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(1=author & date in parentheses)	Citation	Pages	Common Name	Synonyms	Type Locality	Distribution	CITES	IUCN	Comments	column3781	column3791	subtribe	CONCAT_ALTNAMES
line:xlsx:hash://sha256/181a039844a33e66a35a457b7ece741051086608e425a040051b79581d606b97!/Sheet1!/L1375	application/vnd.openxmlformats-officedocument.spreadsheetml.sheet	Rhinolophus ferrumequinum	Rhinolophus ferrumequinum	Rhinolophus ferrumequinum	Rhinolophus ferrumequinum	Rhinolophus ferrumequinum	Rhinolophus ferrumequinum	Rhinolophus ferrumequinum	Rhinolophus ferrumequinum	Rhinolophus ferrumequinum	Rhinolophus ferrumequinum	Rhinolophus ferrumequinum	Rhinolophus ferrumequinum	Rhinolophus ferrumequinum	Rhinolophus ferrumequinum	Rhinolophus ferrumequinum		[MSW2] Revised by Strelkov et al. (1978).; [MSW3] ferrumequinum species group. Revised by Strelkov et al. (1978). Reviewed in part by Yoshiyuki (1989), Harrison and Bates (1991), Paz (1995), Kock (1996), Bates and Harrison (1997), Sinha (1999), Zagorodnyuk (1999), Horácek et al. (2000) and Gaisler (2001a). Subspecies limits are somewhat unclear and there may be more than one species present in this complex; see discussion in Csorba et al. (2003).; [HMW] Vespertilioferrum-equinum Schreber, 1774 , France . Rhinolophusferrumequinum is in the ferrumequinum species group with the extinct species R maghrebensis and R mellali . The ferrumequinum group is included in the Afro-Palearctic clade of Rhinolophus close to the maclaudi , fumigatus , and xinanzhongguoensis groups. Rhinolophusferrumequinum is sister to R clivosus , although some Egyptian specimens attributed to R clivosus cluster within R ferrumequinum . Rhinolophus nippon was previously included as a subspecies of R ferrumequinum , but it is genetically sister to the clade including R ferrumequinum and R clivosus . Exact distributional limit between R nippon and R ferrumequinum is currendy uncertain due to lack of genetic data from specimens in central and southern Asia, and thus, distributional difference here is tentatively marked at Kashmir. Subspecies creticumis, now considered a synonym of the nominate form whereas subspecies Irani is now a synonym of proximus, although genetic tests have not been performed on populations from central and southern Asia. Two subspecies recognized.; [batnames2022]  ferrumequinum species group. Does not include nippon , tragatus or korai ; see Thomas (1997), Zhou et al. (2009), Benda and Vallo (2012), Burgin (2019), and Ikeda et al. (2020). Revised by Strelkov et al. (1978). Reviewed in part by Yoshiyuki (1989), Harrison and Bates (1991), Paz (1995), Kock (1996),Bates and Harrison (1997), Sinha (1999), Zagorodnyuk (1999), Horácek et al. (2000) and Gaisler (2001a). Subspecies limits are somewhat unclear and there may be more than one species present in this complex; see discussion in Csorba et al. (2003). For a review of the species in Iran see Shahabi et al. (2019).; [MDD2022] previously included R. nippon; [IUCN] Although Thomas (1997, unpublished thesis) found very high divergence in mitochondrial DNA sequences, Csorba et al. ;(2003) refrained from splitting Japanese Greater Horseshoe Bats (Rhinolophus nippon ) from R. ferrumequinum . Several subspecies are identified over the range, of which two occur in the western Palaearctic: R. f. creticus (Crete) and R. f. ferrumequinum (rest of western Palaearctic range).; [batnames2023]  ferrumequinum species group. Does not include nippon , tragatus or korai ; see Thomas (1997), Zhou et al. (2009), Benda and Vallo (2012), Burgin (2019), and Ikeda et al. (2020). Revised by Strelkov et al. (1978). Reviewed in part by Yoshiyuki (1989), Harrison and Bates (1991), Paz (1995), Kock (1996),Bates and Harrison (1997), Sinha (1999), Zagorodnyuk (1999), Horácek et al. (2000) and Gaisler (2001a). Subspecies limits are somewhat unclear and there may be more than one species present in this complex; see discussion in Csorba et al. (2003). For a review of the species in Iran see Shahabi et al. (2019).; [MDD2023] previously included R. nippon; [MDD2025_2.0] previously included R. nippon; [batnames2025_1.7] ferrumequinum species group. Does not include nippon, tragatus or korai; see Thomas (1997), Zhou et al. (2009), Benda and Vallo (2012), Burgin (2019), and Ikeda et al. (2020). Revised by Strelkov et al. (1978). Reviewed in part by Yoshiyuki (1989), Harrison and Bates (1991), Paz (1995), Kock (1996),Bates and Harrison (1997), Sinha (1999), Zagorodnyuk (1999), Horácek et al. (2000) and Gaisler (2001a). Subspecies limits are somewhat unclear and there may be more than one species present in this complex; see discussion in Csorba et al. (2003). For a review of the species in Iran see Shahabi et al. (2019).; [MDD2025_2.2] previously included R. nippon						brevitarsus, colchicus, creticum, equinus, fudisanus, germanicus, hippocrepis, homodorensis, homorodalmasiensis, insulanus, irani, italicus, korai, kosidanus, martinoi, mikadoi, nippon, norikuranus, obscurus, ogasimanus, proximus, quelpartis, regulus, rubiginosus, typicus, tragatus, ungula, unihastatus.	ferrumequinum, creticus, proximus, tragatus, nippon	ferrumequinum, creticum, irani, korai, nippon, proximus, tragatus	colchicus, equinus, germanicus, hippocrepis, homodorensis, homorodalmasiensis, insulanus, italicus, major, martinoi , obscurus, perspicillatus, solea, typicus, ungula, unihastatus; irani - rubiginosus; korai - pachyodontus, quelpartis; nippon - fudisanus, kosidianus, mikadoi, norikuranus, ogasimanus; tragatus - brevitarsus, regulus	ferrumequinum, proximus	creticumis; proximus - irani	ferrumequinum, creticum, irani, korai, nippon, proximus, tragatus	ferrumequinum - colchicus, equinus, germanicus, hippocrepis, homodorensis, insulanus, italicus, major, martinoi, obscurus, perspicillatus, solea, typicus, ungula, unihastatus; irani - rubiginosus; korai - pachyodontus, quelpartis; nippon - fudisanus, kosidianus, mikadoi, norikuranus, ogasimanus; tragatus - brevitarsus, regulus	ferrumequinum, equinus, solea, perspicillatus, ungula, major, hippocrepis, major, unihastatus, germanicus, italicus, homorodalmasiensis, homodorensis, obscurus, proximus, typicus, insulanus, colchicus, irani, martinoi, rubiginosus, creticum	Although Thomas (1997, unpublished thesis) found very high divergence in mitochondrial DNA sequences, Csorba et al. ;(2003) refrained from splitting Japanese Greater Horseshoe Bats (Rhinolophus nippon ) from R. ferrumequinum . Several subspecies are identified over the range, of which two occur in the western Palaearctic: R. f. creticus (Crete) and R. f. ferrumequinum (rest of western Palaearctic range).	ferrumequinum, creticum, irani, proximus	ferrumequinum - colchicus, equinus, germanicus, hippocrepis, homodorensis, insulanus, italicus, major, martinoi, obscurus, perspicillatus, solea, typicus, ungula, unihastatus; irani - rubiginosus	ferrumequinum, equinus, solea, perspicillatus, ungula, major, hippocrepis, major, unihastatus, germanicus, italicus, homorodalmasiensis, homodorensis, obscurus, proximus, typicus, insulanus, colchicus, irani, martinoi, rubiginosus, creticum	ferrumequinum, equinus, solea, perspicillatus, ungulus, major, hippocrepis, ferumequinum, major, unihastatus, unifer, libanoticus, germanicus, italicus, rufescens, conchifer, homorodalmasiensis, homodorensis, obscurus, proximus, typicus, insulanus, colchicus, irani, martinoi, rubiginosus, mellali, topali, creticum, tarkoensis, antiquis, creticus	creticus, ferrumequinum, irani, proximus	ferrumequinum - colchicus, equinus, germanicus, hippocrepis, homodorensis, homorodalmasiensis, insulanus, italicus, major, martinoi, obscurus, perspicillatus, solea, typicus, ungula, unihastatus; irani - rubiginosus	ferrumequinum (von Schreber, 1774)|equinus (P. L. S. Müller, 1776)|solea (E. A. W. von Zimmermann, 1777) [placed on index]|perspicillatus (Blumenbach, 1779) [preoccupied]|solea (E. A. W. von Zimmermann, 1783) [nomen nudum]|ungulus (Boddaert, 1785)|major (Kerr, 1792) [preoccupied]|hippocrepis (von Schrank, 1798) [nomen novum]|ferumequinum (Bechstein, 1801) [incorrect subsequent spelling]|major É. Geoffroy Saint-Hilaire, 1803 [preoccupied]|unihastatus É. Geoffroy Saint-Hilaire, 1813|unifer Kaup, 1829 [nomen nudum]|libanoticus Temminck, 1835 [nomen nudum | not used as valid]|germanicus C. Koch, 1863|italicus C. Koch, 1863|rufescens Fitzinger, 1870 [nomen nudum]|conchifer W. C. H. Peters, 1871 [nomen nudum | not used as valid]|homorodalmasiensis Daday, 1885 [nomen nudum]|homodorensis Daday, 1887 [nomen novum]|obscurus Cabrera, 1904|proximus Andersen, 1905|typicus Andersen, 1905 [nomen novum]|insulanus Barrett-Hamilton, 1910|colchicus Satunin, 1912|irani Cheesman, 1921|martinoi B. Petrov, 1941|rubiginosus Gubareff, 1941|mellali Lavocat, 1961|topali M. Kretzoi, 1977|creticum J. Iliopoulou-Georgudaki, 1979 [nomen nudum]|tarkoensis Topál, 1979|antiquis M. Kretzoi in Jánossy, 1986 [nomen nudum]|creticus J. Iliopoulou-Georgudaki & Ondrias, 1986		Corbet, G.B. and Hill, J.E. 1980. A World List of Mammalian Species. British Museum (Natural History), London, 226 pp.	