Andinobates supata, Chaves-Portilla & Salazar & Gil-Acero & Dorado-Correa & Márquez & Rueda-Almonacid & Amézquita, 2021

Chaves-Portilla, Giovanni A., Salazar, Erika Nathalia, Gil-Acero, José, Dorado-Correa, Adriana, Márquez, Roberto, Rueda-Almonacid, José Vicente & Amézquita, Adolfo, 2021, A new species of Andean golden poison frog (Andinobates, Dendrobatidae) from the Eastern Andes of Colombia, Zootaxa 5047 (5), pp. 531-546 : 536-541

Andinobates supata sp. nov.

Figures 1 , 2 , 3 , 4 (E-M) and 5. Table 1.

Andinobates sp. Lötters et al. 2007 (Figures 571–574)

Andinobates sp. “supatá” Brown & Twomey et al. 2011.

Andinobates sp. aff. tolimensis Brown & Twomey et al. 2011 ( Figures 6–7 )

“ Andinobates supatae” Salazar, Márquez, Chaves-Portilla, Dorado-Correa & Amézquita ; In Kahn et al. (2016), species account 290–294.

Holotype. Adult female ( MUJ 6604 ), Colombia, Departamento de Cundinamarca, about 8 km south of the Municipio de Supatá , Vereda San Marcos (ca 5°3’N, 74°14’W, 1890–2000 m elevation), collected by Giovanni Chaves- Portilla, Erika Nathalia Salazar and José Gil-Acero on 3 February 2007.

Paratypes. Three adult males ( MUJ 6605 , 6607,6608 ), one adult female ( MUJ 6609 ), one sub-adult female ( MUJ 6606 ), all collected with the holotype .

Additional referred material. A batch of four tadpoles ( MUJ 6610) bred in captivity.

Etymology. The species is dedicated to Chief Panché Supatá (or Chupatá), who lived during the time of the Spanish Conquest (first half of the 16th Century) in the Gualivá Province, where the municipality of Supatá is locat- ed. He is believed to have died defending his people in the battle of Tocarema, when the Spanish and their Chibcha allies defeated the Panché in 1537. We also dedicate this species to the modern inhabitants of Supatá for embracing the Supatá Golden Frog as a conservation icon and incorporating it into their culture and festivities.

Vernacular name. Supatá Golden Poison Frog (Rana Dorada de Supatá).

Definition. Andinobates supata sp nov. is a small-sized dendrobatid frog assigned to the A. bombetes group based on the following combination of traits: (1) SVL less than 20 mm (SVL 16.0– 18.4 mm; Table 1). (2) Dorsum bicolored, lacking markings; head uniformly ocher or golden; dorsal coloration fades from a brightly colored head to sepia (dark reddish-brown) in the lower dorsum. A cephalic hood covers the sides of the head and a large part of the tympanum ( Figures 3 & 4 ). (3) Conspicuous dorsolateral stripes absent; abdominal flanks sepia with large, round bluish-white spots ( Figures 3 & 4 ). (4) Ventral coloration with distinct, bluish-white spots ( Fig. 4 ). (5) Dorsal skin mostly smooth, becoming slightly granular on the sacral region and thighs. (6) Ventral skin slightly granular. (7) Advertisement calls are “short buzz” type ( Brown & Twomey et al. 2011; Fig. 5 ), note length approximately 1.3 s, with 141–243 pulses per note and a mean dominant frequency of 4.7 kHz. (8) First finger shorter than second; thenar tubercle present. (9) Five toes on foot. (10) Distinct limb reticulation absent. (11) Hands and feet lack webbing. (12) Oral discs of the larvae are normal and non-umbelliform.

Comparisons with other species. Andinobates supata sp. nov. is a close relative of A. opisthomelas , A. virolinensis , A. dorisswansonae , A. tolimensis , A. cassidyhornae , and A. bombetes , all of which occur within roughly 100 km of the type locality of the new species. Of these, only A. tolimensis has yellow dorsal coloration, and therefore is the species most easily confused with Andinobates supata sp. nov. The remainder of these species all have red dorsal coloration (versus yellow in A. supata sp. nov.), including A. virolinensis , which is the only other Andinobates species occurring in the same Eastern Cordillera as Andinobates supata sp. nov. Andinobates tolimensis has dark bluish spots on the venter (versus pale bluish ventral spots in A. supata sp. nov.) and has golden dorsal/cephalic coloration that does not cover the tympanum, leaving the impression of a black stripe running back from the eye (versus golden head coloration complete in A. supata sp. nov.) ( Figures 3 & 4 ). SVL between both species are similar and overlapping although A. tolimensis may be slightly larger (17.7–18.9 mm; n=6) than Andinobates supata sp. nov. (16.0– 18.4 mm).

