Molecular identification of three sympatric species of wood mice (Apodemus sylvaticus,A. flavicollis,A. alpicola) in western Europe (Muridae: Rodentia)

The woodmouse (Apodemus sylvaticus) and yellow‐necked fieldmouse (Apodemus flavicollis) are sympatric and even syntopic in many regions throughout their European range. Their field discrimination on the basis of external characters is a real challenge for many fields of research. The problem is even more complicated in the Alpine chain where they live sympatrically with a third similar species: A. alpicola. A rapid and simple method is proposed to discriminate the three species in processing field‐collected biopsies as well as ethanol‐preserved museum samples.     

Comparative study of the phylogenetic structure in six Apodemus species (Mammalia,Rodentia) inferred from ISSR-PCR data

The utility of the Inter Simple Sequence Repeat-Polymerase Chain Reaction (ISSR-PCR) was explored in order to determine genetic variation in six species of the genus Apodemus (A. flavicollis, A. sylvaticus, A. uralensis, A. agrarius, A. mystacinus and A. epimelas) at the individual level, population level, in separate geographic samples and in the species as a whole. Six optimized primers produced highly reproducible and polymorphic DNA markers with 98.3% polymorphic bands on a total sample of 91 individuals from 32 localities in Europe and Asia. Moreover, each primer allowed for an exact diagnosis of each of the six Apodemus species and thus provides a simple and reliable tool for the hitherto problematic discrimination of species from the subgenus Sylvaemus. Genetic distances between species ranged from 0.079, among the closely related A. flavicollis and A. sylvaticus, to 0.203 between A. mystacinus and A. agrarius. A. flavicollis, A. uralensis and A. sylvaticus display a strong population substructure. The range of genetic distances among geographic samples within last two species reaches the values obtained for closely related species. ISSR markers proved to be a simple and reliable tool for species diagnosis, as well as for estimating genetic diversity below the species level and for closely related species, but they showed questionable reliability for larger genetic distances.     

Genetic variation and evolution in the genus Apodemus (Muridae: Rodentia)

Genetic variation was studied using protein electrophoresis of 28–38 gene loci in 1347 specimens of Apodemus agrar-ius, A. peninsulae, A. flavicollis, A. sylvaticus, A. alpicola, A. uralensis, A . cf. hyrcanicus, A. hermonensis, A. m. mystacinus and A. m. epimelas , representing 121 populations from Europe, the Middle East, and North Africa. Mean values of heterozygosity per locus for each species ranged from 0.02 to 0.04. Mean values of Nel's genetic distance ( D ) between the taxa ranged from 0.06 (between A. flavicollis and A. alpicola) to 1.34 (between A. uralensis and A. agrarius ). The highest values of D were found between A. agrarius and other Apodemus species (0.62-1.34). These values correspond to those generally observed between genera in small mammals. Our data show that A. agrarius and A. peninsulae are sister species, well-differentiated from other taxa. High genetic distance between A. m. mystacinus and A. m. epimelas leads us to consider them distinct species and sister taxa to other Western Palaearctic species of the subgenus Sylvaemus . The data also suggest a recent separation of members of the latter group from a common ancestor, and subsequent rapid radiation, making it difficult to infer phylogenetic relationships. Some taxonomic implications of the results are discussed further.© 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 75 , 395–419.     

Arboreal gaits in three sympatric rodents Apodemus agrarius,Apodemus flavicollis (Rodentia,Muridae) and Myodes glareolus (Rodentia,Cricetidae)

