Discrete characters of the occlusal surface (additional cusps) have been studied to elaborate a new approach to the identification of the Ground Squirrel species Spermophilus odessanus, S. suslicus, S. pygmaeus, S. citellus, and S. xanthoprymnus. Data on the presence/absence of the additional cusps have been represented as star plots and, in addition, have been studied using discriminant function analysis. The species-specific sets of the characters (patterns of bunodonty) have been revealed and are of high diagnostic value. The Citellus-set is defined by the presence of mesostyles and the rareness of the metastylids, paraconules and metaconules, hypostyles and protostyles. The Pygmaeus-set is characterized by the presence of additional cusps in the lower cheek teeth. The Odessanus-oriented set is found in the Spermophilus pygmaeus, S. odessanus, and S. suslicus. The relatively high frequency of additional cusps of the metaloph and the paraloph is characteristic for this set. The Plesiomorphic-set (characters shared by all the studied species and for this reason regarded herein as ancestral) is found in S. xanthoprymnus. The patterns of bunodonty serve as diagnostic criteria only as a whole: the shape of a star plot (relations among the character frequencies), rather than certain character values, is indicative. An optimal level of identification of species is possible based on the combination of the discrete characters mentioned and on the size parameters of the third upper molar. The occlusal sets are intended to remain stable during the time of species existence and seem to correspond to trends in specialization. The functional meaning of the sets can be explained by the dependence between the presence/absence of the discrete characters and the shape of the crown and its main lophs. Each pattern is likely to correspond to a trophic niche, and this niche corresponds to the species. J. Morphol. 277:814-825, 2016. © 2016 Wiley Periodicals, Inc.

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Ground squirrels were an important member of the Pleistocene steppe-tundra mammal community. They evolved ecological specialisations and exhibit behaviours that make them particularly informative subjects to study palaeoenvironmental constraints affecting species distribution and speciation. Interspecific competition and isolating geographical barriers are considered as the principal factors that define species range boundaries.The present paper provides a first comprehensive compilation of the living and extinct Spermophilus species in Europe. These data suggest 'patchwork quilt' model for the expansion and spatial distribution of ground squirrel species. Here we consider mainly small-sized Spermophilus species because large-sized (e.g., S. superciliosus) ground squirrels consist another 'patchwork quilt', which overlap the first one. This overlapping of the species ranges is possible because of the size difference that lowers interspecific competition (Hutchinson's rule).We consider two main types of range boundaries. One type includes roughly 'sub-parallel' boundaries that oscillate in concert with climatic and vegetational changes (a case of climatically controlled competitive exclusion). The other type consists of roughly 'sub-meridional' boundaries corresponding to geographical barriers (e.g., water barriers, mountain ridges); these boundaries are rather stable. Examples of 'sub-parallel range modifications include: oscillations of boundaries between S. pygmaeus and S. suslicus; the immigration of S. citellus into the Pre-Carpathian area; the branching of S. suslicus from S. pygmaeus; the regional appearance of the Late Pleistocene species S. severskensis and S. citelloides. Examples of 'sub-meridional events' are: the crossing of the Danube by S. citellus; the appearance of an isolated population of S. pygmaeus on right bank of the Dnieper during the Late Pleistocene to Middle Holocene; a crossing of the Dnieper river by S. pygmaeus, which resulted in the appearance of S. odessanus; the intrusion of eastern populations of S. pygmaeus into the Trans-Volga areas.

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