93 Crassulaceae r- subfadistribution ::" s p p . tribe m i l y family taxon clade 5 Slnocrassu/a t - l, Meterostachys , f f;mnerate .r ä Orostachyssubs. Append. I ) Hytotetephium Rhodiola Umbilicus Pseudosedum I Rhodiota Phedimus rytvum nrl, - Aeonium ES t I temoerate atiu 1e Medit') i f ) Europe/N.East S. assyrlacum [G] S. mooneyifG] NE Africa Petrosedum Eurooe/Medit. S. ser. Caerulea lGl S. ser. Monanthoidea [G] Near East lll t--l I INorthAfrica ) Monanthes 1e S. magel/ense [G] rP ic ae S. dasyphyllum [G] S. tydium lGl -{ Europe/ Mediterranean/ Near EasV (CentralAsia) l Rosularia 'Leuco- rd liu- Macaronesia ] sedum S. sedoides [G] S. hispanicum [G] S. commixtum lGl Prometheum S. gracle [G] Sedella ES nle ra d 't_ re n, Dudleya -'t )F NorthAmerica Asia Europe Europe S. div. spp. [S] (Asia) S. div. spp. [S] (Europe) S. sexangu/are[S] S. fannosum [S] a Villadia Lenophyllum S. fernafum [S] S. hemsleyanum [S] )r e- North America S. fuffuraceum lSl S. sect. Pachysedum [S] Graptopetalum \r"n",,,," Thompsonella n,o,o Echeveria -,| Pachyphytum Kalanchoe *** Adromischus Kalanchoe Tylecodon Crassula Cotyledon Crassu/a E] l-tl.l l'ltl l=l tll'l l"lI tl l"ll"l tl I l,ll"l tll"lll tl S. obcordafum [S] 3S !l'l tsll'l ll S. oaxacanum lSl 9S tl l.l I S. ser. Macaronesica [S] -|Macaronesia tn E_l lo I t \ Aeonium to I N ; tEI Aichryson .)) r- lslf t!_i Sempervivum S. ser. Pubescens [G] F l'l F]l"lI EIl=l I,l l"l EI to I l Kungia ll 5 t \ | (South)Africa ü )' (South)Africa \ (n I l'l tl l,l l"l lo I !l"l t_lL] FI lAl I E] t-öt td I I!',_JL_l Crassulaceae reli' t r-t )or rtt). {t0 ons ,g a r and )g)', sed ned rted -lus, :hin allv also )s. rbg. th e rd es ;sile 1t0 in ric a Iu m ade rlar rdly :eds 320 r20 109 roots); Ieavesusuallyterete-subulate, * conspic- Byrnesia Rose( 1922). uouslyspurredat base;inflorescences t elongate TacitusMoran (1974). thyrsoids, often spike- or raceme-like, with Herbs (sect.Graptopetalum) to subshrubs(sect. 6-70(-150)cincinnoidpartial inflorescences with (broadly) Byrnesia);Ieavesobovateto spathulate, 1-5(-8) flowers;sepals(almost) free, (always?) usuallymucronatein sect.Graptopetalum; usually spurred at base; petals connate at base, lobes + glaucous-pruinose;inflorescencesthyrsoids spreadingto reflexedor erect,whitish to pink or with few to many cincinnoid partial infloresreddish;stylodiat recurved;fruits erect.n = 9-17, cences,or pleiochasiawith few cincinnoidpartial 20-22,33andhigher.About21 species in southern inflorescences; flowers(4)5(-10)-merous, stamens USA (south-westernTexas),Mexico, Guatemala diplostemonous, rarely haplostemonous; sepals (BajaYerapaz), Peru,at (600-)1,500-4,000 m. appressed, free base to and + equal in size; Divided into seveninformal speciesgroups petals slightly connate at base, spreading stellately, (Thiedein Eggli 2003).Appearsto be closestto basicallywhitishor yellowish(to greenish),usually Sedumsect.Fruticisedum(IJhl and Moran, l.c.). with reddish to brown cross-bandmarkings or Täxaformerly classifiedin sect.Altamiranoa are now placedin Sedum(Moran 1996;Thiede and blotches becoming denser towards tips, rarely uniformly coloured; stamens first erect, after 't Hart 1999). anther dehiscencespreadingand the antesepalous stamensrecurvedbetweenthe petals,after 24. LenophyllumRose anthesisbecoming erect again; carpels shortly Lenophyllum Rose, Smithsonian Misc. Collect. 47:159 connateat base;stylodianormally abruptlyoffset; (190a);Moran, Haseltonia2:l-19 (1994),rev. fruits ascendingto erect;seedsusuallyreticulate. Herbs,rootsfibrousor thickened;leavesdecussate n = 30-32, 34, 35, 62, 64, 66, 68, +93, +170, in few basalpairs, elliptic, roundish or rhombic; +175, 192, +204, +208, +244, +270. Eighteen inflorescencesthyrsoids with several cincinni, species,USA (central and southern Arizona), narrow thyrsoidsof compactcincinni or reduced Mexico (widespreadfrom Sonoraand Chihuahua to racemesor spikesaboveor throughout;sepals to Oaxaca);rocky places,to 2,400m. Divided into sect. Byrnesiaand sect. Graperector ascending, nearlyequal,* aslong asopen corolla;petalsin upperhalf spreadingto recurved, topetalum(incl. Tacitus).Accordingto molecular (ob)lanceolate, yellow(ish); stamens slightly data,Graptopetalum andits sectionsarenot monophyletic, and Tacitus,Cremnophila(= Sedum) exserted; nectary scales subquadrate;carpels and selectedspeciesof Sedumand Echeveriaare erect,narrog taperinginto slenderstylodia;fruits amongits species(Acevedoet al. l.c.). erect.n -- 22,32,33,44.Sevenspecies, from USA interspersed (southernTexas)and north-easternMexico. 26. Thompsonella Britton & Rose Genera25-28 torbg. t al. uhl : al., tml ned :ent ous Ling ;eo:ent Stemsat leastbasallywoody,but many taxa with sessilerosettes;leavesusuallythick and strongly succulent, in t distinctrosettes;inflorescences lateral;petalsat leastbasallyconnate.x - 30-34with secondaryreductions.Centredin Mexico. According to molecular data (Mort et al. 2001),these genera form a distinct American clade(--'Echeveriagroup'),togetherwith Sedum sect. Pachysedum.The latter shares the above characters, exceptfor its choripetalousflowers. 25, GraptopetalumRose GraptopetalumRose,Contr. U.S.Natl Herb. 13'296(1911); Acevedoet al.,Brittonia 56:185-194(2004),morph. phylog.; Acevedoet al., Amer. |. Bot. 91:1099-1104(2004),mol. phylog. Thompsonel/aBritton & Rose, Contr. U.S. Natl Herb. 12.391(1909);Moran, Cact.Succ.I. (U.S.)64:37-44(1992), synopsis. Glabrousherbs or subshrubs;leavesin rosettes, flattish, semi-amplexicaul, often glaucous,margins straight or undulate; inflorescenceserect narrow thyrsoidsor spicateaboveor as a whole, with 10-70, 1-12-floweredcincinni; flowers (almost) sessile;sepalsfree, (sub)equal,clavate; petalsshortly connateat base,imbricatein bud, spreadingfrom middle, outer face pale, inner face * dark purplish-red;nectary scalesminute; carpelsshortly stipitate,shortly connateat base; stylodia slender; fruits erect; seeds reticulate with irregularlongitudinalrows.n = 26,52. Six species,centraland southernMexico;usuallyon limestone. 110 |. Thiede and U. Eggli Genera27 + 28 Petalsupright and connate for most of their length; sepalsoften strongly unequal in size; petals thickfleshy; anthers (light) yellow; fruits divergent. 