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Family PTEROPODIDAE Gray, 1821 REMARK Nine species of Old World fruit bats (Pteropodidae) were encountered during the two field seasons. The maximum number of pteropodid species encountered at one site was five during a single field season (dry) and seven during both field seasons in the Agumatsa valley (Wli). The latter number is identical to what Yeboah (2001) found at Mt. Afadjato Forest (near Tagbo Falls)., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on page 266, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["YEBOAH S. 2001. - The distribution and relative abundance of megachiropteran bats in Southern Ghana. Journal of the Ghana Science Association 3: 99 - 105."]}

Crocidura olivieri (Lesson, 1827) Sorex olivieri Lesson, 1827: 121. COMMON NAME. — African Giant White-toothed Shrew, Olivier’s Shrew. MATERIAL EXAMINED. — Apesokubi • 1 ♀; Sherman trap in forest; ZFMK 2003.1089. Liati Wote (Tagbo Falls) • 1?; Sherman trap between buttress roots in forest; ZFMK 2003.1090. REMARK This species was previously known from several localities in the Volta Region (Grubb et al. 1998). In their Kyabobo expedition report Hurst et al. (1995) mention several individuals of this large shrew (as C. flavescens) from KRNP, but we did not encounter it there. Our specimen from Liati Wote (ZFMK 2003.1090) was included in a recent phylogeographic study of C. olivieri (Jacquet et al. 2015) and grouped with their Clade IIB with Dahomey Gap specimens from Togo and Benin. CONSERVATION STATUS. — Crocidura olivieri is listed as “Least Concern” on the IUCN Red List., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on page 260, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["LESSON R. P. 1827. - Manuel de mammalogie, ou histoire naturelle des mammiferos. J. B Bailliere, Pairs, 442 p. https: // doi. org / 10.5962 / bhl. title. 137811","GRUBB P., JONES T. S., DAVIES A. G., EDBERG E., STARIN E. D. & HILL J. E. 1998. - Mammals of Ghana, Sierra Leone and the Gambia. The Trendrine Press, Zennor, St. Ives, Cornwall, vi + 265 p.","HURST J., BOWES LYON D. & KEEFE E. O' 1995. - Kyabobo' 94: a joint Ghanaian and Oxford University Expedition. Final Report. KCP Publications, Ghana Wildlife Department, Institute for Renewable Natural Resources, 69 p.","JACQUET F., DENYS C., VERHEYEN E., BRYJA J., HUTTERER R., KERBIS PETERHANS J. C., STANLEY W. T., GOODMAN S. M., COULOUX A., COLYN M. C. & NICOLAS V. 2015. - Phylogeography and evolutionary history of the Crocidura olivieri complex (Mammalia, Soricomorpha): from a forest origin to broad ecological expansion across Africa. BMC Evolutionary Biology 15: 71, https: // doi. org / 10.1186 / s 12862 - 015 - 0344 - y"]}

Micropteropus pusillus (Peters, 1868) Epomophorus pusillus Peters, 1868: 870. COMMON NAME. — Peters’ Lesser Epauletted Fruit Bat. MATERIAL EXAMINED. — Agumatsa Wildlife Sanctuary • 3 ♀; ZTNHC 957, SMF 92112, 92113. Apesokubi • 1 ♂; 0.5 km N Apesokubi; USNM 590071 • 4♀; USNM 590070, ZTNHC 966, SMF 89660, 89661. Kyabobo NP • 1 ♂; SMF 92114. Shiare Schoolyard • 1 ♀; SMF 92115. REMARK As in previous studies from Ghana and Togo (De Vree et al. 1969, 1971; Decher 1997a), with 23 individuals caught, Micropteropus pusillus was the most common small fruit bat in our survey, perhaps outnumbered locally only by large colonies of the high-flying Eidolon helvum (Kerr, 1792). It is a typical species of the rainforest-savannah mosaic characteristic for much of the Volta Region. In Ghana east of the Volta River, M. pusillus was previously captured at Akwamufe, Amedzofe, Kalakpa Game Production Reserve, Leklebi Agbesia and Odomi Jongo (Bergmans 1989). In Ghana and Togo, M. pusillus shares a similar distribution pattern with Epomophorus gambianus (see maps in Bergmans [1988, 1989] and Grubb et al. [1998]) but it seems to occur more commonly in or near forest remnants than the latter species, suggesting a greater dependence on forest. In our survey up to eight individuals would hit the nets at the same time indicating that they fly and forage in groups. CONSERVATION STATUS. — Micropteropus pusillus was classified as “Least Concern” on the IUCN Red List. It remains a common fruit bat species in the Ghana-Togo Highlands. However, this species may depend on forest remnants during its foraging flights (Fahr 1996)., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on page 267, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["PETERS W. 1868. - Fortsetzung und der Schluss einer Ubersicht der Flederhunde. Monatsberichte der Koniglich-Preussischen Akademie der Wissenschaften zu Berlin 1868: 865 - 872.","DE VREE F., DE ROO A. & VERHEYEN W. N. 1969. - Contribution a l'etude des chiropteres de la Republique du Togo. Revue de Zoologie et de Botanique Africaines 83: 200 - 207.","DE VREE F. & VAN DER STRAETEN E. 1971. - Contribution a l'etude des chiropteres de la Republique duTogo. 3. Liste preliminaire des Chiropteres recoltes par la troisieme Mission zoologique belge au Togo. R evue de Zoologie et de Botanique Africaines 83: 159 - 164.","DECHER J. 1997 a. - Bat community patterns on the Accra Plains of Ghana, West Africa. Zeitschrift fur Saugetierkunde 62: 129 - 142.","KERR R. 1792. - The animal kingdom, or zoological system, of the celebratrd Sir Charles Linnaeus; class I: Mammalia. J. Murray & R. Faulder, London, 644 p. https: // doi. org / 10.5962 / bhl. title. 57940","BERGMANS W. 1989. - Taxonomy and biogeography of African fruit bats (Mammalia, Megachiroptera). 2. The genera Micropterus Matschie, 1899, Epomops Gray, 1870, Hypsignathus H. Allen, 1861, Nanonycteris Matschie, 1899, and Plerotes Andersen, 1910. Beaufortia 39: 89 - 153.","BERGMANS W. 1988. - Taxonomy and biogeography of African fruit bats (Mammalia, Megachiroptera). 1. General Introduction; material and methods; results: The genus Epomophorus Bennett, 1836. Beaufortia 38: 75 - 146.","GRUBB P., JONES T. S., DAVIES A. G., EDBERG E., STARIN E. D. & HILL J. E. 1998. - Mammals of Ghana, Sierra Leone and the Gambia. The Trendrine Press, Zennor, St. Ives, Cornwall, vi + 265 p.","FAHR J. 1996. - Die Chiroptera der Elfenbeinkuste (unter Berucksichtigung des westafrikanischen Raumes): Taxonomie, Habitatpraferenzen und Lebensgemeinschaften. Diplomarbeit, Julius-Maximilians-Universitat, Wurzburg. 204 p."]}

Macronycteris gigas (Wagner, 1845) Rhinolophus gigas Wagner, 1845: 148. COMMON NAME. — Giant Leaf-nosed Bat. MATERIAL EXAMINED. — Agumatsa Wildlife Sanctuary • 1 ♀; SMF 92135; 11.VIII.2001. REMARK This is the second largest insectivorous bat in West Africa, exceeded in size only by the emballonurid Saccolaimus peli (Temminck, 1853). We follow Foley et al. (2017) in the use of the name Macronycteris Gray, 1866 instead of Hipposideros Gray, 1831. This Macronyteris female was identified as M. gigas by the late D. Kock (in litt.) based on its large forearm measurement of 102 mm. The mean for ♀ M. vittatus (Peters, 1852) is 93.9 mm with a Range of 84-101mm (Happold 2013b). Only one individual of this large hipposiderid bat was collected during the rainy season. Four other Volta Region specimens are known from Odomi Jongo, 2 miles E Nkwanta (USNM 424858-61). Togo highlands specimens are known from Adjido, Agou, and Ezimé (De Vree et al. 1969; Robbins 1980). Our record confirms previous observations (Grubb et al. 1998) that in Ghana this species often occurs in forests, although it is also found in Guinea woodland and even in the coastal savanna like the Accra Plains (Decher 1997a). In Côte d’Ivoire, Fahr (1996) made six of his eight captures in the rain forest zone. Six ectoparasitic Ascodipteron variisetosum Maa, 1965 [Diptera: Streblidae] with (2 × 3) were found on the ventral side of the upper arm on the specimen from Agumatsa. CONSERVATION STATUS. — Listed as “Least Concern” on the IUCN Red List, this species is of some conservation concern in the Ghana- Togo Highlands because it appears to be patchily distributed and may be limited not so much by the availability of large tracts of forest as by the presence of large hollow trees and caves for its roosts (Grubb et al. 1998)., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on page 271, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["WAGNER A. 1845. - Diagnosen einiger neuer Arten von Nagern und Handfluglern. Archiv fur Naturgeschichte Jahrs 11, Bd. 1: 145 - 149.","THOMAS O. 1904. - On mammals from Northern Angola collected by Dr. W. J. Ansorge. The Annals and Magazine of Natural History (ser. 7) 13: 405 - 421. https: // doi. org / 10.1080 / 00222930408562472","TEMMINCK C. J. 1853. - Esquisses zoologiques sur la Cote de Guine, Part I, Mammiferes. E. J. Brill, Leiden, 256 p. https: // www. biodiversitylibrary. org / page / 14835765","FOLEY N. M., GOODMAN S. M., WHELAN C. V., PUECHMAILLE S. J. & TEELING E. 2017. - Towards navigating the Minotaur's Labyrinth: Cryptic diversity and taxonomic revision within the speciose genus Hipposideros (Hipposideridae). Acta Chiropterologica 19: 1 - 18. https: // doi. org / 10.3161 / 15081109 ACC 2017.19.1.001","HAPPOLD D. C. D. 2013 b. - Hipposideros vittatus Striped Leaf-nosed Bat, in HAPPOLD M. & HAPPOLD D. C. D. (eds), The Mammals of Africa - Vol. IV: Hedgehogs, Shrews and Bats. Bloomsbury Publishing, London: 395 - 398.","DE VREE F., DE ROO A. & VERHEYEN W. N. 1969. - Contribution a l'etude des chiropteres de la Republique du Togo. Revue de Zoologie et de Botanique Africaines 83: 200 - 207.","ROBBINS C. B. 1980. - Small mammals of Togo and Benin. I. Chiroptera. Mammalia 44: 83 - 88.","GRUBB P., JONES T. S., DAVIES A. G., EDBERG E., STARIN E. D. & HILL J. E. 1998. - Mammals of Ghana, Sierra Leone and the Gambia. The Trendrine Press, Zennor, St. Ives, Cornwall, vi + 265 p.","DECHER J. 1997 a. - Bat community patterns on the Accra Plains of Ghana, West Africa. Zeitschrift fur Saugetierkunde 62: 129 - 142.","FAHR J. 1996. - Die Chiroptera der Elfenbeinkuste (unter Berucksichtigung des westafrikanischen Raumes): Taxonomie, Habitatpraferenzen und Lebensgemeinschaften. Diplomarbeit, Julius-Maximilians-Universitat, Wurzburg. 204 p."]}

