Sear

Bat

Bat s are mammals of t he order Chiropt era.[a] Wit h t heir forelimbs adapt ed as wings, t hey are t he
only mammals capable of t rue and sust ained flight . Bat s are more agile in flight t han most birds,
flying wit h t heir very long spread-out digit s covered wit h a t hin membrane or pat agium. The
smallest bat , and arguably t he smallest ext ant mammal, is Kit t i's hog-nosed bat , which is 29–34
millimet res (11⁄8–13⁄8 inches) in lengt h, 150 mm (6 in) across t he wings and 2–2.6 g
(1⁄16–3⁄32 oz) in mass. The largest bat s are t he flying foxes, wit h t he giant golden-crowned flying
fox, Acerodon jubatus, reaching a weight of 1.6 kg (31⁄2 lb) and having a wingspan of 1.7 m (5 ft
7 in).
 Bat
Temporal range:

Scient ific classificat ion

Kingdom: Animalia

Phylum: Chordat a

Class: Mammalia

Clade: Scrot ifera

Order: Chiropt era

Blumenbach, 1779

Suborders

(t radit ional):

Megachiropt era

Microchiropt era

(present ):
 Yinpt erochiropt era

Yangochiropt era

Worldwide distribution of bat species

The second largest order of mammals aft er rodent s, bat s comprise about 20% of all classified
mammal species worldwide, wit h over 1,400 species. These were t radit ionally divided int o t wo
suborders: t he largely fruit -eat ing megabat s, and t he echolocat ing microbat s. But more recent
evidence has support ed dividing t he order int o Yinpt erochiropt era and Yangochiropt era, wit h
megabat s as members of t he former along wit h several species of microbat s. Many bat s are
insect ivores, and most of t he rest are frugivores (fruit -eat ers) or nect arivores (nect ar-eat ers). A
few species feed on animals ot her t han insect s; for example, t he vampire bat s feed on blood.
Most bat s are noct urnal, and many roost in caves or ot her refuges; it is uncert ain whet her bat s
have t hese behaviours t o escape predat ors. Bat s are present t hroughout t he world, wit h t he
except ion of ext remely cold regions. They are import ant in t heir ecosyst ems for pollinat ing
flowers and dispersing seeds; many t ropical plant s depend ent irely on bat s for t hese services.

Bat s provide humans wit h some direct benefit s, at t he cost of some disadvant ages. Bat dung
has been mined as guano from caves and used as fert iliser. Bat s consume insect pest s, reducing
t he need for pest icides and ot her insect management measures. They are somet imes numerous
enough and close enough t o human set t lement s t o serve as t ourist at t ract ions, and t hey are
used as food across Asia and t he Pacific Rim. However, fruit bat s are frequent ly considered
pest s by fruit growers. Due t o t heir physiology, bat s are one t ype of animal t hat act s as a nat ural
reservoir of many pat hogens, such as rabies; and since t hey are highly mobile, social, and long-
lived, t hey can readily spread disease among t hemselves. If humans int eract wit h bat s, t hese
t rait s become pot ent ially dangerous t o humans. Some bat s are also predat ors of mosquit oes,
suppressing t he t ransmission of mosquit o-borne diseases.
Depending on t he cult ure, bat s may be symbolically associat ed wit h posit ive t rait s, such as
prot ect ion from cert ain diseases or risks, rebirt h, or long life, but in t he West , bat s are popularly
associat ed wit h darkness, malevolence, wit chcraft , vampires, and deat h.

Etymology

An older English name for bat s is flit t ermouse, which mat ches t heir name in ot her Germanic
languages (for example German Fledermaus and Swedish fladdermus), relat ed t o t he flut t ering
of wings. Middle English had bakke, most likely cognat e wit h Old Swedish natbakka ("night -bat "),
which may have undergone a shift from -k- t o -t- (t o Modern English bat) influenced by Lat in
blatta, "mot h, noct urnal insect ". The word "bat " was probably first used in t he early 1570s.[2][3]
The name "Chiropt era" derives from Ancient Greek: χείρ – cheir, "hand"[4] and πτερόν – pteron,
"wing".[1][5]

Phylogeny and taxonomy

The early Eocene fossil microchiropteran Icaronycteris, from the Green River Formation
Evolution

The delicat e skelet ons of bat s do not fossilise well; it is est imat ed t hat only 12% of bat genera
t hat lived have been found in t he fossil record.[6] Most of t he oldest known bat fossils were
already very similar t o modern microbat s, such as Archaeopteropus (32 million years ago).[7] The
ext inct bat s Palaeochiropteryx tupaiodon (48 million years ago) and Hassianycteris kumari
(48 million years ago) are t he first fossil mammals whose colourat ion has been discovered: bot h
were reddish-brown.[8][9]

Bat s were formerly grouped in t he superorder Archont a, along wit h t he t reeshrews (Scandent ia),
colugos (Dermopt era), and primat es.[10] Modern genet ic evidence now places bat s in t he
superorder Laurasiat heria, wit h it s sist er t axon as Fereuungulat a, which includes carnivorans,
pangolins, odd-t oed ungulat es, even-t oed ungulat es, and cet aceans.[11][12][13][14][15] One st udy
places Chiropt era as a sist er t axon t o odd-t oed ungulat es (Perissodact yla).[16]
   Euarchontoglires (primates, treeshrews, rodents, rabbits)
 

  Eulipotyphla (hedgehogs, shrews, moles, solenodons)

 

  Chiroptera (bats)
 

  Pholidota (pangolins)
 

Ferae Carnivora (cats, hyenas, dogs, bears,

Boreoeutheria
 
   
Laurasiatheria   seals)
 
Scrotifera
 
Fereuungulata Perissodactyla (horses, tapirs, rhinos)
 
 
 

Euungulata
  Cetartiodactyla (camels, ruminants,
 
  whales)

Phylogenetic tree showing Chiroptera within Laurasiatheria, with Fereuungulata as its sister taxon according to a 2013
study[15]

The flying primat e hypot hesis proposed t hat when adapt at ions t o flight are removed, megabat s
are allied t o primat es by anat omical feat ures not shared wit h microbat s and t hus flight evolved
t wice in mammals.[17] Genet ic st udies have st rongly support ed t he monophyly of bat s and t he
single origin of mammal flight .[7][17]
Chiroptera
 
  Megachiroptera Pteropodidae (megabats)
   

  Microchiroptera
   
  Megadermatidae (false vampire bats)
 

  Craseonycteridae (Kitti's hog-nosed bat)

 
 
 
  Rhinolophoidea   Rhinopomatidae (mouse-tailed bats)
     
 
 
  Hipposideridae (Old World leaf-nosed bats)
 
 
 
  Rhinolophidae (horseshoe bats)
 

  Yangochiroptera    
            Miniopteridae (long winged bat)
   

   
   
  Noctilionidae (fisherman bats)
 
 
 
  Mormoopidae (Pteronotus)
 

 
Mystacinidae (New Zealand short-tailed
 
 
bats)
   
 
  Thyropteridae (disc-winged bats)
 

  Furipteridae
   
   
 
  Mormoopidae (Mormoops)
 
 Phyllostomidae (New World leaf-nosed bats)
 
 

  Molossidae (free-tailed bats)

 
 
 
  Emballonuridae (sac-winged bats)
 

  Myzopodidae (sucker-footed bats)

 
 
      Emballonuridae (Taphozous)
   

 
    Natalidae (funnel-eared bats)
 
 
 
  Vespertilionidae (vesper bats)
 

Internal relationships of the Chiroptera, divided into the traditional megabat and microbat clades, according to a 2011
study[18]

Genet ic evidence indicat es t hat megabat s originat ed during t he early Eocene, and belong wit hin
t he four major lines of microbat s.[15] Two new suborders have been proposed; Yinpt erochiropt era
includes t he Pt eropodidae, or megabat family, as well as t he families Rhinolophidae,
Hipposideridae, Craseonyct eridae, Megadermat idae, and Rhinopomat idae.[19] Yangochiropt era
includes t he ot her families of bat s (all of which use laryngeal echolocat ion), a conclusion
support ed by a 2005 DNA st udy.[19] A 2013 phylogenomic st udy support ed t he t wo new
proposed suborders.[15]
   Yangochiroptera (as above)
 

