The Amphibians and Reptiles of Michoacán, México
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Title: The Amphibians and Reptiles of Michoacán, México
Author: William Edward Duellman
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UNIVERSITY OF KANSAS PUBLICATIONS
MUSEUM OF NATURAL HISTORY
Volume 15, No. 1, pp. 1-148, pls. 1-6, 11 figs.
December 20, 1961
[Transcriber's Note: Words surrounded by cedillas, like ~this~ signifies
words in bold. Words surrounded by underscores, like _this_, signifies
words in italics.]
The Amphibians and Reptiles of Michoacan, Mexico
BY
WILLIAM E. DUELLMAN
UNIVERSITY OF KANSAS
LAWRENCE
1961
UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY
Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Theodore H. Eaton, Jr.
Volume 15, No. 1, pp. 1-148, pls. 1-6, 11 figs.
Published December 20, 1961
UNIVERSITY OF KANSAS
Lawrence, Kansas
PRINTED IN
THE STATE PRINTING PLANT
TOPEKA, KANSAS
1961
The Amphibians and Reptiles of Michoacan, Mexico
BY
WILLIAM E. DUELLMAN
CONTENTS
PAGE
INTRODUCTION 3
Acknowledgments 5
Historical Account 7
NATURAL LANDSCAPE 9
GEOGRAPHY OF THE HERPETOFAUNA 13
ANNOTATED LIST OF SPECIES 13
Amphibia 14
Caudata 14
Salientia 20
Reptilia 56
Testudines 56
Crocodilia 58
Sauria 59
Serpentes 88
SPECIES OF QUESTIONABLE OCCURRENCE 124
GAZETTEER 129
SUMMARY 141
LITERATURE CITED 142
INTRODUCTION
For almost 30 years North American herpetologists have been making
extensive collections of reptiles and amphibians in Mexico. Some parts
of the country, because of their accessibility, soon became relatively
well known; other regions lying off the beaten path were bypassed or
inadequately sampled. Principally in the last decade herpetologists have
been entering regions from which no collections previously were
available in an attempt to fill gaps in known distributions and to
discover unknown species of animals. In 1950 Dr. Donald D. Brand led an
exploration party from the University of Texas to the poorly explored
and faunistically unknown region of southwestern Michoacan. James A.
Peters accompanied Brand and collected amphibians and reptiles. In 1951
I welcomed the opportunity to accompany Brand on a second expedition to
southwestern Michoacan. Such was the beginning of my interest in the
herpetofauna of the region. I have been fortunate to return to Michoacan
on four successive trips, all of which had as their purpose the
accumulation of data on the herpetofauna that would result in a survey
of the component species and an analysis of their distribution.
My original intention was to amplify Peters' (1954) study based on the
collections made by him in 1950 and by me in 1951 in the Sierra de
Coalcoman. But it soon became evident that in order to understand the
relationships of the herpetofauna of the Sierra de Coalcoman, the
species inhabiting the Tepalcatepec Valley and adjacent mountain ranges
would have to be studied. In the course of making that study I examined
all specimens from Michoacan already in museums.
There have been few detailed herpetofaunal studies in Mexico. The first
such study of any consequence was that by Bogert and Oliver (1945) on
the herpetofauna of Sonora. In that paper the authors analyzed the fauna
from a geographic view and showed the transition from tropical species
in the southern part of the state to members of the Sonoran Desert
assemblage to the north. Martin (1958) made a detailed study of the
herpetofauna of the Gomez Farias region in southern Tamaulipas; he
emphasized the ecological distribution of amphibians and reptiles in
that region with special reference to cloud forests. Duellman (1958c)
presented a preliminary geographic analysis of the herpetofauna of
Colima with special reference to the continuity of the species
inhabiting the lowlands. Zweifel (1960) discussed in detail the
herpetofauna of the Tres Marias Islands and commented on the derivation
of the fauna. Duellman (1960d) provided a detailed account of the
geographic distribution of the amphibians known to occur in the lowlands
of the Isthmus of Tehuantepec and attempted to account for the present
patterns of distribution.
The present report is the first of two parts dealing with the
herpetofauna of Michoacan. The purpose of this part is to present a full
account of the species of amphibians and reptiles known to inhabit the
state of Michoacan; the accounts of the species are accompanied by a
brief description of the natural landscape and of the various
assemblages of species comprising the major faunistic groups within the
region. A gazetteer of collecting localities is appended. The second
part of the study, now in preparation, deals with the ecological and
historical geography of the herpetofauna. Since the present part will be
of interest primarily to systematic herpetologists, I have decided to
separate it from the more general material of interest to
biogeographers.
One of the major problems that faces the worker undertaking a faunal
study is the presence of species or genera of unsettled systematic
status. My work in Michoacan has been no exception; fifteen separate
studies were undertaken in an attempt to solve systematic problems in
certain groups. Some systematic problems still remain but are of little
consequence insofar as the entire faunal picture is concerned, or are so
involved as to be impractical to undertake at this time. In accounts of
species, such problems are mentioned in the hope that they will interest
some worker who will be inclined to investigate them.
ACKNOWLEDGMENTS
While engaged in the study of the herpetofauna of Michoacan I have built
up a debt of gratitude to many individuals, without whose aid my
ambition to complete my study never would have been realized. I am
especially grateful to those individuals who accompanied me in the
field; Lee D. Beatty, Richard E. Etheridge, Carter R. Gilbert, Fred G.
Thompson, Jerome Tulecke, and John Wellman offered stimulating
companionship and valuable assistance. On many occasions they suffered
hardships on behalf of my interests.
Studies of my own specimens have been augmented by material from other
institutions. For permitting me to examine specimens in their care I am
indebted to W. Frank Blair, Charles M. Bogert, Doris M. Cochran, William
B. Davis, James R. Dixon, the late Emmett R. Dunn, Josef Eiselt, Alice
G. C. Grandison, Norman Hartweg, Robert F. Inger, Arthur Loveridge, the
late Karl P. Schmidt, Hobart M. Smith, Robert C. Stebbins, Margaret
Storey, Edward H. Taylor, and Richard G. Zweifel.
Several people have aided me in the study of specimens and in the
analysis of data; I am grateful to Donald D. Brand, who first introduced
me to Michoacan; since that time I have benefited much from his
knowledge of the area. James A. Peters provided me with essential
information concerning his field work in southern Michoacan in 1950.
James R. Dixon and Floyd L. Downs have permitted me to use freely the
material and data that they accumulated in their recent field work in
Michoacan. Norman E. Hartweg allowed me to use the specimens and data
that he gathered in his survey of the herpetofauna in the region of
Volcan Paricutin. L. C. Stuart, Charles F. Walker, and Richard G.
Zweifel have helped in unraveling some of the systematic and
distributional problems.
I am especially grateful to my wife, Ann, who for six months helped me
track down elusive species and explore new areas. Furthermore, she has
stimulated me to carry this study to completion.
Many people in Michoacan favored the field parties with quarters,
transportation, and valuable information, which greatly facilitated the
field work. In this respect I am especially indebted to Ingeniero Ruben
Erbina of Ingenieros Civiles Asociados, who not only let us use his home
as our headquarters, but through a letter of introduction gave us the
"key" to southern Michoacan. Ingeniero Pedro Tonda aided us in Arteaga
and San Salvador. Ingeniero Anastacio Perez Alfaro of the Comision
Tepalcatepec in Uruapan provided the latest maps of southern Michoacan
and much essential information pertaining to travel conditions in the
area. Senor Nefty Mendoza gave us a home in Dos Aguas; this kindness
allowed us to work in this interesting region during the height of the
rainy season. Mr. and Mrs. Bob Thomas let us make use of their
facilities at Hacienda Zirimicuaro. The naval officers at the Estacion
Marina at Playa Azul made pleasant what might have been a dreadful stay
in that small coastal village. To the managers and pilots of Lineas
Aereas Picho in Uruapan I owe special thanks for going out of their way
on more than one occasion to transport a stranded snake-hunter.
Throughout the months of field work beginning in 1955 I constantly have
been aided by the authorities and workers of the Comision Tepalcatepec,
a subdivision of the Secretaria de Caminos y Obras Publicas, and of the
private corporation, Ingenieros Civiles Asociados. Much of the field
work in Michoacan was made possible only through the co-operation of the
natives who supplied mules, acted as guides, and aided in the collection
of specimens. I have learned a great deal from these people. They will
never see this report. Their work as guides, muleteers, and collectors
greatly assisted me with the mountains of equipment that had to be piled
on the backs of scrawny mules for transportation to places where the
natives seldom trod. Their efforts in behalf of Don Guillermo never will
be forgotten; I extend an especially hearty _muchas gracias_ to
Benjamin, Ignacio, Jesus, Lorenzo, Mariano, and Remigio.
Much of the work on this report was done while I was associated with the
Museum of Zoology at the University of Michigan. I thank Norman E.
Hartweg and T. H. Hubbell for making available to me the facilities of
the museum and for their numerous courtesies that aided me so much.
My field work in Michoacan was supported by the Museum of Zoology at the
University of Michigan (1951), by the Horace H. Rackham School of
Graduate Studies of the University of Michigan (1955), by the Penrose
Fund of the American Philosophical Society (1956), by the Bache Fund of
the National Academy of Sciences (1958), and by the University of Kansas
Endowment Association (1960).
Permits for collecting specimens in Mexico were provided by the
Direccion General de Caza through the courtesy of Ing. Juan Lozano
Franco and Luis Macias Arellano.
Historical Account
Unlike many parts of southern Mexico and northern Central America,
Michoacan received no attention from the collecting expeditions of the
European museums in the last century. The earliest known herpetological
specimens from Michoacan were obtained by Louis John Xantus, who was
appointed U. S. Consul to Colima in 1859. In April, 1863, Xantus
collected at Volcan Jorullo in Michoacan; in April and May of the same
year he collected along the coast of Michoacan between the Rio Cachan
and the Rio Nexpa. His small collection of 19 extant specimens is in the
United States National Museum. Alfredo Duges, a resident of Guanajuato,
Mexico, made early contributions to the knowledge of the herpetofauna of
Michoacan. In 1885 he described _Sonora michoacanensis_, and in 1891 he
described _Eumeces altamirani_; from what is known of the distribution
of these species, he probably had collected in the Tepalcatepec Valley.
During their biological survey of Mexico, Edward W. Nelson and Edward A.
Goldman spent a limited amount of time in Michoacan in 1892 and again in
1903 and 1904. Most of their collecting was done on the plateau in the
north-central part of the state; their collections are in the United
States National Museum. While collecting fishes in southern Mexico, Seth
E. Meek obtained some amphibians and reptiles from Lago de Patzcuaro in
1904; these are in the collections of the Chicago Natural History
Museum. In 1908 Hans Gadow ventured into the then unexplored "tierra
caliente" of the Balsas Valley and collected at Volcan Jorullo and other
localities in the valley. Later in the same year he collected at
Guayabo, San Salvador, and Arteaga in the Sierra de Coalcoman and at
Buena Vista and Cofradia in the Tepalcatepec Valley. His collections
were deposited in the British Museum (Natural History) and the
Naturhistorisches Museum Wien.
The first thirty years of the present century saw little more field work
in Michoacan. In the 1930's Edward H. Taylor and Hobart M. Smith
collected throughout much of Mexico. At various times they worked in
Michoacan, principally along the road from Mexico City to Guadalajara.
In 1935 Hobart M. Smith spent a week at Hacienda El Sabino south of
Uruapan; he revisited the locality again in 1936 and made a large and
important collection of amphibians and reptiles from the upper limits of
the arid tropical scrub forest in the Tepalcatepec Valley. Specimens
collected by Smith and Taylor were incorporated into the Edward H.
Taylor-Hobart M. Smith collection, which subsequently was deposited in
part in the Museum of Natural History at the University of Illinois and
in part in the Chicago Natural History Museum. In 1939 Hobart M. Smith
collected at Patzcuaro and between Uruapan and Apatzingan; these
collections, made while he was a Walter Rathbone Bacon Scholar of the
Smithsonian Institution, are deposited in the United States National
Museum. In 1940 and 1941 Frederick A. Shannon, who was a member of the
Hoogstraal Expeditions under the auspices of the Chicago Natural History
Museum, collected on Cerro de Tancitaro and at Apatzingan; an account of
the specimens collected there was published by Schmidt and Shannon
(1947).
The eruption of Volcan Paricutin in February, 1943, attracted the
attention of many biologists, a group of which from the Museum of
Zoology at the University of Michigan collected in the Cordillera
Volcanica in 1945 and 1947. The amphibians and reptiles were collected
and studied by Norman E. Hartweg. In 1950 James A. Peters accompanied
Donald D. Brand on a preliminary exploration of the western part of the
Sierra de Coalcoman and adjacent Pacific coast of Michoacan; in the same
year Peters collected also on the Mexican Plateau and at Volcan Jorullo.
His specimens are in the Museum of Zoology at the University of
Michigan. Since 1950 many biologists have collected in Michoacan in the
course of work on certain groups of animals or in general surveys. In
this way Raymond Alcorn, Robert W. Dickerman, James R. Dixon, Floyd L.
Downs, Emmet T. Hooper, and Robert R. Miller have contributed to our
knowledge of the herpetofauna.
As stated previously, my own field work in Michoacan began in 1951, when
I accompanied Donald D. Brand on an exploring expedition to the
southern part of the state. In that year a short time was spent on the
Mexican Plateau, principally in the area around Lago de Cuitzeo, and at
Volcan Jorullo. In July and August we made our headquarters at
Coalcoman. From that town the field party travelled southward to Maruata
on the Pacific coast and thence back over the mountains to Coalcoman.
Later in that summer we travelled by mule from Coalcoman southeastward
to the mouth of the Rio Nexpa. In 1955, accompanied by Lee D. Beatty,
Carter R. Gilbert, and Fred G. Thompson, I collected in the Tepalcatepec
Valley and at Coalcoman. We made a mule trip from Coalcoman to Cerro de
Barolosa, where we made the first collections from the pine-fir forests
in the Sierra de Coalcoman. Later in the same summer Carter R. Gilbert
and I spent a week at Playa Azul on the Pacific coast. In March, April,
and May, 1956, my wife and I collected for a short time in the
Cordillera Volcanica and on the Mexican Plateau. In early April we moved
into the Tepalcatepec Valley, where we collected intensively between
Churumuco and Tepalcatepec. In May we collected on the Pacific coast
between Boca de Apiza and La Placita. In July and August, 1956,
accompanied by Richard E. Etheridge, we returned to Michoacan and again
collected on the Mexican Plateau and in the Cordillera Volcanica, before
moving into the Tepalcatepec Valley. In an attempt to fill in gaps in
the known distributions of many species and to sample the fauna in some
previously uncollected areas, I returned to Michoacan in June, 1958.
Accompanied by Jerome B. Tulecke and John Wellman, I collected on the
Mexican Plateau in the northwestern part of the state, on the southern
slopes of the Cordillera Volcanica, and in the Tepalcatepec Valley. Most
of our time was spent in the Sierra de Coalcoman, where we collected at
Aguililla, Artega, and Dos Aguas. In 1960 two days were spent in
Michoacan; a small collection was made in the eastern part of the
Cordillera Volcanica. With the exception of the specimens collected in
1960, which are at the Museum of Natural History at the University of
Kansas, the specimens that I have collected in Michoacan are in the
Museum of Zoology at the University of Michigan.
NATURAL LANDSCAPE
A proper understanding of the geographical distribution of animals in a
given region is possible only after a thorough acquaintance with the
geography of the region. Likewise, in order to gain a knowledge of the
ecological distribution and relationships of the components of the
fauna, it is necessary to study the animals in their natural
environments. In order to give the reader a picture of the physical
features and the major animal habitats within the state of Michoacan,
the following brief description is offered. Each of these facets
mentioned below will be elaborated in detail in my final report on the
herpetofauna of Michoacan.
Physiography
The state of Michoacan comprises an area of 60,093 square kilometers
(Vivo, 1953). Within this area the rugged terrain has a total relief of
nearly 4000 meters. There have been several attempts to classify the
physiographic provinces of Mexico; the classification used here is a
slight modification of the scheme proposed by Tamayo (1949). I have
tried to keep the system as simple as possible, but still useful in
discussing the distribution of animals living in the region. For general
purposes the state of Michoacan can be divided into lowlands and
highlands as follows:
LOWLANDS
Pacific Coastal Plain
Balsas-Tepalcatepec Basin
HIGHLANDS
Mexican Plateau
Cordillera Volcanica
Sierra de Coalcoman
Although the lowlands in the state are continuous, they are only
narrowly connected and thus form two distinct physiographic and biotic
areas. The Pacific Coastal Plain in Michoacan extends for a distance of
about 200 kilometers (airline) from the Rio Coahuayana to the Rio
Balsas. The coastal plain is broad between the Rio Coahuayana and San
Juan de Lima, and between Las Penas and the Rio Balsas, where the hills
rise some 12 kilometers inland from the sea. Between San Juan de Lima
and Las Penas the mountains extend to the sea; in this region rocky
promontories form precipitous cliffs dropping into the sea. Between the
promontories are small sandy or rocky beaches.
Lying to the north of the Sierra de Coalcoman and the Sierra del Sur,
but south of the Cordillera Volcanica, is a broad structural depression,
the Balsas-Tepalcatepec Basin. The western part of this basin, which
separates the Sierra de Coalcoman from the Cordillera Volcanica, is the
valley of the Rio Tepalcatepec, a major tributary of the Rio Balsas. The
eastern part of the basin is the valley of the Rio Balsas. From the
point of junction of the two rivers, the Rio Balsas flows southward
through a narrow gorge, which separates the Sierra de Coalcoman from
the Sierra del Sur, to the Pacific Ocean. In Michoacan the floor of the
Balsas-Tepalcatepec Basin varies from 200 to 700 meters above sea level.
The central part of Mexico is a vast table-land, the Mexican Plateau,
the southern part of which extends into northern Michoacan. In this
region the terrain is rolling and varies from 1500 to 1900 meters above
sea level. Many small mountain ranges rise from the plateau and break
the continuity of the rolling table-land. Located on the southern part
of the Mexican Plateau in Michoacan are several lakes, the largest of
which are Lago de Chapala, Lago de Cuitzeo, and Lago de Patzcuaro.
Bordering the southern edge of the Mexican Plateau is a nearly unbroken
chain of volcanos, the Cordillera Volcanica. The highest peaks in
Michoacan, Cerro San Andres (3930 meters) and Cerro de Tancitaro (3870
meters), are in this range. Parts of the Cordillera Volcanica in
Michoacan are known by separate names; these are, from west to east:
Sierra de los Tarascos, Sierra de Ozumatlan, and Serrania de Ucareo.
Lying between the Tepalcatepec Valley and the Pacific Ocean, and east of
the Rio Coahuayana and west of the Rio Balsas, is an isolated highland
mass, the Sierra de Coalcoman. This mountain range rises to elevations
of slightly more than 3000 meters. It has a length of about 200
kilometers and a width of about 80 kilometers. Except for a relatively
low connection with the Cordillera Volcanica, the Sierra de Coalcoman is
isolated from other mountain ranges in southwestern Mexico.
CLIMATE
The climates in Michoacan vary from tropical in the lowlands to cool
temperate at high elevations in the Sierra de Coalcoman and Cordillera
Volcanica. The highest temperatures are known in the Balsas-Tepalcatepec
Basin, where at Churumuco the mean annual temperature is 29.3 deg. C. and
the range of monthly means is 3.5 deg. C. (Contreras, 1942). Frosts occur
sporadically on the Mexican Plateau, and in the winter snow falls on the
highest mountains.
Precipitation varies geographically and seasonally. Most of the rain
falls between June and October. In the Balsas-Tepalcatepec Basin
rainfall in the rest of the year is negligible. The annual average
rainfall at Coahuayana on the Pacific Coastal Plain is 871 mm.
(Guzman-Rivas, 1957:52). In the Balsas-Tepalcatepec Basin rainfall
seldom exceeds 800 mm. per year. In the mountains precipitation is
heavier and somewhat more evenly distributed throughout the year, but
still definitely cyclic. For example, Uruapan (elevation, 1500 meters)
receives an average annual rainfall of 1674 mm. (Contreras, 1942). The
prevailing winds are from the Pacific Ocean. The southern (windward)
slopes of the Sierra de Coalcoman probably receive more rain than any
other part of the state. The Balsas-Tepalcatepec Basin lies in a rain
shadow of the Sierra de Coalcoman, and the Mexican Plateau lies in a
somewhat less drastic rain shadow of the Cordillera Volcanica; these are
the driest regions in the state.
VEGETATION AND ANIMAL HABITATS
For the purposes of this report I have adopted the classification of
types of vegetation that seem to me most significant in terms of
ecological distribution of reptiles and amphibians in Michoacan. These
types are as follows:
TEMPERATE (1000-4000 meters)
Fir Forest (2400-4000 meters)
Pine-oak Forest (1000-4000 meters)
Mesquite-grassland (1500-2100 meters)
TROPICAL (0-1000 meters)
Arid Tropical Scrub Forest (0-1000 meters)
Tropical Semi-deciduous Forest (150-600 meters)
The vegetation of the Pacific Coastal Plain and the Balsas-Tepalcatepec
Basin consists of arid tropical scrub forest, composed of deciduous
trees, which in many places are stunted and widely spaced. In the dry
season there is little cover provided by this forest. In the rainy
season there is a sparse growth of grasses and some shade provided by
the small leaves of the thorny trees.
In Michoacan the rainfall is heaviest on the southern slopes of the
Sierra de Coalcoman and somewhat less so on the southwestern slopes of
the Cordillera Volcanica. At these relatively low elevations (150 to 600
meters) there is tropical semi-deciduous forest, characterized by
relatively dense shade throughout the year and by a leaf mulch on the
ground. This type of forest forms the gallery forest along the larger
streams in the Balsas-Tepalcatepec Basin and on the Pacific Coastal
Plain.
Rainfall also is heavy on the high mountain ridges, where temperatures
are low. On these ridges, fir forest, often mixed with pine and oaks, is
found. This habitat is characterized by a cool, moist climate, many
rotting logs, and a moist ground cover of leaves and needles.
Most of the mountains are covered with pine-oak forest, which in most
places is decidedly subhumid, but where this forest occurs on the
windward sides of high ridges, it sometimes is noticeably humid. In
this forest the important animal habitats include the needle- and
leaf-litter, and in some areas, bromeliads.
The rolling terrain of the Mexican Plateau supports cacti, small
leguminous trees, and grasses. Like the arid tropical scrub forest, this
type of vegetation, the Mesquite-grassland association, is deciduous and
thus provides little shelter in the dry season. Unlike the areas in
which arid tropical scrub forest is developed, the Mesquite-grassland is
found in areas having warm days and cool nights.
GEOGRAPHY OF THE HERPETOFAUNA
Although the main part of my final report on the herpetofauna of
Michoacan will deal with the geographical and ecological patterns of
distribution of the herpetofauna, a brief summary of the faunal
assemblages is presented here.
In Michoacan there are two major faunal assemblages, one in the
lowlands, and one in the highlands. A large number of the species
inhabiting the lowlands are wide-ranging species, such as _Bufo
marinus_, _Iguana iguana_, and _Boa constrictor_. Sixty-three species
are known to occur on the Pacific Coastal Plain; 41 of these, together
with 36 others occur in the Balsas-Tepalcatepec Basin, a physiographic
region to which several species of reptiles are endemic; for example,
_Enyaliosaurus clarki_, _Urosaurus gadowi_, _Cnemidophorus calidipes_,
and _Eumeces altamirani_.
Generally speaking, the members of the highland faunal assemblage have
more restricted geographic ranges. The major exceptions are those
species that are widely distributed on the Mexican Plateau, such as:
_Bufo compactilis_, _Sceloporus torquatus_, and _Salvadora bairdi_. In
the montane habitats of the Cordillera Volcanica, 45 species of
amphibians and reptiles are known; 34 species have been found in the
Sierra de Coalcoman. Fourteen species are known to occur in both ranges.
Several species are known only from the Cordillera Volcanica and
adjacent highlands, and three species are endemic to the Sierra de
Coalcoman.
ANNOTATED LIST OF SPECIES
In the following pages the 176 species and subspecies of amphibians and
reptiles known to occur in the state of Michoacan are discussed in
relation to their variation, life histories, ecology, and distribution
in the state. Data have been gathered from 9676 specimens. I have not
prolonged the accounts of species with information that has been
presented elsewhere. Consequently, the length and completeness of the
accounts are variable. I have given only the information that I consider
a worthwhile contribution to our knowledge of the particular species.
