RELATIVELY few years ago, Campbell and Smith

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1 Copeia 2009, No. 2, A New Species of Stream-breeding Treefrog of the Genus Charadrahyla (Hylidae) from the Sierra Madre del Sur of Guerrero, Mexico Jonathan A. Campbell 1, J. Cristian Blancas-Hernández 2, and Eric N. Smith 1 A new species of treefrog is described from the central region of the Western Sierra Madre del Sur of Guerrero, Mexico. This frog is a member of the genus Charadrahyla, which contains five other species that are restricted to mesic highlands in Mexico. It possesses extensive webbing and in particular a hypertrophied membrane between toes I and II. This characteristic distinguishes the species from all other Middle American hylid frogs, except C. trux that also occurs in Sierra Madre del Sur of Guerrero. From C. trux the new form differs in being smaller, having relatively longer legs, aspects of coloration, and in having more prominent cloacal ornamentation. Se describe una nueva especie de rana arborícola del área central de la Sierra Madre del Sur del Oeste de Guerrero, México. Esta rana pertenece al género Charadrahyla, que contiene otras cinco especies. La nueva especie posee membranas extensas y en particular, membranas superdesarrolladas entre los dedos I y II. Esta característica distingue la nueva especie de cualquier otra rana hylida de Centro y Norte America excepto. C. trux, habitante de la región este de la Sierra Madre del Sur del Oeste de Guerrero. De C. trux la nueva especie difiere en ser más pequeña, tener piernas más largas, aspectos de coloración, y en tener ornamentos cloacales más prominentes. RELATIVELY few years ago, Campbell and Smith (1992) emphasized that Neotropical hylids were paraphyletic and that it was only a matter of time before individual monophyletic lineages would yield to generic recognition. Faivovich et al. (2005) presented a phylogenetic analysis of the Hylidae and recognized 16 genera in the New World tribe Hylini, proposing the new name Charadrahyla for the Hyla taeniopus group of Duellman (1965, 1970, 2001). The first member of this group to be described was Charadrahyla taeniopus (Günther, 1901) from the highlands of west-central Veracruz and subsequently discovered to be relatively widespread in the central portion of the Sierra Madre Oriental in the states of Hidalgo, Puebla, and Veracruz (Duellman, 2001). Charadrahyla chaneque (Duellman, 1961) was described from the cloudforests of western Chiapas and remains the only member of the genus known east of the Isthmus of Tehuantepec. Duellman (1965) prepared the first review of this group, at that time containing only two recognized species, C. taeniopus and C. chaneque. Lynch and Smith (1966) described Hyla duellmani from the highlands to the north of Zanatepec, Oaxaca, which was shown to be a synonym of C. chaneque (Duellman, 1970). Material collected subsequent to the description of C. chaneque in the Sierra de Juárez was allocated to C. chaneque by Duellman (1970), but was demonstrated to be a distinct species by Mendelson and Campbell (1999), for which they proposed the name C. nephila. Two additional species were added to the group from the Sierra Madre del Sur: C. altipotens (Duellman, 1968) from Oaxaca and C. trux (Adler and Dennis, 1972) from Guerrero. While conducting biodiversity surveys on the amphibians of Mexico, a field party from The University of Texas at Arlington, the Universidad Nacional Autónoma de México (UNAM), and the Universidad Autonoma de Guerrero (UAG) collected a series of a large and handsome species of treefrog in the mountains of western Guerrero. This species unquestionably appears to be a member of the genus Charadrahyla,which comprises relatively large, stream-breeding frogs inhabiting cloudforest or humid pine oak forest at elevations of m in central and southern Mexico. These frogs are characterized by usually having a brownish dorsum with large blotches (except C. altipotens), clear palpebral membranes, nuptial excrescences consisting of large patches of dark spinules (except C. altipotens), well-ossified quadratojugals in contact with the maxillaries (except C. trux), a tarsal fold, tadpoles adapted for mountain streams with two or three upper rows of teeth, three or four lower rows of teeth, and lips completely bordered by papillae (unknown for C. chaneque and the species described herein), and no axillary membrane. MATERIALS AND METHODS Standard