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_.

In the Sierra de los Tuxtlas in southern Veracruz and in the cloud forests along the eastern slopes of the Sierra Madre Oriental northward to Nuevo Leon, _Hyla miotympanum_ seems to be the ecological replacement of _Ptychohyla_. On the Pacific slopes north of Guerrero, Mexico, humid forests in which there are cascading mountain streams are absent; consequently, no _Ptychohyla_ are known from that region. In the mountains of El Salvador _Ptychohyla euthysanota euthysanota_ occurs sympatrically with another small stream-breeding hylid, _Hyla salvadorensis_. To the south of Honduras the highlands diminish into the lowlands of Nicaragua, where habitat suitable for _Ptychohyla_ apparently does not exist. In the mountains of Costa Rica and Panama, the habitats occupied by _Ptychohyla_ in northern Central America are filled by a variety of stream-breeding _Hyla_, such as _Hyla legleri_, _H. rivularis_, _H. rufioculis_, _H. alleei_, and _H. uranochroa_.

Although members of the genus _Ptychohyla_ occur in the southern part of the Mexican Highlands to the west of the Isthmus of Tehuantepec, the greater distribution and differentiation in the genus is in the Chiapan-Guatemalan Highlands. In this respect _Ptychohyla_ is a counterpart of _Plectrohyla_.

Habitat Preference

Frogs of the genus _Ptychohyla_ are ecologically associated with mountain streams at elevations between 650 and 2200 meters; in the geographic region where these frogs occur the vegetation between those elevations consists of cloud forest or pine-oak forest. In some places the frogs have been found in a mixture of oak and semi-deciduous scrub forest. At Vista Hermosa, Oaxaca, _P. leonhardschultzei_ and _P. ignicolor_ were found in cloud forest, whereas at Agua del Obispo, Guerrero, the former species was found in pine-oak forest. _Ptychohyla schmidtorum_ is known only from cloud forest; _P. euthysanota euthysanota_ and _P. spinipollex_ generally are found in cloud forest, but in some places they live in pine-oak forest. _Ptychohyla euthysanota macrotympanum_ has been found in pine-oak forest and in a mixture of oak and semi-deciduous scrub forest. With the possible exception of the members of the _Ptychohyla schmidtorum_ group, which has been found only in cloud forest, it seems as though the type of vegetation is not the controlling factor in the ecological distribution of these frogs.

_Ptychohyla_ has been found only where there are clear, cascading streams overhung by vegetation, on which adults and young perch at night, or even by day. The presence of these streams, in which the tadpoles live, seems to be an important factor in the distribution of _Ptychohyla_. As has been shown previously, the tadpoles of _Ptychohyla_ are adapted for existence in torrential streams, where the water is cool, and the amount of oxygen is high. Clearly these tadpoles are unsuited for life in ponds or sluggish streams in the lowlands, where the temperature of the water is high, a layer of silt on the bottom is deep, and the amount of oxygen is low. The tadpoles cling to rocks on the bottom of the streams; there they move slowly across the rocks, apparently feeding on the thin covering of algae. Tadpoles were not observed on rocks having a thick covering of algae or moss. The tadpoles were observed to swim against the current in torrential streams, in which no fishes were found. Therefore, it seems as though the presence of the stream-habitat for the tadpoles is a significant factor in the ecological distribution of the species of _Ptychohyla_.

Interspecific Competition

At localities where two species of _Ptychohyla_ occur sympatrically (_P. ignicolor_ and _P. leonhardschultzei_ at Vista Hermosa, Oaxaca, and _P. euthysanota euthysanota_ and _P. schmidtorum schmidtorum_ at Finca La Paz, Depto. San Marcos, Guatemala) effort was made to determine what, if any, ecological interspecific relationships existed. Although adults of the sympatric species were found on adjacent leaves or branches of bushes overhanging the streams at both localities, segregation at the time of breeding seems to be maintained by the notably different breeding calls in sympatric species (see discussion of breeding calls). Thus, as has been shown by Blair (1956), Fouquette (1960), and others working on a variety of pond-breeding frogs and toads, the breeding call in _Ptychohyla_ acts as an important reproductive isolating mechanism.

