Skull and dentition The head and teeth shape of bats can vary by species. In general, megabats have a fox-like appearance with long snouts and ears, hence their nickname of "flying foxes". The number of teeth in bats can vary between 38 teeth in small, insect-eating species and as low as 20 in vampire bats. A diet of hard-shelled insects requires fewer but larger teeth along with longer canines and more robust lower jaws. In nectar-feeding bats, the canines are long, while the cheek teeth are reduced. In fruit-eating microbats, the cusps of the cheek teeth are adapted for crushing.
Wings, skin, and flight Bats are the only mammals capable of sustained flight, as opposed to the
gliding of
flying squirrels, colugos and
sugar gliders. The fastest bat, the
Mexican free-tailed bat (
Tadarida brasiliensis), can achieve a
ground speed of . The flexible finger bones of bats have a flattened cross-section and become less mineralised towards the tips. During flight, the bones undergo bending and
shearing stress; the former being less than in terrestrial mammals, and the latter being greater. The wing bones of bats are less resistant to breaking than those of birds. As in other mammals, and unlike in birds, the
radius is the main component of the forearm. Bats have five elongated digits, which all radiate around the wrist. The thumb points forward and supports the
leading edge of the wing, and the other digits support the tension held in the wing membrane. The second and third digits go along the wingtip, allowing the wing to be pulled forward against aerodynamic
drag without having to be thick, as in
pterosaur wings. The fourth and fifth digits go from the wrist to the
trailing edge and repel the bending force caused by air pushing up against the stiff membrane. The knees point upwards and outwards during flight due to the attachment of the femurs, while the ankle joint can bend the trailing edge downwards. and their thin, articulated wings allow them to manoeuvre more accurately than birds and fly with more lift and less drag. By folding the wings in toward their bodies on the upstroke, they save 35 percent energy during flight. Flight muscles used for the upstroke are located on the back, while those for the downstroke are at the chest. This is in contrast to birds, where both muscle types are at the chest. Nectar- and pollen-eating bats can hover in a similar way to
hummingbirds. The sharp leading edges of the wings can create
vortices, which provide
lift. The vortex may be stabilised by the animal changing its wing curvature. The
patagium is the wing membrane, which reaches from the arm and finger bones to the side of the body and the hindlimbs. While the skin on the body of the bat is covered in hair and sweat glands with an
epidermis, a
dermis, and a fatty subcutaneous layer, the patagium is an extremely thin double layer of epidermis separated by a
connective tissue centre rich with
collagen and
elastic fibres. The surface of the wings is equipped with touch-sensitive receptors on small bumps called
Merkel cells. Each bump has a tiny hair in the centre, allowing the bat to detect and adapt to changing airflow; the primary use is to judge the most efficient speed at which to fly, and possibly also to avoid
stalls. Insectivorous bats may also use tactile hairs when manoeuvring to capture flying insects.
Photoluminescence has been reported in at least six North American species based on 60 museum specimens. The wings, uropatagium (around the tail), and hind limbs of these bats glowed green when exposed to UV light. The explanation for this is uncertain, and some have suggested the phenomenon is an artefact of the techniques used to dry specimens for museum storage.
Roosting and gaits When not flying, bats hang upside down from their feet, a posture known as roosting. Most megabats roost with the head tucked towards the belly, whereas most microbats roost with the neck curled towards the back. This difference is due to the structure of the
cervical or neck vertebrae in the two groups, which are clearly distinct. Tendons allow bats to hang from a roost with no effort, which is needed to release. Bats are more awkward when crawling on the ground, though a few species, such as the
New Zealand lesser short-tailed bat (
Mystacina tuberculata) and the
common vampire bat (
Desmodus rotundus), are quite agile. These species move their limbs one after the other, but vampire bats accelerate by bounding, the folded-up wings being used to propel them forward. Vampire bats likely evolved these gaits to stalk their hosts, while short-tailed bats took to the ground due to a lack of competition from other mammals. Terrestrial locomotion does not appear to affect their ability to fly.
Internal systems Bats have an efficient
circulatory system. They seem to make use of particularly strong venomotion, a rhythmic contraction of venous wall muscles. In most mammals, the walls of the veins provide mainly passive resistance, maintaining their shape as deoxygenated blood flows through them, but in bats, they appear to actively support blood flow back to the heart with this pumping action. Because of their small, lightweight bodies, bats are not at risk of blood rushing to their heads when roosting. Compared to a terrestrial mammal of similar size, the bat's heart can be up to three times larger and pump more blood, while blood oxygen levels are twice as much. An active microbat can reach a heart rate of 1000
beats per minute. (
Hypsignathus monstrosus) Bats possess a highly adapted respiratory system to cope with the demands of powered flight. They have relatively large lungs, and many species have proportionally larger alveolar surface areas and pulmonary capillary blood volumes than other mammals. During flight, the respiratory cycle has a one-to-one relationship with the wing-beat cycle. Their mammalian lungs prevent them from flying at high altitudes. Bats can also meet oxygen demands by exchanging gas through the patagium of the wing. When the bat has its wings spread, it allows for an increase in surface area to volume ratio, 85% of the surface area being the wing. The subcutaneous vessels in the membrane lie near the surface and allow for the diffusion of oxygen and carbon dioxide. The
digestive system of bats varies depending on the species of bat and its diet. Digestion is relatively quick to meet the energy demands of flight. Insectivorous bats may have certain
digestive enzymes to better process insects, such as
chitinase to break down their
chitin exoskeleton. Vampire bats, probably due to their diet of blood, are unique among vertebrates in that they do not have the enzyme
maltase, which breaks down
malt sugar, in their intestinal tract. Nectivorous and frugivorous bats have more maltase and
sucrase enzymes than insectivores, 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. Water helps maintain the
ionic balance in their blood,
thermoregulation system and urinary and waste system. 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.
