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908.1 AMPHIBIA: ANURA: BUFONIDAE Atelophryniscus, Atelophryniscus chrysophorus

Catalogue of American and Reptiles.

McCranie, J. R. 2017. Atelophryniscus, Atelophryniscus chrysophorus.

Atelophryniscus McCranie, Wilson, and Williams Quebrada de Oro

Atelophryniscus McCranie, Wilson, and Wil- liams 1989:2. Type species: Atelophrynis- cus chrysophorus McCranie, Wilson, and Williams (1989), by original designation Figure 1. Adult male (USNM 523121; SVL and by monotypy. 35.5 mm) of Atelophryniscus chrysophorus from Aelurophryniscus: Inger 1992:237. Lapsus. Quebrada de Oro, Atlántida, Honduras. Photo- Atelophrynus: Altig and Channing 1993:73. graphed by the author on 17 May 1988. Lapsus. : Pramuk and Lehr 2005:613. Chaunus: Frost et al. 2006:364. By implica- (interorbital distance/eye length in males = tion. 0.848–1.000, in females = 0.914–1.036) and Rhinella: Chaparro, Pramuk, and Gluesen- the tip of snout. The skin of the head is co-os- kamp 2007:211. By implication. sified with the underlying skull. The supra- orbital crests are low and confluent with the CONTENT. One species, Atelophryniscus short and low parietal crests. The postorbital chrysophorus, is recognized (see Com- crests are pustulose and low to indistinct. The ments). supratympanic crests are low and in contact with the parotoid glands. Canthal and preor- DESCRIPTION. Atelophryniscus is a small bital crests are absent to indistinct. The pre- bufonid (in ten adult males, snout-vent tympanic crests are pustulose and low to in- length [SVL] range = 35.4–40.1 mm, mean distinct. The parotid crests are subtriangular [standard deviation] = 37.8 mm [± 1.6 mm]; and slightly larger than, to about equal to, the in seven adult females, SVL range = 49.4– size of the prominent upper eyelids (upper 59.6 mm, mean [standard deviation] = 53.7 eyelid width/interorbital distance in males = mm [3.2 mm]). The head is relatively short 0.739–0.929, in females = 0.672–0.868), and and broad (head length/SVL in adult males = extend posteriorly to a level varying from 0.297–0.334, in adult females = 0.290–0.304; between the axillae to the anterior insertions head width/SVL in males = 0.319–0.359, in of forelimbs. Parotoid glands are usually dis- adult females = 0.303–0.327). The snout is tinct, occasionally indistinct, especially in nearly rounded in dorsal aspect and rounded females. Tympana are absent to indistinct in lateral profile. The top of the head is flat, (usually indistinct or absent in males, usually but slightly concave between the nostrils. The indistinct in females). The pupils are horizon- canthal ridges are angular and the loreal re- tally elliptical and the palpebral membranes gions are slightly concave. The nostrils are are translucent, with scattered silver flecks. directed laterally and are situated at a point The upper arms are more slender than slightly over one-half the distance between the moderately robust forearms. There is no the anterior borders of the prominent eyes transverse dermal fold on the upper surfac- 908.2

BELIZE 88° 86° 84°

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GUATEMALA HONDURAS

NICARAGUA 14°

EL SALVADOR

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Map. Distribution of Atelophryniscus and its only known species Atelophryniscus chrysophorus. Dot marks type locality and the open circle contacting the dot marks other nearby locality records. The second known area of occurrence is marked by the remaining open circle.

es of the wrists, nor dermal ridges along the The hind limbs are relatively long (shank posterior ventrolateral edges of the forearms, length/SVL in males = 0.461–0.510, in females although numerous tubercles are present = 0.419–0.464; foot length/SVL in males = along each forearm. The fingertips are nar- 0.427–0.496, in females = 0.428–0.464). The rowly expanded and there is no digital groove heels narrowly overlap when the hind limbs around the tips of the fingers. The subartic- are held together towards the cloacal open- ular tubercles on the fingers are round to ing at right angles to the body. The vertical ovoid, and globular to nearly conical. Super- dermal fold on the outer lateral edge of each numerary tubercles are absent to indistinct. heel is weak to absent. Numerous tubercles The palmar tubercles are elevated and round are present along the posterior edge of each to ovoid, and much larger than the thenar tarsus. A tuberculate inner tarsal fold extends tubercles. Numerous small accessory palmar about three-quarters the length of each tar- tubercles are present. The thenar tubercles are sus. The subarticular tubercles on the toes are round and barely elevated, not visible from round to ovoid and globular. Supernumerary above, and bear a patch of tiny nuptial excres- tubercles are absent to indistinct. The plantar ences in adult males. Each prepollex is not tubercles are small and indistinct. The inner enlarged, but an extensive thumb pad bearing metatarsal tubercles are elongate, elevated, tiny nuptial excresences is present in adult and visible from above. The outer metatarsal males. The first fingers are shorter than the tubercles are small, round to ovoid, and ele- second fingers; relative length of all fingers = vated. Relative toe length is I

