Old World Leaf-Nosed Bats (Hipposideridae)

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Old World leaf-nosed bats

(Hipposideridae)

Class Mammalia

Order Chiroptera

Suborder Microchiroptera

Family Hipposideridae


Thumbnail description
This large family is characterized by elaborate modifications of the nose and muzzle, forming leaf-like projections that are thought to help focus echolocation signals emitted through the nose

Size
Range in size from small to very large, with head and body lengths of 1.1–4.3 in (28–110 mm), and forearms a similar length

Number of genera, species
9 genera; 66 species

Habitat
Occupy a variety of habitats, from arid to tropical areas throughout much of the Old World

Conservation status
Critically Endangered: 2 species; Endangered: 1 species; Vulnerable: 15 species; Lower Risk/Near Threatened: 23 species; Data Deficient: 7 species

Distribution
Old World tropics and subtropics

Evolution and systematics

Similar in diversity, size, and characteristics to horseshoe bats (family Rhinolophidae), Old World leaf-nosed bats are sometimes considered a subfamily (Hipposiderinae, but sometimes referred to as the subfamily Rhinonycterinae) of Rhinolophidae. Recent analyses of large molecular and morphological datasets place Hipposideridae in a large group along with the bat families Craseonycteridae, Rhinopomatidae, Nycteridae, Megadermatidae, and Rhinolophidae, all of which are exclusively Old World in distribution.

A recent study employing primarily nuclear sequence data suggests that the group, including Hipposideridae (with the exception of Nycteridae), is more closely related to flying foxes (Megachiroptera) than to other microbats (Microchiroptera). This study also provides some morphological support for their assertions. If this finding is supported by additional evidence and analyses, it would mean that either the general characteristics associated with microbats (echolocation and various morphological features) have evolved twice independently or that they have been lost in the megachiropteran lineage.

A systematic analysis, based on morphological features, of relationships among bats in the family Hipposideridae suggested that the genus Hipposideros is actually composed of three distinct groups, which may deserve generic status. In general, relationships among hipposiderid bats are poorly understood and additional research is needed to fully understand the pattern of evolutionary history in this group. These bats are known from the Eocene to Oligocene in the fossil record.

Physical characteristics

Bats in this family vary greatly in size, from small to very large. Head and body lengths are 1.1–4.3 in (2.8–11 cm) and forearms are a similar length. One of the largest insectivorous bat species is a hipposiderid, Commerson's leaf-nosed bat (Hipposideros commersoni). They are characterized by their elaborate, leaf-like nose leaves, which are composed of an anterior, horseshoe-shaped portion and a posterior portion that is often lobed. Lateral leaflets are also present in many species. These elaborate facial appendages seem to be related to their use of nasal echolocation, where nose leaves act to focus and modify emitted echolocation signals. The ears of these bats vary in size but always lack a tragus, the anterior ear appendage found in many microbats. Members of the families Hipposideridae and Rhinolophidae share a unique feature of the premaxillary bones, whereby premaxillae on either side of the skull are not fused to each other or to the maxillae. Fur coloration is generally shades of brown and red. Tail length ranges from zero (nonexistant) to 2.4 in (6 cm).

Their teeth are much like those of other insectivorous bats, the dental formula is I1/2 C1/1 P1–2/2–3 M3/3.

Distribution

Hipposiderids are found mainly in tropical and subtropical regions of the Old World, including Africa, Asia, and Australia.

Habitat

These bats are found in a variety of habitats, from deserts to tropical rainforests. Most are found in moist, lowland areas.

Behavior

The majority of species in the family Hipposideridae are only poorly known. They are a diverse group and exhibit a variety of life history strategies, social structures, and behaviors. Some northern populations of this family hibernate, such as some of the genus Hipposideros, while others are active year-round, and

only one species is thought to migrate. Most roost in groups varying in size from small (as few as 12) to very large (5,000) congregations, though some are solitary. Roosting often occurs in caves and tunnels, but some species also roost in hollow trees, human structures, and the burrows of animals. The fulvus roundleaf bat (Hipposideros fulvus) roosts in African porcupine (Hystrix) burrows.

Many hipposiderid species have a small sac that sits behind the nose leaf. The sac secretes a waxy substance and is mainly found in males, suggesting the possibility that it is used in social or reproductive interactions for attracting mates or for male competiton.

Feeding ecology and diet

Hipposiderids are generally insectivorous. Little information is available on the specific diets of most species, although most seem to capture insects in flight. Many species return to a roost to eat captured prey.

