Hammer-headed bat

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Hammer-headed bat
Hypsignathus monstrosus.png
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Chiroptera
Family: Pteropodidae
Genus: Hypsignathus
H. Allen, 1861
Species:
H. monstrosus
Binomial name
Hypsignathus monstrosus
H. Allen, 1861
Hammer-headed Bat area.png
Hammer-headed bat range
Synonyms
  • Sphyrocephalus labrosus Murray, 1862
  • Zygaenocephalus labrosus (Murray, 1862)

The hammer-headed bat (Hypsignathus monstrosus), also known as hammer-headed fruit bat, big-lipped bat, and hammerhead bat, [2] [3] is a megabat widely distributed in West and Central Africa. It is the only member of the genus Hypsignathus, which is part of the tribe Epomophorini along with four other genera. It is the largest bat in continental Africa, with wingspans approaching 1 m (3.3 ft), and males almost twice as heavy as females. Males and females also greatly differ in appearance, making it the most sexually dimorphic bat species in the world. These differences include several adaptations that help males produce and amplify vocalizations: the males' larynges (vocal cords) are about three times as large as those of females, and they have large resonating chambers on their faces. Females appear more like a typical megabat, with foxlike faces.

Contents

The hammer-headed bat is frugivorous, consuming a variety of fruits such as figs, bananas, and mangoes, though a few instances of carnivory have been noted. Females tend to travel a consistent route to find predictable fruits, whereas males travel more to find the highest quality fruit. It forages at night, sleeping during the day in tree roosts. Individuals may roost alone or in small groups. Unlike many other bat species that segregate based on sex, males and females will roost together during the day. It has two mating seasons each year during the dry seasons. It is believed to be the only bat species with a classical lek mating system, wherein males gather on a "lek", which in this case is a long and thin stretch of land, such as along a river. There, they produce loud, honking vocalizations to attract females. Females visit the lek and select a male to mate with; the most successful 6% of males are involved in 79% of matings. Offspring are born five or six months later, typically a singleton, though twins have been documented. Its predators are not well-known, but may include hawks. Adults are commonly affected by parasites such as flies and mites.

The hammer-headed bat is sometimes considered a pest due to its frugivorous diet and its extremely loud honking noises at night. In Nigeria and the Democratic Republic of the Congo, it is consumed as bushmeat. It has been investigated as a potential reservoir of the Ebola virus, with several testing positive for antibodies against the virus. It is not considered a species of conservation concern due to its large range and presumably large population size.

Taxonomy and etymology

Hypsignathus

Epomops

Nanonycteris

Epomophorus

Micropteropus

Megaloglossus

Myonycteris

Lissonycteris

Position of Hypsignathus within Pteropodidae [4]

The hammer-headed bat was described as a new species in 1861 by American scientist Harrison Allen. Allen placed the species into a newly-created genus, Hypsignathus. [5] The holotype had been collected by French-American zoologist Paul Du Chaillu [5] in Gabon. [6] The genus name Hypsignathus comes from the Ancient Greek húpsos , meaning "high", and gnáthos , meaning "jaw". T. S. Palmer speculated that Allen chose the name Hypsignathus to allude to the "deeply arched mouth" of the species. [7] The species name monstrosus is Latin for "having the qualities of a monster". [8]

A 2011 study found that Hypsignathus was the most basal member of the tribe Epomophorini, which also includes Epomops , Micropteropus , Epomophorus , and Nanonycteris . [4] Initially, Allen identified the hammer-headed bat as a member of the subfamily Pteropodinae of the megabats. [5] However, in 1997, Epomophorini was recognized as part of the subfamily Epomophorinae. [9] Some taxonomists do not recognize Epomophorinae as a valid subfamily and include its taxa, including the Epomophorini, within Rousettinae. [10] [11]

Description

Internal organs as viewed from the side. Note the extent of the laryngeal structure (vocal folds). Hypsignathus monstrosus side cross section.png
Internal organs as viewed from the side. Note the extent of the laryngeal structure (vocal folds).

