|Lesser horseshoe bat (Rhinolophus hipposideros)|
Rhinolophidae is a family of bats commonly known as horseshoe bats. In addition to the single living genus, Rhinolophus, one extinct genus, Palaeonycteris, has been recognized. Horseshoe bats are closely related to the Hipposideridae.
Taxonomy and evolution
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. Attempts were made to divide Rhinolophus into other genera. In 1816, William Elford Leach proposed the genus name Phyllorhina; John Edward Gray proposed Aquias in 1847 and Phyllotis in 1866; and Wilhelm Peters proposed Coelophyllus in 1867. In 1876, George Edward Dobson returned all Asiatic horseshoe bats to Rhinolophus, additionally proposing the subfamilies Phyllorhininae (for the hipposiderids) and Rhinolophinae. Gerrit Smith Miller 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, a few other attempted to split Rhinolophus, naming Rhinolphyllotis in 1934 and Rhinomegalophus in 1951, though both additional genera were returned to Rhinolophus.
Knud Andersen was the first to propose species groups for Rhinolophus, doing so in 1905. 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 have been revised. 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.
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). The biogeography of horseshoe bats is poorly understood. Various studies have proposed that the family originated in Europe, Asia, or Africa. A 2010 study supported an Asian or Oriental origin of the family, with rapid evolutionary radiations of the African and Oriental clades during the Oligocene. A 2019 study found that R. xinanzhongguoensis and R. nippon, both Eurasian species, were more closely related to African species than to other Eurasian species, suggesting that rhinolophids may have a complex biogeographical relationship with Asia and the Afrotropics.
Rhinolophidae is represented by one extant genus, Rhinolophus. Both the family and the genus are confirmed as monophyletic (containing all descendants of a common ancestor). As of 2019, there were 106 described species in Rhinolophus, making it the second-most speciose genus of bat after Myotis. Rhinolophus may be undersampled in the Afrotropical realm, with one genetic study estimating that there could be up to twelve cryptic species in the region. Additionally, some taxa recognized as full species have been found to have little genetic divergence. Rhinolophus kahuzi may be a synonym for the Ruwenzori horseshoe bat (R. ruwenzorii), and R. gorongosae or R. rhodesiae may be synonyms of the Bushveld horseshoe bat (R. simulator). Additionally, Smithers's horseshoe bat (R. smithersi), Cohen's horseshoe bat (R. cohenae), and the Mount Mabu horseshoe bat (R. mabuensis) all have little genetic divergence from Hildebrandt's horseshoe bat (R. hildebrandtii). Recognizing the former three as full species leaves Hildebrandt's horseshoe bat paraphyletic.
The second genus in Rhinolophidae is the extinct Palaeonycteris, represented by one species, Palaeonycteris robustus. Palaeonycteris robustus lived during the Lower Miocene and its fossilized remains were found in Saint-Gérand-le-Puy, France.
Horseshoe bats are considered small or medium microbats. Individuals have a head and body length ranging from 35–110 mm (1.4–4.3 in) and have forearm lengths of 30–75 mm (1.2–3.0 in). One of the smaller species, the lesser horseshoe bat (R. hipposideros), weighs 4–10 g (0.14–0.35 oz), while one of the larger species, the greater horseshoe bat (R. ferrumequinum), weighs 16.5–28 g (0.58–0.99 oz). Fur color is highly variable among species, ranging from blackish to reddish brown to bright orange-red dorsal fur. The underparts are paler than the back fur. The majority of species have long, soft fur, but the woolly and lesser woolly horseshoe bats (R. luctus and R. beddomei) are unusual in their very long, woolly fur. Most have low or very low aspect ratios, which relates wingspan to wing area. Higher aspect ratios are associated with faster flight speed. Their wings are broad and rounded at the tips.
Like most bats, horseshoe bats have two mammary glands on their chests. Adult females additionally have two teat-like projections on their abdomens, called pubic nipples or false nipples, which are not connected to mammary glands. Only a few other bat families have pubic nipples, including Hipposideridae, Craseonycteridae, Megadermatidae, and Rhinopomatidae; they serve as attachment points for their offspring. In a few species, males have a false nipple in each armpit.
Head and teeth
All horseshoe bats have large, leaf-like protuberances on their noses, which are called nose-leafs. The anterior portion of the nose-leaf roughly resembles a horseshoe, earning them the common name of "horseshoe bats". The noseleafs are important in species identification, and are composed of several parts. The lancet is triangular, pointed, and pocketed, and points up between the bats' eyes. The sella is a flat, ridge-like structure at the center of the nose, rising from behind the nostrils, that points out perpendicular from the head. The sella usually has less hair than the lancet or the noseleaf. Their ears are large and leaf-shaped, nearly as broad as they are long, and lack tragi. The antitragi of the ears are conspicuous. Their eyes are very small. The skull always has rostral inflations. The typical dental formula of a horseshoe bat is 18.104.22.168, but the middle lower premolars are often missing, as well as the anterior upper premolars. The young lose their milk teeth while still in utero, with the teeth resorbed into the body. They are born with the four permanent canine teeth erupted, which enables them to cling to their mothers. This is atypical among bat families, as most newborns have at least some milk teeth at birth, which are quickly replaced by the permanent set.
