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diff --git a/38440-h/38440-h.htm b/38440-h/38440-h.htm new file mode 100644 index 0000000..aed90e1 --- /dev/null +++ b/38440-h/38440-h.htm @@ -0,0 +1,3915 @@ +<!DOCTYPE html PUBLIC "-//W3C//DTD XHTML 1.0 Strict//EN" + "http://www.w3.org/TR/xhtml1/DTD/xhtml1-strict.dtd"> +<html xmlns="http://www.w3.org/1999/xhtml" xml:lang="en" lang="en"> + <head> + <meta http-equiv="Content-Type" content="text/html;charset=iso-8859-1" /> + <meta http-equiv="Content-Style-Type" content="text/css" /> + <title> + The Project Gutenberg eBook of A Distributional Study of the Amphibians +of the Isthmus of Tehuantepec, México, by William E. Duellman. + </title> + <style type="text/css"> + +body { + margin-left: 10%; + margin-right: 10%; +} + + h1,h2,h3,h4 { + text-align: center; /* all headings centered */ + clear: both; +} + +p { + margin-top: .75em; + text-align: justify; + margin-bottom: .75em; +} + +p.title { text-align: center; text-indent: 0; + font-weight: bold; + line-height: 1.4; margin-bottom: 1em; } + +hr.tb {width: 45%;} +hr.chap {width: 65%} +hr.full {width: 95%;} + +hr.r5 {width: 5%; margin-top: .2em; margin-bottom: .2em;} + +table { + margin-left: auto; + margin-right: auto; +} + +.pagenum { /* uncomment the next line for invisible page numbers */ + /* visibility: hidden; */ + position: absolute; + left: 92%; + font-size: smaller; + text-align: right; +} + +.blockquot { + margin-left: 5%; + margin-right: 10%; +} + +.center {text-align: center;} + +.smcap {font-variant: small-caps;} + +.u {text-decoration: underline;} + +.caption {font-weight: bold;} + +.figcenter { + margin: auto; + text-align: center; +} + + +.i2 { + display: block; + margin-left: 2em; + padding-left: 3em; + text-indent: -3em; +} + +.transnote {background-color: #E6E6FA; + color: black; + font-size:smaller; + padding:0.5em; + margin-bottom:5em; + font-family:sans-serif, serif; } + + </style> + </head> +<body> + + +<pre> + +The Project Gutenberg EBook of A Distributional Study of the Amphibians of +the Isthmus of Tehuantepec, Mexico, by William E. Duellman + +This eBook is for the use of anyone anywhere at no cost and with +almost no restrictions whatsoever. You may copy it, give it away or +re-use it under the terms of the Project Gutenberg License included +with this eBook or online at www.gutenberg.org + + +Title: A Distributional Study of the Amphibians of the Isthmus of Tehuantepec, Mexico + +Author: William E. Duellman + +Release Date: December 30, 2011 [EBook #38440] + +Language: English + +Character set encoding: ISO-8859-1 + +*** START OF THIS PROJECT GUTENBERG EBOOK A DISTRIBUTIONAL STUDY OF *** + + + + +Produced by Chris Curnow, Joseph Cooper, Diane Monico, and +the Online Distributed Proofreading Team at +http://www.pgdp.net + + + + + + +</pre> + + + + + + +<hr class="tb" /> +<p class="title"> +<span class="smcap">University of Kansas Publications</span><br /> +<span class="smcap">Museum of Natural History</span></p> +<hr class="r5" /> +<p class="title">Volume 13, No. 2, pp. 19-72, pls. 1-8, 3 figs.<br /> + +August 16, 1960</p> +<hr class="tb" /> + +<h1>A Distributional Study of the Amphibians<br /> +of the Isthmus of Tehuantepec, México</h1> + +<p class="title">BY<br /> +WILLIAM E. DUELLMAN<br /><br /><br /> + +<span class="smcap">University of Kansas</span><br /> +<span class="smcap">Lawrence</span><br /> +1960 +</p> + + +<p><span class="pagenum"><a name="Page_19" id="Page_19">[Pg 19]</a></span></p> +<hr class="tb" /> +<p class="title"> +<span class="smcap">University of Kansas Publications</span><br /> +<span class="smcap">Museum of Natural History</span></p> +<hr class="r5" /> +<p class="title">Volume 13, No. 2, pp. 19-72, pls. 1-8, 3 figs.<br /> + +August 16, 1960</p> +<hr class="tb" /> + +<h1>A Distributional Study of the Amphibians<br /> +of the Isthmus of Tehuantepec, México</h1> + +<p class="title">BY<br /> +WILLIAM E. DUELLMAN<br /><br /><br /> + +<span class="smcap">University of Kansas</span><br /> +<span class="smcap">Lawrence</span><br /> +1960 +</p> +<hr class="chap" /> + + +<p><span class="pagenum"><a name="Page_20" id="Page_20">[Pg 20]</a></span></p> +<p class="title"> +<span class="smcap">University of Kansas Publications, Museum of Natural History</span><br /> +<br /> +Editors: E. Raymond Hall, Chairman, Henry S. Fitch,<br /> +Robert W. Wilson<br /> +<br /> +Volume 13, No. 2, pp. 19-72, pls. 1-8, 3 figs.<br /> +Published August 16, 1960<br /> +<br /> +<span class="smcap">University of Kansas</span><br /> +Lawrence, Kansas<br /> +<br /> +<small>PRINTED IN</small><br /> +<small>THE STATE PRINTING PLANT</small><br /> +<small>TOPEKA, KANSAS</small><br /> +<small>1960</small><br /> +<br /> +28-3859<br /> +</p> + + + +<hr class="chap" /><p><span class="pagenum"><a name="Page_21" id="Page_21">[Pg 21]</a></span></p> +<h2>A Distributional Study of the Amphibians<br /> +of the Isthmus of Tehuantepec, México</h2> + +<p class="title">BY<br /><br /> + +WILLIAM E. DUELLMAN</p> + +<hr class="r5" /> + + +<h2><a name="CONTENTS" id="CONTENTS"></a>CONTENTS</h2> + + +<div class="center"> +<table border="0" cellpadding="4" cellspacing="0" summary="toc"> +<tr><th align="left"> </th><th align="right"><small>PAGE</small></th></tr> +<tr><td align="left"><span class="smcap">Introduction</span></td><td align="right"><a href="#INTRODUCTION">21</a></td></tr> +<tr><td align="left"><span style="margin-left: 1em;">Acknowledgments</span></td><td align="right"> <a href="#Page_23">23</a></td></tr> +<tr><td align="left"><span style="margin-left: 1em;">Field Studies in the Isthmus of Tehuantepec</span></td><td align="right"><a href="#Page_23">23</a></td></tr> +<tr><td align="left"><span style="margin-left: 1em;">Sources of Material</span></td><td align="right"><a href="#Page_24">24</a></td></tr> +<tr><td align="left"><span class="smcap">Description of the Isthmus of Tehuantepec</span></td><td align="right"><a href="#Page_25">25</a></td></tr> +<tr><td align="left"><span style="margin-left: 1em;">Physiography</span></td><td align="right"><a href="#Page_25">25</a></td></tr> +<tr><td align="left"><span style="margin-left: 1em;">Climate</span></td><td align="right"><a href="#Page_28">28</a></td></tr> +<tr><td align="left"><span style="margin-left: 1em;">Vegetation</span></td><td align="right"><a href="#Page_29">29</a></td></tr> +<tr><td align="left"><span style="margin-left: 1em;">The Sierra de los Tuxtlas</span></td><td align="right"><a href="#Page_32">32</a></td></tr> +<tr><td align="left"><span class="smcap">Gazetteer</span></td><td align="right"><a href="#Page_33">33</a></td></tr> +<tr><td align="left"><span class="smcap">The Amphibian Fauna of the Lowlands</span></td><td align="right"><a href="#Page_37">37</a></td></tr> +<tr><td align="left"><span style="margin-left: 1em;">Composition of the Fauna</span></td><td align="right"><a href="#Page_37">37</a></td></tr> +<tr><td align="left"><span style="margin-left: 1em;">Ecology of the Fauna</span></td><td align="right"><a href="#Page_38">38</a></td></tr> +<tr><td align="left"><span style="margin-left: 1em;">Distribution of the Fauna</span></td><td align="right"><a href="#Page_42">42</a></td></tr> +<tr><td align="left"><span class="smcap">The Amphibian Fauna of the Foothills and Adjacent Highlands</span></td><td align="right"><a href="#Page_44">44</a></td></tr> +<tr><td align="left"><span class="smcap">Establishment of Present Patterns of Distribution</span></td><td align="right"><a href="#Page_45">45</a></td></tr> +<tr><td align="left"><span class="smcap">Accounts of Species</span></td><td align="right"><a href="#Page_49">49</a></td></tr> +<tr><td align="left"><span class="smcap">Summary</span></td><td align="right"><a href="#Page_68">68</a></td></tr> +<tr><td align="left"><span class="smcap">Literature Cited</span></td><td align="right"><a href="#Page_69">69</a></td></tr> +</table></div> + + + + +<h2><a name="INTRODUCTION" id="INTRODUCTION"></a>INTRODUCTION</h2> + + +<p>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<span class="pagenum"><a name="Page_22" id="Page_22">[Pg 22]</a></span> +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.</p> + +<p>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.</p> + +<p>Among the species of amphibians that I would expect to occur in the +isthmus, the only one not yet found there is <i>Hyla phaeota</i>. 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 <i>Hyla ebraccata</i> with <i>Hyla leucophyllata</i> and <i>Hyla +robertmertensi</i> with <i>Hyla underwoodi</i>. 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<span class="pagenum"><a name="Page_23" id="Page_23">[Pg 23]</a></span> +humid rainforests, as well as in the arid tropical scrub forest? The answers +to these questions and many others must await additional field studies.</p> + +<p>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.</p> + + +<h3><i>Acknowledgments</i></h3> + +<p>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.</p> + + +<h3><i>Field Studies in the Isthmus of Tehuantepec</i></h3> + +<p>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<span class="pagenum"><a name="Page_24" id="Page_24">[Pg 24]</a></span> 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 (<i>Bolitoglossa +veracrucis</i>) in other parts of México.</p> + + +<h3><i>Sources of Material</i></h3> + +<p>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: <i>Bufo +canaliferus</i> Cope, <i>Eleutherodactylus rugulosus</i> Cope, <i>Syrrhophus +leprus</i> Cope, and <i>Hylella sumichrasti</i> 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 +<i>Eleutherodactylus avocalis</i> Taylor and Smith and <i>Diaglena reticulata</i> +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.</p> + +<p>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).</p> + +<p>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 <i>Bolitoglossa platydactyla</i> (USNM 30305, +30344-6, 30528), <i>Bolitoglossa rufescens</i> (10042), <i>Chiropterotriton +chiropterus</i> (30347), <i>Lineatriton lineola</i> (30353), <i>Parvimolge +townsendi</i> (30352), <i>Pseudoeurycea cephalica</i> (30350), <i>Thorius +pennatulus</i> (30348-9), <i>Hyla miotympanum</i> (30302-3),<span class="pagenum"><a name="Page_25" id="Page_25">[Pg 25]</a></span> and <i>Hyla picta</i> +(30304). Because of the poor condition of the specimens, determinations +of those listed as <i>Bolitoglossa rufescens</i> and <i>Pseudoeurycea +cephalica</i> are uncertain. With the exception of the <i>Bolitoglossa +rufescens</i>, 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.</p> + + + + +<h2><a name="DESCRIPTION_OF_THE_ISTHMUS_OF_TEHUANTEPEC" id="DESCRIPTION_OF_THE_ISTHMUS_OF_TEHUANTEPEC"></a>DESCRIPTION OF THE ISTHMUS OF TEHUANTEPEC</h2> + + +<p>The Isthmus of Tehuantepec is a strip of land forming a low pass, which +separates the mountain masses of México proper from those of Central +America, and at the same time provides a continuum of lowlands from the +Gulf of Mexico to the Pacific Ocean. This topography combines with the +climatic conditions to create extremely diverse environments, the +distribution of which can be adequately understood only after an +acquaintance with the topography and climate of the region.</p> + + +<h3><i>Physiography</i></h3> + +<p>In east-central Oaxaca the mountain masses comprising the Sierra Madre +Oriental and the Sierra del Sur terminate in a series of ranges—Sierra +de Juárez, Sierra de los Míjes, and Sierra de Choapam. From lofty +peaks, such as Cerro de Zempoaltepetl (3400 meters), the highlands +diminish eastward to succeedingly lower ridges, until in the middle of +the Isthmus of Tehuantepec the continental divide is about 250 meters +above sea level. Eastward from this low divide the land rises to form +the Sierra Madre de Chiapas, which is continuous with the highland +masses of Guatemala.</p> + +<p>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 (<a href="#fig1">Figs. 1</a> and <a href="#fig2">2</a>).</p> + +<p>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<span class="pagenum"><a name="Page_26" id="Page_26">[Pg 26]</a></span> 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.</p> + +<div class="figcenter" style="width: 416px;"> +<a name="fig1" id="fig1"></a> +<a href="images/i_010g.jpg"> +<img src="images/i_010.jpg" width="416" height="600" alt="Fig. 1. Map of the Isthmus of Tehuantepec based on the +American Geographical Society's Map of Hispanic America on the Scale +of 1:1,000,000. + +The localities shown are numbered in the gazetteer; the numerical +sequence of localities is an arrangement whereby north takes precedence +over south and west over east. 1. Alvarado. 2. Lerdo de Tejada. 3. +Tlacotalpan. 4. Tula. 5. Tecolapan. 6. Amatitlán. 7. Cosamaloapan. 8. +Chacaltianguis. 9. Novillero. 10. Ciudad Alemán. 11. Papaloapan. 12. +Tuxtepec. 13. Cuatotolapam. 14. Hueyapan. 15. Berta. 16. Coatzacoalcos. +17. Ayentes. 18. Río de las Playas. 19. Cosoleacaque. 20. Minatitlán. +21. Acayucan. 22. Aquilera. 23. San Lorenzo. 24. Naranja. 25. Suchil. +26. Jesús Carranza. 27. La Oaxaqueña. 28. Ubero. 29. Donají. 30. +Tolosita. 31. El Modelo. 32. Sarabia, 33. Guichicovi. 34. La Princesa. +35. Santa María Chimalapa. 36. Matías Romero. 37. Santo Domingo Petapa. +38. El Barrio. 39. Palmar. 40. Chivela. 41. Santiago Chivela. 42. +Nizanda. 43. Agua Caliente. 44. Portillo Los Nanches. 45. Ixtepec. 46. +La Ventosa. 47. Zanatepec. 48. Unión Hidalgo. 49. Tres Cruces. 50. +Juchitán. 51. Escurano. 52. Salazar. 53. Santa Efigenia. 54. +Tequisistlán. 55. Cerro de Quiengola. 56. San Pablo. 57. Mixtequilla. +58. Tapanatepec. 59. Zarzamora. 60. Limón. 61. Tehuantepec. 62. +Bisilana. 63. Santa Lucía. 64. Cerro de Arenal. 65. Cerro de San Pedro. +66. La Concepción. 67. Tenango. 68. San Antonio. 69. Huilotepec. 70. +Salina Cruz." /></a> +<span class="caption"><span class="smcap">Fig. 1.</span> Map of the Isthmus of Tehuantepec based on the +American Geographical Society's "Map of Hispanic America on the Scale +of 1:1,000,000."<br /><small>(Click image for larger view.)</small><br /><br /> + +The localities shown are numbered in the gazetteer; the numerical +sequence of localities is an arrangement whereby north takes precedence +over south and west over east. 1. Alvarado. 2. Lerdo de Tejada. 3. +Tlacotalpan. 4. Tula. 5. Tecolapan. 6. Amatitlán. 7. Cosamaloapan. 8. +Chacaltianguis. 9. Novillero. 10. Ciudad Alemán. 11. Papaloapan. 12. +Tuxtepec. 13. Cuatotolapam. 14. Hueyapan. 15. Berta. 16. Coatzacoalcos. +17. Ayentes. 18. Río de las Playas. 19. Cosoleacaque. 20. Minatitlán. +21. Acayucan. 22. Aquilera. 23. San Lorenzo. 24. Naranja. 25. Suchil. +26. Jesús Carranza. 27. La Oaxaqueña. 28. Ubero. 29. Donají. 30. +Tolosita. 31. El Modelo. 32. Sarabia, 33. Guichicovi. 34. La Princesa. +35. Santa María Chimalapa. 36. Matías Romero. 37. Santo Domingo Petapa. +38. El Barrio. 39. Palmar. 40. Chivela. 41. Santiago Chivela. 42. +Nizanda. 43. Agua Caliente. 44. Portillo Los Nanches. 45. Ixtepec. 46. +La Ventosa. 47. Zanatepec. 48. Unión Hidalgo. 49. Tres Cruces. 50. +Juchitán. 51. Escurano. 52. Salazar. 53. Santa Efigenia. 54. +Tequisistlán. 55. Cerro de Quiengola. 56. San Pablo. 57. Mixtequilla. +58. Tapanatepec. 59. Zarzamora. 60. Limón. 61. Tehuantepec. 62. +Bisilana. 63. Santa Lucía. 64. Cerro de Arenal. 65. Cerro de San Pedro. +66. La Concepción. 67. Tenango. 68. San Antonio. 69. Huilotepec. 70. +Salina Cruz.</span> +</div> + +<hr class="tb" /><p><span class="pagenum"><a name="Page_27" id="Page_27">[Pg 27]</a></span></p> + +<div class="figcenter" style="width: 800px;"> +<a name="fig2" id="fig2"></a> +<img src="images/i_011.png" width="800" height="342" alt="Fig. 2. Topographic profile of the Isthmus of +Tehuantepec showing major localities along the Trans-isthmian Highway +and major types of vegetation. Vertical exaggeration approximately 165 +times."/> +<span class="caption"><span class="smcap">Fig. 2.</span> Topographic profile of the Isthmus of +Tehuantepec showing major localities along the Trans-isthmian Highway +and major types of vegetation. Vertical exaggeration approximately 165 +times.</span> +</div> +<p><span class="pagenum"><a name="Page_28" id="Page_28">[Pg 28]</a></span></p> + +<p>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.</p> + +<p>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.</p> + + +<h3><i>Climate</i></h3> + +<p>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 (<a href="#fig3">Fig. 3</a>). 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<span class="pagenum"><a name="Page_29" id="Page_29">[Pg 29]</a></span> +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.</p> + +<div class="figcenter" style="width: 600px;"> +<a name="fig3" id="fig3"></a> +<img src="images/i_013.png" width="600" height="314" alt="Fig. 3. Climatographs for Minatitlán, Veracruz, and +Salina Cruz, Oaxaca, based on data given by Contreras (1942). Plotted +points are for mean monthly temperatures and rainfall; months are +indicated by numbers."/> +<span class="caption"><span class="smcap">Fig. 3.</span> Climatographs for Minatitlán, Veracruz, and +Salina Cruz, Oaxaca, based on data given by Contreras (1942). Plotted +points are for mean monthly temperatures and rainfall; months are +indicated by numbers.</span> +</div> + + +<h3><i>Vegetation</i></h3> + +<p>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.<span class="pagenum"><a name="Page_30" id="Page_30">[Pg 30]</a></span> +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:</p> + +<h4>Gulf Lowlands</h4> + +<p>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.</p> + +<p>On the coastal dunes there are thickets of sea grape, patches of +<i>Cenchrus</i>, and clumps or scattered <i>Opuntia</i>. The lagoons are bordered +by mangrove thickets made up primarily of <i>Lonchocarpus hondurensis</i>. +In the marshes along the lower Río Coatzacoalcos and Río Papaloapan the +tall tough grass, <i>Gynerium sagittatum</i>, is common.</p> + +<p>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 (<i>op. cit.</i>: 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.</p> + +<p>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 (<a href="#pl1-fig1">Pl. 1, fig. 1</a>). According to Williams (<i>op. cit.</i>), the most +common trees in the savannas in southern Veracruz are <i>Ceiba +pentandra</i>, <i>Chlorophora tinctoria</i>, and <i>Byrsonima crassifolia</i>.</p> + +<p>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 (<a href="#pl1-fig2">Pl. 1, fig. 2</a>). Conspicuous trees in this scrub forest are +<i>Acacia cornigera</i>, <i>Bauhinia latifolia</i>, <i>Calliandra bijuga</i>, <i>Cassia +laevigata</i>, <i>Guazuma ulmifolia</i>, and various species of <i>Bursera</i>.