Diversity
Hipposideridae, Old World leaf-nosed bats, is one of eighteen families that make up
the order
Chiroptera
, and consists of nine genera. The largest genus is
Hipposideros
(roundleaf bats), which consists of 76 species. The remaining genera are
Asellia
(trident leaf-nosed bats),
Anthops
(flower-faced bats),
Aselliscus
(trident bats),
Cloeotis
(Percival's trident bat),
Coelops
(tailless leaf-nosed bats),
Paracoelops
(orange leaf-nosed bat),
Rhinonicteris
(orange leaf-nosed bat), and
Triaenops
(trident bats). Hipposiderids live in tropical and subtropical regions of Africa,
south Asia, Australia, the Philippine Islands, and the Solomon Islands. They can be
found in deserts, dunes, savannas, grasslands, forests, rainforests, scrub forests
and mangroves. Most species roost in dark, enclosed spaces, but some do roost in open
areas. Hipposiderids range from 28 to 110 mm in body length, 30 to 110 mm in forearm
(wing) length, and may or may not have a tail, up to 60 mm in length. Colors range
from white to red to dark brown depending on species, geographic area, sex, and age,
and
pelage
also varies in length and texture. They may have small or large ears, and some species’
ears are interconnected along the dorsal surface of the head. The appearance of the
noseleaf is highly variable among genera. Hipposiderids show a great deal of diversity
in roosting behavior and reproductive habits and show slight differences in feeding
habits from genus to genus.
Geographic Range
Members of Hipposideridae are found throughout tropical and subtropical regions of
the Old World. These Old World leaf-nosed bats are found in Africa, southern Asia,
the Philippine Islands, the Solomon Islands, and Australia.
- Biogeographic Regions
- palearctic
- oriental
- ethiopian
- australian
Habitat
Hipposideridae inhabits tropical and subtropical habitats and roosting preferences
vary by genera. Hipposiderids have been found roosting in caves, mines, hollow trees,
buildings, and man-made underground compartments like cellars and tombs. In Africa,
Fulvus round-leaf bats
are often found in the burrows of
Hystrix
(Old World porcupines) and members of the genus
Asellia
roost in the inner walls of wells, in caves, and in man-made structures. Though
Percival's trident bats
live in forests and generally roost in trees, in Taiwan they have been discovered
in abandoned Japanese bomb shelters, also known as pillboxes.
- Habitat Regions
- tropical
- terrestrial
- Terrestrial Biomes
- desert or dune
- savanna or grassland
- forest
- rainforest
- scrub forest
Systematic and Taxonomic History
The phylogenetic relationships between Hipposiderids and their closest relatives are
not well understood. Currently, Hipposideridae is considered 1 of 18 families in the
order
Chiroptera
. Traditionally, the order
Chiroptera
was divided into two sub-orders,
Megachiroptera
and
Microchiroptera
. Recently,
Microchiroptera
was further divided into
Yangochiroptera
and
Yinochiroptera
. Hipposideridae is classified within
Yinochiroptera
. Previously, Hipposideridae was considered a sub-family of the family
Rhinolophidae
. Under this classification, what is now known as Hipposideridae was the sub-family
Hipposiderinae
, and what is known now as
Rhinolophidae
was known as the sub-family
Rhinolophinae
. Recent nuclear sequencing and morphological data suggests that hipposiderids are
more closely related to
Megachiroptera
than
Microchiroptera
. Regardless, more research is needed in order to establish a well-supported phylogenetic
and taxonomic history of this group. The earliest known hipposiderid fossils are from
the
Eocene
.
Physical Description
A defining characteristic of Hipposiderids is their elaborate noseleaf. The noseleaf
consists of fleshy protrusions on top of a U-shaped rhinarium (i.e., the wet surface
surrounding the nostrils). Hipposiderids have an erect transverse leaf within the
noseleaf as well as smaller accessory leaflets. The common name of many genera corresponds
to the shape of the noseleaf. For example,
flower-faced bats
have two circular lateral leaflets, the smaller of which is superimposed onto the
larger, resulting in a noseleaf resembling the petals of a flower. Differences in
noseleaf characteristics are commonly used to discern between genera. These 'appendages'
are thought to be related to nasal echolocation, and may help to focus and modify
echolocation signals.