The synonymies given at the beginning of each account include the first
use of the trivial name by the original author, the first usage of the
combination that I am using, and, if the circumstances make it
necessary, additional names or combinations that have been proposed
since the publication of the checklists of Mexican amphibians and
reptiles by Smith and Taylor (1945, 1948, and 1950b). References cited
only in the synonymies are not listed in the Literature Cited. Preceding
the discussion of each species is an alphabetical list of the localities
in Michoacan from which specimens have been examined. The listing of a
locality means that one or more specimens, as indicated, has been
examined from that locality. Only for those specimens especially
mentioned in the text are catalogue numbers given. Abbreviations for the
various museums and scientific collections are, as follows:
AMNH American Museum of Natural History
ANSP Academy of Natural Sciences of Philadelphia
BMNH British Museum (Natural History)
CNHM Chicago Natural History Museum
EHT-HMS Edward H. Taylor-Hobart M. Smith collection
JRD James R. Dixon collection, College Station, Texas
KU University of Kansas Museum of Natural History
MCZ Museum of Comparative Zoology
MVZ Museum of Vertebrate Zoology
NMW Naturhistorisches Museum Wien
SU Stanford University Museum of Natural History
TCWC Texas Cooperative Wildlife Collection
UIMNH University of Illinois Museum of Natural History
UMMZ University of Michigan Museum of Zoology
USNM United States National Museum
UTNHC University of Texas Natural History Collection
Throughout the accounts of the species all measurements are given in
millimeters; if the range of variation is given, the mean follows in
parentheses.
AMPHIBIA
Caudata
~Ambystoma amblycephalum~ Taylor
_Ambystoma amblycephala_ Taylor, Univ. Kansas Sci. Bull.,
26: 420, November 27, 1940.--Fifteen kilometers west of
Morelia, Michoacan, Mexico.
Fifteen km. W of Morelia (19); 11 km. SSE of Opopeo (12); 8
km. S of Patzcuaro; 24 km. S of Patzcuaro (2); Quiroga (20);
Tacicuaro (167).
Taylor and Smith (1945:530) presented data on 137 specimens collected at
Tacicuaro on October 1, 1939; these are all larvae and metamorphosing
individuals. Aside from these, the largest larva examined (UMMZ 104962
from 15 km. W of Morelia) has a snout-vent length of 70.0 mm. and a tail
length of 53.5 mm. The larvae are pale pinkish tan above and somewhat
paler below; there is a lateral row of cream colored spots. The
tail-fin, which is deepest at mid-length, extends to the back of the
head and is flecked with brown. In small larvae the outer edge of the
tail-fin is dark brown. The eyes are large. Two small metamorphosed
specimens (UMMZ 98967) from 24 kilometers south of Patzcuaro are
tentatively referred to this species. These specimens have body lengths
of 49.0 and 45.0 mm. and tail lengths of 36.0 and 31.5 mm.,
respectively. They have 17-17 and 16-15 vomerine teeth arranged in a
broad arch behind the choanae, 10 costal grooves, and 7 intercostal
spaces between adpressed toes. The dorsal color is uniform brown; that
of the venter is a dusty cream.
Larvae were collected from shallow ponds near Quiroga and 15 kilometers
west of Morelia; metamorphosed individuals were taken from beneath logs
in pine and fir forests at elevations from 2300 to 2800 meters.
~Ambystoma dumerili dumerili~ (Duges)
_Siredon Dumerili_ Duges, La Naturaleza, 1:241, 1870--Lago
de Patzcuaro, Michoacan, Mexico.
_Bathysiredon dumerilii_, Dunn, Notulae Naturae, 36:1,
November 9, 1939.
_Bathysiredon dumerilii dumerilii_, Maldonado-Koerdell, Mem.
y Rev. Acad. Nac. Cien., 56:199, 1948.
_Ambystoma_ (_Bathysiredon_) _dumerili_, Tihen, Bull.
Florida State Mus., 3:3, June 20, 1958.
Lago de Patzcuaro (22);? Morelia.
For many years this unusual salamander was known from only a few
specimens mostly collected in the last century; Smith and Taylor
(1948:7) stated: "It is presumed that this species is extinct owing to
the introduction of exotic game and food fishes." In 1951 and in 1955 I
had been told that _axolotls_ were sold in the market at Patzcuaro;
nevertheless, none was found on my visits there. In 1956 Charles M.
Bogert obtained several large specimens at the market in Patzcuaro.
These establish the continued existence of the salamander in Lago de
Patzcuaro. On January 27, 1955, R. W. Dickerman procured a specimen (KU
41573) in the market at Morelia. Since fish are brought to Morelia from
Lago de Patzcuaro, the specimen probably was from that lake.
Nevertheless, the species may occur in other permanent bodies of water
in Michoacan. Maldonado-Koerdell (1948) described _Bathysiredon dumerili
queretarensis_ from San Juan del Rio, Queretaro. This locality is about
200 airline kilometers northeast of Lago de Patzcuaro and is in the Rio
Moctezuma drainage.
~Ambystoma ordinarium~ Taylor
_Ambystoma ordinaria_ Taylor, Univ. Kansas Sci. Bull.,
26:422, November 27, 1940.--Four miles west of El Mirador,
near Puerto Hondo, Michoacan, Mexico.
Axolotl (56); Cerro San Andres; 22 km. W of Mil Cumbres; 46
km. E of Morelia (34); 8 km. SE of Opopeo (5); Puerto de
Garnica (8); Puerto Hondo (41); San Gregorio (16); San Jose
de la Cumbre (20).
Of 16 specimens (KU 51520-35) collected on June 18, 1955, near San
Gregorio, 15 are adult females with swollen cloacae and minute ovarian
eggs. Possibly these specimens had just recently deposited their mature
eggs. In preservative the specimens are black above and dull creamy gray
below. Measurements for the 15 females are: snout-vent length,
80.0-102.0 (92.5); tail length, 69.0-93.0 (84.2); head width, 15.8-20.5
(17.7); head length, 22.8-26.6 (24.4). A larval specimen with small
gills has a snout-vent length of 72 mm. and a tail length of 62 mm.
Three specimens have 12 costal grooves; the other have 11.
Of 20 specimens from San Jose de la Cumbre (UMMZ 112857 and 115143), 14
are neotenic adults; the others are larvae. In life the salamanders were
blackish to olive-brown above with scattered cream-colored dots on the
dorsum and flanks but in preservative are dull grayish black with
indistinct pale spots and dark reticulations. The belly is pale gray
with indistinct dark spots. Eleven females and three males have the
following measurements, respectively: snout-vent length, 76.0-90.0
(80.7), 64.0-84.0 (74.3); tail length, 70.0-81.0 (75.0), 58.0-71.0
(66.7); head width, 19.5-23.5 (20.7), 17.5-20.5 (19.3); head length,
22.0-25.0 (23.0), 20.0-22.5 (21.5). The smallest larva has a snout-vent
length of 43.0 mm. and a tail length of 38.0 mm. Two individuals have 12
costal grooves; the others have 11. All of the females contained eggs,
the largest of which were 1.5 mm. in diameter. The stomachs of most of
the specimens were distended with oligochaets, aquatic insect larvae,
and small aquatic beetles.
A series of 34 larvae (JRD 5904-37) from 46 kilometers east of Morelia
are tentatively referred to this species. These specimens are
olive-brown above with cream-colored spots on the flanks; the dorsal
tail-fin does not extend onto the body.
This species has been found only at elevations in excess of 2400 meters
in pine and fir forests. At Rancho Axolotl James A. Peters collected
larvae and neotenic individuals in a rocky stream and adults from
beneath rocks and logs in the forest near the stream. Neotenic
individuals and larvae were found in a clear stream in pine-fir forest
at an elevation of 2700 meters near San Jose de la Cumbre; specimens
were collected there in July, 1955, and again in July, 1956. The site
was visited in April, 1956, at which time the stream consisted of only a
few puddles; no salamanders were found.
~Ambystoma tigrinum velasci~ Duges
_Ambystoma velasci_ Duges, La Naturaleza, ser. 2, 1:142,
1888.--Laguna Santa Isabel, near Guadalupe Hidalgo, Distrito
Federal, Mexico.
_Ambystoma tigrinum velasci_, Dunn, Copeia, no. 3:157,
November 14, 1940.
Patzcuaro (5); Tacicuaro (9).
Definite specific assignment of these specimens, all larvae, cannot be
made at this time. They have shovel-shaped heads and laterally
compressed bodies with the dorsal tail-fin extending anteriorly to the
back of the head. The eyes are small. The body is pale tan with dark
mottling on the tail and flanks. The average snout-vent length for nine
specimens from Tacicuaro is 61.0 mm.
The larvae from Tacicuaro (UMMZ 89255) were collected by Dyfrig Forbes
in October, 1939; those from Patzcuaro, presumably Lago de Patzcuaro
(BMNH 1914.1.28-247-8 and CNHM 948), were collected by Hans Gadow and
Seth Meek in 1908.
~Pseudoeurycea belli~ (Gray)
_Spelerpes belli_ Gray, Catalogue Batrachia Gradientia
British Museum, p. 46, 1850.--Mexico. Type locality
restricted to 2 miles east of Rio Frio, Puebla, Mexico, by
Smith and Taylor (1950a:341).
_Pseudoeurycea bellii_, Taylor, Univ. Kansas Sci. Bull.,
30:209, June 12, 1944.
Axolotl (2); Carapan; Cerro Tancitaro (84); Macho de Agua;
22 km. W of Mil Cumbres; Opopeo; Patzcuaro (8); Puerto Hondo
(2): San Jose de la Cumbre; San Juan de Parangaricutiro
(42); Uruapan (5); Zacapu (4).
This salamander seems to reach its greatest abundance in Michoacan in
the Sierra de los Tarascos between Patzcuaro and Tancitaro, where it is
found at elevations from 1500 to 2900 meters. It is found less commonly
in the eastern part of the Cordillera Volcanica in Michoacan, where it
sometimes occurs in association with _Pseudoeurycea robertsi_.
On June 22 and 23, 1955, four clutches of eggs of this species were
found beneath adobe bricks and rocks on the volcanic ash that has buried
the village of San Juan de Parangaricutiro. The eggs were unstalked and
separate, but adherent in clumps of three or four (Pl. 2, Fig. 1). The
outer membranes were covered with fine particles of ash. The ash beneath
the stones where the eggs were found was only slightly moist; one clump
of eggs was partially desiccated. Three complete clutches have 20, 23,
and 34 eggs; one clutch of 15 eggs was being eaten by beetles
(Tenebrionidae: _Eleodes_ sp.). The eggs vary in size from 4.6 to 6.5
mm. and average 5.3 mm. in diameter. They are unpigmented. Surrounding
the embryo is a vitelline membrane, an inner, and an outer envelope
(Fig. 1). In an average-sized egg having an embryo 4 mm. in length, the
diameter of the outer membrane is 5.3 mm., the inner membrane 5.0 mm.,
and the vitelline membrane 4.6 mm. All of the eggs contained embryos in
which the limb buds were developed; in about half of these the eyes were
distinctly visible.
[Illustration: Fig. 1. Diagram of an egg of _Pseudoeurycea belli_ from
San Juan de Parangaricutiro, Michoacan. x 10.]
The first heavy rain of the season occurred on the night of June 22,
1955. Thus, at least sometimes, _Pseudoeurycea belli_ lays its eggs
before the onset of the rainy season. A female having a snout-vent
length of 110 mm., collected on June 22, 1955, contained 36 ovarian eggs
having diameters from 3.0 to 3.5 mm. The fact that small juveniles were
collected on the same date indicates that this salamander lays eggs over
a period of several weeks in late spring and early summer.
The smallest juvenile examined has a snout-vent length of 17.0 mm. and a
tail length of 7.5 mm. Twelve juveniles from the vicinity of San Juan de
Parangaricutiro have an average snout-vent length of 19.4 mm. and an
average tail length of 9.7 mm. In juveniles the adpressed limbs either
touch or overlap by one intercostal space; in adults there are two or
three intercostal spaces between adpressed toes. Therefore the greatest
number of intercostal spaces between adpressed limbs is found in the
largest specimens. A similar relationship between adpressed limbs (=
length of limbs) and snout-vent length was shown for _Plethodon
richmondi_ by Duellman (1954a). The number of vomerine teeth is
variable; the number of teeth seems to be closely correlated with the
size of the salamander (Fig. 2). A similar correlation between the
number of maxillary teeth and body length was reported for
_Chiropterotriton multidentatus_ by Rabb (1958). In 12 juvenile
_Pseudoeurycea belli_ there are 6-13 (8.8) vomerine teeth, and in 11
adults having snout-vent lengths greater than 90 mm. there are 39-49
(44.0) vomerine teeth. The coloration of the juveniles resembles that of
the adults (Pl. 1).
[Illustration: FIG. 2. Correlation between the number of vomerine teeth
and snout-vent length in 79 _Pseudoeurycea belli_ from Michoacan.]
The differences between this species and _Pseudoeurycea gigantea_ are
minor. Taylor (1939a) distinguished _gigantea_ from _belli_ by the
larger size, fewer intercostal spaces between adpressed limbs, more
vomerine teeth, and absence of occipital spots in _gigantea_. Taylor and
Smith (1945) stated that in life the spots in _gigantea_ are orange
instead of red as in _belli_. Five specimens of _Pseudoeurycea belli_
from Michoacan, including one juvenile, lack occipital spots. In the 34
living individuals that I have seen from Michoacan the spots varied from
deep red to orange. Therefore, of the characters listed by Taylor (_op.
cit._) to diagnose _Pseudoeurycea gigantea_, only the over-all larger
size and smaller number of intercostal spaces between adpressed limbs (=
relatively longer limbs) are useful in separating _Pseudoeurycea belli_
and _gigantea_.
~Pseudoeurycea robertsi~ (Taylor)
_Oedipus robertsi_ Taylor, Univ. Kansas Sci. Bull., 25:287,
July 10, 1939.--Nevado de Toluca, Mexico.
_Pseudoeurycea robertsi_ Taylor, Univ. Kansas Sci. Bull.,
30:209, June 12, 1944.
Atzimba (3); Macho de Agua (9); Puerto Lengua de Vaca (14).
Previously this species has been recorded only from the type locality.
In July, 1956, individuals referable to this species were found at two
sites in pine-fir forest immediately to the east of Macho de Agua and in
pine-oak-fir forest at Atzimba. On August 20, 1958, a series was
collected in pine-fir forest at Puerto Lengua de Vaca. These localities
are between 2900 and 3000 meters in the Cordillera Volcanica in eastern
Michoacan.
In life the coloration of these salamanders was highly variable. The
belly and undersurfaces of the tail and hind limbs were pale gray, with
or without silvery white flecks; the chin was a cream-color and flecked
with silvery white in some specimens. The middorsal area was brown,
orange-brown, or dull grayish yellow. The flanks and lateral surfaces of
the tail were black with yellowish flecks or streaks on the flanks and
yellowish or orange-brown flecks on the tail. The iris was golden brown.
Measurements of eight males and two females are, respectively:
snout-vent length, 42.5-56.0 (49.5), 54.0-60.0 (57.0); tail length,
42.0-56.0 (48.1), 52.0-55.0 (53.5). The smallest juvenile has a
snout-vent length of 28.0 mm. and a tail length of 23.0 mm. Of the 26
available specimens, six have 12 costal grooves, and the others have 11.
In comparison with 36 topotypes, the specimens from Michoacan have a
less striking dorsal color pattern; none has a well-defined dorsal
reddish brown area or bold reddish mottling on the tail. Furthermore,
the specimens from Michoacan have paler venters than do topotypic
specimens.
Salientia
~Rhinophrynus dorsalis~ Dumeril and Bibron
_Rhinophrynus dorsalis_ Dumeril and Bibron, Erpetologie
generale, vol. 8:758, 1841.--Veracruz, Veracruz, Mexico.
Mouth of the Rio Balsas (10).
These specimens (BMNH 1914.1.28.181-90) were collected by Gadow in 1908
and reported by him (1930:72): "Whilst this very sluggish termite-eating
toad is common enough in the sweltering hot country of the state of Vera
Cruz, up to an elevation of 1500 feet, it was unknown on the west side
of the Isthmus until I found it in great numbers near the mouth of the
Balsas River, in and near fresh-water pools, where it attracted
attention by its loud peculiar voice during the pairing season in the
month of July." Subsequently, Peters (1954:3) verified the
identification of these specimens. Although torrential rains fell during
the week in July, 1955, that I spent at Playa Azul near the mouth of the
Rio Balsas, the distinctive voice of _Rhinophrynus_ was not heard.
Elsewhere on the Pacific coast of Mexico adult _Rhinophrynus_ have been
reported only from Tehuantepec and a few localities on the coastal
lowlands of Chiapas. Taylor (1942b:37) found on the coast of Guerrero a
tadpole that was referred to the genus _Rhinophrynus_ by Orton (1943).
In the summer of 1960 adults of _Rhinophrynus_ were collected near
Acapulco, Guerrero (Fouquette, _in litt._). These recent collections
verify the existence of the species along the Pacific lowlands of Mexico
at least as far north as Michoacan.
~Scaphiopus hammondi multiplicatus~ Cope
_Scaphiopus multiplicatus_ Cope, Proc. Acad. Nat. Sci.
Philadelphia, 15:52, June 8, 1863.--Valley of Mexico.
_Scaphiopus hammondi multiplicatus_, Kellogg, Bull. U. S.
Natl. Mus., 160:22, March 31, 1932.
Angahuan (5); Cuitzeo (4); Cuseno Station (2); Jiquilpan
(9); Morelia (7); Patzcuaro (3); Quiroga; Tarecuaro; Uruapan
(24); Zacapu.
This small toad has been found at elevations between 1500 and 2500
meters on the Mexican Plateau and associated mountain ranges; it occurs
in mesquite-grassland and in pine forests. Calling males and females
laden with eggs have been collected in the rainy season in the months of
July and August. The call is a medium-pitched snore. In living
individuals the dorsal ground color varies from pale brown to gray with
dark brown or olive-brown markings. In many individuals the tips of the
small dorsal pustules are red.
~Bufo coccifer~ Cope
_Bufo coccifer_ Cope, Proc. Acad. Nat. Sci. Philadelphia,
18:130, 1866--Arriba, Costa Rica.
Apatzingan (27); Lombardia; Nueva Italia (5).
In life the dorsal color pattern consists of a yellowish tan ground
color with dark brown spots; the middorsal stripe is deep yellow or
cream color. The venter is a dusty cream color, and the iris is pale
gold. Males have dark brown horny nuptial tuberosities on the thumb. The
following measurements are of 21 males and four females, respectively:
snout-vent length, 43.5-51.7 (48.1), 55.6-62.6 (59.1); tibia length,
16.6-18.8 (17.6), 18.8-20.3 (19.3); head width, 16.7-19.7 (18.4),
20.6-22.2 (21.4); head length, 13.8-16.6 (14.8), 16.5-18.2 (17.3).
The specimens from the Tepalcatepec Valley differ slightly from
specimens from southeastern Mexico and Central America. Those from
Michoacan have low and narrow cranial crests; in about one-half of the
specimens the occipital crest exists only as a row of tubercles, and in
some the postorbital and suborbital crests are barely discernible.
Specimens from the southern part of the range, Costa Rica and Nicaragua,
have much higher and thicker cranial crests; in these the occipital
crest is well defined and extends posteriorly to a point back of the
anterior edge of the parotid gland; the postorbital and suborbital
crests are well marked. Of 48 specimens from Esquipulas, Guatemala, all
have high crests, but these are not so well developed as in ten
specimens from Matagalpa, Nicaragua, and three from various localities
in Costa Rica. Six specimens from Tehuantepec, Oaxaca, have cranial
crests that are lower than those in specimens from Guatemala. In three
of the specimens from Tehuantepec the occipital crests are reduced to a
series of tubercles. Of six specimens from Agua del Obispo, Guerrero,
four have poorly developed occipital crests. These observations suggest
the presence of a cline in the development of the cranial crests;
specimens have higher crests in the southern part of the range than in
the northern part.
In Mexico _Bufo coccifer_ has been collected only in semi-xeric
habitats, but to the south, from Guatemala to Costa Rica, it has been
found in more upland and humid habitats. Southern specimens are darker
than those from the north, a possible correlation with the differences
in habitat.
These toads probably range throughout the Tepalcatepec Valley, but they
are unknown from the coast of Michoacan. Breeding choruses were found
after heavy rains on June 24, 1955, and on August 2, 1956. The first was
in a muddy ditch; the second was in a flooded grassy field. The call is
a high-pitched, but not loud, "whirrr." Males were calling from the edge
of the water or from clumps of grass in the water. Clasping pairs were
in the water; amplexus is axillary.
~Bufo compactilis compactilis~ Wiegmann
_Bufo compactilis_ Wiegmann, Isis von Oken, 26:661,
1833.--Mexico. Type locality restricted to Xochimilco,
Distrito Federal, Mexico, by Smith and Taylor (1950a:330).
_Bufo compactilis compactilis_, Smith, Herpetologica, 4:7,
September 17, 1947.
Cuitzeo (2); Emiliano Zapata (20); Jiquilpan (5); La Palma
(5); Morelia; Tupataro.
The southwestern terminus of the range of this species is on the Mexican
Plateau in Michoacan. All specimens from the state have spotted venters.
In living toads the dorsal ground color was gray or grayish tan with
olive green spots. The vocal sac was brownish gray; the iris was a
bright golden color.
On June 11, 1958, many individuals were calling from shallow water in a
flooded field at Emiliano Zapata. The call is a slow trill, in which the
individual notes are discernible.
~Bufo marinus~ (Linnaeus)
_Rana marina_ Linnaeus, Systema naturae, ed. 10, 1:211,
1758.--America.
_Bufo horribilis_ Wiegmann, Isis von Oken, 26:654,
1833.--Misantla and Veracruz, Veracruz, Mexico. Taylor and
Smith, Proc. U. S. Natl. Mus., 95:551, January 30, 1945.
_Bufo angustipes_ Taylor and Smith, Proc. U. S. Natl. Mus.,
95:553, January 30, 1945.--La Esperanza, Chiapas, Mexico.
Aguililla; Apatzingan (3); Barranca de Bejuco; Capirio;
Charapendo; Chichihuas; Coahuayana (2); Coalcoman (7);
Cofradia (2); 25 km. S of Cuatro Caminos; El Sabino (10);
Huahua, La Playa (13); Ojos de Agua de San Telmo; Ostula;
Playa Azul (2); Pomaro (2).
This large toad is characteristically found in areas supporting tropical
scrub forest to elevations of about 1000 meters. The species is much
more abundant than the numbers listed above suggest. In the dry season
individuals have been observed in patios, along streams, and by
irrigation ditches. In the rainy season the loud, rattling call of the
males is heard at night throughout the Tepalcatepec Valley and the
coastal lowlands.
Taylor and Smith (1945:552) revived Wiegmann's _Bufo horribilis_ for the
large toads of Mexico that are here referred to _B. marinus_. Their
action was based upon the supposition that the "species _marinus_" is
composite. Although probably true, this supposition has yet to be
proved. Until the large, and apparently related, species of _Bufo_
inhabiting tropical America have been studied systematically as a unit,
the recognition of segments of the population as either species or
subspecies is meaningless. Taylor and Smith (op. cit.:553) based the
description of a new species, _Bufo angustipes_, on one rather
emaciated, formalin-hardened female from La Esperanza, Chiapas. The type
(USNM 116513), when compared with numerous specimens of _Bufo marinus_
from throughout the range of the species in Mexico and northern Central
America, displays no combination of characters to set it off from the
others. Therefore, I suggest that _Bufo horribilis_ Wiegmann and _Bufo
angustipes_ Taylor and Smith be placed in the synonymy of _Bufo marinus_
(Linnaeus) until future systematic study of the genus and this species
in particular establishes the existence of recognizable taxa.
~Bufo marmoreus~ Wiegmann
_Bufo marmoreus_ Wiegmann, Isis von Oken, 26:66,
1833.--Veracruz, Veracruz, Mexico.
Barranca de Bejuco; Coahuayana (11); El Diezmo (2); La
Placita (9); La Orilla (12); Motin del Oro; Ostula (9);
Playa Azul (5); Pomaro (15); Salitre de Estopilas; San Pedro
Naranjestila.
In Michoacan this species is confined to elevations of less than 1000
meters on the coast and foothills of the Sierra de Coalcoman. In this
region in the months of June and July, breeding congregations have been
found in temporary pools and along streams.
Smith and Taylor (1948:39), in their key to the Mexican species of
_Bufo_, placed emphasis on the nature of the supraorbital and
postorbital crests (whether they form a curve or a sharp angle) in
distinguishing _Bufo marmoreus_ from _Bufo perplexus_. In the original
description of _perplexus_, Taylor (1943a:347) characterized the species
as follows: supraorbital and postorbital crests forming a sharp angle,
instead of a curve as in _marmoreus_; supratympanic crest smaller than
in _marmoreus_; diagonal lateral stripe lacking in females;
concentration of dorsal tubercles as found in _marmoreus_ lacking in
males. The discovery of specimens in which the crests form a curve and
others in which the crests form an angle in both the Tepalcatepec Valley
and in the coastal lowlands prompted an investigation of these
characters and others throughout the ranges of the species. An
examination of 410 specimens has resulted in the following conclusions.