terminology, abbreviations, and measurements as described by Duellman (2001) have been used. For ease of comparison, we have formatted the description following those that we have published previously for hylids (Mendelson and Campbell, 1999; Campbell et al., 2000). Measurements were made using digital calipers held under a dissecting microscope and rounded to 0.1 mm. Sex of adult individuals was determined by the presence of secondary sexual characters such as nuptial excrescences. The procedure for describing webbing formulae of hands and feet is that of Savage and Heyer (1967), as modified by Myers and Duellman (1982) and Savage and Heyer (1997). Color pattern of the new species is taken from a series of color images taken for each of three specimens and color notes prepared in the field. Comparisons were made with all other congeners (Table 1). Charadrahyla tecuani, new species Figures 1 3 Holotype. MZFC (original field no. ENS 11909), adult male (Fig. 1A), Sierra Madre del Sur: Carretera Bajos de 1 Department of Biology, The University of Texas at Arlington, Arlington, Texas 76019; (JAC) Campbell@uta.edu; and (ENS) e.smith@uta.edu. Send reprint requests to JAC. 2 Instituto de Investigación Científica, Área de Ciencias Naturales, Universidad Autónoma de Guerrero, Chilpancingo, Guerrero, México; E- mail: streptojcbh@yahoo.com.mx. Submitted: 15 August Accepted: 10 November Associate Editor: D. Kizirian. F 2009 by the American Society of Ichthyologists and Herpetologists DOI: /CH

2 288 Copeia 2009, No. 2 Table 1. Selected Distinguishing Characteristics of Species of Charadrahyla. Data taken from Adler and Dennis (1972), Duellman (1970, 2001), Mendelson and Campbell (1999), and specimens examined. Character C. tecuani C. taeniopus C. nephila C. chaneque C. altipotens C. trux Maximum adult SVL (mm) 57.8 (male) 65.9 (male) 70.9 (male) 60.7 (male) 80.6 (male) 79.1 (male) (female) 70.0 (female) 80.7 (female) 79.3 (female) 78.8 (female) Male snout shape dorsal Acuminate Acuminate Bluntly rounded Bluntly rounded Acuminate Acuminate profile Vocal slits Present a Present Present Absent Absent b Present Nuptial excrescences Present Present Present Present Absent Present Testes Enlarged Enlarged Normal Normal Enlarged Enlarged Webbing foot of males Hypertrophied Normal Normal Normal Normal Hypertrophied Quadratojugal articulating Yes Yes Yes Yes Yes No with maxillary Belly coloration in adults Yellow Usually brownish black with yellow flecks c Purplish tan Dull creamy brown Yellow Dull creamy brown a Three of the five adult males comprising the type-series have vocal slits on both sides; one individual has a slit on one side only. One male (UTA A-58708), apparently an adult based on its well-developed testes, lacks vocal slits. b Reported to be absent in Duellman (1970) and present in Duellman (2001). c Variable. Individuals in some populations with pale venters (Duellman, 1970). Balzamar La Sierrita, Sierra Madre del Sur, Guerrero, Mexico, 1552 m, uN, uW, 6 June 2007, Eric N. Smith, G. Andrea Acevedo, J. Cristian Blancas-Hernández, Ur. O. García-Vázquez, Robert C. Jadin, and Coleman M. Sheehy, III. Paratypes. All adult males, bearing identical locality data as the holotype. MZFC 22091, UTA A , 6 June 2007; UTA A (Fig. 1B), 5 June Diagnosis. A species of Charadrahyla moderate in size (males to 57.8 mm SVL), with a protruding snout and vocal slits in males, a distinct tarsal fold extending the length of the tarsus, hypertrophied webbing on foot, a boldly blotched pattern on dorsum and flanks, distinctly barred limbs, and clear palpebral membrane. Except for C. trux, it may be distinguished from all other members of the genus, as well as all other Middle American hylids, by the presence of hypertrophied webbing between the first and second toes (Table 1). Charadrahyla tecuani is easily distinguished from C. trux, the only other member of the genus which occurs in the Guerreran highlands, by its smaller size (adult males mm vs mm SVL), proportionately longer legs (tibia/svl vs ), pale venter with a few dark marking on throat (vs. dark venter with small pale dots on throat), and larger, more prominent vent tubercles and longer vent sheath. Description of holotype. Body moderately slender, SVL 55.1 mm, tibia length 29.9 mm, foot length 25.1 mm, head length 17.4 mm, head width 16.4 mm, diameter of tympanum 2.5 mm, diameter of eye 5.5 mm, interorbital distance 5.0 mm, eye tympanum distance 2.6 mm. Head about