[205]

Volume 15, No. 5, pp. 205-249, pls. 7-10, 6 figs.

Amphibians and Reptiles of the Rainforests
of Southern El Petén, Guatemala

BY

WILLIAM E. DUELLMAN

University of Kansas

Lawrence

1963

[206]

University of Kansas Publications, Museum of Natural History



Editors: E. Raymond Hall, Chairman, Henry S. Fitch,

Theodore H. Eaton, Jr.

Vol. 15, No. 5, pp. 205-249, pls. 7-10, 6 figs.

Published October 4, 1963

University of Kansas

Lawrence, Kansas

PRINTED BY

JEAN M. NEIBARGER, STATE PRINTER

TOPEKA, KANSAS

1963





29-5935

[207]

Amphibians and Reptiles of the Rainforests
of Southern El Petén, Guatemala

BY

WILLIAM E. DUELLMAN

CONTENTS

INTRODUCTION

Early in 1960 an unusual opportunity arose to carry on biological
field work in the midst of virgin rainforest in southern El Petén,
Guatemala. At that time the Ohio Oil Company of Guatemala had
an air strip and camp at Chinajá, from which place the company
was constructing a road northward through the forest. In mid-February,
1960, J. Knox Jones, Jr. and I flew into El Petén to
collect and study mammals, reptiles, and amphibians. While enjoying
the comforts of the fine field camp at Chinajá, we worked
in the surrounding forest and availed ourselves of the opportunity
to be on hand when the road crews were cutting the tall trees in
the forest, thereby bringing to the ground many interesting specimens
of the arboreal fauna. We stayed at Chinajá until late March,
with the exception of a week spent at Toocog, another camp of the
Ohio Oil Company located 15 kilometers southeast of La Libertad
and on the edge of the savanna. Thus, at Toocog we were able[208] to work both in the forest and on the savanna. In the summer of
1960, John Wellman accompanied me to El Petén for two weeks
in June and July. Most of our time was spent at Chinajá, but a
few days were spent at Toocog and other localities in south-central
El Petén.

Many areas in Guatemala have been studied intensively by
L. C. Stuart, who has published on the herpetofauna of the forested
area of northeastern El Petén (1958), the savannas of central
El Petén (1935), and the humid mountainous region to the south
of El Petén in Alta Verapaz (1948 and 1950). The area studied
by me and my companions is covered with rainforest and lies to
the north of the highlands of Alta Verapaz and to the south of the
savannas of central El Petén. A few specimens of amphibians
and reptiles were obtained in this area in 1935 by C. L. Hubbs
and Henry van der Schalie; this collection, reported on by Stuart
(1937), contained only one species, Cochranella fleischmanni, not
present in our collection of 77 species and 617 specimens.

Acknowledgments

I am grateful to L. C. Stuart of the University of Michigan, who made the
initial arrangements for our work in El Petén, aided me in the identification
of certain specimens, and helped in the preparation of this report. J. Knox
Jones, Jr. and John Wellman were able field companions, who added greatly
to the number of specimens in the collection. In Guatemala, Clark M.
Shimeall and Harold Hoopman of the Ohio Oil Company of Guatemala made
available to us the facilities of the company's camps at Chinajá and Toocog.
Alberto Alcain and Luis Escaler welcomed us at Chinajá and gave us every
possible assistance. Juan Monteras and Antonio Aldaña made our stay at
Toocog enjoyable and profitable. During our visits to southern El Petén, Julio
Bolón C. worked for us as a collector, and between March and June he collected
and saved many valuable specimens; his knowledge of the forest and
its inhabitants was a great asset to our work. Jorge A. Ibarra, Director of
the Museo Nacional de Historia Natural in Guatemala assisted us in obtaining
necessary permits and extended other kindnesses. To all of these people I
am indebted for the essential parts that they played in the completion of this
study.

Field work in the winter of 1960 was made possible by funds from the
American Heart Association for the purposes of collecting mammalian hearts.
My field work in the summer of 1960 was supported by a grant from the
Graduate Research Fund of the University of Kansas.

DESCRIPTION OF THE AREA

A vast lowland region stretches northward for approximately
700 kilometers from the highlands of Guatemala to the Gulf of
Mexico. The northern two-thirds of this low plain is bordered on
three sides by seas and forms the Yucatán Peninsula. The lowlands[209] at the base of the Yucatán Peninsula make up the Departamento
El Petén of Guatemala. The area with which this report is concerned
consists of the south-central part of El Petén.

Physiography

Immediately south of Chinajá is a range of hills, the Serrania de Chinajá,
having an almost due east-west axis and a crest of about 600 meters above
sea level. South of the Serrania de Chinajá are succeedingly higher ridges
building up to the Meseta de Cobán and Sierra de Pocolha and eventually
to the main Guatemalan highlands. The northern face of the Serrania de
Chinajá is a fault scarp dropping abruptly from about 650 meters at the crest
to about 140 meters at the base. From the base of the Serrania de Chinajá
northward to the Río de la Pasión at Sayaxché the terrain is gently rolling
and has a total relief of about 50 meters. North of the Río de la Pasión is
a low dome reaching an elevation of 170 meters at La Libertad; see Stuart
(1935:12) for further discussion of the physiography of central El Petén.
The rocks in southern El Petén are predominately Miocene marine limestones;
there are occasional pockets of Pliocene deposits. There is little evidence
of subterranean solution at Chinajá, but northward in central El Petén karsting
is common. The upper few inches of soil is humus rich in organic matter;
below this is clay.

Climate

The climate of El Petén is tropical with equable temperatures throughout
the year. Temperatures at Chinajá varied between a night-time low of 65° F.
and a daytime high of 91° F. during the time of our visits. In the Köppen
system of classification the climate at Chinajá and Toocog is Af. Rain falls
throughout the year, but there is a noticeable dry season. To anyone who
has traveled from south to north in El Petén and the Yucatán Peninsula, it is
obvious from the changes in vegetation that there is a decrease in rainfall
from south to north. There is a noticeable difference between Chinajá and
Toocog. Although rainfall data are not available for Chinajá and Toocog,
there are records for nearby stations (Sapper, 1932). At Paso Caballos on
the Río San Pedro about 40 kilometers northwest of Toocog the average
annual rainfall amounts to 1620 mm.; the driest month is March (21 mm.),
and the wettest months are June (269 mm.) and September (265 mm.). At
Cubilquitz, Alta Verapaz, about 35 kilometers south-southwest of Chinajá
and at an elevation of 300 meters, the average annual rainfall is 4006 mm.;
the driest month is March (128 mm.), and the wettest months are July (488
mm.) and October (634 mm.).

During the 18 days in February and March, 1960, that we kept records on
the weather at Chinajá moderate to heavy showers occurred on seven days.
During our stay there in June and July rain fell every day, as it did in Toocog.
However, during the week spent at Toocog in March no rain fell.

Vegetation

The vegetation of northern and central El Petén has been studied by
Lundell (1937), who made only passing remarks concerning the plants of the
southern part of El Petén. No floristic studies have been made there. The[210] following remarks are necessarily brief and are intended only to give the
reader a general picture of the forest. I have included names of a few of
the commoner trees that I recognized.

Chinajá is located in a vast expanse of unbroken rainforest. In this forest
there is a noticeable stratification of the vegetation. Three strata are apparent;
in the uppermost layer the tops of the trees are from 40 to 50 meters above
the ground. The spreading crowns of the trees and the interlacing vines form
a nearly continuous canopy over the lower layers. Among the common trees
in the upper stratum are Calophyllum brasiliense, Castilla elastica, Cedrela
mexicana, Ceiba pentandra, Didalium guianense, Ficus sp., Sideroxylon lundelli, Swietenia macrophylla, and Vitex sp. (Pl. 1, fig. 1). The middle layer of
trees have crowns about 25 meters above the ground; these trees in some
places where the upper canopy is missing form the tallest trees in the forest. This
is especially true on steep hillsides. Common trees in the middle layer include Achras zapote, Bombax ellipticum, Cecropia mexicana, Orbignya cohune, and Sabal sp. The lowermost layer reaches a height of about 10 meters; in many
places in the forest this layer is absent. Common trees in the lower stratum
include Crysophila argentea, Cymbopetalum penduliflorum, Casearia sp., and Hasseltia dioica.

The ground cover is sparce; apparently only a few small herbs and ferns
live on the heavily shaded forest floor. Important herpetological habitats
include the leaf litter, rotting stumps, and rotting tree trunks on the forest
floor and the buttresses of many of the gigantic trees, especially Ceiba pentandra (Pl. 2). Epiphytes, especially various kinds of bromeliads, are common.
Most frequently these are in the trees in the upper and middle strata.

At Toocog there is sharp break between savanna and forest (Pl. 7, fig. 2).
The forest is noticeably drier and more open than at Chinajá (Pl. 9). The
crowns of the trees are lower, and there is no nearly continuous canopy between
40 and 50 meters above the ground. Although Swietenia macrophylla and
other large trees occur, they are less common than at Chinajá. Especially
common at Toocog are Achras zapote, Brosimum alicastrum, and various species
of Ficus.

GAZETTEER

The localities from which specimens were obtained are cited below and
shown on the accompanying map (Fig. 1).

Fig. 1. Map of El Petén, Guatemala, showing localities mentioned in text.

THE HERPETOFAUNA OF THE RAINFOREST

In presenting an account of the herpetofauna of southern El Petén three
items need to be considered: (1) The composition of the fauna; (2) the
ecology of the fauna; (3) the relationships of the fauna. Each of these
topics is discussed briefly below. Logically a discussion of the origin of the
fauna should follow, but this is being withheld for inclusion in a report
on the herpetofauna of the entire El Petén by L. C. Stuart and the author;
at that time the above topics will be expanded to cover the herpetofauna of
the whole region.

[212]

Composition of the Fauna

Table 1.—Composition of the Herpetofauna in Southern
El Petén, Guatemala.

A total of 78 species of amphibians and reptiles has been found in the
rainforests in southern El Petén; a break down into families and genera is
given in table 1. Another 13 species probably occur in southern El Petén (see
Hypothetical List of Species). The fauna primarily is composed of typical
humid lowland forest inhabitants, such as:

Nevertheless, the region also provides at least a limited amount of habitat
suitable for some species that are more frequently found in open forest of
a drier nature; such species include:

Because of the absence of sufficiently open habitat or owing to the
presence of competitors, some conspicuous members of sub-humid forests are
not present in southern El Petén. Conspicuous absentees are the following:

PLATE 7

Fig. 1. Edge of rainforest along airstrip at Chinajá, El Petén, Guatemala.

