American leaf-nosed bats

(Phyllostomidae)

Class Mammalia

Order Chiroptera

Suborder Microchiroptera

Family Phyllostomidae

Thumbnail description
Small to relatively large bats, most of which have a fleshy, triangular nose-leaf projecting above the nostrils

Size
Head and body length 1.6–5.3 in (40–135 mm); tail 0–2.2 in (0–55 mm); forearm 1.2–4.1 in (31–105 mm); weight 0.2–6.8 oz (5–190 g)

Number of genera, species
49 genera; 151 species

Habitat
Mainly forests, but also deserts

Conservation status
Extinct: 1 species; Endangered: 4 species; Vulnerable: 25 species; Lower Risk/Near Threatened: 36 species; Data Deficient: 5 species

Distribution
Subtropical and tropical America from southern Arizona and the West Indies to northern Argentina

Evolution and systematics

Members of this family are among the most common mammals in American tropical forests. Rainforests in Central and South America contain 31–49 species of phyllostomids; tropical forests receiving less rainfall contain 20–30 species; and dry regions contain two to three species. In addition to being one of the most taxonomically diverse families of bats, the Phyllostomidae is the most ecologically diverse group of bats. Food habits range from insects, flowers, and fruit to blood and other vertebrates. Sizes of social groups range from monogamous pairs to colonies containing several hundred thousand individuals.

Along with the two small American families, Noctilionidae (one genus, two species) and Mormoopidae (two genera, eight species), the Phyllostomidae is classified in superfamily Noctilionoidea of suborder Microchiroptera. Food habits of noctilionids (also known as bulldog bats) include insects and fish, while mormoopids (also known as moustached or leaf-chinned bats) are insectivorous. These two families are unknown in the fossil record other than from Pleistocene deposits. The earliest known fossil, phyllostomid (Notonycteris), comes from the Miocene of Colombia (about 20 million years ago). It is likely that each of these families evolved in South America. All three families occur in the West Indies as well as in mainland subtropical and tropical America.

Reflecting its substantial dietary diversity, the Phyllostomidae is currently classified into eight subfamilies containing a total of 49 genera and 151 species. Subfamilies and their diversity include: Phyllostominae (11 genera, 37 species), Lonchophyllinae (3, 9), Brachyphyllinae (1, 2), Phyllonycterinae (2, 4), Glossophaginae (10, 23), Carolliinae (2, 7), Stenodermatinae (17, 66), and Desmodontinae (3, 3). Phyllostomines include insectivores and carnivores; lonchophyllines, phyllonycterines (a West Indian group), and glossophagines are nectarivores; brachyphyllines (a West Indian group), carolliinines, and stenodermatines are frugivores; and desmodontines are blood-feeding vampires.

Physical characteristics

Sizes and facial characteristics of American leaf-nosed bats differ substantially among the subfamilies and reflect their diverse

feeding adaptations. Phyllostomines include the largest members of the family (3.2–6.7 oz; 90–190 g) and generally have the longest nose leaves. The nose leaf of the sword-nosed bat (Lonchorhina aurita), for example, is as long as its long, pointed ears. Large members of this subfamily have robust canines and molars for killing and chewing vertebrate prey. At the other end of the size spectrum are flower-visiting bats (Lonchophyllinae and Glossophaginae), which weigh 0.2–0.8 oz (7–25 g). These bats have elongated muzzles, small nose-leaves, long tongues, and dentition that are reduced in size and number. Fruit-eating bats (Stenodermatinae) weigh 0.2–2.8 oz (5–80 g), have medium-sized nose leaves, and generally have flattened faces with dagger-like canines and broad cheek teeth for grabbing and crushing fruit. Vampire bats (Desmodontinae) weigh 0.2–1.7 oz (20–50 g) and have much-reduced nose leaves and a reduced number of morphologically specialized teeth. The two upper incisors are sharp and chisel-like for making incisions in the skin of mammals or birds.

The fur color of phyllostomid bats is generally brown or gray, but one species (Ectophylla alba) is white. Many genera of stenodermatines, which often roost in foliage by day, have white facial stripes, and a few have a white mid-dorsal stripe.

The nose-leaves that give this family its common name are also found in several families of Old World Microchiroptera (e.g., Nycteridae, Rhinolophidae, Hipposideridae, and Megadermatidae). All of these bats emit echolocation sounds

through their nostrils rather than through their mouths. It is thought that these structures serve as an acoustic lens that focuses the outgoing sound into a narrow beam.

Distribution

Phyllostomid bats are currently distributed from the southwestern United States and the West Indies south to northern Argentina and central Chile. Two subfamilies (Phyllonycterinae and Brachyphyllinae) are restricted to the West Indies, which generally lack members of the Phyllostominae and Carolliinae. Lonchophyllinae and Desmodontinae are absent from the West Indies, although the common vampire, Desmodus rotundus (Desmodontinae), is known as a fossil from Cuba. In the late Pleistocene, vampire bats had a much broader distribution and occurred across the southern United States. Highest diversities of phyllostomid bats, in terms of number of subfamilies and species, occur in the lowland rain-forests of northwestern South America and adjacent Central America. Phyllostomid diversity declines with altitude, latitude, and increasing aridity.

Habitat

Most phyllostomid bats are forest-dwellers. They live in a wide range of forest habitats, including tropical rainforests, tropical dry forests, and subtropical cloud forests. A few species, including two species of Leptonycteris and Choeronycteris mexicana (Glossophaginae) in Mexico and Platalina genovensium (Lonchophyllinae) in Peru, inhabit deserts or very dry tropical forests where they are important pollinators of columnar cacti and century plants (agaves).

Daytime roost structures of American leaf-nosed bats are quite diverse. Most species live in caves and/or hollow trees, but alternate roosts include mines, culverts, hollow logs, under tree roots, undercut river banks, houses, abandoned termite nests, and tree foliage. Suitable roosts are extremely important in the ecology of these bats and can sometimes limit their abundance and distribution. The relatively recent local extinction of many species of mormoopid and phyllostomid bats in the West Indies, for example, was likely caused by the inundation of extensive cave systems as a result of post-Pleistocene increases in sea level.

In addition to providing protection from inclement weather and predators, cave roosts provide stable microclimates. Most cave-dwelling phyllostomids live in roosts that are at or slightly below outside ambient temperatures. These temperatures are often below the thermoneutral zones of the bats, which forces them to expend energy to maintain a constant, high body temperature (98.6–100.4°F; 37–38°C). Certain glossophagine bats, however, including Leptonycteris curasoae and Monophyllus redmani, live in hot, humid caves where temperatures reach 91°F (33°C) within their thermoneutral zones. These caves are hot because they trap the body heat of tens of thousands of phyllostomid and mormoopid bats. In addition to reducing individual daily energy costs and rates of evaporative water loss,

this roosting strategy increases the developmental rates of embryos and lactating babies.

