Proteins are ubiquitous macromolecules that display a vast repertoire of chemical and enzymatic functions, making them suitable candidates for chemosignals, used in intraspecific communication. Proteins are present in the skin gland secretions of vertebrates but their identity, and especially, their functions, remain largely unknown. Many lizard species possess femoral glands, i.e., epidermal organs primarily involved in the production and secretion of chemosignals, playing a pivotal role in mate choice and intrasexual communication. The lipophilic fraction of femoral glands has been well studied in lizards. In contrast, proteins have been the focus of only a handful of investigations. Here, we identify and describe inter-individual expression patterns and the functionality of proteins present in femoral glands of male sand lizards (Lacerta agilis) by applying mass spectrometry-based proteomics. Our results show that the total number of proteins varied substantially among individuals. None of the identified femoral gland proteins could be directly linked to chemical communication in lizards, although this result hinges on protein annotation in databases in which squamate semiochemicals are poorly represented. In contrast to our expectations, the proteins consistently expressed across individuals were related to the immune system, antioxidant activity and lipid metabolism as their main functions, showing that proteins in reptilian epidermal glands may have other functions besides chemical communication. Interestingly, we found expression of the Major Histocompatibility Complex (MHC) among the multiple and diverse biological processes enriched in FGs, tentatively supporting a previous hypothesis that MHC was coopted for semiochemical function in sand lizards, specifically in mate recognition. Our study shows that mass spectrometry-based proteomics are a powerful tool for characterizing and deciphering the role of proteins secreted by skin glands in non-model vertebrates.

It is postulated that melanism in ectotherms is adaptive by enhancing thermoregulation, subsequent resource acquisition, and growth. Such effects may differ between the sexes as a result of the differential costs of self-maintenance and reproduction, but empirical support for the sex-specific consequences of melanism remains inconsistent. We studied the effects of melanism on body size and sex ratio in a population of the European grass snake (Natrix natrix) in SE Poland and also carried out a systematic review of the literature on the consequences of melanism in terrestrial snakes. Melanistic grass snakes of both sexes appeared to be smaller than the typical phenotype, which indicates higher predation pressure and minimal thermal benefits for black individuals. A female-biased sex ratio was observed in the typical phenotype, but not in melanistic snakes, suggesting that the costs for females and/or benefits for males are higher in melanistic individuals. In conjunction with earlier studies, our data indicate that the consequences of melanism may be related to the reproductive mode of species. In viviparous species, melanism tends to improve growth and/or body size and is more frequent in females, whereas the opposite holds for oviparous snakes. Further studies on melanism should examine a wider array of species with different reproductive strategies and traits beyond the usual thermal benefits.

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