Predictors of Species Sensitivity to Fragmentation

Biodiversity and Conservation - Tập 13 - Trang 207-251 - 2004
Klaus Henle1, Kendi F. Davies2, Michael Kleyer3, Chris Margules4, Josef Settele5
1Department of Conservation Biology and Natural Resources, Centre for Environmental Research, UFZ-Leipzig-Halle, Leipzig, Germany
2CSIRO Sustainable Ecosystems, Alice Springs, Australia
3Landscape Ecology Group (FB 7), University of Oldenburg, Oldenburg, Germany
4CSIRO Sustainable Ecosystems and the Rainforest Cooperative Research Centre, CSIRO Tropical Forest Research Centre, Atherton, Australia
5Department of Community Ecology, Centre for Environmental Research, UFZ-Leipzig-Halle, Halle, Germany

Tóm tắt

We reviewed empirical data and hypotheses derived from demographic, optimal foraging, life-history, community, and biogeographic theory for predicting the sensitivity of species to habitat fragmentation. We found 12 traits or trait groups that have been suggested as predictors of species sensitivity: population size; population fluctuation and storage effect; dispersal power; reproductive potential; annual survival; sociality; body size; trophic position; ecological specialisation, microhabitat and matrix use; disturbance and competition sensitive traits; rarity; and biogeographic position. For each trait we discuss the theoretical justification for its sensitivity to fragmentation and empirical evidence for and against the suitability of the trait as a predictor of fragmentation sensitivity. Where relevant, we also discuss experimental design problems for testing the underlying hypotheses. There is good empirical support for 6 of the 12 traits as sensitivity predictors: population size; population fluctuation and storage effects; traits associated with competitive ability and disturbance sensitivity in plants; microhabitat specialisation and matrix use; rarity in the form of low abundance within a habitat; and relative biogeographic position. Few clear patterns emerge for the remaining traits from empirical studies if examined in isolation. Consequently, interactions of species traits and environmental conditions must be considered if we want to be able to predict species sensitivity to fragmentation. We develop a classification of fragmentation sensitivity based on specific trait combinations and discuss the implications of the results for ecological theory.

Tài liệu tham khảo

Allee W.C., Emerson A.E., Park O., Park T. and Schmidt K.P. 1949. Principles of Animal Ecology. Saunders, Philadelphia, Pennsylvania. Andrén H., Delin A. and Seiler A. 1997. Population response to landscape changes depends on specialisation to different landscape elements. Oikos 80: 193-196. Angermeier P.I. 1995. Ecological attributes of extinction-prone species: loss of freshwater fishes of Virginia. Conservation Biology 9: 143-158. Bahl A., Pfenninger M., Bamberger H., Frye M. and Streit B.B. 1996. Survival of snails in fragmented landscapes. In: Settele J., Margules C.R., Poschlod P. and Henle K. (eds), Species Survival in Fragmented Landscapes. Kluwer, Dordrecht, The Netherlands, pp. 329-343. Bahl A., Bender C., Gottschalk E., Pfenninger M. and Seufert W. 1999. Populationsgefährdungsanalysen auf verschiedenen räumlichen Ebenen. In: Amler K., Bahl A., Henle K., Kaule G., Poschlod P. and Settele J. (eds), Populationsbiologie in der Naturschutzpraxis. Ulmer, Stuttgart, Germany, pp. 148-153. Belovsky G.E. 1987. Extinction models and mammalian persistence. In: Soulé M.E. (ed), Viable Populations for Conservation. Cambridge University Press, Cambridge, UK, pp. 35-57. Bengtsson J. 1989. Interspecific competition increases local extinction rate in a metapopulation system. Nature 340: 713-715. Bengtsson J. and Milbrink G. 1995. Predicting extinctions: interspecific competition, predation and population variability in experimental Daphnia populations. Oecologia 101: 397-406. Benitez-Malvido J. 1998. Impact of forest fragmentation on seedling abundance in a tropical rain forest. Conservation Biology 12: 380-389. Bentley J.M., Catterall C.P. and Smith G.C. 2000. Effects of fragmentation of araucarian vine forest on small mammal communities. Conservation Biology 14: 1075-1087. Berger J. 1990. Persistence of different-sized populations: an empirical assessment of rapid extinctions in bighorn sheep. Conservation Biology 4: 91-96. Biedermann R., Hoffmann E. and Seufert W. 1999. Auswahl von Tierarten in der Planungspraxis auf populationsökologischer Basis: Das Fallbeispiel Wirbellose in der Porphyrkuppenlandschaft. In: Amler K., Bahl A., Henle K., Kaule G., Poschlod P. and Settele J. (eds), Populationsbiologie in der Naturschutzpraxis. Ulmer, Stuttgart, Germany, pp. 29-37. Blackburn T.M. and Gaston K.J. 1994. Animal body size distributions: patterns, mechanisms and implications. Trends in Ecology and Evolution 9: 471-474. Blackburn T.M. and Gaston K.J. 1997. A critical assessment of the form of the interspecific relationship between