1 AMPHIBIA: ANURA: Craugastor guerreroensis

Catalogue of American and Reptiles 919.

Palacios-Aguilar, R. 2019. Craugastor guerreroensis.

Craugastor guerreroensis (Lynch, 1967) Guerreran Robber

Eleutherodactylus guerreroensis Lynch 1967a: 216. Type locality: “3 km N Agua del Obispo, Guerrero, México, 980 m.” Ho- lotype: Kansas University (KU) 86873, an Figure. Adult female (MZFC 30709; SVL 50.4 adult male 39.2 mm snout-vent length, mm) Craugastor guerreroensis from the vicinity of collected by J. S. Waddick, 8 June 1964 Coaxtlahuacán, Guerrero, Mexico. Photographed (examined by the author). by the author on 3 July 2016. Eleutherodactylus (Craugastor) guerreroensis: Hedges 1989:317. By implication. are directed anterolaterally at the tip of the Craugastor guerreroensis: Duellman 1993:148. snout. Cranial crests absent. The tympanum By implication. is visible, though partially concealed with su- Craugastor guerreoensis: Glaw, Köhler, Hof- pratympanic folds on each side. richter, and Dubois 1998. Lapsus. Arms are slender and long (arm length/ Craugastor guerreroensis: Crawford and SVL in adult males =0.447–0.580, mean Smith 2005:551. By implication. [standard deviation] =0.506 [±0.049]; in adult Craugastor (Hylactophryne) guerreroensis: females =0.444–0.468, mean [standard devia- Hedges, Duellman, and Heinicke 2008:45. tion] =0.456 [±0.017]). Relative length of the Craugastor querreroensis: Liner and Ca- fingers is III-IV-II-I in decreasing length. The sas-Andreu 2008:10. Lapsus. fingers are not webbed and are without any distinctive keel or flare. The finger discs are CONTENT. No subspecies are recognized in broadly expanded and truncated, being wid- this taxon. est on Finger III. The subarticular tubercles are globular and either rounded or ovoid. DESCRIPTION. Craugastor guerreroensis Supernumerary tubercles are absent on the is a moderately sized (in seven adult fingers. The palmar tubercles are elevated and males, snout-vent length [SVL] range =31.7– ovoid, and the bigger one is bifid. Prepollex 39.2 mm, mean [standard deviation] =35.5 in adult males is not enlarged, and nuptial ex- mm [±2.85 mm]; in two adult females, SVL crescences are not present. = 50.4–55.5 mm, mean [standard deviation] The hind limbs are relatively long (shank =53 mm [±3.61 mm]) with a moderately long, length/SVL 0.561–0.608 in adult males, broad head (head length/SVL in adult males 0.513–0.625 in adult females; tarsus length/ =0.373–0.403, in adult females =0.355–0.358; SVL 0.344–0.490 in adult males, 0.367–0.470 head width/SVL in adult males =0.399–0.435, in adult females). Tarsal fold is weakly de- in adult females =0.391–0.423). The snout is veloped. The heels do not overlap when the truncated in lateral view and nearly rounded hind limbs are held together towards the clo- in dorsal view. The top of the head is slightly acal opening at right angles to the body. The concave. The lips are slightly flared. Nostrils only tubercles present on the underside of the 2

Map. Distribution of Craugastor guerreroensis. Black circle represents the type locality, open circles represent other records. The question mark represents a dubious record (CNAR 2386; seeComments ).

