Annotated checklist of bats (Mammalia: Chiroptera) of Mount Cameroon, southwestern Cameroon

ABSTRACT Mount Cameroon is a priority area for global biodiversity conservation, considering its high species richness and endemism across many taxa. As of yet, its Chiroptera fauna is still poorly known. Thus this paper aims to provide an updated checklist of the Chiroptera fauna of Mount Cameroon based on our own field data, previously published data and data from museum collections. Details on published references and conservation statuses are also provided for each species. During our field surveys (2016-2018), 613 individuals and 21 species were recorded. Among them three species are new to the area: Mops (Xiphonycteris) nanulus J. A. Allen, 1917, Mops (Xiphonycteris) thersites (Thomas, 1903) and Glauconycteris egeria Thomas, 1913. These new records bring the number of known bat species in the Mount Cameroon area to 38, belonging to seven families. This includes Pteropodidae Gray, 1821 (eleven species), Hipposideridae Gray, 1821 (six species), Rhinolophidae Gray, 1825 (three species), Miniopteridae Dobson, 1875 (three species), Vespertilionidae Gray, 1821 (seven species), Nycteridae Van der Hoeven, 1855 (five species) and Molossidae Gervais, 1856 (three species). We also observe that 23.7% of bats in the area are frugivorous (nine species), 71.1% are insectivorous (27 species) and 5.3% are nectarivorous (two species). Moreover, because of the increasing rate of deforestation in the area caused by slash-and-burn shifting cultivation and expansion of agro-industrial plantations, knowledge of the regional chiropteran fauna is imperative, especially as this information can provide a base for the development of future management and conservation strategies.


INTRODUCTION
Cameroon is one of the richest biodiversity countries in Africa, with high levels of endemism among various taxa (Alpert 1993). This rich biodiversity can be attributed to the country's diverse ecosystems suited for a wide range of biota. The number of plant species is estimated at over 9,000, of which 160 are endemic (Sunderland et al. 2003). At least 409 mammal species occur, of which 10 are endemic (Tchigio 2007); of these mammals, bat account for approximately 120 species according to ACR (2018). Moreover, most of the country's endemic taxa are concentrated on Mount Cameroon and other Cameroonian Highlands situated in the southwestern part of the country.
Mount Cameroon belongs to an ecoregion located on a volcanic chain called the Cameroon Volcanic Line (CVL). This volcanic chain extends northeast along the border between Cameroon and Nigeria, and southwest towards the islands of Bioko, São Tomé, Príncipe and Annobon. The ecoregion also encompasses the Cameroonian Highlands: a mountain chain that runs north east from Mount Cameroon along the coast through the Bakossi, Bamboutos and Bamenda Highlands, and finally ending further north at Tchabal Mbabo Mountains on the Adamaoua plateau. The Cameroonian Highlands are recognized as an area of high species diversity and endemism across many taxa (Stuart 1986). Moreover, the Mount Cameroon area is considered as a richness and endemism hotspot for bats in Africa (Herkt et al. 2016). This exceptional species richness is attributed to the heterogeneity of the habitats found within the area, which ranges from coastal ecosystems through submontane and montane forests to subalpine grasslands. The significance of Mount Cameroon and other Cameroonian Highlands as important hotspots with biodiversity with many endemic taxa has long being recognized (Bergl et al. 2007). A possible reason for this unique diversity of Mount Cameroon is probably because it was one of the Pleistocene refuges (Maley 1991), together with the relative spatial isolation of the montane environment that resulted in increased speciation rates.
Mount Cameroon is listed among World Biodiversity Hotspots (Myers et al. 2000), and its rich biodiversity is threatened by anthropogenic activities. There are at least 42 plant species including three genera that are strictly endemic and another 50 species are near-endemic (Cable & Cheek 1998). The avifauna is estimated at over 370 species recorded, a number of which are endemic (Fotso et al. 2001). Two of these bird species are strictly endemic to Mount Cameroon notably the Mount Cameroon francolin (Francolinus camerunensis Alexander, 1909) and the Mount Cameroon speirops (Speirops melanocephalus Gray, 1862), which are listed as Endangered and Vulnerable respectively (Fotso et al. 2001).
Despite Mount Cameroon's importance for conservation, its bat fauna still remains under-sampled. Pioneer surveys in the area were conducted in the 1950's and 1960's by Eisentraut (1956Eisentraut ( , 1963Eisentraut ( , 1964Eisentraut ( , 1968 and Hill (1968), with very few surveys carried out since then. There have been a growing number of studies on bat species richness and diversity in the past two decades in Cameroon, greatly adding to our knowledge on this order of mammals (Hutterer et al. 1992;Sedláček et al. 2006;Bakwo Fils 2009Bakwo Fils et al. 2012, 2014Hassanin 2014). These studies focused on other ecoregions of the country and none involved the Cameroonian Highlands in general and Mount Cameroon in particular. Furthermore, Mount Cameroon is characterized by fertile volcanic soils, which has favored the clearing of the lowland forest to make way for large-scale plantation agriculture (Forboseh et al. 2011). This is likely to have a considerable impact on biodiversity of the area. Other threats to biodiversity in the area include burning of vegetation, slash-and-burn agricultural practices, uphill shifting cultivation and poaching. Given these threats, it is important to publish an updated checklist of bats of the area in order to document the species that are most in need of conservation measures. Furthermore, knowledge about the bat ecology of the area is important in the development of future conservation plans for these species, which is currently lacking.
The purpose of this study is therefore to compile an updated checklist of the Chiropteran fauna of Mount Cameroon from field surveys, published literature and museum collections, for the purpose of conservation. Data from previous studies by Eisentraut (1956;1963;1964;1973) and Fedden & MacLeod (1986) were compared to our recent survey in order Manga Mongombe A. et al. to determine whether the populations of the species have increased or decreased during the past few decades.