27, EcheveriaDC. EcheveriaDC., Prodr. Syst. Regni Veg. 3:401 (1828); (1972),rev. Walther,Echeveria Rose( 1903). Oliverella UrbiniaBritton& Rose(1903). Rose( 1905). Oliveranthus Glabrous to hirsute herbs to subshrubs; stem none or tall, branching or not; leaves rarely scattered along the stems, usually (ob)lanceolate and mucronate, often glaucousor highly coloured; inflorescences, racemose, paniculate, or rarely spicate thyrsoids, or cymose with one to several cincinni; pedicels usually with one to several minute bracteoles;sepalsreflexed to appressedbut usually somewhat expanding, almost completely free, equal to strongly unequal; corolla cylindrical to pentagonal to urceolate; petals imbricate (valvate in Ser. Valvatae), white through yellow and orange to red, rarely green(ish), inner surface usually with nectar-cavity at base; stamens 10, 5 attached at top of nectar-cavities, 5 at top of corolla tube between petals; carpels connate at base, erect at anthesis; stylodia slender; fruits widely divergent follicles; seeds reticulate or smooth. n = L2-34, polyploid from 28-25A.About 139 species,centred in (southern) Mexico, also southern USA (Texas) and Central and South America (Venezuela, Colombia, Ecuador, Peru, Bolivia, northern Argentina). Divided into 17series(Kimnach in Eggli 2003). 28. PachyphytumLink, Klotzsch & Otto (sect.Pachyphytum),* oblong to oblanceolate, white to pink, rarely orangeto red(dish),inner face in upper part often with red blotch in sect. Pachyphyt um, Iaterallynear base with appendages which form two free,1-2 mm largescalesbeneath filaments; antetepalousfilaments connate with corolla,antesepalous ones (almost)free; nectary scalesoblong, * yellowish(-white);carpelserect offset at anthesis,f free;stylodiainconspicuously to abruptly narrowing; fruits usually divergent follicles;seedsfairly smooth.n - 3l-33,62,64,66, 96,99,+I24, +128,+160,+186.Fifteenspecies, in easterncentralMexico,(600-)I,200-2,500m. Divided into sectionsDiotostemonandPachyphytum(Thiedein Eggli2003).Themorphologyof the petal scaleswas studiedby Leinfellner(195a); similar scalesalso occur in some Echeveriaspecies.Pachyphytum maybe nestedwithin Echeveria (Thiedein Eggli and closestto its sect.Urceolatae 2003). KaTRNcnoIDEAE A. Berger(1930). II. SuspA,r,r. Shoots* woody*; tissues(always?)with crystal sand*;petalsconnateto strongiydevelopedcorolla tubex;antherswith terminal,f sphericalconnective appendage*;seedswith few (4-6) costae*in sideview,coronate* . x - 9*. Genera29-32. Note that Kalanchoideaeare morphologically highly derived,althoughcladisticallythey are the secondcladebranchingoff from the remainderof the family. 29. AdromischusLem. Adromischus Lem., lard. Fleur. misc. 2:58-59 (1852); Pilbeamet al.,Adromischus(I998), synopsis. Pachyphytum Link, Klotzsch& Otto, Allg. Gartenzeitung Shrubletsto * 20 cm; stemsfleshy-woody;leaves (1841). 9:9-10 Subshrubs; stems first erect, with age usually decumbent to pendent, not or few-branched; leaves obovate, spathulate, elliptic-oblong or lanceolate, usually very thick, usually conspicuously glaucous-farinose; inflorescences almost always simple cincinni, first drooping,later * erect; floral bracts 5-9 mm in sect. Diotostemon or usually 13-30 mm in sect. Pachypltytum; sepals erect, appressed,almost equal (sect.Diotostemon)or (often strongly) unequal (sect. Pachyphytum); petals erect (sect.Diotostemon)or spreadingto divaricate flat to almost terete,glabrousor glandular-hairy, erect often with thick wax bloom; inflorescences spike-likethyrsoids or spikeswithout terminal flower, to 55 cm, with few to numerous, 1-5flowered dichasia;flowers usually erect, rarely pendulous (A. phillippsiae (Marloth) Poelln.); corolla usuallylong and narrow; petalswhite to pink to red, rarelybright orange,Iobesat sinuses joined by thin membrane; filaments slightly exserted or included, papillate where connate with corollatube;carpelselongate;fruits follicles, (always?) dehiscing completely along ventral Crassulaceae lte, ner 3ct. 8es ath 'ith ary 'ect [set ent 66, ies, lry Yof ;a); petria Sgli !0). stal olla rec'in ally the rof t52); tves iry, rect inal -5rely n.); :to rSES htly rate :les, tral epidermal cells with sinuate anticlinal walls, with bladder-cellsidioblasts'usuoften (always?) inflorescences ally completelydrought-deciduous; Fig.28 thyrsoidswith oneto severaldichasia;petalswhite, 30. Kalanchoe Adans. greenish,yellowishor mauve,rarely reddish;filaKalanchoe Adans., Fam. Pl. 2:248 (1763); Hamet, Bull. mentsusuallyexserted,hairy whereconnatewith Herb. Boiss.II,7:870-900(1907)& 8:17-48(1908);Hamet corolla tube; fruits follicles, dehiscing apically & Lapostolle,Genre Kalanchoeau |ardin Botanique "Les only; seedswith irregularcostae.Forty-sixspecies, Cödres"(I96a); Raadts,Willdenowia 8:101-157(1977), rev. winter-rainfall regions of Namibia and South E. Afr.; Fernandes,Bol. Soc. Brot. II, 53:325-442(1980)' Africa, mainly SucculentKaroo. Growing season African taxa; Boiteau & Allorge-Boiteau,Kalanchoe de autumnto earlysummer;floweringt in summer. Madagascar (1995); Gehrig et al., Pl. Sci. 160:827-835 Sister to Cotyledon,accordingto molecular (2001),mol. phylog. data (Mort et al. 2001).The two generahavein Bryophyllun Salisbury( I 805). commonbasallyhairy fi.laments. suture. n = 9. About 28 species,Namibia, South Africa (especiallySucculentand Little Karoo). Kitchingia Baker ( I 881). Shrubletsto shrubs,rarelyrosulateor smalltrees, or biennial to annual; leavesusually decussate, 32. CotyledonL. * flat, Cotyledon rarelyalternate,verticillateor subrosulate, & Koutnik, L., Sp.Pl.:429(1753);van Jaarsveld t incisedor 3- to 5-foliate, Tylecodon sometimes (2004), rarelyterete, rev.;Mort et al.,Amer.