Editing Megabat (section)

==Taxonomy and evolution==
===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>

===Evolutionary history===
====Fossil record and divergence times====
The fossil record for pteropodid bats is the most incomplete of any bat family. Although the poor skeletal record of Chiroptera is probably from how fragile bat skeletons are, Pteropodidae still have the most incomplete despite generally having the biggest and most sturdy skeletons. It is also surprising that Pteropodidae are the least represented because they were the first major group to diverge.<ref>{{Cite journal |last1=Brown |first1=Emily E. |last2=Cashmore |first2=Daniel D. |last3=Simmons |first3=Nancy B. |last4=Butler |first4=Richard J. |date=2019-03-25 |editor-last=Mannion |editor-first=Philip |title=Quantifying the completeness of the bat fossil record |url=https://onlinelibrary.wiley.com/doi/epdf/10.1111/pala.12426 |journal=Palaeontology |language=en |publisher=Paleobiology Database |volume=62 |issue=5 |pages=757–776 |doi=10.1111/pala.12426 |bibcode=2019Palgy..62..757B |s2cid=133901426 |issn=0031-0239}}</ref> Several factors could explain why so few pteropodid fossils have been discovered: tropical regions where their fossils might be found are under-sampled relative to Europe and North America; conditions for fossilization are poor in the tropics, which could lead to fewer fossils overall; and even when fossils are formed, they may be destroyed by subsequent geological activity.<ref>{{cite journal|last1=Eiting|first1=T. P.|last2=Gunnell|first2=G. F.|year=2009|title=Global Completeness of the Bat Fossil Record|journal=Journal of Mammalian Evolution|volume=16|issue=3|pages=157|doi=10.1007/s10914-009-9118-x|s2cid=5923450}}</ref> It is estimated that more than 98% of pteropodid fossil history is missing.<ref name="Teeling 2005">{{cite journal|doi=10.1126/science.1105113|pmid=15681385|title=A Molecular Phylogeny for Bats Illuminates Biogeography and the Fossil Record|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| url=https://courses.cit.cornell.edu/bionb4240/Reprints/Bat%20phylogeny%20Teeling%20et%20al%202005.pdf}}</ref> Even without fossils, the age and divergence times of the family can still be estimated by using [[computational phylogenetics]]. Pteropodidae split from the superfamily [[Rhinolophoidea]] (which contains all the other families of the suborder Yinpterochiroptera) approximately 58&nbsp;Mya (million years ago).<ref name="Teeling 2005"/> The ancestor of the [[crown group]] of Pteropodidae, or all living species, lived approximately 31&nbsp;Mya.<ref name="Almeida 2009"/>

====Biogeography====
[[File:Oceania UN Geoscheme - Map of Melanesia cropped.jpg|thumb|upright=1.2|alt=A map of Oceania with the islands of Melanesia highlighted in pink.|Melanesia, where many megabat subfamilies are likely to have originated]]
The family Pteropodidae likely originated in [[Australasia]] based on [[biogeography|biogeographic reconstructions]].<ref name="Almeida 2016"/> Other biogeographic analyses have suggested that the [[Melanesia]]n Islands, including [[New Guinea]], are a plausible candidate for the origin of most megabat subfamilies, with the exception of Cynopterinae;<ref name="Almeida 2011">{{cite journal|doi=10.1186/1471-2148-11-281|pmid=21961908|pmc=3199269|title=Evolutionary relationships of the old world fruit bats (Chiroptera, Pteropodidae): Another star phylogeny?|journal=[[BMC Evolutionary Biology]]|volume=11|page=281|year=2011|last1=Almeida|first1=F. C.| last2=Giannini|first2=N. P.|last3=Desalle|first3=R.|last4=Simmons|first4=N. B. |issue=1 |doi-access=free |bibcode=2011BMCEE..11..281A }}</ref> the cynopterines likely originated on the [[Sunda Shelf]] based on results of a Weighted Ancestral Area Analysis of six nuclear and mitochondrial genes.<ref name="Almeida 2009">{{cite journal|doi=10.1016/j.ympev.2009.07.035|pmid=19660560|title=The phylogenetic relationships of cynopterine fruit bats (Chiroptera: Pteropodidae: Cynopterinae)|journal=[[Molecular Phylogenetics and Evolution]]|volume=53|issue=3|pages=772–783|year=2009|last1=Almeida|first1=F. C.|last2=Giannini|first2=N. P.|last3=Desalle|first3=Rob|last4=Simmons|first4=N. B.|hdl=11336/74530|hdl-access=free}}</ref> From these regions, pteropodids colonized other areas, including continental Asia and Africa. Megabats reached Africa in at least four distinct events. The four proposed events are represented by (1) ''[[Scotonycteris]]'', (2) ''[[Rousettus]]'', (3) Scotonycterini, and (4) the "endemic Africa clade", which includes Stenonycterini, Plerotini, Myonycterini, and Epomophorini, according to a 2016 study. It is unknown when megabats reached Africa, but several tribes (Scotonycterini, Stenonycterini, Plerotini, Myonycterini, and Epomophorini) were present by the [[Late Miocene]]. How megabats reached Africa is also unknown. It has been proposed that they could have arrived via the [[Middle East]] before it became more arid at the end of the Miocene. Conversely, they could have reached the continent via the [[Gomphotherium land bridge]], which connected Africa and the [[Arabian Peninsula]] to [[Eurasia]]. The genus ''Pteropus'' (flying foxes), which is not found on mainland Africa, is proposed to have dispersed from Melanesia via [[island hopping]] across the [[Indian Ocean]];<ref>{{cite journal|doi=10.1016/j.ympev.2009.02.010|pmid=19249376|title=Multiple colonisations of the western Indian Ocean by Pteropus fruit bats (Megachiroptera: Pteropodidae): The furthest islands were colonised first|journal=Molecular Phylogenetics and Evolution|volume=51|issue=2|pages=294–303|year=2009|last1=O'Brien|first1=J.|last2=Mariani|first2=C.|last3=Olson|first3=L.|last4=Russell|first4=A. L.|last5=Say|first5=L.|last6=Yoder|first6=A. D.|last7=Hayden|first7=T. J.|bibcode=2009MolPE..51..294O }}</ref> this is less likely for other megabat genera, which have smaller body sizes and thus have more limited flight capabilities.<ref name="Almeida 2016"/>

