Plant community structure and function in a swamp forest within the Atlantic rain forest complex: a synthesis

Scarano, Fabio Rubio

doi:10.1590/2175-7860200657308

ABSTRACT

This paper synthesises a decade of research on a swamp forest within the Atlantic forest complex in the state of Rio de Janeiro, Brazil. I propose that this swamp is a fragile ecosystem, since its diversity and functioning are highly dependent on a specific interaction between two functional groups: shade-providers (locally rare trees) and providers of safe germination sites (terrestrial bromeliads). This conclusion is based on a broad set of data regarding plant ecophysiology, population and community ecology and phytogeography, which I review here. I discuss the implications of these findings for conservation and restoration of swamps at the Atlantic forest complex.

Key-words:
clonal growth; epiphytes; functional groups; photosynthesis; swamp forest

RESUMO

Este artigo sintetiza uma década de pesquisas realizadas em um pântano pertencente ao complexo vegetacional atlântico no estado do Rio de Janeiro, Brasil. Eu proponho que este pântano é um ecossistema frágil, na medida em que seu funcionamento e sua diversidade são altamente dependentes da interação específica entre dois grupos funcionais: os geradores de sombra (árvores localmente raras) e os geradores de sítios seguros para germinação (bromélias terrestres). Esta conclusão se baseia num amplo conjunto de dados referentes à ecofisiologia, ecologia de populações e comunidades, e fitogeografia, que são revisados aqui. Eu discuto as implicações destes resultados para a conservação e restauração deste tipo de vegetação.

Palavras-chave:
crescimento clonal; epífitas; estabilidade de ecossistemas; fotossíntese; grupos funcionais; pântano

Full text available only in PDF format.

ACKNOWLEDGEMENTS

Tânia S. Pereira, G. Martinelli, H. C. Lima and C. F. Barros kindly hosted me as a postdoctoral researcher in the Botanical Gardens of Rio de Janeiro from 1992 to 1993, when they introduced me to the beautiful swamp forests of Poço das Antas, and taught me a great deal about them. I also thank my other co-workers and students involved in the studies revised here: E. F. Amado, S. M. B. Barreto, D. D. Biesboer, C. H. Callado, C. G. Costa, H. M. Duarte, A. C. Franco, C. A. Freitas, H. R. P. Lima, U. Lüttge, A. Mantovani, E. A. de Mattos, L. F. D. de Moraes, K. T. Ribeiro, G. Rôças, S. J. Silva Neto and T. Wendt. Finally, I thank the staff at the Biological Reserve of Poço das Antas for support and the Brazilian Research Council (CNPq) for a productivity grant.