The hammer-headed bat is the largest bat in mainland Africa. [12] Males have wingspans up to 90.1 cm (2.96 ft), [13] and all individuals have forearm lengths exceeding 112 mm (4.4 in). [12] It has pronounced sexual dimorphism, more so than any other bat species in the world, [12] with males up to twice as heavy as females. The average weight of males is 420 g (15 oz), compared to 234 g (8.3 oz) for females. [13] Other differences between the sexes relate to their social system, in which males produce loud, honking vocalizations. Therefore, males have greatly enlarged larynges, about three times the size of females', [14] extending through most of the thoracic cavity, and measuring half the length of the spine. The larynx is so large, it displaces other organs, including the heart, lungs, and gastrointestinal tract. [14] Males also have resonating chambers to increase the volume of sound production. These chambers are pharyngeal air sacs connected to a large sinus in the humped snout. [12] These numerous adaptations caused scientists Herbert Lang and James Chapin to remark, "In no other mammal is everything so entirely subordinated to the organs of voice". [15]

Males overall have boxy heads with enormous lips, while the females, with their narrower snouts, have more foxlike faces. [14] Males and females both have dark brown fur, with a paler mantle (sides and back of neck). It has patches of white fur at the base of the ears, though sometimes indistinct. The fur is long and smooth, though somewhat woolly in texture on the mantle. The ears are triangular and blackish-brown, and the eyes are very large. [12] The dental formula is 2.1.2.12.1.3.2 for a total of 28 teeth; very occasionally, individuals have been found with an additional upper premolar on each side of the mouth, for a total of 30 teeth. The skull is larger and more robust than any other megabat in Africa, with a pronounced, massive snout. The tongue is large and powerful, with an expanded, tridentate tip. The tongue has backwards-facing papillae used to extract juice from fruits. [12]

The wings are characterized by low aspect ratio, meaning that it has a smaller wingspan relative to the wing area. The wing loading is considered exceptionally high, meaning that it has a large body weight relative to the wing area. The wings are blackish brown in color. [12] The thumb is approximately 128–137 mm (5.0–5.4 in) long. [14] The wings attach to the hindlimbs at the second toe. It lacks a tail. [12]

Instead of the typical mammalian karyotype where females have two X chromosomes and males have one each of X and Y, males have a single X chromosome and no Y chromosome, known as X0 sex-determination system. [12] Thus, females have 36 chromosomes (34 autosomes and two sex chromosomes), and males have 35 chromosomes (34 autosomes but only one sex chromosome). [16] This is seen in a few other bat genera, including Epomophorus and Epomops . [17] [18]

Biology and ecology

Diet and foraging

The powerful, tridentate tongue is used to extract juice from fruits Hypsignathus monstrosus tongue.png
The powerful, tridentate tongue is used to extract juice from fruits

Hammer-headed bats are frugivores. Figs make up much of their diet, but mangos, bananas and guavas may also be consumed. There are some complications inherent in a fruit diet such as insufficient protein intake. It is suggested that fruit bats compensate for this by possessing a proportionally longer intestine compared to insectivorous species. [14]

Males and females rely on different strategies for foraging. Females use trap-lining, in which they travel an established route with dependable and predictable food sources, even if the food is lower quality. Males, in contrast, search for areas rich with food, traveling up to 10 km (6.2 mi) to reach particularly good food patches. [14] Upon finding suitable fruit, the hammer-headed bat may eat at the tree or pick the fruit and carry it away to another site for consumption. It chews the fruit, swallowing the juice and soft pulp, before spitting out the rest. [14] The guano (feces) typically contains seeds from ingested fruits, indicating that it may be an important seed disperser. [12]

Van Deusan (1968) reports H. monstrosus showing some carnivorous behavior by attacking chickens to drink their blood and scavenging for meat. [19]

Reproduction

Males have massive resonating chambers on their faces to amplify vocalizations (indicated by dashed line) Hypsignathus monstrosus cheeck pouch.png
Males have massive resonating chambers on their faces to amplify vocalizations (indicated by dashed line)

