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Bat
Temporal range: Eocene – Present
Common vampire batHorseshoe batGreater short-nosed fruit batEgyptian fruit batMexican free-tailed batGreater mouse-eared bat
Clockwise from top right: Egyptian fruit bat (Rousettus aegyptiacus), mass of Mexican free-tailed bats (Tadarida brasiliensis), greater mouse-eared bat (Myotis myotis), greater short-nosed fruit bat (Cynopterus sphinx), horseshoe bat (Rhinolophus ferrumequinum), common vampire bat (Desmodus rotundus).
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Clade: Apo-Chiroptera
Order: Chiroptera
Blumenbach, 1779
Suborders
Worldwide distribution of bat species

Bats are mammals of the order Chiroptera (/kaɪˈrɒptərə/; from the Ancient Greek: χείρ – cheir, "hand" and πτερόν – pteron, "wing")[1] whose forelimbs form webbed 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, can only glide for short distances. Bats are less efficient at flying than birds, but are more manoeuvrable, using their very long spread-out digits which are covered with a thin membrane or patagium.

Bats are the second largest order of mammals (after the rodents), representing about 20% of all classified mammal species worldwide, with about 1,240 bat species divided into two suborders: the less specialized and largely fruit-eating megabats, including flying foxes, and the highly specialized and echolocating microbats. About 70% of bat species are insectivores. Most of the rest are frugivores, or fruit eaters. A few species feed from animals other than insects, with the vampire bats being hematophagous, or feeding on blood.

Bats are present throughout most of the world, with the exception of extremely cold regions. They perform the vital ecological roles of pollinating flowers and dispersing fruit seeds; many tropical plant species depend entirely on bats for the distribution of their seeds. Bats are economically important, as they consume insect pests, reducing the need for pesticides. The smallest bat, and arguably the smallest extant mammal, is 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. The largest bats are a few species of Pteropus (fruit bats or flying foxes) and the giant golden-crowned flying fox, Acerodon jubatus, with a weight up to 1.6 kg (4 lb) and wingspan up to 1.7 m (5 ft 7 in).

Bat dung has been mined as guano from caves and used as fertilizer. Bats are natural reservoirs of many diseases including rabies; since they are highly mobile, social, and long-lived, they can readily spread diseases. Bats are sometimes numerous enough to serve as tourist attractions, and are used as food across Asia and the Pacific Rim. In many cultures, bats are popularly associated with darkness, death, witchcraft and malevolence.

Etymology

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 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".[2][3]

The name "Chiroptera" derives from Ancient Greek: χείρ – cheir, "hand"[4] and πτερόν – pteron, "wing".[5][1]

Taxonomy

Classification

See also: List of bats and List of fruit bats

After rodents, bats are the largest order of mammals, making up about 20% of mammal species. There are 1,240 bat species which are traditionally recognized to belong to two suborders of bats: Megachiroptera (megabats), and the Microchiroptera (microbats/echolocating bats).[6] Not all megabats are larger than microbats.[7] Microbats use echolocation, but megabats do not, with the exception of the genus Rousettus.[8] Microbats lack the claw at the second finger of the forelimb.[9][10] The ears of microbats do not close to form a ring; the edges are separated from each other at the base of the ear.[10] Megabats eat fruit, nectar, or pollen. Most microbats eat insects; others may feed on fruit, nectar, pollen, fish, frogs, small mammals, or the blood of animals. Megabats have well-developed visual cortices and good visual acuity, while microbats rely on echolocation for navigation and finding prey.[6]

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

The 2011 classification of bats (which does not include Nycteridae as a separate family, nor the grouping of the Rhinolophoidea families and Pteropodidae into the Yinpterochiroptera) is:[11]

Phylogeny

Bats are placental mammals. They 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 evidence places bats in the superorder Laurasiatheria, with its sister taxon as Fereuungulata, which includes carnivorans, pangolins, odd-toed ungulates, even-toed ungulates, and cetaceans.[12][13][14][15][16] One study places Chiroptera as a sister taxon to odd-toed ungulates (Perissodactyla).[17]

Boreoeutheria
Laurasiatheria

 Eulipotyphla (hedgehogs, shrews, moles, solenodons)

Scrotifera

 Chiroptera (bats)

Fereuungulata
Ferae

 Pholidota (pangolins)

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

Euungulata

 Perissodactyla (horses, tapirs, rhinos)

 Cetartiodactyla (camels, ruminants, whales)

 Euarchontoglires (primates, colugos, treeshrews, rodents, rabbits)

Cladogram showing Chiroptera within Laurasiatheria, with Fereuungulata as its sister taxon[16]
Giant golden-crowned flying fox, Acerodon jubatus

