The _Ptychohyla euthysanota_ Group

Three species in group; adults having moderate amount of webbing

between fingers, and tarsal fold; breeding males having spinous,

horny, nuptial tuberosities on pollex; mouths of tadpoles having

lateral folds in lips and 4/6 or 4/7 tooth-rows; breeding call

consisting of one long note.

=Ptychohyla euthysanota=

_Diagnosis._--Rostral keel absent; nuptial spines in males small;

interorbital region much wider than eyelid.

[Illustration: PLATE 12

_Ptychohyla euthysanota euthysanota_ (KU 58008). x 2.]

=Ptychohyla euthysanota euthysanota= (Kellogg)

_Hyla euthysanota_ Kellogg, Proc. Biol. Soc. Washington,

41:123-124, June 29, 1928 [Holotype.--USNM 73296 from Los

Esemiles, Depto. Chalatenango, El Salvador; Ruben A. Stirton

collector]. Mertens, Senckenbergiana, 33:169-171, June 15,

1952; Abhand. Senckenbergische Naturf. Gesell., 487:29,

December 1, 1952. Stuart, Proc. Biol. Soc. Washington, 67:169,

August 5, 1954.

_Hyla rozellae_ Taylor, Univ. Kansas Sci. Bull., 28:78-80, pl. 9,

fig. 1, May 15, 1942 [Holotype.--USNM 115039 from Salto de

Agua, Chiapas, Mexico; Hobart M. and Rozella Smith

collectors]. Taylor and Smith, Proc. U. S. Natl. Mus., 95:587,

June 30, 1945. Smith and Taylor, Bull. U. S. Natl. Mus.,

194:86, June 17, 1948. Stuart, Proc. Biol. Soc. Washington,

67:169, August 5, 1954.

_Ptychohyla bogerti_ Taylor, Amer. Mus. Novitates, 1437:13-16,

fig. 5, December 7, 1949 [Holotype.--AMNH 51847 from Rio

Grande, Oaxaca, Mexico; Thomas MacDougall collector]. Stuart,

Proc. Biol. Soc. Washington, 67:169, August 5, 1954.

_Ptychohyla euthysanota_, Duellman, Univ. Kansas Publ. Mus. Nat.

Hist., 13:351, April 27, 1961.

_Diagnosis._--Dorsum tan to reddish brown; venter white; rarely

flecked with brown or black; a white stripe on upper lip, on flank,

and usually above anus.

_Description._--The following description is based on KU 58008 from

Finca La Paz, Depto. San Marcos, Guatemala (Pl. 12). Adult male having

a snout-vent length of 35.0 mm.; tibia length, 16.5 mm.; tibia

length/snout-vent length, 47.1 per cent; foot length, 14.2 mm.; head

length, 11.0 mm.; head length/snout-vent length, 31.4 per cent; head

width, 10.7 mm.; head width/snout-vent length, 30.6 per cent; diameter

of eye, 3.3 mm.; diameter of tympanum, 1.8 mm.; tympanum/eye, 54.5 per

cent. Snout in lateral profile nearly square, slightly rounded above,

and in dorsal profile bluntly rounded; canthus pronounced; loreal

region moderately concave; lips thick, rounded, and slightly flaring;

nostrils protuberant; internarial distance, 3.0 mm.; top of head flat;

interorbital distance, 4.1 mm., and approximately a third broader than

width of eyelid, 2.9 mm. Moderately heavy dermal fold from posterior

corner of eye above tympanum to point above insertion of forelimb,

covering upper edge of tympanum; tympanum round, its diameter slightly

more than its distance from eye. Forearm moderately robust, having

distinct dermal fold on wrist; dermal fold, but no row of tubercles

along ventrolateral surface of forearm; pollex only slightly enlarged,

bearing triangular shaped patch of small horn-covered spines (128 on

right, 134 on left); second and fourth fingers equal in length;

subarticular tubercles round, distal one on fourth finger bifid; discs

moderate in size, that of third finger equal to 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; low rounded tarsal fold; inner

metatarsal tubercle large, elliptical, and flat; outer metatarsal

tubercle small and round; low dermal fold from heel to disc of fifth

toe; 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 proximal end of penultimate phalanx; thin dermal fold

from inner metatarsal tubercle to disc of first toe; disc smaller than

on fingers. Anal opening at the level of the upper edge of thighs;

