Small mammal species diversity in brazilian tropical primary and secondary forests of different sizes

Fonseca, Gustavo A. B. da

doi:10.1590/S0101-81751989000300001

Abstract

This study investigated the effects of forest fragment size and habitat structure on species richness and diversity of Brazilian Atlantic forest small mammal communities. During the course of 17 months of live-trapping, a total of 57, 120 trap nights accumulated in six forest plots. There were three size categories: small (60-80 ha.), medium (860 ha) and large (35, 973 ha). At each site two forest sections were selected for small mammal sampling: one in primary and another in secondary succession stage. A total of 692 individuals belonging to 19 species were trapped 1, 366 times. Species diversity was measured using the Shannon-Wiener index H' and also a weighted version of this index that takes into account the number of individuals of each species trapped at the forests. The modified index was developed specifically for this study. The richest and most diverse forest for small mammals was the large secondary fragment, while the least diverse was the small mammal community of the small primary forest. Overall, species diversity is lower among primary forests, and also in forests of decreasing size. However, area sizel species relationship was only observed among secondary forests. It is postulated that primary forests are less complex habitats, when compared to secondary habitats in mid-stages of succession. Therefore, area size/species relationships should only be applied to conservation schemes if habitat structure is taken into account. By using multivariate regression methods in the investigation of the structural and size predictors of small mammal species diversity, it was possible to separate the effects of area size from those of habitat structure. Approximately half of the variance in small mammal species diversity between secondary forest fragments can be explained by area size, while habitat structure accounts for about 45% of the variation.

Small mammal species diversity in brazilian tropical primary and secondary forests of different sizes

Gustavo A. B. da Fonseca

Departamento de Zoologia, Instituto de Ciências Biológicas, Universidade Federal de Minas Gerais. 30000, Belo Horizonte MG Brasil

ABSTRACT

This study investigated the effects of forest fragment size and habitat structure on species richness and diversity of Brazilian Atlantic forest small mammal communities. During the course of 17 months of live-trapping, a total of 57, 120 trap nights accumulated in six forest plots. There were three size categories: small (60-80 ha.), medium (860 ha) and large (35, 973 ha). At each site two forest sections were selected for small mammal sampling: one in primary and another in secondary succession stage. A total of 692 individuals belonging to 19 species were trapped 1, 366 times. Species diversity was measured using the Shannon-Wiener index H' and also a weighted version of this index that takes into account the number of individuals of each species trapped at the forests. The modified index was developed specifically for this study. The richest and most diverse forest for small mammals was the large secondary fragment, while the least diverse was the small mammal community of the small primary forest. Overall, species diversity is lower among primary forests, and also in forests of decreasing size. However, area sizel species relationship was only observed among secondary forests. It is postulated that primary forests are less complex habitats, when compared to secondary habitats in mid-stages of succession. Therefore, area size/species relationships should only be applied to conservation schemes if habitat structure is taken into account. By using multivariate regression methods in the investigation of the structural and size predictors of small mammal species diversity, it was possible to separate the effects of area size from those of habitat structure. Approximately half of the variance in small mammal species diversity between secondary forest fragments can be explained by area size, while habitat structure accounts for about 45% of the variation.

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ACKNOWLEDGEMENTS

First and foremost, I would like to thank John G. Robinson for his dedication to my work. His advice during planning, field work, analysis and writing of this paper were invaluable. John Eisenberg, Larry Harris, Nigel Smith, Melvin Sunquist, and Charles Woods provided support in the planning of this study and improvements in earlier drafts. This research also benefited from discussions with Thomas Lacher, Kent Redford, Russell Mittermeier and Michael Mares. Two other reviewers offered valuable suggestions, some of them adopted here.

Celio Valle encouraged the field work with his enthusiasm, vision, and eternal optimism. Several people helped during field work: Ludmilla Aguiar, Ilmar Bastos, Sonia Rigueira, Carlos Alberto Pinto, Ederson Machado (sometimes), Silverio Machado, Ney Carnevalli, Luiz Fernando Mello, Gisela Hermann, Jairo Vieira, Eduardo Veado, Eduardo Sabato, Luiz Paulo Pinto, and Maria Cristina Alves. My small mammal data from the Rio Doce Park was partially collected by Jody Stallings and his crew.