DEScRIpTION Of STuDy SITE
Mount Cameroon is an active volcano located in the Gulf of Guinea between latitudes 3°57' to 4°27'N and longitudes 8°58' to 9°24'E (Suh et al. 2003) (Fig. 1). The main axis of this roughly elliptical shaped mountain is 45 km long and 30 km wide (Tame 1999), and the mountain occupies an area of about 1500 km² (Tchouto 1995). The mountain's original vegetation was described by Thomas & Cheek (1992) as consisting of a gradual gradient that starts with evergreen lowland rainforest at lower altitudes (0-800 m a.s.l.), through submontane forest (801-1600 m a.s.l.), montane forest (1601-1800 m a.s.l.), montane scrub (1801-2400 m a.s.l.), montane grassland (2000-3000 m a.s.l.) and finally sub-alpine grassland (3001-4100 m a.s.l.) near a bare summit. Unfortunately, most of the lowland forest has already been cleared and transformed into human settlements and large agro-industrial companies that grow mainly export crops such as rubber (Hevea brasiliensis), oil palm (Elaeis guineensis), tea (Camellia sinensis) and banana (Musa spp.) (Forboseh et al. 2011). Small-holder farmers in the region also extensively cultivate crops for subsistence and sale around the villages on the slopes of the mountain.
The climate of Mount Cameroon is described as predominantly humid tropical with heavy precipitation and high temperatures along the coastal belt. There are two distinct seasons: a rainy season that occurs between the months of June and October, and a dry season that extends from November to May. The annual rainfall on the mountain varies between 2085 and 10000 mm and the average annual humidity is high at about 75-80% (Payton 1993). The mean annual temperature is about 25°C, and temperature decreases by 0.6°C per 100 m ascent (Fraser et al. 1998). The hydrological network is poor with few water bodies mainly found at the lower slopes of the mountain.