</p> + +<p>The most extensive type of vegetation on the Gulf Coastal Plain is a +tall evergreen forest resembling tropical rainforest. Although<span class="pagenum"><a name="Page_31" id="Page_31">[Pg 31]</a></span> 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 (<a href="#pl2-fig1">Pl. 2, figs. 1</a>-<a href="#pl2-fig2">2</a>). 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.</p> + +<p>Among the abundant and dominant trees in this forest are <i>Swietenia +macrophylla</i>, <i>Calophyllum brasiliense</i>, <i>Achras zapota</i>, <i>Ceiba +pentandra</i>, <i>Castilla elastica</i>, <i>Cedrela mexicana</i>, <i>Tabebuia +Donnell-Smithi</i>, <i>Calocarpum mammosum</i>, <i>Bombax ellipticum</i>, and a +variety of <i>Ficus</i>. Epiphytes and Ilianas are abundant.</p> + +<h4>Central Ridges</h4> + +<p>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 <i>Ceiba pentandra</i>, <i>Cedrela +mexicana</i>, <i>Swietenia macrophylla</i>, and <i>Ficus</i> 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 +<i>Tabebuia Donnell-Smithi</i>, <i>Zanthoxylum melanostictum</i>, <i>Pithecolobium +arboreum</i>, and a species of <i>Pterocarpus</i>. 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 (<a href="#pl3-fig1">Pl. 3, fig.1</a>). +This type of forest extends from Mogoñe southward to about Matías +Romero.</p> + +<p>In the vicinity of Matías Romero open pine-oak forest (<i>Pinus caribaea</i> +and <i>Quercus</i> sp.) is found on some ridges as low as 250 meters above +sea level.</p> + +<p>On the Plains of Chivela in the southern part of the central region<span class="pagenum"><a name="Page_32" id="Page_32">[Pg 32]</a></span> +the vegetation takes on a semi-arid appearance, especially in a savanna +on the plains. Clumps of small trees and bushes, consisting of <i>Croton +nivea</i>, <i>Cordia cana</i>, <i>Jacquinia aurantiaca</i>, <i>Calycophyllum +candidissimum</i>, and <i>Cassia emarginata</i>, are scattered on a grassy +plain, from which rise widely-spaced palms of an unknown species (<a href="#pl3-fig2">Pl. 3, fig. 2</a>).</p> + +<h4>Pacific Coastal Plain</h4> + +<p>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 (<a href="#pl4-fig1">Pl. 4, figs. 1</a> and <a href="#pl4-fig2">2</a>). +Between Juchitán and La Ventosa few trees are more than two +meters high (<a href="#pl5-fig1">Pl. 5, fig. 1</a>). 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 <i>Acacia +cymbispina</i>, <i>Prosopis chilensis</i>, <i>Caesalpinia coriaria</i>, <i>Caesalpinia +eriostachys</i>, <i>Celtis iguanaea</i>, <i>Cordia brevispicata</i>, <i>Jatropha +aconitifolia</i>, and <i>Crescentia alata</i>.</p> + +<h4>Montane Vegetation</h4> + +<p>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 (<i>Abies religiosa</i>); 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.</p> + + +<h3><i>The Sierra de los Tuxtlas</i></h3> + +<p>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<span class="pagenum"><a name="Page_33" id="Page_33">[Pg 33]</a></span> 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.</p> + +<p>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.</p> + + + + +<h2><a name="GAZETTEER" id="GAZETTEER"></a>GAZETTEER</h2> + + +<p>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 +(<a href="#fig1">Fig. 1</a>).</p> + + +<h3><i>Oaxaca</i></h3> + +<div class="blockquot"><p>Agua Caliente.—Lat. 16° 38'; long. 94° 48'; elev. 140 m. A +hot spring, 6.9 km. north of La Ventosa on the +Trans-isthmian Highway; arid scrub forest [43].</p> + +<p>Arenal, Cerro de.—Lat. 16° 18'; long. 95° 32'; elev. 925 m. +(crest). A ridge northeast of Tenango; scrub forest on +slopes and pine-oak forest on top [64].</p> + +<p>Barrio, El.—Lat. 16° 38'; long. 95° 07'; elev. 314 m. A +village about 10 kilometers southwest of Matías Romero; +transition between scrub forest and broadleaf hardwood +forest [38].</p> + +<p>Bisilana.—Lat. 16° 20'; long. 95° 13'; elev. 35 m. A place +name for a former ranch at the edge of Tehuantepec; open +arid scrub forest [62].</p> + +<p>Chivela.—Lat. 16° 20'; long. 95° 01'; elev. 195 m. A +village on the Trans-isthmian Railroad, 26 kilometers by +rail south of Matías Romero and on the western edge of the +semi-arid Plains of Chivela [40].</p> + +<p>Concepción.—Lat. 16° 17'; long. 95° 29'; elev. 1200 m. A +ranch on the slopes of Cerro Arenal, east-northeast of +Tenango; dry pine-oak forest [66].</p><p><span class="pagenum"><a name="Page_34" id="Page_34">[Pg 34]</a></span></p> + +<p>Coyol.—Exact position unknown; according to Smith and +Taylor (1950: 10), Coyol is "between San Antonio and Las +Cruces."</p> + +<p>Donají.—Lat. 17° 13'; long. 95° 02'; elev. 90 m. A village +at Km. 155 on the Trans-isthmian Highway; rainforest [29].</p> + +<p>Escurano.—Lat. 16° 25'; long. 95° 27'; elev. 500 m. A ranch +about 25 kilometers west-northwest of Tehuantepec; arid +scrub forest [51].</p> + +<p>Guichicovi, San Juan.—Lat. 16° 58'; long. 95° 06'; elev. +250 m. A village on the north slopes of the isthmus, 12 +kilometers north-northwest of Matías Romero; cleared +hardwood forest and coffee plantations [33].</p> + +<p>Huilotepec.—Lat. 16° 14'; long. 95° 09'; elev. 30 m. A +small village on the Río Tehuantepec, 13 kilometers +south-southeast of Tehuantepec; open arid scrub forest [69].</p> + +<p>Ixtepec.—Lat. 16° 34'; long. 95° 06'; elev. 60 m. A town +and railroad junction on the northwestern edge of the Plains +of Tehuantepec; arid scrub forest [45].</p> + +<p>Juchitán.—Lat. 16° 26'; long. 95° 02'; elev. 15 m. A town +on the Plains of Tehuantepec, 22 kilometers by road +east-northeast of Tehuantepec; arid scrub forest [50].</p> + +<p>Limón.—Lat. 16° 20'; long. 95° 29'; elev. 600 m. A former +agrarian colony and now a small ranch about 27 kilometers +west of Tehuantepec; arid scrub forest [60].</p> + +<p>Matías Romero.—Lat. 16° 53'; long. 95° 02'; elev. 200 m. A +town on the Trans-isthmian Highway and railroad in the hills +near the crest of the isthmus; broadleaf hardwood forest and +open pine-oak forest [36].</p> + +<p>Mixtequilla.—Lat. 16° 24'; long. 95° 18'; elev. 40 m. A +village on the Río Tehuantepec, northwest of Tehuantepec; +dense scrub forest [57].</p> + +<p>Modelo, El.—Lat. 17° 07'; long. 94° 43'; elev. 200 m. An +old rubber plantation on the Río Chalchijapa, a tributary to +the Río Coatzacoalcos; rainforest [31].</p> + +<p>Nanches, Portillo Los.—Lat. 16° 35'; long. 95° 37'; elev. +500 m. A place name, about 4 kilometers southeast of +Totolapilla; scrub forest [44].</p> + +<p>Nizanda.—Lat. 16° 42'; long. 95° 02'; elev. 150 m. A +village on the Trans-isthmian Railroad between Chivela and +Ixtepec; dense scrub forest [42].</p> + +<p>Nueva Raza.—Exact location unknown; according to Thomas +MacDougall, this locality is in the lowlands of northern +Oaxaca; rainforest.</p> + +<p>Palmar.—Lat. 16° 43'; long. 94° 40'; elev. 300 m. A small +ranch on the west base of Cerro Atravesado; scrub forest +[39].</p> + +<p>Papaloapan.—Lat. 18° 11'; long. 96° 06'; elev. 25 m. A +small village on the Río Papaloapan in northern Oaxaca; low +evergreen forest and savanna [11].</p> + +<p>Princesa, La.—Lat. 16° 56'; long. 95° 02'; elev. 150 m. A +ranch on the northern slopes of the isthmus, 6 kilometers by +road north of Matías Romero; poorly developed rainforest +[34].</p> + +<p>Quiengola, Cerro de.—Lat. 16° 24'; long. 95° 22'; elev. 900 +m. (crest). A hill 15 kilometers west-northwest of +Tehuantepec; dense scrub forest on slopes and scattered +pines on top [55].</p> + +<p>Salazar.—Lat. 16° 25'; long. 95° 20'; elev. 45 m. A ranch +on the Río Tehuantepec, northwest of Tehuantepec; dense +scrub forest [52].</p> + +<p>Salina Cruz.—Lat. 16° 10'; long. 95° 12'; sea level. A port +on the Golfo de Tehuantepec; open arid scrub forest [70]. +Collections were made in the vicinity of the town and in the +open scrub forest 2.4 kilometers north at an elevation of 20 +meters.</p> + +<p>San Antonio.—Lat. 16° 15'; long. 95° 22'; elev. 40 m. A +ranch about 25 kilometers west-southwest of Tehuantepec; +arid scrub forest [68].</p> + +<p>San Pablo.—Lat. 16° 24'; long. 95° 18'; elev. 40 m. A ranch +on the Río Tehuantepec, northwest of Tehuantepec; dense +scrub forest [56]. Cerro San Pablo probably is the hill +north of this ranch; this is shown on some maps as Cerro de +los Amates.</p><p><span class="pagenum"><a name="Page_35" id="Page_35">[Pg 35]</a></span></p> + +<p>San Pedro, Cerro de.—Lat. 16° 18'; long. 95° 28'; elev. +about 1100 m. (crest). A ridge about 24 kilometers west of +Tehuantepec and east of Cerro Arenal; scrub forest on slopes +and pine-oak forest on top [65].</p> + +<p>Santa Efigenia.—Lat. 16° 25'; long. 94° 13'; elev. 500 m. A +ranch on the southern slopes of the Sierra Madre de Chiapas, +8 kilometers north-northwest of Tapanatepec; scrub forest. +Former home of Francis Sumichrast [53].</p> + +<p>Santa Lucía.—Lat. 16° 18'; long. 95° 28'; elev. 800 m. A +place name for a former ranch on the east slopes of Cerro +Arenal; scrub forest [63].</p> + +<p>Santa María Chimalapa.—Lat. 16° 55'; long. 94° 42'; elev. +296 m. A village on the Río de los Milagros, a tributary to +the Río Coatzacoalcos; rainforest [35].</p> + +<p>Santiago Chivela.—Lat. 16° 42'; long. 94° 53'; elev. 200 m. +A village on the Trans-isthmian Highway, 13.4 kilometers by +road south of Matías Romero; dry, grassy plains and +scattered clumps of scrubby trees and palms [41]. +Collections were made in the vicinity of the village and at +a rocky stream, 11 kilometers south on the Trans-isthmian +Highway at an elevation of 230 m.</p> + +<p>Santo Domingo (Petapa).—Lat. 16° 50'; long. 95° 08'; elev. +225 m. A village about 13 kilometers west-southwest of +Matías Romero; semi-arid scrub forest [37].</p> + +<p>Sarabia.—Lat. 17° 04'; long. 95° 02'; elev. 100 m. A +village 25 kilometers north of Matías Romero on the +Trans-isthmian Highway; rainforest [32]. Collections were +made in the vicinity of the village and in the rainforest +along the Río Sarabia, 5 kilometers north of the village at +an elevation of 80 meters.</p> + +<p>Tapanatepec.—Lat. 16° 32'; long. 94° 12'; elev. 90 m. A +town on the Pan-American Highway on the lower slopes of the +Sierra Madre de Chiapas; dense scrub forest [58].</p> + +<p>Tehuantepec.—Lat. 16° 20'; long. 95° 14'; elev. 35 m. A +large town on the Plains of Tehuantepec; scrub forest [61]. +Collections were made in the vicinity of the town and in the +dense scrub forest 8.6 kilometers west at an elevation of 85 +meters and 14 kilometers west at an elevation of 120 meters.</p> + +<p>Tenango.—Lat. 16° 16'; long. 95° 30'; elev. 1100 m. A town +in the mountains about 40 kilometers west-southwest of +Tehuantepec; scrub forest [67].</p> + +<p>Tequisistlán.—Lat. 16° 24'; long. 95° 37'; elev. 190 m. A +village in the valley of the Río Tequisistlán, a tributary +to the Río Tehuantepec; dense scrub forest [54]. Most +collections were made about one kilometer north of the +village where the Pan-American Highway crosses the Río +Tequisistlán.</p> + +<p>Tolosita.—Lat. 17° 12'; long. 95° 03'; elev. 80 m. A +village on the Río Tortuguero near the Trans-isthmian +Highway; rainforest [30].</p> + +<p>Tres Cruces.—Lat. 16° 26'; long. 95° 51'; elev. 750 m. A +ranch near the Pan-American Highway, 70 kilometers by road +west-northwest of Tehuantepec; dense scrub forest [49].</p> + +<p>Tuxtepec—Lat. 18° 06'; long. 96° 05'; elev. 80 m. A town on +the Río Papaloapan in northern Oaxaca; low evergreen forest +[12].</p> + +<p>Ubero.—Lat. 17° 18'; long. 95° 00'; elev. 80 m. A lumber +camp and railroad station, 8.5 kilometers south of the Río +Jaltepec on the Trans-isthmian Highway; rainforest [28].</p> + +<p>Unión Hidalgo.—Lat. 16° 27'; long. 94° 48'; elev. 7 m. A +village on the railroad, 20 kilometers east-northeast of +Juchitán; open scrub forest [48].</p> + +<p>Ventosa, La.—Lat. 16° 30'; long. 94° 51'; elev. 25 m. A +village at the junction of the Pan-American and +Trans-isthmian highways; open scrub forest [46].</p> + +<p>Zanatepec.—Lat. 16° 28'; long. 94° 22'; elev. 80 m. A +village on the Pan-American Highway at the eastern edge of +the Plains of Tehuantepec; dense scrub forest [47]. Most +collections were made in the scrub forest 5 to 8 kilometers +west-northwest of the village.</p> + +<p>Zarzamora.—Lat. 16° 21'; long. 95° 48'; elev. 800 m. A +ranch between La Reforma (16 kilometers west of +Tequisistlán) and Santa María Ecatepec; scrub forest with +oaks on higher ridges [59].</p></div><p><span class="pagenum"><a name="Page_36" id="Page_36">[Pg 36]</a></span></p> + + +<h3><i>Veracruz</i></h3> + +<div class="blockquot"><p>Acayucan.—Lat. 17° 57'; long. 94° 55'; elev. 160 m. A large +town on the Trans-isthmian Highway; rainforest [21]. +Collections were made in the vicinity of the town, but +principally at Rancho Las Hojitas, 7 kilometers northwest of +town at an elevation of 150 meters.</p> + +<p>Alvarado.—Lat. 18° 47'; long. 95° 47'; sea level. A fishing +village at the mouth of the Río Papaloapan; coastal dunes +and marshes [1]. Most collections were made 1-3 kilometers +southeast of the village in marshes on the leeward side of +the coastal dunes.</p> + +<p>Amatitlán.—Lat. 18° 26'; long. 95° 45'; elev. 4 m. A +village on the bank of the Río Papaloapan; savanna and sugar +plantations [6].</p> + +<p>Aquilera.—Lat. 17° 48'; long. 95° 01'; elev. 150 m. A +village 21 kilometers southwest of Acayucan on the +Trans-isthmian Highway; rainforest [22].</p> + +<p>Ayentes.—Lat. 18° 10'; long. 94° 26'; elev. 2 m. A railroad +station on the east bank of the Río Coatzacoalcos, across +the river from the city of Coatzacoalcos; scrub forest and +marshes [17].</p> + +<p>Berta.—Lat. 18° 07'; long. 94° 27'; elev. 5 m. A ranch just +south of Coatzacoalcos; scrub and low evergreen forest [15].</p> + +<p>Chacaltianguis.—Lat. 18° 18'; long. 95° 52'; elev. 5 m. A +village on the Río Papaloapan; savanna [8].</p> + +<p>Ciudad Alemán.—Lat. 18° 13'; long. 96° 07'; elev. 30 m. A +new government town, headquarters of the Comisión del +Papaloapan; scrub and low evergreen forest [10].</p> + +<p>Coatzacoalcos (formerly Puerto México).—Lat. 18° 10'; long. +94° 27'; elev. 2 m. A seaport at the mouth of the Río +Coatzacoalcos; scrub on coastal dunes; marshes and low +evergreen forest inland [16]. Most collections are from the +forest-savanna ecotone, 8 kilometers southwest of town.</p> + +<p>Cosamaloapan.—Lat. 18° 22'; long. 95° 50'; elev. 4 m. An +agricultural town on the Río Papaloapan; savanna and sugar +plantations [7].</p> + +<p>Cosoleacaque.—Lat. 17° 59'; long. 94° 38'; elev. 55 m. A +village 8 kilometers by road west of Minatitlán; savanna +[19].</p> + +<p>Cuatotolapam.—Lat. 18° 08'; long. 95° 16'; elev. 13 m. A +village on the Trans-isthmian Railroad; savanna and low +evergreen forest along streams [13].</p> + +<p>Hueyapan.—Lat. 18° 08'; long. 19° 09'; elev. 85 m. A town +32 kilometers by road northwest of Acayucan; savanna and low +evergreen forest [14]. Collections were made in the vicinity +of the town and from forest 10 kilometers southeast of town +at an elevation of 135 meters.</p> + +<p>Jesús Carranza (formerly Santa Lucrecia).—Lat. 17° 27'; +long. 95° 02'; elev. 80 m. A town and railroad junction in +the middle of the isthmus; rainforest [26]. Most of +Dalquest's specimens came from varying distances from Jesús +Carranza along the Río Coatzacoalcos and its tributaries.</p> + +<p>Minatitlán.—Lat. 17° 58'; long. 94° 32'; elev. 15 m. An oil +refinery center on the Río Coatzacoalcos; savanna [20].</p> + +<p>Naranjo.—Lat. 17° 35'; long. 95° 07'; elev. 100 m. A +village on the Trans-isthmian Highway, 45 kilometers south +of Acayucan; rainforest and palm forest [24].</p> + +<p>Novillero.—Lat. 18° 16'; long. 95° 59'; elev. 10 m. A +village on the Río Papaloapan; scrub forest and grassland +[9].</p> + +<p>Oaxaqueña, La.—Lat. 17° 26'; long. 94° 53'; elev. 80 m. A +hacienda on the Río Coatzacoalcos about 12 kilometers east +of Jesús Carranza; rainforest [27].</p> + +<p>Playas, Río de las.—Lat. 18° 08'; long. 94° 07'; elev. 3 m. +The river (sometimes known as the Río Tonolá) forming the +boundary between the states of Veracruz and Tabasco; +rainforest [18].</p> + +<p>San Lorenzo.—Lat. 17° 44'; long. 94° 42'; elev. 25 m. A +village on the Río Chiquito, about 30 kilometers southeast +of Acayucan; rainforest [23].</p><p><span class="pagenum"><a name="Page_37" id="Page_37">[Pg 37]</a></span></p> + +<p>Suchil.—Lat. 17° 31'; long. 95° 03'; elev. 40 m. A village +on the Trans-isthmian Railroad, about 10 kilometers north of +Jesús Carranza; rainforest [25].</p> + +<p>Tecolapan.—Lat. 18° 24'; long. 95° 18'; elev. 275 m. A +village on a small river of the same name in the western +foothills of Los Tuxtlas; rainforest [5].</p> + +<p>Tejada, Lerdo de.—Lat. 18° 37'; long. 95° 31'; elev. 60 m. +An agricultural village, 35 kilometers by road +east-southeast of Alvarado; scrub forest, marshes, and sugar +plantations [2]. Collections were made in a marsh, 5 +kilometers west-northwest of the village.</p> + +<p>Tlacotalpan.—Lat. 18° 37'; long. 95° 42'; elev. 3 m. A town +at the confluence of the Río San Juan and Río Papaloapan; +marshes and sugar plantations [3].</p> + +<p>Tula.—Lat. 18° 36'; long. 95° 22'; elev. 150 m. A village +near the western base of Los Tuxtlas; low evergreen forest +and marshes [4]. Collections were made in a marsh 3 +kilometers northwest of the village.</p></div> + + + + +<h2><a name="THE_AMPHIBIAN_FAUNA_OF_THE_LOWLANDS" id="THE_AMPHIBIAN_FAUNA_OF_THE_LOWLANDS"></a>THE AMPHIBIAN FAUNA OF THE LOWLANDS</h2> + + +<p>In presenting an account of the amphibian fauna of the lowlands of the +Isthmus of Tehuantepec three items must be considered:</p> + +<p><span style="margin-left: 2.5em;">1. The composition of the fauna.</span><br /> + +<span style="margin-left: 2.5em;">2. The ecology of the fauna.</span><br /> + +<span style="margin-left: 2.5em;">3. The distribution of the fauna.</span><br /></p> + +<p>These items, together with similar data concerning the amphibians of +the adjacent highlands, will form the basis for the subsequent +discussion of the establishment of present patterns of distribution in +the isthmian region.</p> + + +<h3><i>Composition of the Fauna</i></h3> + +<p>The amphibian fauna of the lowlands of the Isthmus of Tehuantepec +consists of 36 species definitely recorded from the area. These include +one genus and species of caecilian, one genus, including three species +of salamanders, and 14 genera and 32 species of anurans.