TABLE 1.--VARIATION IN THE SHAPE OF THE SUPRAORBITAL AND POSTORBITAL
CRANIAL CRESTS IN BUFO MARMOREUS AND B. PERPLEXUS.
+------------------------+---+-----------+------------+----------+
|LOCALITY | N | Curved |Intermediate| Angular |
+------------------------+---+-----------+------------+----------+
|Tepalcatepec Valley | 50| 10 (20.0%)| 17 (34.0%) |23 (46.0%)|
+------------------------+---+-----------+------------+----------+
|Morelos | 12| 2 (16.6%)| 5 (41.7%) | 5 (41.7%)|
+------------------------+---+-----------+------------+----------+
|Izucar, Puebla | 4| 2 (50.0%)| 0 (0.0%) | 2 (50.0%)|
+------------------------+---+-----------+------------+----------+
|Southern Sinaloa | 1| 1(100.0%)| 0 (0.0%) | 0 (0.0%)|
+------------------------+---+-----------+------------+----------+
|Puerto Vallarta, Jalisco| 2| 2(100.0%)| 0 (0.0%) | 0 (0.0%)|
+------------------------+---+-----------+------------+----------+
|Colima | 45| 25 (55.0%)| 18 (40.0%) | 2 (5.0%)|
+------------------------+---+-----------+------------+----------+
|Coast of Michoacan | 55| 35 (63.6%)| 17 (30.9%) | 3 (5.5%)|
+------------------------+---+-----------+------------+----------+
|Acapulco, Guerrero | 7| 7(100.0%)| 0 (0.0%) | 0 (0.0%)|
+------------------------+---+-----------+------------+----------+
|Chilpancingo, Guerrero | 10| 1 (10.0%)| 4 (40.0%) | 5 (50.0%)|
+------------------------+---+-----------+------------+----------+
|Pochutla, Oaxaca | 13| 6 (46.2%)| 6 (46.2%) | 1 (7.6%)|
+------------------------+---+-----------+------------+----------+
|Tehuantepec, Oaxaca |177| 81 (45.8%)| 67 (37.8%) |29 (16.4%)|
+------------------------+---+-----------+------------+----------+
|Tonola, Chiapas | 1| 0 (0.0%)| 0 (0.0%) | 1(100.0%)|
+------------------------+---+-----------+------------+----------+
|Veracruz | 33| 26 (78.8%)| 6 (18.2%) | 1 (3.0%)|
+------------------------+---+-----------+------------+----------+
| Total |410|198 (48.3%)|140 (34.2%) |72 (17.5%)|
+------------------------+---+-----------+------------+----------+
1. Although the highest percentage of individuals having the
supraorbital and postorbital crests forming a sharp angle is
from localities in the Balsas-Tepalcatepec Basin, numerous
individuals from throughout the range of _marmoreus_ have the
crests forming an angle (Table 1).
2. In all samples of ten or more specimens, some toads have
the supraorbital and postorbital crests forming a sharp
angle, some have the crests forming a curve, and some have
an intermediate condition.
3. The relative size of the supratympanic crest is highly
variable in all samples examined.
[Illustration: FIG. 3. Adult male of _Bufo perplexus_ from Apatzingan,
Michoacan. x 1.5.]
[Illustration: FIG. 4. Adult male of _Bufo marmoreus_ from Pomaro,
Michoacan. x 1.5.]
[Illustration: PLATE 1
Hatchling of _Pseudoeurycea belli_ from San Juan de Parangaricutiro,
Michoacan. x 8.]
[Illustration: PLATE 2
FIG. 1. Nest and eggs of _Pseudoeurycea belli_ beneath a
rock at San Juan de Parangaricutiro. Approx. natural size.
FIG. 2. Multiple egg clutches of _Phyllomedusa
dacnicolor_ from Coalcoman, Michoacan. 1/3 x.
[Illustration: PLATE 3
FIG. 1. Adult male of _Tomodactylus angustidigitorum_
from Paracho, Michoacan. x 4.
FIG. 2. Adult male of _Tomodactylus fuscus_ from Los
Cantiles, Michoacan. x4.
[Illustration: PLATE 4
FIG. 1. Adult male of _Tomodactylus nitidus_ nitidus from
Tuxpan, Michoacan. x.
FIG. 2. Adult male of _Tomodactylus nitidus orarius_ from
Tecolapa, Colima. x 4.]
[Illustration: PLATE 5
FIG. 1. Adult male of _Tomodactylus nitidus_ petersi from
Apatzingan, Michoacan. x 4.
Fig. 2. Adult male of _Tomodactylus rufescens_ from Dos
Aguas, Michoacan. x 4.]
[Illustration: PLATE 6
FIG. 1. Adult male of _Hypopachus caprimimus_ from
Tuxpan, Michoacan. x 2-1/2.
FIG. 2. Adult male of _Hypopachus oxyrrhinus ovis_ from
Tangamandapio, Michoacan. x 3.]
4. A distinct, pale-colored, diagonal lateral stripe is
found in females only from localities outside of the
Balsas-Tepalcatepec Basin; females from the basin have a
spotted dorsum.
5. Males from the Balsas-Tepalcatepec Basin usually have a
broad middorsal line that is yellow or pale tan; those from
outside the basin have either a narrow middorsal line or
none.
6. Males from the Balsas-Tepalcatepec Basin have low,
scattered dorsal tubercles (Fig. 3); males from outside the
basin have a concentration of tubercles in a broad band on
the back (Fig. 4).
Therefore the nature of the cranial crests is of little value in
separating two populations, but the color pattern of the females and the
nature of the dorsal tubercles of the males do show distinct
differences. Furthermore, certain differences in size and proportion are
evident; _Bufo marmoreus_ is a slightly larger toad and has a relatively
longer tibia and longer head than _perplexus_ (Table 2).
TABLE 2.--COMPARISON OF CERTAIN MEASUREMENTS AND PROPORTIONS IN BUFO
MARMOREUS AND B. PERPLEXUS. (MEANS ARE GIVEN IN PARENTHESES BELOW THE
RANGES.)
+-------------------+------------+--+----------+------------+-----------+
| | | | |Tibia length|Head length|
| Species | Sex | N|Snout-vent+------------+-----------+
| | | | length | Snout-vent |Snout-vent |
| | | | | length | length |
+-------------------+------------+--+----------+------------+-----------+
| _B. marmoreus_ | Male |15|61.5-72.5 | 35.9-41.6 | 28.3-33.3 |
| | | | (65.2) | (39.0) | (31.6) |
+-------------------+------------+--+----------+------------+-----------+
| _B. perplexus_ | Male |20|50.0-59.0 | 33.7-38.1 | 26.4-31.1 |
| | | | (54.9) | (36.4) | (29.5) |
+-------------------+------------+--+----------+------------+-----------+
| _B. marmoreus_ | Female | 7|68.0-76.0 | 33.0-36.8 | 26.8-32.6 |
| | | | (70.7) | (34.7) | (29.6) |
+-------------------+------------+--+----------+------------+-----------+
| _B. perplexus_ |Female | 6|64.1-69.8 | 32.4-36.9 | 25.1-29.0 |
| | | | (66.8) | (35.5) | (27.5) |
+-------------------+------------+--+----------+------------+-----------+
Taylor (1943a:347) described _Bufo perplexus_ from Mexcala on the Rio
Balsas in Guerrero. Among the many paratypes are specimens from Tonola,
Chiapas, and Tehuantepec, Oaxaca. These apparently were referred to
_perplexus_ solely on the nature of the cranial crests. All of the
specimens examined during the course of the present study from the
lowlands of Veracruz and from the Pacific lowlands from Sinaloa
southward to Chiapas are referable to _Bufo marmoreus_; those from the
Balsas-Tepalcatepec Basin are referable to _Bufo perplexus_, as defined
above. Ten specimens from Chilpancingo, Guerrero (UMMZ 115352), do not
readily fit either species. Perhaps there is gene exchange between the
inland and coastal populations through the relatively low pass at
Chilpancingo, at the mouth of the Rio Balsas, and near the convergent
headwaters of the Rio Coahuayana and Rio Tepalcatepec in southern
Jalisco. If this can be demonstrated, then _Bufo perplexus_ would have
to be considered as a subspecies of _Bufo marmoreus_, instead of an
allopatric species.
~Bufo perplexus~ Taylor
_Bufo perplexus_ Taylor, Univ. Kansas Sci. Bull., 29:347,
October 15, 1943.--Balsas River near Mexcala, Guerrero,
Mexico.
Aguililla (2); Apatzingan (42); Buena Vista (5); Capirio
(3); La Playa (25); Lombardia (6); Nueva Italia (9); Rio
Cancita, 14 km. E of Apatzingan; Rio Tepalcatepec, 27 km. S
of Apatzingan; San Salvador (4); Tzitzio; Volcan Jorullo.
~Bufo occidentalis~ Camerano
_Bufo occidentalis_ Camerano, Atti R. Accad. Sci. Torino,
14:887, December 31, 1878.--Mexico. Type locality restricted
to Guanajuato, Guanajuato, Mexico, by Smith and Taylor
(1950a:330). Firschein, Copeia, no. 3:220, September 15,
1950.
_Bufo simus_, Smith and Taylor, Bull. U. S. Natl. Mus.,
194:42, 1948.
Barranca Seca (32); Cerro de Barolosa (4); Cerro Tancitaro,
3 km. E of Apo (2); Cerro Tancitaro, 19 km. E. of Apo (10);
Charapendo; Coalcoman (7); Dos Aguas (4); Jacona, Jaramillo
(2); Las Tecatas; Los Reyes (181); Tancitaro (10); Uruapan
(3).
This toad is an inhabitant of pine and oak forests between 900 and 2400
meters. Near Charapendo on the slopes of the Sierra de los Tarascos and
at Coalcoman it apparently reaches its lowest altitudinal limits. At
both of these localities the pine-oak forest is replaced by arid
tropical scrub forest on the lower slopes.
Twenty-four tadpoles were collected on May 3 in a quiet section of a
fast stream near Barranca Seca. The tadpoles have a robust body,
broadest about two-thirds the distance from the snout to the posterior
edge of the body, half again as broad as deep. Eyes dorsolateral;
nostrils dorsal, somewhat directed forward, and about three-fifths the
distance from the tip of the snout to the eye; spiracle sinistral and
lateral, located at about midbody; anus median; tail long and slender;
tail-musculature extends nearly to tip of tail; depth of
tail-musculature at mid-length about one-third total depth of tail;
dorsal tail-fin not extending onto body (Fig. 5); average body length of
ten tadpoles having small hind limb buds, 14.4 mm.; average tail length,
22.0 mm.
Mouth ventral, nearly terminal, about one-third as wide as widest part
of body; anterior lip has no papillae; lower lip bordered by two rows of
papillae and lateral lips by one row of papillae; beaks moderately well
developed, the upper forming a broad arch and finely denticulate; tooth
rows 2/3, the upper rows extending to the edge of the lips, subequal in
length, and slightly longer than lower rows, which also are subequal in
length; inner upper tooth row broken medially; inner lower tooth row
sometimes broken (Fig. 6).
The body is black dorsally and laterally, and bluish gray ventrally; the
tail musculature is brown and stippled with darker brown. The fins are
transparent and stippled with brown, the stippling being most pronounced
on the posterior two-thirds of the upper tail-fin.
[Illustration: FIG. 5. Tadpole of _Bufo occidentalis_ (UMMZ 94269) from
Barranca Seca, Michoacan. x 3.]
[Illustration: FIG. 6. Mouthparts of larval _Bufo occidentalis_ (UMMZ
94269) from Barranca Seca, Michoacan. x 20.]
Forty recently metamorphosed individuals average 18.9 mm. in snout-vent
length.
The relationships of this toad seem to be with _Bufo bocourti_ Brocchi,
an inhabitant of pine and oak forests in the uplands of Chiapas and
Guatemala. In _Bufo occidentalis_ the tympanum usually is indistinct and
sometimes completely covered, and it is absent in _bocourti_. _Bufo
occidentalis_ has a broader interorbital area and relatively shorter and
more rounded parotid glands than _bocourti_. The tadpoles of the two
species are nearly identical (see Stuart, 1943:12).
~Leptodactylus labialis~ (Cope)
_Cystignathus labialis_ Cope, Proc. Amer. Philos. Soc.,
17:90, 1877.--No type locality designated; type locality
restricted to Potrero Viejo, Veracruz, Mexico, by Smith and
Taylor (1950a:350).
_Leptodactylus labialis_, Brocchi, Mission Scientifique au
Mexique et dans l'Amerique Centrale, pt. 3, sec. 2, livr.
1:20, 1881.
Apatzingan (26); Capirio (5); Cofradia (9); El Sabino (4);
Lombardia; Rio Tepalcatepec, 27 km. S of Apatzingan (2).
In the Tepalcatepec Valley this frog reaches the northernmost known
limit of its range in western Mexico. Although the species is abundant
in the valley, it apparently is absent from the coastal lowlands. In the
Tepalcatepec Valley _Leptodactylus melanonotus_ seems to be more
abundant than _labialis_. In the rainy season both species have been
heard calling from the same ponds and flooded fields.
There are only slight differences in size between the sexes;
measurements of 20 males and eight females are, respectively: snout-vent
length, 32.3-39.5 (35.1), 34.1-39.2 (37.2); tibia length, 14.3-17.0
(15.4), 14.9-16.8 (15.8); head width, 11.0-13.6 (12.0), 12.2-13.2
(12.6); head length, 12.8-15.1 (13.3), 12.8-14.6 (13.7).
~Leptodactylus melanonotus~ (Hallowell)
_Cystignathus melanonotus_ Hallowell, Proc. Acad. Nat. Sci.
Philadelphia, 12:485, 1861.--Nicaragua. Type locality
restricted to Recero, Nicaragua, by Smith and Taylor
(1950a:320).
_Leptodactylus melanonotus_, Brocchi, Mission Scientifique
au Mexique et dans l'Amerique Centrale, pt. 3, sec. 2, livr.
1:20, 1881.
Apatzingan (103); Capirio; Charapendo (7); Coahuayana;
Cofradia (10); El Sabino (21); La Playa (3); Lombardia (5);
Maruata; Nueva Italia (7); Ostula (9); Playa Azul (11); Rio
Marquez, 10 km. S of Lombardia; Rio Marquez, 13 km. SE of
Nueva Italia (6); Rio Tepalcatepec, 27 km. S of Apatzingan.
This species is widespread in the lowlands of the state; it has been
collected up to elevations of 1050 meters in the Tepalcatepec Valley. In
the dry season individuals were discovered beneath rocks along streams
and in damp arroyos; in the rainy season they were found wherever there
was water. Males were heard calling from flooded fields, ditches, rocky
streams, and small puddles. The call is a series of individual notes:
"woink, woink, woink."
Adult males are noticeably smaller than females; measurements for 20
males and ten females from Apatzingan are, respectively: snout-vent
length, 29.6-34.6 (32.3), 36.3-44.1 (40.8); tibia length, 12.6-15.1
(14.0), 16.5-19.0 (17.8); head width, 10.8-11.9 (11.3), 12.6-14.8
(13.7); head length, 11.2-13.2 (11.9), 13.1-14.8 (14.0). Brownish yellow
ventral glands are present in some juveniles and in some adults
collected in the dry season as well as in the rainy season.
~Leptodactylus occidentalis~ Taylor
_Leptodactylus occidentalis_ Taylor, Trans. Kansas Acad.
Sci., 39:349, 1937.--Tepic, Nayarit, Mexico.
Five km. W of Tangamandapio.
On the night of June 11, 1958, this species was calling from a
hyacinth-choked ditch. Although numerous individuals were heard, only
one specimen was obtained. The frogs were calling from the tangled mat
of hyacinths along with _Hyla eximia_, _Hypopachus oxyrrhinus ovis_, and
_Rana pipiens_.
Taylor (1936a:352) characterized this species as follows: "The narrow
head, small maximum size (38 mm. for females, 33 mm. for males), the
character of the postaxillary and postfemoral glands, the narrower
groups of vomerine teeth, clearly distinguish this western Mexican form
from the more robust, larger _melanonotus_ to the south. The call is
likewise fainter and different in quality." Concerning the glands,
Taylor (_loc. cit._) remarked: "There is a possibility that the horny
excrescence covering the glands may appear only during the breeding
season. This character is quite as strongly marked in females as in
males." Bogert and Oliver (1945:324) concluded that the population of
_Leptodactylus_ in northwestern Mexico could not be distinguished from
_melanonotus_ in other parts of the country and thus synonymized
_Leptodactylus occidentalis_ with _melanonotus_. Bogert and Oliver (op.
cit.: 324) stated that the extent as well as the presence or absence of
ventral glands was highly variable in all samples examined by them.
Upon seeing numerous living individuals of _Leptodactylus melanonotus_
from many parts of its range in Mexico and individuals of the population
of _Leptodactylus_ in northwestern Mexico (Nayarit and Sinaloa), I was
immediately impressed not so much by the differences in the development
of the ventral glands, but by the color of the glands. The differences
in color are apparent in freshly preserved specimens. With the exception
of _Leptodactylus_ from northwestern Mexico, specimens of _melanonotus_
from throughout Mexico and northern Central America have yellow or
yellowish brown glands. Specimens from northwestern Mexico have black or
brownish black glands that are conspicuously darker than those found in
_melanonotus_. Examination of 653 preserved specimens of _Leptodactylus
melanonotus_ from Mexico and Guatemala has failed to reveal specimens
with black ventral glands, like those found in specimens from
northwestern Mexico, to which the name _Leptodactylus occidentalis_ has
been applied. Furthermore, in _melanonotus_ the glands are less distinct
and more extensive than in _occidentalis_; in the latter species glands
are absent from the throat and midventral area, where they often are
present in _melanonotus_ (Fig. 7).
In some individuals of both species collected in the dry season and in
some collected in the rainy (breeding) season the glands are absent; the
development of these glands, therefore, does not seem to be correlated
with breeding. Likewise, the glands are present or absent in either sex,
and often as not they are present in juveniles. Presence of the glands,
therefore, cannot be correlated either with sexual or ontogenetic
development. Since the glands are found in individuals from all parts of
the range, it is unlikely that there is a correlation between the
development of the glands and the environment.
[Illustration: FIG. 7. Diagrammatic view of ventral surfaces of
_Leptodactylus melanonotus_ (A) and _Leptodactylus occidentalis_ (B),
showing usual position and size of glandular areas. Approx. natural
size.]
Aside from the differences in the ventral glands, the call is different
in the two populations. The call of _Leptodactylus occidentalis_ is a
rather harsh "wack, wack, wack" as contrasted with the more nasal
"woink, woink, woink" of _melanonotus_. Sound spectrographs are needed
to analyze the differences in calls. None of the specimens of
_occidentalis_ examined approaches in size the largest individuals of
_melanonotus_; possibly the size of the frogs is another valid character
for separating the species. On the basis of the above data it is evident
that the frogs in northwestern Mexico show certain characters that
distinguish them from _Leptodactylus melanonotus_, as it is known
throughout the rest of Mexico. It is not known for certain that
_melanonotus_ and _occidentalis_ are sympatric. Several series of old,
poorly preserved specimens from Nayarit and Sinaloa cannot be placed in
either species, for none has visible ventral glands. _Leptodactylus
melanonotus_ is known from Acaponeta, Nayarit (AMNH 43913-25), and the
following localities in Jalisco: Barro de Navidad (UMMZ 118098), La
Concepcion (UMMZ 113081), La Resolana (UMMZ 102104), and Tenachitlan
(UMMZ 113045-6). Records for _Leptodactylus occidentalis_ are: Alamos,
Sonora (AMNH 51356-65); Culiacan (AMNH 49511-9), Chele (UMMZ 110914),
and Rosario (UMMZ 113062) in Sinaloa; Ixtlan del Rio (UMMZ 102108), San
Blas (UMMZ 112814, 112994, 110892, 115543), and Tepic (UMMZ 115544) in
Nayarit; Ameca (UMMZ 102106-7) and La Cofradia on the south shore of
Lago de Chapala (UMMZ 102105) in Jalisco; and Tangamandapio, Michoacan
(UMMZ 119145). From these scattered records it appears that
_Leptodactylus occidentalis_ in the southern part of its range stays in
the uplands, whereas _melanonotus_ is confined to the lowlands.
~Microbatrachylus hobartsmithi~ (Taylor)
_Eleutherodactylus hobartsmithi_ Taylor, Trans. Kansas Acad.
Sci., 39:355, 1937.--Uruapan, Michoacan, Mexico.
_Microbatrachylus hobartsmithi_ Taylor, Univ. Kansas Sci.
Bull., 26:501, November 27, 1940.
Cascada Tzararacua (6); 21 km. W of Ciudad Hidalgo; 29 km. E
of Morelia; Puerto Hondo; San Jose de la Cumbre (13);
Uruapan (2); Zitacuaro.
Of six specimens from Cascada Tzararacua, five are colored like typical
_M. hobartsmithi_, having the anterior and posterior surfaces of the
thighs and the upper arms pale pink in life and a grayish brown dorsum
in preservative. The other specimen (UMMZ 94231) has in preservative a
dark brown dorsolateral line on each side enclosing a pale tan area that
extends from the snout to the vent. One specimen from 29 kilometers east
of Morelia (UIMNH 40338) and 13 specimens from San Jose de la Cumbre
(UMMZ 102111) do not have the prominent tarsal tubercles characteristic
of _M. hobartsmithi_. Also, in these fourteen specimens the palmar
tubercles are larger, and the dark anal patch more distinct, than in
typical _M. hobartsmithi_. Possibly these specimens, which are from the
high mountains in the eastern part of Michoacan, represent another
species of _Microbatrachylus_. However, Taylor (1940d:501) reported a
series of _M. hobartsmithi_ from the mountains 10 miles west of Villa
Victoria in the western part of the state of Mexico.
The largest specimen from Michoacan is a gravid female (UIMNH 16104)
having a snout-vent length of 23.5 mm.
_Microbatrachylus hobartsmithi_ has been found in rocky ravines along
streams in the Cordillera Volcanica and the southwestern escarpment of
these mountains at elevations from 1450 to 2750 meters.
~Microbatrachylus pygmaeus~ (Taylor)
_Eleutherodactylus pygmaeus_ Taylor, Trans. Kansas Acad.
Sci., 39:352, 1937.--1 mile north of Rodriguez Clara,
Veracruz, Mexico.
_Microbatrachylus pygmaeus_ Taylor, Univ. Kansas Sci. Bull.,
26:500, November 27, 1940.
_Microbatrachylus albolabris_ Taylor, Univ. Kansas Sci.
Bull., 26:502, November 27, 1940.--2 miles west of Cordoba,
Veracruz, Mexico.
_Microbatrachylus minimus_ Taylor, Univ. Kansas Sci. Bull.,
26:507, November 27, 1940.--Agua del Obispo, Guerrero,
Mexico.
_Microbatrachylus imitator_ Taylor, Univ. Kansas Sci. Bull.,
28:70, May 15, 1942.--La Esperanza, Chiapas, Mexico.
Arteaga (328).
This large series (UMMZ 119247-8) was collected on June 22 and 23, 1958,
before the onset of the heavy summer rains. The frogs were found in a
shaded ravine at the north edge of Arteaga; they were obtained during
the day, at which time they were actively moving about in the leaf
litter along a small stream.
These frogs are all referred to _M. pygmaeus_, because this is the
earliest name available for frogs showing the variation in
characteristics displayed by this large series. The characters used by
Taylor (1936a, 1940d, 1941a, and 1942b) and Smith and Taylor (1948) to
distinguish the various species of _Microbatrachylus_ include color
pattern, relative length of the hind limb, presence and position of
dorsal dermal folds or pustules, relative size of inner and outer
metatarsal tubercles, and the number of palmar tubercles. All specimens
from Arteaga have two palmar tubercles; the inner and outer metatarsal
tubercles are subequal in size. Furthermore, aside from sexual
difference, there is little variation in the relative length of the hind
limbs (Table 3). However, many color patterns do exist in the series;
each of these color patterns is described below.