DATA cOLLEcTION
Data were gathered by searching internet bases, examining museum collections and by carrying out field surveys. At the onset, we collected data by searching the internet for publications made on bats of Mount Cameroon through Google Scholar (https://scholar.google.com): by typing key words such as "Chiroptera of Mount Cameroon", "montane bat of Cameroon", "Mount Cameroon bats". We also consulted the ACR (2018) for information on type locality of African bat species. Additionally, we examined voucher specimens of bats collected from Mount Cameroon in the following museum collections in Germany: Zoological Research Museum Alexander Koenig (ZFMK), the State Museum of Natural History Stuttgart (SMNS) and the Berlin Zoological Museum (ZMB). These museums house some of the earliest bat specimens collected by Prof. Martin Eisentraut in the Mount Cameroon area.
Field surveys were also conducted over a 24 months period from November 2016 to November 2018 in the four forest types of Mount Cameroon. In each forest type, we established mist net sampling stations, four in lowland forest, six in submontane forest, two in montane forest and three in montane scrub (Table 1). At each sampling station, we deployed six nylon mist nets (Ecotone©, Poland), 12m long, 2.5m high, five shelves, mesh: 16, denier 70/2. We opened mist nets at dusk and operated them for at least six hours during capture nights depending on weather conditions, but we often had to close mist nets early on rainy, very windy nights. We also avoided sampling during full moons. In order to prevent the capture of familiar individuals and to maximize capture success, the location of each mist net was changed on subsequent capture nights within each capture station (Bergallo et al. 2003). The coordinates of sampled sites were recorded using a hand-held GPS (Garmin eTrex). The mist nets were deployed in different habitat types within each sampling station in order to maximize the chances of capturing different species (Table 1; Fig. 2A-I). The mist nets were attended to regularly and each captured bat was carefully extracted to avoid injury. We recorded parameters such as the sex, weight, age (juvenile or adult), and the reproductive status (testicles abdominal or scrotal for males; pregnant, lactating or post-lactating for females) (Racey 1988). Weight was recorded using a Ohaus YA501 electronic pocket scale, 500x0.1g, and standard museum measurements were recorded using a dial caliper (Ecotone©, Poland 150/0.1mm). The following external measurements were used for the identification of each bat to species level: forearm length (FA), tibia length (TiL), tail length (TaL), head and body length (HBL), ear length (EL), hind foot length (HFL), tragus length, shape (TrL) and greatest length of skull (GLS). In addition, the arrangement of palatal ridges was also examined to aid identification of fruit bats (Monadjem et al. 2010;. Identification was based on information and illustrations published in the keys of Rosevear (1965), Hayman & Hill (1971), . After identification, bats were released near the location where they were captured. Species which were difficult to identify were preserved in 70% alcohol for later identification based on close examination of craniodental features, and deposited at the zoological laboratory of the University of Maroua. Craniodental measurements were also recorded from preserved bats for the three species recorded for the first time: Mops (Xiphonycteris) nanulus J. A. Allen, 1917, Mops (Xiphonycteris) thersites (Thomas, 1903) and Glauconycteris egeria Thomas, 1913. The greatest length of skull (GLS), condylo-basal length (CBL), condylo-canine length (GCL), zygomatic breadth (ZB), breath of brain case (BB), maxillary toothrow length (C-M 3 ), mandibular toothrow length (C-M 3 ), and the mandible length (M) were measured following Bates et al. (2005).
HAbITATS AND DISTRIbuTION. -This bat is widely distributed across the lowland rainforest and savannah zones of sub-Saharan Africa from Senegal in the west, to Ethiopia in the east, to southern South Africa (Mickleburgh et al. 2008a), and it has also been recorded on the Arabian Peninsula (Mickleburgh et al. 2008a). It is a species that often roosts amidst human habitations (Webala et al. 2014). This species is gregarious and colonies may number in tens of thousands (Monadjem et al. 2010). According to Happold (1987) they prefer tall dead trees with bare branches of particular species for roosting.
They are threatened across West and Central Africa by harvesting for bushmeat and medicinal use (Mickleburgh et al. 2008a ORIgINAL DATA. -79 individuals (22 males and 57 females) of this species were captured during our field surveys in disturbed habitats at altitudes below 1300 m a.s.l. (Table 1). Both Eisentraut (1963) and Fedden & MacLeod (1986) also recorded this species in lowland, disturbed forest in Mount Cameroon.
HAbITATS AND DISTRIbuTION. -This species is mostly distributed in Central Africa, but can also be encountered in parts of West and East Africa (Kityo & Nalikka 2016). This species inhabits low to mid-elevation rainforest and some relic forest in savannah regions (Happold 1987). Happold (2013a) pointed out that this species prefers secondary to primary forest and is often caught in gardens, and in clearings close to dense undergrowth where it roosts in dense foliage of large trees.
Genus Hypsignathus H. Allen, 1862 Hypsignathus monstrosus H. Allen, 1862 Hypsignathus monstrosus H. Allen, 1862: 158. ORIgINAL DATA. -The hammer-headed fruit bat was not captured during our field surveys, but had previously been recorded in the lowland and montane forest of Mount Cameroon by Eisentraut (1963), and by Fedden & MacLeod (1986) who capture one individual in coastal vegetation at Bimbia.
HAbITATS AND DISTRIbuTION. -This species is mainly distributed in West and Central Africa, extending marginally to East Africa. This species is an inhabitant of lowland tropical rainforest, particularly gallery forest, swamp forest, mangroves, palm forest, and it has also been recorded in forest patches in savannah areas (Happold 1987 This specialist nectar and pollen eater was mostly captured in low altitude disturbed and cultivated habitats, though 11 individuals were also netted in undisturbed lowland forest and one in montane scrub (Table 1). The species had previously been documented on Mount Cameroon (Eisentraut 1963(Eisentraut , 1973Hayman & Hill 1971;Fedden & MacLeod 1986).

HAbITATS AND DISTRIbuTION. -This species is widespread in Central
Africa but marginal to Southern Africa. According to Bergmans (1997), it typically inhabits lowland tropical rainforest, although it has also been recorded from swamp forest, Mangrove forest and mosaic forest-grassland habitats where it commonly forages in cultivated area adjacent to forest.   (2013) also noted that this species prefers secondary to primary forest and is often caught in gardens, and in clearings close to dense undergrowth. They roost singly or in pairs in lower part of trees among dense foliage within few meters of the ground (Happold 1987 ORIgINAL DATA. -The species was the most abundant captured during field surveys, with 174 individuals (18 males and 156 females, Table 1). Individuals were recorded in different habitat types such as cultivated and fallow farmland, entrance of caves, around fruiting mango trees (Mangifera indica) at low to mid-altitudes, and in understory canopy of high altitude primary montane forest (Table 1). Eisentraut (1963;1973) and Fedden & MacLeod (1986) captured this species in both primary and disturbed forest in a wide range of altitudes on Mount Cameroon.

