Towards a more general species–area relationship: diversity on all islands, great and small

Aim To demonstrate a new and more general model of the species–area relationship that builds on traditional models, but includes the provision that richness may vary independently of island area on relatively small islands (the small island effect). Location We analysed species–area patterns for a broad diversity of insular biotas from aquatic and terrestrial archipelagoes. Methods We used breakpoint or piecewise regression methods by adding an additional term (the breakpoint transformation) to traditional species–area models. The resultant, more general, species–area model has three readily interpretable, biologically relevant parameters: (1) the upper limit of the small island effect (SIE), (2) an estimate of richness for relatively small islands and (3) the slope of the species–area relationship (in semi-log or log–log space) for relatively large islands. Results The SIE, albeit of varying magnitude depending on the biotas in question, appeared to be a relatively common feature of the data sets we studied. The upper limit of the SIE tended to be highest for species groups with relatively high resource requirements and low dispersal abilities, and for biotas of more isolated archipelagoes. Main conclusions The breakpoint species–area model can be used to test for the significance, and to explore patterns of variation in small island effects, and to estimate slopes of the species–area (semi-log or log–log) relationship after adjusting for SIE. Moreover, the breakpoint species–area model can be expanded to investigate three fundamentally different realms of the species–area relationship: (1) small islands where species richness varies independent of area, but with idiosyncratic differences among islands and with catastrophic events such as hurricanes, (2) islands beyond the upper limit of SIE where richness varies in a more deterministic and predictable manner with island area and associated, ecological factors and (3) islands large enough to provide the internal geographical isolation (large rivers, mountains and other barriers within islands) necessary for in situ speciation.

[1]  Michael L. Rosenzweig,et al.  Species Diversity in Space and Time , 1997 .

[2]  H. Gleason,et al.  On the Relation Between Species and Area , 1922 .

[3]  J. Cook,et al.  Mammalian biogeography of the Alexander Archipelago of Alaska: a north temperate nested fauna , 1999 .

[4]  R. Cowie.  VARIATION IN SPECIES DIVERSITY AND SHELL SHAPE IN HAWAIIAN LAND SNAILS: IN SITU SPECIATION AND ECOLOGICAL RELATIONSHIPS , 1995, Evolution; international journal of organic evolution.

[5]  Mark V. Lomolino,et al.  Ecology’s most general, yet protean 1 pattern: the species‐area relationship , 2000 .

[6]  J. Neter,et al.  Applied Linear Regression Models , 1983 .

[7]  S. Wright.  Intra-Archipelago Vertebrate Distributions: The Slope of the Species-Area Relation , 1981, The American Naturalist.

[8]  D. Bloch,et al.  Island land bird population densities in relation to island size and habitat quality on the Faroe Islands , 1983 .

[9]  Mark V. Lomolino.  Mammalian community structure on islands: the importance of immigration, extinction and interactive effects , 1986 .

[10]  G. R. Williams.  Aspects of Avian Island Biogeography in New Zealand , 1981 .

[11]  David P. Olson,et al.  Small mammal populations on the islands of Basswood Lake, Minnesota , 1954 .

[12]  M. Rosenzweig.  Species diversity in space and time: Coevolution of habitat diversity and species diversity , 1995 .

[13]  Herold J. Wiens.  Atoll environment and ecology , 1963 .

[14]  James H. Brown,et al.  Island Biogeograhy of Montane Forest Mammals in the American Southwest , 1989 .

[15]  C. Meine.  It’s about Time: Conservation Biology and History , 1999 .

[16]  J. Heller.  The Biogeography of Enid Landsnails on the Aegean Islands , 1976 .

[17]  L F Bersier,et al.  Scaling regions for food web properties. , 1997, Proceedings of the National Academy of Sciences of the United States of America.

[18]  R. Macarthur,et al.  The Theory of Island Biogeography , 1969 .

[19]  Nicholas J. Gotelli,et al.  Statistical Distributions of West Indian Land Bird Families , 1982 .

[20]  Thomas M. Brooks,et al.  Time Lag between Deforestation and Bird Extinction in Tropical Forest Fragments , 1999 .

[21]  B. Wiman,et al.  Serf and waif colonization: distribution and dispersal of invertebrate species in Faroe island settlement areas , 1987 .

[22]  J. Downhower.  The biogeography of the island region of western Lake Erie , 1988 .

[23]  C. L. Shafer,et al.  NATURE RESERVES - Island Theory and Conservation Practice , 1991 .

[24]  James H. Brown,et al.  Turnover Rates in Insular Biogeography: Effect of Immigration on Extinction , 1977 .

