~e anm, the other rwo being repre,cnted only by
t«xl grooves extending down tht· s1des of the cup.
Th~ OrdovlCJan P l1rocn111d.le (e.g. Pomcmms and
Tnht.lftlptmt.(; Ftg. IJ.40a,b) are 1 ~Lrangc &mily which
bYe bt·en pbccd 111 the inadunlte' (Keshng and
rJul, 1968). Figvre 9.39 Petolocnnvs (Sil.), a small inadunate, having the
brachia welded into Rat plates with food grooves. The cross-sec·
In rhc'c rhe calyx i' \trengrhcned by tluckened lion shows the cover plates. (Redrawn from Tosch, Po/eobio/ogy
!l.igcs hnkmg the pl.ltl.' centre\ \\1 tthin rh1s frame- of the Invertebrates, 1973 )
296 Echinoderms
lbl ~uborders Monub.tthriJa .md Dtplnbarhnd.l. I'C~pr~

tivcly. The carhe~t camerat<..'S, \UCh a\ the I O\\CT
Ordovioan Rrtt'tltn/IUS, luve m.llly fe.tturc' r('nnnb-
n:nr of m,HlunJte,, imp!) ing .1 rcbnt~mlup A UUJ
c nnoid such ;~s .-ittinotrillitr.s c\htbito. the.: h.I~ICS of
t amerate strurtun: dear!}, 111 the cxplodnl pl.11c du-
h'T.Im and in tl1e lateral and oral views. The fl'b'lltrn

j, l..trge and dome-shaped .111J has ,, mbtt·ntral .mm
Tht "
•.lrl
Anm anse from the cucul.1r b rachial facets , \dud Ill a<
.trl larcr.tl proJccnons from the calyx Th~ m 10} Son
Figure 9 40 Porocrinus (Ord.). with folded membrane struc- interbrachlaJ p la tes (ll:hllr) and hrJchi.1l plarC$ tOll
tures at the plate junctions: la) lateral view of calyx (x 4 which are UICOrporated intu the c,tlyx makl· .1 brg~
approx ); (b) section through fo ded membrane (Redrawn from pol)
Kesling ond Paul. 1968.) nh'ld theca. h
In borh monobathral .llld (hplobathr.tl l nnot~
there are evolunona.ry trends par.1Ud \\ 1th thme- of "·"·\
pre\
goruo\ptn:~ prohabl) h.1d .1 rc~prratory funcuon. but the Flcxtbthn. Thus in the early g<..·nera \UCh :h tl nn t
of grcatc\l llllcn:st i~ their ~im1larity to or her rypes ot dtplobathrJI Rl!leacrimiS, thC' 1nterbrachul p!Jtt'> arc ,lfl'
folded nH.:Illbram· >trurtun..· in both <..·y~roids and ~mall and irregular. lym~ 111 dcpre~sn.l 1011~~ lurtl
blast01ds. '"Inch aros<..' qllltC indcpcndenth. Some berwccn the arms. In the desc.endanb of RtluV11t1u th:.\i
tunction.tl .mJ other c OINdcranom on ~uch struc- rhq. have b~:comc large. re~ular polygonal ~11d Ieo. and •
tures arc ~1\'cn ldtcr. Tlw b't known undunate\ .1re deprc,scd. Furrhennorc, the- .mal and interhrJchl I that
TnasSll. The \\ dJ-k11UV. 11 Et~.niiiiS /i/fr/c11111is which pl.lte~ tend in the later gencr.t to move upward~ and \llll'l
occun. m thl..' Gemun Mmdwlblk 1s om· of the~e . nut of the cup, pcnnitting J perfect radial ') mmctl) h1e.ll
wtth thc radi<ll~ .tll in cont.JCt. Ma.ny other .lpJMenr ,lbk
Flexible crinoids evoluuonal) trcnd' have hl en reviewed {Smun th.m
The.. FleXJbtlsa (M Ord U. Perm.; e.g . f>rl'f,l\wn·,,s: 1990, 1991 ), but have to be mtcrpr<..tt•d "rth cau- l'l3 h
F1g. 9 4Jr) arc less mm•c-rous .md <..hversc than tion. An evident trend common to allrnnotd' •~ rl, m lo
madunatcs. They .m· ,Ill dicyclic wlth three mcrea~ing prevalence of forms with pinnul.Hc arnt\. ~~ dkt
11tfrabasab, one smaller th.m the tlther two. The Tlus may, however. rcpn·,c·nt only an 1m rc.l\t 1 h.lVl'
lower bra<..lllab 1re mcorpur.tretl mto tlw cup bur are 'vanance' (i.e. the number of types \\ tth pmnullt c 1rn
not ng:.dly JOilled so tlut the ''hole cup 1s Aexiblc arnlS) foUO\\cJ by the cxtmnion of all grmtp' \\1th .thout
The momh :md food ~'Tuovc~ arc on tlu. upper 'ur- non-pmutdate anns, rather than ,1 tntl' trend \u
tace ot thl..' tegmen. Tht·rc i\ .1 radi.tn,tl .mtl an .m.tl 1mpclled by functional nc:cc..·~~tty. St'iror
plate. 'iomc evolution,Lt)' trends, such .1~ thl..' relative There are many unu~ual camcr.ttl.'~. '!'he there
wtdcnmg of the calyx .tml the tcndcnq l'l>r rhc lower Dcvonian <it yplwainiu·s is J r.1cher e>..-rremc fom1111 .llld 1
part of rh~ ·'"''' to droop over UH! cup .ITL parallel wluch not only the lowu brach1ah but .1lro the mt lu\
Wtth thmt• in the inadun.Ht~ . lower pinnular plates have become mcorporated I•II(!Ul
1nto a very large tht:ca. M;tny gem·ra, 'uch 31 tbuuc
Camerote crinoids (Fig. 9.37) Harralldeocrinr~s, developed Luriou~ly clowntumcd rl'giOI
Subclas~ C.unerara (Ord M. Pcnn.) j, dtt' brgl..''t of anns, droopmg over the cup. The ann' hcrL' are .1rc c1
all th1. Pal.tt•ozotc ~rtoup' In dm h'Tuup, whit h very broad, with the plmtulc~ bent ovcr to mw 111 1~01
Jppcarcd \Olllt' 4 Ma .ttrc:r the first in.tdunJtes, the mcdtally ~o that a trough w,1, formed. pr<..-sumabl) I he: I
cup 1~ ngtd, the radi.m.tl p!Jtc 1s ab~t>lll .md <>ome of ,Ktmg as some kmd of food groove. The melcll:ruu• north'
the lower bradual pl.Jt~s .trc mcurpor.1ted into dtt· t1ds arc a group of CamcrJtn Ill which th~ lTOWI\1\ 111~ 6·,
calyx. Tht• mouth anJ foc•d grooves l1e below the n~d .md bt:ars five arms with five ~et~ of p.uallcl hc·t.mt
tegmen, and m some r.lSC\ (c .g. Cwt•mnus) the tlun pmnules between, leadm~ toward~ thl l'C'Iltrc potent
subtegtnm.d tube' lml...mg the bracltl.t With the Cowen (19tH) dre\\ the rc·nurkably dO\l' .malogy lflllOlt
mouth hJvc been prcwrvc:d and c.m be d1~secreJ b1.·twccn tlm .1rrangement and the idLal l.tvout l'i a \uc.
out. The ,mlh .tlway' h.wt· pmnuks. 1 here arc both h.mana plam.HJon! The problcn1s arc the s.tnll', lw- rhL M
monocyclic and d1cvchr genera dJ"1fied in the vcsting an evenly distributed p.trticul.ttL' fi..'\Ollt'IC (ftp;. l)
Subphylum Crinozoo 297
and Its dc:hwry to a Ct'ntr.ll p1 tJCt:~\mg point ""i.th

mirumal expenditure of cm·r~ry. As Cowcn put it,
'cost-bendit analy'"' 'hould pia\ a pan in fimc-
nonal smd1e).
Mesozoic to recent crinoids: articulates

The c:.u·lscst .lnitUI.tte t. nnotds are T na,,ic, and the
car~· gcner,1 bc.1r 1 COINtkrablc resemblance to
madun.1tc,, from wlu~ h tht.T probably evolved.
Some .lrnrul.ttc\, however. could posstbly have
COffil' fmm the ( le..xJb.JJa; tht.• nrdt.•r would thus be
rolyphylcnc.
In artlcui.Ite\ tht.• r.1lyx t\ rdativdy reduced: ftve (a) Encrinus
ba1lb, rivc mtr.Ib.ts.ll~ .llld tlvt.· r.11:iJah usually being
prc1cnt. The lllOuth .111d t(H)d grooves .lrt· ex-posed
on thl· \urt:lce nf tht.· tlt.•xJbk te~mt.·n. but brachials
m· never II)Corpor.ttt.•d 111 tht.• cup. Articulates arc
hutlwr 1nd mmt import.lntly dmlllgmshcd by the
rlrx1biliry n( tht·ir lnng .lflll'. \\hich can move up
~nd down, coil up or t\\l~t .1bour. lt is tlus character
tlut ha, given rlwm .1 grear cvnluuonary advantage.
mcc tht.· m,lll)' gcnt•r.l o( (. )rdn Cnmatuhd.l which
:ul"Jk frcL' from tht·ir \UIJ...\ 111 the early \tagt'' arc
Wit actively to lllll\'t' ,InHitld lll) the \eJ floor rather
115, than hcing tixt.•tl. Of nwtkm cmHlld\ only 25 gen- (b) Saccocoma
IU- m havt• ,ulk,, .mc.l thc,t• .Jrt• r.trdy mor~ than 0 75
•he m long, nmtr.t\ting With 'mnt· of the gJJnt 3 m Figure 9 41 (a) Encnnus {Trios.), on advanced inodunole
ns. ;ulk~d ilmm of t'.1rl1cr tlllll',. Tht· 'tt'ni\ frequently ( '0.75); (b) Soccocomo (Jur.), a po55ibly planktonic articulate
( 5) (Redrawn from M.oore et of., 1953 I
Ill bm: dongatc rim cruptmg from nodt·~ and dtc'>c
!lC om arc muscul.tr, C11.1blin~ tilt' rrinotd tu 'walk
11th lOO\It • on the m. • long feather) anm .md wrious fl.tttcned lappcts,
end Stalked i\ocnJuJ, 'uch ,1, Pt llltunnitrs and bemg cxten,ions of the proximal ann pl.1tc!>.
Sttr,'mlllll arc quite .tbund.Inl in the Jura•"ic. and lr is the mo~t numcrou~ m.Icroto"il m the said
rhe llltre .m: many nllli-Crct.tteou' tom!!>. Mcsozoic lithographic lune1>tollc (Chapter 12) .•md bct.·amc of
1 111 o~nJ l'ari.1ry gcncr.1 .trt• ~cncrall} lung-ranged and m pcndi,1r morphology h.t\ tr.tdttimtJIJy been con-
the mdud~ .1 numbL'I Llf miLrtmiorphlt forms, e.g. sidered a:. a freL·-m imming form, floJting mouth
ated Lu~wi.nri1111s. I'ht· umtalkt•d torn,ltulicb arc very downwards with the lappeb 111tre.l~ing the \Uri:Kc
l •• , rund.1ut tmh1y. thcrt· bt:iug 90 gcncr.1. In certain area and thm retarding stnking. Milsom (1994),
rned rc~om of the world, Antarttic.t m particular, there however, has reinterpreted Snccocouw .ts .1 bcnthic
are it' t'ndemil tonl.lllllid fJtlllJ~ wh1ch have evolved fonn, lying pa~s1vely on tht' !>ea floor with the anm
ne et o l~olatJon ~llllt' the bq~innmg of the Plt:istocene. archmg upwards from thctr nudpoint and fonmng a
ubi) The Ant.lfl ne gencr.1 now M~t'lll to be spreading horizontal collening bowl tor tccding, or med to
Tllll- northwMds .tlong rht· lll.lllY 'ubm.1rine riJges radiat- intercept current~. SamlW/1111 wa' probably able to
.Vll 1' g fiom the Antlrt.tic t'ontint·nt. Comatulids first crawl and, smce there wa~ littk other bcntho~. it
.r:illcl ~lll1C common in tht• Juras\ic; evidently the seem_s to have been Jll llpportuni,t, flourishing in
·urre. trntHI f(lr movcmem inlwrcnt m the unstalked vast numbers dunn~ periods wht·n the wa floor
aloro :nnoad\ anm \\",1 ' rc.1hn·d ~:.1rly. was favourable ro htc. largt.·, plated. stcm.lcs~ genera
t ot .• Sucn:"ivc conuruhd ~t·ner.l continued through (e.g. ,\/arsupitt·s .111J Uiut.lWI/IIl) arl' wdl-known
. har- e Mt·,ozoic and r tTtl,l!!"· Tlw jUIJ\SIC Sa(((l{llllltl Crcraceom tos,ils. Most living gt·ncLJ arc unknown
)tJrce. : ').41 b) from cht.• ',)olnhnt\.·n Lunestone ha5 very as tossik
298 Echinoderms
Ecology of crinoids shallow current-swept water; and nuke me of rh~ R

Modern umr.lll..cd c nnmch live .lt .tll depths from currents in teedmg. (tho
subhrtorJl ro .lbY''·ll, st.llkcd torrm arc normally Dunng the d.ty tndn•tduab may htde 111 c.tvincs. The•
found only bdo\\' ltlllm. Crinoids .m.· uncommon but come out .ll mght to tccd. chmbing wtth th~tr fccdt
in shalJow w.ucr .md tl·nd m be most abundant in clfri onto .1 htgh po~ttron on .1 rock or sea gras-; .md pdrtt
rather in.tcn·,~rbk n.:giom. \U that until recently lit- unfurling thetr arms. Tht• :1011\ spread out tn a ver- rator
tle wa~ known .1bmn their lite and tccding habits. tical filtration fan. wtth the pol.tr axts honzom.tl. uatu1
Fir;t-hand diving ob,crvatiom have illuminated the aboral ~tdc t:Kmg thl· current .md thc pmnul~ mud
nuny ummpl'C:h:d .1d.1ptatiom and stimulated func- spread our to form a gr.tting. The tube fect Jre Jmh
tional ronsideratiom on .111c:iem fonns. Ho-.vever. extended from thl·~t· to form a fine ftltering net. bow
knowkdgc i' srill vcrv limited .•md most observa- Even with a current 'perd of only 2 cm ,-t J surtacc Som1
tions ~o t:1r h.wc bel'n nudl.' l.trgd) on comatulids. of SOD cm~ can filter 1 I ' 1• Such .1 sy\tem i' rcm.trk- JlllOt
Prc~I.'IH-day comatuhds divide into two quite dis- ably etlictent, maktng mc c)f cxmmg currents. tunc
tinct feeding tYpl'S rhcophilic (n1rrent seeking) thmg
and rhcophobic (llment .wo1dmg). IZ.heophilic -----+ current direction simpl
crinoid' (Fig~ 1J.34b. 1J.42, 1>.43.t) live 111 relatively (a) cornp
proba
ter he
Me
proba
of life
(Breu
crate
do SOl
~~~rinp
carbo1
Burlu
<ltes (£
mtcrp
hut At
(c:) the cu
I I of a v
I I
I I tanng
... ... away I
mostc
these c
cicnt t
thl' cu
tn.: knt
\CJ.
So m
.1nd we
bent at
n~mg a
ucal tilt
Mo~
Figure 9 42 Cenocnnus oslenus lmne; a Recent rheophilic Figure 9.43 (a), (b) Stalked rheophihc cnnoids with calyces t.hc cro
crinoid living at depth of 200-300 m of the north coast of arranged to oct as vertical ~ltrotion Ions. (o) with no lih From lhe
aJrrent (b) with lih, (c) rheophobic cnnoid (based 011
tJon. Ft
Jamaica The arms are of rod1us 20-30 cm and form a vertical
~ltration fen . (Photograph reproduced by courtesy of Dr D.B. Protaxocrinus) with heod ocllng as a horizontal bowl. [(o), (b) havc p
.V.Ocurda) modi~ed from Breimer and Webster, 1975 ) rhcoph
Rhcophob•c ainmd~ live m current-free waters rheophobes with the crown actmg a~ a horizontal
'though mmr decp-w.ncr l nn01d\ are rheophilic). collectmg bowl (F1g. Y.43c), though ~omc may have
They rdv cnurd), ,\, f.1r a~ 1~ known, on gravtty been macrophagou' IJke modern ha~ket star\.
~dmg; onh through .1 rommuom ram of detrital Despite the tlcxibtlity of thc1r \tt:m\, no living
punclc~ cJn rhey obt.un enough to e.u. In the labo- erinoids have any mmde' rutllllllg down the col-
ro~tOT)' Aurrd.m beh.l"c' J\ J rheophobe, though it 1s unu1 and it t\ most unlikely that any toml onl'S did
111rurally .1 rhcoph•lc Rht·ophobe' (rig 9.4Jb) are eHher (Donovan, 19HHb, 191-N). The only tibn.'S
much Jcs, common. They ht· on the bonom. the extendmg through the column are non-contracule
II'Jru. out,prl'.td to lonn .1 horizontal coUe cting ligaments of collagen. rhc\C, however, are mutable
bowl, cJptunng orbr.lnll pJmdt'\ t:tlling onto it. and connected to the nervous \Y'>tem. and they are
\•Jntc rheophohe\ h\e 111 dct·per w,lter;. ~o that a fatr involved in conrroll1ng the pmture of the colunm.
Jmount of dernral marcn.1l h.1, .KcumuiJted by the This IS also aided by the form of the articulatwn
time tt Tl'.lrhe' rhcm. b11t they do not consume any- between adjaccnr columnah (l;1g. lJ. H).
:hing hkt: the yu.mnry that rheoph1le\ do. They are Synostosul arnculations, With thetr plane wrfaces,
11111ply cxploitin~ .1 dilfcrt•nt ~ourec of food, .md in allow .tlmost no tlex1bthty of the column. They are
LOillpensation Ll11.'1r gmwth .md met.tbolic rates arc rare and confined to culy Palaeozmc crinoids.
rrobably lc~s. Abyssal t rinoids (and others) may ftl- Symplcxial articuL1nom. with tht•tr radial ridges,
tcr horizontal tun t'lll~. would prevent tw•~tmg of the column .md hence
Most l<mil t rinoiJ~. ~t.llkt:d or unstalked, were snapping, but would alJow 'rocking' between adja-
prob.1bly rheoph1ks or rht•ophohes: hence their mode cent columnals to allow rhe ~tem to bt?nd. They
1f hie n1.1y bt: inlt'f11rt·tt•J from thl'ir morphology would also ,1ct ·" gt11des to allow the columnals to
Br~·imcr, 1<J6lJ; tlrci111a and W t:bster, I CJ75). Cam- return to thctr ongrnal posrnon after mch flcxl.lrc.
mtc lllllOJd, havt: long .mu ven flt·xiblc stcm'i, as Synarthrial articul.mon~. wlth tht.·tr central fulcra!
do ~omc uudun.ltcs. ~omctmH."i uinoid t3unas con- ndgcs, allow ~uh~t:mnal rockmg bc~vccn each
mnug lar~d) of tJilll'J,Ilt:' m:nrr m lugh-encrgy columnal and hence ,, Vl'ry fic\Jblc \tcm.
wbonJ.lt' 'equerKe,, In the Lower Carboniferous A number of VCf) diVCI"il' rnnmd taunas arc
Burlinglon Limt:,Lollt' of MI\\Otm. the lllatly camer- known, ruamly tTom thc USA. and can be tmcr-
l!C\ (fanui.Jt•s A cum)cnmd.lt:, tl.ltocrinidac, etc.) are pretcd t:cologtcJIJy. The Crawfordwlile LimL"itont·
Ultl'rpretcd lar~d) '" theoplttle~. The ~tc1m arc strong of Indiana 1~ full of tht• \talk~ and crowm of
but lkxrblc, .1nd tht·lr Hcx1b1hty mrre;t~e~ towards inadunacc .111d At:xtble trlllOtth .md \\J~ dcpo,ited m
'Dl' cup. 1 he r m\HI 'mrid thu., ,1\\ume the attitude quiet and poorly aerated w,ltt'r' On morphologtcal
~i a vcrncal filtr,mon t:m. w1th the aboral surface grounds mo~t of the cnnoid, were probabl)
~ung the currt•m .1nd tht· long anJ.l tube facing rheophobes (though they rmght po'>~tbly have been
1way from lt ( l1~ 1J.J4b. 'JA 1a). ~ince the arms of
mo~t camcratcs .\rt' brwri,1l wnh veT) nuny pinnulcs,
0•m
thl·sc crino1d~ would set'lll to luve been lughly dft-
l~nt filrcrcrs, str.1rn111~ otltood partJdes borne by
thr currents whit h, .1~ ~~ clear from the sediments,
1rr known to hav(• swept the tlurlinp;ton Li.mcsronc
$,1.
Some omeratt: stalb h.wc cirri on one side only

md were prob.1bly tixl'd tu the 'ea Aoor in a recum-
1\(m attitude, w1rh only the crown and upper stalk
[29 . ·. ... . .
.
rismg abovt• rhe ,t:,l tlnm, .I~Jlll bemg hdd as a ver- .

~al filtering tan ... ,_.__.....
Mo~t flt'XIbk c nnwds haVt.' 'hort rigid stalks, and
es he crown could not have bent mro a verncal posi- Figure 9 44 Different kinds of articulating geomelries in
1e
'100. Furthcnnorc, tht· .mm of the Flcxtbtha do not crinoid columnols (ossides). wrth longitudinal sections through
)0 columns below· (a) synostosial orticulotion, (b) symplexial articu-
b) ::.~re pmnulc,, 'o .111 the evJdt·ncc 1s ag;umt a lation; (c) synarthrial articulation (Redrawn from Donovan,
ill·ophilic mode of ht~. Tht') .1re best mterpreted as l988b.)
300 Echinoderms
able to Crl'.llt' theu own tunenb by undulatory pscudoplankton. hence the pendent life of 'uLh
movemcnl\ 1li rhl' .11111$. The.- verv ditfnent aspects obligate lom1s as these. They are not uncommon!)
of the Burl111gton .md C 1.1wfixdsvilk crinoid fauna found with lo\slltzed dnftwood, and fi.mher e\1-
ddinitd) pomt to .1 thtli:rc..·m nolob': and feedjng dcnce tor tlm ltfc llJbtt come\ from a 'inglc beddtng
type. plane. now ext ,lV:Itcd ,md di\pl.lyed in th(
Perh 'P' rh~· larg~.·~t .111d mo't d1\'crse fauna of Tiibmgen Museum. 111 ' ' hich he \Oilll' 50 crino1d
ec..hmo,lcrrm 1-tHI\\ 11 1\ 111 thc Mtddle Ordovtcian speClmcm. each wuh a cmwn abour RO cm across ;trc,
Uronudc..· f-ormation uf Okl1hnma. ln thi\ extraorch- and a <;tern up m 1 c; m in length. They arc prc-,cm:d rl'n
narilv rid1 dep1"1t ~omc..· hI gem rl bdongmg to Wlth the crowm fjcc down in h),H:k etL'tinic: ,haJes. the
13 dJ\sc..·~ h.t\'l" bc..•c..·n de,nihc..·d (Spnnklc. 1982). One interpret:ltlnn of the~e h that thq lived a \am
They hved 111 ;~ \"Jriet) of \\ .mn-water shelf deposits pscudoplankronir lite. hanb.;.ng downwards from a "va~
with depth~ rangin~ tiu111 ' ln 75 m and were com- Ao:ltlng dnftwood log whtrh slowly became water- LfO'
lllOilC\1 m em·lrc..llllllent\ w11h a mbstratc of lime logged and ~ani-. o,o that the crowm touched down lllll
mud ;~nd skdet.d ddm\, t:\pet t.11ly where these were first. fi)llowed by the \tl:lll~ whkh ~ettkd in loop~ a) CJSC
smaU bf) o:zo,111 nHntnd' provtdmg \UJtabk attach- the log drifred slightly before coming to rest. In a no
ment ~lte~. Altlwu~h the f.HlnJ 1~ cnnmd-dommated, Seirocri1111S the ba\al pan of the ~tcm is the more tll'x-
so m,my other cd11noderm grnups arc present that ible, by contrast with the norm. This too i~ in accor-
tlm '' surely the ne.. hc~t of .111 cchmoderm fitllnas dance witlt a psemloplanktonit mode of life. for the
kllo\nl from .111y pill c nr rune. seem need\ to be flexible at the base to wtthst.tnd
PJbeozmt c..nn01J ~ardc..·m m.1y luvc been strati- ~torms ycr rigid near the calyx to maintain control of
fied like ,, tmpil .tl jungle, p.utiuonmg rc~ourcc~ b)' the inverted poHur~. Spt>dc..:s of Pelllcurinitt's and (
feedmg on d•tlhc..·nt tood n c:hffL·rL'nt kvck St•tromrws sho\\ buk lllurphologJcal change through c
f-m,tl .uncul.nc..·, cm hknvtw bl· mtcrprcred a~ ti.tn~:, and indeed have remarkable longevtt) .1s com-
rheophlle~ ur rhenphnbl'\, llowc..·vcr. m the case pared Wlth t ontl'mpOt.meous benclue t nnotd~. ThtS 'Cy
of the M c-,n:zou· cnnmd t'Jnuly Pcntacriniridae confonns well to an unc-rprctauon a~ pseudoplank- bla-;
(Pc/ltcJmtlitn md St'~r,wi1111s) dtfferl'nt mterpretanvc mnk - a ca~e of stabtlizing selectton for a very spe- tru•
critena havt· hl'l•n ll't·d. l hl',c..' arc..· \'CIJ large. oti:en cialized habn.tt. One ~oup of Pc:mamm1s 'Petiei, l-
spect.lc..ul.lr rnm)ld,. olttu tnund completely pn;- incidemally \\ith a :.horter, densely ctrrate ~tem. ml) Ord
sen·cd 111 bltck slult·, (t'.tt \unm'. 19S9). There arc have departed fiom the p~cudoplanktomc h.1btt 111 Eun
t\\0 po"ihk \llg_J.!;t \tiOll\ .1\ tl1 thl'JT mode of life. have become be nth". It
One i' rh.u rhn wcrt• bemhtt, but their long stems wit!
gave the l-.1lyx .md mm tht• .1dv.mtage of clcvatwn, Formation of crinoidallimestones Str,lt
so that thl' fccdlll~ c..'W\\ 11 ' ' J\ ,tbk to file er from a Many Palaeozotc tnnOJds 5eem to have lived tn plau
reg10n of tlw w.Ht'r nnt l xplottcd b)' other organ- 'crinmd gardens', i.e-. 111 dumps or patche~ i\obtcd upp•
ISm~. A long-\t.tlkl'd tnmnd could have gained lift from other :;uth .m~a\ .md 111 wluch dtversit)' m::ty be hcs 1
from the CUt1'l'tH, provttkd tb.tt it w.t~ able to orien- high or low. Other crinotds mhab1tcd muddier and
tate the panbohc.. tan .1t the correct angle (Fig. water conditions. the
9.43b) The whole 'Y'ltnl c.1n be ltkened to a kite After death, crinotds usu:~Uy bn:Jk up throu~h each
terhert•d hy •• u.m!. But uniJke J ktte the crinoid decay, disaruculation or the JCtH)Il Clf scavenge~ smaJ
could prob,tbly rc..~ui.Ht' tb own degree of lift by The: specific gravity of the vanou~ components tir:~t po111
::tltenn~ tlw curvaturt· of thl' anm, the angle of the mcrea.ses, owing to the remov.1l of mvestmg tis~uc, (bra•
fan or rlw ,tltltllc..k pf the..· pumule~. Given certain cl1en decrca~es whilqt the stereopb~m (msuc pcnc- to id:
condition,, tlw crown (l1uld rt\l or 'mk in rhc water traring the \tereom cavtucs) i~ replaced by ~c..:J water to tl
to exphut fi.lod-11d1 lt·H·k Where Isolated column,1h :Jrc tound they have been The·
Au altern.tnve. curr~..ntly nil.> re popular view transported hy rollmg .111d m:ty haVl' come from zont
(~e1bc..her rr ,,/, 19M{, ':>tmtm. 1YH9. Wtgnall and some dt~tam e Common!) grt•at thtckm•,,c, t>f watc
~tmms, I 990) t~ that thc..'\t ~t.mt cnnmds tived a limestone arc found, formed of ~hort or long lcnb'l~ A
p'cudopl.lllkh'JUL hti.•. han~m~ dtm'll\\'drd'\ from a of stem, Isolated mm le~ or c.1lyc.1l plJtl''i. In tht:' angu
floaun~ log and rhereby feeding from the ~urtace Carbomferom limc~conc..· of England crino1dal the I
pl.mkwn . 0H·r~. W\Hhn!.!; " ,t problem for <;uch debns of tht\ kind c.ln often tonn the bulk of the alon 1
Subphylum Blastozo a 301
rO<k. The~e rocks .trc more.: m lcs!> .Jutochth onou' sm,tll shr known a\ th1.· hydropor eo, pwb.tbl) tlll'
Jnd rcprc~enr the Jebri~ l)f cmH)Id gardens L'ntr;)JlCI.' to thl' w.ltcr-va~c llbr ~y,tem .111d dural-
whne tht· material "' ,1\ bum·d prior to complete tt•n.;ric of cystoid,, is \L't nl'ar th\: pemtllllll '. All
Jmntq.,'Tanon pl.ttc' except thl.''e of the pt.>nstomt tre dcmdy nlv-
When ont·ntate d, 'hurt qt'lll lengths .trc tounu in l'rc.:d with tvvin p1.•rtorat1on,, surround ed by a ni,~.·d
abuud.mn·; they ,..,,.t'rt' roUcd by the current mto an rim .md r~mmJSl\.'llt of the port• p.11r' of t'dllnmth
1n:~ of .l\:cumuh non ..111J till' Jm:cnnn of thl.' cur- Thc'e .m· known J\ diplopor es. The} prnb.1bly In!
rent that movl.'d rhem c.1n tlllrnully he told from to ,111 external 'p.1puiJ', uncakiticd .111d 1\.'\l.'lllbln•g
th~ir oricntat11m. Uno••ent. ttt•d dl.'bri\ found on the Jll cloug.1ted tube ft)Ot extended inro th~.· 'll!Totntd -
same bcddmg plann •pparl.'lltl) bn.nne ~ntck .md lng .''a tu for n:,ptrano n Th1' !,:l'llll' lu~ bl'l'll 'tud-
\\J.\ not movt·d by tht• l'Urrellt. 1 he r.tre case'> when icd in det.ul by Uockdic (IIJH4).
cruwn and \tl'm are presc.:n cJ n·pn.·,ent almmt
imn11.'d1ate buri1l bet'l1rt' decompu sttwn or, as 111 the Structural characteristics
mt: of Sdnlfrllllls, dep11SltlOn 111 quil't and pmstbly ~u1ne ry\tuids ltkc Haph•spltactcll/1.\ were tixcd b)
anoxiC watcn their baw., to thl.' sea Aoor or to :1 'hell: other' h.1c.l
\tems "lut h hJvl' been (Kra~wn.tll) pre\l rn·d \'. 1th
the holdla~c intarr. but tbc'L' stt.'nl~ were .llw.1yo; ~hnn
,111d ne\ L'r rcarhl·d more than a k\\ centimet re' 1n
length. M 1nv of rhe OrdovJCian l'\\Wido, lud tht>cll'
L0111post•d of numerom 1rregul.u plate\ (c.~. the
Classes Diplopo rito and Rhomb ifero: dtplopori te Ari.,IOC)'Sfitel and the rhombifcr.111
cystoids [ :c fllllt'~Jllt•lfnlt'S).
fhe foud groove\ m cy\tOid~ were prot~:cred by
'Cy~t01ds' .m: hetemgc neou' group of Pal ~u.ozoic
.1 lOVer pl,ttl.'\. The~e food groove' 111a) bt 'hnrt a\ 111
bU'tozoam .•1U of \\h1ch h.we re~p1rarory 'pore "mo.-ystr~ (Fig. 9 45) or long. bur onJy 111 F.nnil)
>trucmres' traveNng the pl.He~ of the theca. (. heirocnns dac: (Cl:t~s Rhomhifc :ra) •., thcr1.' .my doc-
1lnplospluuwnis (Cl,l\s Dtploporita: Ftg 9AS) 1~ an umentt:d trend to lengthcn mg th1.· .unbul.ll·r.l \\ ith
OrdoVlCIJil lvsroid k.nown mamly from nnrrhcm nme.
Europ<.'. The t ltnOU\ OrdovJCI.IIl genu\ Asteroltlasllls h.h .1
lt !lJs an ovoid thn .1 tc~~ than 30 mm in height ~t.tr-s luped system of food groove,, very 'imil.tr 111
with .1 fiar ba't' by \\ hsl h it w l' .macht.'d to tht 'ub- :!ppea.r.lnlc to the Jrnbul.1n.1 ofbJ.,,toi ds. Ur.ll"hink'
l:mum. Tht.' theca h.t, 1:\.\ o csrdets ot polyguna1 arose l.tter.illy hom thl'\C. and the drplopore s
pl.lte\, the lower cirdl'r of seven pbtt.'' (LL) ,md the were re\tnctcd to the intl.'r.1mbul.11. ral regtom alone.
upper of sevt.'n perioral plates (POO) Above: these lt has been \llg!;t:\ted that blasto11.h descemk d from
tiC'!~ th<.' pemtome , con,tmn ed of fi Vt' pen oral pl.ltcs ry~toids through •!sremhlctsrlls. ·r lm 1s unlikdy. hm\-
and h.wing ,lt tts cemre the ~hr-like mouth From cver; it is more probably ;J c::asc oft onveq~c:nt l'Volu-
the- mouth r.1d1ate hvl.' tood )..'l'Oo\·e~ or ambui.JCra, non. Br.ll hiole' .trc rardv found m q 'tollh. lllll
.llh of whKh has \t.'\ cnl br.1nchc~ tt.'nmnau ng in where they are known they an: .tlways ,hurt and
.mall smd<; (ambula cral facets) wh1ch wen: the ~lender. rhe gonopor e (for the shedding of gelllt.d
oomts of att,tclune nl of tht• fnod-g.1t henng .mm products) .md thc hydrnpor l' are u,uaJl) Ul\tmct but
1-·-achsoles) ilrach10k ' arc rarely prc~crvcd in cy~ \Ometime s combsncd .
tOid\ but t~re very sinular 111 ~tmcture and function
• the arm<; of crinotd~. though small and sunplc. Pore structures IFig. 9.46)
Th~~ were probabl} ~pread nut in ,, k.Jnd of hori- The porl structures of rysr01ds, wl11d1 art• nl
lontal rolletttng bowl, extr,JCting food from the very many kinm, provide a tinn bas1~ tor d.ml-
· 1ter and p.ls,mg tt to tlw mouth v1.1 the ambulacr a. litarion (Paul. I 968. llJ72). l>1plopore,, a\ in
A \\cU-dcti ned 'anJl pyra111sd' offi\~: or more tn- HaplosfJ/Uirrol/rs, .Ire p.11rs of perpendiCtll,u t<m.ds
Jilb'lll.tr anal plates " set at the junnion of twu of through the the~:::~. carh opening in .111 cxlt'n1.1l
'•l POO plates. The .1naJ pi.He\ could hinge open dl·prcs~ton (the peripore ). Othl'r kinth of port'
mg rheir bases and dose again after t•vacuatto n. A 'tntcture ,\re based on u-~haped tube, of vanOll\
302 Echinoderms
ClASS DIPLOPORITA
Superfemlly
Superfemlly Spheeronltlde Glyptosphaerltlda
hydropore
br.tdrrdl
facRts
pens tome Frgur

drploporus
show
FamriV Sphaeronrudae Famrly Holocyshlldae drplopores

(all have drploporesl (all h.1ve humatrporesl on plate~ ~md
H•plo¥h••ronll Holocystts
lXtCI
mouth
Superfamily L· xtc
Arlstoc:ystltlde brachral facet
~ht-s
(all have brachral facets
drploport<S) T'hcl
\\)J.....,I-"'""0~..,..,- mouth under
the r
cover plates .lre p
p.trt:s,
Anstoc:ytit•s
gonopore T,
boh1mlcus lOrd ) tu re
S inocystis oral regron tu re~
l'Xtcr
CLASS RHOMBIFERA with
ORDER DICHOPORITA
(all have pectrnrrhombs pores
or crvptorhombs) prc\ll
h .IVC
Superfamily thm 1
Glyptoc:ystitida
the 11 ·
n,d. E
Pleurocystittls ORDE=R FISTULIPORITA pore~:
(all have humaurhombs)
mnni:
woult
oral proaecuon '""' back,
t OJ) 1
~
litem ,
cxchar
brachrolP facet
CJH.Io t
torcig~
Echino$ph.,.rit•s pl.1tc-.
hod1e~
pure J
Sr.uroc:ystrs
stnJctu
liable 1
were 1
nf'iurt
Figure 9.45 Cystoid morphology ond clossificolion according to Poul' s system, showing the high taxonomic volue of pore strvctvres Dip!
(Diogroms bosed moinly on Kesling et of. in Treatise on Invertebrate Paleontology, PartS.)
ll umer•
Subphylum Bla stozoa 303
dt·~cribed. ThL·~c arc 'imple p:tired wbcs cro\'llng
the plate perpcndu.:ulJ rly and leading to unc;tloli~..·d
extemal papulae An altern.ltlve kmd of ~tructurc
~~ the huma ripo r e, in wh1ch the paired perpemht-
ular rubes an: linked by ,1 complex of exothL'c.1l
tubes. These Jre either Rmh with rhe surtac~..· ur
Ol ~ upy raised lnunp,. Dtploponte' lt ke Hllft,, }'~fi~
(£ 1g. 9.45) h.JVc their pl.tte~ replete w1th ,uch
Figure 9 46 A pectinirhomb, crossing plate boundaries ond hunutipores .
V!Owing the folded membrane (Based on Paul, 1968 ) Rhombtferan s nuy have exo- or t:ndothecaJ pore
~tmcmres wh1e.: h J.re ah\i;l)'' ,lfranged in p.u.1Jid 'eh
With :t rhomb-shapc d contour. Lach rhomb
crm~es a plate boundary. Pcctinirhom bs are htghly
hmb known .ts t hecal canals \\Inch open lo rhe org:.tmzed tllllt~ cort~tructcd of P••r•llk·l d1chopore~ .
external or tmem.tl 'urf\Ce through th ecal pores. There arc nomullv on!)' rwo to four m any one
Fxtemal pore'> .tre round .md 'lllllplc. sieve-like or cy\tOJd. Each thchopore nuy open 111 a singk ,IJt
sbt-shaped, mtem.tl pnres arc always ~1mple. (conjunct ) (Ptg. Y.46) or h.we the central p.trt
Thecal L,111JI' .tre alw.tys tubular .md may tr:Jver'e root(:d over (disjunct) to form a <,ep.lr.lte entranu:
•he thee,\ pcrpendtcul.l rly or tangcntt.tUv (diplopores and cx1r. Dichoporcs nuy be joined Literally (con -
\re p.tired thecal ~..· anJh tonning U-pairs in tht· soft flu ent) fonmng a singk folded membrane. '-lurh
: lrtS). pt•rnmrhomh s luve a rat~cd cxtcnul nm and nuy bt·
Two quttc di,tint'l lu11ction,11 rypcs of pon.' :.Lruc- l<H~unct or diSJunct. l f di,junct, they have :1 thtrk-
IUn: can ht· distingut\hL'J . Endothecal pore \lruc- enl.'d bar rumHng along the hue of the 'uture c.lrry-
rores (dichopores) h td c.mal, tlur ran bdO\\ the tng .1 r.used ndge on one "de onl). Th" ha~ bl'l'll
\tema) \llr{Jce of till' tht'CJ ,ltld COI11111llllll ,Jtedl'xpl.nncd J\ .1 mechamm• for preventin~ the n·cy-
wtth the ~urrountlill~ ~~.·a water through C'-ler mtl cling of currenr~. The nli,try action of the cellulJr
rorcs If slol water ",,, pumped through the ~..mal, membrane Jimng the dtt hopore~ \\ould produn· a
rresumabl) by nli,t, rL',plratory exchange could one-way current. Small parudes droppmg through
h•ve been d'fc.-cted b) ga'>t'ou' ddn1s11m through the to the base of the d~ehopor~ would be lt~ht enough
:hm calntlcd will of tht• thec.tl r.uul whiCh bathed to be Jispo~ed of by the ciliary current. Endorlwc1l
!he intern:tl body thuds Rcspir,ttion wa' thm mter- cryptorho mbs are made of p.lr.tUcl dichopnrt' S
n.J. By contrast. exothecal pnrt srmcturc' (ti~tuli wJlhout tht· r.tt~ed extcmal stnH. cure of p~..·c
rvrc:~) lud mternal pore' and ,111 external canal tuurhombs. They have "eve-hke mcurrcnt and
running outside tht> theta. Thu, rhc body thuds ~tntph: excurrent pores. Humatirho rubs an: m.ltlc
would have to be brou~ht through thL' c.111al and of L'xothccnl fi~tnliporc~ likewhc p:tralld .md
.-k down again \\.tthm the thn.l, ~o that rc~ptra ,mJngcd in rhomhs.
ltllV c:xdunge wa' outMde thl.' dwc.1 Thc~c rwo
u:ernanvc methods l)f facihLHmg re~ptr:nory ClassiBcation
txchang~..· have thc1r ,ldhmt:tge' .1nd cti~aJv.mr.1ges. Many ta.xononllC scheme~ h.we been propo~ed for
!.udothcL.ll 'tructurcs were liable to chok1ng by tlm complex and d1fficulr group. l he schcmt: uf
re•gn parncks. but they n1.1y ha\'( slm. ''eve Paul (1972) adopted here (hg. 9.45) dttfer.. \01111..'-
:.tte~ or (lthcr dcvtres ro keep 'uch toretgn '\ hat from that of Ke~hng in th<. Trcnti.IC (Ill
'IJdie, out A sievt· muaUy prutt·ct' the 111l1JI.lnt llll'ertebmtt P.drolltology. Paul ha~ gJVeu high-kvd
're J.lone. cJublmg 1l to be iuenti tied. Fxothce:tl t.1xonomic value to tht· pore stnu.. ture' In Cla"
wctures did nor luve thL~ ptubkm but were D1ploporita there are only d•plopores KJttered
uH'r
Jble to abra,ton or hrc.1kage. anJ m any l,tst• they the rhecal ~urf:lCe, manv ro :t pbte, other than
111
iiCre muc:h k~s eftiCJL'llt 111 tenm of rel.Htvc .1reas 1~.unily Holocysutid ae where [here ,1rc hum~Hipu•
cs
l iUrface l'Xchangc. imt~·ad. l n CIJ~s Rhombi fer:.~ rhe members of Ordt•r
Dtplopontc cy,tmd' ~uch a' Jf,IJIIosphaeflltlis h.wc DJChoporit.l aU have pl·crimrhomb~ or Cl)ptu-
:::merous cxotht·c.tl diploport.''• a' already rhombs, wh1ch an: used m dtv1de the order further
304 Echinoderms
intn -.upcrtjmilk,, I hunnirhntllb' lllJr,tctcrize the radial ridges org.mi2'ed in rhomb\. lt live~ 111 cold
mher ,,nk·r. tht• Ft,tuhp~.•r.lt.l. circumpolar watt'!'\, whtrh would ho~ve had a t:urh·
htgh o:-..·ygen roncenrr.mon Pnre \tnacture~ c)ccur-
Ecology ring 111 1ts dc,ccndam'>. m nthn cy,roid~ ••md mdccd
Mmr l '\tmd, "er~.· '""ik .md ti..·d on ,m,ill org.um: in all uther ~·t hinudenns wtth t>Llhor,Jte re,ptrJtor;
partid~.·, U\lll~ tlll'll hr.• ~. hiuks. T hc'c: br.Kh.iuk~ .tre structures, .trc t'tmnd predonunantly 111 shallow,
..,ddom ti.l~\tllll.'d but Ju tlllt nurm.1ll)' sc:c:m to ha\'e rropitJI sea' Nt·.trly 1ll Ord<wtu.m occurrence- of
bc:tn \Cl)' l.•rg~.·. Jud~m~ by the \lte uf the ambu- pore bl'Jrer' .tre berwe~.·n .30°N .md .311°S of the
lacnl fan:h 111 tlw Hul1.1l} 'uud.1e. h1.1\\ ever. the brJ- pabeo-t·quawr. In \uch wann, ,lullow. tmpa:.tl ~ea5
l."h tok' m.t\ 111 thh t.unily h.t\'1.' been of fau ox-ygen tt:mwm .1rc VCI)' lo\\ dunng rh~.· mght. and
dtmcii,H>m Pu"tbl} th~.· do,c:Jy sp.H.ed brachiole'i of · it •~ sugge,r~.·d rlur the pnrt· ~trunurc' \wrc wdl
'uch l!;l'rtc:r.t ·'' l /,,p/tl_,p/llrrtwri~ fonm:d .1 food trap .tdaprcd tor lOpmg with rhi,. f'urthennore. trom
hkc tl1.1t nf \01111.' mmkm Ph~.1romJca. evidence .w:nl.tbk .tt pre\t'nt tt set·ms that the mor.:
Cy,toid~ 1-(t.'IICt.tlly livl.'d in ~lullo\\ '"ater~ . .tnd efliCJcnr pore stntl m re~ (pet tmtrhombs) orcur 111
1:\'tdcnth thplopontt.'' Wl.."rt: .lmon~st the flt'\t colo- genera ltving wtthm I5°N .mtl I '1°S of the cyuatnr;
ntzcr.., of .m.•,,, 111 whtdt 'edtment depo~ttton beg:m the ks~ cffirtcll! onc'> h.t\'l' J \\tdn dt~tributwn.
.tgain ,tltcr .1 br~.otk. Cy'>tOtlh h,td some tolerance of
su~pt•mkd ,cJuncnt. turH.:etvJbly exothel."al pore
,u·uuurc' u1uld h.n 1..' bt•t.•n .td,tptcd to sediment- Class Blastaideo (Figs 9.47, 9 . 48)
laden WJtcr. Whcrt• ~.ondtuum were particular!\
l~vom .tbk, q ~t1.11d~ liH:d to!);cthcr in clump'>, .md Bl;v;rotds (Sil. Pl·ntt.) an· .111 t'xtincr dJ~' of small,
ou;.l\tunall) thl.''l N;cmbl.lge~ \\'l.'re preserved in lite penumer.tlly synll11l'tnc.tl 'pelm.ttozo.lm' dlJr.Ktcr-
po,lllon. izcd by mtem,J) re~parawry stntcturc~ known a1
Some cy't(.lld gl'IH:r.t arc stnkm~ly modified. The hydrospires. An mdsvadu.JI blastmd had J ,hnrt
rhumbtfc:r.tn fJftlllt'I)'Stilt'.' (lsg ') 4:>) h.1\ m JSym- fixed \tem sunnounrt·J by .1 c.tl) x tiom whirh <Ho,,
melntJI. fl.tttt'llt:d th~.·~.. \\ l11d1 h.!' lJrgc trregular a crown of brJrhinlt''· Thl· 'tl.'m .md brachlllks '31'r
pLtte' wsd1 pt•t tintdwmb' on <'I ne 'tdt• and ·mlJil rJrely preserved, an d the nujont\ of ~pcuc:~ art Figu.
Rextble pi.Hc' on tht• otlwr 'I ht• t\\'u brJchJOlcs are kno\\11 from dtt' c.tlyx .tlon~.·. The plating dillen hon
lar~t· .md ftll'\\ 1rd pWJt.'l'tm~: rhe ,111m ., located by from thar of cnnoln.m... In ~nme gem:ra there .uc
the St(k of tht• ~Ct'lll. whtch '' wry tle:-oblc and cxrr.1 and t.:omplc:-. pl.tte,, but only the ,t.mdud
tJpcnng .md l.tck' .1 h<11dt:m. Plnrrt~ry5titt'.' probably pLning 'tmcwre is ~i"en herl.." Some 80 ~encra are bee,
by on the ,c,t tlonr \\'Jth the Acxibk p.m of the known. th.lt
thet.:,l down (P.wl 19117) Thl· 'tcm morphology Pt'lllfl'mitc'.' l omc-. Imm the l'vh\\t,stppun of North ·r
would ,IIlO\\ undul.mn~ nlll\ cmcnc\ \\ htch would Ameri1.a. Near!.,. 70 'pcue~ bdong to tht' g("IIU\ lOll
penntt tor\VJrd propul,tou, .1nd thus the wholl' P. symmctricrt1 (Fig. IJ .47) trom the Lowrr I he
morpholog) of flf,•~trcl()'.llitt'S ~l"C:lll\ to be "PeCt:tli7ed C<trboniferom of lllinoi' h<~s .1 bud-sh.tpcd l\llp sid(
for .1 rnobtle. bt•ntlllt ltfe A~ 'lll'h tt bear' J rcm:uk- with tive pet.tloid ·.unbul.!l'ra' nHuung from tbr ked
Jblc ~upertiL·t,ll rt'\l'mbl.tncc: to \untl.trly ~pecializcd ~ummit lO t wu-thirJ~ or tht• way to the bJse. EJch ,111 J
' honnlozoam' (e.g. ( ;,llfrrtrllt'ry>tis; Ftg. 9.49c) . which ambulatnnn i~ indl.'nll:d by ,\ vertlc~l mt:dun \\ hl
quHe mdependt•mly .ll qum:d \lmiiJr charJctcr; groove. There .trc thtn· ba~.1l pl;~tcs (UU). Abow o l c~
through convcrgl'nt t•volutton. them arc five radial platt:s (R.R.), deeply mdcntt'd by lrO'
Wh) dtd L}''tol<l' nu:d \Uch dah0ratc pore 'truc- the Jmbui.K ra, .llld ~11rmountmg the raduls Jre tivr llh.'l
ntrc' for r~.·sp•ratwn when the cnnOJd,, which out- \ntJII rhomb-\h.tpeJ 111te1 ambu!.H.ral deltoid plates llltll
Lt,tcd thl'lll, ~Clt I)Jl pt•rli..·nly wdl Without them? (t.). Bclov. l'Jth ambulo~uum IJ~.~ J long. spw- tr.tu
Po~ul ( llJ7(,) h,t\ prcst·ntn.l .1 rt',l\Onabk hypothc'i~ ~hapcd lancet pla te, l.tr~el} covered by the snuU ht'Cl
ahom £111, h.ht'd on thl kno\vn p.tl.teo~t·ographtc.tl platt.'' of tht· tlwd j..'Toove,. llltC
d1,mhunon of ry,tmth ( )n~.· of tlw carlie\t known At the summit tht're as a central. \tJr-,haptJ tl·on
C\'\tOith, tht• Tr~.·m.tdnl\111 \l.tcWc)'$1../1,1 (which \\'J\ mouth, .md ~urroundtng it .trc tive large spiracles tcm
onguully thought to h~ an et>ninoad). lJck, .my each \C( at thl \Uillllllt or a deltOid pbte. Om: ·I wall
pore stntnurc,. rhuugh lt' pl.uc' ,lrt.• thin Jnd h.tvc thc:\c {the anispiraclc) 1~ larger than the othen surt:
Subphylum Blostozoo 305
~f< hocet plate
(1) (c)
hydrosplrP folds
f19ure 9.47 Pentremile) a Lower Carboniferous spiroculote blostoid (a) lateral view (x 2); (b) adoral view (x 2); (c) transverse sec-
!ioo with hydrospires (K 2); (d) oblique, three dimensional section showing brochioles and hydrosptre system (x 10).
bc•.auw tt cont.nucd tlw tntemal anm. ~uggl',ttng to the many hydrospire pores by mdivtduJl thm
tlw thl' ~pirack~ were ~om~: kmd of outkt ~y,rem. Lubc~. A p.1ir ofhydrosptn·s tTom adj::acent .unbul.tcra
fhe amhubcr.1l .md ~ub.lmbuiJcral Hnu.turc i~ l.Ollllt'cts with a smgle sp1r.1clc. The ~ubdtn"ol1 of
complex .md i'i b~o:\1 ~fl1dted by making tlun \t'CCiom th~.. tnlll'r f.lce of the hydrmpire 111to four ,malkr
r I 1 .,l ,how dut the l.mcer plate bt:.lri two lateral p.u.tlld rubes results frum rhe mtoldmg of tht.• caki-
1ide plates a~snn,ttl·d wuh rhc elemt"nl\ of both the ticd tlll'mbrane of whKh tr is constructed.
ln·Jmg .md rl',pir.Jtory 'Y'tenK Alnng eadl edge of
;a m .unbulacmm r;an .1 ~mgk Ime of br.t( htolt.-~ winch, Diversity and function of hydrospires
1 1~hl:11 broken off. ldi dto;tmct fJccrs. l'hesc brachi- H ydro~pircs may. a~ in Orbrtre111ires (C.ub.; Ftg.
ub lcJ to food b'TOOVl''i with bt'icrial ll.lVt:r plates ') .4Hc). t on'i't merely of srraight u11divtckd tube~.
(t•>"in~ the ambul.icnun .llld joinm~ \\a rh the u,u.tlh. however, .1~ tn Peurrrmires. the hydmsptrc<.
uteJtan ti>Od b'TOO\'t'' \\ hKh ran \'t'"rtll ally ru the an.: co11volured (Fig. IJ.47e,d). tonmng a hy dro-
mouth. Fcl'ding \\ .1s prob.tbly hke th.tt ot utnoid'i, spiraculum, \\luch n1.1y h:we fi-om two to 'even
trm:' of the watcr-v.1'l'llltr ,yqem h.n ing recently fuld\. The hydrospm· pores were probably lined
l-cc:u dNoven.:d. l'hl· hydropon. w.l\ probablv w tth cthJ, and their heM prcsumJbly crc:-Jtcd a uni-
Ultc:rn.ll. Berwcen .ldJ.In•nt hrachwlt's 1s ,1 narrow ~ltt dtrt.'ctwnal current, drawing water in thrnugh the
tinm '" hiLh a pore k Hh down to the hvdrmptrc \)'S- pun.:\, passing it .1long the hydrospir.Kuhnn .111d
s. tem. bch o[ tltl. p.tirl'd hydwspire-. is simply .1 thm- exltJ.ling tt through the ~p•r.1clc~ fhe toldlllg wouiJ
:;~f wallc:J, ngtd, cakitll.·d rubt.'. wirh a COI1\'oluted 11111er unduubtcdl} aiiO\\ .1 h"'l'.lfCt '>Urlace tor re,ptrJtnry
:rs rurt"Jcc:, lying bdm\ tht.• .tmbulacmm .md connected c:-x~.:h.mgc.
306 Echinoderms
(b) Tin
t'Xtc
r-
by
lnd
s!de plate the1
h,w,
lancet plate
crall
Ion(
w it!
a nu~
lOll•
Coc/aster
tng
Id) 1•1
E,
l3las
t:tun
'>Oill
the 1
Ill
Orln
~tud·
J()5C
Orbitremites 111 er
T
Figure 9 48 (aHc) Codoster a Carboniferous fissiculote blastoid with exposed hydrospire folds: (a) adoral v1ew (y. 3), (b) la~ and
view, (c) ambulacro w1th hydrosp1res 1n section, (d). (e) Orbitremites, a lower Carboniferous spiroculote blastoid· (d) in lateral v1ew Amt
(...: 3); (e) hydrospires in scchon
h.td
Ill ,
HO\\ etlictenr wen: the\e \tntcturt·s? It '' a fairly comcal theca With .1 flatrened upper surt:1ce. The l..Jrb
\lmpk n1.1ttcr m calcui.He mput .tnd output flow upper edges of the radtal plates fonn :1 nng around an: r.
r.ues by tnC.l\Urtn~ rro,\-\ClttOnal .ueas of the the honzontal ddtmd, rad1al :md lancet plate~ . The of po
hydro,pin. pore\ tnd the \pirade\. In Clobtlblasws, anu~ IS off-centre. at the junrnon of two radiJls and Yery
where a complctl' ~tudy through ontogeny has been a deltoid. Parallel with the edge' of the lancet plart..., hcm1
nude (M.tulrd.t, 19(l5b). the potennal outflow rate arc the uppl'r edge~ of till· hydrmp1rl' folds, graded Pt•m
1n ,tn ,tdult could havt· been ~1x ttmcs the inflow in stzc :~way from the l.utect so th.tt thl· hydrosp1rr
rate. Thu' ,H .111 utfiow vdoetty of 11.1 mm s- 1, the system ha~ a h.tlf- rhomb-shaped outltnc. There Jic
water volurnt• could bl' changed completely in I 00 s only .1 few n:duced hydrospire foldl> between the
in :m adult .md 411' m ,1 juvt•tu k. In Claboblastus dif- anus and the atljact·nt !.meets. The side plates Jir
fcrenti.d growth gradlt'nt~ seem to relate to hydro- h1rge and almo't cover the lann:t plates, as shown in
dynamiC .md n.•t•dtng dricicncy by controlling its Fig. 9 .4Hc. It ~~ presumed that the hydrospires fun<:-
external fom1. tioned in .1 m.mncr hro.t<.lly simiJJr to that of the The
spir.tculatt· blastoids, but the system was probabl) ously
Classification and evolution of blastoids lcs~ effictcnt. older
Bla,toilh .tro~e 1n the S1lun.t11 trom an undetermined Tht· Fis~iculat.t are wtdcspread and la\ted until the p erhJ
mce~tor. T\\O ordl'r' an.· dtstmgUJshed: the P l'nnian. Such genera J' Oroplww11m (e.g. Macunil, lu1o"
Fimcubt.l and <;ptr.Kulat.l The Ft~~iculata have 1965a) are well-known ~.:ompom·nts of the ally l
open hydrmp1rc fokb. whcre.t' the Spiraculata (e.g. Mississippian reef knoll and carbonate ~belt f.tUru. Jdfer
Pet~trrmrtcs, Cft,[,(,/Jltlstll.\, Orbttrt'mrtrs) have ~piracles. Bizarre genera include .-lstrocn11m (L. Carb.) which them
f t,,tcul.ttt• bb,tmds arc apparently more primi- has one aberrant ambul.tcntm .md a displaced anu" cc brat
me. C!lci.Hta (Fi~. 9.4~a-c). tor example. has .1 sub- Amongst -,c,eral odd genera from tlte Pennian ol chore
Subphyfum Ho malozoa, o therwise ca lcichordates 307
Timor ~~ 71t.lii11Hlfrl/Jhtwts. whtch hJ.~ inultcmcly chordate~ v.rith echinodenn al)inmcs which ~ave
··xtcnded bnu·ts .md :1 rounded base. ri~t· tO other chordates, three t1n1cs indepc:ndcnrly,
FtssicuLm.-., l ould h:~w ~;wen rise to ~ptr;ll ulatc' by Ios~ of the calc1tc skeleton Udferies, \9H6. 11J97;
" the latcr.ll b'Towrh of the lancet J.nd \tdc plate' Jdfcnes er al .. 1~96). So involved j, rhis cnntrnvu-ry
nd the inward migr:ltlon of the hydro~pin:s, d10ugh th,tt 1 can do no more here than -.ketch out a few
th~rc is no unequivoc.11 e'ndcnce. All ~piracnl atcs dc:t::tils and di rect the student w recent litcr.nurc:.
hm· true hydrmpire porc~ .md sptrac!es. The) gcn- None of the diverse organi~tm included 111 the lar-
mll} have the c har:~crel'\ of Pcmre111ites but may be poid~ has radial ~ymmerry and most ~how no pi:H1e'
long and thm (e.g. Tr,,Mr.-,ainus) or near-~;lobul.tr llf 'ymmctry at all (c:xccpt m the btlaterilly wmmer-
Mth rhc amhuiJna extcndmg nght do\\ n to the ru;.tl 'tail'). They do. however, have proJedmg
l'OtKave ba\e. Gt·ner:~ are dtstmguished by the plat- pl.1tcd arm and tail-like strul'tulcs, intemal C\VItlt''
lllt-i >mtctun: and the number of hydrospin: folds. .111d pores, and the problem h.l\ always becn ht)\\
to interpret them. Here tcnm from both the trJdi-
Ecology and distribution of blostoids nonJI classification of the Tn•d/m 011 lllflall'bmrc
BINoids were never Jn .1huud.mt compom nt of any P.lit'OI/tology and th<ll ofJeffenes arl' both rdC.·n·c:d w
t.lunl cx,c:pt locallv, when· in rhm bands they are The.: Ordov1nan Dc~tdrocysrcwlr.l (CI;c;s HomuiO-
'nmetunes so numerom as ro lll;lkt• up the bulk of slt'lc.t; or Solut::t of Jefferies, 1990) Ius ,, lobed,
the rock. TIK·n..· .m:, for tll~t.mce, well-known bJuds Jsyn1metricJJ, flattened theca of small liTl'gt.ilar
:n the Vhl'Jn of northem England, where pl.lte\, a singlr 'ann' , and an elongated 'tail' w1d1 a
l)i itmmto 1~ so .1bund.mt ,,, to pt'mllt mJthemarical \lOUt po(y~eriaJ prOXImal part ,md .1 long, t.lpl'nng
'llldy ot vanauon and rl·l.•t•vc growth Qoysey. thst.tl poruon. A hydropore (wh1ch indilatl'~ that a
''l'il/). Th~~e mdiv1duJis wen c~pect.1lly .1bundant watl·r--vascul:u \Y'>lcm was present) and ,1 gonoporc
umnoid bJ.nk dt·po~ib c.lpptng rtef knol l ~ lie close to thl· b.l'>l' of the ann, while thl' .tnus 1~
The fl'\'< Stlunan bJa,tmd gctH.'f'J., both ti~stculate nc:.1r, but to the nght of. the 111\ertton of the ta1l.
ol md ~p•rac:ulate. are entird) fl''tncted to the North I here 1s a single: gtll ~ht on rhc left-hand pmrer ior
~
~merican comment, but by thl Devom.111 hl:l~toids \td~:. Ccratocysti.- (Cias~ Scylophor.t, Order CornutJ)
luJ bccnmc worldwide Tlwy rcachl.'d .1 nuximum bc;·,tr' o;omc rc\l'lllhlancc ro Dr~tdrarystordcs dc~pttc
m lbundame .md Jtstribuuon 111 the Lower h.w111g largcr plates. but there :-~re alsu many
• Jrboniternu~ wnh .1bout 4'5 gclll·r.l. Tlwugh they ditfcren~:es, indudmg the prc:sc;•ncc of seven wll ~he<;
;re nrc 111 tlw h1~her CJrbmut\.orous. biJ.~tmd~. often 011 the left-hand \ldt: of the hl•;td. The mo't l'Xtrcmc
of pecuh.tr form, .ue found in proti.J,ton 111 some vt'riiOn of cornurc oq,ranttttton 1s f()und 111
,.!\ loc.thzl·d pi.Kes 111 the Permian of the e:mem CMhrmrocystis. Tlus j, a latcr::tlly Aattencd form with
(mi1pht re ~ta h a~ Timor llv the t•nd of the J boot-slupcd theca ,md a 'tail' not unltkc: that of
'muan they were exrrncl. Vcndrl'cystoidcs. A marginal frame of stout plate'> sur-
trc rounds the thel.l. ~:xtcnding tnto J 'ton~·uc'. 'hcel
ate ~trap· and 'toe' for the boot. On one of the: fiar mr-
the f.ln'' there arc prOJClting studs ,111d .1 stn·ngthemng
.tre strut crossmg the theca. Small rounded phtl'\ fonn-
tl\11 ing <1 Acx:ible cover occupy the ~u rfacc~ of the thela
111 l - w1thm the margmal plates. These; arc ~hghtly d1tfer-
the e calntc-pl.ut•d ,J,}'Tllmcrncal tosstl~ known vari- ent on the two ~1dc~. On the opposite <,Jdc ro the
.Jbl~ ' il~ horruJozoam. c:.tluchordarc~. or by the ~tntt ,ue some 15 sills arranged 111 J curvmg .1rc, each
lh and mort• non-commm.ll name 'c.1rp01d~·. :.re wtrh a hangmg lbp of small pbtc\ hinged above .1nd
I th.:- rhaps the rlltHt biz.1rrc .111d controveN.tl of all ftxcd externally (f<ig. 9.49c). Behind the 'tont,"lle' is
,11m extinc:t im crrebr.ue~. I hev hJvc tr.tdition- a pyranud of platcs closing an onficc.
becu rq~.1rded ,1, t·rhltlodcmlS. but f(,Jiowin!; lt "agreed by Jll .lllthors that Cotllllrttol)'Sfrs pmb-
no (I %H) m.my modern 'cholars hJve .1scribed .1bl} by fiar on the.: sea Aoor 'upporred hy the
.m to Phylum Chord.1t.1 (whtch mdudes the ver- 111:1rgmal spme~ Jnd the studs, with the surt:lcl' \\'lfh
ratn). jtdTt•nl.'~ and others use the tenn 'calct- thl· stmt lying 1n proximity to the sea floor, thl' ~lits
'\in'date,·: thcy believe thl·m to be olntl·-plated l1ppennost and thc aulacophorc fixed in the mud
308 Echinoderms
or
tht
thl
Dondrocystoidos n .' l
Rh
tht
. 1~'1
prt
.1~
l".'\(
Ul·
itth
on
.Ill I
nth
tht•
bro
(a) branchial slits .1u l,
(e)Ceratocystis the
Vt'r.
Ill()
prt..
~·lie
t•xh
.~<.: n
t ht•
or
.Ull )
(d) olfactory nerve Au:
Coli
( h<
E
Cephalodiscus-like ancestor Mitr
whio
thct·
llll'V
this
Echinoderms c:.Hc
plate
of tl
(g)
~itl' I
n..tlly
tht• t
!!rOO
Chordates
Mitrocystella m to
br.111
F1gvre 9 49 (a) Dendrocysto1cles (Ord , ' 1), (b) same, in side view(>< 1); (c) Cothvmocystis (Ord.) in dorsal v1ew the orifice X 1s inll!r· ~tmc
preted by Jefferies os the mouth ond by Uboghs os the onus (x 1.5); (d) bronchial slits seen extemolly, showing the 'hanging curtain'
of covering plates; (e) Cerotocystis (Ord ), in Clorsol view (x 1.5); (f) Mitrocystello (Ord.) in dorsal view with righ t·hond plates removed m at
showing nervous system os interpreted by Jefferies (x 5); (g) Jerferies' view of o Cee.halodiscvs·like ancestor giv1ng nse to o dexiothel•c slit-li
progeni tor of both chordates ond echinoderms [(o), (b), (e), (g) Redrawn from Jefferies, 1990; (c), (d), (f) l:>osed on Jefferies, 1968]
Subphylum Homalozoo, otherwise calcichordates 309
!lr ti-~1!. l:.xpc:nm~•m \\ 1th m odd~ mdicate that the higher nervous complex witlun the vertebrate
the t.ul \\ l\ U'l'd ro pull tht: he.1d rt lr'\Vard' across skull. In mppon of thl\ v1ew, the .l!>ynunemcal
rhe \C:l tloor (Woods .mJ Jdfenes, I'N2). aud the pharynx of a rumcate, for exampk. would t1r neatly
li'Cent di S(ll\'Cry llr ,l tl.lU.' to~'ll with the allied mm the head of a J\ lim>cystclla. Ubagh!> ( llJ7J). on
Rlllllclllacptis l~m~ .1t tht• end of its trail confinu that the other hand, has mrerpn:tcd the frmgcd opcnmg
rfu, \\ ,1\ very hkd" th~ t',l\l', Thert· i' much dis- as a penproct and located rhe mouth where Jeffenes
At,'T\'l'lllCnt .h to hn\\ tht•w -;tmrrure' .lrt' to be inter- h.1s suggested rhc .mm 1~. I li~ .lrt,'tlnwnt th.tt the 'car-
preted. T}w more rr.•dmon.•l ,·icw of Ub.1ghs (1971) paid' skdcron ~~ ~tknncal in \tructure with th.lt of
as to the hnmologtt'' ol dw v.1nou' 'tructures vie~ cchinodcm1s 1s tountcrcd by the ·calcichord.lte
t.xc~cdmgly with th.u ofJdli:rk' .md lm coworkcrs. school' who argue that there '' no rca,on tor it nor
UbJglh h.l\ COil\llktt•d tht• 'hr-hh· pore~ to be the to have been; \llCh .1 ~kcleton ha' bct'n retained in
mhalant opcnlll);,"i of re,pir.ltnry nrg-;lm, the large the echmoderm3, .md lmt 111 the later chordates.
rmfin: hdund tht• 'tongue' to be the .mm, and the Whereas the whok •~sue rcm;nm controvcrstal and
.miJcophnrt• to lx· J ll-t·dmg org.u1. jt ffcncs, on the unresolved, it rcraim its f.Jscin.ltton.
other hand. h.l\ propml'li vt·rtd1rarc humolugies for It has been known for a long time on embryolog-
rh~ vanou' org.lll\. l Jl' h.h ~ugge~reJ that the theca is kal and anatontic.ll grounds that the echinodcmlS and
bro.Jdly homnlogou' With till' chmthlte he:Jd, the chordates arc indt•cd rdatcd, and ~ome .mrhoritit·s
.mlacoplwrt• ~~ t•qui\iJient to the chordate tail, that would group them together ~l\ De";othetK<l. jefferies
the 'bt-lth· port·~ .ut· t•xh.li.lllt md homologous with {lfl8o, I990) and JdTcric~ 1'1 c1/. ( 19%) v1ew the latest
mtrhr.ltl: g~ll \IH~ ••md th 1t the I.Jrgc orifice is rhe common :mcestor ofhoth a~ ~omt·thing like tht· mod-
mouth .111d t•nrrJIH t' tu lht· rt'\pir.Hory 'Y~tem. The cm hcmtchordate Ccpha/odism.1, but lying on ih nght
rrt\tiKt." of th~ h.lll~lllg lllft.llll <>f \lll.l!J plates Out- ~idc in the mud (the dc:x1othel.lte condition) and
Side the ptH·~.:, doe' Ill t:t~· t 'u~gc.,t th.lt the) were thereby lming tht• upentnb~ and ann' oftlw pnrruttvt
cxh.tl.mt \tnltlm~·' · Till' anu' .md gonopon:, nght ... idc. Thc vanml\ rem.umng orgam r.m be
JCCo.mhng to Jt.:tl'l•ric,, art• ,1 \lll.lll pt•liorJtlon near homologized w1th tho'e m tht ·c.lktt hor<.Lte'·. the
th~ ong1n ut tlw \tl'lll. ( ;iiJ-,hb nn rhc ldi:-hand side ~talk of the dexiorhcut .lnn\tor being the cquiv·.llenr
oi tlw head rt't',lll tht· 'tnKturt'' of the larval of tht· tall of Dcudn>t yswidn. CotlllmllllJ'jltl .md by
ampluo:xu~. wh1ch likt•w ''c ha' kft giJJ-,Iit, only. mfcrcnce. modern l hord.ttt''· Dwdr,lC)>ftlldrs. \\ hich
.<\ccordmg to .Jdlcnt·~ buth Dt~ufr,>cy~ t~,,dt's and had developed a calcit~·-platt•d 'kdeton. j, ,t·eu to
C.,tlllllllt>tysti) bclllug to the \lt'lll :,.'luup of the occup} .1 rnucal po~ltlon m cvoluuon of the dun-
Chord.lt.l. dJte/echinodcnn pk·xll\ sltltt 1t ret.uned both a t.ul
Even mon.: nt .1 prl1bl~m I\ the OrJo\'lClan genus and J water-vasrular 'Y'tcm. The iom1er \\J'> lost m
.\litn>cystd/,, (CL\,. ~t\ luphora. Order Mttrata) the cchint,denm. wlulc the lattt·r Jj,,lppeart·d 111 the
11h1th h.l\ .1 l.ug~.:, t!Jctellt·d. thtlllgh ,t,ymmetrical l,Hcr chordate~ along With tht· prmutive lalnte-pl.Jted
theca of irregular pLitt'\ and a '>hurt, \tumpy, down- skeleton. Here .1gain the mttratc~. accordmg tll the
curved \lt'lll or tJtL jdll'rlt'' (I lJ6H) ha3 described 'cale~chordatt• sthool' .tre of l.ntltal unponance, ... mce
tlm fonn .1~ rt·~emblmK .1 'l,uge talut~.:-plated turu- they seem to have l-,'lVcll n'>e respccuvdy to rlll' tulll-
cate t.tdpolc., r ht• tht't',l h.t'> .l lllargin.ll run of large c:Jtes, cephalochordates and cr.uu.ltC,, by mdepcndent
plJtes, wluk rh~ ,\t~htly ~m.tlkr pl.w:s of both sides loss of tlw ta!ute-pbtcd skckton, on thn."c ~t'pJr.lte
ofdw tbn·,• .lrt• tnt•gul,lr. An up~·mng at the oppo- occasions. Hence tht• mitr.llt·~. from the cladi'>tll'
ll!e end to tbt· \lCill k.td~ tn a tcntral GlVlty mter- point or VIeW, ~hou(d be J\Sl~lleJ lll the ~le m group~
nilly d1v1ded by nd~t'3. In tht· tnner dor!>al surfuce of of these import.llll ta.x;l.
th~ tbel.'.l 1rc lmprc\scd \ene' ot ~tdlare .md dongate It I~ or cour~c cqua!Jy pos~tblc that the line~ of
~oovc\; the\~· h.l\'t' bc~·n rct on~tructed bv Jeffcries descent lcadtng tl1 modem c hordJtes took place
mto .1 complex 'l'l'lt' ' of 'tnu.: turc' renuni,n.:nt of the amonbost \Oft-bod1ed Jmmab wlud1 left no fossil
bram and cr.uu.tl lll'r\'C\ of a ti,ll. Tlwsc and other record, and that the 'carpOlds', rather than being 111
structure~ (e.g. tht· ll'fr- l.ttl'ral pom10n of the .mus, as the drren lme of de~u: llt. were· merely .1 rather
,. m a nnucatt' t.tdpnlc) led jl'lli.·nc~ to homologize the peculiar sideline who'e t'\'Oiuuon wJ' m \umc "ays
d
ic !lir-hh· anrcnnr untict• '' tth .1 mouth. the ~tern with paralld. The debatt• rngge1cd ulfby Jdft·ries· ong~
,j ~ \'ertebrate uil .md rhe r.umf) mg canal !>)'~tcm w1th nal wod. ha~ continued fm 311 yc.m, .md ha~ pro.,;cd
31 0 Echinoderms
to
pa
G<
tht
Jl1(
lie
de
"b·
rh
ed
.tin
bio
dat·
har
gro
fro1
~c to t
·g_
Cl)
c
0
den
'
( 19:
1,& ea le
Vlt'\
adu
re~c
cch1
witl
eocl
I ate<
side
ated
togc
bran
midi
cord
on l
gut i
dire<
coelt
Ec
The
s1fied
Cam
Evolution 311
to be perhap~ rht> ltvdit•,t and l crtamly the mo~t mmt pmmuv e belong tu the Cmt t;l, di\l·-~hapcd
pJrtisan conrro vern 111 tht \\ holt of pabcon tology fonm such .h 7 mchoryswes, wluch h;1s ~uull p!Jrt·~
Gee\ (1996) work 1~ the mmt ol~lettiVl: recent syn- surroun ded by a margin al ring of stoutc:r pl.1tes. Jnd
thesU., and trc.m tht· \Ubjett m grcat det.ul. He com- a short tail Amon~t the carhl''t cchmo denm is the
ment' 'my \)'mpat h} fi>r the nlncho llilte theory monog cncric Cl.lss Hehcop i.Jl·uide .l. spcnm cn' of
hL'S in rhe wc,1lrh ami interest of questJOIIS it cngcn- wlm.h wcre h~t dtSCO\ cred in 196 ~ 1nd arc known
dtrs . not hecauw I advot.ltt> ch.lt tt 1s "nght" or from c1 ll'W lm::~littes only in western Nnnh
"better" ·. Admir.1 blv f;ur tomme nr. Amcnc a. The body ts ti1sifon n. l.1ckmg pentam t•ral
5ynune try and wnh rhe plate~ (Jnd llltt'rn,ll mustlL·,)
spirally an.1ngc d. In three re,pcct s it ~lure' chanc-
ter; (:~utapomorph1es) w1th hvmg et hmode nm: the
plate' have typical ~tereom structu re, the ;:~mbul.tt ra
1 here '' no gcQiogi t.ll evtdenc c .Is ro the .mcc\tr y of (three in number ) haw biscnal CO\'er pl.lres, .111d
rchinod cnn,, 'mn· rhc f1n.t n:prco;e ntanves appc.tr they are r.1dially .trrangc d round the momh. 'The
.1lready dit1ercn ti.ltcd 111 the Lnwt-r C:tmbr ian. On mourh 15 laterally positiO ned ,111d tlw .mm ~~ prob.l-
h•olog~ul ground \, 1t ~ccms prob.1h le rh.u the chor-
bly JpicaJ.
d~te-edmlOdt·rm ~ronp (Dexior hcric.l) on the one
Three o ther, sligh rly l.ner Lower CJmbn .ln
lund. and tht lophoph or.1te (brJthi opod-b rvozoa n) cchu10 denns have been descnb ed. Lrpuft,cy.>crs h,,, ,1
group on thc orha, \Wft' mdepcn dcndy derived stalk,:\ cup. ;~nd .1rms w 1th ambula cra runnmg down
from .1 commo n worm- hke ann·,to r. ,1lhed pl'rhap ' them. ch.1rac ten uniLJllt' ro cnno1cb Scr,>lllil l•'tl'!ifltt's
the modern Sipunn t!id.l . ha~ .1 free-hv ing disc-,h. 1ped body w1th fiw .mlbu-
We h.tve .lirc.1dy rl'fcrn·d tu the model ofJdferie~ lacra 011 the first upper ,urface . le look~ mlllh like .111
11'Jl16} and Jdl~ric\ l't •ll (1lJW,). winch would edrioJ~rero1d. C.wapto.\crcJma hke\\oise has a Aat uppa
dcnve t honbte s !Tom cak1churd.Ht:~ by loss of the ~urftce w1th a 1-.ind of pcntam ery, .md a co1mal
Jkite-pl arcd skckto n. There arc, of cour;t'. other body below, but H re tamed a ~pu:tl 111tenul 111U\LII-
l'ltw,. An altcmatJ\'L' propo\J I ~~ that the ancestra
l LHure. Even in the Ll1Wer Cambn .m the\c e.1rly
.1Jult of both cchuao dcrnh and chord.u es closely echinod erms are widely sep.1rated 111 nmrpholnt-.'Y
~mhled the 'diple uruJa lan.11 ~tJ~t· of modern
.md the1r rclation~hip' c.m only be undc~tood
echinodcm1s rim Mn.ill pl.mktl t Llrv:~ ~~ cylindri cal, dadi,tic ally. Smith (19HH). ming thi~ approa ch, 'ee~
1•ith 1 ventral mnuth .md anm and 'Ytnllll' trical Ht'lrrc'Jllams as bemg so ~enerah7cd that tt bdon~ to
mclonllc poul hes un t'ithL·r ~1dc. fhcre '' aho ,, cil- the 'lem group of all lalcr cthmod cnns, ,md from
llt~d b;Jnd fornun g J loop .md nuuun~ down either which evoluti on took place 111 nvo mam d1recti mh.
11Je Accord mg to the 'd1plcuntla theory· . the cili- of which 1r was dose tQ the latc~t commo n ance~tor.
Jt~:d b;1nd w::ts pll'~cnt m the ;ul(c~tral adult.
The first derived group were the 'pelmatozoan~'.
rognhtr with 1ts cuorcL n.mng nen·e plc,-us. In the criuoid~. blastozo .ms and othL-r 'fixl·d' ecluno denm
branch that led tu dwrd.lt es tlm band h.tscd 111 the of which Lepidorystis was an early repn.•sentative.
a1idlinc to provide the rudnne nts of the dor<.al nerve The CIJ." Eocnno idca (Subph ylum illastoz o.l)
orcl In the .1nce~tr.1l bl,lstl)Z O.m-cn nozoan stock, wlmh belongs here becamL' very diver;c by the
cm the nth er h.1nd. turuon kd to the twl\ting of r.he Middle Cambn an and persiste d into rhe ~llurian In
.at intuitS present pu,ition wnh .1 central, up\\ard Jy many re~pcct~ the~~: fo-.sili. re-.emb lc the Cnnoid t•J
d1recred mouth .md a later.ll anus. '"'bile one of the but their plates dose right up to the pensru me .1nd
roelomlc pouche s '' .t, lmt. rlle) do nut ha\'C a tegmen . Subwq uent develo p-
melll~ include d the later Bla~tozoa proper. of wh1ch
the Cocrin0 1dca rn.1y have bet·n the anct>stral sto<.k.
Earlies t echino derms and the ir There arose the ~.-y~to1ds and lart·r d1e blastult h;
radiati ons (Figs 9 .50, 9 . 51) short-li ved fonm of bizarre appearJ nce. the
paracri noah and parabla stoid,, aho origma tcd .1nd
The earht:st crhinodern1~ were alre.1dy highly diver- soou beuune extinct . Me,tnw hik, in the early
l:fied on thl'ir first appeara nce in the Lower Ordovi e1an true crin01d s had origin,n ed ro become
Cnnbnan (P.ml .md Smith. fi)H4). Pc:rh.1ps tht· the domina nt pdm:Jt ozoan group.
312 Echinoderms
\'1.!
Ill
ap
gr
0
Pnm11lve
'ophiuro1d'
Archegonaster
\
•
Cystoids
~n
Stromatocystttes
bu
/ .l '
1111
en
dC'
of
C11mptostroma
IZC
Or
eta
pt•t
tht
r.td
Figure 9.51 A possible scheme for the relationships of early echinoderms. (Redrawn From Poul ond Smith, 1984.)
111
P.tl
Tht· 'ccond luw of dc~cenr led to all clc:uthero- nn~ of pbres. Thts gcncraJJy comtderl·d J~ an
1' tnt
llMm (.1\termd\, nphmrotd~. t:(hmouis ,mJ holo- cdno;l\tcroid (and lllldcmbrcdly J lme of dc\ll'llt to \lll
thunam). Smtth (1 1JHH) tdentitie\ rhe Mitldk true l'drioasteroid' kd off from 1l thn..1ugh grc
C<tmbrian c;,,/1/ll'tWrr .1~ ,1 member of the ~tcm group Sti'OIIItllc>cystites). Smith regard~ it as bdnngmg to ·• th e
which gave me to the ckuchero7o.m~. lt i~ a ~m.1ll, ~lem group which .tl,o gave me to .111 Llter the
di\c-,haped edHnodt'nn with tivc pcntameral deuthcrozoans. The ,l,rerozoam denvt'd from rh1s \1114
ambu!Jcral :mm kJding to .1 ct:mral mouth, near morphology. b} t•xtcmion of the arrm .md n·dul- ma
"hirh lie<; the .1nm. .llld a pronuncnt marginal uon of the margmal (r:unc: of plate~. ·'' 'c:c:n m th,· gl'C
Evolution 313
earl) OrdO\'lLI.lll r ·;udmtl /ciSICI ,llld the comem pora-
c;ech evolut ionary srcp .1rtcd ,ts :1 spnng board for the
llCOll\ ophtu rotd flr<~eduum. Tht·r~· .tre ccrrai n next.
~yn.tpomorphil. eh tr.lcter s "hi eh the
pmmt ive
ophturotd~ ~h.1r1.' with the carlit·st t•t·hino ids:
Jo" of
Ull margm al fr.Hllt:, cndo> un: of
the r.tdial water Evolu tion of the tube feet
vc~".+'. the pn:sl'n n: of arncul .ltcd 'pines .
and the
motht icannn of the oral .unbul .u:ral pl.tte' nuo J jaw Tht:
eflit it·nt me and dJ11i:rcntiarinn of tlw tube feet
appar.ttus. It lu~ bet·n .1rgm·d. theref ore. on dadistiC
lu\ dearly been very impon ant to cchm odenn cvn-
ground;; that the t:Jrlu:st eduno 1ds (rbe late
lutton (Nichol~. I 974). Crino id' have the 'implc
Ordov in.tn A u/cdti11w .md htcun lllliiS) togeth er ~t
Jtl<l appare ntly mmt pnmit lvc sy>rem. In cnnozo,m~
with &11/mandaris tin m the si5ter 6'1"oup of the oph i-
there :ue no ampul lae ,tnd tube feet arc extrud t•d
uroi<k Tht·se hr>t t't'hinm ds h.td a 'good Baupl an'
only by the contr. tction of the radial "arer ve-;sel,
but Jt\ potent tJl wa~ not h1lly w be re.tli7ed untJ1
thl' whtch i~ rhc nnly pressu re gcner Jtor. Edlli1 01d'
\1e\ll20 1< A p<Nitbh: sister group ol the Jnd
cchin0 1ds I\ dcndr or hirotc holot hunJm have unpul
the Oph1o t ystotcLt which h:wc bun-~h.tped inthtc d lae, whtt h
art· rhc main pres,u re genera tors, but toutr. Ktiom
bodte~ and enonn ou' pl.ttt·d tube ket. but
aho poS~l'\\ An\to tk\ bntem s. l"he holoth urians (of
which of the radJal water vessel arc .,nu unpor rant.
Aspidochirote~. on the other h.md, c.m
wh~t:h tht• t'Jrht.·,t " the nl'\'On ian Pctlcll'cl(ll(llllh extend "
lria} rd.n~ing tube tom only when a nctgh bounn
proh,tbly .1rosr !Tom ophuxystmd~ by plntr reduc - g one
contra cts, sendin g water mto the t•xten ding one.
hon; the \tlun 111 R,,r,,_,,,,..,,s IS 111 many \\ .1ys mter-
Astero ids and ophiur01d~ have ampul l.te, but to ,,
nu:dlate bctwt' l'n ophjo cy,tol ds ami holotl uu;;m s.
v;~rying exten t inflate the tube feet by contra
Opluu w•J,, ecl11n uzoam md h1)lot huri.m ' art· thu~ cting
the lateral and radtal "ater v~.-s~cl. The tube font
1een to bl· 1soi.lccd anJ ~nHth ( 198-t.t.b) pl.tcc~ them
exwns or system 111 cxt111ct cchino derms ha' been
together .1, Supcr da" C)'rto synng tda
mtcrre d by analot-.'Y with that of mode m to nth.
I he abtwc: an;tlym (F1g. 9. '1 I), bN:J on Patd .md
~rmth ( I'>H4) .md \mith (I ClHX) IS not t·xluu
stive,
bUilt lC\llftCS to th1.• V tlut• of cJ,Idi~tll ,\nJly~is in ~uch
Why penta rnery?
.ase a' thi,, whcrt · 1t '' 'ough t to undt•r .tand tht>
lntl·rrelatHmShlpS of 'uch tlivl.'rsc and wtJdy ditTer-
Not all ech111odcrms luve pentam cral symm etry.
·nt orgaHI\m' .I\ tht ead~ rJdnn on of tlw cc hmo-
E.1rly cy~toids ,tre rndially symm etncal bm m.ty have
Jenm produ ced.
t'ivc '>ymmetrical plates in tht• anal pyram td. where .l'
In summ ary, we 't e an tnittJl Camb n.m radun on
later cyo,toid~ acqtm c pl'ntam eral thcc.lc . But other
of wr} d1vcrse fonm but livin~ onh 111 small. local- tlun
in 'carpm ds' and other e.1rly group. ,, pentam eral
JZ<'d :tnd 'l Jttcrt• d P<'t kc:t~. A turthc r r.1d1Jt10n 111 tht• S)
111111etry i~ so charac tcnstic that there must surely
lhdovi tian led to .l gre.1t .md r:lptd r.tdiat iun, t'<ipc-
b~: .1 gond rea~on for it. Nicho ls (11.)74} hJ~
Jll> of the pentam cral group ,. Dunn g sub~e4uc.·nt tl1Jt sugge sted
tht• \uturc s bmdin g l'Chm oderm plates must have
pt'rtOdS tJw gn;,lt r,lnge of higJll'T t.tX:l t'StJbl
i,hed Ill acted a\ lmes of wcJkn e''- Only if there
mt Ordov u.ian """·'' dr,,,nr aU) pnmc d The non- odd numb had been .tn
er of plates would 'oppos ite' suture s have
odi.tl fonns were extinc t by the Lnwe r Devon ian.
been ofEet \O as to give greatc.·r 'treng th ag.wN
~r.J m.my of the puuam eral group~ becam
e exrin n brt•akage. A circlet of three plates 1~ too
wthe Penm an if nut bctl.)rc. b·olut 1on 111 the l.ner few, where as
five ~eems to be an 1deal numb er for maxm lllm
l'ibt·ozolt ami .1ftt•rw:trd' Oltllrr c:d only in well-
rc~•stancc.:. A Simila r augub r ot'Ret is neede
·rid and Jdapt. tbk tJX.l. '"htlh \Ur\ IVc:" with great ag.uns d to gu.1rd
t accide ntal ntptur e along lmes of pore,, a:. m
1rcc~s tod:~y. The p.lttem of evolut iOn wnhin
theSL' an echino id, which has been called 'posta ge
;roups varied 1-.'l"CJth. In a..sternzo.lm, for tn\t..tnce, ~tamp
\Wakness·. The ofhett ing of pore rows 111 multip les
ih prJl of tHgam z.ttlon w.l, re..tt hcd e.1rl~. and of five
has been an 1deal comp ronus e betwe en
·rl.' ~et•m lO h,l\'l' be~.·n no major mnov. .trion\
stren~th of te\t and JDa.'\lmum numb er of tube
nr,· the Pal,lt'OZOil. Tn t'l hmoid s, by comra st. fe~:t.
On the other hJnd. damag ed echin oiJ tests r:~rdy
JO}' ne\\ t\ pes emerg ed throug
hout ~un·e\Si\1. brc..tk along the mrurec;, \tnCe
.'C<llogicJI pt'nod s, .md 111 certain hnc\ of de\ccn t the lancr ,tre rt'-
inforc cd with strong collag en fibre~; hence the
314 Echinoderms
~utures are tHJt necc~\.mly the weakest p:1rt of the specialized, luving ~nppresscd or chm.inJt~d IIHit( I' 1u1
tC\t. Another suggesuon i~ LlMl a p.1ttem of tive generalized trait~. Perhaps the genetic pott•ntl.lllor, P.
combmc~ .1 nearly const,lJlt width pn.'~cnted in say, folded membrane ~tntctures was prewnl 111 c
almost any d1rt'ctton in tlw horizontal pl.111e with a many Palaeozo1r cchinodenm, though \Upprr it'd ~0111
~mall number of ray,. 111 the majont) .md only realt?ed in a ft.>\v ''ul.ttfd
.u
groups. Wlutc:ver the ultlm:ltc :mswer. the'e per- Wrij
plexing early invc:rtC'brates, more than those nf Jll\
c
JlJ
Conve r gent e volution ond interme diote other groups. \c:em ro '>potltght some complt' ;~nd
forrn s httle-kno\\ n a'pcns of evolunonary theory t!m
h,we so far received but scant .Htenrion. lr
A strikmg l har.Ktcri,tll 111 the early echmodenns i~
for the \Jillt' kmd of strtu rure ro develop over .1nd Bell,
over again 111 th~tinct though unrelated stocks. An Ec
example ~~ the folded nu·mbr.me stn1cturc found in 21
bbstoids. t y~totLb .md the: cnnotd famil) tu
PorocrinOidt•a, m .1ll of wbu:h tt ~ec1m to have Blak•
Books, treatises ond symposio dfl
evolved mdcpemlently. P.med pore:~ h.we Jnsen
mdependL'Htly in diplopontc cy~toid~ and 111 echi- 25
Gl't' . H. (19%) Rc:(tm· the BMkf,om·: I 11'11'5 011 tl.c Orw11 of
notds. 1 am:colatt' e~mbul.1t r.t and a <;tern art· present Bl.tkc
the· I 'atebmlt'(. l 'h.1pman & ll.tll. London (<. rmqut• • f
po
m blastoi.d' and in tht· pen1har genu' Astrocystrtt'S, cL hmodem1--c:hord ate rclanomh1p~J
)or
which 111 otht'r re~pecb I\ more hke .111 edrioast- J.tck\on, R. r. ( IIJ 12} P/rylll,t?fll)' •!fl/rr hc/11111 walr ,, Rol.'ISt n
Bl.1k<
eroJd. It ha' bt>en propo\ed thJt a nt>w class, the 4 P,r/rozt>rc S/ll'lrcs. Mcmotr. of the llo,toll Snuct1 tU
EdnobiJsto1dea, be est.tbh,hed to accommntltte this for Natural ll tstory No. 7 (I .tr~e mono~.tplt ''' .1St
genus, but it 111:1) ~•mpl~ lll· better to regard tht\ Pabeozotc elhinntds, profusd}' dlustr.ncd)
Bolk
Jdfcncs, R. P ~ (I 9Hl>) Tirt' Itlft'.<lry tl} tlrr I 'ntrbr.tU
genus :t\ .m ednoa~teroid nmvergcnt on bl.1~totds. Nl
13rin~h Muwum (N:mtr.ll Hi~tory). LonJon
Very many uf the LmH'r Palacozoic erhmodem1s Brcin
(V crtcbrJll' .lflirunc~ of hornal~.>2<.><trl~}
.ue thu' of .1 cnmpo:.1te type which, .1\ RegncU of
Moon·. R..C (l•d.) (196C•) J'rrntisc un lrwrrttl•hrt
(1960) h.h po111tcd out. ltt• in the dtrL't'l hneage of Ne
p,,fconrolo..t!Y· Part U, llrltiltc•dmll<lftl 3, Vl'l' l trlll 2
none of the later eciHuudL·rm ~oup~ hlll present (At
Geological Sonery of Anwm ,I ,md UntH'f\lty n(
tos.
features from ~cvcral 1.1i them. Pnlups rhe most Kansas Prc\\, l ~wrenn:, Kiln.
Bn·in
extrcm1. rase may Lie 111 the Cl)nvergente between Moore. R.C. (ed.) (19<>7) f'rCllll51' mr lm·, n.l1Jit
but
the 'carpmd' Cillhunwq•.'l'-' and the t)''toid Plellrll- Pah·outolt>I!Y PJrt S, Ecluuodtnllcllcl 1, Vols I anJ 2
Pro
<ystitl'.>. lnde~.·d the controversy over tht• reLttion- GcologicJJ Soncry of Ana: rt~·a .md Untwr.Hy t•f
Bi
~hip~ of rht·,e s.1mc 'c.1rpotds' illuminatL'' rhc fact K.m~as P ress, I .1wrcncc, K~n.
llrctt,
that within tht· e:1rly echlllodcnn-rho rd.ltl' hnc onJy Moore, R .E..tnd Tt:ichcrt. C (t·d~) (197H) l'rt·.rtt.-<' 111
pre
J few kmds of structure' ~et> m to have been able to
fiii'Crtt·br<IIC f'alt·.mtolc.>.l/}'' P.tr[ r' F..clllm>cll'nllllM 2 'iha
C.cological 'lonny of Ament;t ..tnd Univcr.tt\" of (),
evolve Jnd carry out parttt ular tunttiom. K :m~,1s Pre\\, L.twrence, K.m . (Cnnmd\ .md rdat~d
The 'mtenncdJate fonm' chat have evolved, often (. .Jrlt'l
groups}
with a puzzhng combinJtllm of ch.Jr.ttlcr,, have ofs
Moorc, R.C., Lahcker, C.G. Jml F~her, A.G. (l'J53'
I'a/,
been the h .tm· of echmodenn taxonomy. though l~rvertebrnt!' Pol!'<>lltolt~I/Y · McCraw-1 Iill. New York
md
tlus is bL•ing re,olvcd to ,1 cOimderablt' extent using Mortemen, T H ( 1928-1'>52) l ;\/ort•'l?'tlf'/r ,Y tht (.brk<
dadistit methodology lt 1\ not easy to try to pro- Etlriuoidca, 5 vob. R.e1t1cl .md Oxford UniventtV I'm;
star
duce an obJCl ttvc phylow:m·nc scheme where such Copcuhagcn .111d Oxford. (The l.1rgest \\ork ..-vcr puh-
(UI'
extreme genera an: involved. Quite ap.u t from the lt~hed on cchinoids, with full dl'scriprwm .md photo
('owe
r:.t.xonomic que,tion tlwre arc deeper evolutionary graphic platt>' llf ,\ll known Rl'Ct·nt ~pecie~)
.111
Nu:hols. D. (I'J7-I) t:chitrodmm. llutchm,on, LonJnn
~~~ues. Why, from the gcuettc pomt ofv1ew, ~hould 132
(Invaluable tcX[ on morphology .llld evoluoon ofh\1ng I ><)no·
early echmodenn.' ha\ e produced similar structures,
.md fm~il fonm)
independcnLI) but oficn .1nd so consmently? lt is of frot
Nithols, D. (1975) '17re l 'nrqrwllf'' •!I Erllillt>dcrms Oxtonl
course true that early m em her\ of ,1 group tend to be Pah
Universiry i>re\S, Oxford. (Short, de~cnptlvc. tum-
rather geuerJlJzeJ. whcn·.t~ Lttcr ones Jre frequently Do no'
rional appro,H h)
Bibliography 315
P~ul. C.R.C .•uul Snlllh. A. D. ( l9S8) (c:ds) Ed111/(1derm Crinoidea, in Ethirwd!'llll Phy!t~I!WY arrd E1•o/ulionmy
Plrylol!rny ~~r~d fr•t>luth''""l' Hrt•lt•.l!)' Cbrcndon Pr~ss, Biolo{!y. (eds C.R.C. P.IUI and A.D <,nudl), ClJieudon
Oxford (':I~·Hnl 1111pnrt.111t p.lpt'~"'>) Pre~~. Oxford, pp. :n::;-44
South, A B. (lliH4o~) F,/,,,.,d f'wlt~t't>Lnv/,,~Y· Gc:orgt· Allen Donovan, S.K. (IYHHh) The unprobab1ht) of J muscular
and Unwin, London. (h.,.cdknt tc:.,.t) cnnoid codom. Lt'lhmtl 22. 307-15.
\\'right, J. ( I'JS-t) Jl \(,,,,.~r,rt•lr •!ltlrr Hnus/1 Carl>,>rr{fertlUS Donovan, ~.K ( llJH'J) The \tgmtic.UlCt' of the Bnt"h
Cnu.Jid,•.r I ondon I'.ll.lt'nntol!rJPim al \on~rr. (Fully Ordov1caan cnn01d hun.1 \lt>drm (;,·,,l.>g)' 13. 243-S5.
illmrrJtcd monugr.1ph) Donov.m. S.K .•111d G.tll, A ) ( 1990) Predator; JqerOJds
.md the: dt·t:hm: of tl1l' arm uiJu: brJduopod,. Lctlwra
23 . 77-H6.
Individual papers and other r e ferences D~rh1m. J. W ( llJ66) Evolunon tmong the Echino1dc:a.
Biolt~~iral Rr1•it·u•> 41 , 1M!· lJ I (CI.1ssic study; allustra-
Bell, li.M ( 11J7h) . I ~twly <'{ i\'ortlz Jlmcrirarr nom of early echmmd~)
Edri,•astnonlt·,r. Ne:\~ Yorr... St,lle Museum Memoir N o. E:.rnst, G. ( llJ70) Zur 'lt.lmmge\l ha due und 'rr.Jtlb'Tapht~
21. (Mt>nogr.•plm 111t~·r pn·t.ttrtlll of .mthul.tcral smtc- chcn Ucdcurung dt·r E:chm1dc:tr-G.tttung Hrrraslrr m
curc) dcr non.lwcM dl'Ut)~hl'll Obcrkn:idc:. 11/ittt'ilwt!?t'll dt•r
Bhke, D.B ( I')1(.') '><'J11.t\tl:rotdc:.l. Am:ro1dc:a and the Geolo.f!isdwr-f>alitomolo:.!rSCht' ln ~ltlllf da U11ivrrsitilt
atlimnc:' llf /.r111/111 (1-'ltrta,/rrl<l') latrr.rd~trfa. Palm•olll!>lc))?y Haml111rg 39, I 17-35. (Disrnbutaon of i\ticrt~strr)
25, H>7 lJ I. (Piatmtrn.u nut J 'llllt.l~tc:witl) Fay, R.O. (1961) Ul:hLnid m1di~·~. u,;,,crsity •!( K.wsas
[llakr, O.B. ( l <JX7) A d,t,,lfication ~nd phylogcny of Pt~levmoh~'(ical Coutril111tioll.i 3 , 1- 1·-17. (MonobrrJphic
po\t-1Jabc:o7nit 't'.r-,t.lr' (A,tt'nlltlc:.t, Echinodermata). treatment)
]11unwl of !'J,,,,,,,,/ HIS I M)' 2 1, 4H I 528. Fell, H . B. ( 1963) Phylngl'ny of se.1-sL.1.r;. f>lriiMoplriml
Bl~kc, DB .•md Cut•mbt·r~. T.C ( I1JHH) The: water va.~ Transartroll.\ t>{ tiro? Rt>yd/ .St>ti!'ly o( Lmdo11 B 246,
rular W\tCm .1nd hiiHflml.ll morphology of Palaeozmc .1H1-U5 (J>os,Jblc: cnnmd ongm~ of \Onta~tc:rOJds)
Jitc.>rOid, . Ltotlrtli,, 2 1, I tN .:!Oh. Gale, A.S. ( 1987) l'hylngt•ny and cl.m1ficanon of the
llo<keli!!, F. (I Cl H-I) The: D1plopuritJ of the O~lo Rehrtou Asteroidea (Eclunoclcnnat.l) ZMI<•~ual )<'""''''
t~( tlrr
1\.:orwav. f>,dw<'<>trlt'l<'l!)' 27, 1-<>H. urmra11 StJCicty 89, I 07 32
llremter, A. ( IW1Y) A cnnrnbunou ro rhe pa!Jeoecology Goldnng, R . and Stc:ven,on, D.G . (I'J72) 1 he dq.'o'J-
of P..tl.aeuzoJc \t.llknl ~nllollk Pn.•<n'diiii/S Kotrmklr.l/ tiolul cnvironmcnr of three qarfl\h beds. Smrs
.\'rJtrlllrrds .1/.:.ut. 1111c I l'tlltli<IIIIJlf'tll B 72. 13<).....50. Jalrrluulr flir Gel>lt•l/lt' rmd Ptrlii<llllolv'!ic Al<•rmr~/u:_lit 10.
(Apphnnon nfkmm ktlg<" nl rc:ccnr l nnoid ecology ro 611-24. (Raptd bunal es~cnual tor \tJ.Ifi,h pre,crva-
lo<~il~) rion)
Brenner, A. 1nJ Wd,,tt·r. C J) (IY75) A tiarther conrri- H awk.im. H.L. (I Y12) Cl.l\\llr{Jtllln. morphology and
burion to the p..tlll'~lccolol-') of ti:mil ,uJkc:d crinotds. evolution of th~· Fl hlllllldca, Holt·ctypuidc:J.
/til(t't'<iiiii!S "''""'klt11 t\ctft r/,,,J, . lk,ufnlllc ll'etter:;dwppen Procerdill)?$ o( tlrt' /,o,>ltJ.I!Uill ~.•aft}' t>( f...o11d011 1912,
B78, 1.\1}-(17 (A' tbuvc:) 44(}-<.J7.
B~tt, C.E ami Ed,.~·ll, J D ( ll)l':!) Pal.Jcucwlut,'Y uf a wcll- Hawk.im. H .L. ( 19·13) Fvulull{lll .md h.1bit .llltont; the
prt"ic:rvt·J cnno1d t·olony fn11n the ~tlun,m Rochc~ter Edunoadea: ~omc f.tct~ ,md thconc\. Qu,rrtaly Jounwl of
Sh.1l~ m Ont.mn T.J.I•' ~11'1111' Co11trilomum.< '!f rlrc Rt>yal tire ( :eologrc,tf .Sotiety oj Lo11do11 99, 1-75. (Snmubnng
f Olllnnto Afmt'lllll 131, I :w (Fwdlcnt illustrations) paper on cchino rd phylogeny .lnd .1dJptat1ons)
wn:cr, R.M. ( 1967) nn Llw b1ology and p.a.l:!c:ontology Jcffcric:,, R.P.S. (I '168) The ~ubpltyl um C.llcichord.lta,
ofmme pred.ltO~"'> of OIV.llvtd mollmc~. Paloeogeoj!raplry, primittvc: fossil chordates with cchinndenn alTi.nibcs.
Pd/a,·or!lllllll<>!twy, l'o~lan>t'rt>h~lil' 4, 29-(,5. (Startish habits Btrllctill of thl! /'Jnlislr A/u.ll' lll/1 <?} Ntrtrmt/1/istary (CCtJIOJ!Yi
and mn-~tartish dt'VIn's in biv.ilvl'~) 16, 243-339. (Proposes tiML 'lJI'}Wi<h' .1rc: chord;1tcs)
,. ( hrkr. J.M. ( ll) I J) A rcm.uk.1blt• occurrt'IKt' of Devonic Jdferit.-s, R .P.S. (197\J) The: tmglll of chordates - a
,r.J.rfi,h lillll••tm ll[ till' Nrr11 \'t~rk ,\/t1l1' ,\/11.11'11111 15 , 44--5. methodolog~cal c~~.1y, 111 '17ll' On~!lill of ,\l,!ior lrll'rrtel>ratc
iBavalvt.....,lJrlish ,1\SOCIJlltlll) Crotrps (cd. M.R. llousL·), Sy,Lc:nwirs Association
C.)\', en. R.. ( 1'181) Lnno1d .mm Jml ban.uu piJnt,arions: Special Volume: No. 1:!, Al,ldcnul Prc:ss. London. pp.
•n rcononuc ho~rvc:~un~ .auJlogy. Htlcoh~t~logy 7. 4t3-77. (What art• the: homalozoam, othcrwL\e calci-
1. 331-4' (lnvenovt" Jnd tntt're,ung) chordares?)
rg Oonovan, S.K ( 19H6, I'I HQ) Pdnutozoan columnals Jdferi6, R.P.S. (1990) The ~olut~ Ot'lldr,'<ystitc< sc<>ticus
IT\lm the: Ordovtcaan ol tht• 13nn\h ble\. pam I and .1. from the Uppc.:r Ordovlclan ofScodand and the ances-
:d P.~lat'<'""'I!'"J'Iuc,tl \,¥1t'1)' ,\I,,,,.,I!,,,,,Jr. try ot chord:ucs .and ~·thmodcrms. Pi11ar•otrtology 33.
Donovan. S K. ( l<lHXJ) The: t".Jrly c:volution of the 631-lm.
316 Echinoderms
jdfcnc\ R.P\ (1'1'17) A ddi:ucc ofrhc cJkichord.Ht'\. Mil~om, C. V. (11144) 'lmur((lllftl, .1 bcntllll' cnnutd from Sl'tl:
Lu/r.u.1 30, 1-10 (Cl.tdJ,tlt .tppn">Jch) the Jura~~il Sulnhuf'c:11 lmu:\lone, GcmLln) de
Jcflcm·, RP S. llru\\11 N.A aud Daley, P.E. (19%) Palaromoh>J~)' 37, 121 'J. Se1ld
The c .JTiv phvlot;,·nr ,,r d1ordJtC' md cchmodcnn~ and Moore, R.C . .md l.l\ldon , L.R. (144.'\) J:r•,•lutronnnJ pi
the onh'111 111 ch11rcl.uc lctr nght .1\)'llllllctrv and btlat- Cfi!Ss!funtum ••f Pcllllc<l::t•t< CmwiU5, GcoloKJcal Sooet'O ln
cral wmmt'ln . lt1.1 z.,,,ft~rc.r 77 101-22. (Sununary of of Anll."nca Spcual P.1pcr No. 4h (M..tJOr monobonph p,
l'alul hurJ.ltl." t hcory) on cnnotd lVOiunon) Scil.1
Joy~cy, K.A. (I <J.!'i9) A \tU.!} of \'.lnJtmn .md relative Nicholo;, I) (I 9'i9) Ch.mgc' 111 tht• ch.tllo.. he.ut-UI(hin Ft
growth 111 the: hi t\h1td ( >rl>ttrwrrtrs. Plrilosopltmll .\/icmsta interprt•tcJ 111 re!Juon to hvmg tlmm. Pit~ er
Traw.uttt•m ·~I tlrt R•'Y"I S.•w ty tJ{ Lllldmr B 242 . St>pftic<~l Trni!Mcticllh '?{ tltt 1~<'}'111 'iom ty C!_f L•rtd,,, 8 242 di
99-12::> (U.t\\tt \tUd} of a l.trge popuhoon !Tom bbs- 347---+37. (Cbmc \.lhdauon of t'volunon m ,\(r"r/N or
totd-nch hnn1um) Nichols, D (197 4) Thc '-\JLc:r-v.l\l ul.tr ,\,tcm 111 ll\ u~g '\c ila
K.m.u.tw.t. K (I '142) Ad.q,t.mmh l,f re't shape tor bur- and foml tchmodcnm Pa/,1t'clll/t>i~I!Y 15. 51' ~&_ Se
rowing .md lo<..nnwuvn 111 \pat.lllgoid echinoids. (Structure .md evolution) to
H:~/,u·•>llft1iol!)' 3 'i. 7 ' I 'ltl. l'aul, C.R.C. (1')(,7) The funcnon.1l morphology Jn I l:r
Kemuck. K A (l'h4) A bronu:tnc.tl study of 1\lraaster mode t•f life: uf tht• cy~toad Ph•llrt>(y.Wtt·s E Billing- c.
mrmr.~rrirrttm .Hid \/ i/s<lllllcrtt>ltrl >IIHII/1'115/J. Plulosophical ! H54. Sy111pc1.sia o( tlw 7-ooh~lltrnl ,s,, irty <!f Lt>mlilll 10 Se:
Tram.lrticlll> <'/ tftc' U.c1ya/ St>lll'f)' <!f l.J.llldmt B 237, I 05-23. (lknd11c crawling h.alm pnstul,1red) <..,mu
375-42H. (Sr:ttl't it.ll t'V,llu.ltllll> of,\ liaruter evolution) Paul, C.R.C. ( 1%H) Morpho logy and fullctlon ot P.J
Ke-.lin~. R .tnd P.wl. C..R C (I%R) New ~pecies of dichopurire pore structures 111 cy~road~. Pctl.tcontc•l,>tJ S1m1
PoroLrtnid.tc ,111J btil'l. rt•nurk' upon the'>C unurually 11 , h':I7-7Jtl. (Funcnonal llll)I')Jhology and ic~ bc~nng K
cnn01d~ ( :clllttlllcllicllll 111 /',!l!'tlll/rlhlq)• C!,/thr Unil•rrslly l!f on cJa,,ificanon) Sutu
,\licfll'(•lll 22. I P. (I nldt·d-ntt·mbranc \trut·tun:') Paul. C.R.C. (I 972) Morphology .md fimcuon of Ju
I<.!er, P. 'VI (I 41r'i) E' tllution.lry treiiLh 1r1 Palal"OZOtC cchi- c:xorhecal pore ,trucnart'' 111 n·stmd\. Palarc>lllc'lc->gy 15 Sim1
I!Oith (c>llmcllc>/ l'.lfc•c>tllcJ/,~1!}' 39, 43(, -6.5 (Very impor- 1-2H. (FunCtiO!l.ll morpholob'Y .md at~ beanns on JJ1
t:mt p 1per: numcrom tllthtrlttnn') da,.~itlcarion) Smit
Klcr, P.M . (197J) A Ill\\ <..,iJun.ln edwmid gt•nm from Paul, C.R.C. ( 197(1) P.llac:o~l'O~phy of prinutlvc: echm· lld
Scod.md. ['.,/.n,•111<1h',l!)' I 6 h51-o3. (Dc-.cnprion of odeml\ tn the Onlo\'u:i;\11, tn l11t C)rtlcwinllll 'iysw~ 75
·Jprtfr.-flllllh, With r('t nlhlrlll ttOII) (ed. M.G. U,h,cct), UmWT'>ItY of Wale~' Pre,s, CudiJf. Snut
K1er, I' M. (1'174) Evoluuun.uy trend~ and their func- pp 553 7 4 (T cm per Jlllfl' :u11trol of dl.)tnbuuon of of
oon.ll 'i!{IUIIc.mcc: 111 the ro,t-P.tlJcuzou: echinoids. echmodcrm~ w1th toldcd-mcmbranc 'rrucrurc~) ne
(P alt·,,molo!-'1 Snuet~ l\lem01r No. 5 ) }tllmltll 1!1 Paul. C:.R C. .md Smtth AB (1<JH4) TI1c earl} rJdiAUOb Smul
fla/rcllllr•/•1_1!}' 48 (Suppl) (1\I.IJOr reference work) :md phylogen~ of cc h11wdl m\\ IJI<IIO.I!tcal Rn~ru~ 59, thc
K1cr. l'.i\.1 (I 'IX;!) l~.1p1d evolunon 111 echinoid~. 443-81. 24
Pa/,Julllf•'"'l!)' 25, I 11 (SJnd dollars) Phtlip, G.M. and Fmtcr, R.J. (1471) Mmupaarc: ternary Smn
Klcr, PM .md C.rmt. R L. (1%5) Echm01d distribution echmo1d\ from \outh-c:,l\tc:m Au~trahJ .md thetr Zu<>- \e:
,md h,1h1t' Kt y I .H ~o Cnnl Reef Preserve. Florida. gcographic sav;mfit..mce p,,f,u•t>moh~y 14, o6b-'J5 Snut
S1111ll~>•''"'"' \III,·,JI,IItl'<'"' Ct>llcctwm 149 (6), 1-b':l. {L1fc (Dcscnpoon of m.an.uptatc echmotd~ illmtrar..:d h1 Pa
h.1b1t' of 'evcr.d rncHk·m 'pccic') stereo-pa1n., and contro l~ of d1stnbutmn) Srlllt.
Koch, I) I ..tnd ~mrnple, II.L. (IW>H) A new Upper R.amsbottont, W.li.C. (1% 1) Briti.1h Ordc111icia11 Crinoidca. de
Ocvoni.tn ,·y~told .ttt.cdH:J ll> .1 d1~cuntinuity surface. Palaeontogr.•ph1cal Soc1cty, London, pp. 1-37. Pa
/1111'" Cmfcwu1l .Stm•t')', R~porr, ofl•wc,ttgJtions, No. 5. Regnell, G. (1 %0) ' lntcmtcd•ate' tonm 111 early PllacozOJc Smid
(EdnoJ~tl'mtds .1nd rV'otO rd~ 111 htc pmition) echinodenm. Prorcrrlil(fl' '!ftlw 2/sl lmrmational Grolc',llicn/ ex
M.ll urd.1, I> U ( llJI•'i.1) Tht' tunLtional 111l1rphology and Con{/n•ss, Ct>pl.'llluw·n 22, 71 -80. (Taxonomic problem gt:
\tratiKr.Jplm J,,tt ihur1u11 ni rht• Mi~,l~~ippian blastoid raised by ~omc pccuhar echmodcnn groups) (cc
genm Owpilcltn/1111 /<111111<11 of PnlnlllfcJfogy 39. 1045-96. Ro,enkram, J) (1471) /ua Sc:dirnc:ntologJc und 0'
(Mmpholoh'Y and dyn:tnm:s) Geulogie von EchmoJcrlllcn Lagcrstattcn !\'cut• Sm.itl
MitturJJ, DU. (IW1Sb) 1-iyJroJynaanin of the: ..fahrbucft fUr Ct·ol!ll!lt' rmd 11alilontc'h!I!Jt' 138, 221-SK. ho
Ml\\t\\lppun hl.t\tOJJ ~~Ill'' G/ol•••bla.sws. Jmmwl of (Variou~ kmd~ of cchmoda:nn beds) ·W
Jlalecmul/,1'1)' 39, !209-17. (lnh.tlanr-c:xhalam Row Rowe, A W (I H9!J) An .lnJlym of thl;' ~en m ,\f1mutcr, ~~
rhrou!-th ~ptr.lde)) detemuned b~ n~1d 7tmal colb:ung from thc zone of
Macurd.1. D.U. and Ml'Vt'r, D [ ( I'H4) Fcedmg posture Rltynehonella nw1rn to that of 'll10a<tt'r (<Jral~'(tiiiiUI"
of mod\:m ~talked l'nrwtd\. '\,1/tllt 247. 39+-6. (Life Qumtrrly Jc•um.tl of the Gt·ofc';l/ir.ll ~t>crrty 4 Lm.fo, )5
h1bits ol rhcoph•hc cnno1d\l 494-547 (Cb"1c evoluuon;~rv \tlld))
Bibliography 317
Sctbcher. A. (1979) Comtn1crion.u morphology of sand Smith. A.ll. and Ghtuld.j. (IIJH2) Roles for holes 111 ,,IJld-
doUm. Hlll't>biol.>',!)' 5, Jl}J - 2.:!.:?.. doU.m (l:.chmmde.1): .1 rcv1cw of lunu!C' tuncnon and
St'i!lcher. A. (I YlJO) f he ,,md-dolhr 'yndrome; .1 poly- evolunon. Palmbiolt>(/)' 8. 242-'i.)
phylenc comtntll ic11l.ll hn•.1kthrough. m Er•t>/lllwrwry Smith, A.D and Hollinl-('vorth, N.T .J. (1990) Tooth
lmltwaii•IIH (ed . M . NitclJ..I), Univer\i!Y of Chicago ~trucrurc and phlogen}' of the Upper Pemuan c:chmmd
Prc .,, <.. htc.lgo. pp ..D 1-52. ,\liMtt.ms kq.(rr/111~1 J>ro..·l'cliiii(S <!I tlu \'<>rh/urr
Stdachcr. A., Dm7on·w,ki, { ; , and ll.mde, R . ( 1968) Grolt>_~t<•11 Stl(l!'ty 48 . 47- W
Fonu md functmn ••f tht• 'tem in .1 r~eudoplankwnic Smim. A.B. and Paul. (RC {19H2) Re\i,ion ufthc
COllOid ( XlrlllniiiiS) . P.!ltlt'<ll/ltlh~I!Y 11. 275-82. (HO\\ class Cyclocvsrmde.1 (l:.chmodemut.1). fllu/o_cc>pllical
dJd pertccd} pre,ervcJ cnno1d~ come w he in posinon TramartimH oj rh, Rt>}'tll ,\,>(lfl)' t!{ IA>IIdt>ll R 296.
on the l'J floor?) 577-8~. (Exqui.,ire .Uustr.mom)
scthcher, A . Rei( \V.-E. .mJ Wt.~tpha1. F. (1985) Spnnkk H.J. (I fJ76) Cl.mtfic.mon .md phylogcm of pcl-
Sedimemolo~H.al, ccolob'lC.ll .md temporal pJttcms of matozo.m echmodc.:rm~. Systc'm<~tit z,,oft~~y 25. HJ-41.
ioNI L.lgcr;t:ittl'll, 111 J:.\ lrtiiiTdiii<IT)' l·o.;si/ Biatas <llld tltetr (Bam of cl.mltic.ltton uwd her~·)
Et•oluncmary SJ~IIitr••"'"' (c:J\ II.D. Wlutungton and S.C. Spnnkk ]. ( 19Rtl) An overVIl'W or the tosstl rcl"ord, Ill
Conw.w Morns). fl/uh•.~c>p/uc,,/ Tr,msactlolls of tltr Royal EclrinodL'flll.\, J\'0(1"~ {or 11 Short Ct>IIW" (etbJ. r. DU£ro and
Sooery 4 Lo111lo11 H. J 11 , 5 23. R.S. Boardman), Umvcmry of I ennessee Studic~ in
Stmms, M.J (I 91N) Hritislt Lt>Wt'r jumssic Cri11oids. Geology No. 4, Knoxville, pp. 15-2(1.
PaiJconttlb'l",Jphil.tl Socil'ty Monograph Sprinkle, J. (1982) Echi.J1o1.h:nn Faunas from the Drormdc
Smum, M J. ( I 1190) Crtnolds, in Cllt>lulwllo1T)' Trcr1ds (ed. Fom1ation (Middle Ordovtrtan) of Oklahoma.
K.j McN.1m.u.t), lklh.wc:n, Boqon. pp \HH-204. r lllitll'tsiry <!f !l..tlltS/1.( Pah•t>lll!l/(l~im/ Ctllltrillllfhllls
Stmms. MJ. (I 'NI) P.mt'rm of t•vulutton anwng Lower Mtmo,~raplt 1, 1- 3119. (Very detailed Lomplhllon)
Jurmtc (nnot<k Hi.~ttm(,J/ Bi.•h~~}' 1, 17~4. Sprinkle, J. ( 198.') l'.mam and problem~ 111 echuwdcm1
S.:mm~. M.J .md ~l'\,1\Wpultl, l• 1) (llJlJJ) Thl• ongin of cvoluuon, in Frlllrllld!'nll Stt1dir.~ (cd~ M j.111goux and
arocul.ltl' rnnmds 1'<~/a,..,,/,>lt>.:y 36. '11- 109. J.M. I awrenc<"), B,llkcm,l, Rmtcrdam. pp 1-Ht
Snmh, A. B. (I 97H) A functhm.tl d.l\stltutmn of the: (OfO- Stephcnson, D.G. (11}63) The 'PIIll\ and ditfu,c: lL\cwJc,
nJI pu11:s uf ~~~ul.!r edtlllllllk 1'<1/,rn>llft>lt•.:y 21, of the C'reraccon' c:chmmd l:'c/lillocol')•s .<mtat<l I c'ke.
75<)-.X<J (lllmrr.ltt'd tl.l~'•c) flt~farmllt>lo_~y 6, -l5H- 70. (runnion.ll morphology of a
South. A U ( llJHO) Tht• slntllllrl', funttton .tnd evolution ,hallow burrowa)
of ntht feet .md .uubul.tu•tl pure~ 111 i~TCgular echi- Stokt.~. R.B. {IIJ77) Thl cchrno1d' ,\/J,.,.a;to and cpw.sw
nmds. Hl/,n''"'"'l'l!)' 23, .\11- HJ (St·e text} from thc Turo111.111 .md ')cnoman of Somhcm England.
\mith, A B. (1'1)-11) lmpht.mnn' nf l.mtem morphology for P,r/at't'lll••h'llY 20. Hll5-21
dtc ph}lugcn} t>fplht-P tl.tc:owrc: nlunoitl' P.1lae,mtofogy Stunner, W. and llc:rg.tro111. J (1976) The Jnhropods
2•'- 779 ·HO I (lla'i' tl.>t re\ ~l·J clt.,~ification) .\fmu:ttl.>tcr and ,.,,,/tiJIII>hl lrolll the nn·oman Humruck
~nuth, A U ( 19H2) Tomb ~trut:turc oi the: pygastcroid Shale. P.rliiollt<lh~IJ<"ht'.< /.fitrlllg 50, 7K-I 11.
\t.t-urchm Plt•sir.hillll<. J>,,/,rl'•'"'''h'I!Y 25. H91-{). Ubaghs, G. (19'i3) Cntll>tUe\, 111 Trmti- tit• Paltwllolt~{!ie
Snmh, A.B ( lli!Hb) CINifit Jtivn of the Eclunodernuw. 11 1. Onycltopltorc>, .-I rtlut!p•>tln, L:clmwdt•rmc·s. Sttllllll-
P.1/act>lllc>ltJ.I!Y 27. ·U I 59. c/Jordes (cd. J. Ptvcrc.lU), M.mon, P.1m, pp. (•SK-77.3
Smith, A B (I 9H5) <. .lmbnJn l'ktahern.coan cchino- Ubaghs, G. {1971) Dtvt•r-Jito et 'Pt'U,tli\Jtiou de<~ plm
denm .md the t'.1rly di\'t'r.>iht.ltion of cdrioasteroids. ancicm cchinodtTJilC:\ yuc l'on lOnu.U~~e. Btolt\l!iml
P.liaetJIIft'/'~1/)' 28. 715 5lL RePJCII'S 46, 157 200. (C.trly radi.ltlom of cchinodcmH;
StnJth, A.B. ( IIJHH) Fo~s1l cvKknce tor the rei.ltiOmhtps of di-sclaims th:H carp•)id' .1rc d1ord.uc')
cxtanr cchmoderm rl.1~~e' .tnd rht·tr tunC's of diver- W!gJlall, V.B. and Simlll\, MJ. ( 11)90) p,cudophlnklon.
gene<.', 111 Etlri11t>dc·rm Pilylt~(/t'IIY tllltl El'tlllltilliiiiY)' Bio/o.(/)' Palaccmtolo,(!y 33, 359 7H.
(cds C.R.C. l'.tul .md A U. Sm1th), Clarendon, Woods, I.S., and Jc!Tcnes, R.. P ~ (1 1)1).2) A new qem-
Oxtord. pp 'IH-101. group chordatl' lrom tlw I OWl'r OrdoviclJn of South
~mtth, A U ami ( :Jill'llu, I. ( Jt.)l) I) M1ddlc: Triassic Wales, aud thl· probk•111 oflornmoLilln m boot-,hapcd
holothun.ln\ lrum rmrthtm Sp.un P.rlac,lllttllt~r:y 34. cornutes. fJnf•I<'OIIftll()',!)' 35, I ·2S.
49-76
Graptolites
Abund.mt ~rick-like fmsih are <;Ometilm·~ found m much fine derail is lo~t. Even so. 1r 1s surpmm~ hm1
Lowe1 p,,!Jeozo~r bLlck ~hale~ or more: ra rely in much remai111ng structure can soml'timcs be found
other .ugJIIaceous rocks of tht· same age. These an: t•ven in cru~ h ed and carbonized o;pecuuem.
the brraprolnes, so t.lllcd from their re~emblance eo
wntten nnrk~ on the ,h.Jk (< .reek: A.t9o~ = to
write; fP<lEt V .... 'lone). Tlll'y may bl' stratght ur
Order Graptoloi deo
curved sometiml'S \piral in fom1, smgle-branc hed,
bifid or many-b1am he d. When preserved in tht· Soetograptus chimaera (Fig. 10. 1)
most l '>111111011 ' \ J\, 1 e. flartem·d in 'hJk, one or This Ludlovun spec Jl.'\ (Urbam·k, I<>5R) 1s hmnd
both C:llgcs appeat st•rr.lted hke .1 tiny ~a\\ blade, but only in gl.1nal cnauc boulden, onginJily dcn\'cd
oth<.•rwl~l' nor mud1 \lructural detail m:~y be visible. from rocJ.,, under the Ualtic Sea JIH.l now 'LJttt·rrd
ThL•st• t(mils are of smgul.tr unport.lnct· 111 estab- over the North GemtJn and Pohsh plain: 'Pl uncm
lislung .1 sttatigraphtca l Ullll'\l tlc for the Lower can be isolated from the rock wJth aud.
Palaeozmc .md h.we been of zonal value \lllce thl' As 111 all graptolites the 'skeletou' of Sorr,\~IIIJ!IIIJ
mid-nmeteen th n·ntury. Though much of the consists of a series uf hollow 1nterlmkcd tube,
srrat1):.'1',1phical work. tlut h.l' bel.'n done 1s based ronsm•ctcc.l of a thm shcet-hke nuterial kml\1 n ~~
upon 'f enmem prL'\nveJ as fl.nrened urbnnized or periderm.
kaolinimed film' (L'ven so, thL'> are ~nlltdenutiablc), The tiN-formed p.trt of tht' brr.1pt0loid •~ till' sic-
the ,uJJtomy of w·•ptoutes t.Hl only hl' properly ula: a conical tube with irs aperture pointmg down- f•gure
undL'r\lLlOO with rl'l't·retll'e to the rdauvcly rare wards and terminating at m apex 111 a loug. hollm\, port rer
spec11m m preserved in tluet> dunL·nsiom rod-like nema, extendmg well bevond the upper
hmir of growth of the \ICula. Tht• '>Kula 1' JtvHkd
mto two parts: ,m upper prosicula anJ a lowc:r which
metasicula. The prosicula has a th111 proxuual p.m, l'dgc t
the cauda , and a broader distal con us has Jll urn l· Jcvck
'Graptohtc\' i~ .1 vcrnacubr tL'nll for members of ment of longltudmal :111d ~piral striae ; the mctasJC- bud g
Class Graptolithin.J (Phylum Hemichordat a). They ula, like aH otht'r pJrtS of thl' graptolo1d, 11 fi rst of
arc coloniJI tnJrinc uwcrt\'bralL'' now bdil'vcd to ornamented with well-marked nngs reprc~l'ntmg '>IVC tJ·
bear a distant aJlimtv ro 'cno.:braces, though tor a pl·riodic, perhaps datly, growth mcremeut' Jnd uutil a
long tunc thq Wt'rt' regarded as bcmg allJed to known J\ fuscUae. FuseUar tissue constst., ol tlun lll dir
coral\, \\ hich haw .1 much ~uuplcr gradt· ol"btOiog~ half- ringll or compk•tt' ring; of ~keleral m.1h:nal turc~ ~
cal orgamzation. Wtthin C l.m Graptoltthuu there ~kic ked o ne above the other :md uniting alon~ provid
arc scvtrJI orders, of wh1ch only two arl.' of any zigzag sutures. In the: \!Cub rhe ~ururc on one \Jde The
real unportance· the Graptolo1de.1 (Arcnig-Prag ian) h.1~ the duracterisnc z•gzag fonn .. hm"n m all gT.lp- \ lth cl
and tho.; Dcndnmk.1 (Trcm.ldoc- carbomfcrou ,). tohtcs; on the other ~•de the hJ'>dbr half-nn~ arc other
£.-xampks of really wcU-prc~l.'rvL·d genna of these JOmed to .1 ~tout rod (the virgeUa) which pmJnl!. throug
two m am orders will be descnbeJ bdow, but it has below the aperture and may curve !.hghtly unJc:t 11 thl' par
to be 1111de ckar th.lt graptohtc.:s are rardy preserved From the s1ntla thl're b'TOW up a number ot LUp· thL di•
like thi~; 111 the nomJJI cour\l' of prcscn'JliOn very like thecae (smgular: theca). the tlrst-fomwd of embed
Structure 319
'pi 10 l Morphology of Soetogroplv$ chimaera. (a)loteral view; (b) frontal view; (c) $lrudvre of theca and common canal with
mtemoved (x 40 approx.) (Modified from Urbonek, 1958.)
h bl.'~n' .H J prima ry n o tch m tht> gnlwmg .ls the virgula. The: partioon wJlls themselv~ Jre
~· nt rhr sicul.\ ,\lld on m Vlrgt>llar stdc, bc:fon: the double-lay c:red struLlun:s, the wall b(:mg secrc:tc:J
lopmc:nt of the !alter •~ complete The initial fi·om both ~ides (Fig. l 0. Ic). Each thee\ has twu
Ill .;row~ up from tho: pnm.1ry note h to form the p.1rts: the protheca , proximal to the nema, through
nfrhe thec.lc. whtch an mm b'lH'' nse to ~utccs which the con1Jnon canal run\; and J m e tathcca.
thtcat' incrc.1sang in ~izc :twJy fmm th\.' sicula which is the cxtern.tl pan divided by p.uttrion walh
about thl' htrh or ~i-.th rhec.t, 1trer which thc- Jnd pOS\l'S.\tng apertural spine.:~. The b'1'0Wth lint's of
dimcn iom fl'lnall comrant. rhl' thecal .lpo:r- thl' thecae tend to be more wtdcly spaced towJrd
pmnt up\\ .lrds but .trt· obhqudy mdmt>d Jnd thl' .1perturc:, presumably sigmfyrng a pc nod of fJ~tn
vt,lt\l With paired apcrtural spine~. growth towards the end of thc:cal form.ltlon.
llvirv ot the ftf\t-fomll' d theca conncct5 ~ince gr.lptolites were colomal animal,, with thc
dur of rhc \tcula through ,t fo ramen ..111d all mhabJtants of the thecae and the 'JCub (whatt>Vl'r
1 thccJe hkcwi~t> lmk up wtth one another they may have been like) linked to one another
l!ih a common canal dmc to dw nem.a, smce through the common canal, it rs prc\umed that tood
r anon \\,111~ of the thecJc: do not extend trom lilught by one mdtvtdual would bl' mgested .md
r.il pan l1f the nc:uu. Wht·rc: the IH.'mJ lS 'bared by the: whole colony. Tlus colony i~ known
.ldcd 111 the wall of the graptoloid it 1s known ,,, a rhabdoso mc (.m older and less correct name ts
320 G raptolites
p olypary) ..md the branche' .trc stipcs. S,u•to.~raprus Th. 1' •~ fin.t directed up\Hrd,, thl·n I\ t\\1 d
ha~ onh one stipe. m to a recumbent S-shape ..md tin.tlly ~ll\\~ acrth1
Thnc ., no direct cndL·nu: ahom what sort of rhc from of the stcula, becoming wddnl to it <Jnd
<uumal \\J\ ft)und in each theta . hI\ J"umed rhat in fonmng ,1 crossing canal. Cunoml} enough. the
c,1ch thc~J there was a zooid ,\ 'ilmplc animal with fitscll.1e th:n torm the cro~smg c.mal .trl' laid dol\ n
'ome \Ort ol tood-g-athenng ,tpp.tr.ttm, possibly a not only at the growing aperture ol th. I', but .~11•
lophophore - and tlut all th~· zo01ds were con- a~ J tbnge .tlong the ,tlrcady-tormed tubulJr .. urtJu."
nected Internally by tiS\liC rJte pyntlz~.;d remains of of rh. 11. The two part~ of th. J 2 ullltl' tu lom1 ~
retr:Jctl·d graptohte zootd~••md tube~ connecting tube npt•n :tr borh ends. ~o th.H when lully tom11:d mt
them. have been fOtrnd 111 tht.> Au~tr.than .uus05'Tap- th. I' termin:ltes 111 an upward rutVl', \\ h1lc th. 11
ud Pst~r,lptus (R1cbrds aud ~talt. 19H~). These. amc\ on the oppo~ite side !Tom th. 12• fhen tit 1
howe\ l'r. re\·e.ll no detatl' of tl'JH.ldc' or other fine e.rrow' up from th. 2 1• tom1mg .mother crO">Sifll:
o;trunure. The nature of the zoo1d~ will be con- can.tl. LlkC\\t'ie the hter-fonned thec:1e .111 pm m
\ldcrl·d afrer the atliruuc, of tht• gr.tptolitcs an: dis- from of the ~tcuh or the nenl.l, e.tch nml" fOnmn~ a
cu\\ed m more derail (~ectton 111.3). cro.,~i ng l<lll.ll.
The mode of growth i~ marked by thl fu~ciiJt' et
Diplogroptus leptotheca (Fig. 10.2) l~ig. IOA) so tlut rhc dcvclopment.ll h1~tory IS ckar (d)
Dipl!wrnpms ~~ a CarJdonan gr-.1ptolo1d w1th two from them, bur it is more u~cfiii to ht' .1blc t('l ~cc o~ll
Stlpl'\ 111 the rhabdmomc. Uoth of thcsl· arc united the MJgc' in growth !Tom the devdopmcntJI scne~
b.td. rn back wlth the nemJ m bdween: the rlub- here 1Umtrarcd.
dmome " \,lld to be scanden t .md biserial. Thi'
gr.lpto md 1\ <;ffi1Ctllnll} morl' lomplcx than
<iat'/cl.~t·lfJtlls, c'pccJJllv 111 the proxtnul end. J.e. the Order Dendroideo
\ltu!.t .md the fir>t-fonm·d ther.1c wluch surround 1t.
Though the \tn1cturc ofthl· proxtm.•l end JS dtfrl.cult Tlw dendrOid~ are the mmt .1111: icut of .1ll lh
tn \>\'Mk out m mature ~pcCIIIIl'n<., mdJvJduals in aU graptohte' .md were apparently .llllC\tral ro th
b'TOwth ~rage~ haw been found, ,1nd thC'se can be brr.tptolmd~. How{"ver, their morphology 1s tOll·
.1rran~cd in .1 dcvclopmentJl ~~.·ril'' ~howmg exactly plcx, .md they have various org;lm \\.htch ,11( not
how thl.' different tlwcae orig11I.ltc :md form pm~"''~cd by the descendent ~.tploloiJ\. Nomwl) ,;
(llulnun, 1945 l 04 7). dendrmds .Ire presetved, as ;lfC mmt gt.lptolmd~. ~ I