The phylogenetic relationships of the different groups of bats have been the subject of much debate. The traditional subdivision between 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 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.[18] In the 1980s, a hypothesis based on morphological evidence was offered that stated the Megachiroptera evolved flight separately from the Microchiroptera. The so-called 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 that link them to primates. Although recent genetic studies strongly support the monophyly of bats,[18] debate continues as to the meaning of available genetic and morphological evidence.[19]

Genetic evidence indicates that megabats originated during the early Eocene and should be placed within the four major lines of microbats.[16] Consequently, two new suborders based on molecular data have been proposed. The new suborder of Yinpterochiroptera includes the Pteropodidae, or megabat family, as well as the families Rhinolophidae, Hipposideridae, Craseonycteridae, Megadermatidae, and Rhinopomatidae[20] The other new suborder, Yangochiroptera, includes all of the remaining families of bats (all of which use laryngeal echolocation), a conclusion supported by a 15-base-pair deletion in BRCA1 and a seven-base-pair deletion in PLCB4 present in all Yangochiroptera and absent in all Yinpterochiroptera.[20] One phylogenomic study showed that the two new proposed suborders were supported by analyses of thousands of genes.[16]

The molecular phylogeny of the Chiroptera is controversial, as it points to a microbat paraphyly, 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.[21] 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.[22] Analyses of the sequence of the "vocalization" gene, FoxP2, were inconclusive as to whether laryngeal echolocation was secondarily lost in the pteropodids or independently gained in the echolocating lineages.[23] However, analyses of the "hearing" gene, Prestin, seemed to favor the idea that echolocation developed independently at least twice, rather than there being a secondary loss in the pteropodids.[24]

Chiroptera
Microchiroptera
Rhinolophoidea

Megadermatidae (false vampire bats)

Craseonycteridae (Kitti's hog-nosed bat) File:Craseonycteris thonglongyai.JPG

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)

Natalidae (funnel-eared bats)

Vespertilionidae (vesper bats)

Internal relationships of the Chiroptera, excluding Nycteridae and Cistugidae[11][25]

Fossil record

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

Little fossil remains of bats exist, as their delicate skeletons do not fossilize very well. Only an estimated 12% of the bat fossil record is complete at the genus level.[26] Most of the oldest known bat fossils were already very similar to modern microbats. Archaeopteropus, formerly classified as the earliest known megachiropteran, is now classified as a microchiropteran.[18] The extinct bats Palaeochiropteryx tupaiodon and Hassianycteris kumari are the first fossil mammals to have their coloration discovered: both were reddish-brown.[27][28]

The 2003 discovery of an intermediate fossil bat from the 52 million year old Green River Formation, Onychonycteris finneyi, indicates that flight evolved before echolocative abilities.[29][30] 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 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 likely alternated between flaps and glides while in the air.[18] 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 time climbing or hanging on the branches of trees.[31] The distinctive features noted on the Onychonycteris fossil also support the claim that mammalian flight most likely evolved in arboreal gliders, rather than terrestrial runners. This model of flight development, commonly known as the "trees-down" theory, implies that bats attained powered flight by taking advantage of height and gravity, rather than relying on running speeds fast enough for a ground-level take off.[32]

Echolocation probably first derived in bats from communicative calls. The Eocene bats Icaronycteris and Palaeochiropteryx had cranial adaptations suggesting an ability to detect ultrasound, implying they used echolocation. This may have been used at first mainly for communicative purposes or for mapping out their surroundings during their gliding phase, only being used for hunting insects while foraging on the ground or among vegetation. After the adaptation of flight was established, it may have been refined to target flying prey.[31] Bats may have evolved echolocation through a shared common ancestor, in which case it was then lost in the Old World fruit bats, only to be regained in the horseshoe bats; or, echolocation evolved independently in both the Yinpterochiroptera and Yangochiroptera lineages.[33]

Distribution and habitat

Tent-making bats (Uroderma bilobatum) in Costa Rica

Flight has enabled bats to become one of the most widely distributed groups of mammals.[34] Apart from the high Arctic, the Antarctic and a few isolated oceanic islands, bats exist all over the world.[35] Bats are found in almost every habitat available on Earth. Different species select different habitats 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. Most temperate species additionally need a relatively warm hibernation shelter. Bat roosts can be found in hollows, crevices, foliage, and even human-made structures, and include "tents" the bats construct by biting leaves.[36]

