Primary Results of a Bat Survey in South-Western Ethiopia, with a New Ethiopian Record of Kerivoula Lanosa (Chiroptera: Vespertilionidae)

Russian J. Theriol. 7(2): 71–76 © RUSSIAN JOURNAL OF THERIOLOGY, 2008

Primary results of a bat survey in south-western Ethiopia, with a new Ethiopian record of Kerivoula lanosa (Chiroptera: Vespertilionidae)

Sergei V. Kruskop & Leonid A. Lavrenchenko

ABSTRACT. A short-term bat survey was conducted in spring, 2007, by the Joint Ethiopian-Russian Biological Expedition in the Mejangra Zone of the Gambela Peoples’ National Regional State, south- western Ethiopia. The area studied was adjacent to the Godare forest massif, one of the most lowland forests of Ethiopia. About 100 individuals of fifteen bat species from ten genera and six families were captured. Together with species recorded in 2000, the total bat community in Godare forest includes at least 19 species. One of the most important records is the third capture of Kerivoula lanosa from Ethiopia; the two previous captures in Ethiopia were more than a hundred years ago. The capture of Myotis bocagei also was only the third record from the country. Individuals of Rhinolophus and Miniopterus belong to undetermined species.

KEY WORDS: Chiroptera, Kerivoula, Miniopterus, Rhinolophus, Myotis, Ethiopia, new records.

Sergei V. Kruskop [[email protected]], Zoological Museum of Moscow State University, Ul. Bolshaya Ni- kitskaya 6, Moscow 125009, Russia; Leonid A. Lavrenchenko [[email protected]], A.N. Severtsov Institute of Ecology and Evolution RAS, Leninskii pr. 33, Moscow 119071, Russia. Ïðåäâàðèòåëüíûå ðåçóëüòàòû èçó÷åíèÿ ðóêîêðûëûõ íà þãî-çàïàäå Ýôèîïèè è íîâàÿ ýôèîïñêàÿ íàõîäêà Kerivoula lanosa (Chiroptera: Vespertilionidae)

Ñ.Â. Êðóñêîï, Ë.À. Ëàâðåí÷åíêî

ÐÅÇÞÌÅ. Íåïðîäîëæèòåëüíûå èññëåäîâàíèÿ ôàóíû ðóêîêðûëûõ â çîíå Ìåäæåíãð Äåìîêðàòè÷åñ- êîé Íàöèîíàëüíîé îáëàñòè Ãàìáåëà (þãî-çàïàä Ýôèîïèè) áûëè ïðîâåäåíû âåñíîé 2007 ã. â ðàìêàõ ðàáîòû ñîâìåñòíîé Ðîññèéñêî-Ýôèîïñêîé áèîëîãè÷åñêîé ýêñïåäèöèè. Îáñëåäîâàííàÿ òåððèòîðèÿ íàõîäèòñÿ íà ãðàíèöå ëåñíîãî ìàññèâà Ãîäàðå, îäíîãî èç íàèáîëåå íèçèííûõ ëåñîâ â Ýôèîïèè (âûñîòà íàä óðîâíåì ìîðÿ îêîëî 1300 ì). Áûëî îòëîâëåíî îêîëî ñòà îñîáåé ïÿòíàäöàòè âèäîâ ðóêîêðûëûõ èç äåñÿòè ðîäîâ øåñòè ñåìåéñòâ. Âêëþ÷àÿ âèäû, îòìå÷åííûå â 2000 ã., â öåëîì ñîîáùåñòâî ðóêîêðûëûõ ëåñà Ãîäàðå âêëþ÷àåò íå ìåíåå 19 âèäîâ. Îäíà èç íàèáîëåå çíà÷èìûõ íàõîäîê òðåòèé ñëó÷àé ïîèìêè íà òåððèòîðèè Ýôèîïèè óêðàøåííîãî ãëàäêîíîñà, Kerivoula lanosa; äâå ïðåäûäóùèå íàõîäêè áûëè ñäåëàíû áîëåå ñòà ëåò íàçàä. Ïîèìêà Myotis bocagei òàê æå ïðåäñòàâëÿåò ñîáîé âñåãî ëèøü òðåòüþ íàõîäêó âèäà íà òåððèòîðèè ñòðàíû. Ïîéìàííûå îñîáè ðîäîâ Rhinolophus è Miniopterus ïðèíàäëåæàò ê ïîêà íå óñòàíîâëåííûì âèäàì.

