Genetic variation and climatic selection in the lizard Agama stellio in Israel and Sinai

Allozymic variation in proteins encoded by 25 loci was analyzed electrophoretically in 242 adult specimens representing nine populations of the Levantine lizard, Agama stellio, comprising two subspecies: the Mediterranean A. stellio subsp., and the desert-inhabiting A. stellio brachydactyla from the Negev and Sinai. Likewise, four body traits were measured in the same populations. The nine populations were sampled along a general southward transect of increasing aridity. Agama stellio is above average in both polymorphism, P, and heterozygosity, H, as compared to other reptiles and vertebrates in general, displaying levels of genetic variation characterizing habitat generalist vertebrates. In the populations studied no fixation of alternative alleles was found in any of the 25 loci: rather the commonest allele was either fixed or predominated in 23 of 25 loci examined. Eleven loci (44%) were monomorphic in all nine populations. However, of the remaining 14 polymorphic loci, eight were strongly polymorphic displaying distinct genetic differentiation between populations. Genetic diversity (indexed by P and H) displayed geographic variation and was slightly higher in A.s. brachydactyla than in A. stellio subsp. Nevertheless, genic similarity between populations was high. A statistically significant amount of morphological variation between localities was found for all body characters. In general, body size increased southwards and eastwards with aridity. Selection at some loci is suggested by significant deviation from Hardy-Weinberg expectations and possibly by excess heterogeneity of effective inbreeding coefficients, F_e. Furthermore, allozymic variation at seven loci (Ldh-1, Idh-1, 6-Pgd-1, Aat-1, Pgm-2, Pept-1, and Trf) and geographic variation in body size and weight were significantly correlated with, and predictable by, climatic variables, primarily by water availability and secondarily by temperature. Finally, allozymic and morphological variations were partly correlated. The spatial patterns and ecological correlates of genic and morphological variations in Agama stellio in Israel and Sinai suggest that at least some proteins and body size differentiate geographically and appear to be adaptive, presumably with respect to factors affecting the availability of water.