Article in press - uncorrected proof Mammalia 74 (2010): 163–176
2010 by Walter de Gruyter • Berlin • New York. DOI 10.1515/MAMM.2010.026

Small mammal survey in the upper Seli River valley, Sierra Leone

Jan Decher1,*, Ryan W. Norris1,2 and Jakob Fahr3 1

Department of Biology, University of Vermont, Burlington, VT 05405-0086, USA, e-mail: [email protected] 2 Department of Biology, 208 Mueller Laboratory, Pennsylvania State University, University Park, PA 16802, USA, e-mail: [email protected] 3 Institute of Experimental Ecology (Bio 3), Ulm University, Albert-Einstein Allee 11 D-89069 Ulm, Germany, e-mail: [email protected] *Corresponding author

Abstract In 2006, an inventory of small mammals was conducted on the upper Seli River in central Sierra Leone on the southwestern fringe of the Loma-Man portion of the Guinea Highlands as part of the baseline surveys conducted for the Bumbuna Hydroelectric Project (BHP). We recorded a total of 35 mammal species, including three shrews (Soricomorpha), 19 bats (Chiroptera), 11 rodents (Rodentia), one primate, and one carnivore. New or first well-documented species for Sierra Leone are the shrew Crocidura nigeriae, the bats Myonycteris torquata, Rhinolophus denti knorri, and the rare horseshoe bat Rhinolophus ziama endemic to Upper Guinea. Also reported are two bat specimens apparently representing an undescribed species of Neoromicia related to N. nana. We support the creation of a protected area as an offset for areas lost to the inundation of the BHP reservoir. Keywords: Bumbuna Hydroelectric Project; Chiroptera; Rhinolophus ziama; Rodentia; Soricomorpha; West Africa.

Introduction During the 11 years of civil war in Sierra Leone (1991–2002), most research on small mammals within the country came to a standstill. Universities closed and field stations as well as conservation sites, such as Tiwai Island, were destroyed or abandoned. Glew and Hudson (2007) suggested that the lawlessness and human displacement caused by armed conflict leads to accelerated deterioration of the environment. The impact of conflict on biodiversity hotspots, including the Guinean Forest Hotspot, which includes parts of Sierra Leone, has recently been reviewed by Hanson et al. (2009). The specific case of the effects of the war on

Tiwai Island in Sierra Leone has been described in some detail by Oates (1999). The most recent and thorough review of Sierra Leone mammals is that of Grubb et al. (1999), which is based on pre-civil war publications and museum data. For small mammals these authors list 11 species of ‘‘insectivores’’ (one hedgehog, 10 shrews), 55 species of bats (Chiroptera), 40 rodent species (Rodentia), and one hare (Lagomorpha) for the country. For the four 0.258-squares that encompass our project area to the north of the town of Bumbuna, they mapped two shrew, seven bat, and 12 rodent species, but provided no specific localities. Topography of Sierra Leone consists of coastal and interior lowlands and a high plateau in the northeastern half of the country with 300–600 m elevation dotted by several mountain ranges, with the highest peak being Bintumani Mountain in the Loma range at 1945 m. Ten major rivers drain the plateau region, one of which is the Seli (formerly Rokel) River. The upper Seli River valley, stretching 30 km straight north above Bumbuna Falls before turning 90 degrees east, forms the study area of this survey along with tributaries including the Mawoloko River (Figure 1). This area is characterized by a heterogeneous forest-savanna mosaic, with patches of moist semi-deciduous forest mostly restricted to steep ravines and gallery forest along the water courses. Annual rainfall around Bumbuna (98049N, 118449W) is estimated as 2771 mm with a single wet season from May to November, and a mean monthly temperature of 25.78C (WorldClim 2.59 grid, Hijmans et al. 2005). There are several small traditional villages in the area with rice cultivation in wet depressions and harvesting of non-timber forest products such as oil palm nuts. In this paper, we present results from the first post-war small mammal survey conducted in Sierra Leone. The survey was carried out in the context of the Bumbuna Hydroelectric Project (BHP), which was much delayed by the civil war, but as of November 2009 has been generating electricity for the capital Freetown with the towns of Makeni and Bumbuna to be supplied later. Our study provides baseline data to potentially assess the ecological effects of the Bumbuna reservoir on small mammals.

Material and methods Our study area is located in the Northern Province of Sierra Leone 3 km north of the town of Bumbuna, between the Sula Mountains to the east and the Katabai Hills to the west along a 30-km north-south segment of the upper Seli River

2010/2010011

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(Figure 1). The area is part of the ‘‘Guinea Highlands’’ identified to be of ‘‘extremely high conservation priority’’ for mammals in West Africa (Bakarr et al. 2001). Field work was conducted during 2–28 January 2006 as part of a baseline biodiversity survey commissioned by the BHP. The project involved the sampling of small terrestrial mammals and bats in different vegetation types of the forest-savanna mosaic in the projected reservoir catchment along the upper Seli River. This included riparian forest, ravine forest, grassland away from river, a small rice swamp, and farmbush/secondary forest within a total area of 52.5 km2. Outside of the riparian zone the area is dominated by humid savanna, secondary forest and farmbush, or managed forest remnants, including oil palm stands and sacred groves near villages. At each site terrestrial small mammals were sampled for two or three consecutive nights with two traplines of 50 Sherman live traps (Model LFATDG, 7.5=9=23 cm), 12 Museum Special, five Victor snap traps, and two large Tomahawk live traps. Trap stations were placed 10–15 m apart with two traps per station. Except where rocky conditions prevented it, each trapline also included 5–6 pitfall bucket traps connected by 25 m of plastic drift fencing. The four Tomahawk live traps were used more opportunistically with the assistance of local trapping experts to target larger species such as larger rodents or small carnivores. We recorded species, body mass, sex, and reproductive condition for released animals. To characterize microhabitat at each capture site, percent canopy cover was measured with a concave

spherical densiometer (Robert E. Lemmon, Bartlesville, OK, USA). Percentage of ground cover types (herbs, grass, leaf litter, woody, rock, bare soil) were estimated for a one square meter area centered on each successful trap site and averaged by species. Bats were captured with 2.6=12 m and 2.6=6 m polyester mist nets (19 mm mesh, 75 denier, four shelves; www. avinet.com) set from 18:00 h until 22:00 or 23:00 h along presumed flyways, over watercourses, and along forest edges. Nets were set at ground level with the exception of one 12-m net that was raised up to 1.5 m above ground with a custom-built pulley system. Number of nets ranged from two to eight nets per site. Netting data, weather conditions, and standard measurements (body mass, forearm, head and body, tail, ear, and hindfoot) were recorded. Locations were fixed with a Garmin 12 GPS receiver. Sample site descriptions and representative coordinates are given in Table 1. Species accumulation curves were generated with the program EstimateS, Version 8.0 (Colwell 2006). The samplebased rarefaction curve was calculated with the ‘‘Mao Tau’’ function (Colwell et al. 2004) and graphs rescaled by individuals to allow for comparison of species richness with other studies. The IUCN Red List status was taken from the most recent update (IUCN 2008). Species were identified in the field based mostly on Happold (1987), Hutterer and Happold (1983), and Meester and Setzer (1971). Identifications were subsequently checked with representative voucher specimens. Bat and shrew vouchers are housed at the Zoolo-

Figure 1 Location of the study region in Sierra Leone and sampling sites within the study region (for site numbers refer to Table 1).