Mr. Feliciano Abdalla and Dr. Antonio Cupertino kindly allowed me to work on their farms and provided housing. "Santinho" let us stay in his house at the expense of family problems, and I am thankful to him. The State Forest Institute of Minas Gerais (IEF) provided me with accommodations and gasoline at the Rio Doce State Park. I am very grateful to the staff of the Park, especially Ademir Camara Lopes, Jose Lourenço Ladeira and Hermogenes Ferreira Neto. M. Carleton, K. Creighton, L. Emmons, G. Musser, P. Myers, and J. Patton generously identified the small mammal voucher specimens.

The World Wildlife Fund-US financed the most substantial part of the costs involved in this study. Additional financial support was provided by the Program for Studies in Tropical Conservation, of the University of Florida, and by the Research Council of the Federal University of Minas Gerais, Brazil. The National Research Council of Brazil (CNPq) awarded me with a doctoral fellowship.

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CONNELL, J. H. & R. O. SLATYER. 1977. Mechanisms of sucession in natural communities and their role in community stability and organization. Am. Nat. 111:1119-1144.
DARLINGTON, P. J. 1957. Zoogeography: the Geographical Distribution of Animals. Wiley and Sons, London.
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DAVIS, D. E. 1947. Notes on the life histories of some Brazilian mammals. Bolm Mus. Nac 76:1-8.
DELANY, M. J. 1986. Ecology of small rodents in Africa. Mammal Rev. 16(1):1-41.
DENSLOW, J. S. 1985. Disturbance-mediated coexistence of species. In: S. T. A. PICKETT & P. S. WHITE (eds.), The Ecology of Natural Disturbance and Patch Dynamics. Academic Press, New York.
DIAMOND, J. M. 1975. The island dilema: lessons of modern biogeographical studies for the design of natural reserves. Biol. Conserv. 7:129-146.
DUESER, R. D. & W. C. BROWN. 1980. Ecological correlates of insular rodent diversity. Ecology 61:50-56.
DUESER, R. D. & J. H. PORTER. 1986. Habitat use by insular small mammals: relative effects of competition and habitat structure. Ecology 67(1):195-201.
DUESER, R. D. & H. H. SHUGART. 1978. Microhabitats in a forest-floor small mammal fauna. Ecology 59(1):89-97.
EISENBERG, J. F. 1980. The density and biomass of tropical mammals. In: M. SOULE & B. WILCOX (eds.), Conservation Biology: An Evolutionary-Ecological Perspective. Sinauer Press, Sunderland, Massachussets.
EITEN, G. 1974. An outline of the vegetation of South America. Symp. Congr. Int. Primatol. Soc. 5^th, 529-545.
EMMONS, L. H. 1984. Geographical variation in densities and diversities of non-flying mammals in Amazonia. Biotropica 16(3):210-222.
EMMONS, L. H., A. GAUTIER-HION & G. DUBOST. 1983. Community structure of the frugivorous-folivorous forest mammals of Gabon. J. Zool., Lond. 199:209-222.
FLEMING, T. H. 1971. Population ecology of three species of neotropical rodents. Misc. Publ. Mus. Zool. Univ. Michigan 143:1-77.
FLEMING, T. H. 1975a. The population ecology of two species of Costa Rican heteromyid rodents. Ecology 55:493-510.
FONSECA, G. A. B. 1983. The role of deforestation and private reserves in me conservation of the woolly spider monkey (Brachyteles arachnoides). Masters Thesis, University of Florida, Gainesville, Florida.
FONSECA, G. A. B. 1985. The vanishing Brazilian Atlantic forest. Biol. Conserv. 34(1):17-34.
FONSECA, G. A. B. 1988. Patterns of small mammal species diversity in the Brazilian Atlantic Forest. Ph.D. Dissertation, University of Florida, Gainesville, Florida.
FONSECA, G. A. B., L, F. B. MELLO & G. HERRMANN. 1987. Inventariamento de mamíferos e estudo dos padrões de diversidade de espécies de pequenos mamíferos na EPDA de Peti-Minas Gerais. Report to CEMIG, Belo Horizonte, MG.
