Triaenops menamena
Encyclopedia
Triaenops menamena is a bat in the genus Triaenops
found on Madagascar
, mainly in the drier regions. It was known as Triaenops rufus until 2009, when it was discovered that that name had been incorrectly applied to the species. Triaenops rufus is in fact a synonym
of Triaenops persicus
, a Middle Eastern species closely related to T. menamena—indeed, the Malagasy species had previously been placed as a subspecies
of T. persicus by some authors. Triaenops menamena is mostly found in forests, but also occurs in other habitats. It often roosts in large colonies and eats insects such as butterflies and moths
. Because of its wide range, common occurrence, and tolerance of habitat degradation, it is not considered to be threatened.
With a forearm length of 50 to 56 mm (2.0 to 2.2 in) in males and 46 to 53 mm (1.8 to 2.1 in) in females, this is a medium-sized bat. Its fur color is variable, ranging from reddish-brown to gray, but it is generally darker than the species in the closely related genus Paratriaenops which also occur on Madagascar. The skull contains a pronounced swelling around the nose and the second upper premolar
is displaced outside the toothrow. The maximum frequency of the echolocation
call averages 94.2 kHz
and the species can easily be recognized on the basis of its call.
described two new species in the genus Triaenops
on the basis of specimens supposedly collected by Léon Humblot on Madagascar: the reddish Triaenops rufus and the larger, gray T. humbloti. Jean Dorst
, who reviewed Madagascan Triaenops in 1947, retained both as separate species; however, in another review, published in 1982, John Edwards Hill considered the two to represent the same species. In 1994, Karl Koopman
considered rufus itself to be part of the Middle Eastern and mainland African species Triaenops persicus
, a possibility Hill had discussed, but most authors regarded the two as distinct species. In 2006, Julie Ranivo and Steven Goodman
revised Madagascan Triaenops and found little variation among specimens of T. rufus from throughout the dry parts of the island. Three years later, they published another paper which showed that the original material of T. rufus and T. humbloti was distinct from Madagascan specimens identified as "Triaenops rufus" and more similar to T. persicus. On his journey to Madagascar, Humblot had stayed in Somalia and Yemen, and Goodman and Ranivo concluded that he had probably collected the Triaenops there, after which their provenance was incorrectly recorded. Thus, rufus and humbloti cannot be used for the Madagascan species, and Goodman and Ranivo proposed the new name Triaenops menamena for the species formerly known as T. rufus. The specific name menamena is Malagasy
for "reddish", referring to the animal's coloration. "Rufous Trident Bat" has been used as a common name
for Triaenops rufus.
Triaenops menamena is currently one of four living species in the genus Triaenops; a 2009 revision by Petr Benda and Peter Vallo split off the African T. afer and the Yemeni T. parvus from T. persicus and removed three other species, including two from Madagascar, to the separate genus Paratriaenops. An extinct species, Triaenops goodmani
, is known from northwestern Madagascar. In 2007 and 2008, Amy Russell and colleagues used phylogenetic and coalescent
methodologies to investigate the history of the Triaenops group. They found that T. menamena (as T. rufus) was closest to mainland African Triaenops (but did not study Middle Eastern bats) and concluded that T. menamena and the species of Paratriaenops (then still placed in Triaenops) independently reached Madagascar from Africa; the colonization of the island by T. menamena was dated to about 660,000 years ago. Benda and Vallo also studied phylogenetic relationships in Triaenops and included Middle Eastern T. persicus and T. parvus in their analysis. They found little resolution of relationships within Triaenops, but some evidence suggested that T. menamena is more closely related to the Middle Eastern species than to T. afer in mainland Africa; therefore, T. menamena may have reached Madagascar from the Middle East or northeastern Africa. They also placed the split between T. menamena and the other species much further back, at around 4 million years ago.