rh~ ,jllll.l i~ bro.Jdl) Similar {() th.lt of Sat'tt~l(rapws compre"1om m ~hale. but rare threc-Jum:mwull
bur Ius t'\'.O apertural \pmc,, and thl' lllltt.t! bud .trise~. spcctmens can be ISOlated from the Jock or rudtt'll
not b) the fonnation of J notd1 in thl· growmg edge b) me.m~ of thm section, so thetr deta1lcd morphoi-
Figure I
of the Sllula. but by the n:snrpt10n of J toramcn Og} 1\ qmte \\'ell known. from Bul1
lugh up on the meta~icul.1 bl'fon· huddmg. This sys- The b.l\lc ~mJCturc of dendrotds \\,\\ worked o 1
tem ~~ ,tcntally nH)rc nonnal m ~:,.rr.•ptohte' than the in the l.ttc mnctccnth centlll)' by the S\H'di~b
~pcnahzcd mode of development in Snl'to,iiraptlts. pal.teontologi~t C,1rl Wm1an, m::unly !Tom sen.ll
uolot~
fhl rhC'cac are numbered 111 pa1rs :tecording to a 'ectiom. 1t w.1~ btcr clabor.tted b) 13ulnuu
Wrtl'~ SI
ronwntwnal notariou. so that tht lir..t-formcd theca (l945- IIJ47) and most p:Httcubrly by Ko~7l0\V\~J
.111d , wi
IS th. 1 1, the second which origin.Jtt·s !Torn it is th. (J 1J4H), who h;l\ described in great derail ·' n~·h ~nrl
t!O\t'd (
1',the third 1 1, the fourth 2l. l'lt'. In a bi~erial grap- varied t:wn.t of dendroids and other ~raprohtcs tium
!lm the
tolold \UCh as D1ph\~'''P'"~ the tlu:cae on tme s1de arc the Tn:m.1doc of Poland.
~''lll.1lly
numbc.:red 1 1, 1 1 • 3 . 4 1, ••• and on the other are
numbered Jl . 1!, Y. -1 2• •• • • autothr
Dendrograptus (Fig. 10.3)
In Dtpftl~rt1p111S th. 11 ~'Tow~ dm\ n .llmost vcrti- Dcudn~,gmptm,
'"hich •
J..nnwn !Tom the threc-dunem1onJ
kvll T
olly to about rhc level of the ~llUI.tr .tpcmtre and j, m<tlenal ,wdied by Ko\zlo\'.~k1, l'XIubm rhe reprr.-
~t.IIH \\
then ~harply hooked up\\Jrds to tcnmnate with an sent.ltJve dendroid 'tructurl' adnmahly lb
b~:twec t
.1pcrtural spmc at tts hp Th. I' .Ut\C\ from th. 11 inverted conical ~icui.J ~tood upnght upon the ~
\0 th.tt
n{"ar the s1cular foramen, bcfort· rhe growth ofth. 1 1 floor wirlt its ;lpical base ex-panded 1nto a holdt• t
'lolothe·
is complete. At a po1 11L ::tbout halfway up thl' ~it·uLl arise~ th
h11hccac
Structure 321
(1) (b ) (cl
prosoc:u a---HH
apertural
spines
th. 42 th 4 1
(d) (f)
th . 3 1
th 22
th 2'
th 11 th 1 1
th 2 1
t
y
'
tl
d
Figure 10.2 (aHIJ Development (>< 30 approx.) and (g) adult morphology (x12 approx.) of Diplograptus leptotheca. (Modi~ed
lrom Bulman, 1944-1947 )
lt
h
ai' stolotheca , winch is t•quiv.!lent to the prothccal ously looped or coiled. 1 he autothecae are very
il l
1eries surrounding the common canal in Saetogmptu.s m uch larger and expanded upwards, each terminat-
ki and, with daughter \tolorht:cat•. fonns a continuous ing in an outwardly mclined but often elaborately
!d do~cd chain all tht· way up the rhabdosome. From sculptured aperture with a median tongue.
111
rhis there 1risc rwo kmds of thecae at successive and The arrangement of the amorhccae and bithecae
.qually ~paced nod.tl pomts. Thcst• are the large follows th e 'Wiman rule' of alternating triads; bithe-
autothecae .tnd tht· sm.tller narrower bithec ae, cac usually arise at alternate nodal points on opposite
~~;hich always come otT the stolotheca at the same sides of rhc mam stem and arc earned disral to the
lal lewl. Tht· bichecae in Dc11dr<1\!raptm mamtain con- autothccac. When the rhabdosome branches, the
e- !Jll( width, but t'.Kh 1\ nonnally looped over stolothcca splits and the Wtman rule alternation
he between the ;motntt•J autotht•c;t .111d the stolothcca, continues on each daughter stolothl·ca. In some
.ea 10 rhat the apcrturt· j, on tht• oppmue '>idc of the species the auro- and bithccac ansc m sequence at
angles of90° rather than umo from the one below.
\(,
olothcca eo the ori~rin. In other dcndroids the
he btthecae may be ,jmplt• str;tight cylinder' o r be vari- Within each rhabdo,omc there is an mternal
322 Groptolites
la) lbl
cownng and enveloping the earher tht·r.le. (lt
u~u.lll) prc,cnt in graptaloids a~ '"dJ bur I\ wry thu
and only clc.uly ~een with the ekcrron mlcrO\Wpr
Dendro1d cortical ris~uc ~~ much thicker •md I\
clc:.trly v1sible.) Such cortical ~heet~ fonn the hold-
fast wluch afftxc~ the colony to the \CJ Aoor.
autotheca 2
'--'"'-..(-.IH-+-->l-- botheca (nl -+-f-++- Pre servation and study of graptolites
The three genera described above arc ut \tlllll' ofth(

best-preserved graptolites known. Thl') .tU come
from fine arg.Uaceous limestones. ,md though 'pen-
~-+-- autoth~ 1
mens m.l) be very abundant locall)' 'uch orcut-
4---i-+- botheca hi -+-+--"!V'
renccs are rare. Thrce-dimemiorul ~pt:timcns ot
such .1 kmd must be freed from the rol k ~o that tht ~
cJn be studied in the round. The tOt!...~ .m~ mltl.tll}
broken into fragments, treated ftrst with hydrodllo-
nc .Kid :111d then with hydrofluoril ,11;id, Jnd aJttr
washing may be picked out and indiv1Jually tr.tn~
fcrrcd to a concentrated nitril· .tnd .md potJ'>'ium
chlorate solution wh1ch rende~ them tran<.lucent
Thc..•y .m: then dehydrated in a corKentranng .1kohol
\l'nc\, clc:..lncd 111 ""Ylol and finaUy mountt·d on ,IJd~
in C..an.1da Balsam. Such specimem are then ui cbr
Figure 10.3 Morphology of the prox1mol end of o dendroid, brown hue and V1rtua1Jy transpilrent.
Dendrogroplvs communis: (o) frontal view; (b) reverse view - the With the light microscope. at rd.1l1vdy lo" liiJL!·
cut·awoy panels show the stolons (x 40 approx.). (Modified from
Kozlowski, 1948.) nificanotH, Individual thecae ,~nd thl.' ~itula rnav b~
wry clearly marked with fi.1sell:!r nng~. Fxnlltnt
detail may be \een with scanning cll.'ctron mtcrot.''la·
rubul.1r 'Y~rem ntnmng from tht• 'iulla all the way phy (Fig. lOA).
.1long dw 'tolorheca and the bJ\l of t'ath autothcca Such fusclbe ;trt· occasiOnally seen even m flat·
and btthec.L Tht' stolon system " onl) found in tcned specmtcns, though in typiCal 'peomrn~ ol
wdl-pn.•,ervc:d spc:omen,, but Wun,1n detected it in Sactt~~taptus chmraera the carbonr7.ltrnn of rhr pend-
hi~ \en.rl ~ecnon' and beheved Jl lO have carried an enn ha~ often gone too far to have tOrhrrvcd the
mternal ..lpparatm such as a rH.:rvou~ sy~tem. nticrmtnrcrurc, and growth nng-. nuy lw lmt. yrt
Koszlow~ki confinned tlm ~ugge,tion when study- the thrcc-dimcmional ~tructure of the rhabdmome
mg modern pterobr.mchs, \\ hllh ,ue sm.Ul, tube- is sllll preserved. When carboni2.1t1on 1s complete
dwelling, marine anJmJls, for they c~rt· apparently tl1e graptolite may be isolated from the rock., a~ mth
the neare~r ltvmg relatives ol grapwlites Jnd have an S. cltillltll'fa. Sometimes, however, carbol117.1tlon 11
equ1valent stolon enclosing jmt such ,1 nervous sys- mcornplcrc, and attempts to &cc the gr.1prohrc c;~use
tem as W1man postubred. No \tolon ~y,tcm is tound 1t tu cntrnblc when extracted. Tht·n the only \\ 1y to
m the Gr.lptoloidea, but 1t 1s de\cloped m \omc of clund.tte the structure is to nukc clmeh ~pJccJ ser-
the other nrders, especially tht· ~tolon01dc.1. where Ial 'cctions through the \penmen .lftt'r embeddm~ 11 Figure 10
therl' are Vt'l) nun~ pecuhar and 1rn:gularly branch- m plaster or resin. cutting the 'ecnon' nonnal to the SEM phol
mg stolom. long axis of the rhabdosome (Fig. to ti) If the stm('- width at h
The fmcll.lr rmue of dendrOid~ 1s broadly siuular individual
turc ~~ very complex, especially 111 the rcg~on ui the of Or Petet
m rh.11 of grapwloids. Howt•vcr, there i~ also situla and the fin.t few thecae (thl· proxm1JI cnd),lt
Jnother kmd of pcridennJI m.m·n.1l: the cortical is oftt·n u~cfi1l to build up model~ in w;Jx from thc~e
tissue, wh1ch i~ tud down 111 tlun, llat bandages ~ectiom, by superimposing one l.1yl·r ,1ftcr ;mnthc-r.
Structure 323
(b)
1-
..-
!r
ill
t. In practice tht· wax 1~ cut to show the internal struc-

1 ture of the graptolitc preserved as a core, for in this
es way rclatiomh1ps of the thecae arc more clearly vi~
r lbk (hg. 10.5).
Not mfrequcntly the pendenn is completely
~- dc~troyed, but an mternal mould of the whole rhab-
::>e dosome may remain. retaining the thrce-dunen-
nt ~iona l organization <lS a pyrite core. Sometimes these
a- cores retain c.ubonized peridenn on the outer sur-
6re. bur this tends to Rake off. <;uch preservation ts
l t- common with speumens preserved in black shale,
of the pyrite being derived trom org.1nic matter decay-
d- mg in an anoxit environnll·nt. A~ with the wax
he model the penderm has gone, hut the shape<, ,md
('$ convolutions of the thecae rem;tm m theu ongmal
ne relanonships, ,md mtcmal wall' or ~epta may be
! tt: clcJrly visible. l lowever, the mmt conm1on, and
ith unfortunately the least revealing, of the various
IS
kinds of preserv,\tlon is that where the rhabdosome
l~l'
has been compktd\ flattened and preserved onl)' as
to ,1 compression: ,1 larbon or alummo~ilicatc film on
er- tht• rock. Even here, however, thecal shapes arc
~ it ~~vte 10.4 Cltmocogroplus inuili: (a) a montage built up from usu:1lly retained so that the fossi ls can be readily
:h e ilvl photomicrogrophs showing bonded fusellar tissue (total
~ at top approx 1 mm); (b) corttcol bondoges secreted by
idt•ntttied, and It is from such 'pccimcns that most
lC- graptolite speetes and faunal sequences arc known.
~uol zooids {x. 500). (Photographs reproduced by courtesy
the Or Peter Crowther.)
• it
csc
!Cf.
324 Graptolites
1 he ultr.htrtlt'tun: ot tht· gnptolitc..· pcridc..•Jm,

which lu~ now hcc..·n mtuNvcly ~tudicd (Urbanlk
and J'owe. IY74, 1«:175). \hnh ltght upon tlm com-
plex coptt'. I ratNni"IOn dt·ctron nucro~.Yaphy of
exccpuon;lll) wdl-pr~~l.'rved ~l .lptolOid and dw-
drmd n1.1tcn.11 d1ow' th.H the fil\dlar .md corneal
ri~me~ arc comrrttl'rcd of 'ic..'\'cr.11 kind~ of (tbnc (hg
10.7).
Furrhennore. rht• C'l;l'ift•ncc of corneal tis\uc m
graptolmd\ ·" \\ell ,1, dt·11drmd' ha'i been conhrmed.
'though 111 the bttcr tt m.1y bL· wry thm. Cortical
fabric, wh1d1 makt·' up mmt of the cort1c.1l l1''ue.
comi\t~ ot· rlmd) p.lckt·d p.lr.llld fibrils . In mo1t
brraptoliws rhc~c tihril' arc .1rr.mged 111 'ihort. ribbon-
like cortical bandages deposited one after the
othc..·r in ~ucce~\ivc..· layer-.. This ~vc' the impre"wn,
in thin \LTiion, of' ~t;tcki ng in succc\\ivc layer. wtth
alternating uric·ntation. The manner of p.tcking .md
the 'iize of the wnit.d fih n ls is 'itrikingly rcmumccnt
of the :.ppcarame of th e protein collagen. which''
an imponanr strmtural component of many animal
tissues, and this close rc\Ctnblancc mdccd indicate\
rhat the original pcntlcrm mawn.U wa~ coUagt:n
Figurt
Fusellar fabric , though probabl} aho collagc- showi
uous in nature, 1\ made of tibrils 111 a more open up fro
mesh work. The bulk of the fmcllar ti\\UC io, mJdc: of
thi~ fabnc. There arc aJo,o at least two kmds oi 11<':1-
fibrillar '>hct·t-lih· maccnal. Such sheet fabrics are
Figure 10.5 Reconstructton of port of o Climocq~roplus stipe.
The upper and lower zooids ore withdrawn instde the thecae bot found dclmmu1~ parncular la~Cr<i w1tlun the corul":l!
with extended lophophores, the central one ts extended ond is tissue. fornung a tlun external coat on the ouhide of
'pointing' corttcol bandages on the outside of the rhobdosome. the fu~dl,1c, deposited ,ts a thin sccondJr) ~hret
(Redrawn from the complete model of Crowther and Rickords, inside the thec,le. and occurring inside the \tolon
19771
~heath.
All the~e kmds of fabric llll) m:cur in either cor·
tic:tl or fuscll.1r ussue, but o ne kind is usually pre-
Ultrastruc ture and c h e mis try of domina nt. Thm cortical tissue is largely corttcll
graptolite p e riderm fabric, though sheet-like fabrics form layers withmtt
a~ well as outer and inner limngs. Fusellar tJssue rna\
Fat o~ long lt1ll1.. tt \\,1\ bdkvn{ th.11 tl11.· brraptolite also be sporadically prcsenr in patches.
pcndem1 W,l~ tllll\llllltl'd of ~.:hitin or ,\ related Fuse liar tissue is mainly fL1sclJar fabric and may be
hydrol.uhon. ' I hl· Ll .1mlun:n t bro\\ n appearance of enttrely so. but in some dcndroids laminae of corn-
tsol.ucd ~rr.lplOlmJ,, thl· Wldcspn:.td u~c of chitin eal f:1b n c 111<1)' be prL·scnt. <111d sheet t:tbrics arc nOJ-
throughom the .uunul kinhdum as,, \tructural com- mally pre~cnt.
poncm. and tht• ll'pottnl prc\t"JKt: of ~uco~am.inc Graptolotd~ 111J) h.wt', tn .tddtuon to all the other
whtdl ts .1 kuo\\ 11 t hi tin bu.>.tkdmvn product all fabncs, ,1 peculur virgular fabric wluch has bet"n
~ecmc..·d to 'ug~t·st dllun. Howen•r, more rc~.:eur found. for mstanct·. 111 the monograptld nc:nu. and
work h,l, t:11lc-d tn dctc(t ~lutmarmne, Jnd it i~ now al\o 111 n:tiolind graptolmd~ 111 \\ luch the peridc:rntll
a!-'Tccd rh.u. "h.1tC\'Ct the mi~in.tl matenal of rhc reduced to J mc,hworJ... ot gudcrs. This fabnc lw
b'T.lptohtc pt•rtdl·nn m.1y h.tvt' been, it was not long fibnls o;ct par<lllel wtth one another in an clcc- Figure 1
chiun. rron-deme numx, and tht' tibnb thcmsdvc~ have~ reproduc
Structure 325
(I) (d) (e)
(c)
Fogure 10.6 Morphol<;>gy of Glyptogroptus ovstroclentotus omericonvs, on early (Uonvirn) diplograptid (a) s1mpli~ed diagram
tiiOwing relationships of thecae and sicvla at the proximal end; (b), (c) frontal and reverse views of a wax model ( · 30 approx ) built
~from serial sechons, some of which ore illustrated in (d) ( 25 approx.); (e) odult rhobdosome (I' 6). (Redrawn from Bulmon, 1963 )
.r
gure 10.7 Ultrastructure of Dictyonemo with fusellar tissue (F). cortical tissue (C) and sheet Fabric (S) (x 14 000). (Photograph
3 101oduced by courtesy of K.M. Towe.)
326 Groptolites
r.tdtal Internal ~truaun.· whll h h." not ht:\.'tl found by \t.>l.'retory cdJs "'irllin tht• md .•llld tht'L' mJ
~h~whcrc. han· mtgr.lted over the tip to l.t} dm\ n torn~.
Ulrra~tnKrural studv of numt·rom g~aptolircs h.1s nuten.tl on rhc out,ide. fhi' uHJld, m f.tct. h.avt
)hown rh.u th~ tortlcaJ b.utd.lgt'' w,;,lthc the whole extended m to a cover of extr.trltt'ctl th\\te 1>\ crthe
outt•r .md mrH::r surf.Kc of the rh.1hdmom.: a<> with .1 whole of rhc sJcula .md l'\ t'n oVt'l th\.· rt'~l ol th
mummy, thereby thirkt·niHg and 'tn·ngrhemng the rh,tbdmome.
tubt• w.tlls. In most gr.1ptoloid' .1nd in dt•ndroid~ thl' The ~tudy of the nJture ,111d di't nbutmn of rhn ·
b.utd:t~t· p.tttern is renurk,,bly tllll\lalll dc:spite an ~tructllr.ll fabric~. wh1ch is all!Vtl) b e111~ 111\C\tl·
oti.en 'lllllmt~ course. ,tnd thl'rc '' ,, dost• rcbtion- gated. ~ll!-':gt·~ts .• "\lde v:met) or Jt,tnhutllln,ll and
'htp between the \ndth of the nll ttc1l bandages .Uld compo\ltton.tl p,tttems amon~'t thl' b'T,lptvhtcs
tht> \17<' of the thecae. Moreover, tht: bandage' arc e\pcn.1lly m the Graptolotde.l. 'I ht, '' 11l probJbh
ofi:~n <;een to radiate from the thecal .tpcrturc, .md h t\'l' comHkr.tbk value in ta.xotmm~ (b)
.111 th\. \e t:1crs 'uggest that dw zomd' mh.tbiting th~:
thecJt were dtrecth resp01mblc t(>r 'ccreting
rht· b.md.tge". Some .Hithonlle~ (Crowrher and
Rtckard-., 1977; Crowther. 19X I) believe that the
only way m which sm h b.wdagc\ tould have becn Clas' Gr.tptolithin.t (colloquially ~.lptohtt-s) I' dl\1JlJ
St'Crett•d 1~ t11Jt the zooiJ, crept Ull( or the thecae, into the rwo import.lnt order'i· the I )emlrOJdl'.t and
e.1ch tt•thaed by a long and rlexibk 'talk .•111d slowly Gr.lptolmdt'<l (Fig. I D.ll ). llowever, that .lft' .u~
mov~: d over the surf.1ee. pl.1sterw~ on Ill!\\ cortic.~l \Oillt' short-hved order'i ( l ub,lldl.'.l, SwloHu1Jca
rnatt·n.ll 111 .1 contmuous nbbon as they d1ti \O, as 1f C.un trOttk.t .md Cmsrmdca; hg. 1!1.11) brgd} con-
by ,\ p.unrbnhh (F1g. I ll.5) fht· or.1l d1c;c '' ~ prob- tined w tll\.' T n·madocia.n \\ hidt \nil lw rde!Tt"'d r
ably the ~ctrt'tol)' me. BJmbg~:~ umde the thecae only in p,h\1!1~.
were ,ecn:ted 111 the <..1me w.ty by rhc zomd within
the th ~c al tube. ORDER I DENDROIDEA (M . C.unb.-t'Jrh.l M onv-hrJt h
Novel though th1s modd st•ems, 1t 1c; at least par- ~rJptoht<·, woth l,or~< numbrr~ nf sm.11l tlll'r.ot•, 'nmrnm~
nally mpported by a study of prembranchs. the vmh ''"llt<·c;uug; link> (dissepim.eu ts) b~r" ~' tt 'tiP<'' \nl'tl
wtth two kmd~ ,,f thL't'.H.' (Julllthc<,tc .111d bnhn.te) npcnmg
ne;~rt'\l living relative to t-,'"~·lPtohtcs In the ptero-
nil .c o nntinuou' dosed clum: rht: ~rult~th<'l".l. I tl\\l'r p.trt lf
branch genus Rlwbdc>pleum tht• tuhc' have a c;uperti- rh,tbd(t\OIIIl h.o~ concentric >hct:L\ lll pt:oo<knu (<urn, tl a
cia.l re\Cmbl.tncc to graptohtc fml'I!Jr nsmc, though sue) (Ovcrm~ lht: ,t.md,Jrd ti.rselldt ll"lll.'. Mu'! ~en, u \\~
ultrastructur.tlh. and b1ochcmtt .1lly the rwo are 'hnibbv upnght .md fixed h> th,• ,e.t tluut, lcl Cyst
rather d1tTcrcnt. Tht•rt• i~ nn extrttlll'CJI nsc;ue in the /)mJr,>l/f..IJIICI< Dc:ndro)od~ .m- ' '-'"'fied onto three tjnub
pterobranchs and, in Cc pl!clle~dt.'ws ar lca_~t. the l.tr~dv bJ<t•d upon thcor ntenul .opp<.U"llll <~. fir tl
dctaJicd 'mKmre os known onl~ m il fe\\ gener.t
zoo1d' can \\ ander .tbout O\Tr the surt"Jce of
FAMILY I DENDROGRAPTIDAE tM C.unh -( arh r Figure
the t:oltmy, mm.; ng on the1r cephahc sh1elds. VorJr,>l!r.lptm. Cys~1
Furthermore. although no livmg pterobranch hJs FAMILY 2 PTILOGRAPTIDAE (l ( l11I-U. '>ol. to the c
attually been observed in the act of secreting its flt If, ',l!'•lJJfr1.\ . Sil.). SI<
sh·lcton, the sc~:ond.uy tl,sm· <lt lc.1st could only FAMILY 3. ACANTHOGRAPTIDAE (U Cmoh. M. llrv
From Bt
have been produced by the 'louids during thec;e l" ~ ' lmllllltWIIPtm.
walldt·rings. ln graptoloit.b it is pmb.1ble that both ORDER 2 GRAPTOLOIDEA (L. Ord.- 1. lk\".): lllubJoMIOO Cr.' I
luv~ li.·\\ \tcp<·~. up tu ctght m tht l'.lrl~ tnnm, bur rc.!u Mnchc
fusdlar and tortlcJI ti\\Ue were produced by the:
111 l.u,·r !;<'lll'r.l tu rwu .md finJIIv to one. I ht'< ll 2rc of hut tlll
zoo1d\' t:ephahc di\c.
kunl on I). n1uivalt:nt eo the JUtllth<'LJI' nt ,kndrotJ ~ J'cnden
In the opm10n of 'imllc, tht· fom1 and arrange- lltJ\" bt• lrtang,·d on ouc ,idc.< <•f th• •Up<' ur on both. SUB<
ment ot contcal bat1ddge' nukl'' ir unneceo;sd.l) to UJI.ltnliiY ~~mrnl"tnc.tl lOkHnl"' m: pnnnu n; th, lo oflh I .1111
postul.ue the exhtence of t'Xtr.uhcc.ll ns\tH.' m grap- umdl!lnn m the Monognpntilt' 1' W< und.u"\ . A ~<ub 111.111
tollte\, but the comtructllm ut nem.tr.l, nemal vem~ mg .1 ncm.1 j, .ilw.ty' pr<''t·nt 111 tht• ,oduh ,md I!> u u rulm
,\lld homologous struLturt'\ 111 c.kndroid, ,., not e~ISlly pro11lii1L'l\l I hl' terminology of dlflnt'nl dl<'< 1l duJ~ l.ttt<.;C
g!V('Il 111 hg. III.Kd. t1d rJ
cxplai ned in tlu~ w.ty.
Wtthm the gr.1ptoloich tlwrc ts nt11d1 v.uuuon tc1 ll1c I R,ufi
The nem.1 h.1~ been shown to be a hollow rod. rnnrpholnh'Y· by conrr;~~t with th.at nl dl'ndcucd,. (,rJptolct, somt
open .H 1b di~t.tl end. lt mu~t lt.IVL" hL'L'Il constructed Jr• dJ"1ticd 111 three ~ubordec>. Wllh
Classification 327
(d )
nem<J
rec oned
Kieerogrsptus
(b)
pendent
(e)
Cyrtogreptus
D•crsnogrsptus
Cystogl'llplus
lr
hgwe 10.8 Various dendro1d and graptoloid rhabdosomes: (a) Kiaerograplus (Tremadoc) with rare bithecoe (x 6 approx.); (b)
Cystogroplus vesiculosus (lland.) with a tri~d nemal vane (x 1.5); (c) Dicranograplus ramosus (Corodoc; x 1); (d) terminology applied
1:1 !he different shapes of rhobdosomes in graptoloids; (e) Cyrtogroplus, with webs connecting the stipes (x 1); (f) Cyrtogroplus (M.
·~· ~1), stages in early clodial development (x 15). [(a) Redrawn from Spjeldnaes, 1963; (e) based on Underwood, 1995; (f) redrawn
!tom Bulman in Treatise on Invertebrate Paleontofogy, Port V.]
.).
l'S Graptolmd~. follmvm~ f orwy .111d Cooper ( llJHn) and as in Bry<~l/mpllts, or q u adrirudia tc .Is 111 Stmm•,l/mp111S.
\btcheU ( 19H7). may be ciJ~\IIit•d 111 three ddined ~uborders, The srrucrun· of the tlll'Cit' 1~ l..now11 only 111 a fe\\ amso-
ur the taxonomrc pO<IUOil of <0111l' woup\ (anisograptids, gr.lprids, smcc therr prcserv.mun 1\ nornuUy poor.
~,dent d1chogrJ~md~ 1nd rt·trolwth rt•mam .. unn:rt.1m). Kiaerograp11L1 (Fig. Io.R,l) .HH.I \\lllH: l'Jrly 'PC.:Ut'\ of
SUBORDER 1. (Urumj<ncd) cum.um d1e p:!raphyletrc Bryograpws have both .rutotlwclc Jnu lmhl'< .1~, but other
Fmtilv A111\0!-'r.IPtldJt' ( rrcm.) wluch ha'> charJCtCrtJc
~mny WJ}' mwmrcdr.tlt' bcm:t·cn dt•ndrmd> .1nd !<rOIP- SUBORDER 2. OICHOGRAPTINA (Ord) Early graptol01ds
toloJd., but thc pr~·wun: uf J 1lt:111J !-'l'OUp< them Wldl me lackmg btthecae, Jnd wrthout .1 vrrgdl.t SUPERFAMILY
bttc:r Mo<t JOJ<O~'T.lpnd' rt:IJIII b1tht-cae Th~ ani5ograp- DICHOGR.APTACEA {FAM ILil'l I)JCHt)GRAP-
tld rlubdo,ome<. ma~ ftmn IJrgt· 'preadmg colomt."S .J> m TIDAI: {e.g. 7'rtr<ll/'oli'IIIS, Dtr/lt'l!f<IJllll•. 1..::yl!<~raptus).
R.ult,'j/rotplus. Rlto~l•.lrll<'l'''''' ami ollll'r.. Sm.Uler thabdo- SINOGRAPTIDAE (c.g Sillc'l!r<lJ>Iu~). SI(;MAGRAP-
somc<. \\.,th fcwt·r br.1nt hm~ point< mJV bt." b ilateral (1.c. TIDAE (t•.g. Siii"'·'~'·'JlliH, (.rt•m~raptus) coml\t' m.unlr of
"nh two pnman br.ull ht")) .ts 111 Cl.•tt•'R'•IiJIIIS triradia tc early graptolotth WJth onl\. l tcw dKhotomou\ branchmg
328 Graptolites
pur m, Thev nuy lwn· u~ht. ti.>ur 11r m o 'npc.-s Jnd h;~vc. m.w bt• c\t...n\Jfoml. reclmcd (Uilhl'nall ur p.•rn.lll} son-
J 'unpk proXJm.U end ,tructun·. In ~LI'ERFAMil Y dt nt (lllll-hherial) (DIChllll~~rciJlliiS D~ot//<';l.'tfJtllc ,
GI 0SSl1GRAPTACEA the !-'t.rptolouh always shm' \ 'n"J.I?t•IPIII<, 1-"]tt.>gr"l'tw) FAMil V PSEUDO-
'"o~nptrd 'pnmetry'. "hue tlrl' ~I<UI.r .md th. I tom1 .1 (. I 1.\t\ACOGRAPTIDAE (0rd) «mlpn·n "3ndent bt>- Fo
'YI11111Cin~JI p.11f so tltJt the ,pa, ul 'YIIHIIetry of the rhab- t•ri.ll fom1' with .1 ~rraighr nr l'tW,lt-: lllt'lh.ur ~t·pnun Jlld ~ the
diN>mc p.t"c' bl't\\n·n drcm. fFAMLLIES d1.1r.Klt'n'u' cnmplc.-x pro\llll.ll nlll Stl."lll !,'Tuup Ill<
1\0( atAPTtnAE ~·ilatl•r.tlly W111lllt'fl1l.d. l'.~. lso.~raptlls, PII YLLOGllAI 11'11 >Af. (Ord.) n11np• i,,.,
qu .• dmmal tin
Ollrc~l!'''llllt5, Sku'.l!'•'f'lw), PSEUDOTRI- \Lllldl'nr Aflptoloid\ with 'i111ple thc:-c,I C, .urd 'i\11Kf,1ptiJ'
GONOGRAPTIDAE (s,,mdeall tml·ri 1l m <Jll.ldrueri.tl it
Jewlopmt:nt of the prox1m.ll rnd (Lg l'lly//c~~I·1Jirll>).
/ 1.>t'llcitllni?Oih~Jiraprus): <.LOSS< )(:JlAPTIOAE (,ramlem 'I ht tMwnonuc pomion of the pl'ndcm tlll rogrJpnds Ko
bascnll tom1' m wh~rh the: thn.lt 111.1} lUnc.: m ord.-r ro (c.:.!-(. V•cl)llll<'l!'•lptus) and rhc rctJolrtrJ, (y.v.; <'.g. NrtlcJ/rr,<, ~tn
t·ndmc the sacul.1 m ttont .md belund, "' 1h.u the ~tipt'!> Plr''"'''')!'·'Jllll', <_;,,th<~l/ruptu.<) m \dndl dw pt•ndrml 11 HH
.m~ :ah!-'llCd "de.- b) ''de hut face 111 oppu,atc drre.-trorb, rtdUl·l."d tu .1 rne~hwork rl'maJil\ w ht• J"l"\cd. Wi<
uftc:n 'pmy: l'.g. Glc>.<S•'1:'.11'111!, ( ·'''Y"•'Iolc s. f'wctciiJOgmptu.s) ORDER 3 TUBOIDEA (l. Ord.-:nl.: F1~. lt1.'l.1): Dcn.irclld- Pll'
SUBORDER 3 VIRGElliNA <)rd,-1 >e\' ): A vrrgdiJ a- JIW<I~ lrkc. "uh Jlltn- .md bithcGJl', but \\Jth trrq.,'"liiJr hr•nclung fro1
pn:' nt m tht• \KWl and rn thl· 'm~!k SUI'F.RFAMTl Y 'true tu re: .111d 1 a·du.-ed ,tolmhn~; l' ·!{· /{,n.rllc~1Jplltf
DIPLOGRAPTI\CE.A thtc nur~n of thL \Jrula bears ORDER 4. CAMAROIDEA (Ord: Fig. to 'lb) l:.mrmnng fonm,
Lo'
.lpt·rtur.tl 'pme,. FAMILY MON<. ><.;RAJ' III>AE (Stl.- wuh .lutndwl.IC of peculiar ~hapc:, uch h.wmg dll mHJie.l
..lll
1kv ), wlm h in dude~ the Cyrtuw Jpud,, " utmcnal and balloon-like h.a~c. with .1 vcr1.it.ll collum (dnmncv1 Ko·
"~ml<•nr (e.g. .\fvrw~r.rp/tl), Mtollt>dllll•lti.<, C)'rt<~f(raptu.<). ~rol otht:cJ Jntl bithc:ot' prest"nt, e.g. RitiH'II'ffllllnm. Gr~
fAMILY GLYPTOGRAPTIDA.E (Urd) contpnws ORDER 5 CRUSTOIDEA (L. Ord.-U Ord, F1g. IO.'k) or~
\l',lntknc hi~erial grJptulorJ, ''hit h h.aw .1 duuctcri~tic Encnl\tr ng fonm. w1th dendroid-lrJ..c morplmloKY. but the 1
'glvprogr.1pnd' proxuml l.'lld p.rtt~ rn (c.~-:. Glypt<',llraptus, JlltOthet,ll' h,I\'C moditic.-d apertlll<"', C g. nu/m,uuml<l.l
11101
\!lint Clm~<lu>graprw). FAMII V 1)11'1 OGRAPTIDA!:: ORDER 6 STOLONOIDEA (Ord -M \d. Frg. 10.9,{'
\llll
(Ord <;,11 ) compn'c:- '~andent b1wn.1l tom1>. U>Ual.l} Encna 1111~ tonm of 1rrc.-gu!J.r morphnlu~ ,,,,[orht·c:.~ lu\·e
\nth J rned1Jn scpnun Jnd ".hplogt.•pud' proxmul end 111.111)' 1rrc~hr r.1mlf)'lo~ stolom: Jutothn· lc pn·,ent but no whi
p.lttl'nl (l' g. D•pt•>g•.tptm. ( "hth,'.l:l·lf'IIH. C:luu.,,~~fJI''"-')· In biUll'tJl', c g. Sr,•IJlucJ.Iwdnmr. onJ:
FAMil V DICRANOGRAPTIPAI (Otd ,) rhabdo\nm""' the
Ccp
\0111
(a) Retlt:ulograptus, a tUbOIC1 (b) Blthet:ocamara, a camaro1d R
tS Cl
bltheca
bou
Se
(c) Bulmanlcrusta, a crusto•d
the
bud
stolothec.~
1111--7 stolons
bltheca
autotheca
(d) Stolonodendrum, .1 stoiCliiOMl
Figure I0. 9 Minor groptolite orders. In Slolonodendrum the irregular tubes ore stolons; the enclosing stolothecoe ore rarely pre· Figvre
served [(o), (b) x 50; (c) x 30; (d) >( 15]. (Redrawn from Bulmon in Treatise on Invertebrate Poleontology, Port V.) (Mod if
Evolution 329
each tube I~ sm:t!l ;lJld ha, d palr or tl'Jllacular food-
gathering anm known as the lophophore, though
For a long tinH: it w.t\ .tcct•ptt·d th.lt the aflimne' of th1s is nor nccessanly eqUJvalcnt to the lophophore
h!! GraptolnhinJ lay \\ Hh Phvlum Cmdaria and of other 1nverrcbr.ne~. Each md1v1dual zoo1d 1s
~o,t probahl~ '' tth the hydrozo.m,. The publica- supported by a I."Ontracnle \Wll- wluch hnks it
non of Kos.dm.. ,ki\ work 111 1'1-H~. however, made With a p ecto c a ulus nmmng through tht. colony
·: dear that thl\ \ ic\\' \\a' nu longer acceptable. as .1 stolon system doe\ tn a dcndr01d. lniti<tll)' the
Kmzlowskt nott·d pronounct•d n:\embl.lnces in pectocauJu, i\ \oft, but tt later becomt'\ rigid. The
1tructurc bctwct'll tlw gnpwlnes and the small, pectocaulm with 1ts 'lmil.mty to the dendrOid
modem, Rltt~brlvp/n,,z: .1 Ullonul organism with a stolo,n gtve~ some mdJC.mon of afiinity. but perhap'
,,·ide depth range (usu.dly 101}-JOO m). belongmg to the relationship of the rwo groups " mmt clearly
Vhylum H emtchord,lt.l and \\ ith rdo~tivcs known suggested by the pre~cnce of coU.1gcnou.' fi.l\dlar
!Tom scant} matcn.tl in tlw Cn:taceous and the tissue which fonn<; the peridenH of both brrapto-
Lower P.1laeo:wit. The hemtt hord.1tes are an lites and Rllc1bdople11ra. ~everal genera of rhabdo-
'ldv.mced' phylum ,llht•d to tlw vertebrates; if pleurans have been ti.nmd in the Lower Palaeozoic
Kos1lowski\ 'u~g~.·~tion' .trt torrn:t, then the (Urbanek and lkngtson, 19HC>), e.g. Rhabdot11lws
Gr~ptohthma must be of .1 much higher grade of (M. Cam.), Graptmwmis (Trcmadoc) and some
nrp;aniz;1t1on th.m wa' on~in.tlly tmat,'lned. ,JUthorities ~uggc~r that Stolonoidca are actuaUy
The l lemichordJt.t rompti\e two das~es: the rhabdoplcuram. The earlte~t recorded Rht~bdopll'ura
modem Emcropncmt.t, wh1ch .m: the 'acom worms'. is Middle Cambri:m 111 .1ge (I )unnan and Senn.ikov,
1uch .1s B,lftllll~f!lanw, .tnd the Prerobranchia, to 1993).
which Rfl£1/)(h,piC'Hr,z bdon~'- ln both ca~cs there are The study of livmg ptcrobrand1~ ~erve) ,lS a
onl) a few R.cccnt genera .•md 111 the ptcrobranchs model for many a\pect' of gr::~ptohte bmlogy and
d11~ only 1mport.mt ont:~ .trt' Rltabdopleum .1nd skeletal growth. Sibrnific:mtly. in Rl~c~bdoplc11m
CcpltJiodzsm.'. whil h i' \IIHil.lr to Rlwluf,,pfcura m (Rigby, 199-t) the 7omd' often dte off but .tn:
1omc WJ}"<; hut doe~ not form true colontcs. replaced by other\, budded otr from the stolon,
Rlzabd,,pfrma (fig. I 11 I 0) '' of very ,m.ul ~ize .1nd whJCh lllhabtt the 'ame rube 'lnd conttmtc to .tdd to
1' colonial with indtvidu.ll c:<thkdetal tubes. e.Kh 1t. Such mulnplt.: ot-cupancy 1s marked b" dear
bOU'1ing .1 zooid .md .lrr.mgt'd 111 a t'rt:qnng habit. growth d tscon tinultle\, .md the same arc found in
Sever.1l ntbc' arist· fiom .1 lwriznntJ! ba,aJ tube, graptoloids. 13y analob'Y, thl•refore. gr aptoutes might
the la~t om having .1 dtl\ed t•nd with a terminal well have survtvcd .tdvcf\e condltlons tf the old
bud inside. Tht· zooid whzth ~ecrt'tt'~ and lives m zooids died off, bur with 1mproved condiuom new
buds arose; moreover, old or d.1maged zootds could
have been wnrinually replaced throughout the lite
of the colony. To build any kind of robust and
long-enduri ng restdence i'> cxpcmive and, a~ with
the hnman sphere, it makes stmc to build a house
for ~eria l occup:mcy.
Shape of graptolite rhabdosomes

(fig. 1 0 . 1 1)
The following summary of cvolunon in gr.tptolttes

1~ concemcd w!lh KfO\\ morphology: the kmd of
fi(Ju"! _I 0. I0 The modem hemichordote Rhobclopleuro (x 18). dctail!i that can be \ecn Ul Rattenl·d 'pccimens. Later
1Mod1~ed from Kozlowski 1948 ) 'ome particular det.uls will be added that can only
330 Graptolites
~ he
CAMBRIAN ORQ()\, Jrlb.N SILURIAN DEVONI~N dit
~onodlon
I Middle Uppor rromo<IOC . Atonia Llv-lld .lond. IWen Ludl, IPrld lowor Middle U~per
e Ca
~~!·I
0 Cl
n
-n
)>
-n:I:
)>0 l~ I ! ~ 3
1 the
c
z
)>
CGl
Z;D
)>l>
-o l t t
5' 0 :1 ·~ i ,,fj,
-i 0 Du
~ 0 IOIPI I FAUNA _fA ~
lll.l
I StolonotdOI (()
Cruotold.. l'V('
Coli
rep
r
run
.1bu
Ul1tl
f<:rc
:>- PtnO.nt dichogreptide
wcr
;l l~o
peal
Ord
.H.•ln
bran
SIQmtgrtpttd"
stole
thci1
"' twist
lt) c
snul
\tc:n
l vc:r,
I! .IS<:n(
.Ill ISO
shapl
tr.1l h
\US pc
IIJIII. !)maL
to the
life: S•
nonn
Th
.mac~
bryoz
~------~~
~-----~-~-~-------~~r~------~------~ llotloltttdoe
W:IY ~
mg it
water
11)75)
(I 996:
ot dan
Figure 10. 11 Evolution ond founol sequence in groptolites: o generalized picture. Genero illuslroted: 1, Dithecodendrum (M. rt>pair
Com.); 2, Ptilogroptus (Ord -Sil.); 3, Rhobc/inoporo (Tren1.); 4, Dendrogroptus (U. Com.-l. Com.); 5, Didymogroplus (Aremg); 6 rl'turn
Tetragroptus (Arenig).7, Smogroptus (Arentg); 8, Porogfossogroplus (M Ord.); 9, Corynoides (Corodoc); 10, lsogroptus (Areoig)
11, Monogroplus (Stl.), 12, G/yplogroptus (Uondeilo); 13, Orthogroplus (Corodoc); 14, Dicronogroptus (Corodoc); 15,
Orthoretiofites (M. Ord.); 16, Phyllogroplus (Arenig).
Evolution 331
be esrab!J~hed rhrnugh the examinauon o( d1rec- blc. The earlic~t planlo.m Jc~n~nd.tnts ot Dht}'!111t:Wtl
Jinu.·m1onal maten.1l. Roared apex upward~. and dl\pen,ed wtth the
The earucsr graptulitt.:' J...no" 11 are M1dille attachment; in it~ own \\ av lJUile a maJor evolution-
Ca.mbnan and t"Oilll' from tht ~1bcnan Platform; ary seep. The transtt1on trom bentho~ to pl.111kton b}
the) are ~mall m •g· hkc ~l·ncra dl'lovered .1nd das- gr-aptolites i~ p.tralldc:d b} VIrtually JIJ other plankuc
ufied by Obur ( I '17 -l} in (WO llC\\ ordcf"\, the organisms: no les\ than IH out of 2 L plank tic group~
D1rhecoidea and Archat·odl'ndnd.t, though more had tltetr orig111~ 111 the benrho\ l R..t~by .md Mihom,
marenal m·cd~ to bt· t~mhconung if thJS ra..xonomy is l9'J6}. Such inva~10ns. whteh took place succes-
to be smramed. In tht· Tremadon.m and larcr rhe ~vely through time ~ecm to ha\C been random. and
C\'Oluuon of waprolm·\ 1\ bcttl.'r kJIOWll. The Upper independent of .my global events. fhe ti.f\t planktic
Lambnan graprolin~' .tn: .111 dendw1d\. ;tlong with graprolitcs probably arO\C m deeper, cooler \\atel)
reprc\cntJtl\'es of the ,hort-li \'t•d ordcf"\. and thence rad1.1ted into open peb~ic and neritic
Drndro1d~ wert· qlllt( d1verse 111 Upper Cambrian enVIronments 111 the e.1rl> Tremadoc (Erdttnam1.
rimes, ;md rhougb rht') were thereafter ne\·er a very 1982) .
.1bundam componc:m of .111y taun,1, tht·y continued The earliest plankttc graptolttes have traditionJlly
uuri.l the C.1rhonifernm. Vilnom gt·nera having dif- been reg:Jrded as ari~•ng ti·om the Aoating
ferent time ranges. Dendritic or tw1g-likc fonns Rlrabdirwpora.flabell!f;m,c, presumably because of the
were pcrlups the most common. though there were widespread occurrence of this fos~il just below the
also some pinnate gcner,l (e.g. fllllq~r.zptus). The anisograprid t::luna. R. . .f1abcll!f(mllc, however, IS of
peak of dendroid lOillpkxit} 1~ re,tched in the conical, bell-bke fonn. rJther tl1.1n bemg A..mened.
OrdoV1e1Jn m Dt'VOI11.l11 genu\ Kt~rl'llllli!Yilprus. an A better c:tnd1d::tt«: for the R.lt.~bdinopt>m .mcestor
Jramhograptid \\'11 h .t l nmpln. and .1nastomosing would ~eem to be tht· t1at, r,ldJ::tlly organized
hrJnch ~y,tem. F.lt'h branch has m.mv \Ubparallcl Rllditwmprru, the e:~rhe~t ~JCttbtc graptolitc. whteh
ltolorhec.lt all nnul'd together in columns, with ha~ three or four pnnury ~urc~. Jnd 1n wh1ch, as
thCJr attend.mt .mtorh~·cac .md birhec.1c irregulJ.rly in the :Jl11~ognptJd~. the dJ~~epuncJlt<, arc but
mNcd up togcrlwr tn 1 thtck \tem The VJ'>t major- weJkly developed. When.· th1s gc.-nus occurs. in
:v of dcndro1d, b'TC\\ upnght on the sea floor like Ne\.vfoundl.lnd, 1t undcrlu.'' thl' ZO!ll' of R ff.J[,c/lr-
1null \hmb,, tixcd by holdt.l't' \dm:h. like their }~mnr. I t i'> ~uggc,tcd (C oopt·r .md rorrq. I WQ),
\!em,', were strt'llb'thcncd b) lUrtlc.tl msue. How- that Radul}!r.lpCII( wa~ eh-:- :tlll'C~tor both of the (\'cry
n·cr, um· or t\\ o 'Pl'tlt'' of Rh.llulruvpord (tonnerly s1milJ.r) four-mped am~ograpnds (e.g. Sr,wr,~~mplm)
lilribed tu Dr<~y•>llt·ur.l), .tmong-;r the e.lrlit·~t of the and the thrcc-'>rtped lnt \tli!YtlJIIHS .1~ well as
.nMt,rraptlth (.1 more rig1dly t•ngintTr~·d .md cone- Rlwbd11wpom 11\clf, frtJlll wh1rh m runt Jrosc
11-..tpt·d gcnm) ..1pp.1rwtly tkp.mt·d trom rlm .mee~ Bry,wmpws.
lr.l! lubit .tnd hvcd in .m inwnnl p<Ntton, probably Young ~cage<; of rl1.1bdmopond' 111 whJCh di~~cp•
U\pended by rhl' chu1 m·nu from Ao.tt111g ~cawccd. mcnt~ ,1rc Wl',tk ur .1b~cllt, huwcvcr, arc often found
~mall attachment discs h:tvt• been found :ttt:tched 'di<;coidally' prc~ervcd. i.c. A.tttencd radially, ,\t ,1ll
to the end of rht· lll'lll.t; ht·ncc :1 ~u~pcnded mode of strarigt<~phic levels .•md the~e ;m· sometime~ domi-
bll seems likdy, though it w.ts ccrtJinly nor the nant on p:~rticular lx:dding pl:lnc~. Erdtm.tnn there-
om1 for mm1 gr:~ptolitt·~. fore believe~ that the :mrc~tor~ of thL· planktK
The sessile cruc Orcryollt'll"' spt·cic~ probJbly lived graptolitc~ may bt• sou~ht in p.lcdomorphte
ruchcd w the se.t lloor, .tpt'X down like a conical Rlwbdinopom of.flalll'llijlmuc typl' rJther th,ln neccs-
!:-.ozoan (Chaptn <>), fc cd111g m the ~ame general ~arily in RadiograptrH.
~y by cxtraning tC.md p.trttdes trom current~ Aow- Most .misograptid' .trt' Tr<.'m.ldoetJn m .1gc: there
g in through the mt·~h. ,md t•xpelling exhalant are record' of .1 po~~~ hie l Xtl'll'!On mt0 the later
-:~tcr rhrough rht• open t•ud llf the cone (Rickards. Ordovician, but tht'> ts not ccrt.l111. The) tna) have
i5) In a Stotci'h S,Jun.m t:-tutu de~l nbed by Bull two. three or four pnm.U) hrJnche' as 111
996) ,e,•c:ral DH ry.lrrt'lllll 'peumem sho'" L'\ idcncc Clo11ogmprm, Br)'t>\!mprus Jnd Swtr,~l!f•IJl!IIJ. rc,pec-
cbm~gl' b) prcd.Hitm or di't'.l'-1.'. but \\ere .1blc to rivcly: they arc rarely prc\ervcd other than J'\ com-
(M f;Ur lnJII), though llllt all. lo.md\ of lllJUf) and to prc,siom. However, 'onw c:;c.llldinavi.m genera .trt·
), 6, m to a nonn.tl l,.'TO\\ th p.tttl'Ol ·" \flon J'\ po,si- pyrirized, and the\~: .lrc: of particular interest for
nigl,
15
332 Graptolites
the> show clurJttl'l"\ mtt nncdtatt' hetwcC'n those of from an umpecified rwo-stipcd d1chowapnd
Jcndrotd~ and gr.tptobith at the mJcrO~lOplc level. antestor there aro~t· ,lt \Oilll' tlme during tht• Lower
The Upper Trcrn,tdocian gcner.t Kiarro,qrt1ptus Ordovician the varium \tock~ that cJmt• to donu-
(Fig. I 0.8a) .mJ H'l'<~I[I<IJJIIIS apparently h<tve bid1c- n.tte the Upper Ordovlt ian (Fig. 10. 1 h· t') Tht•se
tae, but the~~ ,trt !ewer in numbcr th.m thme of nlvcr have more th,m t\\O snpcs. fll''\t th~·rl' .1rc the
~tand.1rd dendnlith, .md there ts only nne bttheca tor dtplograpnds. whkh arc ~cmdcnr tonm. The\e han.·
every two .tutothcc.tc: furthcnnort•, they arc the1r on gm well back m Ll.lnVtrnian nmc. thl·•r fiN
reduct:d m ~ize. Why the bitht·cac \\ert dtspensc::d representaove bemg G/ypltlqmpttts dt'lltatm. Thence
wnh during the latt·r evolution c)i b"l'aprolires is not they can be traced to the end of the ll.•ndnvel) . g
cle.tr. The several diplogr,lptid genera arc distlngtmhed
Koszlowskt bl'IH:veJ that the bnhccac and one from anotht:r mainly on tht• ch.lrJrtl'l"\ of their
autothccac homed mak and fcmJk Lomds, respec- thecae. Thus D~~lt~qr<~ptu., ha., ~imple. \tr.ug;ht thcrae.
tively. and that .._.,.,tptoloicb were ht·nnaphrodite. ClrmaCt~(/r<~ptus has thcc.lc: of a 'quJ.n:-cut .lppl'.trJnce
Altemanvcly. autotheot may h.wt· been \ttes of (Ftg. I 0.4), Clyptogmptus has gently l l l n mg ther.w.
feedm~ zoOJd~ only, \dule bithec:te were ~olely etc. All these graptol01d~ arc more or Je,, dlipric:~lm
reproductive. Wh.ttever the functiOn of the bttheca cros~-'lecnon and arc t•asy enough to idnHit)· when
was. it mu~t havr been t.tkcn over by autorhecac; flattened, provided th,lt they .ue pre,ervl'U wuh dtc
thts might h.tvc luppencd through an imennediate 'lon p; aJns' of the dhpsc paralld with the hc:ddmg
kind of graptolitt \Udt as till' Llandeilian plane. for then the thecae are dearly 'c:c.:n 111 l.ttcnl
Cc1lyxdcmintm, 111 wlm h the bnht•c.tt'. rather than VtC'W u: however, the.: long a.'\.iS of the dlipric.•l ern\'
havmg external ~perturc,, open din•c:dy mto the ~ccnon i~ perpendicular to the bedding. tht•n nnl}
aucothccae. The a1mo~rapnds b cc.une extinct by rht: rhc .1pcrtt1res show.•md smcc they alllooJ.. ~111ul.tr 111
end of Llandeili:lll time and the1r progcn1. the 'real thi~ sca)a riform v ie w they arc not ea~) to tdentil) .
gr.1proloids'. b~:cantc dominant. Continuom sec- Other Upper Ordovtc1an genera indude the V-
lions 111 the Tn:madol arc lew Jnd t:1r bl·rwcen: the \hapcd, stoutly built Oic't'lltJgraptus, the morl· ~lender
best i~ m the Yuko11 TcrntOt) .mJ hc1s rc.:n:ntly been bur otherwise sinu!Jr L£'J!Itl.~mptlls, .l!ld Y-sh.tp<'d
descnbed to sho" • l ontmuou~ sCI.Jllt'llLe of b'Tapco- Dirww~rcTpWs findudmg D. mmosw (Ftg. HU'ic
lirc faunas. The t:1una here i~ ntJlttly of anhOb"l'apuds \\ h1ch can be mort' than 30 cm long! . In thl· l.l\t•
.md dendroid~. but there arc aho 'omc.: cc1rly 'rear llll'nnoncd genu~ the ti~t-fom1cd part ot the rhab-
graptoloid~ (dH;hugrapnds). the forerunnen of the dmomc i~ scandent; m ~ub~equcnt ~wwth rhc
m.nn stock th.tt dom.in:1tcJ the lowcnnost hranchc-. diverge. The Ol'C\IITcncc ofd1pk\~, diu.·llo-
Ordovinan. and Jicranograptid~ in tht· Upper OrdovKt.lll gavr
Dichob'Tapttd\ luvc no bithccJe. TIH.~Y arc usually ri~l' to .1 belief that th<.· dtplogr.lptid\ evolwd from 2 Figut
\ytnttlctrical. bt.tm h did10tomou~ly anJ may have Jircllogr;~prid anc:estor by 'ztppmg thcm,clvc\ up' !Red,
many or only a ti.· w bralllhes. A buN of ad.~pmc and that the cond1t1on m Dicmno~raptm r<.·prc\em~J Port 1
rad1:lOOn m rhe t·.Jrly Ordovtcian re~ulrcd in a pro- an Incomplete ztppmg process. Attral tive thout:h
fusiOn oftyp~. e•ghr-, four- or two-bramhcd, with ti11S ~uggcsnon Jppcal"\. 1t is unreahst11:. to• thl.' Gnt
rhabdo~omcs pt•ncknt or dechned, or even scan- diplo!,"l':lprids appt'.lr wdl before the c.ulte~t lhu:Un- \h,lp
dent, like PhyllcwrttJIIII.,, m whJCh the lour ~tipt:~ an: anJ dicranograptids, and in any case the der.1ilcd otltc
<lrranged back to b.1ck .md are in cont.1Ct all the way structures of the 'proxtmal end' - the lt'\\ tiN- lugh
up from the bJsal SICUI:t, or Tristidw.~rap/11), which tonned thecae and the sicul.t. ,,·hich Jrt' all taxo- "otic
h:1' three: sc.mdC'nt ~t1pes. Other genera are nomically diagno,nc - ,1rc quite differt:<nt. ( ·)11/c
Tc:cragraptus. \\ tth tour dcclmed or pt·ndcm stipe~. The diccUo-, d1crano- .md leprograplld' became (dad
and the common. 70nJI1y useful Dtdym.,~ttlptus. in e.\tHu.: t bdore the S•lun.m. though rhe d•plogr.~ptuh V.J
whtch the tv.;m ~ripl'' m:ty be pendent hkr a tuning cun tinucd until thl' end of tht.• Llandovery ·" mtpor· expl<~
fork or cxtendrd in !me. Scver.tl of tlw t:arly multi- tant members ofthc total graptolite f.1llll:l. ing t1
ramous (rnany-br.lttrhed) dichograptid~ had a web- Tile Siluri<tn is dominated by mono!,.rrapnds: ~tan the c:
hkc structure between the supes .u the proximal dent funns with the tht·c:-.t c arranged alon~ onh one part, ,
end, which mcn·a,t•d rhc surt'Jcc are.l and probably \ldc of the ~tipr. They first .tppear JU~t .tbovt· rhr. rh
gave assi!otancc 111 Aor:uion ba'e of the Silurian. Monograprid~ come in \:tnom hete I
Evolution 333
,,::;:;~. ~ ~ t "·~
~
/-'"'
~!!¥ ,-t-~-ff
=---------- Monograptus
generalized diplograptld
Jjt .~ ~
dentatus ~
leptograptid
Figure I0. 13 Spirograplus guerichi: three--d1mens1onal sped·

men showing helical coiling, from the Silurion, Osmundsberg,
Sweden. (SEM photograph reproduced by courtesy of Dr Dennis
Bates.)
c\tablishL•d. l lowever, it i~ based mJmlv upon com-

pre~\iom, and rdcn.-nce to three-d1nH.'mronal mate-
rial cxp.mdo, .md illuminate~ tt to .1 n·markablc
f~ 10.12 Evolution in graptolite proximal end structures. degree.
,Redrawn from Bulman in Treatise on Invertebrate Paleontology,
Port V.)
Proximal end in graptoloids
1hapc~ anJ SJZCs; some arc very long ,111d ~traight, The Graptoloidea differ from thl.' Dendrottkt m the
others short .111J ~tumpy, ami LIH.T L' arL' various absence of bithccae, the rcductmn of cortlt ,tl tissue.
l11ghly rlludifrcd fonm including curvt:d or ~pirally the form of the \icula, the presclll"l' of .1 lll.'lll.t, the
coiled genera (Fig. 10.13). \Omc of whtch, like mode of rhabdosome branching .utd the '>tntnurl.' of
Cynl~rtlptus (Ftg. ltUk,t). may luve lateral anns the thl.'c.tc. T hey arc also uni4m· in the \\,1\ Ill
clldta). wluch tht• tim tcw thecae develop from tht• 'icul.l.
Vanom change) m thecalmorpholog), a' w1.U be In gem·r~l tcnm. the early graptoloid\ llJve .1 rathl.'r
cxplamed m detail later, appeared 111 \UC<:esston dur- sunple kmd of proxtmal end development: the latl.'r
~ rhe St..lunan. The last monograptid\ flounshed in Ordovtctan one~ are much more complex. 'uch .1s
me early Devonian but wen: rc~tm.ted to certain that of DipiO,I?rtlJ'IIIS which may now bt· seen in
p~rts of the world only. cvolunonary per~peetivc. Variom typ1.:~ of prox.tmal
rlJc pKtUre Of graptolite ('VOlUtion prCSCntCd end development arc shown 111 Fig. 10.12, tllu~trat
nere lm been known for a long timl.' ,md is well ing the cv<1lution of structure, but it must be
334 G roptolites
remembt·J c:J tl1.1t tiK· m mid~ reprc~entcd here arc thecae. Norm:tlly 111 Dirc//,,~,,,ptu} .md 1ts relative~
grcatlv ~unphfi~·J ti um thl' .Ktu.JI 'trucrun:~ encoun- the \1cub rcchnc~ ag.1mst one of the snpes.
tered 111 re.1l graptulmd' .111d tiJJt tht· pattem of There arc ~omc vcrv modtfied modes of develop-
proxunal~:nJ l'\olunun 1' much more compkx than ment 111 ccrt.lln 'candent graptolmd genera.
thi~ ~implified p1ctun. would 'u~cst. Mo~t Cryptll,(!mpflls, for 111\t,mt e, ha\ a thecal system quite
dicho~rraptid pro)Umalt·ml, .m· of unmodified forn1. different from th.u of the dtplograpuds: m mo~t 11<
In rl1~:'e (.i~ in Dt.l)'mt~I!"'JIIII> bifidw) th. I~ brrows out peculiar feamrc I' the tt)f\Hm of the tht·cae wluch ter
from rh. 11 ano'' thl· -.iud.J and i' \\Tided m it: it rc~uJt, m the 'erit•, rh. 11• 2 1, 1', .. opening on
tonm a '-mglc nm~ing canal. 13ut in Luer dichograp- oppmtte ~ide' (from.ll .llld rl'vcr.c) of the s1cub to !at
od.~ (e.g. Oulylllc'f!"'Jllll} t.\IU/.111,) th. 2 ., Jho welded th. 12. 2~. Y, . .. "ik1tt~rapt11r has,, modified verswo on
to the i:Ke of the 'icul.1, .md 'o tha..- ,1rc t\vo cro:.'- of this stmcturc \\ htrh i~ even more complex. rlt.
ing cauab. Till' 'D. iliJidm' t) pt· would 'ccm to be The on gm of the ~candent bN.• nal graptoloid.'i lm un
pnnucivt· .md to n:prt·,ent the ruoNock type of been much disputed, but some evidence sugge\ts .1 tor
prox-imal cnJ, in gt-.lptokHJ,. Cooper .1nd Fancy common ongm of bmh dtplogr.lptld and glosso- ial
(1982). however, h.IVI." 'hown that it'' the 'D. cxfell- graptid types fi·om a bm.td le.lt:..Jike scandent rhr
sus · cype wh1ch 1s ,Jl wally pnmitive tor graptoloids. dichograptid, such .\~ .'l Jiic~{!mpws, rhough the prob- .Ho
and 0. biftdus mo1phology •~ 'econdanly derived. In lem of how the crossmg canals ,JfOSt' i~ t~u from wh
~pecies ~uch a~ 'Ji·lmf!mJ•Im bi~sbyi, tor all its apparent clear. est
complexity. there: .uc ~till on I) two crossing canals. Monobrraptids prc,umably arose from a diplo-
since rh. 2' j, uot welded ll> the front of the sicula. graptid ;lnce~tor, hur there arc no dtrcrt link'
Lcptc~~raptu.1 li\U,Ill} h.l' two no,,ing C'Jn,tls, .md bet\veen them. As .tlready shown, the monograp-
though ll nuy hJ\'t' threl' Its dcvdopmcnt is han.lly uds' eh. 11 .tnscs from .1 north lov. on the mctlSIC- Tl' ll '
mod1tied flom dut of rht· bter ,Lchogr.lpttd-;. The ula, and rhe sl'rl.llly arranged th. 2 1, .'\ 1 , 4- 1• . • . grow
dtccUo- .11H.l d•pklgl.tptids lll'\Tr han: less than thret' upward.~ at an angle Bur the change 111 proXImal
crossmg l m.tk T'hl." \tl uc.:turt· uf the proxunalcnd in end stmctures trom tht· compkx .111d twt~tcd W'tt•m
the ~candenr dtplogr.tptlds difli.·r-. m.trkedly from of the diplograpltd pretur.or. to the mnplc ron-
that in thci1 prl'l Uf\lll,, Jnd the thl"cae .ue uften \tmcnon of tht· mnnohrr.1ptid proxunal end '' ven
rwtstcd up rogcrhl'f in .1 f..twr. Dtplograptids lllJ) be suJJLn ..111d rhert· ,\rt' no f..nown mtt·nnt·diatt'\.
a sep tate (t·.g Drjl/c•~r,rplu•). or thq· 111J) have a Tht• earliest known mnnohrrapmh have been
m edia n s e p tum sepJr.ttutg the latt'r thecae so that tdcntlficd Ill the lowt•n nmt Ll,mdnvcry monograp-
there ~n nom1.1ll} l)nlv tlut·c crm'img can.tls rJther nd genus --!ttll'c~l!f<1Jlfll.,, \\ ht·re the thecae .trt' of \1111-
than ..111 mdctimtt' numbt•r. fhm the latcr-tonned pk construction and rhe rh.1bdmomes arc strJighr or 1•1 c.
thec.lc {usu.tlly thmc 'ut·c.:t•t•dmg th ~ ) open on gently curved, but the pro\llll.tl end j, of t)'plcJl
the umc sJdt of thl rlleth,tn I11H: .t\ they Jrtse. A monogr.1pud tUlNI'Ul twn. Thi' b'lVC\ no help w1th
clo~t· :Hl.l!ym of proximal cnJ structure in the the problem of mom>hrraprid nrigm,, but in the
Diplogr.1pt:1l."eJ n·n·.tls no less th:1n nmc separate Engli~h Lake District in the lowermmr zone ofth.:
'astogcneuc' p.mem' (Mat hdl. 1YH7). These sepa- Silurian the re .m: some bl'dding planes that are cm-
rate structttral p.mem~. omt· e\tablished, were con- crcd w1th specimens of , l tlll't~(!rnplll.\ cc•1•:. in wluch (h
served during evolution and thm proVIde a reliable the population appt•;m LO be dithyri al, i.e. conSI\t- (M
guide to phylogl'll} . In p;t'IH:ral terms, wlule new ing of both u ni~eria l and bisl'riJl individuals. the lat-
diplogmpt.tcc.ln proxim.1l-end p.merm appeared ter being ~ i mil.1r in many way~ w Glyptt~~TctJ'IH\
succes,ivdy, rht•y ht'tJ illl lt·~, complex through (Rkkards and Hutl, 1')70). Pt•rh.tp~ it is in 'uch
time. The cl.l,,ltic,ttiOn .1dopted here lS based par- populatiom that monobrr;1ptid origins arc to Pl'
riall) upon \Ut h pm:-.1n1.1lcnd ~rructures. sought (Rtckard~. I974). Prc~umably sclectwe
In CltmcUtl~lclJ'III.I, tor imtanrt. th. J 1 ,1rises from acuon on ~ud1 populatwn' re,ulted in a changed
eh. ~ · , "hcre.J\ 111 Dtpltt~r.tptrls th. 3 1 ansc~ from th. ecolo~cal balamc, wnh tht• Ulll\t'nal fonm ewntu-
2- and th. J- trom th J . ally beconung donunant.
Similar 1n devclopmcnr to dtplo~rrapod~ m many lt •~ worth nonng dJJr there \Vl'n: 'ome 'talsc
way\ an· rht• diccllngr.1pt1ds, whu:h hke\vi'ic have monograpnd.~' 111the carl~ LIJnJovef) as wdl .15 Figure
three crm~ing t-:.Jn.JI\ \\'lth c unomly t\nsred early specie~ truly of monobrr.tpttd t) pt: p,.;,.,tglllf'll/r
change
1958)
Evolution 335
known only limn 'penmt•m ti·om the Ll,tndovcly of markl'd evolutiona ry diversiticatton in the [ udlnw.
Anticosn hl.111d, C.mad.t, ha' tncompktc d1plograp- Thereafter the ~T.lptoloids detlinl.'d but therl' were
nd development \\ ith a 'nll!<ll) th. I' oppoSite a \Oilll.' new rcpiJet.'ment stocks and mnovanom nght
range of Ul11\t.:'ri,tl thecae: 111 Dim•~'1'1W\?ruptus and tu the l.'nd of thear lmtory.
alhed ge-nera the rhJbdo,omc heg~ns hkt> th.lt of a
nonnal monug'r;tpttd (thou~h l,tckmg ~omc d•ara~.:
temric featmc\) but distJlly becomes bisl'rl.ll. Thecal structure (Figs 1 0. 14, 1 0. 1 5)
T he .·ltoll't~l?'•tpllls \tock, '' lurh gave mt: to all the
l.ua monot-~l.lptld\, wa~ cmmcntly succe~sti.tl. and Manv graptolmd\ arc defined Jt the genenc and 'pc-
once the Lll1hcnal pattL'rll h.td hccn c-stabh~hed m otic kvel based upon the \tntrturc of the thcc.u:.
At•ll'cwrclptw 111 the carlic't ~1lun.m. a great bu~t of The dKhograplld' normally have \traight: and sunplc
umscrial ~'Taplolotd~ with m.tny ~hort:-ranged genera thcr,te. Similar thl'c.ll' arc found mdcpcndcutly [n
rook place in parallel with a f1ir expam1011 of blsc-r- ~ome monograptHh and aho in certain b1,cnal
tJl genera l oward' thl• l·nd of the Lhndovery. graptolo1d gcnl'ra. Charactcmnc genera th.ll are
three long-r.mged stabk g.:ncra \Wre e~r.tbhshed. descended from the dKhogr.tpnd~. such ,\S the
.\ltliiO,I?r•tptm ''·· ,\ldllotfllll•lll~ .md Pn.W<~\?rapfrts, baseml Gfypt<wmptll.\ and Ortfw~rapt11s, have gl'ndy
\\ htch survtvt>d .1 severe cxtllK tion penod 111 the lat- curved thecae, wherea~ Lcpto~mptus has exceedingly
~st llandovery to earl) W l'nlock. thin thecae wluch arc more or le~s parallel wtth che
The Wen.lolk was a pl'nod of evolutiOnary con- common canal. In some of thl.' Upper Ordov1uan
~ahd.tnon and \tabihty. but produced bttk new. By groups a sharp-an~lcd bend (genic ulum) '' devel-
mly Ludlow tunes only thl· primograpnd stock oped m the ventral wall; rim i~ usually JS\OCI.tted
remained fhts was the \t1urce of J ~null. but w1th an incurv~.-·d (introtorl.cd) thecal .tpl-rrurc
genoculum
(b) L~ptognptus (d) Glyptogreptus

(1) Dicrsnogrllptus (cl Orthogmptus (e) ClimacograptUI;
If I p !? >
If) Early
M . revolutus
(h) Simple (j) lobate (k) Hooked (m) Triangulate