ÊËÞ×ÅÂÛÅ ÑËÎÂÀ. Ðóêîêðûëûå, Kerivoula, Miniopterus, Rhinolophus, Myotis, Ýôèîïèÿ, íîâûå íàõîäêè.

Introduction highland forests to the north and east and with savan- nahs in west. Bats of this forested area were very briefly Even brief surveys of local bat faunas provide infor- surveyed for the first time in 2000 (Lavrenchenko et al., mation about bat distribution, abundance and environ- 2004), when nine species were recorded. However, mental preferences, forming the basis for further con- results of that survey suggested a more diverse bat servation activities (Fahr & Ebigbo, 2003; Decher & community. Fahr, 2007). They are especially important in the case In spring 2007, a small mammal survey was con- of poorly investigated areas with specific natural condi- ducted by the Ethiopian-Russian Joint Biological Ex- tions. The primary forests of south-western Ethiopia pedition in the vicinity of Bishan Wak’a Lake (Meti undeniably represent such an area, differing from all area of the Mejangra zone, Gambela; N 7º17’, E 35º16’; neighboring territories and the rest of Ethiopia. altitude 1350–1390 m ASL), on the border of Godare The Godare forest massif is in the most eastern part forest massif. About 100 individuals of bats were cap- of Gambela Province, bordering the remains of more tured, representing at least fifteen species from 10 gen- 72 S.V. Kruskop & L.A. Lavrenchenko era and six families. Amongst them, an adult female of For all but three voucher specimens DNA barcodes the woolly bat (Kerivoula) was captured, only the third (sequences of the 5’-fragment of cox-1 mitochondrial record of the genus for Ethiopia and the first from the gene, up to 658 bp) were made as part of the Barcoding south-west of the country. of life project [http://barcodinglife.com], forming basis While there is some progress in the understanding for further molecular studies. A standard cocktail of of Asian Kerivoula (Francis et al., 2007), the African primers acceptable for various mammalian species branch of the genus remains poorly investigated. Infre- (Ivanova et al., 2007) was used to get PCR-products of quent captures and rarity of museum specimens inhibit the 5’-fragment of cox-1 (cytochrome c oxidase second both the investigation of taxonomic relationships be- subunit) mitochondrial gene (up to 658 bp). Analogous tween named forms and ecological research, obscuring data from Kenyan bats were used for comparison. The environmental preferences of this or that species. Kimura 2 parameters model of the neighbour-joining method was accepted for grouping data. More detailed Material and methods analysis of Ethiopian and Kenyan bat DNA barcodes is ongoing (Borisenko, in prep.). The territory on which the investigations were conducted is situated about 23 km W from Tepi at relative- Results ly low elevations (1350–1390 m ASL). It was formerly part of an evergreen tropical forest massif known as Thirteen species of insectivorous bats, from five Godare Forest. Now most of the study area is covered different families, and two species of fruit bats were with coffee plantations (Tepi coffee farm) usually with found. Previously published records from Godare for- canopy trees retained to provide shadow over the cof- est (Lavrenchenko et al., 2004) include two more insec- fee crop. These plantations are divided by variable tivorous bats and two fruit bats, making the overall bat patches of pastures and primary and secondary growth. diversity comparable to that of West African tropical The largest water source in the area is Bishan Wak’a forests (e.g., Fahr & Ebigbo, 2003). Lake — an almost round, flowing lake about 1.5 km in diameter, surrounded by a circle of primary forest. Kerivoula lanosa All the bat research was restricted to that locality, from 31 March to 7 May. Bats were observed during The new record of the wooly bat should be the most the evening, usually between sunset and 21.30–22.00, important result of the given survey. A single adult several times also after midnight or in the morning female of K. lanosa was captured by the flap-trap on 30 hours up to sunrise. All the available biotopes were April. Only two localities of woolly bats were previous- sampled, including variably disturbed primary forest, ly known from Ethiopia: one in northern Ethiopia in the secondary growth, riverine biotopes, road, growth edg- Simien mountains, and the second from the Great Rift es and coffee plantations. Observations were conducted Valley somewhere north of Lake Abaya (Largen et al., visually with the help of electric lanterns, and acousti- 1974; Yalden et al., 1996). Both records were made cally by the use of narrow-band heterodyne ultrasound more than hundred years ago, and each was described detector D100 (Pettersson Elektronik AB, Sweden), as a distinct species, K. eriophora (Heuglin, 1877) and and accompanied with the capture of selected individu- K. harrisoni Thomas, 1901, respectively. Our record als using the flap-trap (Borisenko, 1999). Some indi- was made at least 320 km west from the more southern viduals were captured with the use of typical nylon previously published locality of the genus in Ethiopia mist-nets (size 7 x 2 m), set in bat foraging sites. Total (Fig. 1). trapping effort of this type of trap was relatively small, According to the latest checklist (Simmons, 2005) amounting to only 43 hours/net. there are presumed to be seven African species of All the captured individuals were examined to deter- Kerivoula. Two of them, K. phalaena and K. cuprosa, mine sex, age and physiological condition. Selected spec- were not documented for East Africa. Two species imens of all species captured were collected for further occur in Ethiopia. One of them, K. eriophora, is known investigation, including of verification of their taxonom- only from type locality in northern Ethiopia and once ic position and status. Voucher specimens were fixed in was supposed to be a senior synonym for K. africana alcohol (70% ethanol) or 4% formaldehyde, or prepared (Hayman & Hill, 1971); in his turn Kock (1984) con- as a dry skin and skeleton. Extra bat individuals were cluded that K. eriophora cannot be conspecific with released after the external examination in the same place K. africana and belongs to “lanosa” species group. in which they were captured. Voucher specimens were Based on the description and measurements of the taken for the investigation to the Zoological Museum of only known K. eriophora specimen, Yalden et al. Moscow State University. Acronyms of depositories of (1996) suggest that this form is conspecific with K. the processed collections are as follows: ZMMU — Zoolog- lanosa and a senior name for K. l. harrisoni. Never- ical Museum of Moscow State University; ROM — theless, the northern Ethiopian specimen is still usual- Royal Ontario Museum, Toronto; MHNG — Natural ly treated as uncertain. K. harrisoni is now usually History Museum of Geneva; ZIN — Zoological Institute thought to be a subspecies of K. lanosa (Koopman, of Russian Academy of Sciences, St. Petersburg. 1994; Simmons, 2005). Bats from South-West Ethiopia 73

