“…Trueb (1971) noted the external similarity between the R. margaritifera and R. acrolopha groups, and recent phylogenetic analyses have supported a close relationship between them (e.g., Frost et al, 2006). However, molecular evidence indicates an even closer relationship between the R. acrolopha group and certain species of the R. veraguensis group, specifically R. chavin, R. manu, R. nesiotes, and R. yanachaga (Pramuk, 2006;Chaparro et al, 2007;Pramuk et al, 2008;Van Bocxlaer et al, 2010;Pyron and Wiens, 2011;Peloso et al, 2012;Moravec et al, 2014). Furthermore, the results of those analyses of DNA sequences are consistent with the distribution of several putative morphological synapomorphies, including: (1) occurrence of few, large, unpigmented eggs in the R. acrolopha group and R. chavin, R. justinianoi, R. manu, R. multiverrucosa, R. nesiotes, and R. yanachaga of the R. veraguensis group (Duellman and Toft, 1979;Lehr et al, 2001Lehr et al, , 2005Lehr et al, , 2007Chaparro et al, 2007); (2) m. levator mandibulae externus undivided with trigeminal nerve passing medial (deep) to the muscle in R. paraguas (reported herein) and all other species of the R. acrolopha group (TG, personal observation) and at least R. manu (Chaparro et al, 2007) and R. quechua (TG, personal observation) of the R. veraguensis group; and (3) absence of the m. adductor longus in the R. acrolopha group (Trueb, 1971; TG, personal observation) and R. manu (Chaparro et al, 2007).…”