Editing Megabat (section)

===Taxonomic history===
{{multiple image|perrow = 2|total_width=300
| image1 = Short-nosed Fruit Bat (Cynopterus sphinx) Photograph By Shantanu Kuveskar.jpg
| image2 = Indian Flying Fox (Pteropus giganteus) Kolkata West Bengal India 27042013.png
| image3 = Nyctimene robinsoni.jpg
| image4 = Skraidantis egipto šuo (cropped).jpg
| footer = Megabats of various subfamilies. Clockwise from upper left: [[greater short-nosed fruit bat]] (Cynopterinae), [[Indian flying fox]] (Pteropodinae), [[Egyptian fruit bat]] (Rousettinae), [[eastern tube-nosed bat]] (Nyctimeninae).
}}
{{cladogram|align=right|style=width:30%;line-height:85%|caption=Internal relationships of African Pteropodidae based on combined evidence of [[mitochondrial DNA|mitochondrial]] and [[nuclear DNA|nuclear]] DNA. One species each of Pteropodinae, Nyctimeninae, and Cynopterinae, which are not found in Africa, were included as [[Outgroup (cladistics)|outgroups]].<ref name="Almeida 2016"/>
|cladogram={{clade
|label1=Pteropodidae
|1={{clade
|1={{clade
|1=[[Pteropodinae]]
|2={{clade
  |1=[[Nyctimeninae]]
  |2={{clade
    |1=[[Cynopterinae]]
    |2=[[Eidolinae]] }} }} }}
|label2=[[Rousettinae]]
|2={{clade
    |1=[[Scotonycterini]]
    |2={{clade
      |1=[[Eonycterini]]
      |2={{clade
       |1=[[Rousettini]]
       |2={{clade
         |1=[[Stenonycterini]]
         |2={{clade
           |1={{clade
             |1=[[Plerotini]]
             |2=[[Myonycterini]] }}
           |2=Epomophorini
}} }} }} }} }} }} }} }}
The family Pteropodidae was first described in 1821 by British zoologist [[John Edward Gray]]. He named the family "Pteropidae" (after the genus ''[[Pteropus]]'') and placed it within the now-defunct order Fructivorae.<ref name="Gray">{{cite journal|last=Gray|first=J. E.|year=1821|title=On the natural arrangement of vertebrose animals|url=http://www.rhinoresourcecenter.com/index.php?s=1&act=pdfviewer&id=1175857325&folder=117|journal=London Medical Repository|issue=25|pages=299}}</ref> Fructivorae contained one other family, the now-defunct Cephalotidae, containing one genus, ''Cephalotes''<ref name="Gray"/> (now recognized as a synonym of ''[[Dobsonia]]'').<ref name="Miller, 1907">{{cite journal|last1=Miller|first1=Gerrit S. Jr.|title=The Families and Genera of Bats|journal=United States National Museum Bulletin|date=1907|volume=57|issue=490 |page=63|doi=10.1086/278854 |bibcode=1907ANat...41..671. |url=https://biodiversitylibrary.org/page/51112208}}</ref> Gray's spelling was possibly based on a misunderstanding of the suffix of "''Pteropus''".<ref name="Hutcheon 2006">{{cite journal|last1=Hutcheon| first1= J. M.| last2= Kirsch| first2=J. A.| year=2006| title= A moveable face: deconstructing the Microchiroptera and a new classification of extant bats| journal= Acta Chiropterologica| volume= 8| issue=1| pages= 1–10| doi= 10.3161/1733-5329(2006)8[1:AMFDTM]2.0.CO;2| s2cid= 85948117}}</ref> "''Pteropus''" comes from [[Ancient Greek]] {{Transliteration|grc|pterón}} meaning "wing" and {{Transliteration|grc|poús}} meaning "foot".<ref>{{cite web|url=https://www.merriam-webster.com/dictionary/Pteropus| website=Merriam-Webster|access-date=22 May 2019| title=Definition of PTEROPUS}}</ref> The Greek word ''pous'' of ''Pteropus'' is from the stem word ''pod-''; therefore, Latinizing ''Pteropus'' correctly results in the prefix "''Pteropod-''".<ref name="Jackson">{{cite book| last1=Jackson| first1= S.| last2= Jackson| first2=S. M.| last3= Groves| first3= C.| year=2015| title= Taxonomy of Australian Mammals| publisher= [[Csiro Publishing]]|isbn=9781486300136}}</ref>{{rp|230}} French biologist [[Charles Lucien Bonaparte]] was the first to use the corrected spelling Pteropodidae in 1838.<ref name="Jackson"/>{{Rp|230}}

