Miniopterus Brachytragos
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''Miniopterus brachytragos'' is a
bat Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". 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 bi ...
in the genus ''
Miniopterus ''Miniopterus'', known as the bent-winged or long winged bats, is the sole genus of the family Miniopteridae. They are small flying insectivorous mammals, micro-bats of the order Chiroptera, with wings over twice the length of the body. The genus ...
'' that occurs in northern and western
Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Africa ...
. Populations of this species have historically been included in ''Miniopterus manavi'', but
molecular A molecule is a group of two or more atoms held together by attractive forces known as chemical bonds; depending on context, the term may or may not include ions which satisfy this criterion. In quantum physics, organic chemistry, and bioche ...
data published in 2008 and 2009 indicate this supposed species in fact consists of five separate species, including the newly described ''M. brachytragos''. Up to four species of this group may occur in the same place. ''M. brachytragos'' has been found in dry and wet forests from sea level to altitude. ''Miniopterus brachytragos'' is a small, brown ''Miniopterus''; its forearm length is . The hairs of the underparts have buff tips. The short tragus (a projection in the outer ear) is an important distinguishing feature. The
uropatagium The patagium (plural: patagia) is a membranous body part that assists an animal in obtaining lift when gliding or flight. The structure is found in extant and extinct groups of flying and gliding animals including bats, birds, some dromaeosaurs, ...
(tail membrane) is well-furred and the
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 sepa ...
is concave.


Taxonomy

During the 2000s, molecular studies have revealed that the widely distributed African, Eurasian, and Australian genus ''
Miniopterus ''Miniopterus'', known as the bent-winged or long winged bats, is the sole genus of the family Miniopteridae. They are small flying insectivorous mammals, micro-bats of the order Chiroptera, with wings over twice the length of the body. The genus ...
'' is much more species-rich than previously thought. In a 1995 contribution to '' Faune de Madagascar'' on Malagasy bats, Randolph Peterson and colleagues listed four species of ''Miniopterus'' on Madagascar and the nearby
Comoros The Comoros,, ' officially the Union of the Comoros,; ar, الاتحاد القمري ' is an independent country made up of three islands in southeastern Africa, located at the northern end of the Mozambique Channel in the Indian Ocean. It ...
, including the small ''
Miniopterus manavi The Manavi long-fingered bat (''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 revisi ...
'' with a broad distribution on both Madagascar and the Comoros.Goodman et al., 2009a, p. 340 In 2008 and 2009, however, Steven Goodman and colleagues presented evidence that the former concept of ''M. manavi'' in fact encompassed five morphologically and molecularly distinct species of small ''Miniopterus''. These included ''M. manavi'' itself in the Central Highlands, '' M. griveaudi'' and '' M. aelleni'' in the Comoros and northern and western Madagascar, '' M. mahafaliensis'' in southwestern Madagascar, and ''M. brachytragos'' in northern and western Madagascar. The five recognized species of ''M. manavi''-like bats are not each other's closest relatives, but apparently acquired their similarities through
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
.Goodman et al., 2009b, p. 28 At some places (for example, Namoroka) four
cryptic species In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...
of ''M. manavi''-like bats, including ''M. brachytragos'', may occur together. ''Miniopterus brachytragos'' was described as a new species in the second 2009 paper by Goodman and colleagues.Goodman et al., 2009b, p. 9 The specific name combines the
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic peri ...
''brachys'' "short" and ''tragos'' "goat" and refers to the short tragus (a fleshy projection in the inner side of the outer ear), one of the main distinguishing features of the species; the name of this structure derives from the Greek ''tragos''.Goodman et al., 2009b, p. 17 Analysis of sequences of the
mitochondrial A mitochondrion (; ) is an organelle found in the cells of most Eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is use ...
cytochrome ''b'' gene suggested that ''M. brachytragos'' is most closely related to the
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
of ''M. manavi'' and another Malagasy species, '' M. petersoni''. There was some variation within the species—the most distinctive individual, from the island of
Nosy Komba Nosy Komba is a volcanic island off the north west coast of Madagascar, lying between Nosy Be Nosy Be (formerly Nossi-bé and Nosse Be) is an island off the northwest coast of Madagascar. Nosy Be is Madagascar's largest and busiest tourist re ...
, differed by about 2.1% from other individuals of the same species in its cytochrome ''b'' sequence—but Goodman and colleagues regarded their samples as insufficient for clear conclusions about
phylogeographic Phylogeography is the study of the historical processes that may be responsible for the past to present geographic distributions of genealogical lineages. This is accomplished by considering the geographic distribution of individuals in light of ge ...
structure within the species.Goodman et al., 2009b, p. 16


