Part 7

Specimens examined.--MEXICO: Chiapas: 6 km. NE of Chiapa de Corzo, TCWC 16183; 16 km. E of Chiapa de Corzo, AMNH 62141; Linda Vista, 2 km. NW of Pueblo Nuevo Solistahuacan, KU 58049-51, 59939 (skeleton); Rio Hondo, 9.5 km. S of Pueblo Nuevo Solistahuacan, KU 58047-8, 60046-7, 60048-9 (tadpoles); San Fernando, MZTG 15, 17; Tonina (ruins), KU 41592.

GUATEMALA: Huehuetenango: Jacaltenango, UMMZ 123139 (tadpoles); 0.5 km. E of Jacaltenango, UMMZ 123142-3; 2 km. S of Jacaltenango, UMMZ 123141; 2 km. W of San Pedro Necta, UMMZ 123140 (tadpoles).

=Ptychohyla leonhardschultzei= (Ahl)

Hyla leonhard-schultzei Ahl, Zool. Anz., 106:185-186, fig. 1, April 15, 1934 [Holotype.--ZMB 34353 from Malinaltepec, Guerrero, Mexico; Leonhard Schultze collector]. Smith and Taylor, Bull. U. S. Natl. Mus., 184:87, June 17, 1948.

Hyla godmani, Ahl, Zool. Anz., 106:186, April 15, 1934.

Hyla pinorum Taylor, Proc. Biol. Soc. Washington, 50:46-48, pl. 2, fig. 2, April 21, 1937 [Holotype.--UIMNH 25049 from Agua del Obispo, Guerrero, Mexico; Edward H. Taylor collector]. Smith and Taylor, Bull. U. S. Natl. Mus., 194:87, June 17, 1948.

Ptychohyla adipoventris Taylor, Univ. Kansas Sci. Bull., 30:41-45, May 15, 1944 [Holotype.--UIMNH 25047 from Agua del Obispo, Guerrero, Mexico; Edward T. Taylor collector]. Smith and Taylor, Bull. U. S. Natl. Mus., 194:91, June 17, 1948. Taylor, Amer. Mus. Novitates, 1437:16, December 7, 1949. Stuart, Proc. Biol. Soc. Washington, 67:169, August 5, 1954.

Hyla milleri Shannon, Proc. U. S. Natl. Mus., 101:473-477, figs. 92b, 93a-c, May 17, 1951 [Holotype.--USNM 123700 from San Lucas Camotlan, Oaxaca, Mexico; Walter S. Miller collector].

Ptychohyla leonhard-schultzei, Duellman, Herpetologica, 16:191-197, figs. 1-3, September 23, 1960; Univ. Kansas Publ. Mus. Nat. Hist., 13:351, April 27, 1961.

Diagnosis.--Rostral keel present; snout in lateral profile not rounded above; interorbital region much broader than eyelid; distal subarticular tubercle beneath fourth finger bifid or double; no white stripe on edge of upper lip; flanks white with black spots.

Description.--The following description is based on KU 64117 from Vista Hermosa, Oaxaca, Mexico (Pl. 14). Adult male having a snout-vent length of 35.6 mm.; tibia length, 18.0 mm.; tibia length/snout-vent length, 50.5 per cent; foot length, 14.3 mm.; head length, 10.7 mm.; head length/snout-vent length, 30.1 per cent; head width, 10.6 mm.; head width/snout-vent length, 29.8 per cent; diameter of eye, 3.6 mm.; diameter of tympanum, 1.8 mm.; tympanum/eye, 50.0 per cent. Snout in lateral profile square, not rounded above, and in dorsal profile rounded with pointed tip resulting from vertical rostral keel; canthus pronounced; loreal region barely concave; lips thick, rounded, and barely flaring; nostrils protuberant; internarial distance, 3.2 mm.; top of head flat; interorbital distance, 3.8 mm., and approximately a fifth broader than width of eyelid, 3.2 mm. A moderately heavy dermal fold from posterior corner of eye above tympanum and curving ventrad to anterior edge of insertion of forelimb, covering upper edge of tympanum; tympanum round, its diameter equal to its distance from eye. Forearm moderately robust, having distinct dermal fold on wrist; row of small, low, rounded tubercles along ventrolateral surface of forearm; pollex only slightly enlarged, bearing triangular patch of small horn-covered spines (56 on right, 62 on left); second finger noticeably shorter than fourth; subarticular tubercles round, distal ones on third and fourth toes bifid; discs moderate in size, that of third finger slightly larger than diameter of tympanum; no web between first and second fingers; other fingers one-third webbed. Heels broadly overlap when hind limbs adpressed; tibiotarsal articulation reaches to middle of eye; a low rounded tarsal fold on distal half of tarsus; inner metatarsal tubercle elevated, flat, and elliptical; outer metatarsal tubercle at base of fourth toe, round; row of low, sometimes indistinct, tubercles from heel to base of fifth toe; subarticular tubercles round; length of digits from shortest to longest 1-2-3-5-4, third and fifth being about equal in length; third and fifth toes webbed to base of disc; fourth toe webbed to base of penultimate phalanx; discs of toes much smaller than on fingers. Anal opening near dorsal surface of thighs; short anal flap; opening bordered laterally by heavy dermal fold and ventrolaterally by large tubercles. Skin of dorsum and ventral surfaces of forelimbs and shanks smooth; that of throat, belly, and ventral surfaces of thighs granular. Ventrolateral glands moderately developed, reaching axilla but not to groin and broadly separated midventrally. Tongue cordiform, shallowly notched behind and barely free posteriorly; vomerine teeth 4-3, situated on transverse elevations between ovoid inner nares; openings to vocal sac large, one situated along inner posterior edge of each mandibular ramus.