Little is known about reproduction in hammer-headed bats. In some populations, breeding is thought to take place semi-annually during the dry seasons. The timing of the dry season varies depending on the locality, but in general the first breeding season is from June to August and the second is from December to February. Females may become pregnant up to twice per year, giving birth after five or six months gestation [12] to one offspring at a time, [14] though twins have been reported. [15] Newborns weigh approximately 40 g (1.4 oz) at birth. [15] Females reach sexual maturity faster than males, and can reproduce at six months. Females reach adult size by nine months of age. In contrast, males are not sexually mature until eighteen months. Males and females are similar in size for their first year of life. [14]

This species is often cited as an example of classical lek mating, [20] and is perhaps the only bat species with such. [21] The classical lek is defined by four criteria: [12]

Males form these leks along streams or riverbeds during the mating season, which lasts 13 months. [14] Leks consist of 20135 males in an area about 40 m (130 ft) wide and 400–1,600 m (1,300–5,200 ft) long. [12] Each male claims a display territory of about 10 m (33 ft) in diameter, [13] in which he honks repeatedly and flaps his wings while hanging from a branch. [14] Typically, 60120 honks are produced per minute. [20] Males display for around four hours before foraging, with peaks in lekking activity in the early evening and before dawn. The early evening peak is when the majority of copulation occurs. Females will fly through the lek, selecting a male by landing on a branch next to him. The chosen male emits a "staccato buzz" call, followed immediately by copulation, which lasts 3060 seconds. [14] After copulation, the female immediately departs, and the male resumes displaying. [12] The males at the center of the lek have the most success, and are responsible for the majority of matings: [14] the top 6% of males have 79% of the total matings. [12] In the before-dawn peak in activity, copulation is less frequent, and males spend time jockeying with each other for the best display territory. As the mating season progresses, the importance of the before-dawn peak lessens. [14]

However, some populations of hammer-headed bats in West Africa do not use leks. Instead, they have a harem system. [22]

Behavior

Adult male hammer-headed bat wearing a solar-powered GPS collar to track his movements Hypsignathus monstrosus 2.png
Adult male hammer-headed bat wearing a solar-powered GPS collar to track his movements

During the day, the hammer-headed bat roosts in trees, typically 20–30 m (66–98 ft) above the ground in the forest canopy. Various trees are used for roosting, with no preference for a particular species. It has low fidelity to its roost and will move to a new roost after 59 days. [12] It relies on camouflage to hide from predators. [14] It displays a mix of solitary and social behavior. Individuals of both sexes are frequently found roosting alone, though they may roost in small groups of around four individuals. Occasionally, groups of up to twenty-five have been documented. Groups are of mixed sex and age, unlike other bat species which segregate based on sex. While roosting, individuals in a group are approximately 10–15 cm (3.9–5.9 in) apart, with males on the periphery and females nearer the center. [12] During most of the day, individuals sleep with their noses covered by their wings. [20] Members of the same group show little interaction with each other: they do not "squabble", vocalize, or groom each other. Instead, at sunset, individuals groom themselves then set off independently to forage. [12]

Predators and parasites

Its predators are not well-documented, but may include avian species such as the long-tailed hawk. [23] It has a diverse array of parasites, including such ectoparasites as the bat fly (Nycteribiidae) Dipseliopoda arcuata , the spinturnicid mite Ancystropus aethiopicus , the gastronyssid mite Mycteronyssus polli , and the teinocoptid mite Teinocopties auricularis . [12] Internally, it is known to be affected by the liver parasite Hepatocystis carpenteri . Adults commonly host parasites. [14]

