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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."]}

4. Lander’s Horseshoe Bat Rhinolophus landeri French: Rhinolophe de Lander / German: Lander-Hufeisennase / Spanish: Herradura de Lander Taxonomy. Rhinolophus landeri Martin, 1838, “ Insulâ Fernando Po [= Bioko Island],” Equatorial Guinea. Rhinolophus landeri is in the landeri species group. Rhinolophus lobatus was previously included in R landeri as a subspecies, but recent studies have determined that the two taxa are morphologically and genetically distinct and do not cluster together in phylogenetic reconstructions. Although exact distributions of R landeri and R lobatus are uncertain, arrangement suggested by P. J. Taylor and colleagues in 2018 is followed here, in which all eastern and southern African populations are assigned to R lobatus and all western and central African populations are assigned to R landeri. Monotypic. Distribution. From S Mauritania and Senegal through most ofW & C Africa to S Sudan, South Sudan, and Ethiopia, including Bioko I. Distributional limits are still uncertain. Descriptive notes. Head-body c. 52-60 mm, tail 27 mm, ear 13-20 mm, hindfoot 7 9 mm, forearm 35-49 mm; weight 5-11 g. Pelage is dense and soft, being brownish fawn to grayish brown (gray morph) or golden brown to bright rusty brown (orange morph) dorsally and slightly paler brown (gray morph) or lighter golden brown (orange morph) ventrally. Males often have dark reddish brown to dark reddish axillary tufts that secret sticky yellow substance. Wings and interfemoral membrane are dark grayish brown to blackish brown. Ears are short (34-42% of forearm length). Noseleaf has hastate lancet with pointed tip, being shorter than in Peters’s Horseshoe Bat (A lobatus); connecting process is subtriangular with tip either sharply or blundy pointed, being less erect than in Peters’s Horseshoe Bat; sella is naked and narrow, with slightly concave sides and top being broad and rounded; horseshoe is narrow at 6-8 mm but covers entire muzzle; there are no lateral leaflets; and median emargination is deep notch. Lower lip has well-defined medium groove and two very poorly defined lateral grooves. In general, baculum is shorter with distinctly bulbous tip that in Peters’s Horseshoe Bat Skull is of medium build; zygomatic width is slighdy greater than mastoid width; nasal swellings are narrow compared with Peters’s Horseshoe Bat; braincase is elongated compared with Peters’s Horseshoe Bat; sagittal crest is low to moderately developed anteriorly; and frontal depression is usually shallow. P2 is positioned in tooth row because of its relatively large size, which results in larger space between C1 and P4; P3 is small and slighdy to completely displaced labially; P2 and P4 are separated by narrow gap or are in contact; and P2 is only a litde smaller than P 4. Chromosomal complement has 2n = 58 and FNa = 64 (Senegal). Habitat. Various habitats including lowland and montane primary rainforests, disturbed forests, and riverine woodlands from sea level to elevations of c. 1800 m. Lander’s Horseshoe Bats have been recorded in lowland rainforest and degraded forests in West Africa, occurring in Isoberlinia (Fabaceae) woodland and often near rivers and riverine woodlands throughout