diff options
Diffstat (limited to '35413.txt')
| -rw-r--r-- | 35413.txt | 3617 |
1 files changed, 3617 insertions, 0 deletions
diff --git a/35413.txt b/35413.txt new file mode 100644 index 0000000..9caaea9 --- /dev/null +++ b/35413.txt @@ -0,0 +1,3617 @@ +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] + +Language: English + +Character set encoding: ASCII + +*** 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 + + + + + + + + + + UNIVERSITY OF KANSAS PUBLICATIONS + MUSEUM OF NATURAL HISTORY + + Volume 15, No. 7, pp. 297-349, pls. 11-18, 7 figs. + + October 18, 1963 + + + + + A Review of the Middle American Tree Frogs + of the Genus Ptychohyla + + BY + + WILLIAM E. DUELLMAN + + + + + UNIVERSITY OF KANSAS + LAWRENCE + 1963 + + + + + UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY + + Editors: E. Raymond Hall, Chairman, Henry S. Fitch, + Theodore H. Eaton, Jr. + + Volume 15, No. 7, pp. 297-349, pls. 11-18, 7 figs. + + Published October 18, 1963 + + + UNIVERSITY OF KANSAS + Lawrence, Kansas + + + PRINTED BY + JEAN M. NEIBARGER, STATE PRINTER + TOPEKA, KANSAS + 1963 + + [Union Label] + 29-6531 + + + + + A Review of the Middle American Tree Frogs + of the Genus Ptychohyla + + BY + WILLIAM E. DUELLMAN + + + + CONTENTS + + PAGE + + INTRODUCTION 301 + Acknowledgments 301 + Materials and Methods 302 + + ANALYSIS OF DATA 303 + External Morphology 303 + Color and Pattern 307 + Osteology 307 + Tadpoles 310 + Breeding Call 312 + + SYSTEMATIC ACCOUNTS 314 + _Ptychohyla_ Taylor, 1944 314 + Key to Adults 315 + Key to Tadpoles 315 + _Ptychohyla euthysanota_ Group 315 + _Ptychohyla euthysanota_ 315 + _Ptychohyla euthysanota euthysanota_ (Kellogg) 315 + _Ptychohyla euthysanota macrotympanum_ (Tanner) 320 + _Ptychohyla leonhardschultzei_ (Ahl) 323 + _Ptychohyla spinipollex_ (Schmidt) 327 + _Ptychohyla schmidtorum_ Group 331 + _Ptychohyla schmidtorum_ 331 + _Ptychohyla schmidtorum schmidtorum_ Stuart 331 + _Ptychohyla schmidtorum chamulae_ Duellman 334 + _Ptychohyla ignicolor_ Duellman 337 + + DISTRIBUTION AND ECOLOGY 340 + Geographic Distribution of the Species 340 + Habitat Preference 342 + Interspecific Competition 343 + Reproduction and Development 344 + + PHYLOGENY OF PTYCHOHYLA 345 + _Ptychohyla_ as a Natural Assemblage 345 + Generic Relationships 346 + Interspecific Relationships 347 + + LITERATURE CITED 349 + + + + + INTRODUCTION + + +Probably no ecological group of hylid frogs (some _Hyla_ plus +_Plectrohyla_ and _Ptychohyla_) 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 Mexico 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 _Mission +Scientifique_, no members of the genus _Ptychohyla_ 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 +_Hyla euthysanota_ by Kellogg (1928). Until recently frogs of this +genus were known from few specimens and in the literature by nearly as +many names. + +Although I first collected _Ptychohyla_ 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 Mexico and Guatemala, where every +accessible stream in the cloud forests was searched for tree frogs, +especially _Ptychohyla_ and _Plectrohyla_. Similar, but less +extensive, investigations were carried out in 1961 and 1962. The +result of this field work is a rather large collection of _Ptychohyla_ +representing all of the known species, plus tape recordings of the +breeding calls and tadpoles of all of the species. + +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 "_Ptychohyla_ as a +Natural Assemblage." + + +Acknowledgments + +I am indebted to the following persons for permitting me to examine +specimens in their care: Miguel Alvarez del Toro, Museo Zoologia de +Tuxtla Gutierrez, Mexico (MZTG); Charles M. Bogert and Richard G. +Zweifel, American Museum of Natural History (AMNH); Doris M. Cochran, +United States 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 Zoology (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. + +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. + +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 +_Ptychohyla_. I am indebted to many residents of Mexico, 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 Solistahuacan, +Chiapas, for a pleasant stay at their school; Jordi Julia Z. of the +Comision del Papaloapan, Ciudad Aleman, 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 Cientificas, 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 +Mexico were furnished by the late Luis Macias Arellano of the +Direccion General de Caza. Each of these individuals has my profound +thanks for his indispensable aid. + +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. + + +Materials and Methods + +During the course of this study I have examined 247 frogs that I +assign to the genus _Ptychohyla_, 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 (_P. euthysanota macrotympanum_) to +33 (_P. spinipollex_) living individuals of each species. + +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. + +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. + + + + +ANALYSIS OF DATA + + +Data that are used to arrive at a systematic arrangement of the +species of _Ptychohyla_ 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. + + +External Morphology + +Each of the external morphological characters used in the systematic +treatment of _Ptychohyla_, as well as the nature of the tongue, +is discussed below. + +SIZE AND PROPORTIONS.--Comparisons of size and certain proportions are +given in Table 1. Frogs of this genus are small; the largest specimen +examined is a female of _P. euthysanota euthysanota_ having a +snout-vent length of 53.3 mm. The species comprising the _Ptychohyla +schmidtorum_ group are smaller; the largest specimen examined is a +female of _P. schmidtorum schmidtorum_ having a snout-vent length of +38.0 mm. An analysis of the various measurements and proportions shows +few constant differences. _Ptychohyla ignicolor_ 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. _Ptychohyla +spinipollex_ 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. + +SNOUT.--All species have a blunt snout. In _P. leonhardschultzei_ and +_P. ignicolor_ the snout is nearly square in lateral profile; in _P. +schmidtorum_ the snout is slightly rounded above and below, and in the +other species it is rounded above. _Ptychohyla leonhardschultzei_ and +_P. spinipollex_ have a vertical fleshy rostral keel on the snout; in +these species, because of this keel, the snout in dorsal profile is +pointed. The nostrils are slightly protuberant in all species, and in +_P. schmidtorum_ the internarial region is slightly depressed. + +HAND.--The species in the _Ptychohyla euthysanota_ 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 _P. spinipollex_ +(Fig. 1) and vary in number from 35 to 66 (average 47.4) on each thumb. +In the other species of the _Ptychohyla euthysanota_ 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: _P. +euthysanota euthysanota_, 44-143 (83.8); _P. euthysanota +macrotympanum_, 40-110 (63.0); _P. leonhardschultzei_, 24-80 (54.7). +The species in the _Ptychohyla schmidtorum_ 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 excrescences on the thumbs of +many species of frogs, the horny spines on the thumbs of members of +the _Ptychohyla euthysanota_ group are seasonal in development. + + +TABLE 1.--VARIATION IN CERTAIN CHARACTERS IN THE SPECIES OF PTYCHOHYLA. + (MEANS ARE IN PARENTHESES BELOW THE RANGES.) + + Key to Table Columns + + A) Number of specimens + B) Maximum snout-vent length + C) Tibia length/Snout-vent length + D) Tympanum/Eye + E) Vomerine teeth + +==================+========+======+=======+=========+=========+======= + Species | Sex | A | B | C | D | E +------------------+--------+------+-----------------+----------------- + | | | | | | +_P. euthysanota | [Male] | 17 | 38.1 |44.4-55.0|48.6-63.8| 4-6 + euthysanota_ | | | | (48.7) | (56.3) | (5.1) + | | | | | | + |[Female]| 15 | 53.3 |46.5-56.6|42.9-56.4| 6-18 + | | | | (51.4) | (51.4) | (9.6) + | | | | | | +_P. euthysanota | [Male] | 5 | 38.0 |48.8-52.0|50.0-57.1| 0-4 + macrotympanum_| | | | (50.2) | (54.1) | (2.6) + | | | | | | + |[Female]| 5 | 44.8 |46.4-54.1|48.7-58.9| 8-10 + | | | | (50.2) | (53.7) | (9.2) + | | | | | | +_P. leonhard | [Male] | 16 | 35.6 |48.8-56.1|48.7-61.9| 6-9 + schultzei_ | | | | (51.2) | (52.1) | (6.5) + | | | | | | + |[Female]| 3 | 41.6 |52.3-59.5|47.5-48.6| 7-12 + | | | | (54.7) | (48.1) | (9.5) + | | | | | | +_P. spinipollex_ | [Male] | 32 | 41.2 |46.9-53.1|45.0-55.2| 3-7 + | | | | (49.0) | (49.5) | (4.9) + | | | | | | + |[Female]| 6 | 44.6 |46.1-50.2|47.7-53.8| 6-10 + | | | | (48.8) | (50.4) | (7.6) + | | | | | | +_P. schmidtorum | [Male] | 25 | 32.8 |45.3-52.4|51.5-59.3| 5-11 + schmidtorum_ | | | | (48.1) | (54.7) | (6.2) + | | | | | | + |[Female]| 9 | 38.0 |46.5-49.1|51.3-58.3| 7-11 + | | | | (47.7) | (54.9) | (8.7) + | | | | | | +_P. schmidtorum | [Male] | 40 | 30.5 |46.0-51.9|48.2-65.6| 4-6 + chamulae_ | | | | (48.2) | (54.9) | (4.7) + | | | | | | + |[Female]| 4 | 31.8 |48.1-52.4|51.4-61.7| 4-9 + | | | | (50.5) | (55.7) | (6.2) + | | | | | | +_P. ignicolor_ | [Male] | 13 | 30.5 |45.9-52.2|37.1-47.1| 3-9 + | | | | (49.6) | (43.2) | (6.1) +------------------+--------+------+-------+---------+---------+------- + + + + [Illustration: FIG. 1. Palmar views of right hands of + (A) _Ptychohyla spinipollex_ (KU 58054) and + (B) _Ptychohyla schmidtorum schmidtorum_ (KU 58043).] + + +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 _Ptychohyla_ +reveals considerable intraspecific variation. Bifid tubercles beneath +the fourth finger are found in all species except _P. ignicolor_, +which is known from only two specimens. In _P. euthysanota +euthysanota_ nearly 60 per cent and in _P. schmidtorum schmidtorum_ +about 90 per cent of the specimens have a bifid tubercle beneath the +fourth finger on one or both hands. All specimens of _P. +leonhardschultzei_ have either a bifid or double tubercle beneath the +fourth finger, and some have a bifid distal tubercle beneath the third +finger. + +FEET.--Members of the _Ptychohyla euthysanota_ group have a weak +tarsal fold, whereas in the species comprising the _Ptychohyla +schmidtorum_ 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 _P. ignicolor_, +which has less webbing. + +VENTROLATERAL GLANDS.--Breeding males develop thickened, pigmented +glandular areas on the sides of the body. In living specimens of _P. +schmidtorum_ and _P. ignicolor_ the glands are not readily visible, +but in preservative they show as distinctive orange-colored areas. +These glands are most distinct in _P. euthysanota_; in many specimens +of this species the glands are elevated above the surrounding skin. +The extent of the glands is variable (Fig. 2); probably this +variability is due to different degrees of development in individual +frogs rather than to interspecific differences. All _Ptychohyla +ignicolor_ and some _P. schmidtorum chamulae_ 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. + + + [Illustration: FIG. 2. Normal extent of ventrolateral glands in + (A) _Ptychohyla euthysanota euthysanota_ (KU 58008), + (B) _Ptychohyla schmidtorum schmidtorum_ (KU 58037), and + (C) _Ptychohyla ignicolor_ (UMMZ 119603).] + + +TONGUE.