Chapter II: Part 2

Intergradation between _Ptychohyla euthysanota_ and _P. macrotympanum_ has not been demonstrated. Currently their ranges are separated by the dry valleys of the Rio Grijalva and Rio Cuilco. The similarity in structure of the adults and tadpoles and the indistinguishable breeding calls are the basis for considering the two populations subspecies.

_Distribution._--This species occurs in mixed pine and broad-leafed forests at elevations of 700 to 1700 meters on the southern slopes of the Chiapan Highlands and Sierra de Cuchumatanes, in Guatemala. These forests are on the south facing slopes north of the valleys of the Rio Grijalva and Rio Cuilco.

_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.

_Hyla spinipollex_ Schmidt, Proc. Biol. Soc. Washington, 49:45-46,
May 1, 1936 [Holotype.--MCZ 21300 from the mountains behind
Ceiba, Depto. Atlantidad, Honduras; Raymond E. Stadelman
collector]. Stuart, Misc. Publ. Mus. Zool. Univ. Michigan,
69:32-34, figs. 5-6, June 12, 1948; Contr. Lab. Vert. Biol.
Univ. Michigan, 45:22, 52, 54, 57, May, 1950; Proc. Biol. Soc.
Washington, 67:169, August 5, 1954.

_Ptychohyla spinipollex_, Stuart, Contr. Lab. Vert. Biol. Univ.
Michigan, 68:48, November, 1954. Duellman, Univ. Kansas Publ.
Mus. Nat. Hist., 13:351, April 27, 1961.

_Diagnosis._--Rostral keel present; snout in lateral profile rounded above; eyelid nearly as wide as interorbital region; flanks white with brown spots; belly spotted; nuptial spines pointed and moderate in size.

_Description._--The following description is based on KU 58054 from Finca Los Alpes, Depto. Alta Verapaz, Guatemala (Pl. 15). Adult male having a snout-vent length of 37.7 mm.; tibia length, 18.2 mm.; tibia length/snout-vent length, 48.2 per cent; foot length, 15.8 mm.; head length, 11.7 mm.; head length/snout-vent length, 31.0 per cent; head width, 11.7 mm.; head width/snout-vent length, 31.0 per cent; diameter of eye, 3.6 mm.; diameter of tympanum, 1.9 mm.; tympanum/eye, 52.7 per cent. Snout in lateral profile nearly square, slightly rounded above, and in dorsal profile rounded with a pointed tip resulting from vertical rostral keel; canthus pronounced; loreal region barely concave; lips thick, rounded, and barely flaring; nostrils protuberant; internarial distance, 3.0 mm.; top of head flat; interorbital distance, 3.7 mm., about equal to width of eyelid, 3.6 mm. A heavy dermal fold from posterior corner of eye above tympanum to point above insertion of forearm, covering upper edge of tympanum; tympanum round, its diameter equal to its distance from eye. Forearm moderately robust, having faint dermal fold on wrist; row of low, rounded tubercles along ventrolateral edge of forearm; pollex only slightly enlarged, bearing triangular patch of moderate-sized, pointed, horn-covered spines (38 on right, 43 on left); second finger noticeably shorter than fourth; subarticular tubercles round, distal one on fourth finger bifid; discs of fingers of moderate size, that of third finger slightly smaller than diameter of tympanum; vestigial web between first and second fingers; other fingers one-third webbed. Heels broadly overlapping when hind limbs adpressed; tibiotarsal articulation reaches to middle of eye; distinct, but low, tarsal fold extending length of tarsus; inner metatarsal tubercle elevated, flat, and elliptical; outer metatarsal tubercle small and round, near base of fourth toe; row of low indistinct tubercles from heel to base of fifth toe; subarticular tubercles round; length of toes from shortest to longest 1-2-3-5-4, the 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 slightly smaller than those on fingers. Anal opening near upper edge of thighs; opening bordered laterally by moderately heavy dermal folds and ventrolaterally by tubercles. Skin of dorsum and ventral surfaces of forelimbs and shanks smooth; that of throat, belly, and ventral surfaces of thighs granular. Ventrolateral glands barely evident. Tongue ovoid, barely notched behind and slightly free posteriorly; vomerine teeth 2-3, situated on V-shaped elevations between round 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 grayish tan with dark brown reticulations on head and body and dark brown transverse bars or spots on limbs; first and second fingers cream color; third and fourth fingers and webbing on hands pale grayish brown; anterior surfaces of thighs reddish tan; posterior surfaces of thighs yellowish tan; tarsi and toes pale grayish tan with brown flecks; webbing on foot pale brown; faint white stripe along ventrolateral edges of tarsi and forearms; narrow white line above and beside anal opening; no white stripe on edge of upper lip; axilla pale flesh-color; throat, chest, and ventral surfaces of limbs pale creamy gray; belly white with scattered brown flecks; flanks grayish white with dark brown flecks; ventral surfaces of tarsi dark brown; ventrolateral glands grayish tan.

In life the dorsal ground-color of the head, body, fore limbs, and thighs was yellowish tan (Pinkish Buff); dorsal surfaces of shanks and tarsi pale yellowish tan (Pale Pinkish Buff); markings on head and back brown (Snuff Brown) to dark brown (Chocolate); dark bands on limbs brown (Tawny-Olive); first and second fingers creamy tan (Light Pinkish Cinnamon); posterior surfaces of thighs creamy tan (Light Pinkish Cinnamon); third and fourth fingers and webbing on hand grayish brown (Avellaneous); webbing on feet dark brown (Olive Brown); axilla pale pink (Hydrangea Pink); flanks buff (Cream-Buff) becoming yellow (Lemon Chrome) in groin; spots on flanks dark brown (Clove Brown); iris dull grayish bronze (Orange-Citrine).

