Geographic pattern of cranial differentiation in the Asian Midday Jird Meriones meridianus (Rodentia: Muridae: Gerbillinae) and its taxonomic implications

The existence of cryptic species in the midday jird (Meriones meridianus) has been suggested in literature, although based on little empirical data to support this hypothesis. In this study, a two‐dimensional landmark‐based geometric morphometric approach was used to investigate whether patterns in intraspecific variation in skull shape and size exist, using 110 skull specimens from more than 20 different localities along the distribution range of M. meridianus. This is the first study of morphological differences on such a big sample size and geographical range, and it tries to find whether skull shape variation in this species is best described as being clinal or rather reflecting cryptic diversity. The latter seems to be the case, as a dimorphic skull phenotype was found, reflecting a geographic disparity between the Middle East and the Far East specimens both in shape and in size. Distinct cranial differences were found in the overall cranial size and, also at the level of the inflation of the bulla, the elongation of the nasal, the length of the teeth row and the incisive foramen, as well as the distance in between the latter two. It thus seems that M. meridianus from Middle East is morphologically distinct from that of the Far East. Furthermore, our results also demonstrate that clinal variation could explain variation within Middle East populations, whereas a more heterogenous pattern is found for those of the Far East. The hypothesis that the observed phenotypic variation may reflect cryptic species is discussed, with the recommendation for a thorough taxonomical revision of the genus in the region.     

The <Emphasis Type="Italic">Eumyarion</Emphasis> (Mammalia,Rodentia, Muridae) assemblage from Sandelzhausen (Miocene,Southern Germany): a test on homogeneity

A large sample of Eumyarion cheek teeth (N = 569) from the Early/Middle Miocene boundary locality Sandelzhausen (MN5, Southern Germany), type locality E. bifidus Fahlbusch, 1964, is studied. It is concluded that this collection contains two species: E. bifidus and E. weinfurteri Schaub and Zapfe, 1953. The similarity in size and morphology of the cheek teeth of these two species is so great that only the M1 and M2 can be recognised with certainty. Eumyarion bifidus seems to be a descendant of E. orhani de Bruijn et al., 2006 from the Early Miocene (MN3) of Southwestern Anatolia and is therefore considered to be an immigrant into Central Europe.      

Allelic diversity at the Mhc-DQA locus in cotton rats (Sigmodon hispidus) and a comparison of DQA sequences within the family Muridae (Mammalia: Rodentia)

 The cotton rat (Sigmodon hispidus) is a common murid rodent of the southern United States, Mexico, and Central America. Using single-stranded conformation polymorphism analysis and DNA sequencing techniques, 11 DQA exon 2 alleles were detected among 180 S. hispidus from Caddo County, Oklahoma, USA. The alleles represent a single locus exhibiting a high level of polymorphism. Nucleotide and amino acid distance values among DQA alleles of S. hispidus were higher than those within Mus musculus and species of Rattus. Although the distribution of polymorphic amino acid residues among alleles of S. hispidus was similiar to that of Mus and Rattus, some residues of the α-helix region were more variable in S. hispidus. Comparisons of nonsynonymous and synonymous substitutions indicated a trend toward higher numbers of nonsynonymous substitutions; however, this difference was not significant statistically among S. hispidus alleles. To examine evolution of DQA alleleswithin Muridae, we performed a phylogenetic analysis that included DQA alleles from S. hispidus, Peromyscus leucopus, M. musculus, R. norvegicus, and six Australian species of Rattus. Results depicted monophyly for each genus, and this concordance between species and gene trees represents a lack of evidence for trans-species persistence of alleles among these genera. Received: 22 October 1998 / Revised: 17 March 1999     

Chromosomal evolution in the African Arvicanthine rats (Murinae, Rodentia): comparative cytogenetics of Lemniscomys (L. zebra, L. rosalia, L. striatus) and Arvicanthis dembeensis

