Mantella
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''Mantella'' (also known as golden frogs or Malagasy poison frogs) are a prominent genus of
aposematic Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste o ...
frog A frog is any member of a diverse and largely carnivorous group of short-bodied, tailless amphibians composing the order Anura (ανοὐρά, literally ''without tail'' in Ancient Greek). The oldest fossil "proto-frog" ''Triadobatrachus'' is ...
s in the family
Mantellidae The Mantellidae are a family of the order Anura. These frogs are endemic to the islands of Madagascar and Mayotte. Systematics The family Mantellidae is composed of three extremely ecologically diverse groups of frogs, divided into three sub ...
,
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found else ...
to the island of
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 ...
. Members of ''Mantella'' are diurnal and
terrestrial Terrestrial refers to things related to land or the planet Earth. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on or near the ground, as opposed to ...
, with bright aposematic coloration or cryptic markings.


Natural history

''Mantella'' are an example of
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 ...
—the independent evolution of a similar trait with species of a different lineage—with the
Latin American Latin Americans ( es, Latinoamericanos; pt, Latino-americanos; ) are the citizens of Latin American countries (or people with cultural, ancestral or national origins in Latin America). Latin American countries and their diasporas are multi-eth ...
family
Dendrobatidae Poison dart frog (also known as dart-poison frog, poison frog or formerly known as poison arrow frog) is the common name of a group of frogs in the Family (biology), family Dendrobatidae which are native to tropical Central and South America. T ...
in size, appearance, and some behavioral characteristics. During the description of the first specimens from 1866 to 1872, Alfred Grandidier described both the brown mantella (''Mantella betsileo'') and Malagasy mantella (''Mantella madagascariensis'') and placed them within the genus ''
Dendrobates ''Dendrobates'' is a genus of poison dart frogs native to Central and South America. It once contained numerous species, but most originally placed in this genus have been split off into other genera such as ''Adelphobates'', '' Ameerega'', '' A ...
'' based on their close resemblance. This placement was heavily debated until 1882, when George Albert Boulenger created the genus ''Mantella'' after describing both
Cowan's mantella The Cowan's mantella (''Mantella cowanii'') is a species of frog in the family Mantellidae. It is endemic to Madagascar. Its natural habitats are subtropical or tropical moist montane forests, subtropical or tropical high-altitude grassland, and ...
(''Mantella cowanii'') and, in 1888, Baron's mantella (''Mantella baroni''). ''M. baroni'' was named after the gentleman that collected the specimens, Rev. Richard Baron, a
missionary A missionary is a member of a religious group which is sent into an area in order to promote its faith or provide services to people, such as education, literacy, social justice, health care, and economic development.Thomas Hale 'On Being a Mi ...
and botanist living in Madagascar. Baron was also interested in geology and herpetology, collecting many specimens during his extensive expeditions across the country. This species is incredibly similar in coloration to ''M. madagascariensis,'' except for the ventral/underside markings. In 1889, after the description of ''M. baroni'', French naturalist Alexandre Thominot described ''Phrynomantis maculatus'', with its type locality on
Réunion Island Réunion (; french: La Réunion, ; previously ''Île Bourbon''; rcf, label= Reunionese Creole, La Rényon) is an island in the Indian Ocean that is an overseas department and region of France. It is located approximately east of the island o ...
. However, this locality was later corrected to the off-shore Malagasy islands of Nosy Bé and
Nosy Komba Nosy Komba is a volcanic island off the north west coast of Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an isla ...
and ''P. maculatus'' was synonymized with ''M. baroni''. The genus remained within Dendrobatidae until the late 19th century. The Royal Natural History (1893) by
Richard Lydekker Richard Lydekker (; 25 July 1849 – 16 April 1915) was an English naturalist, geologist and writer of numerous books on natural history. Biography Richard Lydekker was born at Tavistock Square in London. His father was Gerard Wolfe Lydekker, ...
included the genus ''Mantella'' as one of two genera representing Dendrobatidae, saying that they could be "''distinguished by the tip of the tongue being notched; while in'' Dendrobates ''of Tropical America the tongue is entire.''" During the first quarter of the 20th century, another three species of ''Mantella'' were described, including the
golden mantella The golden mantella (''Mantella aurantiaca'') is a small, terrestrial frog endemic to Madagascar. It has an extremely restricted distribution in three distinct areas centered on the town of Moramanga - Beparasy and Ambohibary Communes, Torotorof ...
(''Mantella aurantiaca''), by the French herpetologist François Mocquard in 1900. In his work ''"Synopsis des familles, genres et espèces des reptiles écailleux et des batraciens de Madagascar''" published in 1909, Mocquard gave a detailed description of ''Mantella'' and the species within the genus. Within the document, six species are described, including one unusual description of ''Mantella attemsi'', described in 1901 by Franz Josef Maria Werner, an
Austria Austria, , bar, Östareich officially the Republic of Austria, is a country in the southern part of Central Europe, lying in the Eastern Alps. It is a federation of nine states, one of which is the capital, Vienna, the most populous ...
n
zoologist Zoology ()The pronunciation of zoology as is usually regarded as nonstandard, though it is not uncommon. is the branch of biology that studies the animal kingdom, including the structure, embryology, evolution, classification, habits, and d ...
and
explorer Exploration refers to the historical practice of discovering remote lands. It is studied by geographers and historians. Two major eras of exploration occurred in human history: one of convergence, and one of divergence. The first, covering most ...
. Mocquard's work describes ''M. attemsi'' as follows: "''First digit extends as far as the second. eplilatero-dorsalpresent, starting at the rear of the upper eyelid. Skin very porous, slightly rough on the back and the head, stomach side smooth; lower back of the legs very rough. Back a dark red-brown, rest of the body black''." This species was later synonymised with ''M. betsileo''.


