# Mantella

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{{Short description|Genus of amphibians}}
{{Automatic taxobox
| image = Maurantiacamenalamba.jpg
| image_caption = [Golden mantella](/source/Golden_mantella) (''Mantella aurantiaca'') from the Ramsar Menalamba site, Torotorofotsy Wetlands Reserve.
| taxon = Mantella
| authority = [Boulenger](/source/George_Albert_Boulenger), 1882
| type_species = ''[Dendrobates betsileo](/source/Mantella_betsileo)''
| type_species_authority = [Grandidier](/source/Alfred_Grandidier), 1872
| diversity_link = #Species
| diversity = 16 species
}}

'''''Mantella''''' (also known as '''golden frogs''' or '''Malagasy poison frogs''') are a prominent genus of [aposematic](/source/Aposematism) [frog](/source/frog)s in the family [Mantellidae](/source/Mantellidae), [endemic](/source/Endemism) to the island of [Madagascar](/source/Madagascar).<ref name=Frost/> Members of ''Mantella'' are [diurnal](/source/Diurnality) and [terrestrial](/source/Terrestrial_animal), with bright aposematic coloration or cryptic markings.

== Natural history ==
''Mantella'' are an example of [convergent evolution](/source/convergent_evolution)—the independent evolution of a similar trait with species of a different lineage—with the [Latin American](/source/Latin_America) family [Dendrobatidae](/source/Dendrobatidae) in size, appearance, and some behavioral characteristics. During the description of the first specimens from 1866 to 1872, [Alfred Grandidier](/source/Alfred_Grandidier) described both the [brown mantella](/source/Mantella_betsileo) (''Mantella betsileo'') and [Malagasy mantella](/source/Mantella_madagascariensis) (''Mantella madagascariensis'') and placed them within the genus ''[Dendrobates](/source/Dendrobates)'' based on their close resemblance.

This placement was heavily debated until 1882, when [George Albert Boulenger](/source/George_Albert_Boulenger) created the genus ''Mantella'' after describing both [Cowan's mantella](/source/Cowan's_mantella) (''Mantella cowanii'') and, in 1888, [Baron's mantella](/source/Mantella_baroni) (''Mantella baroni''). ''M. baroni'' was named after the gentleman that collected the specimens, [Rev. Richard Baron](/source/Richard_Baron_(botanist)), a [missionary](/source/missionary) and [botanist](/source/botanist) living in Madagascar. Baron was also interested in geology and [herpetology](/source/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](/source/France) [naturalist](/source/naturalist) [Alexandre Thominot](/source/Alexandre_Thominot) described ''Phrynomantis maculatus'', with its type locality on [Réunion Island](/source/R%C3%A9union). However, this locality was later corrected to the off-shore Malagasy islands of [Nosy Bé](/source/Nosy_Be) and [Nosy Komba](/source/Nosy_Komba) and ''P. maculatus'' was [synonymized](/source/Synonymizing) with ''M. baroni''.

The genus remained within Dendrobatidae until the late 19th century. The Royal Natural History (1893) by [Richard Lydekker](/source/Richard_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.''"<ref>{{Cite book|title=The Royal Natural History Vol. V - Reptiles, Amphibians and Fishes |last=Lydekker|first=Richard|publisher=Frederick Warne & Co.|year=1893|pages=271}}</ref>

During the first quarter of the 20th century, another three species of ''Mantella'' were described, including the [golden mantella](/source/golden_mantella) (''Mantella aurantiaca''), by the French herpetologist [François Mocquard](/source/Fran%C3%A7ois_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](/source/Franz_Werner), an [Austria](/source/Austria)n [zoologist](/source/zoologist) and [explorer](/source/Exploration). Mocquard's work describes ''M. attemsi'' as follows: "''First digit extends as far as the second. [Replilatero-dorsal] present, 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''."<ref>{{Cite journal|last=Mocquard|first=Francois|date=1909|title=Synopsis des familles, genres et espèces des reptiles écailleux et des batraciens de Madagascar (1909)|url=https://www.biodiversitylibrary.org/item/44423#page/72/mode/1up|journal=Masson|pages=65–67}}</ref> This species was later synonymised with ''M. betsileo''.

== Description ==
Species of this genus are small, varying in length between {{convert|18|-|31|mm|in}}.<ref name=":0">{{Cite book|title=A Field Guide to the Amphibians and Reptiles of Madagascar |edition=3 |last1=Glaw|first1=Frank|last2=Vences|first2=Miguel|year=2007|isbn=978-3-929449-03-7|location=Köln|pages=188}}</ref> Most ''Mantella'' species are [sexually dimorphic](/source/Sexual_dimorphism) 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](/source/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](/source/Tibiotarsal_joint) articulation is roughly between the shoulder and the nostrils.  