anal flap short; anal opening bordered above by thin transverse dermal

fold and laterally by heavy dermal fold. 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, not reaching axilla or groin and broadly separated

midventrally. Tongue ovoid, emarginate, and only slightly free

posteriorly; vomerine teeth 2-2, situated on small triangular

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 dull reddish brown with

irregular dark brown reticulations on head and body and dark brown

transverse bands on limbs; dorsal surfaces of first and second fingers

and webbing on hand cream color; dorsal surfaces of third and fourth

fingers dull brown; anterior surfaces of thighs dull creamy yellow;

posterior surfaces of thighs dull brown; tarsi and toes tan with brown

flecks; webbing of feet brown; faint creamy white stripe along lateral

edges of tarsi and forearms; thin white line along edge of upper lip;

distinct white stripe above and beside anal opening; axilla white;

throat, chest, belly, and ventral surfaces of forelimbs creamy white;

flanks white, separated from pale venter by a row of partly connected

dark brown spots; ventral surfaces of thighs dull creamy yellow; feet

grayish brown; ventrolateral glands pale grayish brown; small brown

flecks on periphery of chin.

In life the dorsal ground-color was pale reddish brown

(Orange-Cinnamon); dorsal reticulations dark brown (Chocolate); dorsal

surfaces of first and second fingers and webbing on hands creamy tan

(Light Pinkish Cinnamon); posterior surfaces of thighs reddish brown

(Vinaceous-Tawny); webbing of feet gray (Deep Mouse Gray); throat and

belly grayish white (Pale Gull Gray); ventral surfaces of hind limbs

creamy white (Marguerite Yellow); spots on flanks dark brown (Warm

Sepia); iris reddish bronze (Apricot Orange).

_Variation._--No geographic variation in structural characters is

discernible; variation in size and proportions is given in Table 1. Of

32 adults examined, seven have the tongue shallowly notched

posteriorly; in the others the tongue is emarginate. Twenty specimens

have a bifid subarticular tubercle beneath the fourth finger; in the

others there are no bifid tubercles.

The coloration described above is typical of the 16 specimens

available from Finca La Paz. The living coloration at night, when the

frogs were collected, was somewhat darker than the living colors

described above, which were recorded for the frogs the morning after

collection, at which time one individual had a pale reddish brown

dorsum (Orange-Cinnamon) with dull olive green (Deep Grape Green)

reticulations on the back and transverse bands on the limbs; the

dorsal surfaces of the first and second fingers and the discs on the

third and fourth fingers were orange (Mikado Orange).

More than half of the specimens from Finca La Paz agree in all

essential characters with the description given above. The

distinctness of the white stripe on the upper lip is variable; in two

individuals the stripe is barely discernible. Likewise, in some

individuals the white stripe on the flanks is not distinct, either

because there are few or no brown spots separating the stripe from the

pale venter, or because the ventrolateral gland has diffused the pale

color on the flanks. There is some noticeable variation in dorsal

coloration, either through a greater or lesser development of dark

pigment. One specimen (KU 58007) is grayish tan above with dark brown

markings; the posterior surfaces of the thighs are dull grayish

yellow; the first and second fingers and the webbing on the hands are

pale yellowish gray; the belly and throat are dusty white; the flecks

on the throat are gray; the ventral surfaces of the feet are grayish

brown. Dark individuals, such as KU 58009 have a uniform dark brownish

black dorsum; the belly is cream; the first and second fingers and the

webbing on the hands are dull creamy tan; the dorsal and ventral

surfaces of the feet are dark brown. In KU 58013 there is a heavy

suffusion of brown on the throat and flanks. Two specimens have

scattered white flecks on the dorsum.

The reddish brown dorsal ground-color with dark brown reticulations on

the head and body and dark brown transverse bands on the limbs seems

to be rather constant throughout the range of the subspecies.

Likewise, the presence of the white stripe on the upper lip and the

white stripe around the anal opening are present on most specimens. In

breeding males having well-developed ventrolateral glands the lateral

white stripe often is obliterated.

_Description of Tadpole._--The following description is based on KU

60042 from Finca La Paz, Depto. San Marcos, Guatemala (Figs. 4A and

6A). No limb buds; total length, 35.8 mm.; body length, 11.2 mm.; body

length/total length, 31.3 per cent. Body moderately depressed,

slightly wider than deep, ovoid in dorsal profile; mouth directed

ventrally; eyes small, directed dorsolaterally; nostrils slightly

protuberant and directed anteriorly, closer to eye than snout;

spiracle sinistral and posteroventrad to eye; anal tube dextral.

