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Review of the Hylomyscus Denniae Group (Rodentia: Muridae) in Eastern Africa, with Comments on the Generic Allocation of Epimys Endorobae Heller

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Review of the Hylomyscus Denniae Group (Rodentia: Muridae) in Eastern Africa, with Comments on the Generic Allocation of Epimys Endorobae Heller PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 119(2):293–325. 2006. Review of the Hylomyscus denniae group (Rodentia: Muridae) in eastern Africa, with comments on the generic allocation of Epimys endorobae Heller Michael D. Carleton, Julian C. Kerbis Peterhans, and William T. Stanley (MDC) Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560-0108, U.S.A., e-mail: [email protected]; (JKP) University College, Roosevelt University, Chicago, Illinois 60605, U.S.A.; Department of Zoology, Division of Mammals, The Field Museum of Natural History, Chicago, Illinois 60605, U.S.A., e-mail: [email protected]; (WTS) Department of Zoology, Division of Mammals, The Field Museum of Natural History, Chicago, Illinois 60605, U.S.A., e-mail: [email protected] Abstract.—The status and distribution of eastern African populations currently assigned to Hylomyscus denniae are reviewed based on morpho- logical 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 Lo¨nnberg & 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. Until recently, Hylomyscus denniae, the highlands (cf. anselli group); through the Montane Woodland Mouse, was under- disconnected mountain chains in north- stood to be a highly variable murid ern Zambia and western Tanzania (an- species, broadly if patchily distributed in selli); to Mount Rungwe and the Eastern forested highlands across south-central Arc Mountains of central and eastern and eastern Africa (e.g., Misonne 1974, Tanzania (arcimontensis). In contrast, Bishop 1979, Musser & Carleton 1993). forms of the H. denniae assemblage are Carleton & Stanley (2005), however, centered in eastern African mountains demonstrated that nominal denniae masks (Fig. 1): the central portion of the Alber- a complex of species and perhaps two tine Rift system in easternmost Demo- species groups. Members of the H. anselli cratic Republic of Congo, Burundi, group occur in mountains of south- Rwanda, and southwestern Uganda (vul- central Africa: from the central Angolan canorum); astride the northern section of 294 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Mountainous regions of eastern Africa inhabited by populations of the Hylomyscus denniae complex, as circumscribed herein. The 12 geographic samples (OTUs) used in multivariate analyses are numbered and generally plotted; see Materials and Methods for specific localities comprising each and sample sizes. the Albertine Rift in northeastern Dem- fenestra (tiny or occluded in the denniae ocratic Republic of Congo and western- group; small to medium size in the anselli most Uganda (denniae); and mountains group), and occurrence of pectoral teats surrounding the Gregory Rift Valley in (present in the denniae group, total western Kenya (endorobae). Members of mammae 5 8; absent in the anselli group, the two groups differ in three qualitative total 5 6). characters: relative length of the incisive The primary purpose of the current foramina (longer in the denniae group, study is to evaluate morphological differ- extending between the first molars; entiation among populations of the H. shorter in the anselli group, terminating denniae complex, as a basis for comment- about the anterior border of the first ing on the status of the three forms named molars), patency of the subsquamosal from wet montane forest in eastern VOLUME 119, NUMBER 2 295 African mountains—denniae Thomas breadth of bony palate (BBP), measured (1906), endorobae Heller (1910), and across the maxillary bones above the vulcanorum Lo¨nnberg & Gyldenstolpe second molars; length of incisive foramen (1925). Our secondary goal is to review (LIF); length of diastema (LD); breadth the generic affinity of the Kenyan form of zygomatic plate (BZP); length of endorobae, which, although historically auditory bulla (LAB); coronal length of considered a form of Hylomyscus (e.g., maxillary toothrow (CLM); width of the Hatt 1940, Musser & Carleton 1993), has upper first molar (WM1). Five external been recently transferred to Praomys dimensions (to nearest whole mm) and (Van der Straeten & Robbins 1997). body mass (to nearest 0.5 gm) were transcribed from skin tags or field cata- Materials and Methods logs as given by the collector: total length (TOTL); head and body length (HBL); Specimens reported herein consist prin- tail