====Echolocation====
Megabats are the only family of bats incapable of [[larynx|laryngeal]] echolocation. It is unclear whether the common ancestor of all bats was capable of echolocation, and thus echolocation was lost in the megabat lineage, or multiple bat lineages independently evolved the ability to echolocate (the superfamily [[Rhinolophoidea]] and the suborder [[Yangochiroptera]]). This unknown element of bat evolution has been called a "grand challenge in biology".<ref>{{cite book | vauthors = Teeling EC, Jones G, Rossiter SJ |chapter=Phylogeny, Genes, and Hearing: Implications for the Evolution of Echolocation in Bats |date=2016 |pages=25–54 | veditors = Fenton MB, Grinnell AD, Popper AN, Fay RN |series=Springer Handbook of Auditory Research |publisher=Springer | location = New York |doi=10.1007/978-1-4939-3527-7_2 |isbn=9781493935277 |title=Bat Bioacoustics |volume=54 }}</ref> A 2017 study of bat [[ontogeny]] (embryonic development) found evidence that megabat embryos at first have large, developed [[cochlea]] similar to echolocating microbats, though at birth they have small cochlea similar to non-echolocating mammals. This evidence supports that laryngeal echolocation evolved once among bats, and was lost in pteropodids, rather than evolving twice independently.<ref>{{cite journal|doi=10.1038/s41559-016-0021|pmid=28812602|title=Prenatal development supports a single origin of laryngeal echolocation in bats|journal=Nature Ecology & Evolution|volume=1|issue=2|pages=21|year=2017|last1=Wang|first1=Zhe|last2=Zhu|first2=Tengteng|last3=Xue|first3=Huiling|last4=Fang|first4=Na|last5=Zhang|first5=Junpeng|last6=Zhang|first6=Libiao|last7=Pang|first7=Jian|last8=Teeling|first8=Emma C.|last9=Zhang|first9=Shuyi|s2cid=29068452}}</ref> Megabats in the genus ''Rousettus'' are capable of primitive echolocation through clicking their tongues.<ref>{{cite journal | last1= Holland| first1= R. A.| last2= Waters| first2= D. A.| last3= Rayner| first3= J. M.| title = Echolocation signal structure in the Megachiropteran bat Rousettus aegyptiacus Geoffroy 1810 | journal = [[The Journal of Experimental Biology]] | volume = 207 | issue = Pt 25 | pages = 4361–4369 | date = December 2004 | pmid = 15557022 | doi = 10.1242/jeb.01288 | bibcode= 2004JExpB.207.4361H| s2cid= 2715542}}</ref> Some species&mdash;the [[cave nectar bat]] (''Eonycteris spelaea''), [[lesser short-nosed fruit bat]] (''Cynopterus brachyotis''), and the [[long-tongued fruit bat]] (''Macroglossus sobrinus'')&mdash;have been shown to create clicks similar to those of echolocating bats using their wings.<ref name=Boonman2014>{{cite journal | last1 = Boonman| first1= A.| last2= Bumrungsri| first2= S.| last3= Yovel| first3= Y. | title = Nonecholocating fruit bats produce biosonar clicks with their wings | journal = Current Biology | volume = 24 | issue = 24 | pages = 2962–2967 | date = December 2014 | pmid = 25484290 | doi = 10.1016/j.cub.2014.10.077 | s2cid= 17789233| doi-access = free | bibcode= 2014CBio...24.2962B}}</ref>

