Large congregations were breeding at Tehuantepec on July 5, at

Tapanatepec on July 13, and at Hueyapan on July 24, 1956. The frogs

were breeding in open ponds in scrub forest and savanna; none was found

in the rainforest. Males call while floating on the water (Pl. 7, fig.

1); the call is a soft "do-ing, do-ing" with a rising tone on the last

note. Numerous individual egg masses were along the bank of a pond near

Tehuantepec; one large composite egg mass there had a surface area of

about one square meter (Pl. 7, fig. 2). The large series from Union

Hidalgo was obtained by digging specimens out of a dry sandy river bank

in the dry season. Some of the individuals were buried to a depth of 25

centimeters.

In life individuals from the Pacific lowlands were dull brown and gray;

those from Acayucan were dark chocolate brown to black with pink or red

blotches, forearms, and dorsal stripe. Not all specimens from the

Atlantic lowlands are so colored; individuals from Cordoba and

Mirador, Veracruz, are like those from Tehuantepec.

=Leptodactylus labialis= Cope

_Oaxaca_: Agua Caliente; Chivela (2); Matias Romero (12); 33

km. N of Matias Romero (4); Mixtequilla; Santa Efigenia;

Tapanatepec; Tehuantepec (38); Tolosita (2); 33 km. W of

Zanatepec (49). _Veracruz_: Acayucan (3); Ciudad Aleman;

Cuatotolapam (10); Hueyapan; La Oaxaquena (4); 38 km. SE of

Jesus Carranza; 20 km. ENE of Jesus Carranza; Novillero (3);

San Lorenzo (2).

Although _Leptodactylus labialis_ does not appear to be so abundant as

_Leptodactylus melanonotus_, the former was found throughout the

lowlands of the isthmus. In the dry season individuals were found along

streams, and in the rainy season breeding congregations were found in

rain pools, marshes, ponds, and even small puddles. The call is a slow

"wort, wort, wort." Males call beneath the water and from beneath rocks

and from holes in the ground. The average snout-vent length of eight

adult males is 37.2 mm. A completely metamorphosed juvenile obtained at

Hueyapan on July 24, 1956, has a snout-vent length of 11 mm.

=Leptodactylus melanonotus= Hallowell

_Oaxaca_: Agua Caliente (25); Cerro Arenal (2); Cerro

Quiengola (3); Cerro San Pedro (3); Chivela (2); Coyol;

Juchitan; Matias Romero (11); Mixtequilla (2); Papaloapan

(2); Salazar (9); Salina Cruz; 11 km. S of Santiago Chivela;

Tapanatepec (17); Tehuantepec (176); Tolosita; Union

Hidalgo; 27 km. W of Zanatepec (6). _Veracruz_: Acayucan;

Cuatotolapam (9); Cosoleacaque; 20 km. ENE of Jesus Carranza

(2); 20 km. SE of Minatitlan (2); Novillero; San Lorenzo

(6).

This frog is abundant throughout the lowlands of the isthmus, where in

the dry season individuals were found along streams and beneath rocks

at a spring seepage. In the rainy season males were calling from nearly

every bit of standing water. The call is a soft clicking sound

resembling that made by striking two small stones together. The average

snout-vent length of ten adult males is 41.8 mm. There is considerable

variation in the extent of the yellowish brown glandular areas on the

belly. Some have none, whereas others have a broad area on the chest, a

band along the flanks, and a thin band across the lower abdomen.

Individuals collected in the dry season vary in the same fashion as do

those collected in the rainy season, at which time they were breeding.

The glands are equally well-developed in adults of both sexes, and were

present in some juveniles with snout-vent lengths of less than 20 mm.

Apparently the development of the glands is not associated with

maturity, sex, or size.

=Diaglena reticulata= Taylor

_Oaxaca_: Cerro Arenal; Chivela; Salina Cruz (26); San

Antonio (3); Tehuantepec (2); 8.6 km. W of Tehuantepec (11);

Zarzamora.

Breeding congregations of this rare frog were found 8.6 kilometers west

of Tehuantepec on July 5, 1956, and at Salina Cruz on July 6, 1958.

