Boldness,Dominance, and Territoriality in the Color Polymorphic Tree Lizard,Urosaurus ornatus

Social dynamics in territorial species often reflect underlying variation in aggression and other aspects of social dominance among individuals. In ornate tree lizards (Urosaurus ornatus), males differing in dewlap color differ in social dominance: while blue males are the dominant, aggressive morph and always territorial, yellow males tend to exhibit a less‐aggressive satellite behavioral tactic. However, in habitats with fewer available territorial resources, yellow males defend territories and increase in relative abundance. These observations suggest that consideration of social dominance alone may be insufficient to explain U. ornatus' territorial dynamics in the wild. Here, we tested how both dominance and another important behavioral trait, boldness, contribute to the outcome of territorial disputes in tree lizards. We recorded the territorial behavior of blue and yellow male tree lizards (entered in pairs) in an experimental arena. At the end of each trial, we then approached each male and recorded whether it fled (shy) or not (bold) in response to our approach. As expected, dominant blue males exploited the higher quality perch more often than yellow males. However, when approached by a simulated predator, blue males were more likely to flee than yellow males. Thus, while blue males are more dominant, yellow is likely the bolder morph. As a result, this morph may be better equipped to defend territories in riskier environments. We conclude that although dominance asymmetries may predictably drive initial territorial interactions among competing males, variation in other behaviors (like boldness) may perturb the long‐term outcome of these interactions across variable environments.     

Do Displays Send Information about Ornament Structure and Male Quality in the Ornate Tree Lizard,Urosaurus ornatus?

Displays can transmit information about ornament or male quality; however, few studies have simultaneously explored the relationship between displays, ornament and male quality within a single species. We quantified ornament morphology (five throat color morphs, throat area, and belly area), male quality (bite force, sprint speed, body condition, and body mass), display behavior [percent time displaying (PTD), number of pushups per display, and display duration], and movement behavior among males in a population of the ornate tree lizard, Urosaurus ornatus. Previous studies have shown that male U. ornatus are polymorphic in throat coloration and that morphs differ in behavioral aggression. Our study shows that blue throat morphs use 1.5 more pushups per display than other male color morphs, which suggests that throat color and display behavior act as backup signals for aggression. However, other data support the multiple messages hypothesis, and overall our data do not provide conclusive evidence for any one hypothesis. In addition, we show that body mass is positively related to both PTD and percent time moving and this relationship is independent of color morph. We also found that throat area, belly area, bite force, sprint speed, and body condition are unrelated to display behavior. This result highlights at least some discordance between display behavior, ornaments, and performance in U. ornatus and suggests that these traits may be evolving independently.     

Determinants of embryonic stage at oviposition in the lizard Urosaurus ornatus

Relatively few squamate reptiles oviposit eggs with embryos at developmental stages greater than stage 30. To investigate potential proximate and ultimate bases of this phenomenon, we experimentally induced females of the lizard Urosaurus ornatus to retain their eggs past the normal time of oviposition (NTO). This procedure allowed us to determine whether the length of egg retention is fixed or facultative and to evaluate the effects of retention on embryos, hatchlings, and females. Females were able to retain eggs facultatively for at least 29 d past the NTO. However, retention resulted in arrested development of embryos; arrest occurred at stages 30-30.5, which is only slightly more advanced than that at the NTO (stage 29.5). Embryogenesis was reinitiated when eggs were removed from females and placed in incubation media. Hatching success of these eggs was high (87%), and incubation time was not affected by the number of days that development had been arrested. However, the snout-vent length and water content of hatchlings were negatively related to the length of retention, and they ran slower than hatchlings from control eggs obtained at the NTO. Retention of eggs past the NTO had no detectable effect on the body condition or running speeds of females. Developmental arrest and the adverse effects of retention on hatchling phenotype, if widespread among squamates, would account for the limited range of embryo stages at oviposition and act as major constraints on the evolution of viviparity.     

