Myers 1987:304. Dendrobates Bombetes: Jungfer, Lötters, And

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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 being 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. easily distinguished because of their mostly 6

PHYLOGENETIC RELATIONSHIPS. An- PUBLISHED DESCRIPTIONS. Descrip- dinobates bombetes was originally assigned tions of Andinobates bombetes and taxonom- to the genus Dendrobates (Myers and Daly ic reviews of the family Dendrobatidae were 1980) inside of the “minutus group” sensu published by Brown et al. (2011), Clough and Silverstone (1975). Within this group, Den- Summers (2000), Grant et al. (2006, 2017), drobates bombetes, Dendrobates abditus, and Myers (1987), Myers and Daly (1980), and Dendrobates opisthomelas were hypothesized Santos et al. (2009). Observations of breeding to form a monophyletic subgroup based on behavior and parental care of this species were the presence of a median gap that interrupts recorded by Herkrath (2006), Myers and Daly the papillate fringe on the posterior (lower) (1980), and Súarez-Mayorga (1999, 2004); edge of the oral disc in tadpoles. Later, Den- the diet was described by Gómez-Hoyos et al. drobates bombetes was included in the genus (2014); the effectiveness of red and yellow col- Minyobates by Myers (1987) based on the or in dorsolateral stripes as aposematic signal presence of cephalic amplexus, small body was discussed by Casas-Cardona et al. (2018). size, presence of an oblique lateral stripe, lar- The acoustic communication behavior of in- vae with the cloaca in dextral position, and dividuals in anthropogenic noisy habitats and lateral indentations of the oral disc. The spe- the distribution of individuals according to cies was included in the genus Ranitomeya by habitat features (e.g., number of bromeliads) Grant et al. (2006) after taking into account a were studied by Vargas-Salinas and Amézqui- reduction in the length of Finger I, and oth- ta et al. (2013). Geographic variation in the er characteristics such as conspicuous dorsal advertisement call as a possible adaptation to coloration, the absence of toe webbing, and reduce acoustic masking of the call by stream the presence of lipophilic alkaloids. Final- noise was studied by Vargas-Salinas et al. ly, the genus Andinobates was proposed by (2014a). The advertisement call ofAndino - Brown et al. (2011) using a compendium of bates bombetes was described by Myers and morphological, behavioral, ecological, and Daly (1980) and Súarez-Mayorga (1999). molecular characters. Currently, there are 15 species recognized within the genus Andino- ILLUSTRATIONS. Color photographs of bates (Frost 2020); Andinobates bombetes is dorsal and lateral views of individuals with the type species of the genus. red stripes were published by Amézqui- ta and Vélez [2010], Arcila-Pérez et al. (in CONSERVATION STATUS. Andinobates press), Brown et al. (2011), Castro-Herrera bombetes is considered Vulnerable by the and Bolívar-García (2010), Castro-Herrera IUCN (IUCN SSC Amphibian Specialist et al. (2007), Daly et al. (1997), Diaz-Pulido Group 2017). The threats to this species are et al. (2016), Herrmann (2005), Lötters et al. associated with loss of forest habitats that are (2007), Renjifo [1997], Rodríguez-Suárez and converted for agriculture and cattle grazing. Corredor-Londoño (2012), Rueda-Almonac- This species is listed in Appendix II of CITES id et al. (2004), Ruiz-Carranza et al. (1996), because individuals are collected illegally for Suárez-Mayorga (2004), Vanegas-Guerrero et pet trade (CITES 2020). Populations of And- al. (2016), Vargas-Salinas et al. (2014b), and inobates bombetes are present in several nat- Walls (1994); dorsal views of individuals with ural areas protected by Colombia, including yellow stripes were published by Brown et Bosque de Yotoco in the western Andes (Val- al (2011), Suárez-Mayorga et al. (2016), and le del Cauca), and Parque Regional Natural Vargas-Salinas et al. (2014b); a ventral view of Barbas-Bremem in Central Andes (Quindío). an individual with red stripes was published Account 926 7

