Manavi Long-fingered Bat
Encyclopedia
Miniopterus manavi is a bat
in the genus Miniopterus
that occurs in east-central Madagascar
. First described in 1906, this species was later included in the mainland African M. minor. A 1995 revision united populations of small Miniopterus from Madagascar and the Comoros
as M. manavi, but molecular and morphological studies in 2008 and 2009 showed that this concept of M. manavi in fact included five different species. M. manavi itself was restricted to a few locations in the eastern Central Highlands
and populations in the Comoros and northern and western Madagascar were allocated to different species.
Miniopterus manavi is a small, blackish or reddish-brown Miniopterus; its forearm length is 37.6 to 39.2 mm (1.48 to 1.54 in). The tragus (a projection in the outer ear) is narrow and ends in an angular tip. The uropatagium
(tail membrane) is well-furred and the palate
is flat.
, a widespread genus of bats in Africa, southern Eurasia, and Australia, was first recorded from Madagascar by George Edward Dobson
, who mentioned the larger Miniopterus schreibersii and the smaller M. scotinus (currently M. natalensis) in his 1878 catalog of the bats in the British Museum. In 1906, Oldfield Thomas
named the larger species M. majori
and the smaller M. manavi. He regarded M. manavi as close to the mainland African M. minor, and in 1971, R.W. Hayman and J.E. Hill placed it as a subspecies
of that species. In their 1995 Faune de Madagascar review of Malagasy bats, however, Randolph Peterson and colleagues again separated M. manavi as a species, with M. manavi griveaudi (currently Miniopterus griveaudi
) from Grande Comore
as a subspecies. Peterson, who died before the review was completed, had originally divided M. manavi into several species occurring in different areas, but his collaborators decided conservatively to keep M. manavi as a single species, recommending reassessment of the status of those forms as new material would become available.
In the 2000s, molecular studies helped clarify the systematics of Miniopterus. In 2007, Javier Juste and colleagues, using sequences of the mitochondrial
cytochrome b
gene, found that bats from Madagascar (M. manavi), Grande Comore (M. manavi griveaudi) and São Tomé
(M. minor newtoni; currently Miniopterus newtoni) did not cluster together to the exclusion of other African Miniopterus; however, their samples of "M. manavi" were in fact misidentified M. majori
. The next year, Nicole Weyeneth and colleagues used cytochrome b and mitochondrial D-loop
sequences to assess the relationships of Comoran Miniopterus. They found two unrelated clade
s within Malagasy and Comoran samples of "Miniopterus manavi", neither of which was closely related to M. newtoni or to Tanzanian samples of M. minor.
During 2009, Steven Goodman
and colleagues published two papers that found a total of five genetically and morphologically distinct species within Miniopterus manavi as defined by Peterson and colleagues (1995), up to four of which can be found in a single locality. In order to determine the true identity of M. manavi, Goodman and Claude Maminirina obtained bats near the type locality of M. manavi (the site where the original material was collected, from which the species was described) for inclusion in the analysis; they also sequenced one of Thomas's original specimens. Among the five species they identified, M. griveaudi occurs on Grande Comore and Anjouan
and in northern and western Madagascar; M. aelleni
occurs on Anjouan and in northern and western Madagascar; M. brachytragos
is found in northern Madagascar only; M. mahafaliensis
is confined to the southwestern part of the island; and M. manavi itself is known only from the eastern edge of the Central Highlands
. These five species are not each others' closest relatives according to analyses of cytochrome b sequences and their similarities reflect convergent evolution
. Cytochrome b suggested that the closest relative of M. manavi is the slightly larger M. petersoni
from southeastern Madagascar. Two specimens of M. manavi differed by 1.3% in their cytochrome b sequences and by 2.5% from M. petersoni.
In the single specimen of true M. manavi that Goodman and colleagues could measure, total length is 90 mm (3.5 in), tail length is 39 mm (1.5 in), hindfoot length is 6 mm (0.2 in), tragus length is 6 mm (0.2 in), ear length is 10 mm (0.4 in), and body mass is 6.4 g (0.2 oz). The length of the forearm is known from four specimens; it ranges from 37.6 to 39.2 mm (1.48 to 1.54 in), averaging 38.5 mm (1.52 in).
