A Review of the Middle American Tree Frogs of the Genus Ptychohyla Part 3

_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. Was.h.i.+ngton, 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 a.n.u.s.

_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 p.r.o.nounced; 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. a.n.a.l opening at the level of the upper edge of thighs; a.n.a.l flap short; a.n.a.l 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 a.n.a.l 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 a.n.a.l 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; a.n.a.l 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 h.o.r.n.y 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.

[Ill.u.s.tration: PLATE 13 _Ptychohyla euthysanota macrotympanum_ (KU 58049). 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 a.n.a.l 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 p.r.o.nounced; 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. a.n.a.l opening near upper edge of thighs; short a.n.a.l flap bordered above by thin dermal fold; small tubercles and heavy dermal fold lateral to a.n.a.l 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 a.n.a.l 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 a.n.a.lysis 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 a.n.u.s 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; a.n.a.l 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 h.o.r.n.y 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.