_Variation._--The distal subarticular tubercle beneath the fourth finger is bifid in about two-thirds of the specimens; in the rest it is round. The posterior edge of the tongue varies from being emarginate to shallowly notched. In most specimens the row of tubercles along the outer edge of the tarsus is made up of discrete tubercles, but in some individuals the tubercles form a nearly continuous dermal fold. Most specimens have the vomerine teeth situated on V-shaped elevations, but in some individuals the elevations are more nearly transversely situated between the inner nares.

All 42 specimens from Finca Los Alpes, Guatemala, have dark brown spots and flecks on the venter. Some individuals have only a few flecks on the throat and a few large spots on the flanks, as does KU 64125. Other specimens, such as KU 64132, have dense spotting over the entire venter. The color of the dorsum varies from pale tan to dark brown with darker brown markings; the white line above the anus is present in all specimens, but in some it is indistinct. KU 58053 and 64127 have a dark brown dorsum with large pale tan, square blotches; in life the blotches were pale tan (Pinkish Buff); the rest of the dorsum was dark brown (Sayal Brown). KU 58052 is dark brown with many small white flecks on the dorsum; in life the dorsum was deep olive brown (Dark Olive).

Aside from the differences mentioned above, all specimens from Guatemala are similar in coloration. Three specimens from Honduras (MCZ 21300 and UMMZ 113102-3) have unspotted white venters. MCZ 21300, the holotype of _P. spinipollex_, lacks a white stripe above the anal opening, whereas the stripe is indistinct in UMMZ 113102-3.

_Description of tadpole._--The following description is based on KU 60053 from Fina Los Alpes, Depto. Alta Verapaz, Guatemala (Figs. 4D and 6D). No limb buds; total length, 37.2 mm.; body length, 12.2 mm.; body length/total length, 32.8 per cent. Body rounded, not depressed, as wide as deep, ovoid in dorsal profile; mouth directed ventrally; eyes small, directed dorsolaterally; nostrils barely protuberant and directed anterolaterally, somewhat closer to eye than snout; 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; six or seven rows of papillae laterally; beaks moderately developed, bearing fine blunt serrations; slender lateral projections on upper beak; tooth-rows 4/7; upper rows subequal in length; fourth row interrupted medially; lower rows 1-4 equal in length to upper rows; lower rows 5-7 decreasing in length; first lower row interrupted medially.

Top of head and tip of snout brown; venter creamy gray; caudal musculature tan; caudal fin transparent; faint cream-colored, narrow, crescent-shaped mark on posterior edge of body, not bordered posteriorly by dark brown mark; brown flecks scattered on caudal musculature and caudal fin; only a few flecks on anterior half of ventral caudal fin; eye bronze-color in life.

_Variation._--The variation in size and proportions as compared with tadpoles of other species is given in Table 2. Of the 57 tadpoles of this species that I have examined, 21 have only six lower tooth-rows, although in some of these specimens a faint ridge for a seventh row is present. In those specimens having seven lower rows, the seventh often is broken.

There is considerable variation in coloration. None has a distinct cream-colored, crescent-shaped mark bordered posteriorly by a dark brown bar or triangle, as in the other species in the _Ptychohyla euthysanota_ group. Most specimens have a rather indistinct crescent; some have no crescent. In a few specimens there is a weakly outlined dark mark posterior to the crescent. Some specimens in a series of tadpoles from 32 kilometers north of Morazan, Baja Verapaz, Guatemala, have faint dorsal blotches on the dorsal musculature, much like those in tadpoles of _Ptychohyla leonhardschultzei_.

_Comparisons._--_Ptychohyla spinipollex_ differs from all other species in the genus by having moderate-sized, instead of small, pointed nuptial spines; also it has fewer spines than the other species (see discussion of this character in Analysis of Data). The nearly equal interorbital breadth and width of the upper eyelid also is diagnostic of this species.

Other hylids sympatric with _Ptychohyla spinipollex_ include three species of _Plectrohyla_, each of which has a bony prepollex, heavy body, and rugose skin on the dorsum. The only other sympatric hylid that could be confused with _Ptychohyla spinipollex_ is _Hyla bromeliacea_, which has a round snout and yellowish tan dorsum not marked with dark brown.

_Life History._--At Finca Los Alpes, Guatemala, in July, 1960, and in August, 1961, calling males were found on bushes and trees along cascading mountain streams. The breeding call consists of a soft "wraack," repeated at intervals of 45 seconds to four minutes. Each note has a duration of about .46 of a second and a rate of 147 pulses per second. The dominant frequency is 4300 cycles per second (Pl. 11A).

Tadpoles have been found in cascading mountain streams. Two metamorphosed young have snout-vent lengths of 15.0 and 15.5 mm.

_Remarks._--There is little doubt that all of the specimens herein referred to _Ptychohyla spinipollex_ are conspecific. However, the three specimens from Honduras, including the type of _Ptychohyla spinipollex_, differ from Guatemalan specimens in lacking all dark spotting on the venter. Additional specimens from Honduras and eastern Guatemala may show that two subspecies are recognizable, in which case the nominal subspecies will be the population in Honduras.

_Distribution._--This species lives in cloud forests at elevations of 800 to 1700 meters on the Atlantic side of the Guatemalan Highlands from the Sierra de Cuchumatanes in western Guatemala eastward to central Honduras.