At no locality were _Ptychohyla_ and associated species of hylids found so abundantly as were species of pond-breeding hylids in the lowlands. Apparently reproductive activity is not concentrated in a short breeding season, and it is highly doubtful if the populations of these frogs are as large as those of the lowland pond-breeders. The continual humid conditions and abundance of insect food throughout the year in the cloud forest are perhaps indicative of little interspecific competition among adults of _Ptychohyla_ and other sympatric hylids.

At Finca La Paz, Guatemala, tadpoles of two species of _Ptychohyla_ were ecologically segregated. The tadpoles of _P. euthysanota euthysanota_ were found in riffles in the streams, whereas those of _P. schmidtorum schmidtorum_ were found in slower water, chiefly in small pools in the streams. At Vista Hermosa, Oaxaca, tadpoles of _P. leonhardschultzei_ were found in riffles, and tadpoles of the sympatric _P. ignicolor_ were found in a small pool in a stream. Similar ecological relationships were observed for several species of Costa Rican hylids. Throughout the range of _Ptychohyla_ east of the Isthmus of Tehuantepec, members of the genus occur with species of _Plectrohyla_, all of which are larger than _Ptychohyla_, and all of which have tadpoles that live in torrential streams. Tadpoles of _Ptychohyla spinipollex_ have been found in streams inhabited by the tadpoles of _Plectrohyla guatemalensis_ and _P. quecchi_; tadpoles of _Ptychohyla euthysanota euthysanota_ and _P. schmidtorum schmidtorum_ were found in streams inhabited by tadpoles of _Plectrohyla guatemalensis_, _P. matudai_, and _P. sagorum_. In some streams great numbers of tadpoles occur. The habitat is rather restricted, and the food supply is limited. Consequently, interspecific competition among the various species of hylids whose tadpoles live in the torrential streams probably is highest during the larval stage. Unfortunately, this aspect of salientian population ecology has received no intensive study.

Reproduction and Development

Since the cloud forests inhabited by _Ptychohyla_ are daily bathed in clouds and have a fairly evenly distributed rainfall throughout the year, the frogs living in these forests are active throughout the year. At least some of the species evidently have a long breeding season, for I found calling males of _P. leonhardschultzei_ in February, March, and August, and found tadpoles in February, March, June, and August. Tadpoles of the various species have been obtained throughout much of the year, as follows: _P. euthysanota euthysanota_, February, March, May, and July; _P. euthysanota macrotympanum_, March, June, and August; _P. spinipollex_, February, March, April, June, July, and August; _P. schmidtorum schmidtorum_, March, May, June, July, and August; _P. schmidtorum chamulae_, June and August; _P. ignicolor_, June. I suspect that this temporal distribution more accurately reflects the seasonal activities of collectors than of the frogs.

Calling frogs usually are on vegetation adjacent to or overhanging streams; some calling males of _P. spinipollex_ were on rocks in or by streams. Clasping pairs of _P. euthysanota_ and _P. schmidtorum_ were observed on vegetation by streams. Despite intensive search, no eggs were found. It is doubtful if _Ptychohyla_ deposit eggs on vegetation overhanging streams, as do centrolenids and _Phyllomedusa_, for egg-clutches of these frogs are easily found. Possibly the eggs are laid separately on vegetation above the stream, in which case they could be overlooked easily. In streams where _Ptychohyla_ and other hylid tadpoles occur, empty egg capsules have been found on the lee sides of rocks, but there is no way to determine which species laid the eggs.

Numbers of eggs were counted in gravid females; the largest eggs have diameters ranging from 2.5 to 3.0 mm. The smaller species, comprising the _Ptychohyla schmidtorum_ group, have fewer eggs than do the larger species. Numbers of eggs found in females of the various species are: _P. euthysanota euthysanota_, 108; _P. euthysanota macrotympanum_, 136, 160; _P. leonhardschultzei_, 141; _P. spinipollex_, 119, 134, 143; _P. schmidtorum schmidtorum_, 59, 61, 90; _P. schmidtorum chamulae_, 60, 71, 89.

Duration of the larval stage is unknown. Metamorphosing young have been found from May through August. From two to six completely metamorphosed young are available for each of the species, except for _P. ignicolor_ of which none is available. The smallest young frog is a _P. euthysanota_ having a snout-vent length of 14.2 mm.; the largest young frog is a _P. schmidtorum schmidtorum_ having a snout-vent length of 17.0 mm.