REFERENCES

Abrahamson, W. G. 1980. Demography and vegetative reproduction. In: Solbrig, O. T. (ed.). Demography and evolution in plant populations. Blackwell, Oxford. Pp. 89-106.
Barbosa, F. A. R.; Scarano, F. R.; Sabará, M. G. & Esteves, F. A. 2004. Brazilian LTER: Ecosystem and biodiversity information in support of decision-making. Environmental Monitoring and Assessment 90: 121-133.
Bruno, J. F.; Stachowicz, J. J. & Bertness, M. D. 2003. Inclusion of facilitation into ecological theory. Trends in Ecology and Evolution 18: 119-125.
Callado, C. H.; Silva Neto, S. J.; Scarano, F. R.; Barros, C. F. & Costa, C. G. 2001a. Anatomical features of growth rings in flood-prone trees of the Atlantic rain Forest in Rio de Janeiro, Brazil. IAWA Journal 22: 29-42.
Callado, C. H.; Silva Neto, S. J.; Scarano, F. R. & Costa, C. G. 2001b. Periodicity of growth rings in some flood-prone trees of the Atlantic Rain Forest in Rio de Janeiro, Brazil. Trees Structure and Function 15: 492-497.
Callado, C. H.; Silva Neto, S. J.; Scarano, F. R. & Costa, C. G. 2004. Radial growth dynamics of Tabebuia umbellata (Bignoniaceae), a flood-tolerant tree from the Atlantic forest swamps in Brazil. IAWA Journal 25: 175-183.
Callaway, R. M.; Brooker, R. W.; Choler, P.; Kikvidze, Z.; Lortie, C. J.; Michalet, R.; Paolini, L.; Pugnaire, F. I.; Newingham, B.; Aschehoug, E. T.; Armas, C.; Kikodze, D. & Cook, B. J. 2002. Positive interactions among alpine plants increase with stress. Nature 417: 844-848.
Chase, J. M. & Leibold, M. A. 2003. Ecological niches: linking classical and contemporary approaches. The University of Chicago Press, Chicago, 212p.
Cirne, P. & Scarano, F. R. 2001. Resprouting and growth dynamics after fire of the clonal shrub Andira legalis (Leguminosae) in a sandy coastal plain in southeastern Brazil. Journal of Ecology 89: 351-357.
Colwell, R. K. 1974. Predictability, constancy and contingency of periodic phenomena. Ecology 55: 1148-1153.
Cook, R. 1980. The biology of seeds in the soil. In: Solbrig, O. T. (ed.). Demography and evolution in plant populations. Blackwell, Oxford. Pp.107-129.
Crawford, R. M. M. 1992. Oxygen availability as an ecological limit to plant distribution. Advances in Ecological Research 23: 93-185.
Díaz, S.; Symstad, A. J.; Chapin, F. S. III; Wardle, D. A. & Huenneke, L. F. 2003. Functional diversity revealed by removal experiments. Trends in Ecology and Evolution 18: 140-146.
Dorneles, L. P. P. & Waechter, J. 2004. Fitossociologia do componente arbóreo na floresta turfosa do Parque Nacional da Lagoa do Peixe, Rio Grande do Sul, Brasil. Acta Botanica Brasilica 18: 815-824.
Fargione, J. & Tilman, D. 2002. Competition and coexistence in terrestrial plants. In: Sommer, U. & Worm, B. (eds.). Competition and coexistence. Springer, Berlin. Pp. 165-206.
Freitas, C. A.; Scarano, F. R. & Wendt, T. 1998. Habitat choice in two facultative epiphytes of the genus Nidularium (Bromeliaceae). Selbyana 19: 236-239.
Freitas, C. A. de; Scarano, F. R. & Biesboer, D. D. 2003. Morphological variation in two facultative epiphytic bromeliads growing on the floor of a swamp forest. Biotropica 35: 546-550.
Grime, J. P. 2001. Plant strategies, vegetation processes, and ecosystem properties. 2nd. ed. John Wiley, New York, 417p.