(M. cyphus) (M. loblferus) (M. prlodon) (M trllfngullftus)
I~
(n) Isolate
(Ru tntes/
(p ) With lappets
(Cucullograptus)
-r
F.gure 10.14 la)-(e) Thecal morphology in some Ordov1cion groptoloid genera; (h)-(p) Silurion monc:>graptids- (f) and (g) show
mange in theca shape away from the proximal end of stratigraphically early and la te M reva lutvs (x 2). (Redrawn from Sudbury,
1958)
336 Graptolites
t Iwe.
rt'VI."J
by si
1110d
nwtr
.1rc l
'traig
h.mg
rhe la
lt.\Vl'
In
~ihle
th(' (
I 0.15
~~I
"'::ac:
l
M:
~ I"a
a
~
::.
. •
~
d
~ I from groyp 01
M ausreru:,
Silun
Ri,k:
tion.n
g>
~ • 1 l'tiOll!
lllll'iel
•• ~ si111ila
~k·nt
"'::.
~.., thl'l'.d
l.ttl'. l
triUrlQUI•H thecae ontv pa~red rastnuform thecae - paored rastntoform thecae dl'\ eh
ilp<rturttl sp1nes or horns apenural horns - s•mple apenures
Ll.c11d•
Figure 10.15 Evoluhon of some triangulate end rostritifonn monogroptids in Uondovery times (Redrawn from Ridords e1 al. of the
19n.) dimor
( hher
llll V<ll
tbmt>, often t~1ong JJret:dy into thl· b.tsc of the curving thecae. Above thc~c come faUim \\tth hut al
thcc.t ,tbovc 1t .u1d lcavmg little room tor the zooid si m plc and triangulate thecae, where,,, Upp<'' .md tl·
to cnmc out. 1.\u~ h thecae arc cluractcri\tic of Llandovery genera have mamly lob a te wd hooked y.rJptil
thnano- and dKcllo~.1pt1ds, whcrca\ the bi~crial thcc.tc. Jt Jll c
dtm,Jcogr.lptuh h.tvt· gcnicul.uc thcc.tc With a hori- Tnangulate thecae have tiny apcmrres .1ml m dU\Ull
70ntal .1perturc .md ,, 'tra1ght outu thcc.tl wall above btcr.tl VJcw are mually of triangular fonn T1 l"i lully q
tht• geniculum, p Jr,llld \\!th the mcd1an lmc. rc.1ch their e>..'treme devclopmenr in the isolate rhe- h,,, be.
WJthm the mnnll)..,'1<1ptlds therl J\ often tonsidcr- cac of Rastrites (Fig. 10.14n), where ead1 rhct.l tl IIOnllJ
:thk tht'Cal v.mauon. not only between genera but exceptionally long and IS 1solated from il\ nrigh- \()ft.',ld
;tho within a 'ilngk md1v1dual. In .111 monograptid bouT'\. Hooked thecae have the shape of .10 open the urt
'Pl'l'it'' the thec.1c tlJJngc ~hape ,tlong the rhabdo- hook, whereas lobate thecae are tom1cd ·'' ~hort, tlwrc i
'0111\:, and the pro:\1111.11 thecal m.ty bt• quite differ- th1ck and compact hooks each inturncd in a W\11- R,htritc
ent from thost• of the d1stal end (f-ig. I0.1-lf,g). If hkc manner to embrace rrs O\\n aperture. (whit+
tht·re j, tor\ll"ln in monograpud thecal' 1l i\ mually In the Wen]ock the mo~t abundant graptolt•l ~ prnxin
au nut\\ard tWI\tln~ (retroversion ) ,1s opposed to have hooked thecae, and there are aho cl tiUillhl·r of rion cc
the mtTotoNJOn of tht· Upper Ordov1ci.m genera. cyrtograptids; these are graptoloids wikd in J hdr- ll:lb Ill,
The b,t-;ic form of the thecal 111 monograpticb cal spiral fonn, \VIth rad1ally arranged \tr.right ll! Perhap
scrvl'' to define 'lltrt·s~rve faun~ whtlh can be u~cd curved lateral branche~ (cl,tJia; Fig. IO.Hc, IO.I!iJ i~ the t
'lrat1~•rapluc.tllv (h~"i 1«1.11. IU.Hh-p). Lower C') rtograptids can often be very larg~ Jnd \\tre tlwcac
I l.lndowry mono~tpuds haw \tr,ught or gently apparently free-fioaung. In the Ludlo\\. din:rslt} 1 the tht
Evolution 337
theca.! tom1 rc.tchc~ a pc.1J... l'vbny fonm luve tl1esl.' the new type of theca came in at the proxtnlal
reverted to .1 ~irnpk· thectl t}pt·, bur rhev c.-x1~t side end and spread trom lt, but there are ~omt'
b\ ~1de wHh gcner.1 tn which the .tperrure' .m.· very ca'e' where di~ul mtroduct1on was tltl mk
modt.tled (.l\ 111 Cuudlt~~'"J'"IS) or pminwh ''Ym- Urbanck (19nD) h,l\ posmlared th.tt certain ~'TO\Hh
mcrncal. l'hc "-,-.,pruluid' ot the Low~·r I kvonian ~timul~ting ~ub,t;mcc:s like the .1ux 1m of planb dlf-
.m· quJte dJH'N.' 111 ther.d lonn, with hnok.t·d or t.uscd from the 'icul.1 as the ~raptoloid g-rew,
1tr.1ight th('CJt' h.!Ving cnwl-likc .tpcrtur.il over- promoting ditTcrenttal curvature 111 the proximal
han~. Apparently gr.1ptulu1d' Wt'nt on l'Wllving to thct.le but uot atlt:l'tmg the lattCI thecae. ~inn: rhe1r
the List. and tht·n: j, no dt·Jr n:.t'illll why thq· 'hould qu.mm~ decreased dunng ontogeny In the dc,ccn-
have bccomt• t·xuntt. d.m.b of the~e ~:arly fonns thl' growth stimularors
In ~ilun.m g-r.1ptolotd.' e~pen.dly, 1t tu, bt·t•n pos- continued to be produced from the ~icula unt1l a
\lhk ro make det.ulcJ micnwvnlurion ary \tudie~ of l.ltcr stJgc of development. Where new rypl·~ were
tht· change' 111 thec.1l ,IJJpt through tlllll' (Fig. Introduced diHally, Urbanck a~~umed that J growth
0.15). mlub1tor, rather titan .1 stimulator, dlffi.1~ed from rht
Many detailed lmeage' have been l'\tJbh,hcd for s1cula. Although the operation of tlm ~ystem cannot
S1lun,-Dcvom.1n graptolmd' (Sudbury, 195fl; be: proved. it rcmJIIIS a \~.tluable anJ interesting sug-
ll1~kard~ et ol .• 1977). In the,t• ,\ numbn of evolu- gc~tJon.
UMary rrend~ h.wc been identified, which runouo;ly M.my lines ol dc:\Ccnt arc: now known in grapto-
enough atfertcd '>l'Vcr.ll t'n1lvmg bne~gt''· both llte\ and arc Jl'curatdy tied m eo stratlgraphy
um~crial and b1,enal. at arou11d tlw ~.u11e timL or tor ~omc:nmcs, bowevt:r, LCrtain fi)nm may appe;1r.
unular time 'P·KC<;. ~ucb tl'l'lld' .lrl, of t'Olll'\l', cvi- very \lmilar to tho-.c in the: mam !me of cvolunon,
,k·nt m the p;em·r.d171:d str.ltij!,l aplul ~till e\\lllll of but om of phase with thel>e by ~t'Vct-;-tl millim1 yc:ars
thct.ll type' in Monugrapnd.1e (huokcd, lobate, i'o- (R1ckMds, 1988). Such .m.1chronistic evolution may
llt( ctc). but ,\re .1l'o cv1dent for cx.unplc:, in the bc 'heraldic', 1.c. ,utteddting thl main group, .llld
den~lopment ot thcc.tl don~.llton in the unrelated h.wing ·advanced' lharJctcD too early. On the
lhndowrv gem·r.1 Ccp!l,rh~~'·'f''IIS and R.wmrs, .md other hand it n1.1y be 'echo!(', the graptoht~:s bcmg
1! thecal p,ol.ttHlll m the Ll.mdovcna.n r.l\tntids, 'p.1k echoes' of former succe~~ful rypcs. Till: hc\t
!unorphohrrapt1d' .md non-r.1-.triLid monot,rraptids. ex.tmples here arc the partially biserial st1pc~ of
, )th~o:r trt·uds a!Tect the tonn l'f the rhabJo,Oine, e.g. Dimorpltcwraptus and similar gcm•ra, which appl·ared
ruruture whid1 ''a' domunnt m the Lbndovcry m the early S1lunan, up to 3 Ma after the orib.;n of
but ~ho tound mdcpcndemh until the Dl·volllan, th~ rnonograpttds from which they wne dcnved,
nd tbc ~cp.tr.ttc .md common tcndem y 111 muno- but looking something hke their remoter diplobrr.lp-
i!l.lptld ~tock~ to produce very ~lender rhabdu\lli11C\ tid ancestors. T he timescale of a heraldic-nl,lm-
ltln t•arly 'tagc 111 t'voluunn, .md more robu~t rhab- lim'-cchoic rcbtiomhip may extend over 20 Ma.
1\somcs Jt .1 btt•r ~t.1gc. Tht'\l' .md other trend~ arc The heraldic and echou.- type' an: normally not VCT)
tilll• docurncntt'd b~ RKk.1rd' et ,1/. ( 1':177) .mJ, a~ succe,,fi.,J. Perhaps .•~:~ R.ickardl> tndtcate~. the full
1 been knm' n tc1r \OlllL tune, nc:\v thcta.l t) pes .uc genetic potential for a partiCular morphological rypc:
1om1311) 11Hrodmcd .lt thc proxHnJ.I cuJ, later w.ts ;llways prcst·nr in a particular ltnc but could unly
lpruding .llong tht· rhabdo~ome m de~ceiHlmts of be realized when time and environment were right
''• onginal typl' Thu' tfom \/cll/tl~raptus tnttii\!Uiatus for 1t
rr I\ a trend to\\ ard.~ 1~nl.ltlllll of the thc:t"a~, with
h•mre.s 111.1.\llllll.< ·' ' .m end pomt and R. Jlc'll:~nmts
11luch ha' tht• more t~ol.ltt> thecae confined to the C ladia
l~uXJm.U rq,pon) J\ an inrermrdi.1te; in any popula-
!:On mllectt:d from a ~mgk bt·ddmg pb.ne, tndlvid- Cyrrv,eraptus, amlHlbr<;t other genera, produces lateral
ITlJ) vJry brr~:ltl}
m their st.lte of 'advann:ment·. branches (dadia) at mtt:rvals. The development of
.t'Lip\ tht· b~'t documented of these Stlun.m ~c:rics the~e .:an be followed m well-preserved matcnJl
11c trend from .\f. ar(lt'lllrll.l, where only the c.trly (~1~. 1U.8d). Each theca hJ!i :t hood-like tappet
>CJt are honked, tOW<lrd~ 1 ' l. pritJdt,ll, 111 whiCh all w1th J. pa1r of short lateral whiskers proJecting from
• thcc.1c h.1ve the fom1 nf open hook' In aU of st When a cladium develop~ fi-om such a thel.le,
338 Graptolites
one of th~ two wlmkc" clong.He' and b~comcs a mc,hwork of gtrdcrs (li't~) whtlh lll;ly hwe ;1 rcttm-
nem,t N~w fmdl.1r tmue ~ow' .tlong thl\ '" tf from l.lte net covcnng Jt (Fig. 10.16).
an lllltlll buJ. Tht: .tpcnure of the basal tht.•n j, con- In some renotinds tt ~cems J\ 1f th~: pendl nn 1'
stncted b) the t-.rrowrh of tlw tiN.·U.tr tl,~uc but \imply rcdlll ed to a '>C:l of girder~. wlm:h out.bne the
reman,, open. H" tht· tunc th~· tir~t theca of the new 'icul.l and thecae. In others, however (B.Jte\ .1nd
cladltllll ha, dn dopt·d. the pnman 't1pe h:.t~ Ktrk. I 9H.J.). the s1cul.1 and first ti.'\\ thecae d1·vdop
ad ...-JJit:t'd by sonll' wven or e1ght thec.tt. \u11..c tht.- m a gener.tlly dtptograptid modt•. Then spme' grow
cal fonn 111 C)'lll~~'·'l''''·l ch.mgt·, on gom~ ,1long the from tiH.' sltUI.1r apert11re and l111k to fomt .1 curious
rhabdmomt·, dll' ch.trancr~ ot any one thcr<t on the thin ring, the ancora , lying normal to the .txi, of
pnm.trv \llpe ~:an be m.ut·hed t \.,tcdy with one on the rh.lbdthOI111.' From thts therl' growl> up .1 ,kl'lc- l>
the d.tdltnn; ah\.1\' there j, .1 Llg so th.tt the fif\t ton of hst' .md mesh work wluch mand~ rhc rhelJI t
cladJJI theca j, p.tr.tllekd by .1 theta on the prirnar. skeleton, hut he, well OtHS1de 1t. The primal) thrlll h
stipL seven or eight therae .lht'<ld. Both emh of the l>kclcton 1s rhus J qmtt' ~eparatl' \lt'Urturc !Tom tlm,
rhabdmom~. on the prim.trv \ttpe .md cl.tdnun. have consisting of only tht• ~tcub Jnd .1 few thcr.Je. The p
been grnwmg 111 the ,,tmc w.1~ :H the s.tm~ time. outer ml',hwork wh1ch has elaborate canal~ nmning
Though best km m n in Cythlgrapt11S, cladia are through 1C 1s a second:try structure. and a' Yll it> ti
also present in u:n.un other gcner;1. DPP't'fSc~(!rapf/1.\ function 1~ mcomplcrely under~tood. Possibly the
ha_~ J s1cular dadjuut, \O that tht· stra1ght mam ~ripe P'
ltght, '>prin~ty .,J..dcton m.ty have .tddcd m buo} .1ucy. Fr
grows 111 the opp(Ntt' d1n·uion to tlw c Ltdium. or cnabkd the retlOitnd~ tO colom2c more turbulent fo
There nuy be .tt.\.l'\\Of\ btl'r.tl d.1dia growmg etrhcr ".ltt:l'. Jud~ng by thc1r pemstl.'nce through rime,
from the main ~tlpc or fro111 clw primary cl.tdium. they must h.1vc been .1 h1ghly suc~:essti.11 group.
The rd.ltt:d genm 1/lirs,(!rapttH h,t\ :t number of sicu-
lar cbdt.t .
Structure of retiolitids In general term~ tubmds, camaro1ds and nu,tuid' !>ll

were encru\tmg colonies re~emblmg in hJbtt thl \Cl
The renolttids .trl' a 1-rroup ot \C,tndent biwri.ll grap- rhabdopkura n he mic hordJtec; I kndrot~. vn thC' toI
tohtt·~ whtt: h rangni trom tlw LIJndeih,u1 through other hand, were gcncr.tlly upnght shmbby benrhm
to tht.· ~·nd of th1 '-,iluriJn. l'hey ditrer limn othl'r .md the grJptoloid' to which thl·y gave nse and Rh
graptohtcs in tl1.1r tht. skek-ton l.ugely <..oumt' of J which the> outlasted \.Vcrc the only wholly pl.ml- \VC
(he
(bl the
pl<ll
org
pla1
bite
shal
as t•
Thl
197
the
IHCr•
197.
beea
..tcti\
are
100~~om
SICUJ.
Figure 10 16 Retiolihd ullro$1ruclvre: (o) Orthorelio/,tes homi (Ord ), Oklahoma, proximal end, (b) Gothograptvs d intermeclas by I
(Ord ). dothriol network, from on erratic boulder, northern Germany (Photograph$ courtesy of Or Denni$ Bole$ )
zooi<
How did groptolites live? 339
t<llllt' order. The b,Hrom-dwdhnf.' dl."IH.JwiJs and
t)tht:r> art: found 111.11111) 111 ~1lrs, sand~ .llld lunestont~ Passive drifting
t)f \lullow-w.Jtc•r 1mhnn.· origi11. l'ht:\' have no [f hrrilptoloids were drit-tcrs they would nc<.·d ~c11nc
tim.trionJI ~tructun·, .md tht· corric.11 u~'uc· of den- mechanism to prt:vc:nt then• s111kmg. Pm\tblt· .ld.lp-
droid~ ~uch J\ lJmdr•'l!folpllls appe.u' to h.1vc been t.ltmm tor tln.1ring .1rc: llliiiiCrom: mudl'fll urg.uu'm'
u•c·d pnnuril} for srrcngtht·mng tht· proxim.1l end of .Khie\ c Aor.1tion etthl'r by ha\'mg ~a~ or t:1t bubbles
the rh.1bdmoml.' mto .1 ront-likc hnldf:N. J'hu~ tht: Wlthm thctr ()S~ue~ or h) ha,,ng the ti'\UC\ i,otonit-
Jcndroid' cJn hi.' c'll\"1' tgt•J ,h h,tvmg gfll\\'n upright wJCh th<. '>Urround111g \t' .1 warer. Uulm.m ( JtJCt4).
1lll the \l.'.l floor With the \ICular c•nd down .111d tht' Rid•.trd' ( IIJ75) ..md orl1l'r. believe th.tt g.t, bubble'
'llpt:~ ouhtrc·tchl'd .1\ in ,1 \111,111 shmb. W1th the con- \Vtthnl th<.· twnu, or v.auol.ut·d cxtrathcc.tl ti\~Ue
tmunl addition o( n1rt1t 1l tJ<;suc· tlw ~n:m would cont,linm~ riny g~' pm kers. wott.ld gJVe eJJDttgh
h.l\"l' been grc.1tly 'trcnl:,rtht·ned. but tilt' t'.1rly thecal' buny.tncv. Such C'\tr.tthc<al msue ma) h.t\<.' been
\\ l'TC Ol cJudni cllld the zooids pr<'\lllll:tbl) .ltrO- ,c•netcd by and J\SOCI.ttt•d wirh the open t:nd of rhc:
pJm·d. However, /<,Jdii~I!'·'PIIIS and JVr.r/,ditll'/'''"' art: nl'Jll t; 'upplement.lT} flourion nlechJnl\111\ of \'.In-
wnitJI rathn th.1n ,hruhhy and h.1d little cortical Oli' kmd, may h,lVt' Ktt•d .h .1 support for \UCh \ oll-
ll"lll'. The thn.:.td-lil-1.· nt'lll,J cnt~ld nnt h.we sup- UOI.ttt•d ti ...~lll'. Vane~. onct' thought to bt tl.lttcned
pnrtl'll ,\ rhabdmnlllt' wuwing on the· ~t\1 floor. bbddcrs, art' nt)t uncommon in the KJndent com-
rurthennore. ~tll:lllllllt\ of thin fibre<; .m: ~ometimes p.tl"l d iptobrr aprid~. Tht.•y fomt lateral extt·nstons to
flnmJ tcnninating the· m·m.1. wh1ch could haw thl' 11Clll,l; ~OITlC graptoJOJd\ !1JVC tWO \',\111!'' .11 tl-!0°,
;lltt"J .1' iix.ing dt'\ ''l'' fi,r the attachmt•nt of tht: otht.'T\ h.1\'C three ~t I ?1J 0 [ f rhesc were tht.• ''tt·~ of
lillalruid to tlo.tting w.t\\'ccd. Tlm nuy be ~ug v.Kuol.ned extr.uht.·c.ll ti\\Ue, they Wt)uld h.tve
~'tcJ b) the prt'\t'IH l' ,,f much t·.ubnn Ill rhc hl.lck impartl.'d buoyancy .md \tlhtlity w rhe ~.tpwln1d .
1h.tlc' m \\ hll 11 nl.lll) ~·nptohtt'' .1rc found. Proxun.tl end V.llll.'~ or pbte' occur rJrdy ;md thetr
l'rc,un.ably R/r,d•rlillt'Jir11tl e\'oh-t•cl from dw standard ti.mction is unclear.
ll',~ilt: ckndrnid' b" 111VeJ""\H)n of tlw rh.1bdo~ome, St>llll' dichograptid~ h.1vc CX.11Jnded wdh ~ur
lll"ular end up. thnugh .m \1pnghr' oncm.1tion n.nmJmg the proxim,1l <.'nd which lllCrt',l\C the
M:l'lll' to havl' bct'n n•,wred m the· \t tndl'nt gr.tp- \Ut t.tcc• .uc..t and n1.1y h.l\'e prevc·med \mkmg.
toloiJ, of the i.ltl'r {)rlhl\'iCIIO and 'Ot]un,m. Cert,un 'pir-alJy coiled w.tptoloids ( Cyrr,,l!'•'l''"' .md
A ncmally att.ttht·d JJhHle of lift• " ltkcly for ,\ ltlllt~I!"'J''IIs tum·wlatw aJII\)llb'"'t other;) wen· con-
H./t.r/tdlllopr''" .md proh.1hl} \Otne ntiH:r ~:--rr.lptolotd~ a~ \ldered by Uultnan ( ]1)(,4) J' Aoatmg t<m11s, whJCh
well, hut the nujt,nty Wt'r( probably frt•e-floating would be rotatl.'d round .md round, sptralling
(holnpl.lllktonit). minoph.1gou' ti.·t•lkr,, fonning upw ..trds in the water undn the inAucnn· of cur-
dtc m.tin prt:wrvnl p.lrl of the Lower P,li.H:ozoic renb ,tnd thus buoyt:J up. The pre"il'llll' o( .1 AJt
pbnkton In tlm there "t:rc undoubtt•til) other proxnn.tl membrane 111 Cyrto,\!mptw n1.1y h;l\'<.' been
O~lm~m~. ~omt: of \\ hu.:h , 'ud1 ·"' tlw 'm:11l epi- .111 extr.l flotatJonal :uJ. and j\ I. trrrriwlrllll• 1' now
pbnktonlc br.tt h1upud, ;.llld oct".t,inn.tl 'null trilo- J..nown to have the tight!) coilt:d sttpe' umnt•t:tcd
blt<.',, ;"Ire ~omt:tlllll'' ti.ntnd pre,~:rnd lll brr.1prolit1c by dl\sepnnents; such would "itTt:nsrthcn tht• colony
1luko~. Two oppom1~ 'tl'\\"' have bct'll put K)rward ,J'i it rul.tted.
11 ro how rhe pbnJ..mrllc graptohtc\ .tctu.tlly lived. No ~ap tolitcs have ever bccn f(:mml anmlly
lhc more tradttHln,ll 1dt·' (llu.lman. llJ(,-1; RKkar&•. .ltt~lt hed to .1lgac .mu 'urn~: ..tulhoritlc' (Kmlm'''ki,
11175) has bt:t:n th.lt ~r,tptnloid' j.'J\\1\<.'h tlu.ued 111 llJ7l: Rlckards, 1475) bdieve that lll'lll:ll ,ltt.H"h-
tht' ,,c·t:.an. pnh.1~" at dttli:rcnt lc\'ck .111d at the ment by cxtratheul 11\\Ue l'xuding from tht· np of
mercy of curn."nh. I hl' .11tcrn.ltt\'C vtc·w (K trk. I 1)69. the nt"ma was unhkdy ft1r Jnv graptolitc On the
11172) t' dut gr-.1ptoluak .1lrtr a l,tn•.tl b<.·nthtt '>t.1ge, other h.md, radtally ur,tn~t'd assoCJatmm nf gr.tp-
lx·c,m1c pl.111J..tnnt, .Is young adult,. .md were toloHi rb~bdosomcs. wnh their ncma\ din·ctcd
Ktlwly mobile. R.tthcr than pa\\IV<.I) dnttmg. thl'y tnw.1rd~ the ccntr<.·. h.tVt' been rcportl'd. Thl'Sl' arc
Jr l'IWI~.1ged ,1\ .lttl\'l'l\ ~"WII11111111p;, Wtth the known .1~ synrhabdmolllt:\, .1nd although tht'} .1n·
ucuh upv1:ard' wh1k ti.'t'd111g (urretH\ \\ t'rl.' drawn r.~re (\eC' p. 3-U) ~omt• may be gcnu111t liti.· ·"'ocia-
f11JS
b\' the rhe,-ae ttn\ .trd' the t~h.try umam<.'ntt:d tiom in which pre~umably the rhabdosOJIIl'\ \\ crt•
z(l(mh. connected by a m.l\, of l.':-..tratheca.l ti,,uc from thl'
340 Graptolites
ne mal up~. 13u bbk-hh· tlnt.tnon ~tmnurc~. Origi- and how it chang(:d through tunc are chssic pomte~
nally described at th~ n:mrc~ of 'ynrlubdo~omes, wh1ch hdp in undcn;tanding functional adapt.1tiun~.
have pron~d tu b~.: illmul') but in the C:l,c..' of the graptohtes other rc~·c:mh
The tinal c.:hange to tht.· monobrr.1pod p.lttem pro- developed ind1c1tof\ .uc aho 1 help.
duced long. ,h,-ndcr, 1~:" dt:IN.' colomc,, whiCh The branching ~tructure of multm.unom ~-.rr.lpto
~cemed abk to manag~.: wJtl111ur 'uch cxtr.1 Aot.ltlon hte~. for example, ha~ proved p.uticularly Jmcn,•ble
structures, a' had tht: diptograptids. and probably to computer a1talysis and ~inwbtion (Fig. 10.17)
made do Wtth nc.:mal van1ulatcd ti~~ut'. At the same Multiramous gr..lptolitc\ OCl'ttr rypically 111 the
nme, \mn: the nature of the cxtrathn J.l n<.~ue in Lower Ordov1uan (dichograpnd\) but al'>o .lt h1gher
graptolotds 1~ Js vet not undcr,t<liJd. thJ, v1tal gap in levels (e.g. nemagraptiJs and cyrtograptt.d~) where
our knowledge m.1y well prc\ ent :my further devel- \Ome forms become secondarily convergent on
opment of our understJ.nJin~ uf graptoloid ecology. ruc hograptids. Computer ~imuiJnons haVl' ~hown
that various br:ll1ching pam•m, JUSt like thme m
'real' multir.unou' gr-aptol01(h l.ln be gener.Hed by
Automobility pennuratiom of very simpk mb (Fortey .md 13cll.
J9H7). In mmt d•chograptid<. branchmg " d•choto-
The b.Nc conu.·pt here " rhar fecdmg currents pro- mous. t.e. each growing <;tt.pe ~plits into two equal
duced b)' zonid~ would h.tvc been powt.·rtitl enough daughter stipe~ at the samt: liiTH.', :md :lS the rolony
to propd tht.• colony. (Sml c cortical bandages wen: h'TOWS there \\ 1LI come a further zone of d1choro-
put on from dll· out~ldt· then: '' no hkehhood of cil- mom brancbmg.
Iated t.xtr.ttht.·c..ll tl\\Ut" whJLh could have aided.) l"he simple~t computer-gcnt: rated modt•l, wen·
VeT)' m.m) member\ nf tmby\ pl.tnkton are auto- b.1sed on JUSt such standJrd dichorom•c~
mobile .md there j, 1w rCJ\1111 why gr:1ptoh.11d~ could I n"tructions ..~ere given (I) for branchc~ .1lway'< to
not have lwt.n. Vane~. wdh :md other mch stmc- ,pht dichotomously at nod.ll pomts. bur (2) tn ;lln.:r
ture~ could h.l\t: bt"en ClJU.tlly functll.)nJI in mobik thl' angle of d1choromy in somt• form~ and/or (3) to
graptolotd~ .~, m Jnfi:cf\ The reduc.:uon m sope delay rhe imuanon of dichotomy. The ·graptolo.(h'
nUinber. thl· mcrea~mg wmmctry and the l hange in thm generated bear a sin!,'Ul.lr resemblance 10 the
inchnatwn uf thl" stipe~ art• .111 ~ccn a~ p.lrt of an evo- well-known quadiradiate (StmmwmJ1lus). triraru.1tr
lutiOnary rt:~pomt> to thi' new modt: of hfe. and (Bry(~\!mprus) and biradiatl.' ( ChliiO,t:raptw) gener.l of
though the whole mattt>r " very conttovemal and the Lower O rdovteian. Another permutation pro- figt
has been much debo~tcd ir rcm:um an mteresting duced a 'grapcolo1d' with Jn ~-)haped nuin snpe redt
hypothe~i' . With fan-shaped daughter stJpe\ (approxmmmg did
An .1rgumcnt .1~rain't .nHomobility is that many in ff
Ncwn}!mptr~s). Again, a computer modd \\1th
{n tl
species of waptoloid after ~um·.;ng brCJkagc could 1mrrucrions to generate curvmg stipes produced J Fort.
regenerate .1nd ~rruw m tlu: nppmlte dtrecnon. do~e match wHh Cyrtogmptw, a rnonograptid (Fig. rock
I 0.8e) which ha~ become ~t.·condarily multir.1mou,
by the regular generation of rl.ldia, \Omc ol "hid1
Use of models in interpreting the mode mJy thrmsel ve~ grow secondary cladia. hav
of life of graptoloids B ut w hat may be the ~1gntf1cance of \Uch pJt- .1dja
tern~? From the an:1lysis both of real and cornputn- thcr
There ha,·e to be funcuonal rea:;om why dcndr01d drawn graprohte' 1t IS evtdenl that brancltc' do nnt a~ <11
and graptolo1d rlubdosomc' are the slupc~ they are. tmcrtere with each other. Whereas 'posstble' models tcl!d
Such morpholog-Jc..'S re~ult, a~ m other organisms, C.ln he generated where the branches do overl.lp. W.tt<
from the interplay ofad~tptJuon by Dt!t'WIIIian selec- acluaJ graptoloid~ of this kmc..lnever occur in m tun·. or t
tion and constnint~ impn,t•d by the fundamental Such m ultiramous graptol01d~ must tbcrl'lorc h.1vc thro
design uf the organism' and by the material~ of been planar, ctther AJt or shghtly conical. Morcm·n R
which the)' 1re comtmtted. The ultunatc form of the ,tipcs wtthm th1s flat plane are regular!) arr.nt!(td mou
any such or~.lnl\111 J\ d1U\ a t.ompromJ'e between wilh ~pace" between them .•md m some 1mt.1nre\ been
mherited con~trJints .md .tdaptive posMbJlitiCS. The potential dichotomies with in a rhabdmonll' luvc loJjtl
analysis of the JIMtomic,ll 'tructure oftht.· organism' hcen suppressed where othervvise the ~tiP""' \vnuiG F1g.
How did graptolites live? 341
(1)
-~ F-
--. '=-
--------{ r ---
------.
------, ,-----
~------
------~ LL------
___ _
~~·__1
--~:....1
____1 ~ -
L --
___l LL--
_ _j L-
ldl -1 I
(e)
(h)
Figvre 10.17 Computer-generated graptoloids and their fossil counterparts. (a) triradiate model, as in Tremadoc anisograptids,
redU<ing angle between dichotomies, thereby avoiding overlap; (b) biradiate model, os in later Tremodoc anisograptids, with distal
dichotomies suppressed and slight curvatvre of distal stipes, thereby avoiding overlap, compared with (cl Logonogroplvs logoni; (d)
"'lh1s model stipes are distributed evenly through the area enclosed by the rhabdosome, paralleled by (e) Brochiogroplus etoformis;
!; !he actuol Nemogroplus gracilis which is directly comparable with (g). the computer-generated yin-yong model. (All redrawn from
ftw1ey and Bell, 1987). (h) Computer-generated deformed images of lsogroplus (left) which simulate actual graptolites in deformed
'Oeb, useful m slress~tram measurements (after Williams, 1990)
)
have grown too dmcly. I [we crwt~Jgc the zooids of These were accurately coustntcted m.tinly at life
1djau!nl ~tipc' covering the feeding area m bet\vccn, size, of aluminium tubing covered wtth 'dingfilm'
then .1 multiramous rhabdosomc c.tn b<.' mterprcted to simulate the density of a living graptolitc. The
~ Jll dfn nvc 'harvc~ting mJchuw' It could either models were ,tllowed to fall freely through a column
s twi bv ri~ing or ~inking verncJIIy through the of still water, m horizontJ.I orientation, .tnd their
IUt~r. while the zooid~ ~rrJincd off the nutrients. behavtour was observed. Most of the mulmamou~
r by rcmauun~ stationary whtlc nrrrcm~ passed fonns and Nt'ma~raptus models rotated ~(owl> as they
e duuugh u sank, sptralhng downwards through the water. In
r Reccmly, our conccptton of hO\\ \urh multira- life such fom1s would have been able to ·,weep up·
d ou~ (and other) rhabdosomc'> JctuJI!y behaved has or harvest nutnents as they fell, aided by the spiral
(Cil rcfinetl by the use of cxpcnmcnt:tl model gr::tp- rotation (which allows a longer path th.m stratght
'Jlitc~ (Rigby Jnd RirkarJ.,, 1981); R.t~by, 1991; fall, v.rithout overlap between the p.tths of the
itg. 10.1!:!). descending zooids). The models ,tlso clantlcd how
342 Graptolites
\u
Ill I
wi
\
f
« \) of
\
~ l on
Qt
In
·l
\ ~~ /l \ .........r
fin•
r.ttc
po~
·rh
b'T\1)
"''
~ (C)
rh.1
(a) (b) of'
~pc•
\till
re.tl
rap1
t wn
Jc:cr
ton.
Figure 10 18 Morphological adaptotians which cause groptoloid mobdosomes to rotate, os tested by physical modelling: (a) prox· tolo
imol hook on mooogropltd rhobdosome, (b) vone on biserial mabdosome, (c) thecal olfset ()(1 a dicellograptid; (d) Cyrtograptus, Wilh bv r
spiral form, dodio and thecal offset; (e) Nemogroplus would rotate only if the rhobdasome had this twisted S·shape, rather than dl'll(
being Rot, (f) Logonograptus would rotate if stipes were angled relative to one another, os in a propeller (Redrawn from Rigby and \0111•
Rickords, 1989.)
gr,tp
the ~llpt:~ w~rc probably :trran~ed 111 ltfc. Nellla_~r<~pltls The .u.lvant;lgc of spiral 611 is l'Vtdt·nt both 111
models, for ex.tmplt:. comtrmtc.·d ,,, plan,tr fonm, large mulurarnou~ fonn,, \Weeptng out .1 bro:ul h.tr-
d1d not rot.lt~ . u: howcvt·r. the matn '->-sh.tped ~tipe vestuJ~ path, .md iu rhm, scmdent timm cxplottllll!
\\,1\ bt:nt 'ymmctril'Jll) ,tbovc and bdm' the planc. a n,trrO\\ path. No rh,tbdmomt:. ho\\'t•wr, could The
w tth the. senmdary snpes tomun~ :1 ·-.pJr.tl ~t.ur\.'ase ·. lonttnuc tJIItng mddirutcly· 1t "nuiJ t'\'l'ntuJlly ka.\t
then thl· m odd rotated etfe~. ti\'d\'. In ut ha multira- have lll n\e, which prc,uppmc' 'lllll(' dq.,'Tl'C of (~kl'\
mou' gem·ra (L~I?o711lJ~I•TPtl15) nvt,ting of the supes at automohility, or at k.t~t thc rapacity ro L·h.mge lt\ Ut:rlj
.m .mgk to one another a' 111 .1 pt~.)pdler hkcw1se buoyancy. Tht• nature of \uch pml't'''c' m the cJIIy
faCJht.JteJ rotauon, and m C)'rttlgmptw titrther a1d ab,cncc o( dirccr evidence is snll UIKt'rt.nn; hut the a cu•
wa~ gn t:n hy ~ec:ond.try cladt.t. In this genu~ use of both computer and expcnntl'llt.ll lllodcl-. h.t\ Arcnl
(Undcrwood, I 995) there ts ,, cotttpkx suite of al least p.trti.llly rtTOIKikd nppming hyporhl'st'~ , were
owrl.lpping web\, connect111g tht• t l.tdia with the and added a ncw Jimemion to our undt·r;t.mdm~ nf thl' '
m.tin \npe (F1g. I!U~e). Thr~e Jrt• n11nposed of thin how gt.lptolitl'' lived. 'Pant
pendennal sheet' Jnd gwr tht· rh.t~dmome J screw- Graptolotlh \t'l'm to han. b~.·u1 .1 primll) cnmpl'l- ~imila
hkt· fonu . They m.ty hJve acted to direct water into ncnt of the p!JnJ...ton of dH: PJI,tlO/llil', .md g~ner Engl.J
'ep.tr.ltl' L h.mneh as Cyrroi!'•'Jifl~> rotated. alh tropit al and temper.tt• 111 Jt,tnburinn. 1 hctr Euro!
~:tn y model paucinmom ~.tptohtn also rotated. common ''"m iation ,.,.;rh black \h.tll· (b'T.tptohnc l3oliV1
Dud/cl~l·l[lfm h) thecal otEer. bi\t'ri,tl rhabdosomes facie~) wilh m h1gh c.u-bon tOiltclll and pre,ence of Pacifi.
~y \'Jll~' (\\ hether b!J.ded or t\\'l'itt•d nhbon rype), syngcnt'llt pyntc ~uggest~ that thc~c pl.mkt011ll' Argl'll
.md ntrnng monO!-,'Tlpttd~ by thl· curve itself. organi~m~ were preserved in roll(htwm \\here no 'ighr
Strai~ht monob>Taptids, 011 the ntht·r h.md, tell with- benthm tllnm~hed; then thcy would lw .thk tn Pl' ln:l,\IH
out rolatHln. prew1 wd undisturbed by bioturb.ttmg ,tnimJb. kttl·d
Faunal provinces 343
liinn~ thn: are found in 11tht-r kmd' of ~L·dimt•nt thc:v .fuirly du,c re\cmblancc tu l'.lt h orhcr at thc ~t·ncnc
mu't hav'c been "itk,rn: 1d. but tht•Jr .l'.~nli;Hmn kvd.
w1th .1 p<lrticul<~r argill.Kul\1~ t:lnt'' •~ l.lrgd) .1 m.mer 'I hu~ amon~rst other di,tinl.tiVt. f.ICtOf'i, pemkm
ol pu:~ervarion tiilurc d~L'\\ hl'fl' {tuning f(>rl,.) DiJym<~~rc/)1111.' ~pet. ic~ art: Vl'f)' cnm-
ltJgbv's ( 19tJJ) ,tuJtL·~ of gr.tproloid mit'llt.ltsoa mon 111 the Atl.mttl but not in the P.tcitir pro vi tll't'.
)n bedding ,urfan·, 111 thL Mtddk Ordtl\ iuJn of wluch ha' .1 more cl.J,·cf\c fllln.l \\ ith i~o-. t. lrdm-.
Quc:bt•t ~bowed thJt the t\\ t> 'PCCIL'' prt'\Cill (.m onro- and ~~~~o~pti<b. 13t'cn,JI scandent gr.1pmlmd~
.~mpll'.\()~raprrts and .m ( htl"'.l:'•'l''"s) re.tthcd rhc ,c,1 common to both provide: .1 !otood ba~i' tilr 'tr.Hi-
~oor Jt dttierem nmcs. Thi, 'uggc~t' th.H tht• \t' p.l- gr.lphJc.ll correlation.
utl' ptlpul.lrions lived ·" ntunmpt.•citit ,[1\l,tl\, .tnd The zt·mrh of provmct.~hsm w.1, reached in tlw
poss1bl} Jt dit1crent lkpth' 111 tht• watt·r l olurnn. Arcmg-Ll.mvirn, and 111 tht. btcr Orduviri<ll1 tlw
Th1~ "orlo.. aho gave t'\ ltkiKl' of ,c,t,on.tl growth of two tJun.ll provinces were replaced by J ~ing;k llllL'
!r.tptoloid rhabdmome,, \\ 1th ditl\.·rtllf '17t'' of What could have been tht• tuntroh of prnvinrial-
rhabJo\OilH.' at varmu' tmh..'' t>f tht.• yc,1r. T'ht• n.1turc i,m? W trh trce-floatm~ or~JIU~lll\, gcobrr;tphil.ll h.tr-
of·~ymhabdosomc~·. l l . l.ldt.ll aggrt..');.llt'\ nfllltlllO- ricr; 1rc hkdy ro be le\~ unponant than th~:\· .ue
>JlCOitC ~.lptolmds pn:wnt ht•rt• .md in orhtr lx·ds. ts wnh 'hdf-hvlllg bcntho~. unle~~ thcy crm~ tcmper.t-
mll d1~purcd. Some JuthotHies bclie\'e th.lt the~e arc turc tom:,. Such more or less latitudmal zone'
ffal •moriarious. po,,tbh inditating a method of defining rcgiom of diiTeriug temperature (tropi,al,
:aptd <1\t:xual clonmg, but R.1~b} titvour' tht ~ugge~ subtroptral, wJrm-temp~rature. cold-tcmpct .nun·,
uou that thC\ result fwm ·m.mnc \110\\ •. t.t' . organic boreal. pobr) are the most unportant single 1..111Hml
:ktrin1' \tuck together b) lllltrll~ \H'b' nf zoopl.mk- of the glob.1l d1~tribuuon of orgJmsms tod.l)- ..mJ
ton, tallmg though th~: ".tter .md collt·tting h'T.tp- prc,umJbl) rhey '\ere of the Lower P.ll.tl"o/oh."
tolmd' a~ 1t s.mk. Gr.lptduid, wcrL· l'Vltkntl~ c,Hen plankton .tho. The lantuduul thcrm,ll ~'TJdiL'lll i~
by pn:d<ltors (Uudem O<ld, I ')93). and tht•rt• 1' t•vi- also n·,pomiblc for the d1tf~..·rcntianon of m:c.mtc
j~IKC ti1r rnmching or .tb~tlrpttun of'' lwk· rlubdo- water~ into ~~:paratc w:1tcr 111,1\~c\. 1t is vcty pmbahlc
omes. or dd.Jcatl..' plm kmg tlf indinJu.1l znmd.~; (Finncy. J9R-l; Finney .llld <..:hen Xu. I 9l)ll) that
5f·1ptolmd\ nuy abo have htl\teJ '>mall p.tr,l\ltt'' 'pcotil gr:1prolirc taun.l\ wt•rc cotltincd to p.1rt1t ul.tr
\\'ater nt,l,,c,; mdccd watcr-mJ" 'PCLitint\ h.t\ bl.·en
cited ·'' thl' pnme control of gr.1ptolit~: btogco~rr.l
phy. [ Bt•rry rr al. (19~7) h.wc ~ug_gestcd that \l111le
b'Taptolitc' inlubircd the nt c.1mc oxyg~:n nuuimum
The dtstnbunon of gr.tpwiHc~ on J p;lob,1l ,ok, .ll IJycr. J
lr~>t in rhc Ordov1u.m. \\.1\ not .th\.\y' lll\lform The Pat itic f.mnas of th1..· I owt·r Ordovtn.m Wt'rt•
~kcvin!-,'tOn. 1974; r :lllllt:') ,lltd Clwn Xu. 11)9(1; apparently circumrropK.ll .mJ were cunftm·J
Bcrrpnd W1lde. }lJlJO: Rtt:k.uJHt.tl..ltJ\JO). In the ennrdy bnwccn che l.mnt~..ks J0°N Jnd JO· ">
wly 1\rcmg 1t seem' th 11 1110\t grJptohtt• ~t'IIL'l ,l had Atlanttt f:tun.l\, on the odll'r h.md, lay ~outh of
cosmopolitan d~tnbutH)n, but tram th~ latLr 30° 5, except tor the l::urop~:an region. whne
\rcntg onw;lrd' tmtll tht. end of the Ll,lnddo rh ere AtiJntil fa11na\ arc found nc-.1rcr to the cquatm. lt
\'<Wt.' t\VO well-defined W·lptohtc wun.tl provim.es- h,i~ been propu~cd th.H Lht• t\otherms more or less
he 'Adantic' or 'l.umpcJn' provinct• .tnd the foUuweJ p.tlaeo!Jrirud~:,, t•xcept where ,1 tongue l1t'
Po~eifiL' provmce - whl)\l' tiwn.1s ar~: not dmch coldcr w.1ter projected norrh\\ .trd' in the F.uropt·an
;uml.tr The Jrea of the Atl.mnc pronnc~: 111dudc~ region (a\ \\ ith tod,t} \ Pent t urrcnt), C<liT) in~ wirh
England and Wale\, 'outhe.l'tem lrdand, llltl't uf it ih own mdigcnom titun.t
Europe, Nonh Afnc.1 .md. rJth~:r uddl~. Pnu .md SuLh .1 p.ttrcm could wdl ,Kcoum tc11· 1 owt·r
!i1Jiiv1a. On the odtcr h.md. the graptolJte' ol the Ordoviuan gr<~ptohtc UJ\trthut!On, wnh w.Hcr-m.l'~
l'acific provmcc an.· li1und in Nl.1rth America, spcctfi(lty (ulrimJtcly dependent on tcmpcr.tturt·) .ts
\rgl'nrina, aU of Amtrali.t .tnd Au~tr.tl,ts\.1 .llld (.tt tirst the prim,try Lomrol. but wll\ tht·n d1d the fJull.t\
~~ paradox.icJ!lv) 111 '-lt.otl.md. nurth\\C\tl'm of the l.tler Ordovtcl.ln bt•tomc more unil~mn'
.bnJ and western Non\ ,1) Th~: graptolttc' t:ol- Ont· re.IMIII 1' thJt Famtlic' Oichogr.lptid.IL' .tnd
teJ !Tom any p.ut of ont• of tht·'e prov1ncc' bt'.lr ,, Smograptic.l.ll., whiCh h.td ht.•cn import,lllt Ill
344 Graptolites
detimn~ the two pronnlc,, lud b\ thi~ time pm~ibk '' 1rh (tor m~t.mc.:) .1mmon1tt's: (.\) gnpt~ fa u
bccomc cxnnn, bur 1t 1~ rnmt 'l~nific.:.mt that Jlmmt lne~ 1rt: nornully confinni to bl.tc.:k or ~C) ~hair C.JSt
no late1 OrdovKuu gr.tptohte' .1rc found ouhide the f.tCJl'' by prt:<>crv.tttonal taxtor., but t'lrcn gnw gco
pal.leabtlnldt:s 3U0 N and JO"\: the~ .m: wnflncd thu:knc'~ of OrdovK1an and 'nlunan 'rr.lf:1, whu:h 1
w1th1n the region fonncrh octupu~d b) the circum- nught bt: cxpt:cted to contam graptohtc,, do lltlt m rhn
tropJcal Pacitlc province. Thi~ rcJmtion of the geo- fact have .my fossih at .1ll. Furrhnmor~-, 1t 1s rela- t:nu
graphil.ll distribution of gt"<lptulitl·~ 111 the later tJVcly rare to tlnd !:,'Taprolirc~ in co.mcr-gr.unctl lln.t
Ordovtn.1n may well have been under climanc con- rock\ ,IW,IY irom graptoliric f;Kit:,, ,111d then: ar~ 1
trol. I hgh briruduul realm~ no longer ~upportcd consequl'ntly smgular prublcrm m correlating gra1
grJpcohrcs, benmc the \~:Iter; were roo cold. and bcnn•cn area~ of different fancs. spet
rh~: brraptoltrcs of the trop1c.11 zone' were unable to U~u.lll) gcoJogi~t~ Jool for ,ln.'.l\ \\ hcrl', be~.;.IU•C \\'as
.td.tpt H'l lOid condJtlOm. of 'hordint· oscillations. thert· .uc e~po,td Iern•
B\ rhc l.ue~r OrdO\'lcian (bte A,hgill) only some ~l!qucnccs of alrcmating graptolit1c .md ,hell~ fJtie>, eyuJ
tive nr 'L" ~pcnc' of graptohtc' rcnumed: :. dcct- ~o rh.tr a parncuJar graptlllitc as~embla~c i\ nmc· I Oll~
m.ncd ,rock hv111g Jt a tunc o( '' 1dcsprcad glacial bracketed w1tlun a trilobttt.~brachtOpod .L"t:mb!Jgc dc.u
condmom. from which l'Jme tht: earliest mono- Jnd hence the two may be dirculy corrdJtcd l'XCit
gr.lpnd~. During the milder dima1es of the Silurian Altern,ltJVely. the discovery of m1xeJ gr:.ptolittl Jnd y,
their descendants wcrt: able to exp:md and diversify shdly f.1tm,1s in thc same argillatcous \cdimcnt 11 .tr(.' ~
and to ,pn:ad unitonnly .md widely across the world most helpful. Even so, there ,1re suu;c"1om \\hC!r Tl·rn
once m on:. to rhe regions th,lt rh~:1r :.ncc~rors had rhe shdly f.nmas arc indtgenom to the region or Bryt~.l
occup1l·d m the earlier Ordovtt:l.\11. For a rime, a have very long ranges and where 111 the .lb~t:lllt' of luvc
tropllal Panhc pnwmce ,md .1 cooler Atlantic graptohtt'' prcn~c conclations arc not po\\1ble. r{·cor
provmle \\ere re-c,tabh,hcd. From the Ludlow. Indn:1du.tl graptolite zones arc based md d~lutcd Trcn
howeva. until thetr tinal exrinr1inn in tht Pragian, on the ome ranges of pamtular \hon-t.lugrd fh
grapwhtl·~ appear to h,l\"e bet:n contined to tropical 'pccic\, though the named speues IS uormall) llUh h 111.1
seas alone. one of a number m the total fauna. In the Europt:an tJlk.
sucres~1on 13 OrdoVlcian. 32 Sllunan and three- :tlly ~
Dt:vom.ln zone~ h.we been erected and ,\re in com- rt'Ol~l
mon u~c. Some widespread 'PCCIC\, \Udt 11 ~'T,lptlo
ClYJilt!~''lJliiiS wr~•tillmdtts. Nctllc((/f•lJlfll5 .C!ftldlrs .u1J by tUJ
To geologm~. t-"Taprohtc~ arc primarily of nm:rt·st .1~ Drcdlo,~rt1JIIIIS ,.,liiiJIIutl•lltts. arc norm:tlly rotmdc:·aJ 3! tht.· L
maugraph1c.tlmdicators. The ~eyuctllc of graptolitc n·prt'llcnnng rime-equivalent honzom. but there het am
taunas has been used tor the 'ubdivi~1on of may bt· 'ome di.Khronetty Thc probknt 0: ( arado
Ordovtu.m and Silurian rOlk, 'inn: the timl' of d1achronctry 1' a real one and corrdanom h.wc to bt \\d) I
Lapworth, whme work m \OUti1Lm Scotland m ba\cd upon ~uch events that can bc ~hown to be tht cnd~.:n·
unr.wellmg thc complcxmc~ of geologic.:al ~tructurc lca\t d!Jchronou\. The only w.1y to do tlm 1s ro US{ Lsa~mJl•
IS class1c. Their valut: for long-ran~c correlation i' ·a~ m.tny clmdy 'P·Iet~d events (tirst ,ll1d last <lppc:lr· bed' o
becnu'e {1) they were plankto1ut and widely dis- :~nee~ ol specie~ m time), r,nher th,tn .1 t~·w sekct,•d Amtral
persed: (2) most specics were curythermit and thm "correbcion fimil~"' (Cooper and l mdholm, IYilU). A fi.
not contlned to latitudmal belts; (3) the majotity of Thi~ ltigh-n:solution approach to o,tr.mgr:.ph) u~in~ top of
graptolo1ds were epipelagK. and thu~ may be prc- 130 b'Taprolitc spccit•s, h.1s enahkd thc:~c authors tu nial gl
~crved both 111 dcep-watcr JnJ ~hallow-water fucics, correlate Tremadoc-Llandcilo ~l·gucnce\ on J glob.ll Silun.u
\UbJeCt to preservability: and (4) the 'trattt-'Taphic ba'i'>. with grc.H prcci~ion. hom tl
rangc of nun} 'PeCies ts ~hort. l'hc mam problems Though \Ome graptohtc\ m.ty be very widt- the int
in the strJrigraph1cal use of graptolite\ arc as follows: sprc.u.J, there arc many othl'n. \\ hKh .Ul' g;l'ogrurhl- dtplO!-'T
{1) grapmhtcs are usuallv preserved as compressions. callv re,rnctcd and u\cful fM ,tnctly loa! r ht· c
and thl·re may he problems m pn.·c1sc Identification; corrdauun alouc. Becalt~e of tlm dstlirulry ~c\'~nl Llauvm
(2) the time range of somt: ~'TaptolOJd spec1e~ is palaeontologist~ have tried to dctine faunal Ullltl l--'1 aptolt
rather long, Jnd it is not poss1bk With ~uch specie~ whid1 arc larger than zon\.'~ bm nf mon: or I< 1 f.llma t;
to give the kind of stratibrraph1~,1l preci<>ion that is worldw1dc .:~pplication . Thc~l' ~uccc~,iw graptolite Jitfl·rcn
Bibliography 345
faunas (Fig. lO.ll) arc c.lSil> lllt:nrificd .md 111 any at thi~ time. ev<.·n though they ,m: not nunu:ric.Jily
me epitomiZl' ••1s Uulman ( 11J5~) has shown, the abundant. Lower Caradocian beds contam a
~~ologicallmtol) of the gm up l'lnllti\!TciJlflls-Dicl'lit~I!"'Ptus subbun.t in wh1ch
The foliO\\ mg \cquenn.· nf graptohtc tJuna~ Drcrc~m~i[raptlls is al\o found and "" l11ch contam\ thc
through nme 1s from Bulm.m ( 1958). Tht." v:mous final dichograpnds. !'he OrtiiQ,I?f<1Jlfus-Dicc:llt'l!'•ll'f".l
fJunas and mbfJun.l' hcrl' notl·d arc dctim·d on the subbuna rcplacc\ 1t in the Upper Caradoc and
fir.t appt:arancc or new grapwloid rypc~. Ashgill. In the later A~hgiiJ thi~ subt:tunJ is sonll'-
The Trcmado<.1an i~ chJr.ldcrized by an aniso- what Impoverishl·d a~ 1s the Urthogmptus·
graptid fa una con~t!\ting m.unl) of Rh.r/lcl,opt•ra Climcr(tll!r<IPtlls subfauna of the lowcm1ost S1lun.tn,
•PCC:les and other 1msograptids. <;inn: prov111U.lhty wh1ch unmedJatcly predates the arrival ut the
wa' well marked at tlus time thl·rc arc soml' prob- monobrraptid faunJ .
bm m intcrcontlllental <.orrd.ttion. The tunc This 6una extend' throughout the whole ot the
equivalence of ditfcrent Rlw/Jditwpom species m v.u- Siluri:m (other than m the two lowermosl zones}
IOU\ parts of the wmld is nO\\o belunling rca~nnably and Into the Prag1,111. Though 1t Ius not been
den. evt:n though rhese ~~wdes may be mutually d!vtded into subfaun.1s, different monograprid types.
xcJu,1ve. char.:tcterized mainly by their thecal constructton,
Very few gr:tptolitc~ of Upper TremaJo<.ian age arc found succeSSively and cJn bl' used stratiw.lphi-
1re known , other than one f.luna from the Yukon cally These assemblages allow the SiluriJn to be
r~mmry md anorhcr from NorWJ\ in \\hieh divided mto zonal umt~ of le~~ than a million ye.trs
Btyc.>graptus. Kit~rr.)_\!mpws and po''1bl) a Drdpllt)_\!raptus duration.
ha,·c been dncnbed. But otlwr rhan thj~ doubtful
re,ord there .lrt· no true f.'l"Jpto!OJds k.no\\ n from
rrcmadocian rock~.
The mccecding dichogra ptid fa una of the An:nig
marked by the mconung of the carhe~t dK tograp-
J<h. Tetrctl!mJifll> i~ fimnd nght ,tt the base. and 1\ u~u
Books, treatises and symposia
Uy a\\Oclatcd with didymogr tptKh as wdl ,1~ with
remaming anbo!-,rr.tpnds. The l''-"tensifomt d1dymo- Bulman, O.M.D. (11:14"1-1()47) A lvlom~~rnph ciftlrr Ct!ltldtlr
~pt1d!> of the Arcnig were rt>pl,tccd 111 the Uanvim ( Hcld.uchie) Graplt•lilt'.l from Lime~ tones in Ln,J~}./tlll H11m
~~ turung fl)rJ.. 'PCCH:!> of Drdpllt''<!'"flliiS (though by Ayhflin·. Palaeontot-rraphic al ~oc1ety Monograph (i)
the LlanVlm the mccenling diplograpuJ tauna 1945. 1-42 (ii) 1946. 43-51-' (ii1) lq-17. 5'l-7H.
became eswbh\heJ), and the genus continued mto the (Derailed morphology and OIHo~l·ny, profusely lllm-
Lamdoc bdorc lwcoming l'Xtmct. ProVlllCI.tlity is rrarcd)
weU marh·d 111 thc Lower Ordovici.m, and the CrtlWlhc:r, P.R. ( llJH I) T1te 1-'irrr .'ltwcti/Tl" <?/ Cr.rptttlllc
~ndemic Paciftl gem·r.1 Orlt'c~t:mptm. Cardrt~fclp/11.1 and
Paidcrm. Spcc1al Papl' rs in Palaeontology No. 26.
Ellc\, {; ,L. ,md WooJ, E.M.R. (19.22) -t :\lom~,grapfl .y·
&,\,.uptrts have been tound strangr:tphicalh u~duJ m
Brirrsfl Grapltl/rtn (w1d1 synopnc ~upplement hy I.
bed\ of eqmvah:m age in wc,tl'm North Amerila and Strachan, 1971) l'abeontogro~plu<-11 'iocicty, London .
.~um-alia. (Descnpaon~ and marigraplucal u,~. of all Unmh
A few bbenal b'Taptolnich .m: found ar the very spenc~ then known: swndard work)
or
·np the Arclllg, but the fl'.tl flowering of the bis- P.llrm·r. D. and R..i<k,Jrd\, ~.B. (ed\) (1491) Grapt<•lrtc~:
n.U genera I\ from the Lbmm1 unnl well mto the Wnt111_11 i11 tire RciCkL Ft,ssils illrutr.ltt"cl I . Boydell Pn·~~ .
Sllun3n. The diplograptid fa una <op.ms thc pt·nod Wondbridgc. I H:! pp. (ProfcliSIOn.Jl, though popuiJr
rom the Llanvm1 to the lowcnm.m S1lun.m. before .u;cuunt, 138 photO\ - uncqulvm.11ly rccomnu:ndeJ)
rhe mconling of the monogra ptid fa una. The Rtckards, R.J~.• ).1ck~on, R,£ ..111d Hughcs, C. P. (cds)
hplograptid tJuuJ i' dividl·d 111to tour ~ubt;lllnas. ( l'J74) Craptalitc St11dics ;, Hom111r t•( O.M.B. Hulma11 .
Spccr,1l Papers 111 P.11.1comolugy No. IJ. (20 on~:.'ln.ll
1bc Clyptt>t:r.rprus-:lmplc.wi!T<IJlfll'i ~ubfauna of rhe
PJPl'l'> and bibuowaphy of Hulm.m·~ works)
lJJO\o1m and Llandctlo tontJill\ nuny tunmg fork
Tcid1c:rt. C. (cd.) ( 1970) 7 rentrsc <'11 Im'(:rrcbmtr Pahwlt.l(<~Y·
l1Jptoloids m ,1ddmon to the genera from wlm·h the l'.lli V. Cmptollflmur (revised). Gcolog.caJ Som·l) ol
~una t,tkes m namc; eVldl'IHly the mam pcnod of America and Umvcrsity ofK3ns.1~ Pn-s~. l...1wreun:, K.1n
.btferemiation of thl· bi.,eri.tl ~'T.tptuloid~ tonJ.. place (Upd,ltc:\ Bulnun \ original trcatJ~l' uf 11>55)
346 Graptolites
Fmney, S.C. (llJH..J) limgrngr.1phy of L)rdovaciJn grJpt<l- 1/

Indiv idual papers and other references
litt's 111 tht' Southent App.!IJdu.ms P,J/,rt'<llllolt~iul/ A
Coutnbutwrr> <'(tire l'mau.. rf)' 4 0~1.· 295. 1o7-76. c
ll.itt"t. l>.E.ll ..uad K1rJ.... ".11 (I '~H4) Autccology of Finncy. ~. and Xu t'hen (I 'J'Jtl) rh, rciJtJonshJp of Rtcl
Stlun.m ~-.rr.lptlllold,, 111 < •'•'J'f<•IH< 'lwdr,·s ;,, Hmrom ,~( OrdO\'lCIJil waptohtc pmVIlll'iJh\111 to palaeo~t'Oh'T2-
~
() .\f.H Hu/m.m 'lpo \l.tl PlpCr'\ m Pai.H·omology No. ph). in Pa/m•tJZt•h Hl/m·t':l/n~r<lJdly a/Id Biv~r''II"'P">' (ed.~ 1t
3:!, A, 1Jcm1< Pr,·"· I ondon. pp. 1.:! l-3Q. W.S. McKl'rTO\~ Jnd (. .K Scote,e) GcolOt-'1cal R1d
Bt'ri). W U N .md W aiJc. P. (I 'l<JO) G r.lptolitc bmgeog- Soncry Mcmotr No. 12. pp 1- U'i h<
raph~ . 1mpllt IIW!l\ tor p.ti.H·ogco!!".1PhY .md plbco- Fortey, R.A .lnd Ucll A (I 11X7) BrJnchm~ ~comet!) and Rad
occ.moj.,rraphr. 111 p.,(,u···~··•· 1'.1/,rr,•I!C'I>gr.rplry .md B~t> function ol multJr,lmou' gr.!ptoloJ<k P,rft:,,l.,,,f.l):y 13. (I
!~<'OI!r.rphy (cd\ 'W \ Ml Kern>\\ Jllli C.lt. Scote~c). l- 19. (.
Gcolu!-{lcal Sou<:t\ Mc1mm No 1.:!. pp. I-US. Fum·y. R.A and Cooper. R A ( 19K(•) A phylugeneo, R1d
Bern. W B N., Wtld,·, I' . .rnd Qu1nbr-llunt, M.S. damfication of the gr.lptnloilh. P.rlacorrttt/t~~y 29. E\
(I <JH7\ The oc <'.1111< mHa-\ultida< oxvg<:n minimum 631-54. Ur
zone •• h.1hlt.ll ltlT J.,"f,apwhtn? Bllflt-till d the GeoJ/ogir,,( Kirk, N. (J ')(,')) Some thought\ .~n the etolngy, mode nf 2!!
,<.,ont't} ,,f /Jt'lllllolrk 35. I o.>. 14. life, ;md evolutiOn of the Gr.1ptohthm.1. Pn,rmiitt~; ~{ R•tk
Bulnun. 0 M . B ( I'J')K) 1 h<· \l'l)Ut:nrc of gr.1ptolite fau- tire Cn•lor:iml Svut·ty <'{ Lmdv11 1659. 27 3 92. Gr
11<1'>. Pa/,rnmt!llo~I!Y 1, I 'i'l Tl (Str.otiwaphiral me) Kirk, N. (1972) Mmc dtoup;hts on tbt• .natomobiltry oi h,•
BubnJn, O.M. U. (I %J) On Glyptt•'(r,rpllls den tat us and rhe g:r~prolite~. Qu.mal)' J,,,,,tf •!/till' C:col~f.!ic••l Sudl'ty (<'<
\Oml' .1lhed gcnc·r~ flrJioll'oliii<>/••!!Y 6. M•'i-H'I. <!f L>t1don 128, 127 JJ. (A dt fli:rcnt intl'rpwtJtlon frum So
Hulm.m, O .M B ( I'J(, I) Lnv.er P.ll.te<W<1ir plankton. that of Bulrn.:au) Ri~h
Quart,·rly )<11m1111 ,,f tht• ( .c,•ltll!lf•tl So<tcty •?f u•ndon 119, Ko~zlowsk1, R. (I 'NH) ll"~ l(l':lptolnlu:' tt quclqucs nnu- Hr
~I) 1-1 H. (T IK· ~r.lp!Ohtt' hHlroenn'l' 1s the pre~ervahle ve.lux groupe\ d'.uumaux du rrcm.uloL de Ll Pologne R1~h
pbnkt<lll 111 11\ nnw) PrJ/ncollt<.ll.l!!ill Polomu1 3, 1 :::L'\5. (CIJ\\IC, profusely ll'\
Bull. f:..l ( 1'19h) In arh~ .ltlllll\ 11f th.'IOJJJl Jnd abnonn.u iliU\(f.ltcd \lUdy ot' thrct•-dtllll'llMOnaJ dendr01ds ·md Sh
wowth 'tru~turc' 111 .1 <,,·urt"h Sdun.m dendro1d faun.l. other group~) Rtgh
Plllac,•t~t••l••~y 39 .:! I '1-411 K.mzlow,b, R (I 1171) Early dt•vdopmcnt stJ~~ and the p;ll
Cooper, RA mJ hmn. R.A. (I'JH2) The OrdoviciJJJ mode of lil'i: o( h'1'Jptolrtt''· -!eta P.t/at'on/,,ft,I/IC<I P<~l•'"'•'~~ Jl
~rapwhtt'' o( Spattl•cr~-t~·rr Hu//<'1111 <'{tlrt Bntulr .\Juil'lllll 16. JU-U. H.. 1,;h·
f!.{ .\'tllur.J/llllfllf) f( ;,.,,ft':\y) 36 I"' 7-JII2 (l'roxim.U end Mnchdl, C £::. (I <JH7) Evoluuon Jnd ph)lo~mctlc dl)~l- pl.l
~m1crurc~} ficatJon of the DaplugrJptJlC.l P.1l.rn>Hil•ft'XY JU , till
Cooper, R .A .. funev. R A .md Lmdholm K. ( 1991) 35.3-406. Rtl.(b:
lautu imJI .ami dq,th zntutum of early Ordov1c1an Obut. A.M. (1974) Ne\\ ~lpmhtt."> from the M1ddlt rhl·
!,'1'.tpwtitt''· LetI"'"' 24. I'I' I-~ I H Clmbn.m of tht• "•hcn.ln Pl.ulonn. m l.r.rptolttt StttJ•r (1,,,,
Cooper, R .A. and l1ndholm, K (1')1)11) A preciH" world- m Hm10ur 4 0 ..\1.8 Bulma11 (cd\ R.O. R1rbrd,, 11 E Skt·\ it
wid<: torrdatron of c 1rh 01duvician ~,.rraptolitc Jack~on and C.P llu ~hc,), Spen.1l Papc" 111 rim
sequt•ncc,, C.r,>lt>~h·o/ ,\ hr.~''- "'' 127. 4'J7-525. Palaeontology No. 13, At"adenuc Press. London, pp prn
Crowrha, P .md Rtl!....lrd\. R U (I'J77) Ct,rtical bJn- IJ-1.3. (Oldc\L known W·•ptolite~) Bttf
dag<''i .and till' ~;:raptuht<' /Void Lt'll/t~~~C£1 cl PrJiac- Rick.lrds, H..B. ( 1974) A new monogr,tptid genu~ and the Hu,
''llllll~!iiCo1 lt , 11 lh . ongins of the 111.11r1 rnonogr.lpttd gcnl:r.t, LU Craptoltt• Ac.1
l>1Liy, I' N. ( I'J'JJ) C.cph,rf,llltmr.l .!!Yrljlta/it,>idcs sp. nov .. ,1 Studrcs ;, 1 lnmltll •!f O.i\1.B. Bulmmr (t•tb R.ll. \pjdd
proh.thk t•xt,mt !-'~'·•prnltte. }<11111111/ •!f Lovl~(!)' 229. 69-78. R.ickards. D.E. .Jacl~on and C.P. llughcs), Spew!
Dunmtn, P.N ..1nd t;;t•unlkov, N K. (1 11113) A new rhab- Papers in Pabcuntolot,ry No. I J. Academic Prc<l,
doplcund ht•ma hunbtt' from the Mtddle Cambri:ln of London, pp 141-7 (Monogr.1ptid~ 111ay h.avc origt-
Siht•n.l. fJa/,u·<>lll<li•ti!V 36, ~H \ -9(, nated from dithynal popul.ition)
Erdtm.1nn, B-P (I <)H2) l'.tl.~t· obm~eOl!TJph~ and cnv•- H...ickard'>. R.B. (I CJ75) I'Jl.\t'ot:lology of the
rouniCilt\ of pl.m!...tlt dlrt\'Oill'JnJd f..'1'3ptohte~ during Graptohthrn.!, .111 cxunn cl,a\\ of the Phylum
the clrlu:\t ( lrdt1\ 1\1.111. 111 1111· ( ormlm.m-(>rdtwidan ll em t~horda t..l. Dwh,~iwl Rn•icrt•.> SO, 347436.
R,m,,J,rf)' .'it'< t"''", r·,,,qf Dr•tnbut11•m ""'' Ct~rrdmimu Rick.lrdo;, R.U ( 1977) Pmcm~ tlf .:volutmn m the P"·•r-
(t-d~ M .C. B.•~~,·tt .md W lk.m) , 1\lanonal Mu,eum of colm.-s. Ill Pattcm.1 C)f b•t•/ulit•ll (lod A . llallam), El~e\1t'r,
Wale,, CarJitl pp 'I 2X London,pp 333-'iH
fmnt'y, S t (1<~7 11) Mod,· ol hit- of pl.mkturuc ~l'fJ.pto R.tck.ud,, R.IJ. (19H8) Anaduom,uc. heraldJc, and
I.Jtt's flotanun \liUdun 111 t...)rdovtu.ln Dr((·/1.~1/l<lPIIIS >p. echorc cvoluoon. ne,...· p.lttcm' rt•vnkd by t'''rinct,
Pnlrllf,wf~) 'i , H-'1 pbnktomc hcnuchord.1te\, tll I:.\tlllai<•ll ""'' -:,n·w~lm
Bibliography 347
till FtiSSrl Rt'<tll.l (nl. G.l' l ar\\tll•d). sy~renllri(\ hte\ trmn Non\',1\ . P,,/,lct>/1/t>ft'l/)' 6. I::! 1- J I . (Bilhl'C.ll'
AI~UU.t£11111 ~pcc1.1l Volullll' .34. Cbrcudun Prc". Ill .IIU\0~.1ptids)
l hJi1rd. pp 211- ~~~- 'iudburv, M ( I'J;'i~) Tnangulate mono~,:r.1p11d~ ti orn llw
1\r,l...ml,, R B and Hwr. J.E. (1'170) I he ,·.nh..:'r mono- ,\/''"''1/'''1'''" .l!rt'f!tlnt/) zone: (Lowc::r Ll<tud<•vcry) ol' t h••
.,rr.lpttd [>,,,,,.,·drn~s 4 tlr•· ( ,,.,,h,~rc,rl ·'''"C''}' <'/ Lllldcm Rhetdnl Gor~c (Cardigamlmc). /'lrilc>>c'J'/m,lf
1063. I I 'i-liJ (MonogJ·Jptid nrigu1') Twr 5tllfic11H 4 the Royal Socit'l)' H 24 1, lX5 -::;:;s,
II..Jcl...ml\, R.ll .111d '>t.ut, R.U (I !JH4) l'>!~r.rptw, it'> dassi- (rvnhtrJO JMry lmc.tges)
li<-.llt<HJ, ,·vulunnn 111d zmnd ..•1/drt'fllll/•' 8. lOl-l L Undcr.vnod, ( . ( 1993) The position nf gr.lpLnlires wnlun
lllclo.~rd'-. R. J.j, Cmwdll'r, P. R . .111d ( lupm.m, A.J. the Lmvt.:r P.II,Jeo.wac plauktic ct:mystent /.,·r/r.uct 26.
{I 'I~:!) L ltr,l,tntl rur.1l ~tudJt'' '•I gr.lptnln ' a fl'\'H~\\. l~C) 2112
C:,·,llt\~1111/ \J.ri/,/WII' 119, 35;i-](l, Un.dt'r\\'uod,l (llJ<J'l) lntcr~tlpt• wcbbmg 111 rhc "tluii.Jil
1\Jcl..ird,, R .B .. Hutr. J £ .111d !km·. W . II.N. (1977). gnptohtc:: Cyrtt>l/r•lf'/1/.\ P.J/tlt't>llloh•I/Y 38. 61 ') 2h
Evolunon of thl ~1lun.1n .111d nn·t~lli.Ul l ;raptuloitk Urb.mt:k, A. (l•JSH) Monograpt1d:1e from nr.Htc boulders
Hu/1,1iu O/ tht Hnt~>lr ,\/usmm •I '\clllll.llllr,wry (Gc,•lol!)') in Pol.md. HtfttCt>llt.•l~~gtn l'olcllri<'•l 9. 1-105. (rine \lrtJC-
2R, 1- 1.:!0 rur.JI ckt.ul, m.my tllustratiom)
R•cl...irJ,, R ,l3 .• Rigb). " Jnd llmk J.IL (1990) Urbanek, A ( 19(•0) An attempt Jt brologt(,ll llltl:rpn:t.t-
c.;raptoln1d bmgcogr.1ph)' n:u:nl pro~~n'''· t~Jlur..: Uon of cvnlutionary changc:s m gr·.lptolttt· n~lonw,.
hnpl''• 111 p,,/,Jl'cl;:m,- PulacO,I/t't~l/1<1]'/t )' ol/11/ U''\1/L't'(!f<IJlfry llrta Ptllcrt•••llltllcwia p,,lonica 5, 127-234. (I kvdops thc-
(l'lb W.S. M, Kerrow .mJ L..R Swt<'~l·). Geologic:~! ury ol rht:mil.ll growth rq~tdJtion in ,.,.rapwlitcs)
~cKI<'tY M••nm1r No 12, pp I VI- If, Ut b.uH:k. A. and Bengt:son, S. (l'JHI•) 1{/r,,f>dc>IIII>IIS, .1
ll.1ghy, 'I ( ll)lJ J) h·ed111g 'lratq~tl' 111 ~raptoluids. M1ddlc <..unbnan rhabdoplcurid ht>nm hnrd ltl'. L·1/r,,;,,
P.r/,ll'tlllfc•lc•l/}' 34 , 7Q7 l-\ 14. 19, 29)- \OH
Ru.:l•\ ..., (I 'I'J1) Populanllll .uuly''' .md nnt•ntauon 'rud- lJrlum.:J... A .and Mu:rzesc::w,ki. P. ( 11J'Jl) The line \tnl<-
to·, nf gnpmlotth lrom the ~1td.ill" t )rdm•tt' i.m UricJ tun: uf J , .unarmd ~'T:Iptolitc. Letlrt~r.t 24. 12'J-\H
\b.tlt·. l_2m·he< Hr/.r;·,culc>/,'X)' 36, :!h7-s.:!. Urbanck, A and Towc. K.M (1117-t) Ulrr.htrunur.JI
Rlj,>by, S ( 11.1'14) H cnlll hor.Luc: ,J...dt·t.tl j.,'T<1Wth, 'hMcd \t\Jdit.'' 011 gr.1prohrc--. I: The:: periderm ~nd ir.. Jeriv.t-
p~lt<·m, in Rlwbtf<lJ'kllrcl .md b"' 1ptoln1d,. /..n/r,rr,r 27, uvn 111 thl· Dcndroidca .mbio/o,I!Y 20. I .tH
R"'hV, ~.•tud Mt!,om, C. (llJ11(>) lklllhlt onj.,'lm of zoo- Urb..ln..:k. A .111d Towc, K.M . (lens) Ulnamuuur.~J
pl.tnl..tcHJ. .111 l'JlVICUilllll'IHJJI\ dt•(t'flllllll'd 11l.tcTOeVOIU- ~tudic' 011 ~raptulites. 2: The pemknn and m denv.l-
IIllll.lry l'llt•n, Ct'c'lt~~}' 24, '12--1. tivc' 111 tht· Graproloidca. Smilftso11imr ( .'mrtnlmtio11s 111
ltt,;by. S ..1ttd Rid...mL,, R.B. (JlJHlJ) NI'\' t•v ttlenn: for l'alcol>iolol/)' 22, 1-24. (Transmi~~iou dt·ttrntl mino-
rh•· lifi.· lubit ,,r gr.lplulotd' tl1Hn phY~Il.d mnJcllmg. scope \tlldlc' ol wcll-preservc:d cuudc:, c ~· Ftg. I0.7)
l'~lu•l•ic•lvt')' 15. 402-U. Wtllt.um. c.; 11 ( 19\.JO) Cornputcr-as,i!>tnl j.,"''lpmhrc srud-
mnbotun, I) (I 'J74) Contml, mtlut'll< inp; the <ompmi- 11:~. m ,\/ur<•(ciiiiJllllt'T.' i11 Pal<~e,mtolo..I!Y· (ed\ I) L Umton
aon .md dhtnbunon of Ord,n·it i.m gr.1ptulirc launal .md I ) ,A. I 11.\rpcr). Pt~lc11'0tlllll0,'./i(c1/ C<•ll/nburwm •?/1/rl'
p. pr0\'111<'< '· m Craptc•litt' Studrt'$ irr I ft'll<'llr <'/ ().,\/.H. I ""''""')' '!{ ();[,, 370 . 4fr55.
8ulu111u (l'tb R .H ftt<brd, , D.l . J.,,k,uu ami C.P. WilltJllb, S.H ..md Rtckard,, R.H. (l 'JH-1) l'ahel1ecologr
.
l\~ llught'\ "p<·ual PJ~'<'P• 111 P.tl.tc.'otttolu!-"\ ;--Jo. 13,
AoJcnltl l'rns, londuu. pp. 5 11 73.
of !,"''.Iptnlmc bl.lck shales. l'alctl'tmlt>lt\1/im/ Ct1111ni>11111111S
•!f tltr ( lllll'l'r>ll)' '!f Oslo 295, I 59-(•(•
'.!1ddnan, N. (l'lh.l) "oiiH' upper Tn·nudnc·t.1n gr.1pto-
he
Ill
Arthropods
(
~
Arthropods han·, as dtt'tr mmt Jl,liiKtivc character- a

ll
t\tJC~ •.1 hard outer coat or exoskeleton and jointed
p
appendages wluch they ll~l for movement •.111d Diversity of arthropod types PHYI
fct•du•~· Tho'e livmg today fall 111to three recogniz- Rth t
.lblt• groupm~>s. clu: Crust.Kl'.l. ('hrimps. crabs .md Arthropod' rake their name from the jmntcd phyl
lob~ters). tht Lhdicer,lta (spldt•t '· uutcs and Korpt- appcndagl's rhat arc a comtant fe.1ture of thl·ir In,
ons), ,ltld the Ururamia (111\Cth .mJ other fonns). Of organi,;mon (Greek: cxp9o<j>c; JOII1t, 7tOOo; = be: hi
tlw 1
tht' cxnnn tonm. rnlobttt:' .ltld curyptcrid' (the foot). Thc\C o;amc append.1ges. which includl' the ~d~·.t
ga.mt l'.ll.lt'O:ZOIC w Her 'c:orp10m) are the best jaw \trucwrcs, have become very dlfll:rctm,lted, and sub\t
known . Arthropod' an· pt•rlup' tht· most successful tht•lr number, ,lrr.mgemcnt and morphology .re (lllJI
.tnd di\'CI'\C of alltm·ertt·br.ltt''· .llld \lllCC rhetr tough often oft rirical importance m t.1xonnm}. \o1.tn) dif• 'lOOS
t•xmkdcton offers conSilkr.1hk potential tor fos- fcn.·nt taxonomic 'chemes have been erected for tht roll {
silintton (c,peoally t!" mi11CI.tl11cd) their geological .~rthropod~. and rhcn: is \till mu< h controversy over w.trd
Tl
record 1s good. whether they arose from ~inglc or from multiplt
PI IY
Arthropods arc scgmt·nted cm•lom.1te metazoans, ancestor>. One view is that of Man ton (I IJ73, 1977' b1val•
,1, .lrl' rill' tnnchd wonm w1th wh1ch they posstbly that the arthropods are polyphylcttt. Sht• ronsid.:rl'd Woo
sh.lrt' .1 common aiKt'str. In annchd~ the only that 'arthropodizarion · occurred at lcJ,t three ttmc' ll'rt.ll
'kdcron 1' <1 hydro,tltlt om· provtdcd by the le,tdmg to three or more dJstuH:t phvla. Arwrdm~ tl ~I tc
codnnHc thud. wh1ch dm•, nm gl\'l' .1 rigid anchor- to M.uuon. therefore, the 'Ph)'lum Arthropoda' 11 the c.
.tgt· filr mu,cJc,. The arthropods. however, becaus~: h.l\11."
an unreal entity consisttng of hctcrogt.nCOll\ de·
PHYLL
of thc1r tirrn cxo,kckton, and ll'io c;mce many also nu.·nts. Other authorities (e.g. 13ri~"' and Forte)
j.,'l'OllJ'
h.IVl' .tn mtenul ~kdeton (en doskeleton), possess '' 191:59, 1992) while recogniztn!;!; the three m,1in li\in~ rwo 1
npd b.t~c for .ttt.tehml·nt nf the uuernal muscles thJt t,>Toups. prefer the concept of a \mall nurnb~r ot opistl
mow the ltmh~; heiKt' rhev h.wc the potential for m.~Jor body plans with a common JrthropudJr. cogctl
rap1d lnnlllH>tion. M.111y, fi1rtlwnnorc, h:wc hard .111crstor. The present dehtc, (m· lied by cladi\tit qultl'
J.lw \tructureo, (man d ibles) wh1ch can grind, crush ana l yM~ .111d the discovery of very e.1rly trusta~tdn' <'ntsta.
or btte. Thm arrhropOtk 111 rt·nm of movement and IS hkcly to continue for :1 long lime yt·t. and so fot tronr <
CIJ
tl:t·din~. havt· an ovt·md111~ 'upcnont\ over th~: convemence Manton's system i' folkmcd here. Tlm
ace•
.111ndid wonm and h.1n• ht'l'll .1hle m inv:1dc many cov~:rs both recent and fosstl arthropods, though the fuss
d1tll-rcnr env1ronmrnt' whtch n·nuin dmed to rh~: taxononuc status of tnlobtte\, wh1th arc the m<»l s
Llttt' r. abund.mt foml arthropods, 1s not yet fully r~...oh-ed n
l1
A
SI
1.1
Classification and general morphology 349
PHYlUM UNIRAMIA (<'.1n1.-Rn ) .J.m, b1t. w1th tht 11p ,,f.l CLASS 2 ARACHNIDA (\11.-Rt c) All lrt· ,·t:;lu-lt·grul,
v.h,,J~limb. 'Trunk l.u J.., biramou~ (t\\o-br.lndlt'd) \Uill<' .m~ nunnc, hu1 11111\1 Ill' tcrreMn~l ltHludt·, ~pldt·r,
lppt'nJag~. tlw luuh, .m uniramou~ h,lfVL'\1111<'11 ,md \Uif)'lllll\ , fu"1! ~paJ..:I\ lrt "".1'11111JII\
ClASS 1 ONYCHOPHORA ({ .un Rn ) Tlws<' h~v., 3 tou11d m Uppl'l ( ,lrhomlt' rot l' '\1,11-b ·anng '' •(11<"11• ,..,,
non-ngtd '<'gna·ntt•d lwd~. \\ Hh lti~OIIIIt'd lq;' tlut eau 1'
.llld mJn~ l''\.rruplt•, ~rt· prt"l rd 111 I.Hr 1 t rtl.tl'\ mtbrr ,
~ mftm.·d wnh hlond tnr rhnhmH \\,1lkmg J.'''' ~hem. whiCh \\'.1~ ex,Jdt>d 1< 1 r,•,m trnm rht b.1rk oft omlcr; and
venrrally d1reucd, \\lrh hhdc-hkc tl'lllllnll daw... rr.1ppcd 'Pltlcl'\ .me! mw, '' l')<llogomd.. (,.;,c spt.ICI'\
Exampl~ '"' lud., /',''I'·"" (R ccc m t<'rn·,mll). 1p/rnoJ hJvc \null hndu.:' with four!'·"~"' ot 'llllll ,,r ~pmdh \\;J.lk-
C.l1llbn.ul nunnl lng lq,-s Thcv .lrl' Jll mJrrnt .rnd c'p< tllll~ tclllllll<~n 111
ClASS 2. MYRIAPOOA (C.lrb.-Rcc.l. MyriJpod, and ccn- modt'nt Amarcttc \\.ttn~ \\ hnt· thnc arc tl\Cr LilO
npt•dc,; cnnrch tt ITC\tn.tl, 11.1\ ,. Jllllltni m.mdlblt'> bitlllg >ptctc>. mdudm~-; gum l<ll'lll>. Tht'} hA\'C bct·n rL1!~rdcd
m me tranwcrw pi.\ne. Suml' f:l.lllt fo"1l rcprcscnwm·c~. J' art uult•pt•mknt 'ubphvlum, bur \,1111un \ ,tuJu:, ,fm"
such~ tht• ttrrc,rrl.ll Uppn C.ubumiirou, Art/rr,•pftm,l. rh.lt tht•y .m• mur<' hkt•l) to h.1\'l' dl'nvnl fmm m cJrh
ClASS 3. HEXAPODA (De\, Rn.). ln,etr,. ~cn.ll .md t<:r- gruup of man ne .u ll hrlllh
,..,mal. Tht rno>t thHrw ,111d nulllt'rtlll\ of ,1ll rerrc>tnal

