1 AMPHIBIA: ANURA: fecundus

Catalogue of American and Reptiles 916.

McCranie, J. R. 2018. Craugastor fecundus.

Craugastor fecundus (McCranie and Wilson) Nombre de Dios Streamside

Eleutherodactylus milesi: Schmidt 1933:18 (in part). Eleutherodactylus rugulosus: Meyer 1969:156 Figure 1. Adult female (USNM 497098) of (in part). Craugastor fecundus from Quebrada de Oro, At- Eleutherodactylus milesi / Population System: lántida, Honduras. SVL 33.3 mm. Photograph McCranie et al. 1989:487 (in part). taken by James R. McCranie on 24 May 1988. Eleutherodactylus stadelmani: Campbell 1994:411 (in part). Eleutherodactylus fecundus McCranie and [± 0.8 mm]; in ten adult females, SVL range = Wilson 1997:162. Type locality, “from 29.8–37.3 mm, mean [standard deviation] = Quebrada de Oro (15°38'N, 86°47'W), ele- 33.5 mm [± 2.4 mm]) with a moderately long, vation 880 m, tributary of Río Viejo, south broad head (head length/SVL in adult males = slope of Cerro Búfalo S of La Ceiba, Cor- 0.400–0.462, in adult females = 0.379–0.419; dillera Nombre de Dios, Departamento de head width/SVL in adult males = 0.395–0.455, Atlántida, Honduras.” Holotype, Natural in adult females = 0.369–0.435). The snout is History Museum of Los Angeles County nearly rounded to rounded in dorsal aspect (LACM) 137311, an adult male, collected and rounded to nearly vertical in lateral pro- by J. R. McCranie, K. L. Williams, and L. file. The top of the head is flat. The canthal D. Wilson, 16 August 1982 (examined by ridges are angular and the loreal regions are author). concave. The upper lips are not flared. The Eleutherodactylus aurilegulus: Campbell and nostrils are directed posterolaterally and are Savage 2000:255 (in part). situated at a point slightly less than, to about Eleutherodactylus (Craugastor) fecundus: two-thirds the distance between the anterior Lynch 2000:150. borders of the eyes (interorbital distance/eye Craugastor fecundus: Savage 2002:551. By im- length in males = 0.636–0.867, in females = plication. 0.638–0.833) and the tip of the snout. Cra- Craugastor fecundus: Frost et al. 2006:360. nial crests are absent. The upper eyelids are First use of combination in print. prominent (eyelid width/interorbital distance Craugastor (Campbellius) fecundus: Hedges et in males = 1.040–1.318, in females = 1.031– al. 2008:34. 1.333). Tympana are indistinct or not visible externally in both sexes, tympana located CONTENT. No subspecies are recognized. posteroventral to the eyes and separated from the eyes (in five males, tympana-eye distance DESCRIPTION. Craugastor fecundus is a about one-half to two-thirds tympanum small eleutherodactyline (in ten adult males, length; in four females, distance slightly over snout-vent length [SVL] range = 21.1–23.5 three-fourths tympanum length). A well-de- mm, mean [standard deviation] = 22.2 mm veloped supratympanic fold is usually present 2

Map. Distribution of Craugastor fecundus. The open circle marks the type locality and the black dots represent additional localities.

on each side. about as large as, or slightly larger than the The upper arms are slightly more slen- thenar tubercles. A few small accessory pal- der than the moderately robust forearms. A mar tubercles are present. The thenar tubercle transverse dermal fold is absent on the up- on each hand is suboval, elevated, and visible per surfaces of the wrists. No vertical dermal from above. Each prepollex is not enlarged, folds are present along the elbows. Tubercles but nuptial thumb pads bearing granular, are absent, or tubercles arranged in an irreg- colorless nuptial excresences are present in ular series, or tubercles occasionally arranged adult males. Relative length of the fingers is in a linear row along the posterior ventro- I

