University of Kansas Publications
Museum of Natural History

Volume 13, No. 2, pp. 19-72, pls. 1-8, 3 figs.
August 16, 1960

A Distributional Study of the Amphibians
of the Isthmus of Tehuantepec, México

BY
WILLIAM E. DUELLMAN


University of Kansas
Lawrence
1960

University of Kansas Publications
Museum of Natural History

Volume 13, No. 2, pp. 19-72, pls. 1-8, 3 figs.
August 16, 1960

A Distributional Study of the Amphibians
of the Isthmus of Tehuantepec, México

BY
WILLIAM E. DUELLMAN


University of Kansas
Lawrence
1960

University of Kansas Publications, Museum of Natural History

Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Robert W. Wilson

Volume 13, No. 2, pp. 19-72, pls. 1-8, 3 figs.
Published August 16, 1960

University of Kansas
Lawrence, Kansas

PRINTED IN
THE STATE PRINTING PLANT
TOPEKA, KANSAS
1960

28-3859

A Distributional Study of the Amphibians
of the Isthmus of Tehuantepec, México

BY

WILLIAM E. DUELLMAN

CONTENTS

INTRODUCTION

Few regions in Middle America are so important zoogeographically as is the Isthmus of Tehuantepec, that neck of land connecting North America with Central America, separating the Pacific Ocean from the Gulf of Mexico by a distance of only about 220 kilometers (airline), and forming a low break between the highlands of México and those of Central America. Before World War II the isthmus could be reached readily only by railroad or by ocean vessel to Salina Cruz or Coatzacoalcos. With the advent of roads, principally the Trans-isthmian Highway, vast areas of the interior of the isthmus became accessible to biologists. Nevertheless, long before roads were built in the isthmian region collectors and biologists visited it, especially the town of Tehuantepec, from which collections date back to the 1870's. Therefore, it is rather surprising that no attempt has been made to present a faunal list of the amphibians or reptiles of the isthmus. Ruthven (1912) summarized his collections from the vicinity of Cuatotolapam, Veracruz, and Hartweg and Oliver (1940) presented an annotated list of the species collected by them in the vicinity of Tehuantepec. In recent years there have been only a few papers reporting species from the isthmus (Fugler and Webb, 1957; Langebartel and Smith, 1959). The zoogeographic significance of the Isthmus of Tehuantepec is exemplified by the works of Burt (1931), Duellman (1958), Gloyd (1940), Oliver (1948), and Stuart (1941), who in their discussions of evolution and dispersal of various genera of reptiles, pointed out that the Isthmus of Tehuantepec was a region of zoogeographic importance.

Originally I intended to study the entire herpetofauna of the isthmus. But I have not had opportunity to study all of the reptiles, and I have not had the inclination to solve certain taxonomic problems concerning them. The amphibians that I collected, together with all other known specimens in museums, have been studied. Therefore, the present report is concerned only with the amphibians. Only the amphibians of the lowlands of the isthmus have been sampled adequately. Although I have commented on the highland species in the discussion of distribution, they are not included in the systematic section, which deals solely with the 36 species definitely known to occur in the lowlands of the isthmus.

Among the species of amphibians that I would expect to occur in the isthmus, the only one not yet found there is Hyla phaeota. Sufficient specimens of most of the species are available to show their variation in the isthmus. Consequently, the systematics of these amphibians is on a fairly substantial basis. Probably certain species in the isthmian region will be found to be conspecific with others to the south, for example Hyla ebraccata with Hyla leucophyllata and Hyla robertmertensi with Hyla underwoodi. Nevertheless, such taxonomic changes will not affect the distributional picture presented here. Our greatest lack of knowledge concerning the amphibians is about their life histories, as may be illustrated by the following questions, all of which now are without definite answers. Where do many of the small frogs conceal themselves during the dry season? What amount of, if any, interspecific competition exists among several species of tree frogs, all of which breed in the same ponds? What factors in the environment permit certain amphibians, but not others, to live in the humid rainforests, as well as in the arid tropical scrub forest? The answers to these questions and many others must await additional field studies.

The purpose of this paper is to make known the species of amphibians living in the Isthmus of Tehuantepec, to describe the environments in which they live, and to discuss their distribution in the isthmus. With respect to the distribution of animals in the Isthmus of Tehuantepec I will attempt to explain the present patterns of distribution with special reference to climatic fluctuation in the Pleistocene.