Range and habitat

The hammer-headed bat is a lowland species, always occurring below 1,800 m (5,900 ft) above sea level. [14] Most records of this species occur in rainforest habitat, including lowland rainforest, swamp forest, riverine forests, and mosaics of forest and grassland. While it has been documented in savanna habitats, these records are rare, and it has been speculated that these individuals are vagrants. [12] It has a wide range in West and Central Africa, including the following countries: Angola, Benin, Burkina Faso, Cameroon, Central African Republic, Republic of the Congo, Democratic Republic of the Congo, Ivory Coast, Equatorial Guinea, Ethiopia, Gabon, Ghana, Guinea, Guinea-Bissau, Kenya, Liberia, Nigeria, Sierra Leone, Sudan, Togo, and Uganda. [1]

Interactions with humans

As pests and bushmeat

As a frugivorous species, the hammer-headed bat is sometimes considered a pest of fruit crops. [24] Its ability to produce extremely loud vocalizations means that some consider it one of Africa's most significant nocturnal pests. [12] Humans hunt this large bat and consume it as bushmeat. [24] It is eaten in Nigeria, [25] as well as seasonally in the Democratic Republic of the Congo. [26]

Disease transmission

The hammer-headed bat has been investigated as a potential reservoir of the Ebola virus. Some individuals have tested seropositive for the virus, meaning that they had antibodies against the virus, though the virus itself was not detected. Additionally, nucleic acid sequences associated with the virus have been isolated from its tissues. [27] However, the natural reservoirs of ebolaviruses are still unknown as of 2019. [28] [29] [30] Megabats like the hammer-headed bat tend to be over-sampled relative to other potential Ebola virus hosts, meaning that they may have an unwarranted amount of research attention, and as of 2015, no bat hunter or researcher is known to be the index case ("patient zero") in an Ebola outbreak. [31]

Conservation

As of 2016, the hammer-headed bat is evaluated as a least-concern species by the IUCN its lowest conservation priority. It meets the criteria for this classification because it has a wide geographic range; its population is presumably large; and it is not thought to be experiencing rapid population decline. [1] It is not a common bat species in captivity, though it is kept at the Wrocław Zoo in Poland as of 2020, [32] and was kept at the Bronx Zoo and San Diego Zoo Safari Park in the 1970s and 1980s. In captivity, hammer-headed bats, particularly males, are vulnerable to stress-related illness, especially when moved or placed in new enclosures. [33]

Related Research Articles

<span class="mw-page-title-main">Megabat</span> Family of fruit bats

Megabats constitute the family Pteropodidae of the order Chiroptera (bats). They are also called fruit bats, Old World fruit bats, or—especially the genera Acerodon and Pteropus—flying foxes. They are the only member of the superfamily Pteropodoidea, which is one of two superfamilies in the suborder Yinpterochiroptera. Internal divisions of Pteropodidae have varied since subfamilies were first proposed in 1917. From three subfamilies in the 1917 classification, six are now recognized, along with various tribes. As of 2018, 197 species of megabat had been described.

<span class="mw-page-title-main">Bushmeat</span> Meat hunted in tropical forests

Bushmeat is meat from wildlife species that are hunted for human consumption. Bushmeat represents a primary source of animal protein and a cash-earning commodity for inhabitants of humid tropical forest regions in Africa, Latin America and Asia. Bushmeat is an important food resource in poor, rural communities.

<span class="mw-page-title-main">Giant golden-crowned flying fox</span> Large bat species

The giant golden-crowned flying fox, also known as the golden-capped fruit bat, is a species of megabat endemic to the Philippines. Since its description in 1831, three subspecies of the giant golden-crowned flying fox have been recognized, one of which is extinct. The extinct subspecies was formerly recognized as a full species, the Panay golden-crowned flying fox. Formerly, this species was placed in the genus Pteropus; while it is no longer within the genus, it has many physical similarities to Pteropus megabats. It is one of the largest bat species in the world, weighing up to 1.4 kg (3.1 lb)—only the Indian and great flying fox can weigh more. It has the longest documented forearm length of any bat species at 21 cm (8.3 in).