REMARk. -Recent studies based on molecular analyses revealed that
HAbITATS AND DISTRIbuTION. -This species is widely distributed in West, Central and East Africa, but marginal in southern Africa. The species is an inhabitant of lowland rainforest and montane forest from sea level to 4000 m a.s.l. It has also been recorded in relic forest in savannah. Small colonies roost in hollow trees and entrance of caves (Rosevear 1965;Happold 1987). Thomas (1983) noted that immature male cohorts of this species migrate between forest and savannah habitats in West Africa.
REMARk.    ORIgINAL DATA. -The mainly nectar-feeding Veldkamp's dwarf epauletted fruit bat was represented by six individuals (Table 1): one female was netted in a farmland at an altitude of 20 m a.s.l. and five other individuals (3 males and 2 females) were captured at another cultivated site at an altitude of 470 m a.s.l. (Table 1). The species was previously recorded from Mount Cameroon by Eisentraut (1973).
HAbITATS AND DISTRIbuTION. -This species is widely distributed in West Africa and western parts of Central Africa. It principally inhabits disturbed lowland rainforest, but has also been recorded in savannah habitats, gallery forest and isolated forest patches where it roosts singly or in pairs in the lower parts of shrub and trees among dense foliage (Happold 1987). Thomas (1983) noted that this species shows migratory patterns with both sexes moving between forest and savannah habitats in West Africa. ORIgINAL DATA. -With 137 individuals (131 males and 6 females) the echolocating and cave-dependent Egyptian Rousette was the second most abundant bat species captured during our field surveys (Table 1). The species was trapped in both primary montane forest and lowland secondary and cultivated forest at a wide range of altitudes from 470 m up to 2280 m a.s.l. (Table 1). The species has previously been sampled in large caves on the lower slopes and in montane grassland of Mount Cameroon (Eisentraut 1963;Fedden & MacLeod 1986).
HAbITATS AND DISTRIbuTION. -Rousettus aegyptiacus is widespread in sub-Saharan Africa, North Africa, Middle East, the Arabian Peninsula right across to Iran and Pakistan (Korine 2016). This species has broad habitat preference and has been recorded in arid to moist tropical and subtropical habitats where large colonies roost in caves and artificial structures such as mosques, tombs and mines (Happold 2013c). Happold (1987) noted that the local distribution of this species is influenced more by the availability of suitable roost sites than by vegetation type. The species is threatened in parts of its range by bushmeat trade, cave disturbance and persecution (Korine 2016).
DIAgNOSIS. -The four specimens display the characteristic brown head with prominent white spots on the forehead between nostrils and the eyes, small white spots at the posterior corner of each eye, and lips fringed with white (Fig. 3). The external measurements (Table 3), and description generally agree with those of Fahr (2013b), and Nigerian specimens described by Happold (1987).
HAbITATS AND DISTRIbuTION. -This species has a limited distribution in Central Africa, centered on Cameroon (see Hassanin et al. 2015). According to Bergmans (1990), this species inhabits tropical lowland and montane forest where it lives in the lower strata of the forest about 3 m above the ground (Happold 1987); it roosts singly in trees or bushes (Happold 1987;Fahr 2013b). It is threatened by habitat loss through deforestation and the conversion of primary forest to agricultural use.
REMARk  (Hill 1968), but was not recorded during our field surveys. The habitat of this species is tropical rainforest, but could extend into savannah zones. This species roosts in caves, hollow trees, the roofs of thatched houses and in old mine shafts (Happold 1987   ORIgINAL DATA. -Three individuals of this species were captured during our surveys (Table 1). One was mist-netted in a fallow farmland at an altitude of 1140 m a.s.l, while the other two were captured in primary forest at an altitude of 1070 m a.s.l.
HAbITATS AND DISTRIbuTION. -At Mount Cameroon, this cave dependent species had previously been recorded at altitudes up to 1250 m a.s.l. (Fedden & MacLeod 1986), who also noted that this species inhabits both disturbed forest fringes and montane forest. Eisentraut (1963) also recorded this species in the lowland forest of Mount Cameroon at Bimbia and Limbe area, and in a cave at Buea. This species principally inhabits lowland rainforest, but has also been recorded in savannah and gallery forest (Happold 2013d

