Rapid prey‐induced shift in body size in an isolated snake population (Notechis scutatus,Elapidae)

Abstract Geographic divergence in phenotypic traits between long‐isolated populations likely has a genetic basis, but can phenotypic plasticity generate such divergence rapidly in the initial stages of isolation? Australian tiger snakes (Notechis scutatus, Elapidae) provide a classic model system for the evolution of body size: mean adult sizes are relatively invariant in mainland populations, but many offshore islands have dwarf or giant populations. Previous work has shown a genetic basis to this divergence in long‐isolated islands (>10 000 years), but what of the initial stages of this process? Human translocation of mainland snakes to Carnac Island 90 years ago gives us a unique opportunity to assess the proximate reasons for the giant size of Carnac Island animals compared with mainland conspecifics. Our data suggest a major role for phenotypic plasticity. Feeding trials on captive snakes from both island and mainland populations showed a strong link between food intake and growth rates, similar in the two populations. Snakes given abundant food grew much larger than we have ever recorded in the wild, demonstrating that observed mean body sizes are driven by food availability rather than genetic limits to growth. In combination with earlier work showing genetic divergence in growth rates in snakes from long‐isolated islands, our data suggest that geographical divergence in mean adult body sizes in this system initially is driven by a rapid shift due to phenotypic plasticity, with the divergence later canalized by a gradual accumulation of genetic differentiation.     

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.     

Island colonisation and the evolutionary rates of body size in insular neonate snakes

Island colonisation by animal populations is often associated with dramatic shifts in body size. However, little is known about the rates at which these evolutionary shifts occur, under what precise selective pressures and the putative role played by adaptive plasticity on driving such changes. Isolation time played a significant role in the evolution of body size in island Tiger snake populations, where adaptive phenotypic plasticity followed by genetic assimilation fine-tuned neonate body and head size (hence swallowing performance) to prey size. Here I show that in long isolated islands (>6000 years old) and mainland populations, neonate body mass and snout-vent length are tightly correlated with the average prey body mass available at each site. Regression line equations were used to calculate body size values to match prey size in four recently isolated populations of Tiger snakes. Rates of evolution in body mass and snout-vent length, calculated for seven island snake populations, were significantly correlated with isolation time. Finally, rates of evolution in body mass per generation were significantly correlated with levels of plasticity in head growth rates. This study shows that body size evolution occurs at a faster pace in recently isolated populations and suggests that the level of adaptive plasticity for swallowing abilities may correlate with rates of body mass evolution. I hypothesise that, in the early stages of colonisation, adaptive plasticity and directional selection may combine and generate accelerated evolution towards an ‘optimal'' phenotype.     

COLOR PATTERN POLYMORPHISM IN THE LAKE ERIE WATER SNAKE,NERODIA SIPEDON INSULARUM

Populations of the water snake, Nerodia sipedon, on islands in western Lake Erie are polymorphic for color pattern. These populations include banded, intermediate, and unbanded morphs while surrounding mainland populations consist solely of the banded morph. The hypothesis that this polymorphism is maintained by strong selection and migration pressures is widely accepted. Unbanded morphs are apparently more cryptic along island shorelines while banded morphs are more cryptic on the mainland. Migration of banded morphs from the mainland explains their persistence in island populations. Data collected in a capture-mark-recapture program on six islands provide no evidence of differential selection among morphs; morph frequencies do not differ among age classes, between once-captured and multiply-captured snakes, or between scarred and unscarred snakes. Furthermore, herring gulls, the most common snake predators in the island area, appear to detect banded and unbanded model snakes with equal ease. High site fidelity of water snakes and the distribution of morphs among islands suggest that migration from the mainland is not common. However, islands close to each other are similar in morph frequency, and water snakes have colonized islands elsewhere in the Great Lakes, indicating that some migration does occur. Recently, the frequency of banded morphs has increased in island populations while adult population sizes have declined. This increase in banded morphs is interpreted as reflecting an increased impact of migration from the mainland into these reduced populations. One scenario for the evolution and maintenance of this polymorphism is that selection was important in establishing unbanded morphs in island populations as they became isolated from the mainland. As populations declined to their present size, the impact of migration from the mainland increased and is now swamping the effect of selection. Further declines in island population size may result in fixation of the banded morph.     

