These were traditionally divided into two suborders: 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 s. Bats were formerly grouped in the superorder Archonta , along with the treeshrews Scandentia , colugos Dermoptera , and primates.
Euarchontoglires primates, treeshrews, rodents, rabbits. Eulipotyphla hedgehogs, shrews, moles, solenodons.
Carnivora cats, hyenas, dogs, bears, seals. Perissodactyla horses, tapirs, rhinos. Cetartiodactyla camels, ruminants, whales.
The phylogenetic relationships of the different groups of bats have been the subject of much debate. The traditional subdivision into Megachiroptera and Microchiroptera reflected the view that these groups of bats had evolved independently of each other for a long time, from a common ancestor already capable of flight.
This hypothesis recognised 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 natural or monophyletic group. Megadermatidae false vampire bats.
Hipposideridae Old World leaf-nosed bats. Miniopteridae long winged bat. Mystacinidae New Zealand short-tailed bats. Phyllostomidae New World leaf-nosed bats.
Genetic evidence indicates that megabats originated during the early Eocene , and belong within the four major lines of microbats. In the s, 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,  debate continues about the meaning of the genetic and morphological evidence.
The discovery of an early fossil bat from the 52 million year old Green River Formation , Onychonycteris finneyi , indicates that flight evolved before echolocative abilities.
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.
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.
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. 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.
Bats are placental mammals. Around twenty years later, the German naturalist Johann Friedrich Blumenbach gave them their own order, Chiroptera.
Microbats use echolocation for navigation and finding prey, but megabats apart from those in the genus Rousettus do not, relying instead on their eyesight.
The following classification from Agnarsson and colleagues in reflects the traditional division into megabat and microbat suborders.
The head and teeth shape of bats can vary by species. In general, megabats have longer snouts, larger eye sockets and smaller ears, giving them a more dog-like appearance, which is the source of their nickname of "flying foxes".
Small insect-eating bats can have as many as 38 teeth, while vampire bats have only Bats that feed on hard-shelled insects have fewer but larger teeth with longer canines and more robust lower jaws than species that prey on softer bodied insects.
In nectar-feeding bats, the canines are long while the cheek-teeth are reduced. In fruit-eating bats, the cusps of the cheek teeth are adapted for crushing.
The upper incisors of vampire bats lack enamel , which keeps them razor-sharp. Bats are the only mammals capable of sustained flight, as opposed to gliding , as in the flying squirrel.
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.
The wing bones of bats have a slightly lower breaking stress point than those of birds. 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.
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.
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.
The patagium is the wing membrane; it is stretched between the arm and finger bones, and down the side of the body to the hind limbs and tail. This skin membrane consists of connective tissue , elastic fibres , nerves , muscles , and blood vessels.
The muscles keep the membrane taut during flight. The patagium is an extremely thin double layer of epidermis; these layers are separated by a connective tissue centre, rich with collagen and elastic fibres.
The membrane has no hair follicles or sweat glands, except between the fingers. To achieve flight, a bat exerts force inwards at the points where the membrane meets the skeleton, so that an opposing force balances it on the wing edges perpendicular to the wing surface.
This adaptation does not permit bats to reduce their wingspans, unlike birds, which can partly fold their wings in flight, radically reducing the wing span and area for the upstroke and for gliding.
Hence bats cannot travel over long distances as birds can. Nectar- and pollen-eating bats can hover, in a similar way to hummingbirds. The sharp leading edges of the wings can create vortices , which provide lift.
The vortex may be stabilised by the animal changing its wing curvatures. When not flying, bats hang upside down from their feet, a posture known as roosting.
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.
This difference is reflected in the structure of the cervical or neck vertebrae in the two groups, which are clearly distinct.
Muscular power is needed to let go, but not to grasp a perch or when holding on. When on the ground, most bats can only crawl awkwardly.
A few species such as the New Zealand lesser short-tailed bat and the common vampire bat are agile on the ground.
Both species make lateral gaits the limbs move one after the other when moving slowly but vampire bats move with a bounding gait all limbs move in unison at greater speeds, the folded up wings being used to propel them forward.
Vampire bat likely evolved these gaits to follow their hosts while short-tailed bats developed in the absence of terrestrial mammal competitors.
Enhanced terrestrial locomotion does not appear to have reduced their ability to fly. 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.
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. 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.
When the bat has its wings spread it allows for an increase in surface area to volume ratio. The digestive system of bats has varying adaptations depending on the species of bat and its diet.
As in other flying animals, food is processed quickly and effectively to keep up with the energy demand. Insectivorous bats may have certain digestive enzymes to better process insects, such as chitinase to break down chitin , which is a large component of insects.
