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Vespertilionidae
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{{Short description|Family of microbats}} {{Automatic taxobox | fossil_range = [[Early Eocene]] to recent<ref name=MBF>{{cite book |last=Fenton |first=M. B. |year=2001 |title= Bats|publisher= Checkmark Books|location=New York|pages= 5|isbn= 0-8160-4358-2}}</ref> | image = ComputerHotline - Chiroptera sp. (by) (4).jpg | image_caption = [[Greater mouse-eared bat]] (''Myotis myotis'') | image_alt = The image depicts a bat hanging from a cave wall. | taxon = Vespertilionidae | authority = [[John Edward Gray|Gray]], 1821 | type_genus = ''[[Vespertilio]]'' | type_genus_authority = [[Carl Linnaeus|Linnaeus]], 1758 | display_parents = 3 | subdivision_ranks = Subfamilies | subdivision = *[[Vespertilioninae]] *[[Murininae]] *[[Myotinae]] *[[Kerivoulinae]] }} [[File:Parti-coloured bat (Vespertilio murinus) Slovakia.jpg|thumb|Parti-coloured bat<br>''Vespertilio murinus'', Slovakia]] '''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, ''{{lang|la|[[:wikt:vespertilio|vespertilio]]}}'', derived from the [[Latin]] term ''{{lang|la|[[:wikt:vesper|vesper]]}}'' 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, ''[[evening bat|Nycticeius humeralis]]''.) ==Evolution== They are allied to the suborder [[Microchiroptera]], the families of microbats separated from the [[Pteropus|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]] period.<ref name=Miller2007>{{cite journal | author = Miller-Butterworth, C. M., Murphy, W. J., O'Brien, S. J., Jacobs, D. S., Springer, M. S. & Teeling, E. C. | year = 2007 | title = A family matter: conclusive resolution of the taxonomic position of the long-fingered bats, ''Miniopterus'' | journal = [[Molecular Biology and Evolution]] | volume = 24 | issue = 7 | pages = 1553β1561 | doi = 10.1093/molbev/msm076 | pmid=17449895| doi-access = free }}</ref> The family is thought to have originated somewhere in [[Laurasia]], possibly North America.<ref name=Teeling2005>{{cite journal | author = Teeling, E. C., Springer, M. S., Madsen, O., Bates, P., O'Brien, S. J. & Murphy, W. J. | year = 2005 | title = A molecular phylogeny for bats illuminates biogeography and the fossil record | journal = [[Science (journal)|Science]] | volume = 307 |issue=5709 | pages = 580β584 | doi = 10.1126/science.1105113 | pmid=15681385| bibcode = 2005Sci...307..580T | s2cid = 25912333 }}</ref> A recently extinct species, ''[[Synemporion keana]]'', is known from the [[Holocene]] of [[Hawaii]].<ref name=AMNH>[http://www.amnh.org/content/download/132583/2210546/file/Hawaiian%20bat.pdf 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</ref> ==Characteristics== All species are carnivorous and most are [[insectivore]]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]] of the family varies between species; the dental formula of the family is: {{dentition2|1β2.1.1β3.3|2β3.1.2β3.3}} They rely mainly on [[animal echolocation|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]].<ref name="Richards2012" /> 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 {{convert|3|to|13|cm|in|abbr=on}} 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.<ref name=EoM>{{cite book|editor= Macdonald, D.|year= 1984|title= The Encyclopedia of Mammals|publisher= Facts on File|location= New York|pages= [https://archive.org/details/encyclopediaofma00mals_0/page/807 807]|isbn= 0-87196-871-1|url-access= registration|url= https://archive.org/details/encyclopediaofma00mals_0/page/807}}</ref> Most species roost in caves, although some make use of [[hollow tree]]s, rocky crevices, animal [[burrow]]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 [[hibernation|hibernate]] to avoid cooler weather, while a few of the [[Tropics|tropical]] species employ [[aestivation]] as a method of evading extremes of climate.<ref name=EoM/> ==Systematics== {{cladogram|style=width:250px;font-size:85%;line-height:85%|caption=Subfamily relationships of Vespertilionidae<ref name="Hoofer 2003">{{cite journal| last1=Hoofer| first1= S. R.| last2= Bussche| first2= R. A. V. D.