Rhinolophus clivosus, Cretzschmar, 1826

Burgin, Connor, 2019, Rhinolophidae, Handbook of the Mammals of the World – Volume 9 Bats, Barcelona: Lynx Edicions, pp. 280-332 : 296-297

32 . Geoffroy’s Horseshoe Bat

Rhinolophus clivosus

French: Rhinolophe de Cretzschmar / German: Geoffroy-Hufeisennase / Spanish: Herradura de Geoffroy

Other common names: Arabian Horseshoe Bat, Cretzschmar’s Horseshoe Bat

Taxonomy. Rhinolophus clivosus Cretzschmar in Rùppell, 1828 ,

Red Sea Coast , Saudi Arabia .

Rhinolophus clivosus is in the jfemzTwegwzwra species group and is closely related to R ferrumequinum. Rhinolophus clivosus currently represents a paraphyletic species complex with respect to 7t and is probably composed of at least three species from Arabia and Socotra { clivosus ), East Africa {acrotis), and southern Africa {augurar geoffmyii if the latter is determined to be a valid name). Egyptian populations of R clivosus clustered with R ferrumequinum in phylogenetic studies, but there has been no genetic sampling from the rest of North Africa, east-central Africa, and the Levant so the phylogenetic position of all populations has not yet been determined. There are five genetically distinct populations from southern Africa that could represent subspecies of the southern African species. The name geoffmyii is here recognized as a subspecies because no other name is available for a genetically distinct population in south-western South Africa, but the name is not considered valid by most authors because it lacks a type specimen. Populations originally attributed to R clivosus from Cyrenaica, Libya, recently have been classified as a distinct species, R haraceki . R clivosus traditionally has contained ten subspecies with considerable morphological, ecological, echolocation, and genetic diversity; additional research might result in taxonomic changes. Ten subspecies recognized.

Subspecies and Distribution.

R. c.c clivosus Cretzschmar, 1828 - SWJordan, W & C Saudi Arabia, Yemen, and SW Oman.

R c. acrotis Heuglin, 1862 — Eritrea, Ethiopia, Djibouti, and N Somalia.

R c. augur ÌL. Andersen, 1904 — N & C South Africa.

R c. brachygnathus K. Andersen, 1905 - Israel, Egypt, and N Sudan.

R c. geoffmyii A. Smith, 1829 — SW South Africa.

R c. keniensis Hollister, 1916 - SE Sudan, South Sudan, Uganda, NE DR Congo, Rwanda, Burundi, Kenya, and N Tanzania; other records throughout C Africa need further investigation.

R c. schwarzi Heim de Balsac, 1934 - SE Algeria and W Libya.

7t c., socotranus Benda, Reiter & Vallo, 2017 - Socotra I, Yemen.

AE c. zambesiensis K. Andersen, 1904 - S Tanzania, Malawi, Zambia, and SE DR Congo S to NE South Africa.

R c. zuluensis K. Andersen, 1904 - E & S South Africa, Swaziland, and Lesotho.

There is also a record from W DR Congo with no subspecific affinity and populations from Namibia and SW Angola are not currently assigned to any subspecies but might be associated with subspecies augur or geoffroyii following further morphological and genetic tests.

Descriptive notes. Head—body 46-50 mm, tail 22-40 mm, ear 16-24 mm, hindfoot 11—13 mm, forearm 42-59 mm; weight 10-25 g. Dorsal pelage is highly variable and can be cream, gray, brownish gray, grayish fawn, or reddish brown (hairs usually have dark tips); venter varies from beige and pale brown to gray. Orange-morph (more reddish brown) individuals have been recorded. Adult males lack axillary tufts. Ears are short (30-47% of forearm length). Noseleaf has hastate lancet, with rounded tip; connecting process is rounded and elevated, can be smoothly curved or slighdy angular, and is slighdy to clearly higher than sella tip; sella is naked and narrow and has concave sides and broad and rounded tip; and horseshoe is narrow to moderately wide at 6-6-9- 6 mm, varies widely among subspecies, and has lateral leaflets (rudimentary in some individuals) and variably median emargination, varying between shallow to moderately deep. Lower lip has well-defined medial groove. Wings and uropatagium are light to dark gray. Baculum has shallow dorsal invagination, deep ventral invagination on basal cone, and dorso-ventrally flattened shaft; subspecies augur has slightly expanded tip. Skull is robust; zygomatic width is greater than mastoid width; nasal swellings are relatively low; frontal depression is very shallow, and supraorbital ridges are weak; sagittal crest is low anteriorly and completely absent posteriorly; and interpterygoid groove is absent or very undeveloped. P2 is minute and completely displaced labially or absent all together, allowing C1 and P4 to touch, and P3 is tiny and completely displaced labially or absent when P2 and P4 are in contact. Dental formula is variable: I 1/2, C 1/1, P 2/2, M 3/3 (x2) = 30; I 1/2, C 1/1, P 1/2, M 3/3 (x 2) = 28; I 1/2, C 1/1, P 2/3, M 3/3 (x2) = 32; or 11/2, C 1/1, P 1/3, M 3/3 (x2) = 30. Chromosomal complement has 2n = 58 and FNa = 60 ( South Africa) or 62 (South and East Africa).