Description of the holotype. Adult female, 18.4 mm SVL. Dorsal skin smooth on head and anterior part of the body, slightly granular through sacral region and dorsal surface of hindlimbs. Skin on the belly and flanks weakly granular. Head slightly wider than long, relatively short and rounded snout in lateral and dorsal views. Eyes large, diameter more than 40% of head length. Canthus rostralis not very prominent; flat, vertical loreal region. Small rounded nares situated near the tip of the snout, separated from the orbits by a distance equal or slightly shorter than the diameter of the eye. Pupil nearly rounded; horizontal diameter barely exceeding vertical diameter. Tympanum vertically oval and slightly angled in anteroposterior position, separated from the eye by a distance shorter than its own diameter. Tympanic ring prominent, only visible in the anterior and ventral sides. Premaxillary and maxillary teeth absent. Choanae small and rounded, separated from each other, covered by the maxillary palatal arch when seen from above. Tongue spatulate, fixed to the floor of the mouth and free in most of its extension. Medial lingual process absent.

Relative length of adpressed fingers is: III>IV>II>I; finger I reaches below the disc of II; finger II reaches the distal subarticular tubercle of finger III; tip of digit IV barely exceeds its base. Finger and toe discs smooth, truncated, more expanded on fingers than toes; width is proportional to length. Basal subarticular tubercles large, rounded, slightly elevated, as wide as the digit. Distal subarticular tubercle of fingers half the size of basal ones, rounded, slightly protuberant; supernumerary tubercles absent; palmar tubercles large and rounded; thenar tubercle oval, almost three times smaller than palmar tubercle. Fingers lack webbing. Digits rounded without skin on the edge.

Hind limbs exhibit five well-developed, externally visible toes. Relative length of adpressed toes is: IV>III>II>V>I. Disc of toe I as wide as digit. Subarticular tubercles of toes are elongated, barely prominent. Basal tubercles larger than distal ones; sole of foot smooth with supernumerary tubercles absent; inner metatarsal tubercle elongated and slightly raised, almost two and half times larger than outer one; outer metatarsal tubercle rounded and conical. A small tubercle (about half the size of the external metatarsal tubercle) located in the inner and medial edge of the tarsus. Toes lack webbing. When the hind limbs are close to the sides of the body and stretched forward the heel barely reaches the anterior edge of tympanum. Fingers and toes with paired dermal pads on dorsal surfaces of discs.

Measurements (in mm) and mass of the holotype. SVL 18.4; TL 7.9; GBW 9.5; HW 6.2; IOD 2.2; HL 5.8; TSCN 0.6; NED 1.6; IND 2.3; ED 2.3; HDT 1.0; MET 0.9; HaL 4.6; W3FD 0.8; W3F 0.5; W3TD 0.7; W3T 0.5; W4TD 0.6; W4T 0.4; body mass 0.4 g.

Color in life. Iris black; cephalic hood gold, transitioning to dark brown to sepia near vent and suprascapular region. The remaining body surfaces are sepia. Abdominal flanks, hidden surfaces of the limbs, and belly are predominantly black with large bluish-white spots. Cephalic hood completely covers the back and sides of the head with exception of the black nares and black anterior region of the tympanum.

Color in preservative (ethanol 70%). Head, anterior surface of the trunk, and posterior part of the arm are metallic bluish gray. Posterior surface of the body gray to bronze, gradually fading into sepia. Arms and dorsal surface of the thighs, legs, and foot are sepia or blackish. Lateral surface and belly are sepia or black with large and rounded white to light blue spots. Thighs and hidden surfaces of the forelimbs and hind limbs all black with little bluish to white spots.

Vocalization. Advertisement calls are of the ‘buzz’ type ( Brown & Twomey et al. 2011; Myers & Daly 1976), in which a single trunk muscle contraction produces what humans perceive as a single ‘buzz’. Therefore, the recommended behavioral unit of analysis is one ‘buzz’ ( Erdtmann & Amézquita 2009). In the corresponding oscillogram and sonogram, the buzz consists of a long series of pulses with amplitude but not frequency modulation ( Fig. 5 ), approximately 1.3 s in length. Calls contain 141–243 pulses per call (201.8 ±39.7) and the peak frequency ranges between 4.67–4.84 kHz (4.7 ±0.07 kHz).

Sexual dimorphism and inter-individual variability. Adult females are slightly larger in SVL (both females 18.4 mm in females vs. 16.0– 17.1 mm in males) and slightly heavier than males (0.4 g in females vs. 0.3–0.4 g in males). Dorsal skin is more granular in males than females, however there is considerable variation among individuals. Lastly, on some specimens the yellow coloration of the arm is reduced to a patch on its base ( Figures 3 & 4 ). Measurements of additional specimens are presented in Table 3.

Tadpole description. We describe one specimen from lot MUJ 6610, in stage 36 sensu Gosner (1960). The description was complemented with the remaining three larvae, between stages 32–36. All specimens were obtained from individuals bred and maintained in captivity.

Body elongated, depressed, ventrally flat. Head rounded when viewed from above and from the side. Eyes dorsal, small, not prominent, spaced by about 1.5 times their own diameter. Nares small, located in the anterior dorsal region, spaced from each other and the eyes by a distance roughly equal to the orbital diameter. Operculum located in the latero-ventral margin of the left side of the body. Anal tube dextral and short.