cles. The tubercles on the flanks and posterior portion of the body are larger than those on the remainder of the dorsum. A row of raised skin, with or without tubercles, forms dis- continuous postocular ridges that extend to the level of the scapular region. Dorsolateral ridges are absent on the body. The skin on the chin, throat, and chest is smooth. The skin of the belly and ventral surfaces of the thighs is slightly wrinkled, almost smooth. The ventral Figure 2. Apparent hybrid between Craugastor disc is absent to indistinct in both sexes. The fecundus and Craugastor chrysozetetes (USNM pupil is horizontally ellipitical. The palpebral USNM 497114) from Quebrada de Oro, Atlánti- membrane is translucent and unpatterned. da, Honduras. An adult female with a SVL of 38.3 The tongue is ovoid, not notched posteriorly, mm. Photograph taken by James R. McCranie on and is free posteriorly for about one-fourth of 7 August 1989. its length. The vomerine tooth patches are on elevated, ovoid to nearly triangular-shaped absent. The subarticular tubercles on the toes ridges located posterormedially to round, are ovoid and globular. Supernumerary and tear-shaped, or ovoid choanae. Each tooth plantar tubercles are absent on the toes. Each patch is separated by a distance about equal inner metatarsal tubercle is elongate, elevat- to, or less than the width of either patch in ed, and visible from above. The outer meta- males, and by a distance much less than the tarsal tubercles are small, rounded, and ele- width of either patch in females. Maxillary vated. Relative toe length is I