Preference for Male Traits Differ in Two Female Morphs of the Tree Lizard,Urosaurus ornatus

Non-random female mating preferences may contribute to the maintenance of phenotypic variation in color polymorphic species. However, the effect of female preference depends on the types of male traits used as signals by receptive females. If preference signals derive from discrete male traits (i.e., morph-specific), female preferences may rapidly fix to a morph. However, female preference signals may also include condition-dependent male traits. In this scenario, female preference may differ depending on the social context (i.e., male morph availability). Male tree lizards (Urosaurus ornatus) exhibit a dewlap color polymorphism that covaries with mating behavior. Blue morph males are aggressive and defend territories, yellow males are less aggressive and defend smaller territories, and orange males are typically nomadic. Female U. ornatus are also polymorphic in dewlap color, but the covariation between dewlap color and female behavior is unknown. We performed an experiment to determine how female mate choice depends on the visual and chemical signals produced by males. We also tested whether female morphs differ in their preferences for these signals. Female preferences involved both male dewlap color and size of the ventral color patch. However, the female morphs responded to these signals differently and depended on the choice between the types of male morphs. Our experiment revealed that females may be capable of distinguishing among the male morphs using chemical signals alone. Yellow females exhibit preferences based on both chemical and visual signals, which may be a strategy to avoid ultra-dominant males. In contrast, orange females may prefer dominant males. We conclude that female U. ornatus morphs differ in mating behavior. Our findings also provide evidence for a chemical polymorphism among male lizards in femoral pore secretions.     

Isolation and characterization of microsatellite markers in tree lizards (Urosaurus ornatus)

Tree lizards (Urosaurus ornatus) are a highly accessible species useful in testing theories of life-history evolution and behavioural ecology. A polymerase chain reaction-based method was used to isolate nine polymorphic tetranucleotide microsatellite loci from the genome of these lizards. The isolated loci displayed 5 to 9 alleles in the populations screened. Observed heterozygosity ranged from 0.333 to 0.900. Results indicate these loci now provide a basis to study mate choice, parentage, and population genetic structure within this species.     

Phylogeography of the tree lizard, Urosaurus ornatus: responses of populations to past climate change

Isolation due to both geological barriers and range contractions during the Pleistocene glacial maxima has been an important cause of diversification of arid-adapted species in the North American deserts. Tree lizards, Urosaurus ornatus, are distributed across much of the southwestern arid regions and can tolerate a wide range of environments. Thus, they may have avoided large-scale shifts in distribution caused by Pleistocene climate change and any subsequent evolutionary impacts. Cytochrome b sequences were sampled from U. ornatus across the northern part of their range to test if current structure of these populations resulted from post-Pleistocene range expansion and habitat fragmentation, or prior geological isolation. Phylogenetic analyses found geographical structuring of populations consistent with a model of long-term geographical isolation corresponding to each of the desert regions. The two post-Pleistocene hypotheses were not well supported as estimated times of divergence predated the retreat of the last continental ice sheet. Populations in different regions were impacted by different processes. Southern populations of U. ornatus appear to have remained largely independent of more derived northern and eastern populations during Pleistocene climate change, while populations in regions containing more derived populations showed evidence of more recent range expansion (Colorado Plateau). As populations of U. ornatus attest to, the complex and dynamic history of the southwestern USA has left a deep-rooted and multifaceted imprint on genetic and phylogeographical structure of the species living there.     

Phylogeography and Population Genetic Structure of the Ornate Dragon Lizard, Ctenophorus ornatus

Species inhabiting ancient, geologically stable landscapes that have been impacted by agriculture and urbanisation are expected to have complex patterns of genetic subdivision due to the influence of both historical and contemporary gene flow. Here, we investigate genetic differences among populations of the granite outcrop-dwelling lizard Ctenophorus ornatus, a phenotypically variable species with a wide geographical distribution across the south-west of Western Australia. Phylogenetic analysis of mitochondrial DNA sequence data revealed two distinct evolutionary lineages that have been isolated for more than four million years within the C. ornatus complex. This evolutionary split is associated with a change in dorsal colouration of the lizards from deep brown or black to reddish-pink. In addition, analysis of microsatellite data revealed high levels of genetic structuring within each lineage, as well as strong isolation by distance at multiple spatial scales. Among the 50 outcrop populations’ analysed, non-hierarchical Bayesian clustering analysis revealed the presence of 23 distinct genetic groups, with outcrop populations less than 4 km apart usually forming a single genetic group. When a hierarchical analysis was carried out, almost every outcrop was assigned to a different genetic group. Our results show there are multiple levels of genetic structuring in C. ornatus, reflecting the influence of both historical and contemporary evolutionary processes. They also highlight the need to recognise the presence of two evolutionarily distinct lineages when making conservation management decisions on this species.     