Nanonycteris veldkampi (Jentink, 1888) Epomophorus veldkampii Jentink, 1888a: 51. COMMON NAME. ��� Veldkamp���s Bat. MATERIAL EXAMINED. ��� Agumatsa Wildlife Sanctuary ��� 1 ♀ with 1 embryo; SMF 89665. Apesokubi ��� 1♀ with 1 embryo; SMF 89664. REMARK Previous records of this species were from Odomi Jongo in the Volta Region (USNM) and from Bismarckburg, Misahoh��, Aledjo, Atakpam��, Evou, Ezim��, Fazao, Odjolo, Plateau Akpossi in Togo (De Vree et al. 1969, 1970; De Vree & Van der Straeten 1971). This is a small fruit bat similar in appearance to Micropteropus pusillus from which it can be distinguished by its slender snout and the nine undivided and thinner palatal ridges (see drawings in Bergmans [1997] and Happold [1987]). With just two specimens captured at the beginning of dry season in 1999, N. veldkampi was much less common than M. pusillus (23 specimens; Table 2). The seasonal occurrence could be explained by the migratory behavior of this species, which was shown to follow the progression of the rains northward to savanna areas (Fahr 1996, Thomas 1983). Similarly, at Mount Nimba, Wolton et al. (1982) did not obtain this species at all between early July and early September, whereas it was common there at other times of the year (Monadjem et al. 2016). During the African Small Mammal Project, between January and June 1968 (Robbins 1980), no N. veldkampi were encountered, yet the same Project captured 20 M. pusillus in Togo and Benin. During a study on the Accra Plains between November 1991 and June 1992 no N. veldkampi were encountered, but 45 M. pusillus were captured (Decher 1997a). CONSERVATION STATUS. ��� Nanonyteris veldkampi is classified as ���Least Concern��� on the IUCN Red List. Its migratory behavior and dependence on forest remnants still need to be investigated in more detail in the Ghana-Togo Highlands., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on pages 268-269, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["JENTINK F. A. 1888 a. - Catalogue systematique des Mammiferes. Museum d'Histoire naturelle des Pays-Bas. Leiden 12: 1 - 280. https: // repository. naturalis. nl / pub / 508279","DE VREE F., DE ROO A. & VERHEYEN W. N. 1969. - Contribution a l'etude des chiropteres de la Republique du Togo. Revue de Zoologie et de Botanique Africaines 83: 200 - 207.","DE VREE F., HULSELMANS J. & VERHEYEN W. 1970. - Contribution a l'etude des chiropteres de la Republique du Togo. 2. Liste preliminaire des Chiropteres. Revue de Zoologie et de Botanique Africaines 82: 41 - 46.","DE VREE F. & VAN DER STRAETEN E. 1971. - Contribution a l'etude des chiropteres de la Republique duTogo. 3. Liste preliminaire des Chiropteres recoltes par la troisieme Mission zoologique belge au Togo. R evue de Zoologie et de Botanique Africaines 83: 159 - 164.","BERGMANS W. 1997. - Taxonomy and biogeography of African fruit bats (Mammalia, Megachiroptera). 5. The genera Lissonycteris Andersen, 1912, Myonycteris Matschie, 1899 and Megaloglossus Pagenstecher, 1885; general remarks and conclusions; annex; key to all species. Beaufortia 47: 11 - 90.","HAPPOLD D. C. D. 1987. - The Mammals of Nigeria. Clarendon Press, Oxford, xvii + 402 p.","FAHR J. 1996. - Die Chiroptera der Elfenbeinkuste (unter Berucksichtigung des westafrikanischen Raumes): Taxonomie, Habitatpraferenzen und Lebensgemeinschaften. Diplomarbeit, Julius-Maximilians-Universitat, Wurzburg. 204 p.","THOMAS D. W. 1983. - The annual migrations of three species of West African fruit bats (Chiroptera: Pteropodidae). Canadian Journal of Zoology 61: 2266 - 2272.","WOLTON R. J., PRAK P. A., GODFRAY H. C. J. & WILSON R. P. 1982. - Ecological and behavioral studies of the Megachiroptera at Mount Nimba, Liberia. Mammalia 46: 419 - 448.","MONADJEM A., RICHARDS L. & DENYS C. 2016. - An African bat hotspot: the exceptional importance of Mount Nimba for bat diversity. Acta Chiropterologica 18: 359 - 375. https: // doi. org / 10. 3161 / 15081109 ACC 2016.18.2.005","ROBBINS C. B. 1980. - Small mammals of Togo and Benin. I. Chiroptera. Mammalia 44: 83 - 88.","DECHER J. 1997 a. - Bat community patterns on the Accra Plains of Ghana, West Africa. Zeitschrift fur Saugetierkunde 62: 129 - 142."]}

Hylomyscus pamfi Nicolas, Olayemi, Wendelen & Colyn, 2010 (Fig. 6) Hylomyscus pamfi Nicolas, Olayemi, Wendelen & Colyn, 2010b: 38. COMMON NAME. — Dahomey Gap Woodmouse. MATERIAL EXAMINED. — Agumatsa Wildlife Sanctuary • 1 ♀; SMF 91364; 4.5 km N. Apesokubi (Fig. 6)• 1 ♂; ZTNHC 965 • 2♀; SMF 91434, USNM 590074. Kyabobo National Park • 1 ♂; SMF 91371 • 1 ♀; SMF 91370. Kalakpa Resource Reserve • 1 ♂; USNM 590073. REMARK We follow Nicolas et al. (2010 b, 2020), Monadjem et al. (2015) and Denys et al. (2017) in assigning all Volta Region Hylomyscus Thomas, 1926 to the Dahomey Gap Woodmouse, H. pamfi, which we cross-checked by examining cytochrome b (Cytb) sequences for six of our seven specimens. Three Cytb haplotypes MT311308 - MT311310 were detected at the Apesokubi site, one at Kyabobo (MT311311) and one at Kalakpa (MT311307). These Cytb haplotypes showed a 100-95% identity to sequences reported from H. pamfi by Nicolas et al. (2020) from Benin. The nearest locality in Nicolas et al. (2010b) to our Volta Region study sites was Palimé in Togo, which is about 23 km from Wli Waterfall. Our study represents the first verification that H. pamfi is present in Ghana. There are four additional Hylomyscus specimens from the Volta Region at the USNM (listed as H. alleni (Waterhouse, 1838)), collected in 1968 by the African Mammal Project at Leklebi Agbesia. Hylomyscus pamfi is a forest species with good climbing ability. At least four of our specimens were captured on traps set on horizontal branches above ground, one at Apesokubi in an Umbrella tree (Musanga cecropioides R.Br.). CONSERVATION STATUS. — Hylomyscus pamfi is currently listed as “Data Deficient” on the IUCN Red List.

Order CHIROPTERA Blumenbach, 1779 REMARK Seventeen bat species were recorded from the Ghana-Togo Highlands based on our two expeditions in the Volta Region (19 nights netted). We list all species and their localities encountered in their zoogeographic context., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on page 266, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781

Crocidura cf. foxi Dollman, 1915 (Fig. 3) Crocidura foxi Dollman, 1915: 514. COMMON NAME. — Fox’s White-toothed Shrew. MATERIAL EXAMINED. — Apesokubi • 1 ♀; 26.XI.1999; ZFMK 1999.1145 • 1?; Sherman traps; 27.XI.1999; ZFMK 1999.1146. REMARK Known initially just from Nigeria C. foxi is now known to occur in the Guinea savanna zone and secondary bushland from Senegal to Sudan (Duplantier & Granjon 2013). Our captures are the first from the Volta Region of Ghana, listed as Crocidura cf. foxi, as the identification has not yet been confirmed by molecular genetics. An earlier capture from KRNP, reported by Hurst et al. (1995) as C. foxii [sic], was later re-identified as C. fuscomurina (Heuglin, 1865) (Hutterer 1983; Grubb et al. 1998). CONSERVATION STATUS. — Crocidura foxi is listed as “Least Concern” on the IUCN Red List., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on page 261, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["DOLLMAN G. 1915. - On the African shrews belonging to the genus Crocidura. Journal and Magazine of Natural History (ser. 8) 15: 507 - 527. https: // www. biodiversitylibrary. org / page / 24256132","WATERHOUSE G. R. 1840. - On a new Genus of the Family Muridae and Order Rodentia. Proceeding of the Zoological Society of London 8: 1 - 3.","DUPLANTIER J. - M. & GRANJON L. 2013. - Crocidura foxi Fox's Shrew, in HAPPOLD M. & HAPPOLD D. C. D. (eds), The Mammals of Africa. Volume IV. Hedgehogs, shrews and bats. Bloomsbury Publishing, London: 78 - 79.","HURST J., BOWES LYON D. & KEEFE E. O' 1995. - Kyabobo' 94: a joint Ghanaian and Oxford University Expedition. Final Report. KCP Publications, Ghana Wildlife Department, Institute for Renewable Natural Resources, 69 p.","GRUBB P., JONES T. S., DAVIES A. G., EDBERG E., STARIN E. D. & HILL J. E. 1998. - Mammals of Ghana, Sierra Leone and the Gambia. The Trendrine Press, Zennor, St. Ives, Cornwall, vi + 265 p."]}

Family SORICIDAE G. Fischer, 1814 REMARK Only five shrews belonging to three species were captured during both expeditions (Table 2). However, according to Grubb et al. (1998) only one of these species has previously been recorded from the Volta Region., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on page 260, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["GRUBB P., JONES T. S., DAVIES A. G., EDBERG E., STARIN E. D. & HILL J. E. 1998. - Mammals of Ghana, Sierra Leone and the Gambia. The Trendrine Press, Zennor, St. Ives, Cornwall, vi + 265 p."]}

