Bat (order Chiroptera), any member of the only group of mammals capable of flight. This ability, coupled with the ability to navigate at night by using a system of acoustic orientation (echolocation), has made the bats a highly diverse and populous order. More than 1,200 species are currently recognized, and many are enormously abundant. Observers have concluded, for example, that some 100 million female Mexican free-tailed bats (Tadarida brasiliensis mexicana) form summer nursery colonies in Texas, where they produce about 100 million young in five large caves. The adult males are equal in number to the females, though they do not all range as far north as Texas. Furthermore, this species is found throughout tropical America. Thus, one species alone numbers, at the very least, in the hundreds of millions.
Written by: Don E. Wilson
General features
All bats have a generally similar appearance in flight, dominated by the expanse of the wings, but they vary considerably in size. The order is usually divided into two well-defined suborders: the Megachiroptera (the large Old World fruit bats) and theMicrochiroptera (small bats found worldwide). Among members of the Megachiroptera, flying foxes (Pteropus) have a wingspan of 1.5 metres (about 5 feet) and a weight of 1 kg (2.2 pounds). The largest insectivorous bat is probably the naked, or hairless, bat (Cheiromeles torquatus); it weighs about 250 grams (about 9 ounces). The largest of the carnivorous bats (and the largest bat in the New World) is the spectral bat (Vampyrum spectrum), also known as the tropical American false vampire bat, with a wingspan of over 60 cm (24 inches). The tiny hog-nosed, or bumblebee, bat (Craseonycteris thonglongyai) of Thailand is one of the smallestmammals. It has a wingspan of barely 15 cm (6 inches) and weighs about 2 grams (about 0.07 ounce).
Bats vary in colour and in fur texture. Facial appearance, dominated by the muzzle and ears, varies strikingly between families and often between genera. In several families a complex fleshy adornment called the nose leaf surrounds the nostrils. Although the exact function of these facial appurtenances has yet to be determined, scientists believe they may help to direct outgoing echolocation calls (see belowOrientation). Wing proportions are modified according to mode of flight. The tail and the membrane between the legs also differ, perhaps as adaptations to feeding, flight, and roosting habits. Finally, bats vary in the postures they assume when roosting, particularly in whether they hang suspended or cling to a wall and in the manner in which the wings are folded and used.
Distribution
Bats are particularly abundant in the tropics. In West Africa, for example, more than 30 genera embracing nearly 100 species have been cataloged; in the United States19 genera, totaling about 45 species, are known. Of the 18 bat families, 3—the vesper bats (family Vespertilionidae), free-tailed bats (family Molossidae), and horseshoe bats (family Rhinolophidae)—are well represented in the temperate zones. A few American leaf-nosed bats (family Phyllostomidae) range into mild temperate regions. Several vesper bats range well into Canada.
The Vespertilionidae are found worldwide except in the polar regions and on isolated islands. The brown bats of genus Myotis have a range almost equal to that of the entire order. The free-tailed bats and sheath-tailed bats (family Emballonuridae) also encircle the Earth but are restricted to the tropics and subtropics. The horseshoe bats extend throughout the Old World, the roundleaf bats (family Hipposideridae) and Old World fruit bats (family Pteropodidae) throughout the Old World tropics, and the leaf-nosed bats throughout the New World tropics and slightly beyond. The other families have more restricted ranges.
Importance to humans
Most bats are insectivorous, and they are important to humans primarily for their predation on insects, for pollination, and for seed dispersal. Little is known of the spectrum of insect species consumed, but the sheer quantity is formidable. The Mexican free-tailed bats of Texas have been estimated to consume about 9,100 metric tons (10,000 tons) of insects per year. Bats would thus seem to be important in the balance of insect populations and possibly in the control of insect pests.
Some bats feed on pollen and nectar and are the principal or exclusive pollinators of a number of tropical and subtropical plants. Others feed on fruit and aid in dispersing seeds, although bananas and figs must in some cases be protected from fruit-eating bats by early harvest or by nets.
Vampire bats (family Phyllostomidae, subfamily Desmodontinae) are considered serious pests of livestock in some parts of tropical America because the small wounds they cause provide egg-laying sites for parasites and because the vampires may transmit rabies and trypanosomiasis to cattle. Other bats also carry rabies or related viruses.
The guano (droppings) of insectivorous bats is still used for agricultural fertilizer in many countries and in the past was used as a source of nitrogen and phosphorus for munitions. Large guano deposits, in addition, cover and thus preserve many archaeologically interesting artifacts and fossils in caves.
In tropical regions large colonies of bats often inhabit houses and public buildings, where they attract attention because of their noisiness, guano, and collective odour. In western culture bats have been the subject of unfavourable myths; in parts of the Orient, however, these animals serve as symbols of good luck, long life, and happiness. In some parts of Southeast Asia and on some Pacific islands, flying foxes (Pteropus) are hunted for food. Small bats are also widely but irregularly eaten.
Certain physiological aspects of some bats, particularly those involving adaptations for long hibernation, daily lethargy, complex temperature regulation, acoustical orientation, and long-distance migrations, are of interest to biologists.
In species and numbers, bats constitute an important and generally nonintrusive form of wildlife. Several zoos have established interesting exhibits of bats; indeed, some flying foxes and fruit bats have been exhibited in European zoos since the mid-19th century, and they have been kept widely for research purposes. Bats are interesting pets but require specialized care.
Natural history
ACTIVITY PATTERNS
Nocturnal activity is a major feature of the behavioral pattern of bats: nearly all species roost during the day and forage at night. Carnivorous bats, vampire bats, and perhaps fishing bats (see bulldog bat) may have an advantage at night over inactive or sleeping prey. In addition, nocturnal flight protects bats from visual predators, exposure to the sun, high ambient temperature, and low relative humidity. The large area of naked wing skin might mean that bats would absorb rather than radiate heat if they were active during the day. They would also lose body water required for temperature regulation and would then be forced to forage near water or somehow retain more water (and thus more weight) in their bodies during flight.
The nocturnal activity pattern in bats is probably kept in synchrony with changing day lengths by their exposure to light at dusk or dawn. Bats often awaken and fly from the cave exit well before nightfall. Should they be too early, their internal clock may be reset. A few species of bats, including a flying fox (Pteropus samoensis), theyellow-winged bat (Lavia frons), and the greater sac-winged bat Saccopteryx bilineata, may forage actively during the day, but little is yet known of their special adaptations.
LOCOMOTION
Flight is the primary mode of locomotion in all bats, although the flight styles vary. Some groups (the free-tailed bats, for example) are adapted for flight in open spaces and high altitudes. They have long, narrow wings, swift flight, and a large turning radius. Slit-faced bats (Nycteridae), false vampire bats (Megadermatidae), and others are adapted for hovering as they pick prey off vegetation or feed on flowers. These bats have short, broad wings, slow flight, and a small turning radius. Some bats take flight easily from the ground: members of the genus Macrotus do so simply by flapping, while vampire bats (Desmodus) leap into the air and then spread their wings and fly. The free-tails, however, roost well above the ground because, upon takeoff, they fall before becoming airborne.
Though flight speeds in the wild are hard to measure, four vesper bat species, carefully observed, have been timed on average at 18.7 to 33.3 km (11.7 to 20.8 miles) per hour. In flight the posture of each of the four fingers incorporated into the wing is under precise and individual control. Finger and arm postures, which determine the shape, extension, and angle of the wings, govern such actions as turning, diving, landing, and hovering. Except when interrupted by insect catches or obstacles, bat flight paths are straight. Insects may be pursued and captured at a rate of up to two per second; during each catch the flight path is interrupted and thus appears erratic.
In many cases there is little locomotion other than flight. Bats that hang in caves may move across the ceiling by shifting their toehold, one foot at a time. A few genera, especially among the Old World fruit bats (family Pteropodidae), may crawl along branches in a slothlike posture, using their thumb claws as well as their feet. Thesheath-tailed bats (family Emballonuridae) and mouse-tailed bats (family Rhinopomatidae) hang on vertical surfaces suspended by their hind claws but with their thumbs and wrists propped against the surface. In this orientation they can scramble rapidly up or down and forward or backward, as well as sideways.
Bats of many families walk or crawl on either horizontal or vertical surfaces, using hind feet, wrists, and thumbs. Many move freely either backward or forward, a convenience for entering and leaving crevices. The vampire bats may also leap from roost to roost. The disk-winged bats (family Thyropteridae) and sucker-footed bat(one species, family Myzopodidae), as well as the bamboo bats (Tylonycteris), have specialized wrist and sole pads for moving along and roosting on the smooth surface of leaves or bamboo stalks. Bats are not known to swim in nature except, perhaps, by accident. When they do fall into water, however, they generally swim competently.
ROOSTING
Bats choose a variety of diurnal roosts, although the roost requirements of many bats, which are rather precise in terms of light, temperature, and humidity, limit their distribution. Each species favours a particular kind of roost, though this varies with sex, season, and reproductive activity. Many bats prefer isolated or secure roosts—caves, crevices in cliff faces, the interstices of boulder heaps, tree hollows, animalburrows, culverts, abandoned buildings, portions of buildings inaccessible to humans or infrequently accessed by them (i.e., a roof, attic, or hollow wall), or the hollow core of bamboo stalks. Some species roost externally—on tree trunks or in the branches of trees, under palm leaves, in unopened tubular leaves, or on the surface of rocks or buildings. For some the darkness, stability of temperature and humidity, and isolation from predators provided by caves and crevices seem essential. Others prefer the heat and dryness of sun-exposed roosts. Many bats also occupy nocturnal roosts, often rocky overhangs or cave entrances, for napping, for chewing food, or for shelter from bad weather. Many species likewise choose special nursery or hibernation roosts. Buildings are so widely exploited by bats (especially vesper bats, free-tailed bats, and sheath-tailed bats) that many species have probably become more abundant since the advent of architecture.