Body-size evolution on islands: are adult size variations in tiger snakes a nonadaptive consequence of selection on birth size?

Mean adult size has been used as the traditional measure of body size to explain trends of insular gigantism and dwarfism in a wide array of taxa. However, patterns of variation in body size at birth have received surprisingly little attention, leaving open the possibility that adult body-size differences are nonadaptive consequences of selection acting on neonate body size. Here I used an empirical and correlative approach to test this hypothesis in a mosaic of 12 island and mainland snake populations in Australia. Data collected on 597 adult and 1,084 neonate tiger snakes showed that (1) both adult and neonate mean body sizes varied strongly across populations; (2) prey diversity and size convincingly explained birth-size variations: birth size-notably, gape size-correlated with prey size; (3) neonate snout-vent length was significantly correlated with neonate gape size; and (4) neonate snout-vent length was significantly correlated with adult snout-vent length. Postnatal growth rates recorded under common-garden conditions differed across populations and were correlated with mean prey size. These data collectively suggest that (1) prey size is the main driver for the evolution of body size at birth in gape-limited predators, (2) adult size variations may reflect selective forces acting on earlier life stages, and (3) adult size variations may also reflect resource availability during ontogeny (notably, prey diversity).     

Feeding preferences in 2 disjunct populations of tiger snakes, Notechis scutatus (Elapidae)

Variations at both the genetic and phenotypic levels play animportant role in responses to food and food-related stimuli.Knowledge of such variations is crucial to understanding howpopulations adapt to changing environments. We investigatedthe dietary preferences of 2 tiger snake populations and comparedthe responses of diet-naive animals (laboratory-born neonates),diet-controlled animals (laboratory-reared juveniles), and naturaldiet–experienced animals (wild-caught adults) to visualand chemical cues from 6 prey types (mouse, skink, silver gull,chicken, shearwater, and frog). The mainland population inhabitsa swamp, feeds mostly on frogs, and suffers heavy predation.The second population inhabits a small nearby offshore islandwith no standing water (no frogs); feeds mostly on skinks, mice,and, as adults, silver gull chicks; and suffers no known predation.Although different prey are eaten in the 2 populations, adultwild-caught snakes from both populations showed a significantpreference for 3 types of prey (frog, mouse, and chick), irrespectiveof their natural diet. Neonates responded to all prey cues morethan they did to control stimuli in both populations. However,the island neonates showed significantly higher interest insilver gull chick stimuli (the main prey of the island adultsnakes) than did their mainland conspecifics. Laboratory-bredjuveniles displayed behavioral plasticity by significantly increasingtheir response to mice after being fed baby mice for 7 months.We conclude that genetic-based differences in food-related cuesare important in tiger snakes but that they are also capableof behavioral plasticity. Island adult and neonate snakes exhibitedresponses to prey types no longer consumed naturally (frog),suggesting that behavioral characters may have been retainedfor long periods under relaxed selection. Island neonates showeda strong interest in a novel prey item (silver gull). This resultcomplements previous work describing how island snakes havedeveloped the ability to swallow larger prey than usual, aswell as seemingly developing a taste for them.     

Convergence in reduced body size,head size,and blood glucose in three island reptiles

