ARTICLE IN PRESS www.elsevier.de/mambio Short communication The distribution of Poelagus marjorita (Lagomorpha: Leporidae) in central Africa By D.C.D. Happold and W. Wendelen School of Botany and Zoology, Australian National University, Canberra, A.C.T., Australia and Musée Royale d’Afrique Centrale, Tervuren, Belgium Receipt of Ms. 9.10.2005 Acceptance of Ms. 20.4.2006 Key words: Poelagus marjorita, Lagomorpha, distribution, Africa The ‘‘Bunyoro rabbit’’ or ‘‘Uganda grass hare’’ was described as Lepus marjorita by St. Leger (1929) on the basis of the characteristics of the body and skull. The type series, of four skins and skulls, was collected by Captain C. R. S. Pitman at Masindi, Uganda (01.411N, 31.431E). Three years later, in 1932, St. Leger received a further 11 specimens comprising the skins, skulls and skeletons. On the basis of the skeletal structure, which exhibits characteristics of both hares (Lepus) and rabbits (Oryctolagus, Sylvilagus) and Rock-hares (Pronolagus), St. Leger (1932) described the genus Poelagus, designating Poelagus marjorita as the type species. Corbet (1983) provided a table of 21 character-states for 22 species of leporids, including P. marjorita. Two subspecies have been described: P. m. larkeni from near Diawo, southern Sudan (ca. 06.001N, 29.001E) by St. Leger (1935) and P. m. oweni from Lotti Forest, Imatong Mountains, southern Sudan (04.021N, 32.331E) by Setzer (1956); these subspecies show minor differences in pelage colour and skull from the nominate subspecies P. m. marjorita from Uganda. Neither taxon is currently recognised as a subspecies. Poelagus has been placed as a subgenus of Pronolagus (Ellerman and Morrison-Scott 1951), and as a subgenus of Caprolagus (a monotypic genus from the southern foothills of the Himalaya Mountains; Gureev 1964, cited by Hoffman 1993, Hoffman and Smith 2005). However, because Poelagus has a combination of rabbit-like and hare-like characteristics, it has been retained as a valid genus, with a single species, by Angermann (1966), Corbet (1983), Hoffman (1993), and Hoffman and Smith (2005). Little is known about the biology of P. marjorita, the most detailed study being that of Kingdon (1974). The preferred habitat is woodland savanna and open grassy clearings where trees are dense. Bunyoro rabbits are terrestrial and primarily nocturnal. During the day, individuals rest alone in a form in thick vegetation. Locomotion is more similar to that of a rabbit than of a hare. At night, they forage on flowers and sprouting grasses, preferring pastures that have been heavily grazed by larger mammals, and burnt areas where the grasses are sprouting. They may also be found on rocky inselbergs with rock hyraxes. Accounts from Uganda in the 1920s indicated that they were abundant in certain localities at night grazing on grassy tracks and roads, and very common grazing by roads at night (specimen labels, BMNH). In the 1960s, they were considered as ‘‘not uncommon in north-western Uganda, including the Murchison Falls National Park’’ and ‘‘could always be seen on the 1616-5047/$ - see front matter r 2006 Deutsche Gesellschaft für Säugetierkunde. Published by Elsevier GmbH. All rights reserved. Mamm. biol. 71 (2006) 6
377–383 doi:10.1016/j.mambio.2006.04.004 ARTICLE IN PRESS 378 D.C.D. Happold, W. Wendelen Masindi-Butiaba road after dark’’ (Williams 1967). The geographic distribution of P. marjorita is not well known. This study investigates and analyses the locality records in the literature and specimens in museums, and has resulted in a new distribution map for the species. Prior to this study, the only maps of the geographical distribution of P. marjorita were those of Kingdon (1974), Duthie and Robinson (1990), and Kingdon (1997). The maps in the two latter publications are essentially copied from Kingdon (1974). Duthie and Robinson (1990) describe the distribution as ‘‘from southern Chad eastwards