<span class="mw-page-title-main">Egyptian fruit bat</span> Species of bat

The Egyptian fruit bat or Egyptian rousette is a species of megabat that is found in Africa, the Middle East, the Mediterranean, and the Indian subcontinent. It is one of three Rousettus species with an African-Malagasy range, though the only species of its genus found on continental Africa. The common ancestor of the three species colonized the region in the late Pliocene or early Pleistocene. The species is traditionally divided into six subspecies. It is considered a medium-sized megabat, with adults weighing 80–170 g (2.8–6.0 oz) and possessing wingspans of approximately 60 cm (24 in). Individuals are dark brown or grayish brown, with their undersides paler than their backs.

<span class="mw-page-title-main">Spotted-winged fruit bat</span> Species of bat

The spotted-winged fruit bat is the smallest megabat in the world. It inhabits forests in Brunei, Indonesia, Malaysia, and Thailand.

<span class="mw-page-title-main">Grey-headed flying fox</span> Species of bat

The grey-headed flying fox is a megabat native to Australia. The species shares mainland Australia with three other members of the genus Pteropus: the little red P. scapulatus, spectacled P. conspicillatus, and the black P. alecto. The grey-headed flying fox is the largest bat in Australia.

<span class="mw-page-title-main">XO sex-determination system</span> Biological system that determines the sex of offspring

The XO sex-determination system is a system that some species of insects, arachnids, and mammals use to determine the sex of offspring. In this system, there is only one sex chromosome, referred to as X. Males only have one X chromosome (XO), while females have two (XX). The letter O signifies the lack of a Y chromosome. Maternal gametes always contain an X chromosome, so the sex of the animals' offspring depends on whether a sex chromosome is present in the male gamete. Its sperm normally contains either one X chromosome or no sex chromosomes at all.

<span class="mw-page-title-main">Greater short-nosed fruit bat</span> Species of bat

The greater short-nosed fruit bat, or short-nosed Indian fruit bat, is a species of megabat in the family Pteropodidae found in South and Southeast Asia.

<span class="mw-page-title-main">Straw-coloured fruit bat</span> Species of mammal

The straw-coloured fruit bat is a large fruit bat that is the most widely distributed of all the African megabats. It is quite common throughout its area ranging from the southwestern Arabian Peninsula, across forest and savanna zones of sub-Saharan Africa. It is listed as Near Threatened on the IUCN Red List due to a decreasing population trend. Straw-coloured fruit bats travel in massive colonies of at least 100,000 bats and sometimes massing up to 1 million. From October to end of December every year, in the largest migration of mammals on the planet, up to 10 million straw-coloured fruit bats congregate in Kasanka National Park, Zambia, roosting in a 2 hectare area of Mushitu forest each day. This migration was only discovered in 1980. Their necks and backs are a yellowish-brown colour, while their undersides are tawny olive or brownish.

<span class="mw-page-title-main">Wahlberg's epauletted fruit bat</span> Species of bat

Wahlberg's epauletted fruit bat is a species of megabat in the family Pteropodidae. It is commonly found across southern Africa.

<span class="mw-page-title-main">Buettikofer's epauletted fruit bat</span> Species of mammal

Buettikofer's epauletted fruit bat is a species of megabat in the family Pteropodidae. It is found in Ivory Coast, Ghana, Guinea, Guinea-Bissau, Liberia, Nigeria, Senegal, and Sierra Leone. Its natural habitats are subtropical or tropical moist lowland forest and savanna.

<span class="mw-page-title-main">Peters's dwarf epauletted fruit bat</span> Species of bat

Peters's dwarf epauletted fruit bat is a species of megabat in the family Pteropodidae. It is found in Angola, Benin, Burkina Faso, Burundi, Cameroon, Central African Republic, Chad, Republic of the Congo, Democratic Republic of the Congo, Ivory Coast, Equatorial Guinea, Ethiopia, Gabon, Gambia, Ghana, Guinea, Guinea-Bissau, Kenya, Liberia, Mali, Niger, Nigeria, Rwanda, Senegal, Sierra Leone, Sudan, Tanzania, Togo, Uganda, and Zambia. Its natural habitats are subtropical or tropical moist lowland forests and moist savanna.