Adaptations

Flight

Slow-motion and normal speed of Egyptian fruit bats flying

Bats are the only mammals that can truly fly, as opposed to gliding mammals such as the flying squirrel.[37] The fastest bat, the Mexican free-tailed bat (Tadarida brasiliensis), has a ground speed of 160 kilometres per hour (99 mph).[38]

Bones

A preserved fruit bat showing how the skeleton fits inside its skin

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 minerals, such as 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 signaling, 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 volant locomotion. The relative proportion of extant bat forelimb digits compared with those of Eocene fossil bats have no significant differences in relative digit proportion, suggesting that bat wing morphology has been conserved for over 50 million years.[39] During flight, the bones take on bending and shearing stress; and the bending stresses felt are smaller than terrestrial mammals, however the shearing stress is larger. The wing bones of bats have a slightly lower breaking stress point than those of birds.[40]

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, 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 a strong drag force, without having to be thick like they were 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.[41] Due to their flexible joints, bats are more manoeuvrable and more dextrous than gliding mammals.[42]

Bats are adapted to roosting, hanging upside down from their feet.[43] The femurs are attached at the hips in a way that allows them to bend outward and upward in flight. The ankle joint can flex so as to allow the trailing edge of the wings to bend downwards. However, this design does not permit many movements other than hanging or clambering up trees.[41] Most megabats roost with the head tucked towards the belly, whereas most microbats roost with the neck curled towards the back. This difference is reflected in the structure of the cervical vertebrae in the two groups, which are clearly distinct.[43] 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.[44]

Wings

See also: Bat wing development
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.[45] By folding the wings in toward their bodies on the upstroke, they save 35 percent energy during flight.[46] The membranes are also delicate, ripping easily;[47] however, the tissue of the bat's membrane is able to regrow, such that small tears can heal quickly.[47][48] The surface of their wings is equipped with touch-sensitive receptors on small bumps called Merkel cells, also found on human fingertips. These sensitive areas are different in bats, as each bump has a tiny hair in the center, making it even more sensitive and allowing the bat to detect and adapt to changing airflow; the primary use is to judge what the most efficient speed to fly at is, and possibly also to avoid stalls.[49] Insectivorous bats may also use tactile hairs while performing complex manouvres while attempting to capture prey in-flight.[42]

The patagium is the wing membrane. The patagium is stretched between the arm and hand bones, down the lateral side of the body and down to the hind limbs.[50] This skin membrane consists of connective tissue, elastic fibers, nerves, muscles, and blood vessels. The muscles keep the membrane taut during flight.[50] The skin on the body of the bat, which has one layer of epidermis and dermis, as well as the presence of hair follicles, sweat glands and a fatty subcutaneous layer, is very different from the skin of the wing membrane. The patagium is an extremely thin double layer of epidermis; these layers are separated by a connective tissue center, rich with collagen and elastic fibers. The membrane has no hair follicles or sweat glands, except between the fingers.[49][51] Unlike birds whose stiff wings deliver bending and torsional stress to the shoulders, bats have a flexible wing membrane which can only resist 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. However, this adaptation does not permit bats to reduce their wingspan as birds do, which means they cannot travel over long distances like birds can.[41]

Nectar- and pollen-eating bats are able to hover, in a similar way to hummingbirds. The sharp leading edges of the wings can create vortexes which provide lift. The vortex may be stabilized by the animal changing its wing curvatures.[52]

Gas exchange

Due to this extremely thin membranous tissue, a bat's wing can significantly contribute to the organism's total gas exchange efficiency.[51] 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 wing in an open/spread out position 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 amount of gas exchange.[53] The subcutaneous vessels in the membrane very close to the surface allow for the diffusion of oxygen and carbon dioxide.[54]

Circulatory system

Bats 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.[55][56] Due to their relatively small and lightweight bodies, bats are not at risk of blood flow rushing to their heads when roosting.[57]

Bats possess highly adapted lung systems to cope with the pressures of powered-flight. Flight is an energetically taxing aerobic activity and requires large amounts of oxygen to be sustained. In bats, the relative alveolar surface area and pulmonary capillary blood volume are significantly larger than most other small quadrupedal mammals.[58] Due to the restraints of the mammalian lungs, bats cannot maintain high-altitude flight.[41]

Senses

Echolocation

Main article: Animal echolocation § Bats
Pipistrellus Pulses
Recording of Pipistrellus pipistrellus bat time-expanded echolocation calls and social call.
Problems playing 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. Bat calls are some of the most intense, airborne animal sounds, and can range in intensity from between 60 and 140 decibels.[59][60] Microbats use their larynx to create ultrasound, and emit the sound through their mouth and sometimes their nose. The latter is most pronounced in the horseshoe bats (Rhinolophus spp.). Microbat calls range in frequency from 14,000 to well over 100,000 Hz, extending beyond the range of human hearing (between 20 and 20,000 Hz).[61]

