Lizard Sceloporus Undulatus Research Articles

Fitness Costs of Maternal Ornaments and Prenatal Corticosterone Manifest as Reduced Offspring Survival and Sexual Ornament Expression.

Colorful traits (i.e., ornaments) that signal quality have well-established relationships with individual condition and physiology. Furthermore, ornaments expressed in females may have indirect fitness effects in offspring via the prenatal physiology associated with, and social consequences of, these signaling traits. Here we examine the influence of prenatal maternal physiology and phenotype on condition-dependent signals of their offspring in adulthood. Specifically, we explore how prenatal maternal testosterone, corticosterone, and ornament color and size correlate with female and male offspring survival to adulthood and ornament quality in the lizard Sceloporus undulatus. Offspring of females with more saturated badges and high prenatal corticosterone were less likely to survive to maturity. Badge saturation and area were negatively correlated between mothers and their male offspring, and uncorrelated to those in female offspring. Maternal prenatal corticosterone was correlated negatively with badge saturation of male offspring in adulthood. Our results indicate that maternal ornamentation and prenatal concentrations of a stress-relevant hormone can lead to compounding fitness costs by reducing offspring survival to maturity and impairing expression of a signal of quality in surviving males. This mechanism may occur in concert with social costs of ornamentation in mothers. Intergenerational effects of female ornamentation and prenatal stress may be interdependent drivers of balancing selection and intralocus sexual conflict over signaling traits.

History of predator exposure affects cell-mediated immunity in female eastern fence lizards, Sceloporus undulatus (Squamata: Phrynosomatidae)

Abstract On exposure to stressors, energy is diverted from non-urgent functions towards those important for immediate survival. The degree and nature of reallocation may be affected by the evolutionary history of the animal. The eastern fence lizard (Sceloporus undulatus) coexists in parts of its range with invasive fire ants (Solenopsis invicta), which attack and wound lizards and elevate stress-relevant hormones (corticosterone), whereas other populations have never been exposed to fire ants. We examined how a history of fire ant invasion affected the immune response in female lizards after exposure to exogenous corticosterone (mimicking exposure to a stressor) during gestation (dosing regimens differed among corticosterone-exposed lizards owing to the constraints of the original studies, but we found no evidence that this affected the outcome of the present study). A history of exposure to predatory stressors (fire ants) and corticosterone treatment affected cell-mediated immunity. Lizards from fire ant-invaded sites had a reduced immune response compared with those from uninvaded sites. Corticosterone treatment had no effect on the immune response of lizards from invaded sites but reduced the immune response of lizards from uninvaded sites. This suggests that an evolutionary history of exposure to wounding alters the immune response to corticosterone. Future work on how the immune system responds to environmental threats will be informative for the prediction and management of these threats.

Body Color and Morphological Correlates of Fitness in Eastern Fence Lizards (Sceloporus undulatus): A Spectrophotometric Approach

Many lizards use colorful badges in displays with conspecifics during courtship and aggressive interactions. In Eastern Fence Lizards (Sceloporus undulatus), males and females reveal sexually dimorphic ventral abdominal coloration during social interactions and use these features to signal sex and perhaps other characteristics. However, the extent to which ventral badges or other chromatic features of males and females signal quality remains unresolved in this species. Additionally, adult S. undulatus exhibit temperature-dependent color change of their abdominal badges, a potential confounding variable in studies of the function of these badges. Here, we examined the relationship between spectral variables of ventral and dorsal skin color and morphometric traits linked to fitness in adult male and female S. undulatus under standardized temperature conditions. For males, ventral patch hue tended to decrease (i.e., was blue-shifted) as body size increased, whereas dorsal hue was unrelated to male size. In contrast, there was no relationship between ventral hue and body size in females, and dorsal hue increased (i.e., was red-shifted) with body size. In males, lower ultraviolet (UV) chroma and greater blue chroma of ventral abdominal badges predicted increasing body size. In females, UV chroma of the ventral abdomen was inversely related to body size and better body condition, whereas blue chroma was inversely related to body condition only. Dorsal UV chroma decreased with increasing body size in both males and females, but brightness was not a significant predictor of any morphometric trait in either sex. Overall, these results indicate that both blue and UV reflectance of ventral and dorsal abdominal skin are indices of size and thus age, and could therefore serve as indices of signaler quality in this species.

