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==Biology and ecology== ===Genome size=== Like all bats, megabats have much smaller [[genome]]s than other mammals. A 2009 study of 43 megabat species found that their genomes ranged from 1.86 picograms (pg, 978 Mbp per pg) in the straw-colored fruit bat to 2.51 pg in [[Lyle's flying fox]] (''Pteropus lylei''). All values were much lower than the mammalian average of 3.5 pg. Megabats have even smaller genomes than microbats, with a mean weight of 2.20 pg compared to 2.58 pg. It was speculated that this difference could be related to the fact that the megabat lineage has experienced an extinction of the [[LINE1]]βa type of [[long interspersed nuclear element]]. LINE1 constitutes 15β20% of the human genome and is considered the most prevalent long interspersed nuclear element among mammals.<ref>{{cite journal|doi=10.1098/rsbl.2009.0016|pmid=19324635|pmc=2679926|title=The genome sizes of megabats (Chiroptera: Pteropodidae) are remarkably constrained|journal=[[Biology Letters]]|volume=5|issue=3|pages=347β351|year=2009|last1=Smith|first1=J. D. L.|last2=Gregory|first2=T. R.}}</ref> ===Senses=== ====Sight==== [[File:1977.05.03 Fijian Monkey-faced Bat ,Taveuni, Fiji 3443 ccccr.jpg|thumb|alt=A headshot of a bat with orangish-brown fur looking directly at the camera. Its eyes are piercingly orange.|The reddish-orange eyes of the [[Fijian monkey-faced bat]] (''Mirimiri acrodonta'')]] With very few exceptions, megabats do not [[Animal echolocation|echolocate]], and therefore rely on sight and smell to navigate.<ref name="Muller"/> They have large eyes positioned at the front of their heads.<ref name="Graydon"/> These are larger than those of the common ancestor of all bats, with one study suggesting a trend of increasing eye size among pteropodids. A study that examined the eyes of 18 megabat species determined that the [[common blossom bat]] (''Syconycteris australis'') had the smallest eyes at a diameter of {{cvt|5.03|mm|in}}, while the largest eyes were those of [[large flying fox]] (''Pteropus vampyrus'') at {{cvt|12.34|mm|in}} in diameter.<ref>{{cite journal|doi=10.1038/s41467-017-02532-x|pmid=29311648|pmc=5758785|title=Auditory opportunity and visual constraint enabled the evolution of echolocation in bats|journal=Nature Communications|volume=9|issue=1|page=98|year=2018|last1=Thiagavel|first1=J.|last2=Cechetto|first2=C.|last3=Santana|first3=S. E.|last4=Jakobsen|first4=L.|last5=Warrant|first5=E. J.|last6=Ratcliffe|first6=J. M.|bibcode=2018NatCo...9...98T}}</ref> Megabat irises are usually brown, but they can be red or orange, as in ''[[Desmalopex]]'', ''[[Mirimiri]]'', ''[[Pteralopex]]'', and some ''Pteropus''.<ref>{{cite journal|doi=10.3161/150811008X331054|title=The systematic position of Pteropus leucopterus and its bearing on the monophyly and relationships of Pteropus (Chiroptera: Pteropodidae)|journal=Acta Chiropterologica|volume=10|pages=11β20|year=2008|last1=Giannini|first1=N. P.|last2=Almeida|first2=F. C.|last3=Simmons|first3=N. B.|last4=Helgen|first4=K. M.|hdl=11336/82001|s2cid=59028595|hdl-access=free}}</ref> At high brightness levels, megabat [[visual acuity]] is poorer than that of humans; at low brightness it is superior.<ref name="Graydon">{{cite journal|journal=[[Journal of the Australian Mammal Society]]| pages=101β105| url=https://books.google.com/books?id=yENMUzuisWwC&pg=RA1-PA101| title=Vision in Flying-Foxes (Chiroptera:Pteropodidae)| volume=10| issue=2| year=1987| last1=Graydon| first1=M.| last2=Giorgi| first2=P.| last3=Pettigrew| first3=J.| doi=10.1071/AM87021| s2cid=254716342}}</ref> One study that examined the eyes of some ''Rousettus'', ''Epomophorus'', ''Eidolon'', and ''Pteropus'' species determined that the first three genera possess a ''[[tapetum lucidum]]'', a reflective structure in the eyes that improves vision at low light levels, while the ''Pteropus'' species do not.<ref name="Muller"/> All species examined had [[retina]]e with both [[rod cell]]s and [[cone cell]]s, but only the ''Pteropus'' species had S-cones, which detect the shortest wavelengths of light; because the spectral tuning of the [[opsin]]s was not discernible, it is unclear whether the S-cones of ''Pteropus'' species detect blue or ultraviolet light. ''Pteropus'' bats are [[Dichromacy|dichromatic]], possessing two kinds of cone cells. The other three genera, with their lack of S-cones, are [[Monochromacy|monochromatic]], unable to see color. All genera had very high densities of rod cells, resulting in high sensitivity to light, which corresponds with their nocturnal activity patterns. In ''Pteropus'' and ''Rousettus'', measured rod cell densities were 350,000β800,000 per square millimeter, equal to or exceeding other nocturnal or [[crepuscular]] animals such as the [[house mouse]], [[domestic cat]], and [[domestic rabbit]].