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. This is important because these 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 clearly hear the returning echo.[62] The delay of the returning echoes provides the bat with the ability to estimate the range to their prey.[61]

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. The Doppler 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.[63]

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 not only echolocation signals, but also 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 may be seen in a large variety of unrelated animals, such as the aye-aye, lesser galago, bat-eared fox, mouse lemur, and others.[64][65][66] Bats can estimate the elevation of their target using the interference patterns from the echoes reflecting from the tragus, a flap of skin in the external ear.[61]

The tiger moth (Bertholdia trigona) can jam bat echolocation[67][68]

By repeated scanning, bats can mentally construct an accurate image of the environment in which they are moving and of their prey item.[69] However, some species of moth have exploited this, such as the tiger moths which produces aposematic ultrasound signals to warn bats that they are chemically protected.[67][68] Other moth species 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 maneuvers.[70][71]

Vision

Although the eyes of most microbat species are small and poorly developed, leading to poor visual acuity, no species is blind.[72] Microbats have mesopic vision, meaning that they can only detect light in low levels, whereas other mammals have photopic vision, which allows colour vision. Microbats may use their vision for orientation and while they are travelling between their roosting grounds and their 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.[37][73][74][75]

Megabat species often have eyesight as good as, if not better than, human vision. Their eyesight, unlike that of their microbat relatives, is adapted to both night and daylight vision including some colour vision.[75]

Magnetic field

Microbats possess magnetoreception, in that they have a high sensitivity to Earth's magnetic field, similar to birds. However, microbats use a polarity-based compass, meaning that they differentiate north from south, as opposed to birds which use the strength of the magnetic field to differentiate latitudes, which may be used in long-distance travel. Since microbats generally have poor eyesight, it is thought they use a magnetite-based method for orientation.[76][77]

Thermoregulation

Thermographic image of a bat using trapped air as insulation

Most bats are homeothermic, the exception being the Vespertilionidae, the Rhinolophidae and the Miniopteridae which extensively use heterothermy. Compared to other mammals, bats have a high thermal conductivity. Body heat is mainly lost through the wings as they are filled with blood vessels, but they may be used as an insulator while resting. By wrapping their wings around themselves, they can trap a layer of still air around themselves. Smaller bats generally have a higher metabolic rate than larger bats, and so need to consume more food in order to maintain homeothermy.[78]

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.[79] Unlike birds which have air sacs or other mammals which have sweat glands, bats have no means to cool themselves by evaporation, though they may use saliva to cool themselves in an emergency.[41]

Bats also possess a system of sphincter valves on the arterial side of the vascular network that runs along the edge of their wings. In the fully open state, these allow oxygenated blood to flow through the capillary network across the wing membrane, but when contracted, they shunt flow directly to the veins, bypassing the wing capillaries. This allows bats to control the amount of heat exchanged through the thin flight membrane, allowing them to release heat during flight. Many other mammals use the capillary network in oversized ears for the same purpose.[80]

Torpor

See also: Torpor
A tricoloured bat (Perimyotis subflavus) in torpor

Torpor 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. They generally drop their body temperature in this state to 6–30 °C (43–86 °F), and they may reduce their energy expenditure by 50 to 99%. Around 97% of all microbats use torpor,[81] including tropical bats which may use torpor to avoid predation.[82] Megabats were generally believed to be only homeothermic, however three species of small megabats, with a body mass of about 50 grams (1.8 oz), 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.[83]

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". These adaptations are probably used to decrease the energy costs.[84] Some bats may also aestivate to keep cool in hot summer months.[85]

To conserve energy, heterothermic bats during long migrations may go into a torpid state while roosting in the daytime, and flying at night. Unlike migratory birds which fly during the day and feed during the night, nocturnal bats have a conflict between traveling and eating. Using torpor, bats can save around 90% of energy that they would have spent trying to keep their body temperature at a normal level; thereby reducing the need to feed. It also decreases the duration of migration, which may prevent them from spending too much time in unfamiliar places, and decrease predation. Pregnant individuals of some species may not use torpor.[86][87]