Acknowledgments

My extensive field work in the Isthmus of Tehuantepec was made possible by grants from the Penrose Fund of the American Philosophical Society (1956) and the Bache Fund of the National Academy of Sciences (1958). Furthermore, my field work received the hearty support of the Museum of Zoology at the University of Michigan; for their cooperation I am indebted to Norman Hartweg, T. H. Hubbell, and Henry van der Schalie. In the course of my studies I received helpful suggestions from Norman Hartweg, L. C. Stuart, and Charles F. Walker, to whom I am grateful. For permission to examine specimens in their care I thank Doris M. Cochran, Hobart M. Smith, and Richard G. Zweifel. I am deeply indebted to Thomas MacDougall for many suggestions and for aid in preparing the gazetteer. I am most grateful for the efforts of my field companions, Richard E. Etheridge, Jerome B. Tulecke, John Wellman, and especially my wife, Ann S. Duellman, who spent many long days and nights gathering much of the data on which this report is based. Our work in the isthmus was furthered by the generous help and hospitality of many residents, especially the late Wilbur Barker of Tehuantepec, Fortunado Delgado of Rancho Las Hojitas near Acayucan, César Fárjas of Donají, and Juan Mayol of San Andrés Tuxtla. Profesor Jordi Juliá Z. of the Laboratorio de Entomología, Comisión del Papaloapan, Ciudad Alemán, Veracruz, helped make possible my field work in 1959; for this he has my sincere thanks. In conclusion I express my gratitude to Ing. Juan Lozano Franco, Secretaria de Agricultura y Ganadería, for providing me with the necessary permits.

Field Studies in the Isthmus of Tehuantepec

I first visited the Isthmus of Tehuantepec and collected on the Pacific lowlands of the isthmus in July, 1955. At that time heavy rains and impassable roads restricted travelling. In February and March of 1956 my wife and I concentrated our efforts in the central region between the Río Jaltepec and Matías Romero, but also made several trips across the isthmus to gather ecological data in the dry season. In July of the same year, accompanied by Richard E. Etheridge, we again crossed the isthmus several times in order to gather ecological data in the wet season, and studied especially hylid frogs, most of which had not been seen in the dry season. Accompanied by Jerome B. Tulecke and John Wellman, I collected again in the isthmus in July, 1958, between Salina Cruz and Tehuantepec, and between Coatzacoalcos and Cosoleacaque. In March and April, 1959, I collected at Ciudad Alemán. Nearly 1200 specimens of 30 species of amphibians were thus collected in the Isthmus of Tehuantepec; all specimens are now in the Museum of Zoology at the University of Michigan. Of other species known from the isthmus, I have had field experience with all but one (Bolitoglossa veracrucis) in other parts of México.

Sources of Material

There are in museum collections nearly 3000 specimens of amphibians with reliable data from the Isthmus of Tehuantepec. Among the first herpetological specimens collected in the isthmian region are those assembled by Francis Sumichrast in the 1870's from the vicinity of Santa Efigenia and Tapanatepec, Oaxaca. These specimens were sent to the United States National Museum and the Museum National d'Histoire Naturelle in Paris; many served as the types of new species: Bufo canaliferus Cope, Eleutherodactylus rugulosus Cope, Syrrhophus leprus Cope, and Hylella sumichrasti Brocchi. In 1911 Alexander G. Ruthven collected in the savanna country near Cuatotolapam, Veracruz; the report on his collections (1912) is the first dealing with the herpetofauna of a part of the isthmus. His specimens are in the collection of the University of Michigan Museum of Zoology. Norman Hartweg and James A. Oliver collected for the University of Michigan Museum of Zoology in the vicinity of Tehuantepec, Oaxaca, during the summer of 1936. The results of their work were published as an annotated list of species occurring on the Pacific slopes of the isthmus (1940). Hobart M. Smith collected in the vicinity of Tehuantepec in January, 1940; his specimens are in the United States National Museum. Specimens collected by Smith served as the types of Eleutherodactylus avocalis Taylor and Smith and Diaglena reticulata Taylor. Walter W. Dalquest collected vertebrates for the University of Kansas in southern Veracruz in the winters of 1947 and 1948; he spent about six months on the Gulf lowlands of the isthmus, principally in the vicinity of Jesús Carranza. For the past two decades Thomas MacDougall, a resident of New York City, has spent his winters collecting biological specimens in southern México. He makes his headquarters at Tehuantepec, but his compulsion to see the "back country" has taken him to many remote parts of southern Oaxaca. His earlier collections are in the American Museum of Natural History; the later ones are in the University of Illinois Museum of Natural History.