_Specimens examined._--GUATEMALA: _Alta Verapaz_: Finca Chichen, UMMZ 90876 (tadpoles); Finca Los Alpes, KU 58052-60, 59939 (skeleton), 60053 (tadpoles), 64122-41, 68562, 68563 (tadpoles), 68631-2 (skeletons), MCZ 35000-1, UMMZ 90873, 90874 (3), 90875 (tadpoles); La Primavera, UMMZ 90877 (tadpoles); Panzamala, UMMZ 90878 (tadpoles). _Baja Verapaz_: 32 km. N of Morazan, KU 68564 (tadpoles); _Santa Elena_, UMMZ 98119, 98120 (2). _Huehuetenango_: 1 km. E of Barillas, UMMZ 123136-7 (tadpoles). _Progreso_: Finca Bucaral, UMMZ 106783 (3), 123138 (tadpoles), S-1292 (skeleton).

HONDURAS: _Atlantidad_: Mountains behind Ceiba, MCZ 21300. _Morazan_: Cerro Uyuca, UMMZ 123102-3.

The _Ptychohyla schmidtorum_ Group

Two species in group; adults having only vestige of web between fingers and lacking tarsal fold; pollex of breeding males lacking spinous, horny, nuptial tuberosities; mouth of tadpole greatly expanded, funnel-shaped, lacking lateral folds, and having 3/3 tooth-rows; breeding call consisting of series of short notes.

=Ptychohyla schmidtorum=

_Diagnosis._--Diameter of tympanum more than half of diameter of eye; internarial region depressed; toes three-fourths webbed; no red flash-colors on thighs.

=Ptychohyla schmidtorum schmidtorum= Stuart

_Ptychohyla schmidtorum_ Stuart, Proc. Biol. Soc. Washington,
67:169-172, August 5, 1954 [Holotype.--CNHM 27055 from El
Porvenir (17 kilometers air-line west of San Marcos), Depto.
San Marcos, Guatemala; Karl P. Schmidt collector]. Duellman,
Univ. Kansas Publ. Mus. Nat. Hist., 13:351, 355, April 27, 1961.

_Diagnosis._--Vomerine teeth 5-11; dorsum dark brown; white spot below eye; eye red in life.

_Description._--The following description is based on KU 58043 from Finca La Paz, Depto. San Marcos, Guatemala (Pl. 16). Adult male having snout-vent length of 31.6 mm.; tibia length, 15.0 mm.; tibia length/snout-vent length, 47.5 per cent; foot length, 12.5 mm.; head length, 10.2 mm.; head length/snout-vent length, 32.3 per cent; head width, 9.9 mm.; head width/snout-vent length, 31.3 per cent; diameter of eye, 3.4 mm.; diameter of tympanum, 1.8 mm.; tympanum/eye, 52.9 per cent. Snout in lateral profile nearly square, slightly rounded above and below, and in dorsal profile bluntly squared; canthus pronounced; loreal region concave; lips thick, rounded, and flaring; nostrils protuberant; internarial distance, 2.2 mm.; internarial region depressed; top of head flat; interorbital distance, 3.4 mm., much greater than width of eyelid, 2.5 mm. Thin dermal fold from posterior corner of eye above tympanum to insertion of forelimb, covering upper edge of tympanum; tympanum round, its diameter equal to its distance from eye. Forearm slender, lacking distinct dermal fold on wrist; row of low rounded tubercles along ventrolateral edge of forearm; pollex slightly enlarged; no nuptial spines; second and fourth fingers about equal in length; subarticular tubercles small and round, distal one beneath fourth finger bifid; discs small, that of third finger noticeably smaller than tympanum; no web between first and second fingers; vestige of web between other fingers. Heels overlap when hind limbs adpressed; tibiotarsal articulation reaches to middle of eye; no tarsal fold; inner metatarsal tubercle large, flat, and elliptical; outer metatarsal tubercle small, ovoid, slightly more distal than inner; subarticular tubercles round; length of digits from shortest to longest 1-2-5-3-4; third and fifth toes webbed to base of discs; fourth toe webbed to base of penultimate phalanx; discs of toes smaller than on fingers. Anal opening directed posteriorly at upper edge of thighs; no anal flap; pair of large tubercles below anal opening and smaller tubercles farther below. Skin of dorsum and ventral surfaces of forelimbs and shanks smooth; that of belly and ventral surfaces of thighs granular. Ventrolateral glands well developed, reaching axilla and groin and narrowly separated on chest. Tongue ovoid, emarginate posteriorly, and only slightly free behind; vomerine teeth 3-3, situated on small triangular elevations between ovid inner nares; openings to vocal sac large, one situated along inner posterior edge of each mandibular ramus.

Dorsum of head, body, and limbs reddish brown with indistinct, irregular darker brown markings on body and dark brown transverse bands or spots on limbs; first and second fingers creamy white; third and fourth fingers brown; dorsal surfaces of tarsi and third, fourth, and fifth toes tan with brown spots; first and second toes and webbing on feet creamy tan; enamel-white stripe along edge of upper lip continuing over, and on posterior edge of, forearm to groin, expanded to form spot below eye; belly white, unspotted; ventrolateral glands pale brown; ventral surfaces of hind limbs and anterior and posterior surfaces of thighs cream color; enamel-white stripe on heel; creamy white stripe along ventrolateral edges of tarsi and forearms.

