Bat

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Bat
Temporal range: EocenePresent
Common vampire batGreater horseshoe batGreater short-nosed fruit batEgyptian fruit batMexican free-tailed batGreater mouse-eared bat
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Clade: Scrotifera
Clade: Apo-Chiroptera
Order: Chiroptera
Blumenbach, 1779
Suborders

(traditional):

  • Megachiroptera
  • Microchiroptera

(present):

Worldwide distribution of bat species

Bats are flying

flying foxes, with the giant golden-crowned flying fox
(Acerodon jubatus) reaching a weight of 1.6 kg (3+12 lb) and having a wingspan of 1.7 m (5 ft 7 in).

The second largest order of mammals after

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 direct benefits, at the cost of some disadvantages. Bat dung has been mined as guano from caves and used as fertiliser. Bats consume insect pests, reducing the need for pesticides and other insect management measures. They are sometimes numerous enough and close enough to human settlements to serve as tourist attractions, and they are used as food across Asia and the Pacific Rim. However, fruit bats are frequently considered pests by fruit growers. Due to their physiology, bats are one type of animal that acts as a natural reservoir of many pathogens, such as rabies; and since they are highly mobile, social, and long-lived, they can readily spread disease among themselves. If humans interact with bats, these traits become potentially dangerous to humans. Some bats are also predators of mosquitoes, suppressing the transmission of mosquito-borne diseases.

Depending on the culture, bats may be symbolically associated with positive traits, such as protection from certain diseases or risks, rebirth, or long life, but in the West, bats are popularly associated with darkness, malevolence, witchcraft, vampires, and death.

Etymology

An older English name for bats is

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

Evolutionary history

The delicate

Hassianycteris kumari, both of which lived 48 million years ago, are the first fossil mammals whose colouration has been discovered: both were reddish-brown.[11][12]

Bats were formerly grouped in the superorder

even-toed ungulates, and cetaceans.[14][15][16][17][18] One study places Chiroptera as a sister taxon to odd-toed ungulates (Perissodactyla).[19]

Boreoeutheria

Euarchontoglires (primates, treeshrews, rodents, rabbits)

Laurasiatheria

Eulipotyphla (hedgehogs, shrews, moles, solenodons)

Scrotifera

Chiroptera (bats)

Fereuungulata
Ferae

Pholidota (pangolins)

Carnivora (cats, hyenas, dogs, bears, seals, weasels)

Euungulata

Perissodactyla (horses, tapirs, rhinos)

Cetartiodactyla (camels, ruminants, whales)

sister taxon according to a 2013 study[18]

The flying primate hypothesis proposed that when adaptations to flight are removed, megabats are allied to primates by anatomical features not shared with microbats and thus flight evolved twice in mammals.[20] Genetic studies have strongly supported the monophyly of bats and the single origin of mammal flight.[9][20]

Coevolutionary evidence

An independent molecular analysis trying to establish the dates when bat ectoparasites (bedbugs) evolved came to the conclusion that bedbugs similar to those known today (all major extant lineages, all of which feed primarily on bats) had already diversified and become established over 100 mya (i.e., long before the oldest records for bats, 52 mya), suggesting that they initially all evolved on non-bat hosts and "bats were colonized several times independently, unless the evolutionary origin of bats has been grossly underestimated."[21] Fleas, as a group, are quite old (most flea families formed around the end of the Cretaceous[22]), but no analyses have provided estimates for the age of the flea lineages associated with bats. The oldest known members of a different lineage of bat ectoparasites (bat flies), however, are from roughly 20 mya, well after the origin of bats.[23] The bat-ectoparasitic earwig family Arixeniidae has no fossil record, but is not believed to originate more than 23 mya.[24]

Inner systematic

Chiroptera
Microchiroptera
Rhinolophoidea

Megadermatidae (false vampire bats)

Craseonycteridae (Kitti's hog-nosed bat)

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)

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[25]

Rhinopomatidae.[26] Yangochiroptera includes the other families of bats (all of which use laryngeal echolocation), a conclusion supported by a 2005 DNA study.[26] A 2013 phylogenomic study supported the two new proposed suborders.[18]

Chiroptera

Yangochiroptera (as above)

Yinpterochiroptera

Pteropodidae (megabats)