Description

''Miniopterus brachytragos'' is a small, short-tailed ''Miniopterus'' with short and relatively thin fur. The fur of the upperparts is dark brown and the hairs on the underparts have buff tips.Goodman et al., 2009b, p. 11 ''Miniopterus mahafaliensis'' is similar in color, but other small Malagasy ''Miniopterus'' are darker. The ears are haired above, but virtually naked below and end in a rounded tip. The tragus (a projection on the inner side of the outer ear) is short and has a broad base, sometimes with a crest at the side. It ends in a pointed to slightly rounded tip, which is covered with hairs that are not readily visible to the unaided eye.Goodman et al., 2009b, p. 15 The short tragus easily distinguishes ''M. brachytragos'' from other Malagasy ''Miniopterus''. The wing membrane is also brown, but the
uropatagium The patagium (plural: patagia) is a membranous body part that assists an animal in obtaining lift when gliding or flight. The structure is found in extant and extinct groups of flying and gliding animals including bats, birds, some dromaeosaurs, ...
(tail membrane) is lighter. The wing membrane and uropatagium are attached to the upper leg at the same level, above the ankle. The uropatagium is relatively densely covered with hairs, particularly on the upper side. ''M. manavi'' and ''M. mahafaliensis'' also have a densely haired uropatagium, but in ''M. aelleni'' and ''M. griveaudi'' it is only sparsely haired or even mostly naked.Goodman et al., 2009b, p. 22 In 28 to 30 specimens measured by Goodman and colleagues, total length was , averaging ; tail length was , averaging ; hindfoot length was 5 to 6 mm (about 0.2 in), averaging ; tragus length was , averaging ; ear length was 9 to 11 mm (about 0.4 in), averaging ; forearm length was , averaging ; and body mass was , averaging . There is no evidence for substantial size differences between males and females. In the skull, the rostrum (front part) is short relative to other small Malagasy ''Miniopterus'' and line-shaped. The central groove in the nasal depression is relatively broad and deep. The
frontal bone The frontal bone is a bone in the human skull. The bone consists of two portions.''Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bony part of the forehead, par ...
s are slightly rounded and bear an indistinct
sagittal crest A sagittal crest is a ridge of bone running lengthwise along the midline of the top of the skull (at the sagittal suture) of many mammalian and reptilian skulls, among others. The presence of this ridge of bone indicates that there are exceptiona ...
. Further back on the braincase, the lambdoid crest is also poorly developed. The middle part of the
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 sepa ...
is concave, not flat as in ''M. aelleni'' and ''M. manavi''. At the palate's back margin is a short, blunt posterior palatal spine. There are often
foramina In anatomy and osteology, a foramen (;Entry "foramen"
in
(openings) in the palate near the last molar. ''Miniopterus brachytragos'' has 36 teeth in the
dental formula Dentition pertains to the development of teeth and their arrangement in the mouth. In particular, it is the characteristic arrangement, kind, and number of teeth in a given species at a given age. That is, the number, type, and morpho-physiolo ...
(three
incisor Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, whe ...
s, one canine, three
premolar The premolars, also called premolar teeth, or bicuspids, are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant in the permanent set of teeth, making eight premolars total in the mouth ...
s, and two molars in both upper toothrows and two incisors, one canine, two premolars, and three molars in the lower toothrows). As is characteristic of ''Miniopterus'', the first upper premolar (P1) is smaller and more simplified than the second (P2).


Distribution and ecology

''Miniopterus brachytragos'' is known from several
ecologically Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overlaps wi ...
different sites. In the Namoroka region, it has been recorded at altitude in
dry forest Dry or dryness most often refers to: * Lack of rainfall, which may refer to **Arid regions **Drought * Dry or dry area, relating to legal prohibition of selling, serving, or imbibing alcoholic beverages * Dry humor, deadpan * Dryness (medical) ...
and
gallery forest A gallery forest is one formed as a corridor along rivers or wetlands, projecting into landscapes that are otherwise only sparsely treed such as savannas, grasslands, or deserts. The gallery forest maintains a more temperate microclimate above th ...
. Another specimen was caught in forest in a
karst Karst is a topography formed from the dissolution of soluble rocks such as limestone, dolomite, and gypsum. It is characterized by underground drainage systems with sinkholes and caves. It has also been documented for more weathering-resistant ro ...
region in Bemaraha. Near
Daraina Daraina is a town and commune ( mg, kaominina) in northern Madagascar. It belongs to the district of Vohemar, which is a part of Sava Region. The city is located at the unpaved part of the Route Nationale 5a between Vohemar and Ambilobe. The pop ...
in the northeast, the species occurs in eastern humid forest mixed with dry forest at . At another northeastern site, on the
Masoala Peninsula Masoala National Park, in northeast Madagascar, is the largest of the island's protected areas. Most of the park is situated in Sava Region and a part in Analanjirofo. Created in 1997, the park protects 2,300 square kilometres of rainforest and ...
, it was found in lowland gallery forest near sea level. On Nosy Komba, its habitat consists of dry forest mixed with introduced ''
Mangifera indica ''Mangifera indica'', commonly known as mango, is a species of flowering plant in the family Anacardiaceae. It is a large fruit tree, capable of growing to a height of . There are two distinct genetic populations in modern mangoesthe "Indian type ...
'' (mango tree). Little is known of the ecology of ''M. brachytragos'', but species of ''Miniopterus'' generally feed on insects, breed seasonally, and roost in large colonies in caves.Nowak, 1994, p. 222


References


Literature cited

*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. *Nowak, R.M. 1994. Walker's Bats of the World. Baltimore: The Johns Hopkins University Press, 287 pp. *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. {{Taxonbar, from=Q11936833 Miniopteridae Mammals described in 2009 Endemic fauna of Madagascar Mammals of Madagascar