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===Vision===
===Vision===
Although the eyes of most microbat species are small and poorly developed, leading to poor [[visual acuity]], no species is blind.<ref>{{cite web|author=Nida Sophasarun |title=Experts debunk bats' bad rap |work=Online extra |publisher=National Geographic |url=http://ngm.nationalgeographic.com/ngm/0204/feature7/online_extra.html |accessdate=April 30, 2013}}</ref> Microbats use vision to navigate, especially for long distances when beyond the range of echolocation,<ref>{{cite news|author=Rachel Kaufman |title=Bats Crash More When They Rely on Vision |url=http://news.nationalgeographic.com/news/2010/11/101115-bats-vision-science-animals/ |newspaper=National Geographic Daily News |date=December 15, 2010 |accessdate=April 30, 2013}}</ref> and species that are gleaners—that is, ones that attempt to swoop down from above to ambush insects, like crickets on the ground or moths up a tree, often have eyesight about as good as a rat's. Some species have been shown to be able to detect [[ultraviolet]] light, and most cave-dwelling species have developed the ability to utilize very dim light.<ref name=molecule/>
Although the eyes of most microbat species are small and poorly developed, leading to poor [[visual acuity]], no species is blind.<ref>{{cite web|author=Nida Sophasarun |title=Experts debunk bats' bad rap |work=Online extra |publisher=National Geographic |url=http://ngm.nationalgeographic.com/ngm/0204/feature7/online_extra.html |accessdate=April 30, 2013}}</ref> 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 some microbats have distinct colourations, they may be able to discriminate colours.<ref name=molecule/><ref>{{cite journal|first= Brigitte|last=Müller|first2=Martin|last2=Glösmann|first3=Leo|last3=Peichl|first4=Gabriel C.|last4=Knop|first5=Cornelia|last5=Hagemann|first6=Josef|last6=Ammermüller|year=2009|title=Bat Eyes Have Ultraviolet-Sensitive Cone Photoreceptors|journal=PLoS One|volume=4|issue=7|doi=10.1371/journal.pone.0006390|pmc=2712075}}</ref><ref>{{cite journal|first=Yong-Yi|last=Shen|first2=Jie|last2=Liu|first3=David M.|last3= Irwin|first4=Ya-Ping|last4=Zhang|year=2010|title=Parallel and Convergent Evolution of the Dim-Light Vision Gene ''RH1'' in Bats (Order: Chiroptera)|journal=PLoS One|volume=5|issue=1|doi=10.1371/journal.pone.0008838|pmc= 2809114}}</ref><ref name=wang2004/>


Megabat species often have excellent eyesight as good as, if not better than, human vision. Their eyesight, unlike that of its microbat relatives, is adapted to both night and daylight vision which enables megabats to have some colour vision, whereas microbats see in blurred shades of grey.{{cn|date=September 2017}}
Megabat species often have excellent eyesight as good as, if not better than, human vision. Their eyesight, unlike that of its microbat relatives, is adapted to both night and daylight vision which enables megabats to have some colour vision.<ref name=wang2004>{{cite journal|last=Wang|first=D.|last2=Oakley|first2=T.|last3=Mower|first3=J.|last4=Shimmin|first4=L. C.|last5=Yim|first5=S.|last6=Honeycutt|first6=R. L. |last7=Tsao|first7=H.|last8=Li|first8=W. H.|year=2004|title=Molecular evolution of bat color vision genes|journal=Molecular Biology and Evolution|volume=21|issue=2|pages=295–302|doi=10.1093/molbev/msh015|pmid=14660703}}</ref>


===Thermoregulation===
===Thermoregulation===

Revision as of 00:40, 11 September 2017

For other uses, see Bat (disambiguation) and Bats (disambiguation).

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 Ancient Greek: 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, or 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, such as the fish-eating bat, 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 the Kitti's hog-nosed bat, measuring 29–34 mm (1.14–1.34 in) in length, 15 cm (5.91 in) across the wings and 2–2.6 g (0.07–0.09 oz) in mass. The largest species of bat are a few species of Pteropus (fruit bats or flying foxes) and the giant golden-crowned flying fox with a weight up to 1.6 kg (4 lb) and wingspan up to 1.7 m (5 ft 7 in).