Description

Species of this genus are small, varying in length between . Most ''Mantella'' species are
sexually dimorphic Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...
in size, with females being larger than males. ''Mantella'' vary in shape from streamlined to plump/rounded bodies, with skin that is either smooth or granular. They have small, angular heads, with large eyes that are either entirely dark or have lighter coloration around the edge of the iris. ''Mantella'' have a very distinct tympanum. The tips or discs of the fingers are slightly enlarged, though those of the climbing mantella (''Mantella laevigata'') are distinctly larger than in other members of the genus. They have four fingers on each forelimb and five on each hindlimb; some species have webbed digits, while others do not. The tibiotarsal articulation is roughly between the shoulder and the nostrils. Many species of ''Mantella'' are similar to the
neotropical The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropical terrestrial ecoregions of the Americas and the entire South American temperate zone. Definition In bioge ...
family
Dendrobatidae Poison dart frog (also known as dart-poison frog, poison frog or formerly known as poison arrow frog) is the common name of a group of frogs in the Family (biology), family Dendrobatidae which are native to tropical Central and South America. T ...
in their use of aposematism (from
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
ἀπό ''apo'' away, σῆμα ''sema'' sign), a defense mechanism that uses dramatic coloration to deter potential predators. Coloration and markings vary between species, with combinations of green, red, orange, yellow, blue, brown, white and black. These colorations are often evidence that the specimen produces toxic, pharmacologically active
alkaloid Alkaloids are a class of basic, naturally occurring organic compounds that contain at least one nitrogen atom. This group also includes some related compounds with neutral and even weakly acidic properties. Some synthetic compounds of similar ...
secretions. There are significant similarities between a few species of ''Mantella'' and ''Dendrobatidae'', notably the
golden mantella The golden mantella (''Mantella aurantiaca'') is a small, terrestrial frog endemic to Madagascar. It has an extremely restricted distribution in three distinct areas centered on the town of Moramanga - Beparasy and Ambohibary Communes, Torotorof ...
(''Mantella aurantiaca'') and the
golden poison frog The golden poison frog (''Phyllobates terribilis''), also known as the golden dart frog or golden poison arrow frog, is a poison dart frog endemic to the rainforests of Colombia. The golden poison frog has become endangered due to habitat destr ...
(''Phyllobates terribilis'').
Cowan's mantella The Cowan's mantella (''Mantella cowanii'') is a species of frog in the family Mantellidae. It is endemic to Madagascar. Its natural habitats are subtropical or tropical moist montane forests, subtropical or tropical high-altitude grassland, and ...
(''Mantella cowanii'') and certain variations of the Harlequin poison frog (''Oophaga histrionica'') are also very similar in coloration''.'' Most members of the genus also exhibit aposematism on the
ventral Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
region, excluding the
golden mantella The golden mantella (''Mantella aurantiaca'') is a small, terrestrial frog endemic to Madagascar. It has an extremely restricted distribution in three distinct areas centered on the town of Moramanga - Beparasy and Ambohibary Communes, Torotorof ...
and black-eared mantella (''Mantella milotympanum''). The venter is normally uniform black, dark grey, or brown and are often marked with blueish or white spots, flecks, or blotches. There are similar blueish to white markings in the form of either spots or a continuous horseshoe-shaped marking on the
vocal sac The vocal sac is the flexible membrane of skin possessed by most male frogs and toads. The purpose of the vocal sac is usually as an amplification of their mating or advertisement call. The presence or development of the vocal sac is one way ...
. These characteristics can be used to distinguish between species, such as Baron's mantella (''Mantella baroni'') and the Malagasy mantella (''Mantella madagascariensis''), when locality data is unavailable. ''Mantella'' show a variety in
alkaloid Alkaloids are a class of basic, naturally occurring organic compounds that contain at least one nitrogen atom. This group also includes some related compounds with neutral and even weakly acidic properties. Some synthetic compounds of similar ...
profiles between individual frogs of the Ranomafana region. These same alkaloids have been found to be sequestered by certain insects. It has also been observed that ''Mantella'' retain alkaloids in their skin for years in captivity. This, combined with analyses of stomach contents and diet, suggests that members of ''Mantella'' obtain at least some of their alkaloids from arthropod prey.