Both choruses took place immediately after torrential rains. In both

instances the frogs were in and about open muddy pools in the scrub

forest (Pl. 5, fig. 2); males called from the bank near the water, and

clasping pairs were found only on land (Pl. 8, figs. 1-2). The call is

a loud, nasal "braaa," two to three seconds in duration. Amplexus is

axillary.

The dorsal ground color is light yellowish green tending towards olive

on the head and fading to yellow on the flanks. The ventral surfaces,

including the vocal sac, are white; the iris is golden and flecked with

black. The present series agrees well with the description of

_reticulata_ (based on two specimens) given by Taylor (1942:60). A

detailed analysis of variation, comparison with related species, and

descriptions of tadpoles are reserved for a future report.

=Hyla baudini= Dumeril and Bibron

_Oaxaca_: Bisilana; Cerro Quiengola (2); Cerro San Pedro;

Coyol; Matias Romero (12); Mixtequilla; Rio Sarabia (7);

Salazar; San Antonio; 11 km. S of Santiago Chivela; Santo

Domingo (3); Tapanatepec (2); Tehuantepec (23); Tolosita.

_Veracruz_: Acayucan; Amatitlan; Ciudad Aleman (3);

Cosamaloapan (2); Cuatotolapam (15); 10 km. SE of Hueyapan;

20 km. S of Jesus Carranza; 38 km. S of Jesus Carranza (2);

20 km. ENE of Jesus Carranza (4); La Oaxaquena (2);

Minatitlan (2); Naranja (3); Novillero (9); Rio de las

Playas (2); San Lorenzo (5); Tapalapan (2).

Commonly found on both sides of the isthmus, this large tree frog

nearly always is associated with trees; it is not found in the

savannas, although it breeds in savannas adjacent to rainforest. It

appears to be somewhat more abundant in scrub forest than in

rainforest. In the daytime individuals were found under the outer

sheaths of banana plants, in the axils of leaves of elephant ears

(_Xanthosoma_), in cavities in trees, and on shaded limbs in the

forest. Recently metamorphosed individuals having snout-vent lengths

slightly more than 20 mm. were found in the latter part of July.

=Hyla ebraccata= Cope

_Oaxaca_: Donaji (17); 43 km. N of Matias Romero (27);

Sarabia (6); Tolosita (3); Ubero (17). _Veracruz_: Aquilera.

This small species was found only in forested areas, where calling

males were on bushes and trees around rain pools. The call is a harsh

squawk repeated at intervals of 15 to 20 seconds, followed by a minute

or more of silence, and then repeated. Clasping pairs were found on

bushes and in the water.

The dorsum bears a dark chocolate brown hour glass-shaped mark, which

in some individuals is broken into a large mark posteriorly and a

smaller triangular one on the head and nape. The dorsal ground color

varies from pale cream or ivory to yellow or tan. The intensity of the

dorsal pigmentation is subject to rather rapid change. The flanks,

hands, and anterior part of the venter are lemon yellow; the feet,

thighs, and posterior part of the venter are golden yellow. The dorsal

surface of the shank is yellow to tan with chocolate brown bars or

spots; the heel is pale yellow. There is a dark brown bar in the loreal

region and a dark brown bar extending posteriorly from the eye to a

point above the insertion of the forelimb. The iris is a copper color.

The toes are completely webbed; the fingers, one-third webbed. There is

a small axillary web that is evident when the forelimbs are at right

angles to the body. Twenty males have an average snout-vent length of

28.1 mm.; three females, 35.3 mm. There are no nuptial tuberosities on

the pollex of breeding males.

This species has been collected at Coyame and Catemaco in Los Tuxtlas

and at various localities in Tabasco; it apparently ranges eastward

from southern Veracruz, Mexico, in humid forests to El Peten,

Guatemala.

=Hyla loquax= Gaige and Stuart

_Oaxaca_: Donaji (7); 43 km. N of Matias Romero (21).

_Veracruz_: 19 km. N of Acayucan (4); Aquilera (3); 8 km. SW

of Coatzacoalcos (36); Cuototolapam (11); Naranja (13); San

Lorenzo (8).