length (TL); hindfoot length (HFL); cipally of skins with their associated ear (pinna) length (EL); and weight (WT). skulls and are contained in the following Most external data of the Burundi and museum collections: American Museum Uganda samples were recorded in the of Natural History, New York City field by JKP and WTS, reducing among- (AMNH); Carnegie Museum of Natural collector variability for these big, ex- History, Pittsburgh (CM); Field Museum tremely age-sensitive, dimensions. The of Natural History, Chicago (FMNH); large series from Mount Kenya was Museum of Comparative Zoology, Har- collected early in the 20th century by J. vard University, Cambridge (MCZ); Na- A. Loring and E. A. Mearns (Hollister tional Museum of Natural History, 1919), and specimens lack field-obtained Smithsonian Institution, Washington values for head-and-body length and D.C. (USNM, formerly the U.S. National weight; the former datum was obtained Museum); Natural History Museum of by subtraction of TL from TOTL. Re- Los Angeles County, Los Angeles cording of measurements was limited to (LACM); The Natural History Museum, animals judged to be adult, as based on London (BMNH, formerly the British the possession of fully erupted third Museum of Natural History); and the molars and adult pelage. Three crude Rijksuniversitair Centrum, Antwerpen, age classes of ‘‘adult’’ specimens were Belgium (RUCA). further identified based on degree of Fourteen cranial and two dental vari- upper molar wear, from little (young ables were measured by MDC to adult), to moderate (full adult), to heavy 0.01 mm, using hand-held digital calipers (old adult), generally following the pat- while viewing crania under a stereomicro- terns of coronal change outlined by scope. These measurements, and their Carleton & Martinez (1991). Except for abbreviations as used in text and tables, a priori exclusion of juvenile specimens, are (see Carleton & Van der Straeten 1999 we did not attempt to remove age-related for most landmarks): occipitonasal length size effects (post-weaning growth) in (ONL); greatest zygomatic breadth (ZB); portraying the multivariate results. Sexual breadth of braincase (BBC), measured size dimorphism was statistically unap- across the parietal flanges just behind the preciable in large population samples of zygomatic arches; breadth across occipital H. arcimontensis and H. denniae (Carle- condyles (BOC); interorbital breadth ton & Stanley 2005); measurements of (IOB); length of rostrum (LR); breadth males and females were accordingly of rostrum (BR); postpalatal length combined in analyses and tables. (PPL); length of bony palate (LBP); Throughout the text, we use the abbrevia- 296 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tions M1-3 or m1-m3 to individually Table 1.—External and cranial measurements of reference the upper (maxillary) and lower the holotypes of Epimys endorobae Heller and (dentary) molars, respectively; individual Epimys taitae Heller. Although closely similar in cusps of the M1-3 are referenced using the size, the former is a large example of the genus Hylomyscus (endorobae, here considered a valid positional t-system developed by Miller species) and the latter a small form of Praomys (1912). (taitae, currently a junior synonym of Praomys The following 12 operational taxonom- delectorum). ic units (OTUs) of Hylomyscus, arranged according to country of origin, were used Variable endorobae (USNM 162858) taitae (USNM 181797) to generate the various tabular summaries TOTL 251.0 243.0 and undertake descriptive and multivari- HBL 100.0 105.0 ate analyses. Many of these OTUs are TL 151.0 138.0 HFL 22.5 23.0 allopatric to one another, restricted to EL 19.5 19.0 wet montane forest in isolated mountain ONL 28.2 28.5 systems (Fig. 1). Locality numbers corre- ZB 13.6 13.1 spond to the geographic placenames and BBC 12.1 11.5 coordinates given in Appendix 1; full IOB 4.4 4.3 LR 9.1 9.9 provenance and museum registration BR 4.9 4.9 numbers are provided in the Taxonomic PPL 9.6 9.6 Summary. We included a sample of LBP 4.6 4.9 Praomys delectorum (Kenya, Taita Hills, BBP 5.3 5.3 Mounts Mbololo and Umengo, N 5 29) LIF 6.0 6.2 LD 7.7 7.9 to assess morphometric discrimination BZP 2.4 3.0 between large-bodied taxa of Hylomyscus LAB 4.5 4.6 (i.e., denniae, OTU 8, and endorobae, BOC 6.2 6.0 OTU 2) and a small form of Praomys. CLM 4.4 4.2 WM1 1.4 1.3 Abbreviations are used here and through- out the text for forest reserves (FR), national parks (NP), mount or mountains ity 34; N 5 12). Uganda: OTU 8— (Mt, Mts), and Democratic Republic of Ruwenzori Mts, Kasese District, Mubuku Congo (D.
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