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Table 1 Description of sampling sites, location relative to inundation level, and representative GPS coordinates. Site no.

Site name

Description

Inundation zone

Representative GPS coordinates

1

BHP dam

Inside

9804926.60N, 11843918.10W

2

Tall grass hillside

Outside

9805947.10N, 11843949.60W

3

Forested ravine

Inside

9805934.90N, 11843924.60W

4

Kafogo

Inside

9823946.00N, 11843947.10W

5

Kakutan river crossing

Inside

9809941.10N, 11842933.90W

6

BHP camp ravine BHP camp water source

Riverine forest and forested ravines on left bank of Seli River approximately 500 m north of BHP dam Tall grass hillside and Chromolaena thickets above inundation zone west of BHP dynamite building Forest patch with bamboo and Cola trees along creek in steep ravine below BHP quarry Near Kafogo, Mawoloko river (NW tributary to Seli); riverine forest and edge of rice swamp Near Kakutan, Seli river crossing to Masumarandugu village; floodplain, forested ravine with creek, bordered by farmbush and Chromolaena thicket Steep ravine with rocky creek bed, southwest of Bumbuna camp Higher elevation forest patch at the BHP drinking water source northwest of the BHP camp (bats only)

Outside

9803953.90N, 11843951.60W 9804930.90N, 11844900.90W

7

gisches Forschungsmuseum Alexander Koenig, Bonn (ZFMK) and rodent vouchers at the US National Museum, Washington, DC, USA (USNM). DNA tissue samples and some ectoparasites are stored at the University of Vermont Zadock Thompson Mammal Collection (ZTNH). Previous and partly unpublished records of small mammals from Sierra Leone were queried through MaNIS -http://manisnet.org), with large collections held by the USNM and the Royal Ontario Museum, Toronto, Canada (ROM). These data together with previously published records were used to assess species distributions within the country. Techniques employed for capture and preservation complied with standard methods for measuring and monitoring mammal diversity (Voss and Emmons 1996, Wilson et al. 1996) and guidelines approved by the American Society of Mammalogists (Animal Care and Use Committee 1998). Taxonomy follows Wilson and Reeder (2005) unless otherwise noted.

Results We captured a total of 270 individuals belonging to 31 species: three shrew (Soricomorpha), 18 bat (Chiroptera), and 10 rodent (Rodentia) species. Four additional species were observed (Table 2). The species accumulation curves (Figure 2) for bats and terrestrial small mammals rise steeply and do not reach an asymptotic plateau. This indicates that additional species can be expected in the BHP area with additional sampling effort. Kafogo and Kakutan were the most species rich sites with 16 and 15 total small mammal species, respectively. In the following section, we present an annotated list of the species recorded during this survey, with comments on

Outside

their systematics, known distribution in Sierra Leone, and their conservation status. Order Soricomorpha Family Soricidae Crocidura jouvenetae Heim de Balsac, 1958 Jouvenet’s Shrew

Two males (ZFMK 2006.119, -.120) were captured within or very close to the inundation zone at the northernmost sampling location (Kafogo); one in upland dry forest, the other in elephant grass between the forest edge and a small rice field close to the Mawoloko River. Both sites had a relatively low canopy (17.5%) and considerable grass cover (30%). Previously, this species was included as a subspecies of C. crossei and reported for Sierra Leone from 10.1 km NE Fintonia in Outamba-Kilimi National Park (USNM 546947) and from the Gola region (Grubb et al. 1999). The type locality for this species is Zie´la, Mt Nimba, in Guinea (Heim de Balsac 1958). Crocidura nigeriae Dollman, 1915 Nigerian Shrew

One male (ZFMK 2006.121) was trapped below the inundation line at the Kakutan River crossing. Another male (ZFMK 2006.122) was captured in the ravine west of the BHP camp at the bottom of a steep slope with holes. Crocidura nigeriae has not previously been recorded from Sierra Leone, although it is possible that individuals were included with C. poensis in Grubb et al. (1999) as the taxon nigeriae was previously considered a subspecies of C. poensis (Heim de Balsac and Meester 1977, Hutterer and Happold 1983). Crocidura nigeriae is readily distinguished from C. poensis by its larger overall size. In Nigeria, C. nigeriae

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Table 2 Overview of all species and individuals captured or observed at seven sampling sites. Locality

Date Soricomorpha Crocidura jouvenetae Crocidura nigeriae Crocidura poensis Rodentia Xerus erythropus Cricetomys emini Hybomys planifrons Hylomyscus alleni Lemniscomys striatus Lophuromys sikapusi Malacomys edwardsi Mastomys erythroleucus Mus musculoides/minutoides Mus setulosus Praomys rostratus No. individuals Trap nights Capture success (%) Chiroptera Epomops buettikoferi Micropteropus pusillus Myonycteris torquata Nanonycteris veldkampii Rousettus aegyptiacus Rhinolophus wfumigatusx diversus Rhinolophus guineensis Rhinolophus denti knorri Rhinolophus ziama Hipposideros abae Hipposideros cyclops Hipposideros vittatus/gigas Hipposideros jonesi Hipposideros cf. ruber Nycteris arge Nycteris grandis Miniopterus villiersi Myotis bocagii Neoromicia aff. nana No. individuals 12 m net hours Bats/12 m net hour Carnivora Nandinia binotata Primates Perodicticus potto Total no. of species

1 BHP dam

2 Tall grass hillside

3 Forest ravine

4 Kafogo riverine forest

5 Kakutan ravine and riverine forest

6 BHP camp ravine

7 BHP camp water source

5–9 Jan

9–10 Jan

10–13 Jan

16–19 Jan

20–26 Jan

27–28 Jan

27–28 Jan

2 1

X 2 1 1 2

4 12

2

1 12 18 292 6.2

2

13 268 4.9

58 65 432 15.0

8 6 2 1 9 32 288 11.1

2 2 5 1 1

5

11 23 288 8.0

6 12 160 7.5 1

1 1

1 1

1

n/a n/a n/a

3 4 2 3 31 1

1

7 1 1 1 1 X 6 25 2 2 5 10 2 107 1105.1 0.10

1 1

1

1 1 1 1 1

1 2 1 36 334.9 0.11

5

2 1 1 1

X 2 10 1 1

1 3

2 4

3 8

7 38.8 0.18

7 172.4 0.04

1 9 224.6 0.04

24 256.5 0.09

7 58.4 0.12

17 19.5 0.87

X

10

6

7

16

15

X 10

X 5 4 15 1 24 8 2 1 10 87 163 1728 9.4

2 3 1

31

4 1

Red list

2 2 2

1

2

1

Total

VU EN

NT

n.a.