TABLE 3.--SNOUT-VENT LENGTH EXPRESSED AS A PERCENTAGE OF TIBIA LENGTH IN
ANIMALS OF SIX COLOR PATTERNS OF MICROBATRACHYLUS PYGMAEUS. (LETTERS
REFER TO THE VARIANTS HAVING THE COLOR PATTERN DISCUSSED IMMEDIATELY
BELOW)
+------------+-----------+---------+---------+----+--------+
| | | Number | Range | | Twice |
| Color | Sex | of | of |Mean|standard|
|Pattern | |specimens|variation| | error |
| | | | | |of mean |
+------------+-----------+---------+---------+----+--------+
| A | Male | 25 |51.4-57.5|55.2| 3.34 |
| | Female | 25 |49.3-54.9|51.6| 3.12 |
| B | Male | 20 |51.0-57.1|55.4| 2.44 |
| | Female | 21 |47.3-54.9|51.2| 3.52 |
| C | Male | 6 |54.5-56.2|55.2| .... |
| | Female | 6 |50.0-52.9|51.6| .... |
| D | Male | 17 |52.9-58.2|55.4| 2.64 |
| | Female | 14 |48.5-56.6|52.1| 4.16 |
| E | Male | 10 |50.9-56.9|55.1| 3.40 |
| | Female | 7 |49.6-54.5|51.6| .... |
| F | Female | 2 |51.9-52.6|52.3| .... |
+------------+-----------+---------+---------+----+--------+
A.--225 specimens: Dorsum mottled brown and cream, usually with a dark
spot between the eyes and one or two dark V-shaped marks with the apex
anteriorly on the back; 55 of these have a narrow cream-colored line
from the tip of the snout to the vent and thence onto the posterior
surfaces of the thighs. All are pustulate above; in most specimens the
pustules form no pattern, but in some they tend to form a V in the
scapular region.
B.--41 specimens: Dorsum pale tan or cream-color with brown mottling on
flanks; a brown interorbital bar and a brown chevron in scapular region.
Dorsum irregularly pustulate; in some specimens the pustules tend to
form a V in the scapular region.
C.--12 specimens: Dorsum colored like "A", but having a broad yellow
stripe narrowly bordered by black from the tip of the snout to the vent;
in some specimens there is a narrow yellow stripe on the posterior
surfaces of the thighs. The dorsum is irregularly pustulate.
D.--31 specimens: Dorsum variably streaked with cream-color or pale tan
and brown; usually a broad cream-colored stripe from eyelid to groin
bordered laterally by a somewhat narrower brown stripe; middorsal area
cream-color and separated from dorsolateral cream-colored stripe by a
brown stripe, or middorsal area brown with a cream-colored or yellow,
narrow stripe from tip of snout to vent; a dark stripe from tympanum to
flank; dorsal surfaces of heels creamy white to pale orange; anal patch
brown. A dermal ridge from posterior edge of eyelid to rump; another
ridge extends posteromedially from the eyelid; scattered pustules on the
dorsum in some specimens.
E.--17 specimens: A narrow dark stripe from snout, through nostril and
eye, over tympanum, to vent, enclosing a unicolor dorsum (reddish tan to
yellowish tan in life); heels pale tan or yellow above; anal patch
black. A faint dermal ridge from posterior edge of eyelid to rump, or
part way to rump.
F.--2 specimens: Mottled brown and cream-color above; upper lips and
upper arms white. A dermal fold from posterior edge of eyelid to rump;
scattered pustules on dorsum.
Some of these color variants are assignable to names proposed by Taylor:
"A" and "B" undoubtedly are _M. pygmaeus_ (Taylor, 1936a); "C" probably
is _M. pygmaeus_; "D" is referable to _M. minimus_ (Taylor, 1940d) in
most characteristics, although the coloration is more nearly like that
of _M. lineatissimus_ (Taylor, 1941a), a larger species characterized by
a relatively long hind limb; "E" apparently is _M. imitator_ (Taylor,
1942b); "F" is _M. albolabris_ (Taylor, 1940d). Examination of series of
these frogs from other parts of Mexico shows a similar composition of
color variants. Of 78 specimens from the Rio Sarabia and the village of
Sarabia in Oaxaca (UMMZ 115428-37), 57 are "A," six are "D," three are
"E," and 12 are "F"; of 22 specimens from Teapa, Tabasco (UMMZ 113829),
11 are "A," five are "D," two are "E," and four are "F"; of 33 specimens
from Potrero Viejo, Veracruz (USNM 115447-58, 115461-71, 116840-2,
116864-70), ten are "A," 13 are "E," and ten are "F"; of 31 specimens
from La Esperanza, Chiapas (USNM 115477-9, 116827-39, 116849-63), 28 are
"A" and four are "F."
It is highly doubtful if these color variants are actually distinct
species. Goin (1950 and 1954) in his studies of inheritance of color
pattern in West Indian species of the genus _Eleutherodactylus_ has
shown that similar color pattern variants come from the same clutch of
eggs; furthermore, Goin has worked out the genetic ratios of certain of
these variants. Heathwole (_in litt._) obtained "normal" specimens and
individuals having a broad middorsal stripe ("C" in figure 9) from a
clutch of eggs of _Eleutherodactylus gollmeri_. The presence of a broad
middorsal yellow stripe is common in _Eleutherodactylus rugulosus_.
Perhaps the most interesting aspect of variability in color pattern in
Mexican eleutherodactylids is the parallelism between members of the
_Eleutherodactylus rhodopis_-group and some members of
_Microbatrachylus_. In the former group there are white-lipped
individuals (_Eleutherodactylus beatae_ Boulenger), individuals having a
unicolor reddish or yellowish dorsum (_E. dorsoconcolor_ Taylor), and
individuals having a dorsal pattern of irregular longitudinal brown and
cream-colored streaks (_E. venustus_ Guenther). In the humid forests of
southern Veracruz, northern Oaxaca, and Chiapas members of both groups
occur sympatrically. A proper understanding of the evolutionary
significance of these variants in the two groups, as well as proper
allocation of the presently recognized species, must await experimental
evidence based on studies of the inheritance of color pattern.
Nevertheless, at present it is apparent that certain characters,
especially the nature of the dermal folds and pustules, and the color
pattern, are of little taxonomic value in distinguishing "species" of
_Microbatrachylus_. The data derived from a study of the large series
from Arteaga, together with that from the other series examined,
suggests that _Microbatrachylus albolabris_, _imitator_, _minimus_, and
_pygmaeus_ are morphotypes of one species. Of these names, _pygmaeus_ is
the oldest. Consequently _Microbatrachylus pygmaeus_ has been used here
for the series from Arteaga.
Although _Microbatrachylus hobartsmithi_, a species distinguished from
all of the above by the presence of tubercles on the outer edge of the
tarsus, is known from Michoacan northward into Nayarit,
_Microbatrachylus pygmaeus_ previously has not been known north of
Guerrero, where it occurs in habitats similar to that in which it was
collected at Arteaga.
~Eleutherodactylus augusti cactorum~ Taylor
_Eleutherodactylus cactorum_ Taylor, Univ. Kansas Sci. Bull.,
25:391, July 10, 1939.--20 miles northwest of Tehuacan,
Puebla, Mexico.
_Eleutherodactylus augusti cactorum_, Zweifel, Amer. Mus.
Novitates, 1813:20, December 23, 1956.
Cheran; Coalcoman; Uruapan.
The few specimens indicate that this species occurs at moderate to high
elevations in the state. The specimens from Cheran and Uruapan were
obtained in pine forests; the specimen from Coalcoman was found on a
rocky hillside covered with dense forest and located about 100 meters
below the lower limits of the pine forest in the area. A specimen from
Rancho Reparto (elevation 1850 meters) on the west slope of Cerro
Barolosa was lost.
The specimen from Coalcoman (UMMZ 104728) is a juvenile having a
snout-vent length of 25.0 mm. In life it was tan above, mottled with
olive-green. The ventral surfaces were gray; the hind limbs were
distinctly barred with yellow and brown, and the lips were barred with
yellow and black.
~Eleutherodactylus occidentalis~ Taylor
_Eleutherodactylus occidentalis_ Taylor, Proc. Biol. Soc.
Washington, 54:91, July 31, 1941.--Hacienda El Florencio,
Zacatecas, Mexico.
Arteaga (2); Cascada Tzararacua; Coalcoman (2); 19 km. SW of
Coire (3); La Placita (7); Los Reyes; Ostula (4); Pomaro
(2).
The locality records for this species suggest that it is a member of a
group of animals, the distribution of which includes the western part of
the Mexican Plateau and the Pacific lowlands. In Michoacan this frog
has been collected in pine-oak forest at Cascada Tzararacua and at Los
Reyes, in arid scrub forest at Arteaga and Coalcoman, and in tropical
semi-deciduous forest on the lower Pacific slopes of the Sierra de
Coalcoman. On July 5, 1950, James Peters (1954:6) found calling males at
La Placita.
Most of the specimens are immature; four adult males have snout-vent
lengths of 30.9-33.0 (32.2) mm. In all specimens the first finger is
noticeably longer than the second; the inner metatarsal tubercle is
large, flat, and cream-colored, contrasting with the dark brown sole of
the foot. When the hind limbs are adpressed, the heels broadly overlap.
Characteristically, a dark line extends from the snout, through the eye,
above the tympanum, to a point above the insertion of the forelimb.
Usually there is a dark bar behind the tympanum, two dark brown bars
from the eye to the mouth and thence onto the lower jaw, and another
dark bar on the upper lip between the eye and nostril. One adult from
Arteaga, an adult and a juvenile from La Placita, and one juvenile each
from Coire, Ostula, and Pomaro, have the lower lip barred with dark
brown and white, and have a white stripe extending the length of the
upper lip. In life the dorsum varies from dark gray or olive-brown to
tan or reddish brown.
This species belongs to a group containing two other populations that
are currently recognized as species--_calcitrans_, known only from
Omiltemi, Guerrero, and _mexicanus_, reported from the mountains of
Oaxaca. Another apparently undescribed member of this group has been
collected in the mountains of northern Puebla. The locality records
indicate that the group inhabits the mountains on the periphery of the
Mexican Plateau, except in western Mexico, where _Eleutherodactylus
occidentalis_ extends to the Pacific lowlands.
~Eleutherodactylus rugulosus vocalis~ Taylor
_Eleutherodactylus vocalis_ Taylor, Univ. Kansas Sci. Bull.,
26:401, November 27, 1940.--Hacienda El Sabino, Michoacan,
Mexico.
Arteaga (10); El Sabino (8); Salitre de Estopilas (3);
Tumbiscatio (2); Tzitzio (2).
The distributional data on this frog in Michoacan indicate that it
inhabits riparian situations in arroyos and canyons in the lower slopes
of the Cordillera Volcanica and the Sierra de Coalcoman, where it has
been taken at elevations only below 1100 meters.
The dorsal color of living individuals from Arteaga varied from dark
gray and olive brown to tan and reddish brown. The iris was grayish
brown. In contrast, individuals from Agua del Obispo, Guerrero, had pale
golden eyes; specimens from Matias Romero, Oaxaca, had gold eyes heavily
flecked with gray; and individuals from Volcan San Martin, Veracruz, had
bronze eyes.
The use of the trinomial here is arbitrary. Frogs of the
_Eleutherodactylus rugulosus_ group in Mexico (_rugulosus_, _avocalis_,
and _vocalis_) exhibit only slight differences in size, proportions, and
coloration (Duellman, 1958c:6). Furthermore, the named populations are
allopatric. _Eleutherodactylus rugulosus vocalis_, as defined by
Duellman (_loc. cit._), occurs in the foothills of the Sierra Madre
Occidental and associated ranges from central Sinaloa southward into
Michoacan.
~Tomodactylus angustidigitorum~ Taylor
_Tomadactylus angustidigitorum_ Taylor, Univ. Kansas Sci.
Bull., 26:494, November 27, 1940.--Quiroga, Michoacan,
Mexico.
Angahuan (6); Apo; Carapan (21); 19 km. S of Carapan (13);
Cerro Tancitaro (12); Cheran; Corupu (14); Cuseno Station
(14); Opopeo (3); Paracho (11); Paricutin (2); Patzcuaro
(3); Quiroga (59); San Juan de Parangaricutiro (16);
Tancitaro (25); Uruapan (8); Zacapu (11).
This species is indigenous to the pine-oak forests on the southern rim
of the Mexican Plateau, and has been collected at elevations from 1500
to 2500 meters. Males have been observed to call from rocks, rock
fences, clumps of grass, and low bushes; the call is a single "peep." At
San Juan de Parangaricutiro numerous specimens were found in the daytime
beneath adobe bricks and lava on the volcanic ash derived from Volcan
Paricutin; at Paracho individuals were found by day beneath rocks in a
pine forest.
In most specimens the dorsum is dark reddish brown, and the prominent
inguinal glands are cream-color or pale orange (Pl. 3, Fig. 1). Of eight
individuals collected at Paracho, one was reddish brown, two were
pinkish tan, three were dark brown, and two were black.
~Tomodactylus fuscus~ Davis and Dixon
_Tomodactylus fuscus_ Davis and Dixon, Herpetologica,
11:157, July 15, 1955.--1.5 miles southeast of Huitzilac,
Morelos, Mexico.
Los Cantiles (2); 28 km. E of Morelia.
The range of this species includes the Sierra Ajusco in Mexico and
Morelos and thence westward to the Serrania Ucareo in Michoacan. The
specimen from 28 kilometers east of Morelia was found in an oak forest
on a steep hillside at an elevation of 2100 meters. One from Los
Cantiles was calling from a steep cliff at an elevation of 2200 meters
in pine-oak forest. This specimen (UMMZ 119156) in life had a pale
olive-brown dorsum with irregular dark brown mottling and transverse
bars on the limbs. The interorbital bar, the upper arms, and the tips of
the dorsal pustules were pale orange; the iris was pale grayish gold
(Pl. 3, Fig. 2).
~Tomodactylus nitidus nitidus~ (Peters)
_Liuperus nitidus_ Peters, Monats. Akad. Wiss. Berlin, p.
878, 1869.--Izucar de Matamoras, Puebla, Mexico.
_Tomodactylus amulae_ Guenther, Biologia Centrali-Americana,
Reptilia and Batrachia, p. 219, April, 1900.--Amula,
Guerrero, Mexico.
_Tomodactylus nitidus nitidus_, Dixon, Texas Jour. Sci.,
9:385, December, 1957.
Copuyo (15); Tuxpan (8); Tzitzio (11).
One specimen from Tzitzio (UMMZ 99155) was referred to _Tomodactylus
nitidus petersi_ by Dixon (1957:390). A re-examination of this specimen,
and examination of ten others from the same locality (UMMZ 121571)
reveals that the relatively small size of the tympanum and absence of
dense ventral spotting place these specimens closer to _T. nitidus
nitidus_ than to _T. nitidus petersi_.
The specimens from Tuxpan (UMMZ 114303-4) had in life a gray to olive
tan ground color with dark olive-green markings, bright yellow thighs
with olive-green transverse bands, yellowish tan shanks with olive-green
bars, yellow groin, white inguinal glands with black markings, grayish
white belly with scattered brownish black spots in some specimens, and a
deep golden iris (Pl. 4, Fig. 1). These specimens were found calling
from bushes in a rocky field at an elevation of 1800 meters. The call is
a high-pitched "pee-ee-eep."
~Tomodactylus nitidus orarius~ Dixon
_Tomodactylus nitidus orarius_ Dixon, Texas Jour. Sci.,
9:392, December, 1957.--4.5 miles southwest of Tecolapa,
Colima, Mexico.
La Placita (3); Pomaro.
These specimens, referred to _Tomodactylus petersi_ by Duellman
(1954b:5), were included in _T. nitidus orarius_ by Dixon (1957:392).
Color notes based on living individuals from Tecolapa, Colima (UMMZ
114312 and 116922), are: gray above mottled with brown; venter dirty
white; anterior and posterior surfaces of thighs bright yellow; iris
pale golden (Pl. 4, Fig. 2). The call is a soft "braa" usually followed
by three high notes: "braaa-eep-ee-eep." In Michoacan this subspecies
has been found only in the coastal region and the lower foothills of the
Sierra de Coalcoman, an area in which it replaces _Tomodactylus nitidus
petersi_. This is the only _Tomodactylus_ known to inhabit coastal
lowlands.
~Tomodactylus nitidus petersi~ Duellman
_Tomodactylus petersi_ Duellman, Occ. Pap. Mus. Zool. Univ.
Michigan, 560:5, October 22, 1954.--Coalcoman, Michoacan,
Mexico.
_Tomodactylus nitidus petersi_, Dixon, Texas Jour. Sci.,
9:390, December, 1957.
Aguililla; Apatzingan (8); Cascada Tzararacua: Charapendo
(5); Coalcoman (5); 18 km. E of Dos Aguas (6); El Sabino
(5); La Playa (2); Jiquilpan; Uruapan (2); Volcan Jorullo;
Zamora.
In life, specimens from Apatzingan (UMMZ 114308-9) varied in dorsal
color from grayish tan to pale brown; the dorsal markings were olive
green. The thighs and groin were yellowish orange; the iris was pale
golden, and the vocal sac was purplish gray (Pl. 5, Fig. 1).
Measurements for 13 adult males from the Tepalcatepec Valley are:
snout-vent length, 21.9-26.8 (24.3); tibia length, 8.4-9.9 (9.3); head
width, 7.2-9.2 (7.8); head length, 7.6-8.7 (8.2).
At Apatzingan and Charapendo in the Tepalcatepec Valley males were found
calling from rocks and bushes in open arid tropical scrub forest. The
call, a triple note "peep-ee-eep," is repeated once every 90 to 135
seconds. _Tomodactylus nitidus petersi_ probably ranges throughout the
Tepalcatepec Valley and surrounding foothills. Dixon (1957:392) referred
the specimens from Zamora, Jiquilpan, and Uruapan to this subspecies.
Uruapan is near the lower limits of the pine forest on the slopes of the
Cordillera Volcanica; Zamora and Jiquilpan are on a low part of the
Mexican Plateau southeast of Lago de Chapala.
~Tomodactylus rufescens~ Duellman and Dixon
_Tomodactylus rufescens_ Duellman and Dixon, Texas Jour.
Sci., 11:78, March, 1959.--Dos Aguas, Michoacan, Mexico.
Dos Aguas (14); 18 km. E of Dos Aguas (6).
Fourteen specimens from the pine-oak forests around Dos Aguas (UMMZ
118503-10, 121498-9) have reddish brown dorsal color and a narrow
cream-colored middorsal line (Pl. 5, Fig. 2). Twelve of these specimens
are adult males having snout-vent lengths of 20.7 to 24.6 (22.5) mm. One
female has a snout-vent length of 24.8 mm., and one juvenile has a
snout-vent length of 14.5 mm. Six specimens are from a region of mixture
of pine-oak forest and arid tropical scrub forest at 18 kilometers east
of Dos Aguas (UMMZ 121497, 121500). All are males having snout-vent
lengths of 18.0 to 22.6 (20.7) mm. The dorsum is tan marked with black;
the thighs are yellowish orange.
The specimens from 18 kilometers east of Dos Aguas were found on July
22, 1960, by Floyd L. Downs and John Winklemann, who collected calling
males of _Tomodactylus rufescens_ and _Tomodactylus nitidus petersi_ at
the same locality. Downs (_personal communication_) stated the call was
a single note. At Dos Aguas I heard _T. rufescens_ give two calls, one a
single "peep," the other a triple note--"pee-ee-eep."
In the higher parts of the Sierra de Coalcoman _Tomodactylus rufescens_
seems to fill the same niche as _T. angustidigitorum_ does in the
Cordillera Volcanica. At lower elevations in their respective mountain
ranges the species occur sympatrically with _T. nitidus petersi_.
~Diaglena reticulata~ Taylor
_Diaglena reticulata_ Taylor, Univ. Kansas Sci. Bull.,
28:60, May 15, 1942.--Cerro Arenal, Oaxaca, Mexico.
Nueva Italia (3); Ostula (7).
Until recently frogs of the genus _Diaglena_ were known only from a few
specimens from southern Sinaloa (_Diaglena spatulata_) and from the
Pacific lowlands of the Isthmus of Tehuantepec (_Diaglena reticulata_).
Peters (1955a) reported specimens from Ostula, Michoacan, and compared
these specimens with one _D. reticulata_ from Tehuantepec, Oaxaca, and
four _D. spatulata_ from Sinaloa. This comparison showed that the
specimens from Michoacan, although showing some minor differences from
_D. reticulata_, are closer to that species than to _D. spatulata_.
Subsequent to Peters' work, series of both species of _Diaglena_,
including additional specimens from Michoacan and from Colima, have been
collected, and a more qualified comparison is now possible.
In comparing specimens of _D. spatulata_ from southern Sinaloa (UMMZ
115322) with specimens of _D. reticulata_ from Tehuantepec, Oaxaca (UMMZ
115321), the differences noted by Taylor (1942c:60) were found to be
constant. But specimens from Ostula, Michoacan (UMMZ 104418), and five
individuals from Colima (TNHC 26379-83) were found to be intermediate in
certain characters. The skin of the dorsum in _D. reticulata_ is
granular; that in _D. spatulata_ is smooth. The skin in specimens from
Ostula and Colima is slightly granular. The dorsal ground color of _D.
reticulata_ is yellowish brown with dark reticulations; the dorsal
ground color of _D. spatulata_ is olive-green. Specimens from Ostula and
Colima most closely resemble those from Tehuantepec in coloration, but
the reticulations are more coarse, and the ground color has an
olive-green tint. _Diaglena reticulata_ also differs from _D.
spatulata_ in having a larger over-all size, slightly broader head, a
narrower interorbital distance, and a more pointed snout with a deeper
labial shelf (Table 4). The specimens from Ostula and Colima are
intermediate between _D. reticulata_ from Oaxaca and _D. spatulata_ from
Sinaloa in body proportions.
Of three specimens from the Tepalcatepec Valley (JRD 5991-3), only two
are suitable for measuring. These specimens are smaller than adults from
the coastal areas and have broader heads and snouts, but narrower
interorbital distances, than specimens in the other samples (Table 4).
The texture of the skin is like that of specimens from Ostula and
Colima. The coloration resembles that of _D. reticulata_, but the
reticulations are bold and form indistinct bands on the hind limbs.
TABLE 4.--COMPARISON OF FOUR CHARACTERS IN FIVE SAMPLES OF DIAGLENA.
(ALL DATA ARE FOR MALES; MEANS GIVEN IN PARENTHESES BELOW RANGES.)
+-----------------+---------+----------+----------+---------+-----------+
| | | | Head | Inter- |Internarial|
| | | | width | orbital | distance |
| | Number |Snout-vent| |distance | |
| Locality | of | length |----------+---------+-----------+
| |specimens| |Snout-vent| Head | Head |
| | | | length | width | width |
+-----------------+---------+----------+----------+---------+-----------+
| Oaxaca | 9 |71.1-87.5 |25.4-29.1 |63.0-71.4| 11.9-13.8 |
| | | (80.7) | (27.9) | (67.1) | (12.9) |
+-----------------+---------+----------+----------+---------+-----------+
| Coast of | 5 |72.0-79.2 |24.3-27.2 |67.0-73.8| 13.7-14.4 |
| Michoacan | | (74.8) | (25.6) | (71.4) | (14.1) |
+-----------------+---------+----------+----------+---------+-----------+
| Colima | 4 |71.7-79.6 |26.1-28.6 |70.5-75.3| 16.0-17.9 |
| | | (74.8) | (27.4) | (72.0) | (16.6) |
+-----------------+---------+----------+----------+---------+-----------+
| Tepalcatepec | 2 |63.0-65.4 |28.3-32.2 |57.3-62.4| 17.0-20.2 |
| Valley | | (64.2) | (30.3) | (59.9) | (18.6) |
+-----------------+---------+----------+----------+---------+-----------+
| Sinaloa | 11 |71.9-81.3 |24.0-27.3 |70.5-78.1| 15.0-17.3 |
| | | (77.3) | (25.7) | (73.4) | (16.1) |
+-----------------+---------+----------+----------+---------+-----------+
All specimens from Michoacan and Colima more closely approach _Diaglena
reticulata_ than _D. spatulata_. The acquisition of additional
specimens, especially from the area between Sinaloa and Colima and from
Guerrero, is necessary to determine the relationships among the various
populations known at present. Both species of _Diaglena_ inhabit
tropical scrub forest; none has been found in the more humid and
tropical semi-deciduous forests. Humid forest replaces the scrub forest
in the lowlands of southern Nayarit and northern Jalisco; possibly this
forest acts as a barrier to the distribution of _Diaglena_ and thus
serves as a divider between the ranges of _D. spatulata_ to the north
and _D. reticulata_ to the south.
~Pternohyla fodiens~ Boulenger
_Pternohyla fodiens_ Boulenger, Ann. Mag. Nat. Hist., ser.
5, 10:326, 1882.--Presidio, Sinaloa, Mexico.
Nueva Italia (2).
These specimens (JRD 5994-5) were found on the road near Nueva Italia
during a heavy rain on the night of August 25, 1960, by James R. Dixon.
Both are females having snout-vent lengths of 64.0 and 59.0 mm. They are
typical of the species as it is known from Sinaloa, Nayarit, Jalisco,
and Colima.