Steroid hormone mediation of limbic brain plasticity and aggression in free-living tree lizards, Urosaurus ornatus

The neural mechanisms by which steroid hormones regulate aggression are unclear. Although testosterone and its metabolites are involved in both the regulation of aggression and the maintenance of neural morphology, it is unknown whether these changes are functionally related. We addressed the hypothesis that parallel changes in steroid levels and brain volumes are involved in the regulation of adult aggression. We examined the relationships between seasonal hormone changes, aggressive behavior, and the volumes of limbic brain regions in free-living male and female tree lizards (Urosaurus ornatus). The brain nuclei that we examined included the lateral septum (LS), preoptic area (POA), amygdala (AMY), and ventromedial hypothalamus (VMH). We showed that the volumes of the POA and AMY in males and the POA in females vary with season. However, reproductive state (and thus hormonal state) was incompletely predictive of these seasonal changes in males and completely unrelated to changes in females. We also detected male-biased dimorphisms in volume of the POA, AMY, and a dorsolateral subnucleus of the VMH but did not detect a dimorphism between alternate male morphological phenotypes. Finally, we showed that circulating testosterone levels were higher in males exhibiting higher frequency and intensity of aggressive display to a conspecific, though brain nucleus volumes were unrelated to behavior. Our findings fail to support our hypothesis and suggest instead that plasma testosterone level covaries with aggression level and in a limited capacity with brain nucleus volumes but that these are largely unrelated relationships.     

Eimeria urosauris n. sp., a Coccidium from the Lizard Urosaurus graciosus Hollowell, in California

SYNOPSIS. A new species of coccidium is described: Eimeria urosauris n. sp., in the gall bladder of the lizard Urosaurus graciosus Hollowell, from the Mojave Desert in California. The oocyst of E. urosauris is smooth, bilaminar, nearly cylindrical, with long borders only slightly convex, ends rounded and very nearly hemispherical. It is usually 32 × 20 μ, and its length/width ratio is 1.6. It contains 4 ellipsoid sporocysts, each 10.5 × 9 μ, for which 1/w is 1.17. Each sporocyst contains 2 tapered bent sporozoites with rounded ends, 11 μ long, 4 μ in diameter at the larger end, and 1.5 μ in diameter at the smaller end. Each sporocyst also contains a central granular sporocyst residuum 3.5 μ in diameter. The oocyst lacks a micropyle and oocyst residuum, and there is no Stieda body on the sporocyst. Sporulation time is 6–10 hr. Endogenous development, with reinfection by liberated sporozoites, occurs in the epithelial lining of the gall bladder. E. urosauris is compared to other morphologically similar lacertilian eimerias with which it might be confused.     

Manipulation of egg production reveals costs of reproduction in the tree lizard (Urosaurus ornatus)

Life-history theory predicts that the allocation of energy to current reproduction is associated with a decrement in future fecundity, future survival, or both. I treated this notion as the cost hypothesis, and tested the assumption that current reproduction exacts a cost in future survival and fecundity. Surgical manipulations of egg production were applied to natural populations of the tree lizard, Urosaurus ornatus, in south western New Mexico by yolkectomy surgery in two different years. I reduced the number of eggs produced in the first clutch during vitellogenesis by approximately 50% in yolkectomized females relative to controls. Subsequent survival, fecundity, and growth of females were followed for two or three years, depending on the cohort. Treated females in both cohorts showed significantly higher growth and survivorship throughout the experiment than in controls. After 2 years, yolkectomized females had grown an additional 2 mm (snout-vent length) compared to controls, enough for them to add on average an additional egg to their next clutch. This demonstrated a cost in terms of future fecundity through a reduction in growth and an increase in mortality in these lizards.     