Fig. 2. Rainforest at edge of savanna at Toocog, El Petén, Guatemala.

PLATE 8

Interior of rainforest at Chinajá. Notice size of buttresses on large tree (Ceiba
pentandra).

PLATE 9

Interior of rainforest at Toocog. Notice less dense vegetation as compared
with Pl. 8.

PLATE 10

Fig. 1. Rainforest along Río San Román, 16 kilometers north-northwest of
Chinajá.

Fig. 2. Rain pond in forest at Toocog. This was a breeding site for six species
of frogs.

Ecology of the Herpetofauna

Our two visits to Chinajá and Toocog afforded the opportunity
to gather data on the ecology of the rainforests of southern El Petén
and to study the relationships between the environment and members
of the herpetofauna. Tropical rainforests present the optimum[213] conditions for life, and it is in this environment that life
reaches its greatest diversity. Here, too, biological inter-relationships
are most complex. This complexity is illustrated by the
presence of many species of some genera, all of which are found
together in the same geographic region. In the rainforests of
southern El Petén there are six species of Anolis, five of Hyla,
four of Bothrops, and three of Coniophanes. Obviously, the diversity
of ecological niches in the rainforest is sufficient to support
a variety of related species. Of the examples mentioned above,
fairly adequate ecological data were obtained for most of the
species of Anolis, which will be used to show the ecological diversity
and vertical stratification of sympatric species in the rainforests.

Of the six species of Anolis, all except A. sericeus are typically
found in humid forests. Anolis sericeus sericeus is poorly represented
in the collections from southern El Petén, where it may be
in competition with Anolis limifrons rodriguezi that resembles Anolis
s. sericeus in size, coloration, and habits. Therefore, Anolis sericeus
sericeus is excluded from the following discussion. The common
terrestrial species is Anolis humilis uniformis; sometimes
this small species perches or suns on the bases of small trees or
buttresses of some large trees. When disturbed it takes to the
ground and seeks cover in the leaf litter or beneath logs or palm
fronds. Anolis lemurinus bourgeaei is about twice the size of Anolis humilis uniformis and is usually observed on buttresses of
large trees or on the lower two meters of tree trunks. Individuals
were seen foraging on the ground along with Anolis humilis
uniformis. At no time were Anolis lemurinus bourgeaei observed
to ascend the trunks of large trees; they always took refuge near
the bases of trees. Anolis limifrons rodriguezi is found on the
stems and branches of bushes. It is a small species that sometimes
is observed on the ground but was never seen ascending large
trees. Anolis capito is about the same size as Anolis lemurinus
bourgeaei and lives on the trunks of large trees. In the tops of
the trees lives a large green species, Anolis biporcatus.

Similar segregation habitatwise can be demonstrated for other
members of the herpetofauna. The avoidance of interspecific
competition in feeding is well illustrated by three species of snakes
that probably are the primary ophidian predators on frogs. Drymobius margaritiferus margaritiferus is diurnal and terrestrial;
it feeds on frogs at the edges of breeding ponds by day. Also
during the day Leptophis mexicanus mexicanus feeds on frogs in[214] bushes and trees. At night the activities of both of these species
is replaced by those of Leptodeira septentrionalis polysticta, which
not only feeds on the frogs in the trees and bushes, but descends
to the ground and even enters the water in search of food.

From the examples discussed above, the importance of the
three dimensional aspect of the rainforest is apparent. The
presence of a large and diverse habitat above the ground is of
great significance in the rainforest, for of the non-aquatic components
of the herpetofauna in the rainforests of southern El Petén,
42 per cent of the species spend at least part of their lives in the
bushes and trees. Another important part of the forest is the
subterranean level—the rich mulch, underground tunnels, and
rotting subterranean vegetation. Of the 78 species of amphibians
and reptiles in southern El Petén, seven are primarily fossorial, and
half-a-dozen others are secondarily fossorial. Probably the fossorial
members of the fauna are the least well represented in the
collection, for such widespread species as Dermophis mexicanus
mexicanus, Rhadinaea decorata decorata and Tantilla schistosa
schistosa were expected, but not found.

In the following discussion of the ecological distribution of
amphibians and reptiles in the rainforest I have depended chiefly
on my observations made in southern El Petén, but have taken
into consideration observations made on the same species in other
regions, together with reports from other workers. The reader
should keep in mind that the evidence varies from species to species.
Of some species I have observed only one animal in the
field; of others, I have seen scores and sometimes hundreds of
individuals. For species on which I have few observations or
rather inconclusive evidence, the circumstance of inadequate
data is mentioned.

In analyzing the ecological distribution within the forest, it is
convenient to recognize five subdivisions (habitats); each is
treated below as a unit.

1. Aquatic.—This habitat includes permanent streams and rivers
(Pl. 10, fig. 1), some of which are clear and others muddy. In
the rainy season temporary ponds form in depressions on the forest
floor (Pl. 10, fig. 2); these are important as breeding sites for
many species of amphibians. Aquatic members of the herpetofauna
are here considered to be those species that either spend
the greatest part of their lives in the water or usually retreat to
water for shelter. Seven species of turtles and one crocodilian are[215] aquatic. Of these, Dermatemys mawi, Staurotypus triporcatus,
and Pseudemys scripta ornata inhabit clear water, whereas Chelydra
rossignoni, Claudius angustatus, Kinosternon acutum, and K. leucostomum inhabit muddy water. Crocodylus moreleti apparently
inhabits both clear and muddy water, for in the dry season it
lives along the clear rivers, but in the rainy season inhabits flooded
areas in the forest as well.

2. Aquatic Margin.—Extensive marshes were lacking in the
part of southern El Petén that I visited; consequently, the aquatic
margin habitat is there limited to the edges of rivers and borders
of temporary ponds. Bufo marinus, Rana palmipes, and Rana
pipiens are characteristic inhabitants of the aquatic margin,
although in the rainy reason Bufo marinus often is found away
from water. Observations indicate that Tretanorhinus nigroluteus
lateralis inhabits the margins of ponds and streams and actually
spends considerable time in the water. Although Iguana iguana
rhinolopha is arboreal, it lives in trees along rivers, into which it
plunges upon being disturbed. Species included in this category
are those that customarily spend most of their lives at the edge
of permanent water. Frogs and toads that migrate to the water
for breeding and the snakes that prey on the frogs at that time
are not assigned to the aquatic-margin habitat.

3. Fossorial.—Characteristic inhabitants of the mulch on the
forest floor are Bolitoglossa moreleti mulleri, Lepidophyma flavimaculatum
flavimaculatum, Scincella cherriei cherriei, Ninia sebae
sebae, Pliocercus euryzonus aequalis, and Micrurus affinis apiatus.
Other species of snakes that spend most of their lives above ground
often forage in the mulch layer; among these are Coniophanes
bipunctatus biserialis, Coniophanes fissidens fissidens, Coniophanes
imperialis clavatus, Lampropeltis doliata polyzona, and Stenorrhina
degenhardti. Among the amphibians, at least Hypopachus cuneus
nigroreticulatus, Eleutherodactylus rostralis, and Syrrhophus leprus are known to seek shelter in the mulch.

4. Terrestrial.—One turtle, Geoemyda areolata, is primarily
terrestrial. Among the lizards, conspicuous terrestrial species are Anolis humilis uniformis and Ameiva festiva edwardsi; Anolis
lemurinus bourgeaei and Basiliscus vittatus spend part of their
lives on the ground, but also live on trees and in bushes. Eumeces
schwartzei and E. sumichrasti apparently are terrestrial. The only
terrestrial lizard that is nocturnal is Coleonyx elegans elegans, which[216] by day hides in the leaf litter or below ground. Nocturnal amphibians
that are terrestrial include Bufo marinus, Bufo valliceps
valliceps, Eleutherodactylus rugulosus rugulosus, Syrrhophus leprus,
and Hypopachus cuneus nigroreticulatus. A large number of active
diurnal snakes are terrestrial; these include Boa constrictor imperator, Clelia clelia clelia, Dryadophis melanolomus laevis, Drymarchon
corais melanurus, Drymobius margaritiferus margaritiferus, Pseustes poecilonotus poecilonotus, and Spilotes pullatus mexicanus.
Nocturnal terrestrial snakes include three kinds of Bothrops (B.
atrox asper, B. nasutus, and B. nummifer nummifer), all of which
seem to be equally active by day.

5. Arboreal.—In this habitat the third dimension (height) of
the rainforest probably is the most complex insofar as the inter-relationships
of species and ecological niches are concerned. I
have attempted to categorize species as to microhabitats within
the arboreal habitat; in so doing, I recognize four subdivisions—bushes,
tree trunks, tree tops, and epiphytes.

Bush inhabitants include several species of lizards and snakes,
all of which have rather elongate, slender bodies, and long tails.
Common bush-inhabitants in southern El Petén are Anolis limifrons
rodriguezi, Basiliscus vittatus, Laemanctus deborrei, Leptophis
mexicanus mexicanus, and Oxybelis aeneus aeneus. All of these
are diurnal, and all but Laemanctus have been observed sleeping
on bushes at night.

Tree-trunk inhabitants include five species of lizards. Thecadactylus
rapicaudus lives on the trunks of large trees; Sphaerodactylus
lineolatus lives beneath the bark on dead trees and on
corozo palms. Anolis lemurinus bourgeaei lives on the bases and
buttresses of large trees, from which it often descends to the ground. Corythophanes cristatus and Anolis capito were found only on tree
trunks and large vines.

The least information is available for the species living in the
tree tops. The following species were obtained from tops of trees
when they were felled, or have been observed living in the tree
tops: Anolis biporcatus, Iguana iguana rhinolopha, Celestus rozellae, Leptodeira septentrionalis polysticta, Leptophis ahaetulla
praestans, Sibon dimidiata dimidiata, and Sibon nebulata nebulata.

