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The vortex may be stabilised by the animal changing its wing curvatures. When not flying, bats hang upside down from their feet, a posture known as roosting.
The ankle joint can flex to allow the trailing edge of the wings to bend downwards. This does not permit many movements other than hanging or clambering up trees.
This difference is reflected in the structure of the cervical or neck vertebrae in the two groups, which are clearly distinct.
Muscular power is needed to let go, but not to grasp a perch or when holding on. Bats have an efficient circulatory system.
They seem to make use of particularly strong venomotion, a rhythmic contraction of venous wall muscles. In most mammals, the walls of the veins provide mainly passive resistance, maintaining their shape as deoxygenated blood flows through them, but in bats they appear to actively support blood flow back to the heart with this pumping action.
Bats possess a highly adapted respiratory system to cope with the demands of powered flight, an energetically taxing activity that requires a large continuous throughput of oxygen.
In bats, the relative alveolar surface area and pulmonary capillary blood volume are larger than in most other small quadrupedal mammals.
It takes a lot of energy and an efficient circulatory system to work the flight muscles of bats. Energy supply to the muscles engaged in flight require about double the amount compared to the muscles that do not use flight as a means of mammalian locomotion.
In parallel to energy consumption, blood oxygen levels of flying animals are twice as much as those of their terrestrially locomoting mammals.
As the blood supply controls the amount of oxygen supplied throughout the body, the circulatory system must respond accordingly.
Therefore, compared to a terrestrial mammal of the same relative size, the bat's heart can be up to three times larger, and pump more blood.
With its extremely thin membranous tissue, a bat's wing can significantly contribute to the organism's total gas exchange efficiency.
When the bat has its wings spread it allows for an increase in surface area to volume ratio. The digestive system of bats has varying adaptations depending on the species of bat and its diet.
As in other flying animals, food is processed quickly and effectively to keep up with the energy demand. Insectivorous bats may have certain digestive enzymes to better process insects, such as chitinase to break down chitin , which is a large component of insects.
Nectivorous and frugivorous bats have more maltase and sucrase enzymes than insectivorous, to cope with the higher sugar contents of their diet. The adaptations of the kidneys of bats vary with their diets.
Carnivorous and vampire bats consume large amounts of protein and can output concentrated urine ; their kidneys have a thin cortex and long renal papillae.
Frugivorous bats lack that ability and have kidneys adapted for electrolyte -retention due to their low-electrolyte diet; their kidneys accordingly have a thick cortex and very short conical papillae.
Bats have higher metabolic rates associated with flying, which lead to an increased respiratory water loss. Their large wings are composed of the highly vascularized membranes, increasing the surface area, and leading to cutaneous evaporative water loss.
They are also susceptible to blood urea poisoning if they do not receive enough fluid. The structure of the uterine system in female bats can vary by species, with some having two uterine horns while others have a single mainline chamber.
Microbats and a few megabats emit ultrasonic sounds to produce echoes. 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, localise, and classify their prey in darkness. Bat calls are some of the loudest airborne animal sounds, and can range in intensity from 60 to decibels.
The latter is most pronounced in the horseshoe bats Rhinolophus spp. In low-duty cycle echolocation, bats can separate their calls and returning echoes by time.
They have to time their short calls to finish before echoes return. Bats contract their middle ear muscles when emitting a call, so they can avoid deafening themselves.
The time interval between the call and echo allows them to relax these muscles, so they can hear the returning echo. In high-duty cycle echolocation, bats emit a continuous call and separate pulse and echo in frequency.
The ears of these bats are sharply tuned to a specific frequency range. They emit calls outside this range to avoid deafening themselves.
They then receive echoes back at the finely tuned frequency range by taking advantage of the Doppler shift of their motion in flight.
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.
In addition to echolocating prey, bat ears are sensitive to the fluttering of moth wings, the sounds produced by tymbalate insects, and the movement of ground-dwelling prey, such as centipedes and earwigs.
The complex geometry of ridges on the inner surface of bat ears helps to sharply focus echolocation signals, and to passively listen for any other sound produced by the prey.
These ridges can be regarded as the acoustic equivalent of a Fresnel lens , and exist in a large variety of unrelated animals, such as the aye-aye , lesser galago , bat-eared fox , mouse lemur , and others.