Size

The smallest bat is Kitti's hog-nosed bat, measuring 29–34 millimetres (1.1–1.3 in) in length, 15 centimetres (5.9 in) wingspan and 2–2.6 grams (0.071–0.092 oz) in mass.[88][89] It is also arguably the smallest extant species of mammal, next to the Etruscan shrew.[90] The largest species of bat are a few species of Pteropus megabats and the giant golden-crowned flying fox with a weight up to 1.6 kilograms (3.5 lb) and wingspan up to 1.7 metres (5.6 ft).[91] Larger bats tend to use low-frequency echolocation, and smaller bats high-frequency echolocation, as high-frequency echolocation is more adept at detecting smaller prey. In general with other animals, larger species consume larger prey and smaller species consume higher prey; however in the case of bats, species that use high-frequency echolocation consume smaller prey, and species that use low-frequency echolocation consume larger prey, as low-frequency echolocation does not detect smaller prey items. Small prey may be absent in the diets of large bats as they are unable to detect them.[92] The adaptations in a particular bat species can directly influence what kinds of prey are available to it.[93]

Behaviour and life history

Mexican free-tailed bats on their long aerial migration between Oregon and Mexico

Most microbats are nocturnal[94] while megabats are typically diurnal or crepuscular.[95][96] In temperate areas, bats may migrate hundreds of kilometres to winter hibernation dens,[97] while some pass into torpor in cold weather, rousing and feeding when warm weather allows for insects to be active.[98] Others retreat to caves for winter and hibernate for six months.[98] Microbats rarely fly in rain, it interferes with their echolocation, and they are unable to locate their food.[99] A few species such as the New Zealand short-tailed bat and the common vampire bat are agile on the ground.[100]

Bracken Bat Cave, home to 20 million Mexican free-tailed bats

The social structure of bats varies, with some leading solitary lives and others living in colonies of more than a million bats.[101] Living in large colonies lessens the risk of an individual to predation.[102] Temperate bat species may swarm at hibernation sites as fall apporaches. This may serve to introduce young to hibernation sites, signal reproduction in adults and allow adults with breed with individuals from other groups.[103] The fission-fusion social structure is seen among several species of bats. The term "fusion" refers to a large numbers of bats that congregate in one roosting area, and "fission" refers to breaking up and the mixing of subgroups. Within these societies, bats are able to maintain long term relationships.[104] Some of these relationships consist of matrilineally related females and their dependent offspring.[105] Food sharing and mutual grooming may occur in certain species, such as the common vampire bat (Desmodus rotundus), and these function to strengthen social bonds.[106][107]

Food and feeding

Different bat species have different diets including insects, nectar, pollen, fruit and even vertebrates.[108] Megabats are mostly fruit, nectar and pollen eaters.[95] Due to their small size, high-metabolism and rapid burning of energy though flight, bats must consume large amounts of food for their size. Insectivorous bats may eat over 120 percent of their body weight while frugivorous bats may eat over twice their weight.[109] Predatory bats typically hunt at night, reducing competition with birds, and minimizing contact with certain predators. They can travel large distances, up to 800 kilometres (500 mi), in search of food.[37] Bats are not restricted to any one hunting strategy, but rather use a mix.[92] The bite force of small bats is generated through mechanical advantage, allowing them to bite through the hardened armor of insects or the skin of fruit.[110] Bats get most of their water needs from the food they eat. However, numerous species may drink from water sources like lakes and streams. They may fly over the water surface and dip their tongues.[111]

The Chiroptera as a whole are in the process of losing the ability to synthesize vitamin C: most have lost it completely.[112] In a test of 34 bat species from six major families of bats, including major insect- and fruit-eating bat families, all were found to have lost the ability to synthesize it, and this loss may derive from a common bat ancestor, as a single mutation.[113] Earlier reports of only fruit bats being deficient were based on smaller samples. However, recent results show that there are at least two species of bat, the frugivorous bat (Rousettus leschenaultii) and insectivorous bat (Hipposideros armiger), that have retained their ability to produce vitamin C.[114]

Insects

Most microbats, especially in temperate areas, prey on insects.[108] The diet of an insectivorous bat may span a wide range of species,[115] including flies, beetles, moths, grasshoppers, crickets, termites, bees, wasps, mayflies and caddisflies.[102] Large numbers of Mexican free-tailed bats (Tadarida brasiliensis) fly hundreds of meters above the ground in central Texas to feed on migrating moths.[116] Species that hunt insects in flight, like the little brown bat (Myotis lucifugus), may catch an insect in mid-air with its 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.[117][118] The bat may also take the insect back to its roost and eat it there.[119] Slower moving bat species such as the brown long-eared bat (Plecotus auritus) and many horseshoe bat species, may take or glean insects from vegatation or hunt them from perches.[102] Insectivorous bats living at high latitudes have to consume prey with higher energetic value than tropical bats.[120]