These specimens constitute the southernmost record for the species,
which ranges in semi-arid habitats from southern Arizona southward along
the Pacific lowlands of Mexico to Colima and inland on the Mexican
Plateau in Jalisco.
~Phyllomedusa dacnicolor~ Cope
_Phyllomedusa dacnicolor_ Cope, Proc. Acad. Nat. Sci.
Philadelphia, 16:181, September 30, 1864.--Colima, Colima,
Mexico. Funkhouser, Occ. Pap. Nat. Hist. Mus. Stanford
Univ., 5:37, April 1, 1957.
_Agalychnis alcorni_ Taylor, Copeia, no. 2:31, June 2,
1952.--Rio Tepalcatepec, 17 miles south of Apatzingan,
Michoacan, Mexico.
_Agalychnis dacnicolor_, Duellman, Herpetologica, 13:29,
March 30, 1957.
_Phyllomedusa alcorni_, Funkhouser, Occ. Pap. Nat. Hist.
Mus. Stanford Univ., 5:30, April 1, 1957.
Aguililla (13); Apatzingan (7); Charapendo; Coahuayana (3);
Coalcoman (54); El Sabino; Huetamo Road (2); La Orilla; La
Placita; Nueva Italia (4); 32 km. E of Neuva Italia (2); Rio
Cancita, 14 km. E of Apatzingan; Rio Tepalcatepec, 27 km. S
of Apatzingan; Salitre de Estopilas (2); Tzitzio (4).
This large tree frog has been found only in the lowlands below elevation
of about 1000 meters, usually in arid tropical scrub forest. Calling
males were heard on rainy nights throughout the rainy season; in nearly
every instance both males and females were found in low trees and
bushes. On summer nights when there had been no rain, adults were found
sitting on bushes in the scrub forest.
At Coalcoman on July 1, 1955, a chorus was heard at midday. About forty
_Phyllomedusa dacnicolor_ were found in one guayava bush at the edge of
a recently dried pond. Individual males were calling; clasping males
were silent. The call is a barking groan. Fifteen individual egg masses
were hanging from branches and leaves in tear-drop fashion. Each egg
mass contained 100 to 350 pale green eggs, located only in the exterior
part of the clear gelatinous mass. Two composite egg masses appeared to
have been made up by egg deposition on the part of three to five females
(Pl. 2, Fig. 2).
As shown by Duellman (1957a), the characters used by Taylor (1952) to
diagnose _Phyllomedusa alcorni_ are sexually dimorphic. Funkhouser
(1957) apparently was unaware of this sexual dimorphism, for she
recognized _P. alcorni_ and _P. dacnicolor_ as distinct species.
~Phrynohyas inflata~ (Taylor)
_Acrodytes inflata_ Taylor, Univ. Kansas Sci. Bull., 30:64,
June 12, 1944.--La Venta, Guerrero, Mexico.
_Phrynohyas inflata_, Duellman, Misc. Publ. Mus. Zool. Univ.
Michigan, 96:19, February 1, 1956.
_Phrynohyas corasterias_ Shannon and Humphrey,
Herpetologica, 13:15, March 30, 1957.--4.8 miles east of San
Blas, Nayarit, Mexico.
Barranca de Bejuco.
One specimen of this large species was collected in 1951; it was found
on a low branch in tropical semi-deciduous forest at an elevation of 65
meters. In life there were olive-gray blotches on a pale gray dorsum;
the iris was a dark golden color.
This species, which is known from only a few specimens, seems to be
restricted to the coastal lowlands and low foothills from Guerrero
northward to Nayarit. Shannon and Humphrey (1957) described _Phrynohyas
corasterias_ from Nayarit. Their description was based on a small female
having a snout-vent length of 34.4 mm. The new species was diagnosed as
differing from _P. inflata_ in having less webbing on the feet, a poorly
developed supratympanic fold, a more pustulate dorsum, and marked
differences in dorsal pattern, color, and nature of antebrachial
banding. The significance of the webbing was questioned by Shannon and
Humphrey. The nature of the supratympanic fold and dorsal pustules
changes with age (Duellman, 1956a:31). _Phrynohyas inflata_ is known to
attain a snout-vent length of 95 mm. Dermal structures that undergo
ontogenetic change are of little importance in comparing a juvenile with
a large adult. The only significant difference in color pattern between
_P. inflata_ and _P. corasterias_ is the presence of wide transverse
bands on the limbs of the latter. In this respect _P. corasterias_
approaches _P. latifasciata_, a species known only from two specimens
from southern Sinaloa. The acquisition of additional specimens from
Jalisco, Nayarit, and Sinaloa may show that _P. inflata_ and _P.
latifasciata_ are conspecific, as suggested by Duellman (1956a:21).
Nonetheless, the specimen on which the description of _P. corasterias_
was based is not sufficiently different from the known specimens of _P.
inflata_ to warrant specific recognition.
~Hyla arenicolor~
_Hyla arenicolor_ Cope, Jour. Acad. Nat. Sci. Philadelphia,
ser. 2, 6:84, July, 1866.--Northern Sonora, Mexico. Type
locality restricted to Santa Rita Mountains, Pima County,
Arizona, by Smith and Taylor (1950a:354).
Agua Cerca; Cascada Tzararacua (3); Chinapa; Cojumatlan; Dos
Aguas; El Sabino (25); El Espinal; Lago de Camecuaro;
Lombardia (2); Tupataro; Zinapecuaro.
Altitudinally this frog ranges from 500 to 2100 meters; although the
environments in which it has been found vary from open arid tropical
scrub forest to pine forest, it usually is found near rocky streams in
these habitats. There is great disparity in size between specimens from
the mountains and those from the Tepalcatepec Valley. Seven males from
elevations in excess of 1400 meters have an average snout-vent length of
34.7 mm.; nine from elevations below 1000 meters have an average
snout-vent length of 49.1 mm. In life a male collected at night at
Lombardia (UMMZ 112846) had dark brown spots on a grayish brown dorsum;
the groin, anterior and posterior surfaces of the thighs, and ventral
surfaces of the hind limbs and palms were yellowish orange. The belly
and tips of digits were white; the vocal sac was purplish brown, and the
iris was dark grayish gold. In contrast, a specimen obtained in the
daytime at Chinapa (UMMZ 119204) had indistinct gray spots on a pale
ashy gray dorsum; the flash colors were yellow. After dark the spots
were dark olive-brown on a grayish brown dorsum.
Two males were found calling from a rocky stream near Lombardia on July
12, 1955. The call is a nasal "ah-ah-ah-ah."
~Hyla baudini~ Dumeril and Bibron
_Hyla baudinii_ Dumeril and Bibron, Erpetologie generale,
vol. 8:564, 1841.--Mexico. Type locality restricted to
Cordoba, Veracruz, Mexico, by Smith and Taylor (1950a:346).
Aguililla (5); Apatzingan (30); Arteaga; Buena Vista;
Charapendo; Coahuayana; Cofradia (4); El Sabino (12); La
Placita; La Playa; Maruata; Nueva Italia (3); 32 km. E of
Nueva Italia (2); Ostula (4); Rio Tepalcatepec, 25 km. S of
Apatzingan (3); Salitre de Estopilas; San Jose de la Montana
(2); Tumbiscatio; Tzitzio.
This tree frog is widespread in the coastal lowlands and in the
Tepalcatepec Valley up to elevations of about 1200 meters. It is found
in numbers in the early part of the rainy season, at which time males
were heard calling from bushes and trees along ditches and temporary
ponds. The call is a loud nasal "waank-waank-waank." One individual that
was emitting a long and unusually high-pitched call was found to have
one hind limb engulfed by a _Leptodeira maculata_.
When active at night these frogs usually are pale tan to reddish brown
above with dark brown markings. A specimen found sitting on a maguey
plant in the daytime was pale ashy gray with a pale green upper lip.
~Hyla bistincta~ Cope
_Hyla bistincta_ Cope, Proc. Amer. Philos. Soc., 17:87,
1877.--Veracruz, Mexico. Type locality restricted to
Acultzingo, Veracruz, Mexico, by Smith and Taylor
(1950a:346).
Cerro San Andres; Dos Aguas (2); Los Conejos (3); Uruapan
(50).
In the Parque Nacional at Uruapan this species was found in abundance
during the day. The frogs hide in an entanglement of vines and
vegetation overhanging several small spring-fed streams. Tadpoles were
in the rocky streams, and metamorphosing young were on vegetation at the
edges of the streams.
In life the dorsum is greenish tan with brown mottling; in some
individuals the entire dorsum is dark chocolate brown. The flanks are
pale lemon yellow barred with lavender-brown. Notes on the color of a
living frog from Dos Aguas (UMMZ 119193) are: Dorsal ground color a
medium shade of brown with dark brown flecks; flanks black with silvery
white and pale yellow spots; belly pale yellowish white; throat mottled
with grayish brown; iris pale copper color.
[Illustration: FIG. 8. Tadpole of _Hyla bistincta_ (UMMZ 115231) from
Uruapan, Michoacan. x 2.]
Description of Tadpole: Body somewhat depressed; maximum width of body
slightly more than one-half of body length. Nostrils placed
dorsolaterally and directed anteriorly, situated about midway between
tip of snout and eye. Eyes of moderate size, dorsolateral in position
and directed upwards. Tail about twice as long as body, thrice as long
as deep, and tapering gradually to a rounded tip. Tail-musculature not
extending to tip of tail fin. Spiracle sinistral, lateral, and situated
at midbody. Vent dextral; the cloacal tube extending along ventral part
of tail for a distance equal to about one-eighth of body length (Fig.
8). Average body length of six tadpoles with small hind limb buds, 19.5
mm.; tail length, 38.3 mm. Mouth ventral, its width equal to about
two-thirds of greatest width of body. Lips bordered by two rows of small
papillae; row of larger papillae between upper lip and outer upper
tooth-row, similar row between lower lip and outer lower tooth-row;
laterally these rows degenerating into numerous small papillae. Horny
beaks well developed; upper beak moderately arched and deeply indented;
lower beak slightly indented. Serrations of beaks blunt and peglike,
moderately developed on both beaks, but slightly stronger on lower one.
Tooth-rows 2/3; upper rows nearly equal in length and slightly longer
than lower rows, which are subequal in length; inner upper tooth row
interrupted medially by rounded notch; inner lower tooth-row turned
downward laterally; teeth in all rows about equal in size, but
decreasing in length laterally (Fig. 9).
[Illustration: FIG. 9. Mouthparts of larval _Hyla bistincta_ (UMMZ
115231) from Uruapan, Michoacan. x 15.]
Color in formalin: pale grayish brown dorsally and laterally; pale gray
ventrally; tail-musculature brown; tail-fin translucent with scattered
melanophores most numerous on upper fin.
In most details these tadpoles resemble those of _Hyla robertsorum_
described by Rabb and Mosimann (1955).
Four metamorphosing young have snout-vent lengths of 23.0-23.5 (23.2);
a completely metamorphosed individual has a snout-vent length of 24.8
mm.
In Michoacan this stream-breeding hylid occurs at elevations of 1,600 to
2,400 meters in the Sierra de Coalcoman and in the mountains rising from
the Mexican Plateau.
~Hyla eximia~ Baird
_Hyla eximia_ Baird, Proc. Acad. Nat. Sci. Philadelphia,
7:61, October 20, 1854.--Valley of Mexico. Type locality
restricted to Coyoacan, Distrito Federal, Mexico, by Smith
and Taylor (1950a:329).
_Hyla microeximia_ Maslin, Herpetologica, 13:81, July 10,
1957.--3 miles northwest of Jocotepec, Jalisco, Mexico.
Ciudad Hidalgo (36); Cuitzeo; 29 km. NW of Jacona; Jiquilpan
(2); Lago de Camecuaro (2); Lago de Patzcuaro (129); Los
Reyes; Morelia; Sahuayo (3); San Gregorio (63);
Tangamandapio (4); Temazcal (26); Tupataro; Tuxpan (15);
Undameo (2); Uruapan (20); Zacapu; Zamora (27); Zinapecuaro
(10).
More than 80 per cent of the specimens from Michoacan have brown spots
between the lateral and dorsolateral dark stripes, and more than 50 per
cent have spots between the dorsolateral stripes, at least posteriorly.
In comparison with specimens from the Valley of Mexico, those from
Michoacan have more distinct dorsolateral stripes that extend farther
anteriorly, sometimes to the eyelid, and in this respect are more nearly
like those from Jalisco and Nayarit (Taylor, 1939b:425). Some specimens
from the western part of Michoacan possess certain characters used by
Maslin (1957:81) to distinguish _Hyla microeximia_ from _H. eximia_;
nevertheless, the variation is such that two species cannot be
distinguished in Michoacan. Four series of freshly preserved specimens
have been studied in detail; in the discussion below they are arranged
from west to east; the measurement is for snout-vent length of ten males
from each sample:
_Zamora._--Twenty-two specimens (UMMZ 102083), 24.0-27.6
(26.1) mm. Dorsolateral dark stripe, or row of dashes,
present in all specimens; dark spots in lateral and dorsal
green fields; lateral dark stripe confluent with
dorsolateral stripe posteriorly in 18 specimens; white line
not extending to groin.
_Temazcal._--Thirty-five specimens (UMMZ 119162), 26.5-31.1
(28.2) mm. Dorsolateral dark stripe of row of spots present
only posteriorly in most; both dorsolateral stripes and
dorsal spots lacking in four specimens; heavy spotting
dorsally in three others; lateral and dorsolateral dark
stripes confluent posteriorly in three; lateral white stripe
extending to groin in 16 specimens.
_Ciudad Hidalgo._--Thirty-six specimens (UMMZ 119163),
26.4-30.9 (28.2) mm. Dorsolateral dark stripe or row of
spots present only posteriorly in most; no brown spots in
the green fields of many specimens; large brown inguinal
spot in most specimens; heavy spotting dorsally in four;
lateral and dorsolateral dark stripes confluent posteriorly
in five; lateral white line extending to groin in most
specimens.
_Tuxpan._--Fifteen specimens (UMMZ 115227), 28.7-33.0 (30.5)
mm. Dorsolateral dark stripe or row of dashes in all
specimens; dark spots in lateral green fields, at least
posteriorly in most; dark spots posteriorly in the dorsal
green field in five; lateral dark stripe separated from
dorsolateral stripe in all specimens; lateral white line
extends to the groin in all specimens.
As can be seen from the above descriptions, the distinguishing
characters of _Hyla microeximia_--confluence of lateral and dorsolateral
dark stripes posteriorly, extent of lateral white stripe, and
distribution of dark spots dorsally--are found in individuals from all
of the populations sampled. In the samples from western Michoacan there
is a higher incidence of _microeximia_-like frogs than in those from
other parts of the state. _Hyla eximia_ is a wide-ranging species
varying greatly geographically and individually. A thorough review of
the species and related members of the _Hyla eximia_-group is necessary
before certain populations can justifiably be segregated as subspecies
or species.
In Michoacan _Hyla eximia_ has been collected in mesquite-grassland,
pine-oak forest, and cultivated areas on the Mexican Plateau from 1500
to 2300 meters; apparently it is absent from the Sierra de Coalcoman.
This is the most abundant frog on the southern part of the Mexican
Plateau; in the rainy season breeding choruses are found in temporary
pools and in the marshes adjacent to the permanent lakes.
~Hyla lafrentzi~ Mertens and Wolterstorff
_Hyla lafrentzi_ Mertens and Wolterstorff, Zool. Anz.,
84:235, August 25, 1929.--Desierto de los Leones, Distrito
Federal, Mexico.
Cerro San Andres (26); Opopeo (9).
In March, 1949, James A. Peters collected this species at elevations of
2400 to 2800 meters on the west slope of Cerro San Andres. The frogs
were found beneath logs and rocks in a damp canyon in coniferous forest.
Among the juveniles in this series is a completely transformed
individual (UMMZ 102093) having a snout-vent length of 14.5 mm. Five
adults have snout-vent lengths of 36.2-39.5 (38.0) mm. _Hyla lafrentzi_
has noticeably longer hind limbs than _H. eximia_; in the former, when
the hind limb is brought forward along the body, the tibiotarsal
articulation extends to the snout. There are dark transverse bands on
the hind limbs; the dorsolateral stripe is broken into an anterior and a
posterior segment, and the latter is narrowly bordered by white in most
specimens.
_Hyla lafrentzi_ occurs at higher elevations than any other frog in
Michoacan; the locality records from throughout the range indicate that
it is restricted to pine and pine-fir forests. In these habitats it
replaces _Hyla eximia_, which inhabits the lower pine-oak forests and
mesquite-grassland on the Mexican Plateau. Ponds are absent at places
where _Hyla lafrentzi_ has been collected; possibly the eggs are laid in
streams.
~Hyla smaragdina~ Taylor
_Hyla smaragdina_ Taylor, Copeia, No. 1:18, March 30,
1940.--6 kilometers east of Cojumatlan, Michoacan, Mexico.
_Hylella azteca_ Taylor, Proc. Biol. Soc. Washington, 56:49,
June 16, 1943.--Tepoztlan, Morelos, Mexico.
Cojumatlan (30); Copuyo (7); 18 km. E of Dos Aguas (22);
Ostula (8); Pomaro (3); Sahuayo; Salitre de Estopilas (7).
Taylor (1940a:18) diagnosed this species as having few or no vomerine
teeth, no vocal sac, a rather broad and flat head, two large tubercles
below the anus, a granular venter, and a green dorsum in life. The
specimens on which the description was based are either immature or
non-breeding individuals; all were collected from bromeliads growing on
cacti near Cojumatlan. Another small, flat-headed hylid from Tepoztlan,
Morelos, was described and diagnosed by Taylor (1943b:49) as differing
from _Hyla smaragdina_ in having a vocal sac and a broader head. This
specimen was named _Hylella azteca_. Specimens from the coastal region
of Michoacan and Colima were referred to _Hylella azteca_ by Peters
(1954:7) and Duellman (1958c:8).
Comparison of topotypic _Hyla smaragdina_ and the holotype of _Hylella
azteca_ (UIMNH 25044) with the several series of specimens from
Michoacan has resulted in the conclusion that all pertain to only one
species. Although the type series of _Hyla smaragdina_ consists of
immature specimens, the males in that series do possess vocal sacs.
Since these were not breeding individuals, the sacs are not well
developed. The characters of the anal tubercles and the relative width
of the head are of no value in separating the two species. The
apparently aestivating individuals comprising the type series of _Hyla
smaragdina_, and the type of _Hylella azteca_, which also was found in a
bromeliad, were green in life. Of the calling males found on the coast
of Michoacan, most were yellowish tan when found; two were pale green,
but soon changed to pale tan. Calling males from Copuyo and Dos Aguas
were pale yellowish tan. Therefore the color of the dorsum is of little
significance in distinguishing the two named populations.
Males of _Hyla smaragdina_ have been found calling in the months of June
and July from rocky streams; the call is a nasal "haah-haah-haah,"
repeated quickly and constantly for as long as 30 seconds. As pointed
out by Duellman (1958c:9), this breeding behavior is unlike that
suggested by Taylor (1943b:51). In Michoacan _Hyla smaragdina_ has been
found in tropical semi-deciduous forest, oak forest, and
mesquite-grassland at elevations from 150 to 1500 meters.
~Hyla smithi~ Boulenger
_Hyla smithi_ Boulenger, Zool. Rec. Reptilia and Batrachia,
38:33, 1902.--Cuernavaca, Morelos, Mexico.
Aguililla (14); Apatzingan (104); Arteaga; Charapendo (5);
Coalcoman (11); El Sabino (44); La Playa (6); Lombardia (2);
Nueva Italia (8); Playa Azul; Salitre de Estopilas (2).
This small hylid is abundant in the Tepalcatepec Valley to elevations of
about 1000 meters; it was found infrequently on the coastal lowlands.
Males call from bushes in and around flooded fields and ditches, from
grasses and small herbs in the water and from vegetation overhanging
small streams. The call consists of a series of short, high notes,
somewhat reminiscent of a katydid's song. In the dry season occasional
males were heard calling from irrigated fields near Apatzingan. In the
daytime individuals were found in the axils of leaves of the
elephant-ear plants (_Xanthosoma_).
In living individuals the dorsal color usually is uniform pale yellow;
often the lateral white stripe is barely visible. The vocal sac is
bright yellow, and the iris is pale gold. In some individuals there are
scattered dark brown spots or flecks on the back and upper surfaces of
limbs. Twenty males from Apatzingan have the following measurements:
snout-vent length, 22.8-26.0 (25.0) mm., tibia length, 10.7-13.6 (12.6)
mm.; head width, 7.2-8.0 (7.6) mm., head length, 7.1-8.1 (7.7) mm.
~Hypopachus caprimimus~ Taylor
_Hypopachus caprimimus_ Taylor, Univ. Kansas Sci. Bull.,
26:526, November 27, 1940.--Agua del Obispo, Guerrero,
Mexico.
Buena Vista; Copuyo (6); Charapendo (3); Cofradia;
Jaramillo; Jungapeo; San Salvador; Tuxpan.
Specimens of _Hypopachus_ from the Balsas drainage in Michoacan have
characters consistent with topotypic _H. caprimimus_. Eleven specimens
from the southern edge of the Mexican Plateau all have the flanks darker
than the dorsum, a distinct and continuous dark stripe from the occiput
to the groin, a large dark spot in the inguinal region, and a pair of
dark transverse stripes on the thigh and shank (Pl. 6, Fig. 1). With
the exception of three specimens from Charapendo, all have a
predominantly brown venter with round, cream-colored spots.
Peters (1954:8) referred specimens from Buena Vista and San Salvador to
_Hypopachus oxyrrhinus_. He stated that the specimen (BMNH
1914.1.28.150) from San Salvador had flanks much darker than the dorsum
and a well-defined continuous stripe from the occiput to the groin; this
specimen has the characters of _H. caprimimus_. The specimen (BMNH
1914.1.28.151) from Buena Vista resembles _H. oxyrrhinus_ in some
characters, but it is not like _H. oxyrrhinus ovis_ on the Mexican
Plateau in Michoacan. The specimen has paired transverse stripes on the
hind limbs as does _H. caprimimus_, and is here referred to that
species.
In Michoacan this species has been collected in arid tropical scrub
forest at elevations of 200 to 1800 meters in the northern foothills of
the Sierra de Coalcoman, the Tepalcatepec and Tuxpan valleys, and on the
lower slopes of the Cordillera Volcanica. Calling males have been found
along streams. One specimen from Charapendo was regurgitated by a
_Leptodeira maculata_.
~Hypopachus oxyrrhinus ovis~ Taylor
_Hypopachus ovis_ Taylor, Univ. Kansas Sci. Bull., 26:520,
November 27, 1940.--Tepic, Nayarit, Mexico.
_Hypopachus oxyrrhinus ovis_, Shannon and Humphrey,
Herpetologica, 14:89, July 23, 1958.
Emiliano Zapata; 30 km. NW of Jacona (2); 10 km. NE of
Patzcuaro (2); Tangamandapio (16); 24 km. W of Zamora (16).
Thirty-seven specimens from the Mexican Plateau in northwestern
Michoacan agree well with the diagnosis of _Hypopachus oxyrrhinus ovis_
by Shannon and Humphrey (1958). With the exception of one specimen from
Tangamandapio, all have dark bellies extensively mottled or spotted with
cream-color. Most of the specimens have some form of an irregular,
usually broken, dark line from the occiput to the groin. In eight
specimens there is no line or linear arrangement of spots; instead the
dorsum is spotted or flecked with dark brown. The ground color of the
dorsum and flanks varies from dull reddish brown to grayish brown;
cream-colored spots are evident on the flanks and posterior surfaces of
the thighs in all specimens (Pl. 6, Fig. 2).
In comparison with 14 specimens from Queseria, Colima (UMMZ 80001-2),
individuals from the Mexican Plateau have a darker venter with bolder
markings, and a more mottled dorsum.
In Michoacan this species has been taken between 1500 and 2200 meters on
the Mexican Plateau, where it inhabits mesquite-grassland and cultivated
areas.
~Rana dunni~ Zweifel
_Rana dunni_ Zweifel, Copeia, no. 2:78, July 15, 1957.--Lago
de Patzcuaro, Michoacan, Mexico.
Lago de Patzcuaro (23); Rio de Morelia, near Undameo (8).
Aside from the type series of this species, there are in the Museum of
Zoology at the University of Michigan six specimens taken from "tanks"
at the limnological station at Patzcuaro by Paul S. Martin in 1948, and
eight specimens found in shaded ditches along the Rio de Morelia by
Robert R. Miller on April 4, 1957. The Rio de Morelia flows into Lago de
Cuitzeo; this drainage is separated from Lago de Patzcuaro by a chain of
hills about 2400 meters in elevation. Dr. Richard G. Zweifel has
examined these specimens and has informed me that, although they differ
slightly from typical _Rana dunni_, they are much closer to that species
than to _Rana montezumae_.