Vertical stratification of the arboreal habitat allows the coexistence of several species in a given area, because the complex arboreal strata can be used in different ways by arboreal and scansorial mammals. The present report experimentally investigated the gait metrics on different arboreal substrates, of three sympatric rodents living in a deciduous forest in Poznań, Poland. Arboreal locomotion was compared between the burrowing striped field mouse, Apodemus agrarius, the scansorial bank vole, Myodes glareolus, and the more arboreal yellow-necked mouse, Apodemus flavicollis. We filmed two wild-caught individuals from each species walking on four different substrate diameters (2 mm, 5 mm, 10 mm, 25 mm) and three different inclinations (45° descending, horizontal, 45° ascending) at 240 fps and collected a set of gait parameters from a total of 273 complete cycles. Our results did not demonstrate clear relationships between arboreal locomotion and the ecology of the three species. Only A. flavicollis exhibited locomotor features partly associated with arboreal competence, including lower velocity and diagonality on narrow substrates and asymmetrical gaits on wider ones. On the other hand, the two Apodemus species, despite their different ecologies, shared a few locomotor similarities, such as velocity regulation primarily by stride frequency, and similar effects of substrate size and inclination on diagonality, duty factor, and duty factor index indicating the possibility of a phylogenetic signal. Because the selected gait parameters provided limited insight into the ability of small mammals to move competently through an arboreal habitat, these findings indicate that the relationship between behaviour and ecology is complex.     

Molecular phylogeny of wood mice (Apodemus, Muridae) in East Asia

Sequences of the mitochondrial cytochrome b (1140 bp) and nuclear IRBP (1152 bp) genes were used to assess the evolutionary history of Apodemus , using the complete set of Asian species. Our results indicate that speciation in Asia involved three radiations, which supports an earlier study. The initial radiation yielded A. argenteus (Japanese endemic), A. gurkha (Nepalese endemic), and the ancestral lineage of the remaining Asian species. This lineage subsequently diverged into four groups: agrarius-chevrieri ( agrarius group), draco-latronum-semotus ( draco group), A. peninsulae , and A. speciosus (Japanese endemic). The final step consisted of divergence within two species groups as a consequence of the geography of the Yunnan-Guizhou plateau and Taiwan. The ecological ability of two Apodemus species to inhabit one locality via niche partitioning likely drove the second radiation and shaped the basic geographical pattern seen today: A. argenteus and A. speciosus in Japan, A. agrarius and A. peninsulae in northern China, and the A. agrarius and A. draco groups in southern China. The three radiations are estimated to have occurred 7.5, 6.6, and 1.8–0.8 Mya respectively, using the IRBP clock, based on rat–mouse divergence 12 Mya.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 469–481.     

Bibliography of the genus Apodemus (Rodentia, Muridae)

A complete bibliography of the genus Apodemus for the period 1970–1980 is presented, with a selection of the more important papers from earlier years, and a few references for 1981. A coded index to species and subjects is provided. All titles have been translated into English.     

Population structure and dynamics of sympatric Apodemus species (Rodentia: Muridae)

This paper describes the population biology of sympatric populations of Apodemus sylvaticus and A. flavicollis in western England. Annual changes in population siie closely resembled those seen in allopatric populations such that simultaneous peaks in number occurred only in autumn and early winter. Numbers of A. sylvaticus were low and stable in early summer increasing rapidly in late summer and autumn and remaining high throughout winter before a sharp decline in spring. Numbers of A. flavicollis increased after the start of reproduction and continued to rise throughout summer and autumn. The period of major decline of this species was during early winter. Densities of A. sylvaticus and A. flavicollis were comparable with those cited in allopatric studies. Spring populations of both species consisted of cohorts first appearing in the late summer and autumn of the previous year. Few individuals present in spring survived the summer. Reproduction of A. flavicollis commenced four to six weeks before that of A. sylvaticus . Seasonal variation in mean weight of adult males and females also indicates that those changes, due to the acquisition of secondary sex characteristics and reproductive maturity, occur earlier in A. flavicollis than in A. sylvaticus . Data presented contain no evidence of interspecific competition and it is concluded that interspecific differences, chiefly asynchrony in reproduction and annual population cycles, contribute to their stable coexistence.     