X

8

X 35

Xerus erythropus, Hipposideros vittatus/gigas, Nandinia binotata, and Perodicticus potto are sight records, hence only their presence is given. Red list status except for those species ranked as ‘‘Least Concern’’ by IUCN (2008): NTs‘‘Near Threatened’’, VUs‘‘Vulnerable’’, ENs‘‘Endangered’’, n.a.snot yet assessed. Trap nightssnumber of nights trapped=number of trap units. Net hours were calculated as 12 m-equivalents, including 6 m-nets (=0.5).

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Micropteropus pusillus (Peters, 1868) Peters’s Lesser Epauletted Fruit Bat

One female (ZFMK 2006.125) of this savanna species was captured in a net over the Mawoloko River at Kafogo. Two females and one male were recorded over the creek above the Bumbuna drinking water source. Micropteropus pusillus is known from at least nine localities in Sierra Leone (Bergmans 1989, Grubb et al. 1999; ROM, USNM), mostly from the northern half of the country. Myonycteris torquata (Dobson, 1878) Little Collared Fruit Bat Figure 2 Sample-based rarefaction curves (interpolated species accumulation: Sobs) for bats and terrestrial small mammals rescaled by individuals.

grouped ecologically among the ‘‘dark forest and swamp shrews of large size’’ but it has also been recorded there from ‘‘farmlands in the rainforest zone and grassfields’’ (Hutterer and Happold 1983: 51). Microhabitat data showed that C. nigeriae had the highest association with leaf litter (80%) and the second highest association with canopy cover (45%) of all terrestrial small mammal species caught in the survey.

Crocidura poensis (Fraser, 1843) Fraser’s Musk Shrew

Two individuals (ZFMK 2006.123, -.124) were recorded at Kafogo, one male in dense elephant grass in a rice field and a female in a drainage channel below a small densely herbaceous marshy area above the left bank of the Mawoloko River. Crocidura poensis is known in Sierra Leone from numerous localities in the coastal region. Ecologically C. poensis has been categorized among the ‘‘dark forest and swamp shrewwsx of large size’’ for Nigeria. It has also been recorded ‘‘in various types of grasslands and in some farmlands’’ (Hutterer and Happold 1983: 50), which is reflected in our microhabitat data showing low canopy cover (12.5%), high herbaceous growth (25%), and grass cover (50%). Crocidura poensis has previously been recorded from at least 17 localities throughout Sierra Leone (Grubb et al. 1999).

Order Chiroptera Family Pteropodidae Epomops buettikoferi (Matschie, 1899) Bu¨ttikofer’s Epauletted Fruit Bat

Three specimens were captured outside the inundation zone below the dam. One male (ZFMK 2006.126) was caught over the creek in the BHP camp ravine and two females over the creek at the BHP drinking water source. In Sierra Leone this species is known from at least 11 different localities throughout the country (Bergmans 1989, Grubb et al. 1999; ROM, USNM).

One male (ZFMK 2006.129) was recorded over a small stream in the forested ravine near the Bumbuna camp, and one lactating female was captured over the creek above the Bumbuna drinking water source. These individuals represent the first records for Sierra Leone with precise localities. The taxon leptodon Andersen, 1908, now considered a synonym of M. torquata, was described from Sierra Leone without a specific locality (Bergmans 1997). Nanonycteris veldkampii (Jentink, 1888) Veldkamp’s Dwarf Epauletted Fruit Bat

Three females were captured: one at the Bumbuna dynamite building above the inundation zone (ZFMK 2006.128), one in a net over the Mawoloko River at Kafogo within the inundation zone (ZFMK 2006.127), and one in the BHP camp ravine. Our captures are the second record of N. veldkampii for Sierra Leone, the first being from Bo (Bergmans 1989). Rousettus aegyptiacus (E. Geoffroy, 1810) Egyptian Rousette

This colonial fruit bat was found roosting in large numbers (several hundred) in the large intake tunnel on the northern (reservoir) side of the Bumbuna dam. We captured 31 individuals (three collected: ZFMK 2006.130-.132). Three females were carrying pups in early January, which corresponds fairly well with observations of R. aegyptiacus in a cave on Pic de Fon, Guinea, where parturition was observed in late November (N. Ebigbo and P. Naskrecki personal communication). This species has been previously recorded from the Freetown Peninsula as well as the hilly and mountainous region in the northeastern part of the country (Bergmans 1994, Grubb et al. 1999; ROM, USNM). Sufficiently large caves or cave-like roosting places seem to be essential for this species (Bergmans 1994). Family Rhinolophidae Rhinolophus wfumigatusx diversus Sanborn, 1939 Ru¨ppell’s Horseshoe Bat

One male (ZFMK 2006.136) was captured in a net set in ravine forest at Kakutan inside the inundation zone. This bat was previously known from three localities in the hilly and mountainous region of northeastern Sierra Leone (Rosevear 1965).

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The taxonomy of the fumigatus group and species limits of its members are largely unresolved and in need of revision. Rosevear (1965) already pointed out that the size range from fumigatus Ru¨ppell, 1842 to aethiops Peters, 1869 is very large, indicating that the complex includes more than one species. Both Csorba et al. (2003) and Turni and Kock (2008) emphasized the distinctly larger skull size of one of the syntypes of aethiops (loc. typ.: Otjimbingue, Namibia) in comparison to the lectotype of fumigatus (loc. typ.: Shoa, Ethiopia). The West African taxon foxi Thomas, 1913 (loc. typ.: Kabwir, central Nigeria) is very similar in measurements to fumigatus, whereas diversus Sanborn (1939) (loc. typ.: Bakel, Senegal) is morphometrically in agreement with aethiops. The presence of both large (diversus) and small (foxi) populations in West Africa suggests that this group consists of several independent lineages and further analyses might show that fumigatus, aethiops, foxi, and diversus are distinct species.

