Chiroptera

Cladus: Eukaryota
Supergroup: Opisthokonta
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Superclassis: Tetrapoda
Classis: Mammalia
Subclassis: Theria
Infraclassis: Eutheria
Ordo: Chiroptera
Subordines: Megachiroptera - Microchiroptera

Name

Chiroptera, Blumenbach, 1779

References

* Chiroptera on Mammal species of the World.
* Mammal Species of the World: A Taxonomic and Geographic Reference, 2 Volume Set edited by Don E. Wilson, DeeAnn M. Reeder
* Chiroptera Blumenbach, 1779 Report on ITIS

Vernacular names
Internationalization
Aragonés: Apagacandil
Asturianu: Esperteyu
Български: Прилепи
Català: Ratapinyada
Česky: Letouni
Dansk: Flagermus
Deutsch: Fledertiere
Ελληνικά: Νυχτερίδα
English: Bats
Español: Murciélago
فارسی: خفاش
Français: Chauve-souris
Hawai`i: ‘Ōpe‘ape‘a
עברית: עטלפים
Hrvatski: Netopiri
Bahasa Indonesia: Kelelawar
Ido: Vespertilio
Íslenska: leðurblaka
Italiano: Pipistrelli / Chirotteri
日本語: コウモリ目
Kiswahili: Popo
한국어: 박쥐
Limburgs: Vliermuis
Lietuvių: Šikšnosparniai
Nederlands: Vleermuizen
‪Norsk (bokmål)‬: Flaggermus
Polski: Nietoperze
Português: Morcego
Română: Liliac
Русский: Рукокрылые
Slovenčina: Netopiere
Suomi: Lepakot
Svenska: Fladdermöss
Türkçe: Yarasalar
Українська: Рукокрилі
Vèneto: Babastrel/Babastrìo, Zergnàpola/Signàpola
中文: 翼手目

Bats are flying mammals in the order Chiroptera (pronounced /kaɪˈrɒptərə/). The forelimbs of bats are webbed and developed as wings, making them the only mammals naturally capable of true and sustained flight. By contrast, other mammals said to fly, such as flying squirrels, gliding possums and colugos, glide rather than fly, and can only glide for short distances. Bats do not flap their entire forelimbs, as birds do, but instead flap their spread out digits,[2] which are very long and covered with a thin membrane or patagium. Chiroptera comes from two Greek words, cheir (χείρ) "hand" and pteron (πτερόν) "wing."

There are about 1,100 bat species worldwide, which represent about twenty percent of all classified mammal species.[3] About seventy percent of bats are insectivores. Most of the rest are frugivores, or fruit eaters. A few species such as the Fish-eating Bat feed from animals other than insects, with the vampire bats being the only mammalian parasite species. Bats are present throughout most of the world and perform vital ecological roles such as pollinating flowers and dispersing fruit seeds. Many tropical plant species depend entirely on bats for the distribution of their seeds.

The smallest bat is the Kitti's Hog-nosed Bat, measuring 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,[4][5] The largest species of bat is the Giant Golden-crowned Flying-fox, which is 336–343 mm (13.23–13.50 in) long, has a wingspan of 1.5 m (4 ft 11 in) and weighs approximately 1.1–1.2 kg (2–3 lb).[6]

Classification and evolution
Giant Golden-crowned Flying-fox, Acerodon jubatus.

Bats are mammals. Sometimes they are mistakenly called "flying rodents" or "flying rats", and they can also be mistaken for insects and birds. There are two traditionally recognized suborders of bats:

* Megachiroptera (megabats)
* Microchiroptera (microbats/echolocating bats)

Not all megabats are larger than microbats. The major distinctions between the two suborders are:

* Microbats use echolocation: megabats do not with the exception of Rousettus and relatives.
* Microbats lack the claw at the second toe of the forelimb.
* The ears of microbats do not close to form a ring: the edges are separated from each other at the base of the ear.
* Microbats lack underfur: they are either naked or have guard hairs.

Megabats eat fruit, nectar or pollen while most microbats eat insects; others may feed on the blood of animals, small mammals, fish, frogs, fruit, pollen or nectar. Megabats have a well-developed visual cortex and show good visual acuity, while microbats rely on echolocation for navigation and finding prey.

The phylogenetic relationships of the different groups of bats have been the subject of much debate. The traditional subdivision between Megachiroptera and Microchiroptera reflects the view that these groups of bats have evolved independently of each other for a long time, from a common ancestor that was already capable of flight. This hypothesis recognized differences between microbats and megabats and acknowledged that flight has only evolved once in mammals. Most molecular biological evidence supports the view that bats form a single or monophyletic group.[7]

Researchers have proposed alternate views of chiropteran phylogeny and classification, but more research is needed.

Genetic evidence indicates that megabats originated during the early Eocene and should be placed within the four major lines of microbats.

Consequently, two new suborders based on molecular data have been proposed. The new suborder Yinpterochiroptera includes the Pteropodidae or megabat family as well as the Rhinolophidae, Hipposideridae, Craseonycteridae, Megadermatidae, and Rhinopomatidae families. The new suborder Yangochiroptera includes all the remaining families of bats (all of which use laryngeal echolocation). These two new suborders are strongly supported by statistical tests. Teeling (2005) found 100% bootstrap support in all maximum likelihood analyses for the division of Chiroptera into these two modified suborders. This conclusion is further supported by a fifteen-base pair deletion in BRCA1 and a seven-base pair deletion in PLCB4 present in all Yangochiroptera and absent in all Yinpterochiroptera.[8] The Chiropteran phylogeny based on molecular evidence is controversial because microbat paraphyly implies that one of two seemingly unlikely hypotheses occurred. The first suggests that laryngeal echolocation evolved twice in Chiroptera, once in Yangochiroptera and once in the rhinolophoids.[9][10] The second proposes 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.[11]
Common Pipistrelle, Pipistrellus pipistrellus.