Senses Echolocation and hearing Microbats and a few megabats emit ultrasonic sounds to produce echoes. The sound intensity of these echoes is dependent on subglottic pressure. The bats' cricothyroid muscle, located inside the larynx, controls the orientation pulse frequency, which is an important function. By comparing the outgoing pulse with the returning echoes, bats can learn about their environment and detect prey in darkness. Some bat calls can reach over 140
decibels. Microbats use their
larynx to emit echolocation signals through the mouth or the nose. Bat call frequencies range from as low as 11 kHz to as high as 212 kHz. In addition to echolocating prey, bat ears are sensitive to sounds made by their prey, such as the fluttering of moth wings. The complex geometry of ridges on the inner surface of bat ears helps to sharply focus echolocation signals and to passively listen for any other sound produced by the prey. These ridges can be regarded as the acoustic equivalent of a
Fresnel lens. 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. By repeated scanning, bats can mentally construct an accurate image of the environment in which they are moving and of their prey. Many species of moth have exploited this, such as many
tiger moths, which produce
aposematic ultrasound signals to warn bats that they are chemically protected and therefore distasteful. Some tiger moths can produce signals to
jam bat echolocation. In some moth species, the
tympanum hearing organ causes the insect to move in random evasive manoeuvres when detecting a bat call.
Vision Microbats tend to have small eyes but are still sensitive to light, and no species is truly blind. Most microbats have
mesopic vision, meaning that they can detect light only in low levels, whereas other mammals have
photopic vision, which allows colour vision. Microbats may use their vision for orientation and while travelling between their roosting grounds and feeding grounds, as echolocation is effective only over short distances. Megabat species generally have good eyesight and may have some colour vision to help them distinguish ripe fruits. Some species can detect
ultraviolet (UV). As the bodies of some microbats have distinct colouration, they may be able to discriminate colours.
Smell Among bat species, megabats tend to have a more developed sense of smell, being particularly sensitive to
esters, which are found in ripe fruits. Insectivorous bats have less use for smell during foraging, as they rely on echolocation to search for prey.
Magnetoreception and infrared sensing Like birds, microbats' sensitivity to the
Earth's magnetic field gives them great
magnetoreception. Microbats use a polarity-based compass, which means they can distinguish 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 possibly involves
magnetite particles. Vampire bats are the only mammals that use
infrared sensing; heat sensors around the nose allow them to detect blood vessels near the surface of the skin of their target.
Thermoregulation Tropical bats tend to be
homoeothermic (having a stable body temperature), while temperate and subtropical species which enter
hibernation or
torpor are more
heterothermic (where body temperature can vary). Compared to other mammals, bats have a high
thermal conductivity. They lose heat via the wings when they are spread, so resting bats wrap their wings around themselves to keep warm. Smaller bats generally have a higher metabolic rate than larger bats and so need to consume more food in order to maintain homoeothermy. Bats may avoid flying during the day to prevent overheating in the sun, since they would absorb sun radiation via their dark wing membranes. Bats may not be able to release heat if the ambient temperature is too high; they use saliva to cool themselves in extreme conditions. Among megabats, the flying fox
Pteropus hypomelanus uses saliva and wing-fanning to cool itself while roosting during the hottest part of the day. Among microbats, the
Yuma myotis (
Myotis yumanensis), the
Mexican free-tailed bat (
Tadarida brasiliensis), and the
pallid bat (
Antrozous pallidus) cope with temperatures up to by panting, salivating, and licking their fur to promote evaporative cooling; this is sufficient to release twice their metabolic heat production. (
Perimyotis subflavus) in
torpor During torpor, bats drop their body temperature to , while their energy usage diminishes by 50 to 99%. Tropical bats may use it to reduce the chance of being caught by a predator during foraging. Megabats were generally believed to be homoeothermic, but three species of small megabats, with a mass of about , 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. 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 their body temperature returns to normal, they sometimes enter a shallow torpid state, known as "heterothermic arousal". Some bats become dormant during higher temperatures to keep cool in the summer months (
aestivation). 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 food requirements 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 use a more moderate state of torpor to maintain foetal development, while still saving energy.
Size The smallest bat, and one of the smallest mammals, is
Kitti's hog-nosed bat (
Craseonycteris thonglongyai), which is long with a forearm and weighs . The largest species is the
giant golden-crowned flying fox (
Acerodon jubatus), which can weigh with a wingspan of . Larger bats tend to use lower frequencies and smaller bats higher for echolocation; high-frequency echolocation is better at detecting smaller prey. Small prey may be absent in the diets of large bats as they are unable to detect them. ==Ecology==