Epiphytes, especially the bromeliads, provide refuge for a variety
of tree frogs and small snakes. Of the tree frogs, Hyla picta, Hyla
staufferi, Phyllomedusa callidryas taylori, Similisca baudini, and Similisca phaeota cyanosticta have been found in bromeliads; other[217] species probably occur there. Among the snakes, Imantodes
cenchoa leucomelas, Leptodeira frenata malleisi, Leptodeira
septentrionalis polysticta, Sibon dimidiata dimidiata, and Sibon
nebulata nebulata are frequent inhabitants of bromeliads; all of
these snakes are nocturnal.

Relationships of the Fauna

Most of the 78 species of amphibians and reptiles definitely
known from the rainforest in southern El Petén have extensive
ranges in the Atlantic lowlands of southern México and Central
America; many extend into South America. Sixty-two (80%) of the
species belong to this group having extensive ranges in Middle
America. Three species (Syrrhophus leprus, Leptodeira frenata,
and Kinosternon acutum) are at the southern limits of their distributions
in southern El Petén and northern Alta Verapaz, whereas Eleutherodactylus rostralis and Thecadactylus rapicaudus are at
the northern and western limits of their distributions in El Petén.
Nine (11%) species have the center of their distributions in El
Petén and the Yucatán Peninsula; representatives of this group
include Claudius angustatus, Dermatemys mawi, Laemanctus
deborrei, and Eumeces schwartzei.

In determining a measure of faunal resemblance, I have departed
from the formulae discussed by Simpson (1960) and have analyzed
the degree of resemblance by the following formula used to calculate
an index of faunal relationships:

The herpetofauna of southern El Petén has been compared with
that in the Tikal-Uaxactún area (Stuart, 1958), that in the humid
lowlands of Alta Verapaz (Stuart, 1950, plus additional data), and
that in the Mexican state of Yucatán (Smith and Taylor, 1945,
1948, and 1950). The herpetofaunas of lowland Alta Verapaz and
Yucatán are the largest, having respectively 94 and 91 species,
where as there are 78 species known from southern El Petén and
64 from the Tikal-Uaxactún area. An analysis of faunal relationships
(Table 2) shows that the faunas of the rainforests of southern
El Petén and lowland Alta Verapaz are closely related. The relationships
between these two areas and the Tikal-Uaxactún area[218] in northern El Petén is notably less. Apparently the biggest faunal
changes take place between southern El Petén and the Tikal-Uaxactún
area, and between the latter and Yucatán. As stated
by Stuart (1958:7) the Tikal-Uaxactún is transitional between the
humid rainforests to the south and the dry outer end of the Yucatán
Peninsula. The transitional nature of the environment is exemplified
by a rather depauperate herpetofauna consisting of some
species of both dry and humid environments and lacking a large
fauna typical of either. Contrariwise, the continuity of the environment
from southern El Petén to the lowlands of Alta Verapaz
is reflected in degree of resemblance of the herpetofaunas.

Table 2.—Index of Faunal Relationships Between Southern El Petén
and Other Regions.

Most of the species of amphibians and reptiles found in southern
El Petén are found in humid tropical forests from the Isthmus of
Tehuantepec southeastward on the Atlantic lowlands well into
Central America.

ACCOUNTS OF SPECIES

In the following pages various aspects of the occurrence, life
histories, ecology, and variation of the species of amphibians and
reptiles known from southern El Petén are discussed. Only Cochranella
fleischmanni reported by Stuart (1937) from Río Subín
at Santa Teresa was not collected by us and is excluded. Because
more worthwhile information was gathered for some species than
others, the length and completeness of the accounts vary. All
specimens listed are in the Museum of Natural History at the
University of Kansas, to which institution all catalog numbers refer.
Preceding the discussion of each species is an alphabetical list of[219] localities from which specimens were obtained; numbers after a
locality indicate the number of specimens obtained at each locality.

Bolitoglossa dofleini (Werner)

An adult female having minute ovarian eggs has a snout-vent
length of 81 mm., a tail length of 59 mm., 13 costal grooves, two
intercostal spaces between adpressed toes, 38-35 vomerine teeth
in irregular rows forming a broad arch from a point posterolaterad
to the internal nares to a point near the anterior edge of the
parasphenoid teeth, and 43-44 maxilliary-premaxillary teeth. In
life the dorsum was rusty brown with irregular black and orange
spots and streaks. The flanks were bluish gray with black in the
costal grooves and creamy tan flecks along the ventral edge of
the flank. The belly and underside of the tail were yellowish tan
with dark brown spots laterally. The limbs were orange proximally
and black distally; the pads of the feet were bluish black. The
dorsal and lateral surfaces of the tail were yellowish orange with
black spots. The iris was grayish yellow.

Stuart (1943:17) reported this species from Finca Volcán, Alta
Verapaz. He diagnosed his specimens as having 13 costal grooves
and two or three intercostal spaces between adpressed toes. He
stated that the vomerine teeth were about 12 in number and that
in life the dorsum was mottled gray and black, the sides gray and
brown, and the undersurfaces uniformly dark gray. These specimens
differ noticeably from the individual from Chinajá in the
number of vomerine teeth and in coloration.

In August, 1961, I obtained a specimen of Bolitoglossa dofleini at Finca Los Alpes, Alta Verapaz, approximately 13 kilometers
airline south-southwest of Finca Volcán and at approximately the
same elevation. Although the salamander was dead when found,
it obviously was more heavily pigmented than the individual from
Chinajá. The belly was bluish gray with black spots laterally;
the dorsum was dull brownish gray with some brownish red streaks.
The specimen is a female having small ovarian eggs, a snout-vent
length of 90 mm., 13 costal grooves, and two intercostal spaces
between adpressed limbs. There are 28-29 vomerine teeth, more
than twice as many as in specimens from Finca Volcán (Stuart,
1943:17), but noticeably fewer than in the specimen from Chinajá.

The presence of this species at Chinajá lends support to the idea
that the specimen from the Río de la Pasión listed by Brocchi[220] (1882:116) also is Bolitoglossa dofleini. Furthermore, the confirmed
presence of this species in the lowlands of El Petén suggests
that there may be genetic connection between B. dofleini in the
Alta Verapaz and B. yucatana in the Yucatán Peninsula. Bolitoglossa
yucatana differs from B. dofleini in having five intercostal
spaces between adpressed toes and in having a different color
pattern. Both are robust species having no close relationships to
other species of Bolitoglossa in northern Central America.

The specimen from Chinajá was found in water in the axil of
a large elephant-ear plant (Xanthosoma) by day in March. Its
stomach contained fragments of beetles and a large roach. The
natives did not know salamanders and had no name for them.

Bolitoglossa moreleti mulleri (Brocchi)

One specimen is a female having a snout-vent length of 80 mm.,
a tail length of 82 mm., and a total length of 162 mm. It contains
63 large eggs, the largest of which has a diameter of about three
millimeters. This specimen has 13 costal grooves, four intercostal
spaces between adpressed toes, and 12-13 vomerine teeth. A
juvenile having a snout-vent length of 39 mm. and a tail length
of 33 mm. has 12 costal grooves, three intercostal spaces between
adpressed toes, and 8-8 vomerine teeth. In life these salamanders
were uniformly dull brownish black above with a dull creamy
yellow irregular dorsal stripe beginning on the occiput and continuing
onto the tail. There are no yellow or orange streaks or
flecks on the head or limbs. The specimen from the Río San Román
was taken from the stomach of a Pliocercus euryzonus aequalis and has not been studied in detail, because of its poor condition.

The present specimens show no tendency for the development
of a broad irregular dorsal band that encloses black spots or forms
irregular dorsolateral stripes, as is characteristic of B. moreleti
mexicanus, a subspecies that has been reported from La Libertad
(Stuart, 1935:35) and Piedras Negras (Taylor and Smith, 1945:545)
in El Petén, and from Xunantunich, British Honduras (Neill and
Allen, 1959:20).

Schmidt (1936:151) and Stuart (1943:13) found B. moreleti
mulleri in bromeliads at Finca Samac, Alta Verapaz. Taylor and
Smith's (1945:545) and Neill and Allen's (1959:20) specimens of B. moreleti mexicanus were obtained from bromeliads, but Neill
and Allen (loc. cit.) stated that the natives in British Honduras[221] said that they had found salamanders beneath rubbish on the forest
floor. My specimens were obtained from beneath logs on the
forest floor in the rainy season. Possibly in drier environments the
species characteristically inhabits bromeliads, at least in the dry
season.

Bufo marinus (Linnaeus)

During both visits to Chinajá this large toad was breeding in
a small permanent pond in the camp. During the day the toads
took refuge in crevices beneath the buildings or beneath large
boulders by the pond. At dusk from four to ten males congregated
at the pond and called. Tadpoles of this species were in the pond
in March and in July. One juvenile was found beneath a rock in
the forest, and another was on the forest floor by day.

The natives' name for this species and the following one is sapo.

Bufo valliceps valliceps Wiegmann

This is one of the most abundant, or at least conspicuous,
amphibians inhabiting the forest. Breeding congregations were
found on February 24, March 2, March 11, and June 27. At these
times the toads were congregated at temporary ponds in the forest
or along small sluggish streams. Throughout the duration of both
visits to Chinajá individual males called almost nightly at the
permanent pond at the camp.

The variation in snout-vent length of 20 males selected at
random is 56.7 to 72.5 mm. (average, 64.8 mm.). Two adult females
have snout-vent lengths of 80.4 and 87.6 mm. In all specimens
the parotid glands are somewhat elongated and not rounded as in Bufo valliceps wilsoni (see Baylor and Stuart, 1961:199). My
observations on the condition of the cranial crests of the toads in
El Petén agree with the findings of Baylor and Stuart (op. cit.:198) in
that hypertrophied crests are usual in large females. In the shape
of the parotids and nature of the cranial crests the specimens from
El Petén are like those from the Isthmus of Tehuantepec in
México. As I pointed out (1960:53), the validity of the subspecies Bufo valliceps macrocristatus, described from northern Chiapas by
Firschein and Smith (1957:219) and supposedly characterized by
hypertrophied cranial crests, is highly doubtful.

In the toads from El Petén the greatest variation is in coloration.
The dorsal ground-color varies from orange and rusty tan to
brown, yellowish tan, and pale gray. In some individuals the[222] flanks and dorsum are one continuous color, whereas in others a
distinct dorsolateral pale colored band separates the dorsal color
from dark brown flanks. In some individuals the venter is
uniform cream color, in others it bears a few scattered black spots,
and in still others there are many spots, some of which are fused
to form a black blotch on the chest. In breeding males the vocal
sac is orange tan. All specimens have a coppery red iris.