Guedes-Bruni, R. R.; Silva Neto, S. J.; Morim, M. P. & Mantovani, W. 2006. Composição florística e estrutura de trecho de floresta atlântica sobre planície aluvial na Reserva Biológica de Poço das Antas, Silva Jardim. Rodriguesia 57(3): 413-428.
Gunderson, L. H. 2000. Ecological resilience - in theory and application. Annual Review of Ecology and Systematics 31: 425-439.
Hendry, G. A. F. & Crawford, R. M. M. 1994. Oxygen and environmental stress in plants - an overview. Proceedings of the Royal Society of Edinburgh 102B: 1-10.
Hubbell, S. P. 2001. The unified neutral theory of biodiversity and biogeography. Princeton University Press, Princeton, 375p.
IBDF. 1981. Plano de manejo - Reserva Biológica de Poço das Antas. Brasília, 59p.
Ives, A. R. 1995. Measuring resilience in stochastic systems. Ecological Monographs 65: 217-233.
Joly, C. A. 1990. Flooding tolerance in tropical trees. In: Jackson, M. B., Davies, D. D. & Lambers, H. (eds.). Plant life under oxygen deprivation. SPB Academic, The Hague. Pp. 23-34.
Junk, W. J. 1993. Wetlands of tropical South America. In: Whigham, D. F., Dykyjová, D. & Hejný, S. (eds.). Wetlands of the world I: inventory, ecology and management. Kluwer, Dordrecht. Pp. 679-739.
Kozlowski, T. T. 1984. Response of woody plants to flooding. In: Kozlowski, T. T. (ed.). Flooding and plant growth. Academic Press, San Diego. Pp. 129-163.
Leitão-Filho, H. F. 1993. Ecologia da Mata Atlântica em Cubatão. Editora Unesp, Editora Unicamp, São Paulo, 184p.
Lobo, P. C. & Joly, C. A. 1998. Tolerance to hypoxia and anoxia in neotropical tree species. In: Scarano, F. R. & Franco A. C. (eds.). Plant survival strategies in xerophytic and amphibious habitats in the Neotropics. Oecologia Brasiliensis Series, volume 4. PPGE-UFRJ, Rio de Janeiro, Brazil. Pp. 137-156.
MacArthur, R. H. & Wilson, E. O. 1967. The theory of island biogeography. Princeton University Press, Princeton, 208p.
Metzger, J. P. 2003. Como restaurar a conectividade de paisagens fragmentadas? In: Kageyama, P. Y.; Oliveira, R. E.; Moraes, L. F. D.; Engel, V. L. & Gandara, F. B. (eds.). Restauração ecológica de ecossistemas naturais. Fundação de Estudos e Pesquisas Agrícolas Florestais, Botucatu. Pp. 49-76.
Moraes, L. F. D. & Pereira, T. S. 2003. Restauração ecológica em unidades de conservação. In: Kageyama, P. Y.; Oliveira, R. E.; Moraes, L. F. D.; Engel, V. L. & Gandara, F. B. (eds.). Restauração ecológica de ecossistemas naturais. Fundação de Estudos e Pesquisas Agrícolas Florestais, Botucatu. Pp. 295-305.
Morellato, L. P. C. & Haddad, C. F. B. 2000. Introduction: The Brazilian Atlantic Forest. Biotropica 32: 786-792.
Myers, N.; Mittermeier, R. A.; Mittermeier, C. G.; Fonseca, G. A. B. & Kent, J. 2000. Biodiversity hotspots for conservation priorities. Nature 403: 853-858.
Oliveira-Filho, A. T. & Fontes, M. A. L. 2000. Patterns of floristic differentiation among atlantic forests in southeastern Brazil and the influence of climate. Biotropica 32: 793-810.
Parolin, P.; De Simone, O.; Haase, K.; Waldhoff, D.; Rottenberger, S.; Kuhn, U.; Kesselmeier, J.; Kleiss, B.; Schmidt, W.; Piedade, M. T. F. & Junk, W. J. 2004. Central Amazonian floodplain forests: tree adaptations in a pulsing system. Botanical Review 70: 357-380.
Pillar, V. D. 1999. On the identification of optimal functional types. Journal of Vegetation Science 10: 631-640.
Pillar, V. D. & Sosinski, E. E. Jr. 2003. An improved method for searching plant functional types by numerical analysis. Journal of Vegetation Science 14: 323-332.
