Bats are mammals of the order Chiroptera;[a] with their forelimbs adapted as wings, they are the

only mammals naturally capable of true and sustained flight. Bats are more manoeuvrable
than birds, flying with their very long spread-out digits covered with a thin membrane or patagium.
The smallest bat, and arguably the smallest extant mammal, is Kitti's hog-nosed bat, which is 29–
34 mm (1.14–1.34 in) in length, 15 cm (5.91 in) across the wings and 2–2.6 g (0.07–0.09 oz)
in mass. The largest bats are the flying foxes and the giant golden-crowned flying fox, Acerodon
jubatus, which can weigh 1.6 kg (4 lb) and have a wingspan of 1.7 m (5 ft 7 in).
The second largest order of mammals, bats comprise about 20% of all classified mammal
species worldwide, with over 1,200 species. These were traditionally divided into two
suborders: the largely fruit-eating megabats, and the echolocating microbats. But more recent
evidence has supported dividing the order into Yinpterochiroptera and Yangochiroptera, with
megabats as members of the former along with several species of microbats. Many bats
are insectivores, and most of the rest are frugivores (fruit-eaters). A few species feed on animals
other than insects; for example, the vampire bats feed on blood. Most bats are nocturnal, and many
roost in caves or other refuges; it is uncertain whether bats have these behaviours to
escape predators. Bats are present throughout the world, with the exception of extremely cold
regions. They are important in their ecosystems for pollinating flowers and dispersing seeds;
many tropical plants depend entirely on bats for these services.

Bats provide humans with some benefits, at the cost of some threats. Bat dung has been
mined as guano from caves and used as fertiliser. Bats consume insect pests, reducing the
need for pesticides. They are sometimes numerous enough to serve as tourist attractions, and
are used as food across Asia and the Pacific Rim. They are natural reservoirs of many pathogens,
such as rabies; and since they are highly mobile, social, and long-lived, they can readily spread
disease. In many cultures, bats are popularly associated with darkness, malevolence,
witchcraft, vampires, and death.

An older English name for bats is flittermouse, which matches their name in other Germanic
languages (for example German Fledermaus and Swedish fladdermus), related to the fluttering
of wings. Middle English had bakke, most likely cognate with Old Swedish natbakka ("night-
bat"), which may have undergone a shift from -k- to -t- (to Modern English bat) influenced by
Latin blatta, "moth, nocturnal insect". The word "bat" was probably first used in the early
1570s.[2][3] The name "Chiroptera" 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 delicate skeletons of bats do not fossilise well, and it is estimated that only 12% of bat
genera that lived have been found in the fossil record. [6] Most of the oldest known bat fossils
were already very similar to modern microbats, such as Archaeopteropus (32 million years
ago).[7] The extinct bats Palaeochiropteryx tupaiodon (48 million years ago) and Hassianycteris
kumari (55 million years ago) are the first fossil mammals whose colouration has been
discovered: both were reddish-brown.[8][9]
Bats were formerly grouped in the superorder Archonta, along with
the treeshrews (Scandentia), colugos (Dermoptera), and primates.[10] Modern genetic evidence now
places bats in the superorder Laurasiatheria, with its sister taxon as Fereuungulata, which
includes carnivorans, pangolins, odd-toed ungulates, even-toed ungulates, and cetaceans.[11][12][13][14][15] One
study places Chiroptera as a sister taxon to odd-toed ungulates (Perissodactyla). [16]

Giant golden-crowned flying fox, Acerodon jubatus

In the 1980s, a hypothesis based on morphological evidence stated the Megachiroptera evolved
flight separately from the Microchiroptera. The flying primate hypothesis proposed that, when
adaptations to flight are removed, the Megachiroptera are allied to primates by anatomical
features not shared with Microchiroptera. For example, the brains of megabats have advanced
characteristics. Although recent genetic studies strongly support the monophyly of bats,
[7] debate continues about the meaning of the genetic and morphological evidence. [19]