feet are subarticular tubercles and two paired posteriorly. The vomerine teeth are in two metatarsal tubercles. Subarticular tubercles distinct clumps, separated by a distance equal are round and globular. Metatarsal tubercles to about one fourth of the diameter of each are ovoid and globular, the outer being larger clump. The choanae are ovoid and positioned than the inner metatarsal tubercle. Relative perpendicularly to the vomerine clumps. size of the toes is: 4-5-3-2-1 in decreasing Color in life of an adult female (MZFC length. Toe tips are definite and broadly ex- 30709; Museo de Zoología “Alfonso L. Herre- panded. The toes have vestigial webbing be- ra” de la Facultad de Ciencias, UNAM, Mexico tween them. City) follows. Dorsal surfaces of the head, and The skin on the dorsal surfaces is smooth, body are mottled with dark-olive green and grading to slightly granular on lateral surfac- olive green. Ventral surfaces are mostly white es. A series of greatly enlarged tubercles on and purplish-brown at the level of the junc- the side of the body is always present from tion of the arms and head. The forearms, and the rear end of the tympanum to the groin legs show dark-olive green bars, more definite and occasionally covering the inner and dor- in the forearms. Tympanum dark brown with sal portion of the shanks. The skin on throat a mark that extends to the posterior margin and venter is smooth throughout. The ventral of the eye. Each iris is bronze with a brown disc is absent in both sexes. Males have vo- mark in the anterior and posterior ends. Each cal slits. The pupil is horizontally elliptical. iris is also reticulated with black. The palpebral membrane is translucent and unpatterned. The tongue is round and free DIAGNOSIS. The following diagnosis can 3 differentiate Craugastor guerreroensis from from Coaxtlahuacán, Guerrero, Mexico were other Mesoamerican representatives of the presented by Palacios-Aguilar (2017) and Craugastor alfredi group (as defined by Lynch Johnson et al. (2017). Black-and-white line 1966). Vocal slits are present in males of this drawings representing the dorsal view of the species, absent in all other members of the holotype, details of the hand, and a profile Craugastor alfredi group except Craugastor view of the head were published by Lynch decoratus, Craugastor polymniae, and Crau- (1967a). gastor spatulatus. Females are larger in Crau- gastor guerreroensis than most other species DISTRIBUTION. Craugastor guerreroen- (maximum size >50 mm SVL vs 36–46 mm), sis is known from the humid oak and cloud except Craugastor decoratus (maximum SVL forests of the central portion of the Mexican 60 mm). Additionally, Craugastor decoratus state of Guerrero, on the Pacific versant of the has wide dark bands on the thighs. Craugas- Sierra Madre del Sur at an elevation range tor guerreroensis can be differentiated from of 980–1700 m above sea level. Maps of the Craugastor decoratus by having a distinctive general range were published by Saldaña de la row of lateral tubercles from the rear end of Riva and Pérez Ramos (1987) and Stuart et al. the tympanum to the groin and lacking any (2008). A map showing the type locality and obvious markings on the flanks. Additionally, records of other members of the Craugastor Craugastor guerreroensis is also the only spe- alfredi group was presented by Campbell et al. cies of the Craugastor alfredi group known to (1989), and a simple dot map was published occur on the Pacific versant of Mexico, west by Palacios-Aguilar (2017). of the Isthmus of Tehuantepec. FOSSIL RECORD. None. PHYLOGENETIC RELATIONSHIPS. Crau- gastor guerreroensis is in the Craugastor alfre- PERTINENT LITERATURE. Relevant cita- di group as defined by Lynch (1966, 1967b). tions are listed by topic: checklists and fau- Based on comparisons of jaw musculature nal lists (Campbell 1999; Flores-Villela 1991, with other direct-developing , Craugas- 1993; Flores Villela and Gerez 1988, 1994; tor guerreroensis was placed into the Mid- Frost 1985; Glaw et al. 1998, 2000a, 2000b; dle American clade by Lynch (1986). Based Gorham 1974; Harding 1983; Johnson et al. on morphology, the species was placed into 2017; Liner 1994; Liner and Casas-Andreu the subgenus Hylactophryne by Hedges et 2008; Palacios-Aguilar 2016; Palacios-Agu- al. (2008) and Padial et al. (2014). Based on ilar and Flores-Villela 2018; Pérez-Ramos et morphological similarities, Craugastor guer- al. 2000; Saldaña de la Riva and Pérez Ramos reroensis was assigned to the Craugastor bo- 1987; Smith and Smith 1976, 1993), common courti species group by Hedges et al. (2008). names (Frank and Ramus 1995; Liner 1994; Liner and Casas-Andreu 2008; Mitchell 2017; PUBLISHED DESCRIPTIONS. The species Sokolov 1988; Wrobel 2004), comparisons was described and the holotype was illus- to new species (Campbell et al. 1989; Canse- trated by Lynch (1967a). The morphological co-Márquez and Smith 2004; McCranie and variation, natural history, and distribution of Smith 2006; Smith 2005), conservation sta- the species were reported by Palacios-Aguilar tus (Johnson et al. 2017; Lips et al. 2004; Stu- (2017). art et al. 2008; Wilson et al. 2013), , systematics, and phylogenetics (Crawford ILLUSTRATIONS. A color photograph of and Smith 2005; Frost 2018; Frost et al. 2006; a calling male was published by Stuart et al. Hedges 1989; Hedges et al. 2008; Lynch 1986; (2008); color photographs of an adult female Padial et al. 2014). 4