Graphiurus nagtglasii Jentink, 1888 (Fig. 8) Graphiurus nagtglasii Jentink, 1888b: 38. COMMON NAME. — Nagtglas’s Dormouse; EWE: Kade. MATERIAL EXAMINED. — Apesokubi (Fig. 8) • 1 ♂; USNM 590101; 26.XI.1999; Sherman trap on a horizontal branch at 1.5 m height • 2 ♀; USNM 590102; ZTNHC 972; 27.XI.1999; Sherman and Victor Rat traps. REMARK All three specimens were caught in this dense traditionally protected forest at Apesokubi. There are 22 specimens from Leklebi Agbesia and one specimen from Odomi Jongo (2 miles E Nkwanta), all in the Volta Region in the USNM. Grubb et al. (1998) show two other localities in the Volta Region. The species has also been reported from several localities in the Togo Highlands (Roche 1971; Robbins & Van der Straeten 1996). CONSERVATION STATUS. — Graphiurus nagtglasii is listed as “Least Concern” on the IUCN Red List, however, progressive forest loss will marginalize this arboreal species., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on page 265, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["JENTINK F. A. 1888 b. - Zoological researches in Liberia. A list of mammals, collected by J. Buttikofer, C. F. Sala and F. X. Stampfli, with biological observations. Notes from the Leyden Museum 10: 38 - 41. https: // www. biodiversitylibrary. org / page / 9622343","GRUBB P., JONES T. S., DAVIES A. G., EDBERG E., STARIN E. D. & HILL J. E. 1998. - Mammals of Ghana, Sierra Leone and the Gambia. The Trendrine Press, Zennor, St. Ives, Cornwall, vi + 265 p.","ROCHE J. 1971. - Recherches mammalogiques en Guinee forestiere. Bulletin du Museum national d'Histoire naturelle, 3 e serie 16: 737 - 781.","ROBBINS C. B. & VAN DER STRAETEN E. 1996. - Small mammals of Togo and Benin. II. Rodentia. Mammalia 60: 231 - 242."]}

Epomophorus gambianus (Ogilby, 1835) Pteropus gambianus Ogilby, 1835: 100. COMMON NAME. — Gambian Epauletted Fruit Bat; EWE: Aguto. MATERIAL EXAMINED. — Kalakpa Resource Reserve • 1 ♂; ZTNHC 971; 16.XI.1999. REMARK Only one specimen of this savanna species was captured at a thicket edge in Kalakpa Resource Reserve, a mostly savannacovered legally-protected area in the south of the Volta Region, southeast of the Abutia Hills. Epomophorus gambianus is well documented from Eastern Ghana and Togo and is most common in savanna or secondary savanna areas. During three Belgian expeditions to Togo, including many localities in the Togo Highlands, it was the second most abundant fruit bat after Micropteropus pusillus (Peters, 1868) (DeVree et al. 1969; 1970; De Vree & Van der Straeten 1971). In the Volta Region E. gambianus was previously recorded from Bator, Ho, Keta, Kete Krachi (= Kradji), and Odomi Jongo, 2 miles E Nkwanta (Bergmans 1988). CONSERVATION STATUS. — Epomophorus gambianus is listed as “Least Concern” on the IUCN Red List. Although rare during our survey, the high abundance of this fruit bat in surveys that included savanna areas make it a species of less concern., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on page 266, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["OGILBY W. 1835. - Descriptions of Mammalia and birds from the Gambia. Proceedings of the Society of London 3: 97 - 105.","PETERS W. 1868. - Fortsetzung und der Schluss einer Ubersicht der Flederhunde. Monatsberichte der Koniglich-Preussischen Akademie der Wissenschaften zu Berlin 1868: 865 - 872.","DE VREE F. & VAN DER STRAETEN E. 1971. - Contribution a l'etude des chiropteres de la Republique duTogo. 3. Liste preliminaire des Chiropteres recoltes par la troisieme Mission zoologique belge au Togo. R evue de Zoologie et de Botanique Africaines 83: 159 - 164.","BERGMANS W. 1988. - Taxonomy and biogeography of African fruit bats (Mammalia, Megachiroptera). 1. General Introduction; material and methods; results: The genus Epomophorus Bennett, 1836. Beaufortia 38: 75 - 146."]}

Order RODENTIA Bowditch, 1821 REMARK A total of 238 individuals from 12 rodent species were verified by us during both expeditions to the Volta Region (Table 2)., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on page 261, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781

Megaloglossus azagnyi Nesi, Kadjo & Hassanin, 2013 (Fig. 9) Megaloglossus azagnyi Nesi, Kadjo & Hassanin in Nesi et al. 2013: 134. COMMON NAME. — Western Woermann’s Fruit Bat. MATERIAL EXAMINED. — Agumatsa Wildlife Sanctuary • 2♂; SMF 92117 + 92118; 11.IX.2001. Shiare schoolyard (Fig. 9) • 1 ♀; SMF 92119; 22.VIII.2001 • 1 ♂; SMF 92120; 22.VIII.2001. REMARK We follow Nesi et al. (2013) in the use of the name M. azagnyi as a species separate from M. woermanni Pagenstecher, 1885 for Upper Guinea and the Volta Region of Ghana, which we confirmed with BLAST searches of Cytb sequences for our four specimens. While all four Cytb sequences were of sufficient quality to allow species identification; only two sequences, one from each locality, were of sufficient quality to be entered into Genbank (MT311312 from SMF 92117 and MT311313 from SMF 92120. These sequences had a 100-96% sequence identities to sequences reported from M.azagnyi from the Cote d’Ivoire by Nesi et al. (2013). The sequences obtained from specimens SMF 92118 and 92119 contained numerous “no calls” or “N” resulting from the detection on more than a single nucleotide at a site. Previous records of Megaloglossus Pagenstecher 1885 in the Volta Region were from Amedzofe and Odomi Jongo, 2 miles E Nkwanta (Bergmans 1997). CONSERVATION STATUS. — The IUCN Red List map shows the distribution to include the Ghana-Togo Highlands (Monadjem 2016) but how far this nectar-feeding species extends into the Dahomey Gap before it is replaced by M. woermanni in Lower Guinea is still unknown. The IUCN Red List classification is “Least Concern”., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on page 268, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["NESI N., KADJO B., POURRUT X., LEROY E., PONGOMBO SHONGO C., CRUAUD C. & HASSANIN A. 2013. - Molecular systematics and phylogeography of the tribe Myonycterini (Mammalia, Pteropodidae) inferred from mitochondrial and nuclear markers. Molecular Phylogenetics and Evolution 66: 126 - 137. https: // doi. org / 10.1016 / j. ympev. 2012.09.028","BERGMANS W. 1997. - Taxonomy and biogeography of African fruit bats (Mammalia, Megachiroptera). 5. The genera Lissonycteris Andersen, 1912, Myonycteris Matschie, 1899 and Megaloglossus Pagenstecher, 1885; general remarks and conclusions; annex; key to all species. Beaufortia 47: 11 - 90."]}

Anomalurus beecrofti Fraser, 1852 Anomalurus beecrofti Fraser, 1852: 17. COMMON NAME. — Beecroft’s Anomalure; EWE: Kasanui, Vava. MATERIAL EXAMINED. — Agumatsa Wildlife Sanctuary • 1 ♀; ZTNHC 950. REMARK This specimen was brought to us by a local hunter, on 19 November 1999. In the Volta Region a previous record is from the vicinity of Jasikan (Grubb et al. 1998) and it was observed at Kyabobo National Park by Hurst et al. (1995). In the Togo Highlands A. beecrofti was previously recorded from Apéyémé, Edifou, Ezimé and Misahohé (Robbins & Van der Straeten 1996; USNM). Grubb et al. (1998) followed Rosevear (1969) in using the generic name Anomalurops Matschie, 1914 for this species; we follow Dieterlen (2005) and Kingdon (2013) in the use of Anomalurus Waterhouse, 1843. CONSERVATION STATUS. — Anomalurus beecrofti is listed as “Least Concern” on the IUCN Red List, however, progressive forest fragmentation with the loss of mature trees, specifically oil palms (Elaeis guineensis), will greatly compromise the habitat of this treedependent glider., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on page 265, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["FRASER L. 1852. - Description of a new species on Anomalurus from Fernando Po. Proceedings of the Zoological Society of London 20: 16 - 17. https: // www. biodiversitylibrary. org / page / 30680608","GRUBB P., JONES T. S., DAVIES A. G., EDBERG E., STARIN E. D. & HILL J. E. 1998. - Mammals of Ghana, Sierra Leone and the Gambia. The Trendrine Press, Zennor, St. Ives, Cornwall, vi + 265 p.","HURST J., BOWES LYON D. & KEEFE E. O' 1995. - Kyabobo' 94: a joint Ghanaian and Oxford University Expedition. Final Report. KCP Publications, Ghana Wildlife Department, Institute for Renewable Natural Resources, 69 p.","ROBBINS C. B. & VAN DER STRAETEN E. 1996. - Small mammals of Togo and Benin. II. Rodentia. Mammalia 60: 231 - 242.","ROSEVEAR D. R. 1969. - The rodents of West Africa. Trustees of the British Museum of Natural History, London, xii + 604 p.","DIETERLEN F. 2005. - Family Anomaluridae in WILSON D. E. & REEDEr D. M. (eds) Mammal species of the world. Johns Hopkins University Press, Baltimore: 1532 - 1534."]}

Epomops buettikoferi (Matschie, 1899) Epomophorus büttikoferi Matschie, 1899: 45. COMMON NAME. — Büttikofer’s Epauletted Fruit Bat; EWE: Aguto. MATERIAL EXAMINED. — Apesokubi • 1 adult ♀ with a 53 mm long embryo; 27.VIII.2001; SMF 92116. REMARK Essentially an Upper Guinea forest species with some disjunct populations east of the Dahomey Gap in Nigeria (Bergmans 1989; Thomas & Henry 2013). All previous Ghanaian specimens were from the rainforest zone in SW Ghana west of the Volta River (Bergmans 1989; Grubb et al. 1998). Thus, our specimen is the first record east of the Volta in Ghana. CONSERVATION STATUS. — Epomops buettikoferi is listed as “Least Concern” on the IUCN Red List., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on page 266, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["MATSCHIE P. 1899. - Die Megachiroptera des Berliner Museum fur Naturkunde. G. Reimer, Berlin. https: // doi. org / 10.5962 / bhl. title. 48691","BERGMANS W. 1989. - Taxonomy and biogeography of African fruit bats (Mammalia, Megachiroptera). 2. The genera Micropterus Matschie, 1899, Epomops Gray, 1870, Hypsignathus H. Allen, 1861, Nanonycteris Matschie, 1899, and Plerotes Andersen, 1910. Beaufortia 39: 89 - 153.","THOMAS D. & HENRY M. 2013. - Epomops buettikoferi Buettikofer's Epauletted Fruit Bat in HAPPOLD M. & HAPPOLD D. C. D. (eds) The Mammals of Africa - Vol. IV: Hedgehogs, Shrews and Bats. Bloomsbury Publishing, London: 253 - 255.","GRUBB P., JONES T. S., DAVIES A. G., EDBERG E., STARIN E. D. & HILL J. E. 1998. - Mammals of Ghana, Sierra Leone and the Gambia. The Trendrine Press, Zennor, St. Ives, Cornwall, vi + 265 p."]}