abundance and body size in animals. Journal of Animal Ecology 66: 233-249. Blackburn T.M., Lawton J.H. and Perry J.N. 1992. A method of estimating the slope of the upper bounds of plots of body size and abundance in natural animal assemblages. Oikos 65: 107-112. Blackburn T.M., Brown V.K., Doube B.M., Greenwood J.J.D., Lawton J.H. and Stork N.E. 1993. The relationship between abundance and body size in natural assemblages. Journal of Animal Ecology 62: 519-528. Blackburn T.M., Gates S., Lawton J.H. and Greenwood J.J.D. 1994. Relations between body size, abundance and taxonomy of birds wintering in Britain and Ireland. Philosophical Transactions of the Royal Society of London, Series B 343: 135-144. Bolger D.T., Alberts A.C., Sauvajot R.M., Potenza P., McCalvin C., Tran D. et al. 1997. Response of rodents to habitat fragmentation in coastal southern California. Ecological Applications 7: 552-563. Bonn S. and Poschlod P. 1998. Ausbreitungsbiologie der Pflanzen Mitteleuropas. Grundlagen und kulturhistorische Aspekte. Quelle and Meyer, Wiesbaden, Germany. Bräuer I., Maibom W., Matthies D. and Tscharntke T. 1999. Populationsgröße und Aussterberisiko gefährdeter Pflanzenarten in Niedersachsen. Verhandlungen der Gesellschaft für Ökologie 29: 505-510. Brown J.H. and Kodric-Brown A. 1977. Turnover rates in insular biogeography: effect of immigration on extinction. Ecology 58: 445-449. Burbidge A.A. and McKenzie N.L. 1989. Patterns in the modern decline of Western Australia's vertebrate fauna: causes and conservation implications. Biological Conservation 50: 143-198. Cadenasso M.L., Traynor M.M. and Pickett S.T.A. 1997. Functional location of forest edges: gradients of multiple physical factors. Canadian Journal of Forest Research 27: 774-782. Casagrandi R. and Gatto M. 1999. A mesoscale approach to extinction risk in fragmented habitats. Nature 400: 560-562. Castelletta M., Sodhi N.S. and Subaraj R. 2000. Heavy extinctions of forest avifauna in Singapore: lessons for biodiversity conservation in southeast Asia. Conservation Biology 14: 1870-1880. Chalfoun A.D., Thompson F.R. and Ratnaswamy M.J. 2002. Nest predators and fragmentation: a review and meta-analysis. Conservation Biology 16: 306-318. Chesson P. 1991. Stochastic population models. In: Kolasa J. and Pickett S.T.A. (eds), Ecological Heterogeneity. Springer, New York, pp. 123-143. Cody M.L. 1986. Structural niches in plant communities. In: Diamond J. and Case T.J. (eds), Community Ecology. Harper & Row, New York, pp. 381-405. Cook R.E. 1985. Growth and development in clonal plant population. In: Jackson J.B.C., Buss L.W. and Cook R.E. (eds), Population Biology and Evolution of Clonal Organisms. Yale University Press, New Haven, Connecticut, pp. 259-296. Cook R.R. and Hanski I. 1995. On expected lifetimes of small-bodied and large-bodied species of birds on islands. The American Naturalist 145: 305-315. Corlett R.T. and Turner I.M. 1997. Long-term survival in tropical forest remnants in Singapore and Hong Kong. In: LauranceW.F. and Bierregaard R.O. (eds), Tropical Forest Remnants. University of Chicago Press, Chicago, Illinois, pp. 333-346. Cotgreave P. 1993. The relationship between body size and population abundance in animals. Trends in Ecology and Evolution 8: 244-248. Crooks K.R. 2002. Relative sensitivity of mammalian carnivores to habitat fragmentation. Conservation Biology 16: 488-502. Crooks K.R. and Soulé M.E. 1999. Mesopredator release and avifaunal extinctions in a fragmented system. Nature 400: 563-566. Crooks K.R., Suarez A.V., Bolger D.T. and Soulé M.E. 2001. Extinction and colonization of birds on habitat islands. Conservation Biology 15: 159-172. Crowell K.L. 1973. Experimental zoogeography: introductions of mice to small islands. The American Naturalist 107: 535-558. Cyr H., Downing J.A. and Peters R.H. 1997. Density-body size relationships in local aquatic communities. Oikos 79: 333-346. Davies K.F. and Margules C.R. 2000. The beetles at Wog Wog: a contribution of Coleoptera systematics to an ecological field experiment. Invertebrate Taxonomy 14: 953-956. Davies K.F., Margules C.R. and Lawrence J.F. 2000.Which traits of species predict population declines in experimental forest fragments? Ecology 81: 1450-1461. Davies K.F., Melbourne B.A. and Margules C.R. 2001. Effects of within and between-patch processes on beetle-community dynamics in experimentally fragmented forest. Ecology 82: 1830-1846. Davies K.F., Margules C.R. and Lawrence J.F. A synergistic effect puts rare, specialized species at greater risk of extinction. Ecology, in press. Debinski D.M. and Holt R.D. 2000. A survey and overview of habitat fragmentation experiments. Conservation Biology 14: 342-355. Diamond J.M. 1984. Normal extinctions of isolated populations. In: Nitecki M.H. (ed), Extinctions. University of Chicago Press, Chicago, Illinois, pp. 191-246. Diamond J.M., Bishop K.D. and van Balen S. 1987. Bird survival in an isolated Javan woodland: island or