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The Theory of Insular Biogeography and the Distribution of Boreal Birds and Mammals James H

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The Theory of Insular Biogeography and the Distribution of Boreal Birds and Mammals James H Great Basin Naturalist Memoirs Volume 2 Intermountain Biogeography: A Symposium Article 14 3-1-1978 The theory of insular biogeography and the distribution of boreal birds and mammals James H. Brown Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, Arizona 85721 Follow this and additional works at: https://scholarsarchive.byu.edu/gbnm Recommended Citation Brown, James H. (1978) "The theory of insular biogeography and the distribution of boreal birds and mammals," Great Basin Naturalist Memoirs: Vol. 2 , Article 14. Available at: https://scholarsarchive.byu.edu/gbnm/vol2/iss1/14 This Article is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Great Basin Naturalist Memoirs by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. THE THEORY OF INSULAR BIOGEOGRAPHY AND THE DISTRIBUTION OF BOREAL BIRDS AND MAMMALS James H. Brown 1 Abstract.—The present paper compares the distribution of boreal birds and mammals among the isolated mountain ranges of the Great Basin and relates those patterns to the developing theory of insular biogeography. The results indicate that the distribution of permanent resident bird species represents an approximate equilib- rium between contemporary rates of colonization and extinction. A shallow slope of the species-area curve (Z = 0.165), no significant reduction in numbers of species as a function of insular isolation (distance to nearest conti- nent), and a strong dependence of species diversity on habitat diversity all suggest that immigration rates of bo- real birds are sufficiently high to maintain populations on almost all islands where there are appropriate habitats. In contrast, the insular faunas of boreal mammals represent relictual populations that receive no significant con- temporary immigration. The insular mammal faunas have been derived by extinction from a set of species that colonized the islands when habitat bridges connected them to the continents in the late Pleistocene. A relatively steep species-area curve (Z = 0.326), no effect of isolation on species diversity, and the absence of appropriate species from large areas of apparently suitable habitat all support this conclusion. Measures of habitat diversity that are closely correlated with bird species diversity do not account for much of the variation in number of mammal species among islands. Insular area is the single variable that accounts for most of the variability in both bird and mammal species diversity; this supports the approach of using standard parameters such as area in comparative empirical analyses and general biogeographic theory. The results of this study suggest that extremes of vagility among taxa and a recent history of paleoclimatic and geological changes make it unlikely that equi- librial distributions, of the sort MacArthur and Wilson (1967) propose for the biotas of oceanic islands, are char- acteristic of the insular distributions of terrestial and freshwater vertebrates of western North America. Biogeography is an old science that has draw quantitative relationships and derive made recent advances by assimilating new general principles. concepts and data. Within the last two dec- With the publication of their equilibrium ades information on continental drift, pa- model of insular distribution, MacArthur leoclimatology, and ancient sea levels has and Wilson (1963, 1967) contributed not revolutionized our understanding of histori- only a new theory but also a new approach cal events and their effects on plant and an- to biogeography. Prior to their work, most imal distribution. During the same period, biogeographic research had consisted of de- ecologists and evolutionary biologists have scribing the distributions of particular taxa learned much about the processes of popu- and producing ad hoc, historical explana- lation growth, dispursal, extincton, speci- tions. MacArthur and Wilson advocated a ation, and interspecific interactions which quantitative approach designed to build and are the mechanisms that determine distribu- test general models based on ecological pro- tion. Biogeography appears to be entering cesses. The specific model that they pro- an exciting new period in which the volu- posed suggests that the number of species minous data acquired by systematists and inhabiting an island represents an equilib- descriptive biogeographers can be inter- rium between opposing rates of extinction processes preted in terms of recently understood his- and colonization, and that these torical events and ecological processes to are functions of the size of