Intercontinental karyotypic differentiation of Chironomus entis Shobanov, a Holarctic member of the C. plumosus group (Diptera, Chironomidae).

Analysis of banding sequences of polytene chromosomes in Palearctic (Russian) and Nearctic (North American) Chironomus entis shows strong karyotype divergence between populations on the two continents. Four out of seven chromosomal arms in the North American C. entis karyotype are characterized by sequences found only in the Nearctic. In total, 44 banding sequences are now known for this species across the Holarctic, including 22 exclusively Palearctic, 6 Holarctic, and 16 exclusively Nearctic sequences. The degree of cytogenetic differentiation between Palearctic and Nearctic C. entis populations is an order of magnitude greater than differentiation among populations within either continent, but is only one third as great as the cytogenetic distance between the sibling species C. entis and C. plumosus. C. entis is the only sibling species of C. plumosus uncovered during cytological identification of Chironomus species from more than 50 North American lakes, indicating that the plumosus sibling-species group is much smaller in the Nearctic than in the Palearctic, where a dozen sibling species are known. Cytogenetic distance values calculated between Nearctic and Palearctic representatives of both C. entis and its sibling species C. plumosus are similar, but result from different patterns of karyotype divergence. New World C. entis is distinguished from Old World populations by the 16 uniquely Nearctic sequences, four of which occur in the homozygous state. In contrast, North American C. plumosus has fewer uniquely Nearctic sequences, and only one that occurs as a homozygote. However, four chromosomal arms in C. plumosus that are polymorphic in the Palearctic show fixation, or near fixation, of Holarctic sequences in the Nearctic C. plumosus karyotype. Thus, both the fixation of Holarctic sequences, and the occurrence or fixation of distinctly Nearctic sequences, contribute significantly to karyotype divergence. Patterns of karyotype divergence in Palearctic and Nearctic populations of different Holarctic chironomid species are discussed relative to intercontinental cytogenetic differentiation in other dipterans.     

Molecular phylogeny of the marmots (Rodentia: Sciuridae): tests of evolutionary and biogeographic hypotheses

There are 14 species of marmots distributed across the Holarctic, and despite extensive systematic study, their phylogenetic relationships remain largely unresolved. In particular, comprehensive studies have been lacking. A well-supported phylogeny is needed to place the numerous ecological and behavioral studies on marmots in an evolutionary context. To address this situation, we obtained complete cytochrome (cyt) b sequences for 13 of the species and a partial sequence for the 14th. We applied a statistical approach to both phylogeny estimation and hypothesis testing, using parsimony and maximum likelihood-based methods. We conducted statistical tests on a suite of previously proposed hypotheses of phylogenetic relationships and biogeographic histories. The cyt b data strongly support the monophyly of Marmota and a western montane clade in the Nearctic. Although some other scenarios cannot be rejected, the results are consistent with an initial diversification in North America, followed by an invasion and subsequent rapid diversification in the Palearctic. These analyses reject the two major competing hypotheses of M. broweri's phylogenetic relationships--namely, that it is the sister species to M. camtschatica of eastern Siberia, and that it is related closely to M. caligata of the Nearctic. The Alaskan distribution of M. broweri is best explained as a reinvasion from the Palearctic, but a Nearctic origin can not be rejected. Several other conventionally recognized species groups can also be rejected. Social evolution has been homoplastic, with large colonial systems evolving in two groups convergently. The cyt b data do not provide unambiguous resolution of several basal nodes in the Palearctic radiation, leaving some aspects of pelage and karyotypic evolution equivocal.     

DNA barcoding as a heuristic tool for classifying undescribed Nearctic Myrmica ants (Hymenoptera: Formicidae)

The Palearctic species of the ant genus Myrmica are well studied. In contrast, the taxonomy of the Nearctic species is outdated, making identification impossible. We collected Myrmica samples in the Holarctic and investigated their diversity using mtDNA data. We analysed a barcode sequence of the Cytochrome Oxidase I gene for 57 Palearctic and 293 Nearctic Myrmica samples. We used sequences of known Palearctic species to search for Myrmica barcode patterns. All but one Palearctic species groups were recovered. The Nearctic diversity was much higher than known. We retrieved the punctiventris, crassirugis and incompleta groups, and established nine additional tentative species groups. Genetic distance analysis revealed a large overlap of intra- and inter-specific distances in Palearctic species and species groups. We could not find a variation gap to separate Nearctic sequences into species with COI data only. Variation in scape morphology divided two genetic groups further. Scape morphology correlated with most molecular groups, except three specimens. Our results illustrate that barcoding, using only a limited amount of genetic information, cannot serve as a universal proxy for taxonomy and species demarcation. It should be considered a first step in understanding the taxonomic diversity of an unknown group of organisms.     

