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"Pianka, Eric R."
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Life-History Patterns of Lizards of the World
Identification of mechanisms that promote variation in life-history traits is critical to understand the evolution of divergent reproductive strategies. Here we compiled a large life-history data set (674 lizard populations, representing 297 species from 263 sites globally) to test a number of hypotheses regarding the evolution of life-history traits in lizards. We found significant phylogenetic signal in most life-history traits, although phylogenetic signal was not particularly high. Climatic variables influenced the evolution of many traits, with clutch frequency being positively related to precipitation and clutches of tropical lizards being smaller than those of temperate species. This result supports the hypothesis that in tropical and less seasonal climates, many lizards tend to reproduce repeatedly throughout the season, producing smaller clutches during each reproductive episode. Our analysis also supported the hypothesis that viviparity has evolved in lizards as a response to cooler climates. Finally, we also found that variation in trait values explained by clade membership is unevenly distributed among lizard clades, with basal clades and a few younger clades showing the most variation. Our global analyses are largely consistent with life-history theory and previous results based on smaller and scattered data sets, suggesting that these patterns are remarkably consistent across geographic and taxonomic scales.
Journal Article
Deep History Impacts Present-Day Ecology and Biodiversity
Lizards and snakes putatively arose between the early Jurassic and late Triassic; they diversified worldwide and now occupy many different ecological niches, making them ideal for testing theories on the origin of ecological traits. We propose and test the \"deep history hypothesis,\" which claims that differences in ecological traits among species arose early in evolutionary history of major clades, and that present-day assemblages are structured largely because of ancient, preexisting differences. We combine phylogenetic data with ecological data collected over nearly 40 years to reconstruct the evolution of dietary shifts in squamate reptiles. Data on diets of 184 lizard species in 12 families from 4 continents reveal significant dietary shifts at 6 major divergence points, reducing variation by 79.8%. The most striking dietary divergence (27.6%) occurred in the late Triassic, when Iguania and Scleroglossa split. These two clades occupy different regions of dietary niche space. Acquisition of chemical prey discrimination, jaw prehension, and wide foraging provided scleroglossans access to sedentary and hidden prey that are unavailable to iguanians. This cladogenic event may have profoundly influenced subsequent evolutionary history and diversification. We suggest the hypothesis that ancient events in squamate cladogenesis, rather than present-day competition, caused dietary shifts in major clades such that some lizard clades gained access to new resources, which in turn led to much of the biodiversity observed today.
Journal Article
History and the Global Ecology of Squamate Reptiles
The structure of communities may be largely a result of evolutionary changes that occurred many millions of years ago. We explore the historical ecology of squamates (lizards and snakes), identify historically derived differences among clades, and examine how this history has affected present‐day squamate assemblages globally. A dietary shift occurred in the evolutionary history of squamates. Iguanian diets contain large proportions of ants, other hymenopterans, and beetles, whereas these are minor prey in scleroglossan lizards. A preponderance of termites, grasshoppers, spiders, and insect larvae in their diets suggests that scleroglossan lizards harvest higher energy prey or avoid prey containing noxious chemicals. The success of this dietary shift is suggested by dominance of scleroglossans in lizard assemblages throughout the world. One scleroglossan clade, Autarchoglossa, combined an advanced vomeronasal chemosensory system with jaw prehension and increased activity levels. We suggest these traits provided them a competitive advantage during the day in terrestrial habitats. Iguanians and gekkotans shifted to elevated microhabitats historically, and gekkotans shifted activity to nighttime. These historically derived niche differences are apparent in extant lizard assemblages and account for some observed structure. These patterns occur in a variety of habitats at both regional and local levels throughout the world.
Journal Article
Stable isotope ecology of a hyper-diverse community of scincid lizards from arid Australia
We assessed the utility of stable isotope analysis as a tool for understanding community ecological structure in a species-rich clade of scincid lizards from one of the world's most diverse lizard communities. Using a phylogenetic comparative framework, we tested whether δ15N and δ13C isotopic composition from individual lizards was correlated with species-specific estimates of diet and habitat use. We find that species are highly divergent in isotopic composition with significant correlations to habitat use, but this relationship shows no phylogenetic signal. Isotopic composition corresponds to empirical observations of diet for some species but much variation remains unexplained. We demonstrate the importance of using a multianalytical approach to questions of long-term dietary preference, and suggest that the use of stable isotopes in combination with stomach content analysis and empirical data on habitat use can potentially reveal patterns in ecological traits at finer scales with important implications for community structuring.
Journal Article
How do lizard niches conserve, diverge or converge? Further exploration of saurian evolutionary ecology
Background: Environmental conditions on Earth are repeated in non-random patterns that often coincide with species from different regions and time periods having consistent combinations of morphological, physiological and behavioral traits. Observation of repeated trait combinations among species confronting similar environmental conditions suggest that adaptive trait combinations are constrained by functional tradeoffs within or across niche dimensions. In an earlier study, we assembled a high-resolution database of functional traits for 134 lizard species to explore ecological diversification in relation to five fundamental niche dimensions. Here we expand and further examine multivariate relationships in that dataset to assess the relative influence of niche dimensions on the distribution of species in 6-dimensional niche space and how these may deviate from distributions generated from null models. We then analyzed a dataset with lower functional-trait resolution for 1023 lizard species that was compiled from our dataset and a published database, representing most of the extant families and environmental conditions occupied by lizards globally. Ordinations from multivariate analysis were compared with null models to assess how ecological and historical factors have resulted in the conservation, divergence or convergence of lizard niches. Results: Lizard species clustered within a functional niche volume influenced mostly by functional traits associated with diet, activity, and habitat/substrate. Consistent patterns of trait combinations within and among niche dimensions yielded 24 functional groups that occupied a total niche space significantly smaller than plausible spaces projected by null models. Null model tests indicated that several functional groups are strongly constrained by phylogeny, such as nocturnality in the Gekkota and the secondarily acquired sit-and-wait foraging strategy in Iguania. Most of the widely distributed and species-rich families contained multiple functional groups thereby contributing to high incidence of niche convergence. Conclusions: Comparison of empirical patterns with those generated by null models suggests that ecological filters promote limited sets of trait combinations, especially where similar conditions occur, reflecting both niche convergence and conservatism. Widespread patterns of niche convergence following ancestral niche diversification support the idea that lizard niches are defined by trait-function relationships and interactions with environment that are, to some degree, predictable and independent of phylogeny.