Rhinolophoidea

Megadermatidae (false vampire bats)

horseshoe bats and allies

Internal relationships of the Chiroptera, with the megabats subsumed within Yinpterochiroptera, according to a 2013 study[18]
Giant golden-crowned flying fox, Acerodon jubatus

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.[27][28] 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.[9] 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.[29] 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.[30][31]

The molecular phylogeny was controversial, as it pointed to microbats not having a unique common ancestry, which implied that some seemingly unlikely transformations occurred. The first is that laryngeal echolocation evolved twice in bats, once in Yangochiroptera and once in the rhinolophoids.[32] 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.[33] 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.[34] 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.[29] 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,[35] but ontogenic analysis of the cochlea supports that laryngeal echolocation evolved only once.[36]

Classification

Bats are

eat insects; others feed on fruit, nectar, pollen, fish, frogs, small mammals, or blood.[40]

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

Below is a table chart following the bat classification of families recognized by various authors of the ninth volume of Handbook of the Mammals of the World published in 2019:[45]

Chiroptera
Blumenbach, 1779
Yinpterochiroptera Springer, Teeling, Madsen, Stanhope & Jong, 2001
Pteropodoidea
J. E. Gray, 1821
Family English Name Number of Species Image Figure
Pteropodidae
J. E. Gray, 1821
Old World fruit bats 191
Rhinolophoidea J. E. Gray, 1825
Family English Name Number of Species Image Figure
Rhinopomatidae
Bonaparte, 1838
Mouse-tailed bats 6
Craseonycteridae
Hill, 1974
Hog-nosed bat 1
Megadermatidae H. Allen, 1864 False-vampires 6
Rhinonycteridae J. E. Gray, 1866 Trident bats 9
Hipposideridae Lydekker, 1891 Old World leaf-nosed bats 88
Rhinolophidae
J. E. Gray, 1825
Horseshoe bats 109
Yangochiroptera Koopman, 1984
Emballonuroidea Gervais in de Castelnau, 1855
Family English Name Number of Species Image Figure
Nycteridae
Van der Hoeven, 1855
Slit-faced bats 15
Emballonuridae Gervais in de Castelnau, 1855 Sheath-tailed bats 54
Noctilionoidea J. E. Gray, 1821
Family English Name Number of Species Image Figure
Myzopodidae
Thomas, 1904
Madagascar sucker-footed bats 2
Mystacinidae Dobson, 1875 New Zealand short-tailed bats 2
Thyropteridae
Miller, 1907
Disk-winged bats 5
Furipteridae J. E. Gray, 1866 Smoky bat and thumbless bat 2
Noctilionidae
J. E. Gray, 1821
Bulldog bats 2
Mormoopidae Saussure, 1860 Ghost-faced, naked-backed and mustached bats 18
Phyllostomidae
J. E. Gray, 1825
New World leaf-nosed bats 217
Vespertilionoidea J. E. Gray, 1821
Family English Name Number of Species Image Figure
Natalidae J. E. Gray, 1825 Funnel-eared bats 10
Molossidae
Gervais in de Castelnau, 1855
Free-tailed bats 126
Miniopteridae
Dobson, 1875
Long-fingered bats 38
Cistugidae
Lack et al., 2010
Wing-gland bats 2
Vespertilionidae J. E. Gray, 1821 Vesper bats 496

Anatomy and physiology

Skull and dentition

A preserved megabat showing how the skeleton fits inside its skin

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".[46] Among microbats, longer snouts are associated with nectar-feeding.[47] while vampire bats have reduced snouts to accommodate large incisors and canines.[48]

Small insect-eating bats can have as many as 38 teeth, while vampire bats have only 20. 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.[47] The upper incisors of vampire bats lack enamel, which keeps them razor-sharp.[48] The bite force of small bats is generated through mechanical advantage, allowing them to bite through the hardened armour of insects or the skin of fruit.[49]

Wings and flight

Bats are the only mammals capable of sustained flight, as opposed to

gliding, as in the flying squirrel.[50] The fastest bat, the Mexican free-tailed bat (Tadarida brasiliensis), can achieve a ground speed of 160 km/h (100 mph).[51]

Little brown bat take off and flight

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

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 specialized 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 fifty million years.[54] 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.[55]

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.[56] Due to their flexible joints, bats are more maneuverable and more dexterous than gliding mammals.[57]

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 maneuver more accurately than the latter, and fly with more lift and less drag.

