Longevity of a solitary mole-rat species and its implications for the assumed link between sociality and longevity in African mole-rats (Bathyergidae).

Sociality and cooperative breeding are associated with enhanced longevity in insects and birds, but whether this is also true for mammals is still subject to debate. African mole-rats (Bathyergidae) have recently been claimed to be the only mammalian family in which such an association may exist because cooperatively breeding bathyergids seem to be substantially longer lived than solitary bathyergids. However, although ample longevity data are available for several social bathyergids, almost nothing is known about mortality distribution and lifespan in solitary bathyergids. Here we present robust long-term data on the longevity of a solitary African mole-rat, the silvery mole-rat Heliophobius argenteocinereus. Our findings show that this species is much longer-lived than previously believed. Nonetheless, our comparative analysis suggests that sociality has indeed a positive effect on longevity in this family. We argue that the extreme longevity seen particularly in social bathyergids is probably caused by a combination of subterranean lifestyle and cooperative breeding.

Species limits and phylogeographic structure in two genera of solitary African mole-rats Georychus and Heliophobius.

African mole-rats (Bathyergidae) are an intensively studied family of subterranean rodents including three highly social and three solitary genera. Although their phylogenetic interrelations are clear, genetic diversity and the number of species within each genus is much less certain. Among the solitary genera, Heliophobius and Georychus were for a long time considered as monotypic, but molecular studies demonstrated strong phylogeographic structure within each genus and proposed that they represent complexes of cryptic species. The present study re-evaluates their internal genetic/phylogenetic structure using a combination of methodological approaches. We generated datasets of one mitochondrial and six specifically selected nuclear markers as well as of a large number of double digest restriction site associated (ddRAD) loci and then applied species delimitation analyses based on the multispecies coalescent model or clustering on co-ancestry matrices. The population structure was largely congruent across all analyses, but the methods differed in their resolution scale when determining distinct gene pools. While the multispecies coalescent model distinguished five Georychus and between eleven to thirteen Heliophobius gene pools in both Sanger sequenced and ddRAD loci, two clustering algorithms revealed significantly finer or coarser structure in ddRAD based co-ancestry matrices. Tens of clusters were distinguished by fineRADstructure and one (in Georychus) or two clusters (in Heliophobius) by Infomap. The divergence dating of the bathyergid phylogeny estimated that diversification within both genera coincided with the onset of the Pleistocene and was likely driven by repeated large-scale climatic changes. Based on this updated genetic evidence, we suggest recognizing one species of Georychus and two species of Heliophobius, corresponding to a northern and southern major lineage, separated by the Eastern Arc Mountains. Yet, the final taxonomic revision should await integrated evidence stemming from e.g. morphological, ecological, or behavioral datasets.

Molecular systematics and biogeographic history of the African climbing-mouse complex (Dendromus).

Climbing mice in the genus Dendromus (sensu lato) are widely distributed in Africa, south of the Saharan Desert. The 17 currently recognized species in the genus range from widespread taxa to single-mountain endemics, and there is considerable variation across species with respect to habitats occupied. These habitats range from arid grasslands and savannahs to sub-alpine and alpine vegetation. Using the most comprehensive geographic and genetic survey to date and after reviewing many type specimens, we assess the systematics and biogeography of Dendromus. Given the structure of our molecular phylogenetic hypotheses, in which we recover six major clades, we propose the recognition of three genera within the Dendromus group (sensu lato): in addition to Dendromus (26 lineages), we suggest the retention of Megadendromus (monotypic) and the resurrection of the genus Poemys (six lineages). From our model-based molecular phylogenetic results and morphological comparisons, we suggest that six formerly synonymized taxa should be resurrected, and we highlight 14 previously undescribed lineages. We also constructed time-calibrations on our phylogeny, and performed ancestral area reconstructions using BioGeoBEARS. Based on fossil evidence, Dendromus appears to have had a widespread African distribution dating back to the Late Miocene (8-10 Ma), and our basal ancestral area reconstruction (Ethiopians Highlands + Eastern African Mountains + Zambezian region) supports this. Divergence of the six major clades we recover (Poemys, Megadendromus and four within Dendromus) occurred prior to or at the Miocene-Pliocene boundary 5.3 Ma. Biogeographically, Megadendromus is restricted to the Ethiopian Highlands. The ancestral area for Poemys is reconstructed as the Zambezian region, with species distributions ranging from South Africa to Western Africa. The ancestral area for Dendromus is reconstructed as the Ethiopian Highlands, with the ancestral areas of the four major clades being reconstructed as Ethiopian Highlands, Albertine Rift, South Africa or Western Africa. None of the four Dendromus clades are reciprocally monophyletic with respect to distributional area.