Analyses of the sequence of the "vocalization" gene, FoxP2 was inconclusive of whether laryngeal echolocation was secondarily lost in the pteropodids or independently gained in the echolocating lineages.[12] However, analyses of the "hearing" gene, Prestin seemed to favor the independent gain in echolocating species rather than a secondary loss in the pteropodids.[13]

In addition to Yinpterochiroptera and Yangochiroptera, the names Pteropodiformes and Vespertilioniformes have also been proposed for these suborders.[14][15] Under this new proposed nomenclature, the suborder Pteropodiformes includes all extant bat families more closely related to the genus Pteropus than the genus Vespertilio, while the suborder Vespertilioniformes includes all extant bat families more closely related to the genus Vespertilio than to the genus Pteropus.

In the 1980s, a hypothesis based on morphological evidence was offered that stated that the Megachiroptera evolved flight separately from the Microchiroptera. The so-called flying primates theory proposed that when adaptations to flight are removed, the Megachiroptera are allied to primates by anatomical features that are not shared with Microchiroptera. One example is that the brains of megabats show a number of advanced characteristics that link them to primates. Although recent genetic studies support the monophyly of bats,[16] debate continues as to the meaning of available genetic and morphological evidence.[17]

Little fossil evidence is available to help map the evolution of bats, since their small, delicate skeletons do not fossilize very well. However a Late Cretaceous tooth from South America resembles that of an early Microchiropteran bat. The oldest known definitely identified bat fossils, such as Icaronycteris, Archaeonycteris, Palaeochiropteryx and Hassianycteris, are from the early Eocene period, 52.5 million years ago.[7] These fossil bats were already very similar to modern microbats. Archaeopteropus, formerly classified as the earliest known megachiropteran, is now classified as a microchiropteran.

Bats were formerly grouped in the superorder Archonta along with the treeshrews (Scandentia), colugos (Dermoptera), and the primates, because of the apparent similarities between Megachiroptera and such mammals. Genetic studies have now placed bats in the superorder Laurasiatheria along with carnivorans, pangolins, odd-toed ungulates, even-toed ungulates, and cetaceans.[1]
"Chiroptera" from Ernst Haeckel's Kunstformen der Natur, 1904

The traditional classification of bats is:

* Order Chiroptera
o Suborder Megachiroptera (megabats)
+ Pteropodidae
o Suborder Microchiroptera (microbats)
+ Superfamily Emballonuroidea
# Emballonuridae (Sac-winged or Sheath-tailed bats)
+ Superfamily Molossoidea
# Antrozoidae (Pallid Bat and Van Gelder's Bat)
# Molossidae (Free-tailed bats)
+ Superfamily Nataloidea
# Furipteridae (Smoky bats)
# Myzopodidae (Sucker-footed bats)
# Natalidae (Funnel-eared bats)
# Thyropteridae (Disk-winged bats)
+ Superfamily Noctilionoidea
# Mormoopidae (Ghost-faced or Moustached bats)
# Mystacinidae (New Zealand short-tailed bats)
# Noctilionidae (Bulldog bats or Fisherman bats)
# Phyllostomidae (Leaf-nosed bats)
+ Superfamily Rhinolophoidea
# Megadermatidae (False vampires)
# Nycteridae (Hollow-faced or Slit-faced bats)
# Rhinolophidae (Horseshoe bats)
+ Superfamily Rhinopomatoidea
# Craseonycteridae (Bumblebee Bat or Kitti's Hog-nosed Bat)
# Rhinopomatidae (Mouse-tailed bats)
+ Superfamily Vespertilionoidea
# Vespertilionidae (Vesper bats or Evening bats)

Megabats primarily eat fruit or nectar. In New Guinea, they are likely to have evolved for some time in the absence of microbats. This has resulted in some smaller megabats of the genus Nyctimene becoming (partly) insectivorous to fill the vacant microbat ecological niche. Furthermore, there is some evidence that the fruit bat genus Pteralopex from the Solomon Islands, and its close relative Mirimiri from Fiji, have evolved to fill some niches that were open because there are no nonvolant or non-flying mammals in those islands.
Fossil bats

There are few fossilized remains of bats, as they are terrestrial and light-boned. An Eocene bat, Onychonycteris finneyi, was found in the fifty-two-million-year-old Green River Formation in South Dakota, United States, in 2004.[18] It had characteristics indicating that it could fly, yet the well-preserved skeleton showed that the cochlea of the inner ear lacked development needed to support the greater hearing abilities of modern bats. This provided evidence that flight in bats developed well before echolocation. The team that found the remains of this species, named Onychonycteris finneyi, recognized that it lacked ear and throat features present not only in echolocating bats today, but also in other known prehistoric species. Fossil remains of another Eocene bat, Icaronycteris, were found in 1960.