Praomys misonnei Van der Straeten & Dieterlen, 1987 (Fig. 5) Praomys misonnei Van der Straeten & Dieterlen, 1987: 3. COMMON NAME. — Misonne’s Soft-furred Rat. MATERIAL EXAMINED. — Agumatsa Wildlife Sanctuary • 5 ♂; USNM 590095-590098, SMF 91365 • 12 ♀; USNM 590093, 590094, USNM 590115, 590116, SMF 91435, 91363, 91366, 91367-91369, 93747, ZTNHC 953 • 1?; USNM 590117. Apesokubi • 7 ♂; USNM 590099, SMF 91374 + 91377, ZTNHC 961, 962, 964, UVM VR062 • 5 ♀; USNM 590100, USNM 590118 – 590120, SMF 91375. Kalakpa Resource Reserve (see Fig. 5) • 8 ♂; USNM 590080 + 590081, USNM 590083-590085, USNM 590088, USNM 590091, USNM 590113, ZTNHC 949 • 6 ♀; USNM 590082, USNM 590086 + 590087, USNM 590089 + 590090, USNM 590114 • 1?; USNM 590079. Kyabobo National Park • 3 ♀; USNM 590121-590123. Shiare • 1 ♀; USNM 590124 • 2♀; SMF 91372 + 91373. Tagbo-Falls • 1 ♂; USNM 590103; 1 ♀; USNM 590125. REMARK With 190 captured individuals, Praomys misonnei was the most frequently caught species in the Volta Region, followed by Mus (Nannomys) musculoides Temminck, 1853 (n = 23) and Hylomyscus pamfi (n = 7). All Praomys had been tentatively identified as P. tullbergi in the field, but a recent molecular follow-up study included seven of our specimens from Agumatsa, Apesokubi and Shiare and showed that they all belonged to P. misonnei, reinforcing that P. tullbergi is limited to the Upper Guinea Region, west of the Volta River and P. misonnei to the Dahomey Gap region into Central Africa (Nicolas et al. 2010a, 2011) with the Volta River forming the zoogeographic barrier between the two species. Our specimens were part of their Clade II, which is restricted to the Ghana-Togo Highlands and Dahomey Gap area. CONSERVATION STATUS. — Praomys misonnei is listed as “Least Concern” on the 2019 IUCN Red List, but the species range has not yet been updated to include the Lower Guinea Region based on Nicolas et al. (2010a, 2011)., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on page 262, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["VAN DER STRAETEN E. & DIETERLEN F. 1987. - Praomys misonnei, a new species of Muridae from Eastern Zaire (Mammalia). Stuttgarter Beitrage zur Naturkunde, Serie A (Biologie) 402: 1 - 11. https: // www. biodiversitylibrary. org / page / 33622925","TEMMINCK C. J. 1853. - Esquisses zoologiques sur la Cote de Guine, Part I, Mammiferes. E. J. Brill, Leiden, 256 p. https: // www. biodiversitylibrary. org / page / 14835765","NICOLAS V., AKPATOU B., WENDELEN W. I. M., KERBIS PETER- HANS J., OLAYEMI A., DECHER J., MISSOUP A. - D., DENYS C., BARRIERE P., CRUAUD C. & COLYN M. 2010 a. - Molecular and morphometric variation in two sibling species of the genus Praomys (Rodentia: Muridae): implications for biogeography. Zoological Journal of the Linnean Society 160: 397 - 419. https: // doi. org / 10.1111 / j. 1096 - 3642.2009.00602. x","NICOLAS V., MISSOUP A. D., DENYS C., KERBIS PETERHANS J., KATU- ALA P., COULOUX A. & COLYN M. 2011. - The roles of rivers and Pleistocene refugia in shaping genetic diversity in Praomys misonnei in tropical Africa. Journal of Biogeography 38: 191 - 207. https: // doi. org / 10.1111 / j. 1365 - 2699.2010.02399. x"]}

Scotophilus livingstonii Brooks & Bickham, 2014 Scotophilus livingstonii Brooks & Bickham, 2014: 11. COMMON NAME. — Livingstone’s House Bat. MATERIAL EXAMINED. — Agumatsa (Wli Waterfall) • 1 ♀; SMF 92137; caught on 11.VIII.2001. REMARK Originally identified as Scotophilus dinganii (A. Smith, 1833) due to its bright yellow ventral side side and its forearm length of 57.8 mm, the West African form, which also occurs in western Kenya, has been recently renamed S. livingstonii based on phylogenetic and morphological distinct S. dinganii -like clades (Brooks & Bickham 2014). Our specimen was caught in a net set across the Agumatsa River flanked by secondary forest and nearby small cassava fields. Other Ghana-Togo Highlands specimens are known from Odomi Jongo, 12 Miles E Nkwanta in Ghana (USNM 424888) and from Atakpamé and Ezimé in Togo (Robbins 1980; Robbins et al. 1985). The species was also caught at three locations on the Accra Plains (Decher 1997a and USNM, as S. dinganii) and seems to be most commonly associated with Guinea savanna, forest savanna mosaic, and high forest edge. Records from Côte d’Ivoire are all from the northern tree savanna (Fahr 1996, as S. dinganii). This species also seems to adapt to rooftops and thatched huts for its roosts. CONSERVATION STATUS. — The conservation classification of S. livingstonii on the IUCN Red List is “Least Concern”.

Rhinolophus alcyone Temminck, 1853 (Fig. 10) Rhinolophus alcyone Temminck, 1853: 80. COMMON NAME. — Halcyon Horseshoe Bat. MATERIAL EXAMINED. — Kalakpa Resource Reserve • 1 ♀; SMF 89668. Agumatsa Wildlife Sanctuary • 4 ♂; ZTNHC 952 +958, SMF 89669 +92129 • 2 ♀; SMF 92127 +92128. Apesokubi • 1 ♀; SMF 92130. REMARK Larger than R. landeri Martin, 1838 (weight: 12.5-14.6 g; forearm: 47.5-53 mm). Our specimens confirm that R. alcyone is widely distributed in forest and forest-savanna mosaic. The closest previous records are from Akaniem north of Kete Krachi and from Breniasi and Worawora south of Apesokubi (Grubb 1971). In Côte d’Ivoire Fahr (1996) captured three out of five specimens in savanna. CONSERVATION STATUS. — Listed as “Least Concern” on the IUCN Red List, this species was the most common rhinolophid bat in our survey and is probably of lesser conservation concern in the Ghana-Togo Highlands., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on page 269, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["TEMMINCK C. J. 1853. - Esquisses zoologiques sur la Cote de Guine, Part I, Mammiferes. E. J. Brill, Leiden, 256 p. https: // www. biodiversitylibrary. org / page / 14835765","MARTIN W. 1838. - Description of a new bat (Rhinolophus landeri) from Fernando Po, and a new hedgehog (Erinaceus concolor) from Trebizond. Proceedings of the Zoological Society of London 1838: 101 - 103.","GRUBB P. 1971. - Further records of mammals from Ghana, based on the collections of Angus Booth. Revue de Zoologie et de Botanique Africaines 84: 192 - 202.","FAHR J. 1996. - Die Chiroptera der Elfenbeinkuste (unter Berucksichtigung des westafrikanischen Raumes): Taxonomie, Habitatpraferenzen und Lebensgemeinschaften. Diplomarbeit, Julius-Maximilians-Universitat, Wurzburg. 204 p."]}

Eidolon helvum (Kerr, 1792) Vespertilio vampyrus helvus Kerr, 1792: xvii: 91. COMMON NAME. — Straw-colored Fruit Bat; EWE: Aguto. MATERIAL EXAMINED. — Agumatsa (Wli) • 1 ♀; SMF 89666; brought to us by a local hunter; 19.XI.1999. REMARK A large colony of this gregarious and migratory fruit bat roosts on the trees and rock faces next to Wli Waterfall, recently estimated to include 250 000 individuals (Hayman et al. 2012) with an earlier report estimating the colony to include half a million E. helvum (Rice 1973), which would probably make it the most abundant fruit bat species in the area. Estimating the ecosystem services provided by the seed dispersal activities of E. helvum in Ghana and elsewhere in Africa was the subject of a recent study (van Toor et al. 2019). CONSERVATION STATUS. — Eidolon helvum has been listed as “Near Threatened” on the IUCN Red List because of significant population declines probably resulting from habitat loss and hunting for food and medicine (Peel et al. 2017)., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on page 266, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["KERR R. 1792. - The animal kingdom, or zoological system, of the celebratrd Sir Charles Linnaeus; class I: Mammalia. J. Murray & R. Faulder, London, 644 p. https: // doi. org / 10.5962 / bhl. title. 57940","HAYMAN D. T. S., MCCREA R., RESTIF O., SUU- IRE R., FOOKS A. R., WOOD J. L. N., CUNNINGHAM A. A. & ROWCLIFFE J. M. 2012. - Demography of straw-colored fruit bats in Ghana. Journal of Mammalogy 93: 1393 - 1404.","RICE K. W. 1973. - A faunal survey of Agumatsa Waterfall area, Volta Region. Ghana Department of Game and Wildlife, unpubl. Report.","VAN TOOR M. L., O'MARA M. T., ABEDI- LARTEY M., WIKELSKI M., FAHR J. & DECHMANN D. K. N. 2019. - Linking colony size with quantitative estimates of ecosystem services of African fruit bats. Current Biology 29: R 225 - R 240.","PEEL A. J., WOOD J. L. N., BAKER K. S., BREED A. C., CARVALHO A. D., FERNANDEZ- LORAS A., GABRIELI H. S., GEMBU G. - C., KAKENGI V. A., KALIBA P. M., KITYO R. M., LEMBO T., MBA F. E., RAMOS D., RODRIGUEZ- PRIETO I., SUU- IRE R., CUNNINGHAM A. A. & D. T. S. HAYMAN. 2017. - How does Africa's most hunted bat vary across the continent? Population traits of the Straw-Coloured Fruit Bat (Eidolon helvum) and its interactions with humans. Acta Chiropterologica 19: 77 - 92. https: // doi. org / 10.3161 / 15081109 ACC 2017.19.1.006"]}