Minor collections include those made by Matthew W. Stirling at San Lorenzo, Veracruz, February-April, 1946 (United States National Museum), by Fred G. Thompson on a trip across the isthmus in December, 1955 (University of Michigan Museum of Zoology), by the University of Kansas Museum of Natural History field party under the direction of Rollin H. Baker at Tolosita, Oaxaca, and by David A. Langebartel and associates from southern Oaxaca in June, 1958 (University of Illinois Museum of Natural History).

In the collections of the United States National Museum are several species of amphibians sent to the museum from Tehuantepec by Francis Sumichrast. These include Bolitoglossa platydactyla (USNM 30305, 30344-6, 30528), Bolitoglossa rufescens (10042), Chiropterotriton chiropterus (30347), Lineatriton lineola (30353), Parvimolge townsendi (30352), Pseudoeurycea cephalica (30350), Thorius pennatulus (30348-9), Hyla miotympanum (30302-3), and Hyla picta (30304). Because of the poor condition of the specimens, determinations of those listed as Bolitoglossa rufescens and Pseudoeurycea cephalica are uncertain. With the exception of the Bolitoglossa rufescens, which is stated to have come from Santa Efigenia, all of these specimens are catalogued as having come from Tehuantepec. None of these species has since been recorded from the Pacific slopes of the isthmus; however, they all occur in the vicinity of Orizaba, Veracruz. Probably Sumichrast carried the specimens with him from Orizaba, his home before moving to Santa Efigenia, and shipped them from Tehuantepec to the United States National Museum. These species definitely should not be considered as inhabitants of the Pacific slopes of the Isthmus of Tehuantepec.

DESCRIPTION OF THE ISTHMUS OF TEHUANTEPEC

For the purposes of this description, the lowlands of the isthmus may be divided into three parts—the Gulf Coastal Plain, the central ridges, and the Pacific Coastal Plain, which in the isthmus is called the Plains of Tehuantepec (Figs. 1 and 2).

The Gulf Coastal Plain is broad and fairly level near the coast, but rolling in the interior. The plain, throughout most of its length in the isthmus, is at least 75 kilometers wide. The majority of the region in the isthmus is drained by the Río Coatzacoalcos, which flows in a northerly course to the Gulf of Mexico. The western part is drained by the Río San Juan, the principal tributary of the Río Papaloapan. Behind the coastal dunes are frequent, and sometimes large, lagoons. Immediately inland from Coatzacoalcos and along the lower stretches of the Río Papaloapan are extensive marshes. Essentially the entire coastal plain, with the exception of the coastal dunes, consists of rich alluvial deposits.

The central ridges extend from the Río Jaltepec southward to within 40 kilometers of the Pacific coast. It is in this area that the continuity of the high ridges and volcanic peaks, which extend nearly the entire length of the Americas, is interrupted at a point almost directly in line with the shortest distance between the two oceans. The northern part of this central region consists of hills dissected by tributaries of the Río Coatzacoalcos; the principal ones from north to south are—Río Jaltepec, Río Tortuguero, Río Sarabia, and Río Malatengo. The plains of Chivela are south of these rivers and lie at an elevation of about 200 meters; at the southern edge of these plains a range of hills rises to 250 to 400 meters above sea level. These hills drop abruptly to the Plains of Tehuantepec. In the northern and central parts of this central region the rocks are granitic; the hills to the south of the Plains of Chivela are limestone.

The Pacific Coastal Plain or Plains of Tehuantepec have a maximum width of about 30 kilometers. From the base of the hills at an elevation of about 75 meters the plains slope gradually to the sea. To the west of the Río Tehuantepec and to the east of the Plains of Tehuantepec at the base of the Sierra Madre de Chiapas, the coastal plain becomes much narrower; in these places the continuity of the plain is frequently interrupted by low north-south ridges extending outward from the mountains or by isolated hills. The soil is poor, varying from volcanic rock to gravel and sand.

Climate

The prevailing winds are from the north across the Gulf of Mexico. These moisture-laden winds precipitate most of their moisture north of the central ridges. This results in high rainfall on the northern slopes and Gulf Coastal Plain and relatively little rainfall on the southern slopes and the Pacific Coastal Plain. Precipitation is cyclic; there is a marked wet and a dry season throughout the region, but this is most noticeable on the Pacific lowlands (Fig. 3). At Salina Cruz on the Pacific Ocean the average annual rainfall is 1040 mm. (Contreras, 1942); of this amount, only 15 mm. falls from November through April. On the Gulf Coastal Plain (Minatitlán station) the average annual rainfall is 3085 mm. In this region the driest months are February through May, during which time 236 mm. of rain falls. At Salina Cruz the wettest month is June; at Minatitlán it is September. There is little variation in temperature throughout the isthmus; the average annual temperature at Salina Cruz is 26.6° C.; that at Minatitlán is 26.2° C. During the winter when masses of air from the arctic move southward into the Great Plains of the United States, cool winds blow across the isthmus. These are usually accompanied by overcast sky and sometimes a slight amount of precipitation. These "nortes" may cause a drop in temperature of about six to eight degrees in a few hours.