Pelage
of hipposiderids varies greatly both between and within taxa. Pelage can be white,
light beige, pale yellow, dark yellow, orange, red, red-brown, light brown, dark brown,
gray, or dark gray. Some species have white patches of fur, while others have 2 different
color phases. Often, venter pelage is lighter than dorsum pelage. Pelage also differs
interspecifically by length and texture or silkiness. Hipposiderid skulls have a number
of unique features that differentiate them from other bat families. For example, they
have no post-orbital processes, the nasal portion of the premaxilla is absent, and
the premaxilla is not fused to the maxilla. They have
dilambdodont
molars , and their dental formula is I1/2 C1/1 P1–2/2–3 M3/3, giving them a total
of 28 to 30 teeth. Hipposiderids do not have a tragus, the fleshy protuberance present
at the opening of the ear in many bats. A membrane spanning the dorsal surface of
the head connects their ears, which exhibit a great deal of variation in size.
Hipposideridae shares many traits with the family
Rhinolophidae
, and some accounts consider Hipposideridae a sub-family of Rhinolophidae. Both
hipposiderids
and
rhinolophids
lack post-orbital processes and the nasal portion of the premaxilla, as well as having
a premaxilla that is not fused to the maxilla,
dilambdodont
molars, and a U- or horseshoe-shaped rhinarium. However, hipposiderids can be differentiated
from rhinolophids using a number of different characteristics. Hipposiderids generally
have a more rounded noseleaf, while the noseleaf of rhinolophids is spear-like and
pointed. Hipposiderids have only two bones in each toe, while rhinolophids have three
in all except the first toe, which has two. Rhinolophids always have three lower premolars
on each side of the mandible and hipposiderids have only two. The two families also
differ in the structure of their shoulder and hip girdles. Finally, rhinolophids have
a sella, a flattened leaflet in the middle of the noseleaf structure, that is not
present in hipposiderids.
- Other Physical Features
- endothermic
- heterothermic
- homoiothermic
- bilateral symmetry
- Sexual Dimorphism
- sexes alike
Reproduction
Not enough information is known about hipposiderid mating systems to make accurate
generalizations about the family as a whole; however, research on individual species
provides limited but important insight. Only one example of a polygynous mating in
hipposiderids is known. Colonies of
Commerson's roundleaf bat
, which can contain up to 500,000 individuals, are divided into small harems consisting
of one adult male and several adult females, with whom the male mates. Mating occurs
seasonally, during the fall, and females give birth to a single young during spring
after storing sperm over winter.
- Mating System
- polygynous
Breeding season and birthing season vary among hipposiderid species. For example,
bi-colored leaf-nosed bats
and
ashy roundleaf bats
mate in October and give birth in April. Although birthing season varies slightly,
coinciding with peak rainy season when food is most abundant,
Sundevall's leafnosed bats
give birth in April north of the equator and in October south of the equator.
Fulvus roundleaf bats
mate in November and give birth in late April. Although the specific times vary among
species, birthing among hipposiderids generally occurs during spring. Female hipposiderids
give birth to a single young per pregnancy. Gestation lasts from 90 days in
cyclops roundleaf bats
to 220 days in
Sundevall's leafnosed bats
in South Africa. Females typically carry their young for a few weeks after giving
birth. For example,
Fulvus roundleaf bats
produce a single young, which the female carries for 20 to 22 days. Age at weaning,
age at first flight, and age at independence appears to vary according to latitude.
Species subject to greater seasonality appear to mature more quickly than those resident
to more tropical regions. In at least one species,
Sundevall's leafnosed bats
of Nigeria, delayed implantation occurs. The egg does not implant in the uterine
lining for up to 2 months after fertilization, and as a result, young are born when
prey are more abundant, directly before the rainy season.
- Key Reproductive Features
- iteroparous
- seasonal breeding
- gonochoric/gonochoristic/dioecious (sexes separate)
- sexual
- viviparous
- delayed fertilization
- delayed implantation
Females are the primary care givers in hipposiderids. Female typically carry their
young for a few weeks after birth and prior to weaning. Females have "pubic teats",
which their young hold on to during the carrying period. Little is known of lactation
and weaning in hipposiderids. However, lactation lasts for about 40 days in the genus
Asellia
, and
Taiwanese leaf-nosed bats
are usually weaned at 7 weeks old. Tropical species are thought to be weaned by 8
to 20 weeks and time to independence appears to vary according to latitude, as tropical
species reach sexual maturity between 16 and 24 months, and temperate species reaching
sexual maturity by 6 to 8 months.
- Parental Investment
- precocial
- female parental care
-
pre-hatching/birth
-
provisioning
- female
-
provisioning
-
pre-weaning/fledging
-
protecting
- female
-
protecting
-
pre-independence
-
provisioning
- female
-
provisioning
Lifespan/Longevity
Information regarding the lifespan of hipposiderids is limited, as a majority of species
in this family are not well-known. However, some species have been found to live
more than 10 years.
Behavior
Hipposiderids, like all bats, are motile and capable of flight. They are nocturnal
and roost in caves, mines, hollow trees, buildings, cellars, and abandoned bomb shelters.
