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[Taxonomy of ticks of the genus Microtimyobia (Acariformes: Myobiidae: Radfordia) and their distribution on voles (Rodentia: Cricetidae: Arvicolinae)].

A phylogenetic system of the subgenus Microtimyobia (Myobiidae: Radfordia) elaborated for the first time by means of the software HENNIG-86 is proposed. The subgenus Microtimyobia including three species groups, lemnina, hylandi and zibethicalis, was divided for a cladistic analysis into 6 operation units based mainly on a male genital shield structure. The analysis shows, that the zibethicalis group is a polyphyletic, while the hylandi and lemnina groups are monophyletic. Host-parasite associations of myobiid mite taxa with vole taxa (Arvicolinae) and some peculiarities of mite and host taxa distribution are analysed. The zibethicalis group is represented by two species, associated with the North American rodents of the genera Ondatra and Phenacomys respectively. The hylandi group are most widely distributed among Arvicolinae taxa, both in Eurasia and North America. However, R. hylandi occurs on those vole species of the genera Microtus and Pitymys, which are distributed only in central and southern parts of North America and represent descendants of the earlier migration wave of Microtus from the Eurasia. As far as R. hylandi is also found in the pleistocenus of the Yakutia, that means that its areal was wider than in recent period. The lemnina group lives on hosts of 2 subtribes of the tribe Arvicolini (Arvicolina and Clethrionomyina) and is restricted to Eurasian range, except R. lemnina. This species is also mainly distributed on Eurasian vole species, however it occurs on vole species distributed in Alaska and being decendants of the later wave of Microtus migration. As far as R. lemnina and R. hylandi are associated with representatives of different migration waves of Microtus from the Euroasia, it is suggested that mites of the hylandi group are the original myobiid fauna of the Microtus voles. The species of the lemnina group had apparently originated on voles of the subtribe Clethrionomyina and then migrated onto phylogenetically young hosts of the subtribe Arvicolina (Euroasian species of the Microtus and related genera), where they probably eliminated mites of the hylandi group from these hosts in Euroasia. The recent pattern of myobiid species distribution on vole species is a result of both a mite cospeciation with their hosts and a shift of hosts. Five new myobiid mite species are described and distinguished by characters as follows. R. (M.) abramovi sp. n. from Phodopus roborovskii (Cricetidae) is closely related to R. (M.) triton Fain et Lukoschus, 1977. In both sexes of the new species setae cxI 1, 2 are scale-shaped, while in R. (M.) triton these setae (cxI 1, 2) are hair-like. R. (M.) stekolnikovi sp. n. from Chionomys nivalis (type host) and Ch. gud is similar to R. (M.) lemnina (Koch, 1841). Females of new species have setae ra with 2 apical processes; female tritonymphs with long whip-like setae ic4. R. (M.) lemnina females have setae ra with 3 processes; female tritonymphs with short hair-like setae ic4. R. (M.) stenocrani sp. n. from Microtus gregalis is also closely related to R. (M.) lemnina. Females of the new species have setae ra with 2 processes; female tritonymphs with whip-like setae ic3. R. (M.) lemnina females have seate ra with 3 processes, female tritonymph with short hair-like setae ic3. R. (M.) synaptomysi sp. n. (ABSTRACT TRUNCATED)

Animals↗

The evolutionary radiation of Arvicolinae rodents (voles and lemmings): relative contribution of nuclear and mitochondrial DNA phylogenies.