Epigenetic polymorphism in populations of Apodemus sylvaticus and Apodemus flavicollis (Rodentia, Muridae)

The main object of this study was to use discontinuous variation as a means of measuring divergence between populations of A. sylvaticus (L.) and A. flavicollis (Melchior) from Britain and Eurasia. Samples from 17 populations (671 skulls) were each classified for 20 discontinuous morphological variants. The investigation has shown that certain epigenetic variants have a characteristic incidence in populations of the two species and that the relative incidence of the variants could be used as an aid to specific separation. The calculation of degree of divergence between the populations has made it possible to suggest the order in which populations are related to each other.     

Sex chromosomes at early meiosis in three wood mice species of the genus Apodemus (Rodentia, Muridae)

The prophase of the first meiotic division was studied in field mice of the species Apodemus (Sylvaemus) flavicollis, A. (S.) ponticus, and A. (S.) uralensis by light and electron microscopy. The karyotypes of the species were described on the base of electron microscopy of synaptonemal complexes in spermatocytes I. The axial elements of the sex chromosomes at early-middle pachytene can synapse along the major portion of the Y axis; at late pachytene-early diplotene, the synapsis region shrinks; and at diakinesis-metaphase I, X and Y chromosomes associate tail-to-tail in all species studied. The behavior of sex chromosomes in the synapsis in the species studied was quite uniform. The results are discussed in the context of earlier data on the behavior of sex chromosomes in various rodent species in meiosis prophase I and their banding.     

The mitochondrial genome of Apodemus peninsulae (Rodentia, Muridae)

The complete mitochondrial (mt) genome of the Korean field mouse Apodemus peninsulae was sequenced and found to be 16,266 bp in length. The mt protein-coding genes of A. peninsulae had ATG, GTG, ATC, and ATA as initiation codons and TAA, TAG, TA, and T as termination codons. Two forms each of trnL and trnS and the three tRNA clusters, IQM, WANCY, and HSL were identified, as in the typical Rodentia mt genome. Among tRNAs, abnormal cloverleaf structure of trnS((AGY)) was identified in DHU arm. The l-strand replication origin has the potential to form a stable stem-loop structure and control region has several conserved sequence elements.     

The phylogeny of the Praomys complex (Rodentia: Muridae) and its phylogeographic implications

Among the African Murinae (Rodentia, Muridae), the Praomys complex, whose systematics has been studied by different approaches, has raised numerous taxonomic problems. Different taxa, namely Praomys, Mastomys , Myomys and Hylomyscus have been considered either as separate genera or subgenera of Praomys . In order to understand the relationships within the Praomys complex and to test the monophyly of the genus Praomys , a cladistic analysis was conducted, based on morpho-anatomical factors involving different species of Praomys , Mastomys , Myomys and Hylomyscus. The results indicate that the Praomys complex is monophyletic, as are the genera Hylomyscus , Mastomys and Myomys , whereas the genus Praomys appears paraphyletic. Indeed, a group of species including Praomys jacksoni was found to be more closely related to the genera Mastomys and Myomys than to a Praomys tullbergi -group. The biotopes and the distribution areas of the species were mapped on the phylogeny. It appears that the different clades each present a relative ecological cohesion and are arranged according to a gradient from closed to open habitats. From there, an evolutionary scenario is proposed for the emergence of the different clades and species of the genus Praomys sensu stricto.     

Impact of radio-collars on Yellow-necked Mice, Apodemus flavicollis (Mammalia, Rodentia)

The impact of wearing a radio-collar was investigated for 124 Yellow-necked Mice Apodemus flavicollis, in mixed deciduous woodlands. Collared mice neither lost more weight nor lost weight more frequently than control individuals. The cost of wearing a collar was not greater for small individuals than for larger ones. The impact of wearing a radio-collar was mostly due to the shape and size of the radio-collar and the external antenna rather than the package weight. This aspect was particularly acute in the presence of ticks. One solution to this problem may be to build and use radio-transmitter packages as compact (but nevertheless resistant to gnawing) and symmetrical as possible and to use collars impregnated with tick repellent.     