Caudal fin low, rounded posteriorly; depth of caudal musculature about

one-half greatest 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 six rows of papillae laterally. Beaks

moderately developed, bearing peglike serrations; slender lateral

projections on upper beak; tooth-rows 4/6; upper rows subequal in

length, second longest; fourth row interrupted medially; lower rows

complete; lower rows 1-4 equal in length to upper rows; fifth lower

row somewhat shorter; sixth lower row short.

Body brown above; tip of snout cream color; grayish cream color below;

caudal musculature creamy tan; caudal fin transparent; cream-colored

crescent-shaped mark on posterior edge of body and anterior part of

caudal musculature, bordered posteriorly by dark brown blotch;

scattered brown flecks on caudal musculature and posterior part of

caudal fin. Eye bronze color in life.

_Variation._--The variation in size and proportions is given in Table

2. In some specimens the first upper tooth-row is irregular, sometimes

broken, and often shorter than other upper tooth-rows. Usually the

fourth upper and first lower, and sometimes the sixth lower,

tooth-rows are interrupted medially. One specimen has a short,

irregular, seventh lower tooth-row; all others have six.

The cream-colored crescent-shaped mark usually is distinct. The brown

blotch posterior to this mark is variously shaped ranging from a

narrow vertical bar to a triangular blotch. Brown flecks seldom are

present on the anterior part of the ventral caudal fin.

_Comparisons._--Aside from the characters given in the diagnosis, _P.

euthysanota euthysanota_ can be distinguished from both _P.

spinipollex_ and _P. leonhardschultzei_ by the absence of bold black

and white marbling on the flanks; furthermore, from the former it can

be distinguished by having more and smaller horny nuptial spines and

from the latter by having the snout, in lateral profile, rounded above

and not acutely angulate. _Ptychohyla euthysanota euthysanota_ differs

from _P. euthysanota macrotympanum_ by normally having a darker dorsal

color, broader stripe on upper lip, and a distinct lateral stripe.

Occurring sympatrically with _Ptychohyla euthysanota euthysanota_ are

several species of _Plectrohyla_, all of which differ in having a bony

prepollex, rather rugose skin on the dorsum, and more squat bodies.

Other sympatric species are _Ptychohyla schmidtorum schmidtorum_,

which lacks a tarsal fold and nuptial spines and has a red eye in

life, _Hyla salvadorensis_, which has a green dorsum and lacks spinous

nuptial tuberosities, and _Hyla sumichrasti_, a small yellow frog

usually lacking vomerine teeth.

_Life History._--This subspecies breeds in clear, swift mountain

streams. Males call from stems and leaves of plants at the edge of, or

overhanging, the streams. The breeding call consists of a soft

"wraack," repeated at intervals of three to four seconds. Each note

has a duration of 0.60 to 0.65 seconds and has 91 to 102 pulses per

second; the dominant frequency falls between 3000 and 3200 cycles per

second.

Tadpoles in various stages of development were found at Finca La Paz,

Guatemala, in late July. This indicates that there is either extreme

differential growth, or, more probably, an extended breeding season. A

tadpole having a body length of 6.8 mm. and a total length of 19.1 mm.

has a short median first upper tooth-row; lower tooth-rows 3-6 are

only two-thirds as long as lower rows 1 and 2. Two recently

metamorphosed young have snout-vent lengths of 14.2 and 14.8 mm.; they

are colored like the adults.

_Remarks._--The type specimen of _Hyla euthysanota_ Kellogg (1928:123)

is a female; therefore, when Taylor (1944) proposed the name

_Ptychohyla_ for hylids having ventrolateral glands in breeding males,

he was unaware that _Hyla euthysanota_ was a member of this group. In

his description of _Hyla rozellae_, Taylor (1942) did not compare his

specimens with _Hyla euthysanota_, but instead placed _H. rozellae_

with _H. loquax_ and _H. rickardsi_. The type series of _H. rozellae_

consists of one large adult female and several metamorphosing young.

Taylor (1949:16) based the description of _Ptychohyla bogerti_ on two

males and compared these specimens with _P. adipoventris_ Taylor [=

_P. leonhardschultzei_ (Ahl)]. Thus, in a period of 22 years the

females of this species were given two names and the male another.

