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Skulls of a male (a) and a female (b) of the three species of Eurasian badgers: 1. Meles meles , European form (1a: ZIN 35056, 1b: ZIN 16909, specimens from Leningrad Province, Russia), 2. Meles leucurus , Siberian form (2a: ZIN 1148, 2b: ZIN 1139, specimens from Eastern Kazakhstan), 3. Meles anakuma (3a: NSM 1071, 3b: NSM 27353, specimens from Honshu, Japan). 

Skulls of a male (a) and a female (b) of the three species of Eurasian badgers: 1. Meles meles , European form (1a: ZIN 35056, 1b: ZIN 16909, specimens from Leningrad Province, Russia), 2. Meles leucurus , Siberian form (2a: ZIN 1148, 2b: ZIN 1139, specimens from Eastern Kazakhstan), 3. Meles anakuma (3a: NSM 1071, 3b: NSM 27353, specimens from Honshu, Japan). 

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An analysis of 30 craniological characters of Eurasian badgers (Meles spp.) revealed different levels of sexual size dimorphism (SSD) and geographic variation in the three different species. SSD is displayed mostly in the general size of the skull (condylobasal length, zygomatic width, width of rostrum, and cranial height) and mandible (height of t...

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... Squared Mahalanobis’ distances D 2 between male and female samples summarize the morphological differentiation based on both D - and K axes (Table 2, Fig. 4b). According to this analysis, sexual dimorphism is more pronounced in the Transcaucasian sample of M. meles and Far-Eastern sample of M. leucurus than in the other samples. The European sample of M. meles shows a minimal level of sexual dimorphism. A separation of the sexes by the shape of the skull based on K 1 and K 2 is less clear than by the size of the skull (Fig. 5). The assessment of SSD by D S and S confirmed the above differences in the manifestation of dimorphism in different species and geographical samples of badgers (Fig. 6). Moreover, it was found that in M. anakuma , as well as in the European M. meles , SSD is very weakly expressed, except for the size of the canines. A comparison of different intraspecific geographical samples shows that the value D S depends on the size of females rather than on that of males for the majority of measurements. For example, the difference in the condylobasal length of the skull between males of the relatively large European and the small Transcaucasian M. meles is on average 9.4 mm and that between females is 12.8 mm. Correspondingly, D S for the European sample of M. meles is 2.2 mm ð S 1⁄4 0 : 8 Þ and for the Transcaucasian sample of M. meles is 6.0 mm ð S 1⁄4 2 : 5 Þ . Differences in size between females of both large and small geographical forms are on average greater than those between males. In the European badger, this pattern is more pronounced than in the Asian badger. In M. leucurus , exceptions from this rule are found for the greatest length between the auditory bulla and occipital condyle, postorbital width, neurocranium length, length of the auditory bulla, and length of the lower molar M 2 . The morphological diversity of males in the European sample of M. meles is higher than that of females (Table 3, Fig. 7). At the same time an opposite tendency is observed in the other samples of badgers. No correlation has been found between the level of sexual dimorphism and the degree of morphological diversity within each sex. Morphotypical diversity of teeth of males of the European M. meles , as in the case of the diversity of cranial characters, is higher than in females (Table 4). In the Siberian sample of M. leucurus , an opposite pattern is observed in that females display a greater variability. Size diversity of teeth E D (Table 4) in the European sample of M. meles and Siberian sample of M. leucurus is greater in females (Fig. 8). The degree of morphological diversity of the entire skull correlates positively with the level of sexual dimorphism, expressed by the indices D S and S (Table 3, Fig. 9). Maximum values of morphological diversity are observed in the Transcaucasian M. meles and Far- Eastern M. leucurus . Diversity of the structure, expressed in the number of independent factors of skull variation and reflected in the value ‘‘best-minimum’’ dimension in the MDS model ( d D , d K ), is greatest in the Transcaucasian M. meles . At the species level, the value E D (Table 3), reflecting mostly the diversity of sizes of the skull, and morphotypical diversity E (Table 4) are greater in M. meles than in M. leucurus . At the same time, the diversity of the proportions and size of skulls are greater in M. leucurus (Table 4). There is no direct relationship between the number of characters exhibiting sexual dimorphism and the degree of sexual dimorphism. Thus, at comparable levels of sexual dimorphism assessed by D S and S , the number of characters with sexual dimorphism is only 9 in the European M. meles versus 15 in the Siberian M. leucurus . On the whole, SSD was revealed by a larger number of characters in M. leucurus than in M. meles. In M. anakuma , sexual dimorphism is displayed in only a few cranial characters, such as canine size. Sexual dimorphism of cranial characters are generally correlated with variations in skull size, but exceptions can be expected on theoretical grounds: (1) the dimorphism ratio ( S ) may be constant and not vary with body size, (i.e., isometry); (2) the mean difference between the sexes ( D S ) may be approximately constant, but SSD ratios ( S ) may vary with skull size (i.e., simple allometry); and (3) both D S and S may change (i.e., complex allometry). Our research