Pimm, S. L. 1991. The balance of nature? Ecological issues in the conservation of species and communities. Chicago, The University of Chicago Press, 434p.
Pott A. 2000. Dinâmica da vegetação do Pantanal. In: Cavalcanti, T. B. & Walter B. M. T. (eds.). Tópicos Atuais em Botânica. Sociedade Botânica do Brasil/Embrapa, Brasília. Pp. 172-175.
Rees, M. 1994. Delayed germination of seeds: a look at the effect of adult longevity, the timing of reproduction and population age/stage structure. American Naturalist 144: 43-64.
Rizzini, C. T. 1979. Tratado de fitogeografia do Brasil. 2^a ed. Âmbito Cultural Edições, Rio de Janeiro, 747p.
Rôças, G.; Scarano, F. R. & Barros, C. F. 2001. Leaf anatomical variation in Alchornea triplinervia (Spreng.) Müll. Arg. (Euphorbiaceae) under different light and soil-water regimes. Botanical Journal of the Linnean Society 136: 231-238.
Scarano, F. R. 2002. Structure, function and floristic relationships of plant communities in stressful habitats marginal to the Brazilian Atlantic rain forest. Annals of Botany 90: 517-524.
Scarano, F. R. & Dias, A. T. C. 2004. A importância de espécies no funcionamento de comunidades e ecossistemas. In: Coelho, A. S.; Loyola, R. D. & Souza, M. B. G. (eds.). Ecologia teórica: desafios para o aperfeiçoamento da Ecologia no Brasil. O Lutador, Belo Horizonte. Pp. 43-60.
Scarano, F. R.; Rios, R. I. & Esteves, F. A. 1998. Tree species richness, diversity and flooding regime: case studies of recuperation after anthropic impact in Brazilian flood-prone forests. International Journal of Ecology and Environmental Sciences 24: 223-235.
Scarano, F. R.; Ribeiro, K. T.; Moraes, L. F. D. & Lima, H. C. 1997. Plant establishment on flooded and unflooded patches of a freshwater swamp forest in southeastern Brazil. Journal of Tropical Ecology 14: 793-803.
Scarano, F. R.; Mattos, E. A.; Franco, A. C.; Herzog, B.; Ball, E.; Grams, T. E. E.; Mantovani, A.; Barreto, S.; Haag-Kerwer, A. & Lüttge, U. 1999. Habitat segregation of C3 and CAM Nidularium (Bromeliaceae) in response to different light regimes in the understory of a swamp forest in southeastern Brazil. Flora 194: 281-288.
Scarano, F. R.; Duarte, H. M.; Ribeiro, K. T.; Rodrigues, P. J. F. P.; Barcellos, E. M. B.; Franco, A. C.; Brulfert, J.; Deléens, E. & Lüttge, U. 2001. Four sites with contrasting environmental stress in southeastern Brazil: relations of species, life form diversity, and geographical distribution to ecophysiological parameters. Botanical Journal of the Linnean Society 136: 345-364.
Shrader-Frechette, K. S. & McCoy, E. D. 1993. Method in ecology. Cambridge University Press, Cambridge, 328p.
Smith, T. M.; Shugart, H. H. & Woodward, F. I. (eds.). 1997. Plant functional types. Cambridge University Press, Cambridge, 369p.
Tilman, D.; Knops, J.; Wedin, D. & Reich, P. 2001. Experimental and observational studies of diversity, productivity and stability. In: Kinzig, A. P.; Pacala, S. W. & Tilman, D. (eds.). The functional consequences of biodiversity: empirical progress and theoretical extensions. Princeton University Press, Princeton. Pp. 42-70.
WCMC-World Conservation Monitoring Centre. 1992. Global biodiversity: status of the Earth´s living resources. Chapman & Hall, London, 594p.
Wilson, J. B. 1999. Guilds, functional types and ecological groups. Oikos 86: 507-522.

