Amphibians y Reptiles Guyana Keys

Amphibians and reptiles of Guyana, South America: illustrated
keys, annotated species accounts, and a biogeographic synopsis

Author(s): Charles J. Cole , Carol R. Townsend , Robert P. Reynolds , Ross D.
MacCulloch , and Amy Lathrop
Source: Proceedings of the Biological Society of Washington, 125(4):317-578.
2013.
Published By: Biological Society of Washington
DOI: http://dx.doi.org/10.2988/0006-324X-125.4.317
URL: http://www.bioone.org/doi/full/10.2988/0006-324X-125.4.317
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
125(4):317–620. 2013.
Amphibians and reptiles of Guyana, South America: illustrated keys,

annotated species accounts, and a biogeographic synopsis
Charles J. Cole*, Carol R. Townsend, Robert P. Reynolds, Ross D. MacCulloch,
and Amy Lathrop
(CJC, CRT) Division of Vertebrates (Herpetology), American Museum of Natural History, 200
Central Park West, New York, New York 10024, U.S.A., e-mail: [email protected],
[email protected];
(RPR) Biological Survey Unit, United States Geological Survey, Patuxent Wildlife Research
Center, National Museum of Natural History, P.O. Box 37012, MRC 111,
Washington, D.C. 20013-7012, U.S.A., e-mail: [email protected];
(RDM, AL) Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, 100
Queen’s Park, Toronto, Ontario, M5S 2C6, Canada, e-mail: [email protected], [email protected]
Abstract.—Guyana has a very distinctive herpetofauna. In this first ever

detailed modern accounting, based on voucher specimens, we document the
presence of 324 species of amphibians and reptiles in the country; 148
amphibians, 176 reptiles. Of these, we present species accounts for 317 species
and color photographs of about 62% (Plates 1–40). At the rate that new
species are being described and distributional records are being found for the
first time, we suspect that at least 350 species will be documented in a few
decades.
The diverse herpetofauna includes 137 species of frogs and toads, 11
caecilians, 4 crocodylians, 4 amphisbaenians, 56 lizards, 97 snakes, and 15
turtles. Endemic species, which occur nowhere else in the world, comprise 15%
of the herpetofauna. Most of the endemics are amphibians, comprising 27% of
the amphibian fauna. Type localities (where the type specimens or scientific
name-bearers of species were found) are located within Guyana for 24% of the
herpetofauna, or 36% of the amphibians. This diverse fauna results from the
geographic position of Guyana on the Guiana Shield and the isolated
highlands or tepuis of the eastern part of the Pantepui Region, which are
surrounded by lowland rainforest and savannas. Consequently, there is a
mixture of local endemic species and widespread species characteristic of
Amazonia and the Guianan Region.
Although the size of this volume may mislead some people into thinking
that a lot is known about the fauna of Guyana, the work has just begun.
Many of the species are known from fewer than five individuals in scientific
collections; for many the life history, distribution, ecology, and behavior
remain poorly known; few resources in the country are devoted to developing
such knowledge; and as far as we are aware, no other group of animals in the
fauna of Guyana has been summarized in a volume such as this to document
the biological resources.
We briefly discuss aspects of biogeography, as reflected in samples collected
at seven lowland sites (in rainforest, savanna, and mixed habitats below 500 m
elevation) and three isolated highland sites (in montane forest and evergreen
high-tepui forest above 1400 m elevation). Comparisons of these sites are
preliminary because sampling of the local faunas remains incomplete.
* Corresponding author.
318 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Nevertheless, it is certain that areas of about 2.5 km2 of lowland rainforest can
support more than 130 species of amphibians and reptiles (perhaps actually
more than 150), while many fewer species (fewer than 30 documented so far)
occur in a comparable area of isolated highlands, where low temperatures,
frequent cloudiness, and poor soils are relatively unfavorable for amphibians
and reptiles. Furthermore, insufficient study has been done in upland sites of
intermediate elevations, where lowland and highland faunas overlap
significantly, although considerable work is being accomplished in Kaieteur
National Park by other investigators.
Comparisons of the faunas of the lowland and isolated highland sites
showed that very few species occur in common in both the lowlands and
isolated highlands; that those few are widespread lowland species that tolerate
highland environments; that many endemic species (mostly amphibians) occur
in the isolated highlands of the Pakaraima Mountains; and that each of the
isolated highlands, lowland savannas, and lowland rainforests at these 10 sites
have distinctive faunal elements. No two sites were identical in species
composition. Much more work is needed to compare a variety of sites, and
especially to incorporate upland sites of intermediate elevations in such
comparisons.
Five species of sea turtles utilize the limited areas of Atlantic coastal
beaches to the northwest of Georgetown. All of these are listed by the
International Union for the Conservation of Nature as being of global
concern for long-term survival, mostly owing to human predation. The
categories of Critically Endangered or Endangered are applied to four of the
local sea turtles (80%). It is important to protect the few good nesting beaches
for the sea turtles of Guyana.
We have documented each of the species now known to comprise the
herpetofauna of Guyana by citing specimens that exist in scientific collections,
many of which were collected and identified by us and colleagues, including
students of the University of Guyana (UG). We also re-identified many old
museum specimens collected by others in the past (e.g., collections of William
Beebe) and we used documented publications and collection records of
colleagues, most of whom have been working more recently.
We present dichotomous keys for identifying representatives of the species
known to occur in Guyana, and we present brief annotated species accounts.
The accounts provide the current scientific name, original name (with citation
of the original description, which we personally examined in the literature),
some outdated names used in the recent past, type specimens, type localities,
general geographic distribution, examples of voucher specimens from
Guyana, coloration in life (and often a color photograph), and comments
pointing out interesting subjects for future research.
Keywords: amphibians, biogeography, Guyana, herpetology, keys, rep-

tiles, South America.
VOLUME 125, NUMBER 4 319
Introduction 1–40). A total of 78 of these species were

described based on samples collected at
Amphibians and reptiles are of interest type localities within Guyana and 39 were
to humans for various reasons, ranging described and named in the last 10 years.
from their usefulness for biomedical re- The lack of a modern comprehensive
search and development of pharmaceuti- volume dedicated to the herpetofauna of
cals to fascination with their diverse forms, Guyana has been misunderstood by some
colors, life histories and behavior, and as people to mean that these organisms are
models for scientific research to better unimportant and perhaps ‘‘nothing’’ was
understand the nature of planet Earth. known about the species native to the
Amphibians of one kind or another have country. Actually, the majority of the
existed for the last 365 million yr, having species have been known to science for
evolved from fish. Subsequently, the rep- decades or more (some going back to
tiles evolved from an amphibian, and the Linnaeus 1758), as they occur in adjacent
mammals (including humans) from a countries or broadly across Amazonia.
reptile. These animals are part of our Consequently, this majority of species has
own ancestry, as all living things on Earth been known to science, but most of what
are part of one family tree that extends we know about them is based on research
back for more than 3.5 billion yr. Am- that was done in countries other than
phibians were the first terrestrial verte- Guyana. In order to identify specimens
brates to appear in Earth history, and they found in Guyana, herpetologists have had
and reptiles are important components of to refer to diverse publications scattered
our terrestrial and aquatic ecosystems, widely in the technical literature, which
especially in the tropics. In contrast, can be a daunting task. Summary publi-
humans have existed for considerably less cations (with lists of possibly relevant
than the most recent 1 million yr and have species or comprehensive monographs)
had significant negative impact on plane- concerning the Guianan Region and South
tary ecosystems, particularly in just the last America that have been most broadly
200 yr. In terms of the natural resources of helpful to us include the following: Rivero
Guyana, the amphibians and reptiles (1961), Taylor (1968), Peters & Donoso-
(herpetofauna) are among the most poorly Barros (1970), Peters & Orejas-Miranda
known. (1970), Brazaitis (1973), Hoogmoed (1973,
The first checklist of the herpetofauna of 1979), Chippaux (1986), Ernst & Barbour
Guyana included a total of 164 species, (1989), Avila-Pires (1995, 2005), Barrio-
including 50 frogs and toads, 2 caecilians, Amorós (1998), Starace (1998), Gorzula &
1 amphisbaenian, 4 crocodilians, 33 liz- Señaris (1999), Lescure & Marty (2000),
ards, 63 snakes, and 11 turtles (Beebe Reynolds et al. (2002), Savage (2002),
1919). Today we are confident that twice Campbell & Lamar (2004), Duellman
as many species of amphibians and reptiles (2005), Señaris & MacCulloch (2005), and
occur in Guyana. The present volume Frost (2011). Some of these papers provide
documents 324 species (plus 20 thought skeleton lists of species of the herpetofau-
to occur in Guyana but not yet document- na of Guyana and adjacent areas, but none
ed plus an unknown number of new presents the perspective of Guyana on
species yet to be discovered), including details for confirmed voucher specimens
137 frogs and toads, 11 caecilians, 4 that were identified or re-identified recent-
crocodylians, 4 amphisbaenians, 56 liz- ly, extensive literature citations, identifica-
ards, 97 snakes, and 15 turtles (about 62% tion keys, illustrations of specimens, or
are shown in color photographs in Plates suggestions for future research.
The present volume scratches the sur- raphy, faunal comparisons, and conserva-
face, as considerable research remains to tion.
be accomplished with the wildlife of This volume should be viewed as a
Guyana, including the amphibians and progress report on the status of our
reptiles. Consequently, we are pleased to knowledge today. It seems fitting to quote
see the growing appreciation of Guyana’s Parker (1935:505), who listed 209 species
biodiversity and relatively untouched nat- of frogs, lizards, and snakes from Guyana:
ural environments, as shown in the recent ‘‘Many of the ‘species’ previously said to
outstanding book on the amphibians of occur in the Guianas have already been
Kaieteur National Park by Kok & Kala- relegated to synonymy, others have been
mandeen (2008) and the enthusiastic questioned, and others, again, appear to be
Guyanese students with whom we have based on misidentifications or on speci-
worked. mens with wrong locality data.’’ We have
In 1983 the National Museum of Nat- tried very hard not to perpetuate errors
ural History of the United States (Smith- made previously by others and not to
sonian Institution; NMNH) and the make new errors ourselves. There is still
University of Guyana (UG) initiated a much work to be done.
program now known as the Biological Scholars should consider this volume as
Diversity of the Guiana Shield Program an effort upon which the next generation
(BDG), in order to improve knowledge of should improve. We hope that students
the biota and train students in such and scholars will move quickly to make
this volume obsolete, and we hope the
pursuits. That program, with major fund-
citizens of Guyana will continue to con-
ing from the Smithsonian Institution,
serve their biological resources. It would
supplemented by the American Museum
be a travesty if this volume were to become
of Natural History (AMNH) and Royal
in the future only a record of what once
Ontario Museum (ROM), was the foun-
existed.
dation for developing this publication. In
addition, this volume benefited greatly
from the availability at the AMNH of
significant herpetological collections from Methods
Guyana made early in the 1900s on Each author conducted considerable
various separate trips made by Robert field work in Guyana in the last 20 years
Snedigar and William Beebe. or so, making significant new collections
Objectives.—This volume is intended to from localities that were not well sampled
provide the following: 1) a complete list of previously. In the field, reptiles were
the species of amphibians and reptiles euthanized with nembutal, amphibians
known to occur in Guyana through the with chloretone, usually after being pho-
year 2011 (Appendix 2); 2) keys and tographed in life. Color notes and often
illustrations useful for identifying individ- tissue samples (temporarily stored in liquid
uals of these species; 3) critical references nitrogen or 95% ethanol) were taken prior
to the primary literature; 4) citations of to preservation of the specimens in 10%
specimens in scientific collections that formalin. Upon return to the laboratory,
clearly document presence of the species specimens were soaked in old 70% ethanol,
in Guyana; 5) predictions about species of then after a few days transferred to fresh
possible occurrence but not yet document- 70% ethanol.
ed; 6) remarks about taxonomic and other At each locality visited in the field,
problems that need additional research; geographic coordinates and elevation were
and 7) preliminary comments on biogeog- determined with a GPS receiver (CJC and
CRT used a Sony Pyxis; RPR used a simply obtain a list of specimens from
Magellan and a Garmin; and RDM and collections and expect the identification of
AL used a Garmin); coordinates are specimens to be correct. In many instanc-
presented in World Geodetic System 1984 es the specimens, especially older ones,
(WGS 84). Many of our readings (partic- have not been identified or re-identified to
ularly in the early to mid-1990s) were modern standards, so names applied to
taken unavoidably on days when the specimens need confirmation or correc-
satellite signals were not of the highest tion. This requires examining the speci-
resolution so they include an unknown mens. A good example of what not to do
amount of error. To compensate for this, was unintentionally provided in an ap-
CJC and CRT took the mean of 10 pendix list of amphibians and reptiles
readings per site, and in the process, from Kaieteur National Park, which was
considerable variation was observed. The inappropriately attributed to CJC, CRT,
worst examples noted are as follows: and RPR (Kelloff 2003). Not only is that
readings for N latitude at Konawaruk list far from complete in terms of what
Camp (1998) varied over a span of 7.4 00 ; actually occurs at Kaieteur, but approx-
W longitude at a spring on Dubulay imately half of the names listed are
Ranch (1995) over a span of 4.5 00 ; and incomplete or simply wrong because old
elevation at Dubulay Ranch (1994) over a collection records were not verified prior
span of 51 ft to þ714 ft, where Alexander to publication and the sources cited did
Mendes told us the elevation was mapped not know that the list was being pub-
as about 100 ft. lished.
We identified all of the newly collected We also examined each of the recently
specimens personally and cataloged them published lists of the herpetofauna of
in the scientific collections at the AMNH, Guyana (Reynolds et al. 2002, Avila-Pires
CSBD at UG (specimens specifically col- 2005, Señaris & MacCulloch 2005), and
lected by us), ROM, and NMNH made sure we verified the occurrence of
(¼USNM catalog numbers; see below for each taxon, although none of the previous
a list of abbreviations for scientific collec- lists actually cited individual specimens.
tions). Literature that was most useful for Should users of this volume notice that a
identifying specimens is cited in the keys taxon listed previously for Guyana is not
and annotated species accounts. In addi- included here it is because we were unable
tion, we surveyed major collections that to verify the record, specimens were re-
were known to have specimens from identified to a different taxon, or taxo-
Guyana, and we personally examined nomic revisions resulted in recent name
specimens to confirm or correct identifica- changes. This applies, for example, to the
tions, where necessary. following: Allobates brunneus, Anomalo-
It is important to note, however, that glossus degranvillei, Arthrosaura versteegii,
we listed each of the species of amphib- Atractus badius, Atractus zidoki, Centro-
ians and reptiles that can be documented lene papillahallicum, Chironius cochranae,
with voucher specimens as occurring in Elachistocleis ovalis, Eleutherodactylus ur-
Guyana. We did not list every specimen in ichi, Helicops leopardinus, Hyalinoba-
every collection around the world that has trachium eccentricum, H. ignioculus,
specimens from Guyana. Researchers Mastigodryas bifossatus, Microcaecilia uni-
wanting complete listings of all specimens color, Oxyrhopus formosus, Oxyrhopus
for any taxon collected in Guyana will trigeminus, Phyllomedusa tomopterna, Pipa
need to do their own global surveys of the aspera, Pristimantis fenestratus, Rhinella
various collections. In this context, it is margaritifer, Rhinella typhonius, Thamno-
important to realize that one cannot dynastes strigilis, Trachycephalus venulo-
sus, Typhlops lumbricalis, and others. the caecilian Typhlonectes compressicauda

Comments on such details are presented at the UMMZ that was identified by Mark
in the species accounts, especially where we Wilkinson at the BMNH, or specimens of
explain such matters as re-identification of turtles identified by Peter C. H. Pritchard
specimens that were misidentified previ- at the Chelonian Research Institute. Sim-
ously. Re-identifications applied mostly, ilarly, for species recently described by
but not only, to specimens that were Philippe J. R. Kok at the IRSNB and
collected decades or nearly a century ago collaborators, it was not necessary to
and identified at that time on the basis of reexamine every specimen, although in
outdated literature. In addition, each of some instances we did. Suffice it to say
the bibliographic citations, including the that in the species accounts, we are quite
oldest, going back to Linnaeus (1758), has confident of the accuracy of the identifica-
been verified by at least one of us tions of the specimens listed in the
personally examining the actual publica- vouchers for Guyana. Nevertheless, addi-
tion cited. tional specimens may exist, and there is a
great deal of additional herpetological
work that needs to be done, including re-
Species Accounts identification of older specimens, before
the fauna of Guyana will be known
The brief species accounts are organized reasonably well.
with several topics as follows. Coloration in life.—This is based pri-
Type material.—This includes the orig- marily on our own field notes taken from
inal name used for the taxon, the unique animals while conducting field work in
specimen(s) to which the scientific name Guyana; the same applies to the photo-
is attached, and the type locality. Al- graphs in this volume. Other sources are
though some Amazonian species are used and cited when necessary, including
currently thought to have a very wide Beebe’s color notes on specimens from
distribution, especially the highly mobile Guyana. This category will be useful
species, the species of low vagility often together with the keys and photographs
become known as complexes of cryptic for identifying specimens.
species with several yet to be named, Comments.—This category is used to
following molecular analyses. Conse- explain recent name-changes, to describe
quently, for example, if a Guyanan interesting aspects of life history, and to
species has low vagility and a type indicate areas where additional research is
locality in Bolivia or Peru, this may needed.
suggest that the specimens from Guyana
are of a subtly different species, worthy of
modern investigation. Major Collecting Sites
Distribution.—This general statement of
the overall range of the species allows one For most of our field work, we targeted
to gauge aspects of biogeography and to localities where little or no previous
know whether a species is endemic to scientific collecting had been done. Our
Guyana or the Guianan Region. goal for each trip was to make a general
Vouchers for Guyana.—We personally collection documenting what species of
examined and confirmed the identification amphibians and reptiles occurred at the
of the specimens listed, with a few excep- locality visited. Also, strategies of the
tions where knowledgeable scientists had coauthors differed a bit. For example,
done so. For example, we did not need to CJC and CRT visited sites expected to be
reconfirm identifications of a specimen of best for studying the evolutionary biology
of unisexual species of lizards (thus staying Guyana, we distinguish among the sites
in the lowlands), RDM and AL chose sites by elevation, as follows: lowland sites are
expected to reveal previously unknown those below 500 m elevation; upland sites
species (thus mostly visiting the highlands), are between 500 and 1400 m elevation;
and RPR participated in multidisciplinary and isolated highland sites are above 1400
field trips of the BDG. Consequently, the m. Future workers will want to refine this
herpetofauna of sites from which we made system. For example, elevation alone and
adequate new collections could be com- air pressures associated with it are not as
pared with the herpetofauna of sites that important to most organisms as the more-
had been sampled well previously, and or-less elevation-correlated changes in
herpetofaunas from forest vs. savanna and temperature, rainfall, and soil moisture.
from lowlands vs. isolated highlands can Superimposed on this are additional
be compared to some extent, although effects, such as those based on the nature
considerable comparative work remains to of the local substrate, slope exposure
be done. We hope that scholars at UG will (including proximity of water fall spray
be inspired to improve this effort. and prevailing winds), and extent of
Where logistically feasible, we spent mountain mass. Local plant communities
about a month at each locality and often are more sensitive to these factors
collected both day and night. In addition, than animal communities, which are
where feasible, we dug in pit-fall traps of roughly correlated with the plants, so
20-liter plastic buckets with open tops over ideally, future workers may wish to
which passed vertical plastic sheeting compare herpetofaunal communities
stapled to cut saplings, which formed a more closely with specified plant commu-
drift fence that guided animals into the nities, and depending on all the physical
traps 24 hours per day (Fig. 1). To prevent characters mentioned, plant communities
desiccation of specimens, traps were kept may differ even at the same elevation on
damp and checked at least twice a day. different mountains or tepuis, or even on
This field method provides far better different slopes of the same mountain.
sampling of the herpetofauna than simply Nevertheless, the sites we compare for
walking and attempting to catch animals now are the 10 lowland and isolated
that happen to be seen, as in our experi- highland sites listed below.
ence, pit-falls produced specimens of spe- Lowland sites (below 500 m).—The
cies that we did not see otherwise. lowland sites are the following, going
However, highland sites did not lend approximately from north to south (Figs.
themselves logistically for long-term stays 3, 4). Details on each site follow.
nor for installing pit-fall traps. Informa- Baramita (7822 0 14 00 N, 60829 0 28 00 W; ca.
tion on effort is provided in the following 120 m elevation): RDM collected here,
lists of the sites within Guyana (Figs. 2–4), including use of drift fences and pitfall
and their herpetofaunas are compared traps, from 23 Sep–18 Oct 1992, and RPR
following the species accounts (see Bioge- collected here, without pitfall traps, from
ography). These abbreviated site names 10–18 Jun 1999. This was primarily a
are used to cite vouchers in the species rainforest site.
accounts. Kartabo (6821 0 N, 58841 0 W according to
For general comparisons, we refer to AMNH catalog; ca.100 m elevation [Ste-
lowland, upland, and isolated highland phens & Traylor 1985]): William Beebe
sites, similar to Gorzula & Señaris (1999), and colleagues from the New York Zoo-
because biotic communities vary along logical Society collected ‘‘in one-quarter of
elevational gradients. For convenience, a square mile of jungle’’ for parts of eight
suitable to the available sites within years from 1909–1926 (Beebe 1944:145).
Fig. 1. Trapline of drift fence (plastic sheeting stapled to cut saplings hammered into ground) and pitfall
trap (arrow; 20 liter plastic bucket buried with brim flush to ground). Plastic sheeting crosses the middle of the
open top of the bucket to allow catching things from either side.
Beebe also made collections in Trinidad tion, and this might explain problems
and Venezuela. Many of the specimens he associated with some of the specimens or
collected ultimately were received by the data given for them (Barbour 1920).
AMNH, but often the specimens and/or Identification of representatives of each
data arrived years or decades after collec- taxon was confirmed recently by CJC. This
Fig. 2. South America; rectangle covers Guyana and areas shown in more detail in Figs. 3 and 4.
is the largest fauna known from a single examination. Nevertheless, we examined

site in Guyana, to our knowledge. As the Iwokrama specimens that have USNM
reported by Donnelly et al. (2005), the catalog numbers. The Kartabo collections
fauna at the Iwokrama Forest Reserve were made without the benefit of drift
may be similar in diversity, and perhaps fences and pitfall traps, which might
more diverse, but those collections were explain the apparent absence of some
from a significantly larger area (including species, and if Beebe had a special interest
lowlands and uplands with different hab- in large reptiles, that might explain the
itats); many species were listed with apparent absence from his collections of
incomplete names, and representatives of several species of small frogs that must
some taxa were not available for our occur at Kartabo.
Fig. 3. Map of Guyana, color coded to show topography. The Pakaraima Mountains are west of
Kaieteur NP (¼National Park). We thank Ray Sterner, The Johns Hopkins University Applied Physics
Laboratory for the satellite image base map.
Fig. 4. Map of Guyana, showing the 10 places for which faunas are compared (see Biogeography).
Squares represent the 7 lowland sites, triangles the 3 isolated highland sites. See acknowledgement for Ray
Sterner in Fig. 3.
Dubulay Ranch (5840 0 55 00 N, 57851 0 when we stepped on them, and animal

32 00 W; ca. 30 m elevation): CJC and CRT activity in general was low.
collected here from 25 Feb–24 Mar 1994 Karanambo (3845 0 10 00 N, 59818 0 33 00 W;
and 26 Aug–24 Sep 1995, including the use ca. 115 m elevation): This is the McTurk
of drift fences and pitfall traps. RPR Ranch and vicinity, sometimes spelled
collected here without pitfall traps from Karanambu, on the northern part of the
9–12 Apr 1994. Most collecting was done Rupununi Savanna. CJC and CRT col-
within a 1 km (linear) radius of the ranch lected here and at the nearby village of
house, at ca. 60 m elevation according to Yupukari (ca. 11 km [linear] SSW Kar-
our GPS readings, but which Alexander anambo) from 15 Feb–13 Mar 1992, and
Mendes reports is probably closer to 30 m. RPR collected at Karanambo from 31
In addition, some specimens came from Mar–3 Apr 1994. In lieu of drift fences and
very small sites up to 12 km away. These pitfall traps, we had extensive assistance
sites include the following: Aramatani from local children at Yupukari. This is
Creek (5841 0 14 00 N, 57854 0 59 00 W), ca. 12 mostly savanna habitat, but there is gallery
km (linear) WNW the ranch house; a forest along the waterways, especially the
swamp along Aramatani Creek (5840 0 Rupununi River.
22 00 N, 57855 0 21 00 W); a spring (5839 0 53 00 N, Aishalton (2828 0 31 00 N, 59819 0 16 00 W; ca.
57854 0 31 00 W); Tableau Pond, ca. 6.5 km by 150 m elevation): This is a village on the
road NW the ranch house; and Warniabo southern part of the Rupununi Savanna.
Creek (5839 0 46 00 N, 57853 0 24 00 W), ca. 6.5 km CJC and CRT collected here from 18
by road SW the ranch house. This was an Mar–7 Apr 1993. In lieu of drift fences and
pitfall traps, we had extensive assistance
area of mixed habitats, including gallery
from local children. Most collecting was
rainforest along waterways, and patches of
done within 1 km (linear) of the village.
savanna with bush or forest islands.
This is mostly savanna, but there is gallery
Berbice River Camp (5805 0 06 00 N,
forest along the waterways.
58814 0 14 00 W; ca. 60 m elevation): CJC
Isolated highland sites (above 1400 m).—
and CRT collected here from 14 Feb–15 Locations of these sites are shown in Fig.
Mar 1997 with Gerald and Wesley King, 4. Vegetation descriptions are from
including the use of drift fences and pitfall Huber (1995) and Huber et al. (1995).
traps. Most collecting was done within a 1 Because of the shallow soil, rocky ter-
km (linear) radius of camp, which was on rain, and very dense vegetation, collec-
the east shore of the Berbice River. This tions were made without the use of
was primarily a rainforest site, but highly pitfall traps.
selective logging had been done in the past Mount Ayanganna (summit coordinates
in much of the area sampled. 5823 0 N, 59859 0 W; 2000 m): Mount Ayan-
Konawaruk Camp (5813 0 07 00 N, 59802 0 ganna is the highest peak located entirely
43 00 W; ca. 120 m elevation): CJC and CRT within Guyana. It marks the divide be-
collected here with Gerald and Wesley tween the headwaters of the Mazaruni,
King, including use of drift fences and Potaro, and Ireng Rivers. Most collections
pitfall traps, from 27 Feb–27 Mar 1998. were made on the northeast plateau of
Most collecting was done within a 1 km Ayanganna (5824 0 N, 59857 0 W; elevation
(linear) radius of camp. This was primarily 1490–1550 m), from 25 Oct–2 Nov 2000
a rainforest site, but our observations by RDM, AL, and Carter Cox, with some
suffer because it was unusually dry while help from guides and porters. Habitat on
we were here, during an extreme El Niño the northeast plateau is wet low evergreen
event. The dry leaves on the forest floor high-tepui forest, dominated by Bonnetia
crackled as we walked, sticks snapped roraimae, Schefflera, Clusia and Ilex spp.,
and large terrestrial bromeliads (Brocchi- them in the faunal comparisons (see
nia). More details are provided by Mac- Biogeography, below).
Culloch & Lathrop (2009). Mount Ayanganna Base (870 m;
Mount Wokomung (highest summit at 5825 0 N, 59858 0 W): RDM and AL collected
5805 0 N, 59850 0 W; 1700 m): Unlike Mount here on 3 Nov 2000.
Ayanganna, which has a single peak, Mount Wokomung Slope (1234 m;
Wokomung is a massif with several 5806 0 35 00 N, 59849 0 15 00 W): RDM and AL
connected peaks. Wokomung marks the collected here from 27 Oct–1 Nov 2004.
divide between the headwaters of the Mount Wokomung Base (698 m;
Potaro and Ireng Rivers. Collections 5807 0 46 00 N, 59848 0 37 00 W): RDM and AL
were made by RDM, AL, and Samir collected here from 21–26 Oct 2004.
Khan, with some help from guides and Kaieteur (above Kaieteur Falls; above
porters, on the northeast slope (5805 0 N, 450 m; 5810 0 N, 59829 0 W): RPR collected
59851 0 W; 1411 m), 2–8 Nov 2004. One here from 29 Mar–2 Apr 1989 and RDM
night, 6–7 November, was spent on the collected here from 1–20 Jun 2005. This
summit (5805 0 N, 59850 0 W; 1700 m). Slope area has been visited much more frequent-
habitat is montane forest with some ly by other researchers, however (Kok &
epiphytes and understory. The summit Kalamandeen 2008; their work is still in
is a shallow sloping bowl with very dense progress).
terrestrial bromeliads and woody shrubs
in the center, stunted trees around the
rim. Other collections have been made on
Abbreviations of Scientific
Mount Wokomung, notably by D. B.
Collections Used
Means (Means & Savage 2007).
Mount Roraima, summit, 2600–2810 m Abbreviations are used in the species
(58120 N, 60844 0 W): Roraima is the highest accounts to cite specimens in scientific
peak in the eastern Guiana Shield, and has collections that document occurrence in
been visited by numerous collectors. A Guyana. We follow Sabaj Pérez (2012) for
history of the exploration of Roraima is in collections listed, with few exceptions
McDiarmid & Donnelly (2005). Collections (specified).
were made on the slopes and summit, and
specimens are deposited in many institu- AMNH: American Museum of Natural
tions. Most collecting was done on the History, New York, New York.
Venezuelan portion of the mountain, where ANSP: Academy of Natural Sciences of
access is relatively easy. Two collections Drexel University, Philadelphia,
were made on the Guyanese portion of Pennsylvania.
Roraima: one is at the BMNH (Warren BMNH (¼BM or NHMUK in some
1973) and the other at the NMNH. For the papers): Natural History Museum,
present report we include all species col- London (formerly, British Museum
lected above 1400 m on Mount Roraima, of Natural History).
assuming that they occur in both Venezuela CPI: ‘‘D. Bruce Means field series, to be
and Guyana, including those collected by deposited at USNM’’ (Grant et al.
other investigators. The herpetofauna of 2006:45).
Roraima is summarized by MacCulloch et CSBD: see UG.
al. (2007). EBD: Estación Biológica de Doñana,
Upland sites (between 500–1400 m).— Seville, Spain.
Locations of these sites are shown on Figs. EBRG: Estación Biologia de Rancho
3 and 4, although none of us spent Grande, Museo, Maracay, Vene-
sufficient time at these sites to include zuela.
FMNH: Field Museum of Natural ULAM (¼ULABG in some papers):

History, Chicago, Illinois. Herpetological Collection, Labora-
IBSP: Instituto Butantan, São Paulo, tory of Biogeography, Universidad
Brazil. de Los Andes, Mérida, Venezuela.
IRSNB: Institut Royal des Sciences UMMZ: University of Michigan Muse-
Naturelles de Belgique, Brussels. um of Zoology, Ann Arbor, Michi-
KU: University of Kansas Biodiversity gan.
Institute, Lawrence, Kansas. UPRM (¼UPR-M): University of Puer-
MBUCV: Museo de Biologı́a, Universi- to Rico, Mayagüez.
dad Central de Venezuela, Caracas. USNM: National Museum of Natural
MCZ: Museum of Comparative Zoolo- History, Smithsonian Institution,
gy, Harvard University, Cambridge, Washington, D.C.
Massachusetts. UTA: University of Texas at Arlington,
MHNLS: Museo de Historia Natural Department of Biology.
La Salle, Caracas, Venezuela. UUZM (¼ZIUU or ZMUU in some
MNHN (¼MNHNP in some papers): papers): Uppsala University, Zoolog-
Muséum national d’Histoire Natu- ical Museum (¼Zoologiska Museet),
relle, Paris. Sweden.
MNRJ: Museu Nacional, Universidade ZFMK: Zoologisches Forschungsmuse-
Federal do Rio de Janeiro, Brazil. um Alexander Koenig, Bonn, Ger-
MSNM: Museo Civico di Storia Natu- many.
rale di Milano, Italy. ZISP (¼ZIN): Zoological Institute, Rus-
MZUSP: Museu de Zoologia, Universi- sian Academy of Sciences, St. Peters-
dade de São Paulo, Brazil. burg.
NHMW (¼NMW in some papers): ZIUU: see UUZM.
Naturhistorisches Museum, Wien ZMB: Museum für Naturkunde (for-
(Vienna), Austria. merly Zoologische Museum), Leibniz-
NMNH: National Museum of Natural Institut für Evolutions- und Biodiver-
History, Smithsonian Institution, sitätsforschung an der Humboldt-
Washington, D.C. (USNM for speci- Universität, Berlin, Germany.
men catalog numbers). ZMUU: see UUZM.
NRM (¼NHRM in some papers): Na- ZSM (¼ZSMH in some papers): Zoo-
turhistoriska Riksmuseet, Stockholm, logische Staatssammlung, Munich,
Sweden. Germany.
PCHP: Peter C. H. Pritchard, Chelonian ZUU: Museo Academico Upsalensi,
Research Institute, Oviedo, Florida. Sweden.
RMNH: Netherlands Centre for Biodi-
versity Naturalis, Leiden.
ROM: Royal Ontario Museum, Toron-
to, Canada. Identification of Specimens
SMNS: Staatliches Museum für Natur-
kunde, Stuttgart, Germany. For a variety of reasons, it is often
UF: University of Florida, Florida desirable to identify animals that are under
Museum of Natural History, Gaines- observation, whether in the laboratory or
ville. the field. The use of a dichotomous
UG (¼CSBD or HA in some papers): identification key (or several keys in
Centre for the Study of Biological succession) is an efficient way to accom-
Diversity, University of Guyana, plish this, although good magnification
Georgetown. may be necessary to see some of the
characters, ideally with a dissecting micro- birds (although birds are now classified as
scope. A dichotomous key presents the one of the major groups of reptiles), or
user with a series of paired choices mammals (which have a constant, warm
(couplets), beginning with pair number body temperature, hair, and, in South
one (1) and continuing through a series America, give birth to living young). Once
of numbered choices. At the end of each one has decided one has a vertebrate that is
paired set of statements, the user is led to not one of these, it must be either an
either the next relevant couplet or the amphibian or reptile.
probable name of the taxon identified. At Reptiles have epidermal scales that are
this point, it will be helpful to go to the perfectly conspicuous and readily felt on
relevant species account; each account has the surface of their skin; amphibians do
color notes and each account begins with a not (the hidden dermal scales of caecilians
reference to the plate with the color are very inconspicuous). For those reptiles
photograph, if present. Once this process that have legs, all of the fingers and toes
is completed, the identification obtained almost invariably have claws; amphibian
may or may not be correct, or there may fingers and toes do not. If the animal looks
be questions about why certain characters like a frog or toad, it is an amphibian, even
seen on the specimen do not fit some that if it has a vocalization that sounds like the
were expected in using the keys. quack of a duck. However, if it looks very
When possible, follow-up should be superficially like an earthworm, check it
done by comparing the specimen identified carefully, as there is a possibility that it is
with either photographs or other speci- an amphibian or scaly reptile. Caecilians
mens previously identified as belonging to (Gymnophiona) are wormlike amphibians
the same species. Throughout this process, (see Microcaecilia, Plate 18F) that are
there are several possible sources of error, legless, with a moist body that is not as
such as the following: 1) a mistake in limp in hand as an earthworm. Amphis-
observations or wrong turn in the key; 2) baenians (worm lizards) are legless reptiles
the specimen represents a taxon currently that have epidermal scales and eyelids.
not known to occur in Guyana and Look and feel carefully to see if a wormlike
therefore not included in the key; 3) the animal has the following characters: a
specimen represents a taxon unknown to bone-hard head at one end, which is
science; 4) the specimens used for compar- clearly different from the other end on
ison may have been misidentified previ- close inspection; possibly vestigial eyes (if
ously; 5) the specimen is abnormal in key not absent or concealed by bone), a small
characters; or 6) the key is faulty and does mouth, with tiny teeth, and a vertebral
not work properly in some respect. In column. Some species of caecilians are 30
some cases, it may be useful to compare cm or more in length and not readily
the specimen with the color plates, find a confused with earthworms, but, as if to
species that might be it, then find that make matters worse, one species (see
species in the key and work backwards Typhlonectes compressicauda) is fully
through the key to see where the difficulty aquatic in fresh water and could be
occurs. Frustrating as it may be, it can be confused with an eel.
an interesting challenge to identify a Our focus here is to understand the
difficult specimen when it does not key species that occur in Guyana, without
out directly and easily. emphasis on subspecies. In cases where
Initially, one must choose the right key there are significant issues with possible
with which to begin. In the context of this subspecies, these are discussed in the
volume, our focus is on vertebrates (or- species accounts. Also, our focus is on
ganisms with a backbone) that are not fish, adults and juveniles, excluding identifica-
tion of larval forms (e.g., tadpoles). We 1a. Body very flat; eyelids and tongue
hope that soon someone will follow up absent; fingertips star-shaped;
with a volume dealing with the larval aquatic, even as adults . . . . . . . . . . . . . 2
forms of all the amphibians of Guyana, 1b. Not as 1a . . . . . . . . . . . . . . . . . . . . . . . . . 3
as these are important components of the 2a (1). Irregular shaped dermal flap at
aquatic ecosystems, yet it will require angle of jaw; toes (hind feet)
considerable work to complete such a without keratinous tips. . . . . . . . . . .
volume. Similarly, a volume on the vocal- . . . . . . . . . . . . . . . . . . . . Pipa pipa, p. 435
izations of the frogs, especially their
2b. No dermal flaps at angle of jaw;
advertisement calls, would be very useful
tips of toes I–III with keratinous
to help in the identification of individuals, caps in adults . . . . . . . . . . . . . . . . . . .
supported by recordings and print-outs of
. . . . . . . . . . . . . . . . Pipa arrabali, p. 435
their acoustical characters (e.g., Kok &
3a (1). Toes (hind feet) opposable;
Kalamandeen 2008).
relative toe lengths, adpressed,
IV.V. I.IIIII . . . . . . . . . . . . . . . . . . 4
Key to the Orders of Amphibia 3b. Toes not opposable; relative toe
of Guyana lengths, adpressed, IV.VIII.II
.I or IV.III.V.II.I . . . . . . . . . . . . 5
1a. Legless amphibians . . . . . . . . . . . . . . 4a (3). Larger frogs with green backs,
. . . . . Gymnophiona (caecilians), p. 348 not with warty, toad-like appear-
1b. Four-legged amphibians . . . . . . . . . ance; eardrum (tympanum) distinct
. . . . . Anura (frogs and toads), p. 332 on side of head; pupil of eye
vertical; toes IV and V not fused
Key to the Families and Many Species of at base (Fig. 5B) . . . . . . . . . . . . see sepa-
Anura (Frogs and Toads) of Guyana rate key for Phyllomedusa, p. 345
4b. Small, warty, toad-like appear-
Many of the families and genera of ance; tympanum not visible; pupil
anurans that occur in Guyana are repre- horizontal or round; toes IV and V
sented by only one or a few species that fused at base (Fig. 5A) . . . . . see sepa-
occur in the country (e.g., Atelopus, Pipa). rate key for Bufonidae, p. 338
For those groups, the Guyanese species 5a (3). Discs of fingertips with two
can be identified with this key. For most scale-like scutes on upper surface
families and genera having more represen- (Fig. 5D) . . . . . . . . . . . . . . . . . . . . . . .
tatives in Guyana (e.g., Bufonidae, Hyli- . . . . . . . . see separate key for Aromo-
dae), this key will lead users to relevant batidae and Dendrobatidae, p. 336
specific keys that follow. 5b. Not as 5a . . . . . . . . . . . . . . . . . . . . . . . . . 6
This key is based largely on that of 6a (5). Conspicuous toad-like paro-
Lescure & Marty (2000). In addition, toid glands present (but may be
Dunn (1949), Trueb & Cannatella (1986), small or large) on neck behind ears
and Campbell & Clarke (1998) were (Fig. 5E) and no black spot edged
especially helpful in places. Fingers and with red in groin . . . . . . . . . . . . . . . . . . . 7
toes are numbered by counting from the 6b. No parotoid glands, or, if present,
innermost (thumb or big toe) as I to the groin with large black spot edged
outermost (pinky) as IV or V. Measure- with red . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
ment of the body length is often from the 7a (6). Tympanum (eardrum) not ev-
tip of the snout to the vent, but some ident (Fig. 5E); first toe somewhat
investigators measure to the end of the indistinct, enclosed in skin . . . . . . .
bone felt above the vent (urostyle). . . . . . . . . . . . Atelopus spumarius, p. 375
Fig. 5. A, Underside of foot, Oreophrynella dendronastes, toes IVþV fused at base (Lathrop &
MacCulloch 2007:90). B, Underside of foot, Scinax boesemani, toes webbed, not fused. C, Adelophryne
patamona hand, inconspicuous points on fingertips, pads beneath toes, not tubercles (MacCulloch et al.
2008a:40). D, Upper view, Dendrobates leucomelas toe, paired scale-like scutes shaded. E, Parotoid glands
(shaded) behind tympanum in Rhaebo guttatus (left), neither conspicuous in Atelopus spumarius (right). F,
Hand of Hypsiboas (left), fingertip discs; Leptodactylus (right) without discs. G, Hands of Pristimantis (left),
Scinax (right), fingertip discs. Scale bars: A–D, F, G ¼ 1 mm, E ¼ 1 cm.
7b. Tympanum clearly visible; first toe cles connected by irregular, fine,
distinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 pale line . . . . . . . . . . . . . . . . . . . . . . . . . . 14
8a (6). Body somewhat flattened 14a (13). Upper surface of thigh and
(some species less so than others), rear of thigh mottled with dark
with narrow head (although broad brown spots, with narrow dark
at neck); nose pointed (viewed brown line below; tip of snout
from above); pupil round (some rounded . . . . . Otophryne robusta, p. 433
Bufonidae; all Microhylidae) . . . . . . 10 14b. Upper surface and rear of thigh
8b. Not as 8a . . . . . . . . . . . . . . . . . . . . . . . . 21 not mottled; rear of thigh with row
9a (7). Body length (snout-vent) usu- of large, white tubercles, mostly
ally more than 25 mm; crests on connected by irregular, fine, pale
head; body thick or squat; warty line; pale color above this line,
skin on back . . . . . . . . . . . . see separate broad dark brown line below; tip
key for Bufonidae, p. 338 of snout protrudes sharply . . . . . . .
9b. Body length less than 25 mm; no . . . . . . . . . . . Otophryne pyburni, p. 433
cranial crests; body more slender; 15a (12). Orange spot in groin . . . . . . .
granular skin on back. . . . . . . . . . . . . . . . . . Elachistocleis surinamensis, p. 431
. . . . Dendrophryniscus minutus, p. 375 15b. No orange spot in groin . . . . . . . . . 16
10a (8). Toes (on feet) essentially 16a (15). Snout projecting well past
completely webbed . . . . . . . . . . . . . . . . 11 lower jaw (viewed from side) and
curving downward; nostril beyond
10b. Toes with incomplete or no web-
tip of lower jaw; snout blunt,
bing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
distinctly light in color . . . . . . . . . . . . 17
11a (10). Belly white with large, irreg-
16b. Not as 16a, snout projecting
ular black spots; irregular black
much less . . . . . . . . . . . . . . . . . . . . . . . . . 18
spot in groin . . . . . . . . . . . . . . . . . . . . .
17a (16). Back, sides, and limbs brown
. . . . Chiasmocleis shudikarensis, p. 430
(gray-brown in life) with many light
11b. Belly dark brown with white spots (cream to orange in life); belly
spots; groin basically all dark uniform grayish white in life; dis-
brown to black, not just a spot. . . tinct light stripe from snout over
. . . . . . . . . . . Ctenophryne geayi, p. 431 eye; ear (tympanum) concealed;
12a (10). Tympanum as big as eye; body length of adult males less than
behind eye, light-colored dorsolat- 30 mm. . . Synapturanus salseri, p. 434
eral ridge or fold on skin along 17b. Back brown (reddish brown in
body. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 life) with no discrete spots (or only
12b. Tympanum smaller than eye or tiny flecks of white); brown mottled
concealed; no dorsolateral ridge or pattern on sides and limbs; belly
fold. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 yellow or tan in life; no light stripe
13a (12). Dorsal body coloration (on from snout over eye; ear sometimes
back) black or dark gray, with visible; body length of largest adult
yellow spots or blotches (usually males greater than 30 mm. . . . . . . . .
red in Venezuela); rear of thigh . . Synapturanus mirandaribeiroi, p. 434
with thin but distinct yellow line 18a (16). Back light tan or brown
. . . . . . Otophryne steyermarki, p. 434 with large, irregular, hourglass
13b. Dorsal coloration lighter, reddish shaped darker brown area (which
brown to grayish yellow, with may fade in preservative); viewed
scattered pale tubercles; rear of from above, a dark brown spot at
thigh mottled or with pale tuber- rear on back, in front of hip
(each side) . . . . . . . . . . . . . . . . . . . . . 25b. No dorsolateral fold or ridge . . . . 26

. . . . Hamptophryne boliviana, p. 432 26a (25). Adult body length (snout-
18b. Not as 18a, although possibly vent) usually 50–65 mm; no cream
brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 stripe on side of body; rear of
19a (18). No fingers or toes with thighs with creamy yellow stripe;
swollen tips or discs (Fig. 5F); chest and belly with dark gray
irregular black spot in groin. . . . . . flecks . . . . . . Pseudis paradoxa, p. 414
. . . . Chiasmocleis shudikarensis, p. 430 26b. Adult body length less than 30
19b. Some fingers and toes with swol- mm; cream stripe on side of body;
len tips; no black spot in groin . . . . 20 rear of thighs with cream dots;
20a (19). First (inner) finger and first chest and belly with cream dots . .
toe rudimentary; discs at tips of . . . . . . . . . . . . . . . Lysapsus laevis, p. 408
swollen fingers and toes with 27a (24). Conspicuous, flexible, der-
groove at tip, between upper and mal horn atop upper eyelid . . . . . .
lower surfaces; belly white, with . . . . . . . . . . Ceratophrys cornuta, p. 385
brown stippling on throat, brown 27b. Not as 27a . . . . . . . see separate key
reticulum on limbs . . . . . . . . . . . . . . . for Leptodactylidae, p. 346
. . . . . . . . . Chiasmocleis hudsoni, p. 430 28a (23). Snout pointed (viewed from
20b. First finger not rudimentary, al- above), mouth not at tip; in life,
though first toe reduced; swollen body green above, with cream
tips of fingers and toes without stripe (bordered below by brown
groove; belly cream, heavily spot- stripe) from snout to eye; small
ted with gray . . . . . . . . . . . . . . . . . . . . dermal crest above vent . . . . . . . . . .
. . . . . . . . . . . . . Chiasmocleis jimi, p. 430 . . . . . . Sphaenorhynchus lacteus, p. 418
21a (8). In adults, belly skin translu- 28b. Not as 28a; if body green above,
cent to transparent, some internal snout is rounded or truncate . . . . . . 29
organs clearly visible . . . . . . . . . . . . . . . . 29a (28). Finger I (innermost or
see separate key for Allophrynidae thumb) shorter than finger II. . . . .
and Centrolenidae, p. 335 see separate key for Hylidae, p. 341
21b. Not as 21a . . . . . . . . . . . . . . . . . . . . . . 22 29b. Finger I longer than or equal in
22a (21). Tips of fingers and toes with length to finger II . . . . . . . . . . . . . . . .
conspicuous disc (Fig. 5F, G) . . . . . 23
see separate key for Hemiphractidae
22b. No discs at tips of fingers and
(Stefania), p. 340
toes (or only slightly expanded tips
on animals with adult body length
less than 20 mm). . . . . . . . . . . . . . . . . . 24 Key to Frogs of the Families
23a (22). Toe webbing absent or rudi- Allophrynidae and Centrolenidae
mentary; finger discs wider than of Guyana
long . . . . . . . . . . . see separate key for
Ceuthomantidae, Craugastoridae, The following key is based primarily on
Eleutherodactylidae, p. 339 information provided by Kok & Castro-
23b. Toes webbed at least half way viejo-Fisher (2008). Additional informa-
toward tips; toe discs rounded . . . . 28 tion was obtained from Gorzula & Señaris
24a (22). Toes webbed entirely to tips 25 (1999) and Lescure & Marty (2000), and
24b. Toes not completely webbed . . . . . 27 we benefitted from editing and corrections
25a (24). Body with dorsolateral fold provided by Santiago Castroviejo-Fisher
or ridge of skin from behind eye (pers. comm.).
toward rear . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . Lithobates palmipes, p. 436 1a. Back brown with black spots;
viewed from above, head triangu- servative) and small brown dots . .
lar, with narrow snout; back with . . Hyalinobatrachium cappellei, p. 382
hemispherical pustules having a 6b. Snout rounded; back green with
central spinule; cream markings small light yellow spots . . . . . . . . . .
on lips, sides, arms, and legs . . . . . . . Hyalinobatrachium mondolfii, p. 383
. . . . . . . . . . . Allophryne ruthveni, p. 370
1b. Back primarily green; viewed from
above, snout wider, more rounded;
Key to Frogs of the Families
otherwise also not as 1a. . . . . . . . . . . . 2
Aromobatidae and Dendrobatidae
2a (1). Looking through translucent
of Guyana
belly, anterior (front) one-third of
internal parietal peritoneum (ab-
dominal sheath) white; back light The monophyly and recognition of
lime green with a few small black families and genera of these small frogs
or dark brown flecks; bones green are strongly supported by molecular
in life . . . . . Vitreorana helenae, p. 384 (DNA sequence) data, but in many cases
2b. Parietal peritoneum transparent, there are no, or only inconspicuous,
viewed externally in life, although morphological characters to identify them
some visceral peritonea white; color- (Grant et al. 2006). The following key is
ation basically green on back but not based on information provided by
same as 2a . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Rodrı́guez & Duellman (1994), Grant et
3a (2). Bony spine or protuberance at al. (1997, 2006), Gorzula & Señaris (1999),
base of thumb (Fig. 6A) and on Lescure & Marty (2000), Jungfer & Böhme
upper arm near shoulder in males (2004), Duellman (2005), Kok et al.
(Fig. 6B); bones green in life; body (2006a, 2006b), and Kok et al. (2007).
all dark green above (with numer- 1a. With fingers adpressed, finger I
ous subtle pale dots); heart covered (innermost or thumb) clearly longer
with white pigment . . . . . . . . . . . . . . than finger II . . . . . . . . . . . . . . . . . . . . . . . 2
. . . . . . . . . . Vitreorana gorzulae, p. 384
1b. With fingers adpressed, finger I
3b. Thumb spine concealed and oth-
usually shorter than or equal in
erwise not as 3a. . . . . . . . . . . . . . . . . . . . 4
length to finger II (although may
4a (3). Snout sloping in profile; bones
appear longer without adpressing). . 4
green in life; back green with small
or tiny pale whitish spots . . . . . . . . 2a (1). Upper surfaces of arms and
. . . . Hyalinobatrachium taylori, p. 383 legs green; belly black with blue
4b. Snout truncate or round in profile; spots; broad yellow or greenish
bones white in life; back green but yellow dorsolateral stripe from
otherwise not as in 4a. . . . . . . . . . . . . . 5 snout to groin; adult body (snout-
5a (4). Green back with round yellow vent) length more than 35 mm . . .
spots (cream in preservative) . . . . . . . 6 . . . . . . . . . . . Ameerega trivittata, p. 391
5b. Green back with irregular-shaped 2b. Not as 2a . . . . . . . . . . . . . . . . . . . . . . . . . 3
lime green areas (white in preser- 3a (2). Brown mottling on chest or
vative) and scattered black dots . . belly; adult males with throat dark
. . . . . . Hyalinobatrachium iaspidiense, gray, brown, or black; adult body
p. 382 length 20–34 mm . . . . . . . . . . . . . . . .
6a (4). Snout truncate, viewed from . . . . . . . . . . . Allobates femoralis, p. 370
above or side; body green above 3b. No mottling on chest or belly; adult
with somewhat subtle but large males and females with very pale
round yellow spots (cream in pre- throat; adult body length less than
Fig. 6. A, Vitreorana gorzulae with thumb protuberance. B, Vitreorana with humeral spine. C, Hypsiboas
cinerascens with prepollical spine. D, Calcars on tree frog heels; 1, elongate, pointed on Hypsiboas calcaratus;
2, absent on Dendropsophus marmoratus; 3, small, tubercular on Myersiohyla kanaima; 4, small, conical on
Hypsiboas roraima. E, Caecilians, Siphonops, tentacle near eye (upper), Caecilia, tentacle below nostril
(lower); see Acknowledgments. F, Primary (P) and secondary (S) grooves of Caecilia, marking folds or annuli;
see Acknowledgments. Scale bars: A, C ¼ 1 mm, B, E ¼ 2 mm, D, F ¼ 5 mm.
20 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . 8b. Not as 8a; adults larger; toes not