FONSECA, G. A. B. & K. R. REDFORD. 1984. The mammals of IBGE's ecological reserve and an analysis of the role of gallery forests in increasing diversity. Revta bras. Biol., 44:517-523.
GLITZENSTEIN, J. S., PH. HARCOMBE & D. R. STRENG. 1986. Disturbance, succession, and maintenance of species diversity in an east Texas forest. Ecol. Monogr. 56(3)243-258.
HAIR, J. D. 1982. measurement of ecological diversity. In: S. D. SCHEMNITZ (ed.), Wildlife Management Techiques Manual. The Wildlife Society, Washington, D. C.
HALLETT, J. G., M. A. O'CONNELL & R. L. HONEYCUTT. 1983. Competition and habitat selection: test of a theory using small mammals. Oikos 40:175-181.
HARRIS, L. D. 1984. The Fragmented Forest, Island Biogeography Theory and the Preservation of Biotic Diversity. University of Chicago Press, Chicago.
HORN, H. S. 1974. The ecology of secondary succession. Ann. Rev. Ecol. Syst. 1974:25-37.
HUECK, K. 1972. As Florestas da América do Sul. Editora Polígono, São Paulo, Brasil.
HUNTLY, N. & R. S. INOUYE. 1987 Small mammal populations of an old-field chronosequence: successional patterns and associations with vegetation. J. Mamm. 68(4):739-745.
ISABIRYE-BASUTA, G. & J. M. KASENENE. 1987. Small rodent populations in selectively felled and mature tracts of Kibale forest, Uganda. Biotropica 19(3):260-266.
JAMES, F. C. & N. O. WAMER. 1982. Relationships between temperate forest bird communities and vegetation structure. Ecology 63:159-171.
JOHNS, A. D. & J. P. SKORUPA. 1987. Responses of rain-forest primates to habitat disturbance: a review. Int. J. Primatol. 8:157-191,
JOHNSON, N. K. 1975. Control of number of bird species on montane islands in the Great Basin. Evolution 29:545-567.
KACHIGAN, S. K. 1982. Multivariate Statiscal Analysis. Radius Press, New York.
KARR, J. R. 1982. Avian extinction on Barro Colorado Island: A Reassessment. Amer. Nat. 119(2):220-239.
LACHER, T. E. & C. J. R. ALHO. in press. Small mammal communities in the Brazilian Pantanal: Population densities, microhabitat affinities and species interactions. J. Mammalogy.
LACHER, T. E., M. A. MARES & C. J. R. ALHO in press. The structure of a small mammal community in a central Brazilian savanna. In: K. R. REDFORD (ed.), Mammals of the Americas: Essays in Honor of Ralph M. Wetzel. University of Chicago Press, Chicago.
LAEMMERT, H. W., L. C. FERREIRA & R. M. TAYLOR. 1946. Investigation of Vertebrate Hosts and Arthropod Vectors. Am. J. Trop. Med. 26(suppl.):23-69.
LEIGH, E. G. 1975. Population fluctuations, community stability, and environmental variability. In: M. L. CODY & J. M. DIAMOND (eds.), Ecology and Evolution of Communities. Harvard University Press, Harvard.
LOMOLINO, M. V. 1984. Mammalian island biogeography: effects of area, isolation and vagility. Oecologia 61:376-382.
LOVEJOY, T. E., R. O. BIERREGAARD, A. B. RYLANDS, J. R. MALCOM, C. E. QUINTELA, L. H. HARPER, K. S. BROWN, A. H. POWELL, G. V. N. POWELL, H. O. R. SCHUBART & M. B. HAYS. 1986. Edge and other effects of isolation on Amazon Forest fragments. In: M. E. SOULE (ed.), Conservation Biology. Sinauer Press, Sunderland, Massachussets.
LYNCH, J. F. & D. F. WHIGHAM. 1984. Effects of forest fragmentation on breeding bird communities in Maryland, USA. Biol. Conserv. 28:287-324.
MacARTHUR, R. H. 1957. On the relative abundance of bird species. Proc. Nat. Acad. Sci. 43:293-295.
MacARTHUR, R. H. 1972. Geographical Ecology. Princeton University Press, Princeton.
MacARTHUR, R. H. & E. O. WILSON. 1967. The Theory of Island Biogeography. Princeton University Press, Princeton.
MALCOLM, J. 1987. Small mammal abundances in isolated and non-isolated primary forest reserves near Manaus, Brasil. In: Resumos do Congresso Brasileiro de Zoologia, Juiz de Fora, Minas Gerais, Brasil
MARGULES, C. R. & M. B. USHER. 1981. Criteria used in assessing wildlife conservation potential: a review. Biol. Conserv. 21:79-109.