Stuart (1954:169) suggested that _Hyla euthysanota_ and _Hyla

rozellae_ were _Ptychohyla_. Now that sufficient specimens are

available from throughout the range it is possible to determine that

the various named populations are conspecific.

_Distribution._--This subspecies inhabits cloud forests at elevations

of 660 to 2200 meters on the Pacific slopes of the Sierra Madre from

extreme eastern Oaxaca and western Chiapas, Mexico, through Guatemala

to northern El Salvador; probably it occurs also in southwestern

Honduras. Aside from the specimens listed below, three in the

Frankfurt Museum from Depto. Santa Ana, El Salvador (44571, Hacienda

San Jose; 43040, Hacienda Los Planes; 65119, Miramundo) are listed by

Mertens (1952:29).

_Specimens examined._--MEXICO: _Chiapas: Cascarada, 30 km. W of

Ciltapec_, UMMZ 87851-2; Cerro Ovando, UMMZ 87853-4; Chicomuselo, UMMZ

94439-40; Finca Juarez, 28 km. N of Escuintla, USNM 115052-5; _Las

Nubes, Cerro Ovando_, USNM 115030-8; Salto de Agua, USNM 115039-51.

_Oaxaca_: Cerro Pecho Blanco, UIMNH 40963; between La Gloria and Cerro

Azul, UIMNH 40976-7; Rio Grande, AMNH 51847-8; Santo Tomas Tecpan,

UIMNH 41071.

GUATEMALA: _San Marcos_: Finca La Paz, 2 km. W of La Reforma, KU

58001-14, 59937 (skeleton), 60042-3 (tadpoles), 60044 (4 young), MCZ

34997, UMMZ 107739, 123151-7 (tadpoles); Finca Pirineos, Rio Samala,

CNHM 35066. _Santa Rosa_: Finca La Gloria, UMMZ 123148 (tadpoles),

123150 (tadpoles). _Solola_: Finca Santo Tomas, UMMZ 123149

(tadpoles); Olas de Moca, near Moca, CNHM 20208.

EL SALVADOR: _Chalatenango_: Los Esemiles, USNM 73296. _Santa Ana_:

Miramundo, CNHM 65120.

[Illustration: PLATE 13

_Ptychohyla euthysanota macrotympanum_ (KU 58049). x 2.]

=Ptychohyla euthysanota macrotympanum= (Tanner)

_Hyla macrotympanum_ Tanner, Great Basin Nat., 17:52-53, July 31,

1957 [Holotype.--AMNH 62141 (formerly BYU 13752) from 10 miles

east of Chiapa de Corzo, Chiapas, Mexico; Robert Bohlman

collector].

_Ptychohyla macrotympanum_, Duellman, Univ. Kansas Publ. Mus. Nat.

Hist., 13:351, April 27, 1961.

_Diagnosis._--Dorsum usually pale tan; venter white with scattered

brown or black flecks; a thin white stripe on upper lip and another

above anal opening; no distinct white stripe on flanks.

_Description._--The following description is based on KU 58049 from

Linda Vista, Chiapas, Mexico (Pl. 13). Adult male having a snout-vent

length of 38.0 mm.; tibia length, 19.5 mm.; tibia length/snout-vent

length, 51.3 per cent; foot length, 15.7 mm.; head length, 11.8 mm.;

head length/snout-vent length, 31.1 per cent; head width, 11.7 mm.;

head width/snout-vent length, 30.8 per cent; diameter of eye, 3.8 mm.;

diameter of tympanum, 2.1 mm.; tympanum/eye, 55.2 per cent. Snout in

lateral profile nearly square, slightly rounded above, and in dorsal

profile bluntly rounded; canthus pronounced; loreal region concave;

lips thick, rounded, and slightly flaring; nostrils protuberant;

internarial distance, 3.1 mm.; top of head flat; interorbital

distance, 3.8 mm., and approximately a fourth broader than width of

eyelid, 3.1 mm. A moderately heavy dermal fold from posterior corner

of eye above tympanum to point above insertion of forelimb, covering

upper edge of tympanum; tympanum round, its diameter equal to its

distance from eye. Forearm moderately robust, having a distinct dermal

fold on wrist; dermal fold, but no row of tubercles along

ventrolateral surface of forearm; pollex only slightly enlarged,

bearing triangular patch of small horn covered spines (94 on right,

100 on left); fourth finger slightly longer than second; subarticular

tubercles round, none bifid; discs moderate in size, that of third

finger equal to diameter of tympanum; vestige of 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; weak tarsal fold on distal two-thirds of tarsus; inner