In the skull, the rostrum (front part) is rounded. The central groove in the nasal depression is relatively narrow. The frontal bone
s are inflated and bear a prominent sagittal crest
. Further back on the braincase, the lambdoid crest is poorly developed. The middle part of the palate
is flat, not concave as in M. brachytragos, M. griveaudi, and M. mahafaliensis. At the palate's back margin is a short, thick posterior palatal spine.
, from the vicinity of Ambositra
in the north to Vinanitelo in the south, at 900 to 1500 m (3000 to 4900 ft) above sea level. The 2008 IUCN Red List
assesses the species as "Least Concern", citing its wide distribution, though it is sometimes hunted for food. However, the account predates the recognition of M. aelleni, M. brachytragos, M. griveaudi, and M. mahafaliensis as separate species. Although some ecological data about M. manavi have been published, these need to be reevaluated with the recognition of numerous additional species within M. manavi. Species of Miniopterus generally feed on insects, breed seasonally, and roost in large colonies in caves. The myobiid mite Calcarmyobia comoresensis has been recorded on M. manavi.
Bat
Bats are mammals of the order Chiroptera "hand" and pteron "wing") whose forelimbs form webbed wings, making them the only mammals naturally capable of true and sustained flight. By contrast, other mammals said to fly, such as flying squirrels, gliding possums, and colugos, glide rather than fly,...
in the genus Miniopterus
Miniopterus
Miniopterus is a genus of bats and the only genus of the family Miniopteridae. The genus was previously placed in its own subfamily, Miniopterinae, of the vesper bat family, but is now classified in its own family....
that occurs in east-central Madagascar
Madagascar
The Republic of Madagascar is an island country located in the Indian Ocean off the southeastern coast of Africa...
. First described in 1906, this species was later included in the mainland African M. minor. A 1995 revision united populations of small Miniopterus from Madagascar and the Comoros
Comoros
The Comoros , officially the Union of the Comoros is an archipelago island nation in the Indian Ocean, located off the eastern coast of Africa, on the northern end of the Mozambique Channel, between northeastern Mozambique and northwestern Madagascar...
as M. manavi, but molecular and morphological studies in 2008 and 2009 showed that this concept of M. manavi in fact included five different species. M. manavi itself was restricted to a few locations in the eastern Central Highlands
Central Highlands (Madagascar)
The Central Highlands, Central High Plateau, or Hauts-Plateaux are a mountainous biogeographical region in central Madagascar. They include the contiguous part of the island's interior above 800 m altitude...
and populations in the Comoros and northern and western Madagascar were allocated to different species.
Miniopterus manavi is a small, blackish or reddish-brown Miniopterus; its forearm length is 37.6 to 39.2 mm (1.48 to 1.54 in). The tragus (a projection in the outer ear) is narrow and ends in an angular tip. The uropatagium
Patagium
*In bats, the skin forming the surface of the wing. It is an extension of the skin of the abdomen that runs to the tip of each digit, uniting the forelimb with the body.*The patagium of a bat has four distinct parts:...
(tail membrane) is well-furred and the palate
Palate
The palate is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but, in most other tetrapods, the oral and nasal cavities are not truly separate. The palate is divided into two parts, the anterior...
is flat.
Taxonomy
MiniopterusMiniopterus
Miniopterus is a genus of bats and the only genus of the family Miniopteridae. The genus was previously placed in its own subfamily, Miniopterinae, of the vesper bat family, but is now classified in its own family....
, a widespread genus of bats in Africa, southern Eurasia, and Australia, was first recorded from Madagascar by George Edward Dobson
George Edward Dobson
George Edward Dobson FRS was a zoologist, photographer and army surgeon.-Biography:...
