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Mesoclemmys raniceps (GRAY, 1856)

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Higher TaxaChelidae, Chelinae, Pleurodira, Testudines (turtles)
Common NamesE: Black-lined Toadhead Turtle
Portuguese: Cágado, Cágado-de-Cabeça-de-Sapo-Comum, Lalá

E: Amazon Toadhead Turtle [heliostemma]
Portuguese: Cágado, Perema-da-Cabeça-Amarela [heliostemma] 
SynonymHydraspis raniceps GRAY 1856
Phrynops wermuthi MERTENS 1969
Phrynops (Batrachemys) nasutus wermuthi — ALDERTON 1988
Phrynops raniceps — IVERSON 1992
Batrachemys raniceps — MCCORD et al. 2001
Batrachemys heliostemma MCCORD, JOSEPH-OUNI & LAMAR 2001
Batrachemys heliostemma — MCCORD & JOSEPH-OUNI 2002
Batrachemys raniceps — MCCORD & JOSEPH-OUNI 2002
Batrachemys raniceps — SOUZA 2004
Batrachemys heliostemma — SOUZA 2004
Mesoclemmys heliostemma — BOUR & ZAHER 2005
Mesoclemmys raniceps — BOUR & ZAHER 2005
Batrachemys (Phrynops) raniceps — BONIN et al 2006
Batrachemys heliostemma — CISNEROS-HEREDIA 2006
Batrachemys heliostemma — REYNOLDS et al. 2007
Mesoclemmys heliostemma — VAN DIJK et al. 2011
Mesoclemmys raniceps — VAN DIJK et al. 2011
Batrachemys raniceps — RIVAS et al. 2012
Phrynops (Batrachemys) raniceps — PHILIPPEN 2012
Batrachemys heliostemma — RIVAS et al. 2012
Mesoclemmys heliostemma — MOLINA et al. 2012
Phrynops (Batrachemys) heliostemma — PHILIPPEN 2012
Mesoclemmys raniceps — TTWG 2021 
DistributionSE Colombia, E Peru, N Bolivia, Brazil (Amazon basin, Mato Grosso, Amazonas, Pará, Rondônia), Ecuador (Amazonia, Sucumbíos, Orellana).

wermuthi: Peru; Type locality: “Amazonian Peru”.

Type locality: “Brazil, Para”.

heliostemma: Venezuela, Ecuador, Peru, Brazil (Roraima, Amazonas, Pará, Mato Grosso, Rondônia, Acre); Type locality: base of Pico da Neblina (situated on the Venezuela/Brazil border) on the left bank of Río Baria (=Río Mawarinuma) (4° 95’ N, 66° 10’ W), a tributary of the Rio Negro, Amazonas, Venezuela.  
TypesLectotype: BMNH 1947.3.5.92, formerly NHM; Bates collection, (designated by Bour & Pauler 1987).
Holotype: USNM 541895 [heliostemma] 
DiagnosisM. raniceps and M. gibba can be distinguished by the head width in relation to caparace lenght. This proportion in M. raniceps are more than 20% in adult individuals, and less than 20% in M. gibba (Rueda-Almonacid et al 2007).

Diagnosis (heliostemma). A medium- to large-sized toadhead turtle of the genus Batrachemys that is distinguished from members of other toadhead genera by its large, wide, rounded head; its narrow parietal roof and parieto-squamosal arch; its reduced neural bone series; the combination of its relatively "shallow" carapace, having a median groove, lacking a keel, absence of lateral marginal narrowing, and presence of mild lateral marginal upturning; the presence of a large antero-medial distal tibial scale; and its distinct plastral pattern.
Batrachemys heliostemma is distinguished from its congeners by its distinct head coloration; diminished size of neck tubercles; lack of horizontal line in the iris; greater parietal roof width; its carapace widest at M7; 4th vertebral scute always wider than long; 11th marginals equal to or wider than 12th; mild upturning of lateral marginals; inter-gular scute narrower than gulars; greater width of the anterior plastral lobe; its plastral seam formula; and its plastral pattern.
Among its congeners Batrachemys heliostemma is sympatric with and most similar to B. raniceps. The two forms share a relatively similar head size and shape, a pointed nose, small barbels, a similar neural bone series, a similar plastral seam formula, lack of keels, a shallow carapace groove, a well-developed distal antero-medial tibial scale, and similar plastral patterns. However, B. heliostemma sp. nov. differs from B. raniceps by having a different head pattern and coloration, a broader parietal roof, a more substantial parieto-squamosal arch, an intergular narrower than its gulars (opposite in B. raniceps), a flatter shell, and 11th pair of marginals equal or wider than 12th (equal in B. raniceps). [from MCCORD et al. 2001]. 
CommentFor a color illustration and distribution map see MCCORD & JOSEPH-OUNI 2002.

Distribution: the report of M. raniceps from Peru (FERRONATO et al. 2011) is actually based on a misidentified M. gibba [see erratum to this paper]. Not in Venezuela fide Rivas et al. 2015.

Habitat: freshwater (rivers, swamps)

Synonymy: P. wermuthi has been synonymized with Phrynops raniceps by Bour & Pauler 1987. Cunha et al. 2019 found that eggs of M. raniceps produced offspring that look like raniceps and heliostemma, proving that both are color morphs of the same species. However, neither raniceps nor heliostemma were included in the phylogenetic analysis of Thomson et al. 2021. 
EtymologyNamed after Greek helios, "sun," and Greek stemma (grammatical gender neutral), "wreath," in reference to the bright yellow-orange facial bands found in many juveniles that extend like a wreath from the supra-tympanum over and across the eyes, ending and joining at the tip of the nose. 
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