Laurie J. Vitt

Declining amphibian populations : the problem of separating human impacts from natural fluctuations

Reports of declining amphibian populations in many parts of the world are numerous, but supporting long-term census data are generally unavailable. Census data from 1979 to 1990 for three salamander species and one frog species at a breeding pond in South Carolina showed fluctuations of substantial magnitude in both the size of breeding populations and in recruitment of juveniles. Breeding population sizes exhibited no overall trend in three species and increased in the fourth. Recent droughts account satisfactorily for an increase in recruitment failures. These data illustrate that to distinguish between natural population fluctuations and declines with anthropogenic causes may require long-term studies.

Why tropical forest lizards are vulnerable to climate warming

Biological impacts of climate warming are predicted to increase with latitude, paralleling increases in warming. However, the magnitude of impacts depends not only on the degree of warming but also on the number of species at risk, their physiological sensitivity to warming and their options for behavioural and physiological compensation. Lizards are useful for evaluating risks of warming because their thermal biology is well studied. We conducted macrophysiological analyses of diurnal lizards from diverse latitudes plus focal species analyses of Puerto Rican Anolis and Sphaerodactyus. Although tropical lowland lizards live in environments that are warm all year, macrophysiological analyses indicate that some tropical lineages (thermoconformers that live in forests) are active at low body temperature and are intolerant of warm temperatures. Focal species analyses show that some tropical forest lizards were already experiencing stressful body temperatures in summer when studied several decades ago. Simulations suggest that warming will not only further depress their physiological performance in summer, but will also enable warm-adapted, open-habitat competitors and predators to invade forests. Forest lizards are key components of tropical ecosystems, but appear vulnerable to the cascading physiological and ecological effects of climate warming, even though rates of tropical warming may be relatively low.

Sexual Selection Versus Alternative Causes of Sexual Dimorphism in Teiid Lizards

SummaryThe presence and extent of sexual dimorphisms in body form (size and shape) of adult macroteiid lizards were investigated. Males were significantly larger than females in the temperate species, Cnemidophorus tigris, and in the tropical species, Ameiva ameiva and C. ocellifer. Young adult C. tigris males grew faster than young adult females within and between reproductive seasons. Adult males of all species had larger heads than adult females of the same body size; this difference increased with body size. Moreover, male C. tigris were heavier than females of the same snout-vent length. The causes and consequences of the sexual dimorphisms were also examined. The possible causes of body size are especially numerous, and distinguishing the relative influences of the various causal selection factors on body size is problematical. Nevertheless, observational field data were used to tentatively conclude that intrasexual selection was the cause of larger body size of C. tigris males relative to females because (1) larger males won in male aggressive interactions, (2) the winning males gained access to more females by repelling competitors and by female acceptance, (3) larger males consequently had higher reproductive success, and (4) other hypothetical causes of larger male size were unsupported.

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.

The phylogeny of advanced snakes (Colubroidea), with discovery of a new subfamily and comparison of support methods for likelihood trees

The superfamily Colubroidea (> 2500 species) includes the majority of snake species and is one of the most conspicuous and well-known radiations of terrestrial vertebrates. However, many aspects of the phylogeny of the group remain contentious, and dozens of genera have yet to be included in molecular phylogenetic analyses. We present a new, large-scale, likelihood-based phylogeny for the colubroids, including 761 species sampled for up to five genes: cytochrome b (93% of 761 species sampled), ND4 (69%), ND2 (28%), c-mos (54%), and RAG-1 (13%), totaling up to 5814bp per species. We also compare likelihood bootstrapping and a recently proposed ultra-fast measure of branch support (Shimodaira-Hasegawa-like [SHL] approximate likelihood ratio), and find that the SHL test shows strong support for several clades that were weakly-supported by bootstrapping in this or previous analyses (e.g., Dipsadinae, Lamprophiidae). We find that SHL values are positively related to branch lengths, but show stronger support for shorter branches than bootstrapping. Despite extensive missing data for many taxa (mean=67% per species), neither bootstrap nor SHL support values for terminal species are related to their incompleteness, and that most highly incomplete taxa are placed in the expected families from previous taxonomy, typically with very strong support. The phylogeny indicates that the Neotropical colubrine genus Scaphiodontophis represents an unexpectedly ancient lineage within Colubridae. We present a revised higher-level classification of Colubroidea, which includes a new subfamily for Scaphiodontophis (Scaphiodontophiinae). Our study provides the most comprehensive phylogeny of Colubroidea to date, and suggests that SHL values may provide a useful complement to bootstrapping for estimating support on likelihood-based trees.

