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Lifespan, longevity, and ageing

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Maximum longevity: 30.5 years (wild)
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Morphology

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Rhinolophus ferrumequinum is the largest horseshoe bat in Europe (Schober and Grimmberger, 1997). Its most distinctive feature is the upper saddle process or noseleaf, the upper part of which is pointed while the lower part is horseshoe shaped (Nowak, 1994). Tooth and bone structures distinguish R. ferrumequinum from other rhinolophids. The first premolar on the upper jaw protrudes from the row of teeth. Often this premolar is very small or non-existent. The third and fourth metacarpal bones in the wings are shorter than those of its relatives (Koopman, 1994). The tragus is absent (Simmons and Conway, 1997). The length of the head and body ranges from 57 to 71mm, the tail length ranges from 35 to 43 mm and the forearm from 54 to 61 mm. The wing span ranges from 350 to 400 mm, and the weight from 17 to 34 grams. The Greater horseshoe bat can also be identified by its color. The back is brownish gray with a slight tint of red, while the underside is a lighter gray color. The membrane that connects the forearm and tail is brownish gray. Young R. ferrumequinum are uniformly gray. (Schober and Grimmberger, 1997)

Range mass: 17 to 34 g.

Range length: 57 to 71 mm.

Range wingspan: 350 to 400 mm.

Other Physical Features: endothermic ; bilateral symmetry

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Price, R. 2002. "Rhinolophus ferrumequinum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Rhinolophus_ferrumequinum.html
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Life Expectancy

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Lifespan/ Longevity: The oldest recorded age of an R. ferrumequinum is 30 years (Schober and Grimmberger, 1997)

Range lifespan
Status: wild:
30 (high) years.

Average lifespan
Status: wild:
30.0 years.

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Price, R. 2002. "Rhinolophus ferrumequinum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Rhinolophus_ferrumequinum.html
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Rebecca Price, University of Michigan-Ann Arbor
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Habitat

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Rhinolophus ferrumequinum prefer to dwell in karst caves and cave-like spaces under boulders year-round (DeBlase, 1980). In the winter, R. ferrumequinum roost in caves warmer then 7-10 degrees Celsius. This species prefers warmer regions, which have water, shrubs and trees (Schober and Grimmberger, 1997).

Habitat Regions: temperate ; terrestrial

Terrestrial Biomes: forest

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Price, R. 2002. "Rhinolophus ferrumequinum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Rhinolophus_ferrumequinum.html
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Distribution

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Rhinolophus ferrumequinum are found in northern India, south to northwestern Africa and throughout temperate Eurasia including Great Britain and Japan

(Koopman, 1994). They are also found throughout Turkey, Israel and Jordan (DeBlase, 1980)

Biogeographic Regions: palearctic (Native )

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Trophic Strategy

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Rhinolophus ferrumequinum hunt at nightfall. This species is an insectivore and preys primarily on larger insects such as Lepidoptera (butterflies and moths) and Coleoptera (beetles) (Jones, 1990). In order to obtain their prey, hunting bats fly close to the ground surface (Schober and Grimmberger, 1997). Rhinolophus ferrumequinum does not hunt in the winter unless the air temperature is warm enough for insect flight, and as a result they hunt less during colder and inclement weather (Racey, 1982). Like other microchiropteran bat species, R. ferrumequinum use echolocation to locate their prey. This species emits a call consisting of short blasts at a low frequency (77-81 kHz) (Schober and Grimmberger, 1997). The call is emitted from the nose, not the mouth (Nowak, 1994).

Animal Foods: insects

Primary Diet: carnivore (Insectivore )

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Price, R. 2002. "Rhinolophus ferrumequinum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Rhinolophus_ferrumequinum.html
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Untitled

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There are six subspecies of R. ferrumequinum: R.f. ferrumequinum, R.f. creticus, R.f. proximus, R.f. tragatus, and R.f. nippon. They are found in Europe and northwestern Africa, Crete, southwestern Asia to Kashmir, northern India and southwestern China, and northern and central China, Korea and Japan, respectively (Koopman, 1994).