In life dorsum reddish brown (Terra Cotta) with dark brown (Burnt Umber) markings; first and second fingers and first and second toes orange-yellow (Light Orange-Yellow); posterior surfaces of thighs pale reddish tan (Ochraceous-Salmon); webbing on feet yellowish tan (Deep Colonial Buff); belly white; iris red (Nopal Red).

_Variation._--Little variation in structural characters was observed. All but five specimens have bifid subarticular tubercles beneath the fourth finger. Three specimens have cordiform tongues, and in four others the tongue is ovoid and shallowly notched behind; all other specimens have an emarginate ovoid tongue.

Some individuals when active at night had a pale brown (Ochraceous-Tawny) dorsum with dull olive green (Dark Olive Buff) markings. Otherwise there was no noticeable variation in color.

_Description of tadpole._--The following description is based on KU 60051 from Finca La Paz, Depto. San Marcos, Guatemala (Figs. 5A and 6E). Small hind limbs; total length, 37.9 mm.; body length, 11.6 mm.; body length/total length, 30.6 per cent. Body only slightly depressed, nearly as deep as wide, in dorsal profile ovoid, widest just posterior to eyes; in lateral profile snout rounded; mouth directed ventrally; eyes small, directed dorsolaterally; nostrils barely protuberant, directed anteriorly, somewhat closer to eye than snout; spiracle sinistral and posteroventrad to eye; anal tube dextral. Tail long and slender; caudal fin low and rounded posteriorly; depth of caudal musculature one-half greatest depth of caudal fin; musculature not extending to tip of tail.

Mouth large; thin fleshy lips greatly expanded and forming large funnel-shaped disc; width of mouth two-thirds greatest width of body; outer edge of lips having one row of small papillae; inner surfaces of mouth smooth; scattered large papillae forming one nearly complete row around teeth; other large papillae laterally; beaks moderately developed, bearing long, pointed denticulations; no lateral projections on upper beak; tooth-rows 3/3, all short; second and third upper rows subequal in length; first upper row shorter; first and third upper rows interrupted medially; first lower row interrupted medially, equal in length to second and third upper rows; second lower row slightly shorter; third lower row shortest.

Body mottled brown and creamy gray above and below; mouth colored like body; caudal musculature creamy tan; caudal fin transparent; dark brown streak mid-laterally on anterior third of caudal musculature; rest of tail and all of caudal fin heavily flecked with brown; eye red in life.

_Variation._--The third upper tooth-row is interrupted in all specimens; in some individuals the first upper and first lower rows are complete. The variation in size and proportions is given in Table 2. The dark brown lateral streak on the anterior part of the caudal musculature is distinct on most specimens; the only other variation in coloration is in the amount of brown flecking on the caudal musculature and fin.

_Comparisons._--_Ptychohyla schmidtorum schmidtorum_ differs from _P. schmidtorum chamulae_ as stated in the diagnosis and in having pale creamy tan, as opposed to dark brown, webbing on the feet; and from _P. ignicolor_ in having a depressed, as opposed to a flat, internarial region. Tadpoles of _P. s. schmidtorum_ have a mottled appearance, as opposed to the more uniform brown color of _P. s. chamulae_.

_Ptychohyla schmidtorum schmidtorum_ and several species of _Plectrohyla_ are sympatric. All species of the latter genus have a bony prepollex, rugose skin on the dorsum, and heavy body; also sympatric is _Ptychohyla e. euthysanota_, which has a tarsal fold and in breeding males spinous nuptial tuberosities.

_Life History._--This species breeds in clear mountain streams where males call from vegetation along the stream. The call consists of series of short notes, three to nine notes per series, sounding like "raa-raa-raa." The duration of each note is approximately .065 of a second, and has a rate of 96 to 119 pulses per second; the dominant frequency is about 3400 cycles per second. The call is almost indistinguishable from that of _Ptychohyla schmidtorum chamulae_.

Tadpoles and metamorphosing young were found at Finca La Paz, Guatemala, in late July, 1960. Two young lacking tails but not having completely developed mouths have snout-vent lengths of 14.2 and 14.6 mm. L. C. Stuart collected four metamorphosing young at Finca La Paz on May 6, 1949. By May 10 the frogs were completely metamorphosed, at which time they had snout-vent lengths of 15.5 to 17.0 (average 16.1) mm.

_Remarks._--There is no doubt that this frog is most closely related to _Ptychohyla schmidtorum chamulae_, even though the ranges of the two subspecies are separated by the interior depression of Chiapas. Since at least at Finca La Paz, Guatemala, _P. s. schmidtorum_ occurs with _P. e. euthysanota_, it is surprising that the former species has not been found at more localities along the Pacific slopes on northern Central America. At Finca La Paz in July, 1960, _P. s. schmidtorum_ was more abundant than _P. e. euthysanota_.

_Distribution._--This species is known only from a limited area at elevations between 1300 and 2200 meters on the Pacific slopes of the Sierra Madre in extreme eastern Chiapas and western Guatemala.

_Specimens examined._--MEXICO: _Chiapas_: Finca Irlandia, UMMZ 105429-30.

GUATEMALA: _San Marcos_: El Porvenir, CNHM 20755, 20761, 69904, UMMZ 80918; Finca La Paz, 2 km. W of La Reforma, KU 58016-44, 59940-2 (skeletons), 60050 (3 young), 60051 (tadpoles), 60052 (4 young), MCZ 34998-9, UMMZ 123144-7 (tadpoles).

