Hipposideridae, Old World leaf-nosed bats, is one of eighteen families that make up the order Chiroptera, and consists of nine genera. The largest genus is Hipposideros (roundleaf bats), which consists of 76 species. The remaining genera are Asellia (trident leaf-nosed bats), Anthops (flower-faced bats), Aselliscus (trident bats), Cloeotis (Percival's trident bat), Coelops (tailless leaf-nosed bats), Paracoelops (orange leaf-nosed bat), Rhinonicteris (orange leaf-nosed bat), and Triaenops (trident bats). Hipposiderids live in tropical and subtropical regions of Africa, south Asia, Australia, the Philippine Islands, and the Solomon Islands. They can be found in deserts, dunes, savannas, grasslands, forests, rainforests, scrub forests and mangroves. Most species roost in dark, enclosed spaces, but some do roost in open areas. Hipposiderids range from 28 to 110 mm in body length, 30 to 110 mm in forearm (wing) length, and may or may not have a tail, up to 60 mm in length. Colors range from white to red to dark brown depending on species, geographic area, sex, and age, and pelage also varies in length and texture. They may have small or large ears, and some species’ ears are interconnected along the dorsal surface of the head. The appearance of the noseleaf is highly variable among genera. Hipposiderids show a great deal of diversity in roosting behavior and reproductive habits and show slight differences in feeding habits from genus to genus.
- Hill, J., J. Smith. 1992. Bats: A Natural History. Austin, TX: University of Texas Press.
- Kunz, T., P. Racey. 1998. Bat Biology and Conservation. Washington and London: Smithsonian Institution Press.
- Nowak, R. 1994. Walker's Bats of the World. Baltimore: Johns Hopkins University Press.
- Wilson, D., D. Reeder. 2005. Mammal Species of the World. A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press.
Members of Hipposideridae are found throughout tropical and subtropical regions of the Old World. These Old World leaf-nosed bats are found in Africa, southern Asia, the Philippine Islands, the Solomon Islands, and Australia.
Biogeographic Regions: palearctic (Native ); oriental (Native ); ethiopian (Native ); australian (Native )
- Simmons, N., T. Conway. 1997. "Tree of Life Web Project" (On-line). Rhinolophidae. Horseshoe Bats.. Accessed February 26, 2009 at http://tolweb.org/Rhinolophidae/16126/1997.01.01.
A defining characteristic of Hipposiderids is their elaborate noseleaf. The noseleaf consists of fleshy protrusions on top of a U-shaped rhinarium (i.e., the wet surface surrounding the nostrils). Hipposiderids have an erect transverse leaf within the noseleaf as well as smaller accessory leaflets. The common name of many genera corresponds to the shape of the noseleaf. For example, flower-faced bats have two circular lateral leaflets, the smaller of which is superimposed onto the larger, resulting in a noseleaf resembling the petals of a flower. Differences in noseleaf characteristics are commonly used to discern between genera. These 'appendages' are thought to be related to nasal echolocation, and may help to focus and modify echolocation signals.
Pelage of hipposiderids varies greatly both between and within taxa. Pelage can be white, light beige, pale yellow, dark yellow, orange, red, red-brown, light brown, dark brown, gray, or dark gray. Some species have white patches of fur, while others have 2 different color phases. Often, venter pelage is lighter than dorsum pelage. Pelage also differs interspecifically by length and texture or silkiness. Hipposiderid skulls have a number of unique features that differentiate them from other bat families. For example, they have no post-orbital processes, the nasal portion of the premaxilla is absent, and the premaxilla is not fused to the maxilla. They have dilambdodont molars , and their dental formula is I1/2 C1/1 P1–2/2–3 M3/3, giving them a total of 28 to 30 teeth. Hipposiderids do not have a tragus, the fleshy protuberance present at the opening of the ear in many bats. A membrane spanning the dorsal surface of the head connects their ears, which exhibit a great deal of variation in size.