Bats are usually colonial; indeed, some form very large cave colonies. Generally, large colonies are formed by bats that roost in dense clusters, pressing against one another, although many are widely spaced and do not touch when roosting. Some of the Old World fruit bats strikingly defoliate the trees on which they roost. In trees flying foxes (Pteropus) may form outdoor camps numbering hundreds of thousands of individuals. Many species form smaller groups of several dozen to several hundred. Less commonly, bats are solitary; sometimes the adult female roosts only with its most recent offspring. Occasionally, one sex is colonial and the other is apparently solitary. The advantages of colonial or solitary life and the factors that govern colony size in bats with colonial predilection have not yet been established.
Elaborate communities of other animals are often satellites of cave-bat colonies. Among these are cave crickets, roaches, blood-sucking bugs, a variety of parasites (e.g., fleas, lice, ticks, mites, and certain flies), and dermestid beetles and other insects that feed on cave-floor debris—guano, bat and insect corpses, and discarded pieces of food or seeds. Molds and other fungi are also conspicuous members of the cave-floor community. Bats and their excretions alter the cave environment by producing heat, carbon dioxide, and ammonia.
MIGRATION
Many bats of temperate climates migrate annually to and from summer roosts and winter hibernation sites, with an individual often occupying the same roosts in seasonal sequence each year. Members of the same species may converge on a single hibernation cave or nursery roost from many directions, which indicates that the choice of migration direction to and from these caves cannot be genetically determined. When migration occurs, however, is probably genetically determined (i.e., instinctive) and influenced also by weather conditions and the availability of food. Nothing is known of how bats recognize migration goals or how succeeding generations learn their locations. Female young born at a nursery roost may memorize its location, but how they know where to go at other times is not clear. Likewise, little is yet known of energy storage, navigation, or other specializations for migrations.
Female Mexican free-tailed bats migrate from central Mexico to Texas and adjacent states each spring, returning south in the fall. Mating probably occurs in transient roosts during the spring flight. The migration is believed to remove pregnant and lactating females to a region of high food supply where they need not compete with males of their own species. Presumably they return to Mexico for its suitable winter climate and food supply and to meet their mates.
The North American red and hoary bats (Lasiurus borealis and L. cinereus) and thesilver-haired bat (Lasionycteris noctivagans) migrate in the fall from the northern United States and Canada to the southern United States and beyond, returning in the spring.
ORIENTATION
Bats of the suborder Microchiroptera orient acoustically by echolocation (“sonar”). They emit short high-frequency pulses of sound (usually well above the range of human hearing) and listen to the echoes returning from objects in the vicinity. By interpreting returning echoes, bats may identify the direction, distance, velocity, and some aspects of the size or nature (or both) of objects that draw their attention.Echolocation is used to locate and track flying and terrestrial prey, to avoid obstacles, and possibly to regulate altitude; orientation pulses may also serve as communication signals between bats of the same species. Rousette bats(megachiropteran genus Rousettus) have independently evolved a parallel echolocation system for obstacle avoidance alone. Echolocation pulses are produced by vibrating membranes in the larynx and emitted via the nose or the mouth, depending upon species. Nose leaves in some species may serve to channel the sound.
The echolocation signals spread in three dimensions on emission, the bulk of the energy in the hemisphere in front of the bat or in a cone-shaped region from the nostrils or mouth. When the sound impinges on an intervening surface (an insect or a leaf, for example), some of the energy in the signal is reflected or scattered, some absorbed, and some transmitted and reradiated on the far side of the surface; the proportion of sound energy in each category is a function of wavelength and of the dimensions, characteristics, and orientation of the object. The reflected sound spreads in three dimensions, and some portion of it may impinge on the bat’s ears at perceptible energy levels.
Bats’ external ears are generally large, which probably enhances their value for detecting the direction of incoming signals, and their middle and inner ears are specialized for high-frequency sensitivity. In addition, the bony otic (auditory) complex is often isolated acoustically from the skull, which probably improves signal comparison by both ears. The thresholds andranges of hearing in several genera of bats have been studied, and in each case the region of maximum sensitivity has been found to coincide with the prominent frequencies of the outgoing echolocation signals.
The characteristics of echolocation pulses vary with family and even with species. Echolocation pulses of a substantial number of bat species have been analyzed in terms of frequency, frequency pattern, duration, repetition rate, intensity, and direction. The prominent frequency or frequencies range from 12 kilohertz (1 kilohertz is equivalent to 1,000 hertz, or cycles per second) to about 150 kilohertz or more. Factors influencing frequency may include bat size, prey size, the energetics of sound production, inefficiency of the propagation of high frequencies, and ambient noise levels.
Orientation pulses may be of several types. The individual pulse may include a frequency drop from beginning to end (frequency modulation [FM]), or the frequency may be constant (CF) during part of the pulse, followed by a brief FM sweep; either FM or CF pulses may have high harmonic content. The pulse duration varies with the species and the situation. During cruising flight the pulses of the greater false vampire bat (Megaderma lyra) are 1.5 milliseconds (0.0015 second), those ofWagner’s mustached bat (Pteronotus personatus) 4 milliseconds, and those of thegreater horseshoe bat (Rhinolophus ferrumequinum) 55–65 milliseconds. In goal-oriented flight, such as the pursuit of an insect or the evaluation of an obstacle or a landing perch, the pulse duration is systematically altered (usually shortened) with target distance, sometimes ending with pulses as short as 0.25 millisecond.
During insect pursuit, obstacle avoidance, and landing maneuvers, there are three phases of pulse output design: search, approach, and terminal. The search phase, during which many bats emit about 10 pulses per second, precedes specific attention to a target. In the approach phase, which starts when the bat detects an object to which it subsequently devotes its attention, the bat raises the pulse rate to about 25 to 50 per second, shortens the pulses with decreasing distance, and often alters the frequency pattern. The terminal phase, which often lasts about 100 milliseconds, is characterized by extremely short pulses, repeated as rapidly as 200 or more times per second, and ceases as the bat intercepts the target or passes it (the stimulus being, perhaps, the cessation of echoes); another search phase follows. During the brief terminal phase (a fraction of a second), the bat is engaged in final interception (or avoidance) maneuvers and appears to pay little attention to other objects.
In addition to sensitive ears, the use of echolocation to gain sensory information requires integration of the vocal and auditory centres of the brain. Not only must the nervous system of the bat analyze in a few thousandths of a second the reflected, and thus altered, form of its own pulse, but it must separate this echo from those of other individuals and from others of its own pulses. All of this must be done while the animal (and often the target) is moving in space. In the laboratory, bats have been found to be able to identify, pursue, and capture as many as two fruit flies (Drosophila, about 3 mm [0.12 inch] long) per second and to locate and avoid wires as fine as 0.1 or even 0.08 mm (0.004 or 0.003 inch) in diameter.
Research has provided some information on the mechanisms of bat sonar. There is evidence that the multiple frequencies of FM or harmonic patterns serve in determining target direction. The relative intensities of the various frequencies are different at each ear, which allows the animal to determine the target’s direction when three or more frequencies are received. Target velocity may be measured by CF bats through the use of the Doppler shift, a change in perceived frequency due to the relative motion of the bat and its target. Changes in pulse-echo timing may provide information on target distance and velocity. The ratio of useful signal to background noise is increased by several mechanisms, including specializations of the middle ear and its ossicles (tiny bones), isolation of the cochlea (the area where sound energy is converted into nerve impulses), and adaptations of the central nervous system.
FOOD HABITS
Most bats feed on flying insects. In some cases prey species have been identified from stomach contents or from discarded pieces under night roosts, but such studies have not yet provided an adequate measure of the spectrum of bat diets. Bats identify and track insects in flight by echolocation. Large insects may be intercepted with the wing membranes and pulled into the mouth. Some bats feed on arthropods, such as large insects, spiders, and scorpions, that they find on the ground, on walls, or on vegetation. These bats may either land on and kill their prey before taking off with it or pick it up with their teeth while hovering.
Two genera (Noctilio and Myotis) include at least one species that catches small fishand possibly crustaceans. All fish-eating species also feed on flying insects or have close relatives that do so. Each is specialized in having exceptionally large hind feet armed with long, strong claws with which the fish are gaffed.
The Megachiroptera and many of the phyllostomidgenera feed on a variety of fruits, often green or brown in colour; usually such fruits are either borne directly on wood or hang well away from the bulk of the tree and have a sour or musky odour.
The Old World fruit bat subfamily Macroglossinae (and some other fruit bats) and certain leaf-nosed bats feed, at least in part, on nectar and pollen. Many tropical flowers, adapted forpollination by these bats, open at night, are white or inconspicuous, have a sour, rancid, or mammalian odour, and are borne on wood, on pendulous branches, or beyond or above the bulk of the plant. The phyllostomid Glossophaginae may also feed on flowers. (See Sidebar: Bat-Loving Flowers.)
Several phyllostomid and megadermatid genera are carnivorous, feeding on small rodents, shrews, bats, sleeping birds, tree frogs, and lizards. The true vampires, which feed on the blood of large mammals or birds, land near a quiet prospective victim, walk or jump to a vulnerable spot on it where the skin is relatively exposed—the edge of the ear or nostril, around the anus, or between the toes, for example—make a scooping, superficial bite from which the blood oozes freely, and lap the blood with very specialized tongue movements. Each vampire requires about 15 millilitres (about half an ounce) of blood per night.
The interaction of bats with their food, be it insects, fruit, or flowers, probably has a substantial impact on some biological communities. Many plants are dependent on bats for pollination; other plants benefit from seed dispersal by bats. Moths of two families are known to take evasive or protective action on hearing bat pulses nearby, an adaptation that implies heavy predation.
MAINTENANCE BEHAVIOUR
Bats are meticulous in their grooming, spending a fair part of the day and night combing and grooming their fur and cleansing their wing membranes. Generally, they comb with the claws of one foot while hanging by the other; they remove the combings and moisten their claws with their lips and tongue. On the wing membranes in particular, they use the mouth meticulously, perhaps oiling the skin with the secretions of dermal (skin) glands while cleansing it.