PHYLOGENY OF PTYCHOHYLA

The preceding data on morphology, life histories, and behavior form the basis for the following interpretation of the phylogeny of _Ptychohyla_. Additional data are needed to support some of the ideas discussed below; many of the data that are available for _Ptychohyla_ are lacking for other, possibly related, hylids. The family Hylidae is composed of several hundred species, and most of the species are poorly known. Consequently, any attempt to place _Ptychohyla_ in the over-all scheme of hylid phylogeny would be premature at this time. But, as between the five species of two species-groups here recognized as constituting the genus _Ptychohyla_, some estimate of relationships can be made. First, it is necessary to determine the validity of the genus itself.

Ptychohyla as a Natural Assemblage

As stated in the diagnosis of the genus, the only character that sets this group of species apart from other hylids is the presence of ventrolateral glands in the breeding males. To many systematists the thought of being able to identify to genus only breeding males is sufficiently disturbing to cause them to view with disfavor the recognition of the genus. Nonetheless, the question is raised: Do the five species herein placed in the genus _Ptychohyla_ constitute a natural assemblage? If the genus is considered to be more than a category of convenience, that is to say, a group of related species having a common origin, the primary problem is to determine whether or not the five species form a phylogenetic unit.

The species of _Ptychohyla_ are divided into two groups on the basis of external morphology, breeding calls, and tadpoles. The _Ptychohyla euthysanota_ group seems to be a natural group composed of three species, all of which are more closely related to one another than to any other hylid. Likewise, the species comprising the _Ptychohyla schmidtorum_ group seem to represent a natural unit. If the presence of ventrolateral glands in breeding males is ignored, a student of salientian systematics might derive the _Ptychohyla euthysanota_ group from a hylid stock containing _Hyla miotympanum_ and _Hyla mixomaculata_. Likewise, _Ptychohyla schmidtorum_ could be placed with _Hyla uranochroa_ and related species in Costa Rica. Nonetheless, the fact remains that all of the species assigned to the genus _Ptychohyla_ have ventrolateral glands in the breeding males; furthermore, ventrolateral glands are unknown in other hylids. If the _P. schmidtorum_ group and the _P. euthysanota_ group each evolved from separate hylid stocks, then the ventrolateral glands must have developed independently in both groups. That ventrolateral glands developed independently in five species of frogs in southern Mexico and northern Central America and not in any of the other approximately 500 species of hylids in the world is untenable. It is more logical to assume that the development of the glands took place only once in a stock of hylids that gave rise to the five species herein recognized as members of the genus _Ptychohyla_.

Generic Relationships

The affinities of _Ptychohyla_ apparently are not with any of the other groups that have been generically separated from _Hyla_. Of the daughter genera in Middle America only _Plectrohyla_ has stream-adapted tadpoles, but these large frogs are not closely related to _Ptychohyla_. Stuart (1954:169) suggested that certain montane species of _Hyla_ in lower Central America and _Hyla salvadorensis_ in El Salvador may be related to _Ptychohyla_ or even congeneric. I have had experience with most of these species in the field and believe that Stuart was correct in his suggestion of relationships. The species concerned are four red-eyed stream-breeding _Hyla_ in Costa Rica--_H. alleei_, _H. legleri_, _H. rufioculis_, and _H. uranochroa_, plus _Hyla salvadorensis_ in the mountains of El Salvador. Morphologically all of the species are similar; _Hyla uranochroa_, _H. legleri_, and _H. rufioculis_ have a lateral white stripe that is expanded to form a spot beneath the eye, as in _Ptychohyla schmidtorum_. The tadpoles of _Hyla rufioculis_ and _H. uranochroa_ have large funnel-shaped mouths and long slender tails like those of _Ptychohyla schmidtorum_. Lips of the tadpoles of _H. legleri_ and _H. salvadorensis_ are folded laterally, in this respect resembling those of the _Ptychohyla euthysanota_ group. I do not know the tadpoles and the breeding call of _Hyla alleei_. The breeding calls of _Hyla rufioculis_ and _H. uranochroa_ consist of high melodious notes; the calls of _H. legleri_ and _H. salvadorensis_ consist of series of short notes that have the general characteristics of the call of _Ptychohyla schmidtorum_. Affinities of the genus _Ptychohyla_ seem to me to be with the red-eyed species forming the _Hyla uranochroa_ group in Costa Rica. All of the species in the _Hyla uranochroa_ group have large frontoparietal fontanelles, rather small ethmoids, and small nasals that are not in contact with one another or with the ethmoid. Some species have a complete quadratojugal-maxillary arch; others do not. Assuming that the parental stock that gave rise both to the _Hyla uranochroa_ group and to _Ptychohyla_ was widespread in Central America at a time of cooler, more humid conditions, it is possible that with subsequent warming temperatures and seasonal rainfall in the lowlands the parental stock was restricted to the Costa Rican highlands, where the _Hyla uranochroa_ group developed, and to the Chiapas-Guatemala highlands, where _Ptychohyla_ evolved.

