Head shape evolution in Gymnophthalmidae: does habitat use constrain the evolution of cranial design in fossorial lizards?

Habitat usage comprises interactions between ecological parameters and organismal capacities, and the selective pressures that ultimately determine the outcome of such processes in an evolutionary scale may be conflicting when the same morphological structure is recruited for different activities. Here, we investigate the roles of diet and locomotion in the evolution of cranial design in gymnophthalmid lizards and test the hypothesis that microhabitat use drives head shape evolution, particularly in head-first burrowers. Morphological factors were analysed in relation to continuous ecological indexes (prey hardness and substrate compactness) using conventional and phylogenetic approaches. Results suggest that the evolution of head morphology in Gymnophthalmidae was shaped under the influence of microhabitat use rather than diet: burrowers have shorter heads with lower rostral angulation, independently of the prey consumed. Food preferences appear to be relatively conserved throughout the phylogeny of the group, which may have permitted the extensive radiation of gymnophthalmids into fossorial microhabitats.     

Lizard Assemblages from Natural Cerrado Enclaves in Southwestern Amazonia: The Role of Stochastic Extinctions and Isolation1

We used null model analyses to investigate the existence of structure in lizard assemblages from open vegetation enclaves in Rondónia, southwestern Amazonia, in relation to species richness, species co‐occurrence, diet, and size overlap. These enclaves presumably have been isolated since the Holocene, providing a history of long‐term isolation. We assumed that the presence of structure in lizard assemblages from the Rondónia enclaves is consistent with the notion that extinctions are a deterministic process, some species being more prone to extinction than others. We grouped enclaves into four categories: latosoil cerrado, sandy cerrado, transitional forest, and rocky field. We collected 14 Cerrado lizard species, consisting of five families in all sampled areas. Analyses of species richness, co‐occurrence, diet overlap, and size overlap patterns suggested lack of organization in the assemblages. The assemblages from the rocky fields of Guajarí–Mirim and the sandy cerrados in Vilhena were significantly structured in diet overlap, whereas the remaining assemblages lacked structure. This probably resulted from phylogenetic inertia and not from ecological interactions. Our results suggest that extinctions proceeded in a stochastic fashion and that historical factors had a dominant role in shaping lizard assemblages in detriment of present‐day ecological factors. In addition, we identified endemic species in the enclaves as well as a tight association between unique ecogeographic features of the landscape and species occurrences. We propose that conservation measures in the region must adequately preserve these features to ensure the survival of the species.     

Ontogenetic variation in the autecology of the greater earless lizard Cophosaurus texanus

We investigated the potential for ontogenetic resource partitioning within a population of the Texas earless lizard Cophosaurus texanus Results from focal samples, line transects, haphazard observations, and stomach flushes compiled in summer (July 1993) and autumn (September 1995) revealed differences in microhabitat use and diet between adults and juveniles Juveniles use rock perches more frequently than adult males, and consume smaller prey than adults No ontogenetic differences in thermal ecology were observed Diet differences between juveniles and adults may be attributed to the inability of juveniles to handle large prey items Differences between juveniles and adults in prey size and microhabitat use suggest ontogenetic variation in foraging, predator avoidance, and territory defense     

Niche differences between two sympatric whiptail lizards (Cnemidophorus abaetensis and C. ocellifer, Teiidae) in the restinga habitat of northeastern Brazil.

Differences among sympatric lizard species usually result from differences in the use of three resources: space, time and food or some combination of these three. However, differences in resource utilization among sympatric species may simply reflect their specific ecological needs rather than competitive pressures. In this study, we analyzed the temporal, spatial and food niche of two congeneric teiids (Cnemidophorus abaetensis and C. ocellifer) living sympatrically in the "restinga" habitat of Abaeté in the Salvador Municipality, Bahia State, Brazil to assess the degree of niche differentiation among them. The whiptail species overlapped considerably in an hourly activity (Ojk = 0.93), in microhabitat use (Ojk = 0.97) and in the prey items consumed (Ojk = 0.89). Differences in amount of vegetation in the microhabitats used by both lizard species may have contributed to differences in the activity period and in the distribution of the main prey eaten by these lizards which may, in turn, facilitate their coexistence in Abaeté. Although sympatric C. ocellifer and C. abaetensis in Abaeté differed only slightly in their use of microhabitats, period of activity and diet, the most important niche dimension segregating the two species seemed to be the food niche.     