Circumpolar variation in morphological characteristics of Greater White-fronted Geese Anser albifrons

Capsule Greater White-fronted Geese show significant variation in body size from sampling locations throughout their circumpolar breeding range.

Aims To determine the degree of geographical variation in body size of Greater White-fronted Geese and identify factors contributing to any apparent patterns in variation.

Methods Structural measures of >3000 geese from 16 breeding areas throughout the Holarctic breeding range of the species were compared statistically.

Results Palearctic forms varied clinally, and increased in size from the smallest forms on the Kanin and Taimyr peninsulas in western Eurasia to the largest forms breeding in the Anadyr Lowlands of eastern Chukotka. Clinal variation was less apparent in the Nearctic, as both the smallest form in the Nearctic and the largest form overall (the Tule Goose) were from different breeding areas in Alaska. The Tule Goose was 25% larger than the smallest form. Birds from Greenland (A. a. flavirostris) were the second largest, although only slightly larger than geese from several North American populations. Body size was not correlated with breeding latitude but was positively correlated with temperature on the breeding grounds, breeding habitat, and migration distance. Body mass of Greater White-fronted Geese from all populations remained relatively constant during the period of wing moult. Morphological distinctness of eastern and western Palearctic forms concurs with earlier findings of complete range disjunction.

Conclusions Patterns of morphological variation in Greater White-fronted Geese across the Holarctic can be generally attributed to adaptation to variable breeding environments, migration requirements, and phylo-geographical histories.     

Phylogeny and historical biogeography of Agabinae diving beetles (Coleoptera) inferred from mitochondrial DNA sequences

The Agabinae, with more than 350 species, is one of the most diverse lineages of diving beetles (Dytiscidae). Using the mitochondrial genes 16S rRNA and cytochrome oxidase I we present a phylogenetic analysis based on 107 species drawn mostly from the four main Holarctic genera. Two of these genera (Ilybius and Ilybiosoma) are consistently recovered as monophyletic with strong support, Platambus is never recovered as monophyletic, and Agabus is found paraphyletic with respect to several of the species groups of Platambus. Basal relationships among the main lineages are poorly defined, although within each of them relationships are in general robust and very consistent across the parameter space, and in agreement with previous morphological analyses. In each of the two most diverse lineages (Ilybius and Agabus including part of Platambus) there is a basal split between Palearctic and Nearctic clades, estimated to have occurred in the late Eocene. The Palearctic clade in turn splits into a Western Palearctic clade and a clade containing mostly Eastern Palearctic species, and assumed to be ancestrally Eastern Palearctic but with numerous transitions to a Holarctic or Nearctic distribution. These results suggest an asymmetry in the colonization routes, as there are very few cases of transcontinental range expansions originating from the Nearctic or the Western Palearctic. According to standard clock estimates, we do not find any transcontinental shift during the Pliocene, but numerous speciation events within each of the continental or subcontinental regions.     

IN SEARCH OF A METHOD FOR CLADISTIC BIOGEOGRAPHY: AN EMPIRICAL COMPARISON OF COMPONENT ANALYSIS, BROOKS PARSIMONY ANALYSIS, AND THREE-AREA STATEMENTS

Abstract— Three quantitative cladistic biogeographic methods, namely, component analysis, Brooks parsimony analysis (BPA), and three-area statements (TAS) have been proposed for obtaining general area cladograms from taxon-area cladograms. Available programs implementing these methods include COMPONENT versions 1.5 and 2.0 for component analysis, TAS for three-area statements, and Hennig86 for analysing matrices for both three-area statements and BPA. Ten different data sets were analysed with these programs and items of error were used to evaluate the general area cladograms obtained. None of the computer implementations of the methods compared proved to be more effective than the others.
"…all methodologies, even the most obvious ones, have their limits…"
Feyerabend (1993:231)     

Phylogenetic relationships of Holarctic Teleiodini (Lepidoptera: Gelechiidae) based on analysis of morphological and molecular data