:mirnal ~p<XIl''· M,Uithhlt·, p1IIIIIIIVt"ly rull alld gtind. but Alrhuugh tbt· tJX<HIOIIII C' \t.tW' u 1' 11 ilob1tt.:' .md \lltllhr l11tlll\
m some lal<'~ \tron!\ sc:u1m!Jry tl .lll'\'l'f\t' bmng i~ is yer uncert~lll, tht')' h.l\'t' hn n t·nn'ldt'l cd ·I' .t (H'"'hlt
possible. phylum (,1\ in tht' rhml t•dil i1111 ut tlu, lt ~t) hur 11 prt·,,·tll
P~YLUM CRUSTACEA (l .1111 Rn ) DnnliJJ.mtl} n1;mne 1110\t .ltrtho nru:' would pttlt·• 10 ll'!;·"d thnn ,1\ .1 t 1.1~'• .1
rthropod; with nuny ICI\SI I l t'l'rt'Wilt.ltl\'c'\. A very Y.lrit'd Khemt' provmoru lly •.md wtt h Mllllt' lllltnt.nnry, .1dnptcd
rlwlum with lughly tlttkrmti,Hed ltmb 11111rphnlngy. The hert·
l1li .1dvanced .Jquath 1 nl\l,ll<'.lllS P·"' fi h1d fi.1rw,1rd~ from C l A')S TRII Olil rA (Cnn.-Pcrm.): MJnuc uthropnd~
'lrhind tnw.1rd, the lll<lllth Jl<lll!!, .1 11ied1.111 lond groove ming knnwn only ·" lth\11~. cht ,.llht't knnwn .Jrthropnd,, wuh
dlr lunb,, orb\ fcedcnt; t UITCIII' f:t'llt'r.Htd by cht hmb~. ln hJrd, rhrel'-lnbed exo,kdt•tt'll' t.lgllllturd mtl'norh- ,1, a
;:lrlnced aqumt crmt.I((,Ul\ the fillld " hlted up from the ceph;llon. With free thonnc "Wlll'll£' and .1 pl.m-hkt rlHt-
'Cbtr.~rum by o,pcciJbzt·d .1ppend.1gc' 1 he b1tmg JJW' nor pyg~d1um. I hey h 1\'C .1 p.m of ,uctnmat' md ,.-nail~
!.md!blcs) ue tim11col tfom gnathobasc~ lntt·mal nrcn- rcpcau:d hmt>, ~llthr \\.1\ Jn\\ll tht• hmh
bJrJtnliU\
lh)[b of the a pp( ndJg.:' \du, l1 flll'l'l 111 th< mcdcJn plam· .mol Comld<·nn~ rhc ,.,.,.~ cl dl\'t.:f\11\ of JMhrupod,. ii\'Ul~ Jnd
;ill or gnnd to..:cthn ·" tlw hmb nHl\'l'' bJt kw.trds :mol for- fos,tl. .mol rn ,,cw of thc1r tull trc.1tmt ut "' Man toll ( l'ITi).
udo. The outer pan ,,f the 1··.: lliJ\' dl\.1pp( u dl\CU>\IOII uf drthropud, her<.' '' lmutl'll to trrlt•bltt'1>, c hdKt·l-
The e.uhnt CntiiJti.'Jm ''ere SUBLLASS .ll<.'S and cru,tJtt·an,,
IYLLOCAR!DA (CJm it<"<.), ,,flllh h.1vc ~ Ltrgt.
• rd carapace alm<ht H>vcnng thl' body (l' g. j l•n~ .md
OoJwml IH)!H-1!1'1'1, Rt•lk. lW,:!). Till w, .1hund.1111 in Features of arthropod organization
:tUJn fJ<.'le' tht• <.',IThll I'JIJcotulL, m.l\ luvc been Jncc'-
111
to du: morc .cdv 111n:d ,hrunp-hk<' or llllhtrr-ltkc tom1s. The cxoskdetal c u ticle h.l'> uminuhtcdl) bct•n nnt
c.1rhc,t of whct h .uc lilt'' I Ikvon1.1n 111 ~gl.' .111d which
of the pnnClp.ll f~lCtol"t 111 thl· ~uut'" of the lrthro-
b~comc cxtrcmcl) 1111f'l1rt uH \lntl then
"''llUM CHELICERATA (< .•lm-Rn.) A l.1r~e .mhropod
pod,. lr gtvC\ ,, phy~1C.11 .md dJl'rrlll'.ll b.1mcr
101!p 111 wh1ch .111 rcp rcsuHJii\'l'\ h \\'<" tiH· body d1v1dl.'d m to between the Jmm.ll .md lt\ ~.·nvtronmcnt. )l't .1llnw'
.1 degree of osmorH .1nd lt'lnp~.·r.lturc rq,rul.mnn.
~- o pms rhe prosoma (tmcd ht·ld .md thorax) au d the
~l 't'isthosoma (.1bdt'm c:n). I ht· .JolW' c.m pnmmvcly bite Furthermo re, it supplil•o, gond protcnton ,Jg.umt
n l(tthc r Ill th ~ tr.IIIWt'NC pl.lllt\ IIStll g J h1t111~ m\1\'l'mCnt predators and .1 tirm ha~c for tht· .ltt.tt hnll'nt nf tht·
k· ltCWllike the ~ccomi.Hy trJnSVCI'\t' bumg movement ofrhc inremal muscle~ th.H m ovt· the lirnbs. lt Jlso pw-
lli.ltlCC~m. Tht· prc:SCII l'C of .I p.llr or chelicerae {p111t'Cl'\) Ill
vide~ a sa ri~f.'l cto ry loc .llion fi1r variom kindo, of ~cmt
~t of rhe muurb " t h.11 Jdcn,Lk
ClASS 1. MEROSTOMATA (C.tni.-Rt•r.): A<.JIIJUl' rhehccr- orgam. being linked to th~.· nn\ntl\ syo,rcm through
•tl..,, with rwu 1111(W!i.lllt 'ubd.l\\n \H'll r<:prncntcd as fine tubular canals in the ltttirk .llld 'tr.1tq.,'lt..tll ~
fus)il~: positioned for envtrollllll'llt.JI mon1tonng.
SUBCLASS 1. XIPHOSURA (t .1111 Rt t.) : In~ hale, the Although lht• .tlh.tnt.tgc~ of h;lvmg . 1 t utJl k .lrt•
modem h<li'\C\hm· u Jb I m111b11 md 1t' fn,\11 represent~ clear. there art• .\I\O probkm' wl11t h tht· mlubn.mt
ovr-. (Ordn X1phthund.1) .md th< t ~mbnJn Order of a hard outer ta\lllg h.t, w t Olltl'nd \\'Jth. Of thew.
Agla,p1d~
arriculauun of tht. Juint,, rt•,pir.ltion .md brrowth ,Ill'
SUBCLASS 2 EURYPTERIDA (M <)rJ -Penn.): A gmup of
l.ugt fre\h\\ .llt'r .md nunrw ".Ht·r '' ''11'""" ('innlar 111
perhaps tht· mmt Jnttc.
apJ't'Jr.tnce m rcrrt·,rrul '"'11'11111\, hut uot very do\l'ly All arthropod' h.wl' o,c~mcntni hodrt·'· Tht.•
rcilted) exoskeleton ot most modem one' tnmJ~h of lunl
350 Arthropods
scleri tes, e.tt h mu,tlly comi,ting of a do[',al tergite These m me le\ nuy be .ttt.tcltt•d to llltl'rnal knobs orga
and a ventral stcrnite, n1.1king a nng for each ~cg (a pod e m es). often llmncd by \llllpk Ulflllding of b.JSC
ment. The\e Ill I\ IC.mu .1 rigid cyhndc:r or, altenu- rhe exo.,k.ekwn. or to an cndml...deton. The tnU\· In v
twdy, rhe cxo,k.cktal rint-," nuy be .1ble to move des oper.lte .ltcnrdlltg to tht· noruul JlltJgont n bod)
agamst one .mother by lllt',tm of Ultt'm.tl mmclcs ~y~tem. \O rhat when ont• \t't ol musdcs romracl\ t brgc
runnmg from tllll' 'cgmcm Ln tht: nt'.Xt. The JOint move the ltmh m .1 p.trtltul.tr dtrt·ction. the O~'PO Ht pre<;<
bctwt.>en cat h 'C!-.'11ll'nt.tl ring j, protected by flexi- \t't wtll \lllHtlr.tneou,Jv rrl.!x (.1, wtth the bKt:p~ -and naltl
ble. unmmer.th7cd tlltldc .macht:J to the juncnon tncep' mmde' ot rht· hunt.tn .mn). \\1ht·n th d1an
between 'c~nl"nl\ . ~tnul.trly. t•ach 'leg' (Fig. ILl ) i' oppmmg \Ct t ontr.1c t,. rhe limb 1110\'C:' thr other pods
a ch.1u1 ofhard tvltndcr.; ll.llllll'ltcd by 'hort link~ of WJ.}. rcrrc
~oft .md llextbk m.nen.d .111d p<mered by internal . Re,pir.ltlon 111 .1qu.Htc .trrhrnpod~ normaUy uk~ are l
mmclc,. place through gills. r ht'\l' .lrt• ll\ll.tlly I.Jmcllttc dtrcc
rcc;pi
~lnJ.II
(a)
111 CO
Cr
probl
en cas
ha<. tc
and I.
(5) Arachnida
ts kn
Mthn
HO-<)(
moul
CUttcl
(6) Pycnogomda
1s fon
tlC, !>(
the lo
from
cuta111
(2) Tnlobita (3) Marrella
the ol
(4) Merostomata
but ta
make~
off rhc
1tsclf c
lines <
was tl·
IICSS :lr
(9) Myriapoda
(8) Insecta hody t
water
(b) nu] is
cuttc!e
(7) Crustacea when
rhm, ;:~
m.ty t:a
Dur
both tc
Ecdysi•
Figure 11.1 (o) Diversity of arthropod types with representative examples of major groups, (b) limb joint and internal musculaut o rgarut
in on arthropod leg ((b) Redrawn from Wigglesworth, Principles of Insect Phys,o/ogy, 1965 I ecdys1s
Trilobite 351
org:ms with a vcr, thm nmde. extendtng from the many fossil Jrthropod<., .md by arrangtng the l.1st
b.l\~~ of the append.tgt•s mto the mrroundtng water. shells 1n J ~ize series it ts possible ro eluctttttc the
In very 'm <Ill Jrt hropod~ dttTustOn ovl'r rhc whole various translortnJtium rhar the foss1l .trthropod
hndy surtacc stdhtcs, .md there are nu ~lis, but in went through from the larval ro the J.dult stage. In
l.tr~a ones there Jrl' ~'111s whKh can somcrune~ be trilobttes e~pecially thi, ha' proved to be." nf the
prc:-t~rvcd fos\tl. "'omenme., gtll~ lrt' located m spe- grl'atcst mtcrest.
cialized uucrnal t IIJlnbef\ Modttied gtll, in tntenu1
dumben. are aho loum.l 111 ~ome tt'rrl·srn.11 :lrthro-
.r puJs, \Utb as tht· lung books of .1r.1chmds. Many
terrcstri;l) arthropods breathe by tracheae. which
1re brauchcd, sp1rally thickened tubes bringing air
d1rect trom the outside to the tissues. ~uc h rr.1cheal Trilobites are the carlic~t of .tll known arthropods.
fh~.·Jr first representative-. .trc found 111 ro~.oh of
r~p1rauon is posSible only m .mimals of rebnvdy
mull dimenstom, \\ h1l h is pcrhap' .1 t nucal t:Ktor Lower Cambrian age ju-.t above tht e.trlit'\t non-
mconrrollmg the: uppl'T stzl' limit of IIN~cts. trllobtte (Tonunotian) Cambri.m fauna 1 he lJ,t
d1ed out m the late Penman. Throughout rhctr 350
Growth is pl'rlup' the mmt diHirult phy"ological
problem for arthropods, smce the rtt,.'ld ex~)skcleton millton ye.u:'!. of geolog1tal lustury, they pn:,crvcd ,l
~ncJ~ing tht:m ca1umr eniJrgc once it is fimned. !t
rem,trkablc constancy of f!.mn, though with m.my
ht~ to be shed or moulted at iuterv.lls while .1 new
v.m.1t10m. All trilobites were marine. Well ovl'r
~d lar~er t:xoskclctun tonm. T1m nwulung proct:~s
1500 genera are known. JnJ there arc \ewral thou-
,,tnd spcnes of whit h many, especial!) thme of
:s known as ecdysis It 1\ a ltmiting 'Y'tcm in
1rthropod~ and not .111 cnnrd) pertcct om•, for ~omc
Cambn,m and Ordov1nan rock>. have b'TC.lt 'rratt-
'H-lJ011·u of arthropod mortality UlCllT'\ during
graplm.tl value.
moulting. Before tht· exoskeleton ts ca-.t, a new
111ldt·. somewhat l,tql,et tlun the existing h.m.i shell,
'fonned below tlw o ld one. At th1, \t.tgl' 1t IS elas- General morphology
,, ,oft and wnnkled Wlule thi, ., bt•mg formed,
dte lower pan of the old cuncle ts partulh di"olved Ccrt.1tn ch::tracreno;ttc' Jtsnnguish mloh1tcs from
~om below by l'Orrostve fluid puutl'd out from other :m:hropods. In all mlobtte~ thl he.td
.ut.mcou' glands lwlow the new cuticle. just before (cephalon) is a smgle plate. made up of several
the old exoskeleton 1~ casr, the JJumal ~top\ feeding fu~cd segments. There .m.· mually sense org.1m on
nut t.1h·~ up murh nxygcn .md water As it swells it the hl·Jd, and there arc r1lso cert.tin lim·s of weak-
mA~, spasmod1r movements of tht' bud} to shake ne~~. J...nown as cephalic sutures. wluch look ltke
tl/1" the h,trd shdl, JnJ titully it i~ able to w1thdraw cracf..., on the surtan· but .1pparently taohtated ecdy-
t1df l'omplctcly. Tlw old ~kcleton mar h.wc spenal ,t~. The body (thorax) comt\ts of :t numbtr ol tho-
ne' of weak m·" w lm h tanlit:uc 1ts \phttm~ this racic segntents hmgcd to one another .md
'J~ th~ (J.Se in tnlohltt'\, where the hnl's of weak- allowing some capacity tor cnrollmc:nt of the body.
Ill"' arc known J\ sutures. The tlna1 ~welltng of the The tail (pygidium), ,1ho ~cgrnented, tS fmed tnto a
llody to its full ~in• t.lkt·, place through the upt.tke of smglc plate like the hc.1d. These three primary dlvi-
~ter after the u-.tmg process is complete fhe ani- siom of the body arc set at right angles to the
JIJI i~ now soft-,hdk·d .md cannot move much; unique. three-lobed, longttudinal d1vt'\IOII from
uuck hardening .md tunher secre(lon t.1ke place whirh the class take' 1ts n:tme.
~h(n thi" stage " o-. cr. The mirial cuttcle 1s paper- The hmbs (or append.1gc\) whtch were at~lt'htd to
~~~. and the an~Ul~lllOn of the fi11l cuude tluckness the lower (ventral) mrt:1ce .trc rarely pre.,ervcd When
n~y take ~omt: time. they arc found they ,m: "een to h:we J 'urpmmg
During moultmg the animal is very vulnerable, structural unifomlity within an individu.1l mlohltt.·.
oth to predators.mu m the pos~ibihry of reanng. Except for the flexible uniramous antennae they .lrt•
f dysts is, furthennorc, a wasteful process ,tS much all two-branched {biramous) • .1nd thc1r stntcntre 1'\
loturo tglnic matt:nal " lo~t each omt:. But because of vtrtually 1denrical all the way down the bod)
cd)~ts we have a guud record of the moult \tagt:S of Thcst: ~pecial feature> art• not <>lJJrcd by other
352 Arthropods
arthropod~.and mlobJte' evidt'IHiy fom1 a group of Postcnorl} . the tanal sum re rums out cr.mwcl)el\' ;,tlJ I
thetr own wht(h apparent!} let'i lW Jc,cendanb. on e1 ther ~ide eo tenninate wdl in from t1f cht• genal ~hgl
angle , \\hich '"the most po\tero-Lltcr.ll pomt ot the IUt'J
cepl1.1lo n. Tlm ~ort of suture 1~ satd to be propar- .md
Aca.ste clowningiae (Fig. 11.2) ian. Tht.:· rcb'lon t1tcral to the glabcll.l, though .Lxi.l
within the ~uturc, is the fix:igena ; out\1lk tt I\ thr
Th(• Silurian Ac,hle dtHI'III11,1!"1c, wl11ch • ~ common in librigena . In many trilobites the .mtenor br.u1th ot
the t.lrbon.trc fiKics of thl· bortkr~ of Wales and the fauJ I suture cuts acros~ the .mtert.:l- l ,ltl·r;~l hmdrr,
En~land, ,how' mmt of the ch.1rat tcmtJC te.Hure~ of so thJt there arc two distim t 'uture' rJthcr th.1n a
an ',1d\ .lilt cd' tnlobitc. Tht· bod) h,1, the 'ha pc of.t single t.:onrinuom one. In such c,I\C\, wlm h mduJc:-
laterall) f!Jtrened clllp~e. The hc.1d (ccphalon) is thc IIIJJOnt) of C.1mbn:m ~pccies, thl Cl pl1.1lon mJy
\Oilll'\\ lut larger than the t.1il (pygtdtum), .md the d ...tmtcgratc bcto re burial into duet• cumpontnts
central part of the body (thor.rx) h,l\ I I .mtculared the two hbngenac a11d the c..:ntnl c ranidium,
~t'!(l11l'nt\. wlulh 1\ the glabella plus tixigcna~c. Tlm dl\mtq!ra-
Thl' u:phalon 1s quite \trongl> v<tulted with a tiou doe~ not usually happen 111 Act~Ht' bctJU\e the
pentagonal to semicircular 'hapc and tnnwcrc;e pm- ~uture 1~ contmuous.
terior edges. In the centre of Lhc t eph.1lon is a raised Tht• lowe r surface of the cephalon shows tl1Jt thr
cemral hump (the glabeUa). hounded at the ~ides by a.tlll'ro-latcraJ borderc; are continucJ wntrally '" a
divt·rging axial furrows .md rl'Khtng to the ante- narrow tbnge (doublure), who'e 11111er edgt 11
rior border. Tht· glabdb ,, nakntl·d by three short conccntnt with the edge of the cephJlon .u1d
and more or les~ lTa.tl.,Vef\e p.nf\ of furrow~ (tht• ducctly below the anrero-later al border furrow lt
glabellar furrows) and " timed oiT lt the b.1ck of dol'S nor exrend, howe\er, all the WJ} to the o~:"nr
the u.pho~lun b} an ardll'd occipita l ring. nal nng. being phased our along the po~terior bor·
~tretc hm~ out ~idt:\\ ay~ trom the ocnp•tal ring are der. Two pairs of pronounceJ knob, {th~
the posterior border furrows , \\ hH.. h dehmit a apodemes) project vemrally from thl' postenor two (a)
thtn ~tnp of the ct·phalon .1t tht po~tl·nor edge; thi~ glabellar furrows. These seem to be assou.ued With
i~ the posterior border. Yet .mother indentation the attachment and articulation of the ventr.1l
(the late ral border furrow) runs parallel with the appendages. Identical apodeme~ Jre prt:<~ent on the
semil ircular lateral bordt:r, .md in tlw. tnlobite it thor<~CIC segm ent.~. and evidently there were leg-
joms the na1Tower anterior border furrow round under the head as well as under the thorlx, though
the from of the glabella. ~u c h appe ndages arc not prc~en:ed m . l crlSft'.
Placed lateral!} to the gl.lbeUa .m: the eyes, which A large plate with a central 'bhster' 1s artached to
here are large and cresccntll. The lcme' .m! bome the rear edge of the a.t1terior cephJhc Joublure. Tllli
on a \ tsu.ll surface and arc .1rr.mgcd 111 a 'Y~tem of lS the hypostome, which lie~ belo\\ the glabdb
hexagonal close packmg. There arc Jbnut 100 leme\ The momh apparently lay directly behmd the hyJXIS.
forrmng a compound eye likl· tho\l" of insects and tome, Jnd rrom it the oesoplugm Tlll tor\\':trd tO thr (c)
crustaceans, but they are unu~ually large Jnd sepa- stomach wh1ch by in d1e space between glabella .md
rate from one another. Tlm 1s characteristic of hypmrome. The gm cvidcncly ran posteriorly from
schizochroal eyes, whtch arc contmed to the par- the ~tomac h :~long the thoracic :~xi~ to end under tlu
ticular suborder (Phacopm.1) tu which A castc pygidium . The shape wd pm1tion of the hyposromr
belong... Above the visual surf.Kc lie' the flat palpe- i~ very variJhle amongst trilobites. and many gen~ a
bral lobe . 'ep;1r.1ted by a cr<.:\Cl'llUL palpebral fur- can be identified on the bam of detached hyr*
row from the snul.l ra1sed palpebral area. The tomes found in the rock, even 1f the rest of th~
Jntenor edge of the eye 1s \'Cl} do'c to tlu: glabella. cxo,kdeton ts mi~~mg. A couple of small \wdho~ (e)
the poqenor one t":lrther trom 1t (the mac ulae) lie wwardc; the rt."Jr of the hypos-
A thm though very dmmct ltncat10n (the facial tome. lt h.1s been ~uggcsced (Lmdmom, I<JOt) tlut
suture) rum between the palpl·bral lobe and the these nn: ventral eye~ such as are possessed by vano111
Vl~unl 'urface; thls suture extend'\ torwanh Jround modern :mhro pods, but their tine structure IS nor-
the front of the glabellJ, bcit1g corHu1uo us with the Figure I
mally indisnnct. and this may not have been sn attitude;,
f.1ci.1l 'urure on the other s1de of the hc:ad. In 1lrasrc there are 1 I thoracic ~eg111..:nts. Thev Jr~ nngexp<
Trilobito 353
1ll tdentKal in t(mn, though the pmterior ones are articulating furrow), m front of whtch projects a
>light!} ~omaller. In l'.lt'h then· j, ;~n .uched axial ring ~cmicircubr articulating half-ring which m life fits
identical m fonn to the orupttal nng of the cephalon neatly under rhc aXJal nng 111 &one. The p:ured
:md defined l.m:r,tlly by paired axial furrows. The pleura (smgubr pleuron) proJect honzontally from
:~xul ring j, boumkd .mwnorl) bv a groovt• (the the aXJal ring, thou~h thetr outer extremltl~ are
glabella
glabellar hypostome
furrows
lateral border macula
palpebral
doublure cephalon
lobe eye
fac1al
~
suture
posterior
~:::::~~" g enal
an g le
border
.." thorax
a
occipital
!\
ring apodeme-
:i
axial ring
pygtdium
pleuron
pleural field
0 fa I (b)
h
:tl
eye
.e 3p 2p lp glabellar furrows
;h
;o
.\S
a.
~
l C' (c) articulating facet

ld articulating pleural I
m half ring
I
furrow
I
:1 C
:1 C
·ra
IS-
he
Lg'i (el pleuron (f)
axial ring
)S-
1Jt arterior arch hypostome

Ill\
) t-
Fgure 11 2 Morphology of Acosle downingioe. Silurian, England: (a) dorsal surfoce; (b) ventral surfoce; (c) side view in noturollife
altitude, (d) enrolled specimen, (e) dorsal view of two articulated thoracic segments in on enrolled specimen, with the articulating half.
lTC rong exposed, (f) frontal view showing anterior orch [All x 2 except (e)). For additional terminology see text
354 Arthropods
sharply mmr:d dcw.n l:..tch pkuron ts mdented by an Arastr specnnt·m, like certam other tnlobues. are c
oblique p leural furrow rht'fl' t\ a -.hort doublure often found enrolled wnh the cepha lic and pygtdul Artl
on the nma ~..·dg~..· nt t..•.teh pkuron . l he anterior edge doublure~ in comarr. Th~: outlme of the two ers,
of the dnwnrunwd dl\tal p.trt of r:ach pleuron ts a Rat borders t'> 1dennc.ll. .mc.i the opposmg ~urtaces when Ma1
and mmc.ued articulating facet. When the trilobite 111 cont;lCt .Ire m1rror 1mages of one .mother. ~o bow hydJ
rolled up m a b.tll i(lr pro tee non tht..• paired facers ~lid wa~ ~uch enrollmt.·nt .lch1~:ved? l t is probable thJt tore.
bdO\\ the rt..',lr t'll~t..' nf tht..• prcn~dmg plcur.1. wh1lst rwo lonb'lrudmal ~et~ of pa1r~d mtenul mmcl~ \\1th cons
the <L\.1.11 rcgmn now t..•xp.IIH.kd mtll ,1 hoop. w,\S pro- anragoni~nc action \\>en~ re~p011\tble for holdmg the and
tenc.·c.i b~ th~o: \t.. nt'' of ,lrtlt..Ul.mng half-nng. which body 111 :m extended po~turc or rolhng 1t up. Ont- d)(',
were then cxpmcd. Tht· pkur.a .1re 'eparated by s~r (the fl exors) ran ,11! the w.ry .tlong the body. \Urtj
interplcuraJ furrows. JOining up the apodeme<.. fhere w,h thus J continu- yet t
The pyt~Hlitllll " .1 fl.H pl.ttt' uf fit,ed 'egment> ous p.llr of p.lr;11lcl ltm·~ oflon~tudmal muscle. Thl' .tnd
re,ernblin~ tlllm' of thr: thor.lx. lt h." a ~imilar halt:. o ther sets of 111U\t.lc\ were rhe extensors. l\.1u\ch.· Th t ~
ring ,lt the !runl ,111d " .trtltlll.ttl.'d With the rear tho- scat> on the vcntr.1l ~urf.1u: ,how th.ll they joined
racic segment\ 111 the ,,um• \\',I Y .1\ thc· thoracic the under;ide of each arllcul.tting hal!:.ring to tlw
segment\ .1re ,tlllinhd Thl' pygidul ~xi' h.t~ a ~cries lower surface of the preced111g .txi3l ring. When thl
of fi1rrows eqtuvalen l to ,trtirubting furrow~. flexors cnntr.tetcd the extensors sim ultaneous!~
betOining 111ore dmd> \p.Ked .111d faintt•r toward' relaxed, in the normal p.1ttem of anragonhtiL
the rear. fhe btcr;ll pJm of tht. > pygidium (pleural mmcu lature found 111 tnveru:br.ues .md Vl'rtehrate)
fi elds) .trc ~tulpted bv t-..vu kmd' of lndt..•nt.ltion: one alike. Contr.lt·tion of the fkxor mll\cles would <;()() ...
~crie~ equ 1valent tn tht• r:dgt'~ of the thoraCic ~hortcn the disr.mce betwl'en the Jpodeme,, ~md
segment~ (ulll·rplt·ur.ll furrow\) the other to the since the plcurJl ed~es all formed 'r:nal par.Ukl
thoracic pleur.1l ti.mtl\V\. Ofto:n the l.uter are more hinge~. the thora\. ;md pygJdlUm LOulJ only mo\e
~trongly pronounced in the tnlob1te py~idJUm. The downwards; ti1U~ dw cat! wa~ 'wung uno tlw po,l-
pyg~dtal doublure ha~ about the s.1me width as the tion under the !wad, the kg-. presumabl) having
cephalon. been lifted out of the WJY tir,t. Contr.1ct1on 1)f the
~mce the rnlob1t~· t.. utidt: or ·~hell material' i~ extemor mmcle:- would bnng the bud) b.tck into
fa1rly thm, each furrmv (indencauon) on the dorsal th~: extended pmttton . llent..c the \\hole body of
~urface is nurh·d .1~ .l ndgc on the ventr.ll 'ltde .md Acastt' ~~ finely adapted bllth tor perlcct \phcrotJ.ll
(cl D
v1ce \ er..1. 1 he t Xl cpnon " the tine granulation enrollm~nc and at tlve IJfe m .1 lum:tional. mobile
on the outer \llrt~IC 1.' nt 4 rt~,tt whtch IS not reflected out~trerc hed amtuc.ie.
ventrallv. The Sill.\ I! ~'Tanult:' are rt·mams of se me
org.ms, probably rhe \ltl'~ of \maJI hair; ~u~ceptible
to v1br.mom 111 tht· \.\.\ltl'f.
(d) T•
4rcl.lll' prob,1bly ~pl.'nr mo~t of tt~ umc near the sea Detailed morphology of trilobites
floor 111 .m mmtretrhcd ,mm•dt·. The reconstruction
given here 1' ha,t..·d upon several assumpnons; for Trilobita arc ch aractcri1ed on th~: o ne hand by .1
example, only when the ccphalon is in rhe orienta- confining cvolmionnry comervari~m .md on the
tion illu\tt'<Hed, with tht• ;lntenor border raised to other by a remarkable pla,ticiry w1thm the linub
form an anterior a r c h , \\ill the vmlal tidd of the dictated by the ddlm•d p;lttl'nl of o rganization. The
eye he hori7nnr.tl. llowevc:r. it 'ecm$ to be a reason- overall range m ft)rm is wdl ~h<.)w n in the photo-
able rccommu: 11011, \in a the .1x1~ and the rips of the graphic acl.lsc-s of Whittin ~on ( 1992), Lc:VJ-Setn
anterior pleurat· .1re p.tr,lllcl; the: trilobtte could easily ( 1993) and Lebrun ( I {}lJS). The morphology of th~
rc~t or a.l\vl upt)n tht• \C,l floor m \lKh ;m orienta- earliest and latest trilobite' doe~ not depart verv rad-
tion. Thr: uphftt..•d from of the cephakm (anterior ically from that of Ac.we, and much of the ob~cn~d (el Co
arch) bduw \\ h1ch the hypo,tom~ prOJected, and range in morpholotn can bl' related dm:ctl; to th~
the posterior arch ftmned hy the progressively biological funcnon\ of Jt~ vannu' components. Figure 1
mg outer
\honcned po,tt·nnl thoranr \Cf::.'lllent\ and tail H ence it is appropnatc to coll\idcr the mmphologt- hvely; (c)
upl.Jfted from the \l'J floor, would al!O\\ the fret· cal range in fun~:nonal tt·nm beforr: procecdin!{ 'nth border or
pa\sage of water bel m\ .•lt'r ating the feather. gills. the cla~o,iticanon. From Stor
Trilobite 355
Cuticle (Fig. 11 .3) outer !aver with large calcne (or po,'>tbly l."alcium
-\rthropodan cunde~ nom1ally nm\tSl ot\cvcrallay- pho~phate) cry,tallitcs having thetr c .tws nonnal to
m, and the cunclcs of trilobite' arc· no exception. the surface, :tnd a much tlucker mncr or pnncipal
\bny arthropod cuncle~ are constnu.:t~.:tl of 1.hitin (a layer of rntcrocry~tallinc calcite. 1 he uuwr layer is
hydror.trbon .1llicd to cellulose) ami sometime~ rein- laminated, the tndtvidual lamin,te bcmg concen-
forced by mmeraL In trilobites, howc•vcr, the cuticle trated in three zone~; the outer and umer zone~ have
con~tst' largely of low-magneswu <:akm· (Wilmot closely spaced l:tminae, whereas in the n·ntral zone
.md F.tllick. 19H9) arranged ut m in oLry~t.tlLne nee- they arc much more widely sp.tced. F-rom the bJO-
dles oncnr.ttcd nonnal or ncar-nonn.tl to the outer mechaniral point of view (Wilmot, llJlJO) tnlobtte
\Urfacc and \Ct m an organtc b.m~ \\hose nature has cuuclc~ .trc bc't rebrardcd as ceramic,, havmg a low
y~t to he tktemuned (Daltn~\ater. llJ73b; Teigler percentage of organic matter and behavtng tn .1 ltn-
md To we, 1975). Chicin has not yet been detected. e.trl} da~nc manner. Whereas the thm pn~matic
Tlm ntllt k conststs of two l.tyer.;. a rcbttvcly thin outer layer h,ul good comprcs~we strc.·ngth, tt had
,
~
f
1 (c) Dome Ill
~
(d)
V
Tubercle
\j 7 pseUdotubercles
~/ , (g)
$
J
IC
l~
IC
)-
.u
:\C
l-
.d (e) Composite tuberc:ule
ItS.
figure 11 3 Tnlobite cuhde: (o) section through cuticle of Phocops rano (Dev.); (b) section through terrace ridges of P rono show·
'9 outer cuhculor layer absent over the 'scarp slope' ond the 1wo kinds of pore conols opening on 'scarp' and 'dip' slopes, respec·
!;1- ~.ely; (c) section through dome; (d) section through tubercle; (e) section lhrouah composite tubercle; (f) section through anterior
th border and port ofcepholon of P. rono; (g) culiculor sculpturing of P. rona (x1.5). [(c), (d), (f) Redrawn from Miller. 1976; (e) redrawn
lrom Stormer, 1980.]
356 Arthropods
I.Jnlited tu li.H.'klllg. The undt>rlying

ft''>l\t.llll:l' merely domes w1th 1 span· bdow: othel" Jll' uuc
nucrocry~ralhne prinup.tl l.lycr, hm\ ever, was tubercles e nd o~mg an mtem.ll space connected by
htghly rc~htant lO l rack fomlJtlon. Accordingly pore canah ro the outer and umer surfaces of the
tnlobit..: cunck' couiJ rc,ht romprc~'1ve and tensile cundes. Some kmd~ of true tubercle~ '\eem to haw
forLes. Further n .•,i,t.mce to brcak.lge ts gtven by the been the mes of large numbers of grouped pore
architet rural propcrtit'' nf the t::o..oskekton. for canals or sensory organs of other ti.mcnon. There J.re
strucmrally the cxo,kdewn 1' ctlectJVely a mono- also p seudotubercles wh1th do not h.lVe the dl~
coque 'hell: J strong 'thin \hctr behaving a~ a cretc appearance of ruht:rcle' proper They tend to
'stres~eJ ~km'. Its LOmtrutllLH\ a' a ~erie~ of modified be concenrnted 111 p.uticul,lr area' of thL· exmkde-
domes strengthem the 'hell ..., do doublurcs, cell ton, e\pcn:~lly on the gl.1bdb .md 111 .m:as where the
pol}gons .1m.i tcrr..tte ndge'> The wtide i~ well mp- doublurc ~~ IJkel)- to come in contact with the scJ
plied wtth sen sillae . 'mall 'lructurc' interpreted as floor.
of sensory funcnou . Thm many kind' of tubular The mo~t compkx of the~t'. ,1bo known as com-
canals tr.wer;e the tlllll..k, kading from the 1mide posite tubercles. .1rc found 111 ph,Koptds (Ftg.
to the smf.1ce; the m.uonty of these are straight or l1.3c). Th c~c h.1VL' .1 ~t·rie, of tmy ~phcrical cavmcs
hehcJlh, co1led pore canals, oficn with tnunpct- direcTly below rhe ~u rfJce. from wh1ch fine canJh
shaped outer ends .md m the Cambri.an run radi:tlly inw;ards tn .1 dub-shapt•d ce ntr:~l mm of
E/lipsoct'phtllus wtth dm-hl...e ~we!Jtng.. within the fine branchc~ arrangt·<.l in J !:111 (Stonner, I !)HO).
princ1p.1l layer (0.1hngwJter l't al., 1991 ). These are Their function h:~~ beL·n tk•bated but is unresolved
most dmdy pJL kt•d w here the Luticle is highly Tn the ex:unpk illuo,tr:lted hL·re, Pha!'<lJl.l rana (Ftg
curved Tlll·y wert. prob.1bly mainly ~cnsory. carry- J l.3t), differe nt kimh of \tructurt'' - t~·rract· hne'. Fig
ing mull h:~1r; (setae) cxtenully, each connected to dome~. ruberck~ and port• cmJI opc:nmg<i of dtifcr- met
the cenrr.1l ncr.·ou' ~\ \lCm by a nerve rumung up ent size~ - have been 111,1ppL'd. Dllferent p.uts of rhl ing
Rec
the caml B) :lllalogy wnh modem anhropods. most cuticle were \upphcd With thlferent combmatHlJH t•i
of rhe~c would h.l\ t: hl·~ n ~cm1tive to Vlbrauons or semillae .JCting a' envtror1mellt.11 \Cn\Of\ . ,\1ost
chenuol chang.: in the water. cuticular structure' 'eem to hJvc: been 'c:mory.
Somctiml'\ the pore on,tls arc Js~ociatl'd with a though \pec1fit' func..nom tan only ever he pmru-
~~tem of parallel nd~t·~. do'>dy spaced and forming lated for a b'lven organ b}' .1nalogy with semofl! m
a regul:tr 'tingcrpnnt' omJment over all or pan of modem arthropod\; ~IIKl exact analogil'' 'ddom
the mrtacc. Whl·n ex.1mmed. thlS ')'Stem resolves occur, tlm pron·dure lJll hL· h.wtrdom.
it~clf.1' a ~enc~ ol terrace ridges (hg. ll.Jb) gJving Some other ktml, ol tunn1lar 'om:~mcmatlll~
a senally rcpe.m:d chp-.md-Karp topography Vl.,th were not appan:nrly semof)-. \uch .c, the caecae
the \emory porl t an.11~ npl'lllng at the base of the conunon.ly cncountcrc..·J in Cambnan tnlollltt"\. The
Karps (M1ller, 1lJ7S). llumc experiments have cuticle of many Cambn.111 tnlob1tc gencr.1 ,,.,,, than
~hown th,Jt tl11' rould h,wc time ttoned as a current- and rclanvcly flat. am.l 1t 1s fr'-'4lll'ntly unpn~'cJ
monitoring system, ~c1mt1 ve to change 111 water cur- with a serie~ of r;1di,1l and r.1milymg ndgL'<i tonnmg
rent direcrion f t•rr,ltl' ndgc\ have also been an clabor:1tt' ,md symn1cLnLLl pattern over thC'
interprercd. howcvt:r, Js dev1ces tor tac1litating fric- cephalo n and ~omc tilll cs the r~·~t of the body. Tht''lt'
tio n in hurr0\\'111~ tnlobit<.'' (Schnulti.tss. 1981). caecae seem to bt· m oulded lo the lonn of a tubulu
Such tcrr:~ce raJgc~ Jn· nom1.11ly '-·oncentrated on system originally lying bdow; ctther a drn1latllf} t•r
rhc doublurc, but they nuy Jl\o occur on the upper respiracory appararu~ (lell, l97H), or the divc:rttcul.tc
surface of the tnlobttc. fhe ventraJ terraces may of a complex s ub~t·ph.llir ~t·t of digL'Stive mga111
have been U\ed to comolld,lte the walls of a filter (Opik. 1960). The,e ndgL'' h.we bt•cn rderrcd to JS
chamber unJc..·meath till rt.'\tmg trilobite. In certain alimentary prosopoo and wcm to have been ron-
order\ (e.g. rhc.. ProL·tid.l) tt.·rra(e ndges are &e- nccted wah the oe~ophtlh•m (r1g. 1 l 4a,b)
F1gure
qucntl-,. developed, in other\ h.1rdl} .u Jll. Nutnenrs dige~tcd 111 dll' h"'.tt (Otlld prt-sunuh~ de re,
Man)- rnlob1tc..·' h.I\T cuticular tubercles (Fig. be passed d1recrlv to the other pa.m uf the both b) under~
11 Je-w a~ thl·lr dominant ,urfau.· ..culpture. In thin me am of thi~ S}'Stcm; Jltt·rn,lliVd}. Jnd more probJ- vie\V,
stomoc
section thL'\e Jppc.lr to be of vanous kinds bly. they may h;we ti.1nl tJOned ,1s ~ux.iliJ.l) rc,p1!3- behind
(D.llmg\uler. 1973.t,b; Mtlkr. I 976). Some arc rory srrucmres. ll owevcr. it '' very· r.ll'l' to find jfied Fn
Trilobite 357
alimcnt.uy prosopou 111 po~t-C.1mbn.m trilobtte~. m

which the shell t\ g~o:nn.lll) much th~e:kl·r.
Pre~unubly the mrem.1l orbr.tll\, of w htch the proso-
pon JS the extcmal unprc\,1011, .m: ,tllJ there but
located f.1rthe r belO\\ the cuuc.:k and nn longer par-
tially within Jt. Ltkcwi,e, the lranwc"' ocular
ridges that .ue nonnal Ill C.uubnan nilobttes, hnk-
ing the eye wirh tht• glabt.:Ua (amJ uH:idcntally mu-
ally carrying t\\0 pro,opon c<Jt:cac), an· rJiely
present in poq-( ambnan tnlobitcs. The prc~cnce of
such caec.:ae mu:tlly sugge't' that an unknown tnlo-
bite is of Cambnan age, .ltlt.l the 'culptur,tl p:lttcm~
have been found mcfid 111 t.lxonmny.
Cephalon
Mosr trilobite cephnla h:wc the form uf ~emicircuhr
plates with well-ddincd structures, .1~ reprc~cntcJ in
A mste. T here arc some, h owevt·r, m whkh the fur-
rows limiting the ditTcn·nl strunure~ are all but
efi:Ked, ~o thar the v.1riom pam .1fl' very ind1~tinctly
F;gvre 11.4 Cuticular scvlptunng: (a) Redlich1a (L Cam.), ali· defined, e.g. Trinwrm. Part1cul.1r organs the g1Jbcll.1,
mentary prosapon (in block), (b) Cruocephalus (U. Cam.) show·
•ng peculiar cu.(icular sculpturing and ocular ridges (x 6). [(a) t:1cjaJ mturc~. eye~. hypostome .1nd cert.lln ~pec:i.tl
P.edrawn from Opik, 1960 (b) redrawn lrom Shergold, 1971 .) ized character) - merit further artentton.
Glabella
la)
The ~hape, ~i ze and \tmc.:ture oi thl.' glabella arc
widely varl.lblc. Glabcii.Jc lll.l) or m.1y not re.1ch the
anterior border; 111 \Ome l ,1\l'' thl'}" lllJ\ be greatly
~woUen, havt• latcr.ll lohl'\, hl· cnnrt• or mdeutcd
With up to five pairs of gl.•bdl.1r furrows. ,\\ m the
examples shown m F1g. I l.h. The 'tomach prob.1bly
lay below the glabella and .1boVl' tht hypo~tomc,
and its ~1ze, and hence thl• 'l?e of tht• glabella, wa~
probably related to rhl' tnlobm·\ d•t·t. Some sym-
(b)
metrical mdcntat10m o n the gl.•belb of Lert.lin tnlo-
bttC'I (e.g. Cltasnlt>f'S, where they .lrl' arr.mged m J.
1ntesttne V-shaped pattcm, anJ Pltmaps (h g. 11 .5a), 111 which
7
/ rhcy fom1 a p:~jr of ~ub c lrt" ular p.ttrhl•s] haw be~u
said to be the scars of suspen sory ligam ents or
muscles holding tht• ~tomach in pl.tce and ;\llowmg
tt to expand and comract.
hypostome A rcconstm ction by Eldredge ( 1972,\; Ptg. ll.Sb)
has shown how the\l' rmght h.1vc been arranged.
F.gvre I I .5 (a) Frontal view of Phocops rono (Dev.) with cuti· Cephalic sutures (Fig. 11 .6)
Je removed, showing impressions, possiblt muscular, on the The cephalic sutures, wht1.h .an. un1qm· Jl1l011f,"'t
~mg matr1x ( · 1.5); (b) recoostnx:hon of cephalon in lateral
arthropods, mclude the tiu.:ial 'uturc~ and ventral
~ shOwing possible arrangement of internal organs with
:bnoch supported by muscles (block) and mouth opening cephalic s utures which arc somctm1c~ present.
belund hypOstome [(a) Redrown from Eldredge, 1972o; (b) mod· The faCial sutures .lrc of three nu in kind~. propar-
:>.eel lroni Eldredge, 1972a ]
ian (F•g. 1 1 6hj). \\here the pmtl'nor branch pas~e'
358 Arthropods
t
t
11
(I) Encrinurus g
(c) Redlichia (d) Paradoxides
c
:u
tl
P•
cl
in
h<
(e) Niobe (f) Ptychoparta
C\
gr
gn: ~tf:liD
m
pr
(m) Hypodicranotus
tn
(g) Ca/ymene r- (h) Lygdozoon ~

c.
plc
int
rcg
nn1
(
SL'V
~tit I
,1cn
exa
SUtl
Join
Figure I I .6 Dorsal and ventral views of the cephala of various trilobites, showing relationship of hypostome to cephalon, and whi
course of sutures. The maximum number of ventral sutures is shown in (g) (Co!rmene), Where connective sutures (c). hypostomal sutvre tra1
(h) and rostrol suture (r) enclose the rostrol plate (rp). (a) Holmia (l. Cam.); (b) Xystriduro (U. Cam.); (c) Redlichio (l. Cam.); (d) nor
Porodo1Cides (M Cam ). (e) Niobe (L. Ord ); (f) Ptychoporia (U. Cam.); (g) Ca/ymene (Ord.-SiiJ, (h) Lygdozoon (Sil.); (j) Paladin SUtl
(Carb.); (k) Deiphon (Sil.), (m) Encrinurus (Sil.); (n) Hypoclicronotus (Ord.); (o) Phil/ipsinello (Or .) Rostrol plate in black. (Mainly
redrawn from Whittington, 1988a,b, and in Treatise on Invertebrate Paleontology, Pan 0 , 1959 ) rnlo
ro n
ICaJ
193(
Trilobita 359
in fro Ill of I he gnt.al Jngk or 'Pllle; opisthoparian imporr.1nt ventral 'tructural component m early
!Fig. ll.(lb-d,k), 111 which 1t tlll'i the postenor bor- trilobtte~ ~uch a~ the olenclltd" .md that m many later
d~r anrcrior m tht• gcn.tl .mglt·: and marginal (Fig. lines of descent tt was reduced or lo~t by dtHerent
ll.llla-t .g). whnt' it nlll\ .lk)ng the edge and I'; not evolutionary pathways. In EnmllllniS (Ftg. 11.61) the
''htbk on thl.' dur-.11 'url3n•. In one family, the rostra! plate is very small, .1nd the two connectwe
Cdlymcnidat·. the t:acial 'utun: '' gonatoparian '\Untre' arc dose togt•th l'r. whcrC'a' 111 ,\,'fclbt (Fig.
(Fig. 11 .6) and rum din·nly through the gen.U 11.6c) tht· conm·cnvc suturt'\ .lre rt•prc,cmcd by a
angle. o;mgle median suture (m ). In -tca.,tt' .md it' rdauves
The~c main \Utur.ll ~y,tcniS were believed to there is no tr.tcc of .1 nwdt;m \utUre .•1nd the: two lat-
dehne natural ~roup111b.-; within thL trilobites er,ll fktal 'utun~' unite .1round the front of tht•
though out muth of the nineteenth century and the gl~bdla \O th.tt the ltbngenal.' .1rc fuwd.
fmt pan ufthe t\H'ntactlt u:lllury They \\ere origi-
nally mt'J a~ J prim.tf) b,,,,, of d:t~~tticariort, erect- Hypostome
mg orders: the Prop.aria (prupan,ut and gomtoparian The hypostome nuy be l:trgc or ~m:-all, short or long.
genera). the Upt~tltop.trtJ (opt~Lhop.man genera) lt has proved of ~rrc~ll value in cl.ts~iticltlon. Usually
Jnd the l lypupari.J ( a n.Hglll.ll-~uturcd geuer<l). This it lies din:clly below the most convl.'!x p,•rr of the
cb~stficaoon cventu.all~ came undt·r severe criticism glabe lla. The primitiw condition for trilobite
Jnd w.H tln.1lly ab.tnduncd wht·n it was appreciated hypostome~ seems to have been conterminant
that the Prop.ln.a ami I Jypopan.1 were both com- (Fo rtey, 1990). H em.:, the hypo~tomt• is attad1cd to
posed of vt·ry ht·tt·rngl'llt'Um dcmeuts which had no the c~.:ph :Jlic doublure ..md m ... hape closely corre-
clo~e aurunJ rclauomhtp,. ~tubbldidJ (l':J36), tor sponds with the outline of the front of the glabella.
mswnce, ha-. pc,tntt:d nut rh.n 11 ypoparia could not In the natant cotH.htmn the hypo,tome was not
be a n.ltlll a! grouptng, \nacc: marpnal sum res had attached tO rhc ccph.tliL Joublu re. l mtcJd tt la) on
>1denrh been lkriwd 111dc:pc:mkntly m several the ventral membrane. behind the cephahc dou-
group' of rnlobiw~. the 'llture' beconung more blure and separated from ll by a br:IP it may have
margmal .1s the t'Yt'' wt:re lo~t Tlu·re may be no been anchored by !J~ament~ u~ually the anterior
pnmmwly cycle'' rnlnhitc,, .md all of the blind margin is the same \hapt: as the front of the glabella_
mlobuc' (wHh the pm,ible exception of the A tlurd rondJUon tS an impendent hypo,tome.
Cambnan ,lgtlO\tld') wett· denvc:d from sighted Here the hypo~tome '' Jttached to thl" doublure but
.mn:,ror. tts shape bear<> htde corrc,pondence to that of the
Rt•cenr cl,l\\lttcJtlom h,l\'l' med the whole com- glabella.
plex of mlubttl' lll.lr,\l'tl'f' to dtvJdc the trilobites The natant condi[JOJl t \ denved and ts usually
mto n.mtral groupmg,, l''Pl'C tally thme of the a.XJal conservative. Some tnlobtte~ ongmatmg &om a
rrgton, .mc.l nor nwrely .1 'mgk character, however natanr ~rock become ~ccondanly contenmnant or
unpon.lllt it m.ty .appt·.lr to he impendcnt. lbrcly the hypostomc: proJects back-
On the ventral \tdt• of thl· cepl1.1lon there may be ward~ bcyond rhe ccphJion. Hypostome~ mually
lt'Vl'r.d nther \llttlrt·~. e'pcn.1lly where the fucial have .1 pair of L:ucrnl w m gs wluch may turn up
~uturcs tros~ over thl• .llltl'rtor hnrder and continue inside rhc ccph~l on to n•st dose agaimt its mner
mo~s the doublurt•. Ca/yiiH'III' (Ftg;. I l.ng) is an wall. so that the h ypo~tomc wa' tlrn1ly braced
example 'howtng till.' m.txtmlml number of possible against the dors.1l cxo~kelcton. A linmcd degree of
suture\. I l en· thl' t:llt.tl \Uturc:' .trt· continued and movement may h,wc becn po~~ible with natant
JOIIl wtth the l.ltt•rJI connective sutures (c) with hyposromc~ (Whirrington, 19HHa,b, 19H9). In ~ome
wlurh they m.1y bt hunH,Ingou\. An l'longated ros- cases the hypmtome is ngidly tlxC'd, as 1s particularly
tra! plate (rp) ., 1\tll.\tl·d by the..,t• and by the ante- evident in Er1rn1111nH (F1g. 11 61), m wluch the bt-
rior rosfral suture (rs) and pmtenor hypostomal eral win~ an: pronounced though slender and deh-
Juture (h ). Tlm ro~rral pl.tte '" not prc,ent in .all carely fom1cd, and which ha' " large, three-lobed
rnlobtte\ and ib .tb,entc or n:c.lucuou ha, b'lven ri~e impendcnt hypmtonll' proJCCtmg tC.1rward\ below a
to much phylo~t'IH.'tll' speutlauon. Thc: morpholog- pronounced anterior .1rch lt " connected to the
Jlal sent"> hen: illmtrJtl'd (bawd un Stubblefield. doublurc by a V-shaped hypo\tomal suntrc. and is
1436) ~ugge.,ts that I! wa<; onginallv prc\ent as an pre~umablv immovable. Ltl..cwt~e m the case of
360 Arthropods
\'ic,bt (Fig. ll.nc) .md m rdatn:es no movcmem was each nude of p.mJlel mic rovilli . AltcnloltC plate~
pO\\Ible. ~ince tht• ,um:nor border of the hypostome h.1vc their blocks of tubules amnged Jt n~;ht .1ngles
t~ ~trungly cun·nl .md 1s let tnto tht" re.1r of the tlou- to one another. These tubules are the ~ire of the
blure. Thu' th~.· t )rdm•nan Hyp,,dicrciiiMIIS (Fig. photoreceptive ptgmt·nts whose che11ucal .tltt·r.ltlon
1 I 6m) h.t~ .1 \.lT\ long lwpo,tome extentltng almost by light triggers an electncal dtscharge 111 the
w the pygtdium .tnd prolonged m to ,1 p.m of long o mmatidia! nerves. The nerve unpulse\ art'
bt..J~.·, with a mcdun 'P·lC~-' between. processed m .1 complex o ptic ganglio n deep below
the ommaudia, and \Oilll' kmd of integrJted llllJ~e ~~
Eyes (Figs 11 7, 11.8) produced from th1.• mo<,;uc effect of hght conung
rhl• CYl'\ of triJoblll'\ Jll' the
lllml JllCil'llt VISUJ] sys- down individual and ~l'p,u·are omm,ltldl.l. How
ll'lll known, and tntkcd they .ue the e~trhest of all 'good' the arthropod eye is m contr.lst With Its ver-
wdl-devdoped ''-'11\ory W\tems 1 hetr evolmion tebrate counterpart j, very hard to a'se". for thl." rwo
on be tollm\nl through \Ome 350 nulhon years of .tre d1ficrenr kind' of t•yes performmg b.Nc,tlly dlf-
geulo~l."al time (CI.Irl..,on, 1'J7S. I '179) . TnJobite ti:rent funcriom.
eye\ .1re compound . .md like the l.tta.1l eyes of fhe eyes of triJob1te' nuy also hJvc h.td .. ublcnm
modem cru\tact·am .md imt'C~ dll'y Wl'Tl' composed ommatidia, though tlm may not have bt•t•n the ca~c
of rad1,11ly arranged Vl,u.tl unir' pn1111111g 111 diticrent in all. Little evidence of internal srntcture surv1vcs
directiOns and olkn t•mompamng ,1 wtde-angled The lenses .tlone :m~ prc\ervcd becau~e they. like rh~
\ 1\U,Il field. cutJcle, were constructed of calcite.
fhough cumpouml t:} <:' are typKal of arthro- Mmt trilobite eyes ;tre holoch roal (hg. 11 7a).
pod\. they han• l'\ t'h t•d from \cp.tr.tte bt'b'llllllngs 111 havmg many round or polygonallemeo; who,e cdgl...
•1 numba of .1rthwpod ,rocks; Jnd though the eyeo; .m~ ,1\1 m contact .1110 whtCh .ue covered hv :1 'mgle
ol tnlobltcs .trt: ,IJl,tlogOLI'- to rhmt· of modern corneal membrane. the equivalent of tht• outrr
.mhropods, tht•y .1rc not ncce~sanly homologous. In cutit ul:!r layer w1th wh1ch it i~ laterally connnuous
mmt modt:rn .uthwpmh rhe vi\ual unit' are the l lolochroal eyes an.• the mo~t ancient kind nf eye in
o mmatidia (l:ig. ll.Ht), c1ch bt•mg .1 cylinder of tnlobnes; they are fi.)ttnd 111 the earlic't CamhnJn
t•d), wtth rlu: phutt1\t'll\lll\'l' dt·ment\ (rhabdom) tnlob1te~ and per..1~t until the final exnncnon uf
lm..1tcd deep wtthm tl. E~ch omm,ltldtum io; capped tnlobJtes in the late Penm.m.
In .1 corneal lens. undt:rlymg "h~t.h " 1 \ttb~1diary The oldest well-preserved eye' are kncm n m
d10pmc app.uatu .. (the crystalline con e). The lem l ower C:~mbnan eodm:1d' from China (Zh.mg and
.1nd cone togetht-r ll1tll~ lighr on lht: rhabdom. The C larkson, 1990). The\e are lughly orb':llliLed, though
rlubdom com1st> nf ,, cvhnder of ~l.lt \...ed plate'>, .1 somt·whJt irregular lt'ns-pJcking sy~tem suggc:'t' Figuro
sociao
Huygo
(e)
ofligl
mode
lenses
ponor
oo cl.
Clerks
tlut
grow
were
of ph
.11'1.' p
whoh
sutur
Figure 11.7 (a) Holochroal eye of Paralejurus brongniarti IDev.). Bohemia, Dvorce·Prokop Limestone (x 8); (b) sch•zochrool ey11 ol dt'.lth
Pliocops rano (Dev I Silica Shale. Ohio ( 18) (c) Acernaspis (Eslcospis} sufferto (Sil.), Pen~ond Hills, near Edinburgh, Scotland Joncc
(x 2.5)
throu1
Trilobita 361
(b) (c) (d ) (e)
(f)
lens
crystalli"e core
rhabdom
(h )
(g)
caesura
fgvre 11 8 Tnlobite 11)'!!$: (a) Acoste downingioe, lateral and dorsol views of schizochrool eye (x 7.5); (b), (c) lenses of Dolrnomtino
JOC10f1s (Ord.) and Crozonospis struvei with (shoded) intralensor bowl, confonmng to ideal correcting lenses of Descartes and
1\Jygens, respect•vely- models with a small refroctive index difference between bowl and upper unit Focus light sharply, (d) passage
oll~hl rays thr009h prismatic lenses of Asophus (l. Ord.). (e) Sphoerophrholmus (U Cam.), on olenid; (f) internal organization of a
IIIOdern arthropod eye, (g) Onnothops eye (l Ord ) preserved os on internal mould, and showing irregular disposition of identicol
~ses (x 7.5), (h) Y1suol field of Acoste downingioe eye with individuollens·axis bearings plotted on a lambert equal area net; (j)
panoramic visual field of Bojoscutellum, only beOrings of peripheral lenses plotted. ((a) Redrawn from Clorl<son, 1966; (b). (c) based
on Clarkson and levi·Set1i, 1975, (f) redrawn from Snodgrass in Treatise an Invertebrate Paleonto/ogy, Part 0; (g) redrawn from
dorkson, 1971, (h) redrawn from Clarkson, 1966; (j) redrawn from Clorkson, 1975.)
h.tt the 't·par.ltc dcvelopment;~l progr:unme~ for thereafter welded to the librigcna, the only ~urure
trO\\th of the eye Jnd cmpl.Keml·nt of rhc lense~ then rumung along the top of the eye .l\ the palpe-
1erc not tull) coordm.ued and 'ometimc' gor our bral suture, wh1ch forms part of the dof\JJ f.1c1Jl
olpha>e. In gcnaal. the eye~ of(' unbn:m rnlobne~ c;uture. The post-Cambnan radiation of holochroal
ut poorly kno\\ n, became tn the ma.~onry the trilobite eyes was substantial. The VJmal \urface ma>
hole vi\tt,ll surface i~ cnnrded h\ an ocular be 'mall or large, even hypcrtrophtcd m ~ome
1uture. 'l> th,H the VJ\U.ll 'urtjn· dropped out Jfi:er groups, and encompasses a variable anguttr r.mge.
J(ath or dunng moulting. Th1~ 'Y'rl·m w.1s aban- Lenses in Cambrian trilobites. where found, arc
dnned m most po~t-C.1mbnan ~:,rroup~. probably .111 thm and biconvex (Fig. 11.8e), but in some of
ihrough paedomnrpho~is, <lnd thl· vi,ttal smf:1ce was the Ordov1cian groups, especially thmc with thick
362 Arthropods
runrle~ (Ftg. 11 Xdl. rht· 'leme~· h.t\'c the form llf complex (CIJrk,on, JlJ7'l: M.ilJer Jnd Cl.lrk,uu,
long pnsm~ with .1 tlJtttsb omer ~urtJu: .111d .1 hetnJ- llJHO). Electron microhrraphy ~how~ th.H the bm\1 I.
sphenc;ll inner end: 'urh lenws h,tVl', however, a comim of dense cnlcite whereas the uppt·t unit 1~ (.
\imii.U' focal length to tlm~e rh.H .ne btcnnvcx. Each htult of calcite sheet' .1rranged rad1Jll} wund the 1
km i~ .1 stn~k l ry\t,tl of ta.lcite. wll!d! bt•ing highl~ JXI\ of the lens. E.1ch of these ~heet' in turn t\.111\1\l\ 11
birl'lnngent doe:' nm .H ti~r sight \ecm to be: a vc:ry of .1 p.1lt\,1de of nlnte tibrc~ parallel with tht• ~XI\ 111 .11
,uitJhlt· nutl'ri.ll for .1 dinpmc app.u~lllt' Howc\'er. the lower part hut tummg outwards tJil\\'l,C, near n
.lllthl' lcmes .trc .trr.tn~t·d with their c ,\Xl'\ nonnal to the: nutl'r ~urtJCl' of tht• km. Tlus lllJ) h,l\ e nmu-
thl' \'J\ual ,urfllc. \O hght tr:l\'dhng p.lrJild wtth the mi,.ed the dTcct~ of hircfnngcnce. A n·ntr.tl tort•
.u.i' j, not hro!...l·n 111to rv·.'O t.~ys bm lOtlllllllt'S unal- ~mmJ in P/lawps nuy, ,1\ CJmpbdJ (1'175,t, llJ77) In
tcrnl to the rh.1hdnn1 ( >bhquc r.1y~ lll.IY have been ntllt'd, .tl~o have some <orrccring fi.mct1011. Altt't (S
\l rt·~:ncd out b) p•gtm·nt cyhnde~ lwln\\ tht' lens, .ts et:Jy'•' the lcm I' rc:-tom1cd in stage~. tht: bowl u ..·
in rnndcm .trthrnpod l')'l''· to whiCh holochroal eyes bcmg .1dded lao;t nf all. frj
\l'em to beJr ,1 t:Jir \frt~~:rur.tl .md fum non,1l rc:sem- ~dliLochroallcmt•\, 111 'Pite of bemg nudt• of c:JI-
biJnte Ull', wcrt corrected .tg.1imr .\sngmattsm and pn"ihl)
On the: other lt,11ai, schizochroal eye\ (Fi~ .tl'o aga1mt undu~. bm.~fringenr effects from nbhqut' IPJ
11.7b, lt.Ra-t,g), "h1C'h .ue confined w o;;uborder r<~ys. Uut why wen: they ~o large and of ~uch lngh no
Ph.ll:opin;.t (Ord. l)l'\' ), h.we nn known modt'm opucal 4uality? lt is gcncr:~lly believed (<. '.1mpbdl, \.\ .1
l\Hintcrp.trb; the" .1n .1 umquc '1~nal 'Y~rcm unlike 1'>75..1, Ford)'Cl.' and Cwnin, 1989) th.lt r.tthcr than t.'XJ
.111) thing ehc m tht• .uum.tl kmgdom (llorv tth et ,1/ • .111 ommJtidium I) mg hck1w each !cm. rlwn· \\,1\ ~
1997). In sch11m hnuJ l')'L''· \Uch ,1, rht)\t' of. -Jtt~stt', n:l.mvcly short 'ocdl,1r ~Jp,ule', tloon·d by a tbt
the le: me' arc l.u gt' 111.! <,cp.tr.ttcd from l'.H'h other by I.Jyt•r at narro'" lclb SLhizochroal eye\ .ue rhm
.lll mter,ritial n l.lte11.1l (sclera) ot the ,,tme \tmcturc nlJNdered as Jn J\~embl.Jgl' of '>Imp le eye' nf htgh
•l) rhe rest of the: tundc. E.Kh lcm h.t~ Its own resl)lvmg power .1nJ with overlapping V1Stlal field, .
cornc:~l covering, whit h plunge' throu~h the sclera Such eyes (Stockton and Cowen, 197(,) m.1y have
and inremalh pt nb.1hh tl'mHn,ttt'' m .1 ~ublens:tr been capable of U\11\~ .ld}Kent lemes for ~rcrc~·~
t·omc:JI \tnu::turl', ''I lit h I\ nrch prt''t'rvcd ~coplc vt~ ton through 360°. They would have b~en
The structure 11f the: len,e\ 1' IIHl\t 1nrere\ung. p.1rt1cularly meful in notttmul .mim.lh whose brgc
When·1s holodut,,ll l'Yl'' havt lctht'' r.1ngmg m lem~·., enabled them to '>cc in the dm1 light. Thcs~
dl.liiH."tcr from Jll l\ 1 2!10 111m, though most arc lc~s lughly ~peciahzed VISUal organs arc qtntt· diiTcrtnt
th.111 lOO mm, 'tlul'mhto.lllcnse' Jre U\t1.tlly much from rhmc of ..1ny modcrn compound eyes, .mJ
brgcr, being 111 thl' 1.111g(.· 120-750 111111. They .m: their design may h:~vc ol1cred unique: potenti;ll tor
mually steeply btconvex .wd of compound 'truc- the :maly~is of colour, torm and depth.
tllrt'. \ \ nhw e.Jt h lt·m j, .1 bowl-llkt• umt. \t'p,lr,Jtl'd !'!ch17ochroal eye' ongmatt•d from holochmJI
from the upp<r p.Ht ot the lcn, h~ ·• \\'.IV) ,urface om'\, probably b~ p.lc:domorphoo,i,, \lllCt' l.trv,ll cy~
\duch h,J, bt•t•n \IH>\\'11 m \'mom gcner.1 to be ,jmi- of holochroal-cyed mh.,bHe' are 111 mJny w:~y' hkc
l.lr in 'hapc to rlw 'mtace of apl.m.Hit corn:ning tmy \Chizochroal eyes. The l'Jrltcsl \l hitorhroJI
leme,, .1s dt'Mgm·d by Huyg(.·m (t ig. lt.Rr) .md genu~ (Onll<lflrups; 1 : 1~. ll.Xg) had a ratlt~:r h.aplm-
Dc,c:~rtcs (hg. 11 Hh). Expetlllll'llt.tl models ard and 1rregubr ~y~tem (Jflcns packing. "' hirh .1row
(CI.Hk,on .md ll'Vt-\t•m. IY75) hJvc 'hm\ n thJt J from the geometrical constraints of pat. king leme' of
,IJ~ht d1tlcrencc in rt·fr.ttti\·e index between rh~.· on- ttknnc.ll ~izc on ,1 lurvmg v1sual 'urf.Kl' (C I.Jrhon
crHco olc1te of tlw upper umt .md the bowl \\Ould I 97~). In beer dt•m·am·e\ of Ormt~th<IJH rcguhr
operatt: togethL't Wtth the correcttn~ 'uriarc to pro- p.lt kmg was aclueved by graduating thl· km ,Jze
dlll e .1 ~l1.1rp .m.hll~lll.1tlt to cm. In .1dJition, the De.,pltt' rhe elegance of the eyes or tnlobitn, •t.:-
howl :tppc.lr; to 111l'rt':J~c tr.m~mi\SI\ 11\ of h~ht to nnJ,ll)' blindncs~ is quite common (Forte} .1nd
the pltororcccptor': tndced the whok ~y,tem i' fully Owl·m. 191.}\Ja) :md ch.1r.Ktensric of m.~ur r,lX,l surh
bm-optimizcJ tor m,\XJJll,\1 efficicm y. even dunng .1~ the Agnosrida a.nd Tnnudctda. In othn trilobite
tit\.' \tJge' after moulnng (Hon·ath .111d Clarkson. ~:-rroup' tht· Ios<; of the eye 1s probabl)' rchttt·d to the
I'N3) adoption of a d.Jrk environment or mfaun.tl lubltJt, Figure
upper
Tlw ll'Jht'' of l'h.l~<lPS .lrt' ultr.l\tntt tur.tll) v~:ry· Progrc\\ive eye redlKUOn has been 'hown in t\\O (Pho~
Trilobit a 363
~uccc~~tve hm·.tgc~ of Upper 1kvoJU. III rroptdo co- Cephalic morphology and fringe of trinucleids
ryphml mloblt c\ 111 'lHllhe m I rame (Fct't .1nd (Fig. 11 . 1Oa-f}
Clark~on, I'IXY) ..md t untcmp mam:u mlv Ill ph.t-
Thl' ceph::~ l ic fC,lturc'> of the Ordovu :w1 Fannly
copith IFci\t, llJl)J 'ourh umdlrl.'ctlnnal cvolurt on.
Tnnude~tlae an.· unique . In tnnucle Jds thl' glahell.1 "
mvol\ln g tht• .tdopttu n nf .111 endohe tllhte h.1b1t, i' usu;tlly highly convex , ha~ ~hallow glabell,tr furrows
JIJ ad.lptin · rl.',pomt• tu con,tJlH .md p~:r'1~te
nr env1- and 1\ bounde d bv broad .1x1.tl fum:'IW\. Later.tl ro
ronmen talmflu entt''· the glabdl.t arc the qu.1dr:u11-,hapcd genal lo bes,
which nuy rang~: from gt·ntl) SW11IIen to grt·.arly
Cephalic fringes (Figs 11 9, 11.1 0) infl:w:d .md an· de\ o1d of eye' .1nd \UtUrt.'\ . In
In F.11111ly Tnnudc 11be (Ord.) .utd FJmdy 1-Ltrpid.tt'
Tretaspis and Rccdolitllll.\ then• .m: \111.111 nodes set on
1\ubnrdcr ll.trpm.J Ord ne\ ) the mtero-I .Ht'nl the ~Cll.:tllobcs. fach Ol thC\l 1\ ,1 dnmc-s h,lped tlnn
l"tph,tht bordt·r IS dt•velo pnl inro ,111 I.'Xlt'llSJV I.' pitted are.1 of thl' tc'lt, though !cm-lik e ~rructures h<IVC
fringe , the 'uture h.1~ bl'conJ c lll.trgmal md tht· eyt'' been reporte d Jt the '>ummit uf the dome (~cornwr,
lrc n:n n:dured nr ab't'llt. Two kmd, of fringt•
19JO). Rarely, on1lar ndgc' lllJY hnk the l'ycs w1th
were lmkpt•n dently eV<.JlVl'd ~~~~d arc d1s~imil;tr in the glabella. In .111 tr:inudc ilb the genal spines arc
Jppcar mcc. In .1 ti.·\\' otlter trilobit e groups the antt•- vt·ry long and tcnnina tc well bdunJ the ~hort body.
rior border nuy be pmeJ. though nevt•r in thl· s.1111e
whtch lu~ only Mx thoraci c ~l'!l;lllcnc~ and a 'hort lli-
w:ty nm Sll t'\.tl'mi vdy; yt•t othn tnlnb1te:. posse" angul.u pv~dium.
expanded ;mtcno r tbnge' though withou t tht· plts. The fi 111gl' j, " bilamd lar 'ltructu re in which the
two lamcll.1r counterpart~ arc 'cp;lrJtt ·d h> a mar-
gtn,ll suture, wlm h become s dorsal only \\here the
gmal sp u1c~ join rht• n:phalo n. The uppl'r Ltmella j,
chu' conngu ou' wtth the genal lobe,, wherea ' the
h1wer has the gcn.1l ~pines ;mache d. A ~erie~ of fuu-
nel-,ha pcd fringe pits m dent both !Jmdl.t e. l.',lrh
dor,,JI pit located directly above an mvene d Vl'lltr,11
roumer p.1rt, so that the whole stna.:turt· j, real!). a
hollow ptll.u. The floor of t'.Jch pir (the re1hng in
the ca~c of tht' Vl'ntral onl's) 1~ fanned bv J termina l
disl or 'nozzle" wirh J minmc central pl'rfor. mnn.
and .tt rhc 'llltllrl.' which rum berwee n then1 the two
counrcrp.~rt termina l d"c' an: JUXtaposed. The 'race
betwee n the upper and lov.·er lamel1.1c opens to the
lower sidl' of the Cl"ph.llon by a pair or seril'\ of \'t'll-
tral perfi.)r.1nom. That.• mmt have been nllnnw m-
cation betwee n the fi·mgc c.1viry .md the body
cavity; probab ly part of the thgemv c or urtubt nrv
sy'>tcm w,,.,
hou~cd therein .
Some of the early trmurk tth have an .1lmmt f1.1t
frmgl" with poorly ordered and irre!o,'lllar fnngc pm.
In later ~tocks. howcvl 'r. thl· fringe pib b<.:( .une
orJcn:d 111t<> a synUJJt'tncal p.mem df ord~:rcd •res,
com:cn tric with 1he .tntero-1.\teral border, mtcrst· n-
ing with r.tdial rows. Some 'mall 1rregula1 ·f-pn,·
d
.1rc found in 'null p.ttches latcr.1l to the genal
h lobes in the genera ·\ f,lrrolitll iiS and Cryplll/111111~. The
e
homolog11.:' of the arcs 111 vanou~ rnnuclc ld genera,
H'
Figure I 1 9 Tretospis ceriodes angelina (Ord.), Norway (x 5) wluch is l'~~cnti.1l m t.vconomy and idcntitlcattOn.
t. 14J!ler lamella of cepholon in dorsal ond fronlol loterol views. can be a\'c~sed With relen::n n to a th1ckencd ventr.11
il'hotogrophs reproduced by courtesy of Or Alon Owen.)
ndgc - tlte girder (rJg. 1 I. I Oc) - conccnLric wnh
364 Arthropods
1• 1 (b) (cl (d) Botriodu foveoletus nt.

ho
ge~
WJ
tht
bd
dis·
tcr
ItS
km
Cryptolithus
COl
rJn
1110
f
The
that
(f) Orometopus
into
(h)
Fun
lug!
~m a
the ,
nor
row
t,1po
tht• 1
marg1nal band
rhro1
~upp
p<:rir
cnrn
Figure 11 10 Cephalic fringes: (a), (b), (c) Cryptolithus (Ord.) in lateral, dorsal and ventral views (x 2); (d), (e) reconstruction of 1he tht• I
biim in Botrotdes foveofatus and B. bronnt, (f) Oromefopus (Tremockxion), a possible trinucleid ancestor (x 4.5), (g) Horpes (Dev.) Ill
dorsal and lateral view (x 1); (h) recomtrvcted section through brim and cheek showing perforations and thickened marginal bond l"Otd.
[(o), (b) Redrawn from WhiHmglon 1n Treohse on Invertebrate Poleontology, Port 0; (c) redrawn from Hughes el ol., 1975, (d)mocli- mal.
~ed from StOrmer, 1930; (~. (g) redrawn from Whittington, os above. J hkc J
body
more
the border and now known to be homologous in all trilobites could use their legs to excavate sh.1llo\\
trinudetd~. Arcs external to the gmlcr are numbered burrows, in which imprints of the fnnge. genal En1
£1, .E2, ... , while incern.1l an:~ arc ll, 12, ... . The spine~ and appendage scratch marks ,tre often vt~ible The,
devdopmcm of pseudogirders concentric with the Evidently trinucleids sat in thctr burrows with the l011111
main ~trdcr in some genera h.1~ l·onft,,ed ta:wnomy head down (Campbell, 197Sb). Some sene~ ol as spl
m the p.1st, bur a~ the homolob'll'\ are now clear the supcrpo~cd burrows made by the one a111mal whtle m.t.y 1
tdenufiLation of species m thi~ mo~r 'trangraphically shifting poSition show thJt dunng -;uch 1110\'etnent thor.H
unportanr group. though rime comuming, no the cephalon always faced m the ~ame general dtrec- doub
longer presents the problems it otKC held (Hughes rt tion. lt was probably rheoracocally onented to the there
nf.• 1975). currem in the only stable pmition; l:tter.1l cum:nu fiN fc
11ow did tnnucleids uve. Jlld wlut was the fringe would have ovcnurncd H. H ence the fnnge may be .b J A.
for? Ventral appendages known m Cryptolitl111s have interpreted as a sensory organ, each ptt haVlng \rn- C,u
very long fiJamentous gills and stout walking legs. sory hairs at its base whtch were rc~ponstVc to tsopyg
Trace tosstls made by trinudetd\ ~how that these changes in current direction :md so enabled the .11ll- lllCII t.
Trilobita 365
nul to keep m head to the ntrrcnt Tht' may not. from domg more than curl up m a half-sphere
however, have been the ~ale ftm(·non. Other sug- because the distaJ free edges of the pleurae then
gemom made mdude rhe pm,1b1ltt) that the fringe came m contact. Sucb arcbmg of the body, bow-
was ~omc kind of filter, but tht\ •~ unl1kcly because ever, w::t~ .ln e<;~ential prerequiSite for moultmg, for
the pronounced anterior arch would let m currents only in th1s position, and ati:er opcnmg of the
below .md becau~e the tiny hole~ 111 the tenninal cephahc suntrcs, could the newly moulted tnlobite
dtscs (or nozzles) ,1re not likely to h.wc :~cted as fil- crawl out ameriorly (Wl1ittington. 14YO). While
ters. Our understanding of the trinudcid fringe and ~uch p.1rti:1J (Oiling wa~ necessary tor t'xuvuuon, the
Its funcnom must proceed, as ut ,,lJ ca~c' of this ~tn1cturcs which allowed th1~ were u\efi.llly pre-
klnd. b} extremely detailed morpholngtcal analysis ad~ptcd for th~.: development of tnte cnrollment.
coupled w1th a strattgraphtcu pef\pccttve of the Some C.tmbnan trilobites were able to rot.! m a
range and development through tunc uf the variou~ ICIO\C sp1ral, while m other. !Tom the latest
modttic.ltlons of the baste ~tnu:tural theme. Cambn:m (Sun. 1983) proper enrollment was fi.11ly
achieved. Many Ordovicta.n and later groups. how-
Horpid Fringe (Fig. 11 . 7Og,h) ewr, developed an elaborate enrollment ~ystem,
The fringe 0f trilobites of Suborder H.npina, wuikt: articul:!ting t:lcccs being particularly well developed
that of tnnuckids, 1s flat and prolonged backwards in Ordcr Ph:~cop1da, to which Arnsll' belongs. In
mto a pa1r 0t curvmg hams rather th,ut ~cnal '>pines. addition, within this order there arc many different
Furchcm10re, the dlstnbution or p1h on Its surface is kind~ of 'tooth-and-mcket' stntctures. \eenungly of
fughly Irregular. and the pits thenbdve~ are r-.tther quite mdependcnt origin. which are found on the
IITUl.l and ue tound on the genal regtom a.~ well a.~ on (cphaltc ;md pygiclial doublure and help to 'lock' the
the &tnge .;;ome genera have a 'mall rnangular ante- rolled-up sphere together. These .uc known as
nor arch. but generally the !Ttn~e 1s lbr all the way coapta tivc struc tures (Ciarbon and Hcnr), 1973;
mund The fnnge 1s btlaminar, wtth two closely jLL-x- Henry, I 9BO; Hanunann. 1983). and though partic-
tJposcd lamii1Jl', and the p1ts, wh1ch tn <oecnon have ularly well developed in the PhacopinJ they arc pre-
th~ fonn llf opposed vertical fimnds, perforate right sent also 111 tnnuclcids, calymenids :md .1s.1ph1ds.
through tt. There is a pronoltnced marginal band, Among phacopids Acaste has no n:.1l rooth-and-
1upphcd Wtth many sensillae. round the external socket structure~. though the ccphalon and pygid-
pcnpht•ry nf the fringe. Htupcs and ~~ allies cmtld ium are the s.1mc shape and the oppo~m~ doublurC\
fnrtlll, haVlng a pronounced Aexurc of the thora.-x in fit together neatly. In the related Uppcr Ordoviciau
the rq,'lon of the fi~t icw thoraor ~egmems (dis- Klottcekill, J ~mglc median projccuon m thc ccphaljc
,o,dJI cnrollment). It nuy h,we been .1 sedentary ani- doublure mterlocks with a corresponding excava-
m..ll. U\ll1g the brim to ~pread the wc1ghc oi the body tion tn thc pyg1d1al doublure. Aft,~!!"'"' (F1~. 11. I I g)
likl' ,1 'now,hoe. The pitS undoubtedly l.Jghrened rhe has :1 senes of lateral sockets for tht· rc,eptton of tht·
hod>. but ,uch an dahoratc stntrt\lre may have had pleural ups of the thoractc segmcnc~ and the 11f't
unre than one function . pygtdial ~egment.
In Pltacops a deep marginal groove (the vincular
Enrollment a nd coaptative structures furrow) tS often present on the ccphalJC doublure.
Th~ enrollmcnt ~ystcm exhibtt~d by l l msfl' 1s very The edge of the pygidium tits imo th1s, and the tho-
.ommou 111 po,t-Carnbrian trilohttes .md 1s known racic ~egment~ come to rest in cxtenstons of the vin-
sphero idal e n.rolhnent. More rardv, trilobites cular furrow diVIded into individual lateral pockets.
1\ roll up by tucking the pygtdtttm .md bsr few Such l::ttcral pockets may still be pre\ent even when
· ll'3ctl 'cgmcms under the ceph:·t lon. th1s being the med1an part of the vincular furrow " ab\cnt. a~
Jouble cnr olhncnt; and in harp1d' .md mnuckids in Eskaspis (f1g. ll.llh). Morphologtcal difference~
·re 1~ discoidal c nrolhne nt m wh1ch muy rhe such as thc~e are of some taxonomlt value.
:N f~\\ thoraetc segment~ bend, the r<.·~r being held Technique\ of enrollmcnt have been fully explored
tn flat lanunar plate. by Bruton ,tnd Haas (1997) in Dcvoman phacopids
Lambnan trilobite,, with the exception of the by senal sect10rung. Other e)\'ample~ of quite differ-
l!lpygom .1gnostids, had poor facilint·~ for enroll- ent mechanisms arc shown by t11cri11ttnH, and in the
ll.'llt. Sume, e .g. the nlenellicb, wcrt· prevenrcd cheintnd Placopt1rit1 another stratigraphically docu-
366 Arthropods
llf I
tl)ll
T
·" b·
thl·i
Cllll'
(b l
.an... .
hall:
tttrl'
gt'lll
tl'RH
(c) Leonaspis tnlol
n..·pc
l'H.'l"l
0111.'
llllll
Tl
c.un
(a) Salanopaltis ottcti
~rou
TU\\'
p.1rrs
the l
p.trl\
(g l M orgatia ht·tw
whtco
C.unl
(hi Eshsp/s \tTliCI
(f l Proat&U
,tn.· u•
F1gure 11.11 (a) Selenope/t,s mocrophtolmo (Ord ), an odontopleurid (>< 1), (b), (c) Leonaspis cleflexa (Sil.), in two ohernohve life .tnd a
attitudes for resting and swimming or browsing; (d) Dudleyaspis (Sil.) cepholon in side view, attitude equivalent la (c); (e) Cybe/oide1 the l
(Ord.) in life attitude, lateral view, showtng the body supported on long macropleural spines on the sixth thoracic segment, (f) Proefu~
In life attitude, lateral view, showing high carriage of the body; (g) Morgatia cephalon from below, showing press studs (vmtvlor ;t\soci
notches) for the reception of thoracic pleurae and with the pygidium in place (x 4); (h) Acemaspis (Eskaspis) cephalic doublure wiill bl Ll rt•
vincular furrow and vmcular notches (x 7 5) [(a) Redrawn from Hommonn and Rabono, 1987; (b), (c) redrawn from Clarkson, 1969, Jlll'll t(
(g) redrawn from Henry and Nion, 1970, (h) redrawn from Clarkson et al. , 1977.]
pmt-<
.In.' \r
rhor.u
mented ~cnes ~Jwws .1 progreso;ivc mcrt•Jsc in the .mgular gap between the .mrcnor margu• of th~ prcvl't
number of dcpre,~Hlll~ on the dorsal ~lllt1Ce of the lephalo n .md the pygidi:1l doublure. In solttc gt'llt'rJ rt•st d
cephahc amenm border. When t•nrolled, the forwardly projet ttng ~ptnc~ or rows ot dcllttdc:, on Pan d o
PH~Idtal spun:~ n:,tt•d in rhc~c like dutchtng fmgers the n:ph.llon pJrttally < ovcr thts open 'P·Kl' but ian c
(Henry and Cbrk,nn. 1975). kJvl 'ymmcmcJI g.lp,, \\ hirh presum.thl) JU<l\\etl proh.tl
~omt· other triltlbitc h'Toups (e g. tht• Pmcrida and 'omc Jq,rrce of water urcubtion 111 dtt' enwllcd tttc.:ntc
Agnosndd) .lpptJr to hJvc bet n wdl .1d,1prcd for ammal Enrollment \tntttttrt'' may "I'll bt• 'l'l') Thl
cnrollment, but only thl' PhacoptdJ and C.1lymenida unpot rant iu das~ttirJtion. lnund
rcJched the summtl ol cnrollmettt .tbilny Super!
Enrollmem ~truLturl'S in some d.tlttt,tmnd trilo- Thorax I I I l.t
bttes have been pmtr.t\Td b\ C.nnphdl ( 1977}. In Mmr tnlobitcs .lre 'polymt·rids', tlut ''· rhcr lme l'llrtOll
the\c rhc pygidtun• dot'' nm fonn ,, nghr se.1l \Vlth 'cvcral thoracic \cgment' In the agnmtllk hO\\- .trt eq
the ccphalon hut proje< h lam ard Llf tt, k·;wing a tn- ever. there art· on I) t\\ o ..md in the endt,nch two SlllltJb
Trilobite 367
1r three thlli,H.H: 'L'!:,'l11L'nts: tht' h the 'miom erid' rton. The ( )donto pleund a arL a \'L'f). spm) group of
onditw n. tnlobtt cs. .md e.Kh plcuro n ternnn atcs in t\vn
The .txi~ tlf the thor.r\ 111 mmt tnlnbit e\ i_~ about
\ptncs, fhc .mtenor pleura l 'Plllt: t\ norma l!) \ ertt-
., broad as thl ocnptt :tl nng. The,L ,txi,ll nngs \\ 1th
L al; the pmteri or one 1~ hori.w nral .md
th<:tr h.tlf-rin!--."1 .uttl .tpodt.•me' .1n: ut t:tirly stallli.trd uwally short.
llowc: vcr, in mmhfi cd odontl)pleurid.~ (.: g. Ca.Jit>-
wrmru cnon throug hout the tnlobJt t·,, though then.:
.-tplwla) the vcrtt(.ll ~rmc:s are VC!) long and. being
~re diffcrl'llCL'~ 111 tltt' length ul the .tpod~.·mcs.
the equipp ed with lll.my sccond .m 'pine' vtrtual l}
balf-nn~.., arc \OlllL't llllt:\ .tb~~.·m..ltld surf1cc
~culp dmt' off a o;ubthor.lt ic 'box' from the t·xrcm al envi-
turc and ~putostty .trt· \',m.tble. Cnt.ti n phacop itl ronntL 'nt: a Ullltjlll ' feature '-diOse timctin n LS qllltc
·ncra from rhe D~.: \ onun ut \outh Amcnc.1 - ,1
unkpo " n.
r~!--'1011 long •~ol.ttc:d 'D th.1t a rn h t.·ndcm tc tJUIJ:I
of
nlobttL '' devdup t.·d from . -in1tt- hh alllL'<otOf\
- Pygidium
rrpeatedly devdop~.·d dol"i.1l ~pme~ of .1 p.1rnc ubr l'ht· pygidi um i~ .1 fused plate whil h in the l(lWlf
crcn t'Onstrut.llun, soml'tt mcs sin~h. snmennu:·~
C:amb nJn t'lcm·ll tds m;l) cO!hbt of JU't one 'eg-
ont: for t.•.tch ~cgment of thL• thorax .md on rhe mcnt, bur 111 lat~.·r genera it co n st~t~ uf;t, rn.my a~ 30
oc~.ipital ring antl p\ g~dium :b well
\Cgnw nK It arttLttlares w1th rhc la\l thoraC il se~
Th~. pk ur.t rend tu be rathet Hat 111 some
ment hy m~.·ans ofan arttcul armg ha.ll:..riug, :md tfthl·
Cambn.tn genera . though not tn .tll, but the) are rhor.tt ic 'cgme nt' h.l\'c pleural t"Jcct,, 'iO do~.~ the
Jtim .tr~.hcd or bent dm"n dtst.llly Ill the l:tter
py~ridwm. With thc L'XCep non of tht• Agnn~ti
,;roups and arc nomlJl ly nurkl• d '' itlt .1 pleura l tur- d.t
mn~t Cambr ian trilobttc~ have ~mall (mi
' \\ or le'~ nfren \\ trll .1 ndgc. u~u.tlly tht.: proxm ul c ropy-
gous) pyg-Jdt,l, whilst posc-C :unbn. tn genera tend to
PJrts ot the pleur:t luw p.tr:tlk l edges agaul\( \'1. hJCh have h cter opygo us (\lll.lllcr than the tcplul on) or
-ltr cd~c' nf adj:tc('nt pk·ura .·.m hmge; till' dt\t,tl isopyg ous (cqu.tl-stZL'ti) pyg1di.1. R..trdy pygidt.l
parts do not roud1 .md .m: free. The JUnction ran lw m acrop ygous (l.•q;er rhan the Leph.llnn). "
twn~n till' frt'l' Jnd lun~ed p.trt\ IS the fuJc rum ,
:H 111 tht: Lilhtd a. In Etltfllllm t.\ rhe numb~.·
r of .n,,,tl
wluch the pkurJ .1rc oficn ,Jurply do\>\ ntunw d. scgmc:nts 1~ mpetm tllll'ra ry, prob.tbl-y throug h the
t Jtnbnan genera h.we ,t smtpk artJCu latmg hmge
dtVl\1011 of cxiqm g axi.tl rinh"'·
'rructurc m wh1ch conuguou~ plcut.l l edgt'\ alone
In .tll pywdia rhe pleural fi.trrow' arc homol ogom
r used ti.1r Jrttcul ltton. "pcn.l hzcd htng<- 'tructu n:s With the axtal furrm\ \, "hibr tht mtlrp kurJ!
md arttL uhmng pleural f.tcL·t~ dtd nnt appear until fur-
row\ .tre cquiva lcm to tht: l'llges of the plcur.1; the
rh~ Upper C.tmbn .m: ll\Jil\ of tht'\l" '"'-'m to
be onto~cncttl rcasom tor rhi~ arc dt'< U\sed l.ltcr
I\Ooarcd \\llh enrollm cnt ablltty. The: pkur:.tl dcHJ-
blurc may lw witk or tl.IITO\ \ ,md " oliL•n orna-
Appendages
ncmed with terr.JLC nt!g~.·~. AJ,n to be found in rbc Junb~ .Ire known only Ill a t(·w ~pClll'S uf lri)o-
,ost-C.1mbnan \to( ks, notabh F.unily A~.tplud:te.
httc~ <l' the'c ddtcat c ~tntl tu re~ dtd not ptL"\er.
uc small proLUber.mcc' (ll\ tltl' innl'r f.Kl'' ot the •e
well. Appl'llJagc~ were tirst desLnh cd 111 IH7(, am!
thoracic doublu re,, whtch prL'\ltt nably .1crcd as \lOp\
h.tve bl'L'll reporte d, nti.cn Ill a \ ery fr.tgrtll'lltar.
reventm g O\ crghdm g of the pleura h tht:y caml' to
'tare. m about 20 spectes. They are well kno"" n <Jnd
r~ll dunng cnrolll llcnt I hc~l' .trL' the o rgans
of hwe been fully ~.bcribcd in 0/cnt•idt's (Whjtt lllgtnlt .
Pander , thl \ arc not to b1. lUttfu, t•d Wtth pandc r- t<J75) and Ktwnu a trom the M tddk C.tmbr i.lll
ian openin gs winch. though ~illlilarlv loc.trL·d, Bur~l'\S Sh.tlc of Britt~h Culllm bt.l, ,md ,tlso
111 tiH
wbJbh Wl re tht. onti<L'~ ot 'OIIll kind of \cg- pynuz cd Onlm tct.lll Cryptt•lrt/11" h} R.aym ond
tentrd .md pms1bly cxcret orv org.tn, .
( 1920) .tnd from utrollc d 'Pl'l'im t·m of thL·
The most htghh moditi t·d pkur.t of .111 ,m: tl' be Ordov tcian Ca,wn ts by Stormt :r ( l<JYJ). fhl pyn-
,ound in r amily Chl'lnt nd.JL (Ord.- 1 kv.) .md
tizcd Lower Ordov in.tn 'J'ri,mhms ami De" onian
5upt'rf,unil). Od{lnt npkuro idt.·· (t)rd -1 >e'.; f 'h~ J>l"u''fl·' (I 1g. 11. 14) have yJddcJ much dct.ul
ll.ll.t, 11. lf1). lu the t(mncr the dt~t.1lemh m.l) be throug h dtsscct mn ami X-r,tdJOgr.tphy. In all tht:\L'
uriou\ly poune d. but tht contt~ruom p.trJild t.·d~cs the ~tntcntrc of tnloblt c apptnd Jgn 'hows rl m.uk-
x eqUipped \\itb llltt•rlnckm~ ledge~ whtl h prc- ,tblc: const.l ncy in that there •~ ahvay~ ~ir~t a p;Hr nf
tmably f:.tnlitatcd h1ghh dliut.:n t hingt• .trtiCu b- lllllr.Hnou\ .mtt'lll lJe loc.Jtu l on either \ldc of thL·
368 Arthropods
hypostome, followed b~ p.1ired b1ramous (two-
branched) appcnd.1ges, under the cephalon and one
pa.Jr for each ~egmt·m all the wa) down the body and
also 111 the pyg~d!Um . The~e btramous appendages are
almost Jdcnucal to each other 1n all but s1ze. The
constanC) of morphology here comrasrs with that of
the vanom cmstan·an group,, where there i~ wide
varianon even Wlthlll thl' nnl' gl'nu,. Yet within the
trilobite~ whche appcnd.tgt'\ Jrt' known there is
indeed ~ome dJVeNl). J' \Vtll h( ~hown.
About 15 'Pet till cm of Oltlllllllrs serratus (Figs
11. I2, I I. 13,1 c) \\ tth lunb~ .uc known from the
Burge~~ Shale, and the .tppcnd;tgcs arc preserved J5 a
fine dark ftlm, u\uaUy 111 plJct but 'omerimcs
detached.
They .m~ flattened ,lJld h.!Vc a reflective surface ~o
that many det,nh Gill bl· captured on ftlm, given Light
of suitable indjnation. The :wtcnn.le arc composed of
many JOinted rin~ .•1~ .ue the cerci, which arc poste-
rior equivalent\ of the antctmae. Like the antennae
they almost ccrtmnly had .1 wmory function; they arc
providl'd with nunutc setal h.urs. Bclund the anten-
nae there .1re three pam of btramou' .tppendages
under the ~.cplulon, '>CVI.' ll on the thora.x and Figure I 1. 12 0/enoides serratus from Burgess Shale IM
between tour and 'LX on the pygidium. Each bira- Com.), British Columbia· US Notional Museum no. 58589
(Photograph reproduced by courtesy of Professor H.rl
mous append..t~c wa~ <tppan:ntly attached to an Whittington.)
apodeme, and even where the appcudag~ are not
preser\•cd the number of 1.eph.ilic apodemes will
indiCate the rd:mve number of cephalic appencitges. the last podomcrt.· has three tt"nninal ~pmes. hom
The ba-..11 JOlllt of the .tppcndagc (coxa) g~ves rise to the coxa tht.'rc .tbo prOJCI. rs tht" ~I br,mch, I)in~
two branchc:.. one the walking leg and the other the above the leg branch and dtrected postenorly. The
gill bra nc h. ~m h tcmuuology presuppo~e~ definite mdividu.ll gtll branched ovcrl.tp markedly. Each h;u
function.s, and prob,1bly the gill \'>JS used Cor functions two lobes: a long proximal lobe (thrt>e tunes ,~.,
other th.tn pure!> tor respmtion. However. other long a~ it i~ broad) .mcl a \horter ovoid distal lobe
terrrunology pre,uppme' homolog~es, and though the The fom1cr ha~ a postenor fringe orlong paraUd til-
tenn telopoditc t~ olten u~cd tor the walking leg, the aments dimim~lnng in stze tow:trd~ the di\tal lobt,
corre1.·t tenmnology for the gill is not agreed; for the whereas the bttcr h." no filaments and only a fnngt
moment the term ·g~IJ branch· 1s rct:1ined, though of setae. An anterior marginal rim ori~rinall}
outer ramus i~ .1 good altc:rnative. described on rhc gill branch tS now shown to hJw
The coxa i~ ,tbout half the length of the axial ring been a prcserv:ttional artef<tct.
with winch tl ~~ asson.1ted. and 1ts inner margin is Precisely how rhc coxa fitted on to the npodcmt
supplied wtth long sharp spmes. ~tl':light and curved, is uncertain; presumably thl· whok <1ppendage rould
between which arc numerom shorter spmes. A pre- rotate in a hori:zont.ll phm~ . bur tt is not kn0\\11
coxa. or tnltl,tl 'c-gment other than the coxa. has whether it wa~ capable of any do"o-vcntral or
been dc~cnbed but r~.·c·t·nt work doe~ not <;ccm to transvcr<~c movement.
~ub~tantJatc m prc.·,cncc 1 he walkmg leg ts contin- Before conMdenng how thl' appendages futk-
ued in the direct line !Tom the coxa and consists of tioned. it ~cctm appropn.m to cowr the range 1n
stx JOmtcd segment.. (podomeres). The proximal fonn of known appcnd.1ge,. ~~·ormitl appendagt" art
podomer~ arc ver\ spm> on thetr inner surfaces, fairly '1milar w thme of ()ffll!llcln but an: known only
the more di~tal ones le.,, 'o but wtth fine setae. and from one 'pcctmcn. Tnartluw i.< a Lower OrdonCJan
Trilobita 369
(a)
le) ldl
(b)
(f)
0/enoides sturetus
(g) (h)
~
I
I
,
'if
,
~ <{
*\
I :t Trl11rthus
•
I
r
I
'*I
I I ~,
r
I
•
I
Figure 11 . 13 Trilob1te appendages. (oHc) Olenotdes serratus
+
I
(M. Cam.), showing reconstructed appendages: (a) dorsal view

I
I
, with port of the exoskeleton removed, showing the appendages
~
~
in place and the gill branch lying above the leg; (b) section; (c)
I lateral view; (d)-(f) Triorthrus eotoni (L. Ord.); (Cl) reconstruction
with right side removed to show appendages; (e) frontal view of
walking individual; (f) a detocheCI appendage, (g) Phocops
Phecops
(Dev.) leg with a terminal brush believed to hove mode stellote
markings on either side of a central groove; (h) Cerourus (Ord.):
(h) limb in frontal view (above) and gill branch in dorsal view
(below) [(a)-(c) Redrawn from Whittington, 1975; (d)-{fl
redrawn From Whittington and Almond, 1987; (g) based on
Seilocher, 1962; (h), (j) based on S!Ormer in Treatise on
Invertebrate Poleonto/ogy, Port 0 .)
370 Arthropods
olcnid When. found m rhc ·uor.t \late' (acmally a l•l
bl,tck shale) of Ne\\ York State the appendages arc
pyritizcd. FiN di,covcrcd a century ago (e.g.
ltaymond, 19:!0), lhcy have been tht• st l ~Jt'Ct also of
recent inve~n~.tt1on' (Ct~ne, 1975. wh1..) mcd X-radi-
ography; Whittin~ton .tnd Almond, I tJH7) The latter
authuT\· recon,tntttion (Fig. li.Ud-1) ~haws anten-
na I..' ,md three p.lif\ ut b1ramom n·pluhr .lppendagt."S
umtomt wHh lhmt' of the thorJ.X. 'I he walking legs
arc spiny, wilh a cox.t .md six poJomcn·~. E.tch gill
consi~t\ of tilanH.:nts JOined to J Aexiblc. segmented,
anterior rod, :tnd the dist.J.l lobe ts n·duccd :tnd tiny.
The appendages wnc 'tccplv mdmcd below the
c~oskdeton .md thd not projccc much ouNde it.
The use of soft X-r.1y' Ius aho bct•n u,eful 111 mter- Figure 11. 14 Phocops, on X-radiograph of a slightly crushed
pretlng the apperH.la~c 'tnll"mn: of Dcvom,m Plwwps specimen from the Lower Devonion HunsruckschJefer, Germany,
and .4stcmp):<:c spt·cuuens from till' Nnrth Gem1an showing lightly pyritized gill and leg branches. !Photograph
llunsrilckschierer (Stn nner and Tkrg-;trl\m, 1973). reproduced by courtesy of Professor Wilhelm Sturmer.)
These specimem of PfllltrJp.s (Figs 11 I Jg, 11.14) have
\tOUt walkin~ )Cb'\ \\llh the ll~U;tl \iX rodnmt•res. normal construction, i.e. donbrated thin platl·~
There an: three p.u~ of b1rJIIHHI\ .1ppcndagcs ,\rrangcd subparalld \1.-Jth one anoth er md h:mng
bclov. the tcplulon, the la~r two h.wmg strongly e.lt.h .1 terminaJ bristk.
developed coxae prolongt·d imnn.tll> .tnd an11cd ·r he diversity of fonn in the gtll bra m h unphc.1 (h )
wtth sptne'>. 1 he~c l mrld have functioned as som e differences 111 li.tnLllo n. lt i~ gcner..tlly arn:ptl·d
gnJthobases eqtuvalent to thmc in ~omc modem that the gill branch did in fact function for rc\ptr;l-
.trthropods, ami by rolling tnO\'emcnt'i m J horizon- tory exchange, though Bcrg;rrom ( 1973) hd'
tal plane thev llt.l\ llJve been .1hk tll work ,,, cmsh- pmntcd o ur that suKc they arc rdauvcl> hard smrc-
tng Jaws. There 1s mlt' pair of appt·nd.tgc~ tor ea1.h tun·s they could have functiOned mcrcl) '' .ter.ltor;,
sc~ment of tht thmax and there .trt• ,tJ,o 'cvcral to .tgJtate the water wlulc the anunal wa~ movm~
small ~ets tn tht pygtdJUm. Tht· main difference .md so to feed m o rt· oxygen to the t-,'11h, wh1ch hr
between the appt·nd.tge~ of PfttJfllJl.' Jnd those of imagined as soft s~nl t· turc~ :.tttachcd to the vc.:ntral
nther trilob1tcs 1s thal the gill branch tilamcnt~ me mbrane. The pt~ddle-l.ike g ill br.tmhcs uC
emerge directly ll·um the coxa ami ,1rc transvcr$L Cerattrrts and 0/ctwidc~ could probably h.tvc fiKLh-
and subparaUel w1th t:ach other. The whok· ,1rray of t,ttcd ~wimming, but there i~ no dircLt evidence as ...
gill branch tilameut' fonm a kind of ,l.lttcd curt;lin ro how well trilob1tes could swim. and the quc,tJon
lyrng dorsaJ to tht• \\ alkmg le~~. wlm h may haw rem:ttm open. There IS, however, good evidence as
lud both a resptr.ttul) and a tiJrering fun1. non. Lt·gs ro how the walkmg le~ operated, and tltt' dcwiku
111 the phacoptd tnlobttc R.hctW[I~ h.tVI.' recently been ~tudy of these hJS y1cldl·J a surpmmg amount of
reported (B erg,lri\m and Brasscl. I \)H4). infom1:ttion about tnlobitc behaviour.
Tht' OrdovJcun c heirurid Certllll/1.' (ftg. ll.l3h)
has peculiJr appt·nd.tges, ;15 \hO\\ n b) Storm er
(I ~39) who nJ.ldl sc:n,1l stcttom through enrolled Trilobite tracks and trail s
speomens collt·ned from a lime,tont• of Tremon
(U. Ord.) age. l hl branth structure~ were worked E.trl) in the history of p.tlaeonrology elongated trail- Ftgure 1I
out from a w.tx mndcl bUtlt up l,lyt•r b) l.tycr from hkc markmg:; were described from many dJficrrm by pushir
these ~ections, .ts Ius been done wtth tht• grapcoli le~ st·dimcntary rocks of P:t lacozo ic age. The common- hoe also
oppendo!
(q.v.). The >w.1lk1ng leg 1s rclatJVel} unmoditicd, but est ktnd, tirst called Cm:ia11a (Fig. 11 .15a) .md bter Rusophyc
the gill branch h,t, five scgmcnb mcrea~mg in ~ize rdcrred to by scver,tl ,lltrhors :.ts 'btloblte~ ·. arc nb- (g) termin
dr~tally. the l.m llf whrch tonm a fl.lt paddk-hkc bon-hkc markmg' \\ 1rh p:ured oblique cht:vruu d1stinguisl
strvcted v.
lobe tTmged wJth til.tmenb. Thc~c til.unents arc of ~tructu re~.
1962, (g~
Trilobite 371
(I) (d)
push•ng
marks
I
I Ru$ophycus
~-
Cruzillnll
(cl
Rusophycus
(gl
\ r
'\ (
\ width of set
f length
of set
""\ I'
'-palr-1'
Asaphoidichnus (lsote/us)
' " " - 1mpnnt /
r-
il
f
I-
ll~
n
\.
:d
of DimorphichnitiiS
il- ·"I I 15 Trilobites os trail makers. (o) intersecting Cruziano trails showing how o 'ploughing' trilobite could have mode them
Jmhmg backwards with its legs - the lateral striotions on one trail could result from the gill branches moving across the surfaces
tlt
ulso Fig. 12.12o); (b) Rusophycus from the Combrion of Poland, showing resting marks of coxa, and pushing marks mode by
11- ~ages os the trilobite pulled itself sideways out of its burrow; (c) trilobite in o shallow burrow; (d) leg marks preserved in o
tcr 10phycus; (e) directions ot movement of the legs; (f) o burrow mode by anterior and posterior legs moving in different directions;
h- 'erminology of o set of walking marks; (h) Trochomotichnites, o trilobite (trinucleid?) walking trail in which individual sets con be
""\JUished, (j) Asophoidichnites. sideways movement marks mode by o large (o.sophid?) trilobite, possibly /sotelus; (k) recon·
'01\
rlBd ventral view showmg how D1morphichniles was mode by o sideways-crawling trilobite ((cHO Redrawn from Seilocher,
2. (gHil redrawn from Osgood, 1970; (k) redrawn from Seilocher, 1955 )
372 Arthropods
Thl'} \\ere tir;t bcltc\·ed £•) bt• ot Jlgal on gm. m J\\llt i,mon "ith the Ru(t>p/t)'Cfl) marks (hg. 'fl
l-l1l\\ l' \ l'r,N.Hhol'r ( I hS I) l .tllcd .mcnnon ro rhe ll.l"ib) ,1\