. . . . . . . . Allobates spumaponens, p. 370 webbed; tongue lacks median lin-
4a (1). With fingers adpressed, finger I gual process . . . . . . . . . . . . . . . . . . . . . . . 9
equal in length to finger II . . . . . . . . . 5 9a (8). Back with scattered, small,
4b. With fingers adpressed, finger I roundish light spots; body length
usually shorter than finger II . . . . . . . 7 of adult males less than 30 mm . .
5a (4). Tongue with pimple-like medi- . . . . . Dendrobates nubeculosus, p. 392
an lingual process; belly cream, 9b. Dorsum not as in 9a; body length
yellow, or orange (although ante- of adult males 30 mm or more . . . . 10
rior belly may be gray with black 10a (9). Viewed from above, conspic-
blotches); toes (on feet) with only uous, broad, black band around
basal to moderate webbing . . . . . . . . . 6 neck region; toward rear, conspic-
5b. Tongue without median lingual uous, broad, yellow bands with
process; belly black or dark brown, black intrusions . . . . . . . . . . . . . . . . . .
marbled with light blue; no webbing . . . . . . Dendrobates leucomelas, p. 391
on toes . . . . . Ameerega hahneli, p. 390 10b. Not as 10a; often with blue
6a (5). Adult body length usually less markings also . . . . . . . . . . . . . . . . . . . .
than 20 mm or just a bit more; . . . . . . . . Dendrobates tinctorius, p. 392
fingers II and III without fringing 11a (7). Adults of both sexes with
fold on side; dorsolateral stripe body length of 21 mm or more
begins at posterior corner of eye; (females to 28 mm) . . . . . . . . . . . . . .
no black blotches on anterior part . . . . . . Anomaloglossus megacephalus,
of belly; toes moderately webbed p. 372
. . . . . . . . Anomaloglossus kaiei, p. 372 11b. Adult body length less than 20
6b. Adult body length usually 20 mm mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
or more; fingers II and III with 12a (11). Back bright yellow or orange
keel-like fringing fold on side; to pale tan; webbing between all
dorsolateral light stripe begins at toes (on feet); light dorsolateral
tip of snout; anterior part of belly stripe present, although may be
of males with black blotches; toes obscure; throat and chest yellow . .
webbed only at base . . . . . . . . . . . . . . . . . . . . . Anomaloglossus beebei, p. 371
. . . . . Anomaloglossus praderioi, p. 373 12b. Back gray; no webbing between
7a (4). Coloration on back (dorsum) toes I–III; webbing nearly absent
includes bright colors (gold, yel-
between toes III–V; no light dor-
low, orange, bold white, blue,
solateral stripe; undersides gray or
shiny black); tongue usually with-
brown, not yellow . . . . . . . . . . . . . . .
out pimple-like median lingual
. . . . . . Anomaloglossus roraima, p. 374
process (except in A. beebei). . . . . . . . 8
7b. Dorsal coloration cryptic, dull
brown or gray (sometimes black); Key to Frogs and Toads of the Family
tongue with pimple-like median Bufonidae of Guyana
lingual process . . . . . . . . . . . . . . . . . . . . 11
8a (7). Yellow or orange frog; small This key includes bufonid genera and
(adult body length 19 mm or less); species that were not keyed out in the
toes with webbing; tongue with initial anuran key. The initial key covers
pimple-like median lingual process; Atelopus spumarius and Dendrophryniscus
lives in or near large terrestrial minutus. The following key is based on
bromeliads. . . . . . . . . . . . . . . . . . . . . . . information provided by Hoogmoed
. . . . . . . Anomaloglossus beebei, p. 371 (1977), Lescure & Marty (2000), Frost et
al. (2006), Fouquet et al. (2007a), and brown to black, in some edged
Lathrop & MacCulloch (2007). above by narrow cream stripe or
row of cream warts . . . . . . . . . . . . . .
1a. Conspicuous parotoid glands on . . . . . . . . . . . . . . Rhaebo guttatus, p. 378
neck; toes (on feet) not opposable;
6b. Bony crests present on head, small
usually found in lowland or upland
in some species, very conspicuous
habitats. . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
in others; sides not as 6a . . . . . . . . . . . 7
1b. No conspicuous parotoid glands;
toes opposable; highland species, 7a (6). Protruding bony knob at an-
found above 1000 m elev. . . . . . . . . . . 2 gle of jaws (rear of mouth, extend-
2a (1). Extensive webbing on fingers ing to side); bony crests on head
and toes; distinct bony crest behind most conspicuous . . . . . . . . . . . . . . .
eye . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 . . . . . . . . . . . . . . Rhinella martyi, p. 379
2b. Webbing on fingers and toes only 7b. No protruding bony knob at angle
basal or moderate; bony crest of jaws; crests present on head but
behind eye indistinct or apparently not as prominent . . . . . . . . . . . . . . . . . . 8
absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 8a (7). Edge of lower jaw with white
3a (2). Prominent, short bony crest stripe or row of white spots; paro-
below eye; back blackish brown toid glands on neck moderately
without lengthwise line along mid- large, longitudinally oval, about
dle (vertebral area); belly dark twice as long as wide; upper eyelid
brownish orange with lengthwise spiny, projecting over eye with
black line along middle (midven- distinct vertical surface on side;
tral); palms of hands and soles of sharply pointed snout . . . . . . . . . . . . .
feet orange. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhaebo nasicus, p. 378
. . . . . . Oreophrynella seegobini, p. 377 8b. Not as 8a; parotoid glands large,
3b. Crest below eye indistinct; back triangular in shape . . . . . . . . . . . . . . . . . 9
reddish brown with dark markings 9a (8). Adults are largest toads in the
and a thin, dark brown vertebral world; warts on back without
line; belly light reddish brown spinules; toes up to half webbed;
without black midventral line; belly light cream with gray or
palms and soles light brown. . . . . . brown spots or mottling . . . . . . . . .
. . Oreophrynella weiassipuensis, p. 378 . . . . . . . . . . . . . . Rhinella marina, p. 379
4a (2). Back covered with numerous, 9b. Maximum adult body length, 45
distinct, conical tubercles; fingers mm; some warts on body with
with only moderate webbing . . . . . spinules; toes up to one-third
. . . . . . . . Oreophrynella quelchii, p. 377 webbed; belly grayish . . . . . . . . . . . .
4b. Body with relatively few tubercles; . . . . . . . . . . . . Rhinella merianae, p. 381
only basal webbing on fingers . . . . . . 5
5a (4). Snout somewhat pointed in
profile, projecting beyond front Key to Frogs of the Families
end of lower jaw . . . . . . . . . . . . . . . . . Ceuthomantidae, Craugastoridae, and
. . . . Oreophrynella macconnelli, p. 376 Eleutherodactylidae of Guyana
5b. Snout truncate, not projecting
beyond lower jaw . . . . . . . . . . . . . . . . The following key is based on informa-
. . . Oreophrynella dendronastes, p. 376 tion provided primarily by Lescure &
6a (1). No conspicuous bony crests on Marty (2000), Means & Savage (2007),
head; sides (eye to groin) with MacCulloch et al. (2008a), and Heinicke et
broad, bold band of very dark al. (2009). Helpful information was also
obtained from Rivero (1968a) and Duell- becoming dull white to brown in
man (1997, 2005). preservative) . . . . . . . . . . . . . . . . . . . . . . . . 7
6b. Not as 6a, although a tannish
1a. Fingers and toes end in a point orange vertebral stripe may be
(Fig. 5C); subdigital pads (beneath present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
toes) rather than tubercles; tiny 7a (6). Tympanum (eardrum) distinct;
frogs, adult snout-vent (body) no bars on upper lip area but red
length less than 25 mm. . . . . . . . . . . . . 2 spots may be present; back purplish-
1b. Fingers and toes end in discs (Fig. black with several large red spots;
5G); subdigital tubercles rather undersides bright red, without black
than pads; usually larger frogs, . . . . . Pristimantis dendrobatoides, p. 387
adult body length greater than 25 7b. Tympanum not visible; bars on
mm (but a few species are smaller) . . 3 upper lip areas; back with brilliant
2a (1). Maximum body length of 16 green and red crossbands, possibly
mm; side of face between eye and with an orange vertebral stripe;
nostril (canthus rostralis) straight undersides mottled cream and
or flat; second segment of finger IV black, possibly with cream stripe . .
short, equal to width. . . . . . . . . . . . . . . . . . . . . . . . . . Pristimantis jester, p. 388
. . . . . . . . Adelophryne gutturosa, p. 393 8a (6). Yellow or orange spot in groin
2b. Maximum body length more than . . . . . . . . Pristimantis inguinalis, p. 387
20 mm; canthus rostralis concave; 8b. Not as 8a . . . . . . . . . . . . . . . . . . . . . . . . . 9
second segment of finger IV longer, 9a (8). Toes (on feet) with rudimenta-
about twice width. . . . . . . . . . . . . . . . . ry, basal webbing . . . . . . . . . . . . . . . .
. . . . . . . . Adelophryne patamona, p. 393 . . . . . . Pristimantis marmoratus, p. 388
3a (1). Tiny frogs, adult body length 9b. No webbing on toes . . . . . . . . . . . . . . 10
usually less than 20 mm; paired 10a (9). Tympanum visible but indis-
gland-like protrusions on post- tinct . . . . . . . . . . . . . . . . . . . . . . . . . . . .
temporal and hip regions; viewed . . . . . . Pristimantis saltissimus, p. 389
from above, digital discs notched 10b. Tympanum prominent or clearly
at tip; prominent subconical tuber- visible . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
cle on upper eyelid and heel; row of 11a (10). Large, transversely oval fin-
conical tubercles on outer edge of ger discs; disc on finger III larger
tarsus; bright green markings on than tympanum. . . . . . . . . . . . . . . . . .
head and body. . . . . . . . . . . . . . . . . . . . . . . . . . Pristimantis pulvinatus, p. 389
. . . . Ceuthomantis smaragdinus, p. 385 11b. Smaller, rounded finger discs;
3b. Not as 3a . . . . . . . . . . . . . . . . . . . . . . . . . 4 disc on finger III about half diam-
4a (3), When adpressed, finger I eter of tympanum. . . . . . . . . . . . . . . .
(thumb) longer than finger II. . . . . . . 5 . . . Eleutherodactylus johnstonei, p. 394
4b. When adpressed finger I shorter
than finger II . . . . . . . . . . . . . . . . . . . . . . 6
5a (4). Upper body tan with a grayish Key to Frogs of the Family
tan or brown X across back. . . . . . Hemiphractidae (Stefania) of Guyana
. . . . . Pristimantis chiastonotus, p. 386 The following key is based on informa-
5b. No X across back; back dark tion provided by Duellman & Hoogmoed
brown with or without spots . . . . . (1984), MacCulloch & Lathrop (2002), and
. . . . . . Pristimantis zeuctotylus, p. 390 MacCulloch et al. (2006).
6a (4). Upper body with brilliant col-
ors of red or green or both, in form 1a. Head wider than long; frontopari-
of spots or larger markings (red etal crests (atop head behind eyes)
very prominent . . . . . . . . . . . . . . . . . . no chevrons on back; 6–9 vomerine

. . . . . . . . . . . . . . . . . Stefania coxi, p. 396 teeth. . . . . . . Stefania ackawaio, p. 395
1b. Head length equal to or greater than
width; frontoparietal crests low or Key to Frogs of the Family Hylidae
absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 of Guyana
2a (1). Toe webbing (on foot) exten-
sive, including all but the distal- This key includes hylid genera and
most one (rarely two) phalange(s) species that were not keyed out to species
or joints . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 in the initial anuran key. The initial key
2b. Toe webbing only basal, with at covers the two species of Pseudis and
least two phalanges completely free Sphaenorhynchus lacteus, and it directs
of webbing. . . . . . . . . . . . . . . . . . . . . . . . 4 users to a separate key for Phyllomedusa,
3a (2). Only one toe phalange free of which follows this key.
webbing (but two on toe IV); The following key is based largely on
frontoparietal crests visible; maxi- information provided by Lescure & Marty
mum adult body length (snout- (2000). In addition, we used information
vent) 97 mm; found below 900 m provided by Duellman (2005), original
elev. . . . . . . . . . . Stefania evansi, p. 396 descriptions of recently described species,
3b. More than one toe phalange free and literature for highland species of
of webbing (almost three on toe Guyana and Venezuela that are not known
IV); frontoparietal crests absent; to occur in French Guiana.
maximum adult body length 67 1a. Iris of eye (in moderate-sized
mm; found in Pakaraima high- treefrogs) with alternating narrow
lands above 900 m . . . . . . . . . . . . . . gold and black lines radiating from
. . . . . . . . . . . . . . Stefania scalae, p. 398 area of pupil . . . . . . . . . . . . . . . . . . . . . . . 2
4a (2). Frontoparietal crests absent; 1b. Iris not as 1a, regardless of size,
skin on back smooth; conspicuous although a few dark lines may be in
dorsolateral light stripes present, iris. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
eye to groin. . . . . . . . . . . . . . . . . . . . . . 2a (1). Largest Osteocephalus, adult
. . . . . . . . . . . . Stefania roraimae, p. 397 body length reaching 100 mm or
4b. Frontoparietal crests low; skin more; vocal sacs paired in males;
shagreened (textured between cranial crests (longitudinal bony
smooth and granular) or granular; ridges between eyes) present; toes
bold dorsolateral stripes absent . . . . 5 nearly fully webbed; no row of
5a (4). Mottled color pattern; skin tubercles on outer edge of tarsus
shagreened to granular; found of foot. . . . . . . . . . . . . . . . . . . . . . . . . .
below 900 m . . . . . . . . . . . . . . . . . . . . . . . . . . . Osteocephalus taurinus, p. 412
. . . . . . . . . . . . Stefania woodleyi, p. 398 2b. Smaller Osteocephalus, largest
5b. Color pattern not mottled; skin adults less than 65 mm; vocal sac
shagreened but not granular; single, medial, in males; cranial
found above 1000 m . . . . . . . . . . . . . . . 6 crests not apparent; toes about
6a (5). When adpressed, finger II three-fourths webbed; low row of
reaches base of disc or last joint tarsal tubercles. . . . . . . . . . . . . . . . . . .
of finger I; color pattern of dark . . . . . . Osteocephalus oophagus, p. 411
chevrons on back; five vomerine 3a (1). Skin particularly thick and
teeth. . . . Stefania ayangannae, p. 395 glandular; when handled in life,
6b. When adpressed, finger II does not skin secretions copious, viscous,
reach terminal phalange of finger I; sticky . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3b. Skin of usual thinness of most brown . . . . . . . . . . . . . . . . . . . . . . . . . . .

frogs; secretions not as noxious as . . . . . . . Osteocephalus leprieurii, p. 411
in 3a. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 10b. Prominent tubercles on outer edge
4a (3). No brown bars across thighs; of tarsus and lower arm; back green
black or bluish-black spot behind or brown with green on sides . . . . . 11
insertion of arm; concealed parts 11a (10). Back and head with promi-
beneath arms and legs and web- nent tubercles; toes nearly com-
bing on hands and feet bright red pletely webbed; numerous promi-
. . . . . Trachycephalus coriaceus, p. 418 nent tubercles on outer edge of
4b. Brown bars across thighs; colora- lower arm; back primarily moss-
tion not as 4a. . . . . . . . . . . . . . . . . . . . . . 5 like green . . . . . . . . . . . . . . . . . . . . . . . .
5a (4). Males with single, median vo- . . . . . . . Osteocephalus cabrerai, p. 410
cal sac when expanded; belly with 11b. Tubercles on back and head not
abundant brown speckling . . . . . . . so prominent; toes about two-
. . . . Trachycephalus hadroceps, p. 419 thirds webbed; few prominent tu-
5b. Males with apparently paired vo- bercles on outer edge of lower arm;
cal sacs when expanded, one on back green or brown, with green
each side at rear of mouth; belly on sides. . . . . . . . . . . . . . . . . . . . . . . . . .
lacks significant speckling . . . . . . . . . . 6 . . . . . . . Osteocephalus buckleyi, p. 409
6a (5). Skin on back with low tuber- 12a (7). Discs on fingers and toes
cles; iris with black flecks . . . . . . . .
truncate (viewed from above),
. . . . . Trachycephalus typhonius, p. 419
wider than long (Fig. 5G); loreal
6b. Skin on back with numerous large
region (between eye and nostril
tubercles; iris with dark cross or
and down toward upper lip)
horizontal dark brown to black bars
concave or depressed; can bend
. . . . Trachycephalus resinifictrix, p. 419
finger I (innermost) and toe I off
7a (3). Skin of head co-ossified with
to side; fingers with webbing
skull; males with paired vocal sacs
reduced or absent; webbing re-
at rear of mouth, one on each side . . 8
duced or absent between first two
7b. Skin of head as in most frogs not
co-ossified with skull; males with toes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
single, median vocal sac. . . . . . . . . . . 12 12b. Discs on fingers and toes more
8a (7). Smallest Osteocephalus, adult rounded (viewed from above); oth-
body length less than 43 mm; erwise not as 12a, although certain
posterior surface of thighs black . . species with concave loreal region,
. . . . .Osteocephalus exophthalmus, p. 410 certain species with webbing of
8b. Not as 8a . . . . . . . . . . . . . . . . . . . . . . . . . 9 fingers and toes similar to 12a. . . . . 19
9a (8). Cranial crests present but low; 13a (12). Snout very elongate and
distinct, large membrane in axilla with narrow, projecting, pointed,
(armpit) between body and upper cutaneous appendage; broad black
arm; tympanum (ear drum) about bars over orange background on
half diameter of eye. . . . . . . . . . . . . . rear of thighs . . . . . . . . . . . . . . . . . . . .
. . . . . Osteocephalus phasmatus, p. 412 . . . . . . . . . . Scinax proboscideus, p. 416
9b. No cranial crests; axillary mem- 13b. Not as 13a . . . . . . . . . . . . . . . . . . . . . . 14
brane absent or small; tympanum 14a (13). Top of head, above ears,
about three-fourths or more diam- and back with conspicuous tuber-
eter of eye . . . . . . . . . . . . . . . . . . . . . . . . 10 cles; black spots in groin; black
10a (9). Tubercles absent from outer bars over blue background on rear
edge of tarsus and lower arm; back of thighs . . . . Scinax nebulosus, p. 415
14b. Small to moderate tubercles or and inner side of tarsus . . . . . . . . . .

pustules on head or back, if any; . . . . . . . . . . . . . . . ‘‘Hyla’’ helenae, p. 402
thigh coloration not as 14a. . . . . . . . 15 20b. Tan or brown on back; otherwise
15a (14). Smallest Scinax of Guyana, not as 20a . . . . . . . . . . . . . . . . . . . . . . . . 21
adult body length less than 25 mm; 21a (20). Dorsal (upper) surfaces of
three dark brown longitudinal the upper arms brilliant white; in
stripes on back—one at middle of preservative, back dark brown;
back (vertebral), which may be chin, throat, and belly gray . . . . .
incomplete in length, and two . . Dendropsophus grandisonae, p. 400
dorsolateral (broad, from behind 21b. Not as 21a . . . . . . . . . . . . . . . . . . . . . . 22
eye nearly to groin); indistinct 22a (21). With legs pressed against
paravertebral dark stripes present body in normal resting position,
on some individuals . . . . . . . . . . . . . . cream or white line above vent
. . . . . . . . . . . . . Scinax trilineatus, p. 417 appears similar to those on rear of
15b. Pair of broad dorsolateral stripes ankles; back brown or beige, with
may be present, but otherwise not dark brown markings (usually
as 15a . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 crossbars or chevrons) . . . . . . . . . . .
16a (15). Presence of yellow or orange . . . . . . . Dendropsophus minutus, p. 401
coloration in groin; no light spots 22b. No white line as in 22a, although
on back. . . . . . . . . . . . . . . . . . . . . . . . . . . 18 back may appear similar (in D.
16b. No yellow or orange coloration brevifrons) . . . . . . . . . . . . . . . . . . . . . . . . 23
in groin. . . . . . . . . . . . . . . . . . . . . . . . . . . 17 23a (22). Tympanum distinct; two
small light cream spots or bars
17a (16). Brown stripe, snout to
beneath eyes and a large one on
shoulder; back with light spots on
each side of body; conspicuous
gray, beige, or brown background
dark brown crossbar and/or large
. . . . . . . . . . . . Scinax boesemani, p. 415
dark brown patch on back . . . . . . .
17b. No brown stripe as in 17a; no
. . . . . Dendropsophus brevifrons, p. 399
light spots on back. . . . . . . . . . . . . . .
23b. Tympanum inconspicuous; back
. . . . . . . . . . Scinax cruentommus, p. 415 tan or beige, without conspicuous
18a (16). Two or more dorsolateral, dark markings (except tiny brown
dark brown, longitudinal stripes on dots). . . . . . . . . . . . . . . . . . . . . . . . . . . . .
back; lighter brown on back; yel- . . . . Dendropsophus minusculus, p. 401
low coloration in groin and on rear 24a (19). Dorsum (back) basically
surface of thighs, not orange . . . . . green (light or dark, rarely greenish
. . . . . . . . . . . . . . . . . Scinax ruber, p. 416 brown, although may be white in
18b. Darker brown on back, with preservative), with or without dif-
darker markings but without ferently colored markings (H.
stripes; vivid orange or reddish punctatus vermillion on back at
orange in groin and rear surface of night, with green sides). . . . . . . . . . . . 25
thighs. . . . Scinax cf. x-signatus, p. 417 24b. Basically brown on back, with or
19a (12). Adult body length less than without markings . . . . . . . . . . . . . . . . . 30
25 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 25a (24). Skin on back granular . . . . . . 26
19b. Adult body length greater than 25b. S k i n o n b a c k s m o o t h o r
25 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 shagreened (with texture but not
20a (19). Deep green on back (brown- granular) . . . . . . . . . . . . . . . . . . . . . . . . . 27
ish olive in preservative) with 26a (25). Skin on back with fine gran-
numerous well-defined brown ules; fingers II–IV about one-third
spots; orange on thighs, hind feet webbed; adult males with spine at
base of thumb (Fig. 6C); belly pale rounded, or conical to long, point-
green to bluish green; iris of eye ed, triangular; Fig. 6D) . . . . . . . . . . . 31
orange or reddish orange. . . . . . . . . 30b. Back of heel rounded, without
. . . . . . . . Hypsiboas cinerascens, p. 403 calcar. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
26b. Skin on back with coarse gran- 31a (30). Calcar quite conspicuous,
ules; no webbing on fingers; adult elongate, pointed, triangular . . . . . . 32
males lack spine at base of thumb; 31b. Calcar shorter, although may be
belly blue; iris silver (daytime) or triangular . . . . . . . . . . . . . . . . . . . . . . . . . 33
bronze (night) . . . . . . . . . . . . . . . . . . . 32a (31). Fingers with only basal web-
. . . . . . . . . . . . . . Hypsiboas liliae, p. 406 bing on three outer fingers; sides
27a (25). Middle of back with red- with blue tint and, toward rear,
dish brown marking and longitu- dark brown vertical bars; lower
dinal stripe . . . . . . . . . . . . . . . . . . . . . eyelid without extensive light retic-
. . . . . . Hypsiboas ornatissimus, p. 407 ulation; belly whitish or translu-
27b. Not as 27a . . . . . . . . . . . . . . . . . . . . . . 28 cent gray . . . . . . . . . . . . . . . . . . . . . . . .
28a (27). Narrow vermillion or purple . . . . . . . . . Hypsiboas calcaratus, p. 403
dorsolateral stripe on narrow lon- 32b. Three outer fingers almost but
gitudinal ridge of skin (from eye not completely webbed; blue sides
toward hip), edged above by tan or without bars but with light speckles
yellow stripe . . . . . . . . . . . . . . . . . . . . . instead; lower eyelid covered with
. . . . . . . . . . Hypsiboas punctatus, p. 407 reticulum of light color; belly
28b. No dorsolateral ridge of skin or yellowish orange . . . . . . . . . . . . . . . . .
light stripes (although some H. . . . . . . Hypsiboas geographicus, p. 405
sibleszi have a cream or yellow 33a (31). Large treefrog, adult body
stripe) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 length up to 132 mm; calcar short,
29a (28). Adult body length less than triangular; three outer fingers and
40 mm; found at localities above toes completely webbed; adult
400 m elev.; back with (or with- males with rounded spine at base
out) small reddish brown spots or of thumb . . . . Hypsiboas boans, p. 402
flecks and, on some individuals, 33b. Adult body length 50 mm or less;
small white or yellow spots; belly calcar tubercular or conical, not
pale green with blue tint; no white triangular; fingers with vestigial or
lines above vent or on rear of no webbing; toes not completely
ankles but possible pale yellow webbed to discs; no spine on
line between eyes and from eye thumb . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
toward vent . . . . . . . . . . . . . . . . . . . . . 34a (33). Calcar small, conical (Fig.
. . . . . . . . . . . Hypsiboas sibleszi, p. 408 6D); fingers with vestigial webbing;
29b. Adult body length up to 75 mm; from high elevation or upland
found at lowland localities; back localities higher than 800 m . . . . . .
with or without small dark brown . . . . . . . . . . . Hypsiboas roraima, p. 407
dots; belly orange; with legs 34b. Calcar small or medium, tubercu-
pressed against body in normal lar; fingers without webbing; from
resting position, white line above lowland or highland localities . . . . . 35
vent appears similar to those on 35a (34). Calcar medium, tubercular;
rear of ankles . . . . . . . . . . . . . . . . . . . . toes more than half webbed; eyes
. . . . . . . . . . Hypsiboas crepitans, p. 404 not completely black; from low-
30a (24). Back of heel of foot with a land localities. . . . . . . . . . . . . . . . . . . .
calcar—a dermal appendage (small, . . . . . . . . . . Hypsiboas fasciatus, p. 404
35b. Calcar small, blunt, tubercular 40b. Fingers without webbing or with
(Fig. 6D); toes only about one- basal webbing only; toes about half
third webbed; eyes completely webbed; snout blunt or truncate;
black; from highland or upland somewhat larger frogs, maximum
localities at 650 m or higher. . . . . . adult body length 37–50 mm . . . . . . 41
. . . . . . . . . Myersiohyla kanaima, p. 409 41a (40). Tympanum (ear drum)
36a (30). Finger I half webbed, other about two-thirds diameter of eye;
fingers fully webbed; toes com- width of disc of finger III little
pletely webbed; webbing black at more than half diameter of tympa-
base, yellowish orange distally; num; with arm perpendicular to
belly yellowish orange with numer- body, membrane in armpit extends
ous large black spots. . . . . . . . . . . . . to about one-third length of upper
. . . . Dendropsophus marmoratus, p. 400 arm; undersides light, often with-
36b. Fingers and toes with less web- out brown spots on throat or chest
bing; coloration not as 36a . . . . . . . . 37 . . . . . . . . . . Tepuihyla talbergae, p. 418
37a (36). Upper surfaces of body, 41b. Tympanum about half diameter
arms, and legs bright white to of eye; width of disc of finger III
light beige with sharply contrast- about same diameter as tympa-
ing brown patches; belly bright num; with arm perpendicular to
reddish orange . . . . . . . . . . . . . . . . . . body, membrane in armpit ex-
. . . . . . Dendropsophus leucophyllatus, tends to about middle of upper
arm; undersides light, with
p. 400
brown spots on throat (and
37b. Not as 37a . . . . . . . . . . . . . . . . . . . . . . 38
sometimes chest and belly) . . . . .
38a (37). Smaller frogs (maximum
. . . . . . . . . . . . . ‘‘Hyla’’ warreni, p. 402
adult body length 32–42 mm);
from upland to highland localities
in Pakaraima Mountains . . . . . . . . . . 40 Key to Hylid Frogs of the Genus
38b. Larger frogs (maximum adult Phyllomedusa of Guyana
body length up to 75 mm); from
lowlands . . . . . . . . . . . . . . . . . . . . . . . . . . 39
39a (38). Toes about half webbed; The following key is based on informa-
brown crossbars on back (which tion provided by Lescure & Marty (2000),
is cream by day, light brown by primarily, and by Rodrı́guez & Duellman
night); rear of thighs beige or (1994).
brown without markings . . . . . . . . . 1a. Dark vertical bars on sides of body
. . . . . Hypsiboas multifasciatus, p. 406 toward rear (flanks) and on thighs
39b. T o e s a b o u t t h r e e - f o u r t h s . . . . . . . Phyllomedusa hypochondrialis,
webbed; no brown crossbars on p. 413
back (which is pale cream, green, 1b. Not as 1a . . . . . . . . . . . . . . . . . . . . . . . . . 2
or gray by day, beige to brown at 2a (1). Throat, chest, and underside
night); rear of thighs pale yellow of arms and legs brown; flanks and
with brown bars or spots . . . . . . . . hidden surfaces of limbs green with
. . . . . . . . . . Hypsiboas crepitans, p. 404 small light spots that are not edged
40a (38). Fingers about one-third with dark color . . . . . . . . . . . . . . . . . .
webbed; toes about three-fourths . . . . . . . . . Phyllomedusa tarsius, p. 413
webbed; snout rounded; smaller 2b. Throat and chest gray or grayish
frogs, maximum adult body length brown; light spots on flanks edged
32 mm. . . . . . . Hypsiboas lemai, p. 405 with dark color . . . . . . . . . . . . . . . . . . . . 3
3a (2). The largest Phyllomedusa, 2b. Absence (usually) of bright flash-

maximum adult body length about colors as in 2a, but if present
135 mm; thick glandular fold or (pinkish orange to red), no black
ridge on each side of upper body swollen orbs on sides of body
(dorsolateral) extends to level of posteriorly . . . . . . . . . . . . . . . . . . . . . . . . . 3
groin; discs on fingers large, about 3a (2). Small frogs, maximum body
size of tympanum; first and second length of adults 35 mm or less . . . . . 4
toes (hind foot) equal in length. . . 3b. Larger frogs, adult body length
. . . . . . . . . Phyllomedusa bicolor, p. 413 greater than 35 mm . . . . . . . . . . . . . . . . 8
3b. Dorsolateral ridge extends toward 4a (3). Belly yellow in life; rear sur-
rear only to about midbody; discs face of thigh black with yellow,
on fingers smaller than tympanum; orange, or red spotting or mottling;
first toe longer than second . . . . . . dark brown or black triangular
. . . . . . . Phyllomedusa vaillantii, p. 414 seat patch at vent . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . Adenomera lutzi, p. 421
4b. Not as 4a . . . . . . . . . . . . . . . . . . . . . . . . . 5
Key to Frogs of the Family 5a (4). Inconspicuous flattened discs
Leptodactylidae of Guyana at tips of toes; no dark spots on
back . . . . . Adenomera andreae, p. 420
The following key is based largely on 5b. Toe tips slightly rounded but
information provided by Lescure & Marty without discs; dark spots on back,
(2000). Additional information was ob- although possibly few and widely
tained from Lynch (1970, 1989), Gorzula scattered . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
& Señaris (1999), Heyer & Thompson 6a (5). Small frogs, body length of
(2000), Heyer & Heyer (2006), and Kok adults less than 25 mm; conspicu-
et al. (2007). ous dark bar or two large dark
spots between eyes on top of head;
1a. Tiny frogs, maximum body (snout-
no tubercle on tarsus of foot . . . . .
vent) length less than 20 mm and . . . . . . Adenomera hylaedactyla, p. 420
with one or two tubercles on outer 6b. Larger frogs, adult body length
side of lower arm, fringes on sides more than 25 mm; only very small
of toes, snout pointed (viewing dark spots, if any, between eyes
head from above), tympanum (ear- atop head; rear of tarsus of foot
drum) concealed beneath skin, toes with conical tubercle . . . . . . . . . . . . . . . 7
only basally webbed, and several 7a (6). Skin of back basically smooth,
small tubercles on eyelids . . . . . . . . with a few warts; no parotoid
. . . . Pseudopaludicola boliviana, p. 429 glands on neck; inconspicuous fold
1b. Frogs larger than 20 mm in body of skin from rear edge of eye to
length, or if not, other characters shoulder . . . . . . . . . . . . . . . . . . . . . . . .
in 1a not all present in that . . . . . . . . . . Physalaemus cuvieri, p. 428
combination . . . . . . . . . . . . . . . . . . . . . . . 2 7b. Numerous flat glands and pointed
2a (1). Side of body near rear with tubercles on back; elongate paro-
conspicuous eye-like swollen orbs, toid gland on each side of neck at
black with light gray or blue shoulder; no fold of skin from rear
markings; bright orangish red of eye to shoulder. . . . . . . . . . . . . . . .
flash-marks in groin, behind upper . . . . Engystomops cf. pustulosus, p. 421
leg, behind knee, beneath lower 8a (3). Back essentially black with
leg, and on front of ankle . . . . . . . . brightly contrasting golden yellow
. . . . . . . . Pleurodema brachyops, p. 429 dorsolateral stripe on each side,
snout to groin; bright red spots in yellowish, marbled with black . . . .

groin and on thighs . . . . . . . . . . . . . . . . . . . . Leptodactylus guianensis, p. 423
. . . . . . . . . . . Lithodytes lineatus, p. 428 14b. Toes lack lateral fringes and
8b. Not as 8a . . . . . . . . . . . . . . . . . . . . . . . . . 9 rudimentary webbing; rear of thigh
9a (8). Maximum adult body length black, speckled with light spots. . .
more than 60 mm. . . . . . . . . . . . . . . . . 10 . . . . Leptodactylus rhodomystax, p. 426
9b. Maximum adult body length less 15a (9). Behind eye, conspicuous dor-
than 60 mm . . . . . . . . . . . . . . . . . . . . . . 15 solateral ridge or fold on skin
10a (9). Pattern of bars on upper lips 11 along body . . . . . . . . . . . . . . . . . . . . . . . 16
10b. No bars on upper lips, although 15b. No dorsolateral folds, although a
an elongate spot may be present . . 13 short warty ridge may occur on
11a (10). Behind eye, conspicuous shoulder . . . . . . . . . . . . . . . . . . . . . . . . . . 18
dorsolateral ridge or fold on skin 16a (15). Only two distinct dorsolat-
along body . . . . . . . . . . . . . . . . . . . . . . . 12 eral folds (one on each side);
11b. No dorsolateral fold, although a upper lip area very bright cream
short warty ridge may occur on or white from snout, touching eye
shoulder . . . . . . . . . . . . . . . . . . . . . . . . . and tympanum, extending to an-
gle of jaws . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . Leptodactylus rugosus, p. 427
. . . . . Leptodactylus mystaceus, p. 425
12a (11). Dorsolateral fold long, ex-
16b. More (4 to 6) dorsal and dorso-
tending rearward to level of groin; a
lateral ridges on back, or if only
second fold from above ear (supra-
two, upper lips not as 16a. . . . . . . . . 17
tympanic) slants downward onto
17a (16). Upper lip area mostly dirty
side, accompanied by a black stripe;
cream, snout to angle of jaws; rear
reproductive males with one small
of thigh dark brown or black,
black spine on thumb and no spines marbled with white and having a
on chest; rear of thigh black with line of white dots between vent and
white dots . . . . . . . . . . . . . . . . . . . . . . . . knee . . . . Leptodactylus fuscus, p. 422
. . . . Leptodactylus pentadactylus, p. 425 17b. Upper lip area light with pinkish
12b. Dorsolateral fold usually shorter, coloration in life and possibly
extending only to sacrum; supra- black markings; rear of thigh
tympanic fold and accompanying brown, marbled with white and
black stripe very short; reproductive having one or two yellow lines . . .
males with black spines on thumbs . . . . . Leptodactylus longirostris, p. 424
and chest; rear of thigh black with 18a (15). Toes with slightly expand-
orange bars . . . . . . . . . . . . . . . . . . . . . . . ed, rounded tips; toes without
. . . . . . . . Leptodactylus knudseni, p. 423 fringes on sides; back very rough
13a (10). A dorsolateral fold on each with warts . . . . . . . . . . . . . . . . . . . . . .
side and light band on upper lip; . . . . . . . Leptodactylus rugosus, p. 427
back lacks paired brown spots . . . . 14 18b. Toe tips not expanded, more-or-
13b. Long ridges on back in addition less pointed; toes with fringes on
to dorsolateral folds; elongate dark sides; back less rugose, although
spot may be on upper lip; back warts may be present . . . . . . . . . . . . . 19
with pattern of more-or-less 19a (18). Undersides covered with
paired, conspicuous roundish black anastomosing spots . . . . . . . .
brown spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leptodactylus petersii, p. 426
. . Leptodactylus macrosternum, p. 424 19b. Not as 19a . . . . . . . . . . . . . . . . . . . . . . 20
14a (13). Toes with lateral fringes and 20a (19). Maximum adult body length
rudimentary webbing; rear of thigh greater than 45 mm; often, indis-
tinct bars on upper lip area extend- 6b. Not so small, yet slender relative
ing to rear edge of eye; often, light to length; tentacle essentially below
spots on throat . . . . . . . . . . . . . . . . . . nostril . . . . . Oscaecilia zweifeli, p. 437
. . . . . . Leptodactylus leptodactyloides, 7a (6). Secondary annular grooves on
p. 424 rear of body (Fig. 6F) number 21–
20b. Maximum adult body length less 34; body color purple in life. . . . . .
than 45 mm; often, distinct bars . . . . . . . . . . . Microcaecilia rabei, p. 439
on upper lip area extending to 7b. Number of secondary annular
beneath eye; usually no light spots grooves not in range of 7a; body
on throat . . . . . . . . . . . . . . . . . . . . . . . color brown (may be reddish or
. . . . . . . . Leptodactylus validus, p. 427 grayish brown, approaching pur-
ple) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Key to the Gymnophiona (Caecilians) 8a (7). Secondary annular grooves on
of Guyana rear of body number approximate-
ly 40. . . . . . . Microcaecilia iyob, p. 439
The characters used in the following key 8b. Secondary annular grooves fewer
are based primarily on information pre- than 20 . . . . . . . . . . . . . . . . . . . . . . . . . .
sented by Taylor (1968). . . . . . . . . Microcaecilia cf. rabei, p. 440
9a (5). Body strongly laterally com-
1a. Posterior to vent, a distinct but pressed . . . . . Caecilia pressula, p. 437
short tail is present . . . . . . . . . . . . . . . . 2 9b. Body round in cross-section . . . . . . 10
1b. Tail absent or indistinct . . . . . . . . . . . 3 10a (9). Primary folds on body (Fig.
2a (1). Unicolored body, purple to
6F) more than 150; few secondary
black; long axis of anal opening
folds (numbering 2–25). . . . . . . . . . .
aligned with long axis of body . . .
. . . . . . . . . . . . . . Caecilia gracilis, p. 437
. . . . . . . . . . . . . Epicrionops niger, p. 438
10b. Primary folds fewer than 150;
2b. Not unicolored, a yellow stripe
many secondary folds (25–100). . . .
low on each side of body; anal
opening transverse, its long axis . . . . . . . . . . . Caecilia tentaculata, p. 437
across body . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . Rhinatrema shiv, p. 438 Key to the Major Groups of Reptilia
3a (1). Fully aquatic; posterior part of (Excluding Birds) of Guyana
body somewhat laterally com-
pressed; primary annuli-like folds
on body often difficult to see . . . . . The characters used in the following key
. Typhlonectes compressicauda, p. 440 are based on information presented by
3b. Not as 3a . . . . . . . . . . . . . . . . . . . . . . . . . 4 Savage (2002) and Duellman (2005).
4a (3). Nostrils present, although pos-
sibly small . . . . . . . . . . . . . . . . . . . . . . . . . 5 1a. Body encased in a bony shell; no
4b. No external nostrils visible. . . . . . . teeth on jaws . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . Caecilita iwokramae, p. 438 . . . . . . . . . Testudinata (turtles), p. 366
5a (4). Eyes covered over by bone, 1b. No bony shell around body; teeth
although possibly partly visible. . . . . 6 present . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
5b. Eyes in sockets not covered by 2a (1). Arms and legs present, al-
bone. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 though possibly quite small . . . . . . . . 3
6a (5). Very small and slender animal, 2b. No arms or legs . . . . . . . . . . . . . . . . . . . 4
earthworm-like; tiny tentacle pos- 3a (2). Long axis of anal opening
terior to, not directly below nostril aligned in same direction as long
(Fig. 6E). . . . . . . . . . . . . . . . . . . . . . . . . . . 7 axis of body; toes (hind feet)
webbed; nostrils on top of elongate 2b. Average total length of adults great-
snout . . . . . . . . . . . . . Crocodylia, p. 349 er than 3 m, maximum about 5 m;
3b. Long axis of anal opening perpen- three or more large dark blotches on
dicular to long axis of body; toes sides of jaws; body black, with
not webbed; nostrils usually on yellow or white crossbands; ventral
side of less elongate face . . . . . . . . . collar of 2 transverse rows of slightly
. . . . . . . . . . . . . . . . . . . . . . . lizards, p. 350 enlarged scales; ventral scales in 25–
4a (2). Eyelids absent; scales not rect- 29 transverse rows (collar to vent);
angular and usually not equal in midbelly region white, contrasting
size around body (with exceptions) with black on sides; upper eyelids
. . . . . . . . . . . . . . . . . . . . . . . snakes, p. 356 flat, striated. . . . . . . . . . . . . . . . . . . . . . .
4b. Eyelids present; small rectangular . . . . . . . . . . . . Melanosuchus niger, p. 442
scales around body . . . . . . . . . . . . . . 3a (1). Ear coverlets colored similar to
. . . . . amphisbaenians (worm lizards), side of head; dorsal (upper) scales
p. 349 on body above hind leg inserts
(hips) in a minimum of 4 longitu-
dinal rows; dorsalmost two rows of
Key to the Crocodylia of Guyana
body scales well keeled . . . . . . . . . . .
The characters used in the following key . . . . . . Paleosuchus palpebrosus, p. 442
are based on information presented by 3b. Ear coverlets black or dark
Brazaitis (1973). The four species known brown, darker than color of side
from Guyana are in the family Alligator- of head; dorsal body scales above
idae. hind leg inserts in a minimum of 2
longitudinal rows; lateral body
1a. Transverse bony ridge, well-devel- scales well-keeled but dorsalmost
oped or weak, across face, between two rows of body scales often not
eyes anteriorly; iris of eyes yellow- so strongly keeled . . . . . . . . . . . . . . .
ish or greenish in life; average total . . . . . . . Paleosuchus trigonatus, p. 442
length of adults, including tail,
greater than 1.5 m . . . . . . . . . . . . . . . . . 2
1b. No bony ridge connecting eyes Key to the Amphisbaenians (Worm
anteriorly; iris dark brown, reddish Lizards) of Guyana
brown, or orange; average total
length of adults usually less than The characters used in the following key
1.5 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 are based on information presented by
2a (1). Average total length of adults Gans (1963) and Hoogmoed (1973). The
less than 2 m, maximum about 2.5 four species known from Guyana are all of
m; no large dark blotches on sides the family Amphisbaenidae.
of jaws, although hatchlings have
faint gray ones; no yellow or white 1a. Scales around midbody 39 or
crossbands on body; ventral (un- fewer; preanal pores 6 or fewer
derside) collar (anterior to shoul- (Fig. 7A) . . . . . . . . . . . . . . . . . . . . . . . . . . 2
der) a single transverse series of 1b. Scales around midbody 40 or
enlarged scales; ventral (belly) more; preanal pores 7 or more . . . . . 3
scales in 20–24 transverse rows 2a (1). Preanal pores 6; supralabial
(collar to vent); midbelly region (upper lip) scales 3 (not counting
cream or yellow; upper eyelids rostral scale at tip of snout); body
raised into a high point or tubercle annuli more than 240; mental
. . . . . . . . . . . . Caiman crocodilus, p. 441 (foremost) and postmental scales
on chin fused . . . . . . . . . . . . . . . . . . . . 2a (1). Pupil vertical with lobed mar-

. . . . . . . Amphisbaena stejnegeri, p. 444 gins; toes broadened at least near
2b. Preanal pores 4; supralabial scales base; body length (snout to vent,
2; body annuli fewer than 235; not including tail) may be greater
mental scale separate and distinct than 60 mm . . . . . . . . . . . . . . . . . . . . . . . 3
from postmental . . . . . . . . . . . . . . . . . 2b. Pupil round; toes not broadened at
. . . . . . . Amphisbaena vanzolinii, p. 444 base or only slightly so; small
3a (1). Scales around midbody 65 or lizards, adult snout-vent length less
more; supralabials 4; dorsal (up- than 60 mm . . . . . . . . . . . . . . . . . . . . . . . 5
per) body color almost uniform tan 3a (2). Toes broadened throughout
to dark brown, but with a few light their length; basal webbing be-
areas; belly light . . . . . . . . . . . . . . . . . tween toes; scales on back small,
of basically one size . . . . . . . . . . . . . .
. . . . . . . . . . . . Amphisbaena alba, p. 444
. . . . . . Thecadactylus rapicauda, p. 447
3b. Scales around midbody fewer than
3b. Toes broadened at base, not
60; supralabials 3; body color,
throughout length (Fig. 7B); no
above and below, with patchwork
webbing between toes; scales on
of dark brown or black and light
back (dorsum) include enlarged
(whitish) contrasting areas . . . . . . . tubercles among the granules. . . . . . 4
. . . . . . . Amphisbaena fuliginosa, p. 444 4a (3). Base of fourth toe without
enlarged subdigital lamellae, al-
Key to the Lizards of Guyana though present further along toe;
nearly a 908 angle between fourth
and fifth toes; distance between
The following key is based largely on dorsal tubercles greater than width
that of Avila-Pires (1995) for the lizards of of tubercles . . . . . . . . . . . . . . . . . . . . . .
Brazilian Amazonia. We followed her . . . . . . . Hemidactylus mabouia, p. 446
basic system even where no quotation 4b. Base of fourth toe with enlarged
marks are shown but modified wording subdigital lamellae that continue
for clarity, excluded species not document- along toe (Fig. 7B); very acute
ed to occur in Guyana, and added material angle between fourth and fifth toes;
for Guyanese lizards that were not includ- dorsal (upper body) tubercles rela-
ed in Avila-Pires’ book because they were tively large, distance between them
not known to occur in Brazilian Amazo- equal to or less than width of
nia. Additional references that were useful tubercles . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . Hemidactylus palaichthus, p. 447
in preparing the key include Peters &
5a (2). Claws permanently exposed,
Donoso-Barros (1970), Hoogmoed
with two to four scales at base
(1973), and Savage (2002).
(Fig. 7C); dorsal body scales gran-
ular. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
1a. Top of head covered with very 5b. Claws enclosed in a sheath of tiny
small, granular scales; eyelids ab- scales, but one asymmetrically
sent, represented by a brille (disk of large scale may be part of sheath
immovable transparent skin) cov- (or not); dorsal body scales either
ering eye. . . . . . . . . . . . . . . . . . . . . . . . . . . 2 granular or flat and overlapping. . . . 9
1b. Top of head with some large 6a (5). Ridge of skin above each eye
scales; movable eyelids usually with one very elongate spine . . . . .
present (although absent in Gym- . Gonatodes alexandermendesi, p. 465
nophthalmus). . . . . . . . . . . . . . . . . . . . . . 12 6b. No very elongate spine over eye . . . 7
Fig. 7. A, Preanal pores, Amphisbaena alba (adapted from Gans 1962:11). B, Gekkonid subdigital
lamellae (shaded; Hemidactylus palaichthus). C, Two rows of tiny scales on side of toe (shaded), Gonatodes
humeralis. D, Anolis planiceps subdigital lamellae (shaded). E, Femoral pores (shaded), as in Polychrus. F,
Nasal scales (shaded) either in contact atop snout in Kentropyx calcarata (above) or separated by frontonasal
scales in Neusticurus rudis (below). G, Epicrates cenchria has some upper lip scales in contact with eye (above;
drawing adapted from a photo by Philippe Macquet on the internet); Corallus hortulanus (below) does not
(subocular scales shaded). Scale bars: 1 mm.
7a (6). At base of toes, scales beneath their side; conspicuous, large, sail-
toes narrower than toes; three or like, narrow, dorsal fan of skin on
four rows of tiny scales on sides of body and tail of adult males . . . . .
toes near toe tips; scales under tail . . . . . . . . . . Basiliscus basiliscus, p. 446
moderately enlarged . . . . . . . . . . . . . 14b. Not as 14a . . . . . . . . . . . . . . . . . . . . . . 15
. . . . . . . . . . Gonatodes annularis, p. 465 15a (14). Row of very distinctly en-
7b. At base of toes, scales beneath toes larged scales or spines along verte-
as wide as toes; two rows of tiny bral area (middle of back). . . . . . . . . 24
scales on side of toes near toe tips 15b. Not as 15a, although vertebral
(Fig. 7C); scales under tail much scales may be slightly enlarged in
wider than long . . . . . . . . . . . . . . . . . . . . 8 some species . . . . . . . . . . . . . . . . . . . . . . 16
8a (7). More than 100 granules 16a (15). An extendable, thin, flat
around midbody; no bold dorsal throat fan conspicuous in males
pale stripe on back; light bar near (may be inconspicuous in females);
shoulder preceded by a black spot most fingers and toes appear flat-
. . . . . . . . . Gonatodes humeralis, p. 466 tened top to bottom, along much
8b. Fewer than 100 granules around of their length bearing flat pads
midbody; light vertebral stripe below. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
(conspicuous white with black edg- 16b. No throat fan, but a sac-like
es in males; tan or white edged with throat expansion may be present;
dark spots in females) from snout toes not flattened . . . . . . . . . . . . . . . . . 22
to base of tail . . . . . . . . . . . . . . . . . . . . 17a (16). Widened subdigital lamellae
. . . . . . . . . . . . Gonatodes vittatus, p. 467 relatively narrow (usually less than
9a (5). Dorsal scales, although very twice width of toe; Fig. 7D);
small, flat and overlapping . . . . . . . . 10 ventral (belly) scales relatively
9b. Dorsal scales granular . . . . . . . . . . . . 11 large, keeled, overlapping well. . . . . 18
10a (9). Dorsal scales keeled; claw 17b. Many subdigital lamellae very
sheath comprised of four scales . . wide (clearly twice width of toe or
. . . . . . Chatogekko amazonicus, p. 464 more); ventral scales relatively small,
10b. Dorsal scales smooth; claw usually smooth (but keeled in some
sheath comprised of five scales . . . species), overlapping little . . . . . . . . . . 19
. . Coleodactylus septentrionalis, p. 464 18a (17). Vertebral area with 8–13
11a (9). Claw sheath on toe tips in- longitudinal rows of enlarged,
cludes one asymmetrically large keeled, overlapping scales, abrupt-
scale; spine above eye; dorsal gran- ly demarcated from small granular
ular scales keeled . . . . . . . . . . . . . . . . scales on sides of body; 60–90
. . . . . . . . Sphaerodactylus molei, p. 468 scales around midbody. . . . . . . . . . .
11b. Not as 11a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anolis auratus, p. 460
. . Pseudogonatodes guianensis, p. 467 18b. Two or more rows of somewhat
12a (1). Dorsal scales on head usually but variably enlarged dorsal scales
numerous, irregular, very small. . . . 13 along vertebral area, grading in
12b. All or most dorsal head scales size to granules on sides; 103–175
arranged in plates, relatively few in scales around midbody. . . . . . . . . . .
number and relatively large . . . . . . . 26 . . . . . . . . . . . . . . Anolis planiceps, p. 461
13a (12). A very large, flat, round scale 19a (17). Relatively small lizards (max-
below ear . . . . . . . . Iguana iguana, p. 458 imum snout-vent length 50–60 mm
13b. Not as 13a . . . . . . . . . . . . . . . . . . . . . . 14 or less), with brown or gray body
14a (13). Three outer toes with pro- that is nearly cylindrical; tail round
jecting fringe of flat scales along in cross section. . . . . . . . . . . . . . . . . . . . . 20
19b. Relatively large (maximum 24b. Vertebral crest not extending