Juxtaposition and Disturbance: Disentangling the Determinants of Lizard Community Structure

Disturbance alters the structure and dynamics of communities. Here, we examined the effects of seasonal flooding on the lizard community structure by comparing two adjacent habitats, a seasonally flooded and a non‐flooded forest, in a Cerrado–Amazon ecotone area, the Cantão State Park, Tocantins state, Brazil. Despite the strong potential impact of seasonal flooding, the only significant environmental difference detected was more termite mounds in non‐flooded forests. Species richness was significantly higher in the non‐flooded forest. Colobosaura modesta, followed by Mabuya frenata and Anolis brasiliensis, were the only species that differed in number of captures between sites. Colobosaura modesta was exclusively found in the non‐flooded forest, while Anolis brasiliensis was the most captured in the flooded forest. Mabuya frenata is indicated as an indicator species in the flooded forest, and Colobosaura modesta in the non‐flooded forest. We found a significant association between lizard abundances and habitat characteristics, with flooding, canopy cover, and logs being the best predictors. A phylogenetic community structure analysis indicated a lack of structure in both lizard assemblages. Overall, we show that seasonal flooding can strongly impact species richness and species occurrence patterns, but not phylogenetic community structure. The Amazon–Cerrado transition is undergoing pronounced transformations due to deforestation and climate change. Despite being species‐poor compared with central areas in Amazon or Cerrado, this ecotone harbors species with important adaptations that could hold the key to persistence in human‐disturbed landscapes or during periods of climate change.     

Dominance, Status Signals and Coloration in Male Mandrills (Mandrillus sphinx)

Where individuals contest access to a resource, escalated physical fighting presents a risk to all involved. The requirement for mechanisms of conflict management has led to the evolution of a variety of decision rules and signals that act to reduce the frequency of aggression during competitive encounters. We examined strategies of conflict management in male mandrills (Mandrillus sphinx) living in two semi‐free‐ranging groups in Gabon. Adult male mandrills are large (31 kg), with long canines, making the costs of conflict potentially very high. We found that males formed dominance hierarchies, but that male–male relationships were characterized by avoidance, appeasement and ignoring. Fights were rare, but could result in death. Examination of the relationship between dominance and signaling showed that males use facial and gestural signals to communicate dominance and subordinance, avoiding escalated conflict. Male mandrills also possess rank‐dependent red coloration on the face, rump and genitalia, and we examined the hypothesis that this coloration acts as a ‘badge of status’, communicating male fighting ability to other males. If this is the case, then similarity in color should lead to higher dyadic rates of aggression, while males that differ markedly should resolve encounters quickly, with the paler individual retreating. Indeed, appeasement (the ‘grin’ display), threats, fights and tense ‘stand‐off’ encounters were significantly more frequent between similarly colored males, while clear submission was more frequent where color differences were large. We conclude that male mandrills employ both formal behavioral indicators of dominance and of subordination, and may also use relative brightness of red coloration to facilitate the assessment of individual differences in fighting ability, thereby regulating the degree of costly, escalated conflict between well‐armed males.     

Correlations between habitat use and body shape in a phrynosomatid lizard (Urosaurus ornatus): a population-level analysis

Recent ecomorphological studies have shown that the predicted correlations between morphology and ecology on broad taxonomic levels are often obscured when comparing more closely related groups. Among species, comparisons of lizards often indicate very little support for adaptive radiations into novel habitats. As few population level studies have been performed, we compared body, head and limb shape between four populations of Urosaurus ornatus living in structurally distinct habitats (cliffs, rocks, trees and boulders). Surprisingly, clear correlations between habitat use and body shape among populations were found, most of which were in good accordance with a priori biomechanical predictions (e.g. flat body and head for extreme climbers; long distal hindlimb segments for jumpers and runners; narrow body and long tail for tree dwelling lizards). This indicates that populations of Urosaurus ornatus are seemingly 'adapted' to the habitat they live in. However, quantification of performance and behaviour are needed to determine the adaptive nature of these observations.     