Microsatellites indicate a high frequency of multiple paternity in Apodemus (Rodentia)

Microsatellites were employed to estimate frequency of multiple paternity litters of two species of mice (genus Apodemus): striped field mouse (A. agrarius), and wood mouse (A. sylvaticus). Ten pregnant females of A. agrarius and six of A. sylvaticus were collected from natural populations in the northern Ukraine and analysed with 11 and nine microsatellite loci, respectively. Multiple paternity was indicated in eight of 10 litters in A. agrarius and in three of six litters in A. sylvaticus. Multiple paternity was documented at several loci (ranging from two to 10). In two cases (A. agrarius), three males were estimated to have fathered the litter.     

Chromosomal variability of the field mouse Apodemus agarius (Rodentia, Muridae)

Chromosome sets of 114 Apodemus agrarius mice from 29 localities in Moldova, Ukraine, Siberia, and Far East were studied by means of G-, C-, and NOR-banding. In all populations studied, the Y chromosome was shown to be a medium-size acrocentric chromosome consisting of heterochromatin. Chromosome polymorphism observed in populations from Primorskii krai concerned (1) the morphology of the first two autosome pairs (variants A/A, A/ST, and ST/ST), (2) the number of metacentric chromosomes (from 6 to 8), and (3) heterochromatin localization in the pericentromeric regions of two metacentric chromosome pairs. A karyotype with an additional heterochromatic microchromosome found in all the metaphases studied was described in one mouse from a locality of western Primorye that has not been studied previously. In the karyotype of 15 mice from four populations of Primorye, the pool of nucleolus organizer regions is distributed over three autosome pairs rather than over four, as is the case A. agrarius from Europe. Based on the analysis of literature sources and our own data, the problem of chromosome polymorphism in the field mouse is discussed.     

Discrimination of the sibling species Apodemus flavicollis and A. sylvaticus (Rodentia,Muridae)

Karyotyping and several molecular methods have allowed successful identification of two morphologically similar wide-ranging Western Palearctic species, the yellow-necked field mouse Apodemus flavicollis (Melchior, 1934) and the long-tailed wood mouse A. sylvaticus (Linnaeus, 1758), but reliable species diagnosis on the basis of morphometric characters is particularly problematic. Although they are easily morphologically distinguishable in Central and Northern Europe, this is not the case in southern parts of their distribution areas. Despite that, we have successfully discriminated A. flavicollis and A. sylvaticus from Serbia (Southern Europe) using geometric and traditional morphometric methods on a data set for ventral crania of specimens previously genotyped by the Inter Simple Sequence Repeat-PCR (ISSR-PCR). Discrimination power of applied approaches was more or less similar. The majority of our results were consistent with those obtained for specimens collected across the Czech Republic (Central Europe). Morphological differences observed herein, as well as those already reported between A. flavicollis and A. sylvaticus from the central and northern parts of their distribution areas, could be the outcome of their biology, i.e. ecological discrepancies, different assumed evolutionary scenarios considering biogeography, phylogeny, history and ontogeny.     

Cytogenetic differentiation of Apodemus peninsulae populations (Rodentia, Muridae) in eastern Siberia

Population and cytogenetics studies of geographical karyotype changes in East Asiatic mice in East Siberia showed that each population has unique cytogenetic structure, owing to different variants of B-chromosomes system, including 1 to 18 B-chromosomes differing in morphology and size. For each population, we observed the prevalence of mice with a definite number and morphology of B-chromosomes, which is probably the result of homeostatic processes in the population. Local changes were revealed, due to increase in the number of dot-like B-chromosomes in the direction from the West to the East.     

