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The greater horseshoe bat (''Rhinolophus ferrumequinum'') is an
insectivorous A robber fly eating a hoverfly An insectivore is a carnivorous animal or plant that eats insects. An alternative term is entomophage, which can also refer to the human practice of eating insects. The first vertebrate insectivores we ...
bat Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most bi ...
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 between the winter and summer roosts, with the longest recorded movement being . 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.


Description

The greater horseshoe bat is the largest horseshoe bat in Europe.Schober, W., E. Grimmberger. 1997. It has a distinctive noseleaf, which has a pointed upper part and a horseshoe-shaped lower part. Its horseshoe noseleaf helps to focus the
ultrasound Ultrasound is sound waves with frequencies higher than the upper audible limit of human hearing. Ultrasound is not different from "normal" (audible) sound in its physical properties, except that humans cannot hear it. This limit varies ...
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.Greater horseshoe bat
Animal Diversity Web
For other horseshoe bats, this premolar is very small or non-existent. Also in comparison to its relatives, the greater horseshoe bat has relatively short third and fourth metacarpal bones in its wings. It also lacks a tragus. The Greater Horseshoe Bat is, on average, between long, with a and a ; ''Rhinolophus ferrumequinum'' also exhibits slight
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most anim ...
, with females being a bit larger than males. 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 and can live up to 30 years.


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. The northernmost occurrence is in Wales. Reaches to the southern parts of the Netherlands, Germany, Poland and Ukraine. 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.


Ecology and behavior


Habitat and roosting

Pastures, deciduous temperate woodland, Mediterranean and sub-mediterranean shrubland and woodlands are common foraging habitats for this species. 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. Hutson A. M., Mickleburgh S. P., Racey P. A. 2001. 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. 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. 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.


Diet and hunting

The species feeds preferentially on
lepidoptera Lepidoptera ( ) is an order of insects that includes butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 families and 46 superfamilies, 10 percent of the total described specie ...
ns (moths), making up around 41% of the dietJones, G. 1990 - in particular the
noctuid The Noctuidae, commonly known as owlet moths, cutworms or armyworms, are a family (biology), family of moths. They are considered the most controversial family in the superfamily Noctuoidea because many of the clades are constantly changing, alon ...
species. For example, the species preys on the lesser wax moth by identifying the moth's high frequency mating call.
Coleoptera Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describe ...
ns (beetles) constitute around 33% of the diet, of which
dung beetle Dung beetles are beetles that feed on feces. Some species of dung beetles can bury dung 250 times their own mass in one night. Many dung beetles, known as ''rollers'', roll dung into round balls, which are used as a food source or breeding cha ...
s and
cockchafer The cockchafer, colloquially called Maybug, Maybeetle, or doodlebug, is the name given to any of the European beetles of the genus '' Melolontha'', in the family Scarabaeidae. Once abundant throughout Europe and a major pest in the periodical y ...
s 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 A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. Th ...
. 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.) The remainder of the diet consists of species of
Hymenoptera Hymenoptera is a large order of insects, comprising the sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are parasitic. Females typic ...
and
Diptera Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...
, as well as cave
spider Spiders ( order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species div ...
s. 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 Crane fly is a common name referring to any member of the insect family Tipulidae. Cylindrotominae, Limoniinae, and Pediciinae have been ranked as subfamilies of Tipulidae by most authors, though occasionally elevated to family rank. In the m ...
, to fatten up before hibernation. Breeding females depend on
beetles Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describe ...
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. Some females may not breed until their fifth year. Most matings take place in the fall, however some occur in the spring. Rossiter, S. et al. 2000 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 Polygyny (; from Neoclassical Greek πολυγυνία (); ) is the most common and accepted form of polygamy around the world, entailing the marriage of a man with several women. Incidence Polygyny is more widespread in Africa than in any ...
, 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 Monogamy ( ) is a form of dyadic relationship in which an individual has only one partner during their lifetime. Alternately, only one partner at any one time (serial monogamy) — as compared to the various forms of non-monogamy (e.g., polyg ...
or mate fidelity. Rossiter S. et al. 2005. Curiously, related females have also been found to share sexual partners, which might serve to increase relatedness and social cohesiveness in the colony. Males vary in their reproductive success but do not tend to show reproductive skew within a year. However, over several years, skew becomes strong due to the repeated success of particular males. 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, or to increase the chances that the sperm are retained for successful fertilization. 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. Females raise their young in communal maternity roosts, and show strong fidelity to the sites where they themselves were born (so-called natal
philopatry Philopatry is the tendency of an organism to stay in or habitually return to a particular area. The causes of philopatry are numerous, but natal philopatry, where animals return to their birthplace to breed, may be the most common. The term derives ...
). Each season, a female produces one offspring. Most young are born in June or July. 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". However, the overall greater horseshoe bat population is declining. 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. Less is known about bat trends in other parts of Europe. It is extinct in Malta. 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.Molur, S. et al. 2002.