HAbITATS AND DISTRIbuTION. -This species is widely distributed in West and Central Africa, with a few isolated records from East
Africa. This species principally inhabits undisturbed tropical rainforest (Happold 1987), including lowland, coastal, montane, swamp and mangrove forests (Fahr 2013c as Hipposideros cyclops). It has also been recorded in isolated forest patches on the edges of savannah (Decher & Fahr 2005) where it roosts singly, in pairs or in small groups in cavities of hollow standing trees. They have a preference for cavities located high above the ground, and often return to the roost to consume their prey (Happold 1987). Decher & Fahr (2005) mentioned that this species may be threatened by deforestation in some parts of it range.
REMARk ORIgINAL DATA. -The occurrence of this species in Mount Cameroon was mentioned by Hill (1968), but we did not capture any individual attributed to this species during our field surveys.
HAbITATS AND DISTRIbuTION. -Hipposideros caffer is widespread in most biotic zones of sub-Sahara Africa, particularly woodland savannah habitats, extending into some suitable localities in the rainforest zones (Happold 1987;Bernard & Happold 2013). The species generally prefers savannah habitats but has also been recorded in coastal forest and bushveld habitats close to water bodies (Bernard & Happold 2013). Large colonies roost in caves, mines, buildings and culverts (Happold 1987;Monadjem et al. 2010). REMARk.
-Hipposideros caffer is regarded as a species complex (Kock et al. 2008;Vallo et al. 2008). Three subspecies were recognized by Simmons (2005) ORIgINAL DATA. -A total of 41 specimens of this species were capture during our surveys, of which most were in the understory canopy of primary forest at an altitude of 630 m a.s.l. (Table 1). This corroborates the report by Fedden & MacLeod (1986), who mostly recorded this species at low altitude. Eisentraut (1963) noted that Hipposideros ruber inhabits lowland regions, but could extend into montane forest.
DIAgNOSIS. -The forearm measurement of these specimens are 49.1 to 53.4 mm (Table 3). Other features stated by Happold (1987), such as the variable color phases ranging from orange-rufous to greyish-brown were present in these specimens. Also the ears are large and pointed and the antitragus is well developed. The species is distinguishable from Hipposideros caffer by the larger forearm length of above 50 mm (Patterson & Webala 2012) ( Table 3).
HAbITATS AND DISTRIbuTION. -This species has been recorded throughout much of West, Central, and East Africa and part of southern Africa (Monadjem et al. 2017c). According to Happold (1987), this species mostly inhabits lowland tropical rain forest, but can also be found in relic and riverine forests in savannah regions where large colonies roost in caves, rocky cavities and abandoned mineshafts.
REMARk. -The species is well known for its variable color phases. The pelage gradually changes from grey or brown just after they early molt to bright russet, as it is oxidized by ammonia fumes within the roosts (Fedden & MacLeod 1986). Recent molecular evidence place Hipposideros ruber as belonging to the caffer/ruber complex whose taxonomy is still confused (Vallo et al. 2008(Vallo et al. , 2011Monadjem et al. 2013 where small colonies typically roost in caves, but occasionally could be found roosting in dense vegetation (Happold 1987). In Liberian Mount Nimba this species was captured in forested and disturbed habitats, ranging from 460 m to 1060 m a.s.l. (Monadjem et al.2016). According to Happold (1987) this species does not hunt daily, it preys mostly on large beetles and often return to the roosts to consume prey. Family Nycteridae Van der Hoeven, 1855 Genus Nycteris G. Cuvier and E. Geoffroy, 1795 Nycteris arge Thomas, 1903 Nycteris arge Thomas, 1903: 633. ORIgINAL DATA. -Our recent field surveys did not yield any specimen attributed to this species. HAbITAT AND DISTRIbuTION. -This species is widely distributed over much of West and Central Africa from Sierra Leone in the West, through the Congo Basin to East Africa and the Democratic Republic of the Congo. Nycteris arge is predominantly a lowland rainforest inhabitant, although in southern Africa it can be encountered in forest edges (Monadjem et al. 2010). It roosts singly or in small groups in hollow cavities of trees with large trunks, especially trees with opening near the ground (Happold 1987). According to Rosevear (1965), this species forages in forest clearing and may enter houses to hunt insects attracted to strong light. This species has been listed to occur on Mount Cameroon (Van Cakenberghe & De Vree 1985).

Nycteris grandis Peters, 1865
Nycteris grandis Peters, 1865: 358. HAbITATS AND DISTRIbuTION. -It is predominantly a lowland rainforest species, but it also occurs in savannahs, and along riparian forest where it roosts in hollows of large trees, holes or small caverns in rocks and artificial structures such as disused water tower (Happold 1987;Monadjem et al. 2010). It is broadly distributed in West, Central and East Africa (Monadjem et al. 2017a). It forages close to the ground, in open forests, and near edges of clearings and over streams. Arthropods and small vertebrates such as frogs, birds, fish and small bats constitute the diet of this species making it the only truly carnivorous bat in Africa that eat vertebrate prey (Fenton et al. 1983). This species had previously been signalled in the Mount Cameroon area (Van Cakenberghe & De Vree 1985). During our field surveys, no individuals attributed to this species were recorded. This species may be threatened in some parts of its range by habitat conversion and overharvesting for food (Monadjem et al. 2017a).

Nycteris hispida (Schreber, 1774)
Vespertilio hispidus Schreber, 1774: 169. ORIgINAL DATA. -We did not record any individuals attributed to this species during our field surveys, but the species has previously been recorded at Buea, Mount Cameroon by Eisentraut (1963).