SOCIAL INTERACTIONS
Although social interactions per se have not been observed between adult bats, they are known to often segregate by sex. As noted above, pregnant females in many species occupy special nursery roosts until their young are independent. In some species the sexes occupy the same general roost but gather in separate clusters. In others the sexes intermingle or arrange themselves into a pattern within a group—the females centrally, for example, and the males peripherally. Sexual segregation during foraging has been reported for several species. Among bats that migrate over long distances, such as Mexican free-tailed, red, and hoary bats, the sexes may meet only briefly each year.
Life cycle
Details of the life cycle are known for only a few species. In northern temperate zone species, there is an annual cycle of sexual activity, with birth taking place between May and July. In males the testes, normally located in the abdominal region, descend seasonally into the scrotum, and active spermatogenesis occurs. In females sexual receptivity may be associated with egg maturation and release. Tropical bats may exhibit a single annual sexual cycle or may be diestrous (i.e., have two periods of fertility) or polyestrous (have many).
The sexual cycles of entire populations are closely synchronized, so almost all mating occurs within a few weeks. The periods of gestation, birth, lactation, and weaning are also usually synchronized. Gestation varies in duration: five or six months in flying foxes (Pteropus), more than five months in vampire bats (Desmodus), three months in some small leaf-nosed bats (Hipposideros), and 6 or 7 to 14 weeks in several small vesper bats (family Vespertilionidae). The length of gestation may be influenced by both ambient (surrounding) and body temperature.
In several North American and northern Eurasian vesper and horseshoe bats that hibernate, copulation occurs in the fall, and the sperm are stored in the female genital tract until spring. Ovulation, fertilization, and implantation occur after emergence from hibernation, when the female again has available an abundant food supply and a warm roost. Such favourable environmental conditions greatly enhance the young bat’s chances of survival.
Most bats bear one young, but the big brown bat (Eptesicus fuscus) may bear twins, and the Eastern red bat (Lasiurus borealis) bears litters of one to four.
At birth the young, which may weigh from one-sixth to one-third as much as the mother, usually have well-developed hind legs with which they hold on to their mother or to the roost. Their wings are very immature. The young are hairless or lightly furred and are often briefly blind and deaf. Female bats normally have one pectoral (at the chest) or axillary (at the armpit) mammary gland on each side. Several species that carry their young while foraging also have a pair of false pubic nipples, which the infant may hold in its mouth when its mother flies. The infants are nourished by milk for a period of about five or six weeks in many small bats and for five months in the Indian flying fox (Pteropus giganteus). By two months of age, most smaller bats have been flying and foraging for three or four weeks and have achieved adult size.
In many species females late in pregnancy migrate to special nursery roosts, in which large numbers of pregnant females may aggregate, usually to the exclusion of nonpregnant females, males, and bats of other species. In some cases the nursery roosts seem to be chosen for their high temperature, which may derive from the sun, from the bats themselves, or from decomposing guano. When foraging, some bats (Erophylla) leave their infants hanging quietly, one by one, on the cave wall or ceiling. In the case of the Mexican free-tailed bat and a few others, the closely spaced infants may move about and mingle on the wall. Some bats carry their young with them for a short period of time. Generally, each mother, on returning to her roost, seeks out her own offspring by position, smell, and acoustical exchange.
Some bats achieve sexual maturity in their first year, others in their second. Infant mortality appears to be high. Developmental and genetic errors and disease take their toll, but accidents seem to cause more serious losses—the young may fall from the ceiling or perhaps have serious collisions in early flight attempts. A fair number of bats probably fail to make the transition from dependent infants to self-sufficient foragers.
Adult bats, on the other hand, have low mortality. Predation is rarely serious, especially for cave-dwelling species. Disease, parasitic infestation, starvation, and accidents apparently take small tolls. There are records of several big brown (Eptesicus fuscus), little brown (Myotis lucifugus), and greater horseshoe bats(Rhinolophus ferrumequinum) that have lived more than 20 years, and a few have lived more than 30. Probably many bats in temperate climates live more than 10 years. Longevity has not been established for most tropical species, but a few are known to live for more than 10 years.
Several factors probably contribute to the unusual longevity of bats. Generally isolated roosts and nocturnal flight substantially protect them from predation, from some elements of weather, and from exposure to the sun. Their largely colonial way of life may ensure that entire populations experience contagious infection and subsequent immunity; indeed, such a pattern in the past may have hastened adaptation to disease. The persistent use of various seasonal roosts probably ensures isolation and security, food and water supplies, and access to mates. Many bats, moreover, reduce their body temperature at rest. Not only is there a probability that this conserves some cellular “machinery,” since metabolism is reduced, but fewer hours need to be spent in actively seeking food and water.
Bats are mammals with front limbs modified for flight. The chest and shoulders are large and well-muscled to provide power to the wings. The hips and legs are slender, as they do not usually support any body weight. Wing shape, governed by the relative lengths of the forearm and the fingers, varies greatly, in adaptation to flight characteristics. The fingers, other than the thumb, are greatly elongated and are joined by a membrane that extends from the posterior border of the forearm and upper arm to the side of the body and leg as far as the ankle or foot. The wing membrane consists of two layers of skin, generally darkly pigmented and hairless, between which course blood vessels and nerves. When not fully extended, the wing skin is gathered into wrinkled folds by elastic connective tissue and muscle fibres. Some of the fingers, especially the third, fold over when the bat is not in flight; the wing may then be quite tightly folded or may partly enfold the bat’s undersurface. The thumb, always free of the wing membrane, is used for walking or climbing in some species; in others it is used for handling food. Only the thumb—and occasionally the index finger—ends with a claw. Bats that walk often have pads or suction disks on their thumbs or wrists or both, and many female bats use their thumbs to suspend themselves, hammock fashion, when giving birth.
Most bats have a membrane, consisting of skin like that of the wings, that extends between their legs (the uropatagium, or interfemoral membrane). In the midline the interfemoral membrane is usually supported, at least in part, by the tail, with the distal edges often shaped in flight by greatly elongated heel bones, or calcars. The interfemoral membrane, especially well-developed in insectivorous, carnivorous, and fish-eating bats, is less-well-developed or even absent in the vampires and in fruit- and flower-feeding bats. Many bats, on catching large prey in flight, bring the membrane forward and, by flexing the neck and back, tuck the prey against and into the membrane. With this maneuver the bat takes hold of the victim headfirst and is able to kill or disable it promptly.
At rest a bat’s head, especially the ears, is its most striking feature. The neck is likely to be short and relatively immobile. The projecting portion of the external ear (the pinna) is usually extremely large and often is funnel-shaped. In several genera that feed on terrestrial arthropods, the ears are particularly oversized, probably for highly precise directional assessment. A projection on the front side of the auditory canal (the tragus) or another on the rear side (antitragus) may also be conspicuous. The ears are often highly mobile, sometimes flicking back and forth in phase with the production of sonar signals. In some species the ears are immobile, but in all cases they probably function in tandem for directional analysis.
Bats often have a rodentlike or foxlike muzzle, but in many the face has a pushed-in puglike appearance. In the nectar feeders the snout is elongated to house the long extensible tongue. Many bats have a facial ornament, the nose leaf, which consists of skin and connective tissue. It surrounds the nostrils and extends as a free flap or flaps above the nostrils and in front of the face. The complexity and shape of the nose leaf varies with family; its presence correlates with nasal emission of orientation signals. Thus, it is supposed that the nose leaf influences sound output, perhaps by narrowing the beam, but evidence is sparse.
Most bats are well furred except for the wing membranes. Colours are generally shades of brown, tan, gray, or black on top and lighter shades on the underside. Red, yellow, or orange variants occur in many species. Speckled or mottled patterns are common, as are bright or light-coloured spots or stripes. Bright red, yellow, or orange shading on the head, neck, and shoulders is not unusual. Mottled fur may enable the bat to be inconspicuous on lichen-covered bark or rock. Bright spots may simulate the speckled sunlight of the forest canopy as seen from below. Stripes probably break up contours. The colouring seen while the animal is hanging may be a kind of countershading for concealment, or it may enhance the bat’s simulation of a ripening fruit or a dead leaf. Many bats that roost externally hang from a branch by one foot, which then looks like a plant stem.
Many bats have large dermal glands, the location of which depends on family. These glands secrete odorous substances that may serve as species or sex recognition signals (pheromones). Some glands may also supply oils for conditioning the skin or waterproofing the fur.
When fully active, bats have a body temperature of about 37 °C (98.6 °F). Although some bats maintain fairly even body temperatures, a large number undergo periodic raising or lowering of their temperature. Many of the vesper bats and horseshoe bats and a few free-tailed bats reduce their body temperature to that of their surroundings (ambient temperature) shortly after coming to rest. This condition is called heterothermy. They raise their temperature again on being aroused or when readying themselves for nocturnal foraging. The drop in body temperature, if the ambient temperature is relatively low, results in a lethargic state. Energy is conserved by thus “turning down the thermostat,” but the bat is rendered relatively unresponsive to threats by predators or weather. Heterothermic bats therefore generally roost in secluded sites offering protection, often in crevices. In heterothermic bats one or more sensory systems and the brain remain sensitive at low temperatures and initiate the necessary heat production for arousal. Heat is generated by the metabolism of fat and by shivering.
Many bats that exhibit daily torpor also hibernate during the winter and therefore must store energy as body fat. In the fall these bats increase their weight by 50 to 100 percent. They must also migrate from the summer roost to a suitable hibernation site (often a cave) that will remain cool and humid throughout the winter without freezing. Large populations often aggregate in such caves. Hibernation involves the absence of temperature regulation for long periods in addition to adaptations of circulation, respiration, and renal function and the suspension of most aspects of activity. Bats of hibernating species generally court and mate in the fall when they are at their nutritional peak. During pregnancy, lactation, and juvenile growth, bats probably thermoregulate differently, more closely approximating stability.
Bats of several tropical families maintain a constant body temperature (homeothermy). This, however, depends on the nutritional state as well. A spectrum of degrees of homeothermy and heterothermy probably will be discovered.