through southern Sudan, northeastern Zaire, northwestern Kenya and southwards to the northern tip of Lake Tanganyika. If the current distribution maps are accurate and do not reflect a sampling hiatus, an isolated population would appear to exist in Angola, with a distribution extending from west-central Angola northwards into southern Zaire’’. The (small) map in Kingdon (1974), which shows rivers but not country boundaries, indicates a cluster of 12 localities (as solid circles) centred around Uganda, southern Sudan and northeast Zaire. In addition, there are three isolated single localities: one in eastern Central African Republic, one at the northeastern tip of Lake Tanganyika (in Burundi), and one in Angola south of the Cuanza River and inland from the coast. The presumed range of the species is indicated by stippling; one range links all the locality records in eastern Africa and extends westwards to a point somewhere near the Central African Republic-Cameroon border; a second range is south of the Zaire River in Angola and southern Zaire as described above. The map in Duthie and Robinson (1990) shows only ranges (without any localities) and is very similar to that in Kingdon (1974). Comparison of these published maps with data from the literature and from specimens in museums showed a number of anomalies which significantly reduce the range of the species. The distribution of the species for each country is given below. Uganda: Many of the locality records and specimens come from Uganda (Tab. 1). This country appears to be the central focus of the range (and probably the most important country in terms of conservation of the species). Duthie and Robinson (1990) show the range as most of the country except the southeast. The only specimens in museums (Tab. 1) are from the west-central part of the country in an area bounded in the north by the Victoria Nile and Lake Albert, and extending no further south than about 11N (Tab. 1, Fig. 1). However, the species is probably more widespread than the specimens suggest: Kingdon (in litt. 2005) writes that it ‘‘occurs very patchily but quite extensively throughout northern Karamoja, Acholi, Bunyoro, Mubende [and] parts of Toro and Ankole’’ Districts. These Districts cover a large area from the northeast to the southwest of the country – a much more extensive area than is indicated by museum specimens – but excludes the northwest (West Nile and Madi Districts) and most of the southeast. It is surprising that specimens have not been collected more extensively, and that the presence of the species in many parts of Uganda relies only on sight records. It may be that the species is much less abundant in some parts than in others; Kingdon (in litt. 2005) records that it is ‘‘extremely abundant in certain localities and during certain years, possibly going through boom and bust cycles’’. A specimen collected by Krampitz (1968) from near Tororo (00.371N, 34.101E) extends the range into southeast Uganda, east of the Victoria Nile. This specimen (originally identified by B. Foster, and said to be in the National Museum, Nairobi) has not been examined by us (and hence is shown as a literature record in Fig. 1). Currently the species is present and conserved in two of Uganda’s National Parks – Murchison Falls National Park (ca. 02.101N, 31.451E) and Kipedo Valley National Park (ca. 04.001N, 34.001E) (Wilson 1995). Central African Republic: Duthie and Robinson (1990) included the whole of the Central African Republic in their map. However, there seem to be only three definitive locality records (confirmed by specimens), all in eastern Central African Republic (Tab. 1, Fig. 1). The earliest published evidence for the occurrence of the species is a comment by ARTICLE IN PRESS The distribution of Poelagus marjorita 379 Table 1. Locality records for Poelagus marjorita, arranged by country. BMNH=The Natural History Museum, London, UK; CM=Carnegie