| date= 2003| title= Molecular phylogenetics of the chiropteran family Vespertilionidae| journal= Acta Chiropterologica| volume= 5| issue=1| pages= 1β63| doi= 10.3161/001.005.s101| doi-access= free| hdl= 11244/44678| hdl-access= free}}</ref> |1={{clade |label1='''Vespertilionidae''' |1={{clade |1=[[Vespertilioninae]] |2={{clade |1={{clade |1=[[Myotinae]] |2={{clade |1=[[Kerivoulinae]] |2=[[Murininae]] }} }} }} }} }} }} 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.<ref name="Hoofer 2003"/> Subfamilies that were once recognized as valid, such as the [[Nyctophilinae]], are considered dubious, as molecular evidence suggests they are [[paraphyletic]] in their arrangements.<ref name="Hoofer 2003"/> Within the concept [[Yangochiroptera]], an acknowledged cladistic treatment, the closest relatives to the family are the free-tailed bats of family [[Molossidae]].<ref name="Hoofer 2003"/> The monotypic genus ''Tomopeas'', represented by the blunt-eared bat (''[[Tomopeas ravum]]''), is acknowledged as the potentially closest link between the Vespertilionidae and Molossidae, as it is the most [[basal (phylogenetics)|basal]] member of the Molossidae and has intermediate characteristics of both families.<ref name="Sudman 1994">{{cite journal| doi=10.2307/1382555| jstor=1382555| title=Familial Affinity of Tomopeas ravus (Chiroptera) Based on Protein Electrophoretic and Cytochrome b Sequence Data| journal=Journal of Mammalogy| volume=75| issue=2| pages=365| year=1994| last1=Sudman| first1=P. D| last2=Barkley| first2=L. J| last3=Hafner| first3=M. S}}</ref> ==Classification== {{further|List of kerivoulines|List of myotines|List of murinines|List of vespertilionines}} The grouping of these subfamilies is the classification published by the [[American Society of Mammalogists]].<ref>{{Citation|last=Mammal Diversity Database|title=Mammal Diversity Database|date=2021-08-10|doi=10.5281/zenodo.5175993 |url=https://zenodo.org/record/5175993|access-date=2021-09-17}}</ref> Other authorities raise three subfamilies more: Antrozoinae (which is here the separate family of [[pallid bat]]s), [[Blunt-eared Bat|Tomopeatinae]] (now regarded as a subfamily of the [[free-tailed bat]]s), and Nyctophilinae (here included in Vespertilioninae). Four subfamilies are recognized by ''[[Mammal Species of the World]]'' (2005),<ref name="MSW3" /> 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]].<ref>{{cite journal|last1=Lack|first1=J. B.|last2=Roehrs|first2=Z. P.|last3=Stanley|first3=C. E. Jr.|last4=Ruedi|first4=M.|last5=Van Den Bussche|first5=R. A.|year=2010|title=Molecular phylogenetics of Myotis indicate familial-level divergence for the genus Cistugo (Chiroptera)|journal=Journal of Mammalogy|volume=91|issue=4|pages=976β992|doi=10.1644/09-mamm-a-192.1|doi-access=free}}</ref><ref>{{Catalogue of Life | Vespertilionidae}}</ref> Miniopterinae is additionally no longer recognized as a subfamily, as it was elevated to family status.<ref name="Kulemzina 2011">{{cite journal|doi=10.1159/000328834|last1=Kulemzina| first1= A. I.| last2= Nie| first2= W.| last3= Trifonov| first3= V. A.| last4= Staroselec| first4= Y.| last5= Vasenkov| first5= D. A.| last6= Volleth| first6= M.| last7= Graphodatsky| first7= A. S.| date=2011| title= Comparative chromosome painting of four Siberian Vespertilionidae species with Aselliscus stoliczkanus and Human probes| journal= Cytogenetic and Genome Research| volume= 134| issue=3| pages= 200β205|pmid=21709413|s2cid=8258899}}</ref><ref>{{cite journal |last1=Burgin |first1=Connor J. |last2=Colella |first2=Jocelyn P. |last3= Kahn |first3=Philip L. |last4=Upham |first4=Nathan S. |date=February 1, 2018 |title=How many species of mammals are there? |journal=[[Journal of Mammalogy]] |volume=99 |issue=1 |pages=1β14 |doi=10.1093/jmammal/gyx147 |doi-access=free }}</ref> 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.