  Pteropodidae (megabats)
 

Chiroptera  
   
  Yinpterochiroptera   Megadermatidae (false vampire bats)
     
  Rhinolophoidea
   
  horseshoe bats and allies
 

Internal relationships of the Chiroptera, with the megabats subsumed within Yinpterochiroptera, according to a 2013 study[15]

Giant golden-crowned flying fox, Acerodon jubatus

The 2003 discovery of an early fossil bat from t he 52-million-year-old Green River Format ion,
Onychonycteris finneyi, indicat es t hat flight evolved before echolocat ive abilit ies.[20][21]
Onychonycteris had claws on all five of it s fingers, whereas modern bat s have at most t wo claws
on t wo digit s of each hand. It also had longer hind legs and short er forearms, similar t o climbing
mammals t hat hang under branches, such as slot hs and gibbons. This palm-sized bat had short ,
broad wings, suggest ing t hat it could not fly as fast or as far as lat er bat species. Inst ead of
flapping it s wings cont inuously while flying, Onychonycteris probably alt ernat ed bet ween flaps
and glides in t he air.[7] This suggest s t hat t his bat did not fly as much as modern bat s, but flew
from t ree t o t ree and spent most of it s t ime climbing or hanging on branches.[22] The dist inct ive
feat ures of t he Onychonycteris fossil also support t he hypot hesis t hat mammalian flight most
likely evolved in arboreal locomot ors, rat her t han t errest rial runners. This model of flight
development , commonly known as t he "t rees-down" t heory, holds t hat bat s first flew by t aking
advant age of height and gravit y t o drop down on t o prey, rat her t han running fast enough for a
ground-level t ake off.[23][24]

The molecular phylogeny was cont roversial, as it point ed t o microbat s not having a unique
common ancest ry, which implied t hat some seemingly unlikely t ransformat ions occurred. The
first is t hat laryngeal echolocat ion evolved t wice in bat s, once in Yangochiropt era and once in t he
rhinolophoids.[25] The second is t hat laryngeal echolocat ion had a single origin in Chiropt era, was
subsequent ly lost in t he family Pt eropodidae (all megabat s), and lat er evolved as a syst em of
t ongue-clicking in t he genus Rousettus.[26] Analyses of t he sequence of t he vocalizat ion gene
FoxP2 were inconclusive on whet her laryngeal echolocat ion was lost in t he pt eropodids or
gained in t he echolocat ing lineages.[27] Echolocat ion probably first derived in bat s from
communicat ive calls. The Eocene bat s Icaronycteris (52 million years ago) and Palaeochiropteryx
had cranial adapt at ions suggest ing an abilit y t o det ect ult rasound. This may have been used at
first mainly t o forage on t he ground for insect s and map out t heir surroundings in t heir gliding
phase, or for communicat ive purposes. Aft er t he adapt at ion of flight was est ablished, it may
have been refined t o t arget flying prey by echolocat ion.[22] Analyses of t he hearing gene Prestin
seem t o favour t he idea t hat echolocat ion developed independent ly at least t wice, rat her t han
being lost secondarily in t he pt eropodids,[28] but ont ogenic analysis of t he cochlea support s t hat
laryngeal echolocat ion evolved only once.[29]

Classification

Bat s are placent al mammals. Aft er rodent s, t hey are t he largest order, making up about 20% of
mammal species.[30] In 1758, Carl Linnaeus classified t he seven bat species he knew of in t he
genus Vespertilio in t he order Primat es. Around t went y years lat er, t he German nat uralist Johann
Friedrich Blumenbach gave t hem t heir own order, Chiropt era.[31] Since t hen, t he number of
described species has risen t o over 1,400,[32] t radit ionally classified as t wo suborders:
Megachiropt era (megabat s), and Microchiropt era (microbat s/echolocat ing bat s).[33] Not all
megabat s are larger t han microbat s.[34] Several charact erist ics dist inguish t he t wo groups.
Microbat s use echolocat ion for navigat ion and finding prey, but megabat s apart from t hose in t he
genus Rousettus do not .[35] Accordingly, megabat s have a well-developed eyesight .[33] Megabat s
have a claw on t he second finger of t he forelimb.[36][37] The ext ernal ears of microbat s do not
close t o form a ring; t he edges are separat ed from each ot her at t he base of t he ear.[37]
Megabat s eat fruit , nect ar, or pollen, while most microbat s eat insect s; ot hers feed on fruit ,
nect ar, pollen, fish, frogs, small mammals, or blood.[33]

"Chiroptera" from Ernst Haeckel's Kunstformen der Natur, 1904

Below is a t able chart following t he bat classificat ion of families recognized by various aut hors
of t he nint h volume of Handbook of the Mammals of the World published in 2019:[38]
 Chiropt era Blumenbach, 1779

Yinpt erochiropt era Springer, Teeling, Madsen, St anhope & Jong, 2001

Pt eropodoidea J. E. Gray, 1821

Number of Image
Family English Name
Species Figure

Pt eropodidae J. E. Gray, 1821 Old World fruit bat s 191

Rhinolophoidea J. E. Gray, 1825

Number of Image
Family English Name
Species Figure

Rhinopomat idae Bonapart e,

Mouse-t ailed bat s 6
1838

Craseonyct eridae Hill, 1974 Hog-nosed bat 1

Megadermat idae H. Allen, 1864 False-vampires 6

Rhinonyct eridae J. E. Gray, 1866 Trident bat s 9

Hipposideridae Lydekker, 1891 Old World leaf-nosed bat s 88

Rhinolophidae J. E. Gray, 1825 Horseshoe bat s 109

Yangochiropt era Koopman, 1984

Emballonuroidea Gervais in de Cast elnau, 1855

Family English Name Number of Image

 Species Figure

Nyct eridae Van der Hoeven,

Slit -faced bat s 15
1855

Emballonuridae Gervais in de
Sheat h-t ailed bat s 54
Cast elnau, 1855

Noct ilionoidea J. E. Gray, 1821

Number of Image
Family English Name
Species Figure

Myzopodidae Thomas, 1904 Madagascar sucker-foot ed bat s 2

Myst acinidae Dobson, 1875 New Zealand short -t ailed bat s 2

Thyropt eridae Miller, 1907 Disk-winged bat s 5

Fuript eridae J. E. Gray, 1866 Smoky bat and t humbless bat 2

Noct ilionidae J. E. Gray, 1821 Bulldog bat s 2

Ghost -faced, naked-backed and

Mormoopidae Saussure, 1860 18
must ached bat s

Phyllost omidae J. E. Gray, 1825 New World leaf-nosed bat s 217

 Vespert ilionoidea J. E. Gray, 1821

Number of Image
Family English Name
Species Figure

Nat alidae J. E. Gray, 1825 Funnel-eared bat s 12

Molossidae Gervais in de
Free-t ailed bat s 126
Cast elnau, 1855

Miniopt eridae Dobson, 1875 Free-t ailed bat s 38

Cist ugidae Lack et al., 2010 Wing-gland bat s 2

Vespert ilionidae J. E. Gray, 1821 Vesper bat s 496

Anatomy and physiology

Skull and dentition

A preserved megabat showing how the skeleton fits inside its skin
The head and t eet h shape of bat s can vary by species. In general, megabat s have longer snout s,
larger eye socket s and smaller ears, giving t hem a more dog-like appearance, which is t he source
of t heir nickname of "flying foxes".[39] Among microbat s, longer snout s are associat ed wit h
nect ar-feeding.[40] while vampire bat s have reduced snout s t o accommodat e large incisors and
canines.[41]

Small insect -eat ing bat s can have as many as 38 t eet h, while vampire bat s have only 20. Bat s
t hat feed on hard-shelled insect s have fewer but larger t eet h wit h longer canines and more
robust lower jaws t han species t hat prey on soft er bodied insect s. In nect ar-feeding bat s, t he
canines are long while t he cheek-t eet h are reduced. In fruit -eat ing bat s, t he cusps of t he cheek
t eet h are adapt ed for crushing.[40] The upper incisors of vampire bat s lack enamel, which keeps
t hem razor-sharp.[41] The bit e force of small bat s is generat ed t hrough mechanical advant age,
allowing t hem t o bit e t hrough t he hardened armour of insect s or t he skin of fruit .[42]