Chromosomenevolution bei den afrikanischen arvicanthinen Ratten (Murinae, Rodentia): Vergleichende Cytogenetik von Lemniscomys (L. zebra, L. rosalia, L. striatus) und Arvicanthus dembeensis
Chromosomenbänderungsanalysen (G- und C- Bänderung) wurden an zwei Arten von Lemniscomys ( L. zebra, L. rosalia ) und einer dritten Art aus Benin ( L. striatus ) durchgeführt, um den Verlauf der Chromosomenevolution in dieser Gattung zu verfolgen. Ein Mustervergleich mit der G-Bänderung macht es möglich, die Umstrukturierung der Chromosomen der Karyotypen zu erkennen. Die beiden Arten aus Tansania ( L. rosalia 2n=54, FNa=62; und L. zebra 2n=54, FNa=58) sind durch zwei perizentrische Inversionen voneinander unterschieden. Ein Polymorphismus für das X-Chromosom findet sich in beiden Arten. L. rosalia zeigt außerdem zwei verschiedene Formen des Y-Chromosoms. Das Bänderungsmuster von L. striatus (2n=44, FNa=68) aus Benin läßt im Vergleich zu L. zebra fünf Robertsonsche Fusionen erkennen. Außerdem ist der Karyotyp von L. rosalia aus Tansania verschieden von dem der für Südafrika (2n=48, FNa=62) beschrieben wurde und sollte daher als eine eigene Art betrachtet werden. Eine vergleichende Bänderungsanalyse mit einer anderen arvicanthinen Art ( Arvicanthus dembeensis, 2n=62, FNa=62) zeigt zwischen den beiden Gattungen eine vollständige Homologie in den G-Banden und die Unterschiede beruhen ausschließlich auf Tandem-Fusionen, perizentrischen Inversionen und Robertsonschen Fusionen.     

Chromosomal evidence for a polytypic structure of Arvicanthis niloticus (Rodentia, Muridae)

The analysis of R- and C-banding patterns of chromosomes of Arvicanthis niloticus from five localities of Africa (Egypt, Senegal, Burkina-Faso, Mali and Central African Republic) revealed the existence of three karyotypic forms labelled in the study as ANI-1, ANI-2 and ANI-3. These forms differ from each other by 6 to 8 chromosomal rearrangements such as reciprocal and Robertsonian translocations and pericentric inversions. Moreover, they possess different quantities of C-heterochromatin. The data indicate that these three forms are distinct species, cytogenetically isolated, and that a further taxonomic analysis of the genus Arvicanthis is needed.     

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.     

Molecular Relationships Among Octodontidae (Mammalia: Rodentia: Caviomorpha)

We examined ten species of octodontid rodents and representatives of three outgroups in a complete 13 × 13 DNA-hybridization matrix. The results were indexed as differences in median melting-point depressions (T,_ms), symmetrized, subjected to phylogenetic analysis using FITCH, bootstrapped, and exhaustively taxon-jackknifed. Within Octodontidae, four clades were recovered with 100% bootstrap and complete jackknife support: Tympanoctomys barrerae with Octomys mimax, Octodontomys gliroides alone, Octodon spp., and Aconaemys spp. with Spalacopus cyanus; the latter two clades were closer to each other than either was to Octodontomys or Octomys-Tympanoctomys, but were slightly nearer to and united with Octodontomys, with 89% bootstrap support. However, relationships among the three Aconaemys species and Spalacopus were not completely resolved by our experiments. Ctenomys coyhaiquensis, Abrocoma bennetti, and Lagostomus maximus represented successive outgroups to Octodontidae, while one-way comparisons with Cavia porcellus and Microcavia australis suggested that these caviids are almost as distant from octodontoids as is the chinchilloid Lagostomus. When the data were suitably corrected for percentage hybridization and saturation, division of the distances by the rate previously determined for most amniotes (0.48%/myr) suggested that the basal divergence among the caviomorph rodents examined occurred about 59 myrbp, and that Octodontidae originated 40 myrbp and diversified into extant lineages beginning 14 myr ago. Calibration against the date of the earliest known caviomorph (late Eocene or about 37.5 myrbp) gave a rate of 0.75%/myr, which would suggest later dates for subsequent caviomorph cladogenesis. It is notable that, based respectively on the slower or faster rates, the tetraploid Tympanoctomys barrerae must have diverged from its sister-taxon Octomys mimax 10 or 6.5 myr ago.     

