Genetic diversity, climatic selection and speciation of Sphincterochila landsnails in Israel

TLDR: Allozymic variation in proteins encoded by 29 loci was analysed electrophoretically in 364 adult specimens representing 12 populations and five species of the landsnail Sphincterochila in Israel along a north-south general transect of increasing aridity and suggests that climatic selection plays a conspicuous role in allozymics morphological differentiation into ecologically adaptive patterns.

Abstract: Allozymic variation in proteins encoded by 29 loci was analysed electrophoretically in 364 adult specimens representing 12 populations and five species of the landsnail Sphincterochila in Israel along a north-south general transect of increasing aridity. In addition, geographic variation in three morphological body variables of these snails was also studied. The results indicate that: (i) most loci (86%) are strongly polymorphic; (ii) most loci (65%) display fixation of alternative alleles either within or between species; (iii) most of the variant alleles (51%) are not widespread, and genie differentiation is very high (66%) between populations and species, indicating sharp local and regional geographic differentiation; (iv) clinal patterns are rare or nonexistent; (v) populations of Sphincterochila display average estimates of mean alleles per locus,A=1.53; polymorphism, P (5% criterion) = 0.31; heterozygosity, H=0.07; and genie diversity, He= 0.11; (vi) wide geographic variation within and between species is displayed in A= 1.18-2.07; P=0.11–0.61; H=0.02-0.15, and He = 0.042-0.22. Wright's fixation index, F, ranges from 0.03 to 0.65. (vii) Genie diversity, He, increases southwards with aridity from 0.051 to 0.145. (viii) A differential amount of variation in different functional classes of enzymes follows the Gillespie-Kojima hypothesis, (ix) Coefficients of genetic distance, D, between populations are high, D= 0.34, range 0.09-0.58, and between species, D= 0.27, range 0.12-0.40. D's within species may be higher than between species. Likewise, D's increase clinally southwards, (x) Deviations from Hardy-Weinberg equilibria were found in several loci in some populations and species, (xi) A statistically significant (P< 0.001) amount of morphological variation of body variables exists within and between species. Size between three species increases eastwards and southwards with aridity, (xii) P, H, He, and allozymic variation in several gene loci are significantly correlated with, and predictable by, climatic variables, primarily those related to the moisture index, (xiii) Allozymic and morphological variations are partly correlated, (xiv) Significant microgeographical climatic differentiation was found in three critical tests. The pattern of genetic variation within and between species suggests that: (a) climatic selection plays a conspicuous role in allozymic morphological differentiation into ecologically adaptive patterns; (b) the environmental variation model seems to be a good predictor of genetic variation in Sphincterochila; (c) adaptive radiation of the five species of Sphincterochila in Israel occurred during Pleistocene times in accord with climatic differentiation and apparently involved few changes of structural genes.