Probably the level of the water table in the coastal lowlands and the gradients of the streams in the lowlands and foothills was closely correlated with fluctuation in sea level. If sea level fluctuated as much as 575 feet in the Pleistocene, changes in the level of the water table must have been of considerable magnitude.
During times of glacial advances the lowlands of the isthmus probably were more extensive and had more semi-arid tropical environments than at present, with patches of rainforest existing in sheltered valleys along the major streams. In the course of bio-climatic fluctuation the semi-arid environments (scrub forest and/or savanna) were continuous at times from the Pacific lowlands across the isthmus to the Gulf lowlands. At those times such typical inhabitants of the semi-arid environments as Rhinophrynus dorsalis, Engystomops pustulosus, and Hyla staufferi could have made their way across the isthmus. At times of most extensive glaciation, such as the Illinoian, temperatures in the isthmus probably were low enough to permit the growth of pine-oak forest and cloud forest continuously across the central ridges from the Mexican to the Chiapan-Guatemalan highlands. At those times such highland members of the fauna as Chiropterotriton, Pseudoeurycea, Magnadigita, and the eximia group of Hyla could have crossed the isthmus. During Wisconsin time, climate probably fluctuated less than during previous glaciations; probably no montane environments, except cloud forest, were represented in the isthmus during the Wisconsin. Even at this relatively late date such animals as Lineatriton lineola, Anotheca coronata, and Phyllomedusa moreleti could have crossed the isthmus.
During the interglacial periods, which in the isthmian region were characterized by warmer temperatures, higher sea level and consequently more restricted areas of lowlands, and possibly more rainfall than in the glacial periods, the continuity of pine-oak forest and cloud forest from east to west across the isthmus was interrupted. Probably, too, the semi-arid environments were restricted, and the rainforests were more widespread. At those times animals now inhabiting the rainforests of the Gulf lowlands and those inhabiting the Pacific lowlands of Chiapas and Guatemala could have crossed the isthmus. In this group are species such as Bolitoglossa occidentalis, Eleutherodactylus rhodopis, Microbatrachylus pygmaeus, and Rana palmipes.
The amount of differentiation in isolated populations of amphibians in southern México 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
Gymnopis mexicanus mexicanus Duméril and Bibron
Oaxaca: El Barrio (3); Matías Romero; Tehuantepec (2). Veracruz: Cosamaloapan; Cuatotolapam (2).
The two specimens from Cuatotolapam were collected by Ruthven in an area of mixed savanna and forest. The three specimens (USNM 30535-7) 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 (MCZ 1604) merely bear the data "Tehuantepec, Mexico." They may have come from the town, the district, or from anywhere in the isthmus. The specimen from Matías 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 (1942:475).