Nectivorous and frugivorous bats have more maltase and sucrase enzymes than insectivorous, to cope with the higher sugar contents of their diet.
The adaptations of the kidneys of bats vary with their diets. Carnivorous and vampire bats consume large amounts of protein and can output concentrated urine ; their kidneys have a thin cortex and long renal papillae.
Frugivorous bats lack that ability and have kidneys adapted for electrolyte -retention due to their low-electrolyte diet; their kidneys accordingly have a thick cortex and very short conical papillae.
Bats have higher metabolic rates associated with flying, which lead to an increased respiratory water loss. Their large wings are composed of the highly vascularized membranes, increasing the surface area, and leading to cutaneous evaporative water loss.
They are also susceptible to blood urea poisoning if they do not receive enough fluid. The structure of the uterine system in female bats can vary by species, with some having two uterine horns while others have a single mainline chamber.
Microbats and a few megabats emit ultrasonic sounds to produce echoes. This allows bats to detect, localise, and classify their prey in darkness.
Bat calls are some of the loudest airborne animal sounds, and can range in intensity from 60 to decibels. The latter is most pronounced in the horseshoe bats Rhinolophus spp.
In low-duty cycle echolocation, bats can separate their calls and returning echoes by time. They have to time their short calls to finish before echoes return.
Bats contract their middle ear muscles when emitting a call, so they can avoid deafening themselves. The time interval between the call and echo allows them to relax these muscles, so they can hear the returning echo.
In 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 this range to avoid deafening themselves. They then receive echoes back at the finely tuned frequency range by taking advantage of the Doppler shift of their motion in flight.
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.
In addition to echolocating prey, bat ears are sensitive to the fluttering of moth wings, the sounds produced by tymbalate insects, and the movement of ground-dwelling prey, such as centipedes and earwigs.
The complex geometry of ridges on the inner surface of bat ears helps to sharply focus echolocation signals, and to passively listen for any other sound produced by the prey.
These ridges can be regarded as the acoustic equivalent of a Fresnel lens , and exist in a large variety of unrelated animals, such as the aye-aye , lesser galago , bat-eared fox , mouse lemur , and others.
By repeated scanning, bats can mentally construct an accurate image of the environment in which they are moving and of their prey. The eyes of most microbat species are small and poorly developed, leading to poor visual acuity , but no species is blind.
Microbats may use their vision for orientation and while travelling between their roosting grounds and feeding grounds, as echolocation is only effective over short distances.
Some species can detect ultraviolet UV. As the bodies of some microbats have distinct coloration, they may be able to discriminate colours.
Megabat species often have eyesight as good as, if not better than, human vision. Their eyesight is adapted to both night and daylight vision, including some colour vision.
Microbats use a polarity-based compass, meaning that they differentiate north from south, unlike birds, which use the strength of the magnetic field to differentiate latitudes , which may be used in long-distance travel.
The mechanism is unknown but may involve magnetite particles. Most bats are homeothermic having a stable body temperature , the exception being the vesper bats Vespertilionidae , the horseshoe bats Rhinolophidae , the free-tailed bats Molossidae , and the bent-winged bats Miniopteridae , which extensively use heterothermy where body temperature can vary.
The wings are filled with blood vessels, and lose body heat when extended. At rest, they may wrap their wings around themselves to trap a layer of warm air.
Smaller bats generally have a higher metabolic rate than larger bats, and so need to consume more food in order to maintain homeothermy.
Bats may avoid flying during the day to prevent overheating in the sun, since their dark wing-membranes absorb solar radiation. Bats may not be able to dissipate heat if the ambient temperature is too high;  they use saliva to cool themselves in extreme conditions.
Bats also possess a system of sphincter valves on the arterial side of the vascular network that runs along the edge of their wings. When fully open, these allow oxygenated blood to flow through the capillary network across the wing membrane; when contracted, they shunt flow directly to the veins, bypassing the wing capillaries.
This allows bats to control how much heat is exchanged through the flight membrane, allowing them to release heat during flight.
Many other mammals use the capillary network in oversized ears for the same purpose. Torpor , a state of decreased activity where the body temperature and metabolism decreases, is especially useful for microbats, as they use a large amount of energy while active, depend upon an unreliable food source, and have a limited ability to store fat.
Torpid states last longer in the summer for megabats than in the winter. During hibernation , bats enter a torpid state and decrease their body temperature for Heterothermic bats during long migrations may fly at night and go into a torpid state roosting in the daytime.
Unlike migratory birds, which fly during the day and feed during the night, nocturnal bats have a conflict between travelling and eating.