Hipposideridae shares many traits with the family Rhinolophidae, and some accounts consider Hipposideridae a sub-family of Rhinolophidae. Both hipposiderids and rhinolophids lack post-orbital processes and the nasal portion of the premaxilla, as well as having a premaxilla that is not fused to the maxilla, dilambdodont molars, and a U- or horseshoe-shaped rhinarium. However, hipposiderids can be differentiated from rhinolophids using a number of different characteristics. Hipposiderids generally have a more rounded noseleaf, while the noseleaf of rhinolophids is spear-like and pointed. Hipposiderids have only two bones in each toe, while rhinolophids have three in all except the first toe, which has two. Rhinolophids always have three lower premolars on each side of the mandible and hipposiderids have only two. The two families also differ in the structure of their shoulder and hip girdles. Finally, rhinolophids have a sella, a flattened leaflet in the middle of the noseleaf structure, that is not present in hipposiderids.
Other Physical Features: endothermic ; heterothermic ; homoiothermic; bilateral symmetry
Sexual Dimorphism: sexes alike
- 2009. "Leaf-nosed bat" (On-line). Encyclopaedia Britannica Online. Accessed February 27, 2009 at http://www.britannica.com/EBchecked/topic/693386/leaf-nosed-bat.
- Dewey, T. 2011. "Hipposiderinae (Old World leaf-nosed bats)" (On-line). Grzimek's Animal Life. Accessed April 25, 2011 at http://animals.galegroup.com.
- Jones, G. 2001. Bats. Pp. 754-785 in D MacDonald, ed. The Encyclopedia of Mammals. Oxfordshire, UK: Andromeda Oxford Limited.
Hipposideridae inhabits tropical and subtropical habitats and roosting preferences vary by genera. Hipposiderids have been found roosting in caves, mines, hollow trees, buildings, and man-made underground compartments like cellars and tombs. In Africa, Fulvus round-leaf bats are often found in the burrows of Hystrix (Old World porcupines) and members of the genus Asellia roost in the inner walls of wells, in caves, and in man-made structures. Though Percival's trident bats live in forests and generally roost in trees, in Taiwan they have been discovered in abandoned Japanese bomb shelters, also known as pillboxes.
Habitat Regions: tropical ; terrestrial
Terrestrial Biomes: desert or dune ; savanna or grassland ; forest ; rainforest ; scrub forest
Although little information is available on the diets of most hipposiderid species, they are considered to be primarily insectivorous. Those species that have been studied prefer cicadas, cockroaches, termites, and beetles. The beetle larvae prey of Commerson's roundleaf bats live in wild figs, which results in the addition of small amounts of fruit to their otherwise insectivorous diet.
Hipposiderids have excellent echolocation, and catch most of their prey via aerial hawking and gleaning. They usually fly only a few meters above the ground while echolocating for potential prey. Although most species are thought to prey on flying insects, some occasionally feed on flightless insects such as ants. Hipposiderids are generally territorial and hunt and feed within a specific range. For example, members of the genus Asellia, have been observed flying more than a mile through the desert to their feeding territory. Often, hipposiderids bring captured prey back to their roost prior to consumption. When chewing, the jaws of hipposiderids move side-to-side and up and down, simultaneously. This shearing motion helps break down the chitinous exoskeleton of insect prey.
Primary Diet: carnivore (Insectivore )
- Graham, G., F. Reid. 1994. Bats of the World. New York: St Martin's Press.
- Nowak, R. 1991. Walker's Mammals of the World: Fifth Edition. Baltimore and London: The Johns Hopkins University Press.
- Walton, D., B. Richardson. 1989. Fauna of Australia Volume 1B: Mammalia. Canberra, Australia: Australian Government Publishing Service.
As insectivores, hipposiderids help control insect pest populations. While little information exists on potential endoparasites of hipposiderids, like most bats, they are probably host to a number of ectoparasitic arthropods including lice, mites and fleas.
Hipposiderids are preyed upon by a number of small nocturnal mammals with the ability to capture them mid-flight or locate their roosts. In many localities, the major predator of hipposiderids is snakes, which are sometimes able to locate their roosting sites. During flight, hipposiderids can be captured and eaten by various birds of prey including hawks, falcons, and owls. Furthermore, in Australia, members of the family Dasyuridae have been known to locate hipposiderid roosts. In conjunction with their ability to fly, the nocturnal lifestyle of bats helps reduce predation as does the colonial roosting behavior of many species.