|. Bot. & Cotyledon marginsusuallycrenate,serrateor dentate,partly 92:1170-1176 (2005),mol.phylog. with bulbils (usuallyin sect.Bryophyllum),rarely rarely axillary, corymbose Procumbent to erect shrublets to shrubs, rarely entire; inflorescences woody; leaves or paniculate thyrsoids, partial inflorescences climbers; stems usually becoming lobed or orbicular, rarely or terete, flat decussate, few- to 1-flowered; dichasial,rarelyinflorescences thyr(glandular) inflorescences hairy; or glabrous flowers4-merous,* erect(usuallyin sect.Kalanmonochasia in ending dichasia, several soids with choe)or pendent(usuallyin sects.Kitchingiaand flowers; corolla tube Bryophyllum);sepals free, connate or forming with one to many pendent complex circalyxlcorolla dried hairy or glabrous; long, sometimest inflatedtube (usuallyin sect. exserted, filaments groove; basal along cumscissile Bryophyllum);petals usually brightly coloured, tacarpels tube; with corolla connate hairy where lobesshorterthan corollatube,erect,spreadingor * cuplike. scales stylodia;nectary into erect pering reflexed;filamentsexsertedor included,connate = g.Elevenspecies,southernand easterntropical to corollatube at base(sect.Bryopltyllum)or at or n Arabian Peninsula. abovemiddle (sects.Kalanchoeand Kitchingia); Africa, south-western data (Levsenet al.,l.c.), molecular to According carpelsfree to somewhatconnateat base,erector is polyphyletic. orbiculataL. Cotyledon the variable somewhatspreading(sect.Kitchingia);fruits erect present complex calyx/corolla The circumscissile f o l l i c l e sn. = u s u a l l y1 7 ,a l s o1 8 , 2 0 , 3 4 , 3 5 , 3 6 , 5 1 , TyIeleast some in at is found also in all Cotyledon eastern mainly Madagascar, 85.About 144species, may (Moran 2000), and spp. Kalanchoe codon and and southernAfrica,to tropicalAfrica,Arabiaand genera. three for these represent a synapomorphy Asia;sometaxa (especially tropicaland Southeast K. pinnata (Lam.) Pers.)are neophyticinvaders throughoutthe tropics. Divided into three sections:the widespread III. S usrA ,M.C nR ssur.oID E A EB urnett (183 5) . sect.Kalanchoe,and the Malagasysects.KitchinLeavesdecussate*,rarely ternate or whorled; flowgia andBryophyllum. ers haplostemonous*; anthers slightly introrsex, nucellus tenuinucellate*; fruits opening * comToelken 31. Tylecodon pletely along whole suture, but releasing seeds TylecodonToelken,Bothalia 12'378(197S);van Jaarsveld& through apical pore*; seedssinuate-unipapillate*. Koutnik, Tylecodon& Cotyledon(2004),rev. Genera33 + 34. Shrubletsor dwarf geophytesto pachycauldwarf Note that Crassuloideae are morphologically trees to 2.5m; stems succulent,rarely woody, highly derived, although cladistically they are the usuallywith flaking bark; leavesusuallycrowded first clade branching off from the remainder of the at stem tips, soft-herbaceous,with elongate family. t12 J.Thiede and U. Eggli at base,partly with apex papillateon outer face and with distinct appendage(usually in subg. Crassula)often* whitish;filamentsshortlyadnate to petalsat baseand alternatewith these;carpels usuallyfree;fruits rarely nutlike and indehiscent. n = 8,7 andpolyploids.About 195species; mainly southern Africa, a few speciesin sub-Saharan Africa and south-western Arabia,someephemeral herbs ('TiIIaea')distributed worldwide, and the oniy genusof the family in Australia. Divided into the paraphyleticsubg.Disporocarpa with nine sections(hydathodesof type I, rarelytype II; n = 8, rarely 7) and subg.Crassula with elevensections(hydathodesof type IIl' n - 7 with two satellites;Friedrich 1.973;Toelken1977 l.c.;Martin and von Willert 2000).The ephemeral herbsof sects.Helophytumand Glomeratae,often segregated asgenus Tillaea,arenestedwithin Crassula,accordingto moleculardata ('t Hart unpubl. data). 34. HypagophytumA. Berger HypagophytumA. Berger in Engler & Prantl, Nat. Pflanzenfam., ed. 2, l8a:467-468 (1930); Gilbert, Opera Bot. 12r:47-50(1993). Tuberousgeophytes;stemsone to few, droughtdeciduous;leavesternate,sessileor with petiolelike base, somewhat spurred at base, flat; inflorescences usually with 3 monochasiabelow terminal flower; flowers 10-l2-merous, stellate; sepalsfree; petalswhite or with faint pink tinge; 33. CrqssulaL. carpelsfree,laterallycompressed, Fig.31 constrictedinto two segments, upper part spiny-papillate, with CrassulaL., Sp.Pl;282 (I753); Toelken,Contr. Bolus Herb. long filiform stylodia; fruits 2-seeded, breaking 8 (1977), rev. southern African taxa; Tölken, f. Adelaide Bot. Gard. 3:57-90 (1981), rev. Austral. taxa; Blwater & transversely at the constriction, upper seed Wickens, Kew Bull. 39:699-728(1984), rev. New World dispersedwithin the upper part of the carpel, taxa; Mort et. al., IOS Bull. 12:35-36(2004),mol. phylog. lower seed releasedseparately.Only 1 species, H. abyssinicum(Hochst.ex A. Rich.)A. Bergerin (abstract). north-western Ethiopianhighlands. TillaeaL.(17s3). Characterised by a uniquecombinationof speR o c h e aD C . ( 1 8 0 2 ) . cialisedfeatures,which all occur in Crassula(tuPerennialor rarely annual herbs to (sub)shrubs, bers with annualshoots,whorled leaves,hydathrarely tuberous geophytes;glabrous, papillate odesalongleafmargins,haplostemonous andpolyor hairy; leavesdecussateor rarely in whorls merousflowers,and the peculiarfruits). The seed of 4, partly with bladder-cellidioblastsand leaf surfacestructurewas given as costateby Gilbert bases connate within a pair (usually in subg. (1989andl.c.)and Knapp(1994),whichprompted Crassula);inflorescencesthyrsoids with 1 to 't Hart (1995:169)to placethe genusin his 'Semany dichasia,sometimespartial inflorescences doideae'.However,accordingto Knapp(1997),the glomerate,monochasiaor reduced to solitary seedsurfacestructurein fact correspondsto the flowers;corollaurn-shapedto tubular or stellate; sinuate-papillate (Crassula-)type, clearlyfavourflowers(2-)5(-12)-merous, usuallysmall; sepals ing the placementin Crassuloideae. Hypagophyshortly connate at base;petals shortly connate tum maybe nestedwithin Crassulaand closestto Fig.31. Crassulaceae. Crassulacolumnaris.A Flowering plant. B Flowet openedout. C Youngplant seenfrom above. (Berger1930) .' \ -:. ... :,, ::l Se Crassulaceae ace bg. ate rels )nt. nly :an :ral th e ro]I, uIa )7 7 :ral ten 'asrbl. :en3ot. ;hrrlelat; ow lte; 8e; nto .ith ing :ed rel, ies, 'in Petuthrly:ed ,€ ft ted Sethe the urhyIto 113 its sect.Petrogeton,whichsharesthe tuberous habit Bywater,M., Wickens,G.E. 1983.New world speciesof the genus Crassula.Kew Bull. 39:699-728. and exhibits in some speciesleaveswith short petiCaballero,A., |imdnez, M.S. 1977.Contribuciön al estudio ole and in whorls of 4, monochasial inflorescences anatömicofoliar de las crassuläceas canarias.Vieraea with stellateand polymerous flowers, and long fili7:Il5-132. form stylodia. The samepeculiar fruit type is found Calie,P.l.1981.SystematicstudiesinSedum sectionTernata (Crassulaceae). Brittonia 33:498-507 . in sect.Glomerataep.p.