I ⅓–1½ II ⅓–2 III ¾–3– IV 3–1– V. Lateral fleshy fringes are present on the unwebbed portions of the toes. The vent opening is directed posteroven- trally near upper level of thighs with skin around the vent strongly granular to tubercu- late. The skin of the dorsal surface of the body has numerous, scattered, small to moderate- ly-sized tubercles. The skin of the upper eye- lids has small to moderately-sized tubercles and that of the dorsal surfaces of the limbs has numerous small to moderately-sized tu- Figure 2. Amplectant pair of Atelophryniscus chrysophorus (male USNM 523141, SVL 37.9 bercles. A dorsolateral row of weakly to mod- mm; female USNM 523142, SVL 59.6 mm) from erately enlarged tubercles is usually present. Quebrada de Oro, Atlántida, Honduras. These The skin of the chin, throat, and chest is gran- and other adults collected 12–14 February 1995 ular to strongly granular and those of the bel- were the last adults of this genus and species seen ly and ventral surfaces of thighs are granular alive by a biologist. Photographed by the author to weakly granular. The tongue is elongately on 12 February 1995. ovoid, not notched posteriorly, and is attached for about 20% of its length (anteriorly). Vom- erine and maxillary teeth are absent and cho- anae are ovoid to ellipitical. Males have paired vocal slits and a single, median, vocal sac, but

Figure 3. Drawings of lateral (top) and dorsal (bottom) views of tadpole of Atelophryniscus chrysopho- rus (in lot KU 206748) in Gosner stage 35 (Gosner 1960). 908.4

Figure 5. Lateral view of living tadpole of Ate- lophryniscus chrysophorus (in lot KU 206750) in about Gosner stage 40–41 (Gosner 1960). Photo- graphed by the author at 790 m elevation on 16 August 1984.

Figure 4. Oral disc and ventral view of tad- pole of Atelophryniscus chrysophorus (in lot KU 206748) in Gosner stage 35 (Gosner 1960). that sac is not evident externally. Color in life of the male holotype (KU 206730) follows. The dorsal surfaces of the head and body were pale lime green with Figure 6. Dorsal view of living tadpole of Atelo- olive brown blotching and rusty-red-tipped phryniscus chrysophorus (in lot KU 206750). Gos- ner stage (Gosner 1960) unknown. Photographed pustules. Pale orange pustules outlined the by the author at 790 m elevation on 16 August lower edge of a pale lime-green lateral stripe. 1984. That lateral stripe was bordered below by a dark brown line grading to olive brown ven- trally. The side of head was pale yellow, mot- and the total length was 26.4. The body is de- tled with dark brown in front of and below pressed and 1.5 times wider than high. The the eye. A rusty-red, dark brown-bordered snout is broadly semicircular in dorsal aspect bar extended from each eye to the posteri- and broadly rounded in lateral profile. The or edges of the lips, that bar also with olive eyes are small (eye length / body length about brown blotches. Each iris was pale metalic 0.09), and directed anterolaterally. The inter- green. The limbs were a pale lime-green dor- orbital distance is about 1.2. The nostrils are sally and the toe tips and outer edges of the situated considerably closer to the eyes than webs were orange. The ventral surface of the the tip of snout, and directed anterolaterally. body was pale blue-green, mottled with dark The spiracle is sinistral, directed posterodor- chocolate brown. Color notes of other adults sally, and situated low on the body at a point and subadults were published by McCranie about three-fourths distance from the tip and Wilson (2002). of snout to posterior end of body. The vent A typical tadpole (KU lot 206748) in Gos- tube is medial. The tail musculature is robust, ner stage 37 (Gosner 1960) had the following extending nearly to the tip of the rounded measurements (all measurements in mm, tail. The dorsal fin is slightly arched, highest ratios based on those measurements). The at about midlength of tail, and terminating body length was 9.6, the tail length was 16.8, posterior to body. The ventral fin is not -no 908.5 ticeably arched. The height of the tail muscu- gastromyzophorous tadpole of Atelophry- lature is greater than the height of either the niscus is drastically different from the lentic dorsal or ventral fins at midlength of the tail. water larvae of the remaining Honduran bu- The oral disc is huge (oral disc width / snout fonid genera in having a raised rim laterally width about 1.04) and ventral. Large dorsal and posteriorly, with the rim extending about and ventral gaps are present in the marginal three-fourths the length of the body. papillae, with two rows of marginal papillae present lateral to the dorsal gap (about 10–12 PUBLISHED DESCRIPTIONS, ILLUS- papillae present per mm). The remainder of TRATIONS, DISTRIBUTION, FOSSIL RE- the lateral portion of the oral disc is bordered CORD, AND PERTINENT LITERATURE. by a single row of submarginal papillae. The See species account. oral disc is emarginated laterally. The kerati- nized jaw sheaths are narrow and lack serra- REMARKS. Adults of the monotypic genus tions. The upper jaw sheath is widely arched, Atelophryniscus were last seen in the vicinity with short, lateral processes. The lower jaw of its type locality during February 1995 and sheath is widely V-shaped. The labial tooth tadpoles and a recent metamorph (SVL 12 rows are 2 / 3, all complete, long, subequal, mm) were last seen there during June 1996. and extending to the lateral portion of the The author made unsuccessful subsequent oral disc. The gastromyzophorous suctorial searches in the vicinity of the type locality disc has a raised rim laterally and posteriorly, during February 2003 and June 2005. Neither with the rim extending about three-fourths adults nor tadpoles could be found at its sec- the length of body. ond known locality in the Texíguat region at The color in life of tadpoles in KU lot the same stream as the type locality of Isth- 206748 follow. The dorsal and lateral surfaces mohyla insolita (McCranie, Wilson, and Wil- of the body were black with gold flecking and/ liams 1993; only at a slightly lower elevation or spotting. The tail musculature was black of 1450 m [versus 1550 m]) of the western with pale yellow bands or spots. The tail fins portion of the Cordillera Nombre de Dios on were clear with the dorsal fin lightly flecked three and four day trips during August 1991, with black (all of the above data were taken by August 1993, July 1995, February 2003, July the author and were published by McCranie 2004, and April 2010. Thus, the genus and its et al. 1989, McCranie and Castañeda 2007, only known species are thought to be extinct. and McCranie and Wilson 2002; also see Mc- McCranie and Castañeda (2005, 2007) also Cranie et al. 1989 for color notes on older tad- reached that same grim conclusion (also see poles and metamorphosing individuals). Comments for the species Atelophryniscus chrysophorus). DIAGNOSIS. The combination of having parotoid glands and cranial crests and the ETYMOLOGY. Atelophryniscus is formed absence of maxillary and vomerine teeth from the Greek prefixa - (without), the Greek distinguishes Atelophryniscus, and its single words telo (complete), and phryne (), species, Atelophryniscus chrysophorus, from and the Greek diminuative suffixiskos – . The all other anurans in Honduras, except for the name alludes to the resemblance of the tad- remaining bufonid genera (, Rhinel- pole of this genus with those of the bufonid la, and Rhaebo). Atelophryniscus differs from genus Atelopus and the resemblence of the those genera in Honduras in having the first adult to various species placed in the bufonid finger shorter than the second finger, and in toad genus Bufo at the time of its description, lacking a tympanum, or having an indistinct and to the relatively small size of its adult. The tympanum. Also, the lotic water-dwelling gender is masculine. 908.6