<span class="mw-page-title-main">Little collared fruit bat</span> Species of bat

The little collared fruit bat is a species of megabat in the family Pteropodidae found in Angola, Cameroon, the Central African Republic, the Republic of the Congo, the Democratic Republic of the Congo, Ivory Coast, Equatorial Guinea, Gabon, Ghana, Guinea, Liberia, Nigeria, Rwanda, Sierra Leone, South Sudan, Togo, and Uganda. Its natural habitats are subtropical or tropical moist lowland forests and moist savanna.

<span class="mw-page-title-main">Indian flying fox</span> Species of mammal

The Indian flying fox, also known as the greater Indian fruit bat, is a species of flying fox native to the Indian subcontinent. It is one of the largest bats in the world. It is of interest as a disease vector, as it is capable of transmitting several viruses to humans. It is nocturnal and feeds mainly on ripe fruits, such as mangoes and bananas, and nectar. This species is often regarded as vermin due to its destructive tendencies towards fruit farms, but the benefits of its pollination and seed propagation often outweigh the impacts of its fruit consumption.

<span class="mw-page-title-main">Madagascan flying fox</span> Species of bat

The Madagascan flying fox, Madagascar flying-fox, or Madagascar fruit bat is a species of megabat in the genus Pteropus. It is endemic to Madagascar. Its natural habitats are diverse, and include moist lowland forests, dry forests, succulent woodlands, and spiny thickets, and mangroves. It eats figs and other fruits, flowers, and leaves. It is threatened by habitat loss.

<span class="mw-page-title-main">Large flying fox</span> Species of fruit bat

The large flying fox, also known as the greater flying fox, Malayan flying fox, Malaysian flying fox, large fruit bat, kalang, or kalong, is a southeast Asian species of megabat in the family Pteropodidae. Despite its scientific name, it feeds exclusively on fruits, nectar, and flowers, like the other flying foxes of the genus Pteropus. It is noted for being one of the largest bats. As with nearly all other Old World fruit bats, it lacks the ability to echolocate but compensates for it with well-developed eyesight.

<span class="mw-page-title-main">Bat</span> Order of flying mammals

Bats are flying mammals of the order Chiroptera. With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most birds, flying with their very long spread-out digits covered with a thin membrane or patagium. The smallest bat, and arguably the smallest extant mammal, is Kitti's hog-nosed bat, which is 29–34 millimetres in length, 150 mm (6 in) across the wings and 2–2.6 g in mass. The largest bats are the flying foxes, with the giant golden-crowned flying fox reaching a weight of 1.6 kg and having a wingspan of 1.7 m.

<span class="mw-page-title-main">Bat virome</span> Group of viruses associated with bats

The bat virome is the group of viruses associated with bats. Bats host a diverse array of viruses, including all seven types described by the Baltimore classification system: (I) double-stranded DNA viruses; (II) single-stranded DNA viruses; (III) double-stranded RNA viruses; (IV) positive-sense single-stranded RNA viruses; (V) negative-sense single-stranded RNA viruses; (VI) positive-sense single-stranded RNA viruses that replicate through a DNA intermediate; and (VII) double-stranded DNA viruses that replicate through a single-stranded RNA intermediate. The greatest share of bat-associated viruses identified as of 2020 are of type IV, in the family Coronaviridae.

<span class="mw-page-title-main">Bat as food</span> Bat eaten as food

Bats as food are eaten by people in parts of some North American, Asian, African, Pacific Rim countries, and cultures, including the United States, China, Vietnam, Seychelles, the Philippines, Indonesia, Palau, Thailand, and Guam. Half the megabat species are hunted for food but only eight percent of the insectivorous bat species are. In Guam, Mariana fruit bats are considered a delicacy.

<span class="mw-page-title-main">Rousettinae</span> Subfamily of bats

The Rousettinae are a subfamily of megabats. Taxa within this subfamily include:

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