=Ptychohyla schmidtorum chamulae= Duellman

_Ptychohyla chamulae_ Duellman, Univ. Kansas Publ. Mus. Nat. Hist.,
13: 354-357, pl. 25, fig. 2, April 27, 1961 [Holotype.--KU
58063 from 6.2 kilometers south of Rayon Mescalapa, Chiapas,
Mexico; William E. Duellman collector].

_Diagnosis._--Vomerine teeth 4-6; dorsum bright green; white lateral stripe; eye reddish bronze in life.

_Description._--The following description is based on KU 58069 from 6.2 kilometers south of Rayon Mescalapa, Chiapas, Mexico (Pl. 17). Adult male having a snout-vent length of 27.6 mm.; tibia length, 13.0 mm.; tibia length/snout-vent length, 47.1 per cent; foot length, 10.8 mm.; head length, 9.6 mm.; head length/snout-vent length, 34.7 per cent; head width, 9.2 mm.; head width/snout-vent length, 33.1 per cent; diameter of eye, 3.0 mm.; diameter of tympanum, 1.6 mm.; tympanum/eye, 53.3 per cent. Snout in lateral profile nearly square, slightly rounded above and below, and in dorsal profile blunt, almost square; canthus pronounced; loreal region concave; lips thick, rounded and flaring; nostrils protuberant; internarial distance, 2.5 mm.; internarial region slightly depressed; top of head flat; interorbital distance, 3.3 mm., much greater than width of eyelid, 2.5 mm. Thin dermal fold, from posterior corner of eye above tympanum to insertion of fore limb, covering upper edge of tympanum; tympanum nearly round, its diameter equal to its distance from eye. Forearm slender, lacking distinct fold on wrist; row of low, rounded tubercles on ventrolateral surface of forearm; pollex slightly enlarged, without nuptial spines; second and fourth fingers equal in length; subarticular tubercles round, that under fourth finger bifid; discs small, that of third finger noticeably smaller than tympanum; no web between first and second fingers; vestige of web between other fingers. Heels overlapping when hind limbs adpressed; tibiotarsal articulation reaches to middle of eye; no tarsal fold; inner metatarsal tubercle large, flat, and elliptical; outer metatarsal tubercle small, elliptical, slightly more distal than inner; subarticular tubercles round; length of digits from shortest to longest 1-2-5-3-4; third and fifth toes webbed to base of disc; fourth toe webbed to base of penultimate phalanx; discs smaller on toes than on fingers. Anal opening directed posteriorly at upper edge of thighs; no anal flap; pair of large tubercles below anal opening. Skin of dorsum and of ventral surfaces of forelimbs and shanks smooth; that of throat, belly, and ventral surfaces of thighs granular. Ventrolateral glands well developed, not reaching axilla or groin and broadly separated midventrally. Tongue cordiform, shallowly notched behind and only slightly free posteriorly; vomerine teeth 2-3, situated on small triangular elevations between ovoid inner nares; openings to vocal sac large, one situated along inner posterior edge of each mandibular ramus.

Dorsum of head, body and limbs reddish brown with dark purplish brown markings on back and shanks; first finger creamy tan; other fingers pale brown; dorsal surfaces of tarsi, third, fourth, and fifth toes dull tan with brown spots; first and second toes creamy tan; webbing on feet brown; anterior and posterior surfaces of thighs tan; faint creamy white stripe along ventrolateral edges of tarsi and forearms; enamel-white stripe on heel; axilla and groin gray; enamel-white stripe on edge of upper lip, continuing onto proximal upper surfaces of forelimb and on flanks to groin, widened under eye to form large spot, and bordered below on flanks by dark brown stripe; white stripe above and white spots below anal opening; throat and chest white; belly and ventral surfaces of limbs cream color; brown dash on either side of chin and brown spot on throat near angle of jaws; few brown flecks on belly; ventrolateral glands orange-tan; ventral surfaces of tarsi and feet brown.

In life, dorsal surfaces of head, body, and limbs bright green (Shamrock Green); first and second fingers pale orange (Apricot Yellow); stripe on upper lip and spot below eye enamel-white; stripe on flanks silvery white, bordered below by brown (Saccardo's Umber) brown; anterior and posterior surfaces of thighs yellowish brown (Old Gold); webbing of feet dull brown (Brownish Olive); belly deep yellow (Amber Yellow); iris reddish bronze (English Red).

_Variation._--Tubercles beneath the fourth fingers are bifid in 20 specimens and rounded in all others. The tongue is emarginate in 12 specimens and cordiform in all others. In most specimens the white stripe on the upper lip continues onto the flanks and to the groin; in five specimens the stripe terminates above the forearm, and in three it terminates at mid-flank. The lateral stripe is absent in two specimens. All specimens were uniform green above when found at night; later some changed to pale green (Light Oriental Green) on the dorsum with irregular yellowish tan (Naples Yellow) blotches. Most males have brown flecks on the throat and ventrolateral gland, but some specimens are immaculate below, and one has dark brown mottling on the throat. Several males have a round, orange-tan glandular area on the chin, as does _P. ignicolor_.

_Description of tadpole._--The following description is based on KU 58199 from 6.2 kilometers south of Rayon Mescalapa, Chiapas, Mexico (Figs. 5B and 6F). Hind limbs small; total length, 39.0 mm.; body length, 10.5 mm.; body length/total length, 26.9 per cent. Body barely depressed, only slightly wider than deep, widest just posterior to eyes; in dorsal profile ovoid; mouth directed ventrally; eyes small, directed dorsolaterally; nostrils barely protuberant, directed anterodorsally, slightly closer to eye than snout; spiracle sinistral and posteroventrad to eye; anal tube dextral. Tail long and slender; caudal fin low, rounded posteriorly; depth of caudal musculature one-half greatest depth of caudal fin; musculature not extending to tip of tail.