COMMENTS. Pramuk and Lehr (2005:613) 237. Lapsus. thought Atelophryniscus was closely enough Atelophrynus chrysophorus: Altig and Chan- related to Bufo (= Rhinella) veraguensis that ning 1993:73. Lapsus. they placed Atelophryniscus in the synonymy Atelophryniscus chrysophorous: Coloma and of Bufo (= Rhinella). It would defy all bio- Lötters 1996:69. Lapsus. geographic knowledge gained over the years Bufo chrysophorus: Pramuk and Lehr 2005: about amphibians and reptiles of Honduras 615. to think that a small endemic toad from an Chaunus chrysophorus: Frost 2007: published isolated, montane endemic hotspot in Hon- online. By implication duras would be most closely related to a Rhinella chrysophora: Chaparro, Pramuk, and group of toads otherwise confined “along the Gluesenkamp 2007:211. By implication. Andes from Peru to northwestern Argentina” Rhinella chrysophora: Wilson and Johnson (Haad et al. 2014:184), thousands of miles to 2010:110. First use of combination in print. the south. Similar views were expressed by Köhler (2011) and McCranie and Castañeda CONTENT. No subspecies are recognized. (2007). McCranie et al. (1989) discussed the relationships of Atelophryniscus with other DESCRIPTION AND DIAGNOSIS. See ge- bufonid genera recognized at that time and neric account. Señaris et al. (1994) briefly touched on those same relationships. Because of publication PUBLISHED DESCRIPTIONS. Detailed dates, Frost et al. (2006) did not evaluate the descriptions of the external morphology of taxonomic decisions made by Pramuk and the adult and/or tadpole, based on data gath- Lehr (2005) and listed Atelophryniscus among ered by the author, were published by McCra- the bufonid genera they considered valid; by nie et al. (1989), McCranie and Castañeda implication, the proposed by Frost (2007), and McCranie and Wilson (2002). et al. (2006) synonymized Atelophryniscus The chondrocranium and visceral skeleton of with Chaunus. Frost (2017) synonymized the the tadpole were described by Lavilla and de genus with Rhinella following Chaparro et al. Sá (2001). The pale bands on the tail of the (2007). tadpole was described by Altig and Channing (1993). Brief descriptions of some osteolog- Atelophryniscus chrysophorus McCranie, ical and myological characters of an adult Wilson, and Williams were provided by McCranie et al. (1989; also Quebrada de Oro Toad see Duellman and Schulte 1992), and detailed descriptions of some osteological characters Atelophryniscus chrysophorus McCranie, were published by Pramuk and Lehr (2005). Wilson, and Williams 1989:3. Type local- Basic conditions of inguinal fat bodies and ity, “Quebrada de Oro (15°38'N, 86°47'W), sternal morphology of Atelophryniscus chrys- elevation 880 m, tributary of Río Viejo, ophorus were provided by da Silva and Men- south slope of Cerro Búfalo, Cordillera de delson (1999). Nombre de Dios, Depto. Atlántida, Hon- duras.” Holotype, University of Kansas, ILLUSTRATIONS. Color photographs of Museum of Natural History, Lawrence an adult were published by Köhler (2011). (KU) 206730, an adult male, collected 16 McCranie and Wilson (2002), McCranie and August 1982 by J. R. McCranie, K. L. Wil- Castañeda (2007), Stuart et al. (2008), and liams, and L. D. Wilson (examined by au- Wilson and McCranie (2004a). Black-and- thor). white photographs of the adult were present- Aelurophryniscus chrysophorus: Inger 1992: ed by McCranie et al. (1989) and Pramuk and 908.7