The topography and climate combine to produce drastically different types of climax vegetation on the northern and southern lowlands of the isthmus. The picture is somewhat complicated by the savannas on the Gulf Coastal Plain, which, as will be shown later, are dependent upon edaphic features more than climatic conditions. The following brief account of the vegetation in the Isthmus of Tehuantepec is based on data provided by Williams (1939) and Goldman (1951), supplemented by personal observations. The purpose of this description is not to analyze the flora of the isthmus, but to give the reader a picture of this aspect of the biota of the major environments with which I shall be concerned in the ensuing discourse on the amphibians of the region. The three divisions of the isthmus recognized in the account of the physiography serve equally well in describing the vegetation. Those divisions are as follows:

Gulf Lowlands

On the lowlands north of the continental divide and extending to the Gulf of Mexico are three major types of vegetation—tropical rainforest, arid tropical scrub forest, and savanna. Aside from these, there are marshes and lagoons near the coast.

On the coastal dunes there are thickets of sea grape, patches of Cenchrus, and clumps or scattered Opuntia. The lagoons are bordered by mangrove thickets made up primarily of Lonchocarpus hondurensis. In the marshes along the lower Río Coatzacoalcos and Río Papaloapan the tall tough grass, Gynerium sagittatum, is common.

According to Beard (1953: 291) the development of savanna vegetation is dependent upon soil, topography, and drainage. Level regions having permeable soil horizons lying on top of an impermeable horizon provide poor drainage. In most savanna regions in the Americas the grasslands become waterlogged or even partly flooded during the rainy season and desiccated in the dry season. Many ecologists and phytogeographers have postulated that savannas are either man made or are examples of a fire climax. Beard (op. cit.: 203) provided multitudinous evidence that the association of savanna vegetation and certain types of edaphic and topographic conditions was so strongly marked that grassland is the natural vegetation in these areas.

Savannas are scattered through southern Veracruz eastward to British Honduras. These usually are grasslands having scattered trees or clumps of trees around depressions, which may contain water throughout the year (Pl. 1, fig. 1). According to Williams (op. cit.), the most common trees in the savannas in southern Veracruz are Ceiba pentandra, Chlorophora tinctoria, and Byrsonima crassifolia.

Lying in a rain shadow cast by the Tuxtlas and on sandy and well-drained soils is a dense xerophytic forest. The crown of this deciduous forest usually is little more than ten to twelve meters above the ground (Pl. 1, fig. 2). Conspicuous trees in this scrub forest are Acacia cornigera, Bauhinia latifolia, Calliandra bijuga, Cassia laevigata, Guazuma ulmifolia, and various species of Bursera.

The most extensive type of vegetation on the Gulf Coastal Plain is a tall evergreen forest resembling tropical rainforest. Although this forest is made up of many species of trees that are characteristic of true rainforest, the forest on the Gulf Coastal Plain cannot be classified as true rainforest, neither by the climatic conditions, nor the structure of the forest. The seasonal variation in rainfall probably is the chief factor in hindering the development of a rainforest climax vegetation. Usually a minimum of 65 mm. of rainfall each month is considered essential for the development of true rainforest. At Minatitlán the average rainfall for March (39 mm.) and April (36 mm.) is far below this minimum. Structurally, this forest has a crown about 30-35 meters above the ground but individual trees rising five meters or more above the crown (Pl. 2, figs. 1-2). There is no clear stratification within the forest; in many parts of it there are dense growths of bushes, small trees, and palms. The forest on the Gulf Coastal Plain, therefore, most properly might be referred to as a quasi-rainforest, a term that has been applied to other such forests in tropical America.

Among the abundant and dominant trees in this forest are Swietenia macrophylla, Calophyllum brasiliense, Achras zapota, Ceiba pentandra, Castilla elastica, Cedrela mexicana, Tabebuia Donnell-Smithi, Calocarpum mammosum, Bombax ellipticum, and a variety of Ficus. Epiphytes and Ilianas are abundant.