Natural History Museum, Pittsburgh, USA; MCZ=Museum of Comparative Zoology, Harvard, USA; FMNH=Field Museum of Natural History, Chicago, USA; MNHN=Museum National d’Histoire Naturelle, Paris, France; MRAC=Musée Royale d’Afrique Centrale, Tervuren, Belgium; SMNS=Staatliches Museum für Naturkunde, Stuttgart, Germany; USNM=United States National Museum, Washington, D.C, USA. Locality Country Coordinates Source La Koumbala [=Koumbal] Central African Republic Central African Republic Central African Republic Sudan Sudan Sudan Sudan Sudan 09.201N, 22.401E MNHN 08.001N, 22.251E MNHN 06.331N, 23.101E MNHN ca. 06.001N, 29.001E 04.101N, 32.411E 04.191N, 32.241E 04.021N, 32.471E 04.021N, 32.331E BMNH USNM MCZ USNM CM Sudan Sudan Sudan 04.021N, 32.331E — 04.241N, 32.171E USNM FMNH MCZ Sudan Sudan Sudan 04.401N, 29.451E ca. 06.001N, 29.001E BMNH Sudan Sudan Uganda Uganda Uganda ca. 06.001N, 29.001E 04.001N, 32.441E 02.071N, 31.251E ca. 01.301N, 32.051E 01.231N, 32.271E BMNH SMNS BMNH AMNH AMNH Uganda Uganda Uganda 01.491N, 31.191E ca. 02.161N, 32.031E ca. 01.301N, 31.301E BMNH BMNH BMNH Uganda 01.411N, 31.431E BMNH Ouadda Yalinga Sud Ibba R., 100 miles N Diawa Imela Imurok, 20 miles SE Torit Katire Lotti Forest, Imatong Mts Lotti Forest Lower Imatong Mts Magwe (5 miles N)= 36 miles S Torit Mt. Baginzi Sue R, near Diawo 100 miles N of Diawo, Yambio District, Bahr-elGhazal Province  Ibba R., 100 miles N Diawo Talanga Forest, Imatong Mts Bulisa, NE Lake Albert Nakitoma, Buruli Co Nakasongola, Buruli, Buganda Butiaba Chopi [=Chobi] Isimba Hills, 8 miles S. Masindi Masindi, Bunyoro Kyjanjubija, S of Masindi, Bunyoro Abimva Bagbele, Garamba Uganda ca. 01.301N, 31.451E BMNH Zaire (DRC) Zaire (DRC) 03.091N, 29.501E ca. 04.001N, 29.301E MRAC MRAC Biadimbi Faradje Zaire (DRC) Zaire (DRC) 04.131N, 29.211E 03.441N 29.431E MRAC AMNH, MRAC Mabanga Zaire (DRC) 04.221N, 29.471E MRAC png pfsk/8, Garamba Zaire (DRC) 04.201N, 29.511E MRAC References and notes Petter (1959, 1972) Setzer (1956) Setzer (1956) Setzer (1956) Setzer (1956) Type of P. m. oweni Setzer (1956) Hoogstraal (1956) Setzer (1956) Setzer (1956) St. Leger (1935) Type of P. m. larkeni NW of Nakasongola St. Leger (1929) Type of P. m. marjorita = Kijunjubwa Schouteden (1948) Verheyen and Verschuren 1966 Hatt (1940), Verheyen and Verschuren 1966, Schouteden 1948 Verheyen and Verschuren 1966 Verheyen and Verschuren 1966  St. Leger (1935) recorded the locality as:‘‘100 miles north of Diawo, Yambio District, SW Bahr-elGhazal’’. She did not give any geographical coordinates, but it is calculated to be ca 06.001N, 29.001E, and probably on or near the border with Equatoria Province. ARTICLE IN PRESS 380 D.C.D. Happold, W. Wendelen Fig. 1. Distribution map for museum specimens and published records of Poelagus marjorita. X=specimen(s). =literature records (see text for further details). In the areas south of the Victoria Nile in Uganda, and in the Imatong Mts of S Sudan, a single X may represent more than one collecting locality. Malbrant (1952) that ‘‘l’habitat se trouverait confiné au Sud, et probablement au Sud-Est du Centre africain (Nord-Est de l’Oubangui et sans doubt Sud-East de ce territoire)’’. Malbrant (1952) did not cite any localities. However, Petter (1959b) wrote that the species ‘‘existe certainement dans la region de Fort Sibut – Fort Crampel’’. These two localities (now Sibut [05.461N, 19.101E] and Kaga Bandoro [07.001N, 19.101E]) are further west than the three definite locality records, and are roughly in the centre of the country (Fig. 1). Specimens from Fort Sibut and Fort Crampel have not been located. Thus in the Central African Republic, the species is assumed to be confined to the eastern part of country and may extend to the central part, but is not known to extend to the border with Cameroon. Chad: Presence in Chad seems dubious. No specimens have been located from this country. Petter (1959b), referring