Aside from the breeding congregations, active toads were found
on the forest floor at night; a few were there by day. Some
individuals were beneath logs during the day.

Eleutherodactylus rostralis (Werner)

Because of the multiplicity of names and the variation in coloration,
the small terrestrial Eleutherodactylus in southern México and
northern Central America are in a state of taxonomic confusion.
Stuart (1934:7, 1935:37, and 1958:17) referred specimens from El
Petén to Eleutherodactylus rhodopis (Cope). Stuart (1941b:197)
described Eleutherodactylus anzuetoi from Alta Verapaz and El
Quiché, Guatemala, suggested that the new species was an upland
relative of Eleutherodactylus rostralis (Werner), and used that
name for the frogs that he earlier had referred to Eleutherodactylus
rhodopis. Dunn and Emlen (1932:24) placed E. rostralis in the
synonymy of E. gollmeri (Peters). Examination of series of these
frogs from southern México, Guatemala, and Costa Rica causes me
to think that there are four species; these can be distinguished as
follows:

The presence of webbing between the toes, the absence of tarsal
tubercles, and the coppery red iris distinguish E. rostralis and E.
gollmeri from the other species. Probably E. rostralis and E. gollmeri are conspecific, but additional specimens are needed from Nicaragua
and Honduras to prove conspecificity. On the other hand, the characters
of the frogs from Chinajá clearly show that they are related
to E. gollmeri to the south and not to E. rhodopis to the north in
México.

At Chinajá, Eleutherodactylus rostralis was more abundant than[223] the few specimens indicate, for upon being approached the frogs
moved quickly and erratically, soon disappearing in the leaf litter
on the forest floor. Most of the specimens were seen actively moving
on the forest floor in the daytime; one was found beneath a rock,
and one was on the forest floor at night.

Eleutherodactylus rugulosus rugulosus (Cope)

These frogs were found on the forest floor by day. With the exception
of one female having a snout-vent length of 69.5 mm., all are
juveniles. The apparent rarity of this species at Chinajá may be
due to the absence of rocky streams, a favorite habitat of this frog.
The local name for this frog is sapito, meaning little toad.

Leptodactylus labialis (Cope)

One juvenile having a snout-vent length of 16.4 mm. was found
at night beside a pond in the forest. The scarcity of the species
of Leptodactylus in the southern part of El Petén probably is due
to the lack of permanent marshy ponds.

Leptodactylus melanonotus (Hallowell)

One individual was found beneath a rock beside a stream in
the forest. The local name is ranita, meaning little frog.

Syrrhophus leprus Cope

An adult female having a snout-vent length of 27.5 mm. was
found on the forest floor by day. Two juveniles having snout-vent
lengths of 15.5 and 19.0 mm. were beneath rocks on the forest floor.
The specimens are typical of the species as defined by Duellman
(1958:8).

Hyla ebraccata Cope

This small tree frog congregated in large numbers at a forest
pond at Toocog. Between June 30 and July 2 we collected specimens
and observed the breeding habits of this and other species
at the pond. Calling males were distributed around the pond, where
they called from low herbaceous vegetation at the edge of the pond
or from plants rising above the water. Calling commenced at dusk[224] and continued at least into the early hours of the morning. On one
occasion a female was observed at a distance of about 50 centimeters
away from a calling male sitting on a blade of grass. The
female climbed another blade of grass until she was about eight
centimeters away from the male, at which time he saw her, stopped
calling, jumped to the blade of grass on which she was sitting and
clasped her. Clasping pairs were observed on blades of grass and
leaves of plants above the water; most pairs were less than 50
centimeters above the surface of the pond.

The eggs are deposited on the dorsal surfaces of leaves above
the water. All eggs are in one plane (a single layer) on the leaf. External
membranes are barely visible, as the eggs consist of a single
coherent mass. Eggs in the yolk plug stage have diameters of 1.2
to 1.4 mm. Seventeen eggs masses were found; these contained from
24 to 76 (average 44) eggs. The jelly is extremely viscous and tacky
to the touch. At time of hatching the jelly becomes less viscous;
the tadpoles wriggle until they reach the edge of the leaf and drop
into the water.

Eleven tadpoles were preserved as they hatched; these have total
lengths of 4.5 to 5.0 (average 4.77) mm. Hatchling tadpoles are
active swimmers and have only a small amount of yolk. The largest
tadpoles preserved have total lengths of 13.0 and 13.5 mm. At
this size distinctive sword-tail and bright coloration have developed.

Fig. 2. Tadpole of Hyla ebraccata (KU 59986) from
Toocog, El Petén, Guatemala.

Description of fully developed tadpole (KU 59986): Total
length, 13.5 mm.; tail-length, 8.4 mm., 62 per cent of total length.
Snout, in dorsal view, bluntly rounded; in lateral view less bluntly
rounded; body depressed; head flattened; mouth terminal; eye large,
its diameter 25 per cent of length of body; nostrils near tip of
snout and directed anteriorly; spiracle sinistral and situated postero-ventrad
to eye; cloaca median. Tail-fin thrice depth of tail-musculature,
which extends beyond posterior end of tail-fin giving
sword-tail appearance (Fig. 2). In life, black stripe on each side
of body and on top of head; black band on anterior part of tail[225] and another on the posterior part; body and anterior part of tail
creamy yellow; dark red band between black bands on tail. Mouth
terminal, small, its width about one-fifth width of body; fleshy ridge
dorsally and ventrally; row of small papillae on ventral lip; no lateral
indentations of lips; upper beak massive, convex, and finely serrate;
lower beak small and mostly concealed behind upper; no teeth
(Fig. 3).

Fig. 3. Mouthparts of larval Hyla ebraccata (KU
59986) from Toocog, El Petén, Guatemala.

Hyla loquax Gaige and Stuart

These specimens were found at night when they were calling from
low vegetation in a forest pond. Most of the frogs were several
meters away from the edge of the pond. Although two clasping
pairs were found, we obtained no eggs or tadpoles referable to
this species.

Hyla microcephala martini Smith

The specimen from Chinajá was calling from a small bush at the
edge of a temporary grassy pond in a clearing in the forest. At
Toocog this species was closely associated with Hyla ebraccata;
males were calling from herbaceous vegetation in and around the
forest pond. These frogs were not so abundant in the forest at
Toocog as they were around ponds on the savanna at La Libertad.

[226]

Hyla picta (Günther)

This small tree frog was calling from herbs in a pond in the
forest on June 30 and July 2. The voice is weak; probably greater
numbers of males were present than are indicated by the few
specimens collected, for the din from the more vociferous species
made it impossible to hear Hyla picta unless one was calling
close by.

Hyla staufferi Cope

This individual was calling from a low bush in the clearing at
Chinajá. None was found in the pond in the forest at Toocog.
Stuart (1935:38) and Duellman (1960:63) noted that Hyla staufferi breeds early in the rainy season. Nevertheless, I think early breeding
habits do not account for the near absence of this species in
our collections from southern El Petén. In early July, 1960, a few
individuals were heard at a pond on the savanna at La Libertad. In
mid-July of the same year they were calling sporadically from
temporary ponds in the lower Motagua Valley. Possibly the individual
collected at Chinajá was accidentally transported there in
cargo from Toocog, from which camp at the edge of the savanna
planes fly to Chinajá weekly. My observations on this species
throughout its range in México and Central America indicate that it
inhabits savannas and semi-arid forests and usually is absent
from heavy rainforest. Stuart (1948:34) obtained this species at
Cubilquitz in the lowlands of Alta Verapaz.

Phyllomedusa callidryas taylori Funkhouser

Between June 30 and July 2 this species was abundant at a pond
in the forest at Toocog. Calling males were as high as five meters
in bushes and trees around the pond. At dusk males were observed
descending a vine-covered tree at the edge of the pond; this
strongly suggests that the frogs retreat to this tree and others like
it for diurnal seclusion. Clasping pairs were found on branches
and leaves above the water. The eggs are deposited in clumps
usually on vertical leaves, but sometimes on horizontal leaves or
on branches, vines, and aerial roots above the water. Twenty-six
clutches of eggs contained from 14 to 44 (average 29) eggs. In
a clutch in which the eggs are in yolk plug stage the average
diameter of the embryos is 2.3 mm. and that of the vitelline[227] membranes, 3.4 mm. Most of the eggs are in the external part
of the gelatinous mass; the jelly is clear. The yolk is pale green,
and the animal pole is brown. As development ensues, the yolk
becomes yellow and the embryo first dark brown and then pale
grayish tan. Upon hatching the tadpoles wriggle free of the
jelly and drop into the water. One clutch of 19 eggs was observed
to hatch in three minutes. Apparently, on dropping into the
water the hatchling tadpoles go to the bottom of the pond, for one
or two minutes pass from the time they enter the water until they
reappear near the surface. The average total length of seven
hatchling tadpoles is 7.4 mm. There is a moderate amount of
yolk, but this does not form a large ventral bulge. Large tadpoles
congregate in the sunny parts of the pond, where they were observed
just beneath the surface. Many had their mouths at the
surface. Except for constant fluttering of the tip of the tail, they
lie quietly with the axis of the body at an angle of about 45 degrees
with the surface of the water.

Description of tadpole (KU 60006): total length, 24.5 mm.; tail-length,
15.4 mm.; body broader than deep; head moderately flattened;
snout viewed from above blunt; nostrils close to snout and
directed dorsally; eyes of moderate size and directed laterally; mouth
directed anteroventrally; anus median; spiracle ventral, its opening
just to left of midline slightly more than one-half distance from tip
of snout to vent. Tail-fin slightly more than twice as deep as tail
musculature, which curves upward posteriorly; tail-fin narrowly
extending to tip of tail (Fig. 4). Color in life pale gray; in preservative
white with scattered melanophores; tail-fin transparent.

Fig. 4. Tadpole of Phyllomedusa callidryas taylori (KU 60006) from
Toocog, El Petén, Guatemala.

Upper lip having single row of papillae laterally, but none
medially; lower lip having single row of papillae; no lateral
indentation of lips; two or more rows of papillae at lateral corners
of lips; tooth-rows 2/3; second upper tooth row as long as first,
interrupted medially; inner lower tooth-row as long as upper rows,[228] interrupted medially; second and third lower rows decreasingly
shorter; upper beak moderate in size and having long lateral projections;
lower beak moderate in size; both beaks finely serrate
(Fig. 5).

