Part 3

Remarks.--Shannon (1951:470) named Hyla bistincta labeculata on the basis of a single male from San Lucas Camotlán, Oaxaca; he diagnosed the subspecies as differing from Hyla bistincta bistincta by having "the gray reticulation of the sides entirely broken up into elongate black blotches; tarsal fold moderately elevated." The condition of the tarsal fold is characteristic of the species. The dispersion of dark pigment on the flanks is variable. The type of Hyla bistincta labeculata (USNM 123689) is extreme in the development of dark dashes on the flanks, but this condition is approached in several specimes from Oaxaca and Veracruz. For example, in some specimens from Cumbres de Acultzingo, Veracruz, the mottling on the flanks is bold; in others the flanks are reticulated. The specimen from San Vicente, Oaxaca, has black dashes on the flanks (Smith and Williams, 1963:23), whereas a specimen from Cerro San Felipe, Oaxaca, has no pattern on the flanks. In general, specimens from western México have reticulate mottling on the flanks as compared with the marbling on the flanks in specimens from eastern México. On the basis of available data, the recognition of subspecies in Hyla bistincta is unwarranted.

The tadpoles of this species described by Duellman (1961:47) are like those of Hyla robertsorum in having 2/3 tooth-rows, peglike serrations on the beaks, and long, rounded tails. At Uruapan tadpoles were found in a rocky stream on April 24, 1956, and metamorphosing young were found there on August 2, 1956. A completely metamorphosed juvenile has a snout-vent length of 24.8 mm.

Hyla bistincta is found only along streams, where individuals can be seen clinging to vines and other vegetation closely over-hanging fast-moving parts of the stream.

Distribution.--Hyla bistincta occurs at elevations from 1400 to 2600 meters in the mountains of the Sierra Madre Occidental in western Jalisco southward through the Cordillera Volcánica in Michoacán, México, and Morelos, the Sierra de Coalcomán in Michoacán, and the Sierra Madre del Sur in Guerrero and Oaxaca, and thence northward in the Sierra Madre Oriental to central Veracruz (Fig. 2).

Specimens examined.--GUERRERO: Omiltemi, UIMNH 38023-5. JALISCO: 25 km. SE Autlán, UMMZ 102076. MEXICO: 19 km. W Villa Victoria, UIMNH 28162, USNM 114513. MORELOS: Cuernavaca, USNM 121523; 3 km. N Cuernavaca, UIMNH 28168-70. MICHOACAN: Cerro San Andrés, UMMZ 102075; Dos Aguas, UMMZ 119193; 12.5 km. ENE Dos Aguas, UMMZ 119194; Los Conejos, UMMZ 94238-40; Uruapan, KU 68077-8, 69093 (skeleton), UIMNH 20457, 28167, UMMZ 85452-3, 112838, 112839 (16), 115231 (tadpoles), 115232, 115233 (12), 121515, S-1699 (skeleton), S-1826 (skeleton), USNM 114514-5, 114517-24. OAXACA: Cerro San Felipe, UIMNH 28163; Pluma Hidalgo, AMNH 13447; San Lucas Camotlán, USNM 123689; San Vincente, UIMNH 51346 (Smith and Williams, 1963:23). VERACRUZ: no specific locality, USNM 32261; Cumbres de Acultzingo, CNHM 105482-3, UIMNH 28164-6, 49133-4, USNM 114525.

#Hyla charadricola# new species

Holotype.--University of Kansas Museum of Natural History No. 58414 from the Río Totolapa, 14.4 kilometers by road west of Huachinango, Puebla, México, 2280 meters; obtained by John Wellman on June 8, 1960.

Paratypes.--KU 58415-58423, same data as holotype, and UIMNH 50966, obtained at the type locality by William E. Duellman on February 11, 1961.

Diagnosis.--Maximum snout-vent length in males, 45 mm.; snout in dorsal profile truncate; tarsal fold short and weak; inner metatarsal tubercle moderately large, elliptical, and flat; outer metatarsal tubercle minute, round, and indistinct; webbing extending to base of terminal phalanx of fourth toe; nuptial spines on thumb absent; thoracic fold absent; anal opening at level of middle of thigh; dorsum olive-green with black reticulations; venter cream color with brown flecks on chin; flanks pale grayish green with brown spots; anal stripe absent; row of white flecks above and a row below anus; vocal slits absent.