These bats seem to fly with their mouths closed, emitting ultrasonic pulses through the nose. The many and various modifications of the nose leaf are presumably adaptations to specific modes and frequencies of nasal echolocation. Indeed, some evidence supports a proportional relationship between call frequency and nose leaf width in several species of hipposiderids. This relationship was independent of both body size and generic affiliation. Where studied, echolocation calls tend to be constant frequency and frequency modulated, with frequencies potentially varying with sex and age of individuals.

Reproductive biology

In general, very little is known about reproduction in these bats, but they may be polygynous. Females typically give birth to a single young each year, with breeding occurring seasonally. Hipposiderids mate during the fall and females store the sperm internally during the winter, becoming fertilized and giving birth the following year. Age of weaning, age at first flight, and age at sexual maturity may be correlated with latitude, with tropical species taking longer than temperate species to achieve all of the above landmarks. In Hipposideros, eight tropical species reached sexual maturity at 16–24 months, one subtropical species at less than 12 months, and two temperate species at 6–8 months. The subtropical species (H. terasensis) is weaned at seven weeks old, while tropical species are weaned at 8–20 weeks old.

Conservation status

The majority of bats in this large family are poorly studied, so potential conservation concerns are unknown. Some species are common, others are rare and restricted to islands, making them especially vulnerable to threats such as habitat and roost destruction. The IUCN lists two species as Critically Endangered, one as Endangered, 15 as Vulnerable, 23 as Lower Risk/Near Threatened, and 7 as Data Deficient.

Significance to humans

These primarily insectivorous bats help to control insect pest populations throughout their ranges. Their dung may also be used locally as a fertilizer.

Species accounts

List of Species

Trident leaf-nosed bat
Diadem roundleaf bat
Fulvus roundleaf bat
Noack's roundleaf bat
Golden horseshoe bat

Trident leaf-nosed bat

Asellia tridens

taxonomy

Asellia tridens (Geoffroy, 1813), Egypt.

other common names

None known.

physical characteristics

Medium-sized bats. Total body length 1.8–4.2 in (46–62 mm); forearm length 1.7–2.0 in (45–52 mm); tail length 0.7–2.0 in (18–27 mm); weight 0.2–0.35 oz (6–10 g). Fur coloration varies considerably, ranging from pale yellow to buffy gray and orange-brown, membranes are slightly darker, and the large ears and face are pale. The nose leaf is distinctive, having a large leaf behind the nostrils, with three toothed projections.

distribution

Found throughout Africa north of the Sahara desert, the Arabian Peninsula, and into Pakistan.

habitat

One of the most arid-adapted and common of bats in the areas in which they occur. They have been observed roosting in caves, wells, irrigation culverts, and other man-made structures. They are thought to travel between summer roosts and hibernation sites in caves and tombs. Foraging seems to occur in palm groves and other vegetated areas, the bats sometimes flying many miles (kilometers) across barren terrain to suitable feeding sites.

behavior

A gregarious species, roosts of several hundred individuals have been routinely observed, with one roost numbering as many as

feeding ecology and diet

Eat primarily coleopterans (beetles) and hymenopterans (bees, ants, and wasps) that are taken during flight. They have been recorded foraging mainly in cluttered environments, implying that they are agile in flight and may be able to take prey from the ground and other surfaces.

reproductive biology

Available evidence suggests that females are pregnant in the spring and give birth to a single young in early summer. Gestation time is estimated at 9–10 weeks, and the young nurse for 40 days after birth, after which they become independent. The time of mating is unknown. Most likely polygynous.

conservation status

Common throughout their range, though disturbance to roosting sites could severely impact local populations.

significance to humans

Control insect pest populations throughout their range and are likely to benefit agricultural interests.


Diadem roundleaf bat

Hipposideros diadema

taxonomy

Hipposideros diadema (Geoffroy, 1813), Indonesia.

other common names

English: Large Malayan leaf-nosed bat.

physical characteristics

Medium-sized bat. Weight 1.1–1.7 oz (34–50 g). The fur is dark brown to black and a white spot occurs at the juncture of the forearm and shoulder. The ears are large and the nose leaf is fairly simple, with an anterior horseshoe portion and a posterior, uninterrupted, ridge-like leaflet.

distribution

Found throughout Southeast Asia from Myanmar and Vietnam through Thailand, western Malaysia, and Indonesia to New Guinea, the Bismarck Archipelago, Solomon Islands, Philippines, Nicobar islands, and northeast and north-central Australia.

habitat

Occur primarily in moist, tropical forests.

behavior

Little is known of behavior in this species. They are gregarious, roosting in groups in caves.

feeding ecology and diet

No detailed study of diet has been conducted. Insectivorous, likely to hunt from perches, capturing prey when it is detected by using echolocation.

reproductive biology

Females congregate in March and April, giving birth to a single young during that time. Young probably are weaned and become able to fly between two and three months of age. Nothing is known of their mating behavior; but they are most likely polygynous.

conservation status

Not threatened. They are widespread and common throughout their range.

significance to humans

Consume insect pests.