In 1875, the zoologist [[George Edward Dobson]] was the first to split the order Chiroptera (bats) into two [[suborder]]s: '''Megachiroptera''' (sometimes listed as '''Macrochiroptera''') and [[Microchiroptera]], which are commonly abbreviated to megabats and microbats.<ref name="Dobson 1875">{{cite journal|last=Dobson|first=G. E.| year=1875| title=Conspectus of the suborders, families, and genera of Chiroptera arranged according to their natural affinities| journal=[[Journal of Natural History|The Annals and Magazine of Natural History; Zoology, Botany, and Geology]]|series=4|volume=16|url=https://biodiversitylibrary.org/page/25128227| issue=95}}</ref> Dobson selected these names to allude to the body size differences of the two groups, with many fruit-eating bats being larger than insect-eating bats. Pteropodidae was the only family he included within Megachiroptera.<ref name="Hutcheon 2006"/><ref name="Dobson 1875"/>

A 2001 study found that the dichotomy of megabats and microbats did not accurately reflect their evolutionary relationships. Instead of Megachiroptera and Microchiroptera, the study's authors proposed the new suborders [[Yinpterochiroptera]] and [[Yangochiroptera]].<ref name="Springer"/> This classification scheme has been verified several times subsequently and remains widely supported as of 2019.<ref>{{cite journal|last1=Lei| first1= M.| last2= Dong| first2= D.|year=2016| title= Phylogenomic analyses of bat subordinal relationships based on transcriptome data| journal= [[Scientific Reports]]| volume= 6| issue= 27726| page= 27726| bibcode= 2016NatSR...627726L| doi= 10.1038/srep27726| pmid= 27291671| pmc= 4904216}}</ref><ref>{{cite journal|doi=10.1016/j.cub.2013.09.014|pmid=24184098|title=Phylogenomic Analyses Elucidate the Evolutionary Relationships of Bats|journal=[[Current Biology]]|volume=23|issue=22|pages=2262–2267|year=2013|last1=Tsagkogeorga|first1=G.|last2=Parker|first2=J.|last3=Stupka|first3=E.|last4=Cotton|first4=J.&nbsp;A.|last5=Rossiter|first5=S.&nbsp;J.|s2cid=9133016|doi-access=free|bibcode=2013CBio...23.2262T }}</ref><ref>{{cite journal|doi=10.3109/19401736.2013.809451|pmid=23815317|title=Characterization of the mitochondrial genome of ''Rousettus'' leschenaulti|journal=[[Mitochondrial DNA (journal)|Mitochondrial DNA]]|volume=25|issue=6|pages=443–444|year=2014|last1=Szcześniak|first1=M.|last2=Yoneda|first2=M.|last3=Sato|first3=H.|last4=Makałowska|first4=I.| last5=Kyuwa|first5=S.|last6=Sugano|first6=S.|last7=Suzuki|first7=Y.|last8=Makałowski|first8=W.|last9=Kai|first9=C.|s2cid=207657381}}</ref><ref>{{cite journal|doi=10.1126/science.1105113 |pmid=15681385 |title=A Molecular Phylogeny for Bats Illuminates Biogeography and the Fossil Record |journal=[[Science (journal)|Science]] |volume=307 |issue=5709 |pages=580–584 |year=2005 |last1=Teeling |first1=E. C. |last2=Springer |first2=M. S. |last3=Madsen |first3=O. |last4=Bates |first4=P. |last5=O'Brien |first5=S. J. |last6=Murphy |first6=W. J. |bibcode=2005Sci...307..580T |s2cid=25912333 }}</ref> Since 2005, this suborder has alternatively been called "Pteropodiformes".