1 AMPHIBIA: ANURA: DENDROBATIDAE Andinobates bombetes Catalogue of American Amphibians and Minyobates bombetes: Myers 1987:304. Reptiles 926 Dendrobates bombetes: Jungfer, Lötters, and Jörgens 2000:11. F. Vargas-Salinas, M. A. Atehortua-Vallejo, L. Ranitomeya bombetes: Grant, Frost, Caldwell, F. Arcila-Pérez, G. M. Jiménez-Vargas, Gagliardo, Haddad, Kok, Means, Noonan, C. González-Acosta, S. Casas-Cardona, Schargel, and Wheeler 2006:171. and A. Grajales-Echeverry. 2020. Andinobates bombetes: Brown, Twomey, Andinobates bombetes. Amézquita, de Souza, Caldwell, Lötters, May, Melo-Sampaio, Mejía-Vargas, Pe- Andinobates bombetes (Myers and Daly) rez-Peña, Pepper, Poelman, Sanchez-Ro- Rubí poison frog driguez, and Summers 2011:36. Dendrobates bombetes Myers and Daly CONTENT. No subspecies are recognized. 1980:2. Type locality: “mountains above south side of Lago de Calima, 1580–1600 DESCRIPTION. Individuals of Andinobates meters elevation, about 2 km.,{sic} airline bombetes have a body size (snout-vent length, southwest of Puente Tierra (village), De- SVL) between 16.7– 21.5 mm, with no sexual partment of Valle del Cauca, Colombia.” dimorphism in body size (males: mean SVL Holotype, American Museum of Natural = 17.8 ± 0.1 mm SD, range: 16.7–21.5 mm; History, AMNH 102601, adult female, N = 28; females: mean SVL = 18.6 ± 0.1 mm collected by C. W. Myers, J. W. Daly and SD, range: 17.2–19.8 mm; N = 19) (Myers and E. B. de Bernal on 21 November 1976 (not Daly 1980; Suárez-Mayorga 1999; Vargas-Sa- examined by authors). linas and Amézquita 2013). In adults of Andi- Figure 1. Male of Andinobates bombetes from Finca El Placer, in the municipality of Filandia, depart- ment of Quindío, Colombia. Picture by Cristian González-Acosta. 2 Map. Geographic distribution of Andinobates bombetes in Colombia, South America. Red and yellow dots indicate populations where individuals exhibit red and yellow dorsolateral stripes, respectively. The type locality is highlighted with a black dot in the center of the circle. Account 926 3 Figure 2. Dorsal view (A), lateral view (B), and mouth (C) of a tadpole of Andinobates bombetes (Instituto de Ciencias Naturales [ICN] 42287, Museo de Historia Natural, Univer- sidad Nacional de Colombia, Bogotá). Photographs courtesy of David Sánchez. nobates bombetes, the head is usually narrow- the eyes, typically ending at midbody. Skin is er than the body, especially in gravid females; granular with granulation being particularly greatest head width averaging about 32–33% coarse and strong on lower back and hind of SVL in adults. Nostrils are located near the limbs. The ventral surface is cream white with tip of the snout and are angled in a posterolat- brown blotches. The iris is dark brown in life eral direction, the canthus rostralis is round- with little contrast from the pupil. ed, the loreal region is virtually flat, the eyes The morphology of seven tadpoles at are positioned dorsally, and the eardrum is Gosner stage 25 (Gosner 1960) transported hidden posterodorsally. Relative length of the on the back of parents was described by My- fingers is III>IV>II>I with an expanded ter- ers and Daly (1980), and the morphology of minal disk on all digits except Finger I which four tadpoles at Gosner stage 25, five tadpoles is short; in juveniles, terminal discs are rel- at Gosner stages 25–30, and two tadpoles at atively less expanded. Relative length of the Gosner stages 42–43 was described by Sán- toes is IV>III>V>II>I. chez (2013). At Gosner stage 25, following In life, the upper part of the body and the description of Myers and Daly (1980), the limbs are dark brown. Individuals possess tadpole has a globular shape from a dorsal bright red or gold yellow (rarely light orange) view, with the body width averaging 79.4% stripes that begin at the snout and extend over (74–92%) of the head-body length. The head 4 A B Figure 3. Types of habitats where individuals of Andinobates bombetes have been recorded. A) Humid montane forest; in this habitat individuals lives in places away from streams. B) Tropical dry forest; in this habitat individuals are found alongside streams. Photographs by Fernando Vargas-Salinas. and body are slightly convex above and flat- edges; the lower jaw sheath is V-shaped. Tad- tened below. The eyes and the nostrils are in poles reported by Sanchez (2013) were de- a dorsal position. The spiracle is sinistral and scribed as having a U-shaped jaw sheath, with situated low on the body, and the cloacal tube short guts with visible organs, and the nostrils is dextral. The low-finned tail is 64.3% of the without a projection on the inner margin of total length; the end of the tail is rounded. The the nasal rim. average head-body length is 4.3 mm (range: Individuals of Andinobates bombetes have 4.0–4.6 mm); the average greatest body width toxins