<ref name="Muller">{{cite journal|doi=10.1159/000102971|pmid=17522478|title=Cone Photoreceptor Diversity in the Retinas of Fruit Bats (Megachiroptera)|journal=[[Brain, Behavior and Evolution]]|volume=70|issue=2|pages=90β104|year=2007|last1=MΓΌller|first1=B.|last2=Goodman|first2=S. M.|last3=Peichl|first3=Leo|s2cid=29095435}}</ref> ====Smell==== [[File:EB1911 Chiroptera Fig. 5.jpg|thumb|alt=A scientific illustration of a megabat face in profile with prominent nostrils. Each nostril is a distinct tube projecting away from the face at a right angle.|The nostrils of the [[island tube-nosed fruit bat]] (''Nyctimene major'')]] Megabats use smell to find food sources like fruit and nectar.<ref name="Jones"/> They have keen senses of smell that rival that of the [[domestic dog]].<ref name="Schwab">{{cite journal|doi=10.1136/bjo.2005.077966|pmid=16267906|pmc=1772916|title=A choroidal sleight of hand|journal=[[British Journal of Ophthalmology]]|volume=89|issue=11|page=1398|year=2005|last1=Schwab|first1=I. R.}}</ref> Tube-nosed fruit bats such as the [[eastern tube-nosed bat]] (''Nyctimene robinsoni'') have stereo [[olfaction]], meaning they are able to map and follow odor plumes three-dimensionally.<ref name="Schwab"/> Along with most (or perhaps all) other bat species, megabats mothers and offspring also use scent to recognize each other, as well as for recognition of individuals.<ref name="Jones">{{cite journal|doi=10.3389/fphys.2013.00117|pmid=23755015|pmc=3667242|title=From the ultrasonic to the infrared: Molecular evolution and the sensory biology of bats|journal=[[Frontiers in Physiology]]|volume=4|page=117|year=2013|last1=Jones|first1=G.|last2=Teeling|first2=E. C.|last3=Rossiter|first3=S. J.|doi-access=free}}</ref> In flying foxes, males have enlarged [[androgen]]-sensitive [[sebaceous gland]]s on their shoulders they use for [[Territory (animal)#Spraying|scent-marking]] their territories, particularly during the mating season. The secretions of these glands vary by speciesβof the 65 chemical compounds isolated from the glands of four species, no compound was found in all species.<ref name="Wood 2005"/> Males also engage in [[Urination#Other animals|urine washing]], or coating themselves in their own urine.<ref name="Wood 2005">{{cite journal| last1=Wood| first1= W. F.| last2= Walsh| first2= A.| last3= Seyjagat| first3= J.| last4= Weldon| first4= P. J.| title= Volatile Compounds in Shoulder Gland Secretions of Male Flying Foxes, Genus Pteropus (Pteropodidae, Chiroptera)| year=2005| journal=[[Z Naturforsch C]]| volume=60| issue=9β10| pages= 779β784|url=https://www.researchgate.net/publication/7447695| doi= 10.1515/znc-2005-9-1019| pmid= 16320623| s2cid= 1938648| doi-access= free}}</ref><ref name="Wagner 2008">{{cite journal| last=Wagner| first= J.| year=2008| title= Glandular secretions of male Pteropus (Flying foxes): preliminary chemical comparisons among species|journal=Independent Study Project (Isp) Collection| url=http://digitalcollections.sit.edu/isp_collection/559/}}</ref> ====Taste==== Megabats possess the ''[[TAS1R2]]'' gene, meaning they have the ability to detect sweetness in foods. This gene is present among all bats except [[vampire bat]]s. Like all other bats, megabats cannot taste ''[[umami]]'', due to the absence of the ''[[TAS1R1]]'' gene. Among other mammals, only [[giant panda]]s have been shown to lack this gene.<ref name="Jones"/> Megabats also have multiple ''TAS2R'' genes, indicating that they can taste bitterness.<ref>{{cite journal|doi=10.1093/molbev/mst219|pmid=24202612|pmc=3907052|title=Diet Shapes the Evolution of the Vertebrate Bitter Taste Receptor Gene Repertoire|journal=[[Molecular Biology and Evolution]]|volume=31|issue=2|pages=303β309|year=2014|last1=Li|first1=D.|last2=Zhang|first2=J.}}</ref> ===Reproduction and life cycle=== [[File:Pteropus lylei.jpg|thumb|alt=A female flying fox faces the camera with her wings slightly outstretched. A young flying fox clings to her abdomen, looking at the camera with its eyes open. The mother's eyes are closed and her face is next to her offspring's.