, Volume 137, Issue 1, pp 104–113 | Cite as

Analysis of avian communities in Lake Guri, Venezuela, using multiple assembly rule models

  • Kenneth FeeleyEmail author
Communitity Ecology


This study analyzed the distribution of resident, forest-interior bird species nesting on islands in Lake Guri, Venezuela using several different community assembly rule models. The models that were tested included Diamond's Assembly Rules, Size Structure, Guild Proportionality, Favored States, and Nestedness. It was determined that the species composition of the study communities was only weakly influenced by competition, but that competition did appear to limit the size similarity which is permissible for co-occurring species. There was no tendency for the relative proportion of species within guilds (i.e. insectivore, omnivore, nectivore and frugivore) to remain stable among the islands. When only the insectivorous and omnivorous species were analyzed (using feeding strata as the functional groups) there was some support for the guild proportionality hypothesis. This study found no support for Fox's Favored State hypothesis, possibly due to the overrepresentation of insectivores and omnivores in the species pool. The island communities exhibit a highly nested structure. This high degree of nestedness supports the hypothesis that the assemblages are more strongly determined by differential extinction vulnerability and selective species loss than by interspecific or inter-guild competition. Understanding patterns of community assembly and their underlying forces has important implications for conservation ecology and reserve design.


Competition Nestedness Community composition Guild structure 



I would like to thank Lawrence Lopez, Dan Lebbins, Diana Escalasans, Lisa Davenport, John Terborgh and Primo Rondon for their help collecting the field data. I would also like to thank Luis Balbas for logistical support and Tom Gillespie for collecting the natural history information. Daniel Simberloff and two anonymous reviewers provided valuable comments and suggestions. This work was supported by the Frank M. Chapman Research Grant and the H. Branch Howe, Jr., Graduate Student Research Grant.


