Slaying dragons: limited evidence for unusual body size evolution on islands

Aim Island taxa often attain forms outside the range achieved by mainland relatives. Body size evolution of vertebrates on islands has therefore received much attention, with two seemingly conflicting patterns thought to prevail: (1) islands harbour animals of extreme size, and (2) islands promote evolution towards medium body size (‘the island rule’). We test both hypotheses using body size distributions of mammal, lizard and bird species. Location World‐wide. Methods We assembled body size and insularity datasets for the world’s lizards, birds and mammals. We compared the frequencies with which the largest or smallest member of a group is insular with the frequencies expected if insularity is randomly assigned within groups. We tested whether size extremes on islands considered across mammalian phylogeny depart from a null expectation under a Brownian motion model. We tested the island rule by comparing insular and mainland members of (1) a taxonomic level and (2) mammalian sister species, to determine if large insular animals tend to evolve smaller body sizes while small ones evolve larger sizes. Results The smallest species in a taxon (order, family or genus) are insular no more often than would be expected by chance in all groups. The largest species within lizard families and bird genera (but no other taxonomic levels) are insular more often than expected. The incidence of extreme sizes in insular mammals never departs from the null, except among extant genera, where gigantism is marginally less common than expected under a Brownian motion null. Mammals follow the island rule at the genus level and when comparing sister species and clades. This appears to be driven mainly by insular dwarfing in large‐bodied lineages. A similar pattern in birds is apparent for species within orders. However, lizards follow the converse pattern. Main conclusions The popular misconception that islands have more than their fair share of size extremes may stem from a greater tendency to notice gigantism and dwarfism when they occur on islands. There is compelling evidence for insular dwarfing in large mammals, but not in other taxa, and little evidence for the second component of the island rule – gigantism in small‐bodied taxa.     

The effect of island area on body size in a primate species from the Sunda Shelf Islands

Aim  We examine the effect of island area on body dimensions in a single species of primate endemic to Southeast Asia, the long-tailed macaque ( Macaca fascicularis ). In addition, we test Allen's rule and a within-species or intraspecific equivalent of Bergmann's rule (i.e. Rensch's rule) to evaluate body size and shape evolution in this sample of insular macaques.
Location  The Sunda Shelf islands of Southeast Asia.
Methods  Body size measurements of insular macaques gathered from the literature were analysed relative to island area, latitude, maximum altitude, isolation from the mainland and other islands, and various climatic variables using linear regression.
Results  We found no statistically significant relationship between island area and body length or head length in our sample of insular long-tailed macaques. Tail length correlated negatively with island area. Head length and body length exhibited increases corresponding to increasing latitude, a finding seemingly consistent with the expression of Bergmann's rule within a single species. These variables, however, were not correlated with temperature, indicating that Bergmann's rule is not in effect. Tail length was not correlated with either temperature or increasing latitude, contrary to that predicted by Allen's rule.
Main conclusions  The island rule dictating that body size will covary with island area does not apply to this particular species of primate. Our study is consistent with results presented in the literature by demonstrating that skull and body length in insular long-tailed macaques do not, strictly speaking, conform to Rensch's rule. Unlike previous studies, however, our findings suggest that tail-length variation in insular macaques does not support Allen's rule.     

Life-history traits associated with fragmentation vulnerability of lizards in the Thousand Island Lake, China

Following habitat fragmentation, the remnant faunal community will undergo a period of species loss or 'relaxation.' Theory predicts that species with particular life-history traits, such as a small population size, small geographical range, low fecundity and large body size, should be more vulnerable to fragmentation. In this study, we investigated the relationships between the above life-history traits and the fragmentation vulnerability index (the number of islands occupied) of five lizard species inhabiting recently isolated land-bridge islands in the Thousand Island Lake, China. Data on life-history traits were collected from field surveys (population density) and from the literature (body size, clutch size and geographical range size). The species–area relationships for lizards sampled from the mainland versus on the islands differed significantly (i.e. the number of species inhabiting islands was decreased relative to similar-sized areas on the mainland), indicating that species extinction has occurred on all of the study islands following isolation. For the fragmentation vulnerability index, model selection based on Akaike's information criterion identified natural density at mainland sites as the best correlate of vulnerability to fragmentation, supporting the hypothesis that rare species are most vulnerable to local extinction and will be lost first from fragmented landscapes. In contrast, there was little evidence for an effect of lizards' snout–vent length, clutch size or geographical range size on fragmentation vulnerability. Identification of species traits that render some species more vulnerable to fragmentation than others has important implications for conservation and can be used to aid direct management efforts.     