Most members of the Stenodermatinae usually roost either solitarily or in small groups in foliage and, hence, are constantly exposed to (shaded) ambient temperatures. In addition, several species (e.g., Uroderma bilobatum, Artibeus watsoni, and Ectophylla alba) are known to construct "tents" by clipping the leaves of banana-like herbs, philodendrons, and palms to form shelters from rain and predators. Species of Rhinophylla (Carolliinae) also use tents as day roosts. Tent roosts are short-lived, which forces groups of bats to constantly change roost sites.

Behavior

Most phyllostomid bats are colonial rather than solitary roosters, but roost size varies tremendously within and between species. Modal roost sizes within most species are relatively small and range from a few individuals to a few thousand individuals. A few species (the hot cave bats), in contrast, are highly colonial and roost in groups of tens of thousands to several hundred thousand individuals. Except for the flower-visiting, hot-cave bats, there does not appear to be a strong correlation between feeding habits and average roosting group size in this family. Vampires, fruit-eaters, many flower visitors, and certain carnivore/omnivores (e.g., Phyllostomus hastatus) sometimes live in roosts containing thousands of individuals. Small roost groups occur in pure carnivores, insectivorous phyllostomines, and a variety of flower-visiting and fruit-eating species (especially the tent-making stenodermatines).

Seasonal sexual segregation commonly occurs in many bats, including phyllostomids. Segregation usually involves the formation of maternity roosts by females. This behavior is most strongly developed in migratory glossophagines such as Leptonycteris curasoae, in which females sometimes form maternity roosts containing tens of thousands of adults. Single-sex maternity roosts also occur in non-migratory glossophagines and members of other subfamilies.

Like other Microchiroptera, phyllostomids use ultrasonic sounds for foraging and communication. Unlike other microbats, however, most phyllostomids produce very low intensity sounds that contain only about one-thousandth the sound energy as similar-sized vespertilionid bats. Additional characteristics of their echolocation sounds include multiple harmonics, frequency modulation, and short duration. These sounds provide short-range (3.2–6.5 ft; 1–2 m) information about potential insect or other prey items in areas of high vegetation clutter. In addition to echolocation information, many phyllostomids use other sensory modes (e.g., vision such as the ground-feeding insectivore, Macrotus californicus, and many flower-visiting bats, or olfaction such as many fruit-eating bats) to find food. Certain phyllostomines, for example, locate prey using prey-generated sounds (e.g., singing katydids, Tonatia sylvicola, and singing male frogs, Trachops cirrhosus). Careful experiments have shown that plant-visiting phyllostomids use a combination of vision, olfaction, and echolocation to locate and gain access to their food.

Echolocation and other kinds of vocalizations are also used for communication in these bats. Babies communicate with their mothers using "double-note" calls (rapidly repeated series

of long and short notes); these calls serve to reunite mother and baby in the roost after a female returns from foraging. Female harem-mates in Phyllostomus hastatus communicate while foraging with loud screeching calls. Certain stenodermatine bats (e.g., Artibeus jamaicensis and Uroderma bilobatum) produce intense warning calls that attract conspecifics or other species when they are captured in Japanese mist nets or while being handled.

Territorial behavior away from day roosts appears to be uncommon in phyllostomids bats. Male home ranges of the tent-roosting frugivore (Rhinophylla pumilio) do not overlap and are thought to be territories. The nectar-feeding bat (Glossophaga soricina) defends flowering stalks of agaves, at least in a suburban tropical setting, but no other evidence exists to suggest that flower-visiting or fruit-eating phyllostomids defend their food plants. Instead, it is common to see many individuals of several species feeding together in fruiting or flowering trees. The nectar bat (Leptonycteris curasoae) sometimes feeds in large numbers in isolated patches of flowering columnar cactus plants. Small groups of two to four bats of this species sometimes forage together at cactus and agave blossoms.

Because they live in tropical and subtropical habitats, phyllostomid bats do not hibernate and are active year-round. Although most species are sedentary and do not migrate among habitats during the year, a few species are known to undergo seasonal migrations. Relatively short-distance (<62 mi; 100 km) altitudinal movements are known to occur in three Mexican or Central American species—two frugivores, Carollia perspicillata and Sturnira lilium, and the nectar bat, Leptonycteris curasoae. Longer distance latitudinal migrations occur in three arid-zone nectar bats: Leptonycteris curasoae, L. nivalis, and Choeronycteris mexicana. Movements of over 620 mi (1,000 km) occur in females of L. curasoae as they move between spring maternity roosts in the Sonoran Desert and their late fall mating sites in west central Mexico. The lonchophylline nectar bat (Platalina genovensium) is also thought to undergo substantial migrations in the Andes of Peru.

Feeding ecology and diet

American leaf-nosed bats exhibit a wide variety of feeding habits and much of their morphological and behavioral diversity reflects adaptations for exploiting different kinds of food. The ancestral feeding mode in this family is undoubtedly insectivory, but blood-feeding was an early offshoot and nectarivory and frugivory are widespread in the family.

Foraging styles range from species that have small home ranges (many insectivorous or carnivorous phyllostomines and many frugivorous carolliinines and stenodermatines) to species with very large foraging ranges (the arid-zone glossophagine, Leptonycteris curasoae). Most phyllostomids commute only 0.6–1.2 mi (1–2 km) from their day roosts to feed, but individuals of L. curasoae sometimes commute up to 18.6 mi (30 km). Solitary foraging predominates in the family, but

group foraging also occurs. Mother-young pairs forage together in the common vampire (Desmodus rotundus); female harem-mates forage together in Phyllostomus hastatus; and adults (and possibly young) of L. curasoae and Phyllostomus discolor form small foraging groups when visiting flowers.

Fruit-eating phyllostomids typically harvest one fruit at a time and carry it to a night roost located 65.6–656 ft (20–200m) from the fruiting plant to eat. As a result, substantial piles of fruit remains and seeds can accumulate under these roosts. When eating fruit, phyllostomids use two different feeding methods. Glossophagines, carolliinines, and species of Sturnira are rapid feeders; they consume the pulp and seeds of a fruit in one to three minutes and then harvest another fruit. They tend to feed on succulent fruit produced by early successional plants (e.g., Cecropia species and Muntingia calabura) and understory shrubs (e.g., species of Piper, Solanum, and Vismia). In contrast, stenodermatines are slow feeders and carefully chew fruits into pellets while swallowing fruit juice and a small amount of pulp and seeds. They specialize on fig fruits, which contain high amounts of fiber, and other fruits produced by forest canopy trees.