Malacomys edwardsi Rochebrune, 1885 Malacomys edwardsi Rochebrune, 1885: 87. COMMON NAME. — Edwards’ Swamp Rat. MATERIAL EXAMINED. — Apesokubi • 1 ♀; USNM 59078; 28.VIII.2001; with four embryos (USNM 590105) with a crown -rump length of 19 mm • 2 ♂; USNM 590076, ZTNHC 960 • 1 ♀; USNM 590077. REMARK An obligate forest species in the Upper Guinea and an indicator of intact forest. A recent phylogeographic study confirmed the identification of all four of our M. edwardsi from Apesokubi but showed that these Volta Region specimens represent a lineage (F2) separate from the southwestern Ghanaian lineage (F1), “which may suggest the presence of multiple forest refugia” in Ghana (Bohoussou et al. 2015: 9). Hurst et al. (1995) reported 5 captures of M. edwardsi from moist semi-deciduous and secondary forest in KRNP which appears to be the only previous record of this species from the Ghana-Togo Highlands. We were not able to verify this species again at KRNP. CONSERVATION STATUS. — Malacomys edwardsi is currently listed as “Least Concern” on the IUCN Red List, but - already limited to the densest and best preserved forest in the Volta Region - it would be one of the first rodent species to disappear with continued deforestation., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on page 264, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["ROCHEBRUNE A. T. 1885. - Vertebratorum novorum vel minus cognitorum orae Africae occidentalis incolarum. Bulletin de la Societe Philomathique de Paris (ser. 7) 9: 86 - 100. https: // www. biodiversitylibrary. org / page / 31945680","BOHOUSSOU K. H., CORNETTE R., AKPATOU B., COLYN M., KERBIS PETERHANS J., KENNIS J., SUMBERA R., VERHEYEN E., N'GORAN E., KATUALA P. & NICOLAS V. 2015. - The phylogeography of the rodent genus Malacomys suggests multiple Afrotropical Pleistocene lowland forest refugia. Journal of Biogeography 42: 2049 - 2061. https: // doi. org / 10.1111 / jbi. 12570","HURST J., BOWES LYON D. & KEEFE E. O' 1995. - Kyabobo' 94: a joint Ghanaian and Oxford University Expedition. Final Report. KCP Publications, Ghana Wildlife Department, Institute for Renewable Natural Resources, 69 p."]}

Cricetomys cf. gambianus Waterhouse, 1840 (Fig. 4) Cricetomys gambianus Waterhouse, 1840: 2. COMMON NAME. — Northern Giant Pouched Rat; Ewe: Alegeli. MATERIAL EXAMINED. — Shiare • 1 adult ♀; 24.VIII.2001; Tomahawk trap along Sabu creek; SMF 91437. Apesokubi • 1 subadult ♀; Tomahawk trap; 28.VIII.2001; ZT- NHC 970. REMARK The zygomatic width of the Shiare specimen is 48.2% of occipito-nasal length, which is above the range of “ C. emini ”, following Genest-Villard (1967) (D. Kock pers. comm.), a species that is currently labeled Cricetomys sp. 1 from the region including our study area following the molecular and craniometric study by Olayemi et al. (2012). There is a specimen from the Volta Region at the USNM (USNM 436298, as. C. gambianus), a female from Odomi Jongo (1 miles E Nkwanta), caught during the African Mammal Project in 1968. CONSERVATION STATUS. — Cricetomys gambianus is listed as “Least Concern” on the IUCN Red List., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on pages 261-262, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["WATERHOUSE G. R. 1840. - On a new Genus of the Family Muridae and Order Rodentia. Proceeding of the Zoological Society of London 8: 1 - 3.","OLAYEMI A., NICOLAS V., HULSELMANS J. A. N., MISSOUP A. D., FICHET- CALVET E., AMUNDALA D., DUDU A., DIERCKX T., WEN- DELEN W. I. M., LEIRS H. & VERHEYEN E. 2012. - Taxonomy of the African giant pouched rats (Nesomyidae: Cricetomys): molecular and craniometric evidence support an unexpected high species diversity. Zoological Journal of the Linnean Society 165: 700 - 719. https: // doi. org / 10.1111 / j. 1096 - 3642.2012.00823. x"]}

Lemniscomys striatus (Linnaeus, 1758) Mus striatus Linnaeus, 1758: 62. COMMON NAME. — Striated Grass-Mouse; Ewe: Gbaxlé. MATERIAL EXAMINED. — Agumatsa Wildlife Sanctuary • 1 ♂; USNM 590104; 22.XI.1999 • 1♂; USNM 590075; 13.VIII.2001. Shiare • 1♀; SMF 91436 with 7 embryos; 22.VIII.2001; by hunter. Apesokubi (Site 1) • 1♀; ZTNHC 963; 26.XI.1999 • 1♀; SMF 91376; 28.VIII.2001. REMARK A grassland and farmbush species that also occurs along roads in the forest. There are 43 additional Lemniscomys striatus from the Volta Region at the USNM, collected by the African Mammal Project at Leklebi Agbesia and Odomi Jongo (2 miles E Nkwanta). Hurst et al. (1995) also caught four individuals in riparian forest and tree savanna at KRNP. Although not genetically tested our Lemniscomys striatus east of the Volta River should belong to what Nicolas et al. (2008) called Clade III. CONSERVATION STATUS. — Lemniscomys striatus is listed as “Least Concern” on the IUCN Red List., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on pages 262-264, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["LINNAEUS C. 1758. - Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus differentjis, synonymis, locis, Tomus I. Editio decima, reformata. L. Salvii, Holmiae, 500 p. https: // www. biodiversitylibrary. org / page / 2896589","VAN DER STRAETEN E. & DIETERLEN F. 1987. - Praomys misonnei, a new species of Muridae from Eastern Zaire (Mammalia). Stuttgarter Beitrage zur Naturkunde, Serie A (Biologie) 402: 1 - 11. https: // www. biodiversitylibrary. org / page / 33622925","NICOLAS V., OLAYEMI A., WENDELEN W. & COLYN M. 2010 b. - Mitochondrial DNA and morphometrical identification of a new species of Hylomyscus (Rodentia: Muridae) from West Africa. Zootaxa 2579: 30 - 44.","HURST J., BOWES LYON D. & KEEFE E. O' 1995. - Kyabobo' 94: a joint Ghanaian and Oxford University Expedition. Final Report. KCP Publications, Ghana Wildlife Department, Institute for Renewable Natural Resources, 69 p.","NICOLAS V., MBOUMBA J. - F., VERHEYEN E., DENYS C., LECOMPTE E., OLAYEMI A., MISSOUP A. D., KATUALA P. & COLYN M. 2008. - Phylogeographic structure and regional history of Lemniscomys striatus (Rodentia: Muridae) in tropical Africa. Journal of Biogeography 35: 2074 - 2089. https: // doi. org / 10.1111 / j. 1365 - 2699.2008.01950. x"]}

Rhinolophus landeri Martin, 1838 Rhinolophus landeri Martin, 1838: 101. COMMON NAME. — Lander’s Horseshoe Bat. MATERIAL EXAMINED. — Agumatsa Wildlife Sanctuary • 1♂; SMF 89670; 22.XI.1999. REMARK The single specimen of R. landeri could be distinguished from R. alcyone by its smaller size (weight 7.8 g; forearm 43.2 mm). A British Museum specimen of R. landeri is labeled as originating from “Wraura” (BMNH 55.378, leg. A. H. Booth), which may be the same as Worawora (near Apesokubi). A review of the bats of Côte d’Ivoire, showed that R. landeri occurs in all savanna formations to the northern edge of the Sudan savanna and that rainforest is actually being avoided (Fahr 1996; Fahr & Kalko 2011), contrary to Rosevear’s (1965) assessment of R. landeri as a rainforest species, which at his time included R. guineensis Eisentraut, 1960 as a subspecies. This is also supported by captures from Togo with just one specimen from the eastern edge of the Ghana-Togo Highlands at Atakpamé (De Vree et al. 1969) and 22 specimens caught in Northern Togo at Namoundjoga (De Vree et al. 1970). On the Accra Plains nine R. landeri were caught in forest remnants and in more open savanna (Decher 1997a). CONSERVATION STATUS. — R. landeri is listed as “Least Concern” by IUCN Red List. However, its uncommon occurrence in the Ghana-Togo Highlands and the fact that we found just one specimen in a forested valley make it of some conservation concern for the Volta Region., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on pages 269-270, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["MARTIN W. 1838. - Description of a new bat (Rhinolophus landeri) from Fernando Po, and a new hedgehog (Erinaceus concolor) from Trebizond. Proceedings of the Zoological Society of London 1838: 101 - 103.","NOACK TH. 1893. - Neue Beitrage zur Kenntniss der Saugethierfauna von Ostafrika. Zoologische Jahrbucher 7: 523 - 594.","TEMMINCK C. J. 1853. - Esquisses zoologiques sur la Cote de Guine, Part I, Mammiferes. E. J. Brill, Leiden, 256 p. https: // www. biodiversitylibrary. org / page / 14835765","FAHR J. 1996. - Die Chiroptera der Elfenbeinkuste (unter Berucksichtigung des westafrikanischen Raumes): Taxonomie, Habitatpraferenzen und Lebensgemeinschaften. Diplomarbeit, Julius-Maximilians-Universitat, Wurzburg. 204 p.","FAHR J. & KALKO E. K. V. 2011. - Biome transitions are centres of diversity: habitat heterogeneity and diversity patterns of West African bat assemblages across spatial scales. Ecography 34: 177 - 195. https: // doi. org / 10.1111 / j. 1600 - 0587.2010.05510. x","ROSEVEAR D. R. 1965. - The bats of West Africa. The British Museum (Natural History), London, xvii + 418 p.","DE VREE F., DE ROO A. & VERHEYEN W. N. 1969. - Contribution a l'etude des chiropteres de la Republique du Togo. Revue de Zoologie et de Botanique Africaines 83: 200 - 207.","DE VREE F., HULSELMANS J. & VERHEYEN W. 1970. - Contribution a l'etude des chiropteres de la Republique du Togo. 2. Liste preliminaire des Chiropteres. Revue de Zoologie et de Botanique Africaines 82: 41 - 46.","DECHER J. 1997 a. - Bat community patterns on the Accra Plains of Ghana, West Africa. Zeitschrift fur Saugetierkunde 62: 129 - 142."]}