Body size of insular carnivores: little support for the island rule

Large mammals are thought to evolve to be smaller on islands, whereas small mammals grow larger. A negative correlation between relative size of island individuals and body mass is termed the "island rule." Several mechanisms--mainly competitive release, resource limitation, dispersal ability, and lighter predation pressure on islands, as well as a general physiological advantage of modal size--have been advanced to explain this pattern. We measured skulls and teeth of terrestrial members of the order Carnivora in order to analyze patterns of body size evolution between insular populations and their near mainland conspecifics. No correlations were found between the size ratios of insular/mainland carnivore species and body mass. Only little support for the island rule is found when individual populations rather than species are considered. Our data are at odds with those advanced in support of theories of optimal body size. Carnivore size is subjected to a host of selective pressures that do not vary uniformly from place to place. Mass alone cannot account for the patterns in body size of insular carnivores.     

Area, isolation and body size evolution in insular carnivores

Body sizes of insular mammals often differ strikingly from those of their mainland conspecifics. Small islands have reduced numbers of competitor and predator species, and more limited resources. Such reductions are believed to select for predictable changes in body sizes, with large mammals growing progressively smaller as island area decreases, while small ones grow progressively larger. Medium-sized mammals are thought to be largest on intermediate-sized islands. Increased isolation is seen as promoting insular gigantism. We searched for such patterns using a large database of insular carnivore specimens. Neither small nor large carnivores show a consistent area/body size relationship. Medium-sized carnivores are no more likely to attain large size on medium-sized islands then they are to be small there. We found no consistent patterns of body size variation in relation to isolation.     

Does relaxed predation drive phenotypic divergence among insular populations?

The evolution of striking phenotypes on islands is a well‐known phenomenon, and there has been a long‐standing debate on the patterns of body size evolution on islands. The ecological causes driving divergence in insular populations are, however, poorly understood. Reduced predator fauna is expected to lower escape propensity, increase body size and relax selection for crypsis in small‐bodied, insular prey species. Here, we investigated whether escape behaviour, body size and dorsal coloration have diverged as predicted under predation release in spatially replicated islet and mainland populations of the lizard species Podarcis gaigeae. We show that islet lizards escape approaching observers at shorter distances and are larger than mainland lizards. Additionally, we found evidence for larger between‐population variation in body size among the islet populations than mainland populations. Moreover, islet populations are significantly more divergent in dorsal coloration and match their respective habitats poorer than mainland lizards. These results strongly suggest that predation release on islets has driven population divergence in phenotypic and behavioural traits and that selective release has affected both trait means and variances. Relaxed predation pressure is therefore likely to be one of the major ecological factors driving body size divergence on these islands.     

Evolution and ecology of lizard body sizes

Aim Body size is instrumental in influencing animal physiology, morphology, ecology and evolution, as well as extinction risk. I examine several hypotheses regarding the influence of body size on lizard evolution and extinction risk, assessing whether body size influences, or is influenced by, species richness, herbivory, island dwelling and extinction risk. Location World‐wide. Methods I used literature data and measurements of museum and live specimens to estimate lizard body size distributions. Results I obtained body size data for 99% of the world's lizard species. The body size–frequency distribution is highly modal and right skewed and similar distributions characterize most lizard families and lizard assemblages across biogeographical realms. There is a strong negative correlation between mean body size within families and species richness. Herbivorous lizards are larger than omnivorous and carnivorous ones, and aquatic lizards are larger than non‐aquatic species. Diurnal activity is associated with small body size. Insular lizards tend towards both extremes of the size spectrum. Extinction risk increases with body size of species for which risk has been assessed. Main conclusions Small size seems to promote fast diversification of disparate body plans. The absence of mammalian predators allows insular lizards to attain larger body sizes by means of release from predation and allows them to evolve into the top predator niche. Island living also promotes a high frequency of herbivory, which is also associated with large size. Aquatic and nocturnal lizards probably evolve large size because of thermal constraints. The association between large size and high extinction risk, however, probably reflects a bias in the species in which risk has been studied.     