Distribution

''Mantella'' are endemic to the island of Madagascar and its smaller coastal islands ("Nosy" in Malagasy). They inhabit a wide variety of different habitat types including primary rainforests, secondary rainforests, swamps,
bamboo forest Bamboos are a diverse group of evergreen perennial flowering plants making up the subfamily Bambusoideae of the grass family Poaceae. Giant bamboos are the largest members of the grass family. The origin of the word "bamboo" is uncertain, but ...
s, semi-arid streambeds, slow moving forest streams, seasonal streams,
montane Montane ecosystems are found on the slopes of mountains. The alpine climate in these regions strongly affects the ecosystem because temperatures fall as elevation increases, causing the ecosystem to stratify. This stratification is a crucial ...
grassland A grassland is an area where the vegetation is dominated by grasses ( Poaceae). However, sedge ( Cyperaceae) and rush ( Juncaceae) can also be found along with variable proportions of legumes, like clover, and other herbs. Grasslands occur na ...
savanna A savanna or savannah is a mixed woodland- grassland (i.e. grassy woodland) ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach the ground to ...
h, and wet canyons. Some members of the genus such as Ebenau's mantella (''Mantella ebenaui''), the brown mantella (''Mantella betsileo''), and
Cowan's mantella The Cowan's mantella (''Mantella cowanii'') is a species of frog in the family Mantellidae. It is endemic to Madagascar. Its natural habitats are subtropical or tropical moist montane forests, subtropical or tropical high-altitude grassland, and ...
(''Mantella cowanii'') are highly adaptable and have been reported in a wide variety of habitats. On the island of
Nosy Boraha Nosy Boraha , previously known as Sainte-Marie, main town Ambodifotatra, is an island off the east coast of Madagascar. The island forms an administrative district within Analanjirofo Region, and covers an area of 222 km2. It has a popul ...
(Sainte Marie), ''M. ebenaui'' have been found living in garbage dumps, feeding on flies. Similar behavior has been reported in western Madagascar, with ''M. betsileo'' inhabiting rubbish piles behind human dwellings.