Highlands and listed as ‘‘Vulnerable’’ B2ab (iii) (IUCN 2008).

Rhinolophus guineensis Eisentraut, 1960 Guinean Horseshoe Bat

One female (ZFMK 2006.142) of this recently described species was captured outside the inundation zone over the creek exiting the forest above the Bumbuna drinking water source. Prior to our study, only four specimens were known from two localities in southeastern Guinea (Massif du Ziama) and northwestern Liberia (Wonegizi Mts; Fahr et al. 2002). Our record of R. ziama at Bumbuna is the first from Sierra Leone and extends the range of this bat by 270 km to the southwestern edge of the Guinea Highlands (Figure 3). The species is currently ranked as ‘‘Endangered’’ B1ab (iii) by the IUCN Red List (2008) owing to the very small distribution range, few locations, and an inferred decline in the extent and quality of its habitat. The new record from Sierra Leone increases the extent of occurrence to approximately

Seven individuals were captured: two females and two males over a small creek west of the Bumbuna dynamite building above the inundation zone (coll.: ZFMK 2006.138-.140), one male in riverine forest inside the inundation zone at Kafogo (ZFMK 2006.137), one female in a net perpendicular to the Seli River bank at the Kakutan river crossing, and one male in ravine forest above the BHP drinking water source. In Sierra Leone, R. guineensis was previously known from a single locality 40 km southwest of Bumbuna (Makeni; Grubb et al. 1999, erroneously also listed under R. landeri by Csorba et al. 2003). This species is endemic to the Upper Guinea

Rhinolophus denti knorri Eisentraut, 1960 Knorr’s Horseshoe Bat

One female (ZFMK 2006.141) was caught over a small creek west of the Bumbuna dynamite building above the inundation zone. The taxon knorri is a rare bat endemic to Upper Guinea, known from less than 10 specimens from Guinea, Guinea-Bissau, and northern Ghana. Our specimen is the first record from Sierra Leone. Additional data, in particular genetics, are needed to assess the status of denti Thomas, 1904 and knorri. The large distributional hiatus suggests that both taxa might be specifically distinct, which would necessitate an evaluation of the Red List status of denti. Rhinolophus ziama Fahr, Vierhaus, Hutterer and Kock, 2002 Ziama Horseshoe Bat

Figure 3 Distribution of Rhinolophus ziama in the Upper Guinea highlands with the new locality at Bumbuna. Shaded area in Guinea shows the ‘‘Massif du Ziama Biosphere Reserve’’.

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4500 km2. Current distribution data indicate that R. ziama depends at least partially on forest, and hence the inundation is probably leading to a loss of suitable habitat, particularly owing to flooding of ravine and riverine forest. The criteria for ranking this species as ‘‘Endangered’’ are therefore still met, and we suggest retaining the current threat status. The new record from a topographically complex region supports the suggestion by Fahr et al. (2002) that R. ziama shows a ‘‘paramontane’’ distribution, i.e., being restricted to mountainous and hilly regions although found both at lower and higher elevations. Regions most likely to harbor this species in-between known localities are the Loma Mountains and Tingi Hills in northeastern Sierra Leone as well as the Wologizi Mountains in northwestern Liberia. Family Hipposideridae Hipposideros abae J.A. Allen, 1917 Aba Leaf-nosed Bat

One male (ZFMK 2006.153) was captured in a net set across a trail in ravine forest above the right bank of the Seli River inside the inundation zone. This species was previously known from five localities in the southern half of Sierra Leone at Kent, Njala, 1 mi SE Segbwema, Serabu, and York (Jobling 1952, Rosevear 1965, Grubb et al. 1999; ROM, USNM). Grubb et al. (1999) reported a specimen from Fadugu (USNM 462847) erroneously as H. fuliginosus. Hipposideros cyclops (Temminck, 1853) Cyclops Leaf-nosed Bat

One female (ZFMK 2006.154) of this rather large hipposiderid was caught inside the inundation zone in a net set along a trail in ravine forest above the right bank near the BHP dam. This species was previously known from Kasewe Forest Reserve in central Sierra Leone (Grubb et al. 1999) and from 5.8 km E Fintonia in northern Sierra Leone (USNM 547002-04). Hipposideros vittatus (Peters, 1852) Striped Leafnosed Bat Hipposideros gigas (Wagner, 1845) Giant Leaf-nosed Bat

A member of the commersoni group of very large hipposiderid bats was photographed at Kamasapi village, NW of Kakutan, on 14 April 2006 (Figure 4; C. Ransom unpublished). The individual could represent either the forest species H. gigas or H. vittatus, the latter predominantly occurring in savanna habitats. Both species could occur in the forestsavanna transition zone of central Sierra Leone, and the photographic evidence is insufficient for reliable species identification. Cranial measurements of a specimen from Fadugu (USNM 462846; Koopman et al. 1995, Grubb et al. 1999) kindly provided by M. Carleton (personal communication, 14 Oct 2009) unambiguously allowed identification as H. vittatus. Fadugu is close to our sampling site at Kafogo (Figure 1).

Figure 4 Hipposiderid (either Hipposideros gigas or H. vittatus) roosting in foliage of tree near the village of Kamasapi (photo taken by I. Abu-Bakarr).

Hipposideros jonesi Hayman, 1947 Jones’s Leaf-nosed Bat

Six individuals of this small hipposiderid species were caught: one female (ZFMK 2006.146) in a net across a ravine in riverine forest near the BHP dam, one male (ZFMK 2006.143) over the creek in the forested ravine west of the Bumbuna camp, two males (coll.: ZFMK 2006.145) at the Seli River crossing between Kakutan and Masumarandugu in a net perpendicular to the river, and one male (ZFMK 2006.144) and one female in ravine forest above the Bumbuna camp drinking water source. The type locality of this species is Makeni, approximately 40 km southwest of Bumbuna, and the second record in Sierra Leone is York on the Freetown Peninsula (Hayman 1964). The occurrence of this species appears tightly linked to the availability of caves or artificial substitutes. It is listed as ‘‘Near Threatened’’ (IUCN 2008). Hipposideros cf. ruber (Noack, 1893) Noack’s Leafnosed Bat

With 25 individuals, this was the second most common bat (coll.: ZFMK 2006.147-152), caught in every habitat except on the left bank near the BHP dam. In Sierra Leone, H. ruber has been recorded from numerous localities in the western half of the country (Grubb et al. 1999; ROM). We assigned our specimens to ‘‘H. ruber’’ based on morphology and as traditionally conceived. Vallo et al. (2009) published the first molecular study of bats comprising the caffer-ruber complex, which is notorious for its unresolved and complicated taxonomic relationships. Their study demonstrated substantial cryptic diversity, with at least five distinct clades probably representing distinct species. The Bumbuna specimens might belong to either clade B1 (specimens from Senegal and Benin) or clade D (specimens from Senegal, Ivory Coast, and Benin) as identified by Vallo et al. (2009). As morphological characters have not yet been developed to distinguish among these genetic lineages, the