Triaenops menamena is a medium-sized species with variable fur coloration, ranging from reddish-brown to gray. It is larger and darker than Paratriaenops auritus and P. furculus. Among living Triaenops species, it is smaller than T. persicus and T. afer, but somewhat larger than T. parvus. The extinct Triaenops goodmani, which is known only from three mandible
s (lower jaws), is also larger. In the complexly shaped noseleaf (a group of fleshy structures around the nose and mouth), T. menamena has the trident structure characteristic of Triaenops and Paratriaenops: three adjacent lancets (projecting structures) on the posterior leaf. In T. menamena, the two outer lancets are shorter than the middle one and curved, whereas the three lancets are more equal in Paratriaenops. The anterior leaf contains a broad, flat, horizontal process
and the intermediate leaf contains a pointed process. The ears are small and broad and contain notches on the inner side. The fur is silky and the hairs on the body are about 5 to 6 mm long. The wing membrane is dark and translucent and the end of the tail usually projects from the uropatagium (tail membrane). Males average slightly larger than females. Wingspan is 270 to 305 mm (10.6 to 12.0 in).
The skull is similar to that of Madagascan Paratriaenops, but the premaxilla
is longer. The rostrum (front part of the skull) is well-developed and contains a pronounced nasal swelling, which is less pronounced than in Paratriaenops; the depression behind the swelling is relatively shallow. Compared to other species of Triaenops, the rostrum is relatively narrow and short, similar to T. parvus; T. afer and T. persicus have a broader rostrum. The front margin of the rostrum is cast back between the left and right corners. A transverse line passes over the roof of the rostrum. The infraorbital foramen
, an opening in the skull, is oblong in shape. The zygomatic arch
es (cheekbones) are not broadened towards the sides, are connected to the maxilla
ries by broad bones, and contain clear crests on their upper sides. In T. menamena, these crests are rectangular in form and larger than in Paratriaenops. The braincase is lower than in other species of Triaenops. The sagittal crest
, which is on the roof of the braincase, is poorly developed. In the mandible, the coronoid process
is blunt and rounded, but the angular process (both processes
at the back of the bone) is small.
The upper incisor
s have two cusps and the upper canine has three—a large central one, a small but well-developed one at the back, and a small cuspule at the front. The front upper premolar
is projected outside the toothrow, so that the canine before it and back premolar behind it touch. The first lower incisor has two cusps and the second has three. On the second lower molar, the protoconid cusp is notably taller than the hypoconid; these cusps are about as high in T. goodmani.
The species can easily be identified from recordings of its echolocation
call. The call consists of a component with constant frequency followed by a short one with changing frequency. The call takes 6.5 to 13.5 ms
, averaging 10.1 ms, and the period between two calls is 22.7 to 86.3 ms, averaging 42.7 ms. The maximum frequency averages 94.2 kHz, the minimum frequency averages 82.0 kHz, and the call emits the most energy at a frequency of 93.2 kHz.
of the species is estimated to be about 121,000. Lepidoptera
(butterflies and moths) form the main component of its diet, but it also eats Coleoptera (beetles) and Hemiptera
(bugs) and fewer members of some other insect orders. It is unclear how the ecological niche
s of Triaenops and the less frequently captured Paratriaenops furculus are separated, as both eat lepidopterans and occur in the same regions.
lists this species as "Least Concern
", citing its common occurrence over a wide distribution and tolerance of human modifications of its habitat, even though its forest habitat is being destroyed in places. It has been recorded in numerous protected area
s. Bats, mainly the large Hipposideros commersoni, are sometimes hunted for food in southwestern Madagascar, and T. menamena is also taken incidentally.
Triaenops
Triaenops is a genus of bat in the family Hipposideridae. It is classified in the tribe Triaenopini, along with the closely related genus Paratriaenops and perhaps the poorly known Cloeotis. The species of Paratriaenops, which occur on Madagascar and the Seychelles, were placed in Triaenops until...
found on Madagascar
Madagascar
The Republic of Madagascar is an island country located in the Indian Ocean off the southeastern coast of Africa...
