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Rhabdomys is a genus that occupies a variety of habitats, including forest margins. Among the Rhabdomys taxa, Rhabdomys dilectus chakae has a distribution that covers the eastern seaboard of South Africa, with a poorly defined divergence date from its sister taxon Rhabdomys dilectus dilectus. Here, we study three mitochondrial markers (cytochrome b, cytochrome c oxidase I and partial control region) of R. d. chakae across the Eastern Cape and KwaZulu-Natal forests of South Africa, aiming to determine the cladogenesis (molecular dating) and effective population size of this subspecies through time, in addition to its cladogenesis in relationship to other species within the genus. A phylogenetic analysis revealed six clades within Rhabdomys, confirming that our study area is occupied solely by R. d. chakae, to the exclusion of other recognized sibling species. A fossil-calibrated Bayesian relaxed molecular clock estimated a recent split between R. d. chakae and R. d. dilectus ~1.4 ± 0.35 Mya and between two Rhabdomys pumilio groups, coastal A and B, at 1.16 ± 0.44 Mya. Coalescent Bayesian skyline plots revealed a stable population of R. d. chakae in the study area that was in slow decline until 2500 years ago, when there was an expansion in the late Holocene. Radiation within Rhabdomys dates as far back as 4.27 Mya, and subsequent demographic fluctuations primarily reflect palaeoclimatic changes.

Abstract South African small mammals are under-represented in DNA barcoding efforts, particularly from the eastern forested regions of the country. This study reports DNA barcoding of Rhabdomys taxa from previously unsampled parts of the Eastern Cape and KwaZulu-Natal provinces of South Africa. The complete mitochondrial DNA cytochrome oxidase I (COI) gene was sequenced for 101 Rhabdomys sp. individuals from 16 localities from all three main forest groups (coastal, mistbelt, and scarp forests). Molecular data were supplemented with external morphological measurements, including those deemed potential taxonomically diagnostic characters. Findings indicate the area to be inhabited solely by Rhabdomys dilectus chakae. Haplotypes distributed across the three forest groups were separated by shallow sequence divergences ranging from 0.001–0.015 (Kimura 2-parameter model) and displayed very little population genetic structure (FST= 0.071787). Morphological data revealed some regional metric differences in external morphology, but all the head-and-body to tail (HB: tail) ratios match that of R. d. chakae, and consequently, molecular and morphological data are congruent. These data confirm a range extension of R. d. chakae, supporting the utility of COI barcodes in the identification of small mammalian species.

The Eastern Cape Province harbors almost half of the indigenous forest in South Africa, but these forests are threatened by large-scale agricultural and urban development planned for the coming decade. Additional anthropogenic development is likely to cause further fragmentation and degradation of forests inhabited by the dusky pipistrelle bat (Pipistrellus hesperidus). We used eight microsatellite markers to study the genetic diversity, population structure, and migration, of P. hesperidus (n = 120) across 14 sites in the Eastern Cape and KwaZulu-Natal provinces of South Africa. We examined the effect of contemporary land cover types on genetic differentiation to assess whether current levels of urbanization and agricultural development affect gene flow. High gene flow and low population structure were evident across sampled sites, apart from genetic discontinuities at the northern (Oribi Gorge Nature Reserve) and southern (Alexandria Forest) ends of the seaboard. Genetic discontinuity at Oribi Gorge may relate to anthropogenic modification of two rivers surrounding the forest, while the Alexandria-linked barrier is a climatic break known as the Bedford gap. Migration rates generally were low between sites except for one Scarp forest, Manubi State Forest, from which individuals dispersed to other sites. The Amatole Mistbelt forests supported high genetic diversity, and likely served as a refugium for P. hesperidus during the Last Glacial Maximum. The composition of land cover classes between sites was a poor predictor of genetic differentiation, although it seems likely that P. hesperidus uses riparian habitats and wetlands for dispersal. Lastly, urban and agricultural development did not have a significant effect on genetic differentiation, which may reflect the wide niche breadth and intermediate distribution range of the species. This study provides insights into genetic diversity and gene flow of P. hesperidus across the study region prior to agricultural intensification and large-scale urbanization.