~Rana megapoda~ Taylor
_Rana megapoda_ Taylor, Univ. Kansas Sci. Bull., 28:310,
November 12, 1942.--Chapala, Jalisco, Mexico.
La Palma (8).
These specimens (USNM 113998-114005) are from the marshes along the
southeastern shore of Lago de Chapala. Five females have snout-vent
lengths of 124.0-138.1 (131.5), and one male has a snout-vent length of
110.2 mm. Two juveniles have snout-vent lengths of 49.7 and 56.3 mm. The
coloration of the juveniles is more bold than that of the adults. The
body proportions of these specimens agree with those presented by
Zweifel (1957:80).
~Rana montezumae~ Baird
_Rana montezumae_ Baird, Proc. Acad. Nat. Sci. Philadelphia,
7:61, October 20, 1854.--Mexico City, Distrito Federal,
Mexico.
La Palma; 8 km. NW of Maravatio (10); Sahuayo; Tupataro (7).
This species probably is more abundant and widespread than is indicated
by the few specimens listed above. It has been found only in the
vicinity of permanent water on the Mexican Plateau and the mountains
rising from the plateau at elevations of 1500 to 2000 meters. Its
apparent absence from Lago de Patzcuaro cannot be explained, unless
_Rana dunni_ replaces it there.
~Rana pipiens~ Schreber
_Rana pipiens_ Schreber, Der Naturforscher, Halle, 18:185,
1782.--Raccoon, Gloucester County, New Jersey.
Aguililla (2); Apatzingan (13); Arteaga; Axolotl (16);
Camachines (2); Capirio; Cascada Tzararacua (3); Cerro San
Andres (6); Charapendo (4); Ciudad Hidalgo; Coalcoman (17);
Cuitzeo (3); El Sabino (10); Jacona (3); 29 km. NW of Jacona
(8); Jiquilpan; La Orilla (3); La Palma (5); La Playa (4);
Lago de Chapala (3); Lago de Patzcuaro (6); Lombardia; Los
Conejos (67); Los Reyes (7); Macho de Agua; Maravatio;
Morelia (5); Opopeo (3); Patzcuaro (9); 26 km. S of
Patzcuaro (52); Puerto Hondo (3); Rio Duero, 14 km. E of
Zamora (13); Rio Tepalcatepec, 27 km. S of Apatzingan (2);
San Gregorio (38); San Jose de la Cumbre (5); Tangamandapio;
Zacapu; 18 km. W of Zamora (35).
Except on the Pacific lowlands, this species is abundant throughout the
state. It has been collected from sea level to 2800 meters, the greatest
altitudinal range of any amphibian in Michoacan. It has been found
frequently in the Tepalcatepec Valley; it is not a distinctly highland
species in southern Michoacan, as stated by Peters (1954:9). One
specimen from Aguililla (UMMZ 119257) is an albino. In this specimen
there is a faint pattern on the hind limbs; otherwise the entire body is
creamy white; the eyes are pink.
~Rana pustulosa~ Boulenger
_Rana pustulosa_ Boulenger, Ann. Mag. Nat. Hist., ser. 5,
11:343, 1883.--Ventanas, Durango, Mexico.
Arteaga (4); 21 km. S of Arteaga; Cascada Tzararacua (3);
Coalcoman (3); 12 km. ENE of Dos Aguas (3); El Sabino (53);
Los Reyes (3); Tzitzio (4); Uruapan.
Although _Rana pustulosa_ seems to be absent from the Mexican Plateau in
Michoacan, it has been collected at elevations of 850 to 2150 meters on
the slopes of the Cordillera Volcanico and in the Sierra de Coalcoman.
Usually the frogs are found along rocky streams, but at Coalcoman they
were found in a hyacinth-choked old river channel, and at El Sabino, in
irrigation ditches.
In most specimens the dorsum is dark olive-brown; in some it is pale
olive-tan with dense dark brown mottling on the back and dark transverse
bands on the hind limbs.
Thirteen tadpoles (UMMZ 94271) taken from a seepage pool by a stream
near Uruapan closely resemble the description of tadpoles of this
species given by Taylor (1942b).
REPTILIA
Testudines
~Chelonia mydas~ (Linnaeus)
_Testudo mydas_ Linnaeus, Systema naturae, ed. 10:197,
1758.--Type locality restricted to Ascension Island by
Mertens and Mueller (1928:23).
_Chelonia mydas_, Brongniart, Bull. Sci. Soc. Philom., 2:89,
1800.
Beach between Rio Motin and Rio Colotlan (2); Maruata; Playa
Azul (4).
Green sea turtles are abundant along the coast of Michoacan. Laying
females and fresh nests were found on August 6-12, 1950, July 14-16,
1951, and July 8-10, 1955. The general account of sea turtles on the
coast of Michoacan that was given by Peters (1957) is supplemented here
by my field notes on the actions of one female observed on the night of
July 14, 1951, near Maruata by Donald D. Brand and I. Because of a full
moon, visibility was excellent.
In the course of the day several _Chelonia_ were seen in the surf;
shortly after dark the first turtle was observed on the beach. Several
were observed to come out on the beach and crawl nearly to the strand
line, only to return to the sea.
At 10:20 p. m. one turtle was seen about 15 meters from the water. We
watched this turtle from some distance and observed that by 10:26 p. m.
she had moved about ten meters to a bank of sand about two meters high.
Ten minutes later she had climbed the bank and disappeared over the top
into the brush. We moved closer and remained hidden below the bank.
Although we could not see the turtle, we could hear her movements.
Between 10:37 and 10:57 p. m. the turtle dug, often flipping the dry
sand for a distance of about two meters. When this energetic digging
ceased, we moved up the bank to see that she was facing inland and
sitting in a depression about one and one-half meters in diameter and 30
centimeters in depth. She had cleaned out this depression in the past 20
minutes. Between 11:00 and 11:36 p. m. she dug the nest hole by first
scooping sand with one hind flipper and then with the other; when sand
was thrown by one flipper, there was a similar, but weaker, motion by
the other flipper. At 11:36 p. m. she stopped digging. By crawling up
behind the turtle we were able to examine the nest cavity, which
measured 21 centimeters across the top and 38 centimeters deep. The
diameter of the bottom of the hole was estimated to be about 50
centimeters. At 11:40 p. m. she released the first egg; a minute later
she dropped the second. At 11:42 p. m. the third and fourth eggs were
released; these were coherent, as were the fifth and sixth eggs released
at 11:43 p. m. After this, as many as three eggs were dropped at a
time. After laying about 60 eggs, she paused for a minute and then
continued laying. By 11:55 p. m. she had laid 98 eggs; after this, the
process of deposition slowed considerably. She dropped a fragment of an
egg followed by normal eggs. At midnight she deposited a miniature egg
about 20 mm. in diameter. This terminated the deposition. Immediately
she began to cover the nest.
Within ten minutes after the last egg was deposited the nest had been
covered. The turtle first had been seen at 10:20 p. m.; judging from its
speed and its distance from the water, the turtle probably had been on
land for about ten minutes. About 25 minutes were used in crawling from
the water to the nesting site. One hour and 33 minutes were spent at the
nesting site; of this time twenty minutes were taken for egg deposition.
The turtle was not followed back to the water, but if the return trip
took approximately the same amount of time as required to travel from
the ocean to the nesting site, the total elapsed time from departure to
return to the water was about two and one-half hours.
We collected the eggs as they were deposited. There were 106 eggs, each
having a diameter of about 40 mm., plus one small egg and a fragment of
another. The turtle had a carapace about one meter in length.
From our limited observations of sea turtles and their tracks on the
beaches, and from the accounts of these animals by the residents of the
coastal region, great numbers of sea turtles use these relatively
uninhabited beaches for nesting grounds. However, the turtles do not go
unmolested. The natives capture turtles and collect their eggs. Opened
and emptied nests also showed signs of predatory activity on the part of
other mammals. In the vicinity of Playa Azul several turtles were killed
by dogs.
~Kinosternon hirtipes hirtipes~ Wagler
_Cinosternon hirtipes_ Wagler, Naturl. Syst. Amph., p. 37,
1830.--Mexico. Type locality restricted to Mazatlan,
Sinaloa, Mexico, by Smith and Taylor (1950b:25).
_Kinosternon hirtipes hirtipes_, Schmidt, Check list N Amer.
Amph. Rept., ed. 6, p. 89, 1953.
Eight km. W of Ciudad Hidalgo; Jiquilpan; La Palma; Lago de
Camecuaro (4); Lago de Cuitzeo (3); Lago de Patzcuaro (8);
14 km. E of Zamora (4).
One specimen from eight kilometers west of Ciudad Hidalgo (UIMNH 24707)
is from the Rio Tuxpan, a tributary of the Rio Balsas; this is the only
record for the species from the Balsas drainage. All others are from the
lakes or rivers flowing into the lakes on the southern part of the
Mexican Plateau. This species exists in Lago de Patzcuaro to the
apparent exclusion of the abundant and widespread _Kinosternon
integrum_.
~Kinosternon integrum~ LeConte
_Kinosternon integrum_ LeConte, Proc. Acad. Nat. Sci.
Philadelphia, 7:183, 1854.--Mexico. Type locality restricted
to Acapulco, Guerrero, Mexico, by Smith and Taylor
(1950b:25).
Agua Cerca (3); Aguililla; Arteaga (8); Apatzingan (7);
Barranca de Herradero; Buenavista (20); Capirio (2);
Charapendo (3); Chupio; Coahuayana (2); Coalcoman (169);
Copuyo (4); El Sabino (8); Jacona; Jiquilpan (12); La Orilla
(2); La Playa (2); Lago de Cuitzeo (27); Las Higuertas;
Lombardia (3); Los Reyes (5); Morelia; Ojos de Agua de San
Telmo; San Pedro Naranjestila; Tacicuaro.
Excepting Lago de Patzcuaro, _Kinosternon integrum_ occupies all
permanent and temporary ponds, lakes, and streams below 2200 meters
throughout the state. At Coalcoman the species was in roadside ditches,
small puddles, flooded fields, a hyacinth-choked ox-bow of the Rio
Coalcoman, as well as in the Rio Coalcoman and its tributaries.
Specimens from Arteaga and Barranca de Herradero were found in clear
rocky streams; the one from Las Higuertas was found in a small muddy
pond in pine-oak forest.
On August 26, 1960, James R. Dixon found a copulating pair in a pool at
Capirio. The large series from Coalcoman contains juveniles and adults;
these turtles formed the basis for the study of relative growth of
plastral scutes in this species by Mosimann (1956).
~Geoemyda rubida perixantha~ Mosimann and Rabb
_Geoemyda rubida perixantha_ Mosimann and Rabb, Occ. Pap.
Mus. Zool. Univ. Michigan, 548:1, November 9, 1953.--Eight
kilometers south of Tecoman, Colima, Mexico.
Apatzingan (2); Coahuayana; La Placita; Punta San Juan de
Lima.
These specimens have been discussed in detail by Mosimann and Rabb
(1953). All are from the arid tropical scrub forest; those from the
coastal regions were collected at elevations of less than 40 meters, and
those from the Tepalcatepec Valley were collected at an elevation of 335
meters.
Crocodilia
~Crocodylus acutus acutus~ Cuvier
_Crocodylus acutus_ Cuvier, Ann. Mus. Hist. Nat. Paris,
10:55, 1807.--Santo Domingo.
_Crocodylus acutus acutus_, Mueller and Hellmich,
Ibero-Amerik. Stud., 13:128, 1940.
Boca de Apiza (2); Playa Azul (2).
The crocodile or "caiman" is abundant in the brackish lagoons along the
cost of Michoacan; three large adults and several juveniles were
observed at Estero Pichi at Playa Azul; others were seen at Mexiquillo
and Maruata. Residents of the Balsas-Tepalcatepec Basin frequently have
reported "caimanes" in the Rio Balsas and Rio Tepalcatepec, but the
existence of the crocodile in these rivers has not been verified by
specimens.
Sauria
~Phyllodactylus duellmani~ Dixon
_Phyllodactylus duellmani_ Dixon, Southwest Nat., 5:37,
April 15, 1960.--Rancho El Espinal, Michoacan, Mexico.
Fourteen km. SSW of Apatzingan; Capirio; Cafradia (3); El
Espinal (3).
This species is known only from the Tepalcatepec Valley, where it has
been found in open arid situations from 180 to 500 meters. Specimens
were found in the daytime in stumps, dead cacti, and the hollow branches
of the legume, _Apoplanesia paniculata_. In life adults were pale gray
or grayish tan above and creamy white below. A juvenile having a
snout-vent length of 18 mm. had a pale orange tail with gray
cross-bands. In the adults the tail was colored like the body. The
specimen from 14 kilometers south-southwest of Apatzingan (KU 29764) and
those from Cofradia (BMNH 1914.1.28.28-30) were not listed by Dixon
(1960).
~Phyllodactylus homolepidurus~ Smith
_Phyllodactylus homolepidurus_ Smith, Univ. Kansas Sci.
Bull., 22:121, November 15, 1935.--Five miles southwest of
Hermosillo, Sonora, Mexico.
El Ticuiz (2); La Placita; Ostula (2); Pomaro; San Pedro
Naranjestila.
These specimens have been referred to _Phyllodactylus homolepidurus_ by
James R. Dixon (_in litt._), who is currently studying the American
members of the genus. Geckos of this species have been found in tropical
semi-deciduous forest in the coastal lowlands to elevations of 500
meters. Most specimens were found beneath the bark of standing dead
trees or stumps. Two individuals from El Ticuiz (UMMZ 115102) in life
were dark gray above with brownish tubercles; the belly was a dusty
cream-color. Apparently this species does not enter the Tepalcatepec
Valley, where _Phyllodactylus lanei_ is abundant.
~Phyllodactylus lanei~ Smith
_Phyllodactylus lanei_ Smith, Univ. Kansas Sci. Bull.,
22:125, November 15, 1935.--Tierra Colorado, Guerrero,
Mexico.
Apatzingan (13); 21 km. S of Arteaga: El Sabino (53); La
Playa; Ostula (2); Rio Marquez, 10 km. S of Lombardia (8);
16 km. N of Tafetan.
This widespread species has been taken at elevations of less than 1100
meters in the Balsas-Tepalcatepec Basin, where it occurs in riparian
situations in the foothills. Specimens have been collected in tropical
semi-deciduous forest at Ostula and in oak forest south of Arteaga; both
of these localities are on the Pacific slopes of the Sierra de
Coalcoman, a region inhabited by _Phyllodactylus homolepidurus_. Both
species have been collected at Ostula.
A juvenile from 21 kilometers south of Arteaga (UMMZ 118933) had
alternating black and white bands on the tail. In life most of the
lizards are dull ashy gray or grayish tan above and white below.
According to Dixon (_in litt._), one specimen from Apatzingan (UMMZ
115102) resembles _Phyllodactylus magnus_ in scutellation, but it lacks
the distinctive yellow venter of that species.
Apparently _Phyllodactylus lanei_ is restricted to rather mesic
environments in the Balsas-Tepalcatepec Valley and surrounding
foothills; in the more open arid environments on the floor of the valley
it seems to be replaced by _Phyllodactylus duellmani_.
~Phyllodactylus paucituberculatus~ Dixon
_Phyllodactylus paucituberculatus_ Dixon, Southwest. Nat.,
5:40, April 15, 1960.--Rio Cupatitzio (= Rio Marquez), 6.5
miles south of Lombardia, Michoacan, Mexico.
Rio Marquez, 10 km. S of Lombardia (6).
Two of these specimens (UMMZ 112692-3) were discussed in detail by Dixon
(1960:40) in his description of the species. On August 25, 1960, Dixon
collected four additional specimens at the type locality, a conglomerate
cliff along the Rio Marquez. These will be reported by him in his
forthcoming study of the genus.
~Anolis dunni~ Smith
_Anolis dunni_ Smith, Copeia, no. 1:9, May 10, 1936.--Agua
del Obispo, Guerrero, Mexico.
Arteaga (3); 19 km. S of Arteaga.
Three females from Arteaga (UMMZ 119075) have snout-vent lengths of 41,
41, and 44 mm. In life the pale grayish brown dorsum was marked with
dark brown; the belly was white, and the throat was pale pink. All have
a dark interorbital bar and dark vertical bars on the upper labials. In
two specimens there are only scattered dark flecks on the dorsum; in the
third there is a dark postorbital stripe, a dark lateral stripe, and
four narrow transverse bands on the body. A male from 19 kilometers
south of Arteaga (UMMZ 119076) having a snout-vent length of 49 mm. had
in life a tan dorsum, a broad white stripe from the ear to the groin,
scattered small white spots on the dorsum, and indistinct pale
cream-colored spots on the posterior surfaces of the thighs. This male
has the dark labial bars, but lacks the dark interorbital bar, found in
the females. The large rose-pink throat fan extends to about the middle
of the belly. In all of the specimens the middorsal scales are keeled
and much smaller than the smooth pavementlike or slightly imbricate
ventrals. All have two gulars in contact with the mental, five scales
between the nasals, five scales (not including the first labials) in
contact with the rostral, and four rows of loreals. In these characters
these specimens agree well with _Anolis dunni_ from Guerrero, as
diagnosed by Davis (1954b).
Previously _Anolis dunni_ has been reported only from the vicinity of
Agua del Obispo, Guerrero, a locality situated at an elevation of about
900 meters in pine-oak forest in the Sierra del Sur. All known close
relatives of _Anolis dunni_ occur only in Guerrero: _A. taylori_ Smith
and Spieler from Acapulco, _A. gadowi_ Boulenger from Tierra Colorado,
_A. liogaster_ Boulenger, and _A. omiltemanus_ Davis from Omiltemi. The
present specimens from elevations of about 900 meters in riparian stream
vegetation and oak forest represent the northern known limits of this
group of _Anolis_.
~Anolis nebulosus~ (Wiegmann)
_Dactyloa nebulosa_ Wiegmann, Herpetologia Mexicana, p. 47,
1834.--Mexico. Type locality restricted to Mazatlan,
Sinaloa, Mexico, by Smith and Taylor (1950b:66).
_Anolis nebulosas_, Bocourt, Mission Scientifique au Mexique
et dan l'Amerique Centrale. Reptiles, livr. 2:77, 1873.
Acahuato (3); Agua Cerca; Apatzingan (4); Araparicuaro (3);
29 km. S of Ario de Rosales (3); 20 km. S of Arteaga (2);
Barranca de Bejuco; Cascada Tzararacua (5); Cerro Tancitaro
(13); Cheran; Chupio (5); Coalcoman (10); Cofradia; Dos
Aguas (10); 18 km. E of Dos Aguas (3); El Diezmo; El Sabino
(43); El Ticuiz; Jiquilpan (2); La Orilla; La Placita; La
Playa (3); Los Conejos (2); Los Pozos; Nogueleras (2);
Ostula; 8 km. W of Patzcuaro (2); 8 km. NE of Patzcuaro;
Playa Azul (3); Rio Cachan; Rio Marquez, 10 km. S of
Lombardia; Rio Tepalcatepec, 27 km. S of Apatzingan; San
Juan de Lima (6); San Pedro Naranjestila; Temazcal; Tuxpan
(2); Tzitzio; Uruapan (74); 11 km. N of Uruapan (2); Volcan
Jorullo; 16 km. E of Zacapu (2); 18 km. W of Zamora;
Ziracuaretiro.
Even with the abundance of material the assignment of a specific name to
these anoles is only tentative, for definite determination between
_Anolis nebulosus_ Wiegmann and _A. nebuloides_ Bocourt is uncertain.
Bocourt (1873:75) distinguished _A. nebuloides_ from _A. nebulosus_ by
the following characters: (1) head scales keeled, not smooth; (2) snout
narrower; (3) ear opening larger; (4) supraorbital semicircles separated
by a row of small scales and not in contact; (5) dorsal scales larger
and subequal in size to the belly scales. Boulenger (1885:77) used the
same characters; Smith and Taylor (1950b:58) in their key to the Mexican
species of _Anolis_ stated that the dorsal scales are slightly smaller
than the ventrals in _A. nebulosus_ and markedly smaller in _A.
nebuloides_. Smith (_in litt._) stated that the characters of the
relative sizes of the dorsal and ventral scales were incorrect in that
key.
The application of the above criteria to specimens from Michoacan has
not resulted in the recognition of two species. The majority of the
specimens have the supraorbital semicircles separated by at least one
small scale; the head scales, with the exception of those on the snout
in a few individuals, are smooth; the dorsal scales are only slightly
smaller than the ventrals. In other characters of scutellation the
specimens are highly variable. The males in life have an orange throat
fan. Anoles of this kind have been found in Michoacan, Colima, Jalisco,
Nayarit, and southern Sinaloa. Near Oaxaca, Oaxaca, specimens were
collected that superficially resemble those from Michoacan and farther
north. These have low keels on the snout scales, dorsals somewhat larger
than the ventrals, and a pink throat fan. In ten males from Oaxaca the
size of the dorsal scales relative to that of the ventrals is 1.00:0.83;
the same ratio for 25 males from Michoacan is 1.00:1.08. In both samples
there are specimens in which the dorsal and ventral scales are about
equal in size.
Investigations by Richard E. Etheridge on the osteology of _Anolis_,
including those species here being considered, have revealed relatively
constant differences in the parasternalia and in the caudal vertebrae.
The application of Etheridge's findings to anoline systematics must
await the completion of his study.
The carination of the scales on the snout _versus_ smooth scales there
seems to be the only significant character given by Bocourt that
distinguishes _A. Nebuloides_ from _A. nebulosus_. The difference in the
color of the throat fan, which is apparent only in living individuals,
is more striking. Obviously more than one species is represented, as is
borne out by the differences in the color of the throat fan and in the
osteology, but there is uncertainty about the correct name for each
species. On the strength of Bocourt's diagnosis of keeled snout scales
in _A. nebuloides_, I am applying that name to the population in Oaxaca
and _A. nebulosus_ to the specimens from Michoacan. As arranged here,
the two species can be distinguished, as follows:
_A. nebulosus._--Dorsal scales only slightly smaller than
the ventral scales; snout scales usually smooth; throat-fan
bright orange in adult males.
_A. nebuloides._--Dorsal scales somewhat larger than the
ventral scales; snout scales having a low keel; throat-fan
pink in adult males.
With respect to geographic distribution, _A. nebulosus_ has been
collected from southern Sinaloa southward to Michoacan. The lizards here
referred to _A. nebuloides_ have been taken only in pine-oak forest on
the mountain slopes near Oaxaca City. Zweifel and Norris (1955:233)
reported anoles with pink throat-fans from southern Sonora; possibly
those specimens are _A. nebuloides_; I have not examined them. I have
seen several preserved specimens from the vicinity of Tehuantepec,
Oaxaca. Although they probably belong to this group, those specimens
differ from both _A. nebulosus_ and _A. nebuloides_ in their larger
size, relatively larger head, and much larger throat fan.
Aside from the minor variation in scutellation, specimens of _Anolis
nebulosus_ from Michoacan vary greatly in coloration. Usually the
females have some form of a broad middorsal pale-colored band. In life
this is dull yellow, tan, or orange. Two females from Dos Aguas are
strikingly different; one (UMMZ 119521) has a broad middorsal orange
stripe that is scalloped laterally and bordered by gray. The other (UMMZ
119081) has a narrow middorsal cream-colored line. Males usually are
unicolor brown or olive-tan; sometimes the middorsal region is darker.
Some individuals have dark cross-bands or chevrons on the dorsum. One
male from Dos Aguas (UMMZ 119080) has a cream-colored lateral stripe.
In Michoacan _Anolis nebulosus_ occurs from sea level to elevations
slightly in excess of 2100 meters, usually in areas of dense cover,
whether this be herbaceous, viney, or woody, ordinarily on the ground as
well as in bushes and trees. One was in a bromeliad growing about ten
meters above the ground. In the arid Tepalcatepec Valley anoles of this
species are most frequently found in the tangled growth along streams.
Above Uruapan they were found in pine-oak forest, and on the Mexican
Plateau between Zamora and Zacapu they were found in a bunch grass-scrub
oak association.
~Anolis schmidti~ Smith
_Anolis schmidti_ Smith, Publ. Field Mus. Nat. Hist., zool.
ser., 24:21, January 30, 1939.--Manzanillo, Colima, Mexico.
La Placita; San Juan de Lima.
Peters (1954:11) reported on the specimen from La Placita; another was
secured at San Juan de Lima in 1956. The latter (UMMZ 115078) is a male
having a snout-vent length of 43.0 mm. and a tail length of 70.5 mm. The
dorsal ground color is pale tan; there are five pairs of irregular dark
brown dorsolateral blotches. In life the throat fan was pale orange.
These specimens agree with those from Colima described by Duellman
(1958c:10). The distribution of _Anolis schmidti_ seems to be restricted
to the coastal lowlands from Michoacan to Nayarit.