By repeated scanning, bats can mentally construct an accurate image of the environment in which they are moving and of their prey.
Many moth species have a hearing organ called a tympanum , which responds to an incoming bat signal by causing the moth's flight muscles to twitch erratically, sending the moth into random evasive manoeuvres.
The eyes of most microbat species are small and poorly developed, leading to poor visual acuity , but no species is blind. Microbats may use their vision for orientation and while travelling between their roosting grounds and feeding grounds, as echolocation is only effective over short distances.
Some species can detect ultraviolet UV. As the bodies of some microbats have distinct coloration, they may be able to discriminate colours.
Megabat species often have eyesight as good as, if not better than, human vision. Their eyesight is adapted to both night and daylight vision, including some colour vision.
Microbats make use of magnetoreception , in that they have a high sensitivity to the Earth's magnetic field , as birds do.
Microbats use a polarity-based compass, meaning that they differentiate north from south, unlike birds, which use the strength of the magnetic field to differentiate latitudes , which may be used in long-distance travel.
The mechanism is unknown but may involve magnetite particles. Most bats are homeothermic having a stable body temperature , the exception being the vesper bats Vespertilionidae , the horseshoe bats Rhinolophidae , the free-tailed bats Molossidae , and the bent-winged bats Miniopteridae , which extensively use heterothermy where body temperature can vary.
The wings are filled with blood vessels, and lose body heat when extended. At rest, they may wrap their wings around themselves to trap a layer of warm air.
Smaller bats generally have a higher metabolic rate than larger bats, and so need to consume more food in order to maintain homeothermy.
Bats may avoid flying during the day to prevent overheating in the sun, since their dark wing-membranes absorb solar radiation.
Bats may not be able to dissipate heat if the ambient temperature is too high;  they use saliva to cool themselves in extreme conditions.
Bats also possess a system of sphincter valves on the arterial side of the vascular network that runs along the edge of their wings.
When fully open, these allow oxygenated blood to flow through the capillary network across the wing membrane; when contracted, they shunt flow directly to the veins, bypassing the wing capillaries.
This allows bats to control how much heat is exchanged through the flight membrane, allowing them to release heat during flight. Many other mammals use the capillary network in oversized ears for the same purpose.
Torpor , a state of decreased activity where the body temperature and metabolism decreases, is especially useful for microbats, as they use a large amount of energy while active, depend upon an unreliable food source, and have a limited ability to store fat.
Torpid states last longer in the summer for megabats than in the winter. During hibernation , bats enter a torpid state and decrease their body temperature for Heterothermic bats during long migrations may fly at night and go into a torpid state roosting in the daytime.
Unlike migratory birds, which fly during the day and feed during the night, nocturnal bats have a conflict between travelling and eating.
The energy saved reduces their need to feed, and also decreases the duration of migration, which may prevent them from spending too much time in unfamiliar places, and decrease predation.
In some species, pregnant individuals may not use torpor. The smallest bat is Kitti's hog-nosed bat Craseonycteris thonglongyai , which is 29—34 millimetres 1.
Small prey may be absent in the diets of large bats as they are unable to detect them. Flight has enabled bats to become one of the most widely distributed groups of mammals.
Bat roosts can be found in hollows, crevices, foliage, and even human-made structures, and include "tents" the bats construct with leaves. In temperate areas, some microbats migrate hundreds of kilometres to winter hibernation dens;  others pass into torpor in cold weather, rousing and feeding when warm weather allows insects to be active.
Different bat species have different diets, including insects, nectar, pollen, fruit and even vertebrates. Insectivorous bats may eat over percent of their body weight, while frugivorous bats may eat over twice their weight.
The Chiroptera as a whole are in the process of losing the ability to synthesise vitamin C. Most microbats, especially in temperate areas, prey on insects.
Fruit eating, or frugivory, is found in both major suborders. Bats prefer ripe fruit, pulling it off the trees with their teeth.
They fly back to their roosts to eat the fruit, sucking out the juice and spitting the seeds and pulp out onto the ground. This helps disperse the seeds of these fruit trees, which may take root and grow where the bats have left them, and many species of plants depend on bats for seed dispersal.