Lehr (2005); a black-and-white photograph and McDiarmid 1999a, 1999b; Haad et al. of a juvenile and a tadpole were published by 2014; Hoff et al. 1999; Kaplan 1997; Lehr et Pramuk and Lehr (2005). Black-and-white al. 2005; Lötters 1996; McCranie and Wilson illustrations of the tadpole were published 2000; McDiarmid and Altig 1999; Pereyra et by McCranie et al. (1989), McCranie and al. 2015; Wells 2007), taxonomy, systematics, Castañeda (2007), McCranie and Wilson and phylogenetics (Altig and McDiarmid (2002), and Wilson and McCranie (1993). 1999b; Chaparro et al. 2007; Frost 2007, 2017; Black-and-white illustrations of the adult Frost et al. 2006.; McCranie and Castañeda skull and vertebral column were presented by 2007; McCranie and Wilson 2002; Pramuk Pramuk and Lehr (2005) and the chondro- and Lehr 2005). cranium and hyobranchial skeletons of the tadpole were published by Lavilla and de Sá REMARKS. McCranie et al. (1989) discussed (2001). that the tadpole of Atelophryniscus chrysopho- rus had an amazing resemblence to that of an- DISTRIBUTION. Atelophryniscus chryso- other bufonid genus, Atelopus, and suggested phorus is known to occur from 750 to 1760 m that resemblence was a result of convergent elevation in the Premontane Wet Forest and adaptations to life in fast-flowing streams. Lower Montane Wet Forest formations (see Subsequently, Coloma and Lötters (1996:69) Holdridge 1967) in the central and western wrote that the term convergent adaptation portions of the Cordillera Nombre de Dios, “is unwarranted in the absence of a detailed Honduras (but see Remarks for genus). analysis of bufonid relationships.” Solís et al. (2014:124) wrote “Townsend FOSSIL RECORD. None. and Wilson (2010) listed this species [A. chrys- ophorus] as Rhinella chrysophora, based on PERTINENT LITERATURE. Relevant cita- the opinion of Chaparro et al. (2007).” Chap- tions are listed by topic: checklists and fau- arro et al. (2007) did not specifically mention nal lists (Anonymous 2016; Campbell 1999; Atelophryniscus chrysophorus (nor Rhinella Duellman 1993, 2001; Glaw et al. 1998, 2000a, chrysophora), but instead only implied Rhi- 2000b; Hero et al. 2012; Hutchins et al. 2003; nella included Atelophryniscus chrysophorus McCranie 2006, 2007, 2009, 2015; McCranie by using “R. veraguensis group.” Pramuk and and Castañeda 2005, 2007; McCranie and Lehr (2005) concluded Atelophryniscus chrys- Solís 2013; McCranie and Wilson 2002; Mc- ophorus was closely related to the Rhinella Cranie et al. 2006; Solís et al. 2014; Townsend veraguensis group. Also, Wilson and Johnson and Wilson 2010, 2016; Wilson and Johnson (2010) previously used the new combination 2010; Wilson and McCranie 1994, 2004b; Rhinella chrysophora. Wilson and Townsend 2006, 2010; Wilson et This species was included in the 2012 list al. 2001; Young et al. 2004a, 2004b), compar- of endemic fauna of the Reserva de Vida Sil- isons to new species (Graybeal and Canatella vestre Texíguat (Townsend et al. 2012), but 1995; Lehr et al. 2005), conservation status not in the 2010 list (Townsend et al. 2010). (Johnson et al. 2015; McCranie and Wilson 2004; Stuart et al. 2008; Townsend et al. 2012; ADDITIONAL VERNACULAR NAMES. Wilson and McCranie 1998, 2004a, 2004b; The names Rio Viejo Toad and Rio Viejo Wilson and Townsend 2006, 2010;), tadpole Toads were published by Frank and Ramus keys (McCranie and Castañeda 2007; McCra- (1995), although this toad does not occur nie and Wilson 2002; Wilson and McCranie along that river, but instead along an upper 1993), tadpole morphology/gastromyzo- tributary of that river. Stream Toads and Río phorous larvae (Aguayo et al. 2009; Altig Viejo Toad were listed for the genus and spe- 908.8 cies, respectively, by Wrobel (2004). destruction, caused much larger land- slides, some of which devastated parts of the ETYMOLOGY. The namechrysophorus is de- Quebrada de Oro and the small stream at rived from the Greek noun chrysus (gold) and the second known locality of Atelophryniscus the Greek suffix –phora (bearing). The refer- chrysophorus in the Texíguat region, Yoro. As ence is to the gold markings on the otherwise a result of all of those stream-altering land- black tadpole, to the name of the stream at slides, much of the Quebrada de Oro and the the type locality (Quebrada de Oro), and the small stream at the second known locality fact that the gold-spotted tadpole used to oc- of Atelophryniscus are no longer fit for this cur abundantly in that stream. One can easily species. The slight hope for remaining pop- believe the name of the stream, Quebrada de ulations of Atelophryniscus would most likely Oro, was founded because of the numerous be on the remaining forested north slopes of and obvious distinctly gold-spotted tadpoles the Cordillera Nombre de Dios that have not of Atelophryniscus chrysophorus that were so been damaged by landslides. Unfortunately, obvious during the first few years of my work the majority of those slopes are so steep that in that area before the population crashed few permanent streams exist above about 900 (also see McCranie and Castañeda 2005:10). m elevation.

