Megabat

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Megabats
Temporal range: Oligocene–Recent
Large flying fox, Pteropus vampyrus
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Chiroptera
Suborder: Megachiroptera or Yinpterochiroptera
Dobson, 1875
Family: Pteropodidae
Gray, 1821
Subfamilies

Nyctimeninae
Cynopterinae
Harpiyonycterinae
Macroglossinae
Pteropodinae
Rousettinae
Epomophorinae

Megabats constitute the suborder Megachiroptera, family Pteropodidae of the order Chiroptera (bats). They are also called fruit bats, old world fruit bats,[1] or flying foxes.

Description

Spectacled flying fox (Pteropus conspicillatus)

The megabat, contrary to its name, is not always large: the smallest species is 6 cm (2.4 in) long and thus smaller than some microbats.[2] The largest attain a wingspan of 1.7 m (5.6 ft), weighing in at up to 1.6 kg (3.5 lb).[3] Most fruit bats have large eyes, allowing them to orient themselves visually in twilight and inside caves and forests.

Their sense of smell is excellent. In contrast to the microbats, the fruit bats do not use echolocation (with one exception, the Egyptian fruit bat Rousettus egyptiacus, which uses high-pitched tongue clicks to navigate in caves).[4]

Loss of Echolocation

Megabats make up the only family (Pteropodidae) in order Chiroptera that is not capable of laryngeal echolocation. Echolocation and flight evolved early in the lineage of Chiropterans and echolocation was later lost in family Pteropodidae.[5] Both echolocation and flight are energetically expensive processes for bats.[6] The nature of the flight and echolocation mechanism of bats allows for creation of echolocation pulses with minimal energy use.[7] Energetic coupling of these two processes is thought to have allowed for both energetically expensive processes to evolve in bats. It is hypothesized that the loss of echolocation is due to the uncoupling of flight and echolocation in megabats.[8] The larger average body size of megabats compared to echolocating bats [9] suggests that a larger body size disrupts the flight-echolocation coupling and made echolocation too energetically expensive to be conserved in megabats.[8]

Behavior and ecology

Megabats are frugivorous or nectarivorous, i.e., they eat fruits or lick nectar from flowers. Often the fruits are crushed and only the juices are consumed. The teeth are adapted to bite through hard fruit skins. Large fruit bats must land to eat fruit, while the smaller species are able to hover with flapping wings in front of a flower or fruit.[citation needed]

Frugivorous bats aid the distribution of plants (and therefore, forests) by carrying the fruits with them and spitting the seeds or eliminating them elsewhere. Nectarivores actually pollinate visited plants. They bear long tongues that are inserted deep into the flower; pollen passed to the bat is then transported to the next blossom visited, thereby pollinating it. This relationship between plants and bats is a form of mutualism known as chiropterophily. Examples of plants that benefit from this arrangement include the baobabs of the genus Adansonia and the sausage tree (Kigelia).

As disease reservoirs

Fruit bats have been found to act as reservoirs for a number of diseases which can prove fatal to humans and domestic animals,[10] but the bats themselves sometimes have no signs of infection.

Researchers tested fruit bats for the presence of the Ebola virus between 2001 and 2003. Three species of bats tested positive for Ebola, but had no symptoms of the virus.[10] This indicates the bats may be acting as a reservoir for the virus. Of the infected animals identified during these field collections, immunoglobulin G (IgG) specific for Ebola virus was detected in Hypsignathus monstrosus (Hammer-Headed Bat), Epomops franqueti (Franquet's Epauletted Fruit Bat), and Myonycteris torquata (Little Collared Fruit Bat).

The epidemical Marburg virus was found in 2007 in specimens of the Egyptian fruit bat, confirming the suspicion this species may be a reservoir for this dangerous virus.[11]

Other viral diseases which can be carried by fruit bats include Australian bat lyssavirus and Henipavirus (notably Hendra virus and Nipah virus), both of which can prove fatal to humans. These bats have been shown to infect other species (specifically horses) with Hendra virus in Australian regions. Later, humans became infected with Hendra virus after being exposed to horse body fluids and excretions.[12]

Fruit bats are considered a delicacy by South Pacific Islanders as well as in Micronesia. Consumption has been suggested as a cause of Lytico-Bodig disease on the Micronesian island of Guam, through bioaccumulation of a plant toxin that the bats are immune to.[13]

Classification

Head of a masked flying fox or fruit bat (Pteropus personatus)
Livingstone's fruit bat Pteropus livingstonii
Fox Island, Australia, is believed to be home to the largest colony of flying foxes on the continent.