Greater horseshoe bat	Britain, Morocco – N India, Japan	Honacki, J.H., Kinman, K.E. and Koeppl, J.W. 1982. Mammal Species of the World: A Taxonomic and Geographic Reference. Allen Press, Lawrence, 694 pp.	Rhinolophus ferrumequinum	France.	Schreber	1774	Saugethiere, 1:174, pl. 62.	Distribution: Widely distributed ac ross temperate Eurasia from Britain to Japan south to northwestern Africa, Palestine and nor thern India.		Corbet, G.B. and Hill, J.E. 1991. A World List of Mammalian Species. Third edition. Oxford University Press, London, 243 pp. ISBN 0-19-854017-5	Greater horseshoe bat	Britain, Morocco – N India, Japan	Koopman, K.F. 1993. Order Chiroptera. Pp. 137–242 in Wilson, D.E. and Reeder, D.M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference. Second edition. Smithsonian Institution Press, Washington, 1206 pp.	Schreber	1774	Die Saugethiere, 1:174, pl. 62.	Revised by Strelkov et al. (1978).	S England to Caucasus Mtns south to Morocco and Tunesia (but not Egypt) through Iran and Himalayas to China and Japan; adjacent small islands.	France.		SCHREBER	1774	Metacarpals of third and fourth digits relatively short. Sella pandurate. Horseshoe relatively narrow. Anterior upper premolar extru ded from toothrow, greatly reduced or absent. Anterior nasal swellings fairly small. Periotic bo nes not enlarged. Size fairly large (forearm length, 52-63 mm).	Distribution: Widely distributed ac ross temperate Eurasia from Britain to Japan south to northwestern Africa, Palestine and nor thern India.	Six subspecies are here recognized:	R.f. ferrumequinum (Europe, northwestern Africa), R.f. creticus (Crete), R.f. proximus (= iraní) (southwestern Asia east to Kashmir), R. f. tragatus (= regulus) (northern India, southwestern China), R.f. nippon (northern and cen tral China, Korea, Japan).	54	species	R. ferrumequinum	SCHREBER	1774	Rhinolophus	genus	Rhinolophus ferrumequinum				Metacarpals of third and fourth digits relatively short. Sella pandurate. Horseshoe relatively narrow. Anterior upper premolar extru ded from toothrow, greatly reduced or absent. Anterior nasal swellings fairly small. Periotic bo nes not enlarged. Size fairly large (forearm length, 52-63 mm).	Six subspecies are here recognized:		21. R. ferrumequinum (SCHREBER 1774) [ferrume quinum group],	21	_R. f. creticus_ Iliopoulou-Georgudaki & Ondrias, 1986; _R. f. ferrumequinum_ (Schreber, 1774) (synonyms: _colchicus_ Сатунин, 1912, _conchifer_ Peters, 1871, _equinus_ (Müller, 1776), _germanicus_ Koch, 1863, _hippocrepis_ (Schrank, 1798), _homodorensis_ Daday, 1887, _homorodalmasiensis_ Daday, 1885, _insulanus_ Barrett-Hamilton, 1910, _italicus_ Koch, 1863, _libanoticus_ Temminck, 1835, _major_ (Kerr, 1792), _major_ Geoffroy Saint-Hilaire, 1803, _martinoi_ Petrov, 1941, _obscurus_ Cabrera, 1904, _perspicillatus_ (Blumenbach, 1779), _rufescens_ Fitzinger, 1870, _solea_ (Zimmermann, 1777), _solea_ (Zimmermann, 1783), _typicus_ Andersen, 1905, _ungulus_ (Boddaert, 1785), _unifer_ Kaup, 1829, _unihastatus_ Geoffroy Saint-Hilaire, 1813); _R. f. irani_ Cheesman, 1921 (synonyms: _rubiginosus_ Gubareff, 1941); _R. f. mellali_ Lavocat, 1961 (fossil); _R. f. proximus_ Andersen, 1905; _R. f. tarkoensis_ Topál, 1979 (fossil); _R. f. topali_ Kretzoi, 1977 (fossil)			Don E. Wilson & DeeAnn M. Reeder (editors). 2005. Mammal Species of the World. A Taxonomic and Geographic Reference (3rd ed), Johns Hopkins University Press, 2,142 pp. (Available from Johns Hopkins University Press, 1-800-537-5487 or (410) 516-6900, or at http://www.press.jhu.edu).	CHIROPTERA	Rhinolophidae			Rhinolophus ferrumequinum	Rhinolophus		ferrumequinum	Schreber	y	1774		Die Säugethiere	1		174, pl. 62		Greater Horseshoe Bat	France.	Algeria, Morocco, and Tunisia; S Europe from Portugal to Greece and north to S England, the Netherlands, S Germany, Austria, Czech Republic, Slovakia, and Bulgaria; Turkey, Cyprus, Georgia, and Azerbaijan; Urkrain, Crimea, and Caucacus regions; the Mediterranean coast from Turkey to Israel and Jordan; NE Iraq, Iran, Turkmenistan, Uzbekistan, S Kazakhstan, Afganistan, Pakistan, N India, Nepal, Sikkim, China, Korea, and Japan; adjacent small islands. Records at some localities in northern Europe (e.g., the Netherlands) apparently reflect temporary northern range extensions (Glas and Voûte, 1992a).	IUCN 2003 and IUCN/SSC Action Plan (2001) – Lower Risk (nt).	colchicus Satunin, 1912; equinus Müller, 1776; germanicus Koch, 1865; hippocrepis Schrank, 1798; homodorensis Daday, 1887; homorodalmasiensis Daday, 1885 [nomen nudum]; insulanus Barrett-Hamilton, 1910; italicus Koch, 1865; major E Geoffroy, 1803 [not Kerr, 1792); major Kerr, 1792:99 [not Kerr, 1792:97]; martinoi Petrov, 1940; obscurus Cabrera, 1904; perspicillatus Blumenbach, 1779; solea Zimmermann, 1777 [unavailable; see Bull. Zool. Nomen. (1950)4:547]; typicus K. Andersen, 1905; ungula Boddaert, 1785; unihastatus E. Geoffroy, 1803; creticum Iliopoulou-Georgudaki and Ondrias, 1985; irani Cheesman, 1921; rubiginosus Gubareff, 1941; korai Kuroda, 1938; pachyodontus Kishida, 1931 [nomen nudum]; quelpartis Mori, 1933; nippon Temminck, 1835; fudisanus Kishida, 1940; kosidianus Kishida, 1940; mikadoi Ognev, 1927; norikuranus Kishida, 1940; ogasimanus Kishida, 1940; proximus K. Andersen, 1905; tragatus Hodgson, 1835; brevitarsus Blyth, 1863 [nomen nudum]; regulus K. Andersen, 1905.	ferrumequinum species group. Revised by Strelkov et al. (1978). Reviewed in part by Yoshiyuki (1989), Harrison and Bates (1991), Paz (1995), Kock (1996), Bates and Harrison (1997), Sinha (1999), Zagorodnyuk (1999), Horácek et al. (2000) and Gaisler (2001a). Subspecies limits are somewhat unclear and there may be more than one species present in this complex; see discussion in Csorba et al. (2003).	885887A2FFFC8A1DF84DEF93FABED367	Handbook of the Mammals of the World – Volume 9 Bats, Barcelona: Lynx Edicions	978-84-16728-19-0	hbmw_9_Rhinolophidae.pdf.imf	hash://md5/7461ffdaffcf8a29ffccffa1ff85d963	295	zip:hash://sha256/ec5fd314a06aba1a7b0b72f23e54ac625ae272bd98f82f1d01f4c09627d9e8e0!/treatments-xml-main/data/88/58/87/885887A2FFDF8A38F899FBD8F436D1A0.xml	Rhinolophus ferrumequinum	Rhinolophidae	Rhinolophus	ferrumequinum	Schreber	1774	Greater Horseshoe Bat @en | Grand Rhinolophe @fr | Grosse Hufeisennase @de | Herradura grande @es | German @en | Larger Horseshoe Bat @en | Other commonnames @en	Vespertilioferrum-equinum Schreber, 1774 , France . Rhinolophusferrumequinum is in the ferrumequinum species group with the extinct species R maghrebensis and R mellali . The ferrumequinum group is included in the Afro-Palearctic clade of Rhinolophus close to the maclaudi , fumigatus , and xinanzhongguoensis groups. Rhinolophusferrumequinum is sister to R clivosus , although some Egyptian specimens attributed to R clivosus cluster within R ferrumequinum . Rhinolophus nippon was previously included as a subspecies of R ferrumequinum , but it is genetically sister to the clade including R ferrumequinum and R clivosus . Exact distributional limit between R nippon and R ferrumequinum is currendy uncertain due to lack of genetic data from specimens in central and southern Asia, and thus, distributional difference here is tentatively marked at Kashmir. Subspecies creticumis, now considered a synonym of the nominate form whereas subspecies Irani is now a synonym of proximus, although genetic tests have not been performed on populations from central and southern Asia. Two subspecies recognized.	