In the isthmus this species is known only from the humid forests of the

Gulf lowlands; it is also known from Boca del Rio, Veracruz, and from

Teapa and Villa Hermosa, Tabasco.

Calling males were found on aquatic plants above the water in deep

ponds in the forest where it was necessary for the collector to wade

waist-deep in water to obtain them. The call is a loud "hah-onk."

Individuals, when active at night, are yellowish tan above with light

olive green spots. The flanks, belly, and vocal sac are yellow, and the

anterior and posterior surfaces of the thighs and webbing of the feet

are bright orange-red or tomato red. Individuals found during the day

are grayish brown with olive markings or reddish brown with black

markings. Sleeping individuals are ivory-gray with faint gray markings.

The iris is a bright copper color. Fifteen adult males have an average

snout-vent length of 41.7 mm.; they have no horny nuptial pads on the

pollex.

The relationships of this species are with _Hyla rickardsi_ Taylor, a

species known only from the foothills of the Sierra Madre Oriental in

the states of Puebla and Veracruz. The distinguishing characteristics

of these species are given in Table 1. Living individuals may be

distinguished immediately by the flash colors on the thighs--red in

_loquax_ and yellow in _rickardsi_. The calls of the two species are

distinctly different; that of _rickardsi_ is a high-pitched, loud

rattle continued for several seconds, notably different from the

goose-like honk of _loquax_.

TABLE 1.--COMPARISON OF CERTAIN CHARACTERS IN HYLA LOQUAX

AND HYLA RICKARDSI

=================================+================+====================

CHARACTER | _loquax_ | _rickardsi_

---------------------------------+----------------+--------------------

Toe webbing | Full | Three-fourths

| |

Finger webbing | Three-fourths | One-half

| |

Average snout-vent length (Male) | 41.7 mm. | 37.4 mm.

| |

Tympanum/eye (Male) | 63.2% | 55.8%

| |

Dorsal leg pattern | Barred | Unmarked

| |

Tarsal fold | Tubercular | Absent

| |

Tarsal stripe | Absent or | Broad, indistinct,

| indistinct | or absent

| |

Dorsolateral stripe | Absent | Present

| |

Light line over anus | Broad | Narrow

| |

Flash colors | Red | Yellow

| |

Iris color | Copper | Bronze

---------------------------------+----------------+--------------------

The three specimens from San Lorenzo, Veracruz (USNM 123513-5), were

identified as _Hyla rickardsi_ by Smith (1947:409). The flash colors

have faded in preservative, and so are of no aid in identifying these

specimens. Two are adult females with snout-vent lengths of 35 and 39

mm. In possessing a relatively large tympanum and barred thighs, and in

lacking a dorsolateral stripe they are typical of _loquax_, but in the

amount of webbing on the hands and feet, broad tarsal stripe, and

narrow anal stripe they are like _rickardsi_. The third specimen, a

juvenile, has a snout-vent length of 25 mm. In coloration it resembles

the adults; it has more distinct bars on the limbs. On the basis of

geography these specimens should be _loquax_, for the closest known

record of _rickardsi_ is more than 200 kilometers to the northwest,

whereas _loquax_ is known from several localities around San Lorenzo.

Shannon and Werler (1955:383) described _Hyla axillamembrana_ from the

lower southern slopes of Los Tuxtlas. The unique type is a small male

(27 mm. snout-vent). I have examined the type and find no great

differences between it and small specimens of _loquax_. It is not

possible to determine the color of the thighs, nor was this information

given in the description. _Hyla axillamembrana_ is here considered to

be a synonym of _Hyla loquax_.

=Hyla microcephala martini= Smith

_Oaxaca_: Donaji (15); 43 km. N of Matias Romero (19); Rio

Sarabia (2); Sarabia (11); Tolosita. _Veracruz_: Acayucan

(17); Alvarado (41); Aquilera (21); 8 km. SW of

Coatzacoalcos (10); Cosoleacaque (26); 10 km. SE of

Hueyapan; Naranja (3); Novillero.