stalls.[62] Insectivorous bats may also use tactile hairs to help perform complex maneuvers to capture prey in flight.[57]

The

elastic fibres, nerves, muscles, and blood vessels. The muscles keep the membrane taut during flight.[63] The extent to which the tail of a bat is attached to a patagium can vary by species, with some having completely free tails or even no tails.[47] The skin on the body of the bat, which has one layer of epidermis and dermis, as well as hair follicles, sweat glands and a fatty subcutaneous layer, is very different from the skin of the wing membrane. Depending on the bat species the presence of hair follicles and sweat glands will vary in the patagium.[64] This patagium is an extremely thin double layer of epidermis; these layers are separated by a connective tissue center, rich with collagen and elastic fibers. In some bat species sweats glands will be present in between this connective tissue.[65] Furthermore, if hair follicles are present this supports the bat in order to adjust sudden flight maneuvers.[66][67] For bat embryos, apoptosis (programmed cell death) affects only the hindlimbs, while the forelimbs retain webbing between the digits that forms into the wing membranes.[68] Unlike birds, whose stiff wings deliver bending and torsional stress to the shoulders, bats have a flexible wing membrane that can resist only tension. 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.[56]

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 stabilized by the animal changing its wing curvatures.[69]

Roosting and gaits

Group of megabats roosting

When not flying, bats hang upside down from their feet, a posture known as roosting.

cervical or neck vertebrae in the two groups, which are clearly distinct.[70] 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.[71]

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.[72]

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.[73][74] Since their bodies are relatively small and lightweight, bats are not at risk of blood flow rushing to their heads when roosting.[75]

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.[76] During flight the respiratory cycle has a one-to-one relationship with the wing-beat cycle.[77] Because of the restraints of the mammalian lungs, bats cannot maintain high-altitude flight.[56]

The wings are highly vascularized membranes, the larger blood vessels visible against the light.[78]

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 requires 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

beats per minute.[81]

With its extremely thin membranous tissue, a bat's wing can significantly contribute to the organism's total gas exchange efficiency.[82] 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.[82] The subcutaneous vessels in the membrane lie very close to the surface and allow for the diffusion of oxygen and carbon dioxide.[83]

The

malt sugar, in their intestinal tract. Nectivorous and frugivorous bats have more maltase and sucrase enzymes than insectivorous, to cope with the higher sugar contents of their diet.[85]

The adaptations of the

cutaneous evaporative water loss.[78] Water helps maintain their ionic balance in their blood, thermoregulation system, and removal of wastes and toxins from the body via urine. They are also susceptible to blood urea poisoning if they do not receive enough fluid.[86]

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.[87]

Senses

Echolocation

Microbats and a few megabats emit ultrasonic sounds to produce echoes. Sound intensity of these echos are dependent on subglottic pressure. The bats' cricothyroid muscle controls the orientation pulse frequency, which is an important function. This muscle is located inside the larynx and it is the only tensor muscle capable of aiding phonation.

decibels.[90] Microbats use their larynx to emit echolocation signals through the mouth or the nose.[91] Microbat calls range in frequency from 14,000 to well over 100,000 Hz, extending well beyond the range of human hearing (between 20 and 20,000 Hz).[92] Various groups of bats have evolved fleshy extensions around and above the nostrils, known as nose-leaves, which play a role in sound transmission.[93]

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

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.[94] The delay of the returning echoes allows the bat to estimate the range to their prey.[92] In high-duty cycle echolocation, bats emit a continuous call and separate pulse and echo in frequency using the Doppler effect of their motion in flight. The shift of the returning echoes yields 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.[94][95]

In addition to echolocating prey, bat ears are sensitive to sounds made by their prey, such as the fluttering of moth wings. 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

interference patterns from the echoes reflecting from the tragus, a flap of skin in the external ear.[92]

The tiger moth (Bertholdia trigona) can jam bat echolocation.[99][100]

By repeated scanning, bats can mentally construct an accurate image of the environment in which they are moving and of their prey.

aposematic ultrasound signals to warn bats that they are chemically protected and therefore distasteful.[99][100] Moth species including the tiger moth can produce signals to jam bat echolocation. Many moth species 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.[102][103][104]

Vision

The eyes of most microbat species are small and poorly developed, leading to poor visual acuity, but no species is blind.[105] Most microbats have mesopic vision, meaning that they can detect light only in low levels, whereas other mammals have photopic vision, which allows colour vision. Microbats may use their vision for orientation and while travelling between their roosting grounds and feeding grounds, as echolocation is effective only 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.[50][106][107][108]