</p> + +<p>In comparison with the known amphibian fauna of the forested and +savanna portions of El Petén, Guatemala (Stuart, 1935 and 1958), we +find that there are more species recorded from the isthmus than from El +Petén. Stuart found only 20 species of amphibians in both forest and +savanna habitats in El Petén. Of the 36 species of amphibians known +from the isthmus, 28 occur on the Gulf lowlands and live in forest or +savanna habitats.</p> + +<p>The geographic position of the isthmus with regard to major faunal +areas in Middle America, and the diversity of the environment are +important factors in understanding the presence of a large number of +species of amphibians in the isthmus. The large number of species +probably is a reflection of the diversity of the environment; this +diversity is the result of fluctuation of climate, and thus<span class="pagenum"><a name="Page_38" id="Page_38">[Pg 38]</a></span> +environments, in the not too distant past. In no individual habitat, +such as rainforest, savanna, or scrub forest, does the number of +species approach the total for the region.</p> + + +<h3><i>Ecology of the Fauna</i></h3> + +<p>In the preceding section on the description of the Isthmus of +Tehuantepec I have outlined the major environments in the region. With +respect to the distribution of amphibians we may recognize three major +environments in the isthmus—rainforest, semi-arid scrub forest, and +savanna. Each of these has varying combinations of physical and biotic +factors that are important in the ecology of amphibians. Because of the +importance of moisture, not only for the maintenance of life in these +animals, but in most species their dependence on water for breeding +purposes, this environmental factor is considered the most significant +in the ecological distribution of amphibians. A second factor is the +availability of necessary shelter, especially aestivation sites. These +factors will be compared in the three major environments in the region.</p> + +<p>Moisture is present in the environment in the form of free water or +atmospheric moisture. With respect to the latter, it is well known that +dense shaded forests have a considerably higher relative humidity than +do open plains or areas with only scattered trees. Thus, the +rainforests of the isthmus are characterized by a much higher relative +humidity than are the savannas or semi-arid scrub forests. Although +with regard to rainfall there is a pronounced dry season in the regions +supporting rainforest, there still remains considerable atmospheric +moisture in this environment throughout the year. The dense foliage +provides shade and protection from desiccating effects of wind and +sunlight; furthermore the foliage contributes moisture by +transpiration. The deep alluvial soils mixed with large quantities of +organic matter (decaying leaves and rotting logs) maintain considerable +quantities of moisture.</p> + +<p>Conversely, the savannas and scrub forests have little atmospheric +moisture during the dry season. In the former habitat there are few +trees to provide shade or moisture through transpiration; in the latter +most of the trees lose their leaves during the dry season. Thus, these +environments are desiccated by the dry winds and direct sunlight. +Furthermore, the soils in these environments become dry and caked. +There is little or no terrestrial matter to hold moisture.</p> + +<p>Free water in these environments is present in a variety of forms<span class="pagenum"><a name="Page_39" id="Page_39">[Pg 39]</a></span> at +different times of the year. During the dry season the more extensive +marshes in the savannas persist; many ponds and most of the streams in +the rainforest are permanent throughout the year. In the scrub forest +all except the largest streams become dry during the dry season, and no +ponds exist through the dry season. With the advent of the first heavy +summer rains the stream beds fill with water, marshes expand, and many +depressions become ponds (<a href="#pl5-fig2">Pl. 5, fig. 2</a>). At this time the amount of +free water in the scrub forests and savannas greatly increases, much +more so than that in the rainforests.</p> + +<p>Environments are vertically stratified in the rainforests. There is the +deep alluvial soil, the ground litter of leaves and decaying logs, the +low bushes and small trees, and finally the tall trees of the forest. +Each of these provides certain types of shelter for amphibians. The +moist soil and litter on the forest floor is an important microhabitat +for fossorial and strictly terrestrial species. The dense foliage of +the trees, tree holes, and bromeliads growing on the trees provide +shelter for arboreal species. Arboreal and terrestrial bromeliads and +the terrestrial elephant-ear plants (<i>Xanthosoma</i>) contain water in the +axils of their leaves throughout the year and thus provide an important +habitat for amphibians. The low, spiny, deciduous trees of the scrub +forest and the grasses and scattered trees in the savannas provide +little shelter. In the savannas there are depressions, some of which +contain water throughout the year; these are often surrounded by trees +providing refugia for amphibians during the dry season. In the scrub +forest many species congregate along streams and in moist stream beds +during the dry season.</p> + +<p>Now that the important ecological factors of the major environments +have been outlined, we may examine the local distribution of amphibians +in each of these. Beginning with the rainforest, we find only one +fossorial species, <i>Gymnopis mexicanus</i>. A large number of species are +found on the forest floor; characteristic inhabitants of the leaf +litter are: <i>Bufo valliceps</i>, <i>Eleutherodactylus rhodopis</i>, +<i>Microbatrachylus pygmaeus</i>, and <i>Syrrhophus leprus</i>. Other terrestrial +amphibians usually are not scattered throughout the rainforest, as are +those named immediately above, but instead inhabit areas of forest +adjacent to ponds or streams; these species include: <i>Bufo marinus</i>, +<i>Eleutherodactylus natator</i>, <i>Eleutherodactylus rugulosus</i>, +<i>Leptodactylus labialis</i>, <i>Leptodactylus melanonotus</i>, <i>Rana palmipes</i> +and <i>Rana pipiens</i>. The most striking ecological assemblage<span class="pagenum"><a name="Page_40" id="Page_40">[Pg 40]</a></span> of +amphibians in the rainforest is the arboreal group of species, +including:</p> + +<p> +<span style="margin-left: 2.5em;"><i>Bolitoglossa occidentalis</i></span><br /> +<span style="margin-left: 2.5em;"><i>Bolitoglossa platydactyla</i></span><br /> +<span style="margin-left: 2.5em;"><i>Eleutherodactylus alfredi</i></span><br /> +<span style="margin-left: 2.5em;"><i>Hyla baudini</i></span><br /> +<span style="margin-left: 2.5em;"><i>Hyla ebraccata</i></span><br /> +<span style="margin-left: 2.5em;"><i>Hyla loquax</i></span><br /> +<span style="margin-left: 2.5em;"><i>Hyla microcephala martini</i></span><br /> +<span style="margin-left: 2.5em;"><i>Hyla picta</i></span><br /> +<span style="margin-left: 2.5em;"><i>Phrynohyas modesta</i></span><br /> +<span style="margin-left: 2.5em;"><i>Phrynohyas spilomma</i></span><br /> +<span style="margin-left: 2.5em;"><i>Phyllomedusa callidryas taylori</i></span><br /> +</p> + +<p>In the savannas <i>Rhinophrynus dorsalis</i>, <i>Engystomops pustulosus</i>, and +<i>Gastrophryne usta</i> are fossorial species. <i>Bufo marinus</i>, +<i>Leptodactylus melanonotus</i>, <i>Leptodactylus labialis</i>, <i>Rana palmipes</i>, +and <i>Rana pipiens</i> are found in the vicinity of permanent water in the +savannas. Although the savanna habitat does not provide the ecological +conditions for the existence of an arboreal fauna, many arboreal +species from the surrounding rainforest utilize the extensive marshes +and ponds in the savannas for breeding purposes. Thus, <i>Hyla baudini</i>, +<i>Hyla microcephala martini</i>, <i>Hyla picta</i>, and <i>Phrynohyas spilomma</i> +have been found breeding in savannas. In parts of savannas where clumps +of trees surround depressions containing water throughout the year, +individuals of the species named above, together with <i>Hyla loquax</i> and +<i>Phyllomedusa callidryas taylori</i>, may not only breed, but remain +throughout the year.</p> + +<p>In the semi-arid scrub forest the same fossorial species as exist in +the savannas are found. Likewise, <i>Bufo marinus</i>, <i>Leptodactylus +labialis</i>, <i>Leptodactylus melanonotus</i>, and <i>Rana pipiens</i> are found +near permanent water. Terrestrial species in this semi-arid environment +include <i>Bufo canaliferus</i>, <i>Bufo coccifer</i>, <i>Bufo marmoreus</i>, +<i>Syrrhophus pipilans</i>, and <i>Diaglena reticulata</i>. Of these, <i>Syrrhophus +pipilans</i> sometimes inhabits low trees and bushes; the others may be +fossorial. The arboreal species in the scrub forest include <i>Hyla +baudini</i>, <i>Hyla robertmertensi</i>, <i>Hyla staufferi</i>, and <i>Phyllomedusa +dacnicolor</i>.</p> + +<p><i>Eleutherodactylus rugulosus</i> and <i>Hylella sumichrasti</i> live along +streams in the scrub forest. <i>Hylella sumichrasti</i> lays its eggs in +these streams.</p> + +<p>In comparing the ecological differences in the amphibian assemblages in +the three major habitats, the most obvious difference is the great +percentage of arboreal species in the rainforest as compared with +savanna and scrub forest. Only four arboreal species are found in the +scrub forest, none in the savannas, but eleven in the rainforest. +Likewise, there is an absence of ground-dwelling forms in the arid +habitats; in the latter the only terrestrial<span class="pagenum"><a name="Page_41" id="Page_41">[Pg 41]</a></span> species are those that +are found near water. A possible exception is <i>Syrrhophus pipilans</i>.</p> + +<p>From the above analysis of ecological distribution we may see that the +rainforest provides a variety of habitats for amphibians and that these +habitats are suitable for amphibian life throughout the year. On the +other hand, the savannas and scrub forests are characterized by extreme +conditions of desiccation, a factor of considerable importance in +limiting the ecological distribution of amphibians. However, there +still is a diversity of amphibians in these semi-arid environments. +Obviously, these species are adapted in various ways for survival +during the dry season, at which time environmental conditions are such +that the animals cannot carry on their normal activities.</p> + +<p>Although there is not an abundance of data concerning the seasonal +activity of the fauna, what is available shows some interesting +correlations with the environments. During the dry season in the scrub +forest there is essentially no amphibian activity; an occasional <i>Rana +pipiens</i> may be seen along a river, or a <i>Bufo marinus</i> may be seen at +night. In the rainforest the terrestrial-breeding amphibians are active +during the dry season. <i>Eleutherodactylus rugulosus</i> is found at night +or by day along streams. <i>Eleutherodactylus rhodopis</i>, +<i>Microbatrachylus pygmaeus</i>, and <i>Bufo valliceps</i> are active during the +day; these plus <i>Bolitoglossa occidentalis</i>, <i>Bolitoglossa +platydactyla</i>, <i>Eleutherodactylus alfredi</i>, <i>Eleutherodactylus +natator</i>, and an occasional <i>Hyla</i> are active at night.</p> + +<p>With the onset of the heavy summer rains and the subsequent formation +of breeding ponds, amphibian activity reaches a peak. This is +especially noticeable in the semi-arid environments, where during the +dry season there is little activity.</p> + +<p>Among the anurans in the isthmus the four species of +<i>Eleutherodactylus</i>, the two species of <i>Syrrhophus</i>, and the one +species of <i>Microbatrachylus</i> are either known, or presumed, to lay +eggs on the ground; these develop directly into small frogs. All of the +other anurans deposit their eggs in water or attach them to objects +over water (<i>Phyllomedusa</i>); these hatch into tadpoles, which later +metamorphose into frogs. <i>Hylella sumichrasti</i> is known to breed only +in streams. All of the other species breed in ponds, but at times some +species deposit their eggs in streams; in this last group are <i>Bufo +valliceps</i>, <i>Bufo marmoreus</i>, <i>Phyllomedusa callidryas taylori</i>, and +<i>Rana pipiens</i>.</p> + +<p>Although the ecological data are incomplete, they do show that<span class="pagenum"><a name="Page_42" id="Page_42">[Pg 42]</a></span> +ecological conditions differ greatly in the three major environments, +different species of amphibians inhabit these environments, and that +the fauna is ecologically diversified in each environment.</p> + + +<h3><i>Distribution of the Fauna</i></h3> + +<p>Plotting the distributions of species of amphibians known to live in +the lowlands of the Isthmus of Tehuantepec results in an array of +geographic patterns. These may be analyzed with respect to those +species that are restricted either to the Gulf lowlands or the Pacific +lowlands, or those that occur on both the Gulf and Pacific lowlands. +Furthermore, the distributions may be analyzed with respect to those +species whose ranges extend from México across the Isthmus of +Tehuantepec into Central America, those that reach the isthmus from +Central America but do not extend into México proper, and those that +reach the isthmus from México but do not extend into Central America. +It should be kept in mind that the following analysis is of the lowland +inhabitants only. Species inhabiting the foothills and mountains will +be discussed later.</p> + +<p>1. <span class="smcap">Species Restricted to the Gulf Lowlands.</span> Of the 36 species of +amphibians recorded from the Isthmus of Tehuantepec, nine (25 per cent) +are in this group. Four of these (<i>Eleutherodactylus alfredi</i>, +<i>Syrrhophus leprus</i>, <i>Hyla loquax</i>, and <i>Hyla picta</i>) live in the Gulf +lowlands to the east and to the west of the isthmus. Three others +(<i>Hyla ebraccata</i>, <i>Hyla microcephala martini</i> and <i>Phyllomedusa +callidryas taylori</i>) are primarily Central American in their +distribution and reach the northwestern limits of their ranges in the +Gulf lowlands of the isthmus, whereas <i>Bolitoglossa platydactyla</i> and +<i>Eleutherodactylus natator</i> reach the southern limits of their +distributions in the isthmus.</p> + +<p>2. <span class="smcap">Species Restricted to the Pacific Lowlands.</span> This group includes six +species, or 17 per cent of the amphibian fauna of the isthmus. Two of +these (<i>Bufo coccifer</i> and <i>Syrrhophus pipilans</i>) range to the east and +to the west of the isthmus on the Pacific lowlands. Two others (<i>Bufo +canaliferus</i> and <i>Hyla robertmertensi</i>) range from the isthmus into +Central America, and <i>Diaglena reticulata</i> and <i>Phyllomedusa +dacnicolor</i> range on the Pacific lowlands of México southeastward to +the isthmus.</p> + +<p>3. <span class="smcap">Species That Occur on the Pacific and Gulf Lowlands.</span> This group +includes 19 species, or 53 per cent of the total amphibian fauna. Of +these, nine species (25 per cent of the entire amphibian<span class="pagenum"><a name="Page_43" id="Page_43">[Pg 43]</a></span> fauna) are +widespread throughout the lowlands of México and Central America; these +are:</p> + +<p> +<span style="margin-left: 2.5em;"><i>Gymnopis mexicanus</i></span><br /> +<span style="margin-left: 2.5em;"><i>Rhinophrynus dorsalis</i></span><br /> +<span style="margin-left: 2.5em;"><i>Bufo marinus</i></span><br /> +<span style="margin-left: 2.5em;"><i>Engystomops pustulosus</i></span><br /> +<span style="margin-left: 2.5em;"><i>Leptodactylus labialis</i></span><br /> +<span style="margin-left: 2.5em;"><i>Leptodactylus melanonotus</i></span><br /> +<span style="margin-left: 2.5em;"><i>Hyla baudini</i></span><br /> +<span style="margin-left: 2.5em;"><i>Hyla staufferi</i></span><br /> +<span style="margin-left: 2.5em;"><i>Rana pipiens</i></span><br /> +</p> + +<p>Four species occur on the Gulf lowlands to the east and to the west of +the isthmus, but on the Pacific lowlands they occur only to the east; +this group includes <i>Bufo valliceps</i>, <i>Eleutherodactylus rhodopis</i>, +<i>Phrynohyas modesta</i>, and <i>Phrynohyas spilomma</i>. Three species live to +the east and to the west of the isthmus on the Pacific lowlands, but +only to the west on the Gulf lowlands; these include <i>Eleutherodactylus +rugulosus</i>, <i>Microbatrachylus pygmaeus</i>, and <i>Gastrophryne usta</i>.</p> + +<p>Six species that cross the isthmus live on the humid Gulf lowlands and +on the humid lowlands of Chiapas and Guatemala, but not on the +semi-arid Plains of Tehuantepec; these include <i>Bolitoglossa +occidentalis</i>, <i>Eleutherodactylus rhodopis</i>, <i>Microbatrachylus +pygmaeus</i>, <i>Phrynohyas modesta</i>, <i>Phrynohyas spilomma</i>, and <i>Rana +palmipes</i>. Of these, <i>Microbatrachylus pygmaeus</i> also occurs in +scattered humid environments to the west of the isthmus on the Pacific +lowlands.</p> + +<p>Two species are endemic to the isthmian region. <i>Bolitoglossa +veracrucis</i> is known only from the humid northern slopes of the +isthmus. <i>Hylella sumichrasti</i> occurs on the Pacific slopes of the +isthmus and extends to the east into western Chiapas.</p> + +<p>In analyzing the distribution of the amphibians with respect to those +that are restricted to either the Pacific or Gulf lowlands or those +that cross the continental divide in the isthmus, we find that 25 per +cent of the species are restricted to the Gulf lowlands, 17 per cent +are restricted to the Pacific lowlands, and 53 per cent cross the +isthmus. In analyzing the distribution patterns with respect to those +that extend across the isthmus of Tehuantepec from east to west, we +find that 14 per cent of the species do not extend east of the isthmus +into Central America and that 19 per cent do not range west of the +isthmus into México proper; 61 per cent of the species range to the +east and to the west of the isthmus. Of the 36 species of amphibians +inhabiting the isthmus only nine species (25 per cent) range across the +isthmus, that is, occur on the Gulf and Pacific lowlands, and also +range to the east and to the west of the isthmus. To these wide-ranging +species the diversified environments of the<span class="pagenum"><a name="Page_44" id="Page_44">[Pg 44]</a></span> isthmus do not present a +barrier to distribution. The other 27 species (75 per cent) either do +not cross the isthmus from east to west or from north to south; thus, +probably in one way or another the isthmus presents a barrier to their +distribution.