Publication Dates

Publication in this collection
Sep-Dec 2006

History

Received
Feb 2005
Accepted
May 2006

This is an Open Access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

[1] Abrahamson, W. G. 1980. Demography and vegetative reproduction. In: Solbrig, O. T. (ed.). Demography and evolution in plant populations. Blackwell, Oxford. Pp. 89-106.

[2] Barbosa, F. A. R.; Scarano, F. R.; Sabará, M. G. & Esteves, F. A. 2004. Brazilian LTER: Ecosystem and biodiversity information in support of decision-making. Environmental Monitoring and Assessment 90: 121-133.

[3] Bruno, J. F.; Stachowicz, J. J. & Bertness, M. D. 2003. Inclusion of facilitation into ecological theory. Trends in Ecology and Evolution 18: 119-125.

[4] Callado, C. H.; Silva Neto, S. J.; Scarano, F. R.; Barros, C. F. & Costa, C. G. 2001a. Anatomical features of growth rings in flood-prone trees of the Atlantic rain Forest in Rio de Janeiro, Brazil. IAWA Journal 22: 29-42.

[5] Callado, C. H.; Silva Neto, S. J.; Scarano, F. R. & Costa, C. G. 2001b. Periodicity of growth rings in some flood-prone trees of the Atlantic Rain Forest in Rio de Janeiro, Brazil. Trees Structure and Function 15: 492-497.

[6] Callado, C. H.; Silva Neto, S. J.; Scarano, F. R. & Costa, C. G. 2004. Radial growth dynamics of Tabebuia umbellata (Bignoniaceae), a flood-tolerant tree from the Atlantic forest swamps in Brazil. IAWA Journal 25: 175-183.

[7] Callaway, R. M.; Brooker, R. W.; Choler, P.; Kikvidze, Z.; Lortie, C. J.; Michalet, R.; Paolini, L.; Pugnaire, F. I.; Newingham, B.; Aschehoug, E. T.; Armas, C.; Kikodze, D. & Cook, B. J. 2002. Positive interactions among alpine plants increase with stress. Nature 417: 844-848.

[8] Chase, J. M. & Leibold, M. A. 2003. Ecological niches: linking classical and contemporary approaches. The University of Chicago Press, Chicago, 212p.

[9] Cirne, P. & Scarano, F. R. 2001. Resprouting and growth dynamics after fire of the clonal shrub Andira legalis (Leguminosae) in a sandy coastal plain in southeastern Brazil. Journal of Ecology 89: 351-357.

[10] Colwell, R. K. 1974. Predictability, constancy and contingency of periodic phenomena. Ecology 55: 1148-1153.

[11] Cook, R. 1980. The biology of seeds in the soil. In: Solbrig, O. T. (ed.). Demography and evolution in plant populations. Blackwell, Oxford. Pp.107-129.

[12] Crawford, R. M. M. 1992. Oxygen availability as an ecological limit to plant distribution. Advances in Ecological Research 23: 93-185.

[13] Díaz, S.; Symstad, A. J.; Chapin, F. S. III; Wardle, D. A. & Huenneke, L. F. 2003. Functional diversity revealed by removal experiments. Trends in Ecology and Evolution 18: 140-146.

[14] Dorneles, L. P. P. & Waechter, J. 2004. Fitossociologia do componente arbóreo na floresta turfosa do Parque Nacional da Lagoa do Peixe, Rio Grande do Sul, Brasil. Acta Botanica Brasilica 18: 815-824.

[15] Fargione, J. & Tilman, D. 2002. Competition and coexistence in terrestrial plants. In: Sommer, U. & Worm, B. (eds.). Competition and coexistence. Springer, Berlin. Pp. 165-206.

[16] Freitas, C. A.; Scarano, F. R. & Wendt, T. 1998. Habitat choice in two facultative epiphytes of the genus Nidularium (Bromeliaceae). Selbyana 19: 236-239.

[17] Freitas, C. A. de; Scarano, F. R. & Biesboer, D. D. 2003. Morphological variation in two facultative epiphytic bromeliads growing on the floor of a swamp forest. Biotropica 35: 546-550.

[18] Grime, J. P. 2001. Plant strategies, vegetation processes, and ecosystem properties. 2nd. ed. John Wiley, New York, 417p.

[19] Guedes-Bruni, R. R.; Silva Neto, S. J.; Morim, M. P. & Mantovani, W. 2006. Composição florística e estrutura de trecho de floresta atlântica sobre planície aluvial na Reserva Biológica de Poço das Antas, Silva Jardim. Rodriguesia 57(3): 413-428.

[20] Gunderson, L. H. 2000. Ecological resilience - in theory and application. Annual Review of Ecology and Systematics 31: 425-439.

[21] Hendry, G. A. F. & Crawford, R. M. M. 1994. Oxygen and environmental stress in plants - an overview. Proceedings of the Royal Society of Edinburgh 102B: 1-10.