mens and of the K. eriophora co-type (Yalden et al., 1996). We did not see any specimens of K. africana, but according to the literature this species is smaller and lacks the fringe on the tail membrane (Koopman, 1994; Yalden et al., 1996). Thus we can assign the Godare woolly bat to K. lanosa, and it is to some extent intermediate between K. l. harrisoni and K. l. muscilla; its allocation to a definite geographic form needs investigation of additional material. Our specimen was captured on the border between the coffee plantation and a small triangular patch of primary growth. At least one animal of the same species was observed in the same place two days later. In both cases animals were observed about one and half hours after sunset, flying slowly but extremely maneuverable over the coffee trees and under the large tree growing nearby, usually keeping very close (about 5–30 centi- meters) to tree trunks and to canopies. The tendency to fly very close to obstacles or even between them makes Figure 1. Localities of Kerivoula records in Ethiopia: 1 — woolly bats very hard to capture with the flap trap, the type locality of K. eriophora; 2 — type locality of K. harriso- same habit gives them the ability to escape mist nets. ni; 3 — locality of our field work in Godare forest. This is one of the reasons why Kerivoula are so rare in scientific collections. African woolly bats and particu- The following specimens were used as comparative larly K. lanosa are sometimes thought to be more char- material: K. lanosa muscilla — male and female from acteristic of savannah woodlands (Taylor, 2000); how- Cote d’Ivoire (types of K. l. bellula, MNHG-965.038, ever according to Grubb et al. (1998) and Fahr & 965.039), female from Nigeria (ROM-86938); K. l. Ebigbo (2003), in West Africa this species seems to be harrisoni — male from Kenya (ROM-48531); K. argen- more typical of forests. Our specimen came from rela- tata — male from Kenya (ROM-36421); K. smithi — tively humid forest. male from Kenya (ROM-48495); K. phalaena — male Observed specimens emitted very low intensity, fre- from Rwanda (ROM-59191). quency-modulated, echolocation calls, detected at about Selected measurements of our specimen from Go- 100 kHz and over, which is similar to characteristics dare (ZMMU S-181514) are as follows: forearm: 32.7 known for Asian woolly bats (Kingston et al., 1999) mm, head and body length: 41 mm, tail length: 35.5 and corresponds to the ecological niche of a leaf-glean- mm, hind foot: 6.4 mm, condylocanine length: 11.77 er or species foraging in highly cluttered space (Norb- mm, mastoid width: 7.17 mm, upper tooth row: 5.30 erg, 1994). mm, width between upper canines: 3.16 mm, lower tooth row: 5.68 mm. Externally this individual looks Other species very similar to the specimen of K. l. harrisoni from Kenya, having relatively dark, brown dorsal fur frosted In the direct vicinity of Bishan Wak’a, fruit bats with long very pale-silver hair tips, dark brown ventral were represented by Micropteropus pusillus and Rou- under fur and a well developed fringe of hair on the settus aegyptiacus. Both species seem to be common uropatagium. However our specimen has some minor and abundant, being captured and observed many times difference. Its dorsal fur has shorter silver tips to the during the field work. One specimen of Epomophorus guard hairs; it also looks paler than in K. l. harrisoni sp. was observed but not captured; probably it was E. because of presence of wide brown bands on the hairs. gambianus, which was previously recorded for Godare There is a visible difference in pigmentation of the ears: forest (Lavrenchenko et al., 2004). The list of insectiv- they well pigmented in the ROM specimen and color- orous bats includes, besides Kerivoula, Nycteris hispi- less in our individual. West African specimens of K. da, Rhinolophus sp., Triaenops persicus, Miniopterus lanosa seem to be somewhat smaller and paler than sp., Myotis scotti, M. bocagei, M. welwitschii, Pipist- both Godare and Kenyan specimens. The specimen of rellus cf. aero, P. rusticus, Neoromicia guineensis, N. K. smithi is similar in size but possesses almost no fur cf. nanus and Scotophilus dinganii. Undetermined free- or fringe on tail membrane; its coloration on both back tailed bats from Molossidae family were detected by and belly looks darker because of distinctly shorter and their echolocation calls. Micropteropus pusillus, Tri- more yellowish hair tips. K. argentata has larger pro- aenops persicus, Neoromicia cf. nanus, N. guineensis, portions and is distinctly paler in color, almost white and Scotophylus dinganii were the most common and ventrally. Cranial proportions of the new specimen cor- abundant amongst observed bats. respond well with all measured skulls of K. lanosa as Most of the above listed species are already known well as with published measurements of Kenyan speci- for Gambela Province (Largen et al., 1974) and even 74 S.V. Kruskop & L.A. Lavrenchenko