as wide as long; snout pointed in dorsal profile, without rostral keel; canthus rostralis distinct, angular; loreal region concave; lips not flared; nostrils ovoid, distinctly protuberant, directed posterolaterally; internarial region concave. Top of head flat; interorbital region 30.5% of width of head; diameter of eye 33.5% width of head. Supratympanic fold distinct, thick, extending posteroventrally from posterior margin of orbit, becoming indistinct at level of insertion of forearm; tympanum distinct, round; tympanic annulus mostly distinct, but obscured by supratympanic fold dorsally; width of tympanum 45.5% diameter of eye; width of tympanum 96.2% eye tympanum distance. Axillary membrane absent; thoracic fold absent; dermal fold on wrist present. Fingers long, slender, with broad lateral fringes, bearing large, ovoid terminal discs; relative lengths: I, II, IV, III; discs on Fingers II, III, IV approximately equal in size, as wide as tympanum; disc on Finger I smaller, width 60.0% width of tympanum. Subarticular tubercles small, diameter about one-third width of terminal disc on same finger, rounded, none bifid; supernumerary tubercles smaller than subarticular tubercles, rounded, indistinct. Nuptial excrescences consisting of tiny spinules, forming a large patch on the prepollex of Finger I and extending distally to disc; a narrow patch of spinules also present on the medial side of Finger II; ulnar tubercles large, irregularly coalesced to form low fleshy ulnar ridge. Hand webbing formula: I 2 2J II 1J 2 III 2 1 IV (Fig. 2). Heels of adpressed hind limbs overlap, tibiotarsal articulation extending just past snout; tarsal fold present, extending from heel to disc; tibia length 54.3% SVL; foot length 45.6% SVL. Inner metatarsal tubercle distinct, large, rounded, between times larger than subarticular tubercles; outer metatarsal tubercle absent; subarticular tubercles distinct, large, elevated, rounded, diameter about one-half width of terminal disc on same toe; supernumerary tubercles small, rounded, arranged in rows along axis of proximal portions of phalanges. Toes long, slender, bearing ovoid terminal discs slightly smaller than discs on fingers. Foot webbing formula: I 3/4 1/2 II 1/2 2 III 1 2 IV 2 3/4 V. Cloacal opening directed posteroventrally; a large, yellowish, conical tubercle on either side of vent (Fig. 3A), partially covered by distinctive vent sheath. Skin on dorsal surfaces granular in life, smooth after preservation; skin on ventral surfaces distinctly granular; skin on flanks between forelimbs and hind limbs distinctly thick and glandular. Tongue large, cordiform (notched posteriorly), barely free posteriorly and even less so anteriorly. Vomerine odontoids 5 6 on each side, situated on transverse dentigerous

3 Campbell et al. New Charadrahyla 289 Fig. 1. (A) Holotype of Charadrahyla tecuani, MZFC 22090, 55.1 mm SVL; reproduced from UTADC (B) Paratype, UTA A-58708, 52.5 mm SVL; reproduced from UTADC 1840.

4 290 Copeia 2009, No. 2 Fig. 2. Ventral aspect of hand and foot of Charadrahyla tecuani, male holotype (MZFC 22090). Black arrow indicates hypertrophied webbing between first and second toes. processes at midlevel between choanae, separated medially by length of odontoid process or slightly less; choanae subtriangular, widely separated. In preservative, dorsum of body and head and lateral surfaces of head are dull gray-brown with large distinct irregular dark brown blotches; dorsal surfaces of limbs pale brown with distinct dark brown transverse bars; flanks pale cream with distinctive dark brown markings, some forming irregular ovoids; dorsal surfaces of limbs tan to pale graybrown with distinct dark brown crossbars, three on forearm, four on thigh, and three on tibia; venter of throat cream with few scattered, irregular dark markings; chest immaculate, cream; belly yellow; undersurface of thighs yellow with pink tinge; palpebral membrane clear, dark edged above. Variation. The largest male in the type-series is 57.8 mm in SVL (Table 2), suggesting that this is one of the smallest species of Charadrahyla (Table 1). In three specimens the tympanic annulus is relatively indistinct (not prominently raised), which may be an artifact of preservation. The testes of two individuals (MZFC ) were yellow and relatively large (13.5 and 13.4 mm; % of SVL, respectively), and proportionally slightly larger than the testes size reported for C. trux (Adler and Dennis, 1972; % of