Abstract Phylogenetic relationships of 25 genera of Holarctic Teleiodini (Gelechiidae) are postulated based on morphology and molecular characters, including CO‐I, CO‐II, and 28S genes. The phylogenetic analysis of the morphology matrix yielded four equal most‐parsimonious trees (length 330 steps, CI = 0.36, RI = 0.55) and a strict consensus tree (length 335 steps, CI = 0.36, RI = 0.54) with one polytomy and one trichotomy. The phylogenetic analysis of the combined morphology and CO‐I + CO‐II + 28S matrices yielded two equally most‐parsimonious trees (length 1184 steps, CI = 0.50, RI = 0.42) and a strict consensus tree (length 1187 steps, CI = 0.50, RI = 0.42) that reinforced results from the morphological analysis and resolved the one polytomy present in the morphology consensus tree. Teleiodini are defined as a monophyletic clade with a Bremer support value greater than 5 in the consensus tree based on morphological and molecular data. Twenty‐three clades of genera are defined with Bremer support values provided. An analysis of larval host‐plant preferences based on the consensus tree for combined data indicates derivation of feeding on woody hosts from genera feeding on herbaceous hosts and a single origin of feeding on coniferous hosts. An area cladogram indicates five independent origins of Nearctic genera from Holarctic ancestors and one origin from a Palearctic genus.     

Widespread Taxa and Component 2.0

Resolved area cladograms found by heuristic search with COMPONENT 2.0, with the option “map widespread associates” turned off, correspond to Assumption 1 only if the widespread taxon occupies a terminal position on the taxon-area-cladogram. If the widespread taxon occupies a basal position, the resolved area cladograms correspond to Assumption 2.     

Complex biogeographic history of a Holarctic passerine

Our analysis of the ND2 sequences revealed six clades within winter wrens (Troglodytes troglodytes). These clades corresponded to six geographical regions: western Nearctic, eastern Nearctic, eastern Asia, Nepal, Caucasus and Europe, and differed by 3-8.8% of sequence divergence. Differences among regions explained 96% of the sequence variation in winter wren. Differences among individuals within localities explained 3% of the sequence variation, and differences among localities within regions explained 1%. Grouping sequences into subspecies instead of localities did not change these proportions. Proliferation of the six clades coincided with Early and Middle Pleistocene glaciations. The distribution of winter wren clades can be explained by a series of five consecutive vicariant events. Western Nearctic wrens diverged from the Holarctic ancestor 1.6 Myr before the present time (MYBP). Eastern Nearctic and Palaearctic wrens diverged 1 MYBP. Eastern and western Palaearctic birds diverged 0.83 MYBP. Nepalese and east Asian wrens diverged 0.67 MYBP, and Caucasian birds diverged from European wrens 0.54 MYBP. The winter wren has a much greater degree of inter- and intracontinental differentiation than the three other Holarctic birds studied to date--dunlin (Calidris alpina), common raven (Corvus corax) and three-toed woodpecker (Picoides trydactylus)--and represents an example of cryptic speciation that has been overlooked.     

Plio-Pleistocene climatic oscilations, Holarctic biogeography and speciation in an avian subfamily

Aim In this paper, I discuss the temporal and spatial aspects of historical biogeography and speciation in a widely distributed Holarctic subfamily of birds (Tetraoninae). Location Northern Holarctic. Results Using dated fossils, I calibrated the molecular clock for the mitochondrial control region at 7.23 ± 1.58% nucleotide divergence (maximum likelihood corrected) per million years. The data suggest that grouse (Tetraoninae) originated in the Middle Pliocene, 6.3 Ma. Grouse apparently originated in the northern part of western Nearctic, and Palearctic was colonized independently three times, first by the ancestor of all grouse in the Middle Pliocene, then by the ancestor of forest (Falcipennis, Tetrao and Lyrurus) and prairie (Centrocercus, Dendragapus and Tympanuchus) grouse in the Late Pliocene, and finally by the ancestral Lagopus in the Early Pleistocene. Only once Nearctic was colonized from Palearctic by a common ancestor of forest grouse. Sympatry and range symmetry were positively correlated with molecular divergence. These correlations suggest that peripatric isolation was the predominant mode of speciation throughout grouse history. Main conclusions Speciation events in grouse were driven by climatic oscillations of the Pliocene and Pleistocene. Isolation of small peripheral populations from widely distributed ancestors was the dominant mode of speciation in grouse. Isolations during interglacials both across Beringia, and in southern mountain areas when boreal habitats were restricted to high elevations, suggest an important role for vicariance in grouse speciation.     