(cf. Stopp 1957). Candolle,A.P.de 1828.Mdmoire sur la famille des Crassulacdes.Paris:Treuttel& Würtz. Clausen,R.T. 1959.Sedumof the Trans-MexicanVolcanic SelectedBibliography Belt: an exposition of taxonomic methods. Ithaca: Comstock. Akiyama, S., Ohba, H., Wu, S.-K. 2001.A new variety of Clausen,R.T. 1975.Sedumof North America north of the Mexican Plateau.Ithaca:Cornell UniversityPress. Sinocrassulapaoshingensis(S.H. Fu) H. Ohba et al. Clausen,R.T. 1977.Biennial speciesof Sedumof the Sierra (Crassulaceae). I. |ap. Bot. 76:222-226. Madre Occidentaland the Mexican Plateau.Bull. TorAlm, T. 2004.Ethnobotanyof Rhodiola rosea(Crassulaceae) rey Bot. Club 104:209-217. in Norway.Sida21, l:321-344. Bahadur,8., Ramaswamy,N., Srikanth,R. 1986.Studieson Cronquist,A. 1968.The evolutionand classificationof flowering plants.Boston:Houghton Mifflin. the floral biology and nectar secretionin someKalanIn: Kapil, R.P.(ed.) Pol- Cullen, I. i995. Crassulaceae.In: Cullen, 1., Alexanchoe species(Crassulaceae). der, ).C.M., Brady, A., Brickell, C.D., Green, P.S., lination biology -an analysis.New Delhi: Inter-India Heywood, V.H., Jorgensen, P.-M., Iury, S.L., Publications, pp. 251-259. Knees, S.G., Leslie, A.C., Matthews, V.A., RobBaskin,J.M.,Baskin,C.C.I972. Germinationcharacteristics son, N.K.B., Walters, S.M., The European Garden of Diamorphacymosaseedsand an ecologicalinterpreFlora, IV. Dicotyledons, part II. Dilleniaceae to tation. Oecologia(Berlin) 10:17-28. Krameriaceae. Cambridge: Cambridge University Baskin, |.M., Baskin, C.C. 1977. Germination ecology of Press,pp.170-244. SedumpulchellumMichx. (Crassulaceae). Amer. |. Bot. Davis,G.L. 1966.Seegeneralreferences. 64:1242-1247. Behnke,H.-D. 1991.Distribution and evolutionof forms Deil,U. 1991.Rockcommunitiesin tropicalArabia.In: Engel,T.,Frey,W, Kürschner,H. (eds)Contributionesseplastidsin the dicotyledons. and typesof sieve-element lectaead floram et vegetationemorientis.Berlin: Flora Aliso 13:167-182. et VegetatioMundi, pp. 175-187. In: Engler, A., Prantl, K., Berger, A. 1930. Crassulaceae. Die natürlichen Pflanzenfamilien,ed. 2, 18a.Leipzig: D enton, M.F. I979. Cytological and reproductive differentiBritation in SedumsectionGormania(Crassulaceae). W. Engelmann,pp. 352-483. t o n i a3 l : 1 9 7 - 2 1 I . Bland, K.P. 1995. Phytomyza rhodiolae Griffiths, 1976 (Diptera: Agromyzidae), a leaf-miner in roseroot, Denton, M.F. 1982.Revision of Sedum section Gormania (Crassulaceae). Brittonia 34:48-77. new to Britain. Entomol. Sedumrosea(Crassulaceae), Denton, M.F., Kerwin, J.L. 1980. Survey of vegetative Gaz.46:267-269. flavonoids of Sedum section Gormania (CrassuBoiteau, P., Allorge-Boiteau,L. 1995.Kalanchoe(Crassulaceae).Canad.I. Bot. 58:902-905. lacdes)de Madagascar.Systdmatique,dcophysiologie Ebel, F.,Hagen,A., Kümmel, F. 1991a.Beobachtungenzur et phytochimie.Paris:Karthala, Wuchsrhythmik von Orostachysspinosus(L.) Sweet Böttcher, W., läger, E.J. 1984. Zur Interpretation der (Crassulaceae). Wissensch.Z. Univ. Halle 40:47-68. Verbreitungder Gattung SedumL. s.l. (Crassulaceae) und ihrer Wuchsformtypen. Wissensch.Z. Univ. Halle Ebel, F., Hagen, A., Kümmel, F. 1991b. Beobachtungen zur Wuchsrhythmik und "Knospenbildung" einiger 33:127-14I. Greenovia-und Aeonium-Arten (Crassulaceae). Flora Bowman, R.N. 1983.Intraspecificvariability of leaf cuti85:187-200. cle alkanesin Sedumlanceolatumalong an elevational Eckert, G. 1966. Entwicklungsgeschichtliche und blütengradient.Biochem.Syst.Ecol.11:195-198. anatomischeUntersuchungenzum Problem der ObBraun, U.1987.A monograph of the Erysiphales(powdery diplostemonie.Bot. |ahrb. Syst.85:523-604. mildews).NovaHedwigia,Beih.89:l-700. Braun-Blanquet, |., Sutter, R. 1982. Zur Kenntnis der Eggli,U. 1988.A monographicstudy of the genusRosularia (Crassulaceae-Sedoideae). Bradleyasuppl.6:1- I 19. in den BündCrassulaceen-Pioniergesellschaften ner Alpen. |ahresber. Naturf. Gesell. Graubünden Eggli, U. (ed.) 2003. Illustrated Handbook of Succulent Plants,VI. Crassulaceae. Berlin HeidelbergNew York: 99:75-83. Springer. Burchard,O.1929.Beiträgezur Ökologieund Biologieder Eglinton, G., Gonzalez,A.G., Hamilton, R.|., Raphael,R.A. Kanarenpflanzen. Bibl. Bot. 98:l-262, pls. 1-78. 1962.Hydrocarbon constituents of the wax coatings Byalt, V.V. 1997.Meterostachyssikokianas(Crassulaceae), of plant leaves:a taxonomic survey. Phytochemistry a new speciesand genus for the flora of China (in 1:89-102. Russianwith Englishsummary). Bot. Zhurn. (Moscow Ellenberg,H. 1996.VegetationMitteleuropasmit den Alpen. & Leningrad)82:128-1 30. 5. Auflage.Stuttgart:Ulmer. paradoxa, a rare speciesfrom Byalt, V.V. 1998.Orostachys Endress,P.K.,Stumpf,S.I qgt. The diversityof stamenstructhe Russianfar East.Cact.Succ.I. (U.S.)70:262-263. 'lower' tures in Rosidae(Rosales,Fabales,Proteales, Bywater,M. 1980.Observationson seedsof Crassulasect. Sapindales). Bot. |. Linn. Soc.107:217-293. Rosulares.Kew Bull. 35:401-402. J. Thiede and U. Eggli EngelmannW. 1960.EndogeneRhythmik und photoperi- Hart, H. 't 1994a.The evolution of life-forms, growthodischeBlühinduktion bei Kalanchoe.Planta 55:496forms secondary growth in Eurasian Sedoideae (Crassulaceae). 5 11 . Bradleya12:37-56. Erdtman,G.1952.Seegeneralreferences. Hart. H. 'tI994b. The unilacunartwo-tracenodal structure Fehrenbach, S., Barthlott, W. 1988. MikromorpholoBot. of the caudexof RhodiolaroseaL.(Crassulaceae). gie der Epicuticular-Wachseder Rosales s.l. und |. Linn. Soc.116:235-241. deren systematischeBedeutung. Bot. |ahrb. Syst. Hart, H. 't 1995.Infrafamilial and genericclassificationof 109:407-428. the Crassulaceae. In: Hart, H.'t, Eggli, U. (eds) EvoluFernandes,F.M.1997.Restorationprogrammefor Madeira's tion and systematicsof the Crassulaceae. Leiden:Backendangeredplants.Plant Talk no. 10:19 huys, pp. 159-172. Fdtrd,f., Lebögue,A.1964.Embryogdniedes Crassulacdes. Hart, H. 