Abstract. Sexual selection operates by acting on variation in mating success. However, since selection acts on wholeorganism manifestations (i.e., performance) of underlying morphological traits, tests for phenotypic effects of sexual selection should consider whole-animal performance as a substrate for sexual selection. Previous studies have revealed positive relationships between performance and survival, that is, natural selection, but none have explicitly tested whether performance may influence reproductive success (through more matings), that is, sexual selection. Performance predicts dominance in some species, implying the effects of sexual selection, but how it does so has not been established, nor is it certain whether performance might be a by-product of selection for something else, for example, elevated circulating testosterone levels. We investigated the potential for sexual selection on sprint speed performance in collared lizards (Crotaphytus collaris), considering the potential mediating effects of circulating hormone levels. Among territorial, adult male collared lizards, only sprint speed significantly predicted territory area and number of offspring sired as determined by genetic paternity analysis. Body size, head size, and hind limb length had no effect. Neither plasma testosterone levels nor corticosterone levels correlated with sprint speed, territory area, or number of offspring sired. Thus, our results provide a direct link between whole-animal performance and reproductive success, suggesting that intrasexual selection can act directly on sprint speed performance and drive the evolution of underlying morphological traits.

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Life-history traits such as offspring size, number and sex ratio are affected by maternal feeding rates in many kinds of animals, but the consequences of variation in maternal diet quality (rather than quantity) are poorly understood. We manipulated dietary quality of reproducing female lizards (Amphibolurus muricatus; Agamidae), a species with temperature-dependent sex determination, to examine strategies of reproductive allocation. Females maintained on a poor-quality diet produced fewer clutches but massively (twofold) larger eggs with lower concentrations of yolk testosterone than did conspecific females given a high-quality diet. Although all eggs were incubated at the same temperature, and yolk steroid hormone levels were not correlated with offspring sex, the nutrient-deprived females produced highly male-biased sex ratios among their offspring. These responses to maternal nutrition generate a link between sex and offspring size, in a direction likely to enhance maternal fitness if large body size enhances reproductive success more in sons than in daughters (as seems plausible, given the mating system of this species). Overall, our results show that sex determination in these animals is more complex, and responsive to a wider range of environmental cues, than that suggested by the classification of 'environmental sex determination'.

The green anol (Anolis carolinensis) is an excellent reptilian model for studying reproductive behavior and the neural and muscular morphology that supports it. This lizard has been the subject of behavioral and ecological study for more than 100 yr, and a rich literature exists on its natural history. Both courtship and copulatory behaviors reveal sex and seasonal differences, which allow for the study of mechanisms regulation naturally occurring variation in performance at multiple levels within a single animal model. Green anoles are readily obtained due to their abundance in the wild; once in the laboratory, they are easily maintained, bred, and reared. Background on the natural history and husbandry of this lizard is provided, and the authors' research program on the regulation of reproductive anatomy and behavior is reviewed, Discussion includes the similarities and differences in the mechanisms mediating both structure and function compared with more traditional animal models. This type of comparative research will make it possible to identify the fundamental principles governing reproductive biology, thus advancing both basic and applied knowledge.

Sexual selection operates by acting on variation in mating success. However, since selection acts on whole-organism manifestations (i.e., performance) of underlying morphological traits, tests for phenotypic effects of sexual selection should consider whole-animal performance as a substrate for sexual selection. Previous studies have revealed positive relationships between performance and survival, that is, natural selection, but none have explicitly tested whether performance may influence reproductive success (through more matings), that is, sexual selection. Performance predicts dominance in some species, implying the effects of sexual selection, but how it does so has not been established, nor is it certain whether performance might be a by-product of selection for something else, for example, elevated circulating testosterone levels. We investigated the potential for sexual selection on sprint speed performance in collared lizards (Crotaphytus collaris), considering the potential mediating effects of circulating hormone levels. Among territorial, adult male collared lizards, only sprint speed significantly predicted territory area and number of offspring sired as determined by genetic paternity analysis. Body size, head size, and hind limb length had no effect. Neither plasma testosterone levels nor corticosterone levels correlated with sprint speed, territory area, or number of offspring sired. Thus, our results provide a direct link between whole-animal performance and reproductive success, suggesting that intrasexual selection can act directly on sprint speed performance and drive the evolution of underlying morphological traits.