Journal Article
Toward a Periodic Table of Niches, or Exploring the Lizard Niche Hypervolume
Widespread niche convergence suggests that species can be organized according to functional trait combinations to create a framework analogous to a periodic table. We compiled ecological data for lizards to examine patterns of global and regional niche diversification, and we used multivariate statistical approaches to develop the beginnings for a periodic table of niches. Data (50+ variables) for five major niche dimensions (habitat, diet, life history, metabolism, defense) were compiled for 134 species of lizards representing 24 of the 38 extant families. Principal coordinates analyses were performed on niche dimensional data sets, and species scores for the first three axes were used as input for a principal components analysis to ordinate species in continuous niche space and for a regression tree analysis to separate species into discrete niche categories. Three-dimensional models facilitate exploration of species positions in relation to major gradients within the niche hypervolume. The first gradient loads on body size, foraging mode, and clutch size. The second was influenced by metabolism and terrestrial versus arboreal microhabitat. The third was influenced by activity time, life history, and diet. Natural dichotomies are activity time, foraging mode, parity mode, and habitat. Regression tree analysis identified 103 cases of extreme niche conservatism within clades and 100 convergences between clades. Extending this approach to other taxa should lead to a wider understanding of niche evolution.
Journal Article
Optimal foraging constrains macroecological patterns: body size and dietary niche breadth in lizards
To explore and identify probable mechanisms contributing to the relationships among body size, dietary niche breadth and mean, minimum, maximum and range of prey size in predaceous lizards. Our data set includes species from tropical rainforests, semi-arid regions of Brazil, and from deserts of the south-western United States, Australia and the Kalahari of Africa. We calculated phylogenetic and non-phylogenetic regressions among predator body size, dietary breath and various prey size measures. We found a negative association between body size and dietary niche breadth in 159 lizard species sampled across most evolutionary lineages of squamate reptiles and across major continents and habitats. We also show that mean, minimum, maximum and range of prey size were positively associated with body size. Our results suggest not only that larger lizards tend to eat larger prey, but in doing so offset their use of smaller prey. Reduction of dietary niche breadth with increased body size in these lizards suggests that large predators target large and more profitable prey. Consequently, the negative association between body size and niche breadth in predators most likely results from optimal foraging. Though this result may appear paradoxical and runs counter to previous studies, resources for predators may be predictably more limited than resources for herbivores, thus driving selection for more profitable prey.
Journal Article
Lizard Thermal Biology: Do Genders Differ?
For more than six decades, physiological ecologists have intensively studied diverse aspects of lizard thermal biology. Nevertheless, a recent review notes that prior studies have generally ignored gender differences in body temperatures, thermal sensitivity, or other aspects of thermal biology. We concur that gender differences have been ignored and should be examined: if gender differences prove common, standard protocols for studying lizard natural history, thermal physiology, and ecology will require significant modification. To help resolve this issue, we conducted a retrospective analysis of our huge data set on the thermal biology of many desert lizards (more than 11,000 individuals from 56 species in seven major clades) from Africa, Australia, and North America. Results are unambiguous: gender differences in body temperature, air temperature, and time of activity—and thus in field thermal biology—are almost always minor. In fact, mean body temperatures of males and females differ by less than 1°C in 80.4% of species. For desert lizards, gender differences in thermal biology are the exception, not the rule. Nevertheless, gender differences should be examined when feasible because exceptions—though likely rare—could be biologically interesting.
Journal Article
Body temperature distributions of active diurnal lizards in three deserts
The performance of ectotherms integrated over time depends in part on the position and shape of the distribution of body temperatures (Tb) experienced during activity. For several complementary reasons, physiological ecologists have long expected that Tb distributions during activity should have a long left tail (left‐skewed), but only infrequently have they quantified the magnitude and direction of Tb skewness in nature. To evaluate whether left‐skewed Tb distributions are general for diurnal desert lizards, we compiled and analysed Tb (∑ = 9,023 temperatures) from our own prior studies of active desert lizards in three continents (25 species in Western Australia, 10 in the Kalahari Desert of Africa and 10 species in western North America). We gathered these data over several decades, using standardized techniques. Many species showed significantly left‐skewed Tb distributions, even when records were restricted to summer months. However, magnitudes of skewness were always small, such that mean Tb were never more than 1°C lower than median Tb. The significance of Tb skewness was sensitive to sample size, and power tests reinforced this sensitivity. The magnitude of skewness was not obviously related to phylogeny, desert, body size or median body temperature. Moreover, a formal phylogenetic analysis is inappropriate because geography and phylogeny are confounded (i.e. are highly collinear). Skewness might be limited if lizards pre‐warm inside retreats before emerging in the morning, emerge only when operative temperatures are high enough to speed warming to activity Tb, or if cold lizards are especially wary and difficult to spot or catch. Telemetry studies may help evaluate these possibilities. A plain language summary is available for this article. Plain Language Summary
Journal Article