Maternal stress alters the phenotype of the mother, her eggs and her offspring in a wild-caught lizard.

While biomedical researchers have long appreciated the influence of maternally derived glucocorticoids (GCs) on offspring phenotype, ecologists have only recently begun exploring its impact in wild animals. Interpreting biomedical findings within an ecological context has posited that maternal stress, mediated by elevations of maternal GCs, may play an adaptive role preparing offspring for a stressful or rigorous environment. Yet, the influence of maternal stress on offspring phenotype has been little studied in wild animals. We experimentally elevated GCs to ecologically relevant levels (mimicking increases in maternal stress hormones following a nonlethal predator encounter, a heat challenge, or a chasing or confinement stressor) in female eastern fence lizards Sceloporus undulatus during gestation. We tested the hypothesis that maternally derived stress hormones themselves are sufficient to alter offspring phenotype. Specifically, we examined the effects of experimentally elevated maternal GCs on fitness-relevant traits of the mother, her eggs and her subsequent offspring. We found that daily maternal GC elevation: (a) increased maternal antipredator behaviours and postlaying glucose levels; (b) had no effect on egg morphology or caloric value, but altered yolk hormone (elevated GC) and nutrient content; and (c) altered offspring phenotype including stress-relevant physiology, morphology and behaviour. These findings reveal that maternally derived GCs alone can alter offspring phenotype in a wild animal, changes that may be mediated via maternal behaviour, and egg hormone and nutrient content. Understanding the ecological consequences of these effects under different environmental conditions will be critical for determining the adaptive significance of elevated maternal GCs for offspring.

Asymmetrical mate preference in recently adapted White Sands and black lava populations of Sceloporus undulatus

Abstract Speciation can proceed rapidly when natural and sexual selection act in concert. For example speciation can be accelerated when traits that confer a selective advantage in a particular habitat also influence mate preference. Studying parallel but evolutionarily independent instances of ecological divergence can illuminate the interaction between natural and sexual selection during speciation. Locally adapted populations of the eastern fence lizard Sceloporus undulatus have recently evolved in three different habitats in the Chihuahuan desert: blanched color morphs occur on the gypsum dunes of White Sands, melanic color morphs occur on the Carrizozo lava flow, and brown color morphs occur in the surrounding desert scrubland. In addition to differences in cryptic dorsal coloration, populations also differ in the size and color of ventral patches used for social signaling. This system therefore provides an opportunity to investigate the interplay of natural and sexual selection during rapid ecological speci-ation. We used mate preference experiments to determine whether locally adapted populations may exhibit the early stages of behavioral reproductive isolation. We observed an asymmetrical mate preference in this system; White Sands males preferentially courted local females, while males from dark soils and black lava populations did not exhibit a preference for local mates. We also found that female behavior and ventral patch phenotype were associated with male courtship. Our results suggest that the observed preference for local mates evolved at White Sands, and we discuss the possible link between local adaptation and traits involved in mate preference in this system.

Absence of Climbing-Induced Limb Length Plasticity in the Eastern Fence Lizard,Sceloporus undulatus

Abstract Understanding the processes shaping responses of native species to environmental perturbations, such as the introduction of non-native species, will allow us to better manage such impacts. The imported Red Fire Ant, Solenopsis invicta, invaded the United States ∼70 years ago and co-occurs with Fence Lizards (Sceloporus undulatus) in portions of their range. Lizards from fire ant-invaded populations have longer hind limbs which more effectively remove ants and increase survival following fire ant attack. Prior research on this system suggests that this difference in relative hind limb length (RHL) is fixed at birth; however, RHL is a plastic trait in other lizard species and could help Fence Lizards survive. Lizards from invaded sites are more likely to climb when attacked by fire ants or when stressed (i.e., have elevated levels of corticosterone) than are lizards from uninvaded sites. We tested for climbing-induced plasticity in RHL by manipulating whether or not Fence Lizards had to climb raise...