Molecular phylogeny evidence of altitudinal distribution and habitat adaptation in Korean Ephemera species (Ephemeroptera: Ephemeridae)

Molecular phylogeny of the four Korean Ephemera species, Ephemera orientalis, E. sachalinensis, E. strigata, and E. separigata, was inferred from 630 bp sequences of the partial mitochondrial cytochrome c oxidase I (COI) gene. Results indicated that mean intraspecific sequence divergences were 0.70%, whereas mean interspecific divergences were 15.75%, and 17 samples were distinguished to four species correctly by COI sequences. The results also demonstrated that four species of Korean Ephemera assembled a monophyletic group with high support in maximum parsimony and maximum likelihood analyses. This Ephemera group was divided into two major clades of E. orientalis–E. sachalinensis and E. strigata–E. separigata. Furthermore, we demonstrated that this phylogeny explained altitudinal and habitat adaptations of Korean Ephemera species. The E. orientalis–E. sachalinensis clade, a widespread and lowland‐adapted mayfly group, retained plesiomorphic traits such as paired stripes on abdominal segments and was regarded as plesiotypic in terms of habitat adaptation, compared to the geographically more limited and upland‐adapted E. strigata–E. separigata clade.     

Phylogenetic affinities and species limits within the genus Megadontomys (Rodentia: Cricetidae) based on mitochondrial sequence data

The evolutionary history of the genus Megadontomys, a group of mice allopatrically distributed along the cool‐humid forest in the highlands of México, is controversial. In this study, we examined phylogenetic relationships within the genus using sequences data from the complete mitochondrial cytochrome b gene. This information also allowed us to corroborate species limits, geographic boundaries of taxonomic entities and assess genetic variation within each taxon. The results of the phylogenetic analyses based on maximum likelihood, Bayesian inference and maximum parsimony were largely congruent in that M. nelsoni and M. thomasi were more closely related relative to M. cryophilus. These results are concordant with previous studies based on morphology and allozyme variation. However, testing of the alternative hypothesis of a closer evolutionary affinity between M. nelsoni–M. cryophilus did not produce a significantly less likely tree. The lack of unambiguous support towards one of these previously proposed contending hypotheses is congruent with the alternative scenario of an almost simultaneous diversification of the three species. Application of the phylogenetic species concept and the genetic species concept supports the recognition of three distinct taxonomic entities at the specific level. M. nelsoni inhabits the Sierra Madre Oriental (Hidalgo, Veracruz, and Puebla) including the Sierra Mazateca (Oaxaca); M. cryophilus is restricted to the Sierra de Juárez (Oaxaca); and M. thomasi occurs in portions of the Sierra Madre del Sur (Guerrero) and the Sierra Mixteca (Oaxaca). Our data show that M. thomasi is formed by two genetically distinct lineages that potentially may represent distinct Evolutionary Significant Units.     

Population structure of the large Japanese field mouse, Apodemus speciosus (Rodentia: Muridae), in suburban landscape, based on mitochondrial D-loop sequences

Genetic structure of the large Japanese field mouse populations in suburban landscape of West Tokyo, Japan was determined using mitochondrial DNA control region sequence. Samples were collected from six habitats linked by forests and green tract along the Tama River, and from two forests segregated by urban areas from those continuous habitats. Thirty-five haplotypes were detected in 221 animals. Four to eight haplotypes were found within each local population belonging to the continuous landscape. Some haplotypes were shared by two or three adjacent local populations. On the other hand, two isolated habitats were occupied by one or two indigenous haplotypes. Significant genetic differentiation between all pairs of local populations, except for one pair in the continuous habitats, was found by analysis of molecular variance (amova). The geographical distance between habitats did not explain the large variance of pairwise F(ST)-values among local populations. F(ST)-values between local populations segregated by urban areas were higher than those between local populations in the continuous habitat, regardless of geographical distance. The results of this study demonstrated quantitatively that urban areas inhibit the migration of Apodemus speciosus, whereas a linear green tract along a river functions as a corridor. Moreover, it preserves the metapopulation structure of A. speciosus as well as the corridors in suburban landscape.