PhyllostomidaeNew World leaf-nosed bats

Diversity

Phyllostomidae represents one of the most morphologically diverse bat families, with at least 160 species, divided into 7 subfamilies with more than 55 genera. Commonly known as New World leaf-nosed bats, this family is adapted to a wide range of environments and diets. Members of this family are found as far south as Argentina and north to the southwestern United States, covering diverse habitats including forests, deserts and caves. This family has an immense dietary range, foraging on a variety of foods including insects, fruits, nectar, pollen, small vertebrates and blood. Phyllostomids are nocturnal and echolocate to find food and escape predation. This family generally uses short, broadband echolocation signals of low intensity to perceive their environment. Some species are solitary, while others roost in groups or colonies. ("Encyclopedia Britannica", 2009; Hill and Smith, 1992; Neuweiler, 1993; Peterson, 1964; Slaughter and Walton, 1970; Vaughan, et al., 2011)

Geographic Range

Family Phyllostomidae occupies mostly Neotropical regions of the New World. Phyllostomid bats are found in the extreme southwestern United States to northern Argentina and central Chile (approximately 30˚south latitude) and on several Caribbean islands. Their range extends to regions of the West Indies; members of subfamily Brachyphyllinae are restricted to these islands. ("Encyclopedia Britannica", 2009; Feldhamer, et al., 2007; Hill and Smith, 1992; Neuweiler, 1993; Slaughter and Walton, 1970)

Habitat

Members of family Phyllostomidae are found in many habitats including dense rainforests, deserts and caves, from sea level to areas with high elevations. This family is nocturnal and seeks shelter during the day. Daytime roost structures are quite diverse. Most phyllostomid bats are forest dwellers; however, some species, such as members of genus Leptonycteris, Mexican long-tongued bats and long-snouted bats are desert dwellers and spend much of the day in rock crevices. In the tropics, caves provide ideal day roosts as they maintain an optimal temperature and humidity. Other tropical roosts include hollow trees and crevices in aerial root tangles. Several species of fruit and nectar eating phyllostomids form shelters from large leaves, their preferred materials include the leaves of various Heliconia species and palms. For example, Honduran white bats chew through the horizontal ribs of a Heliconia leaf, causing the leaf to collapse and form a tent. These tents are short-lived and cause bats to change roost locations often. In addition, phyllostomid bats may also roost in buildings or other human-made objects. Carnivorous and frugivorous bats often transport their food to a night roost, found in or near their foraging territory, to eat in relative safety. Ideal night and day roosts share the same characteristics; they are secure from predators and maintain a stable temperature. (Feldhamer, et al., 2007; Hill and Smith, 1984; Neuweiler, 1993; Peterson, 1964; Slaughter and Walton, 1970)

Physical Description

A defining characteristic of family Phyllostomidae is the presence of a noseleaf. This structure is used to send out and interpret incoming high frequency sounds. Some species have developed incredibly large nose-leaves, while they may be completely absent in other species. When present, noseleaves arise from a fleshy plate that surrounds the nasal apertures and stands erect behind these openings. Noseleaves may be long or short, slender or broad. In vampire bats (Desmodontinae) the noseleaf is greatly reduced and modified into a complex series of folds and bumps around the ornamented narial plate, whereas members of subfamily Phyllostominae have highly developed noseleaves. Phyllostomids range dramatically in size; the smallest members, little white-shouldered bats have a forearm length of 26 mm, while the largest members, spectral bats, have forearm lengths of up to 106 mm. Their body size is closely related to food habits and modes of flight. Bats that consume larger food items are generally larger and have long, broad wings capable of lifting heavy weights, although their flight speeds are relatively slow. Insect eating species tend to be smaller and faster, for catching quickly maneuvering insects. When present, tails are enclosed in a uropatagium. Phyllostomids lack postorbital processes and their premaxillae are completely fused. Like body size, head shape and tooth structure generally reflects dietary habits. Nectar and pollen eaters have long, tubular muzzles, fewer and relatively smaller teeth, elongated jaws, and the back of their heads are rounded and low. Their ears vary tremendously in size and shape. All phyllostomids have a tragus. This family uses short, broadband, low intensity echolocation signals. Members of family Phyllostomidae have large eyes and rely somewhat on visual acuity in their nightly activities. Flight is their most common mode of locomotion; however, members of subfamily Desmodontinae have strong, elongated leg bones, which allow them to proficiently walk and jump. Vampire bats are the only phyllostomids capable of launching themselves into flight from the ground. (Gardner, 2008a; Hill and Smith, 1992; Neuweiler, 1993; Peterson, 1964; Slaughter and Walton, 1970; Williams and Genoways, 2008)

