Sources of intraspecific morphological variation in Vipera seoanei: Allometry, sex, and colour phenotype

Snakes frequently exhibit ontogenetic and sexual variation in head dimensions, as well as the occurrence of distinct colour morphotypes which might be fitness-related. In this study, we used linear biometry and geometric morphometrics to investigate intraspecific morphological variation related to allometry and sexual dimorphism in Vipera seoanei, a species that exhibits five colour morphotypes, potentially subjected to distinct ecological pressures. We measured body size (SVL), tail length and head dimensions in 391 specimens, and examined variation in biometric traits with respect to allometry, sex and colour morph. In addition, we analysed head shape variation by recording the position of 29 landmarks in 123 specimens and establishing a low-error protocol for implementing geometric morphometrics to European vipers. All head dimensions exhibited significant allometry, while sexual differences occurred for SVL, relative tail length and snout height. After considering size effects, we found significant differences in body proportions between the sexes and across colour morphs, which suggests an important influence of lowland and montane habitats in shaping morphological variation. By contrast, head shape did not exhibit significant variation across sexes or colour morphs. Instead it was mainly associated to allometric variation, where the supraocular and the rear regions of the head were the areas that varied the most throughout growth and across individuals. Overall, this study provides a thorough description of morphological variability in Vipera seoanei and highlights the relevance of combining different tools (i.e. linear and geometric morphometrics) and analyses to evaluate the relative contribution of different factors in shaping intraspecific variation.