Life History and Behavior
Like most Microchiroptera, members of the family Hipposideridae have relatively small eyes, indicating that vision may not be as important as echolocation for navigation and foraging purposes. However, vision may be used to detect objects past the range of echolocation. Hipposiderids, like all Microchiroptera, do not have color vision.
Unlike most microchiropterans that emit echolocation via the mouth, hipposiderids produce echolocation sounds with the larynx and emit the sound through their nostrils. The sounds produced are considered ultrasonic because they have higher frequencies than the normal range of human hearing. Hipposiderid echolocation calls contain a long constant-frequency (CF) component (i.e., one frequency is maintained throughout the duration of the call) and a much shorter frequency-modulated (FM) component. The CF segment of the call is used to determine the general structure of the local environment and to give a coarse location of potential prey and is preceded or followed by a brief FM segment, which aids in homing in on the location of a target. In general, the calls of larger bats have a tendency to be longer and lower in frequency, whereas the calls of smaller bats have a tendency to be shorter and higher in frequency. Evidence suggests that the calls of hipposiderids are typically higher in frequency relative to body mass than other bat families. Little information is available regarding the use of sound and echolocation for intraspecific communication, though audible sounds may be used to communicate during courtship or between mother and pup.
Many hipposiderid species have a small sac just posterior to the nose leaf. The sac, which is possessed primarily by males, secretes a waxy substance that may be used during mating season to attract mates or fend of potential rivals.
Communication Channels: acoustic ; chemical
Perception Channels: visual ; tactile ; acoustic ; ultrasound ; echolocation ; chemical
- Thomas, J., C. Moss, M. Vater. 2004. Echolocation in Bats and Dolphins. Chicago: University of Chicago Press.
Information regarding the lifespan of hipposiderids is limited, as a majority of species in this family are not well-known. However, some species have been found to live more than 10 years.
Not enough information is known about hipposiderid mating systems to make accurate generalizations about the family as a whole; however, research on individual species provides limited but important insight. Only one example of a polygynous mating in hipposiderids is known. Colonies of Commerson's roundleaf bat, which can contain up to 500,000 individuals, are divided into small harems consisting of one adult male and several adult females, with whom the male mates. Mating occurs seasonally, during the fall, and females give birth to a single young during spring after storing sperm over winter.
Mating System: polygynous
Breeding season and birthing season vary among hipposiderid species. For example, bi-colored leaf-nosed bats and ashy roundleaf bats mate in October and give birth in April. Although birthing season varies slightly, coinciding with peak rainy season when food is most abundant, Sundevall's leafnosed bats give birth in April north of the equator and in October south of the equator. Fulvus roundleaf bats mate in November and give birth in late April. Although the specific times vary among species, birthing among hipposiderids generally occurs during spring. Female hipposiderids give birth to a single young per pregnancy. Gestation lasts from 90 days in cyclops roundleaf bats to 220 days in Sundevall's leafnosed bats in South Africa. Females typically carry their young for a few weeks after giving birth. For example, Fulvus roundleaf bats produce a single young, which the female carries for 20 to 22 days. Age at weaning, age at first flight, and age at independence appears to vary according to latitude. Species subject to greater seasonality appear to mature more quickly than those resident to more tropical regions. In at least one species, Sundevall's leafnosed bats of Nigeria, delayed implantation occurs. The egg does not implant in the uterine lining for up to 2 months after fertilization, and as a result, young are born when prey are more abundant, directly before the rainy season.
Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; viviparous ; delayed fertilization ; delayed implantation
Females are the primary care givers in hipposiderids. Female typically carry their young for a few weeks after birth and prior to weaning. Females have "pubic teats", which their young hold on to during the carrying period. Little is known of lactation and weaning in hipposiderids. However, lactation lasts for about 40 days in the genus Asellia, and Taiwanese leaf-nosed bats are usually weaned at 7 weeks old. Tropical species are thought to be weaned by 8 to 20 weeks and time to independence appears to vary according to latitude, as tropical species reach sexual maturity between 16 and 24 months, and temperate species reaching sexual maturity by 6 to 8 months.
Parental Investment: precocial ; female parental care ; pre-hatching/birth (Provisioning: Female); pre-weaning/fledging (Protecting: Female); pre-independence (Provisioning: Female)
- Dewey, T. 2011. "Hipposiderinae (Old World leaf-nosed bats)" (On-line). Grzimek's Animal Life. Accessed April 25, 2011 at http://animals.galegroup.com.