The appearance and flight movement of bats 52.5 million years ago were different from those of bats today. Onychonycteris had claws on all five of its fingers, whereas modern bats have at most two claws appearing 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 broad, short 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 likely alternated between flaps and glides while in the air. Such physical characteristics suggest that this bat did not fly as much as modern bats do, rather flying from tree to tree and spending most of its waking day climbing or hanging on the branches of trees.[19]
Habitats

Flight has enabled bats to become one of the most widely distributed groups of mammals. Apart from the Arctic, the Antarctic and a few isolated oceanic islands, bats exists all over the world.[20] Bats are found in almost every habitat available on Earth. Different species select different habitat during different seasons — ranging from seasides to mountains and even deserts — but bat habitats have two basic requirements: roosts, where they spend the day or hibernate, and places for foraging. Bat roosts can be found in hollows, crevices, foliage, and even human-made structures; and include "tents" that bats construct by biting leaves. [21]
Anatomy
Skeleton of a Greater Mouse-eared Bat (Myotis myotis).
Scapulae, spine and ribs of Myotis lucifugus (Little Brown Bat).
Echolocation
Spectrogram of Pipistrellus Bat vocalizations. Detail is shown as the pulse duty cycle increases during a close approach to prey. The bat appears to use a hybrid pulse which combines a sharp falling frequency chirp with an extended constant frequency tail. Such a waveform may offer combined benefits of range estimation as well as Doppler shift detection. Spectrogram generated with Fatpigdog's PC based Real Time FFT Spectrum Analyzer.

Pipistrellus Pulses
Play sound
Recording of Pipistrellus bat approaching its prey.
Problems listening to this file? See media help.

Bat echolocation is a perceptual system where ultrasonic sounds are emitted specifically to produce echoes. By comparing the outgoing pulse with the returning echoes the brain and auditory nervous system can produce detailed images of the bat's surroundings. This allows bats to detect, localize and even classify their prey in complete darkness. At 130 decibels in intensity, bat calls are some of the most intense airborne animal sounds.[22]

To clearly distinguish returning information, bats must be able to separate their calls from the echoes they receive. Microbats use two distinct approaches.

1. Low Duty Cycle Echolocation: Bats can separate their calls and returning echos by time. Bats that use this approach time their short calls to finish before echoes return. This is important because these bats contract their middle ear muscles when emitting a call so that they can avoid deafening themselves. The time interval between call and echo allows them to relax these muscles so they can clearly hear the returning echo.[23] The delay of the returning echos provide the bat with the ability to estimate range to their prey.
2. High Duty Cycle Echolocation: Bats emit a continuous call and separate pulse and echo in frequency. The ears of these bats are sharply tuned to a specific frequency range. They emit calls outside of this range to avoid self-deafening. They then receive echoes back at the finely tuned frequency range by taking advantage of the Doppler shift of their motion in flight. The Doppler shift of the returning echos yield information relating to the motion and location of the bat's prey. These bats must deal with changes in the Doppler shift due to changes in their flight speed. They have adapted to change their pulse emission frequency in relation to their flight speed so echoes still return in the optimal hearing range.[24]

The new Yinpterochiroptera and Yangochiroptera classification of bats that are supported by molecular evidence, suggest two possibilities for the evolution of echolocation. It may have been gained once in a common ancestor of all bats and was then subsequently lost in the Old World fruit bats, only to be regained in the Horse-Shoe bats; or echolocation was evolved independent in both the Yinpterochiroptera and Yangochirpotera lineages.[25]

Two groups of moths exploit a bat sense to echolocate: tiger moths produce ultrasonic signals to warn the bats that they (the moths) are chemically protected or aposematic. This was once thought to be the biological equivalent of "radar jamming", but this theory has yet to be confirmed. The moths Noctuidae have a hearing organ called a tympanum, which responds to an incoming bat signal by causing the moth's flight muscles to twitch erratically, sending the moth into random evasive manoeuvres.
Other senses

Although the eyes of most microbat species are small and poorly developed, leading to poor visual acuity, none of them are blind. Vision is used to navigate microbats especially for long distances when beyond the range of echolocation. It has even been discovered that some species are able to detect ultraviolet light. They also have a high quality sense of smell and hearing. Bats hunt at night to avoid competition with birds, and travel large distances at most 800 km, in their search for food.[2]
Wings
Thermographic image of a bat using trapped air as insulation.

The finger bones of bats are much more flexible than those of other mammals. One reason is that the cartilage in their fingers lacks calcium and other minerals nearer the tips, increasing their ability to bend without splintering. The cross-section of the finger bone is also flattened compared to the circular cross section that human finger bones have, and is very flexible. The skin on their wing membranes has more elasticity and so can stretch much more than other mammals.

The wings of bats are much thinner than those of birds, so bats can maneuver more quickly and more accurately than birds. It is also delicate, ripping easily.[26] However the tissue of the bat's membrane is able to regrow, such that small tears can heal quickly.[26][27] The surface of their wings is equipped with touch-sensitive receptors on small bumps called Merkel cells, found in most mammals including humans, similarly found on our finger tips. These sensitive areas are different in bats as each bump has a tiny hair in the center,[28] making it even more sensitive and allowing the bat to detect and collect information about the air flowing over its wings, thereby providing feedback to the bat to change its shape of its wing to fly more efficiently.[28] An additional kind of receptor cell is found in the wing membrane of species that use their wings to catch prey. This receptor cell is sensitive to the stretching of the membrane.[28] The cells are concentrated in areas of the membrane where insects hit the wings when the bats capture them.
Other

The teeth of microbats resemble insectivorans. They are very sharp to bite through the hardened armor of insects or the skin of fruit.

Mammals have one-way valves in veins to prevent the blood from flowing backwards, but bats also have one-way valves in arteries.

One species of bat has the longest tongue of any mammal relative to its body size. This is beneficial to them in terms of pollination and feeding. Their long narrow tongues can reach deep into the long cup shape of some flowers. When their tongue retracts, it coils up inside their rib cage.[29]
Behaviour

Most microbats are nocturnal and are active at twilight. A large portion of bats migrate hundreds of kilometres to winter hibernation dens,[30] some pass into torpor in cold weather, rousing and feeding when warm weather allows for insects to be active.[31] Others retreat to caves for winter and hibernate for six months.[31] Bats rarely fly in rain as the rain interferes with their echo location, and they are unable to locate their food.