mirror? Conservation Biology 1: 132-142. Didham R.K., Lawton J.H., Hammond P.M. and Eggletin P. 1989. Trophic structure stability and extinction dynamics of beetles (Coleoptera) in tropical forest fragments. Philosophical Transactions of the Royal Society of London, Series B 353: 437-451. Diffendorfer J.E., Gaines M.S. and Holt R.D. 1995. Habitat fragmentation and movements of three small mammals (Sigmodon, Microtus, and Peromyscus). Ecology 76: 827-839. Dooley J.L. and Bowers M.A. 1998. Demographic responses to habitat fragmentation: experimental tests at the landscape and patch scale. Ecology 79: 969-980. Durant S.M. and Mace G.M. 1994. Species differences and population structure in population viability analysis. In: Olney P.J.S., Mace G.M. and Feistner A.T.C. (eds), Creative Conservation: Interactive Management of Wild and Captive Animals. Chapman & Hall, London, pp. 67-91. Ebenhard T. 1989. Bank vole [Clethrionomys glareolus (Schreiber, 1780)] propagules of different sizes and island colonization. Journal of Biogeography 16: 173-180. Ehrlén J. and van Groenendal J.M. 1998. The trade-off between dispersability and longevity-an important aspect of plant species diversity. Applied Vegetation Science 1: 29-36. Eisto, A.-K., Kuitunen M., Lammi A., Saari V., Suhonen J., Syrjäsuo S. et al. 2000. Population persistence and offspring fitness in the rare bellflower Campanula cervicaria in relation to population size and habitat quality. Conservation Biology 14: 1413-1421. Eitschberger U., Reinhardt R. and Steiniger H. 1991. Wanderfalter in Europa. Atalanta 22: 1-67. Ellenberg H. 1950. Kausale Pflanzensoziologie auf physiologischer Grundlage. Berichte der Deutschen Botanischen Gesellschaft 63: 24-31. Eriksson O. 1996. Regional dynamics of plants: a review of evidence for remnant, source-sink and metapopulations. Oikos 77: 248-258. Eriksson O. and Jakobsson A. 1999. Recruitment trade-offs and the evolution of dispersal mechanisms in plants. Evolutionary Ecology 13: 411-423. Fagan W.F., Meir E. and Moore J.L. 1999. Variation thresholds for extinction and their implications for conservation strategies. The American Naturalist 154: 510-520. Fahrig L. 1998. When does fragmentation of breeding habitat affect population survival? Ecological Modelling 105: 273-292. Fiedler P.L. and Ahous J.J. 1992. Hierarchies of cause: toward an understanding of rarity in vascular plant species. In: Fiedler P.L. and Jain S.K. (eds), Conservation Biology: The Theory and Practice of Nature Preservation and Management. Cambridge University Press, Cambridge, UK, pp. 23-47. Fischer S.F., Poschlod P. and Beinlich B. 1996. Experimental studies on the dispersal of plants and animals on sheep in calcareous grasslands. Journal of Applied Ecology 33: 1206-1222. Forney K.A. and Gilpin M.E. 1989. Spatial structure and population extinction: a study with Drosophila flies. Conservation Biology 3: 45-51. Foufopoulos J. and Ives A.R. 1999. Reptile extinctions on land-bridge islands: life-history attributes and vulnerability to extinction. The American Naturalist 153: 1-25. Frank K., Drechsler M. and Wissel C. 1994. Ñberleben in fragmentierten Lebensräumen-Stochastische Modelle zu Metapopulationen. Zeitschrift für Ökologie und Naturschutz 3: 167-178. Gabriel W. and Bürger R. 1992. Survival of small populations under demographic stochasticity. Theoretical Population Biology 41: 44-71. Gard T.C. 1984. Persistence in food webs. In: Levin S.A. and Hallam T.G. (eds),Mathematical Ecology. Springer, Berlin, Germany, pp. 208-219. Gascon C., Lovejoy T.E., Bierregaard R.O., Malcolm J.R., Stouffer P.C., Vasconcelos H. et al. 1999. Matrix habitat and species persistence in tropical forest remnants. Biological Conservation 91: 223-229. Gaston K.J. 1994. Rarity. Chapman & Hall, London. Gaston K.J. and Blackburn T.M. 1995. Birds, body size and the threat of extinction. Philosophical Transactions of the Royal Society of London, Series B 347: 205-212. Gaston K.J. and Blackburn T.M. 1996a. Conservation implications of geographic range size-body size relationships. Conservation Biology 10: 638-646. Gaston K.J. and Blackburn T.M. 1996b. Global scale macroecology-interactions between population size, geographic range size and body size in the Anseriformes. Journal of Animal Ecology 65: 701-714. Gaston K.J. and Blackburn T.M. 1996c. Range size-body size relationships: evidence of scale dependence. Oikos 75: 479-485. Gaston K.J. and Blackburn T.M. 1996d. Rarity and body size: importance of generality. Conservation Biology 10: 1295-1298. Gaston K.J., Blackburn T.M. and Spicer J.I. 1998. Rapoport's rule: time for an epitaph? Trends in Ecology and Evolution 13: 70-74. Gentry A.H. 1990. Four Neotropical Rainforests. Yale University Press, New Haven, Connecticut. Goldberg D.E. 1996. Competitive ability in plants: consistency, contingency, and correlated traits. Philosophical Transactions of the Royal Society London, Series B 351: 1377-1385. Goodman D. 1987. The