R f. ferrumequinum Schreber, 1774 - S Europe from Iberian Peninsula and France E to Romania , Bulgaria , and Greece including SW Great Britain , S Germany , S Poland , and Crimea, also on most Mediterranean Is (Balearics, Corsica , Sardinia, Sicily, Malta , Crete, Cyprus , etc.), Turkey ( Anatolia ), NW Africa (N Morocco , N Algeria , N Tunisia , and NW Libya ), and Levant region. R f. proximus K. Andersen, 1905 - Transcaucasia, Mesopotamia , S Turkmenistan , N, W & S Iran , Afghanistan , Tajikistan , Kyrgyzstan , Pakistan , and N India (Kashmir).			Greater Horseshoe Bats forage by slow hawking, fly-catching from perches, and ground-gleaning. When captured, insects are eaten on the wing or taken to a perch. They will sometimes land on the ground to capture dung beetles and other insects in dung piles. Most of their diet comes from species of Lepidoptera, Coleoptern , Hymenoptera , and Diptera (at least throughout Europe), but other arthropods also make up smaller proportions of their diets, including Neuroptera, Trichoptera, and Araneae . A large sample of 1580 feces found that the diet of the Greater Horseshoe Bat in the UK mostly contained Lepidoptera (moths) and changed throughout the year. In spring, they ate cockchafers {Melolontha melolonthd), beetles ( Geotrupe ), caddisflies (Trichoptera), tipulid flies, ichneumonid wasps, and moths. In autumn, dung beetles (Aphodius) and dung flies (various Diptera ) replaced various species of beetles in diets. Coleoptera, Lepidoptera , and Diptera dominated diets in Azerbaijan , and Coleoptera and Lepidoptera were most important in Turkey , Syria , and Jordan . Moths dominated three stomach samples from Iran , and some brachyceran Diptera, Trichoptera , and scarabaeid Coleoptera also were eaten. Greater Horseshoe Bats are versatile feeders and alter their diets based on prey availability throughout the year and among regions. Where they forage also can change; in the UK , they forage mainly in woodlands in spring and over pastures in late summer.	Greater Horseshoe Bats are seasonally monoestrous. Copulation occurs before hibernation; males mate with females in their harems. After mating, males secrete a plug into females’ vaginas to either prevent sperm competition or keep sperm stored in the female until the following spring. Sperm storage occurs in oviducts of females until ovulation and fertilization takes place in March—April. Gestation lasts 2—3 months depending on whether or not embryo development is lengthened due to torpor of the pregnant female; gestation is generally c .9 weeks with limited torpor. Young are bom in late spring or early summer depending on the region. In Algeria , young are bom inJune and volant by the end ofAugust. Litter size is one. Young hold onto their mothers until a little before weaning. Young will forage and hang separate from their mothers before weaning. Greater Horseshoe Bats mature very slowly, similar to many hibernating bats. Females reach sexual maturity at c .3 years of age, but they might not produce their first offspring until 3—5 years of age. Males generally mature at 2-4 years of age. Greater Horseshoe Bats are long-lived; maximum longevities recorded in the wild are 30 years and six months for a male and 27 years and eight months for a female.	Greater Horseshoe Bats are nocturnal and forage throughout the night They generally leave day roosts 15—30 minutes after sunset and begin to forage. They typically linger around day roosts for the early part of the night and fly further from the day roost as the night progresses. Most nightly activity occurs within 5—20 km ofroosts, and adults typically move further thanjuveniles. Greater Horseshoe Bats generally stay out all night or return after 2—3 hours and leave day roosts again for c .30 - 50 minutes before sunrise. They occasionally rest in night roosts. Activity decreases in cold temperatures, markedly below 10°C—the typical thermal threshold for insect activity. During the day, Greater Horseshoe Bats can enter a mild torpor when temperatures are below 22°C. They enter deep torpor (hibernation) from mid-autumn until spring (October-April in Europe), during which they find parts of caves that are warmer than 7 — 10 ° C. During this time, they continue to irregularly awake and leave roosts to forage when weather permits. They do not hibernate in southern parts of the distribution but have been found to hibernate in Iran . Day roosts of Greater Horseshoe Bats are generally in caves and other underground or rocky structures; they commonly inhabit abandoned or unused parts of buildings, particularly roofs. Greater Horseshoe Bats tend to favor houses in northern parts of their distribution but underground structures in the south. They also seem to favor caves as hibernacula in winter. Call shape is FM/ CF /FM, with terminal FM sweep usually having the greatest bandwidth. Aspects of calls vary among seasons and through the lifetime of an individual; juveniles emit lower frequencies than adults. Resting frequencies of mothers and young seem to be similar, indicating that the call is learned from the mother or inherited. F component is 77—83 kHz in southern Europe, 83—84 kHz in the UK , 81-7 kHz in Western Europe, and 84-6 kHz in Morocco . Mean call duration is 53-8 milliseconds in Greece , 21 milliseconds and 31-9 milliseconds in the UK , and 30-3 milliseconds in Morocco .	Greater Horseshoe Bats are highly gregarious, roosting singly, in small groups, or in very large colonies up to 1000 individuals, although they forage alone. Non-matemity colonies have been recorded with up to 500 individuals of both sexes; summer maternity colonies of up to 1000 females have been reported, although 100—300 individuals are more common. Maternity colonies are createdjust before females give birth. When first formed, maternity colonies can still include some persistent males and non-breeding individuals that usually leave after young are bom. Non-breeding individuals disperse like males or can remain in maternity colonies throughout the breeding season. Males form scattered small groups in separate day roosts. All male and all juvenile colonies have been recorded. Males and females start roosting together after young are weaned around late summer (at least in Europe), which leads to copulation in day roosts until the end of October before the beginning of hibernation (in northern populations). From late summer to about mid-autumn, males become territorial in roosting colonies, establishing small harems in the roost. Harems generally include the male and up to eight females, segregating into a single cluster in the roost Populations of Greater Horseshoe Bats are primarily sedentary and hibernate instead of migrating. Nevertheless , they often fly 20—35 km between summer roosts and hibernacula. They typically move to more secluded localities ( e .g. cool, deep parts of caves) where they can enter deep torpor. Individuals wrap themselves completely in their wing membranes during hibernation. Hibernacula generally include fewer individuals than summer roosts but can sometimes include clusters ofup to 100 individuals. Females seem to give birth in the same roosting area each year. Non-matemity summer roosts are often shared with other species of Rhinolophus , Miniopterus , Myotis , Asellia , and Plecotus . Maternity colonies are occasionally shared with other bat species.	Classified as Least Concern on The IUCNed List. The Greater Horseshoe Bat has a wide distribution and is rather abundant throughout much of its distribution. Nevertheless, there are areas that have experienced well-documented declines, including Malta , Belgium , and the Netherlands where it is likely extinct Much of the documented decline has occurred throughout Europe, notably in north-western Europe. Populations in the UK have experienced massive declines, and the overall population is now stable at a low level of c .5000 individuals. Austrian populations have declined by 70% in the last ten years and are down to only c.30 breeding individuals. Population trends throughout the rest of Europe and North Africa are uncertain, but roosts seem to be disappearing throughout the Iberian Peninsula. Populations are considered stable in Croatia . There apparently has been a slow increase in populations in Romania since 1989 due to the reduced use of pesticides. The Greater Horseshoe Bat is rare in Switzerland but considered stable. It is considered relatively rare throughout much of its Asiatic distribution, although it is considerably common throughout Transcaucasia. Primary threats are fragmentation and habitat destruction from deforestation and agricultural expansion. Use of pesticides also negatively affects Greater Horseshoe Bats by targeting some important food sources such as melolonthid beetles, noctuid moth larvae, and crane flies. General roost disturbance is also an evident threat, especially for colonies in buildings where human intolerance can make it difficult for cohabitation. The Greater Horseshoe Bat is widely protected throughout Europe, where many underground roosts have been protected and building roosts have had management agreements to maintain the bat-human relationship. Legislation protects the Greater Horseshoe Bat in some but not all countries in its distribution.	