, who mentioned the larger Miniopterus schreibersii and the smaller M. scotinus (currently M. natalensis) in his 1878 catalog of the bats in the British Museum. In 1906, Oldfield Thomas
Oldfield Thomas
Oldfield Thomas FRS was a British zoologist.Thomas worked at the Natural History Museum on mammals, describing about 2,000 new species and sub-species for the first time. He was appointed to the Museum Secretary's office in 1876, transferring to the Zoological Department in 1878...
named the larger species M. majori
Miniopterus majori
Miniopterus majori is a species of vesper bat in the family Vespertilionidae.It is found only in Madagascar. It is similar to M. schreibersi of Africa, differing by having a shorter forearm, slightly longer digits and a narrow box-shaped skull. The pelage is often a greyish brown colour, and the...
and the smaller M. manavi. He regarded M. manavi as close to the mainland African M. minor, and in 1971, R.W. Hayman and J.E. Hill placed it 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 that species. In their 1995 Faune de Madagascar review of Malagasy bats, however, Randolph Peterson and colleagues again separated M. manavi as a species, with M. manavi griveaudi (currently Miniopterus griveaudi
Miniopterus griveaudi
Miniopterus griveaudi is a bat in the genus Miniopterus found on Grande Comore and Anjouan in the Comoros and in northern and western Madagascar. First described in 1959 from Grande Comore as a subspecies of the mainland African M. minor, it was later placed with the Malagasy M. manavi...
) from Grande Comore
Grande Comore
Grande Comore is an island in the Indian Ocean off the coast of Africa. It is the largest island in the Comoros nation. Most of its population is of the Comorian ethnic group. Its population as of 2006 is about 316,600. The island's capital is Moroni, which is also the national capital...
as a subspecies. Peterson, who died before the review was completed, had originally divided M. manavi into several species occurring in different areas, but his collaborators decided conservatively to keep M. manavi as a single species, recommending reassessment of the status of those forms as new material would become available.
In the 2000s, molecular studies helped clarify the systematics of Miniopterus. In 2007, Javier Juste and colleagues, using sequences of the mitochondrial
Mitochondrial DNA
Mitochondrial DNA is the DNA located in organelles called mitochondria, structures within eukaryotic cells that convert the chemical energy from food into a form that cells can use, adenosine triphosphate...
cytochrome b
Cytochrome b
Cytochrome b/b6 is the main subunit of transmembrane cytochrome bc1 and b6f complexes. In addition, it commonly refers to a region of mtDNA used for population genetics and phylogenetics.- Function :...
gene, found that bats from Madagascar (M. manavi), Grande Comore (M. manavi griveaudi) and São Tomé
São Tomé
-Transport:São Tomé is served by São Tomé International Airport with regular flights to Europe and other African Countries.-Climate:São Tomé features a tropical wet and dry climate with a relatively lengthy wet season and a short dry season. The wet season runs from October through May while the...
(M. minor newtoni; currently Miniopterus newtoni) did not cluster together to the exclusion of other African Miniopterus; however, their samples of "M. manavi" were in fact misidentified M. majori
Miniopterus majori
Miniopterus majori is a species of vesper bat in the family Vespertilionidae.It is found only in Madagascar. It is similar to M. schreibersi of Africa, differing by having a shorter forearm, slightly longer digits and a narrow box-shaped skull. The pelage is often a greyish brown colour, and the...
. The next year, Nicole Weyeneth and colleagues used cytochrome b and mitochondrial D-loop
D-loop
In molecular biology, a displacement loop or D-loop is a DNA structure where the two strands of a double-stranded DNA molecule are separated for a stretch and held apart by a third strand of DNA. The third strand has a base sequence which is complementary to one of the main strands and pairs with...
sequences to assess the relationships of Comoran Miniopterus. They found two unrelated clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...
s within Malagasy and Comoran samples of "Miniopterus manavi", neither of which was closely related to M. newtoni or to Tanzanian samples of M. minor.