Historical forest contractions may have restricted the distributions of forest-utilizing fauna while providing opportunities for range expansions for open-habitat species. We aimed to test if habitat associations have played an important role in determining population genetic structure and demographic responses of six bats to oscillations in forest extent since the Last Glacial Maximum (LGM). We hypothesized that forest-associated species would display high levels of population structure and past population contractions as their distribution is dependent on fragmented forests. By contrast, habitat generalists would demonstrate low geographical structuring and historical population stability as suitable habitats are widely available. We used mitochondrial DNA to generate genetic diversity and population structure metrics of three forest-associated species and three habitat generalists in South Africa. Neutrality tests and Bayesian skyline plots were used to investigate demographic histories. A forest habitat association did not inform the population genetics of the study species. Rather, species-specific traits of roosting requirements, philopatry to the natal range and dispersal ability informed the observed structure. All species demonstrated population expansions during the Pleistocene, with no apparent decline during the LGM. It appears that the lower climate change footprint and refuge-status of eastern South Africa prevented population declines of insectivorous bats during the LGM.

Bats are a highly diverse order with substantial economic and ecological value. Similarly, forests in South Africa form a valuable biome supporting unique biotic diversity, yet forest bat communities have not previously been surveyed. We sampled 17 forests, of seven forest types and three forest groups, in the Eastern Cape and southern KwaZulu-Natal provinces, located within the Maputaland-Pondoland-Albany global biodiversity hotspot. We utilized capture and acoustic survey methods to compile the first bat species inventory for these forests. Species identification was performed with a combined approach of morphology, echolocation, and DNA barcodes. With this we contributed novel DNA barcodes to the Barcode of Life Data System. A total of 25 species was recorded, with range extensions southward into the Eastern Cape for six species, indicating the region to be more diverse than previously thought. Updated modelled distribution maps for these species are presented. We compiled the first reference call library of hand released bats for South African forest habitats, and the southeast region of the country. We compared species richness, diversity, and dissimilarity of the forest types and found that Scarp forests host the highest species richness and diversity. Patterns of species assemblage similarities between forest groups are attributed to forest biogeographical history and historical dispersal routes of forest fauna. A comprehensive survey, such as this, may assist in the compilation and implementation of forest conservation management plans and future monitoring programs.

Pteropodidae and Hipposideridae are 2 of the 9 chiropteran families that occur on Madagascar. Despite major advancements in the systematic study of the island's bat fauna, few karyotypic data exist for endemic species. We utilized G- and C-banding in combination with chromosome painting with Myotismyotis probes to establish a genome-wide homology among Malagasy species belonging to the families Pteropodidae (Pteropus rufus 2n = 38; Rousettus madagascariensis, 2n = 36), Hipposideridae (Hipposideros commersoni s.s., 2n = 52), and a single South African representative of the Rhinolophidae (Rhinolophus clivosus, 2n = 58). Painting probes of M. myotis detected 26, 28, 28, and 29 regions of homology in R. madagascariensis, P. rufus, H. commersoni s.s, and R. clivosus, respectively. Translocations, pericentric inversions, and heterochromatin additions were responsible for karyotypic differences amongst the Malagasy pteropodids. Comparative chromosome painting revealed a novel pericentric inversion on P. rufus chromosome 4. Chromosomal characters suggest a close evolutionary relationship between Rousettus and Pteropus. H. commersoni s.s. shared several chromosomal characters with extralimital congeners but did not exhibit 2 chromosomal synapomorphies proposed for Hipposideridae. This study provides further insight into the ancestral karyotypes of pteropodid and hipposiderid bats and corroborates certain molecular phylogenetic hypotheses.