</p> + + + + +<h2><a name="THE_AMPHIBIAN_FAUNA_OF_THE_FOOTHILLS_AND_ADJACENT_HIGHLANDS" id="THE_AMPHIBIAN_FAUNA_OF_THE_FOOTHILLS_AND_ADJACENT_HIGHLANDS"></a>THE AMPHIBIAN FAUNA OF THE FOOTHILLS AND ADJACENT HIGHLANDS</h2> + + +<p>To amphibians inhabiting the foothills and mountains of southern México +and northern Central America, the isthmus presents a great barrier to +dispersal. For example, salamanders of the genus <i>Thorius</i>, the +<i>mexicanus</i> and <i>augusti</i> groups of the genus <i>Eleutherodactylus</i>, the +<i>bistincta</i> group of the genus <i>Hyla</i>, and the genus <i>Tomodactylus</i> +occur on the Mexican Plateau and southward into the mountains of +Oaxaca. Nevertheless, no members of these groups are present in the +Guatemalan-Chiapan Highlands. The genera <i>Chiropterotriton</i>, +<i>Magnadigita</i>, <i>Pseudoeurycea</i>, and <i>Ptychohyla</i>, as well as the +<i>eximia</i> group of <i>Hyla</i> are represented by different species in the +Guatemalan-Chiapan Highlands than in the mountains of México on the +other side of the isthmus. Several species of <i>Plectrohyla</i> occur in +the Guatemalan-Chiapan Highlands, but none is known from the Mexican +Highlands, although one species occurs in the Tuxtlas.</p> + +<p>Living in the humid forests of the foothills are salamanders of the +genus <i>Lineatriton</i>, frogs of the <i>spatulatus</i> group of +<i>Eleutherodactylus</i>, <i>Anotheca coronata</i>, <i>Hyla miotympanum</i>, and +<i>Phyllomedusa moreleti</i>. All of these occur in the foothills of the +Sierra Madre Oriental in eastern México and in Los Tuxtlas. +<i>Lineatriton</i>, <i>Hyla miotympanum</i>, and the <i>spatulatus</i> group of +<i>Eleutherodactylus</i> do not occur in the foothills of the +Guatemalan-Chiapan Highlands; those amphibians reach the end of their +ranges at the isthmus. <i>Phyllomedusa moreleti</i> and <i>Anotheca coronata</i> +are found in the northern foothills of the Guatemalan-Chiapan +Highlands, and <i>Phyllomedusa moreleti</i> is found in the foothills on the +Pacific slopes of the Chiapan Highlands.</p> + +<p>Although the above analysis is not so detailed as that of the lowland +inhabitants, it does show that all of the genera and species of +amphibians known to inhabit the foothills and highlands adjacent to the +isthmus, only two species of amphibians cross the isthmus from one +highland mass to the other. Thus, it is evident that the Isthmus of +Tehuantepec presents a great barrier to dispersal of these groups of +amphibians.</p> + + + +<p><span class="pagenum"><a name="Page_45" id="Page_45">[Pg 45]</a></span></p> +<h2><a name="ESTABLISHMENT_OF_PRESENT_PATTERNS_OF_DISTRIBUTION" id="ESTABLISHMENT_OF_PRESENT_PATTERNS_OF_DISTRIBUTION"></a>ESTABLISHMENT OF PRESENT PATTERNS OF DISTRIBUTION</h2> + + +<p>From the foregoing analysis of geographical and ecological distribution +in the Isthmus of Tehuantepec we may strive for an interpretation of +the events that led to the establishment of patterns of distribution +displayed not only by the amphibians, but other terrestrial vertebrates +as well. The thesis that I am proposing below is based on the premise +that in southern México and northern Central America climatic +fluctuation during the Pleistocene was of sufficient magnitude to cause +vegetational shifts, both vertically and latitudinally, resulting in +the establishment of alternating continuous and discontinuous lowland +and highland environments, although this climatic fluctuation was not +so great as to eliminate tropical lowland environments from the region. +I feel that the present patterns of distribution of the amphibians in +the Isthmus of Tehuantepec may be explained on this premise.</p> + +<p>Many authors dealing with the herpetofauna of Middle America have +followed Schuchert's (1935) suggestion of a seaway in the isthmus +during the Cenozoic. Thus, Burt (1931), Duellman (1956, 1958a), Gloyd +(1940), Oliver (1948), Smith and Laufe (1946), and Stuart (1941) +employed the presence of a seaway to explain distribution and +speciation in various genera. Durham, Arellano, and Peck (1952), Olson +and McGrew (1941), and Stirton (1954) have provided geological evidence +that there probably was no Cenozoic seaway in the Isthmus of +Tehuantepec. Even if there were a seaway in the Pliocene or Miocene +(the dating of this possible seaway is open to question), its presence +is not necessary to explain the present patterns of distribution in the +isthmus.</p> + +<p>In recent years the study of natural biotic environments, palynology, +and Pleistocene chronology in Middle America has produced a wealth of +data, which although still fragmentary begins to form a picture of past +climatic events in that part of the world. Sedimentary studies by +Hutchinson, Patrick, and Deevey (1956) and Sears, Foreman, and Clisby +(1955) have provided evidence of drastic climatic shifts in México +during the Pleistocene. Further evidence of bioclimatic fluctuation is +provided by Martin and Harrell (1957) and Martin (1958); the latter has +suggested that there was a displacement of the tropical zones in +southern México and northern Central America by as much as 3000 feet +during the glacial maximum. Much of the evidence of such drastic +vertical<span class="pagenum"><a name="Page_46" id="Page_46">[Pg 46]</a></span> shifts in environments is based on the presence of +Pleistocene montane glaciers on Mexican volcanoes (White, 1956) and +Chirripo in Costa Rica (Weyl, 1955). Dorf (1959) supports this idea of +drastic climatic change.</p> + +<p>In his studies of the avifauna of México and Guatemala Griscom (1932 +and 1950) made an important issue of the continuity of the bird fauna +in what he called the Subtropical Life-zone, which essentially consists +of cloud forest, a widespread, but discontinuous, habitat on the Gulf +(windward) slopes of the Mexican and Central American highlands at +elevations between 1000 and 2000 meters. To account for this apparent +uniformity in the avifauna Griscom hypothesized a continuity of cloud +forest environment in the Pleistocene; this would result in the +depression of cloud forests to the coastal lowlands and the +displacement of tropical lowland environments far to the south in +Central America. Stuart (1951) objected to this displacement of lowland +tropical rainforest; he stated that a descent to sea level of a +subtropical zone would have brought about either widespread +extermination of the tropical fauna or acclimatization of that fauna to +subtropical conditions.</p> + +<p>Although palynological studies and some faunal studies of subtropical +and temperate animals suggest a drastic climatic fluctuation that might +have eliminated tropical environments in southern México and northern +Central America, there is much biological evidence indicating the +existence of tropical environments in this region even during the +glacial maximum. Especially significant is the diversity of species +inhabiting the present tropical environments; many of these have +differentiated from related taxa to the south.</p> + +<p>In the Pleistocene, climate fluctuated and vegetation shifted +correspondingly in southern México and northern Central America. Most +of the palynological studies and many studies of Pleistocene chronology +deal with montane regions, either the Mexican Plateau or the mountains +rising from the plateau. No such studies have been made in lowland +tropical environments. During glacial advances the tropical lowland +environments in México probably were not eliminated, for the great +diversity of animals in these environments supports the hypothesis that +they have been in existence for some time, although periodically they +may have been discontinuous.</p> + +<p>In order to understand the nature of bioclimatological events in the +Pleistocene in lowland tropical environments of southern México, +certain factors that are of little importance in the interpretation<span class="pagenum"><a name="Page_47" id="Page_47">[Pg 47]</a></span> of +Pleistocene chronology in the highlands must be considered. These +factors are: 1) climatic moderation by oceans, 2) fluctuation in sea +level, and 3) fluctuation in level of the water table as affected by +sea level.</p> + +<p>It is well-known that large bodies of water moderate the temperature on +adjacent land. Furthermore, it is known that faunas of marine +invertebrates shifted latitudinally in the Pleistocene; Trask, Phleger, +and Stetson (1947) recorded cold-water Foraminifera then as far south +as the Sigsbee Deep in the middle of the Gulf of Mexico. Large bodies +of warm water, such as the Gulf of Mexico, Caribbean Sea, and Pacific +Ocean of today, probably were not sufficiently cooled at the time of +glacial advance to affect greatly the temperature of the winds blowing +across them. Even if these bodies of water were somewhat cooler than +now, the prevailing winds blowing from them onto the lowlands of México +and northern Central America would have aided in maintaining relatively +high temperatures there. These warm winds probably counteracted the +cooling effect of glaciation in the lowlands and thereby maintained +tropical conditions near the seas.</p> + +<p>Although no adequate studies of Pleistocene beach lines have been made +in southern México, such information is available for peninsular +Florida on the other side of the Gulf of Mexico (Cooke, 1945). +Fluctuation in sea level in the Pleistocene has been used by Hubbell +(1954), Goin (1958), and Duellman and Schwartz (1958) to explain +present patterns of distribution of animals in Florida. If Cooke's +interpretations can be applied to the western side of the Gulf of +Mexico, even generally, it would be supposed that sea level varied from +about 300 feet lower than at present during the Illinoian Glacial +Period to about 275 feet higher than at present during the Aftonian +Interglacial Period. Lowering of sea level would expand the lowlands in +the isthmus; rising sea level would restrict them, leaving only the +central ridges and many islands in the isthmus, but never forming a +seaway between the Gulf of Mexico and the Pacific Ocean.</p> + +<p>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.</p> + +<p>During times of glacial advances the lowlands of the isthmus probably +were more extensive and had more semi-arid tropical<span class="pagenum"><a name="Page_48" id="Page_48">[Pg 48]</a></span> 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 <i>Rhinophrynus dorsalis</i>, <i>Engystomops pustulosus</i>, and +<i>Hyla staufferi</i> 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 <i>Chiropterotriton</i>, <i>Pseudoeurycea</i>, +<i>Magnadigita</i>, and the <i>eximia</i> group of <i>Hyla</i> 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 <i>Lineatriton +lineola</i>, <i>Anotheca coronata</i>, and <i>Phyllomedusa moreleti</i> could have +crossed the isthmus.</p> + +<p>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 <i>Bolitoglossa occidentalis</i>, <i>Eleutherodactylus +rhodopis</i>, <i>Microbatrachylus pygmaeus</i>, and <i>Rana palmipes</i>.</p> + +<p>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<span class="pagenum"><a name="Page_49" id="Page_49">[Pg 49]</a></span> 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.</p> + + + + +<h2><a name="ACCOUNTS_OF_SPECIES" id="ACCOUNTS_OF_SPECIES"></a>ACCOUNTS OF SPECIES</h2> + + +<p>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:</p> + +<p> +<span style="margin-left: 2.5em;">AMNH American Museum of Natural History</span><br /> +<span style="margin-left: 2.5em;">KU University of Kansas Museum of Natural History</span><br /> +<span style="margin-left: 2.5em;">MCZ Museum of Comparative Zoology, Harvard College</span><br /> +<span style="margin-left: 2.5em;">UIMNH University of Illinois Museum of Natural History</span><br /> +<span style="margin-left: 2.5em;">UMMZ University of Michigan Museum of Zoology</span><br /> +<span style="margin-left: 2.5em;">USNM United States National Museum</span><br /> +</p> + + +<p class="center"><b>Gymnopis mexicanus mexicanus</b> Duméril and Bibron</p> + +<div class="blockquot"><p><i>Oaxaca</i>: El Barrio (3); Matías Romero; Tehuantepec (2). +<i>Veracruz</i>: Cosamaloapan; Cuatotolapam (2).</p></div> + +<p>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).</p> + + +<p class="center"><b>Bolitoglossa occidentalis</b> Taylor</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Río Sarabia (2); Ubero. <i>Veracruz</i>: La Oaxaqueña; +14 km. E of Suchil.</p></div> + +<p>The specimens from Oaxaca are only tentatively assigned to +<i>occidentalis</i>. All are immature and lack maxillary teeth. Taylor +(1941:147) stated that the maxillary teeth are absent in young +<i>occidentalis</i>. One from Río Sarabia is a male with a body-length of 29 +mm. and a tail-length of 22 mm. The dorsum is reddish brown<span class="pagenum"><a name="Page_50" id="Page_50">[Pg 50]</a></span> streaked +with dark gray; the venter is dark gray. Two small individuals (one +from Sarabia and one from Ubero) have body-lengths of 19 and 21 mm. and +tail-lengths of 10.5 and 11 mm. In life they were pale yellowish tan +above with a brown triangular mark on the occiput, but with no +middorsal stripe. Both were found in the axils of elephant ear plants +(<i>Xanthosoma</i>).</p> + +<p>This species has been noted by Goodnight and Goodnight (1956:146) on +the Atlantic lowlands at Palenque, Chiapas, and by Shannon and Werler +(1955:362) at several localities in Los Tuxtlas, Veracruz. I have +collected it at Vista Hermosa on the eastern slopes of the Sierra Madre +Oriental above Tuxtepec in northern Oaxaca. Both <i>B. occidentalis</i> and +<i>B. rufescens</i> have been reported from Palenque, Chiapas (Taylor and +Smith, 1945:547). Reëxamination of specimens from northern Chiapas and +Tabasco is needed to verify the sympatric occurrence of these two +similar species.</p> + + +<p class="center"><b>Bolitoglossa platydactyla</b> Tschudi</p> + +<div class="blockquot"><p><i>Oaxaca</i>: La Oaxaqueña; Tolosita (2). <i>Veracruz</i>: Acayucan; +Cuatotolapam; 25 km. ESE of Jesús Carranza; 14 km. E of +Suchil; 2.7 km. N of Tula.</p></div> + +<p>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.</p> + + +<p class="center"><b>Bolitoglossa veracrucis</b> Taylor</p> + +<div class="blockquot"><p><i>Veracruz</i>: 35 km. SE of Jesús Carranza (21).</p></div> + +<p>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.</p> + + +<p class="center"><b>Rhinophrynus dorsalis</b> Duméril and Bibron</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Ixtepec; Limón; Salina Cruz (18); Tehuantepec +(57); Tuxtepec (3). <i>Veracruz</i>: Amatitlán (3); Cosamaloapan +(5); Novillero (2); San Lorenzo.</p></div> + +<p>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.</p> + +<p class="center">PLATE 1</p> + +<div class="figcenter" style="width: 600px;"> +<a name="pl1-fig1" id="pl1-fig1"></a> +<img src="images/i_035a.jpg" width="600" height="412" alt="Fig. 1. Savanna about 75 kilometers east of +Coatzacoalcos, Veracruz. Photograph by L. C. Stuart."/> +<span class="caption"><span class="smcap">Fig. 1.</span> Savanna about 75 kilometers east of +Coatzacoalcos, Veracruz. Photograph by L. C. Stuart.</span> +</div> +<hr class="tb" /> +<div class="figcenter" style="width: 600px;"> +<a name="pl1-fig2" id="pl1-fig2"></a> +<img src="images/i_035b.jpg" width="600" height="408" alt="Fig. 2. Low scrub forest near Alvarado, Veracruz. +Photograph by L. C. Stuart."/> +<span class="caption"><span class="smcap">Fig. 2.</span> Low scrub forest near Alvarado, Veracruz. +Photograph by L. C. Stuart.</span> +</div> +<hr class="tb" /> + +<p class="center">PLATE 2</p> + +<div class="figcenter" style="width: 600px;"> +<a name="pl2-fig1" id="pl2-fig1"></a> +<img src="images/i_036a.jpg" width="600" height="392" alt="Fig. 1. Rainforest near Tolosita, Oaxaca. March, 1956."/> +<span class="caption"><span class="smcap">Fig. 1.</span> Rainforest near Tolosita, Oaxaca. March, 1956.</span> +</div> + +<hr class="tb" /> + +<div class="figcenter" style="width: 600px;"> +<a name="pl2-fig2" id="pl2-fig2"></a> +<img src="images/i_036b.jpg" width="600" height="396" alt="Fig. 2. Rainforest along the Río Sarabia, Oaxaca. March, +1956."/> +<span class="caption"><span class="smcap">Fig. 2.</span> Rainforest along the Río Sarabia, Oaxaca. March, +1956.</span> +</div> + +<hr class="tb" /> + +<p class="center">PLATE 3</p> + +<div class="figcenter" style="width: 598px;"> +<a name="pl3-fig1" id="pl3-fig1"></a> +<img src="images/i_037a.jpg" width="598" height="418" alt="Fig. 1. Transition forest near La Princesa, Oaxaca. +March, 1956."/> +<span class="caption"><span class="smcap">Fig. 1.</span> Transition forest near La Princesa, Oaxaca. +March, 1956.</span> +</div> + +<hr class="tb" /> + +<div class="figcenter" style="width: 600px;"> +<a name="pl3-fig2" id="pl3-fig2"></a> +<img src="images/i_037b.jpg" width="600" height="385" alt="Fig. 2. Palm Savanna on the Plains of Chivela, Oaxaca. +March, 1956."/> +<span class="caption"><span class="smcap">Fig. 2.</span> Palm Savanna on the Plains of Chivela, Oaxaca. +March, 1956.</span> +</div> + +<hr class="tb" /> + +<p class="center">PLATE 4</p> + +<div class="figcenter" style="width: 600px;"> +<a name="pl4-fig1" id="pl4-fig1"></a> +<img src="images/i_038a.jpg" width="600" height="392" alt="Fig. 1. Scrub forest on the Plains of Tehuantepec in dry +season. March, 1956."/> +<span class="caption"><span class="smcap">Fig. 1.</span> Scrub forest on the Plains of Tehuantepec in dry +season. March, 1956.</span> +</div> + +<hr class="tb" /> + +<div class="figcenter" style="width: 600px;"> +<a name="pl4-fig2" id="pl4-fig2"></a> +<img src="images/i_038b.jpg" width="600" height="404" alt="Fig. 2. Scrub forest on the Plains of Tehuantepec in +rainy season. View toward the north. In the distance is the Continental +Divide in the hills of the Isthmus. July, 1958."/> +<span class="caption"><span class="smcap">Fig. 2.</span> Scrub forest on the Plains of Tehuantepec in +rainy season. View toward the north. In the distance is the Continental +Divide in the hills of the Isthmus. July, 1958.</span> +</div> + +<hr class="tb" /> + +<p class="center">PLATE 5</p> + +<div class="figcenter" style="width: 600px;"> +<a name="pl5-fig1" id="pl5-fig1"></a> +<img src="images/i_039a.jpg" width="600" height="411" alt="Fig. 1. Low, dense scrub forest near La Ventosa, Oaxaca. +July, 1958."