During 2009, 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....
and colleagues published two papers that found a total of five genetically and morphologically distinct species within Miniopterus manavi as defined by Peterson and colleagues (1995), up to four of which can be found in a single locality. In order to determine the true identity of M. manavi, Goodman and Claude Maminirina obtained bats near the type locality of M. manavi (the site where the original material was collected, from which the species was described) for inclusion in the analysis; they also sequenced one of Thomas's original specimens. Among the five species they identified, M. griveaudi occurs on Grande Comore and Anjouan
Anjouan
Anjouan is an autonomous island, part of the Union of Comoros. The island is located in the Indian Ocean. Its capital is Mutsamudu and its population as of 2006 is about 277,500. The total area of the island is 424 sq. kilometers Anjouan (also known as Ndzuwani or Nzwani) is an autonomous island,...
and in northern and western Madagascar; M. aelleni
Miniopterus aelleni
Miniopterus aelleni is a bat in the genus Miniopterus that occurs on Anjouan in the Comoros and in northern and western Madagascar. It is a small, brown bat; its forearm length is 35 to 41 mm . The long tragus has a broad base and a blunt to rounded tip. The uropatagium is sparsely haired...
occurs on Anjouan and in northern and western Madagascar; M. brachytragos
Miniopterus brachytragos
Miniopterus brachytragos is a bat in the genus Miniopterus that occurs in northern and western Madagascar. Populations of this species have historically been included in Miniopterus manavi, but molecular data published in 2008 and 2009 indicate this supposed species in fact consists of five...
is found in northern Madagascar only; M. mahafaliensis
Miniopterus mahafaliensis
Miniopterus mahafaliensis is a bat in the genus Miniopterus that occurs in southwestern Madagascar. Populations of this species have historically been included in Miniopterus manavi, but molecular data published in 2008 and 2009 indicate that this supposed species in fact consists of five separate...
is confined to the southwestern part of the island; and M. manavi itself is known only from the eastern edge of the Central Highlands
Central Highlands (Madagascar)
The Central Highlands, Central High Plateau, or Hauts-Plateaux are a mountainous biogeographical region in central Madagascar. They include the contiguous part of the island's interior above 800 m altitude...
. These five species are not each others' closest relatives according to analyses of cytochrome b sequences and their similarities reflect convergent evolution
Convergent evolution
Convergent evolution describes the acquisition of the same biological trait in unrelated lineages.The wing is a classic example of convergent evolution in action. Although their last common ancestor did not have wings, both birds and bats do, and are capable of powered flight. The wings are...
. Cytochrome b suggested that the closest relative of M. manavi is the slightly larger M. petersoni
Miniopterus petersoni
Miniopterus petersoni is a bat in the genus Miniopterus which occurs in southeast Madagascar. It was described by Steven M. Goodman et al. in 2008. While M. petersoni is similar to M. sororculus, the two species are not closely related to each other, and possess a number of differing external and...
from southeastern Madagascar. Two specimens of M. manavi differed by 1.3% in their cytochrome b sequences and by 2.5% from M. petersoni.
Description
Miniopterus manavi is a diminutive species with fur of medium length. The upperparts are blackish or reddish brown. Other small Malagasy Miniopterus are lighter. The ears mostly lack hair and end in a rounded tip. The tragus (a projection on the inner side of the outer ear) is thin for most of its length, ends in an angular tip, and has a flange at the medial side (towards the midline of the animal). The tragus is differently shaped in other species. The wings and uropatagium (tail membrane) are blackish and are attached to the upper leg at the same level, above the ankle. The uropatagium is densely haired above and more sparsely below, as in M. mahafaliensis and M. brachytragos; M. griveaudi and M. aelleni have more nearly naked uropatagia.In the single specimen of true M. manavi that Goodman and colleagues could measure, total length is 90 mm (3.5 in), tail length is 39 mm (1.5 in), hindfoot length is 6 mm (0.2 in), tragus length is 6 mm (0.2 in), ear length is 10 mm (0.4 in), and body mass is 6.4 g (0.2 oz). The length of the forearm is known from four specimens; it ranges from 37.6 to 39.2 mm (1.48 to 1.54 in), averaging 38.5 mm (1.52 in).