Species interactions mediate phylogenetic community structure in a hyperdiverse lizard assemblage from arid Australia

Evolutionary history can exert a profound influence on ecological communities, but few generalities have emerged concerning the relationships among phylogeny, community membership, and niche evolution. We compared phylogenetic community structure and niche evolution in three lizard clades (Ctenotus skinks, agamids, and diplodactyline geckos) from arid Australia. We surveyed lizard communities at 32 sites in the northwestern Great Victoria Desert and generated complete species-level molecular phylogenies for regional representatives of the three clades. We document a striking pattern of phylogenetic evenness within local communities for all groups: pairwise correlations in species abundance across sites are negatively related to phylogenetic similarity. By modeling site suitability on the basis of species' habitat preferences, we demonstrate that phylogenetic evenness generally persists even after controlling for habitat filtering among species. This phylogenetic evenness is coupled with evolutionary lability of habitat-associated traits, to the extent that closely related species are more divergent in habitat use than distantly related species. In contrast, lizard diets are phylogenetically conserved, and pairwise dietary overlap between species is negatively related to phylogenetic distance in two of the three clades. Our results suggest that contemporary and historical species interactions have led to similar patterns of community structure across multiple clades in one of the world's most diverse lizard communities.     

Ecological release in lizard assemblages of neotropical savannas

We compare lizard assemblages of Cerrado and Amazonian savannas to test the ecological release hypothesis, which predicts that niche dimensions and abundance should be greater in species inhabiting isolated habitat patches with low species richness (Amazonian savannas and isolated Cerrado patches) when compared with nonisolated areas in central Cerrado with greater species richness. We calculated microhabitat and diet niche breadths with data from 14 isolated Cerrado patches and Amazon savanna areas and six central Cerrado populations. Morphological data were compared using average Euclidean distances, and lizard abundance was estimated using the number of lizards captured in pitfall traps over an extended time period. We found no evidence of ecological release with respect to microhabitat use, suggesting that historical factors are better microhabitat predictors than ecological factors. However, data from individual stomachs indicate that ecological release occurs in these areas for one species (Tropidurus) but not others (Ameiva ameiva, Anolis, Cnemidophorus, and Micrablepharus), suggesting that evolutionary lineages respond differently to environmental pressures, with tropidurids being more affected by ecological factors than polychrotids, teiids, and gymnophthalmids. We found no evidence that ecological release occurs in these areas using morphological data. Based on abundance data, our results indicate that the ecological release (density compensation) hypothesis is not supported: lizard species are not more abundant in isolated areas than in nonisolated areas. The ecology of species is highly conservative, varying little from assemblage to assemblage. Nevertheless, increases in niche breadth for some species indicate that ecological release occurs as well. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Functional and ecological relevance of intraspecific variation in body size and shape in the lizard Podarcis melisellensis (Lacertidae)

Within populations, individual animals may vary considerably in morphology and ecology. The degree to which variation in morphology is related to ecological variation within a population remains largely unexplored. We investigated whether variation in body size and shape among sexes and age classes of the lizard Podarcis melisellensis translates in differential whole-animal performance (sprint speed, bite force), escape and prey attack behaviour in the field, microhabitat use and diet. Male and female adult lizards differed significantly in body size and head and limb proportions. These morphological differences were reflected in differences in bite strength, but not in sprint speed. Accordingly, field measurements of escape behaviour and prey attack speed did not differ between the sexes, but males ate larger, harder and faster prey than females. In addition to differences in body size, juveniles diverged from adults in relative limb and head dimensions. These shape differences may explain the relatively high sprint and bite capacities of juvenile lizards. Ontogenetic variation in morphology and performance is strongly reflected in the behaviour and ecology in the field, with juveniles differing from adults in aspects of their microhabitat use, escape behaviour and diet.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 251–264.     

Historical contingency and animal diets: the origins of egg eating in snakes

Evolutionary changes in animal diets must often begin through the inclusion of a novel food type as a minor component of the diet. An aspect of this initial change that has rarely been studied is the relationship between the existing diet and the use of specific novel foods. We used comparative analyses to test the hypothesis that, in snakes, feeding on squamate (lizard and snake) eggs or bird eggs--items that represent evolutionarily derived and, in most cases, minor components of the diet--is associated with feeding on squamates or birds, respectively. Phylogenetic concentrated-changes tests indicate a significant tendency for predation on eggs to arise in snake lineages characterized by feeding on the corresponding animals. These results also generally hold for analyses including only snake species that are likely to encounter eggs and are large enough to ingest them. The inferred histories of specialized egg eaters also support the hypothesis. Because snakes often use chemical cues to recognize prey, the observed phylogenetic patterns might be explained by chemical similarities between eggs and adult animals. Our results suggest broad effects of predispositions on snake diets and thus illustrate how historical contingencies can shape the ecology of organisms.     