Interspecific Relationships

_Ptychohyla schmidtorum_ is thought to resemble more closely the parental stock of the genus than does any other species of _Ptychohyla_ now extant. This parental stock is discussed above in the account of the generic relationships. _Ptychohyla schmidtorum_ has a red eye, white lateral stripe, frontoparietal fontanelle, funnel-shaped mouth in tadpoles, and lacks nuptial spines; in all of these characters it resembles members of the _Hyla uranochroa_ group. Probably during times of glaciation during the Pleistocene, when climates in Mexico and Central America were depressed, the _Ptychohyla_ stock was more widespread than it is now. Subsequent elevation of climatic zones during interglacial periods would have isolated populations as they are today in regions of cloud forests. Thus, through geographic isolation populations could have differentiated and evolved into the present species. Climatic fluctuation in the Pleistocene must have been of sufficient magnitude to permit the spread of cool, moist forests containing _Ptychohyla_ across the Isthmus of Tehuantepec into the mountains of Oaxaca.

Because of its small nuptial spines, small triangular vomers, coloration, and absence of a rostral keel, _Ptychohyla euthysanota_, more than any of the other species in the _P. euthysanota_ group, resembles _P. schmidtorum_. At the present time _P. euthysanota_ and _P. schmidtorum_ are sympatric.

As I have mentioned previously, ecological segregation and interspecific competition probably is highly developed in the tadpoles of _Ptychohyla_. If this ecological segregation resulted from intraspecific competition in a stock of _Ptychohyla_, possibly _P. euthysanota_ and _P. schmidtorum_ differentiated sympatrically in this way. Specific identity is maintained, at least in part, by different breeding calls in males.

_Ptychohyla spinipollex_ and _P. leonhardschultzei_ seem to be more closely related to one another than either is to _P. euthysanota_. Probably a stock of _P. euthysanota_ was isolated on the Atlantic slopes of northern Central America from _P. euthysanota_ on the southern slopes. The frogs on the Atlantic slopes differentiated and spread into the mountains of Oaxaca, where through isolation by the barrier of the Isthmus of Tehuantepec they developed into _P. leonhardschultzei_, while the stock on the northern slopes of Central America evolved into _P. spinipollex_. Subsequent to the differentiation of _P. leonhardschultzei_ and _P. spinipollex_ from _P. euthysanota_ and during a time of cooler more equable climate than exists now, _P. euthysanota_ and _P. schmidtorum_ invaded the Central Highlands of Chiapas. Subsequent climatic changes isolated populations of each in the Central Highlands, where _P. euthysanota macrotympanum_ and _P. schmidtorum chamulae_ evolved. _Ptychohyla ignicolor_ probably represents stock of _P. schmidtorum_ that crossed the Isthmus of Tehuantepec and became isolated in Oaxaca on the western side of the isthmus.

LITERATURE CITED

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1961. Descriptions of two species of frogs, genus Ptychohyla. Studies of American Hylid Frogs, V. Univ. Kansas Publ. Mus. Nat. Hist., 13:349-357, pl. 25, April 27.

FOUQUETTE, M. J. 1960. Isolating mechanisms in three sympatric treefrogs in the Canal Zone. Evolution, 14:484-497, December.

KELLOGG, R. 1928. An apparently new _Hyla_ from El Salvador. Proc. Biol. Soc. Washington, 41:123-124, June 29.

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STUART, L. C. 1954. Descriptions of some new amphibians and reptiles from Guatemala. Proc. Biol. Soc. Washington, 67:159-178, August 5.

TANNER, W. W. 1957. Notes on a collection of amphibians and reptiles from southern Mexico, with a description of a new _Hyla_. Great Basin Nat., 17:52-56, July 31.

