Animal Diversity Web

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)

• Biogeographic Regions
• nearctic
  • native
• neotropical
  • native

• Other Geographic Terms
• island endemic

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)

• Habitat Regions
• temperate
• tropical
• terrestrial

• Terrestrial Biomes
• desert or dune
• chaparral
• forest
• rainforest

Systematic and Taxonomic History

Bat phylogeny is very controversial among taxonomists. Fossils of family Phyllostomidae date back to the Miocene in Colombia and the Pleistocene in North, Central and South America. This family's debated relationship dates back to 1799, when genus Phyllostomus, the first official grouping of phyllostomid bats, became recognized. By 1820, bats were grouped into separate families, including family Phyllostomata, which became family Phyllostomineae in 1824. By 1904, this grouping had been rearranged many times and became known as family Phyllostomatidae. In the history of phyllostomid bat groupings, many species have been removed or added as new research has been conducted. For instance, family Mormoopidae was emancipated from the phyllostomid grouping in 1961, although it is still considered very closely related, it is even included in superfamily Noctilionidea with Phyllostomidae. Likewise, vampire bats, Desmodontinae, received a subfamily designation in 1970, although some sources still consider it a unique family. (Kwon and Gardner, 2008; Neuweiler, 1993; Simmons, 2005; Slaughter and Walton, 1970; Vaughan, et al., 2011; Wetterer, et al., 2000)

Divisions within family Phyllostomidae are often debated. In the most recent edition of Mammal Species of the World (3rd Edition, 2005), Simmons included seven subfamilies based on morphological and chromosomal data, although she acknowledged the potential for different groupings based on further molecular research. For instance, in 2003, 11 subfamilies were suggested for family Phyllostomidae based on mitochondrial DNA evidence, although this account uses the divisions designated by Simmons 2005. Subfamily divisions are based on morphological traits such as dentition, noseleaf and skull structure, as well as dietary habits. Subfamily Phyllostominae includes 16 genera and 42 species, such as round-eared and spear-nosed bats; however, there is currently a great deal of debate regarding the appropriate number of species and genera within this subfamily. Phyllostominae is the only subfamily that retained the ancestral phyllostomid tuberculosectorial dentition. There are 3 genera and 3 species of vampire bats, all of which are within subfamily Desmodontinae. Research conducted in 2000 suggested that this subfamily may represent a basal lineage of phyllostomids, although this was not fully supported by mtDNA studies. Subfamily Glossophaginae, long-nosed and long-tongued bats, includes 13 genera and 32 species. This subfamily includes two tribes, although, the tribes are sometimes treated as unique subfamilies. Morphologically, tribe Glossophagini and Lonchophyllini can be separated based on their tongue structure. Little fruit or short-tailed bats are included in subfamily Carolliinae, with 2 genera and 9 species, although genus Rhinophylla has historically been grouped in Phyllostominae, Stenodermatinae or in its own unique family and genus Carollia has been grouped with subfamily Glossophaginae. The largest subfamily, Stenodermatinae, fruit-eating, tent-making or yellow-shouldered bats, includes 18 genera and 67 species. Three groups have been recognized within this subfamily, although only 2 tribes are currently valid. Likewise, subfamily Brachyphyllinae, Antillean fruit-eating bats, includes 1 genus and 2 species and subfamily Phyllonycterinae includes 2 genera and 5 species. (Feldhamer, et al., 2007; Gardner, 2008a; Griffiths and Gardner, 2008a; Kwon and Gardner, 2008; McLellan and Koopman, 2008; Neuweiler, 1993; Simmons, 2005; Slaughter and Walton, 1970; Vaughan, et al., 2011; Williams and Genoways, 2008)

• Synonyms

  • Phyllostomatidae
  • Phyllostomus
  • Phyllostomata
  • Phyllostomineae

• Synapomorphies

  • externally fused uterus and oviducts
  • toes have three phalanges

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, 2008b; 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, 2008a; 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)

• Other Physical Features
• endothermic
• bilateral symmetry

• 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)

• Mating System
• monogamous
• polygynous
• cooperative breeder

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)

• Key Reproductive Features
• iteroparous
• seasonal breeding
• year-round breeding
• gonochoric/gonochoristic/dioecious (sexes separate)
• sexual
• viviparous
• sperm-storing
• delayed fertilization
• delayed implantation
• embryonic diapause

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)

• Key Behaviors
• flies
• nocturnal
• crepuscular
• parasite
• motile
• migratory
• sedentary
• aestivation
• daily torpor
• territorial
• social
• colonial
• dominance hierarchies

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)

• Communication Channels
• visual
• tactile
• acoustic
• chemical

• Perception Channels
• visual
• infrared/heat
• tactile
• acoustic
• ultrasound
• echolocation
• chemical

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)

• Primary Diet
• carnivore
  • eats terrestrial vertebrates
  • sanguivore
  • insectivore
• herbivore
  • frugivore
  • nectarivore
• omnivore

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

• Known Predators

  • snakes Serpentes
  • falcons Falconiformes
  • skunks Mephitidae
  • raccoons Procyonidae
  • domestic cats Felis catus
  • owls Strigiformes

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

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)

• Negative Impacts
• injures humans
  • bites or stings
  • causes disease in humans
  • carries human disease
• causes or carries domestic animal disease
• household pest

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.

References

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