Habitat. Wide variety of habitats but generally drier and open environments such as woodland savannas, Mediterranean shrubland, dry savanna, open grasslands, and some semi-desert and desert habitats from sea level to elevations of c. 2300 m. Although Geoffroy’s Horseshoe Bats prefer dry habitats, they are often found associated with water sources, dense vegetation where they can forage, and caves for roosting. In southern Africa, they are found largely in woodland savannas, some desert habitats, and montane grasslands (e.g. Drakensberg). Populations in West Africa are found largely in mountainous and hilly regions. In Malawi, they have been recorded in miombo forest and woodlands and thicket savannas. In Northern Africa, Geoffroy’s Horseshoe Bats are primarily restricted to mountain “islands” throughout the Sahara Desert and sub-desert and savanna habitats in north-eastern Africa. They occur primarily in arid Mediterranean shrubland in the Levant and arid shrubland, steppe, and savanna throughout Arabia and Socotra.

Food and Feeding. Geoffroy’s Horseshoe Bats are insectivorous. They forage by slow hawking, fly-catching, and probably gleaning prey offvegetation and the ground. Foraging occurs around vegetation under tree canopies. After prey is captured, they eat it while perching on branches or other vegetation. They feed primarily on moths and beetles throughout their distribution. Fecal samples from two bats in Zimbabwe included only beetles. Populations in South Africa fed primarily on Lepidoptera and Coleoptera and smaller amounts of Neuroptera, Hemiptera, and Diptera . In Algeria, 46 fecal pellets from five bats included Lepidoptera (mean 62-6% by volume), Coleoptera (29-4%), Hemiptera (2%), Hymenoptera (1-4%), and unknown arthropods (4-6%). Twenty-three fecal samples from two bats in Jordan mainly contained Lepidoptera, Coleoptera , and Diptera (especially Nematocera) and smaller amounts of Hymenoptera, Neuroptera, Trichoptera, and Hemiptera. Captive individuals have eaten mantises. In South Africa, Geoffroy’s Horseshoe Bat seemed to eat larger prey than the Cape Horseshoe Bat ( A capensis ) on average, but prey size of the two species did overlap. Wild Geoffroy’s Horseshoe Bats seem to depend on water sources for drinking; captive individuals drink regularly.

Breeding. Geoffroy’s Horseshoe Bat is seasonally monoestrous, at least in southern Africa ( Zimbabwe and South Africa). Copulation seems to occur in dry season in Zimbabwe (June-July) and autumn in South Africa (April). Females store sperm. Parturition begins in wet season (mid-November) in Zimbabwe and late winter (August) in South Africa. In South Africa, however, not all populations exhibit delayed implantation where females store sperm, and they will copulate in late winter. Gestation lasts 3-5 months, and lactation lasts c.2 months. Most females appear to reach reproductive maturity at c.18 months of age, but it can take as long as 42 months or more.

Activity patterns. Geoffroy’s Horseshoe Bats are nocturnal and forage throughout the night. During the day, they can enter torpor, and in Malawi, they became torpid at ambient temperatures of 21-24°C. Day roosts are usually in caves, rock crevices, and artificial underground structures such as abandoned mine shafts, catacombs, stone huts, and other rocky areas. They will roost in abandoned buildings and hollow baobabs { Adansonia , Malvaceae ). On Socotra, primary day roosts are located in limestone karst caves. Call shape is FM/ CF /FM; they are able to distinguish sex based on call (probably using FM components) and even individual identity (probably using resting frequency). Peak (CF) component has been recorded at 90-100 kHz or 80-85 kHz in various parts of South Africa, 91 -9 kHz in Swaziland, 79-84 kHz (usually 82-84 kHz) in Malawi, 79-8-81 kHz in Mozambique, 92-7 kHz in Algeria, 85-2 kHz in Israel, and 83-5-85 kHz in Jordan. Mean call duration is 32-9 milliseconds in Swaziland, 14-3 milliseconds in South Africa, 54-5 milliseconds in Israel, and 48 milliseconds in Jordan.