|A [[Lyle's flying fox]] (''Pteropus lylei'') with offspring]] Megabats, like all bats, are long-lived relative to their size for mammals. Some captive megabats have had lifespans exceeding thirty years.<ref name="Nowak"/> Relative to their sizes, megabats have low reproductive outputs and delayed sexual maturity, with females of most species not giving birth until the age of one or two.<ref name="Mickleburgh"/>{{rp|6}} Some megabats appear to be able to breed throughout the year, but the majority of species are likely [[seasonal breeder]]s.<ref name="Nowak"/> Mating occurs at the roost.<ref>{{Cite journal| doi = 10.1292/jvms.17-0329| pmid = 28804092| pmc = 5658557| volume = 79| issue = 10| pages = 1667β1674| last1 = Hengjan| first1 = Yupadee| last2 = Iida| first2 = Keisuke| last3 = Doysabas| first3 = Karla Cristine C.| last4 = Phichitrasilp| first4 = Thanmaporn| last5 = Ohmori| first5 = Yasushige| last6 = Hondo| first6 = Eiichi| title = Diurnal behavior and activity budget of the golden-crowned flying fox (''Acerodon jubatus'') in the Subic bay forest reserve area, the Philippines| journal = Journal of Veterinary Medical Science| date = 2017}}</ref> Gestation length is variable,<ref name="Heideman"/> but is four to six months in most species. Different species of megabats have reproductive adaptations that lengthen the period between copulation and giving birth. Some species such as the straw-colored fruit bat have the reproductive adaptation of [[delayed implantation]], meaning that copulation occurs in June or July, but the [[zygote]] does not implant into the [[uterus|uterine]] wall until months later in November.<ref name="Mickleburgh"/>{{rp|6}} The [[Fischer's pygmy fruit bat]] (''Haplonycteris fischeri''), with the adaptation of post-implantation delay, has the longest gestation length of any bat species, at up to 11.5 months.<ref name="Heideman">{{cite journal|doi=10.1111/j.1469-7998.1988.tb02396.x|title=The timing of reproduction in the fruit bat ''Haplonycteris fischeri'' (Pteropodidae): Geographic variation and delayed development|journal=Journal of Zoology|volume=215|issue=4|pages=577β595|year=1988|last1=Heideman|first1=P. D.|hdl=2027.42/72984|hdl-access=free}}</ref> The post-implantation delay means that development of the embryo is suspended for up to eight months after implantation in the uterine wall, which is responsible for its very long pregnancies.<ref name="Mickleburgh"/>{{rp|6}} Shorter gestation lengths are found in the [[greater short-nosed fruit bat]] (''Cynopterus sphinx'') with a period of three months.<ref>{{cite book|first1=R. M.| last1= Nowak| first2= E.| last2= Pillsbury Walker| year=1999|title=Walker's Mammals of the World| volume=1| publisher=JHU Press| isbn=9780801857898|page=287}}</ref> The litter size of all megabats is usually one.<ref name="Mickleburgh">{{cite report|url=https://portals.iucn.org/library/efiles/documents/1992-034.pdf|last1=Mickleburgh| first1= S. P.| last2= Hutson| first2= A. M.| last3= Racey| first3= P. A.| year= 1992| title= Old World fruit bats: An action plan for their conservation| location= Gland, Switzerland| publisher= [[IUCN]]}}</ref>{{rp|6}} There are scarce records of twins in the following species: [[Madagascan flying fox]] (''Pteropus rufus''), [[Dobson's epauletted fruit bat]] (''Epomops dobsoni''), the gray-headed flying fox, the [[black flying fox]] (''Pteropus alecto''), the [[spectacled flying fox]] (''Pteropus conspicillatus''),<ref name="Fox">{{Cite journal| doi = 10.3161/150811008X414845| volume = 10| issue = 2| pages = 271β278| last1 = Fox| first1 = Samantha| last2 = Spencer| first2 = Hugh| last3 = O'Brien| first3 = Gemma M.| title = Analysis of twinning in flying-foxes (Megachiroptera) reveals superfoetation and multiple-paternity| journal = Acta Chiropterologica| date = 2008| s2cid = 83506361}}</ref> the greater short-nosed fruit bat,<ref>{{Cite journal| doi = 10.2307/1379269| volume = 55| issue = 1| pages = 200β202| last1 = Sreenivasan| first1 = M. A.| last2 = Bhat| first2 = H. R.| last3 = Geevarghese| first3 = G.| title = Observations on the Reproductive Cycle of Cynopterus sphinx sphinx Vahl, 1797 (Chiroptera: Pteropidae)| journal = Journal of Mammalogy| date = 1974-03-30| jstor = 1379269| pmid = 4819592}}</ref> [[Peters's epauletted fruit bat]] (''Epomophorus crypturus''), the hammer-headed bat, the straw-colored fruit bat, the [[little collared fruit bat]] (''Myonycteris torquata''), the Egyptian fruit bat, and [[Leschenault's rousette]] (''Rousettus leschenaultii'').