't1997a.Crassulaceae. In: Oldfield,S.,tom. cit., pp. C. R. Acad.Sci.Paris258:5035-5038. 20-23. 't Fishbein,M. et al. 2001.Seegeneralreferences. Hart, H. I997b. Diversity within MediterraneanCrassuFriedrich, H.-C. 1973. Zur Cytotaxonomieder Gattung Craslaceae.Lagascalia 19:93-100. sula. Garciade Orta, Sdr.Bot. l:49-66. Hart, H. 't, Alpinar, K. 1999.Sedum ince (Crassulaceae), Fröderström,H. 1930-1935. The genusSedum.Asystematic a new speciesfrom southern Anatolia. Edinburgh I. essay.I-IV. Acta Horti Gothoburgensis5:1-75,6:I - I I 1, Bot.56:1Bl-194. 7 : l - 1 2 6 ,1 0 : l - 2 6 2 . Hart, H. 't, Arkel, J. van 1985. Quantitative aspects of Gess,S.,Gess,F.,Gess,R. 1998.Birds,waspsandTylecodon. the influence of day-length and temperature on Pollination strategiesof two members of the genus Sedum telephium (Crassulaceae).Acta Bot. Neerl. Tylecodonin Namaqualand.Veld Flora 84:56-57. 34:115-118. Gilbert, M.G. 1985.The genusSedumin Ethiopia.Bradleya Hart, H. 't, Berendsen,W. 1980.Ornamentationof the testa 3:48-52. inSedum (Crassulaceae). Pl. Syst.Evol. I35:I07-117. Gilbert, M.G. 1989. Crassulaceae. In: Hedberg, I., Ed- Hart, H. 't, Bleij, B. 1999.Nieuwe namen in Sempervivum wards, S. (eds) Flora of Ethiopia, III. Pittosporaceae Sect.Jovibarba(Crassulaceae). Succulenta(NL) 78:35to Araliaceae. Addis Ababa: Ethiopian National 42. Herbarium,pp. 5-26. Hart, H.'t, Bleij,8.2003.Phedimus.In:Eggli,U. (ed.)IllusGolding,I. (ed.)2002.SouthernAfrican Plant RedDataList. tratedHandbook of SucculentPlants,VI. Crassulaceae. Pretoria:SABONET. Berlin HeidelbergNew York: Springer,pp.196-203. Gontcharova,S.B.1999.Ornamentationof the testaof some Hart, H.'t, Eggli, U. 1995.Introduction: evolution of CrasEasternAsian Sedoideae(Crassulaceae). Bull. Natl Sci. sulaceaesystematics.In: Hart, H. 't, Eggli, U. (eds) Mus.,Tokyo,ser.B 25:131-141. Evolutionand systematicsof the Crassulaceae. Leiden: Gregory,M. 1998.Crassulaceae. In: Cutler,D.F.,Gregory,M. Backhuys,pp.7-15. (eds)Anatomy of the Dicotyledons,2nd edn. IV. Saxi- Hart, H.'t, Eggli, U. 1998.Cytotaxonomicstudiesin Rosufragales(sensuArmen Takhajan1983).Oxford:ClarenIari a (Cr assulaceae ). Bot. Helvetica 98:223-234. don Press,pp.20l-220. Hart, H. 't, Koek-Noorman,J. 1989.The origin of the woody Günthart, A. 1902.Beiträge zur Blütenbiologieder CruSedoideae(Crassulaceae). Taxon 38:535-544. 't, ciferen,Crassulaceen und der Gattung Saxifraga.Bibl. Hart, H. Sandbrink, |.M., Csikos, I., Ooyen, A. van, B o t .5 8 : 1 - 9 7 . Brederode, |. van 1993. The allopolyploid origin Ham, R.C.H.J. van 1994. Phylogenetic implications of of Sedum rupestre subsp. rupestre (Crassulaceae). chloroplastDNA variation in the Crassulaceae. Thesis, Natural hybrids in Sedum (Crassulaceae) 4. Pl. Syst. University of Utrecht. Evol.184:195-206. 't, Ham, R.C.H.}.van, Hart, H.'t 1998.Phylogeneticrelation- Hart, H. van Ham, R.C.H.J.,Stevens,J.F.,Elema, E.T., ships in the Crassulaceaeinferred from chloroplast Klis, H. van, Gadella, T.W.f. 1999. Biosystematic, DNA restriction-sitevariation. Amer. |. Bot. 85:123molecular and phytochemical evidence for the t34. multiple origin of sympetaly in Eurasian Sedoideae (Crassulaceae). Hart, H.'tl975.The pollenmorphologyof 24EuropeanspeBiochem.Syst.Ecol.27:407-426. ciesofthe genusSedumL. Pollen Spores16:373-387. Hegnauer,R. 1964.Chemotaxonomieder Pflanzen.Band Hart, H. 't 1982.The systematicposition of Sedumtubero3. Dicotyledoneae: Acanthaceae-Cyrillaceae. Basel: suz Coss.& Let.(Crassulaceae). Proc.Koninkl. Nederl. Birkhäuser,pp. 572-584. Akad. Wetensch.,ser.C 85:497-508. Hegnauer,R. 1989.Chemotaxonomieder Pflanzen,Band 't Hart, H. 1985a.Chromosomenumbers in Sedum(Cras8. Nachträge zu Band 3 und Band 4 (Acanthaceae from Greece.Willdenowia15:115-135. sulaceae) bis Lythraceae).Crassulaceae: pp.3a2450, 710.Basel: 't Hart, H. 1985b.Sexualreproduction and hybridisation Birkhäuser. in Sedum telephium (Crassulaceae). Acta Bot. Neerl. Hideux, M.l. 1981.Le pollen. Donndesnouvellesde la mi34:l-4. croscopiedlectroniqueet de l'informatique: structure Hart, H. 't 1985c.The vascular pattern of the flowers of du sporoderme de Rosidae-Saxifragales, dtude com(Crassulaceae). Sedumanacampseros Acta Bot. Neerl. parativeet dynamique.Paris:Agencede Coopdration 34:lI9-121. Culturelle et Technique. Hart, H.'t 1990.Variation in the structure of the flowersof Huber, H. 1961.Crassulaceae. In: Hegi, G. (ed.) Illustrierte Sedum.SedumSoc.Newslett.13:11-17. Flora von Mittel-Europa.2.Auflage.Band IV/2. Teil. 't Hart, H. 1991.Evolution and classificationof the EuTeilbandA.München:CarlHanser, pp.62-125. ropean Sedumspecies(Crassulaceae). Flora Mediter- Hutchinson,I.1973.The families of flowering plants,ed. 3. ranea1:31-61.Madrid: OPTIMA. Oxford:ClarendonPress. Crassulaceae theae ure ,^ . )ut. rof rlu1 pp. sure), ht. of on erl. tsta 7. 'um 351..^ tu5- eae. ras:ds) len: )suody IAn) igin ae). ,ysI. 1.T., rtic, the leae and rsel: and :eae rsel: 1.3. Huxley,A., Griffiths,M., Levy,M. (eds) 1997.TheNew RHS Dictionary of Gardening.4 vols. London: MacMillan. Jaarsveld,E.]. van 1994.The distribution of Tylecodonand in SouthAfrica and Namibia. Cotyledon(Crassulaceae) In: Seyani,J.H.,Chikuni, A.C. (eds)Proc.XIII Plenary MeetingAETFAT,Malawi,pp. 1157-1163. facob,F.H. 1964.A new speciesof Thuleaphisfrom Wales, Scotland,and Iceland (Thuleapäissedi n. sp. on Sedum rosea.Proc. Roy. Entomol. Soc.,London, ser. B, T a x o n o m y3 3 : 1 1 1 - 1 1 6 . läger-Zirn, I. 1989. Zur Kenntnis von Crassula pageae Tölken (syn.Pagellaarcheri).Trop.-subtrop.Pflanzenwelt70:l-71. Mainz: F. Steiner. falas,|., Suominen,|., Lampinen, R., Kurtto, A. 1999.Atlas Florae Europaeae.Distribution of vascular plants in Europe,12.Helsinki:Committeefor Mapping the Flora of Europe& SocietasBiologicaFennicaVanamo. Iay, M. 1971.Quelquesproblömes taxonomiqueset phylogdndtiquesdes Saxifragacdesvus ä la lumiöre de la biochimie flavonique.Bull. Mus. Natl Hist. Nat. II, 42:754-775. fensen, L.C.W. 1966. Comparative anatomical studies in three subfamiliesof the Crassulaceae. Ph.D. Thesis, Universityof Minnesota. JimdnezC., G., SoberdnM., I. 1989.Laboratory rearing of Sandiaxami xami (Lycaenidae, Eumaeini).