Lizard Ecology: Historical and Experimental Perspectives

In a collection rich in implications for all fields of ecology, leading lizard ecologists demonstrate the utility of the phylogenetic approach in understanding the evolution of morphology, physiology, behavior, and life histories. Lizards, which are valued for their amenability to field experiments, have been the subject of reciprocal transplant experiments and of manipulations of resource availability, habitat structure, population density, and entire sections of food webs. Such experiments are rapidly rebuilding ecological theories as they apply to all organisms. As a demonstration of state-of-the-art historical and experimental research and as a call for philosophical engagement, this volume will join its predecessors--Lizard Ecology: A Symposium (Missouri, 1967) and Lizard Ecology: Studies of a Model Organism (Harvard, 1983)--in directing ecological research for years to come.Lizard Ecology contains essays on reproductive ecology (Arthur E. Dunham, Lin Schwarzkopf, Peter H. Niewiarowski, Karen Overall, and Barry Sinervo), behavioral ecology (A. Stanley Rand, William E. Cooper, Jr., Em!lia P. Martins, Craig Guyer, and C. Michael Bull), evolutionary ecology (Raymond B. Huey, Jean Clobert et al., Donald B. Miles, and Theodore Garland, Jr.), and population and community ecology (Ted Case, Robin M. Andrews and S. Joseph Wright, Craig D. James, and Jonathan B. Losos).

Geckos: Adaptive Significance and Energetics of Tail Autotomy

Coleonyx variegatus is adapted to readily sacrifice its tail to predators. This adaptation is associated with characteristic tail behavior and rapid tail regeneration. There is no facultative metabolic increase associated with tail regeneration, and energy normally allocated to body growth and maintenance is diverted to tail regeneration. This supports the contention that tail behavior, autotomy, and rapid regeneration evolved as mechanisms promoting survival in terms of predator escape.

A COMPARISON OF EVOLUTIONARY RADIATIONS IN MAINLAND AND CARIBBEAN ANOLIS LIZARDS

Comparisons between closely related radiations in different environments provide a unique window into understanding how abiotic and biotic factors shape evolu- tionary pathways. Anolis lizards have radiated extensively in the West Indies, as well as mainland Central and South America. In the Caribbean, similar communities of anole species specialized for different habitats (ecomorphs) have evolved independently on each Greater Antillean island. We examined ecological and morphological data on 49 Anolis species (33 Caribbean, 16 mainland) to investigate whether the same set of ecomorphs has arisen in mainland regions. More generally, we investigated whether the relationship between ecology and morphology was similar among anoles in the two regions. Radiations in the two regions are very different. The majority of mainland anole species exhibit morphological characteristics unlike any Caribbean ecomorph. Furthermore, rela- tionships between ecology and morphology are very different between the two sets of anole species. Among mainland anole species, toepad size is positively correlated with perch height, whereas tail length is negatively related to perch diameter. In contrast, among Caribbean anole species, both forelimb length and body mass are positively associated with perch diameter, and both tail length and hindlimb length are negatively related to perch diameter. Biomechanical considerations provide a functional basis for some of these cor- relations, but much variation remains to be explained. These findings demonstrate that factors that caused anole species to converge repeatedly in the West Indies are not present in mainland regions, and that environmental factors can strongly influence the shape of evolutionary radiations.