snout-vent length greater than 70 beyond basal part of tail; fourth
mm), green or gray lizards; body finger longer than third . . . . . . . . . . . 25
and tail somewhat flattened side- 25a (24). Body somewhat flattened
to-side . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 horizontally; tufts of spinose scales
20a (19). Enlarged scales on top of on neck; scales around midbody
head between eyes (supraorbital 121–162. . . . . . . . . . . Plica plica, p. 473
semicircles) of each side of head 25b. Body basically cylindrical; no
broadly in contact between eyes; tufts of spinose scales on neck;
scales on posterior (rear) part of scales around midbody 43–69 . . . .
upper snout (posterior to swollen . . . . . . . . . . . . . . . . . . Plica umbra, p. 474
nasal regions) moderately large, 26a (12). Body scales large, cycloid
smooth, and flat . . . . . . . . . . . . . . . . (with curved rear margin), overlap-
. . . . . . . . . . . . . . . . Anolis ortonii, p. 461 ping, stiff, containing thin dermal
20b. Supraorbital semicircles of each bones; dorsal (upper surface) head
side of head separated from each scales also overlapping . . . . . . . . . . .
other between eyes by 1–3 very . . . . . . . . Mabuya nigropunctata, p. 463
small scales; scales on posterior 26b. Not as 26a; dorsal head scales not
part of upper snout relatively small or only slightly overlapping . . . . . . . 27
or variable in size, mostly with 27a (26). Nasal scales (touching or
small keels . . . . . . . . . . . . . . . . . . . . . . . containing nostrils) in contact on
. . . . . . . . . . . Anolis fuscoauratus, p. 461 middorsal area of snout (Fig. 7F) . . 28
21a (19). Tail somewhat laterally 27b. Nasal scales separated by 1–2
compressed but without conspicu- frontonasal scales on top of snout
ous crest of distinctive scales . . . . . (Fig. 7F). . . . . . . . . . . . . . . . . . . . . . . . . . 33
. . . . . . . . . . . . . . Anolis punctatus, p. 462 28a (27). Ventral (belly) scales dis-
21b. Tail quite laterally compressed tinctly keeled . . . . . . . . . . . . . . . . . . . . . 29
and with conspicuous crest of 28b. Ventral scales smooth . . . . . . . . . . . 31
enlarged scales on top. . . . . . . . . . . . 29a (28). Median dorsal scales much
. . . . . . . . . . . . . . . . Anolis aeneus, p. 459 larger than the lateral scales,
22a (16). Sac-like throat expansion strongly overlapping, keeled, in
present; crest of enlarged, pointed 13–27 longitudinal rows . . . . . . . . . . . 30
scales forming a central row on 29b. Median dorsal scales only slightly
throat; femoral pores under thighs larger than the lateral scales, hex-
present (Fig. 7E); cone- or turret- agonal, juxtaposed, keeled, not in
shaped eyes, lids partially fused . . . longitudinal rows . . . . . . . . . . . . . . . .
. . . . . . . . Polychrus marmoratus, p. 462 . . . . . . . . . . Kentropyx calcarata, p. 471
22b. Not as 22a . . . . . . . . . . . . . . . . . . . . . . 23 30a (29). Enlarged dorsals slightly
23a (22). Tail shorter than body smaller than ventrals, the keels
(snout-vent) length, somewhat flat- forming 13–17 continuous longitu-
tened horizontally, very spiny . . . . dinal ridges . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . Uracentron azureum, p. 475 . . . . . . . . . . . . . Kentropyx striata, p. 472
23b. Not as 23a . . . . . . . . . . . . . . . . . . . . . 30b. Enlarged dorsals much smaller
. . . . . . . . . . . Tropidurus hispidus, p. 475 than ventrals, in 22–27 longitudi-
24a (15). Vertebral crest from neck to nal rows, keels not continuous and
tip of tail; third and fourth fingers directed towards the median line of
about of equal length . . . . . . . . . . . . the back . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . Uranoscodon superciliosus, p. 476 . . . . . . . . . Kentropyx borckiana, p. 471
31a (28). Maximum of ten enlarged 38b. Sides, arms, and legs in life very
ventrals in a row across midbody dark brown to black; tail gray,
(not including smaller ventrolateral bluish gray, or tan above with gray
scales) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 iridescence; from Rupununi Savan-
31b. More than 15 enlarged ventrals in na or not . . . . . . . . . . . . . . . . . . . . . . . . . 39
a row across midbody. . . . . . . . . . . . 39a (38). Anteriorly (toward head),
. . . . . . . . . . Tupinambis teguixin, p. 473 dorsolateral light stripe vivid in
32a (31). Eight ventrals in a row life, yellow or pale gold; upper lip
across midbody . . . . . . . . . . . . . . . . . . with white or otherwise light lon-
. . . . Cnemidophorus lemniscatus, p. 470 gitudinal stripe; occurs on Rupu-
32b. Ten ventrals in a row across nuni Savanna . . . . . . . . . . . . . . . . . . . .
midbody . . . . . . Ameiva ameiva, p. 469 . . . . . . Gymnophthalmus leucomystax,
33a (27). Small, almost worm-like in p. 451
length, diameter, color; limbs very 39b. Dorsolateral light stripe not vivid
much reduced; no ear opening . . . anteriorly, although more conspic-
. . . . . . . . . . . . . Bachia flavescens, p. 450 uous than posteriorly; upper lip
33b. Not as 33a . . . . . . . . . . . . . . . . . . . . . . 34 gray, black, or brown; from Rupu-
34a (33). Inner finger absent or, if nuni Savanna and elsewhere . . . . . . 40
present, clawless . . . . . . . . . . . . . . . . . . 35 40a (39). In life, belly gray or bluish
34b. All five fingers present, with gray, with light copper iridescence;
claws . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 tail gray, bluish gray, rarely gray-
35a (34). Lower eyelids distinct, mov- ish tan above (varying with stage of
able; snout-vent length may be molt); only females exist; from
longer than 50 mm. . . . . . . . . . . . . . . . 36 Rupununi Savanna or not . . . . . . .
35b. Eyelids absent, an immovable . Gymnophthalmus underwoodi, p. 452
brille (disk of transparent skin) 40b. In life, belly cream to tan with
covering eye; snout-vent length less copper iridescence; tail tan or
than 50 mm . . . . . . . . . . . . . . . . . . . . . . 38 reddish tan above, with gray iri-
36a (35). Dorsal and ventral scales in descence; males and females exist;
two longitudinal rows of trans- not from Rupununi Savanna, but
versely enlarged scales . . . . . . . . . . . localities north thereof . . . . . . . . . . .
. . . . . . . . . . . . . . . . Iphisa elegans, p. 454 . . . . . . Gymnophthalmus cf. speciosus,
36b. Dorsal, lateral, and ventral scales p. 452
more similar in size . . . . . . . . . . . . . . . 37 41a (34). Dorsal body scales not of
37a (36). Ventral scales quadrangular uniform size but with distinct
in shape, occurring in neat trans- enlarged tubercles or some longi-
verse and longitudinal rows . . . . . . tudinal rows of enlarged scales; tail
. . . . . . . . . . . Riolama leucosticta, p. 457 with a double dorsal crest or with
37b. Rear edge of ventral scales transverse rows of tubercles . . . . . . . 42
rounded or cycloid in shape . . . . . . 41b. Dorsal scales generally of uni-
. . . . . . . . . . . Tretioscincus agilis, p. 458 form size; tail without crests or
38a (35). Sides, arms, and legs in life transverse rows of tubercles (al-
coppery tan or brown, definitely though scales may be keeled). . . . . . 44
not black; tail bright orange (salm- 42a (41). Rostral and mental scales
on) above in females and juveniles (on frontmost tip of snout and of
(but tan or tannish gray in males or chin, respectively) with striations
if regenerated or about to shed); or grooves; three rows of similar-
occurs on Rupununi Savanna . . . . sized scales on snout between the
. . . . . . Gymnophthalmus vanzoi, p. 454 rostral and frontal (large median
scale between eyes); a pair of scales 47b. Dorsal scales number 35–40; in-
behind the mental, not a single terparietal scale usually not as
large postmental . . . . . . . . . . . . . . . . . wide, with lateral margins parallel
. . . . . Echinosaura sulcarostrum, p. 451 or slightly diverging; only females
42b. Not as 42a . . . . . . . . . . . . . . . . . . . . . . 43 exist . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
43a (42). Tubercles along dorsal sur- . . . . . . Leposoma percarinatum, p. 455
face of tail form a continuous 48a (45). Interparietal scale extends
double row; two transverse rows somewhat posteriorly beyond the
of scales on underside of tail parietal scales, these all followed
correspond to two transverse rows by occipital scales on nape of neck,
on side of tail . . . . . . . . . . . . . . . . . . . . these in turn followed by smaller
. . . . . . . Neusticurus bicarinatus, p. 456 scales of back . . . . . . . . . . . . . . . . . . . . . 49
43b. Tubercles along dorsal surface of 48b. Interparietal and parietal scales
tail form a discontinuous row, with forming a straight margin across
smaller scales in between; two back of head. . . . . . . . . . . . . . . . . . . . . . 50
transverse rows of scales on under- 49a (48). Head black, including upper
side of tail correspond to 3–6 lip scales; belly scales (ventrals)
transverse rows on side of tail. . . . . broadly keeled; one frontonasal
. . . . . . . . . . . . . . Neusticurus rudis, p. 456 scale atop snout between scales
44a (41). Dorsal scales in longitudinal with nostrils . . . . . . . . . . . . . . . . . . . . .
rows on body . . . . . . . . . . . . . . . . . . . . . . . . . Pantepuisaurus rodriguesi, p. 457
. . . . . . . . . . Cercosaura ocellata, p. 451 49b. Head brown, but upper lip scales
44b. Dorsal scales in transverse rows contrasting white; ventrals smooth;
across body or in diagonal rows on frontonasal scale divided, with one
body. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45 on each side of midline of snout . .
45a (44). Large scales atop back of . . . . . . . . . . . . Cercosaura argulus, p. 450
head (parietals and interparietal) 50a (48). Nasal scales divided (¼nostril
followed by smaller scales similar between two scales); ventral scales
to dorsal body scales. . . . . . . . . . . . . . 46 hexagonal, in transverse rows but
45b. Not as 45a; distinct, somewhat not longitudinal rows . . . . . . . . . . . . .
enlarged occipital scales on nape of . . . . . . . . . Kaieteurosaurus hindsi, p. 455
neck . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48 50b. Nostril within one scale; ventrals
46a (45). Large interparietal and pa- quadrangular, in both transverse
rietal scales atop back of head and longitudinal rows . . . . . . . . . . . . . 51
form a straight margin across back 51a (50). Four large supraocular
of head . . . . . . . . . . . . . . . . . . . . . . . . . . scales (above eye) on each side . . . . 52
. . . . . . . Alopoglossus angulatus, p. 448 51b. Three large supraoculars, each
46b. Interparietal and parietal scales side . . . . Arthrosaura reticulata, p. 449
form a curved margin across back 52a (51). Lower eyelid with semi-
of head . . . . . . . . . . . . . . . . . . . . . . . . . . . 47 transparent disc; 45 scales around
47a (46). Dorsal scales (on vertebral midbody; 40–42 keeled temporal
row from nape of neck back to rear scales (on side of head between
edge of hind legs at hip) number postoculars and ear) . . . . . . . . . . . . .
30–35; interparietal scale (largest . . . . . . . Arthrosaura guianensis, p. 448
scale atop back of head) very wide, 52b. Lower eyelid with opaque disc;
with diverging lateral margins; 37 scales around midbody; 32–33
males and females exist . . . . . . . . . . smooth temporals . . . . . . . . . . . . . . . .
. . . . . . . . . . Leposoma guianense, p. 455 . . . . . . . Arthrosaura hoogmoedi, p. 449
Key to the Genera and Most Species of erably with a dissecting microscope (e.g.,
Snakes of Guyana apical pits on upper body scales; grooves
on rear maxillary teeth). Reference also is
Be respectful in the presence of living made to hemipenes. These are the copula-
snakes. Rattlesnakes, other pit vipers, and tory organs of males, which are stored in
coral snakes, whether of colorful species or the base of the tail, paired (one on each
not, are highly venomous and dangerous side), and may be everted through the anal
to humans, as may be a few other species. opening of specimens found dead on the
Also, some of the characters used in this road or preserved in museum collections.
key require getting very close to the animal Characteristics of hemipenes can be very
or picking it up. Therefore, people who are important in snake taxonomy. When
not fully familiar with proper procedures preserving specimens, it is advisable to
for safe handling of venomous snakes fully evert the hemipenes so they will be
should not try to use this key to identify visible for later examination.
living snakes. In addition, snakes that were This key is based largely on that of
killed apparently recently, and perhaps Peters & Orejas-Miranda (1970). In addi-
found dead on the road, sometimes are tion, the following references were espe-
capable of biting in a reflex action when cially helpful: Dixon & Hendricks (1979),
handled, so these also should be handled Chippaux (1986), Savage (2002), Campbell
only with the utmost care, to avoid a & Lamar (2004), and Duellman (2005).
medical emergency.
1a. One deep pit on each side of face
Many of the genera of snakes that occur
between nostril and eye . . . . . . . . . . . . 2
in Guyana are represented by only one or
two species that occur in the country (e.g., 1b. Not as 1a . . . . . . . . . . . . . . . . . . . . . . . . . 6
Boa, Crotalus). For those genera, the 2a (1). Rattle or button of keratin on
Guyanese species can be identified with tip of tail . . . . . . . . . . . . . . . . . . . . . . . .
the key immediately below. For most . . . . . . . . . . . . . Crotalus durissus, p. 522
genera having more representatives in 2b. No rattle or button of keratin on
Guyana (Atractus, Chironius, Dipsas, Lio- tip of tail . . . . . . . . . . . . . . . . . . . . . . . . . . 3
phis, Micrurus, and Oxyrhopus), this key 3a (2). Scales of posterior subcaudals
will lead users to the relevant genus and (under tail, near tip) finely subdi-
then refer to specific generic keys that vided; keels on middorsals (upper
follow. body scales) tubercular or knobby
In some instances, a genus or species . . . . . . . . . . . . . . . . Lachesis muta, p. 523
will appear in more than one place in the 3b. Posterior subcaudal scales single
key. This reflects intrageneric or intraspe- or paired across underside of distal
cific variation in key characters, such as tail; keels (ridges) on middorsal
whether the number of scale rows around scales not knobby or tubercular . . . . 4
midbody is 17 or fewer. Variation in 4a (3). Upper body background color
Guyanese snakes also necessitates using green; yellow stripe low on each
some characters that are difficult to see. side of body . . . . . . . . . . . . . . . . . . . . .
While it would be convenient, it is not . . . . . . . . . . Bothriopsis bilineata, p. 519
possible to design a key based only on 4b. Body not green above; no yellow
conspicuous characters that are readily stripe on each side . . . . . . . . . . . . . . . . . 5
apparent to everyone, including non-spe- 5a (4). Distinct dark stripe behind
cialists. Consequently, reference is of eye; usually fewer than 18 dark
necessity made to some characters that blotches on body (before vent) . . .
are best viewed with magnification, pref- . . . . . . . . . . . . . . . Bothrops atrox, p. 520
5b. Stripe behind eye absent or indis- 12b. Belly scales same size as upper
tinct; usually more than 18 dark body scales; body color not as
blotches on body . . . . . . . . . . . . . . . . above. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
. . . . . . . . . . . . . . Bothrops brazili, p. 521 13a (12). Scales around midbody 14;
6a (1). More than 31 scales around body very slender and short . . . . . . 14
midbody (between head and vent) . . 7 13b. Scales around midbody 15 or
6b. Scales around midbody 30 or more; usually stockier and longer
fewer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 snakes . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
7a (6). All scales on top of snout 14a (13). Enlarged supraocular scales
small, same size as those on top (atop head, between eyes, and in
of head; one longitudinal dark contact with transparent scales
stripe on top of head. . . . . . . . . . . . . covering eyes). . . . . . . . . . . . . . . . . . . . . 15
. . . . . . . . . . . . . . . Boa constrictor, p. 478 14b. Enlarged supraoculars absent;
7b. Small scales on top of head, but scales around middle of tail 10 to
with somewhat enlarged scales 12; upper body background color
between and behind nostrils; head light, with dark brown stripes . . . .
color pattern not as above . . . . . . . . . 8 . . . . Siagonodon septemstriatus, p. 516
8a (7). No pits in labials (scales on 15a (14). Supraoculars rather large,
lips); top of head dark brown with larger than scales (prefrontal and
broad tan or orange stripe behind frontal) centered on top of head;
eye . . . . . . . . . Eunectes murinus, p. 480 upper body color of wide dark
8b. Pits in lip scales; head color stripes separated by inconspicuous,
pattern not as 8a. . . . . . . . . . . . . . . . . . . 9 narrow light lines; small yellow
9a (8). Some scales of upper lip con- spot on tip of snout and tip of tail
tact eye (Fig. 7G); fewer than 10 . . . . . . . . . . . . . . Epictia albifrons, p. 516
scales between the eyes across top 15b. Supraoculars smaller than pre-
of head . . . . . Epicrates cenchria, p. 480 frontal and frontal . . . . . . . . . . . . . . . . 16
9b. Upper lip scales separated from 16a (15). Four scales plus rostral (scale
eye by small scales below eye (Fig. on tip of upper snout) form border
7G); 10 scales or more between of upper lip on each side; belly
eyes across top of head . . . . . . . . . . . 10 without reticulate color pattern . . .
10a (9). Adults with green upper body . . . . . Tricheilostoma dimidiatum, p. 517
(juveniles orange or yellow) with 16b. Five scales plus rostral form
white cross-bars. . . . . . . . . . . . . . . . . . border of upper lip on each side;
. . . . . . . . . . . . . . Corallus caninus, p. 478 reticulate color pattern on belly . .
10b. Upper body brown, gray, or . . . . Tricheilostoma macrolepis, p. 517
yellow, possibly with dark saddles 17a (13). Head covered with small
edged in cream. . . . . . . . . . . . . . . . . . . scales same size as those on body
. . . . . . . . . . . Corallus hortulanus, p. 479 . . . . . . . . Typhlophis squamosus, p. 477
11a (6). Belly scales (ventrals) much 17b. Head with plates larger than
wider than long; 4 scales or more scales on body . . . . . . . . . . . . . . . . . . . . 18
around eye. . . . . . . . . . . . . . . . . . . . . . . . 20 18a (17). Scales around midbody 18 .
11b. Belly scales same size as upper . . . . . . . Typhlops minuisquamus, p. 518
body scales or slightly widened; 18b. Scales around midbody 20. . . . . . . 19
eye contained in or under a single 19a (18). White snout and white tail
scale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 ring; dorsum (upper body) dark
12a (11). Belly scales slightly widened; brown to black; 9 upper rows of
body color red with black rings . . body scales pigmented . . . . . . . . . . .
. . . . . . . . . . . . . . . Anilius scytale, p. 477 . . . . . . . . . . Typhlops reticulatus, p. 518
19b. No white snout or tail ring; 27b. Not as 27a . . . . . . . . . . . . . . . . . . . . .

dorsum yellowish brown; usually . . . . . . . Mastigodryas boddaerti, p. 499
11 dorsal scale rows pigmented. . . 28a (26). Rings of color completely,
. . . . . Typhlops brongersmianus, p. 517 or nearly completely, encircle
20a (11). About one head length in body. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
front of vent, upper body (dorsal) 28b. Body without complete rings of
scale rows at least 2 fewer than color . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
around midbody, not counting 29a (28). Venomous coral snake with
wide belly scale . . . . . . . . . . . . . . . . . . . 21 upper body black or brown and,
20b. Dorsal scale rows same number on belly, white or yellow half-rings
throughout body, rarely reduced in (red under tail); frontmost upper
number immediately before vent . . 23 tooth hollow (with venom canal)
21a (20). Dorsal scales in even num- . . . . . . . . . . . . Micrurus collaris, p. 512
ber of rows around body, not 29b. Not as 29a . . . . . . . . . . . . . . . . . . . . . . 30
counting wide belly scale . . . . . . . . . . 22 30a (29). On each side, largest scale
21b. Dorsal scales in odd number of atop head behind eye (parietal
rows around body . . . . . . . . . . . . . . . . 47 scale) contacts some scales of
22a (21). Dorsal scale rows around upper lip (supralabials). . . . . . . . . . .
midbody 12 or fewer . . . . . . . . . . . . . . . . . . Apostolepis nigrolineata, p. 482
see separate key for genus Chironius, 30b. Parietal scale separated from
p. 364 supralabials by one or two rows
22b. Dorsal scale rows around mid- of scales . . . . . . . . . . . . . . . . . . . . . . . . . . 31
31a (30). Atop snout, internasal scales
body 14 or more . . . . . . . . . . . . . . . . .
(between nostrils) fused into one
. . . . . . . . . . . . . . Spilotes pullatus, p. 507
plate across snout. . . . . . . . . . . . . . . .
23a (20). More than 18 dorsal scale
. . . . . . . . . . Pseudoeryx plicatilis, p. 505
rows at midbody. . . . . . . . . . . . . . . . . . 24
31b. Internasals paired, one on each
23b. Fewer than 18 dorsal scale rows
side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
at midbody . . . . . . . . . . . . . . . . . . . . . . . 25
32a (31). Neck extremely thin, with
24a (23). Fewer than 200 ventrals head very distinct from it; upper-
(wide belly scales from neck to most row of body scales distinctly
vent); scales around midbody 21 wider than scales of adjacent rows
. . . . . . . . . . . . . . . Ninia hudsoni, p. 500 . . . . . . . . . . Imantodes lentiferus, p. 495
24b. Ventrals more than 200; scales 32b. Not as 32a . . . . . . . . . . . . . . . . . . . . . . 33
around midbody 19 . . . . . . . . . . . . . . 33a (32). Fewer than 170 wide belly
. . . . . . . . . . . . . . . . . . Clelia clelia, p. 488 scales (ventrals) from neck to vent
25a (23). Vent covered by one scale as . . . . . . . Tantilla melanocephala, p. 508
wide as belly scales before it . . . . . . 36 33b. Ventrals more than 170 . . . . . . . . .
25b. Anal scale divided into side-by- Elapomorphus quinquelineatus, p. 493
side halves . . . . . . . . . . . . . . . . . . . . . . . . 26 34a (28). Small scale (loreal) present
26a (25). Dorsal scale rows at mid- (Fig. 8D) on side of face between
body 17. . . . . . . . . . . . . . . . . . . . . . . . . . . 27 nasal scale (enclosing or contact-
26b. Dorsal scale rows at midbody ing nostril) and preocular scale (in
fewer than 17 . . . . . . . . . . . . . . . . . . . . . 28 front of eye) . . . . . . . . . . . . . . . . . . . .
27a (26). Neck extremely thin, head . . . . Erythrolamprus aesculapii, p. 493
very distinct from it; uppermost 34b. Loreal scale absent (Fig. 8D),
row of body scales distinctly wider nasal contacts preocular. . . . . . . . . . . 35
than scales of adjacent rows. . . . . . 35a (34). Venomous coral snake;
. . . . . . . . . . . Imantodes cenchoa, p. 495 frontmost upper tooth hollow
(with venom canal). . . . . see separate 42a (36). Across top of snout, pre-
key for genus Micrurus, p. 366 frontal scales (in front of frontal
35b. Frontmost upper tooth lacks scale, the single large plate between
venom canal . . . . . . . . . . . . . . . . . . . . . the eyes) fused into one scale. . . . .
. . . . . . . . . Hydrops triangularis, p. 495 . . . . . . . . . . . Xenopholis scalaris, p. 511
36a (25). Upper body scale rows 17 42b. Prefrontals not fused, but paired,
around midbody . . . . . . . . . . . . . . . . . . 42 one on each side of snout . . . . . . . . . 43
36b. Upper body scale rows fewer 43a (42). Scales beneath tail (subcau-
than 17 at midbody . . . . . . . . . . . . . . . 37 dals) paired, one on each side . . . . . 44
37a (36). Uppermost longitudinal row 43b. Subcaudals single, one wide scale
of body scales (vertebrals) distinct- across underside of tail. . . . . . . . . . .
ly larger than scales of adjacent . . . . . . . . . . Pseudoboa coronata, p. 504
rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 44a (43). Upper body scales with a
37b. Scales of vertebral row on body keel. . . . . . . . . . . . Ninia hudsoni, p. 500
about same size as those of adja- 44b. Upper body scales smooth, with-
cent rows . . . . . . . . . . . . . . . . . . . . . . . . . 40 out a keel. . . . . . . . . . . . . . . . . . . . . . . . . 45
38a (37). Shallow longitudinal groove 45a (44). A labial scale (on upper lip)
or depression (mental groove) pres- behind eye is enlarged (Fig. 8B),
ent behind chin in middle of higher than other labials and in
underside of head (Fig. 8A) . . . . . . . 39 contact with postocular (immedi-
38b. Mental groove absent (Fig. 8A). ately behind eye), anterior temporal
. . see separate key for genus Dipsas, (behind postocular), and posterior
p. 365 temporal (behind anterior tempo-
39a (38). A labial scale (on upper lip) ral) . . . . . . . . . . . Sibon nebulatus, p. 506
behind eye is enlarged (Fig. 8B), 45b. Not as 45a . . . . . . . . . . . . . . . . . . . . . . 46
higher than other labials and in 46a (45). More than 200 wide belly
contact with postocular scale (im- scales (ventrals) neck to vent . . . . .
mediately behind eye), anterior . . . . . . . . . . . . . . . . . . Clelia clelia, p. 488
temporal (behind postocular), and 46b. Fewer than 200 ventrals . . . . . . . .
posterior temporal (behind anterior . . see separate key for genus Atractus,
temporal) . . . . . Sibon nebulatus, p. 506 p. 364
39b. Not as 39a . . . . . . . . . . . . . . . . . . . . . 47a (21). Some or all upper body
. . . . . . . . . . Imantodes lentiferus, p. 495 scales with a keel . . . . . . . . . . . . . . . . . 88
40a (37). Upper body scales with pair 47b. Upper body scales smooth, with-
of small, subtle pits or depressions out a keel. . . . . . . . . . . . . . . . . . . . . . . . . 48
(apical pits) near rear of scale (Fig. 48a (47). Vent covered by one scale as
8C), best seen with magnification wide as belly scales before it . . . . . . 65
. . . . . . . . . Drymoluber dichrous, p. 493 48b. Anal scale divided into side-by-
40b. Apical pits absent . . . . . . . . . . . . . . . 41 side halves . . . . . . . . . . . . . . . . . . . . . . . . 49
41a (40). Upper jaw teeth (on maxil- 49a (48). Scales of uppermost longi-
lary bone) solid, without hollow tudinal row on body (vertebrals)
venom canal or open venom groove distinctly larger than scales of
. . . . . . . . . Drepanoides anomalus, p. 492 adjacent rows . . . . . . . . . . . . . . . . . . . . . 50
41b. Venomous coral snake, with 49b. Scales of vertebral row about same
frontmost maxillary tooth hollow, size as those of adjacent rows . . . . . 51
with enclosed venom canal . . . . . . . 50a (49). Upper body scale rows 15
. . see separate key for genus Micrurus, around midbody . . . . . . . . . . . . . . . . .
p. 366 . . . . . . . . . . Imantodes lentiferus, p. 495
Fig. 8. A, Inconspicuous mental groove either present behind chin (shaded), Sibon nebulatus (left) or
absent, Dipsas catesbyi (right). B, Enlarged labial scale (shaded) behind eye, Sibon nebulatus. C, Tiny apical
pits on rear of body scales, Philodryas. D, Loreal scale (shaded) either present, Erythrolamprus aesculapii
(below; with large eye) or absent, Micrurus (above; with small eye). E, Anterior temporal scales (shaded)
either 2 as in a Thamnodynastes (above) or 1, Philodryas (below); both adapted from photos by W. Wüster on
the internet. F, Body scales in either extremely oblique rows, Xenodon (left) or not, most snakes (right). Scale
bars: A–D ¼ 1 mm, E, F ¼ 2 mm.
50b. Upper body scale rows 17 at ing nostril) and preocular scale (in
midbody. . . . Imantodes cenchoa, p. 495 front of eye) . . . . . . . . . . . . . . . . . . . . . . 54
51a (49). Small scale (loreal) present 51b. Loreal scale absent (Fig. 8D) . . . . 52
(Fig. 8D) on side of face between 52a (51). Very elongate head with
nasal scales (enclosing or contact- pointed snout . . . . . . . . . . . . . . . . . . . . . 53
52b. Head and snout not as 52a . . . . . (Fig. 8C), best seen with magnifica-
. . . . . . . . . Hydrops triangularis, p. 495 tion. . . . . . . Leptodeira annulata, p. 496
53a (52). Upper body color mainly 60b. Apical pits absent . . . . . see separate
tan or brown . . . . . . . . . . . . . . . . . . . . key for genus Liophis, p. 365
. . . . . . . . . . . . . . Oxybelis aeneus, p. 500 61a (54). Rearmost one or two teeth
53b. Upper body color bright green . on maxillary bone with venom
. . . . . . . . . . . . . Oxybelis fulgidus, p. 501 groove . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
54a (51). Two or more anterior tem- 61b. Rear maxillary teeth not grooved
poral scales (Fig. 8E), which are . . . . . . . Mastigodryas boddaerti, p. 499
behind and in contact with post- 62a (61). Upper body scales with one
oculars (behind and in contact with or two small, subtle pits or depres-
eye) and between upper lip scales sions (apical pits) near rear of scale
(supralabials) and parietal (rear- (Fig. 8C), best seen with magnifi-
most large plate atop head) . . . . . . . 61 cation . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
54b. One anterior temporal (Fig. 8E) 62b. Apical pits absent . . . . . . . . . . . . . . . 63
or none . . . . . . . . . . . . . . . . . . . . . . . . . . . 55 63a (62). Usually 17 scales around
55a (54). Upper body scale rows midbody; paired scales beneath tail
around midbody 17 or fewer . . . . . . 60 (subcaudals) of males 85–100, of
55b. Upper body scale rows at mid- females 78–97; male hemipenes
body more than 17. . . . . . . . . . . . . . . . 56 lack enlarged basal hooks, but
56a (55). Upper body scale rows ex- have numerous small spines . . . . . .
tremely oblique (Fig. 8F), as scales . . . . . Thamnodynastes pallidus, p. 508
of lateral rows (on sides) much
63b. Usually 19 scales around mid-
narrower than those of upper and
body; subcaudals of males 61–64,
lower rows. . . . . . . . . . . . . . . . . . . . . . . . 57
of females 54–61; male hemipenes
56b. Upper body scale rows not so
have several basalmost spines no-
oblique, but as in most other
ticeably enlarged . . . . . . . . . . . . . . . . .
snakes (Fig. 8F) . . . . . . . . . . . . . . . . . . 58
. . . . . . . Thamnodynastes ramonriveroi,
57a (56). Upper body scales in 19 or
p. 509
fewer rows at midbody. . . . . . . . . . .
. . . . . . . . . . . . Xenodon merremii, p. 510 64a (62). Belly scales, neck to vent,
57b. Upper body scales in 21 or more more than 205 . . . . . . . . . . . . . . . . . . .
rows at midbody. . . . . . . . . . . . . . . . . . . . . . . . Philodryas viridissimus, p. 503
. . . . . . . . . . . . . . Xenodon severus, p. 510 64b. Belly scales fewer than 205 . . . . .
58a (56). Upper body scales with one . . . . . . . . . . . . Philodryas olfersii, p. 503
or two small, subtle pits or depres- 65a (48). Upper body (dorsal) scale
sions (apical pits) near rear of scale rows fewer than 17 around mid-
(Fig. 8C), best seen with magnifi- body. . . . . . . . . . . . . . . . . see separate key
cation . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59 for genus Dipsas, p. 365
58b. Apical pits absent . . . . . see separate 65b. Dorsal scale rows 17 or more
key for genus Liophis, p. 365 around midbody . . . . . . . . . . . . . . . . . . 66
59a (58). Belly scales, neck to vent, 66a (65). Dorsal scale rows 17 at
more than 205 . . . . . . . . . . . . . . . . . . . midbody . . . . . . . . . . . . . . . . . . . . . . . . . . 67
. . . . . . . Philodryas viridissimus, p. 503 66b. Dorsal scale rows 19 or more at
59b. Belly scales fewer than 205 . . . . . midbody . . . . . . . . . . . . . . . . . . . . . . . . . . 73
. . . . . . . . . . . . Philodryas olfersii, p. 503 67a (66). Body with longitudinal
60a (55). Upper body scales with a stripes, although may be subtle. . .
pair of small, subtle pits or depres- . . . . . Thamnodynastes pallidus, p. 508
sions (apical pits) near rear of scale 67b. Body without stripes . . . . . . . . . . . . 68
68a (67). Upper body scale rows ex- much narrower than those of
tremely oblique (Fig. 8F), as scales upper and lower rows . . . . . . . . . . . . . 77
of lateral rows (on side) much 75b. Body more slender; upper body
narrower than those of upper and scale rows not so oblique, as in
lower rows. . . . . . . . . . . . . . . . . . . . . . . most other snakes (Fig. 8F) . . . . . . . 76
. . . . . . . . . . . . Xenodon merremii, p. 510 76a (75). Scales beneath tail (subcau-
68b. Upper body scale rows not so dals) paired, one on each side . . . .
oblique, but as in most other see separate key for genus Oxyrhopus,
snakes (Fig. 8F) . . . . . . . . . . . . . . . . . . 69 p. 366
69a (68). Shallow longitudinal groove 76b. Subcaudals single, one wide scale
or depression (mental groove) pres- across underside of tail. . . . . . . . . . .
ent behind chin in middle of . . . . . . . . . . Pseudoboa neuwiedii, p. 504
underside of head (Fig. 8A) . . . . . . . 70 77a (75). Scales around midbody 17–
69b. Mental groove absent (Fig. 8A) 19 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
. . see separate key for genus Dipsas, 77b. Scales around midbody 21–23 . .
p. 365 . . . . . . . . . . . . . . Xenodon severus, p. 510
70a (69). Rearmost one or two teeth 78a (77). Teeth on maxillary bone
on maxillary bone (upper jaw) with fewer than 10, including 2 en-
venom groove . . . . . . . . . . . . . . . . . . . . 71 larged ones at rear (ungrooved)
70b. Rear maxillary teeth not grooved 72 . . . . . . . . . . Xenodon merremii, p. 510
71a (70). Scales beneath tail (subcau- 78b. Maxillary teeth 15 or more . . . .
dals) paired, one on each side . . . . . . . . . . Xenodon rabdocephalus, p. 510
. . . . . . . . . . . . . . . . . . Clelia clelia, p. 488 79a (74). Rearmost one or two teeth
71b. Subcaudals single, one wide scale on maxillary bone (upper jaw) with
across underside of tail. . . . . . . . . . . venom groove . . . . . . . . . . . . . . . . . . . . 80
. . . . . . . . . . Pseudoboa coronata, p. 504 79b. Rear maxillary teeth not grooved 85
72a (70). Belly scales, neck to vent, 80a (79). Uppermost longitudinal
more than 180 . . . . . . . . . . . . . . . . . . . row of body scales (vertebrals)
. . . . . . . . . . . Drymarchon corais, p. 492 distinctly larger than scales of
72b. Belly scales, neck to vent, fewer adjacent rows . . . . . . . . . . . . . . . . . .
than 180 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Siphlophis compressus, p. 507
. . . . . . . . . Drymoluber dichrous, p. 493 80b. Scales of vertebral row about the
73a (66). Frontmost scale atop tip of same size as those of adjacent rows 81
snout turned up into a sharp point 81a (80). Scales beneath tail (subcau-
. . . . . . . . . Phimophis guianensis, p. 503 dals) single, with one wide scale
73b. Not as 73a . . . . . . . . . . . . . . . . . . . . . . 74 across underside of tail. . . . . . . . . . .
74a (73). Two or more anterior tem- . . . . . . . . . . Pseudoboa neuwiedii, p. 504
poral scales (Fig. 8E), which are 81b. Most or all subcaudals paired,
behind and in contact with post- one on each side . . . . . . . . . . . . . . . . . . 82
oculars (behind and in contact with 82a (81). All head scales with light
eye) and between upper lip scales edges; sides of body pale yellow
(supralabials) and parietal (rear- with irregular vertical black marks;
most large plate atop head) . . . . . . . 79 subtle diffuse orange-red middor-
74b. Only one anterior temporal scale sal stripe present length of body;
on each side (Fig. 8E). . . . . . . . . . . . . 75 on lower jaw (mandible), tooth
75a (74). Body relatively heavy; upper numbers 3 through 5 (counted
body scale rows extremely oblique from front) very much enlarged . .
(Fig. 8F), as scales of lateral rows . . . . . . . . . . . Siphlophis cervinus, p. 507
82b. Coloration not as 82a; frontmost 89b. Dorsal scale rows 17 or fewer at
mandibular teeth only somewhat midbody . . . . . . . . . . . . . . . . . . . . . . . . . . 91
enlarged if at all, tooth length 90a (89). Upper body pattern with
decreasing toward rear of bone. . . . 83 colorful rings, including red,
83a (82). Belly scales (ventrals), neck around body; upper body scales
to vent, more than 200; body with a pair of small, subtle pits or
coloration not a series of dark depressions (apical pits) near rear
crossbands . . . . . . . Clelia clelia, p. 488 of scale (Fig. 8C), best seen with
83b. Ventrals fewer than 200, or if magnification . . . . . . . . . . . . . . . . . . . .
more, body coloration with dark . . Rhinobothryum lentiginosum, p. 506
crossbands that do not cross belly. 84 90b. Upper body coloration without
84a (83). Body coloration with dark colorful rings including red, but dark
crossbands that do not cross belly; brown bands present; apical pits
belly unmarked . . . . . see separate key absent . . . . . . Helicops angulatus, p. 494
for genus Oxyrhopus, p. 366 91a (89). Upper body scale rows 17
84b. Body coloration with dark stripes around midbody . . . . . . . . . . . . . . . . . . 92
or spots; belly with dark stripes . . 91b. Upper body scale rows 15 at mid-
. . . . . . . Thamnodynastes ramonriveroi, body. . . . . . . Leptophis ahaetulla, p. 496
p. 509 92a (91). Very elongate head with
85a (79). Upper body scales with a pointed snout . . . . . . . . . . . . . . . . . . . . . 93
pair of small, subtle pits or depres- 92b. Head not as 92a. . . . . . . . . . . . . . . .
sions (apical pits) near rear of scale . . . . . . . . . Drymobius rhombifer, p. 492
(Fig. 8C), best seen with magnifi- 93a (92). Upper body color mainly tan
cation . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86 or brown . . . . . Oxybelis aeneus, p. 500
85b. Apical pits absent . . . . . . . . . . . . . . 93b. Upper body color bright green .
. . . . . . Hydrodynastes bicinctus, p. 494 . . . . . . . . . . . . . Oxybelis fulgidus, p. 501
86a (85). All upper body scales 94a (88). Upper body (dorsal) scale
smooth, not keeled. . . . . . . . . . . . . . . rows around midbody 17 . . . . . . . . . 95
. . . . . . . Xenodon rabdocephalus, p. 510 94b. Dorsal scale rows at midbody
86b. At least uppermost body scales more than 17 . . . . . . . . . . . . . . . . . . . . . 96
keeled (with ridge) . . . . . . . . . . . . . . . . 87 95a (94). Snout relatively very long
87a (86). All upper body scales keeled and pointed; body with 3 conspic-
except in lowermost row; usually 3 uous dark longitudinal stripes, one
small scales contact rear edge of along vertebral area (atop body),
eye (postoculars); usually more one on each side . . . . . . . . . . . . . . . . .
than 210 belly scales, neck to vent . . . . . . . . Xenoxybelis argenteus, p. 511
. . . . . . . . . . . Pseustes sulphureus, p. 506 95b. Not as 95a . . . . . . . . . . . . . . . . . . . . .
87b. Scales of several lower body rows . . . . Dendrophidion dendrophis, p. 489
usually smooth, not keeled; usually 96a (94). All dorsal scales keeled (with
2 postoculars; usually fewer than ridge) except in lowermost row;
210 belly scales . . . . . . . . . . . . . . . . . . usually 3 small scales contact rear
. . . . . . . . . Pseustes poecilonotus, p. 505 edge of eye (postoculars); usually
88a (47). Vent covered by one scale as more than 210 belly scales, neck to
wide as belly scales in front of it. . . 94 vent . . . . . . Pseustes sulphureus, p. 506
88b. Anal scale divided into side-by- 96b. Scales of several lower rows of
side halves . . . . . . . . . . . . . . . . . . . . . . . . 89 dorsals usually smooth, not keeled;
89a (88). Upper body (dorsal) scale usually 2 postoculars; usually fewer
rows more than 17 around mid- than 210 belly scales . . . . . . . . . . . . .
body. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90 . . . . . . . . . Pseustes poecilonotus, p. 505
Key to Snakes of the Genus Atractus Key to Snakes of the Genus Chironius
of Guyana of Guyana
The following key is based largely on

The following key is based primarily
information provided by Peters & Orejas-
on information provided by Dixon et al.
Miranda (1970), Chippaux (1986), and (1993) and Kok (2010b).
Kok (2006a).
1a. Upper body (dorsal) scale rows 15 1a. Upper body (dorsal) scales 10
around midbody, excluding wide around midbody, excluding wide
belly scale . . . . . . . . . . . . . . . . . . . . . . . . . . 2 belly scale . . . . . . . . . . . . . . . . . . . . . . . . . 2
1b. Dorsal scales 17 around midbody . . 3 1b. Dorsal scales 12 at midbody . . . . . . . 4
2a (1). Upper body color pattern with 2a (1). Dorsal scales in longitudinal
3 or 4 longitudinal dark stripes; rows to each side of uppermost
wide belly scales, neck to vent, (vertebral) row with a keel (ridge;
125–150; paired scales of underside may be very subtle, but present)
of tail (subcaudals) fewer than 20 . . . . . . . . . . . . Chironius fuscus, p. 487
. . . . . . . . . . . Atractus trilineatus, p. 485 2b. Dorsal scales smooth . . . . . . . . . . . . . . 3
2b. Upper body pattern lacks well- 3a (2). No light crossbands on body;
defined dark stripes; wide belly loreal scale distinctly longer than
scales 153–163; subcaudals more high; small apical pits on scales
than 20 . . . Atractus tamessari, p. 484 on neck. . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . Chironius scurrulus, p. 488
3a (1). Paired scales of underside of
3b. Light crossbands on body; loreal
tail (subcaudals) more than 55;
about as high as long (perhaps
mental scale (foremost one on
slightly longer); no apical pits . . . .
lower jaw) in contact at rear with
. . . . . . . . . . Chironius challenger, p. 486
paired, central chinshields; belly
4a (1). Anal plate entire . . . . . . . . . . . . .
pattern of dark crossbands . . . . . . .
. . . . . . . . . . . . Chironius exoletus, p. 486
. . . . . . . . . . . . . . . Atractus favae, p. 483
4b. Anal plate divided . . . . . . . . . . . . . . . . . 5
3b. Subcaudals fewer than 55; mental
5a (4). Subcaudal scales (beneath tail)
scale separated from chinshields yellow and outlined in black or
by medial contact of first pair of dark brown; dorsal scales with
lower lip scales (infralabials) on light flecks or presence of a broad
each side. . . . . . . . . . . . . . . . . . . . . . . . . . 4 reddish brown stripe on each side
4a (3). Usually two upper lip scales on forward part of body . . . . . . . . .
(supralabials) contact preocular . . . . . . . . . . . Chironius carinatus, p. 485
(scale immediately in front of eye 5b. Subcaudals may be yellow, but not
on side of face) . . . . . . . . . . . . . . . . . . . . 5 completely outlined in black or
4b. Three supralabials contact preoc- dark brown; dorsal coloration not
ular. . . . . . . Atractus torquatus, p. 485 as in 5a . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
5a (4). Upper body color brown with 6a (5). Dorsal color uniform, without
black rectangular markings. . . . . . . distinct stripes or spots. . . . . . . . . . .
. . . . . . . . . . . . . . Atractus schach, p. 483 . . . . . . . . . . . . Chironius exoletus, p. 486
5b. Upper body unicolor and dark 6b. Dorsum brown or olive with light
(but broad dark stripes may appear vertebral stripe edged in darker
over time in preservative) . . . . . . . . brown or olive . . . . . . . . . . . . . . . . . . .
. . . . . . . . . Atractus steyermarki, p. 484 . . . . . . . . Chironius multiventris, p. 487
Key to Snakes of the Genus Dipsas Key to Snakes of the Genus Liophis
of Guyana of Guyana
The following key is based largely on The following key is based largely on
information provided by Peters & Orejas- Dixon (1989).
Miranda (1970) and MacCulloch & Lath-
rop (2004a). 1a. One head length in front of vent,
scale rows around body at least 2
1a. Dorsal (upper body) pattern of fewer than around midbody (be-
rounded, dark-brown or black tween head and vent), excluding
blotches or saddles with interspac- wide belly scale . . . . . . . . . . . . . . . . . . . . 2
es tawny brown . . . . . . . . . . . . . . . . . . . . 2 1b. Scale rows one head length in
1b. Not as 1a . . . . . . . . . . . . . . . . . . . . . . . . . 4 front of vent same count as around
2a (1). Dorsal scales 15 around mid- midbody . . . . . . Liophis reginae, p. 499
body, not counting wide belly scale 2a (1). Dorsal (upper) body scale
. . . . . . . . . . . . . . . . . Dipsas copei, p. 490 rows 17 around midbody . . . . . . . . . . 3
2b. Dorsal scales 13 at midbody . . . . . . . 3 2b. Dorsal scale rows 19 at midbody . . 8
3a (2). Blotches saddle-shaped, wider 3a (2). Upper lip scales (supralabials),
at vertebral row (atop body) than not counting rostral scale (at tip of
laterally (on sides) . . . . . . . . . . . . . . . upper snout), usually 7. . . . . . . . . . .
. . . . . . . . . . . . . . Dipsas pavonina, p. 491 . . . . . . . . . . . . . Liophis breviceps, p. 497
3b. Blotches narrower at vertebral row 3b. Supralabials 8 . . . . . . . . . . . . . . . . . . . . . 4
(may not meet there) than laterally 4a (3). Belly with checkered pattern of
. . . . . . . . . . . . . . . Dipsas catesbyi, p. 489 black and red or yellow . . . . . . . . . . . . 5
4a (1). Dorsal blotches triangular or 4b. Belly not checkered with black and
lozenge-shaped, usually widest at red or yellow, but occasionally
ventrals (edges of wide belly with black marks on sides of
scales), with yellow spot between ventrals (belly scales). . . . . . . . . . . . . . . 7
corners of blotches at ventrals . . 5a (4). Black stripe on side at rear of
. . . . . . . . . . . . . . . Dipsas indica, p. 490 body, extending onto tail. . . . . . . . .
4b. Not as 4a . . . . . . . . . . . . . . . . . . . . . . . . . 5 . . . . . . . . . . . . . . . Liophis reginae, p. 499
5a (4). Dorsal ground color of light 5b. Not as 5a . . . . . . . . . . . . . . . . . . . . . . . . . 6
browns and tans, with narrow 6a (5). Upper body with pattern of
blotches higher than wide, much bands, sometimes reticulations;
narrower than interspaces (at belly and underside of tail check-
least posteriorly or toward rear); ered with black and red or yellow,
upper lip scales (supralabials) 7– seen as dark bands or partial bands
10; chinshields (paired median in older preserved specimens . . . . .
scales of underside of head in . . . . . . . . . . . . . . . Liophis cobella, p. 497
front of wide belly scales) in 2 or 6b. Salt and pepper dorsal pattern,
3 pairs. . . . . . . . . . . . . . . . . . . . . . . . . . obscure in Amazon populations;
. . . . . . . . . . . . Dipsas variegata, p. 491 belly and underside of tail not as 6a
5b. Dorsal color pattern not as 5a; . . . . . . . . . . . . . . Liophis miliaris, p. 498
supralabials 6, last elongate; chin- 7a (4). Belly scales edged with black
shields in 4 pairs; from Mt. on sides. . . . . . Liophis miliaris, p. 498
Ayanganna . . . . . . . . . . . . . . . . . . . . . 7b. Belly scales unmarked on sides . .
. . . . . . . . . . . Dipsas pakaraima, p. 490 . . . . . . . . . . . . . . . Liophis reginae, p. 499
8a (2). One head length in front of black ring present around neck;
vent, scale rows around body only black rings not relatively narrow,
2 fewer than around midbody . . . . about as wide as red ones or wider
. . . . . . . . . . . . . . Liophis lineatus, p. 498 . . . . . . . . . . . . Micrurus psyches, p. 514
8b. One head length in front of vent,
scale rows around body 4 fewer
than around midbody . . . . . . . . . . . . . . 9 Key to Snakes of the Genus Oxyrhopus
9a (8). Upper body (dorsum) bright of Guyana
green . . . . . . . . . Liophis typhlus, p. 499
The following key is based primarily
9b. Dorsum usually brown to nearly on information provided by Bailey (in
black, not green . . . . . . . . . . . . . . . . . Peters & Orejas-Miranda 1970), Starace
. . . . . . . . . . Liophis poecilogyrus, p. 498 (1998), and Zaher & Caramaschi (1992).
1a. Snout black, but may have light
Key to Most Snakes of the Genus spots; preocular scale (in front of
Micrurus of Guyana and contacting eye) in contact with
frontal scale (large plate atop head
One of the species of coral snakes, between eyes); 10 scales along lower
Micrurus collaris, formerly placed in the lip (infralabials), not counting me-
genus Leptomicrurus, was included in the dian foremost scale (mental); adults
larger key to the snakes, above. The with bands on body, but older
following key is based primarily on infor- adults may be melanistic . . . . . . . . . . . . 2
mation provided by Campbell & Lamar 1b. Snout yellow or cream; supraocu-
(2004). lar scale (above eye) separates
1a. Black rings in triads (groups of preocular from frontal; 9 infrala-
black-white [or yellow]-black- bials; body reddish with cream
white-black) separated by red bands in juveniles, but bands ab-
bands, perhaps best visible on sent or obscure in adults . . . . . . . . .
belly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 . . . . . . . . . Oxyrhopus occipitalis, p. 502
1b. Black rings not in triads . . . . . . . . . . . 4 2a (1). Upper body (dorsal) black
2a (1). Vent covered by one scale as bands in triads (black-white-
wide as belly scales in front of it black-white-black, separated by
. . . . . . . . . Micrurus hemprichii, p. 513 red or yellow bands), at least
2b. Anal plate divided into side-by- toward rear of body . . . . . . . . . . . . .
side halves . . . . . . . . . . . . . . . . . . . . . . . . . 3 . . . . . . Oxyrhopus melanogenys, p. 501
3a (2). Head scales red but outlined 2b. Dorsal bands alternating red (may
with black . . . . . . . . . . . . . . . . . . . . . . . be orange or yellow) and black, not
. . . . . . . . Micrurus surinamensis, p. 515 in triads . . . . . . . . . . . . . . . . . . . . . . . . .
3b. Head scales not outlined with black . . . . . . . . . Oxyrhopus petolarius, p. 502
. . . . . . . . . . Micrurus lemniscatus, p. 514
4a (1). Red rings very wide; no black Key to the Testudinata (Turtles and
ring around neck right behind Tortoises) of Guyana
head; black rings on body relative-
ly very narrow . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . Micrurus averyi, p. 512 The characters used in the following key
4b. Red rings about as wide as black are based on information presented by
rings but red ones may be appar- Pritchard (1964), Ernst & Barbour (1989),
ently absent or reduced in size; and Duellman (2005).
1a. Arms and legs flattened as oar-like 7a (6). Webbing between toes; legs
flippers (sea turtles) . . . . . . . . . . . . . . . . 2 not elephantine or columnar . . . . . . . 8
1b. Not as 1a . . . . . . . . . . . . . . . . . . . . . . . . . 6 7b. Toes not webbed but stout; legs
2a (1). Top of shell covered with elephantine or columnar . . . . . . . . . . . 9
leathery skin and having 7 conspic- 8a (7). Red lines on head; 12 or 13
uous longitudinal ridges. . . . . . . . . . large scutes on lower shell. . . . . . . .
. . . . . . . . Dermochelys coriacea, p. 528 . . . . Rhinoclemmys punctularia, p. 528
2b. Top of shell bone-hard and lack- 8b. No red on head; large scutes on
ing 7 conspicuous longitudinal lower shell fewer than 12. . . . . . . . .
ridges. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 . . . . . . . Kinosternon scorpioides, p. 529
3a (2). Pleural scutes on upper shell 9a (7). Background color of upper shell
(scutes beside those along midline) brown; light spots on arms yellow or
4 on each side; cervical scute orange; viewed from above, sides of
(frontmost one on edge of upper upper shell straight in both sexes. .
shell) not in contact with first . . . . . . . . . Chelonoidis denticulata, p. 531
pleural on either side. . . . . . . . . . . . . . . 4 9b. Background color of upper shell
3b. Pleural scutes 5 or more on each more of a black than brown; light
side; cervical scute contacts first spots on arms red; viewed from
pleural on each side . . . . . . . . . . . . . . . . 5 above, sides of upper shell of males
4a (3). One pair of elongate prefrontal indented as if constricted by a large
scales atop head behind nasal rubber band . . . . . . . . . . . . . . . . . . . . .
scales and between eyes; large . . . . . . . Chelonoidis carbonaria, p. 530
plates of upper shell not overlap-
10a (6). A single, usually small, cen-
ping; rear edge of upper shell
tral, nuchal (¼cervical) scute on
smooth; head broader, beak
front edge of upper shell (Fig. 9) . . 12
rounded (viewed from above) . . . .
10b. No central nuchal scute on front
. . . . . . . . . . . . . . Chelonia mydas, p. 526
edge of upper shell (Fig. 9). . . . . . . . 11
4b. Two pairs of prefrontal scales; large
plates of upper shell overlapping; 11a (10). Upper shell relatively flat
rear edge of upper shell serrated; and widest posterior to (behind)
head narrower, beak bluntly point- center; usually two barbels on chin
ed (viewed from above) . . . . . . . . . . . near front of lower jaw; upper jaw
. . . . . . . . Eretmochelys imbricata, p. 527 rather squared off, not notched in
5a (3). Underside of bridge (connect- front; groove on head between eyes
ing upper and lower shells on sides) . . . . . . . . . . Podocnemis expansa, p. 529
with 3 inframarginal scales (on 11b. Upper shell relatively domed and
bridge beside lower shell); upper widest at midbody; usually one
shell somewhat elongate (viewed barbel on chin near front of lower
from above); head relatively large jaw; upper jaw distinctly notched;
. . . . . . . . . . . . . . . Caretta caretta, p. 526 no groove between eyes . . . . . . . . . .
5b. Underside of bridge with 4 infra- . . . . . . . . . . . Podocnemis unifilis, p. 530
marginal scales; upper shell nearly 12a (10). Bizarre turtle with enor-
round (viewed from above); head mous head of broadly triangular
relatively small. . . . . . . . . . . . . . . . . . . shape (viewed from above) and
. . . . . . . . . Lepidochelys olivacea, p. 527 flattened; snout long, narrow, snor-
6a (1). To hide head, neck is pulled kel-like; upper shell with 3 con-
straight back into shell . . . . . . . . . . . . . 7 spicuous lengthwise knobby keels
6b. To hide head, neck is bent side- . . . . . . . . . . . . . Chelus fimbriatus, p. 524
ways to lie along edge of shell. . . . . 10 12b. Head, snout, and shell not as 12a . . 13
13a (12). Upper shell with broad, them for swimming. Adults are tailless.
lengthwise, round-bottomed, ver- About 15 families, 46 genera, and 137
tebral groove on middle of back species of frogs and toads are known to
. . . . . . Platemys platycephala, p. 525 occur in Guyana.
13b. Upper shell without vertebral
groove . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14a (13). Chin and throat yellow with Allophrynidae
black streaks; dark line on side of
head, passing through eye . . . . . . . . We follow Guayasamin et al. (2009) in
. . . . . . . . . . . Phrynops tuberosus, p. 524 recognizing this family and the Allocen-
14b. Throat gray to pale yellow; no troleniae, an unranked taxon above the
dark line through eye . . . . . . . . . . . . family level that contains two families, the
. . . . . . . . . . Mesoclemmys gibba, p. 524 Allophrynidae and Centrolenidae. Their
research agrees with several recent papers
involving molecular phylogeny reconstruc-
Species Accounts: Amphibia tion (e.g., Faivovich et al. 2005, Frost et al.
The approximately 6700 species of 2006) showing clearly that Allophryne
modern amphibians (Hoffmann et al. ruthveni is the sister group of what
2010) are remarkably diverse in body traditionally had been called the Centrole-
form, coloration (some being spectacularly nidae exclusive of the monotypic Allo-
beautiful), camouflage, life history, behav- phryne. Consequently, Frost et al. (2006)
ior, toxicity, actual and potential use to combined the Allophrynidae with the
humans, and so-on. There are three major Centrolenidae, which would eliminate a
groups, as follows: Anura (frogs and monotypic family (the former) and em-
toads); Caudata (salamanders, of which phasize the sister group relationships of
none occurs in Guyana); and Gymnophio- these organisms. However, glass frog
na (the legless caecilians). The earliest specialists prefer retaining the two tradi-
origin(s) of amphibians from fish and the tional family names and showing their
evolutionary relationships of the modern close relationship by naming a new and
species have received extensive modern unranked Allocentroleniae. What is im-
study (e.g., Frost et al. 2006, Anderson et portant in the science involved is for the
al. 2008, Pyron & Wiens 2011), but there is classification to most clearly reflect evolu-
not general agreement on their classifica- tionary relationships of monophyletic
tion yet, and a great deal of work remains groups that are strongly supported by
to be done. The on-line web reference by evidence, rather than have a classification
Frost (2011) is an exceedingly important that obscures or is inconsistent with
resource for accessing the taxonomic relationships. One might still question
literature on all species of amphibians of whether the classification system of Frost
the world. Appendix 2 provides a complete et al. (2006) or Guayasamin et al. (2009)
list of the species we have documented as best accomplishes this for these frogs.
occurring in Guyana. These animals can be difficult to find,
even when males are calling. They are
small and most often found on low
Species Accounts: Amphibia: Anura vegetation at night, usually in primary
(Frogs and Toads) forest ‘‘in the close vicinity of streams’’
(Kok & Kalamandeen 2008:111), often
The anurans have the legs modified for while it is raining. ‘‘Eggs are deposited on
jumping, although the aquatic species use a leaf overhanging water, from which
Fig. 9. Turtle shells; nuchal scute (shaded) either present on front edge of upper shell as in Chelus (below),
or absent as in Podocnemis (above). Scale bars: 1 cm.
tadpoles will fall into the water as they species, after hatching, a parent carries the
hatch’’ (Kok & Kalamandeen 2008:112). tadpoles to water.
Allophryne ruthveni Gaige, 1926 Allobates femoralis (Boulenger, 1884)

Plate 1A, B Plate 1C
Type material.—The holotype is Type material.—Originally named Pros-