Nectar-eating bats have acquired specialised adaptations. These bats possess long muzzles and long, extensible tongues covered in fine bristles that aid them in feeding on particular flowers and plants.
This is beneficial to them in terms of pollination and feeding. Their long, narrow tongues can reach deep into the long cup shape of some flowers.
When the tongue retracts, it coils up inside the rib cage. Around species of flowering plant rely on bat pollination and thus tend to open their flowers at night.
Some bats prey on other vertebrates, such as fish, frogs, lizards, birds and mammals. These bats locate large groups of frogs by tracking their mating calls, then plucking them from the surface of the water with their sharp canine teeth.
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.
A few species, specifically the common, white-winged , and hairy-legged vampire bats, only feed on animal blood hematophagy.
The common vampire bat typically feeds on large mammals such as cattle ; the hairy-legged and white-winged vampires feed on birds.
Bats are subject to predation from birds of prey , such as owls , hawks , and falcons , and at roosts from terrestrial predators able to climb, such as cats.
Speakman argue that bats evolved nocturnality during the early and middle Eocene period to avoid predators. Among ectoparasites , bats carry fleas and mites , as well as specific parasites such as bat bugs and bat flies Nycteribiidae and Streblidae.
White nose syndrome is a condition associated with the deaths of millions of bats in the Eastern United States and Canada.
The fungus is mostly spread from bat to bat, and causes the disease. Bats are natural reservoirs for a large number of zoonotic pathogens ,  including rabies , endemic in many bat populations,    histoplasmosis both directly and in guano,  Nipah and Hendra viruses ,   and possibly the ebola virus.
Compared to rodents, bats carry more zoonotic viruses per species, and each virus is shared with more species. Some bats lead solitary lives, while others live in colonies of more than a million.
This may serve to introduce young to hibernation sites, signal reproduction in adults and allow adults to breed with those from other groups.
Several species have a fission-fusion social structure , where large numbers of bats congregate in one roosting area, along with breaking up and mixing of subgroups.
Within these societies, bats are able to maintain long term relationships. Bats are among the most vocal of mammals and produce calls to attract mates, find roost partners and defend resources.
These calls are typically low-frequency and can travel long distances. Males sing to attract females. Songs have three phrases: Bat songs are highly stereotypical but with variation in syllable number, phrase order, and phrase repetitions between individuals.
Calls differ between roosting groups and may arise from vocal learning. The animals made slightly different sounds when communicating with different individual bats, especially those of the opposite sex.
Bats in flight make vocal signals for traffic control. Greater bulldog bats honk when on a collision course with each other.
Bats also communicate by other means. Male little yellow-shouldered bats Sturnira lilium have shoulder glands that produce a spicy odour during the breeding season.
Like many other species, they have hair specialised for retaining and dispersing secretions. Such hair forms a conspicuous collar around the necks of the some Old World megabat males.
Male greater sac-winged bats Saccopteryx bilineata have sacs in their wings in which they mix body secretions like saliva and urine to create a perfume that they sprinkle on roost sites, a behaviour known as "salting".
Salting may be accompanied by singing. Most bat species are polygynous , where males mate with multiple females.
Male pipistrelle, noctule and vampire bats may claim and defend resources that attract females, such as roost sites, and mate with those females.
Males unable to claim a site are forced to live on the periphery where they have less reproductive success. For temperate living bats, mating takes place in late summer and early autumn.
In hibernating species, males are known to mate with females in torpor. Females of some species have delayed fertilisation, in which sperm is stored in the reproductive tract for several months after mating.
Mating occurs in the autumn but fertilisation does not occur until the following spring. Other species exhibit delayed implantation , in which the egg is fertilised after mating, but remains free in the reproductive tract until external conditions become favourable for giving birth and caring for the offspring.
During the delayed development the mother keeps the fertilised egg alive with nutrients. This process can go on for a long period, because of the advanced gas exchange system.
For temperate living bats, births typically take place in May or June in the northern hemisphere; births in the southern hemisphere occur in November and December.
Tropical species give birth at the beginning of the rainy season. The young emerges rear-first, possibly to prevent the wings from getting tangled, and the female cradles it in her wing and tail membranes.