The vlei rat Otomys irroratus has a wide distribution in southern Africa with several datasets indicating the presence of two putative species (O. irroratus and O. auratus). In the present study we use mitochrondrial cyt b data (∼950 bp) from 98 specimens (including museum material) collected throughout the range of the species to determine the geographical limits of the two recognized species, and we link this to niche modelling to validate these species. Phylogenetic analysis of the DNA sequence data, using maximum parsimony, neighbour joining and Bayesian inference, retrieved two divergent statistically well-supported clades. Clade A occurs in the Western and Eastern Cape while Clade B occurs in the Free State, KwaZulu-Natal, Northern Cape and Mpumalanga provinces of South Africa and Zimbabwe. Mean sequence divergence between the two clades (A and B) was 7.0% and between sub-clades comprising clade B it was 4.8%; the two clades diverged during the Pleistocene. Within Clade A the mean sequence divergence among specimens was 1.91%. Niche modelling revealed that the incipient species occupy distinct bioclimatic niches associated with seasonality of precipitation. Our data allow insightful analysis into the factors that could have led to cladogenesis within this rodent. More significantly, the new data enable us to pinpoint the Eastern Cape province as a contact zone for the divergent species. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104, 192–206.

There is great uncertainty about how Ireland attained its current fauna and flora. Long-distance human-mediated colonization from southwestern Europe has been seen as a possible way that Ireland obtained many of its species; however, Britain has (surprisingly) been neglected as a source area for Ireland. The pygmy shrew has long been considered an illustrative model species, such that the uncertainty of the Irish colonization process has been dubbed ‘the pygmy shrew syndrome'. Here, we used new genetic data consisting of 218 cytochrome (cyt) b sequences, 153 control region sequences, 17 Y-intron sequences and 335 microsatellite multilocus genotypes to distinguish between four possible hypotheses for the colonization of the British Isles, formulated in the context of previously published data. Cyt b sequences from western Europe were basal to those found in Ireland, but also to those found in the periphery of Britain and several offshore islands. Although the central cyt b haplotype in Ireland was found in northern Spain, we argue that it most likely occurred in Britain also, from where the pygmy shrew colonized Ireland as a human introduction during the Holocene. Y-intron and microsatellite data are consistent with this hypothesis, and the biological traits and distributional data of pygmy shrews argue against long-distance colonization from Spain. The compact starburst of the Irish cyt b expansion and the low genetic diversity across all markers strongly suggests a recent colonization. This detailed molecular study of the pygmy shrew provides a new perspective on an old colonization question.

The chiropteran fauna of Madagascar comprises eight of the 19 recognized families of bats, including the endemic Myzopodidae. While recent systematic studies of Malagasy bats have contributed to our understanding of the morphological and genetic diversity of the island's fauna, little is known about their cytosystematics. Here we investigate karyotypic relationships among four species, representing four families of Chiroptera endemic to the Malagasy region using cross-species chromosome painting with painting probes of Myotis myotis: Myzopodidae (Myzopoda aurita, 2n = 26), Molossidae (Mormopterus jugularis, 2n = 48), Miniopteridae (Miniopterus griveaudi, 2n = 46), and Vespertilionidae (Myotis goudoti, 2n = 44). This study represents the first time a member of the family Myzopodidae has been investigated using chromosome painting. Painting probes of M. myotis were used to delimit 29, 24, 23, and 22 homologous chromosomal segments in the genomes of M. aurita, M. jugularis, M. griveaudi, and M. goudoti, respectively. Comparison of GTG-banded homologous chromosomes/chromosomal segments among the four species revealed the genome of M. aurita has been structured through 14 fusions of chromosomes and chromosomal segments of M. myotis chromosomes leading to a karyotype consisting solely of bi-armed chromosomes. In addition, chromosome painting revealed a novel X-autosome translocation in M. aurita. Comparison of our results with published chromosome maps provided further evidence for karyotypic conservatism within the genera Mormopterus, Miniopterus, and Myotis. Mapping of chromosomal rearrangements onto a molecular consensus phylogeny revealed ancestral syntenies shared between Myzopoda and other bat species of the infraorders Pteropodiformes and Vespertilioniformes. Our study provides further evidence for the involvement of Robertsonian (Rb) translocations and fusions/fissions in chromosomal evolution within Chiroptera.