Fulvus roundleaf bat

Hipposideros fulvus

taxonomy

Hipposideros fulvus Gray, 1838, India.

other common names

English: Fulvous leaf-nosed bat.

physical characteristics

Medium-sized bats, with simple nose leaves.

distribution

South Asia, from Pakistan and Vietnam south to Sri Lanka.

habitat

Tropical, lowland forested habitats.

behavior

Found roosting in the burrows of porcupines (Hystrix). Both smaller (10–20) and larger (50–100) colonies have been observed in caves.

feeding ecology and diet

Insectivorous, though detailed studies of their diet have not been conducted.

reproductive biology

In India, they mate in mid November and give birth to single young in late April and early May. The gestation period is 150–160 days. The female carries her young for 20–22 days and sexual maturity is reached at 18–19 months old. A banded female was captured at an age of at least 12 years and was pregnant at the time of capture. This species is probably polygynous.

conservation status

Has not been evaluated by the IUCN. They are common throughout their range, though population trends are unknown.

significance to humans

Consume insect pests.


Noack's roundleaf bat

Hipposideros ruber

taxonomy

Hipposideros ruber (Noack, 1893), Tanzania.

other common names

English: Noack's African leaf-nosed bat.

physical characteristics

These bats have a simple nose leaf; the anterior and posterior leaflets are simple in outline, without indentations. Their fur is long, silky, and light brown, and the ears are large and round.

distribution

Central Africa, from Senegal to Ethiopia and south to Angola, Zambia, Malawi, and Mozambique, including Bioko and São Tomé and Principe.

habitat

Tropical lowland areas.

behavior

Use a constant frequency echolocation call that differs between the sexes. Some evidence suggests that frequencies used correspond with body condition indices, making it possible for females to discern which mates are in the best condition. Frequency was also found to change with humidity. Little is known about roosting and social behaviors.

feeding ecology and diet

Feed on insects that are captured from hunting perches when detected by echolocation.

reproductive biology

Reproduction dynamics are unknown in this species; but they are most likely polygynous.

conservation status

Has not been evaluated by the IUCN. They are common throughout their range, although population trends are unknown.

significance to humans

Controls insect pests.


Golden horseshoe bat

Rhinonicteris aurantia

taxonomy

Rhinonicteris aurantia (Gray, 1845), Northern Territory, Australia.

other common names

Orange leaf-nosed bat.

physical characteristics

Head and body length 1.7–2.0 in (45–53 mm); forearm length 1.8–1.9 in (47–50 mm); tail length 0.9–1.1 in (24–28 mm; weight 0.28–0.35 oz (8–10 g). The fur is long, silky, and typically bright orange above and paler below, though specimens vary from dark brown to almost white. Characterized by their unique nose leaf; the anterior portion is a divided horseshoe and the posterior portion is deeply indented, making it appear a bit like lace.

distribution

Across northern Australia, including north Western Australia, Northern Territory, and northwestern Queensland.

habitat

Found in a variety of habitats, from open grassland to dense forests. They roost in hot, humid mines and caves.

behavior

Do not hibernate and seem unable to become torpid. They become lethargic at temperatures below 68°F (20°C). They emerge up to 1.5 hours after sunset and forage up to 9.8 ft (3m) above vegetation. Their flight is described as somewhat faster than most hipposiderids. They congregate in roosts of several hundred individuals, with up to 5,000 being observed in a single roost.

feeding ecology and diet

Eat primarily moths.

reproductive biology

Breeding occurs during the wet season, from October–April. Probably polygynous.

conservation status

Considered Vulnerable by the IUCN, populations seem to be in decline. Several important roosts have suffered from human disturbance. Cave gating has led to modest increases in the populations of previously declining roosts.

significance to humans

Controls insect pests.

