








                  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 México and northern Central America were named
and described. Despite the extensive collecting by Salvin and Godman,
Nelson and Goldman, and the various expeditions of the _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 México 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 Zoología de
Tuxtla Gutierrez, México (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 México, Guatemala,
and El Salvador for permission to work on their land and for providing
shelter, food, and guides. I am especially grateful to Mr. and Mrs.
Horatio Kelly of "Colegio Linda Vista" at Pueblo Nuevo Solistahuacán,
Chiapas, for a pleasant stay at their school; Jordi Juliá Z. of the
Comisión del Papaloapan, Ciudad Alemán, Veracruz, for arranging for
field work in northern Oaxaca in 1959; Walter Hannstein and Lothar
Menzel for the use of facilities at Finca La Paz, Guatemala, in 1960;
Alan Hempstead for the use of facilities at Finca Los Alpes, Guatemala
in 1960 and 1961; and Julio Aguirre C. of the Instituto Tropical de
Investigaciones Científicas, San Salvador, El Salvador, for providing
comfortable working quarters and transportation and guides to the
mountains in northern El Salvador. Without the cheerful efforts of
Jorge A. Ibarra, Director of the Museo Nacional de Historia Natural in
Guatemala, my field work would have been greatly restricted during
politically precarious times in that country. Permits to collect in
México were furnished by the late Luis Macías Arellano of the
Dirección General de Caza. Each of these individuals has my profound
thanks for his indispensable aid.

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). × 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, México, 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). × 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, México; Hobart M. and Rozella Smith
        collectors]. Taylor and Smith, Proc. U. S. Natl. Mus., 95:587,
        June 30, 1945. Smith and Taylor, Bull. U. S. Natl. Mus.,
        194:86, June 17, 1948. Stuart, Proc. Biol. Soc. Washington,
        67:169, August 5, 1954.

    _Ptychohyla bogerti_ Taylor, Amer. Mus. Novitates, 1437:13-16,
        fig. 5, December 7, 1949 [Holotype.--AMNH 51847 from Río
        Grande, Oaxaca, México; Thomas MacDougall collector]. Stuart,
        Proc. Biol. Soc. Washington, 67:169, August 5, 1954.

    _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, México, through Guatemala
to northern El Salvador; probably it occurs also in southwestern
Honduras. Aside from the specimens listed below, three in the
Frankfurt Museum from Depto. Santa Ana, El Salvador (44571, Hacienda
San José; 43040, Hacienda Los Planes; 65119, Miramundo) are listed by
Mertens (1952:29).

_Specimens examined._--MEXICO: _Chiapas: Cascarada, 30 km. W of
Cíltapec_, UMMZ 87851-2; Cerro Ovando, UMMZ 87853-4; Chicomuselo, UMMZ
94439-40; Finca Juárez, 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; Río Grande, AMNH 51847-8; Santo Tomás 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, Río Samalá,
CNHM 35066. _Santa Rosa_: Finca La Gloria, UMMZ 123148 (tadpoles),
123150 (tadpoles). _Sololá_: Finca Santo Tomás, UMMZ 123149
(tadpoles); Olas de Mocá, near Mocá, CNHM 20208.

EL SALVADOR: _Chalatenango_: Los Esemiles, USNM 73296. _Santa Ana_:
Miramundo, CNHM 65120.




  [Illustration: PLATE 13
        _Ptychohyla euthysanota macrotympanum_ (KU 58049). × 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, México; 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, México (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 Río Hondo, 9.5 kilometers south of Pueblo Nuevo
Solistahuacán, Chiapas, México (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 Río Hondo,
Chiapas, on June 16, 1960. The smallest tadpole has a total length of
24.1 mm.; in this individual the sixth lower tooth row has barely
started to develop. A metamorphosing frog taken at the same time has a
snout-vent length of 19.8 mm., a short remnant of the tail, and the
mouth and tongue developed, whereas another individual having a
snout-vent length of 17.8 mm. and a tail 31.0 mm. in length still has
larval teeth. Three completely metamorphosed juveniles collected by L.
C. Stuart at Jacaltenango, Guatemala, on June 6 and 7, 1955, have
snout-vent lengths of 16.0, 16.0, and 16.1 mm.

_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 Río Grijalva and Río Cuilco. The similarity in
structure of the adults and tadpoles and the indistinguishable
breeding calls are the basis for considering the two populations
subspecies.