Digestion and water conservation
Digestion in bats is unusually rapid. They chew and fragment their food exceptionally thoroughly and thus expose a large surface area of it to digestive action. They may begin to defecate 30 to 60 minutes after beginning to feed and thereby reduce the load that must be carried in flight.
Some bats live in sun-baked roosts without access to water during the day. They may choose these roosts for their heat, and thus conserve their own, but it is not yet known how they hold their body temperature down without using water. In the laboratory, bats die if body temperature rises above about 40–41 °C (104–106 °F).
Senses
In folklore, bats have been considered to be blind. In fact, the eyes in the Microchiroptera are small and have not been well studied. Among the Megachiroptera the eyes are large, but vision has been studied in detail only in flying foxes. These bats are able to make visual discriminations at lower light levels than humans can. The Megachiroptera fly at night, of course, and some genera fly below or in the jungle canopy, where light levels are very low. Except for rousette bats (Rousettus), none are known to orient acoustically.
Studies of several genera of Microchiroptera have revealed that vision is used in long-distance navigation and that obstacles and motion can be detected visually. Bats also presumably use vision to distinguish day from night and to synchronize their internal clocks with the local cycle of daylight and darkness.
The senses of taste, smell, and touch in bats do not seem to be strikingly different from those of related mammals. Smell is probably used as an aid in locating fruit and flowers and possibly, in the case of vampire bats, large vertebrates. It may also be used for locating an occupied roost, members of the same species, and the differentiation of individuals by sex. Many bats depend upon touch, aided by well-developed facial and toe whiskers and possibly by the projecting tail, to place themselves in comforting body contact with rock surfaces or with other bats in the roost.
The fossil record of bats prior to the Pleistocene Epoch (about 2,600,000 to 11,700 years ago) is limited and reveals little about bat evolution. Most fossils can be attributed to living families. Skulls and teeth compatible with early bats are known from about 60 million years ago, during the Paleocene Epoch. These specimens, however, may well have been from insectivores, from which bats are clearly distinguishable only on the basis of flight adaptations. By 45 million years ago (theEocene Epoch), bats with fully developed powers of flight had evolved.
The order Chiroptera is readily divided into two suborders—Megachiroptera (large Old World fruit bats) and Microchiroptera (small bats). The Megachiroptera orient visually and exhibit a number of primitive skeletal features. The Microchiroptera orient acoustically. It is not certain that they have a common origin. The suborders either evolved separately from flightless insectivores or diverged very early in chiropteran history.
The two principal geographic centres of bat evolution appear to be the Australo-Malaysian region, with about 290 species, and the New World tropics, with about 230 species. Comparable ecological niches in the Old World and the New World are occupied largely by different genera of bats, usually of different families.
Classification
Distinguishing taxonomic features
The order Chiroptera is defined by flight and the elongated finger bones and marked pectoral specialization that support it. Weak pelvic and leg development is also a chiropteran feature. The ulna of the forearm is reduced; claws are absent on the fingers except on the thumb (and occasionally the second finger); and the knee is directed rearward and outward. The maximum complement of permanent teeth is 38, the minimum 20.
Annotated classification
The following classification is based on the third edition of Mammal Species of the World, edited by Don E. Wilson and DeeAnn M. Reeder, published in 2005. Subsequent research has shown, however, that the number of bat species continues to increase from the discovery of new forms and from the results of studies using DNA analysis to examine the evolutionary relationships between known bat species.
Order Chiroptera
1,116 species in 18 families belonging to 2 suborders. Found from the tropics into temperate regions worldwide.
Suborder Microchiroptera
930 species in 17 families found from the tropics into temperate regions worldwide.
Family Vespertilionidae (vesper bats)
407 small to medium-sized species in 48 genera found worldwide to tree line and on many oceanic islands. Muzzle plain; eyes small; ears moderate to large; wings generally long and moderately narrow; tail and interfemoral membrane well developed. All walk well, often entering crevices. A few species eat fish. Generally colonial in dense, touching clusters; a few solitary; mostly roost in caves, attics, barns, hollow trees, boulder heaps, the twigs of birds’ nests, or roof thatching; a few (Lasiurus) roost in branches, on tree trunks, or in the hollow core of bamboo (Tylonycteris). Many temperate species hibernate and migrate; many are drably coloured; several that roost externally are either spotted (Euderma) or speckled (Lasiurus). Brown bats (Myotis and Eptesicus) and pipistrelles (Pipistrellus) found worldwide.
Family Phyllostomidae (American leaf-nosed bats, including vampire bats)
160 small to large species in 55 genera found from the southwestern United States through tropical America. Nose leaf simply shaped, ears often large and mobile, wings generally short and broad; tail and interfemoral membrane vary from absent to well developed; fur colour and patterns varied. Includes insectivorous, carnivorous, and fruit- and flower-feeding species. Generally do not walk. Colonial, often densely so; generally roost in touching clusters in caves, tree hollows, buildings, and culverts or in the open under bridges or eaves, in the crests of palm trees, or on the underside of palm leaves. Flight ranges from swift and straight to hovering.
Family Rhinolophidae (horseshoe bats)
77 small to moderately large species in 1 Old World genus. Complex nose leaf; large, highly mobile ears; wings short and rounded; well-developed interfemoral membrane, supported by tail. Unable to walk. Fur generally brown (occasionally red). Dark, humid roosts selected, especially caves, but tree hollows, buildings, and culverts as well. Generally colonial, nontouching. Usually feed on flying insects.
Family Hipposideridae (Old World leaf-nosed bats)
81 small to large species in 9 genera, some ranging from Old World tropics to temperate regions. Complex nose leaf with both anterior and posterior portions, the anterior portion being homologous to the “horseshoe” in the closely related Rhinolophidae. Toes have only two phalanges, owing to fusion; second finger of wing has only metacarpal bones; other fingers have two phalanges. Ears medium, without tragus. Fur colour ranging from dark brown to hues of red and yellow. Mostly colonial in caves, tunnels, or other dark retreats. Forage for flying insects.
Family Molossidae (free-tailed bats)
100 robust, small to very large species in 16 genera, some ranging into mild temperate regions. Tail projects well beyond the well-developed interfemoral membrane; ears large, rather immobile, often fused to one another, and unusually shaped; lips and snout often heavy; eyes tiny. Wings very long and narrow; legs well developed for walking; toes often bearing bristles. Often have a conspicuous odour; fur short, usually dark brown or black. Many highly colonial, with millions clustering in caves, tree hollows, and buildings; dense, touching groups roost pressed against fellows or walls, often in crevices; many prefer hot, dry roosts. Walk exceptionally well. Feed on flying insects; some migrate.
Family Emballonuridae (sheath-tailed, or sac-winged, bats)
51 small to medium-sized species in 13 genera found in the tropics excluding the West Indies and some Pacific islands; each genus restricted to either Old or New World. Ears large but simply shaped; eyes small; muzzle sharp but plain; tail short, perforating the dorsal surface of a well-developed interfemoral membrane. Several genera have a glandular pouch in the wing extensions. Roost on vertical surfaces, such as tree trunks, cliff faces, cave entrances, and walls; some favour buildings. Some densely colonial but not touching one another; others form small groups or are solitary. Hang suspended from toes with wrists propped against wall.
Family Nycteridae (slit-faced, or hollow-faced, bats)
16 small to medium-sized species in 1 genus (Nycteris) distributed through most of tropical Africa and Southeast Asia. Head with peculiar nasal depression, cleft nose leaf, and a deep midline facial cleft behind and above the nostrils. Ears large; wings broad; tail long with split end; heel bones greatly elongated; well-developed interfemoral membrane. Prey mostly on terrestrial insects resting on vegetation, rocks, or walls. Unable to walk. Roosts usually dark and humid; some species roost exposed in forest canopy.
Family Mormoopidae (leaf-chinned bats)
10 small species in 2 genera of tropical Central and South America. Some walk. All lack nose leaf but have elaborate lip leaves. Tail and interfemoral membrane well developed. Colour ranges from brown through orange, red, and yellow. Feed on flying insects. Densely colonial in dark caves, population of colonies often numbering in the tens of thousands.
Family Megadermatidae (false vampire bats)
5 moderately large species in 4 genera of Central Africa, Southeast Asia, and Australia. External ears very large and fused across midline; nose leaf large with truncated end; eyes relatively large; wings broad; interfemoral membrane well developed and supported distally by heel bones; no external tail. Feed principally on terrestrial arthropods, as in Nycteridae; at least 2 species also feed on small vertebrates taken in the same fashion as arthropod prey. Unable to walk. Form small colonies of nontouching individuals usually in dark, secluded caves or abandoned buildings. Yellow-winged bat (Lavia frons) is at least partly diurnal and roosts in trees in the savanna and open forest.
Family Natalidae (funnel-eared bats)
8 species of small, slenderly built bats in 3 genera (Natalus) of Central America, northern South America, and the West Indies. Thick gray, buff, yellow, or reddish fur. Well-developed tail and interfemoral membrane. Ears large; snout plain. Walk clumsily and do not enter crevices; cave-dwelling and colonial in nontouching groups. Feed on flying insects.
Family Rhinopomatidae (mouse-tailed bats)
4 small species in 1 genus (Rhinopoma) of North Africa and tropical Asia. Tail very long and largely free beyond a narrow interfemoral membrane. Ears large; small nose leaf; primitive shoulder girdle.
Family Thyropteridae (disk-winged bats)
3 species in 1 genus (Thyroptera) of Central America and northern South America, excluding West Indies. Base of thumb and sole provided with sucking disk; simple muzzle; ears large; second finger reduced to rudiment. Roost alone or in small groups, often in still-furled banana leaves. Biology poorly known.
Family Mystacinidae (New Zealand short-tailed bats)
2 small species in 1 genus (Mysticina) of New Zealand. Simple head similar to that of vesper bats. Wings fold very compactly; thumb and toe claws long and sharp; highly adapted for walking; tail perforates interfemoral membrane dorsally. Feed on flying and terrestrial insects. Biology poorly known.
Family Furipteridae (smoky bats)
2 small, delicately built species in 2 genera of northern South America and Central America. Snout plain; tail long, ending short of the well-developed interfemoral membrane; thumb vestigial; legs long; feet small. Biology unknown; probably insectivorous.