UMMZ 63419, adult female, from Tukeit therapis femoralis based on two specimens,
Hill below Kaieteur Falls, Guyana. a male and female, from Yurimaguas,
Distribution.—The Guiana Shield re- Huallaga River, northern Peru. BMNH
gion, including Venezuela, Guyana, 1947.2.14.21 was designated as the lecto-
French Guiana, Surinam, and Brazil type (Silverstone 1976).
(Frost 2011, Lescure & Marty 2000). Distribution.—Occurs widely in the Gui-
Vouchers for Guyana.—The holotype; ana Shield Region in eastern Venezuela,
and USNM 531291 from Iwokrama Forest Guyana, French Guiana, Surinam, and
Brazil; also in the Amazon drainage of
Reserve, 5 hr downstream of Kurupukari
Brazil, Colombia, Ecuador, Peru, and
Base Camp on the Essequibo River,
Bolivia (Frost 2011).
Kabocali Camp.
Vouchers for Guyana.—AMNH A-
Coloration in life.—The following quo-
39679–39680, AMNH A-39687, AMNH
tations are from Kok & Kalamandeen
A-39708, AMNH A-70991, AMNH A-
(2008:112), for specimens from Kaieteur
70993, AMNH A-70994, and USNM
National Park. ‘‘Dorsal ground colour and
118075 from Kartabo; USNM 531354–
pattern variable, ranging from greyish- 531355 from Iwokrama Forest Reserve, 13
brown to creamish bronze with dark km S of Kurupukari Base Camp on George-
irregular spots and/or reticulum; often a town-Lethem Road; USNM 535737 from
conspicuous cream spot on posterior face of Baramita, on trail S to Baramita River; and
upper arm;...Ventral surface ...translucent USNM 566054 from the Marudi Moun-
dark grey, with a whitish central area visible tains, about 24 km SE Aishalton.
through the skin in males...Throat black Coloration in life.—The following is
with white spots...Iris dark reddish brown.’’ based on USNM 566054 (CJC field notes,
Usually there are white or cream markings 1993): dorsum black; dorsolateral light
on the sides also (Lescure & Marty 2000). stripe pale metallic tan (very pale copper);
lateral light stripe of similar color anterior
to arm, very pale green posterior to arm;
Aromobatidae arms brown; orange spot in armpit, with
yellow edging; femoral stripe orange with
We follow Grant et al. (2006) in recog-
yellow center, within a black field; other-
nizing this family, which is diagnosed solely
wise, legs brown with black spots; chin,
on DNA sequence data. In the recent past,
throat, and chest black; beneath arms,
Guyanan species in this family have been
abdomen, and legs gray with irregular
assigned to the genera Colostethus, Den-
black markings.
drobates, or Epipedobates, but recently,
generic assignments have been changed.
These are rather small, inconspicuous frogs Allobates spumaponens Kok & Ernst, 2007
that usually occur on the ground or in low Plate 1D
vegetation, such as terrestrial bromeliads.
Eggs are laid hidden in moist situations on Type material.—SMNS 12511, an adult
the ground or in vegetation, and for some male, from Mabura Hill Forest Reserve,
Upper Demerara-Berbice Region, Guyana did not note the partial oblique pale lateral
(5809 0 N, 58841 0 W, elevation ca. 100 m). stripe as mentioned by Kok & Ernst
Distribution.—Known only from central (2007), although this is subtle (not very
Guyana. evident in their photos).
Vouchers for Guyana.—These include Comments.—At times in the recent past,
the holotype and paratopotypes (SMNS this species was referred to as Colostethus
12510, and 12512–12516); also AMNH A- brunneus.
166470–166475 from Berbice River Camp
and AMNH A-166498 from Konawaruk
Camp. Anomaloglossus beebei (Noble, 1923)
Coloration in life.—‘‘Dorsal ground col- Plate 1E
our medium brown with several dark
brown flecks on dorsum; dorsal surface Type material.—Originally named Hy-
of head slightly darker than dorsum. Wide loxalus beebei based on the holotype
lateral black band from tip of snout to AMNH A-18683, an adult female from
vent, containing two-thirds of tympanum near Kaieteur Falls, Guyana.
and not tapering posteriorly from axilla. Distribution.—Known only from the
Pale, narrow, diffuse dorsolateral stripe Pakaraima region of western Guyana.
above the band, extending from tip of Vouchers for Guyana.—The holotype;
snout to vent. Diffuse, pale, partial oblique AMNH A-166132–166134 from the vicin-
lateral stripe imbedded in the black band, ity of Kaieteur Falls; IRSNB 13721–13726
extending from groin to about one-quarter and 13728–13753 from Kaieteur National
of the distance to arm insertion. Flanks Park; ROM 42994 and 43178–43197 from
white with a few irregular brownish grey the Kaieteur plateau; ROM 39629–39636,
blotches; ventrolateral stripe present. Up- and 42388 from Mount Ayanganna Pla-
per lip white, suffused with barely visible teau; and USNM 337769–337771 from the
tiny melanophores. Throat hyaline white Kaieteur Savanna, Potaro River.
with barely visible tiny melanophores Coloration in life.—Adult coloration
laterally and on the chin. Belly immaculate varies, with one of two ground colors;
hyaline white. Upper surfaces of thighs bright yellow to orange or pale brown.
and shanks brown with dark brown ‘‘Cream dorsolateral stripes and dark
blotches, no crossbars discernible. Pale brown markings are present or absent,
paracloacal mark present. Pale yellow spot producing five color patterns, described in
on groin. Forelimbs light brown with a detail by Bourne (2001). In juveniles the
dark brown line posteriorly from elbow to background color is green by night and
wrist and a short dark brown line anteri- yellow-green by day, slightly darker than
orly discontinuous from arm insertion to adult coloration. Palpebral membrane is
elbow. Toes greyish with some light grey unpigmented, with no dark line along its
blotches. Fingers light brown with some upper edge’’ (Kok et al. 2006a:58). ‘‘Ven-
pale blotches. Palm and sole dark brown. tral surface...immaculate yellow to yellow-
Iris golden bronze’’ (Kok & Ernst 2007: ish orange in both sexes’’ (Kok &
27). Kalamandeen 2008:116). In preservative
Specimens in the AMNH series from the the background color fades to pale dull
Berbice River Camp noted by CJC in life yellow and the bright cream dorsolateral
(1997 field notes) had a translucent yellow lines become white. Tadpoles are pale
venter (none white as Kok & Ernst [2007] yellow with variable dark mottling.
noted for males, although translucent gray Comments.—See Kok et al. (2006a) for a
beneath arms and legs), with either the detailed redescription of this species and
throat or abdomen of brighter color. CJC comments on past misuse of the name.
Also, MacCulloch & Lathrop (2009) dis- upper lip. Chin same colour as upper lip;
cuss the specimens from Mount Ayangan- throat and chest medium pink, belly cream
na. Details of habitat, diet, vocalization, stippled with melanophores; underside of
reproductive behavior and parental care thigh and shank creamy yellow. Iris
are in Bourne et al. (2001), including a metallic reddish bronze. Upper arm and
report that adult females will return to forearm orange, posterior edge of the
bromeliad leaf-axils containing water with latter with a black longitudinal stripe
their own tadpoles and deposit unfertilized tapering from wrist to elbow. Dark spot
eggs upon which their tadpoles will feed. on upper surface of wrist. Dorsal surfaces
Details of predator avoidance and forag- of thigh, shank and tarsus same colour as
ing behavior are in Bourne et al. (2001) dorsum, with ill-defined black crossbars,
also. more crossbars on tarsus. Pale paracloacal
mark. Toes and digits with small light blue
dots. Palms and soles black.’’
Anomaloglossus kaiei (Kok, Sambhu, Coloration of the AMNH series from
Roopsind, Lenglet, & Bourne, 2006b) Konawaruk Camp was noted by CJC (field
Plate 1F notes, 1998) as follows: brown above (all
dorsal surfaces), with very dark brown to
Type material.—Originally named Co- black sides; all ventral surfaces yellow
lostethus kaiei, based on the holotype (abdomen translucent); anal area dark
IRSNB 1938, adult male from Kaieteur brown to black, flanked by light tan line
National Park, Guyana, along Tukeit trail (slanted off vertical); some specimens with
(5811 0 06 00 N, 59828 0 51 00 W, elevation ca. 400 two or a few dark brown bands on legs.
m). Comments.—Details of habitat, diet,
Distribution.—Pakaraima region, west- vocalization, tadpoles, and reproductive
ern Guyana. behavior are in Kok et al. (2006b).
Vouchers for Guyana.—These include
the holotype and paratypes (IRSNB
1939–1964), all from Kaieteur National Anomaloglossus megacephalus
Park (Kok et al. 2006b:38–39); ROM Kok, MacCulloch, Lathrop,
42999–43000, also from Kaieteur National Willaert, & Bossuyt, 2010
Park; AMNH A-166483–166497 from Plate 2A
Konawaruk Camp; IRSNB 14922–14924
from Wayalayeng; IRSNB 14925–14931 Type material.—The holotype is IRSNB
from Maringma Tepui; and ROM 43221, 1986, adult female from the eastern base of
43327, 43330, and 43333 from Mount Mount Maringma, Cuyuni-Mazaruni Dis-
Wokomung. trict, Guyana (05812 0 37 00 N, 060833 0 59 00 W,
Coloration in life.—The holotype was 1060 m elevation).
described by Kok et al. (2006b:43–44) as Distribution.—Only three specimens are
follows: ‘‘Dorsal ground colour medium known, all from the Pakaraima region of
brown. A wide black band extending from Guyana.
tip of snout laterally around the body and Vouchers for Guyana.—The holotype
above the vent, containing most of the (see above) and two paratypes, ROM
tympanum and tapering posteriorly from 39637–39638 from Mount Ayanganna
axilla; a thin, white, partly broken dorso- (05824 0 N, 59857 0 W, 1490 m elevation).
lateral stripe above this band, extending Coloration in life.—‘‘Dorsal ground
from posterior edge of orbit to vent. Upper colour medium brown, with a dark brown
lip bar dirty white with a few white and V-shaped interorbital bar slightly outlined
light blue dots. Flank same colour as with light creamish brown, followed by
two less defined dark brown bars, the first representing A. tepuyensis, with the pre-
more or less V-shaped, at the level of arm diction that additional work would dis-
insertion, the second just anterior to cover that it was an undescribed species
sacrum. Small, poorly defined, dark (Grant et al. 2006:120). Based on that
brown flecks on snout, between and on tentative identification, tadpoles from
dorsal bars. Upper surface of arm light Mount Ayanganna were described under
brown with well-defined dark brown the name A. tepuyensis by MacCulloch &
transverse bands on forearm and wrist, a Lathrop (2009:11).
less defined one on elbow; upper surface
of thigh, shank, and foot light brown with
well-defined dark brown transverse Anomaloglossus praderioi
bands. A yellow wash on the anterior (La Marca, 1997)
face of the thigh. Flanks dark brown,
slightly lighter ventrally with a few bluish Type material.—Originally named Co-
irregular spots and a few whitish irregular lostethus praderioi based on the holotype,
blotches on the lower part forming a UALBG 4196, adult male from Mount
broken inconspicuous ventrolateral Roraima (5810 0 N, 60847 0 W; 1950 m),
stripe. A few white spots (covering small Estado Bolı́var, Venezuela.
tubercles) form a broken oblique lateral Distribution.—This species is known
stripe that does not extend to arm from only two localities in southeastern
insertion [but on some specimens the Venezuela and one in western Guyana
stripe is not broken]. No dorsolateral (Kok 2010a).
stripe. Upper lip creamish brown suffused Vouchers for Guyana.—IRSNB 11403–
with melanophores, two bluish marks 11413 ‘‘from the southeastern slope of
below eye; loreal region and side of head Maringma Tepui (05812 0 16 00 N, 060834 0
dark brown; tympanum area light cream- 39 00 W, 1376 m elevation...), Cuyuni-
ish brown concealed in a poorly defined Mazaruni District, Guyana’’ (Kok
light stripe from posterior corner of eye to 2010a:52). Considering that the type local-
arm insertion. A dark brown stripe is ity is on the Venezuelan part of Mount
present on the anterior edge of upper arm, Roraima and that similar habitats occur
tapering from arm insertion to forearm. on the Guyanan parts, we are certain that
Throat light brown, blotched with white this species occurs here as well, but
and dark brown; belly light grey, blotched voucher specimens are needed.
with dark brown, blotches more numer- Coloration in life.—‘‘Dorsal ground
ous and larger laterally. Undersurface of color varies from light grayish brown to
upper arm light brown with a few dark reddish brown or dark brown, usually
markings; undersurface of forearm black- with one to three dark brown to black
ish; undersurface of thigh and shank light triangular, diamond-shaped or diffuse
grey blotched with dark brown; rear of hourglass markings from interorbital to
thigh and cloacal region blackish. Pale presacral region. Upper surface of arm
reddish orange paracloacal marks pre- light brown to yellowish or orangish
sent. Tip of digits whitish. Palms and soles brown; upper surfaces of leg light grayish
black. Iris mostly orange-bronze, dark- brown to dark gray with dark brown to
ened with black suffusion, with two black transverse bands on thigh, shank
oblique metallic white marks separated and foot; in most living specimens one of
by a black triangular mark below pupil’’ the transverse bands is larger and more
(Kok et al. 2010:23). conspicuous than others, which can some-
Comments.—Previously, a specimen of times be absent or barely visible... Adult
the type series was reported tentatively as males with gray to very dark gray throat,
almost solid black, with black blotches, Anomaloglossus roraima

throat color extending onto chest and (La Marca, 1997)
anterior part of belly...; posterior part of
belly bright orange... Adult female with Type material.—Originally named Co-
bright orange throat, almost immaculate lostethus roraima, based on the holotype
except a few lighter blotches and a few ULABG 4197, a juvenile female from ‘‘El
melanophores on chin and lower lip; belly Paso de La Muerte, a unos 60–70 m antes
bright orange... Flanks light gray to light de la cumbre del tepuy, en el camino, ‘La
reddish brown (males) or yellowish brown Rampa,’ que conduce desde su base hasta
(female), usually with some small white or cima del Monte Roraima, Estado Bolı́var,
sky blue flecks more concentrated on the Venezuela.’’
lower part, not forming a straight ventro- Distribution.—This species is known
lateral stripe... However, some inconspic- only from Mount Roraima, Guyana and
uous yellowish elongate spots may occur Venezuela.
and can be interpreted as a broken Vouchers for Guyana.—CPI 10212–
stripe... Distinct thin pale dorsolateral 10217, from ‘‘Mt. Roraima, 1860–2350
stripe present in both sexes,...extending m’’ (Grant et al. 2006:242), which Grant
from tip of snout to tip of urostyle, et al. stated (p. 45) would be deposited at
usually slightly narrower on body than the USNM.
on head... A wide dark brown band of Coloration in life.—In the original de-
variable width present below the dorso- scription, La Marca (1997 [1996]) stated
lateral stripe, extending from tip of snout (as translated from Spanish) that photo-
laterally around the body and above the graphs of the holotype show grayish dorsal
vent, containing most of the tympanum tones and light brown ventral surfaces.
and usually tapering posteriorly from There were no gaudy colors or yellow or
axilla. Upper lip light to dark gray, reddish tones. Colors and pattern of the
suffused with melanophores and some- holotype in alcohol was also given in the
times with a few sky blue flecks in males, original description (p. 41).
yellow with few melanophores in female. Comments.—This species was described
A black stripe extends from posterior from a single juvenile, but the CPI
margin of eye to axilla region and is specimens collected recently in Guyana
usually underlined by a white to light gray (see above) confirm its validity.
(males) or bright yellow (females) line; a
white or light gray (males) or yellow spot
(female) at arm insertion. Usually a black Bufonidae
stripe on anterior and posterior edges of
forearm, tapering from arm insertion to This family, which includes the terres-
forearm; dark area on upper surface of trial, brown and warty anurans most often
wrist. Undersurfaces of arm and thigh referred to as toads, was named about 185
bright to dark orange, undersurface of yr ago (Gray 1825) and its composition
shank yellowish brown with black flecks has changed over many decades. Never-
and spots (more extensive in males). Pale, theless, the family in its modern sense
creamish brown, paracloacal marks pres- received strong support in recent molecu-
ent. Toes and digits with sky blue dots... lar phylogenetic analyses (e.g., Frost et al.
Palms and soles dark brown to black. Iris 2006, Pramuk 2006, Chaparro et al. 2007),
metallic reddish bronze in males, metallic although assignment of generic names in
bronze in female, with fine dark brown this large and complex family is still
reticulation in both sexes’’ (Kok shifting as more and more is learned about
2010a:57–58). relationships and synonymies. The family
probably originated in South America (irregular circles, ellipses, reticulations);

(Pramuk et al. 2008). throat, chest, and anterior abdomen
Most species of bufonids live on the orange; remaining ventral surfaces yellow
ground, whether in savanna or forest, and or pale orange, although pink with black
many of them produce noxious secretions spots in a small male. Kok & Kala-
from their parotoid glands (on back of the mandeen (2008:122) add that the dorsal
head and/or neck) when disturbed. The ground color is dark brown on some
species of one genus, however, are special individuals and the light markings may
in several respects. These are the five have dark brown or black spots within
Guyanan species of Oreophrynella, a re- them.
markable genus of montane species the Comments.—The frogs currently cov-
total global distribution of which includes ered by this name may be a complex of
western Guyana, southeastern Venezuela, cryptic species. They occur widely across
and northern Brazil. These are upland and Amazonia and have a spotty distribution
highland species that occur at the highest and low vagility, which promotes local
elevations, are small, have opposable toes, differentiation. A thorough analysis of
and some of which are found on vegetation geographic variation is needed, following
up to 3 m above the ground. They are up on the recent work analyzing DNA
among the least known frogs in the world. sequences provided by Noonan & Gaucher
Of the five species of Oreophrynella that (2005), which suggests that several taxa
occur in Guyana, four are known from a may be masquerading under this name.
total of 10 or fewer specimens (two species These authors and others (e.g., Lescure &
are known from only one or two speci- Marty 2000, Kok & Kalamandeen 2008)
mens), and two species are endemic to have used various names, including Atelo-
Guyana. pus spumarius hoogmoedi, Atelopus hoog-
moedi, and Atelopus spumarius barbotini,
for different populations of these frogs
Atelopus spumarius Cope, 1871 from the Guianas, but we consider it to be
Plate 2B premature to apply such names until
additional studies are completed. See also
Type material.—The neotype is MNHN comments by Frost (2011). If in the future
1979–8382, from ‘‘Colonia, bassin de la it turns out that the name A. barbotini
rivière Ampiyacu (Dpto Loreto, Pérou)’’ should be applied to some of the frogs we
(Lescure 1981:894). listed above (Vouchers), that name might
Distribution.—‘‘Disjunct populations in apply to the series from Konawaruk
Amazon lowlands of Ecuador and eastern Camp, a matter that needs additional
Peru, to Amazonas, Pará, Amapá (Brazil), investigation.
and the Guianas’’ (Frost 2011). Daly et al. (1994) reported that the
Vouchers for Guyana.—AMNH A- major component of the toxins in the skin
13528–13529, AMNH A-21332–21333, of A. spumarius sampled in Brazil was the
and AMNH A-53304–53305 from Karta- powerful tetrodotoxin.
bo; AMNH A-70997 from Kalacoon; and
AMNH A-166066–166077 and USNM
566005–566010 from Konawaruk Camp. Dendrophryniscus minutus (Melin, 1941)
Coloration in life.—The following is Plate 2C, D
based on the AMNH and USNM speci-
mens from Konawaruk Camp (CJC field Type material.—Originally named Ate-
notes, 1998). All dorsal surfaces black with lopus minutus based on specimens from
bright orange, gold, or red markings Taracuá, Rio Uaupés, Brazil. The website
of the Göteborgs Naturhistoriska Museum 46409–46413, and 46417–46418 from

(www.gnm.se) lists the holotype as Ba.Ex. Mount Wokomung; and IRSNB 1981.
462. Coloration in life.—‘‘Dorsum, top of
Distribution.—Amazonian Colombia head, flanks and limbs uniformly brown-
(from the llanos of Meta south to Ama- ish-orange. Throat, abdomen and ventral
zonia), Ecuador, Peru, Bolivia, Brazil, and surfaces of limbs creamy yellow. Tip of
southern Guianas (Frost 2011). snout, eyes and anterior flanks lighter
Vouchers for Guyana.—AMNH A- shade of yellow. Flanks and upper por-
166120–166125, 166130–166131 and tions of limbs covered in medium sized,
USNM 566041–566044 from Berbice Riv- yellow glandular spots. A loosely orga-
er Camp; USNM 200507 from 24-mile nized line of these spots extends from
Forest Reserve south of Bartica; and scapular region to groin. Posterior aspect
USNM 531329–531353 from several sites of thighs and anal sheath with similar
within Iwokrama Forest Reserve. yellow glandular spots. Upper lip creamy
Coloration in life.—The following is yellow with diffuse brown mottling. Iris
based on the AMNH and USNM series silvery orange with black reticulations’’
from Berbice River Camp (CJC field notes, (Lathrop & MacCulloch 2007:90).
1997). Dorsum reddish brown or dark Comments.—These are small toads,
brown, with or without gray patches that males not exceeding a body size of 27
may be in a reticulum, at times also mm, females 38 mm. ‘‘All specimens were
numerous tiny gray or white dots; tan or found on broad-leaf vegetation at a height
light gray vertebral light line present or of 1.5 m or more...[on] very humid
not; throat and chest black on some, evenings in which it had either rained all
reddish brown on others; belly white with day, or was in the early stages of a
black spots; beneath arms and legs may be downpour’’ (Lathrop & MacCulloch
translucent gray with dark gray or black 2007:91). Additional research is warranted
spots; iris black with narrow copper inner to clarify the differences between this
ring. species and O. macconnelli (Lathrop &
Comments.—These are small toads with MacCulloch 2007, Kok 2009a).
a maximum snout-vent length of about 24
mm (Lescure & Marty 2000) and a body
Oreophrynella macconnelli
shape somewhat similar to that of Atelopus.
Boulenger, 1900
Type material.—The holotype is BMNH
Oreophrynella dendronastes 1947.2.14.49 (formerly 99.3.25.17), from
Lathrop & MacCulloch, 2007 the base of Mt. Roraima, 3500 ft, reported
Plate 2E, F originally as being from Guyana. However
the locality is probably in Venezuela
Type material.—The holotype is ROM (Phelps 1938). See below (Vouchers) for
39645, an adult female from the northeast definite records in Guyana.
slope of Mount Ayanganna, District 7, Distribution.—This species is known
Guyana, 1490 m (5824.11 0 N, 59857.41 0 W). only from Mt. Roraima and Maringma
Distribution.—Known only from Guya- tepui, Guyana, on the Guyana-Brazil bor-
na, in the immediate vicinity of the type der (Kok 2009a), but it probably occurs in
locality. adjacent Brazil and Venezuela also.
Vouchers for Guyana.—The holotype; Vouchers for Guyana.—BMNH 1976.700
ROM 39648–39655 from Mount Ayan- and 1976.702 from northern slopes of
ganna; ROM 46398–46399, 46401–46402, Mount Roraima, 1460 m elevation; IRSNB
14333–14336 from ‘‘Maringma tepui, Vouchers for Guyana.—USNM 118230

southeast slope, 1376 m asl (05812 0 16 00 N, is from Mount Roraima within the Guya-
060834 0 39 00 W)’’ (Kok 2009a:49); USNM na portion. In addition, Señaris et al.
549306 from the north slope of Mount (2005:67) included the following: ‘‘GUYA-
Roraima, 058160 N, 60844 0 W, at 1300 m NA-BRAZIL: Wei-Assipu-tepui,
elevation. (5813 0 1 00 N, 60842 0 19 00 W) 2400 m a.s.l.:
Coloration in life.—‘‘Adult dorsal colour MHNLS 15912.’’
is variable...ranging from light brownish Coloration in life.—Without specifying
orange...to olive brown with darker spot- whether the color notes were taken in life or
ting...Olive brown specimens have distinct not, Boulenger (1895:521) stated the fol-
white spots on the lower flank, while in the lowing: ‘‘black; throat and belly spotted or
orange specimen these spots are yellowish marbled with bright yellow.’’ Kok (2009a:
and less conspicuous. All adult specimens 39) stated that the belly in life is ‘‘bright
with a series of yellow...to brownish orange with black mottling in O. quelchii.’’
orange...spots forming a dorsolateral line, Comments.—This species is very poorly
and series of similar spots on axilla, upper known.
arm, groin and thigh forming short longi-
tudinal stripes on limbs. All adult speci-
Oreophrynella seegobini Kok, 2009a
mens have a white to yellow spot below the
eye, although...inconspicuous [on some]. Type material.—The holotype is IRSNB
Adult ventral colour is creamy yellow with 1979, an adult male from the summit
brown marbling; palm and sole are plateau of Mount Maringma, Cuyuni-
orange’’ (Kok 2009a:43–44). Mazaruni District, Guyana (05812 0 59 00 N,
Comments.—Individuals on Maringma 060835 0 05 00 W, 2088 m elevation).
Tepui were all ‘‘found active by night, Distribution.—Known only from the
walking on...epiphytic ferns...about 1.0– type locality.
3.0 m above the ground, except...[one] Vouchers for Guyana.—The holotype
collected on the ground at 17h00’’ (Kok and a paratopotype, IRSNB 1980.
2009a:44–45). Oreophrynella macconnelli is Coloration in life.—‘‘Dorsum, top of
more poorly known than O. quelchii. head and flanks blackish brown, lower
arm, hindlimbs and top of hands and feet
slightly lighter than dorsum. A bright
Oreophrynella quelchii (Boulenger, 1895) yellowish orange spot at arm insertion on
the left side, lacking on the right side where
Type material.—Originally named Oreo- the area is dark brownish orange. Upper
phryne Quelchii based on several specimens eyelids and lateral surfaces of snout dark
from the summit of Mount Roraima, at brownish orange. Throat, chest and ante-
8500 ft elevation. Apparently the type rior part of belly dark brownish orange,
specimen was collected on the Venezuelan posterior part of belly and area below vent
part of the summit (Phelps 1938). The blackish brown. A black mid-ventral line
syntypes include BMNH 95.4.19.1–5, running from tip of lower jaw to posterior
99.3.25.7–13, KU 126081–82 (formerly part of belly, slightly curved sinistrally.
BMNH 99.3.25.14–15), and MCZ 3500– Ventral surface of limbs brownish orange.
02 (Frost 2011). See below (Vouchers) for Palms and soles orange...Iris dark greenish
records from Guyana. brown with inconspicuous dark grey
Distribution.—‘‘Cerro Roraima, Vene- streaks’’ (Kok 2009a:40).
zuela and Guyana, presumably into Brazil Comments.—This is one of the least
on this same massif’’ (Frost 2011). known species in the world, with only two
specimens known to science. One was Vouchers for Guyana.—AMNH A-

found in late afternoon walking on muddy 136027 from Karanambo; AMNH A-
ground, the other hiding beneath a rotting 141058–141060, AMNH 141063–141064,
bromeliad leaf (Kok 2009a). and USNM 497688 and 566020–566021
from Dubulay Ranch; AMNH A-166084
and A-166087 and USNM 566022–566023
Oreophrynella weiassipuensis Señaris, from Berbice River Camp; and AMNH A-
166089–166093 and USNM 566024–
DoNascimiento, & Villarreal, 2005
566025 from Konawaruk Camp. Also
Type material.—The holotype is USNM 85013 from Pickersgill, Pomeroon
MHNLS #15913 from Wei-Assipu-tepui, River (07 816 0 N, 58843 0 W) ; USNM
Sima de los Guácharos, 280 m north of the 118068–118070 from Kartabo; USNM
heliport, 2280 m a.s.l. (coordinates of the 291103 from Kato, Chiung River; and
heliport 5813 0 1 00 N, 60842 0 19 00 W), Brazil- USNM 531292–531298 from several sites
Guyana border. within Iwokrama Forest Reserve.
Distribution.—Known only from the Coloration in life.—The following is
type locality. based on the AMNH vouchers listed
Vouchers for Guyana.—The holotype above from Karanambo and Dubulay
from the Brazil-Guyana border is the only Ranch, paraphrased from CJC’s field
known specimen. notes (1992, 1994): Tan anterior to paro-
Coloration in life.—‘‘Dorsum reddish toid glands, with or without a few dots of
rust; posterior body similar but warts are
brown with scattered darker brown marks
rust and there may be a wash of rust
and a fine dark mid-dorsal line; flanks dark
posteriorly; broad reddish brown lateral
brown; dorsum of hands light brown with
area from tip of snout nearly to groin;
minute reddish brown spots;...throat and
arms and legs dark brown or gray; ventral
chest reddish brown; belly slightly darker
surfaces gray with cream spots; broad rust
than the remaining ventral surface; fingers
wash may be present on posterior belly,
and toes light brown ventrally’’ (Señaris et groin, and anal region, but on some
al. 2005:62). individuals ventrally across thighs and
Comments.—This is one of the most hips is very pale tan instead. These notes
poorly known animals in the world, being compare well with those of Kok & Kala-
known to science by only a single speci- mandeen (2008:126).
men. The frog ‘‘was found in a forested
area at the bottom of a large pit’’ (Señaris
et al. 2005:66).
Rhaebo nasicus (Werner, 1903)
Plate 3B
Rhaebo guttatus (Schneider, 1799) Type material.—Originally named Bufo

Plate 3A nasicus. The holotype is IRSNB 1.015 and
the type locality was restricted to South
Type material.—Originally named Bufo America, probably along the Atlantic
guttatus with the apparent holotype ZMB drainage by Smith & Laurent (1950).
3517 (Frost 2011). The type locality was Distribution.—The Pakaraima region of
restricted to Surinam (Rivero 1961). western Guyana and southeastern Vene-
Distribution.—Occurs widely through- zuela, 100-1410 m elevation (Hoogmoed
out the Guianan Region and in Amazo- 1977).
nian Brazil, Colombia, Ecuador, Peru, and Vouchers for Guyana.—AMNH A-3791
Venezuela (Frost 2011). from Rockstone, Essequibo River, and A-
18986 and A-18988 from Kamakusa. Also Chiung River; USNM 291157–291171
a series of BMNH specimens cited by from Paramaktoi; USNM 497694–497695
Hoogmoed (1977:267). Also ROM 20648– from Karanambo Ranch, Maricuba Lake
20650 from Tukeit; ROM 39445–39448 (03845 0 08 00 N, 59818 0 36 00 W); USNM 497696
from Mount Ayanganna; and USNM from Lethem, Takutu River (03822 0 38 00 N,
118071 from Kartabo. 59848 0 13 00 W); USNM 497697 from Kaie-
Coloration in life.—The following is teur National Park (near paved airstrip);
based on Hoogmoed (1977) and AL’s USNM 497698 from Georgetown; USNM
color photos of specimens from Guyana: 531299–531303 from several sites within
dorsal ground color reddish brown, often Iwokrama Forest Reserve; and USNM
with darker spots, a dark brown hourglass 535733–535736 from Baramita. In addi-
patch and/or an ochre middorsal stripe. tion, CJC and CRT saw specimens (not
Sides dark brown. Limbs reddish brown collected) at Dubulay Ranch.
with darker crossbands. Throat dark Coloration in life.—‘‘Dorsal ground col-
brown with white spots, venter white with our brown to greyish or reddish brown,
brown mottling. Kok & Kalamandeen with or without dark brown or black
(2008:128) add that ‘‘the dorsal coloration mottling and/or cream spots;...Ventral
can be variable, from grayish brown to surface...creamy white, with or without a
reddish brown, and it may include pale distinct pattern consisting in greyish
blue spots (often present on flanks).’’ brown, dark brown or black spots and/or
Comments.—Essentially nothing is mottling;...Flanks similar to dorsum’’
known about the life history of this species (Kok & Kalamandeen 2008:132).
(Kok & Kalamandeen 2008). Comments.—This is the largest species
of toad in the world, the largest specimens
being from the Guianan Region.
Rhinella marina (Linnaeus, 1758)
Plate 3D
Rhinella martyi Fouquet, Gaucher, Blanc,
Type material.—Originally named Rana & Vélez-Rodriquez, 2007a
marina based on an illustration (Frost Plate 3E, F
2011). The type locality was restricted to
Surinam by Müller & Hellmich (1936). Type material.—The holotype is
Distribution.—Occurs naturally widely MNHN 2006.2601, an adult female from
throughout North America, Central Brownsberg Nature Park, Suriname, Bro-
America, and South America; introduced kopondo district (4856 0 N, 55810 0 W), 510 m
worldwide (Frost 2011). above sea level.
Vouchers for Guyana.—AMNH A- Distribution.—Guyana, Suriname, and
24899–24900 from Kartabo; AMNH A- southwestern French Guiana, ‘‘probably
136024–136026 from Karanambo; AMNH also present in adjacent areas of Brazil and
A-139035–139039 and USNM 566026 may extend into...Venezuela’’ (Fouquet et
from Aishalton; AMNH A-166079 and al. 2007a:24).
USNM 566027 from Berbice River Camp; Vouchers for Guyana.—AMNH A-
and AMNH A-166080 and USNM 566028 24891 and A-39597 from Kartabo;
from Konawaruk Camp. Also ROM 43998 AMNH A-49392 from Isheartun; AMNH
(tadpoles) from Ayanganna; USNM 84491 A-141083–141085, 141089–141090, A-
from Linden; USNM 162894 from Atkin- 145164–145169, and USNM 566029–
son (6827 0 N, 58815 0 W); USNM 162968– 566034 from Dubulay Ranch; AMNH A-
162974 from Enmore Estate (6844 0 N, 166096–166105 and USNM 566035–
57859 0 W); USNM 291104 from Kato, 566037 from Berbice River Camp; AMNH
A-166109–166116 and USNM 566038– Camp (AMNH A-166100) was noted

566040 from Konawaruk Camp; ROM separately as follows: black dorsum with
22813 and 22833 from Baramita (para- gray mottling, the black being quite
types); ROM 20652–20654 from Kurupu- intense anteriorly, beneath the lateral row
kari (4840 0 N, 58839 0 W; paratypes); USNM of warts; chin, throat and chest black with
164164 from Mabaruma Compound gray dots; anterior belly pale yellow with
(08812 0 N, 59847 0 W); USNM 200506 from black mottling; pale yellow beneath arms
Bartica; USNM 531304–531328 from sev- and legs with very little pale black spot-
eral sites within Iwokrama Forest Reserve; ting; iris black and gold, with touch of pale
USNM 535732 from Baramita; USNM copper.
549300 from Waruma River (05820 0 N, Comments.—These toads are part of a
60846 0 W); and USNM 549301–549303 group of cryptic species that occur in the
from the north slope of Mount Roraima. Guianan Region and in the recent past
Coloration in life.—The following is certain populations have been referred to
quoted from the original description (Fou- as Bufo margaritifer and Rhinella margar-
quet et al. 2007a:22), although the authors itifera. Fouquet et al. (2007b) showed with
did not specify that these notes are from DNA sequence data and phylogenetic
animals in life nor did they specify the analyses that there are several lineages of
series of specimens from which the notes this basic kind of toad in French Guiana,
were taken: ‘‘dorsum gray-brown with Suriname, and Guyana that appear to be
dark brown small patches; dark brown on separate evolutionary tracks without
marks also on legs, tarsa and toes; belly evidence of interbreeding, which could
cream slightly orange with more and more signify that they are different cryptic
small grey spots going to the flanks; throat species. Consequently, Fouquet et al.
light grey; interior surface of the tarsa and (2007a) named one of these lineages R.
feet dark brown (except the webbing); no martyi, and they cited specimens from the
middorsal stripe.’’ Fouquet et al. following localities in Guyana: Bartica,
(2007a:24) also noted that ‘‘this species is Baramita, and Kurupukari. The other
highly polymorphic...the back varies from species they recognized were from Suri-
dark brown to light gray and sometimes name, French Guiana, and the southern
even reddish...the patterns...also variable border region of these countries with
with a variety of leaf like patterns and Brazil. However, the confusion about these
successive shades of dark to light brown or animals is such that there may still be
gray. A whitish middorsal stripe can occur unrecognized cryptic species among them.
and can be very thin to 5 mm wide.’’ See Frost (2011) for further discussion.
The following notes are based on the Recently, Avila-Pires et al. (2010), un-
AMNH series from Dubulay Ranch and happy with the changes in taxonomy and
Berbice River Camp (CJC field notes, nomenclature, rejected the new informa-
1994, 1997): tan above (reddish tan or tion and returned to referring to all these
reddish brown before chloretone) with or toads as Bufo margaritifer, synonymizing
without light pale gray or tan vertebral R. martyi with that taxon, but they did so
line; some individuals with dark brown without any molecular data. In the long
spots; line of tan or brown warts on sides; term, this could prove to be more confus-
arms and legs may have dark brown ing rather than enlightening for under-
bands; dark brown stripe on outer lower standing these populations. Consequently,
arm; reddish tan throat and anterior chest; we follow Fouquet et al. (2007a) in
other ventral surfaces pale yellow with referring the Guyanan samples they ana-
black spots; iris a reticulum of black and lyzed to R. martyi. Because there are not
gold. One individual from Berbice River conspicuous reasons to use alternative
names on the other specimens of similar Comments.—After we had reviewed

toads that we have seen from Guyana, specimens and written this part of our
from localities not represented in the study manuscript, Narvaes & Rodrigues (2009)
by Fouquet et al. (2007a), we tentatively published a morphological review of the
use the same name for all the Guyanan toads in this complex, which were previ-
specimens of this group. However, a ously referred to as Rhinella granulosa.
thorough study of these toads throughout They concluded that there are 12 species
the Guianan Region is needed to clarify throughout the overall range of the com-
their status and determine whether there plex, many of which are similar to each
are additional cryptic species present. The other, and three occur in Guyana. Accord-
future study should include morphology, ing to their conclusions, most Guyanan
vocalizations, and DNA sequences ana- localities are occupied by R. merianae, and
lyzed phylogenetically for samples from these include the specimens from Aishal-
throughout the Guianas, including the ton and Dubulay Ranch cited above
entire margaritifera group. (AMNH and USNM catalog numbers).
The other species they recorded from
Guyana are R. humboldti and R. nattereri.
Rhinella merianae (Gallardo, 1965) The single Guyanan locality they cited for
Plate 3C R. nattereri was from Cuyuni-Mazaruni,
Mount Roraima, where they said it is
Type material.—Originally named Bufo sympatric with R. merianae. The only two
granulosus merianae. The holotype, Guyanan localities they cited for R.
AMNH A-46531, is from Head Falls, humboldti were from Cuyuni-Mazaruni,
Essequibo River, Guyana. Bartica; and Upper Takutu-Upper Esse-
Distribution.—Brazil, north of the Am- quibo, Alto Rio Rupununi. We think it is
azon River, eastern Venezuela, Guyana, timely for there to be a thorough review of
Surinam, and French Guiana (Narvaes & the toads of this complex throughout
Rodrigues 2009). Guyana, including ontogenetic develop-
Vouchers for Guyana.—AMNH A- ment of morphology and DNA sequence
70831 from Kartabo; AMNH A-139018– analyses.
139025, A-139028, A-139031, A-139034,
and USNM 566011–566016 from Aishal-
ton; and AMNH A-141065–141073, A- Centrolenidae
141077–141082, A-167103–167104, and
USNM 566017–566019 from Dubulay We follow Guayasamin et al. (2009) in
Ranch. recognizing this family. Its undisputed
Coloration in life.—The following is closest relative is the monotypic Allophry-
based on the AMNH specimens from nidae (see above), in the account for which
Aishalton (CJC field notes, 1993): Dorsum we discussed an alternative opinion of
gray or pale tan with dark brown mark- whether to include Allophryne ruthveni
ings; pale yellow spot in anal region; within the Centrolenidae, as recommended
ventral surfaces cream, abdomen of some by Frost et al. (2006).
individuals with gray, brown, or black Frost et al. (2006) used quotation marks
spots (sometimes subtle); cream warts on for the generic names ‘‘Centrolene’’ and
posterior abdomen; throat and vocal ‘‘Cochranella’’ to indicate that species
pouch of males gold anteriorly, black assigned to these genera at the time did
posteriorly; chin and throat of females not comprise monophyletic groups; this
cream, with or without irregular gray signified that generic assignments would
spots. change, but it would have been premature
to change them in their paper because 58.96 00 W; identification confirmed by Cas-

considerably more research was needed. troviejo-Fisher et al. 2011:44); UG HA 722
Subsequently, Guayasamin et al. (2008) and UTA 51654–51664 from ‘‘a small
showed that the genus Hyalinobatrachium stream on Peters Mountain, 3.6 km north
was not monophyletic either. Taxonomic of Imbaimadai in the Pacaraima Moun-
review of these frogs has been happening tains, 600 m’’ (Noonan & Bonett 2003:92);
quickly, however, as shown by Guayasa- also known from Kaieteur National Park
min et al. (2009) and Castroviejo-Fisher et (Kok & Kalamandeen 2008).
al. (2011). Considering the new generic Coloration in life.—‘‘Dorsum (including
arrangements proposed in the most recent eyelid) light lime green with numerous
papers, which are consistent with the yellow spots, giving appearance of a green
results of their phylogenetic conclusions, ’net’ over a yellow ground color; fine [dark
we drop the use of quotation marks on the brown] melanophores...in green areas;
generic names and use Vitreorana where ventral surfaces. . .[transparent with] parie-
applicable because now the names reflect tal peritoneum clear [translucent]; pericar-
monophyletic groups. dial [peritoneum partly or totally white],
These frogs are difficult to find. They are visceral, and hepatic peritonea white; iris
small and basically green with a high- yellow...and [sometimes] a distinct red
pitched call produced by males perched on ring...encircling the pupil [complete or
green leaves or moss overhanging water, not]; bones white’’ (Noonan & Bonett
often on the underside of a leaf and during 2003:94) [with some details from Kok &
a rain. Egg masses are deposited here, and, Kalamandeen 2008 or Castroviejo-Fisher
if successful, hatching tadpoles drop into et al. 2011, in brackets].
the water below. In some species, particu- Comments.—The latest work on these
larly in the genus Hyalinobatrachium, the frogs was done by Castroviejo-Fisher et al.
male will stay with his clutch of eggs (2011), who showed that Hyalinoba-
during development. Recent research with trachium crurifasciatum Myers & Donnelly
a Costa Rican species has shown that (1997), Hyalinobatrachium eccentricum
clutches with the attendant male have Myers & Donnelly (2001), and Hyalinoba-
better survival rates than those without, trachium ignioculus Noonan & Bonett
and perhaps the male helps discourage (2003) are junior synonyms of H. cappellei.
predation and prevent desiccation (Vock- The latter synonym was named based on
enhuber et al. 2009). specimens from Guyana (UG and UTA
vouchers listed above).
Hyalinobatrachium cappellei
Van Lidth de Jeude, 1904 Hyalinobatrachium iaspidiense
Plate 4A (Ayarzagüena, 1992)
Plate 4B–E
Type material.—The holotype is
RMNH 4463 from the Saramacca River Type material.—The original name was
and vicinity (Gassó Miracle et al. 2007). Centrolenella iaspidiensis, based on the
Distribution.—This species occurs wide- holotype EBD 28803, a male from Que-
ly in the Guiana Shield region, in Brazil, brada Jaspe, San Ignacio de Yuruanı́,
French Guiana, Guyana, Suriname, and Bolivar, Venezuela.
Venezuela (Castroviejo-Fisher et al. 2011). Distribution.—This species occurs wide-
Vouchers for Guyana.—SMNS 12250– ly in the Guianan Region in Brazil, French
12254 from Mabura Hill Forest Reserve, Guiana, Guyana, Suriname, and Venezue-
Maiko Creek (05809 0 19.30 00 N, 58841 0 la, and in the Amazonian lowlands of
Ecuador and Peru. Some records for Vouchers for Guyana.—SMNS 12255–
Guyana are under the name H. noura- 12260 from Mabura Hill Forest Reserve,
guense (see Comments, below). M a i k o C r e e k ( 0 5 80 9 0 1 9 . 3 0 0 00 N ,
Vouchers for Guyana.—SMNS 12242– 58841 0 58.96 00 W); and BMNH 1939.1.1.64
12249 from Mabura Hill Forest Reserve, from the New River. A highly suggestive
Maiko Creek (Kok & Castroviejo-Fisher recording of an advertisement call (vocal-
2008, Yánez-Muñoz et al. 2009, Castro- izing male in Kaieteur National Park) has
viejo-Fisher et al. 2011). been analyzed and a color photograph has
Coloration in life.—Dorsum yellowish been published (see Comments, below).
green with leaf green spots (but white in Coloration in life.—Dorsum pale green
preservative; see also Yánez-Muñoz et al. with numerous tiny yellow dots; belly and
2009) and scattered black dots; belly and parietal peritoneum transparent; pericar-
some peritoneum (e.g., pericardial) trans- dial, hepatic, and visceral peritonea white;
parent (heart visible, red); visceral and iris golden; bones white (Señaris & Ayar-
parietal peritonea white; iris yellow to dull zagüena 2001).
silver; bones white (Lescure & Marty Comments.—Kok & Castroviejo-Fisher
2000:75, under the junior synonym H. (2008:48) compared audiospectograms and
nouraguensis). oscillograms of an uncollected frog that
Comments.—Information on this species was recorded in Kaieteur National Park
has appeared in the literature under the with calls of H. mondolfii from Venezuela
junior synonyms of Hyalinobatrachium and concluded that they were so similar
nouraguensis Lescure & Marty, 2000 and that ‘‘therefore we consider the presence of
H. nouraguense, but recent morphological H. mondolfii in Guyana as likely.’’ Kok &
research suggests that these names are Kalamandeen (2008:143) also stated that
junior synonyms of H. iaspidiense (e.g., ‘‘the presence of Hyalinobatrachium mon-
Yánez-Muñoz et al. 2009). Nevertheless, dolfii in KNP [Kaieteur National Park] is
DNA sequence data indicate there are two probable.’’ Voucher specimens are needed
or more morphologically cryptic lineages to confirm the presence of this species in
that require more detailed analyses Kaieteur National Park. In addition, the
(Guayasamin et al. 2008). See Castrovie- species has been reported for Iwokrama
jo-Fisher et al. (2011) for the latest
(Castroviejo-Fisher et al. 2011).
information.
Hyalinobatrachium mondolfii Hyalinobatrachium taylori (Goin, 1968)

Señaris & Ayarzagüena, 2001
Type material.—The original name was
Plate 5A, B
Centrolenella taylori based on BMNH
Type material.—The holotype is 1939.1.1.65, an adult male from an eleva-
MHNLS 12710, an adult male from ‘‘el tion of 750 ft along the New River,
Primer Raudal del Caño Acoima, afluente Guyana. Frost (2011) noted Hoogmoed’s
del Rı́o Grande (8822 0 N, 61832 0 W), 15 comment in the 1985 edition of the
msnm., estribaciones de la Serranı́a de amphibian checklist volume that the type
Imataca, Estado Delta Amacuro, Vene- locality is within an area that is in dispute
zuela’’ (Señaris & Ayarzagüena 2001:1084). with Suriname.
Distribution.—This species is known Distribution.—‘‘Guianan Shield in
from Bolivia, Brazil, Colombia, French southern Surinam, Guyana, French Gui-
Guiana, Guyana, Suriname, and Venezue- ana, and southeastern Venezuela; presum-
la (Castroviejo-Fisher et al. 2011). ably in adjacent Brazil’’ (Frost 2011).
Vouchers for Guyana.—The holotype Castroviejo-Fisher et al. (2009) for addi-

from the New River (see above); and tional localities.
IRSNB 13981–13987 from Kaieteur Na- Coloration in life.—‘‘Dorsal surfaces
tional Park (Kok & Castroviejo-Fisher dark green; upper lip yellow; hands and
2008). feet yellow green [or bluish green]; bones
Coloration in life.—‘‘Dorsal ground col- green; iris metallic copper with brown
our dark green with pale green spots, reticulations, yellow ring around pupil
usually bearing a white fleck in their centre incomplete laterally; ventral surfaces
[and perhaps some outside the spots], cream; parietal peritoneum clear [transpar-
bronze flecks/lines sometimes present on ent]; visceral peritoneum white; hepatic
dorsal surfaces; iris metallic lavender with peritoneum white; pericardium white’’
dark brown reticulations; ventral sur- (Noonan & Harvey 2000:299).
face...transparent, internal organs visible Comments.—Noonan & Harvey (2000)
through the skin; parietal peritoneum named Centrolene papillahallicum based on
transparent, pericardial peritoneum partly specimens from Guyana (UG and UTA
white, hepatic and visceral peritonea vouchers listed above), but the frogs
white... bones translucent green’’ (Kok & actually were misidentified specimens of
Kalamandeen 2008:146). Centrolene (¼Vitreorana) gorzulae (Kok &
Comments.—Kok & Castroviejo-Fisher Castroviejo-Fisher 2008). The specific ep-
(2008) discussed taxonomic problems with ithet of this frog has been used by some
authors with the ending ai instead of ae as
this species, which are under review, and
used in the original description. This
for which ‘‘at least two species’’ have been
emendation was based on the assumption
masquerading under this name recently.
that the original authors either intended to
See Castroviejo-Fisher et al. (2011) for the
or should have used that spelling. Howev-
latest information.
er, article 32.5.1 of The Code (ICZN
1999:39) states that the original spelling
should be used in cases like this, even if
Vitreorana gorzulae (Ayarzagüena, 1992) certain errors were involved in forming the
Plate 5C name. This phenomenon is discussed
further by Dubois (2007:57). See Castro-
Type material.—The original name was viejo-Fisher et al. (2009) for additional
Centrolenella gorzulae, based on MHNLS details on this species.
11221 from Cerro Auyantepuy (5856 0 N,
62834 0 W; 1850 m), Bolı́var, Venezuela.
Distribution.—‘‘Known from Auyante-
pui, Atapare, and Sierra de Lema, 1800– Vitreorana helenae (Ayarzagüena, 1992)
1900 m elevation, state of Bolı́var, Ven- Plate 5D
ezuela, as well as Kaieteur National Park, Type material.—The original name was
Maringma Tepui, and Peters Mountain in Centrolenella helenae, based on MHNLS
Guyana; presumably to be found in 9431, a male from Quebrada Jaspe, San
adjacent areas of Brazil’’ (Frost 2011). Ignacio de Yuruanı́, Bolı́var, Venezuela.
Vouchers for Guyana.—IRSNB 13974– Distribution.—‘‘Known from the type
13978 from Kaieteur National Park; ROM locality...and Salto Karuay, Bolı́var, Ven-
43982 from Mount Wokomung. Also ezuela, as well as Kaieteur National Park,
CSBD ‘‘HA 721’’ and UTA 52229–52247 Guyana’’ (Frost 2011).
from ‘‘Peters Mountain 3.6 km N of Vouchers for Guyana.—IRSNB 13979–
Imbaimadai in the Pacaraima Mountains, 80 from Kaieteur National Park (Kok &
600 m’’ (Noonan & Harvey 2000:295). See Castroviejo-Fisher 2008); and ROM 43980
from the vicinity of the Potaro River Brown markings consist of irregular spots
(Wokomung Expedition). lateral to the paravertebral rows of tuber-
Coloration in life.—Dorsal surfaces cles and transverse bars on the limbs. The
green to greenish yellow with scattered belly and ventral surfaces of the limbs are
dark brown flecks; eyelids greenish yellow; dull cream, and the throat is brown (black
fingers and toes bluish green; bones green, in calling males)’’ (Duellman 2005:263,
iris yellow; ventral surface transparent; based on frogs from Peru).
parietal peritoneum white, heart not visible Comments.—These frogs of low-eleva-
(Kok & Castroviejo-Fisher 2008:40). tion rain forest have a conspicuous horn
Comments.—See Kok and Castroviejo- over each eye and a huge head and mouth.
Fisher (2008:40) for comments on the They are terrestrial sit-and-wait predators,
value of certain characters for identifying and their exceptionally large gape enables
this species. them to include a wide variety of relatively
large vertebrates and invertebrates as prey.
Ceratophryidae
Ceuthomantidae
We follow Frost et al. (2006) in recog-
nizing this family, which is diagnosed This recently named family (Heinicke et
primarily on the basis of DNA sequence al. 2009) contains three species, all of
data. which occur only in the Guianan High-
lands of Venezuela and Guyana. On the
basis of extensive DNA sequence data and
Ceratophrys cornuta (Linnaeus, 1758) phylogenetic analyses with related frogs,
the ceuthomantids apparently have been
Type material.—The original name was evolving as a clade (branch) on the
Rana cornuta for which the type can be amphibian tree of life separated from their
seen on-line (http://linnaeus.nrm.se/zool/ closest relatives for approximately 60
herp/madamph.html.en, image of NHRM million yr. Only three species of ceutho-
type) (Frost 2011). The type was reported mantids are known to survive today, with
as being from Virginia, obviously in error one of them endemic to Guyana. Among
as the species occurs only in South other Guyanan frogs, their closest relatives
America. are the Craugastoridae, Eleutherodactyli-
Distribution.—Amazonian Colombia, dae, and Hemiphractidae (Pyron & Wiens
Ecuador, Peru, Bolivia, Brazil, Venezuela, 2011).
and the Guianas (Frost 2011).
49247 from Shudikar-wau and A-70873– Ceuthomantis smaragdinus
70874 from Kartabo. Heinicke, Duellman, Trueb, Means,
Coloration in life.—‘‘About 50% of the MacCulloch, & Hedges, 2009
individuals are green with brown mark-
ings, whereas the others are tan with Type material.—The holotype is KU
brown markings...The snout and side of 300000, an adult male from the top of
the head usually are unicolor pale green or Kamana Falls on Mt. Kopinang, part of
pale brown, but in a few individuals dark the Wokomung Massif, Potaro-Siparuni
brown diagonal marks are present in the District, Guyana (05800 0 08 00 N, 59852 0
labial region. In brown individuals, the 47 00 W) at approximately 1540 m elevation.
region demarcated by the interocular and Distribution.—This species is endemic to
paravertebral rows of tubercles is pale tan. Guyana, known from only two localities at
high elevation (1490–1540 m) on Mount (many others occur in the Caribbean), and
Wokomung and Mount Ayanganna (Hei- that is treated in the recently-named family
nicke et al. 2009). Eleutherodactylidae (see below).
Vouchers for Guyana.—Only three spec- These small to medium-sized frogs are
imens are known as follows: the holotype; primarily terrestrial but may be found low
a paratopotype (KU 315000); and ROM on vegetation and rocks. Their eggs are
40161 from Mount Ayanganna (Heinicke laid on land in moist situations (e.g., under
et al. 2009:7). cover on a humid forest floor), not
Coloration in life.—‘‘Dorsum dull olive- necessarily near water. They have direct
brown with diffuse black markings on development; the hatchlings are froglets
body; black transverse bars on limbs; that immediately take on the terrestrial
black longitudinal stripe on inner surface mode of life.
of forearm; black labial bars; broad black
canthal stripe; bright, almost phosphores-
cent green interorbital bar; pair of diago- Pristimantis chiastonotus
nal marks in scapular region; spot on (Lynch & Hoogmoed, 1977)
anterior surfaces of upper arm; distinct
green bar below black canthal stripe. . .; Type material.—The original name was
dorsal surfaces of discs on fingers white; Eleutherodactylus chiastonotus and the
dorsal surfaces of toe pads creamy white holotype is RMNH 17614, an adult male
with black suffusion in terminal notch; from Brownsburg, Brokopondo District,
venter creamy gray, heavily mottled in Surinam, 500 m.
black; throat nearly entirely black. . .; belly Distribution.—Guyana and ‘‘low eleva-
mottled black and gray; iris greenish tion forests in northeastern Brazil, French
bronze heavily flecked with black’’ (Hei- Guiana, and Surinam (below 700 m)’’
nicke et al. 2009:8). (Frost 2011).
Comments.—Essentially nothing is Vouchers for Guyana.—AMNH A-
known about the biology of these frogs. 46564 from Marudi and A-46245 and A-
91649 from Shudikarwau.
Coloration in life.—‘‘Back and legs
Craugastoridae ochre-yellow, beige or yellow-orange, with
a pattern of light-brown or beige-brown
We follow Pyron & Wiens (2011) in bands of varying width, forming a distinct
recognizing this family, which is diagnosed X-shaped mark on the back, chevrons or
primarily on the basis of DNA sequence oblique bands on the back and flanks,
data. Previously, these frogs were consid- transverse bands on the legs, an interorbital
ered as part of the family Leptodactylidae, bar, 2 spots on the snout and 2 bands
then Strabomantidae, but the current radiating from the eye to the upper lip. The
arrangement is based on the most recent elements of the darker pattern are bordered
phylogenetic analyses (Pyron & Wiens by a narrow black or dark-brown and a
2011). In addition, the generic name narrow pale yellow line. Some specimens
Eleutherodactylus was used previously for (from Löe Creek and Saül) have a cream-
most of the species of craugastorids that colored vertebral stripe from the tip of the
occur in Guyana, and today these are all snout to the vent. Canthal and supra-
assigned to the genus Pristimantis (Hei- tympanic stripe dark brown. Dorsally this
nicke et al. 2007), an ancient and diverse stripe is bordered by a narrow pale-brown
South American clade of more than 400 to pale-yellow line passing over the canthus
species. There is now one species of true rostralis and the outer edge of the upper
Eleutherodactylus that occurs in Guyana eyelid; ventrally there is no sharp border
anterior to the tympanum; in some speci- red or dark purple with several bright red
mens nearly the whole loreal region and the spots; posterior of thigh bright red; lower
upper lip are dark-brown. Tympanum arm and dorsal surfaces of hands, feet and
chestnut-brown to light brown. A dark- toes dark purple with bright red spots,
brown spot at the base of the forelimbs is more prominent distally; hand red, finger
sometimes present. Throat white, dirty- discs purple; upper leg surface purple with
white or yellow with gray spots, belly small red spots; three obscure reddish cross
yellowish-white. Groin, ventral, anterior bands on crus; toe discs outlined by
and posterior surfaces of the thighs, poste- purple. Undersurface of the skin over each
rior and ventral surfaces of the lower leg dentary dark purple with four discrete, red
and palms bright orange. The soles may be spots; throat dark purple to black, extend-
orange, gray or dark-brown. Ventral sur- ing onto the anterior pectoral area as a
face of forelimbs transparent flesh-colored. small bib. Venter below the bib uniformly
Iris divided in 3 parts: an upper golden bright red or, in some individuals, marked
part, a silvery gray, gray-brown, or gray- with a faint irregular, mid-ventral suffu-
green lower part, both parts separated by a sion of black pigment from bib to hind
horizontal reddish-brown bar. Around the limb insertions; in some of these, a faint
pupil a narrow band, anteriorly reddish black wash over red is present lateral to the
brown, posteriorly yellow’’ (Lynch & mid-ventral darker pigment (the mid-ven-
Hoogmoed 1977:429–430). tral and lateral dark wash becoming much
more pronounced in preservative); under-
side of legs sprinkled with tiny dark
Pristimantis dendrobatoides punctuations, most heavily on crus; pal-
Means & Savage, 2007 mar and plantar surfaces dark purple. Iris
Plate 17A black with faint reddish punctations, but
Type material.—The holotype is USNM thin and continuously outlining the edge of
563662, an adult male from the Woko- the pupil. No color changes were noted
mung Massif, near Falls Camp, Potaro- between day and night. Specimens found
Siparuni District, west-central Guyana with dry skin...had a blue-gray cast to their
(05805 0 25 00 N, 59850 0 18 00 W, 1385 m). dorsal skin as opposed to wet frogs’’
Distribution.—‘‘Known only from the (Means & Savage 2007:44).
Wokomung Massif in west-central Guya-
na in cloud forest habitat (1385–1570 m)’’
(Means & Savage 2007:45). Pristimantis inguinalis (Parker, 1940)
Vouchers for Guyana.—ROM 43317, Plate 17B
USNM 563661–563662 and USNM
564161–564164, all from the Wokomung Type material.—The original name was
Massif. Eleutherodactylus inguinalis and the holo-
Coloration in life.—‘‘Upper surfaces of type is BMNH 1939.1.1.1, a female from
head and body dark purple; head uniform- the New River, Guyana. Frost (2011)
ly dark purple except for one or two small noted that Hoogmoed (in the Frost 1985
interorbital bright red spots and three or version of the Amphibian Species of the
four bright red lip spots present in USNM World) mentioned that this locality is in an
563661 but absent in the holotype (USNM area under dispute between Surinam and
563662); one distinct, bright red, round Guyana.
middorsal, suprascapular spot on all seven Distribution.—‘‘Several localities in
specimens with 0–4 smaller, round, mid- Guyana, Surinam, and French Guiana’’
dorsal suprasacral spots; upper arm bright (Frost 2011).
Vouchers for Guyana.—The holotype; mostly yellow-green dorsally with four

and ROM 40163–40164 from Mount bright red lateral blotches; head olive green,
Ayanganna. mottled green and black, or with a broad
Coloration in life.—The following is orange medial stripe. Upper lips with red
translated and paraphrased from Lescure and olive green patches set off by thin
& Marty (2000:208), based on specimens yellow green vertical lines; upper arm olive
from French Guiana. Dorsal surface light green; lower arm with one or two red and
brown with large brownish black spots, of green equally wide crossbands; finger and
which one behind the head is in the form of toe discs dark red to dark gray; upper leg
a W and another in the form of a chevron. surface with 3 or 4 alternating red and olive
Black supralabial bars. Ventral surface green crossbands continuing onto the crus
light gray with numerous small black dots. when legs are folded against the body; ...
Yellowish orange spot in groin. Iris gray to throat, belly, and ventral surfaces of arms
reddish with fine, irregular black lines. and legs mottled in cream and black except
Kok & Kalamandeen (2008:240) describe in the specimen with the bold middorsal
frogs from Kaieteur National Park as stripe which had a cream colored, midven-
having a more variable dorsum (‘‘greenish tral longitudinal stripe; upper 2/5 of iris blue
brown, brown or dark brown’’); possible gray against the sclera, changing to metallic
dorsolateral stripe; flecks on ventral sur- brown around and below the iris. No color
faces dark brown or white; iris possibly changes were noted between day and night’’
gold in upper part, reddish gray in lower. (Means & Savage 2007:47).
Pristimantis jester Means & Savage, 2007 Pristimantis marmoratus