ACR (2018) | Aldridge (1986) | Arslan & Zima (2014) | Aulagnier &Thévenot (1986) | Bates & Harrison (1997) | Benda & Gaisler (2015) | Benda & Vallo (2012) | Benda, Abi-Said et al. (2016) | Benda, Andreas et al. (2006) | Benda, Faizolâhi et al. (2012) | Benda, Georgiakakis et al. (2008) | Benda, Hanâk & Cervenÿ (2011) | Benda, Ivanova et al. (2003) | Benda, Lucan et al. (2010) | Benda, Spitzenberger et al. (2014) | Botnariuc &Tatole (2005) | Csorba et al. (2003) | Disca et al. (2014) | landers, Jones et al. (2009) | landers, Wei Li et al. (2011) | Gaisler (2001, 2013b) | Gunnell et al. (2011) | Hanâk et al. (2001) | Jones (1990) | Jones & Ransome (1993) | Jones & Rayner (1989) | Jones & Siemers (2011) | Koh Hung-Sun et al. (2014) | Long & Schnitzler (1975) | Ma Jie, Kobayasi et al. (2006) | Park et al. (2000) | Piraccini (2016a) | Ransome & McOwat (1994) | Rossiter , Benda et al. (2007) | Rossiter, Jones et al. (2000a, 2000b, 2001) | Schnitzler (1973) | Schnitzler & Grinnell (1977) | Stoffberg et al. (2010) | Suga et al. (1976) | Vogler & Neuweiler (1983) | Walters et al. (2012)	https://zenodo.org/record/3750152/files/figure.png	31 . Greater Horseshoe Bat Rhinolophus ferrumequinum French: Grand Rhinolophe / German : Grosse Hufeisennase I Spanish: Herradura grande Other common names: Larger Horseshoe Bat Taxonomy. Vespertilioferrum-equinum Schreber, 1774 , France . Rhinolophusferrumequinum is in the ferrumequinum species group with the extinct species R maghrebensis and R mellali . The ferrumequinum group is included in the Afro-Palearctic clade of Rhinolophus close to the maclaudi , fumigatus , and xinanzhongguoensis groups. Rhinolophusferrumequinum is sister to R clivosus , although some Egyptian specimens attributed to R clivosus cluster within R ferrumequinum . Rhinolophus nippon was previously included as a subspecies of R ferrumequinum , but it is genetically sister to the clade including R ferrumequinum and R clivosus . Exact distributional limit between R nippon and R ferrumequinum is currendy uncertain due to lack of genetic data from specimens in central and southern Asia, and thus, distributional difference here is tentatively marked at Kashmir. Subspecies creticumis, now considered a synonym of the nominate form whereas subspecies Irani is now a synonym of proximus, although genetic tests have not been performed on populations from central and southern Asia. Two subspecies recognized. Subspecies and Distribution. R f. ferrumequinum Schreber, 1774 - S Europe from Iberian Peninsula and France E to Romania , Bulgaria , and Greece including SW Great Britain , S Germany , S Poland , and Crimea, also on most Mediterranean Is (Balearics, Corsica , Sardinia, Sicily, Malta , Crete, Cyprus , etc.), Turkey ( Anatolia ), NW Africa (N Morocco , N Algeria , N Tunisia , and NW Libya ), and Levant region. R f. proximus K. Andersen, 1905 - Transcaucasia, Mesopotamia , S Turkmenistan , N, W & S Iran , Afghanistan , Tajikistan , Kyrgyzstan , Pakistan , and N India (Kashmir). Descriptive notes . Head-body 54-71 mm , tail 31-44 mm , ear 19-25 mm , hindfoot 10- 14 mm , forearm 51-61 mm ; weight 13-44 g . The Greater Horseshoe Bat is the largest rhinolophid in Europe. Nominate ferrumequinum is generally larger than proximus. Dorsal pelage is generally grayish brown to pale brown (hairs are beige, with grayish brown or brown tips) and variably, usually lightly, tinged with red; venter is grayish white to yellowish white. Juveniles are ashy gray. Specimens of proximus are generally pale fawn. There is no orange morph. Males lack axillary tufts. Ears are medium to short ( c .41% of forearm length). Noseleafhas hastate or subtriangular lancet, becoming slightly concave near bluntly pointed tip; connecting process is rounded and much higher than tip of sella; sella is naked, relatively small, and curved forward, making front surface strongly concave, and sides are only slightly concave, and tip is pointed; and horseshoe is narrow at 6-5-9- 9 mm , does not cover muzzle, and has lateral leaflets (although sometimes inconspicuous) and deep median emargination. Lower lip has one or three grooves, and lateral grooves can be inconspicuous if present. Wings and uropatagium are pale brown or grayish brown. Baculum has dorso-ventrally flattened basal cone, with somewhat deep, ventral incision; rim of cone is thickened and forms strong protuberance on both sides ofventral incision ; shaft tapers and is almost cylindrical; and tip is a dorso-ventrally strongly flattened lancet Skull is robust, with sturdy zygomatic arches (zygomatic width is greater than mastoid width); nasal swellings are ofmedium low height; frontal depression is shallow; supraorbital crests are weak; and sagittal crest is well developed anteriorly but absent posteriorly. P2 is tiny and fully displaced labially or absent, allowing C1 and P4 to touch, and P3 is also tiny and fully displaced labially or absent, so that P2 and P are in full contact Dental formula is variable: 11/2, C 1/1, P 2/2, M 3/3 (x2) = 30; 11/2, C 1/1, P 1/2, M 3/3 (x2) = 28; 11/2, C 1/1, P 2/3, M 3/3 ( x 2) = 32; or 11/2, C 1/1’, P 1/3, M 3/3 (x2) = 30. Chromosomal complement has 2n = 58 and FNa = 60-62. Habitat Mainly forested habitats, especially in areas with abundant roosting sites, from sea level to elevations of c. 3500 m . Unlike the related Geoffroy’s Horseshoe Bat ( clivosus ), the Greater Horseshoe Bat is less associated with dry habitats, although it is found in some drier regions through West Asia and North Africa. In Europe, it is found in deciduous temperate woodlands throughout the northern part of its distribution and more Mediterranean and sub-Mediterranean woodlands and shrublands in southern Europe, North Africa, and the Levant Greater Horseshoe Bats are typically associated with forested habitats where they forage near vegetation, but they can be found in open areas, especially in West Asia. Throughout western and central Asia, they are largely associated with highland habitats and only found in temperate forested montane regions of Transcaucasia, Iran , and central Asia from Afghanistan to Kashmir. Food and Feeding. Greater Horseshoe Bats forage by slow hawking, fly-catching from perches, and ground-gleaning. When captured, insects are eaten on the wing or taken to a perch. They will sometimes land on the ground to capture dung beetles and other insects in dung piles. Most of their diet comes from species of Lepidoptera, Coleoptern , Hymenoptera , and Diptera (at least throughout Europe), but other arthropods also make up smaller proportions of their diets, including Neuroptera, Trichoptera, and Araneae . A large sample of 1580 feces found that the diet of the Greater Horseshoe Bat in the UK mostly contained Lepidoptera (moths) and changed throughout the year. In spring, they ate cockchafers {Melolontha melolonthd), beetles ( Geotrupe ), caddisflies (Trichoptera), tipulid flies, ichneumonid wasps, and moths. In autumn, dung beetles (Aphodius) and dung flies (various Diptera ) replaced various species of beetles in diets. Coleoptera, Lepidoptera , and Diptera dominated diets in Azerbaijan , and Coleoptera and Lepidoptera were most important in Turkey , Syria , and Jordan . Moths dominated three stomach samples from Iran , and some brachyceran Diptera, Trichoptera , and scarabaeid Coleoptera also were eaten. Greater Horseshoe Bats are versatile feeders and alter their diets based on prey availability throughout the year and among regions. Where they forage