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Price, R. 2002. "Rhinolophus ferrumequinum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Rhinolophus_ferrumequinum.html
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Behavior

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Perception Channels: tactile ; chemical

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Price, R. 2002. "Rhinolophus ferrumequinum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Rhinolophus_ferrumequinum.html
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Conservation Status

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Rhinolophus ferrumequinum are threatened in western Germany and Austria. Because R. ferrumequinum are sensitive to human disturbances, a special effort has to be made to protect the roosts and to lower the amount of insecticides used on crops in western Germany and Austria. The insecticides kill R. ferrumequinum prey, thus destroying their only food source (Nowak, 1994), as well as harming the bats themselves

US Migratory Bird Act: no special status

US Federal List: no special status

CITES: appendix iii

IUCN Red List of Threatened Species: least concern

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Price, R. 2002. "Rhinolophus ferrumequinum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Rhinolophus_ferrumequinum.html
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Benefits

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Positive Impacts: controls pest population

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Price, R. 2002. "Rhinolophus ferrumequinum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Rhinolophus_ferrumequinum.html
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Reproduction

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Females are not sexually mature until their third year, while males are sexually reproductive at age two. Females often do not have young until their fifth year (Racey,1982). Mating usually occurs in the fall but can also occur in the spring (Rossiter et al. 2000). Fertilization does not take place until later and is controlled by the females (Schober and Grimmberger, 1997). After mating, the vagina of Rhinolophus ferrumequinum females is closed by a vaginal plug secreted by the male. It is speculated that the plug is used to keep unwanted sperm out, as a method of mate selection (Fenton, 1984) or to hold the sperm until fertilization in the spring (Rossiter et al., 2000). Female R. ferrumequinum return to the same roosting area year after year to give birth and care for their young.

Average number of offspring: 1.

Average weaning age: 8 weeks.

Range age at sexual or reproductive maturity (female): 2 to 5 years.

Average age at sexual or reproductive maturity (female): 3 years.

Range age at sexual or reproductive maturity (male): 2 to 5 years.

Average age at sexual or reproductive maturity (male): 3 years.

Key Reproductive Features: seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; fertilization (Internal ); sperm-storing ; delayed fertilization

Average birth mass: 5.8 g.

Average gestation period: 80 days.

Average number of offspring: 1.

A female produces one baby which is typically born during June or July (Schober and Grimmberger, 1997). Young open their eyes at 7 day and can fly during the third to fourth week. After seven to eight weeks, young are ready to leave the roost. Females form maternity roosts in warmer places such as attics to care for the young (Schober and Grimmberger, 1997). Greater horseshoe bats lose their milk teeth before birth (Nowak, 1994).

Parental Investment: altricial ; female parental care

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Price, R. 2002. "Rhinolophus ferrumequinum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Rhinolophus_ferrumequinum.html
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Biology

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Greater horseshoe bats often roost in buildings during the summer, and their presence can be detected by piles of excrement on the ground. They leave their roosts just after sunset, and during the summer, spend about an hour feeding before returning. They often feed again just before dawn. At the end of August they stay on the wing all night.Greater horseshoe bats mate in autumn, sometimes in late winter or early spring. They form maternity roosts in May and the young are born in mid-July. This species reaches maturity at around three years old and they may live for 30 years. Their preferred food is large beetles, such as cockchafers and dung beetles, large moths and caddis flies. They have been observed watching from a regular roost and then flying out to take passing insects. Greater horseshoe bats hibernate in caves, cellars or disused mines, from late September to mid-May. They may emerge to feed during mild spells.
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Conservation

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This species is listed in the UK Biodiversity Action Plans (UKBAPs), and has been included in English Nature's Species Recovery Programme. It belongs to a family of animals that have been much maligned over the centuries. However, more people are realising just how fascinating bats are, and they receive a high level of legal protection. The main effort in their conservation is to encourage landowners and farmers to manage their land in ways that benefit the bats. They are also being asked to limit the use of ivermectin insecticides, commonly used for treating cattle. The chemical in the insecticide also poisons the cattle's dung, and kills the larvae of dung beetles, one of the greater horseshoe bat's principal foods. As more people learn about bats, it is hoped that the efforts to conserve them as a breeding species will gain more support. They are an intriguing group of mammals, and undeserving of their sinister reputation. The Bat Conservation Trust carries out work on surveys and monitoring, and employs many volunteers.
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Description

