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Body Size of Insular Carnivores: Little Support for the Island Rule. Author(S): Shai Meiri, Tamar Dayan, and Daniel Simberloff Source: the American Naturalist, Vol

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Body Size of Insular Carnivores: Little Support for the Island Rule. Author(S): Shai Meiri, Tamar Dayan, and Daniel Simberloff Source: the American Naturalist, Vol The University of Chicago Body Size of Insular Carnivores: Little Support for the Island Rule. Author(s): Shai Meiri, Tamar Dayan, and Daniel Simberloff Source: The American Naturalist, Vol. 163, No. 3 (March 2004), pp. 469-479 Published by: The University of Chicago Press for The American Society of Naturalists Stable URL: http://www.jstor.org/stable/10.1086/382229 . Accessed: 15/04/2013 15:37 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. The University of Chicago Press, The American Society of Naturalists, The University of Chicago are collaborating with JSTOR to digitize, preserve and extend access to The American Naturalist. http://www.jstor.org This content downloaded from 129.11.21.2 on Mon, 15 Apr 2013 15:37:08 PM All use subject to JSTOR Terms and Conditions vol. 163, no. 3 the american naturalist march 2004 ൴ Body Size of Insular Carnivores: Little Support for the Island Rule Shai Meiri,1,* Tamar Dayan,1,† and Daniel Simberloff 2,‡ 1. Department of Zoology, Tel Aviv University, Tel Aviv 69978, introduce exotic species) make islands an excellent arena Israel; for evolutionary research. 2. Department of Ecology and Evolutionary Biology, University of Among the most pronounced microevolutionary Tennessee, Knoxville, Tennessee 37996 changes occurring on islands are changes in body size. Submitted February 20, 2003; Accepted August 12, 2003; These changes are especially apparent in mammals (but Electronically published March 9, 2004 by no means only in mammals; Case 1978; Pregil 1986; Alcover and McMinn 1994; McNab 1994; Benton et al. Online enhancements: tables, literature cited. 1997; Brown and Lomolino 1998; Jianu and Weishampel 1999; Clegg and Owens 2002). Foster (1964) conducted the first systematic analysis of patterns in body size evolution of insular mammals. Sur- abstract: Large mammals are thought to evolve to be smaller on veying the literature, he found that rodents tend toward islands, whereas small mammals grow larger. A negative correlation gigantism on islands, while lagomorphs, carnivores, and between relative size of island individuals and body mass is termed artiodactyls are usually characterized by insular dwarfing. the “island rule.” Several mechanisms—mainly competitive release, Van Valen named these phenomena the “island rule” (Van resource limitation, dispersal ability, and lighter predation pressure on islands, as well as a general physiological advantage of modal Valen 1973, p. 32), according to which small mammals size—have been advanced to explain this pattern. We measured skulls grow larger on islands while large mammals are dwarfed. and teeth of terrestrial members of the order Carnivora in order to Van Valen (1973, p. 72) concluded, “The regular evolution analyze patterns of body size evolution between insular populations of mammalian body size on islands is an extraordinary and their near mainland conspecifics. No correlations were found phenomenon which seems to have fewer exceptions than between the size ratios of insular/mainland carnivore species and any other ecotypic rule in animals.” Several authors (Case body mass. Only little support for the island rule is found when individual populations rather than species are considered. Our data 1978; Heaney 1978; Lawlor 1982; Melton 1982; Davis 1983; are at odds with those advanced in support of theories of optimal Angerbjo¨rn 1985) raised interesting hypotheses about the body size. Carnivore size is subjected to a host of selective pressures selective pressures underlying these patterns, highlighting that do not vary uniformly from place to place. Mass alone cannot such agents as resource limitation, dispersal ability, com- account for the patterns in body size of insular carnivores. petition (or lack thereof), predation, and territoriality. Keywords: body size, Carnivora, geographic variation, island rule, Lomolino’s (1983, 1985) review expanded Foster’s da- optimal body size. tabase to include 375 populations representing more than 74 mammalian species (seven comparisons included in- dividuals of more than one species; Lomolino 1983). Islands have served as models in the study of evolutionary Lomolino reinforced Foster’s conclusions as to the fre- and ecological phenomena ever since Darwin (1845; Wal- quency of dwarfism and gigantism in various mammalian lace 1868, 1880; MacArthur and Wilson 1967). Their