Wings and flight

Bat s are t he only mammals capable of sust ained flight , as opposed t o gliding, as in t he flying
squirrel.[43] The fast est bat , t he Mexican free-t ailed bat (Tadarida brasiliensis), can achieve a
ground speed of 160 km/h (100 mph).[44]

Play media

Little brown bat take off and flight

The finger bones of bat s are much more flexible t han t hose of ot her mammals, owing t o t heir
flat t ened cross-sect ion and t o low levels of calcium near t heir t ips.[45][46] The elongat ion of bat
digit s, a key feat ure required for wing development , is due t o t he upregulat ion of bone
morphogenet ic prot eins (Bmps). During embryonic development , t he gene cont rolling Bmp
signalling, Bmp2, is subject ed t o increased expression in bat forelimbs – result ing in t he
ext ension of t he manual digit s. This crucial genet ic alt erat ion helps creat e t he specialized limbs
required for powered flight . The relat ive proport ion of ext ant bat forelimb digit s compared wit h
t hose of Eocene fossil bat s have no significant differences, suggest ing t hat bat wing
morphology has been conserved for over fift y million years.[47] During flight , t he bones undergo
bending and shearing st ress; t he bending st resses felt are smaller t han in t errest rial mammals,
but t he shearing st ress is larger. The wing bones of bat s have a slight ly lower breaking st ress
point t han t hose of birds.[48]

As in ot her mammals, and unlike in birds, t he radius is t he main component of t he forearm. Bat s
have five elongat ed digit s, which all radiat e around t he wrist . The t humb point s forward and
support s t he leading edge of t he wing, and t he ot her digit s support t he t ension held in t he wing
membrane. The second and t hird digit s go along t he wing t ip, allowing t he wing t o be pulled
forward against aerodynamic drag, wit hout having t o be t hick as in pt erosaur wings. The fourt h
and fift h digit s go from t he wrist t o t he t railing edge, and repel t he bending force caused by air
pushing up against t he st iff membrane.[49] Due t o t heir flexible joint s, bat s are more
maneuverable and more dext erous t han gliding mammals.[50]

Wing membranes (patagia) of Townsend's big-eared bat, Corynorhinus townsendii

The wings of bat s are much t hinner and consist of more bones t han t he wings of birds, allowing
bat s t o maneuver more accurat ely t han t he lat t er, and fly wit h more lift and less drag.[51] By
folding t he wings in t oward t heir bodies on t he upst roke, t hey save 35 percent energy during
flight .[52] The membranes are delicat e, t earing easily,[53] but can regrow, and small t ears heal
quickly.[53][54] The surface of t he wings is equipped wit h t ouch-sensit ive recept ors on small
bumps called Merkel cells, also found on human fingert ips. These sensit ive areas are different in
bat s, as each bump has a t iny hair in t he cent er, making it even more sensit ive and allowing t he
bat t o det ect and adapt t o changing airflow; t he primary use is t o judge t he most efficient speed
at which t o fly, and possibly also t o avoid st alls.[55] Insect ivorous bat s may also use t act ile hairs
t o help perform complex maneuvers t o capt ure prey in flight .[50]

The pat agium is t he wing membrane; it is st ret ched bet ween t he arm and finger bones, and down
t he side of t he body t o t he hind limbs and t ail. This skin membrane consist s of connect ive t issue,
elast ic fibres, nerves, muscles, and blood vessels. The muscles keep t he membrane t aut during
flight .[56] The ext ent t o which t he t ail of a bat is at t ached t o a pat agium can vary by species,
wit h some having complet ely free t ails or even no t ails.[40] The skin on t he body of t he bat , which
has one layer of epidermis and dermis, as well as hair follicles, sweat glands and a fat t y
subcut aneous layer, is very different from t he skin of t he wing membrane. The pat agium is an
ext remely t hin double layer of epidermis; t hese layers are separat ed by a connect ive t issue
cent er, rich wit h collagen and elast ic fibers. The membrane has no hair follicles or sweat glands,
except bet ween t he fingers.[55][57] For bat embryos, apopt osis (cell deat h) affect s only t he
hindlimbs, while t he forelimbs ret ain webbing bet ween t he digit s t hat forms int o t he wing
membranes.[58] Unlike birds, whose st iff wings deliver bending and t orsional st ress t o t he
shoulders, bat s have a flexible wing membrane t hat can resist only t ension. To achieve flight , a
bat exert s force inwards at t he point s where t he membrane meet s t he skelet on, so t hat an
opposing force balances it on t he wing edges perpendicular t o t he wing surface. This adapt at ion
does not permit bat s t o reduce t heir wingspans, unlike birds, which can part ly fold t heir wings in
flight , radically reducing t he wing span and area for t he upst roke and for gliding. Hence bat s
cannot t ravel over long dist ances as birds can.[49]

Nect ar- and pollen-eat ing bat s can hover, in a similar way t o hummingbirds. The sharp leading
edges of t he wings can creat e vort ices, which provide lift . The vort ex may be st abilized by t he
animal changing it s wing curvat ures.[59]

Roosting and gaits

Group of megabats roosting

When not flying, bat s hang upside down from t heir feet , a post ure known as roost ing.[60] The
femurs are at t ached at t he hips in a way t hat allows t hem t o bend out ward and upward in flight .
The ankle joint can flex t o allow t he t railing edge of t he wings t o bend downwards. This does not
permit many movement s ot her t han hanging or clambering up t rees.[49] Most megabat s roost
wit h t he head t ucked t owards t he belly, whereas most microbat s roost wit h t he neck curled
t owards t he back. This difference is reflect ed in t he st ruct ure of t he cervical or neck vert ebrae
in t he t wo groups, which are clearly dist inct .[60] Tendons allow bat s t o lock t heir feet closed
when hanging from a roost . Muscular power is needed t o let go, but not t o grasp a perch or when
holding on.[61]

When on t he ground, most bat s can only crawl awkwardly. A few species such as t he New
Zealand lesser short -t ailed bat and t he common vampire bat are agile on t he ground. Bot h
species make lat eral gait s (t he limbs move one aft er t he ot her) when moving slowly but vampire
bat s move wit h a bounding gait (all limbs move in unison) at great er speeds, t he folded up wings
being used t o propel t hem forward. Vampire bat likely evolved t hese gait s t o follow t heir host s
while short -t ailed bat s developed in t he absence of t errest rial mammal compet it ors. Enhanced
t errest rial locomot ion does not appear t o have reduced t heir abilit y t o fly.[62]

Internal systems

Bat s have an efficient circulat ory syst em. They seem t o make use of part icularly st rong
venomot ion, a rhyt hmic cont ract ion of venous wall muscles. In most mammals, t he walls of t he
veins provide mainly passive resist ance, maint aining t heir shape as deoxygenat ed blood flows
t hrough t hem, but in bat s t hey appear t o act ively support blood flow back t o t he heart wit h t his
pumping act ion.[63][64] Since t heir bodies are relat ively small and light weight , bat s are not at risk
of blood flow rushing t o t heir heads when roost ing.[65]

Bat s possess a highly adapt ed respirat ory syst em t o cope wit h t he demands of powered flight ,
an energet ically t axing act ivit y t hat requires a large cont inuous t hroughput of oxygen. In bat s,
t he relat ive alveolar surface area and pulmonary capillary blood volume are larger t han in most
ot her small quadrupedal mammals.[66] During flight t he respirat ory cycle has a one-t o-one
relat ionship wit h t he wing-beat cycle.[67] Because of t he rest raint s of t he mammalian lungs, bat s
cannot maint ain high-alt it ude flight .[49]
The wings are highly vascularized membranes, the larger blood vessels visible against the light.[68]