to Malbrant (1952), wrote that the habitat for P. marjorita is woodland savanna and open forest between 41N and 81N in ‘‘Oubangui and Tchad’’. These terms are assumed to refer to Provinces in the former French Equatorial Africa before the formation of the modern States of Central African Republic and Chad. Hence this reference to ‘‘Tchad’’ refers to the extent of the habitat of the species, and does not refer to the modern State of Chad, nor to the distribution of the species. On the present evidence, we do not include Chad in the distribution of the species. Rwanda and Burundi: The range in Duthie and Robinson (1990) includes west Rwanda and west Burundi as far as the northern tip of Lake Tanganyika. Kingdon (1974) shows a single locality at the NE tip of Lake Tanganyika (i.e. in Burundi), but no other localities in Rwanda or Burundi. There are specimens originally labelled as P. marjorita from Rwanda and Burundi in the Musée Royale d’Afrique Centrale at Tervuren ARTICLE IN PRESS The distribution of Poelagus marjorita (Belgium). However, these specimens (MRAC 20034–20044 from Kibungu, Rwanda; MRAC 13884–5 from Nyakatare, Rwanda; and MRAC 20039 from Rwinkwavu, Rwanda) were shown by Petter (1959a) to belong to Lepus. Analysis of the craniometric data of these specimens, together with known specimens of Lepus and P. marjorita in the Musée Royale d’Afrique Centrale, confirmed Petter’s (1959a) determinations. In addition, when the skins of these specimens were examined, the ear length was similar to that of Lepus (ca. 120 mm) and unlike that of Poelagus (ca. 60 mm). Montfort (1992) correctly does not list P. marjorita as occurring in Rwanda. On this evidence, Rwanda and Burundi are excluded from the range, so that the southernmost confirmed locality of the species is in Uganda (at ca. 01.001N). Kenya: We are unable to find any locality records or specimens from NW Kenya. The map in Duthie and Robinson (1990) appears to show that the range extends to the western shores of Lake Turkana. It is possible that the population in Kipedo National Park in north Uganda may extend marginally into northern Kenya. Sudan: There are 13 substantiated localities and many specimens from the Imatong Mts, east of the White Nile River, mostly between 41N and 51N (Tab. 1, Fig. 1). Three records from west of the White Nile (100 miles north of Diawo, Bahr-el-Ghazal Province; Ibba River, 100 miles north of Diawo; and Sue River, near Diawo) cannot be located precisely but are thought to be about 06.001N, 29.001E (see Fig. 1). All records in Sudan (both east and west of the White Nile river) are fairly close to the border with Uganda and Zaire (Tab. 1, Fig. 1), but none are as far north (ca. 101N) as indicated in Duthie and Robinson (1990). Ticks have been collected from specimens of P. marjorita at Magwe (04.081N, 32.171E) near Torit (Hoogstraal 1956). Zaire: There are several substantiated records and specimens from Zaire (now Democratic Republic of Congo) where there is woodland savanna similar to that in Uganda and Central African Republic. These records are clustered in the extreme northeast of the country, some of them within Garamba 381 National Park, and close to the border with Sudan (see Hatt 1940; Verheyen and Verschuren 1966). Angola and southern Zaire: The range in Angola appears to be based on information in Petter (1972) where it was recorded that the species occurred, inter alia, at ‘‘Gabela, Angola’’. Gabela (11.001S, 14.241E) is south of the Zaire River and its tributaries, and far removed from all other known localities. It is situated on the escarpment which contains a series of humid forests (running in a north– south direction) between the coastal plains and the highlands (J. Crawford-Cabral, in litt. 2004). Extensive collections at Gabela and other locations in Angola were made by Gerd Heinrich in 1954 and 1955, and are now housed in the Field Museum of Natural History in