Oral disc emarginated, transversally elliptic, located in the ventral and anterior extreme of the head; surrounded by a row of small, digitiform papillae on the sides and bottom. Submarginal papillae absent; upper labium is smooth and lacks papillae on most of its anterior margin. Labial tooth row formula is 2/3. Outer tooth rows well developed compared to the inner rows. Second tooth row of the upper labium is discontinuous towards the sides of the mouth. Beak has tiny serrations on the edges.

Tail long and laterally compressed with the tip slightly rounded and the caudal musculature well developed. Musculature at mid-tail greatly exceeds musculature of the fins. Upper fin barely visible in the anterior third of the tail, reaching maximal width halfway through the tail.

In preservative larvae are dark gray; the tail is lighter than the body. Abdominal surfaces are opaque, with internal organs hardly visible.

Measurements: ToL 26.9 mm; HBL 8.9 mm; BW 6.3 mm; BH 3.2 mm; TaL 18.0 mm; TH 3.7 mm; TMW 2.0 mm; TMH 2.4 mm; MTMW 2.5 mm.

Distribution. Andinobates supata sp. nov. occurs in a few small forest fragments at the type locality, and near the municipality of San Francisco (Cundinamarca, Colombia). Despite numerous expeditions undertaken by our colleagues and us over the last decade in the western versant of the Andes in Cundinamarca, with the specific objective of finding populations of this and other endangered amphibian species (e.g. Atelopus spp. Bolitoglossa spp. ), we are not aware of any other locality where this species occurs. Citizen science-based efforts and publicity campaigns attempting to crowdsource new reports of this species have also been unsuccessful.

Natural history and ecology. Like other dendrobatid frogs, Andinobates supata sp. nov. is a diurnal species with a bimodal calling activity pattern, peaking in early in the morning (6:00 to 9:00 am) and in the mid-afternoon (15:00 to 17:00). However, during several nocturnal surveys, males have been observed to be active and calling from the forest floor. Male density appears to be very high. In a single day we observed 39 individual males (individually identified by their ventral color pattern) within a ~0.6 hectare plot that included both forested and open areas. Andinobates supata sp. nov. is commonly found in the forest floor, among the leaf litter, or occupying small cavities in the ground, under fallen trunks, within tree roots, and even in cavities formed by grass tufts in open areas adjacent to the forest, sometimes up to 75 m away from the forest edge. Males attract females by emitting advertisement calls and actively repel other males by increasing call rate or by direct fighting. During courtship, the female will follow the male into a cave (usually under fallen logs or under the roots of trees), where mating and egg laying presumably occur. Once larvae hatch, males carry up to three tadpoles on their back ( Fig. 6 ). In some instances, we observed males carrying tadpoles of noticeably different sizes, which lead us to suspect that they may be at different developmental stages. Males climb up tree trunks and presumably deposit the larvae in phytotelmata, where they develop until metamorphosis. Actively calling males are more common at sites where the density of bromeliads of the Guzmania genus is higher (Salazar E.N., Gil. J.D., Amézquita. A., unpublished data). Calling individuals are active throughout the year, but males carrying larvae are more often found during the rainy season. Some individuals have been observed feeding on small invertebrates such as Collembola, Protura and Diplura.

Figures

FIGURE 1. Localities of Andinobates supata sp. nov. and six species of Andinobates used in the phylogenetic analysis (A. cassidyhornae, A. opisthomelas, A. bombetes, A. tolimensis, A. virolinensis and A. dorisswansonae).

FIGURE 2. Bayesian mtDNA phylogeny of 11 Andinobates species, based on 16S and Cytb genes, with Ranitomeya ventrimaculata used as outgroup. Nodal support is represented on internodes as Bayesian posterior probability/ML bootstrap. Values for nodes with support below 0.9/0.7 are not shown. Support values for some intraspecific nodes are also not shown to improve visualization.

FIGURE 3. Body coloration of the two yellow species of Andinobates described to date. Note the postocular dark band in A. tolimensis (above) as opposed to the complete cephalic hood in A. supata sp. nov. (below). Photos by Manuel H. Bernal and Jennifer Del Río, respectively.

FIGURE 4. Ventral coloration of Andinobates supata sp. nov. compared to the geographically and genetically closest species of Andinobates. A. bombetes (A), A. virolinensis (B), A. tolimensis (C–D), and A. supata sp. nov. (E–H). Dorsolateral coloration of Andinobates supata sp. nov. (I–M) compared to A. tolimensis (N). Photos in B by Gert Benaets, A, E to M by Giovanni Chaves- Portilla, and C, D and N by Taran Grant.

FIGURE 5. Oscillogram below and sonogram (above) of the advertisement call of Andinobates supata sp. nov. Male size (SVL) = 18.7 mm, body temperature = 18.4° C.

FIGURE 6. Male attracting female to his cave where mating and egg laying presumably occur (A). Male of Andinobates supata sp. nov. carrying three tadpoles presumably to a deposition site such as a bromeliad (B). Photos: Giovanni Chaves-Portilla.

FIGURE 7. View of the degraded habitat of Andinobates supata sp. nov. Photo: Giovanni Chaves-Portilla.