Phyllostomid subfamilies show a great deal of morphological variation based on their dietary habits. Subfamily Phyllostominae has the most ancestral appearance, including tuberculosectorial dentition. Some Phyllostominae species have a tail and all species have a noseleaf. Carnivorous Phyllostominae species generally have a larger body size, which helps them carry their prey; they also have robust canines and molars. Insectivorous species, including Phyllostominae and Stenodermatinae species are usually small and have complex, sharply crested teeth. Members of subfamily Stenodermatinae may also be frugivorous, interestingly; their robust molars are similar to those of Neotropical primates. They often have brightly colored pelage, with a well-developed noseleaf and no tail. Nectivorous species, such as Glossophaginae and Brachyphyllinae are small with long, narrow snouts. They have extremely long tongues tipped with brush-like structures to help them collect pollen and nectar. Their teeth are often small, which helps their tongue move. Members of subfamily Glossophaginae also have round ears, facial whiskers and noseleaves. Likewise, nectivorous Phyllonycterinae species are small to medium-sized with long, thin rostrums and tongues and broad wings, with a small uropatagium. Larger frugivorous species, like members of Stenodermatinae have flattened faces, with rounded mouths, and simple teeth, with sharp canines and flat molars for grabbing and crushing fruit. Members of subfamily Carolliinae are frugivorous, with generalized skulls, reduced molars and long wings. With the most divergent diet, members of Desmodontinae also have the most unusual dentition. Their teeth are morphologically specialized with two sharp upper incisors and bladelike canines to slit their prey’s skin. Their molars are simple and reduced, as they do not chew; instead they lick blood as it flows from the wound. These bats are relatively small, with a short rostrum and rudimentary noseleaf. (Allen, 2004; Findley, 1993; Gardner, 2008b; Griffiths and Gardner, 2008a; Griffiths and Gardner, 2008b; Hill and Smith, 1984; Kwon and Gardner, 2008; McLellan and Koopman, 2008; Vaughan, et al., 2011; Williams and Genoways, 2008)

  • Sexual Dimorphism
  • sexes alike
  • female larger
  • male larger
  • sexes colored or patterned differently
  • female more colorful
  • male more colorful
  • sexes shaped differently
  • ornamentation

Reproduction

Members of family Phyllostomidae have a variety of mating systems based on their social structure and roosting habits, although mating systems are only understood for about 7.5% of the family. Spectral bats are the only currently known monogamous members of this family. More commonly, members of this family are polygynous. Polygynous, harem-based mating systems have been described for common vampire bats, greater spear-nosed bats, pale spear-nosed bats and Jamaican fruit bats. Likewise, seasonal polygynous aggregations such as leks are seen in California leaf-nosed bats and Honduran white bats. (Adams and Pedersen, 2000; Altringham, 1996; Crichton and Krutzsch, 2000; Fenton, 1985; Fenton, 2001; Greenhall and Schmidt, 1988; Nowak, 1994; Slaughter and Walton, 1970)

Most phyllostomids are polyestrous and show no breeding seasonality. Males experience year round spermatogenesis, although members of genus Hipposideros store sperm until copulation. Within this family, both ovaries are generally functional in the egg release, except for subspecies of Waterhouse's leaf-nosed bats (Macrotus waterhoussi californicus). At the northern edges of the family's range in the southwestern United States, this subspecies follows a seasonal reproductive cycle, with a single estrous and ovulation during the summer. Embryonic diapauses can be seen for a period of about 4 months in this northern subspecies and in Jamaican fruit bats. Phyllostomids typically give birth to a single offspring, which weighs about 26.6% of the mother's weight on average. Gestation periods are variable, lasting 8 months in vampire and Waterhouse's leaf-nosed bats to as short as 40 days. Weaning is also variable, ranging from about 6 weeks to 8 to 10 months in vampire bats. The age of reproductive maturity is usually earlier for females, but ranges from 1 to 2 years for both sexes. (Crichton and Krutzsch, 2000; Fenton, 2001; Hill and Smith, 1984; Hill and Smith, 1992; Kleiman, et al., 2004; Neuweiler, 1993)