TAYLOR, E. H. 1937. New species of hylid frogs from Mexico with comments on the rare _Hyla bistincta_ Cope. Proc. Biol. Soc. Washington, 50:43-54, pls. 2-3, April 21.

1942. New tailless amphibia from Mexico. Univ. Kansas Sci. Bull., 28: 67-89, May 15.

1944. A new genus and species of Mexican frogs. Univ. Kansas Sci. Bull., 30:41-45, June 12.

1949. New or unusual Mexican amphibians. Amer. Mus. Novitates, 1437:1-21, December 7.

_Transmitted December 27, 1962._

UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY

Institutional libraries interested in publications exchange may obtain this series by addressing the Exchange Librarian, University of Kansas Library, Lawrence, Kansas. Copies for individuals, persons working in a particular field of study, may be obtained by addressing instead the Museum of Natural History, University of Kansas, Lawrence, Kansas. There is no provision for sale of this series by the University Library, which meets institutional requests, or by the Museum of Natural History, which meets the requests of individuals. Nevertheless, when individuals request copies from the Museum, 25 cents should be included, for each separate number that is 100 pages or more in length, for the purpose of defraying the costs of wrapping and mailing.

* An asterisk designates those numbers of which the Museum's supply (not the Library's supply) is exhausted. Numbers published to date, in this series, are as follows:

Vol. 1. Nos. 1-26 and index. Pp. 1-638, 1946-1950.

*Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest. Pp. 1-444, 140 figures in text. April 9, 1948.

Vol. 3. *1. The avifauna of Micronesia, its origin, evolution, and distribution. By Rollin H. Baker. Pp. 1-359, 16 figures in text. June 12, 1951.

*2. A quantitative study of the nocturnal migration of birds. By George H. Lowery, Jr. Pp. 361-472, 47 figures in text. June 29, 1951.

3. Phylogeny of the waxwings and allied birds. By M. Dale Arvey. Pp. 473-530, 49 figures in text, 13 tables. October 10, 1951.

*4. Birds from the state of Veracruz, Mexico. By George H. Lowery, Jr., and Walter W. Dalquest. Pp. 531-649, 7 figures in text, 2 tables. October 10, 1951.

Index. Pp. 651-681.

*Vol. 4. (Complete) American weasels. By E. Raymond Hall. Pp. 1-466, 41 plates, 31 figures in text. December 27, 1951.

Vol. 5. Nos. 1-37 and index. Pp. 1-676, 1951-1953.

*Vol. 6. (Complete) Mammals of Utah, _taxonomy and distribution_. By Stephen D. Durrant. Pp. 1-549, 91 figures in text, 30 tables. August 10, 1952.

Vol. 7. Nos. 1-15 and index. Pp. 1-651, 1952-1955.

Vol. 8. Nos. 1-10 and index. Pp. 1-675, 1954-1956.

Vol. 9. *1. Speciation of the wandering shrew. By James S. Findley. Pp. 1-68, 18 figures in text. December 10, 1955.

2. Additional records and extension of ranges of mammals from Utah. By Stephen D. Durrant, M. Raymond Lee, and Richard M. Hansen. Pp. 69-80. December 10, 1955.

3. A new long-eared myotis (Myotis evotis) from northeastern Mexico. By Rollin H. Baker and Howard J. Stains. Pp. 81-84. December 10, 1955.

4. Subspeciation in the meadow mouse, Microtus pennsylvanicus, in Wyoming. By Sydney Anderson. Pp. 85-104, 2 figures in text. May 10, 1956.

5. The condylarth genus Ellipsodon. By Robert W. Wilson. Pp. 105-116, 6 figures in text. May 19, 1956.

6. Additional remains of the multituberculate genus Eucosmodon. By Robert W. Wilson. Pp. 117-123, 10 figures in text. May 19, 1956.

7. Mammals of Coahulia, Mexico. By Rollin H. Baker. Pp. 125-335, 75 figures in text. June 15, 1956.

8. Comments on the taxonomic status of Apodemus peninsulae, with description of a new subspecies from North China. By J. Knox Jones, Jr. Pp. 337-346, 1 figure in text, 1 table. August 15, 1956.