|. Res.Lepidoptera 27:268-271. Johnson,S.D.,Ellis,A., Carrick,P.,Swift,P.,Horner,N., |anse van Rensburg,S., Bond, W.J. 1993.Moth pollination and rhythms of advertisementand reward in Crassula S. African |. Bot. 59:511fascicularis(Crassulaceae). 513. |ohri, B.M. et al.1992.Seegeneralreferences. Jorgensen,T.H., Olesen,I.M. 2000.Growth rules basedon the modularity of the Canarian Aeonium (Crassulaceae)and their phylogeneticvalue.Bot. J. Linn. Soc. 132:223-240. forgensen,T.H., Olesen,I.M. 2001. Adaptive radiation of island plants:Evidencefrom Aeonium (Crassulaceae) of the Canary Islands. Perspec.Pl. Ecol. Evol. Syst. 4:29-42. fürgens,N. 1995.Contributions to the phytogeographyof 't, Eggli, (eds) Crassula.In:Hart, H. U. Evolution and systematicsof the Crassulaceae. Leiden:Backhuys,pp. 136-150. Keeley,J.E.1998.CAM photosynthesisin submergedaquatic plants.Bot. Rev.64:12l-175. Kim, |.-H. i994. Pollenmorphology of genusSedumin Korea.I. Pl. Biol. 37, 2:245-252. acid metabolism Kluge,M., Brulfert, J. 1996.Crassulacean in the genusKalanchoe:ecological,physiologicaland biological approaches.In: Winter, K., Smith, A.P., Smith, I.A.C. (eds) Crassulaceanacid metabolism. Biochemistry, ecophysiology and evolution. Berlin HeidelbergNew York: Springer,pp.32a435. Knapp, R. 1973.Die Vegetationvon Afrika. Stuttgart: G. Fischer. Knapp,U.1994.Skulptur der Samenschale und Gliederung Bot. |ahrb. Syst.116:157-187. der Crassulaceae. Knapp,V. 1997.Samenoberflache und Systematikder Saxifragaceaeund Crassulaceae. Ph.D. Thesis,University of Kaiserslautern,pp. l-ßa. Krach, I.E. 1976. Samenanatomieder Rosifloren. 1. Die Samender Saxifragaceae. Bot. |ahrb. Syst.97:l-60. Kurkin, V.A.,Zapesochnaya, G.G. 1986.The chemicalcomposition and pharmacologicalpropertiesof Rhodiola plants (in Russian).Khim.-farm. Zhurn. 20:1231-1244. Leinfellner,W. 1954.Beiträge zur Kronblattmorphologie. I I I. D ie Kronblätter der Gattung Pachyphy t um. O esterr. B o t .Z . 1 0 1 : 5 8 6 - 5 9 1 . Levin, G.A., Mulroy, T.W. 1985.Floral morphology, nectar production, and breedingsystemsin Dudleya subgenusDudleya (Crassulaceae). Trans.San Diego Soc. Nat. Hist. 2l:57-70. Lippert, W. 1995.Familie Crassulaceae. Dickblattgewächse. In: Weber, H.E. (ed.) Gustav Hegi, Illustrierte Flora von Mitteleuropa, Band IV Teil 24: Spermatophyta: Angiospermae,Dicotyledones2(2), 3. Auflage.Berlin: Blackwell,pp.69-129. Lösch,R. 1990.FunktionelleVoraussetzungder adaptiven Nischenbesetzung in der Evolution der makaronesischen Semperviven.Diss.Bot. 146:I-482. Manheim, B.S.lr, Mulroy, T.W.,Hogness,D.K.,Kerwin, |.L. 7979. Interspecific variation in leaf wax of Dudleya. Biochem.Syst.Ecol.7:I7-19. Marchant,T.A.,Alarcon, R., Simonsen,).A., Koopowitz,H. 1998. Population ecology of D udleya mult i caulis ( Crassulaceae):a rare narrow endemic. Madrono 45:215220. Martin, C.E.,Willert, D.l. von 2000.Leaf epidermalhydathodes and the ecophysiologicalconsequences of foliar water uptake in speciesof Crassulafrom the Namib desertin SouthernAfrica. Pl. Biol. 2:229-242. Mauritzon, I. 1930.Beitrag zur Embryologie der Crassulaceen.Bot. Notiser 1930:233-250. Mauritzon, 1.1933.Studienüber die Embryologieder Familien Crassulaceaeund Saxifragaceae. Lund: Hakan Ohlssons. Mayuzumi, S., Ohba, H. 2004.The phylogeneticposition of Eastern Asian Sedoideae(Crassulaceae)inferred from chloroplast and nuclear DNA sequences.Syst. Bot.29:587-598. Merxmüller, H., Friedrich, H.-C., Grau, I. 197I. CytotaxonomischeUntersuchungenzur Gattungsstrukturvon Crassula. Ann. Naturhist.Mus.Wien 75:111-119. Mes,T.H.M. I995.Phylogenetic and systematicimplications of chloroplastand nuclear spacersequencevariation in the MacaronesianSempervivoideae and relatedSedoideae.In: Hart, H. 't, Eggli, U. (eds) Evolution and systematicsof the Crassulaceae. Leiden:Backhuys,pp. 30-44. Mes,T.H.M.1996.Origin and evolutionof the Macaronesian (Crassulaceae). Sempervivoideae Ph.D.Thesis,University of Utrecht. Mes,T.H.M.,Hart, H.'t 1994.SedumsurculosumandS.jaccardianum (Crassulaceae) sharea unique 70 bp deletion in the chloroplastDNA trnL (UAA)-trnF (GAA) intergenicspacer.Pl. Syst.Evol. 193:213-221. Mes,T.H.M.,Hart, H. 't 1996.The evolutionof growth-forms in the Macaronesiangenus Aeonium (Crassulaceae) inferredfrom chloroplastRFLPsand morphology.Mol. E c o l .5 : 3 5 1 - 3 6 3 , Mes, T.H.M., Brederode,f. van, Hart, H. 't 1996.Origin of the woody MacaronesianSempervivoideae and the phylogeneticposition of the East African speciesof Aeonium.Bot. Acta 109:477-49I. Mes,T.H.M.,Wijers,G.|.,Hart, H.'t1997. Phylogeneticrelationships in Monanthes(Crassulaceae) basedon mor- ll6 J. Thiede and U. Eggli phological,chloroplastand nuclear DNA variation. I. Evol.Biol. 10:193-216. Meusel, H., |äger, E., Weinert, E. 1965. Vergleichende Chorologie der zentraleuropäischenFlora. Band 1. Crassulaceae. Jena:G. Fischer. Moran, R. 1949.Graptopetalumbartramii in Chihuahua. DesertPl. Life 2l'53-56. Moran, R. 1996.Altamiranoa into Sedum (Crassulaceae). Haseltonia4:46. Moran, R. 2000.Circumscissionin Cotyledon:withthoughts on what is Cotyledon,and on how A.P.de Candollewas right all along and they should have listened. Cact. Succ.J. (U.S.)72:306-308. Moran,R.,Meyrän,1.1974.Tacitusbellus,un nuevogdnero y especiede Crassulaceae de Chihuahua,Mdxico.Cact. Suc.Mex. l9:75-84. Morgan, D.R.,Soltis,D.E. 1993.Phylogeneticrelationships among membersof Saxifragaceae sensulato basedon rbcl sequencedata.Ann. Missouri Bot. Gard. 80:631660. Mort, M.8., Soltis, D.E., Soltis, P.S.,Francisco-Ortega,|., Santos-Guerra,A. 2001. Phylogenetic relationships and evolution of Crassulaceaeinferred from matK sequencedata.Amer. J. Bot. 88:76-91. Mort, M.E., Soltis, D.E., Soltis, P.S.,Francisco-Ortega,)., Santos-Guerra,A. 2002.Phylogeneticsand evolution of the Macaronesianclade of Crassulaceaeinferred from nuclearand chloroplastsequencedata.Syst.Bot. 27:271-288. Moteetee,A., Nagendran,C.R. 1997.Comparativeanatomical studiesin five southernAfrican speciesof Crassula. I. Structureof the stem and the root. S. African |. Bot. 63:90-94. Nakanishi,H.2002.Splashseeddispersalby raindrops.Ecol. Res.