SVL). The skin on the dorsum is reported to be smooth in all species of Charadrahyla, except C. chaneque, which has tuberculate skin (Mendelson and Campbell, 1999; Duellman, 2001). The texture of the skin of these streaminhabiting frogs, however, appears to be easily relaxed during preservation, which results in loss of tubercularity. Photographs in life of C. tecuani clearly show a tubercular skin, but the dorsum is smooth in the well-preserved typeseries. We have observed images of live C. nephila with highly tubercular skin, but preserved specimens range from smooth skinned to slightly tubercular. Tubercular skin may also vary among individuals, between sexes, and ontogenetically. The type-series consists entirely of adult males, and all have nuptial excrescences on the prepollex and first finger, and all individuals except UTA A have spinules along the medial edge of the second finger. In life, the dorsal ground color of adult males is medium brown to reddish brown. Large irregular blotches on the dorsum are dark brown. These blotches are essentially uniformly dark brown with a trace of pale centers in some specimens; in other specimens the blotches are heavily superimposed with numerous small leaf green blotches that extend into the ground color but are not as numerous outside of the blotches. Dark brown bars on the forearms, thighs, and shanks are more-or-less uniformly colored in individuals with uniformly brown dorsal blotches, but are suffused with considerable green pigment in other specimens. Flanks are bright yellow with large dark brown spots tending to have darker borders; lower portion of flanks grade into a pale salmon color. The dorsa of the digits are marked with several relatively inconspicuous dark brown bars, and the top of the discs grade from yellowish (inner digits) to tan (outer digits). Posterior of thighs are mauve to dark brown with small yellow spots. Webbing is chartreuse to yellowish tan. The loreal region is dark brown, often bordered below with irregular green markings that extend to below the eye. The supratympanic fold is marked with dark brown pigment extending from behind the eye over and posterior to the tympanum and may extend irregularly to a point over the insertion of the forearm. The undersurface of the forelimbs, irregular ulnar tubercles, outer tarsal tubercles, and calcar/ heel tubercles are cream. The large conical cloacal tubercles are yellow, and the posterior of the body above the vent, including the cloacal sheath, is dark brown with small white and/or yellow spots. The palmar and plantar surfaces are reddish proximally to mauve more distally on digits and yellowish on webbing. The throat is cream or pale yellow with a few irregular dark markings, some of these vermiform; the chest is pale yellow grading to a darker yellow in the belly (Fig. 3B). The underside of the thighs are reddish. The iris color is copper to bronze with fine black reticulations. Habitat and natural history. Charadrahyla tecuani is known from a single cloudforest locality in the western portion of the Sierra Madre del Sur of Guerrero (Fig. 4). The locality is a mountain stream that crosses the road between Bajos de Balzamar and La Sierrita, at 1552 m in the headwaters of the Río San Luís (Fig. 5). The vegetation is broad-leaf forest and treeferns mixed with pine oak, which has been heavily logged and trampled in some areas by grazing livestock. The stream has multiple cascades over huge boulders and between relatively quiet stretches with muddy bottoms. The first specimen was found during the day at 1410 hr on 5 June The frog was inactive on the underside of a fern frond between the roots of a tree and at the edge of the stream. The tree was between the beginning of a slow moving section of the stream and the entrance of a cave formed by two huge boulders over the stream. Realization of the uniqueness of this first specimen prompted us to return at night the following day, between 2000 and 2200 hr. On this occasion four more specimens were secured. One of the frogs was calling from a branch over the stream and another from a boulder in the middle of the stream. The call was an infrequent, long and soft boop. One specimen was

5 Campbell et al. New Charadrahyla 291 Fig. 3. (A) Venter of Charadrahyla tecuani (paratype, UTA A-58709, 57.8 mm SVL), reproduced from UTADC (B) Posterior of body (holotype, MZFC 22090), reproduced from UTADC 1866, showing enlarged pale vent tubercles.