[25]  R. Macarthur,et al.  AN EQUILIBRIUM THEORY OF INSULAR ZOOGEOGRAPHY , 1963 .

[26]  Ecological and Evolutionary Determinants of the Species-Area Relation in Caribbean Anoline Lizards , 1996 .

[27]  Edward O. Wilson,et al.  The Nature of the Taxon Cycle in the Melanesian Ant Fauna , 1961, The American Naturalist.

[28]  O. Järvinen,et al.  Colonization of islands by land birds: prevalence functions in a Finnish archipelago , 1983 .

[29]  D. Kohn,et al.  Plant species richness: the effect of island size and habitat diversity , 1994 .

[30]  N. K. Johnson.  CONTROLS OF NUMBER OF BIRD SPECIES ON MONTANE ISLANDS IN THE GREAT BASIN , 1975, Evolution; international journal of organic evolution.

[31]  P. Raven,et al.  Species Number and Endemism: The Gal�pagos Archipelago Revisited , 1973, Science.

[32]  H. Rydin,et al.  Plant species richness on islands over a century of primary succession : Lake Hjälmaren , 1988 .

[33]  P. Morisset,et al.  Floristic richness, area, and habitat diversity in a hemiarctic archipelago , 1988 .

[34]  F. Vuilleumier.  Insular Biogeography in Continental Regions. I. The Northern Andes of South America , 1970, The American Naturalist.

[35]  O. Järvinen,et al.  Dynamics of Protected Bird Communities in a Finnish Archipelago , 1977 .

[36]  D. Srivastava.  Using local–regional richness plots to test for species saturation: pitfalls and potentials , 1999 .

[37]  K. McGuinness.  Species–area curves , 1984 .

[38]  D. Schluter,et al.  Analysis of an evolutionary species–area relationship , 2000, Nature.

[39]  T. Case.  Species Numbers, Density Compensation, and Colonizing Ability of Lizards on Islands in the Gulf of California , 1975 .

[40]  James H. Brown.  Mammals on Mountaintops: Nonequilibrium Insular Biogeography , 1971, The American Naturalist.

[41]  C. Roden.  Persistence, extinction and different species pools within the flora of lake islands in western Ireland , 1998 .

[42]  S. As.  To Fly or Not to Fly? Colonization of Baltic Islands by Winged and Wingless Carabid Beetles , 1984 .

[43]  Mark V. Lomolino,et al.  Island biogeography and landscape ecology of mammals inhabiting fragmented, temperate rain forests , 2001 .

[44]  K. Crowell.  A comparison of relict versus equilibrium models for insular mammals of the Gulf of Maine , 1986 .

[45]  Daniel Simberloff,et al.  Species Number and Compositional Similarity of the Galapagos Flora and Avifauna , 1978 .

[46]  D. Simberloff,et al.  Environmental Determinants of Island Species Numbers in the British Isles , 1974 .

[47]  S. Sfenthourakis.  The Species-Area Relationship of Terrestrial Isopods (Isopoda; Oniscidea) from the Aegean Archipelago (Greece): A Comparative Study , 1996 .

[48]  D. Western,et al.  The future of the savannah ecosystems: ecological islands or faunal enclaves? , 1981 .

[49]  Island Biogeography, Theory Of , 1982 .

[50]  L. Heaney.  Dynamic disequilibrium: a long-term, large-scale perspective on the equilibrium model of island biogeography , 2000 .

[51]  C. Woodroffe.  Vascular plant species‐area relationships on Nui Atoll, Tuvalu, Central Pacific: a reassessment of the small island effect , 1986 .

[52]  J. Montague.  The Ecology of Hawaiian Flower-Breeding Drosophilids. 2. Adult Dispersions and Reproductive Ecology , 1989, The American Naturalist.

[53]  Mark V. Lomolino,et al.  ASSEMBLY AND DISASSEMBLY OF MAMMAL COMMUNITIES IN A FRAGMENTED TEMPERATE RAIN FOREST , 2000 .

[54]  T. Brooks,et al.  Deforestation Predicts the Number of Threatened Birds in Insular Southeast Asia , 1997 .

[55]  Mark V. Lomolino,et al.  Species Diversity in Space and Time. , 1996 .

[56]  T. E. Martin.  Species-Area Slopes and Coefficients: A Caution on Their Interpretation , 1981, The American Naturalist.

[57]  S. M. Glenn.  Regional analysis of mammal distributions among Canadian parks: implications for park planning , 1990 .

[58]  O. Arrhenius,et al.  Species and Area , 1921 .