HAbITATS AND DISTRIbuTION. -This species is widely distributed throughout sub-Saharan Africa (Van Cakenberghe & De
Vree 1993). The species occupies a variety of habitats including savannah, woodland and forest where it roosts in dense bushes, houses, hollow trees, and caves (Monadjem et al. 2010). The occurrence of this species on Mount Cameroon was previously documented by Eisentraut (1963), but no individual was recorded during our field surveys. Fedden & MacLeod (1986) noted that this species has a wide African distribution, and is often associated with human habitation and common in cultivated areas. In Liberian Mt. Nimba it was recorded sparsely in open savannah (Monadjem et al. 2016). This ubiquitous species roosts singly or in groups in holes in the ground or on twigs close to the ground (Happold 1987). Its diet consists of insect such as moths and mantises which are apprehended by slow-hawking or gleaning (Happold 2013f ). Aellen, 1959 Nycteris intermedia Aellen, 1959: 218. cOMMON NAME. -English: Intermediate Slit-faced Bat. French: Nyctère d'Aellen HAbITATS AND DISTRIbuTION. -We did not encounter this species during our field surveys and did not examine any specimen of this species. Nycteris intermedia has been widely documented in West and Central Africa (Van Cakenberghe & De Vree 1985). The species mainly inhabits lowland rainforest and coastal forest (Monadjem et al. 2010;Denys et al. 2013;Fahr 2013d). The species was listed to occur in Mount Cameroon by Van Cakenberghe & De Vree (1985). It forages by gleaning insects close to the ground in the understory of forest (Fahr 2013d).

Nycteris intermedia
Nycteris major (K. Andersen, 1912) Petalia major K. Andersen, 1912: 547. ORIgINAL DATA. -Previously, this species has been recorded at the southwestern slope of Mount Cameroon by Eisentraut (1973), but no individuals were encountered during our field surveys.
HAbITATS AND DISTRIbuTION. -This species is distributed in West and Central Africa (Mickleburgh et al. 2008b). It is principally associated with lowland rainforest and coastal forest (Monadjem et al. 2010;Fahr 2013e). As this species requires large trees for roosting, it is presumably threatened by deforestation (Mickleburgh et al. 2008b MATERIAL ExAMINED. -1 specimen (see Table 1).
ORIgINAL DATA. -During our field surveys, a single male was captured above a stream in cultivated farm at an altitude of 20 m a.s.l. This specimen was the first recorded from Mount Cameroon area. We did not examine any museum specimens belonging to this species.
DIAgNOSIS. -The measurement of the forearm is 42.4 mm (Table 3), that fits well within the range of Chaerephon major (Happold 2013g, as Tadarida major). A distinct central lappet of skin projects between inner bases of ears. The dorsal and ventral pelage is greyish-brown, with a white central area and whitish flank-stripe ventrally. Other external measurements fit within those provided by Happold (2013g) by Happold (2013g) (Table 3).
HAbITATS AND DISTRIbuTION. -This lowland species is found throughout much of West Africa, as well as along the Nile River from Sudan to Uganda. It has also been recorded in Uganda, Kenya and Tanzania (Happold 2013g as Tadarida major). In Cameroon, the species has been recorded previously in the Sahelian zone of northern Cameroon (Bakwo Fils et al. 2014) The species primarily inhabits savannahs but has also been recorded in rainforest habitats where it roosts in hollow trees and holes in walls of buildings (Happold 2013g MATERIAL ExAMINED. -5 specimens (see Table 1).
ORIgINAL DATA. -5 individuals of the Dwarf free-tailed bat were mist-netted over a slow flowing stream in a fallow farmland, at an altitude 470 m a.s.l. These specimens were the first recorded in the Mount Cameroon area. We did not examine any museum specimens attributed to this species.
DIAgNOSIS. -The forearm measurement of this species is 28.7-30.2 mm (Table 3), which concurs with measurements provided by Happold (2013h) as Tadarida nanula. The dorsal and flank pelage is brown, contrasting with creamy-white chest and belly (Fig. 17). Ears are large with large flap of skin joining the inner surfaces. The wing membranes are whitish, and each jaw possesses two lower incisors. External measurements (Table 3) and skull measurements (Table 4) of one preserved bat are within the range given by Happold (2013h as Tadarida nanula).
HAbITATS AND DISTRIbuTION. -The Dwarf free-tailed bat is widely, but patchily, recorded in West, Central and East Africa from Sierra Leone and Guinea through Cameroon, to western Ethiopia and southward to Democratic Republic of the Congo (Happold 2013h as Tadarida nanula). It is principally associated with lowland closed forest habitat (Happold 1987;Monadjem et al. 2010). It has also been recorded in mesic savannah (Kingdon 1974), where it roosts in small colonies in roofs of thatched huts and tree hollows (Happold 1987). (Thomas, 1903) (   stream at an altitude of 470 m a.s.l. This specimen was the first recorded from the Mount Cameroon area, but this species had previously been recorded from Sahelian zone of northern Cameroon (Bakwo Fils et al. 2014). We did not examine any museum specimens.

