Horseshoe bats (Rhinolophidae) are a family of bats. In addition to the single living genus, Rhinolophus, there is one extinct genus, Palaeonycteris. The closely related Hipposideridae are sometimes included within the horseshoe bats as a subfamily, Hipposiderinae. Both families are classified in the suborder Yinpterochiroptera or Pteropodiformes and were previously included in Microchiroptera.

Appearance

All horseshoe bats have leaf-like, horseshoe-shaped protuberances on their noses. In the related Hipposideridae, these noseleafs are leaf- or spear-like. They emit echolocation calls through these structures, which may serve to focus the sound. Their hind limbs are not well developed, so that they cannot walk on all fours; conversely, their wings are broad, making their flight particularly agile. Most rhinolophids are dull brown or reddish brown in color. They vary in size from 2.5 cm to 14 cm in head-body length, and 4 to 120 grams in weight (Macdonald, 1984). Their dental formula is .

The females have a pair of mammary glands and two "false nipples" above and to the side of the genital opening, which newborn bats cling to for a few days after birth.

Ecology

Rhinolophids inhabit temperate and tropical regions of southern Europe, Africa, and Asia south to northern and eastern Australia. All species are insectivorous, capturing insects in flight. Their roost habits are diverse; some species are found in large colonies in caves, some prefer hollow trees, and others sleep in the open, among the branches of trees. Members of northern populations may hibernate during the winter, while a few are known to aestivate; at least one species is migratory. Like many Vespertilionidae bats, females of some rhinolophid species mate during the fall and store the sperm over the winter, conceiving and gestating young beginning in the spring.

Classification

Horseshoe bats are closely related to the family Hipposideridae, which is often included within Rhinolophidae; however, it is now considered a separate family (Simmons, 2005; Hutcheon and Kirsch, 2006). In addition to the sole living genus, Rhinolophus, Rhinolophidae contains one extinct genus, Palaeonycteris (McKenna and Bell, 1997). Many species are extremely difficult to distinguish.

Although horseshoe bats have traditionally been included in the suborder Microchiroptera ("microbats"), genetic evidence suggests they and a few other microbat families are more closely related to Pteropodidae, the only family of "megabats" (Megachiroptera). Therefore, Pteropodidae, horseshoe bats, and related families are now placed in a single suborder, called Yinpterochiroptera or Pteropodiformes (Hutcheon and Kirsch, 2006).

Medical significance

In September 2005, four Rhinolophus species (R. sinicus, R. ferrumequinum, R. macrotis, R. pearsoni) were identified as natural reservoirs of SARS coronavirus-like viruses, the causative agent of SARS outbreaks in 2002–2004 (Li et al., 2005; Lau et al., 2005).

List of species

Genus Rhinolophus

• P. adami species group
  • Adam's Horseshoe Bat, Rhinolophus adami
  • Maendeleo Horseshoe Bat, Rhinolophus maendeleo

• P. capensis species group
  • Cape Horseshoe Bat, Rhinolophus capensis
  • Dent's Horseshoe Bat, Rhinolophus denti
  • Bushveld Horseshoe Bat, Rhinolophus simulator
  • Swinny's Horseshoe Bat, Rhinolophus swinnyi

• P. euryale species group
  • Mediterranean Horseshoe Bat, Rhinolophus euryale
  • Mehely's Horseshoe Bat, Rhinolophus mehelyi

• P. euryotis species group
  • Arcuate Horseshoe Bat, Rhinolophus arcuatus (see also Andersen's Horseshoe Bat)
  • Canut's Horseshoe Bat, Rhinolophus canuti
  • Croslet Horseshoe Bat, Rhinolophus coelophyllus
  • Creagh's Horseshoe Bat, Rhinolophus creaghi
  • Broad-eared Horseshoe Bat, Rhinolophus euryotis
  • Philippine Forest Horseshoe Bat, Rhinolophus inops
  • Large Rufous Horseshoe Bat, Rhinolophus rufus
  • Shamel's Horseshoe Bat, Rhinolophus shameli
  • Small Rufous Horseshoe Bat, Rhinolophus subrufus