~Basiliscus vittatus~ Wiegmann
_Basiliscus vittatus_ Wiegmann, Isis von Oken, 21:373,
1828.--Mexico. Type locality restricted to Veracruz,
Veracruz, Mexico, by Smith and Taylor (1950b:72).
Apatzingan (9); Capirio; Coahuayana (5); El Cerrito; El
Sabino (2); El Ticuiz; La Placita (3); Maruata (2); Motin
del Oro; Ostula; Playa Azul (3).
This species has been found only on the coast and in the low
Tepalcatepec Valley. In the latter area it is restricted to riparian
situations along the larger streams. The lizard is abundant in the
mangrove swamps bordering the brackish lagoons on the coast. In July,
1955, scores of individuals were seen around Estero Pichi at Playa Azul.
Adults, especially the large males, are exceedingly wary and difficult
to collect. At all localities where they were found, the lizards were
most often seen in dense bushes, where they are well camouflaged.
Individuals of all sizes were observed to run across the surface of the
ponds.
~Iguana iguana rhinolopha~ Wiegmann
_Iguana rhinolopha_ Wiegmann, Herpetologia Mexicana, p. 44,
1834.--Mexico. Type locality restricted to Cordoba,
Veracruz, Mexico, by Smith and Taylor (1950b:72).
_Iguana iguana rhinolopha_, Van Denburgh, Proc. Acad. Nat.
Sci. Philadelphia, 1897:461, January 18, 1898.
Apatzingan (8); Capirio (3); El Cerrito; El Ticuiz (2); La
Placita; La Playa (2); Maruata; Playa Azul; Rio Cachan.
Like the preceding species, this lizard is always found near water. It
does not ascend the foothills of the Sierra de Coalcoman, but in the
Balsas Basin it reaches elevations of 800 meters at La Playa. Large
adults are often seen in the large trees making up the gallery forests
along rivers. From high perches the lizards drop into the water with a
terrific splash. Bright green juveniles were abundant in bushes along
the Rio Tepalcatepec in July, 1955.
~Ctenosaura pectinata~ (Wiegmann)
_Cyclura pectinata_ Wiegmann, Herpetologia Mexicana, p. 42,
1834.--Mexico (by inference). Type locality restricted to
Colima, Colima, Mexico, by Bailey (1928:25).
_Ctenosaura pectinata_, Gray, Catalogue of the lizards...
British Museum, p. 191, 1845.
Apatzingan (27); between Ario de Rosales and La Playa;
Barranca de Bejuco; Capirio (2); Coalcoman (4); El Espinal;
El Sabino (2); El Ticuiz; Jazmin (2); La Huacana; La Placita
(8); La Playa (3); Limoncito; Lombardia; Motin del Oro;
Playa Azul; Rio Cancita, 12 km. E of Apatzingan (2); Rio
Marquez, 10 km. S of Lombardia (2);? Uruapan; Volcan
Jorullo.
_Ctenosaura pectinata_ is a common lowland species that ascends the
slopes of the Sierra de Coalcoman and the Cordillera Volcanica to
elevations of about 1050 meters (approximating the lower limits of the
oak forest). The record from Uruapan (USNM 10234, collected by Duges) is
doubtful.
These large lizards are most easily observed on rock fences along roads.
Near Apatzingan innumerable individuals can be seen in mid-morning.
Later in the day, as the sun rises higher in the sky, the lizards
retreat to the shade of the crevices in the fences. The abundance of
these lizards in the Tepalcatepec Valley, together with evidence
gathered from the natives of the valley, indicates that these lizards
are seldom used for human consumption there. On the other hand, several
people in Coalcoman consider the "iguana negra" (local name for
_Ctenosaura_) to be a delicacy and serve it at every opportunity. In
early July, 1951, brilliant green young of the year were collected at La
Playa and at Coalcoman.
~Enyaliosaurus clarki~ (Bailey)
_Ctenosaura clarki_ Bailey, Proc. U. S. Natl. Mus., 73:44,
September 26, 1928.--Ovopeo (= Oropeo), Michoacan, Mexico.
_Enyaliosaurus clarki_, Duellman and Duellman, Occ. Pap.
Mus. Zool. Univ. Michigan, 598:1, February 16, 1959.
Twelve km. SSW of Apatzingan; Capirio (7); Cofradia (3); El
Espinal (2); 32 km. E of Huetamo; Jazmin (5); Oropeo (10);
Rancho Nuevo; Rio Cancita, 12 km. E of Apatzingan (8);
Tepalcatepec (3); Zicuiran (6).
This species is known only from the low areas of the Balsas-Tepalcatepec
Basin between elevations of 200 and 510 meters. It is commonly found in
the open arid tropical scrub forest dominated by _Prosopsis_ sp.,
_Apoplanesia paniculata_, and _Cercidium plurifoliolatum_. Continued
collecting in the Tepalcatepec Valley has borne out the suggestions of
Duellman and Duellman (1959) concerning the distribution and abundance
of this lizard. Also, continued collecting in Colima and on the Pacific
coast has failed to reveal the presence of _Enyaliosaurus_ there.
~Phrynosoma asio~ Cope
_Phrynosoma asio_ Cope, Proc. Acad. Nat. Sci. Philadelphia,
16:178, September 30, 1864.--Colima, Colima, Mexico.
Apatzingan (4); San Salvador.
In Michoacan this species has been obtained only in the Tepalcatepec
Valley and on the northern slopes of the Sierra de Coalcoman between 300
and 700 meters. Apparently the lizard is absent from the coastal
lowlands of Michoacan and Guerrero. The distribution of this species,
therefore, is discontinuous. One population inhabits the lowlands of
Colima and the Balsas-Tepalcatepec Basin inland to northern Guerrero and
Morelos; a southern population inhabits the Plains of Tehuantepec in
Oaxaca.
A juvenile from Apatzingan (USNM 47739) has a snout-vent length of 40.0
mm. and a tail length of 19.5 mm.
~Sceloporus aeneus aeneus~ Wiegmann
_Sceloporus aeneus_ Wiegmann, Isis von Oken, 21:370,
1828.--Mexico. Type locality restricted to Tres Cumbres,
Morelos, Mexico, by Smith and Taylor (1950b:137).
_Sceloporus aeneus aeneus_, Smith, Occ. Pap. Mus. Zool.
Univ. Michigan, 361:6, December 15, 1937.
Angahuan; Araparicuaro (2); Capacuaro (2); Carapan (11);
Cheran (11); 18 km. WNW of Ciudad Hidalgo (10); Cuseno
Station; Jerahuaro; Los Conejos (36); Macho de Agua (7);
Opopeo; Paracho (2); Patzcuaro (4); Pino Gordo; 18 km. W of
Quiroga (2); Tancitaro (49); Uruapan (14); 16 km. NW of
Zacapu (5); between Zacapu and Zamora (2); 13 km. E of
Zinapecuaro; 14 km. SE of Zitacuaro (14).
This small terrestrial species inhabits the pine and fir forests of the
Cordillera Volcanica between elevations of 1850 and 3100 meters;
apparently it is absent from the Sierra de Coalcoman. It seems to prefer
rather open coniferous forests in which there is a more or less
continuous cover of grasses on the ground. On warm sunny days the
lizards can be observed scurrying about in the grass; in the early hours
of the day, or on cold days, they are found beneath stones, logs, or
dead clumps of bunch grass.
~Sceloporus asper~ Boulenger
_Sceloporus asper_ Boulenger, Proc. Zool. Soc. London,
1897:497, October, 1897.--La Cumbre de los Arrastrados,
Jalisco, Mexico.
Apatzingan (3); 10 km. E of Dos Aguas; Uruapan (41).
This strictly arboreal lizard is abundant in the mixed broad-leafed
forest near Uruapan. The lizards are exceedingly wary and can be
approached only with difficulty. In life males have pale blue bellies;
the throat is pale pink. The pale gray dorsum marked with irregular
darker gray blotches blends well with the color of the tree trunks on
which the lizard lives. The one specimen from Dos Aguas was found on a
pine tree; it provides the only record for the species from the Sierra
de Coalcoman.
~Sceloporus bulleri~ Boulenger
_Sceloporus bulleri_ Boulenger, Proc. Zool. Soc. London,
1894:729, April, 1895.--Las Cumbre de los Arrastrados,
Jalisco, Mexico.
Acuaro de las Lleguas (13); Barolosa (9); Dos Aguas (61); 10
km. NE of Dos Aguas (5).
Heretofore this species has been known only from a few specimens from
scattered localities in the Sierra Madre Occidental in southwestern
Jalisco and Sinaloa. The collection of a large series of these lizards
in virgin pine forest at elevations of more than 2000 meters in the
Sierra de Coalcoman now makes possible an analysis of variation in the
species.
Superficially _S. bulleri_ resembles _S. torquatus_, but _S. bulleri_ is
smaller, has more dorsal scales, fewer scales in the dark collar, and
fewer femoral pores. In 88 specimens of _S. bulleri_ there are 36-41
(38.7) dorsal scales and 2 or 3 (2.6) middorsal scales in the collar, as
compared with 28-31 (29.3) dorsal scales and 3 or 4 (3.4) middorsal
scales in the collar of 26 specimens of _S. torquatus_ from Uruapan. In
20 adult males of _S. bulleri_ there are 13-15 (14.3) femoral pores, and
13-16 (14.4) in 11 females; 13 males of _S. torquatus_ have 14-21 (17.3)
femoral pores, and 13 females have 15-21 (16.7). Seventeen adult males
of _S. bulleri_ have snout-vent lengths of 72-91 (82.0); ten females,
71-87 (75.7). In comparison, 13 adult males of _S. torquatus_ have an
average snout-vent length of 88.9 mm., and 13 females, 88.5 mm. In _S.
bulleri_ there is little variation in the head scales. The frontal is in
contact with the interparietal in 63, and not in 24, specimens; the
median frontonasal is in contact with the frontal in 13, and not in 74,
specimens. In 39 specimens there are two canthals, and in 48 there is
one; in 29 specimens there are three preauriculars, and in 58 there are
four.
In life adult males have a pale blue tail, bright blue belly patches, a
purplish blue throat, and pale blue lines on the sides of the head and
neck.
This species was obtained at four localities in the high mountains of
the Sierra de Coalcoman. In this mountain range _Sceloporus bulleri_
apparently replaces _S. torquatus_, a species that is widespread in the
Cordillera Volcanica and on the Mexican Plateau. At Dos Aguas and at
Acuaro de las Lleguas the lizards were abundant in the tall pine forest,
where they were found on standing pine trees, on pine logs, and on rock
outcroppings.
~Sceloporus dugesi intermedius~ Duges
_Sceloporus intermedius_ Duges, La Naturaleza, 4:29,
1877.--La Noria, near Zamora, Michoacan, Mexico.
_Sceloporus dugesii intermedius_, Smith, Univ. Kansas Sci.
Bull., 24:663, February 16, 1938.
Cojumatlan (6); Jiquilpan (11); Lago de Camecuaro; Lago de
Chapala; Morelia (23); Patzcuaro (84); Quiroga (35); Sahuayo
(4); Tacicuaro (2); Tangamandapio (17); Tangancicuaro (9);
Zacapu (4); Zamora (11); Zinapecuaro (9).
This lizard is strictly an inhabitant of the Mexican Plateau, where it
is found in rocky places, sometimes in pine-oak forest, but more
frequently in mesquite-grassland. It is a terrestrial species, and is
most often seen on rock fences at elevations of 1500 to 2200 meters.
This species differs from _S. bulleri_ and _S. torquatus_ in having two
rows of supraoculars, instead of one; also it has more dorsal scales.
Twenty-six specimens of _Sceloporus dugesi intermedius_ from
Tangamandapio and Tangancicuaro have 44-48 (45.7) dorsal scales, as
compared with an average of 38.7 in _S. bulleri_ and 29.3 in _S.
torquatus_. In life _Sceloporus dugesi intermedius_ has a dull greenish
gray dorsum; in males the belly patches are bright blue bordered
medially by black, and the throat is bluish gray. The largest specimen
examined is a male having a snout-vent length of 80 mm.
~Sceloporus gadowae~ Boulenger
_Sceloporus gadoviae_ Boulenger, Proc. Zool. Soc. London,
1905, 2:246, October 7, 1905.--Mezquititlan, Guerrero,
Mexico.
Chupio; El Sabino (77); La Playa (6); Rio Marquez, 10 km. S
of Lombardia (11).
Although this species has a rather extensive range in the
Balsas-Tepalcatepec Basin in the state of Michoacan, Guerrero, Morelos,
and Puebla, it is only locally abundant in that area. Usually these
lizards are found on rocky cliffs in which there are many crevices for
cover. _Sceloporus gadowae_ is abundant on a conglomerate cliff along
the Rio Marquez south of Lombardia. Although the closely related _S.
pyrocephalus_ is abundant in the stream valley and in the hills above
the cliff, _S. gadowae_ has been found only on the cliff; few
individuals of _S. pyrocephalus_ have been observed on the cliff. A
similar situation was discovered on a much more extensive conglomerate
cliff along the Rio Balsas near Mexcala, Guerrero. Near Tehuitzingo,
Puebla, where _S. pyrocephalus_ was not found, _S. gadowae_ was found on
conglomerate cliffs. Probably there is strong competition between the
two species; possibly this has resulted in the restriction of _S.
gadowae_ to isolated cliff-habitats within the extensive range of the
more widespread _S. pyrocephalus_.
In Michoacan _Sceloporus gadowae_ has been found along the lower slopes
of the Cordillera Volcanica at elevations from 250 to 1050 meters. All
of the localities from which this lizard is known lie in the arid
tropical scrub forest.
~Sceloporus grammicus microlepidotus~ Wiegmann
_Sceloporus microlepidotus_ Wiegmann, Herpetologia Mexicana,
p. 51, 1834.--Mexico. Type locality restricted to Mexico,
Distrito Federal, by Smith and Taylor (1950b:120).
_Sceloporus grammicus microlepidotus_, Smith and Laufe,
Trans. Kansas Acad. Sci., 48:332, December, 1945.
Angahuari; Apo (10); Atzimba (3); Carapan (5); Cerro San
Andres (17); Cerro Tancitaro (18); Corupu; Cuseno Station
(2); Jacona; Jerahuaro (10); Macho de Agua; Mil Cumbres; 46
km. E of Morelia; 60 km. E of Morelia (2); Opopeo (14);
Patzcuaro (30); Puerto Hondo (19); San Gregorio (41); San
Jose de la Cumbre (8); Sierra Patamba; Tancitaro (233);
Tupataro; Undameo; Uruapan (180); between Zacapu and Zamora;
24 km. SE of Zitacuaro; between Zurumbeneo and Cerro
Garnica.
This small species of _Sceloporus_ is an ubiquitous inhabitant of the
coniferous forests from 1550 to 3100 meters in the Cordillera Volcanica.
Usually it is seen on tree trunks, but occasionally on the ground. Near
the lower limit of the altitudinal distribution of the species, as at
Uruapan, individuals sometimes are found on broad-leafed trees.
Apparently _Sceloporus heterolepis_ replaces _S. grammicus
microlepidotus_ in the Sierra de Coalcoman.
~Sceloporus heterolepis~ Boulenger
_Sceloporus heterolepis_ Boulenger, Proc. Zool. Soc. London,
1894:731, April, 1895.--La Cumbre de los Arrastrados,
Jalisco, Mexico.
Araparicuaro; Cerro Barolosa (6); Dos Aguas (13); Los
Conejos; 11 km. N of Uruapan (3).
Although Michoacan has not previously been included in the range of this
lizard, it was first collected in the state by Gadow in 1908 (BMNH
1914.1.28.69 from Araparicuaro). The description of _S. heterolepis_
given by Smith (1939:197) can be supplemented by data on the 23
specimens now in the collections of the Museum of Zoology at the
University of Michigan. All have two canthals; there are 55 to 71 (63.6)
scales in the middorsal row; 1 to 3 rows middorsally are somewhat
enlarged and bordered on either side by a row of larger scales bearing
high keels. There are 14 to 20 (16.2) femoral pores. Eight adult males
have snout-vent lengths from 49 to 61 (58.0) mm. and tail lengths from
57 to 74 (66.0) mm.; four adult females have snout-vent lengths from 52
to 57 (55.2) mm. and tail lengths from 60 to 66 (63.5) mm. The smallest
of eight juveniles has a snout-vent length of 28 mm. and a tail length
of 32 mm. The dorsum in adults is pale grayish brown; there are three
irregular chevron-shaped dark marks and a triangular dark brown mark
above the insertion of the hind limbs; on the tail are dark brown rings.
There are scattered faint blue flecks on the flanks and narrow
transverse dark lines on the lower limbs. Males have pale bluish green
belly patches and an orange-salmon-colored throat; the belly in females
is pale orange-tan. The juveniles have a more contrasting color pattern;
the dark chevrons on the dorsum are bordered posteriorly by pale gray.
In Michoacan this species has been obtained in pine and pine-fir forests
from 1800 to 2700 meters. On Cerro Barolosa and at Dos Aguas, both in
the Sierra de Coalcoman, the lizards were found beneath the bark of
dead, standing pines. In the Sierra de Coalcoman _Sceloporus
heterolepis_ seems to fill the niche of the small arboreal _Sceloporus_
in the coniferous forest in southwestern Mexico, a position held by _S.
grammicus microlepidotus_ in the Cordillera Volcanica; the latter
species does not occur in the Sierra de Coalcoman. Five specimens of
_Sceloporus heterolepis_ are known from the Cordillera Volcanica,
whereas 603 of _S. grammicus microlepidotus_ have been collected there.
The ecological relationships that exist between the two species in the
Cordillera Volcanica are not known.
Insofar as is known, _Sceloporus heterolepis_ reaches the southern
limits of its range in the Sierra de Coalcoman and in the western part
of the Cordillera Volcanica. Other records for the species are from the
Sierra Madre Occidental in Jalisco. Langebartel (1959) described
_Sceloporus shannonorum_ from the mountains near the Durango-Sinaloa
border; the single specimen of _S. shannonorum_ differs significantly
from _S. heterolepis_ only in having fewer dorsal scales (48). The
acquisition of additional material, especially from Nayarit and northern
Jalisco, probably will provide a basis for showing that these two
populations are conspecific.
~Sceloporus horridus oligoporus~ Cope
_Sceloporus oligoporus_ Cope, Proc. Acad. Nat. Sci.
Philadelphia, 16:177, September 30, 1864.--Colima, Colima,
Mexico.
_Sceloporus horridus oligoporus_, Taylor, Univ. Kansas Sci.
Bull., 24:520, February 16, 1938.
Aguililla; Apatzingan (50); Arteaga (2); Capirio (2);
Cascada Tzararacua; Charapendo (4); Coahuayana (3);
Coalcoman (32); 19 km. S of Corralito; 27 km. E of Dos
Aguas; El Sabino (55); El Ticuiz; Huetamo (2); Jazmin;
Jungapeo (2); La Orilla (2); La Placita; Limoncito (3);
Playa Azul (5); Tzitzio (8); Uruapan (4); Volcan Jorullo
(2); Ziracuaretiro; Zirimicuaro (13).
All of the specimens from Michoacan seem to be typical _S. horridus
oligoporus_; none has more than six femoral pores.
Characteristically this species is found in open arid scrub forest; it
reaches its greatest abundance in rocky areas in which there are
scattered leguminous trees and bushes. It has been found in these low
trees and bushes almost as frequently as it has been found on the
ground; none has been seen in large trees or far above the ground.
Altitudinally, this species ranges from sea level to about 1600 meters.
~Sceloporus melanorhinus calligaster~ Smith
_Sceloporus melanorhinus calligaster_ Smith, Proc. U. S.
Natl. Mus., 92:360, November 5, 1942.--Acapulco, Guerrero,
Mexico.
Aguililla; Apatzingan (18); Barranca de Herradero; Capirio
(19); Coahuayana (4); Coalcoman (2); Cofradia (4); El
Cerrito; El Sabino (33); El Ticuiz (3); La Placita (6);
Lombardia (4); Playa Azul; Rio Marquez, 10 km. S of
Lombardia (2); Rio Marquez, 13 km. SE of Nueva Italia (4);
Salitre de Estopila; San Juan de Lima (2); Santa Ana;
Tzitzio; Ziracuaretiro.
Smith (1942a:360) diagnosed _Sceloporus melanorhinus calligaster_ as
having fewer femoral pores than the other subspecies of _S.
melanorhinus_ and as having the lateral belly patches in the males
confluent in the midline. Examination of forty specimens from the
Tepalcatepec Valley and the coastal regions of Michoacan does not
substantiate this diagnosis. The number of femoral pores varies from 15
to 22 (18.9); 14 individuals (35%) had 20 or more femoral pores. Smith
(_loc. cit._) stated that _S. melanorhinus_ in Oaxaca had 18 to 27
(21.6) femoral pores and that 77 per cent of the specimens had more than
20 femoral pores. Of the 24 males examined from Michoacan, 18 have the
lateral belly patches separated in the midline. Usually they are
separated by no more than one scale, but in some individuals they are
separated by two or more scales. Although the above data minimize
certain differences between the northern and southern populations of
this species, certain of the color pattern characters seem to be
diagnostic of the subspecies inhabiting the Pacific lowlands from
Guerrero to Nayarit. Large adults of _S. m. calligaster_ have only a
faint dorsal pattern, which in the subspecies _melanorhinus_ and
_stuarti_ consists of a series of large, dark, interconnected triangles
on the back. This pattern is present in young and small adults of _S. m.
calligaster_; furthermore, in this subspecies the ventral coloration of
the males differs from that found in the more southern populations.
Adult males of _S. m. calligaster_ have a black throat, that changes to
brilliant blue posteriorly, and a large white spot medially on the chin.
This spot is present in some specimens from Oaxaca and Chiapas, but, if
present, it is much smaller and less distinct than in specimens from
Michoacan. In _S. m. calligaster_ the chest and midventral area are
orange to salmon-color.
A male from Lombardia in life was colored as follows: Dorsum grayish tan
bearing faint bluish gray flecks; chest deep salmon-orange, this color
continuing down midventral area to the somewhat paler groin; belly
patches pale blue fading to pale green laterally; throat black
anteriorly enclosing a white spot; throat blue posteriorly and bluish
green posterolaterally.
Individual lizards were observed to change in dorsal color from a pale
ashy gray to a rather dull brown. Normally, inactive individuals and
those observed on overcast days were dull brown.
_Sceloporus melanorhinus calligaster_ is found in trees in riparian
situations in the lowlands to elevations of about 1500 meters. It does
not inhabit the arid tropical scrub forest in the Tepalcatepec Valley or
on the coast, but in those areas is found in the gallery forests along
streams and rivers. The lizards are wary and live high in the trees;
they are especially difficult to locate in the rainy season, when the
trees are in full leaf.
~Sceloporus pyrocephalus~ Cope
_Sceloporus pyrocephalus_ Cope, Proc. Acad. Nat. Sci.
Philadelphia, 16:177, September 30, 1864.--Colima, Colima,
Mexico.
Acahuato (2); Apatzingan (142); Arteaga (4); 26 km. S of
Arteaga (4); Capirio (6); Chinapa; Chupio; 19 km. S of
Corralito (5); El Sabino (220); Jazmin (3); La Placita (8);
La Playa (14); La Salada (6); Lombardia (5); Nueva Italia
(14); Ojos de Agua de San Telmo (2); Oropeo (3); Ostula;
Punta de San Telmo (3); Rio Cancita, 14 km. E of Apatzingan
(13); Rio Marquez, 10 km. S of Lombardia (10); Rio Marquez,
13 km. SE of Nueva Italia (3); San Juan de Lima (2); Santa
Ana (2); Tafetan (2); Tepalcatepec (2); Tzitzio (6); Volcan
Jorullo (3).
This small species is extremely common in the Tepalcatepec Valley and
noticeably less so on the coast. It is usually found on the ground in
rocky areas, but males frequently have been seen on the trunks of low
trees in the scrub forest. Altitudinally, it ranges from sea level to
slightly more than 1000 meters. The sexes are readily distinguished in
the field (Oliver, 1937; Smith, 1939; Duellman, 1954b). In the dry
season only males were observed in the Tepalcatepec Valley, but in the
rainy season both sexes were found in approximately the same numbers.
~Sceloporus scalaris scalaris~ Wiegmann
_Sceloporus scalaris_ Wiegmann, Isis von Oken, 21:370,
1828.--Mexico. Type locality restricted to Mexico, Distrito
Federal, by Smith and Taylor (1950b:137).
_Sceloporus scalaris scalaris_, Smith, Occ. Pap. Mus. Zool.
Univ. Michigan, 361:2, December 15, 1937.