_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 Río
Grijalva and Río 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 Solistahuacán, KU 58049-51, 59939 (skeleton);
_Río Hondo, 9.5 km. S of Pueblo Nuevo Solistahuacán_, 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). × 2.]


                 =Ptychohyla leonhardschultzei= (Ahl)


    _Hyla leonhard-schultzei_ Ahl, Zool. Anz., 106:185-186, fig. 1,
        April 15, 1934 [Holotype.--ZMB 34353 from Malinaltepec,
        Guerrero, México; Leonhard Schultze collector]. Smith and
        Taylor, Bull. U. S. Natl. Mus., 184:87, June 17, 1948.

    _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, México; 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, México; Edward T. Taylor collector]. Smith
        and Taylor, Bull. U. S. Natl. Mus., 194:91, June 17, 1948.
        Taylor, Amer. Mus. Novitates, 1437:16, December 7, 1949.
        Stuart, Proc. Biol. Soc. Washington, 67:169, August 5, 1954.

    _Hyla milleri_ Shannon, Proc. U. S. Natl. Mus., 101:473-477,
        figs. 92b, 93a-c, May 17, 1951 [Holotype.--USNM 123700 from
        San Lucas Camotlán, Oaxaca, México; Walter S. Miller
        collector].

    _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, México (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, México (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 Camotlán, UIMNH 3201, USNM 123700-1; Vista Hermosa,
KU 64116-7, 64119, 68560 (tadpoles), 71344, 71717-8 (tadpoles), UMMZ
119604; 5 km. S of Yetla, KU 60045 (tadpoles); _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). × 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 Morazán, 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 Chichén, UMMZ
90876 (tadpoles); Finca Los Alpes, KU 58052-60, 59939 (skeleton),
60053 (tadpoles), 64122-41, 68562, 68563 (tadpoles), 68631-2
(skeletons), MCZ 35000-1, UMMZ 90873, 90874 (3), 90875 (tadpoles); La
Primavera, UMMZ 90877 (tadpoles); Panzamalá, UMMZ 90878 (tadpoles).
_Baja Verapaz_: 32 km. N of Morazán, 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. _Morazán_:
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). × 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). × 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 Rayón Mescalapa, Chiapas,
        México; 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 Rayón Mescalapa, Chiapas, México (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 Rayón Mescalapa, Chiapas, México
(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 Rayón Mescalapa, Chiapas, metamorphosing young
were found on June 16 and August 5, 1960. Each of two completely
metamorphosed young has a snout-vent length of 15.7 mm. Another having
a snout-vent length of 16.2 mm. has a tail stub 2 mm. long and a
completely metamorphosed mouth. Two others have snout-vent lengths of
13.6 and 14.1 mm. and tail lengths of 11.5 and 8.1 mm. respectively;
in these the mouth parts are incompletely metamorphosed.

_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
Solistahuacán_, UMMZ 123325 (4); _16.5 km. N of Pueblo Nuevo
Solistahuacán_, UMMZ 123322 (10); _18 km. N of Pueblo Nuevo
Solistahuacán_, UMMZ 121395-9, 123324 (8), 123326 (5); _18.6 km. N of
Pueblo Nuevo Solistahuacán_, UMMZ 123323 (4); _5.6 km. S of Rayón
Mescalapa_, KU 58062, 58200 (tadpoles); _6.2 km. S of Rayón
Mescalapa_, KU 58063-74, 58199 (tadpole), 58234-8, 59936 (skeleton).




  [Illustration: PLATE 18
        _Ptychohyla ignicolor_ (UMMZ 119603). × 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,
        México; 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, México (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, México (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 México 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 León, _Hyla miotympanum_ seems to be the ecological
replacement of _Ptychohyla_. On the Pacific slopes north of Guerrero,
México, 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 Panamá,
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 México
and northern Central America and not in any of the other approximately
500 species of hylids in the world is untenable. It is more logical to
assume that the development of the glands took place only once in a
stock of hylids that gave rise to the five species herein recognized
as members of the genus _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 México 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. Über 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


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* 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 León,
                  México. 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 León, México. By E. Raymond Hall. Pp. 531-538,
                  1 figure in text. January 14, 1960.

              21. Pleistocene pocket gophers from San Josecito Cave,
                  Nuevo León, México. 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 Michoacán, México. 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,
                  México. 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, México. 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, México. 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 México. 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, México. 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 Michoacán, México.
                  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 México. 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 Petén, 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 "× 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









