AMERICAN MUSEUM Norntates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3157, 44 pp., 14 figures, 8 tables February 14, 1996 Systematic Revision of Sub-Saharan African Dormice (Rodentia: Myoxidae: Graphiurus) Part 1: An Introduction to the Generic Revision, and a Revision of Graphiurus Surdus1 MARY ELLEN HOLDEN2 ABSTRACT African dormice, genus Graphiurus (Rodentia: of Graphiurus surdus, represents the first distilla- Myoxidae), occur from the Cape of Good Hope tion of the large amount of data that has been to Senegal. The genus has never been adequately collected. The completed revision ofthe genus will revised, and thus the species diversity within the form one of the few comprehensive taxonomic genus, interspecific relationships, and biogeogra- revisions for any group ofAfrican rodents in which phy have not been documented. This report con- the intent is to present a reliable estimate ofspecies stitutes the first of seven papers whose collective diversity and document patterns of geographical purpose is to provide a morphological revision of distributions. It will provide a framework for the the genus, explore the phylogenetic relationships future tasks oftesting hypotheses of species limits of species or species groups within the genus, and and interspecific relationships within Graphiurus document biogeographic patterns inferred from using molecular and perhaps other anatomical data, species distributions and relationships. Over 2700 and determining the significance of the docu- museum specimens ofAfrican dormice have been mented geographic distributional patterns relative examined and measured from institutions in the to distributions of other African mammals and United States, Europe, and Africa for this study. environmental associations. This introductory paper, that includes a revision This first report contains an introduction to the 1 This report is dedicated to Guy Graham Musser, who ignited my interest in mammalogy, and provided support and inspiration for my research endeavors. 2 Instructor, Department of Biology, Orange County Community College, 115 South Street, Middletown, New York, 10940; Research Associate, Department of Mammalogy, American Museum of Natural History. Copyright © American Museum of Natural History 1996 ISSN 0003-0082 / Price $5.60 2 AMERICAN MUSEUM NOVITATES NO. 3157 entire revision, including an overview of the in- ences between the two similarly sized species. Al- tended scope and organization, and a review of though the geographic range ofboth extends from historic taxonomic treatments of species of Gra- southern Cameroon to northeast and south-central phiurus. A species account of Graphiurus surdus, Zaire, the two have never been taken at the same and a comparison of that species with G. christyi collecting locality, and may have different micro- and G. lorraineus, follows. Graphiurus surdus was habitat requirements. previously known only from southern Cameroon; Comparisons are also made between G. surdus its range is shown to extend at least to northeast and G. lorraineus, as G. surdus has previously been and central Zaire. Graphiurus schwabi, previously synonymized with G. lorraineus as a subspecies of thought to be a synonym of G. lorraineus, is al- G. murinus. Graphiurus lorraineus is not allied located as a junior synonym of G. surdus. Gra- with the G. murinus complex, and is sympatric phiurus surdus has alternately been recognized as with representatives of that assemblage through- a valid species, included in the problematic species out its range, including G. christyi. Sympatry of complex G. murinus, or obscured along with G. G. lorraineus with G. surdus in southern Came- lorraineus under G. murinus lorraineus. roon and central Zaire is documented. Compari- Comparisons with G. christyi, a member of the sons between the two species show that they are G. murinus complex from the same geographic easily separated by significant size and shape dif- area, show that there are significant shape differ- ferences. RESUME Les loirs africains du genre Graphiurus (Roden- travail et une recapitulation historique de la po- tia: Myoxidae) ont une distribution geographique sition taxonomique des especes de Graphiurus. Suit allant du Cap au Senegal. Ce genre n'a jamais ete une redefinition specifique de G. surdus, et une revise de facon adequate. Par consequent, ni la comparaison de cette espece avec G. christyi et G. diversite specifique intragenerique, ni les relations lorraineus. interspecifiques non plus que la biogeographie ne Jusqu'alors, G. surdus etait connu uniquement sont bien documentees. Ce rapport est la premier au Sud du Cameroun mais nous avons demontre d'une serie de sept publications decrivant la mor- que son aire de distribution