Colonization of novel White Sands habitat is associated with changes in lizard anti-predator behaviour

Colonization of novel habitats is often associated with differences in ecological community composition. For small diurnal animals, differences in predator diversity and abundance can lead to behavioural shifts in the novel habitat. The eastern fence lizard Sceloporus undulatus (Bosc and Daudin, 1801) recently colonized the gypsum dunes of White Sands, a predator-poor community relative to the predator-rich community of the surrounding Chihuahuan dark-soil habitat. We used field experiments to assess S. undulatus anti-predator behaviour in white-sand versus dark-soil habitats, and used laboratory assays to determine whether behavioural differences could be mediated by hormonal regulation. Overall, we found that white-sand lizards were less vigilant but more wary than their dark-soil counterparts; it took them longer to detect a simulated predator, but once detected they were more likely to retreat from their perches than dark-soil lizards. At the proximate level, differences in anti-predator behaviour could not be explained by differences in plasma hormone levels (corticosterone and testosterone); we detected elevated corticosterone for lizards in our stress treatment relative to control treatment, but found no differences between habitats in baseline or acute corticosterone levels. At the evolutionary level, we suggest that differences in anti-predator behaviour may be explained by differences across habitats in predation environment, habituation, and/or the cost of retreating. Our study implicates changes in predator community composition in mediating ecological divergence in behaviour.

Male territoriality and ‘sex confusion’ in recently adapted lizards at White Sands

The evolution of intersexual interactions, like mate choice, during ecological speciation has received widespread attention. However, changes in intrasexual interactions, like male territoriality, during ecological divergence are largely unexamined. We conducted field experiments with adaptively diverged populations of the eastern fence lizard (Sceloporus undulatus) to determine whether territorial males behaved differently towards ecologically similar vs. dissimilar intruders. We performed trials with light-coloured males from White Sands, New Mexico and dark-coloured males from the surrounding desert. We found that intruders from White Sands elicited more aggression than intruders from dark-soil habitat. We also documented a case of 'sex confusion' where white-sand males courted dark-soil intruders. We found population differences in signalling patch size that can explain both aggression bias and sex misidentification. We argue that direct selection (for population recognition or optimal signal transmission) and indirect selection (by-products of ecological adaptation) should influence both intersexual and intrasexual interactions during ecological speciation.

Habitat relationships of reptiles in pine beetle disturbed forests of Alabama, U.S.A. with guidelines for a modified drift-fence sampling method

Abstract Understanding vertebrate habitat relationships is important to promote management strategies for the longterm conservation of many species. Using a modified drift fence method, we sampled reptiles and compared habitat variables within the William B. Bankhead National Forest (BNF) in Alabama, U.S.A from April 2005 to June 2006. We captured 226 individual reptiles representing 19 species during 564 total trap nights. We used canonical correspondence analysis to examine habitat associations for the reptiles sampled and we detected a distinct habitat gradient ranging from sites with greater litter depth and percent canopy cover to more open sites with greater woody, herbaceous, and coarse woody debris (CWD) coverage, and CWD volume. Little brown skinks Scincella lateralis and eastern worm snakes Carphophis a. amoenus were associated with sites with greater litter depth and canopy cover, whereas eastern fence lizards Sceloporus undulatus, copperheads Agkistrodon contortrix, and gray ratsnakes Pantherophis spiloides were associated with sites possessing greater CWD coverage and volume. We found that disturbances due to the southern pine beetle Dendroctonus frontalis were likely important for influencing reptile distributions through the creation of canopy gaps and fallen coarse woody debris. Compared to other studies, our modified drift-fence trap technique was successful for sampling larger snake species (66 snakes in 564 trap nights). We have also provided detailed schematics for constructing drift fence array and box traps used in this study.