- Nowak, R. 1994. Walker's Bats of the World. Baltimore: Johns Hopkins University Press.
- Slaughter, B., D. Walton. 1970. About Bats: A Chiropteran Biology Symposium. Dallas: Southern Methodist University Press.
As a family, hipposiderids are not a particularly threatened group. However, many species are not well understood and as a result, potential conservation needs are unknown. The International Union for Conservation of Nature (IUCN) lists 5 species as either endangered or critically endangered, and another 10 species are listed as vulnerable. Of the 84 species listed, 7 are classified as near threatened, 44 are listed as least concern, and the remaining 18 are classified as data deficient. Habitat loss and deforestation are serious concerns and their greatest threats. In specific cases, habitat loss has been so extreme that several species are now classified as endangered and some local populations are nearing extirpation. For example, Thailand leaf-nosed bats have been subjected to severe range contraction due to deforestation, which has resulted in a population reduction of 20% in just the last 5 years. Durga Das's leaf-nosed bats have had nearly all of their native range destroyed and now only roost in the homes of three different villages in central India. Due to deforestation, Orbiculus leaf-nosed bats are now resident to only two locations in Indonesia and Malaysia. Lamotte's roundleaf bats can be found in a single cave on the island of Guinea, and are classified as critically endangered. If conservation efforts are to be successful, habitat loss must be slowed and reforestation projects should be encouraged in critical habitat areas.
- IUCN, 2008. "IUCN 2008 Red List" (On-line). Accessed February 15, 2009 at http://www.iucnredlist.org/search.
Relevance to Humans and Ecosystems
Hipposiderids cause little economic damage. There are no known pathogens specific to Hipposideridae that are harmful to people or domesticated animals. However, bats occasionally roost in occupied buildings, which can be destructive and has the potential to spread disease. Any species of bat infected with rabies could potentially bite and transmit the pathogen to humans.
Negative Impacts: injures humans (causes disease in humans , carries human disease); causes or carries domestic animal disease ; household pest
As insectivores, hipposiderids help control insect pest populations that might otherwise spread disease or damage crops. The guano of hipposiderids is locally used as a nitrogen rich fertilizer.
Positive Impacts: produces fertilizer; controls pest population
The Hipposideridae are a family of bats commonly known as the Old World leaf-nosed bats. While it has often been seen as a subfamily, Hipposiderinae, of the family Rhinolophidae, it is now more generally classified as its own family. Nevertheless, it is most closely related to Rhinolophidae within the suborder Pteropodiformes (or Yinpterochiroptera).
The Hipposideridae contain 10 living genera and more than 70 species, mostly in the widespread genus Hipposideros. In addition, several fossil genera are known; the oldest fossils attributed to the family are from the middle Eocene of Europe. In their 1997 Classification of Mammals, Malcolm C. McKenna and Susan K. Bell proposed a division of Hipposideridae (called Rhinonycterinae in their work) into three tribes, one with two subtribes, but these tribes turned out to be non-monophyletic and have been abandoned. A different classification was proposed by Hand and Kirsch in 2003. More recently, Petr Benda and Peter Vallo (2009) proposed a separate tribe, Triaenopini, for the genera Triaenops, Paratriaenops, and possibly Cloeotis.