MAY, R. M. 1975. Patterns of species abundance and diversity. In: M. L. CODY & J. M. DIAMOND (eds.), Ecology and Evolution of Communities. Harvard Univ. Press, Harvard, Massachussets.
M'CLOSKEY, R. T. 1976. Community structure in sympatic rodents. Ecology 57:728-739.
MILES, M. A., A. A. de SOUZA & M. M. POVOA. 1981. mammal tracking and nest location in Brazilian forest with an improved spool-and-line device. J. Zool. Lond. 195:331-347.
MITTERMEIER, R. A., A. F. COIMBRA-FILHO, I. D. CONSTABLE, A. B. RY LANDS & C. VALLE. 1982. Conservation of primates in the Atlantic forests of Eastern Brazil. Int. Zoo. Yearbook, 1982.
MOOJEN, J. 1952, Os Roedores do Brasil. Instituto Nacional do Livro, Rio de Janeiro, Brazil.
MULLER, P. 1973. The Dispersal Centers of Terrestrial Vertebrates in the Neotropical Realm. W. Junk Publishers, The Hague.
NITIKMAN, L. Z. & M. A. MARES. 1987. Ecology of small mammals in a gallery forest of Central Brazil. Ann. Carn. Mus. 56:75-95.
PATIL, G. P. & C. TAILLIE. 1983. Diversity as a concent and its measurement. Trans. North. Amer. Wildl. Nat. Resour. Conf. 39:334-353.
PEET, R. K. 1974. The measurement of species diversity. Ann. Rev. Ecol. Syst. 1974:285-307.
PIELGU, E. C. 1974. Population and Community Ecology, Gordon and Breach Publishers, New York.
POOLE, R. H. 1974. An Introduction to Quantitative Ecology. McGraw-Hill, New York,
PRANCE, G. T. 1982. Biological Diversification in the Tropics. Columbia University Press, New York.
QUINN, J. F. & A. HASTINGS. 1987. Extinction in subdivided habitats. Conserv. Biol. 1:198-208.
ROSENZWEIG, M. L. & J. WINAKUR. 1969. Population ecology of desert rodent communities: habitats and environmental complexity. Ecology 50:558-572.
SHAFFER, M. L. 1981. Minimum population sizes for species conservation. Bioscience 31:131-134.
SIMBERLOFF, D. S. 1976. Experimental zoogeography of islands: effects of island size. Ecology 57:629-578.
SIMBERLOFF, D. S. 1982. Big advantages of small refuges. Nat. Hist. 91:6-13.
SIMBERLOFF, D. S. & L. G. ABELE. 1976. Island biogeography theory and conservation pratice. Science 191:285-286.
SIMBERLOFF, D. S. & L. G. ABELE. 1982. Refuge design and island biogeographic theory: effects of fragmentation. Amer. Nat. 120:41-50.
SOKAL, R. R. & ROHLF, F. J. 1981. Biometry. W. H. Freeman Co., New York.
SOULE, M. E. B. 1980. Thresholds for survival: maintaining fitness and evolutionary potential. In: M. E. SOULE & B. A. WILCOX (eds.), Conservation Biology. Sinauer Press, Sunderland, Massachussets.
SOULE, M. E. B. 1983. What do we really know about extinction? In: C. M. SCHONEWALD-WILCOX, S. M. CHAMBERS, B. MACBRYDE & L. THOMAS (eds.), Genetics and Conservation. Benjamin Cummings Co., Menlo Park, California.
SOUZA, W. P. 1984. The role of disturbance in natural communities. Ann. Rev. Ecol. Syst. 15:353-391.
STALLINGS, J. R. 1988. Small mammal comunities of an eastern Brazilian park. Ph. D. Dissertation, University of Florida, Gainesville, Florida.
USHER, M. B. 1986. Wildlife Conservation Evaluation. Chapman and Hall, New York,
WEAVER, M. & M. KELLMAN. 1981. The effects of forest fragmentation on woodlot tree biotas in Southern Ontario. J. Biogeogr. 8:199-210.
WESTMAN, W. E. 1983. Island biogeography: studies on the xeric shrublands of the inner Channel Islands, California. J. Biogeogr. 10:96-118.
WILLIAMS, C. B. 1964. Patterns in the Balance of Nature. Academic Press, New York.
WILLIS, E. O. 1979. The composition of avian communities in remanescent woodlots in southern Brazil. Papeis Avulsos Zool. 33(1):1-25.
ZIMMERMAN, B. L. & R. O. BIERREGAARD. 1986. Relevance of the equilibrium theory of island biogeography and species-areas relations to conservation with a case from Amazonia. J. Biogeogr. 13:133-143.