Adaptive Strategies and Energetics of Tail Autonomy in Lizards

Various methods were used to study and quantify components of tail autotomy adapta- tions in four species of lizards; Coleonyx variegatus (Gekkonidae), Eumeces skiltonianus and Eumeces gilberti (Scincidae), and Gerrhonotus multicarinatus (Anguidae). High frequency of tail breaks in natural populations of these species (65%, 62%, 52%, and 74% respectively) suggests the effectiveness of autotomy for predator escape. In species where tail autotomy is most important for escape, regener- ated tails are as large or larger than the original tails. Regenerated tails were higher in lipids and energy than original tails, which were, in turn, higher than bodies. The species with alternate uses for the tail (other than predator escape via autotomy) exhibited lower rates of energy allocation to regeneration and the regenerated tails were lower in energy content (both per unit weight and total, proportionately) than tails of other species studied. Total growth energy allocated to tail regeneration varied from 24.7% in G. multicarinatus (the species least adapted for autotomy) to 56.5% in E. gilberti (a species highly adapted for autotomy). Regeneration was most rapid in short-lived early-maturing species, those with several breeding opportunities in a given season. Species with high probabilities of future reproduc- tions (long-lived, late-maturing) and short, seasonally restricted breeding seasons, regenerated tails at low rates. Energy allocations to tail regeneration were varied within and among species. Generally, immatures of all species except G. multicarinatus allocated proportionately less to tail regeneration and more to body growth than adults. Energy allocations to tail regeneration in G. multi- carinatus were relatively consistent across age groups. A summary of pathways by which autotomy adaptations may be selected for, or by which retention adaptations may evolve, is presented.

Distribution, extent, and evolution of plant consumption by lizards

Animal diets crucially affect fitness, yet many aspects of their ultimate determinants are unknown. The distribution and extent of herbivory in lizards, its evolutionary history, and ecological factors that may favour it are discussed. Most lizards are exclusively or primarily carnivorous, yet many species eat some plants and a few are almost exclusively herbivorous. Based on a literature survey of diets of over 450 lizard species, the distribution and degree of omnivory and herbivory are described. Some plants occur in the diets of slightly over half of lizard species, and plants formed 10% or more of the dietary volume of 12.1% of species, and 90% or more of the diet of 0.8% of species. The greatest percentage of omnivorous species (> 10% plant diet), over 30% in each, and highest mean percentage plant matter in the diet are in Iguanidae, Corytophanidae, Gerrhosauridae, Agamidae, Xantusiidae, and Tropiduridae. Numerous other omnivores occur in Lacertidae and Scincidae and fewer in several additional families. Herbivorous lizards (> 90% plant volume) tend to be folivorous and to possess adaptations for processing leaves, including specialized dentition for cutting or reducing leaves, elongated intestines, colic valves that slow passage of food, and intestinal flora that digest cellulose. Omnivorous lizards lacking such specializations may eat some leaves, but consume much more fruit, flowers, and seeds, plant parts that are easy to digest, likely to be very abundant seasonally, and may be highly nutritious. Some lizards eat nectar and pollen; even sap is eaten by at least one gecko. Ontogenetic increase in plant consumption and decrease in prey consumption is known, but its generality has been controversial. Such ontogeny has been demonstrated in three iguanid species, a skink, a lacertid, two tropidurids, a phrynosomatid, and two corytophanids, but it does not occur in some other species. Thus, the presences of ontogenetic variation must be evaluated for each species. The importance of specific foods may also vary with age. Omnivory and/or herbivory have originated in many lizard families, with at least nine origins in Iguania and 23 in Scleroglossa. Origins have been rare in Gekkonoidea and Anguimorpha and common in Scincomorpha, especially in Lacertidae and Scincidae. Losses of omnivory have been much less frequent than gains. Only a few origins can account for all the herbivory in lizards. Concentrated changes tests show that there is a significant association in Lacertidae, Lacertiformes, Lacertoidea, Scincidae, and Scleroglossa between insularity and omnivory. Insular lizards may broaden their diets to compensate for limited availability of prey. Addition of other factors that reduce availability of prey, i.e. extreme aridity and cave-dwelling, to insularity, strengthened the relationship to omnivory in Lacertidae and Lacertoidea. We were unable to demonstrate a role of aridity independent of insularity, but present anecdotal evidence suggests that it may promote evolution of plant consumption. Large body size in lizards has long been associated with herbivory, and more recently, with omnivory in lacertid lizards. Using a conventional regression approach in which each species is considered to supply an independent data point, this relationship was confirmed for all lizards. Although larger species have diets with more plants, plant consumption accounts for only 9% of the variation in body length, which is not surprising given that other factors such as predation, competition, and sexual selection affect body size. The frequency of transitions body size associated with transitions to ominivory or carnivory was also examined. In Iguania, Scleroglossa, and all lizards, transitions supporting the hypothesis that omnivory favours increase in body size were significantly more frequent than nonsupporting transitions. This suggests that substantial plant consumption favours evolution of larger size, probably because of the energetic considerations first presented by Pough (1973). Because actively foraging lizards move widely through the habitat to locate prey and tongue-flick to locate prey by chemical cues, we