Plate 17C, D (Boulenger, 1900a)
Plate 17E
Type material.—The holotype is USNM
563631, an adult male from the summit of Type material.—The original name was
the Wokomung Massif called Mt. Woko- Hylodes marmoratus and the holotype is
mung, Potaro-Siparuni District, Guyana BMNH 1947.2.16.92 (formerly 99.3.25.19)
(05804 0 03.3 00 N, 59851 0 41.8 00 W, 1560 m). from the foot of Mt. Roraima, 3500 ft,
Distribution.—Known from the slopes reported originally as being from Guyana.
and tops of the highest peaks of the However, the type specimen was collected
Wokomung Massif and from Mount Ayan- in adjacent Venezuela (Phelps 1938). See
ganna (MacCulloch & Lathrop 2009). below (Vouchers) for records from Guya-
Vouchers for Guyana.—ROM 43303, na.
43306, and USNM 563631–563633, holo- Distribution.—Occurs widely in the Gui-
type and paratypes, all from the Woko- ana Shield region in ‘‘eastern Venezuela,
mung Massif; ROM 40169–40171 from Guyana, Surinam, and French Guiana;
Ayanganna. presumably in adjacent Brazil’’ (Frost
Coloration in life.—‘‘Upper surfaces of 2011).
head and body extremely variable with dark Vouchers for Guyana.—ROM 40159–
red, light green, and black marbling. In 40160 and 40162 from Mount Ayanganna;
some specimens the red predominates but in USNM 549318–549319 from the N slope
others the green and black are more of Mt. Roraima; and USNM 566156 from
extensive. In one specimen (USNM Ceiba Biological Station along the Made-
563632) a bold, wide, pure orange stripe wini River, ca. 42 km S Georgetown on
runs from the tip of the snout to the cloaca; Linden Highway (06829 0 57 00 N, 058813 0
another specimen (USNM 563633) was 06 00 W).
Coloration in life.—The following is mung Massif, near Falls Camp, Potaro-

from Kok & Kalamandeen (2008:242), Siparuni District, Guyana (05805 0 25 00 N,
based on observations in Kaieteur Nation- 59850 0 18 00 W, 1385 m).
al Park: dorsal color variable, brown, Distribution.—Known only from the
greenish brown, grayish brown, reddish Wokomung Massif, Guyana.
brown, or dark brown, with dark markings Vouchers for Guyana.—USNM 563634–
(one W-shaped on neck); ventral surface 563651, 564165–564179, and ROM 43302,
grayish white; upper part of iris gray, 43307, 43309, 43310, 43313, 43314. These
lower part copper. Lescure & Marty are the holotype, paratypes, and referred
(2000:212) said that the iris has black specimens, all from the Wokomung Mas-
vermiculations. sif.
Coloration in life.—‘‘Dorsal color pat-
tern extremely variable due to pattern
Pristimantis pulvinatus (Rivero, 1968a) polymorphism as well as metachrosis...
Frogs may be darker or lighter overall, as
Type material.—The original name was a result of handling, background colora-
Eleutherodactylus pulvinatus and holotype tion, temperature, or time of day. Upper
is MCZ 64741, female from Paso del surfaces of head and body light brown to
Danto, Región de La Escalera around tan with extremely variable dark brown
1400 m above San Isidro, road from El markings that range from small spots or
Dorado to Sta. Elena de Uairén, Bolivar, irregular blotches..., through single trans-
Venezuela. verse suprascapular bar, to a series of
Distribution.—Western Guyana and several oblique bars that extend laterally
‘‘La Escalera and the upland forests of onto the flanks... Many specimens have an
the Gran Sabana region of southeastern unbroken, cream to light tannish orange
Venezuela’’ (Frost 2011). colored middorsal stripe highlighted by a
Vouchers for Guyana.—ROM 40156– suffusion of black pigment laterally...; the
40158 from Mount Ayanganna. stripe may be narrower than the width of a
Coloration in life.—Dorsum medium finger or half as wide as the interorbital
brown, with a dark brown interorbital distance; others have a broad middorsal
bar, a dark W-shaped mark on anterior light tan field bordered laterally by a black
back, and some large blotches and small margin..., and a few have the dorsum
spots on the posterior back; black supra- marked with slightly irregular dark and
tympanic streak; flanks light brown with light longitudinal stripes. Head in many
oblique gray bars; limbs medium brown individuals with a dark brown interorbital
with gray crossbands; upper lip with bar set off by tan color on snout...; canthus
triangular brown blotches on gray back- rostralis usually marked with a strong dark
ground; dark canthal streak may be stripe; upper surfaces of arms and legs the
present. Ventral surfaces dirty white, same color as dorsum; finger and toe discs
darker on throat and distal portion of pinkish tan; upper and lower legs and crus
limbs (Rivero 1968a). on some specimens have a hint of three to
four light gray and tan alternating cross-
bands and on others there are three dark
Pristimantis saltissimus brown bands; throat of males white with
Means & Savage, 2007 black smudging or mottling, often white in
Plate 17F females; belly and undersurfaces of legs
heavily marked with dark pigment some-
Type material.—The holotype is USNM times mottles as on throat, but lighter
563639, an adult female from the Woko- overall...; individuals with a middorsal
stripe also have a mid-ventral, off-white Dendrobatidae

streak...; upper two-fifths of iris dirty gold
color, lower three fifths dark brown to Although this family name has been
black’’ (Means & Savage 2007:51). used for about 135 yr (Cope 1865), the
family has been reorganized and the
Aromobatidae have been split out from
within it. We follow Grant et al. (2006) in
Pristimantis zeuctotylus
their revision of the family, which is
(Lynch & Hoogmoed, 1977)
diagnosed on the basis of five morpholog-
Plate 18A
ical characters as well as considerable
Type material.—The original name was DNA sequence data. In the recent past,
Eleutherodactylus zeuctotylus and holotype the generic name Epipedobates has been
is RMNH 17701 from the west slope, Vier improperly assigned to some of the
Gebroeders Mountain, Sipaliwini, Nick- Guyanan species currently assigned to
erie District, Suriname. Ameerega and Dendrobates.
Distribution.—‘‘Wet rainforests of These small frogs include some species
northeastern South America from extreme with bright aposematic colors that signal
southeastern Colombia through Amazo- to predators that they have skin glands
nian Venezuela and the Guianas, as well as with powerful toxins; these have potential
scattered records from northwestern Brazil for biomedical research (e.g., Daly et al.
and southwestern Brazil’’ (Frost 2011). 2005). In many instances, the frogs do not
Vouchers for Guyana.—‘‘Rupununi Dis- synthesize the toxins but sequester them
trict: N of Acarai Mountains, W of New after they are obtained in small inverte-
River, KU 69663; Shudikar-wau, AMNH brate prey (e.g., Daly et al. 2000, 2002).
43680’’ (Lynch & Hoogmoed 1977:432); The mechanisms of sequestration are
also AMNH A-166327–166336 and unknown. Eggs are laid hidden in moist
USNM 566157–566160 from Berbice Riv- situations on the ground or in vegetation,
er Camp. and for some species, after hatching, a
Coloration in life.—‘‘Back orange- parent carries the tadpoles to water. There
brown, red-brown, brown, gray brown, are reports for several species in this family
or greenish-gray with an indistinct darker (e.g., Brust 1993, Summers et al. 1999) of
pattern of lines and spots; flanks some- females or pairs returning to places where
times yellowish-green; snout, in front of they previously oviposited in nutrient-poor
eyes, paler than dorsum; upper arm of locations and depositing unfertilized or
juveniles orange-brown to yellow; venter even fertilized eggs, which are consumed
dark gray to black; throat with white by their developing tadpoles.
spots; upper lip spotted black and white;
canthal and supratympanic stripe black;
tympanum chestnut-brown; iris gold, di- Ameerega hahneli (Boulenger, 1884)
vided horizontally by reddish-brown band
(lower part of iris sometimes darker than Type material.—The original name was
upper part)’’ (Lynch & Hoogmoed Dendrobates hahneli and BMNH
1977:436–437). Field notes of CJC (1997) 1947.2.15.17 from Yurimaguas, Huallaga
on the AMNH and USNM series from the River, Northern Peru was designated
Berbice River Camp are consistent with lectotype by Silverstone (1976).
the above and include the following: thin Distribution.—‘‘Amazonian lowlands
light yellow ring around outer edge of of...Peru, Ecuador, Colombia, Bolivia,
tympanum; iris copper. Brazil, the extreme south of Venezuela,
French Guiana, and likely adjacent Sur- AMNH A-21401 from Kamakusa, AMNH
inam, and southern Guyana’’ (Frost 2011). A-45774 from Wismar, AMNH A-46232
Vouchers for Guyana.—AMNH A- from Kuyuwini Landing, AMNH A-46236
11667, A-13502–13503, A-23120–23121, from Parabam, and AMNH A-46570 from
A-23126–23127, A-39575–39576, A- Marudi. Additional specimens are USNM
39677–39678, A-39681–39686, A-39692– 535738–535748 from Baramita and USNM
39695, A-39698, A-39702–39704, A- 566055 from 6.4 km SE Aishalton.
39706–39707, A-39709–39717, and A- Coloration in life.—The following notes
70984–70987 all from ‘‘Essequibo: Karta- were recorded on the USNM specimen
bo’’ (quoted from the AMNH catalog). from 6.4 km SE Aishalton (CJC field
Coloration in life.—The following is notes, 1993): Dorsum black; dorsolateral
translated and paraphrased from Lescure light stripe bright goldish yellow; lateral
& Marty (2000:96), based on specimens light stripe same color as this anterior to
from French Guiana: Dorsum black; thin arms, but very pale yellow posterior to
white dorsolateral stripe from snout to arms; arms and legs dark green above;
groin; fine white stripe edging lower lip, hands and feet, above, green with irregular
extending to shoulder; sides of body blue spots; posterior of thighs black with
brownish black; small yellow or orange irregular blue spots; chin, throat, and chest
spot in axilla and groin; ventral surface black; abdomen black with irregular cream
brownish black to black, marbled laterally or pale blue spots; beneath arms and legs
and posteriorly with light blue; yellow or black with irregular blue spots.
orange spot on anterior surface of thighs, Silverstone (1976:47) noted the follow-
extending that of the groin and visible on ing for specimens of the pattern type found
the upper surface; yellow or orange spot in the Guianan Region: ‘‘The lateral
on lower leg; iris dark chestnut. stripes are green, yellow-green, yellow, or
Comments.—Specimens of this species orange. The dorsum is black; there is no
from the Guiana Shield region previously median dorsal stripe...Specimens from
were referred to as Dendrobates or Epi- Suriname...[have] green spots...[on] the
pedobates pictus. dorsal surface of the limbs; light blue spots
adorned the belly and the ventral surface
of the limbs.’’
Ameerega trivittata (Spix, 1824) Comments.—This species occurs in low-
Plate 5E land forests, from ‘‘20 to 680 m’’ elevation
(Silverstone 1976:47).
Hyla Trivittata and RMNH 1836 from Rio
Tefé, Brasil was designated as lectotype by Dendrobates leucomelas
Hoogmoed & Gruber (1983). Steindachner, 1864
Distribution.—‘‘Guianas and the Ama- Plate 5F
zon drainage of Brazil, Peru, Bolivia
(Pando), Colombia (Putumayo and Ama- Type material.—The holotype from Co-
zonia), and Venezuela; presumably in lombia is NMW 19188, according to
Amazonian Ecuador’’ (Frost 2011). Häupl and Tiedemann, 1978.
Vouchers for Guyana.—Silverstone Distribution.—‘‘Guianan Orinoco drain-
(1976:48) listed 11 localities scattered age of Venezuela north to the Rı́o Orino-
around Guyana, each documented with co, east into Guyana to the Essequibo
specimens from various museums, includ- River, south into extreme northern Brazil,
ing the following: AMNH A-13493, A- and west into eastern Amazonian Colom-
24925, and A-70988–70989 from Kartabo, bia’’ (Frost 2011).
Vouchers for Guyana.—AMNH A- from preservative for a few minutes and

43823–43825 from ‘‘Kopang River’’ (Sil- starts to dry several reddish brown areas
verstone 1975:26), but in the AMNH appear on the posterior part of the
catalog as ‘‘Kumpang’’; AMNH A-49942 abdomen, dorsally and laterally on the
from ‘‘Kurupung River’’ (Silverstone thighs and in the interorbital area, which
1975:26), but in the AMNH catalog as might indicate lighter reddish brown col-
‘‘Kurupung, Upper Mazaruni’’; AMNH ouration in the living frog’’ (Jungfer &
A-70983 from Kartabo; and USNM Böhme 2004:102).
118044 from 16.1 km N Kaieteur Falls Comments.—This is one of the most
on Potaro River. poorly known species in the world. Jungfer
Coloration in life.—According to Silver- & Böhme (2004) suggested that it belongs
stone (1975:26), ‘‘most Venezuelan speci- to the tinctorius species group, and its
mens are black with three broad yellow [or closest relative may be D. leucomelas.
orange] transverse dorsal bands containing
black intrusions, bands, or spots. The limbs
are yellow, spotted with black. The venter is Dendrobates tinctorius (Cuvier, 1797)
black, sometimes with a yellow spot.’’
Comments.—This species inhabits low- Type material.—The original name was
land moist forests from ‘‘50 to 800 m’’ Rana tinctoria. The neotype designation
elevation (Silverstone 1975:26). has a complex history (Frost 2011).
Distribution.—‘‘Lowland forests of the
Guianas and adjacent Brazil’’ (Frost 2011).
Dendrobates nubeculosus Vouchers for Guyana.—AMNH A-
Jungfer & Böhme, 2004 49301–49328 from Shudikar-wau and
USNM 164813 from Kamoa Mountains.
Type material.—The holotype is ZFMK Coloration in life.—Color and pattern
45354, an adult male from Rockstone, are extremely variable. Lescure & Marty
Essequibo River, Mazaruni Potaro Dis- (2000) have color photographs of several
trict, Guyana. very different colorations on frogs from
Distribution.—‘‘The collecting site was French Guiana. The following quotes
noted as ‘‘Rockstone (5858 0 N, 58832 0 W) (Silverstone 1975:45–46) are for specimens
. . .7 m above sea level’’ (Jungfer & Böhme geographically relevant to Guyana: ‘‘Spec-
2004:102). imens from the Kayser Mountains, Sur-
Vouchers for Guyana.—Known only iname, are black with dorsolateral stripes;
from the holotype. these stripes are yellow on the snout and
Coloration.—The holotype was collected greenish white on the back, and connect
by ornithologists in the 1980s. ‘‘In alcohol, with a greenish white reticulation on the
ground colouration of all surfaces dark sides of the body [these light areas being
brown. Dorsal surfaces of head and body yellow or yellow-orange on specimens
with numerous irregular, ill-defined light from French Guiana]. The digital disks
grey spots (creamy white or yellow in life?), are light blue. The limbs and venter are
laterally on the body these spots merge to black with dark purplish blue reticula-
form an irregularly reticulate pattern. Legs tion...Most specimens from Guyana and a
and side of head uniformly dark brown, few specimens from Suriname have stripes
although a few small light spots may have composed of reticulation...Specimens from
darkened on the dorsal surfaces of the Shudikar-wau, Guyana, have a dorsal
humeri. Ventrally two light brown spots in pattern transitional between stripes com-
the pectoral area and a light grey spot on posed of reticulation and reticulation
mid-venter. When the specimen is removed covering the whole back; some of these
specimens are similar in pattern and color (Bolı́var), southern Surinam, southern
(blue in life, no yellow [R. Snedigar, field French Guiana, and to Serra do Veado,
notes at AMNH] to D. azureus, but the Amapá, Brazil’’ (Frost 2011).
reticulation is broader (i.e., the interspaces Vouchers for Guyana.—The holotype;
are larger).’’ IRSNB 13784–13793 from Kaieteur Na-
Comments.—Silverstone (1975:46) not- tional Park; ROM 39578 from the base of
ed similarities between the AMNH series Mount Ayanganna; ROM 43036–43037
from Shudikar-wau and Dendrobates azur- from Mount Wokomung; ROM 44050–
eus from Suriname (also see above for 44051 from Meamu River (06816 0 N,
color) but ‘‘the tympanum is larger than in 060830 0 W); and USNM 549315 from the
the type material of D. azureus. Provision- N slope of Mt. Roraima (5817 0 N,
ally, I include the Shudikar-wau popula- 60845 0 W).
tion in D. tinctorius.’’ Later, a molecular Coloration in life.—‘‘Dorsal ground col-
phylogenetic analysis (Wollenberg et al. our medium brown or grey. A middorsal
2006) confirmed the extreme variability in black ‘‘)(‘‘, black canthal and supratym-
color and pattern and concluded that D. panic bars and other scattered black marks
azureus is a junior synonym of the highly may be present; numerous irregular sky
variable D. tinctorius. blue or white dots on dorsum and flanks;
dorsal surfaces of hind limbs and forearms
medium brown or grey, with or without
Eleutherodactylidae black marks or crossbands and small
irregular sky blue or white dots; upper
Frogs of this family, which is diagnosed arm orange...Venter brown or grey with
primarily on DNA sequence data, until small irregular sky blue or white dots; iris
recently were included in the family copper, with a red ring around pupil. In
Leptodactylidae, but molecular phyloge- preservative the orange and sky blue colors
netic analyses have shown that this partic- become white, otherwise there is little
ul ar li neage sh oul d be s pl it out change’’ (MacCulloch et al. 2008a:46).
taxonomically, consistent also with split- Comments.—Although they may be
ting out the Craugastoridae (e.g., Heinicke locally abundant, the small size (maximum
et al. 2007, Hedges et al. 2008, Pyron & body length of 16 mm) and secretive habits
Wiens 2011). These terrestrial frogs are of Adelophryne gutturosa make it difficult
tiny to rather small in size and, for the to find. It has been found at elevations
species studied so far, have direct develop- ranging from 110-1234 m. This species
ment; hatchlings are froglets that immedi- produces large eggs and probably under-
ately take on the terrestrial mode of life. goes direct development (for a review of
natural history, see MacCulloch et al.
2008a).
Adelophryne gutturosa
Hoogmoed & Lescure, 1984
Adelophryne patamona
Type material.—The holotype is BMNH MacCulloch, Lathrop, Kok, Minter,
1983.1139, a male from between camp IV Khan, & Barrio-Amorós, 2008
and V, northern slopes of Mount Rorai- Plate 6A
ma, Guyana (5817 0 N, 60846 0 W, 3000 ft
[914 m]). Type material.—The holotype is ROM
Distribution.—‘‘Guiana Shield, from 43034, an adult male from the north slope
Mount Roraima and Pakaraima Moun- of Mount Wokomung, Guyana (05806 0
tains, Guyana, nea rby Venezuel a 35 00 N, 059849 0 15 00 W, 1234 m).
Distribution.—This species is known Distribution.—‘‘Lesser Antilles; likely

only from Guyana, in the Pakaraima originally from St. Lucia or the Antigua
highlands at elevations ranging from and Barbuda Bank but also known from
1060–1411 m, but probably also occurs in the islands of St. Martin, Saba, St.
adjacent Brazil (MacCulloch et al. 2008a). Eustatius, St. Christopher, Nevis, Guade-
Vouchers for Guyana.—ROM 43034– loupe, Montserrat, Martinique, Bequia,
43035 from Mount Wokomung. Also Anguilla, St. Lucia, St. Vincent, Barbados,
IRSNB 1971–1978 from two sites around Grenada, Mustique, Canouan, and Car-
Mount Maringma (see MacCulloch et al. riacou; definitely introduced on Trinidad,
2008a:37 for details). Jamaica, Bermuda, Dominica, Venezuela,
Coloration in life.—‘‘Ground colour of Panama, Barranquilla (Colombia), San
dorsum, flanks and dorsal surfaces of limbs José (Costa Rica), Curaçao, and George-
medium brown. Central part of dorsum town, Guyana, and coastal French Gui-
dark brown, bounded by a black middorsal ana; likely on coastal Surinam’’ (Frost
‘‘)(’’-shaped mark and interorbital bar. 2011). In Guyana, this species is abundant
Dorsum with a few other small, scattered in urban areas of Georgetown, Linden,
black marks. Loreal and temporal regions and New Amsterdam (Bourne 1997).
dark brown. A black bar extending from tip Vouchers for Guyana.—AMNH A-
of snout through eye to flank; another black 18981–18982, A-21403–21404, and A-
bar in the inguinal region. Dorsal surfaces of 21413; UMMZ 80415, 80443, and 83576;
limbs with black crossbands. Throat, chest and USNM 118685 and 194416–194428,
and underside of arms dark grey with white all from Georgetown.
or pale blue spots. Venter, underside of legs Coloration in life.—The following is
and rear of thighs reddish-brown with white translated and paraphrased from Lescure
or pale blue spots. A black anal spot. & Marty (2000:210): Dorsum brown with
Palpebrum not reticulated, a black band one or two darker brown chevrons, a W,
along its upper rim. Iris copper with a red or two dorsolateral stripes that may be
ring around pupil...In preservative the venter either lighter or darker in color. Ventral
becomes brownish-grey with white spots’’ surface whitish. Iris golden.
(MacCulloch et al. 2008a:39). ‘‘Black dorsal Comments.—Adults usually are smaller
markings are variable; some specimens are than 35 mm in body length. Eleutherodac-
darker with more pale blue spots...In the tylus johnstonei is a native of Caribbean
specimens from Maringma the throat, chest islands and was introduced into Guyana
and underside of arms are black with white prior to 1923 (Hardy & Harris 1979,
or pale blue spots and a few reddish-brown Kaiser et al. 2002). Previously, some
dots; the venter, underside of legs and rear of Guyanan specimens of this species were
thighs are reddish-brown, sometimes with misidentified and incorrectly reported as
very few white or pale blue spots’’ (MacCul- Eleutherodactylus urichi (Kaiser et al.
loch et al. 2008a:41). 1994); CJC also has re-examined many of
Comments.—See MacCulloch et al. these specimens (among the AMNH and
(2008a) for notes on natural history. This UMMZ vouchers cited above), and we are
species reaches a body length of 23 mm, confident that E. urichi does not occur in
the largest known in the genus. Guyana.
Eleutherodactylus johnstonei Barbour, 1914 Hemiphractidae

Type material.—There are two syntypes, We follow Guayasamin et al. (2008) in
MCZ 2759 from St. Georges, Grenada. recognizing this family, which is diagnosed
primarily on the basis of DNA sequence plateau of Mount Ayanganna, Guyana

data, although Frost (2011) commented (05824 0 N, 059857 0 W; elevation 1490 m).
that the question of monophyly for this Distribution.—Known only from Guya-
group needs confirmation through addi- na (MacCulloch & Lathrop 2006a).
tional research. Although these frogs Vouchers for Guyana.—ROM 39473–
appear superficially like large, agile tree 39476 and ROM 40769 from Mount
frogs and previously were included in the Ayanganna; and ROM 42804–42831 from
Hylidae, it now appears that Hylidae is not Mount Wokomung.
their closest relative (see Guayasamin et al. Coloration in life.—‘‘Dorsum, top of
2008, Frost 2011, and Pyron & Wiens 2011 head and loreal region golden brown with
for summaries of recent research). numerous small orange spots. Lips and
Stefania is the only genus of this family suborbital region light brown with distinct
known to occur in Guyana, and it is black mottling on lips and one vertical
represented here by seven species, six of black bar extending from eye to lip.
which are endemic to Guyana. All other Ventral margin of supratympanic fold
locality records for the genus are in black; this line continues forward along
southern and eastern Venezuela, with the canthus past nostrils to tip of snout.
exception of one Brazilian species recently Temporal region and tympanum dark
described (Carvalho et al. 2010). brown. Limbs bronze above, with orange
These frogs can be found on rocks, in flecks more concentrated on forearm.
trees, and on the ground in moist areas. Transverse bars complete on thighs, in-
For the species for which life history is complete on shanks, tarsi, and forelimbs.
known, females carry their clutch of Flanks, belly, throat, posterior thigh and
relatively large fertilized eggs on their ventral surfaces of limbs purple-grey with
back; being large, the eggs are also few creamy white mottling most concentrated
per clutch. There is no brood pouch on chest and throat region and reduced to
holding the eggs in place; instead, they irregular spots on flanks and limbs. Iris
adhere by means of a mucoid glue-like bicoloured, top third gold, lower two
substance. The adult female can jump for thirds reddish brown’’ (MacCulloch &
quite a distance and land abruptly, without Lathrop 2002:335–336). In some individu-
the eggs becoming dislodged, so the als the dorsal background is grey rather
adhesive and mechanics of the attachment than brown (MacCulloch et al. 2006).
warrant further study. During develop- Comments.—There are no data on
ment, if the eggs become somewhat desic- number of eggs or juveniles per clutch,
cated the female can rehydrate them in and the call is unknown.
water. There is no free-living larval stage
or metamorphosis, as tiny froglets emerge
from the eggs. These remain on the back of Stefania ayangannae
the female (attached by seat-belt-like MacCulloch & Lathrop, 2002
membranes) for a short while before Plate 6C
dispersing.
Type material.—The holotype is ROM
39501, an adult male from the northeast
Stefania ackawaio plateau of Mount Ayanganna, Guyana
MacCulloch & Lathrop, 2002 (05824 0 N, 059857 0 W; elevation 1490 m).
Plate 6B Distribution.—Known only from Guya-
na (MacCulloch et al. 2006).
Type material.—The holotype is ROM Vouchers for Guyana.—ROM 39501–
39473, an adult male from the northeast 39551, ROM 39640, and USNM 561833–
561835 from Mount Ayanganna; and Vouchers for Guyana.—ROM 39477–

ROM 42893–42952 from Mount Woko- 39480 from Mount Ayanganna; and
mung. ROM 42856–42860 from Mount Woko-
Coloration in life.—‘‘Dorsal color is mung.
yellowish-brown or yellowish-red with Coloration in life.—‘‘The dorsum is
diagonal paravertebral dark brown marks, medium brown or grey with a few scat-
often joined to form chevrons. A dark tered dark brown marks. Flanks are
brown patch is present on the head, medium brown with darker brown mot-
bordered anteriorly by a cream interorbital tling. The lower limbs, hands and feet are
bar. Canthal, supratympanic and lip bars light brown with dark brown mottling.
are dark brown, with a cream vertical line The distal phalanges of the toes have a red
on the tip of the snout. Flanks have five wash. Dark brown canthal and supra-
tympanic stripes are present. The upper
narrow diagonal dark brown bars. The
lip is creamy white with black mottling; the
venter is pale yellow with dark brown
lower lip is black with a few white spots.
mottling on the throat. Limbs are yellow-
The throat is brown or grey with white
ish-brown with dark brown crossbands mottling; the venter and underside of
dorsally, pale ventrally with increasing thighs are white with a variable amount
dark brown stippling distally. Palms and of brown or grey mottling...Neonates are
soles are darker brown, and the distal dark grey with immaculate limbs and a
phalanges of the toes are red. In some cream interorbital bar; some have a dorsal
individuals the markings are much less chevron pattern’’ (MacCulloch & Lathrop
prominent’’ (MacCulloch & Lathrop 2006c:1).
2006b:1). Comments.—The number of eggs or
Comments.—‘‘Intra- and interpopula- juveniles ranges from four to six per
tion variation in disc size, finger length, clutch. The call is unknown.
skin texture and coloration occurs in
Stefania ayangannae from Wokomung
and Ayanganna’’ (MacCulloch et al. Stefania evansi (Boulenger, 1904)
2006:35). The call is unknown. Number Plate 6E
of eggs or juveniles ranges from four to
nine per clutch. Means et al. (2008) Type material.—The original name was
describe amplexus and oviposition behav- Hyla evansi and the holotype is BMNH
ior, with photographs showing how the 1947.2.13.11 (formerly 1904.5.30.1; Condit
male and female cooperate in getting the 1964:89) from Groete Creek, Essequibo,
eggs placed on the female’s back. Guyana.
Distribution.—This species is known
only from ‘‘forested lowlands of west-
central Guyana, up to 900 m’’ (MacCul-
Stefania coxi loch & Lathrop 2006d:2).
MacCulloch & Lathrop, 2002 Vouchers for Guyana.—The holotype;
Plate 6D AMNH A-166292–166293, A-166297–
166299, A-166302–166307, and USNM
Type material.—The holotype is ROM 566144–566148 from Konawaruk Camp;
39477, an adult female from the northeast IRSNB 13673 and ROM 20555 and 20559
plateau of Mount Ayanganna, Guyana from Tukeit, Kaieteur National Park;
(05824 0 N, 059857 0 W; elevation 1550 m). ROM 39450–39463 from Mount Ayan-
Distribution.—Known only from Guya- ganna; ROM 42861–42892 from Mount
na (MacCulloch et al. 2006). Wokomung; USNM 129530 from Karta-
bo; and USNM 531494–531499 from ous bands on the legs and arms, and
Iwokrama Forest Reserve, Sipuruni River, golden finger discs.
Pakatau Camp. Comments.—Stefania evansi is the larg-
Coloration in life.—Adults have been est member of the genus, up to 97.5 mm
reported (MacCulloch & Lathrop 2006d) snout-vent length, and the most widely
as usually having one of two dorsal color distributed. Up to 25 eggs or 30 juveniles
patterns, either uniform medium brown or on a single female have been reported
dark brown with cream dorsolateral and (Kok & Benjamin 2007, Kok et al. 2006c).
interorbital stripes. Juveniles may have For the AMNH series from Konawaruk
dark brown dorsal chevrons, but these Camp, the following data may someday be
are rarely seen in adults. Black spots are on added to calculations of the relationship
the distal portion of the anterior surface of between female body size and number of
the thighs. The venter is white, often with eggs per clutch: AMNH A-166307, snout-
brown mottling. urostyle length (sul; basically the snout-
The following notes are paraphrased vent length, but using the firm posterior
from CJC’s field notes (1998) for the end of the urostyle bone as the rear point
AMNH and USNM series from Konawa- of the measurement) ¼ 66 mm, 16 eggs in
ruk Camp: Dorsum either brown or tan, at attached, intact mass on back; AMNH A-
times with a wide postocular tan stripe and 166306, sul ¼ 69 mm, 17 eggs in attached,
narrow tan interorbital line, and at times intact mass on back; AMNH A-166303,
with dark or light brown spots on the sul ¼ 72 mm, 19 babies in clutch, most
back; darker brown mask aside face having been detached in preservative;
(nostril to past ear but not evident on AMNH A-166304, sul ¼ 72 mm, 19 term
every individual); canthal and postocular embryos in attached, intact mass on back;
stripes (to beyond ear) darker brown yet; AMNH A-166305, sul ¼ 73 mm, 13 babies
arms somewhat lighter brown than body, that had become detached in preservative.
with few darker brown spots; lower legs
similar to arms, upper legs of ground color
similar to body, with a few more dark Stefania roraimae
brown spots than arms, and very subtle Duellman & Hoogmoed, 1984
brown bands on thighs; fingers and toes Plate 6F
tan, fingers being lightest; toe webbing
very dark brown; lips pale with few dark Type material.—The holotype is UG
brown spots and a yellow wash that [¼CSBD] 10, an adult female from Ro-
extends along the sides half way toward raima, Guyana, 1402 m.
hip or all the way to groin; upper half of Distribution.—Known only from Guya-
iris tan (metallic copper in sunlight), lower na, as follows: ‘‘Mounts Roraima, Ayan-
half bronze; venter cream, palest on throat ganna and Wokomung, Pakaraima
(which may or may not have a copper Mountains...at elevations of 1234–1550
wash), chest, abdomen, and, on some m’’ (MacCulloch & Lathrop 2006e:1–2).
individuals, with extensive gray smudging; We hypothesize that it also occurs in
yellow wash ventrolaterally (sometimes Venezuelan and Brazilian parts of Mount
extending to legs, abdomen, chest); pale Roraima, where similar habitats occur.
gray mottling beneath legs and arms (less Vouchers for Guyana.—The holotype;
so); grayish brown beneath feet; tip of ROM 39466–39469 from Mount Ayan-
snout with yellowish tan vertical stripe. A ganna; ROM 42836–42855 from Mount
male was noted to have extensive dark Wokomung; and USNM 549313–549314
brown mottling on the dorsum, conspicu- from Mount Roraima.
Coloration in life.—‘‘The dorsum is dark brown mottling. This species is similar in

brown or greyish brown. The top of the appearance to Stefania evansi, but smaller.
head and loreal region are golden brown Comments.—Stefania scalae was synon-
speckled with orange. The canthus rostra- ymized with S. evansi by Duellman &
lis is outlined in black. The upper lip is Hoogmoed (1984) but was restored to
mottled in black, with one or two irregular species status by Señaris et al. (1997).
bars extending to the margin of the eye in
some individuals. A black supratympanic
bar extends halfway down the length of the Stefania woodleyi Rivero, 1968b
body. Creamy yellow dorsolateral stripes,
continuous with an interorbital bar of Type material.—The holotype is BMNH
similar color, extend to the groin. Flanks 1967.654, a female (MacCulloch & Lath-
are burnt orange fading to brown near rop 2006f:1) from a rocky stream on the
belly. Belly, throat and thighs are light to slope of Mt. Kanaima, near the Potaro
medium brown or grey with white spots River, Guyana.
which are most numerous on throat and Distribution.—Known only from Guya-
chest, and reduced to a few irregularly na, in the ‘‘eastern portion of the Paka-
shaped spots on thighs, arms and flanks. raima Mountain region...at elevations
Indistinct transverse bars are present on from 200 to 870 m’’ (MacCulloch &
dorsal surfaces of shanks and forearms. Lathrop 2006f:2).
The top third of the iris is golden, the Vouchers for Guyana.—The holotype;
lower two-thirds dark red. Juvenile color- ROM 20570 from Tukeit; ROM 28425–
ation resembles that of adults by day. At 28426 and USNM 561836 from Para-
night juveniles are uniformly yellow-green, makatoi; ROM 39465 from Mount Ayan-
with only the outline of the dorsolateral ganna; and ROM 42832–42835 from
and interorbital stripes visible’’ (MacCul- Mount Wokomung.
loch & Lathrop 2006e: 1). Coloration in life.—‘‘Dorsal surfaces are
Comments.—‘‘No information is avail- medium brown with irregular ochre spots
able on reproduction’’ (MacCulloch & or reticulations. Some individuals have an
Lathrop 2006e:2). ochre or cream interorbital bar and
dorsolateral stripes. Laterally, the head is
ochre with a dark brown canthal stripe and
Stefania scalae Rivero, 1970 irregular medium brown spots in loreal
Plate 7A and temporal regions. A dark brown
supratympanic stripe continues to the
Type material.—The holotype is MCZ groin. Flanks and groin are medium to
64373, a female from the region of La dark brown with irregular ochre spots.
Escalera, road between El Dorado and Dorsal surfaces of thighs and arms are
Sta. Elena de Uairén, around 1200 m, ochre with dark brown transverse bars
Bolivar, Venezuela. continuing onto anterior surfaces of
Distribution.—Known only from Guya- thighs; posterior surfaces of thighs are
na and Venezuela, in the uplands and medium brown with white spots. Shanks
highlands of the Pakaraima Mountains. and tarsi are ochre with dark brown spots
Voucher for Guyana.—ROM 39470 or bars. Throat and ventral surfaces of
from Mount Ayanganna. thighs are medium brown with cream to
Coloration in life.—Dorsum medium ochre mottling; ventral surfaces of shanks,
brown with scattered dark brown spots tarsi and arms are immaculate medium
and dark brown canthal and supratym- brown. The colour pattern on the chest
panic bars. Venter white, possibly with and belly varies from medium brown with
ochre mottling to ochre with brown mot- These are the frogs most commonly
tling...Juvenile coloration is similar to that referred to as tree frogs, most of which
of adults’’ (MacCulloch & Lathrop spend much of their lives in trees or
2006f:1). bushes, and they are exceptional climbers.
Comments.—‘‘Information on habitat Some of the species are so well adapted to
and reproduction is in MacCulloch & arboreal life (e.g., Trachycephalus resin-
Lathrop (2002), MacCulloch et al. (2006), ifictrix) that they can exist high in rain
and Rivero (1968b)’’ (MacCulloch & forest trees without coming to the ground,
Lathrop 2006f:2). as they usually breed in treeholes that are
filled with water, in which the tadpoles
complete their larval life.
Hylidae
The family Hylidae was named about Dendropsophus brevifrons
(Duellman & Crump, 1974)
100 yr ago (Rafinesque 1815). Its content
(genera and species) has changed consid- Type material.—The original name was
erably over time, and recent name-chang- Hyla brevifrons and the holotype is KU
es, in order to reflect relationships of 126370, an adult male from Santa Cecilia,
monophyletic groups, have occurred as a Provincia Napo, Ecuador.
result of modern phylogenetic analyses Distribution.—‘‘The upper Amazon Ba-
based primarily, although not exclusively, sin in Ecuador, Perú, and probably west-
on DNA sequence data (e.g., Faivovich et ern Brazil, to elevations of about 1300 m
al. 2005, Frost et al. 2006). Consequently, on the eastern slopes of the Andes.
significant recent changes in familial and Additionally, the species is known from
generic names include the following: 1) French Guiana and Belém, Brazil; it
many species that previously were included probably occurs throughout the Amazon
in the genus Hyla have been transferred to Basin’’ (Duellman & Crump 1974:17);
other genera (Dendropsophus, Hypsiboas, Frost (2011) also included Amazonian
Myersiohyla, Scinax); 2) species previously Bolivia and Colombia.
in the genus Ololygon are now in Scinax; 3) Vouchers for Guyana.—USNM 566060
species previously in the genus Phrynohyas from Konawaruk Camp.
are now Trachycephalus; 4) frogs of the Coloration in life.—‘‘The dorsum is
olive-tan with brown markings and, in
genus Pseudis previously referred to the
females, distinct creamy tan dorsolateral
family Pseudidae are now in the Hylidae,
stripes. The thighs are dark brown; the
including Lysapsus laevis (previously re-
spots on the thighs and the suborbital bars
ferred to as Lysapsus limellus or Pseudis are creamy yellow. The venter is unpig-
laevis); and 5) the genus Stefania has been mented except for the pectoral region and
transferred to a different family, the Hemi- chin (in females), which is white; males
phractidae (see above). In addition, a have a pale yellow vocal sac. The iris is red
modern phylogenetic analysis on the char- medially and silvery gray peripherally’’
ismatic poster frogs, or Phyllomedusinae, (Duellman & Crump 1974:17), based on
has provided ‘‘a historical framework for a frogs from Ecuador. The following is
discussion of the evolution of characters added from Lescure & Marty (2000:156),
associated with reproductive biology, glid- based on frogs from French Guiana.
ing behavior, the physiology of water- Dorsum beige to brown; cream spot in
proofing, and bioactive peptides’’ groin; iris gray to chestnut, with a narrow
(Faivovich et al. 2010:227). edge of red at the pupil.
Dendropsophus grandisonae (Goin, 1966a) Vouchers for Guyana.—AMNH A-

164124–164133 and USNM 531425–
Type material.—The original name was 531434 from Iwokrama Forest Reserve
Hyla grandisonae and the holotype is (several localities); and USNM 129528–
BMNH 1938.10.3.25 from the forest at 129529 from Kartabo.
Mazaruni, British Guiana. Coloration in life.—The following is
Distribution.—Known only from the from Lescure & Marty (2000:138) based
type locality. on frogs from French Guiana. Dorsum
Vouchers for Guyana.—We are not with light cream to yellowish tan vividly
aware of any specimens other than the contrasting with chestnut or chocolate
holotype. brown; arms and legs with similar colors;
Coloration.—‘‘Color in alcohol. A very flanks chestnut; ventral surface and hidden
strikingly marked little frog. The dorsal parts of legs bright reddish orange; iris
ground color is dark brown with the under reddish chestnut.
surfaces of the chin, throat, and body gray.
There is a rather distinct rounded white
spot on the occipital region, and the dorsal
Dendropsophus marmoratus
surfaces of the upper arms are clear white.
(Laurenti, 1768)
The patagium, while not white like the
Plate 7B, D
dorsal surfaces of the upper arms, is
unpigmented. The left heel has been Type material.—The original name was
injured, but there is a short white line on Bufo marmoratus and type(s) are frog(s)
the right heel’’ (Goin 1966a:40). This was from Surinam ‘‘illustrated as Rana surina-
published more than 25 yr after the frog mensis by Seba, 1734, Locuplet. Rer. Nat.
was collected. Thesaur. Descript. Icon. Exp. Univ. Phys.
Comments.—This is one of the most Hist., 1: pl. 71, figs. 4, 5’’ (Frost 2011).
poorly-known frogs in the world. Efforts Distribution.—‘‘Amazon Basin in Bra-
should be expended to collect additional zil. . ., Colombia, Ecuador, Peru, Bolivia;
specimens for comparative analyses and to southern Venezuela; Guianas’’ (Frost 2011).
learn about the natural history of this frog. Vouchers for Guyana.—AMNH A-
Faivovich et al. (2005:93) provisionally 18989 from Kamakusa; AMNH A-24914,
considered the species to be a member of A-25234, and A-70948 from Kartabo;
the parviceps species group; their comment USNM 102413 from Arakaka; USNM
(p. 156) about the microcephalus group 284388–284389 from Takutu Mountains
was an error (J. Faivovich pers. comm.). (6815 0 N, 059805 0 W); USNM 535749 from
Baramita; and USNM 566092 from Ko-
nawaruk Camp.
Dendropsophus leucophyllatus Coloration in life.—The following is
(Beireis, 1783) based on the USNM specimen from
Konawaruk Camp (CJC field notes,
Type material.—The original name was 1998). Gray above, with very dark gray
Ranam leucophyllatam and the holotype is markings; orange axilla and groin; chin
lost (Frost 2011). The type locality is white with gray smudging and tiny black
‘‘unknown, but believed by Beireis to be dots; chest yellow with black spots; abdo-
from Surinam’’ (Frost 2011). men, beneath upper arms, and beneath
Distribution.—‘‘The Amazon Basin (Co- upper legs orange with black spots; be-
lombia, Ecuador, Peru, Bolivia, and Bra- neath lower arms, hands, lower legs, and
zil) and the Guianas; expected but not yet feet black; basal webbing (hands and feet)
collected in Venezuela’’ (Frost 2011). black, distal webbing bright yellow on
fingers, bright orange on toes; iris light 7456 ‘‘and possibly two unlocated speci-
gray with black lines. Kok & Kala- mens’’ (Frost 2011). The type locality is in
mandeen (2008:160) added that the dorsal Brazil (Frost 2011).
color may be brown in part, the pattern of Distribution.—‘‘Lowlands east of Andes
coloration is ‘‘a network of dark lines and from Colombia, Venezuela, and Trinidad
markings that resemble lichens or bird southward through Ecuador, Peru and
droppings,’’ and the chest may be white Brazil to Bolivia, eastern Paraguay, Ur-
with black spots. uguay, and Argentina, up to 2000 m
elevation’’ (Frost 2011).
Dendropsophus minusculus (Rivero, 1971) 141033–141036, A-141038–141039, A-
Plate 7C 167097–167098, and USNM 566097 from
Dubulay Ranch; AMNH A-166209–
166211 from Konawaruk Camp. Also
Hyla minuscula and the holotype is UPRM
USNM 162854–162864 from Atkinson
3377, a male from Nirgua, Yaracuy,
(¼Timehri Airport); USNM 291171–
Venezuela.
291172 from Paramakatoi; USNM
Distribution.—‘‘Llanos of eastern Meta
531445–531454 from Iwokrama Forest
(Colombia) and lowland savannas of
Reserve (various localities); and USNM
southeastern Venezuela eastward through
535750–535759 from Baramita.
Guianas to Belém, Brazil’’ (Frost 2011).
Coloration in life.—The following is
based on AMNH and USNM specimens
139064–139067 from Aishalton; AMNH
from Dubulay Ranch and Konawaruk
A-141041–141045, A-141050–141051, A-
Camp (CJC field notes, 1994–1998). This
167099–167100, and USNM 566093–
species has considerable variation in dorsal
566096 from Dubulay Ranch. Also
pattern. Usually the dorsal body, arms,
USNM 497715 from Karanambo Ranch;
and lower legs are dark tan with conspic-
USNM 531435–531444 from Iwokrama
uous brown bands on the back. On some
Forest Reserve (various localities).
individuals, the dorsum is tan with very
subtle darker tan markings. One individual
was basically patternless dark tan above
from Aishalton and Dubulay Ranch (CJC
with scattered pale yellowish gold flecks.
field notes, 1993, 1994). Dorsum tan or
Thighs yellow mottled with tan; short
copper, with tiny dark brown dots and a
white or yellow bar atop vent and at each
broad darker brown or copper stripe on
ankle; all ventral surfaces grayish white to
sides (snout almost to groin), lined above
pale yellow, but some individuals translu-
with cream; arms and legs similar in color,
cent tan beneath arms and legs or yellow-
but thighs translucent yellow; hands and
ish green beneath thighs; chin with gray
feet (most inner fingers and toes) pale
smudges on some individuals; iris tan.
yellow to gold; throat and abdomen cream
Comments.—All three of the AMNH
to very pale yellow, but many males with a
specimens from Konawaruk Camp were of
touch of green in the vocal sac; arms and
the patternless morph (one having numer-
legs translucent below.
ous yellowish gold flecks), whereas those
from Dubulay Ranch were of the morph
Dendropsophus minutus (Peters, 1872) with the conspicuous pattern on the back.
Plate 7E, F Hawkins et al. (2007) reported that anal-
yses of DNA sequences from nine popula-
Type material.—The original name was tions sampled across much of the Guianan
Hyla minuta and the syntypes are ZMB Region revealed two significant lineages
that differed from each other by 9%. One The quoted latitude corresponds to a
occurs in the western highlands, one location approximately 35 km north of
spanned the whole area studied, and both Mt. Roraima and according to Warren
occur ‘‘in the area surrounding Imbaima- (1973) the latitude should be 05817 0 N.
dai, Guyana’’ (p. 63). The authors found Distribution.—Known only from high-
no differences in morphology, ecology, or lands of Guyana (MacCulloch & Lathrop
behavior of these frogs, which may be part 2005).
of a complex of cryptic species. Vouchers for Guyana.—The holotype; a
paratopotype, KU 182471; ROM 39487–
39496 from Mount Ayanganna; ROM
‘‘Hyla’’ helenae Ruthven, 1919 43500 from Mount Wokomung; and
USNM 549312 from the north slope of
Mount Roraima.
UMMZ 52681 from Dunoon, valley of
Coloration in life.—‘‘Dorsum ground
the Demerara River, Guyana.
colour greenish-grey with medium brown
Distribution.—Known only from the
blotches outlined in darker brown; fine
type locality (Frost 2011).
dark brown or black reticulations between
Vouchers for Guyana.—The type series
blotches on back. Dorsal dark blotches
included three adults from the type locality.
joined to form two irregular longitudinal
Coloration in life.—‘‘Above deep green
stripes in one female. Dorsal pattern of
with numerous well-defined brown spots;
diffuse brown reticulations and scattered
thighs, hind feet and inner side of tarsus
orange buff, tibiae javel green; fore limbs, dark brown spots on the ground color in
except elbows and inner side of feet, two males. Axilla and flanks reddish-brown
asphodel green, elbows viridine yellow, with dark brown blotches. Posterior thigh
inner side of fore feet orange buff; both surface uniform medium brown with small
fore limbs and hind limbs with light brown white spots. Venter white with reddish-
cross-bars; upper lip viridine yellow with brown mottling on throat and chest,
vertical brown bars; belly white, immacu- changing to small spots on belly. Underside
late. In alcohol the green has faded to a of limbs white with reddish-brown wash.
brownish olive, the light markings to dull Iris gold to pale green with fine radiating
white’’ (Ruthven 1919:11). black reticulations. Palpebral membrane
Comments.—Owing to the lack of spec- not reticulated, upper edge with a broad
imens, Faivovich et al. (2005) and Frost et black band containing tiny white specks’’
al. (2006) were unable to clarify the (MacCulloch & Lathrop 2005:24–25).
relationships of this species, so the generic Comments.—Owing to the lack of spec-
name was placed in quotes to indicate that imens, Faivovich et al. (2005) and Frost et
the taxon is known not to be a natural al. (2006) were unable to clarify the
monophyletic group and future research relationships of this species, so the generic
will result in a change in the generic name. name was placed in quotes to indicate that
the taxon is known not to be a natural
monophyletic group and future research
‘‘Hyla’’ warreni will result in a change in the generic name.
Duellman & Hoogmoed, 1992
Plate 8A
Hypsiboas boans (Linnaeus, 1758)
Type material.—BMNH 1979.561, an Plate 8B
adult female from the north slope of Mt.
Roraima (05838 0 N, 60844 0 W; elevation Type material.—The original name was
1480 m), Rupununi District, Guyana. Rana boans and the types, from unspecified
localities, have had a complicated history body length of about 132 mm, and males,
(summarized by Frost 2011). at least at some localities, are known to
Distribution.—‘‘Lower Amazon Basin, defend their nests in basin-like depressions
upper Orinoco and Magdalena Basins, that they construct near water (Kok &
Guianas, and Pacific lowlands of Colom- Kalamandeen 2008).
bia and adjacent Ecuador in South Amer-
ica; eastern Panama; Trinidad’’ (Frost
2011). Hypsiboas calcaratus (Troschel, 1848)
Vouchers for Guyana.—AMNH A- Plate 8C
70929–70942 and USNM 118051 and
129535 from Kartabo; AMNH A- Type material.—The original name was
141014–141018, USNM 497704, and Hyla calcarata in a book about Guyana
USNM 566056–566057 from Dubulay and based on a type or types ‘‘likely now
Ranch; AMNH A-166135–166137 from lost’’ (Frost 2011).
Berbice River Camp; AMNH A-166138– Distribution.—‘‘Guianas, Amazon Basin
166139 and USNM 566058–566059 from from southern Colombia and adjacent
Konawaruk Camp. Also ROM 22966 Ecuador south through Peru to Bolivia,
from Baramita; ROM 20556–20558 from and Orinoco Basin and delta in Venezuela
Kurupukari; ROM 28420–28421 and and Brazil’’ (Frost 2011).
USNM 291160 from Paramakatoi; USNM Vouchers for Guyana.—AMNH A-
284387 from Takutu Mountains (6823 0 N, 166143–166144 and USNM 566061 from
059805 0 W); USNM 291105 from Chiung Konawaruk Camp. Also USNM 531366–
River at Kato; USNM 291196 from 531375 from Iwokrama Forest Reserve
Kaieteur; USNM 497699–497703 from (several localities).
Kumu (3815 0 52 00 N, 059843 0 34 00 W); USNM Coloration in life.—The following notes
497811 from Karanambo; USNM 497812– are based on AMNH and USNM speci-
497813 from Moco Moco (3818 0 12 00 N, mens from Konawaruk Camp (CJC field
059818 0 36 00 W); and USNM 531356– notes, 1998). All dorsal surfaces very dark
531365 from Iwokrama Forest Reserve tan, with or without dark brown to black
(several localities). vertebral line from tip of snout to posterior
Coloration in life.—The following notes body; posterior sides, groin, and rear of
are based on AMNH and USNM speci- thigh light gray with chocolate brown bars;
mens from Dubulay Ranch (CJC field throat white; other ventral surfaces trans-
notes, 1994). Dorsum brown with cream lucent gray or tan; very narrow pale yellow
dots (plus darker brown spots and/or pale lines may be on upper and lower lips, rear
tan bands on some individuals); legs of arms, and above vent; feet and spurs on
brown with paler tan bands across thighs; ankles dark brown; iris tan.
throat and beneath hands greenish gray;
chest, abdomen and beneath thighs pale
yellow; yellowish gray to pale tan bars on Hypsiboas cinerascens (Spix, 1824)
sides; bones green. Kok & Kalamandeen Plate 8D
(2008:164) add the following for frogs
from Kaieteur National Park: ‘‘Ventral Type material.—The original name was
surface...white to greenish white, throat Hyla cinerascens and the syntypes (from
white to greenish white in females, greyish Brazil) are ZSM 2498/0, now lost (Glaw &
in males. Iris bronze, lower eyelid reticu- Franzen 2006).
lated with silvery gold.’’ Distribution.—‘‘Guianas and Amazon
Comments.—This is the largest species Basin in Brazil, Colombia, Venezuela,
of tree frog in Guyana, with a maximum Ecuador, Peru, and Bolivia’’ (Frost 2011).
Vouchers for Guyana.—AMNH A- USNM 497705–497714, and USNM