Description of Holotype.--Adult male having a snout-vent lenth of 44.4 mm.; tibia length, 22.2 mm., 50 per cent of snout-vent length; foot length (measured from proximal edge of inner metatarsal tubercle to tip of longest toe), 20.4 mm.; greatest width of head, 14.0 mm., 31.5 per cent of snout-vent length; head length, 13.6 mm., 30.6 per cent of snout-vent length; diameter of eye, 5.0 mm.; diameter of tympanum, 1.6 mm., 32.0 per cent of diameter of eye. Snout in lateral profile bluntly rounded, in dorsal profile truncate (Fig. 3); canthus pronounced, rounded, not angular; loreal region slightly concave; lips thick, rounded, not flaring; nostrils slightly protuberant; internarial distance, 3.7 mm.; interorbital distance, 4.1 mm., somewhat broader than width of eyelid, 3.3 mm. A heavy dermal fold from posterior corner of eye above tympanum and then to insertion of forearm; tympanum round, its diameter three-fourths its distance from eye. Forearm rather slender; a short axillary membrane; no fold on wrist; no nuptial excrescence or spines on enlarged prepollex; subarticular tubercles moderately small, round; none bifid; few supernumerary tubercles on proximal segments of digits; large, flat palmar tubercle present; fingers long and slender; length of fingers from shortest to longest, 1-2-4-3; discs moderately large, that on third finger about equal to diameter of tympanum; rudimentary web between first and second fingers; web between third and fourth fingers extending to about one-fifth length of fourth finger (Fig. 1). Heels overlap by about one-third length of tibia when hind limbs adpressed; tibiotarsal articulation extends to middle of eye; tarsal fold weak, extending from moderately large, flat, elliptical inner metatarsal tubercle to middle of tarsus; outer metatarsal tubercle minute, round, indistinct; subarticular tubercles round; single row of small, round supernumerary tubercles on proximal segments of each digit; toes moderately short and slender; length of toes from shortest to longest, 1-2-3-5-4, third and fifth about equal in length; toes about three-fourths webbed; web extending to base of terminal phalanx of fourth toe; discs small, about two-thirds size of those on fingers. Anal opening directed posteroventrally at middle of thighs; two transverse dermal folds above anus; short, thin anal sheath present; many small tubercles lateral and ventral to anal opening. Skin of dorsum, chin, and ventral surfaces of limbs smooth; belly moderately granular; posterior surfaces of thighs heavily granular; no thoracic fold. Tongue nearly round, shallowly notched behind, free posteriorly for about one-fourth its length; vomerine teeth, 3-4, long, situated on posteroventral edges of narrow transverse vomerine ridges between moderately large, round inner nares; no vocal slits.

Color (in alcohol) purplish brown on dorsal surfaces of head, body, and limbs; fine darker reticulations on dorsum; flanks pale tan with dark brown spots; posterior surfaces of thighs tan; chin creamy white with brown spots; belly dusty white; undersides of thighs and shanks yellow; webbing grayish brown; undersides of first two fingers dusty white, of third and fourth fingers and of feet brown; anal stripe absent; small white flecks above and below anal opening.

Color (in life) dark green on dorsal surfaces of head, body, and limbs; darker green reticulations on back; flanks dusty white with dark olive-gray mottling; dark olive-gray stripe from nostril through eye and then to insertion of forelimb; upper lips pale green; inguinal region, anterior and posterior surfaces of thighs dark yellowish orange; ventral surfaces of tarsi and shanks, and webbing of feet dusty yellow; belly white; iris silvery gold.

Variation.--Adult males have snout-vent lengths from 35.3 to 44.4 mm., and adult females from 43.4 to 50.9 mm. No notable variation in structure is displayed in the type series. In some individuals the dark reticulation on the dorsum is faint. Juveniles in life had dorsal colorations varying from rich brown with darker reticulations to pale green or gray with dark green reticulations. Some adults when collected were pale green with faint or no dorsal reticulations; later these individuals darkened. In all specimens the anal stripe is absent, and the flanks are heavily spotted.

Comparisons.--Hyla charadricola differs from all other members of the Hyla bistincta group in the following combination of characters: truncate snout, green dorsum, and absence of a thoracic fold. From other hylids that occur in the same area, Hyla charadricola differs from Hyla miotympanum and H. arborescandens, both of which are green dorsally, by having a truncate snout and longer fingers with less webbing. Hyla eximia, though green, has brown dorsal spots, shorter fingers, and a round snout. Hyla robertsorum differs in having a round snout and brown dorsum, and Hyla taeniopus is much larger, has transverse bands on the limbs, and has extensive webbing between fingers.