Fluctuations in the incubation moisture environment affect growth but not survival of hatchling lizards

Few studies have collected longitudinal data that follow the complete microevolutionary path of an organism linking sources of variation (e.g. environmental versus genetic) to a trait and its subsequent relationship with fitness. Identifying the links within this pathway is imperative for understanding the ecological relevance of effects found at the phenotypic level. Furthermore, experimental studies that examine parts of the pathway in ectothermic organisms often fail to mimic the complexities of the natural developmental environment. Temperature and moisture conditions in reptile nests, for example, can fluctuate greatly on a seasonal and daily basis. Despite the potential effects of fluctuating environments, the vast majority of studies have held environmental treatments constant during the developmental period. We investigated the effects of fluctuating moisture regimes during incubation on eggs, hatchling phenotypes, and subsequent survival in the eastern fence lizard Sceloporus undulatus. Moisture fluctuations during embryonic development caused water absorption by eggs to follow the environmental availability of moisture. Initial hatchling tail length was affected by the pattern of moisture fluctuations, and hatchling growth rates in fluctuating treatments were significantly faster than those in a constant treatment, resulting in larger hatchlings after 4 weeks. A release–recapture experiment conducted in the field did not detect a treatment effect on survival despite the larger body sizes. In summary, although fluctuations affected water absorption by eggs and some hatchling traits, these effects did not have subsequent fitness consequences. The results obtained suggest that egg and hatchling survival are buffered against natural soil moisture fluctuations during incubation, even when egg and hatchling traits are significantly affected. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100, 89–102.

The importance of energetic versus pelvic constraints on reproductive allocation by the eastern fence lizard (Sceloporus undulatus)

In ectothermic species, females often produce larger eggs in colder environments. Models based on energetic con- straints suggest that this pattern is an adaptation to compensate for the slower growth of offspring in the cold. Yet, females in cold environments also tend to be larger than females in warm environments. Consequently, thermal clines in egg size could be caused by pelvic constraints, which stem from the inability of large eggs to pass through a small pelvic aperture. Models based on energetic constraints and models based on pelvic constraints predict similar relationships between maternal size and egg size. However, pelvic constraints should produce these relationships both within and among populations, whereas energetic constraints would not necessarily do so. If pelvic constraints are important, we might also expect small females to compensate by producing eggs that are relatively rich in lipids (i.e. high energy density). The present study aimed to assess whether energetic or pelvic constraints generate geo- graphical variation in egg size of the lizard Sceloporus undulatus . Pelvic width is very highly correlated with body length in S. undulatus , making maternal size a suitable measure of pelvic constraint. Although maternal size and egg mass (dry and wet) covaried among populations, these variables were generally not related within populations. Energetic density of eggs tended to increase with decreasing egg mass (dry and wet), but this relationship was stron- gest in populations where no relationship between maternal size and egg mass was observed. Our results do not sup- port the pelvic constraint model and thus indicate energetic constraints play a greater role in generating geographical variation in egg size. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 513-521.

Incubation temperature and phenotypic traits of Sceloporus undulatus: implications for the northern limits of distribution