/> +<span class="caption"><span class="smcap">Fig. 1.</span> Low, dense scrub forest near La Ventosa, Oaxaca. +July, 1958.</span> +</div> + +<hr class="tb" /> + +<div class="figcenter" style="width: 600px;"> +<a name="pl5-fig2" id="pl5-fig2"></a> +<img src="images/i_039b.jpg" width="600" height="424" alt="Fig. 2. Temporary pond in scrub forest north of Salina +Cruz, Oaxaca. July 7, 1958. Rhinophrynus dorsalis, Bufo marmoreus, +and Diaglena reticulata were breeding here the previous night."/> +<span class="caption"><span class="smcap">Fig. 2.</span> Temporary pond in scrub forest north of Salina +Cruz, Oaxaca. July 7, 1958. <i>Rhinophrynus dorsalis</i>, <i>Bufo marmoreus</i>, +and <i>Diaglena reticulata</i> were breeding here the previous night.</span> +</div> + +<hr class="tb" /> + +<p class="center">PLATE 6</p> + +<div class="figcenter" style="width: 600px;"> +<a name="pl6-fig1" id="pl6-fig1"></a> +<img src="images/i_040a.jpg" width="600" height="392" alt="Fig. 1. Calling male of Rhinophrynus dorsalis, +photographed in a pond north of Santa Cruz, Oaxaca, on July 6, 1958. × +2/3."/> +<span class="caption"><span class="smcap">Fig. 1.</span> Calling male of <i>Rhinophrynus dorsalis</i>, +photographed in a pond north of Santa Cruz, Oaxaca, on July 6, 1958. × +2/3.</span> +</div> + +<hr class="tb" /> + +<div class="figcenter" style="width: 600px;"> +<a name="pl6-fig2" id="pl6-fig2"></a> +<img src="images/i_040b.jpg" width="600" height="414" alt="Fig. 2. Color pattern variation in two adults of Bufo +canaliferus from Juchitán, Oaxaca. × 2/3."/> +<span class="caption"><span class="smcap">Fig. 2.</span> Color pattern variation in two adults of <i>Bufo +canaliferus</i> from Juchitán, Oaxaca. × 2/3.</span> +</div> + +<hr class="tb" /> + +<p class="center">PLATE 7</p> + +<div class="figcenter" style="width: 600px;"> +<a name="pl7-fig1" id="pl7-fig1"></a> +<img src="images/i_041a.jpg" width="600" height="432" alt="Fig. 1. Calling male of Engystomops pustulosus, +photographed in a pond west of Tehuantepec, Oaxaca, on July 5, 1956. × +2."/> +<span class="caption"><span class="smcap">Fig. 1.</span> Calling male of <i>Engystomops pustulosus</i>, +photographed in a pond west of Tehuantepec, Oaxaca, on July 5, 1956. × +2.</span> +</div> + +<hr class="tb" /> + +<div class="figcenter" style="width: 600px;"> +<a name="pl7-fig2" id="pl7-fig2"></a> +<img src="images/i_041b.jpg" width="600" height="422" alt="Fig. 2. Foamy egg mass of Engystomops pustulosus at +the edge of a pond west of Tehuantepec, Oaxaca. July 5, 1956. × 3/8."/> +<span class="caption"><span class="smcap">Fig. 2.</span> Foamy egg mass of <i>Engystomops pustulosus</i> at +the edge of a pond west of Tehuantepec, Oaxaca. July 5, 1956. × 3/8.</span> +</div> + +<hr class="tb" /> + +<p class="center">PLATE 8</p> + +<div class="figcenter" style="width: 600px;"> +<a name="pl8-fig1" id="pl8-fig1"></a> +<img src="images/i_042a.jpg" width="600" height="417" alt="Fig. 1. Calling male of Diaglena reticulata, +photographed at a pond north of Salina Cruz, Oaxaca, on July 6, 1958. × +1/2."/> +<span class="caption"><span class="smcap">Fig. 1.</span> Calling male of <i>Diaglena reticulata</i>, +photographed at a pond north of Salina Cruz, Oaxaca, on July 6, 1958. × +1/2.</span> +</div> + +<hr class="tb" /> + +<div class="figcenter" style="width: 600px;"> +<a name="pl8-fig2" id="pl8-fig2"></a> +<img src="images/i_042b.jpg" width="600" height="386" alt="Fig. 2. Clasping pair of Diaglena reticulata at the +edge of a pond north of Salina Cruz, Oaxaca, on July 6, 1958. × 1."/> +<span class="caption"><span class="smcap">Fig. 2.</span> Clasping pair of <i>Diaglena reticulata</i> at the +edge of a pond north of Salina Cruz, Oaxaca, on July 6, 1958. × 1.</span> +</div> + +<p><span class="pagenum"><a name="Page_51" id="Page_51">[Pg 51]</a></span></p> + +<hr class="tb" /> + +<p>Breeding congregations were found after heavy rains at Tehuantepec on +July 5, 1956, at Cosamaloapan, Novillero, and Amatitlán 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 (<a href="#pl6-fig1">Pl. 6, fig. 1</a>). 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.</p> + + +<p class="center"><b>Bufo canaliferus</b> Cope</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Chivela; Salina Cruz; Santa Efigenia; Tapanatepec +(6); Tehuantepec (10); Zanatepec (4).</p></div> + +<p>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 (<a href="#pl6-fig2">Pl. 6, fig. 2</a>).</p> + + +<p class="center"><b>Bufo coccifer</b> Cope</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Juchitán (5); Tehuantepec.</p></div> + +<p>It is with some degree of hesitancy that these toads are referred to +the species <i>coccifer</i>. Although these and other specimens from +Guerrero and Michoacán display no striking differences from specimens +from Costa Rica, Nicaragua, and southeastern Guatemala, the ranges of +the populations are separated by a broad hiatus in Chiapas and +Guatemala. Possibly this species has utilized the sub-humid corridor +through northern Central America (Stuart, 1954) and subsequently +disappeared from the corridor in Guatemala and Chiapas. Specimens of a +<i>coccifer</i>-like toad collected by Stuart in the vicinity of +Jacaltenango, Departamento Huehuetenango, Guatemala, are much larger +than either the Central American or Mexican specimens of <i>coccifer</i>. A +final commitment on the systematic status must await a thorough study +of this group of toads.</p> + +<p>Males of this species were calling from a grassy rain-pool in open +scrub forest at the edge of Juchitán on July 6, 1956. The call is a +low<span class="pagenum"><a name="Page_52" id="Page_52">[Pg 52]</a></span> "whirrr." The calling males were sitting in the shallow water at +the edge of pond, where they were hidden by the grass. None was +observed in open water, as is characteristic of calling males of <i>Bufo +canaliferus</i> and <i>marmoreus</i>.</p> + + +<p class="center"><b>Bufo marinus</b> Linnaeus</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Agua Caliente; Guichicovi (3); Mixtequilla; +Tolosita (6); Tehuantepec (37); Tuxtepec; Unión Hidalgo. +<i>Veracruz</i>: Ciudad Alemán (4); Cosamaloapan; Cuatotolapam +(19); 20 km. SE of Jesús Carranza (4); 38 km. SE of Jesús +Carranza (10); 20 km. NE of Jesús Carranza (4); Novillero.</p></div> + +<p>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.</p> + + +<p class="center"><b>Bufo marmoreus</b> Wiegmann</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Cerro San Pedro (2); Chivela (5); Escurano (3); +Juchitán; Salina Cruz (101); Santa Lucía (2); 12 km. S of +Santiago Chivela (11); Santo Domingo; Tapanatepec; +Tehuantepec (100); Tequisistlán. <i>Veracruz</i>: Alvarado; +Coatzacoalcos.</p></div> + +<p>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 Río (Langebartel and Smith, 1959:27).</p> + +<p>The similarity in size of <i>Bufo marmoreus</i> and <i>valliceps</i> and their +almost completely allopatric ranges suggest that the two species may be +in competition at any one locality. Nevertheless, both were calling +from a small rocky stream south of Santiago Chivela on July 6, 1956.</p> + +<p>On the night of July 6, 1958, an estimated 400 toads of this species +made up a breeding congregation near Salina Cruz. The site was a +shallow muddy pond about 20 × 40 meters located in an area cleared of +scrub forest; the banks of the pond were devoid of vegetation (<a href="#pl5-fig2">Pl. 5, fig. 2</a>). +Breeding in the same pond were <i>Rhinophrynus dorsalis</i> and +<i>Diaglena reticulata</i>. The following morning no more than a dozen +<i>Bufo</i> were found in the pond, but several individuals were found +beneath debris and in small burrows near the pond. On July 7, 1958, +large numbers of tadpoles and recently metamorphosed young were in a +shallow grassy pool just east of Salina Cruz.</p> + +<p>Taylor (1943b:347) referred certain specimens from Tehuantepec to <i>Bufo +perplexus</i>, a species closely related to <i>Bufo marmoreus</i>.<span class="pagenum"><a name="Page_53" id="Page_53">[Pg 53]</a></span> Evidence to +be presented elsewhere shows that <i>perplexus</i> does not occur in the +isthmus.</p> + + +<p class="center"><b>Bufo valliceps valliceps</b> Wiegmann</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Guichicovi (2); Matías Romero; 32 km. N of Matías +Romero (2); Nueva Raza; Río Sarabia (3); Santa María +Chimalapa (14); Santiago Chivela; 12 km. S of Santiago +Chivela (5); Santo Domingo (5); Tolosita (7). <i>Veracruz</i>: +Acayucan (3); Alvarado; Amatitlán; Ayentes; Cosamaloapan +(3); Cosoleacaque (6); Cuatotolapam (14); Hueyapan; 20 km. +ENE of Jesús Carranza (6); 20 km. S of Jesús Carranza; 25 +km. SE of Jesús Carranza (23); 35 km. SE of Jesús Carranza; +60 km. SW of Jesús Carranza (5); La Oaxaqueña (4); Novillero +(4); San Lorenzo (5).</p></div> + +<p>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.</p> + +<p>I have not been able to recognize individuals referrable to the race +<i>macrocristatus</i>. Firschein and Smith (1957:219) described +<i>macrocristatus</i> from the mountains of eastern Oaxaca and referred to +it specimens from the Gulf lowlands of northern Chiapas. None of the +present material shows the hypertrophied cranial crests supposedly +characteristic of <i>macroaristatus</i>, nor do specimens from the isthmus +resemble the population in the Grijalva Valley being described by L. C. +Stuart, who will discuss the variation in, and the validity of, the +named populations of <i>valliceps</i>.</p> + +<p>Five specimens from San Lorenzo, Veracruz (USNM 123516-20), were +identified as <i>Bufo cristatus</i> by Smith (1947:408). Firschein (1950:83) +redefined the <i>cristatus</i> group of <i>Bufo</i> and assigned these specimens +to <i>valliceps</i>.</p> + + +<p class="center"><b>Eleutherodactylus alfredi</b> Boulenger</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Tolosita (2). <i>Veracruz</i>: 35 km. SE of Jesús +Carranza (6).</p></div> + +<p>These specimens were collected in rainforest. Shreve (1957:247) pointed +out the close resemblance between <i>E. alfredi</i> and <i>E. conspicuus</i> from +Piedras Negras, Guatemala, and treated them as subspecies. Examination +of the specimens from the isthmus, together with seven from central +Veracruz and one from Teapa, Tabasco, suggests an even closer +relationship. <i>Eleutherodactylus conspicuus</i> was diagnosed by Taylor +and Smith (1945:567) as differing from <i>alfredi</i> "in lacking a tarsal +fold, in having shorter hind legs with the tibiotarsal articulation +reaching only to the nostril instead of beyond the tip of the snout; +the vomerine teeth barely reach the posterior level of the choanae." +The specimen from<span class="pagenum"><a name="Page_54" id="Page_54">[Pg 54]</a></span> Teapa has the vomerine teeth reaching to the +posterior edge of the choanae; in the eight specimens from the isthmus +the teeth reach the posterior edge of the choanae in two and to the +middle of the choanae in six; in seven specimens from central Veracruz +the teeth reach the posterior edge of the choanae in two and to the +middle in five. The tibiotarsal articulation extends beyond the tip of +the snout in the specimen from Teapa and in two from central Veracruz; +in three specimens from the isthmus and in one from central Veracruz it +extends only to the nostril; in the others it extends to the snout. The +tarsal fold is absent in the specimen from Teapa, in three from the +isthmus, and in all those from central Veracruz; it is weakly present +in the others.</p> + +<p>In the light of this evidence there seems to be little justification in +recognizing two species or even two subspecies in this group. +Consequently, <i>Eleutherodactylus conspicuus</i> Taylor and Smith (1945) is +here placed in the synonymy of <i>Eleutherodactylus alfredi</i> Boulenger +(1898), a species with a range extending from Cuautlapan and Potrero +Viejo in central Veracruz southward and eastward in forested habitats +to western El Petén, Guatemala.</p> + + +<p class="center"><b>Eleutherodactylus natator</b> Taylor</p> + +<div class="blockquot"><p><i>Veracruz</i>: 35 km. SE of Jesús Carranza (3); 38 km. S of +Jesús Carranza; 55 km. SE of Jesús Carranza.</p></div> + +<p>The snout-vent length is 42.0 mm. in a male and averages 59.5 mm. in +three adult females. The tarsal fold is low and extends about half the +length of the tarsus; the first and second fingers are subequal in +length; the tibiotarsal articulation extends beyond the tip of the +snout. The patches of vomerine teeth lie between the posterior margins +of the choanae. The throat and belly are immaculate, and the soles of +the feet are dark. In the isthmus this species can be distinguished +from <i>Eleutherodactylus rugulosus</i> by less rugose skin on the dorsum +and absence of dark ventral mottling.</p> + +<p>The specimens reported here extend the known range of <i>natator</i> +eastward from Camotlán, Oaxaca; northward in Veracruz the species +inhabits foothills as far north as Huatusco.</p> + + +<p class="center"><b>Eleutherodactylus rhodopis</b> Cope</p> + +<div class="blockquot"><p><i>Oaxaca</i>: 30 km. N of Matías Romero; Río Sarabia (5); +Tapanatepec (87); Tolosita (6); between Zanatepec and +Tapanatepec. <i>Veracruz</i>: 25 km. SE of Jesús Carranza; 35 km. +SE of Jesús Carranza (2); 22 km. SSW of Jesús Carranza; 20 +km. ENE of Jesús Carranza (7); Minatitlán; Tapalapan (5).</p></div> + +<p>For the purposes of the present study I am not recognizing +<i>Eleutherodactylus beati</i>, <i>E. dorsoconcolor</i>, and <i>E. venustus</i> as +specifically,<span class="pagenum"><a name="Page_55" id="Page_55">[Pg 55]</a></span> or even subspecifically distinct from the earlier named +<i>E. rhodopis</i>. Probably these are mere color varieties of a single +species.</p> + +<p>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.</p> + + +<p class="center"><b>Eleutherodactylus rugulosus</b> Cope</p> + +<div class="blockquot"><p><i>Oaxaca</i>: La Princesa (30); Modelo; Santa Lucía (10); +Tapanatepec (26); Tehuantepec (6); Tres Cruces (8). +<i>Veracruz</i>: Tapalapan (5).</p></div> + +<p>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 +(Mirador and Chacalapa) and on several specimens from Los Tuxtlas in +Veracruz (Los Chaneques, 67; Salto de Eyipantla, 35; and San Andrés +Tuxtla, 11).</p> + +<p>Frogs of the <i>Eleutherodactylus rugulosus</i> complex occur from southern +Veracruz and Sinaloa southward through Central America. Taylor +(1940:401) described <i>E. vocalis</i> from Hacienda El Sabino, Michoacán; +Taylor and Smith (1945:580) described <i>E. avocalis</i> from Tres Cruces, +Oaxaca. These have been considered as species distinct from +<i>rugulosus</i>, which is known to occur in Veracruz, Guerrero, and Chiapas +southward into Central America. Although the large number of specimens +collected in the isthmus does not aid in defining the ranges of the +taxa involved, these specimens do give some idea of the variation in +certain characters in a given population.</p> + +<p>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<span class="pagenum"><a name="Page_56" id="Page_56">[Pg 56]</a></span> 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 (1958b:6) discussed the variation in +these characters in populations in Colima, Jalisco, and Michoacán.</p> + +<p>Until the extent of variation of these characters is known throughout +the range of <i>rugulosus</i>, the recognition of populations either as +species or subspecies seems superfluous. Consequently, I have used the +oldest name; this does not necessarily imply, however, that all +populations of <i>rugulosus</i> (<i>sensu lato</i>) are conspecific.</p> + +<p>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.</p> + + +<p class="center"><b>Microbatrachylus pygmaeus</b> Taylor</p> + +<div class="blockquot"><p><i>Oaxaca</i>: La Princesa (5); Matías Romero (9); Río Sarabia +(41); Tolosita (2). <i>Veracruz</i>: Jesús Carranza; 20 km. ENE +of Jesús Carranza.</p></div> + +<p>The specimens listed above vary widely in color patterns; some of the +patterns are characteristic of certain named "species": <i>albolabris</i>, +<i>imitator</i>, <i>lineatissimus</i>, and <i>minimus</i>. The large series from the +Río Sarabia contains all of the color patterns; this series was +obtained in one small ravine in the rainforest. At least in the +isthmian region, this species does not inhabit the Pacific slopes and +lowlands.</p> + + +<p class="center"><b>Syrrhophus leprus</b> Cope</p> + +<div class="blockquot"><p><i>Oaxaca</i>: 33 km. N of Matías Romero; Santa Efigenia. +<i>Veracruz</i>: San Lorenzo.</p></div> + +<p>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 +(1958c:8).</p> + +<p>Smith (1947:408) reported a specimen of <i>Syrrhophus verruculatus</i> +Peters from San Lorenzo, Veracruz; he stated that this specimen<span class="pagenum"><a name="Page_57" id="Page_57">[Pg 57]</a></span> (USNM +123530) could not be <i>S. leprus</i>, because it had a gray belly, nor <i>S. +cystignathoides</i>, because of the dark and light dorsal coloration. +Firschein (1954:57) in his review of the species of <i>Syrrhophus</i> in +eastern México referred the specimen to <i>S. cystignathoides</i>. The +specimen is in poor condition. Nevertheless, specific determination is +possible. Numerous specimens of <i>S. leprus</i> from Los Tuxtlas have gray +bellies; some have heavier pigmentation than the specimen from San +Lorenzo. In preservative the dorsum is dark brown with lighter +mottling. There is little doubt that the specimen from San Lorenzo is a +<i>Syrrhophus leprus</i>, an abundant and widespread species in the humid +Gulf lowlands of southern México, and not <i>verruculatus</i>, if this is a +valid species (see Firschein, <i>op. cit.</i>:58), and not +<i>cystignathoides</i>, a species known from San Luis Potosí southward to +central Veracruz.</p> + + +<p class="center"><b>Syrrhophus pipilans pipilans</b> Taylor</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Cerro Arenal; Cerro San Pedro; 6 km. N of Chivela; +14 km. W of Tehuantepec (2).</p></div> + +<p>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."</p> + + +<p class="center"><b>Engystomops pustulosus</b> Cope</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Chivela; La Ventosa (3); Santo Domingo; +Tapanatepec (14); Tehuantepec (61); Unión Hidalgo (62). +<i>Veracruz</i>: Acayucan; Cuatotolapam (7); 10 km. SE of +Hueyapan (11).