Dorsal ground-color of head, body, and limbs pale tan with large interconnected dark brown blotches on head and body and broad dark brown transverse bands on limbs; dorsal surfaces of first and second fingers and of webbing of hands pale brown; dorsal surfaces of third and fourth fingers dark brown; anterior surfaces of thighs flesh-color; posterior surfaces of thighs brown; dorsal surfaces of tarsi and feet dark brown; narrow white stripe along ventrolateral edges of forearms and tarsi; a faint creamy white stripe above anal opening; axilla white; flanks creamy white, boldly spotted with black; throat and chest white; ventral surfaces of tarsi and feet dark brown; other ventral surfaces dusty cream color; large brown spots on chin and anterior part of abdomen.

In life the dorsum was reddish brown (Orange-Cinnamon) with dark brown (Chocolate) blotches; first and second fingers and webbing on hand pale reddish brown (Cinnamon); webbing on feet dark brown (Clove Brown); flanks pale creamy white (Pale Olive Buff) with dark brown (Bone Brown) spots; iris reddish bronze (Apricot Orange).

Variation.--No noticeable geographic variation is apparent in the few available specimens; variations in proportions are given in Table 1. The distal subarticular tubercle of the fourth finger is either bifid or double in all specimens; that on the third finger usually is bifid, sometimes single. The vertical rostral keel is prominent in all freshly preserved specimens; in some older specimens it is indistinct. The tongue always is notched posteriorly; in some individuals the notch is shallow; in others it is deep.

Some specimens are paler and less boldly marked than the specimen described above. All specimens from Agua del Obispo and some specimens from the northern slopes of the Sierra Madre Oriental in Oaxaca have a pale tan dorsum with brown markings. In most individuals the white color in the axilla extends onto the posterior edge of the upper arm. The creamy white color of the flanks is constant and usually extends slightly dorsad in the inguinal region. The white stripe above, and sometimes continuing down beside, the anal opening varies from a thin indistinct line or row of flecks to a distinct continuous stripe. Two specimens have dark brown spots on the belly; in the others the spots are confined to the flanks and throat.

Description of tadpole.--The following description is based on KU 68556 from 7.5 kilometers south of Yetla, Oaxaca, Mexico (Figs. 4C and 6C). No limb buds; total length, 37.3 mm.; body length, 12.2 mm.; body length/total length, 32.7 per cent; body slightly depressed, barely wider than deep, ovoid in dorsal profile; mouth directed ventrally; eyes small, directed dorsolaterally; nostrils barely protuberant and directed anterolaterally, about midway between snout and eye; spiracle sinistral and posteroventrad to eye; anal tube dextral. Caudal fin low, bluntly rounded posteriorly; greatest depth of caudal musculature about one-half depth of caudal fin; musculature extends nearly to tip of tail.

Mouth large; lips having deep lateral folds; two complete rows of papillae on lips; five to seven rows of papillae laterally; beaks moderately developed, bearing short peglike serrations; moderately wide lateral projections on upper beak; tooth-rows 4/6; upper rows subequal in length; fourth row interrupted medially; length of lower rows 1-4 equal to upper rows; fifth and sixth lower rows shorter; first lower row interrupted medially.