Many oceanic islands harbor diverse species that differ markedly from their mainland relatives with respect to morphology, behavior, and physiology. A particularly common morphological change exhibited by a wide range of species on islands worldwide involves either a reduction in body size, termed island dwarfism, or an increase in body size, termed island gigantism. While numerous instances of dwarfism and gigantism have been well documented, documentation of other morphological changes on islands remains limited. Furthermore, we lack a basic understanding of the physiological mechanisms that underlie these changes, and whether they are convergent. A major hypothesis for the repeated evolution of dwarfism posits selection for smaller, more efficient body sizes in the context of low resource availability. Under this hypothesis, we would expect the physiological mechanisms known to be downregulated in model organisms exhibiting small body sizes due to dietary restriction or artificial selection would also be downregulated in wild species exhibiting dwarfism on islands. We measured body size, relative head size, and circulating blood glucose in three species of reptiles—two snakes and one lizard—in the California Channel Islands relative to mainland populations. Collating data from 6 years of study, we found that relative to mainland population the island populations had smaller body size (i.e., island dwarfism), smaller head sizes relative to body size, and lower levels of blood glucose, although with some variation by sex and year. These findings suggest that the island populations of these three species have independently evolved convergent physiological changes (lower glucose set point) corresponding to convergent changes in morphology that are consistent with a scenario of reduced resource availability and/or changes in prey size on the islands. This provides a powerful system to further investigate ecological, physiological, and genetic variables to elucidate the mechanisms underlying convergent changes in life history on islands.     

Empirical evidence for an optimal body size in snakes

Abstract The concept of optimal size has been invoked to explain patterns in body size of terrestrial mammals. However, the generality of this phenomenon has not been tested with similarly complete data from other taxonomic groups. In this study we describe three statistical patterns of body size in snakes, all of which indicate an optimal length of 1.0 m. First, a distribution of largest body lengths of 618 snake species had a single mode at 1.0 m. Second, we found a positive relationship between the size of the largest member of an island snake assemblage and island area and a negative relationship between the size of the smallest member of an island snake assemblage and island area. Best-fit lines through these data cross at a point corresponding to 1.0 m in body length, the presumed optimal size for a one-species island. Third, mainland snake species smaller than 1.0 m become larger on islands whereas those larger than 1.0 m become smaller on islands. The observation that all three analyses converge on a common body size is concordant with patterns observed in mammals and partial analyses of four other disparate animal clades. Because snakes differ so strikingly from mammals (ectotherms, gape-limited predators, elongate body shape) the concordant patterns of these two groups provide strong evidence for the evolution of an optimal body size within independent monophyletic groups. However, snakes differ from other taxonomic groups that have been studied in exhibiting a body size distribution that is not obviously skewed in either direction. We suggest that idiosyncratic features of the natural history of ectotherms allow relatively unconstrained distributions of body size whereas physiological limitations of endotherms constrain distributions of body size to a right skew.     

Insular shifts in body size of rice frogs in the Zhoushan Archipelago, China

1. Differences in body size between mainland and island populations have been reported for reptiles, birds and mammals. Despite widespread recognition of insular shifts in body size in these taxa, there have been no reports of such body size shifts in amphibians. 2. We provide the first evidence of an insular shift in body size for an amphibian species, the rice frog Rana limnocharis. We found significant increases in body size of rice frogs on most sampled islands in the Zhoushan archipelago when compared with neighbouring mainland China. 3. Large body size in rice frogs on islands was significantly related to increased population density, in both breeding and non-breeding seasons. Increases in rice frog density were significantly related to higher resource availability on islands. Increased resource availability on islands has led to higher carrying capacities, which has subsequently facilitated higher densities and individual growth rates, resulting in larger body size in rice frogs. We also suggest that large body size has evolved on islands, as larger individuals are competitively superior under conditions of harsh intraspecific competition at high densities. 4. Increases in body size in rice frogs were not related to several factors that have been implicated previously in insular shifts in body size in other taxa. We found no significant relationships between body size of rice frogs and prey size, number of larger or smaller frog species, island area or distance of islands from the mainland. 5. Our findings contribute to the formation of a broad, repeatable ecological generality for insular shifts in body size across a range of terrestrial vertebrate taxa, and provide support for recent theoretical work concerning the importance of resource availability for insular shifts in body size.     