demography of chance extinctions. In: Soulé M.E. (ed), Viable Populations for Conservation. Cambridge University Press, Cambridge, UK, pp. 11-34. Grime J.P. 1973. Control of species diversity in herbaceous vegetation. Journal of Environmental Management 1: 151-167. Grime J.P., Hodgson J.G. and Hunt R. 1988. Comparative Plant Ecology. Unwin Hyman, London. Grime J.P., Thompson K., Hunt R., Hodgson J.G., Cornelissen J.H.C., Riorison I.H. et al. 1997. Integrated screening validates primary axes of specialisation in plants. Oikos 79: 259-281. Groombridge B. 1992. Global Biodiversity. Chapman & Hall, London. Haig S.M., Belthoff J.R. and Allen D.H. 1993. Population viability analysis for a small population of red-cockaded woodpeckers and an evaluation of enhancement strategies. Conservation Biology 7: 289-301. Hassell M.P., Lawton J.H. and May R.M. 1976. Patterns of dynamical behaviour in singlespecies populations. Journal of Animal Ecology 45: 471-486. Hawkins B.A. 1994. Pattern and Process in Host-parasitoid Interactions. Cambridge University Press, Cambridge, UK. Henle K. and Streit B. 1990. Kritische Beobachtungen zum Artenrückgang bei Amphibien und Reptilien und zu dessen Ursachen. Natur und Landschaft 65: 347-361. Henle K., Poschlod P., Margules C. and Settele J. 1996. Species survival in relation to habitat quality, size, and isolation: summary, conclusions and future directions. In: Settele J., Margules C.R., Poschlod P. and Henle K. (eds), Species Survival in Fragmented Landscapes. Kluwer, Dordrecht, The Netherlands, pp. 373-381. Henle K., Sarre S. and Wiegand K. 2004. The role of density regulation in extinction processes and population viability analysis. Biodiversity and Conservation 13: 9-52 (this issue). Higgins S.I., Pickett S.T.A. and Bond W. 2000. Predicting extinction risks for plants: environmental stochasticity can save declining populations. Trends in Ecology and Evolution 15: 515-520. Hildrew A.G. 1985. A quantitative study of the life history of a fairy shrimp (Branchiopoda: Anostraca) in relation to the temporary nature of its habitat, a Kenyan rain pool. Journal of Animal Ecology 54: 99-110. Hoffmeister T.S. and Rohlfs M. 2001. Aggregative egg distributions may promote species co-existence-but why do they exist? Evolutionary Ecology Research 3: 37-50. Holt R.D. 1996. Food webs in space: an island biogeographic perspective. In: Polis G.A. and Winemiller K.O. (eds), Food Webs: Integration of Patterns and Dynamics. Chapman & Hall, New York, pp. 313-323. Huston M. 1979. A general hypothesis of species diversity. The American Naturalist 113: 81-101. Huxel G.R. and Hastings A. 1998. Population size dependence, competitive coexistence and habitat destruction. Journal of Animal Ecology 67: 446-453. Iwasa Y. and Roughgarden J. 1986. Interspecific competition in metapopulations with spacelimited subpopulations. Theoretical Population Biology 30: 194-214. Jablonski D. 1987. Heritability at the species level: analysis of geographic ranges of Cretaceous molluscs. Science 238: 360-363. Jackel A.K. and Poschlod P. 1996. Why are some plant species of fragmented continental dry grasslands frequent and some rare?-the role of germination and establishment. In: Settele J., Margules C.R., Poschlod P. and Henle K. (eds), Species Survival in Fragmented Landscapes. Kluwer, Dordrecht, The Netherlands, pp. 194-203. Johst K. and Brandl R. 1997. Body size and extinction risk in a stochastic environment. Oikos 78: 612-617. Jonsson A. and Ebenman B. 2001. Are certain life histories particularly prone to local extinction? Journal of Theoretical Biology 209: 455-463. Karr J.R. 1982a. Population variability and extinction in the avifauna of a tropical land bridge Island. Ecology 63: 1975-1978. Karr J.R. 1982b. Avian extinction on Barro Colorado Island, Panama: a reassessment. The American Naturalist 119: 220-239. Karr J.R. 1990. Avian survival rates and the extinction process on Barro Colorado Island, Panama. Conservation Biology 4: 391-397. Kattan G.H. 1992. Rarity and vulnerability: the birds of the Cordillera Central of Colombia. Conservation Biology 6: 64-70. Kattan G.H., Alvarez-López H. and Giraldo M. 1994. Forest fragmentation and bird extinctions: San Antonio eighty years later. Conservation Biology 8: 138-146. Kiefer S. and Poschlod P. 1996. Restoration of fallow or afforested calcareous grasslands by clear-cutting. In: Settele J., Margules C.R., Poschlod P. and Henle K. (eds), Species Survival in Fragmented Landscapes. Kluwer, Dordrecht, The Netherlands, pp. 209-218. Klein B.C. 1989. Effects of forest fragmentation on dung and carrion beetle communities in central Amazonia. Ecology 70: 1715-1725. Kleyer M. 1999. The distribution of plant functional types on gradients of disturbance intensity and resource supply in an agricultural landscape. Journal of Vegetation Science 10: 697-708. Kleyer M., Kaule G. and Settele J. 1996. Landscape fragmentation and landscape planning. In: Settele J., Margules