[59]  R. Buckley,et al.  Distinguishing the Effects of Area and Habitat Type on Island Plant Species Richness by Separating Floristic Elements and Substrate Types and Controlling for Island Isolation , 1985 .

[60]  Mark V. Lomolino,et al.  CORRIDORS AND MAMMAL COMMUNITY STRUCTURE ACROSS A FRAGMENTED, OLD‐GROWTH FOREST LANDSCAPE , 2000 .

[61]  W. Newmark.  Species‐area relationship and its determinants for mammals in western North American national parks , 1986 .

[62]  M. Willig,et al.  Randomness, Area, and Species Richness , 1982 .

[63]  G. Perry,et al.  The lizard fauna of Guam's fringing islets: Island biogeography, phylogenetic history, and conservation implications , 1998 .

[64]  A. Kratter.  Montane avian biogeography in southern California and Baja California , 1992 .

[65]  James H. Brown.  The theory of insular biogeography and the distribution of boreal birds and mammals , 1978 .

[66]  A. Amerson.  Species Richness on the Nondisturbed Northwestern Hawaiian Islands , 1975 .

[67]  F. Stearns,et al.  Mammals in Forest Islands in Southeastern Wisconsin , 1981 .

[68]  Mark V. Lomolino.  Species Richness of Mammals Inhabiting Nearshore Archipelagoes: Area, Isolation, and Immigration Filters , 1994 .

[69]  Mark V. Lomolino.  The target area hypothesis: the influence of island area on immigration rates of non-volant mammals , 1990 .

[70]  M E Gilpin,et al.  Calculation of immigration and extinction curves from the species-area-distance relation. , 1976, Proceedings of the National Academy of Sciences of the United States of America.

[71]  A. Harcourt.  BIOGEOGRAPHIC RELATIONSHIPS OF PRIMATES ON SOUTH-EAST ASIAN ISLANDS , 1999 .

[72]  F. W. Preston.  The Canonical Distribution of Commonness and Rarity: Part I , 1962 .

[73]  Ian Abbott,et al.  Changes in Species Composition of Floras on Islets Near Perth, Western Australia , 1980 .

[74]  Ted J. Case,et al.  Island biogeography in the Sea of Cortéz , 1983 .

[75]  A. Amerson.  SPECIES RICHNESS ON THE NONDISTURBED NORTHWESTERN , 1975 .

[76]  R. Macarthur.  Species packing and competitive equilibrium for many species. , 1970, Theoretical population biology.

[77]  J. Bengtsson,et al.  Habitat diversity or area per se? species richness of woody plants, carabid beetles and land snails on Islands , 1988 .

[78]  Earl D. McCoy,et al.  The Statistics and Biology of the Species-Area Relationship , 1979, The American Naturalist.

[79]  J. Boomsma,et al.  Insular biogeography and distribution ecology of ants on the Frisian islands , 1987 .

[80]  M. Gilpin,et al.  Metapopulation Biology: Ecology, Genetics, and Evolution , 1997 .

[81]  E. Ranta,et al.  Distribution of carabid beetles in four boreal archipelagoes , 1987 .

[82]  J. Diamond,et al.  Distributional ecology of new hebridean birds a species kaleidoscope , 1977 .

[83]  N. Davies,et al.  MUNROE REVISITED : A SURVEY OF WEST INDIAN BUTTERFLY FAUNAS AND THEIR SPECIES-AREA RELATIONSHIP , 1998 .

[84]  T. Reed.  The Number of Breeding Landbird Species on British Islands , 1981 .

[85]  G. Sugihara.  S = CAz, z ≃ 1/4: A Reply to Connor and McCoy , 1981, The American Naturalist.

[86]  S. Pimm,et al.  Threat from deforestation to montane and lowland birds and mammals in insular South‐east Asia , 1999 .

[87]  Mark V. Lomolino.  Interpretations and Comparisons of Constants in the Species-Area Relationship: An Additional Caution , 1989, The American Naturalist.

[88]  T. Crowe.  Lots of weeds: insular phytogeography of vacant urban lots , 1979 .

[89]  T. Lawlor.  Comparative biogeography of mammals on islands , 1986 .

[90]  I. Abbott.  Biogeography of grasses (Poaceae) on islands of southwestern Australia , 1992 .

[91]  William A. Niering,et al.  Terrestrial Ecology of Kapingamarangi Atoll, Caroline Islands , 1963 .

[92]  M. Oldham,et al.  Historic and Current Amphibian and Reptile Distributions in the Island Region of Western Lake Erie , 1997 .