t:•n dur 'b1lobirc•: ;J)W.l~' lluurrcd ,u the Interface:~ C1
o( ,,lnd: .md ,J1.1ly bycr<> in tht rock . Tlu~. he pre- 110
sumcd, \\:1\ more likd) to h.1n bt•t•n the result of a Walking mo vement in arthropods
In Jll trawling ::arth ropod~ thl' movement of the lnl
mm mg .mim;JI cr:twlmg ova or r.lthn ploughing 111J
limb~ rdntivc to one ano rhcr t::tkl·~ p l.1rc 111 the ~ame
throu!):h .1 tinn muddy ~c.1 Anor, le.wing m,uks
whit h h:1d bel'll tlllnl wath s.llld \oon .1ti:cr. the dif- gencrJ I way. Leg movement\ on c.tch .;ide of the
bod) arc sytKhroni.zcd in rq;ul.tr w.IVl'\ of forward the
t\.•rcth t' 111 \edantt·nt t) pc being rt·,pnmiblc for their \V,!
prc,erv.triou. Nnl intTI.'qm:ndy Cm ::1•111•1 mJrkings movemcnl, 1-.nown Js metac hro nal rhythm. Each
.1\
Jrt• lllt ,lcrms h\ other .md prt'\lllll.lbly l.tter traih kg tome\ forwards in mm, 1\ pl.Ked on tht gmund
and pushc\ backward~. hdpmg to support ,1nd move Tra
nl.ldt• b} the sJnu: kmd ni .tmm.ll,, continuing the fiN
thl' hody an the procco;s bdund. At .my one rime 2
moblk- .mmul hn'othc't~ . Orha kullh of ntJrlang-; l .• ,t
l.'Xl,lln~ on 'hak-..md,tonl: nHt-rf.an~~ could also
lcg will be ~lighrly the one in fi·om of it 111 H' fitr-
1110
w.ud movement. so that m a long-bod!l'd arthropod
h.1n~ bet•n made b> movmg .tll!Tllal' oft
lt ., gt't1lr.ill) bdJt'vt'd tlut CntZitllla m.trking; (e.g. a tl'ntipedt>) successive waVl'' of motion mJy
hmt
were Jll,ldc by rrilobiu:,, tlwu~lt Whitrington (19RO) be ~ecn tr.wclling up the body from the t.1il to tlw
11101
h.l~ cnuttzed pn:viom intcrprl't,ltHms, and suggest\
he.ld. A~ the waves o[ movell!L'nt sweep lorwJrd~.
llttp
th.\t they wert· uton: lil-.cly to h.t\'l' lwen made by evt't) tcnth leg or so is in approxitn,ttdy the 'amt'
(Ftg
\OIIW nther or~ani,m.
posH!On .1nd at rhe s:une Jnglc. Even 111 'hort-bodt~d
Sin~
arthmpods (e.g. woodlice) the p.tttern of mnvemc:nt
I herl· '' htuc Jtrecr C\ idctKl' th.u thc,c and other Ill Ill
IS dc.1r. and the waves of mo\ement ah\ay~ tr.l\·d
kuHh ot nurktn~C> \\'ert• actu.tllv m.tde by Jny one Xtph
forwards .tlong the body. Tlus w.1, evidently thl·
type ,,f org.uu~m. 'u1ce tt.tn· to"''' of anv kmd .tn.
l'.l\l' m tnlobaes, and the walkmg n1.1rk~ espeCially
V,).
r.udv tound wtth bodv to"ik C.tll itit·d ,hell\ are not ,1£ th
lll'mully rct.um·d by tht• pl.:'t1lll.tbk \Jnd\tOnL~ in m.ty be mterpreted on tlm bas1.s.
h.!Vl'
whtch the trace fossib mcur. Ho,,cvt·r. ;t t\.-"v spco- 'I I
meth oi mt:tct trilobtt\.'s lt.IVt' bt'l'll tound in the Different kinds of trilobite trails COil\!
Ordovtnm nf Clllcitut.Hi (Os~ood, 1970). where T he vn riou~ sorts of tracc fossi l~ belaeveJ to havl mark
C.!lymt•m· ~pecunen~ wnh inl.td llllldc~ h:wc bc(•n been m:tdc by trilobites arc clamfied .lt'lOrdmg to bc..·r c
lt)l!nd an thl'lr restmg ext.l\',lttvm (t.tUed Rwopltyws). the binomial ~ystem of nomcndature. The retcnuon ' hnw
Other l ntt·ri.l tor atmbuttn~ p.trllutbr trace tu,sil~ to of lmn.te,ln pr.tctice for \uch fonn genera .md the tr
tnubue movements Jn: mdtrl'l t. though compdling. form species is unavoidable. smcc ar is vef) raff Otplit
TJm, C.1mbn.m and Ordono.tn Cru:::itllhl Jrc numcr- that .1 particular trace fosstl can be dtrel'tl) related to llltl'rf
011' .ltld di\'CI'l'. whert·a, the ~cnt•r.~lly poorer Siluro- a kno\\ n producer, and even when tt l'Jn, both th~ othc.-r
Devllm.m C111:::rmw fauna rdlects rhe dcdinc of the trJII md chc body fossil rctam the1r O\\ n nJme-... 01 Then
tnlohtte,. Although tnlobtte' .tn: nor !otmd in the vanom fi.1nn genera that have been attributed to bl· idt
Cnt~tlll!l-be.tring s:tndsrones. they lliJ) occur in tht• hie activities of triJobttes ~omc of the moq IK·twc
imcrc.tl.ttcd bed, of the ,,mle 'eduncnt,ll")' sequence, im portant .lrl' 1ng le
,,, 1n rite Ordovtn:m of Pol:lnd .1nd the Devonian of tnlnbi
Gt·ma.tn~. Most Cm:::itlllrt .md other 'unilar markings Protidlllitcs, Tracltomatirlmires: w~lktng or srndm~ progn
\\'l'rc dcarl} made by 'le~· of umlonn and undiffer- traih with individual leg tmprc,siom (Fu~. an i m.tl
cnll.ttt•J \tructurc. such ,1\ .u,· pm't:''t:d by rrilobite~. li.ISh); ' u rfJct
and \lltllctinll'~ the 'prod nurk\' n1.1de by walkmg le~ Diplidmitrs, PNafu·lmite_,, Asllphclltliduws: .1\ .tb\lvt In t
cnl be m.ltl hed warh thl· Ktll.tl kno\\ n ~mKture of but usually oblique (Fig. I I .lSJ). WJ lkm
till' dhtJI part of the kg. Furthennorc. th\.' common Cm :::w11t1: bilobed chcvron-m.nl-.t·d rr:u]s wha1h tern1m
bilobt·d .md uonn.tlh tWl'ld tr.ll'c tosstJ... known J.~ arc the trace~ of crawling. lnu~;htng, c;,ho\'clling or H unsri
Ruwp!Jy, w arc ju't tht' nght 'ill m h.lVe been made burrowing movement~ (Fig. I I .I Sa); f>/ltl(clf!~
by tr ilobitc,, Jnd ~nmctimt'\ mdent,mom caused by RIIS<'phyms: btlobed ovoid trJet'\ wht< h ,1r~ prob lnt:tlt I
tht· Hllpre,~iott of .1 ccplulnn, gt'n.tl 'Pllll'S or latcr:ll ably rl·~ting n cm, burrows or ,urt:1n· cxc~vatl{ln' natc 111
p.1rts of Lhc bndv rc~ting in tlw \edirncnt arc preserved (Ftg. l1.15b -f). claw,"'
Trilobite 373
1h~c moVl'mem tr.nl~ would bt: gt:ncr.1lly .::b-... itlcd idt•Jil) with rhe ~tclllt<.' imprc,,mn ' Jrr.mgt.·d in
;r, Rcptchm.l (cr.J\dmg tr.llc') :md Rrt5tlp!lycw as p.mt•d ,crie' on either Side of .1 ml.'tlt.m gro<WC', ~m h
Cubichnt.l (n:~tmg tr.Ke~) ••ttrorditt~ to tht: taxu- JS luvc been fo u nd at the ~ilt-jialld\tllt1C mteri:JCC~ of
uomic 'l'hcme of ")t•tiJchcr (Chaptcr 12). A '>lltglc bt·d, 1t1 the 'i:Jmc ~equl.'ntt: Th1' l\ one nf the few
lrilobttc rould wdl ha\'e nl.ldt d1tl~rt'nt kmd' of cast·~ of uneqluvoc JI m•ltthmg. In the Ordm 1c iJn uf
mrking, tkpcndmg on ~.·x.H:tly wlut tt w.1~ domg. Cmtmn.Hi thL· !JrR<:" ,1saphid l.•••tdw 1\ pn•,em in .1
Though trilobite'' often w.tlked '>tr.ught ahl'.td, ~elJUence that :~ho contaim the H lmolo"1l
rllc:y frequcntly prnwe,~cd obliquely or 1. vl.'n '-tLk- or
.·L~•li'Jtt,idiclmw, wluch t onsists p.1ircd 'et' of trifid
1\S like a uab. In the ca'e uf forward nHwement , unpnn~ (Fig. 11.15j} Tlm trJrc t("'il t\ lar14t'
11 rcprc\enlcd by rhc trad; ~ of Pmtrd111itt'.1 .md enmtg;h to havL' been nude by /~o/dm. ,tnd mdecd tt
TradiOIIIctlirllllifes, the pygidi.tl .tppend.lg<.'' wert: the m;ly give J llliL a~ to the 'tnKtl.Jrt· of tht :lppt nd.t~t·s
first to tom h thL ground. followed by thme of the of du, trilob1tt, whid1 .trc otht'n\ tsc unknm\ tl.
~~ thorauc 'cgnwnt, .md \O on until tht• W;tvt> of l::.vtdcnce of ~trongly obhyuc tllOH mcnt ts
movemcul n.:Jched tht· head, ltleanwhill- new wavt•s known only in r:lrt' case,, The wdl-kno\\ n
•I movement ''en: pro~;,,-cmn~ torwarJ, !Tom the Dwtt'lpllicltmlt's (Fig. 11.1 Cik) !Torn rhc C.unbrian ,,f
nder l'nd The rt·,ult of \liC h ~ucct:\\1\'t' \\'JVL'' of the ~.lit Ran~c of P .lkt\t,l11 is one of d1c't'. In 1. ll'h
novemem i~ a writ•, of 'L't\ of pJt reJ markmg'> ·untt' of Dirnorpltirhmrt•s there .ue two ekllwnb: on
rnpressed lik.e opt·n-cnde d (tnmc Jtcd) V _,ha pes one l>tdc .1 \L'I of round .tnd dccply unprt'\\t·d
~· ll.ISg; Seilacher. 1955, llJ&4; O'gtmd. 19711) impnn£5. on tlw other .1 'et of h~htly unprc"nt and
'mce the tnlobite '' tdens antcnorly from the py~td ~hghtl) sigmotd<ll r.tkmg mark~. ~ct.lacher\ ( JlJCj:'i)
mt, the V is dirt'ctt•d forwards (though ~ome interpretat ion of these ts thar ;~ trilobttc (probably m
1lJihosund' movmg tc1rward~ l1.1d a sent'' uf rcvu-.cd olendud) moving crabw1~e from 1..-tr tu right "ould
1. Where th(;' ~tde' of the \ do not t: un c in tg.un pre'~ tts right-hJud lcb" u1to thl· 'ubsrratL .md. ll\ltl);
nhe from Jt seetm that tht· lw.td appcnd.t~es cannot the,t• tor suppon, would hc:tve tt~ body ~id..:w;~y~ by
1ve been used in v•. tlkmg. comractm g il~ leg tllli~L·k~. Then tt wouiJ dng JL'
1nlobttc\ mm•tng dtrecth tl.ln,·ard, m.1de a tr:IL k lett-hand ll'~ acro.;, tht mud, nnh: u~ing th..:m t'l1r
1qstmg ot a ~cnt' of ~upt•nmpml.'d 'ets of V- minrmal supporl, and b) withdr m · tn~ m rtght-hand
nJrkings, \O thal it j, 11llt e.t\y to ~ort out tht• mun- le~ one aflet Lhc other Jnd repLKm~ tht.•tn ~omc
. of tmpnnt~ per sd, though a good cxamplt: i' di'it.mce to the nght would then be: m a pu\lllou Lu
'.own m I ~~ 11.15h. WhL·n rnlobm.'\ "alked with repe.lt tht· prmedurc. A number of complete mu\·e-
ir movement obhquc to the body .txis (.1, in ment'i of tlu' kmd can hi.' clearly ~c~:n on tht· luwcr
')jpltdmitcs). the intprinb Jrt• dmdy crowded .wd '>Urfo1ce of .1 ~andstonc umt. Tn one CJ.\l' ,1 Lu\\cr
crfenng on om: ,tJc bm ljltlte -;ep.u.tte on the C.1mbrian olendlid, mm mg !Tom the lct'i by sur h ,,
Kr, and \O the) tan more e;1stly bt: counted. modt• of progrcssmn , cn.::ounta ed the tr.Kk of
tre 1~ then sonH.' t h..tnn· th;~t rhc tr.ul m.tkcr t.an .motht'r olendhd. turned round .lga.in ollld movcJ
identtticd. as tlu:rc 'hould be some ktnd of panty b:tck .1gain p.tralld with it'i own 'icr of track' Then it
't'.Veen Imprint number and the numbt:r of\\ 1lk- jumped O\'er tt' own tr Jl b .1nd mm·ed n\ ;t) 11 n~lll
~ leg-;. Olfl/ic/mitc., rmght h.!Vl' been m.tde by J Jngle'i to it~ first track by ~lightly oblJquL' torw.ml
lobtte walking obliquely ,t, it~ normal mode of movt·mcnt~. Seilachcr lntcrpret~.·d rhe~e ·'' grazmg
,rr~s1on, or 1t t.ould have resulttd from tht: tr.Kk 'i 'illlCe the tnlobttt• ohviomly J\'Olllt·d pre\ i
:r..U trymg to keep 'ite.ld> on a ~hppel) mud ou,Jy made trJd(s. Simtl.tr rr.K h h:l\'e bc:t"n fL,und ttt
~bee whtk .1 lateral curreut w:l~ Aow111g. the Ordovil i,tn, apparently m.ldt· by trtnudctds .
In thc~c vanou~ walkm~ tr.tth the m.1rk~ of the (ru::rmltt rs .1 very w1dcly di,rnbutcd tr.1cc fiJ~,il.
ling leg often rctJIIl the detailed impnm ol the .md up to JO different tonn 'pccie, h.1w bt>t:ll
-mmal dtgit whtrh mJdt· tlwm. In rhc Dcvonun UL \en bed, thou~h the~e .:.111 ofi ..·n be attnburcd tn
umikkschiefer of Gcmtany many ~p..-cimem of ~omcwh:~t diHcretlt burrowing tt·rhniqm·, b\ tht·
· pl arc fow1d, \l>llle with the1r JppcnJ.Jgt·~ ,,llllt' animal or ro the mclination of the bod) of tht·
t (Ftg. ll.l3g; 5etlacht·r, llJ62) These temli- amm:tl on dtffrrent f:,'T,Jdtt>ncs. llmYCVt r. the bJ,tc
cma di~tal tuft ofbrmlc,, ennrchng the terrnmal Crtt.:::iall•l 1~ simple: .m dongal<.'U h1lobed tratl. each
., whilh if exwmicd hkc .111 umbrdl.1 would tit lobe being t'Oitvex downward ' .md mu,11ly ~tri.ltcd
374 Arthropods
wtth oblique. clo~dy 'PJced ndgt.'' and grooves been found m the OrdovJCJan of Cinnnn.1t1 ~n11 hu
fonnmg a hemngbonc pattern (F1~ I 1.1 Sa, 12. I 2a). with tnlobttes in them. though only .1 vcr. ft'w Ho
The rrilob1re must have ploughed 1t~ way along the example\ have actually bccn dtslowred. Some py
upper surface of the sediment, HKI~mg the~e mark- mtere~ting examples descnbed by ~ctl.tcher were .tti•
mgs as 1t went; in the be\t-pre<.crvcd ~pccimens clearly produced by the .tntenor appeudJ~(• 13c
there was probably a thj11 byer of~and already over- ~hovclling forwards and the postenor nm:s workut~ ht:l
lying the plastic mud that took Lhe impressions, pro- backwards; h ence RIISophyws ha~ .1 bidm:ctJOnal .dlt
tt.•ctmg tt from current t"ro~ion. More sand later ~rructurc. Other examples wen· lpp.Jrcntly exc;~
filled rhe markings. The two lobe~. with their vatcd by mmg the anterior border it,clt ,,, .1 \ho\d Old.
obltqm· ~tructures, were prob.1bly made by the ;md .;aabbhng ''r1th the postenor JppendJge\. Such lit I•
walkmg legs movmg im.,ards rowarm the median mod1ficanom of nomul pract1te .tllowed the iomla- lril
hne md backwards. The ob!Jqut: \l ratchcs seem to non of deeper burrows than thc nonnal dwUow of';
have been made by div1ded tenmnal dJ.ws. though resnng traces. The trilobHe could e.t,ily ~et out ot Jtl tha
as they are generally supcnmpo~ed one upon nc\t by pushmg Wlth the w:tlktng leh~· but one 'Pr
another, the marks of individual legs are otten diffi- intriguing example from Poland (Orlow'>kt et tTI.• un
cult to distmgmsh. 1970; Fig. 11.15b) shows impnnts of the le~~ out- PHI
V .m ant~ of the basic struclltre :tre often found. side the burrow. though only on one ~1de .as c\'1• ex c..
Sometimes brer.:tl ti.mowed bonkrs arc present, dently the :mimal had levered lt\elf out stdcwm lll<l
probJbly the scratch marks of gills or pleurae. before ~wimming away. This .md other exampb hig
Cerr.un Cm~w1w spccH."S have .111 extra pair of lobcc; ~how the grooves where the coxae had rc\ted. Hot
wHhtn the omer fiJrrows. perhap' the brush nurks Some 'hunnng burrows' or Rmopllyms lrom the Cat
of the g~ll branches as the trilobite ploughed tor- Lower Cambnan of Sweden g~ve t:\'ldence of FfC- I
ward,. Cnt.::lalla has aho been found wah apparent d,mon by trilobites, probably olt:nelhd\, or mtamul .md
pyg~dtal ~cratch marks J\ well One species shows a 'wonns' (Berg-.trom, 1973: Jemen, 19YO). hwm;- Od
hcmngbone pattem clearh subdtvided into sets thrcc known examples of R. drspar ;m: dug do\\11 to (
with between three and mnc parallel ~cratches; each U-~hapcd ~prcitc burrow~. wh1ch were probJhh nnl·
succcsstvc ~et overlap~ .md tnmcares the preceding made by a priapulid. This prey mmt have b~t·n 'ucl
one m ,1 mannt•r ~uggesttve of altcrtute stronger and located. chemically or vistJJlly, .tt the aperture) ol fTm.
weaker pushe,. On ran~ occasions a parttcular the burrows. The Rusop/1 yws tr.tt'C\ .m~ .1lways .llltl
Cm.::1011a 5pcoe~ CJn be matched wtth a particular located in contact with but ~tightly to unc ,.de- ot Sint
trilobtte, where there '' only one ktnd of Cntzitma thl..' wonn burrow. Thi~ suggc~ts that the prcy .,..._, h.\ V•
and one ~pccie\ of a~\OCiated mlobtre (Forrey and captured by the rrilobitc digging down Jeep. thrn part
Set!Jl her, 1997), bm tlm '" uncommon. hookmg one set of le£9> round thl• tubuiJr hod}' of the
Rus(1phycus (Fig. 11 15b-f). mterpreted as trilobite the wonn and finally impaling and -.hrc~ddmg 1t \nth 1mit
burrow\ or temporal) remng rraces, is the name the sptnO\c coxac. Where the 'pn:ite burrow IS wht
b11ven to ~hort ovoid markin~ which are nom1ally short, the trilobite was able to ,thgn itsdf pnnr to the,
bilobed and deepe\t in the centre. These may be diggwg. Lon~er wom1 burrows n·quired 'eved cuul
smooth (coffee bean marks) or hJve herringbone successive diggings by the trilohitt· in order to 1t de
ridges as in Cruziww. Tlwy usually occur singly, adueve the correct alignment. T
implymg that a swtmming trilobite lud landed, 1.-t•(lt,
excavated J temporary 'ne~c· and then swum away. the .
The) may aho terminate Cm:::l,tlta trails, as if thc Life attitudes, habits and ecology .• pl;l
trilobite had crawled, rested .md then ~wum off. and that
they have been found 111 a"onanon with striding Acc~•lc (F1g. 11.2c) shows a stanwrJ pattern of con- <ittltl
trails The smooth Rusopilyrus wa' probably tanned strucoon m s1de view. This attitude IS very c..orr.- purp
beneath the body of a re\tmg mlobtte rheocactically mon. and even in such a spmy tnlobnc a' CybtiCJ ·ts l'r:l\\
dJrceted upstream and hollowed out and smoothed (Fig. 1 I . 11e) the spatial and anguLlr relatton\lup' of ceph
by current ~cour. Some deeply excav:1ted hollows the varil)u~ parts of the body are stmilJr. Cybc/,Jid , rtnr
are best mterpretcd as nc:st~ in which the trilobites however, has both genal spine~ and macroplrmJ! whi<.
l<IY for quite long periods, and it i~ the~e that have spines n o the sixth thoracic segtlH:nl. Thc~c would I I. I
Trilobito 375
hold the body in an outstretl hed attitude on the sea
ftoor wlth the postenor thor.~eic segments .md rhe
pygid1um nsing awJ.y lrom 1t, though some rclax-
lhon of the muscles would lower the 't.ulgatc ·.
Bes1des bcmg ttsed J.S props, the spmes could have
~t:'t'l used 111 burymg the rnloblte by a ~crie<> of
dtern.m: AeJ<.'lon and re!Jx.ltion movement\.
llpmO\It) m trilobltl'S was once regarded a~ an
ldJpt.tnon for a planktou1c mode of !tie, the '>pmes
111h1biting the anim:U's sml...mg. Howcvl.!r, most
mlobitcs arc far too Lirge t(Jr the ~pines to have hccn
1( 1ny real value in dm context, and Jt I\ more likely
thlt they were, as wdl .h being protecuvc, used for
1preadmg the we1ght of the tnlobne \\hen restmg
on the ~c.l bottom. Thm long genal spmes, ..ts 111
Pri'lt'M (F1g. 11.1 1 t). bon. the v. etght of the '"hole
~xoskd eton, bemg the nw't ventral part of the ani-
mJI. the thorax and pyg~dnun were earned slightly
htgha than these and not m contact wath the sea
toor. Thts as a common po,ture. espet:ially in the
Dmbrian, and is probably pnnutive for tnlubtte\.
Jlcrlldps the most smkm~ case where \pinu,tty
llld hfl: attitudes Jrl cnrrcl.tted IS 111 the
Odomopleurotdae (Ftgs 11 I !.1, 11.16). Figure 1l. 16 Leonaspis coronale (SiL), Wenlock l1mestone,
Dudley, England: o complete specimen in dorsal view cepholon
Odontopleunds an· .1lw.1y' ~piny and h.wc not slightly crushed (x 4).
only l.ttcrJI bur also ventr.lll) dtrccted sp1ne' Genera
eh~' Awlnsprs and lJrulfq,upi~ (F1g. 11 ll d) llJvc J
Inn~ of mod1fied denticles cxtendmg alon~ the fom Jrd' lt kc that of Dudlry.rsp1s, resting on the
mum-lateral border on l'llhcr ~tde of the gl.tbcUa. antenor dt·nttcle~. wh1le the body 1~ held out hon-
\me.: they rcrminatc m 1 flu pl.me thl') wl'm to zontally (Fag. I I. 11 d). Tht· fim attitude st·cm~ tu
aw bc.·en adapted for supportmg the ccphalon an .l have been .1 rc~ting po\ition, the second an .IC.tiVl' or
'lrtlcular :mitudc, prc~um .• hly whale fccdmg;. tor 'brow~ing' ~lance.
he 'hort hypostome would thc.·n be in do~c prox- Dudlqasprs WJ\ .1 premmcd dcnvativc of the
mny to the sea floor .md 'n would tht· mouth Lco11<1spis stock, permanent!) spl'n.lhzed for ht~ 111
'l'htdtlay JUSt bc!Und it 'I hl·rc ~~no ,mtcnnr arch in the active .lttitudc, since the poo;tenor chor,tuc
e modaficd gcm·ra rmr m Caal!,aplra{,1, \\ htch spme' .md the pygidial spmt'\ .1rc bent down so a' ro
uld support itself 111 J \llnthr .1tt1tude even though termJIIJtc in the same pl.mc .1~ the antenor dcnnde,.
·doe' not have the.· deutide' (Whmingcon, 1956). ln Crratawplwla it i~ the anterior thoracic sp111l'S char
The.· rootstock gem-r.1 of the odontopkurid,, arc modific.·d; they arc large and thick and terminate
' •lllil>pi;; (Pig. 11.11 b.c.. 1 I. 16) and PmlltiSJlis, have in the '.tflll' plane a\ the ,mtcnor border of the
lnt<:nor denticle\ fy111g 111 ,, cun·e rathl'l th m 111 ceph,tlon, showing hm" spcnalization for the .tctt\C
fhne, .md it has bccn pmcul.ttcd (Clark,on, 1W1lJ) attitude h.l\ bC"cn independently evolved.
these tnlobites could t.1h· up two .11rl'manve ltfe The rd.1ttvdy umpecialin·d 1-A•tmttspH stocl.... with
tudc~. both cquJ.lly ltmtttOtldl but for d1tfcrcnt its du.tl mode of life, was VC!) vcr\atilc. Jnd the
rpo,cs. In om: the tnlobitc lies on the ,c,1 Rnor o r strattgraphtcal range of th1~ one genus h.1~ been
·1wl~ over it with an opt'n Jnterior .uch. \\ 1th rhe estimated ,1s 170 Ma. However. its dcrivatiVt'\
.ph.1lon supported by the gt>nal spine~ and pmte- (Dudlcyaspis, Atidnspis and other~) were of relatively
dcnndes and wath the thorax and py~daum, short durataon, illu:.trating thl" general rule thJt the
eh ,]ope postenorly rc,tmg on the1r 'panes (Ftg. highh spco.1hzed taxa did not mu:~lly bst 'Cl)' long.
llb). In the orhcr attitude the ceph.1lon j, olted The spcoalizcd odontopkund Selcnopcllu (Fig.
376 Arthropods
11.11 J) (wh1cil \.\',1\ l'.lp.•bk lll di,coidJicnrollmcnt)

has bl.'ul mtcrprl'll'll ·'' an .ll ti\'c..' \Wimmcr but capa-
ble of rc\ting f(n 'hm t period, of omc on the ~ea
floor (Hamm.mn .md R.tb.111o. I 91:!7)
Throughout the hi'tol') uf trilobite\ certain ~pc
cialized 'nwrphotypc' · h.wc on:urrcd again and
agam, 111 d1lkn.•nt \IOC:k\ . Thi~ cum·crgcnt C\olution
sugg~~b 'inul.u nulu~il ,,I llll he'. Fortcv and
Owe m ( 1990a,h) h.we donunentcd etght \uch mor-
photype~ re,ttlttn~ front iterati\·e cvoluoon.uy
trends. Thu, 'ph.tlonwrph,' .1re tuberculate. con-
vex. ne trlv 1\0pygnu' trilobite' with wl'U-dcveloped
eye\. ~ut h morphoklg)' " charactcmttc of many
unret1ted Ordm. lll.lll to Pc..·muan tonus. Likewi'e
~ome J...md~ of 'lll.lt'llllllllrph~ · have .1 lugh.ly convex
exoskekton 111 wllll h the hc,td in silk \'icw is hcnu-
~pheritJI and the furrow~ .tre ellac<:d. The body
cannot .mlnd.tte with the head in the nonnal way fbl
but dedi ne~ b.tl J.. ward' wrv ~h.1rply from J highly Figure
mdim:d Olllpll.JI 1111~ (t·.g. the illaenine Broch;
pelogi•
HttiiWSt.,idcs) \)uch .1 dNimll\'c morphology is by eau
Figure 11 . 17 Opipeuter, on Ordovicion pelogic trilobite in (ol
poor),_ ~llllt'O to t'plf.ll lll.ll c rawling or swinuning, dorsol and (b) side vtew. (Redrawn from fortey 1974o.l
and 1t i' h•Ahh, prob.1blL· th.tt thc'e unuHJal tnlobtre-s
hved mf.tunally 111 tint•t \llb\tr,ltt'\ and Wt're largely thJt
sedentary 'u'p~m1nn fi..·L·dcr' (Stm. 11)76: llergscrom, clear confirmation of pd,tgH hit· lubtt for gertm
.1 wluch
197J; We,trop. IYtU). Othc..·r trends mdude Ios~ of ~uch a' Optpmter, Cunl/tmtcs .md Cydt>pygr whtcb arr rna~'es
eye~. development of m.ugin.1l spme-s or pitted vaulted, ott.:n ~pinmc Jnd poorly \trc..·.Jmlincd. Surb ttncnt
fnnges. mmt.Hllnl.ltlllll, .md .1daptanon to a dysaer- mlobite~ werl' prob.1bl) ~IO\'- 'wtnuncr\ like dtt' r.~. " l)ltl[
obic h.tbH.lt (olcmmorph,), or to .1 pelag~c mode of mg ampl11pod Hypm.,; thetr murpholo~y lOilfrl~ ~outhe
life. 1 he mtL·rpretanon of 'ome trilobite!> as pelagtc with the remarl..abl> ~trc.>amJineJ tom1 of cyclop}· ~<.'C:lll\
i~ ba~ed upon t!Ul'l' imkpendent critena: funcrional gids !Jke .'\tlllakdltz. wh1lh I\ ellipt" 111 forrn, brh merid,
morpholob'), Hulogy w1th living arthropods, and spmcs, and h:t~ the eye\ flu~h wtth the surlact' the l.ttt
geolog~cal cvllienn• (I" m rey. 1975. 19~5). the long-~nouted hc.HJ Sw.. h ~hape~ have betn Pos"b!)
TnlobHe' \lleh ·" tlw <'<Ju.uon.ll Optpcurer bave expenment.llly te\ted lo1 ~treamlmmg 111 flun 'omc i1
enonnnu~ hypL·rhoph1ed eyt>,, c ilpable of all-round expenments and ~upport the ltk.l of adJpt.ltton w ~"ouditi<
vision .•1nd long tu'lfoml bodJt'\, a-,;Jally vaulted but rapid ~wimm111g. Tnlob1tes like [\hwal.:t'ila were thw h.tvc .ld·
with reduced rhnr.KIL pkur,1e gwing longitudinal adapted for tast movement tn tht' pd.tgic Zl)ttt Jnd Trilol
Rex1bihry (F1g. 11.17). arc likely to have been pred.ltOI'\ Cydopygtds wcrr llH.:h('~ (
The.• latt'rill protik ~how' that there IS no way they the high-l:wtude equivaJeuts of Upipclllrr .md f11unJ in
could havl' rl'Stl·d on tht• ~,.,, floor like other trilo- Ct1rolinircs. l i t ul.Jr ~.
bite\. ThL'\L' tkr.ul~ ''•ggt·~r tllJt the lifestyle was A pelagic mode of hie Ius also bl'c..'n propm,·d tor ()j lht• I
pclagic , and .ltulo~t·' nuy bt· mtde with IJrgr-cycd m:tny agno~t1d tnlolmes (R.obhon. 1972) The~ thu~c of
pclagic cru,wcc.m-; nf the present d.1y belonging to small, blmd, isopygou~ Cambro-Ordovinan form < hlupac
nonm.lly bcnthll ~roup' (e.g. the amphipod have only two thor.tnc segment~ (F1g. ll.l!i •nd (I >t·voni,J
Hypcna) whow L'ycs .md bodic~ are modified m sim- their hmb morphology (Ch,tpter 12) diffel'\ fro, lt I.\ \\
Ilar \V.lY'· Moreover, unlikt· the nomul, largrly rhar of other tnlobttt'~. hnc beez
facic,-C\)n£rolkd hL·mhir .mcmbl.tges, Oprpemc1 The global dJStnbuuon of these Jgno~ttd~ (\\I • 3Dtl flour
~ccm' to hL' indcpl·ndem of t~tcic,, ,1\ ts Curtl/imus. be<>t known 111 the M1ddk Cambnan) ts ,·en· \lil!r fllt.lbk• en
another l.trge-q c..·d Ordm tnan trilohitc. which to a the same spenes occur 111 ~c..andma,;a and 1\l$: !Urc7Imkt
degrl'c 1' homcumorphlt on Opipcuter. Thi~ gi\'C~ a North Amenc.1. This comra,ts dramancallv \\ \tnglt' hed
Trilobite 377
Figure 11. 18 Agnostid (miomerid) ond polymerid trilobites from the Middle Combrion of Utoh: (o) Ptychognoslus otavus (x 7); (b)
Brochyaspidion microps (x 5); (c) Modocia typ1calis (x 2). Agnostids con be used for intercontinental correlation, and may hove been
pelogic; this can be tied in with o stratigraphic scale for the laurentian continent, based upon the benthic polymerids. (Photographs
by courtesy of Or R.A. Robison.)
hat of contempor.mcnu' polymerid trilobites. Ecdysis and ontoge ny (Fig. 11 . 1 9)

whtch rcm.tm emkmic to th1.•1r own continental
masses. Thm the polymt•rid\ of tht' Lmrentlan con- Like all other arthropod,, tnlob1lc~ had to moult
nncnt {at th.1t time tim1ly .1nrhored on the equator) penodtcally m order ro gro'". l"he ca\t shells (exu-
15 quite ditfen:nt !Tom tlur of B:~.lticJ (i<;obred in viae) were thrown off a~ the the amm.1l ecdy~cd.
IOUCht'm latitudes), yet tlw ·'b'110\tids Jrl' the sJme. It bemg commonly d1sarnculared and broken along
seems re.tson.tblc to condutk that while the poly- the cephahc suntrcs during the moulrmg proce~s.
merid, were bentlllc, thl· a!:tllO\tld\ were pdagtc, bur Meanwh1le, the soft ~hell wh1ch h.1d fom1ed below
the lancr do not appt'.tr ,td.tpted tor .1 planknc life. the old exoskeleton was tnfl.lted to ::t larger s1ze and
Possibly they h.td ver> e:-.H:nded larval stages. but hardened. Occasionally the moulted exuv1ae remain
~~>me indeed may have \\\'lint m ,1 p.trttally enrolled mtact or only slightly d1splaccd mto a parttcular and
condition in the upper \\ .w:t\ nf the sca. others mJ.y fairly constant rebriomhtp. M any differeut kinds of
h1ve adopted dtll'crenr mmk' oflifc. such moultmg contigurat1ons have been described
Trilobites were adapted to spcnalizcd ecological and different trilobites had particular techniques for
niches (Thom·~~ and Lam·. I <.>H4) J.nd .tre mually escaping !Tom the old exoskeleton (Ilcnnlng;moen,
found in characteri,til a~soci.ttiom, confined to par- 1975; McNamara :md Rudkin, 19H3; Whittington,
ticular setli mt:ntary t'nvi ronml·nt~ 01 bathymetries. 1990). Somctimes, cspeetally in very fine argillaceous
Of the many \ynccolo!;ic.tl studic~ m.tdc recently, sedm1enrs, a gradational ~ tze ~erie~ can be collected
:ho\c ofHcnry (19H<.>: Ordovin.111), Thomas (1980), which gives a parnal or complete record of onroge-
Chlupac ( 1987) (Siluri.m) and Chlup.K (1983) nenc devdopmenr from the earliest larval stages
'Devonian) .m: rctomrnt:ndl'll tor lttrthcr readmg. (Whittingron, 1957). The first omogenetic series
It is worrh nonng that 'opporruni~ttc' trilobites kl10wn were described by Barrande 111 1852; one of
.uve been rccordt:d. tht'\l' ,tre \pecit'\ able to invade these, the Cambnan Stw hirsuttl, 1s tllustrated here
llld Rounsh 111 rc,ourt.t.·-rich but otherwi~e inho'- (Fig. 11. 19a) and ~haws the general sv~tem of growth.
~ttablc envtronmcnt,. Thl')' arl' mually recognized The e.1rliesr ~rage IS a p rotaspis. usually some
1 Brezinsk.J, l9H5) bv uumu.tl .tbundancc \\ithin a 0.75 mm m dl.lmcter. Th1s 1s usually a cambered
ngk bed di~c. open vcnrrally. wl111:h carrie~ a segmented
378 Arthropods
~
<"">.
&~ \ ~---~~
lJj
(a) Sao
"
\:
2
- _,.-.../
3
~if:--
4
~-
6
(d)
Fig
(~
\--~-,;t ,,
~
~;
~
~ ~
"\~&;~
(2,
'\ i) -. . : ;;• the
Ore
~J·- .~
~~ .v.·:-..: fron
2 3 4 5
(b) Shumardia
ccn
ar tl
mar
fae~.
kno
ll.l
de se
tlon.
1she!
spine
Ole1
7 6 11.1 '
hkc :
of tu
arc q
As
trans'
CL' ph;
two c
m era
tree. ·
growt
actual
4 they a
rior p.
(c) 0/ene//us tory I
1dcnrit
reJcase
2, 3,.
mcnts
Trilobito 379
101 asaphid
frgure 11.19 Trilobite ontogeny· (a) Soo hirsuto (M Cam.), Bohemia, showing prolospides (1-3; x 14); merospides
(4, 5, >< 10) of
degrees 0 and 6, and holospis (6; x 1 5), (b) Shumardia (Conophrys) solopiensis (L. Ord), Shropshire, England:
( 1-6) meraspides
(degrees o-5; x 30), showing the development of the macropleurol spine; \7) holaspis (x 15); (c) Olenellus gilberti,
(1) early meraspis
12, 3; x 8); intermediate stages (.><: 6, x 3); (4) odult cepholon (x 0.5). The orvol (intergenol) spines are a lmost completely
i-t adult ond ore not homolgous ....nth the genal spines (5) (d) lateral, anterior and ventral views of the late reduced in
OrdoY1tion osophid, show1ng the spiny hyposlorne (x 40). ((a) Aher Borronde in Treatise oo Invertebrate Poleontology prolospis of an
; (b) redrawn
tom fortey and Owens, 1991; (c) redrawn from Pal mer, 1957; (d) redrawn from Evitt, 1961.]
entrallobl•, l,tter to become the glabel1,1. The eye~ transitory py~idium. The process is perhaps best
,. tillS st:lgl' arc tiny .md are located on the :mtenor seen m the developme nt of Sl111mardia (Cmwpl1rys)
~'gm; l.tter they migrate mward~ bnnging the salaplt'IISIS (Fig. IJ.l9b) from the Trcmadoe~an of
~1al suture wirh them. The hypO\LOllte is not Shropshire , England, since the macropleu ral spines
~nown in SM. but in genera wch .lS Asapftus (Fig. on the fourth thoracic segment arc a good marker.
l 19d) .md Grc11'1rtll)'llll'lll', tn wh1ch H has bl'l'll The~c are liberated at merasp1d degree four whik
Scribed, lt ~~ extremely ~Pill\ . ~ugge,nng a funt- the fit'i:h and Sl'<th segmt•nrs are still bemg fonned at
l)llal dlJnge a~ the hypmtom,l l spillC>\llY dimm- rhc front of the tr:ms1tory pyg1dium. Meanwhil e the
llhe<i. M.my protaspide ' arc generally spinv. and thL cephalon is acquiring adult proport1o11S.
f!ues nl.l) later d1,appear totally in the adult. When the adult number of thoracic segments has
' ~(nclhd tnlobites. tor lrutanct· (Palmer, 1957; F1g. been reached the trilobite is now a holapsis, though
I.IIJc). have prota\piJe , \\'llh .1 pair or
long bladc- it may have to pass through m.my more mouJts
l< 'pmcs. whose adult cqUtvalenr j, merely a pa1r before it 1s of fully aduh propomoll S. Rardy, as 111
. :my knobs ly1ng wHhm the genal sp111C\, which A11fnrople11ra wluch had up to 25 rhorac1c segment!>,
• quite ~cp.1rate orgJns dcvclopmg hll'r. new segments .ne added until ,, very bte stage 111
As the protaspidc~ grO\\ by further moult' a developme nt.
:~Sverse fi1rrow develop~. 'cparntm~ the larval \omc cunous pho~ph.1tizcd obJects found 111 a_~so
. ~halon from the pre~umpnve pv~1d1um; tht• ci<H1on with cybclimnd trilobites and smaller than
o can freely articulate aga1mt one another. The the earliest protasptdes have been interpreted as pn·-
Dltraspid ~tage begim when the pygidmm become\ protaspid larval stages (Fortcy and Mom~. 1978) .
.. The thoracic 'eg111ent' then tonn 111 a zone of These phasclus larvae arc ovmd, vaulted bodies
1wth .llong the front of the pygi.dium. They ,1re with .1 ventral free edge and doublurc.
uilly part of the PYbridium for .1 while .•md then Exquisitely preserved details of ~tructure in trilo-
arc rdea~ed in rum and liberated from its antc- bitl larval St.lges are now known fTom ~1 1tciticd
t part. the pygidlllm is thu' known as a transi - speCimens wh1ch has brrcatly added to om knowl-
IOrypygidium as it do<.·~ nor .ltquirt· 1t~ permanl'IH edge (e.g. Chattertou , 19RO). Two basic kinds of
'1oty until the 1,1\t thor.ltH. segment has been prot;l\pides have been di~cint,rul~hed (Spl·yer and
~-ed Mer,lspidc' arL numbered in del.'TCl'S - 0, I, Chatterton , 1989), h1ghly inflated 'non-tnlob itc-
3, ... - .JCcordmg w how tnJny thoraciC ~eg likc' fonns such as rh me of AstlJIII/Is (Fig. 11.19d),
nts have been freed from the .lnterior edge of the and the more typical 'tnlobite-l ikc' cambered disc'
380 Arthropods
(hg 11.1 1J.1) J\ 111 S,l<l. Th~: tom1er Jre believed to C lassification (Fig. 1 1 . 20)
be pbnktK, the l.ttter hcnthic.
QllltC ap.trt ti·om thL'II imnm1c intert~~t. ontoge- The d.lmiit.ltll)ll gwcn be-low ''a moditicd versJon
mc~ arc porenn.11ly li't ful in 'ug._~e\ting phylogenetic
of char 111 the ti)rthulllllng \econd edition of the
rcl.momh1p\ hL·twccn e,t,1bhshed r.rxa. If two dis- Trc•lllsc 1111 hu•t•Hc/mllt fJ.Ift'tlllftlft\\!)'. lr is based upon
uncr 'wck~ h.wt• t'Jirly 'mubr protJsp•des, d1tlering the whole compkx of ax1al .1nd other charactl.'l'\,
wuhout any 0111.' or m on bcmg ,jngkd our ;1.\ bcmg
m 'h.tracter\ tTtllll rho,e of other ~rock~. they ntJy
~h.lrC J COIIIIIHlll ,11\l'l'\try 1.'\'t'll 1f the adults look
of nvcmdmg •mport.llll't' Virtually all rhe character;
qu1rc dilll:renr Thu' whl.'n the L1ntog~:ny of thl.' U\l'd by thl' t'.lrht•r \)''>tl'lll.ltl\t'i Jrt· hl'rcin incorpo-
cory·nL·xodllll rn lob m Rlfh)•Hrisws Jimhriatus wa~ rarl'd, but 111 .1 morc re.tlt,ti( pel'\pl'ctJve. In addi-
duodatL·d (Robt,on, I1)117), 1t beeami.' ckar rh.u' tion, dl.lr.Ktl.'f\ xuch ,1\ rt'SI.'mbl.mcc~ at earl)
ontogl'neric xt,Jgl'' bt•rnmt· me rc.lstngly 1mportJnt
there wert• 'und tn£Jl'' betwt•en it' protaspides
Thl' duractcr .llld lm.ltiOII tltt' hypostome Jic nm\
.md the.: prot,l\ptdL'' of m my ptychop:mids.
Furtlll.'nnorL', .1dulr cOIJIII.'XOdllds show m.tny char- '>l'en bl.'ing :1\ of p.1rtaul.1r 1mport.mcc for h1ght r
.Ktcrs th.1r .1re tound 111 thL' prnt.l'>pid and mt'f.!Spld kvd cLI'i\itil;ltion (Fortt·y. I 'J1Jil) .
stage~ o! ptyt hnp.1md~ .•1nd ~o it has bl'l.'n possible to CLASS TRILOBITA (C.1m Jlmn.): M.mnc .mhropod' w11h
cst.tbh~h .1 n:.il phyktit: rdationship between the IJr~,;elv t.aluut <'Xt>,J..clcton div1d.:d 1nt11 tlun· km~ludHI.d
two ~rroups, ,1\ wdl .1s showing thJt major differ- lohL·~ aud with ,1 <.hstm ct ccph:11on .•lrtu: ui.Jtl·d tbor~x ,nut
cnn:~ 111 thL· hol.1~piJ ntorplwlogy of the two groups PYh'lllium. CL•pll.llun Wllh primiuvcly furrowl•d ~ilbdiA
t:u;J.tJ 'lilUll'S, llllllfJ\llllld C)i<'' (S'llUCtlllll'\ )mt) ,IJld JouJ>hU<'
I1Jw n: ... ultt·d m.unly trLllll thfli:rl.'ntJal growth rate,.
\\ rth frl'l· t>r .ltt.lt hnl hypu,IUII\l Fn.>m 2 tu 40 thorJCit \cg-
Ontogl'lllt'' 111 tnlobJlc' .tho pomt to the impor- rtlt'nt'· l'.ll h wuh ·"'~ ;ul\l plcur .1 tJygrdaun vanJ.blc 10 ~z,
tanc~ of p.1cdomorphos1' J\ an Important factor m .md sll.lpl'. o;;p1nm1l'\ v.m.lhl, Vt'illr.ll ipp<'miAg,.,, .<pJn &om
evolutJtm. l:\peti.lll) where JH.'W kind!\ of organiza- unir.1mnus .lnf<•nn.ll , .Ill' <llll' p.ur p1·r w~'llWnt. b1r.unnw. ;u,d
tiOn 'eem w h.1n• .lml'n vel) r.tpidly and \\'llhout all of rh I." ~.llllt' J..md de, H'J'IIl!!. 111 \l7l' p•htt norly.
prewrvcd Hltl.'rmt•dJ.lte~ (Ml Natmr.t. 1978. 1983, ORDER 1 REDUCHIIDA (L.- M <. .un ) An carh gru11p tll
Wluttint-rton I<JH I) I hu' the \Uturc of trilobite~ nilohlfe" wirh .1 largl' semJnrcuiJr ct'pbJion luvmg mon~
genal ~pme~. numt•rous .1nd usu.1lh· 'l'lllV rhurJ.uc •Cl;·
can be tr.tn d eh rough ontugl·ny a:, uuually prop.u-
mcnr- mJ J nny py~dmm . l·vc' brgc, hypo,tomC' C•'ll!et·
Jan. though 1t ll\.1) l.ner become opisthopanan. m mint. In< ludc, \UIIl>RJ>LR~ I 01 EN El LOIOEA
Smle thc1e '' gt.llld evidence that ph:u.:optd (nu do".tl \Uturn). ~ ~ {)/cud/rh 2. I:.MUElLOilll:::l\,
sch1zochroal C)'l'~ .tro\1.' paedomorphic.ill). it i:. not "·K Emudk ·' RLL>LI<. liOIDl:A. e.g. Rt•,l/r.lua. 4.
unlikely rh.1t the prnpJrl.ln 'urure of the Phacopma PAJlADOXIDOIDEA. t .J.: t'.H.l.ltwi.ft•> - doNl !llCur~
is hkew1~e paL·dmnorphtt prc\cnt
ORDER 2 AGNOSTIDA (I .. C.un.-U Ord.): !>mall tnlot.m1
HO\\ cnlobltl'' reproduced IS unclear. The: occ.l-
with subt>yu.tl tt·ph.,Jon ,111J J'}t,;ldlllm. ll'llall) blinJ
sion:tl dl'>lOVl ry. ho\n:ver. of monmpc:cific. age- ,u[Uro:ll'ss. t'nrollmcnt t~pK.Ii. Thnr.l< 1c "gme111' nurnl>cr
m reed clustered .1~'-cmbLlge' (~pever and llrctt, nnl\ twu (SUTIOROCR. I. AGNOSTINA Itwo 'uht:uu·
19H7) su~ge~r,, :lS \\ 1th \CH11C modem arthropods, ihes]) or thret• (::.UllOK.DER .2, EODISCINA) !three
rhar rnlob1tcs .1ggrcg.ltt'd together prior to moulting ~ubfanuhe~ inrludin~ J>AGCTIIDA.I:; the ouly group '''
Jnd 111::1s~ copul,ltum h,IVe t'Yt'~ .111d sutttr~'l llrpnstnme natJnt. with rihhon·
Stze-frequcnry .malysis of trilobite populations hkc wmp;s; c .p.. ll,~nmtw . Ji,>dism<. Pagt•tict.
ORDER 3 NARAOIIDA (M. < ',un) Um.•lu~ied tnloh1tc·~
h.1w revc:~lcd diflcrcnt reproductive strategies. Thm
WHh no rhnrJCII scgmt·ut~; l'.g . .'\'ar<t<lia. Trg•'JWitt .
r-strJteg1srs h;wc high tccundiry and substantial ORDER 4 CORYNEXOCHIDA (I C.m1 -M Dev.): A nthrr
juvenile morr.1lity. the popuLitmn curve is lcft- hetrrogcnt•om group. <.ibbdiJ ot \'lflrd lonn bu1 usualh
skewed. K'--.rr.Hq~tsts on the other hand, such as pJrJIJl'l Sided or l'XpJnJmg .mrenorlv: suture~ opl\thop.rr·
O~)'l?lllf<llt'l/,,, .m mh.lhltJIIt of t::urly dl·cp water, h:lVe un \Vith \ubpar.JIIcl Jntcnor br.mclll"', thorax mualh \\'llll
broad, flu di~rnburion cun'C~ (Shcldon, 1988). 'even \Jr t:r~ht w~ncnt,, oti.:n 1~opygou,, Hvpo>tom<
t:lllllt:nlllll.lllt IO IIIIJ'Clldl'lll, llldy bl· lliWd With fll'niJ
Here. ti:cundity. and hcnn· rccnutmcnr. is low,
plm·. Jndud,.., SUDORIJ£RS l. COR YNl:.XOLHIINA.
small individu.11, nt'VL'r tom11.'d more than a small
\\ 1rh contanun.ull h\ pmlomc: (t•.g. 0/trr,,,J,., Z.'l(.t•ll/,.
pan of rhc tot.ll f.nuu. 1nd the popul:mon wa.~ main- &llh)'uri.«r•~} • .2 <;( UTELLUI~A \\Jth impcndcr:t hyJll~
tainl.'d in .1 \{C,lll)' \{,He '\uch srratq~ie' were proba- lOillt' (t• g. '\,uttl/ulll. 1/l,t(ltftr, 8111/loUIIH, f'/111/1/•!Jri<~IJ). 3
bly ch:uacreri,ric of p.lrtltul.lr l.'nvironmcnts. LEIO(\TEGIINA (q;. C/"""(1:'•')·
JW~ (/)
51
a::
Upper
I I 11 I I I
~ ~----
~
17
le --'
"'r~ '- -~1~ .
~ Upper I \~ 9
z Middle
~0
~ --.
w 5 13 ~
-
OL
c Lowe<
')
15
I !11& . .•
• -! '2
_./. ~0 i- ~>
~~
14 ~ ~
1 t 0
-.;
0. c:
·=
1-
£
•
tO
1~~.0
~
5101 I
!Iil
Upfler
zi Mlddle 16- .
~
a::
Q)
u
~I towe< ::i5§ •
57
I
Redliclllida AQnostida PNcoptda Ptyct>opa riida Auphkla Proetida
UBRISTOMA
Figure 11.20 Time ranges of orders (e.g. Phacopido) and suborders (e.g. Calymenino) of trilobites Genera illustrated ore os follows: 1 PoecJumios (L Com.l, 2, -;
~-
Porodoxides (M. Cam.); 3, Eoc/iscus (L Cam.); 4, Agnoslus (U. Cam.). 5, Scutellum (L Dev.), 6, Kootenia (M. Cam.); 7, Dicronurus (Dev.); 8, Trochurus (U. Ord.-M. Sil.), 0
9, Cheirurus (U. Ord.-M. Sil.); 10, Ca/ymene (Sii-Dev.); 11, Acasle (Sil.), 12, Olenus (U. Cam.), 13, Harpes (M. Dev.); 14, Ogygiocaris (M. Ord ), 15, Cydopyge (Ord ), e-
16, Trinucleus (Ord.); 17, Paladin (l. Carb.). a-·
(,)
-
00
382 Arthropods
ORDER 5 UCHIDA (M . C.uu.-M. De, ,): u,utlh 'pim llARPINA havt· pnlmlunct"d cephaht fnn~t·' · c ~
rnlobltc:<i. With , nntt•nmu.mt hy1'mtvmn. \uperl:tnu!ic, of
HIIIJIC.i.
1 I H. HOIDEA (I Ord -U Dev.), medllllll-\IZed t0 ORDER 8. ASAPHIDA (U. C..tm.-Sil.): Mcdt.Ln vcmr I iry
very large rrilobue~. \\ tth uum t,takablv dl'tmrnve cephab suture present. only pnmtt1vc tomlS rewm rt.tt.ln\ ,·nnllt-
3nd pv~dta ( cplul,,n \\tth broaJ gl.tb<ll.t, •>ften with uom. mosr bcmg contcrnun.mt or impendC'nt. llt~hcr
fitsed bter.ll UJd ~ltlwlbr lobi."'>, upi,th"p;Jn.lll surur~. ,1\Jptmb h.IVc .1 dbtmcuvc mtbtcd prot,t'Pl' (f'nrtt'\' lutl
Pyg~dturn oftl.'n l.tr~cr th.m cepllll<'ll; tuhl.'rcui:HC.' Cluncrron. 191:11:1). SUPER.FAMILli:.S I J\N0\10- rea
exo,k<.'leton (e .g. l ,,/~ot•. Hrm1J~~t<. Trr.Jio1•p1.>); ' CAROIDEA (e g. ,-!uNIINolris). ::!. ASAI'HOII H!.A (C.' g rea
ODONTOI'LEUlU 'IDEA (U. CJm .-M De" ) Vcrv t l'•'flltu,, Ct"''ai<'PY.!J<'), 1 DLKELOC.EPIIALOli> M
~pmy tnlob1te~ G!Jbdla Wlth three p.uf\ ot l.m·r:~l lobe\,
the
(t' g Prytl111spi,); -1. fU:.MOPLEUIUUOIUL A (c!!
Ollb tdt whJCh on tht hbn~,:cn.H· he thl' m ubr ndge~. Rt•m••ph•,cridcs); 5. CYCLOPYGOIDI:A (e.g. ( r•'•'P)'C<', the
~utures upisthop.ntilll, hypmrome snull Short spiny · Ni/~t11): 6. TR.INUCLEOIOEA (e.g. Trimcr/c'IH, Dtomcl, , bei
pyg1thum. lUbl•r.:ul,H< or 'PIIIY cxmkdctun (c.g 4ddasJ>iS. l'llllflY·'· Or<>lllt'ltlpm} l'VC
L"''"'sl'i>, Ccr.llottp!J,,/,1): 3. DMlESl:LLt>II>I:.A (M.-U. ORDER 9 PROEllDA (Oni-Pcnn.): GIJb~U.t l~rgt .md ll.lt<
Cam.): East A,wt. mu.tlh >pmv tnlob1te,, prnb.1bly 'l'ter vaulted. well defmC'd, u~11.1ll) \\ ith ~enal 'Ptll<:~. at:trr.:>\1 rep
!-'TOll{' to oduntoplt·ur.uJ, and h.lCI.."wudl} Wpt'nng rmrral plarc. opl\thup.tnJn e~t1
ORDER 6. PHACOPIDA (L Clrd.-U. lk\ .): L.1r~..- order of
On
hnlochroal and u'ually l.arg<". long hypo~tnmc Tlwru
domm.mtly prop.m.nt trilobirt·'· Jindt·d inw three clearly wtrh ctght to Ill St'f:lllCnts, t\npygous: pyguiJuu• mu.allr
Tm
rletint"d \ubordcl" turrowcd and not spmy (Fortcy ;md 0wt"m, I \175). div(
SUBORDER 1. CHEIRURINA (L. Ortl. M Dn .). A very l>tstltll'twc proetotd pmt:.lspis. Two SUP.I::RI·AMH IES. HI I'\
v.mable, nnmt.~lly prnpm.u1 gT\.lup. GIJ!-..!IJ \.1-ith up tu I I'ROI:TOIIJEA; .:!. BATIIYUROIDEA (C'.g Prc•·lt" lllte
four patrs of ~I.Jiwl1.1r titrrows .md li\U.tlly .-xpandUJj; fllu/!lpmr, 8atllytmH, .-lu/,t.-,>plclllll). thi\
tonnrd~. ~null h11lm hru.1! e-yC'I. roMrtl pl.ltl' pre~l'nt.
~.·.1rl>
h\ po<>tome- tree. I hM.I' wirh etglu tn 14 wgn1enb.
Py~dtum UStl.lllv lnbcd or 'PIIIY, C' ~. Clrt'mlnt> ,
urga
Evo lution
D"'l'lwn, l.t·r.wru<, 'rl~o~nr.wdrus. v. .ls
SUBORDER 2 CALYMENINA (L. Ord -M Dt·v.): A fairly General pattern of evolution dcvl
homu~t'llCOU\, ll\ll,tlly Ji;Oil,tropartJil WOUf' ,·haracter- AJmost nothing i~ known nbour the .mn·~tor; nt
ized by an mtcrmrly l.lpcrmg gbbdi.J wtth four nr tive
lriloblle'>. In common wtth other Jrthropod~ 1t Ulll'
pain of Luer.ll lnbt·' c.lunumlung m 'Ill.' lorwJrd~. Eyes
'm~L! holoduu.tl Thurax with I 1- 13 'cgn1cnt'
be pre,umcd that mlobttc~ were po,,1bly denved
from thl' <..1me ancestor-. J\ rhe annelid\, .md thl' Ill'\\
PygtdJUm rouudt.",l vr \l.IOtnJn~:uiJr In FAI'v1ll Y
HOMAIONOTII >A the .1..xis IS very broJd .md the fur- EJlatara ·~ofi-bod1ed en lobar~'. recent!\ de~t nbrd ble t
row' Jrc lat~el) t'll.h et! ot Vt'lY ,Jullm\, c ~· t...Tiymmt' , (p. 60, Fig. 3.lj). may have been ~omewhcrl'lllthc wnc
Tnmt'ntS. ante'>tral hne, but thi~ "only '>peculatiOn. I he lJrl te1:f 1
SUBORDER 3 . PHACOPINA (L. Ord U. Dcv.). o thc1
e~t tnlobicc b'l'Oup to appt·ar is Suborder Okm·llm.1,
Ccph.tlon w11h lnm.1rdly t•xpandlll~ w.tb(·U.t, schizo- \llllll.
clun.J..I cyC'I, pmp.m.m \lttUrt'' .md tw rmt ul plate
wh1lh is micropygm". These. together w1th the
~ewndarily blind Agno,tid.t, which Jppe.1red .1 ltttlc I tu!lt
Thorax of 11 •c-gmcnt\, SUPERFAMIL Y PHACO-
later. dm:rsificd through the Lower CJmbmn and lll'UU
PAC EA h.t\ ·• '111311 PH,dtum Jnd "C'll dt·\·doJX'd
cnrullmcm mcdum~m, , Mcmbl.'~" niSUPF.R.FAMil. Y were jo111cd by the C nryncxochidJ and Prycho- pun·
DALMANITIDAc trt' larger. tmpvgou, .md l.ug<"-<"}'l.'d p<tmdJ, which continued through after the dcnuse ,lprq
.md have rnrollfl1(11l stnh:ntrt"S ponrh de\'c:lnped: e-.g. ot the Olcndlina at the l·nd of the Lowt·r C 1mbn.m. lt~hll•
Plwwps. Vclllll.lllfll'.l, lrol\h', C/urslll•'l" The Upper Cambrian t:llln:J is dominJtt•d rn.unly br dtcd
SUBCLASS LIBRISTOMATA (lwtcy, 1~.1411)· lndmk-., .111 trilo- pted 1
bttc' wtth uat.uH hvpmtnmc~. together wtrh ~t·, onJ.mly con- the Pryt hopariina whid1 \how, :lt h~a't supt•rtiu.lllv,
rdati vdy limited variation, by contr.t't Wtth th1c the (
tcmunam or uupcndcm trtlohne, w1th tl.l!Jilt .uKc,wr;
t.':\hibtted by maJl} OrdoVlnan t.lXJ. 1 hi~ LOllld lll.lll}
ORDER 7 PTYCHOPAIIIIDA (l. Cun.- U . l>c\'.). A l.uge.
p.uaphvlctJc tnlobnc urd.:r. lll \\1th tUtanr hvpmtomt'<> pomt etther to a pcnod of cvolunonary sta~l.ltion 10 bcun
.md tncludmg 11\.lll~ htt-:hlY modtht·d !-'lOUP' 1 yprCJI ' \\.hlth there was both !muted genetic pOtl!lUal anJ \Ur\IV
prychopJroid' .,u, h ·" ~UDORDER I lll I:..NIDA hJve low selection prcs,ure, or .dtcmativdy to J mhcr (ul .u
.1 sunpll-. ti.1n\.mlly t.tpt'rlllK ~bbt·U.L wHh nonnally Aluup•
n~orml\ sdection wn·ding our of all but the mm-
strnight glabelbr lintnw,, brgc thorax Jnd ,m,lll pyg~dium to t tht
pholo~'lcally defined fonm. The forml'r .1lrcmJUVt
(e.g. 0/nrm. L.JII•'I'f,,,,.,.,, Tn,mlt~~c~) JSUI'f'RF-AMILLES
1' f.JVourcd. since in tht• few group' m whtlh evoh. p.lrltCl
I ELLIPSOC fPliAI OIDEA .mJ 2 P I'YCHO-
PAR.IOIDEA .1l,o nd•>n~t here (~ g. Hlc1 "'"I'I'J/u.>. 11011 ha' been ~rudit.d m detatl (e.g. the Ukt dJr li1Jl Jt
Pryclu,p.mil, r:,,,,~,,l')'l'fc,•, \lwm,~r.lrll) l ~UUt)ROER. :2. thcrt• is. 111 spite of tht• retention of a standard nwdel hJd pJ
,IJj( .lth
Trilobita 383
of organiZJtion ..1 t:11r dq,,rrn· of cvolutwna ry plastic- The tnlobttc faunJS of tht· Silunan .md De\'OiliJn
ity withm the group. tuntain many of the same dcments: plucoptd,,
At the t•nd ol the C.1111bn.tn there W<lS a major dJimanitid~. cheirurid~. calymcnids , h;lrpids, ctt.
ms1' 111 trilobJt( lmtory. Mo~t of chc compar.mv el) Consequen tly a Stlunan as'ernblage can easih be
unspeuahn ·d Upp~·r C.1mbn.1n o;tod.:., died our. fe,\ tOnlu~ed. at least supcrticiall y, \\lth a Dn·onian unc.
reJeh1ng the Ord(WlCI.l!l (Whmiugt on, I 966). The Encrinurid ae (S·uborder Chcirurin.1), however,
reasom fi>r such cxonct10II arc unJ..nown; perhaps though ~o rh::tractenstlc of many Siluri.m fiumas, du
du.• nunnc rq,rrC\\1011<. ot l.uc C.1mbnan nme and not extend tar into rhc Oevo111.1n.
the me ut pn:darnry cc:ph.1lopods m.l'f bt: invoked as The cxnncttom of tbc- Mtddle ,md Uppe-r
bemg .1nwng~t the nu$c~. Some ~hort-ltved, rapidly Devonian disposed of tht majority of trilobite
evolving Upper 'I rcm,1dotta n group-; then origi- wuup~. y~·r the rcmJlllmg Proctitb Wl'IH .Ill the \\ I)
nated, .md after thcu c'tmcuou t\Jme rht· tlrst through to the Upper Penman. Most of rhe~e. e\pe-
r~pre<.entarivcs of the 1mportJnt .1nd dommant ua lly thmc ltving 111 shallow water<., wt·rc lar~e
Ordovi~:ian brroup~: the lllaeuin,l, Ph.1copida, eyed, robust genera capable of enrollmen t, but there
Tnuudcma and other'\. all h1ghl) thlfacntiat t'd and wen~ also <>peo,1hzed. thin-~helled, hghtly ton-
dJwr..c, mmt of them oft rvprogcnc uc ongm, and 'tmcted Jud often eyeless form<;, e~pect,11ly m the
\Umvin~ for v.1nom penods thcrc.ttlcr. lt 1s an deep wat~:r nlthc- Variscan told belt innorthwe~rern
mtere~ting featun· of tnlublte evolutum th.H after Emopt. Thl'~e Jrc found from the Upper
this grc.u bur<.t of new comtntctio nal them~:' m the l)evoman, ''here n·duced-ey ed proetld\ and plu-
e.rrly Ordovici.m very le\\ entirely Ill'\" p:merns of topids art· pre~ent, ,111d through the Carhonife mm
orgaru~atton ,\ro\t' ••Jtter<.v.trJs evolunon in trilobites m the Culn 1 f.tcie~.
\''ob ver; largd>· .1 matter of expenment<~tiun and ExtinctJon of the last Penman tnlohm:~ w;t~
dcwlopn~t:nt of the ne\\ ~t·netic nuterial th,lt tlNt probably rdated rn the lowering uf 'ea kvel
.::une mt1> being m the e.uly Ordm tu.m. that contempor am·ously :ltfened so nuny other
lt seem~ clear that thc~c 'trong ami do m in mt new uwertebrJte~.
Ordovtuan ~tod-., \H're .1bk· to colomze and t"'<plolt In gc:uerJI, ~omc furly dear evolullmlJ ry trend~
new cnv1rOJm1ent' 111 a w.1) dut lud not been pos~i tan be distinguished (Fortey and Owen~. 1990J,h).
hl<.' for thc:ir C:.unhrian progenitor~. Fur the first such as the origin of new eye types, the improvem ent
ume. tor exan1plc, thtrt: lrl' many g\:naa colonizing nf superior enrollmenr and artiCulating mcclum'm '.
rttf em·tronrnc llt,, 'omc pcNistmg ii.1r a long nme, the change from nurropygy to 1sopygy. the dcvdop-
orhef'\ being repl.u:ed by dtfTerc:nt gl'ner.l of bro.1dly ment of extreme ~pmosiry in certain groups and the
\imiiJr morpholog y. fhc~t· reef faunJs are often dif- reduction in the ro\tr;Jl pbte. Yet trilobite~ as a whole
rirult ro correl.ltc 'traugr:~plm:ally \\ 1th comempor a- remained constntrtcd on the s<~me an.hcrypal pl.m
nrom t:luna~ lnilng in oth~:r en\ lroimlent,. The defined in the earlil.''>t Cambnan and, especially .tfter
pure lune~tolle faun,!,\ of t•arly Ordov~~:ian a~e have the early Ordovician . the number of change~ of n·;tl
l preponderanc~· of brge. nther flattened and rather \tgnificanu : remained mrprismgly low.
ti;hcly fUrrowed a\aphtd,, ii.1r mstJnu~· wheu these
ru~d out. !>CUtdluld'> of ~Jmilar morpholog y OCCU- Microevolution
pied their ecologital nicht•. Desptt<. the su~o:te'' of There have been m,my studit·' on tnlob1tc evolution
I< Urdovinan tnlobttt'' a' a whole, however, (~unm1Jnled tn Eldrcdge. 1977). ~peuJll) on Upper
'lUny dmmct group~ (q~. Famtl) Trinutkld ae) Cambrian, OrdO\Jtian , Devonian .111d Lower
J~c~mc extinct by tht• end nf rhc OrdovKI..tl l. The Carbonifer ous gener.t; ol these only three ::tre sekctt:d
,ur\WOf'i were lon~-rangt•J .md pre,tnnably 'uccc~s for ttmher di~cusston. Other c.1se histone' are don•-
ful and versattk gruup' V et tht•,e 'wcll-tncd ' memed m l.Llrkson ( 19R8).
J;J'OUP' were not devotd ot d!Ver<.tlit.ttlon potential,
tor there is evid~·lll<." of~t'Vl.'ral adaptiVe: radJatiuns in 0/enidoe
p.unwbr i'olated Jrea.s ut tht· world where an urig- fnlobites of Famil} Olcnid.lt' occur 111 V:l~t numbt f '
11.11 nce,tor uw.tdm~ a !Jtrly empt", environme nt m the Upper Cambri.ln of ~cmdinavi,1 and al~o 111
.1ad p.tvcd the w;ly for ,1 bur~t of evolut10n.1ry thvcr- Britain. North America and Argentma. They h.tvc
llfionun m its Je\ccnd.tnt \. been stud1cd in grc,Jt detail ;md are of ~rreat strati-
384 Arthropods
~'T.1ph1cal valuc. I'hc1r cvoluoon ~p.lns over 40 Uppc:r Cambnan zoues 111 ~candinavia. 0/1'1111.1 g.tw \l'l
1111lhon years, ntenJm~ 111to the Ordovician. rise to Lcptoplastinae and JJelturinae wh1ch prohfcr- 'P'
Ht.•
The 'olcmd sc,1' ll.:.~rht.·d from Ea\tt.•rn Canada to atcJ and became very diverse in the Slll"Ct'eding
Sc.mclmavia. It .1ppc.m to have been l11rly shallow, zones, along with ~ome denved Olenmac. 1\omr n(
but may have been !Jr~dy stagnant ne.1r the bottom gener.1, such as the leptopla~rine Ctt'llOJI)~~t. b~(.llllC oft
or JU\t below thc ~t.·dunent surface .md the olenids very ~pmy. There was a rc!Jtwely rapid tumovcr u1 be
\eern to have been .ldo~ptcd to dysat.'rob1c environ- olcmd genera and ~prCie~. mmt of whkh Jrc of dlJ
ments. ~ome of them m.1y have been ,ldapted eo grc.Jt str,ltigraphical v.lhlt.'. At the end of Upper llm
bottom-dwclim~ ,\1\d tC:edmg off tht• nutrient-rich Cambri.1n time the Oknid Sea contracted, and then li l t!
tlocculcnt layer on the ~c.l floor. c,x:p.111ded in the Trcm:~docian. The oh:111d~ of sta~
The overall p.!llt'rn of olemd evolution in Tremadocian time were :1gain quite v:wi.1ble, .md it fr01
S(Jndmavia h.1~ been splendJdly dnrumcnted by was at thi~ time that Subfamily Triarthnnac ;trmr oft
We~terg-lrd (1912) md Henningsmocn (1957), who wluch became dommant in the early Ordoviuan m g-ra1
deal wHh Upper <.. ,1mbnan Swed1~h .md Norwegian North Amcric:J.. Subfam1ly BalnibJrbnd.tc. on the grc•
t'Junas. respecnvdy Forte} (1974b) lm dt.:scribed a other hand, wh1ch .uc known onh. from lm11
nrh Lower Ordov1n.m olcnid fauna from Spitz- Spitzbergcn, evolved rap1dly in virtual ~~olauon m 1.'rs .
berp;cn and Clarkson (1973) clc:1lt with eye mor- the early Ordovici:m, a product of a n:n1c1rkable t'.lg<
phology. A number of olcnid subbmilic' have been bmst of evolutionary activity. Only a tC:w pdturnws thJt
di~ungmshed. of wh1ch the most 1mportant five arc and triarthrines and membcrs of smaller olcmd sub- WCI
1llmtrated here (I 1g 11.21). fanuhcs snll renuined ar th1s time, suborJmate ro t'vol
Subfauuly Olcmn.lc '" the carlit.,t and its type the U,tlmbarbiidae. Man} studieS ha\ c been nude ot the
~c:nm 0/et/11.1 dominate:\ the lnwe\t two (of ~ix) the o,·cra.JI pattem of evolution in uleni1.h, .UJd a thro
nf t."
~tor!
m or
not
tht• (
J,w
whJI
trom
el\ev
grou
I' Jll
S ubfamily Olanlnae Subfamily laptoplestlnu Subfamily Pelturinaa
0/ttnus trunu tus Lttptoplttstus stenotus Pttlturtt scerttbllttOidtn \tudi
treno
Ph
f' Id re
Dcvc
d.I~SII
ltllltl
M1dd
dt.·nvc
!fr,lm.
P. im,
IL's, 111
wry
Sablemlly Balnibarbiinae Subfamily Triarthrlnaa rh.mg
Bttlmbttrbl pulvintt• Trisrthrus BBtoni
much
Figure 11.21 The trilobite fomily Olenidoe representative genera and species of each of the five main subfomilies r.lll,l, (
Trilobito 385
very nnportant cnntnhutt(' lll to our knowledge · of These ~ub,pccico; an: nor vcr, eaw to J1~tlll~'lmh.
'peciatlon in fm,!l-. c.tmc· (mm K.tulnl.lnn (I 1J33). rhetr ddimtmn bell\g mainly ba~ed on eye mor-
He made J ~tati,tiol ~tudy ot· Ulc11w Ill a 1.5 m p:trt pholo!-,ry. Mid ;\Jl cxrcm1Ve multtv;lriatc :ln;llysis w.l\
of ,1 condensed \l'liUerKc' 01 one of tht• lower zones ll'~·d m dulidation of rh1s h1storv. Eldrcdgc\ study
of the Upp.:r Camhnan. 1 ht pyg.tdiJ wen: found ro 'panned a l;~rgc .1rca of easrem !".oonh Amt·rica. m
br pJrncularly v.tlu.lbk, .md lw was .1bh: to t'luud.m· wh1ch the ,eJimcnt~ both of the .;hallo'" sL·J overly-
changes 111 pygidi..J sb,1pl' m no less tlun tour di,tinct ing the continent Jnd of the m:ugitl.ll 't•xogeown -
lmeage' 'W'here the,L tn•nJ, were .tpp.m:nt ''a dun chnt·' were represemed . Tht· regwn I' well
mdiv1d11.ll ~peuc' he dt,tangUJshed the dJ!Ji:rem l nrrdared ~traugraplm .1lly. rha~e ~rag~:~ bcmg pre-
\tage~ ,IJ1tcnosu,, mctlim .111d po~tenosus, gomg o;cnt .. In the oldest (C•zcno\ri: tn) stage fJ. 1(1/lll rra.wtu-
h-om comervativ c to pro~rc'~\lve. n:,pcctlvd y. In all ilermlclltl .md P. ''""' millm were both pre~c nt
1f these the dlar.ttlcr llll 111 111 the pnpu!Jnon ,Jufb. (conceanbl )- thL') could be se~u:tl d11110rphs of a \111-
brradually, and in c.tlh l,l\L' there as .1 gcncral pro- gk· ~pcne~) 111 the L'ptContincnrJI sc.t to the \\'l''L
brre~~l Oll 111 py"-'ridi.LI ~lupe trom broad :111d ~hort to wlule P. fell/cl rana liv~.:d contempor aneously in thl·
lung and n.1rrO\\. Luh·age' 1 .l!ld 4 ovcrl.lp: the oth- t'.l\tCnl are,J. In the .:xtn:mc WC\t P. iou'f//.)b wa' rl''-
"' arc• dMmct. lt \\,\\ p1htul.ttc:d th.n rhc p.m:nt hn- tdcnt. When the Cuenovia n tnlobites h,1d become
.:age of Oh•1111S w,1s .H tbi.; tlllll' hvmg t'l";•when:, .md c·xti iKt P . lllllcl r<111t1 ~prcad from the ea~t IntO the
that e.ll"h of the lour Jt,llnctivt· ltm•;1ges allually cpt•mc ,c,l, and when tlu~ sulhpccic' died out at tht·
I<.:Cil 10 the <;CCtlOil beg,111 \\ lth Jll 111\'.l<;tOn Ol an end of the' laoughmo~.m <>tage P ftlll<l /ltlnl'l.ltldt'IW S'.
emlutton.ll )' ofGhuot of tht• p.uent pnpulanou uHo .1 new mv.tder fro111 the e,t\tt·m exogco~ynclim·.
the are.l. fhe subseq Ut.'IH developme nt of Oh·11tts n.:pl.Kl:'d lt . Thus the tnlobitc' {)(the cp~eonrincnt.tl
through nme gener.1lly t~>lluwed ,m Hl.'r.ltt\'C p.mcm w.1. though they appear tu fom1 .1 mcce"a'·e cvolu-
of C\'olutlon, "hcr~· si1ntl.tr trend' .1n,~· m -.uccc·!>~IVC tiun.IT) o;equence 111 \\Inch the cyt· 1s reduced, .Ill'
1tock~ !Tom ,1 more or le~' untlun!-.'111~ .mce,tot. The .tctu.tlly dcnvcd from .111 .mn~~tr.ll popui.Hton living
morphnlog~tal c h.1r.1cters nf the ~nct·~rrJI ~rock are in the ea~t ••1!> a scrie' of invader,, ori~mally penph-
not kmt\\ n \\'lth cerrauu> but can be interred lrom aal a'ol.lte~ of a parental popul.tnnn whtt h as in th1.·
the carltc\t memtw" of hrtL·agc' I, 2 .111d 4. lrncage casL of the olenids dungcd rebtively httll:' through-
3, which p.trtially overlaps wah lim·age 4. has somt·- out time. This stud y al~u 1llmtrates Eldredgc ,111d
what dttft•rt nt clur.1ner; .md 'cem' to h.we come in Could\ model of Lvolutlon (t• .g. Fldredge and
!Tom a rdarcd bm tla~tmct ance,tor. probabl) hvmg (,ould, 1972) as a senc~ of 'puncruare d cqutlibn,1'
d>ewherc. Tl11~ <;uggcstmn l.)f the ongm of new The indiv1du.1l popubtiom of P. rmw t)nce estab-
group~ fi·om pcriphcr.tl isol.HC) of .1 n1.1in population ltshcd wcrt' rclanvcl) ~table .Jnd underwent lmk
~ amply ronfimwd by othc•r tnlobm· c\·olunona ry l'Volutional)' change lt wa~ onl) when .1 p.ucm
.cmht•,, though nnly tn hne.tgc J d1d rhe ob,erved population became 1.:xtanct chat the dc\cendanr~ of
crcnd<; lc.1d ro tht.' developme nt of .1 nt·w spenc·, ~null, r.1pidly evolv1ng popul:ttion s ltvmg .1~
gamodeme ' pcripher:~l to the main ance.;rr.tl popula-
Phocops rona complex tion could opportums tiLJil) mv.1de the nnw-vat.tl ll
Eldredge\ ( 1972h) ,mdy of evolulion 111 tht• M1ddle temtory. These then became eh..: doman.tnt widL·-
Uevoni.u1 Phacop.• r.uw of North AmL·ric.l dc<;tribe> a ~prcad spccit•s: a succe,~ful balann·J popul.ttion wllh
llSSil C.l\C of allopatric 'Pl'li,Jtion (Ciupter \). P. nt'\.. and \t.lblr charJcter state\.
mJ WJ<; \\;de<;prl'.ld over North Aml.'ncJ dunng
\1Jddlc Devonian nmc,, h.wing probably been Sheldon's Welsh trilobites
derived from a ruropean ,11\l.:estor \Ul h ,\<; J> lat- Ordovician (llandc1han) trilobite'> in Cc ncral W Jb
:r.,s fo the we~t of rht> ar~·.t 111 wh1c h P. ''"'" hved, otcur in ~'Tt:.u numbl'rs in a VIrtually connnuou '
'. l<lll'tll~l.' resided, the t\\ n spenes bl·mg more or ~t·ne' of hl.1rk shale~. Sheldon ( l9R7) studied .1
lt'ls mutually cxdmivc. Wlwrea~ P Wlt't'llsis i~ never 'lt'tllletKC spanning \OIIll' 3 MJ, in which there .1re
el) abundant .md ~howcd uttlc t'VOIUtiOn.lry etght common rrilobtte lineage' (F1g. 11.22).
hange thrnughnu r rhe Mtddle Dcvmu.m. tht•re wa~ In all of thl' eight genera, n\C..I,urcd trom IS 11110
1uch grc.lter var1.1b1ht) amon~ the 'ub~pCCll'' of P. spcnmcm, there was J net incrc,l\e in the numbt·r ol
:>1111, of which iudiv1dual~o uc uti:cn VCT) common.
pyg~dial rib,, J charJctcr used 111 'lpl'cie' dubrnom. lt
386 Arthropods
i~ a \triking ex,unpll of gradual L"volunon occurring becomes less easy to tract.
111parallel 111 tht· v,1rious genera Equ.Jll}, Shcldon fhe distribution of rnlobites in rhe Cunbn,m I\
demonstrated that thert• .m: charactt•r n:venals from complex (Cowie, 1tJ7 1). In a broad ,1nd general
time to time, Hlth J~ temporal)' denc.t~c in rib sense it appears th,tt there were two Lower
number. There •~ m1 re.tson wh> charattcr revcr;als Cambn.lll main taun.1l reJ.Ims. duractcmcJ b)
~uch a~ this ,hould nut rake p!Jc\.', .mJ here they olcnclhd ::md redltchnd tnlobites, respedn·dr. 0)
have been dearly Jemumrrated. Tlm " one of the the Mtddle and Upper Cambrian thl· (pll\\tbl))
be~r examples ot ~raduah,nc genetic ch.111ge known pdagic JgJlostids had .1 Vl'l)' broad dmnbunon wlnle Fr.
from the fossJI rcund, though the sdcctton pressures endemic bcnth1c pol ymcrids remamed Cl)tltlned to ~L'I
that caused tt rt:.'li1Jin uncertain. thctr 1solated continental blocks. (n<
Trilobite disrrihution 111 the Ordovtci.m h,,, been ce1
tltlly explored, and .Jttcmpts to work out the •on- Lu
Founol prov inc e s trol' of distribunon have on rhc whole l'een quite Ch
\UCcc,sfitl (Whitnngtnn and Hughes, 1472~ Cocks hH
Trilobttcl., J.\ much ,1, .my other fauna! group. sho"" ;lnd Fortcy, 1990). lt 1~ known dut the: low~ at i
provinnJ.I diJTt'rl·ncc' throughout the Palaeozoic Ordovician trilobite~ an: clearly dmnburcJ m a sa
wllll h can be uwfi.tl indicators uf the b10geography provinces, apart from a very few gcm·ra of cos- ,lllJ
of rlut time. Then· 'ecms to have becu f.1irly well- mopolitan distributton. Some of thc~c w1despr.:.1d pro
marked provinn.tll\m m the Lambn.m and espe- tnlobttcs (e.g. Gcrcwnosllls, Teleplr11111, '\rlrllt.<'mr) ,,
oally m the e.trlkr Ordovtcian. rim pmvmciahsm could well have been pl.lnJ..'lotuc gener.t. (. )the!'\\1~ tent
decrc.l\ed throughout the later Ordovtcl.ln until, b" no le's than four provmces have been ddmcd the
the !arc AshgJili,m. mlnbite t:mnJ\ were more or less named after char:lctcmtlc groups or genera Each C .1r.
c mmopolitan, a umtlttwn that pt'f\1\tcd through the provmce has endemtl f.umhe~ and endenuc gcner.~ pro,
Stlurian. In the l'.1rl; Oevunian Sl·par,ltC provinces of 11101'1! widely distributed £11111hl·s. ThN tlw 1
began once morl' lO thtl\.·renoatc, most parncularly a proviuces, when plotted on new palleogeo~ph~t·.ll tJu n
\OUthnn Malvino-K.ttTnc provmu:. bur following m.tp' (M cKerrow .md Srotese. 1990), .tnord vctv (mai
dH~ exnocriom of l.ttl' Dt voni.m time provinciahry wdl with Early Ortlov1eian ronrmental plttc:rru. have
and
Tc
~hc~:1
I ltm
C:o\llll
ut' d
Gracilis
Stlat..
M~kh ~
4
Cnemidopyge
7'..
Ogyglocanlla
r
Ordc
A~
llwpmle
LQ 1" ,. tlwrc
. PH ,o,. ---+.- .. p.tJ,Jeo
~EP ~
~TC
('
"
t ·' early
evide1
t,tl ( 01
Fo l
SNL •• ( nnt re
""WL ' \Vt'fl' I
BG !. f.tUil.\S
,1 cold
0 I > I $ 6 f a • 1 1 t n tt n • •onuuu\t•l
h.IVe h
Number of ribs ttatton
D t'\cOil
Figure 11.22 Gradual change in rib number in eight contemporaneous trilobite lineages from the Ordovician of Woles (Redrown .mJ \O
frOm Sheldon, 1987 1 Jatt·d 1
Trilobita 387
.md rontim1 the u11portance of palat·olatitudc tiated 111 rrilobltc\, rhough tlus j, le'' well known .
~ .ls a prunary control The vast conrmcnt of Denv,\llvcs of ::m acastid stock migrated ntto an
I Gondwana bnd \pread trom tht· South Pole to the almm l empty region, and diversified ncet·dul~ly,
tqu.uor and J1,play\ ,, tcmperat1.1re-conttolled l.mtu- constitutm g ont' oi the rno~t remarkably dl\nnct of
dm:~l clme Fir~t. the cold-water h1gh-latitu de all f.mnal province~ (Eidredge and Ommton. 1979).
)' ~hdve~ of tlm contincm earned an mdigenous Provinciali ty in the later rocks 1s poorly known but
'calymena ccan- dalmanita cean · f.."'una (c.g does not ~eem to have been great.
Fr.lncc. Spam, Central Europe and Turkey) In Arct1c North Amenc:~ cosmopoh taniml bcg.m
Secund, the luw-l.ttltud c ~helve' of Gondwana to decrease after Gedinni.m ttme, ~o the Dcvonun a'
(no\v south Chnu and Amtral1.1) carried a 'dikclo- .a whole seems to have been a time of intTl\l\ing
1\ cephalinid ' t:lllna. Tlmd, the trop1cal platforms of provinciali ty.
1- l.JUrcntu (North America. \1bena Jlld north
Chma). ctmed a faun.1 of b athyurid trilobite\.
Fourth, the t~olated U.tltic-Rm~t.m platform, lying
Stratigra phical use
at lntermedu tt' lautudl''• bear<; .111 cndemil fau na of
asaphid mlob1tes. Oth..:r foml\, e.g. hr.lc.:hiopod' Trilobites are of comiderab le ~tratiguphica] value m
and mollmc~. .uc endemic to these ~ame four the Cambrian and Ordovitian but nttH:h less 'o m
prnvmces. later rocks. except perhap~ locally. The C.1mbnan
In the later Ordo\ inan thl· 'cparate provinces ~ystcm 1\ 7oned almost enurdy on mlobite,, other
tended to mmgle as ,\ rt·,ulr of the closer approach nf than the basal (T onu11otian) zone. Cambn:~n trilo-
the Gondw;Jn:-tn and Lhltic contment~. L3y the bite5 obey most of the requiremen ts of good zone
CaraJoc only the 'ulymcn.t n.•m-dalmJm taceJu' fossth, being abundant and easily rerogmt.1b le and
provint e n:t.uned m lntt.'gritJ. Thl.' fate of some of often having short tlme ranges and wtdc honzomaJ
cl1e mJCrocontment~ t.\11 bl.' tr.1ced hy me.1ns of thelr d istribution. Thcy ~1re hmited by bemg di-anbutt·d
fauna\. The ~mall coutu1enral !Tab'llll.'nt of Avalonia in thc faunal pnwinces de\CnbcJ above .md by
(m.unl) ~outhem Hntam). tor I.'X.:tmple, i~ known to being rather f.1<.:1es controlled . Though good
1\ , haw brokt·n utr Gond\\ ;uu, migrared northward~ mtraprovm cial ~cquences are establt~he<L ir " only
and umtt·d with LlUremia. possibk to corrcbte between them where f.mna~ J.rc
1 O\\ ards the end of the Ordovician .1 pol:.u ice mixeJ or by u'mg pelagtc tnlobite\.
sheer fanned and ~eatlv cxp.mded dunng the The Ordovicun -;ystem IS zoned b} b'T.lptohtes
Himantian (the final ~taw:). This t\IU\Cd \\1despread (q.v.), cswbli~hl'd for the offihore sequences.
e-xtmcuon ,1nd i~; conmieted eo be the primary camt' However, the Hratigraphical ~ubdtvision nf the
.. ot the brt•.tkdo\\ , ot provinn.tli ty around the

OrdovH..ian-S1Iuna n boundary .
As well a' ,helf t:mn.ls dunng the Ordovici;m
nearshorc shclly 6cies 1s based upon tnlohttc' and
braduopod s. whteh give a refinemen t of corrl'btlon
of special merit in the Car:~doc and A.,hgill. In the
there were tkeper-wa ter bentluc\ marginal to the British type area no stages have yet been dctined in
rabeoconti ncnb (e.g. the olemJ biofancs in the the Arenig, U.mvim and Llandeilo. but only Upper
early Ordov1oan ); such deeper-wa ter f..1un;1s arc less and Lower divi~ions (the Llanddo has a 1111ddle
evidently controlled by rulaeolatit uJe and continen- d ivision). In the Car.ldonan . however. eight 'tage~
ul contigur.tti om. have been ereued. based mainly on bral h10pods
Following the Htm.mnan glaoation, fauna.~ d1d though with trilobites (espcoally tnnudcid') b'Wing
not recover very qmckly. BccJU\t.' the rontinen~ con fi rm:uory evidence. In the A~hgillian m.my
were dose together bv tlm tune tropical/te mperate kinds of trilobitL'~ have been used tu define 'everal
13un;h became \\ 1dcly d"tnbutcJ , hence . .tp.m from zone~ wtthin the four known stage,, 'o dfec.:ttvely
1 cold-water Clmkera f."'una, S1lun:m t:mna\ seem to that sdccted area~ of htholog~cal monotony havt·
have been more or lcs~ cmmopolit .lll. The Jlffercn- been mapped u~i n g the trilobite fauna~ as marker~.
bmon of the Mah;rw- K.1f:Inc provmcc 111 <iiluro- There arc still problems with correlanon mmg
Dcvonian tunc' (South A.ment.l, Falkl.md Isbnd' Ordov1C1an tttlob1tc~. nlJl!lly becau\C of faLll'\ con-
~nd ~outhenunost Afnca), which 1s so well clun- trol .md provwciali ty; thl're 1s still. for c:xampk, no
datcJ m thL· br.tchiopm h, appeal'\ .1bo tu he mbstan- cl me tune correlation between the faunas of vanous
388 Arthropods
Upper OrdovKI.lll red~ - DaJarna (Sweden), Class Meros tomata
Kci~le)' (northern Engl.md) .md Kildare: (Ireland) -
and tho~e of L"Ontc mpor.1nc:om bedded carbonate or Subclass Xiphosuro
m ud~rone f:lt 1e' Limulus
In ~tlun.m .md I.Hc:r rock' tnlobites arc nor of The char:~cter. of xtpho~ure~ are well d.tsplaycd b)
gre:n ~tr:ltlgr.lphtc.tl ,-.,lm·. for other tosstls have gen- rhe modem Umrtfu~: om· nf three closely related
erally ~horrer tnm.• r.m~t'' or m.l\' othenv1sc be used genera wh1ch .1re rhe only llnng representative' of
m on.· t tTt·crivdv. \11~ h ·'' the: I )e, oman .u11monmds this o;ubcla\\.
and Carbonifi.·r~>tl\ llllt rotm\ll\ On the orher hand, Limulus p,,fyphemus (Fi~. 11.2Ja-e), the type
tor lot11 corrd.mnn thl'') .1re cc:rt.lmly of some use, sp ecies. hve 111 shallO\'\ watl'Ts .tlnng the northwest-
and their v.llul' ,1\ o,rratl~'Taphical mchcators may cm Atlantic shore,.
incrl':tSt (Tho111.1' rl al•• JlJK4) In dorsal vte'W 1t ha' a \ aulted 'emiCJrcular pro-
soma, w h ich ao; 111 all cht:hn:r.Jte!> t\ ,, fu~ed cephalo-
thorax arriculatmg wtth a ll's' v:tulted pl.1te-hke
abdomen , there bemg .t narrow dunncl mosed
obliquely between the two. From the n:ar of the
abdomen ~p n n ~ a strong tl'n ninal ~pine, the telson.
Chehraatl.'~ (C.1m - H.t•c.) IIKiude sp1ders, nn tes aud The axial part of thl' prosom:l (cardiac lobe) is
scorpwm, :ts wdl .1~ the ho~c:shoe crab Umufus and defined by axtal furrows so that it has a superficial
its liv·mg .md fossil .1lhc,, and the extinct l:"urypterids. 'iimilarity to a trilobite glabella. Out~idc: the a.XJal
This dJvt·~c: group of ,lnJm 11, \\ h1ch .1ppear ~o het- furrows ,md parallel with thl'm arl' thl' ophthalmic
c:rogt·nc:otl~ .1TL' tmtted b) P"'sessm~ the following: ridges which curve anrennrly to jom med1ally, at
which point thcrl' arc ,\ pa1r of 'mall o celli. There
Jn .llllt'nor proson1a t)f \IX ~egments whiCh is are abo compo und eyes situated m the mJddk, of
eyu1vah:m to rht• fml'd hc.1d .md thorax of orher the ophthalmic nuge~. Thl'sC l yes h.1Vl' many len\t:.\,
.mhwpods. but rhctr structure " quite unlike that of tnlobite\,
.1 pmtcnor opisthosom a (.1hdomen) \\;th 12 or the dwptnc part~ bemg tompml'd of mvagmaoom
le" \C:gll1l'nt\, of the cunde w)11( h fonn long, tnwardly projecting
a pa1r of JOinted J'llll'l'f\ (tht· chelicerae) w hich cones. These arc ut parabolic comtructtom opn-
give tht• 'ubphylum it~ tuml' and which are mized for maXImum ughr t•ollctU011. Bdo\\ th~~
alw.t}' prl',clll tn rill' tiN (prc-oral) segment of arl:" apposmon-rypc ommatidta, l'ach wtth a markt·d
the prmoma. excentric cell.
The .abdomen arttculatc\ to thl" prmomJ along ~
Two m.un t-,rroup' W1th111 thl' cheliceraces are of hinge wluch cur~ atro'' thl' primary ~egmentatJon.
\mgul.1r p.ll.tl'lltHnlt>~ll.tl .ltld l'vulunonary interest. A.,xial ridges are prc,cnl, ant.! trace' of 'eb"'ltentatlon
are evident Ill the ti-ingl' of six movabk spines. The Fig
CLASS 1. MEROSTOMATA (t .m1 -Re~ ) Aqu.wc chelicerJte,. strong styliform tt:blln. rid~ed .1long its length, 1s OPJ
otrc:n ot l.lrge sm· l'hl'rl' .m: two \llbcbm~s: SUBCLASS sw;
nearly as long a~ th 1.· rest of the body and can tu m m Lim
XIPI IOSURA wluch w1th 1t' ORDER XI PI JOSURIDA
almost any dtrection. Wt
and ORPER Al:LASPI()A mdudc\ the L.imulus group and (x{
the1r rel.•nves; :uH.l ~Ull<. LA')S I·UR Yl'1 ERJDA which T he segm entation of thl' bouy i~ readily worked
red1
Include~ rhl.' w.ua ~lorpion' nf I'Jl.ll'OZOH nme. ou t fro m embryology, but i~ also dt:ar in the: vcntr.~
CLASS 2. ARACHNIDA (~ul. Rl•c.) Donunantly rerre~trial morpholoi-,ry of Limu/us. The prc-oral 'cgrncm car-
fom1~. the \)11dl·r., mm·s .llld \lOrpwm: the latter bear a ries the short, patreu. thrt·e-jointed chelicerae, .md pai
\Upl·rliu.II rewml>L•n~t· to Clll)f'll'nd, but .ue not closely rhe tive pair. of post-oral appendage' arl' arranged
related. Though \UH'J'Illll\ olrl' l.JIO~\ 11 t:i-um the Suunan and arc
radlally around the mouth within the u1dcnted ring .1 p
nlllc\ ami ,p,Jer' li nil\ till' [),·volliJil, .ill are rJrc J> fo,~iJS,
otht'r tl1.1n pt"rh,tp~ tlw lt•rtury 'l'•der. prt... ~rved tn ambl'T. fonned by the antenor uoublure. These five pain of th e
Fos\il "t)rp•on' .uc kll<l\\ n in Ulll\ldt:ublt• morphological post- oral append.1ge' are morphologic.illy sunilir to \
der.ul. but the1r study'' hlyonJ thl Slopt• oithi:. boo~ 1nd one another and fu nt:llon as walkmg legs. Each h.ba .lbd
ar.achmd· wtllntll h,· ~ \111\ldt:n·.l fimhn. basal 'pinose wxa Jnu \CVcral jOJnt,. Thl' fir~t four itl cc
lun-
Phylum Chelicerata 389
(h) Agl..p is
(Jl B«kwltht.
(c) (d)
(e)
:§ -
If) Maollmulu• Wllkhll
(m ) Euproops
.~...-_-·(•
.
"'
Figure I 1.23 Merostomato . (o)-(e) Limulus polyphemus (Rec.): (a) dorsal view (x 1); (b) ventral view (x 1), (c) fourth
appendage (x 1); (d) swimming upside down with the body inclined at on angle to the horizontal; (e) section through prosomal
1wimming Limulus showing recirculoting vortex; (f) Mesolimulus wolchii (Jur.). section through prosoma (more Rol1ened prosomo of
!imulus) showing vortex; (g) Pa/oeomerus (L Cam.), (h) Aglaspis (reconstructed; L. Ca m., x1l; (j} Beckwirhia (L. Cam., than that of
Weinbergino (L. Dev.), Germany (x 0.75}; (m) Euproops (U. Cam.; x1); (n), (p) l.egronde la (Dev.) in la teral and x 0.75), (k}
dorsal views
I>< 0.75). [(dHf) Redrawn from Fishet", 1975; (g) redrawn from Bergstrom,
1971; {h), (j) redrawn aFter
redrawn from S!Ormer in Treatise on Invertebrate Paleontology, Part P, (n), (p) redrawn from Eldredge, 1974.]Roasch, 1939; lkHml
pair~ lcnninatc in pincer-like ch elae; the last pair 111 all chcliccrate \, bears the genital o p enings. The
lfe cqutpped mstc.1d wtth numerous 'pme' and have \cvcmh 'cg~ncnt •~ rudimenta ry and " apparently
1 peg-hh outgrowth (tht.· flabellum ) attJchcd to rcprescmc d only by a pair of reduced pl.ltes. Bd11nd
rbe coxa. the operculum arc the ~uccceding five gill
Wtthin the broad vt.•ntral doublurc of the appendage'> which bear the re\ptrarory gills on thetr
1bdomcn arc \IX ovcrlappm g plates which arc mod- mncr (dor'\al) side. where they arc protected ag,11nst
Ified appcndagt·~ . The tint of thc~c •~ the operc u- desiccation.
lum, morpholog icallv the ctghth segment whtch, as
390 Arthropods
Life hobits of Limulus Unmlus bt:mg dttl'erent Jll\\\Cr; to the ~.unt: prob- Lu-
Lmrul11s r ~ ,1 ~hMdllll' mh,1b1t.mt tolerant of a wide lem. Howcvcr, Um11/11s has more room m which to w
range of ~.lhmttl'~. hvmg Ill 'hallow waters and capa- retract itS legs and 1' morr: dfec.:ovdy adapted tor C.X•
ble of cr.1whng out ti.)r 'hnn dJ,t.lnce~ on land. It can burrowmg. sur
'' aJk on the sl'.l floor on thl' prmomal appendages, Ul
but H c.m al'u ''' 1111 .md right tN:lf wtth the relson 1f BURROWlNG LIMULUS A~
overtumed. lt 1\ .111 t'Xct•cdmgly vcrsattle animal Umulus ts acuve at mght. but dunng the d.ty 1t bur- .1rc
whme hfc lubJt\ h.tVl' been mcd wHh 'omc ~ucce~s rows 111 below the ~cdtmenr ~urf..Kl for .1~ much a_~ 11.
in mtcrprering dlml' of tnlobtrc'. ro whtch H ts the 12 h at a nmc (Eldredgl. 1970). Adulr ~pecimem PY!
c!mc~r hvmg rcl:ltl\'t Jtld to whtdl tt ha~ ~ome mor- have an antenor arch. as do tnlnbtte\ Ar thr: \t,m of art.>
phologic.ll rc,cmbi.HKl'' llo\vcvcr. there musr be the bunal procedurc the arch 15 first lowered so thJt wh
pamcul.1r funcuotul fl',l\011\ for thl''l! \mularitics, the amenor t.'dgl· nf the pro~oma come~ m contact ~lllf
and the 'pcnltl me' of thl v.1nous organs must be With the sea floor; 1t th(.•n dig~ uno the: sand a' tlw t'VO
titlly undeNoud bd(lrt' \UCh .m.tly'e' .m: drawn. prosomal leg-; pcrtom1 normaJ w:~lkmg movements oft
S;md 1s pu~hl·d b:~ckw.1rds, covenng the prosmm a pp•
SWIMMING L/MUWS and the an tenor thml of thl· opi~tho~oma. The tel- ob~c
Lrmulus mmn;~lly \Witm up.,idc down (Fig. 1l.23d), son i~ buried and only makt·~ horizontll mnwmcnt' ann
inclined at .1bout J0° tll the honzontal and moving thereafter. The walking legs t•xcavate a burrow. likel
at .~bout Ill- IS u11' 1 l t " propelled by both the pushing the sand up bct'\vcen the pro~oma and men
pro,omal .md npi,thmom.ll appendages. The opistho~oma. One of these cha nnel~ renwim open refer
opi,thtNII11.tl .md ,l,th p.ur of prn,omal app(:ndages when the :mim.1l 1s compk•tdy b1.med. When thl' Palm
move m antl'nurl) .1dv.mcmg w,tve~ of metachronal first ~tage ofbunalts etT~ccted. the ammal i~ then qut- three
rhythm, wh1lc the tiN f(mr p.ur~ of walking leg; escent and he~ for .1 whrle oxygenaong the g1lk 1.3urg
move m ph,l\c. c.xtcndtn~ ·" th~:) pu'h backward~. There 1~ then a thtrd \tage. when th<..• pro~omal k·g. Tl
withdra'' ing into tlw prmomal cavity or vault as wJlk forward-. once mon• whtlt· the npr~tho,omJ 1' cardj
the} ret ovt•r their po,ltlon Th1s qrokmg cycle of flapped vigoromly, mtng the tcl~on ro some l'Xtcnt. and
the fiN appcmltgc' bq;m' dm:ttl} .tfter the si.xth and stm up a cloud of \and \\ lucb scttll'\ to cova Ther
prosonul p.ur h.t\'1: cnmpk•tt:d the1r 'trokc. the animal complctdy. Thl· f111Jl st.tgc ts om· of
The ll'ydrod) namit' of tlow m the swimming absolute qme,cenu.:. on I',. J \lnglc re\piraton rh.m- ORDEl
U11111lro have bet•n aru(y,t:d U\lll~ ,1 mnJd Lm111lus in nel being left open on ont• 'ilk. SUI
a flo\\ thamblr (f-i,hcr. 1975) In the nom1al lt has been shown expenment.tlly that the dor;JI ol'
inclined \\\'tmmrng .tttttudt: .1 strong rccirculating setae are U')ed ;t' rncrhanorcrepto!'\ wlurh mdic.tre do
vortex tonm '~ rthut the pru,omJ.I vault. Thi5 to the ..tnimal when tt 1\ fl.tlly buned. The td~on lS wh
appear' to brea~ Jm.,n l>r bt: sht:d ,tt intervals into covered from ,t,lrt to flrmh and 1s only u,ed to ~ sue
~CgJ
the ,.,akc of the .uurn,tl pnor to rt:-tonning. The limited extent in the thtrd stage. lt ti.mcnom matnly
app·
vortex ts ,1 nornul ttmst'<.jlll'IKC of flow dynanucs as a stabilizer or ntdder iu the w.tlking or swinmung
tor an obJN't llf sut h ,1 sh.1pe .md inclination, but it individual or tn nght111g one thal i~ overturned. Tbt•
seem~ to be cxpll)iteJ by the ;mutlal in aiding the comparative morphulOb'Y of Li11111lus .tnd trilobites m Much
torward movement of the .mterior prommal such details a~ the prt·~cnce of dorsal sensor~ aml the ~it)r lh
appendages in the fl'lUVl'ry ~troke. \\hi eh would vaulted slupe of the cephalon \ttggl·~t~ th.lt trilobuc~ .lnd th
otherwise hJvc to pu~h agJlll~t the current. At might have 'pent mmc nf their time hurrownl Ill drscov
grt:ater ur lc~st•r mdtn..ttwn' the vortex would be less ~and, and this is amply bome out by trace lchstl IJ.."Oil,
effectiv~..· or Jbsem (rtt-\. ll.23e,t). evidence. Cbl W
Sinul.lr moJds uf thl• Jur.mtc ,\lnlllilllulus u•alchii, ~ynzjp
wltich ha~ a mud1 lllt.'rt: RJttl'ncd morphology, Other xiphosures and their geological history ~Cill'ra
'how that an C4l11\ ,tk·nt voncx would operate in a Xiphosures arc rare in the fo"il rl·cord and, uth :r has a I.
~mul.lr \\',ly provtdt•J th.lt thl· .uumal's angle ofincli- than Um11l11s and m modem rdanve,, arc gctwr.ill} Hr mode
uauon wa' k"'· H cnu.• therl' i, a d.Jrett rdatiomhip confined to non-nurrne sequences. the pc
bet\\ccn the ..lupl' uf the pro,oma .md swinumng The oldest wdl-kno\\11 \.tphmurc~ belong tu the rather~
abtht\ . thc dttli!rem vaulunp of ,\lt•st>lmwlus and Cambnan Order Agl.bptda, wh1rh arc: known Vl't)
largdy from rhc Upper (. '.unbnJn of ~omhwe~rem the Lower Cambri. m llt>linwlus, is known from two
Wiswmi n (R.;ta~ch, I IJ39) Thc<it: h.we ,1 phmph.t nc partial prmoma~ only. it wa.' mannc, as .1re the.·
cX(l\keleron m whH.h the prmom.1 I\ t)1'1llll} xipho- much bcner-prc~crved Devom. m genera. but some
'llrJn. rhou14h wJth Vl'ty pmmJnt :nt compou nd l'yes, of rhe Carbon iferous .llld latcr ~rroups 1nvadcd and
m which untortu1 utd) no ~tnu:nm~ tl·mains. In colonize d brarki~h water hab1t.1t'i. 1 he (_ .ubom-
.4~/a.sprs (F1l-\ 11.23h) the I l m 12 thor.lctl' segnwntc; feratl~ Bdr11mrrs .md errprtlclps (F1g. I I .23m) arl· small
are li-ce .1s m mmt .1glasp1ds, but 111 Bcd•.!tl'itlrill (Fig. forms, often tiHmd 111 nodule\ dc.·pomed m coal
ll.23j) thl posteno r thn:e m four .m.~ tiascd mto a ~"'amps. In the Carbom ferous of llhn01s (Chapte r
pygJthum-M.c pLJte ln ~mne 'penmc m .1ppen<.l.1gc~ 11). whKh ha~ been very thoroug hly tnvc~ug,Jted,
arc pn:serw d. Evid~:utly tl1c.· tiJ'\t p.ur bore chd.1e Ertpr~lop~ almo't .llways ~eell1!> to be .1\\0ci.m:d '"ith
whill the re,r, indudm g the Jbdomm al one~. "l're plant r~:mam~ rarhcr th.ln with other ulvertcbr,Hes of
smgk walkmg Jc~. Jt wuuJd be po~\ibJe W inter th.H ponds .md lagoons. h5her ( l97lJ), adducm g much
evolullon 111 the x1pho\UTl'\ procc.·elkd cowards fi.t,ton other cvtdcnc e, believes th.lt it ma)- have been at
oi the Jbdommal \l~nent\ .md modtficatJc.m of their k:~~r part1ally subacna l, crawling up U)re~r trt:c~. ;lnd
appendage~. Howeve r. xiphmurc.· phyloge ny ic; di~b··uising itself to resembl e the 'urtacc of the rree~.
o~urc, and 1t I' not n:rtam "''ht·tlu:r Jgl.l\ptds wen·
Both Belimrnb and ruprt>Op.> have \\ cll-de\T ioped
aiKC<>tral to .my latt·r Xlphmurc~ or whethe1, as i~ more
cardiac lobes and ophth::tlmi<.. ndge~. BdinunH, hke
bkcly, they were ~1mply an early .md c;terile 'experi- the Devoma.t1 Nmbelinuropsis. has some !Tee ahdom-
mental' group. The oldest .mhropo d wh1ch has bc.•en 111:11 o;egmenb, but in Ertprvc'p~ and Limulus the
rcfern.:d to Subclass Xlphu\u ra i~ the Lower Cambna n whole of Lh~ ,tbdomc n is fused, though ~urtac.c
HriJt'tli/JtTits (Ftg. 11 23g), but it I'> known from only rr.1ce~ of se!:{lm:nratJon are vel)' pronoun ced and
thrn: ~pcdmcns and l-mtld be J rclam·c ol 'ome of the mcident ally v~ry rrilobac -likc. h is of '1.>me mrercst
Burgess Shale arthropo ds ('111crmtomoid~'). tlut the first ti cc larva of Limulus, when hatched
The Xlpho,u nda nonnalk h.wc the dtstincn ve from tht' egg, has a ~tmilar pronoun ced segment a-
md1at lobl and ophthalm ic ndgl'~ wdl develop ed, tion in the fus~:d abdome n to the adult Eupmaps. i t ts
md there .If<.: In or fewer :t.bdomm.ll ~cgments. colloqui aUy known as .1 'trilobite larva' without an)
There arc m o suhorde~. 1mplicJ non of a clme relations hip. The promma l
appenda ges of all Linmlin:~ arc wry 'in11!Jr Cl.)
O~OER XIPHOSURIDA (( Jm.-Rn thmc of Limulrll. The. earliest tme lunulid (Rc1l/e1t1)
SUBORDER 1. SYNZIPHOSURINA (Ord.-De v.): Mo~t ~~r .1U .1rosc !Tom a bclinuri d ancestor m the early
of rhr abJomma l sl.'gtrt~llh .In.' fi-n·. Jnd the .tppeml.t~l'' Carboni t(:rous. One of th e large~t known fm~1l
J,, not app ·ar w h< chcl.•t~· (otha dun the chchcc<n r, llmulmd ' 1s .\m1iopyra111is, descnbc d from northcm
which .1rc not kiHl\\ n Wtth , c<rt.llllt) ).
SUBORDER 2. UMULINA (I l)cv -Rt·c,): Till' .tbdwninal Enghnd (Siveter and Sddcn. 1987). lt is fmmd m
~q:mem' mav be fu~c.-d. though nut alW.l}~· and the ,hallow- marine deltaic o;edimems aud the 'ingle
~pp.:ndagr\ .1rc nomullv chcl.trt'. ~pccimen is pre~crved with J current- scour nurk m