Fig. 5. Mouthparts of larval Phyllomedusa callidryas taylori (KU
60006) from Toocog, El Petén, Guatemala.  

 

Smilisca baudini (Duméril and Bibron)

Individuals of this species were found at night sitting on bushes
and small trees in the forest in February and March and again in
June and July. One was in the axil of a leaf of a Xanthosoma. In
June and July males were heard nearly every night. The series of
specimens from 20 kilometers north-northwest of Chinajá was taken
from a breeding congregation in a shallow muddy pool in the
forest. Tadpoles of this species were in small, often muddy pools
in the forest. To my knowledge Smilisca baudini is the only hylid
to breed in these pools at Chinajá, although perhaps Smilisca
phaeota also utilizes them. The only other amphibian at Chinajá
known to breed in the pools is Bufo valliceps valliceps. Although
two specimens were on bushes at night at Toocog, Smilisca
baudini was not present at the pond where five other species of
hylids were breeding. Nevertheless, Smilisca baudini was calling
from two ponds on the savannas near La Libertad. All of the
specimens from southern El Petén have yellow or yellowish white
flanks and ventrolateral surfaces.

[229]

Smilisca phaeota cyanosticta (Smith)

All specimens were found in February and March. Those from
Chinajá were obtained from Xanthosoma and bromeliads; the
individual from 10 kilometers north-northwest of Chinajá is an
adult male that was calling from a puddle in a fallen tree on
March 13. A juvenile having a snout-vent length of 34.7 mm. lacks
the pale blue spots on the thighs; instead, the anterior and posterior
surfaces of the thighs are bright red.

Hypopachus cuneus nigroreticulatus Taylor

An adult male having a snout-vent length of 41.7 mm. was found
at night on the forest floor at the edge of a temporary pond. In
life the dorsum was dark brown with chocolate brown markings;
the stripe on the side of the head was white; the middorsal stripe
was pale orange; the belly was black and white, and the iris was
a bronze color.

Characteristically this species inhabits savannas and open forest;
thus, its occurrence in the rainforest at Toocog is surprising. This is
the southernmost record for the species in El Petén; to the south in
the highlands it is replaced by the smaller Hypopachus inguinalis,
having rounded, instead of compressed, metatarsal tubercles.

Rana palmipes Spix

With the exception of one recently metamorphosed juvenile having
a snout-vent length of 30.7 mm. that was found on the forest floor
by day on June 24, and one that was found beside a pool in a cave,
all individuals were found at temporary woodland pools or along
sluggish streams at night. The largest specimen is a female having
a snout-vent length of 107 mm.

Rana pipiens Schreber

All specimens were found near water at night. The largest individual
is a female having a snout-vent length of 112.5 mm.

Crocodylus moreleti Duméril and Duméril

One specimen was obtained from a quiet pool in the Río San[230] Román at night; another was found in a small sluggish stream at
Chinajá. Two large individuals were seen in tributaries to the Río
San Román. On the savannas at Toocog two small individuals were
obtained in the dry season, at which time the crocodiles apparently
were migrating to water. The local name for this species is lagarto.

Chelydra rossignoni (Bocourt)

The paucity of specimens of Chelydra from Central America has
resulted in rather inadequate diagnoses of various populations. The
present specimens have carapace lengths of 250 and 238 mm. and
plastral lengths of 185 and 176 mm. The length of carapace/bridge
ratio is 6.0 and 6.1 per cent. Each individual has four barbels, the
median pair of which are extremely long. In KU 55977 the lateral
pair of barbels is forked at the base. The relative length of the
plastral bridge in these specimens compares favorable with the
ratio (.06-.08) given by Schmidt (1946:4) for five specimens from
Honduras. Chelydra serpentina, which may occur sympatrically
with C. rossignoni in some parts of Central America, has a narrower
plastral bridge and only two barbels beneath the chin. Furthermore, C. rossignoni and C. osceola in Florida have long, flat tubercles
on the dorsal and lateral surfaces of the neck, whereas C.
serpentina has short, round tubercles.

The specimen from Chinajá was found in a small sluggish stream;
the other individual was in a muddy pool in the forest. The local
name is sambodanga.

Claudius angustatus Cope

One specimen was unearthed from the bank of a small muddy
stream by a bulldozer. This individual represents the second
record for the species in Guatemala; the first was provided by
specimens, likewise found in muddy waters, at Tikal (Stuart,
1958:19). The local name is caiman.

Kinosternon acutum Gray

These turtles were found on the forest floor, in small sluggish
streams, and in pools in the forest. One adult male had, in life,
the top of the head yellow with black spots; the stripes on the
head and neck were red. Specimens were obtained both in the dry[231] and rainy seasons. The local name for both species of Kinosternon is pochitoque.

Kinosternon leucostomum Duméril and Bibron

Individuals of this turtle were found on the forest floor and in
small sluggish streams. In life most specimens had a tan or
pale brown head with pinkish tan stripes on the head and neck.
All individuals were obtained in February and March. No ecological
differences between this species and K. acutum were evident.

Staurotypus triporcatus (Wiegmann)

This species is represented in the collection by one complete
shell found on the bank of the Río Subín. The carapace has a
length of 292 mm. The local name is Guao. Natives stated that
this turtle was not uncommon in clear rivers and lakes, a habitat
suggested for the species by Stuart (1958:19).

Dermatemys mawi Gray

The record from Chinajá is based on a carapace found in a
chiclero camp, where the turtle evidently had been brought for
food. The four specimens from the Río San Román were obtained
from edges of deep pools in clear water. In adult males the top
of the head was reddish orange in life. One of the specimens from
the Río San Román currently is living in the Philadelphia Zoological
Gardens. The local name for this turtle is tortuga blanca; it is
sought for its meat.

Geoemyda areolata (Duméril and Bibron)

Two specimens were obtained from dense forest at Chinajá.
The local name is mojina.

Pseudemys scripta ornata (Gray)

One subadult was obtained from clear water in the Río Subín.
The stripes on the head and neck were yellow; there was no red
"ear" on the side of the head. The stripes on the forelimbs were
orange, and the ocelli on the carapace were red. The local name
is jicotea.

[232]

Coleonyx elegans elegans Gray

One adult male having a snout-vent length of 89 mm. was found
beneath a log in the forest. Locally this gecko is known as escorpión; the natives believe it to be deadly poisonous. The use
of the name escorpión seems to be restricted to lizards thought to
be venomous. Nearly everywhere in México and Central America
some species of lizard carries this appellation. In El Petén I heard
the name used only for Coleonyx elegans and Thecadactylus rapicaudus;
in the lowlands of Guerrero, México, the name is applied
to geckos of the genus Phyllodactylus. The venomous lizards of
the genus Heloderma in the lowlands of western México are called escorpiónes. In the mountains of southern México various skinks
of the genus Eumeces, as well as lizards of the genus Xenosaurus,
carry the same appellation. Abronia in the mountains of México
and Gerrhonontus throughout México and Central America likewise
are called escorpiónes. Although many people in various parts of
Middle America consider most lizards poisonous, there is a unanimity
of opinion concerning the venomous qualities of the various
kinds of escorpiónes. I know of only two other lizards in Middle
America that are so uniformly regarded in native beliefs; these
are Enyaliosaurus clarki in the Tepalcatepec Valley in Michoacán,
called nopiche, and Phrynosoma asio in western México, called cameleón.

Sphaerodactylus lineolatus Lichtenstein

These small geckos were much more abundant than the few specimens
indicate. They frequently were seen on the trunks of corozo
palms, where they quickly took refuge in crevices at the bases of
the fronds. The specimen obtained at Toocog was under the bark
of a standing dead tree. In life the ventral surface of the tail was
orange. The individual from Chinajá was in the leaf litter on the
ground at the base of a dead tree.

Thecadactylus rapicaudus (Houttuyn)

Two specimens were found beneath the bark of standing dead
trees; another was found in the crack in the trunk of a mahogany
tree about 13 meters above the ground. In life the dorsum was[233] yellowish tan with dark brown markings; the venter was yellowish
tan with brown flecks, and the iris was olive-tan. The largest specimen
is a male having a snout-vent length of 95 mm.; all specimens
have regenerated tails. Individuals when caught twisted their
bodies and attempted to bite; upon grabbing a finger they held
on with great tenacity.

Anolis biporcatus (Wiegmann)

All specimens of this large anole were obtained from trees.
Some individuals were found in the tops of trees immediately after
they were felled. My limited observations on this anole suggest
that it is an inhabitant of the upper levels of the forest. In life an
adult male from 20 kilometers north-northwest of Chinajá was
brilliant green above; the eyelids were bright yellow; the belly was
white. The outer part of the dewlap was pale orange, and the
median part was pinkish blue. A juvenile having a snout-vent length
of 47 mm. and a tail length of 86 mm. was pale grayish green with
pale gray flecks on the dorsum. The largest male has a snout-vent
length of 98 mm. and a tail length of 217 mm.; the same measurements
of the largest female are 89 and 213 mm. This species, together
with all other anoles, is known locally as toloque.

Anolis capito Peters

All individuals were observed on trunks of trees between heights
of three and ten meters above the ground. The largest male has a
snout-vent length of 81 mm. and a tail length of 155 mm.; the same
measurements of the largest female are 87 and 150 mm. The
streaked brown dorsum, combined with the lizards' habit of pressing
the body against the trunks of trees, make this anole especially difficult
to see.

Anolis humilis uniformis Cope

This small dull brown anole is a characteristic inhabitant of the
forest floor, where the lizards move about in a series of quick,
short hops and thus easily evade capture. Three individuals were
found on small bushes, and four were on the bases of trees; otherwise,
all were observed on the ground. Observations indicate that[234] this species is active throughout the day, except during and immediately
after heavy rains. The males have a deep red dewlap
with a dark blue median spot.

Anolis lemurinus bourgeaei Bocourt

This moderate-sized anole characteristically inhabits the low
bushes and bases of trees in the forest. Individuals were most
readily observed on the buttresses of some of the gigantic mahogany
and ceiba trees. When approached the lizards usually ran around
the tree or ducked to the other side of the buttress; if the observer
moved closer, they jumped to the ground and ran off. None was
observed to ascend large trees. Some individuals were observed
foraging on the forest floor; these took shelter on the bases of
trees. One individual was sleeping on a palm frond at night. The
adult males have a uniformly orange-red dewlap.

