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Vespertilionidae is a family of microbats, of the order Chiroptera, flying, insect-eating mammals variously described as the common, vesper, or simple nosed bats. The vespertilionid family is the most diverse and widely distributed of bat families, specialised in many forms to occupy a range of habitats and ecological circumstances, and it is frequently observed or the subject of research. The facial features of the species are often simple, as they mainly rely on vocally emitted echolocation. The tails of the species are enclosed by the lower flight membranes between the legs. Over 300 species are distributed all over the world, on every continent except
Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest cont ...
. It owes its name to the genus '' Vespertilio'', which takes its name from a word for bat, ', derived from the
Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through ...
term ' meaning 'evening'; they are termed "evening bats" and were once referred to as "evening birds". (The term "evening bat" also often refers more specifically to one of the species, '' Nycticeius humeralis''.)


Evolution

They are allied to the suborder Microchiroptera, the families of microbats separated from the flying foxes and fruit bats of the megabat group Megachiroptera. The treatments of bat taxonomy have also included a placement amongst the Vespertilioniformes, or
Yangochiroptera Yangochiroptera, or Vespertilioniformes, is a suborder of Chiroptera that includes most of the microbat families, except the Rhinopomatidae, Rhinolophidae, Hipposideridae, and Megadermatidae. These other families, plus the megabats, are seen as ...
, as suborder Vespertilionoidea. Molecular data indicate the Vespertilionidae diverged from the Molossidae in the early Eocene period. The family is thought to have originated somewhere in
Laurasia Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pa ...
, possibly North America. A recently extinct species, '' Synemporion keana'', is known from the
Holocene The Holocene ( ) is the current geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene togeth ...
of
Hawaii Hawaii ( ; haw, Hawaii or ) is a state in the Western United States, located in the Pacific Ocean about from the U.S. mainland. It is the only U.S. state outside North America, the only state that is an archipelago, and the only stat ...
.Discovery of Extinct Bat Doubles Diversity of Native Hawaiian Land Mammals
at the American Museum of Natural History; published March 21, 2016; retrieved June 20, 2016


Characteristics

All species are carnivorous and most are insectivores, exceptions are bats of genera '' Myotis'' and '' Pizonyx'' that catch
fish Fish are Aquatic animal, aquatic, craniate, gill-bearing animals that lack Limb (anatomy), limbs with Digit (anatomy), digits. Included in this definition are the living hagfish, lampreys, and Chondrichthyes, cartilaginous and bony fish as we ...
and the larger '' Nyctalus'' species known to capture small
passerine A passerine () is any bird of the order Passeriformes (; from Latin 'sparrow' and '-shaped'), which includes more than half of all bird species. Sometimes known as perching birds, passerines are distinguished from other orders of birds by th ...
birds in flight. The
dentition Dentition pertains to the development of teeth and their arrangement in the mouth. In particular, it is the characteristic arrangement, kind, and number of teeth in a given species at a given age. That is, the number, type, and morpho-physiolo ...
of the family varies between species; the dental formula of the family is: They rely mainly on echolocation to navigate and obtain food, but they lack the elaborate nose appendages of microbats that focus nasal emitted
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 ...
. The ultrasound signal is usually produced orally, and many species have large external ears to capture and reflect sound, enabling them to discriminate and extract information. The vespertilionids employ a range of flight techniques. The wing surface is extended to the lower limbs, and the tails of this family are enclosed in an interfemoral membrane. Some are relatively slow-flying genera, such as ''
Pipistrellus ''Pipistrellus'' is a genus of bats in the family Vespertilionidae and subfamily Vespertilioninae. The name of the genus is derived from the Italian word , meaning "bat" (from Latin "bird of evening, bat"). The size of the genus has been con ...
'', that manipulate the configuration of their broader wing shape and may give a fluttery appearance as they forage and glean. Others are specialised as long-winged genera, such as ''
Lasiurus ''Lasiurus'' is a genus of bats in the family Vespertilionidae. Its members are known as hairy-tailed bats or red bats. Phylogeny The following is the relationship of the three genera formerly included within ''Lasiurus'', based on an analysis ...
'' and '' Nyctalus'', that use rapid pursuit to capture insects. The size range of the family is in head and body length; this excludes the tail, which is itself quite long in many species. They are generally brown or grey in colour, often an indiscriminate appearance as a 'little brown bat', although some species have fur that is brightly colored, with reds, oranges, and yellows all being known. The patterns of the superficial appearance include white patches or stripes that may distinguish some species. Most species roost in caves, although some make use of hollow trees, rocky crevices, animal
burrow An Eastern chipmunk at the entrance of its burrow A burrow is a hole or tunnel excavated into the ground by an animal to construct a space suitable for habitation or temporary refuge, or as a byproduct of locomotion. Burrows provide a form of s ...
s, or other forms of shelter. Colony sizes also vary greatly, with some roosting alone, and others in groups up to a million individuals. Species native to temperate latitudes typically hibernate to avoid cooler weather, while a few of the
tropical The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...
species employ
aestivation Aestivation ( la, aestas (summer); also spelled estivation in American English) is a state of animal dormancy, similar to hibernation, although taking place in the summer rather than the winter. Aestivation is characterized by inactivity and a ...
as a method of evading extremes of climate.