Bats are usually thought to belong to one of two monophyletic groups, a view that is reflected in their classification into two suborders (Megachiroptera and Microchiroptera). According to this hypothesis, all living megabats and microbats are descendants of a common ancestor species that was already capable of flight.

However, there have been other views, and a vigorous debate persists to this date. For example, in the 1980s and 1990s, some researchers proposed (based primarily on the similarity of the visual pathways) that the Megachiroptera were in fact more closely affiliated with the primates than the Microchiroptera, with the two groups of bats having therefore evolved flight via convergence (see Flying primates theory).[14] However, a recent flurry of genetic studies confirms the more longstanding notion that all bats are indeed members of the same clade, the Chiroptera.[15][16] Other studies have recently suggested that certain families of microbats (possibly the horseshoe bats, mouse-tailed bats and the false vampires) are evolutionarily closer to the fruit bats than to other microbats.[15][17]

List of species

The family Pteropodidae is divided into seven subfamilies with 186 total extant species, represented by 44 - 46 genera:

FAMILY PTEROPODIDAE

  • Subfamily Nyctimeninae
    • Genus Nyctimene - tube-nosed fruit bats
      • Broad-striped Tube-nosed Fruit Bat, Nyctimene aello
      • Common Tube-nosed Fruit Bat, Nyctimene albiventer
      • Pallas's Tube-nosed Fruit Bat, Nyctimene cephalotes
      • Dark Tube-nosed Fruit Bat, Nyctimene celaeno
      • Mountain Tube-nosed Fruit Bat, Nyctimene certans
      • Round-eared Tube-nosed Fruit Bat, Nyctimene cyclotis
      • Dragon Tube-nosed Fruit Bat, Nyctimene draconilla
      • Keast's Tube-nosed Fruit Bat, Nyctimene keasti
      • Island Tube-nosed Fruit Bat, Nyctimene major
      • Malaita Tube-nosed Fruit Bat, Nyctimene malaitensis
      • Demonic Tube-nosed Fruit Bat, Nyctimene masalai
      • Lesser Tube-nosed Bat, Nyctimene minutus
      • Philippine Tube-nosed Fruit Bat, Nyctimene rabori
      • Eastern Tube-nosed Bat, Nyctimene robinsoni
      • Nendo Tube-nosed Fruit Bat, Nyctimene sanctacrucis (early 20th century †)
      • Umboi Tube-nosed Fruit Bat, Nyctimene vizcaccia
    • Genus Paranyctimene
      • Lesser Tube-nosed Fruit Bat, Paranyctimene raptor
      • Steadfast Tube-nosed Fruit Bat, Paranyctimene tenax
  • Subfamily Cynopterinae
    • Genus Aethalops - pygmy fruit bats
      • Borneo Fruit Bat, Aethalops aequalis
      • Pygmy Fruit Bat, Aethalops alecto
    • Genus Alionycteris
      • Mindanao Pygmy Fruit Bat, Alionycteris paucidentata
    • Genus Balionycteris
      • Spotted-winged Fruit Bat, Balionycteris maculata
    • Genus Chironax
      • Black-capped Fruit Bat, Chironax melanocephalus
    • Genus Cynopterus - dog-faced fruit bats or short-nosed fruit bats
      • Lesser Short-nosed Fruit Bat, Cynopterus brachyotis
      • Horsfield’s Fruit Bat, Cynopterus horsfieldii
      • Peters’s Fruit Bat, Cynopterus luzoniensis
      • Minute Fruit Bat, Cynopterus minutus
      • Nusatenggara Short-nosed Fruit Bat, Cynopterus nusatenggara
      • Greater Short-nosed Fruit Bat, Cynopterus sphinx
      • Indonesian Short-nosed Fruit Bat, Cynopterus titthaecheilus
    • Genus Dyacopterus - Dayak fruit bats
      • Brooks’s Dyak Fruit Bat, Dyacopterus brooksi