Anolis limifrons rodriguezi Bocourt

In dry forests and more open situations than occur at Chinajá
this little anole is abundant, but in the wet forests of southern El
Petén, only three specimens were found. Two were on palm
fronds about two meters above the ground; the other was on a low
bush. I suspect that ecologically this species overlaps A. humilis
uniformis and A. lemurinus bourgeaei, but too few observations are
recorded to justify a definite statement at this time.

Anolis sericeus sericeus Hallowell

This small anole is common and widespread in the Atlantic
lowlands of southern México and northern Central America; usually
it inhabits sub-humid regions. Consequently, its presence in the
wet forests of southern El Petén was unexpected. The specimens
from Chinajá were sleeping on low bushes at night, whereas the
others were found on bushes by day.

Basiliscus vittatus Wiegmann

Individuals of this abundant species were most frequently seen in
dense bushes along the margins of rivers or small streams. None
was observed far from water. These lizards, like the anoles, are
known locally as toloque.

[235]

Corythophanes cristatus (Merrem)

Three individuals were found on tree trunks; the fourth was on
a thick vine about one meter above the ground. The two largest
males have snout-vent lengths of 121 and 115 mm. and tail lengths
of 265 and 243 mm. The largest female (KU 59603), obtained on
June 28, has a snout-vent length of 125 mm. and a tail length of
247 mm. This individual contained eight ova varying in greatest
diameter from 10.6 to 12.2 (average 11.1) mm. Also present are
numerous ovarian eggs having diameters up to about 3.5 mm.

One of the large males displayed a defensive behavior prior to
capture. When first observed the lizard was clinging to a tree
trunk about one and one-half meters above the ground. When I
approached, the lizard turned its flanks towards me; then it flattened
the body laterally, extended the dewlap, opened its mouth, and
made short rushing motions. When touched it bit viciously. On
the ground these lizards have a rather awkward bipedal gait that
is much slower than in Basiliscus vittatus.

In life an adult male (KU 55804) was reddish brown dorsally
with dark chocolate brown markings; the venter was creamy white,
and the iris was dark red. The natives call this lizard piende jente.

Iguana iguana rhinolopha Wiegmann

The iguana, as this lizard is called locally, seems to be uncommon
in the forested areas of southern El Petén. Possibly this is due to
the fact that the flesh of this lizard is relished as food by the natives.
My two specimens were in large trees at the edge of the river.

Laemanctus deborrei Boulenger

On June 26 a female having a snout-vent length of 129 mm.
and a tail length of 502 mm. was found on a bush in the forest.
The lizard, when approached, faced the collector and opened its
mouth. In life the dorsum was bright green; the lateral stripe
was white, and the iris was yellowish brown. This specimen contained
four ova having lengths of 13.4 to 14.2 (average 13.9) mm.

On June 30 at Toocog five white-shelled eggs were found in a
rotting log. Measurements of the eggs are—length, 23.5 to 25.0
(average 24.2) mm.; width, 15.0 to 15.5 (average 15.4) mm. These
eggs hatched on August 30. The five young had snout-vent lengths[236] of 43 to 45 (average 44) mm., and tail lengths of 137 to 140 (average
138) mm. In life the hatchlings had a dull dark green dorsum,
pale bright green venter and stripes on head, and reddish brown
iris. In preservative the hatchlings are creamy tan above with
five or six square dark brown blotches middorsally.

The natives consider this lizard to be one of the anoles; consequently,
it is known as toloque.

Lepidophyma flavimaculatum flavimaculatum Duméril

Individuals were found beneath logs on the forest floor or moving
about in the litter on the forest floor. One was observed crawling
across a trail during a heavy rain. In some adults the tan dorsal
spots are large and distinct; in others the spots are small and indistinct.
Two juveniles, apparently recent hatchlings, were found
on June 28 and July 5. These specimens have snout-vent lengths
of 29 mm. and tail lengths of 38 and 41 mm.

Eumeces schwartzei Fischer

One specimen (KU 59551) was found on the forest floor at midday;
it is an adult female having a snout-vent length of 125 mm.
and a tail length of 210 mm. This specimen is larger than those recorded
by Taylor (1936:99) and extends the known range of the
species south of Ramate, approximately 125 kilometers south-south-westward
to Chinajá.

Eumeces sumichrasti (Cope)

One adult male having a snout-vent length of 82 mm. was found
beneath a palm frond on the forest floor. In life the dorsum was
dull brown; the chin was cream; the belly was yellow, and the underside
of the tail was orange. A juvenile having a black body, yellow
dorsal stripes, and a bright blue tail was observed on the forest floor.

Scincella cherriei cherriei (Cope)

All individuals of this lizard were found in the leaf litter on the
forest floor; many escaped capture. In life the tail is dull bluish
gray. The number of dorsal scales varies from 59 to 61 (average[237] 60); thus, these specimens fall within the range of variation of S.
cherriei cherriei, and thereby differ from S. cherriei stuarti to the
west and S. cherriei ixbaac to the north.

Ameiva festiva edwardsi Bocourt

This abundant terrestrial lizard, locally called lagartijo, is found
throughout the forest. A juvenile obtained on March 14 at Sayaxché
has a snout-vent length of 42 mm. and a prominent umbilical scar.
Other juveniles were observed at Chinajá in February and March,
thereby indicating that the young probably hatch in the early part
of the year. Juveniles have bright blue tails.

Celestus rozellae Smith

Two specimens were obtained from trees by workmen in
February. These lizards have snout-vent lengths of 70 and 83 mm.
and tail lengths of 133 and 135 mm. There are 21 and 23 lamellae
beneath the fourth toe; each has 31 longitudinal rows of scales
around the body.

Boa constrictor imperator Daudin

All specimens were found on the forest floor. One individual
was found in combat with a large Drymarchon corais melanurus.
Apparently, the Drymarchon was attempting to devour the Boa,
which had a total length of 1683 mm. Locally this snake is called masacuata; it is one of the few snakes believed by the local inhabitants
to be non-poisonous.

Clelia clelia clelia Daudin

One specimen is represented only by the head; the snake was
killed on the forest floor by workmen. Another individual was
found in a pool of water at the base of a limestone outcropping in
the forest; this specimen (KU 58167) is a female having a body
length of 2220 mm. and a total length of 2634 mm. This snake
contained 22 ova averaging 56 × 23 mm. Both specimens were
uniform shiny black above and cream-color below. The local name
is sumbadora.

[238]

Coniophanes bipunctatus bipunctatus (Günther)

This snake was found on the forest floor by day; it is a male
having 130 ventrals, an incomplete tail; cream-colored belly, and a
pair of large brown spots on each ventral scute.

Coniophanes fissidens fissidens (Günther)

This male specimen was found beneath a rock in a sink hole.
It has 122 ventrals and 77 caudals. A narrow temporal stripe
extends along the upper edge of the anterior temporal and the
lower edge of the upper secondary temporal. The belly is ashy
white with a pair of small black spots on each ventral.

Coniophanes imperialis clavatus (Peters)

All specimens were found on the forest floor by day. These small
snakes are capable of rapid movement and quickly disappear in
the litter on the ground. Two individuals evaded capture. The
belly is creamy white anteriorly and vermillion red posteriorly.

Dryadophis melanolomus laevis (Fischer)

These snakes, locally known as sumbadora, were found on the
forest floor; two others were seen, but escaped. The variation in
coloration has been a source of confusion in this species in northern
Central America (see Stuart, 1941:86). All of the present specimens
are males: KU 55709 has 178 ventrals, 121 caudals, and a total
length of 914 mm.; the dorsum is olive-tan with six darker cross-bars
on the neck; the belly is creamy white. KU 58160 has 188
ventrals, 123 caudals, and a total length of 1365 mm.; the dorsum is
uniform olive-brown, except that some dorsal scales at midbody
have black anterior borders like D. melanolomus melanolomus has
in the Yucatán Peninsula; the venter is pale yellow. KU 58158 has
179 ventrals, 122 caudals, and a total length of 723 mm.; the dorsum
is rich chocolate brown with eight dark cross-bars on the neck; the
belly is bright orange.

Stuart (1941a:87) stated that in life two distinct color phases
were observed in specimens collected by him in Alta Verapaz,
Guatemala. One had an olive-brown dorsum and the other, a[239] reddish orange dorsum. Stuart made no mention of variation in
the color of the venter. Similar variation is known in D. melanolomus
alternatus in Costa Rica, where some individuals have orange-red
venters. This color phase has been recognized as a distinct
species, Dryadophis sanguiventris, by Taylor (1954:722). Examination
of 18 specimens from Costa Rica shows no differences in
scutellation, nor geographic segregation of two populations. I am
convinced that the red-bellied Dryadophis in Costa Rica, like those
in Guatemala, represent a color phase of the subspecies inhabiting
those areas and that Dryadophis sanguiventris Taylor is a synonym
of Dryadophis melanolomus alternatus (Bocourt).

Drymarchon corais melanurus (Duméril, Bibron and Duméril)

The specimen from Sayaxché was found at the edge of a clearing
in the forest; that from 15 kilometers northwest of Chinajá was found
on the forest floor coiled with a Boa constrictor imperator, which
the Drymarchon apparently was trying to eat. The Drymarchon is
a giant specimen having a total length of 2950 mm. (see Duellman,
1961:368). The Boa with which it was coiled has a total length
of 1683 mm. I was attracted to the snakes by a loud thrashing
noise. When I approached the writhing mass, the snakes separated,
but I was able to see that the Drymarchon had its teeth firmly imbedded
in the posterior part of the head of the Boa. From the Drymarchon I forced the regurgitation of a recently ingested Bothrops nummifer nummifer having a total length of 953 mm.
These observations show that the snake-eating capabilities of Drymarchon can hardly be over-estimated.

In both Drymarchon the anterior one-half of the body is olive-tan,
which changes to bluish black posteriorly. The local name is sumbadora.

Drymobius margaritiferus margaritiferus (Schlegel)

All individuals were obtained in clearings in the forest by day
in the rainy season. Two individuals each contained a Similisca
baudini and another contained a Bufo valliceps valliceps. Locally
this snake is known by the appropriate name of ranera.