Rejection of non-adaptive hypotheses for intraspecific variation in trophic morphology in gape-limited predators

Recent studies have interpreted intraspecific divergence in relative head sizes in snakes as evidence for adaptation of the trophic apparatus in gape-limited predators to local prey size. However, such variation might also arise from non-adaptive processes (such as allometry, correlated response, genetic drift, or non-adaptive phenotypic plasticity). We test predictions from these alternative hypotheses using data on the allometric relationship between head size and body size in two wide-ranging snake species: eight populations of adders ( Vipera berus ) and 30 populations of common gartersnakes ( Thamnophis sirtalis ). Our data enable strong rejection of the alternative (non-adaptive) hypotheses, because the relationship between head and body size differed significantly among populations, the geographic distance separating pairs of populations explained less than 1.5% of their divergence in allometric coefficients, and the within-population allometric coefficients were higher than the among-population coefficients in each species. In addition, the geographical variability of allometric coefficients in females did not parallel that in males, suggesting that allometric coefficients have evolved independently in the two sexes. Phenotypic plasticity also cannot explain the data, because laboratory studies show that the allometric relationship between head size and body size is relatively insensitive to differing growth rates. We conclude that the intraspecific head size divergence in these snakes is better explained by spatially heterogeneous selection to optimize prey handling ability, than by non-adaptive processes.     

Lake Erie water snakes revisited: Morph- and age-specific variation in relative crypsis

Summary Populations of water snakes (Nerodia sipedon insularum) on islands in western Lake Erie are variable in colour pattern, consisting of unbanded, intermediate, and banded morphs. In contrast, mainland populations (N. s. sipedon) consist solely of banded morphs. Previous investigators hypothesized that natural selection favoured unbanded morphs on exposed island shorelines and banded morphs in overgrown mainland habitats and that gene flow from mainland populations was responsible for the persistence of banded morphs on islands. To clarify the potential role of natural selection, I quantified relative crypsis among morphs and age classes of water snakes by comparing the size of patches making up their colour patterns with the size of patches in island and mainland backgrounds. This analysis reveals that if unbanded morphs are more cryptic than intermediate and banded morphs on islands, it is only in the young-of-the-year age class. For older snakes on islands and for all snakes on the mainland, unbanded morphs are consistently less cryptic than intermediate and banded morphs. Given these results, the net direction of selection in island populations should depend on the intensity of predation on different age classes of snakes. Overall, selection may favour unbanded morphs (e.g. if predation occurs primarily on young-of-the-year), intermediate and banded morphs (e.g. if predation occurs primarily on older snakes), or be weak or absent (e.g. certain combinations of predation on young-of-the-year and older snakes). Using estimates of relative crypsis to guide reanalysis of morph frequency data, I find support for the hypothesis that unbanded morphs are favoured by natural selection in island populations.     

Sexual divergence in diets and morphology in Fijian sea snakes Laticauda colubrina (Laticaudinae)

In the Fiji Islands, female yellow‐lipped sea kraits (Laticauda colubrina) grow much larger than males, and have longer and wider heads than do conspecific males of the same body length. This morphological divergence is accompanied by (and may be adaptive to) a marked sex divergence in dietary habits. Adult female sea kraits feed primarily on large conger eels, and take only a single prey item per foraging bout. In contrast, adult males feed upon smaller moray eels, and frequently take multiple prey items. Prey size increases with snake body size in both males and females, but the sexes follow different trajectories in this respect. Female sea kraits consume larger eels relative to predator head size and body length than do males. Thus, the larger relative head size of female sea kraits is interpreted as an adaptation to consuming larger prey items. Our results are similar to those of previous studies on American water snakes (natricines) and Australian file snakes (acrochordids), indicating that similar patterns of sex divergence in dietary habits and feeding structures have evolved convergently in at least three separate lineages of aquatic snakes.     

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.     

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.     

‘Heaven’ for serpents? A mark–recapture study of tiger snakes (Notechis scutatus) on Carnac Island,Western Australia

Abstract Animals resident on small islands provide excellent opportunities to carry out detailed mark–recapture studies. Populations are closed and ecosystems are often simpler than those of mainland sites. These factors enable the study of cryptic species that have otherwise been neglected. Snakes are notable for their secretive nature and, as a result, few natural populations have been accurately described through long‐term mark–recapture monitoring. A population of tiger snakes (Notechis scutatus) was studied on Carnac Island, a small limestone island (16 ha) off the coast of Western Australia. Population estimates show that snake density is very high, with more than 20 adult snakes per ha. This equates to a biomass of more than 100 kg of a top predator concentrated in a very small area. Such a high predator density can be explained because adult snakes feed mainly on chicks of nesting birds that breed in large colonies on Carnac but forage elsewhere. Substantial annual growth rates in body size in most individuals suggest that food availability is high on Carnac. Growth rates decreased more sharply in adult females than in males, whereas annual changes in body mass were similar in both sexes, probably because of the high energetic costs of reproduction experienced by females. Surprisingly, the sex ratio was highly biased, with males largely outnumbering females.     