Common name / Scientific name / Other common namesPhysical characteristicsHabitat and behaviorDistributionDietConservation status
Persian trident bat Triaenops persicusCoat color highly variable, ranging from brown to gray to reds, with some individuals being almost white. The anterior portion of the nose leaf is similar to Rhinonicteris, which is thought to be closely related. The posterior portion of the nose leaf, however, is tridentate. Head and body length 1.4–2.4 in (3.5–6.2 cm), tail length 0.8–1.3 in (2.0–3.4 cm), and forearm length 1.8–2.2 in (4.5–5.5 cm), adults weigh 0.3–0.5 oz (8–15 g).These bats roost in smaller groups, often associated with larger groups of other species. They emerge early in the evening, before dusk, to forage and typically fly close to the ground. They have been observed roosting in caves and underground tunnels. Births have been recorded in January, but may occur at other times as well.Africa, including Somalia, Ethiopia, Eritrea, Kenya, Tanzania Uganda, Angola, Zanzibar, Mozambique, the Congo Republic, and Egypt. The Arabian Peninsula, including Oman, Yemen, and Iran.Insects.Not listed by IUCN
Flower-faced bat Anthops ornatusThis species is known only from a few specimens. The fur is gray-buff, long, and silky. The tail is much reduced, otherwise this species is similar to Hipposideros species. The nose leaf is elaborate, with a tri-lobed posterior portion. Head and body length about 2 in (5 cm), forearm length 1.9–2.0 in (4.8–5.1 cm).Nothing known.Solomon Islands.Presumably insects, though no information is available.Vulnerable
East Asian tailless leaf–nosed bat Coelops frithiFur is variable, from brown to black. Ears are short and rounded and the nose leaf lacks lateral leaflets. Head and body length 1.1–2.0 in (2.8–5.0 cm) and forearm length 1.3–1.9 in (3.3–4.7 cm), weight 0.25–0.32 oz (7–9 g).These bats have been recorded roosting in small numbers in caves, human structures, and hollow trees. They may hibernate in caves. Young have been recorded in March, though births may occur at other times of the year as well.South and Southeast Asia, from India to southern China and Vietnam, and into Malaysia, Taiwan, Java, and Bali.Presumably insects, though no information is available.Not listed by IUCN
Common name / Scientific name / Other common namesPhysical characteristicsHabitat and behaviorDistributionDietConservation status
Temminck's trident bat Aselliscus tricuspidatusFur is bright brown above and buffy below. There are lateral nose leaflets on either side of the anterior portion of the nose leaf and the posterior portion is divided into three, bluntly toothed, portions. The tail extends beyond the tail membrane. Smaller bats, head and body length 1.5–1.8 in (3.8–4.5 cm), forearm length 1.3–1.8 in (3.5–4.5 cm), tail length 0.8–1.6 in (2.0–4.0 cm). Weights have been recorded at 0.1–0.14 oz (3.5–4 g).These bats roost in caves. They hang singly and evenly spaced from other bats, but in distinct groups of usually 40 to 50 bats. Females were recorded both pregnant with single young and lactating in May and June.Pacific islands, including the Moluccas, New Guinea, Bismarck Archipelago, Solomon Islands, Vanuatu, and adjacent islands.Presumably insects, though no information is available.Not listed by IUCN
Percival's trident bat Cloeotis percivaliTwo subspecies are recognized based on fur coloration, which varies from sooty to grayish brown and dark to buffy brown, with lighter tips. The tail is well-developed and the nose leaf has three, distinctly pointed processes arising from the posterior nose leaf. The ears are small and round. Head and body length 1.3–2.0 in (3.3–5.0 cm), forearm length 1.2–1.4 in (3.0–3.6 cm), tail length 0.9–1.3 in (2.2–3.3 cm). These small bats weigh 0.13–0.21 oz (3.8–5.9 g).These bats roost in large numbers in narrow-mouthed caves. Individuals roost in tight clusters. Pregnant females have been recorded in October.Sub-Saharan Africa, including Kenya, Tanzania, Zaire, Mozambique, Zambia, Zimbabwe, Botswana, Swaziland, and the Transvaal of South Africa.In some areas these bats have been reported to eat almost exclusively moths.Lower Risk/Near Threatened
Ashy roundleaf bat Hipposideros cineraceusThese bats weigh 0.25–0.28 oz (7–8 g), making them smaller members of the genus HipposiderosA gregarious species, they can be found roosting in large numbers. Reproduction is seasonal, with mating in October, births occurring in April, and the young capable of flight by June.South and Southeast Asia, from Pakistan and northern India to Vietnam, Borneo, and adjacent small islands. May occur in the Philippines.Little information available, presumably insectivorous.Not listed by IUCN
Commerson's leaf-nosed bat Hipposideros commersoni English: Giant leaf-nosed batThis is one of the largest insectivorous bats worldwide, weighing from 2.5 to 3.5 oz (74–100 g).These bats roost in large numbers of several hundred bats. Pregnant females have been recorded in August and October.Africa, from Gambia and Ethiopia south to South Africa, and Madagascar.These bats are sit-and-wait predators, hanging from a perch and flying to intercept large prey (up to 2.6 in [60 mm] long) that are detected via echolocation. Prey are detected at distances up to 65 ft (20 m) and are taken back to the perch for consumption. They have been observed eating beetle larvae inside wild figs and incidentally take fig pulp as well.Not listed by IUCN
Ridley's roundleaf bat Hipposideros ridleyi English: Singapore roundleaf horseshoe bat; Ridley's leaf-nosed batNo specific information available.Little information is available. Survives only in small patches of heavily logged lowland peat forest. Females give birth in April.Western Malaysia and Bomeo.Unknown, presumably insects.Vulnerable

