Vespertilionid
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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. It owes its name to the genus '' Vespertilio'', which takes its name from a word for bat, ', derived from the Latin 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 The evening bat (''Nycticeius humeralis'') is a species of bat in the vesper bat family that is native to North America. Hunting at night, they eat beetles, moths, and other flying insects. Description The evening bat is a small bat weighin ...
''.)


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, as suborder Vespertilionoidea. Molecular data indicate the Vespertilionidae diverged from the Molossidae in the
early Eocene In the geologic timescale the Ypresian is the oldest age or lowest stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by the Eocene Lutetian Age. The Ypresian i ...
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 Pan ...
, possibly North America. A recently extinct species, '' Synemporion keana'', is known from the Holocene of Hawaii.Discovery of Extinct Bat Doubles Diversity of Native Hawaiian Land Mammals
at the
American Museum of Natural History The American Museum of Natural History (abbreviated as AMNH) is a natural history museum on the Upper West Side of Manhattan in New York City. In Theodore Roosevelt Park, across the street from Central Park, the museum complex comprises 26 inter ...
; published March 21, 2016; retrieved June 20, 2016


Characteristics

All species are carnivorous and most are
insectivore A robber fly eating a hoverfly An insectivore is a carnivorous animal or plant that eats insects. An alternative term is entomophage, which can also refer to the human practice of eating insects. The first vertebrate insectivores wer ...
s, exceptions are bats of genera '' Myotis'' and '' Pizonyx'' that catch fish and the larger '' Nyctalus'' species known to capture small passerine 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. 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'', 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'' 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 sh ...
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 Hibernation is a state of minimal activity and metabolic depression undergone by some animal species. Hibernation is a seasonal heterothermy characterized by low body-temperature, slow breathing and heart-rate, and low metabolic rate. It most ...
to avoid cooler weather, while a few of the tropical 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 ...
as a method of evading extremes of climate.