Phylogeny of the vipers of the Caucasus (Reptilia, Viperidae)

Phylogenetic relationships for five taxa of Palearctic vipers (genus Vipera ) from the Caucasian region were revealed by cladistic analyses of separate and combined morphological and biochemical characters. The different data sets yielded largely congruent cladograms. Vipera berus from Sweden was included as an ingroup and V. aspis was used for outgroup comparison. For V. kaznakovi and V. dinniki , three and four different sub-populations, respectively, were treated as independent terminal taxa in the analyses. The most parsimonious cladograms confirmed the systematic positions of these populations, discussed in a recent study, and support the hypothesis that the montane populations of the western main Caucasus comprise one polymorphie species: V. dinniki.
Analyses of combined biochemical and morphological data generated two equally parsimonious cladograms (for all ingroups compared), but yielded only one fully resolved topology when ingroups were condensed to the species level: ( berus ((renardi ('ursinii'-eriwanensis )))( dinnikikaznakovi )).     

The role of chromosomal polymorphism in divergence of populations and species of the genus Chironomus (Diptera)

The data on the structure and level of chromosomal polymorphism in natural populations of species of the genus Chironomus are summarized. A very high level of chromosomal polymorphism was noted for most species. Paracentric inversions prevailed among the chromosomal rearrangements found in natural populations. Changes in the set and frequency of inversion sequences are the most important factor of cytogenetic divergence of populations. Several cytogenetic types of populations were distinguished. The Palaearctic and Nearctic populations of Holarctic species diverged to a greater extent due to the formation of endemic Palearctic and Nearctic inversion sequences. The sequences common for both regions indicated a common ancestry of the populations. The cytogenetic distances between the Palearctic and Nearctic populations are greater by an order of magnitude than those between populations within each zoogeographic region. Divergence of species karyotypes was found to result from fixation of different inversion sequences in the course of evolution. The karyotypes of Palearctic and Nearctic species mainly differ by the presence of endemic Palearctic and Nearctic banding sequences. Several basic sequences common for some species allow the cytogenetic history of their origin to be revealed. A NJ phylogenetic tree was built for the genus Chironomus, demonstrating chromosomal evolution of its species.     

Phylogeny and historical biogeography of the cynipoid wasp family Ibaliidae (Hymenoptera)

The Ibaliidae are a small family of cynipoid wasps, the members of which parasitize woodboring siricid larvae in hardwoods and conifers. The 19 currently recognized extant species occur mainly in the Northern Hemisphere. No fossils are known despite the presumed old age of the family. We present a cladistic analysis of ibaliid relationships at the species-level, mainly based on external skeletal characters of adults. The results indicate that the three genera ( Eileenella , Heteribalia , Ibalia ) and two subgenera of Ibalia ( Ibalia s. str. and Tremibalia ) recognized in the current classification are monophyletic. Three different categories of characters were compared for their phylogenetic usefulness. Homoplasy was found to be lowest for main structures, higher for sculptural characters, and still higher for colour differences. The historical biogeography of the family was reconstructed using dispersal–vicariance analysis in combination with palaeogeographical data. The results suggest that the family primarily diversified within the eastern Palaearctic–northern Oriental region. The nominate subgenus of Ibalia dispersed early to the western Nearctic, where it radiated; two species later spread throughout the Holarctic. The other subgenus of Ibalia shows an early eastern Palaearctic–eastern Nearctic disjunction which presumably dates back to the Eocene–Oligocene transition.     

Molecular phylogeny of the speciose vole genus Microtus (Arvicolinae, Rodentia) inferred from mitochondrial DNA sequences

Voles of the genus Microtus represent one of the most speciose mammalian genera in the Holarctic. We established a molecular phylogeny for Microtus to resolve contentious issues of systematic relationships and evolutionary history in this genus. A total of 81 specimens representing ten Microtus species endemic to Europe as well as eight Eurasian, six Asian and one Holarctic species were sequenced for the entire cytochrome b gene (1140 bp). A further 25 sequences were retrieved from GenBank, providing data on an additional 23, mainly Nearctic, Microtus species. Phylogenetic analysis of these 48 species generated four well-supported monophyletic lineages. The genus Chionomys, snow voles, formed a distinct and well-supported lineage separate from the genus Microtus. The subgenus Microtus formed the strongest supported lineage with two sublineages displaying a close relationship between the arvalis species group (common voles) and the socialis species group (social voles). Monophyly of the Palearctic pitymyid voles, subgenus Terricola, was supported, and this subgenus was also subdivided into two monophyletic species groups. Together, these groupings clarify long-standing taxonomic uncertainties in Microtus. In addition, the "Asian" and the Nearctic lineages reported previously were identified although the latter group was not supported. However, relationships among the main Microtus branches were not resolved, suggesting a rapid and potentially simultaneous radiation of a widespread ancestor early in the history of the genus. This and subsequent radiations discernible in the cytochrome b phylogeny, show the considerable potential of Microtus for analysis of historical and ecological determinants of speciation in small mammals. It is evident that speciation is an ongoing process in the genus and that the molecular data provides a vital insight into current species limits as well as cladogenic events of the past.     