The social structure of bats varies, with some bats leading a solitary life and others living in caves colonized by more than a million bats.[32] The fission-fusion social structure is seen among several species of bats. The term "fusion" refers to a large numbers of bats that congregate together in one roosting area and "fission" refers to breaking up and the mixing of subgroups, with individual bats switching roosts with others and often ending up in different trees and with different roostmates.

Studies also show that bats make all kinds of sounds to communicate with others. Scientists in the field have listened to bats and have been able to identify some sounds with some behaviour bats will make after the sounds are made.[32]

70% of bat species are insectivorous, locating their prey by means of echolocation. Of the remainder, most feed on fruits.[33] Only three species sustain themselves with blood. Some species even prey on vertebrates: these are the leaf-nosed bats (Phyllostomidae) of Central America and South America, and the two bulldog bat (Noctilionidae) species, which feed on fish. At least two species of bat are known to feed on bats: the Spectral Bat, also known as the American False Vampire bat, and the Ghost Bat of Australia.[33] One species, the Greater Noctule bat, catches and eats small birds in the air.

Predators of bats include bat hawks and bat falcons.
Reproduction
Newborn Common Pipistrelle, Pipistrellus pipistrellus.
Colony of Mouse-eared bats, Myotis myotis.

Most bats have a breeding season, which is in the spring for species living in a temperate climate. Bats may have one to three litters in a season, depending on the species and on environmental conditions such as the availability of food and roost sites. Females generally have one offspring at a time, which could be a result of the mother's need to fly to feed while pregnant. Female bats nurse their youngsters until they are nearly adult size; this is because a young bat cannot forage on its own until its wings are fully developed.

Female bats use a variety of strategies to control the timing of pregnancy and the birth of young, to make delivery coincide with maximum food ability and other ecological factors. Females of some species have delayed fertilization, in which sperm are stored in the reproductive tract for several months after mating. In many such cases, mating occurs in the fall, and fertilization does not occur until the following spring. Other species exhibit delayed implantation, in which the egg is fertilized after mating, but remains free in the reproductive tract until external conditions become favorable for giving birth and caring for the offspring. In yet another strategy, fertilization and implantation both occur but development of the fetus is delayed until favorable conditions prevail. All of these adaptations result in the pup being born during a time of high local production of fruit or insects.

At birth the wings are too small to be used for flight. Young microbats become independent at the age of 6 to 8 weeks, while megabats do not until they are four months old.

A single bat can live over 20 years, but the bat population growth is limited by the slow birth rate.[34]
Hunting, feeding, and drinking

A very young bat in Tamil Nadu.

Newborn bats rely on the milk from their mother’s nipples for sustenance.[35] When they are a few weeks old, bats are expected to fly and hunt on their own. It is up to them to find and catch their prey, along with satisfying their thirst.[36]
Hunting

Most bats are nocturnal creatures. Their daylight hours are spent grooming, sleeping, and resting; it is during the nighttime hours that they hunt. The means by which bats navigate while finding and catching their prey in the dark was unknown until the 1790s, when Lazzaro Spallanzani conducted a series of experiments on a group of blind bats. These bats were placed in a room submerged in total darkness, with silk threads strung across the room. Even then, the bats were able to navigate their way through the room. Spallanzani concluded that the bats were not using their eyes to fly through complete darkness, but something else.

Spallanzani decided that bats were able to catch and find their prey through the use of their ears. To prove this theory, Spallanzani plugged the ears of the bats in his experiment. To his pleasure, he found that the bats with plugged ears were not able to fly with the same amount of skill and precision that they were able to without their ears plugged.

Bats seem to use their ears to locate and catch their prey, but how they accomplish this wasn’t discovered until the 1930s, by one Donald R. Griffin. Griffin, who was a biology student at Harvard College at the time, discovered that bats use echolocation to locate and catch their prey. When bats fly, they produce a constant stream of high-pitched sounds that only bats are able to hear. When the sound waves produced by these sounds hit an insect or other animal, the echoes bounce back to the bat, and guide them to the source.[36]
Feeding and diet

The majority of food consumed by bats includes insects, fruits and flower nectar, vertebrates and blood.[37] Almost three-fourths of the world’s bats are insect eaters. Insects consumed by bats include both aerial insects, and ground-dwelling insects. Each bat is typically able to consume one third of its body weight in insects each night, and several hundred insects in a few hours. This means that a group of one thousand bats could eat four tons of insects each year. If bats were to become extinct, the insect population is calculated to reach an alarmingly high number.[38]
Vitamin C

In a test of 34 bat species from six major families of bats, including major insect and fruit-eating bat families, found that bats in all tested families have lost the ability to make vitamin C, and this loss may derive from a common bat ancestor, as a single mutation.[39]
Aerial insectivores

Watching a bat catch and eat an insect is difficult. The action is so fast that all one sees is a bat rapidly change directions, and continue on its way. Scientist Frederick A. Webster discovered how bats catch their prey. In 1960, Webster developed a high-speed camera that was able to take one thousand pictures per second. These photos revealed the fast and precise way in which bats catch insects.[36] Occasionally, a bat will catch an insect in mid-air with its mouth, and eat it in the air. However, more often than not, a bat will use its tail membrane or wings to scoop up the insect and trap it in a sort of “bug net”.[35] Then, the bat will take the insect back to its roost. There, the bat will proceed to eat said insect, often using its tail membrane as a kind of napkin, to prevent its meal from falling to the ground.[37]
Forage gleaners (diet of non-flying insects)

These bats typically fly down and grasp their prey off the ground with their teeth, and take it to a nearby perch to eat it. Generally, these bats don’t use echolocation to locate their prey. Instead, they rely on the sounds produced by the insects. Some make unique sounds, and almost all make some noise while moving through the environment.[35]
Fruits and flower nectar
A colony of Great Fruit-eating Bats