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.[108]

Magnetoreception

Microbats make use of magnetoreception, in that they have a high sensitivity to the Earth's magnetic field, as birds do. 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.[109][110]

Thermoregulation

Thermographic image of a bat using trapped air as insulation

Most bats are

thermal conductivity. 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.[113]

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;

Pteropus hypomelanus uses saliva and wing-fanning to cool itself while roosting during the hottest part of the day.[115] Among microbats, the Yuma myotis (Myotis yumanensis), the Mexican free-tailed bat, and the pallid bat (Antrozous pallidus) cope with temperatures up to 45 °C (113 °F) by panting, salivating, and licking their fur to promote evaporative cooling; this is sufficient to dissipate twice their metabolic heat production.[116]

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.[117]

Torpor

tricoloured bat (Perimyotis subflavus) in torpor

Torpor, a state of decreased activity where the body temperature and metabolism decreases, is especially useful for bats, as they use a large amount of energy while active, depend upon an unreliable food source, and have a limited ability to store fat. They generally drop their body temperature in this state to 6–30 °C (43–86 °F), and may reduce their energy expenditure by 50 to 99%.[118] Tropical bats may use it to avoid predation, by reducing the amount of time spent on foraging and thus reducing the chance of being caught by a predator.[119] Megabats were generally believed to be homeothermic, but three species of small megabats, with a mass of about 50 grams (1+34 ounces), have been known to use torpor: the common blossom bat (Syconycteris australis), the long-tongued nectar bat (Macroglossus minimus), and the eastern tube-nosed bat (Nyctimene robinsoni). Torpid states last longer in the summer for megabats than in the winter.[120]

During hibernation, bats enter a torpid state and decrease their body temperature for 99.6% of their hibernation period; even during periods of arousal, when they return their body temperature to normal, they sometimes enter a shallow torpid state, known as "heterothermic arousal".[121] Some bats become dormant during higher temperatures to keep cool in the summer months.[122]

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.[123][124]

Size

The smallest bat is

extant species of mammal, next to the Etruscan shrew.[126] The largest bats are a few species of Pteropus megabats and the giant golden-crowned flying fox, (Acerodon jubatus), which can weigh 1.6 kg (3+12 lb) with a wingspan of 1.7 m (5 ft 7 in).[127] Larger bats tend to use lower frequencies and smaller bats higher for echolocation; high-frequency echolocation is better at detecting smaller prey. Small prey may be absent in the diets of large bats as they are unable to detect them.[128] The adaptations of a particular bat species can directly influence what kinds of prey are available to it.[129]

Ecology

Tent-making bats (Uroderma bilobatum) in Costa Rica

Flight has enabled bats to become one of the most widely distributed groups of mammals.

crepuscular.[136][137] Microbats are known to exhibit diurnal behaviour in temperate regions during summer when there is insufficient night time to forage,[138][139] and in areas where there are few avian predators during the day.[140][141]

In temperate areas, some microbats migrate hundreds of kilometres to winter hibernation dens;[142] others pass into torpor in cold weather, rousing and feeding when warm weather allows insects to be active.[143] Others retreat to caves for winter and hibernate for as much as six months.[143] Microbats rarely fly in rain; it interferes with their echolocation, and they are unable to hunt.[144]

Food and feeding

Bats feeding on insects over a lake

Different bat species have different diets, including insects, nectar, pollen, fruit and even vertebrates.

Euderma maculatum), in search of food.[147] Bats use a variety of hunting strategies.[128] Bats get most of their water from the food they eat; many species also drink from water sources like lakes and streams, flying over the surface and dipping their tongues into the water.[148]

The Chiroptera as a whole are in the process of losing the ability to synthesise vitamin C.[149] In a test of 34 bat species from six major families, including major insect- and fruit-eating bat families, all were found to have lost the ability to synthesise it, and this loss may derive from a common bat ancestor, as a single mutation.[150][b] At least two species of bat, the frugivorous bat (Rousettus leschenaultii) and the insectivorous bat (Hipposideros armiger), have retained their ability to produce vitamin C.[151]

Insects

Most microbats, especially in temperate areas, prey on insects.