Does Intraspecific Size Variation in a Predator Affect Its Diet Diversity and Top-Down Control of Prey?

It has long been known that intraspecific variation impacts evolutionary processes, but only recently have its potential ecological effects received much attention. Theoretical models predict that genetic or phenotypic variance within species can alter interspecific interactions, and experiments have shown that genotypic diversity in clonal species can impact a wide range of ecological processes. To extend these studies to quantitative trait variation within populations, we experimentally manipulated the variance in body size of threespine stickleback in enclosures in a natural lake environment. We found that body size of stickleback in the lake is correlated with prey size and (to a lesser extent) composition, and that stickleback can exert top-down control on their benthic prey in enclosures. However, a six-fold contrast in body size variance had no effect on the degree of diet variation among individuals, or on the abundance or composition of benthic or pelagic prey. Interestingly, post-hoc analyses revealed suggestive correlations between the degree of diet variation and the strength of top-down control by stickleback. Our negative results indicate that, unless the correlation between morphology and diet is very strong, ecological variation among individuals may be largely decoupled from morphological variance. Consequently we should be cautious in our interpretation both of theoretical models that assume perfect correlations between morphology and diet, and of empirical studies that use morphological variation as a proxy for resource use diversity.     

Spatial and trophic niche differentiation in two sympatric lizards (Tropidurus torquatus and Cnemidophorus ocellifer) with different foraging tactics

Abstract In a ‘restinga’ habitat of southeastern Brazil, we studied the food habits and the microhabitat use of two lizards with distinct foraging modes: the tropidurid Tropidurus torquatus, a sit-and-wait predator, and the teiid Cnemidophorus ocellifer, a wide forager. The diet of the two species differed strongly, indicating a low level of similarity in their trophic niche. The sit-and-wait predator fed mainly on mobile prey, whereas the wide forager fed mainly on sedentary prey (larvae). The spatial niche breadth of T. torquatus was larger than that of C. ocellifer. Despite interspecific differences, the two species overlapped greatly in micro-habitat use. The data indicate that at Linhares the two lizard species differed more in food resources than in microhabitat, and that most of the food differences reflect the foraging patterns of the species.     

Seasonal variation in terrestrial resource subsidies influences trophic niche width and overlap in two aquatic snake species: a stable isotope approach

Quantifying diet is essential for understanding the functional role of species with regard to energy processing, transfer, and storage within ecosystems. Recently, variance structure in the stable isotope composition of consumer tissues has been touted as a robust tool for quantifying trophic niche width, a task that has previously proven difficult due to bias in direct dietary analyses and difficulties in integrating diet composition over time. We used carbon and nitrogen stable isotope analyses to examine trophic niche width of two sympatric aquatic snakes, banded watersnakes Nerodia fasciata and black swamp snakes Seminatrix pygaea inhabiting an isolated wetland where seasonal migrations of amphibian prey cause dramatic shifts in resource availability. Specifically, we characterized snake and prey isotope compositions through time, space, and ontogeny and examined isotope values in relation to prey availability and snake diets assessed by gut content analysis. We determined that prey cluster into functional groups based on similarity of isotopic composition and seasonal availability. Isotope variance structure indicated that the trophic niche width of the banded watersnake was broader (more generalist) than that of the black swamp snake. Banded watersnakes also exhibited seasonal variation in isotope composition, suggesting seasonal diet shifts that track amphibian prey availability. Conversely, black swamp snakes exhibited little seasonal variation but displayed strong ontogenetic shifts in carbon and nitrogen isotope composition that closely paralleled ontogenetic shifts in their primary prey, paedomorphic mole salamanders Ambystoma talpoideum. Although niche dimensions are often treated as static, our results demonstrate that seasonal shifts in niche dimensions can lead to changes in niche overlap between sympatric species. Such short‐term fluctuations in niche overlap can influence competitive interactions and consequently the composition and dynamics of communities and ecosystems.     

Is dietary or microhabitat specialization associated with environmental heterogeneity in horned lizards (Phrynosoma)?