BACKGROUND: Mitochondrial and nuclear genes have generally been employed for different purposes in molecular systematics, the former to resolve relationships within recently evolved groups and the latter to investigate phylogenies at a deeper level. In the case of rapid and recent evolutionary radiations, mitochondrial genes like cytochrome b (CYB) are often inefficient for resolving phylogenetic relationships. One of the best examples is illustrated by Arvicolinae rodents (Rodentia; Muridae), the most impressive mammalian radiation of the Northern Hemisphere which produced voles, lemmings and muskrats. Here, we compare the relative contribution of a nuclear marker--the exon 10 of the growth hormone receptor (GHR) gene--to the one of the mitochondrial CYB for inferring phylogenetic relationships among the major lineages of arvicoline rodents. RESULTS: The analysis of GHR sequences improves the overall resolution of the Arvicolinae phylogeny. Our results show that the Caucasian long-clawed vole (Prometheomys schaposnikowi) is one of the basalmost arvicolines, and confirm that true lemmings (Lemmus) and collared lemmings (Dicrostonyx) are not closely related as suggested by morphology. Red-backed voles (Myodini) are found as the sister-group of a clade encompassing water vole (Arvicola), snow vole (Chionomys), and meadow voles (Microtus and allies). Within the latter, no support is recovered for the generic recognition of Blanfordimys, Lasiopodomys, Neodon, and Phaiomys as suggested by morphology. Comparisons of parameter estimates for branch lengths, base composition, among sites rate heterogeneity, and GTR relative substitution rates indicate that CYB sequences consistently exhibit more heterogeneity among codon positions than GHR. By analyzing the contribution of each codon position to node resolution, we show that the apparent higher efficiency of GHR is due to their third positions. Although we focus on speciation events spanning the last 10 million years (Myr), CYB sequences display highly saturated codon positions contrary to the nuclear exon. Lastly, variable length bootstrap predicts a significant increase in resolution of arvicoline phylogeny through the sequencing of nuclear data in an order of magnitude three to five times greater than the size of GHR exon 10. CONCLUSION: Our survey provides a first resolved gene tree for Arvicolinae. The comparison of CYB and GHR phylogenetic efficiency supports recent assertions that nuclear genes are useful for resolving relationships of recently evolved animals. The superiority of nuclear exons may reside both in (i) less heterogeneity among sites, and (ii) the presence of highly informative sites in third codon positions, that evolve rapidly enough to accumulate synapomorphies, but slow enough to avoid substitutional saturation.

Animals↗

Comparative functional morphology of mandibular forward movement during mastication of two murid rodents, Apodemus speciosus (Murinae) and Clethrionomys rufocanus (Arvicolinae).

The anatomy of the masticatory apparatus, the direction in which masticatory muscles act during mastication, and jaw muscle forces as estimated by muscle dry weight are compared between two murid rodents, the Japanese field mouse (Apodemus speciosus, subfamily Murinae) and the gray red-backed vole (Clethrionomys rufocanus; subfamily Arvicolinae). The occlusal forces exerted by the deep masseter and the anterior temporalis are large in C. rufocanus. Furthermore, in this species, the angle between the sagittal plane and the occlusal plane of the cheek teeth is larger than in A. speciosus. Therefore, a relatively large occlusal force can be generated in C. rufocanus. The estimated line of action of the anterior temporalis differs markedly between these two species. The functional significance of this difference is discussed relative to the adaptive dental characteristics for food processing, the forces required to masticate different types of food, and the forces that control mandibular forward movement.

Animals↗

Duplication, balancing selection and trans-species evolution explain the high levels of polymorphism of the DQA MHC class II gene in voles (Arvicolinae).

Major histocompatibility complex (MHC) genes play important role in host-parasite interactions and parasites are crucial factors influencing the population dynamics of hosts. We described the structure and diversity of exon 2 of the MHC class II DQA gene in three species of voles (Arvicolinae) exhibiting regular multi-annual fluctuations of population density and analysed the processes leading to the observed MHC polymorphism. By using cloning-sequencing methodology and capillary electrophoresis-single strand conformation polymorphism, we described seven sequences in the water, eight in the common, and seven in the bank voles coming from an area of 70 km(2) around the Nozeroy canton in the Jura Mountains (Franche Comté, France). All exon 2 sequences translate to give unique amino acid sequences and positive selection was found to act very intensively on antigen binding sites. We documented the presence of recombination at vole DQA region but its importance in generating allelic polymorphism seems to be relatively limited. For the first time within rodents, we documented the duplication of the DQA gene in all three species with both copies being transcriptionally active. Phylogenetic analysis of allelic sequences revealed extensive trans-species polymorphism within the subfamily although no alleles were shared between species in our data set. We discuss possible role of parasites in forming the recent polymorphism pattern of the DQA locus in voles.

Alleles↗

Description and morphometric variability of Paranoplocephala serrata n. sp. (Cestoda: Anoplocephalidae) in collared lemmings (Dicrostonyx spp., Arvicolinae) from Arctic Siberia and North America.