Multiple radiations of spiny mice (Rodentia: Acomys) in dry open habitats of Afro-Arabia: evidence from a multi-locus phylogeny.

BACKGROUND: Spiny mice of the genus Acomys are distributed mainly in dry open habitats in Africa and the Middle East, and they are widely used as model taxa for various biological disciplines (e.g. ecology, physiology and evolutionary biology). Despite their importance, large distribution and abundance in local communities, the phylogeny and the species limits in the genus are poorly resolved, and this is especially true for sub-Saharan taxa. The main aims of this study are (1) to reconstruct phylogenetic relationships of Acomys based on the largest available multilocus dataset (700 genotyped individuals from 282 localities), (2) to identify the main biogeographical divides in the distribution of Acomys diversity in dry open habitats in Afro-Arabia, (3) to reconstruct the historical biogeography of the genus, and finally (4) to estimate the species richness of the genus by application of the phylogenetic species concept. RESULTS: The multilocus phylogeny based on four genetic markers shows presence of five major groups of Acomys called here subspinosus, spinosissimus, russatus, wilsoni and cahirinus groups. Three of these major groups (spinosissimus, wilsoni and cahirinus) are further sub-structured to phylogenetic lineages with predominantly parapatric distributions. Combination of alternative species delimitation methods suggests the existence of 26 molecular operational taxonomic units (MOTUs), potentially corresponding to separate species. The highest genetic diversity was found in Eastern Africa. The origin of the genus Acomys is dated to late Miocene (ca. 8.7 Ma), when the first split occurred between spiny mice of eastern (Somali-Masai) and south-eastern (Zambezian) savannas. Further diversification, mostly in Plio-Pleistocene, and the current distribution of Acomys were influenced by the interplay of global climatic factors (e.g., Messinian salinity crisis, intensification of Northern Hemisphere glaciation) with local geomorphology (mountain chains, aridity belts, water bodies). Combination of divergence dating, species distribution modelling and historical biogeography analysis suggests repeated "out-of-East-Africa" dispersal events into western Africa, the Mediterranean region and Arabia. CONCLUSIONS: The genus Acomys is very suitable model for historical phylogeographic and biogeographic reconstructions of dry non-forested environments in Afro-Arabia. We provide the most thorough phylogenetic reconstruction of the genus and identify major factors that influenced its evolutionary history since the late Miocene. We also highlight the urgent need of integrative taxonomic revision of east African taxa.

Genetic distinction between contiguous urban and rural multimammate mice in Tanzania despite gene flow.

Special conditions are required for genetic differentiation to arise at a local geographical scale in the face of gene flow. The Natal multimammate mouse, Mastomys natalensis, is the most widely distributed and abundant rodent in sub-Saharan Africa. A notorious agricultural pest and a natural host for many zoonotic diseases, it can live in close proximity to humans and appears to compete with other rodents for the synanthropic niche. We surveyed its population genetic structure across a 180-km transect in central Tanzania along which the landscape varied between agricultural land in a rural setting and natural woody vegetation, rivers, roads and a city (Morogoro). We sampled M. natalensis across 10 localities and genotyped 15 microsatellite loci from 515 individuals. Hierarchical STRUCTURE analyses show a K-invariant pattern distinguishing Morogoro suburbs (located in the centre of the transect) from nine surrounding rural localities. Landscape connectivity analyses in Circuitscape and comparison of rainfall patterns suggest that neither geographical isolation nor natural breeding asynchrony could explain the genetic differentiation of the urban population. Using the isolation-with-migration model implemented in IMa2, we inferred that a split between suburban and rural populations would have occurred recently (<150 years ago) with higher urban effective population density consistent with an urban source to rural sink of effective migration. The observed genetic differentiation of urban multimammate mice is striking given the uninterrupted distribution of the animal throughout the landscape and the high estimates of effective migration (2Ne M = 3.0 and 29.7), suggesting a strong selection gradient across the urban boundary.