Status in Britain

The species is rare in Britain, confined to just a small number of sites. Its distribution can be found on the National Biodiversity websit
here
Its breeding sites include
Brockley Hall Stables Brockley Hall Stables () is a 0.065 hectare biological Site of Special Scientific Interest near the village of Brockley, North Somerset, England notified in 1987. Biological Interest The SSSI designation applies to part of the former stable blo ...
near
Bristol Bristol () is a city, ceremonial county and unitary authority in England. Situated on the River Avon, it is bordered by the ceremonial counties of Gloucestershire to the north and Somerset to the south. Bristol is the most populous city in ...
, Iford Manor near Bath, and
Littledean Hall Littledean Hall () is a country house in the village of Littledean, Gloucestershire, England. It has been described as one of the most haunted houses in England and is thought to be the oldest house in the United Kingdom which is still occupied. S ...
in the
Forest of Dean The Forest of Dean is a geographical, historical and cultural region in the western part of the county of Gloucestershire, England. It forms a roughly triangular plateau bounded by the River Wye to the west and northwest, Herefordshire to the ...
. Its winter hibernation sites include
Banwell Caves Banwell Caves () are a 1.7-hectare geological and biological Site of Special Scientific Interest near the village of Banwell, North Somerset, England notified in 1963. The site comprises two caves, called 'Banwell Bone Cave' and 'Banwell Sta ...
and Compton Martin Ochre Mine in the
Mendip Hills The Mendip Hills (commonly called the Mendips) is a range of limestone hills to the south of Bristol and Bath in Somerset, England. Running from Weston-super-Mare and the Bristol Channel in the west to the Frome valley in the east, the hills o ...
,
Chilmark Quarries Chilmark Quarries () is a 9.65 hectare biological and geological Site of Special Scientific Interest (SSSI), in the ravine south of the village of Chilmark in Wiltshire, England. The SSSI was first notified in 1977. Its importance as a home ...
in
Wiltshire Wiltshire (; abbreviated Wilts) is a historic and ceremonial county in South West England with an area of . It is landlocked and borders the counties of Dorset to the southwest, Somerset to the west, Hampshire to the southeast, Gloucestershire ...
, and
Combe Down and Bathampton Down Mines Combe Down and Bathampton Down Quarries () make up a 6.22 hectare (15.37 acre) Site of Special Scientific Interest (SSSI) in Bath and North East Somerset, England, important for its bat population. The disused quarries date from the 17th and 18t ...
near Bath. In Dorset, the species roosts at
Bryanston Bryanston is a village and civil parish in north Dorset, England, situated on the River Stour west of Blandford Forum. In the 2011 census the parish had a population of 925. The village is adjacent to the grounds of Bryanston School, an ind ...
, Creech Grange and in Belle Vue Quarry. The species also occurs at Berry Head in
Devon Devon ( , historically known as Devonshire , ) is a ceremonial and non-metropolitan county in South West England. The most populous settlement in Devon is the city of Plymouth, followed by Devon's county town, the city of Exeter. Devon is a ...
and has a monitored roost site at
Woodchester Mansion Woodchester Mansion is an unfinished, Gothic revival mansion house in Nympsfield, Gloucestershire, England. It is on the site of an earlier house known as Spring Park. The mansion is a Grade I listed building. The mansion was abandoned by its b ...
in Stroud. It occurs in Wales, including at Felin Llwyngwair, a
SSSI A Site of Special Scientific Interest (SSSI) in Great Britain or an Area of Special Scientific Interest (ASSI) in the Isle of Man and Northern Ireland is a conservation designation denoting a protected area in the United Kingdom and Isle ...
. 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 Ivermectin (, '' EYE-vər-MEK-tin'') is an antiparasitic drug. After its discovery in 1975, its first uses were in veterinary medicine to prevent and treat heartworm and acariasis. Approved for human use in 1987, today it is used to treat inf ...
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