Figure 4. Males of Andinobates bombetes carrying tadpoles on their backs. A) Individual with red dorsolateral stripes recorded in the Parque Nacional Natural Los Catíos, municipality of Toro, Valle del Cau- ca, Colombia. Photograph by Fernando Vargas-Salinas. B) Individual with yellow dorsolateral stripes recorded in Reserva Natural Altamira, municipality of El Cairo, Valle del Cauca, Colombia. Photograph by Alejandro Grajales-Echeverry. 8

Figure 5. Oscillogram (A) and spectrogram (B) of the advertisement call of Andinobates bombetes. Male body size = 20.09 mm; body temperature of male = 21.6ºC; locality: Parque Nacional Natural Los Catíos, municipality of Toro, Valle del Cauca in the western Andes of Colombia. by Rodríguez-Suárez and Corredor-Londoño Brown et al. (2011). A color photograph of (2012), and similar ventral photographs of an individual with red stripes was mistakenly individuals with yellow stripes were pub- labeled as Ameerega picta by Marent (2008, lished by Brown et al. (2011). Photographs 2010). Color photographs demonstrating the of a red morph male carrying tadpoles on ontogenetic changes in dorsal coloration (red its back were presented by Castaño and Car- morph) were published by Rodríguez-Suárez ranza-Quiceno (2015), Stevens and Ruxton and Corredor-Londoño (2012); color photo- (2012), and Vargas-Salinas et al. (2014b); a graphs of eggs, embryos, and tadpoles in dif- photograph of a yellow morph male carry- ferent developmental stages were published ing tadpoles on its back was published by by Rodríguez-Suárez and Corredor-Lon- Account 926 9 doño (2012). Black-and-white photographs 1998, 2000a, 2000b; Harding 1983; Hutchins of the dorsal view of red-morph adults were et al. 2003; Renjifo [1997]; Ruiz-Carranza et published by Myers and Daly (1980), and the al. 1996), conservation status (Castro-Herre- ornamentation of the margin of the naris at ra and Bolívar-García 2010; Furrer and Corre- stage 27 and the ventral view of the U-shaped dor 2008; Marín Gómez and Gómez Hoyos larval jaw sheaths at stage 28 were published 2011; Rueda-Almonacid et al. 2004; Stuart et by Sánchez (2013). Color illustrations of an al. 2008; Suárez-Mayorga 2004), development adult individual with red stripes was present- of embryos and tadpoles (Rodríguez-Suárez ed by Amézquita et al. (2013), Arcila-Pérez et and Corredor-Londoño 2012; Sánchez 2013), al. (in press), and Brown et al. (2011); color diet (Gómez-Hoyos 2010; Gómez-Hoyos et illustrations of red and yellow morphs were al. 2014; Herrmann 2005), geographic dis- published by Casas-Cardona et al. (2018). tribution (Acosta-Galvis 2000; Castro-Her- A black-and-white illustration of tadpole rera and Bolívar-García 2010; Ruiz-Carranza mouth parts was presented by Walls (1994) et al. 1996; Suárez-Mayorga 2004; Vargas-Sa- and an illustration of the mouth of a tadpole linas et al. 2014b), habitat use and acoustic at stage 25 was published by Myers and Daly communication (Vanegas-Guerrero et al. (1980). 2016; Vargas-Salinas and Amézquita 2013; Vargas-Salinas et al. 2014a), homing behav- DISTRIBUTION. Andinobates bombetes is ior (Arcila-Pérez et al. in press), husbandry found in leaf litter or under fallen logs lo- (Herrmann 2005), natural history (Hessel- cated along the forest edge and in the forest haus 1994; Mattison 1987, 2002; Walls 1994), interior, often adjacent to streams.. Although pet trade (Anonymous 1993a, 1993b), pop- populations in humid montane forests can be ulation ecology (Gómez-Hoyos 2010), skin found away from streams, some populations toxins (Daly 1988; Daly et al. 1987, 1993; established in tropical dry forests are found Daly et al. 1997; Erspamer 1994; Myers and alongside small streams. This species is en- Daly 1976, 1980; Myers et al. 1995), and tax- demic to Colombia and has been found be- onomy (Brown et al. 2011; Grant et al. 2006; tween 867–2100 m elevation on both slopes Ruiz-Carranza and Ramírez-Pinilla 1992; Sil- of the Western Andes and on the western verstone 1975). slope of the