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 Ancient Greek: πτερόν – pteron, "wing".[5])[1]

Taxonomy

Classification

"Chiroptera" from Ernst Haeckel's Kunstformen der Natur, 1904
Townsend's big-eared bat, Corynorhinus townsendii

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 show good visual acuity, while microbats rely on echolocation for navigation and finding prey.[6]

The classification of bats 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 studies have now placed 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 and flying foxes)

Fereuungulata
Ferae

 Pholidota (pangolins)

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

Euungulata

 Perissodactyla (horses, tapirs, rhinos)

 Cetartiodactyla (camels, pigs, ruminants, hippos, 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 reflects the view that these groups of bats have 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 proposes that, when adaptations to flight are removed, the Megachiroptera are allied to primates by anatomical features not shared with Microchiroptera. One example is that the brains of megabats show a number of advanced characteristics that link them to primates. Although recent genetic studies strongly support the monophyly of bats,[19] debate continues as to the meaning of available genetic and morphological evidence.[20]

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[21] 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.[21] 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.[22] 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.[23] 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.[24] However, analyses of the "hearing" gene, Prestin, seemed to favor the the idea that echolocation developed independently in species, rather than there being a secondary loss in the pteropodids.[25]

Chiroptera
Yinpterochiroptera

Pteropodidae (megabats)

Rhinolophidae (horseshoe bats)

Hipposideridae (Old World leaf-nosed bats)

Megadermatidae (false vampire bats)

Rhinopomatidae (mouse-tailed bats)

Yangochiroptera

Tadarida (part of "Molossidae", free-tailed bats)

Vespertilionidae (common or evening bats) File:Lasiurus intermedius.jpg

Taphozous (part of "Molossidae", free-tailed bats)

Internal relationships of the Chiroptera[11][26]

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.[27] 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 colouration discovered, both of a reddish-brown.[28][29]

The 2003 discovery of an intermediary fossil bat from the 52 million year old Green River Formation, Onychonycteris finneyi, indicates that flight evolved before echolocative abilities.[30][31] 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.[32] 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.[33]

Echolocation probably first derived in bats from communicative calls. The Eocene bats Icaronycteris and Palaeochiropteryx had some cranial adaptations suggesting an ability to detecting 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 more refined to target flying prey items.[32]

Distribution and habitat

Mexican free-tailed bats emerging from Carlsbad Caverns, New Mexico

Flight has enabled bats to become one of the most widely distributed groups of mammals.[34] Apart from the 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. In 2006, Sears et al. published a study that traces the elongation of manual bat digits, a key feature required for wing development, 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 offspring's manual digits. This crucial genetic alteration helps create the specialized limbs required for volant locomotion.[39] Sears et al. (2006) also studied the relative proportion of bat forelimb digits from several extant species and compared these with a fossil of Icaronycteris index, an early extinct species from approximately 50 million years ago. The study found no significant differences in relative digit proportion, suggesting that bat wing morphology has been conserved for over 50 million years.[39]

Wings

Image of patagium membrane of a microbat. Specimen from the Pacific Lutheran University Natural History collection.

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.[40] By folding the wings in toward their bodies on the upstroke, they save 35 percent energy during flight.[41] The membranes are also delicate, ripping easily;[42] however, the tissue of the bat's membrane is able to regrow, such that small tears can heal quickly.[42][43] 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 collect information about the air flowing over its wings, and to fly more efficiently by changing the shape of its wings in response. An additional kind of receptor cell is found in the wing membrane of species that use their wings to catch prey. This receptor cell is sensitive to the stretching of the membrane.[44] The cells are concentrated in areas of the membrane where insects hit the wings when the bats capture them.[citation needed]

The patagium is the wing's skin membrane. It covers and is strengthened by the bones of the bat's four long, thin digits, though the thumb is free-moving. The patagium is stretched between the arm and hand bones, down the lateral side of the body and down to the hind limbs.[45] This skin membrane consists of connective tissue, elastic filaments, nerves, muscles and blood vessels. The muscles keep the membrane taut during flight.[45] 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 skin 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 skin also does not have any hair follicles or sweat glands.[46]