      • Rickart's Dyak Fruit Bat, Dyacopterus rickarti
      • Dayak Fruit Bat, Dyacopterus spadiceus
    • Genus Haplonycteris
      • Fischer's Pygmy Fruit Bat, Haplonycteris fischeri
    • Genus Latidens
      • Salim Ali's Fruit Bat, Latidens salimalii
    • Genus Megaerops
    • Genus Otopteropus
      • Luzon Fruit Bat, Otopteropus cartilagonodus
    • Genus Penthetor
      • Dusky Fruit Bat, Penthetor lucasi
    • Genus Ptenochirus - musky fruit bats
      • Greater Musky Fruit Bat, Ptenochirus jagori
      • Lesser Musky Fruit Bat, Ptenochirus minor
    • Genus Sphaerias
      • Blanford's Fruit Bat, Sphaerias blanfordi
    • Genus Thoopterus
      • Swift Fruit Bat, Thoopterus nigrescens
  • Subfamily Harpiyonycterinae
    • Genus Aproteles
      • Bulmer's Fruit Bat, Aproteles bulmerae
    • Genus Dobsonia - bare-backed fruit bats
      • Andersen's Bare-backed Fruit Bat, Dobsonia anderseni
      • Beaufort's Naked-backed Fruit Bat, Dobsonia beauforti
      • Philippine Bare-backed Fruit Bat, Dobsonia chapmani
      • Halmahera Naked-backed Fruit Bat, Dobsonia crenulata
      • Biak Naked-backed Fruit Bat, Dobsonia emersa
      • Sulawesi Naked-backed Fruit Bat, Dobsonia exoleta
      • Solomon's Naked-backed Fruit Bat, Dobsonia inermis
      • Lesser Naked-backed Fruit Bat, Dobsonia minor
      • Moluccan Naked-backed Fruit Bat, Dobsonia moluccensis
      • Panniet Naked-backed Fruit Bat, Dobsonia pannietensis
      • Western Naked-backed Fruit Bat, Dobsonia peroni
      • New Britain Naked-backed Fruit Bat, Dobsonia praedatrix
      • Greenish Naked-backed Fruit Bat, Dobsonia viridis
    • Genus Harpyionycteris
      • Sulawesi Harpy Fruit Bat, Harpyionycteris celebensis
      • Harpy Fruit Bat, Harpyionycteris whiteheadi
  • Subfamily Macroglossinae
    • Genus Macroglossus - long-tongued fruit bats
      • Long-tongued Nectar Bat, Macroglossus minimus
      • Long-tongued Fruit Bat, Macroglossus sobrinus
    • Genus Melonycteris
      • Fardoulis' Blossom Bat, Melonycteris fardoulisi
      • Black-bellied Fruit Bat, Melonycteris melanops
      • Woodford's Fruit Bat, Melonycteris woodfordi
    • Genus Notopteris - long-tailed fruit bats
      • Long-tailed Fruit Bat, Notopteris macdonaldi (Fiji and Vanuatu)
      • New Caledonia Blossom Bat, Notopteris neocaledonica (New Caledonia)
    • Genus Syconycteris - blossom bats
      • Common Blossom Bat, Syconycteris australis
      • Halmahera Blossom Bat, Syconycteris carolinae
      • Moss-forest Blossom Bat, Syconycteris hobbit
  • Subfamily Pteropodinae
    • Genus Acerodon
      • Sulawesi Flying Fox, Acerodon celebensis
      • Talaud Flying Fox, Acerodon humilis
      • Giant Golden-crowned Flying Fox, Acerodon jubatus
      • Palawan Fruit Bat, Acerodon leucotis
      • Sunda Flying Fox, Acerodon mackloti
    • Genus Desmalopex
      • White-winged Flying Fox, Desmalopex leucopterus
      • Small White-winged Flying Fox, Desmalopex microleucopterus
    • Genus Eidolon - straw-coloured fruit bats
      • Madagascan Fruit Bat, Eidolon dupreanum
      • Straw-coloured Fruit Bat, Eidolon helvum
    • Genus Mirimiri
      • Fijian Monkey-faced Bat, Mirimiri acrodonta
    • Genus Neopteryx
      • Small-toothed Fruit Bat, Neopteryx frosti
    • Genus Pteralopex
      • Bougainville Monkey-faced Bat, Pteralopex anceps
      • Guadalcanal Monkey-faced Bat, Pteralopex atrata
