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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."]}