It t akes a lot of energy and an efficient circulat ory syst em t o work t he flight muscles of bat s.
Energy supply t o t he muscles engaged in flight requires about double t he amount compared t o
t he muscles t hat do not use flight as a means of mammalian locomot ion. In parallel t o energy
consumpt ion, blood oxygen levels of flying animals are t wice as much as t hose of t heir
t errest rially locomot ing mammals. As t he blood supply cont rols t he amount of oxygen supplied
t hroughout t he body, t he circulat ory syst em must respond accordingly. Therefore, compared t o
a t errest rial mammal of t he same relat ive size, t he bat 's heart can be up t o t hree t imes larger,
and pump more blood.[69] Cardiac out put is direct ly derived from heart rat e and st roke volume of
t he blood;[70] an act ive microbat can reach a heart rat e of 1000 beat s per minut e.[71]

Wit h it s ext remely t hin membranous t issue, a bat 's wing can significant ly cont ribut e t o t he
organism's t ot al gas exchange efficiency.[57] Because of t he high energy demand of flight , t he
bat 's body meet s t hose demands by exchanging gas t hrough t he pat agium of t he wing. When t he
bat has it s wings spread it allows for an increase in surface area t o volume rat io. The surface
area of t he wings is about 85% of t he t ot al body surface area, suggest ing t he possibilit y of a
useful degree of gas exchange.[57] The subcut aneous vessels in t he membrane lie very close t o
t he surface and allow for t he diffusion of oxygen and carbon dioxide.[72]

The digest ive syst em of bat s has varying adapt at ions depending on t he species of bat and it s
diet . As in ot her flying animals, food is processed quickly and effect ively t o keep up wit h t he
energy demand. Insect ivorous bat s may have cert ain digest ive enzymes t o bet t er process
insect s, such as chit inase t o break down chit in, which is a large component of insect s.[73] Vampire
bat s, probably due t o t heir diet of blood, are t he only vert ebrat es t hat do not have t he enzyme
malt ase, which breaks down malt sugar, in t heir int est inal t ract . Nect ivorous and frugivorous bat s
have more malt ase and sucrase enzymes t han insect ivorous, t o cope wit h t he higher sugar
cont ent s of t heir diet .[74]

The adapt at ions of t he kidneys of bat s vary wit h t heir diet s. Carnivorous and vampire bat s
consume large amount s of prot ein and can out put concent rat ed urine; t heir kidneys have a t hin
cort ex and long renal papillae. Frugivorous bat s lack t hat abilit y and have kidneys adapt ed for
elect rolyt e-ret ent ion due t o t heir low-elect rolyt e diet ; t heir kidneys accordingly have a t hick
cort ex and very short conical papillae.[74] Bat s have higher met abolic rat es associat ed wit h flying,
which lead t o an increased respirat ory wat er loss. Their large wings are composed of t he highly
vascularized membranes, increasing t he surface area, and leading t o cut aneous evaporat ive
wat er loss.[68] Wat er helps maint ain t heir ionic balance in t heir blood, t hermoregulat ion syst em,
and removal of wast es and t oxins from t he body via urine. They are also suscept ible t o blood
urea poisoning if t hey do not receive enough fluid.[75]

The st ruct ure of t he ut erine syst em in female bat s can vary by species, wit h some having t wo
ut erine horns while ot hers have a single mainline chamber.[76]

Senses

Echolocat ion

Pipistrellus pulses (0:05)

Time-expanded recording of Pipistrellus pipistrellus bat echolocat ion calls and social call

Problems playing this file? See media help.

Feeding buzz (0:07)

Time-expanded recording of t he feeding buzz of a bat homing in on it s prey

Problems playing this file? See media help.

Microbat s and a few megabat s emit ult rasonic sounds t o produce echoes. Sound int ensit y of
t hese echos are dependent on subglot t ic pressure. The bat s' cricot hyroid muscle cont rols t he
orient at ion pulse frequency, which is an import ant funct ion. This muscle is locat ed inside t he
larynx and it is t he only t ensor muscle capable of aiding phonat ion.[77] By comparing t he out going
pulse wit h t he ret urning echoes, bat s can gat her informat ion on t heir surroundings. This allows
t hem t o det ect prey in darkness.[78] Some bat calls can reach 140 decibels.[79] Microbat s use
t heir larynx t o emit echolocat ion signals t hrough t he mout h or t he nose.[80] Microbat calls range
in frequency from 14,000 t o well over 100,000 Hz, ext ending well beyond t he range of human
hearing (bet ween 20 and 20,000 Hz).[81] Various groups of bat s have evolved fleshy ext ensions
around and above t he nost rils, known as nose-leaves, which play a role in sound t ransmission.[82]

Principle of bat echolocation: orange is the call and green is the echo.

In low-dut y cycle echolocat ion, bat s can separat e t heir calls and ret urning echoes by t ime. They
have t o t ime t heir short calls t o finish before echoes ret urn.[83] The delay of t he ret urning echoes
allows t he bat t o est imat e t he range t o t heir prey.[81] In high-dut y cycle echolocat ion, bat s emit
a cont inuous call and separat e pulse and echo in frequency using t he Doppler effect of t heir
mot ion in flight . The shift of t he ret urning echoes yields informat ion relat ing t o t he mot ion and
locat ion of t he bat 's prey. These bat s must deal wit h changes in t he Doppler shift due t o
changes in t heir flight speed. They have adapt ed t o change t heir pulse emission frequency in
relat ion t o t heir flight speed so echoes st ill ret urn in t he opt imal hearing range.[83][84]

In addit ion t o echolocat ing prey, bat ears are sensit ive t o sounds made by t heir prey, such as t he
flut t ering of mot h wings. The complex geomet ry of ridges on t he inner surface of bat ears helps
t o sharply focus echolocat ion signals, and t o passively list en for any ot her sound produced by
t he prey. These ridges can be regarded as t he acoust ic equivalent of a Fresnel lens, and exist in a
large variet y of unrelat ed animals, such as t he aye-aye, lesser galago, bat -eared fox, mouse
lemur, and ot hers.[85][86][87] Bat s can est imat e t he elevat ion of t heir t arget using t he int erference
pat t erns from t he echoes reflect ing from t he t ragus, a flap of skin in t he ext ernal ear.[81]

The tiger moth (Bertholdia trigona) can jam bat echolocation.[88][89]

By repeat ed scanning, bat s can ment ally const ruct an accurat e image of t he environment in
which t hey are moving and of t heir prey.[90] Some species of mot h have exploit ed t his, such as
t he t iger mot hs, which produces aposemat ic ult rasound signals t o warn bat s t hat t hey are
chemically prot ect ed and t herefore dist ast eful.[88][89] Mot h species including t he t iger mot h can
produce signals t o jam bat echolocat ion. Many mot h species have a hearing organ called a
t ympanum, which responds t o an incoming bat signal by causing t he mot h's flight muscles t o
t wit ch errat ically, sending t he mot h int o random evasive manoeuvres.[91][92][93]

Vision

The eyes of most microbat species are small and poorly developed, leading t o poor visual acuit y,
but no species is blind.[94] Most microbat s have mesopic vision, meaning t hat t hey can det ect
light only in low levels, whereas ot her mammals have phot opic vision, which allows colour vision.
Microbat s may use t heir vision for orient at ion and while t ravelling bet ween t heir roost ing grounds
and feeding grounds, as echolocat ion is effect ive only over short dist ances. Some species can
det ect ult raviolet (UV). As t he bodies of some microbat s have dist inct colorat ion, t hey may be
able t o discriminat e colours.[43][95][96][97]

Megabat species oft en have eyesight as good as, if not bet t er t han, human vision. Their eyesight
is adapt ed t o bot h night and daylight vision, including some colour vision.[97]

Magnet orecept ion

Microbat s make use of magnet orecept ion, in t hat t hey have a high sensit ivit y t o t he Eart h's
magnet ic field, as birds do. Microbat s use a polarit y-based compass, meaning t hat t hey
different iat e nort h from sout h, unlike birds, which use t he st rengt h of t he magnet ic field t o
different iat e lat it udes, which may be used in long-dist ance t ravel. The mechanism is unknown but
may involve magnet it e part icles.[98][99]