in the skin (Myers and Daly 1980). is 3.4 mm (range = 3.1–3.7 mm); the average Those toxins are rich in piperidine alkaloids, total length of the tadpole is 12.2 mm (range with at least 22 kinds; three of those alka- = 11.1–13.7 mm); the average greatest tail loids (247, 251F, and 265B) are present only depth from upper edge dorsal fin to lower in Andinobates bombetes; the fourth alkaloid edge ventral fin is 1.7 mm (range = 1.6–1.8 (217) is shared with other species within the mm). The morphology of the mouth consists family Dendrobatidae. With the exception of of an oral disc that is oriented anteroventrally two unclassified compounds, all the alkaloids and the labial tooth rows are 2/3; the second found in Andinobates bombetes were assigned upper row of teeth has a central gap. The up- to several classes of pumiliotoxins. Despite per jaw sheath is solid with serrated cutting the abundance of alkaloids in their skin tox- Account 926 5 ins, Andinobates bombetes is not highly toxic. brown cryptic coloration (Grant and Castro This species shows geographic variation in 1998; Marin et al. 2018). the skin toxins through qualitative and quan- The tadpole ofAndinobates bombetes can titative differences of alkaloids found in the be differentiated from other congeners be- skin across different populations (Myers and cause of the presence of a U-shaped upper Daly 1980). jaw sheath, a differentiated short gut with The advertisement call of Andinobates visible organs, the absence of a projection bombetes is a ‘buzz’ with an average duration on the inner margin of the nasal rim, and of 1.28 s (range = 0.9–1.7 s), consists of ap- the presence of a median gap that interrupts proximately 160 pulses (pulse emission rate = the papillate fringe on the lower (posterior) 113–134 per second), and its dominant fre- edge of the oral disc (Myers and Daly 1980; quency varies between 4000–5500 Hz (Brown Sánchez 2013). Tadpoles of Andinobates et al. 2011; Myers and Daly 1980; Vargas-Sali- bombetes develop in small pools of bromeli- nas et al. 2014a). See Etymology. ads, distinguishing them from other syntopic Andinobates bombetes is easily distin- dendrobatid tadpoles (Leucostethus) that de- guished from congeneric species based on velop in terrestrial bodies of water (Lötters et small size, and especially because of the dis- al. 2007). tinctive color pattern and the first finger be- ing shorter than the second finger (Myers and DIAGNOSIS. Andinobates bombetes has the Daly 1980). Despite the descriptions of new following diagnostic characters: vomerine species and the subsequent rearrangement of teeth are absent; width of head is usually less the phylogenetic relationships among species than width of the body; dorsal and ventral (Amézquita et al. 2013; Brown et al. 2011; skin are granular, particularly coarse on lower Márquez et al. 2017), these characters are still back and the hind limbs; snout rounded from useful. For instance, Andinobates bombetes a lateral view and bluntly rounded to truncate exhibits a black or dark brown color with red from a dorsal or ventral view; external nares and yellow dorsolateral stripes, while Andi- situated near the tip of the snout and directed nobates opisthomelas is largely red (Brown et posterolaterally; canthus rostralis is rounded; al. 2011; Stuart et al. 2008), the posterior two- the loreal region is flat; tympanum is hidden thirds of the dorsal surface of Andinobates posterodorsally; the relative length of the fin- virolinensis is dark brown (Ruiz-Carranza gers have the following order III>IV≥II>I; and Ramírez-Pinilla 1992), Andinobates do- distal discs of the fingers are expanded on risswansonae is dark brown or black with ir- all of them except the first one; a large circu- regular red blotches (Rueda-Almonacid et al. lar-to-elliptical outer metacarpal tubercle on 2006), Andinobates tolimensis is a metallic yel- median base of palm; the relative length of low-bronze color anteriorly that fades to dark the toes have the following order IV>III>V brown by midbody (Bernal et al. 2007, Bernal >II>I; first toe short and without expanded & Luna-Mora 2014), and Andinobates cassi- distal disc; the coloration of the back and dyhornae has a bright red dorsum with a ven- limbs is black or dark brown with two red ter that is black with well-defined red blotch- dorsolateral stripes (in some populations the es or spots (Amézquita et al. 2013). Moreover, stripes are yellow), which begins at the snout Andinobates bombetes is not sympatric with and extends to midbody although sometimes any other species of the genus Andinobates. extending the full length of the body; ventral Some populations are syntopic with species of coloration of the head, body, and hind limbs the genus Leucostethus, but those species are is motted black on a pale yellow or pale green.