In the skull, the rostrum (front part) is rounded. The central groove in the nasal depression is relatively narrow. The frontal bone
Frontal bone
The frontal bone is a bone in the human skull that resembles a cockleshell in form, and consists of two portions:* a vertical portion, the squama frontalis, corresponding with the region of the forehead....
s are inflated and bear a prominent 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....
. Further back on the braincase, the lambdoid crest is poorly developed. The middle part of the palate
Palate
The palate is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but, in most other tetrapods, the oral and nasal cavities are not truly separate. The palate is divided into two parts, the anterior...
is flat, not concave as in M. brachytragos, M. griveaudi, and M. mahafaliensis. At the palate's back margin is a short, thick posterior palatal spine.
Distribution and ecology
The currently known distribution of M. manavi extends around the eastern margin of the Central HighlandsCentral Highlands (Madagascar)
The Central Highlands, Central High Plateau, or Hauts-Plateaux are a mountainous biogeographical region in central Madagascar. They include the contiguous part of the island's interior above 800 m altitude...
, from the vicinity of Ambositra
Ambositra
Ambositra is a city in central Madagascar.Ambositra is the capital of the Amoron'i Mania region, and of Ambositra District....
in the north to Vinanitelo in the south, at 900 to 1500 m (3000 to 4900 ft) above sea level. The 2008 IUCN Red List
IUCN 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...
assesses the species as "Least Concern", citing its wide distribution, though it is sometimes hunted for food. However, the account predates the recognition of M. aelleni, M. brachytragos, M. griveaudi, and M. mahafaliensis as separate species. Although some ecological data about M. manavi have been published, these need to be reevaluated with the recognition of numerous additional species within M. manavi. Species of Miniopterus generally feed on insects, breed seasonally, and roost in large colonies in caves. The myobiid mite Calcarmyobia comoresensis has been recorded on M. manavi.
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 July 13, 2010.
- Dobson, G.E. 1878. Catalogue of the Chiroptera in the collections of the British Museum. London: British Museum,
- Goodman, S.M., Maminirina, C.P., Weyeneth, N., Bradman, H.M., Christidis, L., Ruedi, M. and Appleton, B. 2009a. The use of molecular and morphological characters to resolve the taxonomic identity of cryptic species: the case of Miniopterus manavi (Chiroptera: Miniopteridae) (subscription required). Zoologica Scripta 38:339–363.
- Goodman, S.M., Maminirina, C.P., Bradman, H.M., Christidis, L. and Appleton, B. 2009b. The use of molecular phylogenetic and morphological tools to identify cryptic and paraphyletic species: Examples from the diminutive long-fingered bats (Chiroptera: Miniopteridae: Miniopterus) on Madagascar. American Museum Novitates 3669:1–34.
- Hill, J.E. 1993. Long-fingered bats of the genus Miniopterus (Chiroptera: Vespertilionidae) from Madagascar. Mammalia 57(3):401–405.
- Juste, J., Ferrández, A., Fa, J.E., Masefield, W. and Ibáñez, C. 2007. Taxonomy of little bentwinged bats (Miniopterus, Miniopteridae) from the African islands of São Tomé, Grand Comoro and Madagascar, based on mtDNA. Acta Chiropterologica 9:27–37.
- Nowak, R.M. 1994. Walker's Bats of the Old World. Baltimore: The Johns Hopkins University Press, 287 pp. ISBN 9780801849862
- Thomas, O. 1906. New African mammals of the genera Cercopithecus, Scotophilus, Miniopterus, Crocidura, Georychus, and Heliophobius. Annals and Magazine of Natural History (7)17:173–179.
- Uchikawa, K. 1985. Calcarmyobia from the Ethiopian region (Acarina, Myobiidae). Bulletin of the British Museum of Natural History (Zoology) 48(1):45–55.
- Weyeneth, N., Goodman, S.M., Stanley, W.T. and Ruedi, M. 2008. The biogeography of Miniopterus bats (Chiroptera: Miniopteridae) from the Comoro Archipelago inferred from mitochondrial DNA (subscription required). Molecular Ecology 17:5205–5219.