Parental investment is almost solely maternal; it is also energy expensive due to prolonged gestation periods for large, well-developed young. Females provide parental care by nursing and food supplementation. The pre-parturition phase also involves hormone and immunological exchange. Altricial young rely on females for thermoregulation, nursing, transporting, food provisioning and teaching. Common vampire bats nurse up to 9 to 10 months, and greater spear-nosed bats produce 15.7 ml of milk, the same volume a similar-sized rodent might produce for multiple young. One exception to sole female care is seen among monogamous spectral bats. (Adams and Pedersen, 2000; Altringham, 1996; Crichton and Krutzsch, 2000; Fenton, 1985; Fenton, 2001; Greenhall and Schmidt, 1988; Nowak, 1994; Slaughter and Walton, 1970)

  • Parental Investment
  • altricial
  • precocial
  • male parental care
  • female parental care
  • pre-fertilization
    • protecting
      • female
  • pre-hatching/birth
    • provisioning
      • female
  • pre-weaning/fledging
    • provisioning
      • male
      • female
    • protecting
      • male
      • female
  • pre-independence
    • provisioning
      • female
  • post-independence association with parents
  • extended period of juvenile learning
  • inherits maternal/paternal territory

Lifespan/Longevity

As small mammals, bats are generally long-lived for their size. If they survive past their first and most uncertain year, their average lifespan is about 7 years. Members of genus Desmodus are known to live 18 years in the wild and up to 19.5 years in captivity. (Greenhall and Schmidt, 1988; Slaughter and Walton, 1970)

Behavior

Many members of family Phyllostomidae have a low wing aspect ratio and high wing load, which reflects feeding behavior, such as hovering to feed on insects, pollen and fruit. Roosting groups range from single bats in genus Mimon and Macrophyllum, to groups of more than a thousand like genus Phyllostomus, Phyllonycteris and Desmodus. Phyllostomids may roost in drain pipes, buildings, caves and vegetation. Using broad leaves, members of genus Uroderma make tents by chewing along the leaf mid rib, causing it to fold into a tent shape, under which about 10 individuals can roost. Most phyllostomids do not exhibit torpor or reduced body temperature, but aestivation is possible for three species including Pallas's long-tongued bats, Heller's broad-nosed bats and Seba's short-tailed bats. Migration and a semi-dormant hibernation-like state is observed only in Macrotus species. Reciprocity and food sharing is demonstrated by Desmodus species as a fairly complex social interaction between mother and young, related individuals (kin selection) and unrelated individuals (altruism) in the form of blood regurgitation. Aggressive behaviors are displayed by Desmodus species including hair bristling, wing beating, lunging and vocal calls when an individual at a prey wound is approached by another. Similar displays are made by males in roost defense; the presence of bite scars suggests that males often engage in fights over roosts and female defense. A less aggressive and more communal behavior is displayed by greater spear-nosed bats, by making feeding calls to coalesce a larger foraging group on their way to feeding locations. (Altringham, 1996; Crichton and Krutzsch, 2000; Fenton, 1985; Fenton, 2001; Greenhall and Schmidt, 1988; Nowak, 1994; Slaughter and Walton, 1970)

Communication and Perception

Phyllostomids use audible, tactile, olfaction and visual perception. Tactile reception involving vibrissae and thermoperception using their noseleaf has evolved in Desmodus species for prey detection. When they perceive danger, Diaemus species can eject a fine mist with an offensive odor using two oral glands. Female California leaf-nosed bats, greater spear-nosed bats, Seba's short-tailed bats and common vampire bats use olfactory detection to find their young. These species, in addition to Leptonycteris sanborni, also identify young by vocalizations. To echolocate phyllostomids emit low-intensity, larynx-derived sounds through their nostrils, which are well-suited for detecting large or stationary objects such as flowers and fruits. Often referred to as whispering bats, most phyllostomids emit a short duration pulse of multi-harmonic fm sounds. Common vampire bats echolocate by emitting sounds through their mouth, which they keep open when flying, and can detect the sound of their preys’ respiration and movement. Likewise, insectivorous Pteronotus species can judge the distance of fruit flies using echolocation. The variation in echolocation can be attributed to prey selection. (Altringham, 1996; Fenton, 1985; Fenton, 2001; Greenhall and Schmidt, 1988; Nowak, 1994; Slaughter and Walton, 1970)