COMMENTS. The last specimens ofAte - ACKNOWLEDGMENTS. Many thanks to lophryniscus chrysophorus were seen alive Alexander Gutsche for his valuable assistance during 1996, before the author began collect- in acquiring several needed publications. ing tissues. Thus, no tissues are available from Steve W. Gotte provided the measurement for this taxon for molecular analyses. The single the recent metamorph. phylogenetic analysis performed for this spe- cies used only morphological data (Pramuk LITERATURE CITED and Lehr 2005), and resulted in a question- able taxonomic decision. Aguayo, R., E. O. Lavilla, M. F. Vera Candioti, occurring at the type and T. Camacho. 2009. Living in fast-flow- locality of Atelophryniscus chrysophorus ing water: Morphology of the gastromyzo- along the Quebrada de Oro, Atlántida was phorous tadpole of the bufonid Rhinella discussed by McCranie and Castañeda (2005, quechua (R. veraguensis group). Journal 2007), McCranie and Wilson (2002), and of Morphology 270:1431–1442. Wilson and McCranie (1998). Destruction of Altig, R. and A. Channing. 1993. Hypothesis: the forest by humans played a significant role Functional significance of colour and pat- in facilitating landslides that destroyed the tern of anuran tadpoles. The Herpetolog- portions of the Quebrada de Oro in which ical Journal 3:73–75. both adults and larva of Atelophryniscus Altig, R. and R. W. McDiarmid. 1999a. Body chrysophorus lived. Further complicating that plan. Development and morphology. Pp. habitat destruction was the documentation 24–51 in Tadpoles. The Biology of Anu- of the chytrid fungus Batrachochytrium den- ran Larvae (R. W. McDiarmid and R. Al- drobatidis in two of the four anuran species tig, editors). The University of Chicago collected along the Quebrada de Oro in 2003 Press, Chicago, Illinois. (Puschendorf et al. 2006). This chytrid fungus Altig, R. and R. W. McDiarmid. 1999b. Di- has been linked to anuran declines in other versity. Familial and generic characteriza- neotropical regions (Puschendorf et al. 2006). tions. Pp. 295–337 in Tadpoles. The Biol- During October 1998, the torrential rains ogy of Anuran Larvae (R. W. McDiarmid from Hurricane Mitch, aided by previous and R. Altig, editors). The University of 908.9