Skull evolution in the rhinocerotidae (Mammalia,Perissodactyla): Cartesian transformations and functional interpretations

Cartesian transformation, applied as a landmark morphometric method, is used to investigate some of the evolutionary shape changes leading to the skulls of the modern rhinoceroses. The early Oligocene genusSubhyracodon serves as the primitive shape from which the extant genera (Dicerorhinus, Rhinocerso, Diceros, andCeratotherium) have been transformed. Coordinate data for 21 landmarks, defined in lateral view, are analyzed by the computer program Thinplate Splines. Each of the four transformations are interpreted separately as shape change fromSubhyracodon. Computed results forRhinoceros are also compared with previous results obtained by visual interpretation (the classic method). Among the extant genera,Ceratotherium andRhinoceros have the most derived shapes and are opposites with respect to orientation of the occiput and relative size of the mandible angle. The significance of these foci of change is discussed in terms of the functions of the masseter and posterior temporalis muscles. In head positions associated with feeding on short vs. tall grasses, the two skull shapes are consistent with a role for these muscles in support of a large mandible against gravity. This common factor may help to explain both shapes.     

Chromosomal diversity in the genus Arvicanthis (Rodentia, Muridae) from East Africa: a taxonomic and phylogenetic evaluation

In this paper we discuss the contribution of cytogenetics to the systematics of Arvicanthis in East Africa, by reviewing all the known chromosomal cytotypes of the genus in the area. We also provide G‐ and C‐banding comparisons for two recently described karyotypes, provisionally named ANI‐5 (2n = 56, NFa = 62) and ANI‐6 (2n = 60, NFa = 72). This, therefore, brings the total number of known cytotypes in this area to 10. Five of these correspond to the species recognized by the latest rodent checklist, i.e. A. nairobae (2n = 62, NFa = 78), A. neumanni (2n = 52–53, NFa = 62), A. blicki (2n = 48, NFa = 62), A. abyssinicus (2n = 62, NFa = 64) and A. niloticus (2n = 62, NFa = 60–62). The taxonomic status of the remaining five cytotypes (A. cf. somalicus, 2n = 62 NFa = 62–63; ANI‐5, 2n = 56, NFa = 62; ANI‐6/6a 2n = 60, NFa = 72/76; ANI‐7, 2n = 56, NFa = 78; and ANI‐8, 2n = 44, NF = 72) is discussed. Finally, we reconstruct the phylogenetic relationships among all the known karyotypes on the basis of banding data available for the genus in Africa and show the occurrence of two main clades, each characterized by different types of chromosomal rearrangements. The times of the cladogenetic events, inferred by a molecular clock, indicate that karyotype evolution has accomplished almost all the dichotomic events from the end of the Miocene to the present day. The discovery of a large chromosomal differentiation between populations showing low genetic distances and intrapopulation chromosomal polymorphism suggests that the process of chromosomal differentiation in Arvicanthis is still ongoing and may possibly be responsible for speciation.     

Albumin preservation in fossil bones and systematics of Malpaisomys insularis (Muridae,Rodentia), an extinct rodent of the Canary Islands

Collagen and albumin were extracted from subfossil bones of the extinct rodent Malpaisomys insularis; the radiocarbon age of the sample was at least 1.73 Ka. Electrophoretic and immunological techniques indicate the preservation of albumin while collagen is revealed as degraded fragments. Immunological comparisons between Malpaisomys albumin and albumin antisera of four extant rodents indicate that Malpaisomys is more closely related to Mus than to Acomys and Uranomys and that Cavia is the most divergent taxon studied. The branching order of the fossil and the extant rodents is discussed with respect to the relationships derived from morphological criteria.     

Phylogeographic study of Apodemus ilex (Rodentia: Muridae) in Southwest China

Background

The Mountains of southwest China have complex river systems and a profoundly complex topography and are among the most important biodiversity hotspots in the world. However, only a few studies have shed light on how the mountains and river valleys promote genetic diversity. Apodemus ilex is a fine model for investigating this subject.

Methodology/Principal Findings

To assess the genetic diversity and biogeographic patterns of Apodemus ilex, the complete cytochrome b gene sequences (1,140 bp) were determined from 203 samples of A. draco/ilex that were collected from southwest China. The results obtained suggested that A. ilex and A. draco are sistergroups and diverged from each other approximately 2.25 million years ago. A. ilex could be divided into Eastern and Western phylogroups, each containing two sub-groups and being widespread in different geographical regions of the southern Hengduan Mountains and the western Yunnan - Guizhou Plateau. The population expansions of A. ilex were roughly from 0.089 Mya to 0.023 Mya.

