Rhinella sternosignata (Gunther, 1858)

Pereyra, Martín O., Blotto, Boris L., Baldo, Diego, Chaparro, Juan C., Ron, Santiago R., Elias-Costa, Agustín J., Iglesias, Patricia P., Venegas, Pablo J., C. Thomé, Maria Tereza, Ospina-Sarria, Jhon Jairo, Maciel, Natan M., Rada, Marco, Kolenc, Francisco, Borteiro, Claudio, Rivera-Correa, Mauricio, Rojas-Runjaic, Fernando J. M., Moravec, Jiří, De La Riva, Ignacio, Wheeler, Ward C., Castroviejo-Fisher, Santiago, Grant, Taran, Haddad, Célio F. B. & Faivovich, Julián, 2021, Evolution In The Genus Rhinella: A Total Evidence Phylogenetic Analysis Of Neotropical True Toads (Anura: Bufonidae), Bulletin of the American Museum of Natural History 2021 (447), pp. 1-157 : 57

RHINELLA STERNOSIGNATA

DIAGNOSIS: Rhinella sternosignata ( Günther, 1858b) was recovered as the sister taxon of all other species of the R. margaritifera Clade, with poor support (JGC = 49%, JAC = 71%). Phenotypic autapomorphies are: (1) acuminate anterior margins of nasals (char. 6.1), with instances of homoplasy in Incilius coniferus and the R. margaritifera Group; (2) articulation of the zygomatic ramus of the squamosal with the maxilla (char. 15.1), with instances of homoplasy in the R. granulosa Group and Peltophryne lemur ; (3) articulation between the medial ramus of the pterygoid and parasphenoid alae with a jagged suture (char. 22.1) with instances of homoplasy in R. achalensis , R. atacamensis , some species of the R. festae and R. margaritifera Groups , and in the R. marina Group; (4) parotoid gland round to ovoid mostly symmetrical (char. 56.2); (5) large size of adult males with respect to adult females (char. 66.1), with instances of homoplasy in R. yanachaga , and in several species of the R. marina Clade; and (6) unpigmented eggs (char. 88.0), with instances of homoplasy in Ansonia longidigita , Rhinella justinianoi , R. stanlaii , and in the R. festae Group.

DISTRIBUTION: This species inhabits montane forests of the Cordillera de la Costa and the Andean Cordillera de Mérida of Venezuela (La Marca and Mijares-Urrutia, 1996; Barrio- Amorós et al., 2019). See map 5 (available at https://doi.org/10.5531/sd.sp.46) for type and sampled localities .

SISTER CLADE: The clade composed of the Rhinella festae , R. margaritifera , and R. veraguensis Groups.

COMMENTS: This species was tentatively associated with the Rhinella margaritifera ( Cei, 1972a; Hoogmoed, 1990; Duellman and Schulte, 1992) or R. granulosa Groups ( Gallardo, 1962) . Pereyra et al. (2016a) rejected the inclusion of this species in any of these groups, but they could not determine its relationships rigorously due to the poor sampling of Rhinella . This species was wrongly reported for many localities outside the Cordillera de la Costa montane forests region in Venezuela as discussed by La Marca and Mijares-Urrutia (1996). Vélez-Rodríguez (1999) recorded this species in error for Colombia (see Vélez-Rodríguez, 2004b, 2005). Additionally, there are a large number of recent reports of R. sternosignata for Colombia ( Acosta-Galvis et al., 2006; Romero et al., 2008; Acosta-Galvis, 2012a, 2012b). Analyzed specimens tentatively assigned to this species from the eastern slope of the Cordillera Oriental in Colombia (MAR 1314, Boyacá and MAR 1955, Caquetá) were unrelated to the specimen of R. sternosignata from Venezuela in the phylogenetic analyses, and instead, they represent an undescribed species along with other specimens of the R. margaritifera Group from Loreto, Peru, and Miranda, Venezuela ( Rhinella sp. 13, see below). These results, and the absence of comprehensive comparative studies considering topotypical material of R. sternosignata , indicate that there is no evidence to consider its occurrence in Colombia.

THE RHINELLA VERAGUENSIS GROUP DIAGNOSIS: No phenotypic synapomorphies were recovered for this strongly supported group (JGC and JAF = 100%). This is mainly due to the lack of detailed information for one of its two con-