front of the pro~oma in the enclosin g sc:dirncnt. Thts
large indiVIdual, 14 cm .lCrQ\\, was probabl) a bur-
~tu rh conti1~1on ts a\,on.Ht 'd wnh .lttcmpt ' to c l.l~-
rower wtth poor swimmi ng abtllties.
1lfY these ammals, ~lntc the.· iosstl record 1s so ~linty Front the known fossil Limulin J it m:-ty be
;td thl· prcserva nou 1' oltl'll poor In suth ca'e' the inferred char there "'.J-' ,, general cvoluoo n.ll) trend
J1srovery ol a ne\\ 'peomc.· n, t'\peciall y of a new Low.uds a fused short .1bdome n ;md l.1rgcr ~i7c. The
uxon. often ncces~it.ltc\ the raxonom tt rev,swn of Penni:111 Palm•tl/imulus, the Tna~sic Lrmultt,-1/11 and
•b,· whole ,._rroup A n.·cc.·nt cl,t\Sificauon of the rh<..· Jura\SIC Mnalrm ulw differ from modem genera
~ynnphost1nna (Eidredg e, 197 -t) tnclud~,·, only four
mainly 111 their smaUcr dimcm1 0ns and Lhcir pro~o
;!l:nera: 11'dllllt'f'.~illll (fig. I l.2Jk). Lc:,rwuldla (\\ lmh malloba non. 1 hq were aU m.mne [onns. Amongs t
J' a large and VC!) Lmllllm -hkc eye; hK 11.2.1n),
tom! limulids , ~cveral ~pecimens cxm of thc Upper
Bm10dcs and Li11111h•rrlcs. Thl· Lumtlin a. other tlun Juras~ic Alcsc>limrrlus from the Solnhof cn Li1ucsto m·.
n~: pecullJr Fam1h p,ludom sl·1dac. .\re nnmly
preserve d .u the end of .1 tr.ul m whKh the 1mpnm '
nther ~mub1 m comtruc twn ro lrnudrr) , fhe oldl•sr, ol the appcnda~es can be cleJrly seen.
392 Arthropods
Subclass Eurypterida sepJrJted mto .1 mcsosotna .1nd tnctasoma ot ~lX
The eurvptends 1lt rdJuvdy rJIC: but always spec- segments each. I he fom1cr beJrS appt·nd.tgc:~. tlu- bt-
tacular tosstJ... I ht·y r.mgcd from Ordovtciau ro rer does nor. The<;t.' appenda~t.·s look like stenutes bnt
Pemu.1n .md .Ut' ti.ltmd in nunnl'. bradush. hyper- are m t:Kt m1e appenda~t'), The se\'t.'nth )l'ginent IS
saltne and frt·,h\\·,ltt.T lidltli,Ktcs. reduced as the merasroma. wlulc: tht· etghth. a~ m ;ill
furyptt·rid murpholog\" " wdl dtsplayed bv mc:ro,tomc\, Ius the gt'mt.ll :tperture. Tht~ segme1ll1S
rhc: ~tluriJn H.lilc •fiii')'J•It'lll~ ll'lr,tl/•'"''f'/11/11/mrt.~ {Ftg. known a\ the operculum .1nd. together with the ti.1ur
11 .24a-g), tonnerl) known .1s 1-'lll')'ptt'ms_Ji_;c/un. mccecdmg <lppend,1ge<~. j, plate-hh .md ,\CUlhCd
l'ht.· morpholo~y of thts spt·nes was one of the >llong the Jntenor horder so that they all ovc-rbp. In
first to he rt'.lll) wdl under.tood, l.1rgcly !Tom the . the centre of the operculum and dtrccted po.;;tenorly
work llfl lolm (IHlJH: .llld l.ltt:r by Wtlk 1965), who 1s the genital appen dage. ;ln clong.ued and ehbo-
1\0IJtt.d tT.lg.m·nt, of tht· t•xm kdetnn from rock, as r;ltcly ~culpmred rod of \\luch two kmds may lw pre-
ht• d1d with W·lptolitt''· sn thJt they could be Hudied ~t'nt in >1 smglc curyprend population. )ll~'t'~nng
as rr,\mpMetH' le' In dor..1l vtt'\\' the prosoma is sexunl dimurph1~m. In Haltocuryptcnts one kmd (t)pe
l:lrge .1nd trapt.'7\lltl.tl. with promment compound A) ts clong.ued, and the other kmd (type B), rhout;h
eyes hetwt.·en wlm h .lrt.' ~m.tll p .ured organs, p roba- ntbular, is much shorter: othlT g~:nera have gctlltll
bly m·elli. lkhind thi, ~~ the o p isthosoma, consist- appendages that ,1rc of .1pproxim.ucly cqu.1l lcn~:th,
ing of a hroad thttt·m·d pre-a bdomen of seven though d1tTcrcntly shnpccl (fig. I J .24h,j). In
scgmt. nh, .llld 1 n.lrrowl't' .md more cylindncal Plt't')(I?MIIS .md some other gcncr.l type A 1~ l'OIN~
post-a bdome n ol only fi\•e sq.,'11tt·nts, terminating rently :mociatcd wnh 'cbspmg o rgans' on tlw pro~1-
m a ~tout. pn111t~d rd'l'll E.Kh of the abdominal mal ,1 ppcndagc' .md Sel'lll!> to luve heen J null'
scgnllllC~ 1~ lOlllpll\L'd ul J dctr..JI tcrt...-it~ ;md a vt·n- copubtory Ol'g;m: the olhl·r type " bc~t imeq'Jct,·J as
rr.ll \tt.·mltL' Th~ wntr.Jl appcnd.1ges can be ~een .1 tenulc ovtptmtor. Tr.lt.c~ of intcnul duch hl\l'
from the dlll'\,tl \llrt:lt.L' prll.Jeltmg wdl bl·yond tht been distillb'lll\hed 111 bmh kmd~.
bod)- The \'l'ntr:ll llH•rplwlogy j, complex. The The opcrutlum Jnd thl' 'uut•cdmg four
prnsoma h.h .1 bro.td antkncd doublun: with a mar- appendages of the pre-abdomen covered th.lmhcn
ginal 'uturc. u'ed Junng ecd~ ,I,, .md oftt'n \\ ith in \\ htch the g~U, were 'JtuatcJ. The \tnlltUI~ o
patred LOilllCtu\·t.• 'urur~' l,oJ.mng J median dou- tht-sc io; kno'' n in \/ttrll•nicl and better ,nJl 111
blural pl.ltt' (the: c pisto m a) \\ nhin the doublure Tarsoptelt'lfa. ''here the JttTt.•rt.•nt pattt'm.s ot mtcnu
i~ a softcr 1ntq.,'1.11ncm \UITounJmg: the mouth. to and extemal \urt'Jcc sculpture. even in lhttrned
which tht.• .1ppcondJ~l'' uc: attJchcd. The tirst (pn:- spenmens (W atcr.ton. 1975), suggested that th<' g~Th
oral) patr of JppcndJ~C\ are ~mall chdtCt'rae. 13c:hmd were spectaltzed vascul.lr tr<lt.t~ of the ventrallxld
th~se po~r-or.tlh trc lour p.1m of stout. jomted wail protected h) the .lppenJagc' (F1g. 11 24~1
w.ukmg lq~'· tncrt.'.t~tng m \lZc: towards the re.tr legs, M ore recently (Maunm g .llld Dunlop, l Y95) 1t h
which arL' n· hod nul .md split}. Thl" sixth pair of been ~ hown that t•uryptcnds ha\'<.' a dual resparaton FigvrE
prosonul .lpputd.l!!:t.'' Uo..' VL'l) l.1rgc and have the system. The brandual ch.1mbl·rs housed 'book chelicc
dle-lik.
ternun;:tl p.tno. flattem·d hkt· paddles. These last lungs' 111 stacked rlun sheets wlth nbbed ~urt:Ka with fi1
append.1gc' wert' t.'.1p.1ble of beu1g protracted and providing extra poten tt,ll fo r resp1r:ttory exd4Hlf.\' tical el
rctmcted .md Wl'IT probably used .1s in swimming, They .1rc very sinular to tht.• book lungs of scorpt· (k) To1
in .1 bre:mstroke nwmcr. fhe LO'<a of each 'leg' is ons, bu t .1re dehotL' ;md seldom preserv<'d In
Paleon
brge .md Jnlli.Hirt.•d mtenully with ~pines projecting additio n , thl' g1JI-tracts, or more proper!,,
a~ gnathob,l\l'S A sm.tll U-sluped plate (the endos- 'K iemenplatren' on the roof of the branclu.1l chJm· pt..'n11.
toma) border. the mouth; th1' JS nonn.llly covered ber, have downwardly t3ung patches of httl<' spmc:s time n
by ,1 much l.1r~t.'r pl.tte (the mt>tastoma) whiCh ts coverc:d With ~m.1ll prOJCt ttom 111 ht•x,lgorul
accually part of tht• ,1bdmnen Jnd 1~ Jll that remams rosettes. I'hcst> .1re tntetvreted as \\'ater-trappmg Cu1
of the reduced ~evunh \t'~lleiH . lt .1ho covers the deVlc:c:s. hnked to the v.t~c\llar 'Y'tem. and prov1ding The c
proximal p:tm of the coxa . an .lddmon.ll '\urtlce tor ga~eou" exchange Whcr{; men~
Tlw pre- .md pmt-.lhdomt•n o f 'even and five seg- euryptends were pnmarily water dweUers, the} nu} ofcxtJ
ment\, re\pt't'tl\'dy. h.we been defined on the post- well hJve been able to m.1ke excursions on land far borJe
cion of 1hc '' .u,l. hilt tht· .lbdomt•n can aho be: penods of nme If so the · Ktemenplatten·, \\,th thru dld rac
(a) lbl
postabdomen
(hi Type A Ul Type B
Sltmonill IICumin•t• T•rsopter.tl•
Figure 11.24 Eurypterida. (a)-(e) Boltoeurypterus tetrogonophtholmus (Sil.): (o) dorsal view (x 0.35); (b) ventral
view (x 0.35), (c)
diefi<era (i) (x 0.35); (d) walking leg (prasomal appendages (ii)-(iv); X 0.35); (e) ~fth prosomal appendage (X 0.35);
dle-like appendage (x 0.35), (g) section (d1ogrammatic) tnraugh cuticle of B. tetrogonOphthalmus, showing inner,
m sixth, pad-
non laminor layer
wilh fine vertical canals, middle (laminor) layer with laminae more closely spaced ex1emolly, and ovter layer with indications ol
licol elements, (h) 5/imonia ocuminalo (Si I.), type A (?mole) genital appendage; (j) S. acuminolo type B (?female) genital ver·
(k) Tarsopferello (Dev.), simplified reconstruction of o gill chamber. ((a)-(e) Redrawn from Stormer appendages;
in Treatise on
Pa!eonlology, Port P; (g) based on Dalingwoter, 1975, (j) redrawn from Woterslon, 1960; (k) redrawn from Waterslon, Invertebrate
1975.)
pcrmacntly moist watcr-u.w crcd ~urtaces, would mcntary remains in sedimenta ry rock can be imme-
fimction :1.~ cerrcstnal lung.. diately di~cmguished as being ~uryptend 111 origiu.
Ultramtcro graphtc work (D:Ilingwaccr. 197 3.1.
Cuticle I1)75) ha' shown that curyptcnd cutlclcs are pre-
The cuticle of curyptcnd ' I' very thin, and \pcci- served a~ silica, though undetermi ned nrganic m.tte-
mem arc usually crushed. It may be.u different kmds nal wa~ ongmally present too. The outer cuticular
,f external sculpture, notJbly tcrraCl' lincc; round the tone (Fig 11.2-lg) The lanunatlom resemble those
!larder and somcnmcs on the lower surface~ .•md a uf Lin111/1tS Ill nature, which COUld be of phyJogc-
.haractcristic scale-like onumcm from whKh tng- netic import.lnc e.
394 Arthropods
Range in form, evolution ond ecology abducnon, i.e. by comJng togt·thcr :1nd 1110\ rng
(Ftg. 11.25) apart. they could not be swung to Jnd fro ~econd,
Eurypt~:nth are rclao\'l+y r.lrt' timtl... .1ml though the acntal mode of operatlon of rhe leg-. ".1s worked
th~: morpholu~"\ of '0111t:' gull'ra j, ""ell known, out from the posJttOn and nature of the JOillt)
uthl'!~ .1n: represented unh h} \'cry poorly pn.•c;crvcd between the podomere~. ln CUI')'ptends, as in
mall'ri,JI, .md tht:re ,1re .ttute t,I.\.Oilllmic problem' arthropod\ generally, joints .1re t.:itht•r hinges (with
111 cl.t~\llyrn~ such i~olatt·d n:nuim. Rather than ,1 ~111gk arricubtion) or pivots (with rwo arricula-
dcscnbing the ta.J~.ononlit pwbbm in detail, the tiom on oppos1te sides of the joint) The le~ of
ioUowrng dr.,cussron J) l·oruinc:d to a tcw c;ekcccd Ballt.Jt'llryptt•rrts had pivots pro'X1111.1lly ,md hmges dJs-
cvpe~ whtch ~hO\\ notJ.ble and pll''umabl} adaptrvc u11)' ••md l".lch podomerc could rock in one plar1e
dttfcrencc~ Ill ~tu. pro!>om,tl slupc. the loc:1tion of only rd.mve to the next one but, ~.·xrcpt wlth a few
the t: ye,, and the morpholog) ut" tht: bod), the 'Pl'ctaltzed JOint\, could not rot.ttc \urreN\'C JOUlts,
appcnd.1~es and the tc:hon. A lllrrem dac;\itication however, luve tht•ir arricuhuom wr m dttfcrcnr
(Srornwr. I 970-1 97-l) di"ide~ CUI') prLuds m to three pl.111c\ (F1g. I I .26a-("), that of th1.· coxa or tinr
suborders. e.Kh with as~oci.1ted !,uJULes ,lf1d super- podomcn: bemg ~ubvt·rtical.
f:lrmhec;. When the leg moved fonvard' rhe \t't of the
jo111ts .:mured that It w.h liH:cd up and ~er down in J
SUBCLASS EURYPTERIDA.
fut \\lard pmition for the ~urn:cding b:Jckw.trd
ORDER EURYPTERIDA.
SUBORDER 1 EURYPTERINA (t )rd.- l'cnn.): Ucner.1 v.:ich \troJ...c. lt IS probable that onl~ th~: l,\,t three
'null Ullt.h.>thL·d LhdJlll.tc, l' ~ Drqmnopkm<, .tppcndagc~ were acntall) used for w.1lking. mdud-
/l ,Jt,,t·uryrtum, ~tyi.•Hum;, .SimWIII•'· .\lt\•'PII'nl'. iug the paddle-like ~txth appendJ.ge
1/u~/um/l.n.t . The \IXth appendage. how~.·vcr, could also be
SUBORDER 2. PTERYGOTINA (Ord De:' .): EurypcenJ, ll\l·d l(lr 1'0\\c mg the arumal through the \C\ uer. a' ;m
Wtth t'nomtou~ dcnttutiJtc , hdn t•r w: c: g. Prtry,l!tll115.
an.tlyw• ofjoint patterns shO\n. If the paddle were ht
{."kt'/t>J'ItlliS.
SUBORDER 3 HIBBERTOPTERINA (C.trb.): Genera in
fimcuon "' ,m oar it would n~t·d lo pre~em a flat
whtch tlw po~t..-rior prmortt.•l lc:t-" Jt,,,.,..1 ha,,tJ e••tc:u- verllcal ~urface to the water when pu~hing b.Kk-
~tOn: c.~- /Jil>brrroplt'ttl>. <:'""l'YI•'"l'"'"">· w.ml,, but be hdd horizontally during the recovcT}
\troke for minimal resistance; tlm, 1n f.1ct, 1t ran do.
Th t:• Furyptcrina, by f.-n th~· l.tr~cst ,uborder, h,JVc
the srrcat<:~t drvcmty I lt~~J,,;/1,.,,1 1\ re!Jtively small
t()r the JOmts arc so arranged th.ll Jll Olttual rotation
or thl' blade occurs both .ll the prox.mul and rhc dll-
i
.tnd u111noditi~:d. "\/u/1<11/ld h,l\ a qu.1drare prosoma. tal JOtnt' of podomere 6. The bbde " thu~ out-
,mtcro-latt:ral eye' and .1 I.HcrJIIY ~·xp.lnded tdson. stretched and vertical dunng the backward
CttrilltMt'/1111 has a brgc di,nlld.ll prt•-J.bdomen wtth propuls1w ~troke and collapsed and honzontal dur-
a marked wam and a cylmdric.1l pmr-abdomeu and. mg the for.'lc.trd recovery ~trokt·.
hke th<: tiercdy am1oun~d \/r_,,,f'/trm. 1t has a tdson. 1lm rowmg technique has be~·n shown to be func-
lih that of 1 ~corp10n ..1pp.1rcntly mod1fied as a pm- ttonally eqmvalent to the scul.lmg Jt:tJOn of modem
'on 'pim•. The pro~omal .1ppend.1gcc; of Dolirlwptcms \WII11tnmg crabs (pottunids: PlolntL k, 19H5), and l!
.m.• ltf typic1l t{mn thou~h l'IHmnmtsly cxtended, ha\ b<:en c:tlcubted (Selden, l9H4) that BaltClt'llrY/Itrnt
whl•rca~ in Stylttrlt/niS .md lt\ rd:Hivc~ all the legs :1re could have re.Khed a ma.ximum velocity of 2.5 timn
\lendl•r and elong.He, even t lw c;i"Xth prosonu1 it\ body length per second. Ptcrygotids, on the oth~1
appcnd.tge. Jnd prcsunubl) moditied for llghtly h.md, bemg more hydrofi:Hl-shJped prubablv mti1cd
w.1lkrng over muddv \urftn'' .tt low spet:·ds with occas10nal bur..ts ol power, usmg F1gun
Ml,,t cUI")-ptcnd<> seem ro lu\'\:.' been active ben- the tclson tor ~tecnng. a style
thm. though ,\Ltively .rbk to \Wtm. Bafr,wt~ryptem> <,omL' euryprcnds ar lea'l rn.l) have been able to ondb
(glo I
ha' r~.-~.ently been the \llh.Jel'f of .1 dcuiled functioru1 cr.l'' I our onto land tor ~hort pl'noJ\. A largt= rr.ul tllustrc
mrd) b) <:id den ( 1QH l). \\'ho ll\t'd cxqui~itely pre- from the ~1lun;m of Rmgenke, NomJ.y, .,,;tb m· Wote1
'crvcd, 1\0IJted piece' of l'uryptcnd, mcluding plu\t' g:~tt p.tttems, seems to hav~· been made m dill
whole let->:~· fi·om localitic' 1n the Ihltic Silunan. way. lt IS believed to have been made by a Llrt,rc
First . it W;l\ ~hown that whtk the r.1di:~lly ;m,mged l\Jixoptcr11s (Hanken :1nd Stormc1, 11)75, fig.
LOX.tc tmtld rllasticate food hy adduction and 11 .25f-h). The trail is 520 mm lung and about 1611
(b)~
~ (c) Cttmopterus
(I)
(g) (h)
(f ) M ixopterus kiaerl
91re 11.25 Eurypterido. (a), (b) Pterygolus menonioe (Dev.): (o) dorsal view; (b) detail of chelicera (x 0.1); (c)Ctenopterus (Sil.),
atylonurid (x 0.5); (d) Hughmilleria (Si f.;~x 0.35); (e) Campy/ocephafus (Corb.). ventral view of prosomo, showing large metostomo
'lldbosol extensions on posterior legs typical of Hibbertopleridoe (x 0.2); (f) Mixopterus kiaeri (Sil.), dorsal and ventral views (x 0.1);
Jl o troil in Silurion red-beds of Ringerike, NorwC?)', believed to hove been mode by M kioeri (see text) while walking in the position
oo~~lroled in (h) (x 0.25) [(oHd), (f) Redrawn from S!Ormer in Treatise on Invertebrate Paleontology, Port P; (e) redrawn
from
o1eolon, 1957, (g) redrawn from Honken and Srormer, 1975.]
396 Arthropods
an
(b)
hy
0 2
p
.t
11
4 5
...
!> 6
~
6
p
pn
---..t---"''IIM.fl!-- qnathObase ho
leelh (cl ot~
typ
tiOI
Mi.
o[)
the
the;
I
tlOI
podomt>rf!s Ol.CI
Bar
(d) 196
care
nch
w:itl
de pc
Iago
:l~SC J
nuJlc
long
Devo
pod
Figure 11 26 Eurypterid lvncttonol morphology. (aHcl Ba/loevrypferus (Sil.). (a) exploded view of second prosomal appendage
showing hinge and pivots, (b) arrangement oF hinges and pivots between podomeres which Facilitates (c), For the sixth proso;;;.;i
appendage, an oor·like thrust dunng bockword stroke, and minimal resistonce to the water during Forward recovery stroke, (cf]
Parastylonurus (Sil.) m wolkmg posture [(o)-(c) Redrawn from Selden, 1981; (d) redrawn from Woterston, 1979.1 The.
gular
Cr
mm bro.td and t OINst~ of thrc:~: s~:ts of pJired parallel appendages held out 111 front of tt as a kind of cagt• fresh'
tr,tck' m;Hk up nf \l'p,lr.nc \tiCl'C''ivc imprints. Mixclpterlls w:.~ contt•mporancou' .tnd wa~ the nght inciU!
dccrea~mg m '17e mwards The ouccrmost 'A-tracks' size to have made ~liCh .1 trail. Presumably the shrim
.1rc hook-\h,lpl·d .1nd would fit With the imprint of curyptcrid cou ld .tlso sw1m. mmg the rctractabk many
the p.1ddk-\h.1ped t•nd of the 'iiXth prosomal sixth appendage :1~ a paddle. have:
append.1~t· Tht mli. rmcd1.He 'B-tr.Kk'' .md the Special :~d:~pt.mons for mbtlity in long-kggcd is sue
innennmt 'C-tr.ltks' \l'l'lll, however, to have been walking curyptcnds have been dl''>cnbed b) their
m,Jdt• b) appt'IHI,Jgt'' \'vtth A.Jttt·ned 'PII10\e end\. A W.HcNon ( 1979). c~pn ully m the Stlunm and se
long nwdi.m t--'TIH)\T 'l'l'lll' to luvl' been made by ,tylonurid Pamsty/cltlltrll.< (Ftg. 11.26d). Thi" genus recogJ
tht· tip nf tlw g~.·nit.ll lppt•nd,lgl' 'crapmg Jlong the h.ts ·' long prc-.1bdonwn .•llld 1t I\ e~nmatt·d that th<' wdJ
!-,'TOUnJ. Thc'e v~no\1\ l:Kl\ tit wnh the ronccpt of centre of grav1ty would havl lam near the wtdest (Buck.
the rr.1il being m,tdt· by .1 Mixclptcrm walking on the point of the :~bdomcn, at about the third prt· group
fourth, tlfth .md \IXth p.ur. of appendages. while abdomitul ~egmcnt. The hro.td, ~ctmltaHhJpcd ~tudy I
kecpm~ tht' ~r.hping 'cc~lnd .tml tlurd pair\ of Range' (cpimcra) o( the po<>t-Jbdomm.ll segments In ~
Phylum Crustacea 397
.md the broad, keeled [(:)son would have acted as a gcnc1ty of the phylum has mually been a~rrcni, for
hydrofoil, giving lift to rhc pmtcrJOr part of the ani- in all ca'e~ the embryo, developmg by ~piral lk.lv-
nul, and theu:by increasing \t.lbllttv m light currents .tgc, \Ub)equently bc~:omes a nauplius larva. lorn-
With nunimal mu\c ular dTort. mon to all except the most specJahzed groups. Th~.-·
Such generalized type' \' Ha/t,wurypffnl~ were nauphus is a 5mall ovoid larva, •nvanably ~ith thre<.:
probably umpeoahzcd fct'del"o Most euryptends, pdlt"'> of appendage~. These an· the uniramum
however, were probably prcual<m, their prey being antcnnulcs, the b1r:unous antennae and the
other euryptcnd, and fishc' Two different fccdmg mandibles. In later developmental stagt''> thc~t·
types were denvcd from tln. un,pcciahzcd condl- mand1bles become differentiated J\ rruc jaws, but
uon, the enlarged \putme lunb' Tl and Ill of they arc used in the nauplius along wnh the uthn
.\1ixoptm1s. and the grc.1tly cnbrged chelicerae two pall'\ of append.1gcs for ~wummng .md lcedmg.
of Pterygotus. Thl· laner were probably kept folded as The shape and movement~ of the naupl1U5 were
the animal cruised around lool..ing for prey, and descnbed in memorable ccrrn' by Garstang ( 1951)
then rapid!) extended for c.tpture. ' fhc nauplius i\ a wobbly thmg, .1 head Without ,\
In some ca~cs, characten\ta euryptend a\~ooa body. He flops about \vith foolt\h JCrks, a regular
nom can be: dmmgmshed .md related to cnvlron- Tom-Noddy .. .', f(Jr the nauplltl~ IS invanably a
mcntal conditmns. In the Stlunan of the Welsh swunmmg entlty, .111d only after sevcrJl other devel-
Borderland, for example (Kjcllcsvig-Wae nng, opmental stages, which differ in the variom m,nn
1961), there: ~eem to be three \uch a~\ociattom: (1) groups, ts the adult fom1 attaim·d. When the Jdult
arnnosom.1t1ds and pterygouJ,, a~sociated \\1th a stage 1s reached, the crmtacc:m rct.um the antt'n-
nch, norrnll manne fauna; (2) Euryptcrinac found nules and the antennae, and the mandible\ .tre
wtth rare manne fosstl~. 1n ~ediment\ probably JOined by two ~Hher pairs of fl,od-prorc~'lng
deposited m :m mshore envtronmcnt, probably :tppendages, the maxillules and maxillae. The~l· five
IJgoonal or cm1anne; (3) bradu\h to freshwater pall"o of differentiated append.1ge' belong to the
1\\Cmblage~. dommated by stylonunds .md hugh- head, behmd wh1ch there ts u~ually .1 thorax .md an
milleriids. Most representatives arc les!> th.m 20 cm abdomen. These three ragmata (he:td, thorax and
long, but there Jre \cveraJ gtJnt fomts mduding the abdomen) arc pos!>t:~scd by mo't cm,taceans, though
Dcvonian Ptrry,~Mrts (Erettoptcms), the largcst arthro- not by ostracodes. Likewise provtded with append-
pod of all time, which reached nearly 2 m m length. ages, the head may be fused with the thorax and
commonly the two arc covered by a saddle-like
dol"oal ~hteld, the cJrapace, wluch may be free.: po,te-
riorly, as m many shnmp-likc ronm, or, a~ in l r.lh\
and lobsters, may be fused to the body. Despite
The .trthropod Phylum Cm~tacea (Fig. 11.27) IS sin- these similanties there is some pombility th.1t cms-
I(Ularly div<. r;e and h1ghJy succe\sful. taceam are not a natural but a p.1raphykttc i,'TOIIp
Crustaceans an: matnly nunne but there arc \ome (Brigg; :tnd Fortey, 1989), and tl11. pmition of
IR-shwater and a few terrestnal groups; the phylum Cambnan cmstacean-likc arthropotb both from the:
Uldudes not only the famthar cr:~bs, lob~tcrs and Burge~~ Sb:tle and fTOm Swedish 'Orstcn' i~ still
Wimps, but al~o mtracods, copepods, barnacles and much debated.
mny other t<t.xa. Although crustaceans nonnally The classification of crustaceam i~ based on many
.1ve a ch.itinou~ e .. mkelcton, their range and variety characters, which mclude the fom1 of the body,
1 'uch that some b'TOUp) (e.g. adult bamade~ with number of segments, presence or ab,ence of a l ar,J-
bc1r pyramidal external ~kckton of calntc plates pace .md in particular the nature of the appendages.
tnd \Ornc of the parasttic taxa) Jre not inuncd1atdy E1ght classes arc recogn.ized by McLaughlin
recognizable as Cm\taceans at all Fossil barnacles are (1980), ranging from the t111y pnmltlvt·
wrU k.no~ n, especially from Tertiary deposits Ccphalocarida to the largest aud most important
:IIuckeridgc, J9R3); mdt'ed they were the one group, the Malaro~rraca, wluch •~ the only da''
t- ;roup of wluch Charles Darwm made a t.lXonomic (othl·r th,m the o~tracoda, usuaiJy treatcd a~ mlcrn-
d tudy before embarking upon h1s Ori~in t?[ Speri1's. fo~\ih) with a good fossil record. Only four d,1~~cs,
S, In spite of thi~ divel"oity, however, the homo- however, (Rcm1pcdia, Malacostraca, PhyllnpmL
398 Arthropods
(19~
ro a
SUBC
la~e
chc •
JlllC:I
powt
(a) CMatiourl• (Phyllocanda)
.mJ
JhJ01
v1Jed
le.tt:.t,
PO>>il
lo.:.1ri
M.'rvct
Burgt
arc qL
abunr
daUy
tJ.(S. ~
(Fig. I
75 ern
.1bouc
1\'tba/i
SUBCL
mda)'•
i~ J ma
rapron
l.u. du
been n
SUBClJ
of Cr:t~
other<.
(c) THHiocarls IEocanda) (d) Ac.ntlto~illa (Hoplocanda) (• ) Eryma (De<:.ipoda)
dctatl I
>ummJ
Figure I I 27 Molocostrocon d1versity· (o) Cerotiocoris (Phyllocorido; SiI.; x 0.5), (b) Poloeocoris (Syncorido, Corb., x 2), lcl VMIOU\
Teolliocom (Eocondo, Corb.; >< 1), (d) Aconthosquillo (Hoplocorido; Rec.; x 0.25); (e) Erymo (Decopodo; Jur.; >< 0.5.) .•~ tlcti1
much c
SUPE
rat!H
anJ •
and MaxillopoJ.t), .m~ recogmzed by Schram (1986), mous thoracic limb~. unifo rm .1nd unspeculizt'd 10
ingly
who used com puter-based cl.tdbtic methodology. the m on.· primitlVe groups, but oftt:n becommg ,.,ns,
In view of the current Attx m crust.Ke,tn classitlca- greatly dilferentiated 111 the more adv,mced malacc)l- h,lS l:
tion, I here me Lbe more conservative classification tracans. Many of the more successfi.tl predators haw I'~IJr
of M cl :tughlin chelate or pincer-like appendages, .111d indeed much for 1
In the M.tlacostr.tcJ there are generally paired of the success of the malacosrracatlS ts lmked eo eh~ .-lttllSJ
compound eyes, .md the antennules and antennae d.iflerentiation of the umbs for V<trJOU~ purpo~~ nvcn.
lrum
are biramous, the latter havmg a $htdd-hke ante1mal feedmg, sw1mnung and resp1rat1on. The m.Uacostra-
SUPEI
scale or ~caphocente. I he bttmg J.lws (mandibles) can abdomen comtst~ o f ~tx segm en ~ bt>anng five drvcr
are fonned from gnath obases , mtemal extensions btramous pleopods wh1ch are o ften well develop~ .. V.'llll!
of the mandthuhr .tppend.tge that meet in the and used in swtmm tng (though they are lmt m wm~ ek,ng
median plane of the body .1nd roll or gnnd together groups). The last ~egment h.1s J tnangul.u CC'ntnl under
as the ltmb move' hJl kwards .md forwards. The telson Aanked by pa1red b1ramous uropod~. tl.~uet
body
outer part of the mandtbubr ltmb may dtsappear. In Accordmg to M c l aughlm ( 19XO) there .ue th' c
1
[01'\.
the MaiJcmtraca then' .m: u~u.tllv etght p;ur; ofbtra- m.uacosttacan subdJs~es (followed here). Schnm
Phylum Crustacea 399
(1986), on the mhcr h.and, rt•movt:~ the Phylloc.1rids ew .lppt.lr.Htlt' (F•!<· 11 27c) though they h.wt' lOmp.u-J-
ro a \eparate cla,~. ovdy uuspeumtc:d .1ppc:ndJ~c:' whiCh ,~re not prov1dc-d
with cheldt' Tht· eotJnth Wl'rt' undnuhredlv .mc(Str.U to
SUBCLASS 1 PHYLLOCARIDA {Cam-Rc:c): In tlm group a
all the mu<km t nlslJCl'JII !,.'Tt!Up~ fhc'e IJttn m dud<' the
l.trt,'C. blvJlvnltJrJp.l<t', \\ICh .1.11 .lrtlnlhtmg rmtrum. cover.;
followmg two supt'rord~:f\,
the lntc:nor pan of the bud\ hut 1\ nut fu~ed to it. The
SUPERORDER 3 PERACARIDA (?CJrb./Pcnu.-Rn .): Com-
JJllennae Jnd lntennult·\ art• shun. wlulc: the: nundible, ii1m1
pme,, :unong oth..r group,, rh,· my-o•J, (c,mdotd). I'Opod;
powerfi1l biung J·''''' Uehmd tht·\l .lrt the: c•ght btr.llnou~
(dor;o-vcntr.llly tf.merwd) .tml Jmplupud' (btl·ra.l.lv tfJ.t-
and tohaeemh thm.tu•J'•"" u,,·d m h.ltcr tccJmg. The
tcned) Thcv arc generally rJthcr 'nuU u u,t.u.:eJm wuh
~bdomcn pmJt'cr' pn,ll·nNly lr••m tlw CJrJ.pacc; it '' pro-
unnunerahzcd ruri,·lc,, .llld hen<t' With J rJLhc:r poor fu>-
,,ded \'vlth up tn "" tppend.lgl"'> .1nJ temllllJtCs 111 a pltr of
s•l record. The tl:nulc-; h~vc .1 brl'I<XI pou(h or rno~nupium
leat:hke p!.ltt'\ (furca l flw c:.trht"'t kuuwn phvUocand IS
ui wh1ch the c~" dcvdop .md '' hc:nc.:e the: young Mt'
po~s1blv Lower t .unbn.u1 m .1~e. E'-tJlll\ltd) preserved phyl-
rt'leased a .. llllm,\ntrc adults.
locand-l.tkc c ru•t.tce.ul,, C~m~<l•i•~>f'" 111J p,.,~ric.ms, with prc-
SUPERORDER 4. EUCARIDA- In tlm <.tt<·gnn come the grc:.!t
>erved .lppend.tge' .lrt' kno\\11 tmm tht' Middlt· Cambrian
llldJtlrtf) of common modnn ausL.In·.tm. divJtlc:d into
Burgc:s> ~hale (tln~J:!!S. l'J7H) .111d ~e11ero~ 'lU.:h ,1., Hym~nocans
two mJul oldt•r>, ORDI:.R LUI'IIAU'-IA\1-A (Rec.)
~re lJlllle tUnunon 111 the l;m C.1mhrt.tn l'hvllot.trid, 111ay be
Jnd ORDER. DECAPODA [rna~.-Rl'r.). Th<'
Jbundant in ct·rt,un bc1es m tht eHly p,,l,,t'ozotc, and c:\pt:-
euphausiid' .m: tl<l'alllt' ~wunmm).( ond01d~ nf cm-
cially in 111Jr!{lll,tl lll.lrllll' .1ml po~stbly br.1cki~h-wJter habi-
mopolitan distnbutwn which Jrl' vc:ry •mporr.mt in
IJt>. Some: of thc:Sl' ~rcw very brgc.-; the '>iluri.lll Ccr111iowris
tod.1y'~ oreJII~. hut wuh J m:p:Ji~blt• t(lsstl rt'cord. The:
(Fig, 11.27a), ti.1r r:xJmpk. h.ul some rep1 "'e11t.1tivt."s reJching
dt·capod~. whidt .tll h.tvc five P'"r.. of thoracic
71 rm in length, '~hcrl·.t~ tht' common rr:ccnt '\lt·f,alia ,, Oltly
1ppcnd.1gt'~. indudt· ~Wi11u1un~ tom1' (\hrunp~ .u1d
.1bout I cm long, md dtl lar~l'\t modem phyllocarid
.\'ebctliop.il.\ i' on!) 4 lilt 111 lt'u~th . prawns) ·" well '" Lwntllll rl'prt~t'lltJUVC\ (the crabs .111d
lobsters, Jnd the frf'\hw.IL<·r rrJvti\h).
SUBCLASS 2 HOPLOCARIDA (l.1rh.-Rr:c): ReprNented by
rodav·~ 'i.Juil/,,, rhe m.mt"-'hrimp. wlud1 hk<' :ill hoploc.Hid,
IS l mJnne prcd.nnr with J dhti.'ntl.-.1 ~bdum<'n dnd modified Shnmp-hke cmstac.:can\ of Ji,tHlltly modern type
t•ptonal rhoncopoth (< I. f1g. 11.2id) Some ~em:r.illv simi- ('cocarid~')
tir<:t appeared 1n the l..atc Dc.:von..m .md
!Jr. though 'nullcr ti1nn~ trnm th<· l Jtbom1t:mm have .i.l•o
underwent a lllaJOr adapltvc rJdi.Hion in the
been referred ro th" ,uh,l."'·
SUBCLASS 3. EUMALACOSTRACA (nn -K,...., ): The VJ\t .may Carbomternu' They arc umally L'llllnd tn 'cd1mt:nts
llf 'rab~. lobo.tcr;. shnmp,, my,~t·k I'>Opu<h.•unph1puds .tnd of nurgnul-mannc and br,td,J,h-w:m:r ong:m
•>thers which mak<· up tlm !,'Tl' 1t grnup 1r<· tno numcrom tO (coasul pia m .md lo\\ -lv111g ~wamp utvtronmcJW•),
leuil here Mel:.lu~;h.lm ( I'JkO) pro\'t.fc:, .1 good modem ~inccit i'i on I} the,e cnv1ronmt:nh \\ htth pro\ided
•umm.ll) Jnd •llmtr.unl Jncnpnon' of reprc,cntJttv(S of the excepuonJI nrcumst.utcc' rcqturcd for th~:
nnom gruup\. There •re f.tur L'Um.J!acmtraun <upcrorder; . preservation of thetr utumncrahzcd chttll1llUS
.t~ detint'd hv h<·r. but cuuui.Jco,tl".tCtn t.txonom~ " >till
murh dcbatl·tl exmkdcrom. ft 1~ ,tctually qune ltkdy that \hrimp,'
SUPERORDER 1 SYNCARIDA (M. C.1rb.-Rec.): Small, were common m conrcmporaueou~ sc.1s. but wc•rc
rather clclllg.ltcd <fll\t.ILt',lll\, pnmJnJv l.tckmg .1 tanpace, not preserved (Hnggs and Cbd,,on, [CJWJ, 19Wl).
I •
dnd \\1th b1r:trnum rlhtr.lnt .tppt•ntlJgc>. They h.we seem- These P::al::teo?OlC fonm were l:lrgdy ext1nct by the
ingly evolved by ,, ,hffl'r,·•H p.tth to orhrr cum.llacosrra- late Penman. The .appe.mancc of c,lluiied cutidcs in
cart~. m which rlw l.tr.tp.tte "prun.tl) Jnd. Jf not pre•em. Mc~ozoic marim· cuc:1nd~ allow~ much grc.:ttcr
ha~ b<.'<.'n ~<'Cond.mlv Inst. Sv~tl.lrid, tk1umhed m the IJtc
preservation pownt1.'11.
PabcozOtl (hg. 11 ..:!7h), .11;d \\l"rt' only k•tuwn J~ losuls
for many Yl',\f\. In IH92, hll~~e..,er a h"mg genus,
All c·ucarids hJ ve the carap.1ce fi.1sc·d wtth the
l
Anaspidrs, w.1s disrowrt•d l.vmg 1n T.151JIJ.JU.lll bke, and thor:lcic somite~. The compound eye~ ,1re stalked,
nver<., .1ml ,, few mhn ~l'lll'r.l hJvc- \illle been reponed and the ern:,rs ,1re earned :Ht.lclwd to the abdonunaJ
&urn the ..1mc art .1. appendage~. Th(· young, pnmat1vcly h.nched as nau-
SUPERORDER 2 EOCARIDA (M. 11n ·Pern1.)· A rc:lativdy plii, may undergo v.1nom frcc-,w•mnung tr.ms-
e divene group of \IIIJU, \hnmp-hkl· m caridoid form~.
j fomlations, and arc often of hi1 JJTt: appearance
S\\,mnung <rmrJn·.tll• 111 ''h•<h dtt•rc " J cuaplle, an
before bccommg adult,. Lobster\ .md crabs have
elongatt'd budy wuh tht· ~lxiumcn oticn partly n1rled
under the body ..md ,, wl (ju <ompmt•d of the rdson and appendages .:tnd .trc· htghly 'iuccc•s,fiJI marine preda-
flattened uropoJ, wlut h .au b,· rlpuily flu: ked undt'r the tof'>. lt i~ probable dl.lt the pos'c~sion of such grasp-
bodv <o dlJr tht• tnmlll d.lrtl'll out nf the w.1y of preda- ing pmceN (ab,cnt m the1r P.llaco70Jc prccuNof'>;
c
tors. The c:ocan<h .lre lutihh or~tJmn·d .llld oi verv mod- Fig. ll.27c) ha' been the mo~t unportant smgle
11
400 Arthropods
factor in d1c1r \UCCt''' · Tht y .Ire the l.:rrgest manne LcVl-Setti. R . ( 191J3) l'ri/tlbrtt•s: et Plwto)lmplric Atlru. 2nd
cru~taccans hvmg todJy .md. \IOCC their skeletom cdn. Univcr.1ty ot ( h1cabro Pre~~. Chicago. (LJrgt·
arc rcmfon:ed "' Hh '.1lnum carbonate. they have a numbers of full p~ge pl.1tc~. Wtth di~CU\\t011 of eyes)
good fos~il n:wrd Although \orne Triassic genera Mc.Laughhn, P . (1980) Co111parnfiP1.' l\.-torplrol~ '!f R11tlll
arc known, thl· fiN n.·.tl bur.t of .-.dapnve radianon Cmst<lfCfl. Freeman, San Fran~•,co. ( lllu~tratiom Jlld
damficanon~)
was delayed until the e.trl} Jura\\tC. Most of these
Manron, ~ M . ( I Y77) 17rr -irtfm,pmL<: H.tbits , FmltiMtlll Ur
were of rather luh,ter-hke fcmll. though some, such
Morphohl.fly a11d Er,olutiOII Oxfi1rd Uruvc~lt) Prt'\,,
as the Eryomdac (l g. f1g 1.2.10), had dor;o-ven-
Oxtord. (Dcfimovc work on arthropod morphology
trally comprc\scd ldr.1pacc' Mmt of the Jurasstc and cvoluoon)
group\ have ~unwed to the pre\ent day but ~ome ~oorc, R.C (cd) (1955) Trt·att>f 011 lt~r•t•rtrhhllc'
have dccl•ncd 111 number\ The cryomd~. for exam- Palt'olllola.i/Y Part P, Artlrr<>pvdcl 2. GcologJcal Sonety of Dn
ple. wh1ch formerly inh,1bited 'haJJow waters, are Amc;-rica Jnd Umvcrs•ry of K.ln'kl~ Press. L.1wnmte,
known today only J\ bhnd 'pccil.!s from abyssal Kan. (Chcl iLcr.ttl·~. pycnoKomd\ and sp1ders)
depths. Although •noucrn lob,ter' an~ widespread Moo re, R.C. (cd.) ( L959) Trealut.· 011 luvntchr.rtc Uu(
and sucu:ssful, they art· represented by many fewer Paleontola.qy P.trt 0, .'1rtlrropoda I .Gcolog.cal Sonety of 'I
genera lhan the c;.rab~ (Brachyura), in which the America and Unive rsity of Kansas Press, Llwrence (
abdomen i~ very reduced .llld permanently turned

Kan. (Trilobites .111d related group~) IK.teslt:r, R.L. (cd.) C.11
(1997) Vol. I or the revised version ts now published. I r
under the thorax. Tlll:'\1.! originated in the Lower Moo re, R .C (ed) ( l (j(,J) Treatise ou lrwcrtrlmltf /•
Juras~ic and have expanded Jnd dtvcr;ified ever Palemllology P:m Q !Jrtlrwpodo 3. Geological Society of I•
since, having many kmc:b of .1dipuvcly specialized Arncric.t and University or Kans:b Prcs~. WWTC:OlC, Can
characterisncs llttmg them for lJfe m innumerable K311. 01
habitats, and rhese tan bl.' \tttdied from the Mesozoic Moore, R.C. (ed.) ( 196Y) Treatise 011 lm~·rt.-llr.llf fr
onwards and c~pco,dly through the Tcrti.1ry. From Paleontolo,Ry P;ut R, .4rtlrmpotls, Vob I and 2 Can
their Devono-Carbomferom origim the cumalacm- Gcologtcal Souety of America .rnd Umver.ll) of d'
rrac~m have t'xpanded tnonnously into diverse Kansa..~ Press, Lawrence, Kan AI
enV1ronment~ a \tnkmg example of the rad1ation of Schram. F.R. ( 1~H(,) CntsldCI.'cl Oxford Umwrsiry Pr~' 0
Oxford. en
a group 111 wh1ch ' Uauplan' .md physiology com-
Whittington, H.U. (ICJ'J2) Tn/o~rtl's. Fosst.ls JJlum;~tcd ·! Chat
bined the reqmred degree of t:liolutionary plasticicy
Boyddl Press, Woodhndge (120 phntob~T.~Ph•c pl.tt~) Sil
ro aJJow them ro become one of the most successful
life fonn~ m the \CJ roda>
w
ChJ.tt
Individual papers and atfler references Ot
M,
Bcrgo;trom, J. (1971) PalatomrnH - merostomc or nu:ro.... 17
tomoid? Let/rain 4, 3<)3-401 ChiUJ
Bcrgstrom,]. and Bra~~d, C. (l~H4) Leg. in the tnlobicc of1
Rlrenops fmm the Lower Dcvonian ll unsrck Slatt VlrC
Uergstrom. J. (1973) OrH.\IIi,auon, ltfe .tnd systemati c~ of Lerlroia 17, 67-72 Chl ur
rnlobttes. Pom"Ls a11d Srrara 2. (l:lrge-scale work; pre- Drezinski, D.K. (1985) An opportun.u.oc Upper a~se
sent~ a different clJss•f•catJOn from th.lt in the 'Trt·atise') OrdovicJan tn lobrtc aS\emblage from Mtssouri. u·thora Ner
Uowes, D .R .•md Watt.•r.ton, C.D. (eds) (1985) Fosstl 19,315-25. Cisnc,
Arthropods as l.Jllit~ct / lnimals. TratiSactrom of the Royal Driggs, D .E.G. (LY71:!) The n1orphology, mode ofl.tfe md sh1p
Societ>· ~f J:'di"l>"~l/lr 76, 1o1-3~':1 (24 pJpcrs) affinities of CarrodtlSpis perferta (Cru~tacca: PhyUo- rddi
Ungg;, D E.G .111d L.me, P.D. (c(b) (1983) Trilobites arul carida), M iddle Camhnan Burgess Shale. Bntt~h Cl ark~
Otlrrr t4rtlm•potls SpeuaJ PJper. m Palaeontology No Columbia. P/ri/omplrwrl Tmnsmtiom •"!f tire Royal &•crrty so m
30. AcademiC l're", l ondon (16 papers) <if Lorul o11 B 281 , 439-87. Cl arks<
Clarke, K. U ( 1973) '17!r Bit>!.wy 11{ ,frthropods. Edward Briggs, D .E.G . 311d Cl.trkson, E..N.K. (19159) Envmm· SiiUI
Amold, I ondon ( hort mnple text) ment.1l control\ on the t.aphonomy and distnbuuon et 329-
Gmt.ang. W ( 11J5 I) Lm•al I :<>m~> , 1111d Otlrrr 7..ool<>glllll Carboniferous nubco~tracan cru~taccm!\. Trc~ruachoru cf odoJ
Verses Blackwcll, <hford . the Roynl Society of bdr11buf'.l/lt. I:artlt SrittueJ 80, 293-\IJI Cl.1rksc
Lebrun, p ( 19~5) 1 n/,ohrtr<. ,\fmrr.:lll.\ rt rossl!t'S ) Bngg;. D.E.G. Jnd CIJrk,on, E.N.K ( 1'190) The btr of (
EdJtiom CEDIM, Pan\. (nccllcnt colour plates) Palacozote radJallOll or m.tlaeostraC<.Ul crustace~m. Ill Hurt'•
Bibliography 401
AflltiiT I:r•c>lrttioii<Jry R.JclrcJIIrll/1 (1-d, v.n. I lylm .md G.P. Cbrkson, E.N.K. (!IJ73) Morphology and l'VUJUUOJI or
Larwood). Systt>matln A~\ucwion Spl:oal Vol. 42, the eye in Upper C rmhn.m OlenicL!e (T rilobna).
Clarcndon Prc~\, Oxfi>rd, pp l C>5-R6. Pohtt'OIIIolo,l/y 16, 735-63.
Uril(h'S, D.E.G. aud fortl'Y· RA (lYIN) l he early radia- C IJ.rk.\Oll, E.N.K. (IIJ75) The evolution of the eye m
tion and rclauomru1)~ nl the maJor .trthropoJ )o{l'Oups. trilobttC\. Fossils a11d Strala 4 , 7-31. (Swnmary .1ntl btb-
'lnt'llrt 246, 241-3. hography on eye funcuou)
Bn~. 0 E.G. and Furtq, R.A. ( JIJCJ2) Tht· early Cbrbon. E.N.K. (1979} The vt~ual s~tem of rnlobttC\.
Cambnan rad.Jauon of Jrthmpods. m Orlf(lll mu/ Early Palarotltoh~y 22, 1-22. {Mo't recent ~ummary)
br'<!llltrl)ll r.if the 'lfrtcJ,•'" (c<h J.H . Ltpp,, and P.W. Cl:ukson, LN.K. (19HH) The ongin of m.trill(: mvcnc-
Stgnor), Plenum Prc\\, Nnv York ami London, pp. hr.lte specie~: a cntical review of microevoluhon;~ry
J3(r-74. tr~nsformations. ProCI!t•drii.I!S of the C'J~?oiClgisrs • A ~scJd11tiun
13ruton, D. :md Haas. W ( IYY7) Functron.tl morphology 99. 153-71.
of Phacoprnae (Tnlobrt.t) 1nd the mcch.11uc' of cnroll- Clarkson, c.N.K. and H enry,] L. (l973) Structures co.ap-
rnent PalacotlltiJ!raplum A/1/ri/un.f{l'tt -l 245. 1-U tatlvcs et eruoulement chcz quclques trilobtte~ orJovt-
Buckcndgc J.S ( IIJHJ) 1"-m,tl bamacl~ (Ctppipcdia: ncm Cl \tluncns. Lrtl1111a 6. I 05-32. (Enrollment and
1 horacica) of New lnl.mJ and Ausrralta 1\'rw L<'aland mterlodang mechantsms)
Gcolo,i!IWI Sun1cy flahtt'Cllll!>foi(IClll Bul/eun 50, 1- 151 . Uark~on, E.N.K. and li.'VJ-Scttl, R. (1975) Tnlobl[e eyes
('.unphdl K.S.W. (11J7"i,l) The functional ltMtomy of .111d th.: optics of Des Cartes ;rnd Huygem. Naltrrt' 254,
phacoprd trilobl[e<>: mu,cubnm: and eyc·s. .Jt~umal and 063- 7. ( Lens structure .md function)
Pmrf'tdin.f{s r.if the Rt>ylll SI>Cit'IY f:!{ New S<>utlr I!titles 108. Clark~on, F N.K., Eldredge, N . and Henry. J.L. (I'J77)
168-88 (Lens struc.:turt;) ::iOllll' Phacopina (Trrlob1ta) from the Stlunan of
C.unpbell, K.S.W. (1975h) The function.1l morpholobry Sc.:otl.tnd. Palaeo/Jtology 20, 11 'J-42.
of Cryptolitlws Fo5stl( mul StratcJ 4. 65-86. (Tnnudcid Cock~. L R.M. and Fortcy, R.A. (1988) Lower
tnnge function ..md ichnulugy) Paheo7oic facie~ and Eaunas Jround Gondwana, m
C~mpb.:ll, K.S.W. (l'J77) fnlolmc<> ofthe ll.lra~;an, llois Gomlwana a11d Tethys (~ds M.G. Audlcy-Charle' ~nd A
d'Arc and fn~co l~om1~cions (Early D cvoman), Hallam) , Geological Soctery o( London Spcctal
Arbuckle Mountaim l~c){io n, Oklahoma. Bulletrll q{ the Pubhc~tion No. 37. pp. l83-200.
Oklalwma Ccolo.~tiull \mvry 123, 1-227. (Eye~ and Cocks, L.R. M. and Fortcy, R..A. (19HH) An:mg to
cnrollmem ~trunun·\) Llandovcry fauna! d1~tnbutions m the Caledorudt·'· rn
Chlrtcrton, B.D.I:. ( \971) T;n;ouomy .111d umngeny of 71tf' Ca/t'doman-Appaltulua/1 (Jro.~m (em A.L. H.um .md
)rluro-Devonian tnlobttt.~ from ne.u Ya,~. Ne\\- South D.J . Fctu~s), Geolog~cal Soctct) of London \penal
Wale,. PalaeotliOJ:rcJplllld 137, 1-108. Pubhcauon No. 38, pp. 22J-4o.
Chatterton, B.O .E. ( llJHO) Ontogenetic studlt\ l)f Middle Coch, L.l-t.M .•md Fortey, R .A (1990) Biogcojo{raphy uf
Ordovician trilobitl.'' front the E5battaotinl' form:mon, Ordovtcian and Siluriau faunas, in l'lllaeozt•ic
M:rckenzre Mounwius. Canada. p,tftli'OIII<J.!!rnphim .'\ Paftrt't!f!t'o,!!raplty and Bio,I(CDJ!raphy (cds W.S. M cKcrrow
171, 1-74. (SiHcificd tauna •llusrratcd) and C.R. Scotcsc), GeologlC:II Society Memo•r No
( hlup.tc, I (1983) Tnlobtte a.N:mblagt~ tn the Dcvonian 12. pp 47-104.
of the Barranctian arl'l and chctr relattom to palaeoen- Cowte,J W. {1971 ) Lower C.1mbnan faunJl provtnce~. Ill
vrronmcnt. Gt:Cl/o~ic,J rl Pa/clf'Otl/t>logtut 17, 4S-73 Fatmlll Provin«s in Spare aud Timf', C~ological }t>tlmal
Chluplc, I. (1987) Ecmtrangraphy of Silunan trilobite Spectal ls~ue No. 4, pp. 2 1-46. ( Recent sllldy uftrilo-
assemblages of the lhrranctian area, Czecho~lovak.ia. bne provinces)
Nnvslerter itt Strat(l/rlfJ!It y 17, I 69-86. Dalingwater, J.E. (1973a) The cuticle of a curyptcnd.
rtsnc, S. (1975) Anatomy of Tt1ttrtlmo Jnd the rclation- Ll'tlllfia 6 , 179-86. (Morphology)
'hips of the trilobttes h1ssi/s and Strata 4 , t 5 64. ()(- Dalmgw:~rcr, J.E . (l973b) Tnlobitc cuticle nucro\tnrcrurc
radiography, with 11mb mnrphology) J.nd lOlllposition. PalaeomoiCJ<ily 16, 827-39. (F111e ~true
Cl.trbon. E.N.K. (I'Jfl6) \cht7nchroal eye<> and vtston of cure)
'omc S1lurian aca~t1d rnlobrte~ Palm'<llllt>I.~!!Y 9 , I ·29. DahnJ.,>W.lter, J.E. (197'i) Further ob~crvauon' of
I'IJrkson, I:..N K. (ICJh1)) A functional \tudy of the euryptend cuticle~ Fomls rmd Srrata 4, 271-80.
Siluri:w trilobite Ll·mttllflis dt:{ltxa (Lake). Ll•rlrtlla 2, (Eutyplcrid cuciclcs were \lhtcous)
329-44. (Lateral vtl'W .1nd tunction,1l evolution of Dalmgwatcr, J.E., Hutchm,on, S.J., Mutvct, 11 and
odontoplru rids) Swctcr, D .J. (1991) C..utieular ultra.~tmcturl' of the
Cl.lrk\on, E N.K. (I'J71) On the early ~dll7ochroal eyes tnlobitc Ellipsocepltalu.s polytomus from the M1ddlc
of Onnatltops (TnlobttJ /.dlszkellinae) ll,ft·mom·s du C.unbnan of Oland, \wedcn. Palm·ontology 34 205- I 7.
Hun·m1 dr Rrchfrrht.., Cl'l>l~tqun tl .\111u:r,s 73 . 'i l -63. El dredge, N (1970) Obscrv:~uons on burrowrng bcluv-
402 Arthropod s
tour tn Lim11/m ptl/ypii<'IIIIU (( lu:lt, rrH.t, Mt>ro~tonut.l) Spttlbcr~l.'n , I. Olcnidae. Sorsk l'c•f,mllstrtw Skr!tur 11
with unphcauom on the lutKIIun.tl .m.ttumy of tnJo- 160. l - 12H.
btt~·~ /\'oi'Hralt's o( rhc Amrnum ,\twwm 4 ,\'nrw,ll 1-ortcy. R. A. (1':175) Early Ordovtcldll tnJobtt<' conmm-
H1s1oty 2436,1-17. (Valu.thle stud) ol lt"mg Jmmals) nitic~. Fossils and Strata 4 , 331-52. ( rlm:t t>ll\hurt'-Otl- lie
Udredgc. N. (1972.t) Parrems of rcph.tla mu..cuiJture in 'horc bcnth tc and one planktomc ,ts\c:mhl.agt•)
rhe l'lucopin~ (Tnlobita) .wd dn·it playlogcnctic stg- Fortcy. R .A. ( I'JMS) Pelagic rnlobitc~ .1~ ,\11 cx.uuple uf
nJflcMlt:C. Jcmrtl<ll of PtTicoutoh~I!Y 45. 52-67. (Rcron- deduc ing t he life habib of c.:xtinct .Jrthropods.
structwn of internal organ~) T tcl/1.\alfi{)m of the Royal Socit'l}' 11{ hdwbur~h: E1rth 11 ~
f:ldrcdgc, N. (I 972b) ~y~tcnl.ltl(' ,111d evolution of Somc't'.' 76. 2 I IJ-JO.
PiltiWJ'i rcma (Green 1~32). ;uu.l f>f~tluJps iowmm (Delo Fortq:. R .A. ( IIJIJO) Ontogl'ny. hypo\tome anachment
I 935) (Tnlubtt.t) from the Mtddll· Dcvom.m of North ;md tnlobate d.l\\lfication. P.Jiat:t•rllt>lo(!}' 33. 52')-7(,,
Amcnu. Hu/11'1111 "f rltr Amrnuw \tu~cum tij' Nallmtl Furtcy. R.A. .md Chatterton. B D .E 1lJXli)
111<1<11) 147. -l'+-113 (Allopatnc cvnluuon) C!Js,tftt JtJon of the niilobJte 'uhonlt r A,JphmJ. f ie
Eldrcd~l, N. (197-l) Rl'vt,ion oi the ~uborder Pa/dl'<lllllllc!(1y. 3 1. 165-222
!:>ynztphmunnl (Chdtrl'r.UJ, Mcro,romat.l) wtth Fortt'y R A .md Mom~. S. F. (I <J7K) DJ,covcn nl n:tu· t
rcnurb on merosrom(· phylogt·ny \;,wiriate.s of the phu~-likc tnlobJtC brvac. Pohu·omoln.I!Y 2 1, X2J- 33. (
,-lua'tlul/1 .\tust'lllll c•f 1\/,1/ur.ll Histvry 2 543 . 1-n. {Pre-proCl~pl~ •t.1gcs) I lc1
(Taxonomy and morphology) Fortcy. R.. A. and Owens, R.M. (IIJ75) PruccidJ: a new ~I
Eldrcdg~·. N. ( 1'177) T'nlohucs .and evtllutaonary pattern~, urder of trilobites. Fossils mu/ Strtlltl 4 . 227-40. I'
In Pollft•ms •!{ f;p,,fuli<'ll (ed A. H.tii.Hn). E l,evtcr. (T axonomy; mcd m the cla,slfic<ttlon pr~'cntcd here) Jlol
AnMad.un. pp. 305 Jl. (Wh.at Jo tnlobJtes have to Fortcy , R.A. ;md Owcns, R.M . (l'J90.a) Tnlubttt:,, m d
tdl u' .1buut cvoluunnary proce~sc\?) Ftot•lutitltwry Trmd5 (cd. KJ McNJnun) 13elh.wrn fl
cldrl'dg< N .tnd Gould. S.J (1972) l'um·tu.ttcJ equilib- lJO\tllll, pp 121-42. llur
ri.l: .ut .lltem.tove tll phvlettc gr..tJu.tlmu m .\J,otfd.s 111 Fortey, R A tnd Owens. R.M. ( llJ90h) Evnluuun.•ry Tl
Pc~ft·,,l·r,,f,'I!Y (cd. T.J. \11 Schopt). I rccm.m. Cooper :mrl r.lru.mon~ m thl' Trilobita. m ,\.[,,j,•r Et•tlluth•nary l:l
l.o . \.111 Fr.tnn,co. pp H2-ll 'I (!\llnparnc evolutim1) RtUiratrort.l (cds P .D. Taylor .md (;.P. I arv.nod) l•t
l~ldrl'dg(, N ..md Onm~ton, R (llJ7 11) llao~co~phy of ~y,tcm:mc~ A~,ociation Speet:'al Vol. 42. Chrc.:nJun J)
Stlun.m .md Ot'voniJn tnlubttn ot' the M.1h1nokaflrit. Press, Oxford, pp. 139-64 . _lo
Rl'Jint, in HistcJricc~l BtO.I!<'cll!r.ljllly, fJI,ltl' Tl'rrCittt(S autf tlw Fortt'y, R A. and Owcns, R.M. (I'J'Jl) A tnlolmc faun• llor
Gh111r_t:11(~ l~lll'lfcJIIIIII!IIf (ed~ J Cr.1y .1nd A tio u cot), fro m the ha~h cst Sh inc to n Shale, ll1 Shrnp\hll c, .amltltl' St
Orc~;nn St.u,· U111vt•r~iry Jltc"• Portland. Orl'., pp. rorrd.atm ll nf the T remadoc ( .roh•l!tutl Af,~f!tl~lllf 128. t•v
147 (\7 437-(14 l'fl
EvlU. W R. (I '161) Ltrh nntngcnv 111 the rnlobitc family Fortcy, R .A. Jnu Scuachcr. A. ( ltJ 1J7) l h~· tr.Kc lu111l I lug!
A,.,plud.Jc _l.lllmdl ~~r H!hwttc•l·~~y 35 <JH6-95 C:nt.;;hllltl ~cmrplicata, and the tnlobatc that m .• dt• I\ lllt
Fct~t. R (llJlJI) 11te Late Dcvtllll.lll trilobite cnst.><.. uth<111t, 3 0. XlJ-16H. lr
fli~lt•ncal Bwli>I!Y 5. 197-214 Fortey. R .A. .md Whitongton, I I B (JIIXIJ) The RtJ
Fcl\t, R Jnd c.L.ukson, E N K ( 11JH<J) F.nvironmentall) T nJobtt.l as ,, uaturJl b'Toup fiHI<>Ilftlf Htc>lt")' 2 WC
,·ontrnlk•d phykn( evoluuon. bhndnc'' .111J cxnncnon 125-\H c." VI
111 L.11c l)l'voman trupidocoryphine mlob11". Lrtl1mu llammJnn. W . (1':1~3) Calymcn.u.:t:J (fnlobttJ) .m, Jcm JcU I
22, 351) 73. OrdovJ.ti um von Sparucn: 1hrc Blmrr:ltlgr.tplur Ail
Ftshcr J) (1475) \wtmmmg .md hurrowmg m Limulus 0 "-oloj;,'il' und Sysremali k. o1bhcmdlul~(/fll J,, .J.:ml'l
.mcl .\lr•sc>limlllus. Fo., .,;ts ,,,,tf .'~trara .S. 281-90. &11rkfllllt'l;f!isd•c Natur:forsdmult· Cl'wll~t llc~fr 542, I Jn l)IJ
(lmport.ant funcunn.tl ~tudy) (Enrollmcnt >tm ctures) Mi1

Ftsltcr, D ( 1974J) FV1denre lnr ~uh.wml activity of H .llll TII ,IIlrl, W and R abano. I. ( JIIH7) Morphnlngit und _l one~.
/'upr•'•'P·' diltltl<', u1 .\fa;:(llt Crrd: h'·'·"'~ (cJ. M. Nitccki), Lt•bcn,WCISC dcr Gattung w/rllclpd/IJ (Trilubll.J) und \ lv.
Ac.tdumc Press. Nl'w York. pp. J7'1 447 1hrc Vorkomrnen in1 Ordovt:uutu vt>n SpJntl'll (Pit
Fordyct D Jnd Cronm. T W (IIJ!N) A tomparison ot St:lltkclll>t'I)!WIIt1 L:tltara 68. 91 -137 Lu1·
the iil\'lh,cd compound C) t'' nt' plt;tt up1d tnJobucs H.mk~n. N.M lnd Sronner, L. ( 197'l) The cr.til uf tl.utt K.lufn
WHh !h~ eyes Of modt'CO IIIJntlC .lrthropmh. jounra/ o( ~Jlun.tn t•uryptl'rid FIJ.ssils c111d Stratrl 4 , 2'i5-70. (Hl''' j;t'll
GrttJI•l•<'rlll l3r<•lt•~y 9 . 55.J.-hCJ l'uryprcmh mcd their le~) llbc
Ft,rtev. R.A. (I '17 4a) A ne\\ pdJj..'1C rnlob11c from the H cuniu~utotn, G. (1957) Thc rnlobttc t:um ly Ulcmd.~e .Jbfr
OrdovJCr~n ot Sp11zbcrgen, lrd.md, .l!ld Utah. N(lrskl' l"idellsl.wps :'tmdrmit ,\lmlnmali;nr I r,,.
Palat'tllllolc!C/Y 17 . 111- 24. Natun•itft'tllkaps Klasse Skrfftc I , I 303 (MorphuiO!o')', t1011
Forrcy, R.A. (1'174h) The Ordovwa:an tri lobites of taXO IH) IIlY d lld cvoJutJOil) KJdk~·
Bibliogra phy 403
Henmng;mo~:n, G (l'rJ'l) '\1oulung m tnlobitl"' hm1/s of th~.: Welsh Bordt<rbnd. Jounlll/ <1( H1ft,utt.•IO.I/}' 35 ,
.mci Strata 4, 17?-:!00, (Exuvt.ll .membl.ag~s and v.anou" 784-835. (Euryprcri d .t~sl·mbbgl'S)
w.1ys in whu h trilobl(t:\ moult~d) Ltndstrl'lm. G_ (1901) Re~earchcs on the visu.llorg- Jn' of
Henrv, J.-L. ( 1980) Tnloblte\ mdoviltCII S du Massif rhe tnlobJtc~. k.ou.l/ltf/<l 'll'wsk I 't·tcmk.lps 4kadrmu·
am10ricun . .\Ihn,>m·< dt· /,t Soot'li tit· Cc,>J.wn· t1 Handlm.</ar 34, l-B5. (Ptrst work on rnloh!te eyt''·
A/inhllh>fi!tC de Breto~m 22. 1 250. (.l:.nrnllment \ln.IC- hypmtom c may have vcntr&.l eye~)
ture': cxquNcc phutu') Manmng, P.L and Dun.lop, J.A. (I'N5) Th1.· rl.">piraml)
Henry, J-L. ( 14R9) PJltncnvt ronncmen t' et dynJmtqu c orgam of curypu:rid s. Polactmlt>h~flY· 38, 287 'JH.
de f.1Unc' dc mlobttt·~ dJn' I'Ordovtn cn (LJnvim M an ton, S.M. ( 11J71) Arthropod phylogcny - ,\ modern
'upcril'ur- C.Jradoc hJ,J.!) du Ma~~•f Amlllncai u synthC\l\ jottnt<lf <!{the Zoo/ol/!f•tl '\.>ol'ly <!/ l-LIIIdtlll 171,
FrJncc) f>.t/.~t·t~flCt~l('<lf'lly, f><lilll'•'' lmlllt••h•,n·, Paltlt'<>rwlogy 1.11-30 (Clamfi,a non of an-hropods )
73. 139-53. McKerruw , S. ,md Scotcse, C.R. (ed~) (ltJ'JO) Palart)~••ir
H:nry,J.- L .md Cbrk.,,m, E.N.K. (l'J7:i) Enrollmen t Palaco~r,wraplty <tttd Bio,l!t't>ttmplry. Geolog~ta1 Sout·t~
lnd coaptJtiOil.\ 111 ~ome 'P\:OCS of the:- Ordo\ tcian Memotr No. 12, 2-'0 pp
mloh1tc ~i.'nm Pl<lWJ'tlll<l. Ft>s .t/s 1111d 'lrr.tta 4, H7-9(>. MLNanuro~, K.J. (1978) P~edomorphu~i.~ m Scotmh
(Lntcrlockmg rnrchatmm s and their evolutiOn) okncllid trilohttcs (early Cunbrian ). Palof'<llll<llt~'()' 2 1,
H~nl')·. J.-L. lnd Nion, J (1CJ711) Nouvdlc, observ.ttio m 635-5(,
'ur quelque' ZeJ.a,Lkdhnac er PhJlOptdd Juui.' de McNamar. 1, K .J . ( 1983) l'rogcne~Js m tnlobJtcs, 111
I'Ordoviu en de Brc:t.Jgnc. Lt•t!l.lia 3. 2U 24. Trilobircs aud Othl'r Arlhn>pods (ed~ D.E.G Hngg> .md
Holm. G. ( IH'JH) Ober dtc Organt\at ion de' L:uryptamjis- PD. L1ne), ~peoal P.1pCT\ m PJ.lacontulot-n Nu. 311
··ltcn Elchw Hu/lt-1111 t>{ rhr 4r,l.frmy •!I Scit'ttccs tJj St Acadcn11c Pet:~~. Loudon, PP- SlJ-<1H.
Pw·d'''"~l! 4, 369-72. McNamar a. K.J. ,md R.udkm, D. (l9X3) Techniqu e' ol
Hnrvath, G. Jnd Cbr~'on. E N.K (I'>'JJ) (omput.1t1 0nal trilobttl' exuvratiun . Lt:tltaw 11, 153-73.
rccumrruc tmn of the probabk chang~· nl fonn tlf the Miller, J (1975) Stmcrure .md funcunn of trilobite tLrt.lu'
tome.ll lt'm ,md nuturanon nf opt!•' m the pust- lines. Possi/s ami Strata 4, 155-78. (Env1ronmcnt~l
ecdy,ial developm ent of the \lhizochro ,tl eye of the monitorin g)
Devonian mlobite 1'/l.k·,,p; ''""! mtllm \rcwart 1927. Miller, J. (197(,) The semory ticJd, and hie modt' of
}MII'It.ll oj 11uwt'ltml Hw/.1~)' 160, 3-U-7:\ Pit amps m11a (Green 1R.,2) (Tnlobna) . Trall.mC/IortS of tlrr
HorvJth, G., C larksnn. E N.K .•md Pix, W . (l'JlJ7) Royal St!(iety of [;'tllrJbtJr.'l!lt 69. 337 -f>7. (1-unrhon~ of
Survey uf modem ,oumcrpJ rt' of sduzuchro al mlobttc cuttcular sensory organs)
c\·c~ srrucmral and funnwn.u ~•mtbntte~ and thffer- Mtllcr,J. and Cbrhon, E N.K. (ICJHO) The pt~\t-ecdy~tal
cn.-es. I /t.lltlllt<tl Bialc~~y 12, 22'1-63. cievclopm ent of the l'Uncle and the eve of tht•
Hught'\, C.P., lngham, J K Jnd Addison, R ( 1975) The Dcvoru.m rrilobtte Plraro1u r111111 milleri Stt•wan- 1927.
morpholog y cl.J'>\Iht.lU Oll 111d evo)uoon of the Pllilosoplrical Tmmaaio11s ,,, lht• Roy.1/ Socirry t!( Lottd,>tt
lnml(/ctdm• (Tnlobtt.t) Pht/o,opltual 'Jr,m.wclimzs t?f tlte 288, 4l, l -80.
RtJyol So<icly of Lt>tu/mt B 212. 537-{107. (Dctimriv e Nathorst. A.G. ( I HH1) Om spar af rt.t!,rra t'Vl'rtebrer,tdt•
work wnh dcscriptto m of all trinmlc1d gener.l .1nd djur oc h der.\~ paleontolo giska berydd~c Kon~lt.l!<'
evolunotu ry chJrts) Sr'f'11Sk Vctcmk<tp:. ,1kadcmrl' Hm1dli11,1/<lT 18. I 104.
JcU, P.A. (1lJ7R) Tnlubtte re~ptr.lttOn Jnd gen&.l caeca. t~pik. A . ( 1960) Alimentar y c,\eca of agnostid and other
Jll,/lrnttl(•1 2. 2S 1-(111. trilob1te~. Palat'tlllllllogy 3, 41 (}-38. (Surface ~culptur~:
1rmtn,
S ( 1')90) Pred.mon b\ c .1rly C.unbnan rnlobite~ .md tts tnrcrprew uon)
on mfaun.1l worn~ evJdc:ncc from rhc Swedish Osgood, R .G. (1970) Trace fm~i ls uf the Cinunnat i :m·a.
Milkwitzta Sandstone: L•tlto~i<t 23 , 29 42 PalactlllfO)!rap/um -lmericawt 41. 2H 1-4-44 (Trilnbtte
>n T.R and \f.!uodw.m l. H. (1HHX-18lJ9) ...t tratls, burrow) and method' oflocomo non)
\f!llll>gmp/r •>/ 1/11 Bnlt~lt Palac·o~,,,. Pllyllc>J>Otia Orlowsk1, S., Radwansk t, A. and RomcwK 2, P. (llJ711)
!Piryllomritla. Pac/.:.tr,/), Pal.\eontog-raphic.u Son cry. The trilobite ichnococn mes in the Cambn.tn sequenre
London, pp 1-211 of the Holy cm~~ Mountam s Pol.md. in TrLICC Ft•~>rh
K:tuhnJnn. R ( llJ33) V m.1rion'-'uti~nc hl UnteT\uch un- (eds r .P. Crimes and J.L. Harper) Sed H ome Pre\\,
~rn Uher dtc Artabw.mdlun~ and Arluc~tbildung .m dcr Live.,,ool, pp. 345-60. (l)c~criptton of Rll.\llf'ltycus .md
ooerl..amb n,thcn 1 nlobnen~.tttun.: Ofnms D.\lrn. Cru;::uma)
,1!Jh.tudlurt~ tlrr Gt·,,f,~tSclrn·l'.t/.i,>lllt>/o,.:t-'• ltt' l11<111111 Jl'r
Palmer, A.IL ( 1'J'i7) OntoKcne nc dcvdoptn l'nt of rwo
( rtit•fJStliil Cnc{swald 10, 1-54. ( Chwc study of evolu-
olend!ttl tnlobtle~. )o11mal of Palwlltolo.~y 3 1 , I05-::!H.
tion in Oh·m11) (Changes in proportion during ontOgt·ny)
Kl!'Ul"'''-1);;- W.tcnng, E 1':. (I% I) The Stlun.m Eurypterid a Plum~ek, R . (19H5) Ltfr ba,l·d mech.llll\t ns for \Wtmmin~
404 Arthropods
111 Clll"\'ptcnd~ :111d portunid crnh\. Tr.msartitms f!( rite Sivecer, DJ. and ~ddcn, P. (19!>7) A new bri.lllt XIpho-
R•'Y•ll S,•orrt) ·~I Fdru/•mvlr · E.m/r S..rn'cn 76. 325-JS. ~und from the Lower NJmunan of Wc·arrulc, County
Raa~ch. G.O ( JI)JlJ\ C.u11hrwr .\/rn>liOm<l/•1 Gcological Durlnm. Pf(l(Cfdiii~S 4 tilt' ' '••rk.•lmc Cl'olo.tZ'l•11 '"'" ty 4(1
'-loci c.'!"\' of Anll"nl'.l, ')p<'CJJJi~t Po1pe"' No. I<,~, 15.3-61:!. Th
Geolog.c.tl 'm• icry n! Amcril.l. LJwrence. Kan. Spt'yer, S. and Brctt, C (, (19H7) Clu\tend truobttc (
(Agl.hp•d~) J.)~cmblo~gt'\ 111 the Mtddlc.: Dc.:vomm I!Jmilron tirour
R.1ymond. P. E. ( 1'120) TIll' app••ndag<''· .macomy and urh,,;, 1s. H5-J03 w.
relaaon,htp' nf mlobtte,. Cot~~recllotl .-lcadn11y of Arts ~pcycr, S., .md Ch.mcrwn. B D.E ( I'HN) Trilobite brnt I
aud '\.rt'lll<' .\lt-m.•rr• 7, I ·lhlJ. (Pe,.:npnum of rrilobtte and larv.J CClllllg}' ll15tt•ncttl BroiOI()' 3. 27-611.
append.!~,., wtth lln<' phutogr.tph\) Sritt, J. (1976) Funcnnnal morphology .md ht'l· habtrs 01 Wa
Rob• son R .A. ( llJh 7) <. )ntogcny of &lthyunsrus funbriaws the Late C.1mbri.m mlolmc Stmoprltl> JlltllflH R..t1 ntul!d .t
,111d tt' be.mng on allllllltc' ot' .:oryoexod11d tnlobiceo;. · Joumal ~I H1h'OIIIt>fo.l?y 50, '1(>1- 76. I
)<>llm<lf v( P<lfcvlllt>ft~l/)' 41 , 213-21. (U~l' of Ontogeny m Stitt, J. ( IIJHJ) Enrolled 1 arc C ambn.m tnJobttc~ frmu the Wa
phylogenecil affinttu:s) Davts Fomunon, ~nutht•.lq Mt~~ourt )ourn.1l vJ I
Robi~on, R.A ( 11172) Mndc of hti.· of Jgnosttd trilobtb:\. Palmllfc•fll,~tY 57, 9 ~ 105
lutt·m•llltllltll Gt',,fologuctl Ct•ll,l/1<'5.1 24, Ses\iou 7, JJ-+0. Stockton, W.L. .md Cowen. R. (1976) Stcreo,,opt• Wa
RoUt-. W.D.I. (1%2) Grum:r mmpholo~:-ry of the Scottish vision m onr eyt· - pal.tcophy>to1ob'Y of rhe schtzo- c
Siluriau phylloc.u-id C"nl\t,lccan Ccmtroc.rris ptrpilio Salter chtOa1 eye of lnlohttcs. Pa!twohi,>I•\1/Y 2. 304-15. (Ne11 (•
i11 Murchi~nn Jarmwl ..( Pah·olltolv_~)' 36, 1.)12-32. interprel.alton of' eye funclton) s
(De~criprion) Stonner, L. (19JO) Scandlll.IVtan l'nnude!dae l'wr;kc ~
Schmalfu\,, 11. ( l'l>-1 I) Stnl\.turc. pmcm' and function of l·ldcm.skapr .-lkadcmil' Mtlllwmatik-Nt11111Vidrll.<kcrp5·1\lc•i( We
cuocular tcn·rc' m rnlobitc' Lrtl~ttia 14. 33 1-t I. Skrj{te11 4, 1-1 J 1. (Dct.Hblmorpbolo~:-ry offrmgt .111d (
Seihcher, A. (I 955) Spurn1 und Lebcnswei;.e dcr scmory orgam) \I
Trilob1tcn: )purc.:n unJ r JliC\ un Untcrkambrium. in Smm1cr, L. (I 'JJlJ) Studtc\ nn tnlobne morphnloro. Pm We
Hritro~r :ur ~."'"'m Jn 1\.,Jm/mum.r '" der Salt Ra11'(C 1. The thorJcJc appendage' a11d th<·ir phylogcncttt ••g- il
(Pakw.m) (cd' 0 11 "idundcwoll md A <;eilacher). ruficancc. 1\m:.k Ccoh'.~t>k TrdHJ.m(t 19. IH-273 Wh
.-lhrdwur flh rt H.H'II<dr.tjt tmd Lltmlltlr .\lam::: (Ceraums .lppcnd.igl·~; J very dct;ulcd 'tudy) t1
.Uatlmu.lllk· '-·"""~'"·'I'HSt b.!ftm ~/,tss.: Ab/Joudlrm(/, Stomlcr, L. (1 9711-1 1J74) Arthropod\ !Tom the lov.er [
~6-I·U . (fll'\t repnrc of ,,,kway'l-l'I'Jwling triJobite~} Dn·onrdn (Lower ErmrJm) of Alken .111 dcr \lo·d fi
Sc1lachcr. A. (llJ1,2) £\mu .tnd l--unktion de~ Tnlobiten - Gcnnany. 1970 Pm I. Ar;~clwida ; 1971: P.ut 2 Wh
Oak!) Ius. Pa/,;,,,,,lv_t.:Bdtr Znt~. hn(tt' (Herra Schnudt Xtpho~ura; I 1>7J : I'art J. Euryptenda HughmiUcnd.Jc n
Fe,tb.tnd). 21ti-::?7 (Muchtn~ of tnlobttc tra.th ,.,.,th 1974· P.1rt 4. LunptcridJ Dtcp:tnoptendte '\or·hn· Wh
;appendage ~rrunurl·) brl]?i,ma L·tllltt\t 51, J35 69. 53. I 2\J: 54. ll 'l-2~6. c
Se•l:!chcr, A . ( l4h4) Utugcm~ wduncntJry stmcrures. iu 359-451. (Cb"thc.mon ot curvptt:nlh) 6'
."lpprt~arlrn to !'tllc!l't't'Wlt'S)' (cd.' 1. 1mbne and N. Scormcr, L. (I 'JRO) Sl'ulpturc and mtcrmtrucnrrc of rh~ pi
Newell). John Wtley, Nn\ York. pp. 296- 31/l. cxthkeleton 111 dum1Uptmd .lnd phacopid trih,bucs Whr
(lncludc' di~c US\11)11 nf trilnhlll' tr.tl k,) Palaeonh>lt~I/Y 23, 2]7 7'2. IT
St>klcn, P. (I'}~ I} hlllll!Otl.ll morphology of the prosoma StOrmer, W ..md 1krgsrrl\m ( 1971) New dt~covcnes 1111 c
of Baltot'II")'Jllrftl\ tl'll•l\!OIIOJ'htlrallllltJ (Ftscher) rnlobttes hy X-ray•. Pttltrcmtolo_l!it• 47, ltl4-ll. (Append- .tr
(Chehcer.tl.t I'urvptc.:miJ) I;m''''cliom ·~F the Royal age morphology) Wht
ScJudy v[ Erllltf,ur~lt fi<llllt Snmm 72, Q-18. Stubblefidd, J. (I 'J.\6) Ccph,thc ~ur11rc\ ,111d the1r bc.umg ar
Scldcn, P.A. ( 1\ll-14) Autl'Clllt~~.')' ofStluri.tn euryptcrid~. in on current da~~iticntton nf tn1obtre~. Hit•/o,~irc1/ Rer•rciiiJ c
A11taohlg}' o(Silllri1111 ()~~·minm (t:d~ MJ. D.1~~et andJ.D. 1 1, 407-40. (Cl.mit p.tpl'r porntmg <)Ill limitJtmm nf
l Jw'ion), Ciped.tl P.lpl"r. 111 p,,],H:L•nwlogy No. 32. 'urrure' .ts taxonorn.tt trlll'rta)
Ac.1dcmit Prc),, London. pp 3'1-54. Tcig1cr. DJ and l owc, K.M (JIJ7'l) Mtcro\tntcturr ~nd
Shcldon, I'.R (I 'JH7) PJr.llld gr.~duJ.INic evolution of compo,ioon of the ll tlobtle cxnskdl·ton. r,,._,;/; ctnJ
Ordm·tn.ln mlulml.'~ \Jc~~urc 330, 561 - 3. Strata 4 , 137--19. (Cuudc.: of \Cn:rJ.l ~pcul"' descnhcJ
ShdJon, P.R (19HH) Trilubtte \lLl- fn:yucnc; t.hstribu- Thom:ll., A.T. (19H(I) l'nlobtcc· a"nctJJtons 111 the Bntuh
t.tom, rccu~utton of m~r.m . .md phvletic qze change~ . Wcnlock, m 17rt· Calrdmndo t>f the Hntr<lr l.i/rs -
Lrthatd 21, 2'H- \oc,. Ri'l 1u'wcJ (t:th A .L. H.mt'. C...H. Holl.wd .md ll.f.
Shcrgnld. J 11 (1971 ) L.at• Uppci C.!IllbriJ.U rrilobitl"' Lcakc), Spcctal Public~uon' ol the: C.cologK.&l Som~
from the Gol..t Unh. \Vl',tlnt Quccn,l.lnd. Bureau •?f oi London. Vol. X, pp. 447- 51
.\/mn.r/ Rrsom•(S ( ,.·,•1.•1/)' .md (,r,>phyncs Bulletm 112. Thon1.1s. A.T. Jnd Larll'. P.D. (llJM-t) Autccolog) of
l-12h. Siluri.1n trilobttc\, ut • lult'<'•'lt~r t>{ <:;,[un.m Qrx.tmsm>
Bibliography 405
(eels M.J. Ba~sct .md J.D. L.1w~on). Spcn,•l Papers in 171-204. (Sccpncal about Cw::iarw bemg of trilt)btte
Palacoutolo!,')' No. 3:!, Ac.1dcmJc Prcss. London, pp. origin)
55-69. Wluttington, H.D. ( 1981) Pacdomorphosi' a.nd crvproge-
1 homas, A.T .. Uwem, R M . and Rmhton, A.W.A. nesil> in rrilobitc\. Geological ,\Ja~a::lllr 118, 591 <o02.
(1984) Tnlob1tc' 10 Bnmh straograph) Geologtcal Whmington, H B. (1988a) Hypostoml'S of pmt-
Society oflondon SpewJ Report No. 16, pp. 1-78. C:unbrian rnlob1tes. New Alexrco Bureau ~f Afiurs
Waterston, C.D. (1957) fht: Scorti~h Carboniferous Mi11cral Rescarrh Memoirs 44, 321-3<).
Eurypterida Trrmsactwm of tlrr Royal Society of Edinburglr Whirtington, ll.U. ( 1988b) I I ypmtomes and ventral
63 .265--RH cephalic sutures m Cambrian trilobites. Palanmlt,fogy
Watcmou, C .D (1 %0) fhc: median o~bdominal 31,577-609
.1ppcndage of rhe ~iluron eurypterid Sfmtoma aomri- Wlurnngton, H .B. (1989) OleneUoid trilobites. type
llata. Pt1/acolll<llo~ 3, 245--SIJ. spectcs, functional morphology and higher cl:bslfica-
Watc:-rston, C.O. (I !J75) Gtll structures in the Lower tion. Philosoplriraf Tramactio11s cif tire Royal Sclril'ty of
Dcvoman curypterid Ttlr.wptcrelfa )CIJtica. Possils and Loudo11 B 324, 1 I l-147.
'irraca 4. 2 1I -'l4 (Detailed morphology of gills) Whittington, H.B. (1990) Arocul.uion and exuv1at.ion m
Waterston, CD. 1979) Problems offunct1on.1l morphol- Cambrian tnlobites. Pltilosophtcal Tramactious of tltr
ogy and cla\>ification 111 \tylonur01d curypterids Royal Society '?f Lottdtm B 329, 27-46.
(Chchcerata Merostomata) with observations on th e Whittington, H ..O and Almond, J.E. (1987) App~nd.1ges
Scottish Stlurian Stylonuroidca. Transarrions 4 tlu~ Royal and habits of the Upper Ordovician trilobite Triart/111.<
Society of Edin/Hr~h 70, 251-322. ealtllti. Pllilosopl!ica f Transactrons of tiH• Royal .\c)(ltty of
We~tcrg;\rd, A ( 1422) Sverigc' Olen.idskitfcr. Svmges Lo11don B 317, 1 -46.
GtofogiSk,, l'11tmolmm!! 18, I 205. (Cia<s1c taxonomic Whmington, H.B. and Hughes. C.P. (1972) Ordov1cian
study u( olerud-beanng sh.llc\ of ~canduuvta) geograph y and faurul province~ deduced from tnlobite
We,trop, S.R. (19H3) The ltk h.lbirs of the Ordovician distribution. Piu'lnsophical Transactrmu of thr Ro>ynl S~>rtt'IY
illaenine tnlnlmc Brmuutoitlrs. utllaia 16, 15-24. r!f London B 263, 235-78. (With detailed 1mps; upd.ltl'S
Wluttmgton. H B. ( 1956) S..hdied Middle Ordovician Wh.irtington, 1966)
trilobit~:s: the OdontoplcuncLe (Trilob1ta) ]014maf if Whittington, H .D. and Hughes, C.P (1974) (,eograpby
Pafeolltt•h~y 30, 304--520. (Deuiled morphology and and fauna! provmccs, in Tire Trr11r.uloc Epoch (ed C .A.
functional rntcrprctaaon) Ro~s) . SoCiety of Econorruc Paleoncology a.nd Mlm.r.U-
Wluttington, H .B ( 1957) The onto~ny of triJobites. Ob')' Specialist Publication No. 21, Society of Econom1c
Biologiwl Rc11iews 32, 421 -69. (~tand.lrd reference) Paleontology aud Mineralogy. Tul~a. Ok.la., pp. 20J IH.
Whittington, II.U (1966) Phylogeny and di~tnbution of Will~. LJ. (1965) A supplement tO Gerhard Holm\ 'Ober
OrdovlCJJn tnlobites. .foumal if Palrontology 40 . dJe Org:misaoon des Euryptcms .fisclreri Eicbcr'. \Vlth
696-737. (Evoluoonary cll.lrt~ and discuss1on of fauna! speetal reference to the orgaru of 'ight, re>plrJtion and
provinn...,) reproduction. Arlcrvjur Zool~lt. 18, 93-145.
Whtttington, I I. B. ( 1975) Tnlobires wtth appendages Wilmot, N.V. (1990) Biomechanics of trilobite cxo,kcle-
from the Middle Ounbri.111, Burgess Sh,1le, British tons. Palaeolllology 33, 749-68.
Columbta. Ft'S5tls a11d Strata 4 , 97-136. (Description Wilmot, N. V. and Fallick, A. (1989) OnginaJ JTuncralogy
and funcaon:tl morpholob')' of Olmoitks appcndabrt:s) of trilobite cx.oskeletons. Palaeo111ology 32, 297 304
Whitcington, H U (1980) l:xoskderon, moult stage. Zhang, X.-G. and Cbrlcson, E.N.K. (1990) The eye<> of
appenda~l· morphology Jnd lubits of the Middle Lower Cambnan eodiscid trilobncs. Pafaro11tolc11/)' 33 ,
Cambn.m tnlobttc Olwaitfe< scrral1tS. Palc1L'OIIItllogy 23 , '1 11 -32.
Exceptional faunas; Jf.
ton
ichnology filh
sed
ties
gen
e~"a
part
In th~ hnal chapter wt· cu1Nd~t two qwte different numbers of lt)"tls arc pre\crvcd. The\e include the mat
and c01 tr<llltmg tllllll'll\luns of p.1la~ontology. The following: ITICC
firsL concerns t'xcepllon.uly well-preserved fossil rest<
faunas. how they umH: to be prt'llcrvcd and what condensation deposit\, \\here Lhe ratL ut sedt- the :
th~y have to tell u~. The sccond ts ichnology, the mem<ltion has been 'o slow, that an extended sott-
study of trace f()~,lk the track~. trails, nunes and gal- time period i~ represented by a thin layer only: auth
leries nude by long-vanishc·J t rcaturcs, 011 or in the th1s may be crowded, for example, with ofci
sediment. Wherca'> rhc· tr.u-c 1nakcrs arc not nor- ammomtc shell~ which had been \lowly accumu- be p
mally thcmsdvt''> prt'~crvc·d. p~thaps paradoxically, lating fo r many thousand years; alter
th~ir bt havwur p.Htcl m Jrc ] he se arc two end placn depos1ts, Hit h Js bouebcd,, where fr"'J! pyrit
point~ 111 pJI.Jcontolot-,')', e.1th of wlm:h illuminate~ maten::~J ha~ been tOIH.t:nlratt'd, tor ex.mtplc. b\ may
uruqud) the hl\tory nl hie on this plJnet. tidal tag cIfet r~; ln
concentration rr.1p' ~uc h a~ ti"urc tillin~ mm JllOX
wh1ch small vcrtcbr:~tc~ !Jvcd or fell Jnd were ~ott-]
buried. cases
nfau
'Only J fi-.1rnon of tht· myn.1d crcatt1re~ that have These concemunon depmit~ havt' a greater num- the o
lived 011 the E:mh h.l\'C left bchmd traces of the1r ber. or a more d1ve~e a\~embl.1gc of fo,~ih than at thl
ex1stcnce, and only ~petltir parr' of thtNc' orgarusms usual. but nornully prc~CrYl' no more than shell~ or con d.
have been prc\t'rn•tl' (Bngt-,"S, ll)l) l). Th1~ ~tatemem skcletom. is inh
is indct•d rme. and 1t t•mpll.lsi7c~ th,Jt the vast Second, there art• conservation deposits .md Strat1t
111<lJOnt'\ of fo...sll .mt·mbl.lgt·~ arc composed only of ir i~ these that m:Jy. and often do, prov1de mlorm - ktnds
shelb or \kcktom Nnnn.11ly we· ex-pect to see rion of much higher quahty, .md can mduJc AJJjso
no more than J n \ITO\\ h.md of 'pre\ervable' organ- soft-parr pre~crv;Jtion. Among-;t these arc the rate o
t~ms from .m ong111.tlly much broader biotic follo\vi ng: the gl
spectrum. Acrnrdmgly n11r conception of the comb
course nf orgamc evolution through time is based stagnation deposits, where anoxiC bottom condi- (Fig.
upon ·,1 snull ~ample t<lll\1\hng .tlmmt entirely of tions have precluded tl..- :lctivitlc~ of 'iCJvcngm; (c:'llctt
anim.tls w1th prcwrvahle h.ml p.ms' Oohnson and in bbck ~hale~ thus depo'ilted may be ti.1und Pyr
l-ttchanhon, 1969) undi~mrbed and exqu1S1tdy preserved rt.'mJILll l>f rapid
Then: arc, howt'Vt'r, r.trl' nrcum~tances where gr:1ptohre~ or fi'ih; or nea
fmsih on·ur c1thtT 111 rcmark.1hk umct·ntr.ltion or ohmtion dcpmm, where a nch b1ot:l may bc pre- .lV:'liJa~
in out~t.lndmg prt·,erv.H ion. The gcolog~cal hori- ~erved by rap1d bun,JI: 111 ~Ul h tmt.mce'i ~tartish or place ·
zons pre~crvmg \lll h remaim were tenned by crinmd~. wh1rh u~ually brc·ak up very soon at1cr high. I
Seilacht: r ( 1970) FmSII-l agcrsr:ittcn, ·whKh may be death, rcm.1111 mr:1ct .lftcr hcmg 'mothered bphc depm11
rough!} cramlart•d '' 'tm,il-bonanza,· or 'rock bod- inmshmg sediment and arc fmsthzed; pre~cn
ies unu,ually nch 111 p.tlacomolog~cal mfonnation'. comcrvanon tr.lp~. a das'iiC ex.1mple bemg the ti:z.JtioJ
Of tl C\C, lir...t tht'll' .UT con centration d eposits rrappmg of m\ett\ m tht' \Ocky reo-m of pme treo lugh 01
m whKh, .Ha parcicul.u honwn. ntepnonally large wh1ch 'iubsc'lucndy ll.lrdt•n, .l~ amber: saiJmty
Introduction 407
concr et1om . wh~rc the lo~~tls art· prese rved Calci um phosp hate (.1pame or fluora patite ) pre-
wttlun " diagc nettc rwdu k
\erves detail \ of structLirc bette r than .my
other
authig enctic miner al. Howe ver. 1t will nut
If, in the\c ea~~.·-.. anyrhm~ other th.m shells or skcle- tom1
where the conce ntrati on of HCO , 1ons ~~
tom IS to he preser ved. l>ne ronch uon rnu't bt: h1gh:
ful- such aJk.1linc condl tlons wiU al CIVJte imtca
lillcd: that diagt·netll ch.mg<..·~ wtthr n the t•nclo d the
~mg prl·cipitatlOn of calcm m carbo nate. and tlm 'dct\u
'cdim ent \t:t in npidl y, Withi n hour\ of death . No-., lt
a mech amsm ' has to be switc hed off if Lalnu m
freshly depos ited ~cd1ment nuy comt\ t ol very heter pho5-
o- phatc is to form (Llri~ and W1lby .
geneous comp onent '\, clu\.lWn together, and not llJ%)
nec- Experiment~ on dl.'cay mg ~hrimp' in
noml dl 'open '
e,sarily 111 J st.1tc of chl.'mical cqlllli bnurn . Thi'
is condi tions show that CaCO , cryMals be~in to
p:utlc ubrly tnl<..' tf tt conta im the remLum of pre-
hvmg ctpit.l te wtdu n 3 d,tys of death . Phosp hate, howc
nmen al. '1 he bre~kdo\\ 11 of tlm org;Huc maten wr,
:U nnly precip itates m low-p H condi rions, a~ favou
medJated by b.Kteria m.1y 'l't up re;tl ttom wh1ch red
by 'close d' anOXIC condi twns, ,md mflue nccd
rc~torc chcmic;U balan ce but may alter the by
char.t etcr of micro bial activi ty (Bri~ and Kear, 1994) The
the \t>dum.·nr profo undly rbj~ ts di.lgenc\JS, 111 wh1ch bal-
ance, howe ver. is delica te, and initial ly pho~phanzc
~oft-part fLw>iliz.ttion mvolv e' the grow"th d
of new. 1.e. m~ue m:ty be overg rown by, or 1~ fimnd
.luthigemc nuncr ah. In Cl'ftaiJl unu~u:U tomb inatio m do~e
ns a~sociation with bundl es of CaCO , crysta
,f ciroll11stancc the \ofi or non-m mcrah zcd p.un. may ls the
result of·' loc::thzcd rise of p i I.
he prc~ervt:d a.~ fthm of phosp hate. ~ihcatc or kaolin
ite: One rema rkabk exam ple of st:que nual authig esm
.Utem.uively cerui n org:Hts mJ) ht: wholl y or pama
lly miner ahzat ion ha~ been descr ibed from the Jur..t\~
pyriti1ed. R1rdy, even whole urgam ~ut h a' mu,dc il
~ of La Voult e :;ur R.hnn e, Fram e (Wilb y,
nYy he pn:~ ·rvt:d in phmp hate. 199(,)
Here lhl·rc is n divt:r~e f.1una of marin e inver tebr.t
ln gcm·ral tenm , rhc npid bunal of fo,"l' , .md tc'
pre~t:n•ed in three dimcm 1ons, 111 a ~haly
.tnoxtc condiriul15 (whjc h mh1b1t ~c.tvenging) f.wou depos it
r with str:tt. lhoun d nuner als, laid down in a restri<
<.oft-p.lrt prcse natm n, but 111 tht: va~t maJo nt) ..tcd
of ba1>m. In concr euom !Tom wrthm the
oses wft parts arc prc~ervcd only by rhe ~rrowt shall'~.
h uncnL~hcd antmals are uniqu ely pre\er
of authig enenc nuncr ah VC I) \oon ati:e1 the death ved Apati te
of fanne d l',trly a\ a t~mpl.ttc upon which other mme
the org:mimlS. V cry often ~eJimcnt~ are m:y!!,cuatcd r-
als fomte d m succc~sion: calcit e, then ± gypo;t
~~ the surfac e and ht:rc Jt:cay 1~ rap1d, but un.
:macr obic pyrite ± chalc opynt c and finally galen a. and
condi tiom .ue prese nt bdm\ In anox; c layer.. differ-
dec.ty ent kind\ of tl\sue wen: prese rved by differ ent
IS mlHb ited but not haltt•d. Tht·r e m.ty min-
he cht:m.ically eral~.
\tr.ltiiled zones with m thl· scdm1e11t \\her e tliffen
:nt There arc three comm on mode~ of presc rvnno n
kind' of bacte n.tl 1nilic:ttors opcr.H e (Ht:rn er, by
19!-\ I: amhi genk miner als: pemli ncrali z.ttion of ~oft
Allison, I 98~a.b, I 'JIJO) Three main param ris~ues,
eters: miner al coat~ and nssuc mould s. Penni
rate of burial. \:tlin.ity and or~anic tonre nt, contr ncrall zatton ,
ol the replac emen t of sud1 \mtet ure as muscl
dte geochcmtSrl") of the c;eduu ent\, .md m tiltrer< e fibres. s:.
..'nt very rare and wher e found (as in
comb irutio ns le.td to diiTcrcnt Juthig cw:ri c miner the muscl es ot
als Creta ceous fish from Brazil ; Mans ll, 1988) must
{Ftg. 12.11 ). These nun crab .1re pynte . c.11 bonat have
es r.l.ken place wnhi n hours of death . 011ly phosp
(calcjte or \ideri te). phmp hatl'S and ~ihc1. h::ttes,
the earhl'l>t authig encO l miner als to fom1, can
Pynte preserv::ttion of soft pam i~ f.lVoured by per-
miner alize in thi~ way, but they may also affect
rapid bunal . ,1 low orgam c conte nt and norm more
al dur,1ble maten al such as cellul ose or chitm
or near-n om1a l sal111ity ..md ~ulphatc ions must .
be Mine ral coats, the commonc~t mode of occur
.tvailable. Carbo nate prcop lt.ltlo n ts likely to -
take rence , may be of phmp hatc, c.ubm~<tte
pbce when burial h rapid and org.u m conte or pyrite
nt is The surlac c of the oq.,r.m.ism act~ as a
htgh. In low-, ahmt ) cond mom . sJtlent<..' tends to templ ate tor
be the form. ttion of nliner als, whos e prec1
dcpo~ired whcrc:lS noml. ll 'i.llimrv lc:JJ, pitano n j,
ro calnt e often uuuat ed or media ted by bacte
preser vation . The condttton~ reqm red ltlr phosp na. These min-
ha- l'rah tonn co.w. 011 the surutc e. and
tlZJrion of sofi pJrts arc a low rate ot bunal thoug h thl' soft
and a parts decay . their cxtem al coaun g does
htgh orgam c conte nt. VJvia111tc 1~ Jepm itcd m not, rema m-
low ing as a pst:ud ornor ph of the ~oft tissue~
\Jlimty. apatit e m norm.tl m,mn e cond ltwm . Well-
know n example~ 'uch as the Burgess Shale
. the
408 Exceptionol faunas; ichnology
Swcdt\h anthracomte\, the llun~riickschtefer and many occurrences there arc of fm~il a">\cmbla~es ,..
the Mc\\cl OJI Shale are \O prc,erved. At certain wtth soft-part preservation; over 60 ~lles have E
periods m tht· Earth's hmory. parncular mmerals been doc umented, and such Lagcr\t:itten occur in d
seem lo have been pJrttcul.trly abund.mt. There virtually all tl1e geological o;y,term. The following (~
wa\, fiH tmtancc:, a remark::tblc amount of phosphate list (modified from Briggs, 1991) gives Jn mdication
in the early and Mid-Cambnan (Cook and of where some of tbe more important one' he.
Shcrgold, 19R4), hence phmphatc coatings were Some sttes of greater interest from the pJIJeobot:mi-
partu.:ularly common at thi~ time. Sihca may also cal/early tcrreslrial fuunal and vertebrate point of l.
fom1 nuncral coats, cncru,ting ~mall trilobites and v1ew arc included here also.
brat·h10pods, for example, and fo,~tl pl,mts are some-
tunc~ found in three dimem10m. wtth cellular detail EOCENE. Grubc Mes~el. Frankfurt, C;emUII}
"
(e
prcosely replicated by \Utca coatmg. Lake dcpom, plmt:>, vertebrates, insect~ M

(Fr.UlZCD, 1985)
Tts~uc cast:. are formed by tht· \tabthnoon of scd- AI
tmcnt'> through diagcncm but bt:fore the rock C.:ll
EOCENE. Green R.iver, Wyomang (1::1
finally hthifies. Faunas found m 'lliccou~ o r calcare- Large l.ake depo\11. Fish and other vcrtt·bratt.,
ous nodules (e.g. the Mazon Creek and similar bio- (Grandc, 19!l4) M.
tas) are fonned through such proCC\SCS. Finally the Fh
Edtacaran faunall of South Australia (Chapter 3) CRETACEOUS. Sicrrra de Momscch. 'ipam (R
were preserved by lithification of the containing Web-we.1vmg ~p1dcrs, mseccs, crustaceam, vcrtcbrJtes
sand, whtch underwent very little compaction so (Se!den, l 990) l s
that the tonn of the oq,r.mt,ms \\a\ retamcd. An
CRETACEOUS S.mtanJ, Brazil (M
foSSII-Lagerstlttcn can tell U\ tiN much about the
Fi~h with pr~erved mu.sclc fibre\, ptero~u~ \'Hh \\11\~'\
anatomy and relauonshtp\ of ammals otherwise (MuniJ, 198H, 19<13)
known only from bard parts. The discovery of
pho~ph 1tized 'conodont ammah' 10 the Scottish U. JURASSIC. Solnhofcn, Germany
Carbomfcrous illu~trates thi' nicely (Briggs et al., (sec text)
1 983); the tooth-like conodont~, the only preserv- l.O
abk· parts of an otherwise soft-bodied creature, had M. JURASSIC. Chrhl1an Malford, En~l.md Pyn
been known for 125 ye:m. prevwusly but not the Sofi-bodicu sq•aids (Cts
(Allison, I')!!H;I)
ammal that bore them. Second, they tell us about
the n::1ture of entirely soft-bodied or otherwise nor- uc
L JURASSIC Holzmadcn, Gcnnmy (~cc
mally non-preservable ammal~. and 10 some ca~cs Rcpnle\, crmtacc:ul.S, cepbalopod'
show the complete spectrum of ,mtmals 10 ancient (Hauff .1nd Hauff. 1981) MC
communities. ln tlus context, the \tudy ofCambnan l·qu1
Lagcrstarten has totally altered our per\pcctive on TRIASSIC. Gres J Voltzia, France
early PhanerozOic life. What ha\ emerged so clearly DeltaiC deposit\, plant:>, msects, and tcrn-..tual orprusrru arc
is that the 'explos10n' ofhfe in Cambnan times gave also aquacic h\h and crustaceans
(Unggs ;md Gall, 1990)
nsc to 1 diversity of tonns surpassing that of living
creatures today. This concept, nowadays almost PREC
U. CARBONIFEROUS. Mazon Creek, llhnoi~
(~ee (
taken for granted, wouJd have been inconceivable DeltaiC dnd ncar10horc marine biot.1s, .md eqUJvalcnl!\ m
w1thour the Lagerstlitten The thud potential of Engl.md and Fr.mcc (see text)
Fosql-L.lgerst!itten lies in molcwlar palaeontology. A nt
whtch wtll not be pursued further here. l. CARBONIFEROUS. Scottish Carbomfcrous 'shnmp bt:Js' si on,
Thcrl' has been much tntere\t m 'extraordinary' Cru~uccans. conodom animals. tomuptcnd wurms, fish Burg
foml b10us of late (e.g. Whltungton and Conway chordates weU
(Bnggs md Clark,on, 1983, 1985; Bn~ rt al ., IIJ'llh) (19X.
Moms, I !JHS; Gould. 19lJO: Stmonctta and Conway
Moms, 1991; Bnggs, I Y<Jl; Allison ,md l3riggs, Bn~
L. CARBONIFEROUS. East Klrkton, Scod.llld
1991 ), both in the processes of foss1lizauon and lhe tive 1
Hot \pring depoSit:>, plants, arnprub1Jns, rcpulo. scurpt(>n<
nature of the biotas tllemselves. lt is surprising how (R.olfe. 1988)
Burgess Shale fauna 409
M. OEVONIAN Gtlboa, Nt:w York Sute
l:.arly terrestnal biot:a. spu.lcr. (wtth ~•lk \pinnercts), )>'>CU-
dolcorpions
:!\hear er al., I<J!l4; Selden fl ,1/., 1991)
In 1909 the American geologist Charles D. Wakott
DEVONIAN. Humrilcksdlll•ft:r, Gem1any
cc text} wa\ engaged 111 a reconnat~~ance 'urvcy of thc
Cambrian geology of the Mount Field area m
l. DEVONIAN. Rhynie, Scotl.md
l!ot 1pring dcpm•t, plants, c.trly terrestrial cwsystcm 1•1
g Selden and Edwards, 19H9)
M, SllURIAN. Hcrcfordshtre, England

Anhropods and worms from CaC0 conucuom m .1 vol-
3
liiiC ash
llriggs tt al., I 'J96)
M SllURIAN. Ll-:.mahaf,'OW, Scotland

h;h, arthropods, chordates
Rttchie, 1985)
SllURIAN. Wtttkesha, W~wmm

-.rn,ropods, conodont amnul
:'Atkulic et al., 1985)
VORDOVICIAN Soom Shale, South Afrtca

l nodont ~runuls, other 111vcrtebntes
[Gilibon et al., 1<J95)
ORDOVICIAN. 'Bcechcr's 13ed', Utica, Nl·w York

·· nozed tnJob•tcs with .:1ppcnd.1ges
Clme, 1972, Bn~>s l't a/, ICJIJ!a)
J CAMBRIAN. Swedish '01'\tcn' fauna

t text)
~ CAMBRIAN Buff,'CSS Shall• F~una of Bnmh Columbia md

(b)
~mvalents 111Utah (see text)
CAMBRIAN Chengjiang t:IUn.l, CtunJ

r~xt)
ltECAMBRIAN. lldiacan and Avalon faunas

eChapter ])
~ number o( these arc selected for fi.mher discus-

lOO, beginmng with the ex.hausllvcly researched
Burgess Shale. (N.B. Smcc the fauna has become Figure 12. 1 (a) Morrello sp/enclens (M. Cam.), Burgess Shale,
.ll known through the syntheses of Con way Morris British Columbia, s~imen from rile Geoloslicol Survey of
'J82), Whtttlllgton (1 985), Gould (1990) and Conodo with most of the body preserved though posterior
nggs et al. (1994), rcfrrcnces to mdiVtduaJ desctip- cephalic horns ore damaged (xA); (b) Opobinio regolis (M.
Com.J, Burgess Shale complete specimen in lateral view (xl
;ve papers arc only giVen where din..•ctly relevant.) approx.). (US Notional Museum no. 57683.)
41 0 Excephonal faunas; ichnology
British Columht.t. l.Hud.l. In tlw process he found a expla111s rhc atnntdt') of the anun.Js m the ~edunent
singll' dt;;Jodged ~lth ot rock contatmng a remarkable .tnd the completeness ot" mo~t of the 'Pectmem •
•mc.:mbl.Jge of prt'\ tomly unknown fossils, and he
went b.tck tht.• nt. 't )'l' u to find the horizon fi-om
,.,Juch H CJlllt' W ht.•n thl' productive sequence in Arthropods (Fig. 12. 2)
the Middll' ( tmhn.m Burgt.''' ~hak had been suc-
cc~,fi.tll'l' lm.ttl:d, ht.• t''t t'v.ltt•d Jnd mtemtvcly qtur- Arthropod' .1noum for the l,trgc't fraction o f the
ricd tt. dl't·ovcnng nut only tnlobnc., wuh their biorJ (37"'u). The tnloblll'\, of whtch Okll!>itfo Ius
appcnd.tge' prcwrn·d but Jl\n t \'l'f) dtH'f'C suite of preserved appcnd,tge.,, art• m.nnl> bt•nthtc bur there
other arthropod,, numt•rt>lh wonn\. :1 tunous crea- are :.tlso 'omc pl'lagtt Jb'll<.>stids and t:.od1~1ds
ture hke tiK mndl·m onyt.hophoran Prripatus. ;1\'arc~oiLJ, wh1t h ha., an unminerahzed c:xm~dc:ton. IS
cchinodcrnl\, bratltH>pud' ,tnd l'.trly molluscs. as a unique kmd of trilubm• w1th J l.trgc tepluklll and
well ,t, \pongc' .md algal. Thnc \\ere .1lso many a posterior ,\htcld (thor.KOPY!Pdium). However. 11
other aninuf, whtllt ,lrt' tlllt .1\\i~•n.lblc to .my living has un i r:.unou~ ,ltltenll.le Jnd typically rnlt>bltJP
phylulll. Walcotl\ l(ll.lrrytng actJvitics between appendages. Tc_eapdre tS .1 ~econd soft-bodied rnlv-
1910 Jnd 1917 produn:J ~l'vt•ral tt•m of thousand~ bitc and ir is very l::tr~e. It hJs a n~phalon .md pygJd-
of specimens fi·on• what he <.-.tlkd the 'Phyllopod ium, b ut th e thrL·e divisiom of the thor.lx embrace
bed' .md rc,ulLcd 111 diver'e publir.\tiom by himsdf three or four hotly \Omitt·~.
and other authnN. In JW,7 196H, under the direc- T he non-trilobite .trthropods of the Burgc:~' %.1k
tion ot 11.13 Wlllttlllgton, m.my thous.mds of new were untji recently dassi lied togt.'thcr as C!Jss
speumem. mdud1ng p.1rt~ ,ll!d countt.'rparts, were Trilobitoidea, smce thc1r Jppendages seemed ttl bt·
collected li om W alwtt \ phy llopod bt•d and from a generally bir:llnous with leg and glll brancht·~ ltkc
level (rhe lt.Iynwnd <~uJrry) 23 m hi~her in the those of mlob1res. I Jo-.ve\'cr, recent work h;_~
section and the.,c h .IH' been 111ten~t\'dy studied shown that rim ~11ppmed re~emblance ~~ ev1Jen
Sll1Ce. only 111 a tt'W genera and even 111 the~t> 1s not e'pe-
All the ~olt-budied Jnd rhm-~helled fossils are cially close. The poc;sc~~lon of btramom limm ~~ n
flattened, prc~crved ,1\ ,1 dJik tilm which wholly or any case no more than .l '>ympleHomorphr: a hC't-
partly rdil'et\ lighr; pyrite mJy be a~o;octated "'ith ttage from an ongmal arthropod ancestor with ~c.-n
the film. This film romi\L\ of calctum J.lummosilt- ally umfom1 ltmb~. :111d therefore no gwde to
c.lte' wuh addinonal nugnc\lum m the more refiec- affimry.
nve art.ls. Tht• m.~joriry nf ~pcomcns be fiat upon ,\l,me//,1 sple11d1'11s, the commonest and perhap, tht'
the beddmg phne,. but maJw othcl"'i be Jt an most clcgam of rhc arthropod taunJ, has a wedge-
obliqul .mgle to tlw bcddmg wtth their spines and shaped ceph.tltc 'h1eld With tour long. backwardly
.tppend.l~c' .1t d1tli:rem kveJ, (hg. 12. 1). In either directed \pmc\, the: po,tenor pau of whtch h.J\e
oncntarion nf pn'\l'TV,ltlon, dlffcrenr 'urt3ces of the crenul.ued nurgtns. On rhc ventral surface 1s a two-