(Tokyo)17:663-67l. Newton, D.J.,Chan, J. 1998.South Africa's trade in southern African succulentplants.fohannesburg:TRAFFIC East/SouthernAfrica. Nyffeler, R. 1992.A taxonomic revision of the genus MonanthesHaworth (Crassulaceae). Bradleya10:49-82. Ohba,H. 1978.Genericand infragenericclassificationof the Old World Sedoideae(Crassulaceae). J. Fac.Sci.,Univ. Tokyo III, Bot. 12139-198. Ohba,H. 1989.Biogeographyof the genusRhodiola (Crassulaceae),with specialreferenceto the floristic interaction betweenthe Himalalya and the Arctic Region. In: Ohba,H., Hayami, I., Mochizuki, K. (eds) Current aspectsof biogeographyin WestPacificand EastAsian regions.UniversityofTokyo,pp. 115-133. Oldfield, S. (ed.) 1997.Cactusand SucculentPlants - Status Surveyand ConservationAction Plan. Gland and Cambridge:IUCN/SSCCactusand SucculentSpecialist Group. Olfelt, |.P.,Furnier, G.P.,Luby,].L. 1998.Reproductionand development of the endangered Sedum integrifolium ssp.leedyi(Crassulaceae). Amer. J. Bot. 85:346-351. Olfelt, |.P.,Furnier, G.P.,Luby, |.L. 2001.What data determine whethera plant'staxonis distinct enoughto merit legal protection?A casestudy of Sedumintegrifulium (Crassulaceae). Amer.f. Bot. 88:401-410. Parnell,I. 199I. Pollen morphology of JovibarbaOpiz and SempervivunL. (Crassulaceae). Kew Bull. 46:733-738. Parra,V., Vargas,C.F.,Eguiarte,L.E. 1993.Reproductivebiology, pollen and seed dispersal,and neighborhood size in the hummingbird-pollinated EcheveriagibbiAmer. |. Bot. 80:153-159. flora (Crassulaceae). Parra,V.,Vargas,C.F.,Eguiarte,L.E. 1998.Is Echeveriagibbiflora (Crassulaceae)fecundity limited by pollen availability? An experimentalstudy. Funct. Ecol. t2:59t595. Pörezde Paz,P.L. 1980.Contribuciön al atlaspalinolögico de dndemismosCanario-Macarondsicos. Part III. Bot. Macar.7:77-112.Gran Canaria:|ardin Botänico. Pilon-Smits,E.A.H. 1992.Yariationand evolutionof CrassulaceanAcid MetabolisminSedumand Aeonium (Crassulaceae).Ph.D.Thesis,Universityof Utrecht. Quimby, M.W. 1971.The floral morphology of the Crassulaceae.Ph.D.Thesis,Cornell University,Ithaca. Raadts,E. 1979.Rasterelektronenmikroskopische und anatomische Untersuchungen an Konnektivdrüsen von Kalanchoe(Crassulaceae). Willdenowia 9:169-175. Riefnet R.E.)r, Bowler,P.A.,Mulroy,T.W.,Wishner,C.2003. Lichenson rock and biologicalcrustsenhancerecruitment successof rare Dudleyaspecies(Crossulaceae) in SouthernCalifornia.Crossosoma29:l-36. Rivas-Martinez,S., Wildpret de la Torre, W., del Arco, M., Rodriguez, O., Pörez de Paz, P.L., Garcia-Gallo,A., Acebes,f.R., Fernändez-Gonälez, F. 1993.Las comunidadesvegetalesde la Islade Tenerife(IslasCanarias). Itinera Geobot.7:169-374. Rocdn,T. 1928.Beitragzur Embryologieder Crassulaceen. SvenskBot. Tidsskr.22:368-376. Rocher, E.J. de, Harkins, K.R., Galbraith, D.W., Bohnert, H.J. 1990. Developmentallyregulated systemic endopolyploidy in succulentswith small genomes. Scienc2 e50:99-101. Rünger,W, Wehr,B. 1969.Über den Einflussder Tageslänge und der Temperatur auf die Blütenbildung einiger Echeveria-Arten.Gartenbauwissenscha ft 3q:t1I - I 43. Said,C. 1982.LesnectairesfloreauxdesCrassulacdes. Etude morphologique,histologiqueet anatomique.Bull. Soc. Bot. FranceLett. Bot. 129:231-240. Savolainen,V., Chase,M.W. et al. 2000.Seegeneral references. Savolainen,V., Fay,M.F.et al. 2000.Seegeneralreferences. Schönland,S. 1894.Crassulaceae. In: Engler,A., Prantl, K., Die natürlichenPflanzenfamilien,ed. 1, 3,2a.Leipzig: W. Engelmann, pp. 23-38. Sharma,A.K.,Gosh,S.i967. Cytotaxonomyof Crassulaceae. Biol. Zentralbl.suppl.86:313-336. Sin, |.-H., Yoo,Y.-G.,Park, K.-R. 2002.A palynotaxonomic study of the Korean Crassulaceae. Korean ). Electron Micro scopy 32:345 -360. Soltis,D.E.,Soltis,P.E.1997.Phylogeneticrelationshipsin Saxifragaceae sensulato: a comparison of topologies basedon 18SrDNA and rbcL sequences. Amer. f. Bot. 84:504-522. Soltis,D.E.et al. 2000.Seegeneralreferences. Soltis,D.E.,Fishbein,M., Kuzoff, R.K. 2003.Re-evaluating the evolution of epigyny: data from phylogeneticsand floral ontogeny.Intl |. Pl. Sci.164:5251-5264. Souöges,R. 1936.Les relationsembryogdniquesdes Crassulacdes, Saxifragacdes et Hypdricacdes. Bull. Soc.Bot. France83:317-329. Stevens,|.F. 1995a.Chemotaxonomyof the Eurasian Sedoideaeand Sempervivoideae. In: Hart, H. 't, Eggli,U. (eds) Evolution and systematicsof the Crassulaceae. Leiden:Backhuys,pp. 30-44. Crassulaceae bbiibb'ail9l;ico Bot. ;suras- nge Lger 43. ude loc. fer- I17 Stevens,J.F.1995b.The systematicand evolutionarysignif- Torrey,1., Gray,A. 1838.A flora of North America, I. New York: Wiley & Putnan. icance of phytochemical variation in the Eurasian Se(Crassulaceae). Gronin- Troll, W. 1964.Die Infloreszenzen.Erster Band. Stuttgart: doideaeand Sempervivoideae G. Fischer. gen: Rijks University. Stevens, |.F.,Hart, H.'t, Hendriks,H., Malingrd,T.M.1992. Troll, W. 1969.Die Infloreszenzen.Zweiter Band, 1. Teil. Stuttgart:G. Fischer. Alkaloids of some European and MacaronesianSePhyto- Troll, W., Weberling,F. 1989.Infloreszenzuntersuchungen doideaeand Sempervivoideae(Crassulaceae). an monotelenFamilien.Stuttgart:G. Fischer. chemistry 3 I:3917-3924. Stevens,J.F.,Hart, H.'t, Hendricks,H., Malingrd,T.M. 1993. Uhl, C.H. 1948.Cytotaxonomicstudiesin the subfamilies Pl. Crassuloideae,Kalanchoideae,and Cotyledonoideae Alkaloids of the Sedumacre-grovp(Crassulaceae). -217 . Amer. J. Bot.35:695-706. of the Crassulaceae. Syst.Evol. 185:207 't, Bolck, A., Zwaving, I.H., Ma- Uhl, C.H. 1961.The chromosomesof the Sempervivoideae Stevens,J.F.,Hart, H. (Crassulaceae). Amer. ]. Bot. 48:lI4-I23. lingrd, T.M. 1994. Epicuticular wax composition of some European Sedum species.Phytochemistry Uhl, C.H. 1970.Chromosomesof Graptopetalumand ThoAmer. I. Bot. 57:1115mpsonella (Crassulaceae). 35:389-399. 't, rt2l. Stevens,|.F.,Hart, H. van Ham, R.C.H.J.,Elema,E.T.,van den Ent, M.M.V.X.,Wildeboer,M.,Zwaving,J.H. 1995. Uhl, C.H. 1976-1992.Chromosomesof Mexican Sedum. -402, I-VI. Rhodora 79:629-640,80:49I-512, 82:377 Distribution of alkaloids and tannins in the Crassu0. laceae.Biochem.Syst.Ecol.23:157-165. 