6 292 Copeia 2009, No. 2 Table 2. Morphometric Variation in Type-Series of Charadrahyla tecuani, Consisting of Five Adult Males. Mean 6 SD above range (in parentheses); all measurements in mm. SVL ( ) Head length ( ) Head width ( ) Eye diameter ( ) Tympanum diameter ( ) Tibia length ( ) Foot length ( ) floating downstream with the legs spread, and another was sitting with eyes open on a branch. Two species of Charadrahyla occur in the Sierra Madre Oriental (C. taeniopus, C. nephila) from northern Hidalgo to north-central Oaxaca; one species occurs east of the Isthmus of Tehuantepec (C. chaneque) in the mountains of extreme southeastern Oaxaca and northeastern Chiapas; and three species inhabit the Sierra Madre del Sur, one in Oaxaca (C. altipotens) and two in Guerrero (C. trux, C. tecuani). The distributions of species of Charadrahyla are disjunct no case of sympatry among congeners is known. Charadrahyla taeniopus and C. nephila approach each other to within about 100 km. The other species of Charadrahyla known from the Sierra Madre del Sur of Guerrero, C. trux, has been found some 70 km to the east in the highlands near Cerro Teotepec. The presence of another member of the genus from these mountains was not anticipated. Several tadpoles belonging to other species of frogs were collected in the stream at the type-locality, many of them with obliterated and unkeratinized mouth parts, probably due to chytrid fungus infection. These larvae represent species of Plectrohyla, Ptychohyla, and Rana. Etymology. The specific epithet is taken from the Náhuatl term tecuáni, a noun referring to the jaguar (Panthera onca), a species formerly widespread in much of Mexico including Guerrero and possessing a bold pattern of dark spots. DISCUSSION Even though the Guerreran highlands had been largely ignored by biologists at the time, Taylor (1942) noted the isolated nature of this region and its high degree of endemicity. Except for the famous collecting site of Omilteme (sometimes Omiltemi ) near Chilpancingo, the uplands comprising the Sierra Madre del Sur in Guerrero received scant attention from herpetologists prior to about Since that time, expeditions led primarily by Kraig Adler and associates and by personnel from the University of Texas at Arlington have made numerous forays into more remote sections of these mountains. These trips have Fig. 4. Distribution of members of the genus Charadrahyla in the Sierra Madre del Sur of Guerrero and western Oaxaca; records exist for C. altipotens further to the east than shown.

7 Campbell et al. New Charadrahyla 293 Fig. 5. Type locality of Charadrahyla tecuani in the western portion of the Sierra Madre del Sur of Guerrero, showing stream where all known specimens were collected, as seen from the road between Bajos de Balzamar and La Sierrita at 1552 m. The black animal at the center of the image is Canis lupus familiaris. yielded a number of discoveries (Adler and Dennis, 1972; Smith and Savitsky, 1974; Campbell and Armstrong, 1979; Myers and Campbell, 1981; Adler, 1996). The Sierra Madre del Sur of Western Guerrero is relatively isolated from other highlands in Mexico: to the south lies the Pacific Coastal Plain, to the north and west is situated the deeply entrenched Rı o Balsas Basin with xerophytic vegetation, and to the east lies the dry pine oak and dry forest entrenchment of the Rı o Papagayo, including the pass in which lies Chilpancingo. The main axis of the Sierra Madre del Sur of Guerrero extends eastward into Oaxaca, but the crest of this range descends to elevations below 1500 m, in several places with dry valleys extending to the crest, effectively isolating cloudforest habitats. It has been previously suggested that the major disjunction of habitat and associated faunas occurred to the east of Chilpancingo (see Myers and Campbell, 1981, for a discussion). A picture is now emerging that suggests that significant breaks in faunal distributions may also occur in the major uplifts to the west of Chilpancingo, resulting in isolation