, mainly in the drier regions. It was known as Triaenops rufus until 2009, when it was discovered that that name had been incorrectly applied to the species. Triaenops rufus is in fact a synonym
Synonym (taxonomy)
In scientific nomenclature, a synonym is a scientific name that is or was used for a taxon of organisms that also goes by a different scientific name. For example, Linnaeus was the first to give a scientific name to the Norway spruce, which he called Pinus abies...
of Triaenops persicus
Triaenops persicus
Triaenops persicus, also known as the Persian Trident Bat or Triple Nose-leaf Bat, is a species of bat in the genus Triaenops. It occurs in southwestern Pakistan, southern Iran, the United Arab Emirates, Oman, and Yemen. In the last country, it occurs together with the much smaller Triaenops parvus...
, a Middle Eastern species closely related to T. menamena—indeed, the Malagasy species had previously been placed as a subspecies
Subspecies
Subspecies in biological classification, is either a taxonomic rank subordinate to species, ora taxonomic unit in that rank . A subspecies cannot be recognized in isolation: a species will either be recognized as having no subspecies at all or two or more, never just one...
of T. persicus by some authors. Triaenops menamena is mostly found in forests, but also occurs in other habitats. It often roosts in large colonies and eats insects such as butterflies and moths
Lepidoptera
Lepidoptera is a large order of insects that includes moths and butterflies . It is one of the most widespread and widely recognizable insect orders in the world, encompassing moths and the three superfamilies of butterflies, skipper butterflies, and moth-butterflies...
. Because of its wide range, common occurrence, and tolerance of habitat degradation, it is not considered to be threatened.
With a forearm length of 50 to 56 mm (2.0 to 2.2 in) in males and 46 to 53 mm (1.8 to 2.1 in) in females, this is a medium-sized bat. Its fur color is variable, ranging from reddish-brown to gray, but it is generally darker than the species in the closely related genus Paratriaenops which also occur on Madagascar. The skull contains a pronounced swelling around the nose and the second upper premolar
Premolar
The premolar teeth or bicuspids are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant, making eight premolars total in the mouth. They have at least two cusps. Premolars can be considered as a 'transitional tooth' during chewing, or...
is displaced outside the toothrow. The maximum frequency of the echolocation
Animal echolocation
Echolocation, also called biosonar, is the biological sonar used by several kinds of animals.Echolocating animals emit calls out to the environment and listen to the echoes of those calls that return from various objects near them. They use these echoes to locate and identify the objects...
call averages 94.2 kHz
Hertz
The hertz is the SI unit of frequency defined as the number of cycles per second of a periodic phenomenon. One of its most common uses is the description of the sine wave, particularly those used in radio and audio applications....
and the species can easily be recognized on the basis of its call.
Taxonomy
In 1881, Alphonse Milne-EdwardsAlphonse Milne-Edwards
Alphonse Milne-Edwards was a French mammalologist, ornithologist and carcinologist. He was English in origin, the son of Henri Milne-Edwards and grandson of Bryan Edwards, a Jamaican planter who settled at Bruges .Milne-Edwards obtained a medical degree in 1859 and became assistant to his father...
described two new species in the genus Triaenops
Triaenops
Triaenops is a genus of bat in the family Hipposideridae. It is classified in the tribe Triaenopini, along with the closely related genus Paratriaenops and perhaps the poorly known Cloeotis. The species of Paratriaenops, which occur on Madagascar and the Seychelles, were placed in Triaenops until...
on the basis of specimens supposedly collected by Léon Humblot on Madagascar: the reddish Triaenops rufus and the larger, gray T. humbloti. Jean Dorst
Jean Dorst
Professor Dr Jean Dorst was a French ornithologist.Dorst was born at Mulhouse and studied biology and paleontology at the Faculty of Sciences of the University of Paris. In 1947 he joined the staff of the Muséum national d'Histoire naturelle...