iso- orders. He then represented the relative size of the insular lation, the relatively low species richness, and, in many forms (the ratio between a species insular and mainland cases, their clear faunal history (such as when humans sizes, averaged for all populations) as a function of body mass, obtaining a graded trend from gigantism in the * E-mail: [email protected]. smaller species to dwarfism in the larger ones. Lomolino † E-mail: [email protected]. (1985, p. 314) reformulated the island rule in these terms, ‡ E-mail: [email protected]. and this is the accepted modern form of the rule. Am. Nat. 2004. Vol. 163, pp. 469–479. ᭧ 2004 by The University of Chicago. The clear, fascinating pattern observed by Lomolino 0003-0147/2004/16303-30069$15.00. All rights reserved. (1985) and the elegant models of Heaney (1978) and Case This content downloaded from 129.11.21.2 on Mon, 15 Apr 2013 15:37:08 PM All use subject to JSTOR Terms and Conditions 470 The American Naturalist (1978) have been widely discussed in the ecological and Material and Methods evolutionary literature. For example, they were advanced as support for the concept of an optimal mammalian body In order to analyze body size variation in insular and size (Brown et al. 1993; Damuth 1993). They have fostered mainland carnivores, we measured skulls of carnivores in controversy over the possibility of different evolutionary the following collections: Natural History Museum, Lon- patterns and forces for mammals and birds (Clegg and don; Zoology Museum of Cambridge University; New Owen 2002). Furthermore, they have spawned a host of Walk Museum; National Monuments Archeozoological partially contradictory explanations for the perceived rule collections, London; Harrison Zoological Museum; the based on ecology (reviewed by Angerbjo¨rn 1986; Dayan Royal Museum, Edinburgh; Muse´um National d’Histoire and Simberloff 1998), paleobiology (Gordon 1986), phys- Naturelle, Paris; Laboratoire d’Anatomie Compare´e, Paris; iology (Lovegrove 2000), and evolution (Demetrius 2000). Institut Royal des Sciences Naturelles de Belgique; Na- We aimed to see whether patterns of size in members tional Museum of Natural History Leiden “Naturalis”; University of Amsterdam, Zoological Museum; Zoolog- of the Carnivora accord with the island rule and, if so, for ische Staatssammlung, Mu¨nchen; Zoological Museum what reasons. Carnivore evolution on islands has been University of Copenhagen; Tel Aviv University, Zoological widely studied ever since Darwin. In his letter to the Lin- Museum; Raffles Museum of Biodiversity Research; Na- nean society in which he outlined his theory of evolution tional Science Museum, Tokyo; Primate Research Institute, by natural selection, Darwin (Darwin and Wallace 1858, Kyoto University; University of Alaska Fairbanks, Museum p. 49) wrote, “To give an imaginary example from changes of Natural History; Royal British Columbia Museum; in progress on an island: Let the organization of a canine Royal Ontario Museum; Canadian Museum of Nature; animal … become slightly plastic … those individuals with Field Museum; Carnegie Museum of Natural History; Na- the lightest forms … would be slightly favored … these tional Museum of Natural History–Smithsonian Institu- causes would … produce a marked effect, and adapt the tion; American Museum of Natural History; Museum of form of the fox or dog.” The Carnivora are particularly Comparative Zoology, Harvard University; National Mu- suitable for studies of body size evolution for several rea- seum of Ireland; Collections of the Department of Zoology, sons: they span a large range of body masses. Members University College, Cork, and the Ulster Museum. of the order are found on many islands throughout the Measurements were taken using digital calipers to 0.01- world (135 species on more than 500 different islands; S. mm precision or vernier calipers to 0.02-mm precision Meiri, unpublished data). They are extremely varied in (for measurements exceeding 300 mm). We chose three diet, from almost strict vegetarians through omnivores to measurements as measures of body size: skull length (con- piscivores, insectivores, and strict carnivores. Interspecific dylobasal length [CBL]; Von den Driesch 1976), the max- competition and predation, two selective pressures likely imum diameter of the upper canine, and the length of the to promote size change, are well documented for many lower first molar (M1, the lower carnassial tooth). The carnivores (Dayan et al. 1989, 1990, 1992; Thurber et al. CBL is a common measure of body size in biogeographic 1992; Dayan and Simberloff 1994; Palomares and Caro research (e.g., Rausch 1963; Ralls and Harvey 1985; Ellison 1999; Arjo et al. 2002). In addition, body size
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