Gerbilliscus kempi (Wroughton, 1906) Tatera kempii Wroughton, 1906: 375. COMMON NAME. — Kemp’s Gerbil. MATERIAL EXAMINED. — Kalakpa Resource Reserve • 1 ♂; brought to us by a Ghana Wildlife Division game scout; 18.XI.1999; ZT- NHC 973. REMARK This species is known from other locations in the Volta Region and the Togo Highlands (Robbins & Van der Straeten 1996; Grubb et al. 1998). Notwithstanding the debate voiced in Granjon et al. (2012) regarding the identity of West African G. kempi, we retain the name here, due to the far southern Dahomey Gap occurrence of our single specimen from Kalakpa Resource Reserve. CONSERVATION STATUS. — Gerbilliscus kempi is listed as “Least Concern” on the IUCN Red List., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on page 262, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["WROUGHTON R. C. 1906. - On some mammals collected by Mr. Robin Kemp in S. Nigeria. The Annals and Magazine of Natural History (ser. 7) 17: 375 - 379.","ROBBINS C. B. & VAN DER STRAETEN E. 1996. - Small mammals of Togo and Benin. II. Rodentia. Mammalia 60: 231 - 242.","GRUBB P., JONES T. S., DAVIES A. G., EDBERG E., STARIN E. D. & HILL J. E. 1998. - Mammals of Ghana, Sierra Leone and the Gambia. The Trendrine Press, Zennor, St. Ives, Cornwall, vi + 265 p.","GRANJON L., COLANGELO P., TATARD C., COLYN M., DOBIGNY G. & NICOLAS V. 2012. - Intrageneric relationships within Gerbil- liscus (Rodentia, Muridae, Gerbillinae), with characterization of an additional West African species. Zootaxa 3325: 1 - 25. https: // doi. org / 10.11646 / zootaxa. 3325.1.1"]}

Myonycteris angolensis smithii (Thomas, 1908) Rousettus smithi Thomas, 1908: 375. COMMON NAME. — Angolan Fruit Bat. MATERIAL EXAMINED. — Agumatsa Wildlife Sanctuary • 2 ♀; SMF 89667 + 92121. Apesokubi • 1 ♀; SMF 92122. Kyabobo NP • 1♀; SMF 92123. Liati-Wote • 2 ♀; SMF 92124 + 92125. REMARK We follow Nesi et al. (2013) in the use of the name Myonycteris Matschie, 1899 instead of Lissonycteris K. Andersen, 1912 for this fruit bat. Myonycteris angolensis smithii has previously been recorded in the Volta Region from Leklebi Agbesia, Akaniem, and Odomi Jongo (Bergmans 1997) and in neighboring Togo from Bismarckburg, Ahoué-houé, Aledjo, Odjolo, Pewa, and the Région d’Atakpamé (De Vree et al. 1969, 1970; Robbins, 1980; Grubb et al. 1998). Myonycteris a. smithii appears to be a relatively common forest and forest edge species throughout the Ghana-Togo Highlands and does not appear too affected by the widespread forest fragmentation.Most specimens in Ghana, Togo and Côte d’Ivoire have been recorded from the forest savanna mosaic and dry forests, but only a few from evergreen rain forest (Fahr 1996). We captured no males and only one female during the dry season in 1999 versus eight females from four localities in 2001. This may resemble the migration and possibly sexual segregation in this species observed at Mount Nimba, Liberia (Wolton et al. 1982) but details have to be verified for the Ghana-Togo Highlands. This fruit bat quickly attenuates to human handling. A female which stayed around our field camp at Apesokubi after release readily took banana pieces from our hands while hanging on a laundry line. An ectoparasite found on a M. s. smithii from Liati-Wote was a male Dipseliopoda biannulata (Oldyroyd, 1953) [Diptera: Nycteribiidae]. CONSERVATION STATUS. — Myonycteris angolensis is listed as Least Concern on the IUCN Red List (as Lissonycteris angolensis (Bocage, 1898)). Though not strictly a high forest species, it seems to be associated with dry forest of different types or gallery and island forests in the savanna zone and may depend on the existence of forest remnants in the Volta Region to survive., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on pages 267-268, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["THOMAS O. 1908. - A new fruit-bat from Sierra Leone. The Annals and Magazine of Natural History (ser. 8) 2: 375 - 376.","NESI N., KADJO B., POURRUT X., LEROY E., PONGOMBO SHONGO C., CRUAUD C. & HASSANIN A. 2013. - Molecular systematics and phylogeography of the tribe Myonycterini (Mammalia, Pteropodidae) inferred from mitochondrial and nuclear markers. Molecular Phylogenetics and Evolution 66: 126 - 137. https: // doi. org / 10.1016 / j. ympev. 2012.09.028","MATSCHIE P. 1899. - Die Megachiroptera des Berliner Museum fur Naturkunde. G. Reimer, Berlin. https: // doi. org / 10.5962 / bhl. title. 48691","BERGMANS W. 1997. - Taxonomy and biogeography of African fruit bats (Mammalia, Megachiroptera). 5. The genera Lissonycteris Andersen, 1912, Myonycteris Matschie, 1899 and Megaloglossus Pagenstecher, 1885; general remarks and conclusions; annex; key to all species. Beaufortia 47: 11 - 90.","DE VREE F., DE ROO A. & VERHEYEN W. N. 1969. - Contribution a l'etude des chiropteres de la Republique du Togo. Revue de Zoologie et de Botanique Africaines 83: 200 - 207.","DE VREE F., HULSELMANS J. & VERHEYEN W. 1970. - Contribution a l'etude des chiropteres de la Republique du Togo. 2. Liste preliminaire des Chiropteres. Revue de Zoologie et de Botanique Africaines 82: 41 - 46.","ROBBINS C. B. 1980. - Small mammals of Togo and Benin. I. Chiroptera. Mammalia 44: 83 - 88.","GRUBB P., JONES T. S., DAVIES A. G., EDBERG E., STARIN E. D. & HILL J. E. 1998. - Mammals of Ghana, Sierra Leone and the Gambia. The Trendrine Press, Zennor, St. Ives, Cornwall, vi + 265 p.","FAHR J. 1996. - Die Chiroptera der Elfenbeinkuste (unter Berucksichtigung des westafrikanischen Raumes): Taxonomie, Habitatpraferenzen und Lebensgemeinschaften. Diplomarbeit, Julius-Maximilians-Universitat, Wurzburg. 204 p.","WOLTON R. J., PRAK P. A., GODFRAY H. C. J. & WILSON R. P. 1982. - Ecological and behavioral studies of the Megachiroptera at Mount Nimba, Liberia. Mammalia 46: 419 - 448."]}

Hypsignathus monstrosus H. Allen, 1861 Hypsignathus monstrosus H. Allen, 1861: 157. COMMON NAME. — Hammer-headed Bat. REMARK Males of this species were only heard performing their characteristic lek-calling behaviour (Bradbury 1977) in a nearby mango tree at Agumatsa Wildlife Sanctuary in August 2001. Hypsignathus monstrosus has been previously recorded in the Volta Region from Odomi Jongo, 2 miles E Nkwanta and Leklebi Agbesia (USNM). CONSERVATION STATUS. — Hypsignathus monstrosus is listed as “Least Concern” on the IUCN Red List., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on page 269, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["ALLEN H. 1861. - Descriptions of new pteropine bats from Africa. Proceedings of the Academy of Natural Sciences of Philadelphia 13: 157 - 160. https: // www. biodiversitylibrary. org / page / 1683285","TEMMINCK C. J. 1853. - Esquisses zoologiques sur la Cote de Guine, Part I, Mammiferes. E. J. Brill, Leiden, 256 p. https: // www. biodiversitylibrary. org / page / 14835765","BRADBURY J. W. 1977. - Lek mating behavior in the hammerheaded bat. Zeitschrift fur Tierpsychologie 45: 225 - 255. https: // doi. org / 10.1111 / j. 1439 - 0310.1977. tb 02120. x"]}

Crocidura cf. eburnea Heim de Balsac, 1958 Crocidura (bottegi) eburnea Heim de Balsac, 1958: 327. COMMON NAME. — Ivory Coast White-toothed Shrew. MATERIAL EXAMINED. — Apesokubi • 1 ♀; pitfall; 27.XI.1999; ZFMK 1999.1144. REMARK Initially identified as Crocidura cf. douceti Heim de Balsac, 1958, this tiny shrew has been re-assigned to Crocidura eburnea, which has only recently been elevated from a synonym of C. obscurior (Hutterer 2005) to species level (Jacquet et al. 2014). This is the first record of C. cf. eburnea from the Volta Region and since the Volta River is considered the eastern boundary of the C. obscurior complex (Jacquet et al. 2014), this possible range extension awaits further study. CONSERVATION STATUS. — Crocidura eburnea was included in the IUCN Red List in 2019 and assessed as “Least Concern”., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on page 261, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["HEIM DE BALSAC H. 1958. - La reserve integrale du Mont Nimba, XIV. Mammiferes insectivores. Memoires de l'Institut Francaise d'Afrique Noire 53: 301 - 337.","HUTTERER R. 2005. - Order Insectivora in WILSON, D. E. & REEDER D. M. (eds), Mammal species of the world: a taxonomic and geographic reference. 3 rd Edition. Smithsonian Institution Press, Washington, D. C.: 212 - 311","JACQUET F., NICOLAS V., COLYN M., KADJO B., HUTTERER R., DECHER J., AKPATOU B., CRUAUD C. & DENYS C. 2014. - Forest refugia and riverine barriers promote diversification in the West African pygmy shrew (Crocidura obscurior complex, Soricomorpha). Zoologica Scripta 43: 131 - 148.","DOLLMAN G. 1915. - On the African shrews belonging to the genus Crocidura. Journal and Magazine of Natural History (ser. 8) 15: 507 - 527. https: // www. biodiversitylibrary. org / page / 24256132"]}