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The greater horseshoe bat is the larger of the two horseshoe bats found in Britain. They are so-named from the horseshoe shaped nose 'leaf', used as part of the bat's echolocation system. The ears are leaf-shaped and have a sharply pointed tip. The fur is thick, and coloured ash-grey above, and buff underneath. Bats are not blind as was once popularly thought. They have good eyesight but rely on their echolocation to navigate and to detect their insect prey. They emit a succession of high-pitched squeaks and judge their position and the location of their prey from the reflected echoes.
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Habitat

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The greater horseshoe bat prefers traditionally managed farmland, with grazing pasture and broad-leaved woodland.
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Range

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This bat is found in central and southern Europe but has declined significantly in northern Europe. In the UK it is restricted to southern England and South Wales.
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Status

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Classified as Endangered in the UK, listed under Appendix II of the Bonn Convention, Appendix II of the Berne Convention, Annexes II and IV of the EC Habitats Directive, Schedule 2 of the Conservation Regulations 1994 and protected in the UK under Schedule 5 of the Wildlife and Countryside Act (as amended).
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Threats

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The greater horseshoe bat has declined by over 90% in numbers during the last 100 years. This is due largely to habitat loss, caused by modern intensive farming methods. The destruction of woods, roosting sites, old pastureland, and the use of chemical insecticides, which have seriously reduced the abundance of their insect prey, have all contributed to this decline.
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Associations

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Animal / parasite / ectoparasite
Basilia nana ectoparasitises Rhinolophus ferrumequinum
Other: minor host/prey

In Great Britain and/or Ireland:
Animal / parasite / ectoparasite
Phthiridium biarticulata ectoparasitises Rhinolophus ferrumequinum
Other: major host/prey

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Greater horseshoe bat

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The greater horseshoe bat (Rhinolophus ferrumequinum) is an insectivorous bat of the genus Rhinolophus. Its distribution covers Europe, Northern Africa, Central Asia and Eastern Asia.[1] It is the largest of the horseshoe bats in Europe and is thus easily distinguished from other species. The species is sedentary, typically travelling up to 30 kilometres (19 mi) between the winter and summer roosts, with the longest recorded movement being 180 km (110 mi). The frequencies used by this bat species for echolocation lie between 69–83 kHz, have most energy at 81 kHz and have an average duration of 37.4 ms.[2][3]

Description

The greater horseshoe bat is the largest horseshoe bat in Europe.[4] It has a distinctive noseleaf, which has a pointed upper part and a horseshoe-shaped lower part.[5] Its horseshoe noseleaf helps to focus the ultrasound it uses to 'see'. The greater horseshoe bat also has tooth and bone structures that are distinct from that of other rhinolophids. Its first premolar on the upper jaw protrudes from the row of teeth.[6] For other horseshoe bats, this premolar is very small or non-existent.[6] Also in comparison to its relatives, the greater horseshoe bat has relatively short third and fourth metacarpal bones in its wings.[7] It also lacks a tragus. The Greater Horseshoe Bat is, on average, between 57–71 millimetres (2.2–2.8 in) long, with a 35–43 mm tail (1.4–1.7 in) and a 350–400 mm wingspan (14–16 in);[6] Rhinolophus ferrumequinum also exhibits slight sexual dimorphism, with females being a bit larger than males.[8] The fur of the species is soft and fluffy, with the base of hairs being light grey, the dorsal side hair grey brown and the ventral side grey-white, with juvenile bats having more of an ash-grey tint to their fur. Wing membranes and ears are light grey-brown. The greater horseshoe bat weighs up to 30 grams (1.1 oz)[9] and can live up to 30 years.[10]

Distribution

The greater horseshoe bat ranges 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.[11] The northernmost occurrence is in Wales. Reaches to the southern parts of the Netherlands, Germany, Poland and Ukraine.[12] Generally the bats live below 800 m asl (above sea level), but it also lives, depending on roost availability and humidity, 3,000 m asl in the Caucasus.[1]

Ecology and behavior

Habitat and roosting

Pastures, deciduous temperate woodland, Mediterranean and sub-mediterranean shrubland and woodlands are common foraging habitats for this species.[1] In northern parts of its range, the horseshoe uses warm underground sites, both natural and artificial, as summer roosts as well as attics. Where the species occupies buildings, proximity to good foraging areas and underground sites for torpor at various times of year and for winter hibernation as well as the building's own features are important.[13]