This frog is abundant in the Gulf lowlands of the isthmus, where large

breeding congregations were found in grassy ponds on the savannas and

in openings in the forest. Most frequently males were calling from

grasses and reeds in the ponds; many individuals were perched

precariously on thin blades as high as one meter above the water. The

call is a series of low squeaks.

Individuals found at night were pale yellow above with light brown

lines arranged in an irregular pattern on the back, but often forming a

cross or an X-shaped mark in the scapular region. There is a brown

stripe from the nostril to the eye and thence to the groin. Anteriorly

this stripe is bordered above by a thin white or cream-colored line.

Numerous small brown flecks are scattered on the back and dorsal

surface of the shank. In most specimens there are thin transverse brown

bars on the shank. The thighs and undersides of the limbs are golden

yellow; the belly and vocal sac are lemon yellow. The iris is yellowish

brown. During the day individuals assume a pale reddish tan ground

color with darker brown markings. Twenty-five adult males from Alvarado

have an average snout-vent length of 24.1 mm.

=Hyla picta= Guenther

_Oaxaca_: Donaji (8); Sarabia (11); Tolosita (15); Ubero

(6). _Veracruz_: 19 km. N of Acayucan (4); Alvarado (5);

Aquilera; 8 km. SW of Coatzacoalcos; 10 km. SE of Hueyapan

(7); Lerdo de Tejada; Tula (3).

Widespread in the forests, scrub, and savannas on the Gulf lowlands of

the isthmus, these frogs were found breeding at numerous localities.

Males call from grasses and bushes growing in and about ponds. The

call is a high-pitched insect-like trill. At night these frogs are pale

yellow above; they change to light grayish tan during the day. A dark

stripe extends from the nostril to the eye and thence posteriorly to a

point between the axilla and groin. Above this dark stripe is a broader

white stripe. Scattered on the dorsum are brown flecks or spots; the

shanks are marked with poorly-defined cross-bars. The thighs are deep

yellow below and paler above with scattered dark flecks. The belly is

white, and the vocal sac is yellow. The iris is golden. Twenty males

have an average snout-vent length of 21.5 mm.; three females, 24.0 mm.

=Hyla robertmertensi= Taylor

_Oaxaca_: Tapanatepec (28); 7.5 km. NW of Tapanatepec (38);

7.2 km. WNW of Zanatepec (77).

This species was found in the isthmian region only on the Pacific

lowlands at the southern base of the western part of the Sierra Madre

de Chiapas. On July 13, 1956, many large choruses were discovered. The

calling males were on reeds and thorn scrub in and at the edge of

temporary ponds; the call is a cricket-like "creak-creack," quickly

followed by a series of notes "creak-eek-eek-eek-eek."

At night the dorsal ground color is pale yellow; this changes to

pinkish buff during the day. There is a grayish or brown dark stripe

from the nostril to the eye; the stripe continues to the groin. This

dark stripe is bordered above by a narrow white stripe. The belly is

white, and the vocal sac is yellow. The iris is dull reddish brown.

Twenty-five males have an average snout-vent length of 24.7 mm.

=Hyla staufferi= Cope

_Oaxaca_: Chivela; Huilotepec (5); Juchitan (4); Matias

Romero (4); 25 km. N of Matias Romero; Mixtequilla (4); Rio

Sarabia (11); 11 km. S of Santiago Chivela; Sarabia (3);

Tapanatepec (67); Tehuantepec (66); Tolosita (2); Ubero;

Union Hidalgo; Zanatepec (6). _Veracruz_: Acayucan (7);

Alvarado (3); Amatitlan; Aquilera; Ciudad Aleman (3); 8 km.

SW of Coatzacoalcos (9); Cosamaloapan (4); Cosoleacaque (8);

10 km. SE of Hueyapan; Lerdo de Tejada; Novillero (6); Tula

(2).

This is the only species of small hylid that crosses the isthmus.