85:243-252, 87:38I- 423,94:362-37 't, Elema, E.T., Bolck, A. 1996. Uhl, C.H. 1992. Polyploidy, dysploidy, and chromosome Stevens,I.F., Hart, H. and its hybrids. pairing in Echeverla(Crassulaceae) Flavonoid variation in Eurasian Sedum and SemperAmer. f. Bot. 79:556-566. v ivum. Phytochemistry41:503-5I 2. Uhl, C.H. 1993-1995.Intergeneric hybrids in the MexiStevens,P.F.2005.Seegeneralreferences. I-V. Cact.Succ.f. (U.S.)65:27I-273, can Crassulaceae. Stopp, K. 1957.Aberrante Dehiszenzformenbei Früchten 66:74-80, 175-179, 214-217, 67:144-147. einiger Crassula-Arten.Beitr. Biol. Pflanzen 34165175. Uhl, C.H. 1994-2005.Chromosomesand hybrids of Echeveri a (Cr assulaceae Supratman,U., Fujita, T., Akiyama, K., Hayashi,H. 2001. ). I - IX. Haseltonia 2:79 -80, 3:25-33, 3z34- 48,4:66- 88,5:2I - 36, 6:63-9 0, 8:7I -82, 9:l2l - | 45, Insecticidalcompounds from Kalanchoedaigremon11:138-149. tiana x tubiflora.Phytochemistry58:31I-314. Takhtajan,A.L.1969.Floweringplants,origin and dispersal. Uhl, C.H. 1996.Chromosomesand polyploidy in LenophylAmer. J. Bot. 83:216-220. lum (Crassulaceae). Washington,DC: SmithsonianInstitution Press,pp. 1Uhl, C.H., Moran, R. 1972.Chromosomesof Crassulaceae 31 0 . from |apan and South Korea.Cytologia37:59-81. Teeri, f.A., Overton, |. 1981.Chloroplastultrastructure in two Crassulaceanspeciesand an F1hybrid with differ- Uhl, C.H.,Moran, R. 1973.The chromosomesof Pachyphytum (Crassulaceae). Amer. f. Bot. 60:648-656. ing biomassdelta 13Cvalues.Pl. Cell Environ. 4:427Uhl, C.H., Moran, R. 1999. Chromosomes of Villadia 43r. and Altamiranoa (Crassulaceae).Amer. l. Bot. Teeri, |.A., Stowe,L.G., Murawski, D.A. 1978.The clima86:387-397. tology of two succulent plant families: Cactaceaeand U.S.Fish and Wildlife Service2004.Threatenedand EndanCanad.J. Bot. 56:L750-1758. Crassulaceae. gered SpeciesSystem (TESS).http:/endangered.fws. Thiede, I. 1995. Quantitative phytogeography, species gov/wildlife.html richness, and evolution of American Crassulaceae. 't, Eggli, U. (eds) Evolution and sys- Visser, J. 1981.South African parasitic flowering plants. In: Hart, H. CapeTown: |uta. Leiden: Backhuys,pp. tematics of the Crassulaceae. Vogel,S. 1954.BlütenbiologischeStudienals Elementeder 89-r23. dargestelltan der Flora Südafrikas. Sippengliederung, Thiede,1.2004.The genusDudleyaBritton & Rose(CrassuBot. Stud.1:1-338. laceae):its systematicsand biology.Cact.Succ.I. (U.S.) Wakabayashi, M., Ohba,H. 1999.Chromosomenumbersof 76:4-ll. 't sevenspeciesof Sedumand Sinucrassulaindica (CrasThiede,f., Hart, H. 1999.Transferof four PeruvianAltamisulaceae)in EastHimalaya.l. Iap. Bot 74:228-235. Novon 9:124ranoa speciesto Sedum(Crassulaceae). Wassmer, A. 1955. Vergleichend-morphologischeUnr25. tersuchungen an den Blüten der Crassulaceen. Thorne, R.F. 1968.Synopsisof a putatively phylogenetic Winterthur: P.G.Keller. classificationof the floweringplants.Aliso 6:57-66. Thorne, R.F. 1983.Proposednew realignmentsin the an- Wickens,G.E.,Bywater,M. 1980.Seedstudiesin Crassula subgen. Disporocarpa. Kew Br:Jl.34:629-637 . giosperms.Nordic |. Bot. 3:75-II7. Thorne, R.F. 1992.An updated phylogeneticclassification Wikström, N., Savolainen,V., Chase,M.W. 2001.Evolution of the angiosperms:calibrating the family tree. Proc. of the floweringplants.Aliso 13:365-389. Roy.Soc.London, ser.B 268:2211-2219. Tillson,A.H. 1940.The floral anatomyofthe Kalanchoideae. Wyatt, R., Stoneburger,A. 1981.Patternsof ant-mediated Amer. |. Bot. 27:595-600. pollen dispersalin Diamorpha smallii (Crassulaceae). Toelken,H.R. 1977.A revision of the genus Crassulain Syst.Bot. 6:l-7. southernAfrica. Parts 1 & 2. Contr.BolusHerb. 1-331, Xu, f.F.,Liu, C.8.,Han, A.M., Feng,P.S.,Su,Z.G.1998.Strate332-595. gies for the improvement of salidrosideproduction in Toelken, H.R. 1986. Crassulaceae. In: )essop, J.P., Pl. cell suspensionculturesof Rhodiola sachalinensis. Toelken,H.R. (eds) Flora of South Australia. Part I. Cell Rep. 17:288-293. Adelaide: Government Printing Office, pp. aß-a28. 118 ). Thiede and U. Eggli T.,Haruna,M., Yan,X.-Z.,Chang,|.-f.,Lee,K.-H. Yamagishi, 1989.Antitumor agents,i 10.1,2bryophyllin B, a novel potent cytotoxic bufadienolide from Bryophyllum pinn a t u m . l .N a t .P r o d .5 2 : 1 0 7 I - 1 0 7 9 . Yoshikawa,M., Shimadä,H., Shimoda,H., Murakami, N., Yamahara,J.,Matsuda,H. 1996.Bioactiveconstituents of Chinesenatural medicines.II. RhodiolaeRadix (1): chemical structures and antiallergic activity of rhodiocyanosidesA and B from the underground part (Pall.) Fisch.et Mey. (Crasof Rhodiola sachalinerzsis sulaceae).Chem. Pharmaceut.Bull. (Tokyo) 44:2086209t. c d TheFamilies and Genera of VascularPlants Edited bv K. Kubitzki T \f FloweringPlants. Eudicots I Berberidopsidales, Buxales, Crossosomatales, /\ Fabalesp.p., Geraniales,Gunnerales,Myrtales p.p., Pr oteales, Saxifr agales, VitaIes,Zygophyll aIes, Clusi aceae Alliance, Passifl oraceae Alli ance, Dilleniaceae, Huaceae,Picramniaceae, Sabiaceae VolumeEditor: K. Kubitzki in Collaborationwith C. Bayerand P.F. Stevens With 174Figures üspringer P r t . r t es sDr or . K r a u s K u s rrz rr L'niversitätHamburg BiozentrumKlein-Flottbekund BotanischerGarten O hnhor s t s t r aß1e8 22609Hamburg Germany Library of CongressControl Number: 2006928744 ISBN-10 3-540-32214-0 SpringerBerlin HeidelbergNew york ISBN-13 978-3-540-32214-6 SpringerBerlin HeidelbergNew york This work is subject to copvright. All rights are reserved,whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilmor in any other waF and storage in data banks. Duplication of this publication or parts thereof is permitted only under the provisions of the German Copyright Larv of September 9, 1965,in its current version, and permissions for use must always be obtained from Springer-Verlag. \riolations are liable for prosecution under the German Copyright Law. Springer is a part of Springer Science+BusinessMedia springer.com @ Springer-Verlag Berlin Heidelberg 2007 The use of general descriptive names, registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. Cover design: WMXDesign, Heidelberg, Germany Typesetting and production: LE-TEX lelonek, Schmidr & vöckler GbR, Leipzig, Germany Printedonacid-freepaper 3tl3t50lYL- 5 432t0