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Community Ecology Species/area relationship Click here for supplemental materials for today (PDF) Outline: 1. Discussion of the ubiquitous relationship between species richness and area A. Patterns associated with k and z 2. Four possible explanations for this pattern 3. How loss of A may unexpectedly affect S via supersaturation 4. How universal is the S/A relationship? Terms/people: S=kAz passive sampling hypothesis area per se Preston Darlington’s Rule supersaturated habitat heterogeneity hypothesis Arrhenius extinction debt (Tilman et al.) faunal relaxation (Diamond) As area increases, richness increases: one of the few “laws” in ecology, according to Schoener (1976). First attributed to H.C. Watson (1859). Lots of empirical evidence for S/A relationship e.g. Darlington’s Rule Formalization of the S/A relationship - Arrhenius 1921, Preston 1962: S = kAz where k = y-intercept, z = slope, S=number of species, A=area To linearize: log(S) = zlog(A) + log (k) See Scheiner (2003) for a discussion of 6 types of species-area curves, based on various aspects of sampling. Properties of k and z can provide insights about isolation, extinction risk, etc. In general, data are plotted and a curve is then fitted retroactively, which allows calculation of k and z. Explanations for species-area relationship: 1) area per se with respect to colonization and extinction 2) habitat heterogeneity hypothesis (see Fig. 2.14 in Mittelbach text) 3) passive sampling phenomenon 4) disturbance BUT... The species-area relationship is not as straightforward as it first appears… Habitat loss, supersaturation, extinction debt, and faunal relaxation What are we to conclude about species/area relationships? S is related to A because of a variety of factors that vary with A, including habitat, sampling, and other properties. There is considerable scatter about the S/A regression line (supersaturation). Nonetheless, the S/A relationship is a reasonably robust pattern––it is the process behind it that is unclear. Next time: the relationship between biodiversity and environmental productivity. References: Abbott, I. 1983. The meaning of z in species/area regressions and the study of species turnover in island biogeography. Oikos 41:385-390. Arrhenius, O. 1921. Species and area. J. Ecol. 9:95-99. Becker, P. 1992. Colonization of islands by carnivorous and herbivorous Heteroptera and Coleoptera: effects of island area, plant species richness, and ‘extinction’ rates. J. Biogeogr. 19:163-171. Boecklen, W.J. 1986. Effects of habitat heterogeneity on the species-area relationships of forest birds. J. Biogeogr. 13:59-68. Connor, E.F., and E.D. McCoy. 1979. The statistics and biology of the species-area relationship. Am. Nat. 113:791-833. Darlington, P.J. 1957. Zoogeography: The Geographical Distribution of Animals. Wiley, New York, NY. den Boer, P.J. 1968. Spreading of risk and stabilization of animal numbers. Acta Biotheor. 18:165- 194. Diamond, J.M. 1972. Biogeographic kinetics: estimation of relaxation times for avifaunas of southwest Pacific islands. Proc. Nat. Acad. Sci. USA 69:3199-3203. Drakare, S., J.J. Lennon, and H. Hillebrand. 2006. The imprint of the geographical, evolutionary and ecological context on species-area relationships. Ecology Letters 9:215-227. Dunn, C.P., and C. Loehle. 1988. Species-area parameter estimation: testing the null model of lack of relationship. J. Biogeogr. 15: 721-728. Gitay, H., S.H. Roxburgh, and J.B. Wilson. 1991. Species-area relationships in a New Zealand tussock grassland, with implications for nature reserve design and for community structure. J. Veg. Sci. 2:113-118. Matter, S.F., I. Hanski, M. Gyllenberg. 2002. A test of the metapopulation model of the speciesarea relationship. J. Biogeogr. 9:977-983. McGuinness, K.A. 1984. Equations and explanations in the study of species-area curves. Biol. Rev. 59:423-440. Ney-Nifle, M., and M. Mangel. 2000. Habitat loss and changes in the species-area relationship. Conserv. Biol. 14:893-898. Palmer, M.W., and P.S. White. 1994. Scale dependence and the species-area relationship. Am. Nat. 144:717-740. Preston, F.W. 1948. The commonness and rarity of species. Ecology 29:254-283. Preston, F.W. 1962. The canonical distribution of commonness and rarity of species. Ecology 43:185-215. Rosenzweig, M.L. 1995. Species Diversity in Space and Time. Cambridge Univ. Press, New York, NY. Scheiner, S.M. 2003. Six types of species-area curves. Global Ecology & Biogeography 12:441-447. Schoener, T.W. 1976. The species-area relationship within archipelagos: models and evidence from island land birds. Proceedings of the 16th International Ornithological Congress (Canberra), pp. 629- 642. Sugihara, G. 1981. S=CAz, z=¼: a reply to Connor and McCoy. Am. Nat. 117:790-793. Tilman, D., R.M. May, C.L. Lehman, and M.A. Nowak. 1994. Habitat destruction and the extinction debt. Nature 371:65-66. Whittaker, R.J., and J.M. Fernández-Palacios. 2007. Island Biogeography: Ecology, Evolution, and Conservation (2nd ed.). Oxford Univ. Press, Oxford. Williamson, M. 1988. Relationship of species number to area, distance and other variables. Pp. 91-115 in: Analytical Biogeography (A.A. Myers and P.S. Giller, eds.). Chapman and Hall, London.