Figure 2. Kimura 2 parameter tree for East African Miniopterus and some Rhinolophus; fruit bats were taken as outgroup; bootstraps over 70% calculated from 1000 replicas are mentioned over appropriate branches. for Godare forest (Lavrenchenko et al., 2004). For the similar to that of Rh. alcyone from West Africa. Ac- Ethiopian endemic Myotis scotti our record is new; cording to the preliminary results of cox 1 gene analy- however, there is a previous known locality less than 30 sis, these two specimens lie at a distance from Kenyan km eastward, on the Beko River in the vicinity of Tepi ones close to species level (average Kimura distance (Lavrenchenko et al., 2004). Thus the record of major 4.77%, observed distance 4.57%; Fig. 2). faunistic interest is M. bocagei. This species, not rare in Three captured specimens of medium-size Min- the East Africa (Kingdon, 1974), has been previously iopterus were compared with other African bent-winged found in Ethiopia only three times, in Addis-Ababa bats in ZMMU and ROM collections. They demon- (Demeter & Topal, 1982), near Arba Minch and at strate similarity with M. minor in fur structure and Djala (Yalden et al., 1996). During our observations, coloration, but resemble M. natalensis in external and Bocage’s mouse-eared bat was common on and nearby cranial size. Average Kimura distances between newly Bishan Wak’a Lake. Possibly its habit of hunting over found cox I haplotypes and the clade comprising Min- water has led to it being under-recorded. iopterus natalensis, M. minor, and M. africanus from Both captured horseshoe bats demonstrate qualita- Kenya is 9.78% (observed distance 8.93%; Fig. 2). tive features, such as definitely hastate lancet, relatively narrow horseshoe not covering the whole muzzle, and Discussion lacking of the small upper premolars, which allocate them to Rh. clivosus (Csorba et al., 2003). However Summarizing results of this survey and previous these individuals compared with reference material in data, we can speak about the bat community containing the Royal Ontario Museum and Moscow Zoological at least 19 species. Insectivorous bats are represented, Museum, look unlike other Rh. clivosus from Ethiopia as it was already mentioned, by majority of known and Kenya, especially in coloration which is greatly ecological forms: gleaners, trawlers and aerial hawkers Bats from South-West Ethiopia 75