9. Extensions of known ranges of Mexican bats. By Sydney Anderson. Pp. 347-351. August 15, 1956.

10. A new bat (Genus Leptonycteris) from Coahulia. By Howard J. Stains. Pp. 353-356. January 21, 1957.

11. A new species of pocket gopher (Genus Pappogeomys) from Jalisco, Mexico. By Robert J. Russell. Pp. 357-361. January 21, 1957.

12. Geographic variation in the pocket gopher, Thomomys bottae, in Colorado. By Phillip M. Youngman. Pp. 363-387, 7 figures in text. February 21, 1958.

13. New bog lemming (genus Synaptomys) from Nebraska. By J. Knox Jones, Jr. Pp. 385-388. May 12, 1958.

14. Pleistocene bats from San Josecito Cave, Nuevo Leon, Mexico. By J. Knox Jones, Jr. Pp. 389-396. December 19, 1958.

15. New subspecies of the rodent Baiomys from Central America. By Robert L. Packard. Pp. 397-404. December 19, 1958.

16. Mammals of the Grand Mesa, Colorado. By Sydney Anderson. Pp. 405-414, 1 figure in text, May 20, 1959.

17. Distribution, variation, and relationships of the montane vole, Microtus montanus. By Sydney Anderson. Pp. 415-511, 12 figures in text, 2 tables. August 1, 1959.

18. Conspecificity of two pocket mice, Perognathus goldmani and P. artus. By E. Raymond Hall and Marilyn Bailey Ogilvie. Pp. 513-518, 1 map. January 14, 1960.

19. Records of harvest mice, Reithrodontomys, from Central America, with description of a new subspecies from Nicaragua. By Sydney Anderson and J. Knox Jones, Jr. Pp. 519-529. January 14, 1960.

20. Small carnivores from San Josecito Cave (Pleistocene), Nuevo Leon, Mexico. By E. Raymond Hall. Pp. 531-538, 1 figure in text. January 14, 1960.

21. Pleistocene pocket gophers from San Josecito Cave, Nuevo Leon, Mexico. By Robert J. Russell. Pp. 539-548,1 figure in text. January 14, 1960.

22. Review of the insectivores of Korea. By J. Knox Jones, Jr., and David H. Johnson. Pp. 549-578. February 23, 1960.

23. Speciation and evolution of the pygmy mice, genus Baimoys. By Robert L. Packard. Pp. 579-670, 4 plates, 12 figures in text. June 16, 1960.

Index. Pp. 671-690

Vol. 10. 1. Studies of birds killed in nocturnal migration. By Harrison B. Tordoff and Robert M. Mengel. Pp. 1-44, 6 figures in text, 2 tables. September 12, 1956.

2. Comparative breeding behavior of Ammospiza caudacuta and A. maritima. By Glen E. Woolfenden. Pp. 45-75, 6 plates, 1 figure. December 20, 1956.

3. The forest habitat of the University of Kansas Natural History Reservation. By Henry S. Fitch and Ronald R. McGregor. Pp. 77-127, 2 plates, 7 figures in text, 4 tables. December 31, 1956.

4. Aspects of reproduction and development in the prairie vole (Microtus ochrogaster). By Henry S. Fitch. Pp. 129-161, 8 figures in text, 4 tables. December 19, 1957.

5. Birds found on the Arctic slope of northern Alaska. By James W. Bee. Pp. 163-211, plates 9-10, 1 figure in text. March 12, 1958.

*6. The wood rats of Colorado: distribution and ecology. By Robert B. Finley, Jr. Pp. 213-552, 34 plates, 8 figures in text, 35 tables. November 7, 1958.

7. Home ranges and movements of the eastern cottontail in Kansas. By Donald W. Janes. Pp. 553-572, 4 plates, 3 figures in text. May 4, 1959.

8. Natural history of the salamander, Aneides hardyi. By Richard F. Johnston and Gerhard A. Schad. Pp. 573-585. October 8, 1959.

9. A new subspecies of lizard, Cnemidophorus sacki, from Michoacan, Mexico. By William E. Duellman. Pp. 587-598, 2 figures in text. May 2, 1960.

10. A taxonomic study of the middle American snake, Pituophis deppei. By William E. Duellman. Pp. 599-610, 1 plate, 1 figure in text. May 2, 1960.

Index. Pp. 611-626.

Vol. 11. Nos. 1-10 and index. Pp. 1-703, 1958-1960.