Gas exchange

Due to this extremely thin membranous tissue, a bat's wing can significantly contribute to the organism's total gas exchange efficiency.[46] 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.[47] The subcutaneous vessels in the membrane very close to the surface allow for the diffusion of oxygen and carbon dioxide.[48]

Circulatory system

Scapulae, spine and ribs of Eptesicus fuscus (big brown bat)

Bats seem to make use of particularly strong venomotion (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.[49][50]

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 flight membrane (i.e. wing surface), but when contracted, they shunt flow directly to the veins, bypassing the wing capillaries. This is likely an important tool for thermoregulation, allowing the bats to control the amount of heat exchanged through the thin flight membrane (many other mammals use the capillary network in oversized ears for the same purpose).[51]

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.[52]

Echolocation

Main article: Animal echolocation § Bats
Spectrogram of Pipistrellus pipistrellus (common pipistrelle) bat vocalizations, forming a social call[53]
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. At 130 decibels in intensity, bat calls are some of the most intense, airborne animal sounds.[54]

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.[55] The delay of the returning echoes provides the bat with the ability to estimate the range to their prey.

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 of this range to avoid self-deafening. They then receive echoes back at the finely tuned frequency range by taking advantage of the Doppler shift of their motion in flight. The Doppler shift of the returning 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.[56]

A diagram illustrating bat echolocation, orange is the call and green is the echo

The new Yinpterochiroptera and Yangochiroptera classification of bats, supported by molecular evidence, suggests two possibilities for the evolution of echolocation. It may have been gained once in a common ancestor of all bats and was then subsequently lost in the Old World fruit bats, only to be regained in the horseshoe bats, or echolocation evolved independently in both the Yinpterochiroptera and Yangochiroptera lineages.[57]

Two groups of moths exploit a bat sense to echolocate: tiger moths produce ultrasonic signals to warn the bats that they (the moths) are chemically protected or aposematic, other moth species 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.

Plecotus auritus, the brown long-eared bat

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, earwigs, etc. 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.[58][59][60]

By repeated scanning, bats can mentally construct an accurate image of the environment in which they are moving and of their prey item.[61]

Vision

Although the eyes of most microbat species are small and poorly developed, leading to poor visual acuity, no species is blind.[62] 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 some microbats have distinct colourations, they may be able to discriminate colours.[37][63][64][65]

Megabat species often have excellent eyesight as good as, if not better than, human vision. Their eyesight, unlike that of its microbat relatives, is adapted to both night and daylight vision which enables megabats to have some colour vision.[65]

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, 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.[66]

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.[67]

Size

The smallest bat is the 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.[68][69] It is also arguably the smallest extant species of mammal, next to the Etruscan shrew.[70] 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).[71] 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.[72] The adaptations in a particular bat species can directly influence what kinds of prey are available to it.[73]

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 has 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.[74][75] 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. 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.[75][76]

Behaviour

Most microbats are nocturnal[77] and are active at twilight. A large portion of bats migrate hundreds of kilometres to winter hibernation dens,[78] while some pass into torpor in cold weather, rousing and feeding when warm weather allows for insects to be active.[79] Others retreat to caves for winter and hibernate for six months.[79] Bats rarely fly in rain, as the rain interferes with their echolocation, and they are unable to locate their food.

The social structure of bats varies, with some leading solitary lives and others living in caves colonized by more than a million bats.[80] 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, with individual bats switching roosts with others and often ending up in different trees and with different roostmates.

Studies also show that bats make various sounds in order to communicate with one another. Scientists in the field have listened to bats and have been able to associate certain sounds with certain behaviours that bats make after the sounds are made.[80]

Insectivores make up 70% of bat species and locate their prey by means of echolocation. Of the remainder, most feed on fruits.[81] Only three species sustain themselves with blood.

Some species even prey on vertebrates. The leaf-nosed bats (Phyllostomidae) of Central America and South America, and the two bulldog bat (Noctilionidae) species feed on fish. 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.[81] One species, the greater noctule bat, catches and eats small birds in the air.