Linkage Groups of Protein-Coding Genes in Western Palearctic Water Frogs Reveal Extensive Evolutionary Conservation

Among progeny of a hybrid (Rana shqiperica X R. lessonae) X R. lessonae, 14 of 22 loci form four linkage groups (LGs): (1) mitochondrial aspartate aminotransferase, carbonate dehydratase-2, esterase 4, peptidase D; (2) mannosephosphate isomerase, lactate dehydrogenase-B, sex, hexokinase-1, peptidase B; (3) albumin, fructose-biphosphatase-1, guanine deaminase; (4) mitochondrial superoxide dismutase, cytosolic malic enzyme, xanthine oxidase. Fructose-biphosphate aldolase-2 and cytosolic aspartate aminotransferase possibly form a fifth LG. Mitochondrial aconitate hydratase, α-glucosidase, glyceraldehyde-3-phosphate dehydrogenase, phosphogluconate dehydrogenase, and phosphoglucomutase-2 are unlinked to other loci. All testable linkages (among eight loci of LGs 1, 2, 3, and 4) are shared with eastern Palearctic water frogs. Including published data, 44 protein loci can be assigned to 10 of the 13 chromosomes in Holarctic Rana. Of testable pairs among 18 protein loci, agreement between Palearctic and Nearctic Rana is complete (125 unlinked, 14 linked pairs among 14 loci of five syntenies), and Holarctic Rana and Xenopus laevis are highly concordant (125 shared nonlinkages, 13 shared linkages, three differences). Several Rana syntenies occur in mammals and fish. Many syntenies apparently have persisted for 60-140 X 10(6) years (frogs), some even for 350-400 X 10(6) years (mammals and teleosts).     

Chironomidae of the Holarctic region: a comparison of ecological and functional traits between North America and Europe

Chironomidae (Diptera) are widespread, abundant, diverse and ubiquitous, and include genera and species that are distributed across the Holarctic region. However, the geographical barriers between continents should have resulted in intraspecific population differentiation with reflection on individual biological and ecological traits. Our aim was to test for potential differences in Chironomidae species/genus and traits between the Nearctic and Palearctic regions. We compared the Chironomidae trait information gathered in two databases; one database was developed in Europe and the other in North America. Common genera and species of both databases were selected, and the common traits were adjusted into the same trait categories. Data were transformed into presence/absence and divided into Eltonian (biological/functional) and Grinnellian (ecological) traits. Common genera and common species were analyzed using Fuzzy correspondence analysis (FCA). Differences between databases occur for all trait domains. Yet, Eltonian traits showed lower level of concordance than Grinnellian traits at the species level. Different biological characteristics in the Nearctic and Palearctic regions may indicate that Chironomidae have different adaptions to similar ecological environments due to intraspecific variability or even trait plasticity.     

Worldwide distribution and theoretical spreading of Trichoferus campestris (Coleoptera: Cerambycidae) depending on the main climatic elements

The velvet longhorned beetle, Trichoferus campestris (Coleoptera: Cerambycidae) is a serious wood‐boring pest that is a major threat to the phytosanitary condition of forests and orchards. Its worldwide expansion is a major concern for plant health. We have collected all bibliographical references, phytosanitary reports and authentic photographic evidence from entomological websites to determine the worldwide distribution of T. campestris. The theoretical directions of the spreading and actually occupied area of this arthropod pest were determined over the whole Holarctic range. Furthermore, the potential distribution area was calculated using cumulated temperature in the growing season averaged over 15 years both in the Palearctic and Nearctic regions. Holarctic expansion of the species, including the main parts of North America and Eurasia, is clearly indicated. Its populations occur in 29 countries to date, supported by documentation from 64 publications and 30 online forums. Its spread is continuously westward in the Palearctic; in the Nearctic, the spread was first southward from the Great Lakes region then eastward from Utah. The species has excellently adapted to circumstances of freight by ship and plane, as wood is often used, ensuring optimal conditions for the pest. In addition, the active spreading achieved by flying is an important factor contributing to its expansion. The primary criterion for controlling the species would be the introduction of a monitoring system in affected and exposed areas.     