The energy saved reduces their need to feed, and also decreases the duration of migration, which may prevent them from spending too much time in unfamiliar places, and decrease predation.
In some species, pregnant individuals may not use torpor. Small prey may be absent in the diets of large bats as they are unable to detect them.
Flight has enabled bats to become one of the most widely distributed groups of mammals. Bat roosts can be found in hollows, crevices, foliage, and even human-made structures, and include "tents" the bats construct with leaves.
In temperate areas, some microbats migrate hundreds of kilometres to winter hibernation dens;  others pass into torpor in cold weather, rousing and feeding when warm weather allows insects to be active.
Different bat species have different diets, including insects, nectar, pollen, fruit and even vertebrates. Insectivorous bats may eat over percent of their body weight, while frugivorous bats may eat over twice their weight.
The Chiroptera as a whole are in the process of losing the ability to synthesise vitamin C. Most microbats, especially in temperate areas, prey on insects.
Fruit eating, or frugivory, is found in both major suborders. Bats prefer ripe fruit, pulling it off the trees with their teeth.
They fly back to their roosts to eat the fruit, sucking out the juice and spitting the seeds and pulp out onto the ground. This helps disperse the seeds of these fruit trees, which may take root and grow where the bats have left them, and many species of plants depend on bats for seed dispersal.
Nectar-eating bats have acquired specialised adaptations. These bats possess long muzzles and long, extensible tongues covered in fine bristles that aid them in feeding on particular flowers and plants.
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 the tongue retracts, it coils up inside the rib cage. Around species of flowering plant rely on bat pollination and thus tend to open their flowers at night.
Some bats prey on other vertebrates, such as fish, frogs, lizards, birds and mammals. These bats locate large groups of frogs by tracking their mating calls, then plucking them from the surface of the water with their sharp canine teeth.
A few species, specifically the common, white-winged , and hairy-legged vampire bats, only feed on animal blood hematophagy. The common vampire bat typically feeds on large mammals such as cattle ; the hairy-legged and white-winged vampires feed on birds.
Bats are subject to predation from birds of prey , such as owls , hawks , and falcons , and at roosts from terrestrial predators able to climb, such as cats.
Speakman argue that bats evolved nocturnality during the early and middle Eocene period to avoid predators. Among ectoparasites , bats carry fleas and mites , as well as specific parasites such as bat bugs and bat flies Nycteribiidae and Streblidae.
White nose syndrome is a condition associated with the deaths of millions of bats in the Eastern United States and Canada.
The fungus is mostly spread from bat to bat, and causes the disease. Bats are natural reservoirs for a large number of zoonotic pathogens ,  including rabies , endemic in many bat populations,    histoplasmosis both directly and in guano,  Nipah and Hendra viruses ,   and possibly the ebola virus.
Compared to rodents, bats carry more zoonotic viruses per species, and each virus is shared with more species. Some bats lead solitary lives, while others live in colonies of more than a million.
This may serve to introduce young to hibernation sites, signal reproduction in adults and allow adults to breed with those from other groups.
Several species have a fission-fusion social structure , where large numbers of bats congregate in one roosting area, along with breaking up and mixing of subgroups.
Within these societies, bats are able to maintain long term relationships. Bats are among the most vocal of mammals and produce calls to attract mates, find roost partners and defend resources.
These calls are typically low-frequency and can travel long distances. Males sing to attract females. Songs have three phrases: Bat songs are highly stereotypical but with variation in syllable number, phrase order, and phrase repetitions between individuals.
Calls differ between roosting groups and may arise from vocal learning. The animals made slightly different sounds when communicating with different individual bats, especially those of the opposite sex.
Bats in flight make vocal signals for traffic control. Greater bulldog bats honk when on a collision course with each other.
Bats also communicate by other means. Male little yellow-shouldered bats Sturnira lilium have shoulder glands that produce a spicy odour during the breeding season.
Like many other species, they have hair specialised for retaining and dispersing secretions. Such hair forms a conspicuous collar around the necks of the some Old World megabat males.
Male greater sac-winged bats Saccopteryx bilineata have sacs in their wings in which they mix body secretions like saliva and urine to create a perfume that they sprinkle on roost sites, a behaviour known as "salting".
Salting may be accompanied by singing. Most bat species are polygynous , where males mate with multiple females. Male pipistrelle, noctule and vampire bats may claim and defend resources that attract females, such as roost sites, and mate with those females.
Males unable to claim a site are forced to live on the periphery where they have less reproductive success. For temperate living bats, mating takes place in late summer and early autumn.
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