Mouth large; thin fleshy lips greatly expanded and forming funnel-shaped disc; outer edge of lips having one row of small papillae; inner surfaces of mouth smooth; scattered large papillae forming nearly one complete row around teeth; other papillae laterally; beaks moderately developed, bearing long, pointed denticulations; no lateral projections on upper beak; tooth-rows 3/3, all short; second and third upper rows subequal in length; first upper row shorter; first and third upper rows interrupted medially; first lower row interrupted medially, equal in length to second and third upper rows; second lower row slightly shorter; third lower row shortest.

Body dark brown above and dark gray below; fleshy part of mouth creamy gray mottled with dark brown; caudal musculature pale tan with heavy suffusion of brown flecks; caudal fin transparent with brown spots; dark brown streak mid-laterally on anterior one-fifth of caudal musculature, bordered below by cream-colored spot; eye brown in life.

_Variation._--The third upper tooth-row is interrupted in all specimens, but in some individuals the first upper row and first lower row are complete. The only noted variation in color is the intensity of brown pigmentation on the caudal musculature, which in most specimens is sufficiently dense to make the tail look brown. In some specimens the mid-lateral streak is indistinct, and the pale spot below the streak is absent.

_Comparisons._--Aside from the characters listed in the diagnosis, _Ptychohyla schmidtorum chamulae_ differs from _P. schmidtorum schmidtorum_ by having dark brown webbing on the feet, instead of pale creamy tan webbing, and in having in life a yellow venter, instead of a white venter. _Ptychohyla ignicolor_ also is green in life, but has red flash-colors on the thighs, red webbing on the feet, and lacks the white lateral stripe diagnostic of _P. schmidtorum chamulae_.

_Plectrohyla matudai matudai_ and _P. guatemalensis_ are sympatric with _Ptychohyla schmidtorum chamulae_. Each of the first two has a bony prepollex, rugose skin on the dorsum, and heavy body. Also living with _Ptychohyla chamulae_ are _Hyla chaneque_, a large species having a tuberculate dorsum and webbed fingers, and _Hyla bivocata_, a small yellow species having a broad, flat head, small indistinct tympanum, and an axillary membrane.

_Life History._--Calling males were found on leaves of herbs and bushes by cascading streams in cloud forest. The call consists of series of short notes, three to nine notes per series, sounding like "raa-raa-raa." The duration of each note is .054 to .070 of a second, and has a rate of 96 to 110 pulses per second. The dominant frequency falls between 3350 and 3450 cycles per second (Pl. 11D). The call is almost indistinguishable from that of _Ptychohyla schmidtorum schmidtorum_.

Tadpoles were found in the cascading streams; the smallest tadpole has a total length of 17.2 mm. and has only 3/2 tooth-rows. At a stream 6.2 kilometers south of Rayon Mescalapa, Chiapas, metamorphosing young were found on June 16 and August 5, 1960. Each of two completely metamorphosed young has a snout-vent length of 15.7 mm. Another having a snout-vent length of 16.2 mm. has a tail stub 2 mm. long and a completely metamorphosed mouth. Two others have snout-vent lengths of 13.6 and 14.1 mm. and tail lengths of 11.5 and 8.1 mm. respectively; in these the mouth parts are incompletely metamorphosed.

_Remarks._--Duellman (1961:354) described _Ptychohyla chamulae_ and stated that it probably was most closely related to _P. schmidtorum_. Further study has revealed additional resemblance in morphological and behavioral details. It is concluded that the two populations are more realistically treated as subspecies than as species. The geographic ranges, as now known, are disjunct. _Ptychohyla schmidtorum chamulae_ inhabits cloud forest on the Atlantic slopes of the Chiapan Highlands, whereas _P. s. schmidtorum_ lives in cloud forest on the Pacific slopes of the Sierra Madre in Chiapas and Guatemala. Between their known geographic ranges are the pine clad Sierra Madre and Chiapan Highlands, and intervening sub-humid Grijalva Valley.

_Distribution._--This species is known only from elevations between 1500 and 1700 meters on the Atlantic slopes of the Chiapan Highlands; it is to be expected in cloud forests on the northern slopes of the Sierra de Cuchumatanes in Guatemala.

_Specimens examined._--MEXICO: _Chiapas: 15 km. N of Pueblo Nuevo Solistahuacan_, UMMZ 123325 (4); _16.5 km. N of Pueblo Nuevo Solistahuacan_, UMMZ 123322 (10); _18 km. N of Pueblo Nuevo Solistahuacan_, UMMZ 121395-9, 123324 (8), 123326 (5); _18.6 km. N of Pueblo Nuevo Solistahuacan_, UMMZ 123323 (4); _5.6 km. S of Rayon Mescalapa_, KU 58062, 58200 (tadpoles); _6.2 km. S of Rayon Mescalapa_, KU 58063-74, 58199 (tadpole), 58234-8, 59936 (skeleton).

=Ptychohyla ignicolor= Duellman

_Ptychohyla ignicolor_ Duellman, Uni. Kansas Publ. Mus. Nat. Hist.,
13:352-353, pl. 25, fig. 1, April 27, 1961 [Holotype.--UMMZ
119603 from 6 kilometers south of Vista Hermosa, Oaxaca,
Mexico; Thomas E. Moore collector].