[22] Hubbell, S. P. 2001. The unified neutral theory of biodiversity and biogeography. Princeton University Press, Princeton, 375p.

[23] IBDF. 1981. Plano de manejo - Reserva Biológica de Poço das Antas. Brasília, 59p.

[24] Ives, A. R. 1995. Measuring resilience in stochastic systems. Ecological Monographs 65: 217-233.

[25] Joly, C. A. 1990. Flooding tolerance in tropical trees. In: Jackson, M. B., Davies, D. D. & Lambers, H. (eds.). Plant life under oxygen deprivation. SPB Academic, The Hague. Pp. 23-34.

[26] Junk, W. J. 1993. Wetlands of tropical South America. In: Whigham, D. F., Dykyjová, D. & Hejný, S. (eds.). Wetlands of the world I: inventory, ecology and management. Kluwer, Dordrecht. Pp. 679-739.

[27] Kozlowski, T. T. 1984. Response of woody plants to flooding. In: Kozlowski, T. T. (ed.). Flooding and plant growth. Academic Press, San Diego. Pp. 129-163.

[28] Leitão-Filho, H. F. 1993. Ecologia da Mata Atlântica em Cubatão. Editora Unesp, Editora Unicamp, São Paulo, 184p.

[29] Lobo, P. C. & Joly, C. A. 1998. Tolerance to hypoxia and anoxia in neotropical tree species. In: Scarano, F. R. & Franco A. C. (eds.). Plant survival strategies in xerophytic and amphibious habitats in the Neotropics. Oecologia Brasiliensis Series, volume 4. PPGE-UFRJ, Rio de Janeiro, Brazil. Pp. 137-156.

[30] MacArthur, R. H. & Wilson, E. O. 1967. The theory of island biogeography. Princeton University Press, Princeton, 208p.

[31] Metzger, J. P. 2003. Como restaurar a conectividade de paisagens fragmentadas? In: Kageyama, P. Y.; Oliveira, R. E.; Moraes, L. F. D.; Engel, V. L. & Gandara, F. B. (eds.). Restauração ecológica de ecossistemas naturais. Fundação de Estudos e Pesquisas Agrícolas Florestais, Botucatu. Pp. 49-76.

[32] Moraes, L. F. D. & Pereira, T. S. 2003. Restauração ecológica em unidades de conservação. In: Kageyama, P. Y.; Oliveira, R. E.; Moraes, L. F. D.; Engel, V. L. & Gandara, F. B. (eds.). Restauração ecológica de ecossistemas naturais. Fundação de Estudos e Pesquisas Agrícolas Florestais, Botucatu. Pp. 295-305.

[33] Morellato, L. P. C. & Haddad, C. F. B. 2000. Introduction: The Brazilian Atlantic Forest. Biotropica 32: 786-792.

[34] Myers, N.; Mittermeier, R. A.; Mittermeier, C. G.; Fonseca, G. A. B. & Kent, J. 2000. Biodiversity hotspots for conservation priorities. Nature 403: 853-858.

[35] Oliveira-Filho, A. T. & Fontes, M. A. L. 2000. Patterns of floristic differentiation among atlantic forests in southeastern Brazil and the influence of climate. Biotropica 32: 793-810.

[36] Parolin, P.; De Simone, O.; Haase, K.; Waldhoff, D.; Rottenberger, S.; Kuhn, U.; Kesselmeier, J.; Kleiss, B.; Schmidt, W.; Piedade, M. T. F. & Junk, W. J. 2004. Central Amazonian floodplain forests: tree adaptations in a pulsing system. Botanical Review 70: 357-380.

[37] Pillar, V. D. 1999. On the identification of optimal functional types. Journal of Vegetation Science 10: 631-640.

[38] Pillar, V. D. & Sosinski, E. E. Jr. 2003. An improved method for searching plant functional types by numerical analysis. Journal of Vegetation Science 14: 323-332.