Publication Dates

Publication in this collection
11 Aug 2009
Date of issue
1989

This work is licensed under a Creative Commons Attribution-NonCommercial 4.0 International License.

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[52] LACHER, T. E., M. A. MARES & C. J. R. ALHO in press. The structure of a small mammal community in a central Brazilian savanna. In: K. R. REDFORD (ed.), Mammals of the Americas: Essays in Honor of Ralph M. Wetzel. University of Chicago Press, Chicago.

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[60] MacARTHUR, R. H. & E. O. WILSON. 1967. The Theory of Island Biogeography. Princeton University Press, Princeton.

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[62] MARGULES, C. R. & M. B. USHER. 1981. Criteria used in assessing wildlife conservation potential: a review. Biol. Conserv. 21:79-109.

[63] MAY, R. M. 1975. Patterns of species abundance and diversity. In: M. L. CODY & J. M. DIAMOND (eds.), Ecology and Evolution of Communities. Harvard Univ. Press, Harvard, Massachussets.

[64] M'CLOSKEY, R. T. 1976. Community structure in sympatic rodents. Ecology 57:728-739.

[65] MILES, M. A., A. A. de SOUZA & M. M. POVOA. 1981. mammal tracking and nest location in Brazilian forest with an improved spool-and-line device. J. Zool. Lond. 195:331-347.

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[70] PATIL, G. P. & C. TAILLIE. 1983. Diversity as a concent and its measurement. Trans. North. Amer. Wildl. Nat. Resour. Conf. 39:334-353.

[71] PEET, R. K. 1974. The measurement of species diversity. Ann. Rev. Ecol. Syst. 1974:285-307.

[72] PIELGU, E. C. 1974. Population and Community Ecology, Gordon and Breach Publishers, New York.

[73] POOLE, R. H. 1974. An Introduction to Quantitative Ecology. McGraw-Hill, New York,

[74] PRANCE, G. T. 1982. Biological Diversification in the Tropics. Columbia University Press, New York.

[75] QUINN, J. F. & A. HASTINGS. 1987. Extinction in subdivided habitats. Conserv. Biol. 1:198-208.

[76] ROSENZWEIG, M. L. & J. WINAKUR. 1969. Population ecology of desert rodent communities: habitats and environmental complexity. Ecology 50:558-572.

[77] SHAFFER, M. L. 1981. Minimum population sizes for species conservation. Bioscience 31:131-134.

[78] SIMBERLOFF, D. S. 1976. Experimental zoogeography of islands: effects of island size. Ecology 57:629-578.

[79] SIMBERLOFF, D. S. 1982. Big advantages of small refuges. Nat. Hist. 91:6-13.

[80] SIMBERLOFF, D. S. & L. G. ABELE. 1976. Island biogeography theory and conservation pratice. Science 191:285-286.

[81] SIMBERLOFF, D. S. & L. G. ABELE. 1982. Refuge design and island biogeographic theory: effects of fragmentation. Amer. Nat. 120:41-50.

[82] SOKAL, R. R. & ROHLF, F. J. 1981. Biometry. W. H. Freeman Co., New York.

[83] SOULE, M. E. B. 1980. Thresholds for survival: maintaining fitness and evolutionary potential. In: M. E. SOULE & B. A. WILCOX (eds.), Conservation Biology. Sinauer Press, Sunderland, Massachussets.

[84] SOULE, M. E. B. 1983. What do we really know about extinction? In: C. M. SCHONEWALD-WILCOX, S. M. CHAMBERS, B. MACBRYDE & L. THOMAS (eds.), Genetics and Conservation. Benjamin Cummings Co., Menlo Park, California.

[85] SOUZA, W. P. 1984. The role of disturbance in natural communities. Ann. Rev. Ecol. Syst. 15:353-391.

[86] STALLINGS, J. R. 1988. Small mammal comunities of an eastern Brazilian park. Ph. D. Dissertation, University of Florida, Gainesville, Florida.

[87] USHER, M. B. 1986. Wildlife Conservation Evaluation. Chapman and Hall, New York,

[88] WEAVER, M. & M. KELLMAN. 1981. The effects of forest fragmentation on woodlot tree biotas in Southern Ontario. J. Biogeogr. 8:199-210.

[89] WESTMAN, W. E. 1983. Island biogeography: studies on the xeric shrublands of the inner Channel Islands, California. J. Biogeogr. 10:96-118.

[90] WILLIAMS, C. B. 1964. Patterns in the Balance of Nature. Academic Press, New York.

[91] WILLIS, E. O. 1979. The composition of avian communities in remanescent woodlots in southern Brazil. Papeis Avulsos Zool. 33(1):1-25.

[92] ZIMMERMAN, B. L. & R. O. BIERREGAARD. 1986. Relevance of the equilibrium theory of island biogeography and species-areas relations to conservation with a case from Amazonia. J. Biogeogr. 13:133-143.