Fruit-eating, or frugivory, is a specific habit found in two families of bats. Megachiropterans and microchiropterans both include species of bat that feed on fruits. These bats feed on the juices of sweet fruits, and fulfill the needs of some seeds to be dispersed. The fruits preferred by most fruit-eating bats are fleshy and sweet, but not particularly strong smelling or colorful.[35] To get the juice of these fruits, bats pull the fruit off the trees with their teeth, and fly back to their roost with the fruit in their mouth. There, the bat will consume the fruit in a specific way. To do this, the bats crush open the fruit and eat the parts that satisfy their hunger. The remainder of the fruit; the seeds and pulp, are spat onto the ground. These seeds take root and begin to grow into new fruit trees.[38] “Over one hundred and fifty types of plants depend on bats in order to reproduce”.[38]

Some bats prefer the nectar of flowers to insects or other animals. These bats have evolved specifically for this purpose. For example, these bats possess long muzzles and long extrusible tongues covered in fine bristles that aid them in feeding on particular flowers and plants.[35] When they sip the nectar from these flowers, pollen gets stuck to their fur, and is dusted off when the bat takes flight, thus pollinating the plants below them.[38] The rainforest is said to be the most benefitted out off all the biomes that bats live in, because of the large variety of appealing plants.[40] Because of their specific eating habits, nectar-feeding bats are more prone to extinction than any other type of bat.[41] However, according to a study, bats benefit from eating fruits and nectar just as much from eating insects.[42]
Vertebrates

Although most bats are not included in this group, there is a small group of carnivorous bats which feed on other vertebrates and are considered the top carnivores of the bat world.[35] These bats typically eat a variety of animals, but normally consume frogs, lizards, birds, and sometimes other bats.[38] For example, one vertebrate predator, Trachops cirrhosus, is particularly skilled at catching frogs. These bats locate large groups of frogs by distinguishing their mating calls from other sounds around them. They follow the sounds to the source and pluck them from the surface of the water with their sharp canine teeth.[35] Another example is the Greater Noctule bat which is believed to catch birds on the wing.

There are also several species of bat that feed on fish. These types of bats are found on almost all continents. They use echolocation to detect tiny ripples in the water’s surface to locate fish. From there, the bats swoop down low, inches from the water, and use specially enlarged claws on their hind feet to grab the fish out of the water. The bats then take the fish to a feeding roost and consume the animal.[35]
Blood

There are a few species of bat that consume blood exclusively as their diet. This type of diet is referred to as hematophagy, and three species of bat exhibit this behavior. These species include the Common Vampire Bat, the White-winged Vampire Bat, and the Hairy-legged Vampire Bat. The Common Vampire Bat typically consumes the blood of mammals, while the Hairy-legged and White-winged feed on the blood of birds.[43]
Results of eating

Bats’ dung, or guano, is so rich in nutrients, that it is mined from caves, bagged, and used by farmers to fertilize their crops. Also, guano was used in the U.S. Civil War to make gunpowder.[38]

There comes a time in the year that some bats will not eat to supply themselves with food for the night, but for the coming months. These bats are beginning to hibernate. To do this, the bat will eat as much food as its body can contain, being as fat as possible. The bat’s body then takes from the supply of fat for energy, but very slowly, because all body activities have slowed down. This supply of fat will last until the spring season arrives.[36]
Drinking

Generally, bats drink water. In 1960, Frederic A. Webster discovered how bats are able to acquire this water. To do this, Webster developed a high-speed camera and flashgun that could take one thousand photos per second. Webster's camera captured the bat's method of skimming the surface of a body of water, and lowering its jaw to get just one drop of water. It then skims again to get a second drop of water, and then again to get a third, and so on, until it has had its fill of water. Its precision and control is very fine, and it almost never misses.[36] Other bats such as the flying fox or fruit bat gently skim the water's surface, then land nearby to lick water from their chest fur.[44]
Conservation efforts

Through conservancy efforts, such as the Organization for Bat Conservation, bats are becoming better understood and people beginning to understand the crucial role bats play in insect control and pollination.

In the United Kingdom all bats are protected under the Wildlife and Countryside Acts, and even disturbing a bat or its roost can be punished with a heavy fine.

In Sarawak, Malaysia bats are protected species under the Wildlife Protection Ordinance 1998 (see Malaysian Wildlife Law). The large Naked bat (see Mammals of Borneo) and Greater Nectar bat are consumed by the local communities.

Bats can be a tourist attraction. The Congress Avenue Bridge in Austin, Texas is the summer home to North America's largest urban bat colony, an estimated 1,500,000 Mexican free-tailed bats, which eat an estimated 10,000 to 30,000 pounds of insects each night. An estimated 100,000 tourists per year visit the bridge at twilight to watch the bats leave the roost.
Artificial roosts
Very large bat house, Tallahassee, Florida, United States.

Many people put up bat houses to attract bats just like many people put up birdhouses to attract birds. Reasons for this vary, but mostly center around the fact that bats are the primary nocturnal insectivores in most if not all ecologies. Bat houses can be made from scratch, made from kits, or bought ready made. Plans for bat houses exist on many web sites, as well as guidelines for designing a bat house.[45] Some conservation societies are giving away free bat houses to bat enthusiasts worldwide .