caddisflies.[47][153][154] Large numbers of Mexican free-tailed bats (Tadarida brasiliensis) fly hundreds of metres above the ground in central Texas to feed on migrating moths.[155] Species that hunt insects in flight, like the little brown bat (Myotis lucifugus), may catch an insect in mid-air with the mouth, and eat it in the air or use their tail membranes or wings to scoop up the insect and carry it to the mouth.[156][157] The bat may also take the insect back to its roost and eat it there.[158] Slower moving bat species, such as the brown long-eared bat (Plecotus auritus) and many horseshoe bat species, may take or glean insects from vegetation or hunt them from perches.[47] Insectivorous bats living at high latitudes have to consume prey with higher energetic value than tropical bats.[159]

Fruit and nectar

An Egyptian fruit bat (Rousettus aegyptiacus) carrying a fig

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.[160][161] The Jamaican fruit bat (Artibeus jamaicensis) has been recorded carrying fruits weighing 3–14 g (1812 oz) or even as much as 50 g (1+34 oz).[162]

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.[161][163] The tube-lipped nectar bat (Anoura fistulata) 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 the tongue retracts, it coils up inside the rib cage.[163] Because of these features, nectar-feeding bats cannot easily turn to other food sources in times of scarcity, making them more prone to extinction than other types of bat.[164][165] Nectar feeding also aids a variety of plants, since these bats serve as pollinators, as pollen gets attached to their fur while they are feeding. Around 500 species of flowering plant rely on bat pollination and thus tend to open their flowers at night.[161] Many rainforest plants depend on bat pollination.[166]

Vertebrates

The greater noctule bat (Nyctalus lasiopterus) uses its large teeth to catch birds.[167]

Some bats prey on other vertebrates, such as fish, frogs, lizards, birds and mammals.

Noctilio leporinus) hunt fish. They use echolocation to detect small ripples on the water's surface, swoop down and use specially enlarged claws on their hind feet to grab the fish, then take their prey to a feeding roost and consume it.[170] At least two species of bat are known to feed on other bats: the spectral bat (Vampyrum spectrum), and the ghost bat (Macroderma gigas).[171]

Blood

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

A few species, specifically the common, white-winged, and hairy-legged vampire bats, feed only 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.[172] Vampire bats target sleeping prey and can detect deep breathing.[173] Heat sensors in the nose help them to detect blood vessels near the surface of the skin.[174] They pierce the animal's skin with their teeth, biting away a small flap,[175] and lap up the blood with their tongues, which have lateral grooves adapted to this purpose.[176] The blood is kept from clotting by an anticoagulant in the saliva.[175]

Predators, parasites, and diseases

Bats are subject to predation from

falcons, and at roosts from terrestrial predators able to climb, such as cats.[177] Low-flying bats are vulnerable to crocodiles.[178] Twenty species of tropical New World snakes are known to capture bats, often waiting at the entrances of refuges, such as caves, for bats to fly past.[179] J. Rydell and J. R. Speakman argue that bats evolved nocturnality during the early and middle Eocene period to avoid predators.[177] The evidence is thought by some zoologists to be equivocal so far.[180]

white nose syndrome

As are most mammals, bats are hosts to a number of internal and external parasites.

lice, possibly due to competition from more specialised parasites that occupy the same niche.[183]

mid-Atlantic states have, since 2006, witnessed entire species completely extirpated and others with numbers that have gone from the hundreds of thousands, even millions, to a few hundred or less.[187] Nova Scotia, Quebec, Ontario, and New Brunswick have witnessed identical die offs, with the Canadian government making preparations to protect all remaining bat populations in its territory.[188] Scientific evidence suggests that longer winters where the fungus has a longer period to infect bats result in greater mortality.[189][190][191] In 2014, the infection crossed the Mississippi River,[192] and in 2017, it was found on bats in Texas.[193]

Bats are

zoonotic viruses (which can be transmitted to humans) than other mammal groups, though the differences among the aforementioned three groups were not significant (bats have no more zoonotic viruses than rodents and primates).[206] Another 2020 review of mammals and birds found that the identity of the taxonomic groups did not have any impact on the probability of harboring zoonotic viruses. Instead, more diverse groups had greater viral diversity.[207]

They seem to be highly resistant to many of the pathogens they carry, suggesting a degree of adaptation to their immune systems.

