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COMUNICACIONES 1 71 -1 73, 1 98 2 Rev. Bio!. Trop., 30(2): Habiat of Neoregelia cuenta (Bromeliaceae) n coastal sand duns oí Maria, Brazl Luiz Drude de Lacerda* and John DuVall Hay** Deparamento de Ecoloia, Universidade Federal do Rio de Jneiro, I1ha da Cídade niversitaria, Río de Janeiro, R.J. , 2 1 941 (Received for publication arch 25 , 1982) Ren: e realizó n estudio de los habitats uiliads por . cuenta (R. Graham) L. Sínith en un ecosistema costero de dunas de arena en Barra de Maricá, Rio de Janeiro . Los resultads demuestan que esa especie tiene la capacidad de mantener una élevada densidad n ambientes normalmente adversos, aravs de epii ismo facultativo. The family Bromeliaceae is c ompoition widespread However, it can also be subdivided and horizontal verticl the two intermedia' zones 2 and 3. Once in �Íle other exteme zones, plant communities are herbaceous, thus precluding the possibility of epiphytic habitats. Zone 2 ig located in the into ecological groups based on mechanisns of water and nutrient absorption (Pittendrigh, 1 948). The types of habitats used by different spcies of bromeliads are many (pittendrigh, 1948, valley behind the pimary dunes and exhibits a clumped plant distribution (± 1 2 m in height, coverage ± 25%). It gradualIy gives way to Zone 3 which is denser, featuring saller and fewer open areas (± 6-7 m in height, coverage > 75%). These transitions occur within 300 m. Habitat use by . cuenta was quantiied uing a line-intercept technique. Two line tranects, perpendicular to the ocean and the vegetation zones, were established in Zones 2 and 3. ll individuals of . cruenta within 2 m of each transect were counted and their habitat r ecorded. Specifically, we noted if the individual was associated with another species, and if so, whether it was growing as an epiphyte An individual was considered to e associated if it was within 1 5 cm of at least one - Aragao, 1 967; Medina, 1 974). One aspect o f habitat u se in the Bromeliaceae is facultative epiphytism, present in each of the subfamilies (Medina 1 974). The present study deals with habitat use by Neoregelia cuenta (R . Graham) L. Smith in the Barra de Maricá, a coastal sand dune ecosystem, approximately 35 km E of Rio de Janeiro, Brazil (220 5 7 ' S, 430 08' W). The locl bromeliad lora consists of . cuenta, Bromelia an tiacan tha Bertol, Tilansia spp. and Aechmea sp . . Of these only . cruenta exploits both terrestial and epiphytic habitats. This species is one of the dominants in this system (acerda and Hay 1 977) anrl important in improving soil chemistry (Hay and acerda other species, and classiied as epiphytic when no part was in contact with he soil surface . Neoregelia cruenta can be found in all zones 1 980; Hay et al. , 1 981 ). The area stuied can b e divided into four distinct zones, with increaing distance from the ocean, based on differences in plant species * and structure (see Hay and Lacerda, 1980 for a complete descrtption). The study was done in throughout the Neotropics. It is subdivided into three subfamilies distinguished by loral and seed orphology (Smith and Downs, 1 974). of the Barra de Maricá, but it is most abundant in Zones 2 and 3 where it attains a similar Present address : Insituto de Biof{sica, Universidade Federal do Rio de Janeiro, llha da Cidade Univrsitária' Rio de Janeiro, RJ, 2 1 94 1 . ** Present address : Departamento de Bioloia Vegetal, Universidade de Brasilia, Brasília, D.F. 7091 9. 171 1 72 REVISTA DE BIOLOGIA TROPICAL Fig. 1 . Group of terrestrial individuals of Neoregel1 cuenta in the open areas of the second vgetation zone of the Barra de Maricá RJ). Epiphytic forms are also present in the backround, along with the third vegetation zone. Fig. 2. Epiphyic forms of . cuenta in the second vegetation zone of the Barra de Maricá (RJ). Fig. 3. Epiphytic form of N. cruenta in the canopy of the third veget'ion zone of the Barra de Maricá RJ). density of approximately 86 individuals/1 00 m 2 (Lacerda and Hay 1977). This maintainance . of density with a change in habitats appeas to be accomplished