Many species of ''Mantella'' are similar to the [neotropical](/source/Neotropical_realm) family [Dendrobatidae](/source/Dendrobatidae) in their use of [aposematism](/source/aposematism) (from [Greek](/source/Greek_language) 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](/source/alkaloid) secretions. There are significant similarities between a few species of ''Mantella'' and ''Dendrobatidae'', notably the [golden mantella](/source/golden_mantella) (''Mantella aurantiaca'') and the [golden poison frog](/source/golden_poison_frog) (''Phyllobates terribilis''). [Cowan's mantella](/source/Cowan's_mantella) (''Mantella cowanii'') and certain variations of the [Harlequin poison frog](/source/Harlequin_poison_frog) (''Oophaga histrionica'') are also very similar in coloration''.'' Most members of the genus also exhibit aposematism on the [ventral](/source/ventral) region, excluding the [golden mantella](/source/golden_mantella) and [black-eared mantella](/source/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](/source/vocal_sac). These characteristics can be used to distinguish between species, such as [Baron's mantella](/source/Mantella_baroni) (''Mantella baroni'') and the [Malagasy mantella](/source/Madagascan_mantella) (''Mantella madagascariensis''), when locality data is unavailable. 

''Mantella'' show a variety in [alkaloid](/source/alkaloid) 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.<ref>{{Cite journal |last1=Clark |first1=V.C. |last2=Rakotomalala |first2=V. |last3=Ramilijaona |first3=O. |last4=Abrell |first4=L. |last5=Fisher |first5=B.L. |date=2006-10-01 |title=Individual Variation in Alkaloid Content of Poison Frogs of Madagascar (Mantella; Mantellidae) |url=https://doi.org/10.1007/s10886-006-9144-6 |journal=Journal of Chemical Ecology |volume=32 |issue=10 |pages=2219–2233 |doi=10.1007/s10886-006-9144-6 |pmid=17013661 |s2cid=12171450 |issn=1573-1561|url-access=subscription }}</ref> 

== Distribution ==
thumb|Habitat that beautiful mantella (''Mantella pulchra'') can be found around at Vohimana Experimental Reserve, Madagascar
thumb|Bronze variation of yellow mantella (''Mantella crocea'') at the Bakozetra locality, north of the Torotorofotsy Wetlands Reserve
''Mantella'' are endemic to the island of Madagascar and its smaller coastal islands ("Nosy" in [Malagasy](/source/Malagasy_language)). They inhabit a wide variety of different habitat types including primary [rainforest](/source/rainforest)s, secondary rainforests, [swamp](/source/swamp)s, [bamboo forest](/source/bamboo_forest)s, semi-arid [streambeds](/source/Stream_bed), slow moving forest [stream](/source/stream)s, seasonal streams, [montane](/source/Montane_ecosystems) [grassland](/source/grassland) [savanna](/source/savanna)h, and wet canyons.<ref name=":0" /><ref name=":1">{{Cite web|url=http://www.iucnredlist.org|title=The IUCN Red List of Threatened Species|date=2017|website=The IUCN Red List of Threatened Species}}</ref><ref>{{Cite book|title=Mantellas|last=Staniszewski|first=Marc|publisher=Edition Chimaira|year=2001|isbn=978-3930612543}}</ref>

Some members of the genus such as [Ebenau's mantella](/source/Mantella_ebenaui) (''Mantella ebenaui''), the [brown mantella](/source/brown_mantella) (''Mantella betsileo''), and [Cowan's mantella](/source/Cowan's_mantella) (''Mantella cowanii'') are highly adaptable and have been reported in a wide variety of habitats. On the island of [Nosy Boraha](/source/%C3%8Ele_Sainte-Marie) (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](/source/sympatry). For example, there are populations of [yellow mantella](/source/yellow_mantella) (''Mantella crocea'') and [black-eared mantella](/source/black-eared_mantella) (''Mantella milotympanum'') found in [Fierenana](/source/Fierenana%2C_Moramanga), Andriabe, [Ambohitantely](/source/Ambohitantely_Reserve) Reserve and Savakoanina that have green, red and yellow colourations. This often makes it difficult to distinguish between the two species.<ref name=":0" />

Populations of [Baron's mantella](/source/Mantella_baroni) (''Mantella baroni'') have also been reported at [Pic d'Ivohibe Reserve](/source/Pic_d'Ivohibe_Reserve), 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.](/source/aff.) baroni''.<ref>{{Cite journal|last1=Vences|first1=Miguel|last2=Glaw|first2=Frank|last3=Mausfeld|first3=Patrick|last4=Böhme|first4=Wolfgang|date=October 1998|title=Comparative osteology of Malagasy poison frogs of the genus ''Mantella'' |url=http://zoologicalbulletin.de/BzB_Volumes/Volume_48_2/205_215_BZB48_2_Vences_Miguel_et_al.PDF|journal=Zoological Bulletin|volume=48|pages=206}}</ref>