Food Habits

The seven subfamilies of Phyllostomidae are distinctive mainly due to their dietary specializations and related adaptations. Phyllostominae, the most primitive subfamily, includes carnivorous, insectivorous and frugivorous species. Carnivorous species feed on small vertebrates like birds, frogs, rodents and other bats. Subfamily Glossophaginae includes nectar and pollen feeders and Carolliinae species are frugivorous, both subfamilies are very rapid feeders, consuming food items within minutes. Glossophaginae species in particular have remarkable adaptations for harvesting nectar, and, like hummingbirds, their diurnal counterpart, they are capable of hovering flight. Frugivorous members of subfamily Stenodermatinae specialize on larger, high fiber fruits and are slow feeders, carefully chewing the fruit to digest it more efficiently, although Stenodermatinae species may also be insectivorous as well. Members of Brachyphyllinae are frugivorous and nectivorous but show different specializations. Likewise, subfamily Phyllonycterinae is nectivorous. Finally, Desmodontinae species are the only mammal species that feed exclusively on blood. They use infrared sensors to determine a suitable place to bite an animal, and then make an incision and lick the blood that flows due to an anticoagulant in their saliva. Vampire bats feed mainly on birds and livestock, but humans are sometimes bitten as well. When vampire bats fail to feed they may try to induce a food donation from a conspecific, this altruistic act can keep an adult vampire from starving. (Allen, 2004; Findley, 1993; Hill and Smith, 1984; Vaughan, et al., 2011)

Predation

There is no evidence of any predators specializing in phyllostomid bats, although many animals such as skunks, raccoons, snakes, domestic cats and some owls and raptors are known opportunistic bat predators. Bats generally avoid predation with their cryptic coloration, diurnal habits and lithe night flight. Bats are therefore most susceptible if discovered in their roosts, or when they are arriving or departing their roosts. Some predators like snakes, owls and hawks may wait near their cave entrances, attacking bats as they leave their roosts in large numbers, but individual bats are probably difficult to catch. A few species are specialized for preying on other bats, including spectral bats and big-eared woolly bats. (Allen, 2004; Findley, 1993; Hill and Smith, 1984; Tuttle, 2002)

  • Anti-predator Adaptations
  • cryptic

Ecosystem Roles

Nectivorous and frugivorous subfamilies pollinate and disperse seeds in the New World. Over 1,000 plants are pollinated by members of Phyllostomidae. Likewise, fruit-eating species disperse seeds of many plants, such as fig trees, palms, peppers, agaves and many types of columnar cacti. Some bats prefer to move the fruit before eating, apparently to avoid predators, and therefore have a larger seed dispersal territory than other dispersing animals. These bats assist secondary plant succession and forest regeneration. Insectivorous members of Phyllostomidae prey on insects that would otherwise become pests. Desmodontinae are parasitic on their host prey, usually livestock or bird species, by living on their blood. They can also carry and transfer rabies to their hosts. (Allen, 2004; LaVal and Rodriguez, 2002)

  • Ecosystem Impact
  • disperses seeds
  • pollinates
  • creates habitat
  • parasite
Species Used as Host
Mutualist Species
  • common figs (Ficus carica)
  • Agaves
  • balsa trees (Ochroma pyramidale)
  • Javan cotton trees (Ceiba pentandra)
  • calabash trees (Crescentia cujete)
Commensal/Parasitic Species

Economic Importance for Humans: Positive

Through pollination and seed dispersal alone, Phyllostomidae species greatly impact the economy. Thousands of plant species rely on phyllostomid pollination and seed dispersal for propagation, and many of those species are used by humans, such as figs, agaves, balsa, kapok, and calabash. Bats are also key insect predators, preying on species that cost farmers and foresters billions of dollars annually and are a natural alternative to pesticide use. Their feces are harvested by humans for use as a natural fertilizer. Although Desmodontinae species are sometimes considered a threat to humans because they can transmit rabies, the anticoagulant protein in their saliva is being studied in an effort to help prevent blood clots in humans. (LaVal and Rodriguez, 2002)

  • Positive Impacts
  • source of medicine or drug
  • research and education
  • produces fertilizer
  • pollinates crops
  • controls pest population

Economic Importance for Humans: Negative

Desmodontinae species are often seen as an economic threat to human interests because they can transmit rabies to livestock, birds and even humans. Even if bats do not transmit rabies, injuries can incur from their bites. Other non-vampire species of family Phyllostomidae can also become a nuisance if they begin roosting in human dwellings and need to be removed. (LaVal and Rodriguez, 2002)