Food sharing is known to occur in the carnivorous bat (Vampyrum spectrum); it also occurs in the common vampire. Adult female vampires share blood meals with their young and with unrelated adult females in their roost. Blood-sharing among adults is thought to prevent individuals that have been unsuccessful in finding food on a particular night from starving. This form of food-sharing is one of the few examples of reciprocal altruism known to occur in non-primate mammals.

The feeding and foraging behavior of plant-visiting phyllostomids bats has considerable ecological and economic importance. Flower-visiting species, for example, are known to pollinate nearly 1,000 species of Neotropical plants, including trees, shrubs, vines, bromeliads, and arid-zone succulents such as columnar cacti and agaves. Economically important trees that are pollinated by these bats include balsa, kapok, and calabash, as well as agaves. Likewise, fruit-eating species disperse the seeds of hundreds of species of trees and shrubs. Important tree families containing bat-dispersed species include Anacardiaceae (cashew), Saptoaceae (chicle), Moraceae (figs), and Arecaceae (palms). Important understory plant families that are bat-dispersed include Piperaceae (pepper) and Solanaceae (nightshade). Bats such as Carollia perspicillata, Sturnira lilium, and Artibeus jamaicensis are very important for dispersing seeds to disturbed habitats where secondary plant succession and forest regeneration can begin.

Reproductive biology

In part, roosting group size reflects the mating systems of American leaf-nosed bats. At one extreme is the carnivore Vampyrum spectrum, which roosts in small family groups consisting of a pair of adults and up to three of their recent offspring. This species has a monogamous mating system, the only known example so far in this family, and adults share the prey they capture with each other and their offspring. It would not be surprising to learn that other carnivorous species (e.g., Chrotopterus auritus) are also monogamous. At the other extreme are colonially roosting species with harem polygynous mating systems, the most common mating system in this family. In these systems, single males aggressively defend groups of females against the intrusions of other males. Only a fraction (about 20% in Carollia perspicillata) of adult males possesses a harem at any given time; the majority of adult males are bachelors. Harems either form seasonally or remain stable in their female composition year-round. The most stable harems occur in the greater spear-nosed bat (Phyllostomus hastatus), in which groups of unrelated females may spend their entire adult lives together. More ephemeral groups of females occur in two common fruit-eating species, Carollia perspicillata and Artibeus jamaicensis. Genetic studies indicate that harem males father most, but not all, of the babies born in their harem.

Another variation on the theme of polygyny in phyllostomids is the multiple-male/multiple-female group. An example of this mating system occurs in the common vampire, Desmodus rotundus. In this species, stable groups of eight to 12 adult females live together in roosts with several adult males, which form a dominance hierarchy regarding mating rights. When roosting in hollow trees, males fight with each other for access to the top of the roost, where most matings take place. In the phyllostomine (Macrotus californicus), seasonal aggregations of adult males and females form in which males defend preferred roosting sites against other males. They attract females for mating with wing flapping and vocalization displays.

Like all bats, American leaf-nosed bats are low-fecundity animals. Females almost always give birth to a single young once or twice a year. Insectivorous or carnivorous species and vampires tend to be monestrous and undergo a single pregnancy each year. Most plant-visiting species, in contrast, are polyestrous and undergo two pregnancies a year. In Central America, one birth occurs in the late dry season (March–April) and another occurs in the middle of the wet

season (July–August) in these species. Gestation periods are relatively long and last about four months in carolliinines and many stenodermatines. It lasts about seven months in the common vampire. In at least two species, Macrotus californicus and Artibeus jamaicensis, gestation is prolonged as a result of delayed embryonic development.

The timing of reproduction in the glossophagine (Leptonycteris curasoae) shows interesting intraspecific variation. Its general pattern is monestry, but the timing of births varies geographically. In northern South America and northern Mexico, births occur in May after a five-to-six-month gestation period. In southern Mexico, births occur in December after a similar gestation period. May births coincide with the flowering seasons of columnar cacti in the Sonoran Desert of northern Mexico and in the arid regions of northern South America. December births in southern Mexico coincide with the flowering seasons of tropical dry forest trees and shrubs. In Mexico, most of the mating activity that produces these babies occurs in caves located in south central Mexico, far from the maternity sites. After mating, females migrate north (in the spring) or south (in the fall) to form maternity colonies. Genetic studies indicate that populations belonging to these reproductive demes currently undergo substantial gene flow. Different reproductive schedules have not resulted in genetic isolation in this species in Mexico.

With a few notable exceptions, parental care is not extensive in this family. Most young phyllostomids are weaned about six weeks after birth, and few young have any further contact with their mothers (or fathers). Extended parent-offspring contact occurs in at least two species, the spectral (Vampyrum spectrum) and the common vampire (Desmodus rotundus). In the former species, young bats remain with their parents long after weaning and are provisioned with vertebrate prey while they are learning to hunt for themselves. Young vampires remain with their mothers for up to a year after birth. Their mothers sometimes feed them regurgitated blood after they reach three months of age, and they continue to forage with their mothers until they are one year old.

Conservation status

In terms of relative abundance, American leaf-nosed bats range from rare to very common. Rarity or commonness in these bats is mostly associated with their food habits or trophic position. As expected, carnivores such as Vampyrum spectrum are rare. Insectivorous phyllostomines in general are also far less common in any habitat than frugivorous carolliinines or stenodermatines. Flower visitors tend to be substantially less common than fruit-eaters. The local abundance of common vampires varies tremendously and is correlated with the availability of domestic animals. In primary forest, vampires are uncommon, but they can be very common in disturbed habitats whenever they have regular access to livestock.

In 2001, the IUCN/SSC Chiroptera Specialist Group listed four species of phyllostomids as Endangered and 25 species as Vulnerable. The endangered species included Phyllonycteris aphylla on Jamaica, Chiroderma improvisum on Guadeloupe and Montserrat, and Sturnira thomasi on Guadeloupe, as well as Leptonycteris nivalis in Mexico. The vulnerable species included seven phyllostomines, five lonchophyllines, four glossophagines, and nine stenodermatines. Only two of these species, Ariteus flavescens and Stenoderma rufum, occur on islands.