Predators

Predators of bats include bat hawks, bat falcons, snakes, and even spiders.[82][83]

Reproduction

Newborn common pipistrelle, Pipistrellus pipistrellus
Colony of mouse-eared bats, Myotis myotis

Most bats have a breeding season, which is in the spring for species living in a temperate climate.[84] Bats may have one to three litters in a season, depending on the species and on environmental conditions, such as the availability of food and roost sites. Females generally have one offspring at a time, which could be a result of the mother's need to fly to feed while pregnant. Female bats nurse their young until they are nearly adult size, because a young bat cannot forage on its own until its wings are fully developed.

Female bats use a variety of strategies to control the timing of pregnancy and the birth of young, to make delivery coincide with maximum food ability and other ecological factors. Females of some species have delayed fertilization, in which sperm is stored in the reproductive tract for several months after mating. In many such cases, mating occurs in the fall, and fertilization does not occur until the following spring. Other species exhibit delayed implantation, in which the egg is fertilized after mating, but remains free in the reproductive tract until external conditions become favorable for giving birth and caring for the offspring.

In yet another strategy, fertilization and implantation both occur, but development of the fetus is delayed until favorable conditions prevail, during the delayed development the mother still gives the fertilized egg nutrients, and oxygenated blood to keep it alive. However, this process can go for a long period of time, because of the advanced gas exchange system.[85] All of these adaptations result in the pup being born during a time of high local production of fruit or insects.

At birth, the wings are too small to be used for flight. Young microbats become independent at the age of six to eight weeks, while megabats do not until they are four months old. Newborn bats feed solely on their mother's milk.[86][87]

Feeding

Bats typically hunt at night, reducing competition with birds, minimizing contact with certain predators, and travel large distances, up to 800 kilometres (500 mi), in search of food.[37] Their daylight hours are spent grooming and sleeping; they hunt during the night. The majority of food consumed by bats includes insects, fruits and flower nectar, vertebrates and blood.[88] Before going into hibernation, some species build up large reserves of body fat, both as fuel and as insulation.[87] Bats are not restricted to any one hunting strategy, but rather use a mix.[72]

A Philippines bat circles the sweet manzanitas fruits of the Aratiles tree (Muntingia).

The Chiroptera as a whole are in the process of losing the ability to synthesize vitamin C: most have lost it completely.[89] 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.[90] 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.[91]

Insects

Almost three-fourths of the world's bats are insect eaters. Bats consume both aerial and ground-dwelling insects. Each bat is typically able to consume one-third of its body weight in insects each night, and several hundred insects in a few hours, meaning that a group of a thousand bats could eat 3.6 metric tons (4 short tons) of insects each year.[92] Insectivorous bats may occasionally catch an insect in mid-air with its mouth, and eat it in the air. However, more often than not, bats use their tail membranes or wings to scoop up the insect.[86] Then, the bat takes the insect back to its roost and eats it there.[88][93] The bite force of these small bats is generated through mechanical advantage, in that it is side-independent, through the hardened armor of insects or the skin of fruit.[94] Insectivorous bats living at high latitudes have to consume prey with higher energetic value than tropical bats.[95]

Forage gleaners typically fly down and grasp their prey off the ground with their teeth, and take it to a nearby perch to eat it. Generally, these bats do not use echolocation to locate their prey. Instead, they rely on the sounds produced by the insects. Some make unique sounds, and almost all make some noise while moving through the environment.[86]

Fruits and nectar

An Egyptian fruit bat (Rousettus aegyptiacus) carrying a fig
Fruit bats roosting by day

Fruit eating, or frugivory, is found in particular species from both major suborders. These bats favor fleshy and sweet fruits, but not those particularly strong smelling or colorful.[86] They pull the fruit off the trees with their teeth, then fly back to their roosts to consume them, 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 over 150 types of plants depend on bats for seed dispersal.[92]

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

Some Chiropterans consume nectar instead, for which they 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.[86] 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 its rib cage.[96] 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.[97][98]

Nectar feeding also aids a variety of plants, since these bats serve as pollinators: pollen gets stuck to the bats' fur while they sip the nectar, and is transferred to the next flower they visit (or dusts off in flight).[92] Rainforests are said to benefit the most from bat pollination, because of the large variety of plants that depend on it.[99]

Vertebrates

Some bats are primarily carnivorous, feeding on vertebrates.[86] These bats typically eat a variety of animals, especially frogs, lizards, birds, and sometimes other bats.[92]

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.[86] Another example is the greater noctule bat, which is believed to catch birds in flight.