Resources

Books

Bogdanowicz, Wieslaw, and Robert D. Owen. "In the Minotaur's Labryinth: Phylogeny of the Bat Family Hipposideridae." In Bat Biology and Conservation, edited by Thomas H. Kunz and Paul A. Racey. Washington, DC: Smithsonian Institution Press, 1998.

Hill, John E., and James D. Smith. Bats, A Natural History. London: British Museum of Natural History, 1984.

Koopman, Karl F. "Chiroptera: Systematics." In Handbook of Zoology. Volume VIII, Mammalia, edited by J. Niethammer, H. Schliemann, and D. Starck. Berlin: Walter de Gruyter, 1994.

McKenna, Malcolm C., and Susan K. Bell, eds. Classification of Mammals Above the Species Level. New York: Columbia University Press, 1997.

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

Simmons, Nancy B., and Jonathan H. Geisler. "Phylogenetic Relationships of Icaronycteris, Archaeonycteris, Hassianycteris, and Palaeochiropteryx to Extant Bat Lineages, with Comments on the Evolution of Echolocation and Foraging Strategies in Microchiroptera." Bulletin of the American Museum of Natural History, Number 235. New York: American Museum of Natural History, 1998.

Periodicals

Brosset, A. "Recherches sur la Biologie des Chiropteres Troglophiles dans le Nord-est du Gabon." Biologie du Gabon 5 (1969): 93–116.

Cheng, H. C., and L. L. Lee. "Postnatal Growth, Age Estimation, and Sexual Maturity in the Formosan Leaf-nosed Bat (Hipposideros terasensis)." Journal of Mammalogy 83 (2002): 785–793.

Churchill, S. H. "Reproductive Ecology of the Orange Horseshoe Bat, Rhinonycteris aurantius (Chiroptera: Hipposideridae), a Tropical Cave-dweller." Wildlife Research 22 (1995): 687–698.

Habersetzer, J., and G. Marimuthu. "Ontogeny of Sounds in the Echolocating Bat Hipposideros speoris." Journal of Comparative Physiology, A. Sensory, Neural, and Behavioral Physiology 158 (1986): 247–257.

Hill, J. E. "A Revision of the Genus Hipposideros." Bulletin of the British Museum of Natural History, Zoology 11 (1963): 1–129.

Jones, G., M. Morton, P. M. Hughes, and R. M. Budden. "Echolocation, Flight Morphology, and Foraging Strategies of Some West African Hipposiderid Species." Journal of the Zoological Society of London 230 (1993): 385–400.

Robinson, Mark F. "A Relationship between Echolocation Calls and Nose-leaf Widths in Bats of the Genera Rhinolophus and Hipposideros." Journal of the Zoological Society of London 239 (1996): 389–393.

Teeling, Emma C., Ole Madsen, Ronald A. Van Den Bussche, Wilfried W. de Jong, and Michael J. Stanhope. "Microbat Paraphyly and the Convergent Evolution of a Key Innovation in Old World Rhinolophoid Microbats." Proceedings of the National Academy of Sciences 99 (2002): 1431–1436.

Organizations

Gulf of Guinea Conservation Group. CP289, Sao Tome, Sao Tome e Principe. Phone: (230) 225428. E-mail: [email protected] Web site: <http://www.ggcg.st/>

Tanya Dewey

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