Phyllostomid bats, and Latin American bats in general, suffer from the "vampire problem," meaning most Latin Americans consider all bats to be vampiros that should be destroyed. Millions of cave-dwelling bats in Mexico alone have been killed in recent decades as a result of misguided vampire-control programs. Until enlightened vampire-control methods become widespread in Latin America, all colonially roosting bats are vulnerable to local destruction. Bat Conservation International, located in Austin, Texas, United States, has made a major effort to disseminate information about the selective control of vampires in areas where they pose an economic threat in Latin America.

Significance to humans

Because it sometimes transmits rabies to livestock, the common vampire is the most notorious phyllostomid bat. It has been estimated that 100,000 or more cattle die annually, at a cost of $40 million, as a result of vampire-transmitted rabies in Latin America. On the other hand, other phyllostomid bats can have an important positive impact on humans through their pollination and seed dispersal activities. Many of the market fruits in both the New and Old World tropics are bat-dispersed species. Sisal and tequila plants originally relied on phyllostomids bats for their pollination. Forest regeneration resulting from seed dispersal and enhanced fruit and seed set resulting from pollination are two of the beneficial ecosystem services provided by phyllostomid bats.

Species accounts

List of Species

California leaf-nosed bat

Macrotus californicus

subfamily

Phyllostominae

taxonomy

Macrotus californicus Baird, 1858, California, United States.

other common names

None known.

physical characteristics

Head and body length 2.1–2.5 in (53–64 mm); tail 1.4–1.6 in (35–41 mm); forearm 1.9–2.1 in (48–54 mm); weight 0.4–0.7 oz (12–20 g); upper body is brown or gray, lower body is brown or buff with a silvery wash.

distribution

Southwestern United States and northwestern Mexico, including Baja California.

habitat

Arid subtropical lowlands; roosts in caves, mines, and abandoned buildings.

behavior

A relatively sedentary, nonmigratory species that is active year-round at its northern distributional limit, where it roosts in geothermally heated mines in the winter; forms roosts of a few hundred to 1,000 or more individuals; occurs in many small, scattered colonies throughout its range.

feeding ecology and diet

Feeds on insects, including relatively large orthopterans and moths, which it captures in flight and by gleaning from vegetation and the ground; begins to forage well after sunset.

reproductive biology

This species is monestrous and probably polygynous. Males and females form colonies in which mating takes place in September and October; males attract females with wing-flapping and vocal displays. Embryonic development is slow during the winter and faster in spring; gestation period is eight months. Females form maternity colonies, and babies are born in June.

conservation status

Considered Vulnerable in the United States by the IUCN because of its small roost sizes.

significance to humans

None known.

Fringe-lipped bat

Trachops cirrhosus

subfamily

Phyllostominae

taxonomy

Trachops cirrhosus (Spix, 1823), Pernambuco, Brazil.

other common names

None known.

physical characteristics

Head and body length 2.6–3.5 in (65–88 mm); tail 0.4–0.8 in (10–20 mm); forearm 2.2–2.6 in (57–65 mm); weight 0.9–1.3 oz (24–36 g); upper body is dark reddish brown or cinnamon brown, lower body is dull brown washed with gray.

distribution

Southern Mexico to Bolivia and southern Brazil, and Trinidad.

habitat

Humid tropical lowlands; roosts in caves, hollow trees, culverts, and abandoned buildings.

behavior

Relatively sedentary and nonmigratory. Roosts in small, mixedsex colonies.

feeding ecology and diet

Diet includes insects, frogs, and lizards. Flies low to the ground in foraging. Preys on singing male frogs that it captures on the wing; can discriminate between palatable and unpalatable frogs on the basis of vocalizations.

reproductive biology

Monestrous and polygynous, with babies being born in May or June. Mating system is unknown.

conservation status

Not threatened. Widespread and locally common to uncommon; vulnerable to cave disturbance and habitat fragmentation.

significance to humans

None known.

Greater spear-nosed bat

Phyllostomus hastatus

subfamily

Phyllostominae

taxonomy

Phyllostomus hastatus (Pallas, 1767), Suriname.

other common names

None known.

physical characteristics

Head and body length 4.1–4.9 in (103–124 mm); tail 0.4–1.1 in (10–29 mm); forearm 3.1–3.7 in (80–93 mm); weight 2.8–3.9 oz (78–110 g); upper body is dark brown or reddish brown, lower body is somewhat paler.

distribution

Honduras to Paraguay and southeastern Brazil, and Trinidad.

habitat

Lowland tropical forests; roosts in caves, hollow trees, culverts, and abandoned buildings.

behavior

Relatively sedentary and nonmigratory; roosts in colonies of dozens to thousands of individuals. Females communicate with harem-mates by giving audible screeching calls.

feeding ecology and diet

Omnivorous and eats insects (beetles), small vertebrates (reptiles, mammals), nectar, and pollen (especially Ochroma), and fruit (especially Cecropia). Individuals forage within 6.2 mi (10 km) of their roosts; female harem-mates may forage in groups but not with their harem males. Different groups of females within a roost forage in different areas.

reproductive biology

Monestrous and polygynous, with mating and birth being quite synchronous within a roost. Births occur in April or May. Mating system is harem-polygynous with single harem males defending groups of up to about 20 females. Female harem composition is very stable with most individuals remaining together their entire lives. Recently weaned females form new social groups and do not join groups of older females. Harem male turnover rates are relatively high.

conservation status

Not threatened. Widespread and locally common to uncommon; vulnerable to cave disturbance and habitat fragmentation.

significance to humans

Important pollinator and seed-disperser of certain tropical trees.

Spectral bat

Vampyrum spectrum

subfamily

Phyllostominae

taxonomy

Vampyrum spectrum (Linnaeus, 1758), Suriname.

other common names

None known.

physical characteristics

Head and body length 5.3–6.0 in (135–152 mm); forearm 3.9–4.3 in (98–110 mm); weight 5.2–6.8 oz (145–190 g); upper body is reddish brown, lower body is slightly paler.

distribution

Southern Mexico to Peru and central Brazil, and Trinidad.

habitat

Lowland tropical forests; roosts in hollow trees.

behavior

Sedentary and nonmigratory; roosts in single family groups.

feeding ecology and diet

Carnivorous species that feeds on a wide range of vertebrates, including lizards, birds, and mammals (rodents, other bats). Adults feed within several hundred feet (meters) of their roost

and bring prey back to the day roost to share with roost-mates. In one roost studied in detail, bats fed mostly on birds ranging from 0.7–5.3 oz (20–150 g) in weight; major prey included one species of parakeet and the groove-billed ani, two species that sleep in groups at night.

reproductive biology

Monestrous and monogamous; young bats remain in their natal roost long after they are weaned.

conservation status

Not threatened, though uncommon throughout its range. Roost trees and foraging habitat threatened by forest destruction.

significance to humans

None known.