ILLUSTRATIONS. Color photographs of made between 1980 and 1989, the last year an adult were presented by Köhler (2011), Craugastor fecundus was seen alive. Craugas- McCranie and Castañeda (2007), McCranie tor fecundus was seen on five of those six trips. and Wilson (2002), and Wilson and McCra- After the August 1989 trip, I did not return to nie (2004). A black-and-white photograph Quebrada de Oro until February 1995 and de- of an adult and a black-and-white drawing spite efforts targeting streamside Craugastor, of a hind foot were published by McCranie not a single streamside Craugastor was seen. and Wilson (1997). A color photograph Subsequent trips were made to the Quebrada of an apparent hybrid between Craugastor de Oro in 1996, 2003, and 2005, all without chrysozetetes and Craugastor fecundus was seeing a single streamside Craugastor. Thus, published by McCranie and Wilson (2002). sometime between August 1989 and Febru- ary 1995 the population of Craugastor fecun- DISTRIBUTION. Craugastor fecundus is dus at Quebrada de Oro completely crashed. known to occur from 200 to 1260 m elevation As a result, Craugastor fecundus is feared to be in the Lowland Moist Forest and Premontane extirpated from that, its best known, locality. Wet Forest formations (see Holdridge 1967) The in the Quebrada in the central and northeastern portions of de Oro region was discussed by McCranie the Cordillera Nombre de Dios, Honduras. (2017). Additionally, the chytrid fungus Ba- trachochytrium dendrobatidis was recorded in FOSSIL RECORD. None. two of the four anuran species that were col- lected along the Quebrada de Oro in 2003 by PERTINENT LITERATURE. Relevant cita- Puschendorf et al. (2006). This chytrid fungus tions are listed by topic: checklists and fau- has been linked to anuran declines in other nal lists (Anonymous 2016; Campbell 1999; neotropical regions (Puschendorf et al. 2006; Frost 2018; Glaw et al. 1998, 2000a, 2000b; also see recent summary by Whifield et al. Heinicke et al. 2007; Hutchins et al. 2003; Mc- 2016 and references cited therein) and likely Cranie 2006, 2007, 2009, 2015; McCranie and played a role in the extirpation of this species Castañeda 2005, 2007; McCranie and Solís at that locality. 2013; McCranie and Wilson 2002; McCranie The best chance for a surviving population et al. 2006; Padial et al. 2014; Solís et al. 2014; of Craugastor fecundus might be at its only Townsend and Wilson 2010, 2016; Wilson other known locality, from along streams at and Johnson 2010; Wilson and Townsend low elevations of Cerro Calentura south of 2006, 2010; Wilson et al. 2001), comparsions Trujillo, Colón. Four specimens of this spe- to new species (McCranie and Wilson 1997; cies were collected at that locality on one day Savage 2001), conservation status (Johnson during June 1989 at 460 m elevation. The au- et al. 2015; McCranie and Castañeda 2005; thor has not visited that site since that year. McCranie and Wilson 2004; Mitchell 2017; Two Quebrada de Oro specimens (USNM Stuart et al. 2008; Townsend et al. 2012; Wil- 497113–114) appear to represent hybrids son and McCranie 1998, 2003, 2004; Wilson between Craugastor chrysozetetes and Crau- and Townsend 2006, 2010), identification gastor fecundus. The following identifying keys (McCranie and Castañeda 2007; McCra- remarks were listed under Eleutherodactylus nie and Wilson 1997, 2002), and , chrysozetetes by McCranie and Wilson (2002) systematics, and phylogenetics (Frost et al. and taken from the author's field notes for 7 2006; Lynch 2000; Padial et al. 2014). August 1989. The latter specimen (USNM 497114; Figure 2) was identified as a hybrid in REMARKS. Six trips to the Quebreda de Oro the field because the specimen had yellow in region, the type locality of the species, were the groin and on the ventral surfaces (a trait 6 of Craugastor fecundus; purple in Craugastor character was based on more than one species chrysozetetes), red pustules on the dorsal sur- (based on his own data included in McCranie faces (a trait of Craugastor fecundus) on an et al. 1989). The jaw musculature discussed by otherwise typical dorsal pattern for Craugas- Lynch (1993; as Eleutherodactylus) for Crau- tor chrysozetetes, broadly expanded finger and gastor milesi actually applies to Craugastor toe discs (traits of Craugastor chrysozetetes), fecundus (based on KU 209105). Discussion and well-developed toe webbing (approach- of this jaw musculature character was repeat- ing the character state found in Craugastor ed by Campbell and Savage (2000), using KU chrysozetetes). The iris lacked red marks in 209105 as the source of those data. Specimens this specimen (a trait of Craugastor fecundus), of Craugastor fecundus (all as Eleutherodac- and it had well-developed lateral fringes on tylus) were variously identified as Craugas- the unwebbed portions of the toes as occurs tor aurilegulus (Campbell and Savage 2000), in Craugastor chrysozetetes. The second spec- Craugastor milsei (Lynch 1993; Lynch and imen (USNM 497113; Figure 3) resembled Fugler 1965; Meyer 1969; Meyer and Wilson USNM 497114 by "having nearly uniformly 1971; Savage 1975; Schmidt 1933), Craugas- pale cream ventral and groin surfaces in pre- tor rugulosus (Meyer 1969; Meyer and Wilson servative, broadly expanded finger and toe 1971), or Craugastor stadelmani (Campbell discs, well-developed lateral fringes on the 1994). toes, and a comparable amount of webbing" The suggestion thatCraugastor Cope (McCranie and Wilson 2002:376). Addition- might be available as a subgenus or genus ally, both of these presumed hybrids most name for the complex of containing the closely resemble Craugastor chrysozetetes in subsequently described Craugastor fecundus size. (Lynch’s Eleutherodactylus rugulosus group) was made by Lynch (1986, 1993). Craugastor ADDITIONAL VERNACULAR NAMES. fecundus was listed as belonging to the sub- “Ranita de Arroyo de la Cordillera de Nom- genus Craugastor by Lynch (2000). The use of bre de Dios” was used by McCranie and Craugastor as a genus name was advocated by Castañeda (2007) as a Spanish equivilant of Savage (2002), but Craugastor was used as a the English name suggested herein. "Elevated subgenus of Eleutherodactylus by Crawford Patterlove" was suggested by Mitchell (2017). and Smith (2005). The subgenus nameCamp - bellius was erected for the group of frogs con- ETYMOLOGY. The name fecundus is Lat- taining Craugastor fecundus by Hedges et al. in, meaning fertile or fruitful. The name was (2008); however, several errors occur in their used in reference to the Cordillera Nombre literature-based morphological description de Dios, the mountain range where the type of that subgenus. The speculation that the series of Craugastor fecundus was discovered. subgenus Campbellius “will ultimately be That mountain range has proved to be fertile considered” a genus was made by Padial et al. or fruitful ground for discovery of other un- (2014). However, that bold speculation was described anuran species (see McCranie in based solely on genetic data available from press). only two of the 13 species included in that subgenus by Padial et al. (2014). COMMENTS. The disagreement in tym- The holotype of Craugastor fecundus panum characters of Craugastor fecundus (LACM 137311) was not listed by Wright et as discussed by Savage (2001) in relation to al. (2008) in the type specimen list for the McCranie and Wilson (1997), was a result of Natural History Museum of Los Angeles Savage not realizing that his discussion of that County (LACM). 7

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