<ref name="Douglass"/>{{rp|85β87}} In the cases of twins, it is rare that both offspring survive.<ref name="Fox"/> Because megabats, like all bats, have low reproductive rates, their populations are slow to recover from declines.<ref>{{Cite book| edition = 2nd| publisher = Oxford University Press| isbn = 978-0-19-920711-4| last1 = Altringham| first1 = John D.| last2 = McOwat| first2 = Tom| last3 = Hammond| first3 = Lucy| title = Bats: from evolution to conservation| location = Oxford and New York| date = 2011|page=xv}}</ref> At birth, megabat offspring are, on average, 17.5% of their mother's post-partum weight. This is the smallest offspring-to-mother ratio for any bat family; across all bats, newborns are 22.3% of their mother's post-partum weight. Megabat offspring are not easily categorized into the traditional categories of [[altricial]] (helpless at birth) or [[precocial]] (capable at birth). Species such as the greater short-nosed fruit bat are born with their eyes open (a sign of precocial offspring), whereas the Egyptian fruit bat offspring's eyes do not open until nine days after birth (a sign of altricial offspring).<ref name="Kunz 1987">{{cite journal|last1=Kunz| first1= T. H.| last2= Kurta| first2= A.| year=1987| title= Size of bats at birth and maternal investment during pregnancy| journal= Symposia of the Zoological Society of London| volume=57| url=http://www.bu.edu/cecb/files/2009/08/Symp.pdf}}</ref> As with nearly all bat species, males do not assist females in parental care.<ref name="Safi 2008">{{cite journal|doi=10.1644/08-MAMM-S-058.1|title=Social Bats: The Males' Perspective|journal=[[Journal of Mammalogy]]|volume=89|issue=6|pages=1342β1350|year=2008|last1=Safi|first1=K.|s2cid=85733862|doi-access=free}}</ref> The young stay with their mothers until they are [[Weaning|weaned]]; how long weaning takes varies throughout the family. Megabats, like all bats, have relatively long nursing periods: offspring will nurse until they are approximately 71% of adult body mass, compared to 40% of adult body mass in non-bat mammals.<ref>{{cite book|title=Reproductive Biology of Bats|url=https://archive.org/details/reproductivebiol00cric_666|url-access=limited| editor1-first=E. G.| editor1-last= Crichton| editor2-first= P. H.|editor2-last= Krutzsch| page=[https://archive.org/details/reproductivebiol00cric_666/page/n445 433]| publisher=Academic Press| year=2000| isbn=9780080540535}}</ref> Species in the genus ''[[Micropteropus]]'' wean their young by seven to eight weeks of age, whereas the [[Indian flying fox]] (''Pteropus medius'') does not wean its young until five months of age.<ref name="Douglass">{{cite book|url=https://books.google.com/books?id=yQzSe71g2AcC&pg=PA89| title=Asdell's Patterns of Mammalian Reproduction: A Compendium of Species-specific Data| first1=V.| last1= Douglass Hayssen| first2= A.| last2= Van Tienhoven| first3= A.| last3= Van Tienhoven| year=1993| publisher=[[Cornell University Press]]| isbn=9780801417535|page=89}}</ref> Very unusually, male individuals of two megabat species, the [[Bismarck masked flying fox]] (''Pteropus capistratus'') and the [[Dayak fruit bat]] (''Dyacopterus spadiceus''), have been observed [[male lactation|producing milk]], but there has never been an observation of a male nursing young.<ref>{{cite journal|doi=10.1016/j.tree.2009.03.008|pmid=19427057|title=Galactorrhoea is not lactation|journal=[[Trends in Ecology & Evolution]]|volume=24|issue=7|pages=354β355|year=2009|last1=Racey|first1=D. N.|last2=Peaker|first2=M.|last3=Racey|first3=P. A.|bibcode=2009TEcoE..24..354R }}</ref> It is unclear if the lactation is functional and males actually nurse pups or if it is a result of [[stress (biology)|stress]] or [[malnutrition]].<ref name="Kunz 2009">{{cite journal|doi=10.1016/j.tree.2008.09.009|pmid=19100649|title=Male lactation: Why, why not and is it care?|journal=Trends in Ecology & Evolution|volume=24|issue=2|pages=80β85|year=2009|last1=Kunz|first1=T. H|last2=Hosken|first2=David J|bibcode=2009TEcoE..24...80K }}</ref> ===Behavior and social systems=== [[File:Flying foxes.png|thumb|alt=A colony of megabats roosting in a tree during the daytime. They appear as black shapes evenly dispersed throughout the canopy of the tree.