Chicago Press, Chicago, Illinois. World: An Online Reference. Version Anonymous. 2016. Herpetofaunal list for Me- 5.0. American Museum of Natural Histo- soamerica. Mesoamerican Herpetology ry, New York, New York. [Cited in Frost (www.mesoamericanherpetology.com/ 2017. Not seen; no on-line archive of da- taxonomic-list.html; last assessed 5 May tabase is available]. 2017). Frost, D. R. 2017. Rhinella chrysophora (Mc- Campbell, J. A. 1999. Distribution patterns Cranie, Wilson, and Williams, 1989) | of amphibians in Middle America. Pp. Species of the World: An 111–210 in Patterns of Distribution of Online Reference. Version 6.0. American Amphibians. A Global Perspective (W. Museum of Natural History, New York, E. Duellman, editor). The John Hopkins New York. Available at http://research. University Press, Baltimore, Maryland. amnh.org/vz/herpetology/amphibia/ Chaparro, J. C., J. B. Pramuk, and A. G. Amphibia/Anura/Bufonidae/Rhinella/ Gluesenkamp. 2007. A new species of ar- Rhinella-chrysophora. Archived by Web- boreal Rhinella (Anura: Bufonidae) from Cite at http://www.webcitation.org/6qIQ- cloud forest of southeastern Peru. Herpe- vAt4g on 7 May 2017. tologica 63:203–212. Frost, D. R., T. Grant, J. Faivovich, R. H. Bain, Coloma, L. A. and S. Lötters. 1996. The tad- A. Hass, C. F. B. Haddad, R. O. de Sá, A. pole of Atelopus balios (Anura: Bufonidae) Channing, M. Wilkinson, S. G. Donnel- from the Pacific lowlands of Ecuador. lan, C. J. Raxworthy, J. A. Campbell, B. Herpetologica 52:66–70. L. Blotto, P. Moler, R. C. Drewes, R. A. da Silva, H. R. and J. R. Mendelson, III. 1999. Nussbaum, J. D. Lynch, D. M. Green, and A new organ and sternal morphology in W. C. Wheeler. 2006. The amphibian tree toads (Anura: Bufonidae): Descriptions, of life. Bulletin of the American Museum taxonomic distribution, and evolution. of Natural History 297:1–370, + foldout Herpetologica 55:114–126. phylogenetic tree. Duellman, W. E. 1993. Amphibian species of Glaw, F., J. Köhler, R. Hofrichter, and A. Du- the world: Additions and corrections. The bois. 1998. Systematik der Amphibien: University of Kansas Museum of Natu- Liste der rezenten Familien, Gattungen ral History, Special Publication 21:i–iii, und Arten. Pp. 252-258 in Amphibien. 1–372. Evolution, Anatomie, Physiologie, Ökol- Duellman, W. E. 2001. The Hylid of ogie und Verbreitung, Verhalten, Bed- Middle America. Volume 2. Society for rohung und Gefährdung (R. Hofrichter, the Study of Amphibians and Reptiles, editor). Naturbuch Verlag, Augsburg, Contributions to Herpetology 18:v–x, Germany. 695–1158, plates 1–92. Glaw, F., J. Köhler, R. Hofrichter, and A. Du- Duellman, W. E. and R. Schulte. 1992. De- bois. 2000a. Amphibian Systematics: List scription of a new species of Bufo from of recent families, genera, and species. northern Peru with comments on phenet- Pp. 252-258 in Amphibians: The World ic groups of South American toads (Anu- of Frogs, Toads, Salamanders, and Newts. ra: Bufonidae). Copeia 1992:162–172. (R. Hofrichter, editor). Firefly Books Frank, N. and E. Ramus. 1995. A Complete (U.S.), Inc., Buffalo, New York. Guide to Scientific and Common Names Glaw, F., J. Köhler, R. Hofrichter, and A. Du- of Reptiles and Amphibians of the World. bois. 2000b. Amphibian Systematics: List N G Publishing Inc., Pottsville, Pennsyl- of recent families, genera, and species. Pp. vania. 377 pp. 252-258 in The Encyclopedia of Amphib- Frost, D. R. 2007. Amphibian Species of the ians (R. Hofrichter, editor). Key Porter 908.10

Books Limited, Toronto, Ontario, Cana- an Society 105:225–237. da. Johnson, J. D., V. Mata-Silva, and L. D. Wil- Gosner, K. L. 1960. A simplified table for son. 2015. A conservation reassessment staging anuran embryos and larvae with of the Central American herpetofauna notes on identification. Herpetologica based on the EVS measure. Amphibian & 16:183–190. Reptile Conservation 9(2)[General Sec- Graybeal A. and D. C. Cannatella. 1995. A tion]:1–94(e100). new taxon of Bufonidae from Peru, with Kaplan, M. 1997. Internal and external anato- descriptions of two new species and a my of the abdominal disc of Atelopus (Bu- review of the phylogenetic status of su- fonidae) larvae. Caldasia 19:61–69. praspecific bufonid taxa. Herpetologica Köhler, G. 2011. Amphibians of Central 51:105–131. America. Herpeton, Verlag Elke Köhler, Haad, M. B., F. Vera Candioti, and D. Baldo. Offenbach, Germany. 378 pp. 2014. The stream tadpoles ofRhinella Lavilla, E. O. and R. de Sá. 2001. Chondocra- rumbolli (Anura: Bufonidae). Herpeto- nium and visceral skeleton of Atelopus logica 70:184–197. tricolor and Atelophryniscus chrysophorus Hero, J.-M., C, Morrison, J. Chanson, S. Stu- tadpoles (Anura, Bufonidae). Amphib- art, and N. A. Cox. 2012. Phylogenetic ia-Reptilia 22:167–177. correlates of population decline and ex- Lehr, E., J. B. Pramuk, and M. Lundberg. tinction risk in amphibians. Pp. 3539- 2005. A new species of Bufo (Anura: Bu- 3551 in Amphibian Biology. Volume 10. fonidae) from Andean Peru. Herpetolog- Conservation and Decline of Amphib- ica 61:308–318. ians: Ecological Aspects, Effect of Hu- Lötters, S. 1996. The Neotropical Toad Genus mans, and Management (H. Heatwole Atelopus. Checklist — Biology — Distri- and J. W. Wilkinson, editors). Surrey Be- bution. M. Vences & F. Glaw Verlags GbR, atty & Sons Pty Limited, Baulkham Hills, Köln, Germany. 143 pp., 70 plates. New South Wales, Australia. McCranie, J. R. 2006. Specimen Locality Hoff, K. vS., A. R. Blaustein, R. W. McDiar- Data & Museum Numbers / Ubicación y mid, and R. Altig. 1999. Behavior. Inter- Números de Museo de los Especímenes, actions and their consequences. Pp. 215– Información Complementaria for/a la 239 in Tadpoles. The Biology of Anuran “Guía de Campo de los Anfibios de Hon- Larvae (R. W. McDiarmid and R. Altig, duras” by/por James R. McCranie and editors). The University of Chicago Press, Franklin E. Castañeda. Smithsonian Her- Chicago, Illinois. petological Information Service 137:1–39. Holdridge, L. R. 1967. Life Zone Ecology. Re- McCranie, J. R. 2007. Distribution of the am- vised Edition. Tropical Science Center, phibians of Honduras by departments. San José, Costa Rica. 206 pp. Herpetological Review 38:35–39. Hutchins, M., W. E. Duellman, and N. McCranie, J. R. 2009. Amphibians and rep- Schlager (editors). 2003. Grizmek’s Ani- tiles of Honduras. Listas Zoológicas Actu- mal Life Encyclopedia. Second Edition. alizadas UCR. Museo de Zoología UCR, Volume 6. Amphibians. The Gale Group, San Pedro, Costa Rica. 20 pp. http://mu- Inc., Farmington Hills, Michigan. xvi + seo.biologia.ucr.ac.cr/Listas/Anteriores/ 507 pp. HerpHon.htm (last accessed 1 May 2017). Inger, R. F. 1992. Variation of apomorphic McCranie, J. R. 2015. A checklist of the am- characters in stream-dwelling tadpoles phibians and reptiles of Honduras, with of the bufonid genus Ansonia (Amphibia: additions, comments on taxonomy, Anura). Zoological Journal of the Linne- some recent taxonomic decisions, and 908.11