Niche breadth is predicted to correlate with environmental heterogeneity, such that generalists will evolve in heterogeneous environments and specialists will evolve in environments that vary less over space and time. We tested the hypothesis that lizards in a heterogeneous environment were generalists compared to lizards in a homogeneous environment. We compared niche breadths of greater short‐horned lizards by quantifying resource selection in terms of two different niche axes, diet (prey items and trophic level), and microhabitat (ground cover and shade cover) between two populations occurring at different elevations. We assessed the heterogeneity of dietary and microhabitat resources within each population's environment by quantifying the availability of prey items, ground cover, and shade cover in each environment. Overall, our results demonstrate that despite differences in resource heterogeneity between elevations, resource selection did not consistently differ between populations. Moreover, environmental heterogeneity was not associated with generalization of resource use. The low‐elevation site had a broader range of available prey items, yet lizards at the high‐elevation site demonstrated more generalization in diet. In contrast, the high‐elevation site had a broader range of available microhabitats, but the lizard populations at both sites were similarly generalized for shade cover selection and were similarly specialized for ground cover selection. Our results demonstrate that environmental heterogeneity of a particular resource does not necessarily predict the degree to which organisms specialize on that resource.     

Microhabitat Variation Explains Local‐scale Distribution of Terrestrial Amazonian Lizards in Rondônia,Western Brazil

We investigate the role of ecology and phylogeny in the association between lizard abundance and microhabitat variables in an Amazon rain forest site. Using pitfall trap arrays, we collected data from 349 individuals belonging to 23 lizard species. After accounting for spatial autocorrelation and using a canonical correspondence analysis (CCA), we found that lizard captures were significantly associated with microhabitat variables, which accounted for 48 percent of the observed variation. Furthermore, a canonical phylogenetic ordination (CPO) indicated that microhabitat variables are more important in determining the distribution of lizard species than phylogenetic relationships among species. Termite nests, canopy openness, and tree circumference were strongly associated with the number of captures of certain lizard species. Our results confirm autecology studies of individual lizard species for which data are available. We suggest that maintaining heterogeneous forested microhabitats should be a central goal for sustaining a high lizard biodiversity in Amazon rain forests.     

Change your diet or die: predator-induced shifts in insectivorous lizard feeding ecology

Animal feeding ecology and diet are influenced by the fear of predation. While the mechanistic bases for such changes are well understood, technical difficulties often prevent testing how these mechanisms interact to affect a mesopredator’s diet in natural environments. Here, we compared the insectivorous lizard Acanthodactylus beershebensis’ feeding ecology and diet between high- and low-risk environments, using focal observations, intensive trapping effort and fecal pellet analysis. To create spatial variation in predation risk, we planted “artificial trees” in a scrubland habitat that lacks natural perches, allowing avian predators to hunt for lizards in patches that were previously unavailable to them. Lizards in elevated-risk environments became less mobile but did not change their microhabitat use or temporal activity. These lizards changed their diet, consuming smaller prey and less plant material. We suggest that diet shifts were mainly because lizards from risky environments consumed prey items that required shorter handling time.     

Resource partitioning in sympatric langurs and macaques in tropical rainforests of the central Western Ghats, south India

In a competitive sympatric association, coexisting species may try to reduce interspecific interactions as well as competition for similar resources by several ecological and behavioral practices. We studied resource utilization of three sympatric primate species namely, lion-tailed macaques (Macaca silenus), bonnet macaques (M. radiata) and Hanuman langurs (Semnopithecus entellus) in a tropical rainforest of the central Western Ghats, south India. We studied resource use, tree-height use, foraging height, substrate use when consuming animal prey and interspecific interactions. The results revealed that across the year, there was very limited niche overlap in diet between each species-pair. Each primate species largely depended on different plant species or different plant parts and phenophases from shared plant species. Primate species used different heights for foraging, and the two macaque species searched different substrates when foraging on animal prey. We also recorded season-wise resource abundance for the resources shared by these three primate species. While there was low dietary overlap during the dry season (a period of relatively low resource abundance), there was high dietary overlap between the two macaque species during the wet season (a period of high resource abundance for the shared resources). We observed only a few interspecific interactions. None of these were agonistic, even during the period of high niche overlap. This suggests that the sympatric primate species in this region are characterized by little or no contest competition. Unlike in some other regions of the Western Ghats, the lack of interspecific feeding competition appears to allow these primates, especially the macaques, to remain sympatric year-round.     