Calling males were found in and about ponds on the savannas in southern

Veracruz, in ponds in open forest in northern Oaxaca (not in forest

pools), and in temporary pools in the scrub forest on the Pacific

lowlands. Individuals usually called from bushes and reeds in or at the

edge of ponds. The call is a short "braaa." Dates of breeding choruses

indicate that by the time the other small species of hylids in the Gulf

lowlands reach the peak of their breeding season, that of _H.

staufferi_ is essentially over; no large breeding congregations were

found in July. On July 8, 1956, two metamorphosing young were found

clinging to blades of grass in a pond; they had snout-vent lengths of 8

and 9 mm. and tail stumps less than 3 mm. in length. Others were found

on July 13 and 26. The juveniles are nearly unicolor olive green above

and white below.

In life the adults vary greatly in color pattern. The dorsal ground

color is yellowish tan to olive brown with olive brown or dark brown

spots, some of which in certain individuals are connected to form

longitudinal dark stripes. On the posterior surface of the thighs are

small white flecks. The belly is white, and the vocal sac is a rich

yellow. Twenty males have an average snout-vent length of 26.3 mm.;

they have no horny nuptial pads. No noticeable differences in either

color or body proportions were found between the populations on either

side of the isthmus.

=Hylella sumichrasti= Brocchi

_Oaxaca_: Cerro Arenal (5); Cerro San Pedro (2); Escurano;

La Concepcion (41); Portillo Los Nanches (6); San Antonio

(16); 11 km. S of Santiago Chivela (18); Santa Lucia (7);

Tapanatepec (5); Tehuantepec (8); Tenango (49); Tres Cruces

(19).

With the exception of the series from 11 kilometers south of Santiago

Chivela, most of these specimens were found in small arboreal

bromeliads during the dry season. Males were found along a clear,

shallow, rocky stream south of Santiago Chivela on July 6, 1956. The

frogs were calling from bushes and rocks in and along the stream. When

disturbed, they jumped into the water and floated downstream until they

were able to hold onto a rock or other object. The call is a loud

"bra-a-ah." In breeding individuals the dorsum is pale yellow; the

belly is white, and the vocal sac is yellow. The iris is pale golden

yellow. Eighteen males have an average snout-vent length of 25.2 mm.

All have dark brown nuptial tuberosities on the pollex.

Certain diagnostic characters of this species as given by Taylor

(1943a:50) and Taylor and Smith (1945:598) are in need of revision.

_Hylella sumichrasti_ has been characterized as having no vocal sac,

rarely having vomerine teeth, and as having a relatively smooth throat.

The vocal sac in breeding males is quite evident; it is single, median,

and when expanded, spherical. The openings into the vocal sac are

narrow slits along the inner posterior border of the jaw rami. Of 151

specimens studied, 74 have vomerine ridges between the choanae, and 36

of these have one to three teeth on each ridge. The belly and

undersurfaces of the thighs are granular; the throat is only somewhat

less so. The granular condition may be correlated with breeding, for

specimens obtained from bromeliads in the dry season had rather smooth

throats. It seems that the vocal sac atrophys in the non-breeding

season. These seasonal changes may account for the diagnoses given by

Taylor (_op. cit._) and Taylor and Smith (_op. cit._); likewise, since

many of the specimens obtained by Smith in the dry season were

juveniles and subadults, the development of the vomerine ridges could

not be diagnosed properly.

The range of this species encompasses the Pacific slopes of the Isthmus

of Tehuantepec eastward to the upper Cintalapa Valley and vicinity of

Tonala in western Chiapas. Priscilla Starrett collected tadpoles of _H.

sumichrasti_ from a stream 19 km. N of Arriaga, Chiapas. These limited

observations on the ecology of this frog suggest that it breeds in the

fast-moving streams of the Pacific slopes, and that it seeks shelter in

arboreal bromeliads during the dry season.

=Phrynohyas modesta= Taylor and Smith

_Oaxaca_: Tuxtepec. _Veracruz_: 20 km. S of Jesus Carranza;

20 km. ENE of Jesus Carranza (2); Minatitlan.

