Read Ebook: A Distributional Study of the Amphibians of the Isthmus of Tehuantepec México by Duellman William Edward

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The amount of differentiation in isolated populations of amphibians in southern M?xico and northern Central America gives some idea of relative lengths of time of isolation from related populations. Those populations inhabiting high mountain environments on either side of the isthmus are specifically distinct. Some populations inhabiting cloud forests lower on the mountains are specifically distinct from related populations on the other side of the isthmus; between others there is no recognizable differentiation. Even though many populations are isolated from other populations of the same species in the lowlands of the isthmus, there is no apparent speciation. This indicates that the lowland environments and their inhabitants have been isolated from one another for a shorter time than have the highland environments and their inhabitants.

ACCOUNTS OF SPECIES

For each species of amphibian known to occur in the lowlands of the Isthmus of Tehuantepec, localities where one or more specimens were collected are listed, and variation, ecology, and life histories are discussed. A total of 2833 specimens has been examined for the purposes of this study. Individual specimens cited in the text are listed with catalogue numbers and abbreviations of the name of the museum, as follows:

AMNH American Museum of Natural History KU University of Kansas Museum of Natural History MCZ Museum of Comparative Zoology, Harvard College UIMNH University of Illinois Museum of Natural History UMMZ University of Michigan Museum of Zoology USNM United States National Museum

The two specimens from Cuatotolapam were collected by Ruthven in an area of mixed savanna and forest. The three specimens listed above from El Barrio were collected by Sumichrast; possibly they came from another locality. The city of Tehuantepec is divided into seven districts called "barrios." The two specimens listed from Tehuantepec merely bear the data "Tehuantepec, Mexico." They may have come from the town, the district, or from anywhere in the isthmus. The specimen from Mat?as Romero has 109 primary and 67 secondary annuli, a length of 400 mm., and a diameter of 19 mm.; the one from Cosamaloapan has 106 primary and 58 secondary annuli, a length of 397 mm., and a diameter of 19 mm. Data on the other specimens were recorded by Dunn .

Known only from the Gulf lowlands in the isthmian region, this species has been taken in a variety of habitats within the humid forest area: under outer leaves of banana plants, under a rock along a stream, under a log in a plowed field, and on a reed in a pond at night. Three adult males have an average snout-vent length of 44 mm. and a tail-length of 41 mm. In life the color of the dorsum varied from orange-yellow to orange-tan, usually being more orange on the tail. The iris was a reddish orange.

This species is known only from the type series collected at night on a limestone cliff by Walter W. Dalquest. If this salamander is restricted to this type of habitat, it should be found in the region of extensive limestone outcroppings in northern Chiapas and southern Tabasco.

This species inhabits the scrub forests of the Pacific coastal plain and the savannas in southern Veracruz; apparently it does not occur in rainforest. Consequently, its distribution in the isthmus is discontinuous.

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Breeding congregations were found after heavy rains at Tehuantepec on July 5, 1956, at Cosamaloapan, Novillero, and Amatitl?n on July 26, 1956, and at Salina Cruz on July 6, 1958. The call is a long "worrp" made while the male is floating on the surface of the pond. The small heads, small limbs, and greatly inflated bodies cause the calling males to resemble miniature caricature balloons . Amplexus is inguinal. These toads are notably wary, even when calling. Often the beam of a flashlight or the slightest disturbance of the water will cause the males to stop calling. The body is deflated with one last nauseous note, and the frog sinks beneath the surface of the water and swims away with short slow kicks of the hind feet.

This small toad apparently is restricted to the Pacific lowlands from the Isthmus of Tehuantepec eastward to Guatemala. At Zanatepec on July 13, 1956, males were calling from a flooded field bordered by scrub forest. The call is a rather loud nasal racket. Living individuals vary greatly in coloration. Some have yellowish tan flanks and dorsum and an orange middorsal stripe; others have a pale red dorsum, yellow flanks, and a cream middorsal stripe .

This large toad is abundant throughout the lowlands of the isthmus. The loud rattling call of males was heard on rainy nights throughout the summer. In March, 1956, several adults were found in a small cave back of a spring at Agua Caliente.