Both echolocation and [[Bat flight|flight]] are energetically expensive processes separately, although no increase in flight energy expenditure was found for two species of echolocating bats compared with other bats and birds .<ref>{{cite journal | last1= Speakman| first1= J. R.| last2= Racey| first2= P. A. | title = No cost of echolocation for bats in flight | journal = Nature | volume = 350 | issue = 6317 | pages = 421–423 | date = April 1991 | pmid = 2011191 | doi = 10.1038/350421a0 | bibcode = 1991Natur.350..421S | s2cid= 4314715}}</ref> Echolocating bats couple sound production with the mechanisms engaged for flight, allowing them to reduce the additional energy burden of echolocation. Instead of pressurizing a bolus of air for the production of sound, laryngeally echolocating bats likely use the force of the downbeat of their wings to pressurize the air, cutting energetic costs by synchronizing wingbeats and echolocation.<ref>{{cite journal | last1=  Lancaster| first1= W. C.| last2= Henson| first2= O. W.| last3= Keating| first3= A. W. | title = Respiratory muscle activity in relation to vocalization in flying bats | journal = The Journal of Experimental Biology | volume = 198 | issue = Pt 1 | pages = 175–191 | date = January 1995 | doi= 10.1242/jeb.198.1.175| pmid = 7891034| bibcode= 1995JExpB.198..175L|url=http://jeb.biologists.org/content/jexbio/198/1/175.full.pdf }}</ref> The loss of echolocation (or conversely, the lack of its evolution) may be due to the uncoupling of flight and echolocation in megabats.<ref name="Book" /> The larger average body size of megabats compared to echolocating bats<ref name="Hutcheon 2004" /> suggests a larger body size disrupts the flight-echolocation coupling and made echolocation too energetically expensive to be conserved in megabats.<ref name="Book">{{cite book |last=Altringham |first=J. D. | name-list-style = vanc |year=2011 |chapter=Echolocation and other senses| title=Bats: From Evolution to Conservation|location=New York |publisher=[[Oxford University Press]] |isbn=9780199207114 }}</ref>

=== List of genera ===
{{Main|List of pteropodids}}
[[File:Spotted-winged fruit bat Balionycteris maculata.jpg|thumb|upright|alt=A small brown bat with black wings is hanging upside down on a tree branch. Its wings have small, pinkish spots.| The [[spotted-winged fruit bat]] (''Balionycteris maculata'')]]
[[File:Bat Week 2017 - Congressional Reception (37237943654) (cropped).jpg|thumb|alt=A bat with large eyes and a dog-like face in profile. Its fur is a tawny yellow, while the side of its neck is bright yellow.|The [[straw-coloured fruit bat]] (''Eidolon helvum'')]]
[[File:Macrog sobrin 120912-0073 tdp.jpg|thumb|upright|alt=A small, yellowish brown bat clings upside down to a branch with one foot. Its wings are slightly spread and it has a narrow snout.| The [[long-tongued fruit bat]] (''Macroglossus sobrinus'')]]
[[File:Epomophorus wahlbergi in Ruaha National Park 6.jpg|thumb|upright|alt=A bat with its wings wrapped around its body. Its eyes are tawny brown and prominent, and the sun shines through its ear membranes.|[[Wahlberg's epauletted fruit bat]] (''Epomophorus wahlbergi'')]]
The family Pteropodidae is divided into six [[subfamilies]] represented by 46 [[genus|genera]]:<ref name="Almeida 2016"/><ref name="Almeida 2011"/>