through epiphytism since insuicient open ground remais to support an equiva1ent number of terrestrial individuals. Typical habitat use by . cuenta in Zones 2 and 3 is shown in Figs. 1 , 2 , 3 . Figure 1 shows a group of individuals of . cuenta in the open areas of Zone 2. Figures 2 and 3 show examples of the use of the epiphytic habitat. In Fig. 2 N cuenta is epiphytic on a community group in Zone 2, while in Fig. 3 it is present within the tree canopy in Zone 3. When . cuenta was present in Zone 3 as a terrestrial, it occurred on the periphery of the closed communities or within the open areas. The results of this study are shown in Table 1 . A total of 222 individuals of . cuenta were encountered in Zone 2 and 298 in Zone 3. A comparison (Dixon and Massey 1968) of the percentages of the total number of plants associated and of the epiphytes showed that in both cases Zone 3 was significantly different from Zone 2; z 2.63 (p < 0.0 1) for the percentage of plants associated and z 2.32 (p < 0.0 1 ) for the percentage of epiphytes, supporting our hypothesis that the degree of epiphytism increased between the second and thrid zone<. A po ssible advantage for facultative epiphytism would be reduced competition for light. ight has been shown by Pittendri� ( 1 948), Veloso and Klein (1 957) and Aragao (1967) to be important in determining bromeliad distribution in ecosystems. Being a = = 1 73 COMUNICACIONES Chavarría and an anonymous reviewer for their constructive criticism of the manuscript. TBLE 1 Comparison 01 rhe prcentage o[ Neoregelia ruenta in arra de Mricá (R. J.) The igures in arentheses refer fa the percentage ofthe rubtotal ZONE 2 Nwner % ZONE 3 Numer LITERATURE CITED % Associated Terrestrial 92 (92) Epiphytic 8( 8) Subtotl 138 (81 .6) 31(18.4) lOO 45.05 122 222 169 56.71 54.95 1 29 43.29 10.0 298 100.0 Not �sociated Terrestrial Toal: A ragao , M. B. Considerayoes de habitat e distribui�o geográfica de algunas Bromeliáceas. Sellowia , 1 9 : 83-95. '.'tank" bromeliad, . cruenta is independent of the soil and receivs nutrients ann water from atmospheric sources. It is also po�sible to speculate on the evolutionary historY of the Bromeliaceae. In this family , the obligatory epiphytes are considered to be the most highly evolved (Smith and Downs 1974) ; thus the capability for farultative epiphytism might represent a transitory phase in the evolutionary history of the family. . In summary, observations on habitat use by . cuen ta show that it is capable of exp loitation of different habitats within the coastal sand dune ecosystem. The mechanism ued is facultative epiphytism, possibly in response to a requirement of high light intensity for germination or rowth. ACKNOWLEDGEMENTS We thank the Conselho Nacional de Desenvolvimento Cientíico e Tecnológico (CNPq) for support (grants 1074 10/79 to LDL and 2222. 1 9 1 5/78 to JDH) of this project. We also thank Drs. D. Benzing, J. Utley , M. Dixon, W.J., & F .J . Massey. 1 968. lntroducion of statistical analysis. McGraw-Hill, New York. Hay, J . D., & L.D . de Lacerda. 1 980. A lterayoes nas caracter ísias do solo aps a fixa�io de Neoregela cuenta R. Graham) L. Smith em'. m ec ossistema de dunas de areia. ienc. e Cult, 32 : 863-867 . Hay , J.D., L.D. d e Laeerda, & A.L. Tan. 1981. Soil eation inerease in a tropical sand dune eeosystem due to a terresrial bromeliad. 'Eeology , 62: 1 392-1 395 . Lacerda, L.D. de & J.D. Hay, 1 977. Importancia e a l g u m a s espécies '{egetais peren.s em um ecossistema de dunas de areia. Cien. e Cult. (Supl. , Resumos), 29: 491492. Medina, E . , 1 974. Dark C02 ixation, habitat p r e f e r e n ce and e v o l u t i on within t he Bromeliaceae, Evolution, 2 8 : 677-i86. Pittendrigh , C., 1 948. The bromeliad-anopheles­ malaria eomplex n Trinidad. I. The bromeliad flora. Evolution, 2 : 5 8-89. Smith, L.B. & R.J. Downs, 1 974. Flora Neotropica. Monoraph NO. 1 4 (Piteairnioidae). Hafner Press, New York. Veloso, H.P., & R .M. Klein, 1 957. As comunidades e aso cia�oes vegetais de mata pluvial do sul do Brasil. I. s comunidades do Município de Brusque, Estado de anta atarina. ellowia, 8 : 81-235.