[39] Pimm, S. L. 1991. The balance of nature? Ecological issues in the conservation of species and communities. Chicago, The University of Chicago Press, 434p.

[40] Pott A. 2000. Dinâmica da vegetação do Pantanal. In: Cavalcanti, T. B. & Walter B. M. T. (eds.). Tópicos Atuais em Botânica. Sociedade Botânica do Brasil/Embrapa, Brasília. Pp. 172-175.

[41] Rees, M. 1994. Delayed germination of seeds: a look at the effect of adult longevity, the timing of reproduction and population age/stage structure. American Naturalist 144: 43-64.

[42] Rizzini, C. T. 1979. Tratado de fitogeografia do Brasil. 2^a ed. Âmbito Cultural Edições, Rio de Janeiro, 747p.

[43] Rôças, G.; Scarano, F. R. & Barros, C. F. 2001. Leaf anatomical variation in Alchornea triplinervia (Spreng.) Müll. Arg. (Euphorbiaceae) under different light and soil-water regimes. Botanical Journal of the Linnean Society 136: 231-238.

[44] Scarano, F. R. 2002. Structure, function and floristic relationships of plant communities in stressful habitats marginal to the Brazilian Atlantic rain forest. Annals of Botany 90: 517-524.

[45] Scarano, F. R. & Dias, A. T. C. 2004. A importância de espécies no funcionamento de comunidades e ecossistemas. In: Coelho, A. S.; Loyola, R. D. & Souza, M. B. G. (eds.). Ecologia teórica: desafios para o aperfeiçoamento da Ecologia no Brasil. O Lutador, Belo Horizonte. Pp. 43-60.

[46] Scarano, F. R.; Rios, R. I. & Esteves, F. A. 1998. Tree species richness, diversity and flooding regime: case studies of recuperation after anthropic impact in Brazilian flood-prone forests. International Journal of Ecology and Environmental Sciences 24: 223-235.

[47] Scarano, F. R.; Ribeiro, K. T.; Moraes, L. F. D. & Lima, H. C. 1997. Plant establishment on flooded and unflooded patches of a freshwater swamp forest in southeastern Brazil. Journal of Tropical Ecology 14: 793-803.

[48] Scarano, F. R.; Mattos, E. A.; Franco, A. C.; Herzog, B.; Ball, E.; Grams, T. E. E.; Mantovani, A.; Barreto, S.; Haag-Kerwer, A. & Lüttge, U. 1999. Habitat segregation of C3 and CAM Nidularium (Bromeliaceae) in response to different light regimes in the understory of a swamp forest in southeastern Brazil. Flora 194: 281-288.

[49] Scarano, F. R.; Duarte, H. M.; Ribeiro, K. T.; Rodrigues, P. J. F. P.; Barcellos, E. M. B.; Franco, A. C.; Brulfert, J.; Deléens, E. & Lüttge, U. 2001. Four sites with contrasting environmental stress in southeastern Brazil: relations of species, life form diversity, and geographical distribution to ecophysiological parameters. Botanical Journal of the Linnean Society 136: 345-364.

[50] Shrader-Frechette, K. S. & McCoy, E. D. 1993. Method in ecology. Cambridge University Press, Cambridge, 328p.

[51] Smith, T. M.; Shugart, H. H. & Woodward, F. I. (eds.). 1997. Plant functional types. Cambridge University Press, Cambridge, 369p.

[52] Tilman, D.; Knops, J.; Wedin, D. & Reich, P. 2001. Experimental and observational studies of diversity, productivity and stability. In: Kinzig, A. P.; Pacala, S. W. & Tilman, D. (eds.). The functional consequences of biodiversity: empirical progress and theoretical extensions. Princeton University Press, Princeton. Pp. 42-70.

[53] WCMC-World Conservation Monitoring Centre. 1992. Global biodiversity: status of the Earth´s living resources. Chapman & Hall, London, 594p.

[54] Wilson, J. B. 1999. Guilds, functional types and ecological groups. Oikos 86: 507-522.

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