Carapan (2); Cheran; Ciudad Hidalgo; Huingo (3); Jacona (3);
Jiquilpan (2); Lago de Camecuaro (2); Lago de Chapala; Lago
de Cuitzeo (5); Morelia (4); Patzcuaro (4); Querendaro;
Quiroga; Tacicuaro (5); Tarecuaro; Zacapu (4); Zamora (4);
Zinapecuaro (11).
This small terrestrial species does not seem to be abundant anywhere in
the state. It sometimes is found in open pine, oak, or pine-oak forest,
but usually it is observed in areas supporting bunch grass. In such
places the lizards sun and forage on the open ground and quickly take
refuge in the large clumps of grass. Altitudinally, the species ranges
from 1550 to 2300 meters. Although _Sceloporus scalaris scalaris_ has
been found in association with _S. dugesi intermedius_, _S. spinosus_,
and _S. torquatus_, it does not seem to form any close ecological
association with any of these species. In the pine forests of the
Cordillera Volcanica _S. s. scalaris_ is replaced by _Sceloporus aeneus
aeneus_, another small terrestrial species that occurs in great
abundance throughout the coniferous forests of the Cordillera Volcanica.
~Sceloporus siniferus siniferus~ Cope
_Sceloporus siniferus_ Cope, Proc. Amer. Philos. Soc.,
11:159, 1869.--Pacific side of the Isthmus of Tehuantepec.
Type locality restricted to Tehuantepec, Oaxaca, Mexico, by
Smith and Taylor (1950b:134).
_Sceloporus siniferus siniferus_, Smith and Taylor, Bull. U.
S. Natl. Mus., 199:134, October 26, 1950.
Twenty-six km. S of Arteaga; Barranca de Bejuco (2);
Coahuayana; El Ticuiz (2); La Mira; La Orilla (2); La
Placita (9); Maruata; Ojos de Agua de San Telmo; Ostula (4);
Playa Azul (6); Pomaro (2); Puerto de las Higuerita; Santa
Ana (3).
This small terrestrial species inhabits the dense arid tropical scrub
forest on the coast and lower foothills of the Sierra de Coalcoman to
elevations of about 150 meters. It also occurs in the lower Balsas
Valley, but it has not been found in the scrub forest of the broad
Tepalcatepec Valley. Perhaps the large number of _Sceloporus siniferus_
on the coastal lowlands is responsible for the small number there of _S.
pyrocephalus_, another terrestrial species of about the same size. The
latter is abundant in the Tepalcatepec Valley, where _S. siniferus
siniferus_ has not been found. _Sceloporus siniferus siniferus_ is a
fast runner and difficult to collect; consequently, the small number of
specimens available is not indicative of its abundance.
~Sceloporus spinosus spinosus~ Wiegmann
_Sceloporus spinosus_ Wiegmann, Isis von Oken, 21:370,
1828.--Mexico. Type locality restricted to Puebla, Puebla,
Mexico, by Smith and Taylor (1950b:116).
_Sceloporus spinosus spinosus_, Martin del Campo, Anal.
Inst. Biol. Mexico, 8:262, 1937.
Cojumatlan (2); Huetamo Road; Lago de Cuitzeo (4); Maravatio
(8); Tupataro (2).
Although this species is widespread on the southern part of the Mexican
Plateau, it is uncommon in Michoacan. It has been collected only in
rather open situations in the mesquite-grassland on the plateau between
1500 and 2300 meters, where it has been found in association with
_Sceloporus dugesi intermedius_ and _S. scalaris scalaris_. Most
specimens of _Sceloporus spinosus spinosus_ have been observed on rock
fences. In this habitat the species is the larger member of a pair of
species, the smaller of which is _Sceloporus dugesi intermedius_.
~Sceloporus torquatus torquatus~ Wiegmann
_Sceloporus torquatus_ Wiegmann, Isis von Oken, 21:369,
1828.--Mexico. Type locality restricted to Mexico, Distrito
Federal, by Smith and Taylor (1950b:126).
_Sceloporus torquatus torquatus_, Cope, Proc. Amer. Philos.
Soc., 22:402, 1885.
Angahuan (31); Araparicuaro; Capacuaro (3); Carapan (11);
Cerro Tancitaro; Cheran; Ciudad Hidalgo; Cojumatlan;
Copandaro (2); Corupu (4); Cuseno Station (9); El Alamo;
Jacona (6); Jiquilpan (2); Jungapeo (3); Lago de Camecuaro;
Lago de Chapala; Lago de Cuitzeo (3); La Palma (2); Los
Conejos (3); Los Reyes (3); Maravatio (9); Morelia (17);
Paracho (3); Patzcuaro (27); Pino Gordo; Querendaro (2);
Quiroga; Sahuayo (3); San Jose de la Cumbre; San Juan de
Panangaricutiro; Tacicuaro (10); Tancitaro (200);
Tangamandapio; Tangancicuaro (3); Temazcal (2); Tupataro
(5); Uruapan (136); Zacapu; Zinapecuaro (10); Zirimicuaro
(12); Zitacuaro.
This large species inhabits the Mexican Plateau and the Cordillera
Volcanica, but not the Sierra de Coalcoman, where apparently it is
replaced by _Sceloporus bulleri_. _Sceloporus torquatus torquatus_
usually is found in pine or pine-fir forests at elevations between 1450
and 3000 meters. In many places it is almost entirely arboreal, but in
areas where there are many fallen trees or rock fences and rock piles,
many individuals have been found on the ground near the rocks or logs.
In the coniferous forests this species is associated with _S. grammicus
microtepidotus_ and _S. aeneus aeneus_.
The distinction made by Smith (1938:572) between the subspecies _S.
torquatus torquatus_ and _melanogaster_ is slight. Individuals with pale
bluish spots are found throughout the range of the species in Michoacan;
spotting is especially evident in the young. Individuals having an
incomplete nuchal collar have been found at Maravatio and at Zinapecuaro
in the northern part of the state; in this character these specimens
resemble _S. torquatus melanogaster_, which is found to the north from
Guanajuato to Zacatecas and San Luis Potosi.
~Sceloporus utiformis~ Cope
_Sceloporus utiformis_ Cope, Proc. Acad. Nat. Sci.
Philadelphia, 16:177, September 30, 1864.--Colima, Colima,
Mexico.
Nineteen km. S of Arteaga (2); Cascada Tzararacua (17);
Coahuayana (3); Coalcoman (6); El Sabino (2); El Ticuiz (2);
Ostula (3); Pomaro; Rio Cachan; San Juan de Lima; Uruapan
(26).
In Michoacan the range of this species is discontinuous. It has been
found between 1050 and 1550 meters on the slopes of the Cordillera
Volcanica, and on the coast and seaward slopes of the Sierra de
Coalcoman up to an elevation of 900 meters. It is absent from the
Tepalcatepec Valley. At Uruapan and at Cascada Tzararacua this lizard
was found on the ground in oak forest or in open pine-oak forest; on the
coast and foothills of the Sierra de Coalcoman it was found on the
ground in the gallery forests along streams, and not in the scrub
forest.
~Urosaurus bicarinatus tuberculatus~ (Schmidt)
_Uta tuberculata_ Schmidt, Amer. Mus. Novitates, 22:4,
December 1, 1921.--Colima, Colima, Mexico.
_Urosaurus bicarinatus tuberculatus_, Mittleman, Bull. Mus.
Comp. Zool., 91:169, September, 1942.
Twenty-six km. S of Arteaga; Cascada Tzararacua (2); Chupio;
Coahuayana; Coalcoman (8); El Sabino (2); Jungapeo; La
Orilla (2); La Placita (4); Playa Azul (4); Pomaro (2); San
Salvador (16);? Tupataro; Uruapan (12); Tzitzio; Zamora.
The known distribution and geographic variation of _Urosaurus
bicarinatus_ in southwestern Mexico presents a confused picture. In
general rugosity, specimens from the coastal region of Michoacan
(Coahuayana, La Orilla, La Placita, Playa Azul, and Pomaro) resemble _U.
bicarinatus tuberculatus_ to the north along the Pacific coast.
Furthermore, specimens from the coast have less stippling in the gular
region than do those from the Sierra de Coalcoman and the slopes of the
Cordillera Volcanica. Specimens from the mountains have greatly carinate
enlarged dorsals, large lateral tubercles, and heavily stippled throats;
in these characters they resemble specimens from Morelos, Guerrero, and
Oaxaca. As mentioned by Peters (1954:14), some specimens from La Orilla
and San Salvador are like _U. bicarinatus bicarinatus_ in certain
characters, and one specimen has the blue ventral patches restricted to
the sternal area, a characteristic of _U. bicarinatus anonymorphus_ of
Oaxaca and eastern Guerrero. Examination of all available specimens from
Michoacan indicates that the nature of the dorsal scales is of little
value in separating the subspecies. The specimens from Michoacan are
here provisionally referred to _U. bicarinatus tuberculatus_, because
cursory examination of specimens from several localities between Nayarit
and Oaxaca shows that there are only minor differences between the named
populations. Individuals from the northern part of the range are more
rugose and have larger blue ventral patches and less gular stippling
than those from the south.
In Michoacan _Urosaurus bicarinatus tuberculatus_ is found in wooded
areas, not in open scrub forest, in the coastal area to elevations of
about 900 meters, and along the slopes of the Cordillera Volcanica and
the southern edge of the Mexican Plateau at elevations from 1000 to 1700
meters. The record for Tupataro probably is erroneous, for no other
specimens of this species are known from the central plateau.
Essentially, the distribution of this species parallels that of
_Sceloporus utiformis_, a strictly terrestrial species. _Urosaurus
bicarinatus tuberculatus_ lives on tree trunks. Below 1000 meters in the
Tepalcatepec Valley _Urosaurus bicarinatus tuberculatus_ is replaced by
_Urosaurus gadowi_.
~Urosaurus gadowi~ (Schmidt)
_Uta gadovi_ Schmidt, Amer. Mus. Novitates, 22:3, December
1, 1921.--Cofradia, Jalisco, Mexico (in error) = Cofradia,
Michoacan, Mexico (Duellman, 1958b:49).
_Urosaurus gadowi_, Mittleman, Bull. Mus. Comp. Zool.,
91:154, September, 1942.
Acahuato (2); Apatzingan (56); 12-16 km. S of Apatzingan
(12); Buenavista (7); Capirio (23); Cofradia (21); El Sabino
(13); Guayabo; Jazmin; La Playa; La Salada (3); Nueva Italia
(7); Rancho Nuevo; Rio Cancita, 14 km. E of Apatzingan (5);
Rio Marquez, 10 km. S of Lombardia (2); Rio Marquez, 13 km.
SE of Nueva Italia (3); San Salvador (2); Santa Ana;
Tepalcatepec; Volcan Jorullo (3); Zicuiran (2);
Ziracuaretiro.
Although individuals of this species have been collected at elevations
slightly exceeding 1200 meters on Volcan Jorullo and at 1100 meters at
Ziracuaretiro on the southern slopes of the Cordillera Volcanica, for
the most part these lizards are found at elevations of less than 800
meters, where they inhabit the open arid scrub forest of the
Tepalcatepec Valley, a region to which this species is endemic
(Duellman, 1958b:49). These small lizards usually are found on the
trunks and main branches of the small trees in the scrub forest; in this
habitat they are associated with _Sceloporus horridus oligoporus_, a
much larger species.
Males have a pale orange spot on the throat and a pale blue belly;
females have immaculate venters.
A specimen from Guayabo on the northern slopes of the Sierra de
Coalcoman was referred to _Urosaurus irregularis_ (Fischer) by Peters
(1954:15). I have studied this specimen (BMNH 1914.1.28.110), a female
having a snout-vent length of 46 mm., and agree with Peters that it
closely resembles Fischer's description and figure (1882: pl. 17, fig.
1). This specimen and those seen of _Urosaurus gadowi_ all have
pavementlike enlarged dorsal scales that are complete across the
vertical line. In _U. gadowi_ the enlarged dorsals usually are in four
to six irregular rows; in the specimen from Guayabo the dorsals are in
two rows. Although none of the other specimens of _U. gadowi_ examined
has only two rows of enlarged dorsals, I prefer to consider the specimen
from Guayabo as an aberrant individual of that species, rather than _U.
irregularis_. Guayabo is in the known range of _U. gadowi_. _Urosaurus
irregularis_ is known only from the type specimen in the Bremen Museum;
the type locality, according to Fischer (1882:232), is "Aus dem
Hochlande von Mexico." If an examination of the type specimen of _U.
irregularis_ shows it to be identical with _U. gadowi_, then _U.
irregularis_ would be the name for the lizards here referred to _U.
gadowi_.
~Mabuya brachypoda~ Taylor
_Mabuya brachypoda_ Taylor, Univ. Kansas Sci. Bull., 38
(1):308, December 20, 1956.--Four kilometers east-southeast
of Los Angeles de Tilaran, Guanacaste, Costa Rica.
El Sabino (42); La Placita; Playa Azul; Tzitzio (3).
Previously this species has been reported from La Placita as _Mabuya
mabouya alliacea_ by Peters (1954:15). Webb (1958:1311) provided
evidence that Mexican specimens were conspecific with _Mabuya
brachypoda_, as described from Costa Rica by Taylor (1956:308). The
large series in the Taylor collection studied by Webb and listed by him
as being from Uruapan actually is part of a series collected by Hobart
M. Smith at El Sabino at an elevation of 1050 meters, 30 kilometers
south of Uruapan.
This species probably ranges throughout the coastal region of the state;
individuals from La Placita and Playa Azul were taken in dense scrub
forest near sea level.
~Scincella assata taylori~ (Oliver)
_Leiolopisma assatum taylori_ Oliver, Occ. Pap. Mus. Zool.
Univ. Michigan, 360:12, November 20, 1937.--Santiago,
Colima, Mexico.
_Scincella assata taylori_, Mittleman, Herpetologica, 6:20,
June 5, 1950.
Twenty-one km. S of Arteaga; Ostula.
The specimen from Ostula was obtained in semi-deciduous broad-leaf
forest at an elevation of 120 meters; that from 21 kilometers south of
Arteaga was taken in oak forest at an elevation of 830 meters. Both
localities are on the coastal slopes of the Sierra de Coalcoman.
Probably the species inhabits the heavy forests on the lower slopes of
these mountains. The specimen from south of Arteaga (UMMZ 119117) in
life had a tan dorsum and a bright orange-pink tail.
~Eumeces altamirani~ Duges
_Eumeces altamirani_ Duges, La Naturaleza, ser. 2, 1:485,
1891.--Apatzingan, Michoacan, Mexico.
Twelve km. E of Apatzingan; El Sabino (4).
One specimen of this rare species was found beneath a rock in the open
scrub forest 12 kilometers east of Apatzingan on July 3, 1955. Another
skink, presumably of this species, was seen at Capirio. The specimen
from east of Apatzingan is a male having a snout-vent length of 97 mm.
and an incomplete tail. In most respects it compares favorably with
accounts of the species given by Taylor (1936b:55 and 1936c:102). The
frontal is divided by a transverse suture; the enlarged dorsal scales
are arranged in 11 pairs anteriorly, followed by 48 unpaired enlarged
scales. The head and middorsal area are brown; there is a pale tan
stripe on the edges of the vertebral and paravertebral rows, bordered by
a dark brown stripe on the paravertebral row, which, in turn, is
bordered by a pale tan stripe on the lateral edge of the paravertebral
scale row and the median edge of the adjacent scale row. The stripes
extend from the neck to the base of the tail. The flanks are mottled
with brown and cream-color; the labials are cream-color barred by brown;
the venter is a pale cream-color.
Duges (1891:485) described _Eumeces altamirani_ from "las regiones
calidas del Estado de Michoacan" and subsequently (1896:480) gave
Apatzingan as a locality for the species. Presumably he had only one
specimen. In 1935 Hobart M. Smith collected the species at El Sabino on
the lower slopes of the Cordillera Volcanica bordering the Tepalcatepec
Valley. All of the known specimens are from this valley and the adjacent
slopes, an area to which the species apparently is endemic.
~Eumeces colimensis~ Taylor
_Eumeces colimensis_ Taylor, Publ. Field Mus. Nat. Hist.,
zool. ser., 20:77, May 15, 1935.--Colima, Colima, Mexico.
Coalcoman; Salitre de Estopila.
The species was reported by Peters (1954:16); no additional material has
been discovered. The species is known only from foothills and low
mountains at elevations between 130 and 950 meters in Michoacan and
Colima.
~Eumeces copei~ Taylor
_Eumeces copei_ Taylor, Proc. Biol. Soc. Washington, 46:133,
June 5, 1933.--10 miles southeast of Asuncion, Mexico,
Mexico. Cerro Tancitaro (3); Zacapu.
This member of the _Eumeces brevirostris_-group has been found only in
pine or pine-fir forests at elevations from 1800 to 2700 meters. It
probably ranges throughout the high mountains of the state north of the
Tepalcatepec Valley; its apparent absence in other parts of the
Cordillera Volcanica, other than on Cerro Tancitaro, is surprising. The
species has been taken near Asuncion in the state of Mexico and at
Lagunas de Zempoala in Morelos.
In this species the lateral pale yellow stripe, which is bordered below
by dark brown, extends to the groin and onto the base of the tail. The
dorsolateral stripe is separated from the copper-colored middorsum by a
narrow brown stripe.
~Eumeces dugesi~ Thominot
_Eumeces Dugesii_ Thominot, Bull. Soc. Philom. Paris, ser.
7, 7:138, 1883.--Guanajuato. Type locality restricted to
Guanajuato, Guanajuato, Mexico, by Smith and Taylor
(1950b:169).
Carapan (6); Cheran (5); Opopeo (2); 17 km. S of Patzcuaro
(3); San Jose de la Cumbre (2); Tancitaro (2);
Tangancicuaro; Uruapan; Zacapu.
Individuals of this species frequently have been found beneath rocks and
logs in pine-oak, pine, or fir forests from elevations of 1550 to 1850
meters. To judge from specimens available, _E. dugesi_ probably is the
most abundant and widespread species of skink in the state.
In this species the lateral yellow stripe is indistinct and is
persistent only in the axilla; the dorsolateral stripes terminate
anterior to the hind limbs and are not separated from the tan dorsum.
~Eumeces indubitus~ Taylor
_Eumeces indubitus_ Taylor, Univ. Kansas Sci. Bull., 21:257,
November 27, 1934.--Near Cuernavaca, Morelos, Mexico.
Puerto Hondo.
The one specimen of this species from Michoacan was collected by Edward
H. Taylor in pine forest at Puerto Hondo, near Zitacuaro, at an
elevation of about 2750 meters (Taylor, 1935:466). The species is known
from the high mountains of eastern Michoacan, western Mexico, and
northern Morelos.
~Eumeces parvulus~ Taylor
_Eumeces parvulus_ Taylor, Proc. Biol. Soc. Washington,
46:175, October 26, 1933.--Tepic, Nayarit, Mexico.
El Ticuiz; La Placita; Pomaro (2); San Pedro Naranjestila
(3).
Aside from the specimens reported by Peters (1954:17), one other
specimen was obtained at El Ticuiz. It has 22 scale rows, 3 supraoculars
in contact with the frontal, 2 postlabials, and a unicolored olive-tan
dorsum. In life the anterior dorsolateral stripes were pale pinkish tan,
the labials cream color, the throat white, and the tail pale blue. All
specimens were found in semi-deciduous broad-leaf forest at elevations
of less than 500 meters on the seaward slopes of the Sierra de
Coalcoman.
~Ameiva undulata sinistra~ Smith and Laufe
_Ameiva undulata sinistra_ Smith and Laufe, Univ. Kansas
Sci. Bull., 31 (1):59, May 1, 1946.--Manzanillo, Colima,
Mexico.
Apatzingan (9); 19 km. S of Arteaga (3); Barranca de Bejuco
(2); Coahuayana (6); Coalcoman (3); El Ticuiz (10); La
Placita (2); Limoncito (3); Ostula (2); Playa Azul; Salitre
de Estopila; San Juan de Lima (2); San Pedro Naranjestila
(4).
Six males and six females from the Tepalcatepec Valley have more femoral
pores than do 16 males and nine females from the coastal lowlands; the
ranges and average number of femoral pores in the former are 40-50
(44.8) for males and 38-40 (38.6) for females; males from the coast have
34-44 (39.2), and females have 32-40 (36.2) femoral pores. In all
specimens the number of lamellae beneath the fourth toe varies from 26
to 33 (29.7). In life juveniles have a pale olive-tan dorsum and a
dorsolateral dark band, superimposed on which is a row of darker brown
spots. The dorsolateral band is bordered below by a narrow cream-colored
stripe. The tail is tan above and grayish white below; the belly is pale
bluish white. Adult males are brilliantly colored in life. A male having
a snout-vent length of 108 mm. had a rusty brown dorsum and bright blue
bars on the flanks separated by dark brown interspaces. The side of the
head was pale green, and the chin and throat were golden yellow. In some
specimens the throat is orange. Juveniles and subadults have dark flecks
on the brown or tan middorsal area, but these are absent in the largest
males.
This species inhabits the heavily wooded areas in the lowlands to
elevations of about 950 meters. In the Tepalcatepec Valley it has been
found only in gallery forests along streams. In both the Tepalcatepec
Valley and the coastal lowlands there is a noticeable absence of large
adults in the dry season.
~Cnemidophorus calidipes~ Duellman
_Cnemidophorus calidipes_ Duellman, Occ. Pap. Mus. Zool.
Univ. Michigan, 574:1, December 23, 1955.--Capirio,
Michoacan, Mexico.
Apatzingan (56); 12-20 km. S of Apatzingan (5); 19 km. E of
Apatzingan (5); 25 km. S of Arteaga; Capirio (57); El
Espinal (13); Jazmin (9); 11 km. S of Lombardia; Nueva
Italia.
This small, distinctive species of the _sexlineatus_-group of
_Cnemidophorus_ was discovered in the Tepalcatepec Valley in 1955
(Duellman, 1955); subsequent field studies showed it to be widespread in
the valley (Duellman, 1960c). One specimen (KU 29747) is from the
relatively arid, low Pacific slope of the Sierra de Coalcoman, 25
kilometers south of Arteaga. All other specimens have been taken at
elevations of 200 to 650 meters in the Tepalcatepec Valley, where the
species characteristically inhabits the open scrub forests of the valley
floor, especially the _Cercidium-Prosopis-Apoplanesia_ associations,
where there is a sparse growth of grasses. In this habitat it is most
frequently seen in association with _Cnemidophorus costatus zweifeli_
and _C. deppei infernalis_.
Aside from the characters given in Table 5, _Cnemidophorus calidipes_
differs from other species of _Cnemidophorus_ in Michoacan by possessing
a complete (or nearly so) supraorbital semicircle-series of granules; in
other species the granules seldom extend anteriorly beyond the posterior
border of the frontal.
~Cnemidophorus communis communis~ Cope
_Cnemidophorus communis_ Cope, Proc. Amer. Philos. Soc.,
17:95, 1877.--No type locality given; type locality
restricted to Colima, Colima, Mexico, by Zweifel (1959a:74).
_Cnemidophorus communis communis_, Zweifel, Bull. Amer. Mus.
Nat. Hist., 117:74, April 27, 1959.
Aguililla (2); Apatzingan (6); 13 km. S of Arteaga (2); 19
km. S of Arteaga (3); Capirio (3); Coahuayana (3); Coalcoman
(44); El Ticuiz; between El Ticuiz and Ojos de Agua de San
Telmo; La Placita (6); Pomaro (2); Rio Cachan; Salitre de
Estopila; San Juan de Lima.
The specimens from Coalcoman and the coastal localities were referred to
_Cnemidophorus sacki copei_ by Peters (1954:18) and Duellman (1954b:12).
Zweifel (1959a) referred these specimens to _Cnemidophorus communis
communis_ and pointed out the probable sympatry of _C. communis_ and _C.
costatus_ (= _sacki_ of Zweifel) in the Tepalcatepec Valley.
There is considerable geographic variation in the number of dorsal
granules around the midbody. Sixteen specimens from the coastal regions
of Michoacan have 129-146 (136.3) granules; nine from the Tepalcatepec
Valley have 124-137 (128.3), and 44 from Coalcoman at an elevation of
950 meters in the Sierra de Coalcoman, intermediate geographically
between the coast and the Tepalcatepec Valley, have 105-144 (119.7). The
number of granules in specimens from the coast of Michoacan compares
favorably with the range of 118-154 (137.8) for 34 specimens from
Colima, Colima (Zweifel, 1959a:107). Aside from the characters given in
Table 5, _C. communis communis_ can be distinguished from other members
of the _Cnemidophorus sexlineatus_-group (_calidipes_, _costatus_, and
_scarlaris_) by its relatively small post-antebrachial scales.
TABLE 5.--COMPARISON OF THE TEN SPECIES AND SUBSPECIES OF CNEMIDOPHORUS
IN MICHOACAN (SCALE COUNTS ARE FOR SPECIMENS FROM MICHOACAN ONLY)
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