, who reviewed Madagascan Triaenops in 1947, retained both as separate species; however, in another review, published in 1982, John Edwards Hill considered the two to represent the same species. In 1994, Karl Koopman
Karl Koopman
Karl Koopman was a zoologist with a special interest in bats. He worked for many years in the Mammalogy Department of the American Museum of Natural History in New York. Species named after him include Monticolomys koopmani, Rattus koopmani, and Sturnira koopmanhilli.-Further reading:...
considered rufus itself to be part of the Middle Eastern and mainland African species Triaenops persicus
Triaenops persicus
Triaenops persicus, also known as the Persian Trident Bat or Triple Nose-leaf Bat, is a species of bat in the genus Triaenops. It occurs in southwestern Pakistan, southern Iran, the United Arab Emirates, Oman, and Yemen. In the last country, it occurs together with the much smaller Triaenops parvus...
, a possibility Hill had discussed, but most authors regarded the two as distinct species. In 2006, Julie Ranivo and Steven Goodman
Steven Goodman
Steven Goodman is an American Conservation Biologist, and field biologist on staff in the Department of Zoology at the Field Museum of Natural History....
revised Madagascan Triaenops and found little variation among specimens of T. rufus from throughout the dry parts of the island. Three years later, they published another paper which showed that the original material of T. rufus and T. humbloti was distinct from Madagascan specimens identified as "Triaenops rufus" and more similar to T. persicus. On his journey to Madagascar, Humblot had stayed in Somalia and Yemen, and Goodman and Ranivo concluded that he had probably collected the Triaenops there, after which their provenance was incorrectly recorded. Thus, rufus and humbloti cannot be used for the Madagascan species, and Goodman and Ranivo proposed the new name Triaenops menamena for the species formerly known as T. rufus. The specific name menamena is Malagasy
Malagasy language
Malagasy is the national language of Madagascar, a member of the Austronesian family of languages. Most people in Madagascar speak it as a first language as do some people of Malagasy descent elsewhere.-History:...
for "reddish", referring to the animal's coloration. "Rufous Trident Bat" has been used as a common name
Common name
A common name of a taxon or organism is a name in general use within a community; it is often contrasted with the scientific name for the same organism...
for Triaenops rufus.
Triaenops menamena is currently one of four living species in the genus Triaenops; a 2009 revision by Petr Benda and Peter Vallo split off the African T. afer and the Yemeni T. parvus from T. persicus and removed three other species, including two from Madagascar, to the separate genus Paratriaenops. An extinct species, Triaenops goodmani
Triaenops goodmani
Triaenops goodmani is an extinct bat from Madagascar in the genus Triaenops. It is known from three lower jaws collected in a cave at Anjohibe in 1996, and described as a new species in 2007. The material is at most 10,000 years old. A bat humerus from the same site could not be identified as...
, is known from northwestern Madagascar. In 2007 and 2008, Amy Russell and colleagues used phylogenetic and coalescent
Coalescent theory
In genetics, coalescent theory is a retrospective model of population genetics. It attempts to trace all alleles of a gene shared by all members of a population to a single ancestral copy, known as the most recent common ancestor...
methodologies to investigate the history of the Triaenops group. They found that T. menamena (as T. rufus) was closest to mainland African Triaenops (but did not study Middle Eastern bats) and concluded that T. menamena and the species of Paratriaenops (then still placed in Triaenops) independently reached Madagascar from Africa; the colonization of the island by T. menamena was dated to about 660,000 years ago. Benda and Vallo also studied phylogenetic relationships in Triaenops and included Middle Eastern T. persicus and T. parvus in their analysis. They found little resolution of relationships within Triaenops, but some evidence suggested that T. menamena is more closely related to the Middle Eastern species than to T. afer in mainland Africa; therefore, T. menamena may have reached Madagascar from the Middle East or northeastern Africa. They also placed the split between T. menamena and the other species much further back, at around 4 million years ago.