Movements, Home range and Social organization. Geoffroy’s Horseshoe Bat roosts singly or in groups of up to 50 individuals in most cases, although colonies of up to 10,000 individuals have been recorded in Malawi and southern Africa. Roosting bats generally hang in small clusters not touching each another; many small clusters are formed in large colonies.

Status and Conservation. Classified as Least Concern on The IUCNRed List. Geoffroy’s Horseshoe Bat is relatively common throughout much of its distribution, although it might be declining in Palearctic parts of its distribution. There are no overarching threats, but some populations might be locally threatened by habitat destruction and roost disturbance. Geoffroy’s Horseshoe Bat might be locally threatened by indirect poisoning from insecticides, pesticides, and similar chemicals. It is protected inJordan by national legislation.

Bibliography. ACR (2018), Benda & Vallo (2012), Benda, Dietz eta /. (2008), Benda, Lucan eta/. (2010), Benda, Nasher eta/. (2017), Benda, Spitzenberger eta/. (2014), Bernard (1983), Bernard & Happold (2013a), Csorba et al. (2003), Dulie & Mutere (1974), enton et al. (1977), Finger et al. (2017), Hackett et al. (2017), Jacobs, Barclay et al. (2007), Jacobs, Catto et al. (2017), Linden et al. (2014), Monadjem, Reside & Lumsden (2007), Monadjem, Schoeman et al. (2010), Monadjem, Shapiro et al. (2017), Monadjem, Taylor, Jacobs, Kock et al. (2017), Nader (1982), Rautenbach eta/. (1993), Richards eta/. (2016), Schoeman & Jacobs (2003), Stoffberg et al. (2012), Taylor (1999, 2000), Taylor, Sowler et al. (2013), Wessels & van der Merwe (1997), Whitaker et al. (1994), Wingate (1986).

Figures

On following pages: 27. African Forest Horseshoe Bat (Rhinolophus silvestris); 28. Upland Horseshoe Bat (Rhinolophus hillorum); 29. Sakeji Horseshoe Bat (Rhinolophus sakejiensis); 30. Bokhara Horseshoe Bat (Rhinolophus bocharicus); 31. Greater Horseshoe Bat (Rhinolophus ferrumequinum); 32. Geoffrey's Horseshoe Bat (Rhinolophus clivosus); 33. Greater Japanese Horseshoe Bat (Rhinolophus nippon); 34. Horacek's Horseshoe Bat (Rhinolophus horaceki); 35. Maclaud's Horseshoe Bat (Rhinolophus maclaudi); 36. Ziama Horseshoe Bat (Rhinolophus ziama); 37. Hill's Horseshoe Bat (Rhinolophus hilli); 38. Kahuzi Horseshoe Bat (Rhinolophus kahuzi); 39. Willard's Horseshoe Bat (Rhinolophus willardi); 40. Ruwenzori Horseshoe Bat (Rhinolophus ruwenzorii); 41. Chinese Horseshoe Bat (Rhinolophus xinanzhongguoensis); 42. Big-eared Horseshoe Bat (Rhinolophus macrotis); 43. Osgood's Horseshoe Bat (Rhinolophus osgoodi); 44. Allen's Horseshoe Bat (Rhinolophus episcopus); 45. Thai Horseshoe Bat (Rhinolophus siamensis); 46, Schnitzler's Horseshoe Bat (Rhinolophus schnitzten); M. King Horseshoe Bat (Rhinolophus rex); 48. Marshall’s Horseshoe Bat (Rhinolophus marshalli).

Subspecies and Distribution. R. c.c clivosus Cretzschmar, 1828 - SWJordan, W & C Saudi Arabia , Yemen , and SW Oman . R c. acrotis Heuglin, 1862 — Eritrea , Ethiopia , Djibouti , and N Somalia . R c. augur ÌL. Andersen, 1904 — N & C South Africa . R c. brachygnathus K. Andersen, 1905 - Israel , Egypt , and N Sudan . R c. geoffmyii A. Smith, 1829 — SW South Africa . R c. keniensis Hollister, 1916 - SE Sudan , South Sudan , Uganda , NE DR Congo , Rwanda , Burundi , Kenya , and N Tanzania ; other records throughout C Africa need further investigation. R c. schwarzi Heim de Balsac, 1934 - SE Algeria and W Libya . 7t c., socotranus Benda, Reiter & Vallo, 2017 - Socotra I, Yemen . AE c. zambesiensis K. Andersen, 1904 - S Tanzania, Malawi , Zambia, and SE DR Congo S to NE South Africa. R c. zuluensis K. Andersen, 1904 - E & S South Africa , Swaziland , and Lesotho . There is also a record from W DR Congo with no subspecific affinity and populations from Namibia and SW Angola are not currently assigned to any subspecies but might be associated with subspecies augur or geoffroyii following further morphological and genetic tests.