Miniopterus Brachytragos
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''Miniopterus brachytragos'' is a
bat 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 ...
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, officially the Republic of Madagascar, is an island country that includes the island of Madagascar and numerous smaller peripheral islands. Lying off the southeastern coast of Africa, it is the world's List of islands by area, f ...
. Populations of this species have historically been included in ''Miniopterus manavi'', but
molecular A molecule is a group of two or more atoms that are held together by attractive forces known as chemical bonds; depending on context, the term may or may not include ions that satisfy this criterion. In quantum physics, organic chemistry, ...
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 (: patagia) is a membranous body part that assists an animal in obtaining lift when gliding or flying. The structure is found in extant and extinct groups of flying and gliding animals including bats, theropod dinosaurs (including b ...
(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 sep ...
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, is an archipelagic country made up of three islands in Southeastern Africa, located at the northern end of the Mozambique Channel in the Indian Ocean. Its capital and largest city is Moroni, ...
, including the small '' Miniopterus manavi'' 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 comm ...
.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 Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
combines the
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
''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 used ...
cytochrome ''b'' gene suggested that ''M. brachytragos'' is most closely related to the
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
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 (; ), also known as Nosy Ambariovato, is a small island in Madagascar, situated between the island of Nosy Be and the northwestern coast of the main island of Madagascar. Roughly circular, it rises sharply towards a plateau and the su ...
, 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 (: patagia) is a membranous body part that assists an animal in obtaining lift when gliding or flying. The structure is found in extant and extinct groups of flying and gliding animals including bats, theropod dinosaurs (including b ...
(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 In the human skull, the frontal bone or sincipital bone is an unpaired bone which consists of two portions.'' Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bo ...
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 excepti ...
. 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 sep ...
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 (; : foramina, or foramens ; ) is an opening or enclosed gap within the dense connective tissue (bones and deep fasciae) of extant and extinct amniote animals, typically to allow passage of nerves, arter ...
(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-physiology ...
(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, wher ...
s, one
canine Canine may refer to: Zoology and anatomy * Animals of the family Canidae, more specifically the subfamily Caninae, which includes dogs, wolves, foxes, jackals and coyotes ** ''Canis'', a genus that includes dogs, wolves, coyotes, and jackals ** Do ...
, three
premolar The premolars, also called premolar Tooth (human), teeth, or bicuspids, are transitional teeth located between the Canine tooth, canine and Molar (tooth), molar teeth. In humans, there are two premolars per dental terminology#Quadrant, quadrant in ...
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 natural science of the relationships among living organisms and their environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere levels. Ecology overlaps with the closely re ...
different sites. In the Namoroka region, it has been recorded at altitude in
dry forest The tropical and subtropical dry broadleaf forest is a habitat type defined by the World Wide Fund for Nature and is located at tropical and subtropical latitudes. Though these forests occur in climates that are warm year-round, and may receive ...
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 carbonate rocks such as limestone and Dolomite (rock), dolomite. It is characterized by features like poljes above and drainage systems with sinkholes and caves underground. Ther ...
region in Bemaraha. Near Daraina 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 an evergreen species of flowering plant in the family Anacardiaceae. It is a large fruit tree, capable of growing to a height and width of . There are two distinct genetic populations in modern m ...
'' (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