also can change; in the UK , they forage mainly in woodlands in spring and over pastures in late summer. Breeding. Greater Horseshoe Bats are seasonally monoestrous. Copulation occurs before hibernation; males mate with females in their harems. After mating, males secrete a plug into females’ vaginas to either prevent sperm competition or keep sperm stored in the female until the following spring. Sperm storage occurs in oviducts of females until ovulation and fertilization takes place in March—April. Gestation lasts 2—3 months depending on whether or not embryo development is lengthened due to torpor of the pregnant female; gestation is generally c .9 weeks with limited torpor. Young are bom in late spring or early summer depending on the region. In Algeria , young are bom inJune and volant by the end ofAugust. Litter size is one. Young hold onto their mothers until a little before weaning. Young will forage and hang separate from their mothers before weaning. Greater Horseshoe Bats mature very slowly, similar to many hibernating bats. Females reach sexual maturity at c .3 years of age, but they might not produce their first offspring until 3—5 years of age. Males generally mature at 2-4 years of age. Greater Horseshoe Bats are long-lived; maximum longevities recorded in the wild are 30 years and six months for a male and 27 years and eight months for a female. Activity patterns. Greater Horseshoe Bats are nocturnal and forage throughout the night They generally leave day roosts 15—30 minutes after sunset and begin to forage. They typically linger around day roosts for the early part of the night and fly further from the day roost as the night progresses. Most nightly activity occurs within 5—20 km ofroosts, and adults typically move further thanjuveniles. Greater Horseshoe Bats generally stay out all night or return after 2—3 hours and leave day roosts again for c .30 - 50 minutes before sunrise. They occasionally rest in night roosts. Activity decreases in cold temperatures, markedly below 10°C—the typical thermal threshold for insect activity. During the day, Greater Horseshoe Bats can enter a mild torpor when temperatures are below 22°C. They enter deep torpor (hibernation) from mid-autumn until spring (October-April in Europe), during which they find parts of caves that are warmer than 7 — 10 ° C. During this time, they continue to irregularly awake and leave roosts to forage when weather permits. They do not hibernate in southern parts of the distribution but have been found to hibernate in Iran . Day roosts of Greater Horseshoe Bats are generally in caves and other underground or rocky structures; they commonly inhabit abandoned or unused parts of buildings, particularly roofs. Greater Horseshoe Bats tend to favor houses in northern parts of their distribution but underground structures in the south. They also seem to favor caves as hibernacula in winter. Call shape is FM/ CF /FM, with terminal FM sweep usually having the greatest bandwidth. Aspects of calls vary among seasons and through the lifetime of an individual; juveniles emit lower frequencies than adults. Resting frequencies of mothers and young seem to be similar, indicating that the call is learned from the mother or inherited. F component is 77—83 kHz in southern Europe, 83—84 kHz in the UK , 81-7 kHz in Western Europe, and 84-6 kHz in Morocco . Mean call duration is 53-8 milliseconds in Greece , 21 milliseconds and 31-9 milliseconds in the UK , and 30-3 milliseconds in Morocco . Movements, Home range and Social organization. Greater Horseshoe Bats are highly gregarious, roosting singly, in small groups, or in very large colonies up to 1000 individuals, although they forage alone. Non-matemity colonies have been recorded with up to 500 individuals of both sexes; summer maternity colonies of up to 1000 females have been reported, although 100—300 individuals are more common. Maternity colonies are createdjust before females give birth. When first formed, maternity colonies can still include some persistent males and non-breeding individuals that usually leave after young are bom. Non-breeding individuals disperse like males or can remain in maternity colonies throughout the breeding season. Males form scattered small groups in separate day roosts. All male and all juvenile colonies have been recorded. Males and females start roosting together after young are weaned around late summer (at least in Europe), which leads to copulation in day roosts until the end of October before the beginning of hibernation (in northern populations). From late summer to about mid-autumn, males become territorial in roosting colonies, establishing small harems in the roost. Harems generally include the male and up to eight females, segregating into a single cluster in the roost Populations of Greater Horseshoe Bats are primarily sedentary and hibernate instead of migrating. Nevertheless , they often fly 20—35 km between summer roosts and hibernacula. They typically move to more secluded localities ( e .g. cool, deep parts of caves) where they can enter deep torpor. Individuals wrap themselves completely in their wing membranes during hibernation. Hibernacula generally include fewer individuals than summer roosts but can sometimes include clusters ofup to 100 individuals. Females seem to give birth in the same roosting area each year. Non-matemity summer roosts are often shared with other species of Rhinolophus , Miniopterus , Myotis , Asellia , and Plecotus . Maternity colonies are occasionally shared with other bat species. Status and Conservation. Classified as Least Concern on The IUCNed List. The Greater Horseshoe Bat has a wide distribution and is rather abundant throughout much of its distribution. Nevertheless, there are areas that have experienced well-documented declines, including Malta , Belgium , and the Netherlands where it is likely extinct Much of the documented decline has occurred throughout Europe, notably in north-western Europe. Populations in the UK have experienced massive declines, and the overall population is now stable at a low level of c .5000 individuals. Austrian populations have declined by 70% in the last ten years and are down to only c.30 breeding individuals. Population trends throughout the rest of Europe and North Africa are uncertain, but roosts seem to be disappearing throughout the Iberian Peninsula. Populations are considered stable in Croatia . There apparently has been a slow increase in populations in Romania since 1989 due to the reduced use of pesticides. The Greater Horseshoe Bat is rare in Switzerland but considered stable. It is considered relatively rare throughout much of its Asiatic distribution, although it is considerably common throughout Transcaucasia. Primary threats are fragmentation and habitat destruction from deforestation and agricultural expansion. Use of pesticides also negatively affects Greater Horseshoe Bats by targeting some important food sources such as melolonthid beetles, noctuid moth larvae, and crane flies. General roost disturbance is also an evident threat, especially for colonies in buildings where human intolerance can make it difficult for cohabitation. The Greater Horseshoe Bat is widely protected throughout Europe, where many underground roosts have been protected and building roosts have had management agreements to maintain the bat-human relationship. Legislation protects the Greater Horseshoe Bat in some but not all countries in its distribution. Bibliography. ACR (2018), Aldridge (1986), Arslan & Zima (2014), Aulagnier &Thévenot (1986), Bates & Harrison (1997), Benda & Gaisler (2015), Benda & Vallo (2012), Benda, Abi-Said et al. (2016), Benda, Andreas et al. (2006), Benda, Faizolâhi et al. (2012), Benda, Georgiakakis et al. (2008), Benda, Hanâk & Cervenÿ (2011), Benda, Ivanova et al. (2003), Benda, Lucan et al. (2010), Benda, Spitzenberger et al. (2014), Botnariuc &Tatole (2005), Csorba et al. (2003), Disca et al. (2014), landers, Jones et al. (2009), landers, Wei Li et al. (2011), Gaisler (2001, 2013b), Gunnell et al. (2011), Hanâk et al. (2001), Jones (1990), Jones & Ransome (1993), Jones & Rayner (1989), Jones & Siemers (2011), Koh Hung-Sun et al. (2014), Long & Schnitzler (1975), Ma Jie, Kobayasi et al. (2006), Park et al. (2000), Piraccini (2016a), Ransome & McOwat (1994), Rossiter , Benda et al. (2007), Rossiter, Jones et al. (2000a, 2000b, 2001), Schnitzler (1973), Schnitzler & Grinnell (1977), Stoffberg et al. (2010), Suga et al. (1976), Vogler & Neuweiler (1983), Walters et al. (2012).	