Male mate choice and the potential for complex mating dynamics in the tree lizard (Urosaurus ornatus)

A growing body of literature is recognizing that males may also play a role in the mating process by behaving non‐randomly toward potential female mates during courtship. In numerous species, discrete color polymorphisms in males are inferred to represent alternative mating tactics, which often correspond with concomitant asymmetries in ecology and behavior. In terms of their mating behavior, these ecological outcomes of a color polymorphism should affect a morph's likelihood and frequency of encountering females in a population, possibly favoring the evolution of morph‐specific mating preferences. Knowledge of how male morphs contribute to a species’ overall mating dynamics will improve our understanding of how sexual selection shapes phenotypic diversity in color polymorphic systems. We conducted a mate choice experiment to evaluate the extent and morph specificity of non‐random mating preferences by male ornate tree lizards, Urosaurus ornatus. We observed the behavior of blue and yellow males in an experimental arena in response to a choice between an orange or yellow female. We found that blue males preferred yellow females over orange females, and although yellow males visited females more often than blue males overall, their attention was not morph‐specific. Given male morph differences in choosiness, and their differences in social dominance, we conclude that female throat color may be partly under sexual selection in U. ornatus. However, a lack of concordance between male and female mating preferences (drawn from an earlier study) suggests that overall mating dynamics may serve to maintain, rather than enhance, color morph differences in this species.     

Aggression frequency and intensity, independent of testosterone levels, relate to neural activation within the dorsolateral subdivision of the ventromedial hypothalamus in the tree lizard Urosaurus ornatus

The mechanisms by which testosterone regulates aggression are unclear and may involve changes that alter the activity levels of one or more brain nuclei. We estimate neural activity by counting immunopositive cells against phosphorylated cyclic AMP response element binding protein (pCREB). We demonstrate increased pCREB immunoreactivity within the dorsolateral subdivision of the ventromedial hypothalamus (VMHdl) following an aggressive encounter in male tree lizards Urosaurus ornatus. This immunoreactivity is induced both by exposure to and performance of aggressive behaviors. This dual activation of the VMHdl suggests its possible role as an integration center for assessment and expression of aggressive behavior. Furthermore, pCREB induction was greater in encounters involving higher frequency and intensity of aggressive display, demonstrating a direct relationship between neural activation and behavior. The VMHdl is also rich in steroid receptors. In a second experiment involving hormone manipulations, testosterone treatment increased aggression levels, though it did not increase the number of pCREB positive cells within the VMHdl. This lack of an effect of testosterone on pCREB induction within the VMHdl may be due to induction arising from the behaviors of conspecifics (especially in low-testosterone, low-aggression individuals), variation in aggression mediated by other variables, or regulation of aggression by circuits outside of the VMHdl. Together, these findings support a notion of the VMHdl as a nucleus involved in integrating afferent and efferent information within the neural aggression-control circuit.     

The Relative Importance of Fungal Infection,Conspecific Density and Environmental Heterogeneity for Seedling Survival in a Dominant Tropical Tree

A gap remains in our understanding of how host‐specific fungal pathogens impact negative density dependence (NDD). Here, we investigated survival of Cinnamomum subavenium Miq. seedlings, the dominant canopy species in a seasonal tropical evergreen forest, Thailand. It is infected by a host‐specific fungus that is easily identifiable in the field. We quantified the effects of conspecific seedling and adult density on fungal infection and seedling survival over a wide range of environmental heterogeneity in elevation, understory vegetation and presence of forest gaps. Generalized linear mixed models (GLMMs) for seedling survival revealed that fungal infection significantly reduced survival and had the strongest effect on seedling survival as compared with conspecific density and environmental heterogeneity. Adult conspecific density was not, however, significantly correlated with the probability of infection, and conspecific seedling density was positively associated with increased infection only at high elevations. In contrast to infection, we found a significant positive correlation between conspecific seedling density and the probability of seedling survival. Consequently, our results demonstrate that fungal infection can have major impacts on seedling survival, but not in a manner consistent with local NDD effects on seedlings, as assumed in the Janzen–Connell hypothesis. Our study provides an example of how quantifying the interaction between environmental heterogeneity and a host‐specific plant‐pathogen can yield unexpected insights into the dynamics of seedling populations. The combined effects of host‐specific pathogens and environmental heterogeneity on survival of dominant seedling species may ultimately provide a chance for rarer species to recruit.     