Recent genetic studies have challenged the traditional view that the ancestors of British Celtic people spread from central Europe during the Iron Age and have suggested a much earlier origin for them as part of the human recolonization of Britain at the end of the last glaciation. Here we propose that small mammals provide an analogue to help resolve this controversy. Previous studies have shown that common shrews ( Sorex araneus ) with particular chromosomal characteristics and water voles ( Arvicola terrestris ) of a specific mitochondrial (mt) DNA lineage have peripheral western/northern distributions with striking similarities to that of Celtic people. We show that mtDNA lineages of three other small mammal species (bank vole Myodes glareolus , field vole Microtus agrestis and pygmy shrew Sorex minutus ) also form a ‘Celtic fringe’. We argue that these small mammals most reasonably colonized Britain in a two-phase process following the last glacial maximum (LGM), with climatically driven partial replacement of the first colonists by the second colonists, leaving a peripheral geographical distribution for the first colonists. We suggest that these natural Celtic fringes provide insight into the same phenomenon in humans and support its origin in processes following the end of the LGM.

The status of the pygmy shrew (Sorex minutus L.) as a native or an introduced species in Ireland has been subject to much debate. To examine this and other aspects of the colonization history of the Irish pygmy shrew, genetic variation was determined in 247 pygmy shrews collected throughout Ireland, using mitochondrial control region sequences and five polymorphic microsatellite loci. Genetic diversity was low for both types of marker. The median-joining network for control region sequences was star-like, suggesting that the colonization of Ireland involved a small number of founders and rapid population expansion thereafter; this was supported by other statistics. Molecular dating with both mitochondrial DNA and microsatellite data is consistent with a human introduction. This would have been several thousand years ago; a recent colonization within historical times can be ruled out. This is the first detailed population genetic study of the pygmy shrew anywhere in its range. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 97, 918–927.

Chromosomal polytypy, morphological conservatism and absence of data have frustrated the taxonomic revision of two species of southern African-endemic laminate-toothed rats (Otomys irroratus and Otomys saundersiae s.l.). New cytogenetic (G-banding and fluorescence in situ hybridization), DNA sequence [cytochrome b (cyt b) gene] and geometric morphometric data demonstrate the synonymy of O. saundersiae from Grahamstown (Eastern Cape, South Africa) under O. irroratus, and the validity of Otomys karoensis from the Fynbos Biome of the Western Cape. Phenotypic dimorphism in pelage colour and cranial morphology in O. irroratus from the climatically unpredictable Albany Thicket (=Savanna) Biome of the Eastern Cape results from the retention of allometric paedomorphic traits in some adults (saundersiae morph) but not others. The same paedomorphic traits are associated with speciation and karyotypic and genetic differentiation in O. karoensis. Within O. irroratus, two phenotypically and genotypically (cyt b divergence=6.4%) divergent lineages correspond with the Fynbos/Albany Thicket and Grassland biomes. Incipient speciation in O. irroratus seems to be associated with ecology rather than karyotype.

Aspects of the anatomy and histology of the alimentary canal of Thryonomys swinderianus were studied to gain a better understanding of this animal’s ability to digest large quantities of fibre. Morphometric measurements of the gut regions were taken with the aid of light, transmission and scanning electron microscopy. The stomach is fully glandular and unilocular with a low pH and is possibly the initial site of protein digestion. The duodenum, with its closely packed columnar cells with many mitochondria, may be the main site of enzymatic digestion. Owing to coprophagy, food and fermentation products are recycled for further digestion in the small intestine. The caecum is by far the largest region in mass and the apex ceci may be the main site of fermentation. A ridge-lined furrow is present inside the proximal colon. Thick muscle layers are present in the vicinity of the ridge-lined furrow. It is suggested that the furrow is part of the colonic separation mechanism that transports bacteria back to the caecum. The longitudinal folds and long villi in the proximal colon enlarge the surface area, where the columnar cells of the villi have many mitochondria, probably to absorb fermentation products and to transport water and electrolytes across the mucosa. In the distal colon many mucous cells open in tubular glands of Lieberkühn. Reabsorption of water probably occurs through the dense microvillous border.Keywords: Thryonomys, alimentary canal, anatomy, histology, cane rat