Imantodes cenchoa leucomelas Cope

With the exception of one that was found dead in camp, all[240] individuals were taken from low vegetation by day. The dorsum
is creamy tan with 28 to 35 (average 32) chocolate brown blotches,
and the venter is ashy white with small brown flecks. Three males
have 238 to 248 (average 244) ventrals and 148 to 154 (average
151) caudals; one female has 239 ventrals and 142 caudals. The
largest specimen, a male, has a body length of 660 mm. and a total
length of 943 mm.

Lampropeltis doliata polyzona Cope

One female (KU 57156) having 230 ventrals and 54 caudals
was found on the forest floor by day. This individual has a black
snout with a white bar across the nasals and prefrontals, a white
spot in the middle of the frontal, and a white band across the
temporals and parietals that is bordered posteriorly by a black
band. There are 28 white and 28 red rings on the body. The
tips of the red scales are darkened. The black rings between the
white and red rings are not so expanded as to interrupt the white
rings dorsally as in L. doliata abnorma as identified by Stuart
(1948:70). Locally this snake, like all red, black, and white or
yellow banded snakes, is called coral or coralillo.

Leptodeira frenata malleisi Dunn and Stuart

This specimen, a male having 173 ventrals and 69 caudals, was
found beneath the bark on a log in the forest. In life the dorsum
was pinkish tan with 36 chocolate brown blotches on the body;
the venter was rosy pink.

Leptodeira septentrionalis polysticta Günther

If numbers of specimens are indicative of abundance, this is the
most common snake in southern El Petén. All were found at night
in the rainy season. At a pond in the forest at Toocog these snakes
were observed on low vegetation, on the ground, and in the water.
Evidently they congregate at breeding choruses of frogs. One Leptodeira contained a Smilisca baudini and another contained
eggs of Phyllomedusa callidryas taylori. The natives call this snake nahuyaca.

[241]

Leptophis ahaetulla praestans (Cope)

Both specimens were obtained from trees when they were felled.
One individual (KU 55716) has a body length of 1345 mm. and a
total length of 2035 mm. In life the entire snake was uniform bright
green; the eye was yellow. In preservative the dorsum is dark
blue, and the venter is green.

Leptophis mexicanus mexicanus Duméril, Bibron and Duméril

All specimens came from low trees in the forest. The largest
specimen is a male having a body length of 724 mm. and a total
length of 1236 mm. In life the middorsum was a golden tan; the
top of the head was a vivid green. One individual had ingested a Smilisca baudini. The local name is bejuquillo.

Ninia sebae sebae (Duméril, Bibron and Duméril)

This specimen, a male having 144 ventrals and 55 caudals, was
found beneath bark on a log in the forest. There is a black band
five scales in length on the nape followed posteriorly by a red
band six scales in length and then by a complete black band one
and one-half scales in length. The rest of the body is dull red with
16 incomplete black bands one to one and one-half scales in length
on the anterior two-thirds of the body.

Oxybelis aeneus aeneus (Wagler)

One individual was found in a low tree; the other was in a
bush. Both specimens are males; the largest has a body length
of 754 mm. and a total length of 1286 mm. Bogert and Oliver
(1945:388) distinguished O. aeneus aeneus in Central and South
America from O. aeneus auratus in México in that the diameter of
the eye is more than the length of the internasal, whereas in O.
aeneus auratus the diameter of the eye is less than the length of the
internasal. Stuart (1958:27) stated that on the basis of this
character three specimens from Tikal in northeastern El Petén
definitely were O. aeneus aeneus. Of the present specimens from
southern El Petén, one has an internasal:eye ratio of 1.08; the other
has a ratio of 0.87. A careful review of these snakes is needed to
verify the validity of the characters used to separate the subspecies[242] and to determine areas of intergradation. The local name for the
vine-snake is bejuquillo.

Pliocercus euryzonus aequalis Salvin

These specimens are tentatively referred to P. euryzonus. KU
57160 is a female having 130 ventrals, 87 caudals, and 23 black
rings on the body; KU 58150 is a juvenile having 128 ventrals, 79
caudals, and 27 black rings on the body. In both specimens the
tip of the snout is yellow; a broad yellow band on the parietals and
temporals is bordered posteriorly by a black band on the nape.
The black rings on the body are not bordered by yellow, but
black rings on the tail have yellow borders ventrally. In the red
interspaces between the black rings, black flecks and spots, especially
posteriorly, tend to form secondary black rings (Fig. 6a).
According to Stuart (1948:71), P. euryzonus aequalis has 25 to 27
black rings on the body, whereas P. elapoides salvini, which also
occurs in El Petén, has 15 to 23 black rings.

Fig. 6. Dorsal color patterns of Pliocercus euryzonus aequalis (A) and Micrurus
affinis apiatus (B).

 

The specimen from the Río San Román contained a partly
digested Bolitoglossa moreleti mulleri. Locally Piocercus is called coral or coralillo.

[243]

Pseustes poecilonotus poecilonotus (Günther)

Two juveniles were on the forest floor; one juvenile and an adult
were on low bushes. The juveniles have a tan dorsum with reddish
brown blotches; the belly is gray, and the iris is cream-color
above and brown below. The one adult is olive-brown above and
creamy white below on the anterior three-fourths of the body;
posteriorly it is black above and below. There are no paravertebral
dark stripes nor pale spots on the dorsal scales.

Two specimens (one juvenile and the adult) when encountered
compressed the anterior part of the body laterally and struck repeatedly.
Locally the adults are called sumbadora.

Sibon dimidiata dimidiata (Günther)

Both snakes were obtained from trees when they were felled.
In life the dorsum was pinkish orange with dark chocolate brown
blotches narrowly edged with black.

Sibon nebulata nebulata (Linnaeus)

This specimen, a male having a body length of 544 mm. and
a tail length of 198 mm., was found in a felled tree. In life the
belly was pink and black; the dorsal black blotches were narrowly
outlined with pink.

Spilotes pullatus mexicanus (Laurenti)

This large snake, locally called mica, seems to be equally at
home on the ground and in low trees and bushes. It is fast moving
for a large snake; two individuals escaped capture. The natives
said that this snake eats other snakes, but examination of stomachs
revealed no supporting evidence.

Stenorrhina degenhardti (Berthold)

This specimen, a female having 158 ventrals, 37 caudals, and a
total length of 489 mm., was found on the forest floor. On the
olive-brown dorsum are 27 irregular, narrow, dark brown, transverse
bands. The head is uniform olive-brown; the chin and labials are[244] cream-color. The venter is cream-color with a row of brown spots
forming a midventral stripe. A large spider was found in the
stomach.

I have refrained from assigning a subspecific name to this snake.
Cursory examination of specimens from throughout México and
Central America reveals a bewildering array of variation in coloration
that suggests that the subspecies mexicanus is not recognizable,
or that two species occur sympatrically in parts of southern México
and northern Central America.

Tretanorhinus nigroluteus lateralis Bocourt

A single male having 136 ventrals, 75 caudals, and a total length
of 407 mm. was found by a stream in camp. The dorsum is pale
grayish tan with 34 pairs of small chocolate brown spots, some of
the anterior ones of which are connected across the back. A cream-colored
lateral stripe is on the third and fourth dorsal scale-rows
anteriorly and the second and third rows posteriorly. The lower
dorsal scale rows are black. The venter is dark grayish brown with
cream-colored flecks anteriorly and creamy gray posteriorly where
the dark color is restricted to the midventral region and the lateral
edges of ventrals and first dorsal scale-row.

Xenodon rabdocephalus mexicanus Smith

Both individuals were found on the forest floor. An adult male
having a total length of 420 mm. has a cream-colored venter with
brown flecks. A juvenile having a total length of 172 mm. has a
creamy white belly with black crossbands.

At the suggestion of L. C. Stuart, I am following Schmidt
(1941:501) in placing X. mexicanus as a subspecies of X. rabdocephalus.

Micrurus affinis apiatus (Jan)

All specimens were found beneath litter on the forest floor.
All are males having 202 to 211 (average 205) ventrals, 53 to 56
(54.6) caudals, and 34 to 48 (41) primary black rings on the body.
There are no yellow rings, and black spots in the red interspaces
tend to form secondary black rings (Fig. 6b), the same as in Pliocercus euryzonus aequalis. The local name is coral or coralillo.

[245]

Bothrops atrox asper (Garman)

Although we found only two specimens, natives and workmen
at the camp at Chinajá stated that the barba amarilla, as this snake
is known locally, had been abundant when the camp had been
established less than two years before our visit.

Bothrops nasutus Bocourt

This specimen, a male having a total length of 415 mm., was
found on the forest floor. The dorsum is brown with dark brown
blotches separated middorsally by a narrow orange-tan stripe extending
from the nape to the base of the tail. The belly is grayish
tan with white flecks on the lateral edges of the ventrals. The local
name is nahuyaca.

Bothrops nummifer nummifer (Rüppell)

Two individuals were found on the forest floor, and one adult,
having a total length of 953 mm., was removed from the stomach
of a large Drymarchon corais melanurus. There is considerable
variation in color and pattern. A juvenile (KU 58104), having a
total length of 332 mm., has a tan dorsum with 19 interconnected
dark brown, diamond-shaped, middorsal blotches, the lateral extensions
of which are black; the belly is a cream-color with brown
squares. An adult female (KU 55706), having a total length of
779 mm., has a dorsal coloration like the preceding specimen, except
that the lateral extensions of the dorsal blotches are brown;
the belly is a uniform cream-color. A second adult female (KU
55707), having a total length of 953 mm., has a brown dorsum
with 21 interconnected black, diamond-shaped, middorsal blotches,
the lateral extensions of which are black; the belly is a cream-color
with black squares.

The local name for this species is braza de piedra.

Bothrops schlegeli schlegeli (Berthold)

This specimen was taken from the thatched roof of a house at
the edge of the forest and contained the remains of a small mammal.
The local name is nahuyaca.

[246]

HYPOTHETICAL LIST OF SPECIES

Listed below are thirteen species that have not been found in
southern El Petén but that probably occur there.

SUMMARY

A study of the amphibians and reptiles in the rainforests of
southern El Petén, Guatemala, reveals the presence of 78 species;
an additional 13 species probably occur there. In this tropical area
having a high amount of rainfall most of the species of amphibians
and reptiles have extensive ranges in the wet forests on the Atlantic
lowlands of southern México and northern Central America; some
species that more frequently are found in sub-humid forests also
occur.