Mops (Xiphonycteris) thersites
DIAgNOSIS. -This specimen possesses the characteristic dark colored wings and conspicuous blackish flanks below wings between the leg and arm described by Patterson & Webala (2012) and Happold (2013i as Tadarida thersites). Other features mentioned by Happold (2013i) are also typical such as a forearm length of 35 to 42 mm. The external and skull measurements are provided in Table 3 and Table 4, respectively.
HAbITATS AND DISTRIbuTION. -This species has been widely recorded from much of western and Central Africa (Happold 2013i). It has been recorded mostly from lowland rainforest and secondary forest, but also occurs in mesic savannah and adjacent grassland habitats (Monadjem et al. 2010;Happold 2013i). It roosts in small groups in tree hollows, under roof of houses and cracks of building (Happold 1987;Monadjem et al. 2010 MATERIAL ExAMINED. -11 specimens (see Table 1).
ORIgINAL DATA. -A total of 11 individuals attributed to this species were obtained from a cave at an altitude of 1400 m a.s.l. during our field surveys. DIAgNOSIS.
-The species has a forearm length of 41.0-44.9 mm (Table 3), and possesses features typical of the genus in having a dome-shaped forehead (Fig. 18), and a second phalanx of third finger which is greatly elongated and folds back completely on metacarpals and first phalanx when at rest. The pelage is brownish dorsally and a paler brown ventrally. The interfemoral and wing membranes are brownish. The skull is characteristically large with a greatest length of 13.9-16.5 mm (Table 3), other external measurements are presented in Table 3.
REMARk. -The genus Miniopterus has a number of cryptic species with two listed to occur in Cameroon by   Thomas, 1903 Miniopterus inflatus Thomas, 1903: 634. cOMMON NAME. -English: Greater Long-fingered bat. French: Grand minioptère africain

Miniopterus inflatus
HAbITATS AND DISTRIbuTION. -We did not examine any specimen attributed to this species. This species was also not recorded during our field surveys, but has been signalled in the Mount Cameroon Highlands by Eisentraut (1956Eisentraut ( , 1963 Eger (2013), this species can be encountered in lowland forest, montane forest and farmlands where it roosts in caves. The species had previously been recorded from Mount Cameroon (Eisentraut 1963(Eisentraut , 1973Hill 1968;Fedden & MacLeod 1986), but we did not capture any specimen during our field surveys. Eisentraut (1963) recorded this species in primary montane forest at elevations from 1200 m up to 2260 m a.s.l., where it roosted in large caves.  ORIgINAL DATA. -This species was not netted during our field surveys, but was previously recorded from Mount Cameroon by Eisentraut (1973).

HAbITATS AND DISTRIbuTION. -This species is distributed in Central and
East Africa from Cameroon through Uganda to northwestern Angola. Happold (2013j) indicated that this species inhabits rainforest habitats, miombo woodland and coastal forests where it roosts in vegetation. Thomas, 1913 (Table 3) Glauconycteris egeria Thomas, 1913: 144. cOMMON NAMES. -English: Bibundi Bufferfly Bat. French: Glauconyctère de Bibundi.
ORIgINAL DATA. -A single female individual was captured in a grassland patch at the submontane forest, at an altitude of 1590 m a.s.l. This represented the first record of this species in the Mount Cameroon area. We did not examine any museum specimen attributed to this species.
DIAgNOSIS. -The species is a small microbat with a forearm length of 38.5 mm, and head body length of 43.8 mm ( They have also been recorded to roost between "hands" of bunches of banana fruit, thatch huts, oil palms, roofs of houses and road culverts (Happold 1987). Parahypsugo eisentrauti (Hill, 1968) (Table 3) Pipistrellus eisentrauti Hill, 1968 (Table 1) was caught at the ecotone between montane forest and montane grassland at an altitude of 2130 m a.s.l. This species has previously been referred to as Hypsugo eisentrauti (Hill, 1968) and it was recorded from Mount Cameroon by Eisentraut (1968) as Pipistrellus eisentrauti. The new generic name is based on recent molecular and morphological results by Hutterer et al. (2019). Fedden & MacLeod (1986) mist-netted a total of 22 individuals in the montane forest of Mount Cameroon.
DIAgNOSIS. -The species is very small with a forearm length of 35.4 mm. The pelage is reddish-brown dorsally and slightly paler ventrally. The anterior upper premolar is visible above gum. The wings and interfemoral membrane are also reddish-brown. External measurements are presented in Table 3.
MATERIAL ExAMINED. -We did not examine any specimen attributed to this species.
ORIgINAL DATA. -The tiny pipistrelle was not captured during our field surveys, but has previously been recorded from Mount Cameroon by Fedden & MacLeod (1986), who captured a single male in a primary forest at an altitude of 860 m a.s.l.
HAbITATS AND DISTRIbuTION. -This species is widespread in West and Central Africa, with an isolated population occurring in East Africa. According to Van Cakenberghe & Happold (2013), this species principally inhabits tropical lowland rainforest, but has also been recorded in riverine forest, tropical dry forest and lowland and moist savannah. In Nigeria it is found in the rainforest zone extending northwards into derived and Guinea savannah zones and may be associated with banana plantations (Happold 1987).