</p></div> + +<p>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 (<a href="#pl7-fig1">Pl. 7, fig. 1</a>); +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 (<a href="#pl7-fig2">Pl. 7, fig. 2</a>). The large series from Unión +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.</p> + +<p>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<span class="pagenum"><a name="Page_58" id="Page_58">[Pg 58]</a></span> from Cordoba and +Mirador, Veracruz, are like those from Tehuantepec.</p> + + +<p class="center"><b>Leptodactylus labialis</b> Cope</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Agua Caliente; Chivela (2); Matías Romero (12); 33 +km. N of Matías Romero (4); Mixtequilla; Santa Efigenia; +Tapanatepec; Tehuantepec (38); Tolosita (2); 33 km. W of +Zanatepec (49). <i>Veracruz</i>: Acayucan (3); Ciudad Alemán; +Cuatotolapam (10); Hueyapan; La Oaxaqueña (4); 38 km. SE of +Jesús Carranza; 20 km. ENE of Jesús Carranza; Novillero (3); +San Lorenzo (2).</p></div> + +<p>Although <i>Leptodactylus labialis</i> does not appear to be so abundant as +<i>Leptodactylus melanonotus</i>, 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.</p> + + +<p class="center"><b>Leptodactylus melanonotus</b> Hallowell</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Agua Caliente (25); Cerro Arenal (2); Cerro +Quiengola (3); Cerro San Pedro (3); Chivela (2); Coyol; +Juchitán; Matías Romero (11); Mixtequilla (2); Papaloapan +(2); Salazar (9); Salina Cruz; 11 km. S of Santiago Chivela; +Tapanatepec (17); Tehuantepec (176); Tolosita; Unión +Hidalgo; 27 km. W of Zanatepec (6). <i>Veracruz</i>: Acayucan; +Cuatotolapam (9); Cosoleacaque; 20 km. ENE of Jesús Carranza +(2); 20 km. SE of Minatitlán (2); Novillero; San Lorenzo +(6).</p></div> + +<p>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.</p><p><span class="pagenum"><a name="Page_59" id="Page_59">[Pg 59]</a></span></p> + + +<p class="center"><b>Diaglena reticulata</b> Taylor</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Cerro Arenal; Chivela; Salina Cruz (26); San +Antonio (3); Tehuantepec (2); 8.6 km. W of Tehuantepec (11); +Zarzamora.</p></div> + +<p>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 (<a href="#pl5-fig2">Pl. 5, fig. 2</a>); males called from the bank near the water, and +clasping pairs were found only on land (<a href="#pl8-fig1">Pl. 8, figs. 1</a>-<a href="#pl8-fig2">2</a>). The call is +a loud, nasal "braaa," two to three seconds in duration. Amplexus is +axillary.</p> + +<p>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 +<i>reticulata</i> (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.</p> + + +<p class="center"><b>Hyla baudini</b> Duméril and Bibron</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Bisilana; Cerro Quiengola (2); Cerro San Pedro; +Coyol; Matías Romero (12); Mixtequilla; Río Sarabia (7); +Salazar; San Antonio; 11 km. S of Santiago Chivela; Santo +Domingo (3); Tapanatepec (2); Tehuantepec (23); Tolosita. +<i>Veracruz</i>: Acayucan; Amatitlán; Ciudad Alemán (3); +Cosamaloapan (2); Cuatotolapam (15); 10 km. SE of Hueyapan; +20 km. S of Jesús Carranza; 38 km. S of Jesús Carranza (2); +20 km. ENE of Jesús Carranza (4); La Oaxaqueña (2); +Minatitlán (2); Naranja (3); Novillero (9); Río de las +Playas (2); San Lorenzo (5); Tapalapan (2).</p></div> + +<p>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 +(<i>Xanthosoma</i>), 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.</p> + + +<p class="center"><b>Hyla ebraccata</b> Cope</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Donají (17); 43 km. N of Matías Romero (27); +Sarabia (6); Tolosita (3); Ubero (17). <i>Veracruz</i>: Aquilera.</p></div> + +<p>This small species was found only in forested areas, where calling +males were on bushes and trees around rain pools. The call<span class="pagenum"><a name="Page_60" id="Page_60">[Pg 60]</a></span> 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.</p> + +<p>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.</p> + +<p>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, México, in humid forests to El Petén, +Guatemala.</p> + + +<p class="center"><b>Hyla loquax</b> Gaige and Stuart</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Donají (7); 43 km. N of Matías Romero (21). +<i>Veracruz</i>: 19 km. N of Acayucan (4); Aquilera (3); 8 km. SW +of Coatzacoalcos (36); Cuototolapam (11); Naranja (13); San +Lorenzo (8).</p></div> + +<p>In the isthmus this species is known only from the humid forests of the +Gulf lowlands; it is also known from Boca del Río, Veracruz, and from +Teapa and Villa Hermosa, Tabasco.</p> + +<p>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<span class="pagenum"><a name="Page_61" id="Page_61">[Pg 61]</a></span> 41.7 mm.; they have no horny nuptial pads on the +pollex.</p> + +<p>The relationships of this species are with <i>Hyla rickardsi</i> 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 +<i>loquax</i> and yellow in <i>rickardsi</i>. The calls of the two species are +distinctly different; that of <i>rickardsi</i> is a high-pitched, loud +rattle continued for several seconds, notably different from the +goose-like honk of <i>loquax</i>.</p> + +<p class="center"><span class="smcap">Table 1.—Comparison of Certain Characters in Hyla loquax and Hyla rickardsi</span></p> + + +<div class="center"> +<table border="1" cellpadding="6" cellspacing="0" summary="table1"> +<tr><th align="center"><span class="smcap">Character</span></th><th align="center"><i>loquax</i></th><th align="center"><i>rickardsi</i></th></tr> +<tr><td align="left">Toe webbing</td><td align="left">Full</td><td align="left">Three-fourths</td></tr> +<tr><td align="left">Finger webbing</td><td align="left">Three-fourths</td><td align="left">One-half</td></tr> +<tr><td align="left">Average snout-vent length (♂)</td><td align="left">41.7 mm.</td><td align="left">37.4 mm.</td></tr> +<tr><td align="left">Tympanum/eye (♂)</td><td align="left">63.2%</td><td align="left">55.8%</td></tr> +<tr><td align="left">Dorsal leg pattern</td><td align="left">Barred</td><td align="left">Unmarked</td></tr> +<tr><td align="left">Tarsal fold</td><td align="left">Tubercular</td><td align="left">Absent</td></tr> +<tr><td align="left">Tarsal stripe</td><td align="left">Absent or indistinct</td><td align="left">Broad, indistinct, or absent</td></tr> +<tr><td align="left">Dorsolateral stripe</td><td align="left">Absent</td><td align="left">Present</td></tr> +<tr><td align="left">Light line over anus</td><td align="left">Broad</td><td align="left">Narrow</td></tr> +<tr><td align="left">Flash colors</td><td align="left">Red</td><td align="left">Yellow</td></tr> +<tr><td align="left">Iris color</td><td align="left">Copper</td><td align="left">Bronze</td></tr> +</table></div> + +<p>The three specimens from San Lorenzo, Veracruz (USNM 123513-5), were +identified as <i>Hyla rickardsi</i> 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 <i>loquax</i>, but in the +amount of webbing on the hands and feet, broad tarsal stripe, and +narrow anal stripe they are like <i>rickardsi</i>. The third specimen, a +juvenile, has a snout-vent length of 25 mm. In coloration it resembles +the adults; it has more distinct<span class="pagenum"><a name="Page_62" id="Page_62">[Pg 62]</a></span> bars on the limbs. On the basis of +geography these specimens should be <i>loquax</i>, for the closest known +record of <i>rickardsi</i> is more than 200 kilometers to the northwest, +whereas <i>loquax</i> is known from several localities around San Lorenzo.</p> + +<p>Shannon and Werler (1955:383) described <i>Hyla axillamembrana</i> 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 <i>loquax</i>. It is not +possible to determine the color of the thighs, nor was this information +given in the description. <i>Hyla axillamembrana</i> is here considered to +be a synonym of <i>Hyla loquax</i>.</p> + + +<p class="center"><b>Hyla microcephala martini</b> Smith</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Donají (15); 43 km. N of Matías Romero (19); Río +Sarabia (2); Sarabia (11); Tolosita. <i>Veracruz</i>: Acayucan +(17); Alvarado (41); Aquilera (21); 8 km. SW of +Coatzacoalcos (10); Cosoleacaque (26); 10 km. SE of +Hueyapan; Naranja (3); Novillero.</p></div> + +<p>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.</p> + +<p>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.</p> + + +<p class="center"><b>Hyla picta</b> Günther</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Donají (8); Sarabia (11); Tolosita (15); Ubero +(6). <i>Veracruz</i>: 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).</p></div> + +<p>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<span class="pagenum"><a name="Page_63" id="Page_63">[Pg 63]</a></span> 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.</p> + + +<p class="center"><b>Hyla robertmertensi</b> Taylor</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Tapanatepec (28); 7.5 km. NW of Tapanatepec (38); +7.2 km. WNW of Zanatepec (77).</p></div> + +<p>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."</p> + +<p>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.</p> + + +<p class="center"><b>Hyla staufferi</b> Cope</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Chivela; Huilotepec (5); Juchitán (4); Matías +Romero (4); 25 km. N of Matías Romero; Mixtequilla (4); Río +Sarabia (11); 11 km. S of Santiago Chivela; Sarabia (3); +Tapanatepec (67); Tehuantepec (66); Tolosita (2); Ubero; +Unión Hidalgo; Zanatepec (6). <i>Veracruz</i>: Acayucan (7); +Alvarado (3); Amatitlán; Aquilera; Ciudad Alemán (3); 8 km. +SW of Coatzacoalcos (9); Cosamaloapan (4); Cosoleacaque (8); +10 km. SE of Hueyapan; Lerdo de Tejada; Novillero (6); Tula +(2).</p></div> + +<p>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<span class="pagenum"><a name="Page_64" id="Page_64">[Pg 64]</a></span> breeding season, that of <i>H. +staufferi</i> 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.</p> + +<p>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.</p> + + +<p class="center"><b>Hylella sumichrasti</b> Brocchi</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Cerro Arenal (5); Cerro San Pedro (2); Escurano; +La Concepción (41); Portillo Los Nanches (6); San Antonio +(16); 11 km. S of Santiago Chivela (18); Santa Lucía (7); +Tapanatepec (5); Tehuantepec (8); Tenango (49); Tres Cruces +(19).</p></div> + +<p>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.</p> + +<p>Certain diagnostic characters of this species as given by Taylor +(1943a:50) and Taylor and Smith (1945:598) are in need of revision. +<i>Hylella sumichrasti</i> 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<span class="pagenum"><a name="Page_65" id="Page_65">[Pg 65]</a></span> 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 (<i>op. cit.</i>) and Taylor and Smith (<i>op. cit.</i>); 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.</p> + +<p>The range of this species encompasses the Pacific slopes of the Isthmus +of Tehuantepec eastward to the upper Cintalapa Valley and vicinity of +Tonalá in western Chiapas. Priscilla Starrett collected tadpoles of <i>H. +sumichrasti</i> 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.</p> + + +<p class="center"><b>Phrynohyas modesta</b> Taylor and Smith</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Tuxtepec. <i>Veracruz</i>: 20 km. S of Jesús Carranza; +20 km. ENE of Jesús Carranza (2); Minatitlán.</p></div> + +<p>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.</p> + +<p>The acquisition of several specimens of this species in southern +Veracruz, Tabasco, and Oaxaca, together with a knowledge of the +variation displayed by <i>Phrynohyas spilomma</i>, suggests that <i>modesta</i> +may be a color variety of <i>spilomma</i>.</p> + + +<p class="center"><b>Phrynohyas spilomma</b> Cope</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Tapanatepec (3). <i>Veracruz</i>: Amatitlán (12); +Chacaltianguis (2); Ciudad Alemán (6); Cosamaloapan; +Novillero (3).</p></div> + +<p>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 <i>P. modesta</i>. Furthermore, this frog<span class="pagenum"><a name="Page_66" id="Page_66">[Pg 66]</a></span> was not +detected in the rainforests of the Gulf lowlands; in that region it was +found only in scrub forest and savanna.</p> + +<p>On July 26, 1956, numerous choruses of these frogs were heard between +Ciudad Alemán 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 Alemán.</p> + + +<p class="center"><b>Phyllomedusa callidryas taylori</b> Funkhouser</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Donají (9); Sarabia (8); Tolosita (6); Ubero (27). +<i>Veracruz</i>: Alvarado (7); Aquilera; Berta; Coatzacoalcos +(9); 10 km. SE of Hueyapan (5); Naranja (17).</p></div> + +<p>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.</p> + +<p>Throughout the month of July, 1956, <i>Phyllomedusa</i> 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.</p> + + +<p class="center"><b>Phyllomedusa dacnicolor</b> Cope</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Escurano; Tehuantepec.</p></div> + +<p>Although it is abundant on the Pacific lowlands to the northwest in +Guerrero, Michoacán, and Colima, this species is known only from two +specimens from Tehuantepec. There is no apparent<span class="pagenum"><a name="Page_67" id="Page_67">[Pg 67]</a></span> 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.</p> + + +<p class="center"><b>Gastrophryne usta</b> Cope</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Santa Efigenia; Tehuantepec (10); 24 km. W of +Tehuantepec; Tolosita (2). <i>Veracruz</i>: Ayentes (6); La +Oaxaqueña; Novillero (2); San Lorenzo.</p></div> + +<p>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).</p> + +<p>Specimens from the Pacific lowlands are typical of <i>Gastrophryne usta +gadowi</i> Boulenger in possessing a thin line on the posterior surface of +the thighs and a thin line from the snout to the vent. Of nine +specimens from the Gulf lowlands (Ayentes, Novillero, and San Lorenzo), +seven have a middorsal line; this is narrow in four and wide in three. +Five have the stripes on the thighs. Two specimens from the middle of +the isthmus (Tolosita) have no stripes on the thighs; one has a thin +middorsal line, and the other has a broad line. The adult males have a +black throat; females have a mottled one. The brown reticulations on +the bellies of specimens from the Gulf lowlands is bolder than on +specimens from the Pacific lowlands. The presence of certain characters +supposedly diagnostic of the subspecies <i>gadowi</i> (line on dorsum and +thighs) in the population of <i>usta</i> in southern Veracruz suggests that +a redefinition of the ranges of these subspecies will be in order when +sufficient material is available to delimit them accurately. For the +present I prefer to consider all specimens from the isthmus solely as +<i>Gastrophryne usta</i> without referring them to subspecies.</p> + + +<p class="center"><b>Rana palmipes</b> Spix</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Matías Romero (11); 11 km. S of Santiago Chivela; +Santo Domingo; Sarabia. <i>Veracruz</i>: Coatzacoalcos; +Cuatotolapam; 25 km. SE of Jesús Carranza (4); Tlacotalpan +(2); Tula.</p></div> + +<p>Adults were found along streams and in marshes in savannas and +rainforest. These frogs are wary and difficult to capture, even at +night. <i>Rana palmipes</i> is another species that has a discontinuous +distribution in the isthmus. The species does not occur on the Pacific +lowlands of the isthmus, but does occur on the more humid Pacific +slopes of Chiapas and Guatemala.</p> + +<p>Tadpoles were found in a small sluggish tributary to the Río Sarabia.</p><p><span class="pagenum"><a name="Page_68" id="Page_68">[Pg 68]</a></span></p> + + +<p class="center"><b>Rana pipiens</b> Schreber</p> + +<div class="blockquot"><p><i>Oaxaca</i>: Agua Caliente; Cerro Quiengola; Escurano (14); Río +Sarabia (2); Tapanatepec (5); Tehuantepec (24). <i>Veracruz</i>: +Acayucan; Cuatotolapam (15); Jesús Carranza (2); 20 km. S of +Jesús Carranza (11); 25 km. SE of Jesús Carranza; 20 km. ENE +of Jesús Carranza (10); San Lorenzo (10).</p></div> + +<p>As in most other places in México and northern Central America, this +species occurs wherever there is permanent water. Males were heard +calling from woodland ponds and from savanna ponds.</p> + + + + +<h2><a name="SUMMARY" id="SUMMARY"></a>SUMMARY</h2> + + +<p>Investigations of the amphibians and their environments in the Isthmus +of Tehuantepec have been presented with the aim of gaining an +understanding of the present biological and of the historical events +responsible for the present patterns of distribution of amphibians in +this region.</p> + +<p>The Isthmus of Tehuantepec embraces three major environments—savanna, +semi-arid scrub forest, and quasi-rainforest. The rainforest presents +an environment noticeably different from the other two and has a +different amphibian fauna.</p> + +<p>Analysis of present patterns of distribution shows that certain species +are restricted to the rainforests on the Gulf lowlands; others live +only in the semi-arid scrub forests on the Pacific lowlands. A third +group of species lives on both the Gulf and Pacific lowlands; most of +these species occur only in the scrub forests or savannas on the Gulf +lowlands, but some also inhabit the rainforest. In one way or another +the isthmus presents a barrier to the distribution of 75 per cent of +the species of amphibians living in the lowlands; it is a greater +barrier still to the species inhabiting the highlands on either side.</p> + +<p>Present patterns of distribution are attributed to bioclimatic +fluctuation in the Pleistocene. In the course of these climatic shifts, +tropical environments and their amphibian inhabitants seem to have +survived in the isthmian region.</p> + +<p>The amphibian fauna of the lowlands of the Isthmus of Tehuantepec +consists of 16 genera and 36 species. Systematic studies of all +available specimens from the region show that <i>Eleutherodactylus +conspicuus</i> Taylor and Smith is a synonym of <i>Eleutherodactylus +alfredi</i> Boulenger and that <i>Hyla axillamembrana</i> Shannon and Werler is +a synonym of <i>Hyla loquax</i> Gaige and Stuart.</p><hr class="chap" /><p><span class="pagenum"><a name="Page_69" id="Page_69">[Pg 69]</a></span></p> + + + + +<h2><a name="LITERATURE_CITED" id="LITERATURE_CITED"></a>LITERATURE CITED</h2> + + +<p><span class="smcap">Beard, J. S.</span></p> + +<p class="i2">1953. The savanna vegetation of northern tropical America. +Ecol. Mono., vol. 23 (2):149-215.