their distribution. In Ethiopia, they have been recorded at elevations of 515-1800 m. They also occur in montane vegetation at 1400 m on Mount Cameroon in Cameroon and primary rainforest at 1000 m on Mount Bintumani in Sierra Leone. Food and Feeding. Lander’s Horseshoe Bat is insectivorous. It is a slow hawker that often forages within 6 m of the ground at night. Although fly-catching and gleaning have not been reported, they are expected to occur. Moths and beetles seem to make up a large proportion of diets. In Sudan, small beetles made up a large proportion of diets. Breeding. Lander’s Horseshoe Bat is seasonally monoestrous throughout its distribution, copulating in dry seasons. In Nigeria, copulation generally occurs around November but probably into December because ovulation occurs in late November. Implantation is delayed for c.2 months and does not occur until late dry season around February. Births often occur in early wet season but vary throughout the distribution, being in late April in Nigeria. Lactation in Nigeria occurred in April-June. Litters have a single young. Weaning occurs at c.2 months ofage, and for a short period of the early life ofyoung, mothers carry them when they forage. Activity patterns. Lander’s Horseshoe Bats are active during parts of the night. They roost during the day in colonies or, less commonly, alone and can enter torpor to save energy. Lander’s Horseshoe Bats have great maneuverability and can take offfrom the ground. They are able to fly slowly with bursts of speed and are able to briefly hover. Roosts are commonly found in caves, mine shafts, and crevices in rocks, although some individuals have been found in trees (particularly hollow baobabs, Adansonia, Malvaceae), thatched buildings, wells, and other holes in the ground. Call shape is FM/CF/FM, with F component of 103-3 kHz in Guinea, 108-5 kHz (one male) and 105-6 kHz (one female) in Burkina Faso, and 104-3 kHz in Liberia; average F component seems to be lower than in Peters’s Horseshoe Bat. Movements, Home range and Social organization. Lander’s Horseshoe Bats roost alone and in groups of more than 1000 individuals, being largely gregarious. In Nigeria, a cave with more than 1000 individuals has been recorded. Males and females usually are found in the same caves, but before birth, males leave the colony. They often share day roosts with a variety of other bats, including species of Lissonycteris, Nycteris, Coleura, Hipposideros, and other species of Rhinolophus. Status and Conservation. Classified as Least Concern on The IUCN Red List. Lander’s Horseshoe Bat is widespread and relatively common with no major known conservati ^ ® threats. Bibliography. ACR (2018), Aggundey & Schütter (1984), Brown & Dunlop (1997), Csorba et al. (2003), Dool, Puechmaille, Foley et al. (2016), Happold, M. (2013s), Hayman et al. (1966), Kangoyé et al. (2015), Koopman (1975,1989), Koopman, Kofron & Chapman (1995), Koopman, Mumford & Heisterberg (1978), Menzies (1973), Monadjem, Taylor, Jacobs & Cotterill (2017 b), Rosevear (1965), Taylor (1998), Taylor, Macdonald et al. (2018)., Published as part of Burgin, Connor, 2019, Rhinolophidae, pp. 280-332 in Handbook of the Mammals of the World – Volume 9 Bats, Barcelona :Lynx Edicions on page 282, DOI: 10.5281/zenodo.3748525