Description
| Sex | nNumber of specimens measured. | Total length | Tail | Hindfoot | Ear | Forearm | Mass |
|---|---|---|---|---|---|---|---|
| Male | 2827 for total length. | 90–104 | 27–38 | 6–9 | 14–17 | 50–56 | 8.2–15.5 |
| Female | 67 | 86–98 | 28–39 | 6–9 | 12–17 | 46–53 | 6.6–11.5 |
| Measurements are in millimeters (except weight in grams) and in the form "minimum–maximum". | |||||||
Triaenops menamena is a medium-sized species with variable fur coloration, ranging from reddish-brown to gray. It is larger and darker than Paratriaenops auritus and P. furculus. Among living Triaenops species, it is smaller than T. persicus and T. afer, but somewhat larger than T. parvus. The extinct Triaenops goodmani, which is known only from three mandible
Mandible
The mandible pronunciation or inferior maxillary bone forms the lower jaw and holds the lower teeth in place...
s (lower jaws), is also larger. In the complexly shaped noseleaf (a group of fleshy structures around the nose and mouth), T. menamena has the trident structure characteristic of Triaenops and Paratriaenops: three adjacent lancets (projecting structures) on the posterior leaf. In T. menamena, the two outer lancets are shorter than the middle one and curved, whereas the three lancets are more equal in Paratriaenops. The anterior leaf contains a broad, flat, horizontal process
Process (anatomy)
In anatomy, a process is a projection or outgrowth of tissue from a larger body. The vertebra has several kinds of processes,such as: transverse process, prezygapophysis, postzygapophysis.-Examples:Examples of processes include:...
and the intermediate leaf contains a pointed process. The ears are small and broad and contain notches on the inner side. The fur is silky and the hairs on the body are about 5 to 6 mm long. The wing membrane is dark and translucent and the end of the tail usually projects from the uropatagium (tail membrane). Males average slightly larger than females. Wingspan is 270 to 305 mm (10.6 to 12.0 in).
The skull is similar to that of Madagascan Paratriaenops, but the premaxilla
Premaxilla
The incisive bone is the portion of the maxilla adjacent to the incisors. It is a pair of small cranial bones at the very tip of the jaws of many animals, usually bearing teeth, but not always. They are connected to the maxilla and the nasals....
is longer. The rostrum (front part of the skull) is well-developed and contains a pronounced nasal swelling, which is less pronounced than in Paratriaenops; the depression behind the swelling is relatively shallow. Compared to other species of Triaenops, the rostrum is relatively narrow and short, similar to T. parvus; T. afer and T. persicus have a broader rostrum. The front margin of the rostrum is cast back between the left and right corners. A transverse line passes over the roof of the rostrum. The infraorbital foramen
Infraorbital foramen
Above the canine fossa is the infraorbital foramen, the end of the infraorbital canal; it transmits the infraorbital artery, vein, and infraorbital nerve.-External links: *...
, an opening in the skull, is oblong in shape. The zygomatic arch
Zygomatic arch
The zygomatic arch or cheek bone is formed by the zygomatic process of temporal bone and the temporal process of the zygomatic bone , the two being united by an oblique suture; the tendon of the Temporalis passes medial to the arch to gain insertion into the coronoid process...
es (cheekbones) are not broadened towards the sides, are connected to the maxilla
Maxilla
The maxilla is a fusion of two bones along the palatal fissure that form the upper jaw. This is similar to the mandible , which is also a fusion of two halves at the mental symphysis. Sometimes The maxilla (plural: maxillae) is a fusion of two bones along the palatal fissure that form the upper...
ries by broad bones, and contain clear crests on their upper sides. In T. menamena, these crests are rectangular in form and larger than in Paratriaenops. The braincase is lower than in other species of Triaenops. The sagittal crest
Sagittal crest
A sagittal crest is a ridge of bone running lengthwise along the midline of the top of the skull of many mammalian and reptilian skulls, among others....
, which is on the roof of the braincase, is poorly developed. In the mandible, the coronoid process
Coronoid process of the mandible
The mandible's coronoid process is a thin, triangular eminence, which is flattened from side to side and varies in shape and size....
is blunt and rounded, but the angular process (both processes
Process (anatomy)
In anatomy, a process is a projection or outgrowth of tissue from a larger body. The vertebra has several kinds of processes,such as: transverse process, prezygapophysis, postzygapophysis.-Examples:Examples of processes include:...
at the back of the bone) is small.