Family '''Pteropodidae'''
* subfamily [[Cynopterinae]]<ref name="Almeida 2011"/>
** genus ''[[Aethalops]]'' – pygmy fruit bats
** genus ''[[Alionycteris]]''
** genus ''[[Balionycteris]]''
** genus ''[[Chironax]]''
** genus ''[[Cynopterus]]'' – dog-faced fruit bats or short-nosed fruit bats
** genus ''[[Dyacopterus]]'' – Dayak fruit bats
** genus ''[[Haplonycteris]]''
** genus ''[[Latidens]]''
** genus ''[[Megaerops]]''
** genus ''[[Otopteropus]]''
** genus ''[[Penthetor]]''
** genus ''[[Ptenochirus]]'' – musky fruit bats
** genus ''[[Sphaerias]]''
** genus ''[[Thoopterus]]''
* subfamily [[Eidolinae]]<ref name="Almeida 2016"/>
** genus ''[[Eidolon (genus)|Eidolon]]'' – straw-colored fruit bats
* subfamily Harpyionycterinae<ref name="Almeida 2016"/>
** genus ''[[Aproteles]]''
** genus ''[[Boneia]]''
** genus ''[[Dobsonia]]'' – naked-backed fruit bats
** genus ''[[Harpyionycteris]]''
* subfamily Nyctimeninae<ref name="Almeida 2011"/>
** genus ''[[Nyctimene (genus)|Nyctimene]]'' – tube-nosed fruit bats
** genus ''[[Paranyctimene]]''
* subfamily [[Pteropodinae]]
*** genus ''[[Melonycteris]]''<ref name="Almeida 2011"/>
** tribe [[Pteropodini]]<ref name="Almeida 2011"/>
*** genus ''[[Acerodon]]''
*** genus ''[[Pteralopex]]''
*** genus ''[[Pteropus]]'' – flying foxes
*** genus ''[[Styloctenium]]''
* subfamily [[Rousettinae]]
** tribe [[Eonycterini]]<ref name="Almeida 2016"/>
*** genus ''[[Eonycteris]]'' – dawn fruit bats
** tribe [[Epomophorini]]<ref name="Almeida 2016"/><ref name="Almeida 2011"/>
*** genus ''[[Epomophorus]]'' – epauletted fruit bats
*** genus ''[[Epomops]]'' – epauletted bats
*** genus ''[[Hypsignathus]]''
*** genus ''[[Micropteropus]]'' – dwarf epauletted bats
*** genus ''[[Nanonycteris]]''
** tribe ''[[incertae sedis]]<ref>{{Cite journal|last1=Nesi|first1=Nicolas|last2=Tsagkogeorga|first2=Georgia|last3=Tsang|first3=Susan M|last4=Nicolas|first4=Violaine|last5=Lalis|first5=Aude|last6=Scanlon|first6=Annette T|last7=Riesle-Sbarbaro|first7=Silke A|last8=Wiantoro|first8=Sigit|last9=Hitch|first9=Alan T|last10=Juste|first10=Javier|last11=Pinzari|first11=Corinna A|date=2021-03-04|title=Interrogating Phylogenetic Discordance Resolves Deep Splits in the Rapid Radiation of Old World Fruit Bats (Chiroptera: Pteropodidae)|url=http://dx.doi.org/10.1093/sysbio/syab013|journal=Systematic Biology|volume=70|issue=6|pages=1077–1089|doi=10.1093/sysbio/syab013 | pmc=8513763|pmid=33693838|issn=1063-5157}}</ref>''
***genus ''[[Pilonycteris]]''
**tribe [[Myonycterini]]<ref name="Almeida 2016" />
*** genus ''[[Megaloglossus]]''
*** genus ''[[Myonycteris]]'' – little collared fruit bats
** tribe [[Plerotini]]<ref name="Almeida 2016"/>
*** genus ''[[Plerotes]]''
** tribe [[Rousettini]]<ref name="Almeida 2016"/> 
*** genus ''[[Rousettus]]'' – rousette fruit bats
**tribe [[Scotonycterini]]<ref name="Almeida 2016" />
*** genus ''[[Casinycteris]]''
*** genus ''[[Scotonycteris]]''
** tribe [[Stenonycterini]]<ref name="Almeida 2016"/>
*** genus ''[[Stenonycteris]]''
* ''[[Incertae sedis]]''
** genus ''[[Notopteris]]'' – long-tailed fruit bats<ref name="Almeida 2011"/>
** genus ''[[Mirimiri]]''<ref name="Almeida 2011"/>
** genus ''[[Neopteryx]]''<ref name="Almeida 2011"/>
** genus ''[[Desmalopex]]''<ref name="Almeida 2011"/>
** genus ''{{extinct}}[[Turkanycteris]]''<ref>{{Cite journal| doi = 10.1016/j.jhevol.2018.01.001| pmid = 29628118| issn = 0047-2484| pages = 4| last1 = Gunnell| first1 = Gregg F.| last2 = Manthi| first2 = Fredrick K.| title = Pliocene bats (Chiroptera) from Kanapoi, Turkana Basin, Kenya| journal = Journal of Human Evolution| date = April 2018| volume = 140| s2cid = 206143059| doi-access = free}}</ref>
** tribe [[Macroglossini (Chiroptera)|Macroglossini]]<ref name="Almeida 2011"/>
*** genus ''[[Macroglossus]]'' – long-tongued fruit bats
*** genus ''[[Syconycteris]]'' – blossom bats