Systematics

The four subfamilies of Vespertilionidae separate the presumably related taxa, tribes, and genera of extant and extinct taxa. The subfamilial treatments, based on morphological, geographical, and ecological comparisons have been recombined since the inclusion of the phylogenetic implications of molecular genetics; only the Murininae and Kerivoulinae have not been changed in light of genetic analysis. Subfamilies that were once recognized as valid, such as the Nyctophilinae, are considered dubious, as molecular evidence suggests they are
paraphyletic In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In ...
in their arrangements. Within the concept
Yangochiroptera Yangochiroptera, or Vespertilioniformes, is a suborder of Chiroptera that includes most of the microbat families, except the Rhinopomatidae, Rhinolophidae, Hipposideridae, and Megadermatidae. These other families, plus the megabats, are seen as ...
, an acknowledged cladistic treatment, the closest relatives to the family are the free-tailed bats of family Molossidae. The monotypic genus ''Tomopeas'', represented by the blunt-eared bat ('' Tomopeas ravus''), is acknowledged as the potentially closest link between the Vespertilionidae and Molossidae, as it is the most basal member of the Molossidae and has intermediate characteristics of both families.


Classification

The grouping of these subfamilies is the classification published by the American Society of Mammalogists. Other authorities raise three subfamilies more: Antrozoinae (which is here the separate family of pallid bats),
Tomopeatinae The blunt-eared bat or Peruvian crevice-dwelling bat (''Tomopeas ravus'') is a species of bat in the family Molossidae. It is monotypic within the genus ''Tomopeas'' and subfamily Tomopeatinae. It is Endemism, endemic to Peru, where it is conside ...
(now regarded as a subfamily of the
free-tailed bat The Molossidae, or free-tailed bats, are a family of bats within the order Chiroptera. The Molossidae is the fourth-largest family of bats, containing about 110 species as of 2012. They are generally quite robust, and consist of many strong-flyi ...
s), and Nyctophilinae (here included in Vespertilioninae). Four subfamilies are recognized by '' Mammal Species of the World'' (2005), the highly diverse Vespertilioninae are also separated as tribes. Newer or resurrected genera are noted. The genus '' Cistugo'' is no longer included following its move to the separate family Cistugidae. Miniopterinae is additionally no longer recognized as a subfamily, as it was elevated to family status. A 2021 study attempted to resolve the systematic relationships among the pipistrelle-like bats of sub-Saharan Africa and Madagascar, with systematic inferences based on genetic and morphological analyses of more than 400 individuals across all named genera and the majority of described African pipistrelle-like bat species, with a focus on previously unstudied samples of East African bats. The study proposed a revision of the pipistrelle-like bats in East Africa and described multiple new genera and species. Family Vespertilionidae * subfamily Kerivoulinae ::* genus ''
Kerivoula ''Kerivoula'' is a genus of vesper bat Vespertilionidae is a family of microbats, of the order Chiroptera, flying, insect-eating mammals variously described as the common, vesper, or simple nosed bats. The vespertilionid family is the most div ...
'' – painted bats ::* genus '' Phoniscus'' *subfamily Myotinae ::* genus ''
Eudiscopus The disk-footed bat (''Eudiscopus denticulus'') is a species of vesper bat in the Vespertilionidae family found in Laos and Myanmar. The disc-footed bat was recorded in the northeastern Indian state's (Meghalaya), Lailad area near the Nongkhylle ...
'' ::*genus '' Myotis'' – mouse-eared bats ::* genus ''
Submyotodon ''Submyotodon'' is a genus of vespertilionid bats, published as a new taxon in 2003 to describe a Miocene fossil species. Extant species and subspecies previously included in ''Myotis'' were later transferred to this genus. Species in this genus ...