Thermoregulation

Thermographic image of a bat using trapped air as insulation

Most bat s are homeot hermic (having a st able body t emperat ure), t he except ion being t he vesper
bat s (Vespert ilionidae), t he horseshoe bat s (Rhinolophidae), t he free-t ailed bat s (Molossidae),
and t he bent -winged bat s (Miniopt eridae), which ext ensively use het erot hermy (where body
t emperat ure can vary).[100] Compared t o ot her mammals, bat s have a high t hermal conduct ivit y.
The wings are filled wit h blood vessels, and lose body heat when ext ended. At rest , t hey may
wrap t heir wings around t hemselves t o t rap a layer of warm air. Smaller bat s generally have a
higher met abolic rat e t han larger bat s, and so need t o consume more food in order t o maint ain
homeot hermy.[101]

Bat s may avoid flying during t he day t o prevent overheat ing in t he sun, since t heir dark wing-
membranes absorb solar radiat ion. Bat s may not be able t o dissipat e heat if t he ambient
t emperat ure is t oo high;[102] t hey use saliva t o cool t hemselves in ext reme condit ions.[49] Among
megabat s, t he flying fox Pteropus hypomelanus uses saliva and wing-fanning t o cool it self while
roost ing during t he hot t est part of t he day.[103] Among microbat s, t he Yuma myot is (Myotis
yumanensis), t he Mexican free-t ailed bat , and t he pallid bat (Antrozous pallidus) cope wit h
t emperat ures up t o 45 °C (113 °F) by pant ing, salivat ing, and licking t heir fur t o promot e
evaporat ive cooling; t his is sufficient t o dissipat e t wice t heir met abolic heat product ion.[104]

Bat s also possess a syst em of sphinct er valves on t he art erial side of t he vascular net work t hat
runs along t he edge of t heir wings. When fully open, t hese allow oxygenat ed blood t o flow
t hrough t he capillary net work across t he wing membrane; when cont ract ed, t hey shunt flow
direct ly t o t he veins, bypassing t he wing capillaries. This allows bat s t o cont rol how much heat is
exchanged t hrough t he flight membrane, allowing t hem t o release heat during flight . Many ot her
mammals use t he capillary net work in oversized ears for t he same purpose.[105]

Torpor
A tricoloured bat (Perimyotis subflavus) in torpor

Torpor, a st at e of decreased act ivit y where t he body t emperat ure and met abolism decreases, is
especially useful for microbat s, as t hey use a large amount of energy while act ive, depend upon
an unreliable food source, and have a limit ed abilit y t o st ore fat . They generally drop t heir body
t emperat ure in t his st at e t o 6–30 °C (43–86 °F), and may reduce t heir energy expendit ure by 50
t o 99%. Around 97% of all microbat s use t orpor.[106] Tropical bat s may use it t o avoid predat ion,
by reducing t he amount of t ime spent on foraging and t hus reducing t he chance of being caught
by a predat or.[107] Megabat s were generally believed t o be homeot hermic, but t hree species of
small megabat s, wit h a mass of about 50 grams (13⁄4 ounces), have been known t o use t orpor:
t he common blossom bat (Syconycteris australis), t he long-t ongued nect ar bat (Macroglossus
minimus), and t he east ern t ube-nosed bat (Nyctimene robinsoni). Torpid st at es last longer in
t he summer for megabat s t han in t he wint er.[108]

During hibernat ion, bat s ent er a t orpid st at e and decrease t heir body t emperat ure for 99.6% of
t heir hibernat ion period; even during periods of arousal, when t hey ret urn t heir body t emperat ure
t o normal, t hey somet imes ent er a shallow t orpid st at e, known as "het erot hermic arousal".[109]
Some bat s become dormant during higher t emperat ures t o keep cool in t he summer mont hs.[110]

Het erot hermic bat s during long migrat ions may fly at night and go int o a t orpid st at e roost ing in
t he dayt ime. Unlike migrat ory birds, which fly during t he day and feed during t he night , noct urnal
bat s have a conflict bet ween t ravelling and eat ing. The energy saved reduces t heir need t o feed,
and also decreases t he durat ion of migrat ion, which may prevent t hem from spending t oo much
t ime in unfamiliar places, and decrease predat ion. In some species, pregnant individuals may not
use t orpor.[111][112]

Size

The smallest bat is Kit t i's hog-nosed bat (Craseonycteris thonglongyai), which is 29–34 mm
(11⁄8–13⁄8 in) long wit h a 150-millimet re (6 in) wingspan and weighs 2–2.6 g
(1⁄16–3⁄32 oz).[113][114] It is also arguably t he smallest ext ant species of mammal, next t o t he
Et ruscan shrew.[115] The largest bat s are a few species of Pteropus megabat s and t he giant
golden-crowned flying fox, (Acerodon jubatus), which can weigh 1.6 kg (31⁄2 lb) wit h a wingspan
of 1.7 m (5 ft 7 in).[116] Larger bat s t end t o use lower frequencies and smaller bat s higher for
echolocat ion; high-frequency echolocat ion is bet t er at det ect ing smaller prey. Small prey may be
absent in t he diet s of large bat s as t hey are unable t o det ect t hem.[117] The adapt at ions of a
part icular bat species can direct ly influence what kinds of prey are available t o it .[118]

Ecology

Tent-making bats (Uroderma bilobatum) in Costa Rica

Flight has enabled bat s t o become one of t he most widely dist ribut ed groups of mammals.[119]
Apart from t he Arct ic, t he Ant arct ic and a few isolat ed oceanic islands, bat s exist in almost
every habit at on Eart h.[120] Tropical areas t end t o have more species t han t emperat e ones.[121]
Different species select different habit at s during different seasons, ranging from seasides t o
mount ains and desert s, but t hey require suit able roost s. Bat roost s can be found in hollows,
crevices, foliage, and even human-made st ruct ures, and include "t ent s" t he bat s const ruct wit h
leaves.[122] Megabat s generally roost in t rees.[123] Most microbat s are noct urnal[124] and
megabat s are t ypically diurnal or crepuscular.[125][126] Microbat s are known t o exhibit diurnal
behaviour in t emperat e regions during summer when t here is insufficient night t ime t o
forage,[127][128] and in areas where t here are few avian predat ors during t he day.[129][130]

In t emperat e areas, some microbat s migrat e hundreds of kilomet res t o wint er hibernat ion
dens;[131] ot hers pass int o t orpor in cold weat her, rousing and feeding when warm weat her allows
insect s t o be act ive.[132] Ot hers ret reat t o caves for wint er and hibernat e for as much as six
mont hs.[132] Microbat s rarely fly in rain; it int erferes wit h t heir echolocat ion, and t hey are unable
t o hunt .[133]

Food and feeding

Different bat species have different diet s, including insect s, nect ar, pollen, fruit and even
vert ebrat es.[134] Megabat s are most ly fruit , nect ar and pollen eat ers.[125] Due t o t heir small size,
high-met abolism and rapid burning of energy t hrough flight , bat s must consume large amount s of
food for t heir size. Insect ivorous bat s may eat over 120 percent of t heir body weight , while
frugivorous bat s may eat over t wice t heir weight .[135] They can t ravel significant dist ances each
night , except ionally as much as 38.5 km (24 mi) in t he spot t ed bat (Euderma maculatum), in
search of food.[136] Bat s use a variet y of hunt ing st rat egies.[117] Bat s get most of t heir wat er
from t he food t hey eat ; many species also drink from wat er sources like lakes and st reams, flying
over t he surface and dipping t heir t ongues int o t he wat er.[137]

The Chiropt era as a whole are in t he process of losing t he abilit y t o synt hesise vit amin C.[138] In a
t est of 34 bat species from six major families, including major insect - and fruit -eat ing bat
families, all were found t o have lost t he abilit y t o synt hesise it , and t his loss may derive from a
common bat ancest or, as a single mut at ion.[139][b] At least t wo species of bat , t he frugivorous
bat (Rousettus leschenaultii) and t he insect ivorous bat (Hipposideros armiger), have ret ained
t heir abilit y t o produce vit amin C.[140]