_Diagnosis._--Diameter of tympanum less than one-half diameter of eye; internarial region flat; 3-7 vomerine teeth; toes one-half webbed; no white spot below eye; no lateral white stripe; in life dorsum green; groin and thighs having bright red flash-colors.

_Description._--The following description is based on UMMZ 119603 from 6 kilometers south of Vista Hermosa, Oaxaca, Mexico (Pl. 18). Adult male having a snout-vent length of 30.0 mm.; tibia length, 14.6 mm.; tibia length/snout-vent length, 48.7 per cent; foot length, 12.3 mm.; head length, 9.2 mm.; head length/snout-vent length, 30.7 per cent; head width, 9.3 mm.; head width/snout-vent length, 31.0 per cent; diameter of eye, 3.2 mm.; diameter of tympanum, 1.3 mm.; tympanum/eye, 40.6 per cent. Snout in lateral profile square, and in dorsal profile rounded; canthus pronounced; loreal region slightly concave; lips moderately flaring; top of head flat; nostrils protuberant; internarial distance, 2.8 mm.; internarial region flat; interorbital distance, 3.3 mm., much broader than width of eyelid, 2.8 mm. A heavy dermal fold from posterior corner of eye above tympanum to insertion of forelimb, covering upper edge of tympanum; tympanum elliptical, its greatest diameter equal to its distance from eye. Forearm moderately robust having distinct dermal fold on wrist; pollex moderately enlarged without nuptial spines; second and fourth fingers equal in length; subarticular tubercles round, none is bifid; discs on fingers moderate in size, that on third finger slightly larger than tympanum; no web between first and second fingers; vestige of web between other fingers. Heels overlap when hind limbs adpressed; tibiotarsal articulation extends to anterior corner of eye; no tarsal fold; inner metatarsal tubercle large, flat, and elliptical; outer metatarsal tubercle near inner one and triangular in shape; subarticular tubercles round; length of digits from shortest to longest 1-2-5-3-4; third toe webbed to proximal end of penultimate phalanx; fourth toe webbed to distal part of antepenultimate phalanx; fifth toe webbed to middle of penultimate phalanx; discs on toes smaller than on fingers. Anal opening directed posteriorly at upper edge of thighs; no anal flap; pair of large tubercles below anal opening; small tubercles ventral and lateral to these. Skin of dorsum and ventral surfaces of limbs smooth; that of throat and belly granular. Ventrolateral glands noticeably thickened, extending from axilla nearly to groin and only narrowly separated midventrally on chest; skin of anterior part of chin thickened and glandular. Tongue cordiform, shallowly notched behind and only slightly free posteriorly; vomerine teeth 0-3, situated on rounded elevations between somewhat larger, round inner nares; openings to vocal sac large, one situated along posterior margin of each mandibular ramus.

Dorsal ground-color of head, body, and limbs dull brown with dark brown reticulations on head and body and dark brown transverse bands or spots on limbs; first and second fingers cream color; third and fourth fingers dull tan; anterior surfaces of thighs pale brown; posterior surfaces of thighs cream color with heavy suffusion of brown; dorsal surfaces of tarsi and third, fourth, and fifth toes dull brown with dark brown spots; first and second toes creamy white; webbing on foot brown; axilla and groin cream color; flanks brown; no white stripes on edge of upper lip or on flank; faint, barely discernible tan streak above anal opening; faint creamy tan line on ventrolateral edges of tarsi; throat, belly, ventral surfaces of limbs, inner edges of tarsi, and first toes cream color; outer ventral surfaces of tarsi and other toes brown; chest and throat spotted with brown; ventrolateral and chin glands orange-brown.

In life the dorsum was uniform green (Cosse Green) becoming paler green (Bright Green-Yellow) on flanks, later changing to paler green (Javel Green) on dorsum with irregular darker green (Lettuce Green) markings and greenish yellow (Green-Yellow) on flanks; anterior and posterior surfaces of thighs, ventral surfaces of shanks, anterior surfaces of tarsi, and upper proximal surfaces of first, second, and third toes red (Coral Red); venter pale creamy yellow (Sulfur Yellow); iris pale golden color (Aniline Yellow).

_Variation._--Of 13 specimens, six have a cordiform tongue; the others have an emarginate tongue. Five specimens have round subarticular tubercles beneath the fourth fingers; six specimens have a bifid tubercle on one hand, and two specimens have bifid tubercles on both hands. A round gland is present on the chin of all specimens; in some the gland is barely visible, but in others it is large and distinct. In two specimens the ventrolateral glands are weakly developed; in the others the glands are well developed and orange-tan. The white anal stripe varies from a thin line to a series of white flecks. Dark brown or black flecks are present on the throat, chest, and flanks of all specimens. In some the flecks are small and widely scattered; in others the flecks are larger and more numerous. All specimens were pale green above when calling at night; later they changed to dull green with darker green reticulations. The flash-colors on the thighs and in the groin vary from red to orange-red or brownish red.

_Description of tadpole._--The following description is based on KU 71716 from Vista Hermosa, Oaxaca, Mexico (Figs. 5C and 6G). Hind limbs small; total length, 39.6 mm.; body length, 11.8 mm.; body length/total length, 29.8 per cent. Body moderately depressed, only slightly wider than deep, in dorsal profile ovoid, widest just posterior to eyes; in lateral profile snout rounded; mouth directed ventrally; eyes small, directed dorsolaterally; nostrils barely protuberant, directed anteriorly, somewhat closer to eye than snout; spiracle sinistral and posteroventrad to eye; anal tube dextral. Tail long and slender; caudal fin low and rounded posteriorly; depth of caudal musculature about one-half greatest depth of caudal fin; musculature not extending to tip of tail.