|A group of roosting megabats of the genus ''Pteropus'']] Many megabat species are highly [[gregarious]] or social. Megabats will vocalize to communicate with each other, creating noises described as "trill-like bursts of sound",<ref>{{cite journal|doi=10.3161/150811012X661729|title=Vocalizations in the Malagasy Cave-Dwelling Fruit Bat, Eidolon dupreanum: Possible Evidence of Incipient Echolocation?|last1=Schoeman| first1= M. C.| last2= Goodman| first2= S. M. |journal=Acta Chiropterologica|volume=14|issue=2|page=409|year=2012|s2cid=86613252}}</ref> honking,<ref>{{cite magazine|url=http://www.batcon.org/resources/media-education/bats-magazine/bat_article/1500| title=Hammer-headed Fruit Bat| volume=34| issue=1| magazine=BATS Magazine| year=2015| access-date=30 May 2019}}</ref> or loud, bleat-like calls<ref>{{cite journal|doi=10.1093/mspecies/sex007|title=Nyctimene robinsoni (Chiroptera: Pteropodidae)|journal=[[Mammalian Species]]|volume=49|issue=949|pages=68β75|year=2017|last1=Loveless|first1=A. M.|last2=McBee|first2=K.|s2cid=89828640|doi-access=free}}</ref> in various genera. At least one species, the Egyptian fruit bat, is capable of a kind of [[vocal learning]] called vocal production learning, defined as "the ability to modify vocalizations in response to interactions with conspecifics".<ref>{{Cite journal|last1=Prat|first1=Yosef|last2=Taub|first2=Mor|last3=Yovel|first3=Yossi|date=2015|title=Vocal learning in a social mammal: Demonstrated by isolation and playback experiments in bats|journal=Science Advances|language=en|volume=1|issue=2|pages=e1500019|doi=10.1126/sciadv.1500019|issn=2375-2548|pmc=4643821|pmid=26601149|bibcode=2015SciA....1E0019P}}</ref><ref>{{cite journal|doi=10.3758/s13423-016-1060-3|pmid=27368623|pmc=5325843|title=What bats have to say about speech and language|journal=[[Psychonomic Bulletin & Review]]|volume=24|issue=1|pages=111β117|year=2017|last1=Vernes|first1=S. C.}}</ref> Young Egyptian fruit bats are capable of acquiring a [[dialect]] by listening to their mothers, as well as other individuals in their colonies. It has been postulated that these dialect differences may result in individuals of different colonies communicating at different frequencies, for instance.<ref>{{Cite journal|last1=Prat|first1=Yosef|last2=Azoulay|first2=Lindsay|last3=Dor|first3=Roi|last4=Yovel|first4=Yossi|date=2017|title=Crowd vocal learning induces vocal dialects in bats: Playback of conspecifics shapes fundamental frequency usage by pups|journal=PLOS Biology|language=en|volume=15|issue=10|pages=e2002556|doi=10.1371/journal.pbio.2002556|issn=1545-7885|pmc=5663327|pmid=29088225 |doi-access=free }}</ref><ref>{{cite web|url=https://www.the-scientist.com/notebook/what-bat-quarrels-tell-us-about-vocal-learning-30141| title=What Bat Quarrels Tell Us About Vocal Learning| last=Zimmer| first=K.| date=1 January 2018| access-date=23 May 2019|website=The Scientist}}</ref> Megabat social behavior includes using sexual behaviors for more than just reproduction. Evidence suggests that female Egyptian fruit bats take food from males in exchange for sex. Paternity tests confirmed that the males from which each female scrounged food had a greater likelihood of fathering the scrounging female's offspring.<ref>{{cite journal|doi=10.1016/j.cub.2019.04.066|pmid=31130455|title=Food for Sex in Bats Revealed as Producer Males Reproduce with Scrounging Females|journal=Current Biology|volume=29|issue=11|pages=1895β1900.e3|year=2019|last1=Harten|first1=L.|last2=Prat|first2=Y.|last3=Ben Cohen|first3=S.|last4=Dor|first4=R.|last5=Yovel|first5=Y.|s2cid=162183551|doi-access=free|bibcode=2019CBio...29E1895H }}</ref> Homosexual fellatio has been observed in at least one species, the [[Bonin flying fox]] (''Pteropus pselaphon'').<ref name="Sugita 2016">{{cite journal|doi=10.1371/journal.pone.0166024|pmid=27824953|pmc=5100941|title=Homosexual Fellatio: Erect Penis Licking between Male Bonin Flying Foxes Pteropus pselaphon|journal=PLOS ONE|volume=11|issue=11|page=e0166024|year=2016|last1=Sugita|first1=N.|bibcode=2016PLoSO..1166024S|doi-access=free}}</ref><ref name="Tan 2009">{{cite journal|doi=10.1371/journal.pone.0007595|pmid=19862320|pmc=2762080|title=Fellatio by Fruit Bats Prolongs Copulation Time|journal=PLOS ONE|volume=4|issue=10|page=e7595|year=2009|last1=Tan|first1=M.|last2=Jones|first2=G.|last3=Zhu|first3=G.|last4=Ye|first4=J.