Family Noctilionidae (bulldog bats)
2 medium-sized species in 1 genus (Noctilio) of the American tropics. Muzzle heavy but unadorned; lips full; internal cheek pouches; ears large, pointed, and mobile; wings long and narrow. Tail well developed, extending to midpoint of large interfemoral membrane, which is pierced dorsally by tail tip; interfemoral membrane supported by very well-developed bones. Insectivorous, but one species also uses its very large feet to capture fish. Musky odour. Walk well; often roost in crevices, tree hollows, attics, grottoes, and caves; colonial, in touching clusters.
Family Craseonycteridae (hog-nosed, or bumblebee, bat)
1 tiny species of Thailand, Craseonycteris thonglongyai, perhaps the smallest living mammal.
Family Myzopodidae (Old World sucker-footed bat)
1 species in 1 genus (Myzopoda) endemic to Madagascar. Small, plain muzzle; large ears with peculiar mushroom-shaped lobe. Thumb and sole with adhesive disks; vestigial thumb claw; tail extends free beyond interfemoral membrane. Probably insectivorous; biology unknown.
Suborder Megachiroptera
Family Pteropodidae (flying foxes and other Old World fruit bats)
186 generally large species in 42 fruit- or flower-feeding genera found in the Old World tropics and subtropics, including many Pacific islands. Lack acoustic orientation except rousette bats (Rousettus); ears small; eyes large, with vision well developed; generally roost in trees; often colonial; often show countershading, cryptic markings, or bright fur colours or patterns. Index finger generally clawed; tail short or lacking; interfemoral membrane reduced. Muzzle simple in appearance, except in the hammer-headed fruit bat (Hypsignathus monstrosus). Generally cannot walk but can move along branches in hanging posture. Teeth modified for feeding on fruit and flowers.
All bats have a generally similar appearance in flight, dominated by the expanse of the wings, but they vary considerably in size. The order is usually divided into two well-defined suborders: the Megachiroptera (the large Old World fruit bats) and theMicrochiroptera (small bats found worldwide). Among members of the Megachiroptera, flying foxes (Pteropus) have a wingspan of 1.5 metres (about 5 feet) and a weight of 1 kg (2.2 pounds). The largest insectivorous bat is probably the naked, or hairless, bat (Cheiromeles torquatus); it weighs about 250 grams (about 9 ounces). The largest of the carnivorous bats (and the largest bat in the New World) is the spectral bat (Vampyrum spectrum), also known as the tropical American false vampire bat, with a wingspan of over 60 cm (24 inches). The tiny hog-nosed, or bumblebee, bat (Craseonycteris thonglongyai) of Thailand is one of the smallestmammals. It has a wingspan of barely 15 cm (6 inches) and weighs about 2 grams (about 0.07 ounce).
Bats vary in colour and in fur texture. Facial appearance, dominated by the muzzle and ears, varies strikingly between families and often between genera. In several families a complex fleshy adornment called the nose leaf surrounds the nostrils. Although the exact function of these facial appurtenances has yet to be determined, scientists believe they may help to direct outgoing echolocation calls (see belowOrientation). Wing proportions are modified according to mode of flight. The tail and the membrane between the legs also differ, perhaps as adaptations to feeding, flight, and roosting habits. Finally, bats vary in the postures they assume when roosting, particularly in whether they hang suspended or cling to a wall and in the manner in which the wings are folded and used.
Distribution
Bats are particularly abundant in the tropics. In West Africa, for example, more than 30 genera embracing nearly 100 species have been cataloged; in the United States19 genera, totaling about 45 species, are known. Of the 18 bat families, 3—the vesper bats (family Vespertilionidae), free-tailed bats (family Molossidae), and horseshoe bats (family Rhinolophidae)—are well represented in the temperate zones. A few American leaf-nosed bats (family Phyllostomidae) range into mild temperate regions. Several vesper bats range well into Canada.
The Vespertilionidae are found worldwide except in the polar regions and on isolated islands. The brown bats of genus Myotis have a range almost equal to that of the entire order. The free-tailed bats and sheath-tailed bats (family Emballonuridae) also encircle the Earth but are restricted to the tropics and subtropics. The horseshoe bats extend throughout the Old World, the roundleaf bats (family Hipposideridae) and Old World fruit bats (family Pteropodidae) throughout the Old World tropics, and the leaf-nosed bats throughout the New World tropics and slightly beyond. The other families have more restricted ranges.
Importance to humans
Most bats are insectivorous, and they are important to humans primarily for their predation on insects, for pollination, and for seed dispersal. Little is known of the spectrum of insect species consumed, but the sheer quantity is formidable. The Mexican free-tailed bats of Texas have been estimated to consume about 9,100 metric tons (10,000 tons) of insects per year. Bats would thus seem to be important in the balance of insect populations and possibly in the control of insect pests.
Some bats feed on pollen and nectar and are the principal or exclusive pollinators of a number of tropical and subtropical plants. Others feed on fruit and aid in dispersing seeds, although bananas and figs must in some cases be protected from fruit-eating bats by early harvest or by nets.
Vampire bats (family Phyllostomidae, subfamily Desmodontinae) are considered serious pests of livestock in some parts of tropical America because the small wounds they cause provide egg-laying sites for parasites and because the vampires may transmit rabies and trypanosomiasis to cattle. Other bats also carry rabies or related viruses.
The guano (droppings) of insectivorous bats is still used for agricultural fertilizer in many countries and in the past was used as a source of nitrogen and phosphorus for munitions. Large guano deposits, in addition, cover and thus preserve many archaeologically interesting artifacts and fossils in caves.
In tropical regions large colonies of bats often inhabit houses and public buildings, where they attract attention because of their noisiness, guano, and collective odour. In western culture bats have been the subject of unfavourable myths; in parts of the Orient, however, these animals serve as symbols of good luck, long life, and happiness. In some parts of Southeast Asia and on some Pacific islands, flying foxes (Pteropus) are hunted for food. Small bats are also widely but irregularly eaten.
Certain physiological aspects of some bats, particularly those involving adaptations for long hibernation, daily lethargy, complex temperature regulation, acoustical orientation, and long-distance migrations, are of interest to biologists.
In species and numbers, bats constitute an important and generally nonintrusive form of wildlife. Several zoos have established interesting exhibits of bats; indeed, some flying foxes and fruit bats have been exhibited in European zoos since the mid-19th century, and they have been kept widely for research purposes. Bats are interesting pets but require specialized care.
Natural history
ACTIVITY PATTERNS
Nocturnal activity is a major feature of the behavioral pattern of bats: nearly all species roost during the day and forage at night. Carnivorous bats, vampire bats, and perhaps fishing bats (see bulldog bat) may have an advantage at night over inactive or sleeping prey. In addition, nocturnal flight protects bats from visual predators, exposure to the sun, high ambient temperature, and low relative humidity. The large area of naked wing skin might mean that bats would absorb rather than radiate heat if they were active during the day. They would also lose body water required for temperature regulation and would then be forced to forage near water or somehow retain more water (and thus more weight) in their bodies during flight.
The nocturnal activity pattern in bats is probably kept in synchrony with changing day lengths by their exposure to light at dusk or dawn. Bats often awaken and fly from the cave exit well before nightfall. Should they be too early, their internal clock may be reset. A few species of bats, including a flying fox (Pteropus samoensis), theyellow-winged bat (Lavia frons), and the greater sac-winged bat Saccopteryx bilineata, may forage actively during the day, but little is yet known of their special adaptations.
LOCOMOTION
Flight is the primary mode of locomotion in all bats, although the flight styles vary. Some groups (the free-tailed bats, for example) are adapted for flight in open spaces and high altitudes. They have long, narrow wings, swift flight, and a large turning radius. Slit-faced bats (Nycteridae), false vampire bats (Megadermatidae), and others are adapted for hovering as they pick prey off vegetation or feed on flowers. These bats have short, broad wings, slow flight, and a small turning radius. Some bats take flight easily from the ground: members of the genus Macrotus do so simply by flapping, while vampire bats (Desmodus) leap into the air and then spread their wings and fly. The free-tails, however, roost well above the ground because, upon takeoff, they fall before becoming airborne.
Though flight speeds in the wild are hard to measure, four vesper bat species, carefully observed, have been timed on average at 18.7 to 33.3 km (11.7 to 20.8 miles) per hour. In flight the posture of each of the four fingers incorporated into the wing is under precise and individual control. Finger and arm postures, which determine the shape, extension, and angle of the wings, govern such actions as turning, diving, landing, and hovering. Except when interrupted by insect catches or obstacles, bat flight paths are straight. Insects may be pursued and captured at a rate of up to two per second; during each catch the flight path is interrupted and thus appears erratic.
In many cases there is little locomotion other than flight. Bats that hang in caves may move across the ceiling by shifting their toehold, one foot at a time. A few genera, especially among the Old World fruit bats (family Pteropodidae), may crawl along branches in a slothlike posture, using their thumb claws as well as their feet. Thesheath-tailed bats (family Emballonuridae) and mouse-tailed bats (family Rhinopomatidae) hang on vertical surfaces suspended by their hind claws but with their thumbs and wrists propped against the surface. In this orientation they can scramble rapidly up or down and forward or backward, as well as sideways.
Bats of many families walk or crawl on either horizontal or vertical surfaces, using hind feet, wrists, and thumbs. Many move freely either backward or forward, a convenience for entering and leaving crevices. The vampire bats may also leap from roost to roost. The disk-winged bats (family Thyropteridae) and sucker-footed bat(one species, family Myzopodidae), as well as the bamboo bats (Tylonycteris), have specialized wrist and sole pads for moving along and roosting on the smooth surface of leaves or bamboo stalks. Bats are not known to swim in nature except, perhaps, by accident. When they do fall into water, however, they generally swim competently.