and restriction of species to a single highland block. Charadrahyla tecuani is known from a single locality along a small stream that forms part of the headwaters of the Rı o San Luis. This watershed drains the Pacific slope and several peaks and crests in the region exceed 2500 m. There is an exceedingly tight interdigitation of habitats in the Sierra Madre del Sur, especially in Guerrero. Prevailing moisture-laden winds come from the south off the Pacific Ocean for most of the year, resulting in mesic broadleaf forests at the upper elevations. These forests often extend to the top of the crest or may be replaced at the higher elevations by fir pine oak forests. The cloudforests of Guerrero are not the dripping, soggy forests that are found in a few places in southern Mexico and Central America, usually on the Atlantic versant. It has been our experience that the Guerrero forest tend to be relatively dry for at least several months at a time during a period between January and May when rainfall is intermittent. MATERIAL EXAMINED Charadrahyla altipotens: Mexico, Oaxaca: Jalatengo, 0.2 km N, KU ; Jalatengo, 3.9 km N, KU ; Jalatengo, 4 km N, KU ; Jalatengo, 5.1 km S, KU ; Rı o Jalatengo, 0.8 km S Jalatengo, KU ;

8 294 Copeia 2009, No. 2 San Gabriel Mixtepec, 35.9 km N, KU , ; San Gabriel Mixtepec, 37 km N, KU , Charadrahyla chaneque: Mexico, Chiapas: 6.2 km S of Rayón Mescalapa, ca m, UMMZ ; Puerto Viento, 3.6 mi W along Mex Hwy 195, stream on steep slope, Selva Negra, CAS ; Puerto Viento, 4.0 mi W along Mex Hwy 195, stream on steep slope, Selva Negra, CAS ; Rayón Mescalapa, 5.6 km S, KU 58444; Rayón Mescalapa, 6.2 km S, KU ; Oaxaca: Sierra Madre, Zanatepec, UIMNH 56821; Zanatepec, above Sierra Madre, KU Charadrahyla nephila: Mexico, Oaxaca: Cerro Pelon, 10.0 mi N, UTA A-4769; Cerro Pelon, 9.6 mi (by road) N of crest, N slope Sierra de Juarez, UTA A-4770; Sierra Mixes, 3.6 mi (by road) N Totontepec, ca ft, UTA A-5887; Sierra Mixes, 5.6 km W Totontepec, 2121 m, UTA A ; Sierra Mixes, 5.8 km (by road) W Totontepec, headwaters of Río de la Lana, 2103 m, UTA A-5796; Teotitlan, 40.1 km E on road to Huautla, UTA A ; Totontepec, 3.1 mi W, Sierra Mixe, UTA A , ; Totontepec, 3.6 mi (by road) W, headwaters of Río de la Lana, Sierra Mixe, UTA A , ; Valle Nacional, 10.5 km S, KU ; Valle Nacional, 12.7 km S, KU ; Valle Nacional, 21.2 mi S, UTA A ; Valle Nacional, 9.1 km S, KU ; Vista Hermosa, 11.1 km S, KU ; Vista Hermosa, 11.6 km S, KU ; Vista Hermosa, 4.2 km S, KU , , , ; Vista Hermosa, 7.8 km S, KU ; Vista Hermosa, 9.4 km S, KU ; Vista Hermosa, 2.5 mi S, Sierra de Juarez, UTA A-6836; Vista Hermosa, 8 km S, Sierra Juarez, CAS , ; Vista Hermosa, 11 km S, KU 86968; Vista Hermosa, 11.9 km S, KU ; Vista Hermosa, 12.3 km S, KU ; Vista Hermosa, 13 km S, KU ; Vista Hermosa, 15.8 km S, KU ; Vista Hermosa, 16 km S, KU 86971; Vista Hermosa, 16.6 km S, KU ; Vista Hermosa, 4.1 km S, KU ; Vista Hermosa, 6 km S, KU 58445; Vista Hermosa, 6.5 km S, Arroyo Buena Vista, KU ; Vista Hermosa, 7.5 km S, KU 86956; Vista Hermosa, 9.3 km S, KU ; Yetla, 8 km S, KU 86967; Yetla, 9 km S, KU Charadrahyla taeniopus: Mexico, Hidalgo: Tianguistengo, 2.5 km SW, KU ; Tianguistengo, 4 km SW, KU 53830; Tlanchinol, 4.7 km SW, UTA A ; Tlanchinol, Carretera Tlanchinol-Sierra Colorada, 1375 m, uN, uW, UTA A-56957; Tlanchinol: La Cabaa, 1538 m, uN, uW, UTA A ; Xochicoatlan, 3 km W, KU , ; Puebla: Cuetzalan, Cueva de Guayateno, KU ; Cuetzalan, Santa Lucia, Cueva de Guayateno, KU ; Jonotla, Cueva de Aguayaco, KU ; Río Octapa, 3.7 km NNE Tezuitlan, KU , 57827, 65062; Sierra Norte: Cuetzalan: Hotel Villas Cuetzalan, 1250 m, uN, uW, UTA A ; Sima Esteban, 8 km SW Cuetzalan, KU ; Tlatlauquitepec, 1.5 km SW, KU 65057; Veracruz: Huatusco, KU ; Huatusco, 3 km SW, KU Charadrahyla trux: Mexico, Guerrero: 120 km by road E Chilpancingo, stream jct W of Mezones, 17u32.879N, 098u53.409W, UTA A-54815; 16.5 km SW of Puerto del Gallo (on road to Atoyac), 1700 m, UTA A ; Puerto del Gallo, 10.4 km SW by road, KU ; Puerto del Gallo, 11.4 km SW