In contrast to a substantial inventory of mitochondrial D-loop sequences from the West European part of the range of the western house mouse Mus musculus domesticus, there are few such data from close to the Middle Eastern origin of the subspecies. This paper attempts to rectify this situation, with details of D-loop sequences of 92 mice from 45 localities in Turkey, adding to the six mice from three localities previously studied from that country. The molecular data were found to be consistent with Turkey being close to the site of origin of M. m. domesticus. The 54 Turkish haplotypes were distributed throughout a phylogenetic tree incorporating representative sequences described previously, and there was high nucleotide diversity among the Turkish mice. There was weak genetic structure in Turkey, although haplotypes of the 'Main Turkish Clade', while common over much of Turkey, were absent from Southern Anatolia. Haplotypes of the Main Turkish Clade were also found in western Europe and it is believed that they derived from Turkey, which thus represents one of the source areas for the westwards expansion of M. m. domesticus.

The phylogeography of the African four-striped mouse, Rhabdomys pumilio, was investigated using complete sequences of the mtDNA cytochrome b gene (1140 bp) and a combination of fluorescence in situ hybridization (FISH) and conventional cytogenetic banding techniques (G- and C-banding). Two cytotypes (2n=46 and 2n=48) were identified by cytogenetic analysis. There is no evidence of diploid number variation within populations, difference in gross chromosome morphology or of subtle interchromosomal rearrangements at levels detected by ZOO-FISH. Analysis of the mtDNA cytochrome b resulted in two major lineages that correspond roughly to the xeric and mesic biotic zones of southern Africa. One mtDNA clade comprises specimens with 2n=48 and the other representatives of two cytotypes (2n=48 and 2n=46). The mean sequence divergence (12%, range 8.3-15.6%) separating the two mtDNA clades is comparable to among-species variation within murid genera suggesting their recognition as distinct species, the prior names for which would be R. dilectus and R. pumilio. Low sequence divergences and the diploid number dichotomy within the mesic lineage support the recognition of two subspecies corresponding to R. d. dilectus (2n=46) and R. d. chakae (2n=48). Our data do not support subspecific delimitation within the nominate, R. pumilio. Molecular dating places cladogenesis of the two putative species at less than five million years, a period characterised by extensive climatic oscillations which are thought to have resulted in habitat fragmentation throughout much of the species range.

Fluorescence in-situ hybridization was used to construct a comparative chromosome map between the laboratory mouse, Mus musculus and the African four-striped mouse, Rhabdomys pumilio. A high degree of homology between the species was detected using both FISH and G-banding. Ten mouse chromosomes (2–4, 7, 14–16, 18, 19 and the X) were retained as chromosomal arms or intact chromosome blocks. Six mouse chromosome painting probes that correspond to mouse autosomes 5, 6, 8, 11, 12 and 13, produced double signals; the remaining four painting probes (1, 9, 10 and 17) hybridized to three or more R. pumilio chromosomes respectively. In total, the 20 mouse chromosome paints revealed 40 segments of conserved synteny in the R. pumilio genome. Most of the mouse chromosomes that produced single signals in R. pumilio have previously been shown to be conserved in the Black and Norwegian rats and the Chinese hamster. Eight contiguous segment associations appear to be R. pumilio specific, two were shared by R. pumilio and the Black and Norwegian rats, but to the exclusion of the Chinese hamster. Our data suggest that mouse chromosomes 1, 10, and 17 have undergone extensive rearrangements during genome evolution in the murids and may be useful markers for enhancing our understanding of the mode and tempo of chromosome evolution in rodents.