Insect s

Most microbat s, especially in t emperat e areas, prey on insect s.[134] The diet of an insect ivorous
bat may span many species,[141] including flies, mosquit os, beet les, mot hs, grasshoppers,
cricket s, t ermit es, bees, wasps, mayflies and caddisflies.[40][142][143] Large numbers of Mexican
free-t ailed bat s (Tadarida brasiliensis) fly hundreds of met res above t he ground in cent ral Texas
t o feed on migrat ing mot hs.[144] Species t hat hunt insect s in flight , like t he lit t le brown bat
(Myotis lucifugus), may cat ch an insect in mid-air wit h t he mout h, and eat it in t he air or use t heir
t ail membranes or wings t o scoop up t he insect and carry it t o t he mout h.[145][146] The bat may
also t ake t he insect back t o it s roost and eat it t here.[147] Slower moving bat species, such as
t he brown long-eared bat (Plecotus auritus) and many horseshoe bat species, may t ake or glean
insect s from veget at ion or hunt t hem from perches.[40] Insect ivorous bat s living at high lat it udes
have t o consume prey wit h higher energet ic value t han t ropical bat s.[148]

Fruit and nect ar

An Egyptian fruit bat (Rousettus aegyptiacus) carrying a fig

Fruit eat ing, or frugivory, is found in bot h major suborders. Bat s prefer ripe fruit , pulling it off t he
t rees wit h t heir t eet h. They fly back t o t heir roost s t o eat t he fruit , sucking out t he juice and
spit t ing t he seeds and pulp out ont o t he ground. This helps disperse t he seeds of t hese fruit
t rees, which may t ake root and grow where t he bat s have left t hem, and many species of plant s
depend on bat s for seed dispersal.[149][150] The Jamaican fruit bat (Artibeus jamaicensis) has
been recorded carrying fruit s weighing 3–14 g (1⁄8–1⁄2 oz) or even as much as 50 g (13⁄4 oz).[151]

Nect ar-eat ing bat s have acquired specialised adapt at ions. These bat s possess long muzzles and
long, ext ensible t ongues covered in fine brist les t hat aid t hem in feeding on part icular flowers
and plant s.[150][152] The t ube-lipped nect ar bat (Anoura fistulata) has t he longest t ongue of any
mammal relat ive t o it s body size. This is beneficial t o t hem in t erms of pollinat ion and feeding.
Their long, narrow t ongues can reach deep int o t he long cup shape of some flowers. When t he
t ongue ret ract s, it coils up inside t he rib cage.[152] Because of t hese feat ures, nect ar-feeding
bat s cannot easily t urn t o ot her food sources in t imes of scarcit y, making t hem more prone t o
ext inct ion t han ot her t ypes of bat .[153][154] Nect ar feeding also aids a variet y of plant s, since
t hese bat s serve as pollinat ors, as pollen get s at t ached t o t heir fur while t hey are feeding.
Around 500 species of flowering plant rely on bat pollinat ion and t hus t end t o open t heir flowers
at night .[150] Many rainforest plant s depend on bat pollinat ion.[155]

Vert ebrat es

The greater noctule bat (Nyctalus lasiopterus) uses its large teeth to catch birds.[156]

Some bat s prey on ot her vert ebrat es, such as fish, frogs, lizards, birds and mammals.[40][157] The
fringe-lipped bat (Trachops cirrhosus,) for example, is skilled at cat ching frogs. These bat s
locat e large groups of frogs by t racking t heir mat ing calls, t hen plucking t hem from t he surface
of t he wat er wit h t heir sharp canine t eet h.[158] The great er noct ule bat can cat ch birds in
flight .[156] Some species, like t he great er bulldog bat (Noctilio leporinus) hunt fish. They use
echolocat ion t o det ect small ripples on t he wat er's surface, swoop down and use specially
enlarged claws on t heir hind feet t o grab t he fish, t hen t ake t heir prey t o a feeding roost and
consume it .[159] At least t wo species of bat are known t o feed on ot her bat s: t he spect ral bat
(Vampyrum spectrum), and t he ghost bat (Macroderma gigas).[160]

Blood
The common vampire bat (Desmodus rotundus) feeds on blood (hematophagy).

A few species, specifically t he common, whit e-winged, and hairy-legged vampire bat s, feed only
on animal blood (hemat ophagy). The common vampire bat t ypically feeds on large mammals such
as cat t le; t he hairy-legged and whit e-winged vampires feed on birds.[161] Vampire bat s t arget
sleeping prey and can det ect deep breat hing.[162] Heat sensors in t he nose help t hem t o det ect
blood vessels near t he surface of t he skin.[163] They pierce t he animal's skin wit h t heir t eet h,
bit ing away a small flap,[164] and lap up t he blood wit h t heir t ongues, which have lat eral grooves
adapt ed t o t his purpose.[165] The blood is kept from clot t ing by an ant icoagulant in t he saliva.[164]

Predators, parasites, and diseases

Bat s are subject t o predat ion from birds of prey, such as owls, hawks, and falcons, and at roost s
from t errest rial predat ors able t o climb, such as cat s.[166] Low-flying bat s are vulnerable t o
crocodiles.[167] Twent y species of t ropical New World snakes are known t o capt ure bat s, oft en
wait ing at t he ent rances of refuges, such as caves, for bat s t o fly past .[168] J. Rydell and J. R.
Speakman argue t hat bat s evolved noct urnalit y during t he early and middle Eocene period t o
avoid predat ors.[166] The evidence is t hought by some zoologist s t o be equivocal so far.[169]
A little brown bat with white nose syndrome

As most mammals, bat s are host s t o a number of int ernal and ext ernal parasit es.[170] Among
ect oparasit es, bat s carry fleas and mit es, as well as specific parasit es such as bat bugs and bat
flies (Nyct eribiidae and St reblidae).[171][172] Bat s are among t he few non-aquat ic mammalian
orders t hat do not host lice, possibly due t o compet it ion from more specialised parasit es t hat
occupy t he same niche.[172]

Whit e nose syndrome is a condit ion associat ed wit h t he deat hs of millions of bat s in t he East ern
Unit ed St at es and Canada.[173] The disease is named aft er a whit e fungus, Pseudogymnoascus
destructans, found growing on t he muzzles, ears, and wings of afflict ed bat s. The fungus is
most ly spread from bat t o bat , and causes t he disease.[174] The fungus was first discovered in
cent ral New York St at e in 2006 and spread quickly t o t he ent ire East ern US nort h of Florida;
mort alit y rat es of 90–100% have been observed in most affect ed caves.[175] New England and
t he mid-At lant ic st at es have, since 2006, wit nessed ent ire species complet ely ext irpat ed and
ot hers wit h numbers t hat have gone from t he hundreds of t housands, even millions, t o a few
hundred or less.[176] Nova Scot ia, Quebec, Ont ario, and New Brunswick have wit nessed ident ical
die offs, wit h t he Canadian government making preparat ions t o prot ect all remaining bat
populat ions in it s t errit ory.[177] Scient ific evidence suggest s t hat longer wint ers where t he fungus
has a longer period t o infect bat s result in great er mort alit y.[178][179][180] In 2014, t he infect ion
crossed t he Mississippi River,[181] and in 2017, it was found on bat s in Texas.[182]
Bat s are nat ural reservoirs for a large number of zoonot ic pat hogens,[183] including rabies,
endemic in many bat populat ions,[184][185][186] hist oplasmosis bot h direct ly and in guano,[187] Nipah
and Hendra viruses,[188][189] and possibly t he ebola virus,[190][191] whose nat ural reservoir is yet
unknown.[192][193] Their high mobilit y, broad dist ribut ion, long life spans, subst ant ial sympat ry
(range overlap) of species, and social behaviour make bat s favourable host s and vect ors of
disease.[194] Reviews have found different answers as t o whet her bat s have more zoonot ic
viruses t han ot her mammal groups. One 2015 review found t hat bat s, rodent s, and primat es all
harbored significant ly more zoonot ic viruses (which can be t ransmit t ed t o humans) t han ot her
mammal groups, t hough t he differences among t he aforement ioned t hree groups were not
significant (bat s have no more zoonot ic viruses t han rodent s and primat es).[195] Anot her 2020
review of mammals and birds found t hat t he ident ify of t he t axonomic groups did not have any
impact on t he probabilit y of harboring zoonot ic viruses. Inst ead, more diverse groups had great er
viral diversit y.[196]