areas of further studies needed. Zootaxa search. Materials and techniques. Pp. 3931:352–386. 7–23 in Tadpoles. The Biology of Anuran McCranie, J. R. and F. E. Castañeda. 2005. The Larvae (R. W. McDiarmid and R. Altig, herpetofauna of Parque Nacional Pico editors). The University of Chicago Press, Bonito, Honduras. Phyllomedusa 4:3–16. Chicago, Illinois. McCranie, J. R. and F. E. Castañeda. 2007. Pereyra, M. O., M. F. Vera Candioti, J. Guía de Campo de los Anfibios de Hon- Faivovich, and D. Baldo. 2015. Egg clutch duras. Bibliomania!, Salt Lake City, Utah. structure of Rhinella rumbolli (Anura: x + 304pp. + 147 plates. Bufonidae), a toad from the Yungas of McCranie, J. R. and J. M. Solís. 2013. Addi- Argentina, with a review of the repro- tions to the amphibians and reptiles of ductive diversity in Rhinella. Salamandra Parque Nacional Pico Bonito, Honduras, 51:161–170. with an updated nomenclatural list. Her- Pramuk, J. B. and E. Lehr. 2005. Taxonomic petology Notes 6:239–243. status of Atelophryniscus chrysophorus McCranie, J. R. and L. D. Wilson. 2000. A new McCranie, Wilson, and Williams, 1989 species of high-crested toad of the Bufo (Anura: Bufonidae) inferred from phylog- valliceps group from north-central Hon- eny. Journal of Herpetology 39:610–618. duras. Journal of Herpetology 34:21–31. Puschendorf, R., F. Castañeda, and J. R. Mc- McCranie, J. R. and L. D. Wilson. 2002. The Cranie. 2006. Chytridiomycosis in wild Amphibians of Honduras. Society for the frogs from Pico Bonito National Park, Study of Amphibians and Reptiles, Con- Honduras. EcoHealth 3:178–181. tributions to Herpetology 19. i–x + 625 Señaris, J. C., J. Ayarzagüena, and S. Gorzula. pp., 1–20 plates. 1994. Los sapos de la familia Bufonidae McCranie, J. R. and L. D. Wilson. 2004. The (Amphibia: Anura) de las tierras altas de Honduran amphibian fauna: Perched on la Guayana Venezolana: Descripcion de the brink of decline, Pp. 71–75 in Col- un nuevo genero y tres especies. Publi- lected DAPTF Working Group Reports: caciones de la Asociación de Amigos de Ten Years On (J. W. Wilkinson, editor). Doñana 3:1–37. DAPTF, Milton Keynes, United King- Solís, J. M., L. D. Wilson, and J. H. Townsend. dom. [Apparently mailed in 2006]. 2014. An updated list of the amphibians McCranie, J. R., L. D. Wilson, and K. L. Wil- and reptiles of Honduras, with comments liams. 1989. A new genus and species on their nomenclature. Mesoamerican of toad (Anura: Bufonidae) with an ex- Herpetology 1:122–144. traordinary stream-adapted tadpole from Stuart, S. N., M. Hoffmann, J. S. Chanson, northern Honduras. Occasional Papers of N. A. Cox, R. J. Berridge, P. Ramani, and the Museum of Natural History, The Uni- B. E. Young (editors). 2008. Threatened versity of Kansas 129:1–18. Amphibians of the World. Lynx Edicions, McCranie, J. R., L. D. Wilson, and K. L. Wil- Barcelona, Spain. xv + 758 pp. liams. 1993. New species of tree of the Townsend, J. H. and L. D. Wilson. 2010. Con- genus Hyla (Anura: Hylidae) from north- servation of the Honduran herpetofau- ern Honduras. Copeia 1993:1057–1062. na: Issues and imperatives. Pp. 460–487 McCranie, J. R., J. H. Townsend, and L. D. in Conservation of Mesoamerican Am- Wilson. 2006. The Amphibians and Rep- phibians and Reptiles (L. D. Wilson, J. H. tiles of the Honduran Mosquitia. Krieger Townsend, and J. D. Johnson, editors). Publishing Company, Malabar, Florida. Eagle Mountain Publishing, LC., Eagle i–x + 291 pp. Mountain, Utah. McDiarmid, R. W. and R. Altig. 1999. Re- Townsend, J. H. and L. D. Wilson. 2016. Am- 908.12