The genera included in Hipposideridae are (species counts only include living species):
- Archerops (Miocene of Australia)
- Anthops (one species; Solomon Islands and Bougainville Island)
- Asellia (two species; Africa and southwestern Asia; Miocene fossils from Europe)
- Aselliscus (two species; southeastern Asia and Melanesia)
- Brachipposideros (Oligocene to Miocene of Europe and Miocene of Australia; sometimes included in Hipposideros)
- Brevipalatus (Miocene of Australia)
- Cloeotis (one species; Africa)
- Coelops (at least two species; southeastern Asia; Miocene fossils from Africa)
- Hipposideros (more than sixty species; Africa, southern Eurasia, and Australasia; oldest fossils from the Eocene of Europe; includes Pseudorhinolophus, sometimes considered a separate genus)
- Miophyllorhina (Miocene of Australia)
- Palaeophyllophora (Eocene to Miocene of Europe)
- Paracoelops (one species; Vietnam)
- Paraphyllophora (Eocene or Oligocene to Miocene of Europe)
- Paratriaenops (previously included in Triaenops; three species; Madagascar and Seychelles)
- Rhinonicteris (one species; Australia; earliest fossils from the Miocene)
- Riversleighia (Miocene of Australia)
- Triaenops (four species; Africa and southwestern Asia)
- Vaylatsia (Oligocene of Europe)
- Xenorhinos (Miocene of Australia)
List of species
- Family Hipposideridae — Old World leaf-nosed bats
- Genus Anthops
- Flower-faced bat, Anthops ornatus
- Genus Asellia — trident leaf-nosed bats
- Genus Aselliscus — Tate's trident-nosed bats
- Genus Cloeotis
- Percival's trident bat, Cloeotis percivali
- Genus Coelops — tailless leaf-nosed bats
- Genus Hipposideros — roundleaf bats
- Aba roundleaf bat, Hipposideros abae
- Great roundleaf bat, Hipposideros armiger
- Dusky roundleaf bat, Hipposideros ater
- Benito roundleaf bat, Hipposideros beatus
- Boeadi’s roundleaf bat, Hipposideros boeadii
- Bicolored roundleaf bat, Hipposideros bicolor
- Short-headed roundleaf bat, Hipposideros breviceps
- Sundevall's roundleaf bat, Hipposideros caffer
- Spurred roundleaf bat, Hipposideros calcaratus
- Greater roundleaf bat, Hipposideros camerunensis
- Fawn leaf-nosed bat or fawn roundleaf bat, Hipposideros cervinus
- Ashy roundleaf bat, Hipposideros cineraceus
- Commerson's leaf-nosed bat or Commerson's roundleaf bat, Hipposideros commersoni
- Large Mindanao roundleaf bat, Hipposideros coronatus
- Telefomin roundleaf bat, Hipposideros corynophyllus
- Cox's roundleaf bat, Hipposideros coxi
- Timor roundleaf bat, Hipposideros crumeniferus
- Short-tailed roundleaf bat, Hipposideros curtus
- Cyclops roundleaf bat, Hipposideros cyclops
- Makira roundleaf bat, Hipposideros demissus
- Diadem roundleaf bat, Hipposideros diadema
- Fierce roundleaf bat, Hipposideros dinops
- Borneo roundleaf bat, Hipposideros doriae
- Khajuria's leaf-nosed bat, Hipposideros durgadasi
- Dayak roundleaf bat, Hipposideros dyacorum
- Hill's roundleaf bat, Hipposideros edwardshilli
- Hipposideros fasensis
- Sooty roundleaf bat, Hipposideros fuliginosus
- Fulvus roundleaf bat, Hipposideros fulvus
- Cantor's roundleaf bat, Hipposideros galeritus
- Giant roundleaf bat, Hipposideros gigas
- Grand roundleaf bat, Hipposideros grandis
- Thailand roundleaf bat, Hipposideros halophyllus
- Kolar leaf-nosed bat, Hipposideros hypophyllus
- Crested roundleaf bat, Hipposideros inexpectatus
- Arnhem leaf-nosed bat, Hipposideros inornatus
- Jones's roundleaf bat, Hipposideros jonesi
- Phou Khao Khouay leaf-nosed bat, Hipposideros khaokhouayensis
- Khasian leaf-nosed bat, Hipposideros khasiana
- Lamotte's roundleaf bat, Hipposideros lamottei
- Indian roundleaf bat, Hipposideros lankadiva
- Intermediate roundleaf bat, Hipposideros larvatus
- Large Asian roundleaf bat, Hipposideros lekaguli
- Shield-faced roundleaf bat, Hipposideros lylei
- Big-eared roundleaf bat, Hipposideros macrobullatus
- Maduran leaf-nosed bat, Hipposideros madurae