      • Greater Monkey-faced Bat, Pteralopex flanneryi
      • Montane Monkey-faced Bat, Pteralopex pulchra
      • New Georgian Monkey-faced Bat, Pteralopex taki
    • Genus Pteropus - flying foxes
      • P. alecto species group
        • Black Flying Fox, Pteropus alecto
      • P. caniceps species group
        • Ashy-headed Flying Fox, Pteropus caniceps
      • P. chrysoproctus species group
        • Silvery Flying Fox, Pteropus argentatus
        • Moluccan Flying Fox, Pteropus chrysoproctus
        • Makira Flying Fox, Pteropus cognatus
        • Banks Flying Fox, Pteropus fundatus
        • Solomons Flying Fox, Pteropus rayneri
        • Rennell Flying Fox, Pteropus rennelli
      • P. conspicillatus species group
      • P. livingstonii species group
        • Aru Flying Fox, Pteropus aruensis
        • Kei Flying Fox, Pteropus keyensis
        • Livingstone's Fruit Bat, Pteropus livingstonii
        • Black-bearded Flying Fox, Pteropus melanopogon
      • P. mariannus species group
        • Okinawa Flying Fox, Pteropus loochoensis
        • Mariana Fruit Bat, Pteropus mariannus
        • Pelew Flying Fox, Pteropus pelewensis
        • Kosrae Flying Fox, Pteropus ualanus
        • Yap Flying Fox, Pteropus yapensis
      • P. melanotus species group
        • Black-eared Flying Fox, Pteropus melanotus
      • P. molossinus species group
        • Lombok Flying Fox, Pteropus lombocensis
        • Caroline Flying Fox, Pteropus molossinus
        • Rodrigues Flying Fox, Pteropus rodricensis
      • P. neohibernicus species group
        • Great Flying Fox, Pteropus neohibernicus
      • P. niger species group
      • P. personatus species group
        • Bismark Masked Flying Fox, Pteropus capistratus
        • Masked Flying Fox, Pteropus personatus
        • Temminck's Flying Fox, Pteropus temminckii
      • P. poliocephalus species group
        • Big-eared Flying Fox, Pteropus macrotis
        • Geelvink Bay Flying Fox, Pteropus pohlei
        • Grey-headed Flying Fox, Pteropus poliocephalus
      • P. pselaphon species group
        • Chuuk Flying Fox, Pteropus insularis
        • Temotu Flying Fox, Pteropus nitendiensis
        • Large Palau Flying Fox, Pteropus pilosus (19th century †)
        • Bonin Flying Fox, Pteropus pselaphon
        • Guam Flying Fox, Pteropus tokudae (1970s †)
        • Insular Flying Fox, Pteropus tonganus
        • Vanikoro Flying Fox, Pteropus tuberculatus
        • New Caledonia Flying Fox, Pteropus vetulus
      • P. samoensis species group
        • Vanuatu Flying Fox, Pteropus anetianus
        • Samoa Flying Fox, Pteropus samoensis
      • P. scapulatus species group
        • Gilliard's Flying Fox, Pteropus gilliardorum
        • Lesser Flying Fox, Pteropus mahaganus
        • Little Red Flying Fox, Pteropus scapulatus
        • Dwarf Flying Fox, Pteropus woodfordi
      • P. subniger species group
        • Admiralty Flying Fox, Pteropus admiralitatum
        • Dusky Flying Fox, Pteropus brunneus (19th century †)
        • Ryukyu Flying Fox, Pteropus dasymallus
        • Nicobar Flying Fox, Pteropus faunulus
        • Gray Flying Fox, Pteropus griseus
        • Ontong Java Flying Fox, Pteropus howensis
        • Small Flying Fox, Pteropus hypomelanus
        • Ornate Flying Fox, Pteropus ornatus
        • Little Golden-mantled Flying Fox, Pteropus pumilus
        • Philippine Gray Flying Fox, Pteropus speciosus