phibians of the Cordillera Nombre de conservation status of the herpetofau- Dios, Honduras: COI barcoding suggests na of Honduras. Amphibian and Reptile underestimated taxonomic richness in a Conservation 3:6–33. [Published online threatened endemic fauna. Mesoameri- in 2003]. can Herpetology 3:909–928. Wilson, L. D. and J. R. McCranie. 2004b. The Townsend, J. H., J. M. Butler, L. D. Wilson, herpetofauna of the cloud forests of Hon- and J. D. Austin. 2010. A distinctive new duras. Amphibian and Reptile Conserva- species of moss salamander (Caudata: tion 3:34–48. [Published online in 2003]. Plethodontidae: Nototriton) from an im- Wilson, L. D. and J. H. Townsend. 2006. The periled Honduran endemism hotspot. herpetofauna of the rainforests of Hondu- Zootaxa 2434:1–16. ras. Caribbean Journal of Science 42:88– Townsend, J. H., L. D. Wilson, M. Medi- 113. na-Flores, E. Aguilar-Urbina, B. K. Atkin- Wilson, L. D. and J. H. Townsend. 2010. The son, C. A. Cerrato-Mendoza, A. Contre- herpetofauna of Mesoamerica: Biodi- ras-Castro, L. N. Gray, L. A. Herrera-B., versity significance, conservational sta- I. R. Luque-Montes, M. McKewy-Mejía, tus, and future challenges. Pp. 760–812 A. Portillo-Avilez, A. L. Stubbs, and J. D. in Conservation of Mesoamerican Am- Austin. 2012. A premontane hotspot for phibians and Reptiles (L. D. Wilson, J. H. herpetological endemism on the wind- Townsend, and J. D. Johnson, editors). ward side of Refugio de Vida Silvestre Eagle Mountain Publishing, LC., Eagle Texíguat, Honduras. Salamandra 48:92– Mountain, Utah. 114. Wilson, L. D., J. R. McCranie, and M. R. Es- Wells, K. D. 2007. The Ecology and Behavior pinal. 2001. The ecogeography of the of Amphibians. The University of Chica- Honduran herpetofauna and the design go Press, Chicago, Illinois. xi + 1148 pp. of biotic reserves. Pp. 109–158 in Me- Wilson, L.D. and J. Johnson. 2010. Distri- soamerican Herpetology: Systematics, butional patterns of the herpetofauna of Zoogeography, and Conservation (J. D. Mesoamerica, a biodiversity hotspot. Pp. Johnson, R. G. Webb, and O. Flores-Vil- 30–235 in Conservation of Mesoameri- lela, editors). Centennial Museum, The can Amphibians and Reptiles (L. D. Wil- University of Texas at El Paso, Special son, J. H. Townsend, and J. D. Johnson, Publication 1. editors). Eagle Mountain Publishing, LC., Wrobel, M. (compiler). 2004. Elsevier’s Dic- Eagle Mountain, Utah. tionary of Amphibians in Latin, English, Wilson, L. D. and J. R. McCranie. 1993. Pre- German, French and Italian. Elsevier B. liminary key to the known tadpoles of V., Amsterdam, The Netherlands. ix + 396 anurans from Honduras. Royal Ontario pp. Museum, Life Sciences Occasional Paper Young, B. E., S. N. Stuart, J. S. Chanson, N. 40:1–12. A. Cox, and T. M. Boucher. 2004a. Disap- Wilson, L. D. and J. R. McCranie. 1994. Sec- pearing Jewels. The Status of New World ond update on the list of amphibians and Amphibians. NatureServe, Arlington, reptiles known from Honduras. Herpeto- Virginia. 54 pp. logical Review 25:146–150. Young, B. E., S. N. Stuart, J. S. Chanson, N. Wilson, L. D. and J. R. McCranie. 1998. Am- A. Cox, and T. M. Boucher. 2004b. Joyas phibian population decline in a Hondu- que Están Desapareciendo. El Estado de ran National Park. Froglog 25:1–2. los Anfibios en el Nuevo Mundo. Nature- Wilson, L. D. and J. R. McCranie. 2004a. The Serve, Arlington, Virginia. 54 pp. 908.13

JAMES R. MCCRANIE, 10770 SW 164 Street, Miami, FL 33157–2933 (jmccrani@ bellsouth.net).

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

Published 22 May 2017 and Copyright © 2017 by the Society for the Study of Amphibians and Reptiles.