[Malagasy mantella](/source/Madagascan_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](/source/Anosibe_An'ala_District) of the [Alaotra-Mangoro](/source/Alaotra-Mangoro) 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.<ref name=Frost/><ref name=":0" /> Most species are easily identifiable by their color patterns, although there are a number of locality variations with an uncertain taxonomic status.<ref name=":0" />

{| class="wikitable collapsible"
|-
! Species group
! Image
! Binomial (scientific name)
! Common name
!IUCN conservation status
! Distribution
|-
! rowspan="2" style="text-align:center;"| ''Mantella laevigata'' group
| 120px 
|''[Mantella laevigata](/source/Mantella_laevigata)'' {{small|Methuen & Hewitt, 1913}}
| Climbing mantella 
|[Least Concern](/source/Least-concern_species)
|northeastern Madagascar
|-
|
|''[Mantella manery](/source/Mantella_manery)'' {{small|Vences, Glaw & Böhme, 1999}}
| Marojejy mantella
|[Vulnerable](/source/Vulnerable_species)
|northern Madagascar
|-
! rowspan="4" style="text-align:center;"| ''Mantella betsileo'' group
| 120px 
|''[Mantella betsileo](/source/Mantella_betsileo)'' {{small|Grandidier, 1872}}
| Brown mantella
|Least Concern
|western & central Madagascar
|-
| 120px 
|''[Mantella ebenaui](/source/Mantella_ebenaui)'' {{small|Boettger, 1880}}
| Ebenau's mantella
|Least Concern
|northeastern Madagascar
|-
| 120px 
|''[Mantella expectata](/source/Mantella_expectata)'' {{small|Busse & Böhme, 1992}}
| Blue-legged mantella 
|[Endangered](/source/Endangered_species)
|southwestern Madagascar
|-
| 120px 
|''[Mantella viridis](/source/Mantella_viridis)'' {{small|Pintak & Böhme, 1988}}
| Green mantella 
|Endangered
|northern Madagascar
|-
!''Mantella bernhardi'' group
| 120px 
|''[Mantella bernhardi](/source/Mantella_bernhardi)'' {{small|Vences, Glaw, Peyrieras, Böhme & Busse, 1994}}
| Bernhard's mantella
|Vulnerable
|southeastern Madagascar
|-
! rowspan="4" style="text-align:center;"| ''Mantella cowanii'' group
| 120px 
|''[Mantella cowanii](/source/Mantella_cowanii)'' {{small|Boulenger, 1882}} 
| Cowan's or harlequin mantella 
|Endangered
|central Madagascar
|-
| 120px  
|''[Mantella baroni](/source/Mantella_baroni)'' {{small|Boulenger, 1888}}
| Baron's mantella
|Least Concern
|central Madagascar
|-
| 120px 
|''[Mantella haraldmeieri](/source/Mantella_haraldmeieri)'' {{small|Busse, 1981}}
| Harald Meier's mantella
|Endangered
|southern Madagascar
|-
| 120px 
|''[Mantella nigricans](/source/Mantella_nigricans)'' {{small|Guibé, 1978}}
| Guibe's mantella
|Least Concern
|northeastern Madagascar
|-
! rowspan="5" style="text-align:center;"| ''Mantella aurantiaca'' group
| 120px  
|''[Mantella aurantiaca](/source/Mantella_aurantiaca)'' {{small|Mocquard, 1900}}
| Golden mantella
|Endangered
|central Madagascar
|-
| 120px 
|''[Mantella crocea](/source/Mantella_crocea)'' {{small|Pintak & Böhme, 1990}}
| Yellow mantella 
|Vulnerable
|central Madagascar
|-
| 120px 
|''[Mantella madagascariensis](/source/Mantella_madagascariensis)'' {{small|Grandidier, 1872}}
| Malagasy mantella
|Vulnerable
|central Madagascar
|-
| 120px 
|''[Mantella milotympanum](/source/Mantella_milotympanum)'' {{small|Staniszewski, 1996}}
| Black-eared mantella 
|[Critically Endangered](/source/Critically_Endangered)
|eastern Madagascar
|-
| 120px 
|''[Mantella pulchra](/source/Mantella_pulchra)'' {{small|Parker, 1925}}
| Beautiful mantella
|[Near Threatened](/source/Near-threatened_species)
|eastern Madagascar
|-
|}