Vol. 12. 1. Functional morphology of three bats: Sumops, Myotis, Macrotus. By Terry A. Vaughan. Pp. 1-153, 4 plates, 24 figures in text. July 8, 1959.

*2. The ancestry of modern Amphibia: a review of the evidence. By Theodore H. Eaton, Jr. Pp. 155-180, 10 figures in text. July 10, 1959.

3. The baculum in microtine rodents. By Sydney Anderson. Pp. 181-216, 49 figures in text. February 19, 1960.

*4. A new order of fishlike Amphibia from the Pennsylvanian of Kansas. By Theodore H. Eaton, Jr., and Peggy Lou Stewart. Pp. 217-240, 12 figures in text. May 2, 1960.

5. Natural history of the bell vireo. By Jon C. Barlow. Pp. 241-296, 6 figures in text. March 7, 1962.

6. Two new pelycosaurs from the lower Permian of Oklahoma. By Richard C. Fox. Pp. 297-307, 6 figures in text. May 21, 1962.

7. Vertebrates from the barrier island of Tamaulipas, Mexico. By Robert K. Selander, Richard F. Johnston, B. J. Wilks, and Gerald G. Raun. Pp. 309-345, pls. 5-8. June 18, 1962.

8. Teeth of Edestid sharks. By Theodore H. Eaton, Jr. Pp. 347-362, 10 figures in text. October 1, 1962.

More numbers will appear in volume 12.

Vol. 13. 1. Five natural hybrid combinations in minnows (Cyprinidae). By Frank B. Cross and W. L. Minckley. Pp. 1-18. June 1, 1960.

2. A distributional study of the amphibians of the Isthmus of Tehuantepec, Mexico. By William E. Duellman. Pp. 19-72, pls. 1-8, 3 figures in text. August 16, 1960.

3. A new subspecies of the slider turtle (Pseudemys scripta) from Coahulia, Mexico. By John M. Legler. Pp. 73-84, pls. 9-12, 3 figures in text. August 16, 1960.

4. Autecology of the copperhead. By Henry S. Fitch. Pp. 85-288, pls. 13-20, 26 figures in text. November 30, 1960.

5. Occurrence of the garter snake, Thamnophis sirtalis, in the Great Plains and Rocky Mountains. By Henry S. Fitch and T. Paul Maslin. Pp. 289-308, 4 figures in text. February 10, 1961.

6. Fishes of the Wakarusa river in Kansas. By James E. Deacon and Artie L. Metcalf. Pp. 309-322, 1 figure in text. February 10, 1961.

7. Geographic variation in the North American cyprinid fish, Hybopsis gracilis. By Leonard J. Olund and Frank B. Cross. Pp. 323-348, pls. 21-24, 2 figures in text. February 10, 1961.

8. Descriptions of two species of frogs, genus Ptychohyla; studies of American hylid frogs, V. By William E. Duellman. Pp. 349-357, pl. 25, 2 figures in text. April 27, 1961.

9. Fish populations, following a drought, in the Neosho and Marais des Cygnes rivers of Kansas. By James Everett Deacon. Pp. 359-427, pls. 26-30, 3 figs. August 11, 1961.

10. Recent soft-shelled turtles of North America (family Trionychidae). By Robert G. Webb. Pp. 429-611, pls. 31-54, 24 figures in text, February 16, 1962.

Index. Pp. 613-624.

Vol. 14. 1. Neotropical bats from western Mexico. By Sydney Anderson. Pp. 1-8. October 24, 1960.

2. Geographic variation in the harvest mouse. Reithrodontomys megalotis, on the central Great Plains and in adjacent regions. By J. Knox Jones, Jr., and B. Mursaloglu. Pp. 9-27, 1 figure in text. July 24, 1961.

3. Mammals of Mesa Verde National Park, Colorado. By Sydney Anderson. Pp. 29-67, pls. 1 and 2, 3 figures in text. July 24, 1961.

4. A new subspecies of the black myotis (bat) from eastern Mexico. By E. Raymond Hall and Ticul Alvarez. Pp. 69-72, 1 figure in text. December 29, 1961.

5. North American yellow bats, "Dasypterus," and a list of the named kinds of the genus Lasiurus Gray. By E. Raymond Hall and J. Knox Jones, Jr. Pp. 73-98, 4 figures in text. December 29, 1961.