Central Andes in the departments of Quindío, Risaralda, and Valle del Cauca ETYMOLOGY. The specific epithet bombe- (Ruiz-Carranza et al. 1996; Súarez-Mayorga tes refers to the loud buzzing sounds made 2004; Vargas-Salinas et al. 2014b). by calling males, similar to those made by insects. The specific epithet is Latinized from FOSSIL RECORD. None. the Greek βομβητηϛ meaning ‘buzzer’ (Myers and Daly 1980). PERTINENT LITERATURE. Published ref- erences to the species are listed by topic: COMMENTS. Two studies of Andinobates advertisement call (Myers and Daly 1980; bombetes are being performed by students Myers 1982), antipredatory behavior in at the University of Quindío, Colombia. One adults (Casas-Cardona et al. 2018), breeding study is focused on the intraspecific geo- behavior and parental care (Herkrath 2006; graphic variation of temporal and spectral Hesselhaus 1994; Suárez-Mayorga 1999), features of the advertisement call, including checklists (Anonymous 1993a; Castro-Her- comparisons between morphs (i.e., popula- rera and Vargas-Salinas 2008; Duellman 1993; tions whose individuals exhibit red vs. yel- Frank and Ramus 1995; Frost 1985; Glaw et al. low dorsolateral stripes) and populations 10 separated by geographic barriers (i.e., low- von Humboldt. Unpaginated foldout lands between Western and Central Andes) sheet [8 pp.]. [No publication date print- (C. Conzález-Acosta, unpublished data). A ed on sheet; date provided by Instituto second study examines how individuals deal Humbolt]. with acoustic anthropogenic disturbance in Amézquita, A., R. Márquez, R. Medina, D. their habitat; specifically, whether males have Mejía-Vargas, T. R. Kahn, G. Suárez, and the capacity to increase the frequency of the L. Mazariegos. 2013. A new species of advertisement call when they are exposed to Andean poison frog, Andinobates (An- noise by traffic on roads (G. M. Jiménez-Var- ura: Dendrobatidae), from the north- gas, unpublished data). western Andes of Colombia. Zootaxa 3620(1):163–178. ADDITIONAL VERNACULAR NAMES. Anonymous. 1993a. Dendrobatid species. Cauca-Baumsteiger (Herrmann 2005), Cauca American Dendrobatid Group Newslet- Poison Frog (Suárez-Mayorga et al. 2016; Wro- ter 8:3–6. bel 2004), Cauca Poison-arrow Frog (Wrobel Anonymous. 1993b. Adds: for sale. American 2004), Elevated Fondle-shun (Mitchell 2017), Dendrobatid Group Newsletter 10:2–3. rana rubí (Castro-Herrera and Bolívar-García Arcila-Pérez, L. F., M. A. Atehortua-Vallejo, 2010; Herkrath 2006), Rana venenosa del and F. Vargas-Salinas. In press. Hom- Calima (Amézquita and Vélez [2010]), rana ing in the rubí poison frog Andinobates venenosa del Cauca (Suárez-Mayorga 2004), bombetes. Copeia. and Red-banded Poison Arrow Frog (as Bernal, M. H. and V. F. Luna-Mora. 2014. An- Краснополосый Древолаз) (Sokolov 1988). dinobates tolimensis. Catálogo de Anfibi- os y Reptiles de Colombia 2(1):1–4. ACKNOWLEDGMENTS. We thank the Bi- Bernal, M. H., V. F. Luna-Mora, O. Gallego, ology program of the University of Quindío, and A. Quevedo. 2007. A new species of Colombia, for logistic support during the poison frog (Amphibia: Dendrobatidae) preparation of this manuscript. We thank from the Andean mountains of Tolima, Dina Lucia Rivera for her help with building Colombia. Zootaxa 1638:59–68. the distribution map. Thanks to J. D. Lynch Brown, J. L., E. Twomey, A. Amézquita, M. B. for allowing access to tadpole vouchers in the de Souza, J. P. Caldwell, S. Lötters, R. von Collection of Amphibians of the ICN, Bogotá, May, P. R. Melo-Sampaio, D. Mejía-Var- Colombia. Finally, we also give thanks to the gas, P. Perez-Peña, M. Pepper, E. H. Poel- Asociación Colombiana de Herpetología man, M. Sanchez-Rodriguez, and K. ACH (Grant Botas al Campo BC-2018-03) Summers. 2011. A taxonomic revision of for economic support for our studies with the Neotropical poison frog genus Ran- Andinobates bombetes. itomeya (Amphibia: Dendrobatidae). Zootaxa 3083:1–120. LITERATURE CITED Casas-Cardona, S., R. Márquez, and F. Var- Acosta-Galvis, A. R. 2000. Ranas, salaman- gas-Salinas. 