Roosting practices of hipposiderids vary widely between genera and species. While
some roost as individuals, others, such as certain species of
Hipposideros
, roost in small groups. Some species roost in colonies of hundreds or even thousands
of individuals.
Pilbara leaf-nosed bats
have been found in groups of more than 5,000 individuals, and Sundevall’s roundleaf
bats were once discovered in a group of more than 500,000 in a cave in Gabon, Africa.
It has been suggested that large colonies of Sundevall’s roundleaf bats separate into
harem groups consisting of seven adult females and one adult male. Alternatively,
cyclops roundleaf bats
congregate in groups of twelve or fewer females, and no males are usually present.
One species is thought to migrate.
While roosting, hipposiderids do not touch each other, including those species that
roost in large groups. A group of 5,000
Pilbara leaf-nosed bats
were observed to each be spaced at least 15 cm from each other. Other genera are
reported to maintain an interindividual roosting distance of 30 to 40 cm. Hipposiderids
are generally found evenly spaced while roosing. Some species roost with bats of other
genera or even other families. For example,
trident leaf-nosed bats
have been found roosting with
trident bats
as well as
Kuhl's pipistrelle
, and
Pilbara leaf-nosed bats
often roosts with
Australian false vampire bat and ghost bats
. In addition, some members of the genera
Hipposideros
,
Asellia
, and
Coelops
hibernate.
Trident leaf-nosed bats
in the deserts of Iraq were found hibernating in cellars and tombs between mid-September
and mid-November. When roosting, hipposiderids fold their wings around themselves.
Communication and Perception
Like most
Microchiroptera
, members of the family Hipposideridae have relatively small eyes, indicating that
vision may not be as important as echolocation for navigation and foraging purposes.
However, vision may be used to detect objects past the range of echolocation. Hipposiderids,
like all
Microchiroptera
, do not have color vision.
Unlike most
microchiropterans
that emit echolocation via the mouth, hipposiderids produce echolocation sounds with
the larynx and emit the sound through their nostrils. The sounds produced are considered
ultrasonic because they have higher frequencies than the normal range of human hearing.
Hipposiderid echolocation calls contain a long constant-frequency (CF) component (i.e.,
one frequency is maintained throughout the duration of the call) and a much shorter
frequency-modulated (FM) component. The CF segment of the call is used to determine
the general structure of the local environment and to give a coarse location of potential
prey and is preceded or followed by a brief FM segment, which aids in homing in on
the location of a target. In general, the calls of larger bats have a tendency to
be longer and lower in frequency, whereas the calls of smaller bats have a tendency
to be shorter and higher in frequency. Evidence suggests that the calls of hipposiderids
are typically higher in frequency relative to body mass than other bat families. Little
information is available regarding the use of sound and echolocation for intraspecific
communication, though audible sounds may be used to communicate during courtship or
between mother and pup.
Many hipposiderid species have a small sac just posterior to the nose leaf. The sac,
which is possessed primarily by males, secretes a waxy substance that may be used
during mating season to attract mates or fend of potential rivals.
- Perception Channels
- visual
- tactile
- acoustic
- ultrasound
- echolocation
- chemical
Food Habits
Although little information is available on the diets of most hipposiderid species,
they are considered to be primarily insectivorous. Those species that have been studied
prefer cicadas, cockroaches, termites, and beetles. The beetle larvae prey of
Commerson's roundleaf bats
live in wild figs, which results in the addition of small amounts of fruit to their
otherwise insectivorous diet.
Hipposiderids have excellent echolocation, and catch most of their prey via aerial
hawking and gleaning. They usually fly only a few meters above the ground while echolocating
for potential prey. Although most species are thought to prey on flying insects, some
occasionally feed on flightless insects such as
ants
. Hipposiderids are generally territorial and hunt and feed within a specific range.
For example, members of the genus
Asellia
, have been observed flying more than a mile through the desert to their feeding territory.
Often, hipposiderids bring captured prey back to their roost prior to consumption.
When chewing, the jaws of hipposiderids move side-to-side and up and down, simultaneously.
This shearing motion helps break down the chitinous exoskeleton of insect prey.
- Primary Diet
- carnivore
Predation
Hipposiderids are preyed upon by a number of small nocturnal mammals with the ability
to capture them mid-flight or locate their roosts. In many localities, the major predator
of hipposiderids is
snakes
, which are sometimes able to locate their roosting sites. During flight, hipposiderids
can be captured and eaten by various birds of prey including
hawks
,
falcons
, and
owls
. Furthermore, in Australia, members of the family
Dasyuridae
have been known to locate hipposiderid roosts. In conjunction with their ability
to fly, the nocturnal lifestyle of bats helps reduce predation as does the colonial
roosting behavior of many species.