</p> + +<p><span class="smcap">Boulenger, G. A.</span></p> + +<p class="i2">1898. Fourth report on additions to the batrachian +collection in the Natural History Museum. Proc. Zool. Soc. +London, 1898, pp. 473-482, pls. 38-39.</p> + +<p><span class="smcap">Burt, C. E.</span></p> + +<p class="i2">1931. A study of the teiid lizards of the genus +<i>Cnemidophorus</i> with special reference to their phylogenetic +relationships. Bull. U. S. Nat. Mus., No. 154, pp. viii + +286.</p> + +<p><span class="smcap">Contreras A., A.</span></p> + +<p class="i2">1942. Mapa de las provincias climatologias de la Republica +Mexicana. México, D. F., Dirección de Geografía, Meterología +e Hidrología, pp. i-xxviii, tables 1-54, 2 maps.</p> + +<p><span class="smcap">Cooke, C. W.</span></p> + +<p class="i2">1945. Geology of Florida. Florida Geol. Surv., Geol. Bull., +No. 29:1-339.</p> + +<p><span class="smcap">Dorf, E.</span></p> + +<p class="i2">1959. Climatic changes of the past and present. Contrib. +Mus. Paleo. Univ. Michigan, vol. 13 (8):181-210.</p> + +<p><span class="smcap">Duellman, W. E.</span></p> + +<p class="i2">1956. The frogs of the hylid genus <i>Phrynohyas</i> Fitzinger, +1843. Miscl. Publ. Mus. Zool. Univ. Michigan, No. 96:1-47, +pls. 1-6.</p> + +<p class="i2">1958a. A monographic study of the colubrid snake genus +<i>Leptodeira</i>. Bull. Amer. Mus. Nat. Hist., vol. 114:1-152, +pls. 1-31.</p> + +<p class="i2">1958b. A preliminary analysis of the herpetofauna of Colima, +Mexico. Occas. Papers Mus. Zool. Univ. Michigan, No. +589:1-22.</p> + +<p class="i2">1958c. A review of the frogs of the genus <i>Syrrhophus</i> in +western Mexico. Occas. Papers Mus. Zool. Univ. Michigan, No. +594:1-15, pls. 1-3.</p> + +<p><span class="smcap">Duellman, W. E.</span>, and <span class="smcap">Schwartz, A.</span></p> + +<p class="i2">1958. Amphibians and reptiles of southern Florida. Bull. +Florida State Mus., vol. 3 (5):181-324.</p> + +<p><span class="smcap">Dunn, E. R.</span></p> + +<p class="i2">1942. The American caecilians. Bull. Mus. Comp. Zool., vol. +91 (6):439-540.</p> + +<p><span class="smcap">Durham, J. W.</span>, <span class="smcap">Arellano, A. R. V.</span>, and <span class="smcap">Peck, Jr., J. H.</span></p> + +<p class="i2">1952. No Cenozoic Tehuantepec seaways. Bull. Geol. Soc. +Amer., vol. 63:1245.</p> + +<p><span class="smcap">Firschein, I. L.</span></p> + +<p class="i2">1950. A new toad from Mexico with a redefinition of the +<i>cristatus</i> group. Copeia, 1950 (2):81-87, pl. 1.</p> + +<p class="i2">1954. Definition of some little-understood members of the +leptodactylid genus <i>Syrrhophus</i>, with a description of a +new species. Copeia, 1954 (1):48-58.</p> + +<p><span class="smcap">Firschein, I. L.</span>, and <span class="smcap">Smith, H. M.</span></p> + +<p class="i2">1957. A high-crested race of toad (<i>Bufo valliceps</i>) and +other noteworthy reptiles and amphibians from southern +Mexico. Herpetologica, vol. 13 (3):219-222.</p> + +<p><span class="smcap">Fugler, C. M.</span>, and <span class="smcap">Webb, R. G.</span></p> + +<p class="i2">1957. Some noteworthy reptiles and amphibians from the +states of Oaxaca and Veracruz. Herpetologica, vol. 13 +(2):103-108.</p> + +<p><span class="smcap">Gloyd, H. K.</span></p> + +<p class="i2">1940. The rattlesnakes, genera <i>Sistrurus</i> and <i>Crotalus</i>. +Chicago Acad. Sci. Special Publ., No. 4, vii + 266 pp., pls. +1-31.</p><p><span class="pagenum"><a name="Page_70" id="Page_70">[Pg 70]</a></span></p> + +<p><span class="smcap">Goin, C. J.</span></p> + +<p class="i2">1958. Comments upon the origin of the herpetofauna of +Florida. Quart. Jour. Florida Acad. Sci., vol. 21 (1):61-70.</p> + +<p><span class="smcap">Goldman, E. A.</span></p> + +<p class="i2">1951. Biological investigations in Mexico. Smithsonian Misc. +Publ., vol. 115, xiii + 476 pp., pls. 1-71.</p> + +<p><span class="smcap">Goodnight, C. J.</span>, and <span class="smcap">Goodnight, M. L.</span></p> + +<p class="i2">1956. Some observations in a tropical rain forest in +Chiapas, Mexico. Ecology, vol. 37 (1):139-150.</p> + +<p><span class="smcap">Griscom, L.</span></p> + +<p class="i2">1932. The distribution of bird-life in Guatemala. Bull. +Amer. Mus. Nat. Hist., vol. 64:1-439.</p> + +<p class="i2">1950. Distribution and origin of the birds of Mexico. Bull. +Mus. Comp. Zool., vol. 103:341-382.</p> + +<p><span class="smcap">Hartweg, N.</span>, and <span class="smcap">Oliver, J. A.</span></p> + +<p class="i2">1940. A contribution to the herpetology of the Isthmus of +Tehuantepec. IV. An annotated list of the amphibians and +reptiles collected on the Pacific slope during the summer of +1936. Misc. Publ. Mus. Zool. Univ. Michigan, No. 47:1-31.</p> + +<p><span class="smcap">Hubbell, T. H.</span></p> + +<p class="i2">1954. Relationships and distribution of <i>Mycotrupes</i>. In The +burrowing beetles of the genus <i>Mycotrupes</i>, Olson, A. L., +Hubbell, T. H., and Howden, H. F. Misc. Publ. Mus. Zool. +Univ. Michigan, No. 84:1-59, pls. 1-8.</p> + +<p><span class="smcap">Hutchinson, G. E.</span>, <span class="smcap">Patrick, R.</span>, and <span class="smcap">Deevey, E. S.</span></p> + +<p class="i2">1956. Sediments of Lake Patzcuaro, Michoacán, Mexico. Bull. +Geol. Soc. Amer., vol. 67:1491-1504.</p> + +<p><span class="smcap">Langebartel, D. A.</span>, and <span class="smcap">Smith, P. W.</span></p> + +<p class="i2">1959. Noteworthy records of amphibians and reptiles from +eastern Mexico. Herpetologica, vol. 15 (1):27-29.</p> + +<p><span class="smcap">Martin, P. S.</span></p> + +<p class="i2">1958. Pleistocene ecology and biogeography of North America. +Zoogeography. Amer. Assoc. Advanc. Sci., Publ. No. +51:375-420.</p> + +<p><span class="smcap">Martin, P. S.</span>, and <span class="smcap">Harrell, B. E.</span></p> + +<p class="i2">1957. The Pleistocene history of temperate biotas in Mexico +and eastern United States. Ecology, vol. 38:468-480.</p> + +<p><span class="smcap">Oliver, J. A.</span></p> + +<p class="i2">1948. The relationships and zoogeography of the genus +<i>Thalerophis</i> Oliver. Bull. Amer. Mus. Nat. Hist., vol. +92:157-280, pls. 16-19.</p> + +<p><span class="smcap">Olson, E. C.</span>, and <span class="smcap">McGrew, P. O.</span></p> + +<p class="i2">1941. Mammalian fauna from the Pliocene of Honduras. Bull. +Geol. Soc. Amer., vol. 52:1219-1244.</p> + +<p><span class="smcap">Ruthven, A. G.</span></p> + +<p class="i2">1912. The amphibians and reptiles collected by the +University of Michigan—Walker Expedition in southern Vera +Cruz, Mexico. Zool. Jahrbuch, vol. 32 (4):295-332, pls. +6-11.</p> + +<p><span class="smcap">Schuchert, C.</span></p> + +<p class="i2">1935. Historical geology of the Antillean-Caribbean region. +New York, xxvi + 811 pp.</p> + +<p><span class="smcap">Sears, P. B.</span>, <span class="smcap">Foreman, F.</span>, and <span class="smcap">Clisby, K. H.</span></p> + +<p class="i2">1955. Palynology in southern North America. Bull. Geol. Soc. +Amer., vol. 66:471-530.</p> + +<p><span class="smcap">Shannon, F. A.</span>, and <span class="smcap">Werler, J.</span></p> + +<p class="i2">1955. Notes on amphibians of the Los Tuxtlas Range of +Veracruz, Mexico. Trans. Kansas Acad. Sci., vol. 58 +(3):360-386.</p> + +<p><span class="smcap">Shreve, B.</span></p> + +<p class="i2">1957. Reptiles and amphibians from the Selva Lacandona. <i>In</i> +Biological Investigations in the Selva Lacandona, Chiapas, +Mexico. Paynter, R. A. (editor). Bull. Mus. Comp. Zool., +vol. 116 (4):193-298.</p><p><span class="pagenum"><a name="Page_71" id="Page_71">[Pg 71]</a></span></p> + +<p><span class="smcap">Smith, H. M.</span></p> + +<p class="i2">1947. Notes on Mexican amphibians and reptiles. Jour. +Washington Acad. Sci., vol. 37:408-412.</p> + +<p><span class="smcap">Smith, H. M.</span>, and <span class="smcap">Laufe, L. E.</span></p> + +<p class="i2">1946. A summary of Mexican Lizards of the genus <i>Ameiva</i>. +Univ. Kansas Sci. Bull., vol. 31 (2):7-73.</p> + +<p><span class="smcap">Smith, H. M.</span>, and <span class="smcap">Taylor, E. H.</span></p> + +<p class="i2">1950. An annotated checklist and key to the reptiles of +Mexico exclusive of the snakes. Bull. U. S. Natl. Mus., No. +199, v + 253 pp.</p> + +<p><span class="smcap">Stirton, R. A.</span></p> + +<p class="i2">1954. Late Miocene mammals from Oaxaca, Mexico. Amer. Jour. +Sci., No. 252:634-638.</p> + +<p><span class="smcap">Stuart, L. C.</span></p> + +<p class="i2">1935. A contribution to a knowledge of the herpetology of a +portion of the savanna region of central Petén, Guatemala. +Misc. Publ. Mus. Zool. Univ. Michigan, No. 29:1-56, pls. +1-4.</p> + +<p class="i2">1941. Studies of Neotropical Colubridae VIII. A revision of +the genus <i>Dryadophis</i> Stuart, 1939. Misc. Publ. Mus. Zool. +Univ. Michigan, No. 49:1-106, pls. 1-4.</p> + +<p class="i2">1951. The herpetofauna of the Guatemalan Plateau, with +special reference to its distribution on the southwestern +highlands. Contrib. Lab. Vertebrate Biol., Univ. Michigan, +No. 49:1-71, pls. 1-7.</p> + +<p class="i2">1954. A description of a subhumid corridor across northern +Central America, with comments on its herpetofaunal +indicators. Contrib. Lab. Vertebrate Biol., Univ. Michigan, +No. 65:1-26, pls. 1-6.</p> + +<p class="i2">1958. A study of the herpetofauna of the Uaxactun-Tikal area +of northern El Petén, Guatemala. Contrib. Lab. Vertebrate +Biol., Univ. Michigan, No. 75:1-30.</p> + +<p><span class="smcap">Taylor, E. H.</span></p> + +<p class="i2">1940. New species of Mexican Anura. Univ. Kansas Sci. Bull., +vol. 26 (11):385-405.</p> + +<p class="i2">1941. New amphibians from the Hobart M. Smith Mexican +collections. Univ. Kansas Sci. Bull., vol. 27 (8):141-167.</p> + +<p class="i2">1942. The frog genus <i>Diaglena</i> with a description of a new +species. Univ. Kansas Sci. Bull., vol. 28 (4):57-65.</p> + +<p class="i2">1943a. A new <i>Hylella</i> from Mexico. Proc. Biol. Soc. +Washington, vol. 56:49-52.</p> + +<p class="i2">1943b. Herpetological novelties from Mexico. Univ. Kansas +Sci. Bull., vol. 29 (8):343-361.</p> + +<p><span class="smcap">Taylor, E. H.</span>, and <span class="smcap">Smith, H. M.</span></p> + +<p class="i2">1945. Summary of the collections of amphibians made in +Mexico under the Walter Rathbone Bacon Traveling +Scholarship. Proc. U. S. Natl. Mus., vol. 95:521-613, pls. +18-32.</p> + +<p><span class="smcap">Trask, P. D.</span>, <span class="smcap">Phleger, F. B.</span>, and <span class="smcap">Stetson, H. C.</span></p> + +<p class="i2">1947. Recent changes in sedimentation in the Gulf of Mexico. +Science, vol. 106:460-461.</p> + +<p><span class="smcap">Weyl, R.</span></p> + +<p class="i2">1955. Vestigios de una glaciacion del Pleistoceno en la +Cordillera de Talamanca, Costa Rica, A. C. Informe +Trimestral, Inst. Geog. de Costa Rica, pp. 9-32.</p> + +<p><span class="smcap">White, S. E.</span></p> + +<p class="i2">1956. Probable substages of glaciation on Ixtaccihuatl, +Mexico. Jour. Geol., vol. 64:289-295.</p> + +<p><span class="smcap">Williams, L.</span></p> + +<p class="i2">1939. Arboles y arbustos del Istmos de Tehuantepec, Mexico. +Lilloa., vol. 4:137-171.</p> + +<p> +<span style="margin-left: 1em;"><i>Transmitted May 23, 1960.</i></span> +</p> +<hr class="full" /> + + +<p><span class="pagenum"><a name="Page_72" id="Page_72">[Pg 72]</a></span></p> +<h2><a name="UNIVERSITY_OF_KANSAS_PUBLICATIONS_MUSEUM_OF_NATURAL_HISTORY" id="UNIVERSITY_OF_KANSAS_PUBLICATIONS_MUSEUM_OF_NATURAL_HISTORY"></a>UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY</h2> + + +<p>Institutional libraries interested in publications exchange may obtain +this series by addressing the Exchange Librarian, University of Kansas +Library, Lawrence, Kansas. Copies for individuals, persons working in a +particular field of study, may be obtained by addressing instead the +Museum of Natural History, University of Kansas, Lawrence, Kansas. +There is no provision for sale of this series by the University +Library, which meets institutional requests, or by the Museum of +Natural History, which meets the requests of individuals. However, when +individuals request copies from the Museum, 25 cents should be +included, for each separate number that is 100 pages or more in length, +for the purpose of defraying the costs of wrapping and mailing.</p> + +<p>* An asterisk designates those numbers of which the Museum's supply +(not the Library's supply) is exhausted. Numbers published to date, in +this series, are as follows:</p> + +<p> +<span style="margin-left: 0.5em;">Vol. 1. Nos. 1-26 and index. Pp. 1-638, 1946-1950.</span><br /> +<br /> +*Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest.<br /> +<span style="margin-left: 6.5em;">Pp. 1-444, 140 figures in text. April 9, 1948.</span><br /> +<br /> +<span style="margin-left: 0.5em;">Vol. 3. *1. The avifauna of Micronesia, its origin, evolution, and</span><br /> +<span style="margin-left: 6.5em;">distribution. By Rollin H. Baker. Pp. 1-359, 16 figures</span><br /> +<span style="margin-left: 6.5em;">in text. June 12, 1951.</span><br /> +<br /> +<span style="margin-left: 4.5em;">*2. A quantitative study of the nocturnal migration of birds.</span><br /> +<span style="margin-left: 6.5em;">By George H. Lowery, Jr. Pp. 361-472, 47 figures in text.</span><br /> +<span style="margin-left: 6.5em;">June 29, 1951.</span><br /> +<br /> +<span style="margin-left: 5em;">3. Phylogeny of the waxwings and allied birds. By M. Dale</span><br /> +<span style="margin-left: 6.5em;">Arvey. Pp. 473-530, 49 figures in text, 13 tables.</span><br /> +<span style="margin-left: 6.5em;">October 10, 1951.</span><br /> +<br /> +<span style="margin-left: 5em;">4. Birds from the state of Veracruz, Mexico. By George H.</span><br /> +<span style="margin-left: 6.5em;">Lowery, Jr., and Walter W. Dalquest. Pp. 531-649, 7</span><br /> +<span style="margin-left: 6.5em;">figures in text, 2 tables. October 10, 1951.</span><br /> +<br /> +<span style="margin-left: 5em;">Index. Pp. 651-681.</span><br /> +<br /> +*Vol. 4. (Complete) American weasels. By E. Raymond Hall. Pp. 1-466,<br /> +<span style="margin-left: 6.5em;">41 plates, 31 figures in text. December 27, 1951.</span><br /> +<br /> +<span style="margin-left: 0.5em;">Vol. 5. Nos. 1-37 and index. Pp. 1-676, 1951-1953.</span><br /> +<br /> +*Vol. 6. (Complete) Mammals of Utah, <i>taxonomy and distribution</i>. By<br /> +<span style="margin-left: 6.5em;">Stephen D. Durrant. Pp. 1-549, 91 figures in text, 80</span><br /> +<span style="margin-left: 6.5em;">tables. August 10, 1952.</span><br /> +<br /> +<span style="margin-left: 0.5em;">Vol. 7. *1. Mammals of Kansas. By E. Lendell Cockrum. Pp. 1-303, 73</span><br /> +<span style="margin-left: 6.5em;">figures in text, 37 tables. August 25, 1952.</span><br /> +<br /> +<span style="margin-left: 5em;">2. Ecology of the opossum on a natural area in northeastern</span><br /> +<span style="margin-left: 6.5em;">Kansas. By Henry S. Fitch and Lewis L. Sandidge. Pp.</span><br /> +<span style="margin-left: 6.5em;">305-338, 5 figures in text. August 24, 1953.</span><br /> +<br /> +<span style="margin-left: 5em;">3. The silky pocket mice (Perognathus flavus) of Mexico. By</span><br /> +<span style="margin-left: 6.5em;">Rollin H. Baker. Pp. 339-347, 1 figure in text. February</span><br /> +<span style="margin-left: 6.5em;">15, 1954.</span><br /> +<br /> +<span style="margin-left: 5em;">4. North American jumping mice (Genus Zapus). By Philip H.</span><br /> +<span style="margin-left: 6.5em;">Krutzsch. Pp. 349-472, 47 figures in text, 4 tables.</span><br /> +<span style="margin-left: 6.5em;">April 21, 1954.</span><br /> +<br /> +<span style="margin-left: 5em;">5. Mammals from Southeastern Alaska. By Rollin H. Baker and</span><br /> +<span style="margin-left: 6.5em;">James S. Findley. Pp. 473-477. April 21, 1954.</span><br /> +<br /> +<span style="margin-left: 5em;">6. Distribution of Some Nebraskan Mammals. By J. Knox Jones,</span><br /> +<span style="margin-left: 6.5em;">Jr. Pp. 479-487. April 21, 1954.</span><br /> +<br /> +<span style="margin-left: 5em;">7. Subspeciation in the montane meadow mouse. Microtus</span><br /> +<span style="margin-left: 6.5em;">montanus, in Wyoming and Colorado. By Sydney Anderson.</span><br /> +<span style="margin-left: 6.5em;">Pp. 489-506, 2 figures in text. July 23, 1954.</span><br /> +<br /> +<span style="margin-left: 5em;">8. A new subspecies of bat (Myotis velifer) from southeastern</span><br /> +<span style="margin-left: 6.5em;">California and Arizona. By Terry A. Vaughan. Pp. 507-512.</span><br /> +<span style="margin-left: 6.5em;">July 23, 1954.</span><br /> +<br /> +<span style="margin-left: 5em;">9. Mammals of the San Gabriel mountains of California. By</span><br /> +<span style="margin-left: 6.5em;">Terry A. Vaughan. Pp. 513-582, 1 figure in text, 12</span><br /> +<span style="margin-left: 6.5em;">tables. November 15, 1954.</span><br /> +<br /> +<span style="margin-left: 4.5em;">10. A new bat (Genus Pipistrellus) from northeastern Mexico.</span><br /> +<span style="margin-left: 6.5em;">By Rollin H. Baker. Pp. 583-586. November 15, 1954.</span><br /> +<br /> +<span style="margin-left: 4.5em;">11. A new subspecies of pocket mouse from Kansas. By E.</span><br /> +<span style="margin-left: 6.5em;">Raymond Hall. Pp. 587-590. November 15, 1954.</span><br /> +<br /> +<span style="margin-left: 4.5em;">12. Geographic variation in the pocket gopher, Cratogeomys</span><br /> +<span style="margin-left: 6.5em;">castanops, in Coahuila, Mexico. By Robert J. Russell and</span><br /> +<span style="margin-left: 6.5em;">Rollin H. Baker. Pp. 591-608. March 15, 1955.</span><br /> +<br /> +<span style="margin-left: 4.5em;">13. A new cottontail (Sylvilagus floridanus) from northeastern</span><br /> +<span style="margin-left: 6.5em;">Mexico. By Rollin H. Baker. Pp. 609-612. April 8, 1955.</span><br /> +<br /> +<span style="margin-left: 4.5em;">14. Taxonomy and distribution of some American shrews. By</span><br /> +<span style="margin-left: 6.5em;">James S. Findley. Pp. 613-618. June 10, 1955.</span><br /> +<br /> +<span style="margin-left: 4.5em;">15. The pigmy woodrat, Neotoma goldmani, its distribution and</span><br /> +<span style="margin-left: 6.5em;">systematic position. By Dennis G. Rainey and Rollin H.</span><br /> +<span style="margin-left: 6.5em;">Baker. Pp. 619-624. 2 figures in text. June 10, 1955.</span><br /> +<br /> +<span style="margin-left: 5em;">Index. Pp. 625-651.</span><br /> +<br /> +<span style="margin-left: 0.5em;">Vol. 8. 