Overview

Brief Summary

Biology

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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Comprehensive Description

Description

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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Distribution

Range Description

This species has a wide range in the Palaearctic, occuring 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).
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Geographic Range

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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Range

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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Physical Description

Morphology

Physical Description

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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Ecology

Habitat

Habitat and Ecology

Habitat and Ecology
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 gestion period of 72 days (Bates and Harrison, 1997).

Systems
  • Terrestrial
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Habitat

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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Habitat

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

Food Habits

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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Associations

Known prey organisms

Rhinolophus ferrumequinum preys on:
Insecta

This list may not be complete but is based on published studies.
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Associations

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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Life History and Behavior

Life Expectancy

Lifespan/Longevity

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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Lifespan, longevity, and ageing

Maximum longevity: 30.5 years (wild)
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Reproduction

Reproduction

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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Molecular Biology and Genetics

Molecular Biology

Statistics of barcoding coverage: Rhinolophus ferrumequinum

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 0
Species: 9
Species With Barcodes: 1

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Conservation

Conservation Status

IUCN Red List Assessment


Red List Category
LC
Least Concern

Red List Criteria

Version
3.1

Year Assessed
2008

Assessor/s
Aulagnier, S., Hutson, A.M., Spitzenberger, F., Juste, J., Karataş, A., Palmeirim, J. & Paunovic, M.

Reviewer/s
Hutson, A.M., Racey, P.A. (Chiroptera Red List Authority) & Temple, H. (Global Mammal Assessment Team)

Justification
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.

History
  • 2000
    Lower Risk/near threatened
  • 1996
    Lower Risk/conservation dependent
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Conservation Status

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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Status

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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Population

Population
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 gone extinct in some countries (eg. Belgium, Netherlands). However, there are signs of stabilisation 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 gone 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), in Croatia the population is thought to be stable (N. Tvrtkovic pers. comm.), and in Romania the population 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.

Population Trend
Decreasing
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Threats

Threats

Major Threats
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).
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Threats

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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Management

Conservation Actions

Conservation Actions
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).
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Conservation

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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Relevance to Humans and Ecosystems

Benefits

Economic Importance for Humans: Positive

Positive Impacts: controls pest population

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Wikipedia

Greater Horseshoe Bat

The Greater Horseshoe Bat (Rhinolophus ferrumequinum) is a European bat of the Rhinolophus genus. Its distribution covers Europe, Africa, South Asia and Australia.[2] It is the largest of the European Horseshoe Bats and is thus easily distinguished from other species. The species is sedentary, travelling between 20 and 30 km between the winter and summer roosts, with the longest recorded movement being 180 km. The species is notable as having the oldest recorded age for any European bat, with a bat living for over 30 years. 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.[3][4]

Contents

Physical description

Grosse Hufeisennase-drawing.jpg

Its distinctive horseshoe noseleaf helps to focus the ultrasound it uses to 'see'.

The Greater Horseshoe Bat is on average between 57 and 71 mm long, with a 35–43 mm tail and a 350–400 mm wingspan. 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. It weighs up to 30 grams.[5] During the winter the bats hibernate in caves, abandoned mines or other underground places.

Habitat

Grand Rhinolophe 2.jpg

The Greater Horseshoe Bat lives in warmer regions of areas of open trees and scrub, near areas of standing water (e.g., ponds), areas of limestone and human settlement. The species is mainly house-dwelling in the north and cave-dwelling in the south. In the mountains nests are normally at below 800 metres above sea level. They are faithful to their summer and winter roosts, returning to the same sites each year.

Other favoured habitat include woodland with grassy rides and glades, old orchards, old 'veteran' trees with holes, permanent pasture, hedgerows linking the different features together.

Mating

Grand Rhinolophe.jpg

Females normally produce their young when around 4 years old in England and 3 years old in the south of Europe, with males becoming mature around the end of their second year. Mating season is from autumn to spring, with nurseries of up to 200 females clustered together with their young. The maternity roosts are usually found in old buildings, occasionally caves or abandoned mines.

The babies are born in June or July. Each bat only has one baby. Young Greater Horseshoe Bats open their eyes at about 4 days, are able to fly after three weeks and become independent at 7–8 weeks, during August. The false nipples of female bats, which newborn bats cling to after birth, are not fully developed until after the first birth.

Diet and hunting

The species feeds preferentially on lepidoptera (moths), making up around 41% of the diet by volume.[6] - in particular the noctuidae species [7], Coleoptera (beetles) constitute around 33% of the diet[6], of which dung beetles 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 food source and habitat for the larvae. Up to 100 larvae can be found in a single cow pat. The beetle is most abundant in August when the young bats begin their first feeeding flights).[8] The remainder of the diet being hymenoptera and diptera.[6] Cockchafers also form an important part of its diet.

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, and its flying is made up of slow, fluttering travel with short glides, normally between 0.3 and 6 metres above the ground, with little hunting during wet and windy weather. It hunts in terrain with poor tree cover such as hillsides, cliff faces and in gardens, locating insects from its resting place and then intercepting 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.

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 Biodivesity 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.

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 further and face increased dangers. Habitat loss is primarily 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.

Gallery

References

  1. ^ Chiroptera Specialist Group (2000). "Rhinolophus ferrumequinum". IUCN Red List of Threatened Species. Version 2.3. International Union for Conservation of Nature. http://www.iucnredlist.org/apps/redlist/details/19517. Retrieved 11 May 2006. 
  2. ^ Maurice Burton, Robert Burton The international wildlife encyclopedia, Volume 9 Marshall Cavendish, 2002, ISBN 0-7614-7266-5
  3. ^ Parsons, S. and Jones, G. (2000) 'Acoustic identification of twelve species of echolocating bat by discriminant function analysis and artificial neural networks.' J Exp Biol., 203: 2641–2656.
  4. ^ Obrist, M.K., Boesch, R. and Flückiger, P.F. (2004) 'Variability in echolocation call design of 26 Swiss bat species: Consequences, limits and options for automated field identification with a synergic pattern recognition approach.' Mammalia., 68 (4): 307–32.
  5. ^ Nature, English (1998). Managing Landscapes For The Greater Horseshoe Bat. Ruddocks (Lincoln) Ltd. ISBN 1-85716-416-4. 
  6. ^ a b c Jones, G. (1990), "Prey selection by the greater horsehoe bat (Rhinolophus ferrumequinum): Optimal foraging by echolocation?", Journal of Animal Ecology, 59: 587–602. 
  7. ^ Bat Conservation Trust Greater Horseshoe bat Species information leaflet
  8. ^ Wild Devon The Magazine of the Devon Wildlife Trust, page 14, Winter 2009 edition

Further reading

  • Schober, Wilfried; Eckard Grimmberger (1989). Dr. Robert E. Stebbings. ed. A Guide to Bats of Britain and Europe (1st ed.). UK: Hamlyn Publishing Group. ISBN 0-600-56424-X. 
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