We describe Paranoplocephala serrata n. sp. (Cestoda, Anoplocephalidae) from collared lemmings Dicrostonyx torquatus and D. groenlandicus (Arvicolinae, Rodentia) in Arctic Siberia and North America. The new species was recorded from the Yamal Peninsula (type-locality), Yana Delta, Kolyma Delta, Wrangel Island, Alaska and Victoria Island/Kent Peninsula (Northwest Territories). P. serrata n. sp. is characterised by a long, ribbon-like strobila, distinctly serrated segments, a small scolex, unilateral or infrequently alternating genital pores and testes confined to the antiporal part of the segment. It differs from the related species (Andrya bairdi, Parandrya feodorovi and Paranoplocephala manseri) by several morphological features, including the distribution of testes (several testes antiporal to ventral longitudinal osmoregulatory canal), structure of the cirrus-sac and vagina, and large eggs (0.053-0.068 mm in the type-material). The material of P. serrata n. sp. from North America differs from the Siberian material by the shorter cirrus-sac, smaller dimensions of the female reproductive organs, larger seminal receptacle and larger eggs. However, the statistical differences in the dimensions of reproductive organs mainly reflect the larger size of mature segments in Siberian specimens compared with North American specimens. The main diagnostic features, i.e. the size and form of scolex and suckers, number and distribution of testes, position of female glands, vagina/cirrus-sac ratio and morphology of reproductive organs, do not differ markedly between the Palaearctic and Nearctic specimens. According to the structure of the early-stage uterus, A. bairdi Schad, 1954 belongs to the genus Paranoplocephala. Parandrya Gulyaev & Chechulin, 1996 is probably a synonym of Paranoplocephala. A redescription is provided for Paranoplocephala bairdi n. comb.

Animals↗

Biogeography of helminth parasitism in Lemmus Link (Arvicolinae), with the description of Paranoplocephala fellmani n. sp. (Cestoda: Anoplocephalidae) from the Norwegian lemming L. lemmus (Linnaeus).

We describe the gastrointestinal helminth fauna of true lemmings (Lemmus spp., Arvicolinae) based on published and original material throughout the Holarctic range of these hosts. According to the existing data, the helminth fauna of true lemmings consists of three widespread and/or locally common taxa: Hymenolepis horrida (sensu lato) (Hymenolepididae), Anoplocephaloides lemmi (Anoplocephalidae) and Heligmosomoides spp. (Heligmosomidae). Despite the taxonomic boundaries and ancient phylogenetic splits in the hosts, there are no major faunistic differences for parasites within western (Siberian) L. sibiricus and L. bungei, and eastern (North American) L. trimucronatus throughout their distribution range. In contrast, the Norwegian lemming L. lemmus, which is a Fennoscandian endemic and closely related to the western populations of L. sibiricus, has only a single host-specific helminth, the cestode Paranoplocephala fellmani n. sp. (Anoplocephalidae). We describe the new species and show that it differs consistently from related species by its long and slender cirrus-sac. However, there are also a number of other significant differences, e.g., P. fellmani n. sp. and Andrya primordialis in Tamiasciurus hudsonicus (Sciuridae) evidently have a unique (sub)type of uterine development among Andrya/Paranoplocephala spp. Because P. fellmani n. sp. was also found to occur in Alaska (host L. trimucronatus), this species seems to follow the same biogeographical pattern as the other specialist helminths of Lemmus. We suggest alternative explanations for the absence of three major helminth taxa in the Norwegian lemming in Fennoscandia.

Animals↗

Morphological characterisation of Paranoplocephala bairdi (Schad, 1954) (Cestoda: Anoplocephalidae) in heather voles Phenacomys spp. and tree voles Arborimus spp., and related species in voles and lemmings (Muridae: Arvicolinae).

The taxonomical status of Paranoplocephala bairdi (Schad, 1954)-like cestodes (Anoplocephalidae) in heather voles Phenacomys spp. and tree voles Arborimus spp. (Muridae: Arvicolinae) and their discrimination from five related species of Paranoplocephala is assessed using uni- and multivariate morphometrics. The analyses support the independent status and conspecificity of specimens from Phenacomys spp. and Arborimus spp., and P. bairdi is therefore suggested to be a host-specialist species of heather and tree voles with a wide geographical distribution in North America. A redescription is presented for P. bairdi.

Analysis of Variance↗

Characterization of the satellite DNA Msat-160 from the species Chionomys nivalis (Rodentia, Arvicolinae).

The satellite DNA Msat-160 has been previously characterized in several species of the genus Microtus. Here we present the characterization of Msat-160 from Chionomys nivalis, a species with a very primitive karyotype. As in other Microtus species analyzed, C. nivalis Msat-160 is AT rich, has a monomer length of 160 bp, is undermethylated and is mainly located in all the pericentromeric heterochromatin of all autosomes and the X chromosome, but is completely absent from the Y chromosome. Hence, our results support the hypothesis that Msat-160 was initially distributed in the pericentromeric heterochromatin of all autosomes and the X chromosome. The taxonomic status of the genus Chionomys in relation to the genus Microtus is a very interesting issue, so we constructed phylogenetic dendrograms using Msat-160 sequences from several Microtus species. Although the results were not informative about this issue, the presence of Msat-160 in C. nivalis and Microtus species suggested that both genera are closely related and that this satellite DNA was present in the common ancestor. Studies of Msat-160 in different arvicoline species could help to determine the origin of this satellite and, perhaps, to establish the phylogenetic relationships of some arvicoline groups.