Simmons, N.B. and A.L. Cirranello. 2022B. Bat Species of the World: A taxonomic and geographic database. Accessed on 10/11/2022.	Rhinolophidae	Rhinolophus ferrumequinum	Rhinolophus		ferrumequinum	Schreber	1774	1	Die S&auml;ugethiere	1: 174, pl. 62	Greater Horseshoe Bat	 colchicus Satunin, 1912; equinus M&uumlller, 1776; germanicus Koch, 1865; hippocrepis Schrank, 1798; homodorensis Daday, 1887; homorodalmasiensis Daday, 1885 [ nomen nudum ]; insulanus Barrett-Hamilton, 1910; italicus Koch, 1865; major E. Geoffroy, 1803 [not Kerr, 1792); major Kerr, 1792:99 [not Kerr, 1792:97]; martinoi Petrov, 1940; obscurus Cabrera, 1904; perspicillatus Blumenbach, 1779; solea Zimmermann, 1777 [unavailable; see Bull. Zool. Nomen. (1950) 4:547]; typicus K. Andersen, 1905; ungula Boddaert, 1785; unihastatus E. Geoffroy, 1803; <b>creticum</b> Iliopoulou-Georgudaki and Ondrias, 1985; <b>irani</b> Cheesman, 1921; rubiginosus Gubareff, 1941; <b>korai</b> Kuroda, 1938; pachyodontus Kishida, 1931 [ nomen nudum ]; quelpartis Mori, 1933; <b> nippon </b> Temminck, 1835; fudisanus Kishida, 1940; kosidianus Kishida, 1940; mikadoi Ognev, 1927; norikuranus Kishida, 1940; ogasimanus Kishida, 1940; <b> proximus </b> K. Andersen, 1905; <b>tragatus</b> Hodgson, 1835; brevitarsus Blyth, 1863 [ nomen nudum ]; regulus K. Andersen, 1905.	France.	Algeria, Morocco, and Tunisia; S Europe from Portugal to Greece and north to S England, the Netherlands, S Germany, Austria, Czech Republic, Slovakia, and Bulgaria; Turkey, Cyprus, Georgia, and Azerbaijan; Urkraine, Crimea, and Caucacus regions; the Mediterranean coast from Turkey to Israel and Jordan; NE Iraq, Iran, Turkmenistan, Uzbekistan, S Kazakhstan, Afghanistan, Pakistan and N India. Records at some localities in northern Europe (e.g., the Netherlands) apparently reflect temporary northern range extensions (Glas and Voûte, 1992a).	Not listed.	Least Concern	 ferrumequinum species group. Does not include nippon , tragatus or korai ; see Thomas (1997), Zhou et al. (2009), Benda and Vallo (2012), Burgin (2019), and Ikeda et al. (2020). Revised by Strelkov et al. (1978). Reviewed in part by Yoshiyuki (1989), Harrison and Bates (1991), Paz (1995), Kock (1996),Bates and Harrison (1997), Sinha (1999), Zagorodnyuk (1999), Horácek et al. (2000) and Gaisler (2001a). Subspecies limits are somewhat unclear and there may be more than one species present in this complex; see discussion in Csorba et al. (2003). For a review of the species in Iran see Shahabi et al. (2019).	Mammal Diversity Database. (2023). Mammal Diversity Database (Version 1.11) [Data set]. Zenodo. https://doi.org/10.5281/zenodo.7830771 released 15 April 2023	Rhinolophus ferrumequinum	23	Greater Horseshoe Bat	Larger Horseshoe Bat	Theria	Placentalia	Boreoeutheria	Laurasiatheria	CHIROPTERA	PTEROPODIFORMES	NA	NA	RHINOLOPHOIDEA	RHINOLOPHIDAE	NA	NA	Rhinolophus	NA	ferrumequinum	von Schreber	1774	1						France.			ferrumequinum (von Schreber, 1774)|equinus (P. L. S. Müller, 1776)|solea (E. A. W. Zimmermann, 1777) [unavailable name]|perspicillatus (Blumenbach, 1779)|ungula (Boddaert, 1785)|major (Kerr, 1792) [Kerr, 1792:99, not Kerr, 1792:97]|hippocrepis (Schrank, 1798)|major É. Geoffroy Saint-Hilaire, 1803 [preoccupied]|unihastatus É. Geoffroy Saint-Hilaire, 1803|germanicus L. Koch, 1865|italicus L. Koch, 1865|homorodalmasiensis Daday, 1885 [nomen nudum]|homodorensis Daday, 1887|obscurus Cabrera, 1904|proximus K. Andersen, 1905|typicus K. Andersen, 1905|insulanus Barrett-Hamilton, 1910|colchicus Satunin, 1912|irani Cheesman, 1921|martinoi Petrov, 1940|rubiginosus Gubareff, 1941|creticum Iliopoulou-Georgudaki & Ondrias, 1985	previously included R. nippon	Ohdachi, S. D. I, Ishibashi, Y., Iwasa, M. A., & Saitoh, Takashi (2009). The Wild Mammals of Japan, Shoukadoh, Kyoto.|Koh, H. S., Jo, J. E., Oh, J. G., Kweon, G. H., Ahn, N. H., Sin, W. H., & Sin, D. S. (2014). Little genetic divergence of the greater horseshoe bat Rhinolophus ferrumequinum from far-eastern Asia, with a preliminary report on genetic differentiation of R. ferrumequinum from Eurasia and northern Africa examined from cytochrome b sequences. Russian Journal of Theriology, 13, 97-103.|Ikeda, Y., Jiang, T., Oh, H., Csorba, G., & Motokawa, M. (2020). Geographic variations of skull morphology in the Rhinolophus ferrumequinum species complex (Mammalia: Chiroptera). Zoologischer Anzeiger, 288, 125-138.	Morocco|Algeria|Tunisia|Libya|United Kingdom|Portugal|Spain|France|Luxembourg|Germany|Switzerland|Liechtenstein|Italy|Malta?|Austria|Czech Republic|Poland|Slovakia|Hungary|Slovenia|Croatia|Bosnia & Herzegovina|Serbia|Kosovo|Montenegro|Albania|North Macedonia|Greece|Bulgaria|Romania|Ukraine|Russia|Greece|Armenia|Azerbaijan|Turkey|Cyprus|Syria|Lebanon|Israel|Palestine|Jordan|Iraq|Iran|Turkmenistan|Tajikistan|Kyrgyzstan|Pakistan|India	Africa|Asia|Europe	Palearctic	LC	0	0	0	Rhinolophus_ferrumequinum	0	sciname match	Rhinolophus_ferrumequinum	0	IUCN. 2022. The IUCN Red List of Threatened Species. Version 2022-1. https://www.iucnredlist.org. Accessed on [28 September, 2022].	19517	Rhinolophus ferrumequinum	ANIMALIA	CHORDATA	MAMMALIA	CHIROPTERA	RHINOLOPHIDAE	Rhinolophus	ferrumequinum	(Schreber, 1774)	Although Thomas (1997, unpublished thesis) found very high divergence in mitochondrial DNA sequences, Csorba et al. ;(2003) refrained from splitting Japanese Greater Horseshoe Bats (Rhinolophus nippon ) from R. ferrumequinum . Several subspecies are identified over the range, of which two occur in the western Palaearctic: R. f. creticus (Crete) and R. f. ferrumequinum (rest of western Palaearctic range).	20000000	Rhinolophus ferrumequinum	Least Concern		2016	2016-04-25 00:00:00 UTC	3.1	English	This species has a large range. Although there have been marked and well-documented declines in some areas, the species remains widespread, abundant, and apparently stable in other areas. Assessed as Least Concern.	It forages in pastures, deciduous temperate woodland, Mediterranean and sub-mediterranean shrubland and woodland. Important foraging habitat and landscape features includes woodland, particularly early in the year, and permanent summer-grazed pasture, particularly late in the summer. It feeds on beetles, moths and other insects at low level in pastures and in trees up to 2 to 3 km from the roost each night (by aerial hawking or perch feeding). Summer roosts are located in warm natural and artificial underground sites, and attics in the northern part of the range. The species will use caves all year, but particularly in northern Europe it uses buildings for summer maternity colonies. Where the species occupies buildings, it requires particular features of the building itself, as well as proximity to good foraging areas and underground sites for torpor at various times of year and for winter hibernation (Hutson et al. 2001). In winter it hibernates in cold underground sites (usually large caves). The requirements for hibernation appear to be within limited confines of temperature and humidity, but vary with age, sex and condition. The mean temperatures in spring, at the end of hibernation, may influence the time of birth at the summer maternity colonies; late parturition in turn increases mortality rates of juveniles (Hutson et al. 2001). In southern parts of the range, they are active all year-round. A sedentary species, distances of 20-30 km between winter and summer roosts are typical (longest distance recorded 180 km: de Paz et al. 1986). In South Asia, this species is found in montane forests among the mountains and valleys of the Himalaya. It is gregarious and roosts in caves, old temples, old and ruined buildings in tight clusters. It has a slow and fluttering flight and feeds on small insects, lacewings, small moths, spiders and grasshoppers. A single young is born after a gestation period of 72 days (Bates and Harrison 1997).	