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).

Figures

FIGURE 3. A consensus Bayesian phylogeny based on 1011 base pairs of aligned mitochondrial DNA sequences of the 12S (12s rRNA), 16S (16s rRNA) and cytb (cytochrome-b gene) regions. Thickened branches represent nodes with posterior probabilities 90 and greater, other values are shown on nodes. Taxon labels depict current specific epithet, number in tree, the epithet being used prior to this revision (contained in parentheses), and the collection locality. A. Top segment. B. Middle segment. C. Bottom segment of phylogeny.

FIGURE 4. Putative species tree for Andinobates, Excidobates, and Ranitomeya. Placement of species where molecular data were lacking (A. altobueyensis, A. viridis, A. abditus, A. daleswansoni and R. opisthomelas) was based on morphology. Andinobates altobueyensis and A. viridis were placed as sister taxa due to the absence of dark pigmentation on dorsal body and limbs and overall similar dorsal coloration and patterning. These species were placed as sister to A. fulguritus (sequenced) on the basis of similar dorsal coloration (bright green to greenish-yellow). Andinobates opisthomelas was placed in the bombetes group in a polytomy with A. bombetes and A. virolinensis (both sequenced) due to their similar advertisement calls and morphology, particularly their red dorsal pattern and marbled venter. Andinobates daleswansoni was placed as sister to A. dorisswansonae due to the absence of a well-defined first toe in both species. Andinobates abditus was placed in the bombetes group based on a larval synapomorphy which appears to be diagnostic of that group (wide medial gap in the papillae on the posterior labium). However, A. abditus was placed as the sister species to all other members of the bombetes group due to the absence of bright dorsal coloration and isolated geographic distribution. Andinobates abditus is currently the only species of its genus known to occur in the east-Andean versant, thus its placement remains speculative until molecular data become available. Photo credits: Thomas Ostrowski, Karl-Heinz Jungfer, Victor Luna-Mora, Giovanni Chaves-Portilla.

FIGURE 5. Andinobates Plate 1. minutus group: A–G: Andinobates claudiae and habitat (all from Bocas del Toro, Panama. Photos T. Ostrowski); A & B: Buena Esperanza; C–F: Isla Colon; G: Cerro Brujo; H: tadpole in phytotelm; I: habitat in Bocas del Toro, Panama. J–M: Andinobates minutus (all from Colombia. Photos DMV unless noted): J & K: Buenaventura, Valle del Cauca; L: Quibdó, Chocó; M: Baudó, Chocó (photo J. Mejía-Vargas). fulguritus group: N–V: Andinobates fulguritus (all from Colombia, photos DMV unless noted): N: Baudó, Chocó (photo J. Mejía-Vargas); O: Playa de Oro, Chocó (type locality); P–R: Uraba, Chocó. S–V: Anchicayá, Valle del Cauca. (nΦ = number of individual in phylogeny, Ω = population sampled in phylogeny).

FIGURE 6. Andinobates Plate 2. fulguritus group: A–D: Andinobates fulguritus (all from Risaralda, Colombia. photos DMV). bombetes group: E: Andinobates abditus type locality (photo W.E. Duellman). F–I: Andinobates daleswansoni and habitat (photos J. Mejía-Vargas); F & G: from type locality; H: type locality habitat; I: overview of habitat – human encroachment continues to threaten the habitat of this species. J–M: Andinobates dorisswansonae from Tolima, Colombia (photos DMV and T. Ostrowski). N–P: Andinobates tolimense from Tolima, Colombia (photos V. Mora-Luna). Q–R: Andinobates sp. aff. tolimense from Supatá, Colombia (photos G. Chaves-Portilla and T. Ostrowski).

FIGURE 7. Andinobates Plate 3. bombetes group: A–B: Andinobates sp. aff. tolimense; A: from Supatá, Colombia (Photo credits: G. Chaves-Portilla); B: Tolima, Colombia (Photo credits: Fundacion ProAves-Alonso Quevedo). C–H: Andinobates bombetes, all from Colombia (photo credits: DMV and T. Ostrowski); C: Quebrada a la Chapa, Boyacá; D-E: Valle del Cauca, T. Ostrowski; F: Yotoco, Valle del Cauca; G: Lowland habitat of A. bombetes near Quebrada a la Chapa; H: Yotoco, Valle del Cauca. I: Andinobates sp. aff. bombetes undisclosed locality in Colombia (photo credits: Dennis Nilsson and DMV). J–N: Andinobates virolinensis; Charalá, Virolin, Colombia. O–Q: Andinobates opisthomelas. O: Guatape, Antioquia, Colombia (photo credits: Dennis Nilsson and DMV); P-Q: Carmen de Atrato, Choco, Colombia.

FIGURE 8. Advertisement calls for species of Andinobates. A. Andinobates bombetes from Bosque Yotoco, Valle del Cauca, Colombia (type locality), recorded at 18-20° C; B. Andinobates claudiae from Isla Colón, Panama, recorded at 25° C (call courtesy Thomas Ostrowski); C. Andinobates fulguritus from Itauri, Colombia, unknown temperature; D. Andinobates fulguritus from Kuna Yala, Panama, recorded in captivity at 24° C (call courtesy T. Ostrowski); E. Andinobates dorisswansonae from “El Estadero”, Caldas, Colombia (type locality), recorded at 19-20° C; F. Andinobates minutus, unknown locality or temperature; G. Andinobates opisthomelas from Guatapé, Antioquia, Colombia, unknown temperature.