A bat house constructed in 1991 at the University of Florida campus next to Lake Alice in Gainesville, Florida has a population of over 100,000 free-tailed bats.[46]

In Britain, British hardened field defences of World War II have been converted to make roosts for bats. Pillboxes that are well dug-in and thick walled are naturally damp and provide a stable thermal environment that is required by bats that would otherwise hibernate in caves. With a few minor modifications, suitable pillboxes can be converted to artificial caves for bats.[47][48]

Again in the UK, purpose-built bat houses are occasionally built when existing roosts are destroyed by developments such as new roads; one such has been built associated with bat bridges on the new (2008) A38 Dobwalls bypass.
Threats
A little brown bat with white nose syndrome.

While conservation efforts are in place to protect bats, many threats still remain.
White nose syndrome
Main article: White nose syndrome

White nose syndrome is a condition associated with the deaths of more than a million bats in the Northeastern United States.[49] The disease is named after a white fungus found growing on the muzzles, ears, and wings of some afflicted bats, but it is not known if the fungus is the primary cause of the disease or is merely an opportunistic infection.[50] Mortality rates of 90–100% have been observed in some caves.[50] At least six species of hibernating bats are affected, including the endangered Indiana bat.[51] Because the affected species have a long lifespan and a low birth rate of only about one offspring per year, it is not expected that populations will recover quickly.[51]
Barotrauma and wind turbines

Evidence suggests Barotrauma is causing bat fatalities around wind farms.[52] The lungs of bats are typical mammalian lungs, and unlike the lungs of birds it has been hypothesized they are more sensitive to sudden air pressure changes in their immediate vicinity such as wind turbines, and are more liable to rupture them to explain their apparent higher rate of mortality rate with such devices.[53] Bats suffer a higher death rate than birds in the neighborhood of wind turbines[54][55][56] since there are no signs of external trauma, the cause has been hypothesized to be a greater sensitivity to sudden pressure fluctuations in the mammalian lung than in that of birds.[57] In addition, it has been suggested that bats are attracted to these structures, perhaps seeking roosts, and thereby increasing the death rate.[53]
Pathogens and role in the transmission of zoonoses

Among ectoparasites, bats occasionally carry fleas, but are one of the few mammalian orders that cannot host lice (most of the others are water animals).

Bats are natural reservoir for a large number of zoonotic pathogens[58] including rabies,[59] severe acute respiratory syndrome (SARS),[60] Henipavirus (i.e. Nipah virus and Hendra virus)[61] and possibly ebola virus.[62][63] Their high mobility, broad distribution, and social behaviour (communal roosting, fission-fusion social structure) make bats favourable hosts and vectors of disease. Many species also appear to have a high tolerance for harbouring pathogens and often do not develop disease while infected. However, contrary to folklore, this is not true of rabies, which is as fatal to bats as it is to all other species. However, a bat may be ill with rabies for a longer time than other mammals.

In regions where rabies is endemic, only 0.5% of bats carry the disease. However, of the few cases of rabies reported in the United States every year not caused by dogs, most are caused by bat bites.[64] Those that are rabid may be clumsy, disoriented, and unable to fly, which makes it more likely that they will come into contact with humans. Although one should not have an unreasonable fear of bats, one should avoid handling them or having them in one's living space, as with any wild animal. If a bat is found in living quarters near a child, mentally handicapped person, intoxicated person, sleeping person, or pet, the person or pet should receive immediate medical attention for rabies. Bats have very small teeth and can bite a sleeping person without being felt. There is evidence that it is possible for the bat rabies virus to infect victims purely through airborne transmission, without direct physical contact of the victim with the bat itself.[65][66]

If a bat is found in a house and the possibility of exposure cannot be ruled out, the bat should be sequestered and an animal control officer called immediately, so that the bat can be analysed. This also applies if the bat is found dead. If it is certain that nobody has been exposed to the bat, it should be removed from the house. The best way to do this is to close all the doors and windows to the room except one that opens to the outside. The bat should soon leave.

Due to the risk of rabies and also due to health problems related to their faecal droppings (guano), bats should be excluded from inhabited parts of houses. The Center for Disease Control and Prevention provides full detailed information on all aspects of bat management, including how to capture a bat, what to do in case of exposure, and how to bat-proof a house humanely.[67] In certain countries, such as the United Kingdom, it is illegal to handle bats without a license.

Where rabies is not endemic, as throughout most of Western Europe, small bats can be considered harmless. Larger bats can give a nasty bite. They should be treated with the respect due to any wild animal.
Mythology
"Nightwing," a work of art by Dale Whistler in Austin, Texas.

The bat is sacred in Tonga and is often considered the physical manifestation of a separable soul . Bats are closely associated with vampires, who are said to be able to shapeshift into bats, fog, or wolves. Bats are also a symbol of ghosts, death, and disease. Among some Native Americans, such as the Creek, Cherokee and Apache, the bat is a trickster spirit.

Chinese lore claims the bat is a symbol of longevity and happiness, and is similarly lucky in Poland and geographical Macedonia and among the Kwakiutl and Arabs. The bat is also a heraldic animal of the Spanish autonomous community of Valencia.

Pre-Columbian cultures associated animals with gods and often displayed them in art. The Moche people depicted bats in their ceramics.[68]

In Western Culture, the bat is often a symbol of the night and its foreboding nature. The bat is a primary animal associated with fictional characters of the night, both villains like Dracula and heroes like Batman. The association of the fear of the night with the animal was treated as a literary challenge by Kenneth Oppel, who created a best selling series of novels, beginning with Silverwing, which feature bats as the central heroic figures much as anthropomorphized rabbits were the central figures to the classic novel Watership Down.

An old wives' tale has it that bats will entangle themselves in people's hair. One likely source of this belief is that insect-eating bats seeking prey may dive erratically toward people, who attract mosquitoes and gnats, leading the squeamish to believe that the bats are trying to get in their hair.
Mesoamerica
Bat, Moche Culture 100 A.D. Larco Museum Lima, Peru.