Emmonsiosis of subterranean rodents (Bathyergidae, Spalacidae) in Africa and Israel.

The presence of adiaspores of the fungal genus Emmonsia was examined in the lungs of 85 mole rats representing 3 subterranean genera: blind mole rats (Spalax galili and S. golani) from Israel, Ansell's mole-rats (Cryptomys anselli) from Zambia, and silvery mole-rats (Heliophobius argenteocinereus) from Malawi and Zambia. Emmonsiosis was found in 28% of the blind mole rats, 100% of the Ansell's mole-rats, but in none of the silvery mole-rats. Infection in African mole-rats was caused by Emmonsia parva, and infection in Israeli blind mole rats was caused by E. parva and E. crescens. The study indicates that the perennial burrow system of the Ansell's mole-rat forms an appropriate microhabitat for the saprophytic growth of E. parva in Lusaka region, Zambia. We suggest that factors contributing to the striking difference in prevalence of emmonsiosis between the two African mole-rat genera (Cryptomys, Heliophobius) may be their differing burrow types, burrow longevity, and social lives.

Three new species of Eimeria (Apicomplexa: Eimeriidae) from the silvery mole rat Heliophobius argenteocinereus Peters, 1846 (Rodentia: Bathyergidae) from Malawi.

Three species of Eimeria Schneider are described from feces of the African bathyergid rodent, Heliophobius argenteocinereus, from Malawi. Oocysts of Eimeria heliophobii n. sp. are broadly ellipsoidal; 27.9 (22-31) x 22.3 (18-24.5) microm with a brownish, heavily pitted oocyst wall, and vacuolar oocyst residuum. Sporocysts are oval, 12.8 (12-14) x 8.4 (8-9) microm with Stieda and substieda bodies. Eimeria nafuko n. sp. has subspherical oocysts; 15.5 (15-16) x 12.8 (12-13) microm with a smooth, colorless oocyst wall. Sporocysts are oval, 9.2 (9-10) x 5.3 (5-6) microm, with a small Stieda body; the substieda body is not visible. Oocysts of Eimeria yamikamiae n. sp. are broadly ellipsoidal to subspherical; 20.8 (19-22) x 17.5 (15.5-19) microm, with slightly yellowish, very faintly pitted oocyst wall. The majority of oocysts contained a single spherical vesicular oocyst residuum and numerous very small granules. Sporocysts are oval, 10.7 (10-11) x 6.8. (6-7) microm, with a dome -like Stieda body and a subspherical to lentil-like substieda body. Typically, infected rodents shed oocysts of more than 1 species of Eimeria.

Eimeria burdai sp. n. (Apicomplexa: Eimeriidae), a new parasite species from subterranean African silvery mole-rat, Heliophobius argenteocinereus.

A new coccidian parasite of the genus Eimeria Schneider, 1875 is described from the subterranean African silvery mole-rat Heliophobius argenteocinereus Peters, 1846. Oocysts of Eimeria burdai sp. n. were subspherical to broadly ellipsoidal 17.8 (16-19) x 14.1 (12-15), with a shape index 1.2 (1.1-1.4). Oocyst wall was bilayered, smooth and colourless, approximately 1.0 thick. Outer layer was significantly thicker than inner one. A micropyle and oocyst residuum were absent. One or two ellipsoidal or spherical polar granules were present. Sporocysts were ellipsoidal, 10.8 (9-12) x 6.2 (5-8) with a shape index 1.7 (1.5-1.9). Sporocyst wall was single-layered, thin, smooth and colourless, with small Stieda body at the pointed end. In freshly sporulated oocysts, spherical sporocyst residuum was composed of small granules enclosed by a thin membrane. Sporozoites were elongate, lying length-wise in the long axis of the sporocyst, partially curled around each other, with single large refractile body located posteriorly. Faintly distinguishable nucleus was in the central part of the sporozoite. This eimerian represents the first coccidian species described from subterranean African silvery mole-rat (Rodentia: Bathyergidae).