Remarks.--At the type locality, a shallow rocky stream in pine forest, Hyla charadricola was found beneath rocks at the edge of fast moving sections of the stream and beneath rocks in shallow riffles in the stream. Most of the frogs were in water. At night they were found sitting on rocks in the stream. Hyla miotympanum, which is abundant at the type locality, lives in bushes and beneath rocks along the stream but usually is not found in the riffles inhabited by Hyla charadricola. At Lago de Tejocotal Hyla charadricola was found beneath rocks near the shore of the lake and by a stream in the pine forest. Individuals were found on low vegetation over-hanging a small stream in pine-oak forest four kilometers southwest of Tianguistengo.

Five recently metamorphosed young (KU 58424-9) found at the type locality on June 8, 1960, have snout-vent lengths of 22.4 to 24.0 (average 23.2) mm. The young are colored like the adults, except that in life the dorsum is a brighter green and the flanks are more yellow than tan and have less dark spotting than in adults.

Distribution.--Hyla charadricola inhabits streams in pine and pine-oak forests at elevations of 2000 to 2300 meters in northern Puebla and eastern Hidalgo (Fig. 2).

Specimens examined.--HIDALGO: Lago de Tejocotal, 11 km. E Acaxochitlán, KU 58438, UMMZ 104032, 118165; 4 km. SW Tianguistengo, KU 53811-2. PUEBLA: 11.7 km. W Huachinango, UMMZ 121567 (5); Río Totolapa, 14.4 km. W Huachinango, KU 53813-5, 55624, 58414-37, 59813 (skeleton), 59886 (skeleton), MCZ 34964-5, UIMNH 50966, UMMZ 118166 (5), S-2242 (skeleton).

#Hyla robertsorum# Taylor

Hyla robertsorum Taylor, Univ. Kansas Sci. Bull., 26:393-396, figs. 5-6, November 27, 1940 [Holotype.--CNHM 100124 (formerly EHT-HMS 16264) from El Chico Parque Nacional, Hidalgo, México; Mr. and Mrs. Radclyffe Roberts and Edward H. Taylor collectors]; Univ. Kansas Sci. Bull., 28:310, November 15, 1942. Taylor and Smith, Proc. U. S. Natl. Mus., 95:589, June 30, 1945. Smith and Taylor, Bull. U. S. Natl. Mus., 194:87, June 17, 1948; Univ. Kansas Sci. Bull., 33:333, March 20, 1950. Rabb and Mosimann, Occas. Papers Mus. Zool. Univ. Michigan, 563:1-9, March 29, 1955. Duellman, Univ. Kansas Publ. Mus. Nat. Hist., 15:48, December 20, 1961.

Diagnosis.--Maximum snout-vent length in males, 48 mm.; snout in dorsal profile rounded; tarsal fold short and weak; inner metatarsal tubercle moderate in size and elliptical; outer metatarsal tubercle small, round, and indistinct; webbing extending to base of penultimate phalanx of fourth toe; nuptial spines on thumb small; weak thoracic fold present; anal opening above level of middle of thighs; dorsum dark brown with dark reticulations; venter brown with cream-colored flecks; flanks brown with creamy white spots; anal stripe absent; small white spots in anal region; vocal slits absent.

Description.--The following description is based on KU 57651 from El Chico Parque Nacional, Hidalgo. Adult male having a snout-vent length of 45.1 mm.; tibia length, 22.1 mm., 49.0 per cent of snout-vent length; foot length (measured from proximal edge of inner metatarsal tubercle to tip of longest toe), 21.5 mm.; greatest width of head, 13.7 mm., 30.4 per cent of snout-vent length; head length, 12.6 mm., 27.9 per cent of snout-vent length; diameter of eye, 4.0 mm.; diameter of tympanum, 1.8 mm., 45 per cent of diameter of eye. Snout short, in lateral profile blunt, in dorsal profile round; canthus rounded; loreal region slightly concave; lips thick, round, and not flaring; nostrils slightly protuberant; internarial distance, 3.6 mm.; interorbital distance, 4.0 mm., slightly broader than width of eyelid, 3.5 mm. A moderately heavy dermal fold from posterior corner of eye above tympanum and curving downward towards insertion of forearm; tympanum nearly round, covered by dermal fold above, its diameter slightly less than its distance from eye. Forearm moderately robust; distinct fold on wrist; prepollex much enlarged with patch of small nuptial spines continuous on side of digit; similar line of nuptial spines on inner edge of second finger; subarticular tubercles round, moderate in size, none bifid; supernumerary tubercles small and present only proximally; fingers long and slender; length of fingers from shortest to longest, 1-2-4-3; discs moderately large, that on third finger about size of tympanum; no web between first and second fingers; rudimentary web between other fingers. Legs robust; heels over-lap by about one-fourth length of shank when hind limbs adpressed; tibiotarsal articulation extending to posterior corner of eye; tarsal fold weak, extending to about middle of tarsus; inner metatarsal tubercle moderately large, flat, and elliptical; outer metatarsal tubercle small, round, and indistinct; subarticular tubercles round; supernumerary tubercles small, in single row on proximal segments of each digit; toes moderately long and slender; length of toes from shortest to longest, 1-2-5-3-4, the fifth nearly as long as third; toes nearly fully webbed; web extending to base of penultimate phalanx of fourth toe and to discs on other toes; discs small, about two-thirds size of those on fingers. Anal opening above middle of thighs; anal sheath short, deeply creased medially; heavy transverse dermal fold above anus; no large anal tubercles. Skin of all dorsal surfaces, chin, and ventral surfaces of limbs, except proximal parts of thighs, smooth; belly and proximal parts of thighs areolate; thoracic fold present, weak. Tongue elliptical, slightly longer than wide, not notched behind, and free posteriorly for about one-fourth of its length; vomerine teeth 3-3, situated on small, widely separated, transverse ridges between rather small elliptical inner nares; no vocal slits.