Literature cited

* Cordes, N., Engqvist, L., Schmoll T., Reinhold, K. 2014. Sexual signaling under predation: attractive moths take the greater risks. ''
Behavioral Ecology Behavioral ecology, also spelled behavioural ecology, is the study of the evolutionary basis for animal behavior due to ecological pressures. Behavioral ecology emerged from ethology after Niko Tinbergen outlined four questions to address when ...
''. 25(2):409–414. * Csorba G. P., Ujhelyi P., Thomas, N. 2003. ''Horseshoe Bats of the World''. Alana Books, Shropshire, England. * De Paz, O., Fernández, R., Benzal, J. 1986
El annilamiento de qirópteros en el centro de la Península Ibérica durante el periodo 1977-1986.
''Boletín de la Estación Central de Ecología''. 30:113–138 * Jones, G. 1990. Prey selection by the greater horseshoe bat (''Rhinolophus ferrumequinum''): Optimal foraging by echolocation? ''
Journal of Animal Ecology The ''Journal of Animal Ecology'' is a bimonthly peer-reviewed scientific journal publishing research in all areas of animal ecology. It began publication in 1932, and as such is the second oldest journal of the British Ecological Society (after ' ...
''. 59:587–602. * Koopman, K. 1994. ''Chiroptera: Systematics''. New York: Walter de Gruyter. * Obrist, M.K., Boesch, R., 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 Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur o ...
''. 68(4) 307–32 * Hutson A. M., Mickleburgh S. P., Racey P. A. 2001
Microchiropteran Bats - Global Status Survey and Conservation Action Plan.
IUCN/SSC Chiroptera Specialist Group, Gland, Switzerland and Cambridge, U.K. * Parsons, S. & Jones, G. 200
Acoustic identification of twelve species of echolocating bat by discriminant function analysis and artificial neural networks.
''
The Journal of Experimental Biology ''Journal of Experimental Biology'' (formerly ''The British Journal of Experimental Biology)'' is a peer-reviewed scientific journal in the field of comparative physiology and integrative biology. It is published by The Company of Biologists. ...
''. 2000 203: 2641-2656. * Molur, S., Marimuthu, G., Srinivasulu, C., Mistry, S. Hutson, A. M., Bates, P. J. J., Walker, S., Padmapriya, K. and Binupriya, A. R. 2002. Status of South Asian Chiroptera: Conservation Assessment and Management Plan (C.A.M.P.) Workshop Report. Zoo Outreach Organization/CBSG-South Asia, Coimbatore, India. * Nature English, 1998. ''Managing Landscapes For The Greater Horseshoe Bat''. Ruddocks (Lincoln) Ltd. * Nowak, R. 1994. ''Walker's Bats of the World''. Baltimore, Maryland: The Johns Hopkins University Press. * Racey, P. 1982. Ecology of Bat Reproduction. Pp. 57-93 in T. Kruz, ed. ''Ecology of Bats''. New York: Plenum Press. * Ransome, R.D. 1995. Earlier Breeding Shortens Life in Female Greater Horseshoe Bats. ''
Philosophical Transactions of the Royal Society B ''Philosophical Transactions of the Royal Society B: Biological Sciences'' is a biweekly peer-reviewed scientific journal published by the Royal Society. The editor-in-chief is John Pickett (Cardiff University). Overview Each issue covers a spe ...
''. 350:153–161. * Rossiter, S., Jones, J., Ransome, R., Barratt, E. 2000. Genetic variation and population structure in the endangered greater horseshoe bat ''Rhinolophus ferrmequinum''. ''
Molecular Ecology Molecular ecology is a field of evolutionary biology that is concerned with applying molecular population genetics, molecular phylogenetics, and more recently genomics to traditional ecological questions (e.g., species diagnosis, conservation and ...
''. 9:1131–1135 * Rossiter S., Ransome, R.D., Faulkes, C.G., Le Comber, S.L, Jones, G. 2005. Mate-fidelity and intra-lineage polygyny in greater horseshoe bats. ''
Nature Nature, in the broadest sense, is the physical world or universe. "Nature" can refer to the phenomena of the physical world, and also to life in general. The study of nature is a large, if not the only, part of science. Although humans are p ...
'' 437:408–411 * Rossiter, S., Ransome, R.D., Faulkes, C.G., Dawson, D.A., Jones, G. 2006. Long-term reproductive skew in male greater horseshoe bats. ''Molecular Ecology'' 15:3035–3043 * Schober, W., E. Grimmberger. 1997. ''The Bats of Europe and North America: Knowing Them, Identifying Them, Protecting Them''. New Jersey: TFH Publications Inc.


Further reading

*


External links


Greater Horseshoe BatWoodland Management For Bats Guide
{{Taxonbar, from=Q737413 Rhinolophidae Bats of Europe Bats of Asia Mammals of Afghanistan Mammals of Pakistan Horseshoe Bat, Greater Mammals of Bhutan Fauna of the British Isles Mammals described in 1774 Bats of India