--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 _P. leonhardschultzei_ +and _P. schmidtorum_; 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. + + + +Color and Pattern + +The dorsum in living frogs of the genus _Ptychohyla_ is primarily +yellowish or reddish brown, except in _P. schmidtorum chamulae_ and +_P. ignicolor_ in which it is green. Usually there are some darker +blotches or reticulations on the dorsum. The venter usually is white; +in _P. ignicolor_ it is yellow. The venter is spotted in all members +of the _Ptychohyla euthysanota_ group; the species, arranged from +least to most spotting ventrally, are: _P. euthysanota euthysanota_, +_P. euthysanota macrotympanum_, _P. leonhardschultzei_, and _P. +spinipollex_. The last two species also have bold dark spots on the +flanks. _Ptychohyla schmidtorum_ lacks spots on the venter, whereas +_P. ignicolor_ has small dark flecks ventrally. + +_Ptychohyla euthysanota_ and _P. schmidtorum_ 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 _P. ignicolor_, +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 _P. schmidtorum schmidtorum_. + +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. + + + +Osteology + +The following description of the skull, hyoid, sternum, and prepollex +is based on a male specimen of _P. spinipollex_ (KU 68632) that has +been cleared and stained. The broad, flat skull (Fig. 3) 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 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. + + + [Illustration: FIG. 3. Dorsal aspect of skull of _Ptychohyla + spinipollex_ (KU 68632). Arrow indicates reduced + quadratojugal.] + + +The posteromedian part of the hyoid plate is calcified; from this +plate the long bony, posterior cornua (thyrohyales) extend +posterolaterally. + +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. + +The swollen thumb is supported by a dorsoventrally flattened spine +that does not extrude through the skin. + +VARIATION.--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 _P. spinipollex_ the quadratojugal +articulates with the maxillary and forms a complete +quadratojugal-maxillary arch on each side of the skull. One specimen +of _P. leonhardschultzei_ has a complete arch on one side and an +incomplete arch on the other. + +Only _P. spinipollex_ has lateral processes anteriorly on the +xiphisternum; in the other species the xiphisternum is deeply +bell-shaped. + +_Ptychohyla schmidtorum_ and _P. ignicolor_ have slightly longer +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 _P. schmidtorum_ and 10 teeth in one _P. +ignicolor_, 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: _P. euthysanota euthysanota_, 43; _P. euthysanota +macrotympanum_, 38; _P. leonhardschultzei_, 38 and 40; _P. +spinipollex_, 34 and 40; _P. schmidtorum schmidtorum_, 37 and 43; _P. +schmidtorum chamulae_, 40 and 41; _P. ignicolor_, 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. + +Variation in number of vomerine teeth is shown in Table 1. + + + [Illustration: FIG. 4. Tadpoles of the _Ptychohyla euthysanota_ group: + (A) _P. euthysanota euthysanota_ (KU 60042), + (B) _P. euthysanota macrotympanum_ (KU 60049), + (C) _P. leonhardschultzei_ (KU 68556), and + (D) _P. spinipollex_ (KU 60053).] + + +Tadpoles + +Tadpoles of the genus _Ptychohyla_ are adapted to live in mountain +streams. The bodies are streamlined, and the tails are long and have +low fins (Figs. 4 and 5). The mouths are large and directed ventrally. +Tadpoles of the two groups of species have strikingly different +mouthparts. + + + [Illustration: FIG. 5. Tadpoles of + (A) _Ptychohyla schmidtorum schmidtorum_ (KU 60051), + (B) _P. schmidtorum chamulae_ (KU 58199), and + (C) _P. ignicolor_ (KU 71716).] + + +Lips of tadpoles of the _Ptychohyla euthysanota_ group (Fig. 6 A-D) +are folded laterally; there are 4/6 or sometimes 4/7 tooth-rows. A +lateral "wing" projects on either side of the upper beak. The beaks +have blunt, peglike serrations. Lips of tadpoles of the _Ptychohyla +schmidtorum_ group (Fig. 6 E-G) are greatly expanded and form a +funnel-shaped disc; there are 3/3 short tooth-rows. There is no +lateral projection or "wing" on either side of the upper beak. The +beaks have long, pointed serrations. + + [Illustration: FIG. 6. Mouthparts of tadpoles of _Ptychohyla_: + (A) _P. euthysanota euthysanota_ (KU 60042), + (B) _P. euthysanota macrotympanum_ (KU 60049), + (C) _P. leonhardschultzei_ (KU 68556), + (D) _P. spinipollex_ (KU 60053), + (E) _P. schmidtorum schmidtorum_ (KU60051), + (F) _P. schmidtorum chamulae_ (KU 58199), and + (G) _P. ignicolor_ (KU 71716). x 10.] + +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 +Table 2. + + +TABLE 2.--COMPARISON OF CERTAIN LARVAL CHARACTERS IN THE SPECIES OF +PTYCHOHYLA. (MEANS ARE IN PARENTHESES BELOW THE RANGES.) + +==============================+=========+=======+============+========== + | Number |Maximum| Head length| + Species | of | total |------------|Tooth-rows + |specimens| length|Total length| +------------------------------+---------+-------+------------+---------- +_P. euthysanota euthysanota_ | 23 | 40.8 | 30.9-37.3 | 4/6 + | | | (33.5) | +_P. euthysanota macrotympanum_| 13 | 43.3 | 30.6-33.4 | 4/6 + | | | (32.7) | +_P. leonhardschultzei_ | 7 | 47.5 | 29.2-32.7 | 4/6 + | | | (31.1) | +_P. spinipollex_ | 32 | 45.0 | 30.2-35.9 | 4/7 + | | | (33.0) | +_P. schmidtorum schmidtorum_ | 14 | 42.5 | 28.9-31.2 | 3/3 + | | | (29.9) | +_P. schmidtorum chamulae_ | 4 | 45.0 | 26.9-29.3 | 3/3 + | | | (27.8) | +_P. ignicolor_ | 2 | 39.6 | 29.6-29.8 | 3/3 + | | | (29.7) | +------------------------------+---------+-------+------------+---------- + + +Evidence on the pattern of development of tooth-rows indicates that +the inner rows develop first. A small tadpole of _P. euthysanota +euthysanota_ has six lower rows and three fully developed upper rows +and only the beginning of the first (outer) upper row. A small tadpole +of _P. euthysanota macrotympanum_ has four upper rows and five lower +rows. In a small tadpole of _P. leonhardschultzei_ 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 _P. spinipollex_, the sixth lower row is +poorly developed, and the seventh row is absent; large individuals +normally have seven lower rows. A small tadpole of _P. schmidtorum +chamulae_ has 3/2 tooth-rows. + + + +Breeding Call + +Breeding calls of all species and subspecies of _Ptychohyla_ were +recorded in the field. Obtaining series of calls of _Ptychohyla_ 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 _P. +spinipollex_ were recorded, but only one recording was sufficiently +free of background noise to be analyzed. + +Analysis of breeding calls supports the division of the genus +_Ptychohyla_ into two groups of species. The call of each member of +the _Ptychohyla euthysanota_ group consists of a single long note, +whereas the call of species in the _Ptychohyla schmidtorum_ 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 Plate 11, and comparisons of the characteristics of the calls +are given in Table 3. + + + [Illustration: PLATE 11 + + Audiospectrographs of the breeding calls of the + five species of _Ptychohyla_: + + (A) _P. spinipollex_ (KU Tape No. 41), + (B) _P. euthysanota macrotympanum_ (KU Tape No. 48), + (C) _P. leonhardschultzei_ (UMMZ Tape No. 525), + (D) _P. schmidtorum chamulae_ (KU Tape No. 52), + (E) _P. ignicolor_ (UMMZ Tape No. 526).] + + + TABLE 3.--COMPARISON OF THE BREEDING CALLS OF PTYCHOHYLA + + -------------------------------------------------------------------------- + ----------------+----+--------+-----------+---------+---------+----------- + | | Notes | Duration | Pulses |Frequency| Dominant + Species |Num-|per call| of note | per | range | frequency + | ber| group | (seconds) | second | (cps) | (cps) + ----------------+----+--------+-----------+---------+---------+----------- + _P. spinipollex_| 1 | 1 | .46 | 147 |3000-5100| 4300 + | | | | | | + _P. euthysanota_| 7 | 1 | .62 | 95.3 |1800-4200| 3070 + | | | (.60-.65) | (91-102)| |(3000-3200) + | | | | | | + _P. leonhard- | 2 | 1 | .79 | 77 |1500-3500| 2750 + schultzei_| | | (.62-.95) | (76-78) | |(2700-2800) + | | | | | | + _P. schmidtorum_| 6 | 8.5[A] | .064 | 110 |1400-5800| 3400 + | | (8-9) |(.054-.070)| (96-121)| |(3350-3450) + | | | | | | + _P. ignicolor_ | 2 | 12[A] | .079 | 126 |1000-5000| 3150 + | |(11-13) |(.078-.080)|(123-129)| |(3100-3200) + ----------------+----+--------+-----------+---------+---------+----------- + + [Footnote A: Only an analysis of the long series of calls is given + here; see text for explanation.] + + +_P. spinipollex_ (Pl. 11A).--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. euthysanota_ (Pl. 11B).--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. leonhardschultzei_ (Pl. 11C).--One long note is repeated once +after 10 to 13 seconds and has an average dominant frequency of 2750 +cycles per second. + +_P. schmidtorum_ (Pl. 11D).--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. ignicolor_ (Pl. 11E).--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 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. + + + + + SYSTEMATIC ACCOUNTS + + +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 (Fig. 7) are in +roman type; those that are not represented on the map, because +overlapping of symbols would have occurred, are in italic type. + + + =Ptychohyla= Taylor, 1944 + + + _Ptychohyla_ Taylor, Univ. Kansas Sci. Bull., 30:41, May 15, 1944. + Type, _Ptychohyla adipoventris_ Taylor, 1944 [= _Hyla + leonhardschultzei_ (Ahl), 1934]. + + +_Diagnosis._--Small hylids having stream-adapted tadpoles and +differing from other hylid genera in having large ventrolateral glands +in breeding males. + +_Composition._--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. + +_Distribution._--Moderate elevations from southern Guerrero and +northern Oaxaca, Mexico, to northern El Salvador and central Honduras. + + + + + KEY TO ADULTS + + +1. 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 + _P. euthysanota_ group--2 + + 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 _P. schmidtorum_ group--5 + +2. 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 3 + + 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 4 + +3. 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 _P. leonhardschultzei_ + + 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 + _P. spinipollex_ + +4. A distinct, broad, white lateral stripe usually present; + usually a distinct white line above anal opening; a distinct + white stripe on upper lip _P. euthysanota euthysanota_ + + No white lateral stripe; a faint white stripe above anal + opening; no distinct white stripe on upper lip + _P. euthysanota macrotympanum_ + +5. 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 feet not red in life; internarial region + slightly depressed; diameter of tympanum greater than one-half + diameter of eye 6 + + 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 _P. ignicolor_ + +6. Webs of feet and posterior surfaces of thighs pale cream + color; dorsum in life reddish brown; iris bright red + _P. schmidtorum schmidtorum_ + + Webs of feet and posterior surfaces of thighs pale brown; + dorsum in life green; iris reddish bronze color + _P. schmidtorum chamulae_ + + + KEY TO TADPOLES + +1. Lips greatly expanded forming a funnel-shaped mouth; + tooth-rows 3/3 _P. schmidtorum_ group--2 + + Lips folded laterally, not forming a funnel-shaped mouth; + tooth-rows 4/6 or more _P. euthysanota_ group--4 + +2. Belly and mouth mottled; tail cream color heavily blotched + with brown 3 + + Belly dark gray; tail cream color with dense brown flecking, + giving brown appearance _P. schmidtorum chamulae_ + +3. Belly cream color with brown mottling; no large tubercle at + each end of first lower tooth-row _P. schmidtorum schmidtorum_ + + Belly grayish green with brown mottling; a large tubercle at + each end of first lower tooth-row _P. ignicolor_ + +4. Tooth-rows 4/6; cream-colored crescent-shaped mark on + posterior part of body bordered posteriorly by large brown + mark 5 + + Tooth-rows usually 4/7 (sometimes 4/6); cream-colored + crescent-shaped mark on posterior part of body usually + indistinct, not bordered posteriorly by large brown mark + _P. spinipollex_ + +5. Caudal musculature uniformly flecked with brown; lower + tooth-rows 1-4 about equal in length to upper rows + _P. euthysanota euthysanota_ + + Caudal musculature having brown square blotches dorsally on + anterior one-half of tail; lower tooth-rows 1-4 usually + slightly shorter than upper rows 6 + +6. Dorsal caudal blotches well defined and extending onto sides + of tail; moderately large brown flecks on caudal fin; eye in + life pale reddish brown _P. leonhardschultzei_ + + Dorsal caudal blotches faint, not extending onto sides of + tail; small brown flecks on caudal fin; eye in life silvery + bronze _P. euthysanota macrotympanum_ + + + + + The _Ptychohyla euthysanota_ Group + + +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 4/6 or 4/7 tooth-rows; breeding call +consisting of one long note. + + + + + =Ptychohyla euthysanota= + +_Diagnosis._--Rostral keel absent; nuptial spines in males small; +interorbital region much wider than eyelid. + + + + + [Illustration: PLATE 12 + _Ptychohyla euthysanota euthysanota_ (KU 58008). x 2.] + + + =Ptychohyla euthysanota euthysanota= (Kellogg) + + _Hyla euthysanota_ 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, 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. + + _Hyla rozellae_ Taylor, Univ. Kansas Sci. Bull., 28:78-80, pl. 9, + fig. 1, May 15, 1942 [Holotype.