Conservation Status

As of 2001, the IUCN/SSC Chiroptera Specialist Group listed four species of phyllostomids as endangered and 25 species as vulnerable. Endangered species included Jamaican flower bats, Guadeloupe big-eyed bats, Thomas's yellow-shouldered bats and greater long-nosed bats. Vulnerable species include seven Phyllostominae species, five Brachyphyllinae species, four Glossophaginae species and nine Stenodermatinae species. Phyllostomid bats are at risk because of the prejudice against “vampires” in Latin America. Many humans are intolerant of Desmodontinae species and feel all vampire bats should be destroyed. Due to a lack of education, many view all members of Phyllostomidae as vampires. Misguided vampire-control programs can eradicate millions of bats, both vampire and not, in a short amount of time, leaving all species, especially colonially roosting bats, vulnerable to extinction. While many Phyllostomidae species are not currently threatened, they are vulnerable to deforestation, habitat loss, mass eradication programs and pesticide use. Species with small geographic ranges or ecological specializations are at greatest risk. ("2008 IUCN Red List of Threatened Species", 2008)

  • IUCN Red List [Link]
    Not Evaluated

Contributors

Leanne Burns (author), University of Michigan-Ann Arbor, Vanessa Hutzley (author), University of Michigan-Ann Arbor, Zach Laubach (author), University of Michigan-Ann Arbor, Leila Siciliano Martina (editor), Animal Diversity Web Staff.

Glossary

Nearctic

living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.

World Map

Neotropical

living in the southern part of the New World. In other words, Central and South America.

World Map

acoustic

uses sound to communicate

altricial

young are born in a relatively underdeveloped state; they are unable to feed or care for themselves or locomote independently for a period of time after birth/hatching. In birds, naked and helpless after hatching.

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

carnivore

an animal that mainly eats meat

causes disease in humans

an animal which directly causes disease in humans. For example, diseases caused by infection of filarial nematodes (elephantiasis and river blindness).

causes or carries domestic animal disease

either directly causes, or indirectly transmits, a disease to a domestic animal

chaparral

Found in coastal areas between 30 and 40 degrees latitude, in areas with a Mediterranean climate. Vegetation is dominated by stands of dense, spiny shrubs with tough (hard or waxy) evergreen leaves. May be maintained by periodic fire. In South America it includes the scrub ecotone between forest and paramo.

chemical

uses smells or other chemicals to communicate

colonial

used loosely to describe any group of organisms living together or in close proximity to each other - for example nesting shorebirds that live in large colonies. More specifically refers to a group of organisms in which members act as specialized subunits (a continuous, modular society) - as in clonal organisms.

cooperative breeder

helpers provide assistance in raising young that are not their own

crepuscular

active at dawn and dusk

cryptic

having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.

delayed fertilization

a substantial delay (longer than the minimum time required for sperm to travel to the egg) takes place between copulation and fertilization, used to describe female sperm storage.

delayed implantation

in mammals, a condition in which a fertilized egg reaches the uterus but delays its implantation in the uterine lining, sometimes for several months.

desert or dunes

in deserts low (less than 30 cm per year) and unpredictable rainfall results in landscapes dominated by plants and animals adapted to aridity. Vegetation is typically sparse, though spectacular blooms may occur following rain. Deserts can be cold or warm and daily temperates typically fluctuate. In dune areas vegetation is also sparse and conditions are dry. This is because sand does not hold water well so little is available to plants. In dunes near seas and oceans this is compounded by the influence of salt in the air and soil. Salt limits the ability of plants to take up water through their roots.

dominance hierarchies

ranking system or pecking order among members of a long-term social group, where dominance status affects access to resources or mates

drug

a substance used for the diagnosis, cure, mitigation, treatment, or prevention of disease

echolocation

The process by which an animal locates itself with respect to other animals and objects by emitting sound waves and sensing the pattern of the reflected sound waves.

embryonic diapause

At about the time a female gives birth (e.g. in most kangaroo species), she also becomes receptive and mates. Embryos produced at this mating develop only as far as a hollow ball of cells (the blastocyst) and then become quiescent, entering a state of suspended animation or embryonic diapause. The hormonal signal (prolactin) which blocks further development of the blastocyst is produced in response to the sucking stimulus from the young in the pouch. When sucking decreases as the young begins to eat other food and to leave the pouch, or if the young is lost from the pouch, the quiescent blastocyst resumes development, the embryo is born, and the cycle begins again. (Macdonald 1984)

endothermic

animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.

female parental care

parental care is carried out by females

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

frugivore

an animal that mainly eats fruit

herbivore

An animal that eats mainly plants or parts of plants.

infrared/heat

(as keyword in perception channel section) This animal has a special ability to detect heat from other organisms in its environment.

insectivore

An animal that eats mainly insects or spiders.

island endemic

animals that live only on an island or set of islands.