• P. ferrumequinum species group
  • Bokhara Horseshoe Bat, Rhinolophus bocharicus
  • Geoffroy's Horseshoe Bat, Rhinolophus clivosus
  • Darling's Horseshoe Bat, Rhinolophus darlingi
  • Decken's Horseshoe Bat, Rhinolophus deckenii
  • Greater Horseshoe Bat, Rhinolophus ferrumequinum
  • Upland Horseshoe Bat (Hill's Horseshoe Bat), Rhinolophus hillorum
  • Sakeji Horseshoe Bat, Rhinolophus sakejiensis
  • Forest Horseshoe Bat, Rhinolophus silvestris

• P. fumigatus species group
  • Eloquent Horseshoe Bat, Rhinolophus eloquens
  • Rüppell's Horseshoe Bat, Rhinolophus fumigatus
  • Hildebrandt's Horseshoe Bat, Rhinolophus hildebrandtii

• P. hipposideros species group
  • Lesser horseshoe bat, Rhinolophus hipposideros

• P. landeri species group
  • Halcyon Horseshoe Bat, Rhinolophus alcyone
  • Blasius's Horseshoe Bat, Rhinolophus blasii
  • Guinean Horseshoe Bat, Rhinolophus guineensis
  • Lander's Horseshoe Bat, Rhinolophus landeri

• P. maclaudi species group
  • Maclaud's Horseshoe Bat, Rhinolophus maclaudi
  • Ruwenzori Horseshoe Bat, Rhinolophus ruwenzorii

• P. megaphyllus species group
  • Intermediate Horseshoe Bat, Rhinolophus affinis
  • Bornean Horseshoe Bat, Rhinolophus borneensis
  • Sulawesi Horseshoe Bat, Rhinolophus celebensis
  • Malayan Horseshoe Bat, Rhinolophus malayanus
  • Smaller Horseshoe Bat, Rhinolophus megaphyllus
  • Neriad Horseshoe Bat, Rhinolophus nereis
  • Lesser Brown Horseshoe Bat, Rhinolophus stheno
  • Yellow-faced Horseshoe Bat, Rhinolophus virgo

• P. pearsonii species group
  • Pearson's Horseshoe Bat, Rhinolophus pearsonii
  • Dobson's Horseshoe Bat, Rhinolophus yunanensis

• P. philippinensis species group
  • Big-eared Horseshoe Bat, Rhinolophus macrotis
  • Marshall's Horseshoe Bat, Rhinolophus marshalli
  • Timorese Horseshoe Bat, Rhinolophus montanus
  • Bourret's Horseshoe Bat, Rhinolophus paradoxolophus
  • Large-eared Horseshoe Bat, Rhinolophus philippinensis
  • King Horseshoe Bat, Rhinolophus rex

• P. pusillus species group
  • Acuminate Horseshoe Bat, Rhinolophus acuminatus
  • Andaman Horseshoe Bat, Rhinolophus cognatus
  • Convex Horseshoe Bat, Rhinolophus convexus
  • Little Japanese Horseshoe Bat, Rhinolophus cornutus
  • Imaizumi's Horseshoe Bat, Rhinolophus imaizumii
  • Blyth's Horseshoe Bat, Rhinolophus lepidus
  • Formosan Lesser Horseshoe Bat, Rhinolophus monoceros
  • Osgood's Horseshoe Bat, Rhinolophus osgoodi
  • Least Horseshoe Bat, Rhinolophus pusillus
  • Shortridge's Horseshoe Bat, Rhinolophus shortridgei
  • Little Nepalese Horseshoe Bat, Rhinolophus subbadius