ROOSTING
Bats choose a variety of diurnal roosts, although the roost requirements of many bats, which are rather precise in terms of light, temperature, and humidity, limit their distribution. Each species favours a particular kind of roost, though this varies with sex, season, and reproductive activity. Many bats prefer isolated or secure roosts—caves, crevices in cliff faces, the interstices of boulder heaps, tree hollows, animalburrows, culverts, abandoned buildings, portions of buildings inaccessible to humans or infrequently accessed by them (i.e., a roof, attic, or hollow wall), or the hollow core of bamboo stalks. Some species roost externally—on tree trunks or in the branches of trees, under palm leaves, in unopened tubular leaves, or on the surface of rocks or buildings. For some the darkness, stability of temperature and humidity, and isolation from predators provided by caves and crevices seem essential. Others prefer the heat and dryness of sun-exposed roosts. Many bats also occupy nocturnal roosts, often rocky overhangs or cave entrances, for napping, for chewing food, or for shelter from bad weather. Many species likewise choose special nursery or hibernation roosts. Buildings are so widely exploited by bats (especially vesper bats, free-tailed bats, and sheath-tailed bats) that many species have probably become more abundant since the advent of architecture.
Bats are usually colonial; indeed, some form very large cave colonies. Generally, large colonies are formed by bats that roost in dense clusters, pressing against one another, although many are widely spaced and do not touch when roosting. Some of the Old World fruit bats strikingly defoliate the trees on which they roost. In trees flying foxes (Pteropus) may form outdoor camps numbering hundreds of thousands of individuals. Many species form smaller groups of several dozen to several hundred. Less commonly, bats are solitary; sometimes the adult female roosts only with its most recent offspring. Occasionally, one sex is colonial and the other is apparently solitary. The advantages of colonial or solitary life and the factors that govern colony size in bats with colonial predilection have not yet been established.
Elaborate communities of other animals are often satellites of cave-bat colonies. Among these are cave crickets, roaches, blood-sucking bugs, a variety of parasites (e.g., fleas, lice, ticks, mites, and certain flies), and dermestid beetles and other insects that feed on cave-floor debris—guano, bat and insect corpses, and discarded pieces of food or seeds. Molds and other fungi are also conspicuous members of the cave-floor community. Bats and their excretions alter the cave environment by producing heat, carbon dioxide, and ammonia.
MIGRATION
Many bats of temperate climates migrate annually to and from summer roosts and winter hibernation sites, with an individual often occupying the same roosts in seasonal sequence each year. Members of the same species may converge on a single hibernation cave or nursery roost from many directions, which indicates that the choice of migration direction to and from these caves cannot be genetically determined. When migration occurs, however, is probably genetically determined (i.e., instinctive) and influenced also by weather conditions and the availability of food. Nothing is known of how bats recognize migration goals or how succeeding generations learn their locations. Female young born at a nursery roost may memorize its location, but how they know where to go at other times is not clear. Likewise, little is yet known of energy storage, navigation, or other specializations for migrations.
Female Mexican free-tailed bats migrate from central Mexico to Texas and adjacent states each spring, returning south in the fall. Mating probably occurs in transient roosts during the spring flight. The migration is believed to remove pregnant and lactating females to a region of high food supply where they need not compete with males of their own species. Presumably they return to Mexico for its suitable winter climate and food supply and to meet their mates.
The North American red and hoary bats (Lasiurus borealis and L. cinereus) and thesilver-haired bat (Lasionycteris noctivagans) migrate in the fall from the northern United States and Canada to the southern United States and beyond, returning in the spring.
ORIENTATION
Bats of the suborder Microchiroptera orient acoustically by echolocation (“sonar”). They emit short high-frequency pulses of sound (usually well above the range of human hearing) and listen to the echoes returning from objects in the vicinity. By interpreting returning echoes, bats may identify the direction, distance, velocity, and some aspects of the size or nature (or both) of objects that draw their attention.Echolocation is used to locate and track flying and terrestrial prey, to avoid obstacles, and possibly to regulate altitude; orientation pulses may also serve as communication signals between bats of the same species. Rousette bats(megachiropteran genus Rousettus) have independently evolved a parallel echolocation system for obstacle avoidance alone. Echolocation pulses are produced by vibrating membranes in the larynx and emitted via the nose or the mouth, depending upon species. Nose leaves in some species may serve to channel the sound.
The echolocation signals spread in three dimensions on emission, the bulk of the energy in the hemisphere in front of the bat or in a cone-shaped region from the nostrils or mouth. When the sound impinges on an intervening surface (an insect or a leaf, for example), some of the energy in the signal is reflected or scattered, some absorbed, and some transmitted and reradiated on the far side of the surface; the proportion of sound energy in each category is a function of wavelength and of the dimensions, characteristics, and orientation of the object. The reflected sound spreads in three dimensions, and some portion of it may impinge on the bat’s ears at perceptible energy levels.
Bats’ external ears are generally large, which probably enhances their value for detecting the direction of incoming signals, and their middle and inner ears are specialized for high-frequency sensitivity. In addition, the bony otic (auditory) complex is often isolated acoustically from the skull, which probably improves signal comparison by both ears. The thresholds andranges of hearing in several genera of bats have been studied, and in each case the region of maximum sensitivity has been found to coincide with the prominent frequencies of the outgoing echolocation signals.
The characteristics of echolocation pulses vary with family and even with species. Echolocation pulses of a substantial number of bat species have been analyzed in terms of frequency, frequency pattern, duration, repetition rate, intensity, and direction. The prominent frequency or frequencies range from 12 kilohertz (1 kilohertz is equivalent to 1,000 hertz, or cycles per second) to about 150 kilohertz or more. Factors influencing frequency may include bat size, prey size, the energetics of sound production, inefficiency of the propagation of high frequencies, and ambient noise levels.
Orientation pulses may be of several types. The individual pulse may include a frequency drop from beginning to end (frequency modulation [FM]), or the frequency may be constant (CF) during part of the pulse, followed by a brief FM sweep; either FM or CF pulses may have high harmonic content. The pulse duration varies with the species and the situation. During cruising flight the pulses of the greater false vampire bat (Megaderma lyra) are 1.5 milliseconds (0.0015 second), those ofWagner’s mustached bat (Pteronotus personatus) 4 milliseconds, and those of thegreater horseshoe bat (Rhinolophus ferrumequinum) 55–65 milliseconds. In goal-oriented flight, such as the pursuit of an insect or the evaluation of an obstacle or a landing perch, the pulse duration is systematically altered (usually shortened) with target distance, sometimes ending with pulses as short as 0.25 millisecond.
During insect pursuit, obstacle avoidance, and landing maneuvers, there are three phases of pulse output design: search, approach, and terminal. The search phase, during which many bats emit about 10 pulses per second, precedes specific attention to a target. In the approach phase, which starts when the bat detects an object to which it subsequently devotes its attention, the bat raises the pulse rate to about 25 to 50 per second, shortens the pulses with decreasing distance, and often alters the frequency pattern. The terminal phase, which often lasts about 100 milliseconds, is characterized by extremely short pulses, repeated as rapidly as 200 or more times per second, and ceases as the bat intercepts the target or passes it (the stimulus being, perhaps, the cessation of echoes); another search phase follows. During the brief terminal phase (a fraction of a second), the bat is engaged in final interception (or avoidance) maneuvers and appears to pay little attention to other objects.
In addition to sensitive ears, the use of echolocation to gain sensory information requires integration of the vocal and auditory centres of the brain. Not only must the nervous system of the bat analyze in a few thousandths of a second the reflected, and thus altered, form of its own pulse, but it must separate this echo from those of other individuals and from others of its own pulses. All of this must be done while the animal (and often the target) is moving in space. In the laboratory, bats have been found to be able to identify, pursue, and capture as many as two fruit flies (Drosophila, about 3 mm [0.12 inch] long) per second and to locate and avoid wires as fine as 0.1 or even 0.08 mm (0.004 or 0.003 inch) in diameter.
Research has provided some information on the mechanisms of bat sonar. There is evidence that the multiple frequencies of FM or harmonic patterns serve in determining target direction. The relative intensities of the various frequencies are different at each ear, which allows the animal to determine the target’s direction when three or more frequencies are received. Target velocity may be measured by CF bats through the use of the Doppler shift, a change in perceived frequency due to the relative motion of the bat and its target. Changes in pulse-echo timing may provide information on target distance and velocity. The ratio of useful signal to background noise is increased by several mechanisms, including specializations of the middle ear and its ossicles (tiny bones), isolation of the cochlea (the area where sound energy is converted into nerve impulses), and adaptations of the central nervous system.
FOOD HABITS
Most bats feed on flying insects. In some cases prey species have been identified from stomach contents or from discarded pieces under night roosts, but such studies have not yet provided an adequate measure of the spectrum of bat diets. Bats identify and track insects in flight by echolocation. Large insects may be intercepted with the wing membranes and pulled into the mouth. Some bats feed on arthropods, such as large insects, spiders, and scorpions, that they find on the ground, on walls, or on vegetation. These bats may either land on and kill their prey before taking off with it or pick it up with their teeth while hovering.
Two genera (Noctilio and Myotis) include at least one species that catches small fishand possibly crustaceans. All fish-eating species also feed on flying insects or have close relatives that do so. Each is specialized in having exceptionally large hind feet armed with long, strong claws with which the fish are gaffed.
The Megachiroptera and many of the phyllostomidgenera feed on a variety of fruits, often green or brown in colour; usually such fruits are either borne directly on wood or hang well away from the bulk of the tree and have a sour or musky odour.
The Old World fruit bat subfamily Macroglossinae (and some other fruit bats) and certain leaf-nosed bats feed, at least in part, on nectar and pollen. Many tropical flowers, adapted forpollination by these bats, open at night, are white or inconspicuous, have a sour, rancid, or mammalian odour, and are borne on wood, on pendulous branches, or beyond or above the bulk of the plant. The phyllostomid Glossophaginae may also feed on flowers. (See Sidebar: Bat-Loving Flowers.)