Food partitioning and community organization in a mountain lizard guild of Northern Mexico

Summary Food niche relationships among four sympatric Sceloporus species were studied in the Sierra Madre Occidental, N.E. Mexico. Although some very high food-niche overlap values were observed, this does not prove that interspecific competition is currently important in organizing this lizard assemblage. Moreover, explaining habitat segregation among the coexisting species as an ecological result of interspecific competitive pressure is unlikely: the overall ecology and behaviour of these species is too much dependent on their microhabitat or substrate specialization to allow such an exclusive interpretation. Thus, this community is probably not mainly organized by species interactions but rather through the specific ecological needs of each species.     

Emergent impacts of cannibalism and size refuges in prey on intraguild predation systems

Many organisms undergo ontogenetic niche shifts due to considerable changes in size during their development. These ontogenetic shifts can alter the trophic position of individuals, the type and strength of ecological interactions across species, and allow for cannibalism within species. In this study we ask if and how the interaction of a size refuge and cannibalism in the prey alters the dynamics of intraguild predation (IGP) systems. By manipulating the composition of large cannibalistic (Aeshna umbrosa) and predatory (Anax junius) dragonfly larvae in mesocosms we show that the interaction of cannibals and predators was non-linear and increased the survival of prey. The structure of the final resource community shared by prey and predator differed between small and large dragonfly treatments but not within size classes across species. In general, the small prey stage showed similar shifts in microhabitat use and refuge use when exposed to either conspecific cannibals or predators, while large cannibals showed no clear anti-predator response. However, further behavioral experiments revealed that specific behavioral components, such as distances between individuals or number of movements, differed when individuals were exposed to either cannibals or predators. This indicates that individuals discriminated between conspecific or heterospecific predators. Furthermore, in similar experiments large cannibals and predators showed different behaviors when exposed to conspecifics rather than to each other. These changes in behavior are consistent with the observed increase in prey survival. In general, the results indicate that cannibalism and ontogenetic niche shifts can result in behavior-mediated indirect interactions that reduce the impact of the predator on the mortality of its prey and alter the interactions of IGP systems. However, they also indicate that size is not the sole determinant and that we also need to account for the species identity when predicting the dynamics of communities.     

Prey use of wetland benthivorous sunfishes: ontogenetic, interspecific and seasonal variation

The intensity of competitive interactions between fishes is partly determined by prey use and ontogenetic niche shifts. In a wetland where distinct habitat shifts are missing we compared prey use of three generalist benthivorous sunfishes to look for evidence of ontogenetic, interspecific, and “seasonal” variation in prey composition. Diet analysis revealed evidence of diet ontogeny in warmouth (Lepomis gulosus, 30–152 mm standard length, SL), but not in bluespotted sunfish (Enneacanthus gloriosus, 30–47 mm SL) or dollar sunfish (Lepomis marginatus, 30–60 mm SL). Bluespotted and dollar sunfishes consumed small dipteran and amphipod prey and had similar diets in both seasons suggesting a potential for strong interspecific competition. In the dry season, warmouth shifted from using smaller insect prey to larger decapod and fish prey with increasing size. This shift to prey types that were little used by the other species reduced dietary niche overlap with the other sunfishes. After drought and re-flooding (in the wet season), decapods and small fish were less abundant in the wetland and the warmouth ontogenetic shift was less distinct. When matched for gape width, prey composition differed between warmouth and both dollar and bluespotted sunfishes in the wet season, suggesting differences in sunfish foraging modes, but prey use differences were less clear in the dry season when prey were abundant. Both warmouth ontogenetic diet shifts and seasonal variation in prey use (probably mediated by prey abundance) had strong influences on diet overlap and therefore the potential for intra- and interspecific competition between sunfishes in this wetland ecosystem.     

Fixed prey cue preferences among Dusky Pigmy Rattlesnakes (<Emphasis Type="Italic">Sistrurus miliarius barbouri</Emphasis>) raised on different long-term diets

Chemoreception is often crucial to the interaction between predators and their prey. Investigating the mechanisms controlling predator chemical preference gives insight into how selection molds traits directly involved in ecological interactions between species. In snakes, prey cue preferences are influenced by both direct genetic control and experience-based plasticity. We assessed prey preference in a group of Dusky Pigmy Rattlesnakes that had eaten only mice or lizards over a 5 year period to test whether genetics or plasticity primarily determine the preference phenotype. Our results provide evidence for genetic determination of preference for lizard chemical cues in pigmy rattlesnakes. Snakes preferred the scent of lizards, regardless of their initial diet, and the response to mouse scent did not differ from the water-only control. We discuss these findings in light of previous studies that manipulated snake diets over shorter timescales.