Fruit and nectar

File:Choeronycteris mexicana, Mexican long-tongued bat (7371567444).jpg
Mexican long-tongued bat (Choeronycteris mexicana) drinking from a cactus

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 numerous species of plants depend on bats for seed dispersal.[121][122] The Jamaican fruit bat (Artibeus jamaicensis) has been recorded carrying fruits weighing 3–14 g (0.11–0.49 oz) or even as much as 50 g (1.8 oz).[123]

An Egyptian fruit bat (Rousettus aegyptiacus) carrying a fig

Nectar-eating bats have acquired specialized adaptations. These bats possess long muzzles and long, extensible tongues covered in fine bristles that aid them in feeding on particular flowers and plants.[122][124] 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.[124] However, 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 any other type of bat.[125][126] Nectar feeding also aids a variety of plants, since these bats serve as pollinators (as pollen can get 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.[122] Rainforests are said to benefit the most from bat pollination, because of the large variety of plants that depend on it.[127]

Vertebrates

The greater noctule bat (Nyctalus lasiopterus) uses its large teeth to catch birds.[128]

Some bats prey on vertebrates, such as fish, frogs, lizards, birds and mammals.[102][129] The fringe-lipped bat (Trachops cirrhosus,) for example, is particularly skilled at catching frogs. 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.[130] Another example is the greater noctule bat, which can catch birds in flight.[128] Some species, like the greater bulldog bat (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.[131] At least two species of bat are known to feed on other bats: the spectral bat, also known as the American false vampire bat, and the ghost bat of Australia.[132]

Blood

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

A few species, specifically the common, white-winged, and hairy-legged vampire bats, exclusively consume animal blood (hematophagy). The common vampire bat typically feeds on large mammals such as cattle, while the hairy-legged and white-winged vampires feed on birds instead.[133] Vampire bats target sleeping prey and can detect deep breathing.[134] Heat sensors in the nose help them to detect blood vessels near the surface of the skin.[135] They pierce the animal's skin with their teeth, biting away a small flap,[136] and lap up the blood with their tongues, which have lateral grooves adapted to this purpose.[137] The blood is kept from clotting by an anticoagulant in the saliva.[136]

Reproduction and lifecycle

Bats employ a number of reproductive strategies. Most 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 that are unable to claim a site are forced to live on the periphery where they have less reproductive success.[138][102] Promiscuity, where both sexes mate with multiple partners, exists in species like the Mexican free-tailed bat and the little brown bat.[139][140] Nevertheless, there does appear to be bias towards certain males among females in these bats.[102] In a few species, such as the yellow-winged bat and spectral bat, adult males and females form monogamous pairs.[102][141] Lek mating, where males aggregate and compete for female choice though display, is rare in bats[142] but notably occurs in the highly sexually dimorphic hammer-headed bat (Hypsignathus monstrosus).[143]

Newborn common pipistrelle, Pipistrellus pipistrellus

For temperate living bats, mating takes place in late summer and early autumn.[144] Tropical bats may mate during the dry season.[145] After copulation, the male may leave behind a mating plug to insure his paternity. In hibernating species, males are known mate with females in torpor.[102] 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 is stored in the reproductive tract for several months after mating. While mating occurs in the fall 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.[146] In another strategy, fertilization and implantation both occur, but development of the fetus is delayed until favorable conditions prevail, during the delayed development the mother still gives the fertilized egg nutrients, and oxygenated blood to keep it alive. However, this process can go on for a long period of time, because of the advanced gas exchange system.[147]

Wahlberg's epauletted fruit bat (Epomophorus wahlberg) with young

For temperate living bats, births typically take place in May or June in the northern hemisphere while births in the southern hemisphere occur in November and December. Tropical species give birth at the beginning of the rainy season.[148] In most bat species, female carry and give birth to one pup per litter.[149] For bat embryos, apoptosis only affects the hindlimbs, while the forelimbs retain webbing between the digits which form into the wing membranes.[150] At birth, a bat pup can be up to 40 percent of the mother's weight,[102] and hence the pelvic girdle of the female expands during birth as the two halves are connected by a flexible ligament.[150] Females typically give birth in a heads-up or horizontal position, using gravity to make birthing easier. The young emerges rear-first, possibly to prevent the wings from getting tangled, and the females cradles it in her wing and tail membranes. In many species, females give birth and raise their young in maternity colonies and individuals may assist others in their birthing process.[151][146][150]

Most of the care for a young bat comes from the mother. However, in monogamous species, the father plays a role. In addition, allo-suckling, where a female suckles young that is not hers, occurs in several species. This may serve to increase colony size in species where females return to their natal colony to breed.[102] 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 likely due to the species' blood-based diet which is difficult to obtain on a nightly basis.[152]