2018. Different colour dras y caecilias (Tetrapoda: Amphibia) morphs of the poison frog Andinobates de Colombia. Biota Colombiana 1:289– bombetes (Dendrobatidae) are similar- 319. ly effective visual predator deterrents. Amézquita, A. and A. Vélez M. [2010]. Ra- Ethology 124:245–255. nas Venenosas de Colombia. Serie Espe- Castaño, J. H. and J. A. Carranza-Quiceno. cies Colombianas 6. Instituto de Investi- 2015. CampoAlegre. 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Suárez-Mayorga, A. M. 1999. Comportamien- Vargas-Salinas, F., C. L. Rodríguez-Collazos, to reproductivo de Minyobates bombetes and M. A. Suárez-Mayorga. 2014b. Andi- (Amphibia: Anura: Dendrobatidae). Un- nobates bombetes. Catálogo de Anfibios y published undergraduate thesis, Depar- Reptiles de Colombia 2(2):13–18. tamento de Biología, Facultad de Cien- Walls, J. G. 1994. Jewels of the Rainforest – cias, Universidad Nacional de Colombia, Poison Frogs of the Family Dendrobati- Bogotá Colombia. ix + 136 pp. dae. T. F. H. Publications, Inc., Neptune Suárez-Mayorga, Á. M. 2004. Rana veneno- City, New Jersey. 288 pp. sa del Cauca Dendrobates bombetes. Pp. Wrobel, M., Compiler. 2004. Elsevier’s Dic- 302–307 in Libro Rojo de los Anfibios de tionary of Amphibians in Latin, English, Colombia (J. V. Rueda-Almonacid, J. D. German, French and Italian. Elsevier B. Lynch, and A. Amézquita, editores). Se- V., Amsterdam, The Netherlands. ix + rie Libros Rojos de Especies Amenazadas 396 pp. de Colombia. Conservación Internacio- nal Colombia, Instituto de Ciencias Na- See page 16 for author affiliations and turales – Universidad Nacional de Co- contact information. lombia, Ministerio del Medio Ambiente, Bogotá, Colombia. Suárez-Mayorga Á. M., T. R. Kahn, and D. A. Gómez Hoyos. 2016. Cauca Poison Frog. Andinobates bombetes (Myers and Daly, 1980). Pp. 251–256 in Aposematic Poison Frogs (Dendrobatidae) of the An- dean Countries: Bolivia, Colombia, Ec- uador, Perú and Venezuela (T. R. Kahn, E. La Marca., S. Lötters, J. L Brown, E. Twomey, and A. Amézquita [editors]). Conservation International Tropical Field Guide Series, Conservation Inter- national, Arlington, Virginia. Vanegas-Guerrero, J., C. Fernández, W. Bui- trago-González, and F. Vargas-Salinas. 2016. Urban remnant forests: Are they important for herpetofaunal conservation in the central Andes of Colombia? Herpetological Review 47:180–185. Vargas-Salinas, F. and A. Amézquita. 2013. Traffic noise correlates with calling time but not spatial distribution in the threatened poison frog Andinobates bombetes. Behaviour 150:569–584. Vargas-Salinas, F., A. Dorado-Correa, and A. Amézquita. 2014a. Microclimate and stream noise predict geographic diver- gence in the auditory signal of a threatened poison frog. Biotropica 46:748-755. 16

FERNANDO VARGAS-SALINAS, Pro- grama de Biología, Universidad del Quindío, Carrera 15 Calle 12 N, Armenia, Colombia ([email protected]); MICHELLE A. ATEHORTUA-VALLEJO, Grupo de In- vestigación en Evolución, Ecología y Con- servación EECO. Universidad del Quindío, Carrera 15 Calle 12 N, Armenia, Colombia ([email protected]); LUISA F. ARCILA-PÉREZ, Grupo de Investigación en Evolución, Ecología y Conservación EECO. Universidad del Quindío, Carrera 15 Cal- le 12 N, Armenia, Colombia (arcilaluisap@ gmail.com); GINA M. JIMÉNEZ-VAR- GAS, Grupo de Investigación en Evolución, Ecología y Conservación EECO. Universidad del Quindío, Carrera 15 Calle 12 N, Arme- nia, Colombia (ginamarcelajimenez@gmail. com); CRISTIAN GONZÁLEZ-ACOS- TA, Grupo de Investigación en Evolución, Ecología y Conservación EECO. Universidad del Quindío, Carrera 15 Calle 12 N, Arme- nia, Colombia ([email protected]); SANTIAGO CASAS-CARDONA, Colec- ción de Anfibios y Reptiles de la Universidad del Quindío, Programa de Biología, Univer- sidad del Quindío, Carrera 15 Calle 12 N, Armenia, Colombia (santiagocasasc@gmail. com); and ALEJANDRO GRAJALES-ECH- EVERRY, Birding and Herping, Cra 6 #19-25 Apto 303 bloque 6, Torres del Río, Armenia, Colombia (alejandrobirdingandherping@ gmail.com).

Primary editors for this account, Christopher J. Bell and Travis J. LaDuc.