166182–166192 and USNM 566086– 566069–566072 from Dubulay Ranch;
566090 from Berbice River Camp; AMNH and AMNH A-145153 from Berbice River,
A-166198–166200 and USNM 566091 1.6 km downriver from Dubulay Ranch.
from Konawaruk Camp; ROM 22750– Also ROM 20560 from Kurupukari; ROM
22753 from Baramita; ROM 43208–43212 28436–28438 and USNM 291106 and
and USNM 291197–291211 from Kaie- USNM 291161–291168 from Paramaka-
teur; USNM 118053–118054 from Karta- toi; USNM 146365 from Lethem; USNM
bo; USNM 164178 from Mabaruma 162846–162853 from Atkinson (¼Timehri
Compound (8812 0 N, 059847 0 W); and Airport); USNM 291107–291108 from
USNM 531405–531424 from Iwokrama Kawa Valley east of Kato; USNM
Forest Reserve (several localities). 291109–291113 from Chiung River at
Coloration in life.—The following notes Kato; and USNM 531376–531380 from
are based on AMNH and USNM speci- Iwokrama Forest Reserve, Muri Scrub
mens from Berbice River Camp (CJC field Camp (4825 0 12 00 N, 058850 0 58 00 W).
notes, 1995). Dorsum light green, with Coloration in life.—The following is
conspicuous or inconspicuous yellow spots based on AMNH and USNM specimens
and possibly one or more irregular chest- from Aishalton and Dubulay Ranch (CJC
nut brown patches; chestnut brown bar field notes, 1993, 1994). Awake at night in
often on head between eyes; chestnut the field, the dorsum is brown with darker
brown horseshoe-shaped mark often on brown markings. By day these frogs are
snout; arms and legs pale green; ventral very pale gray or green on the back; rear of
surfaces translucent bluish green; iris pale thigh with bars of pale yellow and orangish
coppery silver. Kok & Kalamandeen yellow; sides with translucent orangish
(2008:168) mentioned ‘‘outer edge of upper yellow bars; hands and feet orange above
eyelid yellow,’’ and referred to iris color as and below; throat extremely pale green;
‘‘light orange to reddish orange.’’ chest, abdomen, underside of hind legs and
Comments.—Until recently, this species feet orange; bones green.
was referred to as Hypsiboas granosus.
Kok (2006b) stated that there may be two
or more cryptic species currently referred Hypsiboas fasciatus (Günther, 1858)
to as H. cinerascens.
Hypsiboas fasciata and the holotype from
Hypsiboas crepitans (Wied-Neuwied, 1824) the Andes of Ecuador is BMNH 58.4.25.22
Plate 8E, F (Frost 2011).
Distribution.—‘‘Guianas and upper Am-
Type material.—The original name was azon Basin from southeastern Colombia
Hyla crepitans and the lectotype is AMNH and eastern Ecuador through eastern Peru
785 from a locality restricted to Tamburil, to Bolivia; possibly to be found in south-
Bahia, Brazil (Frost 2011). ern Venezuela’’ (Frost 2011).
Distribution.—‘‘Relatively dry habitats Vouchers for Guyana.—AMNH A-
in eastern Panama,. . .Colombia, Venezue- 122257–122258 from Isheartun, identified
la, the Guianas. . .northern Brazil, below by M. S. Hoogmoed; and USNM 531381–
1700 m elevation’’ (Frost 2011). 531384 from Iwokrama Forest Reserve,
Vouchers for Guyana.—AMNH A- Cowfly Camp (4820 0 15 00 N, 058849 0 37 00 W).
139041–139054, AMNH A-139060– Coloration in life.—The following is
139061, and USNM 566062–566068 from from Lescure & Marty (2000:128), based
Aishalton; AMNH A-141019–141026, on frogs from French Guiana. Dorsal
surface grayish brown, yellowish brown, thigh as dorsal body or gray with lighter
or chestnut, with subtle cross bands and gray spots; hands and feet mostly yellow-
small black spots; vertebral area may have ish tan; lips pale yellow, lower ones with
black spots or a stripe. Flanks and anterior gray speckling; throat very pale yellow;
and posterior surfaces of thighs brownish anterior chest cream; orange on abdomen
gray, lightly tinged with black spots. and beneath thighs, hands, and feet,
Ventral surface whitish, translucent. Iris brighter on males than females; brown
white to dull silver. flecks on abdomen; cream to pale yellow
beneath arms; dark brown middorsal
stripe from snout to midbody; some
Hypsiboas geographicus (Spix, 1824) individuals with several very pale yellow,
Plate 9A lichen-like round spots on back; iris
copper; bones green. Juveniles can look
Type material.—The original name was rather different from adults. AMNH A-
Hyla geographica and the holotype is ZSM 166212 was young (SVL ¼ 30 mm) and was
35/0 from Brazil but now lost (Glaw & noted as being tan above with very subtle
Franzen 2006). darker tan markings; lower flanks black;
Distribution.—Very widely distributed anterior parts of the hands and feet black,
in ‘‘tropical South America east of the but yellowish orange beneath innermost
Andes, excluding coastal region from fingers and toes; webbing dark gray; black
Alagoas to Santa Catarina, Brazil; includ- along anterior and posterior edges of the
ing Trinidad’’ (Frost 2011). hind legs; throat and chest cream with gray
Vouchers for Guyana.—AMNH A- smudging; abdomen cream with yellowish
70943 from Kartabo; AMNH A-141052– orange smudge; gray beneath arms; yel-
141054, AMNH A-141056, AMNH A- lowish orange beneath thighs; iris copper.
167108, USNM 497817, and USNM
566073 from Dubulay Ranch; AMNH A-
166146–166157 and USNM 566074– Hypsiboas lemai (Rivero, 1972)
566079 from Berbice River Camp; and
AMNH A-166164–166175, AMNH A- Type material.—The original name was
166212–166213 (juveniles), and USNM Hyla lemai and the holotype is UPRM
566080–566085 from Konawaruk Camp; 3179, a female from ‘‘Paso del Danto, La
ROM 28439–28449 and USNM 291169– Escalera, entre El Dorado y Sta. Elena de
291170 from Paramakatoi; USNM 163071 Uairén, Serranı́a de Lema,’’ Bolivar, Ven-
from Takama; USNM 291114 from 3.2 km ezuela, 1300–1400 m (Rivero 1972:183).
E of Kato; USNM 531385–531404 from Distribution.—Eastern Venezuela, in-
Iwokrama Forest Reserve (several locali- cluding Mount Roraima and Guyana
ties); and USNM 564214 and 564218 from (Frost 2011).
the summit of Mount Kopinang Vouchers for Guyana.—ROM 39567–
(5800 0 08 00 N, 059852 0 47 00 W, ca. 1538 m 39571 from Mount Ayanganna (MacCul-
elevation). loch & Lathrop 2005:37).
Coloration in life.—The following is Coloration in life.—‘‘Dorsum medium
based on AMNH and USNM specimens brown, with dense melanophores on back
from Dubulay Ranch, Berbice River and upper surfaces of limbs, extending
Camp, and Konawaruk Camp (CJC field onto fingers III and IV, and toes IV and V;
notes, 1994–1998). Upper surfaces orang- diffuse black interorbital and canthal lines
ish tan to brown with dark brown or light and a faint black supratympanic line.
yellowish tan bands; flanks gray with Palpebral membrane unpigmented except
cream dots and bars; dorsal surface of for a thin dark brown line along upper
edge. Limbs with dark brown crossbands. Hypsiboas multifasciatus (Günther, 1859)
Ventral surfaces white. Diffuse whitish Plate 9C
patches present on dorsum and limbs of
one individual, similar to the photograph Type material.—The original name was
Hyla multifasciata and the holotype, from
in Rivero (1972:189). Scattered white spots
Belém, Pará, Brazil is BMNH 1947.2.23.6
present on dorsum in three specimens; one
(Condit 1964:93, Frost 2011).
of these also has white dorsolateral lines
Distribution.—‘‘Lowlands of eastern
extending almost to the groin, and anteri-
Venezuela, through the Guianas to Ceará
orly to the nostrils’’ (MacCulloch &
and Goiás, Brazil, limited by the Amazon,
Lathrop 2005:25–26). the Orinoco, and Rio Negro’’ (Frost 2011).
49360–49363 from Isheartun; AMNH A-
Hypsiboas liliae Kok, 2006b 141040 and A-167116–167118 from Ceiba
Plate 9B Biological Station, Madewini River, ca. 4.8
km (linear) east of Timehri Airport
Type material.—The holotype is IRSNB (06829 0 57 00 N, 058813 0 06 00 W); AMNH A-
1965, an adult male from between Boy 145154 from Dubulay Ranch; AMNH A-
Scout View and Johnson View on the 166203–166206 and USNM 566098–
Kaieteur Plateau (5810 0 51 00 N, 59828 0 566099 from Konawaruk Camp; ROM
57 00 W), 400 m elevation, Kaieteur National 22755–22759 and USNM 535760–535761
Park, Guyana. from Baramita; ROM 28435 and USNM
Distribution.—Endemic to Guyana, 291173–291176 from Paramakatoi;
known from only two localities. USNM 118043 from Bartica; USNM
Vouchers for Guyana.—IRSNB 1965– 118049–118050 from Kartabo; USNM
1968 from the Kaieteur plateau, and UTA 291116–291121 from Chiung River at
Kato; USNM 497716 from Moco Moco
A-53052–53054 from scrubby mountains
(3818 0 12 00 N, 059839 0 00 00 W); UTA A-55292–
NE of Imbaimadai, Mazaruni-Potaro,
55293 from Mabaruma, ridge N side of
Guyana.
town, NW District, and UTA A-55294–
Coloration in life.—‘‘Colouration is
55295 from Mabaruma, along road NE of
highly dependent on light intensity. By
town ca. 2.0 km, NW District.
day: dorsal surfaces bright yellowish green; Coloration in life.—The following notes
irregular brown blotch between the eyes; are based on AMNH and USNM speci-
throat and venter blue; central portion of mens from Ceiba Biological Station, Du-
abdomen translucent, white peritoneum bulay Ranch, and Konawaruk Camp (CJC
visible through ventral skin; posterior part field notes, 1994–1998). Tan above (body,
of flank yellowish orange; ventral surfaces arms, legs) with darker tan bands and a
of limbs translucent greenish blue; web- brown face mask (snout through tympa-
bing, fingers and disks greenish orange; num); a gray stripe over the tympanum
palm greenish orange; sole greenish blue; fades quickly posteriorly; gray stripe
upper eyelid yellow; iris silver with black around chin; darker tan to brown on
periphery; bones green...At night: dorsum posterior surface of thighs and on feet;
and dorsal surfaces of hind limbs greenish dark brown on lateral edge of forearm and
brown and iris bronze. At intermediate lower leg; anal region black with cream to
light levels, the granules may be greenish orangish yellow stripe above vent; under-
brown and brown dots may appear on sides yellow to orangish yellow, brightest
snout, lips, and upper surfaces of arms and on throat and chest, the throat having
legs’’ (Kok 2006b:195–196). numerous tiny brown dots, but beneath
arms and legs, tan; iris copper or mixed Coloration in life.—The following is
gold and copper. from Lescure & Marty (2000:136) based
on frogs from French Guiana. Dorsal
surfaces yellowish green by day, vermillion
Hypsiboas ornatissimus (Noble, 1923) at night; small yellow or sometimes ver-
million spots on back; thin dorsolateral
Type material.—The original name was folds vermillion, edged with yellow; ventral
Hyla ornatissima and the holotype from surfaces bluish gray but belly yellow; iris
Meamu, Mazaruni River, Guyana is grayish.
AMNH A-13491, an adult female.
Distribution.—‘‘Guianas and Amapá,
Brazil, to Guainı́a, Colombia’’ (Frost Hypsiboas roraima
2011). (Duellman & Hoogmoed, 1992)
Vouchers for Guyana.—The holotype. Plate 9E, F
from Lescure & Marty (2000:134) based Type material.—The original name was
on frogs from French Guiana. Dorsum Hyla roraima and the holotype is BMNH
light green to yellowish green, with small 1979.560, an adult female from the north
reddish brown to yellow spots; dark slope of Mt. Roraima (05838 0 N, 60844 0 W;
reddish brown bar edged with yellow elev. 1480 m), Rupununi District, Guyana.
According to Warren (1973) the latitude
across top of head from one eyelid to the
should be 05817 0 N, as the latitude quoted
other; similarly colored longitudinal verte-
corresponds to a location some 35 km
bral stripe on body, split into a circle or
north of Roraima.
diamond shape in the middle; dark reddish
Distribution.—All known specimens are
brown spot on nostrils; reddish brown
from Guyana, but the species probably
spots on arms and legs; venter greenish
also occurs in the Venezuelan part of
yellow; iris reddish brown near pupil, with Mount Roraima (MacCulloch & Lathrop
black outer edge. 2005, MacCulloch et al. 2007).
Vouchers for Guyana.—The holotype;
the paratypes, KU 182470 and UGDB 14
Hypsiboas punctatus (Schneider, 1799) from Mount Roraima; and ROM 39596–
39627 and USNM 561830–561831 from
Type material.—The original name was Mount Ayanganna (MacCulloch & Lath-
Calamitae punctatus. The syntypes from rop 2005:37).
Surinam are at ZUU and ZISP (Frost Coloration in life.—The following is
2011). based on ROM specimens from Mount
Distribution.—‘‘The Amazon Basin of Ayanganna (MacCulloch & Lathrop
Ecuador, Peru, Bolivia, and Brazil and the 2005:26). ‘‘Dorsal ground colour medium
Orinoco Basin...; Chaco of Paraguay, and reddish-brown with a pattern varying
Argentina; Guianas, northern Colombia in between two extremes: the first having
the Sierra Santa Marta, Venezuela (Ama- four dark brown patches—on top of
zonas, Apure, Bolı́var, and Sucre), and snout, between orbits, in scapular region,
Trinidad’’ (Frost 2011). and in sacral region; the second having
Vouchers for Guyana.—UMMZ 83552 only a few small scattered dark brown
from Yarikita River (2 specimens) and marks. Dark brown middorsal stripe
UMMZ 83553 from Yarikita River mouth from tip of snout to posterior dorsum
(4 specimens), identified by M. S. Hoog- present in 12 individuals; usually, but not
moed. always, extending to the groin; middorsal
stripe never present on individuals with and V, and Finger IV. Diffuse larger dark
dorsal pattern of dark patches. Colour brown patches, made up of a concentra-
pattern not correlated with sex or size. tion of minute spots, between orbits, on
Scattered dorsal white spots present in dorsum or on shanks in some individuals.
some individuals. Limbs with dark brown Minute dark brown spots on the edge of
crossbands, usually most prominent on the lower jaw below tympanum and eye,
hind limbs; crossbands reduced to small extending anteriorly to form a continuous
dark brown spots in some individuals. line along edge of lower jaw in a few
Axilla pink, flanks pink with dark brown individuals. Scattered round white dorsal
bars or spots and white spots posteriorly spots in several individuals. Pale yellow
in some individuals. Rear of thigh pale interorbital and dorsolateral lines present
brown with small dark brown spots. in two males. Digital discs and iris orange;
Upper lip yellow. Venter white; under- limb bones green. In preservative the
sides of forearms, hands, tarsi and feet dorsal ground color is white and the
shading to reddish-brown. A row of dark interorbital and dorsolateral lines are
spots along underside of tarsus and onto barely visible.’’ Kok & Kalamandeen
toe V. Iris pale green to yellow, pupil (2008:174) presented the ventral colora-
horizontal. Palpebral membrane with tion as ‘‘pale green to bluish green,
white reticulations, its upper edge with translucent in the central portion of
a brown-and-white speckled band.’’ abdomen (internal organs visible).’’
Hypsiboas sibleszi (Rivero, 1972) Lysapsus laevis Parker, 1935

Plate 10A, B Plate 9D
Type material.—The original name was Type material.—The original name was
Hyla sibleszi and the holotype is UPRM Pseudis laevis and the holotype is a male,
3177, a male from Paso del Danto, La BMNH 1947.2.25.85 (Condit 1964) from
Escalera, ‘‘entre El Dorado y Sta. Elena de the Rupununi Savannah, Guyana.
Uairén, 1300–1400 m; Serranı́a de Lema,’’ Distribution.—‘‘Southwestern Guyana
Bolı́var, Venezuela (Rivero 1972:182). and adjacent northern Brazil’’ (Frost
Distribution.—‘‘Moderate [to high] ele- 2011).
vations (900-1850 m) in highlands in Vouchers for Guyana.—The holotype;
eastern Venezuela and adjacent Guyana’’ AMNH A-136028 and USNM 497779–
(Frost 2011). 497788 from Karanambo; AMNH A-
Vouchers for Guyana.—ROM 39552– 139159–139170, A-139178–139181, and
39566 and USNM 561832 from Mount USNM 566237–566243 from Aishalton.
Ayanganna; ROM 28450 from Paramaka- Coloration in life.—The following is
toi; ROM 43229–43232 from the Kaieteur based on AMNH and USNM specimens
plateau; and ROM 43408–43409 and from Karanambo and Aishalton (CJC
43414 from Mount Wokomung. field notes, 1992, 1993). Pale olive green
Coloration in life.—The following is above (tan after immersion in chloretone),
based on ROM specimens from Mount often with gray, brown, or black flecks
Ayanganna (MacCulloch & Lathrop forming an irregular pattern; some indi-
2005:29). ‘‘Dorsum greenish-yellow at viduals with a rust or dark tan vertebral
night, green by day. Both sexes have stripe; tiny cream dots around anal region
small brown chromatophores of varying and rear of thigh; cream stripe on side of
intensity on the dorsum and along upper body, broken or not; arms similar to body;
surfaces of limbs, extending onto Toes IV legs paler, with darker bands of body
color; chin and throat pale green or small white pustules on posterior thigh
greenish yellow with cream or yellow dots; surface. Axilla white with a reddish wash.
chest and abdomen pale yellowish cream Venter white with small brown spots on
with white or cream dots extending rear- throat, variable diffuse brown blotches on
ward to hind legs; arms and legs translu- chest and belly. Palpebral membrane with
cent below; bones green. white reticulations, upper edge with a
Comments.—This aquatic species occurs white band outlined in black. Iris black.
in savannas, and in the recent past was In preservative dorsal ground color tan to
referred to as Pseudis laevis. medium brown’’ (MacCulloch & Lathrop
2005:30).
Myersiohyla kanaima
(Goin & Woodley, 1969) Osteocephalus buckleyi (Boulenger, 1882)
Plate 10C, D
Type material.—The original name was Hyla buckleyi and the lectotype is BMNH
Hyla kanaima and the holotype is BMNH 1947.2.13.44 with the type locality restrict-
1965.230, an adult female from Mt. ed to Canelos, Ecuador (Frost 2011).
Kanaima, near Amatuk Falls on the Distribution.—‘‘An arc around the pe-
Potaro River, Guyana, at 2300 ft eleva- riphery of the Amazon Basin from Co-
tion. lombia to east-central Bolivia, the Guianas
Distribution.—Known only from Guya- and northeastern Brazil (Amapá), at ele-
na (MacCulloch & Lathrop 2005). vations less than 700 m, and to the
Vouchers for Guyana.—The holotype; Orinoco Delta of Venezuela’’ (Frost 2011).
the paratypes; KU 182469 from Mount Vouchers for Guyana.—AMNH A-
Kanaima; ROM 39272–39277, ROM 13492, A-144983, and A-70971 from Kar-
39279–39295, ROM 39580, ROM 39584– tabo; AMNH A-46233 from Marudi;
39585, ROM 39592, and USNM 561828– AMNH A-46283 from Kuyuwini Landing;
561829 from Mount Ayanganna; and and AMNH A-49252 from Shudikar-wau,
USNM 549311 from the north slope of mostly identified by W. E. Duellman and
Mount Roraima. confirmed recently by CJC. Also UTA A-
Coloration in life.—‘‘Dorsal background 51630–51632 from 9.7 km (by road) S
golden to medium brown with dark brown Bartica, near air strip, identified by Smith
or black longitudinal lines. In many & Noonan (2001:356).
individuals the dark lines form a ‘‘)(’’ Coloration in life.—The following, based
dorsal pattern, with additional scattered on frogs from Kaieteur National Park, is
black marks and narrow canthal and from Kok & Kalamandeen (2008:180).
interorbital lines; dorsal lines are broken ‘‘Dorsal ground colour variable, from
and/or irregular in some individuals. Dark greenish brown to green, with dark brown
dorsal lines not present in juveniles. Upper markings often in the shape of a black
eyelid and snout with small black specks. inverted triangle between the eyes followed
Scattered dorsal white spots present in on the back by a ( ) or a X and two large
some individuals. Flanks white to pale dark lumbar spots; ventral surface...grey-
yellow with irregular black spots increas- ish white, with brown blotches located
ing in size posteriorly. Limbs with irregular mainly on throat, chest, and sides of belly;
black crossbands, most pronounced on ventral surface sometimes entirely covered
dorsal surface of thighs, and small spots by brown flecks. Broad, irregular, green
extending onto outer fingers and toes. subocular spot...Iris greenish bronze to
Crossbands fade to fine mottling with gold without radiating black lines, but
with irregular black vermiculations...ingui- (or very pale tan) with distinct tan
nal region and inner thigh blue.’’ markings; tan beneath legs; posterior
Comments.—See comments in account lateral body (anterior to groin) with black
for O. oophagus. In addition, in a recent bars or other markings alternating with
review of frogs of this group, Jungfer pale green; tympanum translucent copper;
(2010:46) stated that ‘‘O. buckleyi was iris greenish gold; bones green. Lescure &
believed to be a widespread Amazonian Marty (2000:182) pointed out that there
and Guianan species. . .but in fact it is a may be blue on the axillary membrane, in
complex of several species. Four species in the groin, and on the posterior surface of
the buckleyi complex are dealt with here.’’ the thigh.
However, Jungfer did not include any Comments.—In a recent review of frogs
specimens of O. buckleyi from the Guiana of this group, Jungfer (2010:33) stated that
Shield in his list of specimens examined. ‘‘Osteocephalus cabrerai is known
This would indicate that the specimens from. . .Colombia. . .and Peru. . .There are
from the Guiana Shield need further study also specimens from the Guiana Shield
to determine whether they are in fact this and the Orinoco Delta. . .that I refrain
species or an as yet unrecognized cryptic from including in the present study.’’ This
species. would indicate that the specimens from the
Guiana Shield need further study to
determine whether they are in fact this
Osteocephalus cabrerai species or an as yet unrecognized cryptic
(Cochran & Goin, 1970) species.
Plate 10E
Type material.—The original name was Osteocephalus exophthalmus
Hyla cabrerai and the holotype is USNM Smith & Noonan, 2001
152759 from Caño Guacayá, a tributary of
the lower Rio Apaporis, Amazonas, Co- Type material.—The holotype is UG/
lombia. CSBD HA-722, adult male cranium of
Distribution.—‘‘Amazonian Colombia UTA A-51620 from about 30 km SE
and Ecuador as well as adjacent north- Imbaimadai, Mazaruni-Potaro District,
eastern Peru. . .; French Guiana and likely Guyana, about 585 m (5837 0 30 00 N,
in adjacent Brazil and Surinam’’ (Frost 60814 0 42 00 W).
2011). Distribution.—Known only from Guya-
Vouchers for Guyana.—AMNH A- na (Kok & Kalamandeen 2008).
166220–166229 and USNM 566104– Vouchers for Guyana.—The holotype;
566108 from Konawaruk Camp. Also and IRSNB 14644, 14662, and 14664–
UTA A-51621–51623 from ‘‘Imbaimadai, 14665 from Kaieteur National Park.
ca. 0.8 km NW Mazaruni camp’’ (Smith & Coloration in life.—‘‘Light Buff (124)
Noonan 2001:356). dorsum with suffusions of Raw Sienna
Coloration in life.—The following is (136); loreal area Raw Sienna; area below
based on the AMNH and USNM speci- canthus rostralis and dorsal blotches and
mens from Konawaruk Camp (CJC field bars Vandyke Brown, somewhat dark
notes, 1998). Dorsum and arms mottled edged; flanks Jet Black (89) with light Buff
very dark brown and moss green to light areolations; anterior area of insertion of
brown (but pale green on sides and anal limbs Jet Black; iris dark Buff with Jet
area); legs and sometimes arms with same Black cross radiation from black pupil;
colors but patterned in bands; chin, throat, area below eye cream-white with small
chest, beneath arms and abdomen white Vandyke Brown markings; Jet Black col-
oration behind posterior corner of eye, Smith & Noonan (2001:356) identified the
bordering transparent tympanum, and following specimens as this species:
continuing to area of insertion of fore- AMNH A-70967–70968 from Kartabo;
limbs; upper area of lower arms dark Buff UTA A-51618 from ‘‘Imbaimadai, ca 100
(24); hand webbing transparent; anterior m W Mazaruni camp’’; and UTA A-51619
area of thighs, foot webbing, and anal from ‘‘Imbaimadai, Mt. NE Imbaimadai.’’
patch Sepia (119), almost black; white Coloration in life.—The following is
coloration surrounding anal patch, includ- based on AMNH and USNM specimens
ing tubercles, and extending as lines of from Berbice River Camp (CJC field notes,
posterodorsal reticulations to about mid 1997). Brown above with darker brown
thigh; posterior area of thighs Jet Black; markings; dark brown canthal stripe or
axillary and groin area Pearl Gray (81); face mask; anterior and posterior surfaces
throat white with Sepia (119) spotting, of thighs orange; varying degrees of
stronger on sides; chest, belly and arm translucent orange on ventral surfaces of
ventral areas Drab-Gray (119D); chest legs and toe webbing; belly varies from
with fine Natal Brown (219A) spotting on light to bright yellow; iris gold and black
sides near insertion of limbs; palms Olive on upper half, brown below. Kok &
Brown (28); ventral surface of thighs Olive Kalamandeen (2008:184) referred to the
Brown (28); ventral shank and tarsal areas dorsum as ‘‘ranging from ochre, tan or
pearl Gray; sole of feet Hair Brown reddish tan to dark tan, usually with up to
(119A); jaw and hind limb bones Emerald four narrow, usually fragmented, dark
Green (163)’’ (Smith & Noonan 2001:351– brown transverse lines, and a narrow dark
352). The numbers associated with colors brown interorbital bar.’’
in the quote refer to ‘‘the Naturalist’s Color
Guide (Smithe 1975)’’ (Smith & Noonan
2001:348). Osteocephalus oophagus
Jungfer & Schiesari, 1995
Plate 11A
Osteocephalus leprieurii
(Duméril & Bibron, 1841) Type material.—The holotype is
Plate 10F MZUSP 69852, an adult male from
Reserva Florestal Adolfo Ducke (2855 0 S,
Type material.—The original name was 59859 0 W), km 26 on the Rodovia AM-010
Hyla leprieurii and the holotype from (Manaus-Itacoatiara), Amazonas, Brazil.
Cayenne, French Guiana is MNHN 4629 Distribution.—‘‘The species is found in
(Guibé 1950). the Guiana Shield and in the Amazon
Distribution.—‘‘It occurs on the Guiana Basin, from east and north of State of Pará
Shield in French Guiana, Surinam, Guya- to eastern Colombia’’ (Kok & Kala-
na, Venezuela, and northern Brazil south- mandeen 2008:186).
ward at least to the Rio Negro and Rio Vouchers for Guyana.—AMNH A-
Uaupés’’ (Jungfer & Hödl 2002:34). 166214–166216 and USNM 566102 from
Vouchers for Guyana.—AMNH A- Berbice River Camp; AMNH A-166219
46387 from Onora; AMNH A-70967– and USNM 566103 from Konawaruk
70968, A-70976, and USNM 118052 from Camp; and ROM 44202–44230 from
Kartabo; AMNH A-166235–166239, A- Kaieteur.
166247–166254, and USNM 566109– Coloration in life.—The following is
566115 from Berbice River Camp; AMNH based on the AMNH and USNM speci-
A-166255 from Konawaruk Camp; and mens from Berbice River Camp (CJC field
ROM 42997 from Kaieteur. In addition, notes, 1997). Dorsum brown or grayish tan
with dark brown markings; all ventral reticulations. Rear of thigh brown with
surfaces tan, chin, throat, and chest with small white spots. Venter white with dark
darker tan markings; iris gold, with brown reticulations on throat and chest.
contrasting black radiating lines. Kok & Palms and soles with scattered melano-
Kalamandeen (2008:186) stated that ‘‘in- phores. Long limb bones green. Iris gold
dividuals with white spots [on the dorsum] with dense black reticulations and a
are reported.’’ median horizontal brown bar’’ (MacCul-
Comments.—This species is smaller than loch & Lathrop 2005:22).
O. taurinus, which also has the alternating
gold and black radiating lines on the iris,
but males of oophagus have a single, Osteocephalus taurinus Steindachner, 1862
median vocal sac, not paired sacs as in Plate 11C
taurinus. There are no supraorbital tuber-
cles and there are small tarsal tubercles, Type material.—The holotype is
but each of the latter usually has a white NHMW 16492 (Häupl & Tiedemann
spot that makes it appear more conspicu- 1978) from Manáus, Brazil (Frost 2011).
ous than the tubercle itself really is – not Distribution.—‘‘Amazon Basin of Ecua-
near as large as the tarsal tubercles of O. dor, Brazil, Bolivia, Peru, Colombia, as
buckleyi, which also has 1–8 enlarged well as the upper Orinoco Basin of
supraorbital tubercles. Venezuela, and the Guianas’’ (Frost 2011).
Fertilized eggs are usually laid in small Vouchers for Guyana.—AMNH A-
pockets of water in vegetation, where the 145156–145158 and USNM 566166 from
tadpoles develop, although few nutrients Dubulay Ranch; AMNH A-166256–
may be present. However, the parents 166263, A-166268 and USNM 566117–
often return to the site, undergo amplexus 566120 from Berbice River Camp; AMNH
again, and deposit fresh fertilized eggs, A-166269 and USNM 566121 from Kona-
which the tadpoles consume (Jungfer & waruk Camp; ROM 22737–22739 from
Weygoldt 1999). Baramita; ROM 42996 from Kaieteur;
USNM 118057 from Kartabo; USNM
291274–291275 from Paramakatoi; and
Osteocephalus phasmatus USNM 531455–531474 from Iwokrama
MacCulloch & Lathrop, 2005 Forest Reserve (various localities). In
Plate 11B addition, Smith & Noonan (2001:357)
identified the following specimens as this
Type material.—The holotype is ROM species: AMNH A-21418–21419 from Ka-
39481, an adult female from the northeast makusa; UTA A-51628–51629 from ca. 1.0
plateau of Mount Ayanganna, Guyana, km N Imbaimadai, Peters Mt.; UTA A-
(05824 0 N, 059857 0 W; 1490–1550 m). 51625–51627 from ca. 3.4 km N Imbaima-
Distribution.—Known only from Guya- dai, Peters Mt.; and UTA A-51624 from
na, in the Pakaraima highlands (MacCul- 1.6 km (by road) W Kartabu Point.
loch & Lathrop 2005). Coloration in life.—The following notes
Vouchers for Guyana.—ROM 39481– are based on AMNH and USNM speci-
39485 and ROM 43852 from Mount mens from Dubulay Ranch (CJC field
Ayanganna (MacCulloch & Lathrop notes, 1995). Tan above, with brown
2005). markings; spots on sides and anal spot
Coloration in life.—‘‘Dorsum medium black; throat and chest cream with gray or
brown with scattered black flecks. Tym- brown spots, but throat pale yellow on
panic region and upper lip dark brown. some; bones green; iris gold with black
Flanks and groin white with dark brown radiating lines. Kok & Kalamandeen
(2008:188) noted that the dorsum may be from Amazonian Colombia, Peru, Ecua-
dark brown and some individuals have a dor, or Bolivia’’ (Frost 2011).
yellow middorsal stripe or small cream Vouchers for Guyana.—AMNH A-
spots. 70980 from Kartabo; AMNH A-141106
from Ceiba Biological Station, Madewini
River, ca. 4.8 km (linear) east of Timehri
Phyllomedusa bicolor (Boddaert, 1772) Airport (06829 0 57 00 N, 058813 0 06 00 W);
Plate 11D AMNH A-141107–141112, A-141115–
141116, A-145152, and USNM 566124–
Type material.—The original name was 566125 from Dubulay Ranch; AMNH A-
Rana bicolor, the type locality was restrict- 166271–166276 and USNM 566126–
ed to Surinam, and the holotype is not 566129 from Berbice River Camp; and
known to exist (Frost 2011). AMNH A-166280–166281 from Konawa-
Distribution.—‘‘Amazon Basin in Brazil, ruk Camp.
Colombia, Bolivia, and Peru; the Guianan Coloration in life.—The following is
region of Venezuela and the Guianas’’ based on AMNH and USNM specimens
(Frost 2011). Kok & Kalamandeen (2008) from Ceiba Biological Station, Dubulay
included Amazonian Ecuador also. Ranch, and Konawaruk Camp (CJC field
Vouchers for Guyana.—AMNH A- notes, 1994–1998). Leaf green above, set
70978–70979 from Kartabo; ROM 22740 off on lower arms and legs by thin cream
from Baramita; ROM 28419 from Para- stripe; posterior half of sides, upper arms,
makatoi; and USNM 531597–531606 from and thighs (but for narrow green dorsal-
Iwokrama Forest Reserve (various locali- most area) and feet bright orange with
ties). dark blue or purple bars and reticulate
Coloration in life.—‘‘Dorsal colour markings; lips cream, the lower lips edged
green, dark green by night;...Ventral sur- posteriorly with row of black dots; chest
face...light grey, sometimes with a few very pale yellow or orange; abdomen and
white ocelli. White ocelli on lower lip, ventral surfaces of thighs pale yellow or
flank, and hidden surfaces of thigh. Iris darker pale orange; iris metallic tan with
silvery grey’’ (Kok & Kalamandeen black reticulation.
2008:192). Comments.—In the recent past, the
Comments.—This species is the largest specific epithet has been incorrectly spelled
of the leaf frogs, attaining a body length of without the middle h as hypocondrialis
115 mm (Kok & Kalamandeen 2008). (Lescure et al. 1995).
Phyllomedusa hypochondrialis Phyllomedusa tarsius (Cope, 1869)

(Daudin, 1800)
Plate 11E Type material.—The original name was
Pithecopus tarsius and the holotype is
Type material.—The original name was USNM 6652 (now lost, according to
Hyla hypochondrialis based on material Duellman 1974) from the Rio Napo,
from Surinam. See Frost (2011) for a Departamento Loreto, Peru.
discussion about possible type specimens. Distribution.—‘‘Amazon Basin in south-
Distribution.—‘‘Eastern Colombia east ern Colombia, Ecuador, Peru, and Brazil,
through northern and eastern Venezuela, . . .as well as. . .eastern Venezuela’’ (Frost
through the Guianas and throughout 2011); recently reported from Guyana
Brazilian Amazonia; currently not known (Forlani et al. 2012).
Vouchers for Guyana.—AMNH A- Forest Reserve (various localities); and

13552 from ‘‘British Guiana’’ (quoted USNM 531781–531783 from Baramita.
from the AMNH catalog; no additional Coloration in life.—The following is
locality data). The specimen was collected based on AMNH and USNM specimens
by W. Beebe, identified by D. Cannatella from Berbice River Camp (CJC field notes,
in 1978, and recently confirmed by CJC. 1997). Green dorsum, but greenish brown
Most recently Forlani et al. (2012) report- on snout, blending gradually (posteriad)
ed specimens (with color photographs) into the green of the midbody; yellow spots
from the vicinity of the hydropower dam ventrolaterally, becoming orangish red in
site on the Kuribrong River, Potaro- groin; orangish red on wrist, behind upper
Siparuni District, Guyana; vouchers are arms, on anterior and posterior surfaces of
UG (¼CSBD) HA 1000, 1001, 1100, and thighs, and anterior lower legs; chestnut
1101. brown posteriolaterally on lower arms and
Coloration in life.—The following is ventrally on lower legs and feet; chin,
from Duellman (1978:178), based on throat, chest, and abdomen pale chestnut
specimens from Ecuador. ‘‘The dorsal brown; small white spots often on throat
surfaces and sides of the head are green. and chest; on body, inconspicuous dorso-
The flanks are dull green or bluish green lateral line of tiny yellow dots; iris light
with discreet or confluent cream, pink, or copper. Kok & Kalamandeen (2008:194)
pale orange spots. The anterior and refer to the iris as ‘‘silvery grey’’ and the
posterior surfaces of the thighs are brown lower lip as white.
or dull green with cream, pale orange, or
bluish-gray spots. The throat, chest, and
ventral surfaces of the limbs are dark Pseudis paradoxa (Linnaeus, 1758)
brown or grayish-brown; the belly is gray
or pale orange-brown. Ventral markings Type material.—The original name was
consist of a pair of white spots on the Rana paradoxa and the syntypes from
throat, a median white spot on the chest, Surinam are NHRM 144–148 (Frost
and usually a pair of para-anal white 2011).
spots. The iris is bright orange with bold Distribution.—‘‘Lower Rı́o Magdalena
black reticulations.’’ Valley as well as Arauca and Meta in
Colombia; Guianas from northeastern
Venezuela east through Guyana, Surinam,
Phyllomedusa vaillantii Boulenger, 1882 and French Guiana to the mouth of the
Plate 11F Amazon, thence west in the Amazon
Valley to northern Bolivia and southeast-
Type material.—The holotype is BMNH ern Peru; Trinidad’’ (Frost 2011).
1947.2.22.34 (Condit 1964:98), a female Vouchers for Guyana.—AMNH A-
from Santarem, Brazil. 70918–70920 from Kartabo (identified by
Distribution.—‘‘Guianas and Amazon R. G. Zweifel); and USNM 162975–
Basin from Guyana, Surinam, French 162981 from Enmore Estate (6844 0 N,
Guiana, and northeastern Brazil to Co- 057859 0 W).
lombia, Ecuador, Peru, and northern Coloration in life.—The following is
Bolivia; reported from Cerro Neblina, from Lescure & Marty (2000:284) based
Amazonas, Venezuela’’ (Frost 2011). on frogs from French Guiana. Dorsal
Vouchers for Guyana.—AMNH A- surface green, especially in the anterior
166282, A-166286–166290, and USNM part, and brown, especially in the posterior
566130–566132 from Berbice River Camp; part, speckled with dark spots; flanks,
USNM 531607–531616 from Iwokrama limbs, and posterior surface of thighs
brown, spotted with white; ventral surface Scinax cruentommus (Duellman, 1972)
white, more or less flecked with brown
spots and with 3–4 brown lines more or Type material.—The original name was
less anastomosing or converging at the Hyla cruentomma and the holotype is KU
level of the thighs; brown reticulation on 126587, an adult male from Santa Cecilia
the underside of the limbs; iris golden on the Rı́o Aguarico, Provincia Napo,
yellow with a horizontal chestnut bar. Ecuador (0802 0 N, 76858 0 W; 340 m).
Comments.—‘‘These are moderately Distribution.—‘‘Amazon Basin in Ecua-
large aquatic frogs; at least one species, dor and northern Peru (drainages of the
Pseudis paradoxa, is well known for its Rı́o Napo, Rı́o Marañon, and Rı́o Ucaya-
gigantic tadpole’’ (Duellman 2005:261). li); reported from 70–90 km north of
Manaus, Brazil; possibly in Amazonian
southeastern Colombia’’ (Frost 2011).
Scinax boesemani (Goin, 1966b) 49447 from ‘‘British Guiana’’ (quoted from
Plate 12A AMNH records, with no additional local-
ity data available), collected by R. Snedi-
Type material.—The original name was gar on the Terry-Holden Expedition,
Hyla boesemani and the holotype is 1937–1938, identified by Julian Faivovich.
RMNH 12601, an adult male found near Coloration in life.—‘‘When collected [at
Zanderij, Suriname District, Suriname. night] pale yellow above with indistinct tan
Distribution.—‘‘Northern Guyana, Sur- dorsolateral stripes. Later [day] changed to
inam, and French Guiana, and Amazonas, tan with olive-brown markings. Vocal sac
Venezuela, to Amazonas and Pará, Brazil’’ bright yellow. Iris pale bronze with median
(Frost 2011). horizontal reddish copper streak. Palpe-
Vouchers for Guyana.—AMNH A- brum clear. Bones green...[the foregoing
145160, A-167101–167102, and USNM for some individuals; the following for
566133 from Dubulay Ranch; ROM others] Dorsum varies from yellow to
43219–43228, USNM 291215–291219, brown to tan to gray-green with or without
291221–291234, and 497717 from Kaieteur distinct markings. Markings are brown to
National Park; and USNM 531475– yellow-brown to greenish brown. Flanks
531492 from Iwokrama Forest Reserve yellow with or without brown spots. Iris
(several sites). silver with red horizontal streak. Posterior
Coloration in life.—The following is surfaces of thighs yellow-brown. Vocal sac
based on the AMNH specimens from deep yellow; venter pale yellow (males).
Dubulay Ranch (CJC field notes, 1994). Belly and throat of female white’’ (Duell-
Brown above on all surfaces; dark brown man 1972:268).
mask from snout, through eye, over ear,
then slanting downward and breaking up
into spots; throat yellow; chest and abdo- Scinax nebulosus (Spix, 1824)
men cream; translucent tan beneath arms;
translucent green beneath legs; iris copper Type material.—The original name was
speckled with black; bones green. Kok & Hyla nebulosa and the neotype is MNRJ
Kalamandeen (2008:198) add that the 4055 from Belem, Para, Brazil (Hoogmoed
dorsum may have cream or yellow spots, & Gruber 1983, Glaw & Franzen 2006).
flanks may have dark brown spots, and the Distribution.—‘‘Southeastern Venezue-
‘‘hidden surfaces of thighs [are] greyish la, through the Guianas and the lower
brown.’’ Amazon region to Alagoas in northeastern
Brazil; also Amazonian Bolivia’’ (Frost Reserve, Siparuni River, 4845 0 17 00 N,

2011). 050801 0 28 00 W.
Vouchers for Guyana.—AMNH A- Coloration in life.—The following is
46497 and A-93335–93336 from the ‘‘Up- from Lescure & Marty (2000:166), based
per Rupununi River’’; and AMNH A- on frogs from French Guiana. Dorsum
141126–141128, A-141131–141134 and brown, with a darker interorbital triangle
USNM 566134–566135 from Dubulay and bars; flanks and anterior surface of
Ranch, all identified by Julian Faivovich; thighs greenish, more or less marbled with
USNM 129534 from Kartabo; USNM black; posterior surface of thighs orange
162842–162845 from Atkinson (¼Timehri with broad black bars, at times extending
Airport); USNM 162913–162914 from onto the posterior surface of the shank;
Enmore Estate (6844 0 N, 057859 0 W); and ventral surface reddish brown, but the
USNM 497718 from Kumu, at base of throat is white in females, grayish in males,
Kanuku Mountains (3815 0 52 00 N, 059843 0 and the belly is white; iris gold to silver
34 00 W). with a horizontal brown band across the
Coloration in life.—The following is middle.
mens from Dubulay Ranch (CJC field
notes, 1994). Grayish tan to brown above
(after chloretone; had looked gray at Scinax ruber (Laurenti, 1768)
night); some individuals with dark brown
markings, including bands on thighs; Type material.—The original name was
throat cream or pale gray, with dark gray Hyla rubra and the neotype is RMNH
or black speckles; abdomen cream or light 15292B from Paramaribo, Surinam (Frost
gray; undersides of arms and legs translu- 2011).
cent gray. Lescure & Marty (2000:164) add Distribution.—‘‘Amazon Basin of Brazil,
that specimens may be brown or brown Peru, Colombia, and Ecuador; Guianas;
and green above, with an inconspicuous coastal northern South America; eastern
interorbital triangle, there are black bars Panama; Tobago, Trinidad, and St. Lucia
over dark blue on the hidden surfaces of (Lesser Antilles). Introduced on Puerto
the hind limbs, and there is a black spot in Rico’’ (Frost 2011).
the groin. Vouchers for Guyana.—AMNH A-
141121, A-141124, A-145161, and USNM
566136 from Dubulay Ranch; also
USNM 118055 from Kartabo; USNM
Scinax proboscideus (Brongersma, 1933)
162865–162871 from Atkinson (¼Timehri
Type material.—The original name was Airport); USNM 162915–162938 from
Hyla proboscidea and the holotype from Enmore Estate (6844 0 N, 057859 0 W);
Surinam is number Z. 354 of the Zoolog- USNM 164165–164177 from Mabaruma
ical Museum of Amsterdam (in the process Compound (8812 0 N, 059847 0 W); USNM
of being transferred to the RMNH). 284390–284391 from Takutu Mountains
Distribution.—‘‘Interior of the Guianas’’ (6815 0 N, 059805 0 W); USNM 497719 from
(Frost 2011) and Brazil (M. S. Hoogmoed, Karanambo Ranch; USNM 497720 from
pers. comm.). Georgetown, 300 New Garden Street
Vouchers for Guyana.—AMNH A- (6848 0 N, 058810 0 W); and USNM 535762–
49248–49249, 49251, and 49254 from 535772 from Baramita.
Shudikar-wau, identified by W. E. Duell- Coloration in life.—The following is
man and confirmed recently by CJC; also based on AMNH specimens from Dubulay
USNM 531493 from Iwokrama Forest Ranch (CJC field notes, 1994). Dorsum
tan, with dark brown stripes (two each Distribution.—‘‘Non-forest habitats of

side); arms and legs tan above with brown northern Colombia and Venezuela to
spots; yellow below (brightest on throat), Surinam; eastern, southern, and southeast-
but greenish yellow beneath hind legs. ern Brazil’’ (Frost 2011).
Comments.—Following analyses of Vouchers for Guyana.—AMNH A-
DNA sequences of frogs from the Guianan 139068–139076, A-139086, and USNM
Region, Fouquet et al. (2007b) suggested 566137–566143 from Aishalton; AMNH
that several cryptic species might be A-141120, A-141123, A-141125, A-145162,
masquerading under the name Scinax and A-167107 from Dubulay Ranch, most
ruber. of which were identified by Julian Faivo-
vich; and ROM 22745–22749 from Bar-
amita.
Scinax trilineatus based on AMNH specimens from Aishal-
(Hoogmoed & Gorzula, 1979) ton and Dubulay Ranch (CJC field notes,
Type material.—The original name was 1993, 1994). Brown or tannish gray above
Ololygon trilineata and the holotype, from (body, arms, lower legs) with dark gray,
12 km SE El Manteco, Bolivar, Venezuela, brown, or black markings (although uni-
is RMNH 18257 a female. form dark brown when photographed);
Distribution.—‘‘Scattered localities in thighs with very narrow stripe colored as
the lowlands of eastern Venezuela and upper body, but anterior and posterior to
this are black-edged yellow or gold spots
savannas of Guyana and Surinam, as well
or mottling (also below shanks and on
as adjacent Brazil’’ (Frost 2011).
anterior of ankle and upper foot); white or
cream on chin and chest, chin with gray
43673 from Isheartun; AMNH A-46248
dots; abdomen translucent gray; fleshy
from Rupununi River near Mora, east of
pink where translucent (especially on
Mt. Tirke; and AMNH A-97944–49 from
thigh); iris copper with black reticulation.
Parabam, identified by M. S. Hoogmoed. Lescure & Marty (2000:174) add that there
Coloration.—We found no description are black and orange patches on the flanks
of coloration in life for these very small and in the groin, and the ventral surface
frogs. ‘‘Colour in preservative light brown may be yellow in males from French
to greyish brown with a pattern of dark Guiana.
brown longitudinal stripes: a narrow [dark Comments.—Application of the name S.
brown] vertebral stripe, wider dorsolateral x-signatus to specimens from Guyana
bands. Flanks either uniform dark brown needs additional research (J. Faivovich,
or with a dark lateral band along the lower pers. comm.).
part. A dark brown canthal stripe. Ventral
parts immaculate, white to pale brown’’
(Hoogmoed & Gorzula 1979:193).
Scinax sp.
Plate 12B
Scinax cf. x-signatus (Spix, 1824) Voucher for Guyana.—AMNH A-