Locality variations

There are several populations of ''Mantella'' species that exhibit unusual coloration, some of which are intermediates between species living in sympatry. For example, there are populations of yellow mantella (''Mantella crocea'') and black-eared mantella (''Mantella milotympanum'') found in Fierenana, Andriabe, Ambohitantely Reserve and Savakoanina that have green, red and yellow colourations. This often makes it difficult to distinguish between the two species. Populations of Baron's mantella (''Mantella baroni'') have also been reported at
Pic d'Ivohibe Reserve Pic d'Ivohibe Reserve is a wildlife reserve of Madagascar. It was created in 1964. Geography Ivohibe is in the southern part of the Andringitra Massif, in the region of the Ihorombe. The reserve covers 3 453 ha. It is situated near the village ...
, being almost entirely green in coloration with black patches and spotting, and lacking their distinctive orange and irregular black crossbands. These specimens are referred to as ''Mantella aff. baroni''. Malagasy mantella (''Mantella madagascariensis''), a species similar in appearance to ''M. baroni'', is also notably variable among different localities. Niagarakely is one such locality within the
Anosibe An'ala District Anosibe An'ala District is a district in the Alaotra-Mangoro region in Madagascar. Its capital is Anosibe An'ala. Communes The district is further divided into 11 communes: * Ambalaomby * Ambatoharanana Ambatoharanana is a rural municipality ...
of the
Alaotra-Mangoro Alaotra-Mangoro is a region in eastern Madagascar. It borders Sofia Region in north, Analanjirofo in northeast, Atsinanana in east, Vakinankaratra in southwest, Analamanga in west and Betsiboka in northwest. The capital of the region is Ambat ...
Region. Here, ''M. madagascariensis'' exhibit highly broken yellow/green and mottled black dorsal coloration.


Species

There are currently 16 species of ''Mantella'', with five recognized species groups. Most species are easily identifiable by their color patterns, although there are a number of locality variations with an uncertain taxonomic status.


Threats

Several species in the genus are threatened because of
habitat loss Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...
(due to subsistence agriculture, timber extraction and charcoal production, fires, draining of wetlands, the spread of invasive eucalyptus, and expanding human settlements),
mining Mining is the extraction of valuable minerals or other geological materials from the Earth, usually from an ore body, lode, vein, seam, reef, or placer deposit. The exploitation of these deposits for raw material is based on the economic ...
, hybridization and over-collection for the international pet trade. As a result of these threats, various ''Mantella'' sp. are listed as
least concern A least-concern species is a species that has been categorized by the International Union for Conservation of Nature (IUCN) as evaluated as not being a focus of species conservation because the specific species is still plentiful in the wild. T ...
,
near threatened A near-threatened species is a species which has been categorized as "Near Threatened" (NT) by the International Union for Conservation of Nature as that may be vulnerable to endangerment in the near future, but it does not currently qualify f ...
, vulnerable,
endangered An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching and in ...
, and critically endangered by the IUCN Red List of Threatened Species. Species in this genus have tested positive for ''Batrachochytrium dendrobatidis'' (''Bd''). As of yet, there have been no negative effects observed within amphibian populations in Madagascar, suggesting that the ''Bd'' strain has a low virulence level but should be closely monitored.


Gallery

Image:Mantella aurantiaca58.JPG, ''Mantella aurantiaca'' Image:Mantella baroni15.JPG, ''Mantella baroni'' Image:Mantella bernhardi47.JPG, ''Mantella bernhardi'' Image:Mantella cowanii01.jpg, ''Mantella cowanii'' Image:Mantella expectata01.jpg, ''Mantella expectata''


References


External links

* * eb application 2008. Berkeley, California
''Mantella''
AmphibiaWeb, available at http://amphibiaweb.org/. (Accessed: June 8, 2008). * taxo
''Mantella''
at http://www.eol.org. * Taxo
''Mantella''
at https://web.archive.org/web/20160606043808/http://www.itis.gov/index.html. (Accessed: June 8, 2008). * Taxo
''Mantella''
at https://web.archive.org/web/20080501142231/http://data.gbif.org/welcome.htm
''Mantella'' ConservationPoison Frogs of Madagascar: ''Mantella''
*Vences, M.; Glaw, F.; Böhme, W. (1999).
A review of the genus Mantella (Anura, Ranidae, Mantellinae): taxonomy, distribution and conservation of Malagasy poison frogs
(PDF). ''Alytes''. 17 (1–2): 3–72. {{Taxonbar , from=Q1627754 Mantellidae Amphibian genera Frogs of Africa Endemic fauna of Madagascar Taxa named by George Albert Boulenger Species endangered by habitat loss Species endangered by the pet trade