Several bones in their thoraxes are fused—the presternum, first rib, partial second rib, seventh cervical vertebra, first thoracic vertebra—making a solid ring. Except for the first digit, which has two phalanges, all of their toes have three phalanges. This distinguishes them from hipposiderids, which have two phalanges in all toes. The tail is completely enclosed in the uropatagium (tail membrane), and the trailing edge of the uropatagium has calcars (cartilaginous spurs).
Biology and ecology
Echolocation and hearing
Horseshoe bats have some of the most sophisticated echolocation of any bat group. To echolocate, they produce sound through their nostrils. While some bats use frequency-modulated echolocation, horseshoe bats use constant-frequency echolocation (also known as single-frequency echolocation). They have high duty cycles, meaning that when individuals are calling, sound production composes more than 30% of total time. The use of high-duty, constant-frequency echolocation aids in distinguishing prey items based on size. These echolocation characteristics are typical of bats that search for moving prey items in cluttered environments. They echolocate at particularly high frequencies for bats, though not as high as hipposiderids relative to their body sizes, and the majority concentrate most of the echolocation energy into the second harmonic. The king horseshoe bat (R. rex) and the large-eared horseshoe bat (R. philippensis) are examples of outlier species that concentrate energy into the first harmonic rather than the second. Their highly furrowed noseleaves likely assist in focusing the emission of sound, reducing the effect of environmental clutter on sound. The noseleaf in general acts like a parabolic reflector, aiming the production of sound while simultaneously shielding the ear from some of it.
Horseshoe bats have sophisticated senses of hearing via well-developed cochlea, and are able to detect Doppler-shifted echoes. This allows them to produce and receive sounds simultaneously. Within horseshoe bats, there is a negative relationship between ear length and echolocation frequency: Species with higher echolocation frequencies tend to have shorter ear lengths. During echolocation, the ears can move independently of each other in a "flickering" motion characteristic of the family, while the head simultaenously moves up and down or side to side.
Diet and foraging
Horseshoe bats are insectivorous, though consume other arthropods like spiders, and employ two main foraging strategies. The first strategy is fly slow and low over the ground, hunting among trees and bushes. Some species who use this strategy are able to hover over prey and glean them from the substrate. The other strategy is known as perch feeding: Individuals roost on feeding perches and wait for prey to fly past, then fly out to capture it. While vesper bats may catch prey in their uropatagia and transfer it to their mouths, horseshoe bats do not use their uropatagia to catch prey. At least one species, the greater horseshoe bat, has been documented catching prey in the tip of its wing by bending the phalanges around it, then transferring it to their mouths.
Reproduction and life cycle
The mating systems of horseshoe bats are poorly understood. A review in 2000 noted that only about 4% of species had published information about their mating systems; along with the free-tailed bats (Molossidae), they had the least attention relative to their species diversity. At least one species, the greater horseshoe bat, appears to have a polygynous mating system where males attempt to establish and defend territories, attracting multiple females. Rhinolophus sedulus, however, is a rare species of bat that is believed to be monogamous (only 17 bat species are recognized as such as of 2000). Some species, particularly temperate species, have an annual breeding season in the fall, while other species mate in the spring. Many horseshoe bat species have the adaptation of delayed fertilization through female sperm storage. This is especially common in temperate species. In hibernating species, the sperm storage timing coincides with hibernation. Gestation takes approximately seven weeks before a single offspring is born, called a pup. Individuals reach sexual maturity by age two. While lifespans typically do not exceed six or seven years, some individuals may have extraordinarily long lives. A greater horseshoe bat individual was once banded and then rediscovered thirty years later.
Various levels of sociality are seen in horseshoe bats. Some species are solitary, with individuals roosting alone, while others are highly colonial, forming aggregations of thousands of individuals. The majority of species are moderately social. In some species, the sexes segregate annually when females form maternity colonies, though the sexes remain together all year in others. Individuals hunt solitarily. Because its hind limbs are poorly developed, it cannot scuttle on flat surfaces nor climb adeptly like other bats.
Range and habitat
Horseshoe bats have a mostly Paleotropical distribution, though some species are in the southern Palearctic realm. They are found in the Old World, including Africa, Australia, Asia, Europe, and Oceania. The greater horseshoe bat has the greatest geographic range of any horseshoe bat, occurring across Europe, North Africa, Japan, China, and southern Asia. Other species are much more restricted, like the Andaman horseshoe bat (R. cognatus), which is only found on the Andaman Islands. They roost in a variety of places, including buildings, caves, tree hollows, and foliage. They occur in both forested and unforested habitat.