Rule reversal: Ecogeographical patterns of body size variation in the common treeshrew (Mammalia,Scandentia)

There are a number of ecogeographical “rules” that describe patterns of geographical variation among organisms. The island rule predicts that populations of larger mammals on islands evolve smaller mean body size than their mainland counterparts, whereas smaller‐bodied mammals evolve larger size. Bergmann's rule predicts that populations of a species in colder climates (generally at higher latitudes) have larger mean body sizes than conspecifics in warmer climates (at lower latitudes). These two rules are rarely tested together and neither has been rigorously tested in treeshrews, a clade of small‐bodied mammals in their own order (Scandentia) broadly distributed in mainland Southeast Asia and on islands throughout much of the Sunda Shelf. The common treeshrew, Tupaia glis, is an excellent candidate for study and was used to test these two rules simultaneously for the first time in treeshrews. This species is distributed on the Malay Peninsula and several offshore islands east, west, and south of the mainland. Using craniodental dimensions as a proxy for body size, we investigated how island size, distance from the mainland, and maximum sea depth between the mainland and the islands relate to body size of 13 insular T. glis populations while also controlling for latitude and correlation among variables. We found a strong negative effect of latitude on body size in the common treeshrew, indicating the inverse of Bergmann's rule. We did not detect any overall difference in body size between the island and mainland populations. However, there was an effect of island area and maximum sea depth on body size among island populations. Although there is a strong latitudinal effect on body size, neither Bergmann's rule nor the island rule applies to the common treeshrew. The results of our analyses demonstrate the necessity of assessing multiple variables simultaneously in studies of ecogeographical rules.     

Frugivory in polychrotid lizards: effects of body size

As more data have become available on lizard diets in the past few decades, researchers have stressed the importance of lizards as pollinators and seed dispersers. Whereas large body size has been traditionally put forward as a major biological factor allowing herbivory and frugivory in lizards, a recent review of frugivory and seed dispersal by lizards showed that frugivory might be considered to be a typical island phenomenon, independent of body size. Here we show that frugivory is correlated with lizard body size among a group of syntopic Anolis species in Jamaica, with larger species eating more fruit. Additionally, the size of the fruits consumed is significantly related to lizard body size. Multiple regression analyses show that this is largely a pure body size effect as head shape or residual bite force are uncorrelated to overall fruit size. Moreover, we demonstrate that among polychrotid (Anolis-like) lizards in general, those that consume fruit are on average larger than those that do not. Lizards from the mainland were not significantly different in body size from island species. We thus suggest that fruit consumption in polychrotid lizards is mediated by large body size whether living on islands or not.     

Body size evolution in insular vertebrates: generality of the island rule

Aim My goals here are to (1) assess the generality of the island rule – the graded trend from gigantism in small species to dwarfism in larger species – for mammals and other terrestrial vertebrates on islands and island‐like ecosystems; (2) explore some related patterns of body size variation in insular vertebrates, in particular variation in body size as a function of island area and isolation; (3) offer causal explanations for these patterns; and (4) identify promising areas for future studies on body size evolution in insular vertebrates. Location Oceanic and near‐shore archipelagos, and island‐like ecosystems world‐wide. Methods Body size measurements of insular vertebrates (non‐volant mammals, bats, birds, snakes and turtles) were obtained from the literature, and then regression analyses were conducted to test whether body size of insular populations varies as a function of body size of the species on the mainland (the island rule) and with characteristics of the islands (i.e. island isolation and area). Results The island rule appears to be a general phenomenon both with mammalian orders (and to some degree within families and particular subfamilies) as well as across the species groups studied, including non‐volant mammals, bats, passerine birds, snakes and turtles. In addition, body size of numerous species in these classes of vertebrates varies significantly with island isolation and island area. Main conclusions The patterns observed here – the island rule and the tendency for body size among populations of particular species to vary with characteristics of the islands – are actually distinct and scale‐dependent phenomena. Patterns within archipelagos reflect the influence of island isolation and area on selective pressures (immigration filters, resource limitation, and intra‐ and interspecific interactions) within particular species. These patterns contribute to variation about the general trend referred to as the island rule, not the signal for that more general, large‐scale pattern. The island rule itself is an emergent pattern resulting from a combination of selective forces whose importance and influence on insular populations vary in a predictable manner along a gradient from relatively small to large species. As a result, body size of insular species tends to converge on a size that is optimal, or fundamental, for a particular bau plan and ecological strategy.     