C., Poschlod P. and Henle K. (eds), Species Survival in Fragmented Landscapes. Kluwer, Dordrecht, The Netherlands, pp. 138-151. Köhler G. 1996. The ecological background of population vulnerability in Central European grasshoppers and bush crickets: a brief review. In: Settele J., Margules C.R., Poschlod P. and Henle K. (eds), Species Survival in Fragmented Landscapes. Kluwer, Dordrecht, The Netherlands, pp. 290-298. Kunin W.E. and Gaston K.J. 1993. The biology of rarity: patterns, causes and consequences. Trends in Ecology and Evolution 8: 289-301. Lande R. 1987. Extinction thresholds in demographic models of territorial populations. The American Naturalist 130: 624-635. Lande R. 1993. Risks of population extinction from demographic and environmental stochasticity and random catastrophe. The American Naturalist 142: 911-927. Lande R. and Orzack S.H. 1988. Extinction dynamics of age-structured populations in a fluctuating environment. Proceedings of the National Academy of Science USA 85: 7418-7421. Laurance W.F. 1991. Ecological correlates of extinction proneness in Australian tropical rain forest mammals. Conservation Biology 5: 79-89. Laurance W.F. and Bierregaard R.O. 1997. Tropical Forest Remnants. University of Chicago Press, Chicago, Illinois. Laurance W.F., Ferreira L.V., Rankin-de-Merona J.M. and Laurance S.G. 1998. Rain forest fragmentation and the dynamics of Amazonian tree communities. Ecology 79: 2032-2040. Laurance W.F., Lovejoy T.E., Vasconcelos H.L., Bruna E.M., Didham R.K., Stouffer P.C. et al. 2002. Ecosystem decay of Amazonian forest fragments: a 22-year investigation. Conservation Biology 16: 605-618. Lavorel S., McIntyre S., Landsberg J. and Forbes T.D.A. 1997. Plant functional classifications: from general groups to specific groups based on response to disturbance. Trends in Ecology and Evolution 12: 474-478. Lawler S.P. 1993. Species richness, species composition and population dynamics of protists in experimental microcosms. Journal of Animal Ecology 62: 711-719. Lawton J.H. 1994. Population dynamic principles. Philosophical Transactions of the Royal Society of London, Series B 344: 61-68. Lawton J.H. 1995. Population dynamics principles. In: Lawton J.H. and May R.M. (eds), Extinction Rates. Oxford University Press, Oxford, UK, pp. 147-163. Leach M.K. and Givnish T.J. 1996. Ecological determinants of species loss in remnant prairies. Science 273: 1555-1558. Leigh E.G. 1981. The average lifetime of a population in a varying environment. Journal of Theoretical Biology 90: 213-239. Levins R. 1970. Extinction. In: Gerstenhaber M. (ed), Some Mathematical Questions in Biology. American Mathematical Society, Providence, Rhode Island, pp. 77-107. Lima M., Marquet P.A. and Jaksic F.M. 1996. Extinction and colonisation processes in subpopulations of five Neotropical small mammal species. Oecologia 107: 197-203. Lindenmayer D.B. and Lacy R.C. 1995a. Metapopulation viability of arboreal marsupials in fragmented old-growth forests: comparison among species. Ecological Applications 5: 183-199. Lindenmayer D.B. and Lacy R.C. 1995b. Metapopulation viability of Leadbeater's possum, Gymnobelideus leadbeateri, in fragmented old-growth forests. Ecological Applications 5: 164-182. Lindenmayer D.B., Cunningham R.B. and Smith A.P. 1990. The conservation of arboreal marsupials in the montane ash forests of the central highlands of Victoria, South-East Australia: II. The loss of trees with hollows and its implications for the conservation of Leadbeater's possum Gymnobelideus leadbeateri McCoy (Marsupialia: Petauridae). Biological Conservation 54: 133-145. Liu J. 1993. Discounting initial population sizes for prediction of extinction probabilities in patchy environments. Ecological Modelling 70: 51-61. Lynch J.F. 1987. Responses of breeding bird communities to forest fragmentation. In: Saunders D.A., Arnold G.W., Burbidge A.A. and Hopkins A.J.M. (eds), Nature Conservation: The Role of Remnants of Native Vegetation. Surrey Beatty, Chipping Norton, Australia, pp. 123-140. Mac Nally R. and Bennett A.F. 1997. Species-specific predictions of the impact of habitat fragmentation: local extinction of birds in the box-ironbark forests of Central Victoria, Australia. Biological Conservation 82: 147-155. Main A.R. 1987. Management of remnants of native vegetation-a review of the problems and the development of an approach with reference to the wheatbelt of Western Australia. In: Saunders D.A., Arnold G.W., Burbidge A.A. and Hopkins A.J.M. (eds), Nature Conservation: The Role of Remnants of Native Vegetation. Surrey Beatty, Chipping Norton, Australia, pp. 1-13. Margules C.R. 1996. Experimental fragmentation. In: Settele J., Margules C.R., Poschlod P. and Henle K. (eds), Species Survival in Fragmented Landscapes. Kluwer, Dordrecht, The Netherlands, pp. 128-137. Margules C.R., Milkovits G.A. and Smith G.T. 1994. Contrasting effects of habitat fragmentation on the scorpion Cercophonius squama and an amphipod. Ecology 