Communication

Acoustics of the songs of Mexican free-tailed bats[153]

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 vast distances.[102][154] 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 among individuals.[153] 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.[155]

In captive Egyptian fruit bats, 70% of the directed calls could be identified as to which 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 individuals, especially one of the opposite sex.[156] Male hammerheaded bats produce deep, resonating, monotonous calls to attract females. Bats in flight make vocal signal for traffic control. Individual greater bulldog bats honk when on a collision course with each other.[154]

Bats also communicate by other means. Male little yellow-shouldered bats (Sturnira lilium) have shoulder glands which produce a spicy odor during the breeding season. Like many other species, they have hair specialized for retaining and dispersing secretions. Such hair forms a ruff around the necks of the some Old World megabat males, forming a conspicuous collar. 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 which they sprinkle on roost sites, a behaviour known as "salting". Salting may be accompanied by singing.[154]

Life expectancy

The maximum lifespan of bats is three-and-a-half times larger than other mammals of a similar size. Five species have been recorded living over 30 years in the wild: the brown long-eared bat (Plecotus auritus), the little brown bat (Myotis lucifugus), Brandt's bat (Myotis brandti), the lesser mouse-eared bat (Myotis blythii) and the greater horseshoe bat (Rhinolophus ferrumequinum). One hypothesis as to why bats can live so long is because they slow down their metabolic rate while hibernating, being consistent with the rate-of-living theory; bats that hibernate, on average, have a longer lifespan than bats that do not.[157][158] Another hypothesis is that flying has reduced their mortality rate, which would also be true for birds and gliding mammals. Bat species which 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. The oldest recorded bat is a 41-year-old male Brandt's bat.[158][159]

Conservation

See also: List of bats by population
Bat roost with Charles Augustus Rosenheimer Campbell in San Antonio, Texas in 1915

Groups such as the Organization for Bat Conservation and Bat Conservation International aim to increase awareness of bats' ecological roles and the environmental threats they face. 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, some bats are protected under the Wildlife Protection Ordinance 1998, but the hairless bat (Cheiromeles torquatus) 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. An estimated 100,000 tourists per year visit the bridge at twilight to watch the bats leave the roost.[160]

Many people put up bat houses to attract bats.[161] The 1991 University of Florida bat house is the largest occupied artificial roost in the world, with around 300,000 residents.[162][163] In Britain, thickwalled and partly underground World War II pillboxes have been converted to make roosts for bats,[164][165] and purpose-built bat houses are occasionally built to mitigate damage to habitat from road or other developments.[166][167]

Gates are sometimes installed at cave entrances 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.[168]

Threats

White nose syndrome

Main article: White nose syndrome
A little brown bat with white nose syndrome

White nose syndrome is a condition associated with the deaths of millions of bats in the Eastern United States and Canada.[169] The disease is named after a white fungus, Pseudogymnoascus destructans, found growing on the muzzles, ears, and wings of afflicted bats. This fungus, which is mostly spread from bat to bat, is the sole cause of the disease.[170] The fungus was first discovered in central New York State in 2006 and spread quickly to the entire Eastern US north of Florida; mortality rates of 90–100% have been observed in most caves.[171] New England and the 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.[172] The provinces of 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.[173] Scientific evidence suggests that longer winters where the fungus has a longer period of time to infect bats results in greater chances of mortality.[174][175][176] In 2014, infection crossed the Mississippi River,[177] but species native to northern Mexico and the West had not yet been affected.[178]

Use as food

Main article: Bat as food
Paniki is a Minahasan dish made from fruit bat meat.

Bats are eaten in countries across Asia and the Pacific Rim. In some cases, such as in Guam, flying foxes have become endangered through being hunted for food.[179]

Barotrauma and wind turbines

Evidence suggests that barotrauma is causing bat fatalities around wind turbines.[180] The lungs of bats are typical mammalian lungs, and are thought to be more sensitive to sudden air pressure changes than the lungs of birds, making them more liable to fatal rupture.[181][182][183][184][185] In addition, it has been suggested that bats are attracted to these structures, perhaps seeking roosts, and thereby increasing the death rate.[181] Acoustic deterrents may help to reduce bat mortality at wind farms.[186]

Interactions with humans

Guano

Main article: Guano
Aerial view of Guano Point, Arizona, site of a large accumulation of bat guano. Old tramway headhouse is at the end of dirt road (right). Bat Cave mine is 2,500 feet (760 m) below, across the canyon.

