Horseshoe bat

Horseshoe bats are bats in the family Rhinolophidae. In addition to the single living genus, ''Rhinolophus'', which has about 106 species, the extinct genus '' Palaeonycteris'' has been recognized. Horseshoe bats are closely related to the Old World leaf-nosed bats, family Hipposideridae, which have sometimes been included in Rhinolophidae. The horseshoe bats are divided into six subgenera and many species groups. The most recent common ancestor of all horseshoe bats lived 34–40 million years ago, though it is unclear where the geographic roots of the family are, and attempts to determine its biogeography have been indecisive. Their taxonomy is complex, as genetic evidence shows the likely existence of many cryptic species, as well as species recognized as distinct that may have little genetic divergence from previously recognized taxa. They are found in the Old World, mostly in tropical or subtropical areas, including Africa, Asia, Europe, and Oceania.

Horseshoe bats are considered small or medium-sized microbats, weighing , with forearm lengths of and combined lengths of head and body of . The fur, long and smooth in most species, can be reddish-brown, blackish, or bright orange-red. They get their common name from their large nose-leafs, which are shaped like horseshoes. The nose-leafs aid in echolocation; horseshoe bats have highly sophisticated echolocation, using constant frequency calls at high duty cycles to detect prey in areas of high environmental clutters. They hunt insects and spiders, swooping down on prey from a perch, or gleaning from foliage. Little is known about their mating systems, but at least one species is monogamous, while another is polygynous. Gestation is approximately seven weeks and one offspring is produced at a time. A typical lifespan is six or seven years, but one greater horseshoe bat lived more than thirty years.

Horseshoe bats are relevant to humans in some regions as a source of disease, as food, and for traditional medicine. Several species are the natural reservoirs of various SARS-related coronaviruses, and data strongly suggests they are a reservoir of SARS-CoV, though humans may face more exposure risk from intermediate hosts such as masked palm civets.

They are hunted for food in several regions, particularly sub-Saharan Africa, but also Southeast Asia. Some species or their guano are used in traditional medicine in Nepal, India, Vietnam, and Senegal.

Taxonomy and evolution

Taxonomic history

''Rhinolophus'' was first described as a genus in 1799 by French naturalist Bernard Germain de Lacépède. Initially, all extant horseshoe bats were in ''Rhinolophus'', as well as the species now in '' Hipposideros'' (roundleaf bats). At first, ''Rhinolophus'' was within the family Vespertilionidae. In 1825, British zoologist John Edward Gray subdivided Vespertilionidae into subfamilies, including what he called Rhinolophina. English zoologist Thomas Bell is credited as the first to recognize horseshoe bats as a separate family, using Rhinolophidae in 1836. While Bell is sometimes recognized as the authority for Rhinolophidae, the authority is more often given as Gray, 1825. Horseshoe bats are in the superfamily Rhinolophoidea, along with Craseonycteridae, Hipposideridae Megadermatidae, Rhinonycteridae, and Rhinopomatidae.

Attempts were made to divide ''Rhinolophus'' into other genera. In 1816, English zoologist William Elford Leach proposed the genus name ''Phyllorhina''; Gray proposed ''Aquias'' in 1847 and ''Phyllotis'' in 1866; and German naturalist Wilhelm Peters proposed ''Coelophyllus'' in 1867. In 1876, Irish zoologist George Edward Dobson returned all Asiatic horseshoe bats to ''Rhinolophus'', additionally proposing the subfamilies Phyllorhininae (for the hipposiderids) and Rhinolophinae. American zoologist Gerrit Smith Miller Jr. further divided the hipposiderids from the horseshoe bats in 1907, recognizing Hipposideridae as a distinct family. Some authors have considered ''Hipposideros'' and associated genera as part of Rhinolophidae as recently as the early 2000s, though they are now most often recognized as a separate family. After the split into Rhinolophidae and Hipposideridae, further divisions were proposed for ''Rhinolophus'', with ''Rhinolphyllotis'' in 1934 and ''Rhinomegalophus'' in 1951, though both additional genera were returned to ''Rhinolophus''.

Danish mammalogist Knud Andersen was the first to propose species groups for ''Rhinolophus'', doing so in 1905. Species groups are a way of clustering species to reflect evolutionary relationships. He recognized six species groups: ''R. simplex'' (now ''R. megaphyllus''), ''R. lepidus'', ''R. midas'' (now ''R. hipposideros''), ''R. philippinensis'', ''R. macrotis'', and ''R. arcuatus''. The species have been frequently rearranged among the groups as new groups are added, new species are described, and relationships among species are revised. Fifteen species groups were given by Csorba and colleagues in 2003. Various subgenera have been proposed as well, with six listed by Csorba ''et al''. in 2003: ''Aquias'', ''Phyllorhina'', ''Rhinolophus'', ''Indorhinolophus'', ''Coelophyllus'', and ''Rhinophyllotis''. Informally, the rhinolophids can be split into two major clades: the mostly African clade, and the mostly Oriental clade.

Evolutionary history

The most recent common ancestor of ''Rhinolophus'' lived an estimated 34–40 million years ago, splitting from the hipposiderid lineage during the Eocene. Fossilized horseshoe bats are known from Europe (early to mid-Miocene, early Oligocene), Australia (Miocene), and Africa (Miocene and late Pliocene

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