Body brown above; tip of snout brown; venter creamy gray; caudal musculature creamy tan; caudal fin transparent; cream-colored crescent-shaped mark on posterior edge of body; dark brown flecks on caudal musculature and all except anterior two-thirds of ventral caudal fin; large brown square blotches on dorsum of caudal fin; eye reddish brown in life.

Variation.--The variation in size and proportions is given in Table 2. With the exception of one specimen having a short, broken, seventh tooth-row, all specimens have 4/6 tooth-rows that are like those described above. In some specimens the brown blotches on the dorsum of the caudal musculature are fused and marked with cream-colored flecks.

Comparisons.--Ptychohyla leonhardschultzei differs from all other members of the Ptychohyla euthysanota group in having a square snout, and further differs from P. spinipollex in more numerous and smaller nuptial spines and in transverse, instead of posteromedially slanting, vomerine processes between the inner nares. Ptychohyla leonhardschultzei differs from P. euthysanota in having a rostral keel and in having white flanks boldly spotted with black.

All small hylids that are sympatric with Ptychohyla leonhardschultzei are either yellow (Hyla dendroscarta and H. melanomma) or green (Hyla erythromma, which has a red eye, Hyla hazelae, which has a black line on the canthus, and Ptychohyla ignicolor, which has red flash colors on the thighs).

Life History.--This frog has been found along streams in cloud forests and in pine-oak forest. Males call from vegetation along the stream or from rocks in and at the edge of the stream. The call is a single, long, soft "wraack," repeated at intervals of anywhere from several seconds to three or four minutes. Each note has a duration of 0.62 to 0.95 of a second and a rate of 76 to 78 pulses per second; the dominant frequency falls between 2700 and 2800 cycles per second (Pl. 11C).

Tadpoles were found in several streams in northern Oaxaca. A tadpole having a total length of 21.1 mm. has three upper and four lower tooth-rows well developed; the fourth upper and fifth lower rows are weakly present, and the sixth lower row has not started to develop. Two metamorphosed young have snout-vent lengths of 15.2 and 15.5 mm.

Remarks.--Four specific names have been applied to this species. Ahl (1934:185) based his description of Hyla leonhardschultzei on a small, poorly preserved female. Taylor (1944:41) proposed the generic name Ptychohyla for a species (named therein as P. adipoventris) of hylid having ventrolateral glands and horn-covered nuptial spines. Obviously, Taylor was unaware that Hyla leonhardschultzei was the same species. Earlier Taylor (1937:46) described Hyla pinorum. The types of all of these species came from the Pacific slopes of the Sierra del Sur in Guerrero. Examination of the types and other available specimens shows that they are representatives of a single species. The type of Hyla pinorum is an immature male having a snout-vent length of 26.7 mm. All of these specimens have the square snout and black and white flanks characteristic of Ptychohyla leonhard-schultzei. Although Shannon (1951:473) based his description of Hyla milleri on a male having well-developed ventrolateral glands, he overlooked the presence of these glands in his description and discussion of relationships. The acquisition of more specimens from northern Oaxaca has shown that Hyla milleri is the same as Ptychohyla leonhardschultzei.

Distribution.--This species is known from pine-oak forest and cloud forest on the Pacific slopes of the Sierra Madre del Sur in Guerrero and Oaxaca and from the Atlantic slopes of the Sierra Madre Oriental in northern Oaxaca. Specimens have been collected at elevations between 700 and 1650 meters. Probably the species occurs in humid forests at similar elevations around the eastern end of the Mexican Highlands in Oaxaca.

Specimens examined.--MEXICO: Guerrero: Agua del Obispo, CNHM 123489-90, 126651, 106300, MCZ 29639, UIMNH 25047, 25049, USNM 114551; Malinaltepec, ZMB 34351, 34353. Oaxaca: 2.5 km. N of La Soledad, KU 58061; San Lucas Camotlan, UIMNH 3201, USNM 123700-1; Vista Hermosa, KU 64116-7, 64119, 68560 (tadpoles), 71344, 71717-8 (tadpoles), UMMZ 119604; 5 km. S of Yetla, KU 60045 (tadpoles); 7.5 km. S of Yetla, KU 64118, 68556-7 (tadpoles), 68559 (tadpoles), 68561 (2 young), 68630 (skeleton), UMMZ 115514-5, 118863 (tadpoles); 9 km. S of Yetla, KU 68558 (tadpoles).

=Ptychohyla spinipollex= (Schmidt)

Hyla euthysanota, Dunn and Emlen, Proc. Acad. Nat. Sci. Philadelphia, 84:25, March 22, 1932.