Cold environmental temperature is detrimental to reproduction by oviparous squamate reptiles by prolonging incubation period, negatively affecting embryonic developmental processes, and by killing embryos in eggs directly. Because low soil temperature may prevent successful development of embryos in eggs in nests, the geographic distributions of oviparous species may be influenced by the thermal requirements of embryos. In the present study, we tested the hypothesis that low incubation temperature determines the northern distributional limit of the oviparous lizard Sceloporus undulatus. To compare the effects of incubation temperature on incubation length, egg and hatchling survival, and hatchling phenotypic traits, we incubated eggs of S. undulatus under temperature treatments that simulated the thermal environment that eggs would experience if located in nests within their geographic range at 37 degrees N and north of the species' present geographic range at latitudes of 44 and 42 degrees N. After hatching, snout-vent length (SVL), mass, tail length, body condition (SVL relative to mass), locomotor performance, and growth rate were measured for each hatchling. Hatchlings were released at a field site to evaluate growth and survival under natural conditions. Incubation at temperatures simulating those of nests at 44 degrees N prolonged incubation and resulted in hatchlings with shorter SVL relative to mass, shorter tails, shorter hind limb span, slower growth, and lower survival than hatchlings from eggs incubated at temperatures simulating those of nests at 37 and 42 degrees N. We also evaluated the association between environmental temperature and the northern distribution of S. undulatus. We predicted that the northernmost distributional limit of S. undulatus would be associated with locations that provide the minimum heat sum (approximately 495 degree-days) required to complete embryonic development. Based on air and soil temperatures, the predicted northern latitudinal limit of S. undulatus would lie at approximately 40.5-41.5 degrees N. Our predicted value closely corresponds to the observed latitudinal limit in the eastern United States of approximately 40 degrees N. Our results suggest that soil temperatures at northern latitudes are not warm enough for a sufficient length of time to permit successful incubation of S. undulatus embryos. These results are consistent with the hypothesis that incubation temperature is an important factor limiting the geographic distributions of oviparous reptile species at high latitudes and elevations.

Seasonal effects on circulating leptin in the lizard Sceloporus undulatus from two populations

Characterizing leptin's structure and function in mammals has been the subject of thousands of studies since 1994. Recently, the study of leptin has expanded to include its distribution in non-mammalian taxa, and the role that leptin plays in the reproductive axis. We demonstrated in a previous study that Sceloporus undulatus, fence lizards (ectotherms), express a leptin-like protein. In the current study we quantified seasonal variation in this putative leptin among free-ranging fence lizards from two populations characterized by early and late reproductive maturation (after one or two years, respectively). Immunoblots were performed on whole blood samples to detect leptin and estimate its titer. Leptin titers were higher in the reproductive population of S. undulatus (early maturing: 2.5 ± 0.2 μg/mL; late-maturing 2.2 ± 0.3 μg/mL; mean ± 2 S.E.), but both populations showed the same seasonal pattern. Leptin titers were lowest in fall when fat stores are expected to be highest (spring: 2.6 ± 0.3 μg/mL; summer: 2.6 ± 0.3; μg/mL; fall: 1.8 ± 0.3 μg/mL), consistent with findings of seasonal variation in free-ranging mammals. Our data support previous work asserting that lizards express leptin and that it has a similar physiological function in endotherms and ectotherms. Our long-term goal is to use leptin to manipulate age at maturity and to test fundamental questions in the evolution of life-history strategies.

CONVERGENT EVOLUTION OF EMBRYONIC GROWTH AND DEVELOPMENT IN THE EASTERN FENCE LIZARD (SCELOPORUS UNDULATUS)

Theory predicts that cold environments will select for strategies that enhance the growth of ectotherms, such as early emergence from nests and more efficient use of resources. We used a common garden experiment to detect parallel clines in rates of embryonic growth and development by eastern fence lizards (Sceloporus undulatus). Using realistic thermal conditions, we measured growth efficiencies and incubation periods of lizards from five populations representing two distinct clades. In both clades, embryos from cold environments (Indiana, New Jersey, and Virginia) grew more efficiently and hatched earlier than embryos from warm environments (Florida and South Carolina). Because eggs from cold environments were larger than eggs from warm environments, we experimentally miniaturized eggs from one population (Virginia) to determine whether rapid growth and development were caused by a greater maternal investment. Embryos in miniaturized eggs grew as efficiently and incubated for the same duration as embryos in unmanipulated eggs. Taken together, our results suggest countergradient variation has evolved at least twice in S. undulatus.