Vampire bat

Desmodus rotundus

subfamily

Desmodontinae

taxonomy

Desmodus rotundus (E. Geoffroy, 1810), Asuncion, Paraguay.

other common names

None known.

physical characteristics

Head and body length 2.7–3.7 in (68–93 mm); forearm 2.1–2.6 in (53–65 mm); weight 0.7–1.5 oz (20–43 g); upper body is

dark gray-brown, lower body is paler and sometimes with a buffy wash. Nose leaf is reduced in size.

distribution

Northern Mexico to central Chile, Argentina, and Uruguay, and Trinidad.

habitat

A wide variety of tropical and subtropical habitats but most common where livestock densities are high; not common in primary forest. Roosts in caves, hollow trees, mines, and abandoned buildings.

behavior

A sedentary and nonmigratory bat that lives in roosts containing a few dozen up to about 2,000 individuals. Highly social animals that often groom each other, a behavior that is uncommon in most other bats.

feeding ecology and diet

Morphologically, behaviorally, and physiologically specialized for feeding on vertebrate blood (mostly mammals but also birds). Flies only during the darkest part of the night and avoids flying when the moon is bright. Has long, narrow wings for flying quickly from roost to feeding areas up to 12.4 mi (20 km) away; often flies near the ground. Once it has located a sleeping mammal, it often approaches it on the ground. Its elongated thumbs serve as front feet and make this species one of the most agile bats on the ground. Makes a bite with its sharp upper incisors on the ears, neck, anus, or ankles of its victim. Laps blood from free-flowing wound and ingests about 0.7 fl oz (20 ml) of liquid. Its stomach and kidneys are adapted for quickly removing excess water (ballast) from its blood meal. Vampires will not survive if they miss more than one night of feeding. Adult females sometimes share a blood meal with other adult roost-mates that have not fed successfully.

reproductive biology

Monestrous, and single babies are born after a gestation period of about seven months. The mating system involves polygyny, but males do not defend harems. Instead, they compete for locations in roosts (e.g., the highest points inside hollow trees), which give them access to the largest number of females. Females live in stable groups of up to about 20 individuals. When they disperse, young females sometimes join mixed-age groups of females; young males disperse from their natal roosts. Young bats remain with their mothers for some time after they are weaned and sometimes share a blood meal with them. They also feed from the same wounds with their mothers.

conservation status

Not threatened. Can be too common in areas of high livestock density. In this situation, other species of bats can suffer for two reasons: vampires are aggressive and can supplant other bats from roosts and unselective vampire-control programs often involve the destruction of bat roosts; many kinds of bats die when this happens. Effective methods for controlling vampires involve applying anticoagulants directly on the fur of netted bats or injecting anticoagulants into livestock. Other vampires will ingest the anticoagulant whenever they groom each other or when they receive a blood meal. Selective control of vampires is possible because of its high degree of sociality.

significance to humans

These bats cause substantial (tens of millions of dollars annually) economic damage to livestock by transmitting rabies. Humans sometimes die from vampire-transmitted rabies.

Antillean fruit-eating bat

Brachyphylla cavernarum

subfamily

Brachyphyllinae

taxonomy

Brachyphylla cavernarum Gray, 1934, St. Vincent.

other common names

None known.

physical characteristics

Head and body length 2.6–4.6 in (65–118 mm); tail vestigial; forearm 2–2.7 in (51–69 mm); weight 1.4–1.8 oz (40–50 g); upper body is ivory-yellow with black-tipped hairs, lower body is brown. Nose-leaf reduced in size.

distribution

Puerto Rico, Virgin Islands, and the Lesser Antilles south to St. Vincent and Barbados.

habitat

Tropical forest. Roosts primarily in caves but also in buildings.

behavior

A colonial bat that lives in colonies of a few thousand individuals. Bats are strong and aggressive among themselves and other species.

feeding ecology and diet

Diet is generally broad and includes insects as well as fruit, nectar, and pollen. Like most phyllostomids, probably takes fruit to a night roost some distance from the fruiting plant to eat.

reproductive biology

Monestrous; babies born in late May or early June. Nothing is known about mating system, but most likely is polygynous.

conservation status

Not threatened. Common throughout its range but vulnerable to habitat and roost destruction.

significance to humans

None known.

Buffy flower bat

Erophylla sezekorni

subfamily

Phyllonycterinae

taxonomy

Erophylla sezekorni (Gundlach, 1860), Cuba.

other common names

None known.

physical characteristics

Head and body length 2.6–3.0 in (65–75 mm); tail 0.5–0.7 in (12–17 mm); forearm 1.8–2.2 in (45–55 mm); weight 0.6 oz (16–18 g); upper body is yellowish brown or buffy, lower body is slightly paler. Nose leaf larger than that of Brachyphylla, but still small.

distribution

Bahamas, Greater Antilles, and Grand Cayman.

habitat

Subtropical and tropical forests, including pine woodlands. Roosts in caves.

behavior

Likely to be a relatively mobile bat that sometimes appears briefly on the coast of south Florida near Miami. In the northern Bahamas, females may move between islands seasonally to form maternity roosts. Roosts contain a few hundred to a few thousand individuals.

feeding ecology and diet

Probably omnivorous, although its somewhat elongated muzzle suggests that it regularly visits flowers. Feces sometimes contain insects and seeds (Piper on Puerto Rico; Ficus, Solanum, and Tetrazygia on Grand Bahama).

reproductive biology

Monestrous; babies born in late May to early June. Mating system likely involves harem-polygyny. Single males seen with groups of females in solution holes in cave ceilings. Females appear to form maternity colonies to rear their young. One such colony on Grand Bahama contained adults and young that were well spaced out from each other on a cave wall.

conservation status

Though not threatened, habitat loss is a long-term threat. Has managed to avoid extinction on islands that have lost other species of bats owing to sea level rise and flooding of hot caves.

significance to humans

None known.