Animals↗

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.

Animals↗

Historical biogeography at the crossroads of the northern continents: molecular phylogenetics of red-backed voles (Rodentia: Arvicolinae).

Evolutionary relationships of red-backed voles and their relatives were examined and used to test biogeographic hypotheses. Sequences of the mitochondrial cytochrome b gene were obtained for 25 individuals representing Alticola macrotis, Clethrionomys californicus, C. gapperi, C. glareolus, C. rutilus, and C. rufocanus. These were combined with 21 partial sequences from GenBank for C. regulus, C. rex, C. rufocanus, C. rutilus, Eothenomys imaizumii, E. melanogaster, Phaulomys andersoni, and P. smithii. Complete sequences of three species of Microtus (M. montanus, M. oeconomus, and M. pennsylvanicus), representative species of other arvicoline genera (Myopus, Synaptomys, Arvicola, Ellobius, Ondatra, Lemmus, Dicrostonyx, and Phenacomys), and a sigmodontine representative (Peromyscus) were included as outgroups. We used maximum parsimony, maximum likelihood, distance, and Bayesian based methods and conducted statistical tests on proposed hypotheses of phylogenetic relationships and biogeographic histories. A close relationship of species representing the genera Alticola, Clethrionomys, and Eothenomys was supported (Clethrionomyini); however, the genus Clethrionomys was paraphyletic with respect to both Alticola and Eothenomys. Three major clades were identified as Asian (Eothenomys andersoni, E. smithii, C. rex, C. regulus, and C. rufocanus), Trans-beringian (Alticola macrotis, C. californicus, C. gapperi, C. glarelolus, and C. rutilus), and Taiwanese (E. melanogaster). These results are consistent with the fossil record which indicates an initial diversification in Asia followed by colonization of the Nearctic on at least two occasions. The holarctic species, C. rutilus, appears to have either reinvaded Asia from North America or colonized North America more recently (late Pleistocene) than the two species of Clethrionomys (C. gapperi and C. californicus) that are endemic to North America (early to mid-Pleistocene). Finally, C. gapperi, appears to be comprised of an eastern and a western species, the former with affinities to the Asian C. glareolus and the latter more closely related to C. californicus.

Animals↗

First report of Siphonaptera infesting Microtus (Microtus) cabrerae (Rodentia-Muridae-Arvicolinae) in Cuenca, Spain and notes about the morphologic variability of Ctenophthalmus (Ctenophthalmus) apertus personatus (Insecta-Siphonaptera-Ctenophthalmidae).

The fleas infesting Microtus (Microtus) cabrerae from three different areas of Cuenca province (Spain) have been studied. It is the first time that on ectoparasitological study of this badly known rodent has been done. Four Siphonaptera species have been detected: Rhadinopsylla (Actenophthalmus) pentacantha, Peromyscopsylla spectabilis spectabilis, Nosopsyllus fasciatus and Ctenophthalmus (Ctenophthalmus) apertus personatus, which was the most abundant species (26 males and 31 females of a total of 28 males and 35 females). Considering the great morphologic variability within the male processus basimerus ventralis (p.b.v.) of segment IX of C. personatus subspecies, three morphotypes have been recognised. The male polymorphism detected, would be the result of both host confinement and genetic selection acting on the parasite. It should be pointed out that C. (C.) apertus personatus is not narrowly host-specific, therefore further studies are required to clarify this taxonomic situation.

Animals↗

Artificial life and speciation, a case study: heterochromatin and speciation in the Microtus savii Group (Rodentia-Arvicolinae).

Artificial life is a tool which is used for simulation of peculiar cases of evolutionary events. The main characteristic of artificial life is that with this technique it is possible to simulate for a high number of generations the evolution of a population of individuals. Each individual is characterised by a small number of parameters, but each individual has its own evolutive story. So far it is possible to simulate the evolution of a population of some thousands specimens, for a high number of generations. The realistic aspect of the simulation is that each specimen is taken individually. In our opinion this instrument is very useful to simulate the evolution of the hybrids barrier during speciation. For this reason it is applied to a peculiar case of speciation, that of the Savi pine vole (Microtus savii) whose experimental data were recently investigated.

Animals↗

[The taxonomic status of the Tuva vole Alticola (A.) tuvinicus Ognev (Mammalia: Arvicolinae)].