Conclusions

Our result suggested that A. ilex is a valid species rather than synonym of A. draco. As a middle-high elevation inhabitant, the phylogenetic pattern of A. ilex was strongly related to the complex geographical structures in southwest China, particularly the existence of deep river valley systems, such as the Mekong and Salween rivers. Also, it appears that the evolutionary history of A. ilex, such as lineage divergences and population expansions were strongly affected by climate fluctuation in the Late Pleistocene.     

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.     

Taxonomic hypotheses regarding the genus Gerbillus (Rodentia,Muridae, Gerbillinae) based on molecular analyses of museum specimens

Methodological improvements now allow routine analyses of highly degraded DNA samples as found in museum specimens. Using these methods could be useful in studying such groups as rodents of the genus Gerbillus for which i) the taxonomy is still highly debated, ii) collection of fresh specimens may prove difficult. Here we address precise taxonomic questions using a small portion of the cytochrome b gene obtained from 45 dry skin/skull museum samples (from 1913 to 1974) originating from two African and three Asian countries. The specimens were labelled Gerbillus gerbillus, Gerbillus andersoni, Gerbillus nanus, Gerbillus amoenus, Gerbillus perpallidus and Gerbillus pyramidum, and molecular results mostly confirmed these assignations. The close relationship between Gerbillus nanus (Asian origin) and Gerbillus amoenus (African origin) confirmed that they represent vicariant sibling species which differentiated in allopatry on either side of the Red Sea. In the closely related Gerbillus perpallidus and Gerbillus pyramidum, specimens considered as belonging to one Gerbillus pyramidum subspecies (Gerbillus pyramidum floweri) appeared closer to Gerbillus perpallidus suggesting that they (Gerbillus pyramidum floweri and Gerbillus perpallidus) may represent a unique species, distributed on both sides of the Nile River, for which the correct name should be Gerbillus floweri. Furthermore, the three other Gerbillus pyramidum subspecies grouped together with no apparent genetic structure suggesting that they may not yet represent genetically differentiated lineages. This study confirms the importance of using these methods on museum samples, which can open new perspectives in this particular group as well as in other groups of interest.     

Taxonomic identity of Chionomys nivalis spitzenbergerae (Mammalia: Rodentia)

The most genetically divergent populations of the European snow vole Chionomys nivalis occupy the extreme eastern range of the Black Sea and the Caspian Sea. It was recently suggested that subspecies C. n. spitzenbergerae from the Central Taurus Mountains (Turkey) represents a highly divergent lineage of C. nivalis from the Alada? Range which induced us to address its status by examining topotype specimens. Two females karyotyped displayed 54 mitotic chromosomes of 25 acrocentric autosomal pairs. One of the smallest autosomal pairs was heteromorphic in both specimens, consisting of a submetacentric and an acrocentric chromosomes (NFa = 53). Cytochrome b sequences however unambiguously clustered both individuals with reference sequences from C. nivalis from Turkey. We conclude that a deviant haplotype from the Alada? Range does not represent C. n. spitzenbergerae. Its taxonomic identity was not resolved in our study. Further attention should be devoted to snow voles from Central Anatolia and Western Iran, which are characterized by cranial peculiarities.     

Evolution of MHC-<Emphasis Type="Italic">DRB</Emphasis> class II polymorphism in the genus <Emphasis Type="Italic">Apodemus</Emphasis> and a comparison of <Emphasis Type="Italic">DRB</Emphasis> sequences within the family Muridae (Mammalia: Rodentia)

Allelic diversity at major histocompatibility complex (MHC) genes is thought to be maintained by balancing selection over long periods of time, even across multiple speciation events. Trans-species sharing of MHC alleles among genera has been supported by many studies on mammals and fish, but in rodents, the results are ambiguous. We investigated natural levels of MHC-DRB variability and evolutionary processes in the wood mouse (Apodemus sylvaticus) and the yellow-necked mouse (Apodemus flavicollis), which are common, sympatric murid rodents in European forests. Using single-strand conformation polymorphism analysis and DNA sequencing, 38 DRB exon 2 alleles were detected among 162 A. sylvaticus from nine different locations in Germany and Switzerland, and 15 DRB exon 2 alleles were detected among 60 A. flavicollis from three different locations in northern Germany. There was evidence for balancing selection in both species. Phylogenetic analysis, including additional murid taxa, showed that the DRB exon 2 sequences did not separate according to species, consistent with trans-species evolution of the MHC in these taxa.