  1. Alatalo RV (1982) Bird species distribution in the Galapagos and other archipelagoes: competition or chance. Ecology 63:881−887Google Scholar
  2. Alvarez E, et al (1986) Aspectos ecologicos del embalse Guri. Interciencia 11:325−333Google Scholar
  3. Atmar W, Patterson BD (1993) The measure of order and disorder in the distribution of species in fragmented habitat. Oecologia 96:373−382Google Scholar
  4. Atmar W, Patterson BD (1995) The nestedness temperature calculator. A visual basic program, including 294 presence-absence matrices. AICS Research, University Park, N.M./The Field Museum, Chicago, Ill.Google Scholar
  5. Blake JG (1991) Nested subsets and the distribution of birds on isolated woodlots. Conserv Biol 5:58−66Google Scholar
  6. Bolger DT, Alberts AC, Soule MS (1991) Occurrence patterns of bird species in habitat fragments: sampling, extinction, and nested species subsets. Am Nat 137:155−166CrossRefGoogle Scholar
  7. Case TJ, Faaborg J, Sidell R (1983) The role of body size in the assembly of West Indian bird communities. Evolution 37:1062−1074Google Scholar
  8. Christman SP (1984) Plot mapping: estimating densities of breeding bird territories by combining spot mapping and transect techniques. Condor 86:237−241Google Scholar
  9. Connor EF, Simberloff D (1979) The assembly of species communities: chance or competition. Ecology 60:1132−1140Google Scholar
  10. Diamond JM (1975). Assembly of species communities. In: Cody ML, Diamond JM (eds) Ecology and evolution of communities. Harvard University Press, Cambridge, Mass., pp 342−444Google Scholar
  11. Diamond JM, Gilpin ME(1982) Examination of the "null" model of Connor and Simberloff for species co-occurrences on islands. Oecologia 52:64−74Google Scholar
  12. Diamond JM, Marshall AG (1977) Distributional ecology of New Hebridean birds: a species kaleidoscope. J Anim Ecol 46:703−727Google Scholar
  13. Faaborg J (1982) Trophic and size structure of West Indian bird communities. Proc Natl Acad Sci USA 79:1563−1567Google Scholar
  14. Fox BJ (1987) Species assembly and the evolution of community structure. Evol Ecol 1:201−213Google Scholar
  15. Fox BJ, Brown JH (1993) Assembly rules for functional groups in North American desert rodent communities. Oikos 67:358−370Google Scholar
  16. Gillespie TW, Feeley JK, Walter HS (2003) Can forest bird composition be predicted in tropical forest fragments: a case study from Lake Guri, Venezuela. Conserv Biol (in press)Google Scholar
  17. Gilpin ME, Diamond JM(1982) Factors contributing to non-randomness in species co-occurrences. Oecologia 52:75−84Google Scholar
  18. Gotelli NJ (2000) Null model analysis of species co-occurence patterns. Ecology 81:2606–2621Google Scholar
  19. Gotelli NJ, Entsminger GL (2001a) EcoSim: null models software for ecology. Acquired Intelligence and Kesey-Bear. Scholar
  20. Gotelli NJ, Entsminger GL (2001b) Swap and fill algorithms in null model analysis: rethinking the knight's tour. Oecologia 129:281−291CrossRefGoogle Scholar
  21. Gotelli NJ, McCabe DJ (2002) Species co-occurrences: a meta-analysis of J.M. Diamond's assembly rule model. Ecology 83:2091−2096Google Scholar
  22. Gotelli NJ, Entsminger GL (2003) Swap algorithms in null model analysis. Ecology 84:532–535Google Scholar
  23. Hutchinson GE (1959) Homage to Santa Rosalia, or why are there so many kinds of animals? Am Nat 93:145−159Google Scholar
  24. Lomolino MV (1996) Investigating causality of nestedness of insular communities: selective immigrations or extinctions? J Biogeog 23:699−703Google Scholar
  25. MacArthur RH, Wilson EO (1967) The theory of island biogeography. Princeton University Press, Princeton, N.J.Google Scholar
  26. Manly BFJ (1995) A note on the analysis of species co-occurrences. Ecology 76:1109−1115Google Scholar
  27. Manly B, Sanderson JG (2002) A note on null models: justifying the methodology. Ecology 83:580−582Google Scholar
  28. Mikkelson GM (1993) How do food webs fall apart? A study of changes in trophic structure during relaxation on habitat fragments. Oikos 67:1993Google Scholar
  29. Morales LC, Gorzula S (1986) The interrelations of the Caroni River Basin ecosystems and hydroelectric power projects. Interciencia 11:272−277Google Scholar
  30. Patterson BD (1987) The principle of nested subsets and its implications for biological conservation. Conserv Biol 1:323−334Google Scholar
  31. Patterson BD (1990) On the temporal development of nested subset patterns of species composition. Oikos 59:330−342Google Scholar