1. Life history and ecology of the five-lined skink, Eumeces</span><br /> +<span style="margin-left: 6.5em;">fasciatus. By Henry S. Fitch. Pp. 1-156, 26 figures in</span><br /> +<span style="margin-left: 6.5em;">text. September 1, 1954.</span><br /> +<br /> +<span style="margin-left: 5em;">2. Myology and serology of the Avian Family Fringillidae, a</span><br /> +<span style="margin-left: 6.5em;">taxonomic study. By William B. Stallcup. Pp. 157-211, 23</span><br /> +<span style="margin-left: 6.5em;">figures in text, 4 tables. November 15, 1954.</span><br /> +<br /> +<span style="margin-left: 5em;">3. An ecological study of the collared lizard (Crotaphytus</span><br /> +<span style="margin-left: 6.5em;">collaris). By Henry S. Fitch. Pp. 213-274, 10 figures in</span><br /> +<span style="margin-left: 6.5em;">text. February 10, 1956.</span><br /> +<br /> +<span style="margin-left: 5em;">4. A field study of the Kansas ant-eating frog, Gastrophryne</span><br /> +<span style="margin-left: 6.5em;">olivacea. By Henry S. Fitch. Pp. 275-306, 9 figures in</span><br /> +<span style="margin-left: 6.5em;">text. February 10, 1956.</span><br /> +<br /> +<span style="margin-left: 5em;">5. Check-list of the birds of Kansas. By Harrison B. Tordoff.</span><br /> +<span style="margin-left: 6.5em;">Pp. 307-359, 1 figure in text. March 10, 1956.</span><br /> +<br /> +<span style="margin-left: 5em;">6. A population study of the prairie vole (Microtus</span><br /> +<span style="margin-left: 6.5em;">ochrogaster) in northeastern Kansas. By Edwin P. Martin.</span><br /> +<span style="margin-left: 6.5em;">Pp. 361-416, 19 figures in text. April 2, 1956.</span><br /> +<br /> +<span style="margin-left: 5em;">7. Temperature responses in free-living amphibians and</span><br /> +<span style="margin-left: 6.5em;">reptiles of northeastern Kansas. By Henry S. Fitch. Pp.</span><br /> +<span style="margin-left: 6.5em;">417-476, 10 figures in text, 6 tables. June 1, 1956.</span><br /> +<br /> +<span style="margin-left: 5em;">8. Food of the crow, Corvus brachyrhynchos Brehm, in</span><br /> +<span style="margin-left: 6.5em;">south-central Kansas. By Dwight Platt. Pp. 477-498, 4</span><br /> +<span style="margin-left: 6.5em;">tables. June 8, 1956.</span><br /> +<br /> +<span style="margin-left: 5em;">9. Ecological observations on the woodrat, Neotoma floridana.</span><br /> +<span style="margin-left: 6.5em;">By Henry S. Fitch and Dennis G. Rainey. Pp. 499-533, 3</span><br /> +<span style="margin-left: 6.5em;">figures in text. June 12, 1956.</span><br /> +<br /> +<span style="margin-left: 4.5em;">10. Eastern woodrat, Neotoma floridana: Life history and</span><br /> +<span style="margin-left: 6.5em;">ecology. By Dennis G. Rainey. Pp. 535-646, 12 plates, 13</span><br /> +<span style="margin-left: 6.5em;">figures in text. August 15, 1956.</span><br /> +<br /> +<span style="margin-left: 4.5em;">Index. Pp. 647-675.</span><br /> +<br /> +<span style="margin-left: 0.5em;">Vol. 9. 1. Speciation of the wandering shrew. By James S. Findley.</span><br /> +<span style="margin-left: 6.5em;">Pp. 1-68, 18 figures in text. December 10, 1955.</span><br /> +<br /> +<span style="margin-left: 5em;">2. Additional records and extensions of ranges of mammals</span><br /> +<span style="margin-left: 6.5em;">from Utah. By Stephen D. Durrant, M. Raymond Lee, and</span><br /> +<span style="margin-left: 6.5em;">Richard M. Hansen. Pp. 69-80. December 10, 1955.</span><br /> +<br /> +<span style="margin-left: 5em;">3. A new long-eared myotis (Myotis evotis) from northeastern</span><br /> +<span style="margin-left: 6.5em;">Mexico. By Rollin H. Baker and Howard J. Stains. Pp.</span><br /> +<span style="margin-left: 6.5em;">81-84. December 10, 1955.</span><br /> +<br /> +<span style="margin-left: 5em;">4. Subspeciation in the meadow mouse, Microtus pennsylvanicus,</span><br /> +<span style="margin-left: 6.5em;">in Wyoming. By Sydney Anderson. Pp. 85-104, 2 figures in</span><br /> +<span style="margin-left: 6.5em;">text. May 10, 1956.</span><br /> +<br /> +<span style="margin-left: 5em;">5. The condylarth genus Ellipsodon. By Robert W. Wilson.</span><br /> +<span style="margin-left: 6.5em;">Pp. 105-116, 6 figures in text. May 19, 1956.</span><br /> +<br /> +<span style="margin-left: 5em;">6. Additional remains of the multituberculate genus</span><br /> +<span style="margin-left: 6.5em;">Eucosmodon. By Robert W. Wilson. Pp. 117-123, 10 figures</span><br /> +<span style="margin-left: 6.5em;">in text. May 19, 1956.</span><br /> +<br /> +<span style="margin-left: 5em;">7. Mammals of Coahuila, Mexico. By Rollin H. Baker. Pp.</span><br /> +<span style="margin-left: 6.5em;">125-335, 75 figures in text. June 15, 1956.</span><br /> +<br /> +<span style="margin-left: 5em;">8. Comments on the taxonomic status of Apodemus peninsulae,</span><br /> +<span style="margin-left: 6.5em;">with description of a new subspecies from North China. By</span><br /> +<span style="margin-left: 6.5em;">J. Knox Jones, Jr. Pp. 337-346, 1 figure in text, 1</span><br /> +<span style="margin-left: 6.5em;">table. August 15, 1956.</span><br /> +<br /> +<span style="margin-left: 5em;">9. Extensions of known ranges of Mexican bats. By Sydney</span><br /> +<span style="margin-left: 6.5em;">Anderson. Pp. 347-351. August 15, 1956.</span><br /> +<br /> +<span style="margin-left: 4.5em;">10. A new bat (Genus Leptonycteris) from Coahuila. By Howard</span><br /> +<span style="margin-left: 6.5em;">J. Stains. Pp. 353-356. January 21, 1957.</span><br /> +<br /> +<span style="margin-left: 4.5em;">11. A new species of pocket gopher (Genus Pappogeomys) from</span><br /> +<span style="margin-left: 6.5em;">Jalisco, Mexico. By Robert J. Russell. Pp. 357-361.</span><br /> +<span style="margin-left: 6.5em;">January 21, 1957.</span><br /> +<br /> +<span style="margin-left: 4.5em;">12. Geographic variation in the pocket gopher, Thomomys</span><br /> +<span style="margin-left: 6.5em;">bottae, in Colorado. By Phillip M. Youngman. Pp. 363-384,</span><br /> +<span style="margin-left: 6.5em;">7 figures in text. February 21, 1958.</span><br /> +<br /> +<span style="margin-left: 4.5em;">13. New bog lemming (genus Synaptomys) from Nebraska. By J.</span><br /> +<span style="margin-left: 6.5em;">Knox Jones, Jr. Pp. 385-388. May 12, 1958.</span><br /> +<br /> +<span style="margin-left: 4.5em;">14. Pleistocene bats from San Josecito Cave, Nuevo León,</span><br /> +<span style="margin-left: 6.5em;">México. By J. Knox Jones, Jr. Pp. 389-396. December 19,</span><br /> +<span style="margin-left: 6.5em;">1958.</span><br /> +<br /> +<span style="margin-left: 4.5em;">15. New Subspecies of the rodent Baiomys from Central America.</span><br /> +<span style="margin-left: 6.5em;">By Robert L. Packard. Pp. 397-404. December 19, 1958.</span><br /> +<br /> +<span style="margin-left: 4.5em;">16. Mammals of the Grand Mesa, Colorado. By Sydney Anderson.</span><br /> +<span style="margin-left: 6.5em;">Pp. 405-414, 1 figure in text. May 20, 1959.</span><br /> +<br /> +<span style="margin-left: 4.5em;">17. Distribution, variation, and relationships of the montane</span><br /> +<span style="margin-left: 6.5em;">vole, Microtus montanus. By Emil K. Urban. Pp. 415-511.</span><br /> +<span style="margin-left: 6.5em;">12 figures in text, 2 tables. August 1, 1959.</span><br /> +<br /> +<span style="margin-left: 4.5em;">18. Conspecificity of two pocket mice, Perognathus goldmani</span><br /> +<span style="margin-left: 6.5em;">and P. artus. By E. Raymond Hall and Marilyn Bailey</span><br /> +<span style="margin-left: 6.5em;">Ogilvie. Pp. 513-518, 1 map. January 14, 1960.</span><br /> +<br /> +<span style="margin-left: 4.5em;">19. Records of harvest mice, Reithrodontomys, from Central</span><br /> +<span style="margin-left: 6.5em;">America, with description of a new subspecies from</span><br /> +<span style="margin-left: 6.5em;">Nicaragua. By Sydney Anderson and J. Knox Jones, Jr. Pp.</span><br /> +<span style="margin-left: 6.5em;">519-529. January 14, 1960.</span><br /> +<br /> +<span style="margin-left: 4.5em;">20. Small carnivores from San Josecito Cave (Pleistocene),</span><br /> +<span style="margin-left: 6.5em;">Nuevo León, México. By E. Raymond Hall. Pp. 531-538, 1</span><br /> +<span style="margin-left: 6.5em;">figure in text. January 14, 1960.</span><br /> +<br /> +<span style="margin-left: 4.5em;">21. Pleistocene pocket gophers from San Josecito Cave, Nuevo</span><br /> +<span style="margin-left: 6.5em;">León, México. By Robert J. Russell. Pp. 539-548, 1 figure</span><br /> +<span style="margin-left: 6.5em;">in text. January 14, 1960.</span><br /> +<br /> +<span style="margin-left: 4.5em;">22. Review of the insectivores of Korea. By J. Knox Jones,</span><br /> +<span style="margin-left: 6.5em;">Jr., and David H. Johnson. Pp. 549-578. February 23, 1960.</span><br /> +<br /> +<span style="margin-left: 4.5em;">23. Speciation and evolution of the pygmy mice, genus Baiomys.</span><br /> +<span style="margin-left: 6.5em;">By Robert L. Packard. Pp. 579-670, 4 plates, 12 figures in</span><br /> +<span style="margin-left: 6.5em;">text. June 16, 1960.</span><br /> +<br /> +<span style="margin-left: 4.5em;">Index Pp. 671-690.</span><br /> +<br /> +Vol. 10. 1. Studies of birds killed in nocturnal migration. By<br /> +<span style="margin-left: 6.5em;">Harrison B. Tordoff and Robert M. Mengel. Pp. 1-44, 6</span><br /> +<span style="margin-left: 6.5em;">figures in text, 2 tables. September 12, 1956.</span><br /> +<br /> +<span style="margin-left: 5em;">2. Comparative breeding behavior of Ammospiza caudacuta and</span><br /> +<span style="margin-left: 6.5em;">A. maritima. By Glen E. Woolfenden. Pp. 45-75, 6 plates,</span><br /> +<span style="margin-left: 6.5em;">1 figure. December 20, 1956.</span><br /> +<br /> +<span style="margin-left: 5em;">3. The forest habitat of the University of Kansas Natural</span><br /> +<span style="margin-left: 6.5em;">History Reservation. By Henry S. Fitch and Ronald R.</span><br /> +<span style="margin-left: 6.5em;">McGregor. Pp. 77-127, 2 plates, 7 figures in text, 4</span><br /> +<span style="margin-left: 6.5em;">tables. December 31, 1956.</span><br /> +<br /> +<span style="margin-left: 5em;">4. Aspects of reproduction and development in the prairie</span><br /> +<span style="margin-left: 6.5em;">vole (Microtus ochrogaster). By Henry S. Fitch. Pp.</span><br /> +<span style="margin-left: 6.5em;">129-161, 8 figures in text, 4 tables. December 19, 1957.</span><br /> +<br /> +<span style="margin-left: 5em;">5. Birds found on the Arctic slope of northern Alaska. By</span><br /> +<span style="margin-left: 6.5em;">James W. Bee. Pp. 163-211, pls. 9-10, 1 figure in text.</span><br /> +<span style="margin-left: 6.5em;">March 12, 1958.</span><br /> +<br /> +<span style="margin-left: 5em;">6. The wood rats of Colorado: distribution and ecology. By</span><br /> +<span style="margin-left: 6.5em;">Robert B. Finley, Jr. Pp. 213-552, 34 plates, 8 figures</span><br /> +<span style="margin-left: 6.5em;">in text, 35 tables. November 7, 1958.</span><br /> +<br /> +<span style="margin-left: 5em;">7. Home ranges and movements of the eastern cottontail in</span><br /> +<span style="margin-left: 6.5em;">Kansas. By Donald W. Janes. Pp. 553-572, 4 plates, 3</span><br /> +<span style="margin-left: 6.5em;">figures in text. May 4, 1959.</span><br /> +<br /> +<span style="margin-left: 5em;">8. Natural history of the salamander, Aneides hardyi. By</span><br /> +<span style="margin-left: 6.5em;">Richard F. Johnston and Schad Gerhard. Pp. 573-585.</span><br /> +<span style="margin-left: 6.5em;">October 8, 1959.</span><br /> +<br /> +<span style="margin-left: 5em;">9. A new subspecies of lizard, Cnemidophorus sacki, from</span><br /> +<span style="margin-left: 6.5em;">Michoacán, México. By William E. Duellman. Pp. 587-598,</span><br /> +<span style="margin-left: 6.5em;">2 figures in text. May 2, 1960.</span><br /> +<br /> +<span style="margin-left: 4.5em;">10. A taxonomic study of the Middle American Snake, Pituophis</span><br /> +<span style="margin-left: 6.5em;">deppei. By William E. Duellman. Pp. 599-610, 1 plate, 1</span><br /> +<span style="margin-left: 6.5em;">figure in text. May 2, 1960.</span><br /> +<br /> +<span style="margin-left: 4.5em;">Index Pp. 611-626.</span><br /> +<br /> +Vol. 11. 1. The systematic status of the colubrid snake, Leptodeira<br /> +<span style="margin-left: 6.5em;">discolor Günther. By William E. Duellman. Pp. 1-9, 4</span><br /> +<span style="margin-left: 6.5em;">figs. July 14, 1958.</span><br /> +<br /> +<span style="margin-left: 5em;">2. Natural history of the six-lined racerunner, Cnemidophorus</span><br /> +<span style="margin-left: 6.5em;">sexlineatus. By Henry S. Fitch. Pp. 11-62, 9 figs., 9</span><br /> +<span style="margin-left: 6.5em;">tables. September 19, 1958.</span><br /> +<br /> +<span style="margin-left: 5em;">3. Home ranges, territories, and seasonal movements of</span><br /> +<span style="margin-left: 6.5em;">vertebrates of the Natural History Reservation. By Henry</span><br /> +<span style="margin-left: 6.5em;">S. Fitch. Pp. 63-326, 6 plates, 24 figures in text, 3</span><br /> +<span style="margin-left: 6.5em;">tables. December 12, 1958.</span><br /> +<br /> +<span style="margin-left: 5em;">4. A new snake of the genus Geophis from Chihuahua, Mexico.</span><br /> +<span style="margin-left: 6.5em;">By John M. Legler. Pp. 327-334, 2 figures in text.</span><br /> +<span style="margin-left: 6.5em;">January 28, 1959.</span><br /> +<br /> +<span style="margin-left: 5em;">5. A new tortoise, genus Gopherus, from north-central</span><br /> +<span style="margin-left: 6.5em;">Mexico. By John M. Legler. Pp. 335-343. April 24, 1959.</span><br /> +<br /> +<span style="margin-left: 5em;">6. Fishes of Chautauqua, Cowley and Elk counties, Kansas. By</span><br /> +<span style="margin-left: 6.5em;">Artie L. Metcalf. Pp. 345-400, 2 plates, 2 figures in</span><br /> +<span style="margin-left: 6.5em;">text, 10 tables. May 6, 1959.</span><br /> +<br /> +<span style="margin-left: 5em;">7. Fishes of the Big Blue River Basin, Kansas. By W. L.</span><br /> +<span style="margin-left: 6.5em;">Minckley. Pp. 401-442, 2 plates, 4 figures in text, 5</span><br /> +<span style="margin-left: 6.5em;">tables. May 8, 1959.</span><br /> +<br /> +<span style="margin-left: 5em;">8. Birds from Coahuila, México. By Emil K. Urban. Pp.</span><br /> +<span style="margin-left: 6.5em;">443-516. August 1, 1959.</span><br /> +<br /> +<span style="margin-left: 5em;">9. Description of a new softshell turtle from the</span><br /> +<span style="margin-left: 6.5em;">southeastern United States. By Robert G. Webb. Pp.</span><br /> +<span style="margin-left: 6.5em;">517-525, 2 pls., 1 figure in text, August 14, 1959.</span><br /> +<br /> +<span style="margin-left: 4.5em;">10. Natural history of the ornate box turtle, Terrapene</span><br /> +<span style="margin-left: 6.5em;">ornata ornata Agassiz. By John M. Legler. Pp. 527-669,</span><br /> +<span style="margin-left: 6.5em;">16 pls., 29 figures in text. March 7, 1960.</span><br /> +<br /> +<span style="margin-left: 4.5em;">Index will follow.</span><br /> +<br /> +Vol. 12. 1. Functional morphology of three bats: Eumops, Myotis,<br /> +<span style="margin-left: 6.5em;">Macrotus. By Terry A. Vaughan. Pp. 1-153, 4 plates, 24</span><br /> +<span style="margin-left: 6.5em;">figures in text, July 8, 1959.</span><br /> +<br /> +<span style="margin-left: 5em;">2. The ancestry of modern Amphibia: a review of the</span><br /> +<span style="margin-left: 6.5em;">evidence. By Theodore H. Eaton, Jr. Pp. 155-180, 10</span><br /> +<span style="margin-left: 6.5em;">figures in text. July 10, 1959.</span><br /> +<br /> +<span style="margin-left: 5em;">3. The baculum in microtine rodents. By Sydney Anderson.</span><br /> +<span style="margin-left: 6.5em;">Pp. 181-216, 49 figures in text. February 19, 1960.</span><br /> +<br /> +<span style="margin-left: 5em;">4. A new order of fishlike Amphibia from the Pennsylvanian</span><br /> +<span style="margin-left: 6.5em;">of Kansas. By Theodore H. Eaton, Jr., and Peggy Lou</span><br /> +<span style="margin-left: 6.5em;">Stewart. Pp. 217-240, 12 figures in text. May 2, 1960.</span><br /> +<br /> +<span style="margin-left: 5em;">More numbers will appear in volume 12.</span><br /> +<br /> +Vol. 13. 1. Five natural hybrid combinations in minnows (Cyprinidae).<br /> +<span style="margin-left: 6.5em;">By Frank B. Cross and W. L. Minckley. Pp. 1-18. June 1,</span><br /> +<span style="margin-left: 6.5em;">1960.</span><br /> +<br /> +<span style="margin-left: 5em;">2. A distributional study of the amphibians of the isthmus</span><br /> +<span style="margin-left: 6.5em;">of Tehuantepec, México. By William E. Duellman.</span><br /> +<span style="margin-left: 6.5em;">Pp. 19-72, pls. 1-8, 3 figs. August 16, 1960.</span><br /> +<br /> +<span style="margin-left: 5em;">More numbers will appear in volume 13.</span><br /> +</p> + +<hr class="full" /> + + +<p class="transnote">Transcriber's Notes<br /><br /> + + + +Page <a href="#Page_26">26</a>: Changed "19. Cosaleacaque" to "19. Cosoleacaque".<br /> +<br /> +Page <a href="#Page_30">30</a>: Changed "Brysonima crassifolia" to "Byrsonima crassifolia".<br /> +<br /> +Page <a href="#Page_34">34</a>: Changed "long. 95' 29°;" to "long. 95° 29';".<br /> +<br /> +Page <a href="#Page_35">35</a>: Changed "Matías Romera" to "Matías Romero".<br /> +<br /> +Page <a href="#Page_47">47</a>: Changed "kown" to "known".<br /> +<br /> +Pages <a href="#Page_50">50</a> and <a href="#Page_59">59</a>: Changed "axills" to "axils".<br /> +<br /> +<a href="#pl1-fig2">Plate 1, Fig. 2</a>: Changed "Veracuz" to "Veracruz".<br /> +<br /> +Page <a href="#Page_53">53</a>: Changed "valadity" to "validity".<br /> +<br /> +Page <a href="#Page_67">67</a>: Changed "refering" to "referring".<br /> +<br /> +Page <a href="#Page_68">68</a>: Changed "survided" to "survived".<br /> +<br /> +Page <a href="#Page_71">71</a>: Changed "subhimid" to "subhumid" and "Amerca" to "America".<br /> +<br /> +Moved University of Kansas <a href="#UNIVERSITY_OF_KANSAS_PUBLICATIONS_MUSEUM_OF_NATURAL_HISTORY">Publications list</a> to end of report.<br /> +Vol. 9, No. 12: Changed pages from "363-387" to "363-384".<br /> +Vol. 10, No. 10: Changed pages from "599-612" to "599-610".<br /> +</p> + + + + + + + + + + + +<pre> + + + + + +End of the Project Gutenberg EBook of A Distributional Study of the +Amphibians of the Isthmus of Tehuantepec, Mexico, by William E. 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