Horseshoe bats hibernate in cold underground sites during the winter. The bats require a certain temperature and humidity limit, but this can vary with age, sex and condition.[1] Horseshoe bats are active throughout the year in the southern parts of their range. Horseshoe bats commonly travel distances of 20–30 km between winter and summer roosts, with longest distance recorded being 180 km.[14] Horseshoe bats also live in montane forests among the mountains and valleys of the Himalaya in South Asia and roosts in caves, old temples, old and ruined buildings in tight clusters.[1]

Diet and hunting

The species feeds preferentially on lepidopterans (moths), making up around 41% of the diet[15] - in particular the noctuid species.[16] For example, the species preys on the lesser wax moth by identifying the moth's high frequency mating call.[17] Coleopterans (beetles) constitute around 33% of the diet,[15] of which dung beetles and cockchafers are often taken. Aphodius rufipes is one such dung beetle forming an especially important part of its diet. (Cow pats are part of its life cycle, acting as a food source and habitat for the larvae. Up to 100 larvae can be found in a single cow pat; while the adult beetle is most abundant in August when the young bats begin their first feeding flights.)[18] The remainder of the diet consists of species of Hymenoptera and Diptera, as well as cave spiders.[15][8]

The feeding area from the maternity roost is typically of radius 4 km, as neither the lactating females or young can travel far. In late August and September the bats feed on cranefly, to fatten up before hibernation. Breeding females depend on beetles from April until June, and moths from June to August.

The greater horseshoe bat leaves its roost at dusk. Its flying is made up of slow, fluttering travel with short glides, normally between 0.3 and 6 metres above the ground. Little hunting is done during wet and windy weather. It hunts in terrain with poor tree cover such as hillsides and cliff faces, and in gardens where it locates insects from a resting place and then intercepts them. The species has the ability to pick food up off the ground while still in flight, and indeed drinks during low-level flight or while hovering. The feeding range of colonies in England is between 8 and 16 kilometres.

Mating and reproduction

Female bats become sexually mature at the age of three years while males are sexually mature at two years.[19] Some females may not breed until their fifth year.[20] Most matings take place in the fall, however some occur in the spring.[21] The reproductive behaviour of this species has been studied in detail. During the mating period, females visit males that are roosting in small caves. The mating system is best described as polygynous, with multiple females visiting males. However, genetic evidence has shown that some female greater horseshoe bats will visit and mate with the same male partner over successive years, indicating monogamy or mate fidelity.[22] Curiously, related females have also been found to share sexual partners, which might serve to increase relatedness and social cohesiveness in the colony.[22] Males vary in their reproductive success but do not tend to show reproductive skew within a year.[22] However, over several years, skew becomes strong due to the repeated success of particular males.[23]

After mating, seminal fluid coagulates in the female’s vulva to form a plug, which probably functions either to prevent subsequent matings by other males,[24] or to increase the chances that the sperm are retained for successful fertilization.[21] There is some evidence that females can eject these plugs, suggesting that they may be able to exert some control over fertilizations which take place.[4] Females raise their young in communal maternity roosts, and show strong fidelity to the sites where they themselves were born (so-called natal philopatry). Each season, a female produces one offspring. Most young are born in June or July.[4] When they are seven days old, young can open their eyes and at their third or fourth week they can fly. Young can leave the roost their seventh or eighth week.

Status and conservation

In general the greater horseshoe bat is listed as Least Concern by the IUCN because: "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".[1] However, the overall greater horseshoe bat population is declining.[1] They are largely uncommon in much of their range. Despite this, they appear to be abundant and widespread in at least parts of south-west Asia and the Caucasus. Also in some northwest European countries, there appears to be some stabilisation and/or recovery.[13] Less is known about bat trends in other parts of Europe. It is extinct in Malta.[1] Fragmentation/isolation of habitats, change of management regime of deciduous forests and agricultural areas, loss of insects caused by pesticides, and disturbance and loss of underground habitats and attics are the major threats to the greater horseshoe bat. Deforestation, mostly caused by logging operations and the conversion of land for agricultural and other uses, threatens the species in South Asia.[25]

Status in Britain

Woodcut from R. A. Sterndale, 1884

The species is rare in Britain, confined to just a small number of sites. Its distribution can be found on the National Biodiversity website here. Its breeding sites include Brockley Hall Stables near Bristol, Iford Manor near Bath, and Littledean Hall in the Forest of Dean. Its winter hibernation sites include Banwell Caves and Compton Martin Ochre Mine in the Mendip Hills, Chilmark Quarries in Wiltshire, and Combe Down and Bathampton Down Mines near Bath. In Dorset, the species roosts at Bryanston, Creech Grange and in Belle Vue Quarry. The species also occurs at Berry Head in Devon and has a monitored roost site at Woodchester Mansion in Stroud. It occurs in Wales, including at Felin Llwyngwair, a SSSI.