(e.g., Schnitzler & Kalko, 1998; Kruskop, 1999). Foli- and thus depend on anthropogenous transformation of age gleaners are represented by Kerivoula lanosa, the landscape. Retaining more or less untouched patch- ground hovering gleaners by Nycteris hispida. Poten- es, even small, of primary forest vegetation allowed the tially gleaning behavior could be expected in Myotis forest bat fauna to survive in the zone of coffee plant- welwitschii, however in our observations it seems to be ing. However we did not observed foraging bats – a highly maneuverable aerial hawker. Amongst other either forest or eurytopic species – in places of continu- members of this ecological group, Neoromicia cf. na- ous plantations with greatly destroyed forest cover. nus, N. guineensis and probably Rhinolophus sp. ex- As for outstanding taxonomic questions, the identi- ploit highly cluttered space between growths, while ties of Rhinolophus and Miniopterus remain problem- both true Pipistrellus, Myotis scotti and Triaenops per- atic. Five species of Miniopterus usually thought to sicus use edges of the growth, ecologically comparable occur in East Africa (Simmons, 2005), however it is to other typical pipistrelles, whiskered bats and large possible that there are still present some undescribed serotines respectively. One of the most interesting as- forms (Appleton et al., 2004). Number of horseshoe pects is the foraging behavior of N. cf. nanus, which bats known for East and North-East Africa is higher: exploits the periphery of canopies and bushes, often about twelve species (Simmons, 2005). Rhinolophus seen flying close to vegetation. Maximum energy of the clivosus itself represents a highly complex morpholog- echolocation call of this species was at over 70 kHz, ically variable taxon (Cotterill, 2002; Csorba et al., which does not correspond to published data on N. 2003) which actually may include more then one spe- nanus (Taylor, 2000). Scotophilus dingani, Miniopter- cies. In any case both horseshoe bats and bent-wing us natalensis and the undetermined free-tailed bat rep- bats are members of taxonomically problematic groups, resent typical high-speed hawkers, exploiting unclut- and identification of the Godare specimens need further tered space. Finally, Myotis bocagei demonstrate for- and more comprehensive investigation of comparative aging behavior almost identical to that of M. daubento- material and molecular data. nii and similar forms (Kalko & Schnitzler, 1989), of a typical trawler or water gleaner. Thus, in comparison to ACKNOWLEDGEMENTS. We would like to thank the other palaeotropical bat communities, we were missing Manager Office and staff of the Tepi Coffee Plantation for only two ecological forms, namely nectarivorous and the use of the Plantation buildings and equipment. We are carnivorous bats. In tropical Africa these types are especially indebted to our Project Coordinators Dr. Andrei represented by the fruit bat Megaloglossus woermanni; Darkov (Joint Ethiopian-Russian Biological Expedition, Sec- ond Phase - JERBE II) and Ato Girma Yosef (Ethiopian by giant slit-faced bat Nycteris grandis and probably Science and Technology Commission) for management of Cardioderma cor (Norberg & Fenton, 1988) Taking the expedition in the field and in Addis Ababa. The access to into account the overall richness of this community, the collections of the Royal Ontario Museum was kindly these elements could probably be found during further provided to the first author by Dr. Judith Eger. We are research. sincerely grateful to Dr. Alex Borisenko, Dr. Nataly Ivanova In respect of environment type, revealed fauna main- and director of Canadian Centre for DNA Barcoding (Guel- ly consist of species with variable biotopical preferenc- ph, Canada) Dr. Paul Hebert for preliminary molecular anal- es, living both in forests and open areas. According to ysis of Godare animals. We also would like to express our Grubb et al. (1998) Micropteropus pusillus and Neoro- thanks to Dr. Ivan Kuzmin (Centers for Disease Control and micia guineensis are mainly inhabitants of savannah Prevention, Atlanta, USA) who provided his Kenyan material for comparison, and Dr. Vladimir Lebedev for his help in woodlands (and previously both species were known in calculating molecular distances and building dendrograms Ethiopia from such habitats); while Nycteris hispida for Miniopterus and Rhinolophus. thought to be mainly forest species. 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