REMARKS. Craugastor guerreroensis is a Although the species is associated with cave poorly known species with scarce informa- systems in Guerrero, the locality in the Sier- tion on its biology, and is represented by less ra de Juárez in northern Oaxaca seems un- than a dozen specimens deposited in three likely and probably is a misidentification [not scientific collections: CNAR (Colección Na- shown on Map]. cional de Anfibios y Reptiles, Instituto de A single specimen (CNAR 2386) appar- Biología, Universidad Nacional Autóno- ently was collected near the town of Telo- ma de México (UNAM), Mexico City), KU loapan in the Balsas Depression of Guerrero, (University of Kansas Institute), a region characterized by having lower ele- and MZFC (Museo de Zoología “Alfonso L. vations and drier habitats (see Map; locality Herrera” de la Facultad de Ciencias, UNAM, labeled as "?"), however, it is possible that the Mexico City). One female collected in Cue- locality data associated with that specimen va La Pedrera de Cajeles, Guerrero (CNAR represent a cataloguing error because the as- 4285-1) on 18 December 1975 contained sev- sociated name on the field tag isRana aurora eral developed ova. draytonii (Palacios-Aguilar 2017). The specific microhabitat in which the type series was obtained is unknown, and the ACKNOWLEDGEMENTS. L. Welton and original description by Lynch (1967a) only R. Glor provided information on the type mentioned that the type series was collected series. O. Flores-Villela (MZFC) and V. H. from the Chilpancingo region of Guerrero in Reynoso (CNAR) provided access to the col- the Sierra Madre del Sur. Most of the speci- lections under their care. Access to crucial mens I collected and examined were obtained literature was facilitated by O. Flores-Ville- from caves in humid oak and cloud forests la and A. Ibarra-Reyes. L. M. Ochoa-Ochoa in the southcentral portion of the mountain provided comments on a preliminary version ranges south of the city of Chilpancingo. of the manuscript.

ADDITIONAL VERNACULAR NAMES. LITERATURE CITED “Rana Ladradora de Guerrero” was used by Campbell, J. A. 1999. Distribution patterns Liner and Casas-Andreu (2008) as a Spanish of amphibians in Middle America. Pp. equivalent of the English vernacular name 111-210 in Patterns of Distribution of proposed herein. The name “Rana Ladro- Amphibians. A Global Perspective (W. na de Guerrero” used by Liner (1994) is an E. Duellman, editor). The Johns Hopkins incorrect translation of the same vernacular University Press, Baltimore, Maryland. name. “Guerrero Patterlove” was provided by Campbell, J. A., W. W. Lamar, and D. M. Hil- Mitchell (2017). lis. 1989. A new species of diminutive Eleutherodactylus (Leptodactylidae) from ETYMOLOGY. The nameguerreroensis is Oaxaca, Mexico. Proceedings of the Bio- a toponym referring to the Mexican state of logical Society of Washington 102:491– Guerrero where the type series was collected 499. and to which the species is restricted. Canseco-Márquez, L. and E. N. Smith. 2004. A diminutive species of Eleutherodactylus COMMENTS. A record of Craugastor guer- (Anura: Leptodactylidae), of the alfredi reroensis from “Cueva del Nacimiento del Río group, from the Sierra Negra of Puebla, San Antonio, Oaxaca” was reported by Red- Mexico. Herpetologica 60:358–363. dell (1981: p. 249, Figure 87). That locality Crawford, A. J. and E. N. Smith. 2005. Ceno- would place Craugastor guerreroensis in sym- zoic biogeography and evolution in di- patry with Craugastor decoratus purpurus. rect-developing frogs of Central America 5

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