--USNM 115039 from Salto de + Agua, Chiapas, Mexico; 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. + + _Ptychohyla bogerti_ Taylor, Amer. Mus. Novitates, 1437:13-16, + fig. 5, December 7, 1949 [Holotype.--AMNH 51847 from Rio + Grande, Oaxaca, Mexico; Thomas MacDougall collector]. Stuart, + Proc. Biol. Soc. Washington, 67:169, August 5, 1954. + + _Ptychohyla euthysanota_, Duellman, Univ. Kansas Publ. Mus. Nat. + Hist., 13:351, April 27, 1961. + + +_Diagnosis._--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. + +_Description._--The following description is based on KU 58008 from +Finca La Paz, Depto. San Marcos, Guatemala (Pl. 12). 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 width/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 to vocal sac large, one +situated along inner posterior edge of each mandibular ramus. + +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. + +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). + +_Variation._--No geographic variation in structural characters is +discernible; variation in size and proportions is given in Table 1. 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. + +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). + +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 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. + +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. + +_Description of Tadpole._--The following description is based on KU +60042 from Finca La Paz, Depto. San Marcos, Guatemala (Figs. 4A and +6A). 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. + +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 4/6; 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. + +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. + +_Variation._--The variation in size and proportions is given in Table +2. 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. + +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. + +_Comparisons._--Aside from the characters given in the diagnosis, _P. +euthysanota euthysanota_ can be distinguished from both _P. +spinipollex_ and _P. leonhardschultzei_ 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. _Ptychohyla euthysanota euthysanota_ differs +from _P. euthysanota macrotympanum_ by normally having a darker dorsal +color, broader stripe on upper lip, and a distinct lateral stripe. + +Occurring sympatrically with _Ptychohyla euthysanota euthysanota_ are +several species of _Plectrohyla_, all of which differ in having a bony +prepollex, rather rugose skin on the dorsum, and more squat bodies. +Other sympatric species are _Ptychohyla schmidtorum schmidtorum_, +which lacks a tarsal fold and nuptial spines and has a red eye in +life, _Hyla salvadorensis_, which has a green dorsum and lacks spinous +nuptial tuberosities, and _Hyla sumichrasti_, a small yellow frog +usually lacking vomerine teeth. + + +_Life History._--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. + +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. + +_Remarks._--The type specimen of _Hyla euthysanota_ Kellogg (1928:123) +is a female; therefore, when Taylor (1944) proposed the name +_Ptychohyla_ for hylids having ventrolateral glands in breeding males, +he was unaware that _Hyla euthysanota_ was a member of this group. In +his description of _Hyla rozellae_, Taylor (1942) did not compare his +specimens with _Hyla euthysanota_, but instead placed _H. rozellae_ +with _H. loquax_ and _H. rickardsi_. The type series of _H. rozellae_ +consists of one large adult female and several metamorphosing young. +Taylor (1949:16) based the description of _Ptychohyla bogerti_ on two +males and compared these specimens with _P. adipoventris_ Taylor [= +_P. leonhardschultzei_ (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 _Hyla euthysanota_ and _Hyla +rozellae_ were _Ptychohyla_. Now that sufficient specimens are +available from throughout the range it is possible to determine that +the various named populations are conspecific. + +_Distribution._--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, Mexico, 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 Jose; 43040, Hacienda Los Planes; 65119, Miramundo) are listed by +Mertens (1952:29). + +_Specimens examined._--MEXICO: _Chiapas: Cascarada, 30 km. W of +Ciltapec_, UMMZ 87851-2; Cerro Ovando, UMMZ 87853-4; Chicomuselo, UMMZ +94439-40; Finca Juarez, 28 km. N of Escuintla, USNM 115052-5; _Las +Nubes, Cerro Ovando_, USNM 115030-8; Salto de Agua, USNM 115039-51. +_Oaxaca_: Cerro Pecho Blanco, UIMNH 40963; between La Gloria and Cerro +Azul, UIMNH 40976-7; Rio Grande, AMNH 51847-8; Santo Tomas Tecpan, +UIMNH 41071. + +GUATEMALA: _San Marcos_: 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, Rio Samala, +CNHM 35066. _Santa Rosa_: Finca La Gloria, UMMZ 123148 (tadpoles), +123150 (tadpoles). _Solola_: Finca Santo Tomas, UMMZ 123149 +(tadpoles); Olas de Moca, near Moca, CNHM 20208. + +EL SALVADOR: _Chalatenango_: Los Esemiles, USNM 73296. _Santa Ana_: +Miramundo, CNHM 65120. + + + + + [Illustration: PLATE 13 + _Ptychohyla euthysanota macrotympanum_ (KU 58049). x 2.] + + + =Ptychohyla euthysanota macrotympanum= (Tanner) + + + _Hyla macrotympanum_ 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, Mexico; Robert Bohlman + collector]. + + _Ptychohyla macrotympanum_, Duellman, Univ. Kansas Publ. Mus. Nat. + Hist., 13:351, April 27, 1961. + + +_Diagnosis._--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. + +_Description._--The following description is based on KU 58049 from +Linda Vista, Chiapas, Mexico (Pl. 13). 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 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. + +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. + +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). + +_Variation._--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. + +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. + +_Description of Tadpole._--The following description is based on KU +60049 from Rio Hondo, 9.5 kilometers south of Pueblo Nuevo +Solistahuacan, Chiapas, Mexico (Figs. 4B and 6B). 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 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. + +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 4/6; 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. + +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. + +_Variation._--The variation in size and proportions is given in Table +2. All specimens have 4/6 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. + +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. + +_Comparisons._--_Ptychohyla euthysanota macrotympanum_ can be +distinguished from both _P. spinipollex_ and _P. leonhardschultzei_ 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. _Ptychohyla euthysanota macrotympanum_ differs +from _P. e. euthysanota_ by normally having a paler dorsum, narrower +stripe on upper lip, and no distinct lateral stripe. + +_Ptychohyla euthysanota macrotympanum_ occurs sympatrically with +_Plectrohyla guatemalensis_ and _P. matudai matudai_. Each of the last +two has a bony prepollex, rather rugose skin on the dorsum, and more +squat body. Other sympatric species are _Hyla walkeri_, which has a +green dorsum with brown markings and a rather pointed snout, and _Hyla +sumichrasti_, a small yellow frog usually lacking vomerine teeth. + +_Life History._--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 (Pl. 11B). The call is +indistinguishable from that of _P. e. euthysanota_. + +Tadpoles in various stages of development were found in the Rio 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. + +_Remarks._--Tanner (1957:52) based the description of _Hyla +macrotympanum_ on a single female, which, of course, lacked the +characters diagnostic of _Ptychohyla_. On the basis of general +external characters Tanner suggested that _Hyla macrotympanum_ was +related to _H. miotympanum_, 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 _Hyla macrotympanum_ is a _Ptychohyla_. + +Intergradation between _Ptychohyla euthysanota_ and _P. macrotympanum_ +has not been demonstrated. Currently their ranges are separated by the +dry valleys of the Rio Grijalva and Rio Cuilco. The similarity in +structure of the adults and tadpoles and the indistinguishable +breeding calls are the basis for considering the two populations +subspecies. + +_Distribution._--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 Rio +Grijalva and Rio Cuilco. + +_Specimens examined._--MEXICO: _Chiapas_: 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 Solistahuacan, KU 58049-51, 59939 (skeleton); +_Rio Hondo, 9.5 km. S of Pueblo Nuevo Solistahuacan_, KU 58047-8, +60046-7, 60048-9 (tadpoles); San Fernando, MZTG 15, 17; Tonina +(ruins), KU 41592. + +GUATEMALA: _Huehuetenango_: 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). + + + + + [Illustration: PLATE 14 + _Ptychohyla leonhardschultzei_ (KU 64117). x 2.] + + + =Ptychohyla leonhardschultzei= (Ahl) + + + _Hyla leonhard-schultzei_ Ahl, Zool. Anz., 106:185-186, fig. 1, + April 15, 1934 [Holotype.--ZMB 34353 from Malinaltepec, + Guerrero, Mexico; Leonhard Schultze collector]. Smith and + Taylor, Bull. U. S. Natl. Mus., 184:87, June 17, 1948. + + _Hyla godmani_, Ahl, Zool. Anz., 106:186, April 15, 1934. + + _Hyla pinorum_ Taylor, Proc. Biol. Soc. Washington, 50:46-48, + pl. 2, fig. 2, April 21, 1937 [Holotype.--UIMNH 25049 from + Agua del Obispo, Guerrero, Mexico; Edward H. Taylor + collector]. Smith and Taylor, Bull. U. S. Natl. Mus., 194:87, + June 17, 1948. + + _Ptychohyla adipoventris_ Taylor, Univ. Kansas Sci. Bull., + 30:41-45, May 15, 1944 [Holotype.--UIMNH 25047 from Agua del + Obispo, Guerrero, Mexico; 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. + + _Hyla milleri_ Shannon, Proc. U. S. Natl. Mus., 101:473-477, + figs. 92b, 93a-c, May 17, 1951 [Holotype.--USNM 123700 from + San Lucas Camotlan, Oaxaca, Mexico; Walter S. Miller + collector]. + + _Ptychohyla leonhard-schultzei_, Duellman, Herpetologica, + 16:191-197, figs. 1-3, September 23, 1960; Univ. Kansas Publ. + Mus. Nat. Hist., 13:351, April 27, 1961. + + +_Diagnosis._--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. + +_Description._--The following description is based on KU 64117 from +Vista Hermosa, Oaxaca, Mexico (Pl. 14). 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 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. + +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. + +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). + +_Variation._--No noticeable geographic variation is apparent in the +few available specimens; variations in proportions are given in Table +1. 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. + +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. + +_Description of tadpole._--The following description is based on KU +68556 from 7.5 kilometers south of Yetla, Oaxaca, Mexico (Figs. 4C and +6C). 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. + +Mouth large; lips having deep lateral folds; two complete rows of +papillae on lips; five to seven rows of papillae laterally; beaks +moderately developed, bearing short peglike serrations; moderately +wide lateral projections on upper beak; tooth-rows 4/6; 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. + +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. + +_Variation._--The variation in size and proportions is given in Table +2. With the exception of one specimen having a short, broken, seventh +tooth-row, all specimens have 4/6 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. + +_Comparisons._--_Ptychohyla leonhardschultzei_ differs from all other +members of the _Ptychohyla euthysanota_ group in having a square +snout, and further differs from _P. spinipollex_ in more numerous and +smaller nuptial spines and in transverse, instead of posteromedially +slanting, vomerine processes between the inner nares. _Ptychohyla +leonhardschultzei_ differs from _P. euthysanota_ in having a rostral +keel and in having white flanks boldly spotted with black. + +All small hylids that are sympatric with _Ptychohyla +leonhardschultzei_ are either yellow (_Hyla dendroscarta_ and _H. +melanomma_) or green (_Hyla erythromma_, which has a red eye, _Hyla +hazelae_, which has a black line on the canthus, and _Ptychohyla +ignicolor_, which has red flash colors on the thighs). + +_Life History._--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 +(Pl. 11C). + +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. + +_Remarks._--Four specific names have been applied to this species. Ahl +(1934:185) based his description of _Hyla leonhardschultzei_ on a +small, poorly preserved female. Taylor (1944:41) proposed the generic +name _Ptychohyla_ for a species (named therein as _P. adipoventris_) +of hylid having ventrolateral glands and horn-covered nuptial spines. +Obviously, Taylor was unaware that _Hyla leonhardschultzei_ was the +same species. Earlier Taylor (1937:46) described _Hyla pinorum_. 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 _Hyla pinorum_ 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 _Ptychohyla +leonhard-schultzei_. Although Shannon (1951:473) based his description +of _Hyla milleri_ 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 _Hyla milleri_ is the same as +_Ptychohyla leonhardschultzei_. + +_Distribution._--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. + +_Specimens examined._--MEXICO: _Guerrero_: Agua del Obispo, CNHM +123489-90, 126651, 106300, MCZ 29639, UIMNH 25047, 25049, USNM 114551; +Malinaltepec, ZMB 34351, 34353. _Oaxaca_: 2.5 km. N of La Soledad, KU +58061; San Lucas Camotlan, 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); _7.5 km. S of Yetla_, +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). + + + + + [Illustration: PLATE 15 + _Ptychohyla spinipollex_ (KU 58054). x 2.] + + + =Ptychohyla spinipollex= (Schmidt) + + + _Hyla euthysanota_, Dunn and Emlen, Proc. Acad. Nat. Sci. + Philadelphia, 84:25, March 22, 1932. + + _Hyla spinipollex_ 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. + + _Ptychohyla spinipollex_, Stuart, Contr. Lab. Vert. Biol. Univ. + Michigan, 68:48, November, 1954. Duellman, Univ. Kansas Publ. + Mus. Nat. Hist., 13:351, April 27, 1961. + + +_Diagnosis._--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. + +_Description._--The following description is based on KU 58054 from +Finca Los Alpes, Depto. Alta Verapaz, Guatemala (Pl. 15). 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 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 V-shaped elevations between round inner nares; openings +to vocal sac large, one situated along inner posterior edge of each +mandibular ramus. + +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. + +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 (Cream-Buff) becoming +yellow (Lemon Chrome) in groin; spots on flanks dark brown (Clove +Brown); iris dull grayish bronze (Orange-Citrine). + +_Variation._--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 V-shaped elevations, but in some individuals the +elevations are more nearly transversely situated between the inner +nares. + +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). + +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 _P. spinipollex_, lacks a white stripe above the anal +opening, whereas the stripe is indistinct in UMMZ 113102-3. + +_Description of tadpole._--The following description is based on KU +60053 from Fina Los Alpes, Depto. Alta Verapaz, Guatemala (Figs. 4D +and 6D). 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. + +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 4/7; 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. + +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. + +_Variation._--The variation in size and proportions as compared with +tadpoles of other species is given in Table 2. Of the 57 tadpoles of +this species that I 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. + +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 _Ptychohyla +euthysanota_ 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 Morazan, Baja Verapaz, Guatemala, +have faint dorsal blotches on the dorsal musculature, much like those +in tadpoles of _Ptychohyla leonhardschultzei_. + +_Comparisons._--_Ptychohyla spinipollex_ 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. + +Other hylids sympatric with _Ptychohyla spinipollex_ include three +species of _Plectrohyla_, 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 _Ptychohyla spinipollex_ is _Hyla +bromeliacea_, which has a round snout and yellowish tan dorsum not +marked with dark brown. + +_Life History._--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 (Pl. 11A). + +Tadpoles have been found in cascading mountain streams. Two +metamorphosed young have snout-vent lengths of 15.0 and 15.5 mm. + +_Remarks._--There is little doubt that all of the specimens herein +referred to _Ptychohyla spinipollex_ are conspecific. However, the +three specimens from Honduras, including the type of _Ptychohyla +spinipollex_, 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. + +_Distribution._--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. + +_Specimens examined._--GUATEMALA: _Alta Verapaz_: Finca Chichen, 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); Panzamala, UMMZ 90878 (tadpoles). +_Baja Verapaz_: 32 km. N of Morazan, KU 68564 (tadpoles); _Santa +Elena_, UMMZ 98119, 98120 (2). _Huehuetenango_: 1 km. E of Barillas, +UMMZ 123136-7 (tadpoles). _Progreso_: Finca Bucaral, UMMZ 106783 (3), +123138 (tadpoles), S-1292 (skeleton). + +HONDURAS: _Atlantidad_: Mountains behind Ceiba, MCZ 21300. _Morazan_: +Cerro Uyuca, UMMZ 123102-3. + + + + +The _Ptychohyla schmidtorum_ Group + + +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 3/3 tooth-rows; +breeding call consisting of series of short notes. + + + + +=Ptychohyla schmidtorum= + + +_Diagnosis._--Diameter of tympanum more than half of diameter of eye; +internarial region depressed; toes three-fourths webbed; no red +flash-colors on thighs. + + + + + [Illustration: PLATE 16 + _Ptychohyla schmidtorum schmidtorum_ (KU 58043). x 2.] + + + =Ptychohyla schmidtorum schmidtorum= Stuart + + + _Ptychohyla schmidtorum_ 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. + + +_Diagnosis._--Vomerine teeth 5-11; dorsum dark brown; white spot below +eye; eye red in life. + +_Description._--The following description is based on KU 58043 from +Finca La Paz, Depto. San Marcos, Guatemala (Pl. 16). 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 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. + +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. + +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). + +_Variation._--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. + +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. + +_Description of tadpole._--The following description is based on KU +60051 from Finca La Paz, Depto. San Marcos, Guatemala (Figs. 5A and +6E). 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. + +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 3/3, all short; second and third +upper rows subequal in length; first upper row shorter; first 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. + +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. + +_Variation._--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 Table +2. 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. + +_Comparisons._--_Ptychohyla schmidtorum schmidtorum_ differs from _P. +schmidtorum chamulae_ as stated in the diagnosis and in having pale +creamy tan, as opposed to dark brown, webbing on the feet; and from +_P. ignicolor_ in having a depressed, as opposed to a flat, +internarial region. Tadpoles of _P. s. schmidtorum_ have a mottled +appearance, as opposed to the more uniform brown color of _P. s. +chamulae_. + +_Ptychohyla schmidtorum schmidtorum_ and several species of +_Plectrohyla_ are sympatric. All species of the latter genus have a +bony prepollex, rugose skin on the dorsum, and heavy body; also +sympatric is _Ptychohyla e. euthysanota_, which has a tarsal fold and +in breeding males spinous nuptial tuberosities. + +_Life History._--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 _Ptychohyla schmidtorum chamulae_. + +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. + +_Remarks._--There is no doubt that this frog is most closely related +to _Ptychohyla schmidtorum chamulae_, even though the ranges of the +two subspecies are separated by the interior depression of Chiapas. +Since at least at Finca La Paz, Guatemala, _P. s. schmidtorum_ occurs +with _P. e. euthysanota_, it is surprising that the former species has +not been found at more localities along the Pacific slopes on northern +Central America. At Finca La Paz in July, 1960, _P. s. schmidtorum_ +was more abundant than _P. e. euthysanota_. + +_Distribution._--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. + +_Specimens examined._--MEXICO: _Chiapas_: Finca Irlandia, UMMZ +105429-30. + +GUATEMALA: _San Marcos_: 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). + + + + + [Illustration: PLATE 17 + _Ptychohyla schmidtorum chamulae_ (KU 58069). x 2.] + + + =Ptychohyla schmidtorum chamulae= Duellman + + + _Ptychohyla chamulae_ 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 Rayon Mescalapa, Chiapas, + Mexico; William E. Duellman collector]. + + +_Diagnosis._--Vomerine teeth 4-6; dorsum bright green; white lateral +stripe; eye reddish bronze in life. + +_Description._--The following description is based on KU 58069 from +6.2 kilometers south of Rayon Mescalapa, Chiapas, Mexico (Pl. 17). +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 between ovoid inner nares; openings to vocal sac +large, one situated along inner posterior edge of each mandibular +ramus. + +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. + +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). + +_Variation._--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 _P. ignicolor_. + +_Description of tadpole._--The following description is based on KU +58199 from 6.2 kilometers south of Rayon Mescalapa, Chiapas, Mexico +(Figs. 5B and 6F). 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. + +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 3/3, +all short; second and third upper rows subequal in length; first upper +row shorter; first 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. + +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. + +_Variation._--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. + +_Comparisons._--Aside from the characters listed in the diagnosis, +_Ptychohyla schmidtorum chamulae_ differs from _P. schmidtorum +schmidtorum_ 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. _Ptychohyla ignicolor_ 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 _P. schmidtorum chamulae_. + +_Plectrohyla matudai matudai_ and _P. guatemalensis_ are sympatric +with _Ptychohyla schmidtorum chamulae_. Each of the first two has a +bony prepollex, rugose skin on the dorsum, and heavy body. Also living +with _Ptychohyla chamulae_ are _Hyla chaneque_, a large species having +a tuberculate dorsum and webbed fingers, and _Hyla bivocata_, a small +yellow species having a broad, flat head, small indistinct tympanum, +and an axillary membrane. + +_Life History._--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 (Pl. 11D). The call is +almost indistinguishable from that of _Ptychohyla schmidtorum +schmidtorum_. + +Tadpoles were found in the cascading streams; the smallest tadpole has +a total length of 17.2 mm. and has only 3/2 tooth-rows. At a stream +6.2 kilometers south of Rayon 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. + +_Remarks._--Duellman (1961:354) described _Ptychohyla chamulae_ and +stated that it probably was most closely related to _P. schmidtorum_. +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. _Ptychohyla schmidtorum chamulae_ +inhabits cloud forest on the Atlantic slopes of the Chiapan Highlands, +whereas _P. s. schmidtorum_ 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. + +_Distribution._--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. + +_Specimens examined._--MEXICO: _Chiapas: 15 km. N of Pueblo Nuevo +Solistahuacan_, UMMZ 123325 (4); _16.5 km. N of Pueblo Nuevo +Solistahuacan_, UMMZ 123322 (10); _18 km. N of Pueblo Nuevo +Solistahuacan_, UMMZ 121395-9, 123324 (8), 123326 (5); _18.6 km. N of +Pueblo Nuevo Solistahuacan_, UMMZ 123323 (4); _5.6 km. S of Rayon +Mescalapa_, KU 58062, 58200 (tadpoles); _6.2 km. S of Rayon +Mescalapa_, KU 58063-74, 58199 (tadpole), 58234-8, 59936 (skeleton). + + + + + [Illustration: PLATE 18 + _Ptychohyla ignicolor_ (UMMZ 119603). x 2.] + + + =Ptychohyla ignicolor= Duellman + + + _Ptychohyla ignicolor_ 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, + Mexico; Thomas E. Moore collector]. + +_Diagnosis._--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. + +_Description._--The following description is based on UMMZ 119603 from +6 kilometers south of Vista Hermosa, Oaxaca, Mexico (Pl. 18). 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; 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. + +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. + +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). + +_Variation._--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 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. + +_Description of tadpole._--The following description is based on KU +71716 from Vista Hermosa, Oaxaca, Mexico (Figs. 5C and 6G). 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. + +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 3/3, 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. + +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. + +_Variation._--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. + +_Comparisons._--From _P. schmidtorum chamulae_ and _P. s. +schmidtorum_, _P. ignicolor_ differs as follows: Tympanum smaller; +snout more nearly square; less webbing on toes; internarial region +flat instead of depressed; white lateral stripes lacking. + +_Ptychohyla ignicolor_ and several small and moderate sized hylids are +sympatric. From _P. ignicolor_ these hylids can be distinguished as +follows: _Hyla dendroscarta_ has a round snout and yellow dorsum; +_Hyla erythromma_ has a round snout, green dorsum, white flanks, and a +red eye; _Hyla hazelae_ has a tarsal fold, green dorsum, and a black +line on the canthus; and _Ptychohyla leonhardschultzei_ has a tarsal +fold, brown dorsum, black and white flanks, and horny nuptial spines +in breeding males. + +_Life History._--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 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 (Pl. 11E). + +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. + +_Remarks._--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 _Ptychohyla ignicolor_ in the _P. +schmidtorum_-group. + +_Distribution._--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. + +_Specimens examined._--MEXICO: _Oaxaca: Vista Hermosa_, KU 71334, +71716 (tadpoles), UMMZ 119602; 6 km. S of Vista Hermosa, KU 71335-42, +71343 (skeleton), UMMZ 119603, 123327 (2). + + + + + DISTRIBUTION AND ECOLOGY + + Geographic Distribution of the Species + + +The distribution of species of _Ptychohyla_ reflects the distribution +of cloud forest in southern Mexico and northern Central America. +The frogs are restricted to mountainous areas, usually at elevations +higher than 1000 meters above sea level. _Ptychohyla_ 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 +_Hyla bistincta_ group, and in Chiapas and Guatemala by species of +_Plectrohyla_. + +Frogs of the _Ptychohyla euthysanota_ group have a greater combined +geographic range than the species comprising the _Ptychohyla +schmidtorum_ 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 _P. leonhardschultzei_ 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 _P. euthysanota euthysanota_ is along +the Pacific slopes of the Sierra Madre in Chiapas, Guatemala, and +El Salvador, whereas that of _P. euthysanota macrotympanum_ is +along the southern interior slopes of the Central Highlands of +Chiapas and the Sierra de Cuchumatanes in Guatemala. _Ptychohyla +spinipollex_ occurs on the wet Atlantic slopes of the Guatemalan and +Honduranean Highlands; the range of the species in Honduras is +poorly known. + + + [Illustration: FIG. 7. Map showing locality records for the species + and subspecies of _Ptychohyla_.] + + +The frogs of the _Ptychohyla schmidtorum_ group have more restricted +geographic ranges than members of the former group. _Ptychohyla +schmidtorum schmidtorum_ occurs on the Pacific slopes of the Sierra +Madre in Chiapas and Guatemala, where it occurs with _P. euthysanota +euthysanota_; _P. schmidtorum chamulae_ 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, _P. euthysanota +macrotympanum_. On the Atlantic slopes of the Sierra Madre Oriental in +northern Oaxaca _P. ignicolor_ occurs with _P. leonhardschultzei_. + +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 Leon, _Hyla miotympanum_ seems to be the ecological +replacement of _Ptychohyla_. On the Pacific slopes north of Guerrero, +Mexico, humid forests in which there are cascading mountain streams +are absent; consequently, no _Ptychohyla_ are known from that region. +In the mountains of El Salvador _Ptychohyla euthysanota euthysanota_ +occurs sympatrically with another small stream-breeding hylid, _Hyla +salvadorensis_. To the south of Honduras the highlands diminish into +the lowlands of Nicaragua, where habitat suitable for _Ptychohyla_ +apparently does not exist. In the mountains of Costa Rica and Panama, +the habitats occupied by _Ptychohyla_ in northern Central America are +filled by a variety of stream-breeding _Hyla_, such as _Hyla legleri_, +_H. rivularis_, _H. rufioculis_, _H. alleei_, and _H. uranochroa_. + +Although members of the genus _Ptychohyla_ 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 +_Ptychohyla_ is a counterpart of _Plectrohyla_. + + + + + Habitat Preference + +Frogs of the genus _Ptychohyla_ 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, _P. leonhardschultzei_ and _P. +ignicolor_ were found in cloud forest, whereas at Agua del Obispo, +Guerrero, the former species was found in pine-oak forest. _Ptychohyla +schmidtorum_ is known only from cloud forest; _P. euthysanota +euthysanota_ and _P. spinipollex_ generally are found in cloud forest, +but in some places they live in pine-oak forest. _Ptychohyla +euthysanota macrotympanum_ 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 _Ptychohyla schmidtorum_ 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. + +_Ptychohyla_ 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 +_Ptychohyla_. As has been shown previously, the tadpoles of +_Ptychohyla_ 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 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 +_Ptychohyla_. + + + + + Interspecific Competition + +At localities where two species of _Ptychohyla_ occur sympatrically +(_P. ignicolor_ and _P. leonhardschultzei_ at Vista Hermosa, Oaxaca, +and _P. euthysanota euthysanota_ and _P. schmidtorum schmidtorum_ 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 _Ptychohyla_ acts +as an important reproductive isolating mechanism. + +At no locality were _Ptychohyla_ 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 _Ptychohyla_ and other +sympatric hylids. + +At Finca La Paz, Guatemala, tadpoles of two species of _Ptychohyla_ +were ecologically segregated. The tadpoles of _P. euthysanota +euthysanota_ were found in riffles in the streams, whereas those of +_P. schmidtorum schmidtorum_ were found in slower water, chiefly in +small pools in the streams. At Vista Hermosa, Oaxaca, tadpoles of _P. +leonhardschultzei_ were found in riffles, and tadpoles of the +sympatric _P. ignicolor_ 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 _Ptychohyla_ east of the +Isthmus of Tehuantepec, members of the genus occur with species of +_Plectrohyla_, all of which are larger than _Ptychohyla_, and all of +which have tadpoles that live in torrential streams. Tadpoles of +_Ptychohyla spinipollex_ have been found in streams inhabited by the +tadpoles of _Plectrohyla guatemalensis_ and _P. quecchi_; tadpoles of +_Ptychohyla euthysanota euthysanota_ and _P. schmidtorum schmidtorum_ +were found in streams inhabited by tadpoles of _Plectrohyla +guatemalensis_, _P. matudai_, and _P. sagorum_. 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. + + + + + Reproduction and Development + +Since the cloud forests inhabited by _Ptychohyla_ 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 _P. leonhardschultzei_ 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: _P. euthysanota euthysanota_, +February, March, May, and July; _P. euthysanota macrotympanum_, March, +June, and August; _P. spinipollex_, February, March, April, June, +July, and August; _P. schmidtorum schmidtorum_, March, May, June, +July, and August; _P. schmidtorum chamulae_, June and August; _P. +ignicolor_, June. I suspect that this temporal distribution more +accurately reflects the seasonal activities of collectors than of the +frogs. + +Calling frogs usually are on vegetation adjacent to or overhanging +streams; some calling males of _P. spinipollex_ were on rocks in or by +streams. Clasping pairs of _P. euthysanota_ and _P. schmidtorum_ were +observed on vegetation by streams. Despite intensive search, no eggs +were found. It is doubtful if _Ptychohyla_ deposit eggs on vegetation +overhanging streams, as do centrolenids and _Phyllomedusa_, 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 _Ptychohyla_ 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. + +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 _Ptychohyla schmidtorum_ group, have fewer eggs than do the larger +species. Numbers of eggs found in females of the various species are: +_P. euthysanota euthysanota_, 108; _P. euthysanota macrotympanum_, 136, +160; _P. leonhardschultzei_, 141; _P. spinipollex_, 119, 134, 143; +_P. schmidtorum schmidtorum_, 59, 61, 90; _P. schmidtorum chamulae_, +60, 71, 89. + +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 +_P. ignicolor_ of which none is available. The smallest young frog is +a _P. euthysanota_ having a snout-vent length of 14.2 mm.; the largest +young frog is a _P. schmidtorum schmidtorum_ having a snout-vent +length of 17.0 mm. + + + + + PHYLOGENY OF PTYCHOHYLA + + +The preceding data on morphology, life histories, and behavior form +the basis for the following interpretation of the phylogeny of +_Ptychohyla_. Additional data are needed to support some of the ideas +discussed below; many of the data that are available for _Ptychohyla_ +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 _Ptychohyla_ 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 _Ptychohyla_, some estimate of relationships +can be made. First, it is necessary to determine the validity of the +genus itself. + + + Ptychohyla as a Natural Assemblage + +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 _Ptychohyla_ 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. + +The species of _Ptychohyla_ are divided into two groups on the basis +of external morphology, breeding calls, and tadpoles. The _Ptychohyla +euthysanota_ 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 the _Ptychohyla +schmidtorum_ 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 _Ptychohyla euthysanota_ group +from a hylid stock containing _Hyla miotympanum_ and _Hyla +mixomaculata_. Likewise, _Ptychohyla schmidtorum_ could be placed with +_Hyla uranochroa_ and related species in Costa Rica. Nonetheless, the +fact remains that all of the species assigned to the genus +_Ptychohyla_ have ventrolateral glands in the breeding males; +furthermore, ventrolateral glands are unknown in other hylids. If the +_P. schmidtorum_ group and the _P. euthysanota_ 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 Mexico +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 _Ptychohyla_. + + + + + Generic Relationships + +The affinities of _Ptychohyla_ apparently are not with any of the +other groups that have been generically separated from _Hyla_. Of the +daughter genera in Middle America only _Plectrohyla_ has +stream-adapted tadpoles, but these large frogs are not closely related +to _Ptychohyla_. Stuart (1954:169) suggested that certain montane +species of _Hyla_ in lower Central America and _Hyla salvadorensis_ in +El Salvador may be related to _Ptychohyla_ 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 _Hyla_ in Costa +Rica--_H. alleei_, _H. legleri_, _H. rufioculis_, and _H. uranochroa_, +plus _Hyla salvadorensis_ in the mountains of El Salvador. +Morphologically all of the species are similar; _Hyla uranochroa_, _H. +legleri_, and _H. rufioculis_ have a lateral white stripe that is +expanded to form a spot beneath the eye, as in _Ptychohyla +schmidtorum_. The tadpoles of _Hyla rufioculis_ and _H. uranochroa_ +have large funnel-shaped mouths and long slender tails like those of +_Ptychohyla schmidtorum_. Lips of the tadpoles of _H. legleri_ and _H. +salvadorensis_ are folded laterally, in this respect resembling those +of the _Ptychohyla euthysanota_ group. I do not know the tadpoles and +the breeding call of _Hyla alleei_. The breeding calls of _Hyla +rufioculis_ and _H. uranochroa_ consist of high melodious notes; the +calls of _H. legleri_ and _H. salvadorensis_ consist of series of +short notes that have the general characteristics of the call of +_Ptychohyla schmidtorum_. Affinities of the genus _Ptychohyla_ seem to +me to be with the red-eyed species forming the _Hyla uranochroa_ group +in Costa Rica. All of the species in the _Hyla uranochroa_ 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 _Hyla +uranochroa_ group and to _Ptychohyla_ 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 _Hyla uranochroa_ group developed, and to the +Chiapas-Guatemala highlands, where _Ptychohyla_ evolved. + + + + + Interspecific Relationships + +_Ptychohyla schmidtorum_ is thought to resemble more closely the +parental stock of the genus than does any other species of +_Ptychohyla_ now extant. This parental stock is discussed above in the +account of the generic relationships. _Ptychohyla schmidtorum_ 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 _Hyla uranochroa_ group. +Probably during times of glaciation during the Pleistocene, when +climates in Mexico and Central America were depressed, the +_Ptychohyla_ 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 _Ptychohyla_ +across the Isthmus of Tehuantepec into the mountains of Oaxaca. + +Because of its small nuptial spines, small triangular vomers, +coloration, and absence of a rostral keel, _Ptychohyla euthysanota_, +more than any of the other species in the _P. euthysanota_ group, +resembles _P. schmidtorum_. At the present time _P. euthysanota_ and +_P. schmidtorum_ are sympatric. + +As I have mentioned previously, ecological segregation and +interspecific competition probably is highly developed in the tadpoles +of _Ptychohyla_. If this ecological segregation resulted from +intraspecific competition in a stock of _Ptychohyla_, possibly _P. +euthysanota_ and _P. schmidtorum_ differentiated sympatrically in this +way. Specific identity is maintained, at least in part, by different +breeding calls in males. + +_Ptychohyla spinipollex_ and _P. leonhardschultzei_ seem to be more +closely related to one another than either is to _P. euthysanota_. +Probably a stock of _P. euthysanota_ was isolated on the Atlantic +slopes of northern Central America from _P. euthysanota_ 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 _P. +leonhardschultzei_, while the stock on the northern slopes of Central +America evolved into _P. spinipollex_. Subsequent to the +differentiation of _P. leonhardschultzei_ and _P. spinipollex_ from +_P. euthysanota_ and during a time of cooler more equable climate than +exists now, _P. euthysanota_ and _P. schmidtorum_ invaded the Central +Highlands of Chiapas. Subsequent climatic changes isolated populations +of each in the Central Highlands, where _P. euthysanota macrotympanum_ +and _P. schmidtorum chamulae_ evolved. _Ptychohyla ignicolor_ probably +represents stock of _P. schmidtorum_ that crossed the Isthmus of +Tehuantepec and became isolated in Oaxaca on the western side of the +isthmus. + + + + + LITERATURE CITED + + + AHL, E. + 1934. Ueber eine sammlung von Reptilien und Amphibien aus + Mexiko, Zool. Anz., 106:184-186, April 15. + + BLAIR, W. F. + 1956. Call difference as an isolation mechanism in + southwestern toads (genus _Bufo_). Texas Jour. Sci., + 8:87-106, March. + + DUELLMAN, W. E. + 1956. The frogs of the hylid genus _Phrynohyas_ Fitzinger, + 1843. Misc. Publ. Mus. Zool. Univ. Michigan, 96:1-47, + pls. 1-6, February 21. + + 1960. Synonymy, variation, and distribution of _Ptychohyla + leonhardschultzei_ Ahl. Studies of American Hylid Frogs, + IV. Herpetologica, 16:191-197, September 23. + + 1961. Descriptions of two species of frogs, genus Ptychohyla. + Studies of American Hylid Frogs, V. Univ. Kansas Publ. + Mus. Nat. Hist., 13:349-357, pl. 25, April 27. + + FOUQUETTE, M. J. + 1960. Isolating mechanisms in three sympatric treefrogs in the + Canal Zone. Evolution, 14:484-497, December. + + KELLOGG, R. + 1928. An apparently new _Hyla_ from El Salvador. Proc. Biol. + Soc. Washington, 41:123-124, June 29. + + MERTENS, R. + 1952. Die Amphibien und Reptilien von El Salvador. + Senckenbergischen Naturf. Gesell., 487:1-120, pls. 1-16, + December 1. + + RIDGWAY, R. + 1912. Color standards and color nomenclature. Washington, + D. C., 44 pp., 53 pls. + + SHANNON, F. A. + 1951. Notes on a herpetological collection from Oaxaca and + other localities in Mexico. Proc. U. S. Nat. Mus., + 101:465-484, May 17. + + STUART, L. C. + 1954. Descriptions of some new amphibians and reptiles from + Guatemala. Proc. Biol. Soc. Washington, 67:159-178, + August 5. + + TANNER, W. W. + 1957. Notes on a collection of amphibians and reptiles from + southern Mexico, with a description of a new _Hyla_. + Great Basin Nat., 17:52-56, July 31. + + TAYLOR, E. H. + 1937. New species of hylid frogs from Mexico with comments on + the rare _Hyla bistincta_ Cope. Proc. Biol. Soc. + Washington, 50:43-54, pls. 2-3, April 21. + + 1942. New tailless amphibia from Mexico. Univ. Kansas Sci. + Bull., 28: 67-89, May 15. + + 1944. A new genus and species of Mexican frogs. Univ. Kansas + Sci. Bull., 30:41-45, June 12. + + 1949. New or unusual Mexican amphibians. Amer. Mus. Novitates, + 1437:1-21, December 7. + + +_Transmitted December 27, 1962._ + + + + + UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY + + +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. + +* 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: + + + Vol. 1. Nos. 1-26 and index. Pp. 1-638, 1946-1950. + + *Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest. + Pp. 1-444, 140 figures in text. April 9, 1948. + + Vol. 3. *1. The avifauna of Micronesia, its origin, evolution, + and distribution. By Rollin H. Baker. Pp. 1-359, + 16 figures in text. June 12, 1951. + + *2. A quantitative study of the nocturnal migration of + birds. By George H. Lowery, Jr. Pp. 361-472, + 47 figures in text. June 29, 1951. + + 3. Phylogeny of the waxwings and allied birds. By M. + Dale Arvey. Pp. 473-530, 49 figures in text, + 13 tables. October 10, 1951. + + *4. 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. + + Index. Pp. 651-681. + + *Vol. 4. (Complete) American weasels. By E. Raymond Hall. + Pp. 1-466, 41 plates, 31 figures in text. + December 27, 1951. + + Vol. 5. Nos. 1-37 and index. Pp. 1-676, 1951-1953. + + *Vol. 6. (Complete) Mammals of Utah, _taxonomy and distribution_. + By Stephen D. Durrant. Pp. 1-549, 91 figures in text, + 30 tables. August 10, 1952. + + Vol. 7. Nos. 1-15 and index. Pp. 1-651, 1952-1955. + + Vol. 8. Nos. 1-10 and index. Pp. 1-675, 1954-1956. + + Vol. 9. *1. Speciation of the wandering shrew. By James S. + Findley. Pp. 1-68, 18 figures in text. + December 10, 1955. + + 2. 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. + + 3. A new long-eared myotis (Myotis evotis) from + northeastern Mexico. By Rollin H. Baker and Howard J. + Stains. Pp. 81-84. December 10, 1955. + + 4. Subspeciation in the meadow mouse, Microtus + pennsylvanicus, in Wyoming. By Sydney Anderson. + Pp. 85-104, 2 figures in text. May 10, 1956. + + 5. The condylarth genus Ellipsodon. By Robert W. Wilson. + Pp. 105-116, 6 figures in text. May 19, 1956. + + 6. Additional remains of the multituberculate genus + Eucosmodon. By Robert W. Wilson. Pp. 117-123, + 10 figures in text. May 19, 1956. + + 7. Mammals of Coahulia, Mexico. By Rollin H. Baker. + Pp. 125-335, 75 figures in text. June 15, 1956. + + 8. 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. + + 9. Extensions of known ranges of Mexican bats. By + Sydney Anderson. Pp. 347-351. August 15, 1956. + + 10. A new bat (Genus Leptonycteris) from Coahulia. By + Howard J. Stains. Pp. 353-356. January 21, 1957. + + 11. A new species of pocket gopher (Genus Pappogeomys) + from Jalisco, Mexico. By Robert J. Russell. + Pp. 357-361. January 21, 1957. + + 12. Geographic variation in the pocket gopher, Thomomys + bottae, in Colorado. By Phillip M. Youngman. + Pp. 363-387, 7 figures in text. February 21, 1958. + + 13. New bog lemming (genus Synaptomys) from Nebraska. + By J. Knox Jones, Jr. Pp. 385-388. May 12, 1958. + + 14. Pleistocene bats from San Josecito Cave, Nuevo Leon, + Mexico. By J. Knox Jones, Jr. Pp. 389-396. + December 19, 1958. + + 15. New subspecies of the rodent Baiomys from Central + America. By Robert L. Packard. Pp. 397-404. + December 19, 1958. + + 16. Mammals of the Grand Mesa, Colorado. By Sydney + Anderson. Pp. 405-414, 1 figure in text, + May 20, 1959. + + 17. 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. + + 18. 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. + + 19. 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. + + 20. Small carnivores from San Josecito Cave (Pleistocene), + Nuevo Leon, Mexico. By E. Raymond Hall. Pp. 531-538, + 1 figure in text. January 14, 1960. + + 21. Pleistocene pocket gophers from San Josecito Cave, + Nuevo Leon, Mexico. By Robert J. Russell. + Pp. 539-548,1 figure in text. January 14, 1960. + + 22. Review of the insectivores of Korea. By J. Knox + Jones, Jr., and David H. Johnson. Pp. 549-578. + February 23, 1960. + + 23. 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. + + Index. Pp. 671-690 + + Vol. 10. 1. 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. + + 2. Comparative breeding behavior of Ammospiza caudacuta + and A. maritima. By Glen E. Woolfenden. Pp. 45-75, + 6 plates, 1 figure. December 20, 1956. + + 3. 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. + + 4. 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. + + 5. 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. + + *6. 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. + + 7. 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. + + 8. Natural history of the salamander, Aneides hardyi. + By Richard F. Johnston and Gerhard A. Schad. + Pp. 573-585. October 8, 1959. + + 9. A new subspecies of lizard, Cnemidophorus sacki, + from Michoacan, Mexico. By William E. Duellman. + Pp. 587-598, 2 figures in text. May 2, 1960. + + 10. 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. + + Index. Pp. 611-626. + + Vol. 11. Nos. 1-10 and index. Pp. 1-703, 1958-1960. + + Vol. 12. 1. Functional morphology of three bats: Sumops, Myotis, + Macrotus. By Terry A. Vaughan. Pp. 1-153, 4 plates, + 24 figures in text. July 8, 1959. + + *2. 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. + + 3. The baculum in microtine rodents. By Sydney Anderson. + Pp. 181-216, 49 figures in text. February 19, 1960. + + *4. 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. + + 5. Natural history of the bell vireo. By Jon C. Barlow. + Pp. 241-296, 6 figures in text. March 7, 1962. + + 6. Two new pelycosaurs from the lower Permian of + Oklahoma. By Richard C. Fox. Pp. 297-307, 6 figures + in text. May 21, 1962. + + 7. Vertebrates from the barrier island of Tamaulipas, + Mexico. By Robert K. Selander, Richard F. Johnston, + B. J. Wilks, and Gerald G. Raun. Pp. 309-345, + pls. 5-8. June 18, 1962. + + 8. Teeth of Edestid sharks. By Theodore H. Eaton, Jr. + Pp. 347-362, 10 figures in text. October 1, 1962. + + More numbers will appear in volume 12. + + + Vol. 13. 1. Five natural hybrid combinations in minnows + (Cyprinidae). By Frank B. Cross and W. L. Minckley. + Pp. 1-18. June 1, 1960. + + 2. A distributional study of the amphibians of the + Isthmus of Tehuantepec, Mexico. By William E. + Duellman. Pp. 19-72, pls. 1-8, 3 figures in text. + August 16, 1960. + + 3. A new subspecies of the slider turtle (Pseudemys + scripta) from Coahulia, Mexico. By John M. Legler. + Pp. 73-84, pls. 9-12, 3 figures in text. + August 16, 1960. + + 4. Autecology of the copperhead. By Henry S. Fitch. + Pp. 85-288, pls. 13-20, 26 figures in text. + November 30, 1960. + + 5. 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. + + 6. 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. + + 7. 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. + + 8. Descriptions 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. + + 9. 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. + + 10. 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. + + Index. Pp. 613-624. + + Vol. 14. 1. Neotropical bats from western Mexico. By Sydney + Anderson. Pp. 1-8. October 24, 1960. + + 2. 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. + + 3. Mammals of Mesa Verde National Park, Colorado. + By Sydney Anderson. Pp. 29-67, pls. 1 and 2, + 3 figures in text. July 24, 1961. + + 4. 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. + + 5. 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. + + 6. 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. + + 7. 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. + + 8. A new subspecies of ground squirrel (Spermophilus + spilosoma) from Tamaulipas, Mexico. By Ticul Alvarez. + Pp. 121-124. March 7, 1962. + + 9. 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. + + 10. 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. + + 11. A new subspecies of wood rat (Neotoma) from + northeastern Mexico. By Ticul Alvarez. Pp. 139-143. + April 30, 1962. + + 12. 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. + + 13. A new bat (Myotis) from Mexico. By E. Raymond Hall. + Pp. 161-164, 1 figure in text. May 21, 1962. + + 14. The mammals of Veracruz. By E. Raymond Hall and + Walter W. Dalquest. Pp. 165-362, 2 figures. + May 20, 1963. + + 15. The recent mammals of Tamaulipas, Mexico. By Ticul + Alvarez. Pp. 363-473, 5 figures in text. + May 20, 1963. + + More numbers will appear in volume 14. + + Vol. 15. 1. The amphibians and reptiles of Michoacan, Mexico. + By William E. Duellman. Pp. 1-148, pls. 1-6, + 11 figures in text. December 20, 1961. + + 2. Some reptiles and amphibians from Korea. By Robert + G. Webb, J. Knox Jones, Jr., and George W. Byers. + Pp. 149-173. January 31, 1962. + + 3. A new species of frog (Genus Tomodactylus) from + western Mexico. By Robert G. Webb. Pp. 175-181, + 1 figure in text. March 7, 1962. + + 4. Type specimens of amphibians and reptiles in the + Museum of Natural History, the University of Kansas. + By William E. Duellman and Barbara Berg. Pp. 183-204. + October 26, 1962. + + 5. Amphibians and Reptiles of the Rainforests of + Southern El Peten, Guatemala. By William E. Duellman. + Pp. 205-249, pls. 7-10, 6 figures in text. + October 4, 1963. + + 6. 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. + + 7. 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. + + More numbers will appear in volume 15. + + + * * * * * * * + +Transcriber's Notes. + +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. + +In the original, the Plates were grouped together between pages 328 +and 329. Here the Illustration: block which contains the text +associated with the Plates were moved just above the text in their +respective Systematic Account listing. The Plate text contain the +notation "x 2" after the caption to let the reader know that the image +was enlarged by a factor of two. + + +Typographical Errors Corrected: + +Several minor typographical corrections were made (missing periods, +commas, misspelling of 'and', etc.); but are not indicated here. More +substantial changes are listed below: + + References to the Plate 11 (Audiograms): + Pl. 1A, Pl. 1B, etc.=> Pl. 11A, Pl. 11B, etc. + Page 301, Paragraph 1: know => known + Page 302, Paragraph 1: Zoolegy => Zoology + Page 303, Paragraph 5: speces => species + Page 305, Paragraph 1: excresences => excrescences + Page 308, Paragraph 6: xiphisterum => xiphisternum + Page 316, Paragraph 3: with => width + Page 327, Paragraph 1: leonhard-schultzei => leonhardschultzei + to match remaining report text + Page 331, Paragraph 1: skelton => skeleton + Publication List Vol. 13, No. 8.: Decriptions => Descriptions + + + + + + + + + +End of the Project Gutenberg EBook of A Review of the Middle American Tree +Frogs of the Genus Ptychohyla, by William E. Duellman + +*** END OF THIS PROJECT GUTENBERG EBOOK MIDDLE AMERICAN TREE FROGS--PTYCHOHYLA *** + +***** This file should be named 35413.txt or 35413.zip ***** +This and all associated files of various formats will be found in: + https://www.gutenberg.org/3/5/4/1/35413/ + +Produced by Chris Curnow, Tom Cosmas, Joseph Cooper and +the Online Distributed Proofreading Team at +https://www.pgdp.net + + +Updated editions will replace the previous one--the old editions +will be renamed. + +Creating the works from public domain print editions means that no +one owns a United States copyright in these works, so the Foundation +(and you!) can copy and distribute it in the United States without +permission and without paying copyright royalties. Special rules, +set forth in the General Terms of Use part of this license, apply to +copying and distributing Project Gutenberg-tm electronic works to +protect the PROJECT GUTENBERG-tm concept and trademark. Project +Gutenberg is a registered trademark, and may not be used if you +charge for the eBooks, unless you receive specific permission. If you +do not charge anything for copies of this eBook, complying with the +rules is very easy. You may use this eBook for nearly any purpose +such as creation of derivative works, reports, performances and +research. They may be modified and printed and given away--you may do +practically ANYTHING with public domain eBooks. Redistribution is +subject to the trademark license, especially commercial +redistribution. + + + +*** START: FULL LICENSE *** + +THE FULL PROJECT GUTENBERG LICENSE +PLEASE READ THIS BEFORE YOU DISTRIBUTE OR USE THIS WORK + +To protect the Project Gutenberg-tm mission of promoting the free +distribution of electronic works, by using or distributing this work +(or any other work associated in any way with the phrase "Project +Gutenberg"), you agree to comply with all the terms of the Full Project +Gutenberg-tm License (available with this file or online at +https://gutenberg.org/license). + + +Section 1. General Terms of Use and Redistributing Project Gutenberg-tm +electronic works + +1.A. By reading or using any part of this Project Gutenberg-tm +electronic work, you indicate that you have read, understand, agree to +and accept all the terms of this license and intellectual property +(trademark/copyright) agreement. If you do not agree to abide by all +the terms of this agreement, you must cease using and return or destroy +all copies of Project Gutenberg-tm electronic works in your possession. +If you paid a fee for obtaining a copy of or access to a Project +Gutenberg-tm electronic work and you do not agree to be bound by the +terms of this agreement, you may obtain a refund from the person or +entity to whom you paid the fee as set forth in paragraph 1.E.8. + +1.B. "Project Gutenberg" is a registered trademark. It may only be +used on or associated in any way with an electronic work by people who +agree to be bound by the terms of this agreement. There are a few +things that you can do with most Project Gutenberg-tm electronic works +even without complying with the full terms of this agreement. See +paragraph 1.C below. There are a lot of things you can do with Project +Gutenberg-tm electronic works if you follow the terms of this agreement +and help preserve free future access to Project Gutenberg-tm electronic +works. See paragraph 1.E below. + +1.C. The Project Gutenberg Literary Archive Foundation ("the Foundation" +or PGLAF), owns a compilation copyright in the collection of Project +Gutenberg-tm electronic works. Nearly all the individual works in the +collection are in the public domain in the United States. If an +individual work is in the public domain in the United States and you are +located in the United States, we do not claim a right to prevent you from +copying, distributing, performing, displaying or creating derivative +works based on the work as long as all references to Project Gutenberg +are removed. Of course, we hope that you will support the Project +Gutenberg-tm mission of promoting free access to electronic works by +freely sharing Project Gutenberg-tm works in compliance with the terms of +this agreement for keeping the Project Gutenberg-tm name associated with +the work. You can easily comply with the terms of this agreement by +keeping this work in the same format with its attached full Project +Gutenberg-tm License when you share it without charge with others. + +1.D. The copyright laws of the place where you are located also govern +what you can do with this work. Copyright laws in most countries are in +a constant state of change. If you are outside the United States, check +the laws of your country in addition to the terms of this agreement +before downloading, copying, displaying, performing, distributing or +creating derivative works based on this work or any other Project +Gutenberg-tm work. The Foundation makes no representations concerning +the copyright status of any work in any country outside the United +States. + +1.E. Unless you have removed all references to Project Gutenberg: + +1.E.1. The following sentence, with active links to, or other immediate +access to, the full Project Gutenberg-tm License must appear prominently +whenever any copy of a Project Gutenberg-tm work (any work on which the +phrase "Project Gutenberg" appears, or with which the phrase "Project +Gutenberg" is associated) is accessed, displayed, performed, viewed, +copied or distributed: + +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 + +1.E.2. If an individual Project Gutenberg-tm electronic work is derived +from the public domain (does not contain a notice indicating that it is +posted with permission of the copyright holder), the work can be copied +and distributed to anyone in the United States without paying any fees +or charges. If you are redistributing or providing access to a work +with the phrase "Project Gutenberg" associated with or appearing on the +work, you must comply either with the requirements of paragraphs 1.E.1 +through 1.E.7 or obtain permission for the use of the work and the +Project Gutenberg-tm trademark as set forth in paragraphs 1.E.8 or +1.E.9. + +1.E.3. If an individual Project Gutenberg-tm electronic work is posted +with the permission of the copyright holder, your use and distribution +must comply with both paragraphs 1.E.1 through 1.E.7 and any additional +terms imposed by the copyright holder. Additional terms will be linked +to the Project Gutenberg-tm License for all works posted with the +permission of the copyright holder found at the beginning of this work. + +1.E.4. Do not