Pallas's long-tongued bat

Glossophaga soricina

subfamily

Glossophaginae

taxonomy

Glossophaga soricina (Pallas, 1766), Suriname.

other common names

None known.

physical characteristics

Head and body length 1.8–2.3 in (45–59 mm); tail 0.2–0.4 in (5–10 mm); forearm 1.3–1.5 in (33–38 mm); weight 0.3–0.4 oz (7–12 g); upper body is dark brown to reddish brown, lower body is slightly paler.

distribution

Northern Mexico to Paraguay and northern Argentina, Trinidad, Grenada, and Jamaica.

habitat

Widespread and common in many tropical lowland habitats; more common in dry forests than in wet forests where it is replaced by Glossophaga commissarisi. Roosts in wide variety of sites, including caves, hollow trees, mines, culverts, and abandoned houses.

behavior

A colonial species that lives in colonies of a few hundred to a few thousand individuals. Sedentary and nonmigratory.

feeding ecology and diet

Despite having an elongated snout and a long tongue, this bat is an omnivore. Its primary food is nectar and pollen from many species of trees, shrubs, and epiphytes, but it switches to eating fruit (of early successional trees and shrubs) when flower availability is low and eats many insects. Individuals forage solitarily, probably relatively close to their day roosts. Flies for about four hours and for about 28 mi (45 km) to feed each night.

reproductive biology

This is a polyestrous, seasonal breeder. Females undergo two pregnancies a year with births occurring in December–February and April–June in Costa Rica. The mating system has not been described but undoubtedly involves polygyny, though perhaps not harem polygyny. Females form maternity colonies to produce their young.

conservation status

Given its generally high abundance and tolerance of disturbed habitats, this species is not currently threatened with extinction.

significance to humans

It is an important pollinator and seed disperser of many tropical plants.

Geoffroy's tailless bat

Anoura geoffroyi

subfamily

Glossophaginae

taxonomy

Anoura geoffroyi Gray, 1838, Rio de Janeiro, Brazil.

other common names

None known.

physical characteristics

Head and body length 2.3–2.9 in (58–73 mm); forearm 1.6–1.8 in (40–45 mm); weight 0.5–0.6 oz (13–18 g); upper body dull brown with a silvery wash, lower body is gray-brown.

distribution

Mexico to southeastern Brazil and northwestern Argentina, and Trinidad.

habitat

Relatively common in tropical forests, especially at mid-montane elevations; occurs from 820–8,200 ft (250–2,500 m) above sea level. Roosts in caves and tunnels.

behavior

Forms relatively small colonies of 100 individuals or fewer. Likely to undergo seasonal altitudinal migrations.

feeding ecology and diet

Although it frequently visits flowers of trees, shrubs, and epiphytes, it routinely eats many insects and occasionally eats fruit. Often visits more than one species of flower during a foraging bout. Efficiently digests pollen and prefers sucrose nectar

to equicaloric solutions of fructose and glucose in lab trials. A related species (A. caudifer) flies for about five hours and for about 43 mi (70 km) while foraging each night.

reproductive biology

Monestrous; babies are born late in the year, during or just before the dry season, on Trinidad. Laboratory studies show that the seasonal male testicular cycle follows an endogenous rhythm that is not regulated by photoperiod. Mating system unknown but likely involves some form of polygyny since bats roost in small clusters in their roosts. Females form maternity roosts separate from males prior to giving birth. Young grow rapidly and are weaned by five to six weeks of age.

conservation status

Vulnerable to roost disturbance and habitat destruction, but its numbers are currently high in many locations and it is not considered threatened.

significance to humans

Pollinates tropical plants.

Southern long-nosed bat

Leptonycteris curasoae

subfamily

Glossophaginae

taxonomy

Leptonycteris curasoae Miller, 1900, Curaçao.

other common names

None known.

physical characteristics

Head and body length 2.6–3.4 in (67–86 mm); forearm 2.1–2.2 in (53–57 mm); weight 0.7–1.0 oz (20–27 g); upper body reddish brown, lower body cinnamon. Moderately elongated snout and small nose leaf.

distribution

Southern Arizona and New Mexico to El Salvador, Aruba, Bonaire, Curaçao, and adjacent Colombia and Venezuela.

habitat

Relatively common in semiarid and arid habitats in Mexico and the southwestern United States; more common in lowlands than in montane drylands where it is replaced by its sister species, L. nivalis. Roosts in caves and mines; is a hot cave bat.

behavior

A very colonial bat forming colonies containing tens to hundreds of thousands of individuals. Not highly social (no allogrooming or food-sharing) in day roosts. Many populations are highly migratory; latitudinal and altitudinal migrations are known.

feeding ecology and diet

Highly specialized flower visitor that subsists on nectar and pollen during the maternity season; also eats fruit but rarely insects. Feeds heavily on flowers of columnar cacti and paniculate agaves, when available. Also visits flowers of tropical trees and shrubs. Wide-ranging forager that spends about five hours in flight and flies about 62 mi (100 km) each night; sometimes commutes 12.4–18.6 mi (20–30 km) from day roosts to feeding areas. Although it leaves its day roost at sunset, it visits flowers mostly between midnight and 2 a.m. Sometimes forages in small groups of two to four when visiting cactus and agave

flowers. Juvenile bats likely forage with their mothers after they are weaned.

reproductive biology

Monestrous, but timing of mating and births varies geographically. In Mexico, births occur in mid-May in the Sonoran Desert and in December in tropical dry forest in Chiapas. Both sexes gather seasonally at specific mating caves. Mating system unknown but likely involves promiscuous mating in both males and females. Gestation is relatively long and lasts five to six months; females migrate while pregnant.

conservation status

Listed as Vulnerable by the IUCN; declared federally Endangered in the United States in 1988; received official protection in Mexico in 1994. Current numbers and genetic evidence, however, indicate that its population sizes are large. Because of its highly colonial roosting behavior, it is very vulnerable to roost disturbance. Protection is also needed for the habitat and food plants (columnar cacti, agaves) that support long-distance migrations.

significance to humans

Important pollinators of columnar cacti and paniculate agaves, two keystone plants of arid habitats in Mexico, southwestern United States, and northern South America.

Long-snouted bat

Platalina genovensium

subfamily

Lonchophyllinae

taxonomy

Platalina genovensium Thomas, 1928, Peru.

other common names

None known.

physical characteristics

Head and body length 2.8 in (70–72 mm); tail 0.3 in (8–9 mm); forearm 1.7–1.9 in (44–47 mm); weight 0.6–0.8 oz (16–23 g); upper and lower body color pale brown. Very long snout and small nose-leaf.

distribution

Peru.

habitat

Arid regions from sea level to about 8,200 ft (2,500 m) in the Andes; mostly a montane species. Roosts in caves and mines.

behavior

Roosts in small groups of 20 individuals or less. Nowhere is it common, and it is poorly represented in museum collections. Undoubtedly undergoes latitudinal and altitudinal migrations but these have not yet been documented.

feeding ecology and diet

Carbon stable isotope data suggest that this bat is a specialized feeder at flowers of columnar cacti. They abandon roosts whenever cactus flower supplies in the area decrease.

reproductive biology

Probably monestrous, but this topic is unstudied, as is the form of its mating system. Most likely polygynous. Pregnant females have been captured in September.

conservation status

Listed as Vulnerable by the IUCN, this species clearly warrants concern because of its low numbers and mobile lifestyle.

significance to humans

None known.