severe acute respiratory syndrome (SARS) in China, since they serve as natural hosts for coronaviruses, several from a single cave in Yunnan, one of which developed into the SARS virus.[198][210][211] However, they neither cause nor spread COVID-19.[212]

Behaviour and life history

Social structure

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

Some bats lead solitary lives, while others live in colonies of more than a million.[213] For instance, the Mexican free-tailed bat fly for more than one thousand miles to the 100-foot (30 m) wide cave known as Bracken Cave every March to October which plays home to an astonishing twenty million of the species,[214] whereas a mouse-eared bat lives an almost completely solitary life.[215] Living in large colonies lessens the risk to an individual of predation.[47] Temperate bat species may swarm at hibernation sites as autumn approaches. This may serve to introduce young to hibernation sites, signal reproduction in adults and allow adults to breed with those from other groups.[216]

Several species have a

mutual grooming may occur in certain species, such as the common vampire bat (Desmodus rotundus), and these strengthen social bonds.[219][220] Homosexual fellatio has been observed in the Bonin Flying Fox Pteropus pselaphon[221] and the Indian Flying Fox Pteropus medius,[222]
though the function and purpose of this behaviour is not clear.

Communication

Acoustics of the songs of Mexican free-tailed bats[223]

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.[47][224] Mexican free-tailed bats are one of the few species to "sing" like birds. Males sing to attract females. Songs have three phrases: chirps, trills and buzzes, the former having "A" and "B" syllables. Bat songs are highly stereotypical but with variation in syllable number, phrase order, and phrase repetitions between individuals.[223] Among greater spear-nosed bats (Phyllostomus hastatus), females produce loud, broadband calls among their roost mates to form group cohesion. Calls differ between roosting groups and may arise from vocal learning.[225]

In a study on captive Egyptian fruit bats, 70% of the directed calls could be identified by the researchers as to which individual bat made it, and 60% could be categorised into four contexts: squabbling over food, jostling over position in their sleeping cluster, protesting over mating attempts and arguing when perched in close proximity to each other. The animals made slightly different sounds when communicating with different individual bats, especially those of the opposite sex.[226] In the highly sexually dimorphic hammer-headed bat (Hypsignathus monstrosus), males produce deep, resonating, monotonous calls to attract females. Bats in flight make vocal signals for traffic control. Greater bulldog bats honk when on a collision course with each other.[224]

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.[224]

Reproduction and life cycle

Group of polygynous vampire bats

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.[227][47] Promiscuity, where both sexes mate with multiple partners, exists in species like the Mexican free-tailed bat and the little brown bat.[228][229] There appears to be bias towards certain males among females in these bats.[47] In a few species, such as the yellow-winged bat and spectral bat, adult males and females form monogamous pairs.[47][230] Lek mating, where males aggregate and compete for female choice through display, is rare in bats[231] but occurs in the hammerheaded bat.[232]

For temperate living bats, mating takes place in late summer and early autumn.

delayed implantation, in which the egg is fertilised after mating, but remains free in the reproductive tract until external conditions become favourable for giving birth and caring for the offspring.[236] In another strategy, fertilisation and implantation both occur, but development of the foetus is delayed until good conditions prevail. During the delayed development the mother keeps the fertilised egg alive with nutrients. This process can go on for a long period, because of the advanced gas exchange system.[237]

Newborn common pipistrelle, Pipistrellus pipistrellus

For temperate living bats, births typically take place in May or June in the northern hemisphere; births in the southern hemisphere occur in November and December. Tropical species give birth at the beginning of the rainy season.

maternity colonies and may assist each other in birthing.[241][242][240]

Most of the care for a young bat comes from the mother. In monogamous species, the father plays a role. Allo-suckling, where a female suckles another mother's young, occurs in several species. This may serve to increase colony size in species where females return to their natal colony to breed.[47] A young bat's ability to fly coincides with the development of an adult body and forelimb length. For the little brown bat, this occurs about eighteen days after birth. Weaning of young for most species takes place in under eighty days. The common vampire bat nurses its offspring beyond that and young vampire bats achieve independence later in life than other species. This is probably due to the species' blood-based diet, which is difficult to obtain on a nightly basis.[243]

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 bats is three-and-a-half times longer than other mammals of similar size. Six species have been recorded to live over thirty years in the wild: the brown long-eared bat (Plecotus auritus), the little brown bat (Myotis lucifugus), the

hibernating; bats that hibernate, on average, have a longer lifespan than bats that do not.[245][246]