The 2003 discovery of an early fossil bat from the 52 million year old Green River
Formation, Onychonycteris finneyi, indicates that flight evolved before echolocative abilities.[20]
[21] Onychonycteris had claws on all five of its fingers, whereas modern bats have at most two
claws on two digits of each hand. It also had longer hind legs and shorter forearms, similar to
climbing mammals that hang under branches, such as sloths and gibbons. This palm-sized bat
had short, broad wings, suggesting that it could not fly as fast or as far as later bat species.
Instead of flapping its wings continuously while flying, Onychonycteris probably alternated
between flaps and glides in the air.[7] This suggests that this bat did not fly as much as
modern bats, but flew from tree to tree and spent most of its time climbing or hanging on
branches.[22] The distinctive features of the Onychonycteris fossil also support the hypothesis
that mammalian flight most likely evolved in arboreal locomotors, rather than terrestrial
runners. This model of flight development, commonly known as the "trees-down" theory,
holds that bats first flew by taking advantage of height and gravity to drop down on to prey,
rather than running fast enough for a ground-level take off.[23][24]
The molecular phylogeny is controversial, as it points to microbats not having a unique common
ancestry, which implies that some seemingly unlikely transformations occurred. The first is that
laryngeal echolocation evolved twice in bats, once in Yangochiroptera and once in the
rhinolophoids.[25] The second is that laryngeal echolocation had a single origin in Chiroptera,
was subsequently lost in the family Pteropodidae (all megabats), and later evolved as a
system of tongue-clicking in the genus Rousettus.[26] Analyses of the sequence of the
vocalization gene FoxP2 were inconclusive on whether laryngeal echolocation was lost in the
pteropodids or gained in the echolocating lineages.[27] Echolocation probably first derived in
bats from communicative calls. The Eocene bats Icaronycteris (52 million years ago)
and Palaeochiropteryx had cranial adaptations suggesting an ability to detect ultrasound. This may
have been used at first mainly to forage on the ground for insects and map out their
surroundings in their gliding phase, or for communicative purposes. After the adaptation of
flight was established, it may have been refined to target flying prey by echolocation. [22] Bats
may have evolved echolocation through a shared common ancestor, in which case it was then
lost in the Old World megabats, only to be regained in the horseshoe bats; or, echolocation
evolved independently in both the Yinpterochiroptera and Yangochiroptera lineages.
[28] Analyses of the hearing gene Prestin seem to favour the idea that echolocation developed
independently at least twice, rather than being lost secondarily in the pteropodids. [29]

Classification
See also: List of bats and List of fruit bats

Bats are placental mammals. After rodents, they are the largest order, making up about 20% of
mammal species.[30] In 1758, Carl Linnaeus classified the seven bat species he knew of in the
genus Vespertilio in the order Primates. Around twenty years later, the German naturalist Johann
Friedrich Blumenbach gave them their own order, Chiroptera.[31] Since then, the number of
described species has risen to over 1,200, traditionally classified as two
suborders: Megachiroptera (megabats), and Microchiroptera (microbats/echolocating bats).[32] Not
all megabats are larger than microbats.[33] Several characteristics distinguish the two groups.
Microbats use echolocation for navigation and finding prey, but megabats apart from those in
the genus Rousettus do not, relying instead on their eyesight.[34] Accordingly, megabats have a
well-developed visual cortex and good visual acuity.[32] Megabats have a claw on the second finger
of the forelimb.[35][36] The external ears of microbats do not close to form a ring; the edges are
separated from each other at the base of the ear.[36] Megabats eat fruit, nectar, or pollen, while
most microbats eat insects; others feed on fruit, nectar, pollen, fish, frogs, small mammals,
or blood.[32]
 "Chiroptera" from Ernst Haeckel's Kunstformen der Natur, 1904

The following classification from Agnarsson and colleagues in 2011 reflects the traditional
division into megabat and microbat suborders.[17]

Order Chiroptera[17]
Suborder Megachiroptera

The finger bones of bats are much more flexible than those of other mammals, owing to their
flattened cross-section and to low levels of calcium near their tips. The elongation of bat digits,
a key feature required for wing development, is due to the upregulation of bone morphogenetic
proteins (Bmps). During embryonic development, the gene controlling Bmp signalling, Bmp2, is
subjected to increased expression in bat forelimbs—resulting in the extension of the manual
digits. This crucial genetic alteration helps create the specialised limbs required for powered
flight. The relative proportion of extant bat forelimb digits compared with those of Eocene
fossil bats have no significant differences, suggesting that bat wing morphology has been
conserved for over 50 million years.[43] During flight, the bones
undergo bending and shearing stress; the bending stresses felt are smaller than in terrestrial
mammals, but the shearing stress is larger. The wing bones of bats have a slightly lower
breaking stress point than those of birds.[44]
As in other mammals, and unlike in birds, the radius is the main component of the forearm.
Bats have five elongated digits, which all radiate around the wrist. The thumb points forward
and supports the leading edge of the wing, and the other digits support the tension held in the
wing membrane. The second and third digits go along the wing tip, allowing the wing to be
pulled forward against aerodynamic drag, without having to be thick as in pterosaur wings. The
fourth and fifth digits go from the wrist to the trailing edge, and repel the bending force caused
by air pushing up against the stiff membrane.[45] Due to their flexible joints, bats are more
manoeuvrable and more dexterous than gliding mammals.[46]