Ecosystem Roles
As insectivores, hipposiderids help control insect pest populations. While little
information exists on potential endoparasites of hipposiderids, like most
bats
, they are probably host to a number of ectoparasitic arthropods including
lice
,
mites
and
fleas
.
- fleas ( Siphonaptera )
- lice ( Phthiraptera )
- mites ( Acari )
Economic Importance for Humans: Positive
As insectivores, hipposiderids help control insect pest populations that might otherwise
spread disease or damage crops. The guano of hipposiderids is locally used as a nitrogen
rich fertilizer.
- Positive Impacts
- produces fertilizer
- controls pest population
Economic Importance for Humans: Negative
Hipposiderids cause little economic damage. There are no known pathogens specific
to Hipposideridae that are harmful to people or domesticated animals. However, bats
occasionally roost in occupied buildings, which can be destructive and has the potential
to spread disease. Any species of bat infected with rabies could potentially bite
and transmit the pathogen to humans.
- Negative Impacts
-
injures humans
- causes disease in humans
- carries human disease
- causes or carries domestic animal disease
- household pest
Conservation Status
As a family, hipposiderids are not a particularly threatened group. However, many
species are not well understood and as a result, potential conservation needs are
unknown. The International Union for Conservation of Nature (IUCN) lists 5 species
as either endangered or critically endangered, and another 10 species are listed as
vulnerable. Of the 84 species listed, 7 are classified as near threatened, 44 are
listed as least concern, and the remaining 18 are classified as data deficient. Habitat
loss and deforestation are serious concerns and their greatest threats. In specific
cases, habitat loss has been so extreme that several species are now classified as
endangered and some local populations are nearing extirpation. For example,
Thailand leaf-nosed bats
have been subjected to severe range contraction due to deforestation, which has resulted
in a population reduction of 20% in just the last 5 years.
Durga Das's leaf-nosed bats
have had nearly all of their native range destroyed and now only roost in the homes
of three different villages in central India. Due to deforestation,
Orbiculus leaf-nosed bats
are now resident to only two locations in Indonesia and Malaysia.
Lamotte's roundleaf bats
can be found in a single cave on the island of Guinea, and are classified as critically
endangered. If conservation efforts are to be successful, habitat loss must be slowed
and reforestation projects should be encouraged in critical habitat areas.
Additional Links
Contributors
Lauren Hall (author), University of Michigan-Ann Arbor, Laura Jadwin (author), University of Michigan-Ann Arbor, Ian Winkelstern (author), University of Michigan-Ann Arbor, Phil Myers (editor), University of Michigan-Ann Arbor, John Berini (editor), Animal Diversity Web Staff.
- Palearctic
-
living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- oriental
-
found in the oriental region of the world. In other words, India and southeast Asia.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Ethiopian
-
living in sub-Saharan Africa (south of 30 degrees north) and Madagascar.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Australian
-
Living in Australia, New Zealand, Tasmania, New Guinea and associated islands.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- tropical
-
the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.
- terrestrial
-
Living on the ground.
- 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.
- tropical savanna and grassland
-
A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.
- savanna
-
A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.
- temperate grassland
-
A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.
- forest
-
forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.
- 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.
- scrub forest
-
scrub forests develop in areas that experience dry seasons.
- 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.
- heterothermic
-
having a body temperature that fluctuates with that of the immediate environment; having no mechanism or a poorly developed mechanism for regulating internal body temperature.
- 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.
- polygynous
-
having more than one female as a mate at one time
- 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).
- seasonal breeding
-
breeding is confined to a particular season
- sexual
-
reproduction that includes combining the genetic contribution of two individuals, a male and a female
- viviparous
-
reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.
- 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.
- young precocial
-
young are relatively well-developed when born
- female parental care
-
parental care is carried out by females
- nocturnal
-
active during the night
- motile
-
having the capacity to move from one place to another.
- hibernation
-
the state that some animals enter during winter in which normal physiological processes are significantly reduced, thus lowering the animal's energy requirements. The act or condition of passing winter in a torpid or resting state, typically involving the abandonment of homoiothermy in mammals.
- solitary
-
lives alone
- social
-
associates with others of its species; forms social groups.
- 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.
- acoustic
-
uses sound to communicate
- chemical
-
uses smells or other chemicals to communicate
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- ultrasound
-
uses sound above the range of human hearing for either navigation or communication or both
- 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.
- chemical
-
uses smells or other chemicals to communicate
- 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
- carnivore
-
an animal that mainly eats meat
- insectivore
-
An animal that eats mainly insects or spiders.
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