DISCUSSION
Our study provides the most recent species list of the bat fauna of Mount Cameroon, with three new species added to the previous record. This emphasizes the importance of carrying out surveys that target different habitat types. However, the surveys conducted by Eisentraut (1963Eisentraut ( , 1964Eisentraut ( , 1968Eisentraut ( & 1973 recorded more species (30) when compared to our surveys (21 species) ( Table 2). This difference can be attributed to several factors, principal among which is the duration of capture which was markedly shorter in our field surveys. Also, the decline in species richness observed during our field surveys may be attributed to anthropogenic activity, which has led to changes in the natural landscape of the region over the past 40 years. Indeed, the natural environments of the Mount Cameroon region have suffered from extensive deforestation, with most of the lowland rainforest being replaced by farmland, agro-industrial plantation and human settlements (Forboseh et al. 2011). All these activities inevitably have a negative impact on the flora and fauna of the region. For instance, in our recent surveys, forest-dependent species such as Hypsignathus monstrosus and Casinycteris ophiodon were not recorded; this could be a sign of a heavily disturbed environment. As pointed out by Cosson et al. (1999), the fragmentation of tropical forest often lead to a rapid decline in the diversity of forest-dependent species in favor of generalist species. The generalists are able to thrive in modified habitats, due to their inherent ability to access additional resources such as food resources and manmade shelters (Coleman & Barclay 2012).
Moreover, there are further indications to support the assertion that deforestation and degradation of the lowland and montane forest of Mount Cameroon may have altered the chiropteran fauna. For example, few highly cluttered space insectivorous bats such as nycterids and vespertilionids were recorded during the recent surveys. Indeed, bats species that prefer cluttered habitat for foraging are more vulnerable to population decline with increase in deforestation due to a decline in resource availability (Threlfall et al. 2012). Additionally, the low number of insectivorous bats captured during our field surveys may be attributed to the use of ground-level mist nets only. As pointed out by Arita (1993), most vespertilionids and molossids are canopy foragers and as such are less frequently captured with ground-level mist nets. Also, insectivorous bats are known to easily escape from mist nets by chewing a hole through the mist nets if the nets are left unwatched even for just a short time (Vaughan 1986). Nevertheless we captured some vespertilionids and molossids mostly as they foraged over slow flowing streams (Table 1).
In order to conserve this exceptional biodiversity, Mount Cameroon National Park (MCNP) was created in 2010. One of the park's main priorities is to protect large mammals that are becoming rare in recent years. We are advocating that bats and other small mammals should also benefit from some of these conservation measures. Among the chiropteran fauna of Mount Cameroon, some species such as Rousettus aegyptiacus, Myonycteris angolensis, Rhinolophus landeri and Hipposideros cf. ruber roost in caves and rock crevices. We recommend that measures should be put in place not only to discourage poaching, but to protect caves against human encroachments. Other species such as Eidolon helvum and Casinycteris ophiodon are red-listed as Near Threatened while Nycteris major, Parahypsugo eisentrauti, Glauconycteris egeria and Doryrhina camerunensis are listed as Data Deficient. These species are particularly in need of dedicated conservation measures to prevent extinction. Thus establishing a checklist of bats will not only provide information on species richness, but can also be an important base for designing future studies and identifying species in need for conservation measures and management consideration.
Indeed, bats provide vital economic and ecological services to humans. Insectivorous bats consume between 50% and 100% of their body mass in insects each night, as such help to regulate nocturnal insect populations, including many agricultural pests (Boyles et al. 2011). Boyles et al. (2011) estimated that bats provide an estimated $3.8 billion per year in North America alone in economic benefits to agriculture as natural control of agricultural pest. Also, Taylor et al. (2018) estimated economic benefits of natural regulation of agricultural pests by bats in South African Macadamia orchids at approximately $613 per hectare, by reducing cost value between 9% and 23% of annual estimated cause damage by stink bugs. Plant-visiting bats typically play a very important role in ecosystem functioning and forest succession after anthropogenic landscape disturbances. Frugivorous bats are excellent seed dispersers that influence the plant species that will potentially recolonize a given region (Henry & Jouard 2007). Nectarivorous and pollinivorous bats pollinate flowers of many species thus influence reproductive success of plants (Quesada et al. 2003). Konaté & Kampmann (2010) noted that almost 100 plant species in Africa are pollinated and or dispersed by bats including trees such as Baobab (Adansonia digitata), African Sausage Tree (Kigelia africana), Silk-cotton Tree (Ceiba pentandra), and the Iroko (Milicia excelsa). Moreover, bats are excellent bioindicators that show measurable responses to environmental stressors such climate change and habitat degradation which reflect the health of the entire ecosystem (Jones et al. 2009).
Our study adds three new species to the Chiropteran fauna of Mount Cameroon, increasing the number of known species to 38. Mount Cameroon now ranks second in terms of species richness on West African mountains after the Upper Guinean Mount Nimba with 59 species (Monadjem et al. 2016). Mount Cameroon is followed by the Simandou Range in Guinea with 35 bat species (Decher et al. 2015), and Mount Mulanje with 30 species (Curran et al. 2012). Our studies conducted at Mount Cameroon were mostly carried out on the seaward aspect of the mountain. We anticipate that further surveys targeting other aspects of the mountain, and using complementary study techniques such as a harp traps and ultrasound detectors will lead to the discovery of additional bat species.