Myotis bocagii cupreolus Thomas, 1904 (Fig. 13) Myotis bocagei cupreolus Thomas, 1904: 407. COMMON NAME. — Rufous Mouse-eared Bat. MATERIAL EXAMINED. — Agumatsa (Wli Waterfall) • 2 ♀; SMF 89673, SMF 92136. REMARK Only two specimens were collected at the same river crossing at Agumatsa, one in the dry season on 22 Nov. 1999 and one in the wet season on 11.VIII.2001. These appear to be the first record of M. bocagii from the Volta Region and from the Ghana-Togo Highlands. The closest records in Ghana are from the lower Volta River near Kpong (Decher et al. 1997), from Jukwa in the Central Region (USNM 4215; 4216) and from Boti Falls in the Eastern Region (ROM 55137, 60227). The only known record from Togo is from Borgou in Northern Togo (De Vree & Van der Straeten 1971). CONSERVATION STATUS. — Classified as “Least Concern”on the IUCN Red List. This bat depends on water for its foraging flight (Brosset 1976), as well as riverine forest vegetation and banana and other musaceous plants whose furled leaves it uses as a roost (Happold 2013c). Its survival in the Ghana-Togo Highlands is thus of some conservation concern if forest loss continues., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on pages 271-272, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["THOMAS O. 1904. - On mammals from Northern Angola collected by Dr. W. J. Ansorge. The Annals and Magazine of Natural History (ser. 7) 13: 405 - 421. https: // doi. org / 10.1080 / 00222930408562472","DECHER J., SCHLITTER D. A. & HUTTERER R. 1997. - Noteworthy records of small mammals from Ghana with special emphasis on the Accra Plains. Annals of Carnegie Museum 66: 209 - 227.","DE VREE F. & VAN DER STRAETEN E. 1971. - Contribution a l'etude des chiropteres de la Republique duTogo. 3. Liste preliminaire des Chiropteres recoltes par la troisieme Mission zoologique belge au Togo. R evue de Zoologie et de Botanique Africaines 83: 159 - 164.","BROSSET A. 1976. - Social organization in the African bat, Myotis bocagii. Zeitschrift fur Tierpsychologie 42: 50 - 56. https: // doi. org / 10.1111 / j. 1439 - 0310.1976. tb 00955. x","HAPPOLD M. 2013 c. - Myotis bocagii Rufous Myotis (Rufous Mouse-eared Bat), in HAPPOLD M. & HAPPOLD D. C. D. (eds), The Mammals of Africa - Vol. IV: Hedgehogs, Shrews and Bats. Bloomsbury Publishing, London: 692 - 694."]}

Epomops franqueti (Tomes, 1860) Epomophorus franqueti Tomes, 1860: 54. COMMON NAME. — Franquet’s Epauletted Fruit Bat; EWE: Aguto. MATERIAL EXAMINED. — Kalakpa Resouce Reserve • 1 ♀; SMF 89662). Agumatsa Valley • 2 ♀; SMF 89663, ZTNHC 959. REMARK Although this is a mostly forest-dwelling fruit bat distributed from eastern Côte d’Ivoire into Central Africa (Happold 2013a), our specimen from Kalakpa Resource Reserve shows that it can cope with forest remnants and gallery forest in more savanna-dominated areas. Epomops franqueti was previously known in the Volta Region from Amedzofe, Bator, Leklebi Agbesia, and Odomi Jongo (Bergmans 1989). CONSERVATION STATUS. — Epomops franqueti is listed as “Least Concern” on the IUCN Red List., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on page 266, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["TOMES R. F. 1860. - A monograph of the genus Epomophorus, with the description of a new species. Proceedings of the Zoological Society of London 28: 42 - 58. https: // www. biodiversitylibrary. org / page / 12866542","HAPPOLD M. 2013 a. - Epomops franqueti Franquet's Epauletted Fruit Bat, in HAPPOLD M. & HAPPOLD D. C. D. (eds). The Mammals of Africa - Vol. IV: Hedgehogs, Shrews and Bats. Bloomsbury Publishing, London: 256 - 258.","BERGMANS W. 1989. - Taxonomy and biogeography of African fruit bats (Mammalia, Megachiroptera). 2. The genera Micropterus Matschie, 1899, Epomops Gray, 1870, Hypsignathus H. Allen, 1861, Nanonycteris Matschie, 1899, and Plerotes Andersen, 1910. Beaufortia 39: 89 - 153."]}

Nycteris arge Thomas, 1903 Nycteris arge Thomas, 1903: 633. COMMON NAME. — Bate’s Slit faced Bat. MATERIAL EXAMINED. — Apesokubi • 1 ♀; SMF 92126; 26.VIII.2001. REMARK Although this bat is known from the Upper Guinea Region west of the Volta River and from central and southern Nigeria (Happold 1987), it does not seem to have been recorded from the Volta Region, or in the Dahomey Gap. The closest records are two BMNH specimens from Akaniem (Akaniem, Buem) between the Volta and Oti River branches of presentday Lake Volta (Grubb 1971). Van Cakenberghe & De Vree (1985) apparently list these specimens as being from “Akenim”, Togo. In Côte d’Ivoire, Fahr (1996) recorded six out of seven specimens of N. arge within the rainforest zone and one in forest-savanna mosaic. Our specimen of N. arge appears to be the first one reported from the Volta Region. CONSERVATION STATUS. — Although classified as a species of “Least Concern” on the IUCN Red List, as a mostly forest-dependent species, N. arge is of some conservation concern in the Ghana-Togo Highlands., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on page 269, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["THOMAS O. 1903. - Three new bats from the Cameroons, discovered by Mr. G. L. Bates. The Annals and Magazine of Natural History (ser. 7), 12: 633 - 635. https: // doi. org / 10.1080 / 00222930309487045","HAPPOLD D. C. D. 1987. - The Mammals of Nigeria. Clarendon Press, Oxford, xvii + 402 p.","GRUBB P. 1971. - Further records of mammals from Ghana, based on the collections of Angus Booth. Revue de Zoologie et de Botanique Africaines 84: 192 - 202.","VAN CAKENBERGHE V. & DE VREE. F. 1985. - Systematics of African Nycteris (Mammalia: Chiroptera), in SCHUCHMANN K. L (ed.), International Symposium on African Vertebrates, Systematics, Phylogeny and Evolutionary Ecology, 15 - 18 May 1984. Zoologisches Forschungsinstitut und Museum Alexander Konig, Bonn, Selbstverlag. Bonn, Germany: 35 - 90.","FAHR J. 1996. - Die Chiroptera der Elfenbeinkuste (unter Berucksichtigung des westafrikanischen Raumes): Taxonomie, Habitatpraferenzen und Lebensgemeinschaften. Diplomarbeit, Julius-Maximilians-Universitat, Wurzburg. 204 p."]}

Monadjem, Ara, Richards, Leigh R., Decher, Jan, Hutterer, Rainer, Mamba, Mnqobi L., Guyton, Jen, Naskrecki, Piotr, Markotter, Wanda, Wipfler, Benjamin, Kropff, Anna S., Dalton, Desire L. (2021): A phylogeny for African Pipistrellus species with the description of a new species from West Africa (Mammalia: Chiroptera). Zoological Journal of the Linnean Society 191 (191): 548-574, DOI: 10.1093/zoolinnean/zlaa068/5901436