Mouth large; thin fleshy lips greatly expanded and forming large funnel-shaped disc; width of mouth about two-thirds greatest width of body; outer edge of lips having one row of small papillae; inner surface of mouth smooth; scattered large papillae forming one nearly complete row around teeth; other papillae laterally; one large papilla just above each end of first lower tooth-row; beaks moderately developed bearing long, pointed denticulations; no lateral projections on upper beak; tooth-rows 3/3, all short; second and third upper rows subequal in length; first upper row shorter; first lower row equal in length to second and third upper rows; second lower row slightly shorter; third lower row shortest.

Body creamy gray with dark brown flecks above and below; mouth colored like body; caudal musculature creamy tan; caudal fin transparent; dark brown streak on anterior third of caudal musculature; rest of tail and all of caudal fin, except anterior two-thirds of ventral fin, heavily flecked with brown; iris silvery bronze color in life.

_Variation._--The only other known tadpole, which was collected with the individual described above, differs in having only two upper tooth-rows. The first upper tooth-row seems not to have developed.

_Comparisons._--From _P. schmidtorum chamulae_ and _P. s. schmidtorum_, _P. ignicolor_ differs as follows: Tympanum smaller; snout more nearly square; less webbing on toes; internarial region flat instead of depressed; white lateral stripes lacking.

_Ptychohyla ignicolor_ and several small and moderate sized hylids are sympatric. From _P. ignicolor_ these hylids can be distinguished as follows: _Hyla dendroscarta_ has a round snout and yellow dorsum; _Hyla erythromma_ has a round snout, green dorsum, white flanks, and a red eye; _Hyla hazelae_ has a tarsal fold, green dorsum, and a black line on the canthus; and _Ptychohyla leonhardschultzei_ has a tarsal fold, brown dorsum, black and white flanks, and horny nuptial spines in breeding males.

_Life History._--At Vista Hermosa, Oaxaca, males were calling on vegetation above small streams on March 30, 1959, and on June 28, 1962; males were found on vegetation overhanging a stream 6 kilometers south of Vista Hermosa on June 27 and July 3, 1962. The call consists of a series of short notes, three to thirteen notes per series, sounding like "raa-raa-raa." The duration of each note is about .08 of a second and has a rate of 123 to 129 pulses per second. The dominant frequency of notes in short series is about 2100 cycles per second, whereas the dominant frequency of notes in long series is about 3150 cycles per second (Pl. 11E).

On June 28, 1962, two tadpoles of this species were found in a quiet pool in a spring-fed rivulet at Vista Hermosa, Oaxaca. Females are unknown.

_Remarks._--The absence of a tarsal fold and of nuptial spines in breeding males, the nature of the breeding call, and the form of tadpole are characters that place _Ptychohyla ignicolor_ in the _P. schmidtorum_-group.

_Distribution._--This species is known from only two localities at elevations of 1500 and 1850 meters in the cloud forest on the northern (Atlantic) slopes of the Sierra Madre Oriental in northern Oaxaca.

_Specimens examined._--MEXICO: _Oaxaca: Vista Hermosa_, KU 71334, 71716 (tadpoles), UMMZ 119602; 6 km. S of Vista Hermosa, KU 71335-42, 71343 (skeleton), UMMZ 119603, 123327 (2).

DISTRIBUTION AND ECOLOGY

Geographic Distribution of the Species

The distribution of species of _Ptychohyla_ reflects the distribution of cloud forest in southern Mexico and northern Central America. The frogs are restricted to mountainous areas, usually at elevations higher than 1000 meters above sea level. _Ptychohyla_ does not range to great heights in the mountains, where west of the Isthmus of Tehuantepec the mountain streams are inhabited by frogs of the _Hyla bistincta_ group, and in Chiapas and Guatemala by species of _Plectrohyla_.

Frogs of the _Ptychohyla euthysanota_ group have a greater combined geographic range than the species comprising the _Ptychohyla schmidtorum_ group (Fig. 7). No two species in the same group are sympatric, but members of different groups are sympatric in at least parts of their ranges. Apparently _P. leonhardschultzei_ ranges around the southern edge of the Mexican Highlands, where the species occurs on both Atlantic and Pacific slopes; as can be seen from the distribution map, there are many gaps in the known range of this species. The range of _P. euthysanota euthysanota_ is along the Pacific slopes of the Sierra Madre in Chiapas, Guatemala, and El Salvador, whereas that of _P. euthysanota macrotympanum_ is along the southern interior slopes of the Central Highlands of Chiapas and the Sierra de Cuchumatanes in Guatemala. _Ptychohyla spinipollex_ occurs on the wet Atlantic slopes of the Guatemalan and Honduranean Highlands; the range of the species in Honduras is poorly known.

The frogs of the _Ptychohyla schmidtorum_ group have more restricted geographic ranges than members of the former group. _Ptychohyla schmidtorum schmidtorum_ occurs on the Pacific slopes of the Sierra Madre in Chiapas and Guatemala, where it occurs with _P. euthysanota euthysanota_; _P. schmidtorum chamulae_ is known from only two localities on the Atlantic slopes of the Central Highlands of Chiapas, where it occurs close to, but as now known not with, _P. euthysanota macrotympanum_. On the Atlantic slopes of the Sierra Madre Oriental in northern Oaxaca _P. ignicolor_ occurs with _P. leonhardschultzei_.