Reptiles use pterin and carotenoid pigments to produce yellow, orange, and red colors. These conspicuous colors serve a diversity of signaling functions, but their molecular basis remains unresolved. Here, we show that the genomes of sympatric color morphs of the European common wall lizard (Podarcis muralis), which differ in orange and yellow pigmentation and in their ecology and behavior, are virtually undifferentiated. Genetic differences are restricted to two small regulatory regions near genes associated with pterin [sepiapterin reductase(SPR)] and carotenoid [beta-carotene oxygenase 2(BCO2)] metabolism, demonstrating that a core gene in the housekeeping pathway of pterin biosynthesis has been coopted for bright coloration in reptiles and indicating that these loci exert pleiotropic effects on other aspects of physiology. Pigmentation differences are explained by extremely divergent alleles, and haplotype analysis revealed abundant transspecific allele sharing with other lacertids exhibiting color polymorphisms. The evolution of these conspicuous color ornaments is the result of ancient genetic variation and cross-species hybridization.

Evidence accumulated over the last few decades demonstrates that all reptiles examined thus far continue to add neurons at a high rate and in many regions of the adult brain. This so-called adult neurogenesis has been described in the olfactory bulbs, rostral forebrain, all cortical areas, anterior dorsal ventricular ridge, septum, striatum, nucleus sphericus, and cerebellum. The rate of neuronal production varies greatly among these brain areas. Moreover, striking differences in the rate and distribution of adult neurogenesis have been noted among species. In addition to producing new neurons in the adult brain, lizards, and possibly other reptiles as well, are capable of regenerating large portions of their telencephalon damaged as a result of experimentally-induced injuries, thus exhibiting an enormous potential for neuronal regeneration. Adult neurogenesis and neuronal regeneration take advantage of the same mechanisms that are present during embryonic neurogenesis. New neurons are born in the ependyma lining the ventricles and migrate radially through the brain parenchyma along processes of radial glial cells. Several lines of evidence suggest that radial glial cells also act as stem cells for adult neurogenesis. Once they reach their final destination, the young neurons extend axons that reach appropriate target areas. Tangential migration of neurons alongside the ventricular ependyma has also been reported. Most of these tangentially migrating neurons seem to be destined for the olfactory bulbs and are, thus, part of a system similar to the mammalian rostral migratory stream. The proliferation and recruitment of new neurons appear to result in continuous growth of most areas showing adult neurogenesis. The functional consequences of this continuous generation and integration of new neurons into existing circuits is largely conjectural, but involvement of these phenomena in learning and memory is one likely possibility.

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Reptiles use pterin and carotenoid pigments to produce yellow, orange, and red colors.These conspicuous colors serve a diversity of signaling functions, but their molecular basis remains unresolved. Here, we show that the genomes of sympatric color morphs of the European common wall lizard, which differ in orange and yellow pigmentation and in their ecology and behavior, are virtually undifferentiated. Genetic differences are restricted to two small regulatory regions, near genes associated with pterin (SPR) and carotenoid metabolism (BCO2), demonstrating that a core gene in the housekeeping pathway of pterin biosynthesis has been co-opted for bright coloration in reptiles and indicating that these loci exert pleiotropic effects on other aspects of physiology. Pigmentation differences are explained by extremely divergent alleles and haplotype analysis revealed abundant trans-specific allele sharing with other lacertids exhibiting color polymorphisms. The evolution of these conspicuous color ornaments is the result of ancient genetic variation and cross-species hybridization.To investigate the genetic and evolutionary bases of the vivid colors displayed by reptiles, and to test hypothesis about how and why color polymorphisms and correlated trait variation persist within populations, we studied the European common wall lizard (Podarcis muralis) (Fig. 1A)a polymorphic lizard in which the ventral scales of males and females exhibits one of three distinct colors (orange, yellow, and white) or a mosaic pattern combining two colors (orange-yellow and orange-white) (12,13). Each of these five color morphs can be found throughout most of the broad geographic distribution of the species (Fig. 1B), and are shared by intraspecific sub-lineages thought to have diverged up to 2.5 million years ago (14). While the white morph is typically the most common (>50%), the relative frequency of morphs is highly variable even at small regional scales and the yellow or orange morphs may occasionally prevail (15,16) (SI Appendix, Fig. S1). The widespread distribution and persistence of color variation is thought to be due to balancing selection and the product of an interplay between natural and sexual selection (17). Previous work has shown that morphs mate assortatively with respect to ventral color (~75% of pairs) and differ in additional traits, including morphology, behavior, physiology, immunology, and reproduction (12,(18)(19)(20)(21)(22). The mode of inheritance of the color morphs is unknown. RESULTS Carotenoid and pterin pigments underlie pigmentation differencesWe began by determining the biochemical and cellular basis of pigmentation differences among morphs. Using electron microscopy (TEM), we found that the ventral skin of all morphs contained the same set of dermal pigment cells arranged as three superimposed 6 layers (xantophores, iridophores, and melanophores; Fig. 1C). The iridophore layer was thinner in orange individuals compared to yellow and white, but the most noticeable difference among morphs was observed in the...

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Numerosity discrimination, the ability to distinguish between sets with more and less items, is recognised as the foundation for higher numerical abilities. Understanding numerosity discrimination from a comparative perspective is hence pivotal in tracing the evolution of numerical representation systems. However, numerosity discrimination has been well studied only in vertebrates, where two innate systems of number representation have been described: an 'analog magnitude system' used to discriminate among numerosities by representing them as cardinal magnitudes and a 'parallel individualisation system' that allows precise discrimination among small arrays of items (< or =4) by representing objects individually. We investigated the existence of quantity discrimination in an insect species (Tenebrio molitor) by using a spontaneous two-choice procedure in which males were exposed to substrates bearing odours from different numbers of females (< or =4) in increasing numerosity ratios (1:4, 1:3 and 1:2). We show that males can discriminate sources of odours reflecting 1 versus 4 and 1 versus 3 females, but not 2 versus 4 or 1 versus 2, indicating that T. molitor males exhibit a marked preference for sources reflecting more female donors only when numerosity ratios are below 1:2. We discuss the functional significance of this finding and whether our pattern of results could be best explained by summation of a non-numerical continuous variable or by the existence of a numerosity discrimination mechanism with an operational signature ratio of 1:2.

Colour signals play a key role in regulating the intensity and outcome of animal contests. Males of the common wall lizard (Podarcis muralis) show conspicuous ventrolateral ultraviolet (UV)-blue and black patches. In addition, some populations express a striking ventral colour polymorphism (i.e., discrete orange, white and yellow morphs). In this study, we set out to evaluate the potential signalling function of these colour patches by staging pairwise combats between 60 size-matched adult lizards (20 per morph). Combats were held in a neutral arena, with each lizard facing rivals from the three morphs in a tournament with a balanced design. We then calculated a fighting ability ranking using the Bradley–Terry model, and used it to explore whether ventral colour morph, the size of UV-blue and black patches or the spectral characteristics of UV-blue patches (i.e., brightness, hue, chroma) are good predictors of fighting ability. We did not find an effect of the UV-blue patches on contest outcome, but the size of black patches emerged as a good predictor of fighting ability. We also found that winners were more aggressive when facing rivals with black patches of similar size, suggesting that black patches play a role in rival assessment and fighting rules. Finally, we found that orange males lost fights against heteromorphic males more often than yellow or white males. In light of these results, we discuss the potential signalling function of ventrolateral and ventral colour patches in mediating agonistic encounters in this species.