metatarsal tubercle large, elliptical, and flat; outer metatarsal

tubercle small and round; no dermal fold from heel to disc of fifth

toe; subarticular tubercles round; length of digits from shortest to

longest 1-2-5-3-4; third toe webbed to base of disc; fifth toe webbed

to middle of penultimate phalanx; fourth toe webbed to proximal end of

penultimate phalanx; no fold of skin from inner metatarsal tubercle to

base of disc on first toe; discs much smaller than on fingers. Anal

opening near upper edge of thighs; short anal flap bordered above by

thin dermal fold; small tubercles and heavy dermal fold lateral to

anal opening. Skin of dorsum and ventral surfaces of fore limbs and

shanks smooth; that of throat, belly, and ventral surfaces of thighs

granular. Ventrolateral glands weakly developed, not reaching axilla

or groin and broadly separated midventrally. Tongue ovoid, shallowly

notched posteriorly, and barely free behind; vomerine teeth 2-2,

situated on small triangular 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 pinkish tan with the

greatest part of head and body covered by large gray interconnected

blotches; black flecks over most of dorsum; grayish brown transverse

bands on shanks; dorsal surfaces of first and second fingers pale

grayish yellow; dorsal surfaces of third and fourth fingers and

webbing on hand pale grayish tan; anterior surfaces of thighs pale

flesh-color; posterior surfaces of thighs pale grayish yellow; dorsal

surfaces of tarsi and toes pale grayish tan with black flecks; webbing

of feet pale gray; faint creamy white stripes along ventrolateral

edges of tarsi and forearms; a very thin white line along edge of

upper lip; a narrow grayish white stripe above anal opening; axilla

gray; throat, chest, belly, and ventral surfaces of forelimbs pale

grayish white; ventral surfaces of hind limbs cream color; flanks gray

flecked with brown; ventral surfaces of feet grayish tan;

ventrolateral glands pinkish tan; anterior one-half of chin flecked

with brown.

In life the dorsum was pale tan (Pinkish Buff); the dark markings on

dorsum dull brown (Tawny-Olive); tarsi pale tan (Pale Pinkish Buff);

flanks pinkish tan (Pale Cinnamon-Pink); iris coppery bronze (Capucine

Orange).

_Variation._--The few specimens from a limited geographic region

preclude any analysis of geographic variation. All specimens, except

the one described above, have the fifth toe webbed to the base of the

disc. Many individuals have a bifid subarticular tubercle beneath the

fourth finger. The shape of the posterior edge of the tongue varies

from nearly straight and shallowly notched to bluntly rounded and

emarginate. Two females (KU 58050-1) have more pointed snouts in

dorsal profile than do males.

Some specimens, such as KU 58048, are notably darker than the specimen

described above; in dark specimens the dorsum is brown with dark brown

markings; all fingers and the webbing on the hand are brown. The tarsi

and feet are like those in the specimen described above, but the

posterior surfaces of the thighs are yellowish tan heavily suffused

with brown; the venter is cream color. In life KU 58048 had a brown

(Verona Brown) dorsum with dark brown (Chocolate) markings. KU 58047

is slightly darker than KU 58048 and has scattered small white flecks

on the dorsum. All specimens have the thin white line on the upper

lip, but in some individuals it is indistinct. The grayish white line

above the anus is present in all specimens.

_Description of Tadpole._--The following description is based on KU

60049 from Rio Hondo, 9.5 kilometers south of Pueblo Nuevo

Solistahuacan, Chiapas, Mexico (Figs. 4B and 6B). No limb buds; total

length, 36.2 mm.; body length, 11.1 mm.; body length/total length,

30.6 per cent. Body moderately depressed, slightly wider than deep,

ovoid in dorsal profile; mouth directed ventrally; eyes small,

directed dorsolaterally; nostrils slightly protuberant and directed

anteriorly, somewhat closer to eye than snout; spiracle sinistral and

posteroventrad to eye; anal tube dextral. Caudal fin low, acutely

rounded posteriorly; depth of caudal musculature slightly more than

one-half greatest depth of caudal fin; caudal musculature extending

nearly to tip of tail.