Seba's short-tailed bat

Carollia perspicillata

subfamily

Carolliinae

taxonomy

Carollia perspicillata (Linnaeus, 1758), Suriname.

other common names

None known.

physical characteristics

Head and body length 1.9–2.8 in (48–70 mm); tail 0.3–0.6 in (8–16 mm); forearm 1.6–1.8 in (41–45 mm); weight 0.5–0.9 oz (15–25 g); upper body dark brown with silvery wash, lower body lighter brown.

distribution

Southern Mexico to Paraguay and southern Brazil, Trinidad and Tobago, and Grenada.

habitat

Tropical forests of all kinds, mostly in the lowlands. Roosts in many sites, including caves, hollow trees, mines, culverts, and abandoned houses.

behavior

One of the most common bats in Latin America. Roosts contain dozens to a few thousand individuals. A relatively sedentary bat, but in western Costa Rica, females make seasonal altitudinal migrations.

feeding ecology and diet

Mostly a fruit-eater but also visits flowers opportunistically and eats insects. Individuals forage within about 1.2 mi (2 km) of their day roost. Harvests one fruit at a time and eats it in a sheltered night roost. Feeds selectively on fruits of understory shrubs (especially Piper and Vismia) and early successional trees.

reproductive biology

Polyestrous; females give birth to a single young twice a year (March–April and July–August in Central America); gestation period is about four months. Degree of synchrony of births is relatively low within populations. Mating system involves harem-polygyny with single males defending groups of up to about 20 females. Most adult males in a roost are bachelors. Sexes usually do not segregate during the maternity period.

conservation status

Not threatened. Thrives in disturbed habitats but is vulnerable to roost destruction.

significance to humans

Important seed disperser that helps to promote tropical forest regeneration resulting from natural or human disturbances.

Dwarf little fruit bat

Rhinophylla pumilio

subfamily

Carolliinae

taxonomy

Rhinophylla pumilio Peters, 1865, Bahia, Brazil.

other common names

None known.

physical characteristics

Head and body length 1.7–1.9 in (43–48 mm); forearm 1.1–1.5 in (29–37mm); weight 0.3–0.4 oz (8–10 g); upper body grayish brown, lower body paler brown.

distribution

Colombia, Ecuador, Peru, and Bolivia to the Guianas and eastern Brazil.

habitat

Lowland tropical forests. Roosts in abandoned tents made by stenodermatine bats. Much less common than species of Carollia when they occur together.

behavior

A sedentary bat that lives in small groups (fewer than 10) in forest understory tents formed from the leaves of a variety of plants (Heliconia, Philodendron, and palms). Group-foraging ranges are small (24.7–37 acres; 10–15 ha). Males may defend these areas from the intrusions of other males.

feeding ecology and diet

Mostly frugivorous but, unlike species of Carollia, does not feed on fruits of Piper shrubs; instead eats fruits of epiphytes (Macgravia and Philodendron).

reproductive biology

Probably polyestrous, but not yet studied in detail. Mating system likely involves harem-polygyny with a single adult male and two to three females and their young living in stable social units in tents. These groups remain together when they change tent locations.

conservation status

Not threatened, though, because of their specialized roosting behavior, they are vulnerable to habitat destruction.

significance to humans

None known.

Little yellow-shouldered bat

Sturnira lilium

subfamily

Stenodermatinae

taxonomy

Sturnira lilium (E. Geoffroy, 1810), Asuncion, Paraguay.

other common names

None known.

physical characteristics

Head and body length 2.1–2.6 in (54–65 mm); forearm 1.5–1.7 in (37–42 mm); weight 0.5–0.6 oz (13–18 g); upper body buff orange with a brown wash, lower body paler; males have conspicuous tufts of stiff yellow or red hairs on the shoulders.

distribution

Northwestern and northeastern Mexico to northern Argentina, southern Lesser Antilles, and Jamaica.

habitat

Most common in lowland forests, especially disturbed sites.

behavior

A relatively sedentary bat that roosts in small, inconspicuous colonies in hollow trees, clusters of vines, and palm fronds. Can be a very common bat, especially in habitats dominated by Piper and Solanum shrubs and treelets. Undergoes seasonal altitudinal migrations in southwestern Mexico and probably elsewhere.

feeding ecology and diet

Mostly frugivorous but also occasionally visits flowers. Morphologically adapted for feeding on relatively hard fruits. Selectively feeds on fruits of Solanum and Piper shrubs and treelets.

reproductive biology

Polyestrous with females giving birth to a single baby twice a year. In Costa Rica, most births occur in May and December. Mating system of this species is unknown, but, to judge from other fruit-eating phyllostomids, likely involves some form of polygyny.

conservation status

Not currently threatened, but habitat destruction and loss of hollow roost trees are two major threats.

significance to humans

Has an important role in forest regeneration through its seed dispersal activities.

Tent-making bat

Uroderma bilobatum

subfamily

Stenodermatinae

taxonomy

Uroderma bilobatum Peters, 1866, São Paulo, Brazil.

other common names

None known.

physical characteristics

Head and body length 2.3–2.7 in (59–69 mm); forearm 1.6–1.7 in (40–44 mm); weight 0.5–0.7 oz (13–20 g); upper and lower body pale gray, two bright white stripes on top of head, a single mid-dorsal white stripe.

distribution

Southern Mexico to Bolivia and southeastern Brazil; Trinidad.

habitat

Wide variety of lowland tropical forests containing palms that it uses for tent roosts.

behavior

Roosts solitarily or in small groups in leaf tents that it fashions from palm or banana-like leaves in the forest understory. Often uses palmate-leafed palms for tent sites. Occupancy of a particular tent can last as long as two months.

feeding ecology and diet

Mostly frugivorous and, like many stenodermatines, feeds heavily on fig fruits. On Barro Colorado Island, Panama, feeds mostly on small figs. Details of its foraging behavior are unknown, but this species sometimes feeds in swarms at fruiting fig trees.

reproductive biology

Seasonally polyestrous with females producing a single baby twice a year. Birth peaks occur in February and July in Panama; degree of birth synchrony is high. Mating system is unstudied but undoubtedly involves harem-polygyny. Single adult males and a few females and their young occupy one tent.

conservation status

Not currently threatened, though because of its specialized roosting requirements, it is vulnerable to deforestation.

significance to humans

It is a disperser of fig seeds.