Color (in alcohol) dark brown with irregular darker reticulations on dorsal surfaces of head, body, and limbs; flanks brown with small creamy white spots; posterior surfaces of thighs dark brown; chin creamy tan; belly grayish brown with cream-colored flecks; ventral surfaces of limbs pale brown; webbing on feet gray; small white spots in anal region.

Color (in life) chocolate brown with darker brown reticulations and irregular blotches above; flanks brown with yellow spots; belly gray to grayish brown with faint cream-colored spots; iris a deep bronze color.

Variation.--In males the total number of vomerine teeth varies from 4 to 7. In many specimens the vomerine ridges are larger and more closely approximated medially than in the specimen described above. Females attain snout-vent lengths of 51 mm., have as many as 9 vomerine teeth, and have a proportionately larger tympanum than males. Some of the largest specimens of both sexes have indistinct cream-colored pustules scattered on the ventral surface of the forearm. Some individuals have nearly uniform grayish brown ventral surfaces; in others the chin, as well as the abdomen, is brown with cream-colored spots. The dorsal surfaces of some specimens are nearly uniform dark brown with no reticulations. In others the dorsum is paler brown with distinct darker mottling; in some of these there is little mottling laterally, so that there is the effect of an irregular, pale brown, dorsolateral stripe.

Tadpoles.--The tadpoles of this species were described by Rabb and Mosimann (1955). Tadpoles obtained from streams at 3.3 kilometers north and at 8.5 kilometers southeast of Zacualtipán, Hidalgo, are like those described by Rabb and Mosimann in having 2/3 tooth rows, peglike serrations on the beaks, and long rounded tails. The largest tadpole (KU 60078) has small hind legs, a body length of 22 mm., and a total length of 61 mm.

Remarks.--Taylor (1940:393) found frogs of this species in plants along small spring-fed rivulets in an open meadow at El Chico Parque Nacional. Also, he noted that active frogs dove into the streams and took refuge in the mud on the bottom. Rabb and Mosimann (1955:1) found this species along banks of tiny streams in open meadows and noted that the frogs sought refuge in the water. At El Chico Parque Nacional on June 8, 1960, I found Hyla robertsorum under rocks along small rivulets by day; at night, when the temperature was 14° C., frogs were sitting on rocks and in junipers overhanging a small stream. At the same locality on June 23 and 24, 1962, frogs of this species were found on rainy nights, when the temperature varied from 10 to 12° C. At this time the frogs were sitting on the grassy banks of rivulets in the meadow. During the day Hyla robertsorum was found on the earthen banks of the rivulets in places where dense growths of grass overhung the streams. On December 23, 1959, one specimen of Hyla robertsorum was found beneath a rock in a small stream in pine forest at 3.3 kilometers north of Zacualtipán.

Rabb and Mosimann (1955:1) obtained tadpoles of Hyla robertsorum from quiet pools of a stream at El Chico Parque Nacional. I found tadpoles in pools in rocky streams in pine forest at 3.3 kilometers north and at 8.5 kilometers southeast of Zacualtipán. Four completely metamorphosed juveniles obtained on June 8, 1960, at El Chico Parque Nacional have snout-vent lengths of 30.6 to 32.0 mm. Gravid females were found at the same locality on June 8, 1960, and June 23, 1962.