Ecologically the fauna is divided into five major habitats—aquatic,
aquatic margin, fossorial, terrestrial, and arboreal. Forty-two
per cent of the 78 species are wholly or partly arboreal. The
fauna is most closely related to that in Alta Verapaz, Guatemala,
but includes many species that occur in the Tikal-Uaxactún area in
northeastern Guatemala.

Eleutherodactylus rostralis (Werner) and E. rhodopis (Cope)
are redefined and their relationships are suggested. The color
phases of Dryadophis melanolomus laevis and D. m. alternatus are
discussed; Dryadophis sanguiventris Taylor is synonymized with Dryadophis melanolomus alternatus (Bocourt).

The breeding habits, eggs, and tadpoles of the hylid frogs Hyla
ebraccata and Phyllomedusa callidryas taylori are described, as
are the eggs and juveniles of Laemanctus deborrei.

LITERATURE CITED

Baylor, E. R. and Stuart, L. C.

Bogert, C. M. and Oliver, J. A.

Brocchi, P.

[248]

Duellman, W. E.

Dunn, E. R. and Emlen, J. T.

Firschein, I. L. and Smith, H. M.

Lundell, C. L.

Neill, W. T. and Allen, R.

Sapper, K.

Schmidt, K. P.

Simpson, G. G.

Smith, H. M. and Taylor, E. H.

Stuart, L. C.

Taylor, E. H.

Taylor, E. H. and Smith, H. M.

Transmitted November 29, 1962.

29-5935

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8. Comments on the taxonomic status of Apodemus peninsulae, with description

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10. A new bat (Genus Leptonycteris) from Coahuila. By Howard J. Stains.

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11. A new species of pocket gopher (Genus Pappogeomys) from Jalisco, Mexico.

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12. Geographic variation in the pocket gopher, Thomomys bottae, in Colorado.

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13. New bog lemming (genus Synaptomys) from Nebraska. By J. Knox Jones,

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14. Pleistocene bats from San Josecito Cave, Nuevo León, México. By J. Knox

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17. Distribution, variation, and relationships of the montane vole, Microtus montanus.

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5. Birds found on the Arctic slope of northern Alaska.  By James W. Bee.

Pp. 163-211, plates 9-10, 1 figure in text. March 12, 1958.



*6. The wood rats of Colorado: distribution and ecology. By Robert B. Finley,

Jr. Pp. 213-552, 34 plates, 8 figures in text, 35 tables. November 7, 1958.



7. Home ranges and movements of the eastern cottontail in Kansas. By Donald

W. Janes. Pp. 553-572, 4 plates, 3 figures in text. May 4, 1959.



8. Natural history of the salamander, Aneides hardyi. By Richard F. Johnston

and Gerhard A. Schad. Pp. 573-585. October 8, 1959.



9. A new subspecies of lizard, Cnemidophorus sacki, from Michoacán, México.

By William E. Duellman. Pp. 587-598, 2 figures in text. May 2, 1960.



10. A taxonomic study of the middle-American snake, Pituophis deppei. By

William E. Duellman. Pp. 599-610, 1 plate, 1 figure in text. May 2, 1960.



Index. Pp. 611-626.



Vol. 11. Nos. 1-10 and index. Pp. 1-703, 1958-1960.



Vol. 12.  1. Functional morphology of three bats: Sumops, Myotis, Macrotus. By Terry

A. Vaughan. Pp. 1-153, 4 plates, 24 figures in text. July 8, 1959.



*2. The ancestry of modern Amphibia: a review of the evidence. By Theodore

H. Eaton, Jr. Pp. 155-180, 10 figures in text. July 10, 1959.



3. The baculum in microtine rodents. By Sydney Anderson. Pp. 181-216, 49

figures in text. February 19, 1960.



*4. A new order of fishlike Amphibia from the Pennsylvanian of Kansas. By

Theodore H. Eaton, Jr., and Peggy Lou Stewart. Pp. 217-240, 12 figures in

text. May 2, 1960.



5. Natural history of the bell vireo. By Jon C. Barlow. Pp. 241-296, 6 figures

in text. March 7, 1962.



6. Two new pelycosaurs from the lower Permian of Oklahoma. By Richard C.

Fox. Pp. 297-307, 6 figures in text. May 21, 1962.



7. Vertebrates from the barrier island of Tamaulipas, México. By Robert K.

Selander, Richard F. Johnston, B. J. Wilks, and Gerald G. Raun. Pp. 309-345,

pls. 5-8. June 18, 1962.



8. Teeth of Edestid sharks. By Theodore H. Eaton, Jr. Pp. 347-362, 10 figures

in text. October 1, 1962.



More numbers will appear in volume 12.



Vol. 13.  1. Five natural hybrid combinations in minnows (Cyprinidae). By Frank B.

Cross and W. L. Minckley. Pp. 1-18. June 1, 1960.



2. A distributional study of the amphibians of the Isthmus of Tehuantepec,

México. By William E. Duellman. Pp. 19-72, pls. 1-8, 3 figures in text.

August 16, 1960.



3. A new subspecies of the slider turtle (Pseudemys scripta) from Coahuila,

México. By John M. Legler. Pp. 73-84, pls. 9-12, 3 figures in text. August

16, 1960.



4. Autecology of the copperhead. By Henry S. Fitch. Pp. 85-288, pls. 13-20,

26 figures in text. November 30, 1960.



5. Occurrence of the garter snake, Thamnophis sirtalis, in the Great Plains and

Rocky Mountains. By Henry S. Fitch and T. Paul Maslin. Pp. 289-308,

4 figures in text. February 10, 1961.



6. Fishes of the Wakarusa river in Kansas. By James E. Deacon and Artie L.

Metcalf. Pp. 309-322, 1 figure in text. February 10, 1961.



7. Geographic variation in the North American cyprinid fish, Hybopsis gracilis.

By Leonard J. Olund and Frank B. Cross. Pp. 323-348, pls. 21-24, 2 figures

in text. February 10, 1961.



8. Descriptions of two species of frogs, genus Ptychohyla; studies of American

hylid frogs, V. By William E. Duellman. Pp. 349-357, pl. 25, 2 figures

in text. April 27, 1961.



9. Fish populations, following a drought, in the Neosho and Marais des Cygnes

rivers of Kansas. By James Everett Deacon. Pp. 359-427, pls. 26-30, 3 figs.

August 11, 1961.



10. Recent soft-shelled turtles of North America (family Trionychidae). By

Robert G. Webb. Pp. 429-611, pls. 31-54, 24 figures in text. February

16, 1962.



Index. Pp. 613-624.



Vol. 14.  1. Neotropical bats from western México. By Sydney Anderson. Pp. 1-8.

October 24, 1960.



2. Geographic variation in the harvest mouse. Reithrodontomys megalotis, on

the central Great Plains and in adjacent regions. By J. Knox Jones, Jr.,

and B. Mursaloglu. Pp. 9-27, 1 figure in text. July 24, 1961.



3. Mammals of Mesa Verde National Park, Colorado. By Sydney Anderson.

Pp. 29-67, pls. 1 and 2, 3 figures in text. July 24, 1961.



4. A new subspecies of the black myotis (bat) from eastern Mexico. By E.

Raymond Hall and Ticul Alvarez. Pp. 69-72, 1 figure in text. December

29, 1961.



5. North American yellow bats, "Dasypterus," and a list of the named kinds

of the genus Lasiurus Gray. By E. Raymond Hall and J. Knox Jones, Jr.

Pp. 73-98, 4 figures in text. December 29, 1961.



6. Natural history of the brush mouse (Peromyscus boylii) in Kansas with

description of a new subspecies. By Charles A. Long. Pp. 99-111, 1 figure

in text. December 29, 1961.



7. Taxonomic status of some mice of the Peromyscus boylii group in eastern

Mexico, with description of a new subspecies. By Ticul Alvarez. Pp. 113-120,

1 figure in text. December 29, 1961.



8. A new subspecies of ground squirrel (Spermophilus spilosoma) from Tamaulipas,

Mexico. By Ticul Alvarez. Pp. 121-124. March 7, 1962.



9. Taxonomic status of the free-tailed bat, Tadarida yucatanica Miller. By J.

Knox Jones, Jr., and Ticul Alvarez. Pp. 125-133, 1 figure in text. March 7,

1962.



10. A new doglike carnivore, genus Cynaretus, from the Clarendonian Pliocene,

of Texas. By E. Raymond Hall and Walter W. Dalquest. Pp. 135-138,

2 figures in text. April 30, 1962.



11. A new subspecies of wood rat (Neotoma) from northeastern Mexico. By

Ticul Alvarez. Pp. 139-143. April 30, 1962.



12. Noteworthy mammals from Sinaloa, Mexico. By J. Knox Jones, Jr., Ticul

Alvarez, and M. Raymond Lee. Pp. 145-159, 1 figure in text. May 18,

1962.



13. A new bat (Myotis) from Mexico. By E. Raymond Hall. Pp. 161-164,

1 figure in text. May 21, 1962.



14. The mammals of Veracruz. By E. Raymond Hall and Walter W. Dalquest.

Pp. 165-362, 2 figures. May 20, 1963.



15. The recent mammals of Tamaulipas, México. By Ticul Alvarez. Pp. 363-473,

5 figures in text. May 20, 1963.



More numbers will appear in volume 14.



Vol. 15.  1. The amphibians and reptiles of Michoacán, México. By William E. Duellman.

Pp. 1-148, pls. 1-6, 11 figures in text. December 20, 1961.



2. Some reptiles and amphibians from Korea. By Robert G. Webb, J. Knox

Jones, Jr., and George W. Byers. Pp. 149-173. January 31, 1962.



3. A new species of frog (Genus Tomodactylus) from western México. By

Robert G. Webb, Pp. 175-181, 1 figure in text. March 7, 1962.



4. Type specimens of amphibians and reptiles in the Museum of Natural History,

the University of Kansas. By William E. Duellman and Barbara Berg.

Pp. 183-204. October 26, 1962.



5. Amphibians and Reptiles of the Rainforests of Southern El Petén, Guatemala.

By William E. Duellman. Pp. 205-249, pls. 7-10, 6 figures in text. October

4, 1963.



More numbers will appear in volume 15.