They seem t o be highly resist ant t o many of t he pat hogens t hey carry, suggest ing a degree of
adapt at ion t o t heir immune syst ems.[194][197][198] Their int eract ions wit h livest ock and pet s,
including predat ion by vampire bat s, accident al encount ers, and t he scavenging of bat carcasses,
compound t he risk of zoonot ic t ransmission.[185] Bat s are implicat ed in t he emergence of severe
acut e respirat ory syndrome (SARS) in China, since t hey serve as nat ural host s for coronaviruses,
several from a single cave in Yunnan, one of which developed int o t he SARS virus.[187][199][200]
However, t hey neit her cause nor spread COVID-19.[201]

Behaviour and life history

Social structure
 Play media

Bracken Bat Cave, home to twenty million Mexican free-tailed bats

Some bat s lead solit ary lives, while ot hers live in colonies of more t han a million.[202] For inst ance,
t he Mexican free-t ailed bat fly for more t han one t housand miles t o t he 100-foot (30 m) wide
cave known as Bracken Cave every March t o Oct ober which plays home t o an ast onishing t went y
million of t he species,[203] whereas a mouse-eared bat lives an almost complet ely solit ary
life.[204] Living in large colonies lessens t he risk t o an individual of predat ion.[40] Temperat e bat
species may swarm at hibernat ion sit es as aut umn approaches. This may serve t o int roduce
young t o hibernat ion sit es, signal reproduct ion in adult s and allow adult s t o breed wit h t hose
from ot her groups.[205]

Several species have a fission-fusion social st ruct ure, where large numbers of bat s congregat e in
one roost ing area, along wit h breaking up and mixing of subgroups. Wit hin t hese societ ies, bat s
are able t o maint ain long-t erm relat ionships.[206] Some of t hese relat ionships consist of
mat rilineally relat ed females and t heir dependent offspring.[207] Food sharing and mut ual
grooming may occur in cert ain species, such as t he common vampire bat (Desmodus rotundus),
and t hese st rengt hen social bonds.[208][209]

Communication
Acoustics of the songs of Mexican free-tailed bats[210]

Bat s are among t he most vocal of mammals and produce calls t o at t ract mat es, find roost
part ners and defend resources. These calls are t ypically low-frequency and can t ravel long
dist ances.[40][211] Mexican free-t ailed bat s are one of t he few species t o "sing" like birds. Males
sing t o at t ract females. Songs have t hree phrases: chirps, t rills and buzzes, t he former having "A"
and "B" syllables. Bat songs are highly st ereot ypical but wit h variat ion in syllable number, phrase
order, and phrase repet it ions bet ween individuals.[210] Among great er spear-nosed bat s
(Phyllostomus hastatus), females produce loud, broadband calls among t heir roost mat es t o
form group cohesion. Calls differ bet ween roost ing groups and may arise from vocal learning.[212]

In a st udy on capt ive Egypt ian fruit bat s, 70% of t he direct ed calls could be ident ified by t he
researchers as t o which individual bat made it , and 60% could be cat egorised int o four cont ext s:
squabbling over food, jost ling over posit ion in t heir sleeping clust er, prot est ing over mat ing
at t empt s and arguing when perched in close proximit y t o each ot her. The animals made slight ly
different sounds when communicat ing wit h different individual bat s, especially t hose of t he
opposit e sex.[213] In t he highly sexually dimorphic hammer-headed bat (Hypsignathus
monstrosus), males produce deep, resonat ing, monot onous calls t o at t ract females. Bat s in
flight make vocal signals for t raffic cont rol. Great er bulldog bat s honk when on a collision course
wit h each ot her.[211]

Bat s also communicat e by ot her means. Male lit t le yellow-shouldered bat s (Sturnira lilium) have
shoulder glands t hat produce a spicy odour during t he breeding season. Like many ot her species,
t hey have hair specialised for ret aining and dispersing secret ions. Such hair forms a conspicuous
collar around t he necks of t he some Old World megabat males. Male great er sac-winged bat s
(Saccopteryx bilineata) have sacs in t heir wings in which t hey mix body secret ions like saliva and
urine t o creat e a perfume t hat t hey sprinkle on roost sit es, a behaviour known as "salt ing". Salt ing
may be accompanied by singing.[211]

Reproduction and lifecycle

Group of polygynous vampire bats

Most bat species are polygynous, where males mat e wit h mult iple females. Male pipist relle,
noct ule and vampire bat s may claim and defend resources t hat at t ract females, such as roost
sit es, and mat e wit h t hose females. Males unable t o claim a sit e are forced t o live on t he
periphery where t hey have less reproduct ive success.[214][40] Promiscuit y, where bot h sexes
mat e wit h mult iple part ners, exist s in species like t he Mexican free-t ailed bat and t he lit t le
brown bat .[215][216] There appears t o be bias t owards cert ain males among females in t hese
bat s.[40] In a few species, such as t he yellow-winged bat and spect ral bat , adult males and
females form monogamous pairs.[40][217] Lek mat ing, where males aggregat e and compet e for
female choice t hrough display, is rare in bat s[218] but occurs in t he hammerheaded bat .[219]

For t emperat e living bat s, mat ing t akes place in lat e summer and early aut umn.[220] Tropical bat s
may mat e during t he dry season.[221] Aft er copulat ion, t he male may leave behind a mat ing plug
t o block t he sperm of ot her males and t hus ensure his pat ernit y.[222] In hibernat ing species, males
are known t o mat e wit h females in t orpor.[40] Female bat s use a variet y of st rat egies t o cont rol
t he t iming of pregnancy and t he birt h of young, t o make delivery coincide wit h maximum food
abilit y and ot her ecological fact ors. Females of some species have delayed fert ilisat ion, in which
sperm is st ored in t he reproduct ive t ract for several mont hs aft er mat ing. Mat ing occurs in lat e
summer t o early aut umn but fert ilisat ion does not occur unt il t he following lat e wint er t o early
spring. Ot her species exhibit delayed implant at ion, in which t he egg is fert ilised aft er mat ing, but
remains free in t he reproduct ive t ract unt il ext ernal condit ions become favourable for giving birt h
and caring for t he offspring.[223] In anot her st rat egy, fert ilisat ion and implant at ion bot h occur, but
development of t he foet us is delayed unt il good condit ions prevail. During t he delayed
development t he mot her keeps t he fert ilised egg alive wit h nut rient s. This process can go on for
a long period, because of t he advanced gas exchange syst em.[224]

Newborn common pipistrelle, Pipistrellus pipistrellus

For t emperat e living bat s, birt hs t ypically t ake place in May or June in t he nort hern hemisphere;
birt hs in t he sout hern hemisphere occur in November and December. Tropical species give birt h
at t he beginning of t he rainy season.[225] In most bat species, females carry and give birt h t o a
single pup per lit t er.[226] At birt h, a bat pup can be up t o 40 percent of t he mot her's weight ,[40]
and t he pelvic girdle of t he female can expand during birt h as t he t wo-halves are connect ed by a
flexible ligament .[227] Females t ypically give birt h in a head-up or horizont al posit ion, using gravit y
t o make birt hing easier. The young emerges rear-first , possibly t o prevent t he wings from get t ing
t angled, and t he female cradles it in her wing and t ail membranes. In many species, females give
birt h and raise t heir young in mat ernit y colonies and may assist each ot her in birt hing.[228][229][227]

Most of t he care for a young bat comes from t he mot her. In monogamous species, t he fat her
plays a role. Allo-suckling, where a female suckles anot her mot her's young, occurs in several
species. This may serve t o increase colony size in species where females ret urn t o t heir nat al
colony t o breed.[40] A young bat 's abilit y t o fly coincides wit h t he development of an adult body
and forelimb lengt h. For t he lit t le brown bat , t his occurs about eight een days aft er birt h. Weaning
of young for most species t akes place in under eight y days. The common vampire bat nurses it s
offspring beyond t hat and young vampire bat s achieve independence lat er in life t han ot her
species. This is probably due t o t he species' blood-based diet , which is difficult t o obt ain on a
night ly basis.[230]

Life expectancy

The bat scientist Lauri Lutsar is checking the age of the bat he is holding as part of a national monitoring program in
Estonia