- Maggie Taylor's roundleaf bat, Hipposideros maggietaylorae
- Aellen's roundleaf bat, Hipposideros marisae
- Ethiopian large-eared roundleaf bat, Hipposideros megalotis
- Fly River roundleaf bat, Hipposideros muscinus
- Malayan roundleaf bat, Hipposideros nequam
- Philippine forest roundleaf bat, Hipposideros obscurus
- Orbiculus leaf-nosed bat, Hipposideros orbiculus
- Biak roundleaf bat, Hipposideros papua
- Hipposideros parnabyi
- Peleng leaf-nosed bat, Hipposideros pelingensis
- Pomona roundleaf bat, Hipposideros pomona
- Pratt's roundleaf bat, Hipposideros pratti
- Philippine pygmy roundleaf bat, Hipposideros pygmaeus
- Ridley's leaf-nosed bat, Hipposideros ridleyi
- Laotian leaf-nosed bat, Hipposideros rotalis
- Noack's roundleaf bat, Hipposideros ruber
- Hipposideros scutinares
- Semon's roundleaf bat, Hipposideros semoni
- Sorensen's leaf-nosed bat, Hipposideros sorenseni
- Schneider's leaf-nosed bat, Hipposideros speoris
- Northern leaf-nosed bat or narrow-eared roundleaf bat, Hipposideros stenotis
- Sumba roundleaf bat or Sumban leaf-nosed bat, Hipposideros sumbae
- Hipposideros tephrus
- Saõ Tomé leaf-nosed bat, Hipposideros thomensis
- Lesser great leaf-nosed bat, Hipposideros turpis
- Striped leaf-nosed bat, Hipposideros vittatus
- Wollaston's roundleaf bat, Hipposideros wollastoni
- Genus Paracoelops
- Vietnam leaf-nosed bat, Paracoelops megalotis
- Genus Rhinonicteris
- Orange leaf-nosed bat, Rhinonicteris aurantia
- Genus Paratriaenops
- Genus Triaenops — triple leaf-nosed bats
- Genus Anthops
- Simmons, 2005, p. 365
- Hutcheon and Kirsch, 2006
- Simmons, 2005, pp. 365–379
- McKenna and Bell, 1997, p. 306
- McKenna and Bell, 1997, pp. 306–307
- Hand and Kirsch, 2003, table 3
- Benda and Vallo, 2009, p. 33
- Simmons, 2005, pp. 365–379; McKenna and Bell, 1997, pp. 306–307; other sources cited for specific genera
- Hand and Kirsch, 2003
- Hand and Archer, 2005
- Archer et al., 2006, p. 7
- Benda and Vallo, 2009, p. 34
- Ziegler, 2000, p. 652; Hand and Kirsch, 2003, table 3; cf. McKenna and Bell, 1997, p. 305 (excluded from Rhinonycterinae)
- Archer, M., Arena, D.A., Bassarova, M., Beck, R.M.D., Black, K., Boles, W.E., Brewer, P., Cooke, B.N., Crosby, K., Gillespie, A., Godthelp, H., Hand, S.J., Kear, B.P., Louys, J., Morrell, A., Muirhead, J., Roberts, K.K., Scanlon, J.D., Travouillon, K.J. and Wroe, S. 2006. Current status of species-level representation in faunas from selected fossil localities in the Riversleigh World Heritage Area, northwestern Queensland. Alcheringa Special Issue 1:1-17. ISBN 0-9757894-5-7
- Benda, P. and Vallo, P. 2009. Taxonomic revision of the genus Triaenops (Chiroptera: Hipposideridae) with description of a new species from southern Arabia and definitions of a new genus and tribe. Folia Zoologica 58(Monograph 1):1–45.
- Hand, S.J. and Archer, M. 2005. A new hipposiderid genus (Microchiroptera) from an early Miocene bat community in Australia. Palaeontology 48(2):371–383.
- Hand, S.J. and Kirsch, J.A.W. 2003. Archerops, a new annectent hipposiderid genus (Mammalia: Microchiroptera) from the Australian Miocene. Journal of Paleontology 77(6):1139–1151.
- 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.
- 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
- 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
- Ziegler, R. 2000. The bats (Chiroptera, Mammalia) from the Late Oligocene fissure fillings Herrlingen 8 and Herrlingen 9 near Ulm (Baden-Württemberg). Senckenbergiana Lethaea 80(2):647–683.
- This name technically has priority over Hipposiderinae Lydekker, 1891, and some have consequently used "Rhinonycteridae" or "Rhinonycterinae" for this (sub)family; however, Hipposideridae/inae has been in common use since 1907 and is currently retained pending action by the International Commission on Zoological Nomenclature.
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