by road, KU ACKNOWLEDGMENTS We are indebted to various museum staff for the loan of critical material: C. Franklin (UT Arlington), C. Phillips (Illinois Natural History Survey/UIMNH), and L. Trueb (University of Kansas). We are particularly thankful to E. Beltrán Sánchez for providing logistical and field support during our stay in the State of Guerrero. U. García Vázquez, C. Sheehy, III, R. Jadin, and G. Acevedo (UTA) participated in the fieldwork that led to the discovery of the new species. O. Flores-Villela provided logistical support and help with permits. This paper is based on work supported by the National Science Foundation (grant no. DEB [ENS and JAC], DEB [JAC]) and Instituto Bioclon (ENS). Collecting permits were issued by the Secretaria de Medio Ambiente y Recursos Naturales (SEMARNAT). LITERATURE CITED Adler, K The salamanders of Guerrero, Mexico, with descriptions of five new species of Pseudoeurycea (Caudata: Plethodontidae). Occasional Papers of The Natural History Museum, University of Kansas 177:1 28. Adler, K., and D. M. Dennis New tree frogs of the genus Hyla from the cloud forests of western Guerrero, Mexico. Occasional Papers of The Natural History Museum, University of Kansas 7:1 19. Campbell, J. A., and B. L. Armstrong Geographic variation in the Mexican Pygmy Rattlesnake, Sisturus ravus, with the description of a new subspecies. Herpetologica 35: Campbell, J. A., and E. N. Smith A new frog of the genus Ptychohyla (Hylidae) from the Sierra de Santa Cruz, Guatemala, and description of a new genus of Middle American stream-breeding treefrogs. Herpetologica 48: Campbell, J. A., E. N. Smith, and M. Acevedo A new species of fringe-limbed treefrog (Hylidae) from the Sierra Los Cuchumatanes of Northwestern Guatemala. Herpetologica 56: Duellman, W. E Description of a new species of tree frog from México. Studies of American hylid frogs VI. Herpetologica 17:1 5. Duellman, W. E Frogs of the Hyla taeniopus group. Copeia 1965: Duellman, W. E Descriptions of new hylid frogs from Mexico and Central America. University of Kansas Publications, Museum of Natural History 17: Duellman, W. E Hylid frogs of Middle America. Museum of Natural History, University of Kansas, Monographs 1: Duellman, W. E The hylid Frogs of Middle America, 2 volumes. Society for the Study of Amphibians and Reptiles, Contributions in Herpetology 18: Faivovich, J., C. F. B. Haddad, P. C. A. Garcia, D. R. Frost, J. A. Campbell, and W. C. Wheeler Systematic review of the frog family Hylidae, with special reference to Hylinae: phylogenetic analysis and taxonomic revision. Bulletin of the American Museum of Natural History 294: Günther,A.C.L.G.1901 [ ]. Reptilia and Batrachia, p In: Biologia Centrali-Americana: Zoology. F. D. Godman and O. Salvin (eds.). Dulau and Co., London. Lynch, J. D., and H. M. Smith New or unusual amphibians and reptiles from Oaxaca, Mexico, II. Transactions of the Kansas Acadademy of Science 69: Mendelson, J. R., and J. A. Campbell The taxonomic status of populations referred to Hyla chaneque in southern Mexico, with the description of a new treefrog from Oaxaca. Journal of Herpetology 33:80 86.

9 Campbell et al. New Charadrahyla 295 Myers, C. W., and J. A. Campbell A new genus and species of colubrid snake from the Sierra Madre del Sur of Guerrero, Mexico. American Museum Novitates 2708:1 20. Myers, C. W., and W. E. Duellman A new species of Hyla from Cerro Colorado, and other tree frog records and geographical notes form western Panama. American Museum Novitates 2752:1 25. Savage, J. M., and R. W. Heyer Variation and distribution of the tree-frog genus Phyllomedusa in Costa Rica. Beiträge zur Neotropical Fauna 5: Savage, J. M., and R. W. Heyer Digital webbing formulae for anurans: a refinement. Herpetological Review 28:131. Smith, H. M., and A. H. Savitsky Another cryptic associate of the lizard Sceloporus formosus in Guerrero, Mexico. Journal of Herpetology 8: Taylor, E. H Island faunas on the Mexican Plateau. Proceedings of the Eighth American Scientific Congress 3: