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Duellman. + </title> + <style type="text/css"> +/*<![CDATA[*/ + p {margin-top: .75em; text-align: justify; text-indent: 2em; margin-bottom: .75em;} + + hr { color:#000; } + .hr2 {width: 30%; height: 2px; text-align: center;} + + table {margin-left: auto; margin-right: auto; vertical-align:text-top;} + .data {text-align:center; padding-left: 4px; padding-right: 4px; white-space: nowrap;} + .bt {border-top:solid #000 1px;} + .bb {border-bottom:solid #000 1px;} + .bl {border-left:solid #000 1px;} + .btlb {border-top:solid #000 1px; border-left:solid #000 1px; border-bottom:solid #000 1px;} + + body {margin-left: 10%; margin-right: 10%; background:#fff;} + + .pagenum {position: absolute; + left: 92%; text-indent: 0em; + font-size: 0.8em; color: #808080; + text-align: right;} + + .dividend { font-size:0.65em; position:relative; top:-4px; } + .divisor { font-size:0.65em; position:relative; bottom:-2px; } + + .center {text-align: center;} + .img_left {float: left; margin-left:-10%; clear: right;} + .justify {text-align: justify;} + .text_lf {text-align: left; margin-left:0;} + .text_rt {text-align: right;} + .smcap {font-variant: small-caps;} + .smaller {font-size:0.85em;} + + .caption1 {font-weight: bold; font-size:2.50em; text-align: center;} + .caption2 {font-weight: bold; font-size:1.50em; text-align: center;} + .caption3 {font-weight: bold; font-size:1.15em; text-align: center;} + .caption3nb {font-size:1.15em; text-align: center;} + .caption4 {font-weight: bold; font-size:0.75em; text-align: center;} + .technt {background:#d0d0d0; padding: 7px; border:solid black 1px;} + + .footnote {font-size: 0.9em;} + .footnote .label {text-align: left;} + .fnanchor {vertical-align: super; font-size: .8em; text-decoration: none;} + + .toc {text-align:left; width:100%; border:1px; background:#fff;} + .pub_list td {vertical-align: top;} + .vtop {vertical-align: top;} + .descrp2 {text-align: right; float:right;} + .species {margin-left: 6em; text-indent: -2em;} + .references p {padding-left: 5em; text-indent:-3em; margin-right: 10%;} + + /*]]>*/ + </style> + </head> +<body> + + +<pre> + +The Project Gutenberg EBook of A Review of the Middle American Tree Frogs +of the Genus Ptychohyla, 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 Review of the Middle American Tree Frogs of the Genus Ptychohyla + +Author: William E. Duellman + +Release Date: February 26, 2011 [EBook #35413] +[Last updated: October 16, 2011] + +Language: English + +Character set encoding: UTF-8 + +*** START OF THIS PROJECT GUTENBERG EBOOK MIDDLE AMERICAN TREE FROGS--PTYCHOHYLA *** + + + + +Produced by Chris Curnow, Tom Cosmas, Joseph Cooper and +the Online Distributed Proofreading Team at +https://www.pgdp.net + + + + + + +</pre> + + + + + +<span class='pagenum'><a name="Page_i" id="Page_i">[Cover]</a></span> +<div class="center"> +<img src="images/cover_treefrog.jpg" width="371" height="600" border="0" alt="Cover" title="Cover" id="coverpage" /> +</div> +<p> </p> +<p> </p> + +<span class='pagenum'><a name="Page_297" id="Page_297">[Pg 297]</a></span> +<div class="center"> +<p> </p> +<img src="images/bar_double.png" width="100%" height="15" border="0" alt="double bar" /> +<div class="caption2"><div class="smcap">University of Kansas Publications<br /> +Museum of Natural History</div></div> +<hr class="hr2" /> +<br /> +<div class="caption2">Volume 15, No. 7, pp. 297-349, pls. 11-18, 7 figs.</div><br /> +<div class="center"><img src="images/bar_single.png" width="28%" height="15" title="bar" alt="bar" /> <span class="caption2">October 18, 1963</span> <img src="images/bar_single.png" width="28%" height="15" title="bar" alt="bar" /></div> +<p> </p> +<p> </p> +<p> </p> +<div class="caption1"> +A Review of the Middle American Tree Frogs<br /> +of the Genus Ptychohyla<br /> +</div> +<p> </p> +<p> </p> + +<div class="caption3"> +BY<br /> +<p> </p> + +WILLIAM E. DUELLMAN<br /> +</div> +<p> </p> +<p> </p> +<p> </p> +<p> </p> +<p> </p> +<p> </p> + +<div class="caption2"> +<span class="smcap">University of Kansas</span><br /> +<span class="smcap">Lawrence</span><br /> +1963 +<p> </p> +<p> </p> +</div> +</div> + + +<span class='pagenum'><a name="Page_298" id="Page_298">[Pg 298]</a></span> +<div class="center caption3"> +<span class="smcap">University of Kansas Publications, Museum of Natural History</span><br /> +<br /> +Editors: E. Raymond Hall, Chairman, Henry S. Fitch,<br /> +Theodore H. Eaton, Jr.<br /> +<p> </p> +<p> </p> + +Volume 15, No. 7, pp. 297-349, pls. 11-18, 7 figs.<br /> +<br /> +Published October 18, 1963<br /> +<p> </p> +<p> </p> +<p> </p> + +<span class="smcap">University of Kansas</span><br /> +Lawrence, Kansas<br /> +<br /> +</div> +<p> </p> +<p> </p> +<div class="caption4"> +PRINTED BY<br /> +JEAN M. NEIBARGER, STATE PRINTER<br /> +TOPEKA, KANSAS<br /> +1963<br /> +<img src="images/union_label.png" width="71" height="26" border="0" alt="Look for the Union Label" title="Look for the Union Label" /><br /> +29-6531<br /> +</div> +<p> </p> +<p> </p> + +<span class='pagenum'><a name="Page_299" id="Page_299">[Pg 299]</a></span> +<div class="caption2">A Review of the Middle American Tree Frogs<br /> +of the Genus Ptychohyla<br /> +</div> + +<div class="caption3">BY<br /> +WILLIAM E. DUELLMAN</div> +<p> </p> + +<a name="TOC"></a> +<div class="caption2">CONTENTS</div> +<br /> +<table class="toc" summary="Table of Contents"> +<tr><td><span class="smcap"><a href="#introduction">Introduction</a> </span></td><td style="width:5em" class="text_rt">301</td></tr> +<tr><td> <a href="#acknwldgmnts">Acknowledgments</a></td><td class="text_rt">301</td></tr> +<tr><td> <a href="#matrls_mthds">Materials and Methods</a></td><td class="text_rt">302</td></tr> +<tr><td><span class="smcap"><a href="#analysis_of_data">Analysis of Data</a></span></td><td class="text_rt">303</td></tr> +<tr><td> <a href="#extrnl_morph">External Morphology</a></td><td class="text_rt">303</td></tr> +<tr><td> <a href="#color_pattern">Color and Pattern</a></td><td class="text_rt">307</td></tr> +<tr><td> <a href="#osteology">Osteology</a></td><td class="text_rt">307</td></tr> +<tr><td> <a href="#tadpoles">Tadpoles</a></td><td class="text_rt">310</td></tr> +<tr><td> <a href="#breeding_call">Breeding Call</a></td><td class="text_rt">312</td></tr> +<tr><td><span class="smcap"><a href="#systm_accs">Systematic Accounts</a></span></td><td class="text_rt">314</td></tr> +<tr><td> <a href="#pytch_taylor"><i>Ptychohyla</i> Taylor, 1944</a></td><td class="text_rt">314</td></tr> +<tr><td> <a href="#key_to_adults">Key to Adults</a></td><td class="text_rt">315</td></tr> +<tr><td> <a href="#key_to_tadpoles">Key to Tadpoles</a></td><td class="text_rt">315</td></tr> +<tr><td> <a href="#ptych_group"><i>Ptychohyla euthysanota</i> Group</a></td><td class="text_rt">315</td></tr> +<tr><td> <a href="#ptych_euth1"><i>Ptychohyla euthysanota</i></a></td><td class="text_rt">315</td></tr> +<tr><td> <a href="#ptych_euth2"><i>Ptychohyla euthysanota euthysanota</i> (Kellogg)</a></td><td class="text_rt">315</td></tr> +<tr><td> <a href="#ptych_euth3"><i>Ptychohyla euthysanota macrotympanum</i> (Tanner)</a></td><td class="text_rt">320</td></tr> +<tr><td> <a href="#ptych_leonh"><i>Ptychohyla leonhardschultzei</i> (Ahl)</a></td><td class="text_rt">323</td></tr> +<tr><td> <a href="#ptych_spin"><i>Ptychohyla spinipollex</i> (Schmidt)</a></td><td class="text_rt">327</td></tr> +<tr><td> <a href="#ptych_schm1"><i>Ptychohyla schmidtorum</i> Group</a></td><td class="text_rt">331</td></tr> +<tr><td> <a href="#ptych_schm2"><i>Ptychohyla schmidtorum</i></a></td><td class="text_rt">331</td></tr> +<tr><td> <a href="#ptych_schm3"><i>Ptychohyla schmidtorum schmidtorum</i> Stuart</a></td><td class="text_rt">331</td></tr> +<tr><td> <a href="#ptych_schm4"><i>Ptychohyla schmidtorum chamulae</i> Duellman</a></td><td class="text_rt">334</td></tr> +<tr><td> <a href="#ptych_schm5"><i>Ptychohyla ignicolor</i> Duellman</a></td><td class="text_rt">337</td></tr> +<tr><td><span class="smcap"><a href="#dist_and_ecology">Distribution and Ecology</a> </span></td><td class="text_rt">340</td></tr> +<tr><td> <a href="#geogr_dist">Geographic Distribution of the Species</a></td><td class="text_rt">340</td></tr> +<tr><td> <a href="#habitat_pref">Habitat Preference</a></td><td class="text_rt">342</td></tr> +<tr><td> <a href="#intersp_comp">Interspecific Competition</a></td><td class="text_rt">343</td></tr> +<tr><td> <a href="#repro_dev">Reproduction and Development</a></td><td class="text_rt">344</td></tr> +<tr><td><span class='pagenum'><a name="Page_300" id="Page_300">[Pg 300]</a></span><a href="#phylogeny"><span class="smcap">Phylogeny of Ptychohyla</span></a></td><td class="text_rt">345</td></tr> +<tr><td> <a href="#nat_assemb"><i>Ptychohyla</i> as a Natural Assemblage</a></td><td class="text_rt">345</td></tr> +<tr><td> <a href="#generic_rel">Generic Relationships</a></td><td class="text_rt">346</td></tr> +<tr><td> <a href="#intersp_rel">Interspecific Relationships</a></td><td class="text_rt">347</td></tr> +<tr><td><span class="smcap"><a href="#literature_cited">Literature Cited</a> </span></td><td class="text_rt">349<br /> +</td></tr></table> +<p> </p> +<p> </p> + +<span class='pagenum'><a name="Page_301" id="Page_301">[Pg 301]</a></span> +<hr style="width: 65%;" /> +<a name="introduction"></a><div class="caption2">INTRODUCTION</div> + +<p>Probably no ecological group of hylid frogs (some <i>Hyla</i> plus +<i>Plectrohyla</i> and <i>Ptychohyla</i>) in Middle America is so poorly known +as those species that live in the cloud forests on steep mountain +slopes and breed in cascading mountain streams. During the last +half of the nineteenth century most of the species of hylids living in +the lowlands of southern México and northern Central America +were named and described. Despite the extensive collecting by +Salvin and Godman, Nelson and Goldman, and the various expeditions +of the <i>Mission Scientifique</i>, no members of the genus <i>Ptychohyla</i> +were obtained until 1927, when in the mountains of El Salvador +Ruben A. Stirton found a small tree frog that subsequently was +described and named <i>Hyla euthysanota</i> by Kellogg (1928). Until +recently frogs of this genus were <a name="known"></a><a href="#typos">known</a> from few specimens and in +the literature by nearly as many names.</p> + +<p>Although I first collected <i>Ptychohyla</i> in 1956, it was not until 1960 +that special efforts were made to obtain specimens of this genus. +The summer of 1960 was spent in southern México and Guatemala, +where every accessible stream in the cloud forests was searched for +tree frogs, especially <i>Ptychohyla</i> and <i>Plectrohyla</i>. Similar, but less +extensive, investigations were carried out in 1961 and 1962. The +result of this field work is a rather large collection of <i>Ptychohyla</i> +representing all of the known species, plus tape recordings of the +breeding calls and tadpoles of all of the species.</p> + +<p>Previously, I have discussed the nomenclature of one of the +species (Duellman, 1960) and have described two new species +(Duellman, 1961). In the latter paper I made reference to a future +account (this one) that would deal with the systematics and biology +of the entire genus. Although I have series of specimens, tadpoles, +osteological preparations, and recordings of breeding calls, thereby +having a wide array of data at my disposal, much still remains to be +learned about these frogs, especially about various aspects of their +life histories. Even the validity of the genus is open to question; +this problem is discussed at length in the section beyond entitled +"<i>Ptychohyla</i> as a Natural Assemblage."</p> +<p> </p> + +<a name="acknwldgmnts"></a> +<div class="caption3">Acknowledgments</div> + +<p>I am indebted to the following persons for permitting me to examine specimens +in their care: Miguel Alvarez del Toro, Museo Zoología de Tuxtla +Gutierrez, México (MZTG); Charles M. Bogert and Richard G. Zweifel, American +Museum of Natural History (AMNH); Doris M. Cochran, United States +<span class='pagenum'><a name="Page_302" id="Page_302">[Pg 302]</a></span> +National Museum (USNM); Norman Hartweg and Charles F. Walker, University +of Michigan Museum of Zoology (UMMZ); Robert F. Inger, Chicago +Natural History Museum (CNHM); Hobart M. Smith, University of Illinois +Museum of Natural History (UIMNH); Heinz Wermuth, Zoologisches Museum +Berlin (ZMB); and Ernest E. Williams, Museum of Comparative <a name="Zoology"></a><a href="#typos">Zoology</a> +(MCZ). The abbreviations following names of institutions will be used +throughout the text; the Museum of Natural History at the University of Kansas +is abbreviated KU.</p> + +<p>Throughout my work on these frogs I have profited from discussions with +L. C. Stuart, who has made many valuable suggestions and with his characteristic +generosity has placed at my disposal his extensive collections of tadpoles +from Guatemala. For his aid I am indeed grateful. I am grateful to +Thomas E. Moore for tapes of breeding calls of two species.</p> + +<p>My own field work was made more enjoyable and profitable through the +assistance of Dale L. Hoyt, Craig E. Nelson, Jerome B. Tulecke, and John +Wellman, all of whom spent many hours in often unsuccessful attempts to +collect specimens and record breeding calls of <i>Ptychohyla</i>. I am indebted to +many residents of México, Guatemala, and El Salvador for permission to work +on their land and for providing shelter, food, and guides. I am especially +grateful to Mr. and Mrs. Horatio Kelly of "Colegio Linda Vista" at Pueblo +Nuevo Solistahuacán, Chiapas, for a pleasant stay at their school; Jordi Juliá +Z. of the Comisión del Papaloapan, Ciudad Alemán, Veracruz, for arranging +for field work in northern Oaxaca in 1959; Walter Hannstein and Lothar +Menzel for the use of facilities at Finca La Paz, Guatemala, in 1960; Alan +Hempstead for the use of facilities at Finca Los Alpes, Guatemala in 1960 +and 1961; and Julio Aguirre C. of the Instituto Tropical de Investigaciones +Científicas, San Salvador, El Salvador, for providing comfortable working quarters +and transportation and guides to the mountains in northern El Salvador. +Without the cheerful efforts of Jorge A. Ibarra, Director of the Museo Nacional +de Historia Natural in Guatemala, my field work would have been greatly restricted +during politically precarious times in that country. Permits to collect +in México were furnished by the late Luis Macías Arellano of the Dirección +General de Caza. Each of these individuals has my profound thanks for his +indispensable aid.</p> + +<p>Field work on hylid frogs in Middle America has been supported by the +National Science Foundation, Grant NSF-G9827, and this is the 9th publication +on the results of study of the material from America.</p> + +<a name="matrls_mthds"></a> +<div class="caption3">Materials and Methods</div> + +<p>During the course of this study I have examined 247 frogs that I assign to +the genus <i>Ptychohyla</i>, plus 40 lots of tadpoles and 12 skeletal preparations. +Furthermore, I have examined all of the type specimens. I have studied each +of the species and subspecies in the field and have examined from seven (<i>P. +euthysanota macrotympanum</i>) to 33 (<i>P. spinipollex</i>) living individuals of each +species.</p> + +<p>Measurements given in the analysis of data and in the descriptions of the +species are those described by Duellman (1956). In the descriptions of living +colors the capitalized names are from Ridgway (1912). All interpretations of +osteological characters are based on specimens cleared in potassium hydroxide +and stained with alizarin red.</p> + +<p><span class='pagenum'><a name="Page_303" id="Page_303">[Pg 303]</a></span> +Recordings of the breeding calls were made with a Magnemite Portable +Tape-recorder; audiospectrographs were made on a vibralyzer (Kay Electric +Company) using normal pattern and wide bandwidth.</p> +<p> </p> + +<a name="analysis_of_data"></a> +<div class="caption2">ANALYSIS OF DATA</div> + +<p>Data that are used to arrive at a systematic arrangement of the +species of <i>Ptychohyla</i> are analyzed and discussed below for the +values inherent in the analysis. These data are of some value also +in the recognition of species and subspecies but if employed for that +purpose the data must be used in combination with the keys and the +diagnoses of the individual species and subspecies.</p> + +<a name="extrnl_morph"></a> +<div class="caption3">External Morphology</div> + +<p>Each of the external morphological characters used in the systematic +treatment of <i>Ptychohyla</i>, as well as the nature of the tongue, +is discussed below.</p> + +<p><span class="smcap">Size and Proportions.</span>—Comparisons of size and certain proportions +are given in <a href="#Table_1">Table 1</a>. Frogs of this genus are small; the largest +specimen examined is a female of <i>P. euthysanota euthysanota</i> having +a snout-vent length of 53.3 mm. The species comprising the +<i>Ptychohyla schmidtorum</i> group are smaller; the largest specimen +examined is a female of <i>P. schmidtorum schmidtorum</i> having a +snout-vent length of 38.0 mm. An analysis of the various measurements +and proportions shows few constant differences. <i>Ptychohyla +ignicolor</i> differs from all of the other species in having the head +slightly wider than long and the tympanum noticeably less than half +the size of eye. <i>Ptychohyla spinipollex</i> has a relatively narrow interorbital +distance, approximately equal to the width of the eyelid, +whereas in all of the other species that distance is much more than +the width of the eyelid.</p> + +<p><span class="smcap">Snout.</span>—All species have a blunt snout. In <i>P. leonhardschultzei</i> +and <i>P. ignicolor</i> the snout is nearly square in lateral profile; in <i>P. +schmidtorum</i> the snout is slightly rounded above and below, and in +the other species it is rounded above. <i>Ptychohyla leonhardschultzei</i> +and <i>P. spinipollex</i> have a vertical fleshy rostral keel on the snout; in +these <a name="species"></a><a href="#typos">species</a>, because of this keel, the snout in dorsal profile is +pointed. The nostrils are slightly protuberant in all species, and in +<i>P. schmidtorum</i> the internarial region is slightly depressed.</p> +<p> </p> +<p> </p> + +<a name="Table_1"></a> +<span class='pagenum'><a name="Page_304" id="Page_304">[Pg 304]</a></span> +<div class="center"> +<span class="smcap smaller">Table 1.—Variation in Certain Characters in the Species of Ptychohyla. +(Means Are in Parentheses Below the Ranges.)</span> +<p><br /></p> + +<table summary="frame"><tr><td class="bt"> +<table class="data" summary="Species Variations"> +<tr><td class="bt bb">Species</td><td class="data btlb">Sex</td><td class="btlb">Number of specimens</td><td class="btlb">Maximum snout-vent length</td><td class="btlb">Tibia length<br /><span style="border-top:solid #000 1px;white-space:nowrap;">Snout-vent length</span></td><td class="btlb"><span style="border-bottom:solid #000 1px;">Tympanum</span><br />Eye</td><td class="btlb">Vomerine teeth</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td></tr> +<tr><td class="text_lf"><i>P. euthysanota euthysanota</i></td><td class="bl">♂</td><td class="bl">17</td><td class="bl">38.1</td><td class="bl">44.4-55.0<br />(48.7)</td><td class="bl">48.6-63.8<br />(56.3)</td><td class="bl">4-6<br />(5.1)</td></tr> +<tr><td> </td><td class="bl">♀</td><td class="bl">15</td><td class="bl">53.3</td><td class="bl">46.5-56.6<br />(51.4)</td><td class="bl">42.9-56.4<br />(51.4)</td><td class="bl">6-18<br />(9.6)</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td></tr> +<tr><td class="text_lf"><i>P. euthysanota macrotympanum</i></td><td class="bl">♂</td><td class="bl">5</td><td class="bl">38.0</td><td class="bl">48.8-52.0<br />(50.2)</td><td class="bl">50.0-57.1<br />(54.1)</td><td class="bl">0-4<br />(2.6)</td></tr> +<tr><td> </td><td class="bl">♀</td><td class="bl">5</td><td class="bl">44.8</td><td class="bl">46.4-54.1<br />(50.2)</td><td class="bl">48.7-58.9<br />(53.7)</td><td class="bl">8-10<br />(9.2)</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td></tr> +<tr><td class="text_lf"><i>P. leonhardschultzei</i></td><td class="bl">♂</td><td class="bl">16</td><td class="bl">35.6</td><td class="bl">48.8-56.1<br />(51.2)</td><td class="bl">48.7-61.9<br />(52.1)</td><td class="bl">6-9<br />(6.5)</td></tr> +<tr><td> </td><td class="bl">♀</td><td class="bl">3</td><td class="bl">41.6</td><td class="bl">52.3-59.5<br />(54.7)</td><td class="bl">47.5-48.6<br />(48.1)</td><td class="bl">7-12<br />(9.5)</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td></tr> +<tr><td class="text_lf"><i>P. spinipollex</i></td><td class="bl">♂</td><td class="bl">32</td><td class="bl">41.2</td><td class="bl">46.9-53.1<br />(49.0)</td><td class="bl">45.0-55.2<br />(49.5)</td><td class="bl">3-7<br />(4.9)</td></tr> +<tr><td> </td><td class="bl">♀</td><td class="bl">6</td><td class="bl">44.6</td><td class="bl">46.1-50.2<br />(48.8)</td><td class="bl">47.7-53.8<br />(50.4)</td><td class="bl">6-10<br />(7.6)</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td></tr> +<tr><td class="text_lf"><i>P. schmidtorum schmidtorum</i></td><td class="bl">♂</td><td class="bl">25</td><td class="bl">32.8</td><td class="bl">45.3-52.4<br />(48.1)</td><td class="bl">51.5-59.3<br />(54.7)</td><td class="bl">5-11<br />(6.2)</td></tr> +<tr><td> </td><td class="bl">♀</td><td class="bl">9</td><td class="bl">38.0</td><td class="bl">46.5-49.1<br />(47.7)</td><td class="bl">51.3-58.3<br />(54.9)</td><td class="bl">7-11<br />(8.7)</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td></tr> +<tr><td class="text_lf"><i>P. schmidtorum chamulae</i></td><td class="bl">♂</td><td class="bl">40</td><td class="bl">30.5</td><td class="bl">46.0-51.9<br />(48.2)</td><td class="bl">48.2-65.6<br />(54.9)</td><td class="bl">4-6<br />(4.7)</td></tr> +<tr><td> </td><td class="bl">♀</td><td class="bl">4</td><td class="bl">31.8</td><td class="bl">48.1-52.4<br />(50.5)</td><td class="bl">51.4-61.7<br />(55.7)</td><td class="bl">4-9<br />(6.2)</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td></tr> +<tr><td class="text_lf"><i>P. ignicolor</i></td><td class="bl">♂</td><td class="bl">13</td><td class="bl">30.5</td><td class="bl">45.9-52.2<br />(49.6)</td><td class="bl">37.1-47.1<br />(43.2)</td><td class="bl">3-9<br />(6.1)</td></tr> +<tr><td class="text_lf bb bb"> </td><td class="bl bb"> </td><td class="bl bb"> </td><td class="bl bb"> </td><td class="bl bb"> </td><td class="bl bb"> </td><td class="bl bb"> </td></tr> +</table> +</td></tr></table> +</div> +<p> </p> + +<p><span class="smcap">Hand.</span>—The species in the <i>Ptychohyla euthysanota</i> group have a +vestige of web between the first and second fingers; the other fingers +are about one-third webbed. Breeding males have a cluster of +horny nuptial spines on the thumb. The spines are largest in <i>P.</i> +<i>spinipollex</i> (<a href="#Fig_1">Fig. 1</a>) and vary in number from 35 to 66 (average +47.4) on each thumb. In the other species of the <i>Ptychohyla +euthysanota</i> group the spines are smaller and usually more numerous; +the numbers of spines on each thumb (means in parentheses) +in members of this group are: <i>P. euthysanota euthysanota</i>, 44-143 +(83.8); <i>P. euthysanota macrotympanum</i>, 40-110 (63.0); <i>P. leonhardschultzei</i>, +<span class='pagenum'><a name="Page_305" id="Page_305">[Pg 305]</a></span> +24-80 (54.7). The species in the <i>Ptychohyla schmidtorum</i> +group have no web between the first and second fingers and only a +vestige of web between the other fingers. Furthermore, these +species lack nuptial spines in breeding males. Like the usual horny +<a name="excrescences"></a><a href="#typos">excrescences</a> on the thumbs of many species of frogs, the horny +spines on the thumbs of members of the <i>Ptychohyla euthysanota</i> +group are seasonal in development.</p> +<p> </p> + +<a name="Fig_1"></a> +<div class="center"> + <a href="images/fig_1_lg.png"><img src="images/fig_1_sm.png" width="400" height="249" border="0" alt="Palmar Views" title="Palmar views of right hands of (A) Ptychohyla spinipollex and (B) Ptychohyla schmidtorum schmidtorum" /></a> +<p><span class="smcap">Fig. 1.</span> Palmar views of right hands of (A) <i>Ptychohyla spinipollex</i> +(KU 58054) and (B) <i>Ptychohyla schmidtorum schmidtorum</i> (KU 58043). × 4.</p> +</div> +<p> </p> + +<p>Many workers have used the presence of a bifid subarticular +tubercle beneath the fourth finger as a diagnostic character of certain +species of hylids. Examination of the subarticular tubercles +in <i>Ptychohyla</i> reveals considerable intraspecific variation. Bifid +tubercles beneath the fourth finger are found in all species except +<i>P. ignicolor</i>, which is known from only two specimens. In <i>P. +euthysanota euthysanota</i> nearly 60 per cent and in <i>P. schmidtorum +schmidtorum</i> about 90 per cent of the specimens have a bifid +tubercle beneath the fourth finger on one or both hands. All +specimens of <i>P. leonhardschultzei</i> have either a bifid or double +tubercle beneath the fourth finger, and some have a bifid distal +tubercle beneath the third finger.</p> + +<p><span class="smcap">Feet.</span>—Members of the <i>Ptychohyla euthysanota</i> group have a +weak tarsal fold, whereas in the species comprising the <i>Ptychohyla +schmidtorum</i> group the tarsal fold is absent. Webbing on the +foot extends to the discs of the third and fifth toes and to the base +of the penultimate phalanx of the fourth toe, except in <i>P. ignicolor</i>, +which has less webbing.</p> + +<p><span class='pagenum'><a name="Page_306" id="Page_306">[Pg 306]</a></span> +<span class="smcap">Ventrolateral Glands.</span>—Breeding males develop thickened, +pigmented glandular areas on the sides of the body. In living +specimens of <i>P. schmidtorum</i> and <i>P. ignicolor</i> the glands are not +readily visible, but in preservative they show as distinctive orange-colored +areas. These glands are most distinct in <i>P. euthysanota</i>; +in many specimens of this species the glands are elevated above +the surrounding skin. The extent of the glands is variable (<a href="#Fig_2">Fig. +2</a>); probably this variability is due to different degrees of development +in individual frogs rather than to interspecific differences. All +<i>Ptychohyla ignicolor</i> and some <i>P. schmidtorum chamulae</i> have a +small, round glandular area on the chin; to my knowledge this does +not occur in the other species. Superficial examination of microscopic +preparations of the glands reveals no histological differences +between species. The glands occupy most of the thickened area +and have narrow ducts leading to the exterior. Detailed studies of +the histology will be reported elsewhere. Since the glands are +developed only in breeding males, it is surmised that the glands +are associated with some phase of the breeding activity.</p> +<p> </p> + +<a name="Fig_2"></a> +<div class="center"> +<img src="images/fig_2.png" width="600" height="471" border="0" alt="Normal extent of ventrolateral glands" title="Normal extent of ventrolateral glands in (A) Ptychohyla euthysanota euthysanota, (B) Ptychohyla schmidtorum schmidtorum" /> +<p><span class="smcap">Fig. 2.</span> Normal extent of ventrolateral glands in (A) <i>Ptychohyla euthysanota +euthysanota</i> (KU 58008), (B) <i>Ptychohyla schmidtorum schmidtorum</i> (KU +58037), and (C) <i>Ptychohyla ignicolor</i> (UMMZ 119603). × 2<span class="dividend">1</span>⁄<span class="divisor">4</span>.</p> +</div> +<p> </p> +<p> </p> + +<p><span class='pagenum'><a name="Page_307" id="Page_307">[Pg 307]</a></span> +<span class="smcap">Tongue.</span>—The shape of the tongue varies intraspecifically. +Usually the tongue is ovoid; in some specimens it is barely notched +posteriorly, whereas in others it is deeply notched, making the +tongue cordiform. Deeply notched cordiform tongues are found +in <i>P. leonhardschultzei</i> and <i>P. schmidtorum</i>; with the exception of +these two species, some individuals of all species have emarginate +tongues. Some individuals of all species have tongues that are +shallowly notched posteriorly.</p> +<p> </p> + +<a name="color_pattern"></a> +<div class="caption3">Color and Pattern</div> + +<p>The dorsum in living frogs of the genus <i>Ptychohyla</i> is primarily +yellowish or reddish brown, except in <i>P. schmidtorum chamulae</i> +and <i>P. ignicolor</i> in which it is green. Usually there are some darker +blotches or reticulations on the dorsum. The venter usually is +white; in <i>P. ignicolor</i> it is yellow. The venter is spotted in all members +of the <i>Ptychohyla euthysanota</i> group; the species, arranged +from least to most spotting ventrally, are: <i>P. euthysanota euthysanota</i>, +<i>P. euthysanota macrotympanum</i>, <i>P. leonhardschultzei</i>, and +<i>P. spinipollex</i>. The last two species also have bold dark spots on +the flanks. <i>Ptychohyla schmidtorum</i> lacks spots on the venter, +whereas <i>P. ignicolor</i> has small dark flecks ventrally.</p> + +<p><i>Ptychohyla euthysanota</i> and <i>P. schmidtorum</i> have white stripes +on the upper lips and on the flanks. All species have some form +of a pale stripe above the anus and usually rather distinct white or +pale stripes along the ventrolateral edges of the tarsi and forearms. +There are no bright or boldly marked flash-colors (colors +that are revealed only when the hind limbs are extended), except +in <i>P. ignicolor</i>, which has bright red flash-colors in the groin and +on the thighs. In life the iris varies from several different shades +of bronze color to deep red in <i>P. schmidtorum schmidtorum</i>.</p> + +<p>The degree of metachrosis is moderate. Usually any change of +color in life consists only of change in the intensity of color. At +times when the over-all coloration is darkened some markings are +obscured.</p> +<p> </p> + +<a name="osteology"></a> +<div class="caption3">Osteology</div> + +<p>The following description of the skull, hyoid, sternum, and +prepollex is based on a male specimen of <i>P. spinipollex</i> (KU +68632) that has been cleared and stained. The broad, flat skull +(<a href="#Fig_3">Fig. 3</a>) has a large frontoparietal fontanelle. The ethmoid is large +and has a flange laterally. The nasals are of moderate size and in +broad contact with the ethmoid, but are separated from one another +<span class='pagenum'><a name="Page_308" id="Page_308">[Pg 308]</a></span> +medially. The anterior half of the maxillary bears a thin, high +flange. The anterior process of the squamosal is short and widely +separated from the maxillary. The quadratojugal is a small spine-shaped +element projecting anteriorly from the ventral base of the +quadrate; the quadratojugal does not articulate with the maxillary.</p> + +<a name="Fig_3"></a> +<div class="center"> +<img src="images/fig_3_lg.png" width="420" height="311" border="0" alt="Dorsal aspect of skull of Ptychohyla spinipollex" title="Dorsal aspect of skull of Ptychohyla spinipollex" /> +<p><span class="smcap">Fig. 3.</span> Dorsal aspect of skull of <i>Ptychohyla spinipollex</i> (KU +68632). Arrow indicates reduced quadratojugal. × 6.</p> +</div> + +<p>The posteromedian part of the hyoid plate is calcified; from this +plate the long bony, posterior cornua (thyrohyales) extend posterolaterally.</p> + +<p>The omosternum is calcified, widest anteriorly, and has a convex +anterior edge. The calcified xiphisternum is roughly bell-shaped +having short lateral processes anteriorly and a deep notch posteriorly.</p> + +<p>The swollen thumb is supported by a dorsoventrally flattened +spine that does not extrude through the skin.</p> + +<p><span class="smcap">Variation.</span>—In general, the skull varies little. Usually the quadratojugal +is present only as a short element attached to the quadrate, +but in one specimen of <i>P. spinipollex</i> the quadratojugal articulates +with the maxillary and forms a complete quadratojugal-maxillary +arch on each side of the skull. One specimen of <i>P. leonhardschultzei</i> +has a complete arch on one side and an incomplete arch +on the other.</p> + +<p>Only <i>P. spinipollex</i> has lateral processes anteriorly on the xiphisternum; +in the other species the <a name="xiphisternum"></a><a href="#typos">xiphisternum</a> is deeply bell-shaped.</p> + +<p><i>Ptychohyla schmidtorum</i> and <i>P. ignicolor</i> have slightly longer +<span class='pagenum'><a name="Page_309" id="Page_309">[Pg 309]</a></span> +premaxillaries than the other species. The longer premaxillary is +reflected in the larger number of teeth on the bone—9 to 11 (average +10) in four specimens of <i>P. schmidtorum</i> and 10 teeth in one +<i>P. ignicolor</i>, as compared with 6 to 10 (average 7.9) in seven specimens +of the other species. The number of maxillary teeth in the +various species are: <i>P. euthysanota euthysanota</i>, 43; <i>P. euthysanota +macrotympanum</i>, 38; <i>P. leonhardschultzei</i>, 38 and 40; <i>P. spinipollex</i>, +34 and 40; <i>P. schmidtorum schmidtorum</i>, 37 and 43; <i>P. schmidtorum +chamulae</i>, 40 and 41; <i>P. ignicolor</i>, 43. The teeth on the premaxillary +and anterior part of the maxillary are long, pointed, and terminally +curved backwards. Posteriorly on the maxillary the teeth become +progressively shorter and blunter.</p> + +<p>Variation in number of vomerine teeth is shown in <a href="#Table_1">Table 1</a>.</p> +<p> </p> +<p> </p> +<p> </p> + +<span class='pagenum'><a name="Page_310" id="Page_310">[Pg 310]</a></span> +<a name="tadpoles"></a> +<div class="caption3">Tadpoles</div> + +<p>Tadpoles of the genus <i>Ptychohyla</i> are adapted to live in mountain +streams. The bodies are streamlined, and the tails are long +and have low fins (Figs. <a href="#Fig_4">4</a> and <a href="#Fig_5">5</a>). The mouths are large and +directed ventrally. Tadpoles of the two groups of species have +strikingly different mouthparts.</p> +<p> </p> + +<a name="Fig_4"></a> +<div class="center"> +<a href="images/fig_4_lg.png"><img src="images/fig_4_sm.png" width="458" height="500" border="0" alt="Tadpoles" title="Tadpoles of the Ptychohyla euthysanota group" /></a> +<p><span class="smcap">Fig. 4.</span> Tadpoles of the <i>Ptychohyla euthysanota</i> group: (A) <i>P. euthysanota +euthysanota</i> (KU 60042), (B) <i>P. euthysanota macrotympanum</i> +(KU 60049), (C) <i>P. leonhardschultzei</i> (KU 68556), and (D) <i>P. spinipollex</i> +(KU 60053). ×2½.</p> +</div> +<p> </p> + +<a name="Fig_5"></a> +<div class="center"> +<a href="images/fig_5_lg.png"><img src="images/fig_5_sm.png" width="550" height="392" border="0" alt="Tadpoles" title="Tadpoles of (A) Ptychohyla schmidtorum schmidtorum" /></a> +<p><span class="smcap">Fig. 5.</span> Tadpoles of (A) <i>Ptychohyla schmidtorum schmidtorum</i> (KU 60051), +(B) <i>P. schmidtorum chamulae</i> (KU 58199), and (C) <i>P. ignicolor</i> (KU 71716). +× 2½.</p> +</div> +<p> </p> + +<p>Lips of tadpoles of the <i>Ptychohyla euthysanota</i> group (<a href="#Fig_6">Fig. 6 +A-D</a>) are folded laterally; there are <span class="dividend">4</span>⁄<span class="divisor">6</span> or sometimes <span class="dividend">4</span>⁄<span class="divisor">7</span> tooth-rows. +A lateral "wing" projects on either side of the upper beak. The +beaks have blunt, peglike serrations. Lips of tadpoles of the +<i>Ptychohyla schmidtorum</i> group (<a href="#Fig_6">Fig. 6 E-G</a>) are greatly expanded +and form a funnel-shaped disc; there are <span class="dividend">3</span>⁄<span class="divisor">3</span> short tooth-rows. There +is no lateral projection or "wing" on either side of the upper beak. +The beaks have long, pointed serrations.</p> +<p> </p> +<p> </p> + +<span class='pagenum'><a name="Page_311" id="Page_311">[Pg 311]</a></span> +<a name="Fig_6"></a> +<div class="center"> +<a href="images/fig_6_lg.png"><img src="images/fig_6_sm.png" width="470" height="600" border="0" alt="Mouthparts of tadpoles" title="Mouthparts of tadpoles of Ptychohyla group" /></a> +<p><span class="smcap">Fig. 6.</span> Mouthparts of tadpoles of <i>Ptychohyla</i>: (A) <i>P. euthysanota euthysanota</i> +(KU 60042), (B) <i>P. euthysanota macrotympanum</i> (KU 60049), (C) +<i>P. leonhardschultzei</i> (KU 68556), (D) <i>P. spinipollex</i> (KU 60053), (E) <i>P. +schmidtorum schmidtorum</i> (KU 60051), (F) <i>P. schmidtorum chamulae</i> (KU +58199), and (G) <i>P. ignicolor</i> (KU 71716). × 10.</p> +</div> + +<p>Variation in certain structural details and in coloration is discussed +for each species and subspecies in the systematic accounts +that follow. Sizes, proportions, and numbers of tooth-rows are +tabulated in <a href="#Table_2">Table 2</a>.</p> +<p> </p> +<p> </p> + +<a name="Table_2"></a> +<span class='pagenum'><a name="Page_312" id="Page_312">[Pg 312]</a></span> +<div class="center"> +<span class="smcap smaller">Table 2.—Comparison of Certain Larval Characters in the Species of +Ptychohyla. (Means Are in Parentheses Below the Ranges.)</span><br /><br /> + +<table summary="frame"><tr><td class="bt"> +<table class="data" summary="Larval Characteristics"> +<tr><td class="bt bb">Species</td><td class="data btlb">Number of specimens</td><td class="data btlb">Maximum length</td><td class="data btlb">Head length<br /><span style="border-top:solid #000 1px;white-space:nowrap;">Total length</span></td><td class="data btlb">Tooth-rows</td></tr> +<tr><td class="text_lf"><i>P. euthysanota<br /> euthysanota</i></td><td class="bl">23</td><td class="bl">40.8</td><td class="bl">30.9-37.3<br />(33.5)</td><td class="bl"><span class="dividend">4</span>⁄<span class="divisor">6</span></td></tr> + +<tr><td class="text_lf"><i>P. euthysanota<br /> macrotympanum</i></td><td class="bl">13</td><td class="bl">43.3</td><td class="bl">30.6-33.4<br />(32.7)</td><td class="bl"><span class="dividend">4</span>⁄<span class="divisor">6</span></td></tr> + +<tr><td class="text_lf"><i>P. leonhardschultzei</i></td><td class="bl">7</td><td class="bl">47.5</td><td class="bl">29.2-32.7<br />(31.1)</td><td class="bl"><span class="dividend">4</span>⁄<span class="divisor">6</span></td></tr> + +<tr><td class="text_lf"><i>P. spinipollex</i></td><td class="bl">32</td><td class="bl">45.0</td><td class="bl">30.2-35.9<br />(33.0)</td><td class="bl"><span class="dividend">4</span>⁄<span class="divisor">7</span></td></tr> + +<tr><td class="text_lf"><i>P. schmidtorum<br /> schmidtorum</i></td><td class="bl">14</td><td class="bl">42.5</td><td class="bl">28.9-31.2<br />(29.9)</td><td class="bl"><span class="dividend">3</span>⁄<span class="divisor">3</span></td></tr> + +<tr><td class="text_lf"><i>P. schmidtorum<br /> chamulae</i></td><td class="bl">4</td><td class="bl">45.0</td><td class="bl">26.9-29.3<br />(27.8)</td><td class="bl"><span class="dividend">3</span>⁄<span class="divisor">3</span></td></tr> + +<tr><td class="text_lf"><i>P. ignicolor</i></td><td class="bl">2</td><td class="bl">39.6</td><td class="bl">29.6-29.8<br />(29.7)</td><td class="bl"><span class="dividend">3</span>⁄<span class="divisor">3</span></td></tr> + +</table> +</td></tr></table> +</div> +<p> </p> + +<p>Evidence on the pattern of development of tooth-rows indicates +that the inner rows develop first. A small tadpole of <i>P. +euthysanota euthysanota</i> has six lower rows and three fully developed +upper rows and only the beginning of the first (outer) +upper row. A small tadpole of <i>P. euthysanota macrotympanum</i> +has four upper rows and five lower rows. In a small tadpole of +<i>P. leonhardschultzei</i> the three upper and four lower tooth-rows are +well developed; the first upper and fifth lower rows are beginning +to develop, and the sixth lower row is absent. In small tadpoles +of <i>P. spinipollex</i>, the sixth lower row is poorly developed, and the +seventh row is absent; large individuals normally have seven lower +rows. A small tadpole of <i>P. schmidtorum chamulae</i> has <span class="dividend">3</span>⁄<span class="divisor">2</span> tooth-rows.</p> + +<a name="breeding_call"></a> +<div class="caption3">Breeding Call</div> + +<p>Breeding calls of all species and subspecies of <i>Ptychohyla</i> were +recorded in the field. Obtaining series of calls of <i>Ptychohyla</i> is +difficult because these frogs call mostly from vegetation along roaring +mountain streams and only by locating a calling frog some +distance from the water or along a quiet stretch of the stream can +good recordings be obtained. For example, four individuals of +<span class='pagenum'><a name="Page_313" id="Page_313">[Pg 313]</a></span> +<i>P. spinipollex</i> were recorded, but only one recording was sufficiently +free of background noise to be analyzed.</p> + +<p>Analysis of breeding calls supports the division of the genus +<i>Ptychohyla</i> into two groups of species. The call of each member +of the <i>Ptychohyla euthysanota</i> group consists of a single long note, +whereas the call of species in the <i>Ptychohyla schmidtorum</i> group +consists of a series of short notes. Since no differences were found +between the calls of subspecies of any given species, the following +discussion of breeding calls pertains to species. These calls are described +briefly below and at greater length in the systematic accounts +farther on. Audiospectrographs of the breeding calls are +shown in <a href="#Plate_11">Plate 11</a>, and comparisons of the characteristics of the calls +are given in <a href="#Table_3">Table 3</a>.</p> +<p> </p> + +<div class="center"> +<a name="Table_3"></a> +<span class="smcap smaller">Table 3.—Comparison of the Breeding Calls of Ptychohyla</span><br /><br /> + +<table summary="frame"><tr><td class="bt"> +<table class="data" summary="Breeding Calls"> + +<tr><td class="bt bb">Species</td><td class="data btlb">Number</td><td class="data btlb">Notes per<br />call group</td><td class="data btlb">Duration of<br />note (seconds)</td><td class="data btlb">Pulses per<br />second</td><td class="data btlb">Frequency<br />range (cps)</td><td class="data btlb">Dominant<br />frequency (cps)</td></tr> + +<tr><td class="text_lf"><i>P. spinipollex</i></td><td class="bl">1</td><td class="bl">1</td><td class="bl">46</td><td class="bl">147</td><td class="bl">3000-5100</td><td class="bl">4300</td></tr> + +<tr><td class="text_lf"><i>P. euthysanota</i></td><td class="bl">7</td><td class="bl">1</td><td class="bl">62<br />(.60-.65)</td><td class="bl">95.3<br />(91-102)</td><td class="bl">1800-4200</td><td class="bl">3070<br />(3000-3200)</td></tr> + +<tr><td class="text_lf"><i>P. leonhardschultzei</i></td><td class="bl">2</td><td class="bl">1</td><td class="bl">79<br />(.62-.95)</td><td class="bl">77<br />(76-78)</td><td class="bl">1500-3500</td><td class="bl">2750<br />(2700-2800)</td></tr> + +<tr><td class="text_lf"><i>P. schmidtorum</i></td><td class="bl">6</td><td class="bl">8.5<a name="FNanchor_A_1" id="FNanchor_A_1"></a><a href="#Footnote_A_1" class="fnanchor">[A]</a><br />(8-9)</td><td class="bl">.064<br />(.054-.070)</td><td class="bl">110<br />(96-121)</td><td class="bl">1400-5800<br />(3350-3450)</td><td class="bl">3400.</td></tr> + +<tr><td class="text_lf"><i>P. ignicolor</i></td><td class="bl">2</td><td class="bl">12<a href="#Footnote_A_1" class="fnanchor">[A]</a><br />(11-13)</td><td class="bl">.079<br />(.078-.080)</td><td class="bl">126<br />(123-129)</td><td class="bl">1000-5000</td><td class="bl">3150<br />(3100-3200)</td></tr> +</table> +</td></tr></table> +<br /> +<div class="footnote"> +<a name="Footnote_A_1" id="Footnote_A_1"></a><a href="#FNanchor_A_1"><span class="label">[A]</span></a> Only an analysis of the long series of calls is given here; see text for explanation. +</div> +</div> +<p> </p> + +<span class='pagenum'><a name="Plate_11" id="Plate_11">[Pl 11]</a></span> +<div class="center"> +<div class="caption3">Plate 11.</div> +<img src="images/plate_11_sm.png" width="418" height="600" border="0" alt="Audiospectrographs" title="Audiospectrographs of the breeding calls of the five species of Ptychohyla" /><br /><br /> +</div> +Audiospectrographs of the breeding calls of the five species of <i>Ptychohyla</i>: +(A) <i>P. spinipollex</i> (KU Tape No. 41), (B) <i>P. euthysanota macrotympanum</i> +(KU Tape No. 48), (C) <i>P. leonhardschultzei</i> (UMMZ Tape No. 525), (D) <i>P. +schmidtorum chamulae</i> (KU Tape No. 52), (E) <i>P. ignicolor</i> (UMMZ Tape +No. 526).<br /> +<p> </p> + +<p><i>P. spinipollex</i> (<a href="#Plate_11">Pl. 11A</a>).—One long note is repeated at intervals of 45 +seconds to four minutes and has an average dominant frequency of 4300 cycles +per second.</p> + +<p><i>P. euthysanota</i> (<a href="#Plate_11">Pl. 11B</a>).—One long note is repeated six to nine times at +intervals of 2.7 to 3.4 seconds and has an average dominant frequency of 3070 +cycles per second.</p> + +<p><i>P. leonhardschultzei</i> (<a href="#Plate_11">Pl. 11C</a>).—One long note is repeated once after 10 +to 13 seconds and has an average dominant frequency of 2750 cycles per +second.</p> + +<p><i>P. schmidtorum</i> (<a href="#Plate_11">Pl. 11D</a>).—The complete call consists of one short series +of notes alternating with two long series. Numbers of notes per series in one +individual having a typical call were 5-8-8-3-9-9. The average dominant +frequency of notes in the short and long series alike is 3400 cycles per second.</p> + +<p><i>P. ignicolor</i> (<a href="#Plate_11">Pl. 11E</a>).—The complete call consists of a short series of notes +alternating with a long series. In one complete recording the numbers of +notes in these series were 4-13-3-11. The notes in the short series have an +average dominant frequency of 2100 cycles per second, whereas the notes in +<span class='pagenum'><a name="Page_314" id="Page_314">[Pg 314]</a></span> +the long series have an average dominant frequency of 3150 cycles per second. +The four series of notes were given in one minute and 15 seconds.</p> +<p> </p> +<p> </p> + +<hr style="width: 65%;" /> + +<a name="systm_accs"></a> +<div class="caption2">SYSTEMATIC ACCOUNTS</div> + +<p>The museum catalogue numbers of the specimens examined, together +with the localities from which they came, are listed at the +end of the account of each subspecies or monotypic species. The +localities that are represented by symbols on the distribution map +(<a href="#Fig_7">Fig. 7</a>) are in roman type; those that are not represented on the +map, because overlapping of symbols would have occurred, are in +italic type.</p> +<p> </p> + +<a name="pytch_taylor"></a> +<div class="caption3"><b>Ptychohyla</b> Taylor, 1944</div> +<br /> +<div class="smaller"> + <div class="species"><i>Ptychohyla</i> Taylor, Univ. Kansas Sci. Bull., 30:41, May 15, 1944. Type, + <i>Ptychohyla adipoventris</i> Taylor, 1944 [= <i>Hyla leonhardschultzei</i> (Ahl), 1934].</div> +</div><br /> +<p><i>Diagnosis.</i>—Small hylids having stream-adapted tadpoles and differing from +other hylid genera in having large ventrolateral glands in breeding males.</p> + +<p><i>Composition.</i>—Five species, two of which are made up of two subspecies, +arranged in two groups of species on the basis of morphological characters of +adults and tadpoles and on the basis of breeding calls.</p> + +<p><i>Distribution.</i>—Moderate elevations from southern Guerrero and northern +Oaxaca, México, to northern El Salvador and central Honduras.</p> +<p> </p> + +<a name="key_to_adults"></a> +<div class="caption3"><span class="smcap">Key to Adults</span></div> + +<table width="100%" summary="Key to Adults"> +<tr><td class="vtop">1.</td><td class="text_lf">A weak tarsal fold; outer fingers one-third webbed; males having spiny +nuptial tuberosities; color in life tan or brown with blotches or reticulations, +never green; iris bronze color<span class="descrp2"><i>P. euthysanota</i> group—2</span></td></tr> +<tr><td class="vtop"> </td><td class="text_lf">No tarsal fold; outer fingers having only vestige of web; males lacking +nuptial tuberosities; color in life green or brown; iris red or golden +color<span class="descrp2"><i>P. schmidtorum</i> group—5</span></td></tr> + +<tr><td class="vtop">2.</td><td class="text_lf">Chest, throat, and flanks usually having brown or black spots; no distinct +white stripe on upper lip or on flanks; a faint white line usually +present above anal opening; a rostral keel<span class="descrp2">3</span></td></tr> +<tr><td class="vtop"> </td><td class="text_lf">Chest, throat, and flanks usually unspotted; distinct white line on upper +lip and on flank present or not; white line above anal opening faint or +well defined; no rostral keel<span class="descrp2">4</span></td></tr> + +<tr><td class="vtop">3.</td><td class="text_lf">Interorbital region much wider than eyelid; spots on throat and +chest black; spots only occasionally present on belly; flanks marbled +with black and white; nuptial spines small, as many as 80 on one +thumb<span class="descrp2"><i>P. leonhardschultzei</i></span></td></tr> +<tr><td class="vtop"> </td><td class="text_lf">Interorbital region about as wide as eyelid; spots on chest and throat +brown or black; spots usually present on belly; flanks having round +brown or black spots; nuptial spines moderate in size, conical, seldom +more than 60 on one thumb<span class="descrp2"><i>P. spinipollex</i></span></td></tr> + +<tr><td class="vtop">4.</td><td class="text_lf">A distinct, broad, white lateral stripe usually present; usually a distinct +white line above anal opening; a distinct white stripe on upper +lip<span class="descrp2"><i>P. euthysanota euthysanota</i></span></td></tr> +<tr><td class="vtop"> </td><td class="text_lf">No white lateral stripe; a faint white stripe above anal opening; no +distinct white stripe on upper lip<span class="descrp2"><i>P. euthysanota macrotympanum</i></span></td></tr> + +<tr><td class="vtop">5.</td><td class="text_lf">A distinct, broad, lateral stripe; a white stripe on upper lip expanded to +form a large spot below eye; hidden surfaces of thighs and webs of +<span class='pagenum'><a name="Page_315" id="Page_315">[Pg 315]</a></span> +feet not red in life; internarial region slightly depressed; diameter of +tympanum greater than one-half diameter of eye<span class="descrp2">6</span></td></tr> +<tr><td class="vtop"> </td><td class="text_lf">No lateral white stripe; no stripe on upper lip; in life dorsum green, +hidden surfaces of thighs and webs of feet orange tan to bright red, +and eye golden color; internarial region flat; diameter of tympanum less +than one-half diameter of eye<span class="descrp2"><i>P. ignicolor</i></span></td></tr> + +<tr><td class="vtop">6.</td><td class="text_lf">Webs of feet and posterior surfaces of thighs pale cream color; dorsum +in life reddish brown; iris bright red<span class="descrp2"><i>P. schmidtorum schmidtorum</i></span></td></tr> +<tr><td class="vtop"> </td><td class="text_lf">Webs of feet and posterior surfaces of thighs pale brown; dorsum in life +green; iris reddish bronze color<span class="descrp2"><i>P. schmidtorum chamulae</i></span></td></tr> + +</table> + +<a name="key_to_tadpoles"></a> +<div class="caption3"><span class="smcap">Key to Tadpoles</span></div> + +<table width="100%" summary="Key to Tadpoles"> +<tr><td class="vtop">1.</td><td class="text_lf">Lips greatly expanded forming a funnel-shaped mouth; tooth-rows <span class="dividend">3</span>⁄<span class="divisor">3</span><span class="descrp2"><i>P. schmidtorum</i> group—2</span></td></tr> +<tr><td class="vtop"> </td><td class="text_lf">Lips folded laterally, not forming a funnel-shaped mouth; tooth-rows +<span class="dividend">4</span>⁄<span class="divisor">6</span> or more<span class="descrp2"><i>P. euthysanota</i> group—4</span></td></tr> + +<tr><td class="vtop">2.</td><td class="text_lf">Belly and mouth mottled; tail cream color heavily blotched with +brown<span class="descrp2">3</span></td></tr> +<tr><td class="vtop"> </td><td class="text_lf">Belly dark gray; tail cream color with dense brown flecking, giving +brown appearance<span class="descrp2"><i>P. schmidtorum chamulae</i></span></td></tr> + +<tr><td class="vtop">3.</td><td class="text_lf">Belly cream color with brown mottling; no large tubercle at each end +of first lower tooth-row<span class="descrp2"><i>P. schmidtorum schmidtorum</i></span></td></tr> +<tr><td class="vtop"> </td><td class="text_lf">Belly grayish green with brown mottling; a large tubercle at each end +of first lower tooth-row<span class="descrp2"><i>P. ignicolor</i></span></td></tr> + +<tr><td class="vtop">4.</td><td class="text_lf">Tooth-rows <span class="dividend">4</span>⁄<span class="divisor">6</span>; cream-colored crescent-shaped mark on posterior part of +body bordered posteriorly by large brown mark<span class="descrp2">5</span></td></tr> +<tr><td class="vtop"> </td><td class="text_lf">Tooth-rows usually <span class="dividend">4</span>⁄<span class="divisor">7</span> (sometimes <span class="dividend">4</span>⁄<span class="divisor">6</span>); cream-colored crescent-shaped +mark on posterior part of body usually indistinct, not bordered posteriorly +by large brown mark<span class="descrp2"><i>P. spinipollex</i></span></td></tr> + +<tr><td class="vtop">5.</td><td class="text_lf">Caudal musculature uniformly flecked with brown; lower tooth-rows +1-4 about equal in length to upper rows<span class="descrp2"><i>P. euthysanota euthysanota</i></span></td></tr> +<tr><td class="vtop"> </td><td class="text_lf">Caudal musculature having brown square blotches dorsally on anterior +one-half of tail; lower tooth-rows 1-4 usually slightly shorter than upper +rows<span class="descrp2">6</span></td></tr> + +<tr><td class="vtop">6.</td><td class="text_lf">Dorsal caudal blotches well defined and extending onto sides of tail; +moderately large brown flecks on caudal fin; eye in life pale reddish +brown<span class="descrp2"><i>P. leonhardschultzei</i></span></td></tr> +<tr><td class="vtop"> </td><td class="text_lf">Dorsal caudal blotches faint, not extending onto sides of tail; small +brown flecks on caudal fin; eye in life silvery bronze<span class="descrp2"><i>P. euthysanota macrotympanum</i></span></td></tr> +</table> +<p> </p> +<p> </p> + +<a name="ptych_group"></a> +<div class="caption3">The <i>Ptychohyla euthysanota</i> Group</div> +<br /> +<p>Three species in group; adults having moderate amount of webbing between +fingers, and tarsal fold; breeding males having spinous, horny, nuptial +tuberosities on pollex; mouths of tadpoles having lateral folds in lips and <span class="dividend">4</span>⁄<span class="divisor">6</span> or +<span class="dividend">4</span>⁄<span class="divisor">7</span> tooth-rows; breeding call consisting of one long note.</p> +<p> </p> + +<a name="ptych_euth1"></a> +<div class="caption3">Ptychohyla euthysanota</div> +<br /> +<p><i>Diagnosis.</i>—Rostral keel absent; nuptial spines in males small; interorbital +region much wider than eyelid.</p> +<p> </p> + +<a name="ptych_euth2"></a> +<table cellpadding="4" summary="mini_plate frame"> +<tr> +<td class="img_left center"> +Plate 12<br /> +<a href="#Plate_12"><img src="images/plate_12_sm.png" width="164" height="300" border="0" alt="Plate 12 small" /></a><br /> +Click to View Larger. +</td><td> +<div class="center"> + <span class="caption3">Ptychohyla euthysanota euthysanota</span> <span class="caption3nb">(Kellogg)</span> +</div> +<p> </p> +<div class="smaller"> + <div class="species"><i>Hyla euthysanota</i> Kellogg, Proc. Biol. Soc. Washington, 41:123-124, June + 29, 1928 [Holotype.—USNM 73296 from Los Esemiles, Depto. Chalatenango, + El Salvador; Ruben A. Stirton collector]. Mertens, Senckenbergiana, + <span class='pagenum'><a name="Page_316" id="Page_316">[Pg 316]</a></span> + 33:169-171, June 15, 1952; Abhand. Senckenbergische Naturf. + Gesell., 487:29, December 1, 1952. Stuart, Proc. Biol. Soc. Washington, + 67:169, August 5, 1954.</div> + + <div class="species"><i>Hyla rozellae</i> Taylor, Univ. Kansas Sci. Bull., + 28:78-80, pl. 9, fig. 1, May + 15, 1942 [Holotype.—USNM 115039 from Salto de Agua, Chiapas, + México; Hobart M. and Rozella Smith collectors]. Taylor and Smith, + Proc. U. S. Natl. Mus., 95:587, June 30, 1945. Smith and Taylor, Bull. + U. S. Natl. Mus., 194:86, June 17, 1948. Stuart, Proc. Biol. Soc. + Washington, 67:169, August 5, 1954.</div> + + <div class="species"><i>Ptychohyla bogerti</i> Taylor, Amer. Mus. Novitates, 1437:13-16, fig. 5, December + 7, 1949 [Holotype.—AMNH 51847 from Río Grande, Oaxaca, + México; Thomas MacDougall collector]. Stuart, Proc. Biol. Soc. Washington, + 67:169, August 5, 1954.</div> + + <div class="species"><i>Ptychohyla euthysanota</i>, Duellman, Univ. Kansas Publ. Mus. Nat. Hist., + 13:351, April 27, 1961.</div> +</div> +</td></tr> +</table> +<p> </p> + +<div class="smaller"> +<p><i>Diagnosis.</i>—Dorsum tan to reddish brown; venter white; rarely flecked with +brown or black; a white stripe on upper lip, on flank, and usually above anus.</p> + +<p><i>Description.</i>—The following description is based on KU 58008 from Finca +La Paz, Depto. San Marcos, Guatemala (<a href="#Plate_12">Pl. 12</a>). Adult male having a snout-vent +length of 35.0 mm.; tibia length, 16.5 mm.; tibia length/snout-vent length, +47.1 per cent; foot length, 14.2 mm.; head length, 11.0 mm.; head length/snout-vent +length, 31.4 per cent; head width, 10.7 mm.; head <a name="width"></a><a href="#typos">width</a>/snout-vent length, +30.6 per cent; diameter of eye, 3.3 mm.; diameter of tympanum, 1.8 mm.; +tympanum/eye, 54.5 per cent. Snout in lateral profile nearly square, slightly +rounded above, and in dorsal profile bluntly rounded; canthus pronounced; +loreal region moderately concave; lips thick, rounded, and slightly flaring; +nostrils protuberant; internarial distance, 3.0 mm.; top of head flat; interorbital +distance, 4.1 mm., and approximately a third broader than width of eyelid, +2.9 mm. Moderately heavy dermal fold from posterior corner of eye above +tympanum to point above insertion of forelimb, covering upper edge of tympanum; +tympanum round, its diameter slightly more than its distance from eye. +Forearm moderately robust, having distinct dermal fold on wrist; dermal fold, +but no row of tubercles along ventrolateral surface of forearm; pollex only +slightly enlarged, bearing triangular shaped patch of small horn-covered spines +(128 on right, 134 on left); second and fourth fingers equal in length; subarticular +tubercles round, distal one on fourth finger bifid; discs moderate in +size, that of third finger equal to diameter of tympanum; no web between first +and second fingers; other fingers one-third webbed. Heels broadly overlap +when hind limbs adpressed; tibiotarsal articulation reaches to middle of eye; +low rounded tarsal fold; inner metatarsal tubercle large, elliptical, and flat; +outer metatarsal tubercle small and round; low dermal fold from heel to disc +of fifth toe; subarticular tubercles round; length of digits from shortest to longest +1-2-5-3-4; third and fifth toes webbed to base of disc; fourth toe webbed to +proximal end of penultimate phalanx; thin dermal fold from inner metatarsal +tubercle to disc of first toe; disc smaller than on fingers. Anal opening at the +level of the upper edge of thighs; anal flap short; anal opening bordered above +by thin transverse dermal fold and laterally by heavy dermal fold. Skin of +dorsum and ventral surfaces of forelimbs and shanks smooth; that of throat, +belly, and ventral surfaces of thighs granular. Ventrolateral glands moderately +developed, not reaching axilla or groin and broadly separated midventrally. +Tongue ovoid, emarginate, and only slightly free posteriorly; vomerine teeth +2-2, situated on small triangular elevations between ovoid inner nares; openings +<span class='pagenum'><a name="Page_317" id="Page_317">[Pg 317]</a></span> +to vocal sac large, one situated along inner posterior edge of each mandibular +ramus.</p> + +<p>Dorsal ground-color of head, body, and limbs dull reddish brown with +irregular dark brown reticulations on head and body and dark brown transverse +bands on limbs; dorsal surfaces of first and second fingers and webbing on hand +cream color; dorsal surfaces of third and fourth fingers dull brown; anterior +surfaces of thighs dull creamy yellow; posterior surfaces of thighs dull brown; +tarsi and toes tan with brown flecks; webbing of feet brown; faint creamy white +stripe along lateral edges of tarsi and forearms; thin white line along edge of +upper lip; distinct white stripe above and beside anal opening; axilla white; +throat, chest, belly, and ventral surfaces of forelimbs creamy white; flanks +white, separated from pale venter by a row of partly connected dark brown +spots; ventral surfaces of thighs dull creamy yellow; feet grayish brown; +ventrolateral glands pale grayish brown; small brown flecks on periphery of +chin.</p> + +<p>In life the dorsal ground-color was pale reddish brown (Orange-Cinnamon); +dorsal reticulations dark brown (Chocolate); dorsal surfaces of first +and second fingers and webbing on hands creamy tan (Light Pinkish Cinnamon); +posterior surfaces of thighs reddish brown (Vinaceous-Tawny); webbing +of feet gray (Deep Mouse Gray); throat and belly grayish white (Pale +Gull Gray); ventral surfaces of hind limbs creamy white (Marguerite Yellow); +spots on flanks dark brown (Warm Sepia); iris reddish bronze (Apricot +Orange).</p> + +<p><i>Variation.</i>—No geographic variation in structural characters is discernible; +variation in size and proportions is given in <a href="#Table_1">Table 1</a>. Of 32 adults examined, +seven have the tongue shallowly notched posteriorly; in the others the tongue +is emarginate. Twenty specimens have a bifid subarticular tubercle beneath +the fourth finger; in the others there are no bifid tubercles.</p> + +<p>The coloration described above is typical of the 16 specimens available from +Finca La Paz. The living coloration at night, when the frogs were collected, +was somewhat darker than the living colors described above, which were recorded +for the frogs the morning after collection, at which time one individual +had a pale reddish brown dorsum (Orange-Cinnamon) with dull olive green +(Deep Grape Green) reticulations on the back and transverse bands on the +limbs; the dorsal surfaces of the first and second fingers and the discs on the +third and fourth fingers were orange (Mikado Orange).</p> + +<p>More than half of the specimens from Finca La Paz agree in all essential +characters with the description given above. The distinctness of the white +stripe on the upper lip is variable; in two individuals the stripe is barely discernible. +Likewise, in some individuals the white stripe on the flanks is not +distinct, either because there are few or no brown spots separating the stripe +from the pale venter, or because the ventrolateral gland has diffused the pale +color on the flanks. There is some noticeable variation in dorsal coloration, +either through a greater or lesser development of dark pigment. One specimen +(KU 58007) is grayish tan above with dark brown markings; the +posterior surfaces of the thighs are dull grayish yellow; the first and second +fingers and the webbing on the hands are pale yellowish gray; the belly and +throat are dusty white; the flecks on the throat are gray; the ventral surfaces +of the feet are grayish brown. Dark individuals, such as KU 58009 have a +uniform dark brownish black dorsum; the belly is cream; the first and second +<span class='pagenum'><a name="Page_318" id="Page_318">[Pg 318]</a></span> +fingers and the webbing on the hands are dull creamy tan; the dorsal and +ventral surfaces of the feet are dark brown. In KU 58013 there is a heavy +suffusion of brown on the throat and flanks. Two specimens have scattered +white flecks on the dorsum.</p> + +<p>The reddish brown dorsal ground-color with dark brown reticulations on +the head and body and dark brown transverse bands on the limbs seems to be +rather constant throughout the range of the subspecies. Likewise, the presence +of the white stripe on the upper lip and the white stripe around the anal opening +are present on most specimens. In breeding males having well-developed +ventrolateral glands the lateral white stripe often is obliterated.</p> + +<p><i>Description of Tadpole.</i>—The following description is based on KU 60042 +from Finca La Paz, Depto. San Marcos, Guatemala (Figs. <a href="#Fig_4">4A</a> and <a href="#Fig_6">6A</a>). No +limb buds; total length, 35.8 mm.; body length, 11.2 mm.; body length/total +length, 31.3 per cent. Body moderately depressed, slightly wider than deep, +ovoid in dorsal profile; mouth directed ventrally; eyes small, directed dorsolaterally; +nostrils slightly protuberant and directed anteriorly, closer to eye than +snout; spiracle sinistral and posteroventrad to eye; anal tube dextral. Caudal +fin low, rounded posteriorly; depth of caudal musculature about one-half +greatest depth of caudal fin; musculature extends nearly to tip of tail.</p> + +<p>Mouth large; lips having deep lateral folds; two complete rows of papillae +on lips; five to six rows of papillae laterally. Beaks moderately developed, +bearing peglike serrations; slender lateral projections on upper beak; tooth-rows +<span class="dividend">4</span>⁄<span class="divisor">6</span>; upper rows subequal in length, second longest; fourth row interrupted +medially; lower rows complete; lower rows 1-4 equal in length to upper rows; +fifth lower row somewhat shorter; sixth lower row short.</p> + +<p>Body brown above; tip of snout cream color; grayish cream color below; +caudal musculature creamy tan; caudal fin transparent; cream-colored crescent-shaped +mark on posterior edge of body and anterior part of caudal musculature, +bordered posteriorly by dark brown blotch; scattered brown flecks on caudal +musculature and posterior part of caudal fin. Eye bronze color in life.</p> + +<p><i>Variation.</i>—The variation in size and proportions is given in <a href="#Table_2">Table 2</a>. In +some specimens the first upper tooth-row is irregular, sometimes broken, and +often shorter than other upper tooth-rows. Usually the fourth upper and first +lower, and sometimes the sixth lower, tooth-rows are interrupted medially. One +specimen has a short, irregular, seventh lower tooth-row; all others have six.</p> + +<p>The cream-colored crescent-shaped mark usually is distinct. The brown +blotch posterior to this mark is variously shaped ranging from a narrow vertical +bar to a triangular blotch. Brown flecks seldom are present on the anterior +part of the ventral caudal fin.</p> + +<p><i>Comparisons.</i>—Aside from the characters given in the diagnosis, <i>P. euthysanota +euthysanota</i> can be distinguished from both <i>P. spinipollex</i> and <i>P. leonhardschultzei</i> +by the absence of bold black and white marbling on the flanks; +furthermore, from the former it can be distinguished by having more and +smaller horny nuptial spines and from the latter by having the snout, in +lateral profile, rounded above and not acutely angulate. <i>Ptychohyla euthysanota +euthysanota</i> differs from <i>P. euthysanota macrotympanum</i> by normally +having a darker dorsal color, broader stripe on upper lip, and a distinct lateral +stripe.</p> + +<p>Occurring sympatrically with <i>Ptychohyla euthysanota euthysanota</i> are several +species of <i>Plectrohyla</i>, all of which differ in having a bony prepollex, rather +<span class='pagenum'><a name="Page_319" id="Page_319">[Pg 319]</a></span> +rugose skin on the dorsum, and more squat bodies. Other sympatric species +are <i>Ptychohyla schmidtorum schmidtorum</i>, which lacks a tarsal fold and nuptial +spines and has a red eye in life, <i>Hyla salvadorensis</i>, which has a green dorsum +and lacks spinous nuptial tuberosities, and <i>Hyla sumichrasti</i>, a small yellow +frog usually lacking vomerine teeth.</p></div> + +<p><i>Life History.</i>—This subspecies breeds in clear, swift mountain +streams. Males call from stems and leaves of plants at the edge of, +or overhanging, the streams. The breeding call consists of a soft +"wraack," repeated at intervals of three to four seconds. Each note +has a duration of 0.60 to 0.65 seconds and has 91 to 102 pulses per +second; the dominant frequency falls between 3000 and 3200 cycles +per second.</p> + +<p>Tadpoles in various stages of development were found at Finca +La Paz, Guatemala, in late July. This indicates that there is either +extreme differential growth, or, more probably, an extended breeding +season. A tadpole having a body length of 6.8 mm. and a total +length of 19.1 mm. has a short median first upper tooth-row; lower +tooth-rows 3-6 are only two-thirds as long as lower rows 1 and 2. +Two recently metamorphosed young have snout-vent lengths of 14.2 +and 14.8 mm.; they are colored like the adults.</p> + +<p><i>Remarks.</i>—The type specimen of <i>Hyla euthysanota</i> Kellogg +(1928:123) is a female; therefore, when Taylor (1944) proposed the +name <i>Ptychohyla</i> for hylids having ventrolateral glands in breeding +males, he was unaware that <i>Hyla euthysanota</i> was a member of this +group. In his description of <i>Hyla rozellae</i>, Taylor (1942) did not +compare his specimens with <i>Hyla euthysanota</i>, but instead placed +<i>H. rozellae</i> with <i>H. loquax</i> and <i>H. rickardsi</i>. The type series of <i>H. +rozellae</i> consists of one large adult female and several metamorphosing +young. Taylor (1949:16) based the description of <i>Ptychohyla +bogerti</i> on two males and compared these specimens with <i>P. +adipoventris</i> Taylor [= <i>P. leonhardschultzei</i> (Ahl)]. Thus, in a +period of 22 years the females of this species were given two names +and the male another. Stuart (1954:169) suggested that <i>Hyla +euthysanota</i> and <i>Hyla rozellae</i> were <i>Ptychohyla</i>. Now that sufficient +specimens are available from throughout the range it is possible to +determine that the various named populations are conspecific.</p> + +<div class="smaller"> +<p><i>Distribution.</i>—This subspecies inhabits cloud forests at elevations of 660 to +2200 meters on the Pacific slopes of the Sierra Madre from extreme eastern +Oaxaca and western Chiapas, México, through Guatemala to northern El +Salvador; probably it occurs also in southwestern Honduras. Aside from the +specimens listed below, three in the Frankfurt Museum from Depto. Santa Ana, +El Salvador (44571, Hacienda San José; 43040, Hacienda Los Planes; 65119, +Miramundo) are listed by Mertens (1952:29).</p> + +<p><span class='pagenum'><a name="Page_320" id="Page_320">[Pg 320]</a></span> +<i>Specimens examined.</i>—<span class="smcap">Mexico</span>: <i>Chiapas: Cascarada, 30 km. W of Cíltapec</i>, +UMMZ 87851-2; Cerro Ovando, UMMZ 87853-4; Chicomuselo, UMMZ 94439-40; +Finca Juárez, 28 km. N of Escuintla, USNM 115052-5; <i>Las Nubes, Cerro +Ovando</i>, USNM 115030-8; Salto de Agua, USNM 115039-51. <i>Oaxaca</i>: Cerro +Pecho Blanco, UIMNH 40963; between La Gloria and Cerro Azul, UIMNH +40976-7; Río Grande, AMNH 51847-8; Santo Tomás Tecpan, UIMNH 41071.</p> + +<p><span class="smcap">Guatemala</span>: <i>San Marcos</i>: Finca La Paz, 2 km. W of La Reforma, KU +58001-14, 59937 (skeleton), 60042-3 (tadpoles), 60044 (4 young), MCZ +34997, UMMZ 107739, 123151-7 (tadpoles); Finca Pirineos, Río Samalá, +CNHM 35066. <i>Santa Rosa</i>: Finca La Gloria, UMMZ 123148 (tadpoles), +123150 (tadpoles). <i>Sololá</i>: Finca Santo Tomás, UMMZ 123149 (tadpoles); +Olas de Mocá, near Mocá, CNHM 20208.</p> + +<p><span class="smcap">El Salvador</span>: <i>Chalatenango</i>: Los Esemiles, USNM 73296. <i>Santa Ana</i>: +Miramundo, CNHM 65120.</p> +</div> +<p> </p> + +<a name="ptych_euth3"></a> +<table cellpadding="4" summary="mini_plate frame"> +<tr> +<td class="img_left center"> +Plate 13<br /> +<a href="#Plate_13"><img src="images/plate_13_sm.png" width="164" height="300" border="0" alt="Plate 13 small" /></a><br /> +Click to View Larger. +</td><td> +<div class="center"> + <span class="caption3">Ptychohyla euthysanota macrotympanum</span> <span class="caption3nb">(Tanner)</span> +</div> +<p> </p> +<div class="smaller"> + <div class="species"><i>Hyla macrotympanum</i> Tanner, Great Basin Nat., 17:52-53, July 31, 1957 + [Holotype.—AMNH 62141 (formerly BYU 13752) from 10 miles east of + Chiapa de Corzo, Chiapas, México; Robert Bohlman collector].</div> + + <div class="species"><i>Ptychohyla macrotympanum</i>, Duellman, Univ. Kansas Publ. Mus. Nat. Hist., + 13:351, April 27, 1961.</div> +</div> +</td></tr> +</table> +<p> </p> + +<div class="smaller"> +<p><i>Diagnosis.</i>—Dorsum usually pale tan; venter white with scattered brown or +black flecks; a thin white stripe on upper lip and another above anal opening; +no distinct white stripe on flanks.</p> + +<p><i>Description.</i>—The following description is based on KU 58049 from Linda +Vista, Chiapas, México (<a href="#Plate_13">Pl. 13</a>). Adult male having a snout-vent length of +38.0 mm.; tibia length, 19.5 mm.; tibia length/snout-vent length, 51.3 per cent; +foot length, 15.7 mm.; head length, 11.8 mm.; head length/snout-vent length, +31.1 per cent; head width, 11.7 mm.; head width/snout-vent length, 30.8 per +cent; diameter of eye, 3.8 mm.; diameter of tympanum, 2.1 mm.; tympanum/eye, +55.2 per cent. Snout in lateral profile nearly square, slightly +rounded above, and in dorsal profile bluntly rounded; canthus pronounced; +loreal region concave; lips thick, rounded, and slightly flaring; nostrils protuberant; +internarial distance, 3.1 mm.; top of head flat; interorbital distance, +3.8 mm., and approximately a fourth broader than width of eyelid, 3.1 mm. +A moderately heavy dermal fold from posterior corner of eye above tympanum +to point above insertion of forelimb, covering upper edge of tympanum; tympanum +round, its diameter equal to its distance from eye. Forearm moderately +robust, having a distinct dermal fold on wrist; dermal fold, but no +row of tubercles along ventrolateral surface of forearm; pollex only slightly +enlarged, bearing triangular patch of small horn covered spines (94 on right, +100 on left); fourth finger slightly longer than second; subarticular tubercles +round, none bifid; discs moderate in size, that of third finger equal to diameter +of tympanum; vestige of web between first and second fingers; other fingers +one-third webbed. Heels broadly overlap when hind limbs adpressed; tibiotarsal +articulation reaches to middle of eye; weak tarsal fold on distal two-thirds +of tarsus; inner metatarsal tubercle large, elliptical, and flat; outer +metatarsal tubercle small and round; no dermal fold from heel to disc of fifth +toe; subarticular tubercles round; length of digits from shortest to longest +1-2-5-3-4; third toe webbed to base of disc; fifth toe webbed to middle of +penultimate phalanx; fourth toe webbed to proximal end of penultimate +phalanx; no fold of skin from inner metatarsal tubercle to base of disc on first +<span class='pagenum'><a name="Page_321" id="Page_321">[Pg 321]</a></span> +toe; discs much smaller than on fingers. Anal opening near upper edge of +thighs; short anal flap bordered above by thin dermal fold; small tubercles +and heavy dermal fold lateral to anal opening. Skin of dorsum and ventral +surfaces of fore limbs and shanks smooth; that of throat, belly, and ventral +surfaces of thighs granular. Ventrolateral glands weakly developed, not +reaching axilla or groin and broadly separated midventrally. Tongue ovoid, +shallowly notched posteriorly, and barely free behind; vomerine teeth 2-2, +situated on small triangular elevations between ovoid inner nares; openings to +vocal sac large, one situated along inner posterior edge of each mandibular +ramus.</p> + +<p>Dorsal ground-color of head, body, and limbs pale pinkish tan with the +greatest part of head and body covered by large gray interconnected blotches; +black flecks over most of dorsum; grayish brown transverse bands on shanks; +dorsal surfaces of first and second fingers pale grayish yellow; dorsal surfaces +of third and fourth fingers and webbing on hand pale grayish tan; anterior +surfaces of thighs pale flesh-color; posterior surfaces of thighs pale grayish +yellow; dorsal surfaces of tarsi and toes pale grayish tan with black flecks; +webbing of feet pale gray; faint creamy white stripes along ventrolateral +edges of tarsi and forearms; a very thin white line along edge of upper lip; +a narrow grayish white stripe above anal opening; axilla gray; throat, chest, +belly, and ventral surfaces of forelimbs pale grayish white; ventral surfaces of +hind limbs cream color; flanks gray flecked with brown; ventral surfaces of +feet grayish tan; ventrolateral glands pinkish tan; anterior one-half of chin +flecked with brown.</p> + +<p>In life the dorsum was pale tan (Pinkish Buff); the dark markings on dorsum +dull brown (Tawny-Olive); tarsi pale tan (Pale Pinkish Buff); flanks +pinkish tan (Pale Cinnamon-Pink); iris coppery bronze (Capucine Orange).</p> + +<p><i>Variation.</i>—The few specimens from a limited geographic region preclude +any analysis of geographic variation. All specimens, except the one described +above, have the fifth toe webbed to the base of the disc. Many individuals +have a bifid subarticular tubercle beneath the fourth finger. The shape of +the posterior edge of the tongue varies from nearly straight and shallowly +notched to bluntly rounded and emarginate. Two females (KU 58050-1) +have more pointed snouts in dorsal profile than do males.</p> + +<p>Some specimens, such as KU 58048, are notably darker than the specimen +described above; in dark specimens the dorsum is brown with dark brown +markings; all fingers and the webbing on the hand are brown. The tarsi and +feet are like those in the specimen described above, but the posterior surfaces +of the thighs are yellowish tan heavily suffused with brown; the venter is +cream color. In life KU 58048 had a brown (Verona Brown) dorsum with +dark brown (Chocolate) markings. KU 58047 is slightly darker than KU +58048 and has scattered small white flecks on the dorsum. All specimens +have the thin white line on the upper lip, but in some individuals it is indistinct. +The grayish white line above the anus is present in all specimens.</p> + +<p><i>Description of Tadpole.</i>—The following description is based on KU 60049 +from Río Hondo, 9.5 kilometers south of Pueblo Nuevo Solistahuacán, Chiapas, +México (Figs. <a href="#Fig_4">4B</a> and <a href="#Fig_6">6B</a>). No limb buds; total length, 36.2 mm.; body +length, 11.1 mm.; body length/total length, 30.6 per cent. Body moderately +depressed, slightly wider than deep, ovoid in dorsal profile; mouth directed +ventrally; eyes small, directed dorsolaterally; nostrils slightly protuberant and +<span class='pagenum'><a name="Page_322" id="Page_322">[Pg 322]</a></span> +directed anteriorly, somewhat closer to eye than snout; spiracle sinistral and +posteroventrad to eye; anal tube dextral. Caudal fin low, acutely rounded +posteriorly; depth of caudal musculature slightly more than one-half greatest +depth of caudal fin; caudal musculature extending nearly to tip of tail.</p> + +<p>Mouth large; lips having deep lateral folds; two complete rows of papillae +on lips; five or six rows of papillae laterally. Beaks moderately developed, +bearing small peglike serrations; moderately wide lateral projections on upper +beak; tooth-rows <span class="dividend">4</span>⁄<span class="divisor">6</span>; upper rows subequal in length; fourth row interrupted +medially; length of lower rows 1-4 subequal to upper rows; fifth and sixth +lower rows decreasing in length; first lower row interrupted medially.</p> + +<p>Body brown above; tip of snout cream color; venter creamy white; caudal +musculature creamy tan; caudal fin transparent; cream-colored crescent-shaped +mark on posterior edge of body and anterior part of caudal musculature, +bordered posterodorsally by dark brown blotch; four dark brown blotches on +dorsum of anterior part of caudal musculature; scattered brown flecks on +caudal musculature and fin; eye silvery bronze in life.</p> + +<p><i>Variation.</i>—The variation in size and proportions is given in <a href="#Table_2">Table 2</a>. All +specimens have <span class="dividend">4</span>⁄<span class="divisor">6</span> tooth-rows; in some the first lower row is interrupted +medially. Specimens from Jacaltenango and two kilometers west of San +Pedro Necta, Depto. Huehuetenango, Guatemala, have weakly developed sixth +lower tooth-rows.</p> + +<p>The cream-colored crescent-shaped mark is distinct in all specimens; the +dorsal blotches on the anterior part of the caudal musculature are narrow and +do not extend far onto the sides of the tail. The blotches are most distinct in +small tadpoles and sometimes indistinct in large ones.</p> + +<p><i>Comparisons.</i>—<i>Ptychohyla euthysanota macrotympanum</i> can be distinguished +from both <i>P. spinipollex</i> and <i>P. leonhardschultzei</i> by the absence of bold black +and white marbling on the flanks, as well as by the characters given in the +diagnosis; furthermore, from the former it differs in having more and smaller +horny nuptial tuberosities and from the latter by having the snout, in lateral +profile, rounded above instead of angulate. <i>Ptychohyla euthysanota macrotympanum</i> +differs from <i>P. e. euthysanota</i> by normally having a paler dorsum, narrower +stripe on upper lip, and no distinct lateral stripe.</p> + +<p><i>Ptychohyla euthysanota macrotympanum</i> occurs sympatrically with <i>Plectrohyla +guatemalensis</i> and <i>P. matudai matudai</i>. Each of the last two has a bony +prepollex, rather rugose skin on the dorsum, and more squat body. Other +sympatric species are <i>Hyla walkeri</i>, which has a green dorsum with brown +markings and a rather pointed snout, and <i>Hyla sumichrasti</i>, a small yellow +frog usually lacking vomerine teeth.</p> +</div> + +<p><i>Life History.</i>—This species breeds in clear mountain streams in +mixed pine and broad-leafed forest. Males call from trees and +bushes along the streams. The breeding call consists of a soft +"wraack," repeated three to nine times with intervals of 2.7 to 3.4 +seconds between notes. Each note has a duration of 0.60 to 0.65 +second, and a rate of 92 to 100 pulses per second; the dominant +frequency falls between 3000 and 3200 cycles per second (<a href="#Plate_11">Pl. 11B</a>). +The call is indistinguishable from that of <i>P. e. euthysanota</i>.</p> + +<p><span class='pagenum'><a name="Page_323" id="Page_323">[Pg 323]</a></span> +Tadpoles in various stages of development were found in the Río +Hondo, Chiapas, on June 16, 1960. The smallest tadpole has a total +length of 24.1 mm.; in this individual the sixth lower tooth row has +barely started to develop. A metamorphosing frog taken at the +same time has a snout-vent length of 19.8 mm., a short remnant of +the tail, and the mouth and tongue developed, whereas another +individual having a snout-vent length of 17.8 mm. and a tail 31.0 mm. +in length still has larval teeth. Three completely metamorphosed +juveniles collected by L. C. Stuart at Jacaltenango, Guatemala, on +June 6 and 7, 1955, have snout-vent lengths of 16.0, 16.0, and +16.1 mm.</p> + +<p><i>Remarks.</i>—Tanner (1957:52) based the description of <i>Hyla +macrotympanum</i> on a single female, which, of course, lacked the +characters diagnostic of <i>Ptychohyla</i>. On the basis of general external +characters Tanner suggested that <i>Hyla macrotympanum</i> was +related to <i>H. miotympanum</i>, from which it differed in having a +larger tympanum and a bifid subarticular tubercle beneath the +fourth finger. The collection of additional females, together with +males of the species, has shown that <i>Hyla macrotympanum</i> is a +<i>Ptychohyla</i>.</p> + +<p>Intergradation between <i>Ptychohyla euthysanota</i> and <i>P. macrotympanum</i> +has not been demonstrated. Currently their ranges are +separated by the dry valleys of the Río Grijalva and Río Cuilco. +The similarity in structure of the adults and tadpoles and the indistinguishable +breeding calls are the basis for considering the two +populations subspecies.</p> + +<div class="smaller"> +<p><i>Distribution.</i>—This species occurs in mixed pine and broad-leafed forests +at elevations of 700 to 1700 meters on the southern slopes of the Chiapan Highlands +and Sierra de Cuchumatanes, in Guatemala. These forests are on the +south facing slopes north of the valleys of the Río Grijalva and Río Cuilco.</p> + +<p><i>Specimens examined.</i>—<span class="smcap">Mexico</span>: <i>Chiapas</i>: 6 km. NE of Chiapa de Corzo, +TCWC 16183; 16 km. E of Chiapa de Corzo, AMNH 62141; Linda Vista, 2 km. +NW of Pueblo Nuevo Solistahuacán, KU 58049-51, 59939 (skeleton); <i>Río +Hondo, 9.5 km. S of Pueblo Nuevo Solistahuacán</i>, KU 58047-8, 60046-7, +60048-9 (tadpoles); San Fernando, MZTG 15, 17; Tonina (ruins), KU 41592.</p> + +<p><span class="smcap">Guatemala</span>: <i>Huehuetenango</i>: Jacaltenango, UMMZ 123139 (tadpoles); +0.5 km. E of Jacaltenango, UMMZ 123142-3; 2 km. S of Jacaltenango, UMMZ +123141; 2 km. W of San Pedro Necta, UMMZ 123140 (tadpoles).</p> +</div> +<p> </p> + +<a name="ptych_leonh"></a> +<table cellpadding="4" summary="mini_plate frame"> +<tr> +<td class="img_left center"> +Plate 14<br /> +<a href="#Plate_14"><img src="images/plate_14_sm.png" width="164" height="300" border="0" alt="Plate 14 small" /></a><br /> +Click to View Larger. +</td><td> +<div class="center"> + <span class="caption3">Ptychohyla leonhardschultzei</span> <span class="caption3nb">(Ahl)</span> +</div> +<p> </p> +<div class="smaller"> + <div class="species"><i>Hyla leonhard-schultzei</i> Ahl, Zool. Anz., 106:185-186, fig. 1, April 15, 1934 + [Holotype.—ZMB 34353 from Malinaltepec, Guerrero, México; Leonhard + Schultze collector]. Smith and Taylor, Bull. U. S. Natl. Mus., 184:87, + June 17, 1948.</div> + + <div class="species"><i>Hyla godmani</i>, Ahl, Zool. Anz., 106:186, April 15, 1934.</div> + + <div class="species"><i>Hyla pinorum</i> Taylor, Proc. Biol. Soc. Washington, 50:46-48, pl. 2, fig. 2, + <span class='pagenum'><a name="Page_324" id="Page_324">[Pg 324]</a></span> + April 21, 1937 [Holotype.—UIMNH 25049 from Agua del Obispo, Guerrero, + México; Edward H. Taylor collector]. Smith and Taylor, Bull. U. S. + Natl. Mus., 194:87, June 17, 1948.</div> + + <div class="species"><i>Ptychohyla adipoventris</i> Taylor, Univ. Kansas Sci. Bull., 30:41-45, May 15, + 1944 [Holotype.—UIMNH 25047 from Agua del Obispo, Guerrero, + México; Edward T. Taylor collector]. Smith and Taylor, Bull. U. S. + Natl. Mus., 194:91, June 17, 1948. Taylor, Amer. Mus. Novitates, + 1437:16, December 7, 1949. Stuart, Proc. Biol. Soc. Washington, 67:169, + August 5, 1954.</div> + + <div class="species"><i>Hyla milleri</i> Shannon, Proc. U. S. Natl. Mus., 101:473-477, figs. 92b, 93a-c, + May 17, 1951 [Holotype.—USNM 123700 from San Lucas Camotlán, + Oaxaca, México; Walter S. Miller collector].</div> + + <div class="species"><i>Ptychohyla leonhard-schultzei</i>, Duellman, Herpetologica, 16:191-197, figs. + 1-3, September 23, 1960; Univ. Kansas Publ. Mus. Nat. Hist., 13:351, + April 27, 1961.</div> +</div> +</td></tr> +</table> +<p> </p> + +<div class="smaller"> +<p><i>Diagnosis.</i>—Rostral keel present; snout in lateral profile not rounded above; +interorbital region much broader than eyelid; distal subarticular tubercle +beneath fourth finger bifid or double; no white stripe on edge of upper lip; +flanks white with black spots.</p> + +<p><i>Description.</i>—The following description is based on KU 64117 from Vista +Hermosa, Oaxaca, México (<a href="#Plate_14">Pl. 14</a>). Adult male having a snout-vent length +of 35.6 mm.; tibia length, 18.0 mm.; tibia length/snout-vent length, 50.5 per +cent; foot length, 14.3 mm.; head length, 10.7 mm.; head length/snout-vent +length, 30.1 per cent; head width, 10.6 mm.; head width/snout-vent length, +29.8 per cent; diameter of eye, 3.6 mm.; diameter of tympanum, 1.8 mm.; +tympanum/eye, 50.0 per cent. Snout in lateral profile square, not rounded +above, and in dorsal profile rounded with pointed tip resulting from vertical +rostral keel; canthus pronounced; loreal region barely concave; lips thick, +rounded, and barely flaring; nostrils protuberant; internarial distance, 3.2 mm.; +top of head flat; interorbital distance, 3.8 mm., and approximately a fifth +broader than width of eyelid, 3.2 mm. A moderately heavy dermal fold from +posterior corner of eye above tympanum and curving ventrad to anterior edge +of insertion of forelimb, covering upper edge of tympanum; tympanum round, +its diameter equal to its distance from eye. Forearm moderately robust, having +distinct dermal fold on wrist; row of small, low, rounded tubercles along +ventrolateral surface of forearm; pollex only slightly enlarged, bearing triangular +patch of small horn-covered spines (56 on right, 62 on left); second finger +noticeably shorter than fourth; subarticular tubercles round, distal ones on +third and fourth toes bifid; discs moderate in size, that of third finger slightly +larger than diameter of tympanum; no web between first and second fingers; +other fingers one-third webbed. Heels broadly overlap when hind limbs adpressed; +tibiotarsal articulation reaches to middle of eye; a low rounded tarsal +fold on distal half of tarsus; inner metatarsal tubercle elevated, flat, and elliptical; +outer metatarsal tubercle at base of fourth toe, round; row of low, sometimes +indistinct, tubercles from heel to base of fifth toe; subarticular tubercles +round; length of digits from shortest to longest 1-2-3-5-4, third and fifth being +about equal in length; third and fifth toes webbed to base of disc; fourth +toe webbed to base of penultimate phalanx; discs of toes much smaller than on +fingers. Anal opening near dorsal surface of thighs; short anal flap; opening +bordered laterally by heavy dermal fold and ventrolaterally by large tubercles. +Skin of dorsum and ventral surfaces of forelimbs and shanks smooth; that of +throat, belly, and ventral surfaces of thighs granular. Ventrolateral glands +<span class='pagenum'><a name="Page_325" id="Page_325">[Pg 325]</a></span> +moderately developed, reaching axilla but not to groin and broadly separated +midventrally. Tongue cordiform, shallowly notched behind and barely free +posteriorly; vomerine teeth 4-3, situated on transverse elevations between ovoid +inner nares; openings to vocal sac large, one situated along inner posterior edge +of each mandibular ramus.</p> + +<p>Dorsal ground-color of head, body, and limbs pale tan with large interconnected +dark brown blotches on head and body and broad dark brown +transverse bands on limbs; dorsal surfaces of first and second fingers and of +webbing of hands pale brown; dorsal surfaces of third and fourth fingers dark +brown; anterior surfaces of thighs flesh-color; posterior surfaces of thighs +brown; dorsal surfaces of tarsi and feet dark brown; narrow white stripe along +ventrolateral edges of forearms and tarsi; a faint creamy white stripe above +anal opening; axilla white; flanks creamy white, boldly spotted with black; +throat and chest white; ventral surfaces of tarsi and feet dark brown; other +ventral surfaces dusty cream color; large brown spots on chin and anterior +part of abdomen.</p> + +<p>In life the dorsum was reddish brown (Orange-Cinnamon) with dark +brown (Chocolate) blotches; first and second fingers and webbing on hand +pale reddish brown (Cinnamon); webbing on feet dark brown (Clove Brown); +flanks pale creamy white (Pale Olive Buff) with dark brown (Bone Brown) +spots; iris reddish bronze (Apricot Orange).</p> + +<p><i>Variation.</i>—No noticeable geographic variation is apparent in the few available +specimens; variations in proportions are given in <a href="#Table_1">Table 1</a>. The distal +subarticular tubercle of the fourth finger is either bifid or double in all specimens; +that on the third finger usually is bifid, sometimes single. The vertical +rostral keel is prominent in all freshly preserved specimens; in some older +specimens it is indistinct. The tongue always is notched posteriorly; in some +individuals the notch is shallow; in others it is deep.</p> + +<p>Some specimens are paler and less boldly marked than the specimen described +above. All specimens from Agua del Obispo and some specimens +from the northern slopes of the Sierra Madre Oriental in Oaxaca have a pale +tan dorsum with brown markings. In most individuals the white color in the +axilla extends onto the posterior edge of the upper arm. The creamy white +color of the flanks is constant and usually extends slightly dorsad in the +inguinal region. The white stripe above, and sometimes continuing down beside, +the anal opening varies from a thin indistinct line or row of flecks to a +distinct continuous stripe. Two specimens have dark brown spots on the +belly; in the others the spots are confined to the flanks and throat.</p> + +<p><i>Description of tadpole.</i>—The following description is based on KU 68556 +from 7.5 kilometers south of Yetla, Oaxaca, México (Figs. <a href="#Fig_4">4C</a> and <a href="#Fig_6">6C</a>). No +limb buds; total length, 37.3 mm.; body length, 12.2 mm.; body length/total +length, 32.7 per cent; body slightly depressed, barely wider than deep, ovoid +in dorsal profile; mouth directed ventrally; eyes small, directed dorsolaterally; +nostrils barely protuberant and directed anterolaterally, about midway between +snout and eye; spiracle sinistral and posteroventrad to eye; anal tube dextral. +Caudal fin low, bluntly rounded posteriorly; greatest depth of caudal musculature +about one-half depth of caudal fin; musculature extends nearly to tip of +tail.</p> + +<p>Mouth large; lips having deep lateral folds; two complete rows of papillae +on lips; five to seven rows of papillae laterally; beaks moderately developed, +<span class='pagenum'><a name="Page_326" id="Page_326">[Pg 326]</a></span> +bearing short peglike serrations; moderately wide lateral projections on upper +beak; tooth-rows <span class="dividend">4</span>⁄<span class="divisor">6</span>; upper rows subequal in length; fourth row interrupted +medially; length of lower rows 1-4 equal to upper rows; fifth and sixth lower +rows shorter; first lower row interrupted medially.</p> + +<p>Body brown above; tip of snout brown; venter creamy gray; caudal musculature +creamy tan; caudal fin transparent; cream-colored crescent-shaped mark on +posterior edge of body; dark brown flecks on caudal musculature and all except +anterior two-thirds of ventral caudal fin; large brown square blotches on dorsum +of caudal fin; eye reddish brown in life.</p> + +<p><i>Variation.</i>—The variation in size and proportions is given in <a href="#Table_2">Table 2</a>. With +the exception of one specimen having a short, broken, seventh tooth-row, all +specimens have <span class="dividend">4</span>⁄<span class="divisor">6</span> tooth-rows that are like those described above. In some +specimens the brown blotches on the dorsum of the caudal musculature are +fused and marked with cream-colored flecks.</p> + +<p><i>Comparisons.</i>—<i>Ptychohyla leonhardschultzei</i> differs from all other members +of the <i>Ptychohyla euthysanota</i> group in having a square snout, and further +differs from <i>P. spinipollex</i> in more numerous and smaller nuptial spines and in +transverse, instead of posteromedially slanting, vomerine processes between +the inner nares. <i>Ptychohyla leonhardschultzei</i> differs from <i>P. euthysanota</i> in +having a rostral keel and in having white flanks boldly spotted with black.</p> + +<p>All small hylids that are sympatric with <i>Ptychohyla leonhardschultzei</i> are +either yellow (<i>Hyla dendroscarta</i> and <i>H. melanomma</i>) or green (<i>Hyla erythromma</i>, +which has a red eye, <i>Hyla hazelae</i>, which has a black line on the +canthus, and <i>Ptychohyla ignicolor</i>, which has red flash colors on the thighs).</p> +</div> + +<p><i>Life History.</i>—This frog has been found along streams in cloud +forests and in pine-oak forest. Males call from vegetation along +the stream or from rocks in and at the edge of the stream. The call +is a single, long, soft "wraack," repeated at intervals of anywhere +from several seconds to three or four minutes. Each note has a +duration of 0.62 to 0.95 of a second and a rate of 76 to 78 pulses +per second; the dominant frequency falls between 2700 and 2800 +cycles per second (<a href="#Plate_11">Pl. 11C</a>).</p> + +<p>Tadpoles were found in several streams in northern Oaxaca. A +tadpole having a total length of 21.1 mm. has three upper and four +lower tooth-rows well developed; the fourth upper and fifth lower +rows are weakly present, and the sixth lower row has not started to +develop. Two metamorphosed young have snout-vent lengths of +15.2 and 15.5 mm.</p> + +<p><i>Remarks.</i>—Four specific names have been applied to this species. +Ahl (1934:185) based his description of <i>Hyla leonhardschultzei</i> on +a small, poorly preserved female. Taylor (1944:41) proposed the +generic name <i>Ptychohyla</i> for a species (named therein as <i>P. adipoventris</i>) +of hylid having ventrolateral glands and horn-covered +nuptial spines. Obviously, Taylor was unaware that <i>Hyla leonhardschultzei</i> +was the same species. Earlier Taylor (1937:46) described +<span class='pagenum'><a name="Page_327" id="Page_327">[Pg 327]</a></span> +<i>Hyla pinorum</i>. The types of all of these species came from the +Pacific slopes of the Sierra del Sur in Guerrero. Examination of +the types and other available specimens shows that they are representatives +of a single species. The type of <i>Hyla pinorum</i> is an immature +male having a snout-vent length of 26.7 mm. All of these +specimens have the square snout and black and white flanks characteristic +of <a name="leonhardschultzei"></a><a href="#typos"><i>Ptychohyla leonhardschultzei</i></a>. Although Shannon +(1951:473) based his description of <i>Hyla milleri</i> on a male having +well-developed ventrolateral glands, he overlooked the presence of +these glands in his description and discussion of relationships. The +acquisition of more specimens from northern Oaxaca has shown +that <i>Hyla milleri</i> is the same as <i>Ptychohyla leonhardschultzei</i>.</p> + +<div class="smaller"> +<p><i>Distribution.</i>—This species is known from pine-oak forest and cloud forest +on the Pacific slopes of the Sierra Madre del Sur in Guerrero and Oaxaca and +from the Atlantic slopes of the Sierra Madre Oriental in northern Oaxaca. +Specimens have been collected at elevations between 700 and 1650 meters. +Probably the species occurs in humid forests at similar elevations around the +eastern end of the Mexican Highlands in Oaxaca.</p> + +<p><i>Specimens examined.</i>—<span class="smcap">Mexico</span>: <i>Guerrero</i>: Agua del Obispo, CNHM +123489-90, 126651, 106300, MCZ 29639, UIMNH 25047, 25049, USNM +114551; Malinaltepec, ZMB 34351, 34353. <i>Oaxaca</i>: 2.5 km. N of La Soledad, +KU 58061; San Lucas Camotlán, UIMNH 3201, USNM 123700-1; Vista Hermosa, +KU 64116-7, 64119, 68560 (tadpoles), 71344, 71717-8 (tadpoles), +UMMZ 119604; 5 km. S of Yetla, KU 60045 (tadpoles); <i>7.5 km. S of Yetla</i>, +KU 64118, 68556-7 (tadpoles), 68559 (tadpoles), 68561 (2 young), 68630 +(skeleton), UMMZ 115514-5, 118863 (tadpoles); 9 km. S of Yetla, KU 68558 +(tadpoles).</p> +</div> +<p> </p> + +<a name="ptych_spin"></a> +<table cellpadding="4" summary="mini_plate frame"> +<tr> +<td class="img_left center"> +Plate 15<br /> +<a href="#Plate_15"><img src="images/plate_15_sm.png" width="164" height="300" border="0" alt="Plate 15 small" /></a><br /> +Click to View Larger. +</td><td> +<div class="center"> + <span class="caption3">Ptychohyla spinipollex</span> <span class="caption3nb">(Schmidt)</span> +</div> +<p> </p> +<div class="smaller"> + <div class="species"><i>Hyla euthysanota</i>, Dunn and Emlen, Proc. Acad. Nat. Sci. Philadelphia, + 84:25, March 22, 1932.</div> + + <div class="species"><i>Hyla spinipollex</i> Schmidt, Proc. Biol. Soc. Washington, 49:45-46, May 1, + 1936 [Holotype.—MCZ 21300 from the mountains behind Ceiba, Depto. + Atlantidad, Honduras; Raymond E. Stadelman collector]. Stuart, Misc. + Publ. Mus. Zool. Univ. Michigan, 69:32-34, figs. 5-6, June 12, 1948; + Contr. Lab. Vert. Biol. Univ. Michigan, 45:22, 52, 54, 57, May, 1950; + Proc. Biol. Soc. Washington, 67:169, August 5, 1954.</div> + + <div class="species"><i>Ptychohyla spinipollex</i>, Stuart, Contr. Lab. Vert. Biol. Univ. Michigan, 68:48, + November, 1954. Duellman, Univ. Kansas Publ. Mus. Nat. Hist., + 13:351, April 27, 1961.</div> +</div> +</td></tr> +</table> +<p> </p> + +<div class="smaller"> +<p><i>Diagnosis.</i>—Rostral keel present; snout in lateral profile rounded above; +eyelid nearly as wide as interorbital region; flanks white with brown spots; +belly spotted; nuptial spines pointed and moderate in size.</p> + +<p><i>Description.</i>—The following description is based on KU 58054 from Finca +Los Alpes, Depto. Alta Verapaz, Guatemala (<a href="#Plate_15">Pl. 15</a>). Adult male having a +snout-vent length of 37.7 mm.; tibia length, 18.2 mm.; tibia length/snout-vent +length, 48.2 per cent; foot length, 15.8 mm.; head length, 11.7 mm.; +head length/snout-vent length, 31.0 per cent; head width, 11.7 mm.; head +width/snout-vent length, 31.0 per cent; diameter of eye, 3.6 mm.; diameter +of tympanum, 1.9 mm.; tympanum/eye, 52.7 per cent. Snout in lateral profile +<span class='pagenum'><a name="Page_328" id="Page_328">[Pg 328]</a></span> +nearly square, slightly rounded above, and in dorsal profile rounded with a +pointed tip resulting from vertical rostral keel; canthus pronounced; loreal +region barely concave; lips thick, rounded, and barely flaring; nostrils protuberant; +internarial distance, 3.0 mm.; top of head flat; interorbital distance, +3.7 mm., about equal to width of eyelid, 3.6 mm. A heavy dermal fold from +posterior corner of eye above tympanum to point above insertion of forearm, +covering upper edge of tympanum; tympanum round, its diameter equal to +its distance from eye. Forearm moderately robust, having faint dermal fold +on wrist; row of low, rounded tubercles along ventrolateral edge of forearm; +pollex only slightly enlarged, bearing triangular patch of moderate-sized, +pointed, horn-covered spines (38 on right, 43 on left); second finger noticeably +shorter than fourth; subarticular tubercles round, distal one on fourth +finger bifid; discs of fingers of moderate size, that of third finger slightly +smaller than diameter of tympanum; vestigial web between first and second +fingers; other fingers one-third webbed. Heels broadly overlapping when +hind limbs adpressed; tibiotarsal articulation reaches to middle of eye; distinct, +but low, tarsal fold extending length of tarsus; inner metatarsal tubercle +elevated, flat, and elliptical; outer metatarsal tubercle small and round, near +base of fourth toe; row of low indistinct tubercles from heel to base of fifth toe; +subarticular tubercles round; length of toes from shortest to longest 1-2-3-5-4, +the third and fifth being about equal in length; third and fifth toes webbed +to base of disc; fourth toe webbed to base of penultimate phalanx; discs of toes +slightly smaller than those on fingers. Anal opening near upper edge of +thighs; opening bordered laterally by moderately heavy dermal folds +and ventrolaterally by tubercles. Skin of dorsum and ventral surfaces of +forelimbs and shanks smooth; that of throat, belly, and ventral surfaces of +thighs granular. Ventrolateral glands barely evident. Tongue ovoid, barely +notched behind and slightly free posteriorly; vomerine teeth 2-3, situated +on <span style="white-space:nowrap;"><img src="images/v_shape.png" width="16" height="14" border="0" alt="V-shape" title="V-shape" />-shaped</span> elevations between round inner nares; openings to vocal sac +large, one situated along inner posterior edge of each mandibular ramus.</p> + +<p>Dorsal ground-color of head, body, and limbs grayish tan with dark brown +reticulations on head and body and dark brown transverse bars or spots on +limbs; first and second fingers cream color; third and fourth fingers and +webbing on hands pale grayish brown; anterior surfaces of thighs reddish tan; +posterior surfaces of thighs yellowish tan; tarsi and toes pale grayish tan with +brown flecks; webbing on foot pale brown; faint white stripe along ventrolateral +edges of tarsi and forearms; narrow white line above and beside anal +opening; no white stripe on edge of upper lip; axilla pale flesh-color; throat, +chest, and ventral surfaces of limbs pale creamy gray; belly white with +scattered brown flecks; flanks grayish white with dark brown flecks; ventral +surfaces of tarsi dark brown; ventrolateral glands grayish tan.</p> + +<p>In life the dorsal ground-color of the head, body, fore limbs, and thighs +was yellowish tan (Pinkish Buff); dorsal surfaces of shanks and tarsi pale +yellowish tan (Pale Pinkish Buff); markings on head and back brown (Snuff +Brown) to dark brown (Chocolate); dark bands on limbs brown (Tawny-Olive); +first and second fingers creamy tan (Light Pinkish Cinnamon); posterior +surfaces of thighs creamy tan (Light Pinkish Cinnamon); third and fourth +fingers and webbing on hand grayish brown (Avellaneous); webbing on feet +dark brown (Olive Brown); axilla pale pink (Hydrangea Pink); flanks buff +<span class='pagenum'><a name="Page_329" id="Page_329">[Pg 329]</a></span> +(Cream-Buff) becoming yellow (Lemon Chrome) in groin; spots on flanks +dark brown (Clove Brown); iris dull grayish bronze (Orange-Citrine).</p> + +<p><i>Variation.</i>—The distal subarticular tubercle beneath the fourth finger is +bifid in about two-thirds of the specimens; in the rest it is round. The posterior +edge of the tongue varies from being emarginate to shallowly notched. In +most specimens the row of tubercles along the outer edge of the tarsus is +made up of discrete tubercles, but in some individuals the tubercles form a +nearly continuous dermal fold. Most specimens have the vomerine teeth +situated on <span style="white-space:nowrap;"><img src="images/v_shape.png" width="16" height="14" border="0" alt="V-shape" title="V-shape" />-shaped</span> elevations, but in some individuals the elevations are +more nearly transversely situated between the inner nares.</p> + +<p>All 42 specimens from Finca Los Alpes, Guatemala, have dark brown +spots and flecks on the venter. Some individuals have only a few flecks on +the throat and a few large spots on the flanks, as does KU 64125. Other +specimens, such as KU 64132, have dense spotting over the entire venter. +The color of the dorsum varies from pale tan to dark brown with darker brown +markings; the white line above the anus is present in all specimens, but in +some it is indistinct. KU 58053 and 64127 have a dark brown dorsum with +large pale tan, square blotches; in life the blotches were pale tan (Pinkish +Buff); the rest of the dorsum was dark brown (Sayal Brown). KU 58052 +is dark brown with many small white flecks on the dorsum; in life the dorsum +was deep olive brown (Dark Olive).</p> + +<p>Aside from the differences mentioned above, all specimens from Guatemala +are similar in coloration. Three specimens from Honduras (MCZ 21300 and +UMMZ 113102-3) have unspotted white venters. MCZ 21300, the holotype +of <i>P. spinipollex</i>, lacks a white stripe above the anal opening, whereas the +stripe is indistinct in UMMZ 113102-3.</p> + +<p><i>Description of tadpole.</i>—The following description is based on KU 60053 +from Fina Los Alpes, Depto. Alta Verapaz, Guatemala (Figs. <a href="#Fig_4">4D</a> and <a href="#Fig_6">6D</a>). +No limb buds; total length, 37.2 mm.; body length, 12.2 mm.; body length/total +length, 32.8 per cent. Body rounded, not depressed, as wide as deep, ovoid +in dorsal profile; mouth directed ventrally; eyes small, directed dorsolaterally; +nostrils barely protuberant and directed anterolaterally, somewhat closer to eye +than snout; spiracle sinistral and posteroventrad to eye; anal tube dextral. +Caudal fin low, bluntly rounded posteriorly; greatest depth of caudal musculature +about one-half depth of caudal fin; musculature extends nearly to tip of +tail.</p> + +<p>Mouth large; lips having deep lateral folds; two complete rows of papillae +on lips; six or seven rows of papillae laterally; beaks moderately developed, +bearing fine blunt serrations; slender lateral projections on upper beak; tooth-rows +<span class="dividend">4</span>⁄<span class="divisor">7</span>; upper rows subequal in length; fourth row interrupted medially; lower +rows 1-4 equal in length to upper rows; lower rows 5-7 decreasing in length; +first lower row interrupted medially.</p> + +<p>Top of head and tip of snout brown; venter creamy gray; caudal musculature +tan; caudal fin transparent; faint cream-colored, narrow, crescent-shaped +mark on posterior edge of body, not bordered posteriorly by dark brown mark; +brown flecks scattered on caudal musculature and caudal fin; only a few flecks +on anterior half of ventral caudal fin; eye bronze-color in life.</p> + +<p><i>Variation.</i>—The variation in size and proportions as compared with tadpoles +of other species is given in <a href="#Table_2">Table 2</a>. Of the 57 tadpoles of this species that I +<span class='pagenum'><a name="Page_330" id="Page_330">[Pg 330]</a></span> +have examined, 21 have only six lower tooth-rows, although in some of these +specimens a faint ridge for a seventh row is present. In those specimens having +seven lower rows, the seventh often is broken.</p> + +<p>There is considerable variation in coloration. None has a distinct cream-colored, +crescent-shaped mark bordered posteriorly by a dark brown bar or +triangle, as in the other species in the <i>Ptychohyla euthysanota</i> group. Most +specimens have a rather indistinct crescent; some have no crescent. In a few +specimens there is a weakly outlined dark mark posterior to the crescent. +Some specimens in a series of tadpoles from 32 kilometers north of Morazán, +Baja Verapaz, Guatemala, have faint dorsal blotches on the dorsal musculature, +much like those in tadpoles of <i>Ptychohyla leonhardschultzei</i>.</p> + +<p><i>Comparisons.</i>—<i>Ptychohyla spinipollex</i> differs from all other species in the +genus by having moderate-sized, instead of small, pointed nuptial spines; also +it has fewer spines than the other species (see discussion of this character in +Analysis of Data). The nearly equal interorbital breadth and width of the +upper eyelid also is diagnostic of this species.</p> + +<p>Other hylids sympatric with <i>Ptychohyla spinipollex</i> include three species of +<i>Plectrohyla</i>, each of which has a bony prepollex, heavy body, and rugose skin +on the dorsum. The only other sympatric hylid that could be confused with +<i>Ptychohyla spinipollex</i> is <i>Hyla bromeliacea</i>, which has a round snout and +yellowish tan dorsum not marked with dark brown.</p> +</div> + +<p><i>Life History.</i>—At Finca Los Alpes, Guatemala, in July, 1960, and +in August, 1961, calling males were found on bushes and trees along +cascading mountain streams. The breeding call consists of a soft +"wraack," repeated at intervals of 45 seconds to four minutes. Each +note has a duration of about .46 of a second and a rate of 147 pulses +per second. The dominant frequency is 4300 cycles per second +(<a href="#Plate_11">Pl. 11A</a>).</p> + +<p>Tadpoles have been found in cascading mountain streams. Two +metamorphosed young have snout-vent lengths of 15.0 and 15.5 mm.</p> + +<p><i>Remarks.</i>—There is little doubt that all of the specimens herein +referred to <i>Ptychohyla spinipollex</i> are conspecific. However, the +three specimens from Honduras, including the type of <i>Ptychohyla +spinipollex</i>, differ from Guatemalan specimens in lacking all dark +spotting on the venter. Additional specimens from Honduras and +eastern Guatemala may show that two subspecies are recognizable, +in which case the nominal subspecies will be the population in +Honduras.</p> + +<div class="smaller"> +<p><i>Distribution.</i>—This species lives in cloud forests at elevations of 800 to +1700 meters on the Atlantic side of the Guatemalan Highlands from the Sierra +de Cuchumatanes in western Guatemala eastward to central Honduras.</p> + +<p><i>Specimens examined.</i>—<span class="smcap">Guatemala</span>: <i>Alta Verapaz</i>: Finca Chichén, UMMZ +90876 (tadpoles); Finca Los Alpes, KU 58052-60, 59939 (skeleton), 60053 +(tadpoles), 64122-41, 68562, 68563 (tadpoles), 68631-2 (skeletons), MCZ +35000-1, UMMZ 90873, 90874 (3), 90875 (tadpoles); La Primavera, UMMZ +90877 (tadpoles); Panzamalá, UMMZ 90878 (tadpoles). <i>Baja Verapaz:</i> 32 +<span class='pagenum'><a name="Page_331" id="Page_331">[Pg 331]</a></span> +km. N of Morazán, KU 68564 (tadpoles); <i>Santa Elena</i>, UMMZ 98119, 98120 +(2). <i>Huehuetenango</i>: 1 km. E of Barillas, UMMZ 123136-7 (tadpoles). +<i>Progreso</i>: Finca Bucaral, UMMZ 106783 (3), 123138 (tadpoles), S-1292 +(<a name="skeleton"></a><a href="#typos">skeleton</a>).</p> + +<p><span class="smcap">Honduras</span>: <i>Atlantidad</i>: Mountains behind Ceiba, MCZ 21300. <i>Morazán</i>: +Cerro Uyuca, UMMZ 123102-3.</p> +</div> +<p> </p> + +<a name="ptych_schm1"></a> +<div class="caption3">The <i>Ptychohyla schmidtorum</i> Group</div> +<br /> +<p>Two species in group; adults having only vestige of web between fingers +and lacking tarsal fold; pollex of breeding males lacking spinous, horny, nuptial +tuberosities; mouth of tadpole greatly expanded, funnel-shaped, lacking lateral +folds, and having <span class="dividend">3</span>⁄<span class="divisor">3</span> tooth-rows; breeding call consisting of series of short notes.</p> +<p> </p> + +<a name="ptych_schm2"></a> +<div class="caption3"><b>Ptychohyla schmidtorum</b></div> +<br /> +<div class="smaller"> +<p><i>Diagnosis.</i>—Diameter of tympanum more than half of diameter of eye; +internarial region depressed; toes three-fourths webbed; no red flash-colors on +thighs.</p> +</div> +<p> </p> + +<a name="ptych_schm3"></a> +<table cellpadding="4" summary="mini_plate frame"> +<tr> +<td class="img_left center"> +Plate 16<br /> +<a href="#Plate_16"><img src="images/plate_16_sm.png" width="164" height="300" border="0" alt="Plate 16 small" /></a><br /> +Click to View Larger. +</td><td> +<div class="caption3"><b>Ptychohyla schmidtorum schmidtorum</b> Stuart</div> +<br /> +<div class="smaller"> + <div class="species"><i>Ptychohyla schmidtorum</i> Stuart, Proc. Biol. Soc. Washington, 67:169-172, + August 5, 1954 [Holotype.—CNHM 27055 from El Porvenir (17 kilometers + air-line west of San Marcos), Depto. San Marcos, Guatemala; Karl + P. Schmidt collector]. Duellman, Univ. Kansas Publ. Mus. Nat. Hist., + 13:351, 355, April 27, 1961.</div> +</div> +</td></tr> +</table> +<p> </p> + +<div class="smaller"> +<p><i>Diagnosis.</i>—Vomerine teeth 5-11; dorsum dark brown; white spot below +eye; eye red in life.</p> + +<p><i>Description.</i>—The following description is based on KU 58043 from Finca +La Paz, Depto. San Marcos, Guatemala (<a href="#Plate_16">Pl. 16</a>). Adult male having snout-vent +length of 31.6 mm.; tibia length, 15.0 mm.; tibia length/snout-vent length, +47.5 per cent; foot length, 12.5 mm.; head length, 10.2 mm.; head length/snout-vent +length, 32.3 per cent; head width, 9.9 mm.; head width/snout-vent length, +31.3 per cent; diameter of eye, 3.4 mm.; diameter of tympanum, 1.8 mm.; +tympanum/eye, 52.9 per cent. Snout in lateral profile nearly square, slightly +rounded above and below, and in dorsal profile bluntly squared; canthus pronounced; +loreal region concave; lips thick, rounded, and flaring; nostrils protuberant; +internarial distance, 2.2 mm.; internarial region depressed; top of +head flat; interorbital distance, 3.4 mm., much greater than width of eyelid, +2.5 mm. Thin dermal fold from posterior corner of eye above tympanum to +insertion of forelimb, covering upper edge of tympanum; tympanum round, +its diameter equal to its distance from eye. Forearm slender, lacking distinct +dermal fold on wrist; row of low rounded tubercles along ventrolateral edge +of forearm; pollex slightly enlarged; no nuptial spines; second and fourth +fingers about equal in length; subarticular tubercles small and round, distal +one beneath fourth finger bifid; discs small, that of third finger noticeably +smaller than tympanum; no web between first and second fingers; vestige of +web between other fingers. Heels overlap when hind limbs adpressed; +tibiotarsal articulation reaches to middle of eye; no tarsal fold; inner metatarsal +tubercle large, flat, and elliptical; outer metatarsal tubercle small, ovoid, +slightly more distal than inner; subarticular tubercles round; length of digits +from shortest to longest 1-2-5-3-4; third and fifth toes webbed to base of +discs; fourth toe webbed to base of penultimate phalanx; discs of toes smaller +<span class='pagenum'><a name="Page_332" id="Page_332">[Pg 332]</a></span> +than on fingers. Anal opening directed posteriorly at upper edge of thighs; +no anal flap; pair of large tubercles below anal opening and smaller tubercles +farther below. Skin of dorsum and ventral surfaces of forelimbs and shanks +smooth; that of belly and ventral surfaces of thighs granular. Ventrolateral +glands well developed, reaching axilla and groin and narrowly separated on +chest. Tongue ovoid, emarginate posteriorly, and only slightly free behind; +vomerine teeth 3-3, situated on small triangular elevations between ovid inner +nares; openings to vocal sac large, one situated along inner posterior edge of +each mandibular ramus.</p> + +<p>Dorsum of head, body, and limbs reddish brown with indistinct, irregular +darker brown markings on body and dark brown transverse bands or spots +on limbs; first and second fingers creamy white; third and fourth fingers brown; +dorsal surfaces of tarsi and third, fourth, and fifth toes tan with brown spots; +first and second toes and webbing on feet creamy tan; enamel-white stripe +along edge of upper lip continuing over, and on posterior edge of, forearm to +groin, expanded to form spot below eye; belly white, unspotted; ventrolateral +glands pale brown; ventral surfaces of hind limbs and anterior and posterior +surfaces of thighs cream color; enamel-white stripe on heel; creamy white stripe +along ventrolateral edges of tarsi and forearms.</p> + +<p>In life dorsum reddish brown (Terra Cotta) with dark brown (Burnt +Umber) markings; first and second fingers and first and second toes orange-yellow +(Light Orange-Yellow); posterior surfaces of thighs pale reddish tan +(Ochraceous-Salmon); webbing on feet yellowish tan (Deep Colonial Buff); +belly white; iris red (Nopal Red).</p> + +<p><i>Variation.</i>—Little variation in structural characters was observed. All but +five specimens have bifid subarticular tubercles beneath the fourth finger. +Three specimens have cordiform tongues, and in four others the tongue is +ovoid and shallowly notched behind; all other specimens have an emarginate +ovoid tongue.</p> + +<p>Some individuals when active at night had a pale brown (Ochraceous-Tawny) +dorsum with dull olive green (Dark Olive Buff) markings. Otherwise +there was no noticeable variation in color.</p> + +<p><i>Description of tadpole.</i>—The following description is based on KU 60051 +from Finca La Paz, Depto. San Marcos, Guatemala (Figs. <a href="#Fig_5">5A</a> and <a href="#Fig_6">6E</a>). Small +hind limbs; total length, 37.9 mm.; body length, 11.6 mm.; body length/total +length, 30.6 per cent. Body only slightly depressed, nearly as deep as wide, +in dorsal profile ovoid, widest just posterior to eyes; in lateral profile snout +rounded; mouth directed ventrally; eyes small, directed dorsolaterally; nostrils +barely protuberant, directed anteriorly, somewhat closer to eye than snout; +spiracle sinistral and posteroventrad to eye; anal tube dextral. Tail long and +slender; caudal fin low and rounded posteriorly; depth of caudal musculature +one-half greatest depth of caudal fin; musculature not extending to tip of tail.</p> + +<p>Mouth large; thin fleshy lips greatly expanded and forming large funnel-shaped +disc; width of mouth two-thirds greatest width of body; outer edge +of lips having one row of small papillae; inner surfaces of mouth smooth; +scattered large papillae forming one nearly complete row around teeth; other +large papillae laterally; beaks moderately developed, bearing long, pointed +denticulations; no lateral projections on upper beak; tooth-rows <span class="dividend">3</span>⁄<span class="divisor">3</span>, all short; +second and third upper rows subequal in length; first upper row shorter; first +<span class='pagenum'><a name="Page_333" id="Page_333">[Pg 333]</a></span> +and third upper rows interrupted medially; first lower row interrupted medially, +equal in length to second and third upper rows; second lower row slightly +shorter; third lower row shortest.</p> + +<p>Body mottled brown and creamy gray above and below; mouth colored +like body; caudal musculature creamy tan; caudal fin transparent; dark brown +streak mid-laterally on anterior third of caudal musculature; rest of tail and +all of caudal fin heavily flecked with brown; eye red in life.</p> + +<p><i>Variation.</i>—The third upper tooth-row is interrupted in all specimens; in +some individuals the first upper and first lower rows are complete. The variation +in size and proportions is given in <a href="#Table_2">Table 2</a>. The dark brown lateral +streak on the anterior part of the caudal musculature is distinct on most specimens; +the only other variation in coloration is in the amount of brown flecking +on the caudal musculature and fin.</p> + +<p><i>Comparisons.</i>—<i>Ptychohyla schmidtorum schmidtorum</i> differs from <i>P. schmidtorum +chamulae</i> as stated in the diagnosis and in having pale creamy tan, as +opposed to dark brown, webbing on the feet; and from <i>P. ignicolor</i> in having +a depressed, as opposed to a flat, internarial region. Tadpoles of <i>P. s. schmidtorum</i> +have a mottled appearance, as opposed to the more uniform brown color +of <i>P. s. chamulae</i>.</p> + +<p><i>Ptychohyla schmidtorum schmidtorum</i> and several species of <i>Plectrohyla</i> +are sympatric. All species of the latter genus have a bony prepollex, rugose +skin on the dorsum, and heavy body; also sympatric is <i>Ptychohyla e. euthysanota</i>, +which has a tarsal fold and in breeding males spinous nuptial tuberosities.</p> +</div> + +<p><i>Life History.</i>—This species breeds in clear mountain streams +where males call from vegetation along the stream. The call consists +of series of short notes, three to nine notes per series, sounding +like "raa-raa-raa." The duration of each note is approximately .065 +of a second, and has a rate of 96 to 119 pulses per second; the +dominant frequency is about 3400 cycles per second. The call is +almost indistinguishable from that of <i>Ptychohyla schmidtorum +chamulae</i>.</p> + +<p>Tadpoles and metamorphosing young were found at Finca La +Paz, Guatemala, in late July, 1960. Two young lacking tails but +not having completely developed mouths have snout-vent lengths +of 14.2 and 14.6 mm. L. C. Stuart collected four metamorphosing +young at Finca La Paz on May 6, 1949. By May 10 the frogs were +completely metamorphosed, at which time they had snout-vent +lengths of 15.5 to 17.0 (average 16.1) mm.</p> + +<p><i>Remarks.</i>—There is no doubt that this frog is most closely related +to <i>Ptychohyla schmidtorum chamulae</i>, even though the ranges +of the two subspecies are separated by the interior depression of +Chiapas. Since at least at Finca La Paz, Guatemala, <i>P. s. schmidtorum</i> +occurs with <i>P. e. euthysanota</i>, it is surprising that the former +species has not been found at more localities along the Pacific slopes +<span class='pagenum'><a name="Page_334" id="Page_334">[Pg 334]</a></span> +on northern Central America. At Finca La Paz in July, 1960, <i>P. s. +schmidtorum</i> was more abundant than <i>P. e. euthysanota</i>.</p> + +<div class="smaller"> +<p><i>Distribution.</i>—This species is known only from a limited area at elevations +between 1300 and 2200 meters on the Pacific slopes of the Sierra Madre +in extreme eastern Chiapas and western Guatemala.</p> + +<p><i>Specimens examined.</i>—<span class="smcap">Mexico</span>: <i>Chiapas</i>: Finca Irlandia, UMMZ 105429-30.</p> + +<p><span class="smcap">Guatemala</span>: <i>San Marcos</i>: El Porvenir, CNHM 20755, 20761, 69904, +UMMZ 80918; Finca La Paz, 2 km. W of La Reforma, KU 58016-44, 59940-2 +(skeletons), 60050 (3 young), 60051 (tadpoles), 60052 (4 young), MCZ +34998-9, UMMZ 123144-7 (tadpoles).</p> +</div> +<p> </p> + +<a name="ptych_schm4"></a> +<table cellpadding="4" summary="mini_plate frame"> +<tr> +<td class="img_left center"> +Plate 17<br /> +<a href="#Plate_17"><img src="images/plate_17_sm.png" width="164" height="300" border="0" alt="Plate 17 small" /></a><br /> +Click to View Larger. +</td><td> +<div class="caption3"><b>Ptychohyla schmidtorum chamulae</b> Duellman</div> +<br /> +<div class="smaller"> + <div class="species"><i>Ptychohyla chamulae</i> Duellman, Univ. Kansas Publ. Mus. Nat. Hist., 13: + 354-357, pl. 25, fig. 2, April 27, 1961 [Holotype.—KU 58063 from 6.2 + kilometers south of Rayón Mescalapa, Chiapas, México; William E. Duellman + collector].</div> +</div> +</td></tr> +</table> +<p> </p> + +<div class="smaller"> +<p><i>Diagnosis.</i>—Vomerine teeth 4-6; dorsum bright green; white lateral stripe; +eye reddish bronze in life.</p> + +<p><i>Description.</i>—The following description is based on KU 58069 from 6.2 +kilometers south of Rayón Mescalapa, Chiapas, México (<a href="#Plate_17">Pl. 17</a>). Adult male +having a snout-vent length of 27.6 mm.; tibia length, 13.0 mm.; tibia +length/snout-vent length, 47.1 per cent; foot length, 10.8 mm.; head length, +9.6 mm.; head length/snout-vent length, 34.7 per cent; head width, 9.2 mm.; +head width/snout-vent length, 33.1 per cent; diameter of eye, 3.0 mm.; +diameter of tympanum, 1.6 mm.; tympanum/eye, 53.3 per cent. Snout in +lateral profile nearly square, slightly rounded above and below, and in dorsal +profile blunt, almost square; canthus pronounced; loreal region concave; lips +thick, rounded and flaring; nostrils protuberant; internarial distance, 2.5 mm.; internarial +region slightly depressed; top of head flat; interorbital distance, 3.3 +mm., much greater than width of eyelid, 2.5 mm. Thin dermal fold, from +posterior corner of eye above tympanum to insertion of fore limb, covering +upper edge of tympanum; tympanum nearly round, its diameter equal to its +distance from eye. Forearm slender, lacking distinct fold on wrist; row of +low, rounded tubercles on ventrolateral surface of forearm; pollex slightly enlarged, +without nuptial spines; second and fourth fingers equal in length; subarticular +tubercles round, that under fourth finger bifid; discs small, that of +third finger noticeably smaller than tympanum; no web between first and +second fingers; vestige of web between other fingers. Heels overlapping when +hind limbs adpressed; tibiotarsal articulation reaches to middle of eye; no +tarsal fold; inner metatarsal tubercle large, flat, and elliptical; outer metatarsal +tubercle small, elliptical, slightly more distal than inner; subarticular tubercles +round; length of digits from shortest to longest 1-2-5-3-4; third and fifth toes +webbed to base of disc; fourth toe webbed to base of penultimate phalanx; +discs smaller on toes than on fingers. Anal opening directed posteriorly at +upper edge of thighs; no anal flap; pair of large tubercles below anal opening. +Skin of dorsum and of ventral surfaces of forelimbs and shanks smooth; that +of throat, belly, and ventral surfaces of thighs granular. Ventrolateral glands +well developed, not reaching axilla or groin and broadly separated midventrally. +Tongue cordiform, shallowly notched behind and only slightly free +posteriorly; vomerine teeth 2-3, situated on small triangular elevations +<span class='pagenum'><a name="Page_335" id="Page_335">[Pg 335]</a></span> +between ovoid inner nares; openings to vocal sac large, one situated along inner +posterior edge of each mandibular ramus.</p> + +<p>Dorsum of head, body and limbs reddish brown with dark purplish brown +markings on back and shanks; first finger creamy tan; other fingers pale +brown; dorsal surfaces of tarsi, third, fourth, and fifth toes dull tan with +brown spots; first and second toes creamy tan; webbing on feet brown; +anterior and posterior surfaces of thighs tan; faint creamy white stripe along +ventrolateral edges of tarsi and forearms; enamel-white stripe on heel; axilla +and groin gray; enamel-white stripe on edge of upper lip, continuing onto +proximal upper surfaces of forelimb and on flanks to groin, widened under eye +to form large spot, and bordered below on flanks by dark brown stripe; white +stripe above and white spots below anal opening; throat and chest white; +belly and ventral surfaces of limbs cream color; brown dash on either side of +chin and brown spot on throat near angle of jaws; few brown flecks on belly; +ventrolateral glands orange-tan; ventral surfaces of tarsi and feet brown.</p> + +<p>In life, dorsal surfaces of head, body, and limbs bright green (Shamrock +Green); first and second fingers pale orange (Apricot Yellow); stripe on upper +lip and spot below eye enamel-white; stripe on flanks silvery white, bordered +below by brown (Saccardo's Umber) brown; anterior and posterior surfaces of +thighs yellowish brown (Old Gold); webbing of feet dull brown (Brownish +Olive); belly deep yellow (Amber Yellow); iris reddish bronze (English Red).</p> + +<p><i>Variation.</i>—Tubercles beneath the fourth fingers are bifid in 20 specimens +and rounded in all others. The tongue is emarginate in 12 specimens and +cordiform in all others. In most specimens the white stripe on the upper lip +continues onto the flanks and to the groin; in five specimens the stripe terminates +above the forearm, and in three it terminates at mid-flank. The lateral +stripe is absent in two specimens. All specimens were uniform green above +when found at night; later some changed to pale green (Light Oriental Green) +on the dorsum with irregular yellowish tan (Naples Yellow) blotches. Most +males have brown flecks on the throat and ventrolateral gland, but some +specimens are immaculate below, and one has dark brown mottling on the +throat. Several males have a round, orange-tan glandular area on the chin, +as does <i>P. ignicolor</i>.</p> + +<p><i>Description of tadpole.</i>—The following description is based on KU 58199 +from 6.2 kilometers south of Rayón Mescalapa, Chiapas, México (Figs. <a href="#Fig_5">5B</a> and <a href="#Fig_6">6F</a>). Hind limbs small; total length, 39.0 mm.; body length, 10.5 mm.; +body length/total length, 26.9 per cent. Body barely depressed, only slightly +wider than deep, widest just posterior to eyes; in dorsal profile ovoid; mouth +directed ventrally; eyes small, directed dorsolaterally; nostrils barely protuberant, +directed anterodorsally, slightly closer to eye than snout; spiracle +sinistral and posteroventrad to eye; anal tube dextral. Tail long and slender; +caudal fin low, rounded posteriorly; depth of caudal musculature one-half +greatest depth of caudal fin; musculature not extending to tip of tail.</p> + +<p>Mouth large; thin fleshy lips greatly expanded and forming funnel-shaped +disc; outer edge of lips having one row of small papillae; inner surfaces of +mouth smooth; scattered large papillae forming nearly one complete row +around teeth; other papillae laterally; beaks moderately developed, bearing +long, pointed denticulations; no lateral projections on upper beak; tooth-rows +<span class="dividend">3</span>⁄<span class="divisor">3</span>, all short; second and third upper rows subequal in length; first upper row +shorter; first and third upper rows interrupted medially; first lower row +<span class='pagenum'><a name="Page_336" id="Page_336">[Pg 336]</a></span> +interrupted medially, equal in length to second and third upper rows; second lower +row slightly shorter; third lower row shortest.</p> + +<p>Body dark brown above and dark gray below; fleshy part of mouth creamy +gray mottled with dark brown; caudal musculature pale tan with heavy suffusion +of brown flecks; caudal fin transparent with brown spots; dark brown +streak mid-laterally on anterior one-fifth of caudal musculature, bordered below +by cream-colored spot; eye brown in life.</p> + +<p><i>Variation.</i>—The third upper tooth-row is interrupted in all specimens, but +in some individuals the first upper row and first lower row are complete. The +only noted variation in color is the intensity of brown pigmentation on the +caudal musculature, which in most specimens is sufficiently dense to make the +tail look brown. In some specimens the mid-lateral streak is indistinct, and +the pale spot below the streak is absent.</p> + +<p><i>Comparisons.</i>—Aside from the characters listed in the diagnosis, <i>Ptychohyla +schmidtorum chamulae</i> differs from <i>P. schmidtorum schmidtorum</i> by having +dark brown webbing on the feet, instead of pale creamy tan webbing, and in +having in life a yellow venter, instead of a white venter. <i>Ptychohyla ignicolor</i> +also is green in life, but has red flash-colors on the thighs, red webbing on +the feet, and lacks the white lateral stripe diagnostic of <i>P. schmidtorum +chamulae</i>.</p> + +<p><i>Plectrohyla matudai matudai</i> and <i>P. guatemalensis</i> are sympatric with +<i>Ptychohyla schmidtorum chamulae</i>. Each of the first two has a bony prepollex, +rugose skin on the dorsum, and heavy body. Also living with <i>Ptychohyla +chamulae</i> are <i>Hyla chaneque</i>, a large species having a tuberculate dorsum and +webbed fingers, and <i>Hyla bivocata</i>, a small yellow species having a broad, flat +head, small indistinct tympanum, and an axillary membrane.</p> +</div> + +<p><i>Life History.</i>—Calling males were found on leaves of herbs and +bushes by cascading streams in cloud forest. The call consists of +series of short notes, three to nine notes per series, sounding like +"raa-raa-raa." The duration of each note is .054 to .070 of a second, +and has a rate of 96 to 110 pulses per second. The dominant +frequency falls between 3350 and 3450 cycles per second (<a href="#Plate_11">Pl. 11D</a>). +The call is almost indistinguishable from that of <i>Ptychohyla schmidtorum +schmidtorum</i>.</p> + +<p>Tadpoles were found in the cascading streams; the smallest tadpole +has a total length of 17.2 mm. and has only <span class="dividend">3</span>⁄<span class="divisor">2</span> tooth-rows. At a +stream 6.2 kilometers south of Rayón Mescalapa, Chiapas, metamorphosing +young were found on June 16 and August 5, 1960. +Each of two completely metamorphosed young has a snout-vent +length of 15.7 mm. Another having a snout-vent length of 16.2 mm. +has a tail stub 2 mm. long and a completely metamorphosed mouth. +Two others have snout-vent lengths of 13.6 and 14.1 mm. and tail +lengths of 11.5 and 8.1 mm. respectively; in these the mouth parts +are incompletely metamorphosed.</p> + +<p><i>Remarks.</i>—Duellman (1961:354) described <i>Ptychohyla chamulae</i> +<span class='pagenum'><a name="Page_337" id="Page_337">[Pg 337]</a></span> +and stated that it probably was most closely related to <i>P. schmidtorum</i>. +Further study has revealed additional resemblance in morphological +and behavioral details. It is concluded that the two +populations are more realistically treated as subspecies than as +species. The geographic ranges, as now known, are disjunct. +<i>Ptychohyla schmidtorum chamulae</i> inhabits cloud forest on the +Atlantic slopes of the Chiapan Highlands, whereas <i>P. s. schmidtorum</i> +lives in cloud forest on the Pacific slopes of the Sierra Madre +in Chiapas and Guatemala. Between their known geographic +ranges are the pine clad Sierra Madre and Chiapan Highlands, and +intervening sub-humid Grijalva Valley.</p> + +<div class="smaller"> +<p><i>Distribution.</i>—This species is known only from elevations between 1500 and +1700 meters on the Atlantic slopes of the Chiapan Highlands; it is to be expected +in cloud forests on the northern slopes of the Sierra de Cuchumatanes +in Guatemala.</p> + +<p><i>Specimens examined.</i>—<span class="smcap">Mexico</span>: <i>Chiapas: 15 km. N of Pueblo +Nuevo Solistahuacán</i>, UMMZ 123325 (4); <i>16.5 km. N of Pueblo Nuevo +Solistahuacán</i>, UMMZ 123322 (10); <i>18 km. N of Pueblo Nuevo Solistahuacán</i>, +UMMZ 121395-9, 123324 (8), 123326 (5); <i>18.6 km. N of Pueblo Nuevo +Solistahuacán</i>, UMMZ 123323 (4); <i>5.6 km. S of Rayón Mescalapa</i>, KU 58062, +58200 (tadpoles); <i>6.2 km. S of Rayón Mescalapa</i>, KU 58063-74, 58199 (tadpole), +58234-8, 59936 (skeleton).</p> +</div> +<p> </p> + +<a name="ptych_schm5"></a> +<table cellpadding="4" summary="mini_plate frame"> +<tr> +<td class="img_left center"> +Plate 18<br /> +<a href="#Plate_18"><img src="images/plate_18_sm.png" width="164" height="300" border="0" alt="Plate 18 small" /></a><br /> +Click to View Larger. +</td><td> +<div class="caption3"><b>Ptychohyla ignicolor</b> Duellman</div> +<br /> +<div class="smaller"> + <div class="species"><i>Ptychohyla ignicolor</i> Duellman, Uni. Kansas Publ. Mus. Nat. Hist., 13:352-353, + pl. 25, fig. 1, April 27, 1961 [Holotype.—UMMZ 119603 from 6 + kilometers south of Vista Hermosa, Oaxaca, México; Thomas E. Moore + collector].</div> +</div> +</td></tr> +</table> +<p> </p> + +<div class="smaller"> +<p><i>Diagnosis.</i>—Diameter of tympanum less than one-half diameter of eye; +internarial region flat; 3-7 vomerine teeth; toes one-half webbed; no white +spot below eye; no lateral white stripe; in life dorsum green; groin and thighs +having bright red flash-colors.</p> + +<p><i>Description.</i>—The following description is based on UMMZ 119603 from +6 kilometers south of Vista Hermosa, Oaxaca, México (<a href="#Plate_18">Pl. 18</a>). Adult male +having a snout-vent length of 30.0 mm.; tibia length, 14.6 mm.; tibia length/snout-vent +length, 48.7 per cent; foot length, 12.3 mm.; head length, 9.2 mm.; +head length/snout-vent length, 30.7 per cent; head width, 9.3 mm.; head +width/snout-vent length, 31.0 per cent; diameter of eye, 3.2 mm.; diameter +of tympanum, 1.3 mm.; tympanum/eye, 40.6 per cent. Snout in lateral profile +square, and in dorsal profile rounded; canthus pronounced; loreal region +slightly concave; lips moderately flaring; top of head flat; nostrils protuberant; +internarial distance, 2.8 mm.; internarial region flat; interorbital distance, +3.3 mm., much broader than width of eyelid, 2.8 mm. A heavy dermal fold +from posterior corner of eye above tympanum to insertion of forelimb, covering +upper edge of tympanum; tympanum elliptical, its greatest diameter equal +to its distance from eye. Forearm moderately robust having distinct dermal +fold on wrist; pollex moderately enlarged without nuptial spines; second and +fourth fingers equal in length; subarticular tubercles round, none is bifid; +<span class='pagenum'><a name="Page_338" id="Page_338">[Pg 338]</a></span> +discs on fingers moderate in size, that on third finger slightly larger than +tympanum; no web between first and second fingers; vestige of web between +other fingers. Heels overlap when hind limbs adpressed; tibiotarsal articulation +extends to anterior corner of eye; no tarsal fold; inner metatarsal tubercle +large, flat, and elliptical; outer metatarsal tubercle near inner one and triangular +in shape; subarticular tubercles round; length of digits from shortest +to longest 1-2-5-3-4; third toe webbed to proximal end of penultimate phalanx; +fourth toe webbed to distal part of antepenultimate phalanx; fifth toe webbed +to middle of penultimate phalanx; discs on toes smaller than on fingers. Anal +opening directed posteriorly at upper edge of thighs; no anal flap; pair of +large tubercles below anal opening; small tubercles ventral and lateral to these. +Skin of dorsum and ventral surfaces of limbs smooth; that of throat and belly +granular. Ventrolateral glands noticeably thickened, extending from axilla +nearly to groin and only narrowly separated midventrally on chest; skin of +anterior part of chin thickened and glandular. Tongue cordiform, shallowly +notched behind and only slightly free posteriorly; vomerine teeth 0-3, situated +on rounded elevations between somewhat larger, round inner nares; openings +to vocal sac large, one situated along posterior margin of each mandibular +ramus.</p> + +<p>Dorsal ground-color of head, body, and limbs dull brown with dark brown +reticulations on head and body and dark brown transverse bands or spots on +limbs; first and second fingers cream color; third and fourth fingers dull tan; +anterior surfaces of thighs pale brown; posterior surfaces of thighs cream +color with heavy suffusion of brown; dorsal surfaces of tarsi and third, fourth, +and fifth toes dull brown with dark brown spots; first and second toes creamy +white; webbing on foot brown; axilla and groin cream color; flanks brown; no +white stripes on edge of upper lip or on flank; faint, barely discernible tan +streak above anal opening; faint creamy tan line on ventrolateral edges of +tarsi; throat, belly, ventral surfaces of limbs, inner edges of tarsi, and first +toes cream color; outer ventral surfaces of tarsi and other toes brown; chest +and throat spotted with brown; ventrolateral and chin glands orange-brown.</p> + +<p>In life the dorsum was uniform green (Cosse Green) becoming paler green +(Bright Green-Yellow) on flanks, later changing to paler green (Javel Green) +on dorsum with irregular darker green (Lettuce Green) markings and greenish +yellow (Green-Yellow) on flanks; anterior and posterior surfaces of thighs, +ventral surfaces of shanks, anterior surfaces of tarsi, and upper proximal surfaces +of first, second, and third toes red (Coral Red); venter pale creamy +yellow (Sulfur Yellow); iris pale golden color (Aniline Yellow).</p> + +<p><i>Variation.</i>—Of 13 specimens, six have a cordiform tongue; the others have +an emarginate tongue. Five specimens have round subarticular tubercles +beneath the fourth fingers; six specimens have a bifid tubercle on one hand, +and two specimens have bifid tubercles on both hands. A round gland is +present on the chin of all specimens; in some the gland is barely visible, but +in others it is large and distinct. In two specimens the ventrolateral glands +are weakly developed; in the others the glands are well developed and orange-tan. +The white anal stripe varies from a thin line to a series of white flecks. +Dark brown or black flecks are present on the throat, chest, and flanks of all +specimens. In some the flecks are small and widely scattered; in others the +flecks are larger and more numerous. All specimens were pale green above +<span class='pagenum'><a name="Page_339" id="Page_339">[Pg 339]</a></span> +when calling at night; later they changed to dull green with darker green +reticulations. The flash-colors on the thighs and in the groin vary from red +to orange-red or brownish red.</p> + +<p><i>Description of tadpole.</i>—The following description is based on KU 71716 +from Vista Hermosa, Oaxaca, México (Figs. <a href="#Fig_5">5C</a> and <a href="#Fig_6">6G</a>). Hind limbs small; +total length, 39.6 mm.; body length, 11.8 mm.; body length/total length, 29.8 +per cent. Body moderately depressed, only slightly wider than deep, in dorsal +profile ovoid, widest just posterior to eyes; in lateral profile snout rounded; +mouth directed ventrally; eyes small, directed dorsolaterally; nostrils barely +protuberant, directed anteriorly, somewhat closer to eye than snout; spiracle +sinistral and posteroventrad to eye; anal tube dextral. Tail long and slender; +caudal fin low and rounded posteriorly; depth of caudal musculature about +one-half greatest depth of caudal fin; musculature not extending to tip of +tail.</p> + +<p>Mouth large; thin fleshy lips greatly expanded and forming large funnel-shaped +disc; width of mouth about two-thirds greatest width of body; outer +edge of lips having one row of small papillae; inner surface of mouth smooth; +scattered large papillae forming one nearly complete row around teeth; other +papillae laterally; one large papilla just above each end of first lower tooth-row; +beaks moderately developed bearing long, pointed denticulations; no +lateral projections on upper beak; tooth-rows <span class="dividend">3</span>⁄<span class="divisor">3</span>, all short; second and third +upper rows subequal in length; first upper row shorter; first lower row equal +in length to second and third upper rows; second lower row slightly shorter; +third lower row shortest.</p> + +<p>Body creamy gray with dark brown flecks above and below; mouth colored +like body; caudal musculature creamy tan; caudal fin transparent; dark brown +streak on anterior third of caudal musculature; rest of tail and all of caudal fin, +except anterior two-thirds of ventral fin, heavily flecked with brown; iris silvery +bronze color in life.</p> + +<p><i>Variation.</i>—The only other known tadpole, which was collected with the +individual described above, differs in having only two upper tooth-rows. The +first upper tooth-row seems not to have developed.</p> + +<p><i>Comparisons.</i>—From <i>P. schmidtorum chamulae</i> and <i>P. s. schmidtorum</i>, <i>P. +ignicolor</i> differs as follows: Tympanum smaller; snout more nearly square; less +webbing on toes; internarial region flat instead of depressed; white lateral +stripes lacking.</p> + +<p><i>Ptychohyla ignicolor</i> and several small and moderate sized hylids are +sympatric. From <i>P. ignicolor</i> these hylids can be distinguished as follows: +<i>Hyla dendroscarta</i> has a round snout and yellow dorsum; <i>Hyla erythromma</i> +has a round snout, green dorsum, white flanks, and a red eye; <i>Hyla hazelae</i> has +a tarsal fold, green dorsum, and a black line on the canthus; and <i>Ptychohyla +leonhardschultzei</i> has a tarsal fold, brown dorsum, black and white flanks, and +horny nuptial spines in breeding males.</p> +</div> + +<p><i>Life History.</i>—At Vista Hermosa, Oaxaca, males were calling on +vegetation above small streams on March 30, 1959, and on June 28, +1962; males were found on vegetation overhanging a stream 6 kilometers +south of Vista Hermosa on June 27 and July 3, 1962. The +call consists of a series of short notes, three to thirteen notes per +<span class='pagenum'><a name="Page_340" id="Page_340">[Pg 340]</a></span> +series, sounding like "raa-raa-raa." The duration of each note is +about .08 of a second and has a rate of 123 to 129 pulses per second. +The dominant frequency of notes in short series is about 2100 cycles +per second, whereas the dominant frequency of notes in long series +is about 3150 cycles per second (<a href="#Plate_11">Pl. 11E</a>).</p> + +<p>On June 28, 1962, two tadpoles of this species were found in a +quiet pool in a spring-fed rivulet at Vista Hermosa, Oaxaca. Females +are unknown.</p> + +<p><i>Remarks.</i>—The absence of a tarsal fold and of nuptial spines in +breeding males, the nature of the breeding call, and the form of +tadpole are characters that place <i>Ptychohyla ignicolor</i> in the <i>P. +schmidtorum</i>-group.</p> + +<div class="smaller"> +<p><i>Distribution.</i>—This species is known from only two localities at elevations +of 1500 and 1850 meters in the cloud forest on the northern (Atlantic) slopes +of the Sierra Madre Oriental in northern Oaxaca.</p> + +<p><i>Specimens examined.</i>—<span class="smcap">Mexico</span>: <i>Oaxaca: Vista Hermosa</i>, KU 71334, 71716 +(tadpoles), UMMZ 119602; 6 km. S of Vista Hermosa, KU 71335-42, 71343 +(skeleton), UMMZ 119603, 123327 (2).</p> +</div> +<p> </p> + +<a name="dist_and_ecology"></a> +<div class="caption2">DISTRIBUTION AND ECOLOGY</div> +<p> </p> + +<a name="geogr_dist"></a> +<div class="caption3">Geographic Distribution of the Species</div> + +<p>The distribution of species of <i>Ptychohyla</i> reflects the distribution +of cloud forest in southern México and northern Central America. +The frogs are restricted to mountainous areas, usually at elevations +higher than 1000 meters above sea level. <i>Ptychohyla</i> does not range +to great heights in the mountains, where west of the Isthmus of +Tehuantepec the mountain streams are inhabited by frogs of the +<i>Hyla bistincta</i> group, and in Chiapas and Guatemala by species of +<i>Plectrohyla</i>.</p> + +<p>Frogs of the <i>Ptychohyla euthysanota</i> group have a greater combined +geographic range than the species comprising the <i>Ptychohyla +schmidtorum</i> group (Fig. 7). No two species in the same group +are sympatric, but members of different groups are sympatric in at +least parts of their ranges. Apparently <i>P. leonhardschultzei</i> ranges +around the southern edge of the Mexican Highlands, where the +species occurs on both Atlantic and Pacific slopes; as can be seen +from the distribution map, there are many gaps in the known range +of this species. The range of <i>P. euthysanota euthysanota</i> is along +the Pacific slopes of the Sierra Madre in Chiapas, Guatemala, and +El Salvador, whereas that of <i>P. euthysanota macrotympanum</i> is +along the southern interior slopes of the Central Highlands of +Chiapas and the Sierra de Cuchumatanes in Guatemala. <i>Ptychohyla</i> +<span class='pagenum'><a name="Page_341" id="Page_341">[Pg 341]</a></span> +<i>spinipollex</i> occurs on the wet Atlantic slopes of the Guatemalan and +Honduranean Highlands; the range of the species in Honduras is +poorly known.</p> +<p> </p> + +<a name="Fig_7"></a> +<div class="center"> +<img src="images/fig_7.png" width="600" height="291" border="0" alt="Map showing record localities." title="Map showing locality records for the species and subspecies of +Ptychohyla" /> +<p><span class="smcap">Fig. 7.</span> Map showing locality records for the species and subspecies of +<i>Ptychohyla</i>.</p> +</div> +<p> </p> + +<p>The frogs of the <i>Ptychohyla schmidtorum</i> group have more restricted +geographic ranges than members of the former group. +<i>Ptychohyla schmidtorum schmidtorum</i> occurs on the Pacific slopes +of the Sierra Madre in Chiapas and Guatemala, where it occurs +with <i>P. euthysanota euthysanota</i>; <i>P. schmidtorum chamulae</i> is +known from only two localities on the Atlantic slopes of the +Central Highlands of Chiapas, where it occurs close to, but as +now known not with, <i>P. euthysanota macrotympanum</i>. On the +Atlantic slopes of the Sierra Madre Oriental in northern Oaxaca <i>P. +ignicolor</i> occurs with <i>P. leonhardschultzei</i>.</p> + +<p>In the Sierra de los Tuxtlas in southern Veracruz and in the cloud +forests along the eastern slopes of the Sierra Madre Oriental northward +to Nuevo León, <i>Hyla miotympanum</i> seems to be the ecological +replacement of <i>Ptychohyla</i>. On the Pacific slopes north of +Guerrero, México, humid forests in which there are cascading mountain +streams are absent; consequently, no <i>Ptychohyla</i> are known +from that region. In the mountains of El Salvador <i>Ptychohyla +euthysanota euthysanota</i> occurs sympatrically with another small +stream-breeding hylid, <i>Hyla salvadorensis</i>. To the south of Honduras +the highlands diminish into the lowlands of Nicaragua, where +habitat suitable for <i>Ptychohyla</i> apparently does not exist. In the +mountains of Costa Rica and Panamá, the habitats occupied by +<i>Ptychohyla</i> in northern Central America are filled by a variety of +<span class='pagenum'><a name="Page_342" id="Page_342">[Pg 342]</a></span> +stream-breeding <i>Hyla</i>, such as <i>Hyla legleri</i>, <i>H. rivularis</i>, <i>H. rufioculis</i>, +<i>H. alleei</i>, and <i>H. uranochroa</i>.</p> + +<p>Although members of the genus <i>Ptychohyla</i> occur in the southern +part of the Mexican Highlands to the west of the Isthmus of +Tehuantepec, the greater distribution and differentiation in the +genus is in the Chiapan-Guatemalan Highlands. In this respect +<i>Ptychohyla</i> is a counterpart of <i>Plectrohyla</i>.</p> + +<a name="habitat_pref"></a> +<div class="caption3">Habitat Preference</div> + +<p>Frogs of the genus <i>Ptychohyla</i> are ecologically associated with +mountain streams at elevations between 650 and 2200 meters; in +the geographic region where these frogs occur the vegetation between +those elevations consists of cloud forest or pine-oak forest. +In some places the frogs have been found in a mixture of oak and +semi-deciduous scrub forest. At Vista Hermosa, Oaxaca, <i>P. leonhardschultzei</i> +and <i>P. ignicolor</i> were found in cloud forest, whereas +at Agua del Obispo, Guerrero, the former species was found in pine-oak +forest. <i>Ptychohyla schmidtorum</i> is known only from cloud +forest; <i>P. euthysanota euthysanota</i> and <i>P. spinipollex</i> generally are +found in cloud forest, but in some places they live in pine-oak forest. +<i>Ptychohyla euthysanota macrotympanum</i> has been found in pine-oak +forest and in a mixture of oak and semi-deciduous scrub forest. +With the possible exception of the members of the <i>Ptychohyla +schmidtorum</i> group, which has been found only in cloud forest, +it seems as though the type of vegetation is not the controlling +factor in the ecological distribution of these frogs.</p> + +<p><i>Ptychohyla</i> has been found only where there are clear, cascading +streams overhung by vegetation, on which adults and young perch +at night, or even by day. The presence of these streams, in which +the tadpoles live, seems to be an important factor in the distribution +of <i>Ptychohyla</i>. As has been shown previously, the tadpoles of +<i>Ptychohyla</i> are adapted for existence in torrential streams, where +the water is cool, and the amount of oxygen is high. Clearly +these tadpoles are unsuited for life in ponds or sluggish streams in +the lowlands, where the temperature of the water is high, a layer +of silt on the bottom is deep, and the amount of oxygen is low. The +tadpoles cling to rocks on the bottom of the streams; there they +move slowly across the rocks, apparently feeding on the thin covering +of algae. Tadpoles were not observed on rocks having a thick +covering of algae or moss. The tadpoles were observed to swim +against the current in torrential streams, in which no fishes were +<span class='pagenum'><a name="Page_343" id="Page_343">[Pg 343]</a></span> +found. Therefore, it seems as though the presence of the stream-habitat +for the tadpoles is a significant factor in the ecological distribution +of the species of <i>Ptychohyla</i>.</p> + +<a name="intersp_comp"></a> +<div class="caption3">Interspecific Competition</div> + +<p>At localities where two species of <i>Ptychohyla</i> occur sympatrically +(<i>P. ignicolor</i> and <i>P. leonhardschultzei</i> at Vista Hermosa, Oaxaca, +and <i>P. euthysanota euthysanota</i> and <i>P. schmidtorum schmidtorum</i> +at Finca La Paz, Depto. San Marcos, Guatemala) effort was made +to determine what, if any, ecological interspecific relationships +existed. Although adults of the sympatric species were found on +adjacent leaves or branches of bushes overhanging the streams at +both localities, segregation at the time of breeding seems to be +maintained by the notably different breeding calls in sympatric +species (see discussion of breeding calls). Thus, as has been shown +by Blair (1956), Fouquette (1960), and others working on a variety +of pond-breeding frogs and toads, the breeding call in <i>Ptychohyla</i> +acts as an important reproductive isolating mechanism.</p> + +<p>At no locality were <i>Ptychohyla</i> and associated species of hylids +found so abundantly as were species of pond-breeding hylids in the +lowlands. Apparently reproductive activity is not concentrated in a +short breeding season, and it is highly doubtful if the populations +of these frogs are as large as those of the lowland pond-breeders. +The continual humid conditions and abundance of insect food +throughout the year in the cloud forest are perhaps indicative of +little interspecific competition among adults of <i>Ptychohyla</i> and +other sympatric hylids.</p> + +<p>At Finca La Paz, Guatemala, tadpoles of two species of <i>Ptychohyla</i> +were ecologically segregated. The tadpoles of <i>P. euthysanota +euthysanota</i> were found in riffles in the streams, whereas those of +<i>P. schmidtorum schmidtorum</i> were found in slower water, chiefly +in small pools in the streams. At Vista Hermosa, Oaxaca, tadpoles +of <i>P. leonhardschultzei</i> were found in riffles, and tadpoles of the +sympatric <i>P. ignicolor</i> were found in a small pool in a stream. +Similar ecological relationships were observed for several species of +Costa Rican hylids. Throughout the range of <i>Ptychohyla</i> east of +the Isthmus of Tehuantepec, members of the genus occur with +species of <i>Plectrohyla</i>, all of which are larger than <i>Ptychohyla</i>, and +all of which have tadpoles that live in torrential streams. Tadpoles +of <i>Ptychohyla spinipollex</i> have been found in streams inhabited by +the tadpoles of <i>Plectrohyla guatemalensis</i> and <i>P. quecchi</i>; tadpoles +of <i>Ptychohyla euthysanota euthysanota</i> and <i>P. schmidtorum schmidtorum</i> +<span class='pagenum'><a name="Page_344" id="Page_344">[Pg 344]</a></span> +were found in streams inhabited by tadpoles of <i>Plectrohyla +guatemalensis</i>, <i>P. matudai</i>, and <i>P. sagorum</i>. In some streams great +numbers of tadpoles occur. The habitat is rather restricted, and +the food supply is limited. Consequently, interspecific competition +among the various species of hylids whose tadpoles live in the torrential +streams probably is highest during the larval stage. Unfortunately, +this aspect of salientian population ecology has received +no intensive study.</p> + +<a name="repro_dev"></a> +<div class="caption3">Reproduction and Development</div> + +<p>Since the cloud forests inhabited by <i>Ptychohyla</i> are daily bathed +in clouds and have a fairly evenly distributed rainfall throughout +the year, the frogs living in these forests are active throughout the +year. At least some of the species evidently have a long breeding +season, for I found calling males of <i>P. leonhardschultzei</i> in February, +March, and August, and found tadpoles in February, March, +June, and August. Tadpoles of the various species have been obtained +throughout much of the year, as follows: <i>P. euthysanota +euthysanota</i>, February, March, May, and July; <i>P. euthysanota +macrotympanum</i>, March, June, and August; <i>P. spinipollex</i>, February, +March, April, June, July, and August; <i>P. schmidtorum +schmidtorum</i>, March, May, June, July, and August; <i>P. schmidtorum +chamulae</i>, June and August; <i>P. ignicolor</i>, June. I suspect that this +temporal distribution more accurately reflects the seasonal activities +of collectors than of the frogs.</p> + +<p>Calling frogs usually are on vegetation adjacent to or overhanging +streams; some calling males of <i>P. spinipollex</i> were on rocks in +or by streams. Clasping pairs of <i>P. euthysanota</i> and <i>P. schmidtorum</i> +were observed on vegetation by streams. Despite intensive +search, no eggs were found. It is doubtful if <i>Ptychohyla</i> deposit +eggs on vegetation overhanging streams, as do centrolenids and +<i>Phyllomedusa</i>, for egg-clutches of these frogs are easily found. +Possibly the eggs are laid separately on vegetation above the stream, +in which case they could be overlooked easily. In streams where +<i>Ptychohyla</i> and other hylid tadpoles occur, empty egg capsules +have been found on the lee sides of rocks, but there is no way to +determine which species laid the eggs.</p> + +<p>Numbers of eggs were counted in gravid females; the largest eggs +have diameters ranging from 2.5 to 3.0 mm. The smaller species, +comprising the <i>Ptychohyla schmidtorum</i> group, have fewer eggs +than do the larger species. Numbers of eggs found in females of +the various species are: <i>P. euthysanota euthysanota</i>, 108; <i>P. euthysanota</i> +<span class='pagenum'><a name="Page_345" id="Page_345">[Pg 345]</a></span> +<i>macrotympanum</i>, 136, 160; <i>P. leonhardschultzei</i>, 141; <i>P. +spinipollex</i>, 119, 134, 143; <i>P. schmidtorum schmidtorum</i>, 59, 61, 90; +<i>P. schmidtorum chamulae</i>, 60, 71, 89.</p> + +<p>Duration of the larval stage is unknown. Metamorphosing young +have been found from May through August. From two to six completely +metamorphosed young are available for each of the species, +except for <i>P. ignicolor</i> of which none is available. The smallest +young frog is a <i>P. euthysanota</i> having a snout-vent length of +14.2 mm.; the largest young frog is a <i>P. schmidtorum schmidtorum</i> +having a snout-vent length of 17.0 mm.</p> +<p> </p> +<p> </p> + +<a name="phylogeny" id="phylogeny"></a> +<div class="caption2">PHYLOGENY OF PTYCHOHYLA</div> + +<p>The preceding data on morphology, life histories, and behavior +form the basis for the following interpretation of the phylogeny of +<i>Ptychohyla</i>. Additional data are needed to support some of the +ideas discussed below; many of the data that are available for +<i>Ptychohyla</i> are lacking for other, possibly related, hylids. The +family Hylidae is composed of several hundred species, and most +of the species are poorly known. Consequently, any attempt to +place <i>Ptychohyla</i> in the over-all scheme of hylid phylogeny would +be premature at this time. But, as between the five species of two +species-groups here recognized as constituting the genus <i>Ptychohyla</i>, +some estimate of relationships can be made. First, it is necessary +to determine the validity of the genus itself.</p> + +<a name="nat_assemb"></a> +<div class="caption3"><i>Ptychohyla</i> as a Natural Assemblage</div> + +<p>As stated in the diagnosis of the genus, the only character that +sets this group of species apart from other hylids is the presence of +ventrolateral glands in the breeding males. To many systematists +the thought of being able to identify to genus only breeding males +is sufficiently disturbing to cause them to view with disfavor the +recognition of the genus. Nonetheless, the question is raised: Do +the five species herein placed in the genus <i>Ptychohyla</i> constitute a +natural assemblage? If the genus is considered to be more than a +category of convenience, that is to say, a group of related species +having a common origin, the primary problem is to determine +whether or not the five species form a phylogenetic unit.</p> + +<p>The species of <i>Ptychohyla</i> are divided into two groups on the +basis of external morphology, breeding calls, and tadpoles. The +<i>Ptychohyla euthysanota</i> group seems to be a natural group composed +of three species, all of which are more closely related to one +another than to any other hylid. Likewise, the species comprising +<span class='pagenum'><a name="Page_346" id="Page_346">[Pg 346]</a></span> +the <i>Ptychohyla schmidtorum</i> group seem to represent a natural unit. +If the presence of ventrolateral glands in breeding males is ignored, +a student of salientian systematics might derive the <i>Ptychohyla +euthysanota</i> group from a hylid stock containing <i>Hyla miotympanum</i> +and <i>Hyla mixomaculata</i>. Likewise, <i>Ptychohyla schmidtorum</i> +could be placed with <i>Hyla uranochroa</i> and related species in +Costa Rica. Nonetheless, the fact remains that all of the species +assigned to the genus <i>Ptychohyla</i> have ventrolateral glands in the +breeding males; furthermore, ventrolateral glands are unknown in +other hylids. If the <i>P. schmidtorum</i> group and the <i>P. euthysanota</i> +group each evolved from separate hylid stocks, then the ventrolateral +glands must have developed independently in both groups. +That ventrolateral glands developed independently in five species +of frogs in southern México and northern Central America and not +in any of the other approximately 500 species of hylids in the world +is untenable. It is more logical to assume that the development of +the glands took place only once in a stock of hylids that gave rise +to the five species herein recognized as members of the genus +<i>Ptychohyla</i>.</p> + +<a name="generic_rel"></a> +<div class="caption3">Generic Relationships</div> + +<p>The affinities of <i>Ptychohyla</i> apparently are not with any of the +other groups that have been generically separated from <i>Hyla</i>. Of +the daughter genera in Middle America only <i>Plectrohyla</i> has stream-adapted +tadpoles, but these large frogs are not closely related to +<i>Ptychohyla</i>. Stuart (1954:169) suggested that certain montane +species of <i>Hyla</i> in lower Central America and <i>Hyla salvadorensis</i> +in El Salvador may be related to <i>Ptychohyla</i> or even congeneric. +I have had experience with most of these species in the field and +believe that Stuart was correct in his suggestion of relationships. +The species concerned are four red-eyed stream-breeding <i>Hyla</i> in +Costa Rica—<i>H. alleei</i>, <i>H. legleri</i>, <i>H. rufioculis</i>, and <i>H. uranochroa</i>, +plus <i>Hyla salvadorensis</i> in the mountains of El Salvador. Morphologically +all of the species are similar; <i>Hyla uranochroa</i>, <i>H. +legleri</i>, and <i>H. rufioculis</i> have a lateral white stripe that is expanded +to form a spot beneath the eye, as in <i>Ptychohyla schmidtorum</i>. The +tadpoles of <i>Hyla rufioculis</i> and <i>H. uranochroa</i> have large funnel-shaped +mouths and long slender tails like those of <i>Ptychohyla +schmidtorum</i>. Lips of the tadpoles of <i>H. legleri</i> and <i>H. salvadorensis</i> +are folded laterally, in this respect resembling those of the <i>Ptychohyla +euthysanota</i> group. I do not know the tadpoles and the +breeding call of <i>Hyla alleei</i>. The breeding calls of <i>Hyla rufioculis</i> +<span class='pagenum'><a name="Page_347" id="Page_347">[Pg 347]</a></span> +and <i>H. uranochroa</i> consist of high melodious notes; the calls of +<i>H. legleri</i> and <i>H. salvadorensis</i> consist of series of short notes that +have the general characteristics of the call of <i>Ptychohyla schmidtorum</i>. +Affinities of the genus <i>Ptychohyla</i> seem to me to be with +the red-eyed species forming the <i>Hyla uranochroa</i> group in Costa +Rica. All of the species in the <i>Hyla uranochroa</i> group have large +frontoparietal fontanelles, rather small ethmoids, and small nasals +that are not in contact with one another or with the ethmoid. Some +species have a complete quadratojugal-maxillary arch; others do +not. Assuming that the parental stock that gave rise both to the +<i>Hyla uranochroa</i> group and to <i>Ptychohyla</i> was widespread in +Central America at a time of cooler, more humid conditions, it is +possible that with subsequent warming temperatures and seasonal +rainfall in the lowlands the parental stock was restricted to the +Costa Rican highlands, where the <i>Hyla uranochroa</i> group developed, +and to the Chiapas-Guatemala highlands, where <i>Ptychohyla</i> +evolved.</p> + +<a name="intersp_rel"></a> +<div class="caption3">Interspecific Relationships</div> + +<p><i>Ptychohyla schmidtorum</i> is thought to resemble more closely the +parental stock of the genus than does any other species of <i>Ptychohyla</i> +now extant. This parental stock is discussed above in the +account of the generic relationships. <i>Ptychohyla schmidtorum</i> +has a red eye, white lateral stripe, frontoparietal fontanelle, funnel-shaped +mouth in tadpoles, and lacks nuptial spines; in all of these +characters it resembles members of the <i>Hyla uranochroa</i> group. +Probably during times of glaciation during the Pleistocene, when +climates in México and Central America were depressed, the +<i>Ptychohyla</i> stock was more widespread than it is now. Subsequent +elevation of climatic zones during interglacial periods would have +isolated populations as they are today in regions of cloud forests. +Thus, through geographic isolation populations could have differentiated +and evolved into the present species. Climatic fluctuation +in the Pleistocene must have been of sufficient magnitude to +permit the spread of cool, moist forests containing <i>Ptychohyla</i> across +the Isthmus of Tehuantepec into the mountains of Oaxaca.</p> + +<p>Because of its small nuptial spines, small triangular vomers, +coloration, and absence of a rostral keel, <i>Ptychohyla euthysanota</i>, +more than any of the other species in the <i>P. euthysanota</i> group, resembles +<i>P. schmidtorum</i>. At the present time <i>P. euthysanota</i> and +<i>P. schmidtorum</i> are sympatric.</p> + +<p>As I have mentioned previously, ecological segregation and +<span class='pagenum'><a name="Page_348" id="Page_348">[Pg 348]</a></span> +interspecific competition probably is highly developed in the tadpoles +of <i>Ptychohyla</i>. If this ecological segregation resulted from intraspecific +competition in a stock of <i>Ptychohyla</i>, possibly <i>P. euthysanota</i> +and <i>P. schmidtorum</i> differentiated sympatrically in this way. +Specific identity is maintained, at least in part, by different breeding +calls in males.</p> + +<p><i>Ptychohyla spinipollex</i> and <i>P. leonhardschultzei</i> seem to be more +closely related to one another than either is to <i>P. euthysanota</i>. +Probably a stock of <i>P. euthysanota</i> was isolated on the Atlantic +slopes of northern Central America from <i>P. euthysanota</i> on the +southern slopes. The frogs on the Atlantic slopes differentiated and +spread into the mountains of Oaxaca, where through isolation by +the barrier of the Isthmus of Tehuantepec they developed into <i>P. +leonhardschultzei</i>, while the stock on the northern slopes of Central +America evolved into <i>P. spinipollex</i>. Subsequent to the differentiation +of <i>P. leonhardschultzei</i> and <i>P. spinipollex</i> from <i>P. euthysanota</i> +and during a time of cooler more equable climate than exists now, +<i>P. euthysanota</i> and <i>P. schmidtorum</i> invaded the Central Highlands +of Chiapas. Subsequent climatic changes isolated populations of +each in the Central Highlands, where <i>P. euthysanota macrotympanum</i> +and <i>P. schmidtorum chamulae</i> evolved. <i>Ptychohyla +ignicolor</i> probably represents stock of <i>P. schmidtorum</i> that crossed +the Isthmus of Tehuantepec and became isolated in Oaxaca on the +western side of the isthmus.</p> +<p> </p> +<p> </p> + +<p><span class='pagenum'><a name="Plate_12" id="Plate_12">[Pl 12]</a></span></p> +<div class="center"> +<div class="caption3">PLATE 12</div> +<img src="images/plate_12_lg.png" width="329" height="600" border="0" alt="Ptychohyla euthysanota euthysanota" title="Ptychohyla euthysanota euthysanota" /> +<br /> +<i>Ptychohyla euthysanota euthysanota</i> (KU 58008). +× 2.<br /> +</div> +<p> </p> +<p> </p> + +<span class='pagenum'><a name="Plate_13" id="Plate_13">[Pl 13]</a></span> +<div class="center"> +<div class="caption3">PLATE 13</div> +<img src="images/plate_13_lg.png" width="288" height="600" border="0" alt="Ptychohyla euthysanota macrotympanum" title="Ptychohyla euthysanota macrotympanum" /> +<br /> +<i>Ptychohyla euthysanota macrotympanum</i> (KU 58049). × 2.<br /> +</div> +<p> </p> +<p> </p> + +<span class='pagenum'><a name="Plate_14" id="Plate_14">[Pl 14]</a></span> +<div class="center"> +<div class="caption3">PLATE 14</div> +<img src="images/plate_14_lg.png" width="254" height="600" border="0" alt="Ptychohyla leonhardschultzei" title="Ptychohyla leonhardschultzei" /> +<br /> +<i>Ptychohyla leonhardschultzei</i> (KU 64117). × 2.<br /> +</div> +<p> </p> +<p> </p> + +<span class='pagenum'><a name="Plate_15" id="Plate_15">[Pl 15]</a></span> +<div class="center"> +<div class="caption3">PLATE 15</div> +<img src="images/plate_15_lg.png" width="257" height="600" border="0" alt="Ptychohyla spinipollex" title="Ptychohyla spinipollex" /> +<br /> +<i>Ptychohyla spinipollex</i> (KU 58054). × 2.<br /> +</div> +<p> </p> +<p> </p> + +<span class='pagenum'><a name="Plate_16" id="Plate_16">[Pl 16]</a></span> +<div class="center"> +<div class="caption3">PLATE 16</div> +<img src="images/plate_16_lg.png" width="287" height="600" border="0" alt="Ptychohyla schmidtorum schmidtorum" title="Ptychohyla schmidtorum schmidtorum" /> +<br /> +<i>Ptychohyla schmidtorum schmidtorum</i> +(KU 58043). × 2.<br /> +</div> +<p> </p> +<p> </p> + +<p><span class='pagenum'><a name="Plate_17" id="Plate_17">[Pl 17]</a></span></p> +<div class="center"> +<div class="caption3">PLATE 17</div> +<img src="images/plate_17_lg.png" width="276" height="600" border="0" alt="Ptychohyla schmidtorum chamulae" title="Ptychohyla schmidtorum chamulae" /> +<br /> +<i>Ptychohyla schmidtorum chamulae</i> (KU +58069). × 2.<br /> +</div> +<p> </p> +<p> </p> + +<p><span class='pagenum'><a name="Plate_18" id="Plate_18">[Pl 18]</a></span></p> +<div class="center"> +<div class="caption3">PLATE 18</div> +<img src="images/plate_18_lg.png" width="272" height="600" border="0" alt="Ptychohyla ignicolor" title="Ptychohyla ignicolor" /> +<br /> +<i>Ptychohyla ignicolor</i> (UMMZ 119603). +× 2.<br /> +</div> +<p> </p> +<p> </p> + +<span class='pagenum'><a name="Page_349" id="Page_349">[Pg 349]</a></span> + +<hr style="width: 65%;" /> +<a name="literature_cited"></a> +<div class="caption2">LITERATURE CITED</div> +<p> </p> + +<span class="smcap">Ahl, E.</span><br /> +<div class="references"><p>1934. Über eine sammlung von Reptilien und Amphibien aus Mexiko,<br /> +Zool. Anz., 106:184-186, April 15.</p></div> + +<span class="smcap">Blair, W. F.</span><br /> +<div class="references"><p>1956. Call difference as an isolation mechanism in southwestern toads<br /> +(genus <i>Bufo</i>). Texas Jour. Sci., 8:87-106, March.</p></div> + +<span class="smcap">Duellman, W. E.</span><br /> +<div class="references"><p>1956. The frogs of the hylid genus <i>Phrynohyas</i> Fitzinger, 1843. Misc.<br /> +Publ. Mus. Zool. Univ. Michigan, 96:1-47, pls. 1-6, February 21.</p></div> + +<div class="references"><p>1960. Synonymy, variation, and distribution of <i>Ptychohyla leonhard-schultzei</i><br /> +Ahl. Studies of American Hylid Frogs, IV. Herpetologica,<br /> +16:191-197, September 23.</p></div> + +<div class="references"><p>1961. Descriptions of two species of frogs, genus Ptychohyla. Studies of<br /> +American Hylid Frogs, V. Univ. Kansas Publ. Mus. Nat. Hist.,<br /> +13:349-357, pl. 25, April 27.</p></div> + +<span class="smcap">Fouquette, M. J.</span><br /> +<div class="references"><p>1960. Isolating mechanisms in three sympatric treefrogs in the Canal<br /> +Zone. Evolution, 14:484-497, December.</p></div> + +<span class="smcap">Kellogg, R.</span><br /> +<div class="references"><p>1928. An apparently new <i>Hyla</i> from El Salvador. Proc. Biol. Soc. Washington,<br /> +41:123-124, June 29.</p></div> + +<span class="smcap">Mertens, R.</span><br /> +<div class="references"><p>1952. Die Amphibien und Reptilien von El Salvador. Senckenbergischen<br /> +Naturf. Gesell., 487:1-120, pls. 1-16, December 1.</p></div> + +<span class="smcap">Ridgway, R.</span><br /> +<div class="references"><p>1912. Color standards and color nomenclature. Washington, D. C., 44<br /> +pp., 53 pls.</p></div> + +<span class="smcap">Shannon, F. A.</span><br /> +<div class="references"><p>1951. Notes on a herpetological collection from Oaxaca and other localities<br /> +in Mexico. Proc. U. S. Nat. Mus., 101:465-484, May 17.</p></div> + +<span class="smcap">Stuart, L. C.</span><br /> +<div class="references"><p>1954. Descriptions of some new amphibians and reptiles from Guatemala.<br /> +Proc. Biol. Soc. Washington, 67:159-178, August 5.</p></div> + +<span class="smcap">Tanner, W. W.</span><br /> +<div class="references"><p>1957. Notes on a collection of amphibians and reptiles from southern<br /> +Mexico, with a description of a new <i>Hyla</i>. Great Basin Nat.,<br /> +17:52-56, July 31.</p></div> + +<span class="smcap">Taylor, E. H.</span><br /> +<div class="references"><p>1937. New species of hylid frogs from Mexico with comments on the rare<br /> +<i>Hyla bistincta</i> Cope. Proc. Biol. Soc. Washington, 50:43-54, pls.<br /> +2-3, April 21.</p></div> + +<div class="references"><p>1942. New tailless amphibia from Mexico. Univ. Kansas Sci. Bull., 28:<br /> +67-89, May 15.</p></div> + +<div class="references"><p>1944. A new genus and species of Mexican frogs. Univ. Kansas Sci.<br /> +Bull., 30:41-45, June 12.</p></div> + +<div class="references"><p>1949. New or unusual Mexican amphibians. Amer. Mus. Novitates,<br /> +1437:1-21, December 7.</p></div> + +<p><i>Transmitted December 27, 1962.</i></p> +<p> </p> +<p> </p> + +<p><span class='pagenum'><a name="Pub_1" id="Pub_1">[Pub_1]</a></span></p> +<hr style="width: 65%;" /> +<a name="UNIVERSITY_OF_KANSAS_PUBLICATIONS" id="UNIVERSITY_OF_KANSAS_PUBLICATIONS"></a> +<div class="caption2">UNIVERSITY OF KANSAS PUBLICATIONS<br /> +MUSEUM OF NATURAL HISTORY</div> + +<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. +Nevertheless, 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> + +<table class="pub_list" summary="pub_list"> +<tr><td class="text_rt"> Vol. 1.</td><td> </td><td>Nos. 1-26 and index. Pp. 1-638, 1946-1950.</td></tr> + +<tr><td class="text_rt">*Vol. 2.</td><td colspan="2" class="justify">(Complete) Mammals of Washington. By Walter W. Dalquest. Pp. 1-444, 140 +figures in text. April 9, 1948.</td></tr> + +<tr><td class="text_rt">Vol. 3.</td><td class="text_rt">*1.</td><td class="justify">The avifauna of Micronesia, its origin, evolution, and distribution. By Rollin +H. Baker. Pp. 1-359, 16 figures in text. June 12, 1951.</td></tr> + +<tr><td> </td><td class="text_rt">*2.</td><td class="justify">A quantitative study of the nocturnal migration of birds. By George H. +Lowery, Jr. Pp. 361-472, 47 figures in text. June 29, 1951.</td></tr> + +<tr><td> </td><td class="text_rt">3.</td><td class="justify">Phylogeny of the waxwings and allied birds. By M. Dale Arvey. Pp. 473-530, +49 figures in text, 13 tables. October 10, 1951.</td></tr> + +<tr><td> </td><td class="text_rt">*4.</td><td class="justify">Birds from the state of Veracruz, Mexico. By George H. Lowery, Jr., and +Walter W. Dalquest. Pp. 531-649, 7 figures in text, 2 tables. October 10, 1951.</td></tr> + +<tr><td> </td><td colspan="2" class="justify">Index. Pp. 651-681.</td></tr> + +<tr><td class="text_rt">*Vol. 4.</td><td colspan="2" class="justify">(Complete) American weasels. By E. Raymond Hall. Pp. 1-466, 41 plates, 31 +figures in text. December 27, 1951.</td></tr> + +<tr><td class="text_rt">Vol. 5.</td><td colspan="2" class="justify">Nos. 1-37 and index. Pp. 1-676, 1951-1953.</td></tr> + +<tr><td class="text_rt">*Vol. 6.</td><td colspan="2" class="justify">(Complete) Mammals of Utah, <i>taxonomy and distribution</i>. By Stephen D. +Durrant. Pp. 1-549, 91 figures in text, 30 tables. August 10, 1952.</td></tr> + +<tr><td class="text_rt">Vol. 7.</td><td colspan="2" class="justify">Nos. 1-15 and index. Pp. 1-651, 1952-1955.</td></tr> + +<tr><td class="text_rt">Vol. 8.</td><td colspan="2" class="justify">Nos. 1-10 and index. Pp. 1-675, 1954-1956.</td></tr> + +<tr><td class="text_rt">Vol. 9.</td><td class="text_rt">*1.</td><td class="justify">Speciation of the wandering shrew. By James S. Findley. Pp. 1-68, 18 +figures in text. December 10, 1955.</td></tr> + +<tr><td> </td><td class="text_rt">2.</td><td class="justify">Additional records and extension of ranges of mammals from Utah. By +Stephen D. Durrant, M. Raymond Lee, and Richard M. Hansen. Pp. 69-80. +December 10, 1955.</td></tr> + +<tr><td> </td><td class="text_rt">3.</td><td class="justify">A new long-eared myotis (Myotis evotis) from northeastern Mexico. By Rollin +H. Baker and Howard J. Stains. Pp. 81-84. December 10, 1955.</td></tr> + +<tr><td> </td><td class="text_rt">4.</td><td class="justify">Subspeciation in the meadow mouse, Microtus pennsylvanicus, in Wyoming. +By Sydney Anderson. Pp. 85-104, 2 figures in text. May 10, 1956.</td></tr> + +<tr><td> </td><td class="text_rt">5.</td><td class="justify">The condylarth genus Ellipsodon. By Robert W. Wilson. Pp. 105-116, 6 +figures in text. May 19, 1956.</td></tr> + +<tr><td> </td><td class="text_rt">6.</td><td class="justify">Additional remains of the multituberculate genus Eucosmodon. By Robert +W. Wilson. Pp. 117-123, 10 figures in text. May 19, 1956.</td></tr> + +<tr><td> </td><td class="text_rt">7.</td><td class="justify">Mammals of Coahulia, Mexico. By Rollin H. Baker. Pp. 125-335, 75 figures +in text. June 15, 1956.</td></tr> + +<tr><td> </td><td class="text_rt">8.</td><td class="justify">Comments on the taxonomic status of Apodemus peninsulae, with description +of a new subspecies from North China. By J. Knox Jones, Jr. Pp. 337-346, +1 figure in text, 1 table. August 15, 1956.</td></tr> + +<tr><td> </td><td class="text_rt">9.</td><td class="justify">Extensions of known ranges of Mexican bats. By Sydney Anderson. Pp. +347-351. August 15, 1956.</td></tr> + +<tr><td> </td><td class="text_rt">10.</td><td class="justify">A new bat (Genus Leptonycteris) from Coahulia. By Howard J. Stains. +Pp. 353-356. January 21, 1957.</td></tr> + +<tr><td> </td><td class="text_rt">11.</td><td class="justify">A new species of pocket gopher (Genus Pappogeomys) from Jalisco, Mexico. +By Robert J. Russell. Pp. 357-361. January 21, 1957.</td></tr> + +<tr><td> </td><td class="text_rt">12.</td><td class="justify">Geographic variation in the pocket gopher, Thomomys bottae, in Colorado. +By Phillip M. Youngman. Pp. 363-387, 7 figures in text. February 21, 1958.</td></tr> + +<tr><td> </td><td class="text_rt">13.</td><td class="justify">New bog lemming (genus Synaptomys) from Nebraska. By J. Knox Jones, +Jr. Pp. 385-388. May 12, 1958.</td></tr> + +<tr><td> </td><td class="text_rt">14.</td><td class="justify">Pleistocene bats from San Josecito Cave, Nuevo León, México. By J. Knox +Jones, Jr. Pp. 389-396. December 19, 1958.</td></tr> + +<tr><td> </td><td class="text_rt">15.</td><td class="justify">New subspecies of the rodent Baiomys from Central America. By Robert +L. Packard. Pp. 397-404. December 19, 1958.</td></tr> + +<tr><td> </td><td class="text_rt">16.</td><td class="justify">Mammals of the Grand Mesa, Colorado. By Sydney Anderson. Pp. 405-414, +1 figure in text, May 20, 1959.</td></tr> + +<tr><td> </td><td class="text_rt">17.</td><td class="justify">Distribution, variation, and relationships of the montane vole, Microtus montanus. +By Sydney Anderson. Pp. 415-511, 12 figures in text, 2 tables. +August 1, 1959.</td></tr> + +<tr><td> <span class='pagenum'><a name="Pub_2" id="Pub_2">[Pub_2]</a></span></td><td class="text_rt">18.</td><td class="justify">Conspecificity of two pocket mice, Perognathus goldmani and P. artus. By +E. Raymond Hall and Marilyn Bailey Ogilvie. Pp. 513-518, 1 map. January +14 1960.</td></tr> + +<tr><td> </td><td class="text_rt">19.</td><td class="justify">Records of harvest mice, Reithrodontomys, from Central America, with description +of a new subspecies from Nicaragua. By Sydney Anderson and +J. Knox Jones, Jr. Pp. 519-529. January 14, 1960.</td></tr> + +<tr><td> </td><td class="text_rt">20.</td><td class="justify">Small carnivores from San Josecito Cave (Pleistocene), Nuevo León, México. +By E. Raymond Hall. Pp. 531-538, 1 figure in text. January 14, 1960.</td></tr> + +<tr><td> </td><td class="text_rt">21.</td><td class="justify">Pleistocene pocket gophers from San Josecito Cave, Nuevo León, México. +By Robert J. Russell. Pp. 539-548, 1 figure in text. January 14, 1960.</td></tr> + +<tr><td> </td><td class="text_rt">22.</td><td class="justify">Review of the insectivores of Korea. By J. Knox Jones, Jr., and David H. +Johnson. Pp. 549-578. February 23, 1960.</td></tr> + +<tr><td> </td><td class="text_rt">23.</td><td class="justify">Speciation and evolution of the pygmy mice, genus Baimoys. By Robert L. +Packard. Pp. 579-670, 4 plates, 12 figures in text. June 16, 1960.</td></tr> + +<tr><td> </td><td colspan="2" class="justify">Index. Pp. 671-690</td></tr> + +<tr><td class="text_rt">Vol. 10.</td><td class="text_rt">1.</td><td class="justify">Studies of birds killed in nocturnal migration. By Harrison B. Tordoff and +Robert M. Mengel. Pp. 1-44, 6 figures in text, 2 tables. September 12, 1956.</td></tr> + +<tr><td> </td><td class="text_rt">2.</td><td class="justify">Comparative breeding behavior of Ammospiza caudacuta and A. maritima. +By Glen E. Woolfenden. Pp. 45-75, 6 plates, 1 figure. December 20, 1956.</td></tr> + +<tr><td> </td><td class="text_rt">3.</td><td class="justify">The forest habitat of the University of Kansas Natural History Reservation. +By Henry S. Fitch and Ronald R. McGregor. Pp. 77-127, 2 plates, 7 figures +in text, 4 tables. December 31, 1956.</td></tr> + +<tr><td> </td><td class="text_rt">4.</td><td class="justify">Aspects of reproduction and development in the prairie vole (Microtus ochrogaster). +By Henry S. Fitch. Pp. 129-161, 8 figures in text, 4 tables. December +19, 1957.</td></tr> + +<tr><td> </td><td class="text_rt">5.</td><td class="justify">Birds found on the Arctic slope of northern Alaska. By James W. Bee. +Pp. 163-211, plates 9-10, 1 figure in text. March 12, 1958.</td></tr> + +<tr><td> </td><td class="text_rt">*6.</td><td class="justify">The wood rats of Colorado: distribution and ecology. By Robert B. Finley, +Jr. Pp. 213-552, 34 plates, 8 figures in text, 35 tables. November 7, 1958.</td></tr> + +<tr><td> </td><td class="text_rt">7.</td><td class="justify">Home ranges and movements of the eastern cottontail in Kansas. By Donald +W. Janes. Pp. 553-572, 4 plates, 3 figures in text. May 4, 1959.</td></tr> + +<tr><td> </td><td class="text_rt">8.</td><td class="justify">Natural history of the salamander, Aneides hardyi. By Richard F. Johnston +and Gerhard A. Schad. Pp. 573-585. October 8, 1959.</td></tr> + +<tr><td> </td><td class="text_rt">9.</td><td class="justify">A new subspecies of lizard, Cnemidophorus sacki, from Michoacán, México. +By William E. Duellman. Pp. 587-598, 2 figures in text. May 2, 1960.</td></tr> + +<tr><td> </td><td class="text_rt">10.</td><td class="justify">A taxonomic study of the middle American snake, Pituophis deppei. By +William E. Duellman. Pp. 599-610, 1 plate, 1 figure in text. May 2, 1960.</td></tr> + +<tr><td> </td><td colspan="2" class="justify">Index. Pp. 611-626.</td></tr> + +<tr><td class="text_rt">Vol. 11.</td><td> </td><td colspan="2" class="justify">Nos. 1-10 and index. Pp. 1-703, 1958-1960.</td></tr> + +<tr><td class="text_rt">Vol. 12.</td><td class="text_rt">1.</td><td class="justify">Functional morphology of three bats: Sumops, Myotis, Macrotus. By Terry +A. Vaughan. Pp. 1-153, 4 plates, 24 figures in text. July 8, 1959.</td></tr> + +<tr><td> </td><td class="text_rt">*2.</td><td class="justify">The ancestry of modern Amphibia: a review of the evidence. By Theodore +H. Eaton, Jr. Pp. 155-180, 10 figures in text. July 10, 1959.</td></tr> + +<tr><td> </td><td class="text_rt">3.</td><td class="justify">The baculum in microtine rodents. By Sydney Anderson. Pp. 181-216, 49 +figures in text. February 19, 1960.</td></tr> + +<tr><td> </td><td class="text_rt">*4.</td><td>A new order of fishlike Amphibia from the Pennsylvanian of Kansas. By +Theodore H. Eaton, Jr., and Peggy Lou Stewart. Pp. 217-240, 12 figures in +text. May 2, 1960.</td></tr> + +<tr><td> </td><td class="text_rt">5.</td><td class="justify">Natural history of the bell vireo. By Jon C. Barlow. Pp. 241-296, 6 figures +in text. March 7, 1962.</td></tr> + +<tr><td> </td><td class="text_rt">6.</td><td class="justify">Two new pelycosaurs from the lower Permian of Oklahoma. By Richard C. +Fox. Pp. 297-307, 6 figures in text. May 21, 1962.</td></tr> + +<tr><td> </td><td class="text_rt">7.</td><td class="justify">Vertebrates from the barrier island of Tamaulipas, México. By Robert K. +Selander, Richard F. Johnston, B. J. Wilks, and Gerald G. Raun. Pp. 309-345, +pls. 5-8. June 18, 1962.</td></tr> + +<tr><td> </td><td class="text_rt">8.</td><td class="justify">Teeth of Edestid sharks. By Theodore H. Eaton, Jr. Pp. 347-362, 10 figures +in text. October 1, 1962.</td></tr> + +<tr><td> </td><td colspan="2" class="justify">More numbers will appear in volume 12.</td></tr> + +<tr><td class="text_rt">Vol. 13.</td><td class="text_rt">1.</td><td class="justify">Five natural hybrid combinations in minnows (Cyprinidae). By Frank B. +Cross and W. L. Minckley. Pp. 1-18. June 1, 1960.</td></tr> + +<tr><td> </td><td class="text_rt">2.</td><td class="justify">A distributional study of the amphibians of the Isthmus of Tehuantepec, +México. By William E. Duellman. Pp. 19-72, pls. 1-8, 3 figures in text. +August 16, 1960.</td></tr> + +<tr><td> </td><td class="text_rt">3.</td><td class="justify">A +new subspecies of the slider turtle (Pseudemys scripta) from Coahulia, +México. By John M. Legler. Pp. 73-84, pls. 9-12, 3 figures in text. August +16, 1960.</td></tr> + +<tr><td> </td><td class="text_rt">4.</td><td class="justify">Autecology of the copperhead. By Henry S. Fitch. Pp. 85-288, pls. 13-20, +26 figures in text. November 30, 1960.</td></tr> + +<tr><td> </td><td class="text_rt">5.</td><td class="justify">Occurrence of the garter snake, Thamnophis sirtalis, in the Great Plains and +Rocky Mountains. By Henry S. Fitch and T. Paul Maslin. Pp. 289-308, +4 figures in text. February 10, 1961.</td></tr> + +<tr><td> </td><td class="text_rt">6.</td><td class="justify">Fishes of the Wakarusa river in Kansas. By James E. Deacon and Artie L. +Metcalf. Pp. 309-322, 1 figure in text. February 10, 1961.</td></tr> + +<tr><td> </td><td class="text_rt">7.</td><td class="justify">Geographic variation in the North American cyprinid fish, Hybopsis gracilis. +By Leonard J. Olund and Frank B. Cross. Pp. 323-348, pls. 21-24, 2 figures +in text. February 10, 1961.</td></tr> + +<tr><td> <span class='pagenum'><a name="Pub_3" id="Pub_3">[Pub_3]</a></span></td><td class="text_rt">8.</td><td class="justify"><a name="descriptions"></a><a href="#typos">Descriptions</a> of two species of frogs, genus Ptychohyla; studies of American +hylid frogs, V. By William E. Duellman. Pp. 349-357, pl. 25, 2 figures +in text. April 27, 1961.</td></tr> + +<tr><td> </td><td class="text_rt">9.</td><td class="justify">Fish populations, following a drought, in the Neosho and Marais des Cygnes +rivers of Kansas. By James Everett Deacon. Pp. 359-427, pls. 26-30, 3 figs. +August 11, 1961.</td></tr> + +<tr><td> </td><td class="text_rt">10.</td><td>Recent soft-shelled +turtles of North America (family Trionychidae). By +Robert G. Webb. Pp. 429-611, pls. 31-54, 24 figures in text, February +16, 1962.</td></tr> + +<tr><td> </td><td colspan="2" class="justify">Index. Pp. 613-624.</td></tr> + +<tr><td class="text_rt">Vol. 14.</td><td class="text_rt">1.</td><td class="justify">Neotropical bats from western México. By Sydney Anderson. Pp. 1-8. +October 24, 1960.</td></tr> + +<tr><td> </td><td class="text_rt">2.</td><td class="justify">Geographic variation in the harvest mouse. Reithrodontomys megalotis, on +the central Great Plains and in adjacent regions. By J. Knox Jones, Jr., +and B. Mursaloglu. Pp. 9-27, 1 figure in text. July 24, 1961.</td></tr> + +<tr><td> </td><td class="text_rt">3.</td><td class="justify">Mammals of Mesa Verde National Park, Colorado. By Sydney Anderson. +Pp. 29-67, pls. 1 and 2, 3 figures in text. July 24, 1961.</td></tr> + +<tr><td> </td><td class="text_rt">4.</td><td class="justify">A new subspecies of the black myotis (bat) from eastern Mexico. By E. +Raymond Hall and Ticul Alvarez. Pp. 69-72, 1 figure in text. December +29, 1961.</td></tr> + +<tr><td> </td><td class="text_rt">5.</td><td class="justify">North American yellow bats, "Dasypterus," and a list of the named kinds +of the genus Lasiurus Gray. By E. Raymond Hall and J. Knox Jones, Jr. +Pp. 73-98, 4 figures in text. December 29, 1961.</td></tr> + +<tr><td> </td><td class="text_rt">6.</td><td class="justify">Natural history of the brush mouse (Peromyscus boylii) in Kansas with +description of a new subspecies. By Charles A. Long. Pp. 99-111, 1 figure +in text. December 29, 1961.</td></tr> + +<tr><td> </td><td class="text_rt">7.</td><td class="justify">Taxonomic status of some mice of the Peromyscus boylii group in eastern +Mexico, with description of a new subspecies. By Ticul Alvarez. Pp. 113-120, +1 figure in text. December 29, 1961.</td></tr> + +<tr><td> </td><td class="text_rt">8.</td><td class="justify">A new subspecies of ground squirrel (Spermophilus spilosoma) from Tamaulipas, +Mexico. By Ticul Alvarez. Pp. 121-124. March 7, 1962.</td></tr> + +<tr><td> </td><td class="text_rt">9.</td><td class="justify">Taxonomic status of the free-tailed bat, Tadarida yucatanica Miller. By J. +Knox Jones, Jr., and Ticul Alvarez. Pp. 125-133, 1 figure in text. March 7, +1962.</td></tr> + +<tr><td> </td><td class="text_rt">10.</td><td class="justify">A new doglike carnivore, genus Cynaretus, from the Clarendonian Pliocene, +of Texas. By E. Raymond Hall and Walter W. Dalquest. Pp. 135-138, +2 figures in text. April 30, 1962.</td></tr> + +<tr><td> </td><td class="text_rt">11.</td><td class="justify">A new subspecies of wood rat (Neotoma) from northeastern Mexico. By +Ticul Alvarez. Pp. 139-143. April 30, 1962.</td></tr> + +<tr><td> </td><td class="text_rt">12.</td><td class="justify">Noteworthy mammals from Sinaloa, Mexico. By J. Knox Jones, Jr., Ticul +Alvarez, and M. Raymond Lee. Pp. 145-159, 1 figure in text. May 18, +1962.</td></tr> + +<tr><td> </td><td class="text_rt">13.</td><td class="justify">A new bat (Myotis) from Mexico. By E. Raymond Hall. Pp. 161-164, +1 figure in text. May 21, 1962.</td></tr> + +<tr><td> </td><td class="text_rt">14.</td><td class="justify">The mammals of Veracruz. By E. Raymond Hall and Walter W. Dalquest. +Pp. 165-362, 2 figures. May 20, 1963.</td></tr> + +<tr><td> </td><td class="text_rt">15.</td><td class="justify">The recent mammals of Tamaulipas, México. By Ticul Alvarez. Pp. 363-473, +5 figures in text. May 20, 1963.</td></tr> + +<tr><td> </td><td colspan="2" class="justify">More numbers will appear in volume 14.</td></tr> + +<tr><td class="text_rt">Vol. 15.</td><td class="text_rt">1.</td><td class="justify">The amphibians and reptiles of Michoacán, México. By William E. Duellman. +Pp. 1-148, pls. 1-6, 11 figures in text. December 20, 1961.</td></tr> + +<tr><td> </td><td class="text_rt">2.</td><td class="justify">Some reptiles and amphibians from Korea. By Robert G. Webb, J. Knox +Jones, Jr., and George W. Byers. Pp. 149-173. January 31, 1962.</td></tr> + +<tr><td> </td><td class="text_rt">3.</td><td class="justify">A new species of frog (Genus Tomodactylus) from western México. By +Robert G. Webb. Pp. 175-181, 1 figure in text. March 7, 1962.</td></tr> + +<tr><td> </td><td class="text_rt">4.</td><td class="justify">Type specimens of amphibians and reptiles in the Museum of Natural History, +the University of Kansas. By William E. Duellman and Barbara Berg.<br /> +Pp. 183-204. October 26, 1962.</td></tr> + +<tr><td> </td><td class="text_rt">5.</td><td class="justify">Amphibians and Reptiles of the Rainforests of Southern El Petén, Guatemala. +By William E. Duellman. Pp. 205-249, pls. 7-10, 6 figures in text. October +4, 1963.</td></tr> + +<tr><td> </td><td class="text_rt">6.</td><td class="justify">A revision of snakes of the genus Conophis (Family Colubridae, from Middle +America). By John Wellman. Pp. 251-295, 9 figures in text. October 4, +1963.</td></tr> + +<tr><td> </td><td class="text_rt">7.</td><td class="justify">A review of the Middle American tree frogs of the genus Ptychohyla. By +William E. Duellman. Pp. 297-349, pls. 11-18, 7 figures in text. October +18, 1963.</td></tr> + +<tr><td> </td><td colspan="2" class="justify">More numbers will appear in volume 15.</td></tr> +</table> +<p> </p> +<p> </p> + +<div class="technt"> +<div class="caption3">Transcriber's Notes.</div> +<p>This file was derived from scanned images. With the exception of the +list of typographical errors that were corrected below, the original +text is presented.</p> + +<p>In the copy of the original, the Plates were grouped together between +pages 328 and 329. Here a smaller version of the illustration was placed +beside the text of the Systematic Account listing with a link to the +'full-sized' Plates which were moved to the end of the article's text +just above the listing of the Literature Cited. The Plate text contains +the notation "× 2" after the caption to let the reader know that the +image was enlarged by a factor of two. The images displayed will most +likely NOT be in 1:1 scale with the original printed Plates.</p> +<p> </p> + +<a name="typos"></a> +<span class="caption3">Typographical Errors Corrected:</span> +<p>Several minor typographical corrections were made (missing periods, +commas, misspelling of 'and', etc.); but are not indicated here. More +substantial changes are listed below:</p> + + References to <a href="#Plate_11">Plate 11</a> (Audiograms): Pl. 1A, Pl. 1B, ...=> Pl. 11A, Pl. 11B, ...<br /> + References to the other Plates: Pl. 2, Pl. 3,... => Pl. 12, Pl. 13,...<br /> + Page 301, Paragraph 1: <a href="#known">know => known</a><br /> + Page 302, Paragraph 1: <a href="#Zoology">Zoolegy => Zoology</a><br /> + Page 303, Paragraph 5: <a href="#species">speces => species</a><br /> + Page 305, Paragraph 1: <a href="#excrescences">excresences => excrescences</a><br /> + Page 308, Paragraph 6: <a href="#xiphisternum">xiphisterum => xiphisternum</a><br /> + Page 316, Paragraph 3: <a href="#width">with => width</a><br /> + Page 327, Paragraph 1: <a href="#leonhardschultzei">leonhard-schultzei => leonhardschultzei</a><br /> to match remaining report text<br /> + Page 331, Paragraph 1: <a href="#skeleton">skelton => skeleton</a><br /> + Publication List Vol. 13, No. 8.: <a href="#descriptions">Decriptions => Descriptions</a><br /> +</div> +<p> </p> + + + + + + + + + + +<pre> + + + + + +End of the Project Gutenberg EBook of A Review of the Middle American Tree +Frogs of the Genus Ptychohyla, by William E. 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