|last5=Hong|first5=T. |last6=Zhou|first6=S. |last7=Zhang|first7=S. |last8=Zhang|first8=L. |bibcode=2009PLoSO...4.7595T|doi-access=free}}</ref> This same-sex fellatio is hypothesized to encourage colony formation of otherwise-antagonistic males in colder climates.<ref name="Sugita 2016"/><ref name="Tan 2009"/> Megabats are mostly [[nocturnal]] and [[crepuscular]], though some have been observed flying during the day.<ref name="Nowak 1994">{{cite book| last1= Nowak| first1= R. M.| last2= Walker| first2= E. P.| last3= Kunz| first3= T. H.| last4= Pierson| first4= E. D.| year= 1994| title= Walker's bats of the world| publisher= JHU Press| page= [https://archive.org/details/walkersbatsofwor00rona/page/49 49]| isbn= 9780801849862| url-access= registration| url= https://archive.org/details/walkersbatsofwor00rona/page/49}}</ref> A few island species and subspecies are [[diurnality|diurnal]], hypothesized as a response to a lack of [[predators]]. Diurnal taxa include a subspecies of the [[black-eared flying fox]] (''Pteropus melanotus natalis''), the [[Mauritian flying fox]] (''Pteropus niger''), the [[Caroline flying fox]] (''Pteropus molossinus''), a subspecies of ''[[Pteropus pelagicus]]'' (''P. p. insularis''), and the [[Seychelles fruit bat]] (''Pteropus seychellensis'').<ref name="Pierson 1992"/>{{rp|9}} ====Roosting==== A 1992 summary of forty-one megabat genera noted that twenty-nine are tree-roosting genera. A further eleven genera roost in caves, and the remaining six genera roost in other kinds of sites (human structures, mines, and crevices, for example). Tree-roosting species can be solitary or highly [[colony (biology)|colonial]], forming aggregations of up to one million individuals. Cave-roosting species form aggregations ranging from ten individuals up to several thousand. Highly colonial species often exhibit roost fidelity, meaning that their trees or caves may be used as roosts for many years. Solitary species or those that aggregate in smaller numbers have less fidelity to their roosts.<ref name="Mickleburgh"/>{{rp|2}} ===Diet and foraging=== [[File:Eye contact with flying fox.jpg|thumb|upright=1.2|alt=A flying fox with reddish-yellow fur and a dark brown snout is in flight facing the viewer. The background is white.|[[Indian flying fox]] (''Pteropus medius'') in flight]] Most megabats are primarily [[frugivore|frugivorous]].<ref name="Dumont"/> Throughout the family, a diverse array of fruit is consumed from nearly 188 plant genera.<ref name="Yin"/> Some species are also [[nectarivore|nectarivorous]], meaning that they also drink nectar from flowers.<ref name="Dumont">{{cite journal|doi=10.1644/BOS-107|title=Food Hardness and Feeding Behavior in Old World Fruit Bats (Pteropodidae)|journal=Journal of Mammalogy|volume=85|pages=8β14|year=2004|last1=Dumont|first1=E. R.|last2=O'Neal|first2=R.|s2cid=27275791|doi-access=free}}</ref> In Australia, ''[[Eucalyptus]]'' flowers are an especially important food source.<ref name="Nelson"/> Other food resources include leaves, shoots, buds, pollen, seed pods, sap, cones, bark, and twigs.<ref>{{cite journal|doi=10.1046/j.1365-2907.1998.00033.x|title=Dietary strategies of Old World Fruit Bats (Megachiroptera, Pteropodidae): How do they obtain sufficient protein?|journal=[[Mammal Review]]|volume=28|issue=4|pages=185β194|year=1998|last1=Courts|first1=S. E.|doi-access=free|bibcode=1998MamRv..28..185C }}</ref> They are prodigious eaters and can consume up to 2.5 times their own body weight in fruit per night.<ref name="Yin">{{cite journal|doi=10.1371/journal.pone.0146274|pmid=26735303|pmc=4703304|title=Molecular Evolution of the Nuclear Factor (Erythroid-Derived 2)-Like 2 Gene Nrf2 in Old World Fruit Bats (Chiroptera: Pteropodidae)|journal=PLOS ONE|volume=11|issue=1|page=e0146274|year=2016|last1=Yin|first1=Q.|last2=Zhu|first2=L.|last3=Liu|first3=D.|last4=Irwin|first4=D. M.|last5=Zhang|first5=S.|last6=Pan|first6=Y.|bibcode=2016PLoSO..1146274Y|doi-access=free}}</ref> Megabats fly to roosting and foraging resources. They typically fly straight and relatively fast for bats; some species are slower with greater maneuverability. Species can commute {{cvt|20-50|km|mi}} in a night. [[animal migration|Migratory species]] of the genera ''Eidolon'', ''Pteropus'', ''Epomophorus'', ''Rousettus'', ''Myonycteris'', and ''Nanonycteris'' can migrate distances up to {{cvt|750|km|mi}}. Most megabats have below-average [[Bat flight#Aspect ratio|aspect ratios]],<ref name=Norberg1987>{{cite journal |author1=Norberg, U.M. |author2=Rayner, J.M.V. |name-list-style=amp | date = 1987 | title = Ecological morphology and flight in bats (Mammalia: Chiroptera): wing adaptations, flight performance, foraging strategy and echolocation | journal = Philosophical Transactions of the Royal Society B | volume = 316 | issue = 1179 | pages = 382β383 | doi = 10.1098/rstb.1987.0030|bibcode=1987RSPTB.316..335N }}</ref> which is measurement relating wingspan and wing area.