Bat dung, a type of guano, is rich in nitrates and is mined from caves for use as fertilizer.[187] During the U.S. Civil War, saltpeter was collected from caves to make gunpowder; it was thought that this was bat guano, but most of the nitrate comes from nitrifying bacteria.[188]

Disease transmission

Further information: Bat-borne virus

Bats are natural reservoirs for a large number of zoonotic pathogens,[189] including rabies,[190] histoplasmosis both directly and in guano,[191] Nipah Hendra viruses,[192] and possibly ebola virus.[193][194] Their high mobility, broad distribution, long life spans, substantial sympatry, and social behaviour make bats favourable hosts and vectors of disease. Compared to rodents, bats carry more zoonotic viruses per species, and each virus is shared with more species.[195] They seem to be highly resistant to many of the pathogens they carry, suggesting a degree of adaptation to bats' immune systems.[195][196][197] Furthermore, their interactions with livestock and pets, including predation by vampire bats, accidental encounters, and the scavenging of bat carcasses, compound the risk of zoonotic transmission.[198]

Among ectoparasites, bats carry fleas and mites, as well as specific parasites such as bat bugs and bat flies (Nycteribiidae and Streblidae).[199][200] However, they are one of the few non-aquatic mammalian orders that do not host lice. This may be due to competition from more effective, specialized parasites which occupy the same niche.[200]

They are also implicated in the emergence of SARS (severe acute respiratory syndrome), since they serve as a natural host for the type of virus involved (the genus Coronavirus, whose members typically cause mild respiratory disease in humans). A joint CAS/CSIRO team using phylogenetic analysis found that the SARS Coronavirus originated within the SARS-like Coronavirus group carried by the bat population in China.[201] However, note that they only served as the source of the precursor virus (which "jumped" to humans and evolved into the strain responsible for SARS): bats do not carry the SARS virus itself.[191]

Rabies

Bats present a significant hazard in areas where the rabies virus is endemic.[198] For example, in the United States, bats typically constitute around a quarter of reported cases of rabies in wild animals, and their bites account for the vast majority of cases in humans.[202][203] Rabies is fully preventable if the patient is vaccinated before the onset of symptoms, but bat bites are small and may remain unnoticed.[203] The most severe threat to humans and domestic animals comes from sick, downed, or dead bats.[198][191] Bat rabies virus can rarely infect victims purely through airborne transmission ("cryptic rabies").[204][205] All active widespread rabies strains appear to have evolved from strains endemic to bats. Through zoonosis, these mutated and "jumped" to other species. In North America, for example, this reportedly occurred in the mid-1600s.[198]

In certain countries, such as the United Kingdom, it is illegal to handle bats without a license and advice should be sought from an expert organisation, such as the Bat Conservation Trust, if a trapped or injured bat is found.[206]

Cultural significance

Francisco Goya, The Sleep of Reason Produces Monsters, 1797

In many cultures, including in Europe, bats are associated with darkness, death, witchcraft, and malevolence.[207] Because bats are mammals, yet can fly, they are liminal beings in many traditions.[208] Among Native Americans such as the Creek, Cherokee and Apache, the bat is a trickster spirit. In Tanzania, a winged bat cryptid known as Popobawa, is believed to be a shapeshifting evil spirit that assaults and sodomises its victims.[209] In Aztec mythology, bats symbolized the land of the dead, destruction, and decay.[210][211][212][213] 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.[214]

Zapotec bat god, Oaxaca, 350–500 AD

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.[215] The bat is sacred in Tonga and is often considered the physical manifestation of a separable soul.[216]

The Weird Sisters in Shakespeare's Macbeth used the fur of a bat in their brew.[217] 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, such as Dracula, and heroes, such as Batman.[218][219] Kenneth Oppel's Silverwing novels narrate the adventures of a young bat,[220] based on the silver-haired bat of North America.[221]

The bat is sometimes used as a heraldic symbol in Spain and France, appearing in the coats of arms of the towns of Valencia, Palma de Mallorca, Fraga, Albacete, and Montchauvet.[222][223][224] Three U.S. states have an official state bat. Texas and Oklahoma are represented by the Mexican free-tailed bat; Virginia is represented by the Virginia big-eared bat.[225]

See also

Notes

References

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Bibliography

  • Altringham, J. (2011). Bats: From Evolution to Conservation. Oxford University Press. ISBN 978-0199207114.
  • Fenton, M. B. (2001). Bats. Checkmark Books. ISBN 0-8160-4358-2.
  • Feton, M. B.; Simmons, N. B. (2015). Bats: A World of Science and Mystery. University of Chicago Press. ISBN 978-0226065120.{{cite book}}: CS1 maint: multiple names: authors list (link)
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