The maximum lifespan of bat s is t hree-and-a-half t imes longer t han ot her mammals of similar
size. Six species have been recorded t o live over t hirt y years in t he wild: t he brown long-eared
bat (Plecotus auritus), t he lit t le brown bat (Myotis lucifugus), t he Siberian bat (Myotis sibiricus),
t he lesser mouse-eared bat (Myotis blythii) t he great er horseshoe bat (Rhinolophus
ferrumequinum), and t he Indian flying fox (Pteropus giganteus).[231] One hypot hesis consist ent
wit h t he rat e-of-living t heory links t his t o t he fact t hat t hey slow down t heir met abolic rat e
while hibernat ing; bat s t hat hibernat e, on average, have a longer lifespan t han bat s t hat do
not .[232][233]

Anot her hypot hesis is t hat flying has reduced t heir mort alit y rat e, which would also be t rue for
birds and gliding mammals. Bat species t hat give birt h t o mult iple pups generally have a short er
lifespan t han species t hat give birt h t o only a single pup. Cave-roost ing species may have a
longer lifespan t han non-roost ing species because of t he decreased predat ion in caves. A male
Siberian bat was recapt ured in t he wild aft er 41 years, making it t he oldest known bat .[233][234]
Interactions with humans

Conservation

Conservation statuses of bats as of 2020 according to the IUCN (1,314 species in total)[235]

   Critically endangered (1.6%)

   Endangered (6.3%)
   Vulnerable (8.3%)
   Near-threatened (6.7%)
   Least concern (58.0%)
   Data deficient (18.4%)
   Extinct (0.7%)

Groups such as t he Bat Conservat ion Int ernat ional[236] aim t o increase awareness of bat s'
ecological roles and t he environment al t hreat s t hey face. In t he Unit ed Kingdom, all bat s are
prot ect ed under t he Wildlife and Count ryside Act s, and dist urbing a bat or it s roost can be
punished wit h a heavy fine.[237] In Sarawak, Malaysia, "all bat s"[238] are prot ect ed under t he
Wildlife Prot ect ion Ordinance 1998,[238] but species such as t he hairless bat (Cheiromeles
torquatus) are st ill eat en by t he local communit ies.[239] Humans have caused t he ext inct ion of
several species of bat in modern hist ory, t he most recent being t he Christ mas Island pipist relle
(Pipistrellus murrayi), which was declared ext inct in 2009.[240]
Many people put up bat houses t o at t ract bat s.[241] The 1991 Universit y of Florida bat house is
t he largest occupied art ificial roost in t he world, wit h around 400,000 resident s.[242] In Brit ain,
t hickwalled and part ly underground World War II pillboxes have been convert ed t o make roost s
for bat s,[243][244] and purpose-built bat houses are occasionally built t o mit igat e damage t o
habit at from road or ot her development s.[245][246] Cave gat es are somet imes inst alled t o limit
human ent ry int o caves wit h sensit ive or endangered bat species. The gat es are designed not t o
limit t he airflow, and t hus t o maint ain t he cave's micro-ecosyst em.[247] Of t he 47 species of bat s
found in t he Unit ed St at es, 35 are known t o use human st ruct ures, including buildings and bridges.
Fourt een species use bat houses.[248]

Bat s are eat en in count ries across Africa, Asia and t he Pacific Rim. In some cases, such as in
Guam, flying foxes have become endangered t hrough being hunt ed for food.[249] There is
evidence t hat wind t urbines creat e sufficient barot rauma (pressure damage) t o kill bat s.[250] Bat s
have t ypical mammalian lungs, which are t hought t o be more sensit ive t o sudden air pressure
changes t han t he lungs of birds, making t hem more liable t o fat al rupt ure.[251][252][253][254][255]
Bat s may be at t ract ed t o t urbines, perhaps seeking roost s, increasing t he deat h rat e.[251]
Acoust ic det errent s may help t o reduce bat mort alit y at wind farms.[256]

Cultural significance
Francisco Goya, The Sleep of Reason Produces Monsters, 1797

Since bat s are mammals, yet can fly, t hey are considered t o be liminal beings in various
t radit ions.[257] In many cult ures, including in Europe, bat s are associat ed wit h darkness, deat h,
wit chcraft , and malevolence.[258] Among Nat ive Americans such as t he Creek, Cherokee and
Apache, t he bat is ident ified as a t rickst er.[259] In Tanzania, a winged bat like creat ure known as
Popobawa is believed t o be a shapeshift ing evil spirit t hat assault s and sodomises it s
vict ims.[260] In Azt ec myt hology, bat s symbolised t he land of t he dead, dest ruct ion, and
decay.[261][262][263] An East Nigerian t ale t ells t hat t he bat developed it s noct urnal habit s aft er
causing t he deat h of his part ner, t he bush-rat , and now hides by day t o avoid arrest .[264]

More posit ive depict ions of bat s exist in some cult ures. In China, bat s have been associat ed wit h
happiness, joy and good fort une. Five bat s are used t o symbolise t he "Five Blessings": longevit y,
wealt h, healt h, love of virt ue and peaceful deat h.[265] The bat is sacred in Tonga and is oft en
considered t he physical manifest at ion of a separable soul.[266] In t he Zapot ec civilisat ion of
Mesoamerica, t he bat god presided over corn and fert ilit y.[267]

Zapotec bat god, Oaxaca, 350–500 CE

The Weird Sist ers in Shakespeare's Macbeth used t he fur of a bat in t heir brew.[268] In West ern
cult ure, t he bat is oft en a symbol of t he night and it s foreboding nat ure. The bat is a primary
animal associat ed wit h fict ional charact ers of t he night , bot h villainous vampires, such as Count
Dracula and before him Varney the Vampire,[269] and heroes, such as t he DC Comics charact er
Bat man.[270] Kennet h Oppel's Silverwing novels narrat e t he advent ures of a young bat ,[271] based
on t he silver-haired bat of Nort h America.[272]

The bat is somet imes used as a heraldic symbol in Spain and France, appearing in t he coat s of
arms of t he t owns of Valencia, Palma de Mallorca, Fraga, Albacet e, and Mont chauvet .[273][274][275]
Three US st at es have an official st at e bat . Texas and Oklahoma are represent ed by t he Mexican
free-t ailed bat , while Virginia is represent ed by t he Virginia big-eared bat (Corynorhinus
townsendii virginianus).[276]

Economics

Insect ivorous bat s in part icular are especially helpful t o farmers, as t hey cont rol populat ions of
agricult ural pest s and reduce t he need t o use pest icides. It has been est imat ed t hat bat s save
t he agricult ural indust ry of t he Unit ed St at es anywhere from $3.7 billion t o $53 billion per year in
pest icides and damage t o crops. This also prevent s t he overuse of pest icides, which can pollut e
t he surrounding environment , and may lead t o resist ance in fut ure generat ions of insect s.[277]

Bat dung, a t ype of guano, is rich in nit rat es and is mined from caves for use as fert iliser.[278]
During t he US Civil War, salt pet re was collect ed from caves t o make gunpowder. At t he t ime, it
was believed t hat t he nit rat e all came from t he bat guano, but it is now known t hat most of it is
produced by nit rifying bact eria.[279]

The Congress Avenue Bridge in Aust in, Texas, is t he summer home t o Nort h America's largest
urban bat colony, an est imat ed 1,500,000 Mexican free-t ailed bat s. About 100,000 t ourist s a year
visit t he bridge at t wilight t o wat ch t he bat s leave t he roost .[280]

See also

Bat det ect or

Explanatory notes

a. Pronounced /kaɪˈrɒptərə/; from the Ancient Greek: χείρ – cheir, "hand" and πτερόν – pteron, "wing".[1]

b. Earlier reports that only fruit bats were deficient were based on smaller samples.[140]

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External links

Wikimedia Commons has media relat ed t o Chiroptera.

The Wikibook Dichotomous Key has a page on t he t opic of: Chiroptera

Wikispecies has informat ion relat ed t o Chiroptera.

UK Bat Conservat ion Trust (ht t p://www.bat s.org.uk/pages/about _ bat s.ht ml)

Tree of Life (ht t p://t olweb.org/t ree?group=Chiropt era&cont group=Eut heria)

Microbat Vision (ht t p://www.fladdermus.net /t hesis.ht m)

Analyses of several kinds of bat echolocat ion (ht t p://www.hscot t .net /t he-dsp-behind-bat -ech
olocat ion/)

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