Rhinolophus landeri Martin, 1838 During this study, this species was recorded from forested habitat in Liberian Nimba at altitudes ranging from 550 m to 1,000 m. Despite its wide distribution (Monadjem et al., 2010 a), this species had not been recorded from Mt Nimba prior to this survey. This species closely resembles R. guineensis but was distinguished from it by its smaller size (forearm and greatest skull lengths were 42.0– 43.9 mm and 18.8–18.9 mm, n = 3). Rhinolophus alcyone is typically far larger in size., Published as part of Monadjem, Ara, Richards, Leigh & Denys, Christiane, 2016, An African bat hotspot: the exceptional importance of Mount Nimba for bat diversity, pp. 359-375 in Acta Chiropterologica 18 (2) on page 366, DOI: 10.3161/15081109ACC2016.18.2.005, http://zenodo.org/record/3942481

Rhinolophus landeri Martin, 1837 New material ZFMK 2008.0282 +0283, ♀♀, W1, 3 March 2008; ZFMK 2009.0021, ♂, PF, 9 December 2008. A total of four individuals of this common West African rhinolophid were captured, two over the creek in the W1 ravine, one at PF, and one in the WSV. One female caught at W1 on 3 March 2008 carried a 23 mm long embryo (crown-rump length). Small aggregations of this bat are most likely roosting together with other species in the cliffs, caves, and hollow trees found at the top of the W1 ravine and in the rocks on the west side of the hilltop at PF, which had already been partially clear-cut and mark- ed for bulldozing when we re-visited it in December 2008. One individual of this species was caught in a harp trap during the Pic de Fon RAP at the Banko forest site (Fahr and Ebigbo, 2004). This species roosts in caves, abandoned mines, and occasionally hollow trees or roofs (Fahr, 1996; Happold, 2013 b). At Mount Cameroon R. landeri was found at elevations up to 1,250 m (Fedden and McLeod, 1986). Conservation status Least Concern. Population trend is unknown (IUCN, 2015)., Published as part of Decher, Jan, Hoffmann, Anke, Schaer, Juliane, N Orris, Ryan W., Kadjo, Blaise, Astrin, Jonas, Monadjem, Ara & Hutterer, Rainer, 2015, Bat diversity in the Simandou Mountain Range of Guinea, with the description of a new white-winged vespertilionid, pp. 255-282 in Acta Chiropterologica 17 (2) on page 264, DOI: 10.3161/15081109ACC2015.17.2.003, http://zenodo.org/record/3943621, {"references":["FAHR, J. 1996. Die Chiroptera der Elfenbeinkuste (unter Berucksichtigung des westafrikanischen Raumes): Taxonomie, Habitatpraferenzen und Lebensgemeinschaften. Diploma Thesis, Julius-Maximilians-Universitat, Wurzburg, 204 pp. + 36 maps.","HAPPOLD, M. 2013 b. Rhinolophus landeri Lander's horseshoe bat. Pp. 340 - 341, in The mammals of Africa, Volume IV: hedgehogs, shrews and bats (M. HAPPOLD and D. C. D. HAP- POLD, eds.). Bloomsbury Publishing, London, 800 pp.","IUCN 2015. The IUCN Red List of Threatened Species. Version 2014.1. Available at http: // www. iucnredlist. org. Downloaded on 12 June 2015."]}

Rhinolophus landeri Martin 1838 This species was recorded at three sites in central and northern Mozambique. The male had bright orange axillary tufts in the armpits. Peak echolocation frequencies were 102.2 kHz (female; ANABAT, n = 1) and 104 kHz (male; Pettersson D240x, n = 1) 3–5 kHz lower than in South Africa (Schoeman and Jacobs, 2008; Monadjem et al., 2010) Field measurements: FA (adult male) 46.3 (1); Bm (adult male) 9.5 (1); FA (adult female) 46.3 (1); Bm (adult female) 10.0 (1). Nose-leaf width was 7.9 for the single female., Published as part of Monadjem, Ara, Schoeman, M. Corrie, Reside, April, P Io, Dorothea V., Stoffberg, Samantha, Bayliss, Julian, (Woody) Cotterill, F. P. D., Curran, Michael, Kopp, Mirjam & Taylor, Peter J., 2010, A recent inventory of the bats of Mozambique with documentation of seven new species for the country, pp. 371-391 in Acta Chiropterologica 12 (2) on page 378, DOI: 10.3161/150811010X537963, http://zenodo.org/record/3944583, {"references":["SCHOEMAN, M. C., and D. S. JACOBS. 2008. The relative influence of competition and prey defenses on the phenotypic structures of insectivorous bat ensembles in southern Africa. PLoS ONE 3 (11): e 3715. doi: 10.1371 / journal. pone. 0003715.","MONADJEM, A., P. J. TAYLOR, F. P. D. COTTERILL, and M. C. SCHOEMAN. 2010. Bats of Southern and Central Africa: a biogeographic and taxonomic synthesis. University of the Witwatersrand, Johannesburg, 596 pp."]}