In many species, females give birth and raise their young in maternity colonies and may assist each other in birthing.
Most of the care for a young bat comes from the mother. In monogamous species, the father plays a role. Allo-suckling, where a female suckles another mother's young, occurs in several species.
This may serve to increase colony size in species where females return to their natal colony to breed. For the little brown bat, this occurs about eighteen days after birth.
Weaning of young for most species takes place in under eighty days. The common vampire bat nurses its offspring beyond that and young vampire bats achieve independence later in life than other species.
This is probably due to the species' blood-based diet, which is difficult to obtain on a nightly basis. The maximum lifespan of bats is three-and-a-half times longer than other mammals of similar size.
Five species have been recorded to live over 30 years in the wild: One hypothesis consistent with the rate-of-living theory links this to the fact that they slow down their metabolic rate while hibernating ; bats that hibernate, on average, have a longer lifespan than bats that do not.
Bat species that give birth to multiple pups generally have a shorter lifespan than species that give birth to only a single pup. Cave-roosting species may have a longer lifespan than non-roosting species because of the decreased predation in caves.
A male Brandt's bat was recaptured in the wild after 41 years, making it the oldest known bat. Groups such as the 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 disturbing a bat or its roost can be punished with a heavy fine.
Many people put up bat houses to attract bats. The gates are designed not to limit the airflow, and thus to maintain the cave's micro-ecosystem.
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.
In many cultures, including in Europe, bats are associated with darkness, death, witchcraft, and malevolence. In Tanzania, a winged batlike creature known as Popobawa is believed to be a shapeshifting evil spirit that assaults and sodomises its victims.
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": The Weird Sisters in Shakespeare's Macbeth used the fur of a bat in their brew.
The bat is a primary animal associated with fictional characters of the night, both villainous vampires , such as Count Dracula and before him Varney the Vampire ,  and heroes , such as Batman.
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.
Texas and Oklahoma are represented by the Mexican free-tailed bat, while Virginia is represented by the Virginia big-eared bat Corynorhinus townsendii virginianus.
Insectivorous bats in particular are especially helpful to farmers, as they control populations of agricultural pests and reduce the need to use pesticides.
This also prevents the overuse of pesticides, which can pollute the surrounding environment, and may lead to resistance in future generations of insects.
Bat dung, a type of guano , is rich in nitrates and is mined from caves for use as fertiliser. The Congress Avenue Bridge in Austin, Texas , is the summer home to North America's largest urban bat colony, an estimated 1,, Mexican free-tailed bats.
About , tourists a year visit the bridge at twilight to watch the bats leave the roost. From Wikipedia, the free encyclopedia.
Order of flying mammals. For other uses, see Bat disambiguation and Bats disambiguation. List of bats and List of fruit bats. Recording of Pipistrellus pipistrellus bat time-expanded echolocation calls and social call.
List of bats by population. Retrieved 9 September Retrieved 24 June Journal of Mammalian Evolution. Retrieved 10 September University of California Museum of Paleontology.
Molecular Biology and Evolution. Several molecular studies have shown that Chiroptera belong to the Laurasiatheria represented by carnivores, pangolins, cetartiodactyls, eulipotyphlans, and perissodactyls and are only distantly related to dermopterans, scandentians, and primates Nikaido et al.
Journal of Molecular Evolution. Retrieved 17 December University of Chicago Press: Narrowing the Field of Plausible Hypotheses".
The Quarterly Review of Biology. Proceedings of the National Academy of Sciences. The Biology of Bats.
Journal of Experimental Biology. Trends in Ecology and Evolution. Interactive Management of Wild and Captive Animals. A Bat Man in the Tropics: University of California Press.
The Encyclopedia of Mammals 2nd ed. The molecules and mechanics of flight in animals". Royal Society Open Science. Morphologic and molecular evolution of bat wing digits".
Modelling the Flying Bird. Retrieved 31 October Proceedings of the Royal Society B: Archived from the original PDF on 22 July Bats Chiroptera as Vectors of Diseases and Parasites: Philosophical Transactions of the Royal Society of London.
American Journal of Physiology. Regulatory, Integrative and Comparative Physiology. Retrieved 10 June World Journal of Zoology. A Test of the Allotonic Frequency Hypothesis".
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