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identification of our specimens should be considered tentative until inclusion of tissue samples in future genetic studies or a detailed morphological study will allow a more precise identification. Family Nycteridae Nycteris arge Thomas, 1903 Bates’s Slit-faced Bat

One female (ZFMK 2006.133) was caught in the inundation zone at Kafogo in a net set in riverine forest close to the Mawoloko River. A male (ZFMK 2006.134) was caught in the inundation zone at the Seli River crossing between the villages of Kakutan and Masumarandugu. Nycteris arge is known from two other localities in Sierra Leone: one from Kassewe Forest Reserve (Van Cakenberghe and De Vree 1985) and one from 10.1 km NE Fintonia (USNM 546966). Nycteris grandis Peters, 1865 Large Slit-faced Bat

One male (ZFMK 2006.135) was caught in the inundation zone at Kafogo in a net set in riverine forest. One female was caught in the inundation zone in a net set in ravine forest at the Kakutan Seli River crossing. This is the second record for Sierra Leone, the other being from 5.7 km NE of Fintonia (Grubb et al. 1999; USNM 546964). Family Miniopteridae Miniopterus villiersi Aellen, 1956 Villiers’s Longfingered Bat

Five individuals were captured. One male in a raised 12-m net in front of a large water intake tunnel behind the BHP dam (ZFMK 2006.161), one male in riverine forest in a net set over the Mawoloko River at Kafogo (ZFMK 2006.162), and two males and one female in a net across the creek exiting the forest above at the BHP drinking water source (coll.: ZFMK 2006.163, -.164). Wilson and Reeder (2005) listed the taxon villiersi as a subspecies of Miniopterus schreibersii (Kuhl, 1817), but Fahr et al. (2006) argued for its treatment as a separate species pending a taxonomic revision of sub-Saharan Miniopterus. Individuals that probably represent the same species of Miniopterus have been found at 1 mi E Bumban (USNM 462856) and 5.8 km E Fintonia (USNM 547033), Northern Province, as well as 10 mi N Panguma, Eastern Province (USNM 463167, -68, 463312, -13). It has not yet been assessed as a separate species by the Red List (IUCN 2008). Family Vespertilionidae Myotis bocagii (Peters, 1870) Rufous Mouse-eared Bat

All 10 individuals were captured within the inundation zone: one male and one female behind the Bumbuna dam (ZFMK 2006.155, -.156), and five males and three females in a net over a creek near the Kakutan Seli River crossing (coll.: ZFMK 2006.157, -.158). This species is frequently captured over water. Our finding is the second record of this species from Sierra Leone, the first being from 5.8 km E of Fintonia

on the Kaba River in the northern part of the country (Grubb et al. 1999; USNM 547013-15). Neoromicia aff. nana (Peters, 1852)

Two individuals were captured, both within the inundation zone. One female (ZFMK 2006.159) was caught at the southern end of the study area behind the BHP dam, another female (ZFMK 2006.160) at the northern end in a net over the Mawoloko River at Kafogo. The pelage of the specimens is bicolored: dorsal hairs with medium (chestnut) brown tips and dark brown bases, ventral hairs with pale beige/straw-colored tips and almost black bases. The ears are rather rounded, anterior margin evenly convex, posterior margin with a concave indentation at half the length and with a well-developed antitragal lobe at the base. The anterior margin of the tragus is slightly concave, the posterior margin is irregularly shaped but overall is almost straight, the tip is diagonally truncated. There are large skin pads at the base of the thumb and on the inner side of hindfoot; the wing membranes are dark brown. The calcar is large with a moderately developed lobe at the proximal third of its length. The inner upper incisor (I1) is narrow, unicuspid and with a secondary ‘‘step’’; the outer upper incisor (I2) is narrow and long, almost the same length as I1 and reaching around three-quarters the height of I1. The incisors and the canine are separated by a wide diastema. The anterior upper premolar (PM1) is present on both sides, well-developed, and visible from the outside through a rather large gap between the canine and PM2. The rostrum is narrow, distinctly angular against the frontale of the bulbous brain skull; the anterior palatal emargination is rhombical in dorsal view and very narrow posteriorly; the antorbital foramina are rather large. The lower incisors are tricuspid, and only the inner pair is slightly imbricated. Both specimens do not fit in characteristics or measurements with any currently recognized ‘‘pipistrelle’’ species known to occur in West Africa. Pipistrellus hesperidus, Hypsugo crassulus, and H. wcrassulusx bellieri are craniodentally distinctly larger (e.g., in mastoid width, zygomatic width, C-C, M3-M3, C-M3, mandible length, C-M3). Pipistrellus inexspectatus is craniodentally fairly similar but has a longer upper and lower toothrow (C-M3, C-M3). Pipistrellus rusticus, Hypsugo nanulus, and H. musciculus all have shorter forearms; P. rusticus additionally differs from our specimens in having very small outer upper incisors (I2). Pipistrellus nanulus is further characterized by very broad, deeply bifid inner upper incisors. West African specimens (ns31 measured) of Neoromicia nana have a smaller forearm than the two specimens from Bumbuna and are also smaller than the latter in most craniodental measurements (greatest skull length, condylobasal length, width of braincase, height of braincase, zygomatic width, M3-M3, C-M3, mandible length, C-M3). Tissue samples of both specimens were included in a molecular study of African ‘‘pipistrelles’’ (M. Helbig, F. Mayer, and J. Fahr, unpublished data) and, based on mitochondrial as well as nuclear data, group as a branch separate from several lineages of N. nana. However, the Bumbuna

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specimens are distinctly larger in forearm and most craniodental measurements than N. nana and also lack the ‘‘hatchet’’-shaped tragus typical for this species. In conclusion, these specimens probably represent a species new to science, which appears to have close relationships with N. nana based on genetic data, but from which it clearly differs in morphology. Riccucci and Lanza (2008) pointed out that Neoromicia is of feminine gender, hence N. nana rather than N. nanus should be used. Order Rodentia Family Sciuridae Xerus erythropus (E. Geoffroy, 1803) Striped Ground Squirrel

Frequent sightings were made along the BHP construction roads, e.g., near the tall grass hillside. This is a very common ground squirrel throughout Sierra Leone (Grubb et al. 1999). Xerus erythropus thrives near cultivated lands and can become an agricultural pest in Africa (Herron and Waterman 2004). Family Nesomyidae Cricetomys emini Wroughton, 1910 Forest Giant Pouched Rat