body ":par.ne on part .1nd counterpart, so both are ~pincd l.lbrum ne;lr which .1re :lttJrhcd the two pam
needed fi)r .1 fi1ll inttTprct;'ltion of 'tmcn1re. Since of amcmuc: rhe tir<>t p;'ltr arc long, Ae>.;ble, multt-
Wakott d1d nnt m.1ke use of borh faces for study, jointcd rod\; the second pair are stx-sq,.,netJted '>'1th
'ome ofh1' intt.'rpn·t.Hwn' h."'Vt' been found to be in sctose distal joint,. lkhind the he::td is the cyhndnCJ.!
error. segmented body which is devoid ofpleurat·. Each of
Thl· Bmgc'~ \hak WJ\ app:trendy deposited in the 25 somitcs h.:~~ a pair of biramom appcn<hgt"\
relativdy uc.:c.:p w.ttn nff a submarine limestone which possc~s .1 joimcd walking leg :md a fc<tthcn
c'rarpmcm on or near .1 deep-,ca fan. T he fauna gill branch Jbove it. T he l.ttter wa' probably capahlt Figvt
lived on the mud o,urt:lll' below the bank or swam of rotation backward, Jnd fc>rv•arth about a hon· Plene
JUSt above it, .1nd ir ".l' on·rwhelmed by a turbidity zontJl a.-a._, and the combmed rot.lry effect- of ;ill gill oppe
(x l );
current nf hne \U~pL·ndl'd o;edlmt·nr which flowed branches could have en.thled \lmrl'ila to ,._,,m. RJrr Bruto
down the ,Jopc Jnd trampurreJ the mimals to an specimen\ art' preserved with the intestine J~olate-d
anaerob1c envJrolmtc:nt, nut f'Jt .lway, where rhey from the hod~ The affimty of ,\ fmrdla with other
were prc:\crvcd. lr i' thu.,, w1th reference to arthropods IS \ttJI umcrtain. The rescmbl.mct
Se1ladwr\ '~hc:mc. ;~n ubrution deposit. This between the Cambnan .\l,mdla Jnd the Oevonun
(b) Odaraia
(c) Plenocaris
(a) Marrella
(d) Canadaspis
(e) Burgessia
r:;~
' (g) Actaeus
(f) Sidneyia
(h) Emeraldella
(j) Leanchoilia (I) Naraola
Figure I 2 2 Arthropods from the Bursess Shale: (a) Morrella, dorsal view (x 2 approx.); (b) Odaraio, lateral view (x 0.75); (c)
Plenocaris, dorsal view (x 1 ), (d) Conodospis, lateral view, showing structure of the appendages (x 1 ), (e) Burgessia, dorsal view and
appendage structure( >< 1.5), (f) Sidneyia, dorsal view (x 0.5); (g) Actaeus, lateral view (x 2 .5); (h) Emeroldello, antero-lateral view
(x 1); (j), (k) Leonchoilio, dorsal and lateral views (x 1); (I) Norooio, dorsal view, showing appendages (x 0 6). (Redrawn from Briggs,
Brutan, Conway Morris, Hughes and Whittingtan, see text.)
412 Exceptional faunas; ichnology
Mimt•tastt•r (4. v .) Ill ay pomt to .1 real affinity and it. There i~ a large anterior pair of eyes and a mamve
indicate the 'urv1val ul thi' \tOtk until much later, tail with three large flukes which extends pmtenorly
but on the other h.aml the 'imil.mttcs may be sym- from rhe carapace.
plesiomorphlt Srdneyia is the mo~t Jbundant )Jrgc arthropod
The \econd t ommonc\t ~pcue~. Ca11adaspis pcr- from the Burgess ShaJe. I ~ body t~ \Upcrficially
ftaa, I\ Jt prt:\Cilt regarded .lS the C.liUeSt po~invdy mcrostomc-likc and rcnmnate~ m a Em-hkc tail, but
idcnnficd cru\t.llt:Jn, 1t I'> a phyllocand w1th a cara- the presence of antennae and the Jb~ence of che- 1
pace covenn~ lllO\t of the body. There are two pairs licerae suggest that lt IS n ot a nH:ro,tome. The legs,
of antennae. a rnand1ble p.ur and ten pairs of bira- however, wh1ch are prov1ded with gills posteriorly. t
mous appendage,. h wa~ probably a benthic feeder are m some respect~ not d1ssnnilar to those of L
and IS often found 111 clmters. Limulw. Gut contents, whtch mdude small trilobttes tl
Hur;{essia hdla hJ, .1 l.1rgt\ convex carapace cover- and hyolitbtds, show that this large animal was a t
mg the whole body extept for thc terminal tail predator. The smaller Emaalcldla is again rather tl
spine, "" hu.:h emerge~ under a posterior indcntatiou mermtome-like, with a long tenmnaJ tail spme, bur JJ
and 1s nearly tWilC a' Ion~;~, a~ thc body. The body is has antennae and biramous limbs. Actacus ts of le
rod-like, and the tephalk re~on ts followed by somewhat similar morphology. e:
eight segment~ ;md .l telson in front of the spine. A There are several o ther, rare, small arthiopod p.
pair of very crenulated k1dney-shaped organs occu- genera, such as Habt'lia and Molaria, which extend tl
ptes the lateral parts of the c.lrapace. These bave the range of diversity, :md the list gtven here is b) d;
been mterpreted .1s dige~tlve divcrriculae. There is a no means ex.hausnve. re
pa1r oflong, multuomted, umr.1mous antennae proat
jectmg m from of the carapace. The cephalic region PI
bears three pa1rs of b1ramom, jomted leg.;. The coxa Lobopods tn
of each bears :m mncr branch of o;1x podomeres. w
tdentical Wtth the ~even pam of leg. on the rrunk. One of the invertebrate!> originally described h} fo
The outer branc-h of the,e appendages has the form Walcott tS Aysllt!ia (Fig. 12.3h). lt t'> an annu!Jted. 111
of wh1p-ltke flagellac The walkmg legs of the rrunk catcrpillar-\hapcd creature, wnh an anterior parr an
bear small. latcr.ll. kaf-ltke plates which were prob- of branched appendage\ .md ten followtng p.un of
ably gtlh, o;ccmmgl) attached to the coxa. uf short comcJI, and ltkcwi\e annulated 'legs' lt
PlcntJ<arit m.1y be a pnnunve crustacean. lt<; resembles the modem PenpatuJ (Class Onycho- be
btvalvcd carapace covcl'! the antenor part of the phora): a terrc~trial umranuan which lives in cbmp te'
body, which heal'> une pa1r of antennae and proba- soil in the JUngles of Mina\ Geral\ and elsewhere in an
bly three pan~ of mdetcnmnate appendages. Behind Brazil. The chamcters of Penpatu.> arc tn many way" ~P I
this the 12-set,rntented body terminates in a caudal intermediate between those of anndids and arthro- he.
furca; little IS known of the hmb morphology. The pods, prcservmg, for example, the p.1ired scgment..U dts
affinitic~ of Plr11oraris arc unclear. It is not a phyllo- excre tory organs of the segmented womts, together m
cariJ. even though lt ~upcrficially appears to be so. wjth an annelid-like eye and a muscular body wall Ch
Waptia is a ~hnmp-likc fom1 with a carapace and covered by a thin c uticle. Yet the coelom i~ arthro- Ha
long antennae. Lt'alldwilia (Ftg. 12.2j), which was podan, and the presence of tracheae - ramifying (R .
probably 01 Jctntus fl'cdcr, has a long trilobed body capillaries connecti ng to the outside and bringmg m /In
with a well-dcfmeJ u:phalir part and trunk seg- air to the tissue like those of insects - links Peripatru rhe
ments wtth pleura. The great appcnd1ge of the head with arthropods. The l e~:.rs of Pcripatus move m app
has a b.1.sal JOint wJth (i)ur podomercs of which the metachronal rhythm but Jre Lli1JOinted. Their ng~•l. ~CC I
dist.al one j, dawed and with a long extension. The ity depends upon the <tntagormtic operation of nnu· 'rer
biramous trunk .lppcnJage' lllvc a leg and gill des against coelomic flUid, so that they can lengthen frot
branch. and be rigid wh1le pres\ing backwards, thereafter po~
The rare Otlaraia pu,sCS\t'\ a long body wtth up to ~hortening before the next forward stroke. PaipatllS Ha I
45 trunk somttes, With btramou\ hmbs and a is highly adapted for terrestnal life and ts wdl lob<
remarkable carapatc 'u \tructured that 1t resembles a defended, being capable of entanglmg predaton b'TOI
cylinder with the appendage 'erie~ ~:nclosed within with a sttcky ~ecrenon from ~ands beiO\\ the t'ye 1\ k
.-lys/te,,ia differs from Pmp.1tw 10 '011\c rJthcr funda- plates
along tht· body. each \VJth a m·r-lik~.· oma-
mental anaton ucal respects, and nor only 111 being ment
and usually a short spine. Such plate~. \Vh1ch
marine. It is probab ly not a true onyrho phoran . but often
appC<lr m residut:s of dissolv ed Lower
•~ best rot1Slden:d a~ .1 ·Jobop oJ', a tenn
which Cambr ian lunc~tone, wen: d(;~cnbcd long
embrac es all such unir.m uam. mdud ing Perip<1fus. before riH'
~oft-bodieu lobopo ds were known , bur sinn~
-lyshe111 1s often found .l~~ortated w1th sponge~ .1nd plates the
are now unders tood to be part of the 'lobo-
may have fed upon dwm
pod', the whole animal now b1.M''> the name. In
Opt~lmna 't:l1•1lls ha' m donga tcd, ~cgmented additio n, the 'spmes ' of Hal111nge111a (onpna lly
body, in whiCh the hc.td po"c~ses five mu~hroom though
t to be the 'leg<>') are homol ogou-; w1th tho\c
hke eyes. From tht" front of the head extend~ a long,
same net-hk e plates, but w1th a much more elon-
flexibk- proces~ tennm ;tting m two groups of spines gated
centra l 'thorn '.
tacmg each other m a pmcer- hkt" fashion . Behind
Lobop ods \\erl' a <;lgmficant compo nent of tlw
the head i' an elonga ted cylindr ical body of 15 \Cg-
Cambr ian r.1diation. Some of the other pi.'Clui,lr
ments with 1 t.ulpicn • havmg upward ly turned Burges
s Sh.tle animal-. may belong here, mclud ing
lobes. All the segmcn l5 bear .1 pair of append ages, (Budd
, 1996) the large~t of all, A110111alomris. Tlm Ius
(lch being a lanceo late gill-bla de overly ing a Rat
a segme nted, Ratteo ed. diamo nd-sha ped body w1th
paddle-like lc)be. These wcrl' tixed and quite ng~d,
11 pairs of closely '>p.lccd, overla ppmg lateral tins.
though pm~ibly co~pable of movem ent in an up-and -
Thc~e probab ly undul.H ed in a st·ncs of waves,
down plane. The AcXIbk frontal proces s could reach thus
propel ling the animal forwar d. Near the mouth arc a
round to tht• momh , \\IHch was loc.ncd \'entral ly
pau of g~ant. o,piny 'egme med appl·ndage<>, the
~nd m the pmtcn or part ~1f the ht•ad. and 1t
was catchm g appara tus of J predat onal hunter
probably u~cd to explort • for .md convey t(1od to the structu . fhew
res .1re sometime~ prc~ervcd in J~obnon. and
mouth. Budd ( 1996). when romp. mng Opalmua were
onrc believe d to repre~ent the .1bdom en ot .1
with new and smular m.Ht'ri;tl from Grcenl .lnd, crustac
ean. A circlet of plates mrrou ndmg the
found lunb' belo\\ the bod). These an· nngcd . mouth
, and tonnm g a d1aphr agm wtth ~erratl·d cut-
mflatable Lrnbs of lobopu u type .•md tln~ bizarre
tmg let:rh. was prcviou~ly like\" i\e known only 111
Jn1Jlla1 i~ rcmtcr pn:tt•d a\ .1 lnhopo d, md
presum ably •solaoo n. For some 70 years tillS
ufbent hic habit. org-.m, n,tmeu
Pqwic1. h,1d been beltew d tn be a Jellyli, h!
AnO[her Burgc' i' \hale ('UnmHy. Hal/utrl!llllll, m::~y
hrlong ro th1~ ~amc ~tock. K.rwwn !Tom only a very
lew spccm1 em. 1t was on~nally rcnms rructcd a_~ an
Othe r inve rtebr ates (Figs 12.3 , 1 2.4)
.mimal that walked on seven pain of stiff movab k
'pines. 'uppor ting a ('yhnd ricallm nk \\,th a globul ar The t•arhcst
oLul crino1ds. &/mraf l>mllllj (Fig. IJ.51). I\
head, trom wluch 'even ventta l tent.Kies arose. T h e found in the
Burges s Shale. ll lu~ a l.lrgc conic.U calyx
discovery. howev er, of spmy tJterp11lar-like anunal s of
1rrcgular plate~ .tnd plated unist·nal :mm. There arc
m the Lower Cambn :m Cheng jiang Lagerstiittc in
.-llso cocnno ids and a possible holorh unan and
China, whilh have mauv feature s 111 comm on with cdrioas
terotd. Mollusc~ arc rare other than hyohthid~
1-falluogrma, suggc't .' an alterna tive mterpr etatwn (if these arc m
deed molluscs).
(R.amsk5ld and ! I nu. 19lJ 1; Oengt~on. 191:1 I) For
The curiou s plated 11'irmx~c~. \Vlth 1ts vertu .tl
H,JI/uc~~elllll, turned ups1de -do\\ u I' very sm11lar
to defens ive spme~. may welJ belong to a d1stmct phy-
the Chines e an imal The latter Ius paired lum. Lophop
horate~ mdude \t,mdar d Cambr ian
1ppendages each wnh a tenmn al cbw; and it now inamc
ulate brachi oporls and .1ho the pt·culu r
1ccm~ most likdy that the unpair ed appcar ann·
of Odt>llfOJ(rip/ms. This organi sm ts about 6 cm loug
cntacles' (in wall-.mg legs) of Hallungm111 re\ult.~ .md
1t~ body j~ Rat and annula ted wtth a poor!}
lrom impert ect prtserv anon ol o n e '>et due to the
define d hcJd or \Cm.iClrcular tonn. On the he.ld arc
rosition m \' hich tt was burit:J Tht: b1zarrc a pair ofbrer
al palp~ (~cmory orgam ) of rather mcll.'-
Hallun.~cmn IS therefo re probab ly .m 'armou
red tinct morpholob'Y and also a bil,ttcrally ~ymmctncal
lobopod'. an onydlO phoran -hl-.e animal of the same median structu
re fomun g a p.ljr of loop,. Thss .lppa-
lifOUP as the Clunc 'c fos~ti. Th1s latter. mnden t,illy, ranss lie~ ;lt the front end of tubula r gut and bear'
1' known a\ Mia,•tilttyrm. It bt:ars
.1 dum of llVal some 25 thorn- like 'teeth' The teeth were oog~-
(d) Opabinia
(a) Burgessochaeca
(a) l
Fi9UI
pulid
(;.c 2)
~
("~,:~~ /1~
10 Odontognphus \..:._J (j) Lingulelfa T,
dly
prob
Figure 12 3 Worms, lobopods and other invertebrates From the Burgess Shale: (a) Burgessochoeto (x 3), (b), (c) Anomofocoris (bl puhd
ventral view and (c) great appendage of some rn lateral view (x 0.25); (d) Opobino, interpreted os a lobopod, ontero-loterol view porta
(x I); (e) Holfucigemo, interpreted os a lobopod (x 1.5); (f) Odontogriphus (x 0.5), and (g) reconstructron o( its lophophore and ten
todes: (h)Aysheio, a lobopOd ()( 1,5), (j) Lmgule/lo, a linguliform brochiopod (x 3). (Based on various sources, chiefly Briggs el of,
one
1994; (d) based on Budd, J996; (e) based on Romskold and Hou, 1991.) more
have
by pr
nJJ!y thought to bt• pmsibll' conodont'>, but since fonn and i.~. incidentally , bilaterally looped likt: ti1Jt Th
the d1scovcry of" tntc 'conodont ammal' (Briggs Cl of OdotlfO.~riplllts. There scent\ w b~ a good ca~e for and t
al., 19H3) of <.JllltC difft·rent morpholob'Y· this now aligning Odottlt>~riplws with the Lophophorata as an l ~1 11XI
seems unlikely. Thl• 'tt•eth' of O(hllltl~{!lipltus were early dcriv:~tive of the ~.tm c supcrphylum. spo n~
prob::tbly not bitinp; or rasptng tet·th; they have been The only annchd~ n.·prc,~ntcd arc polychaerr gul:~re
interpreted a~ tht· ~upports for :1 food-gathering worms (e.g. Btt~(!t'sso(/racta) .md the~<.· played a rcb~ biolo~
Jpp.lrJtus hJVln~ tht• fimn of .1 'tt·nt.lcul.lr lophophorc' nvcly minor rok- 111 the f:lun.t None of the~c hJvc like /·
(Conway Mom\, I'J79) \ur h rcnt.1culJr lophophorcs J.lws, for these were not ,1cqwn·d untJI the Ordo- chcvrc
arc found 111 modern br.Khwpods, tube-dwelling vician. Cmradic~, ltke modem polych.lctt..'\, ha' two ea] of
phoromd 'wonm' .llld bryozo.1m, whiCh arc aJl paired 'iet~ o f fldrapodu (bundle' of \tiff bmtb) for The
commonly bnh•d ro~t·th~r m \)upt..•rphylum Lopho- e.Kh segment. Th1<; " the onl> gcnm th.lt might be n:n·tn
phoraca. rn rh c.·\~ rlw lophophor~' lt ka\t Ill it<> iniri.tl related to .1ny modem f.1mily: none of the others w:1ys 1
sr.1g~s of devdopmt•nr, 1s of remJrkably comtant arc. be .1 '>U
Burgess Shale fauna 41 5
(b) Ottoia
(d) Dinomls chus
111 Thaumaptilon
(c) Wiwaxia
''!P.V 12.4 Variovs other Bvr9ess Shale invertebrates. (a) Thoumaptilon, a possible
pU~.d worm in its burrow (x 3); (c) Wiwoxia, in lateral view (x 1); (d) Ediacaran survivor (x 3); (b) Ottoia, a pria-
Dinomischus (x 3); (e) Crvmillospongea (x 1)· Vauxio, a sponge
·21 (Redrawn from Briggs el a/, 1994, also Conwoy Morris, Rigby and Whittington.)
There are tive spi.'Lte' o( pnapuhd~ whos1. prc\ent -

\ representative~ have .1 simtl.u retr.lCt abk spiny Significance af the Burge ss shale fauna s
INI\CIS and annulat ed hody to Lambn an pria- Ecology
uhd' (~·.g. Orwia). Modem priJrull ds ;~re very unim- The commu
mty strut"tu re of thc Burgt'\S Sh,1le
:r.llt. ltvin~; mautJy J\ wld-w Her bc:nthO\. though
faurJJ at Its type lol.ality has been arulyse d by
~ ~rroup 1s endopa rJ\Il ic. C.1mbn Jn t~mm arc
Conwa y M oms (l986). Shdly fuunas compri se 110
;~>r~ divel'\c but. as inbuna l omivo re\, 'ecm to
more than 20"{, of the genera (and perhap ' no more
JVc been dtspbct 'd from their onginJI cnvmm
mc:nc than 2% of individ uals). The soft-bo died or lightly
prcdatorial polycha ctc:' later tn gc:ologJ.cal ttme. skclc:to nin·d faunas are far more abunda nt and thc\e
lhm~ arc a fe" crud.m. 1m .md a vel) .1bunda nt
can gcm·ral ly be analy~cJ in terms of feeding habtts,
nd nch f.mn:1 of spongt· s, ((' g. Crulllll/tlSJlllllgia. .md J food
web can be establis ht'd. In the benthic
.(t.l) mmt of whtch L.!ll be referred to the dcmo-
as~cmblagc there wert' vagrant bentluc deposit feed-
)nt,;es and hexactm dlids; tht•rc .ue Jlso some liner~ (m:~inly
arthropod~). inf.1unal camivo rcs and
1lt~ brac hiopods . At the other end llf the
scavc:ngt·l'\ (chtefly priapuh ds). epihun al ~u,pcmtOn
~'teal 'Pt:ctru m IS the: prim1t1ve and ratht•r fi~h-
feeden (~ponge~ and brachiopods) and iul;wna l ses-
' Pikaia, which scetm w h.wc a notoch ord and sile suspl·m
ion feeders (hcmic hordate s). There ,1re
·otHhap cd blocks ut mmdt (myown u.•,) rypi- also large
obv1ou s predato r' such .1s Arwmalocaris.
of chordates. and Ottoia ~cems ro have hcen c.nu1ibali~tic. The
The po,mon of Duwmi~dws I\ undear . bur tht dJVersit)
of feedmg types and the1r indepe ndence
mtly dc~cnbed ntarllltllp Ciitlll, so suml..tr 111 many shows qu1tc
clearly that the fuudanl.eut<tl trophic
I' to the Ed.iac ,uan Clwmiodrsms. llll\ prm c
to stnt\.Ut re of m:~rme mctazo .ws was alread}
c~tab
a~urvtvor from the lJtt'r Protero zo1c.
l.tshcd by the MHidle Cambn an and poss1bl} earlier.
Geographical distribution mg divcrm;. Even though Ha/luq~ttllcl (formerly

In M1ddk Cambnan tllllt:\ th~: LJurenti.m conti- ~een as the most bizarre and peculiar of ill thl"
nelll. in "'11ll h tlw Uur~"'' Sh.tk o~:curs. lay isolated Burgc:~s anim.1ls) ts now bc,t interpreted ,,, a much
cllld in Jll lqu.Honal pO\Itll\11 It nught be expected more 'rc~pcctablc' o nylhophoran-hke 'lobopod',
that other equt\.',lk·nt .1nd lOntempor,lJ1eous faunas the spectrum of d1versav 1\ still cxtraorduury. Thcrc
would be tound 111 L.lllrentiJ . .1nd thio; has indeed thus arises a raxononul conundnun wh1ch lcd
proved robe: \O . In Untl\h Columb1a. new discover- Whimngton (19HS) to 'JY ' If we knt:\\ more ofd1c
Jes from .u lca~t .1 dozen \HCs (C..olhm et .tl., 1983) soft-bodied t:Hma~ of rhe Phanerozoit, our rnJJOr ~VJ
show that loc.lU) .•1t lc.m , the: Burgess Shale fauna subdivisiom ofinvertcbrarcs nughc be \'iewed ~Mfcr :1111
was Widespread. J'hc tau11.1S of these locahtics arc ently'. More recently Gould ( 1990) has eloquently W<l
broadh \lllHI.tr to tllO\l' of Walrott\ qu.my, though explored tlus whole eo m ept. l n trJllllg the lustory Shl
some qlllte 11C\\ .ltlllllah. ~:.g. the cJrly chcliccratc of rese.1rch ht: \hows how Walcott. in h1~ othcrwts~ bra
SallCTtlcuns ('~.mr.1 Cl.lw,'; 13nggs .md Collins, l 1>HH) :tdmirablc work, had tncd to \hoehorn' rhe .unnJJls Sllf
have been reponed mto established systc m:~tic categone~. He (hd nor nst
Further :~tic:ld thl'rl' Me: othc:r Lmrcntian Middle realize that the va~t dl\•cr<:iry wh•ch the Uur~c~s a re
C:~mbri.1n I agcro;c:ittl'n, \~ hid1 arc less well known Shale animals rcprcsentt•d was the ke\ to ,1 new bit\
bur of ('Xceprmml intl'Tl''lt. In Utah (Rohison, J 991) dimemion of evolutio nary th.inkmg. I h: 'interpreted 111:1
there arc no l es~ th.ln four. in the: Spence, Wheeler them alo ng the path ofleast resistance' a~ 'pnllUtiVC' aJJC
and M.ujum Fmm.lnoiiS, where: ~oft-bod1ed animals vero;ions of larcr 1mprovcm cnts'. Only wht•n rbry Bw
.1lso occur with Cl hinodc:nm. trilobites, bra- hJd been restudicd by Whittington and his tt•am and bea
c hiopad\, hyohrh' and sponge' Of the non-trilobite new concepts had t.1kcn hold, was the full 'lgfllli- r/111:
anhropods, wnrm' .md ~pon~t·~. many also occur rance of thcse Cambnnn orgamsm~ realized lt ~~ l\Ja1
111 tht· Burgt'" \ll.lle. The Kmzer' Shale 111 thc1r divcrsiry m cenm of body pl.ms that is unpm- bnc
Pc:nmyh am;!, poorly cxpmed and terronizcd ram; rhe gencr.l seldom have: more rlun one spent- son
I hough lL 1\ h,J, hkn... l\l' de 1r \tnulantic:~. The avail- and they arc 'epararcd one from another by "tdc ~•de
able cvtdcnn· rht·rct(1rl' \ltggc:~t~ that. at least in morphological gap,. Thu~ c:.1rly Phanerozoic lili: des<
Laurenna, Burgt'\\ \h.lk b10t.\\ arc only unusual i.n shows an cxplmH)n of d1\iemry. far greater m \l,mrt) Mo
the WJY thl > .IH' prcsavc:J md not m thl'lr ta.xo- than today·~ whole (\Una We do not find a t,rrJdu:t1 and
nomil romcnt :md Ji,tribution. Such ftu nas may in crease of divcro;iry bm lJUHc the oppmite. The .lnd
mdced haVl' hl't'll ~lnb.tl m rhe1r disrnbutwn. The 'trc:c of life' •• ri,ing from a root and ldrer flouri,hmg W ltl
new dt\cOvt·m·, of lower Cambrian but ,i_nular fau- into many branchc~ 1s unrcali\nc - a better g:~rdcn tcre
na' m Chuu .utd d~l'Whl•rc (q. v ) nmduce ro the .malogy \.\-ould be a 'grJ\\ l,twn'. The: E.triV 11• ditfc
vtcw (Conw.1y Morn~. 1'>1-N) that Burgc~~ Shale Middle Camhrian record rc:vc:ah a gre.H buNt of C:Jn
1:) pc f.1una' rt·prl'Wilt .1 "" 1ddy distributed offShore radi:~tion in \.\hich marine orgam~ms divcrsilicd tos~1
bcnchic fauna, t)Uitt· lnn~-r.tngl·d (L.- M. Cam.) and ,Jlong man y parallel though d1stinct Iincs. The elim- and
w id1 an t:voluuon.ll ily tllmcrvati vc: a~pcct. ination of many of thc~c typl'' through increa.,mg mg t
competitio n kft the survivor-; to become the prog- icJ I
Diversity enitors of today\ phyb. Thc:rc: i~ no indication ofJ ex cc
The Burgc~, Sh<Jk· t:tull.l Jt Jb tvpt: loc.sliry m Briti~h single conunu11 ancestor, a nd m any auLhoriUl"~ now nic h
Columbia is n.r1 aordlll,\1 ily Jivcr~c, with over HO suggest a po lyp hyle tic organ for the metazoan phyb ll!Ul
specie' 111 11 tJ 't·pat,llc gcnc1<1. lt fonm J major part (i.e. broadly ~ imil:1r kimb of animals arising indc- A
of the: 'CJmbnan cvoluuon;tr) CtunA' (Chapter 3), pendcmly in difTcrt·nt part\ of the world) ratht·r tlun CJm
which othcrwl,t' j, knowu only from the bard- a single JnceHral form for c:~1 c.:h phylum. Tlm
~helled trilobite,, m.lllitul.ne bracbiopoch, etc. Ben~
Whereas 'ome ~em:r.l, \lllh a' the crinoid Persistence pods
Echlll<lfllcM7/II), the cru~t.lt:can Cm~<~daspts Jlld the The Burgess Shall' tawu was ongmally known on!) Burg
che!Jceratc Smur,,c.ms, ar~: the earhe~t known repre- from the M1ddle Ca111brian ut Briti.'b Colwnh1J m ad
sentanve~ of late!, unportant group~. ''ery many oth- Some remarkable rc:u~nt d1\covenc~. ho" ever, 'ho~\ h all..1
ers cannot bt: d.\"Jitctl. In term' ot the number of that many of us c:lcmcnc~ can be traced right do" n mg I
different kin&. of body pl.lm the) show a bewilder- ro the ba.~al Lower Cambnan. Of these, the n< hat kno\1
Burgess Shale Fauna 417
fauna~ .~re .lt Cheng ji;mg in \Outh em Chm.1
and m Camb nan 'small shelly f.1Un,l\'. In the
Peary Land, Green land; tht-re arc aho occurr ences Green land
matL'ria.l arc comp1 etc halkie riid anim,tls (Fig. 12.5).
m a boreh olc in northe .l\tcm Polan d. and in the
They are slug-u kc fonm some 3 ~m long w1th a
SH."rra Mon: n:J ol south cm Sp:1111.
dor-;al annou r consi, ting of imbric ating row:. of the
At Chengjian~ a 1111xed t:mna ofLo\ \Cr C.unb n.m
sdenrc s. Such an orgar untion had actu.1lly been
shdly and ~oft-bodied ,1nima.ls was chscov crcd in
predic ted, but what was quite uncxpectL•d \\<1'> till'
llJ~4 111 a mud~tnne \lllUC IKl' ckpos ttcd in
~hallow presen ce of a large round shell, one ,11 each end. The
wata. lt 1~ the nldcst tlf .111 Camb 1;an Lagt·rstattcn .
afl:inities of the: halkie riids an~ still very obswr e.
.rnd proba bly of late Adtab aman age. Str.n1-
Reg.1 rding these .md other ~uch orgam sms,
!:,rr.lpruc.llly below 1t ., a nch a'~cmblagc of ~maU
llengL~on (19lJO) stated that 'Natu re ha'i
'hcUy tossih Thl Cheng Jiang fmna I'; dom111ated by .1 \\,1} of
outs1tin..ing our most 'pecul atJve hypml w'ics'. Wlw,
hr.1donids, ~mall bJv.1lvcd arthro pod' \\hich are
in comid L•nng the Burge ss Shale faun.1s, could possi-
'upert1C1alh 'inul.u to, bm nor dosch n:latcd to
bly d1~agret"?
mtraco des. Over ~{(In,, nfthl· mdivJ du,ll' iu Lhi' fauna
lu bro,Jd pcrspc cttvc the C.unb rian cvnlu tionar y
Jrc br.1donrd\. There are only three genera of tnlo-
fauua, wh1ch mclud ed the Burges~ Shale fi•nm \\"ith
bltes (cf 401Yr, of the (llln,t m the Burges~ Shale) but
large pred.u ors suLh a' 4notll•downs, spelle d thL
many algat". 'Pllllge-;, medu~llld,, chond ropho rans
end for the qutlte d org.mism~ of the 'Gard en ol
and hyolith1ds; tht'fe .m~ also eleme nts typic.1l of the
Ediaca ra'. Once it had originate!'d, the Burge s\ ShalL•
Burgess Sh.1lc. Theo;c: mdud e pn.1puhd~. the ,c,tle-
faun.l remai ned very conserv::~tivl!. It may have
bt·aring onvch ophor an-lik l· Alim,dictytllt. Di11omis-
ari~cn in shallo w watc!"'>, :.tnd then '' ith the
"i111s and Ar~oma(,,cam, H.dfll,~l!<"""'· Lt·,wrfl,,fia and new ri'c ot
fJun"' migra ted to Jeep w<~tcr, until 1t ton wa'
Nnraoia. but tht•rt' an· also ~l'vcr.rl other non-t nlo-
driven to cxtinc tiou by later Camb rian .md C!'arly
bne arthropod~. One of thew Fu.\·''""""'' looks
~omcwhat like .1 trilobt te, but h.t\ a long.
parall d-
'rded, 'cgrnt• ntt•d tall. The,e taunao; .1re \till under
dt·~cripuon· recent rdert•ncc:s includ e
Conw ay
Moms (19HlJ), Chcn and Erdtm ann (19lJ1), Hou
and Bt•rgstrom (1990 ), R..1mskold and Hou (llJ91)
md Hou and ller~triim (19 117). Desprte stmrlantli.!S
with the Burgc!ts Sh.1k 6un.1. tht•re arc m.uke d dif-
l~rences, proba bly n:lkc nng an origir ul et
ologrcal
Jitfcrc nce. While it ~~ lnterc: ,ting ro comp. tre
Camb n:m fl\Scmblage~ 111 seque ntc, i.L'. sm.1ll ~hdly
tossils. Cheng jiang tjuna and Burge " Sh.tle t~lllna~.
and tirully the 'orsten " fllln.ts, rt would be mislc. td-
mg to Interp ret these lOo dm~ctl} as a global ecolog -
ltal succl'SSion. l:ach rs ~1mply .1 'srup~hot' of an
except ional tauna m tunc, 'Pl'CJ.thzcd tor a p.1rncular
mehe, and little ts know n of con tcmpo raneo
us,
equiv.tlent t:mna ' clsewh er{ 111 the world .
A more renmtl'y di,cov ered ( 1989) Lowe r
Camb rian Ltgcrs t:ine t Olllt'' !Tom north Grt:cn l.md.
This fauna (Conw ay Mnrri \ and Pet:l, 1(}l)();
llen~tson, 1991, Uudt!. 191Jh) indud~:, man}
anhro -
pods, polych acte\ ami pir.1puhds like thmL' of the
llurge'>\ Shale and spnng e,. An uncxp tltl'd bonus ,
m add1tion, solve~ thl· proble m of thL· ori~111 of
the
h.tlkierulb. These are mKnN:aplr sckntl "' of v,lry-
mg form li~t ck~cnbcd 111 11>(,7 .md prcvio mly Figure 12.5 A fully preserved holk.eriid from Greenland with
known onl, a_o; l~obted l'knll'nt~ in man\ Lowe r anterior and posterior shells, and o body covered by dennol
sclerites (x1 .5). (Redrawn from Conwoy Morris and Peel, 1990)
41 8 Exceptional faunas; ichnology
Ordovtl ian l omp~·tttol\ Pm\lbly the peculiar V:istergotland, Oland). "iome ye.trs ,tgo, \\bile
.\limctcl'itrr from th~· I k\'onian Humrticbchiefer sear<. hmg for conodonl\ 111 etcht'd re\tdue~ of the,e
tmght represetll the l,t\l survl\·or of thi, mmt extra- rocb, Professor KJ Mtilkr ofUonn madt' a chance
ordmary btota of ,tll time. diKovery of phosphatited mtracodes and other
Cnt\tal t.:ans, ts wdl as ,tgnostid trilobit~. .'v1any
thou~ands of spt'CIInen' h.tve been isoLued. fre-
quently '' nh the limbs tmacr. ' J he phosphattc to~''''
are of two kmds, hrstly thme \\ tth primanly pho,-
phatic hard p.trt~ (phmph.ttocopmt' ostracode<>, inar-
ricuLtte bracluopoth ,tnd conodonts) and secondly
The Upper C.tmbn,lll rocks ot <.,weden are highl) rhe arthropods 111 \\hie h the bodtes and appendage'
tossthferous .tnd ~.·omain .tbundant olenid trilobite' were preserved by ,1 senmd.try phmplutic roatin~ (a) f
(Chaptt·r 11 ), Ll\lt,tlly ti.nmd m c.tlcareom concre- which must have formed .tt the time of buriaL In
tions or lcmes ktW\\ n .ts 'or~ ten' or \ti nkstones'. some tmtanu:s the shell 1\ repl.tced by pho~phate,
The~c occur ovt·r qttHl' ,1 \\ tdc area (Skane, Preservation " nonnally cxqlllsitc, though in ~ome
(a) (b)
4
Figun
Bredo
( · 70)
W alo!
Icl case~
whic
COa l it
W hc
dtagc
orig ix
Ot
nun
q uite
Aocet
bcri n1
they ·
their I
WHh,
'shell'
Figure 12.6 Elements of the small phosphatized 'orsten' founa of the Swedish Upper Cambrion: (a) Agnoslvs, a partially enrolled when
trilobite showing a phosphotized appendage, (b) Skoro, a cnJsloceon, (c) phosphotized appendages of o phosphatocopine ostrocod. shcU i
(SEM photographs reproduced by courtesy of Professor Klaus Muller.)
baccer
Upper Comb rion of southern Sweden 419
fa) Rehbachiella
(dl limb
(e) Heymo nsicam bria
(c) Skara
(b) Bredoc aris
((0 larva
Figure 12.7 Elements of the Swedish Upper Cambrion 'orsten'

BiecJocons, o moxillopod crustacean ( · 90), (c) Skoro, a moxillopodfauna · (a) Rehbochiella, a bronchiopod crvstoceon (>< 65); (b)
crustacean (x 70); (d) isolated limb of a hesslandonid ostracod
(.< 70); (e) Heymonsicambria, o pentostomid parasite (>< 45); m
o crustacean nouplius larva (x 130). (Redrawn from Muller and
Walossek, 1985o,b , 1988; Wolossek, 1993· Walossek and Muller,
1994.)
c,l\l'S, ~~pcn,dly .lt the amen or end of the bnd).

The arthro pod\ wen: .tll \Winu mng lom1s. By far
'' h1ch was 111 life rhe mo~t h1ghl) pho~phatic, the the LOITIJllcmest are the adont pnmiti ve ostraco des
w.1ting IS rd.Hiv dy thll k .md obsl'llrl'~ the jomt~. l 't•strcJ.\!llt/ua, Falitcs .md Ht'.\5/,mdoll<l Thr'e bdong to
Where tillS mru~ the spmcs .md sm.1ll hairs m,1y be rhe Order Phosp hatoro pina, 111 wluch the angina l
diagcn encallv thickl· ned up tn thrl'C times their hard par~ were probab ly pho,ph.ltt.'. The abdom rn
on~:,'lnal thamet er.
whiLh may not have been pnrnan ly phmph ati7cd , ~~
Only small growth suges <.:1f the .1rthropod~ (<2 not u~ualh preserv ed. In 'P•te of the :.ccond ary
mm long) s<.:em to be to~siliznl in this way. ]( is pho~phatic thicke mug, the mform .ltlon g~ven
quite probab k tlut rhese mlubit ed the nutnen t-rich .1bout
the nalurr of the hmb\ of tht'\e pnnuti vt• ostr.1rode'
floccult:m layer on the 'e" floor, '" 1mmmg, d.un- 1' of the h1ghc!>t quality ; 1t 1' parnc ularly
bcrmg about .md feedin g Within it. When thev died mtrre\tin~
that these pho~ph.ltocopine limbs .Ire clmcly 'mubr
dH:y sltd dm\ n to the lllOXlt -.:a floor JUSt bduw
ro onl anurhe r "' itblll the \amt· .lllllliJI, and arc
the1r hab1tat .111d WtTl' covac d very soon ,1ftcrw,1rd~
comp. uativd y umpeci.1lized.
'' 1th .1 coaun~ of phosrh .Hlc h.Ktcn .l 1t 1'\ tlu, dun
1 here .1re now 25 lulO'\\n ,mhrop od gcner.1
·~hell' of dc.1d, umhgu ous lut tcn,1 th.lt surv1ve
~ mcluding 'J large variety of primord1al lfll\tac ram
''ben the rest oftht• ani nul h" dcc.lyt d ~here the
whose sy~tclnatic pomw n i.' ,1, yet unccn am' (M tiller.
'hell i\ rcplau~d by phmph are H w,l'i 111\'Jtkd by tlw
19~3). Numc rom nauplia r l.trval ~rages of v.mou
b.u:teri.1. '
umdl·n tified crmt.K ean\ arc ltkcwl \l pre\l'n t.
Skam ha~ a ccph.tlon wilh tive pair. of appendagcs ~upport to the idc.t of a 111lHloph~ lctic origin of the
and a long tlc~obk trunk and tail. It was a filter cmstaceam, thctc are also ~imilariues bem.een their
feeder and ~~ knm\ n on I~ from .1dult\ . •'\J,minmmia i~ limb~ and thme of !'i.f!IIO.<tll:o, which may {'X'alossek
supertin,1Uy 'imiiJr, thou~h it ha~ no speciahud and MUller, 19YO) \hed new light on the sy)tenunc
cephalic futenng .lppar.ltu!.. lt wa' probably a bot- position of the latrcr.
tom dweller ti.ocding un detrital particle<> that it Finally, the ~\\cd1'h fauna,, like those of the
stuTed up from the \CJ floor. The juven1le stages are liurgess Shale. show th<H the non-tnlobtte arthro-
also known. [),lf,z h rcprl'\C:nted only by thorax and pod faunas played a much more unportam role than
abdomen; tt m.ty hJVl' been a free swtmmcr. has generally been appreCiated, m Cambnan nm~.
Bmf,,mns ha\ .1 l.tr~l' Ulll\'.llwd hcadsludd, paired and the apparent predonunanct· of rnlobttes must
eye~ and comp.1r.mvcly umpccialized cephalic limbs. have been .ltc..cntu.Hed bv the ht~her preservabihty
There .m: seven p.uf\ l)f thoranc lunbs. All larval of their exo~kcletom. lt •., de at also that these Upper
stages an.· known. lt 1~ unhkdy to have been a filter Cambrian cn1st.1ce.1m bear lmlc rc~cmblance to
feeder and m:~y h:wc \\VlllllJllSt above the ftoccuJem those of the 13urges\ ~hale. Although dm n!Jy.
hyer. ll'lllosst•kia 1' not d1ssuml.1r, whtk Rehba£hiclla again, be purely due to prescrvatjon, the 'morpho-
has a l:trgc bt:.lchllll·ld t•ndo\mg much of the body logical and tempor:tl d t~conncct10n' of the Burge\S
.md nt.llly limb~. All thl' L',trly growth stJgcs arc Shale: fauna rcferrcd to by l.krgstrom ( 191'!0), in
known, but rh<: l:trgc'>t speL'UlH'ns arc still probably orh er words its d tsS1111ilarity from later faunas mJy in
larval. Tlm cnlst:lcc.m 1s .m .lllCL'Stral branchwpod, fact be a real evolutiOll::try phenom cnon .
.md 1ts complex tiltcnng appar:ttus has some rescm-
bl.mce to th.tt nf molkm fonm. Recently described
(Walo~'t k ,md Mtillcr. 191J4) arc ~mall pcnta~tomids.
and arc rcmark.lhlv suml.1r to thl•tr nwdcm countcr-
pam '' luch p.1r.1,1tin· thl' lunh" of crocodile' today.
It ~~ to be '''omkrl·d wh.lt they ti:d on 111 Ca.mbtian
times. The Lower De\onun Humriick.~clucfcr of the
The: ·of\tcn' cnl\t.ll't' 111' ,Ire thu' v·ery dtverse. Rhjncbnd Ius long yielded e'>qlll~itely pre,en·ed
some being progcnllof\ of modcm ta.:xa . o thers Marfish, conoid,, tnlobtte,, crust.1ceans Jnd
belonging to t'.Xtinn h'T\HIP'· Thcy were adapted to ccphalopods. m.unly pre~en·ed in pyrite. In these
ditTercm mirrolubit.tt' '' 1tlun or above the ftoccu- stagnation depO\It'i rhc pyrite wa~ fom1ed "ht're
lcnt laycr. organic compound~ rich m sulphur were pn:scnt in
The trilobite ,'l,CIIII~ Im rev cal' 'urpri~ing details of decaying matcnal; rhc'IC comhmed \\1th tron ion> m
appendage: comtrultwn. Thl·re ''one p.1ir of anten- the water to prenp1r:ne re:::, .1nd FeS,. M,my 'uch
nae and cight pair\ of lmamous .tppt:ndagcs, but fossils have been known for a long time, but surprh-
unlike those of ·,t,md.1rd' trilobites these arc much ingly complete details of\tmcture have been proved
diversified. New Jct.ul' h.tvt• also emerged about th rough the :~ppl ic.1t i on of \oft' X-rays, often \\lth
the nature or tht VLIItrJI mtegumcnt .md the stages p hotographic expmurc' taking sewral hour,. Th11
of early ontogt'll}. lg11m111~ tould not have stretched tec hnique W<lS hcbrtln in the I()J()s by W.M.
out ti1lly and H is ht!;hly hkdy th:~t these trilobites leh mann , am\ was resu med tn the early 196os to
lived for mm h of rhc time i11 .1 state of parti:tl enroll- great effect by Profc~sor W. Stunner of Erlaugt:n,
ment With J ~lightly gaping cepbalon .md pygidium. yielding <;pcctacular res ul ts: a fimd of infonnatiun
The dc,cnpuvc wo1l- on the~e 'orsten' taunas has been obtaim•d, IHH only on thl' hard p.m~ but
(Mi.ilkr I'J71J, IIJl-12. I'JH3, 19~5; Miiller aud also, in many casl's, on thl· lightly pyritized soft th· vane ['I
Walmsck, JIJHS. Jt)H6. IYH7, 1':11:1~: MUller and sue' (Srl.lm1cr, 1970; Sti.irmer .md lkrb"'trom, 1973. tjve~ l
Hinz, ]9l) I: W.1Jo\Sck, 1993) mcludes perhaps the 1976. J97R. l9H I; <)ri.i11ncr r1 ,r/., 19HO; Ber~tron now c
most preca~ely documentl'd of aJI palaeontological and Brassel, J9H4). in ha•
smdte\. it gin·s J dc.lr umght into the nature and Details of limb morpholub'Y has been rewalcd m bchm<
ecology of a C.1mhmn flocculem-layer community. Phawp.s, .-l.;tclclJI)'Ct' .md other rnlob1te gcnt:rJ (!j ,,) life: on
It pronde, new llllornunon about the limbs of the and the dct.likd \tmctun· of thl' phvllor and lrth- (sea sp
annenr crmtJCl':tlh and ! 1.1!110jfll5. While tht) g1ves tacean .'\,'ahaans h.t\ hl·en made clear (Fi!,!. 12.Sl). .A ,l\ we!
Hunsruckschiefer fauna 421
Figure 12.8 Hunsruckschiefer fauna (Dev.), Germany: (a) Nahecaris, a phyllacarid (x 0.5); (b) Mimclosler, on arthropod associ-
ated with an ophiuroid (length of bar is 10 mm). (X-radiographs reproduced by courtesy of Professor W ilhelm Sti.irmer.)
\ariety of other arthropoth llldudt· t'.lrly representa- rcprcsent.ltive of the extant pycnogonid Order
uvc~ of extant group~ or mt•mbel"\ of t:txa \\ hich are Pantopoda.
DO\\ extinct. H'l'iu/lr~l?lll<l, .1 s~ 1111phmund, 1~ unique Somt• of the arthropoth c.mnot be related to any
m having \IX, r.lthl·r than ti\'e, prmom.tl limbs present-day ta.xon. The large ovmd Chrlo11idlo11 has
belund tlu.· chl'iicerae. lt w,1, eviuently ad.lpted for a head with a pair of antcmue prOJecting forwards,
hfc on a soft \UbstrJtl'. L1rgt' primitive pycnogomds nine tnmk \egment'i and a cunom comcal rclson.
~ea spider\) bdonging co extinct ordt'l"\ .1lso occur, There is one pair of ante1mac, ,1 second pre-oral pair
~ weU ·" the tiny }J.,/,Itw/n·,,, tlw mo't .metent of .1ppcndages. .md tour pairs of uniramous
Jppcnd.lgt'' w1rh gnathob.N?\ 1n tht· ht·ad. Etght

p.uT\ ut' bir.1mou' appt·nd.t~l'' .m: prnent in the
both The ,\l,mdla-!Jke uthropod \ limL'ft~sccr, about Tht• Penmylvanian bed~ of lUmo" l1.1n~ long been
h.tlr'tltt' known 'Pt"nmt·m of\\ htt h .m: found asso-
oatt•d Wl{h ,m ophmro1d (hg 12.Hh). w.t~ known in
'trip-nuncd for coal. Over w1Je .1rea' rh<.· noduiJr
\hale ov~·rly111g the coal has been n·movcd ro 'poil (
brrm~ morpholob') pnor to \:)rtinncr .111d l:kr~rrom's tip~. Crom which the weathered ironstonc nodules
work ..IS w.t~ I achonisitl. ,} large .lfthropod with a o r conaclions have proved the sourn of .111 mll-
btv.tlvt•J t.tr.lpace JHitllt't,JSII'r, thwugh X-ray pho- ~tandingly divt·r~c flora and f..Jun.l (Nttt•tlo.J, 197'1,
log;r.tph,, '' now kno-.vn to lun• .1 pJir of ~talked SchrJm, 1974, 1979b: Thompson. I 1J77, I1J79).
eye,, 'lout JOintcJ Jppcml.lgc' on the head wirh
pynttzt·d 'tr:md' of mthck .•md .1bour 30 pairs of
The\e noduk~ provide a valuablc rt:cord or the ,ofi:-
p.lrt .matomy of man) kind, of org.llll\111\
bir.mlllll\ .tppend.tg;c' do\\ n tht· body. I adh>IIISI<l h.LS rhe \cdtmt·nt~ bdong to the Fr.utt.i' Ctcck <;h1ll.
tumeJ out to h,t\'C .1 ~urpn,mgl) 'uml.u body under depos1tcd m a generall)- ddtJit t•m·uunmcnr
tht· ldrgt• cJr.1pacc :md '' prob.tbly rd.l£ed. The~e betwecn .t northerly coal ~wamp and the 'l'J , .. h,, h
two gt·ucra could be \Ur\ivor; of tht• U,mC'fhl stock, lay to thc ~outh west. Two mam Hm.1l .1nd f.~u11.1l
pt'T\JMJng ro thc Dt·voJu.m .md known only from groupm~-ors t..m be ddincd. the non-manm•
tlu.'SI.' t·:-.u:pt ion a! fJunas, though ~nmc .1uthoritics Braidwood .md the marine Essex .~~~cmbbgcs. All
r~:g;ard tht·lr chJractcr~ ,t\ sympk,iomorphk. the fos,il~ li·om thcse Jrc prcscrvcd in iromwnc wu-
OdtL'r ck·ment' 111 the 6un.1 lilt lude e.1rly rcpre- crctJons. fhe non-marine (brat khh to lrt•shw ltcr)
~cntJtt vc' of ~cvcral 1-,'l'OliP'· t.g. thl.' floattng l 'del/a- Braid\\iOOd .1sscmblagc. wluch Js dummall.:d h)
like h-. drozu,111 Plcct<ldlst 11~. Amung the moUu~t' cxqul\ttdy pre,erved land plant<. (3511 \pctie,) .md
thcr~· ,tr th1n-~hdltd ctenodum bi\ .11\.·c,, g.t~rropods
tn~crts ( 140 spcncs), was eVIdent!) Jt•pu~ited m a
with \OIL p.1n' pre,cnt. JIJ\Jllt.llHlld\ .md bJctririd~ swamp)' lowbnd region clo~t· to tht• ,hon:. I hne
(CllJptt'r H) wl11ch give mlll h int"lmn.tnon .1bour tht: .1rc .tho 'corp1om. 'piders, nullipcdl'' .md Lt'n-
e.trl) l'\ olut1on of tlus group . .llld ~tr.ughr-,helled ttpl·dt·'· dcnved from thL' \horc\V.lrd rt'Jl he, ol th~
ceph.1lopod\. ~ume of rhe~e were n tuLochlcar, with co.tl forc~t. SnlJll ponds and th.ullleh were the
tilt· \ltdl uuhtde tlte bud\. wht'rl'.l' udteT\ were homt· of mtr.tcodes, and of .-.m.11l ~hnmp-hke tru\-
endocOt h le;tr wtth the outmh: t•f the ~lll:ll invcstcd taccam, tl ~h .md :~mphibians. Tlw Xtpho,urau
with \Oft tis~ue TentJcles .1nd lltht•r cXtl·mal organs Euprovps. nfi:cn found with tht• mort• tcrrc~m,tl el~-
Jppt.tr 111 some r:tdwgrJph\, lK>th uf the orthocone 111l'nto.,, may h.wc· ~pent some of Its hfc ~ub.1enall). Figvfl
l.of,,,!J,utntt•s J.nd in c..;,.,,,ll//(',1, .mJ ctnbryoruc (,., 2);
The conrr:t.-.nng E~scx faunJ mh.1b1t~·d rhe manu~ (, I 2
n:ph.tlopods luve :tho :tppt:Jrcd um xpectedly on f.lcic' <lf the delt.1 front. lt ~~ astotmhmgly dJvef'~. Povci1
the phorogr:tplllc pbte'>. Culcolth llJve aho been Jnd 111 m.my CJ\CS the soft p.tns ot the .1ninl.ll~ are Thom1
found tn rhe Hun~ruck~chtd~r. f>f!lt,,,,,/af<'a'rtls ha., prc,en cd .1~ C:".lrbon film.-.. ThL' verrehr.ue record
\Olllt' ii. .mtrc' rcnlllmcent of orthucunc ccphalo-
ft.'Vl'.l)\ Jll.ll1)' tl~h \pCCiCS ,1\ well ,1<. h.uchJing COeJl-
pods, hm pm~es~cs J ro~tmm, wht·reJs two other C.lllth\ w1th 1 he yolk-s:tc st1ll prt''l'rvcd, nne of .he
C\\l'S
gcm't\1 'how nurked ~uml.mttes to Recent reuthids t'arlicst known Ltmprcy~ .•111d other J.1wle~~ h~hc'. boda
•md would ,n·m to be thtlr e.trlll'Sl known rcpre- Ion~ .
Tltc invcrtd1ratc t':tunJ 1s domm,ltl·d hy l.trgl' lnl\-
wm:uive\. l11 'ome bed~ ~penm~ m t1f thL' cnignlJtlc taccam (cg. Be/Me/son). otht•r .trthropod~ (t.g.
,1 p.til
cnnit.tl-,ht·lkd tcnt:tculitJth .lrt· .1bund.1nt. On X-ray Euprot>ps. Kot11.Wrxes) and O} l'plt:nm.tl polych.wte smndedmtt) nt < 1n11vorou' llHO I
tt nd' m ~upport the Vlt"W tlf \Ollll' workers that l.ttcr.tl
h.1bit and complete with bn,tks ,111dj.1\\.,, 1 here .1re
tht'\t" \cpt.uc ,hdh wtth their l,m~ body chambers .1ho jt:ll} ti\h, hvdrmds (e.g. ,U.J ::,>Itp/rt~), cluctClg-
hnllu1
"n~· .m ~·xnnrt group of ccph.llnpo~k Oll\h
nJth' (c .g. ]J,wr~JclWIIIIII) , ncmcrt1nc wonm (e.g
s~~'1lll
Ardmymplcctt's). holothun.mo.,, b.IOl.tdt·, \\ uh rhni-
prob,t
nou~ pl.ltt'\ .md ccphJlopod' with .lrm;, .tnd hook\
markt
attachl·d. Among other tu~'1h thl'rl .trt• \penmem or
thl' ~·niglllatlt" Trtllimvlls/111111 (Tully monsrm), Trt/
whil h
munc.:d ;Jitl'r their discOVL'rt'r.•tnd unique to thr
probw
Mazon Creek fauna 423
(b) Kottixerx es (cl Fossundecima
(d) Tullimon strum
lhl Paucijac ulum
(g) Mazoh ydra

(f) Archisym p/ectes
Figure I 2. 9 Some s,oh-bodied elements ol the Mozon Creek fauna (Corb.). Illinois:
(a) Belotelson, a molocostrocon crustacean
(x 2); (b) Koltixerxes, a euthycorcinoid arthropod (>< 3); (c) Fossundec
imo, a polychoete worm, with jaws (x 1); (d) Tullimonslrum
(x 1.25); (e) Lotzelio, a centipede (x 1), (f) Archisymplectes, a nemertine
worm; (g) Mozohydro, a hydrozoon (x 1 ), (h) choelc>gnoth
Poucijocu/um (- 3); (j) Leviseltius. a polychoele worm ( • 1). [(a). (b), (f), (h)
Redrawn From Schrom, 1971, 1973; (c). (j). redrawn from
Thomps,on, 1977; others redrawn !Tom Foster, Mogel and other authors
in N itecki, 1979.)
Essex t:lUnJ. l"u//,,t,ll.,tntllt (F1g 12. 9d) w." a soft- most authonti cs have cons1dered '/itlliiJil)/l lfllll/1 tl) he a
bodied, bil.ltt'r,lUy <;ynunt:trit:al anin1Jl, so111c X nn member of an entirely exttnct phylum. f·mter ( 11>79),
long. lt\ he.1d taper\ to an dongaw d probme~s w1th how~ver, poinc. out that TullitlllliiSintm share~ many
a pair of pinn•r-l ike JoiW' ,lt the end, the1r tnner [~et· fcawres in conuuon with living hetcropo ds, .1 1-.TJ'oup
amu:d with mmute \{\let- Wht·re the head gr.tdl'\ of actively swimmi ng g<l\trOpod~ wluch h,lVe l,lrgel)
mto till trunk there; '' ,, tranwer ,e bar projectm g lmt the sht·U. These include overall shape, prohmet s
laterally from the hndy, t·.tth end tt·rrnm.Hing in .1 f(>rlll, raduJ,\r teeth and pos~ibJy also the C} C\. fhese
hoUO\\. globular b.lr-orga n: thew llJvc been \'Jn- corre<ipondences arc not, however , ex.Kt, and most
ously mterprn ed a' t'ye~ or \tab•hzt'l"'i. The trunk '" p.1Licontologists Mill hold the v tew tlw the btZ:ItTe
~cgmented, often <;howlllg ,l metban unpre~sion, Tu/limomtn1111 is a member of a long--exuntt phylum.
probably thl' b'llt At tht• re.ar a 'Patuht~ t.ul t~ The degree of nuxing of tlw Esse' and
marked with paired ltns. llraidwo od clemem s vanes: mmetim es unly a fnv of
1itllmwm tnllll wa' prob.lbl} a pdag~c c.lmivorc the more typilal Br.1idwood clcmt·nt s are t<nmd
wluch t,IUght 11:\ prc•y w1th the pmbmn s. ~mce the With mt·mber s orthe Essex fauna; elsc:where, more
probO\JC S and th~: lr:JJtW<'f" t' bar appt\lr to bt• uniqtJe, truly mixed faun.JS n"lay have re~ulted from pl'riodac
swmt ~urges v. hidt tllO\'ed the marine animah over time. 1\ a primary t.l,J... for p.al.aeontology 111 the ye.1~
the delta, suamJing th~..·m 011 ih mr6cc. Increased that lie ahead.
Jr
ram dunng such ,1 \torm 'urge is thought to have
T
swept do\\ 11 m,h\C\ ut wdtnH.'Ilt to bury thl' animals
lil
almmr tmml'dlatdy. Thl' c.:oncrcuom mmr have
fanned round tlw 'PI..'t'imem '-'l'ry rapidly after bur-
c
T
ial, and rhe prl'\l'f\ .ation it~df j, r.tthcr unmual, tor
Cl
wnhm the \H.h:ritt> nlllut:tium caluum larbonacc is
In Ba\ana there i\ a n:mark.1bh: and ""ide!>predd w
nonnaJiv alN:nt "hi le dmut j, unalt~:n:d, .md the
limestont' ut Upper JurJS\Il (f1thunun) age, for- (F
soft par;., .tre prcwn·ed .1~ c.:<Jrbon tiJtm. Th~: nodule~
merly quarried as lirhograplm 'tone. lt Th vel)' tine er
nught hav~: tonnt>d ati:t>r .1 ~uddt>n infltL" of ~tlr and
textured, and Liglu h'Tt'} to butr coloun·d. and
fresh water uno the .arc.t .1ltcr hc.t\'\ rams or due ro
:1lthough tl.1~\ll~ .tre r.tther uncommon thereiu, they m
channel dtver~IOn r he ,tbund.llll e. of Fe'+ derived
are excepnonally well p1e~ened .1ncl very di\er;e. d
from tltt> co.tl sw.unp~ tugl.'ther '' tth th~: raptd bunal of
Some ~00 speue\ ot .tn11nab and pl.tnt~ have bet:n
may h we provl·d an tde:~l environment tor the th.
recorded from lamutar bcdd1ng planes. and nearly alJ
siderite concrctlllll tot m,ttion.
the antnu l foss1ls arc nekttl marine invertebr.tte~ and w;
The Ma7on Crcd, Cu111.ts are of unusual compo-
vertebrates. The vcrtcbr:1rc fauna includes many fish be
sltlon, tor the common Carbonilerous m.uine fos<;ils
species, m:~ny aqu.ltlc (and ,a few terre~trial) reptiles, lat
such ,\s br.tclllopod' .m.· .tb~enl, even in tbe manne
prerodacryls :tnd the early bird -ltrlwrtlpleryx of
Essex f.1c1e~. and cmtold~. g.tstropods ;llld bivalves
(1:3.mhel. 197X. I t)7<.J; B.1rthel rt ,1/, I <.)l)O). tic
are r.tn·. The great hull,. nftht> fos,1ls ,1re, indeed. of
Ba
uncommon type
thl
In the 6unal ·"'cmblagl' seme, the Es~ex taun.l is
So
unusual. lt is not. however, umqul', except m the
sea
abundance and dn·l•r,it: oi fmo;11\ found therein
Sp<
Sim1l.tr C'trhollltcrom m·ar\hore faunas e:x.ist else-
de;
where. 111 Amcnc.1, l:uropt• and other place~. mhab-
no
mng shallow-nurim· l'nvtronmcnts ,tnd nearshore
COl
lagoom between '"' tmpy Im\ l.mdo; .1nd rhe sea.
ba<
Simtla1 bionr ·'"oo.uinm .tn· J..:nown elsewhere m
deJ
Tlhn01s (Langt•rl .md Rid1.mhon, llJ6.,), !Tom the
COl
Bear Gukh \t'l(\ll'IKt of Monr m a (\V 11liams. 19SJ),
!Tom Monrre.lll-k,-Mme,, l ranee (Heyler .tnd
Poplin. 11JHH) and from tht' C...trhomterous of
Scotland (">chram, I'J79.a, HnR-.l<i and CIJrkson,
1983. I'J85) wh1th tl·mt~ to .m ecological continu-
ity ovt·r ,\ Wldl· .arc.t dunng the Carboniferous.
Indeed a \imil.1r b1ot1l' .1~son.mnn can be rccogn11ed
as far back ,,., tht• Silurun 111 .111 equivalent ncarshore
habitar.
Such f.n111.1~ ,1\ tht''>l' .Ire tln\\ bl'J:,'1tming to be
undcrstoud, not jll\l dl'~criptlvely and taxonomi-
c.tlly, but Clolog~l.tll~. ·" h10tic .md trophic associa-
tiom cxrendmg thwugh \ptte .md time. In the
word, of Sthr.un (I'J79b), 'Wt c.m no longer be
content wHh trl',tting the M.1zon Cn:ck biora~ as
baroyut• bnc-.t-br:tt: tht') m 111 tal t pan of a com-
bination w unlol k thl· \l'l'rl'h of ( .1rboniferous his-
Figure 12 10 Cycleryon propinquus, a Jurossic crab from !he Figu
tory'. The undeN.mding 11f 'udt Ctun.\\, \\ hich givl' shOllow·woter Solnholen limestone, Germany ( 0.5) The only
and
'uch a detailed p4..'f\J1l'l tiH· on thl· \\ hnk biot.t of the living eryonids ore deep·woter species
ing 1
Solnhofen lithog raphic limestone, Bavaria 425
The mn:rr ebr.lle t.wna •~ parncul.lrl> nch m Smcc the c..linutc wa' very hot the lagoon.1l water
arthropods. \ome 70 spL•dcs bclon~mg to ~5 gent:ra.
~omerime~ becam e hyper.~abnt:< throug h evapo ra-
They were numl) swtmm cr;, ~uch .ts rhe ~hrimp
tiOn. The sea Aoor may h,we oftt'n been ~tagn.mt,
likc:: Acl!er, but there is al~o J larger m.1cntr.1n,
.md both these factors 'eem tO have been respom1blc
Cydrryon, .ltld ~ome ~mall hmuhnc~ .tl~o ocntr. tor tht· frequc nr ,1bsencc of epi- .md intaun.1 [thoug
!"here arc al~o m.llly ~pccie' of Aymg insect~. e~pc h
Sa((!lW III<l (q.v.) ma) luve been a
Cially dragonA1e~. broug ht in dunng ,c.lson:~l ram- hcutlu c oppor -
tunbt] , and also tor the remar kable paucit y of tr,lcks
'' .1ter Aoods. Other •m·crtL·br:ltcs imlud c jdlyfi 'h .md trail~. There arc indeed a tcw tr.l1ls, ~hort ones
{TVti:::ll.ltomitc~). aphal opod , and the ,1bund
.mc made by the dl.'tap od i\tlrmchi111s aud longe r trails
cnnoid Sacctl(tl/11<1
resulti ng from the movcmen~ of hmulmcs. Many
TIH!rc ha' been much disrus~1nn of the enviro
n- ~pcc 11ncm of 1\lcsolilllllitls rmlclli are preser
ment ol dcpmi rion of the Solnh ofl'n Limcs tonl'. le 1s ved at tht·
end of the1r tra1k Mode m l..mru/11~ c; highl> tolera nt
dt•arly a mann c scd1m ent, a~ ,hown by the presen ce
of ch.mg e 111 salmiry, tempe rature and oxyge n ten-
of cocco lidl,, ceplu lopod s anJ crinoid~ lr c; pos~1ble \ton, and the same wa' proba bly also true ofJur:.s~JC
tlut dtc bme-m ud tonrun g the hmest onc w.1s l~tnulines. ~o that at Jc:ast ~orne indivi duah
w.tshed into the l.tgoo n by stonm , but 1t may h.we could tol-
erate to some extent the locally tluctu .tring and
been at least parn.tlly ,Ill aJ~ral or chemiC<ll precip i-
often 1nimical condi tions of the Solnh ofen ~ea fioor.
t.ltc. Some author s h.lvc invok ed pcriod tc ~trandmg And althou gh these \amc cond1 tions prove d too
of the fossil~ on mud lbts to cxpl.1111 the1r prcscr va-
much for ~omc indrvi duab. the circumstJnce~ of
nnn. bur a more recent mtcrp rcwtJo n .tdvam.ed by
thetr demi' e 140 M.1 ago, record ed m dL·t.lil 111 the
Bilrthcl (1 Y7lJ) ,H'J:,'1.1C~ for a contmuo11s St.'J cover rock~. provid e :1 cleare r tlun nonua l pe~pe
throug hout Jcpmt tlon Acc..onlmg ro B.1rrh d the crive of
the hfe of that time.
Solnhofcn Lunes tone was appare ntly tkpos ited on J All these excl'p tionaJ £1un.1~ are arypicJJ onl} in
\C.I ftoor of 1rre~•ular rdicf ongin. illy
consi, nng of thetr preservatHm. lt is thi~ alone wh1ch di,tin -
~pongc-algal reef~ and motmd~. Maxu num
water bruishcs them trom othe~. What j, ~o unpor unt
depth may have been from 30 to nO m. "To the ahout them is that they provid e ne:lr-c omple te pic-
uorth there were b,1rricr isl,tnds, ~md to the \Outh a tures of the divers ity of .mima b living at panitu lar
LOr<~l-hydrozoan barrit.•r reef. wluch cut
off the time~, thoug h rhey were forme d under
b.tck- n:cfla goona l ,1re.1 in \Vh1rh the hmc\t one was dtfrer cm
lOndit ions. There secnts . howev er. to be no genera
dc:po~1tcd as a carbon atl' mud in a ~y,tcm l
ot intcr- patter n for the preser vation of soli: p:1m, and there is
lOnnccrcd b.1sim frin~,..;ng the deep 'pon~c-rccls. no dc<lr unde~tanding of how decJ} is mhtbl ted.
Figvre 12. 11 Depositional porometers for early diagene sis

of soft ports of fossils: (a) pyritization, requiring o very high
orld low organic content; (b) phosphotizohon, requiring o burial rote
low burial rote and a high organic content; (c) within
1ng o very high rote of burial and high organic content (Redraw carbon ates, requir·
n from Alii son, 1988.)
426 Exceptional faunos; ichnology
Much more hmgl'Ol'hcmical re~card1 (e.g. Fig. \tudymg trace fu,sil~ .md lwnre d•tferelll 'Y'tum ftf
12.1 I) need~ to lK· utHk-rr.!ken to rL'\olvc m.tny cur- clas\lfymg them (S t mp~t)l1, 1976). Thn:t· of thc:o;e m
r~.·nt q nc.:\tion~. parllrul.u are important: morphological :md prcser-
The further ~tudy of Fm~il-1 a~cm:iLtc:n would vau o nal , behaviour.tl, .tnd phylogenctic.
\C.:elll to be one of tlw 1111p0rtant t.l\1,., tor paJaeon-
toiOg) 10 the lillurl'. Morphological and preservational classification
fratc fosstls arc all gwcn 'tc)nn-genenc n.unn'. that
1s, the n:unes an:- mcd on I> tor the d"tinnwn of var-
lOll~ rype~ and do not Jtlcmpt to 1dentil) or 'uggcsr
thl..'lr nnkl..'r. A simplifil..'d system of t-,'T~Hlping them
khnology is the ~tudy of the bc.:hJviour of o nce-liv- with lllterpretation is giwn here.
Ing Jnimals b) cx.u11in.ltllln ofth1.· tr.Kk\, tr.HI~. bor-
Ing' .md markm~"' that the) nude wlll·n ahvc. The~e tr.lcks or tro~ib on a bcddmg plane origm.tlly nude
.1r~ called trace fossils or ichnofossils They arr upon the sediment- v.ater interface. c.~. Cnt::hJtM
usuall) preserYed onh .u rh~.- mt~.-·rface of two rypes (rnlobltC trails), \ 1acllt'S, Pfi)'C<lS1plum, c.l.llll<lfQ]'/It·
ot ~t:dnnent, tor 111\l,lllll ''"here th1.· marktngs made (worm tr-atls). Gywt!l(lJ tc (possible g.t,tropod rr.ul):
nn J mud surfan: by ,1 uawling trilobill' were filled mdially symmetrical horizontal m.trkin~,..,, e.g.
111 by ~ine s.111d "' hid1 bter lurdcnc.:d. khnology is 1~trria(itcs (re~ting mark~ of startl~h):
by no means a relent Je, dopnu:nt 111 geological wnnel~ <lnd ~lufts within the sedlllll'llt, e.g.
~c1cnn: - mdeed Jt ~.-.tn be tr.tted bJLk tn the 1820~ "'kctlirlws (verncal womt rube,), Cht•m/,to (br.utch-
but ll j, onl~ \\tthlll tl11.' l."t two dctadc' dur it h," lllg galleries probJbl-y made by a probin~ wunnJ:
~.ome to be a re.1lh p1.m ertitl tool tor undcf'r.mding tran~~ with J spreitc (a web-like \tfll( ltln:. usu-
cerr.tm past ~edm1enur) ~:nviromJl\.'llh .md one of ,,Jiy with a sene~ of roncenrric marku1~") J\ otkn
our m;lJOr due' to th1. behaviou1 of thl• ,1ninuls rh.n fimn d joining t\\'O br,ll1chcs of a U-tube .md
lived within thcw t'J WJroll lll t:nt~ . Tlw different rt·pre~enring fonner positions of the tube within
kind~ of traces made 111 'hJllo\\ -wJt~r \cdiment' the o;cdiment. e.g. Rhi::occmtllilllll (hon;wnul U-
have 1-lcen fi.1ll) dtll\llllented in the v.l\t mudflats of tubcs), Diplvtratcrittll (wrtical U-tube,), Ztl<tp/1)'~<'-'
rhe North ea b~ Gt•nn:~n ,-..orkcf' ·" \Ummarized (mchncd spiral~) - ,tll of which could h tve bt'l'fl
b) c;;t-h1ter (147 ~) ..md dm knowblj.!;t' h.1~ been nude by different kmds of anunah:
.lpph~.·d to the gn)log1~.1l record. \tnte Vt'l) good pouch-, haped markmg-., e.g. P<'lcqp(lditl1111t• (l)J\.llve
(f)
sumn1.1nes of tr,tlt' to,sil work ex1st ehewhcrc burrows). Rusaphyms (rnlob1te resrmg tral.t'').
(Cnnw~ and ll.1rpcr. 1970; llantz,rhd, 1975; Frcy. other~. e.g. f>a/(lcadirl y<•ll (net-likt· structurl..' of
197(1; Donov,m. l 1 >t>l: Bromley. llJlJ(>). only a tcw unccrt.1in orib>in)
potnt' will he m.Jdt hat' ro ~hrl\\ snmt•thm~ of the
'l'ope aud methods of tchnolog}. Although rhi' de~rnptivc rl.lssitirauon h.t'i 1r' U\t'-
In p.1mcubr I \\ould reter smdum tu Uromley's fu lnc\\, the ch,tractc" ,eJccccd tor dcscnpuon mmt
degant and hwlo~l.tll> b.1~ed rreJtm~.:nt In this arc be arbitrary and therefore f:t r rrom obJCCtlV(' A d;l\-
to be found lll.lll} illll\trared exampks of the life sJliration based on prc,crvattonal fe,lture~ .1 lonc 1s
h.tbit"> and tr.tn·s nC modern burrow111A .lllimals, and equall y hard to apply.
,111 .1pplication of hinlo~:.ric.ll pnnripk' to trace toSSJl (h) l
studll'S. Behavioural classi~cation

Sei!Jdter ( l 964) dc,igncd a very u'ditl \dtcmt
bJ\Cd upon th e behaVIOUr of the orgJnl\JH th.u nude
Class ification of trace fossils the tr.lle~.
In moditi~.·d timn thi~ d,l\\thc.ltlnn now FifJUI
h.1~ ,Hleast six catcgoneo;, but \in cc th~ v overiJp Ill m me
star~
rr.tCe fos~i)s :1re aJI St'dlllll'llt,tr}' \[rt1LltlrC1> made by som e c:xtent this s1. hc1nc i\ ,d,o not .t p~·rtcct o;ysrem. Rhizc
the .lCtivities of onu.: hvlllg .mim.tls, 111.\ln ly 111\'ertc- though it i~ useful. Th~· tat~.·go n c\ an: ,t'i follows I, nor
brJtes (ltgs 1.2.12 1.2. U). inrlia
However. ~lllul.tr tr.let."> tan be nude by tJUite d1t:. move
Cr~w.lmg traLl~ (R epu;hni,l): thco;e ML' rr:tl k' uf
ti.·rt•nt kmds of org.llll\111\. Thcr~.· arc \",tnou~ ways of
Polec
mov1ng ~urtace dwc:llc~. e.g. Cm:::~<lll•l . 1962
lchnology 427
(b) Nereites
(c) Chondrites
(a) Cruziana
(d) Cosmoraphe
(e) Asteriacites
",.;: .
(g) Gyrochorte
(f) Phycosiphon
(J) Diplocratenon
I 11 Ill V VI VII
Figure 12. 12 Trace fossils (o) Crvziono, mode by ploughing trilobites (x 1}; (b) Nereifes, o surface trail (>< 0.3}; (c) Chom:lrifes,
mines within sediment, mode by o probing worm (>< 0.5); (d) Cosmoraphe, surface trail (x 0.5}; (e) Asterracites, resting troce of
slomsh (x 1); (f) Phycostphon, surface marking, the originating worm marked in block (x 5); (g) Gyrochorte, surface trail (v I); (h)
Rhizocorollium, oblique loterol view, os if through transparent sediment (>< 0. 1); (j} Dipfocroterion, in vertical view From left to right:
I, normal U-tube; 11, U·tube descending os secltment is removed; Ill, U-tube ascending os sediment is rapidly deposited, IV-VII, U-tubes
101tioted at various times and showing movement 1n relation to erosion or depos1tion of sediments on top; arrows show directions of
movement before ~nol plone erosion and deposition of overlying sediment. !Redrawn chieRy From Treatise of Invertebrate
Paleontology; (a) based on Fill1on and Pieke rill, Pofoeontographico Canodiono, 1990, (f) From Sromley, 1996; (j) from Gold ring,
1962.)
(a) Dictyodors (b) Zoophycos (c) Skolithos
(d) Palseodtctyon (e) Zoophycos (0 Pelecypodichnus
Figure 12 13 (a) Oictyoc/oro, reconstructed (x 1), (b), (e) loophycos, in surface v1ew and 1n section, respedJVely ( x 0.1 ). (c)
Slcolithos in vertical section (x 0.2) (d) Poloeoclictyon in surfoce view(>< 0 5); (f) PelecypocJichnus, surface view ( 0 5)
Ge
Rc,tlng tr.ltl'~ (C ub1d1111.l): the'c are exemplified sprettes, wluch ~how th,lt the U-burrow has bt·<·n of.
by Rus,,pftyws, lsttn·.,utn. progrcsstvdy deepened. There are also vanous lun
Gra7tng nr 'urt:1n' f(:cdn tr.tee\ (P,l_\lichnia): sur- kind~ of ramal ~tniClure~. mclurung the many- ph)
face r(:cdcf"> m,th Lhar.u:tcri'>tH.: p.ltterned mark- branched Clw11drite>. whlth ""''~ probably made b) of•
mgs o n tlw \L'dunent; the ~>.tmc \Urt of pattem~ a wonn exploitmg a nch depostt succcss1vrly. 111.11
m.1y be SLL'l1 lll.ltk by Rt•ccnt mails brrazing on again .watding any are,t or
previously ingested ~ed IllS(
algal on '>tom'' 1n rod, pook Nereites, for imcnt. An intcrcsung Stlunan tr.:tce foNI, R11•
inmncc, h .1 tr,\l~ .1ppc~mng ,1s ~~ series of parallel Dtctyvdorn, tonm a sinuou~ nbbon w1th a fbttened ind.
hncs with latcrallobt·~. jmned by tight loops, each T -shaped lower edge. m eandeting through the ca.ti
line being l'l]Utdlst,lnl tn.)m its neighbours. T hts is rock nom1al to the bedding (hg. 12.13). lt must .md
madL' by ~~ stu{KL' t(:L·dcr econom icaJiy exploiring have been con~tntcted by .m :tnmulmoving below
<l fond supply. Slt(:h a~ baLtt•ria. upon the sediment the ~urfacc as it fed, yet mamt.1inu1g contact wtth No
o,urfacc. coVL'rtng .1~ rnud1 ground as pos~ible but the water by meam of ,1 vert1cal tt1be or pipe, and f.'lCt
w1th thl' minum1111 of dlort .111d llever recro~sing it 1s the passage of the latter through the sedmwnt ;my
it\ own trail or f(:cdm~ along previously grazed which created what t\ now pre\crved a' the vertical oftl
,urfaLL '· Pit yn)~iphtl/1 pertonm the same kind of ribbon; the \vonn· made thl' T-shaped ba~c
huKtton m ,1 ditl~nmt \\.ay (llemon and Trcwm, 191-10).
F~:tdmg trJu.•\ (fodtnlt hnl.l): these :tre excavations Dwelling \truCtures (Domtdlllia): thc~e .ue p~r
l..
made by dcpo'll l~·edc-1'). i.e .•munals liVing on the manenr burrow\ .md bonnb" of 'mpenston feed-
surface but Jtmcl~ muung wnlun 1t for thetr tood. ers. i.e. anm1ah, ltvm~ \Vlthin the !>edimenr but
They mdude \tllll\.' U-'>haped tube' wuh intem.ll srrammg otT parttcle' !Tom the water .1bow. The\
lchnology 429
ma} be subcy lindri nl mb~:s (e.g. <;J.:t,frrhth). bul
fimnd 111 pla..:e. lfrhe enclos mg sednn cnt had move d
there .1rc ,tlso some U-~haped burrow s, thoug h
thq would have been destroyed They often OlCUr
nom1ally '''t£ho ur 'pn.He s. 111 sedun em.uy su1tes wher~: then~ Jre
E~cap1.· \tntCtt !re' (Fugtt ·hnia): vanou s kinds ofvn no body fm~1ls
- prc~ent .md espel tally 111 clastic 'equc:n
m:al n1be-. 'ugg~:~t rh.1t .tmmaJ, moved upw.u<h lO ces. Rathe r
than bemg de~troyed by d1agcne~ts they .m: com-
r~capc from beit1~ bum•d by .tn influx nf c;edun
cut. mon!~ unpro ved by ir. ~mce htholo gJCal contra~t
or dO\vnwards when the <;eduncnt .thovc wa~
bnwet •n the fo,\tl :ll1d the ntclos mg ~edmtcnt may
crod~:tl. The origlllal tube~ are U\Ually rherefo
r1.· thereb y bt: 1.'11h:mced. furthermore: thLy tend to
modified Donu duua. Diplorllllt'Mil. !i)r exam pk, 1s be
rc,tnc ted to ,1 narrow f.1cies range. A~ glllde~ to the
.1 vertic.tl U-tub c found in high-~:ncrgy
cm·lro n- nature of the sedim entary enviro nmen t m lCrt.tm
ments, wnh ~pn~ltt• \true tun:s 1:1ther above or t} pes of segue nce they hav~.: prove d mv;tlu
bdo"' it 01 both lt j, uttcrp rcted a' the penna ncnl ablc.
If trace tchsih are presen t in a cla~tlc seque nce
Jwdlm g burro" " of an .mim.tl th.n lived bdow the then the sedim ent must h.1ve hL·en well aerate d.
~cdiment. When ,eJmw nt ''as n:mov rd b..,
rapid C'onvcr<~dy, a seque nce lackm g m trJlt'S may \\ell
ero:uon the amma l buJT\l wed down more deeply 111
h.wc been anoxic . Thus black shales w1th abund .mt
order to n·mai u lOl1c1.·.Ued at lb habttu al depth.
trace'\. even tf deep- water m origm , canno t poss1h
lc.wing a ~pn:lle .tbnvc . Raptd ~cdinwntanon ly
have been .mox1c. Grapt olitic shales and their
cauc;ed the orgaJll\111 to mm e upwar d'. produ cing Me~o70lC equiv.1lems, wh1ch c:ont:un only
a ~prcltl' below of ~onwwhilt d1tf~:rt·nt morph ology thm-
shelled plank lonic bivalv es, ,tre gener,1lly lal king in
'ince 1t wa<; no long...·r lmk.111~ parallel Jmts. Such a
tr.tce tossils, which sugge st' a cu:o.mic ongin .
\trucn1rc gwes 111ionn.1tion about tht• cJwtro n-
Rates of \cdim entati on can often be inferr ed
ment. ~uggl·,nng ven rap1d cro~1on .md depo, l-
from thl" relativ e .1bund ance of trace tossils, as With
nun, ,md the related up-:tn d-dow n move ment of
Diplocmterio11, and the presen Le of defim tc boring ' 111
the orgaru<;m that m.ltk ir led Goldn ng ( 19(}2) to
'lurdg round ,' shows clearly that the <;ed1mem 111\1\t
give the orga.tliMil the approp1 iarcly de~cnptive h~ve been
indura ted when the boril1 g\ w<.:re
'pecifi c name D }'•'}'•' tonn~:d. a~
may somet m1es be ~..·otlfimt1.·d by
cncrm ters upon 1ts surtac e. lnevit:~bly a tempo rary
Phylogenetic classification break Ill dcpos itton must be rccog mzed to have
Generally the1c are few llldiL.Itlom as w the identi ty allowe
d such indura non. Mino r varianom m tauc"
of a parncu l:n tr:J.ce nuJ..a .•md otren tV1.'n it' phy-
can often be mfcrrc d from the presen ct' of partiw
lum can hardly be l'\tabl i,hcd \liKe ,Hlim.lls of many tr.1cc l.u
fossil ,1\sem blage, , even if the nature of their
ph\ la <.:an nuke tlw \.IITil' kmd of tr,lCe,, In the ca'e
m.1ke n ts not kn•wvn.
of ~omc .11thropod 1chnofo~~Jb, howev er, the trau:
The .maly 'i' of trace foss1l assemblages Js lcs' \1111-
:nakcr ts knmv n \\,th cert,un ty; tnlobJ tc,, for
pk than it .tppcar.., since: the pott:ntia1 for prescr v.l-
m\lJnce. have ,H:tu.dly been found w•tlun
tion ot 'fo~~il bch.wiour' depen d' upon the depth .1t
R.Jm•phyws burro w,. Ho\\'e vcr. ~incc this 1' rare \\ htch
the trace- maker hvcd wllhm the sedim1.·nt.
mtked there ~~ no point in tn mg to er1.'l't a cJ,ISSlfi- Most
living lllfaun al amma h arc vertica lly zoned .md
C'.&Uon based npon tht: 1dcntH) of the tr.lcc maker ,
inh.tb lt dll.f<:rcnt levels or tJc~ within the ~cdim~:nt
.md tonn-g eneric n.1m~:' h IVt' to be giv~.·n . (Fig. 12.14)
WHIH n t\ldt of these ticl"'' the-re may be dctlnc d
No me:Jns of dassif ication '' thcrct urc ~.·nrirely ~aci' il:hno
gmlds (Brom lq:, lY96), 1.e. group s of Kh-
IJctory or comp lcw, and the one Lhat '' tollo"
c:d w nmpc ues which behav e in a sinnla r nuwtc r, ,l!ld
any partic ular c.tse dcpl'lldc; rather upon the purpo se on·up
y l sumla r Jocau on w1Lhin the substrat1.. In the
of the \tudy. upper layer~ there may be highly mobik .mima ls
''hose hfe al!ivit tcs tot,tUy obhtc:r:ltc other prinu ry
~tructurcs. They may .tho sometime~ make
it d1fli-
Uses of ichno logy culr for the mhabl tJJlts of lower ner.. ro milinu m
contac t w1th the surf.1cc. While the upper byer..
Sedimentary environment of
\cdim cnt .m.: usu.tlly very mrrcd up (biotm batcd) ,
By contf. l)t wtth body fus~ih, trilce fos,lls .m: always
eh..• trace\ in the lowcr mo'it layer~ (~:.g.
77~cr/,'-1SIII!Hdcs) .1lunc.• .n~ thl' ones nw\t like!~ to be U\C

prcsc.'r\'l'd. Bmmky\ ram 'dttc.· tr:~n· lo~stls' apphes tiOI
to rhc.•sc: dwy lllJ\ .llsn be.· cnh.mu:d by dia~Cnl''l~. Th,
Thl' tf.lll' li.hstl ol'>'>lll\bl.t~c.· that 1'> ti.w;ilizc:d IS thu~ [JI)
nor ~ true.• retll'c. uon of the.• nnguul b10log]cal com-
mumtv Wh.lt \\ c sc.•c.• '' thus t.lphonom.ically 'tu-
tcn:d' .tnd .11! tlur rc.•m.tuts of the rich .md di\·erse
.1sscmbbgc.• of tlw uppc.•r laH.'f'i " \ll11pl) nuxed and
b10turb.1tl'd ,l'duncnt.
Thc.·re .trc.• .1J,o r.lthcr ~c.'lll'r.lltzc.·d trace foml IN.
Uur
.!\\l.'lllhltg~s \\ h1d1 gt\1.' tnti.,nn.~uon of a grosser
( J9t
4uall(\. l'spc.•u,Jll) pc.• rt.unm~ to dll· on~'lllJl depth of
arc
dq,osltlnn (~ul.tt·ha. 11ll>i). Smu: phy~K.ll protec-
tion 1s HI) unpc.,tlllll hl ,JJJllow-watcr dweller>,
partly Juc l1• turbuk-nu.: Jnd p.utly because the light
d11uws other .111 ituab to we.· thl'm. ,1 lugh proportion
of HlVertcbr.Hc~ arc.· blll roww~; m~pc:ns.ion feeders
(tiJtc.·n:f\) .111J the tLilCs .Ire lllJJnh Cubtchnia and
Oui11Jthnu. r arth~:r 1\\.IV Ill l)tllt'ter waters, dcpo~lt All"
fc~:dn~ ('wallown~) Jn' abumbnt, and hence T
Fodimt hm.t b~:c.omc.· 111~ t c.'.t<oinp;ly common, likew1-;e re
p~
Rcpll'hlll,l. In the.· lil..\htlc,~ ~;·nvimnment oi the deep
Butl
\Ca thc1c j, 1~:" n~:c.·d fot PH'tedmu. and becau'e of
D
the rich \ttpph ot \llrf. ttl'-<1\\ dhn~ bactena there are pi.
vcrv m.un f mhnllllllu .md t:\pl'c.: i.tlly Pasnchma. lhrtl
ThL" bua nuke c.t'IIIPkx 'P1r.1l ur meandering pat- s
ran' tor ')'tcnl.ltll ,tllY gl.tZint: from tood-rich sur- (..
Gee b) en.. On tlth bas1' therc '"ere erected tour Figure 12 14 Tiering of troce loS$ils, Creloceous example. (1) Bngl
main (\l'll.lt hl'll.lll) ll hm.1l,ll ie' ba,ed un bathymt:- _~;,!
bioturbotion from echinoids, goslropods, bivolves, elc , (2)
rry. the.: '>J.w/IIIIM, Cm::i.w.I. z,,,,P,Y"'~ and ,\'t'fcllc$ Thalassinoides (crustacean burrow system), (3) Taenidium (bur· w
t'Jctc,. Otlll'r trhnot:tllt'' h.t\'1.' been added bv differ- rowing worm), (A) Zoophycos; (5) Chondrites, lorge-bore and C on•
smoll·bore forms, respectively (Redrawn from Bromley, 1996) Sh
cm authon. \lit h ·'' dtl -~"'}'t'llht n:dbed 6cies. The
SJ.:olultM t:1-.i1.", uHtsl\l\ l,1r~dy ut vcmctl rubes in ph
Done
d~.:Jn, ~lullow-\\,llCt \,tmk the shillow-water
Ue
Crtt'::l<lll<l ichnut:ICil'\ (dq'll\Hc.·d .tbovl' storm wave-
Eiml.'
base.·) i, of bwrm"'· Jnhropod tratks and resting good zone fmsib. Yel ~ume -;hort-ranged ewes Su
tr.H.:C\; tilt. uttl."ttttl'di.ttl." L,,,,p/Iy(os Goes is highly arc useti1l in local srrali~raphy, .md 1110\t parti~.:ul.uly Coul
bioturh.ucd and "'·" nude pnuupally by sediment trace fmsils have been or immense value m under- l'l/r,
miners; ,111d th1.: dl.'c.'p-\\,llt r ~\'cintn t~1cics ;~ssociatcd standmg rlll' cxplt.hJOn ol litt- a1 Lhe PrecHnbri.m- (A
\\ ith turbid1H.~ h,t, ~r.11in~ tr.1il\ of tompkx form. Cambrian bound;~ry (<....haptt:r 3). and particubrl) ol ll,wfi
nr
Tht·~l' ll hlltll.lc.'ll'\ .Ire.· ll~l' in a brcnd ,wd gen- llll'tazoa n lift·, Ior many trace fossils can only br Fd
eral \1."11\0.:, hut 111dtv1du.tl tt.lt.c.\ Ilia\ occur out of m.1de by Jn .lJtimaJ \.VIlh .1 ~ut. 0Lht·r exJmples have Marn
J·o1
context, .111d th..: ,1\\c.lllhl.t~..:' thc.'nN.'h'l'S Jrc subject been given by Cmncs .md 1 Ltrper (l 1l71l).
Gu
lO t.lphtJIHlllllt lllt..:nn~.
Mo01
Fossil behaviour
H1/
Strahgrophy In a s~mc, all 1dmulogy rdatc.·~ to the study of Iima \ne
Cctll:ralh tl.ltc. lo"th .u~..· ol n ry htdl" ~tratigrapluc behaviOur The complex grazmg tratls d1Jt ha,·e La,
m~;. The \'c.'!')' dlat.lltc.•r; th.lt mJkc them 'o vaJuabll' alre,tdy been llll'nttoned t~'tit)· to a d~..·gree ofbchJ\- "I[('(
~:n\'ironmcur 1ll). t.e. Inn~ rJn~c. 'trid taocs con- IOUraJ complc:XJf\ in their ancic.·nt nuke" like thJt ol Ac
trol. c.'t1. ., Ul' Jl"l the uppu,Hc.' uf '' h.u '' neetkd for many pre~ent-tby oq~.mmm. Trace h)\\lls have been p.tp
Bibliography 431
U~l·d Jm:~.:tlv m .tu.tJy,mg the: hfc h.1b1t' ,md lo..:omo- Stmon.:rt.l, A.M. and C'onw.ty 'vtoms, S. (eds) ( 1'Jot I) /7re
oon of tn.lobitc:~ .111d 11thcr Jnhropods (l'h,tptcr 11). J:arl)' Er•nlulitllt of ,\Jcta::c~ot and tltr Sr.~tnr/1<'<111<1 c!{
'1 heir u~c: in intcrprenng ltfc habits and pN ~c:Jimcn l'rohlt•mtlfir Trt.\'cJ. (Procl'l'drng:; of .111 lnt<..•m.llion,tl
tary cnvirorullt'nt~ l .111 but mcrease n1 tlw tu cure. 'iyrnposnrm held at thl· UniVl'l'\tty ofCaml·nnu• .27 31
M mh I98'J). CambrrJ~c Umvc:NitY Prcs\, ( 'amhrrdgc.
21111 pp
\~ hntlll~'ttlll, H.B (I t.JXS) T7rc· Brtrl!css Slr.tl•• V .tk•
Umwrsrry Pre~~. 167 pp.
WhHlln~>ton, H.l3. .mJ (on way Mom\, \ ( 1'JH5)
[N. B. Smcc rctcrt•n<.:t'' ll> ori~mal Jc~u·iplHlll' of the F.\trtl<>rdiwrry h>.isif Hiotm. /1l('lt E,-,,f,~~~~,r/ ,,,,[
13urgc~s Shale t:mn.l .ne found in Conw.1y Morris til'cllrttitlllnry Si'(nijic<IIICt' Rlwa.l "iocil:'tv. London, I-1J~.
(19!-12), Whittin~ton ( l tJHS) .md GoulJ (II)IJD), they ( 19 p.lpl'l'S)
art• nor meluded hc:rt· .[
Individual papers and other references

(exceptional faunas)
(exceptional faunas) Alhson, P.A. (1988a) Plrosphamcd soft-hodll'd "luid~
frun1 thc: Jur.mic Oxford CLly. Lctlrclltl 21. 40' I 0
Alh~Qn, P A. .md lln&."'· D.E.G. (l·th) (1991) Alll\on, P. (IIJHHb) Kllii\I.'I"V.It-L.igcm:lrtcn: t:.IU\l' md
l.1phonomy: rdcJ,Ill~ the d.tu locked 111 the tm;;il d.tmhntion. Pai<lt'<•b•o/,~y 14. JJl_,g.
n:~ord. Plenum Prc~'· NI.'\\ YClrJ..: and l ondun Scveral Alll\on, P. (1':190) Decay procc,)c:'. m p,,[,,,·,•ltrcJ/,•t:r: ,,
papc:r.) .')yllllr<"5r.> (c:d<o D.l:..(, Brig~"' .md P.A. Cnn\thcr).
BJrth<.'l. K W. (I •PH) '>c•lllf~<•f(·u ,.,, Hl11k ;, die Bllckwcll, Oxford, pp. :!I'\ 6.
Ert~s:o.iutfllrc. On V1·r1Jg, 1 hun (E:-lt•llc:lll rolour B.mhd. K.W. (197<J) Solnhofcn Foml,ltton, 111 Tin
pl.uc:s) L:myd••!wtlicr <!flJa[e,mttllCJ,S:)' (ed~ R W. Paicbridt-;c .llld D .
llmhcl, KW., Swmburm. NI LM . .md Com\.1) Morri~. J.tblonsk•). l lutchimun. Oowdcn and Rtm,
S ( 1490) Solufw{fu , I Hll<i) 111 MnCJ.::orr /',r/,rwulclltJ.~?Y· \troud,burg, Penn.. pp 757 {•4
(,mtbndgc: 245 pp Dcrnc:r R A. ( 19R I) Autlugt·nrc nuner.1l l~1rm.1tion re,ulr-
Bn~. D.E.G., Erwm n H .md CuU1c:r. FJ. ( 1<J'J4) Tirf rng lrom orga.ruc nutter tll'l omplNtron in modem ,l•d-
JcH~i/s •!I rlrc Bw.:n.• ..,,,,,,,, ~nuthsonran lmnn1rion Pr~'· lllll'IH~ f',•rst>e/rri.fit'.flir ,\frll<hlh~lt' 59 . 117-.h
\V,l'\luUK{:Uil. Bcngtsun, S (1991) Oddb:tU, from the C.nubrr.m sr.1r1 tu
Conw.1y Morri~. ~.( (t•d.) (JQ82) .411<1.' t>(tlu Bu~~css !4t:l c:wu. Nnttrrr 35 1, IH~-'l.
S/r,rlr. Pa.lacontolo~tl d A~souJtion, Lllndon. (Folio of Bc:rg,tri)m, J. (1'J72) Morpholoro- uf fOssil arthmptHh .ts .1
photogr-Jphs and .: xpl.m.mon) ~mlt> to phylogenc:trc rclmomh1ps. 111 Irtl,,,,,,,d
Donovan, S.K. (c:d.) (1 ')'I 1) Tirt' Proii'«rs t•/ Ft>.•sr/iscltlttrt. J>lrylvl!tiiY (cd. A.P Cupt.1), V.1n !\lostr:md Rc:mhold.
lklh1vcn. Bo,ton M,,,,, ( 1.2 uphomonic p.1pl'l'<>) New Vorl.. pp. 1-56
Em~e1c, G and Sc1l.ldtc:r, A. (cJ,) (I'JH2) <.yrlic .uul Er•,•m 1-krg,trilm. J. (1980) Morplwlog) .md "''tem.tuc' uf t·.•rlv
StroJtrfri.1titm. ~pnn~er. lkrltn. .1rtluopods. . ~blr.mJ/rll~f! du i\',uun'''·'·'''"'' lr11/lrd;<
Guuld, S..J (1 t.JIJO) Wt>mln/ul Llli': tin Blll.l!t'SS Slr.tlr a11d tire I tl!'tll(~ull,l/ Hnmluux 23. 7 42
,''\lalrllc' c?[Histtll/'· Hutdumon Rldiu,, Londt11l, 347 pp. Bcrgstri)rn, J. (1990) H uns•ikk Slate, m /'t~hlt'llbio/,>.~y: ,,
(A mine of d.1t.1 .md phdmophy) Syllt/rc.>t~ (.:Js D.E G. Unggs and l'.A. CrowthlT),
I!Juff. B..tnd llaufl'. R B ( 191:-ll) D,r, Hcl/;:m,tdcflbrtclr. l11.11:kwell. O.'.ford. pp. "177 ·9
c h-llbark (Rcpro-Unllk) IJh pp. Hl'fb"'triirn J .111d Brd~~d. 11. ( JI)X4) Legs m thl· tnlobrte
.\!mill, D A (1 1111.\). lwsrJ., •!f thr ,..,,,,,, a111l Crato Rlrnr<'p.> from the Lower Dc\'OmJn Hull\rllll. \I.Hc:
f'()mr.tlrcm; Br.t ·tl I' Jl.tnmcologtral A"uo.mon: F1dd L.tfltlltl 17,67-72.
(;mdt' m Fo~<;ih No. 11. Ungg., D.E G. (I9H3) Aflmitil·' .md early nolunon of
Mnore, R.C. (cd) (1 115'1) Tre.tiiSI' cm iltr•crtcb•••tt· the Cru~tacea: the cVItknce of the Cambri.m to\sd~. 111
Pnlt·cmtolo.t:Y, P.m t.), /'ri/,•l•itcllllt>lphrt. Gl•ological CntsltJH'atr Issues /· ( ;,u,t,rrr<lll Ph ylo,I!~'~~Y, (cd. F. R..
Socu::ry of Amt'm., .tml Uruver.iry of K.ms.t~ Press, Schr.lln). Balkema, Rottt·nl.ull, pp. 1-:.!2.
L.m rl'lll t'. K.Ul Bngg,, D E.G. ( i9'J I) b.:tr.1urdmary fo\\tk .tlmmitlll
Nttetl.i, fo.\.H (cd.) ( 1'179) .\1<~.::•'" Crn J.: Ft>SSI/s. .\dwti.'t 79. Ull-H
A.c;.~demrc Press, Ne:\\ York. (Excdllnr lWC:r\'tC\\ - I() 13ngg-.. D.E G . .llld CIJrk,on. E.K.K (1'JX.l) 1he
p.lpc.-) C.,rbomtt:rous Granton ,hnmp-bcd, Edmhurgh . m
.md Orlr" .!~tltr•'Jl••rls (.:tb O.E.G. Bri!,tg; JnJ

Tri/,•bltt:,; Systl'llllltlcs 10, )27 4'1 (Very guod ovcn·tl'\\)
P.D. Lm.:), \p.:u.1l P 1pcr; 10 I' 1hconrology No. JO, Comv.tv Morm, S (l'lH6) Tht• rmnrnun•r.: ~tmeture nt
At'.ldl"nlll I'll , lnndon. lhl-77. (Cntst.ltl';m the MKklle C.unbn.m Phyllopnd lkd (llurg"'~' Shalt).
Lagt·N:iuc:) Plflll'tlllhl/c~~Y 29, 42.\-!>7.
Bng,L."'· D.J:.t •. 111d Cl.trl.,nn. t:.N.K. (I '>!\5) The Lower Com'a" Morm, \ (I 'J!\'J) l'ht· pcNstcnn~ of Burge,,
C.ubontfi:rous 'lmmp I C•lllh~.ms trom Gull.me. Eao;t -.halc-C)·pe l;.unl\ unpht·~unll' t(n tht· C\'l1lunnn ot
L,~thun. Sc<>tl.tnd fr,,m,uft,>ll> ·~f tlrt.' R,>)•al :),>a ay of dc~:p.:r \\ Jtl'r tJUII.I\ rr.lll>tltCil111S ,,, tin· Roy.tl Si"-1• '>' of
£,/.,hurl!h: E.mlr '" lho 76, 17J-2lll. c•lmbur~lz: bmlr ,\;rnH'> 80, 2" I-8J.
Bri~. D.!: t, md ( •0thn,. D. I9XX) A M1ddlc C.onway Mom~. \ II'J'Jfl) L.ne l'rccunbnan and
Canthn.m tit tit, t.lle from ~1nunt ')cepheu. Briush Cambnan ~nft-bnd•ed iJunas. Amw,,/ Rrl'lfll' 4 Et~rt/,
Columbta, Pal,lnt/ltoi,•I!Y 31, "'7<J -9~ n11d Pl.111rt"ry '1.1n11r.' 18. 101· 22.
Hn~g-;. D.£.(,_ mcl C:.11l. J.-C ( I'NO) The conunuum in Conway Morn~. \ .md Peel J ~ (llJlJO) Arucubred
soti-botht•d hll>t.l' lrllm UJII\Ittnn.ll euvtronme-nts: a halk.ieriid~ trom thl" Ltm er ( Jmbri.m of north
quantJtJtlvt tontpari,on of l'na,,i, and Carboniterous Grcl'nland. ;'\,wm· 345, H02 "i.
Kon· erv.n-l .tgt•N:iuen Jl.,/,tt'l'lm•/,!1!)' 16, 21 l4-l X. Conw:~y-Mom~. ~ l . ,\lld Wh1tt111gton. 11 B (197iJ) fht:
Brigg.. D.P.G. .1nd Kl'.u. A (11JlJ4) Dec;ly of Jmmals of tltt· llur~e~s Sh.de Sut'llt!/u .·lmmca11 241,
Bramhltl}ft•ma. mlpliL.Ililllb ti_,, ~L,tt-ti,~uc preservation 1:!2-33. (Colour phtli:S ,md tl'tumtnJctiLm~)
in Ct>lltH.Innt~ .wd mht:t pnnuti\T churd.ates. Lcchaia 26, Cook, P. and Shagold, J. (I'JH~) Pho~phoru~. pho~pho
27">-X7. rite~ .111d -.krlt'1.1l rvolution ;tt lhc Cambri.m-
Brig~. D.E G .ulll W1lby P. (19%) Tht: role of the c.al- Prnambri.lll boumhlry. :V,IIIItt' 308. 231-o.
ctum C,HbllllJ[l' <.lkllllll phO\ph.lt~o: \WltCh Ill the llUfl- Fo~tcr, M. (llJ7 1J) A rc,tppr.m,tl of 'l'ullimotrsfrmu,l?rtl?a"um,
Cr.'Lh\.l(iOil nt o;nti lwdll'd t(l,~d". }rlllntrll,i tire Geohtgical 111 ,\ /a::o11 Crrck f-'M_,;[, (ed. M 11. Ntt~:ckt). Ac.tdenuc
Som·ty of ulilcf,,,, 153. hh"> H. Prt:~s. New Y L>rk . pp. ;!(,() ~02. (W'.t~ tht) ,, )l,~twpuJ?)
Ori~. I) E C. Cl.lrk,oll, E. N K .md Aldridgt\ R.J Fr.mtcn, J.L. (llJHS) fxn·puunal prcwrv.ltlon of Eoc.xnc.:
(li.JH1) l he ti)Jlnd<1nf .mtm.ll [,.·t/1.1111 16. 1-14 (Fmt vcrtcbratc~ m thc hk< dq'o'tt of Grubc Tv1c~'cl (\Ve~t
kno\\ n 'pnuuen) Gcnnany). in l ',,,,,,.{,,,lry p,,nr/ Hr. Her.- thri• E.,•f.'~''<ll M
Ori~"'· D E.G, Ottttrdl, S 11., md R.u,wcU. R. (191}1a) ar11l El'o/llliott.IY)' Srl?mfic.wu• (c,h H B Whtmngton .md
Pynn,.Hion nf o;olt-bndtni to•.-tl,_ lkcchcr\ T nJobJrc S. ConWJ\' Mom~). Philc'WJ'Itir.ll 7 r,w, '' ''•'liS of rht
Bed. Upper Ordo\Jfi.Hl, Ne\\ Yno. <,ucc Gc,>/~11)' 19. R0)'•1l '>cVIt"ty •!/ L>u,fclll B 31 1 [!q-h_
1:!21--t Gabbott. S. E . Aldnd)o:e. R J .md Th ... ron. J !'. (1495) A M
Bn~. D.E.G .. Cl.lrk, Kl l L. and Cl.uk,on. E.N.K gtJnt conodont wich prc\cn·cd mmcle tJ~'ue ti-om the
(l'l~lb) Th,· (;r.mr.m · hnmp-1-ed', Edinburgh - a Upper Ord.-wtci.ln c1t Somh Atr1c.1 Slllllr• 374 Hl~l-3.
Lower C.u·burull'r•'l" Ktllht'rvJt- W!o(t'rstattt'. Tmusat'tious C rande, L (14~-1) P.tbt·umnlob') of the Grt>t'n R1vcr
•{the R,•)'•'l S..\1l'IY •1 J:,/i,rl•••rxll: E.rrtl; S..1met:s 82. 65-85. 1:-omuoon \\ 11h J n.:\'lt:\" of ehc.: ti\h t~una. Bul/uu, •1 M
Bnggs, D.f (,,, St\'i.'tcr, l>.lVIl.li?Jnll .'iun•c )' 11/ 11 ·Y'"""'t: 63 , I -JJJ.
(I Ql)(,) Suit hll,ht•d h.l\)Jb from ..1 Suun.m vole .mic J5h. Ht>yb, D and Poplw. C. (14RR) The fo~~ih oi
Nlllllrf 382, 2-Hi-Sil. Monr.tl'au-lt·~-M llll'S. Sciwt!/lc AlllfnColll, Sl:pt.. 71.l-6.
Budd, l; r. ( Jll%) I h,· rnmphnloh') of Op.1/•i11ia re.t:•llts Hon. X-(~ .111d lkr~strl\m, J. (IYlll) I he .mhropods oi Ra
and chc rl<Omtru. tll'n nt' the .uthropod stem-group. tht> Lowt>r C.unbn.111 ChengjiJ.Ilg ttunJ, '~Hh rdanon-
u:t/",;,,29. 1 14 shlps J nd evolut10n.1ry \l).tlllfil.Ullt', m Tile Early
Chcu, J.-Y. .u1t.l 1 rdtmJIIII, D.-D. (J')'Jl) Lower E1•tli11111>11 •!( MNtl~clrl .wd 1/11 S(~lli/1(<111(1' c>( PmMemaci, Ri1
C.unhn.111 l.ab'Cr'it:ittt• from l'h<·ngpang. Chin.1: insights Tn.\rl (c·cls A.M. Slllltl lll!lt.l .md S. Conw.\y Moms),
tor rccomtrulllll)l: l'.lrly llll't.ILn.m lifl·, in The E.trly Co~-rnbridgc Unlvu-.lly Pre~~. C.unbnJgc. pp. 171J-88.
E••o/111,..11 4 i\lrt,,;:,," .md rlrr S(t:''!fi"mt'r ~~r Pr,>blcmntic H ou, X.-G .md Jjl•rgstrllm. J (19'>7) Lower Cambn;m Ro
'J'<J.\<1 (cd~ A M '\mH1m tt.l .md S ConWJ)' Mom5), Jrthrupods of \Ollth-wt'st l 'hm.l. f·,,s,i/.1 .md Stmta, m
C1u1hru.l~c Ulll\l'Nt)i Prt·~s. I..'Jmbndgc. pp. 57-76. pr<-s~.
Ctsnc, J L. (I'J72) Bn·thcr\ Tnloh•tc Bed rev1s.itcd: Johmon, R.G . .rnd l~.xh.ml~on, E.S. (I%'J) Pennsvl-
el'olob'Y of .111 Urdn'VICI.IIl dt•epwarcr fmna. P.>sci/1,1 v1m.m mverrchr.l!cs nf tht• .'VI.I7on Crtt."k .trt'.1, llhnoa:
160. 1-2'>. the morphology .lnd .lllinitlcs or
TIIIIIHhlllllnlm !to
Colli'"· D. On~"'- D E (., ~nd Conwa:- Moms. S. Fll·lt/,,,,, Lt'c•J,,.l!Y 1 2, I IIJ-4'1. (M.mm Crct•k f;aun~) d
(1911}) 1'.1:\\ tlu~l"" \Ink ti1s'1l sttes n.•vt>al M1ddle MamU. D M. ( IIJHH) I hl pn·scrvauon of fi~h~ 111 con- Ro
Can hn.ln hunal cmnplt'\ \o1r11" 222. 163-7.
ConwJy .\-1urm, S C. (JI> ''11 Tht Uurt:•·~, Shale (Middk
Cambrian) hun.1 .l11mr.1/ Rn•1n1' •{ [;(ll[,'gy .111d
crl'tlom from the '),un.m.J Fonu.ltlon (Cretaceou,) of
Dr.lllJ. PtiiJrc•lltc>/,,1!)' 31 , l-18. (Piw,pllJtu.IOon of
muKk)
"
I!
l
Bibl iography 433
'vlartill, n .M (1941) h"'1ls nl dw -.;.lntan.• .md Craro Speml Paper No 244. pp 13-24.
Fomunom.. Br.1ztl. Ftdd Guitlt• ''' F,•.•sil.< 5. 1-159. Schram, F.R (1471) A man~c .trthropod !Tom the
PJbconrolog1nl A"<H IICHlll Mazon Creek of lllinoi\ ""d the tram-Permo- Tn.tmc
Mikuhc. I> t... Bn~"· ll.b c; .md h.Luc"~ndor{ J. Mermtom01de:1 (I nlnbltmde.l). hcldrcura Gn>l£1:1!)· 20,
(1 <J85) A Silun:m 'nli-bodu:d b1ot.1 St~tll« 228, ~G-l 02.
715-17 Schrarn. F.R. (1473) P\eudutoc:lumo~le' .md a nememne
Muller. K J ( I97'J) l'ht"pluwwpmc <),rncodc~ wnh !Tom the lllmo1' Penn,ylv.ml.ln /tJuma/ •!f P.1h-cmtt•l.~r.
prt·~crved 1ppend.1~<.:s I rum the Upper Cambnan of 47, ~H5-IJ.
Sweden. Lt·tlr""' 12 , l-27. Schram. F. (197-t) Tht• Maznu C...rc:c:k ( mdo1d ( m~tacea .
'vHillrr. K J. (l')l\2) \Vt'll'ht<:llt• \Oil F-o"1hen aus Fu:ldimw Gt·oiOI!)' 30, 'J hS
de m F rd.lltt·rrum . ,\,uum'l«t'INhll/tm 69, 241.J-54. Schram. F. (I 'J7lJ.l) Hnn'h Carbon•tcrou' to. \.tlal'oqrJ.o.
(Phmph.ltlll:d t:mn .•, ) fll·itfi<11111 40, J-J.:! 11.
MUller. K.J (I '>H l) Cnl,t.ar~.l '' 1th prl·savcd mfi pans Schram, F. ( 197'Jb) The M.won Ctcck biut.l in the con-