Several phyllostomid and megadermatid genera are carnivorous, feeding on small rodents, shrews, bats, sleeping birds, tree frogs, and lizards. The true vampires, which feed on the blood of large mammals or birds, land near a quiet prospective victim, walk or jump to a vulnerable spot on it where the skin is relatively exposed—the edge of the ear or nostril, around the anus, or between the toes, for example—make a scooping, superficial bite from which the blood oozes freely, and lap the blood with very specialized tongue movements. Each vampire requires about 15 millilitres (about half an ounce) of blood per night.
The interaction of bats with their food, be it insects, fruit, or flowers, probably has a substantial impact on some biological communities. Many plants are dependent on bats for pollination; other plants benefit from seed dispersal by bats. Moths of two families are known to take evasive or protective action on hearing bat pulses nearby, an adaptation that implies heavy predation.
MAINTENANCE BEHAVIOUR
Bats are meticulous in their grooming, spending a fair part of the day and night combing and grooming their fur and cleansing their wing membranes. Generally, they comb with the claws of one foot while hanging by the other; they remove the combings and moisten their claws with their lips and tongue. On the wing membranes in particular, they use the mouth meticulously, perhaps oiling the skin with the secretions of dermal (skin) glands while cleansing it.
SOCIAL INTERACTIONS
Although social interactions per se have not been observed between adult bats, they are known to often segregate by sex. As noted above, pregnant females in many species occupy special nursery roosts until their young are independent. In some species the sexes occupy the same general roost but gather in separate clusters. In others the sexes intermingle or arrange themselves into a pattern within a group—the females centrally, for example, and the males peripherally. Sexual segregation during foraging has been reported for several species. Among bats that migrate over long distances, such as Mexican free-tailed, red, and hoary bats, the sexes may meet only briefly each year.
Life cycle
Details of the life cycle are known for only a few species. In northern temperate zone species, there is an annual cycle of sexual activity, with birth taking place between May and July. In males the testes, normally located in the abdominal region, descend seasonally into the scrotum, and active spermatogenesis occurs. In females sexual receptivity may be associated with egg maturation and release. Tropical bats may exhibit a single annual sexual cycle or may be diestrous (i.e., have two periods of fertility) or polyestrous (have many).
The sexual cycles of entire populations are closely synchronized, so almost all mating occurs within a few weeks. The periods of gestation, birth, lactation, and weaning are also usually synchronized. Gestation varies in duration: five or six months in flying foxes (Pteropus), more than five months in vampire bats (Desmodus), three months in some small leaf-nosed bats (Hipposideros), and 6 or 7 to 14 weeks in several small vesper bats (family Vespertilionidae). The length of gestation may be influenced by both ambient (surrounding) and body temperature.
In several North American and northern Eurasian vesper and horseshoe bats that hibernate, copulation occurs in the fall, and the sperm are stored in the female genital tract until spring. Ovulation, fertilization, and implantation occur after emergence from hibernation, when the female again has available an abundant food supply and a warm roost. Such favourable environmental conditions greatly enhance the young bat’s chances of survival.
Most bats bear one young, but the big brown bat (Eptesicus fuscus) may bear twins, and the Eastern red bat (Lasiurus borealis) bears litters of one to four.
At birth the young, which may weigh from one-sixth to one-third as much as the mother, usually have well-developed hind legs with which they hold on to their mother or to the roost. Their wings are very immature. The young are hairless or lightly furred and are often briefly blind and deaf. Female bats normally have one pectoral (at the chest) or axillary (at the armpit) mammary gland on each side. Several species that carry their young while foraging also have a pair of false pubic nipples, which the infant may hold in its mouth when its mother flies. The infants are nourished by milk for a period of about five or six weeks in many small bats and for five months in the Indian flying fox (Pteropus giganteus). By two months of age, most smaller bats have been flying and foraging for three or four weeks and have achieved adult size.
In many species females late in pregnancy migrate to special nursery roosts, in which large numbers of pregnant females may aggregate, usually to the exclusion of nonpregnant females, males, and bats of other species. In some cases the nursery roosts seem to be chosen for their high temperature, which may derive from the sun, from the bats themselves, or from decomposing guano. When foraging, some bats (Erophylla) leave their infants hanging quietly, one by one, on the cave wall or ceiling. In the case of the Mexican free-tailed bat and a few others, the closely spaced infants may move about and mingle on the wall. Some bats carry their young with them for a short period of time. Generally, each mother, on returning to her roost, seeks out her own offspring by position, smell, and acoustical exchange.
Some bats achieve sexual maturity in their first year, others in their second. Infant mortality appears to be high. Developmental and genetic errors and disease take their toll, but accidents seem to cause more serious losses—the young may fall from the ceiling or perhaps have serious collisions in early flight attempts. A fair number of bats probably fail to make the transition from dependent infants to self-sufficient foragers.
Adult bats, on the other hand, have low mortality. Predation is rarely serious, especially for cave-dwelling species. Disease, parasitic infestation, starvation, and accidents apparently take small tolls. There are records of several big brown (Eptesicus fuscus), little brown (Myotis lucifugus), and greater horseshoe bats(Rhinolophus ferrumequinum) that have lived more than 20 years, and a few have lived more than 30. Probably many bats in temperate climates live more than 10 years. Longevity has not been established for most tropical species, but a few are known to live for more than 10 years.
Several factors probably contribute to the unusual longevity of bats. Generally isolated roosts and nocturnal flight substantially protect them from predation, from some elements of weather, and from exposure to the sun. Their largely colonial way of life may ensure that entire populations experience contagious infection and subsequent immunity; indeed, such a pattern in the past may have hastened adaptation to disease. The persistent use of various seasonal roosts probably ensures isolation and security, food and water supplies, and access to mates. Many bats, moreover, reduce their body temperature at rest. Not only is there a probability that this conserves some cellular “machinery,” since metabolism is reduced, but fewer hours need to be spent in actively seeking food and water.
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At rest a bat’s head, especially the ears, is its most striking feature. The neck is likely to be short and relatively immobile. The projecting portion of the external ear (the pinna) is usually extremely large and often is funnel-shaped. In several genera that feed on terrestrial arthropods, the ears are particularly oversized, probably for highly precise directional assessment. A projection on the front side of the auditory canal (the tragus) or another on the rear side (antitragus) may also be conspicuous. The ears are often highly mobile, sometimes flicking back and forth in phase with the production of sonar signals. In some species the ears are immobile, but in all cases they probably function in tandem for directional analysis.
Bats often have a rodentlike or foxlike muzzle, but in many the face has a pushed-in puglike appearance. In the nectar feeders the snout is elongated to house the long extensible tongue. Many bats have a facial ornament, the nose leaf, which consists of skin and connective tissue. It surrounds the nostrils and extends as a free flap or flaps above the nostrils and in front of the face. The complexity and shape of the nose leaf varies with family; its presence correlates with nasal emission of orientation signals. Thus, it is supposed that the nose leaf influences sound output, perhaps by narrowing the beam, but evidence is sparse.
Most bats are well furred except for the wing membranes. Colours are generally shades of brown, tan, gray, or black on top and lighter shades on the underside. Red, yellow, or orange variants occur in many species. Speckled or mottled patterns are common, as are bright or light-coloured spots or stripes. Bright red, yellow, or orange shading on the head, neck, and shoulders is not unusual. Mottled fur may enable the bat to be inconspicuous on lichen-covered bark or rock. Bright spots may simulate the speckled sunlight of the forest canopy as seen from below. Stripes probably break up contours. The colouring seen while the animal is hanging may be a kind of countershading for concealment, or it may enhance the bat’s simulation of a ripening fruit or a dead leaf. Many bats that roost externally hang from a branch by one foot, which then looks like a plant stem.
When fully active, bats have a body temperature of about 37 °C (98.6 °F). Although some bats maintain fairly even body temperatures, a large number undergo periodic raising or lowering of their temperature. Many of the vesper bats and horseshoe bats and a few free-tailed bats reduce their body temperature to that of their surroundings (ambient temperature) shortly after coming to rest. This condition is called heterothermy. They raise their temperature again on being aroused or when readying themselves for nocturnal foraging. The drop in body temperature, if the ambient temperature is relatively low, results in a lethargic state. Energy is conserved by thus “turning down the thermostat,” but the bat is rendered relatively unresponsive to threats by predators or weather. Heterothermic bats therefore generally roost in secluded sites offering protection, often in crevices. In heterothermic bats one or more sensory systems and the brain remain sensitive at low temperatures and initiate the necessary heat production for arousal. Heat is generated by the metabolism of fat and by shivering.
Many bats that exhibit daily torpor also hibernate during the winter and therefore must store energy as body fat. In the fall these bats increase their weight by 50 to 100 percent. They must also migrate from the summer roost to a suitable hibernation site (often a cave) that will remain cool and humid throughout the winter without freezing. Large populations often aggregate in such caves. Hibernation involves the absence of temperature regulation for long periods in addition to adaptations of circulation, respiration, and renal function and the suspension of most aspects of activity. Bats of hibernating species generally court and mate in the fall when they are at their nutritional peak. During pregnancy, lactation, and juvenile growth, bats probably thermoregulate differently, more closely approximating stability.
Bats of several tropical families maintain a constant body temperature (homeothermy). This, however, depends on the nutritional state as well. A spectrum of degrees of homeothermy and heterothermy probably will be discovered.
Digestion and water conservation
Digestion in bats is unusually rapid. They chew and fragment their food exceptionally thoroughly and thus expose a large surface area of it to digestive action. They may begin to defecate 30 to 60 minutes after beginning to feed and thereby reduce the load that must be carried in flight.
Some bats live in sun-baked roosts without access to water during the day. They may choose these roosts for their heat, and thus conserve their own, but it is not yet known how they hold their body temperature down without using water. In the laboratory, bats die if body temperature rises above about 40–41 °C (104–106 °F).
In folklore, bats have been considered to be blind. In fact, the eyes in the Microchiroptera are small and have not been well studied. Among the Megachiroptera the eyes are large, but vision has been studied in detail only in flying foxes. These bats are able to make visual discriminations at lower light levels than humans can. The Megachiroptera fly at night, of course, and some genera fly below or in the jungle canopy, where light levels are very low. Except for rousette bats (Rousettus), none are known to orient acoustically.