'' – broad-muzzled bats * subfamily Murininae ::* genus ''
Harpiocephalus The lesser hairy-winged bat (''Harpiocephalus harpia'') is a species of vesper bat in the family Vespertilionidae. It can be found in India, Indonesia, the Philippines, and Taiwan Taiwan, officially the Republic of China (ROC), is a coun ...
'' – hairy-winged bats ::* genus ''
Harpiola ''Harpiola'' is a genus (or possibly subgenus) of vesper bats within the subfamily Murininae. It contains Peters's tube-nosed bat (''Harpiola grisea'') and the Formosan golden tube-nosed bat The Formosan golden tube-nosed bat (''Harpiola iso ...
'' ::* genus ''
Murina ''Murina'' is a genus of vesper bats. They are found throughout temperate and tropical regions of Asia. Taxonomy Species list based on American Society of Mammalogists and ITIS The Integrated Taxonomic Information System (ITIS) is an Ame ...
'' – tube-nosed insectivorous bats * subfamily Vespertilioninae ** tribe
Antrozoini Antrozoini is a tribe of bats in the subfamily Vespertilioninae of the family Vespertilionidae. It contains the pallid bat (''Antrozous pallidus''), Van Gelder's bat (''Bauerus dubiaquercus''), the genus '' Rhogeessa'', and the fossil '' Anzany ...
***genus '' Antrozous'' ***genus '' Bauerus'' ***genus ''
Rhogeessa ''Rhogeessa'' is a genus of bats within the vesper bats family, Vespertilionidae. Species * Yucatan yellow bat (''R. aeneus'') * Allen's yellow bat (''R. alleni'') * Bickham's little yellow bat (''Rhogeessa bickhami'') * Genoways's yellow bat ( ...
'' **tribe Eptesicini *** genus ''
Arielulus ''Arielulus'' is a genus of vesper bat Vespertilionidae is a family of microbats, of the order Chiroptera, flying, insect-eating mammals variously described as the common, vesper, or simple nosed bats. The vespertilionid family is the most div ...
'' *** genus '' Eptesicus'' – house bats *** genus ''
Glauconycteris ''Glauconycteris'' is a genus of vespertilionid bats found in Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles ...
'' – butterfly bats *** genus ''
Hesperoptenus Hesperoptenus is a genus of bats within the Vespertilionidae or vesper bat family. The species within this genus are: * Blanford's bat (''Hesperoptenus blanfordi'') * False serotine bat (''Hesperoptenus doriae'') * Gaskell's false serotine (''H ...
'' – false serotine bats ***genus ''
Histiotus ''Histiotus'' (meaning "sail ears") is a genus of South American vesper bats with species that include: * Strange big-eared brown bat, ''Histiotus alienus'' * Cadena-García's big-eared brown bat, ''Histiotus cadenai'' * Colombian big-eared ...
'' – big-eared brown bats ***genus '' Ia'' ***genus ''
Lasionycteris The silver-haired bat (''Lasionycteris noctivagans'') is a solitary migratory species of vesper bat in the family Vespertilionidae and the only member of the genus ''Lasionycteris''. Etymology The species name translates as night-wandering, r ...
'' ***genus '' Scoteanax'' – greater broad-nosed bats ***genus ''
Scotomanes The harlequin bat (''Scotomanes ornatus'') is a species of 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 tru ...
'' ***genus '' Scotorepens'' – lesser broad-nosed bats ***genus ''
Thainycteris ''Thainycteris'' is a genus of vesper bat with two species, one found in Indochina and the other found in Taiwan. Its members were previously classified within ''Arielulus'', but more recent studies support ''Thainycteris'' as being a distinct ge ...
'' ** tribe '' incertae sedis'' ***genus '' Rhyneptesicus'' **tribe
Lasiurini Lasiurini is a tribe of bats in the family Vespertilionidae. It contains three genera of bats found in the Americas. All three genera were previously considered one genus, ''Lasiurus'', but have since been split from one another. However, the val ...
*** genus ''
Aeorestes ''Aeorestes'' is a genus or subgenus of vesper bat commonly known as the hoary bats. As a genus, it includes species that were formerly included in the genus ''Lasiurus''. Taxonomy The genus was initially described by Austrian zoologist Leopol ...
'' – hoary bats *** genus ''