Plate 12C, D 166499 from Berbice River Camp, identi-
fied by Julian Faivovich.
Type material.—The original name was Coloration in life.—The following is
Hyla x-signata and the holotype is ZSM based on the only known specimen (above;
2494/0, now lost (Glaw & Franzen 2006), CJC field notes, 1997). All dorsal surfaces
from Provincia Bahia, Brazil. tan (darker and lighter); all ventral surfac-
es very pale translucent tan, but chin and Distribution.—Known only from Guya-
throat yellow; iris a reticulation of black na, endemic to Kaieteur National Park
and copper. (Kok & Kalamandeen 2008).
Comments.—This is probably a new Vouchers for Guyana.—The holotype;
species. Additional specimens should be BMNH 1938.10.3.26 from Kaieteur; KU
collected and studied, and the species 221735 collected with the holotype; and
should be named. ROM 42983–42984 and USNM 291235–
291237 from Kaieteur National Park.
Coloration in life.—‘‘Dorsum grayish
tan with minute dark brown flecks on
Sphaenorhynchus lacteus (Daudin, 1800) body and faintly darker transverse bars
on limbs—one on forearm, three on
Type material.—The original name was shank, and two on tarsus. Dark brown
Hyla lactea and the holotype, from an canthal stripe from nostril to eye; narrow
unspecified locality in Amérique, is in the dark brown line from posterior corner of
MNHN (Frost 2011). orbit to point above tympanum; narrow
Distribution.—‘‘Amazon and Orinoco creamy white labial stripe continuous
basins of Colombia, Venezuela, Ecuador, with creamy white venter on anteroven-
Peru, Bolivia, and Brazil; Guianas, and tral edge of insertion of arm. Tubercles in
Trinidad’’ (Frost 2011). cloacal region and tips of discs on digits
Vouchers for Guyana.—AMNH A- creamy white; iris pale silvery bronze with
1327–1328 and A-13048 from ‘‘British black flecks’’ (Duellman & Yoshpa
Guiana’’ (no additional data in AMNH 1996:278). Kok & Kalamandeen
catalog); ROM 11081 and 11090 from (2008:204) provide additional details on
the vicinity of Georgetown; USNM coloration.
127888 from Akyma, Demerara River
(5853 0 N, 059818 0 W); and USNM 162942
from Enmore Estate (6844 0 N, 0578 Trachycephalus coriaceus (Peters, 1867)
59 0 W). Plate 12E
based on Lescure & Marty (2000:178)
Hyla coriacea and the syntypes are ZMB
for frogs from French Guiana. Dorsal
5807 (3 specimens; Bauer et al. 1995) from
and lateral surfaces green to yellowish Surinam.
green, with a light dorsolateral line; thin Distribution.—‘‘French Guiana, Suri-
brown stripe from snout to eye; ventral nam, and Upper Amazon Basin in Co-
surfaces bluish gray, except belly white; lombia, Ecuador, Peru, Bolivia, and
males with yellow vocal sac; outer areas of downstream to near Manaus, Brazil’’
limbs and above vent with light edging; (Frost 2011).
iris orange. Vouchers for Guyana.—IRSNB 13672
from Kaieteur National Park (Kok et al.
2005); USNM 566122 from Berbice River
Tepuihyla talbergae Camp.
Duellman & Yoshpa, 1996 Coloration in life.—The following is
based on the USNM specimen from
Type material.—The holotype is CSBD Berbice River Camp (CJC field notes,
198920, an adult male from Kaieteur Falls, 1997). Dorsal surfaces tan, with greenish
366 m (05810 0 N, 59828 0 W), Mazaruni- brown markings (including large dorsal
Potaro District, Guyana. hourglass); sides yellow; black patch on
shoulder; chin, throat, and chest yellow; from Mission of Santo Antonio do Prata
abdomen of yellow warts surrounded by at the River Maracanã (interior of the
reticulation of bright orangish red; ventral State of Pará).
posterior thigh area likewise; anterior of Distribution.—‘‘Widespread from east-
arms, hands, narrow throat band, narrow ern Ecuador, Peru and northern Bolivia
chest band, anterior and posterior of through the Amazon Basin to the Guiana
thighs, ventral surface of lower legs, and Shield’’ (Kok & Kalamandeen 2008:210).
feet all bright orangish red; iris copper. Vouchers for Guyana.—IRSNB 14756
Kok & Kalamandeen (2008:208) add that from Kaieteur National Park.
the dorsal ground color is reddish brown Coloration in life.—‘‘Dorsal ground col-
by night and there is ‘‘a saddle-shaped our dark brown, with one large whitish,
dark brown blotch over the sacrum.’’ tan or greenish brown blotch narrowly
outlined with a creamy border on the flank
and another on the top of the head, the
Trachycephalus hadroceps latter often having the shape of a triangle;
(Duellman & Hoogmoed, 1992) tubercles [on back] usually with white tip;
ventral surface...greenish white to light
Type material.—The original name was brown; iris golden with four radiating
Hyla hadroceps and the holotype is KU black lines (a black ‘Maltese cross’); finger
69720, an adult male from area north of webbing...[and] toe webbing greenish blue’’
Acarai Mountains, west of New River (ca. (Kok & Kalamandeen 2008:210).
028N, 588W), Rupununi District, Guyana.
Distribution.—‘‘Extreme southern
Guyana; French Guiana; presumably in Trachycephalus typhonius (Linnaeus, 1758)
adjacent Surinam’’ (Frost 2011).
Vouchers for Guyana.—The holotype is Type material.—The original name was
the only specimen known from Guyana. Rana typhonia and the holotype is UUZM
Coloration in life.—The following is 134; the type locality was restricted to the
based on frogs from French Guiana vicinity of Paramaribo, Surinam (Lavilla
(Lescure & Marty 2000:192). Dorsal sur- et al. 2010).
face light chestnut with a dark brown Distribution.—‘‘Lowlands of Mexico,
preorbital area and a large, equally dark Central America (to 2500 m in Guatema-
brown dorsal area across middle of back la), to Amazon Basin of Colombia, Ecua-
and extending onto sides at axilla and dor, Peru, Venezuela, and Brazil, Guianas,
groin; limbs with dark brown transversal and south to Paraná (Brazil), Paraguay,
bars; toe discs light green; ventral surface northern Argentina; Trinidad and Toba-
greenish, speckled with brown, almost go’’ (Frost 2011).
marbled on throat; iris chestnut to beige, Vouchers for Guyana.—AMNH A-
with a brown horizontal bar and at times a 70977 from Kartabo; AMNH A-141142–
vertical one below the pupil, thus making 141143, A-141145, A-167105–167106, and
an incomplete cross. USNM 566123 from Dubulay Ranch;
USNM 162939–162941 from Enmore Es-
tate (6844 0 N, 057859 0 W); and USNM
Trachycephalus resinifictrix (Goeldi, 1907) 1 6 4 1 7 9 f r o m M a b a r u m a ( 8 81 2 0 N ,
Plate 12F 059847 0 W).
Type material.—The original name was based on the AMNH and USNM speci-
Hyla resinifictrix and the holotype is mens from Dubulay Ranch (CJC field
BMNH 1947.2.23.24 (Condit 1964:94) notes, 1994). Dark greenish brown above,
with tan or yellowish tan markings (body, Vouchers for Guyana.—AMNH A-

arms, legs); yellow below, brightest ven- 46264, 46268–46269, 46272–46273, 80028,
trolaterally (and on chin of some). Using and A-87877–87888 from Shudikar-Wau,
the name Phrynohyas venulosa, Lescure & identified by W. R. Heyer. Also AMNH
Marty (2000:198) stated that the toe discs A-166320 from Berbice River Camp, and
are emerald and iris is gold with black USNM 531500 (identified by W. R. Heyer)
coalescing vermiculations. from Iwokrama Mountain Gorge Trail,
Comments.—Until recently, this species Iwokrama Forest Reserve.
was referred to as Phrynohyas venulosa or Coloration in life.—According to Les-
Trachycephalus venulosus. cure & Marty (2000:224), based on spec-
imens from French Guiana, the dorsum is
grayish brown to beige, occasionally with
dark brown spots (usually without), and
Leptodactylidae rarely with a vertebral dark brown stripe
This essentially neotropical family of and/or dorsolateral orangish yellow stripe.
frogs was named more than 110 yr ago The ventral surfaces are white; iris chest-
(Werner 1896) and its validity has been nut.
confirmed by phylogenetic analyses based Comments.—This species is so similar
on DNA sequence data, although compo- to A. hylaedactyla (see characters in the
sition of the family has changed signifi- key) that some specimens are very diffi-
cult to identify. In southeastern Peru
cantly (Frost et al. 2006, Pyron & Wiens
these two taxa are part of a complex of
2011).
four cryptic species that were distin-
These frogs range in size from tiny to
guished by vocalizations and subtle char-
very large. Most species are terrestrial or
acters (Angulo et al. 2003). These are
semi-aquatic, and, depending on the spe-
small frogs, usually less than 30 mm in
cies, may be found in habitats ranging adult body length.
from savanna to primary forest. Breeding
often involves depositing eggs in foam
nests formed during amplexus, which may
be in a burrow near the edge of a pond or Adenomera hylaedactyla Cope, 1868
depression such that subsequent rainfall Type material.—The original name was
may flood the nest and the tadpoles can Cystignathus Hylaedactylus and the holo-
swim out. Pseudopaludicola boliviana is an type is ANSP 2240 (Malnate 1971) from
exception in this regard. the Napo or upper Marañon River, Peru.
Distribution.—‘‘Amazonian forests of
southeastern Colombia and Venezuela east
through the Guianas and south into
Adenomera andreae Müller, 1923 northern, eastern, and central Brazil, and
south through Amazonian Ecuador and
Type material.—Originally there were Peru to Bolivia, 0–1000 m elevation’’
five specimens from the type locality of (Frost 2011).
‘‘Peixeboi (a. d. Bragançabahn) Staat Parà, Vouchers for Guyana.—AMNH A-
Brasilien’’ (Müller 1923:41). Heyer (1973) 39671 and A-39696 from Kartabo; and
designated ZSM 145/1911 as lectotype. AMNH A-46565–46568 from Marudi, all
Distribution.—‘‘Lowlands of northern identified by W. R. Heyer. Also AMNH
South America east of the Andes including A-13494 from Kartabo; and A-141118 and
and limited to Amazonia to the south and A-145151 from Dubulay Ranch. Also
east’’ (Frost 2011). ROM 22760 from Baramita; UMMZ
52506 from Dunoon; UMMZ 85148– patterns among our sample: (Pattern A)
85149 from Membaru Creek, upper Ma- dorsal ground color dark brown, grey or
zaruni River; and UMMZ 83581–83582 black with no or a barely discernible
from Yarikita River, mouth and Haul pattern; (Pattern B) dorsal ground color
Over; USNM 163070 from Kamuni Creek; light to medium grey with well-defined
USNM 291122 from Chiung River at pattern, characterized by a strongly
Kato; USNM 497721, 497723–497724, marked dark brown to black interorbital
497726–497729 from Karanambo; and bar and postorbital ridges, chevron be-
USNM 566155 from Berbice River Camp, tween shoulders almost continuous with
all identified by W. R. Heyer. round lumbar spots and irregular dark
Coloration in life.—The following is brown to black markings on back; (Pattern
based on frogs from French Guiana C) similar to the preceding pattern, but
(Lescure & Marty 2000:226). Dorsum with dorsal ground color light grey, brown
grayish brown to beige with a pattern of or reddish brown, with lighter, smaller,
dark brown to black spots that may be and much less evident markings on back
discrete and oval or joined. Some individ- (some of the markings may be absent);
uals with dorsolateral or lateral light (Pattern D) rather similar to pattern B but
stripes. Dorsal surfaces of limbs usually with dark brown to black mottling on
lighter with dark brown bands. Ventral dorsum instead of well-defined markings;
surfaces white. Iris chestnut. (Pattern E) dorsal ground color dark grey
Comments.—See the comments for
to black with large orange to orangish
Adenomera andreae.
brown oblique lateral stripe; darker mark-
ings on back may be discernible or not’’
(Kok et al. 2007:49–50). In addition, these
Adenomera lutzi Heyer, 1975 authors noted that many individuals have
Plate 14B a narrow light yellow middorsal stripe;
ventral surfaces are yellow to orangish
Type material.—The holotype is BMNH yellow, usually with dark gray stippling on
1905.11.1.17, an adult male from China- throat and chest; dorsal surfaces of arms
poon R., upper Potaro (probably Chena- and legs usually with dark bands; dark
powu River), Guyana. brown or black seat patch.
Distribution.—All specimens known are
‘‘from the Pakaraima Mountains of Guya-
na’’ (Kok & Kalamandeen 2008:220).
Vouchers for Guyana.—The holotype; Engystomops cf. pustulosus (Cope, 1864)
the paratypes from Demerara Falls, Guya-
na (BMNH 1872.10.16.43–1872.10.16.44); Type material.—The original name was
AMNH A-3789 from Kai Camp, Kaieteur Paludicola pustulosus and the holotype is
Region, identified by W. R. Heyer; ROM USNM 4339 (apparently lost) from New
40155, 40165, 40167, and USNM 546152 Grenada, on the River Truando, Colom-
from Mount Ayanganna; ROM 43437– bia (Frost 2011).
43455 from Mount Wokomung; ROM Distribution.—‘‘Eastern and southern
44123–44125 from Partang; and ROM Mexico (Veracruz and Oaxaca) south and
44126–44127 from the Merume Moun- east through Central America to Colombia
tains. and Venezuela’’ (Frost 2011).
Coloration in life.—Adenomera lutzi Voucher for Guyana.—BMNH 1977.885
‘‘exhibits a remarkable color-pattern poly- from Bartica District (no additional local-
morphism. We recognize five major color ity data).
Coloration in life.—The following is Leptodactylus fuscus (Schneider, 1799)

from Savage (2002:224) based on frogs Plate 13E
from Costa Rica. ‘‘Ground color brown-
ish to gray above, nearly uniform or with Type material.—The original name was
dark brown blotches, a narrow middorsal Rana fusca based on syntypes thought to
cream to orange stripe or paired dorso- be from Surinam (Frost 2011).
lateral light stripes or a combination of Distribution.—‘‘Savannas from Panama
these; warts on anterior part of body throughout South America, east of the
Andes, south to southern Brazil, Bolivia,
usually reddish, orange or yellow; often
Paraguay, and Argentina’’ (Frost 2011).
an orange spot between shoulders; male
vocal sac black with a medium [median?]
70909–70916 from Kartabo (identified by
light line when inflated; chest dark; upper
R. G. Zweifel); AMNH A-136031 and
surface of leg with several transverse dark
USNM 497732–497733 from Karanambo;
bars; venter dirty white, usually with large
AMNH A-139087–139099, A-139107, and
black spots posteriorly; iris tan or pale
USNM 566173–566179 from Aishalton;
brown.’’
AMNH A-141091–141095, A-141098, A-
Comments.—Colin McCarthy sent
145140–145146, A-167112–167113 and
CJC photographs of the BMNH vouch- USNM 497735–497739 and 566180–
er from Guyana. These photographs 566184 from Dubulay Ranch; USNM
were compared with specimens of sever- 162880–162893 from Atkinson (¼Timehri
al related frogs from Venezuela (Gorzu- Airport); USNM 162943–162965 from
la & Se ñaris 1999; also compared with Enmore Estate (6844 0 N, 057859 0 W);
AMNH A-70631, A-70635, and A- USNM 497734 from Takutu River at
70639, E. pustulosus from Venezuela), Lethem.
and characters of the pustulosus species Coloration in life.—The following is
group as discussed by Lynch (1970) and based on the AMNH and USNM speci-
Cannatella & Duellman (1984). The mens from Karanambo and Aishalton
BMNH voucher clearly does not have (CJC field notes, 1992, 1993). Dorsum
the diagnostic characters of Physalae- reddish brown with dark brown spots and
mus cuvieri, which also occurs in Guya- with or without a light gray vertebral
na, and does have the following stripe; lateral body greenish brown with
characters that fit E. pustulosus: dark dark brown spots; paravertebral ridges
markings above ankles and wrists; dark reddish brown; dorsolateral and lateral
rounded markings on abdomen; pres- ridges yellowish tan; lateralmost warts
ence of parotoid glands (possible, but cream; arms and legs yellowish tan with
photo not absolute); presence of ellipti- gray or black markings; posterior thigh
cal flank glands (possible, but photo not warts pale yellow; chin and throat cream
absolute). The main character that is with gray specks; black line posteriorly on
problematical is that the BMNH vouch- lower lip; chest pale yellow; abdomen
er does not appear to be sufficiently brighter yellow; underneath arms and legs
warty, although this could be affected by translucent pale yellow.
age and method of storage in preserva- Comments.—This species is abundant in
tive. Owing to the uncertainties, we refer savannas and disturbed habitats. Analyses
to this taxon as cf. pustulosus and of DNA sequence data suggest that there
encourage future field work and collect- are distinct evolutionary lineages of these
ing to confirm or reject the hypothesis frogs from localities north versus south of
that this species occurs in the Bartica the Amazon, although no significant dif-
District. ferences in morphology or vocalizations
have been found to distinguish these beneath eye and ear to corner of mouth,
populations (Wynn & Heyer 2001, Camar- perhaps beyond; lower lips brown with
go et al. 2006). cream spots; on some individuals, a rust,
pale orange, or yellow wash also on
uppermost arm, on light warts laterally
Leptodactylus guianensis on body, and as a streak on anterior of
Heyer & de Sá, 2011 thigh; some warts cream in anal region;
Plate 13B chin, throat, chest, and abdomen cream,
yellowish cream, tan, or gray, with or
Type material.—USNM 531509 from without tan or gray mottling; translucent
Pakatau Camp (4845 0 17 00 N, 59801 0 28 00 W; under legs and with or without yellow
85 m elevation), Sipuruni River, Iwokrama wash in groin.
Forest Reserve, Guyana. Comments.—This species is morpholog-
Distribution.—Lowlands of the Guiana ically similar to the other two species in the
Shield, in Brazil, Guyana, Suriname, and bolivianus complex, from which it differs
Venezuela. significantly in molecular characters
Vouchers for Guyana.—Heyer & de Sá (Heyer & de Sá 2011). Adult males of L.
(2011) listed numerous specimens from guianensis differ from males of the others
Guyana in addition to the holotype, para- by having a single modestly chisel-shaped
topotypes, and paratypes. The following spine on each thumb. The other two
are specimens that we examined: AMNH species of the bolivianus complex do not
A-3990 from Kalacoon; AMNH A-10397, occur in Guyana.
A-10423, and A-11709 from Kartabo;
AMNH A-136032–136034 from Karanam-
bo; AMNH A-145130–145131, A-145133– Leptodactylus knudseni Heyer, 1972
145134, and USNM 566161 from Dubulay Plate 13F
Ranch; AMNH A-166341–166349, A-
166356–166358, and USNM 566162– Type material.—The holotype is LACM
566167 from Berbice River Camp; AMNH 72117, a juvenile female from Limoncocha
A-166359–166366 and USNM 566168– (0824 0 S, 76837 0 W; 260 m), Provincia de
566172 from Konawaruk Camp; ROM Napo, Ecuador.
20580–20590 from Kurupukari; USNM Distribution.—‘‘Greater Amazon basin
146369–146374 and 497731 from Moco of Bolivia and Brazil north to Colombia,
Moco, at the base of the Kanuku Moun- Venezuela, Trinidad, and the Guianas’’
tains; USNM 497730 from Karanambo (Frost 2011).
Ranch; and USNM 531501–531511 from Vouchers for Guyana.—AMNH A-
Iwokrama Forest Reserve (various locali- 70880–70881 from Kartabo (identified by
ties). W. R. Heyer); AMNH A-141100–141104,
Coloration in life.—The following is and A-145136 from Dubulay Ranch;
based on the AMNH and USNM speci- AMNH A-166372–166373 and USNM
mens from Karanambo and Dubulay 566185–566186 from Berbice River Camp;
Ranch (CJC field notes, 1992–1995). Light and AMNH A-166376 and USNM
brown with darker brown markings on 566187–566188 from Konawaruk Camp.
upper surfaces of head, body, and arms; Also ROM 22762–22765 and USNM
legs similar but also may have light olive 535773 from Baramita; and ROM 28451–
green mottling and spots on rear of thigh 28452 from Paramakatoi, identified by W.
and a pale yellow or olive green wash on R. Heyer. Also USNM 531512–531520
thigh; dorsolateral folds with reddish from Iwokrama Forest Reserve (several
brown stripe; light cream, tan, or rust line localities).
Coloration in life.—The following is chest, and in some individuals the anterior

based on the AMNH specimens from part of the belly. The iris is bronze with
Dubulay Ranch (CJC field notes, 1994). fine black reticulations.’’
Brown above with light tan markings;
upper arms black on rear, brown above
with wash of rust; rust laterally on body; Leptodactylus longirostris Boulenger, 1882
upper legs (thighs) brown above with light
tan to black bands, black on rear with light Type material.—BMNH 76.5.26.4 was
orange-rust, reticulation, or bars; dorsal designated as the lectotype reportedly
surface of lower legs as dorsal body; gray from Santarem, Brazil, but see Frost
throat, chest, anterior abdomen; yellow (2011) for discussion.
area about waist; legs cream below, except Distribution.—‘‘Guianas, eastern Vene-
thighs pink. zuela, eastern Amazonian Colombia
Comments.—These large frogs can grow (Guainı́a), and Brazilian Amazonia’’
to an adult body length of 170 mm (Kok & (Frost 2011).
Kalamandeen 2008:214). Vouchers for Guyana.—AMNH A-
139109–139112 and USNM 566189 from
Aishalton; ROM 20591–20594, USNM
Leptodactylus leptodactyloides 291238–291243 and 497740–497741 from
(Andersson, 1945) Kaieteur; USNM 118065–118066 from
Kartabo; USNM 291124–291126 from
Type material.—The original name was Chiung River at Kato; and USNM
Eleutherodactylus leptodactyloides and the 291177 from Paramakatoi.
holotype, unnumbered at NHRM, was Coloration in life.—The following notes
from the Rio Pastaza, eastern Ecuador are from Kok & Kalamandeen (2008:218),
(Andersson 1945). based on specimens from Kaieteur Na-
Distribution.—‘‘Greater Amazon Basin tional Park. ‘‘Dorsal ground colour and
and the Guianas, 15–400 m elevation’’ pattern very variable, light brown to
(Frost 2011). reddish brown, sometimes greyish or dark
Vouchers for Guyana.—AMNH A- brown, uniform or with irregular dark
39625 from Kartabo, identified by W. R.
brown markings, middorsal and paraver-
Heyer.
tebral stripes absent or present;...throat
Coloration in life.—The following (and
and chest cream or white, turning yellow
quotes) are from Duellman (2005:284),
on belly...upper lip with a white, cream,
based on frogs from Peru. Dorsum gray,
light brown or pinkish stripe below eye,
tan, or reddish brown, darkest individuals
always entering eye anteriorly. Black stripe
with snout lighter than body; dorsolateral
from tip of snout to eye.’’
fold dark brown; narrow dark brown or
Comments.—‘‘Found in open areas like
black interorbital bar; most specimens
with some irregular black spots on sides; savannah’’ (Kok & Kalamandeen 2008:
indistinct light bars on lips, the posterior- 218).
most of which extends from the posterior
corner of the eye; posterior of thigh
mottled (dark brown to black with cream Leptodactylus macrosternum
to pale tan), many individuals also having Miranda-Ribeiro, 1926
a distinct light stripe (mostly distally on
thigh); ‘‘throat...brown or gray with cream Type material.—The original name was
flecks; ...rest of...venter...white with brown Leptodactylus ocellatus macrosternus and
or dark gray reticulations on...hind limbs, the holotype is MZUSP 448 (Frost 2011),
probably from Salvador, Bahia, Brazil Ranch; AMNH A-166384–166393 and

(Frost 2011). USNM 566193–566198 from Berbice Riv-
Distribution.—‘‘Amazonian Colombia, er Camp; AMNH A-166401–166403 and
most of Venezuela, and the Guianas south USNM 566199 from Konawaruk Camp;
through Brazil to Bolivia; Trinidad’’ (Frost ROM 22767 from Baramita; USNM
2011). 118058 from Kartabo; and USNM
Vouchers for Guyana.—AMNH A- 531521–531541 from Iwokrama Forest
136035–136038, USNM 497742–497763, Reserve (several localities).
and 566190–566192 from Karanambo; Coloration in life.—The following is
AMNH A-139114–139115 from Aishal- based on the AMNH and USNM
ton; and USNM 146368 from Lethem. specimens from Dubulay Ranch, Ber-
Coloration in life.—The following is bice River Camp, and Konawaruk
based on the AMNH specimens from Camp (CJC field notes, 1994–1998).
Karanambo (CJC field notes, 1992). Tan Dorsum reddish brown (possibly bright-
or greenish brown with darker brown er on sides, but sides may be gray) to
spots on head, body, and arms; legs brown, with darker brown markings
similar but wash of green superimposed surrounded by a dirty cream line; arms
(in groin also, and on some, posteriorlat- and legs tan with brown bands, legs at
erally on body); chin and throat cream, times also with yellow wash; upper lip
mottled with gray; chest and abdomen light tan, pinkish tan, or copper, above
cream to pale yellow, sometimes with which is a dark brown or black mask
gray mottling, the latter and thighs with from the snout through the eye and ear;
wash of pale greenish yellow; underside a black stripe may be present on the
of lower legs translucent gray. Lescure & posterior surfaces of the arms; rear of
Marty (2000:242) stated that the rear of thigh may have a yellow stripe (possibly
the thigh is spotted with black and broken) with a thin very dark brown or
yellow. black line above it; vent black with
Comments.—Specimens of this species vertical yellow bar; ventral surfaces
from the Guianas were formerly referred yellow (possibly brighter on abdomen
to as Leptodactylus ocellatus. It occurs in and legs), with small gray or tan spots
savannas. or flecks on the throat; possibly a few
brown to black spots on underside of
lower legs.
Leptodactylus mystaceus (Spix, 1824)

Plate 14C
Leptodactylus pentadactylus
(Laurenti, 1768)
Plate 14D
Rana mystaceus and the lectotype, ap-
parently lost, is ZSM 2504/0 (Hoogmoed Type material.—The original name was
& Gruber 1983), with the type locality Rana pentadactyla and a neotype (RMNH
restricted to Solim ões, Brazil (Frost 29559) from Surinam, Marowijne, Lelyge-
2011). bergte, Suralcokamp V was designated by
Distribution.—‘‘Amazon basin from its Heyer (2005).
southern limit in Brazil to Bolivia, Peru, Distribution.—‘‘Amazon Basin from
Ecuador, Colombia, Venezuela, and the southern Colombia south to Peru, north-
Guianas’’ (Frost 2011). ern Bolivia, and northern Brazil to French
Vouchers for Guyana.—AMNH A- Guiana; Honduras to western Colombia
141105 and A-145147 from Dubulay and northwestern Ecuador’’ (Frost 2011).
Vouchers for Guyana.—AMNH A- 057859 0 W); and USNM 531542–531557

70118–20 from Shudikar-wau, a headwater from Iwokrama Forest Reserve (several
stream of the Essequibo River (1815 0 N, localities).
58835 0 W); these frogs were previously Coloration in life.—The following is
identified by W. R. Heyer (1978) and based on AMNH specimens from Dubulay
recently confirmed by CJC. Ranch and Berbice River Camp (CJC field
Coloration in life.—The following is notes, 1995–1997). Dorsum dark tan with
based on Lescure & Marty (2000:248) for brown markings on arms and legs, black
frogs from French Guiana. Dorsum red- spots on sides; light rust line below
dish brown with 5 or 6 transversal, darker tympanum; throat, chest, and belly white
brown bands with light rims; sides lighter or cream with gray spots, mottling, or
than the back, with some black tubercles; reticulation; some with yellow in groin and
posterior surface of thighs black with on posterior surface of thigh; underside of
white dots (but thighs banded above this); legs with black reticulation. Kok & Kala-
venter cream, marbled with black, espe- mandeen (2008:224) stated that the ground
cially toward the rear; iris reddish copper, color of the dorsum is more variable, from
except the upper third golden. greenish or grayish brown to reddish
Comments.—These frogs, which usually brown.
occur in forest, become quite large, up to Comments.—Previously this species was
185 mm in body length (Lescure & Marty confused with Leptodactylus podicipinus.
2000). They are similar to Leptodactylus
knudseni.
Leptodactylus rhodomystax
Boulenger, 1884
Leptodactylus petersii (Steindachner, 1864) Plate 14F
Plate 14E
Type material.—BMNH 1947.12.17.81
Type material.—The original name was from Yurimaguas, Huallaga River, North-
Platymantis Petersii and the holotype from ern Peru was ‘‘designated lectotype by
Marabitanas, Amazonas, Brazil has been Heyer, 1979, Smithson. Contrib. Zool.,
lost (Frost 2011). 301: 30’’ (Frost 2011).
Distribution.—‘‘Known from the Gui- Distribution.—‘‘Guianas through north-
anas, the Amazon basin (including one ern and central Brazil to Amazonian
locality in Ecuador), and isolated localities Ecuador, Peru, and Bolivia’’ (Frost 2011).
from the Cerrado open formations in Vouchers for Guyana.—AMNH A-
central Brazil reported from Pando and 13489 and A-70903 from Kartabo;
Santa Cruz, Bolivia’’ (Frost 2011). AMNH A-49246 and A-51942 from Shu-
Vouchers for Guyana.—AMNH A- dikarwau (identified by W. R. Heyer);
39630–39631 and A-39633 from Kartabo AMNH A-166412–166414, A-166418–
(identified by W. R. Heyer); AMNH A- 166419, and USNM 566205–566207 from
136050 and USNM 497725, 497775, and Berbice River Camp; AMNH A-166420
566149–566154 from Karanambo; AMNH from Konawaruk Camp; ROM 40153–
A-145150 from Dubulay Ranch; AMNH 40154 from the base of Mount Ayangan-
A-166308–166318 and A-166405–166406 na; and USNM 531558–531571 from
from Berbice River Camp; AMNH A- Iwokrama Forest Reserve (several locali-
166409–166411 and USNM 566202– ties).
566204 from Konawaruk Camp; ROM Coloration in life.—The following is
20613–20616 from Kurupukari; USNM based on AMNH and USNM specimens
162950 from Enmore Estate (6844 0 N, from Berbice River Camp (CJC field notes,
1997). Dorsum reddish brown or tan, ish tan blotches and black markings;...
brightest on sides, with few (anterior) dark ventral surface...variable in pattern, white
brown markings; tiny white specks on to light greyish brown with diffuse brown
sides; snout and side of face dark brown; mottling (...in juveniles and subadults...
light stripe on lip tan, becoming white ventral surface of thighs orangish); throat
posteriorly; upper groin with black patch white, heavily marked with grey mottling
having white spots; upper surfaces of arms in juvenile and female, grey in adult
and legs reddish brown, arms with row of male...dirty white, cream or brownish lines
white specks posteriorly, from wrist to from eye to lip (ill-defined in some
elbow; rear of thighs black with white, specimens)...dirty white, cream or brown-
yellow, and/or gray spots; chin and throat ish interorbital stripe,’’ based on specimens
very dark gray or grayish tan, with white from Kaieteur National Park (Kok &
spots; chest gray or grayish tan, with white Kalamandeen 2008:228).
spots or reticulation; some with gray
reticulation on abdomen; lower groin and
posterior abdomen yellow or orangish Leptodactylus validus Garman, 1888
rust; translucent light tan or white beneath
arms, light yellow beneath legs; black Type material.—See Frost (2011) for a
beneath feet; iris deep reddish brown on discussion of the many syntypes from
lower half, mixed copper and black above. Kingston, St. Vincent, and citation of
Heyer’s designation of MCZ 71920 as
lectotype.
Leptodactylus rugosus Noble, 1923 Distribution.—‘‘Lowlands of Venezuela
Plate 15A and the Guianas, and north-central Ro-
raima, Brazil, and the continental islands
Type material.—The holotype is of Tobago and Trinidad, from elevations
AMNH A-1169, an adult male from near of 0–360 m; Lesser Antillean islands of St.
Kaieteur Falls, Guyana. Vincent, Bequia, and Grenada’’ (Frost
Distribution.—‘‘Southern Venezuela east 2011).
to western Guyana; likely in intervening Vouchers for Guyana.—AMNH A-
adjacent Brazil’’ (Frost 2011). 53435–53437 from Isheartun (identified
Vouchers for Guyana.—Specimens in by W. R. Heyer); AMNH A-136043–
addition to the holotype include paratypes 136046, A-136049, USNM 497722,
AMNH A-1166–1168 from Tukeit and 497764–497774, and 566200–566201 from
Kaieteur, with identifications confirmed Karanambo; AMNH A-145148–145149
by W. R. Heyer. Also ROM 20597– from Dubulay Ranch; ROM 20619–
20606, 43233–43252, USNM 291245– 20632 from Kurupukari; ROM 40183–
291266, and USNM 342151–342160 from 40187 from Baramita; USNM 118059–
Kaieteur National Park; ROM 40150– 118062 from Kartabo; USNM 146366–
40151 from the base of Mount Ayangan- 146367 from Moco Moco (3820 0 N,
na; USNM 118063–118064 and 129537– 059839 0 W); USNM 162872–162879 from
129538 from Kartabo; and USNM 118684 Atkinson (¼Timehri Airport); and USNM
from Membaro Creek, Upper Mazaruni 162966–162967 from Enmore Estate
River. (6844 0 N, 057859 0 W).
Coloration in life.—‘‘Dorsal ground col- Coloration in life.—The following is
our variable, ranging from greyish or based on AMNH and USNM specimens
greyish brown to olive brown or reddish from Karanambo (CJC field notes, 1992).
brown (dorsum bright red in some sub- Brown or greenish brown above, with
adults), with irregular pale cream to grey- inconspicuous black or blackish gray
bands on arms and legs; tan mottling on with light tan bands and spots; orangish
posterior of thighs; tips of fingers white; red spots behind and below knee; all
venter cream (but pale yellow on abdo- ventral surfaces dark gray to black with
men), mottled with dark gray, sometimes white spots.
extensively on chin and throat; beneath Comments.—‘‘Found exclusively in pri-
arms and legs translucent pinkish gray; mary forest, often associated with large
anal warts tan. The face may be gray nests of leaf cutting ants (Atta spp.)’’ (Kok
anterior to the eyes and a short gray line & Kalamandeen 2008:216).
may go posteromedially behind each eye;
also there may be a black interorbital line.
Comments.—This species was referred Physalaemus cuvieri Fitzinger, 1826
to as Leptodactylus pallidirostris, but re- Plate 13A
cent research, including DNA sequence
comparisons, shows that L. pallidirostris is Type material.—‘‘Type(s): [from Brazil]
a junior synonym of L. validus (Yanek et presumably originally in NHMW, but not
al. 2006). noted in recent type lists’’ (Frost 2011).
Distribution.—‘‘Northeastern, central,
and southern Brazil; Misiones, Argentina;
Lithodytes lineatus (Schneider, 1799) eastern Paraguay; Departments of Beni
Plate 14A and Santa Cruz in Bolivia; possibly low-
lands of southern Venezuela (see com-
Type material.—The original name was ment)’’ (Frost 2011).
Rana Lineata based on material in the Vouchers for Guyana.—AMNH A-
‘‘Musei Lampiani’’ with no locality infor- 52875 from Isheartun and A-52876 from
mation given. Kuyuwini Landing, identified by A. J.
Distribution.—‘‘Eastern Peru, Ecuador, Cardoso as ‘‘cf. cuvieri’’ (quoted from the
Bolivia, and Brazil through southern AMNH catalog). Also USNM 291178–
Venezuela to the Guianas; possibly Trini- 291183 from Paramakatoi; USNM
dad’’ (Frost 2011). 535775–535780 from Baramita; USNM
Vouchers for Guyana.—AMNH A- 531577–531583 from Iwokrama Forest
10408, A-11664–11666, A-13504, A- Reserve, ca. 3 mi S of Kurupukari base
13564, A-23122, A-39589–39590, A- camp on Georgetown-Lethem Road;
70917, and A-70992 from Kartabo; USNM 566212 from Dubulay Ranch;
AMNH A-141147 from Dubulay Ranch; and UTA 55526 from Moses.
AMNH A-166428, AMNH A-166433– Coloration in life.—The following is
166435, and USNM 566208–566211 from based on USNM 566212 from Dubulay
Berbice River Camp; AMNH A-166436 Ranch (CJC field notes, 1994). Brown
from Konawaruk Camp; and ROM above, with tan snout (anterior to eyes),
22786–22787 from Baramita. tan spot on upper back, and, posteriorly,
Coloration in life.—The following is tan vertebral line starting at about mid-
based on the AMNH specimen from body; dark brown stripe slants ventrally
Dubulay Ranch (CJC field notes, 1994). from behind eye past midbody, on side;
Dorsum black, with pale tan to gold arms and legs tan with brown bands, lower
dorsolateral light stripe; small white spots arms with dark brown stripe on outer
on side of face and body; arms banded edge; pinkish orange (bright) in groin,
black and light tan; bright orangish red behind knee, down rear of lower leg, and
spot in groin and two on upper posterior anteriorly on ankle; throat, anterior chest,
thigh; rest of upper and anterior thigh and beneath lower legs gray; abdomen and
black with light tan bars; lower legs black beneath and behind thigh very pale yellow.
Comments.—The vouchers for Guyana Distribution.—‘‘Disjunct regions of east-

compare favorably with characters given ern Colombia, northern Brazil (Roraima),
by Gorzula & Señaris (1999:67–79) and and Amazonian Venezuela through south-
also with AMNH A-23586 and A-54511 ern Guyana to southern Surinam; eastern
(P. cuvieri from Brazil). Bolivia, Paraguay, southwestern Brazil,
and northern Argentina’’ (Frost 2011).
Pleurodema brachyops (Cope, 1869) 136029–136030 and USNM 497776–
Plate 13C, D 497777 from Karanambo; AMNH A-
139182–139191, A-139197–139198, and
Type material.—The original name was USNM 566219–566223 from Aishalton;
Lystris brachyops and syntypes (ANSP and USNM 497778 from Moco Moco at
2260–2264) were from the Magdalena base of Kanuku Mountains (3818 0 12 00 N,
River, New Grenada (Malnate 1971). 059839 0 00 00 W).
Distribution.—‘‘Guyana and northern Coloration in life.—The following is
Brazil (Roraima) through Venezuela (in- based on the AMNH and USNM speci-
cluding Isla Margarita) and Colombia into mens from Karanambo and Aishalton
Panama in dry llanos and savanna areas; (CJC field notes, 1992, 1993). Dorsum
also Curaçao, Aruba, Bonaire, and Klein light tan to dark grayish tan or tannish
Bonaire in the Netherlands Antilles’’ gray, with or without darker tan or brown
(Frost 2011). markings; some individuals with cream,
Vouchers for Guyana.—AMNH A- light tan, or rust vertebral stripe; side of
139116–139127, A-139134–139135, and face with very pale streaks; arms and legs
USNM 566213–566218 from Aishalton. pale tan with gray bands; conspicuous
Coloration in life.—The following is broad lateral tan stripe, complete from
based on the AMNH and USNM speci- snout to vent; chin, throat, chest, and belly
mens from Aishalton (CJC field notes, cream with tiny dark gray specks, although
1993). Dorsum yellowish tan or gray, with on some individuals chin and throat are
gray markings and some individuals hav- gray or throat copper; translucent gray
ing small orange dots; most individuals beneath arms and legs, the latter some-
have a largely broken (clearest posteriorly) times with tiny black specks.
cream to orangish tan, thin vertebral Comments.—These are very small frogs,
stripe; posterior lateral abdomen with adults being less than 20 mm in body length.
swollen orbs (‘‘eyespots’’), black with light
gray markings; upper arms and legs as
body, but on legs most marks are black;
Microhylidae
throat (all are males) gray or tan, with
darker gray or tan speckling; abdomen Both the intrafamilial and interfamilial
cream, on some individuals translucent relationships of this family have been
posteriorly (as legs), fleshy pink; bright discussed recently by Frost et al. (2006)
orangish red flash marks in groin, behind and van der Meijden et al. (2007), based on
thigh, behind knee, beneath lower leg, and phylogenetic analyses of DNA sequence
on anterior of ankle. data. Some of these frogs, especially of the
genera Synapturanus and Otophryne, can
be very difficult to find, as they live on or
Pseudopaludicola boliviana Parker, 1927a within the forest floor and are cryptically
colored. Some, such as species of Synap-
Type material.—The holotype is BMNH turanus would appear to be rare, based on
1927.8.1.1, a female from Santa Cruz, Bolivia. the few specimens in collections; yet they
may be abundant in nature but extremely Chiasmocleis jimi

hard to find as males vocalize from Caramaschi & Cruz, 2001
beneath the ground during rains and Plate 15B
quickly escape in burrows as one tries to
dig them up. We still have much to learn Type material.—The holotype is MNRJ
about the life history, ecology, and popu- 15459, an adult female from Brazil, Am-
lation genetics of these frogs. azonas, Municipality of Humaitá, Igarapé
do Banheiro (07831 0 S, 63800 0 W).
vicinity of the type locality, in the states
Chiasmocleis hudsoni Parker, 1940 of Amazonas and Para, Brazil (Frost 2011)
Type material.—The holotype is BMNH and Guyana.
1939.1.1.3, a male, from the New River,
166437–39 and USNM 566224 from Ber-
Guyana (750 ft elevation). This locality is
bice River Camp.
in an area of dispute between Guyana and
Coloration in life.—‘‘In life, color of
Surinam, but it is likely that any species
dorsum of body, arms and legs uniform
occurring here occurs also at other local-
reddish brown with minute irregular white
ities in both countries.
dots; no light mid-dorsal stripe; ventrolat-
Distribution.—‘‘Cerro Neblina in
eral region and venter cream, heavily
Guianan Venezuela, as well as in Guyana,
grayish spotted; gular region grayish, with
Surinam, and French Guiana; presumably
irregular white dots; no femoral light
to be found in adjacent Brazil’’ (Frost stripe’’ (Caramaschi & Cruz 2001:3).
2011). Individuals of the AMNH and USNM
Vouchers for Guyana.—The holotype; series from Berbice River Camp were
paratypes, 3 males and 2 females with the noted in life (CJC field notes, 1997) as
same data as the holotype, BMNH follows: ‘‘all dorsal surfaces chestnut
1939.1.1.4–1939.1.1.8. brown; venter different shades of gray,
Coloration in life.—The following is throat being tannish gray, but hind legs
based on Lescure & Marty (2000:266) for with mottling of orange on the underside.’’
frogs from French Guiana: dorsal surface USNM 566224 ‘‘differs in having a thin
chestnut with light markings at times white stripe on the posterior femoral
forming a dorsolateral line from the snout surfaces, continuous across the anus
to the flanks; ventral surface light, spotted [¼vent]; yellow ground [color] on throat
with brown; sole of foot black. and beneath arms; and white with black
We examined a preserved specimen mottling beneath hind legs, abdomen, and
(MNHN 2001.0322) from French Guiana. chest.’’
In alcohol, the dorsum is dark grayish tan, Comments.—All individuals are smaller
with numerous white dots. There is no than 20 mm in body length.
light vertebral stripe nor is there one on the
posterior surface of the thighs. The tan
throat has irregularly shaped small light
Chiasmocleis shudikarensis Dunn, 1949
spots. The pale belly has a tan reticulation.
Palmar and plantar surfaces of hands and Type material.—AMNH A-43674 from
feet, respectively, are very dark. Shudikar-wau, upper Essequibo River,
Comments.—These tiny frogs (maxi- Guyana.
mum body length about 20 mm) live on Distribution.—Guyana, Surinam,
the ground in primary forest (Lescure & French Guiana, and Brazil (Amazonas)
Marty 2000:266). (Frost 2011).
Vouchers for Guyana.—The holotype the thighs. The palmar and plantar surfac-
(see above) is the only specimen known es of the hands and feet, respectively, are
from Guyana. very dark.
based on Lescure & Marty (2000:268) for
frogs from French Guiana: dorsal surfaces Ctenophryne geayi Mocquard, 1904
light brown to grayish brown, with or Plate 15C
without a fine yellow vertebral line that
meets a similar line on the posterior Type material.—The holotype from the
surfaces of the thighs; irregular black Sarare River, Colombia is registered as
patch in groin; brown throat; venter and MNHN 1903.84 (Frost 2011).
undersides of legs white with large irregu- Distribution.—‘‘Northern South Ameri-
lar black patches; iris reddish chestnut. ca from Surinam, Guyana, and Brazil to
Dunn (1949:8) stated the following in Amazonian Venezuela, Colombia, Ecua-
the original description, based on the dor, and Peru’’ (Frost 2011).
holotype, probably in alcohol: ‘‘above gray Vouchers for Guyana.—AMNH A-
with fine light flecking and a narrow, light, 71399 from Kartabo (identified by R. G.
middorsal line; throat gray with a narrow Zweifel); AMNH A-145159 from Dubulay
light median line; belly and undersurfaces Ranch and AMNH A-166441–166446 and
of limbs cream with black spotting or USNM 566225–566227 from Berbice Riv-
mottling, increasing in amount posteriorly; er Camp. Also ROM 22769 from Barami-
a black inguinal spot, rather vague and
ta.
irregular.’’
Coloration in life.—Specimens from the
We examined a series of French
AMNH series from Berbice River Camp
Guianan specimens in alcohol (MNHN
were noted in life (CJC field notes, 1997) as
2001.0831–2001.0836), from which we
follows: grayish brown body with numer-
took the following notes. The largest
ous tiny white dots; thin light tan vertebral
(2001.0836; SVL ¼ 25 mm) is a female
and it is the only one to have a very thin line; dark chestnut brown on lateral
light vertebral stripe, as occurs on the surfaces beneath a white lateral stripe on
holotype. It also has a light tan throat, head, arms, body, and legs; hands and feet
whereas some of the smaller ones (e.g., with some yellowish tan; all ventral sur-
2001.0832) have a dark gray to black faces black with irregular white spots.
throat and may be males. The dorsum is
dark grayish tan to dark tannish gray,
covered with tiny light dots. The posterior Elachistocleis surinamensis
of the thigh is dark with a thin light stripe (Daudin, 1802a)
along its length from the hip, which may Plate 15D
continue down the leg to the foot. The
throat and chest have irregular light Type material.—The original name was
stippling. A light V (apex to the rear) is Bufo surinamensis based on an unspecified
on the chest of all specimens, but only on individual from Surinam.
the large female is it bisected by a light Distribution.—Venezuela and Guyana
midventral line that extends from the to French Guiana; Trinidad (Frost 2011;
anterior edge of the lower jaw posteriorly M. S. Hoogmoed, pers. comm.).
to the rear of the abdomen. The basically Vouchers for Guyana.—AMNH A-
white abdomen has very dark brown to 139136–139147, A-139154–139155, and
black spots and reticulations, which also USNM 566228–566233 from Aishalton;
are on the ventral and anterior surfaces of AMNH A-141135–141137, A-141139–
141140, A-167110, and USNM 566234 The AMNH specimens from Aishalton
from Dubulay Ranch. and Dubulay Ranch are in general similar to
Coloration in life.—The following is what has been described for E. surinamensis
based on field notes of CJC (1993, 1994) and E. surumu. However, they exhibit
for the AMNH and USNM specimens from considerably more individual variation in
Aishalton and Dubulay Ranch: dark gray ventral coloration than usually is attributed
above with very light gray speckling; mottled to local populations. After several years in
gold and gray below; orange spots in groin, preservative those with the lightest bellies
behind knee (some becoming a stripe behind have a rather abrupt transition on the sides
femur), and on upper side of foot, although between the darker dorsal coloration and
this last spot may be gold in color. lighter ventral coloration. These frogs also
Comments.—The following four names lack a vertebral stripe and have light spots in
have been applied to Elachistocleis-like the groin and behind the knee plus light
frogs of the Guianan Region and Trinidad striping behind the thigh. The largest
for many years: E. bicolor, E. ovalis, E. individual from Aishalton has a snout-vent
surinamensis, and Relictivomer or Elachis- length of 35 mm, that from Dubulay Ranch,
tocleis pearsei. Caramaschi (2010) added a 39 mm, which is larger than what Carama-
new name to the list, Elachistocleis surumu, schi (2010) reported for specimens from
which he applied to frogs from the state of Roraima.
Roraima, Brazil; in the same paper he We conclude that the name E. surina-
named what he called four additional new mensis is most applicable to the specimens
we have seen from Guyana. Furthermore,
species from Brazil. The number of real
the taxonomic problems involving these
species enveloped by all these names and
frogs will be resolved only by doing
how to apply the names remains unknown
extensive analyses of local variation, geo-
to us today.
graphic variation, and DNA sequence
For the Guianan Region, overall varia-
data. In addition, specimens need to be
tion includes whether individuals have an
analyzed from throughout the ranges of
immaculate venter or one with pronounced the so-called species without limitations of
light spotting and dark reticulations; wheth- political boundaries.
er there are light spots in the groin and
behind the knee; whether there is light
striping behind the thigh; and whether there
is a fine light (or dark) vertebral stripe. Hamptophryne boliviana (Parker, 1927b)
However, what is missing from most studies Type material.—The original name was
is careful analysis of local individual varia- Chiasmocleis boliviana and the holotype is
tion, geographic variation, and use of BMNH 1927.8.1.1, a female from Buena
comparative samples from throughout the Vista, Santa Cruz, Bolivia.
geographic ranges of the proposed species. Distribution.—‘‘Northern and western
None of the studies to date has involved sides of the Amazon basin: State of Bolivar
modern molecular analyses. (Venezuela), French Guiana, Surinam,
All of the specimens we have seen from Guyana, Brazil, Ecuador, Peru, Bolivia,
Guyana have the characters usually and Colombia’’ (Frost 2011).
thought to represent E. surinamensis, Vouchers for Guyana.—BMNH 1934.1.
which is consistent with the recent report 4.1 from Upper Cuyuni River (no addi-
of Lavilla et al. (2003). Contrary to tional locality data).
Caramaschi’s (2010) statements, Lavilla Coloration in life.—The following notes
et al. (2003) did not state that E. surina- are from Lescure & Marty (2000:274),
mensis has a light vertebral stripe. based on specimens from French Guiana:
Dorsum brown or light brown with a large from below the vent toward the rear of the
darker brown middorsal area having an knee]. The throat is dark brown, grading
irregular edge (two wavy notches) that to a paler brown on the chest and to
enlarges in width posteriorly; small black orange at about the mid-belly. The abdo-
spot above groin; a blackish brown band men and lower surface of the hind limb
from snout to cloacal area and on poste- and foot are also orange’’ (Campbell &
rior surface of thigh neatly separates Clarke 1998:311).
dorsal and ventral colorations; ventral
surface white (but throat brownish), tinged
with blue on the belly, and very spotted Otophryne robusta Boulenger, 1900a
with brown; sole of the foot and tarsus Plate 15E, F
blackish brown.
1899.3.25.18 from the foot of Mt. Rorai-
Otophryne pyburni ma, 3500 ft, reported originally as being
Campbell & Clarke, 1998 from Guyana. However, the type locality
is probably in Venezuela (Phelps 1938).
Type material.—The holotype is UTA See below (Vouchers) for specimens from
A-3814 from rainforest at 213 m elevation Guyana.
at Wacará, Vaupés Colombia. This site is Distribution.—The region of the Paka-
east of Mitú and close to the Brazilian raima Mountains of western Guyana and
border at approximately 01809 0 N, southeastern Venezuela, at elevations of
69855 0 W. approximately 400-1200 m; probably also
Distribution.—‘‘From Vaupés and Am- in adjacent Brazil (Campbell & Clarke
azonas, Colombia, across the lowlands of 1998).
southern Venezuela and through the Vouchers for Guyana.—Possibly the
Guianan region to Amapás, Brazil’’ holotype; ROM 39678–39681 from Mount
(Campbell & Clark 1998:311). Ayanganna; ROM 42962–42963 from
Vouchers for Guyana.—AMNH A- Mount Wokomung; ROM 44053–44054
71015 from Kartabo. from Meamu; UMMZ 85140 from Mak-
Coloration in life.—‘‘Dorsum...varying reba Falls; and USNM 549320 from the
from reddish brown, brown, gray, grayish north slope of Mount Roraima. For more
yellow...[or] yellow-ochre. The sides of the details, see MacCulloch et al. (2008b).
head and body and the concealed lower Coloration in life.—The following notes
parts of the limbs are dark brown and are drawn from MacCulloch et al.
sharply contrast with the pale dorsum. The (2008b:251). Dorsum variable, from dark
exposed upper parts of the limbs are, for orange with dark brown middorsal stripe
the most part, similar in color to the and lateral bars to almost uniform dark
dorsum. A grayish, whitish, or pinkish, brown with faint orange mottling; narrow
somewhat irregular line extends along the pale yellow dorsolateral stripe from snout
lateral ridge. The upper third of the iris is tip passing above eye to groin, continuing
pale bronze and the lower portion is dark onto anterior thigh in some individuals;
brown similar to the color on the side of lateral surfaces dark orange with small
the head. Most individuals have cream- black spots, some with white centers;
colored dots (tubercles) scattered over the dorsal surfaces of arms dark brown with
dorsum of the body and limbs. The vent is small white spots; dorsal surfaces of
surrounded with a dark brown spot. The thighs, shanks and feet dark orange with
large tubercles on the posterior surface of dark brown mottling and occasional dark
the thigh are white or pinkish [in a row brown bars; posterior thigh orange with
variable dark brown mottling and a few Synapturanus mirandaribeiroi

white spots, often connected by a white Nelson & Lescure, 1975
line; throat dark brown, chest orange with
dense brown speckles in males; throat and Type material.—The holotype is
venter orange with brown speckles in MZUSP 49981, an adult female from
females; digits pink; upper third of iris Kanashen (a Waiwai Indian village and
pale yellow, matching color of dorsolateral mission) on the Upper Essequibo River,
stripe; remainder of iris dark orange with Rupununi District, Guyana.
black reticulations. Distribution.—‘‘Northern Brazil, Co-
lombia, and the Guianas in humid tropical
forests at low elevations’’ (Frost 2011).
Otophryne steyermarki Rivero, 1968c
and paratopotypes include the following:
Plate 16A, B
AMNH A-90935–90943, MNHNP 1974–
Type material.—The original name was 379, and UMMZ 136147.
Otophryne robusta steyermarki and the Coloration in life.—‘‘The dorsal colora-
holotype is FMNH 74031, a male from tion in life ranged from light yellowish
the forest along Rı́o Tiricaclusia, 5900 ft, brown to light reddish brown and to dark
falls below summit camp, Chimantatepui brown’’ (Nelson & Lescure 1975:396).
massif, Venezuela.
Distribution.—Western Guyana and
southeastern Venezuela, at elevations of Synapturanus salseri Pyburn, 1975
1234–2140 m (MacCulloch et al. 2008b). Plate 16C
Vouchers for Guyana.—ROM 39677
from Mount Ayanganna; ROM 44052 Type material.—The holotype is UTA
from Partang; and ROM 42953–42961, A-4011, an adult male from Timb ó,
44778–44779, and 45271 from Mount Vaupés, Colombia.
Wokomung. Distribution.—‘‘Known from the type
Coloration in life.—‘‘Dorsum of body locality..., Cerro Neblina and 34 km south
and limbs black or dark grey with irregu- of Puerto Ayacucho, Amazonas, Venezue-
lar, variable yellow spots. A yellow line la; also reported from Kaieteur National
from tip of snout across eyelid to groin, Park, Guyana, and near Manaus, Ama-
continuing onto anterior thigh and some- zonas, Brazil’’ (Frost 2011).
times onto shank. Lateral surfaces usually Vouchers for Guyana.—AMNH A-
black or dark grey, occasionally dark 166450 and USNM 566235 from Kona-
orange, with yellow or orange spots, waruk Camp; IRSNB 14905–14911 (P. J.
usually smaller than dorsal spots. Venter R. Kok, pers. comm.) and ROM 11731
variable, ranging from black with small from Kaieteur National Park; and USNM
orange spots to orange with black or grey 118067 from Kartabo.
spots or mottling. Rear of thigh with a thin Coloration in life.—The following is
horizontal yellow line, continuing onto from field notes of CJC (1998) based on
shanks and feet. Forelimbs with a thin the AMNH specimen from Konawaruk
pale yellow line on the anterior and Camp: dark tan above with tiny light tan
posterior surfaces, from insertion of arm dots; venter light tan, but abdomen and
to toes. Palms and soles dark yellow or posterior of chest translucent gray. ‘‘In life
orange. Upper third of iris gold with black the dorsal aspect is medium brown with
reticulations, remainder of iris dark red light cream to orange spots over the back,
with black reticulations’’ (MacCulloch et sides, arms and legs. There is a light cream
al. 2008b:255). line along the canthus rostralis and upper
eyelid that becomes a series of irregular Coloration in life.—Dorsal color medi-

spots from the eye to the shoulder. The um brown, with irregular dark brown
venter is pearl white, has no pattern and spots; a dark bar from eye to shoulder;
the gula is light brown’’ (Pyburn ventral surface light brown, shaded with
1975:441). Individuals from Kaieteur Na- orange or pink, with irregular dark brown
tional Park are the same, but the dorsum spots.
on some may be grayish brown (Kok & Comments.—Pipa arrabali and Pipa
Kalamandeen 2008:232). aspera (a rare frog known only from
Comments.—Kok & Kalamandeen Suriname and French Guiana; Lescure &
(2008:232) provided taxonomic comments Marty 2000) were thought to be possibly
and also stated the following: ‘‘Males call conspecific, but Trueb & Massemin (2001)
exclusively during rain, from small bur- demonstrated that they are distinct species,
rows in the ground, below the leaf litter. despite their similarities.
Eggs are laid in burrows below the soil
surface; tadpoles do not feed and com-
plete their development within the bur- Pipa pipa (Linnaeus, 1758)
row.’’ Plate 16E
Pipidae Rana Pipa based on syntypes from Sur-
inam (see http://linnaeus.nrm.se/zool/
This family of frogs has been taxonom-
herp/madamph.html.en).
ically stable for a long time, and is based
Distribution.—‘‘Amazonian Bolivia to
on many morphological and DNA se-
Colombia and the Guianas to Peru,
quence characters (Frost et al. 2006). The
Ecuador; Trinidad’’ (Frost 2011).
species in Guyana are highly aquatic frogs
with rather flat bodies, and occur in ponds,
25191 from Kartabo, A-53492 from Mar-
ox-bow lakes, or slow-moving streams in
udi, A-53493–53494 from Onora, A-
low-elevation rain forest. Females carry
53495–53499 from Rupununi head;
their eggs embedded in their backs, from
USNM 531587–531596 from Iwokrama
which froglets emerge.
Forest Reserve; and USNM 566236 from
Berbice River Camp.
Pipa arrabali Izecksohn, 1976 Coloration in life.—Dorsal surface dark
Plate 16D brown to medium grayish brown, some-
times mottled; flanks and ventral surface
Type material.—The holotype, from light brown, with yellow or whitish mot-
Vila Amazônia, Municı́pio de Parintins, tling; at times, a large T-shaped ventral
Amazonas, Brasil, was a male in the marking composed of a midventral black
author’s private collection (EI 5311) and line from the cloaca forward to the middle
was destroyed (Frost 2011). of a ventral transverse black line between
Distribution.—‘‘Guyana, western Suri- the head and chest (Trueb & Cannatella
nam, eastern Venezuela, and northern, 1986:439, Lescure & Marty 2000:282).
central, and western Brazil’’ (Frost 2011).
52829–52831 from Kartabo; ROM 22809 Ranidae
from Baramita, ROM 22379–22398 and
USNM 291184 from Paramakatoi; ROM Although this family was named nearly
43991 from Mount Wokomung; and 200 yr ago (Rafinesque 1814), our under-
USNM 284392 from Takutu Mountains. standing of relationships (both intrafamil-
ial and interfamilial) has greatly improved on largest; other ventral surfaces light gray
only recently (Hillis & Wilcox 2005, Frost with irregular darker gray or black spots.
et al. 2006). This has involved numerous Comments.—Most individuals in the
name-changes, particularly among species AMNH and USNM series from Konawa-
previously included in the genus Rana. ruk Camp and the USNM specimens from
Kato and Kumu were found in or near
water flowing in streams. The ROM
Lithobates palmipes (Spix, 1824) specimens from Paramakatoi are from a
Plate 16F man-made pond.