unlink or detach or remove the full Project Gutenberg-tm +License terms from this work, or any files containing a part of this +work or any other work associated with Project Gutenberg-tm. + +1.E.5. Do not copy, display, perform, distribute or redistribute this +electronic work, or any part of this electronic work, without +prominently displaying the sentence set forth in paragraph 1.E.1 with +active links or immediate access to the full terms of the Project +Gutenberg-tm License. + +1.E.6. You may convert to and distribute this work in any binary, +compressed, marked up, nonproprietary or proprietary form, including any +word processing or hypertext form. However, if you provide access to or +distribute copies of a Project Gutenberg-tm work in a format other than +"Plain Vanilla ASCII" or other format used in the official version +posted on the official Project Gutenberg-tm web site (www.gutenberg.org), +you must, at no additional cost, fee or expense to the user, provide a +copy, a means of exporting a copy, or a means of obtaining a copy upon +request, of the work in its original "Plain Vanilla ASCII" or other +form. Any alternate format must include the full Project Gutenberg-tm +License as specified in paragraph 1.E.1. + +1.E.7. Do not charge a fee for access to, viewing, displaying, +performing, copying or distributing any Project Gutenberg-tm works +unless you comply with paragraph 1.E.8 or 1.E.9. + +1.E.8. You may charge a reasonable fee for copies of or providing +access to or distributing Project Gutenberg-tm electronic works provided +that + +- You pay a royalty fee of 20% of the gross profits you derive from + the use of Project Gutenberg-tm works calculated using the method + you already use to calculate your applicable taxes. The fee is + owed to the owner of the Project Gutenberg-tm trademark, but he + has agreed to donate royalties under this paragraph to the + Project Gutenberg Literary Archive Foundation. Royalty payments + must be paid within 60 days following each date on which you + prepare (or are legally required to prepare) your periodic tax + returns. Royalty payments should be clearly marked as such and + sent to the Project Gutenberg Literary Archive Foundation at the + address specified in Section 4, "Information about donations to + the Project Gutenberg Literary Archive Foundation." + +- You provide a full refund of any money paid by a user who notifies + you in writing (or by e-mail) within 30 days of receipt that s/he + does not agree to the terms of the full Project Gutenberg-tm + License. You must require such a user to return or + destroy all copies of the works possessed in a physical medium + and discontinue all use of and all access to other copies of + Project Gutenberg-tm works. + +- You provide, in accordance with paragraph 1.F.3, a full refund of any + money paid for a work or a replacement copy, if a defect in the + electronic work is discovered and reported to you within 90 days + of receipt of the work. + +- You comply with all other terms of this agreement for free + distribution of Project Gutenberg-tm works. + +1.E.9. If you wish to charge a fee or distribute a Project Gutenberg-tm +electronic work or group of works on different terms than are set +forth in this agreement, you must obtain permission in writing from +both the Project Gutenberg Literary Archive Foundation and Michael +Hart, the owner of the Project Gutenberg-tm trademark. Contact the +Foundation as set forth in Section 3 below. + +1.F. + +1.F.1. Project Gutenberg volunteers and employees expend considerable +effort to identify, do copyright research on, transcribe and proofread +public domain works in creating the Project Gutenberg-tm +collection. Despite these efforts, Project Gutenberg-tm electronic +works, and the medium on which they may be stored, may contain +"Defects," such as, but not limited to, incomplete, inaccurate or +corrupt data, transcription errors, a copyright or other intellectual +property infringement, a defective or damaged disk or other medium, a +computer virus, or computer codes that damage or cannot be read by +your equipment. + +1.F.2. LIMITED WARRANTY, DISCLAIMER OF DAMAGES - Except for the "Right +of Replacement or Refund" described in paragraph 1.F.3, the Project +Gutenberg Literary Archive Foundation, the owner of the Project +Gutenberg-tm trademark, and any other party distributing a Project +Gutenberg-tm electronic work under this agreement, disclaim all +liability to you for damages, costs and expenses, including legal +fees. YOU AGREE THAT YOU HAVE NO REMEDIES FOR NEGLIGENCE, STRICT +LIABILITY, BREACH OF WARRANTY OR BREACH OF CONTRACT EXCEPT THOSE +PROVIDED IN PARAGRAPH 1.F.3. YOU AGREE THAT THE FOUNDATION, THE +TRADEMARK OWNER, AND ANY DISTRIBUTOR UNDER THIS AGREEMENT WILL NOT BE +LIABLE TO YOU FOR ACTUAL, DIRECT, INDIRECT, CONSEQUENTIAL, PUNITIVE OR +INCIDENTAL DAMAGES EVEN IF YOU GIVE NOTICE OF THE POSSIBILITY OF SUCH +DAMAGE. + +1.F.3. LIMITED RIGHT OF REPLACEMENT OR REFUND - If you discover a +defect in this electronic work within 90 days of receiving it, you can +receive a refund of the money (if any) you paid for it by sending a +written explanation to the person you received the work from. If you +received the work on a physical medium, you must return the medium with +your written explanation. The person or entity that provided you with +the defective work may elect to provide a replacement copy in lieu of a +refund. If you received the work electronically, the person or entity +providing it to you may choose to give you a second opportunity to +receive the work electronically in lieu of a refund. If the second copy +is also defective, you may demand a refund in writing without further +opportunities to fix the problem. + +1.F.4. Except for the limited right of replacement or refund set forth +in paragraph 1.F.3, this work is provided to you 'AS-IS' WITH NO OTHER +WARRANTIES OF ANY KIND, EXPRESS OR IMPLIED, INCLUDING BUT NOT LIMITED TO +WARRANTIES OF MERCHANTIBILITY OR FITNESS FOR ANY PURPOSE. + +1.F.5. Some states do not allow disclaimers of certain implied +warranties or the exclusion or limitation of certain types of damages. +If any disclaimer or limitation set forth in this agreement violates the +law of the state applicable to this agreement, the agreement shall be +interpreted to make the maximum disclaimer or limitation permitted by +the applicable state law. The invalidity or unenforceability of any +provision of this agreement shall not void the remaining provisions. + +1.F.6. INDEMNITY - You agree to indemnify and hold the Foundation, the +trademark owner, any agent or employee of the Foundation, anyone +providing copies of Project Gutenberg-tm electronic works in accordance +with this agreement, and any volunteers associated with the production, +promotion and distribution of Project Gutenberg-tm electronic works, +harmless from all liability, costs and expenses, including legal fees, +that arise directly or indirectly from any of the following which you do +or cause to occur: (a) distribution of this or any Project Gutenberg-tm +work, (b) alteration, modification, or additions or deletions to any +Project Gutenberg-tm work, and (c) any Defect you cause. + + +Section 2. Information about the Mission of Project Gutenberg-tm + +Project Gutenberg-tm is synonymous with the free distribution of +electronic works in formats readable by the widest variety of computers +including obsolete, old, middle-aged and new computers. It exists +because of the efforts of hundreds of volunteers and donations from +people in all walks of life. + +Volunteers and financial support to provide volunteers with the +assistance they need are critical to reaching Project Gutenberg-tm's +goals and ensuring that the Project Gutenberg-tm collection will +remain freely available for generations to come. In 2001, the Project +Gutenberg Literary Archive Foundation was created to provide a secure +and permanent future for Project Gutenberg-tm and future generations. +To learn more about the Project Gutenberg Literary Archive Foundation +and how your efforts and donations can help, see Sections 3 and 4 +and the Foundation web page at https://www.pglaf.org. + + +Section 3. Information about the Project Gutenberg Literary Archive +Foundation + +The Project Gutenberg Literary Archive Foundation is a non profit +501(c)(3) educational corporation organized under the laws of the +state of Mississippi and granted tax exempt status by the Internal +Revenue Service. The Foundation's EIN or federal tax identification +number is 64-6221541. Its 501(c)(3) letter is posted at +https://pglaf.org/fundraising. Contributions to the Project Gutenberg +Literary Archive Foundation are tax deductible to the full extent +permitted by U.S. federal laws and your state's laws. + +The Foundation's principal office is located at 4557 Melan Dr. S. +Fairbanks, AK, 99712., but its volunteers and employees are scattered +throughout numerous locations. Its business office is located at +809 North 1500 West, Salt Lake City, UT 84116, (801) 596-1887, email +business@pglaf.org. Email contact links and up to date contact +information can be found at the Foundation's web site and official +page at https://pglaf.org + +For additional contact information: + Dr. Gregory B. Newby + Chief Executive and Director + gbnewby@pglaf.org + + +Section 4. Information about Donations to the Project Gutenberg +Literary Archive Foundation + +Project Gutenberg-tm depends upon and cannot survive without wide +spread public support and donations to carry out its mission of +increasing the number of public domain and licensed works that can be +freely distributed in machine readable form accessible by the widest +array of equipment including outdated equipment. Many small donations +($1 to $5,000) are particularly important to maintaining tax exempt +status with the IRS. + +The Foundation is committed to complying with the laws regulating +charities and charitable donations in all 50 states of the United +States. Compliance requirements are not uniform and it takes a +considerable effort, much paperwork and many fees to meet and keep up +with these requirements. We do not solicit donations in locations +where we have not received written confirmation of compliance. To +SEND DONATIONS or determine the status of compliance for any +particular state visit https://pglaf.org + +While we cannot and do not solicit contributions from states where we +have not met the solicitation requirements, we know of no prohibition +against accepting unsolicited donations from donors in such states who +approach us with offers to donate. + +International donations are gratefully accepted, but we cannot make +any statements concerning tax treatment of donations received from +outside the United States. U.S. laws alone swamp our small staff. + +Please check the Project Gutenberg Web pages for current donation +methods and addresses. Donations are accepted in a number of other +ways including including checks, online payments and credit card +donations. To donate, please visit: https://pglaf.org/donate + + +Section 5. General Information About Project Gutenberg-tm electronic +works. + +Professor Michael S. Hart was the originator of the Project Gutenberg-tm +concept of a library of electronic works that could be freely shared +with anyone. For thirty years, he produced and distributed Project +Gutenberg-tm eBooks with only a loose network of volunteer support. + + +Project Gutenberg-tm eBooks are often created from several printed +editions, all of which are confirmed as Public Domain in the U.S. +unless a copyright notice is included. Thus, we do not necessarily +keep eBooks in compliance with any particular paper edition. + + +Most people start at our Web site which has the main PG search facility: + + https://www.gutenberg.org + +This Web site includes information about Project Gutenberg-tm, +including how to make donations to the Project Gutenberg Literary +Archive Foundation, how to help produce our new eBooks, and how to +subscribe to our email newsletter to hear about new eBooks. |