Testosterone has opposite effects on male growth in lizards(Sceloporusspp.) with opposite patterns of sexual size dimorphism

Sexual size dimorphism (SSD) has received considerable attention from evolutionary biologists, but relatively little is known about the physiological mechanisms underlying sex differences in growth that lead to SSD. Testosterone (T) stimulates growth in many male-larger vertebrates, but inhibits growth in the female-larger lizard Sceloporus undulatus. Thus, opposite patterns of SSD may develop in part because of underlying differences in the hormonal regulation of male growth. In the present study, we examined the effects of T on male growth in two sympatric congeners with opposite patterns of SSD (S. virgatus: female-larger; S. jarrovii: male-larger). During the mating season, yearling males of both species have higher plasma T levels than females, but whereas yearling males of S. virgatus grow only half as fast as females, yearling males of S. jarrovii grow more quickly than females. Thus, we hypothesized that T inhibits growth in yearling S. virgatus males, but promotes growth in yearling S. jarrovii males. In support of this hypothesis, we found that castrated (CAST) males of S. virgatus grew faster than castrated males given T implants (TEST). In contrast, TEST males of S. jarrovii grew faster than CAST males. Our results provide the first direct evidence for opposite effects of T on male growth in closely related species with opposite patterns of SSD. We speculate that growth inhibition by T reflects an energetic trade-off between growth and reproductive investment, and propose that such ;costs' of male reproduction may help explain the evolution of female-larger SSD in Sceloporus.

Embryonic responses to variation in oviductal oxygen in the lizard Sceloporus undulatus from New Jersey and South Carolina, USA

Viviparity in reptiles is hypothesized to evolve in cold climates at high latitudes and high elevations through selection for progressively longer periods of egg retention. Oxygen consumption of embryos increases during development and therefore longer periods of egg retention should be associated with maternal or embryonic features that enhance embryonic oxygen availability. We tested the hypotheses that embryos of the oviparous lizard Sceloporus undulatus from a high-latitude population in New Jersey are oviposited at more advanced developmental stages and have a higher growth rate at low oxygen partial pressures (pO2) than embryos from a low-latitude population in South Carolina. These hypotheses were rejected; embryos from the two populations did not differ in embryonic stage at oviposition, survival, rate of differentiation or growth in mass when incubated under simulated in utero conditions at low oxygen concentrations. We also estimated the effective pO2 experienced by lizard embryos in utero. At an effective pO2 of 8.6 kPa (9% O2), development of S. undulatus embryos is arrested at Dufaure and Hubert stage 30 and at a dry mass of 0.8 mg. Physiological and morphological features of gravid females, embryos, or both, that facilitate oxygen uptake for developing embryos appear to be a critical early step during the evolution of reptilian viviparity. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83, 289–299.

Patterns of body size and habitat use in a lizard assemblage

:We quantified differences in the distribution and behaviours of three territorial species of phrynosomatine lizard (Urosaurus ornatus, Uta stansburiana, and Sceloporus undulatus) between two habitats on the elevated mesas of western Colorado. Community lizard density was greater on rock ridges dissecting pinyon-juniper woodland than in the surrounding flat, open areas. The frequency of community-level interactions, or behaviours directed towards another lizard of any species, was also greater in the rock compared to the flatland habitat for S. undulatus. Snout-vent length frequency distributions of all three species differed between habitats, with larger lizards residing in the rock habitat. For each species, the distribution of hatchling lizards between habitats was opposite to that expected based on the distribution of conspecific adult females, with the majority of adult females residing in the rock habitat and the majority of hatchlings in the flatland. Indices of juvenile recruitment (number of hatchlings per adult female) in the flatland habitat approximated average clutch size for U. ornatus and U. stansburiana despite known high rates of juvenile mortality in these species. We suggest that ontogenetic habitat shifts may explain the observed age and size distributions of species between habitats. Hatchlings may disperse from rock to flatland to avoid aggression and predation from adult lizards in the rock habitat and return when they have attained larger body size and thus face reduced predation risk, being presumably more able to compete within social hierarchies.