        • Small Mauritian Flying Fox, Pteropus subniger (19th century †)
      • P. vampyrus species group
        • Indian Flying Fox, Pteropus giganteus
        • Andersen's Flying Fox, Pteropus intermedius
        • Lyle's Flying Fox, Pteropus lylei
        • Large Flying Fox, Pteropus vampyrus
      • incertae sedis
        • Small Samoan Flying Fox, Pteropus allenorum (19th century †)
        • Large Samoan Flying Fox, Pteropus coxi (19th century †)
    • Genus Styloctenium
  • Subfamily Rousettinae
    • Genus Eonycteris - dawn fruit bats
      • Greater Nectar Bat, Eonycteris major
      • Cave Nectar Bat, Eonycteris spelaea
      • Philippine Dawn Bat, Eonycteris robusta
    • Genus Rousettus - rousette fruit bats
      • Subgenus Boneia
        • Manado Fruit Bat, Rousettus (Boneia) bidens
      • Subgenus Rousettus
        • Geoffroy's Rousette, Rousettus amplexicaudatus
        • Sulawesi Rousette, Rousettus celebensis
        • Egyptian Rousette (Egyptian Fruit Bat), Rousettus aegyptiacus
        • Leschenault's Rousette, Rousettus leschenaulti
        • Linduan Rousette, Rousettus linduensis
        • Comoro Rousette, Rousettus obliviosus
        • Bare-backed Rousette, Rousettus spinalatus
      • Subgenus Stenonycteris
        • Long-haired Rousette, Rousettus (Stenonycteris) lanosus
        • Madagascar Rousette, Rousettus (Stenonycteris) madagascariensis
  • Subfamily Epomophorinae
    • Tribe Epomophorini
      • Genus Epomophorus - epauletted fruit bats
        • Angolan Epauletted Fruit Bat, Epomophorus angolensis
        • Ansell's Epauletted Fruit Bat, Epomophorus anselli
        • Peters's Epauletted Fruit Bat, Epomophorus crypturus
        • Gambian Epauletted Fruit Bat, Epomophorus gambianus
        • Lesser Angolan Epauletted Fruit Bat, Epomophorus grandis
        • Ethiopian Epauletted Fruit Bat, Epomophorus labiatus
        • East African Epauletted Fruit Bat, Epomophorus minimus
        • Minor Epauletted Fruit Bat, Epomophorus minor
        • Wahlberg's Epauletted Fruit Bat, Epomophorus wahlbergi
      • Genus Epomops - epauletted bats
        • Buettikofer's Epauletted Fruit Bat, Epomops buettikoferi
        • Dobson's Fruit Bat, Epomops dobsoni
        • Franquet's Epauletted Fruit Bat, Epomops franqueti
      • Genus Hypsignathus
      • Genus Micropteropus - dwarf epauletted bats
        • Hayman's Dwarf Epauletted Fruit Bat, Micropteropus intermedius
        • Peters's Dwarf Epauletted Fruit Bat, Micropteropus pusillus
      • Genus Nanonycteris
        • Veldkamp's Dwarf Epauletted Fruit Bat, Nanonycteris veldkampii
    • Tribe Myonycterini
      • Genus Lissonycteris
        • Angolan Rousette, Lissonycteris angolensis
      • Genus Megaloglossus
        • Woermann's Bat, Megaloglossus woermanni
      • Genus Myonycteris - little collared fruit bats
        • São Tomé Collared Fruit Bat, Myonycteris brachycephala
        • East African Little Collared Fruit Bat, Myonycteris relicta
        • Little Collared Fruit Bat, Myonycteris torquata
    • Tribe Plerotini
      • Genus Plerotes
        • D'Anchieta's Fruit Bat, Plerotes anchietae
    • Tribe Scotonycterini
      • Genus Casinycteris
        • Short-palated Fruit Bat, Casinycteris argynnis
      • Genus Scotonycteris
        • Zenker's Fruit Bat, Scotonycteris zenkeri
        • Pohle's Fruit Bat, Scotonycteris ophiodon