<ref name="Monadjem">{{cite journal|author1=Ara Monadjem|author2=Terrence C Demos|author3=Desire L Dalton|author4=Paul W Webala|author5=Simon Musila|author6=Julian C Kerbis Peterhans|author7=Bruce D Patterson|title=A revision of pipistrelle-like bats (Mammalia: Chiroptera: Vespertilionidae) in East Africa with the description of new genera and species|journal=Zoological Journal of the Linnean Society|volume=191|issue=4|year=2021|pages=1114β1146|url=https://doi.org/10.1093/zoolinnean/zlaa087|doi=10.1093/zoolinnean/zlaa087|hdl=2263/84301|hdl-access=free}}</ref> '''Family Vespertilionidae''' * subfamily [[Kerivoulinae]] ::* genus ''[[Kerivoula]]'' β painted bats ::* genus ''[[Phoniscus]]'' *subfamily [[Myotinae]] ::* genus ''[[Disk-footed bat|Eudiscopus]]'' ::*genus ''[[Myotis]]'' β mouse-eared bats ::* genus ''[[Submyotodon]]'' β broad-muzzled bats * subfamily [[Murininae]] ::* genus ''[[Harpiocephalus]]'' β hairy-winged bats ::* genus ''[[Harpiola]]'' ::* genus ''[[Murina]]'' β tube-nosed insectivorous bats * subfamily [[Vespertilioninae]] ** tribe [[Antrozoini]] ***genus ''[[Pallid bat|Antrozous]]'' ***genus ''[[Van Gelder's bat|Bauerus]]'' ***genus ''[[Rhogeessa]]'' **tribe [[Eptesicini]] *** genus ''[[Arielulus]]'' *** genus ''[[Eptesicus]]'' β house bats *** genus ''[[Glauconycteris]]'' β butterfly bats *** genus ''[[Hesperoptenus]]'' β false serotine bats ***genus ''[[Histiotus]]'' β big-eared brown bats ***genus ''[[Ia (genus)|Ia]]'' ***genus ''[[Lasionycteris]]'' ***genus ''[[Scoteanax]]'' β greater broad-nosed bats ***genus ''[[Scotomanes]]'' ***genus ''[[Scotorepens]]'' β lesser broad-nosed bats ***genus ''[[Thainycteris]]'' ** tribe ''[[incertae sedis]]'' ***genus ''[[Rhyneptesicus]]'' **tribe [[Lasiurini]] *** genus ''[[Aeorestes]]'' β hoary bats *** genus ''[[Dasypterus]]'' β yellow bats *** genus ''[[Lasiurus]]'' β hairy-tailed bats ** tribe [[Nycticeiini]] *** genus ''[[Nycticeius]]'' β evening bats ** tribe [[Perimyotini]] ***genus ''[[Parastrellus]]''<ref name="newgenus">{{cite journal|last1=Hoofer|first1=Steven R.|last2=Van Den Bussche|first2=Ronald A.|last3=HorΓ‘Δek|first3=Ivan|date=2006-10-01|title=Generic Status of the American Pipistrelles (Vespertilionidae) with Description of a New Genus|journal=Journal of Mammalogy|volume=87|issue=5|pages=981β992|doi=10.1644/05-MAMM-A-425R1.1|issn=0022-2372|jstor=4094268|doi-access=free}}</ref> *** genus ''[[Perimyotis]]''<ref>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.</ref> ** tribe [[Pipistrellini]] *** genus ''[[Glischropus]]'' β thick-thumbed bats *** genus ''[[Nyctalus]]'' β noctule bats *** genus ''[[Pipistrellus]]'' β true pipistrelles *** genus ''[[Scotoecus]]'' β house bats ***genus ''[[Scotozous]]'' ***genus ''[[RΓΌppell's bat|Vansonia]]'' ** tribe [[Plecotini]] *** genus ''[[Barbastella]]'' β barbastelles or barbastelle bats *** genus ''[[Corynorhinus]]'' β American lump-nosed bats *** genus ''[[Euderma]]'' *** genus ''[[Idionycteris]]'' *** genus ''[[Otonycteris]]'' *** genus ''[[Plecotus]]'' β lump-nosed bats ** tribe [[Scotophilini]] ***genus ''[[Scotophilus]]'' β Old World yellow bats **tribe [[Vespertilionini]] *** genus ''[[Afronycteris]]'' *** genus ''[[Cassistrellus]]'' β helmeted bats ***genus ''[[Chalinolobus]]'' β wattled bats *** genus ''[[Falsistrellus]]'' β false pipistrelles *** genus ''[[Hypsugo]]'' β Asian pipistrelles *** genus ''[[Laephotis]]'' β long-eared bats *** genus ''[[Mimetillus]]'' β mimic bats ***genus ''[[Mirostrellus]]'' *** genus ''[[Neoromicia]]'' ***genus ''[[Nycticeinops]]'' ***genus ''[[Nyctophilus]]'' β New Guinean and Australian big-eared bats ***genus ''[[Pharotis]]'' *** genus ''[[Philetor (genus)|Philetor]]'' *** genus ''[[Pseudoromicia]]'' ***genus ''[[Tylonycteris]]'' β bamboo bats<!-- third species Tylonycteris pygmaeus ("Pygmy Bamboo Bat" perhaps) described in early 2008 --> *** genus ''[[Vespadelus]]'' *** genus ''[[Vespertilio]]'' β frosted bats ==References== {{Reflist|refs= <ref name="Richards2012">{{cite book |last1=Richards |first1=G.C. |last2=Hall |first2=L.S. |last3=Parish |first3=S. (photography) |title=A natural history of Australian bats : working the night shift |date=2012 |publisher=CSIRO Pub |isbn=9780643103740 |page=48}}</ref> <ref name="MSW3">{{MSW3 Chiroptera|id=13801791}}</ref>}} ==Further reading== {{Commons category|Vespertilionidae}} {{Wikispecies|Vespertilionidae}} * 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: 221β234. * Wilson DE, Reeder DM. 2005. Mammal species of the world. Smithsonian Institution Press, Washington DC. {{Taxonbar|from=Q210358}} {{Authority control}} {{DEFAULTSORT:Vesper Bat}} [[Category:Vesper bats| ]] [[Category:Eocene bats|*]] [[Category:Extant Eocene first appearances]] [[Category:Taxa named by John Edward Gray]] [[Category:Bat families]]
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