Some forms united under species Alticola tuvinicus s. lato (4 samples), A. semicanus (1 sample), A. argentatus (8 samples) are compared by 16 characters by means of principal component method and canonical analysis. Species distinctness of A. semicanus is confirmed. Specific status of the form olchonensis is supposed. The results concerning tuvinicus s. str. and kosogol are not clear-cut: these forms may be well differentiated subspecies within either A. tuvinicus s. lato or A. argentatus.

Animals↗

[A rare case of chromosome mutation in the ground vole Terricola majori (Arvicolinae, Rodentia)].

A rare case of chromosome variability is described for the ground vole, Terricola majori, from the Lagodekhi Nature Reserve (Georgia). This species is characterized by a stable karyotype (2n = 54, NF = 60), but we found an animal in which 2n = 53, NF = 60. The change in the diploid number resulted from a Robertsonian translocation accompanied by a local amplification of satellite sequences in centrometric heterochromatin. This structural rearrangement is probably a neutral mutation and is described for forest T. majori voles for the first time.

Animals↗

Molecular phylogeny of European muroid rodents based on complete cytochrome b sequences.

Phylogenetic relationships among 18 species of mainly European muroid rodents that belong to three subfamilies were estimated using complete sequences of the mitochondrial cytochrome b gene. The inferred monophyly of the subfamilies Murinae (mice and rats) and Arvicolinae (voles, lemmings, and muskrats) is in agreement with previous studies. Within the Murinae, the morphology-based division of the genus Apodemus into three subgenera is supported by these DNA sequence data. The relationships among the different genera of the Murinae were generally poorly resolved, and the relationships of Micromys and Acomys to the other murine genera remained unresolved. Within the subfamily Arvicolinae, the relations of the genera Arvicola, Clethrionomys, and Microtus remained tentative with our data. However, within the Microtus group, there is a good molecular support for the phylogenetic relationships. These findings suggest that the origin of the different murine and arvicoline lineages was rapid, indicating an adaptive radiation with fast speciation.

Animals↗

Relationships of the chromosomal species in the Eurasian mole rats of the Spalax ehrenbergi group as determined by DNA-DNA hybridization, and an estimate of the spalacid-murid divergence time.

DNA-DNA hybridization was used to measure the average genomic divergence among the four chromosomal species of the Eurasian mole rats belonging to the Spalax ehrenbergi complex (Rodentia: Spalacidae). The percent nucleotide substitutions in the single-copy nuclear DNA among the species ranged from 0 to 5%, suggesting that speciation has occurred with minor genomic changes in these animals. The youngest chromosomal species appear to differ by 0.2-0.6% base pair mismatch, which is only between one and three base differences in a 500-bp fragment. The interspecific values of percent nucleotide differences permit the recognition of two well-separated speciation events in the S. ehrenbergi complex, the older (of Lower Pleistocene age) having isolated the chromosomal species 2n = 54 before the divergence of the three other species. DNA-DNA hybridization was also used to compare the Spalacinae (Eurasian mole rats), Murinae (Old World rats and mice), and Arvicolinae (voles and lemmings). These data enabled us to estimate the time of divergence of the spalacids at ca. 19 million years ago. The dates of divergence among the other rodent lineages, as predicted by DNA hybridization results, agree well with paleontological data. These dates of divergence are obtained by the relation between geological time and single-copy nuclear DNA change, a relation that was calibrated by Catzeflis et al. (1987) through the use of fossil Arvicolinae and Murinae data.

Animals↗

Evolutionary diversification of protein-coding genes of hantaviruses.

Phylogenetic analyses of the S:, M, and L: genes of the hantaviruses (Bunyaviridae: Hantavirus) revealed three well-differentiated clades corresponding to viruses parasitic on three subfamilies (Murinae, Arvicolinae, and Sigmodontinae) of the rodent family Muridae. In rooted trees of M: and L: genes, the viruses with hosts belonging to Murinae formed an outgroup to those with hosts in Arvicolinae and Sigmodontinae. This phylogeny corresponded with a phylogeny of the murid subfamilies based on mitochondrial cytochrome b sequences, supporting the hypothesis that hantaviruses have coevolved with their mammalian hosts at least since the common ancestor of these three subfamilies, which probably occurred about 50 MYA. The nucleocapsid protein (encoded by the S: gene) differentiated among the viruses parasitic on the three subfamilies in such a way that a high frequency of amino acid residue charge changes occurred in a hypervariable (HV) portion of the molecule, and nonsynonymous nucleotide differences causing amino acid charge changes in the HV region occurred significantly more frequently than expected under random substitution. Along with evidence that at least in some hantaviruses the HV region is a target for host antibodies and the known importance of charged residues in determining antibody epitopes, these results suggest that changes in the HV region may represent adaptation to host-specific characteristics of the immune response.

Animals↗