Rhinolophus landeri Martin, 1838. Proc. Zool. Soc. Lond., 1837:101 [1838]. TYPE LOCALITY: Equatorial Guinea, Bioko. DISTRIBUTION: Senegal to Ethiopia and Somalia, south to South Africa and Namibia; Bioko; Zanzibar. SYNONYMS: angolensis, axillaris, dobsoni, lobatus. COMMENTS: Includes angolensis, dobsoni, and guineensis, but not brockmani according to Hayman and Hill (1971) and Koopman (1975:388); but see Böhme and Hutterer (1979:306-307) who correctly treated guineensis as a separate species., Published as part of Karl F. Koopman, 1993, Order Chiroptera, pp. 137-241 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 166, DOI: 10.5281/zenodo.7353060, {"references":["Hayman, R. W., and J. E. Hill. 1971. Order Chiroptera. Part 2. Pp. 1 - 73, in The mammals of Africa: An identification manual (J. Meester and H. W. Setzer, eds.) [issued 15 Jul 1971]. Smithsonian Institution Press, Washington, D. C., not continuously paginated.","Koopman, K. F. 1975. Bats of the Sudan. Bulletin of the American Museum of Natural History, 154: 353 - 444.","Bohme, W., and R. Hutterer. 1979. Kommentierte Liste einer Saugetier-Aufsammlung aus dem Senegal. Bonner Zoologische Beitrage, 29: 303 - 323."]}

Two species of leaf-nosed bats were netted as they emerged from their roosting cave at dusk. Collecting was repeated four times in 1966 and twice in 1967. Color of pelage, age, and reproductive condition of the bats were noted. From these data I conclude that the bats copulate late in the year, undergo a brief period of delayed implantation, give birth about April, and molt between May and July. The natural hair color of the adult is gray but quickly changes to reddish gold. Association between the breeding cycle and climate of the region is suggested.

Rhinolophus landeri Martin, 1838. Proc. Zool. Soc. Lond., 1837:101. TYPE LOCALITY: Equatorial Guinea, Bioko. DISTRIBUTION: Senegal to Ethiopia, Somalia and Kenya, south to South Africa; Namibia; W. Angola; Bioko; Zaire; Zanzibar. COMMENT: Includes angolensis and guineensis, but not brockmani; see Hayman and Hill, 1971, Part 2; but also see Bohme and Hutterer, 1979, Bonn. Zool. Beitr., 29:306, 307; and Koopman, 1975, Bull. Am. Mus. Nat. Hist., 154:388, who also included dobsoni. ISIS NUMBER: 5301405007003033001., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Chiroptera, pp. 111-215 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on pages 144-145, DOI: 10.5281/zenodo.7352990, {"references":["Hayman, R. W., and J. E. Hill. 1971. Order Chiroptera. Part 2. Pp. 1 - 73, in The mammals of Africa: An identification manual (J. Meester and H. W. Setzer, eds.) [issued 15 Jul 1971]. Smithsonian Institution Press, Washington, D. C., not continuously paginated.","Bohme, W., and R. Hutterer. 1979. Kommentierte Liste einer Saugetier-Aufsammlung aus dem Senegal. Bonner Zoologische Beitrage, 29: 303 - 323.","Koopman, K. F. 1975. Bats of the Sudan. Bulletin of the American Museum of Natural History, 154: 353 - 444."]}

20. Rhinolophus landeri. Rhinolophus landeri, Martin, P. Z. S. 1837, p. 101; Peters, MB. Akad. Beri. 1871, p. 308. Ears shorter than the head, not attenuated at their extremities, the inner margin of the ear-conch convex to the tip; antitragus short, evenly convex, separated posteriorly by a very shallow notch; nose �� leaf about the same size as in Rh. affinis, the horizontal leaf broader than in Rh. euryale, but not concealing the muzzle; vertical process of the sella slightly narrowed in the centre, broader and rounded-off above, exceeded in height by the upper margin of the posterior connecting-process, which forms an angular elevation more or less acute, but much broader at its base and much shorter than in Rh. euryale; posterior leaf rather short and acutely pointed (Plate VII. fig. 9). Wings from the ankles or from the tibia close to the ankle-joint. Interfemoral membrane short, slightly triangular behind. Fur varying from deep ferruginous brown, above and beneath, to sulphur or pale yellowish brown. The first upper premolar blunt, in the tooth-row, but closely appressed between the canine and second premolar. Length (of the type specimen), head and body 1 "" 6, tail 0������85, ear 0''���65, nose-leaf 0',-45 X O''-28, forearm "7, thumb 0"���28; third finger���metacarp. " ���15, 1st ph. 0"-5, 2nd ph. O' '-9; fifth fingermetacarp. l''-2, 1st ph. 0"-3, 2nd ph. 0���-55; tibia 0"-65, foot 0"-3. Hah. Africa (Femando-Po Island, Gaboon, Damara Land). a. ♀ ad., al. (type). Fernando Po. Zool. Soc. Coll. b. ♀ ad., al. Elloby district, Gaboon. H. A. Ansell, Esq. c. ♀ ad., al. Dr. Livingstone [C.]., Published as part of Dobson, George Edward, 1878, Rhinolophus, pp. 100-122 in Catalogue of the Chiroptera in the collection of the British Museum, London :British Museum on pages 118-119, DOI: 10.5281/zenodo.3758443