6. Natural history of the brush mouse (Peromyscus boylii) in Kansas with description of a new subspecies. By Charles A. Long. Pp. 99-111, 1 figure in text. December 29, 1961.

7. Taxonomic status of some mice of the Peromyscus boylii group in eastern Mexico, with description of a new subspecies. By Ticul Alvarez. Pp. 113-120, 1 figure in text. December 29, 1961.

8. A new subspecies of ground squirrel (Spermophilus spilosoma) from Tamaulipas, Mexico. By Ticul Alvarez. Pp. 121-124. March 7, 1962.

9. Taxonomic status of the free-tailed bat, Tadarida yucatanica Miller. By J. Knox Jones, Jr., and Ticul Alvarez. Pp. 125-133, 1 figure in text. March 7, 1962.

10. A new doglike carnivore, genus Cynaretus, from the Clarendonian Pliocene, of Texas. By E. Raymond Hall and Walter W. Dalquest. Pp. 135-138, 2 figures in text. April 30, 1962.

11. A new subspecies of wood rat (Neotoma) from northeastern Mexico. By Ticul Alvarez. Pp. 139-143. April 30, 1962.

12. Noteworthy mammals from Sinaloa, Mexico. By J. Knox Jones, Jr., Ticul Alvarez, and M. Raymond Lee. Pp. 145-159, 1 figure in text. May 18, 1962.

13. A new bat (Myotis) from Mexico. By E. Raymond Hall. Pp. 161-164, 1 figure in text. May 21, 1962.

14. The mammals of Veracruz. By E. Raymond Hall and Walter W. Dalquest. Pp. 165-362, 2 figures. May 20, 1963.

15. The recent mammals of Tamaulipas, Mexico. By Ticul Alvarez. Pp. 363-473, 5 figures in text. May 20, 1963.

More numbers will appear in volume 14.

Vol. 15. 1. The amphibians and reptiles of Michoacan, Mexico. By William E. Duellman. Pp. 1-148, pls. 1-6, 11 figures in text. December 20, 1961.

2. Some reptiles and amphibians from Korea. By Robert G. Webb, J. Knox Jones, Jr., and George W. Byers. Pp. 149-173. January 31, 1962.

3. A new species of frog (Genus Tomodactylus) from western Mexico. By Robert G. Webb. Pp. 175-181, 1 figure in text. March 7, 1962.

4. Type specimens of amphibians and reptiles in the Museum of Natural History, the University of Kansas. By William E. Duellman and Barbara Berg. Pp. 183-204. October 26, 1962.

5. Amphibians and Reptiles of the Rainforests of Southern El Peten, Guatemala. By William E. Duellman. Pp. 205-249, pls. 7-10, 6 figures in text. October 4, 1963.

6. A revision of snakes of the genus Conophis (Family Colubridae, from Middle America). By John Wellman. Pp. 251-295, 9 figures in text. October 4, 1963.

7. A review of the Middle American tree frogs of the genus Ptychohyla. By William E. Duellman. Pp. 297-349, pls. 11-18, 7 figures in text. October 18, 1963.

More numbers will appear in volume 15.

* * * * * * *

Transcriber's Notes.

This file was derived from scanned images. With the exception of the list of typographical errors that were corrected below, the original text is presented.

In the original, the Plates were grouped together between pages 328 and 329. Here the Illustration: block which contains the text associated with the Plates were moved just above the text in their respective Systematic Account listing. The Plate text contain the notation "x 2" after the caption to let the reader know that the image was enlarged by a factor of two.

Typographical Errors Corrected:

Several minor typographical corrections were made (missing periods, commas, misspelling of 'and', etc.); but are not indicated here. More substantial changes are listed below:

References to the Plate 11 (Audiograms): Pl. 1A, Pl. 1B, etc.=> Pl. 11A, Pl. 11B, etc. Page 301, Paragraph 1: know => known Page 302, Paragraph 1: Zoolegy => Zoology Page 303, Paragraph 5: speces => species Page 305, Paragraph 1: excresences => excrescences Page 308, Paragraph 6: xiphisterum => xiphisternum Page 316, Paragraph 3: with => width Page 327, Paragraph 1: leonhard-schultzei => leonhardschultzei to match remaining report text Page 331, Paragraph 1: skelton => skeleton Publication List Vol. 13, No. 8.: Decriptions => Descriptions