William T. Stanley

A New Genus of African Monkey, Rungwecebus: Morphology, Ecology, and Molecular Phylogenetics

A new species of African monkey, Lophocebus kipunji, was described in 2005 based on observations from two sites in Tanzania. We have since obtained a specimen killed by a farmer on Mount Rungwe, the type locality. Detailed molecular phylogenetic analyses of this specimen demonstrate that the genus Lophocebus is diphyletic. We provide a description of a new genus of African monkey and of the only preserved specimen of this primate. We also present information on the animals ecology and conservation.

A new species of Congosorex from the Eastern Arc Mountains, Tanzania, with significant biogeographical implications

A new species of Congosorex is described. The new species occurs in Tanzania and is larger in external characteristics than the other two known species in the genus, with the longer tail being especially distinctive. The skull of the new species is wider, on average, than other Congosorex, but intermediate in greatest length. Previously undocumented characteristics that differentiate Congosorex from Myosorex are described. The presence of Congosorex in the Eastern Arc Mountains supports the theory that these mountains are refuges of ancient lineages of organisms with ties to the Guineo-Congolian region of Africa and that there was once a continuous swath of forest connecting these two areas.

Review of the Hylomyscus denniae complex (Rodentia: Muridae) in Tanzania, with a description of a new species

ABSTRACT Based on morphological and morphometric comparisons, a new species of Hylomyscus, H. arcimontensis (Muridae: Murinae), is identified from Mount Rungwe and the Eastern Arc Mountains of central and eastern Tanzania. The new species is confined to wet montane forests of these mountains and represents another vertebrate endemic to the Tanganyika-Nyasa Montane Forest biotic region (sensu Moreau 1966). It is most closely related to H. anselli, a form described from mountains in northern Zambia as a subspecies of H. denniae and here elevated to species rank. Morphological evidence supports the view that the nominal species H. denniae is a composite of several species whose interrelationships and differentiation patterns indicate two species complexes: the H. denniae group centered in Central East African mountains (Burundi, Kenya, Rwanda, Uganda, Zaire) and the H. anselli group distributed across more southern mountains (Angola, Malawi, Tanzania, Zambia).

Phylogeny, phylogeography, and geographic variation of Sylvisorex howelli (Soricidae), an endemic shrew of the Eastern Arc Mountains, Tanzania

The Eastern Arc Mountains of eastern Africa are notable for the high levels of endemism exhibited by various forest-dwelling organisms of this ancient montane archipelago. There has been virtually no assessment of the variation among populations of small mammal species living on these unique mountains, but recent faunal surveys have produced sufficient material to initiate such studies. Cranial morphometric and DNA sequence data were examined from six populations of Sylvisorex howelli Jenkins, 1984, an endemic shrew found in several different massifs of the Eastern Arc Mountains, to assess variation across the archipelago in the context of various hypotheses of historical biogeography. Twenty-two cranial measurements were analysed using principal components analysis. Age classes (based on tooth wear) and sex had little effect on the variation exhibited by the variables studied. Overall, specimens of S. howelli from the East Usambara Mountains are smaller than specimens from other known populations. The mitochondrial ND2 and 12S rRNA genes from representatives of each montane population of S. howelli in addition to several crocidurine taxa from eastern Africa and three soricine outgroup species were sequenced to assess phylogenetic relationships among these taxa. Neither maximum likelihood, maximum parsimony, nor Bayesian analyses support monophyly of the genus Sylvisorex, but S. howelli populations were consistently recovered as a well-supported clade. Over 40 individuals of S. howelli from six disjunct montane ranges, comprising the entire known distribution of the species, were sequenced for 504 base pairs of ND2 to investigate phylogeographic patterns. Phylogenetic analysis recovered six reciprocally monophyletic haplotype clades grouped by locality. Branch lengths are consistent with relatively long periods of isolation among populations from the Uluguru, Ukaguru, Nguru, Nguu, East Usambara and West Usambara Mountains, with low levels of diversity observed within each population. These results are interpreted within the historical context of the Eastern Arc Mountains.

Small mammals of the Eastern Arc Mountains, Tanzania

ABSTRACT The Eastern Arc Mountains (EAM) contain some of the most biologically diverse and endemic-rich montane ecosystems in all of Africa. Because of the staggering degree of biodiversity, how little we know about the biota of the EAM and the rapid degradation of the remaining montane forests of these mountains, we are studying the natural history and biogeography of the small mammals (shrews and rodents) of this archipelago. We are also interested in specific questions such as the effects of forest fragmentation on and elevational distribution of EAM small mammals. To date we have surveyed the South Pare, East Usambara, West Usambara, Nguru, Uluguru and Udzungwa Mountains. We have documented a rodent fauna that is widespread across the archipelago, although restricted to montane and submontane habitats. Shrew species are much more patchy in distribution with each EAM or subset of mountains holding at least one seemingly endemic species, suggesting that speciation has contributed to the current pattern of species richness in insectivores. However, further analysis is required to discern how dispersal, extinctions and/or speciation have contributed to this pattern. Importantly, we have found very few introduced, exotic or savannah dwelling species in our surveys of undisturbed forests, even in forests in close proximity to disturbed areas or human habitation.