Also, several bat species, found on all continents, feed on fish. They use echolocation to detect tiny 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.[86]

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 mammals, while the hairy-legged and white-winged vampires feed on birds instead.[100] These species are found throughout Central and South America, as well as in Mexico and on the island of Trinidad.

Drinking

In 1960, Frederic A. Webster discovered some bats' method of drinking water using a high-speed (1000 FPS) camera and flashgun. He captured one skimming just above the surface of the water, lowering its jaw to collect a small quantity of water on each pass, taking repeated passes until it drank its fill.[87]

Other bats, such as the flying fox or fruit bat, gently skim the water's surface, then land nearby to lick the water from their chest fur.[101]

Communication

The fruit bat is one of only a few animals known to direct its calls at specific individuals in a colony rather than broadcast like birdsongs and alarm calls. 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.[102]

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, otherwise known as guano, is so rich in nutrients that it is mined from caves, bagged, and used by farmers to fertilize their crops. During the U.S. Civil War, guano was used to make gunpowder.[92]

Disease transmission

Further information: Bat-borne virus

Bats are natural reservoirs for a large number of zoonotic pathogens,[103] including rabies,[104] histoplasmosis both directly and in guano,[105] Nipah Hendra viruses[106] and possibly ebola virus.[107][108]

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.[109] They seem to be highly resistant to many of the pathogens they carry, suggesting a degree of adaptation to bats' immune systems.[109][110][111] 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.[112]

Among ectoparasites, bats carry fleas and mites, as well as specific parasites called bat bugs.[113] 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, such as the bat bugs which occupy the same niche.[114]

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.[115] 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.[105]

Rabies

As of 2016, bats present a significant hazard in areas where the rabies virus is endemic, such as the southern United States, where they serve as natural reservoirs.[112] In the United States, bats typically constitute around a quarter of reported cases of rabies in wild animals. However, their bites account for the vast majority of cases of rabies in humans.[116] Of the 36 cases of domestically acquired rabies recorded in the country in 1995–2010, two were caused by dog bites and four patients were infected by receiving transplants from an organ donor who had previously died of rabies. All other cases were caused by bat bites.[117]

Rabies is fully preventable if the patient is vaccinated before the onset of symptoms. However, bat bites may go ignored or unnoticed and hence untreated. Many victims may not realize they have been bitten, because bats have very small teeth and do not always leave obvious marks. Victims may also be bitten while sleeping or intoxicated, and children, pets, and the mentally handicapped are especially vulnerable.[117] Rabid bats are broadly distributed throughout the United States; in 2008–2010, cases were reported in every state except Alaska and Hawaii, and Puerto Rico.

The most severe threat to humans and domestic animals comes from sick, downed, or dead bats, which typically have a very high infection rate (e.g. 70% for the Austin bats).[112] Furthermore, since they may be clumsy, disoriented, and unable to fly, these stricken bats are much more likely to come into contact with humans.

Public health organizations such as the CDC generally recommend that any contact with a potentially infected animal (including any bat) be reported promptly, and those at risk of infection are treated with a post-exposure prophylaxis (PEP) regimen to prevent contraction of the virus, which is near-universally fatal with very few exceptions. 30,000 PEP treatments are performed each year in the US, in large part due to contact with bats.[105][112]

The Centers for Disease Control and Prevention provide fully detailed information on all aspects of bat management in North America, including how to capture a bat, what to do in case of exposure, and how to bat-proof a house humanely.[117] 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.