<ref name="Jackson"/>{{rp|520–521}} Yinpterochiroptera contained species formerly included in Megachiroptera (all of Pteropodidae), as well as several families formerly included in Microchiroptera: [[Megadermatidae]], [[Rhinolophidae]], [[Nycteridae]], [[Craseonycteridae]], and [[Rhinopomatidae]].<ref name="Springer">{{cite journal|doi=10.1073/pnas.111551998|pmid=11353869|pmc=33452|title=Integrated fossil and molecular data reconstruct bat echolocation|journal=Proceedings of the National Academy of Sciences|volume=98|issue=11|pages=6241–6246|year=2001|last1=Springer|first1=M. S.|last2=Teeling|first2=E. C.|last3=Madsen|first3=O.|last4=Stanhope|first4=M. J.|last5=De Jong|first5=W. W.|bibcode=2001PNAS...98.6241S|doi-access=free}}</ref> Two [[superfamily (taxonomy)|superfamilies]] comprise Yinpterochiroptera: Rhinolophoidea—containing the above families formerly in Microchiroptera—and Pteropodoidea, which only contains Pteropodidae.<ref>{{cite book|last=Ungar|first=P.| title=Mammal Teeth: Origin, Evolution, and Diversity| year=2010|publisher=[[JHU Press]]|isbn=9780801899515|page=166}}</ref>

In 1917, Danish [[mammalogist]] [[Knud Andersen (mammalogist)|Knud Andersen]] divided Pteropodidae into three subfamilies: Macroglossinae, Pteropinae (corrected to [[Pteropodinae]]), and Harpyionycterinae.<ref>{{cite journal|doi=10.1111/j.1096-0031.2003.tb00385.x|title=A phylogeny of megachiropteran bats (Mammalia: Chiroptera: Pteropodidae) based on direct optimization analysis of one nuclear and four mitochondrial genes|journal=[[Cladistics (journal)|Cladistics]]|volume=19|issue=6|pages=496–511|year=2003|last1=Giannini|first1=N. P.|last2=Simmons|first2=N. B.|pmid=34905855|s2cid=84696546}}</ref>{{Rp|496}} A 1995 study found that Macroglossinae as previously defined, containing the genera ''[[Eonycteris]]'', ''[[Notopteris]]'', ''[[Macroglossus]]'', ''[[Syconycteris]]'', ''[[Melonycteris]]'', and ''[[Megaloglossus]]'', was [[paraphyly|paraphyletic]], meaning that the subfamily did not group all the descendants of a common ancestor.<ref>{{cite journal|doi=10.1093/sysbio/44.2.209|title=A Phylogeny of Indo-West Pacific Megachiroptera Based on Ribosomal DNA|journal=[[Systematic Biology]]|volume=44|issue=2|pages=209–220|year=1995|last1=Colgan|first1=D. J.|last2=Flannery|first2=T. F.}}</ref>{{Rp|214}} Subsequent publications consider Macroglossini as a tribe within Pteropodinae that contains only ''Macroglossus'' and ''Syconycteris''.<ref>{{cite journal|last1=Bergmans|first1=W.|year=1997|title=Taxonomy and biogeography of African fruit bats (Mammalia, Megachiroptera). 5. The genera Lissonycteris Andersen, 1912, Myonycteris Matschie, 1899 and Megaloglossus Pagenstecher, 1885; general remarks and conclusions; annex: key to all species|journal=Beaufortia|volume=47|issue=2|pages=69}}</ref><ref name="Almeida 2011"/> ''Eonycteris'' and ''Melonycteris'' are within other tribes in Pteropodinae,<ref name="Almeida 2016"/><ref name="Almeida 2011"/> ''Megaloglossus'' was placed in the tribe Myonycterini of the subfamily Rousettinae, and ''Notopteris'' is of uncertain placement.<ref name="Almeida 2011"/>