I have not collected this species in the isthmus. The locality records

indicate that the range is discontinuous (Duellman, 1956:27). The

species occurs on the humid Pacific slopes from south-central Chiapas

eastward to El Salvador and on the humid Gulf lowlands from southern

Veracruz eastward into Tabasco, but is unknown from the dry Pacific

slopes and plains in the isthmus.

The acquisition of several specimens of this species in southern

Veracruz, Tabasco, and Oaxaca, together with a knowledge of the

variation displayed by _Phrynohyas spilomma_, suggests that _modesta_

may be a color variety of _spilomma_.

=Phrynohyas spilomma= Cope

_Oaxaca_: Tapanatepec (3). _Veracruz_: Amatitlan (12);

Chacaltianguis (2); Ciudad Aleman (6); Cosamaloapan;

Novillero (3).

Like the preceding species, this frog is unknown from the arid Pacific

lowlands of the isthmus; its presence at Tapanatepec, a locality

situated in more mesic conditions than prevail on the Plains of

Tehuantepec, indicates that it may have a distribution on the Pacific

slopes much like that of _P. modesta_. Furthermore, this frog was not

detected in the rainforests of the Gulf lowlands; in that region it was

found only in scrub forest and savanna.

On July 26, 1956, numerous choruses of these frogs were heard between

Ciudad Aleman and Tlacotalpan, Veracruz. The call is a loud, nasal

"grawl" repeated continuously. The males call from the water. Several

clasping pairs were found in shallow grassy ponds amidst the scrub

forest. The ground color varies from reddish brown to tan with dark

brown dorsal markings. The iris is golden, and the vocal sacs are dark

olive brown. After a light shower during the dry season, six

individuals were found on the low branches of trees at night near

Ciudad Aleman.

=Phyllomedusa callidryas taylori= Funkhouser

_Oaxaca_: Donaji (9); Sarabia (8); Tolosita (6); Ubero (27).

_Veracruz_: Alvarado (7); Aquilera; Berta; Coatzacoalcos

(9); 10 km. SE of Hueyapan (5); Naranja (17).

In life this frog presents a striking array of colors. The dorsum

varies from pale green to dark olive green; there may be scattered

whitish or cream-colored spots on the back. On the flanks are bright

yellow to deep cream-colored vertical bars separated by pale blue or

purple interspaces. The thighs and undersurfaces of the hind limbs are

golden orange; the belly is yellow, and the throat is cream-colored.

The iris is crimson; the transparent part of the lower eyelid has

golden reticulations. When the frog is resting, the forefeet are folded

beneath the throat, and the limbs are folded tightly against the body.

In this position and with the eyes closed and head flattened, this

gaudy frog assumes the appearance of a small elliptical green leaf.

Throughout the month of July, 1956, _Phyllomedusa_ was breeding in

ponds in or adjacent to the rainforest in northern Oaxaca and in

southern Veracruz. Only at Alvarado was it found breeding in a grassy

pond. Males and females alike were found on bushes and trees in and

around the ponds. The call is a single "wank." Amplexing males continue

to call, but the call is softer and less nasal in quality. The eggs are

encased in pale green gelatin and attached to leaves on branches

overhanging the water. Three egg clutches contained 38, 41, and 46

eggs.

=Phyllomedusa dacnicolor= Cope

_Oaxaca_: Escurano; Tehuantepec.

Although it is abundant on the Pacific lowlands to the northwest in

Guerrero, Michoacan, and Colima, this species is known only from two

specimens from Tehuantepec. There is no apparent physical barrier to

their distribution in the isthmus; in the Balsas Basin the species

lives in a hotter, more arid environment than that at Tehuantepec.

=Gastrophryne usta= Cope

_Oaxaca_: Santa Efigenia; Tehuantepec (10); 24 km. W of

Tehuantepec; Tolosita (2). _Veracruz_: Ayentes (6); La

Oaxaquena; Novillero (2); San Lorenzo.

Calling males were found in open scrub forest near Tehuantepec and in

savannas near Novillero. The specimens from Tolosita were found under

cover in a clearing in the forest (Fugler and Webb, 1957:106).

A Distributional Study of the Amphibians of the Isthmus of Tehuantepec, México · 第 4 / 6 段