<ref name=Norberg1987/>{{rp|348}} Wing loading, which measures weight relative to wing area,<ref name=Norberg1987/>{{rp|348}} is average or higher than average in megabats.<ref name=Norberg1987/> ====Seed dispersal==== Megabats play an important role in [[seed dispersal]]. As a result of their long evolutionary history, some plants have evolved characteristics compatible with bat senses, including fruits that are strongly scented, brightly colored, and prominently exposed away from foliage. The bright colors and positioning of the fruit may reflect megabats' reliance on visual cues and inability to navigate through clutter. In a study that examined the fruits of more than forty fig species, only one fig species was consumed by both birds and megabats; most species are consumed by one or the other. Bird-consumed figs are frequently red or orange, while megabat-consumed figs are often yellow or green.<ref>{{cite journal|doi=10.1111/j.1744-7429.2003.tb00606.x|title=Fruit Bats (Chiroptera: Pteropodidae) as Seed Dispersers and Pollinators in a Lowland Malaysian Rain Forest1|journal=[[Biotropica]]|volume=35|issue=4|pages=491β502|year=2003|last1=Hodgkison|first1=R. |last2=Balding|first2=S. T.|last3=Zubaid|first3=A. |last4=Kunz|first4=T. H.|bibcode=2003Biotr..35..491H |s2cid=86327074}}</ref> Most seeds are excreted shortly after consumption due to a rapid gut transit time, but some seeds can remain in the gut for more than twelve hours. This heightens megabats' capacity to disperse seeds far from parent trees.<ref>{{cite journal|doi=10.1098/rspb.1999.0625|pmc=1689670|title=Old World fruit bats can be long-distance seed dispersers through extended retention of viable seeds in the gut|journal=[[Proceedings of the Royal Society of London. Series B: Biological Sciences]]|volume=266|issue=1416|pages=219β223|year=1999|last1=Shilton|first1=L. A.|last2=Altringham|first2=J. D.|last3=Compton|first3=S. G.|last4=Whittaker|first4=R. J.}}</ref> As highly mobile frugivores, megabats have the capacity to restore forest between isolated forest fragments by dispersing tree seeds to deforested landscapes.<ref>{{cite journal|doi=10.1016/j.gecco.2015.02.012|title=High-resolution GPS tracking reveals habitat selection and the potential for long-distance seed dispersal by Madagascan flying foxes Pteropus rufus|journal=Global Ecology and Conservation|volume=3|page=690|year=2015|last1=Oleksy|first1=R. |last2=Racey|first2=P. A.|last3=Jones|first3=G.|doi-access=free|bibcode=2015GEcoC...3..678O }}</ref> This dispersal ability is limited to plants with small seeds that are less than {{cvt|4|mm|in}} in length, as seeds larger than this are not ingested.<ref>{{cite journal|doi=10.1016/j.gecco.2017.04.007|title=Frugivory and seed dispersal by vertebrates in tropical and subtropical Asia: An update|journal=Global Ecology and Conservation|volume=11|page=13|year=2017|last1=Corlett|first1=R. T.|doi-access=free|bibcode=2017GEcoC..11....1C }}</ref> ===Predators and parasites=== [[File:Hardwicke's science-gossip - an illustrated medium of interchange and gossip for students and lovers of nature (1886) (14769069502).jpg|thumb|alt=A drawing of a small insect with spider-like legs|An example of a [[Nycteribiidae|bat fly]], a flightless [[Diptera|fly]] that parasitizes bats, including megabats]] Megabats, especially those living on islands, have few native predators. Non-native predators of flying foxes include domestic [[cat]]s and [[rat]]s. The [[mangrove monitor]], which is a native predator for some megabat species but an introduced predator for others, opportunistically preys on megabats, as it is a capable tree climber.<ref name=":0">{{cite journal|doi=10.3897/zookeys.345.5840|pmid=24194666|pmc=3817444|title=Taxonomy, distribution, and natural history of flying foxes (Chiroptera, Pteropodidae) in the Mortlock Islands and Chuuk State, Caroline Islands|url=https://archive.org/details/pubmed-PMC3817444|journal=[[ZooKeys]]|issue=345|year=2013|last1=Buden|first1=D. |last2=Helgen|first2=K. M.