an island and its 'Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, Ariz 209 210 GREAT BASIN NATURALIST MEMOIRS No. 2 distance from a source of colonists, respec- 1946, Miller and Bussell 1956, Van Bos- tively. sem 1936; for mammals Durrant 1952, Dur- MacArthur and Wilsons's choice of is- rant et al. 1955, Grinnell 1933, Hall 1946). lands for their revolutionary approach to The paleoclimatic history of the region also biogeography was not a fortuitous one. Ever is becoming increasingly well understood since the pioneering work of Darwin and (e.g., Hubbs and Miller 1948, Martin and Wallace, islands have played a preeminent Mehringer 1965, Smith this symposium, role in the development of the science. It is Wells and Berger 1967, Wells and Jorgen- difficult to do experiments in biogeography, sen 1964). This excellent data base has been but islands and insular habitats represent used in quantitative analyses of the insular natural experiments; they are small, replica- distributions of lacustrine fishes (Barbour ted systems among which species and envi- and Brown 1974) and montane birds (John- ronmental parameters are distributed in dif- son 1975) and mammals (Brown 1971). ferent combinations. Often particular taxa Much more work remains to be done. Some are distributed among insular habitats in of the most interesting contributions can be patterns that imply the operation of general expected when additional kinds of organ- mechanisms of dispersal, extinction, and in- isms and insular habitats are studied, so that terspecific interaction. The last decade has it is possible to make comparisons among seen several attempts to test the model of taxa which differ in ecological requirements MacArthur and Wilson, using a variety of and dispersal abilities, and among habitats organisms that inhabit both true islands and that differ in environmental parameters and analogous isolated habitats (e.g., Barbour history of isolation. and Brown 1974, Brown 1971, Brown and The present paper discusses the distribu- Kodric-Brown in review, Culver 1970, Cul- tion of boreal birds and mammals among ver et al. 1973, Diamond 1969, 1970a, isolated mountain ranges in the Great Basin 1971, Johnson 1975, Harper et al. this sym- in relation to the theory of insular bio- posium, Schoener 1974, Seifert 1975, Sim- geography. It attempts to relate distribu- berloff 1974, Simberloff and Wilson 1970, tional patterns to mechanisms of dispersal Simpson 1974, Terborgh 1973, Vuilleumier and extinction and to differentiate relictual 1970, 1973). Not all of these studies have patterns that are the legacy of historical supported the model, but the exceptions events from equilibrial ones that can be at- have contributed importantly to our under- tributed to contemporary ecological pro- standing of the patterns of insular distribu- cesses. The paper tries to develop a general tion and the historical events and ecological conceptual basis for analyzing and predict- processes that produce them. ing insular distributions. It first discusses the Analysis of the distribution of vertebrates current state of insular biogeographic theo- among isolated habitats in the Inter- ry and then utilizes empirical data on the mountain Begion of western North America distribution of boreal birds and mammals to has contributed significantly to the devel- test the theory and search for general mech- opment and testing of biogeographic theory. The dedicated field work of several gener- ations of systematists has documented in de- The Theory of Insular Biogeography tail the species distributions of most verte- brate groups (e.g., for fishes Miller 1948, Colonization, extinction, and speciation Hubbs and Miller 1948, Hubbs et al. 1974, are the primary processes that determine Smith, this symposium; for birds Behle the composition of insular biotas. An island 1943, 1955, 1958, this symposium, Grinnell can be defined as a patch of suitable habitat and Miller 1944, Johnson 1965, 1970, 1973, surrounded by unfavorable environment that 1974, 1975, Linsdale 1936, Miller 1935, limits the dispersal of individuals. New spe- 1978 INTERMOUNTAIN BIOGEOGRAPHY: A SYMPOSIUM 211 cies colonize an island either by dispersing tion of island size. The model predicts: first, across the habitat barriers or by immigrat- that relatively constant insular species di- ing sometime when the barriers were tem- versity is maintained by the continual turn- porarily absent. Speciation entirely within over (extinction and colonization) of individ- an island potentially is another source of ual species; second, that the equilibrium new species, but there is no evidence that number of species is positively correlated this process has contributed significantly to with island size and negatively correlated the diversity
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