75: 2033-2042. Matthies D. 1991. Räumliche und zeitliche Dynamik in Populationen der seltenen Art Melampyrum arvense L. In: Schmid B. and Stöcklin J. (eds), Populationsbiologie der Pflanzen. Birkhäuser, Basel, Switzerland, pp. 109-121. May R.M. and Oster G.F. 1976. Bifurcations and dynamic complexity in simple ecological models. The American Naturalist 110: 573-599. McArdle B. and Gaston K.J. 1993. The temporal variability of populations. Oikos 67: 187-191. McCarthy M.A., Lindenmayer D.B. and Drechsler M. 1997. Extinction debts and risks faced by abundant species. Conservation Biology 11: 221-226. McDade L.A., Bawa K.S., Hespenheide H.A. and Hartshorn G. 1994. La Selva-Ecology and Natural History of a Neotropical Rain Forest. University of Chicago Press, Chicago, Illinois. McIntyre S., Lavorel S., Landsberg J. and Forbes T.D.A. 1999. Disturbance response in vegetation-towards a global perspective on functional traits. Journal of Vegetation Science 10: 621-630. McKelvey K., Noon B.R. and Lamberson R.H. 1993. Conservation planning for species occupying fragmented landscapes-the case of the northern spotted owl. In: Kareiva P.M., Ingsolver J.G. and Huey R.B. (eds), Biotic Interactions and Global Change. Sinauer, Sunderland, Massachusetts, pp. 424-450. Menges E.S. 1990. Population viability analysis for an endangered plant. Conservation Biology 4: 52-62. Metzger J.P. 2000. Tree functional group richness and landscape structure in a Brazilian tropical fragmented landscape. Ecological Applications 10: 1147-1161. Mikkelson G.M. 1993. How do food webs fall apart? A study of changes in trophic structure during relaxation on habitat fragments. Oikos 67: 539-547. Mühlenberg M., Hovestadt T. and Röser J. 1991. Are there minimal areas for animal populations? In: Seitz A. and Loeschcke V. (eds), Species Conservation: A Population-Biological Approach. Birkhäuser, Basel, Switzerland, pp. 227-264. Nantel P., Gagnon D. and Nault A.A. 1996. Population viability analysis of American ginseng and wild leek harvested in stochastic environments. Conservation Biology 10: 608-621. Noon B.R. and McKelvey K.S. 1996. A common framework for conservation planning: linking individual and metapopulation models. In: McCullough D.R. (ed), Metapopulations and Wildlife Conservation. Island Press, Washington, DC, pp. 139-165. Oedekoven K.-H. 1992. Forstgeschichte des Nahen Osten. Schriftenreihe Wald und Umwelt: die Welt von morgen. Waldschutz-Weltweit 1992: 19-37. Ogle C.C. 1987. The incidence and conservation of animal and plant species in remnants of native vegetation within New Zealand. In: Saunders D.A., Arnold G.W., Burbidge A.A. and Hopkins A.J.M. (eds), Nature Conservation: The Role of Remnants of Native Vegetation. Surrey Beatty, Chipping Norton, Australia, pp. 79-87. Oostermeijer J.G.B. 2000. Population viability analysis of the rare Gentiana pneumonanthe: the importance of genetics, demography and reproductive biology. In: Young A.G. and Clarke G.M. (eds), Genetics, Demography and Viability of Fragmented Populations. Cambridge University Press, Cambridge, UK, pp. 313-334. Paine R.T. 1988. Habitat suitability and local population persistence of the sea palm Postelsia palmaeformis. Ecology 69: 1787-1794. Patterson B.D. 1984. Mammalian extinctions and biogeography in the Southern Rocky Mountains. In: Nitecki M. (ed), Extinctions. University of Chicago Press, Chicago, Illinois, pp. 247-293. Patterson B.D. 1987. The principle of nested subsets and its implications for biological conservation. Conservation Biology 1: 323-334. Pimm S.L. 1991. The Balance of Nature? University of Chicago Press, Chicago, Illinois. PimmS.L. and Lawton J.H. 1977. Number of trophic levels in ecological communities. Nature 268: 329-331. Pimm S.L., Jones H.L. and Diamond J. 1988. On the risk of extinction. The American Naturalist 132: 757-785. Pimm S.L., Diamond J., Reed T.M., Russell G.J. and Verner J. 1993. Times to extinction for small populations of large birds. Proceedings of the National Academy of Science USA 90: 10871-10875. Pollard E. and Yates T.J. 1992. The extinction and foundation of local butterfly populations in relation to population variability and other factors. Ecological Entomology 17: 249-254. Poschlod P., Kiefer S., Tränkle U., Fischer S. and Bonn S. 1998. Plant species richness in calcareous grasslands as affected by dispersability in space and time. Applied Vegetation Science 1: 75-90. Poschlod P., Kleyer M. and Tackenberg O. 2000. Data bases on life history traits as a tool for risk assessment in plant species. Zeitschrift für Ökologie und Naturschutz 9: 3-18. Preston F.W. 1949. The commonness and rarity of species. Ecology 29: 254-283. Quinn J.F., Wolin C.L. and Judge M.L. 1989. An experimental analysis of patch size, habitat subdivision, and extinction in a marine intertidal snail. Conservation Biology 3: 242-251. Rabinowitz D., Cairns S. and Dillon T. 1986. Seven forms of rarity and their frequency in the flora