Factors explaining increased body size in common shrews (Sorex araneus) on Scottish islands

Aim  Island populations of small mammals are often characterized by a larger body size compared with neighbouring mainland or continental populations of the same species. A number of reasons have been put forward to explain this phenomenon. The aim of this study was to test which of these hypotheses can best explain the increase of body size in common shrews ( Sorex araneus ) on islands.
Location  The fieldwork for this study was carried out on the islands of the Inner Hebrides, Clyde Islands and the west coast of Scotland.
Methods  This study compared body sizes of common shrews from mainland and island sites on the west coast of Scotland, based on measurements of hind foot lengths. On 10 of the 13 islands sampled, common shrews were significantly larger than on the mainland. Body size did not vary significantly among mainland populations. We used the directional contrasts method to test the relative contributions of possible factors explaining the large body size observed in the island populations.
Results  We found that body size of common shrews on islands was positively related to distance from mainland, negatively related to average annual temperature, negatively related to island size, and may also be influenced by the presence or absence of pygmy shrews ( Sorex minutus ) on the island.
Main conclusions  Our results suggest a role for founder events, Bergmann's rule and K -selection in determining body size of common shrews on islands.     

Morphological shifts in island-dwelling birds: the roles of generalist foraging and niche expansion

Abstract Passerine birds living on islands are usually larger than their mainland counterparts, in terms of both body size and bill size. One explanation for this island rule is that shifts in morphology are an adaptation to facilitate ecological niche expansion. In insular passerines, for instance, increased bill size may facilitate generalist foraging because it allows access to a broader range of feeding niches. Here we use morphologically and ecologically divergent races of white-eyes (Zosteropidae) to test three predictions of this explanation: (1) island populations show a wider feeding niche than mainland populations; (2) island-dwelling populations are made up of individual generalists; and (3) within insular populations there is a positive association between size and degree of foraging generalism. Our results provide only partial support for the traditional explanation. In agreement with the core prediction, island populations of white-eye do consistently display a wider feeding niche than comparative mainland populations. However, observations of individually marked birds reveal that island-dwelling individuals are actually more specialized than expected by chance. Additionally, neither large body size nor large bill size are associated with generalist foraging behavior per se. These latter results remained consistent whether we base our tests on natural foraging behavior or on observations at an experimental tree, and whether we use data from single or multiple cohorts. Taken together, our results suggest that generalist foraging and niche expansion are not the full explanation for morphological shifts in island-dwelling white-eyes. Hence, we review briefly five alternative explanations for morphological divergence in insular populations: environmental determination of morphology, reduced predation pressure, physiological optimization, limited dispersal, and intraspecific dominance.     

Island/mainland body size differences in Australian varanid lizards

Island varanids seem to be an exception to the rule that territorial vertebrate taxa often become gigantic relative to mainland relatives when on islands, whereas non-territorial species become dwarfed (Case 1978). However, no systematic island/mainland studies have examined the empirical size trends in this group of carnivorous lizards. We perform such an analysis for the Australian region and critically evaluate various selective agents that might be responsible for size changes in several island populations. Insular gigantism occurs at least four times among the island populations examined. The magnitude of size change is positively correlated to prey abundance on the islands (as indirectly measured through a condition index of the lizards, essentially a measure of how fat they arc) and the size of prey: islands with large prey have large varanids and vice versa. Since the island population with the largest size change, the Reevesby Varanus rosenbergi, was introduced less than 100 years ago, these size changes can be quite rapid. This might indicate that selective coefficients are strong; however, we can not exclude the possibility that these size differences have no genetic component and simply reflect environmental differences in growth rate and shifts in age structure between island and mainland locations.     

Biogeography of body size in Pacific island birds

Many insular vertebrates have undergone rapid and dramatic changes in body size compared to their mainland counterparts. Here we explore the relationship between two well known patterns of island body size – the tendency for large‐bodied species to dwarf and small‐bodied species to get larger on islands, known as the “island rule”, and the scaling of maximum and minimum body size of island assemblages with island area. Drawing on both fossil and modern data, we examined the relationship between body size and island area in Pacific island birds, both within clades and at the island assemblage level. We found that the size of the smallest bird on each island decreased with island area while the maximum body size increased with island area. Similarly, within clades the body size of small‐bodied groups decreased and large‐bodied groups increased from small to large islands, consistent with the island rule. However, the magnitude of size change within clades was not sufficient to explain the overall scaling of maximum size with island area. Instead, the pattern was driven primarily by the evolution of very large, flightless birds on large islands. Human‐mediated extinctions on islands over the past few millennia severely impacted large, flightless birds, to the effect that this macroecological pattern has been virtually erased. After controlling for effects of biogeographic region and island area, we found island productivity to be the best predictor of maximum size in flightless birds. This result, and the striking similarities in maximum body size between flightless birds and island mammals, suggests a common energetic mechanism linking body size and landmass area in both the island rule and the scaling of island body size extremes.     