Digit length ratios are organized during embryonic development and may show sexual dimorphism related to steroid exposure. The second-to-fourth digit length ratio (2D:4D) has received the most attention. In the present study, we measured 2D:4D of all four feet of adult male and female green anoles ( Anolis carolinensis Voigt, 1832) to determine whether it is sexually dimorphic and whether results are repeatable across laboratories. Lizards were housed at Michigan State University (MSU) and Oklahoma State University (OSU), and one investigator at each institution used digital calipers to measure the 2D:4D of each foot. At both MSU and OSU, we found that males had a significantly larger 2D:4D on the back right foot than females did, and that no sex difference existed in either the back left or the front right foot. Furthermore, although no sex difference in the front left foot was found at MSU, the 2D:4D on this foot was larger in females at OSU. Our results demonstrate both sexual dimorphism in 2D:4D and repeatability between laboratories, but they also suggest the importance of verifying such repeatability if 2D:4D or any other digit length ratio is used as a potential indicator of the early steroid environment.

The subventricular zone (SVZ) is greatly expanded in primates with gyrencephalic cortices, and is thought to be absent from vertebrates with three-layered, lissencephalic cortices, such as the turtle. Recent work in rodents has shown that Tbr2-expressing neural precursor cells in the SVZ produce excitatory neurons for each cortical layer in the neocortex. Many excitatory neurons are generated through a two-step process in which Pax6-expressing radial glial cells divide in the VZ to produce Tbr2-expressing intermediate progenitor cells, which divide in the SVZ to produce cortical neurons. We investigated the evolutionary origin of SVZ neural precursor cells in the prenatal cerebral cortex by testing for the presence and distribution of Tbr2-expressing cells in the prenatal cortex of reptilian and avian species. We found that mitotic Tbr2+ cells are present in the prenatal cortex of lizard, turtle, chicken and dove. Furthermore, Tbr2+ cells are organized into a distinct SVZ in the DVR of turtle forebrain, and in the cortices of chicken and dove. Our results are consistent with the concept that Tbr2+ neural precursor cells were present in the common ancestor of mammals and reptiles. Our data also suggest that the organizing principle guiding the assembly of Tbr2+ cells into an anatomically distinct SVZ, both developmentally and evolutionarily, may be shared across vertebrates. Finally, our results indicate that Tbr2 expression can be used to test for the presence of a distinct SVZ, and to define the boundaries of the SVZ in developing cortices.

Environmental conditions that reproductive females experience can influence patterns of offspring provisioning and fitness. In particular, prey availability can influence maternal reproduction and, in turn, affect the viability of their offspring. Although such maternal effects are widespread, the mechanisms by which these effects operate are poorly understood. We manipulated the amount of prey available to female brown anole lizards (Anolis sagrei) to evaluate how this factor affects patterns of reproductive investment (total egg output, egg size, yolk steroids) and offspring viability (morphology, growth, survival). Experimental reduction of yolk in a subset of eggs enabled us to evaluate a potential causal mechanism (yolk investment) that mediates the effect of maternal prey availability on offspring viability. We show that limited prey availability significantly reduced egg size, which negatively influenced offspring size, growth, and survival. Experimental yolk removal from eggs directly reduced offspring size, which, in turn, negatively affected offspring growth and survival. These findings show that maternal environments (i.e., low prey) can affect offspring fitness via an indirect effect of yolk investment on offspring size and highlight the complex set of indirect effects by which maternal effects can operate.