Bat rabies virus can rarely infect victims purely through airborne transmission ("cryptic rabies"). This has occurred among victims breathing virus-infected air in caves, after long exposure.[118][119]

Evidence suggests that all active widespread rabies strains 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.[112]

Conservation efforts

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.[120]

Artificial roosts

Many people put up bat houses to attract bats.[121] The 1991 University of Florida bat house is the largest occupied artificial roost in the world, with around 300,000 residents.[122][123] In Britain, thickwalled and partly underground World War II pillboxes have been converted to make roosts for bats,[124][125] and purpose-built bat houses are occasionally built to mitigate damage to habitat from road or other developments.[126][127]

Threats

A little brown bat with white nose syndrome

While conservation efforts are in place to protect bats, many threats still remain.

White nose syndrome
Main article: White nose syndrome

White nose syndrome is a condition associated with the deaths of millions of bats in the Eastern United States and Canada.[128] 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.[129] 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.[130] 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.[131] 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.[132] Scientific evidence suggests that longer winters where the fungus has a longer period of time to infect bats results in greater chances of mortality.[133][134][135] In 2014, infection crossed the Mississippi River,[136] but species native to northern Mexico and the West had not yet been affected.[137]

Bats being prepared for cooking
Barotrauma and wind turbines

Evidence suggests that barotrauma is causing bat fatalities around wind turbines.[138] 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.[139][140][141][142][143] In addition, it has been suggested that bats are attracted to these structures, perhaps seeking roosts, and thereby increasing the death rate.[139] Acoustic deterrents may help to reduce bat mortality at wind farms.[144]

Use as food
Main article: Bat as food

Bats are eaten in countries across Asia and the Pacific Rim. In some cases, such as in Guam, flying foxes have become endangered through hunting for food.[145]

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.[146] Because bats are mammals, yet can fly, they are liminal beings in many traditions.[147] 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.[148] In Aztec mythology, bats symbolized the land of the dead, destruction, and decay.[149][150][151][152][153] In Oaxacan mythology, the bat became nocturnal through jealousy of birds' feathers.[153] 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.[154]

Zapotec bat god, Oaxaca, 350–500 AD

The Weird Sisters in Shakespeare's Macbeth used the fur of a bat in their brew.[155] 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.[156]

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.[157] The bat is sacred in Tonga and is often considered the physical manifestation of a separable soul.[158]

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.[159][160][161] 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.[162]

See also

Notes

References

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Bibliography

  • Altringham, J.D. 1998. Bats: Biology and Behaviour. Oxford: Oxford University Press.
  • Dobat, K.; Holle, T.P. 1985. Blüten und Fledermäuse: Bestäubung durch Fledermäuse und Flughunde (Chiropterophilie). Frankfurt am Main: W. Kramer & Co. Druckerei.
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  • Findley, J.S. 1995. Bats: a Community Perspective. Cambridge: Press Syndicate of the University of Cambridge.
  • Fleming, T.H. 1988. The Short-Tailed Fruit Bat: a Study in Plant-Animal Interactions. Chicago: The University of Chicago Press.
  • Greenhall, Arthur H. 1961. Bats in Agriculture. A Ministry of Agriculture Publication. Trinidad and Tobago.
  • Klimpel, S.; Mehlhorn, H. 2014. Bats (Chiroptera) as Vectors of Diseases and Parasites: Facts and Myths. Heidelberg New York Dordrecht London: Springer.
  • Kunz, T.H. 1982. Ecology of Bats. New York: Plenum Press.
  • Kunz, T.H.; Racey, P.A. 1999. Bat Biology and Conservation. Washington: Smithsonian Institution Press.
  • Kunz, T.H.; Fenton, M.B. 2003. Bat Ecology. Chicago: The University of Chicago Press.
  • Neuweiler, G. 1993. Biologie der Fledermäuse. Stuttgart: Georg Thieme Verlag.
  • Nowak, Ronald M. 1994. Walker's Bats of the World. Baltimore: The Johns Hopkins University Press.
  • John D. Pettigrew's summary on Flying Primate Hypothesis
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  • Teeling, E.C. 2009. Chiroptera. Oxford University Press.
  • Twilton, B. 1999. My Life as The Bat. Liverpool Hope University Press.

External links

The Wikibook Dichotomous Key has a page on the topic of: Chiroptera

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