Maximum body size among insular Komodo dragon populations covaries with large prey density

This study documents variation in maximum body size of Komodo dragons ( Varanus komodoensis ) among the four extant island populations in Komodo National Park and compares an indirect measure of deer density, the major prey item for large dragons, to differences in maximum body size among islands. The largest 15% of dragons from the large islands of Komodo and Rinca were significantly longer and heavier than the largest 15% of dragons on the small islands of Gili Motang and Nusa Kode. There was a 33% difference in snout vent length (SVL) between dragons found on Komodo and those found on Gili Motang, with mass varying by more than four-fold. Density of deer pellet groups between islands ranged from 5.86±0.75 groups per transect on Gili Motang to 20.73±1.02 groups per transect on Komodo Island. Maximal dragon SVL and mass was highly positively correlated with this index of deer density. Low prey density on the two small islands could constrain body size via energetic constraints. At present we can not deduce if insular body size variation has arisen through genotypic or phenotypic mechanisms.     

The generality of the island rule reexamined

Aim  M.V. Lomolino and colleagues have recently reviewed the island rule in mammals and other vertebrates, claiming it is a general pattern. They have portrayed our recent analysis as weakly supporting the island rule, seeing weakness in our use of what they considered to be inadequate size indices (skulls and teeth, rather than mass or body length) and in our use of large islands. They argue that size evolution on islands points to a bauplan-specific fundamental size. We aim to test the generality of the rule and the adequacy of some of the data used to support it.
Location  Insular environments world-wide.
Methods  We collate and analyse data on skull sizes of carnivores and body masses of mammals in general to see whether there is a graded trend from dwarfism in large species to gigantism in smaller ones.
Results  The island rule is not supported with either the carnivore or the mammal data sets. Island area does not influence size change.
Main conclusions  Our results suggest that data recently advanced in support of the island rule are inadequate and that the island rule is not a general pattern for all mammals.     

Of mice and mammoths: evaluations of causal explanations for body size evolution in insular mammals

Aim We investigated the hypothesis that the insular body size of mammals results from selective forces whose influence varies with characteristics of the focal islands and the focal species, and with interactions among species (ecological displacement and release). Location Islands world‐wide. Methods We assembled data on the geographic characteristics (area, isolation, maximum elevation, latitude) and climate (annual averages and seasonality of temperature and precipitation) of islands, and on the ecological and morphological characteristics of focal species (number of mammalian competitors and predators, diet, body size of mainland reference populations) that were most relevant to our hypothesis (385 insular populations from 98 species of extant, non‐volant mammals across 248 islands). We used regression tree analyses to examine the hypothesized contextual importance of these factors in explaining variation in the insular body size of mammals. Results The results of regression tree analyses were consistent with predictions based on hypotheses of ecological release (more pronounced changes in body size on islands lacking mammalian competitors or predators), immigrant selection (more pronounced gigantism in small species inhabiting more isolated islands), thermoregulation and endurance during periods of climatic or environmental stress (more pronounced gigantism of small mammals on islands of higher latitudes or on those with colder and more seasonal climates), and resource subsidies (larger body size for mammals that utilize aquatic prey). The results, however, were not consistent with a prediction based on resource limitation and island area; that is, the insular body size of large mammals was not positively correlated with island area. Main conclusions These results support the hypothesis that the body size evolution of insular mammals is influenced by a combination of selective forces whose relative importance and nature of influence are contextual. While there may exist a theoretical optimal body size for mammals in general, the optimum for a particular insular population varies in a predictable manner with characteristics of the islands and the species, and with interactions among species. This study did, however, produce some unanticipated results that merit further study – patterns associated with Bergmann’s rule are amplified on islands, and the body size of small mammals appears to peak at intermediate and not maximum values of latitude and island isolation.     