Other subfamilies and tribes within Pteropodidae have also undergone changes since Andersen's 1917 publication.<ref name="Almeida 2011"/> In 1997, the pteropodids were classified into six subfamilies and nine tribes based on their [[morphology (biology)|morphology]], or physical characteristics.<ref name="Almeida 2011"/> A 2011 genetic study concluded that some of these subfamilies were paraphyletic and therefore they did not accurately depict the relationships among megabat species. Three of the subfamilies proposed in 1997 based on morphology received support: Cynopterinae, Harpyionycterinae, and Nyctimeninae. The other three [[clade]]s recovered in this study consisted of Macroglossini, Epomophorinae + Rousettini, and Pteropodini + ''Melonycteris''.<ref name="Almeida 2011"/> A 2016 genetic study focused only on African pteropodids (Harpyionycterinae, Rousettinae, and Epomophorinae) also challenged the 1997 classification. All species formerly included in Epomophorinae were moved to Rousettinae, which was subdivided into additional tribes. The genus ''[[Eidolon (genus)|Eidolon]]'', formerly in the tribe Rousettini of Rousettinae, was moved to its own subfamily, [[Eidolinae]].<ref name="Almeida 2016">{{cite journal|doi=10.3161/15081109ACC2016.18.1.003|title=The Evolutionary History of the African Fruit Bats (Chiroptera: Pteropodidae)|journal=Acta Chiropterologica|volume=18|pages=73–90|year=2016|last1=Almeida|first1=F.|last2=Giannini|first2=N. P.|last3=Simmons|first3=N. B.|s2cid=89415407|url=https://www.researchgate.net/publication/303742900|hdl=11336/12847|hdl-access=free}}</ref>

In 1984, an additional pteropodid subfamily, Propottininae, was proposed, representing one extinct species described from a fossil discovered in Africa, ''[[Propotto leakeyi]]''.<ref>{{cite journal|last= Butler| first= P. M.| year= 1984| title= Macroscelidea, Insectivora and Chiroptera from the Miocene of east Africa| journal=Palaeovertebrata| volume= 14| issue=3| pages= 175|url=https://palaeovertebrata.com/Articles/sendFile/116/published_article}}</ref> In 2018 the fossils were reexamined and determined to represent a [[lemur]].<ref>{{cite journal|doi=10.1038/s41467-018-05648-w|pmid=30131571|pmc=6104046|title=Fossil lemurs from Egypt and Kenya suggest an African origin for Madagascar's aye-aye|journal=[[Nature Communications]]|volume=9|issue=1|page=3193|year=2018|last1=Gunnell|first1=G. F.|last2=Boyer|first2=D. M.|last3=Friscia|first3=A. R.|last4=Heritage|first4=S.|last5=Manthi|first5=F. K.|last6=Miller|first6=E. R.|last7=Sallam|first7=H. M.|last8=Simmons|first8=N. B.|last9=Stevens|first9=N. J.|last10=Seiffert|first10=E. R.|bibcode=2018NatCo...9.3193G}}</ref> As of 2018, there were 197 described species of megabat,<ref>{{Cite journal|last1=Burgin|first1=Connor J|last2=Colella|first2=Jocelyn P|last3=Kahn|first3=Philip L|last4=Upham|first4=Nathan S|date=2018|title=How many species of mammals are there?|journal=Journal of Mammalogy|volume=99|issue=1|pages=1–14|doi=10.1093/jmammal/gyx147|s2cid=90797674|issn=0022-2372|doi-access=free}}</ref> around a third of which are flying foxes of the genus ''Pteropus''.<ref>{{cite web|url=https://www.iucnredlist.org/search/stats| title=<nowiki>Taxonomy=Pteropus</nowiki>| website=IUCN Red List of Threatened Species|year=2019| access-date=19 May 2019}}</ref>