In Mesoamerican mythology during the Classic-Contemporary period, bats symbolized the land of the dead, which was considered to be the underworld.[69] They also symbolized destruction and decay. Bats may have symbolized in this way because they fly only at night and dwell in caves during the daytime and are associated with human skulls and bones by classic Maya ceramists. Central Mexicans sometimes depicted bats having snouts that looked like sacrificial knives and carrying human head in the Postclassic era.[70] Bat images were engraved onto funerary urns and were emphasized with large claws and round ears by Zapotecs. They were commonly associated with death.[71] The depiction of bats on funeral urns and goods took on some the characteristics of the jaguar which was and still is another entity of the night and the underworld. There have also been instances where bats are portrayed next to other unseemly animals including scorpions and other nocturnal animals such as owls.

A gigantic, life-size ceramic bat-man has been discovered and dug up from the Templo Mayor. The Templo Mayor is located in the center of the Mexica capital of Tenochtitlan. Known as a god of death, this statue has the clawed feet and hands of a bat, but the body of a man. The statue's human-like eyes bulged out from the bat-like head, making the Zapotec images very realistic and living. It was said that in the 1930s the Kaqchikel Maya proclaimed that the bat was the Devil’s provider. Kaqchikel would leave the Devil’s underworld home and collect blood from the animals to be used for scrumptious meals to feed the Devil. “In the myths, the beast of prey and the animal that is preyed upon play two significant roles. They represent two aspects of life—the aggressive, killing, conquering, creating aspect of life, and the one that is the matter or, you might say, the subject matter”.[72] In the Devil’s underworld, dead sinners would work off their sins in order to get to heaven, indicating that the bat was too a sinner and worked under the authority of the Devil.[73]
Oaxaca

Oaxacans believe that the jealousy of the bat in wanting birds' feathers that gently fit their bodies led him to become nocturnal. The bat feeling isolated and undesirable spoke to God after that he complained he was extremely cold. God, fair and just turned to birds in the animal kingdom and asked if they would show compassion and donate a feather to the bat so the feathers would keep him warm. The birds all agreed, and began to pluck one feather from their bodies to give to the bat. With all of the feathers, the bat became much magnificent looking than all birds, even able to spread color to the night sky. During daylight, the bat created rainbows that reflected vibrant colors from the sun. The bat soon became overly arrogant and conceited, due having this new and improved look. The birds grew tired of the bat’s self conceitedness and glorification, and so decided to fly up to heaven and speak to God to do something. The birds informed to God of the bat's behaviour, God was surprised and so decided to take a look himself. When on Earth, God called on the bat to show him what he was doing. The bat began to fly across the light blue sky, where one by one each feather began to fall out, uncovering the bat’s natural ugly looking body. The bat became ashamed and distressed of his appearance after all feathers came off, missing the beautiful, plentiful feathers that he had, that he decided to hide in caves during the day. He would only come out during the night, searching high and low for the feathers to avoid embarrassment that he will not be seen during his search.[73]
East Nigeria

According to a particular East Nigerian tale, the bat developed its nocturnal habits after causing the death of his partner the bush-rat. The bat and the bush-rat would share activities such as rummaging through the grass and trees, hunting, talking and bonding during the day. When at night, the bat and the bush-rat would alternate in cooking duties cooking what was caught, and eat together. It appeared to a dedicated partnership, however the bat hated the bush-rat immensely. The bush rat always found the bat’s soup more appetising so when eating dinner one night asked the bat why the soup tasted better than his own and also asked how it was made. The bat agreed to show him how to make it the next day but instead was forming a malicious plan.

Next day as bat prepared his soup, the bush-rat came, greeting him and asked if he could be shown what was agreed yesterday. Earlier, the bat has found a pot looking exactly like the one he used usually, but it held warm water and so decided to use this instead. The bat explained to the bush-rat that to make his soup, he had to boil himself prior to serving the soup where sweetness and flavor of the soup came from the flesh. The bat jumped in the pot seemingly excited, with the bush-rat mesmerised. After a few minutes the bat climbed out and while the bush-rat was distracted, switched pots. The bat then served his soup out of the soup pot, both tasted it. Over anxious and eager, the bush-rat, jumped into the pot of warm water. He stayed much longer in the pot dying in the process.

When the bush-rat’s wife returned that night to find her husband dead, she wept and ran to the chief of the land's house telling him about what happened and what she was sure what the bat had done. In hearing this, the chief became angry, ordering for the immediate arrest of the bat. It just so happened that the bat was flying over the house and overheard what was just said. He quickly went into hiding high up in a tree. When the chief’s men went looking for the bat, he could not be found. The search to arrest the bat carried on over several days, but still could not be found. The bat needed to eat, so flew out of hiding every night to hunt for food to escape of being arrested. This, according to Eastern Nigeria mythology, is why bats only fly at night.[74]
Heraldry
Main article: Bat (heraldry)
Burgee of the Royal Valencia Yacht Club.

The bat is sometimes used as a heraldic symbol. The coats of arms of certain cities in eastern Spain, like Valencia, Palma de Mallorca and Fraga have the bat over the shield. Formerly the Barcelona city coat of arms also had a bat crowning it, but the bat has been removed in the present-day versions.

The heraldic use of the bat in Valencia, Catalonia and the Balearic Islands has its origins in a winged dragon (vibra or vibria), which featured in King James I of Aragon's helmet or cimera reial. This is the most widely accepted theory, although there is also a legend that says that due to the intervention of a bat, King James was able to win a crucial battle against the Saracens that allowed him to win Valencia for his kingdom.