Systematics

The four
subfamilies In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...
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 (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
in their arrangements. Within the concept Yangochiroptera, 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 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 endemic to Peru, where it is considered criti ...
''), is acknowledged as the potentially closest link between the Vespertilionidae and Molossidae, as it is the most
basal Basal or basilar is a term meaning ''base'', ''bottom'', or ''minimum''. Science * Basal (anatomy), an anatomical term of location for features associated with the base of an organism or structure * Basal (medicine), a minimal level that is nec ...
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 The American Society of Mammalogists (ASM) was founded in 1919. Its primary purpose is to encourage the study of mammals, and professions studying them. There are over 4,500 members of this society, and they are primarily professional scientists ...
. 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 endemic to Peru, where it is considered criti ...
(now regarded as a subfamily of the free-tailed bats), 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'' – painted bats ::* genus ''
Phoniscus ''Phoniscus'' 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 ...
'' *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 cou ...
'' – hairy-winged bats ::* genus ''
Harpiola ''Harpiola'' is a genus (or possibly subgenus) of vesper bats within the subfamily Murininae The Murininae are a subfamily of bats in the family Vespertilionidae. They include the tube-nosed bats and hairy-winged bats in the genera ''Murin ...
'' ::* 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 Am ...
'' – tube-nosed insectivorous bats * subfamily
Vespertilioninae The Vespertilioninae are a subfamily of vesper bats from the family Vespertilionidae. Classification Subfamily Vespertilioninae *Tribe Antrozoini **Genus '' Antrozous'' *** Pallid bat, ''Antrozous pallidus'' **Genus ''Bauerus'' ***Van Gelder ...
** tribe
Antrozoini Antrozoini is a tribe (taxonomy), 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 f ...
***genus '' Antrozous'' ***genus ''
Bauerus Van Gelder's bat or Van Gelder's big-eared bat (''Bauerus dubiaquercus'') is a species of vesper bat in the family Vespertilionidae. It is found in Belize, Costa Rica, Honduras, and Mexico. The species is monotypic within its genus. It is part of ...
'' ***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 di ...
'' *** 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 Vespertilionidae, vesper bat family. The species within this genus are: *Blanford's bat (''Hesperoptenus blanfordi'') *False serotine bat (''Hesperoptenus doriae'') *Gaskell's fals ...
'' – 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 b ...
'' – big-eared brown bats ***genus '' Ia'' ***genus '' Lasionycteris'' ***genus '' Scoteanax'' – greater broad-nosed bats ***genus ''
Scotomanes The harlequin bat (''Scotomanes ornatus'') is a species of 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 simp ...
'' ***genus ''
Scotorepens ''Scotorepens'' is a genus of bats within the Vespertilionidae family. Species within this genus are widely distributed across Australia and to the north at Papua New Guinea and Indonesia. Taxonomy The genus was erected by Ellis Troughton in h ...
'' – 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 vali ...
*** 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'' – 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 '' Nycti ...
'' – 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 the ...
***genus '' Parastrellus'' *** 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, th ...
''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 Pipistrellini is a tribe of bats in the family Vespertilionidae. It contains several genera found throughout the Old World and Australasia, including the pipistrelles, noctules and related species. Species Species in the tribe include: * Gen ...
*** 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 bucephalu ...
'' – thick-thumbed bats *** genus '' Nyctalus'' – noctule bats *** genus '' Pipistrellus'' – true pipistrelles *** genus ''
Scotoecus ''Scotoecus'' is a genus of bats 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 vespertilion ...
'' – house bats ***genus '' Scotozous'' ***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 b ...
*** genus '' Barbastella'' – barbastelles or barbastelle bats *** genus '' Corynorhinus'' – American lump-nosed bats *** genus ''
Euderma The spotted bat (''Euderma maculatum'') is a species of vesper bat and the only species of the genus ''Euderma''. Description The spotted bat was first described by zoologist Joel Asaph Allen from the American Museum of Natural History in 1891. ...
'' *** genus ''
Idionycteris Allen's big-eared bat (''Idionycteris phyllotis'') is a species of vesper bat in the monotypic genus ''Idionycteris''. It occurs in Mexico and in Arizona, California, Nevada, New Mexico, Utah, and ColoradoHayes, M. A., et al. (2009)Allen's big-ea ...
'' *** 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-ea ...
'' *** genus ''
Plecotus ''Plecotus'' is a genus of Vespertilionidae, vesper bat, commonly called long-eared bats. They are found throughout Eurasia and North Africa, northern Africa. Many species in the genus have only been described and recognized in recent years. Spe ...
'' – lump-nosed bats ** tribe Scotophilini ***genus '' Scotophilus'' – 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 '' Afronyct ...
*** genus ''
Afronycteris ''Afronycteris'' is a genus of Vespertilionidae, vesper bat containing two species, both of which are found in sub-Saharan Africa. Its members were previously classified in ''Neoromicia'' before phylogenetic analysis found them to comprise a separ ...
'' *** genus ''
Cassistrellus The helmeted bats are two species of vesper bat in the genus ''Cassistrellus''. They are found in South and Southeast Asia. The two species in this genus are: * Surat helmeted bat, ''Cassistrellus dimissus'' * Yok Don helmeted bat, ''Cassistrel ...
'' – 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 ''Mimetillus'' is a genus of Vespertilionidae, vesper bat in the family Vespertilionidae. It contains two species, both known as mimic bats and found in sub-Saharan Africa. There are two species in this genus: * Moloney's mimic bat, ''Mimetillus ...
'' – mimic bats ***genus '' Mirostrellus'' *** genus '' Neoromicia'' ***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, ''Nycti ...
'' ***genus '' Nyctophilus'' – 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 ''Pha ...
'' *** genus '' Philetor'' *** genus '' Pseudoromicia'' ***genus '' Tylonycteris'' – 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