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

The wings of bats are much thinner and consist of more bones than the wings of birds,
allowing bats to manoeuvre more accurately than the latter, and fly with more lift and less
drag.[47] By folding the wings in toward their bodies on the upstroke, they save 35 percent
energy during flight.[48] The membranes are delicate, tearing easily,[49] but can regrow, and
small tears heal quickly.[49][50] The surface of the wings is equipped with touch-sensitive
receptors on small bumps called Merkel cells, also found on human fingertips. These sensitive
areas are different in bats, as each bump has a tiny hair in the centre, making it even more
sensitive and allowing the bat to detect and adapt to changing airflow; the primary use is to
judge the most efficient speed to fly at, and possibly also to avoid stalls.[51] Insectivorous bats
may also use tactile hairs to help perform complex manoeuvres to capture prey in flight. [46]
The patagium is the wing membrane; it is stretched between the arm and finger bones, and
down the side o

Roosting adaptations

Group of megabats roosting

When not flying, bats hang upside down from their feet, a posture known as roosting. [56] The
femurs are attached at the hips in a way that allows them to bend outward and upward in
flight. The ankle joint can flex to allow the trailing edge of the wings to bend downwards. This
does not permit many movements other than hanging or clambering up trees. [45] Most
megabats roost with the head tucked towards the belly, whereas most microbats roost with
the neck curled towards the back. This difference is reflected in the structure of the cervical or
neck vertebrae in the two groups, which are clearly distinct.[56] Tendons allow bats to lock their
feet closed when hanging from a roost. Muscular power is needed to let go, but not to grasp a
perch or when holding on.[57]

Internal systems
Bats have an efficient circulatory system. They seem to make use of particularly strong
venomotion, a rhythmic contraction of venous wall muscles. In most mammals, the walls of the
veins provide mainly passive resistance, maintaining their shape as deoxygenated blood flows
through them, but in bats they appear to actively support blood flow back to the heart with
this pumping action.[58][59] Since their bodies are relatively small and lightweight, bats are not
at risk of blood flow rushing to their heads when roosting.[60]
Bats possess a highly adapted respiratory system to cope with the demands of powered flight, an
energetically taxing activity that requires a large continuous throughput of oxygen. In bats,
the relative alveolar surface area and pulmonary capillary blood volume are larger than in
most other small quadrupedal mammals.[61]Because of the restraints of the mammalian lungs,
bats cannot maintain high-altitude flight.[45]
 The wings are highly vascularized membranes, the larger blood vessels visible against the light. [62]

It takes a lot of energy and an efficient circulatory system to work the flight muscles of bats.
Energy supply to the muscles engaged in flight require about double the amount compared to
the muscles that do not use flight as a means of mammalian locomotion. In parallel to energy
consumption, blood oxygen levels of flying animals are twice as much as those of their
terrestrially locomoting mammals. As the blood supply controls the amount of oxygen
supplied throughout the body, the circulatory system must respond accordingly. Therefore,
compared to a terrestrial mammal of the same relative size, the bat's heart can be up to three
times larger, and pump more blood.[63] Cardiac output is directly derived from heart rate
and stroke volume of the blood;[64] an active microbat can reach a heart rate of 1000 beats per
minute.[65]
With its extremely thin membranous tissue, a bat's wing can significantly contribute to the
organism's total gas exchange efficiency.[53]Because of the high energy demand of flight, the
bat's body meets those demands by exchanging gas through the patagium of the wing. When
the bat has its wings spread it allows for an increase in surface area to volume ratio. The
surface area of the wings is about 85% of the total body surface area, suggesting the
possibility of a useful degree of gas exchange.[53] The subc