Chicago, USA The collection includes several Lepus specimens, but none of P. marjorita (W. T. Stanley, in litt. 2004). Francis Petter (in litt. 2004) is uncertain how Gabela came to be cited as a locality for P. marjorita (as given in Petter 1972), and João Crawford-Cabral (in litt. 2004) has doubts regarding the validity of P. marjorita in Angola. However, because Gabela is situated in a rather unique and special environment, there may be an interesting lagomorph, as yet undescribed, on the escarpments of Angola (João Crawford-Cabral, in litt. 2004). The single ‘‘record’’ for Angola was plotted by Kingdon (1974), and surrounded by a range (indicated by stippling) which included a large area south of Gabela and northwards into southern Zaire, south of the Zaire River. Duthie and Robinson (1990) showed a similar range, but without showing Gabela as a locality. Because there appears to be no specimen or reliable record from Gabela, yet alone any range in Angola and southern Zaire, we discount all of the range south of the Zaire River. Mozambique and Namibia: In the American Museum of Natural History, New York, there are specimens labelled as P. marjorita from ‘Manika, Mozambique’ (18.581S, 32.591E), ‘Sofala, Mozambique’ (9.301S, 32.591E) and ‘Okamahoro, near Okohandja, Namibia’ (Okahandja: 22.001S, 16.591E), all collected by R. G. Van Gelder (D. Lunde, in litt. 2003). These specimens have not been ARTICLE IN PRESS 382 D.C.D. Happold, W. Wendelen examined and hence we are uncertain of their taxonomic placement. These records were not plotted by Kingdon (1974, 1997) nor by Duthie and Robinson (1990). On biogeographical grounds, they are highly unlikely to be P. marjorita and hence are discounted here. The geographic distribution of P. marjorita has been shown to be much less extensive than previously recorded. The species is restricted to Uganda, extreme southern Sudan, northeastern Zaire, and eastern Central African Republic (although the range may extend to central Central African Republic). The overall distribution appears to have four separate clusters (Fig. 1) and without confirmed records between them; however, further careful investigation and additional specimens may reveal that the species occurs in the intervening country between these clusters and may be slightly more widespread than the present records suggest. There are no confirmed records or specimens to indicate that P. marjorita occurs in Angola, southern Zaire, northwestern Kenya, Rwanda and Burundi as previously reported. All records are in a region of rainforest-savanna mosaic north of the Equator between about 11N and 91N, and between about 201E and 331E. The current status of the species is uncertain; although considered as common in certain parts of Uganda in the 1920s to the 1960s, much of its range is in areas which have suffered civil war and a breakdown of law and order in recent years. Acknowledgments We acknowledge with thanks the following curators who have sent information on specimens in their museums: Mike Carleton (United States National Museum, Washington, DC), Judy Chupasko (Museum of Comparative Zoology, Harvard), Jacques Cuisin (Museum National d’Histoire Naturelle, Paris), Fritz Dieterlen (Staatliches Museum für Naturkunde, Stuttgart, Germany), Darrin Lunde (American Museum of Natural History, New York), Suzanne McClaren (Carnegie Natural History Museum, Pittsburgh), and Bill Stanley (Field Museum of Natural History, Chicago). Anke Hoffman (Museum für Naturkunde, Berlin, Germany) and Dieter Kock (Senckenberg Museum, Frankfurt, Germany) kindly confirmed that their museums did not have any specimens of P. marjorita. DCDH is grateful to The Natural History Museum, London, and to the Musée Royal d’Afrique Centrale, Tervuren, for permission to examine specimens in their care. We are also grateful to João Crawford-Cabral, Jonathan Kingdon, Robert Kityo, Dieter Kock and Francis Petter for