The main threats are fragmentation and isolation of habitats, change of management regime of deciduous forests and agricultural areas, loss of insects due to pesticide use, and disturbance and loss of underground habitats and attics. In northwest Europe, habitat change is likely to have been amongst the major causes of declines, the conversion of woodland and small-field landscape to large-scale agricultural land being particularly damaging. While declines elsewhere, particularly in eastern Europe, may currently not be so marked, the loss of cultural landscapes in those countries as they move towards western-style economies may have significant effects in the near future. The use of pesticides has been a recognized threat to the insect food, particularly where these have been directed against the larvae of favoured food items, such as melolonthid beetles, larvae of noctuid moths or crane-flies. Favoured prey may be affected secondarily by pesticide use, such as the loss of dung fauna from the use of persistent anti-parasitic drugs (avermectins) on cattle. Populations in caves and other underground habitats have suffered from increased disturbance (for example by tourist visits to caves). In buildings, colonies may be affected by human intolerance, renovation work or the application of pesticides, such as some of those used for the remedial treatment of timbers (Hutson et al. 2001). In South Asia, this species is threatened by deforestation, generally resulting from logging operations and the conversion of land for agricultural and other uses. Disturbance to roosting sites is likely to be a potential threat to the populations of this species (Molur et al. 2002).	It is an infrequent species in most parts of its range, although in at least parts of south-west Asia and the Caucasus it is abundant and widespread (it is the most frequently reported species in Turkey: A. Karatash pers. comm. 2005), with populations in Iran and Turkey considered to be stable (A. Karatash, M. Sharifi and K. Tsytsulina pers. comm. 2005), although it may be decreasing in Russian parts of the Caucasus (S. Kruskop pers. comm. 2005). Summer colonies of c.30-200 individuals (up to 400 animals), and winter clusters of up to 500 animals are typical.  In Europe, the two most widespread Rhinolophus species, R. ferrumequinum and R. hipposideros , are of particular conservation concern and are the subject of considerable research and monitoring. R. ferrumequinum has shown marked declines in range in northwest Europe within the last 100 years (e.g. United Kingdom, Germany, Austria), and has become extinct in some countries (e.g. Belgium, Netherlands). However, there are signs of stabilization and/or recovery in some northwest European countries (Hutson et al 2001). For example, in the UK the species declined massively in the past but it is now stable at a low population level (around 5,000 individuals) (Ransome and Hutson 2000). However, in Austria declines continue, with population reductions of 70% in the last 10 years (from 100 to 30 breeding individuals: Spitzenberger 2002, F. Spitzenberger pers. comm. 2006). In other parts of Europe, trends vary and are generally less well known: in Malta the species has become extinct, in Portugal and Spain the trend is not known (although some colonies have disappeared in Spain) (Palomo and Gisbert 2002, Cabral et al. 2005) ;, while it is currently considered extinct in Gibraltar ;(S. Finlayson pers. comm. 2015); ;in Croatia the population is thought to be stable (N. Tvrtkovic pers. comm.), and in Romania it has been slowly increasing since 1989 due to reduced use of pesticides and a return to traditional agriculture with colonies of up to 800 individuals. In Switzerland the species is very rare (3 maternity roosts with some 200 individuals), but the population trend appears stable (H. Kraettli pers. comm. 2006). In its north African and south Asian range the population size and trends are unknown.	Decreasing	This species has a wide range in the Palaearctic, occurring from North Africa and southern Europe through south-west Asia, the Caucasus, Iran, Afghanistan, Pakistan and the Himalayas to south-eastern China, Korea, and Japan (Csorba et al. 2003, Abe et al. 2005). It usually occurs below 800 m asl, but can be found up to 3,000 m asl in the Caucasus depending on roost availability and humidity (K. Tsytsulina pers. comm. 2005).		Terrestrial	The greater horseshoe bat has been the subject of widespread conservation activity, especially in Europe. Until recently, this has concentrated on roosts in buildings and caves. Many buildings used as roosts have management agreements and many underground sites have been protected. Nevertheless, sites continue to be lost or damaged. More recently, attention has turned to identifying more precisely the food and foraging requirements. A European meeting (Germany, May 1995) discussed the status and conservation needs for the species on a pan-European scale. The Bern Convention has commissioned a Europe-wide Species Action Plan under the Pan-European Biological and Landscape Diversity Strategy (Ransome and Hutson 2000).  It is protected by national legislation in some range states. There are international legal obligations for its protection through the Bonn Convention (Eurobats) and Bern Convention in parts of its range where these apply. It is included in Annex II (and IV) of the European Union Habitats Directive, and hence requires special measures for conservation including designation of Special Areas for Conservation. There is some habitat protection through Natura 2000 (some roosts are already protected by national legislation). There are no specific conservation measures in place for the species in North Africa or South Asia. Populations should ;be monitored to record changes in abundance and distribution (Molur et al. 2002).	Palearctic		FALSE	FALSE	Global	Simmons, N. B., & Cirranello, A. L. (2023). Batnames.org Species List Version 1.4 (1.4). Zenodo. https://doi.org/10.5281/zenodo.8136157 	Rhinolophidae	Rhinolophus		ferrumequinum	Schreber	1774	1	Die S&auml;ugethiere	1: 174, pl. 62	Greater Horseshoe Bat	 colchicus Satunin, 1912; equinus M&uumlller, 1776; germanicus Koch, 1865; hippocrepis Schrank, 1798; homodorensis Daday, 1887; homorodalmasiensis Daday, 1885 [ nomen nudum ]; insulanus Barrett-Hamilton, 1910; italicus Koch, 1865; major E. Geoffroy, 1803 [not Kerr, 1792); major Kerr, 1792:99 [not Kerr, 1792:97]; martinoi Petrov, 1940; obscurus Cabrera, 1904; perspicillatus Blumenbach, 1779; solea Zimmermann, 1777 [unavailable; see Bull. Zool. Nomen. (1950) 4:547]; typicus K. Andersen, 1905; ungula Boddaert, 1785; unihastatus E. Geoffroy, 1803; <b>creticum</b> Iliopoulou-Georgudaki and Ondrias, 1985; <b>irani</b> Cheesman, 1921; rubiginosus Gubareff, 1941; <b>korai</b> Kuroda, 1938; pachyodontus Kishida, 1931 [ nomen nudum ]; quelpartis Mori, 1933; <b> nippon </b> Temminck, 1835; fudisanus Kishida, 1940; kosidianus Kishida, 1940; mikadoi Ognev, 1927; norikuranus Kishida, 1940; ogasimanus Kishida, 1940; <b> proximus </b> K. Andersen, 1905; <b>tragatus</b> Hodgson, 1835; brevitarsus Blyth, 1863 [ nomen nudum ]; regulus K. Andersen, 1905.	France.	Algeria, Morocco, and Tunisia; S Europe from Portugal to Greece and north to S England, the Netherlands, S Germany, Austria, Czech Republic, Slovakia, and Bulgaria; Turkey, Cyprus, Georgia, and Azerbaijan; Urkraine, Crimea, and Caucacus regions; the Mediterranean coast from Turkey to Israel and Jordan; NE Iraq, Iran, Turkmenistan, Uzbekistan, S Kazakhstan, Afghanistan, Pakistan and N India. Records at some localities in northern Europe (e.g., the Netherlands) apparently reflect temporary northern range extensions (Glas and Voûte, 1992a).	Not listed.	Least Concern	 ferrumequinum species group. Does not include nippon , tragatus or korai ; see Thomas (1997), Zhou et al. (2009), Benda and Vallo (2012), Burgin (2019), and Ikeda et al. (2020). Revised by Strelkov et al. (1978). Reviewed in part by Yoshiyuki (1989), Harrison and Bates (1991), Paz (1995), Kock (1996),Bates and Harrison (1997), Sinha (1999), Zagorodnyuk (1999), Horácek et al. (2000) and Gaisler (2001a). Subspecies limits are somewhat unclear and there may be more than one species present in this complex; see discussion in Csorba et al. (2003). For a review of the species in Iran see Shahabi et al. (2019).	Rhinolophus ferrumequinum	1004684	23	Greater Horseshoe Bat	Larger Horseshoe Bat	Theria	Placentalia	Boreoeutheria	Laurasiatheria	CHIROPTERA	PTEROPODIFORMES	NA	NA	RHINOLOPHOIDEA	Rhinolophidae	NA	NA	Rhinolophus	NA	ferrumequinum	von Schreber	1774	1						France.			