Five individuals were captured using Tomahawk live traps, which were set on runways of these large rats with the assistance of local guides. One female (USNM 584880) was caught in forested ravine (Site 3) along the transition from forest to adjacent farmbush under a dense stand of the invasive weed Chromolaena odorata. A female and a male (USNM 584878, -81) were captured in riverine forest at Kafogo above the inundation line. Another male (USNM 584879) and female (escaped) were caught on both sides of the creek in the ravine near the Kakutan Seli River crossing. One of us (J.D.) also observed Cricetomys in the BHP camp ravine. Our captures of C. emini were associated with dense canopy cover and leaf litter. Dermapteran parasites of the genus Hemimerus were collected from four of the five rats captured. On the map in Grubb et al. (1999), C. emini and C. gambianus are shown for 34 quarter-degree squares across most of Sierra Leone, including several individuals not assigned to either species. Family Muridae Hybomys planifrons (Miller, 1900) Liberian Forest Mouse

Four individuals of this species endemic to Upper Guinea were collected: two on the left bank of the Seli River in steeply sloped riverine forest (USNM 584888) and on the riverflats (USNM 584889) approximately 0.5 km north of the Bumbuna dam and two (coll.: USNM 584890) at the Kakutan River crossing. Averaged microhabitat data show a preference for modest canopy cover (35%) and herbaceous ground cover (21.3%), and a strong association with leaf litter (61.3%) closely matching earlier observations of a preference for ‘‘luxuriant understory and dense ground cover’’

reported for this species from Outamba-Kilimi National Park in Sierra Leone (Carleton and Robbins 1985: 987). Our records are the first from central Sierra Leone. At least nine other localities are known for this species from Sierra Leone (Carleton and Robbins 1985, Grubb et al. 1999). Hylomyscus alleni (Waterhouse, 1883) Allen’s Climbing Wood Mouse

A total of 14 individuals of this partly arboreal forest rodent were captured: one (USNM 584919) on the steeply sloping left bank of the Seli River in riverine forest near the Bumbuna dam, four individuals (coll.: USNM 584891) in ravine forest on the right bank of the Seli River, four individuals (coll.: USNM 584892) at the Kakutan Seli River crossing, and five individuals along the creek in the ravine west of the BHP camp. This partly arboreal species was associated with relatively dense canopy cover (31.2%) and dense leaf litter ground cover (56.7%). Hylomyscus alleni is known from numerous localities in the Eastern and Northern Provinces of Sierra Leone (Grubb et al. 1999; USNM). Lemniscomys striatus (Linnaeus, 1758) Striped Grass Rat

One individual (USNM 584893) of this grassland species was captured in ravine forest-farmbush transition, under closed Chromolaena weed cover on the uppermost end of a trapline set along a creek near the Kakutan River crossing. We expected this species to be common on the tall grass hillside but found only Lophuromys there (see below). Our capture site had relatively low canopy cover (25%) and 45% bare soil ground cover. There are many records of L. striatus throughout Sierra Leone except for a small section to the southeast (Grubb et al. 1999; USNM). Lophuromys sikapusi (Temminck, 1853) Rusty-bellied Brush-furred Rat

Twenty-four individuals of this mostly insectivorous rodent were caught: one male (USNM 584884) along a small creek in riverine forest on the left bank of the Seli River near the BHP dam, 12 individuals (coll.: USNM 584882-883, -885, -918) in tall elephant grass and thickets on the hillside east of the BHP dynamite building, two individuals at the edge of a forest ravine below the BHP quarry (coll.: USNM 584886), one female (USNM 584887), and seven males mostly near or in a small rice field at Kafogo, and one female (USNM 584917) on rocks adjacent to a creek near the Kakutan river crossing. Our findings confirm that this species thrives in dense grasslands (64.8%) and low canopy cover (13.3%), conditions that were more common above the inundation line of the reservoir. Its preference for tall grasses (Pennisetum purpureum) was also confirmed by a study conducted in Saama, Eastern Province (Barnett et al. 2000). Lophuromys sikapusi has previously been recorded from numerous localities in Sierra Leone, except in the far north. T. S. Jones found L. sikapusi in Sierra Leone to be ‘‘the

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fourth commonest after Mastomys spp., Praomys tullbergi and Rattus rattus’’ (Grubb et al. 1999: 192).

common rodents captured (60% of total) in the study Barnett et al. (2000) conducted at Saama in Eastern Province.

Malacomys edwardsi Rochebrunne, 1885 Edwards’s Swamp Rat

Mus setulosus Peters, 1876 Peters’s Pygmy Mouse

Eight individuals of this large and long-legged forest rodent were captured all within or very close to the inundation zone: two males (USNM 584894, -96) in riverine forest on the left bank of the Seli River near the BHP dam and six individuals (coll.: USNM 584895, -97) in upper bank riverine forest of the Mawoloko River at Kafogo. Microhabitat data show an association with relatively high canopy cover (35%) and leaf litter ground cover (48.8%). There are at least seven previous collecting localities for M. edwardsi scattered throughout Sierra Leone with ‘‘most records from forest reserves, e.g., Kasewe, Loma Mountains, Bambawo, Freetown Peninsula, and Tiwai’’ (Grubb et al. 1999: 199). Mastomys erythroleucus (Temminck, 1853) Reddish-white Multimammate Mouse

Two females (USNM 584898, -99), one carrying 11 embryos, were caught in elephant grass (average 65% grass cover) along a small rice field surrounded by riverine forest at Kafogo. There are numerous localities for this rodent mapped by Grubb et al. (1999), except for the forested southeastern part of Sierra Leone. Mastomys has been studied in many localities in Sierra Leone in connection with its role as a reservoir for Lassa fever (Monath et al. 1974, Robbins et al. 1983). Robbins et al. (1983) caught 80% of M. erythroleucus, characterized by a chromosome number of 2ns38, in ‘‘bush’’, near villages, and 20% in houses, as compared to 28% captures of the more commensal M. natalensis, characterized by 2ns32, in bush and 72% in houses. During their study in 1978 both species occurred at Kafogo wvillagex. Mus minutoides Smith, 1834 Southern African Pygmy Mouse Mus musculoides Temminck, 1853 Sub-Saharan Pygmy Mouse