Mouth large; lips having deep lateral folds; two complete rows of

papillae on lips; five or six rows of papillae laterally. Beaks

moderately developed, bearing small 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 subequal to upper rows; fifth and sixth lower rows decreasing

in length; first lower row interrupted medially.

Body brown above; tip of snout cream color; venter creamy white;

caudal musculature creamy tan; caudal fin transparent; cream-colored

crescent-shaped mark on posterior edge of body and anterior part of

caudal musculature, bordered posterodorsally by dark brown blotch;

four dark brown blotches on dorsum of anterior part of caudal

musculature; scattered brown flecks on caudal musculature and fin; eye

silvery bronze in life.

_Variation._--The variation in size and proportions is given in Table

2. All specimens have 4/6 tooth-rows; in some the first lower row is

interrupted medially. Specimens from Jacaltenango and two kilometers

west of San Pedro Necta, Depto. Huehuetenango, Guatemala, have weakly

developed sixth lower tooth-rows.

The cream-colored crescent-shaped mark is distinct in all specimens;

the dorsal blotches on the anterior part of the caudal musculature are

narrow and do not extend far onto the sides of the tail. The blotches

are most distinct in small tadpoles and sometimes indistinct in large

ones.

_Comparisons._--_Ptychohyla euthysanota macrotympanum_ can be

distinguished from both _P. spinipollex_ and _P. leonhardschultzei_ by

the absence of bold black and white marbling on the flanks, as well as

by the characters given in the diagnosis; furthermore, from the former

it differs in having more and smaller horny nuptial tuberosities and

from the latter by having the snout, in lateral profile, rounded above

instead of angulate. _Ptychohyla euthysanota macrotympanum_ differs

from _P. e. euthysanota_ by normally having a paler dorsum, narrower

stripe on upper lip, and no distinct lateral stripe.

_Ptychohyla euthysanota macrotympanum_ occurs sympatrically with

_Plectrohyla guatemalensis_ and _P. matudai matudai_. Each of the last

two has a bony prepollex, rather rugose skin on the dorsum, and more

squat body. Other sympatric species are _Hyla walkeri_, which has a

green dorsum with brown markings and a rather pointed snout, and _Hyla

sumichrasti_, a small yellow frog usually lacking vomerine teeth.

_Life History._--This species breeds in clear mountain streams in

mixed pine and broad-leafed forest. Males call from trees and bushes

along the streams. The breeding call consists of a soft "wraack,"

repeated three to nine times with intervals of 2.7 to 3.4 seconds

between notes. Each note has a duration of 0.60 to 0.65 second, and a

rate of 92 to 100 pulses per second; the dominant frequency falls

between 3000 and 3200 cycles per second (Pl. 11B). The call is

indistinguishable from that of _P. e. euthysanota_.

Tadpoles in various stages of development were found in the Rio Hondo,

Chiapas, on June 16, 1960. The smallest tadpole has a total length of

24.1 mm.; in this individual the sixth lower tooth row has barely

started to develop. A metamorphosing frog taken at the same time has a

snout-vent length of 19.8 mm., a short remnant of the tail, and the

mouth and tongue developed, whereas another individual having a

snout-vent length of 17.8 mm. and a tail 31.0 mm. in length still has

larval teeth. Three completely metamorphosed juveniles collected by L.

C. Stuart at Jacaltenango, Guatemala, on June 6 and 7, 1955, have

snout-vent lengths of 16.0, 16.0, and 16.1 mm.

_Remarks._--Tanner (1957:52) based the description of _Hyla

macrotympanum_ on a single female, which, of course, lacked the

characters diagnostic of _Ptychohyla_. On the basis of general

external characters Tanner suggested that _Hyla macrotympanum_ was

related to _H. miotympanum_, from which it differed in having a larger

tympanum and a bifid subarticular tubercle beneath the fourth finger.

The collection of additional females, together with males of the

species, has shown that _Hyla macrotympanum_ is a _Ptychohyla_.

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).

[Illustration: PLATE 14

_Ptychohyla leonhardschultzei_ (KU 64117). x 2.]

=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).

[Illustration: PLATE 15

_Ptychohyla spinipollex_ (KU 58054). x 2.]

=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.

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