Following the 2002 – 2004 SARS outbreak, several animal species were examined as possible natural reservoirs of the causative coronavirus, SARS-CoV. Several horseshoe bats were seropositive for SARS-like coronaviruses (testing positive for antibodies associated with it), tested positive for the viruses, or both. The least horseshoe bat (R. pusillus) was seropositive, the greater horseshoe bat tested positive for the virus only, and the big-eared horseshoe bat (R. macrotis), Chinese rufous horseshoe bat (R. sinicus), and Pearson's horseshoe bat (R. pearsoni) were seropositive and tested positive for the virus. The bats' viruses were highly similar to SARS-CoV, with 88–92% similarity. Though horseshoe bats appeared to be the natural reservoir of SARS-like coronaviruses, humans likely became sick through contact with infected masked palm civets, which were identified as intermediate hosts of the virus. From 2003–2018, forty-seven SARS-like coronaviruses were detected in bats, forty-five of which were found in horseshoe bats. Thirty SARS-like coronaviruses were from Chinese horseshoe bats, nine from greater horseshoe bats, two from big-eared horseshoe bats, two from the least horseshoe bat, and one each from the intermediate horseshoe bat (R. affinis), Blasius's horseshoe bat (R. blasii), Stoliczka's trident bat (Aselliscus stoliczkanus), and the wrinkle-lipped free-tailed bat (Chaerephon plicata).
Horseshoe bats have also tested positive for Mammalian orthoreovirus (MRV). A type 1 MRV was isolated from the lesser horseshoe bat and a type 2 MRV was isolated from the lesser horseshoe bat. The specific MRVs found in horseshoe bats have not been linked to human infection, though humans can become ill through exposure to other MRVs.
The rufous horseshoe bat (R. rouxii) has tested seropositive for Kyasanur Forest disease, which is a tick-borne viral hemorrhagic fever known from southern India. Kyasanur Forest disease is transmitted to humans through the bite of infected ticks, and has a mortality rate of 2–10%.
- Critically endangered: 1 species (Hill's horseshoe bat R. hilli)
- Endangered: 7 species
- Vulnerable: 6 species
- Near threatened: 8 species
- Least concern: 52 species
- Data deficient: 13 species
List of species
- Rhinolophus adami species group
- R. capensis species group
- R. euryale species group
- R. euryotis species group
- Arcuate horseshoe bat, R. arcuatus (see also Andersen's horseshoe bat)
- Rhinolophus belligerator
- Canut's horseshoe bat, R. canuti
- Croslet horseshoe bat, R. coelophyllus
- Creagh's horseshoe bat, R. creaghi
- Broad-eared horseshoe bat, R. euryotis
- Philippine forest horseshoe bat, R. inops
- Rhinolophus mcintyrei
- Rhinolophus proconsulis
- Large rufous horseshoe bat, R. rufus
- Shamel's horseshoe bat, R. shameli
- Small rufous horseshoe bat, R. subrufus
- R. ferrumequinum species group
- Bokhara horseshoe bat, R. bocharicus
- Geoffroy's horseshoe bat, R. clivosus
- Damara horseshoe bat, R. damarensis
- Darling's horseshoe bat, R. darlingi
- Decken's horseshoe bat, R. deckenii
- Greater horseshoe bat, R. ferrumequinum
- Hill's horseshoe bat, upland horseshoe bat, R. hillorum
- Sakeji horseshoe bat, R. sakejiensis
- Forest horseshoe bat, R. silvestris
- R. fumigatus species group
- R. hildebrandtii species group
- R. hipposideros species group
- Lesser horseshoe bat, R. hipposideros
- R. landeri species group
- R. maclaudi species group
- R. megaphyllus species group
- Intermediate horseshoe bat, R. affinis
- Homfray's horseshoe bat R. andamanensis
- Bornean horseshoe bat, R. borneensis
- Sulawesi horseshoe bat, R. celebensis
- Insular horseshoe bat, R. keyensis
- Madura horseshoe bat, R. madurensis
- Malayan horseshoe bat, R. malayanus
- Smaller horseshoe bat, R. megaphyllus
- Neriad horseshoe bat, R. nereis
- Peninsular horseshoe bat,R. robinsoni
- Lesser brown horseshoe bat, R. stheno
- Yellow-faced horseshoe bat, R. virgo
- R. pearsonii species group
- R. philippinensis species group
- R. pusillus species group
- Acuminate horseshoe bat, R. acuminatus
- Andaman horseshoe bat, R. cognatus
- Convex horseshoe bat, R. convexus
- Little Japanese horseshoe bat, R. cornutus
- Imaizumi's horseshoe bat, R. imaizumii
- Blyth's horseshoe bat, R. lepidus
- Formosan lesser horseshoe bat, R. monoceros
- Osgood's horseshoe bat, R. osgoodi
- Least horseshoe bat, R. pusillus
- Shortridge's horseshoe bat, R. shortridgei
- Little Nepalese horseshoe bat, R. subbadius
- R. rouxii species group
- R. trifoliatus species group
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