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

male parental care

parental care is carried out by males

migratory

makes seasonal movements between breeding and wintering grounds

monogamous

Having one mate at a time.

motile

having the capacity to move from one place to another.

native range

the area in which the animal is naturally found, the region in which it is endemic.

nectarivore

an animal that mainly eats nectar from flowers

nocturnal

active during the night

omnivore

an animal that mainly eats all kinds of things, including plants and animals

parasite

an organism that obtains nutrients from other organisms in a harmful way that doesn't cause immediate death

polygynous

having more than one female as a mate at one time

rainforest

rainforests, both temperate and tropical, are dominated by trees often forming a closed canopy with little light reaching the ground. Epiphytes and climbing plants are also abundant. Precipitation is typically not limiting, but may be somewhat seasonal.

sanguivore

an animal that mainly eats blood

seasonal breeding

breeding is confined to a particular season

sedentary

remains in the same area

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

sexual ornamentation

one of the sexes (usually males) has special physical structures used in courting the other sex or fighting the same sex. For example: antlers, elongated tails, special spurs.

social

associates with others of its species; forms social groups.

sperm-storing

mature spermatozoa are stored by females following copulation. Male sperm storage also occurs, as sperm are retained in the male epididymes (in mammals) for a period that can, in some cases, extend over several weeks or more, but here we use the term to refer only to sperm storage by females.

tactile

uses touch to communicate

temperate

that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).

terrestrial

Living on the ground.

territorial

defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement

tropical

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

ultrasound

uses sound above the range of human hearing for either navigation or communication or both

visual

uses sight to communicate

viviparous

reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.

year-round breeding

breeding takes place throughout the year

young precocial

young are relatively well-developed when born

References

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Gardner, A. 2008. Subfamily Stenodermatinae P. Gervais, 1856. Pp. 300-376 in A Gardner, ed. Mammals of South America: Marsupials, Xenarthrans, Shrews, and Bats, Vol. 1. Chicago: University of Chicago Press.

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

Griffiths, T., A. Gardner. 2008. Subfamily Glossophaginae Bonaparte, 1845. Pp. 224-244 in A Gardner, ed. Mammals of South America: Marsupials, Xenarthrans, Shrews, and Bats, Vol. 1. Chicago: University of Chicago Press.

Griffiths, T., A. Gardner. 2008. Subfamily Glossophaginae Griffiths, 1982. Pp. 244-255 in A Gardner, ed. Mammals of South America: Marsupials, Xenarthrans, Shrews, and Bats, Vol. 1. Chicago: University of Chicago Press.

Hill, J., J. Smith. 1992. Bats: A Natural History. United States of America: University of Texas Press.

Hill, J., J. Smith. 1984. Bats A Natural History. London: British Museum of Natural History.

Kleiman, D., V. Geist, M. McDade. 2004. American leaf-nosed bats (Phyllostomidae). Pp. 307-318, 413-434 in Grzimek's Animal Life Encyclopedia, Vol. 13, Second Edition. Farmington Hills, Michigan: Gale.

Kwon, M., A. Gardner. 2008. Subfamily Desmodontinae J.A. Wagner, 1840. Pp. 218-224 in A Gardner, ed. Mammals of South America: Marsupials, Xenarthrans, Shrews, and Bats, Vol. 1. Chicago: University of Chicago Press.

LaVal, R., B. Rodriguez. 2002. Costa Rica Bats. Costa Rica: Editorial National Institute of Biodiversity.

McLellan, L., K. Koopman. 2008. Subfamily Carolliinae Miller, 1924. Pp. 208-218 in A Gardner, ed. Mammals of South America: Marsupials, Xenarthrans, Shrews, and Bats, Vol. 1. Chicago: University of Chicago Press.

Neuweiler, G. 1993. The Biology of Bats. New York: Oxford University Press.

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

Peterson, R. 1964. Silently, By Night. United States of America: Russell Peterson.

Simmons, N. 2005. Order Chiroptera. Pp. 312-529 in D Wilson, D Reeder, eds. Mammal Species of the World, Vol. 1, 3 Ed. Baltimore: The Johns Hopkins University Press.

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