Another hypothesis is that flying has reduced their mortality rate, which would also be true for birds and gliding mammals. Bat species that give birth to multiple pups generally have a shorter lifespan than species that give birth to only a single pup. Cave-roosting species may have a longer lifespan than non-roosting species because of the decreased predation in caves. A male Siberian bat was recaptured in the wild after 41 years, making it the oldest known bat.[246][247]

Interactions with humans

Conservation

Conservation statuses of bats as of 2020 according to the IUCN (1,314 species in total)[248]

  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 the

Wildlife and Countryside Acts, and disturbing a bat or its roost can be punished with a heavy fine.[250]
In Sarawak, Malaysia, "all bats"[251] are protected under the Wildlife Protection Ordinance 1998,[251] but species such as the hairless bat (Cheiromeles torquatus) are still eaten by the local communities.[252] Humans have caused the extinction of several species of bat in modern history, the most recent being the Christmas Island pipistrelle (Pipistrellus murrayi), which was declared extinct in 2009.[253]

Many people put up bat houses to attract bats.[254] The 1991 University of Florida bat house is the largest occupied artificial roost in the world, with around 400,000 residents.[255] In Britain, thickwalled and partly underground World War II pillboxes have been converted to make roosts for bats,[256][257] and purpose-built bat houses are occasionally built to mitigate damage to habitat from road or other developments.[258][259] Cave gates are sometimes installed to limit human entry into caves with sensitive or endangered bat species. The gates are designed not to limit the airflow, and thus to maintain the cave's micro-ecosystem.[260] Of the 47 species of bats found in the United States, 35 are known to use human structures, including buildings and bridges. Fourteen species use bat houses.[261]

the lungs of birds, making them more liable to fatal rupture.[264][265][266][267][268] Bats may be attracted to turbines, perhaps seeking roosts, increasing the death rate.[264] Acoustic deterrents may help to reduce bat mortality at wind farms.[269]

The diagnosis and contribution of barotrauma to bat deaths near wind turbine blades have been disputed by other research comparing dead bats found near wind turbines with bats killed by impact with buildings in areas with no turbines.[270]

Cultural significance

Francisco de Goya
, The Sleep of Reason Produces Monsters, 1797

Since bats are mammals, yet can fly, they are considered to be

mythology, bats symbolised the land of the dead, destruction, and decay.[275][276][277] An East Nigerian tale tells that the bat developed its nocturnal habits after causing the death of his partner, the bush-rat, and now hides by day to avoid arrest.[278]

More positive depictions of bats exist in some cultures. In China, bats have been associated with happiness, joy and good fortune. Five bats are used to symbolise the "Five Blessings": longevity, wealth, health, love of virtue and peaceful death.

Zapotec civilisation of Mesoamerica, the bat god presided over corn and fertility.[281]

Zapotec bat god, Oaxaca, 350–500 CE

The

Weird Sisters in Shakespeare's Macbeth used the fur of a bat in their brew.[282] 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 villainous vampires, such as Count Dracula and before him Varney the Vampire,[283] and heroes, such as the DC Comics character Batman.[284] Kenneth Oppel's Silverwing novels narrate the adventures of a young bat,[285] based on the silver-haired bat of North America.[286]

The bat is sometimes used as a

state bat. Texas and Oklahoma are represented by the Mexican free-tailed bat, while Virginia is represented by the Virginia big-eared bat (Corynorhinus townsendii virginianus).[290]

Economics

Insectivorous bats in particular are especially helpful to farmers, as they control populations of agricultural pests and reduce the need to use

$3.7 billion to $53 billion per year in pesticides and damage to crops. This also prevents the overuse of pesticides, which can pollute the surrounding environment, and may lead to resistance in future generations of insects.[291]

Bat dung, a type of

saltpetre was collected from caves to make gunpowder. At the time, it was believed that the nitrate all came from the bat guano, but it is now known that most of it is produced by nitrifying bacteria.[293]

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. About 100,000 tourists a year visit the bridge at twilight to watch the bats leave the roost.[294]

See also

  • Bat detector – device used to detect the presence of bats

Explanatory notes

  1. Ancient Greek: χείρ – cheir, "hand" and πτερόν – pteron, "wing".[1]
  2. ^ Earlier reports that only fruit bats were deficient were based on smaller samples.[151]

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Sources

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