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.     

Sex-specific niche partitioning and sexual size dimorphism in Australian pythons (Morelia spilota imbricata)

Sexual dimorphism is usually interpreted in terms of reproductive adaptations, but the degree of sex divergence also may be affected by sex-based niche partitioning. In gape-limited animals like snakes, the degree of sexual dimorphism in body size (SSD) or relative head size can determine the size spectrum of ingestible prey for each sex. Our studies of one mainland and four insular Western Australian populations of carpet pythons ( Morelia spilota ) reveal remarkable geographical variation in SSD, associated with differences in prey resources available to the snakes. In all five populations, females grew larger than males and had larger heads relative to body length. However, the populations differed in mean body sizes and relative head sizes, as well as in the degree of sexual dimorphism in these traits. Adult males and females also diverged strongly in dietary composition: males consumed small prey (lizards, mice and small birds), while females took larger mammals such as possums and wallabies. Geographic differences in the availability of large mammalian prey were linked to differences in mean adult body sizes of females (the larger sex) and thus contributed to sex-based resource partitioning. For example, in one population adult male snakes ate mice and adult females ate wallabies; in another, birds and lizards were important prey types for both sexes. Thus, the high degree of geographical variation among python populations in sexually dimorphic aspects of body size and shape plausibly results from geographical variation in prey availability.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 113–125.     

Morphological divergence in giant fossil dormice

Insular gigantism—evolutionary increases in body size from small-bodied mainland ancestors—is a conceptually significant, but poorly studied, evolutionary phenomenon. Gigantism is widespread on Mediterranean islands, particularly among fossil and extant dormice. These include an extant giant population of Eliomys quercinus on Formentera, the giant Balearic genus †Hypnomys and the exceptionally large †Leithia melitensis of Pleistocene Sicily. We quantified patterns of cranial and mandibular shape and their relationships to head size (allometry) among mainland and insular dormouse populations, asking to what extent the morphology of island giants is explained by allometry. We find that gigantism in dormice is not simply an extrapolation of the allometric trajectory of their mainland relatives. Instead, a large portion of their distinctive cranial and mandibular morphology resulted from the population- or species-specific evolutionary shape changes. Our findings suggest that body size increases in insular giant dormice were accompanied by the evolutionary divergence of feeding adaptations. This complements other evidence of ecological divergence in these taxa, which span predominantly faunivorous to herbivorous diets. Our findings suggest that insular gigantism involves context-dependent phenotypic modifications, underscoring the highly distinctive nature of island faunas.     

Island tameness: living on islands reduces flight initiation distance

One of Darwin''s most widely known conjectures is that prey are tame on remote islands, where mammalian predators are absent. Many species appear to permit close approach on such islands, but no comparative studies have demonstrated reduced wariness quantified as flight initiation distance (FID; i.e. predator–prey distance when the prey begins to flee) in comparison with mainland relatives. We used the phylogenetic comparative method to assess influence of distance from the mainland and island area on FID of 66 lizard species. Because body size and predator approach speed affect predation risk, we included these as independent variables. Multiple regression showed that FID decreases as distance from mainland increases and is shorter in island than mainland populations. Although FID increased as area increased in some models, collinearity made it difficult to separate effects of area from distance and island occupancy. FID increases as SVL increases and approach speed increases; these effects are statistically independent of effects of distance to mainland and island occupancy. Ordinary least-squares models fit the data better than phylogenetic regressions, indicating little or no phylogenetic signal in residual FID after accounting for the independent variables. Our results demonstrate that island tameness is a real phenomenon in lizards.