G- and C-banding delimits two cytogenetic groups within the vlei rat, Otomys irroratus. One has a diploid number of 2n = 24, resulting from a centric fusion of chromosomes 7 and 12 of the O. irroratus standard coupled with a tandem fusion to chromosome 8. The second has a diploid number of 2n = 28, lacks the compound chromosome, and appears to have a far wider geographic distribution within South Africa. Additionally, the two groups differ through the presence of cytotype-specific heterozygous centric fusions and one to three B chromosomes which appear as floating polymorphisms in the 2n = 28 complex.

Desmodillus and Gerbilliscus (formerly Tatera) comprise a monophyletic group of gerbils (subfamily Gerbillinae) which last shared an ancestor approximately 8 million years ago; diploid chromosome number variation among the species ranges from 2n = 36 to 2n = 50. In an attempt to shed more light on chromosome evolution and speciation in these rodents, we compared the karyotypes of 7 species, representing 3 genera, based on homology data revealed by chromosome painting with probes derived from flow-sorted chromosomes of the hairy footed gerbil, Gerbillurus paeba (2n = 36). The fluorescent in situ hybridization data revealed remarkable genome conservation: these species share a high proportion of conserved chromosomes, and differences are due to 10 Robertsonian (Rb) rearrangements (3 autapomorphies, 3 synapomorphies and 4 hemiplasies/homoplasies). Our data suggest that chromosome evolution in Desmodillus occurred at a rate of ∼1.25 rearrangements per million years (Myr), and that the rate among Gerbilliscus over a time period spanning 8 Myr is also ∼1.25 rearrangements/Myr. The recently diverged Gerbillurus (G. tytonis and G. paeba) share an identical karyotype, while Gerbilliscus kempi, G. afra and G. leucogaster differ by 6 Rb rearrangements (a rate of ∼1 rearrangement/Myr). Thus, our data suggests a very slow rate of chromosomal evolution in Southern African gerbils.

Pericentric inversions are important for evolutionary biology because of their potential role in speciation. They may result in reproductive isolation due to illegitimate pairing of homologues at meiosis which leads to the production of aneuploid gametes (containing deletions or duplications of chromosomal segments), and consequently mediate chromosomal divergence. In this study, we describe the prevalence of pericentric inversions in the African vlei rat, Otomys irroratus (OIR). The species is characterized by intraspecific chromosomal variation (2n = 23–32) across its distribution in southern Africa. Here, we analyzed 55 individuals collected from 7 localities in South Africa by G- and C-banding and chromosome painting with flow sorts of Myotomys unisulcatus. Of the 55 specimens that were analyzed, 47% contained inversions or centromeric shifts on 4 autosomes (OIR1, 4, 6 and 10) which were present singly in specimens (i.e. none of the specimens contained all 4 inversions concurrently). These inversions were found in both homozygous and heterozygous state over a wide geographic range suggesting that they are floating polymorphisms. Given the potential role of inversions in post-mating isolation (through production of aneuploid gametes), the prevalence of inversions as floating polymorphisms in the vlei rats suggests that they are probably retained in the population through suppression of recombination in the inverted regions of the chromosomes.

The postglacial recolonization of the pygmy shrew (Sorex minutus, n = 68) across continental Europe was examined using control region sequences (342 bp) and concatenated sequences (2,939 bp) of four paternally inherited introns, specific for the Y chromosome. Phylogenetic analysis (NJ, MP, ML, and Bayesian inference) of the control region retrieved five major lineages (Northern, Western, Italian, Balkan, and Spanish) in Europe, in agreement with the previous cytochrome b data. Our study identifies a new refugium for the pygmy shrew located in France during the Pleistocene glacial cycles and the most important refugium with regards to the recolonization of Eurasia was probably located in the Carpathians. In contrast, Y chromosome intron sequences retrieved four major lineages (Northern, Western, Italian, and Balkan). This may indicate contact between Western and Spanish mtDNA lineages after expansion from their respective refugia post-LGM. Our results also revealed differential introgression of mtDNA and Y chromosome markers at a local level when populations from different glacial refugia come into contact. These results highlight the importance of adopting a multiple marker approach when inferring phylogeographic structure and colonization history of species.