Dasypterus ''Dasypterus'' is a genus of or subgenus of vesper bat. As a genus, it includes species that were formerly in the genus ''Lasiurus''. Collectively, members of ''Dasypterus'' are referred to as the yellow bats. Taxonomy Based on genetic diver ...
'' – yellow bats *** genus ''
Lasiurus ''Lasiurus'' is a genus of bats in the family Vespertilionidae. Its members are known as hairy-tailed bats or red bats. Phylogeny The following is the relationship of the three genera formerly included within ''Lasiurus'', based on an analysis ...
'' – hairy-tailed bats ** tribe Nycticeiini *** genus ''
Nycticeius ''Nycticeius'' is a small genus of bats in the vesper bat family, Vespertilionidae, and the only member of the tribe Nycticeiini. It contains three species, the evening bat (''N. humeralis''), the Cuban evening bat (''N. cubanus'') and ''Nyctice ...
'' – evening bats ** tribe
Perimyotini Perimyotini is a tribe of bats in the family Vespertilionidae. It contains two species of bats found in North America, each in their own monotypic genus. Although this name is already in use by taxonomic authorities, such as the ''Handbook of ...
***genus ''
Parastrellus The canyon bat (''Parastrellus hesperus''), also known as the western pipistrelle, is a species of vesper bat. It is found in Mexico and in the western United States. The species has historically been placed in the genus ''Pipistrellus ''Pip ...
'' *** genus ''
Perimyotis The tricolored bat (''Perimyotis subflavus'') is a species of microbat native to eastern North America. Formerly known as the eastern pipistrelle, based on the incorrect belief that it was closely related to European ''Pipistrellus'' species, t ...
''Solari, S. 2018. Perimyotis subflavus. The IUCN Red List of Threatened Species 2018* e.T17366A22123514. https://doi.org/10.2305/IUCN.UK.2018-2.RLTS.T17366A22123514.en. Downloaded on 05 March 2019. ** tribe Pipistrellini *** genus ''
Glischropus ''Glischropus'' is a genus of bats within the vesper bat family, Vespertilionidae. Species within this genus are: * '' Glischropus aquilus'' (Görföl, Wiantoro, Kingston, Bates & Huang, 2015) — dark thick-thumbed bat * ''Glischropus bucephal ...
'' – thick-thumbed bats *** genus '' Nyctalus'' – noctule bats *** genus ''
Pipistrellus ''Pipistrellus'' is a genus of bats in the family Vespertilionidae and subfamily Vespertilioninae. The name of the genus is derived from the Italian word , meaning "bat" (from Latin "bird of evening, bat"). The size of the genus has been con ...
'' – true pipistrelles *** genus ''
Scotoecus ''Scotoecus'' is a genus of bats in the family Vespertilionidae. Species * '' Scotoecus albigula'' - white-bellied lesser house bat * '' Scotoecus albofuscus'' - light-winged lesser house bat * '' Scotoecus hindei'' - Hinde's lesser house bat * ...
'' – house bats ***genus ''
Scotozous Dormer's bat or Dormer's pipistrelle (''Scotozous dormeri'') is a species of vesper bat. It is the only species in its genus. It is found in Bangladesh, Bhutan, India, and Pakistan. Its natural habitats are subtropical and tropical dry forests ...
'' ***genus '' Vansonia'' ** tribe
Plecotini Plecotini is a tribe of bats in the family Vespertilionidae. It contains several genera found throughout the Northern Hemisphere, in Eurasia, North Africa, and North America. Several genera in this tribe are known as big-eared bats or long-eared ...
*** genus ''
Barbastella ''Barbastella'' is a genus of vespertilionid bats. There are seven extant species in this genus and one only known from fossil remains. Species The genus consists of the following species: * '' Barbastella barbastellus'' – western barbastell ...
'' – barbastelles or barbastelle bats *** genus '' Corynorhinus'' – American lump-nosed bats *** genus '' Euderma'' *** genus '' Idionycteris'' *** genus ''
Otonycteris ''Otonycteris'' is a genus of vesper bats. Members of this genus are found in Northern Africa and Central Asia. Until recently, it was thought to be monotypic, but in 2010, the Turkestani long-eared bat was distinguished from the desert long-e ...
'' *** genus ''