PHYLOGENETIC COMPARATIVE ANALYSIS OF LIFE-HISTORY VARIATION AMONG POPULATIONS OF THE LIZARD SCELOPORUS UNDULATUS: AN EXAMPLE AND PROGNOSIS

Over the past 15 years, phylogenetic comparative methods (PCMs) have become standard in the study of life-history evolution. To date, most studies have focused on variation among species or higher taxonomic levels, generally revealing the presence of significant phylogenetic effects as well as residual variation potentially attributable to adaptive evolution. Recently, population-level phylogenetic hypotheses have become available for many species, making it possible to apply PCMs directly to the level at which experiments are typically used to test adaptive hypotheses. In this study, we present the results of PCMs applied to life-history variation among populations of the widespread and well-studied lizard Sceloporus undulatus. Using S. undulatus (which may represent four closely related species) as an example, we explore the benefits of using PCMs at the population level, as well as consider the importance of several thorny methodological problems including but not limited to nonindependence of populations, lack of sufficient variation in traits, and the typically small sample sizes dictated by the difficulty of collecting detailed demographic data. We show that phylogenetic effects on life-history variation among populations of S. undulatus appear to be unimportant, and that several classic trade-offs expected by theory and revealed by many interspecific comparisons are absent. Our results suggest that PCMs applied to variation in life-history traits below the species level may be of limited value, but more studies like ours are needed to draw a general conclusion. Finally, we discuss several outstanding problems that face studies seeking to apply PCMs below the species level.

Consequences of Extended Egg Retention in the Eastern Fence Lizard (Sceloporus undulatus)

Egg retention beyond the normal time of oviposition occurs frequently in oviparous squamate reptiles and is thought to be a response to unfavorable nesting conditions. During studies of the Eastern Fence Lizard (Sceloporus undulatus), we obtained data on the effects of extended egg retention on embryonic development, hatchling phenotypes, and posthatching survival under natural field conditions. Females that retained eggs beyond the normal time of oviposition produced heavier eggs with embryos more advanced (by one stage unit) at the time of oviposition than females that did not retain eggs for extended periods. Egg retention did not affect any hatchling phenotype (i.e., body size, thermal preference, running speed, desiccation rate, growth rate) but had a significant positive effect on posthatching survival in the field. However, the mechanism by which extended egg retention affects posthatching survival remains unclear. Our results have implications for the evolution of viviparity, but carefully designed experiments are needed to further understand the causes and consequences of extended egg retention.

Laboratory and field experiments identify sources of variation in phenotypes and survival of hatchling lizards

To understand the process of natural selection, relationships between phenotype and fitness and sources of phenotypic variation must be known. We examined the importance of incubation moisture conditions, maternal yolk investment, and clutch (genotype) to phenotypic variation in hatchlings of the lizard Sceloporus undulatus. Eggs were distributed among two moisture treatments and a third treatment in which yolk was removed. After hatching, mass, snout–vent length, tail length, body shape, thermal preference, running speed, desiccation rate, and growth rate were measured for each hatchling in the laboratory. Hatchlings were then released at a field site in order to monitor growth and survival under natural conditions. Hatchlings from the dry and yolk-removed treatments were significantly smaller than those from the wet treatment. However, neither performance nor survival were affected by moisture or yolk removal. All phenotypes were affected by clutch. Clutches that produced relatively large hatchlings had higher survival than clutches that produced relatively small hatchlings. Furthermore, clutches that produced relatively slow growing individuals and fast runners had higher survival rates than clutches that produced relatively rapid growing individuals and slow runners. Our results emphasize the overriding importance of clutch (genotype) to variation in phenotypes and survival in hatchling S. undulatus.