|last3=Wiles|first3=G.|pages=97β135|doi-access=free|bibcode=2013ZooK..345...97B }}</ref> Another species, the [[brown tree snake]], can seriously impact megabat populations; as a non-native predator in [[Guam]], the snake consumes so many offspring that it reduced the [[recruitment (biology)|recruitment]] of the population of the [[Mariana fruit bat]] (''Pteropus mariannus'') to essentially zero. The island is now considered a [[Sourceβsink dynamics|sink]] for the Mariana fruit bat, as its population there relies on bats immigrating from the nearby island of [[Rota (island)|Rota]] to bolster it rather than successful reproduction.<ref name="Esselstyn 2006">{{cite journal|doi=10.1353/psc.2006.0027|title=Impact of Posttyphoon Hunting on Mariana Fruit Bats (Pteropus mariannus)|journal=[[Pacific Science]]|volume=60|issue=4|pages=531β532|year=2006|last1=Esselstyn|first1=J. A. |last2=Amar|first2=A. |last3=Janeke|first3=D. |s2cid=55543225}}</ref> Predators that are naturally [[sympatric]] with megabats include reptiles such as [[crocodilian]]s, snakes, and large lizards, as well as birds like [[falcon]]s, [[hawk]]s, and [[owl]]s.<ref name="Mickleburgh"/>{{rp|5}} The [[saltwater crocodile]] is a known predator of megabats, based on analysis of crocodile stomach contents in northern Australia.<ref>{{cite journal|doi=10.1371/journal.pone.0197159|pmid=29874276|pmc=5991389|title=Estuarine crocodiles in a tropical coastal floodplain obtain nutrition from terrestrial prey|journal=PLOS ONE|volume=13|issue=6|page=e0197159|year=2018|last1=Adame|first1=Maria Fernanda|last2=Jardine|first2=T. D.|last3=Fry|first3=B. |last4=Valdez|first4=D. |last5=Lindner|first5=G. |last6=Nadji|first6=J. |last7=Bunn|first7=S. E.|bibcode=2018PLoSO..1397159A|doi-access=free}}</ref> During extreme heat events, megabats like the [[little red flying fox]] (''Pteropus scapulatus'') must cool off and rehydrate by drinking from waterways, making them susceptible to opportunistic depredation by [[freshwater crocodiles]].<ref>{{cite AV media| url=https://www.youtube.com/watch?v=wi30w-Mk2yQ| archive-url=https://ghostarchive.org/varchive/youtube/20211107/wi30w-Mk2yQ| archive-date=2021-11-07 | url-status=live| date=10 April 2015|medium=video| publisher=BBC Earth|title=Flying Foxes Vs Freshwater Crocodile|access-date=22 May 2019}}{{cbignore}}</ref> Megabats are the hosts of several [[parasite]] taxa. Known parasites include [[Nycteribiidae]] and [[Streblidae]] species ("bat flies"),<ref>{{cite journal|doi=10.1186/s13071-017-2582-x|pmid=29284533|pmc=5747079|title=Hidden diversity of Nycteribiidae (Diptera) bat flies from the Malagasy region and insights on host-parasite interactions|journal=[[Parasites & Vectors]]|volume=10| issue=1|page=630| year=2017|last1=Ramasindrazana| first1=B. |last2=Goodman|first2=S. M.|last3=Gomard|first3=Y. |last4=Dick|first4=C. W.|last5=Tortosa|first5=P. |doi-access=free }}</ref><ref>{{cite journal|doi=10.1186/s13071-018-2918-1|pmid=29859123|pmc=5984742|title=Rates of hematophagous ectoparasite consumption during grooming by an endemic Madagascar fruit bat|journal=Parasites & Vectors|volume=11|issue=1|page=330|year=2018|last1=Ramanantsalama|first1=R. V.|last2=Andrianarimisa|first2=A.|last3=Raselimanana|first3=A. P.|last4=Goodman|first4=S. M. |doi-access=free }}</ref> as well as [[Acari|mites]] of the genus ''[[Demodex]]''.<ref>{{cite journal|last1=Desch| first1= C. E.| year=1981| title= A new species of demodicid mite (Acari: Prostigmata) from Western Australia parasitic on Macroglossus minimus (Chiroptera: Pteropodidae)| journal= [[Records of the Western Australian Museum]]| volume= 9| issue=1| pages= 41β47| url=http://museum.wa.gov.au/sites/default/files/A%20NEW%20SPECIES%20OF%20DEMODICID%20MITE%20(ACARI%20PROSTIGMATA)%20FROM%20WESTERN%20AUSTRALIA%20PARASITIC%20ON%20MACROGLOSSUS%20MINIMUS%20(CHIROPTERA%20PT.pdf}}</ref> Blood parasites of the family [[Haemoproteidae]] and intestinal nematodes of [[Toxocaridae]] also affect megabat species.<ref name="Nelson"/><ref>{{cite journal|doi=10.1051/parasite/2012192137|pmid=22550624|pmc=3671437|title=The haemosporidian parasites of bats with description of ''Sprattiella'' alectogen. Nov., sp. Nov|journal=Parasite|volume=19|issue=2|pages=137β146|year=2012|last1=Landau|first1=I.|last2=Chavatte|first2=J. M.|last3=Karadjian |first3=G.|last4=Chabaud|first4=A.|last5=Beveridge|first5=I.}}</ref>
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