  32. Patterson BD, Atmar W (2000) Analyzing species composition in fragments. In: Rheinwald G (ed) Isolated vertebrate communities in the Tropics. Bonn Zool Monogr 96:9−24Google Scholar
  33. Roth VL (1981) Constancy and size ratios of sympatric species. Am Nat 118:394−404CrossRefGoogle Scholar
  34. Sanderson JG, Moulton MP, Selfridge RG (1998) Null matrices and the analysis of species co-occurrences. Oecologia 116:275−283CrossRefGoogle Scholar
  35. Schluter D (1984) A variance test for detecting species associations, with some example applications. Ecology 65:998−1005Google Scholar
  36. Shields WM (1976) Avian census techniques: an analytical review. The role of insectivorous birds in forest ecosystems. Academic Press, New YorkGoogle Scholar
  37. Sieving KE, Karr JR (1997) Avian extinction and persistence mechanisms in lowland Panama. In: Laurance WF, Bierregaard RO (eds) Tropical forest remnants. University of Chicago Press, Chicago, Ill., pp 156−170Google Scholar
  38. Simberloff D, Boecklen W (1981) Santa Rosalita reconsidered: size ratios and competition. Evolution 35:1206−1228Google Scholar
  39. Simberloff D, Dayan T (1991) The guild concept and the structure of ecological communities. Annu Rev Ecol Syst 22:115−143CrossRefGoogle Scholar
  40. Sokal RR, Rohlf FJ (1995) Biometry. Freeman, New YorkGoogle Scholar
  41. Stone L, Roberts A (1990) The checkerboard score and species distribution. Oecologia 85:74−79Google Scholar
  42. Stone L, Dayan T, Simberloff D (1996) Community-wide assembly patterns unmasked: the importance of species' differing geographical ranges. Am Nat 148:997−1015CrossRefGoogle Scholar
  43. Stone L, Dayan T, Simberloff D (2000) On desert rodents, favored states, and unresolved issues: scaling up and down regional assemblages and local communities. Am Nat 156:322−328CrossRefGoogle Scholar
  44. Stouffer PC, Bierregaard RO (1995) Use of Amazonian forest fragments by understory insectivorous birds. Ecology 76:2429−2445Google Scholar
  45. Svensson SE (1978) Census efficiency and number of visits to a study plot when estimating bird densities by the territory mapping technique. J Appl Ecol 16:61−68Google Scholar
  46. Terborgh J (1974) Preservation of natural diversity: the problem of extinction prone species. Bioscience 24:715−722Google Scholar
  47. Terborgh JW, Faaborg J (1980) Saturation of bird communities in the West Indies. Am Nat 116:178−195CrossRefGoogle Scholar
  48. Terborgh J, Winter B (1980) Some causes of extinction. In: Soule ME, Wilcox BA (eds) Conservation biology: an evolutionary-ecological perspective. Sinauer, Sunderland, Mass., pp 199−133Google Scholar
  49. Terborgh J, et al (1997a) Bird communities in transition: The Lago Guri islands. Ecology 78:1494−1501Google Scholar
  50. Terborgh J, et al (1997b) Transitory states in relaxing ecosystems of land bridge islands. In: Laurence WF, Berregaard RO (eds) Tropical forest remnants. University of Chicago Press, Chicago. Ill., pp 256−274Google Scholar
  51. Warburton NH (1997) Structure and conservation of forest avifauna in isolated rainforest remnants in tropical Australia. In: Laurence WF, Bierregaard RO (eds) Tropical forest remnants. University of Chicago Press, Chicago, Ill., pp 190−208Google Scholar
  52. Weiher E, Keddy P (2001) Ecological assembly rules. Cambridge University Press, CambridgeGoogle Scholar
  53. Williams JD, et al (1998) Ecology and status of piscovers in Guri, an oligotrophic tropical reservoir. N Am J Fish 18:274−285Google Scholar
  54. Wilson JB (1989) A null model of guild proportionality, applied to stratification of a New Zealand temperate rain forest. Oecologia 80:263−267Google Scholar
  55. Wilson JB (1995) Null models for assembly rules: the Jack Horner effect is more insidious than the Narcissus effect. Oikos 72:139−144Google Scholar
  56. Wilson JB, Whittaker RJ (1995) Assembly rules demonstrated in a saltmarsh community. J Ecol 83:801−807Google Scholar
  57. Wright DH, et al (1998) A comparative analysis of nested subset patterns in species composition. Oecologia 113:1−20CrossRefGoogle Scholar
  58. Yeaton RI (1974) An ecological analysis of chaparral and pine forest bird communities of Santa Cruz island and mainland California. Ecology 55:959−973Google Scholar
  59. Yeaton RI, Cody ML (1974) Competitive release in island song sparrow populations. Theor Pop Biol 5:42−58Google Scholar

Copyright information

© Springer-Verlag 2003

Authors and Affiliations

  1. 1.Department of BiologyDuke UniversityDurhamUSA

Personalised recommendations