13Snakes frequently exhibit ontogenetic and sexual variation in head dimensions, as well as 14 the occurrence of distinct colour morphotypes which might be fitness-related. In this 15 study, we used linear biometry and geometric morphometrics to investigate intraspecific 16 morphological variation related to allometry and sexual dimorphism in Vipera seoanei, a 17 species that exhibits five colour morphotypes, potentially subjected to distinct ecological 18 pressures. We measured body size (SVL), tail length and head dimensions in 391 19 specimens, and examined variation in biometric traits with respect to allometry, sex and 20 colour morph. In addition, we analysed head shape variation by recording the position of 21 29 landmarks in 123 specimens and establishing a low-error protocol for implementing 22 geometric morphometrics to European vipers. All head dimensions exhibited significant 23 allometry, while sexual differences occurred for SVL, relative tail length and snout 24 height. After considering size effects, we found significant differences in body 25 proportions between the sexes and across colour morphs, which suggests an important 26 influence of lowland and montane habitats in shaping morphological variation. By 27 contrast, head shape did not exhibit significant variation across sexes or colour morphs.
28Instead it was mainly associated to allometric variation, where the supraocular and the 29 rear regions of the head were the areas that varied the most throughout growth and across 30 individuals. Overall, this study provides a thorough description of morphological 31 variability in Vipera seoanei and highlights the relevance of combining different tools 32 (i.e. linear and geometric morphometrics) and analyses to evaluate the relative 33 contribution of different factors in shaping intraspecific variation.34 35 36 37 38 Understanding how morphological variation across individuals arises and how it is 39 distributed both temporally and spatially have attracted the attention of biologists over 40 the years (Verwaijen, Van Damme and Herrel, 2002; Harmon et al., 2003; 41 Kaliontzopoulou, Pinho and Martínez-Freiría, 2018). 42 Size is the predominant axis of morphological variation within and among populations 43 (Rohlf, 1990). As such, allometry, the dependence of shape on size, is a major framework 44 for understanding how and why different traits vary (Klingenberg, 2016). First, changes 45 in size and shape occurring during growth and their relationship (i.e. ontogenetic 46 allometry) are of essential importance for investigating the developmental processes 47 producing the structures of interest (McNamara, 2012). Second, allometric variation 48 across individuals at the same developmental stage within a population (i.e. static 49 allometry) can be informative on the selective processes acting on individuals, since 50 allometric parameters can be directly linked to both ecological adaptation (Gould, 1966) 51and sexual selection (Bonduriansky and Day, 2003). Indeed, body structures that are of 52 partic...