their comments and advice, and to Alain Reygel (Musée Royal d’Afrique Centrale) for drawing the map. References Angermann, R. (1966): Beiträge zur Kenntnis der Gattung Lepus, I: Abgrenzung der Gattung Lepus. Mitt. Mus. Naturkunde, Berlin 42, 45–63. Corbet, G. B. (1983): A review of the classification in the family Leporidae. Acta Zool. Fenn. 174, 11–15. Duthie, A. G.; Robinson, T. J. (1990). The African Rabbits. In: Rabbits, Hares and Pikas. Status Survey and Conservation Action Plan. Ed. by J. A. Chapman and J. E. C. Flux. Gland, Switzerland: International Union for the Conservation of Nature, Gland, Switzerland. Pp. 121–127. Ellerman, J. R.; Morrison-Scott, T. C. S. (1951): Checklist of Palearctic and Indian Mammals 1758 to 1946. London: British Museum (Natural History). Hatt, R. T. (1940): Lagomorpha and Rodentia other than Sciuridae, Anomaluridae and Idiuridae collected by the American Museum Congo Expedition. Bull. Am. Mus. Nat. Hist. 76, 457–604. Hoffman, R. S. (1993): Order Lagomorpha. In: Mammal Species of the World: a Taxonomic and Geographic Reference. 2 edn. Ed. by D. E. Wilson and D. M. Reeder. Washington DC: Smithsonian Institution Press. Pp. 807–827. Hoffman, R. S.; Smith, A. T. (2005): Order Lagomorpha. In: Mammal Species of the World: a Taxonomic and Geographic Reference. 3 edn. Ed. by D. E. Wilson and D. M. ARTICLE IN PRESS The distribution of Poelagus marjorita Reeder. Baltimore: John Hopkins University Press. Pp. 185–211. Hoogstraal, H. (1956): African Ioxidea. Ticks of the Sudan (with special reference to Equatoria Province and with preliminary reviews of the genera Boophilus, Margaropus and Hyalomma). Washington, DC: US Navy. Kingdon, J. (1974): East African Mammals. An Atlas of Evolution in Africa. Vol. IIB. Hares and Rodents. London: Academic Press Pp. 343–704. Kingdon, J. (1997): The Kingdon Guide to African Mammals. London: Academic Press. Krampitz, H. E. (1968): Felduntersuchungen zur Verbreitung der Flohfauna in einer Kulturlandschaft Ost-Ugandas mit besonderer Beachtung des Menschen. Z. Tropenmed. Parasit. 19, 297–305. Malbrant, R. (1952): Faune du Centre africain franc- ais (Mammifères et Oiseaux). Encylopédie Biologique XV. Paris: Lechevalier. Montfort, A. (1992): Première liste commentée des mammifères du Rwanda. Rev. Zool. Africaine 106, 141–151. Petter, F. (1959a): Éléments d’une révision des lièvres Africains du sous-genre Lepus. Mammalia 23, 41–46. Petter, F. (1959b): Capture d’un Poelagus marjorita en Oubangui. Mammalia 23, 123–124. Petter, F. (1972): Part 5. Lagomorpha. In: The Mammals of Africa an identification manual. Ed. by J. Meester, H. W. Setzer. Washington, DC: Smithsonian Institution Press Pp. 1-7. Schouteden, H. (1948): Faune du Congo Belge et du Ruanda-Urundi. I. Mammifères. Annals 383 Musée Congo Belge, Tervuren, ser 81. Zoologie 1, 1–331. Setzer, H. W. (1956): Mammals of the AngloEgyptian Sudan. Proc. US Nat. Mus. 106, 447–587. St. Leger, J. (1929): An interesting collection of mammals, with a remarkable new species of Hare from Uganda. Ann. Mag. Nat. Hist. Ser. 10 (4), 290–294. St. Leger, J. (1932): A new genus for the Uganda Hare (Lepus marjorita). Proc. Zool. Soc. London 1932, 119–123. St. Leger, J. (1935): A new race of Poëlagus from the Bahr-el-Ghazal. Ann. Mag. Nat. Hist. Ser 10 (16), 598–600. Verheyen, W., Verschuren, J. (1966): Rongeurs et Lagomorphes. Exploration du Parc National de la Garamba. Fasc. 50. Institut des Parcs Nationaux du Congo. Williams, J. G. (1967): A Field Guide to the National Parks of East Africa. London: Collins. Wilson, S. E. (1995): Bird and Mammal Checklists for Ten National Parks in Uganda. Kampala: Uganda National Parks. Authors’ addresses: D. C. D. Happold, School of Botany and Zoology, Australian National University, Canberra, A. C. T., 0200, Australia (e-mail: David.Happold@anu.edu.au) Wim Wendelen, Musée Royale d’Afrique Centrale, Leuvensesteenweg 13, B-3080 Tervuren, Belgium