ferrumequinum (von Schreber, 1774)|equinus (P. L. S. Müller, 1776)|solea (E. A. W. Zimmermann, 1777) [unavailable name]|perspicillatus (Blumenbach, 1779)|ungula (Boddaert, 1785)|major (Kerr, 1792) [Kerr, 1792:99, not Kerr, 1792:97]|hippocrepis (Schrank, 1798)|major É. Geoffroy Saint-Hilaire, 1803 [preoccupied]|unihastatus É. Geoffroy Saint-Hilaire, 1803|germanicus L. Koch, 1865|italicus L. Koch, 1865|homorodalmasiensis Daday, 1885 [nomen nudum]|homodorensis Daday, 1887|obscurus Cabrera, 1904|proximus K. Andersen, 1905|typicus K. Andersen, 1905|insulanus Barrett-Hamilton, 1910|colchicus Satunin, 1912|irani Cheesman, 1921|martinoi Petrov, 1940|rubiginosus Gubareff, 1941|creticum Iliopoulou-Georgudaki & Ondrias, 1985	previously included R. nippon	Ohdachi, S. D. I, Ishibashi, Y., Iwasa, M. A., & Saitoh, Takashi (2009). The Wild Mammals of Japan, Shoukadoh, Kyoto.|Koh, H. S., Jo, J. E., Oh, J. G., Kweon, G. H., Ahn, N. H., Sin, W. H., & Sin, D. S. (2014). Little genetic divergence of the greater horseshoe bat Rhinolophus ferrumequinum from far-eastern Asia, with a preliminary report on genetic differentiation of R. ferrumequinum from Eurasia and northern Africa examined from cytochrome b sequences. Russian Journal of Theriology, 13, 97-103.|Ikeda, Y., Jiang, T., Oh, H., Csorba, G., & Motokawa, M. (2020). Geographic variations of skull morphology in the Rhinolophus ferrumequinum species complex (Mammalia: Chiroptera). Zoologischer Anzeiger, 288, 125-138.				Morocco|Algeria|Tunisia|Libya|United Kingdom|Portugal|Spain|France|Luxembourg|Germany|Switzerland|Liechtenstein|Italy|Malta?|Austria|Czech Republic|Poland|Slovakia|Hungary|Slovenia|Croatia|Bosnia & Herzegovina|Serbia|Kosovo|Montenegro|Albania|North Macedonia|Greece|Bulgaria|Romania|Ukraine|Russia|Greece|Armenia|Azerbaijan|Turkey|Cyprus|Syria|Lebanon|Israel|Palestine|Jordan|Iraq|Iran|Turkmenistan|Tajikistan|Kyrgyzstan|Pakistan|India	Africa|Asia|Europe	Palearctic	LC	0	0	0	Rhinolophus_ferrumequinum	0	sciname match	Rhinolophus_ferrumequinum	0	Burgin, C. J., Zijlstra, J. S., Becker, M. A., Handika, H., Alston, J. M., Widness, J., Liphardt, S., Huckaby, D. G., and Upham, N. S. (2025). How many mammal species are there now? Updates and trends in taxonomic, nomenclatural, and geographic knowledge. Journal of Mammalogy in revision: TBD. https://doi.org/10.1101/2025.02.27.640393	Rhinolophus_ferrumequinum	1004684	23	Greater Horseshoe Bat	Larger Horseshoe Bat	Theria	Placentalia	Boreoeutheria	Laurasiatheria	Chiroptera	Yinpterochiroptera	NA	NA	Rhinolophoidea	Rhinolophidae	NA	NA	Rhinolophus	NA	ferrumequinum	von Schreber	1	Vespertilio ferrum-equinum	Schreber, J.C.D. von. 1774. pl. 62. P. pl. 62 in Schreber, J.C.D. von. 1774-1855. Die Säugthiere in Abbildungen nach der Natur, mit Beschreibungen. Walther, Erlangen.	https://www.biodiversitylibrary.org/page/31060128				France.			previously included R. nippon	Ohdachi, S. D. I, Ishibashi, Y., Iwasa, M. A., & Saitoh, Takashi (2009). The Wild Mammals of Japan, Shoukadoh, Kyoto.|Koh, H. S., Jo, J. E., Oh, J. G., Kweon, G. H., Ahn, N. H., Sin, W. H., & Sin, D. S. (2014). Little genetic divergence of the greater horseshoe bat Rhinolophus ferrumequinum from far-eastern Asia, with a preliminary report on genetic differentiation of R. ferrumequinum from Eurasia and northern Africa examined from cytochrome b sequences. Russian Journal of Theriology, 13, 97-103.|Ikeda, Y., Jiang, T., Oh, H., Csorba, G., & Motokawa, M. (2020). Geographic variations of skull morphology in the Rhinolophus ferrumequinum species complex (Mammalia: Chiroptera). Zoologischer Anzeiger, 288, 125-138.				Morocco|Algeria|Tunisia|Libya|United Kingdom|Portugal|Spain|France|Luxembourg|Germany|Switzerland|Liechtenstein|Italy|Malta?|Austria|Czech Republic|Poland|Slovakia|Hungary|Slovenia|Croatia|Bosnia and Herzegovina|Serbia|Kosovo|Montenegro|Albania|North Macedonia|Greece|Bulgaria|Romania|Ukraine|Russia|Armenia|Azerbaijan|Turkey|Cyprus|Syria|Lebanon|Israel|Palestine|Jordan|Iraq|Iran|Turkmenistan|Tajikistan|Kyrgyzstan|Pakistan|India	Africa|Asia|Europe	Palearctic	LC	0	0	0	Rhinolophus_ferrumequinum	0	sciname match	Rhinolophus_ferrumequinum	0	Simmons, N. B., & Cirranello, A. L. (2025). Batnames.org Species List Version 1.7 (1.7). Zenodo. https://doi.org/10.5281/zenodo.14796586	Rhinolophidae	Rhinolophus		ferrumequinum	Schreber	1774	1	Die S&auml;ugethiere	1: 174, pl. 62	Greater Horseshoe Bat	colchicus Satunin, 1912; equinus M&uumlller, 1776; germanicus Koch, 1865; hippocrepis Schrank, 1798; homodorensis Daday, 1887; homorodalmasiensis Daday, 1885 [nomen nudum]; insulanus Barrett-Hamilton, 1910; italicus Koch, 1865; major E. Geoffroy, 1803 [not Kerr, 1792); major Kerr, 1792:99 [not Kerr, 1792:97]; martinoi Petrov, 1940; obscurus Cabrera, 1904; perspicillatus Blumenbach, 1779; solea Zimmermann, 1777 [unavailable; see Bull. Zool. Nomen. (1950) 4:547]; typicus K. Andersen, 1905; ungula Boddaert, 1785; unihastatus E. Geoffroy, 1803; creticum Iliopoulou-Georgudaki and Ondrias, 1985; irani Cheesman, 1921; rubiginosus Gubareff, 1941; korai Kuroda, 1938; pachyodontus Kishida, 1931 [nomen nudum]; quelpartis Mori, 1933; nippon Temminck, 1835; fudisanus Kishida, 1940; kosidianus Kishida, 1940; mikadoi Ognev, 1927; norikuranus Kishida, 1940; ogasimanus Kishida, 1940; proximus K. Andersen, 1905; tragatus Hodgson, 1835; brevitarsus Blyth, 1863 [nomen nudum]; regulus K. Andersen, 1905.	France.	Algeria, Morocco, and Tunisia; S Europe from Portugal to Greece and north to S England, the Netherlands, S Germany, Austria, Czech Republic, Slovakia, and Bulgaria; Turkey, Cyprus, Georgia, and Azerbaijan; Urkraine, Crimea, and Caucacus regions; the Mediterranean coast from Turkey to Israel and Jordan; NE Iraq, Iran, Turkmenistan, Uzbekistan, S Kazakhstan, Afghanistan, Pakistan and N India. Records at some localities in northern Europe (e.g., the Netherlands) apparently reflect temporary northern range extensions (Glas and Voûte, 1992a).	<a href='https://cites.org/eng/app/appendices.php' target='_blank'>Not Listed</a>	<a href='https://www.iucnredlist.org/species/19517/21973253/' target='_blank'>Least Concern</a>	ferrumequinum species group. Does not include nippon, tragatus or korai; see Thomas (1997), Zhou et al. (2009), Benda and Vallo (2012), Burgin (2019), and Ikeda et al. (2020). Revised by Strelkov et al. (1978). Reviewed in part by Yoshiyuki (1989), Harrison and Bates (1991), Paz (1995), Kock (1996),Bates and Harrison (1997), Sinha (1999), Zagorodnyuk (1999), Horácek et al. (2000) and Gaisler (2001a). Subspecies limits are somewhat unclear and there may be more than one species present in this complex; see discussion in Csorba et al. (2003). For a review of the species in Iran see Shahabi et al. (2019).		Mammal Diversity Database. (2025). Mammal Diversity Database (Version 2.2) [Data set]. Zenodo. https://doi.org/10.5281/zenodo.15007505	NA	Rhinolophus ferrumequinum; Rhinolophus ferrumequinum; Rhinolophus ferrumequinum; Rhinolophus ferrumequinum; Rhinolophus ferrumequinum; Rhinolophus ferrumequinum; ferrumequinum; creticum; irani; korai; nippon; proximus; tragatus; colchicus; equinus; germanicus; hippocrepis; homodorensis; homorodalmasiensis; insulanus; italicus; major; martinoi; obscurus; perspicillatus; solea; typicus; ungula; unihastatus; irani - rubiginosus; korai - pachyodontus; quelpartis; nippon - fudisanus; kosidianus; mikadoi; norikuranus; ogasimanus; tragatus - brevitarsus; regulus; ferrumequinum; proximus; creticumis; proximus - irani; creticum; irani; korai; nippon; proximus; tragatus; colchicus; equinus; germanicus; hippocrepis; homodorensis; insulanus; italicus; major; martinoi; obscurus; perspicillatus; solea; typicus; ungula; unihastatus; irani - rubiginosus; korai - pachyodontus; quelpartis; nippon - fudisanus; kosidianus; mikadoi; norikuranus; ogasimanus; tragatus - brevitarsus; regulus; ferrumequinum; equinus; solea; perspicillatus; ungula; major; hippocrepis; major; unihastatus; germanicus; italicus; homorodalmasiensis; homodorensis; obscurus; proximus; typicus; insulanus; colchicus; irani; martinoi; rubiginosus; creticum; Greater Horseshoe Bat; Grand Rhinolophe; Grosse Hufeisennase; Herradura grande; German; Larger Horseshoe Bat; Other commonnames; Greater Horseshoe Bat; Larger Horseshoe Bat; Greater Horseshoe Bat; Greater Horseshoe Bat; R. ferrumequinum