PIPISTRELLUS SIMANDOUENSIS MONADJEM, RICHARDS, DECHER AND HUTTERER, SP . NOV . SIMANDOU SEROTINE LSID: http://zoobank.org/urn:lsid:zoobank.org:pub:A1F9BAC5-40BA-42FD-8D76-5284691656F4 Holotype: ZMFK-MAM-2008.0302, field number: JD 614. The bat was collected by Jan Decher and team. It is an adult male fixed in formalin and currently preserved in 70% alcohol, with the skull extracted and cleaned. Photographs of the head and tragus are illustrated in Figures 4 and 5, while the mandible and skull of the holotype are illustrated in Figure 6. Images of the entire skeleton of the holotype, including the skull, are presented in Supporting Information (Fig. S 1). The glans penis is illustrated in Figure 8 and the dorsal, ventral and lateral views of the baculum are presented in Figure 9. Type locality: Guinea, Macenta, Simandou Range, “Whiskey 1” as mapped in Decher et al. (2015) (Fig. 1). The bat was netted on 25 February 2008 across a small creek in a forested ravine on the east slope of the Simandou Range (08°32’53.84”N, 08°53’48.07”W) at an elevation of 950 m above sea level. Paratypes: T w o o t h e r b a t s, a f e m a l e (ZMFK MAM2008.0300; field number: JD 656) and a male (ZMFK-MAM-2008.0301; field number: JD 661) identified as belonging to this species were captured 5.9 km to the south-west of the type locality at “Foko confluence” on the west slope of the Simandou Range, on 8 and 9 March 2008, respectively, as mapped in Decher et al. (2015), in more humid forest habitat and at 737 m above sea level. Both these specimens have been sequenced and clearly group with the holotype (Fig. 2A) and can be considered as paratypes. Etymology: This species is named after the type region, the Simandou mountain range in eastern Guinea. Diagnosis: A m e d i u m - s i z e d p i p i s t r e l l o i d b a t, assignable to the genus Pipistrellus on the basis of the presence of a small anterior upper premolar (Hill & Harrison, 1987) and phylogeny (Fig. 2). Some members of the genus Parahypsugo may also have this anterior upper premolar but differ in skull morphology and shape of rhinarium (Hutterer et al., 2019), the cranium being more inflated in Pi. simandouensis than in any Parahypsugo species. Pi. simandouensis is readily distinguishable from Pi. hesperidus (its sister taxon) by its unicoloured pelage (Fig. 4) and shape of baculum (Fig. 5); however, these two species are indistinguishable in craniodental or external measurements. It is significantly larger in external and cranial features (see below) than Pi. nanulus Thomas, 1904, Pi. rusticus (Tomes, 1861) and Pipistrellus deserti Thomas, 1902. It differs from Pipistrellus inexspectatus Aellen, 1959 by its unicoloured pelage (bicoloured in the latter) and lack of white on the trailing edge of the wing membrane. Pi. (Vansonia) rueppellii is readily distinguished from all other Pipistrellus species by its pure white underparts and is currently placed in a separate genus Vansonia Roberts, 1946 (Moratelli & Burgin, 2019). Finally, the poorly diagnosed Pi. musciculus Thomas, 1913 (which may not even be a Pipistrellus; Hill & Harrison, 1987), is far smaller in external and cranial measurements. Description: External characters: Pi. simandouensis is a medium-sized pipistrelloid bat (similar in size to that of Pi. hesperidus), but large for the genus Pipistrellus (in fact it is the largest within the genus in Africa), with a total length of 80–86 mm and forearm length 31–34 mm (Table 3). The pelage is bright yellowishbrown and paler below than above (Fig. 4), with the individual hairs being unicoloured on both the upper and under parts. The patagium and uropatagium are both dark brown. Typically for the genus Pipistrellus, the ears are subtriangular in shape, rounded at the tip, and dark brown in colour (Fig. 4C). The tragus is also typical of the genus and is moderately long (roughly half of the length of the ear), relatively broad with a straight leading edge and convex outer edge (Fig. 9). It bears a markedly pointed projection, situated near the base and lying immediately below the indentation/notch of the outer margin. The rhinarium is as illustrated for the genus Pipistrellus in Hutterer et al. (2019), with the nostrils rounded in shape and obviously protruding from the snout. The external measurements of the holotype and other specimens of Pi. simandouensis are shown in Table 3. Craniodental characters: The skull is relatively robust for a Pipistrellus, while the rostrum is neither particularly broad nor narrow. The brain case is moderately inflated and rises distinctly and sharply above the level of the rostrum (Fig. 6) in contrast to the relatively flatter skulls of Parahypsugo spp. (Hutterer et al., 2019). The posterior of the skull does not end in an extended parietal/supraoccipital crest and the sagittal and lambdoid crests are poorly developed. The zygomatic arch is moderately robust. Cranial measurements for the holotype and other specimens of Pi. simandouensis are shown in Table 4. The dentition of Pi. simandouensis is I 2/3, C 1/1, P 2/3, M 2/3, which is typical of the genus Pipistrellus (Van Cakenberghe & Happold, 2013a). In the upper tooth row, I 1 is not bifid and I 2 is relatively smallsized, not reaching half the length of I 1 (Fig. 7A). P 1 is relatively small in all specimens examined, and in the toothrow, creating an obvious gap between C and P 2 (Fig. 7B). Dental measurements for the holotype and other specimens of Pi. simandouensis are shown in Table 5. The penis of Pi. simandouensis is long (8.4 mm in the holotype), straight and covered by long hairs all over its length (Fig. 8). These hairs are mostly white, with only a few on the dorsal side are brown. The terminal glans is wider than the shaft (Fig. 8). Total length of the penis is about 10% of total body length (Table 3). In its long and straight shape, the penis of Pi. simandouensis is similar to other species of the genus, such as Pipistrellus abramus (Temminck, 1840) and other Asian (Francis, 2019) and African Pipistrellus species (Benda et al., 2004). The bacula of Pi. simandouensis, Pi. grandidieri, Pi. hesperidus, Pi. nanulus and Pi. rusticus are presented in Figure 9. The baculum of Pi. simandouensis was notably shorter than the other species, with a total length of 1.32 mm. Baculum total length for the remaining species was: 2.00 mm (Pi. grandidieri), 1.47 mm (Pi. hesperidus), 2.98 mm (Pi. nanulus) and 1.52 mm (Pi. rusticus). Pi. simandouensis, Pi. hesperidus and Pi. rusticus were overall similar in morphology, presenting a slender-shafted baculum with a bilobed base and an expanded tip with two distinct prongs. However, Pi. simandouensis can be differentiated from Pi. hesperidus and Pi. rusticus on the basis of its robust, triangular base measuring 0.32 mm. Similarly, the projections of the tip in Pi. simandouensis are more pointed than in Pi. hesperidus and Pi. rusticus. The full suite of bacula measurements for all five species are provided in Supporting Information (Table S4). Biology: Pipistrellus simandouensis is currently known from two localities in West Africa, based on the four sequenced specimens (appearing in Fig. 2A), one specimen from Mount Nimba in northern Liberia (Monadjem et al., 2013) and three from the Simandou Range in south-eastern Guinea (Decher et al., 2015). This species has also been reported from the Fouta Djallon in central Guinea (Decher et al., 2015); however, this has not been confirmed genetically and we have not been able to examine these specimens. However, all these sites are from upland regions of West Africa, ranging in elevation from the foothill slopes at around 450 m a.s.l., to the summit of some of the peaks at over 1200 m a.s.l. and higher. Therefore, we suspect that this is a species closely associated with these upland forested habitats. Because such upland habitats are relatively restricted in West Africa, and few are legally protected (Monadjem et al., 2016), we furthermore predict that this is a species under threat. Hence, we recommend that the conservation status of this species be assessed as a matter of urgency., Published as part of Monadjem, Ara, Richards, Leigh R., Decher, Jan, Hutterer, Rainer, Mamba, Mnqobi L., Guyton, Jen, Naskrecki, Piotr, Markotter, Wanda, Wipfler, Benjamin, Kropff, Anna S. & Dalton, Desire L., 2021, A phylogeny for African Pipistrellus species with the description of a new species from West Africa (Mammalia: Chiroptera), pp. 548-574 in Zoological Journal of the Linnean Society 191 (191) on pages 559-563, DOI: 10.1093/zoolinnean/zlaa068/5901436, http://zenodo.org/record/4451311, {"references":["Decher J, Hoffmann A, Schaer J, Norris RW, Kadjo B, Astrin J, Monadjem A, Hutterer R. 2015. Bat diversity in the Simandou Mountain Range of Guinea, with the description of a new white-winged vespertilionid. Acta Chiropterologica 17: 255 - 282.","Tamura K, Stecher G, Peterson D, Filipski A, Kumar S. 2013. MEGA 6: molecular evolutionary genetics analysis version 6.0. Molecular Biology and Evolution 30: 2725 - 2729.","Hutterer R, Decher J, Monadjem A, Astrin J. 2019. A new genus and species of vesper bat from West Africa, with notes on Hypsugo, Neoromicia, and Pipistrellus (Chiroptera: Vespertilionidae). Acta Chiropterologica 21: 1 - 22.","Moratelli R, Burgin CJ. 2019. Family Vespertilionidae (vesper bats). In: Wilson DE, Mittermeier RA, eds. Handbook of the mammals of the world - Vol. 9. Barcelona: Lynx Edicions.","Thomas O. 1913. On African bats and shrews. Annals and Magazine of Natural History 11: 314 - 321.","Van Cakenberghe V, Happold M. 2013 a. Genus Pipistrellus. Pipistrelles. In: Happold M, Happold DCD, eds. The mammals of Africa. Vol. IV: hedgehogs, shrews and bats. London: Bloomsbury Publishing, 600 - 604.","Francis CM. 2019. Field guide to the mammals of South-East Asia. London: Bloomsbury Publishing.","Benda P, Hulva P, Gaisler J. 2004. Systematic status of African populations of Pipistrellus pipistrellus complex (Chiroptera: Vespertilionidae), with a description of a new species from Cyrenaica, Libya. Acta Chiropterologica 6: 193 - 217.","Monadjem A, Richards L, Taylor PJ, Stoffberg S. 2013. High diversity of pipistrelloid bats (Vespertilionidae: Hypsugo, Neoromicia, and Pipistrellus) in a West African rainforest with the description of a new species. Zoological Journal of the Linnean Society 167: 191 - 207.","Monadjem A, Richards L, Denys C. 2016. An African bat hotspot: the exceptional importance of Mount Nimba for bat diversity. Acta Chiropterologica 18: 359 - 375."]}

Madrider Mitteilungen, Vol. 53 (2012), Im Rahmen des von der Volkswagenstiftung finanzierten Projekts ���Sp��tquart��rer Landschafts- und Nutzungswandel im semiariden Nordosten Marokkos��� fanden in den Jahren 2006 bis 2008 geoarch��ologische Feldforschungen in den alluvialen Sedimenten der Unteren Moulouya (Nordost-Marokko) statt. Ziel des Projekts war es, in einem semiariden und daher umweltsensiblen Raum die Reaktion des Menschen auf Umweltver��nderungen zu untersuchen. Die bis zu 15 m hohen Auensedimente lagerten sich in einen Zeitraum von ca. 10.000��� 800 calBP ab und repr��sentieren damit nahezu das ganze Holoz��n im Arbeitsgebiet. Eingebettet in diese Sedimente finden sich arch��ologische Fundstellen, von denen 40 dokumentiert und teilweise ausgegraben wurden. Initiale Bodenbildungen und zahlreiche epipal��olithische und fr��hneolithische Fundstellen deuten auf optimale Siedlungsbedingungen zu Beginn des Holoz��ns hin. Fehlende Bodenbildungen und ein R��ckgang der Besiedlung markieren das Ende dieser Gunstphase um 6300 calBP. Eine noch dramatischere Verschlechterung setzt ab 3500 calBP ein. G��nzlich fehlende Hinweise auf eine Besiedlung und hohe Fluth��ufigkeit legen einen Siedlungsabbruch im Arbeitsgebiet nahe. Das ��ndert sich erst wieder mit dem Beginn der islamischen Periode ab ca. 1250 calBP.

International audience; Our work entitled: Diversity of Rodents (Rodentia: Mammalia) in the Kandolo Forest Reserve (Maniema Province, DR Congo) aimed to assess the biodiversity of Rodents in the Kandolo Forest Reserve while comparing the specific diversity during two capture sessions and in the two prospected habitats (Mixed primary forest (FPM) and Gilbertiodendron dewevrei primary forest (FPG) Finally, assess the distribution of Rodents in the two habitats (FPG and FPM) prospected by sex. These objectives have been achieved. To achieve this, only one method was used in the field, the only in-line trapping using two types of traps including Sharmen and Pitfall. Two habitats were explored (primary mixed forest and primary forest at Gilbertiodendron dewevrei) during 2 capture sessions. After processing the data, the results presented 153 captured Rodents synthesized as follows: For a total of 153 individuals captured in the two habitats prospected during the two capture sessions, the first capture session carried out in the FPM, totaled 40 individuals of Rodents including: Praomys cf. jacksoni is the most represented with 22 individuals captured or 55 %. In contrast, Graphiurus lorraineus, Lophuromys luteogaster and Saccostomus campestris are the least represented with a score of an individual captured, 2, 5 %. On the other hand, during the second session which took place in the FPG, it emerges the following: For the 113 individuals captured, Praomys cf. jacksoni is the most represented with 69 individuals captured, or 61, 06 %. Lophuromys dudui, on the other hand, is the least represented with a score of one specimen, ie 0, 88 %. In comparison with habitats and capture sessions, the following is concluded: the FPG is the best represented quantitatively and qualitatively (113 specimens, TS = 7, 06 %, Ep = 1600 trap nights and 10 species) with Praomys cf. jacksoni which ranks first with 69 individuals out of 113 carcasses captured. Unlike the FPM which occupies the second position (40 specimens, TS = 2, 5 %, EC = 1600 trap nights,) with 9 species caught. But for the indices of specific diversity, it appears that, in the two habitats there is the probability of randomly drawing 2 individuals to have 2 different species because Simpson tends towards1 everywhere. And in the two habitats, the species are not evenly distributed (E tends towards 1). Also, the specific richness evaluated at the level of the two different habitats reveals that there is not a significant difference between the populations of Rodents (Kuskal-Walis test, H = 2, 802; p = 0, 09055). Finally, taking into account the sex distribution of the Rodents caught in the two habitats during the two sessions, it has been shown that, the Rodents sampled are marked by the dominance of the males in percentage as the females. Thus, in the FPM the dominance of the males is expressed at 70 % against 30 % of the females and in the FPG 69, 91 % of the males against 30, 09 % of the females.