A single male (body mass: 4.5 g; USNM 584900) belonging to the minutoides-musculoides complex was captured in elephant grass and ferns along a rice field at Kafogo, where it occurred side by side with the slightly larger M. setulosus (9.5–13.5 g). The smallest West African Mus specimens have been traditionally treated as M. musculoides (Rosevear 1969, Wilson and Reeder 2005), but recent genetic results indicate that both M. minutoides and M. musculoides are present in West Africa (Kan Kouassi et al. 2008). These two species are extremely difficult to distinguish morphologically (Kan Kouassi et al. 2008) and the precise identity of our specimen probably requires genetic determination. Mus minutoides and/or M. musculoides are known from many localities throughout Sierra Leone, whereas M. setulosus appears to be more restricted to the northeastern third of the country (Grubb et al. 1999). Mice in the genus Mus were the most

This small rodent was caught 10 times in the study area. One male (USNM 584901) was caught alongside Lophuromys sikapusi on the tall grass hillside near the BHP dynamite building. The other nine individuals (coll.: USNM 584902, -20) were caught in upland riverine forest and in a rice field at Kafogo. Averaged microhabitat data show a relatively strong association with herbaceous growth (30%) and grasses (36.8%). Mus setulosus was previously recorded from at least six localities in the northeast of Sierra Leone (Grubb et al. 1999). Praomys rostratus (Miller, 1900) Forest Soft-furred Mouse

Our Praomys vouchers were initially identified as P. tullbergi based on morphology, but inclusion of 15 tissue samples in a comparative study of the cytochrome b sequences of P. tullbergi and P. rostratus conducted by Nicolas et al. (2008) grouped them with their Clade II or Clade XII of P. rostratus. With 90 individuals captured, this was the most common rodent in the study. It was absent only from the tall grass hillside and thickets west of the dynamite building and the drier riverine forest at Kafogo. With 58 captures (coll.: USNM 584909, -13 -15, -22), P. rostratus was most common along the creek in the high forest ravine below the BHP quarry. Twelve captures (coll.: USNM 584903-08, -11, -21) were made on the riverflats of the left-bank of the Seli River and steep sloping forest near the BHP dam, 11 captures at the Kakutan River crossing (coll.: USNM 584910, -16), and six captures in the BHP camp ravine. Averaged microhabitat data show a strong association with canopy cover (44.5%), leaf litter (46.7%), and rocks along creeks (26.9%). All Praomys in Grubb et al. (1999) from numerous localities in Sierra Leone were identified as P. tullbergi but may also include P. rostratus. Praomys has been the focus of a population study in lowland forest at Kasewe Forest Reserve and Bambawo near Kenema. At both sites it was the most abundant rodent (Cross 1977). In a study of cultivated sites at Saama, Praomys was most abundant in 3-year-old cocoa groves with dense undergrowth (Barnett et al. 2000). Additional mammal observations

As a separate team conducted a year-long survey of primates and other large mammals at Bumbuna (Ransom et al. 2006), our observations are limited to two species observed and photographed near our traplines. Order Carnivora Family Nandiniidae Nandinia binotata (Gray, 1830) African Palm Civet

At least two individuals of this arboreal carnivore were observed and photographed at approximately 21:45 h foraging on flowers of large Parkia biglobosa trees along the

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forested ravine near the BHP camp. One individual was observed catching a small fruit bat, possibly Micropteropus pusillus or Nanonycteris veldkampii, both of which have been documented to feed on the spherical capitula of P. biglobosa (Baker and Harris 1957, Hopkins 1983). Nandinia binotata is known from Sierra Leone from at least 19 localities throughout the country (Grubb at al. 1999). Order Primates Family Lorisidae Perodicticus potto (Mu¨ller, 1766) Potto

One individual of this nocturnal prosimian was observed at 20:00 h on 26 January 2006 in vines above the creek in the ravine west of the BHP camp below the dam and subsequently photographed. Perodicticus potto is known from at least 13 localities mostly in the forested southeast of Sierra Leone (Grubb et al. 1999).

Discussion We confirmed the presence of 35 species of small mammals, including three shrew (Soricomopha), 19 bat (Chiroptera), and 11 rodent (Rodentia) species in and near the BHP inundation zone, an area that had not previously been surveyed for small mammals. Our results highlight the importance of small mammal inventories prior to large development projects in Sierra Leone given the long hiatus of mammal research imposed by the civil war on the country. Our species accumulation curve for terrestrial small mammals (Figure 2) did not reach an asymptotic plateau, and our sampling included two species recorded by a single individual as well as four species based on two individuals. These results suggest that additional sampling from the area would yield additional species. However, we report the first record of Crocidura nigeriae for Sierra Leone, although earlier records of C. poensis may include individuals properly attributed to C. nigeriae (see above). Additional species, which have been recorded from the immediate vicinity (i.e., Bumban, Fadugu, Kafogo, Kamagbengbe) but were not sampled in our survey include Crocidura olivieri (Lesson, 1827) (USNM 463363), Gerbilliscus kempi (Wroughton, 1906) (USNM 462858-61, 462867-77, 538489), Mastomys natalensis (Smith, 1834) (many USNM specimens), Uranomys ruddi Dollman, 1909 (USNM 463072-77, 538782-87), and the commensal Rattus rattus (Linnaeus, 1758) (USNM 462885-86, 463408-18), which adds up to four shrew and 15 rodent species known from the wider Bumbuna area. The terrestrial small mammal fauna sampled in the BHP area is composed of a forest-savanna ecotone assemblage, with forest species clearly dominating the riverine and ravine forests. No predominantly commensal or invasive species were encountered, indicating a rather undisturbed species assemblage in the study area, with trap lines located between 0.5 km (BHP camp ravine) and several kilometers from human habitations. Our rodent trapping success (1728 trap nights, 163 captures, 10 species) is comparable to that of Cross (1977) at Kasewe (2271 trap nights, 274 captures,