of the British Isles. In: Soulé M.E. (ed), Conservation Biology: The Science of Scarcity and Diversity. Sinauer Associates, Sunderland, Massachusetts, pp. 182-204. Recher H.F., Shields J., Kavanagh R. and Webb G. 1987. Retaining remnant mature forest for nature conservation at Eden, New South Wales: a review of theory and practice. In: Saunders D.A., Arnold G.W., Burbidge A.A. and Hopkins A.J.M. (eds), Nature Conservation: The Role of Remnants of Native Vegetation. Surrey Beatty, Chipping Norton, Australia, pp. 177-194. Reed J.M. 1999. The role of behavior in recent avian extinctions and endangerments. Conservation Biology 13: 232-241. Restrepo C., Renjifo L.M. and Marples P. 1997. Frugivorous birds in fragmented Neotropical montane forests: landscape pattern and body mass distribution. In: Laurance W.F. and Bierregaard R.O. (eds), Tropical Forest Remnants. University of Chicago Press, Chicago, Illinois, pp. 171-189. Ricklefs R.E. and Latham R.E. 1992. Intercontinental correlation of geographic ranges suggests stasis in ecological traits of relict genera of temperate perennial herbs. The American Naturalist 139: 1305-1321. Robinson G.R. and Quinn J.F. 1988. Extinction, turnover and species diversity in an experimentally fragmented Californian annual grassland. Oecologia 76: 71-82. Robinson G.R., Holt R.D., Gaines M.S., Hamburg S.P., Johnson M.L., Fitch H.S. et al. 1992. Diverse and contrasting effects of habitat fragmentation. Science 257: 524-526. Roff D.A. 1992. The Evolution of Life Histories: Theory and Analysis. Chapman & Hall, New York. Roland J., Keyghobadi N. and Fownes S. 2000. Alpine Parnassius butterfly dispersal: effects of landscape and population size. Ecology 81: 1642-1653. Rosenberg K.V., Lowe J.D. and Dhondt A.A. 1999. Effects of forest fragmentation on breeding tanagers: a continental perspective. Conservation Biology 13: 568-583. Rosenzweig M.L. and Clark C.W. 1994. Island extinction rates from regular censuses. Conservation Biology 8: 491-494. Salisbury E. 1976. Seed output and the efficacy of dispersal by wind. Proceedings of the Royal Society London 192: 323-329. Samietz J., Berger U. and Köhler G. 1996. A population vulnerability analysis of the stripewinged grasshopper, Stenobothrus lineatus (Caelifera: Acrididae). In: Settele J., Margules C.R., Poschlod P. and Henle K. (eds), Species Survival in Fragmented Landscapes. Kluwer, Dordrecht, The Netherlands, pp. 299-311. Sarre S., Smith G.T. and Meyers J.A. 1995. Persistence of two species of gecko (Oedura reticulata and Gehyra variegata) in remnant habitat. Biological Conservation 71: 25-33. Sarre S., Wiegand K. and Henle K. 1996. Survival of a specialist and generalist gecko in the fragmented landscape of the Western Australian wheatbelt. In: Settele J., Margules C.R., Poschlod P. and Henle K. (eds), Survival of Species in Fragmented Landscapes. Kluwer, Dordrecht, The Netherlands, pp. 39-51. Saunders D.A. 1996. Habitat networks in the fragmented landscape of the Western Australian wheatbelt: preliminary results, involvement with landcare groups, and experience in implementation. In: Settele J., Margules C.R., Poschlod P. and Henle K. (eds), Species Survival in Fragmented Landscapes. Kluwer, Dordrecht, The Netherlands, pp. 69-80. Saunders D.A., Hobbs R.J. and Margules C.R. 1991. Biological consequences of ecosystem fragmentation: a review. Conservation Biology 5: 18-32. Schoener T.W. and Schoener A. 1983. The time to extinction of a colonizing propagule of lizards increases with island area. Nature 302: 332-334. Schoener T.W. and Spiller D.A. 1987. High population persistence in a system with high turnover. Nature 330: 474-477. Schoener T.W. and Spiller D.A. 1992. Is extinction rate related to temporal variability in population size? An empirical answer for orb spiders. The American Naturalist 139: 1176-1207. Schwartzman S., Moreira A. and Nepstad D. 2000. Rethinking tropical forest conservation: perils in parks. Conservation Biology 14: 1351-1357. Settele J. 1998. Metapopulationsanalyse auf Rasterdatenbasis. Teubner, Stuttgart, Germany. Settele J. and Poethke H.J. 1996. Towards demographic vulnerability categories of butter-flies. In: Settele J., Margules C.R., Poschlod P. and Henle K. (eds), Species Survival in Fragmented Landscapes. Kluwer, Dordrecht, The Netherlands, pp. 282-289. Shaffer M.L. 1981. Minimum population sizes for species conservation. BioScience 31: 131-134. Sieving K.E. and Karr J.R. 1997. Avian extinction and persistance mechanisms in lowland Panama. In: Laurance W.F. and Bierregaard R.O. (eds), Tropical Forest Remnants. University of Chicago Press, Chicago, Illinois, pp. 156-170. Soulé M.E., Bolger D.T., Alberts A.C., Sauvajot R.S., Wright J., Sorice M. et al. 1988. Reconstructed dynamics of rapid extinctions of chaparral-requiring birds in urban habitat islands. Conservation Biology 2: 75-92. Stearns S.C. 1976. Life history tactics: a review of the ideas. Quarterly Review of Biology 51: 3-47. Stöcklin