This toad is abundant on the Pacific lowlands, where it inhabits both open and dense scrub forest. On the Gulf lowlands its distribution seems to be limited to xeric coastal habitats. Aside from the specimens from Alvarado and Coatzacoalcos, it is known in Veracruz only from Boca del R?o .

Individuals were found in both wet and dry seasons. In the dry season they were most frequently found in rainforest, whereas in the rainy season breeding congregations were found in savannas as well. This toad occurs throughout the Gulf lowlands and on the Pacific slopes and in the Grijalva Valley of Chiapas and Guatemala, but not on the Pacific lowlands of the isthmus.

In the dry season frogs of this species were in humid forests, where they were most frequently found along small streams and in ravines. The species is widespread in the Gulf lowlands, but does not occur on the Plains of Tehuantepec. It does inhabit the Pacific slopes on the foothills of the Sierra Madre de Chiapas, the western part of which extends into eastern Oaxaca near Tapanatepec.

In addition to the specimens from the lowlands of the isthmus, for the purposes of the following discussion, I have included data on two specimens from the southern slopes of the Sierra del Sur in Oaxaca and on several specimens from Los Tuxtlas in Veracruz .

In specimens from Los Tuxtlas the tarsal fold is well-developed and extends two-thirds to three-fourths the length of the tarsus; the tibiotarsal articulation reaches the nostril and sometimes slightly beyond the tip of the snout. In males the tympanum is nearly equal to the diameter of the eye; in females it is about one-half the diameter of the eye. The posterior surfaces of the thighs are dark brown or black with whitish or cream-colored spots, flecks, or irregular mottling. The tarsal fold is dark; the throat is pale in some individuals, but in most is mottled with dark brown or gray flecks. Individuals from La Princesa near the continental divide in Oaxaca show the same variation in body proportions and development of the tarsal fold. The posterior surfaces of the thighs are dark brown indistinctly mottled with lighter brown. The throat is dark brown. Specimens from the Pacific slopes of Oaxaca, including the Plains of Tehuantepec, have dark brown thighs mottled with dusty cream. The tibiotarsal articulation extends slightly beyond the tip of the snout in all specimens. In males the tympanum is equal to about two-thirds the diameter of the eye. Duellman discussed the variation in these characters in populations in Colima, Jalisco, and Michoac?n.

Of the 200 specimens examined, 15 have a middorsal stripe that is red or yellow. The iris varies from a copper to a dark golden color and shines bright red at night. Many of the specimens are juveniles; these were collected in the dry season, at which time they were found beneath rocks along streams, in road culverts where there was some water, and in holes in banks and cliffs.

Although the type locality is stated to be Santa Efigenia on the Pacific slopes of the Sierra Madre de Chiapas in eastern Oaxaca, the type specimen probably came from the northern slopes of the mountains. All other known specimens are from the Gulf slopes and lowlands, and from several localities in Los Tuxtlas. Details concerning specimens from the isthmus and other parts of the range were given by Duellman .

In the isthmian region this frog is known only from the Pacific slopes and the Plains of Tehuantepec. Males call from the ground and from trees to heights of about four meters. The call is a single, high, long "peep."

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 ; 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 . The large series from Uni?n 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.

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.

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 ; males called from the bank near the water, and clasping pairs were found only on land . The call is a loud, nasal "braaa," two to three seconds in duration. Amplexus is axillary.

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 , 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.

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, M?xico, in humid forests to El Pet?n, Guatemala.

In the isthmus this species is known only from the humid forests of the Gulf lowlands; it is also known from Boca del R?o, 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.

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

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.

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.

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.

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.

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.

I have not collected this species in the isthmus. The locality records indicate that the range is discontinuous . 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.

On July 26, 1956, numerous choruses of these frogs were heard between Ciudad Alem?n 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 Alem?n.

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.

Although it is abundant on the Pacific lowlands to the northwest in Guerrero, Michoac?n, 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.

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 .

Tadpoles were found in a small sluggish tributary to the R?o Sarabia.

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