== Threats ==
[[File:Tavy Fire.jpg|thumb|[Slash-and-burn](/source/Slash-and-burn) farming method (known locally as ''[tavy](/source/Deforestation_in_Madagascar)'') being used near the Vohimana Experimental Reserve, Madagascar.]]
Several species in the genus are threatened because of [habitat loss](/source/habitat_loss) (due to [subsistence agriculture](/source/subsistence_agriculture), timber extraction and charcoal production, fires, draining of wetlands, the spread of invasive eucalyptus, and expanding human settlements), [mining](/source/mining), [hybridization](/source/Hybridization_(biology)) and over-collection for the international pet trade.<ref name=":1" /> As a result of these threats, various ''Mantella'' sp. are listed as [least concern](/source/least-concern_species), [near threatened](/source/near-threatened_species), [vulnerable](/source/vulnerable_species), [endangered](/source/endangered_species), and [critically endangered](/source/critically_endangered) by the [IUCN Red List of Threatened Species](/source/IUCN_Red_List_of_Threatened_Species).

Species in this genus have tested positive for [''Batrachochytrium dendrobatidis'' (''Bd'')](/source/Batrachochytrium_dendrobatidis). 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.<ref name=":1" />

== Gallery ==
<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''
</gallery>

==References==
{{Reflist|30em|refs=
<ref name=Frost>{{cite web |url=http://research.amnh.org/vz/herpetology/amphibia/Amphibia/Anura/Mantellidae/Mantellinae/Mantella |title=''Mantella'' Boulenger, 1882 |author=Frost, Darrel R. |year=2017 |work=Amphibian Species of the World: an Online Reference. Version 6.0 |publisher=American Museum of Natural History |access-date=6 January 2018}}</ref>
}}

== External links ==
* {{cite journal |last1=Clark |first1=Valerie C. |last2=Raxworthy |first2=Christopher J. |last3=Rakotomalala |first3=Valérie |last4=Sierwald |first4=Petra |last5=Fisher |first5=Brian L. |title=Convergent evolution of chemical defense in poison frogs and arthropod prey between Madagascar and the Neotropics |journal=Proceedings of the National Academy of Sciences |date=16 August 2005 |volume=102 |issue=33 |pages=11617–11622 |doi=10.1073/pnas.0503502102 |pmid=16087888 |pmc=1187980 |bibcode=2005PNAS..10211617C |doi-access=free }}
* {{aut|AmphibiaWeb: Information on amphibian biology and conservation.}} [web application]. 2008. Berkeley, California: [http://amphibiaweb.org/cgi-bin/amphib_query?query_src=aw_lists_genera_&table=amphib&where-genus=Mantella&where-species= ''Mantella'']. AmphibiaWeb, available at http://amphibiaweb.org/. (Accessed: June 8, 2008). 
* {{aut|eol - Encyclopedia of Life}} taxon [https://web.archive.org/web/20081023152914/http://www.eol.org/taxa/16141138 ''Mantella''] at http://www.eol.org.
* {{aut|ITIS - Integrated Taxonomic Information System on-line database}} Taxon [https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=208087 ''Mantella''] at https://web.archive.org/web/20160606043808/http://www.itis.gov/index.html. (Accessed: June 8, 2008).
* {{aut|GBIF - Global Biodiversity Information Facility}} Taxon [http://data.gbif.org/species/13183703/ ''Mantella''] at https://web.archive.org/web/20080501142231/http://data.gbif.org/welcome.htm 
*[https://web.archive.org/web/20070823043059/http://www.mantella-conservation.org/ ''Mantella'' Conservation]
*[https://web.archive.org/web/20070928155421/http://mantella.amphibiancare.com/ Poison Frogs of Madagascar: ''Mantella'']
*Vences, M.; Glaw, F.; Böhme, W. (1999). "[http://www.zsm.mwn.de/docs_zsm/htdocs/her/pdf/70_Vences_et_al_1999_Mantella_revision_ALYTES.pdf 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}}

Category:Mantella
Category:Mantellidae
Category:Amphibian genera
Category:Frogs of Africa
Category:Endemic amphibians of Madagascar
Category:Taxa named by George Albert Boulenger
Category:Species that are or were threatened by habitat loss
Category:Species that are or were threatened by the pet trade

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Adapted from the Wikipedia article [Mantella](https://en.wikipedia.org/wiki/Mantella) by Wikipedia contributors ([contributor history](https://en.wikipedia.org/wiki/Mantella?action=history)). Available under [Creative Commons Attribution-ShareAlike 4.0 International](https://creativecommons.org/licenses/by-sa/4.0/). Changes may have been made.