• P. rouxii species group
  • Rufous Horseshoe Bat, Rhinolophus rouxii
  • Chinese Rufous Horseshoe Bat, Rhinolophus sinicus
  • Thomas's Horseshoe Bat, Rhinolophus thomasi

• P. trifoliatus species group
  • Lesser Woolly Horseshoe Bat, Rhinolophus beddomei
  • Formosan Woolly Horseshoe Bat, Rhinolophus formosae
  • Woolly Horseshoe Bat, Rhinolophus luctus
  • Lesser woolly Horseshoe Bat, Rhinolophus sedulus
  • Trefoil Horseshoe Bat, Rhinolophus trifoliatus

• incertae sedis
  • Mitred Horseshoe Bat, Rhinolophus mitratus

• Rhinolophus chiewkweeae
• Hill's Horseshoe Bat, Rhinolophus hilli
• Rhinolophus huananus
• Insular Horseshoe Bat, Rhinolophus keyensis
• Madura Horseshoe Bat, Rhinolophus madurensis
• Rhinolophus microglobosus
• Rhinolophus robinsoni
• Rhinolophus schnitzleri
• Thai Horseshoe Bat, Rhinolophus siamensis
• Rhinolophus thailandensis
• Rhinolophus xinanzhongguoensis
• Ziama Horseshoe Bat, Rhinolophus ziama

References

• Corbet, G.B. and Hill, J.E. 1992. The mammals of the Indomalayan region: a systematic review. Oxford: Oxford University Press.
• Hutcheon, J.M. and Kirsch, J.A.W. 2006. A moveable face: deconstructing the Microchiroptera and a new classification of extant bats. Acta Chiropterologica 8(1):1–10.
• Kock, D., Csorba, G. and Howell, K.M. 2000. Rhinolophus maendeleo n. sp. from Tanzania, a horseshoe bat noteworthy for its systematics and biogeography (Mammalia, Chiroptera, Rhinolophidae). Senckenbergiana biologica 80:233–239.
• Lau, S., Woo, P., Li, K., et al. 2005. Severe acute respiratory syndrome coronavirus-like virus in Chinese horseshoe bats. Proceedings of the National Academy of Sciences 102(39):14040–14045.
• Li, W., Zhengli, S., Meng, Y., et al. 2005. Bats are natural reservoirs of SARS-like coronaviruses. Science 310(5748):676–679.
• Macdonald, D. 1984. The Encyclopedia of Mammals. New York: Facts on File, 805 pp. ISBN 0-87196-871-1
• McKenna, M.C. and Bell, S.K. 1997. Classification of Mammals: Above the species level. New York: Columbia University Press, 631 pp. ISBN 978-0-231-11013-6
• Schober, W. and Grimmberger, A. 1989. A Guide to Bats of Britain and Europe. Hamlyn Publishing Group. ISBN 0-600-56424-X
• Simmons, N.B. 2005. Order Chiroptera. Pp. 312–529 in Wilson, D.E. and Reeder, D.M. (eds.). Mammal Species of the World: a taxonomic and geographic reference. 3rd ed. Baltimore: The Johns Hopkins University Press, 2 vols., 2142 pp. ISBN 978-0-8018-8221-0
• Corbet, G.B. 2008. Taxonomy of the Horseshoe bats of the World (Chiroptera: Rhinolophidae). http://dea.unideb.hu/dea/bitstream/2437/89636/4/ertekezes_angol.pdf
• Zhou, Z.-M., Guillén-Servent A., Kim, B.K., Eger, J.L., Wang, Y.Y. and Jiang, X.-L. 2009. A new species from southwestern China in the Afro-Palearctic lineage of the horseshoe bats (Rhinolophus). Journal of Mammalogy 90:57–73.
• Wu, Y., Harada, M. and Motokawa, M. 2009. Taxonomy of Rhinolophus yunanensis Dobson, 1872 (Chiroptera: Rhinolophidae) with a description of a new species from Thailand. Acta Chiropterologica 11(2):237–246.

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