s'etend au moinsjusq- phologie du genre Graphiurus, les relations phy- u'aux regions centrales et du Nord-Est du Zaire. logenetiques entre especes ou groupes d'especes, Graphiurus surdus a et6 successivement retenu ainsi que les modeles biogeographiques qui de- comme espece valide, inclu dans le complexe spe- coulent des distributions specifiques et des rela- cifique douteux de G. murinus, ou regroupe comme tions phylogenetiques. sous-espece avec G. lorraineus. Graphiurus schwa- Pour realiser cette etude, plus que 2700 speci- bi doit etre mis en synonymie avec G. surdus. mens conserves dans des institutions scientifiques Des comparaisons avec G. christyi, un membre americaines, europeennes, et africaines ont ete ex- du complexe specifique de G. murinus, habitant amines et mesures. le meme region geographique et de meme taille, Cette introduction, qui comprend une revision ont montre qu'il y a des difflerences morpholo- de Graphiurus surdus, doit etre consideree comme giques importantes. Bien que la distribution geo- le premier "distillat" de la masse de donnees dont graphique de ces deux especes couvre une region nous disposons. La revision complete du genre allant du Sud du Cameroun au Nord-Est du Zaire, formera une des rares etudes taxonomiques ex- elles n'ont jamais ete recoltees dans la meme lo- haustives d'un groupe de rongeurs africains dans calite mais pourraient exiger des microhabitats dif- le but d'en presenter une evaluation fidele de sa ferents. diversite specifique et de formuler des schemas de Des comparaisons de G. surdus et de G. lorrai- distribution geographique. Elle fournira le cadre neus, s'imposaient aussi parce que le premier a ete approprie pour tester les hypotheses futures con- mis en synonymie et regroupe avec le second sous cernant les limites specifiques et les relations in- le nom de G. murinus lorraineus. Toutefois, G. terspecifiques dans le genre Graphiurus a l'aide de lorraineus ne peut etre inclu dans le complexe G. donnees moleculaires et, peut-etre d'autres don- murinus. Il est sympatrique, sur toute son aire de nees anatomiques, ainsi que pour evaluer la valeur distribution, des representants de ce groupe d'es- des modeles biogeographiques par rapport 'a ceux peces, y compris G. christyi. La sympatrie de G. d'autres Mammiferes africains et les associations lorraineus et de G. surdus au Sud du Cameroun et avec l'environnement. au Zaire central a ete demontree. Ces deux especes Ce premier texte comprend outre une introduc- peuvent etre distinguees aisement par des diffTer- tion 'a la revision globale, un resume du plan de ences de taille et de forme. 1 996 HOLDEN: SUB-SAHARAN AFRICAN DORMICE. PART 1 3 INTRODUCTION Recent quality systematic revisions of in- sons for following the conceptual approach adequately studied groups often result in the taken in this revision are detailed below un- recognition of a greater diversity of species der Materials and Methods. in those groups than was previously acknowl- Workers who adopt the same theoretical edged. For some smaller and relatively well- species concept may obtain different esti- known groups, such as mammals, species di- mates of species diversity for a given group, versity estimates might produce higher num- because there are often several subjective es- bers and be made more accurate by focusing timates of diversity that can be justified by more attention on recent systematic revisions differing operational practices consistent with in addition to the relatively few descriptions the same theoretical concept. Therefore, the of new taxa that are published each year. operational concept followed in a given work These estimates may then provide finer doc- should be clearly stated. In addition, it is es- umentation of biogeographic patterns, and sential that adequate data are presented for aid in assessing conservation priorities. reasonable geographic samples (populations) Objections might be made that estimates of each alleged species, if such samples are of species diversity depend on the particular available. Researchers who adopt different species concepts used by the authors of sys- operational or theoretical species concepts tematic revisions. The same arguments apply may then make use ofdata contained in such to descriptions of new species; one worker revisions and possibly build upon them, even might reason that a "new species" is simply if they reject the overall estimate of species a diagnosable population that should not be diversity, or species diagnoses. recognized as a separate species, whereas oth- New species ofsmall African mammals are ers may feel that the population should be described each year (for example, look at spe- recognized as a species precisely because it is cies accounts