Footnotes

  1. Mickleburgh, Hutson and Racey. "Old World Fruit Bats:Introduction". International Union for Conservation of Nature. Retrieved July 19, 2013. 
  2. E.g., the Mauritian Tomb Bat. See Garbutt, Nick. "Mauritian Tomb Bat." Mammals of Madagascar: A Complete Guide. Yale University Press. 2007. p. 67.
  3. Nowak, R. M., editor (1999). Walker's Mammals of the World. Vol. 1. 6th edition. Pp. 264-271. ISBN 0-8018-5789-9
  4. Matti Airas. "Echolocation in bats". HUT, Laboratory of Acoustics and Audio Signal Processing. p. 4. Retrieved July 19, 2013. 
  5. De Jong (2001). "Integrated fossil and molecular data reconstruct bat echolocation". Proceedings of the National Academy of Science 98 (11): 6241–6246. Bibcode:2001PNAS...98.6241S. doi:10.1073/pnas.111551998.  More than one of |author= and |last= specified (help)
  6. Racey (1991). "No cost of echolocation for bats in flight". Nature 350 (6317): 421–423. Bibcode:1991Natur.350..421S. doi:10.1038/350421a0. PMID 2011191.  More than one of |author= and |last= specified (help)
  7. Lancaster, W.C., O.W. Henson, and A.W. Keating (1995). "Respiratory muscle activity in relation to vocalization in flying bats". Journal of Experimental Biology 198 (Pt 1): 175–191. PMID 7891034. 
  8. 8.0 8.1 Altringham, J.D. (2011). Echolocation and other senses. New York: Oxford University Press. 
  9. Hutcheon, J.M., and T.J. Garland (1995). "Are megabats big?". Journal of Mammalian Evolution 11 (3/4): 257–277. doi:10.1023/B:JOMM.0000047340.25620.89. 
  10. 10.0 10.1 National Geographic, October 2007. "Deadly Contact," David Quammen, pp. 78-105.
  11. "Deadly Marburg virus discovered in fruit bats". msnbc. August 21, 2007. Retrieved 2008-03-11. 
  12. "Hendra Virus Disease & Nipah Virus Encephalitis Fact Sheet". CDC. 
  13. Monson, C. S.; Banack, S. A.; Cox, P. A. (2003). "Conservation implications of Chamorro consumption of flying foxes as a possible cause of amyotrophic lateral sclerosis-parkinsonism dementia complex in Guam". Conservation Biology 17 (3): 678–686. doi:10.1046/j.1523-1739.2003.02049.x. 
  14. Pettigrew JD, Jamieson BG, Robson SK, Hall LS, McAnally KI, Cooper HM (1989). "Phylogenetic relations between microbats, megabats and primates (Mammalia: Chiroptera and Primates)". Philosophical Transactions of the Royal Society B 325 (1229): 489–559. Bibcode:1989RSPTB.325..489P. doi:10.1098/rstb.1989.0102. 
  15. 15.0 15.1 Eick, GN; Jacobs; Matthee (September 2005). "A nuclear DNA phylogenetic perspective on the evolution of echolocation and historical biogeography of extant bats (chiroptera)" (Free full text). Molecular Biology and Evolution 22 (9): 1869–86. doi:10.1093/molbev/msi180. PMID 15930153. Archived from the original on 2009-02-13.  More than one of |author2= and |last2= specified (help); More than one of |author3= and |last3= specified (help)
  16. Simmons, NB; Seymour, KL; Habersetzer, J; Gunnell, GF (2008-02-14). "Primitive Early Eocene bat from Wyoming and the evolution of flight and echolocation". Nature 451 (7180): 818–21. Bibcode:2008Natur.451..818S. doi:10.1038/nature06549. PMID 18270539. "recent studies unambiguously support bat monophyly" 
  17. Adkins RM, Honeycutt RL (1991). "Molecular phylogeny of the superorder Archonta" (PDF). Proceedings of the National Academy of Sciences of the U.S.A. 88 (22): 10317–10321. Bibcode:1991PNAS...8810317A. doi:10.1073/pnas.88.22.10317. PMC 52919. PMID 1658802. 

References

External links

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