eight rodent species) and Bambawo (1410 trap nights, 147 captures, nine rodent species), or Barnett et al. (2000) at Saama (4315 trap nights, 138 captures, 10 rodent species). However, much more intensive sampling can yield additional shrew species as demonstrated by Nicolas et al. (2009), who reported 11 species from 2509 shrews captured at Massif du Ziama Biosphere Reserve in Guinea with a trapping effort of 20,160 bucket-nights spread over 4 months and six habitat types. The bat data exhibit an even stronger indication that several additional species would be recovered with additional effort. The species accumulation curve (Figure 2) rises steeply without a plateau, and our sampling includes five species represented by a single individual as well as three species represented by two individuals. Three species of bats (Rhinolophus denti knorri, R. ziama, and Neoromicia aff. nana) represent first records for the country, and five species (Nanonycteris veldkampii, Myonycteris torquata, Rhinolophus guineensis, Nycteris grandis, and Myotis bocagii) are second records. Three additional species have been previously recorded from the immediate vicinity of our study area: Pipistrellus inexspectatus Aellen, 1959 (USNM 462852), Neoromicia somalica (Thomas, 1901) (USNM 462850), and Mops condylurus (A. Smith, 1833) (USNM 462857), all from Fadugu. The combined total richness of 22 bat species recorded from the Bumbuna region is in the range of more intensively surveyed areas in southeastern Guinea, where 16–30 species have been registered in five protected areas (Fahr and Ebigbo 2003, Fahr et al. 2006). The species composition of the bat assemblage is characterized by a mixture of both savanna- and forest-dwelling species, with a slight dominance of the latter. Conspicuous is the large number of species known or suspected to depend on caves or artificial substitutes as day roosts (Rousettus aegyptiacus, Rhinolophus guineensis, R. denti, R. ziama, Hipposideros abae, H. jonesi, and Miniopterus villiersi). High-flying species such as emballonurids and molossids were completely missed by the present survey owing to a lack of mist nets set at canopy level. The comparatively low number of vespertilionids is more difficult to explain but might have been compensated if a harp trap had been employed. Koopman et al. (1995) provided the first recent summary of bat species documented for Sierra Leone. They listed 57 species plus one considered questionable (Epomops franqueti). Of these, three species have been re-identified as taxa otherwise known from Sierra Leone (Nycteris thebaica rather than N. gambiensis, see Van Cakenberghe and De Vree 1988; Hipposideros cf. caffer rather than H. lamottei, see Decher and Fahr 2007; Mops wbrachypterusx leonis rather than M. petersoni, ROM 77463) and are thus deleted from the country list, while one species was omitted (Scotophilus nux, Robbins et al. 1985). Grubb et al. (1999) largely followed Koopman et al., and additionally listed Hypsugo musciculus for Sierra Leone and Ghana based on specimens in the BMNH and USNM, a species currently known from Central Africa, but without further justification. We accept a total of 55 species as confirmed, whereas questionable species and specimens should be re-examined. Our study added three

Article in press - uncorrected proof 174 J. Decher et al.: Small mammals of the Seli River valley, Sierra Leone

species, resulting in a current total of 58 bat species confirmed for Sierra Leone. The presence of globally threatened bat species, in particular Rhinolophus ziama (‘‘Endangered’’) and Rhinolophus guineensis (‘‘Vulnerable’’), indicates that the BHP area is of considerable concern for biodiversity conservation. Although these species were partly found outside the inundation zone of the reservoir, flooding of the Seli River valley will displace forest species to higher elevation areas that are more heavily affected by human uses and encroaching bushfires. Loss of forest patches owing to the reservoir will result in a more homogeneous landscape and probably reduced mammal diversity as forest-dependent species are expected to be vulnerable to this habitat loss. The creation of a proposed Bumbuna Conservation Area, designed mainly to protect the chimpanzee population of the area, would also benefit many other mammals. Ideally, a protected area should include viable sections of all habitats lost by the flooding of the Seli River valley either in the immediate vicinity or as an offset in another region of Sierra Leone, such as at Loma Mountains, which have previously been proposed as a new national park (Atkinson et al. 1996). In conclusion, our results show that the small mammal fauna of the upper Seli River Valley was poorly known. This conclusion also holds true for the mammal fauna of Sierra Leone, as evidenced by our first and second country records of several species. Given the brevity and single-season effort of our survey, we are confident that additional species will be found in this region and we recommend additional surveys in the future. Our data on small mammals provide a baseline to assess the ecological effects of flooding by the dam on these communities, a situation that has been rarely studied in Africa compared to the Neotropics (Granjon et al. 1996, Cosson et al. 1999, Meyer and Kalko 2008).

Appendix Gazetteer of localities. Bambawo (near Kenema) Bo Bumban, 1 mi E Fadugu Fintonia, 5.7 km NE (Outamba-Kilimi National Park) Fintonia, 5.8 km E, Kaba River (Outamba-Kilimi NP) Fintonia, 10.1 km NE (Outamba-Kilimi NP) Freetown Peninsula (Freetown) Kafogo Kakutan Kamagbengbe Kamasapi (NW of Kakutan) Kasewe Forest Reserve wsKassewax Kent Loma Mountains Makeni Njala Panguma, 10 mi N Saama (Eastern Province) Segbwema, 1 mi SE Serabu Tiwai Island York Bakel, Senegal Wonegizi Mountains, Liberia Massif du Ziama, Guinea Pic de Fon, Guinea Zie´la, Mount Nimba, Guinea Kabwir, Nigeria Otjimbingue, Namibia

88009 78589 98079 98239 98429

N, N, N, N, N,

118079 118459 118549 118469 128119

W W W W W

98409 N, 128109 W 98459 N, 128109 W 88309 98249 98099 98119 98119 88199

N, N, N, N, N, N,

138159 118449 118439 118509 118459 128119

W W W W W W

88109 N, 138109 W 98139 N, 118079 W 88539 N, 128039 W 88079 N, 128059 W 88179 N, 118069 W 88109 N, 118009 W 78599 N, 108569 W 78489 N, 128039 W 78319-359 N, 118209-229 W 88179 N, 138119 W 148549 N, 128279 W 88089 N, 98289 W 88239 N, 98179 W 88329 N, 88549 W 78439 N, 88219 W 98249 N, 98349 E 228219 S, 168089 E

Acknowledgements This project was funded by Nippon Koei UK (NKUK) and supported by the Sierra Leone Government Project Implementation Unit (PIU) in Freetown. Sierra Leone Director of Forests, B. M. Kamara, kindly issued export permits for voucher specimens. We are grateful to Paul Driver and Derek Johnson (NKUK) as well as Abdulai Barrie (PIU) for addressing all logistical needs, and to John Maxen, Natural History Museum, London, for organizing the field teams. Thanks to local counterparts Alie B. Fofanah (Forestry Department, Wildlife Conservation Branch) and Joseph B. Bangalie (Center for Biodiversity Research), and to guides Mohamed Koroma, Foday Marah, and Minkailu Bantama for their invaluable assistance. We are grateful to the people of Kafogo and Kakutan for allowing us to work on their land. Thanks to the staff of Salini Construttori for hospitality at the BHP camp and much logistic support. We are particularly grateful to Rainer Hutterer (ZFMK), Michael Carleton, Linda Gordon and Dave Schmidt (USNM), and C. William Kilpatrick (University of Vermont) for voucher specimen identification and preservation, and equipment loans. Thanks to Ibrahim Abu-Bakarr for the photograph of the large Hipposideros. J.F. was supported by the BIOTA program, funded by the German Federal Ministry of Education and Research (BMBF, project 01LC0617E1). We appreciate the comments of three anonymous reviewers that helped to improve the manuscript.

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