J. and Fischer M. 1997. Local extinctions of plants in remnants of extensively used calcareous grasslands 1950-1985. Conservation Biology 11: 727-737. Stouffer P.C. and Bierregaard R.O. 1995. Effects of forest fragmentation on understory hummingbirds in Amazonian Brazil. Conservation Biology 9: 1085-1094. Strykstra R.J., Verweij G.L. and Bakker J.P. 1997. Seed dispersal by mowing machinery in a Dutch brook valley system. Acta Botanica Neerlandica 46: 387-401. Tabarelli M., Mantovani W. and Peres C.A. 1999. Effects of habitat fragmentation on plant guild structure in the montane forest of southeastern Brazil. Biological Conservation 91: 119-127. Temple S.A. and Carey J.R. 1988. Modeling dynamics of habitat-interior bird populations in fragmented landscapes. Conservation Biology 2: 340-347. Terborgh J. 1974. Preservation of natural diversity: the problem of extinction prone species. BioScience 24: 715-722. Terborgh J. 1999. Requiem for Nature. Island Press, Washington, DC. Terborgh J. and Winter B. 1980. Some causes of extinction. In: Soulé M.E. and Wilcox B.A. (eds), Conservation Biology: An Evolutionary-Ecological Perspective. Sinauer, Sunderland, Massachusetts, pp. 119-134. Terborgh J., Lopez L., Tello J., Yu D. and Bruni A.R. 1997. Transitory states in relaxing ecosystems of land bridge islands. In: Laurance W.F. and Bierregaard R.O. (eds), Tropical Forest Remnants. University of Chicago Press, Chicago, Illinois, pp. 256-274. Thomas J.A., Moss D. and Pollard E. 1994. Increased fluctuations of butterfly populations towards the northern edges of species ranges. Ecography 17: 215-220. Thompson K., Hillier S.H., Grime J.P., Bossard C.C. and Band S.R. 1996. A functional analysis of a limestone grassland community. Journal of Vegetation Science 7: 371-380. Tilman D. 1997. Community invasibility, recruitment limitation, and grassland biodiversity. Ecology 78: 81-92. Tilman D., Lehman C.L. and Yin C.J. 1997. Habitat destruction, dispersal, and deterministic extinction in competitive communities. The American Naturalist 149: 407-435. Tracy C.R. and George T.L. 1992. On the determinants of extinction. The American Naturalist 139: 102-122. Troll W. 1937-1943. Vergleichende Morphologie der höheren Pflanzen. Vol. 1: Vegetationsorgane, Part 1-3. Borntraeger, Berlin, Germany. Tullberg B.S. and Hunter A.F. 1996. Evolution of larval gregariousness in relation to repellent defences and warning coloration in tree-feeding Macrolepidoptera: a phylogenetic analysis based on independent contrasts. Biological Journal of the Linnean Society 57: 253-276. Usher M.B. 1987. Effects of fragmentation on communities and populations: a review with applications to wildlife conservation. In: Saunders D.A., Arnold G.W., Burbidge A.A. and Hopkins A.J.M. (eds), Nature Conservation: The Role of Remnants of Native Vegetation. Surrey Beatty, Chipping Norton, Australia, pp. 103-121. Vos C.C., Verboom J., Opdam P.F.M. and Ter Braak C.J.F. 2001. Toward ecologically scaled landscape indices. The American Naturalist 183: 24-41. Vucetich J.A., Waite T.A., Qvarnemark L. and Ibargüen S. 2000. Population variability and extinction risk. Conservation Biology 14: 1704-1714. Warburton N.H. 1997. Structure and conservation of forest avifauna in isolated rainforest remnants in tropical Australia. In: Laurance W.F. and Bierregaard R.O. (eds), Tropical Forest Remnants. University of Chicago Press, Chicago, Illinois, pp. 190-206. Warner R.R. and Chesson P.L. 1985. Coexistence mediated by recruitment fluctuations: a field guide to the storage effect. The American Naturalist 125: 769-787. Wehausen J.D. 1999. Rapid extinction of mountain sheep populations revisited. Conservation Biology 13: 378-384. Whitcomb R.F. 1987. North American forests and grasslands: biotic conservation. In: Saunders D.A., Arnold G.W., Burbidge A.A. and Hopkins A.J.M. (eds), Nature Conservation: The Role of Remnants of Native Vegetation. Surrey Beatty, Chipping Norton, Australia, pp. 163-176. Wiegand K., Sarre S., Henle K., Stephan T., Wissel C. and Brandl R.M. 2001. Demographic stochasticity does not predict persistence of gecko populations. Ecological Applications 11: 1738-1749. Wilcove D.S., McLellan C.H. and Dobson A.P. 1986. Habitat fragmentation in the temperate zone. In: Soulé M.E. (ed), Conservation Biology. The Science of Scarcity and Diversity. Sinauer, Sunderland, Massachusetts, pp. 237-256. Willis E.O. 1979. The composition of avian communities in remanescent woodlots in southern Brazil. Papeis Avulsos de Zoologia 33: 1-25. Wissel C., Stephan T. and Zaschke S. 1994. Modelling extinction and survival of small populations. In: Remmert H. (eds), Minimum Viable Populations. Springer, Heidelberg, Germany, pp. 67-103. With K.A. and King A.W. 1999. Extinction thresholds for species in fractal landscapes. Conservation Biology 13: 314-326. Woodroffe R. and Ginsberg J.R. 1998. Edge effects and the extinction of populations inside protected areas. Science 280: 2126-2128.