from th< Upprr C.1111hn.1n ,,t \w~:den . l..Ftlt,lla 16, texr of .1 Carbomtcrou' t:\Un,ll continuum. m .\1a::o11
93-109. Crrt•k FM51/s (t•d. M 11 N1tecki), At.lliemiC Pre\s, Ne-w
MUller, K.J. (I9X5) t:-u:plwll.d prcwrv.lllon 111 calcart·ous York, pp I 59-'111
noduks. 111 E\ltotolrrillloll}' h•~1i/ IJ111/III.' tht•ll l:wltWIC<Tl a11d St'ilo.~cher, A. (I 1J70) Bt:t;nfT und lkd~utung drr Fo~sil
SvolutwTI"')' Str:ll!jit·,urct (nb II.U. Whlttinb'l:On ,\nd S. Lagl·r;t.'ittcn. Nt'IIC$ Jalllilllr/1 jllr (;(·u/o~~it 1111d
Conway Morri~). f'/ti/,•wpluwl Trclll~ctcliom o( tire Royal Pa/ii(>/110/t~l/il' !11tltlcltslldic• 1, ]4-9. (What are
St><Wty 4 Lcmdo11 IJ 3 1l , h7 -7-t. Lagcmattl·n?)
Muller, K.J ~nd l. llin1 ( 19lJ I) Upper C.unbnan con- Scilacher, A, R <·il. W . h . .md We~tphal. F. ( 14H5)
odolm fn,m S\\ ctkn. h·~~''' c~ml Stro~la 28. 1-152. Sedimcntolo~icJJ, en•logic.tl and tcmporaJ pattern~ of
Muller, K.J and W.l)o,wk. D . (I<J!lS) Sbr..tcand.l, a new fossil-Ll);Cr~r:ilten. in P.\tr.tt>tdill<ll')' Pc~:>sil BrotaJ · /h(-ir
order of (, n.-t.llt t lnun thl· Uppcr C.1mbn<tn l'•f Bcolo,~iml ami [;l'clllllimtJI')' SwllflCdll(( (ed., H.B.
Va~tt:rgOLI.Hlll. ~\\ol'd<'ll. /',•Hi/, ,uu/ .\tr.l/.1 17. 1-{>5. Whitnngton .md S ( onwJy Mom,). P/11/osoplric<~f
Muller. K.J md W.1lu"t-k. D. ( l'>tlh) \f,nfiiJSSt>llill d<11(11•11.1 TmnscJCIIOih <1{ tltc R<'Y•11 \NI<'IY c•f L111dtllr B 311. 5-23
gen. et p. nnv a rm\t,tccm-llkc cuJnthropod !Tom Sddcn, P A ( IIJ'JII) Lowa Crct.11:eou' 'Ptdcn from the
the Uppt:r C.unl>ri.m Or-tt·n' ul sw~den ZMitl:l!iCcl S1erra de Mon~ech. 1\Jorch-l~m lo,p.lm J>dlou'olltc>IOJ!}'
smpr" t 5 7J •n 33. '157 H4.
Miillc:r. K.l .md WtJo,,ck. D . (l'l};"'J Morphology. Seldcn, I' Jnd Edw.1rd,, 0 (19H'J) ColotlliJtJon of the
onro~;en) md lit.: h.1h11 ol ~~\'"•"'" ; I'' !l•'"'"' &om the hnd. 11) r,,,,U/UJII .w.f lht /·owl Rt'((lrd (t·ds K. L. Alien
Upper C.unbnan lll 1\\vnk·n f'tl<SIU lllld .'itralil 19. and D E.G Un~q,r<;), Bdh.wen. Bo,wn. Ma".. pp.
1-12 ~ . 122-S::!.
Miillcr, K.J .llld W.llo"ck, J) ( I'>HH) E.xn.·m.tl morphol- Seldt•n, !>., '>heJr, W.A .md Bon.11110, l'.M. (l'.JCJI) A spi-
ogy .md lar\.11 devdl)pll\ent ol the Upper CJmbrim der J.nd other .lr.tclumh from till· Dl'\ on1.tn of New
ma..xillopoJ a,,.,f.,ums ,ulmrr,,/li/rs. r:.•ssrL< .111d Stmta 23, York, .md reuucrprctJtlOm ot Dcvouun Ar.mcac.
t-70. Palaeouwlo.I/Y 34, 24 1- Hl.
Ramsk6ld. L .1nd Hnu X (llJ91) Ne\\ early Cambrian Shear, W.A, Bon.1mo, PM .. c;ncr,on,J.D. er c~l. (ICJH4)
ammal .111d onytwphor.ul allinitle\ of cru~matic meta- Early laud .uumab 111 Nurrh Amenc1: ev1dence from
zoans. t\:aturr 351 . ..,.,<; 7 Dcvoman a~~ .lrthropocl~ fru111 G1lboa, New York.
R.irchic, A. ( I~H';) limktcl chllli<~l,'tlllt'l/11 Scourfidd, .t pro- Scietsa· 224, 492-4.
cochord.nc' fmm the S1lUnJu ofSLotlmd. Aldrcrit((ia 9 , Shc::ar, W.A.. Palnwr, J.M., Coddingtnu, J.A. and
11 7--42 BonJmo, P.M. (1lJHCJ) A 11t•vonl.ll1 spnmerct: early
R.ob1~on, RA (1~41) M11idlc- C.unbnan bmt1c divet,Jty: e-vidence of 'p1dcrs .md s1lk use. Scirllfl' 246, 4 79-H I.
example' from lou1 U1.1h Llgt'r<it:irren, m 17rr Early Stiirmcr, W. ( 1970) Suft parts ol <C:ph.llopods ,md trilo-
Er•olullo/1 of '\lrt.r.;:M .md tfrt• 'il.l!lli/i<'tlll((' o( Problcmc~tic bit.:s: some ;urpri>lllg re\ults of X - r.t)' exJnunanon of
Tax.1 (ed~ A M \unnnc:tr.l .md ";. <. 'onwJy Morris). Devoni.m ~btcs. Srima• 1170, 1300-2. (Fmt work hy
C.m1bnd~.: UniVt'N!\' Pre''· ( ,unbnd!(e. pp. 77-98. Stiim1<.:r on X-r.1y .lpplllallons, illu~tr.Ht'd)
Rolfc. W.D.l (I'JHH) Ftrh hf~· onl.md- thl E.t~t K.trkton Stiinncr, W. (198J) lntl.'rdl~llp!mJry p.llaeontolot,ry.
dt~covene' blfllr S<tt ,.,., Cc>ll<l'n'•1liclll 25. 12-R. lrmrdimJil"'"'Y Sni'IICn l~l'l'll'll' 9, 1- 14. (U\t' of X- ravs
R.olfe, W D.t.. Durmt. GP.. r .1U1tk, AT .-t al. (1990) and orhc1 rechmque~)
An ~Jrl) t.:rrl!\tn.tl hlllt.l prt·,~ncd b\ V&t:an vukaruc- Stunner, W and lkr~tromJ . (1973) Ne'" d1scovene' on
iry Ill Scorhnd. 10 I ,,fr,wiull 11/lol/ chsi/ 8il'l.ZS (cd.s M. G. tnlobttc, by X r.w,. H1/ii,,,,,,,"X''c/sn 7.ntsdn1{t 47.
Lockley md A . Rile), G ..l,J'-'!;Il-11 \<>ll<'t\' uf Amenc<t 104-41 .
C:,rtiml<'r, W .md lk•rb"trom , J (I ll711) The Jrthrupo&. Books, treatises and symposia
.\!umt,l.S/n and I ~~~Jt,,,;,,,, lro111 th( I )~vonian liOnsruck (ichnology)
'iluk P.t/;i!llllcl/c'gil, /11• 7.ut•dm_ttr 50. 7H-III.
Stliml<'r, W. 1nd llerg-;tri)m, J (I•PK) The ,u"thropod Bromlcv, R.( ;, ( ll.)l)(,) 'J'r.t.r FM<Ii< · llir•h~s:y, ·t:zpl~c'll<•my,
C/trf,,,, 1/,,, lrunt thl n,·\·oni.m Hun,ruck Shak. rmd Appftc,l/1<111.1, 2nd cdn Cho~pm.1n & Hill. London
}Jafil,,.,",f''l!tstlr,• Zmrml! 52 'i7-X I. (Esscmi.1l reJthng. bmlo~nl 'hnt)
~t(lnncr, \\'. .m cl lkn..,rri\111 J. (I ')X 1) ll'rilrbcrgirt.l, :1 Cnmcs, T.P..tnd lllrpa. _I .C. (1'>70) lr.zet· IMHis. ~et'l
xtphtl\ltran .mhrop,,,( tr"m tlw De\ oui.m Humrilck I louse Prc", ltverp<'''' (Pn>t'l'CUII1g'- of .111 tntcnu-
Slate. flc,(;lt•lll••(,,,,, thtJ Zr111111~ 55 , 237-55. nonal coniC:n.:nle; ll>llt.lim J5 pJpcr.)
\runner, W .. \, h.ur,rhnudt, r. .md /1.1inmev<"r, H.l•. Donovan, S.K (cd.) ( 1')1H) '11u flctf.m•f,j,,f,>l!)' ·~( TrJtt
14HO) Vl'l"'tl'll1l rte~ I l'l't:ll 1111 lti.itH~t'tilic.ht. Kfmu l·omfs Wtler. Chtehc-rcr. ( 11 p.1per;) l"ur
Smckm/•oJ: Rnltt' 11 , 1-Xo. (Exc~.·U ..·n; illmtr.mons of Fnry, R.. W (eJ.) ( JIJ76) 'f1tr Stud)• •'I Trtkt Ft>Sltk
llumrilrk,rludi:r t 111n.1) !:>pnngcr-V crl.t~. lkrhn. (I m .tlu.1hk cumptl.ttton \,f
Thomp,nn, I. ( I() 77) I· rr.tnt polvr:h.Jlte' (Anndtda) from ongtnJI p1pcrs and anrrodurtnry ch.1ptcr')
the P ..·nnsylv.ull.lll r\\.:X f .IUIM of Northenl Illinois. Hamzschd. W. (tY7C,) Tr,Kt' lu"tl' .md Prublcm.tticJ, in
Jl.tlncollf<~~r,,J,ftic.t I 163, I (l<) <)<) Tmllist' 1111 IIIJJcrtrllftllr P.!lnllltc>f(lt:Y· l'.trt W, :\lmdhmm,
l'homp~on, l (1 'J7'J) M 111111 ('reek. 111 17u· l; myc/,,petlin qj Supplemmt (cd C. r ctchcrt). Geolo~ric,tl !:!octt'ty of
J>ah·,mt.lft'l!)' (•·d, R W F.urbmlg..· .md n. Jablonski), Amcnc.1 .md Un !Vcf\lty of K,m,,,., Prt'S\. Lawn.'ncc,
DmvJcn. Hutthu!SOII ;and Ross, StrouJshurg, l'cnn., Km. (De~oiptiom Jnd rig-urt·~ o[ all l..nown ~cucra)
pp. ~h '\ -~J (U,dul Sllllllll.lry) Sch:UC:r, W. (1972) Ec,•!tw>' mu/ Pt~/an•l'ccl!t~r:y t1( M<lrilll' tw
Wa)o,wk , ]) ( I 'J<JJ) Tltl Uppl't Cu11bra,m Rch/oac!lid/" Jlrii'irc>wm·rl/.1 , Oliver &. l3oyd, Edinbur!;h. lrl'l
•l t~J
,aud the phylu~-:cny of Br.111cluopud.a and CrustJcca.
hmtl.c a11d Stftll•l 32 1-202 A en
ilil\1
Walo,~;ek, [) md Mi'!lk1, K,l. (I'Jllll) Uppl·r Cunbri.an Individual pape rs (ichnology) 4ltft1
~r,·m-lmt·.a~t· l rtl\tall 111' Jnd thnt bt·Jnn~o: upon the .. ;fht
mmloph\·lc:nl nng111 ol <. ru,t.lt e.t md the po~1tion of Bcmon, MJ .llld Trt'WIIt, N 11 ( IIJHO) Dittyc>dtlr.l lro111 Atrt
.-l![lhl.</11>, 1-t·tlt.lltl 23, -lll'l ::!7 the Silun.1n of Pcl·bk,htre, ",lotl.md. P.IIJWIII••f~l)' 23 . lrr••
W.llu"ek. D. ,111d MliUet, K.l. (IY'J4). Pmt.t\mmtd para- 501-13. ,·Jar,
'irt·, !Tom tilt" I cm u P 1Licozoac of Sweden. Goldring. R (I 91>.:?) Tr.1n· 1\,"j(, .md tht· wduncntu\ lr/11
~urt'au: m ,hJUtl\~ \\.tll'r .mJ mannc \l'dimcnl,, m A.ru.

I''"' .lC//<1111 •!I tltr R..Jy,ll '''' 11'1)' ••/ b/rubur-.:11: E.mh Acru
"'lt'll<tf 85, 1-J7. Dt'lratc mu/ '>ll<lffc•u• ,H.tnllt lJcp<•SII> (ed. l-J.M L. Vm A,w
\Vilby. P.R. ( l'l'!h) l\1mcr.tli,.mon of,oft-bodied mvcrtc- Str.1.1ten), [1.,.·\'acr. Am,terdam. pp I Vr-43 ,,,,.
br.ltl'' m .1 _lur.t,,ic llll'(l)hkrnu~; dt·po\lt. Cc·c•f.>._l(y 24. (De,criprion of Drph><l<lft'IWII JnJ other tra(e io.,th)
H-l7-C,II PtekenU, lt. ( I '194) Nomcncl.tture ~nd tJxonom) ot '""'
lrrm
Willi.um, L A. ( 1'1:-U) I kpo,ition ol the Bear Gulch invcrtcbr.lte rr.act• fn~'ill\, in /ht' J>.llacc>lllt>h~l!Y <!I 'fr.z;r ltlth"•
it't't't
Lime\tolll' .1 Ctrbc,tut'l n1ll\ {Jf,mwkalk ti-nm ccntr.1l Ft•>Si(; (ed. S.K. l>onm·.1n). W1lt>y, ChidJe,ter, pp.
• I.tit'/••
Mont.111a Sc•d1111t "''''•~~)' 30. H-t' ·llll 1-1.2 . Agl.t·
Z.mgcrl. R. uul Ril h.trd,uu, .E:..~ ( (9113) Pakoccology of Scilacher, A. ( I'JI.~) liaogt•mc scdtmcntJ.ry structure~. m lgl«
two 11cnn .. ylv,mi.ln l::ll.td;. 5h.dc, ltddral/<1, Cwf·~~y .-!ppma.-fw, t•• P,lf<li'<IN<II<II!Y (eds J lmbnt!' .1nd N.D. AI'm
.\felll<ltrs 4, I-JS2 (Cl.l\Sit p.tpcr) NL'\ovcll), Wrlcy. Nt:w Yotk. pp. 2Wr-316. Al{nc
(CI.t~'\ific,tti()n of rr.1n: f(lsstls) A~-:11<
Sc,Jachcr, A. ( (9(,7) ll.lth y ltl l'try of Lr.1cc fm\lk ,\J,trillt' .·lgm>·

AKOll
Ceuf,,_~y 5, 4l 3 2H.
lt<Uir
Simpson, S. ( 1''7h) Cl:a>~1hCJlto n nl tr.acc fmstls, m 711r Alcv,
Study '!f l-,.,,a• l·<~ui/; (cd. R W l·rl'v), "lpnnger-VcrlJg, lit Y'''
Ut:rlin, pp ..W-54 1/,/,,,,
J/r·rc.lt
Arn111
Arnm
imph
Amp I
.-fltiJ>i<
1onru
' "'l'Y·
~•loll•.r
An.lh.
Systematic index
.4l•~rsc>,~?rJ1,tu• .nx Att."<J 14"> J·tucno' .:!X'J

Ar.mth,wrr I .\' ''"'-'1'"'" J'J'I 11lhn'Mit\llb2't')
Ac.uuhogr~pud.cc .t!f, Ancyloccranda 247 A'tcrmdc.c :!11'1. :!IlK
.lc.:mh<!flr.tptw .\2h ,,,,.h,..t',llJ ~46 r1<lnc'f'Y~t· 4211
.1c.rntll<>sqmll.t 11.;!7 Anunalr.o57 A,r,•ruma 1711,254 , Dllll
Ar<llltlmtlllrll 1111~. 1 1 1~ Atmograpudae 327. 3~5 A'rrocnl'ntllu I U
A cam 3'i2ff. \t>5. J7:!. JK2. /1 .!, I 1.111. .rlttiSt~~riiJ>IIIS 327. 331 • I J/Mmlll.l J07
1/.20 , l11iwtrypo1 h.8, 1~9 .1 "''''l''llll'< 9. W, \ll I. J I 1
Arc·ma'J'is I/ , /I 11 ~'"""''''~·.ms ~ 13. 415. 11. 7, .J 17 1/.11'<\~'•'J'IItl 1.\.ifl
.'lcm•11lana I 17, l21J, ' 7 Anumalodc;mata 210 Ardu<tOIII.lt.l :!HO
.lcida>pidl>S, \!12 lttMtflo>py~us 9.19 ltlr/C'f,, R IX
Acroechmou.k.c 2711 1111.-d<'>l 9.35. 291 Arhvntl111a 11!~
."irrc>l"'''' 1.l2fT. ; 17 Anthozoa 1114if ltlrytl.• IK.!
Atrcl~alrm~l 270, V7 lutlrrt~am.u,J 1.20 . lrt.rir,>,tf/,r ICII!
.·lm>trrtJ 1!!1 tr..~er.1p11H 33~ ~ltrn>~ 7. 7, 17~. lR2, 1>6
Aaom:nda 11\1 Af>OdJ :!)!(, .-lrryt•l'l/.r 1'1.'
~liJ<'Ih ~ 12, 1.! 2 Arr, po.l.t I ~2
!J''"'""s 22h
~ ll~<'ltrll.t li Hi ~rtrltduuu.< 264. 9. 'I Alllnol 2.\:i, 2.\K, 251
.lam<>e.:lll.l.\ 254. 8 Jl t\r.~<hnida H9. 3AA ~~~~~ ..l(( IJS .:!SJ. ~ !,!
Afliii<'Wr.tJ 2.,5 ~r~.I""'J'h)·llum 11-l, 12ft Aul.c-<>e<·rr.l.o.!S 1
Acnn•>ct'r:Jiln~ .!Vi -I mat~" Y J, 21u. no. v 15 .lu/,r"'/ lwrJ \')'•1, \X:!
Aronnc:nmd.lt> 2'16 '"" 2«1!>. :! Ill lu/r,/111111> ;!711, Y 7, '1./l, 290, J l J
.lrruuvnmft~ ~~)fl ,JrdtJf<'<ldolti> :!70, 274, 9 1-1. 'I 15 Aulm<l ~.Ill
A am« y.ulmt 5 Ill Arcl>a~"', Jth.. 88. 951T ....,.,.,,)'llltiH 11 ~. ~ 10, 12(1if
'!rttllO>t"""·' '11, .J Y, I I 11, -1 11 A re h.tc1xkndnda 331 ~tu/,,r,lr.t 126. 2~ 7
.~.J•.-n.~ y.•tttcs 2K7 Art:hacoh-a'rropod.l224 ,lut.wm/1., -1. If,
.!~~fr425 .l rdlJt'<>•prr.r J.J lu'"''''(2~..t ..~ J2
~~c>liumtitn 2k7 Ardut''}utpJ.,,re--122. 11.'1 Ju,/f•"/'111/cr lb'>
Agh,pcd.c .Htl lrdttmcdes 6.13. 149tT A)'h,·c.I.H'J . .!lJ. 1.!.7
Ag/a,pu 11 2.1, .1'11 lrdritmtlus 252
Agnma 70 .olrunJt'•l 210 IJ.tbllllicr ~ 12
A~'11mtitb I I 20, .:11>2 • .\l•hlf•.'lh • .\KO ~''''""' ,, 215, R. 13 lloltlnll'\ 211>
Aglll.>\ttn.c JKO ~f\!~1111111>1 252, 8.31 UJcmnn.t :!35. 24<•
.l.~llMtrtdHO, 11 .!U, 12 6, 1::!0 .lnstaqsmes 9.45, 301 B.olncll.lrbiitl.cl' /I .21, \H-1
Agomamin.1 2-ih Arthroplcura .349 /l.llttwm~·l•tml• \'1 1 , \•n. \'/ol. 11 ..!•/,
·lt~ricy«tlws 95, I 17 Arthropod.t HXtf /I .u.
Akynnact'<l I OK ArttculJta IJ.rrr.mdc,•cnlllt< 21)(,
.1/cymrtoi.mu 150 13r.Jduopoda 159 lk<kll•ir/ua \Ill
At.lattdhr bl, ~ 1 l nnoukt297 lklt·mnlll'' 2~~
4/l'r••ltlt'• 121! <\•aphocda Ju7. 3!!2 l:kkmnmd.l .:!5~
Ammomuda 24211, 2·17 • u.•phord1dmus 1 I .5, 372 J-ldrnll~»tmtllls 25S
Ammonocdea 2311, 2JKtt .1sJp/uu 379. 1 1.19, J82 Hdmuru• \'1 I
AmpiiiC<IIIol 7. , .:!' no lltOit(J< 2J'i Hdlcrc'/'ltt•ll .:!.21•
Amphincura (I' oh plJ< "t'l" '"' I'i'l <\••:n<L·nd.t 2J5 Udkruphntll3Cl'J 2:!t•tf
~mf'/t'.\IH ;, /I/, 11:i A•cophor.a 145 8.-lc>Jcpi 1 .\55
, lmpul/,•tWtfolii IX itsr<rylllttm8 1.?.21'1 Hdc>tr/sc•ll I :!2
''"'I'~ .\t(! A•r•dodurora 28u li<rcttlu 1 6 I I, 14'). h.!
Au.rb..nt()ul, J.l 'l.ll~nt 5.12. 214 n,,,,, 7 ''
Anarn·,nda 2-16 IJitnJ,~If.> 2o. 11.12 llrllw.~·<ll,, • 1 !, 182
436 Systematic index
Birh •..-.~.w"'" /!i/1 (",uyc ... nmtt.~ 9,-/.f OyJ'I'IU:!71l

Btv.tlna 1113, IQ'I , ;!11.'11 Co~ryophyllio~ HIR. 1 4. I :!S, U:! l nrd.ma 10211'
BIJ>!Otdo :?.t•:!, .\1 q C.uyophyUma 13:! CO<!tht·t"' 2-l'l
Hb•r<>zoJ .!lt:?., 'Ill,'"" CaNduloHh 2711 c:,,,i,hto Y.4X
8.\IJJ,JIIllr'i :!50 r..lj(JihiJiclrl , :! • 1511 t {>do:nt~r•ta "' l 'n•dJrt•
IJ.•r<><wlr/lum 11 X c;.,r.e }'\?" I!\2 c;,,,.,,,.,/c,rrr"' 'I IIJ
&Jttr.J 211\, H 14 CJ!..tp)')!lll 9 7. 'J.!-1 C,'IIIJ'<>JIIol ;,8, 11!:!
B.•rlm,,lJ.m, •J I 'l 27f>. 31 ' C.w,,.n,, 9.42 C:.•rult.•J'<ItH Hli
B.•rrr,•tlts 1/.111 CNIIf•'tldLI 1X2 c;,,,hluHtlUIW .!.:! 1' N I h
&•,.n/·,ml:l,, 6 I, 11 liT C•·nrr<>ncllodm~ 174, l!Q (.',,,..,,,..,,,I!! I
Hnduopod.> 1•7. I H, ISHrl, II\ HI c,.,,,~,t•rh.mus !J.lJ c;..,,/,lf!J 107
Br.Jdt)'"'l'l·lwtt /I I~ C cplulo.:-and.t 397 CuuulttJ ltl7
13raclwura 41~1 CtJ•lo<l/o•dt<(lb .\0(), ':!h c:~.•tmltu Y .!.2.
Hr~dunod• ~ 17 Ct'plooJI<..craptu> '137 (""II'IS 2::! 4
Hr.w.-lth~Jn < 4llX, 11 t. Ccph.Uopoda :!1)). :!::!IJ!T <..or~l,uHorph.t 1.\K
Bri'd<'WH 12.1. 4:!11 c~r.llllvf~lil 149 C.~rf•u/r 21!1
B"'"'l'"' !J 1
I, i /'i (.,·wt<>Jmtt.J 10-IIT 1/. 7, ::!IIH, .2111, 21 ::!. (.'c.Jfrthf'"J l.l, ].~(, o.,
Bry<'~'"l'"" J27 l' I \411 21-l ( omut.l \ll7tl v.r
Bryozo.t 1.>7. I~ 311 Cer.ui.xo~ri, I 1.1 i. 3'J!! CuryncxuduJ.t JH(J Oo1
B,.WIIIIIII .!2, X./ r., :?:?4 Cl'rJUtid.t 242 r,,,.,,,,.,;t/r•.• .l27. Ill. I/ D,1
Btwllrll-llt,l51 (.', ro~tortplwlu JG7, 375, 380 c;,,,lll<lh1Jrlll· 421>, 12. 12 v ..
814/m,JIII!ntsl,s 1/) 11 Ct-r.J(j1{ySIIS I) 49, 307 Cosristricklmufitl I HI! DJ
Bwuam1s 'llO Cn.wnrs 11. 13, 3 70, 382 C.•rl"'"""' 5' I 9. lJR DJ
Bwwtlt> 3'12 Cen.mupathari o I 07tT ( l1lhOIIItlllitdJ I()<) I I X, 1>8 Dt1
Bu~~""''' 412, I2 ..! c..,,,,,, 224 Cr•llm""~Y<II• 9.4'1, 104. \117 ~1.; /),1
Bu~~e"''"'·"'lor -1 H. I.: .J C/o,lt·trfcs 90, 4 ., Cwro/ri,r252 De·
Clomurfl,>ri.s 3.2 Cmu.r 7.4, IH4, 111:; o,,
C.m.•mriiiJ 2lJCt, 'J 17 C/tolr11111J1<11H 54 l / 1 I08, 1J7 0 4 I ' Ct.IIIIUII IK I TAl
C.t./omr/1,, I S2 Choulllo>fi 357, 31:12 ( rAilltfimm•J 151'. 17Xtl; llJ.."'>IrnpuJ~ 22 ~ Chnlmromara 145, ISO Cruti•>rs 171l, I X I Vrt
CakarcJ \CJI<t>pnnj,\<a) t)Htl Cbcmx:nnid.tc 2'1i Cr.LC.oltr/1~2111 De~
C!l.t,,/., ; IIJ, 120 Chl"mtnn<~ 382 C:rt<\(l.\l~t •'
2:!U o..
C~<1<hurJat1 25!tf Chunrnu 3b2. 11 . .!U Cr.u mr./rmu• 2711, 272 Der
C..:a/r,,l"''·' 1 I4 12h Chchccuca J~<J • .l!!Htf Crrphlot"llltl•• 6 .!, 151) De1
c..1.~-...u s 24, K.JI Cltdomor//t1114:!1 Cn~n/w.J 1511 o.,
CJ)C"" hJ.t<' 12~ Clull' ·.n. IRI \.nhrunnrph~ 15.\, h Y Dt·t
r..,/n<lyl" 12•1 ( 'lutllolllohlrr 28') Cnnotd~J 2n2, 2'JIIi 01'1,
Cttlyllr<ru '4'1, Jh2, 11 6, \~:!. /I ..!1/ Uur,,,. X.3. 168. ll-12 CnnozoJ 2h2Jf. 2'llli o,.,
C.tl).mcrunJ Jloh Chtkau 11!1 Cn•rot 14'1 Dt'l
CJlypwhh.wu 1!14 ChlidNtophor-,o 185 Ctt<lolld/.rl-17 o...
C.t/y.dur.lnlllr J.\2 Cho:mofi.Igdhd.t 1\5 Cro:<lllo1'J"' 11 li On
(..,un,uodum,t .!711 Choncndina 18:! r.nurlfJI/r•lu . . l('f!''~ I K2. 7 I 7. 1'J I CnNau.t .\I'J, J<J7tf o;,,
l'.unbr,bln 'I 1 Chord.tta 311':1, 311! ( 'nM111 dCJ .\.!!! U tb
Cambr~d•"'"' i 2 C/o,>riSIOC<'W 2~'1' 8.30 CntZI<IIhl 370£f. 42Ci, 11 . 15, 12. 12 01111
Cm,,(,r4JtHiwlu.\ hi Cul~m :!70, 9. i. 9./9, 9.22 Cry pt .. dunl.l 210 Ora·
Camer~t.• 2'1JIT. 1lJh C td,Jrotd.t 270, 273 Cr~ pru~apttdJ~ J.H D1re
Campto\trtllll,lfOtd,·.o Y I I ,,\ 1 I CJJuldJ 2lJJ Crypro~~'''I'IIIS 334. 10.12 Did
C.ltttpy!.•rtpltollw ~!!4, I 1..!5 C/mkfi,r J93, 3!16 Crypwlit/w, 31iJfl. J(•7. 1 J.Jn Dirh
Campyl.vrra• .~ :? ~ Clt't>lloyri,/,.,,, 7 9 ( :rvpm,tmn.tc.r 149 Dtdt
Cmr<lil<hJ'r' 11 2. 1:!.:!. 41h c:lmta<OJi'•lJ'IIh J2!!, 11!.4, JIJ .:i, /(1,/-/ Crenodonu 2 I 0 Die•
C:anatl•~ ! I 1 Clr~ll.r8<> Ctcnophor~ 101 Dtm
r:auim',, l.:!tl, 1 ~~ Cltwml>~mte•
182 Crmo•py~r \f\4 Dicr,
Cat1t1Jlt.lcclmh f ! ClotJmbumudin.I IR:! ( 'reumtom.tta 14\, I 511 Drrto
Cll'f1cltlllt'l•l~2t• f .l~ll••gr.lptus 327. J31 , .Htl Cul11d1111.1 4:!1> Drrr1
C:.mlt,'<.-rJ< .:!47 CINurdJ 11!8 Cttrttllo,~••rrul Ill 14. H7 Drn1
Cardto..crJtrcLc .:!4~ <:l,'llamJ t>l C:urul.t./n·., lr ..l 15u. I:;:; Dr.-r
C.lrJt ..'):r,zptur ,34:::; ("[)"IIC!II<I:! 12, 24fl (.ul'ul,,nlltll:!'l'. SI 16, <; ~~~ D•tt)
(.,.,.Jro/.r ~I o ( 'h mo:nuJ.l 242. :!46 C)•.t~lr.tWIIIoiiN,1.:!.:! D td)
C.milr.t 211l Clrmmt>u.urr,Ju, 23!> Cl'•"'""nmt<S 9 lS Dttf)•
C,mfumuu C.u,r~d1111~1 ClyJ'•'•'''" 9.19. 1i0 C 1·,Jtlho(}'>lll 2K7 I.Jrtl.
f:,Jf;,.l/HUif "\~h \ h pr:t>trrot.h 2711 C}·l~/.u,ks \71, 11 1/ V(~
Systematic index 437
r.yd.uuutlr.m.J IX:!, 7. 1-, ~ I 7 Drrwrplrt,/lllt/o 373, 11. 15 Elcuthcmblamu.t lll4
Cydcf)Wt 125 , 12. 1! n"'"''l'lllllitts 8 15 Ekuthaozoa 2115
C yclocy<toJd< a .:!hJ Utmorphob'T3prid.lc :US Flr.ttlwt.lli2
CydomdJh.l '\'l, l I Dt>IIO'J'h-~"'P"U 10 12. 315 Lmtr.ziJrll.t 112, 12. 1.!
c:nl•·t•rx• lX:! 11 111 nm~b.·/,, till Emrrr/111 ,\HO
( \'ciO>tODLI!J I ~~~ LJw,>muduu 11.-1,-115,41 7 l:ncnoundJc JS<J
Cypr.tFJ ~:!4 o.,,,,,, 188 l:lllnmmo5 I I . I>, 3''i'J 'h:;, .lb7
Cyt!l.r ~. H. 17.:?, Ill.?, IX'\ Dirhrrhrll•m• J.to /;tliTIIIIH 2'11t, 9,-/ I
Cj'riN'•'f'lll< l~!l .U.l, .\.11>11 , ;\.Nif , 111. /i D•pl~hmtrs 373 F.nJ,~nl\ 2.35
10. 18 /JJpliktr.u 8.25 Endo.:l·raodl 235
Cy<~•phylhw 1111 Drp/.•mtmorr 11. 1:!, -l:!o, -129 buis/J. /1.:?14
Cymr•hyllu"' 115. 12111 , ~to D1plo~pcacn 328. H-1 bmd~tl<:l':l 171
Cy>IC'\!r.IJ'(Itj Ill .~ D•plngrJpnru 321! hllt<•lt•t•h)•llrrm 5 ~, 117, 120
Cy>tOJd, <<t D1plnponu, RIHllllbJtrr.• LJ.pf<~~r,1p111S 10.2, 3:!1), J3l l::.ntt·r(>pnrll)tJ 3:!9
Cv>toporau 14CJ D1ploponu 262. 30 I tr EocJtldJ .\99
Diplcm.r 133 EmiiOtl('/c' I 0
V.zl.z4:!0 /)UIIIM :!7() / ;',l(,lr/ld li!ll
o,,fmmrf/1,, IHl L>1~m~roic.lcil 270 l::.ocn1101dc.• 262
o,,fmmlllt$ lR2 D1K1nub 181 l::.ow<einJ 1r.7
0Jhn.tniud,t<' lH.:! Dmilli.•rcc 177,331 f:t>dt.<llls I I .X, JKO, // ..!()
[J,,f,wmri•w I I .N Dmc'<'T05 235 h>lillrrlilll3<.J2
n~mestlla 382 Disrc>id•·.r -13 r.·~"''"~iui/er.r I !12
D.u yolvsiu 7. 9 /Ji<wson•s 235 r:~llfto.to<clomto 10, Holff, 7 5, 7.6, 173, Ill:?
f>.ltlliJOIIIU 182 D"p mJA :!93 J-'.>JIO'flllt159
Dny•.t 182 Dithc•;ndmdnmt tu. I/ 1-i<l[>tm.•ll!l
Oc,Jpodl (Crult.llca) Jll<l D11heco1dc.l 30 ltf bmhunJ .:?75, :!K(o, .:!1)11
Drtt''"'" /I 6, lR2 Dlln•r·• 5 4, IOR £p,.,,..,9 ,r,2xo~
D<k.Jyr.J 6 8 JJw,m,tfl,, X. 11, 2 14 Endnph~llu111 I"'
Demmpon~J Ktltf Dll't'hc').!rJpllls 331! 1-rymcl I I .!7
DmJt.JJttr 211') Vu~·opl•ylluml:!l, 5 . 1! bryomdtc -11111
Ocndro• hirot4 21!1• D,•/r.ht•[ll(no; 3'H uk."t'" 11 i, I I 11 , 1t.5
1Jr,,f,,,.nt,h 2•no· Dom1chnaJ -128 T:tlund 215
0.'tldro.:yu., ..ltr 9.49. .\'17 o.....,x :!0'1, 21-1 t' uctnd~ 111<1
Drndl'\lgupud.le J21t I )rcp.mochuc~ 3.3 t.:uc,hono•deJ :!711
O.nJr<')lr.Jplw• _\,?(), .l.'!h, /Cl I, Ill / I DrepJnoptc:ru> 39-1 cJt(!llll.llntlll! 2')tt
DrnJrmdt-:t J18. 320tf, .\.'!1> o,,,JI')'ol>f'l' 11 11, 375 [uniJI.Itc"ln<J 3 1111
O.uJrt•pltyllr.l IJ2. U5 [>,,,.,[, 25-1, 8 ~ 1 f:u,mrphalus 224, 8. 1.\
Dtutalmm ll -1, 19'1 l:.uphaU\IJCCl J99
DruttrO>t"m•~ hb Echm•ccJ 270 J.ul•lc.trlla 114
O.t\'llolllrr 21!11 Fcllltlfl{,ll•/111111 '267, 275, 2l:l5, !1.5 9- F.upmnp' //.H. Wt, 4.:!.:!
Dexwrhcoc~ 30'1 F.dtlllc>i<lrys 'I ]9, 284 Euryabc :!')(I
Drn.lemu 270, 9 :! f [;, /llll<'i)'.illltS 270 Euryptcnd~ 349 3!/'l, V/4
D1Jdem.1tat c• .270 Edtinocy~mc:uJc:a 2711 Furyptcrtnd 19·1
O..zJrmopru :!!i<l l:.dunudennat:a 263ff rurypl• nH 392, W4
Dtbrancru~rc 230 F dunotdc~ 263 Ew~yr.r 210
Di~tmopl•yllum ~. I() Ec!Jirrosl'ilaerilrs 9.45• .30 I
f)t(£//i>jlrolplru 10. /1 , Ill Ill, l:!I! • •U:!. 3-1~ Erlwwtlmri•• 270 fod~(<ld 11.2 5
01,11as:! lfl, R IJ, 217 Edtinmo.l 2C>3tf 1-a/,,f(r 7. 17, I H5
D1chogr;~pnn,1 327, J I~ /•'drrtllls26'.:!ff, 9.1, 9.2. 9 .3, 9.7, 9./9, Pavima 132
DichoxrJplu.< J27 q !:! l'a••u•ite.< 1 2~1T. 'i I.J, 127. 1v.tr
D•.km.w11ia 59, J . I fdmutl•••rirtus 293, 9. 'it, 413.416 J'JV0\1(111.1 127
D1mnograpuwc 328 Frtl'llt.iiiiiiiS 9,9, 272, :!90, 313 f't•lt! ')
OiJJII<I~raptrts J21\, I 0. /1, Ill /I, .U:! l:ccocochlel .:?3P hllfstr/J.r 6 I J, I 411fT
Drmmmu 11 ,!0 E.ropmcta I·H l"tncm;~r.l 119
Ot.:tn•o.nru11 2 1H, 9.J4 hllllnmll•'.:!lfl r'""·ul.\1.1 'Oh
0..7yo>t-/ol5/llj I!!.'! EJnt>JJtlr ~ - f.:!, 287 h•rultJI<>rJ 14'J
DiayoJora -12H, 12 .I! E'dnoasterotdcd '.:!63 rJ'tuhpont.l 14'1
0J£t)'OntllidinJ 11\J Ednobll<midc1 317 FlcXJb1li 1 2'1:\tf. :?91,
!XI)•'"'''"" Ill. 7, J.l I F.i<hu~ld1o1 183 n,,.,,,,l54
Duh mogr.apuru .1:!8 Eh"podJ::!llo Fodnuchnu -l'.:!ll
o..lym".'(r~ptu• 111. 11, /() I:!. l:!h ,.Hl L.llmntrtl(er.U 235 h•lh>lllf111119J
Dltl.um.J I ~J Elle\meroct-ratinol 2.15 f',•mitolwlloJ I l
Digomtlla I KJ f:lllt'""'l'''olllh 356. 3ll2 f,.,,ft/la 2liJ, 210
rlH~ul.t-;1(, 2).( 1 ll.trl• •rlt.tno>tiiJ •; 4 '· Jn I u...,,, 21'1 I

I C'''IIUiltfUI t4~.2. f2 'I I ldl•f>rhrm I tJ m., IIH' \xo L
J-u~-:~d1nu 1:!'1 lluro 11 Ill, JIQ, 11 .!0 ln.1clttn ttd 2'1.\lf l
l-ung~ ~7 IIJrpmJ .\c.S, 1X:! louru.Jtl.tta 15X I
flii1!!J3 5.16. IJ:! 1/d,,.,,.r//J b.l-• .!<10. 221, 221r lu~ ur.,,u, :nu L
rungnn~ l \:! I ftolt{upl~(otdc·J .!••.\ '11 ln"•to.'\1'1 L
hi\ 11111111111 117 11 lr..yl.uw Y. 50 CJ 'il. Jll lrrq.'lli~H \ ''"'' l
//d1"btd 10~. 5 1~. 1~1> lr"t,'llldna 271) I
G.thxC'.l ' I>. 1:\11 lkholmru 1~3. 1:!~ I> lli1Jt I I J.:! I
(;.1\tru~urLI ~ I !i 11 lr<'f'l>yllwn 1()'1, > .J, 1211, Ut., > IS I·•'X'~I''"' \.:!-, I tl 11, I /1 I- L
CJ\lnopo.l 2f~l ~IT 1111<'1~"·'1(11! I,.,,,(,H 17'. I I IS' l
Cd•w~<•-u !!711 11 rttJ,J1f!tt \H:! L
(-;,,,,(1/Jr." 111:!,:? 17 llcnudt.lrd.lu .1 I x. 3:!'1· / rd:rlo>J''(III \<1-1 I
(,c-rhurou-r,UIIIL 'Ut, Ht•m,,,f,m, :!IIJ L
( tlfol~lh'.!fu 'Sh Hc•111u o'nund:~t' r; 4:; "'·'f't'~'''f'/lt( \27 10 ..\ L
Gtn•rllr.r2IO Homr/rrp<"~lll·• 221 Krll•ll•lt••rllylh" :; I'J f.
( ,i/•J.orhprr/u,, IK.2 flt~u~f:.!J,l., ,,1 X 2 3. :?3H. :!~ I K1lhudurpl1)111d.o lfl'J. UH L
C;,J:•""•'J'h'·flltlth I H 1 Ht"'>p<'rotth" 7 11. I X>i 1\'lllli>l'll·/l.r ,I). 5 I. I 07 l·
(;tgflllftl'r'lll\!lol tJI) Hc•tm\mlroltts 27~. 9.!2 1-.rtklllrlllll - /!, 17.:! [,
G/,ol>r~nll~llllllll 11. IS 1lclc fOCfYoiS 2-1') J..:l,utrt'hl ltrS I~
G/ol>llol 11-\2 1kt~rocm.,(h,l I0':1, I JH J..:,,,Jr;IIIOolllt/11• -.17. I fl7 L
(;/, "''""''")'"'"" ')() ll t·tc·rtulonHA 203. 21(1 1\o•tlollltiJIII)'I/um I I~. 5 7, ' Ill Lt
G/,>/lr•M•""'' iltlo 1/tll ro>plryllr.t 5 I <J /<..,>,>11'/tlol \1>7, I 1..!(1 L.
(;]o<"X~t.l• .2.1S. S .!J 1/rrtr,>rrhi.< I !13 1-.clltiii•I~I•IJIIII• .ti I l;
lrl"""lo:IJI 1111.1 \.:!K I ln.llllllelltdJ -lil, l\XtT. 4 I~. 9-ltT J\.•HIH4'i I'M\ ~ \H L
( ,/(lj}tl~'·'l'"h \~ti Hr\ r~~'-'ll1lfhf 1CJlJ ~t•HI\0.\f'\ !.2..!, J.! 'J L'
Ctutndlo li(> ll""•pnd.l 3-1'1 Krrt•''.~"'''.., I I, lXI l.
Glr I"""
m .:!tlll, HIll .:!ltt :!I\ 1/~o~tcll.r ~./.!, 21'l K utt>q.:u 1.!.1 I K I h
(.lypt<I<),UCI<ll ., Ill llthberrnrterina 39-1 L
GIYJII~I·IJ'tl" Ill lt Ill 11 Ill u '~~ H·l·l..tc>]'IWh 34-1 lol/oftlll•l 111, -1 Ill
33.:!. '1\ Hrbomc ::!S-1, li 32 l..l/l('dllr/1<1 111 If
Glrr'''"'' - 1' /Jrl•l•'<f3 8 2/ l <atdlt l!\.1 \I
t ,/IJ'l'-"J'I' frtt '14-i I ftl'l'""'' 5 2111, 211 I ~""'1'''1'1/llm I 21 , ' I 2 \I
(,,~UJ 301 lllppuntur.l• :!I it I un~l!thrcm< l111 " lil\ Jl\ 1.1 \1
( "-'IIIJliM"a ; 1,. llmt ""'·' 1'13 Lunuun• 1-1.1 \I
C.•m.mtcs ~~~, 122 /1,•11)1(1 270 loiJ'Ih>rl/lt/l.o l ! , I I \1
(1UIII.UUhJ.i ~ .. ::! .:!4h llu!J tnn•.L 27() t.rt /r,r I.!J , U 'I \1
CLllltllJ'h\ h:rn I "~tt I l4>l<Ct\ puodJ 27P 1. uu/l,rr/r,t 112, 12 2, -117 \I
( oOrf!OIIJl"~l 101-1 1/.•lt<I)'J'II! 2711. 11 ! I' 'I .! ' l"l''''"'lr//J '\SJ, I/ .! ! Ill
(ri.tr~llllf•t lnH i j 1/,>lml,l 11 6 L,,·u.u~y,,,,. '""() t\1
C:o•llh'glollllill /11 /lr, 1111 H•IVC)'<IIIc < 9.-i'i. "\fl) /~''"'''1'" 11 /I, /I /fo, )7'\ lS:! M
Lr..rmmy~r,J ~N•J H,•lc•p/u,,~.,.,, I:!U. i I! 14·rr.lo tlu·< 2711 1\1
c,,,,,,,,., •.u i'i 2- f/,•h•rlrurht '1.30, ::!1>5 L~ ptd~'thtdJ~ 2741 M
(;r,tpwhrhl!l.l 1 I tllf llnlollmrmJt'J :!h), :!!15 I' pHiliH'IIllld ... :!71) M
( ;r,opwl<Htl~.t J I Kll l2r.tl I Iun1.Jinrund.IC \x2 I ,.1""'''"1""' '7-1 ,I/,
Cr;WI<JI) lllcllt.' .17'1 Homaloznd :!62 I t'pHl<ll }'\IOt<kt .:'Id . .' I I H ,\{.
Crrrr'mkirl l2 l llomorosrdra Jll'l Uf>/c'llillol 7 11 If,
c'"'·H•HWIM H 2 i 1/o•t•fit.,; 2-17 Le!''''""' IH2. ftl.! .H
(;ry,),}lflfl :"1 i llnpln,·.mol.o.'\'111 I~J'I"~"'I'''" I(} J.l. l.!H, Hl M,
C,ryplrMol.;! 17, N I I 1111~/mull•·rt•l /I !5 . .1'14 l t·ptnpJ.t,tlll.ll 11 ..! I, 31-\\ ,\I,
( ;1')'/11111> 7 12 I lva(o,poti!(C.I !>tl f.oJII<>J<l,l•lll\ )!12, ,ltll, I/ :!11 ,\J,
(;ylllnllhi.I\UIII ltl•l
(;ymnullult.trt I I\, hCI
I l)'.rllcolmun 9. 9
Hj·I·''•Y•tr•I\X
I"'""'"'''
l
4 r.
llftl, ... .!2, I.!
r l'l~r 1
1
.\I·
,\/,
( •)'TJIIIU\ .!.!H Hy,/llo>Jdf,I(,J ), { .~ lo bn llllll.ltJ \K,! ,\lo
c;>''"'"'"' l;!c,, 12 t:? Hr·"·' 101 LJo!r ,, 'H.:! Mo
I h ,frnu>r.lllmJ 10 lrf IHhc·n.llllll.l 1.:!7 ,\I,
H.r/o.l1.r -11 1 H\llwodJ lll-1 lt hotd.t 11.7, \H2 ,I/,
I IJik.-•ra 1.2 '· -ll7 Hy.lr•>Z<lJ 1\I.Jtl /JIIf,f2(1'1il, 21H ,\{,
lJJ//,•IW•l 14'1 //ymrrr<'imir 3'111 luuulmJ .'l'l.? ,\I,
/IJ//u,,~~~~~a 4 I '. I:: !, 41 1/yomiiJr< 2'1.'\ IJIIIIIIr•ltft.,. l'12 Mt
HJipll<~ 121fT,> I I I >s Hn·''~"w' ,,, <( .lo IJtmrlu l-1'1, 3J'Stl. 11 !1, l'l:? .e_~ \/,
HJh,tnnJ l.!!i Hrr<••1rcr.trro>llls I 1 11• Jt•l lJII~U/.s-17, 7 10, 17:\11'.- 1-1 llil, IX'i, 1\.lo
I I ""''1""'2-1'1, ~ Ill 11Yf'<'Hit'm.r 'I I 4 Ill!> \1,
lm~o:ultiJ I '>X, 177 1!0 i\h<n~l,trop•..L 224 r\rnw r.12-l, 2.2.7, ,,, 1.\'
Ut!fttftl/.t 411, 12 I \/, >/wru/us 11 2J. ,WOfi, -125 Sif,,, ~t12
Lnl!-'1.1hd.i ISI I lu.r/.1• tr< o/ I X, 1J I \'11•/ ~ I I 8, H''
Llll~hlomlel 1 ~2. 17~£r. llil
u,,],r.'!!<-1
""'" 1'''~'''"'" ; 11/ \'•pptmlfr., !'cl
Ht~lrrlllrroll27 ·'"'l•'lltfllltrllf 274
l•npfllot I ~2 \J, 1,. lutoceroJ.< 2 v;, 237 Nt•san.t I t.x. 175, 17 ~
1
.,\
lrthN1d.1 X~ ,\f.,T111>.za.t ; I; :\',,, ,,krlloJ ,~6
Utlwr/1"~":!111, :!11'1 \(,., tc• 2H21T. 9.2i, 'I 2~ ',,f..,frr~.t. 2t>2, '' I 'J
Ullttlit,,l,,~., 11.2, it. J'll M to r.>.ltll)\•11 ~ U. ·117 ,,rwd" 20'1,..! tu, \ Ill
Utwln 2J'), ~ .! J 1/toMIII/r.l lii'i,- 4. ll>l l'uculmd) .:!10
lttUIO<il<.' .::!.)5 lt.//qw" \03, 5.1, 5.2 1\:u.ll X71T
Vl>.•b.l.mtrs8 . .!4, 1.::!1 \lmr.:J~·'IU rllt1 R 2 1\:udthr.ut, lu r ~:!I
L~'"''~'·'f'IIH ; !I, I 11 I 7 Ill I \
L,bt;~l ~52U
\11111(/ .,,,.,-Ill. 12 11
\f,.,,,/,ms 'J.t-. 2711 2H2
·'""'"'"'Ill·' 177. 7 14, I SI
[\;ulludt.tphllhd.! 10 1, IJ4 1
L.m~\tb.mltrt·s b' ~" :l.tttr,tt.tJU'J ·' rr•lrl>tdft 177. 'i

L,.,,,,/,t/,,., I~~~ 1/,,,,,,y.rd/.t Y -19. 31~1
Lvrlrdw f. I 'i I J2, I \Sit IIL'•'P" "'' I I .! ;, W-1 ( lbtlt.! l03 . .'i '· 1114
lophophor.u.t lt7, I H \/,•l•o.~··/1,; 3.2, J . .J. 70 0/•,,f,J/,, IHI
L.•J•It<!J>lr )•llidiwtr > I I I \1\uJtolnp•l' 212 ( lhulh·l.n.a lXI
:!R5
U>f'rlliot \l,>dtt>lll<210, 2 12, R 12, .::!l'i C11~<>111• 177, IK I
/.umrn .:!Ill, 211 \1\udtnmnrphtdJ 210 llcwcor.lUlJ I 07ti
Lucin.acr.l 211 ,\1,,,/,>od I I 111 Ortrll'"' 25'i
Lu.lwt~i,, 11.2l, 8 1/ \1,>/~rt.r 412 Od.rr,,,, 11.1, 12 ..!
Luada.1 2'10 \h•llu•ca hlltf, 1<17{}' r Mt>IH•~~ril'/'"·' 4 1.\, 1.! 3
Lrg.l~z,,,m II h \\1\n(rt 57 Odolllt)pkuwHll \1!2
l )Topor.t b.:! ll<•th•.-iulltlM 427 ( \'1:~"~,,,;, \HO, /I ]0
L)'t•''''" 2-17, X.2'J \1'"'"b'l'~pud, 327tT. H4 ()l,·t~dlu>d.a.\X!I
l)tnn~n.JJ<' 24.::!, .::!H lli•II•'I!IIJ'IIH 127. Ill I'· JJ2 ( Jlr ,, 11111 11 ./ '1, ,\1\Cl
Lyr :or·n,_,r,H 21'1, :'1 I/ \ 1onupiJCupborJ I <)7. 111'1 ( )kmtl.lc \!\2, '\H5tl
\/o"""''•'I'''J 149 ( )knuw· \X2
\l,k'<•')'•' 27J llt>rlllll'tJirtt 'i I>, 132 ()f,>k>rJ,, lil, 1/ 12 11 /I \t.S '\Xil
.ll.z.lrmtr 2:!4 \1nndl\~1tudl<· 132 Olt~rrb \!\2, \X4
,IJ,u.>tllol 212 \l<'r~,lll~ I 1.1 I. 31>5 ( )U{t\( f,h 1.'\5
.\L,rX)'Stell.r )< 14
"""" pm/a I 82 OtU•'Jll•lf'IUJ. Ill 14, \27
,\J.rro•..,rlur.s .:!.SO, 8 10 ~ IJ \lunlu>elltJ .'/ Ill <)n) dwphurJ ll'l
\ 1~ar.r .:! If) \turf:'\:!:! I 01 rl•rm., 12. I 12 I, 4 I \
\ldJl·''lmJ 1s~ .\/yt., lU. •\ 11. :!1(1, :!12 Oph!O•)'told<.'~ .11!11
\l,,~rll,mtoJ l.'i'tlf.- I,~ 2, 17.:!. lll3 !\.l}tlldJ 210 ( )ph111Tt>Hk,l Jto2, 21\!., 2'111
i\I.Likh.tucllJ ! I ~vh nJpo,b '-1'1 o,.1~"'" 11 11 17~>
\hbw•<rJ<.t :\'l7tT 1/yn.Nr,ltrs 27'2 ( )pi'thobr.ln< htlt.l 221
t\.\J!Udll.l 131. 1\\~rtlo.>!d.t 210 Orl•~tlllo•rtlt t 177,- 14 IHI, lli">
Manocon:n' 8 24, 2411 \/ytt/11 l'IH, 21tl, 212. 2l'i,:!20,1\ Ill, ( '""~)'1/olllol y I 'I
Marrdl.1 1111, 12 .:!, 12.:! 8.11 Otltt~IIY•'I\"'' h !, bO
M.umlllht" 11•.1 Orbrtrtllllh'> 'n~tf~ 'J o/~
J\/UbUJHfiS 2 1)7 \',thrurm +10 1 12.1! Ori''"''·'"'" ,, I 14 I I 'i I I!\
AT.tniiiSS<>tti.t -~~~~ ,~,u.drtt"' 2:211 Omt.rtlwf" 11 .~. \~t)
.\l,sww,itl ~() :\,,,,,,,, 'Htl -1111. 1:!.2, ·117 ( ht>llll'l<>l'"' /I /Cl, \h2
,\.f,Jzt>ftydr.r .:!2. I~. 'I •'>'allt<Ill-1 Or~pftrJ<ri11m .101>
,\./e,mrlnlllr '·I i, .>./6 N.1unhd1 23otl On had.• II>] I1!2
.\1t'.mdr.IJh'l4f I ~~1 1\. .1\llllm.L. 230 Onhorl11o.l I Id, I H2
'·'"'''"' 20::!. 2JOff. S.:!IJ, IS.! I, 251 ( )rtftt• IH2
·"'"'"'""" 12'i
,1/t',//umltol .::!-17 \t'b.r/l<r .\'11) ( )rthot a,1, .:!J'i
.\lr,/wimrrs 3 .1, 5X Ncm.tgr.>phruc .Ul ( )fllo(l• <'Filii d./ 2 ';;
,\lt'tk<'ITI•l< .~ 2t> \'nllt~"'l'"'' /017, Ill /X,J:!X.J il, ( )fl}l!l~l·llllll\
11/ I/ , 11) 14
Mq~JtlwnJ." 17~ H-t Orth,lf.tr.>lirr, I I) /1 , /Cl /I•
,\lt/lrt.1 fJ I, 111'(11, r11,
Y :0'1<-og.t,rrnpcxb 224 I )ttlh>lo I•' 1!\2
.\fd,.rdtittus :!7tl, 27.i "((1~/)•//flol(U.I• S.~fo ( ><tlllt~ '"''' .:!1'1, ,\ 211
.11&>1111<$ 270 Ncn~u.uhmtomata 271 o,,,.,.:!lll \ H,.::!.:!O
.\lt~~II>r.uttl >tJ H \, I ill ~·(\WUO!ttll .:!2U n,rr~"''L 1'17, 4ll-, 1.! 1,, 12 -
1\knnu :!7•1 \roprlm.1 I'Ji'. 11.2. 212 O•UCI'I~ 2(U
.llcnrtd/.1 I 1\2 "fOtn~''"'·'llll\, S 111, .::! Ill, :!I ' ( lllo>ll I.! 4, •11 ~
Mcnl\tnnuu 3 1'1 1»8 \'ant.~-126, 12.12 ( ,,.,.,,~)fU.J ... fJ
,\f,,,,""l'"'" t> IZ Ncnuacuhc 22i ( ))'lh lltl'f,L\ 8 ~~~
I'•" Jtmuaa If 111 Plurcrrnmd.1 93 f'r.xluctus 182

fJ.z~•lt•lJKI) f'l!,u.t.~olls Ill. 5.12 Proeuda .'M•
l'a~<·ttad~c IKII J>lulllphHITolcd ll~J. Il-l I f. 1/, 37S,
l'r,>rfuj '~:!
f'~l.tJtll JtC, If to, 1/.:!fl Plulllp•hf J71, 382 Prolttumsrs 24 7 I
P.J.bcdunt•ld~ .:!711 Phtlltp>mella If. 6, .\HO Prolccaniricb 247 f
fJ.JI,sr,'l(.m\ 11 '""" /'ltt</,,domyJ 210 Prota;ococrinus 296, 9 4 l }.
l't!lotrol(!to Ullltlllol 2~h, '1J fllt<<lm :!to. 8.12 ProrostomiJ ()(, I•
l'alat'o()•du, ~I, 1211. 121,5.12 Pl<(llidcrllillll< 9. 9 PfMl'rDCdHit'r()(<'I',1S H. 1 f
l'.tf,,emlirry,m 421o, 12 I l l'hnrwuda 67, 1-14 PrmcrocidaridJc :!7(1 ,.f,
Pot/,(('()/oll~l1flo 127 Phn\p h~cocopUla 41 1) Pr.>tm•odaru 9 11), 'I 11. 2711, :!7'1 I•
I'Jl.an>h<'t<·rt>dunta :! Ill l'hrJWJlOteuthis :!55 Pr<llrrluutr• 372 I<
P~/Jr••/imulou J91 Plty.~·•r""" 4:!tJ. 12.12 Prorocti>w 57 R
l'o~l.rr•>mmt ~\IJ I. If.! l Phyl.<noiJ~IIl.IU 147 Pr..r,>Jrcrt;,in~
J .2 /(
1'~1 '""""''~ 1211 l'h\ Uonnda 34'1. 39'1 P,,uul~rlul.•
1114, i R
l'.lbc\lU"' l<lnt.o 2111 f'/<yl/<~'tfol> 247, 8 29 l'»tmnuhimtl 9 21 R
P.u.ututrMif CJ I() Ph) ll<>araoib ~43. ~-17 f',wJopy~.mt.u '1,/9 R
Ptlfil4tlWII,,,,.,' t l'lly/1.~/tulu, 226, 8 IJI i>sl'ttd<'lnl,'<'riOJ;roiJ'IIO 3:!7 R
f',uaolo•Holllllll ::!1> I l'hyll<•xr·•ptus 3::!~. JU. 1 1 l'!(~rolpl115 327 H
/'ar,,.t,>\I.IIS I I b, 1/.211. J80 P/lyllhi.•Miol:!711 Ptma2ltt,.:!15 R
f'ollot/(/11111> If./ l'!k.w! ~1h l'tmtlin iwll 59, fll H
Pll'•"'I'"I~U.~ll\ 2M I f>llma JQQ Pcrrineo1 289 it
P.mutrr 211 I l'macoccrar:acea 243 PtrmwJ•crtrtt 2 18 R
l'ar.•~tylonurus I I .26, J% Pimw2 10 Pt~rimda 2 10 R
Jl,lrHm••tutl I 0~ fll,sf<•Jinn·a 365 l'rcriomorpha 210 R
P~rv.ut< <lniiJ hO I I 1 11,!~•·•~/yptoJ l 9lJ PtcrobrJochiJ J:!? R
l'aM tchlltJ 127 l'bntJe 57 PteropucL .::!26 R
l'atc/1., 22 I f>lo~MHrn.U 2CJ\) l'ro"foth«a 22h R
l'~ttnll~ l ri l ['/,uy.t~a< 22-1 Pr<>cygoona Jl):! R
P~tmnat3 177, l XI l'latyo.honra 98tT l'r~ow; II.Z.'i, 392 /{.
P.lU<IJd'Uium 422. 12.'1 I'IJcyhrlnunthe-.. 65 Pttln~<rJpOd.11: 32ft R
Pecten 2U'I, 210, 2lt!,!! 1-1 pt,~,..,,lwutt; I fQ 1111/ogrufllll, IIJ 11. '21,, JJ I R
Pttrd~t•IJ'III l'\4 IT, /11 12 l'lc·< todl\cm -122 Prycllll~mmu.• I 1. lli R.
Pelecypnd.a •rt· R•val"'' f>/usMit}'ns I tU Ptyd"'JI·"i" 11 to
l'dt•cvpotltt hnm 126, I!. I i fl/l'lltiJIIMfhU 8 ,5, :!0.\, 2JJ. ~ ,.:; Ptychopantda .'K:!
l'rltiiiii<'I"''J l ~~~. 6.9 J'/t'll!><<lnS •Il l. /2.2 l'tychophyUum 115, ~ IIJ
Pt1111Jl01!1 I :!1·2A 1'1<'•11 .Jumu 270, 277, 'i. I 'I f>u.~llrtX 11!2
Ptflot.<lt'< 271> l'lt•umbr.ulli<~ 5.3 PuhnonJra I'17. 224
l't·lturuuc 11-14 Pkttr•ll)'SIItt.< 9. 45. 3fl4. J 14 Punctmptrtft'r I H~
l'olltl•llltlll 5 J PlrumJirtyum 127. ~ 14 Pswula 7. I I
P~llll.JtUh>C'~ 5'), l!l(o 1'/tun•m r•• 2 I 0 f'stN51J 8.!6
,,,,,,,,,, :!'17, ''"' l'ltllr.lltliiiJil~ 224 p, mo~omd~ .H'I, 1.H
l'cnunwnJ~ l 1'1.::! ,.,,,,.,,,//,, 1-17 Py..:.ol,., 2711 'I ~. IJ !U, 27H
235. I! 2 I
l'tlll~lllt'to)o (MS P.>J<•rlh•>ol 2KU Pyg;stcroJcb 270
Pnrr.mum• l 1-o2. 1X); I 1D<Tilt;,<lml~ llb, !I 19 Pyc~pt 1HJ
/),,,,,,,,.~HI , '1.26 1\'itW<l 2111 Pytgocy<t" 2R7
l 1olllltllllltS Jll-1 . 9-17 l'olyp!Jcophora (AmpluneurJ) IWIT
Per.1c.1ncL 3'1'1 Polym.1 1 B RJdio,(!r.~ptus 327, 3.11. l:\ 1)
P..rtt'·'"" \-IQ, ·11:! I '•>t•llll/oatlllt' S 183 Radtolaria 57
L't!mt hot'< h"""J,.,, 2o111f ronfcrl li5ff [{adiolitri 210, 211<, 11. I I
f'rnsl<lullrtn 1/ 27, .:!47 Jl,>ntr< S. 17 Rofiuesquill<l I Hl
P<'•••mJdf,, •n, 4.1• l'nnKnntddl' 9.40,295, J 14 R<1pludollem.t 1)J, 4.f>, ~.11
Prr,tu• .m' ''l"l l'ornmvace;t 210 RapluophomiAe \KO
l'<lotlioluut<' \71 p, ..,,J.•ma:!IB R.writu 3JbtT, Ill Jol, 111 15
l'tt.<lo.l<mW i :!'15, Y 1\1 Pnurt tlc>td 270 Rayoutllllftol< 2.35
1\ Y""" 41'\ l'n~,ardtodtcb 2111 Rdlidu" tl 4,Jti<l
l>l•·•·•"·'l'"r""'" s 10 l'nnombndtum LO, J R~dlichud~t' ,\1\!l
l'il.l(llltl(\ 8 11. ~ 14 l'r<~r/t.I,J
2'18, 8./0 Rrdoma 210
Ph~cora•< 1 \H::! l'ntpuhda 4 H. 417 RmJo/11/ws Jc.3
Pharor1d.1 16(,, .\1\2 Pnm4hJ'U ]75 Reguluc:~ 95tf
Phacoptna 15:>., \H2 1 11SIIt<~r.1pt115 332
1
'Regt•lJna' :!hQ
l'ltoWI" 4'1 11 I. 357,362,365, // . 7, Probns<JdactyiJ I 115 Rrlti'IUimlld 4.!0, 1.! "
Jlt7, 11 11, J70, J73. JH~. 1/ 1-+. ProduwJa 1!$2 Rt:tnipedi~ 397
\R'i. 1"11 l'rudncodma 7. 11 Rrllt<'J'Itundt~ J!!2
llcpJchm l 42(J \rph,•m,;IIIS, 4.6
Rtrr.•mulr• JJ'a I=; 'ltTophom enau If! 11r
"'I'''•'II••Jmdr.m 11:!, I U. 117, 120. 5.1•. Sll'Qphomen~tb If>" , lbl
Rr/I'Mltllll ::!46 ~ 7, i 1U
Rrtrp••M 15 1 StyiNt•n na 1!1-l
'•phnnop hvrJ l0-1 St\ lunurm JIJ4
RtiiWiiJniJ 1 8:! "•J•Ir.•rl<.>lt.t.l 177, 1111
Rtrrn.to}irJJ>I'" Ill Y
!>rylophora 3119
>;k,,,, l.?.fi, 11.7, 41'J Symmrtr<>r<tpulll! H. /8
Reriolili.l.,,. 3:!8 Hlltf 'iltr,t~f,IJ'III< \28. J.H
RrtztJ 18~ Svn.-Jnda W'l
\~·,•lulr<>> 42ft, 1.? I.? 'iynzrrhm unn.t J?
R/r,Jbd(l()•.-111<5. 7. 114 'ilrmmll<l I 1.24, 3931T
Rhabdt•lltr"'" 149 SyntJ,Q<lpcm 5 1-1, 121111
'mwmo~ r. .2 145tf I :;o SynM<>thyro• ~ I)
RJrabJ,•pl u J HI./IJ, .l:!t•, l~'l Stttlll~odc ,, ft -I , 6 5
lthJbmup vra 3:!7, /0.11 . HI, 3.1'111 s•• lnnyn 2 1(1 TJbuiJco ntd.t 109. IlK, 131'<
Rhti[tlluplryllum 1.:! I, 'i. 12 !'tolcntYtcile 21 n T~bulara 1(1'.1 12-lff
Rf fi<K)' " :!!!7
Rh"'''!" no "" "~·-~
Sllllla~teroadca 2t>.!. :!H!HT
T.1dmtl<•n.~ra 21 7
T.rrmd11mr 1.! 14
Rluzomr,•llirmr42h. 12 11 "'""·r/Jyrlla l !12
R.III:Zt.lJft.•,.,.,, --~5 Tarplryttm.; 2.'5
Spaungo•t:b 270
Rhombd't·r• :U•::!. \Ill 1 ''"•'P'•·rrlla I I 24 .l'U
Sf'"'•"(il"' 9 19 Tr.rllio.:aris I I 2 7
Rlt)'lldtollc/1,, 7. 1.!. 18:! S['lloJ,.,rXt>tlms 382
R.hynchnndhcL I k.:! Tt"o!•'l''lte Jl)(l, 410
SphJ~ronmda 9.-1 > Trlq>lrrlltl Jl!l>
Rh~nchoncllifomwJ 15X, I tlltf Sphacrophth,1lmu' I I .8
RhynchnpordCCJ 17 1 7cllma210, :!I~
Sphin,·w7u.l CJ2, 4 I.? Tdlnuce a ~ 14
Rlryr•:l••"f'"lliJ liS.:! SrwtkllS• tl • r.u 8.2 i
Rolvrota 2:!1 T t•nmoplcurou:b 271>
~plr.tcu1Ju JOC> Tcnt~culwdJc 2.29, I ~2
Rltlrrlr~{rma 1R::! "ipon.fn 7 ~ 17~. I X:! Terat.UI'" Jl!2
RICbd•<•l; nud.t~ 11!:! Spmfen.t~ 1112 Tm·~tatrl/a Ill~
R~lfno .W 1
R.e>ltrocondu~ 200, 221
"l''''"wm 2-IIJ, B.JO rwbmtuln 'l. 7. 11, IH.i
'lpn••~r~plll> /{) I J Tcrcbrmt hd.t 15Q, ll!i
Rct.IJ,!UJI< 2 lt, J.j .1
<;1111\lf'•'fJ J52 Tml>mru/w.r 7 l, 11>.'1 17'J
Rorul.l:!~u
·' I''"'''' 8 .ll, 232. :!55 Tm1lo1 210, :! I H
RugmJ ltl'!tf 'iplfuliMI11tl.l 2541} 1/ l.?
Rk><>plryul 1 3 . .'72tT, /I 1'
Tcu.tbran dUAIJ 2311
SIWio/y/u. -~ /0,21111,2111 Tctr,utmr/1,, I K::!
!'>f""'RI0111CIJ')'htd• IIH TrltJ.~"'J>III$ '27. 10. 11 , 33::!
S.Jb!'/lrdll•'-1 h.:! -'1'''"~•'1'/ry//tdJ 122
~•.,·umJ <; 41,2'17, -l.:!">
Trlt,rrlrytr<lll,r ll-!2
\1'"~'1/WJ Nl , I I Teuthud. t 2'5
'-•rr·~rapll" 10. 1, ·' 1/ltf, J 22 '·lllffi,,_W•l n,.,,~"mOldC'~ 4JO, 12 to/
\,rlmia 271• 1211
\ '(,11/JiJ
111>111111/IS/tr•J I 12
~lportg. '"'"' 1 8 I f. '>t.nrmd.t 1:!1
.\.J/r,,./1., -~~ 77oJttllh'J"'r,r I ~IJ , 124
St.um•ry.•rtro 'l.·H 61,-1 1'i, 12.4
7/t,lll//hlpll ft>ll
,-;,IIC/aloln< 416 'ilmmwro~prrn I 0.11 '27, 331 , .14(1
'i.JJ37 7 /I IQ 71r.umwlobl"'"'' J(\7
Srdlerotd e3 262 17ttntlr.t ll:O
.'\Jrcw.l.t P 7 St<·nurl'IIJ.Jt;tcea 11!2 Th~miciduu lllJ. 11!4
'Vophilt'! 24 7 Stcnolacn ul3 147
S.:~phopuJ.1 1 9'~
Tirt'1 ••>milia I :\2
''' rr.>lo1JIIIJ i I 0 Thccmpi nd.l 182
&}IIZol.>lft ::!K4
S!tr<)dontJ 2711 17tnmoa f; . Jil. 152
S.!u<~plr,>na 182 Stol'"'''dt '"lr.•u 111 !J 'T7t<"l11 2111
\, ·<r.lcttm• 111'1, 1.:!9tT
'>r••lcmotdc.t J::!IS Tit<•/o.urut/1,, -1 /6
'><:lcro\pn~l lllt, ll'ltf 'ii<>III<II<>J.I<ll<t 1>. 10, lo 11 , 1411, l52tf Ttlti,, 8, 18
·'''!''"''''/"' I l '\t~mrcllimo• :no Tommtllr'a J l
.'i..•yrm• ~ 1tl '>II<J<IdJ•III·I 1211
\iur.-llwn .:!-••. 31111, I 1..?11
y.,,httrt.1.• ~4h
'ltrq:>tcl.l<manna 1211 Tormtcs 8.1 l, .:!17
\c)'plrocmllll'> 9..111, :!%
Stnltopo t ·' 124 1o.rbtwlut• I '~
Scypho~o.o I H-IIJ '\tn.Hudl l 11!:!
'-~rMIIIII k~t
Tra.ltmtdl ldlllllt < I/ I~. J7~
<;tntt~•'f<]>lr~ltu
18.' T r,,, hylma I 0-1
'wlrwwur 3~1, ..,,,,,,,,,,,,,..t/,t90 1 rep<htomJU l-1!111'
.'><·lw,•J'<lll• .1-5 /I I I \rr,•trt<llcvy>trrr< .:!R7 'I.JZ, 9,5/ Tr>·w.spi$ I 1 I''· 3b.}
'~·~ 8 .?I, 'n.!, .:!~~ \(l</l/loi/Oft11<ll)) , o//i, ~. 9, 4 /0
.\lrtk,m•m•t217 Tn.mhru1J<' 'H-I
'-ttr<lllt.~toporoid.t 901T Tnarrlmu 11 13 1 1.:! I . 3f•7, '11.:! , 3H4
'iJIIIIII;IrJo,J 1/./ Q, J7CJ, ,'\11;! \rrurnbu~ .:!2-'
~oJrrcyo• /.? 1, 412
T ni•Md11.-lumr htl, l I, .?S7
'itr,ltll(}'f,,, "''"''"' ~711, 9.::! 5 Trrttplr<II<>J~<.•r<~ 9 , 9
Sir/om·/111 l H2 Stro•t>lrt>cltuttt•tr> Ill:! TnJIIttl.t 21"> 217
~t~rnJgrJIIIS 3.?7 '\tr.•rlr<>dNII r lli~
~ini>C)SILI V ·F ,,Ill TnX•'III" 2U~ :!H•
.'>tr••rlromm.t 7. 11 Tnji;01110ld., 210
442 Systema tic index
I
Tnlob1rJ 3-t'l• .\511{ ,\XOtl ( 'mr.tcrinus 297 J l ~tcr~""~·•rc.lror I l-l2. 1!15. - I-
'Tnlobaoi,ka' -till l'rrarm/u• I!12 ll '"'~'"''l'lry//rmr 120
Tnmrrrll.r 17J, 7 14, 17tl, lXI (',.,,., 172 ll ~rllo>llhl 1711
Tnmt•rdhtll17'1, lXI { 'mQ.:!JO. 220 lt:rrll.r -1 12
Tnmenu 11.6, J57, ,,!1:2 Uruomw<: :210 llnrrl>n~r"" 11 !J,YJ0.421
Tnnucleuu J62ff. 11 'J, 11 Ill Umr:mu.1 .H9 111/..•m.r I H:?
Tnmtdr'U> 3S2, 11 111 llm·.r"''"' 11. I:! l
Trh'l'' 641 v.,,-/r,•rrwo~-122 u·... klllmt·n.t 2-th
"I nples~o~ IS:? l'ul.'lll<l.'t'r.Js 227, 8.23
TriJlldtCJ;l!r.Jptu.< 1!8 Vulamrc.r I!12 -'·""''I'Y'•""''J'II
TrcJ<Ih>t)'•lllr• J 11 l'<~ldwioulryn> !SI X1pho•ur.1 '1 1 1
Trt~<lmrur I I 20 I 'ul~J'<'IIgt'Q 4. J.l X•pho,uncU .\-1'1

JrcJ<-JUH J.:!-1 1-';mxi<~-115, 12.4 X y•rridur.J I I li
Trtlt'>l<>alnlb '(17 f 'I'IIIJ \99
Tropl!.\ct:,l "117
I'""''·' hll \ ',.. 1.. ''"''" 1/,, j :! N
Tryh/rdrrtm 1'1'1 Vcner01da 203. 210 \ ',•1/o,r H. I.?, 214
Tryp/<1SIIIol 120 l'nms 210, 8.11. 21-t \ 'vom~olwlu• 25 I A
Tuhip~r.r 5 J, IOH l'mrrflw 8.18. 227 A
Tubnllk,r 121-l l 'mmntlc1nn 227 L.u,urtl~e>itb JHO A
fubuhpor.ua 14'1 l'il/(hnmoiSiff 289 z,,1,Jnmrir.·.r JO<JU. 5.4, Jt'J. 1:20 A
Tu//irnc>rHfrrrm ('Tully ll\<llh!cr<') -12211, Virgdlin~ 328 Zn.uoth,lrJ,l 10711 A
12.9 l'iyella 7.4, ltl3tT. 17'1, IH2 /.c"'l'loyu>< -121>, I 2 /J, 1:!. 14 A.
'/1mi/llc'l :2 l 7 I wip.rrus 2'27 A•
Jimitdlo1 H 1ii I 'o/drovi,, Y.50 A•
Tylc'lldcms '1. 15 l'ollllol 224 A,
IJ•Ic>lllym ,1) A,
A,
Ac
Ac
A,
AI
At
A,
At
At
A,
At
At
A~
A~
AI
AJ.
AL
All
All
Alr
All
All
All
All
All
Ah
Ah
All
An
An
An
An
An
An
An
An
An.
General index
Numb,·" m nahc, r~kr r,~ ll!(urt·,, (S'"' ) 1s .1, ru•,-r~kre11c~ to th~ systematic ind~x
Ahnul21i~
Anal (X) plate 295
At>,vlutc ag.·, (• A\St'mbiJgc ""'"' 2 1
An.u pyr.umd 30 I A,~ou.111on I '
Abund.mce I~ Anal mbc 2'1-1
Abys<.tl I I A•trJeotd 1 I I
Aruptvchm (-i) 2-lSif A,trmhlla<' 'I I
Ahv".li cnn01d, 2'17tr Ant<',tnlb I H
Acanthmryk, I .'Ill Atolh 133
A.ncora 3J7 AtrJdHucnt rh ~n U I
Accc,sory 'uckt:t 173 Ani~ngrapuJ fauna 345
Ac< n·flnn I9 Aul.twphorc 307
Anh<llll]Jnan ::!09 Auludonr l.lmcrn :!ll!l
Acme (pt•ak) tone' 22 AnhpLraclc 305
Amurch• 57 Auloporo1d 12h
Anl..llmed 'uturc' 359 Aulo> 115
Adlmbul.lt ul rl Ill'> ..!tl'l Aundld>5'J
Ad.tpllal :!Id Auntk 2XI
Anugnrmoc rnu,cles I ltl Alllc<ulogy 7
Ad.Jptauon 27 Anu:nn.te 35 I, 3'17
AdJpnvc Llnd•c•p•· 411 Auth•g<'ll<''" -1, 407
Antt'flliUit·, I.Jit Auromululm. \411
Ad.Jpmc rJdlJ!IUII .W, -17 Amcr 1~1,
Ad.Jpuv~ >tc~t~~r·, 31>
Autntht'C.I .l21lf
An tenor .uububcruru (111) ::!7tiif Autotroph I c.
.o\..Wuctorruu\dt• (,«1,.,.) ILl. 205 Antenor Jrdt 354
Adju,tor mu,.J~ (<c.oro} I hI AurotO<><<I.I 1-li.J
Amenur l:>u,h "·'IJ 161 AutolumJ, IIIX
Adoral :!h.\ Ant~n<~r border 352
A•~eulanum (-J) I ~6
Advenunom J\1<'\.!Un 1 147 Antt·nor border furrow J'i:!
.Advcnuuou> lnlle\ 2~:! Ax1al column 115
Amhr.rcorutc -118 Ax1Jl turrow' J521T
Aeon 2.2 c\p•ritc -107
Age 22 Axt21uKrc"''' 117
Arwrnrr.J >Pill"' 319 A '\JJ.l nrj.::ln ..!:'14
Agrr. n.V.I~>~. JS7, 1'1:! Apt•rturc (gmmpod) :!Ctl
Ah<mtJtvplc (llralo IJ I If AxJJ) nn~-t 353
Aphrotd Il-l A.\.lal <Lru<turc 111'1 11 ~
Alar fo"nlt Hl<l Api<Jl ,11" ::!63, 276tf. 9. 19
Alar (A) "·pnun 1119 Axr.tl vnrl<''< 115
Apudcme' 350, 352
Albcrmdr, L.l'. 12·1 Apnphy>~S 274, 281
""'(If wtlrng 8.6
Albnnon, C. C. \ Axon ')4
Append1).\<'' 3<•7
Alimcntaf) proM\)'011 hh Ap,Kiiuc 172
AUd..-,J..!fT U.1b11t, C • 211
Aptychop<td 237 B<tbtock, I E. 107
Allbon, P.A. 15, :!55, 107. 12'i, 1.!,1/ Aprychus (-i} 245fT'
AUnton. W. D. 221 Uack-rcct f.u;ic> /. r.
Augonite -lff, I08. 115, 232 1Jactcri;1 I
Allnmetry 4 ~ Arc h.1ch.1<rcna 55 if O,tkt't, P I!Btl
Allup.ltnr •pcciauun J.j Arducocyatham 18
Alrcmacion of~""''' ltltlll' 102tl
l3JJJ JIIO ;ockctl(lllll' (Fc·hiiH>IdcJ) 266,
Archet\p<'' (oflff 'J4
Alvarcz, l W. 7J, 7711 Archunollu,, 6!1
Alveo)u, 254 Uall'to""' 99
Art·tll.• 9.-1 B.tlwn. W.l C)(,
A1nb1!up1C ICJ4, 1/'l'i An,totlc:'s l.um:m. set• Lantern
Ambmi>2h3 UJ.l,un. I' ':> I 52
Amcuhtmg ha)f:..nng 35.1 H.1mbach, R K 219
Ambuucr.~ :!t..J
Artlcttluing (pleural) facet 35-l Uank U5
:\mbul~"'fl' tube fc, t 21>.\ ""'' ul.non (cnno1d) 297
Arruttom•cal ~unll 252tf B.wu. WC. !52
A,<·nn ISl>. 4.1 13Jrt>~m. M SI
,\mmorutc <eptum 2 \X A>CU\ 1-16
Amoebocvte' !i5
U;~l):hoom. r S ~(,
A•eputc dtpl<'b'T'apnd 33-llf 0Jrl..c:r, M 2~"
Arnpulu105. 2~>5 A>ptdnhnnch jnlls 22-l
Anlclmd7:! BJrr.tn•kJ. J77
A'•cmhlage 13 H•rncr rccf- l.H
444 General index
l:iJmn~on, EJ ..H B1.1d)tehc cvolutmn 47ff ca,t4 (

Barilid, K . 42-1 1:\r-Ancht•l chamber :!311 c~ll'lll fnnlt l:!ll (
0Jul pbte I OH, :!'XI fir.u1~r. M. h~. IJ6 LJutiA JIK (
B.1,;,l (DB) pl.H<'' :!'lltl Brenner. A 299 Cell 'tru<turt• :!•ltT, .!. I (
B~"ctl, :vt. IM Urt'ndtl~\. P U. 15.75 ( dlui.Jr ep•thdtum I hI (
H.ucs, I) L U \JKII Brcn, C E. ;!91, 3!«> vr •I<•• Mandl' <pnheltum (
DlthuN. R l· c 'us Brcvtcone 235 Lt'llll'llUUilll 4 (
fiJth) 1l I Bn~"· D.E.G 3-IR. 397. J99tr. -105. Bl\·3lvn 215 (
H..th)Pd.ltQl' I of .Jto"ff. -124 llr•clunpod• I R5 (
B.ncn. ll I . 2~7 ilnggs.J C 16 ( t'lllrn-tlm-.al piJtl' ~<)I (
D~~er. I" M. In. :!-I~> l:lnttlc->Lm, sl't OphmrcHdel (")\!) LephJIJ.: &mgc> Jl~3lf (
Bcd::!U Urnm1d~ Fom1ation 31JO ( eph.ilic \Uil.lrc' J'i t tl (
'Bdemrutc bJnletid.f,' .h I Brumle)l. R.. G 4::!f>tT C<"phJlon HI 3'i71f (
Bdrmmtc>. (< Brlunnmd 1 I")'' ) Brood-pouchC\ 17'1 Lt•r.mml I 12 (
Odl, B.::vt. \!<et Brouwer. A. 2fo t'crCI .1MI c
lkntl lk·l Brumun. C.H.C 158 C.-n01d Ill tl, I ::!1,, 'i.J (
\ hi, ~otl, IHX, 11 I, I I 7
Bl.'n~•hull, Urutun, f) 165 l'lt.llunn. W ( 7.1 c
Bcmluc mcmbl.lllc' (i'lA) I'll Buc~ .1 rube-reer :?!!5 ChJrnbct' ~7tf. 2 Utf (
Ucmhi, (h<ntlui') 14 Duckt:ridg~. J. .397 Ch.lr.•ctcr d•~pl.occmcnt 4 11tt (
lkmnn , M 20. ~ I 'ill 71, , IH Burld. G. 413 Ch.nwrton . U.I1.E. '7!J c
Bcrgqtmt. P R ~'> llut!dmg 117,13 1 Chd.o(-~) Jl'l') (_
Bcr!!,<tr\1111._1. C.7. 2')11. 171!. 174H iluli.E. E.331 Clll'ltcc rac 144, JKH l
Herry, W 11 N H1 l'lulman, O. M.B. 320tT. .139, 345 Che nucal lo~'il h c
Ho.!J,•r, { ;.J•. H7 B1onyancy 2251f, :?44, 149 Clu·n~ia nA l~un.1 uS, 413, 417 (
·tlJiobne'' 171 Buf'b>e'' ~hale Faun• 41. 1>7. 36H. 41ll/ff l'herm. I 17(• l
lill>ji;CiliC 1'1lVlr0Jlt11Ciit' 17 13urhngton Llmestonc 2')<J C luorulub u•lcr 77 c
RH')!.'·nu. .. du. ~ 1 Uurrowm)l bivalve' 21-llf (. luhd1al pine~ 171 (
UlCIIll'mh I H, 1 1~11 Burrowm~ edun"'id' 267, 27hiT ('hum .\24 c
Btonu" lt 11 Bu<k.G 1511 t hmnuphr"J'IHtlt vahn 15K 17111 L
Btomnc 21 By,...U notch :? 15 L 'llupJt', 1.13K, '77 (.
UonutukLuiJt p~l.oconlnlog< 7 B'"us ::!15 (;hoJm>cytc• (tuUu cell<) HS c
t:l~<"t.rJill-'T phv I•. lOll ( h~,mdmphurr 2119 c
Bllhtntm'""' I ~If ( Jrucone 20::!, 241 Chronto>nllln 2Rif c
D•mrron~<-s 1K, ,,~tr c~ccum(-~c) 171. 140. 35t. t hrunmtr•usnph} 20ft::!~ c
Dtnturb.ltiiHI 1-l C.1lu·olotd cor.illum I 12 C hron•nonc 22 c
HhJtnnt: 21 c~Jnmng lbh C•ruroHllmtrm 21!0 (_
Btr.1mo1h ~rrcnd~g<s 14''· 1:,1 C.Udcr ~. 57 Ctlu I·H,Ihl c.
B"cml 1'15, J::!O C.Wcc 109, 115 C.tlmy lr1(t< 144, 21)2 c
Onh<"l.':l \2Jtf C.illomon.J. 244. 149 C~rcum-ora.l nng 265. 2!!6, 292 c
Billet ~pnu~-. fumt olmn 5•· c~'" 29 1 Cim2Cll c
tllakc. IJ I~."!. 11'1'111 C:un;~rooont lwtcm 2h6 C"'1t',j 370 c
BhKht, A .md W li!'i C•mhnan 'explo~on of ltfe' 'ill. h-Ill Cbde 48 c
Blnc-grt·~n •IJ(.oc' (< \.tnnh." h'n t) I Cuner-.1e (chambers} :no C ladoa 48 c
Hu~n.lnun R '> 14 ' LJntaal dt'posits 237ff Cl~d"m HliT c
l..lod.doc,l. Jllll CJmcr.U ltyuid 235 Cldtlogr•m 1. 1 c
llody c~ viry I lol C~tnpbeU.J. H . 51 C larkc, J .M ::!!!9 c
Budy dwubct :!.:\Cl Cunpbdl, K.S.W . 3fl2. 365 C IJokson , Ii.N.K. 4211', 128, 221o. 31l0ff. G
B<•dy ftiNI b l ';unpb.,U, N .S 70 37'i, J H41T, W<l, -IOH, -1::!4 c.
Uonncr.J. I. :!Ct Larapace 349ff CIJ>~ 10 c.
Boring and ne~tlon)\ b1valw\ 21 H Cardt·lb 145 CJ,,thm 321! c.
Ot"t'llll' fl :?2 C.trdiat.: lobe 3t!tl CIJvubc: 268 c.
Boucot A 1'\, 17,7 1 l'klll C•rdm•l fossula IC).ltl Cloud. I' 55 Co
Dnwen.J .•~.12 C~rdma.l (C) ;eprum IO'J Crud.uia 51:!tl I0::! c.
l:lo\'ll, [) ::!llo C.udm;tl teeth .205 (. 111dnnl 102 c.
I:JrJchtJ 141 C.r.!nuha 172 Co-~d~pt:~twH I 2li c.
tlrJchiJic tul 1111 Cmdotd 3'19 Co2pt.1m·c \rructure' .\1>5 c.
lk•dH.tl (llrllr) pill<~ ~'If 11 c~nnJe 111 ff Ccme'O, A G 121, IJ."!. 137 c:.
Brac h11l v d~ ~ lt~ l C .mu..U pl.tt..., ::!811 <obb. V R Q~ c.
Hnch1dmm Ho I L.upmd, 3U7tf C..:ubb~n. W A 243 Cc
BuchiuJe, .2•)1, 3{)1 C.muthc:n. R.G . 119 Coc IJc , •re CcrNodcnnol (Sy<t ) (\
Br.chwphcon• 1h4, 171 C.mcr, G.S 39 Co<k\, L RM. ~~~. 19).Jflft (",
BrJrhtc•ph"f<' bo~>n I tH Cancr, R. M.::!51l. 2K'I ( ud<ltll l•b, ::!h·l c..
General index 445
Codonutc« f·6tf C11u.,MitT DotTU\t" W\ctulcl 2l-!3fl

Codorno~ytc• 1611. :!h:, Cr.tnotft.j 11 Dtmorphllln (Amm•)noodcJ) 244tT
Coencnthymc •>7. ;!1>~ Crug. G.Y. 15. 176 Dtmyanan 20b
Coenour.: I Ct4 CnnodtUm 352 Ooplcurub J 11
Coeno\teum '11, 1.\ I Cr-Jwtords~..Ue Lmte,rone 2<1<1 Uaplobla~o< 65fr, I 5,1112
Co~nluuon .J' Cnme<., T P 63. 426 Otplognpnd f~unJ .HS
COiled \hdl morpholo~"' .!tltt Cnnoo<W hmestonc~ .}(M) Doplood .}()
Co~n3.:!~ Cm<..-d-bmellar <hell 2ll7 Doplupor.: JO I
Colln ~elL, (choJoW< yt~) ts'i Cnw••ng caJul 320 Dort"<IIV<" mc>cntcn~ 130
CoUccton. (dcmtu• li:t•tkn) If• Cro,>.mg owr 30 Dt~couhl 1 13
CoUtm. V 416tf Cro\\ thcr, P.R.. 326ff Dt<cutJ.J.I enrollment '65
Collum32!1 Crura 174 Dt"cpuncnunum 1()<1
Coloru.l.l n>n.l~ 112tT Cmr.tlium 173 DI\)CpllllCIJ[\ '>5fr. I()C), 115, 326
Colnnoal uut·~ttoun l.\7tl Cryprorhombs 303 Dt1~oconch 221
Cohunella !'15, :!21-o Crysrilline C11ne 351 Dothyrial pnpu!Jrion\ 3}4
Column;~.!, 2'11 Ctcnidial mmcles :?.12 Dovc~oty 14
Conuous,uJ e I bO Ctenophore> Jl)2 Doxon, J t: 133
Common c.n1.ol J I'ltl Lurry. G.B. 158. 17<1, 1!:!4 OobzhJn~ky, T 11 :!off
Comomon1ty c<·ology IJ Cuticle 349fT, 3551r Ondtl,J R. 1'\
Compenm.nx 145 Cuttlc-bonc 252 Dollll!l 35(,
Competitioo1 28, 49ff Cuttldi~h 252 lJon1inanr a.llclc> 3Jtf
Complex cro,<ed-lamellH 1ltcll 201> Cuvtenan organs 286 Dommcrgtte\,J. 248
Compome prisnutic \hell :!07 Cyanobacteria I Douuvan, D .T 226, 289
Compo,.tc tubncJc, JS<o Cylmdrical cordllum 1 12 Donov.111, S. K. 73, 11!4, 289. 299, 326
Compound tnr,tl< 112ff Cyrtocone' 234 Uorsal (pcd.Jde) valve 160
Cmnpound <ye> Jootl, JXH Cysod 143 Dot'io-btcra1 plates 288
Compound plaung :m~ cy,riphragms 149 DoMtlll 2:\Rff. 24 1
Computer modelling 92, .HO 'Cy<t,,id;' 299fT Dott, R 11 27
Conccntr•twn Jcpn\11\ 40'i Double cnrollment 365
Concerted cvoluu(>n J 7 Oactylozoood I 05 Doublure )52
Conch10lin .:!OS D.ilingwater,J.E. 355, 393 Dover. G n
Concreoon\ (, Dm' in, C. 21\fr. 133 Doylc. 1' 253tT
Concurrent rJnge-z.mcs 21 Da\'id>on. T. 187. - - J.)urbam.j W 269
Conne<'UnK nngs 235tf Da,,·km~. R. 26.51 Dumw1 P 32<)
Connccovo: •ururn 3S'J Debrenne, F. 96 Oyrr, 13 P 56tT
Conodontuphon.l' 5K Oechned /0 8 DvsO<iom :!OH
Conodont> '\H Ddthvnum 160. 171 DLlk.J 223
Con.crv~oon dcpout> IO'i Delndu.lplatcs 160,171
Contcmundnt hvpo1tomc -'~'J Dcltid.Jum 160. 171 'l:'.on' 215
Conunenul hdf 14 Deltoid (A) plates 304 EcJy'" (moulong) 1'1, 351. 377ff
Conunenul 'l"P<' 14 Demeru114 Edunopluteul larva 264
Couus 3111 Derrupbtt"S 278 Eckrn,J D 291
Convergent cvoluuon 12, h7tl, .376 Demtpyramids 272 lkologo.-al omhe 15
Conw~y-Morn,, '> 40Kff Dmdnt~e339 l:.cologtd zonanon (cor:ll~) 133fT
Cnok. P M. HI!\ o~ndrotd corals 112, 5.J l::co~~tcm 13
Cooper. R. 33 I. 344 Dental plate< 164 Ecu:oco~hlrar 23:\
Cope,J.C W 26, 42,210 Denticle\ 164 l:.cll><lcm o 6'i, 1021r
Copper, P.l22. 117 Dcntltton 208 bdgn~onc 12!1
Coppic~ 1J5 Dcnton, E.J. 232 EdiacarJ fauna 58fT. 408
Coral reds I'J. IJJtl Dcnvcd charnctcrs 11 tT EJ...m.m. ~. I ll
Cor.Uitc 112 ne~can:n, R . 362 Eldrcdgc, N 11, W, 41:!. 357, 385. 3H7
Cor:illutn II)R Dc,ma Kt:!, 4.6 EIIC~, G.L 344
Cor.~h 102ff Ue~modom 209 ElloptKone 8.21
Core 4ff Dctor.non 227 Enu~. C 177
Coruc.l.l b.tn.UgM .'2-1 Dextoth.-catt' 320 Emkn unl) 16
Cnrt1c ,] fJbm U4 Doachronou~ 20 Endocochktr :!35
Cllrtonl tmu, 322tT Daagenem 4, 407tT EndoconC\ 236, 8 21
Cosue IJU Du.uem 4 E.nJun·clac :?.66
Cosrcll~e 164 Dochograptod iituna 336 l::odod<·rtn 65. IU2ff
C11unter (K) -.cprum IO'J OtChopore 303 Endobun.J.I brachtopods 18f
Coumcr-l~ter.J.I (KL) ~cpnuu Ill' I DJChoromom branching 331 J;nd<lpunl tate >hell 171
CO\'C( pl.ttC> :!tlb, 2'12 Ot~v.:hc cnrtmd> 293 F.ndo"phunculu >tructurr> 235
Cowcn. R l'iCltf, IK4, 2'1h, .ltJ2 [),Juctor mwcle, (md >C:m) I 61 l::ndo,l.dNon 34~
446 General index
Endn,pan"' 171, ~ /1 I ft·nc.,tr:alt' '~PIJ 121< ( ;,.m•linw J~li

EnJo,tuma 3'11 1'<'11<':\tmlt"' ISO (~Cl H. rlun.,: l·un·l· :!I l2
EnJt"ymbJthl~ 5(, hbl"\" h~c~tk·, 11-ltf Gt:llt"r.lU\t~ IOIIC" I h'-l
En~c·,t·r, T, :!..\') hlthr~n~h ~11, 207 l tl'IH:flc n UliC '' I
Enrollmcnt 354, \t.Sff' Fllteref' (,U,pt>ll\1011 fc-.-Jcr>) 1 (jl'lh'~ :!')If ~
l::merun Ill.:! hnnn. S 3.J3 <;,."' lll <t><k 2•111 I-
J:mocod 1 }1.1 h•ha. D. 3'){) Ccn•·uc .Jnh .H I
f·nw,.-pu I "\ll f-"ruhpMe-> 301 Ccruculum H~ I
l!onothcnt :!~ FI'U!;t"ll.l J52 (;,nu~ I app< ndJgcs 3'12 I-
EphH.t lar.> 1117 hxo'<''>lk I SS C"nul opcnlnl-" 3 11<1 I'
Epll.ttm.t I Ill I)(:, Fbl>dlum 3lSH ( ;,.1111.11 phtC> :!1~1 I
Ep•ph~"'" 2ijll rl~dl~te dumber 67 -1 .4 Ccmt.1l pun.' ~h \ I
EpNum.t J•}2 H~l(rllum (-~) 8S (,cnomr 2'1 1-
Fpah.-,a 1mm H~tWOnl\\ 65 Gcnot)pc .111
Epnhdnam I I>X, 7 /11 Flextble rest 270 c;,."'" ( er.t) I 0
Epoch 22 f'ln1>r mthdes 354 ( ;,..,, hwnult>KY 7
Equi.utgul.tr •pan I 1 11111 I lnonng plates :!Ho ( ico, hronul1ll'IT't 7 115 )..,
f'.<JIIthhnllm \pt'Ut'\ 15 I Howcr. R.J:-1 235 (1)uold. I 21t9 H
l:iqauvalvc' :!04 Fluctuaung \ea le' eJ, 74 (, ,ll ,tppt.'nd.tg.-.. W1 11
Era 22 f'!ul(d. E. 98 ( ;lllln.tlll h J(>X 11
I:r.uh.-m 11 F<Jidcd rw:rnbranc st:rllcrurc; 2':11>, :?'J!!ff, C! lllll.m ll'nt' 205 11
Erbt•n. H.J... :nx 312,314 C:all, (i.A. 1.\2 11
Erdtmann, ll.lJ JJI foli acd 'hell 207 (;,11 no!tlll'' 21• I, 2XI H
Enm. G :!~I l-oud !,.'TOO VC 26H (,cll< :!(,\,\SI H
F."·lltdH'IHI:!!111 hwd web In (i•lrm Bro'' n,J.ll 2J2 H
Eub~cren.t ::;:, foot (Molht<ca) IC)f!, ~05tl (,ud<·r )h.1 H
Enklrym.-, 2X. !.1. '>I• rorm f!COCTJ .172 ( il.hdlt h.:!tl, .\~711 11
Euhmcllobr.tll< h f:'ll' 2117 r un11 'PC\.ll~, 37.2 ct.hdl~r tun''"' .1'>.!. 357 ll
F.ul) In lull.' I'\ f<'munon:!O Gl2c"'lcr. M 5tH!, r.5 H
Evaptmtn IX I OrtCY. R..A..J3. 142, .HI. J.Jil. J~H. ,\62. (jn.nhnh.l""' H'l ,\'IH
Evolurc 2.1~. ::>~o ( onl.lnnJl. R 1 11<1, 42<1
J74, '"·
Evolun"ll th<'<ll) loll Fn"tl Ltf!e"--ltren 4U5ii; 411l!tl Cun.1d' 11• I, :!h..' 11
Evuluuuou" c'\Cl!Juon 11 h>">hz~onn I tf. I. I (;un.lrnp..lnJn .'5'1 H
f,olunnnary l.tnn.t, 71 Fn•ter. RJ. :!Ill. -123 (H~UU'hplre':\ ~9:; 11
Evoluno>n.t') r.ttcs .\'I FnuuJn pnn<.:aplc -1 I CunUJ't'rl~ 1 'o H
Evoluuoour. t."nlllllll) [011 Fr:t.ke•. l. A. 7'!_ Cnnu1hna 10-1 H
FH>Illllt>O 11'\ [Tl'llth lh hct.~lymg btv.tlvc> 215 <..ort'J\1, 1 E. b7, ~·1. I\:! H
f:xh.tlem \lphun' 21l he e-h lllf! br;tchwpod.s Ill~ Cur~n1n.m' I h~ H
frn R.V 4~h
E"""'"k Ull
E:<:oc y< he ~hS f range-pat' Jl>3
Could '\j !1, \<1, lh ,~II.:!~7 . .!1X,411J
GrJnt, R 1:. IX7, I'll "'Ho
E"""PI 1 I 11 I l'nngmg reds 133 (ir.lptolttl' IJUJul \t'qurnct' J.J4 H,
Exc"kdcron J4xtt rront.lf wall 1-16 c;r.lpt111Jl" Jh.alt• '11 11<
Exothccal nur~rowrh1 'IIJ
fx,rrt apac al tli" 27h
f.x\t'rt '"PU 1'\tl
rum UllllJimurphulngy 7
IUO)..'l 'i7
huucular tube 1.J3
<ir:ty, l) 110
Gr.I\',J 17. n
Grt'Jt ha.lllu.JI cu1.d 17J, 7.1::!
"'
He
J-1,
Exrensoti>nn /0 /I Fumculu' 143 Ciroml·d 1ntl1 27' lit
Exrenwr mu,dn .15 I Funnd-buaklmg (Ed uunad.-.a) 21>~1T c:rnrllll~lr,.J, 1'. 61 Hc
El\tt·nr.tlut~JIIId Ill. I h7 f-umr>h. H 240 Croup 20 !I•
EM111Lll011 ·11-ilf. 7 Ill han.rch, FT.IS (irowth linl'' 11,11, 2114 JJ,
Exrr:uemacubrbuddtnf• I 11 Fu;ellar UNtt' 31 R. 324 ( iu.trd (ht•lc·JliiHC t·s) 'S-1 li e
Exu vtJt' .I77 UlllllTrt''I-MJrw,J 211
Ey~'
Ceplulopt>d.a.!\1111
(,all'. A Hl·t. lll<J
t:Jllacma. I. 286 11.1.1\, W llo'i
"'
Ht
li e
Tnlt>btr.t 36, .JI• . .\52, \toOII (;;llloway. CB. :\(1 H.~tltl 11 'fl
C.uuete \OIT H.t<•lkl·l . P. -17 He
FJuJI <lltlll<, 15:!11 C•rnvJernc 1·1 l hc·n1JI '"'rrrn :'ll4 He
b<'l~\ 21 c~rc21-l l hllnn, A. ll 1, 17 21•. ~:!. 71, 1~7 Ht
Fanuh 10 G~T\Uill(. w. 2:!..1. \<J7 1-fZIIIIII~IIII \V \11~. J/h lie
f'a<etml.ttl' 112, l.:!lo l •.arwor>d, F. J 1'14 I hmmunJ,I.., 1~6 He
Fa\Oolt• :!I .X, :'X~ l•.l>tTOZOOtd 1115 ll.lnt..cn, N 11.1 \'1-lfi H"
f'1un.U pr<wtncei ll,tl, 3~5 Gcc.ll 65. J11 I Um1" hrl, W ~21• Ho
Fcm.R 4\,Jto\ Gcn.J! .m)llc 352 lh)'luulll Ho
FeU. If 0 2'111 C: .. nal 'P'"" 363 I hrdttn>und' 21 ~ Hu
General index 447
HHdy-WctnbcrJ.t t'4UJ!I<ln H llu!(h"'· C.P 3!16 hour~ ,n.u1 ~it'~
llarLmd, W 0 211 Hugh<">. N 1- 17 hopygom .11•7
HJrp~r. n AT Ill, h, 7.,, I''' Hum.1npor.·s 303 h" mvc: nc•luuun 247
H.trpcr. E 211~. :\l"i, 251o I lum;~urhoml" 303
1-larper.J.C. 4:!1> llumb<lldt <urrem lh J.ocl"'" 1>.1:
llarmun, \V I) 117, X'l I htnlo~ptlkr, T. 37 jKk\On,J U (. -J<J, 121. 15'i
Ha\\ km,, H L 2n HwL<rud-chidC:r 102. :!35. .!Jtl, 255. J11 l,,on, T '> ~-o. ;!HO
H•)"'ard. I' J I:;·' 373. 4.:!0tT J twh, I 211, .11,
Hcad-fout .:>:!~ Hun.J.E. 334 jJm,-.., I' 'llo
lleckd. P. 'J<J Hu'k>.J. :!6 ).1"> (Ccph.llopod•) 245
Hecto<OI'ilu, 2J2, :!"il Hurg.:ns, C 362 Jdkm·, , R P 'i :!t •'I, 21 !1, Jti8tT
Hedhng. Hll .:!I I lydnophoroid 21i Jdi,J ~. I IK
Hcdgpcth,J \V .:!4 HyJrJnth tn-1 Jdl. P .A 1.\t;, 351>
I ldicJl cnihnfl 20hT. 2:!1 H ydrotd-\Crpulod symb10'1> HIS jclhll,h, .<rr '>qphllWJ ('iy\t
Hrnm~. W. ll llvdroporc 2!!6. 30 I ]t'll\1'11, ~ \7~
HcmHng~nll.lw, G .177. 31:\.1 llvdrospiraculum 305 Jcp,l'n . C.L. ~:!
Hcnry,J.-L. JhSfl; 377 llytltnspirt•s 304, 9.47 J<'Trl'. I I07
l lennJphrndllt' 22J Hydrost.ltlc 'kclcron 66. 384 Jcr prnpul'""' :!:!'l
Hem1.1typ•r n•r.1h I lttr Hydrorbcca 10+ Johmu11 J (; IX, 217
HetcmchronNI\ hunwomurphy IItH llvnlithclnunthid~ 61 Joh"'""· R .l,.; 405
H eterochrony 41 fl llyuhthid< 61 Jomtcc.l Jppcndagn .\47
Hnen>dom d.-nu tu m '11"> 'lOH Hnwrchnt 7.11 j <lllC\, j. S 229
Ht•tt'llllllUIT'h<::!Jil ll ypo11ome 230 IOY\l'~. K A 107
Hcrcropvgom 367 H}pmwm.u sulurt:s J5'J jugu111 15. 175
licu:rmtcle 3!17 Hvpo,tonw 352 jull, R K 117
HetetUzygulc 1J ]umpmg !;l'"'' (trJn<postltl\) .\~
He\mt. R. .:>.H. l·U l chnolojzy 7
Htll, 0. 1:!.:! lmpendent hypo<tome J5'1 K.11U/.IV.~ . K. 2X5
Hmp;e J11d .!nl<ltllllPII hnplm10o .:!2CJ. 2451f KJutnunn, R Ji15
Or.~c hmpmh I hlltl lmpunctarc !>hell 171 Kcc:I2JX
Ill\·~ h-e, .:>t hiT lnd1v"mJ 114 Ktdnl t•rth :!HO
Hmge pbtc :!115 ll1JU,lrlalmd.lmsm J(, K"U~. ~. 21'1
n,pi.."'· R. G I 11, I nc4u•l~tcr.1l 15!5 KcnncJ, W J. :!4 I
HtmJnnln gi.1L!1Uull 1'1.1 11\7 lni.IUIU 151f Kcmu, l. K A 2R I
Ho.M-\\' 51 lnt~unal ''phon-reedm!). 21_l 1\cr.lt:Aw,~ '12
Hoffitun, A ~(. lntT-.,b.lS.ll (188) pbtM .:!')3 Kc·,hn~ot. R. 10.1
Hob'Jm J7'J hllrlrJJt.J IRR) plate' 2'.14 Ktonmpbncn J'l:!
HoiiJuJ. CH 7~. l..lO, :!.Brl, 2."i l lntundohultfunn t2bubc 126 K•,·r, P..'\1 :!h'ltl
Hnlm. (, .\'12 ln~ham,JK. 14:! !<.or.. N \.\11
Holrncr. l.E 17711 lnhJknt \lphun' 213 1\.Jtthcr,!) :!i
Holo.:hrnal C)l' l(,otl lnhrntJJKt' 2tllf Ktdbvog-W Jcnn~. 1.. J'l7
Hol..,tonuwu• 22h lruu.tl bud .1211 Klov.111, J f 12\tJ'
Holorype 'I Ink \JC 232 KnoB, A.H. 'ilt, 511, hi
Homcobox.'H ln\t'n .lpteal dose 276 KtKh. l> L 2l!7
llomcmldndtum 17'1 lntt•r.unbuJacrum (-a) 2/lJ, 27 11 Knz\low,ko. R J2tJtl, .129, .\J'I
Ht>nteonmrphy I~~, I hX lntcrarca I &4, 172 Krc:lh, 'W 12211
Homcouc gt·n•·, JH Internal mnuiJ 4fT
Homogeneou< <hdllllh hn<·rplcut.t l furrows 354 LJ U.1ebcrJ, M. ll!l!
I t.~m\Jiugoth thHllll<""" '"' IH lmcrradJalsurure '1. JJ 1.1hi.1 l palp' :!tJ'i
Hnmoluh'Y Ill lmerthecal ;cptum I 0. I I Jhn11 11 '''"
Humuzyj.,'<.Hl'> 1.:!11 Interval 70IIC:\ 22 L.ldd. 11.~ 112
Hnod (Naurilu') 2:10 lmt·rvJllum !B LJ~.l.IIJ, R . 155
liun~ ed Utt'lJ<' Hh lntcrzootdJI alflculariJ 147
'Hupdi.1l muu,tt•r' 4(1
'·""·"'l· 111 :mr
lnnJtcntJcul~r budding 1.11 I nllp-\ht•ll,. '<'I' HDdunpodJ (~:>"1.)
HonzomaJ collccttnK bowl .:!Hh, .:!'1'1 Jntrmor.ton J35 l.on.;,·r pbtc·, .111°1
Horizontal \tipcs (CM<'Il,llnnn) /1/. X lnvolmc.:>JK I .tndnun "' , .\0
Horotch,- c\ oluunn I 7 lndmm bvcr 77 bu.-, P J) 177
Hof'eo.hoe .:rJb, >If lmoulus (~\,!.) lroiL\tune• .1 l.m~t. \V () 1 I X
Horvath, G. Jh:! lrrc~'\lbr cthmoi.h :lf,7tf I anrcm :!lo.l, :!SI•tf
I lorweU, Lnnl"\tune 227 lrrt'~"'l.tr g.Amopods :?J<) L.lppct' 2.\tl
Hnu,X HJ lsoJont :!OR I 11Jcw•. J I 22'1
HOU\C,M.:!H hol.u.- th,·uc .lJIJ l.1r.~on, K 22'1
HudK>tl,JO li'Jtf hollrwn HIT l.lrwt>e..i,lol'. 7;\ I-lXII
448 General index
Lncr.Jl border lillnlW JS2 M .m d.- ov•ry Ho I, I '17 Mol~.:ubr ~cntto" .\7
Latcr:ll rn< rc~'" 117 t.hntle ep1thelium lh1 :vlllllbm hl\'~IVC'< 21 Q, If I 'i
Lucral rccth 205 Mantle tiNon 212 J\.1l>II~X~>n IlK
l:ue" r~rl•cJnon 1t.t, Manton. '>.M 341<tT Mon~rJ 57

Ltw,on.J n 1941 Manubnum 103 M<moh.t.,.l 277
L~yttun, M .~ X Mu~ual plm.-s :!8::1 Mull<K)th( ttlllnld' 2•Htf
Lebrun. P J'i-1 .\h.r~Qrul 'llturt"'i 35'1 Monotv.apn.l fnma 34 ~

Lcft valve (BI\•alvta) 21•4 MJrgtnJrium 116 Monomprun 20'1
Lch1mnn U 2-1;; M.u-guill, L. 57 Monuphyly 12
L.-nholf. H.M. U2 M:mup1ac 281 MnllUtulc' 12X, 152
Lcnnon. C D :!17 Mmill, D. 23-1, 407 Morpholo~ ll
Lt"ucon lrlul(\~nll!7, -1 I M.c.\lvc corals I 12. 12h MoN J W M
Lcnntou, J 2h ~tthCW>, ~.C. 61 M<>rtcrr•l'n, T 2to'l
lcV1-~cm ll 35-1. \h.! MoL-cwcll, W. D. 7h Mm11tl, liT, lh'i
Lrbcros~ml~ 120 IllS M•"'·eU, W G H . 133 Mnulttnl:! (ct:d}~") ,,S liT, l771T
lihrigen~ \=t2 MAyrurd-Smn:h.J. 2h Mu1r-Wood, f1 17'J
Lrcan, C.R. 51• M.l)T.£. 26 Mullr•, K J. 41 'n
ltg-dniCIIl 20:ill Mazon Creek Famu -122tr Multljol\'lll'f;unil or• 17
L1~ment plt 2ll5 Mcandrn1d 128ff MulurdtltOlll graptl1lold'-' n.
J.j(l
l iljedahl , L. 2 tl)tf Med1an sumre 359 MutJ.I pore> 12b
lmdstrilrn. c; \'i2 Medma 51:), 102fT Mtl\<":\lln~. l. 1.32
LmdstrOm, M 10k Mega-evolution 45 Mu,tk platlortll\ 171
LinnJcm, C.G I! Mcg.~>dcre> 94 Mu>clc lt:Jrs 162
Lm.\lc), R M. 202 Me10bcnrho; 68 Mu"·ulo-cpothcli.ll .:ells IIJ2ff
l1rhofinnon 2 Me''"" 3illf. :!.J Mut.~tum 11fT
Luhosrr.mgrJphy 2lltl Mdvtll~. R. 269 Myophorc IB
Lmoral14 Member20 Myoltr~cum :!07
'l.tV\111! fo"t1\' 4H, 1711 Mcndcl, C... 2!S

Llantlowry br.tcluupod ~"t·mhl•t:~ 11N Mcr;c,pod 37!:i Na" t•um \hrll 2117
Lob•te thcoe .I \ll Ml"t:nrhvme R7 NJUnt hvpt"h'lll<' .\''i'l
lob~ (.unmonuc ~mur.ll) :!42 Me.cnren~ I02tf N~thol'\l, AL. 372
Lobopoth 412 M~odcrm 65, 262 l\o.Jtur.ll >election 281f
Lodde\', M 1J. 1~t! M~odcmul ~kdcron 26::! "Juphm luv.t .\117, I:! -
Lo..r.m;hllll< ~p1r.U l 11, 20 I M('>Og_IOt'd 102iT '\lelr.oc (ncl.tnn) I 4
Long~c<>nto 235 fo, k>OWIII2 392 NcmJ31h
Lophoph.,rt· h7, 1 n, 151!, 17\11 Mc-,ozooecu 14'.1 "'rn1.atocv•t IO:!fl
lucuto1d X I I .\.1ccachronal rh)'thm 372 N~:n-n.rrwuu\111 27tf
Lunam JS(), 6 12 !1-\t-rapanAn >Unrro; 359 Neoteny 41
Lung-honk' 351 Met~pta 108 Ncurypc'l
L\mule .::!09, 2toK Mct.mcula 31 i! NcphnJ1Uill I hi
Mcta,onu 392 Ncntr. I I
McAle~ter, L 212 Mt·t.J.>toma 3q~ NculfiJII, U I:! I
M<Ghe• G 7\ 7<o, .NI! Mcwheca 31') Nlci\C!I, ( (,7
McKcrrow, " · l'i,1 7, ll!li,144,JKto Meuzo:ms 58ff Nttt'ci..I,M 422
McKmm·y F 41 411, I !;5 Mer~oritc impact 73fT, 251 Ncwell, N [) IX
McLJu~hlin, P 3'17 Mrcroconch1 2-14 Ntchob. D. 2lH, \ I '
Mcleod, N 77 Minocvoluoon 38ff Nnn-mnplm shell 163
M cMt'namut, M. Jnd IJ 57, t.O, M M!Cropygous 367 Nnh lt hynum If..!
McNam.m, K.J. 41, 4Cltl 177, 31'111 M tcro,cJeres 94 Nudds.J 117
Macntwudt> 244 M1ddlenms. FA 17 Nudibr~nch> (sca-~lugs)
2.!4
Macmnulufi<Jn NIT, 4 I Mdlcr, A.K.. 8.14 Nuntrntal wxonomy HilT
Macropkur~l 'P"'l'• \7'l /I ltl Mlllcr,J. 356, 3(>2
Macrop) go us 31• 7 MJ.lsnm. C. 297, 331 Obar. R A 56
Macula (-c) .352 Mmt'rJ!ized <kdcrom 65 O'Uncn, (., D 1) 1!
Mo~curdJ D B .\llh Mmton. R.P. 21!l ( lrnp11.1l furrow 352
Madrcpnnie 26.\ :vliomend. 367 On 1p11.oiJJn): .\'i2
Makow>ll , 11. 2-1-1 Ml•..arzhev,k\·. V. f>2 { ktop1. H< C rph.tlnp<•d.. (~"'t )
M;~l\lm>-KJtfn' pruvmc,· .\Hio Mnchell. C. H. 31!1, 3~t.. 333 Otul.ar pl.ai<'' :!1'>\
.\.IJmelnn 911. 9 4, 27' MitOSh Jt.lff, 2 2 n. uiJr pure\ 2113
Mandlblt· .\411. 3 117 Model waptotitci :laOtf 0.-ub · nd~<" \57
Manmn~;. P. W Jn MoJc- of evoluoon 38fT O.:ubr 'uturc .\h i
Mandt' lhlfi: 1'1711 Modes ot incre.1.•e (corJl>) 117 O'Dor, R K. ::!.H
Mm tic onAI' I hI Molecubr dnv.: 37 l1hvcr, W.l 1.!(1, U7
General index 449
Orunutidi.t Jhll PAUl' 12 I'LlyiMd, G. 12JIT
OntO!It:ll} 17'1 l'c~l (acme} zone~ 22 Plc<tnlupltc 175
OptTculum 112. 145, 22\, \~8 l't·arly tnutiJu,. se< Nautt.lus Plrmtroprc l(t'll<'' JO
Optk. A . .\Sh Pntnubratu:h 224 l'lcrupcduncu!Jrc I 1'6
OpL<thO!-'Yr.U ;!(1 1/
Pccururhomb> .\03, 9.46 l'l<"ur.U (antcuhnng) l>•·ct J5.J
C)pa•thopman 'utur~ \5CI l'~eto.:~ulus 329 Pleurallll'ld JS4
Opa•thmonu .\Kh, \'J2 l'cdaceUande 266 Pleural turrow 35-1
Opponuni,u, '!'<"< •<-. I'\\ PrJtcle lhO, 171 l'lcuron 354
Order 10 Pc:dtdc· tonmen 1611 l'locoid 1211
Org:meU..-.. ::?s .! I, 51J Peruclc opening 1no. I i I Pndt~ (tuhc-fl·o:t) 2Cd, ::!7h
Organ> of I'And~r lh 7 l'erude (vrnrr.U} v..J\'c 160 f'<>dnmcrc' 3tlH
Ont.:m oflife ;5 PC'd J S 417 l'ogonophor~n\ Sf!
Orl~w,ki.S 174.11,1) PciJgtC H l'o;cu,J ,,Cl
,21
Onn"ton, A IX7 l't·ndcnr Ill. H Pul~ gctH'' \0
'Onten' 41l!ff Pentwtubct-am 58 l'olymtnd 166
Orthoconc' l.l~ Pc:rmmeral symmetry 21l21T, J IJ l'olymnrpht'm I::?, U7
Q,culum K~ Pt·rJmorphom 41 Polyp 11\10"
(),good. R .C l72tf Perched 1-:tuna.' 48 Polvp.try (rlubdmotn<') 3!9
o,phrad'~ 2::?.\ Pt·rforaced <epw JJ2 Pnlyphyly 12
O~<acle' 2'11 l'<'ndenn 311!, 322fT l'olyptdc I .JC.
O,ti~ RS Pt·ng~mhic gtrdle 2M. 280 Pulyplmcly 3.\
OvKdh 145 Pen haemal <y<tem 264 Pol y>t\llltod~,·.tl I J 1, 1.17
0\•tpomor :\92 Pertud 2::? l'"phn. C. 424
Owcn,A W 17~ Pen-oral (POO) plate• 30 I Pore-catul' Vilt
Owem. R .M \(,2 l'cn-or.~l nng 286 Pore- p.ur. 211.\, 27()
Oxy,one 241 I'COll\traCaJ bnt\h 161! Pure 'lnl< tu re\ ('cy\tOtd., ') 303
Oy'>ter<. srt•l )mcan.a (SY'f) Pemmr.onmt 168, 2tl5 Por<K}'tC\ 1!5
Pt>npho:r.U mcrea,e 118 l'opnv, LE 177
P.l<hydom 20'! l'cnpheral uobtes 34 f'o,t-.thd•>mcn 3'12
Pacl..;.rd. A. .25.:! l'cnpor<" 301 Pcht-.od.tpt.ttion -!()
P~c:domorpho " ~I , Jl,.\ Pcnpnxt 21.>3 l'c"tn l-16
P.U~eobwgengr•ph\ 7 l't·marc 1114 i'o,rcnor horcicr 1''>.:!
P.lbeobtulugy 1.' l'trutome ::!llJ Prc-•btlnnll·n .\'12
P.&l~t'ocommumue' 1.\, I KR Pemtorm..J plate' 281). 21:1'1 l'rc·~.,,I.Jpucmn 46
Pal.lcoccologv 7, IJ, 111-l Pcnnun n·d-complex 18. 191 l'n·dmnn 25511"
Pabeohrtt'md<•nt 211'1 Pcmun~r.tlaunon 5 PnmJn'<nn,um...-, 1o
'PuJeunrl'l"~ .&I borunu\' 4•"• Pcr-rJdW -ururc 263 Pnnun· nutt·h 31'1
P&eontulotpc-.1 rcb\ ::!~K ('et.11utd ;unbul•na ::!.n''ff Pnn1.1n produ,c<·n 1h
l'abcou.xodom 211il Petcf\cn, C.. C.J. 15 Pnmary ,hclll.tyrr I t•K
Po~bcutt'ntp<'r.mtrc 25~ Pho~<elmd I 1::?. 5..1 l'nnubrac h' 9. Ill
P•L'I<"c>wogn,~'T:lphy 7 l'ha•du, IJrvJ ~79 Pnnnttvt• ch~r.lct<·n 11 ti
Pab 130 l'h<"nen.- I 11f Probmm (Gamopnda) 223
l'.liintrope l (,0 Pht'tH>type 30 !'roeh o~ m tiC 2.\il
Pallt.tllmc ::?115 l'htlop. C..M. 281 Pmc)jnl' ~.12
PJUi.U •inu' .2115 l'hosphariunon I• • .J07ff Progt·n~"s ~I
l'almcr. AJ, . .2.:!. J7CJ PhragtliOClllte 240. :?54 Prok.tryotc• 55
Pahncr, T M 'JK Phyletic gradu:llt~m 39£1' l'r0-0\tr.JCUnt ;!S.j
PJ.!pebr.tl 'l'i.!, \61 l'hyllodcs '2.71) PmpJt t .t 177
PALSTAI 1J l'hylogcnetic <y•temacic' 1 l ff Prop;artJn Mttnrc, 352, 3'i 7
Pmdenan upcnm~" lli7 Phylu~cny 11 Proso:pta IOIJ
Papcr-naunltl\ '"' Argun.outJ Ph}lum (- a) 1(1 l'rosicub J 1!1
1'.1pui.Je 2illJ l'tckmll, R. 13, 427 l'rtNtt-CyrJJ 209
P•rabl.t>tOJd' ,l 1-l l'uucocyt<" f\5 Prm<UIIJ .11'11!, .1'12
Paracnnoul.s .11 I PmJmdcnn 85 Prou'P" 377ft
PJrag:tstl'r 1!5 Pmn;~t<' 331 Prntc:~-:ulunt lltll, 179
Par•ph\h 12 Pmnulc\ 291 l'mthcc1 J 1'1
Paraporua 60 P!Jm<pir~l 202 PmtnbratKh ~~~~ 207
Par.~typ.,9
Pl.mkn,· (pl.mktonj H l'rotnnmch :?ol
Par.uoam RS Planu-.:onvc>. shell I 64 Proto<orallttc 117
P~mudal buddmg 117 l'bn\IU 104 Protclcll,t.l 57
Pa.rvlCO\tdLc - 5 Pbnubtt> ::!41 Proto,cpu 108
Patdl.tte I 12 PI oltron 268 Protozoam 57. 115
Paul. C.R C .t2. 7ft, .lolll, .3111tf P!Jt\ cone 202 Proxmul <'nJ J.Hif, Ill 1.!
450 General index
P,~uJu.lduJtun• 7 ll R.lut-t<'ll (k\JC·on) !S7, 4 1 \c.•-<·UL uml>l'f\, •c r llnlnthun"de• (Sy>t.) sr

P<cud<1"'mkr .\h-1 Rhr, •philic ;?Q8 '\c·J lrvd,lws~:t·7:?. ~p
P,cudnonl<."r.art"~ 17'1 PJ1cnphnhi, 2'18 '•·•-J•tn,, ''' l'l"nn.aru!Jn·J ("V'I J Sp

Pwud<•pt><ILI 57 Rhunp<'dL:ncuiJre IRS .,,•• _,h'b"· •re NudahrJnclu~ (S\>r) '>p
p,~udopunnar<" •h<ll 17 ' RhnJJ~. I) C h4 \cJ-urdun,, iff EdunotdC2 (\v,t.) Sp
P,cud,mal->crd~ J'it. Radurd,on E.S. -toe. St nn,!.rv ,Jac·lll.•r•-r ll•'i Sp
Ptt·rnp<~<I:!.!S RachJrd'<>n .1 R. 11'1-. -l:?-1 \t umlahr.ad1' 'I lli \p
PC)< huloplu· 17'> RarlwrJ, R.B 3211 32f>,.UI,JJ-l,J31>tf, s,·abdtn, A :;•ltf, :!h'ltl, 2'~ I, '\lj(l, '7.~tf. 'ip
Punna< 171 J-11 ~115tl. <l:?l·tl Sp
Puncm•rcd cqmlabnl ~Uti R1der.j. 150 Scldrn, P .•W-1 :...r.
Pupafimu 8 I'' Radang. R. lOS 'tdt"diVt' 3<1\ dlll.l!(l' \\If ~p
P\.~>tJnun 35 I R.adlcy J\.1 12. 2o Sdt'lll'""" 221• Sp
I') mm,ul , urJIJum 11.! Rtgby.J.K. lB. 9!l!f s,·r~>albc -'·'" ~r
Rtgbv. 'i . \:l9£f. 331, J-11. \.D St·pt..u,l.t J 1•'1. 71 Sp
R1ght valve (B.-.th ia) :!04 O..,<'ptJ 'I~ llllltl" 11 ~.::!tn '\p
Rt!Q.."· t.A. Hl7 Septal llullllp: 242tl "P
QuadnrJtiLuc .1.!7 Rob"on. R. 376. 3HO. 411• '>cpt..sl m·, k 2.'12 'r
Quamun• cn•lutu•n 40 Rnlfr. W.D.I. 349 ScpuhrJmla ~·11' 2117 Sq
Qua\1-IU(.,., 1l 1117
RaJ ben, M 1:. '17

RnnK,J -Y. 75, 193
ltoniCWICZ, I' , 11 15
R.(lse•tkrantz, D. :NO
'it:rll''2l
Scrp<'amcont 11.::!5
S..v.l\l<lfltdu, G. 29.1
""
'><j
... t;
RaJ.\ch. G.O. Wl Rustral pl.tle 359 'it•xu.al thmurpl111n1 U. :!4~ Sr

RJh,um. I \71, R<l~tr~l <Utur~ 3'i'l 'ilwds.m, I' M . 7~ 'it.
ltddacbo,·uf. P. I'll Rostrum 172 Sheer f.1hrit~ 324
'Rac1.~1 <CnNCCII(t•" ::!.~11 Rc•wt', A.W. 282 Shl'J,inn. PR 21•, -1~. J!\0, ·'ll~tT St
Radnl (Rit) phtc' .2 11I tl -.h,··~"l.l, 1. M 1011 St
RadtJI WJte• VI'\\<"I' 211'\ .2'1:!
Rtmdl, A 1i:>3
Rudt,uJ b"•lves l•J. :w~ !:>hsdl>. K 1117
....,
..,,
H.JthJn.al CIL'\) plJt•· .2'•4 Rudwtck, Mj.~. \'I, lf!7ff. 17Jt1, l!H. Sablang 'I'"'"'' I~
R~d•nrncmc d.atmg h, ::!.\ 1<)2 <;,.ul• 'Ill ~{I
R•dul• :!::!~. ::!4'i Rumu:~ar. 0. :?!In. 210. 2.:!1 'tadt pl.att"' \(15 Sto
Rad\.an•kt, 1\, /I I ' R y.m. I' n u St.bt.t I. 5, ~07tf '\u
lulL R.A h7 Rvbnd.J.!:> I·Ptf. lS:!JI 'ttlvcr,L 7J \tc
Rmlfhe I~(, \unna, M. 7h, 2'1Jtf, 2'll•, '00 St<
RAnhl'utltllll, V.' H (' fl lr. SJddle l•mur.th 24::! \1111nm·tu,/\ -ICII- Stc
RAm,knld, I 41 \ ~.a.l.un tY I Sff. 407 "nnpk pn,nUtK 'hcU 207 '>tc
Range zone• ,!n ~Jnd~~oUJr. :!Mti Sunpk thc<~t' 3.\5, /0 11. 111 14 \((
Stc
Rat~ 01 c~oluuou :Ntf, 47
R~ur. n M .:>u::!, .2 n
\.uu:.an tul><" 265
\.mnd.-1"\, P T. 51
Sunr"'ll ( •. G 2t•. W
\uup"•ll '> l:!h
,,,
RA\\\Uil, I' I' :?~ Saundel', W.B 230 "•rhooul cJn.•l 2.\2 ...{.
RJvmond, I' E Jrt7 SJV.UZl. E. :!O::!, 213 S1phorul ruhc 2.\~ 'ir.
R.lzor <hdl, ~c.: .'>.•it "· l.wu (:'' •I.) St.dantonn view 332 "tpfHHUhtt1111.1tt'll' 2_2h Sn
R~. Jf'lnii.Hil\11 17 \l' aUnp <r•· Pt·.:tcn (Sv" ) '•rhonuwuu.l 137 '\u
Rece"t''' .tilde, Htl '>candcnt 320, IU.I.'? ~aphon' :!OO 'iu
R.cclmcd IU .•~ Sch.ifer. W. 1::!. 4211 \aphllllt'k 2J() Sto
R~t:l>mbmation 1::! Schmdel, [), 42 '"tcr j.:rc>liJ" I I Sr<
Reay>t.•llazJtlllll HT 'Khmuewolf, 0. 2511 '-,kchon. P W .!h 42.21 7, 2'ill St<
Renun.lr~;>n.IIL" I 117 S•ruzo•hroalt-y<·s 36. 46, 352. 362£1 Skcvmgton. D 343 Stt
'Red Qut•cn' hvpothcm H 'irl112ndont~OS '>kmll<'r. H.J 15 Stt
Reef 17tT, I~-\ I Uti I~<., Scluwlophe 174 '-ht-b.uuJ 2:>' Stc
Reb'Tldl (, 111 Sdmullt\ ass, H. 35 5 'llugo;. rrl' ( ;.a,tmp.,,b (Sy,t.) SI<
Rci(Ul.1r C< h1noad, 211otl Schopf.j.W. 5(, '\IIJJ II ~ht>lly IO\\ill b lH . 71 Sto
Rcid, R .E 11 H7tl <), hnm, F.R. 398, -!~2fT ~mnh. A.ll. I ~. 2t>-'lf. .!6<J, .!7(1, :!!ill ~[l
Retf. w ::!o. :nx "rhulz. P. 73 "!l!h, I \till Su

RCJllYflll'~tCIIt{', ,~ O..,dente> 2::15. 3511 Stull>. n·(' l..altr<'podJ (Sv,t ) ~~~
Rrp!J.c,·muu ~ '>rlemhlJ~t' X7 Sot ~~·r- I I•~, 2H'i Stt
Rcprodu,·t•vc 1\ol.mon .\41[ 'iloffin, T.P 99, 1:!7, l.H ..,.,hi N r . .2 11• Su
Rq>t.•nl 1211 'it<'I~((IIJ 112 s,,Jatsrv «>rah lllllf Sn
R .-"htun 21 X ~corpaoru. 3~S Solnl ofcn Luuntone ~'17, -!2<1R Su
R,·,orptwn lllr.uncn .\1 11 C.R.. 17, 193. J81J
'>•·nt~~c, ~<•rauf, J.E 111, l:?~tl Sn
Rc<parawry tree• ::!!;h "nobacuiJr tuherd<" 27 .t 'p.ad" 2.'2 Sn
Retrudtt\JIIIll< .2 1i'IIT S<natttm. C 1 1115. Htt!. l26tl. 1.3bfi !:>p<'tll'> h, btf 'iu
RctroHr.tun ,\.lh Scvph~>tnmJ 1117 Spt'< 1c' ~dc~u<m 4-1 \u
Rhllxlowmc ] 1'1 Sca-JJJemone<, set H yJruzna (S) 't ) <.,pntlit llllll<"'.llf Su
General index 451
SpcnnJtoplwr~ 2J::! '-luhblll~.:!.:! ltlllth-<hdl,, ,,.,. ..,, •Ph••r••d.t (Sy<t.)
Spncr. S ,17'l•T "'udbury. M 3J6 Tui'\IOtl (CJ>tropu<l.t) 22J
Sphacrn.->tll' 211 :O.uku, If~ 'Tnwl', I' M \21, \5~
Sphcndt3 ;!hiJ \up~rphhmt 1-J) HI ruhnu!Jc HS, I H
\phcwiJ.I.} ~nruUrn,•nt '\115 \u perpo,tuon .!11 J r.tlC-flh'll, (tdtuuhl,_,,J,) h. •Cl off
Spt.:uk" R5, SKit, IllS ..,upnr.tdul (SSR pbtcs)2'J-I 1 r.ll h~.l<" JSI
Spuade 1114 "'UrJn.tl pl11c TC> r r.Jll,lllll) 1'~1->~·huttl ,\7:-.
\ptrah~ I x:., - -, -.1.: ~u,pcn,lun tCecicf' 1l• 'I""' m, N I! 4:!~
4
Sptrolnphc.· 175, • I.! \uturJI ltt!;l<· 1114 1 n.lllb'U)JtC the<-:It' ,Hh
Sptcldtue-. N /11,7 \utur,·luw 2.\0, .\.'\1 rnhJ-..11277
Spo>n,Jvlnmt 17'\ :O.t>nlf<'> :!JII, 2-11 tf. 35 I I nlt•hn•· t'H"' ~lo, -\to, '1>11!1
.Spun~·· retl' 'I'll 'I\\ .tUtmTI'\ (dt•po ' I nluhll<' Ill'\~· (Lillwlu•) 3'111
Spou~n <>5. S'itf \\\ olll, A '1:!, ~4!j Tnplobl.\<11< ~~~. l.'
Spnn~r-•pt' uk• )oliltT \\\cJ,,h Upper Cambnau -115 Tnv1~l ('I'"''"') tWllt' 'I
'>pnnt-.'ln!6 Sw1mmm~ bt nl vc~ :! 17 1 rw lutunn .'( 111
Sprcrtc -1211 Svwn RI•. -1 I I rudttnd nlfaUum I 12
Spnnklc.J 2r.J. \tKI \\tllpamt -I<J I rmholnpht· 171
S4uamul.tc 121, S)mplc<tomorphy 11 Trnt hnphw .. l.orv.t 117, l·H, 22J
S<llllll,, <a l'tplo.tlnpod;t ('I) •I) '>yn,tpolllorph} 11 rmph~< lbtl
Squtrc<. J) I· I \', :'!yu.tplluJ iae DO I n•t•man. A. b. 250
Stabthzmg ,,•)c, 1'"" J!'> \ym•cnlt>gy 7 Tuhc-tn·t (pnd1.1) 2f•.\ 271,
Stdgf' 22 S)1trlubdmomcs ]-\5 luh~r.lt• Hlo
Stdtllt"). '>M 2htl. 11, IX , 70, lol.!, II..!U, .,y,tc.·m 22 lurhu1~tc 112
243 ... VIl<'lll,lllnl!tf I urbn.l U.tllll 22'1
\wuon. R.J I> Typ< 'fll'<ml~n' lJ
Starfi,h. '" A'l '""'''·' (Sv'1 l I'JhcUac 115
'Swrft,lt Lwc.h' 1110 Ttbul.ot"91, 109,114. 12-l Uh•gh<. (; .lll'ltl
Suo,tJcal mcthNh 12 I Jl•u l.mum ICJ') Umhli1CU• 22h. :!4(1
Stc.tm. C. W '11 tT, Ill'\ I ldtYtchc: Cl'ltlunon 47 Umbu(•IIC<) I 1~1. I x~. 211+
'itt'~m,, \.{ • \to I ~lco!Jt" I t.-1, 171 Umlfr.<ond. C .\.12
'>tc.·de-Pt•trm tf, 1-1, 17h T~phonomy 2. 14 Umr.uno1h tpp<:ndJ~.:' J-I'J
Stegtdturn I i 2 Tuch P. 2'15 Urm,oht· >hell• 200
Stcll{<'r. T 'lk I a.x. I• Vrhau,·k. A Jll\. .\2~. \\7
Sten<, ~.2u J ...,,,..t,mt 21ll! Uuc.t >IJt<' ,\ 70
..,tcnoh.ilinc 15 l .L"\OilOtllY lofT U-wb·· -1.!4
Stc111cl, 11.13 21.5 I ~\lnr,J l> X.~
Stephen<M, n (. lX';, .!'.10 l•vlur I'D 152tf Yl.<'lllln<",j.W 2h
Stere.. m Do r C'~·th H..t. 2os V.tlvf~ I ~X 2tl5
\t<·reuwm· 11 r; TC~ll<ll 2q:. v~n !ten. I! 1117
'ltcmueJ~II 1 ('tdlrrt, 74 c. V.111 Valcn , L 4-1
\ope J~rt 1 dopodtt<' JI>H V.lnl'' (!-., .tpltlhtt·) l.\<1
\urud<~nt lmt<'tn ~)>Ill TcJ,on :\Hfl V.tr.tn~un ,.:l.an.lnnn ~7. ~H, fd , 72
Sttn.J 3Cd, Ph Tmtpnnlp••lymorpht<m IO:!ff Vm.lllon28tl
Smckmn. \V I•~ I <"lll:lrllhtuh 229, -122 Vane<'\ ::!25
C.,toddard, D ( I }2 I er~ll<' J'iO V.lllgh.ul, A I h
Stol.c.·'· R 21q T <'M'Jce radgc' .\5(, Velot:t't Lu·v• .:!2.\, .'1 17
'>wlon '\~2 TN (cooona) 21>J Velu111 ~2.1
StolothccJ 32 1 T•·tr.ob,,.,,1 277 Vcndmo,tns f,Otf, 7 1
Stnrnod;leuon )117 f l'lr .IX Oil !:IX v~nu·• :!JH
Stnnt cnl.ll 2<• 'i f'hJIIIII NCroid ll.j Vt·ntr;t l (1wdoci<-) v.olvc I f•ll
Sturm<"r. L. J5h,JIJJ. \h7 \'14 I hwn. C h'l. 176. 185 Vcnm.'IJ. (;, 44, 25511
Strati~r~phy 211 I hCLt(-··) 2Xh. 318 Vcnm·ufi•nn ).;J,ttnpllll, :!:!X
<,lrttkbt·l);t'r, R 2h 1 h<cl.ct 1.\5 Vcwor.J. 1:. N u 11r
<,mJ~berg '> 2l7. 2-l I Thom.1<, A 1 . 377, JRI! 'Vcnd,r.w' (ophturood) .!'Ill
Smmplc, II.L 2S7 ThtHIII'· R.n K. :!0. 213 Vcrttlal hltram•n IJII 2 1)1\
!'>twbiJJtwn I117 I h"'HJ'••.m. J'A. W. l'l. 2(11 Vcrtr• .tlpllt<" I Id
"itmmat<lhte' I >l, ".7 l'homp•lUI, I -122 Vc'" ubr "'"~J'liiiCIIh IJ2
Stmph1r 'hell 11):\ Thurox ~.11 Vrbr...·ulum (·l) 1-11,
Srubbkfidd, C.J WJ TilOf\nu, C. I h Vtl' tnou' lVI(UI.tn• I-ll•
Stumt~r. W 1112 .!V> .:!'ltl \~11. ~~tllf l~<•gkr, D. 355 \'u.l.1l. C !'H
">ub-;uul Ll\unle .:!l•>i 'T•·~~nng-fork' evolutoun2~>! Vmtul.tr furruw Jh'i
..,ub-pet.tlt•td l(,'l r\ltnllllliUil 58 \'r~.l.lu21\'J
~ub'J'<'Cl~ 111 I llllllllO!lld\ t•l Vr~dl.t.\11<
452 General index
Vugul<l319 Westennann, G. 233ff, 243, 251 'World-m\Llnt' (chmnozone) 22

Vtrgular tabnc 324 Westrop, S. 73, 376fT Wnght, A. 182
Vi~er.lll't3<< 198 Whelk. su Buccmum (Syst.) Wnght,J 273
Vivtanue t071T Whmaker, R.H. 57 Wnght, S. 40
Vogd. ~ <>b Whittmgton, H.B. 73. 354, 359, 365.
Vo~buqth. f 132 367,377,408 X-ndtography 421 ff
Whor1201
Wade. M 59fT, 107 Wiedmann,J. 250 Yancey, T E. 218
Walcott, C. D. 4<~1 Wign.all, P. 300 Yodtd<on, E. 70, 200,210, 226
Walker, K.R. 124 Wilbur, K. 206 Yong<'. C .M IJ2£1
Walo<sd.., (). 4201T Wtlby, P. R. 407 Yudomtan 63
Waiter, M.R. . S7 Williams, A. 158, 168
Ward, P 230fT, 251 Williams, H. 332 Zangerl, R 424
Water va\Cubr 'Y'tem 262 Wtlliams, L.A. 424 Zhang, X.-G. 360
Watel"''ton, C D J92tT, \(/(, Wllliamson, 0.1. 67 Zhunvlev, Yu A. 88. 99ff
Wat:.on,JE 11!7 Williamson,P.42,229 Ziegler, A.M 15, 188
Wax-modelling 322 Willmer,P. 12,64,66 Ztg-zJg ··nmmmurc' 168
Webers, G.r. 260 Wills, L.J. 392 Zoanum 143fT
Webster, C.D 299 Wilmot, N. 355ff Zone 20ff
Weedon, M 228 Wilson,J.B. 108 foss•h 20fT
Wclls,J.W. 120, 132. 136 Wiman, C. 320fT Zonedunt 143fT
Wells, M . j. 233, 2'i 1 Wood. E.M.R. 345 Zooid 143, 320
Wendt,J 115 Wood. R. 92ff, 99 ZooxantheU.te 121, 13lff,215
Wcst,IUUS Woods, H. 8.10 Zygolnphe 17'i
Westcrglrd, A. 31!4 Woodward, H.B. 349

Invertebrate Paleontology and Evolution. Fourth Edition (Clarkson, 1998)

Uploaded by

Document Information

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Invertebrate Paleontology and Evolution. Fourth Edition (Clarkson, 1998)

Uploaded by

Copyright:

Available Formats

Ill 11

9 780632 052387 >

Part One General Palaeontological Concepts 1

2 Evolution and the fossil r ecord 26

Microev lution 39 Part

2.4 omperition and it eflects ~9

3 Major events in the history of life 55 4

tn1ctural characteristics 301 11

U c of model in interpreting the mode of life of grapt loids 34il

12 Exceptional faunas; ichnology 406

12.7 Ichnol gy 426

PC or MA , minuuum 6Mb RAM, D-R M dnve, 640 X 480 colour monitor.

Ftgurc 1.2 The various subdisciplines of palaeontology.

F~~ cold tempera te

made up entirely of sedtmcnt-tr;lppmg and po ,,bly l1 Cllll.: ,

lime-secreting cy<mClb,1ctcna: the prokaryotic tro- t••WJtds i1

i~ the rhiMIH'Imia 'm.!JIIf,lf,, zone . ·a eh

re ci nal biostratib'T:lphic unit Stoge• Age htl-C:renr F.

1tr.1t1graphy A stage, on the other hand, b a group

Evo ufon and the fossil 1! e~ not pc

pr vided J ~tarting point r r later developments. \trun~ H gc

ula.r characters. ( teanlS, 1977). Furthermore, the concept of t:h redundant,

C;lse~ tllu~tr:~tin g various pattern' of microevnllltiCln- I hcrt• an

cx:~mples of long-tcml sta'h and of the Otlgll1 ol tllu~ \VCtc i

::111d that speciation w3s rare and probably raptd. Uu ll~tigr.tphy.

~ren,Jii7~:d (~:.~;. in food prl-fen·nt:cs) tt may increase orgJtusms. may n>

Evolutulnary COIIVl'rgcnce m.1y bt• mtcrpreted .1~ .1

w.1s so 1111portant, for rhcy respond ed d11Tercntl> tt cvl"nl. Cc

tht<rc.lfter. Hl're, of courc;e, dinc·ct competl tion fi • tl ll' tllech

"·,_ Some of the earlv protem-hkc sub~tances proba-

1:3 soh-bodied tnloblte

mnelids and some pccuhar orgJnisms of unkno\\ n Pennotuloceons

(h) Chancel/aria (j) Yochelcionella

(I) Cambroc/avus (n) Canopoconus (o) orthtd brachiopod

Precarnbrian trace fossi l s

The e.1rlicst appearance of rr.1ce fossils may help to

(c) mon ancestry m 'mega-~roup~· or \uperphyla'. "1rd

ism\. R .uher more cnvi,agc some kmd of rree-b ''1wn

brate~. the vanoll\ phyla or ~uperphyb bem~ deri\ '" be

from a prot070,l n precursor) Js comidered to hav bvbri\

Tnlobite' i, for example, whteh have been t•xn In

morpholo gy of their eyes and other envirorun c Tht·

arthropo d groups. The term 'prim.inv c', bO\.WH le-. c.

succc~<;fu llife forms of their time. Ill cl

originate , brc:tking away from chc fimcttonaJ \}'t I 1tt

archetype s, which were likt·wi~c continin~ p.llll

mollu~cs, marttcul.lte brachwpod~ and others that

Agmata (Yochclson, llJ77, l9H3). Salterclla is wide- 'ncs ofcxti

Extinctio n s Possible c auses o f m ass extinctions

UniverstC) Prl.'~'• NcwhJvt•n. (S) mbtotK ong;m Awr,uml.., ').11

.\I 0 '1979) Th,· (. .unbn.ln r tdiatmn ~ Vl'nt, 111

J'i tht• Choanoftagel11da art· m form \'Jrtu,tUy Jdt'IHI-

~I uprmng, but thl.' v.t~l lllJJority of ~ponges are of

nbrr of ~ylOn-hkc: dcm~:nts opc:n into the large

JllllUh m~y hvt• 111 the p.u.tg.tstc:r. Jnd ~uch assona-

PHYlUM PORIFERA (l .1111.-Rn.) MultllellulJ r anunal~ ''

(c) Demosponge spicules

(a) S/phon/11 (b) Uthlatld desmas

(e) Peronk»ll11, with spicules

ogmnd ,,, 1. hhd} rd.1t1.'d tn ,cJcru~ponge'. Strornatoporoids

f-gwe 4.8 (o) Stromoloporo (Dev.), showmg lominoe in o froc-

tnl- diffm~nt!ltcd \tntCflll'l' i' often lmt and only J rctic-

Figvre 4.12 A generalized sphinctozoon sponge, shoW1"'8

li'gure 4.13 External appearance of on unusual twinned

ltm,, (Order Chetlo~cum.HJ, "iuborder A~co