The upper incisor
Incisor
Incisors are the first kind of tooth in heterodont mammals. They are located in the premaxilla above and mandible below.-Function:...
s have two cusps and the upper canine has three—a large central one, a small but well-developed one at the back, and a small cuspule at the front. The front upper premolar
Premolar
The premolar teeth or bicuspids are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant, making eight premolars total in the mouth. They have at least two cusps. Premolars can be considered as a 'transitional tooth' during chewing, or...
is projected outside the toothrow, so that the canine before it and back premolar behind it touch. The first lower incisor has two cusps and the second has three. On the second lower molar, the protoconid cusp is notably taller than the hypoconid; these cusps are about as high in T. goodmani.
The species can easily be identified from recordings of its echolocation
Animal echolocation
Echolocation, also called biosonar, is the biological sonar used by several kinds of animals.Echolocating animals emit calls out to the environment and listen to the echoes of those calls that return from various objects near them. They use these echoes to locate and identify the objects...
call. The call consists of a component with constant frequency followed by a short one with changing frequency. The call takes 6.5 to 13.5 ms
Millisecond
A millisecond is a thousandth of a second.10 milliseconds are called a centisecond....
, averaging 10.1 ms, and the period between two calls is 22.7 to 86.3 ms, averaging 42.7 ms. The maximum frequency averages 94.2 kHz, the minimum frequency averages 82.0 kHz, and the call emits the most energy at a frequency of 93.2 kHz.
Distribution and ecology
Triaenops menamena mainly occurs in the dry regions of western Madagascar, but has also been recorded in humid areas in the far southeast and northeast, and is found up to 1300 m (4300 ft) above sea level. It mainly occurs in forests, but has also been recorded outside forest and is not dependent on it. An obligate cave dweller, it is known to roost in large colonies, with one cave colony estimated to contain over 40,000 bats; this cave also contained an estimated 10,000 Paratriaenops furculus. The effective population sizeEffective population size
In population genetics, the concept of effective population size Ne was introduced by the American geneticist Sewall Wright, who wrote two landmark papers on it...
of the species is estimated to be about 121,000. Lepidoptera
Lepidoptera
Lepidoptera is a large order of insects that includes moths and butterflies . It is one of the most widespread and widely recognizable insect orders in the world, encompassing moths and the three superfamilies of butterflies, skipper butterflies, and moth-butterflies...
(butterflies and moths) form the main component of its diet, but it also eats Coleoptera (beetles) and Hemiptera
Hemiptera
Hemiptera is an order of insects most often known as the true bugs , comprising around 50,000–80,000 species of cicadas, aphids, planthoppers, leafhoppers, shield bugs, and others...
(bugs) and fewer members of some other insect orders. It is unclear how the ecological niche
Ecological niche
In ecology, a niche is a term describing the relational position of a species or population in its ecosystem to each other; e.g. a dolphin could potentially be in another ecological niche from one that travels in a different pod if the members of these pods utilize significantly different food...
s of Triaenops and the less frequently captured Paratriaenops furculus are separated, as both eat lepidopterans and occur in the same regions.
Conservation status
Under Triaenops rufus, the IUCN Red ListIUCN Red List
The IUCN Red List of Threatened Species , founded in 1963, is the world's most comprehensive inventory of the global conservation status of biological species. The International Union for Conservation of Nature is the world's main authority on the conservation status of species...
lists this species as "Least Concern
Least Concern
Least Concern is an IUCN category assigned to extant taxon or lower taxa which have been evaluated but do not qualify for any other category. As such they do not qualify as threatened, Near Threatened, or Conservation Dependent...