Plecotus ''Plecotus'' is a genus of vesper bat, commonly called long-eared bats. They are found throughout Eurasia and northern Africa. Many species in the genus have only been described and recognized in recent years. Species Genus ''Plecotus'' � ...
'' – lump-nosed bats ** tribe Scotophilini ***genus ''
Scotophilus ''Scotophilus'' is a genus of vesper bat, vespertilionid bats commonly called yellow bats. They are found in southern Asia and Africa. They are the only members of the Tribe (biology), tribe Scotophilini. Species * East African yellow bat, ' ...
'' – Old World yellow bats **tribe
Vespertilionini Vespertilionini is a tribe of bats in the family Vespertilionidae. The largest of the tribes in Vespertilioninae, it contains many genera found throughout the Old World and Australasia. Species Species in the tribe include: * Genus '' Afrony ...
*** genus ''
Afronycteris ''Afronycteris'' is a genus of vesper bat containing two species, both of which are found in sub-Saharan Africa Sub-Saharan Africa is, geographically, the area and regions of the continent of Africa that lies south of the Sahara. These in ...
'' *** genus '' Cassistrellus'' – helmeted bats ***genus '' Chalinolobus'' – wattled bats *** genus ''
Falsistrellus ''Falsistrellus'' is a genus of vespertilionid family of bats, small predatory flying mammals. They are known from Australia. The poorly researched species have been variously placed by authors, and revised again by studies of their distinct char ...
'' – false pipistrelles *** genus '' Hypsugo'' – Asian pipistrelles *** genus '' Laephotis'' – long-eared bats *** genus '' Mimetillus'' – mimic bats ***genus '' Mirostrellus'' *** genus ''
Neoromicia ''Neoromicia'' is a genus of vesper bat in the family Vespertilionidae Vespertilionidae is a family of microbats, of the order Chiroptera, flying, insect-eating mammals variously described as the common, vesper, or simple nosed bats. The vespe ...
'' ***genus ''
Nycticeinops ''Nycticeinops'' is a genus of vesper bat in the family Vespertilionidae. It contains the following species: * Bellier's serotine, ''Nycticeinops bellieri'' * Broad-headed serotine, ''Nycticeinops crassulus'' * Eisentraut's serotine, ''Nyc ...
'' ***genus ''
Nyctophilus ''Nyctophilus'' is a genus of the vespertilionids or vesper bats. They are often termed Australian big-eared bats or long-eared bats, as the length of their ears often greatly exceeds that of the head. This genus occurs in the New Guinean- Aus ...
'' – New Guinean and Australian big-eared bats ***genus ''
Pharotis The New Guinea big-eared bat or Papuan big-eared bat, (''Pharotis imogene''), is a vesper bat endemic to Papua New Guinea. It is listed as a critically endangered species due to ongoing habitat loss. It is the only known member of the genus ''P ...
'' *** genus '' Philetor'' *** genus ''
Pseudoromicia ''Pseudoromicia'' is a genus of vesper bat in the family Vespertilionidae. All species in this genus are native to sub-Saharan Africa. It contains the following species: * Dark-brown serotine, ''Pseudoromicia brunnea'' * Isabelline serotine, ...
'' ***genus ''
Tylonycteris The bamboo bats are genus of vesper bats Vespertilionidae is a family of microbats, of the order Chiroptera, flying, insect-eating mammals variously described as the common, vesper, or simple nosed bats. The vespertilionid family is the most ...
'' – bamboo bats *** genus ''
Vespadelus ''Vespadelus'' is a genus of Australian bats in the family Vespertilionidae. The genus name was first published by Ellis Le Geyt Troughton and Tom Iredale in 1934, but without an adequate description; the type was cited as ''Scotophilus pumilus ...
'' *** genus '' Vespertilio'' – frosted bats


References


Further reading

* Corbet, GB, Hill JE. 1992. The mammals of the Indomalayan region: a systematic review. Oxford University Press, Oxford. * Karim, C., A.A. Tuen and M.T. Abdullah. 2004. Mammals. Sarawak Museum Journal Special Issue No. 6. 80: 221–234. * Wilson DE, Reeder DM. 2005. Mammal species of the world. Smithsonian Institution Press, Washington DC. {{DEFAULTSORT:Vesper Bat * Extant Eocene first appearances Taxa named by John Edward Gray Bat families