The use of the bat as a heraldic symbol is prevalent in the territories of the former Crown of Aragon and it is little used elsewhere. However, it can be found in a few places, as in the coats of arms of the city of Albacete, in Spain, as well as the town of Montchauvet (Yvelines), in France.

Certain Spanish soccer clubs including Valencia CF and Levante UD use bats in their badges.

The Burgee of the Royal Valencia Yacht Club (Reial Club Nàutic de València) displays a bat on a golden field in its center.
In popular culture

In Western Culture, bats are a symbol of darkness and forbidden nature, something that is associated with fictional dark characters. For example: superheroes like Batman, villains like Dracula and videogame characters like Rouge the Bat of Sonic the Hedgehog videogame series.

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63. ^ Charles Q. Choi (March 2006). "Going to Bat". Scientific American: pp. 24, 26. http://www.sciam.com/article.cfm?id=going-to-bat. Retrieved 2007-12-29. "Long known as vectors for rabies, bats may be the origin of some of the most deadly emerging viruses, including SARS, Ebola, Nipah, Hendra and Marburg." Note: This could be considered a lay summary of the various scientific publications cited in the preceding sentence.
64. ^ Gibbons, Robert V.; Charles Rupprecht (2000). "Twelve Common Questions About Human Rabies and Its Prevention" (PDF). Infectious Diseases in Clinical Practice (Lippincott Williams & Wilkins) 9: 202–207. doi:10.1097/00019048-200009050-00005. http://www.cdc.gov/rabies/docs/12_questions_rabies.pdf. Retrieved 2007-12-29. "Excluding dog bites that occurred outside of the country, 22 of the 31 (71%) human cases of rabies in the United States since 1980 have been associated with bat rabies virus variants.". Note: the 71% figure in the quote would be for the 20 year period from 1980 to c.2000.
65. ^ Constantine, Denny G. (April 1962). "Rabies transmission by nonbite route" (PDF). Public Health Reports (Public Health Service) 77 (4): 287–289. PMID 13880956. PMC 1914752. http://www.pubmedcentral.nih.gov/picrender.fcgi?artid=1914752&blobtype=pdf. Retrieved 2007-12-29. "These findings support consideration of an airborne medium, such as an aerosol, as the mechanism of rabies transmission in this instance.".
66. ^ Messenger, Sharon L.; Jean S. Smith and Charles E. Rupprecht (2002-09-15). "Emerging Epidemiology of Bat-Associated Cryptic Cases of Rabies in Humans in the United States". Clinical Infectious Diseases 35 (6): 738–747. doi:10.1086/342387. PMID 12203172. http://www.journals.uchicago.edu/doi/abs/10.1086/342387. Retrieved 2007-12-29. "Cryptic rabies cases are those in which a clear history of exposure to rabies virus cannot be documented, despite extensive case‐history investigation. Absence of a documented bite history reflects inherent difficulties in obtaining accurate animal‐contact information.... <gap> Thus, absence of bite‐history data does not mean that a bite did not occur.".
67. ^ Center for Disease Control's website on bats and rabies
68. ^ Berrin, Katherine & Larco Museum. The Spirit of Ancient Peru:Treasures from the Museo Arqueológico Rafael Larco Herrera. New York: Thames and Hudson, 1997.
69. ^ http://www.aztec-history.net/aztec_symbols
70. ^ Kay Almere Read and Jason J. Gonzalez. 2000. Mesoamerican Mythology. Oxford University Press. pp. 132
71. ^ http://www.oaxacanwoodcarving.com/mythnatr.html
72. ^ Joseph Cambell and Bill Moyers. 1988. The Power of Myth. Doubleday. pp. 91
73. ^ a b Kay Almere Read and Jason J. Gonzalez. 2000. Mesoamerican Mythology. Oxford University Press. pp. 132–134
74. ^ Arnott, Kathleen. 1962. African Myths and Legends. Oxford University Press. Pp. 150–152

General references

* Greenhall, Arthur H. 1961. Bats in Agriculture. A Ministry of Agriculture Publication. Trinidad and Tobago.
* Nowak, Ronald M. 1994. " Walker's BATS of the World". The Johns Hopikins University Press, Baltimore and London.
* John D. Pettigrew's summary on Flying Primate Hypothesis
* Altringham, J.D. 1998. Bats: Biology and Behaviour. Oxford: Oxford University Press.
* Dobat, K.; Holle, T.P. 1985. Blüten und Fledermäuse: Bestäubung durch Fledermäuse und Flughunde (Chiropterophilie). Frankfurt am Main: W. Kramer & Co. Druckerei.
* Fenton, M.B. 1985. Communication in the Chiroptera. Bloomington: Indiana University Press.
* Findley, J.S. 1995. Bats: a Community Perspective. Cambridge: Press Syndicate of the University of Cambridge.
* Fleming, T.H. 1988. The Short-Tailed Fruit Bat: a Study in Plant-Animal Interactions. Chicago: The University of Chicago Press.
* Kunz, T.H. 1982. Ecology of Bats. New York: Plenum Press.
* Kunz, T.H.; Racey, P.A. 1999. Bat Biology and Conservation. Washington: Smithsonian Institution Press.
* Kunz, T.H.; Fenton, M.B. 2003. Bat Ecology. Chicago: The University of Chicago Press.
* Neuweiler, G. 1993. Biologie der Fledermäuse. Stuttgart: Georg Thieme Verlag.
* Nowak, R.M. 1994. Walker's Bats of the World. Baltimore: The Johns Hopkins University Press.
* Richarz, K. & Limbruner, A. 1993. The World of Bats. Neptune City: TFH Publications.
* Teeling, E.C. 2009. Chiroptera. Oxford University Press.
* Twilton, B. 1999. My Life as The Bat. Liverpool Hope University press