Effects of Prior Residence and Previous Cohabitation on the Polistes dominulus Christ Dominance Hierarchy

We investigated the effects of prior residence and previous cohabitation on the hierarchical relationships established between two wasps which were placed together in hitherto unknown surroundings. When two wasps which were both previously of alpha rank are brought together on the nest already occupied by one of them, the prior resident is likely to become the alpha individual in the newly established hierarchy. Prior residence is not a decisive factor however when two beta-ranked wasps come together. When two wasps which previously belonged to the same social hierarchy were separated for one week, during which time they both became alpha individuals before being reunited, they tended to revert to their original status. The low intensity level of their reactions suggests here that a persistent mnesic trace may have led them to adopt their previous respective attitudes.     

Relative Importance of Sex,Pre-Starvation Body Mass and Structural Body Size in the Determination of Exceptional Starvation Resistance of Anchomenus dorsalis (Coleoptera: Carabidae)

In nature, almost all animals have to cope with periods of food shortage during their lifetimes. Starvation risks are especially high for carnivorous predatory species, which often experience long intervals between stochastic prey capturing events. A laboratory experiment using the common predatory carabid beetle Anchomenus dorsalis revealed an exceptional level of starvation resistance in this species: males survived up to 137 days and females up to 218 days without food at 20°C. Individual starvation resistance was strongly positively affected by pre-starvation body mass but only slightly by beetle structural body size per se. Females outperformed males even when the effect of gender was corrected for the effects of structural body size and pre-starvation body mass. The better performance of females compared to males and of beetles with higher relative pre-starvation body mass could be linked to higher fat content and lean dry mass before starvation, followed by a greater decrease in both during starvation. There was also a difference between the sexes in the extent of body mass changes both during ad libitum feeding and following starvation; the body masses of females fluctuated more compared to males. This study stresses the need to distinguish between body mass and structural body size when investigating the ecological and evolutionary consequences of body size. Investigation of the net effects of body size and sex is necessary to disentangle the causes of differences in individual performances in studies of species with significant sexual size dimorphism.     

Effects of Limb Length,Body Mass,Gender, Gravidity,and Elevation on Escape Speed in the Lizard Psammodromus algirus

Most animals rely on their escape speed to flee from predators. Here, we test several hypotheses on the evolution of escape speed in the lizard Psammodromus algirus. We test that: (1) Longer limbs should improve speed sprint. (2) Heavier lizards should be impaired regarding their sprint speed ability, suggesting a trade-off between fat storage and escape capability. (3) Males should achieve faster speeds due to their higher exposure to predators. (4) Gravid females, with increased body mass, should perform lower speed than non-gravid females. And (5) there are inter-population differences in sprint speed across an elevational gradient. We measured lizards sprint speed in a lineal raceway in the laboratory, filming races in standardized conditions and then calculating their maximal speed. We found that hind limb length greatly determined maximal sprint speed, lizards with longer limbs being faster. In parallel, higher body masses reduced maximal speed, which points to a trade-off between fat storage and escaping capability. Sexual differences also arose, as males were faster than females, as a consequence of males having longer limbs. Regarding females, gravidity did not impair maximal sprint speed, suggesting adaptations which compensate for the increased body mass. Finally, we found no elevational trend in both limbs length and sprint speed. In any case, this study suggests that selection on escape capacity may cast morphological evolution, and affect other life-history traits, such as fat storage and reproduction.