Studies of several genera of Microchiroptera have revealed that vision is used in long-distance navigation and that obstacles and motion can be detected visually. Bats also presumably use vision to distinguish day from night and to synchronize their internal clocks with the local cycle of daylight and darkness.
The senses of taste, smell, and touch in bats do not seem to be strikingly different from those of related mammals. Smell is probably used as an aid in locating fruit and flowers and possibly, in the case of vampire bats, large vertebrates. It may also be used for locating an occupied roost, members of the same species, and the differentiation of individuals by sex. Many bats depend upon touch, aided by well-developed facial and toe whiskers and possibly by the projecting tail, to place themselves in comforting body contact with rock surfaces or with other bats in the roost.
The fossil record of bats prior to the Pleistocene Epoch (about 2,600,000 to 11,700 years ago) is limited and reveals little about bat evolution. Most fossils can be attributed to living families. Skulls and teeth compatible with early bats are known from about 60 million years ago, during the Paleocene Epoch. These specimens, however, may well have been from insectivores, from which bats are clearly distinguishable only on the basis of flight adaptations. By 45 million years ago (theEocene Epoch), bats with fully developed powers of flight had evolved.
The order Chiroptera is readily divided into two suborders—Megachiroptera (large Old World fruit bats) and Microchiroptera (small bats). The Megachiroptera orient visually and exhibit a number of primitive skeletal features. The Microchiroptera orient acoustically. It is not certain that they have a common origin. The suborders either evolved separately from flightless insectivores or diverged very early in chiropteran history.
The two principal geographic centres of bat evolution appear to be the Australo-Malaysian region, with about 290 species, and the New World tropics, with about 230 species. Comparable ecological niches in the Old World and the New World are occupied largely by different genera of bats, usually of different families.
Classification
Distinguishing taxonomic features
The order Chiroptera is defined by flight and the elongated finger bones and marked pectoral specialization that support it. Weak pelvic and leg development is also a chiropteran feature. The ulna of the forearm is reduced; claws are absent on the fingers except on the thumb (and occasionally the second finger); and the knee is directed rearward and outward. The maximum complement of permanent teeth is 38, the minimum 20.
Annotated classification
The following classification is based on the third edition of Mammal Species of the World, edited by Don E. Wilson and DeeAnn M. Reeder, published in 2005. Subsequent research has shown, however, that the number of bat species continues to increase from the discovery of new forms and from the results of studies using DNA analysis to examine the evolutionary relationships between known bat species.
1,116 species in 18 families belonging to 2 suborders. Found from the tropics into temperate regions worldwide.
930 species in 17 families found from the tropics into temperate regions worldwide.
407 small to medium-sized species in 48 genera found worldwide to tree line and on many oceanic islands. Muzzle plain; eyes small; ears moderate to large; wings generally long and moderately narrow; tail and interfemoral membrane well developed. All walk well, often entering crevices. A few species eat fish. Generally colonial in dense, touching clusters; a few solitary; mostly roost in caves, attics, barns, hollow trees, boulder heaps, the twigs of birds’ nests, or roof thatching; a few (Lasiurus) roost in branches, on tree trunks, or in the hollow core of bamboo (Tylonycteris). Many temperate species hibernate and migrate; many are drably coloured; several that roost externally are either spotted (Euderma) or speckled (Lasiurus). Brown bats (Myotis and Eptesicus) and pipistrelles (Pipistrellus) found worldwide.
160 small to large species in 55 genera found from the southwestern United States through tropical America. Nose leaf simply shaped, ears often large and mobile, wings generally short and broad; tail and interfemoral membrane vary from absent to well developed; fur colour and patterns varied. Includes insectivorous, carnivorous, and fruit- and flower-feeding species. Generally do not walk. Colonial, often densely so; generally roost in touching clusters in caves, tree hollows, buildings, and culverts or in the open under bridges or eaves, in the crests of palm trees, or on the underside of palm leaves. Flight ranges from swift and straight to hovering.
77 small to moderately large species in 1 Old World genus. Complex nose leaf; large, highly mobile ears; wings short and rounded; well-developed interfemoral membrane, supported by tail. Unable to walk. Fur generally brown (occasionally red). Dark, humid roosts selected, especially caves, but tree hollows, buildings, and culverts as well. Generally colonial, nontouching. Usually feed on flying insects.
81 small to large species in 9 genera, some ranging from Old World tropics to temperate regions. Complex nose leaf with both anterior and posterior portions, the anterior portion being homologous to the “horseshoe” in the closely related Rhinolophidae. Toes have only two phalanges, owing to fusion; second finger of wing has only metacarpal bones; other fingers have two phalanges. Ears medium, without tragus. Fur colour ranging from dark brown to hues of red and yellow. Mostly colonial in caves, tunnels, or other dark retreats. Forage for flying insects.
100 robust, small to very large species in 16 genera, some ranging into mild temperate regions. Tail projects well beyond the well-developed interfemoral membrane; ears large, rather immobile, often fused to one another, and unusually shaped; lips and snout often heavy; eyes tiny. Wings very long and narrow; legs well developed for walking; toes often bearing bristles. Often have a conspicuous odour; fur short, usually dark brown or black. Many highly colonial, with millions clustering in caves, tree hollows, and buildings; dense, touching groups roost pressed against fellows or walls, often in crevices; many prefer hot, dry roosts. Walk exceptionally well. Feed on flying insects; some migrate.
51 small to medium-sized species in 13 genera found in the tropics excluding the West Indies and some Pacific islands; each genus restricted to either Old or New World. Ears large but simply shaped; eyes small; muzzle sharp but plain; tail short, perforating the dorsal surface of a well-developed interfemoral membrane. Several genera have a glandular pouch in the wing extensions. Roost on vertical surfaces, such as tree trunks, cliff faces, cave entrances, and walls; some favour buildings. Some densely colonial but not touching one another; others form small groups or are solitary. Hang suspended from toes with wrists propped against wall.
16 small to medium-sized species in 1 genus (Nycteris) distributed through most of tropical Africa and Southeast Asia. Head with peculiar nasal depression, cleft nose leaf, and a deep midline facial cleft behind and above the nostrils. Ears large; wings broad; tail long with split end; heel bones greatly elongated; well-developed interfemoral membrane. Prey mostly on terrestrial insects resting on vegetation, rocks, or walls. Unable to walk. Roosts usually dark and humid; some species roost exposed in forest canopy.
10 small species in 2 genera of tropical Central and South America. Some walk. All lack nose leaf but have elaborate lip leaves. Tail and interfemoral membrane well developed. Colour ranges from brown through orange, red, and yellow. Feed on flying insects. Densely colonial in dark caves, population of colonies often numbering in the tens of thousands.
5 moderately large species in 4 genera of Central Africa, Southeast Asia, and Australia. External ears very large and fused across midline; nose leaf large with truncated end; eyes relatively large; wings broad; interfemoral membrane well developed and supported distally by heel bones; no external tail. Feed principally on terrestrial arthropods, as in Nycteridae; at least 2 species also feed on small vertebrates taken in the same fashion as arthropod prey. Unable to walk. Form small colonies of nontouching individuals usually in dark, secluded caves or abandoned buildings. Yellow-winged bat (Lavia frons) is at least partly diurnal and roosts in trees in the savanna and open forest.
8 species of small, slenderly built bats in 3 genera (Natalus) of Central America, northern South America, and the West Indies. Thick gray, buff, yellow, or reddish fur. Well-developed tail and interfemoral membrane. Ears large; snout plain. Walk clumsily and do not enter crevices; cave-dwelling and colonial in nontouching groups. Feed on flying insects.
4 small species in 1 genus (Rhinopoma) of North Africa and tropical Asia. Tail very long and largely free beyond a narrow interfemoral membrane. Ears large; small nose leaf; primitive shoulder girdle.
3 species in 1 genus (Thyroptera) of Central America and northern South America, excluding West Indies. Base of thumb and sole provided with sucking disk; simple muzzle; ears large; second finger reduced to rudiment. Roost alone or in small groups, often in still-furled banana leaves. Biology poorly known.
2 small species in 1 genus (Mysticina) of New Zealand. Simple head similar to that of vesper bats. Wings fold very compactly; thumb and toe claws long and sharp; highly adapted for walking; tail perforates interfemoral membrane dorsally. Feed on flying and terrestrial insects. Biology poorly known.
2 small, delicately built species in 2 genera of northern South America and Central America. Snout plain; tail long, ending short of the well-developed interfemoral membrane; thumb vestigial; legs long; feet small. Biology unknown; probably insectivorous.
2 medium-sized species in 1 genus (Noctilio) of the American tropics. Muzzle heavy but unadorned; lips full; internal cheek pouches; ears large, pointed, and mobile; wings long and narrow. Tail well developed, extending to midpoint of large interfemoral membrane, which is pierced dorsally by tail tip; interfemoral membrane supported by very well-developed bones. Insectivorous, but one species also uses its very large feet to capture fish. Musky odour. Walk well; often roost in crevices, tree hollows, attics, grottoes, and caves; colonial, in touching clusters.
1 tiny species of Thailand, Craseonycteris thonglongyai, perhaps the smallest living mammal.
1 species in 1 genus (Myzopoda) endemic to Madagascar. Small, plain muzzle; large ears with peculiar mushroom-shaped lobe. Thumb and sole with adhesive disks; vestigial thumb claw; tail extends free beyond interfemoral membrane. Probably insectivorous; biology unknown.
186 generally large species in 42 fruit- or flower-feeding genera found in the Old World tropics and subtropics, including many Pacific islands. Lack acoustic orientation except rousette bats (Rousettus); ears small; eyes large, with vision well developed; generally roost in trees; often colonial; often show countershading, cryptic markings, or bright fur colours or patterns. Index finger generally clawed; tail short or lacking; interfemoral membrane reduced. Muzzle simple in appearance, except in the hammer-headed fruit bat (Hypsignathus monstrosus). Generally cannot walk but can move along branches in hanging posture. Teeth modified for feeding on fruit and flowers.
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