The species has disappeared from over half of its former range within the United Kingdom, with about 1% of the population surviving. Like all horseshoe bats it is sensitive to disturbance, and is threatened by the use of insecticides and the elimination of beetles by the changing agricultural practices.

There are seventeen recorded species of bat in Britain as of recent survey results. The greater horseshoe bat is one of the rarest. There are currently 35 recognised maternity and all-year roosts and 369 hibernation sites. Current estimates range between 4000 and 6600 individuals. Greater Horseshoes have declined for numerous reasons ranging from the use of agrichemicals (Ivermectin in particular) to loss of habitat and redundancy of farming methods. Avermectin kills off insect larvae and thus a decrease in the abundance of food for the Horseshoes, causing them to travel farther and face increased dangers.

Habitat loss is primarily due to the lack of established hedgerows and deciduous woodland-pasture ecotones. Modern farming methods have seen the reduction of cattle-grazing and this has impacted the Horseshoes who previously found that dung attracted insects and sustained entomogenous populations, giving their prey a stable population.

References

  1. ^ a b c d e f g h i Piraccini R. (2016). "Rhinolophus ferrumequinum". IUCN Red List of Threatened Species. 2016: e.T19517A21973253. doi:10.2305/IUCN.UK.2016-2.RLTS.T19517A21973253.en. Retrieved 17 February 2022.
  2. ^ Parsons, S. & Jones, G. 2000
  3. ^ Obrist, M.K. et al. 2004
  4. ^ a b c Schober, W., E. Grimmberger. 1997.
  5. ^ Nowak, R. 1994.
  6. ^ a b c Greater horseshoe bat Animal Diversity Web
  7. ^ Koopman, K. 1994.
  8. ^ a b Antoni Alcover, J., ed. (1988). "Mamífers Actuals". els Mamífers de les Balears. Palma de Mallorca: Editorial Moll. pp. 88–90. ISBN 84-273-0265-7.
  9. ^ Nature English, 1998
  10. ^ "A Guide to Bats of Britain and Europe. by Schober, W. & Grimmberger, E.: (1989) | C. Arden (Bookseller) ABA". www.abebooks.co.uk. Retrieved 2020-04-27.
  11. ^ Csorba G. P., Ujhelyi P. and Thomas, N. 2003.
  12. ^ Rhinolophus ferrumequinum - Science for Nature Foundation
  13. ^ a b Hutson A. M., Mickleburgh S. P., Racey P. A. 2001.
  14. ^ De Paz, O., Fernández, R., Benzal, J. 1986.
  15. ^ a b c Jones, G. 1990
  16. ^ Bat Conservation Trust Greater Horseshoe bat Species information leaflet
  17. ^ Cordes, N. et al. 2014.
  18. ^ Wild Devon The Magazine of the Devon Wildlife Trust, page 14, Winter 2009 edition
  19. ^ Ransome, R.D. 1995.
  20. ^ Racey, P. 1982.
  21. ^ a b Rossiter, S. et al. 2000
  22. ^ a b c Rossiter S. et al. 2005.
  23. ^ Rossiter, S. 2006.
  24. ^ Fenton, M. 1994. Just Bats. Toronto: University of Toronto Press.
  25. ^ Molur, S. et al. 2002.
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Greater horseshoe bat: Brief Summary

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The greater horseshoe bat (Rhinolophus ferrumequinum) is an insectivorous bat of the genus Rhinolophus. Its distribution covers Europe, Northern Africa, Central Asia and Eastern Asia. It is the largest of the horseshoe bats in Europe and is thus easily distinguished from other species. The species is sedentary, typically travelling up to 30 kilometres (19 mi) between the winter and summer roosts, with the longest recorded movement being 180 km (110 mi). The frequencies used by this bat species for echolocation lie between 69–83 kHz, have most energy at 81 kHz and have an average duration of 37.4 ms.

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