shows that sexual dimorphism of the skull in badgers is displayed mostly in the form of complex allometry (variant 3), whereas isometric variation (variant 1) is nearly absent. Based on our results, we can reject the statistical null hypotheses of sexual dimorphism being absent in the various species and geographical samples of the Eurasian badger. However, only some skull characters exhibit sexual dimorphism in the three species of Eurasian badgers. Sexual dimorphism is most clearly expressed in the overall size of the skull and mandible and in the length of the upper molar M 1 (see Fig. 10), but was not detectable in the length of the auditory bulla, the interorbital width, the length of lower molar M 2 , and the length of the lower carnassial tooth M 1 . In all studied samples of badgers, there is a pronounced dimorphism in the size of the upper canines, which is typical of many mammalian taxa that possess these teeth, including all carnivores (see Gittleman and Van Valkenburgh 1997). In both the European and Asian badgers, two forms that differ in the degree of SSD have been identified. In the larger badgers (i.e., the European M. meles and Siberian M. leucurus ), sexual dimorphism is weaker in both absolute and relative terms than in the smaller badgers (i.e., the Transcaucasian M. meles and Far-Eastern M. leucurus ). The degree of sexual dimorphism in Eurasian badgers is related mainly to a variation of the females. For the majority of characters, including those for which SSD is statistically non- significant, the absolute difference between females of the large and small forms is greater than the analogous difference between males. This pattern is particularly pronounced in the two forms of M. meles . In the Far-Eastern M. leucurus, an increase of an absolute mean difference between the sexes ( D S ), and particularly of SSD ratios ( S ), of some characters (e.g., viscerocranium length, zygomatic breadth, breadth of rostrum, length and width of upper canine, etc.) is accompanied by a general decline in the values of these characters in both sexes relative to the Siberian M. leucurus . In the Japanese badger M. anakuma (the smallest representative of Meles ), a decline of skull size relative to that of the continental M. leucurus is accompanied by a decline of sexual dimorphism in absolute terms (except canine size). Isometry, however, is complicated by virtue of a more basic decline of skull size in both sexes. As a consequence, the relative index of sexual dimorphism ( S ) is greater in the Japanese badger than in the larger European M. meles and Siberian M. leucurus . The Transcaucasian M. meles and the Far-Eastern M. leucurus are characterized by a complex allometry related to a change in both relative and absolute differences in skull size between males and females. But for all that, sexual dimorphism is maximal in the Transcaucasian M. meles in absolute terms, but it is maximal in the Far-Eastern M. leucurus in relative terms. The morphological diversity of the skull, as defined in this study, is generally greater in female badgers than in males, except in the European M. meles , in which it is greater in males than in females. The degree of size diversity of teeth appears to be greater in females in all forms of the three badger species. On the whole, the degree of morphotypical diversity is apparently correlated with the degree of diversity of tooth shape and does not depend on the size of teeth. A comparison of the morphotypical diversity of teeth of the European and Asian badgers shows that dental variation is relatively independent of skull variation. The degree of morphological diversity of the skull on the whole is positively correlated with the degree of sexual dimorphism expressed by the indices of the absolute mean difference between the sexes ( D S ) and SSD ratios ( S ). At the species level, the diversity of skull size is greater between the two forms of M. meles than between the two forms of M. leucurus , but the diversity of skull proportions is greater in M. leucurus . Therefore, M. meles has more a stable shape of the skull across the studied forms than M. leucurus . The characteristics of sexual dimorphism in badgers do not quite conform to the data known for other representatives of the family Mustelidae (Shubin and Shubin 1975; Zyll de Jong 1992; Abramov and Baryshnikov 2000; Abramov and Tumanov 2003), in which sexual dimorphism is more pronounced in large forms than in smaller ones. In the Eurasian badgers, in contrast, the smaller intraspecific forms display a greater degree of sexual dimorphism than the larger forms. Only the Japanese badger, the smallest representative of the genus Meles , conforms to the pattern observed in other Mustelidae. Even though numerous investigations of sexual dimorphism of Mustelidae have been published, the reasons for sexual dimorphism in the sizes of the skull and body, and for the variation in the degree of sexual dimorphism within a species, have remained unclear. The resource partitioning model attributes differences in sizes to differences in the diet of males and females (Brown and Lasiewski 1972; Shubin and Shubin 1975; Dayan et al. 1989; Dayan and Simberloff 1994; Gittleman and Van Valkenburgh 1997). According to this rule, the sexes achieve optimal sizes, and differences in body size correlate with the size of the food items ingested. It is believed that differences in the feeding apparatus and, in particular, in canine size, indicate some level of selection for niche separation between the sexes (Dayan ...

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