Rhinolophus landeri Martin Rhinolophus landeri Martin, Proc. Zool. Soc. London, for 1837, p. 101, 25 May 1838. Fernando Po., Published as part of Allen, Glover M., 1939, Family Rhinolophidae. Horseshoe Bats in Checklist of African Mammals, pp. 72-78 in Bulletin of the Museum of Comparative Zoology at Harvard College 81 on page 75, DOI: 10.5281/zenodo.3759519

Between 10 January and 14 February 1976, activity patterns, habitat use, and selection of prey by some insectivorous bats were studied in mopane and brachystegia deciduous woodlands in the Sengwa Wild Life Research Area of the Hostes Nicolle Institute of Wild Life Research in Rhodesia (18?10' S, 23013' E), using ultrasonic (=bat) detectors, light tags, and analysis of insect remains from bat feces. Some bats were active throughout the night, in part feeding, but in general bat activity was reduced from around midnight until just before dawn. Rain tended to suppress bat activity, although the timing of the rain was important. Since insects remained active in the rain, we suspect that the responses of the bats to rain reflected problems of thermoregulation associated with wet fur, and the effect of multiple echoes and attenuation of high-frequency sound on echolocation. Bright moonlight suppressed bat activity and altered the foraging patterns of light-tagged Scotophilus viridis, Eptesicus capensis, and Nycticeius schlieffeni which on dark nights fed above the canopy and along the grassland sides of meadow/woodland interfaces, but which were mainly restricted within the woodland on bright moonlight nights. Since a pair of bat hawks (Macheiramphus alcinus) nested near the study area and fed mainly on bats, we suggest that the bats' responses to moonlight are predator-avoidance behaviors. S. viridis (15 20 g), E. capensis (5 6 g), and N. schlieffeni (5 6 g) appeared to feed on concentrations of insects, and responded quickly to aggregations of insects around lights. These bats fed mainly on beetles (S. viridis-85%%; E. capensis-68%; N. schlieffeni-72%), although E. capensis also took moths (18.5%) and N. schlieffeni took flies (19%). Other insectivorous bats in the study area fed mainly on moths (Hipposideros caffer, Laephotis angolensis, Ga4uconycteris variegata) or beetles (Scotophilus nigrita), or both (Rhinolophus hildebrandti, Pipistrellus nanus). Data for some insectivorous bats from the vicinity of Salisbury, Rhodesia, indicated a similar separation for some bats (moths-Nycteris thebaica, Rhinolophus landeri; beetles-Rhinolopus clivosus) in addition to two species of Miniopterus which took mainly aquatic Diptera. The data from this study, with those from other works, are used to illustrate partitioning of insect prey by the species of bats known to occur in the Sengwa Wild Life Research Area. KNOWLEDGE OF THE PARTITIONING of available resources is central to the understanding of the ecological interactions between sympatric species of animals. While substantial data are available for some birds, in terms of what food is consumed and where (e.g, MacArthur 1958, Lack 1966. Keast 1968). this information is not available for most bats. There are reports of the insects eaten by different bats (e.g.. Ross 1967; Black 1972, 1974; Whitaker and Black 1976), but these are usually based on analysis of stomach contents or feces, and do not indicate where the bats had been feeding. Other studies (e.g., Kunz 1973) have shown that different insectivorous bats have different periods of activity that may be related