Central Ridges

The vegetation of the central ridges of the isthmus is, for the most part, transitional between the tall rainforest of the Gulf Coastal Plain and the low xerophytic scrub forest of the semi-arid Pacific Coastal Plain. On the northern slopes of the ridges the rainforest is more poorly developed than on the plains to the north. Many of the same species of trees are present, including Ceiba pentandra, Cedrela mexicana, Swietenia macrophylla, and Ficus sp.; nevertheless, these seldom are as large as members of the same species in the forest on the plains. Other species present on the forested slopes include Tabebuia Donnell-Smithi, Zanthoxylum melanostictum, Pithecolobium arboreum, and a species of Pterocarpus. The structure of this forest differs from that on the Gulf Coastal Plain in that there is no continuous upper canopy and there is a dense undergrowth (Pl. 3, fig.1). This type of forest extends from Mogoñe southward to about Matías Romero.

In the vicinity of Matías Romero open pine-oak forest (Pinus caribaea and Quercus sp.) is found on some ridges as low as 250 meters above sea level.

On the Plains of Chivela in the southern part of the central region the vegetation takes on a semi-arid appearance, especially in a savanna on the plains. Clumps of small trees and bushes, consisting of Croton nivea, Cordia cana, Jacquinia aurantiaca, Calycophyllum candidissimum, and Cassia emarginata, are scattered on a grassy plain, from which rise widely-spaced palms of an unknown species (Pl. 3, fig. 2).

Pacific Coastal Plain

The vegetation of the Pacific lowlands definitely is semi-arid in character. Most of the trees are deciduous, thorny, and short. During the dry season the landscape presents a barren appearance, but shortly after the first summer rains dense green foliage appears (Pl. 4, figs. 1 and 2). Between Juchitán and La Ventosa few trees are more than two meters high (Pl. 5, fig. 1). In many areas the trees and bushes form an almost impenetrable tangle, whereas on especially rocky soils or on slopes those plants are more widely spaced. Abundant and widespread species of trees on the Plains of Tehuantepec include Acacia cymbispina, Prosopis chilensis, Caesalpinia coriaria, Caesalpinia eriostachys, Celtis iguanaea, Cordia brevispicata, Jatropha aconitifolia, and Crescentia alata.

Montane Vegetation

In order to illustrate the interruption of subtropical and temperate types of vegetation by the lowlands of the Isthmus of Tehuantepec, it is necessary to digress for a moment from the isthmus and consider the types of vegetation present on the adjacent highlands. On the higher peaks, such as Cerro de Zempoaltepetl, above about 2500 meters is fir forest (Abies religiosa); lower on the slopes are extensive pine forests, which on some slopes are mixed with oak or replaced entirely by oaks. Subtropical cloud forest, characterized by relatively cool temperatures and high humidity, is found at elevations usually between 1000 and 1800 meters on the windward slopes of the Sierra Madre Oriental in Veracruz and northern Oaxaca and on the northern and southern slopes of the Chiapan-Guatemalan Highlands. None of these forest types is continuous across the Isthmus of Tehuantepec.

The Sierra de los Tuxtlas

Although actually located in the region of the Isthmus of Tehuantepec, the Sierra de los Tuxtlas, because of its isolated position, need not be considered in great detail in analyzing the distribution of animals inhabiting the lowlands of the isthmus. Nevertheless because some species living in the highlands adjacent to the isthmus also live in the Tuxtlas, this range is briefly described here. The Sierra de los Tuxtlas is a range of volcanos lying near the Gulf Coast in southern Veracruz between the mouths of the Río Papaloapan and the Río Coatzacoalcos. Volcán San Martín, the highest peak, rises above 1800 meters. This range of volcanos is surrounded by lowlands, which immediately to the south and west are covered with savanna and in places by scrub forest. The luxuriant nature of the vegetation on these volcanos indicates that this range receives much more rainfall than the surrounding lowlands. Especially on the northern slopes, tropical rainforest is well developed; this is replaced at about 1200 meters by cloud forest. The southern and western slopes are drier, for the lower slopes are covered with a scrubby, but evergreen, forest.

Detailed comments on the herpetofauna of the Tuxtlas have been omitted purposefully, for the reptiles and amphibians of the region currently are being studied by Douglas Robinson.

GAZETTEER

The following localities are those referred to in the text. The name of the locality (listed alphabetically by states) is followed by latitude, longitude, elevation, general description (town, ranch, etc.), and general type of habitat. Unless otherwise noted, distances are straight-line (airline) distances in kilometers. The localities have been plotted from the American Geographical Society's "Map of Hispanic America on the Scale of 1:1,000,000" (Millionth Map). Numbers in brackets identify the position of a locality on the accompanying map (Fig. 1).

Oaxaca

Veracruz