An evaluation of the influence of environment and biogeography on community structure: the case of Holarctic mammals

Aim To evaluate the influence of environment and biogeographical region, as a proxy for historical influence, on the ecological structure of Holarctic communities from similar environments. It is assumed that similarities among communities from similar environments in different realms are the result of convergence, whereas their differences are interpreted as being due to different historical processes. Location Holarctic realm, North America and Eurasia above 25° N. Methods Checklists of mammalian species occurring in 96 Holarctic localities were collected from published sources. Species were assigned to one of 20 functional groups defined by diet, body size and three‐dimensional use of space. The matrix composed of the frequencies of functional groups in the 96 localities is used as input data in a correspondence analysis (CA). The localities are classified into nine groups according to Bailey's ecoregions (used as a surrogate of regional climate), and the positions of the communities in the dimensions of the CA are compared in relation to ecoregion and realm. Partial regression was used to test for the relative influence of ecoregion and realm over each dimension and to evaluate the effect of biogeographical realm on the variation in the factor scores of the communities of the same ecoregion. Results In some cases, mammalian communities from areas with similar regional climates exhibit convergence in community structure, irrespective of the biogeographical realm where they are located. However, all of them are clearly subdivided into Nearctic and Palearctic subsets. Differences in the composition of the regional pools only partially explain differences in local communities between realms. Main conclusions Holarctic mammalian communities from regions with widely different climates differ in ecological structure irrespective of their biogeographical location. On the other hand, the structures of Nearctic and Palearctic communities from regions of similar climate radically differ in some features. Thus, although present climatic conditions influence community structure, contingent historic processes associated with each region also play a major role in determining community structure.     

Molecular and morphological evidence for the Holarctic distribution of Urogonimus macrostomus (Rudolphi, 1803) Monticelli, 1888 (Digenea: Leucochloridiidae)

Species of Urogonimus Monticelli, 1888 (Leucochloridiidae Poche, 1907) are difficult to distinguish using adult morphology, and their taxonomy has been repeatedly subjected to revision. Some Nearctic species have been regarded as synonymous with the Palearctic type species Urogonimus macrostomus (Rudolphi, 1803) Monticelli, 1888. This implies that U. macrostomus is present in the Nearctic, but there is no additional evidence for this putative distribution. We collected trematodes morphologically indistinguishable from U. macrostomus from a house sparrow (Passer domesticus) in Edmonton, Alberta, Canada. Sequences 2958 bp in total length from the small and large subunits of ribosomal DNA from 2 specimens were 99.8-100% identical to those of U. macrostomus in the Ukraine and Japan. In light of the lack of morphological differences and small degree of genetic variation, we consider the specimens we collected to be conspecific with U. macrostomus in the Palearctic, and the Holarctic range of the species is thus supported. Sequences from a more rapidly evolving gene, cytochrome c oxidase 1, were obtained to aid future study of this and related species.     

Evolutionary implications of morphological variation in the lower carnassial of red fox<Emphasis Type="Italic">Vulpes vulpes</Emphasis>

Research on the morphological variability of the occlusal surface of M₁ talonid in the red foxVulpes vulpes (Linnaeus, 1758) in the Holarctic has been carried out on 2271 specimens originating from 42 populations. The Nearctic was represented by 666 specimens belonging to 13 populations, whereas Palearctic was represented by 1605 specimens from 29 populations. Analyses of the developmental level and formation of cristids between the hypoconid and entoconid allowed the differentiation of 34 shape variants of the occlusal surface of the talonid in the red fox. Because of the complicated variation of cristids, 34 variants were assigned to 5 morphotypes of group P. In the Palearctic and Nearctic a significant geographic variation occurred of P morphotypes and their variants. Primitive variants of the talonid structure on M₁ are predominant in populations from the south of the Asian range of the red fox, while more progressive characters of the occlusal surface of the lower carnassial are typical of the northern and centrally located red fox populations in the Palearctic and Nearctic. The geographic differentiation is probably connected with different Pleistocene histories of particular populations.