One size does not fit all: no evidence for an optimal body size on islands

Aim Optimal body size theories predict that large clades have a single, optimal, body size that serves as an evolutionary attractor, with the full body size spectrum of a clade resulting from interspecific competition. Because interspecific competition is believed to be reduced on islands, such theories predict that insular animals should be closer to the optimal size than mainland animals. We test the resulting prediction that insular clade members should therefore have narrower body size ranges than their mainland relatives. Location World‐wide. Methods We used body sizes and a phylogenetic tree of 4004 mammal species, including more than 200 species that went extinct since the last ice age. We tested, in a phylogenetically explicit framework, whether insular taxa converge on an optimal size and whether insular clades have narrow size ranges. Results We found no support for any of the predictions of the optimal size theory. No specific size serves as an evolutionary attractor. We did find consistent evidence that large (> 10 kg) mammals grow smaller on islands. Smaller species, however, show no consistent tendency to either dwarf or grow larger on islands. Size ranges of insular taxa are not narrower than expected by chance given the number of species in their clades, nor are they narrower than the size ranges of their mainland sister clades – despite insular clade members showing strong phylogenetic clustering. Main conclusions The concept of a single optimal body size is not supported by the data that were thought most likely to show it. We reject the notion that inclusive clades evolve towards a body‐plan‐specific optimum.     

The ‘Island Rule’ Acting on Anuran Populations (Bufonidae: Rhinella ornata) of the Southern Hemisphere

Darwin and Wallace, in the mid‐nineteenth century, were the first to document examples of natural selection acting on island dwellers. A century later a pattern of morphological differences among organisms on islands was coined the ‘island rule’, which states that on islands species with small individuals tend toward gigantism and large individuals tend toward dwarfism. Selective pressures such as limited resources and increased intraspecific competition modulate the size of organisms in these environments. Of the several works that have tested vertebrates for adherence to the island rule only two have addressed amphibians. This work is the third record of body size variation of island amphibian populations, and the first for the Southern Hemisphere. The islands investigated were once continuous with mainland, and now are isolated as a result of sea level fluctuations that took place in the Pleistocene and Holocene. This study compared morphometric variation in populations of Rhinella ornata (Bufonidae) occurring on three islands of the Costa Verde to populations on five continental areas in Rio de Janeiro, Brazil. We measured 18 morphometric variables of 177 individuals. There was a shift toward smaller body size (dwarfism) in two of the three island populations studied. We attribute this general pattern to geographic factors, verifying the expression of the island rule in tropical frogs populations (insular dwarfism) operating inversely in relation to those of temperate environments (island gigantism).     

Body Size Evolution in Insular Speckled Rattlesnakes (Viperidae: Crotalus mitchellii)

Background

Speckled rattlesnakes (Crotalus mitchellii) inhabit multiple islands off the coast of Baja California, Mexico. Two of the 14 known insular populations have been recognized as subspecies based primarily on body size divergence from putative mainland ancestral populations; however, a survey of body size variation from other islands occupied by these snakes has not been previously reported. We examined body size variation between island and mainland speckled rattlesnakes, and the relationship between body size and various island physical variables among 12 island populations. We also examined relative head size among giant, dwarfed, and mainland speckled rattlesnakes to determine whether allometric differences conformed to predictions of gape size (and indirectly body size) evolving in response to shifts in prey size.

Methodology/Principal Findings

Insular speckled rattlesnakes show considerable variation in body size when compared to mainland source subspecies. In addition to previously known instances of gigantism on Ángel de la Guarda and dwarfism on El Muerto, various degrees of body size decrease have occurred frequently in this taxon, with dwarfed rattlesnakes occurring mostly on small, recently isolated, land-bridge islands. Regression models using the Akaike information criterion (AIC) showed that mean SVL of insular populations was most strongly correlated with island area, suggesting the influence of selection for different body size optima for islands of different size. Allometric differences in head size of giant and dwarf rattlesnakes revealed patterns consistent with shifts to larger and smaller prey, respectively.

Conclusions/Significance

Our data provide the first example of a clear relationship between body size and island area in a squamate reptile species; among vertebrates this pattern has been previously documented in few insular mammals. This finding suggests that selection for body size is influenced by changes in community dynamics that are related to graded differences in area over what are otherwise similar bioclimatic conditions. We hypothesize that in this system shifts to larger prey, episodic saturation and depression of primary prey density, and predator release may have led to insular gigantism, and that shifts to smaller prey and increased reproductive efficiency in the presence of intense intraspecific competition may have led to insular dwarfism.