White bat

Ectophylla alba

subfamily

Stenodermatinae

taxonomy

Ectophylla alba H. Allen, 1892, Honduras.

other common names

None known.

physical characteristics

Head and body length 1.6–1.9 in (40–47 mm); forearm 0.9–1.2 in (23–31 mm); weight 0.1–0.3 oz (4–7 g); entire body white, ears and nose leaf edged in yellow.

distribution

Eastern Honduras to western Panama.

habitat

Moist or wet lowland tropical forests. Roosts in understory in tents.

behavior

Constructs tents about 6.5 ft (2 m) above the ground using banana-like leaves in disturbed or wet areas. Males and females use the same tent.

feeding ecology and diet

Foraging behavior unstudied. Feeds on figs and other fruit.

reproductive biology

Details unstudied, but may be monestrous. Young born in April in Costa Rica. Mating system likely involves harem polygyny.

conservation status

Listed as Lower Risk/Near Threatened. Vulnerable to habitat destruction.

significance to humans

None known.

Jamaican fruit-eating bat

Artibeus jamaicensis

subfamily

Stenodermatinae

taxonomy

Artibeus jamaicensis Leach, 1821, Jamaica.

other common names

None known.

physical characteristics

Head and body length 2.8–3.3 in (70–85 mm); forearm 2.2–2.6 in (55–67 mm); weight 1.0–1.8 oz (29–51 g); upper body gray or gray-brown, lower body paler with hairs frosted with silver. Faint facial stripes on head.

distribution

Central Mexico to Paraguay and central Brazil, Trinidad and Tobago, Greater and Lesser Antilles, and southern Bahamas.

habitat

Lives in wide variety of habitats, mostly in the lowlands.

behavior

One of the most common bats in Latin America. Often roosts in foliage, less often in caves.

feeding ecology and diet

Mostly frugivorous, but also occasionally visits flowers of canopy trees and eats protein-rich leaves. Diet is known to contain many kinds of fruits but concentrates on fruits produced by the Moraceae (the fig family). In some locations, most of diet comes from figs. Eats primarily large figs in central Panama, where bats commute 0.6–1.2 mi (1–2 km) from their day roosts to fruiting fig trees. Full moon suppresses foraging activity.

reproductive biology

Seasonally polyestrous with females producing a single baby twice a year. Birth peaks occur in March and July in Panama; births are highly synchronous. Babies born in March are conceived in August; delayed development produces an eight-month gestation period. Mating system involves harem polygyny with adult males defending groups of two to 18 females. Some large harems contain a dominant and a subordinate male, which are likely to be related. Stability of the female composition of harems is relatively low. It is unlikely that young females recruit into their natal harems.

conservation status

Not threatened. A very adaptable bat that can tolerate deforestation better than most species.

significance to humans

An extremely important disperser of the seeds of forest trees.

Wrinkle-faced bat

Centurio senex

subfamily

Stenodermatinae

taxonomy

Centurio senex Gray, 1842, Chinandega, Nicaragua.

other common names

None known.

physical characteristics

Head and body length 2.1–2.7 in (54–68 mm); forearm 1.6–1.8 in (41–45 mm); weight 0.5–0.9 oz (13–26 g); upper body yellowish brown with a white spot on shoulder, lower body paler; face grotesquely wrinkled with no nose leaf; flap of skin under chin hangs over face when bat is at rest.

distribution

Northern coastal Mexico south to Venezuela and Trinidad and Tobago.

habitat

Lowland tropical forests, including second growth, up to about 4,595 ft (1,400 m). Roosts in foliage and vine tangles.

behavior

An uncommon bat that roosts in small groups of less than a dozen individuals. Bats are inconspicuous in their roosts. They apparently change roost sites regularly; new roosts are often located just several hundred feet (meters) from old roosts.

feeding ecology and diet

Thought to eat soft fruit and use its facial wrinkles to direct fruit juices into its mouth.

reproductive biology

Details unknown but records of pregnant or lactating females (February–August in Mexico and Central America) suggest seasonal polyestry. Mating system is likely to involve harem polygyny.

conservation status

Not currently threatened, but vulnerable to habitat destruction.

significance to humans

None known.

Resources

Books

Altringham, J. D. Bats: Biology and Behaviour. Oxford: Oxford University Press, 1996.

Dobat, K. Bluten and Fledermause. Frankfurt am Main: Dr. Waldemar Kramer, 1985.

Fleming, T. H. The Short-tailed Fruit Bat. Chicago: University of Chicago Press, 1988.

Fleming, T. H., and J. Nassar. "Population Biology of the Lesser Long-nosed Bat (Leptonycteris curasoae) in Mexico and Northern South America." In Columnar Cacti and Their Mutualists, edited by T. H. Fleming and A. Valiente-Banuet. Tucson: University of Arizona Press, 2002.

Greenhall, A. M., and U. Schmidt, eds. Natural History of Vampire Bats. Boca Raton: CRC Press, 1988.

Heideman, P. D. "Environmental Regulation of Reproduction." In Reproductive Biology of Bats, edited by E. G. Crichton and P. H. Krutzsch. San Diego: Academic Press, 2000.

McCracken, G. F., and G. S. Wilkinson. "Bat Mating Systems." In Reproductive Biology of Bats, edited by E. G. Crichton and P. H. Krutzsch. San Diego: Academic Press, 2000.

Nowak, R. M. Walker's Bats of the World. Baltimore: Johns Hopkins University Press, 1994.

Wilkinson, G. S. "Information Transfer in Bats." In Ecology, Evolution and Behaviour of Bats, edited by P. A. Racey and S. M. Swift. Oxford: Clarendon Press, 1995.

Periodicals

Handley Jr., C. O., D. E. Wilson, and A. L. Gardner, eds. "Demography and Natural History of the Common Fruit Bat, Artibeus jamaicensis, on Barro Colorado Island, Panama." Smithsonian Contributions to Zoology 511 (1991).

Simmons, N. B., and R. S. Voss. "The Mammals of Paracou, French Guiana: A Neotropical Lowland Rainforest Fauna, Part 1. Bats." Bulletin of the American Museum of Natural History 237 (1998).

Wetterer, A. L., M. V. Rockman, and N. B. Simmons. "Phylogeny of Phyllostomid Bats (Mammalia: Chiroptera): Data From Diverse Morphological Systems, Sex Chromosomes, and Restriction Sites." Bulletin of the American Museum of Natural History 248 (2000).

Organizations

Bat Conservation International. P.O. Box 162603, Austin, TX 78716 USA. Phone: (512) 327-9721. Fax: (512) 327-9724. E-mail: [email protected] Web site: <http://www.batcon.org>

Theodore H. Fleming, PhD