Andinobates Twomey, Brown, Amézquita & Mejía-Vargas, 2011

Brown, Jason L., Twomey, Evan, Amézquita, Adolfo, Souza, Moisés Barbosa De, Caldwell, Jana- Lee P., Lötters, Stefan, May, Rudolf Von, Melo-Sampaio, Paulo Roberto, Mejía-Vargas, Daniel, Perez-Peña, Pedro, Pepper, Mark, Poelman, Erik H., Sanchez-Rodriguez, Manuel & Summers, Kyle, 2011, A taxonomic revision of the Neotropical poison frog genus Ranitomeya (Amphibia: Dendrobatidae) 3083, Zootaxa 3083 (1), pp. 1-120 : 20-30

Andinobates Twomey, Brown, Amézquita & Mejía-Vargas , gen. nov.

Account authors: E. Twomey, J.L. Brown, A. Amézquita, D. Mejía-Vargas

Figs. 3–8

Tables 2 –6

Type species. Dendrobates bombetes Myers & Daly 1980

Etymology. From the Spanish adjective Andino (of the Andes) + the Greek bates (a walker), referring to the distribution of this genus, being primarily in the northern Andes. Gender masculine. The second half of the name is common among dendrobatine poison frogs.

Proposed sister group. Ranitomeya .

Definition and diagnosis. Unambiguous synapomorphies include: 15 nuclear and 11 mitochondrial synapomorphies (based upon the dataset used in this study, Fig. 3, Tables 2 and 3); vertebrae 2 and 3 fused (known only in A. claudiae , A. minuta , A. opisthomelas , and A. virolinensis ; trait also present in Oophaga ). Other characteristics include: adult SVL less than 20 mm; adults typically with bright coloration, often with bright red, green, or yellow dorsal coloration (either uniform or as stripes); Andinobates abditus an exception, dorsum mostly black or brown, ...... continued on the next page ...... continued on the next page ...... continued on the next page orange spots in axilla and groin. When present, complete dorsolateral stripe ending before thigh (type ‘A’ in Grant et al. 2006); ventrolateral stripe absent; when present, oblique lateral stripe incomplete. Limbs usually dark with or without pale longitudinal stripes, lacking discrete pale reticulation as in most species of Ranitomeya . Ventral coloration variable, usually with distinct, bright markings. Colored throat patch absent. Head narrower than body; teeth absent; vocal slits present in males. First finger distinctly shorter than second; finger discs II and III weakly to moderately expanded; toe disc III and IV weakly expanded; toe V unexpanded; toe webbing absent; median lingual process absent; tadpoles with LTRF 2(2)/3 (sometimes with gap in first posterior row, Table 4); larval oral disc emarginated; larvae with complete papillae on the posterior labium (so far known in A. minuta and A. claudiae ) or wide medial gap in papillae on posterior labium (so far known in A. abditus , A. bombetes , A. opisthomelas , A. virolinensis and A. tolimensis ).

Distribution. This genus occurs within the rainforests of Colombia (Departments: Antioquia, Chocó, Santander, Cundinamarca, Caldas, Cauca, Córdoba, Tolima, Valle del Cauca, Quindío, Risaralda), Ecuador (Provinces: Napo) and Panama (Provinces: Bocas del Toro, Colón, Coclé, Kuna Yala, Veraguas).

Vocalizations. Two main call groups within Andinobates correspond to the Andean and Chocoan clades ( Fig. 8, Table 5). Species in the former ( A. bombetes , A. opisthomelas , A. daleswansoni , A. dorisswansonae , A. tolimensis , A. virolinensis ) produce calls that can be described as an extended rattle or buzz. These calls are typically over 1 sec in length and are relatively atonal with low energy. Chocoan species ( A. claudiae , A. fulguritus , A. minutus ) produce high-energy, short calls (less than 1 sec.) that are more reminiscent of calls produced by Ranitomeya species of the variabilis group. Data on A. abditus , A. viridis , A. virolinensis and A. altobueyensis are lacking.

Species included (12). Dendrobates abditus Myers & Daly 1976 ; Dendrobates altobueyensis Silverstone 1975 ; Dendrobates bombetes Myers & Daly 1980 ; Dendrobates claudiae Jungfer, Lötters & Jörgens 2000 ; Dendrobates daleswansoni Rueda-Almonacid, Rada, Sánchez-Pacheco, Velásquez-Álvarez & Quevedo-Gil 2006 ; Dendrobates dorisswansonae Rueda-Almonacid, Rada, Sánchez-Pacheco, Velásquez-Álvarez & Quevedo-Gil 2006 ; Dendrobates fulguritus Silverstone 1975 ; Dendrobates minutus Shreve 1935 ; Dendrobates opisthomelas Boulenger 1899 ; Ranitomeya tolimensis Bernal-Bautista, Luna-Mora, Gallego & Quevedo-Gil 2007 ; Dendrobates viridis Myers & Daly 1976 ; Dendrobates virolinensis Ruiz-Carranza & Ramírez-Pinilla 1992 .

Remarks. Our definition of Andinobates is essentially equal to Myers’ (1987) definition of Minyobates minus steyermarki . When molecular data became available for Minyobates steyermarki ( Vences et al. 2003; Roberts et al. 2006a), it was evident that this species was not affiliated with the other members of Minyobates sensu Myers (1987) , but rather was more closely related to the Oophaga + Dendrobates + Adelphobates clade (see discussion for more information). The ancestor to Andinobates apparently diverged during the mid-Miocene from Excidobates about 17 mya and from Ranitomeya sensu this paper about 14 mya ( Santos et al. 2009).