The biogeography of introgression in the critically endangered African monkey Rungwecebus kipunji

In the four years since its original description, the taxonomy of the kipunji (Rungwecebus kipunji), a geographically restricted and critically endangered African monkey, has been the subject of much debate, and recent research suggesting that the first voucher specimen of Rungwecebus has baboon mitochondrial DNA has intensified the controversy. We show that Rungwecebus from a second region of Tanzania has a distinct mitochondrial haplotype that is basal to a clade containing all Papio species and the original Rungwecebus voucher, supporting the placement of Rungwecebus as the sister taxon of Papio and its status as a separate genus. We suggest that the Rungwecebus population in the Southern Highlands has experienced geographically localized mitochondrial DNA introgression from Papio, while the Ndundulu population retains the true Rungwecebus mitochondrial genome.

The biogeography of Miniopterus bats (Chiroptera: Miniopteridae) from the Comoro Archipelago inferred from mitochondrial DNA

The endemic fauna of the Comoro Archipelago is composed of a mixture of taxa originating from Africa and Madagascar. Bats are the only native land dwelling mammals on this archipelago, but the biogeographical origins for the vast majority of species within this group are ambiguous. We report here genetic analyses based on two mitochondrial DNA markers to infer the origin of Comorian bats belonging to a reputed species complex of Miniopterus that is further distributed across Africa and Madagascar. Phylogenetic reconstructions show that east African M. minor are not closely related to the insular Miniopterus of Madagascar and the Comoros (Grande Comore and Anjouan). The latter cluster into two distinct, monophyletic clades (Clade 1 and Clade 2). Representatives of these clades occur sympatrically both on the Comoros and on Madagascar, and are distinguished by a large genetic distance (K2P: 9.9% for cytochrome b). No haplotypes are shared between any islands, suggesting the absence of contemporary gene flow. Populations of the widespread Clade 1 are furthermore characterized by a significant inter‐island structure (ΦCT = 0.249), and by high haplotype and nucleotide diversities (h = 0.90–0.98, π = 0.04–0.06). Demographic analyses of Clade 1 suggest secondary contact between two distinct phylogroups (Subclade 1 A and 1B) that reached Grande Comore and Anjouan, and a large, stable population with a long evolutionary history on Madagascar. These results and the current distribution of related lineages suggest that the Comoros were colonized independently at least two or three times by ancestors from Madagascar.

Additional molecular evidence strongly supports the distinction between the recently described African primate Rungwecebus kipunji (Cercopithecidae, Papionini) and Lophocebus

Department of Mammalogy, University of Alaska Museum, 907 Yukon Drive, Fairbanks, AK 99775, USA b Institute of Arctic Biology, University of Alaska Fairbanks, Fairbanks, AK 99775, USA Department of Anthropology, Yale University, P.O. Box 208277, New Haven, CT 06520-8277, USA Division of Vertebrate Zoology, Peabody Museum of Natural History, Yale University, New Haven, CT 06520-8118, USA Department of Zoology, Field Museum of Natural History, 1400 S Lake Shore Drive, Chicago, IL 60605, USA Wildlife Conservation Society, Tanzania Program, P.O. Box 1475, Mbeya, Tanzania

Differences in abundance and species richness between shrews and rodents along an elevational gradient in the Udzungwa Mountains, Tanzania

Small mammals (shrews and rodents) were surveyed along an elevational transect in the Udzungwa Scarp Forest Reserve, in the Udzungwa Mountains, Tanzania. Trap lines and pitfall lines were installed at 600, 910, 1460, and 2000 m a.s.l. In a total of 10341 sample nights (7448 trap-nights and 2893 bucket-nights) 343 specimens (148 shrews, 205 rodents) were captured representing 9 shrew and 14 rodent species for a total of 23 species. While overall species diversity generally increased with elevation, this pattern was not constant for each group sampled. For rodents, both species richness and abundance were lowest at 600 m and greatest at 2000 m a.s.l., and were significantly correlated with elevation. While the highest species number and abundance for shrews was at 2000 m, there was no correlation of these two values with elevation. Rainfall appears to have affected the capture of shrews, but not rodents, and capture success of individual buckets and traps indicated a lack of capture independence. Eastern Arc endemics such asCrocidura desperata Hutterer, Jenkins and Verheyen, 1991 andMyosorex kihaulei Stanley and Hutterer, 2000 were more abundant at 2000 m a.s.l., than at lower elevations. Implications of results of this survey for analyses of future biotic surveys are discussed.

Review of the Hylomyscus denniae group (Rodentia: Muridae) in eastern Africa, with comments on the generic allocation of Epimys endorobae Heller

ABSTRACT The status and distribution of eastern African populations currently assigned to Hylomyscus denniae are reviewed based on morphological and morphometric comparisons. Three species are considered valid, each confined largely to wet montane forest above 2000 meters: H. denniae (Thomas, 1906) proper from the Ruwenzori Mountains in the northern Albertine Rift (west-central Uganda and contiguous D. R. Congo); H. vulcanorum Lönnberg & Gyldenstolpe, 1925 from mountains in the central Albertine Rift (southwestern Uganda, easternmost D. R. Congo, Rwanda, and Burundi); and H. endorobae (Heller, 1910) from mountains bounding the Gregory Rift Valley (west-central Kenya). Although endorobae has been interpreted as a small form of Praomys, additional data are presented that reinforce its membership within Hylomyscus and that clarify the status of Hylomyscus and Praomys as distinct genus-group taxa. The 12 species of Hylomyscus now currently recognized are provisionally arranged in six species groups (H. aeta, H. alleni, H. anselli, H. baeri, H. denniae, H. parvus) based on 8 qualitatitive characters. Biogeography of the three species of the H. denniae group is discussed in the context of broad distributional patterns and area relationships evident among other terrestrial small mammals also confined to the Afromontane biotic region in eastern Africa.