Seasonal variations of plasma testosterone among colour-morph common wall lizards (Podarcis muralis).

Sexual steroids influence reproductive behaviours and promote secondary sexual traits. In male lizards, increasing levels of testosterone (T) bolster conspicuous colouration, stimulate territoriality, and trigger antagonistic interactions among rivals. Moreover, in colour polymorphic species, reproductive strategy, aggressiveness and T levels can differ between morphs. Therefore, T level is considered as an important mechanism that regulates the expression of colour polymorphism and sexual behaviours of males. But in the polymorphic territorial wall lizard (Podarcis muralis), a lack of relationship between morphs and aggressiveness challenges the notion that T plays such a role. To examine this issue, we compared adult T levels among three colour morphs (white, yellow and red) through repeated sampling during the mating season. High T levels were observed at the onset of the mating season followed by a significant decrease, a pattern documented in other lizard species. Mean T levels did not differ among morphs. However, yellow males maintained significantly higher T levels over time and displayed a stronger subsequent decline. Overall, in this species, seasonal T patterns differ among morphs, not mean values. Previous studies revealed that T suppresses the immune response; suggesting that a strong initial investment promoted by high T levels may trade-off against immunity (maintenance). Further experimental investigations are required to clarify the relationship between T and reproductive effort in polymorphic species that exhibit complex temporal pattern of T levels.

Refugia within refugia as a key to disentangle the genetic pattern of a highly variable species: the case of Rana temporaria Linnaeus, 1758 (Anura, Ranidae).

Two distinct lineages of Rana temporaria are known in the Palaearctic region, but it is uncertain whether this species persisted in one or more Pleistocene refugia. We resolved the phylogeographic history and genetic variability of R. temporaria in the Italian peninsula, a 'traditional' Pleistocene refugium, and related our findings to patterns described for other European populations. We sequenced the mitochondrial markers Cox I and cytochrome b. Phylogenetic reconstruction only indicated the presence of haplotypes belonging to the Western lineage in the Italian peninsula. Overall, the genetic variability of Italian populations was higher than other European populations, which shared haplotypes with the Alpine populations. We demonstrated subdivision into five main Italian sublineages, which was associated with a geographical structure of populations in two divergent groups. In particular, one Apennine group might have resulted from bottlenecks during the last interglacials ages. In contrast, Alpine populations were recently diverged and showed incomplete lineage sorting. Our data indicate that the Italian peninsula served as refugium for the Western lineage of R. temporaria. Dispersion towards Central Europe probably started only from the western slope of the Alps via a rapid leading edge expansion. The identified structure is partially congruent with traditional peripheral refugia identified for plants. This evolutionary scenario does not support any taxonomic distinction at the subspecific level for R. temporaria.

Sex-, morph- and size-specific susceptibility to stress measured by haematological variables in captive common wall lizard Podarcis muralis.

In polymorphic species of animals, colour morphs may show alternative physiological properties, and hence evolve or be maintained as an indirect response to selection exerted on these physiological attributes. In this study, we investigated if different colour morphs (white, red and yellow) of the polymorphic common wall lizard differed in their physiological responses to a long-term stress by determining variation between capture and release in leukocytes profiles, haemoparasite loads and body condition of male and females maintained in captivity throughout the breeding season. We found that most blood parameters of lizards varied significantly following captivity, and this variation was sex-, morph- and size-dependent. In particular, the heterophil:lymphocyte ratio (H:L), a sensitive measure of immunodepression and long-term stress, varied significantly among yellow females, larger individuals significantly increasing and smaller individuals decreasing their H:L ratio after captivity. This trend was reversed in red females, where smaller individuals presented raised H:L index at release. Our study indicated that response to long-term stressful conditions, such as those induced by captivity, differed among common wall lizard colour morphs, implying a sex-, size-(i.e. age) and morph-specific sensitivity to stress, and hence a different physiological profile of colour morphs, which may contribute to the maintenance of colour polymorphism in this species.

Male-male combats in a polymorphic lizard: residency and size, but not color, affect fighting rules and contest outcome.

Theoretical models predict that the outcome of dyadic agonistic encounters between males is influenced by resource-holding potential, resource value, and intrinsic aggressiveness of contestants. Moreover, in territorial disputes residents enjoy a further obvious competitive advantage from the residency itself, owing to the intimate familiarity with their territory. Costs of physical combats are, however, dramatically high in many instances. Thus, signals reliably reflecting fighting ability of the opponents could easily evolve in order to reduce these costs. For example, variation in color morph in polymorphic species has been associated with dominance in several case studies. In this study, we staged asymmetric resident-intruder encounters in males of the common wall lizard Podarcis muralis, a species showing three discrete morphs (white, yellow, and red) to investigate the effects of asymmetries in color morph, residency, and size between contestants on the outcome of territorial contests. We collected aggression data by presenting each resident male with three intruders of different color morph, in three consecutive tests conducted in different days, and videotaping their interactions. The results showed that simple rules such as residency and body size differences could determine the outcome of agonistic interactions: residents were more aggressive than intruders, and larger males were competitively superior to smaller males. However, we did not find any effect of color on male aggression or fighting success, suggesting that color polymorphism in this species is not a signal of status or fighting ability in intermale conflicts.

Sexual dimorphism in digit length ratios in two lizard species.

Sexual dimorphism in digit length ratios has been reported for humans, a few other mammals, and two bird species. This dimorphism is thought to arise via an interaction between the prenatal exposure of the embryo to sex hormones and the Hox genes, which are highly conserved among vertebrates and control the development of both the appendices, including fingers and toes, and the urogenital system. In this study, we report on sexual dimorphism in 2D:3D, 2D:4D, and 3D:4D contralateral ratios of the forelimbs in two species of oviparous lizards, the common wall lizard (Podarcis muralis) and the tree skink (Mabuya planifrons), as measured on museum specimens. We found that male P. muralis had a larger 2D:4D ratio on both sides and larger 2D:3D ratio on the left side than females, whereas in M. planifrons, males had lower 2D:3D ratios than females on the left side. The two species show opposite patterns of sexual dimorphism in body size, males being larger than females in P. muralis, and the reverse in M. planifrons, suggesting that interspecific variation of sex differences in digit ratios could be associated with sex-specific growth trajectories. There was a limited evidence for directional asymmetry in digit ratios. Therefore, our findings provide the first evidence that digit ratios are sexually dimorphic in any reptile species and are consistent with the idea that the genetic link between limb development and the urogenital system had been established with the evolution of the earliest terrestrial tetrapods. Importantly, many lizard species with genetic sex determination, including the ones we studied, are oviparous and may represent valuable animal models for experimental tests of the association between prenatal exposure to androgens or estrogens and digit ratios.