", citing its common occurrence over a wide distribution and tolerance of human modifications of its habitat, even though its forest habitat is being destroyed in places. It has been recorded in numerous protected area
Protected area
Protected areas are locations which receive protection because of their recognised natural, ecological and/or cultural values. There are several kinds of protected areas, which vary by level of protection depending on the enabling laws of each country or the regulations of the international...
s. Bats, mainly the large Hipposideros commersoni, are sometimes hunted for food in southwestern Madagascar, and T. menamena is also taken incidentally.
Literature cited
- Andriafidison, D., Cardiff, S.G., Goodman, S.M., Hutson, A.M., Jenkins, R.K.B., Kofoky, A.F., Racey, P.A., Ranivo, J., Ratrimomanarivo, F.H. and Razafimanahaka, H.J. 2008. . In IUCN. IUCN Red List of Threatened Species. Version 2009.2. <www.iucnredlist.org>. Downloaded on May 29, 2010.
- Benda, P. and Vallo, P. 2009. Taxonomic revision of the genus Triaenops (Chiroptera: Hipposideridae) with description of a new species from southern Arabia and definitions of a new genus and tribe. Folia Zoologica 58(Monograph 1):1–45.
- Garbutt, N. 2007. Mammals of Madagascar: A Complete Guide. A & C Black, 304 pp. ISBN 978-0-7136-7043-1
- Goodman, S.M. 2006. Hunting of Microchiroptera in south-western Madagascar (subscription required). Oryx 40(2):225–228.
- Goodman S.M. and Ranivo, J. 2008. A new species of Triaenops (Mammalia, Chiroptera, Hipposideridae) from Aldabra Atoll, Picard Island (Seychelles). Zoosystema 30(3):681–693.
- Goodman, S.M. and Ranivo, J. 2009. The geographical origin of the type specimens of Triaenops rufus and T. humbloti (Chiroptera: Hipposideridae) reputed to be from Madagascar and the description of a replacement species name (subscription required). Mammalia 73:47–55.
- Kofoky, A.F., Andriafidison, D., Ratrimomanarivo, F.H., Razafimanahaka, H.J., Rakotondravony, D., Racey, P.A. and Jenkins, R.K.B. 2007. Habitat use, roost selection and conservation of bats in Tsingy de Bemaraha National Park, Madagascar. Biodiversity and Conservation 16:1039–1053.
- Kofoky, A.F., Randrianandrianina, F., Russ, J., Raharinantenaina, I., Cardiff, S.G., Jenkins, R.K.B. and Racey, P.A. 2009. Forest bats of Madagascar: results of acoustic surveys (subscription required). Acta Chiropterologica 11(2):375–392.
- Milne-Edwards, A. 1881. Observations sur quelques animaux de Madagascar. Comptes rendus hebdomadaires des séances de l'Académie des sciences, Paris 91:1034–1038 (in French).
- Olsson, A., Emmett, D., Henson, D. and Fanning, E. 2006. Activity patterns and abundance of microchiropteran bats at a cave roost in southwest Madagascar (subscription required). African Journal of Ecology 44:401–403.
- Peterson, R.L., Eger, J.L. and Mitchell, L. 1995. Chiroptères. Faune de Madagascar 84:1–204 (in French).
- Racey, P.A., Goodman, S.M. and Jenkins, R.K.B. 2010. The ecology and conservation of Malagasy bats. Pp. 369–404 in Fleming, T.H. and Racey, P.A. (eds.). Island Bats: Evolution, Ecology, and Conservation. University of Chicago Press, 549 pp. ISBN 9780226253305
- Rakotoarivelo, A.A., Ranaivoson, N., Ramilijaona, O.R., Kofoky, A.F., Racey, P.A. and Jenkins, R.K.B. 2007. Seasonal food habits of five sympatric forest microchiropterans in western Madagascar (subscription required). Journal of Mammalogy 88(4):959–966.
- Ranivo, J. and Goodman S.M. 2006. Révision taxinomique des Triaenops malgaches (Mammalia, Chiroptera, Hipposideridae). Zoosystema 28(4):963–985 (in French).
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