Rana Palmipes based on material from the Species Accounts: Amphibia:
Amazon River, Brazil. None of the types Gymnophiona (Caecillians)
exist (Glaw & Franzen 2006).
Distribution.—‘‘Northern and Amazo- Frost et al. (2006:165) obtained very
nian South America east of the Andes, strong support for this group, based on
including Colombia, Ecuador, Peru, Boli- phylogenetic analyses of DNA sequence
via, Venezuela, the Guianas, and Brazil data, and characterized the group as
south to Alagoas, Bahia, and Mato follows: ‘‘Caecilians are a bizarre group
Grosso’’ (Frost 2011). of legless amphibians, primitively ovipa-
Vouchers for Guyana.—AMNH A- rous with aquatic larvae (Rhinatremati-
11655, A-11669–11671, and A-71006– dae, Ichthyophiidae), although some
71008 from Kartabo; AMNH A-139156 species are ovoviparous (with or without
from Aishalton; AMNH A-166453– direct development) and burrowing, as
166459, A-166466–166469, and USNM reflected by considerable numbers of oste-
566244–566249 from Konawaruk Camp; ological modifications.’’ Superficially, in-
ROM 28836–28970 from Paramakatoi; dividuals resemble earthworms (in some
ROM 42641 from the Ayanganna airstrip cases very large ones) more than they do
(05818 0 10 00 N, 059849 0 55 00 W); USNM other vertebrates, and it is unusual to find
291127–291128 and 291130 from Kato; one crawling on the surface of the ground,
USNM 497790 from Kumu at the base of except during or after a rain.
the Kanuku Mountains (3815 0 52 00 N, The most recent taxonomic revision,
059843 0 34 00 W); and USNM 531617– consistent with the phylogeny reconstruc-
531619 from Iwokrama Forest Reserve, tion, concluded that globally there are nine
Sipuruni River (4845 0 17 00 N, 059801 0 28 00 W). families that have existed for about 100
Coloration in life.—The following notes million yr (Wilkinson et al. 2011). Repre-
were taken on specimens of the AMNH sentatives of four families are known to
and USNM series from Konawaruk Camp occur in Guyana, including seven genera
(CJC field notes, 1998): top of head green, and 11 species.
fading on body (from anterior to shoulders
posteriorly to midbody) to become brown
posteriorly on the dorsum with irregular Caeciliidae
black spots; arms brown; legs same as
posterior dorsum and with dark brown Frost et al. (2006) and Wilkinson et al.
subtle bands on lower legs; posterior of (2011) found strong support for this family
thighs black with contrasting reticulation based on DNA sequence data and mor-
of pale blue, cream, or tan; throat cream phology. Most species are terrestrial,
on smallest, extensively smudged with gray burrowing forms.
Caecilia gracilis Shaw, 1802 each fold. A median ventral gray-slate

stripe, lighter than dorsum, scarcely dis-
Type material.—Shaw (1802:598) stated: tinguishable under neck. No light spots at
‘‘In the Museum Adolphi Friderici it is vent, tentacle, eyes, lips, or nostrils’’
figured under the name of C. tentaculata, (Taylor 1968:433).
and appears to be confounded with that
species.’’ It was said to be from South
America.
Distribution.—‘‘Surinam, and French Caecilia tentaculata Linnaeus, 1758
Guiana; Pará and Maranhão, Brazil; Plate 18B
northeastern Peru, presumably into adja-
cent southeastern Colombia, western Bra- Type material.—See Frost (2011) for a
zil, and eastern Guyana’’ (Frost 2011). discussion of the syntypes for which the
Vouchers for Guyana.—BMNH RR1987. only locality datum given was America.
946 from Demerara and 1887.1.22.30 from Distribution.—‘‘The greater part of wet-
Maccasseema. Also FMNH 35116 ‘‘from the forested South America east of the Andes,
Arenoque River,’’ Guyana, 2842 0 N Lat. south to central Peru and Amazonian
(Taylor 1969:282); UMMZ 47410–47411 Brazil’’ (Frost 2011).
and 52507 from Dunoon; UMMZ 76676 Vouchers for Guyana.—AMNH A-
from Wismar; and USNM 58750 from 49962 from Kamakusa; and USNM
‘‘Guyane’’ (no additional data). 566003 from Dubulay Ranch.
Coloration in life.—The following is Coloration in life.—The following is
from Lescure & Marty (2000:290), based based on the USNM specimen from
on specimens from French Guiana. Bluish Dubulay Ranch (CJC field notes, 1995).
lavender or lilac, a little lighter at the level The animal was dark gun-metal blue,
of the head and on the belly. Nussbaum & lighter below than above.
Hoogmoed (1979:223) stated that a spec-
imen from Surinam ‘‘RMNH 18696 was
said to be ‘dark purplish blue’ in life.’’
Comment.—Diameter of the body is Oscaecilia zweifeli Taylor, 1968
extremely slender relative to the body
length in this species.
AMNH A-20079 from a small creek
tributary to Rio Mazaruni, Guyana.
Distribution.—Known from Guyana
Caecilia pressula Taylor, 1968 and the vicinity of Cayenne, French
Type material.—The holotype is Guiana (Lescure & Marty 2000).
AMNH A-49475, a male from the Marudi Vouchers for Guyana.—The holotype;
Mountains, Guyana. and AMNH No. 20080 from the type
Distribution.—Known only from Guya- locality.
na, at the type locality (Frost 2011). Coloration.—The following is from
Vouchers for Guyana.—The holotype Lescure & Marty (2000:296), without
and AMNH A-49470–49474, 49476, all reference to source or whether it is
from the Marudi Mountains. coloration in life or in preservative.
Coloration.—In preservative, ‘‘dorsally Dorsum grayish lavender; belly whitish
dark grayish-slate, the head olive; from with some gray specks; a yellowish spot
high on the sides down to venter, yellow- near the nostril; a yellowish band on the
ish-olive or olive-yellow, the dark lines side of the head, ending at the aperture for
following the grooves interrupting this on the tentacle.
Rhinatrematidae Vouchers for Guyana.—The holotype;

BMNH 1976.102 from Winiperu, (6810 0 N,
Frost et al. (2006) and Wilkinson et al. 58834 0 W); BMNH 2008.601, BMNH
(2011) found strong support for this family 2008.602, IRSNB 1992–1996, and IRSNB
based on DNA sequence data and mor- 1998–1999 from at or in the vicinity of the
phology. These are oviparous animals with type locality; and USNM 566004 from
aquatic larvae. Adults have conspicuous Konawaruk Camp.
annuli and numerous secondary and ter- Coloration in life.—The following is
tiary grooves. based on the USNM specimen from
1998). Brown above with tiny yellow
speckles; yellow patch on head; yellow
Epicrionops niger (Dunn, 1942)
lateral stripe, with irregular edges; venter
Plate 18C, D
brown, gradually darkening to black pos-
Type material.—The original name was teriorly.
Rhinatrema nigrum and the holotype was Comments.—Kok & Kalamandeen
lost. The neotype is MBUCV 5360 from El (2008:246) described specimens from Kaie-
Dorado, Bolivar, Venezuela (Taylor 1968). teur National Park.
Distribution.—‘‘Central Guyana to
northeastern Venezuela; presumably in
adjacent Brazil’’ (Frost 2011). Siphonopidae
Vouchers for Guyana.—ROM 39682–
Wilkinson et al. (2011) recognized this
39683 ‘‘from the northeast plateau of
neotropical family of oviparous species, of
Mount Ayanganna, 1490 m’’ (MacCulloch
which the Guyanan forms are terrestrial
& Lathrop 2009:16).
burrowers.
Coloration in life.—‘‘The body is bluish-
black...with diffuse medium-gray mottling
on the sides of the head; a line of small
blue spots laterally; chin with three longi- Caecilita iwokramae
tudinal white bars, one at midline and one Wake & Donnelly, 2010
along the underside of each mandible; the Type material.—The holotype is CSBD
anus [¼ vent] white’’ (MacCulloch & HA 1500, an adult female from near Top
Lathrop 2009:17). Camp, Iwokrama Forest, Guyana (4820 0 N,
58848 0 W; ca. 1000 m).
Distribution.—Known only from Guya-
Rhinatrema shiv Gower, Wilkinson, na.
Sherratt, & Kok, 2010 Voucher for Guyana.—The holotype is
Plate 18E the only specimen known.
Coloration.—Coloration in preservative
Type material.—The holotype is IRSNB only was given by Wake & Donnelly
1991, a transformed male (Gower et al. (2010:917). ‘‘Skin colour nearly uniform
2010:49) from Sir Donald’s trail ca. 2 km light yellow-brown dorsally and ventrally.
from a tributary of the Elinkwa River, Melanophore pattern mottled, surround-
ESE Kaieteur National Park, Guyana, ca. ing light skin glands. Melanophores on
540 m elevation (5808 0 N, 59825 0 W) (Gower dorsum of head more concentrated on
et al. 2010:49). snout, fewer over parietal region of head.
Distribution.—Known only from Guya- Throat cream-coloured. Annuli slightly
na. darker posteriorly adjacent to annular
groove. Annuli darker and much narrower small pale spots. Cloacal disc, including
posteriorly.’’ small ventral area of terminal cap, pale.’’
Comments.—This is one of the most Comments.—This is another of the most
poorly known species in the world, as it is poorly known species in the world. The
known from only one extremely distinctive holotype and only known specimen was
specimen. It was found in ‘‘mixed low- identified by Taylor (1968) as a represen-
stature scrub forest on high hills’’ (Wake & tative of Microcaecilia unicolor. However,
Donnelly 2010:916) in a terrestrial, rather ‘‘Taylor’s description of this species was
than aquatic, situation. The specimen is as based on a specimen from Guyana that
small as the smallest species of caecilians differs so substantially from the type series
known, and it is lungless, as is only one [of unicolor]... that it most probably
other species of caecilian in the world. We represents an as yet undescribed species’’
know essentially nothing about its biology. (Wilkinson et al. 2009:417).
Microcaecilia iyob Wilkinson & Kok, 2010 Microcaecilia rabei (Roze & Solano, 1963)
Type material.—The holotype is FMNH Type material.—The original name was
35117, a mature male from the Oko River Gymnopis rabei and the holotype is
at Cuyuni River, Guyana. MBUCV 5126 from the base of Cerro
Distribution.—Known only from Guya- Lema, Rı́o Chicanán, Bolı́var, Venezuela.
na. Distribution.—‘‘State of Bolı́var, eastern
Voucher for Guyana.—The holotype is Venezuela, and Surinam; presumably in
the only specimen known. intervening Guyana and adjacent Brazil’’
Coloration in preservative.—‘‘The speci- (Frost 2011).
men [FMNH 35117] is now nearly a Vouchers for Guyana.—BMNH
uniform brown, the head somewhat cloud- 1968.1283 from Winiperu, identified by
ed with yellowish-brown back to the M. Wilkinson.
collars, which are somewhat darker brown Coloration in life.—The following is
than the following annuli; chin and throat from Nussbaum & Hoogmoed
are very light brown to second collar; vent (1979:225), based on specimens from
area whitish’’ (Taylor 1968:546). More Surinam. ‘‘In life...snout pinkish, anterior
than 40 yr later the same specimen was part of the body purple-blue, darkening
described (Wilkinson & Kok 2010:36–37) towards the posterior part of the body.’’
as follows: ‘‘Gray-brown, more brown ‘‘In preservative...blue-gray to slate above
anteriorly and where superficial epidermis and below, somewhat lighter anteriorly.
missing, darker, more plumbous gray The tip and sides of the snout are cream.
posteriorly, perhaps slightly paler on The chin and throat region is tan. The area
lateral flanks especially posteriorly where immediately surrounding the vent is white.
each annulus has a paler anterior and Apparently the Surinam and Venezuelan
darker posterior half. Head paler than specimens are nearly identical in colour
body especially ventrally, except for nar- pattern’’ (cf. Taylor 1968:541).
row darker lines, approximately level with Comments.—Most of the RMNH spec-
medial aspect of mandibles, central pale imens from Surinam ‘‘were collected in the
colour extending onto C1 [first collar]. daytime by digging in the upper soil layer
Collar grooves and AGs [annular grooves] in rainforest, between the buttresses of
edged in white, especially laterally. TAs t r e e s ’’ ( N u s s b a u m & H o o g m o e d
[tentacular apertures] and nares lie within 1979:225).
Microcaecilia cf. rabei Camp, it is by specimens from Konawaruk

Plate 18F Camp. The range in number of secondary
annular grooves for Guyanan specimens
Vouchers for Guyana.—AMNH A- superficially appears not to overlap those
166045–166053 from Berbice River Camp for specimens from Venezuela and Sur-
and AMNH A-166054–166058 from Ko- iname, but the differences are small and
nawaruk Camp. These specimens superfi- need confirmation. Considering that Mi-
cially are similar to ones reported from crocaecilia are small burrowing animals,
Kaieteur National Park (Kok & Kala- they may have low vagility, and they may
mandeen 2008:244–245). be prone to local differentiation of cryptic
Coloration in life.—The AMNH speci- species. Consequently, considerable molec-
mens from Berbice River Camp were ular work is necessary to determine their
noted in life (CJC field notes, 1997) as evolutionary history and to estimate the
being reddish brown, but with the distal number of species and how they differ
20–25% being grayish brown. The AMNH from each other. One specimen from our
specimens from Konawaruk Camp (CJC Berbice River Camp was found burrowing
field notes, 1998) were noted in life (CJC in a sandy dirt pile.
field notes, 1998) as being similar, but with
the additional note that the head was pink
(lighter below), which, as recalled, seemed
Typhlonectidae
similar to those from Berbice River Camp
as well; also, one was noted as having the This is another strictly neotropical
distal 40% grayish brown. In alcohol, the family (Wilkinson et al. 2011), of which
head color is brown or yellowish brown; the one species known from Guyana is
throat from snout through first collar is aquatic, with a somewhat vertically com-
yellow; and the cloacal area is cream to pressed tail.
pale yellow.
Comments.—These small caecilians gen-
erally have a body length of less than 160
Typhlonectes compressicauda
mm; primary annuli numbering 101–109;
(Duméril & Bibron, 1841)
and secondary annular grooves numbering
6–18, of which only 1–7 are complete. Type material.—The original name was
Teeth were counted on only one or two Coecilia compressicauda and MNHN 4269
specimens as follows: premaxillary and from Cayenne, French Guiana is regarded
maxillary teeth, 20–26; vomeropalatine as the lectotype (Frost 2011).
teeth, 29; and dentary teeth, 15–19. These Distribution.—‘‘Amazon Basin from the
data are similar to those for M. rabei, eastern coast of Brazil to the eastern slopes
although not identical in some compari- of Peruvian and Colombian Andes and the
sons, but one must allow for complications rivers of the Guyana Shield region’’ (Frost
owing to possible geographic variation, 2011).
small sample sizes, and data having been Vouchers for Guyana.—UMMZ 82854
recorded by different specialists. Neverthe- from ‘‘Demerara River Harauruni’’ (quot-
less, the range of primary annuli overlaps ed from the UMMZ records, although
that of M. rabei from Suriname (and cited by Taylor [1968:241] as ‘‘Hahauruni
specimens from Konawaruk Camp fall Creek, Demerara River, Guiana’’); identi-
entirely within the range for Suriname), fication recently confirmed by Mark Wil-
and although the range of data for kinson.
specimens from Venezuela is not over- Coloration in life.—The following is
lapped by specimens from Berbice River from Lescure & Marty (2000:300). Bluish
gray to brownish gray; white spot tinged in Guyana. Today, none of these is
with blue at the level of the eyes and considered to be critically endangered
cloaca. (http://www.iucncsg.org/).
Comments.—This species is aquatic.
Taylor (1968) used the UMMZ voucher
cited above for his detailed description of Alligatoridae
the species.
Caiman crocodilus (Linnaeus, 1758)
Species Accounts: Reptilia Lacerta Crocodilus and Hoogmoed &
The approximately 9100 species of Gruber (1983) selected as lectotype the
modern non-avian reptiles (Hoffmann et ZMUU specimen of a certain size and
al. 2010) are remarkably diverse in body condition they specified. The type locality
form, coloration (some being spectacular- is unknown (Duellman 2005).
ly beautiful), camouflage, life history, Distribution.—‘‘Pacific lowlands from
behavior, toxicity, and actual or potential the Isthmus of Tehuantepec, Mexico, to
use to humans. There are three major southern Ecuador; Atlantic lowlands from
groups represented in Guyana, as follows: extreme eastern Honduras to northern
Crocodylia (crocodilians; 4 species); Squa- Colombia, Venezuela, and the Guianas
mata (amphisbaenians or worm lizards, and throughout the Amazon basin to
lizards, and snakes; 157 species); and eastern Peru and central Brazil’’ (Savage
Testudinata (turtles and tortoises; 15 2002:778). The subspecies C. c. crocodilus
species). The earliest origin of reptiles occurs in ‘‘Guiana, Venezuela, Trinidad,
from an amphibian approximately 300 the Amazon drainage, eastern Colombia
million yr ago and the evolutionary to Peru’’ (Brazaitis 1973:64).
relationships of the modern species have Vouchers for Guyana.—AMNH R-
received extensive modern study, but a 32344 from Kartabo. In addition, CJC,
great deal of work remains to be done RPR (USNM field series specimen number
before we will adequately understand 207159 donated to UG), and CRT saw
these organisms. Appendix 2 provides a representatives of this species (not collect-
complete list of the species we have ed) at Dubulay Ranch and CJC and CRT
documented as occurring in Guyana. saw them (not collected) at their Berbice
River camp.
from Brazaitis (1973:64): ‘‘Dorsally olive
Species Accounts: Reptilia: Crocodylia
green [or brown] with numerous dark
Molecular and morphological analyses brown or black flecks on the head, body,
indicate that there are three major groups and tail. Dark brown or black crossbands
of crocodilians (about two dozen species) on the back and tail. Ventral surfaces are
that have survived to modern times unpatterned, uniform cream, or yellow.
(Gatesy et al. 2003, Harshman et al. No large dark blotches on the sides of the
2003, Brochu et al. 2009). These are the jaws...Hatchlings display a series of faded
alligators and caimans (primarily of the bars on the sides of the jaws, no longer
Western Hemisphere but including the visible at about 35 cm total length.’’
Chinese alligator); crocodiles (pantropical Duellman (2005:328) described the dorsal
in distribution); and the gavials or gharials coloration as ‘‘tan to pale olive-gray...Ju-
(of southern Asia). Four species (3 genera) veniles tend to be more tan...and small
of 1 family of crocodilians are represented juveniles have distinct transverse dark
brown bars on the dorsum. The iris is pale Coloration in life.—‘‘Reddish brown
brown with many black flecks.’’ above, blotched with brown on the ventral
surfaces. Very dark, nearly black individ-
uals are not uncommon. Light areas of the
Melanosuchus niger (Spix, 1825) head and body may be light brown or
Plate 19A, B yellow. The ear coverlets match the color
of the cranial table which is noticeably
Type material.—The original name was reddish brown. The dorsal surface of the
Caiman niger and ZSMH 2480/0 was snout is uniform reddish brown. Some
designated as lectotype by Hoogmoed & large red blotches on the sides of the jaws.
Gruber (1983). The type locality is ‘‘flumi- The iris of the eyes are reddish brown or
nis Amazonium et Solimoëns [Brazil]’’ orange’’ (Brazaitis 1973:68).
(Spix 1825:4). Comments.—With respect to the two
Distribution.—‘‘Eastern Colombia, Pe- species of Paleosuchus that occur in
ru, south to Bolivia, the Brazilian Amazon Guyana, Brazaitis (1973:69) stated the
basin north to Guiana’’ (Brazaitis following: ‘‘Both species appear to occupy
1973:68). the same habitat within the same distribu-
Vouchers for Guyana.—AMNH R-15171 tion. Specific niches are not fully under-
from Georgetown. In addition, CJC, RPR, stood. Many individuals display over-
and CRT saw representatives of this species lapping physical characteristics, although
(not collected) at Karanambo. P. palpebrosus shows a greater degree of
Coloration in life.—The following is from dermal ossification.’’
Brazaitis (1973:67): ‘‘Jet black above with
narrow yellow crossbands on the body and
tail, neck and base of skull light tan or Paleosuchus trigonatus (Schneider, 1801)
yellow. From three to five large dark blotches Plate 19C
on the sides of the jaws... Ear coverlets black,
as is the median dorsal surface of the snout.
Crocodilus trigonatus and ZMB 243 from
Median ventral surfaces immaculate white
South America is the lectotype (Bauer &
bordered by jet black.’’
Günther 2006).
Distribution.—‘‘South America. From
Bahia, Brazil, in the south; west to
Paleosuchus palpebrosus (Cuvier, 1807) northern Bolivia and the tributaries of
the Amazon river in eastern Peru; north
Type material.—The original name was through Colombia to Ciudad Bolivar on
Crocodilus palpebrosus and type material the Rio Orinoco, Venezuela; eastward
from Cayenne, French Guiana, may be at through Guiana, Surinam, and Mexiana
the MNHN. Island’’ (Brazaitis 1973:69).
Distribution.—‘‘The whole of tropical Vouchers for Guyana.—AMNH R-
South America, from the Guianas in the 16048 and R-64823–64825 from Kartabo;
north and Mexiana Island; south to Sao BMNH 1872.10.16.89 from Demerara
Paulo, Brazil; the Rio Pastaza in Ecuador Falls; and UMMZ 86158 from Kurupung
to the east coast’’ (Brazaitis 1973:69). River, with thanks to Colin McCarthy and
Vouchers for Guyana.—UMMZ 46110 Greg Schneider for identifying the BMNH
from Dunoon, Demerara River; UMMZ and UMMZ specimens, respectively. In
55873 from Moruca River; and UMMZ addition, CJC and CRT saw representa-
80435 from Wismar, with thanks to Greg tives of this species (not collected) at their
Schneider, who confirmed identifications. Konawaruk camp.
Coloration in life.—The following is and snakes split away from lizards. In

from Duellman (2005:329–330), based on addition, there are disagreements and new
specimens from Peru: ‘‘Adults are dull nomenclature for formal names to apply to
brown with fainter darker brown crossbars the diverse higher-level taxonomic groups
on the body and tail, the posterior third of of lizards excluding amphisbaenians and
which is tan. The venter is cream, and the snakes (see references above), as origins of
sides of the head are tan. Juveniles are the amphisbaenians and snakes within
reddish brown with irregular black trans- them could make the overall group of
verse marks on the body and tail; the snout lizards paraphyletic. Nevertheless, we refer
is black, and the parietal shelf is to lizards (excluding amphisbaenians and
brown...The iris is brown.’’ Brazaitis snakes) by their popular name, which, for
(1973:69) also noted ‘‘some individuals present purposes, allows us to discuss in
laterally tinged with yellow. Cranial table general the amphisbaenians or worm
is dark brown, ear coverlets contrasting lizards, lizards, and snakes as separate
black or darker brown. A dark brown or groups within Squamata.
black median dorsal stripe on the snout. The amphisbaenians (worm lizards) are
Ventral surfaces densely blotched with highly adapted to burrowing, and most
brown. Iris of the eyes reddish brown or species are limbless. They have a solid,
orange.’’ compact skull, ‘‘an elongated body, short
Comments.—Pritchard (1995:43) men- tail, and a unique modification of the inner
tioned an individual (photo PCHP 002
ear that allows the detection of low
deposited at UF) that he saw emerge from
frequency sounds’’ (Kearney 2003:1). They
the sea and rest on Almond Beach near
‘‘burrow in loose or sandy soils [and
Waini Point, Guyana. He also noted that
riverbanks] and ostensibly come to the
this was the ‘‘first marine record for
surface only rarely...Many species are only
species; closest records are from Lower
found serendipitously after heavy rains or
Demerara and Lower Essequibo rivers,
Guyana (Medem 1983, Crocodylia de Sur when plowing or moving the soil. The
America, Vol. 2:95).’’ ecology and life history of amphisbaenians
are poorly known due to their fossorial
lifestyle and the scarcity of researchers
studying the group’’ (Kearney 2003:2). A
Species Accounts: Reptilia: Squamata: recent phylogenetic analysis of many
Amphisbaenians or Worm Lizards Brazilian amphisbaenians concluded that
The Squamata is the largest group of several genera recognized previously
reptiles (more than 8500 species) and should be synonymized with the genus
includes the beak heads or tuataras of Amphisbaena (Mott & Vieites 2009). Pres-
New Zealand, amphisbaenians or worm ently, 1 family, 1 genus, and 4 species of
lizards, lizards, and snakes. Several mod- amphisbaenians are known to occur in
ern studies based on phylogenetic analyses Guyana.
of morphology and/or DNA sequence
data have shown clearly that the amphis-
baenians and snakes are both monophy- Amphisbaenidae
letic groups that have evolved from within
the lizards (Kearney 2003, Townsend et al. As for the species from Surinam
2004, Vidal & Hedges 2005, Conrad 2008, (Hoogmoed 1973:371), the species from
Wiens et al. 2010). However, there remain Guyana are ‘‘legless, having reduced eyes
some uncertainties as to phylogenetically covered by the oculars and the scales
where the lineages of the Amphisbaenia arranged in annuli [around the body].’’
Amphisbaena alba Linnaeus, 1758 AMNH R-137423–137424 from Kartabo;

AMNH R-140977, R-141875, and USNM
Type material.—Gans (1967) cited syn- 566299 from Dubulay Ranch; and USNM
types from America in the Museum 164181 from Mabaruma Compound
Drottningholm. Hoogmoed (1973) re-
(8812 0 N, 059847 0 W).
stricted the type locality to the confluence
of the Cottica River and Perica Creek,
based on the AMNH and USNM spec-
Surinam.
imens from Dubulay Ranch (CJC field
Distribution.—‘‘Forested lowlands of
South America. . .Venezuela, Trinidad, notes, 1994). They are black and white,
and the Guianas; Colombia, Peru, and with more black above, more white
Bolivia east of the Andes; Brazil and below.
northern Paraguay’’ (Gans 1962:16).
Vouchers for Guyana.—AMNH R-
61223 from the Kuyuwini River and
Amphisbaena stejnegeri Ruthven, 1922
AMNH R-21279 from Kartabo.
Coloration in life.—‘‘Light- to dark- Type material.—The holotype is
brown above, fading to a dirty white on UMMZ 55858, from a sand reef at
the ventral parts (Gans 1969). Juvenile Vreeden Rust, Demerara River, Guyana.
specimens show white spots in the brown Distribution.—Known only from the
dorsal area, which, in adults, is uniform. type locality (Gans 1963:13).
Tip of the tail entirely white’’ (Hoogmoed Vouchers for Guyana.—The holotype is
1973:376–377). Beebe (1945:29) described the only known specimen having locality
a specimen from Kartabo as follows. data, and only one other specimen, with-
‘‘Snout back to the eyes and the chin out data, exists (Gans 1963).
shields pale pink. Above dark brownish-
Coloration.—The only color notes are
yellow shading to old ivory below, the tail
for specimens in preservative, which are
mottled with dead white. Eye showing
‘‘yellowish, with an irregular dorsal scat-
pink beneath its scale.’’
tering of individual dark segments...[more
so on one specimen than the other],
thinning out anteriorly toward an immac-
Amphisbaena fuliginosa Linnaeus, 1758 ulate head. The anterior half of some
Plate 19D, E [most] infracaudal segments is strongly
Type material.—Gans (1967) cited syn- pigmented, while the infracaudal segments
types in the Stockholm Museum and of the ninth postcloacal annulus are
Uppsala Museum from America. Hoog- entirely pigmented’’ (Gans 1963:6–7).
moed (1973) restricted the type locality to
the confluence of the Cottica River and
Perica Creek, Surinam. Amphisbaena vanzolinii Gans, 1963
Distribution.—‘‘Surinam, Guyana,
French Guyana, Venezuela’’ (Hoogmoed Type material.—The holotype is
1973:383). Vanzolini (2002:625) included AMNH R-60975, a female from ‘‘Marudi,
Brazil, Colombia, Ecuador, and Peru. British Guiana... Charles M. Bogert in-
Vouchers for Guyana.—AMNH R-1085 forms me that W. Hassler, who went over
from Kaieteur Falls; AMNH R-7295 and the route of the expedition, placed the
R-17688–17689 from Georgetown; locality as ‘on a tributary of the Kuyuwini
AMNH R-25088 from Kamakusa; River, at 285 0 North and 598 East’’’ (Gans
AMNH R-60907 from Marudi River; 1963:13).
Distribution.—Brazil, Guyana, and Sur- that occur primarily in the Western Hemi-
inam (Hoogmoed 1973, Hoogmoed & sphere. Frost & Etheridge (1989) conduct-
Mott 2003). ed the first cladistic analysis of the group
Vouchers for Guyana.—The holotype; as a whole, and they partitioned the large
the paratopotype AMNH R-60778; and traditional family into eight smaller fam-
KU 69817–69818 from ‘‘Rupununi Dis- ilies, many of which had been recognized
trict, N. of Acarai Mountains, W. of New previously as natural groups below the
River’’ (Hoogmoed 1973:384). family level. By recognizing the eight
Coloration.—The following is based on families, Frost & Etheridge (1989) empha-
specimens from Surinam (Hoogmoed sized that they had strong evidence that
1973:387). ‘‘RMNH 15144 from Paloemeu each of these families represented a mono-
was purplish all over, slightly darker phyletic group of genera and species;
dorsally than ventrally. The specimens however, this also emphasized that at the
from Sipaliwini were light- to dark-brown time they could not establish firmly the
above, with a transparent pink belly, the monophyly of the larger group of those
underside of the tail white. In preservative eight families with respect to other lizards.
RMNH 15144 is dirty white all over, with Recently, the monophyly of the eight
a brown tinge on the dorsal parts. All families partitioned from the larger, tradi-
Sipaliwini specimens are light- to dark- tional Iguanidae has received improved
brown above, white below.’’ support (e.g., Schulte et al. 2003, Town-
Comments.—‘‘All Surinam specimens send et al. 2004, 2011), and some authors
(except RMNH 15191) were collected in have suggested returning to the original
rotten logs lying on the rain-forest floor. concept of one large family (e.g., Schulte et
RMNH 15191 (from Sipaliwini) was found al. 2003), with the eight groups of Frost &
while digging in the upper soil-layers. Only Etheridge (1989) being treated as subfam-
a special type of rotten log seems to be ilies. Both versions of family groupings
favoured by this species, namely logs that proposed (Frost & Etheridge 1989 vs.
are very soft and when pulled apart Schulte et al. 2003) are consistent with
rendered a pulp of small wood-fragments’’ formal taxonomic and nomenclatural pro-
(Hoogmoed 1973:387). cedures, so neither is conceptually flawed
or necessarily wrong. However, the ar-
rangement of families proposed by Frost &
Species Accounts: Reptilia: Etheridge (1989) has received such wide-
Squamata: Lizards spread use, including among authors
writing about Guianan taxa (e.g., Avila-
Lizards are non-avian, non-crocodilian, Pires 1995, Gorzula & Señaris 1999 [1998],
and non-testudinatan reptiles (without a Savage 2002, Conrad 2008), that we follow
bony shell) that, for most species, are their usage of multiple smaller families.
terrestrial or arboreal and have four legs Most recently, the earlier Iguanidae has
and a tail. However, some lizards that lack been divided up into even more than eight
limbs, and some of these and others are smaller families (Townsend et al. 2011).
burrowing organisms. At the moment, at The following four families of iguanian
least 9 families, 34 genera, and 56 species lizards have representatives living in Guya-
of lizards are known to occur in Guyana. na: Corytophanidae (introduced by hu-
mans), Iguanidae, Polychrotidae, and
Iguanian Lizards Tropiduridae. Consistent with our treat-
ment of families in this volume, these
Traditionally, until 1989, the Iguanidae groups are considered below in alphabet-
was treated as a very large family of lizards ical order by family. However, see Appen-
dix 1 for a most recent splitting of CJC by M. S. Hoogmoed, according to

Dactyloidae (for Anolis) from the Poly- whom (pers. comm.), Mr. Hoevers ‘‘said
chrotidae (Townsend et al. 2011). there was a reproducing population at the
airport.’’ The presence of this population
should be confirmed.
Corytophanidae
Representatives of this family of igua- Gekkonidae
nian lizards occur in southern ‘‘Mexico,
southward through Central America, to Until recently, many genera of geckoes
Ecuador and Venezuela’’ (Frost & Ether- were included in the large family Gekko-
idge 1989:35). One species has been intro- nidae (Gray 1825). However, recent phy-
duced to Guyana by humans (see logenetic analyses (Gamble et al. 2008),
Comments for Basiliscus). including considerable DNA sequence
data, indicate that the geckoes of Guyana
should now be considered as representative
Basiliscus basiliscus (Linnaeus, 1758) of the two following families: Gekkonidae
(with Hemidactylus and Thecadactylus)
Type material.—The original name was and Sphaerodactylidae (with Chatogekko,
Lacerta Basiliscus and the type material Coleodactylus, Gonatodes, Pseudogonato-
from South America is not known to us. des, and Sphaerodactylus).
Distribution.—‘‘Lowlands and adjacent
slopes on the Pacific versant from south-
western Nicaragua to northwestern Co- Hemidactylus mabouia
lombia, and on the Atlantic versant from (Moreau de Jonnès, 1818)
central Panama to northwestern Venezue- Plate 20A
la’’ (Savage 2002:430). Avila-Pires
(2005:33) stated that this species was Type material.—The original name was
introduced into Guyana by humans (see Gecko Mabouia and the holotype is
Comments, below). MNHN 6573 (Avila-Pires 1995:229). The
Vouchers for Guyana.—RMNH 40127 type locality was restricted to St. Vincent
from Timehri Airport, identified by M. S. Island, Lesser Antilles, according to Avila-
Hoogmoed. Pires (1995).
Coloration in life.—The following is Distribution.—Africa south of 108N,
based on specimens from Costa Rica Madagascar, the Antilles and South Amer-
(Savage 2002:428–429). Adults with ‘‘body ica, where it occurs in Brazil the Guianas,
brown, olive, or bronze, with darker cross- Ecuador, and Peru (Avila-Pires 1995:233).
bands and usually cream to yellow light The species has been introduced to other
stripes along lips and a pair of similarly localities in the Western Hemisphere also,
colored lateral stripes. Juveniles: yellowish although it was originally a ‘‘native to sub-
green with brown cross-bands, lip and Saharan Africa’’ (Bugoni & Welff-Neto
lateral stripes usually present, all markings 2008:226).
more vivid than in adults; throat with three Vouchers for Guyana.—AMNH R-
distinct longitudinal light lines. . .; iris 137421 from Kartabo; AMNH R-
brown to bronze in all age-classes.’’ 140930–140936, R-141899 and USNM
Comments.—The RMNH voucher was 566313–566314 from Dubulay Ranch;
collected by L. G. Hoevers in ‘‘swampy USNM 162983–162999 from Enmore Es-
bush’’ (quoted from RMNH records). A tate (6844 0 N, 057859 0 W); and USNM
photograph of the specimen was sent to 291268 and 497791 from Georgetown.
Coloration in life.—The following is Thecadactylus rapicauda (Houttuyn, 1782)

based on AMNH and USNM specimens Plate 20C, D
from Dubulay Ranch (CJC field notes,
1994, 1995). Very light tan, yellowish tan, Type material.—The original name was
or grayish tan on body, arms, legs, and tail Gekko Rapicauda and type locality was
(the latter lighter), with dark gray, tan, restricted to Paramaribo, Surinam by
brown, or black bands, which may be Hoogmoed (1973). Avila-Pires (1995:239)
inconspicuous; venter translucent light stated that the ‘‘holotype is unknown, but
gray, cream, or yellow. possibly still preserved in the Zoological
Museum Amsterdam’’ (M. S. Hoogmoed,
pers. comm.).
Distribution.—‘‘Venezuela, Guyana,
Hemidactylus palaichthus Kluge, 1969 Suriname, French Guiana, Brazil, on both
Plate 20B sides of the Andes in Ecuador and
Colombia, on eastern side in Peru and
Type material.—The holotype is Bolivia; Central America north to Mexico;
AMNH R-60931, an adult male from Lesser Antilles’’ (Avila-Pires 1995:245).
Kurupukari, Guyana. Vouchers for Guyana.—AMNH R-
Distribution.—This species is known 8106–8109 from Kalacoon; AMNH R-
only from the Western Hemisphere, from 21302–21304 and R-46437–46438 from
‘‘northern Brazil, coastal Suriname, Guya- Kartabo; AMNH R-60914–60915 from
na, central and northeastern Venezuela, Isheartun; AMNH R-60927–60928 from
and on the islands of Trinidad and St. Onora; AMNH R-61382 from Marudi;
Lucia’’ (Avila-Pires 1995:237). AMNH R-140956–140959 from Dubulay
Vouchers for Guyana.—The holotype; Ranch; AMNH R-151743, R-151745, and
AMNH R-60900–60906 from Isheartun USNM 566331 from Berbice River Camp;
(on the upper Rupununi River, about 30 and AMNH R-151746–151753, R-151755,
km [linear] NW Kuyuwini Landing); and USNM 566332–566336 from Kona-
AMNH R-137925–137926, R-137929, R- waruk Camp. Also USNM 84486 from
137931–137938, R-137940 and USNM Tumatumari; USNM 164184–164185 from
566315–566319 from Karanambo; AMNH M a b a r u m a C o m p o u n d ( 8 81 2 0 N ,
R-138064 from Yupukari; AMNH R- 059847 0 W); USNM 531640–531641 from
139741–139753, 139759–139760, and Iwokrama Forest Reserve; USNM
USNM 566320–566326 from Aishalton; 561449–561452 from CEIBA Biological
AMNH R-140948 from Dubulay Ranch; Center, 26 mi S of Georgetown on Linden
ROM 20521–20524 from Kurupukari. Highway (06829 0 57 00 N, 058813 0 06 00 W).
Coloration in life.—The following is Coloration in life.—The following is
based on AMNH and USNM specimens based on AMNH specimens from Dubulay
from Karanambo, Aishalton, and Dubu- Ranch (CJC field notes, 1994). Yellowish
lay Ranch (CJC field notes, 1992–1994). tan above with brown markings (all dorsal
Gray, pinkish gray tan or dark brown surfaces); ventral surfaces yellow or very
above (including body, arms, legs, tail) pale greenish yellow; regenerated tail (above
with grayish brown or dark gray, brown, and below) tan with dark brown markings.
or black markings (mostly bands) that are
edged behind by light gray; ventral surfac-
es cream or pale yellow, with or without Gymnophthalmidae
gray or tan specks, but with gray or brown
flecks or mottling beneath tail, which may Until recently, the microteiid and mac-
otherwise be gray or yellow. roteiid lizards were all included within the
large family Teiidae. However, recent Alopoglossus angulatus (Linnaeus, 1758)

phylogenetic analyses, including consid- Plate 20E
erable data from DNA sequencing, have
shown that the microteiids probably are a Type material.—The original name was
monophyletic group that can be treated as Lacerta angulata and Hoogmoed (1973)
a separate family (Gymnophthalmidae), designated RMNH 15200, a female, as the
and its closest relatives are the Teiidae neotype. This specimen is from Brown’s
(Pellegrino et al. 2001, Castoe et al. 2004). Mountain, Surinam.
Distribution.—‘‘Amazonian region in
Recent revisionary work has shown that
Brazil..., French Guiana, Suriname, Guya-
one Guyanan species that formerly was
na, Colombia, Ecuador and Peru’’ (Avila-
assigned to the genus Prionodactylus is
Pires 1995:313).
actually a member of the genus Cerco-
saura (C. argulus), and four new species
61381 from Marudi; and USNM 566400
and genera known only from Guyana
from Berbice River Camp.
have been described recently (Arthrosaura
hoogmoedi, Echinosaura sulcarostrum,
Kaieteurosaurus hindsi, and Pantepuisau-
Berbice River Camp (CJC field notes,
rus rodriguesi).
1997). All dorsal surfaces dark brown,
There are approximately 200 species in with very inconspicuous dark tan dorso-
approximately 45 genera of gymnoph- lateral stripes, most visible on the neck and
thalmids, and they occur primarily in as spots on the hips and base of tail; all
South America. The group is exceedingly ventral surfaces very pale yellow, with
interesting because of its diversity, fre- dark gray to black spots on preanal area
quent examples of convergent evolution, and base of tail. The following is from
and limited geographic ranges of many Hoogmoed (1973:222), based on speci-
species. For taxa occurring in Guyana, mens from Suriname. ‘‘In life dark-brown
there are examples of unisexual (all- above with faint orange-brown dorsolat-
female) species that reproduce by par- eral stripes, tail with light-brown and dark-
thenogenetic cloning (by means of un- red spots. Ventral parts dirty white with
fertilized eggs), and species with extreme two dark-brown spots on the preanal
elongation of the body, loss of limbs, plate. Iris green. Tongue bluish black.’’
and loss of eyelids, owing to multiple Avila-Pires (1995) stated that males also
(convergent) origins of a burrowing life have a light lateral stripe that is absent in
style. Most of the species, while being females.
immensely interesting, are small and
cryptic in the habitats in which they
live, having secretive lives that are very
Arthrosaura guianensis
difficult to study. As a consequence, of
MacCulloch & Lathrop, 2001
the 21 Guyanan species, 12 are known
from only 5 specimens or fewer that Type material.—The holotype is ROM
have been found in Guyana, and about 39471, an adult male from the northeast
half of these are endemic to Guyana. In plateau of Mount Ayanganna, Guyana
addition, some genera (e.g., Gymnoph- (05824 0 N, 059857 0 W; 1490 m).
thalmus and Leposoma) have taxonomi- Distribution.—This species is known
cally cryptic species, which are only from Guyana.
morphologically so similar to one anoth- Vouchers for Guyana.—Only two speci-
er that they are difficult to identify mens are known in addition to the
correctly to species. holotype. These are ROM 39472 from
the type locality and USNM 549323 from about midbody; flanks black, lower part of
the north slope of Mount Roraima flanks peppered with reddish brown, and
(5817 0 N, 060845 0 W; 700 m). with some small pale brown spots ventro-
Coloration in life.—‘‘Dark brown dor- laterally, forming an irregular ventrolater-
sally and laterally, with a light dorsolateral al line. Axilla region and underside of arms
line from above the eye to the groin. A fine marbled with reddish brown and black.
line of small white dots from neck to Head blackish brown with tan dorsolateral
midbody, parallel to dorsolateral line and lines extending above eyes, from the
passing just above forelimb. A second row anterior part of the loreals to the parietals.
of similar spots, also parallel to dorsolat- Tip of snout paler; diffuse paler blotches
eral line, extending from behind forelimb on lower part of lateral surface of head.
insertion to about halfway to the hindlimb. Tympanum black, surrounded by some
Chin white, throat white with orange tan scales. Arms black, legs and tail
wash. Venter and underside of limbs and blackish brown. Iris black...Tongue dark
tail reddish-orange, fading to white distally gray. Palms and soles black. Chin, throat,
on limbs and tail. Dark mottling extending chest, belly and ventral surface of upper
from lateral surfaces onto ventral surfaces, legs [arms] cream with dark mottling;
decreasing in intensity so that midventral lower legs ventrally reddish brown with
areas are immaculate or nearly so; dark dark mottling; tail ventrally black, except
mottling most intense in gular region. proximally where it is dirty white peppered
Underside of tail light with dark mottling with black’’ (Kok 2008:11).
proximally, becoming darker distally.
Some irregular light mottling on sides of
neck. Upper lips dark, with two vertical Arthrosaura reticulata
white bars below eye. Diffuse light mark- (O’Shaughnessy, 1881)
ings on anterior supralabials, frenocular, Plate 21A, B
and first subocular’’ (MacCulloch & Lath-
rop 2001:178). Type material.—The original name was
Cercosaura (Pantodactylus) reticulata and
the holotype is BMNH 1946.9.1.5 (Avila-
Arthrosaura hoogmoedi Kok, 2008 Pires 1995:335) from Canelos, Ecuador.
Distribution.—‘‘Amazonian region in
Type material.—The holotype is IRSNB Brazil, French Guiana, Suriname, Guya-
2653, an adult female from the summit na, southern Colombia, Ecuador and
plateau of Mount Maringma, Cuyuni- northern Peru’’ (Avila-Pires 1995:342).
Mazaruni District, Guyana (5812 0 60 00 N, Vouchers for Guyana.—AMNH R-
60835 0 06 00 W; 2112 m). 57454 from Kuyuwini Landing; AMNH
Distribution.—Known only from the R-70623 from Isheartun; AMNH R-
type locality. 152005–009, R-152016, and USNM
Vouchers for Guyana.—This is another 566421 from Berbice River Camp.
of the least known species in the world, as Coloration in life.—The following is
it is known from only the type specimen. based on AMNH and USNM specimens
Coloration in life.—‘‘Dorsum dark from Berbice River Camp (CJC field notes,
brown, with a black middorsal stripe 1997). Dark brown above, with three
(about two–three dorsal scales wide) ex- lighter brown stripes (one, the vertebral
tending from nape to tail, where it quickly stripe, is very irregular on one individual,
becomes inconspicuous due to the dark and all stripes can be very inconspicuous).
coloration of the tail; subtle, irregular, tan Light gray spots on sides of lips and neck;
dorsolateral line extending from nape to light tan spots on sides of body. Top of
head, arms, and legs very dark brown; all males have white to very pale blue spots on
ventral surfaces bright orange, throat with the preanal scales, posterior abdominals,
black spots. Hind legs above with incon- and basal subcaudals.
spicuous tan dots. The coloration given by Comments.—Avila-Pires (1995) dis-
Hoogmoed & Avila-Pires (1992:466) is cussed taxonomic problems and conclu-
somewhat different, but within the realm sions concerning this locally and
of individual and geographic variation. geographically variable species.
They also noted the following: ‘‘iris brown,
brown with an orange rim, or golden;
tongue anteriorly blue-grey, posteriorly
pink-white. Adult males may have brightly Cercosaura argulus Peters, 1863a
orange undersides, possibly related to Plate 21E
breeding condition.’’
Cercosaura (Pantodactylus) argulus and
the holotype is ZMB 4555 from ‘‘moun-
Bachia flavescens (Bonnaterre, 1789) tainous regions around Santa Fé de
Plate 20F Bogotá, Cundinamarca, Colombia’’ (Avi-
la-Pires 1995:453).
Type material.—The original name was Distribution.—‘‘French Guiana, Colom-
Chalcides Flavescens based on material bia, Ecuador, Peru, Bolivia, and Brazil’’
from an unknown locality. Avila-Pires (Avila-Pires 1995:458).
(1995:344) stated that the type was in the
MNHN, but was lost.
21265 from Bartica; AMNH R-61386 from
Distribution.—‘‘Brazil, French Guiana,
Marudi; and AMNH R-61434 from Shu-
Suriname, Guyana, and Colombia’’ (Avi-
dikar-wau, headwater stream, Essequibo
la-Pires 1995:349).
River.
Coloration in life.—The following notes
137365–137367 and R-137369 from Kar-
tabo; AMNH R-140923–140927 and are based on AMNH R-119427 from
USNM 566422 from Dubulay Ranch; Brownsberg Nature Park, Suriname (CJC
AMNH R-151916–151925 and USNM field notes, 1980). Dorsum, posteriorly
566423–566426 from Berbice River Camp; (including tail), brown; head and neck
AMNH R-151931–151932 and USNM gray, grading into the brown dorsum
566427 from Konawaruk Camp; ROM posterior to shoulder; three largely com-
20515 from Tukeit; and USNM 85012 plete rows (vertebral and dorsolateral) of
from Pickersgill, Pomeroon River. tiny, black dots on back, fading on base of
Coloration in life.—The following is tail; dark brown lateral stripe on body
based on AMNH and USNM specimens originating at eye, followed below by a
from Dubulay Ranch, Berbice River yellow stripe, in turn followed below by a
Camp, and Konawaruk Camp (CJC field thin black line originating anterior to arm;
notes, 1994–1998). Brown above, with on body above arm are two tiny white dots
dark tan dorsolateral stripe bordered encircled with black; one cream dot on
above and sometimes below by broken each elbow plus one above humerus;
(dashed) fine dark brown or black lines; venter beige, lightest anteriorly.
venter translucent pinkish gray or light to Comments.—Previously, this species was
medium brown on body, brown on tail. treated as being in the genus Prionodacty-
Dorsally, the coloration may be intensified lus, but Doan (2003) showed that this
on the base of the tail. Ventrally, adult genus is a junior synonym of Cercosaura.
Cercosaura ocellata Wagler, 1830 Distribution.—Known only from Guya-

Plate 21D na.
Type material.—Avila-Pires (1995) stat- ROM 22892 and ROM 22894 from the
ed that the holotype is ZSMH 643/0 from type locality; and ROM 43805 from
‘‘probably somewhere in northeastern Mount Wokomung.
South America; Surinam?’’ (Ruibal Coloration in life.—‘‘Dark brown dor-
1952:494). sally and laterally, with pale yellow spots.
Distribution.—‘‘Eastern Amazonia, in Spots join to form short longitudinal
Guyana, Suriname, French Guiana, and stripes on the nape and transverse blotches
Brazil’’ (Avila-Pires 1995:364). posteriorly. Venter mottled brown/yellow;
Vouchers for Guyana.—AMNH R- chin and throat more yellow than brown,
21264 from Kartabo; AMNH R-61385 brown increasing posteriorly. Underside of
from Marudi; AMNH R-139827–139830 limbs pale yellow with brown mottling.
and USNM 566428 from Aishalton; Proximal portion of ventral tail brown
AMNH R-140947 from Ceiba Biological with yellow spots. Regenerated portion of
Station (Madewini River, ca. 5 km. E tail uniform reddish brown’’ (Donnelly et
Timehri Airport; 06829 0 57 00 N, 058813 0 al. 2006:400).
06 00 W); AMNH R-141877 from Dubulay
Ranch; ROM 28352 from Paramakatoi;
and UMMZ 46771–46772 from Dunoon, Gymnophthalmus leucomystax
Demerara River. Vanzolini & Carvalho, 1991
Coloration in life.—The following is Plate 21C
from Aishalton, Ceiba Biological Station, Type material.—The holotype is
MZUSP 69301 from Fazenda Salvamento,
and Dubulay Ranch (CJC field notes,
Roraima, Brazil.
1993–1995). Dorsum brown or yellowish
Distribution.—Known only from savan-
dark tan; orange lateral stripe or incon-
na regions in the Roraima area of northern
spicuous tan or cream dorsolateral stripe;
Brazil and southern Guyana (within or
sometimes black stripe to each side of
near the Rupununi Savanna).
dorsolateral stripe; sides green below this; Vouchers for Guyana.—AMNH R-
head, arms, and legs brown or tan; 131991–93 from Isheartun; and AMNH
sometimes black flecks on lips; chin, R-139856–139860, AMNH R-139863–
throat, and chest white or cream; abdomen 139866, and USNM 566453–566454 from
and beneath arms, legs, and base of tail Aishalton; both males and females are
grayish tan or yellow; tail of adult male represented.
with orange on sides, pale yellow ventrally Coloration in life.—The following is
at base, grading into orange posteriorly. based on AMNH and USNM specimens
from Aishalton (CJC field notes, 1993).
Dorsum grayish brown to shiny tan;
Echinosaura sulcarostrum Donnelly, dorsolateral light stripe distinct, yellow
MacCulloch, Ugarte, & Kizirian, 2006 or pale gold anteriorly, becoming cream
Plate 21F or tan posteriorly, continuous to base of
tail; black stripe above and alongside the
Type material.—The holotype is ROM light stripe, becoming line of black dots
22893, an adult male from Guyana, posteriorly; sides very dark brown to
Baramita, approximately 1 km S airstrip black; cream to tan lateral (or essentially
(07822 0 N, 060829 0 W; 100 m). ventrolateral) light stripe, on some pres-
ent only anterior to arm, on others Comments.—The more we study Gym-

extending to midbody or hip; tail dark nophthalmus, the more we realize that
gray or bluish gray above (although may there are morphologically similar, cryptic
be tan at base), grayish blue below; arms species that are difficult to distinguish
and legs brown; females with chin and from each other and which usually have
throat white, other ventral surfaces (ex- rather restricted geographic ranges. Con-
cluding tail) gray with tiny black dots; sequently, most recent authors doubt that
adult males orange below. a single species, G. speciosus, actually
Comments.—This species occurs on the ranges from southern Mexico to northern
southern part of the Rupununi Savanna in South America (e.g., Cole et al. 1990,
the vicinity of Aishalton, where the habitat Vanzolini & Carvalho 1991, Carvalho
is similar to that of localities where it 1997, Kizirian & Cole 1999). Neverthe-
less, specimens from bisexual (gonochor-
occurs in Brazil.
istic) populations in northern South
America are similar to those from Nicar-
agua, and there are no clear-cut objective
Gymnophthalmus cf. speciosus data yet for concluding which populations
(Hallowell, 1861) should be named as a separate species. In
Plate 22C fact, there may be several species mas-
querading under the name G. speciosus
Type material.—The original name was within its broad range. This will be
Blepharactisis speciosa and the type spec- resolved when somebody does careful
imen(s) from Nicaragua are unknown to comparative analyses of samples from
us. throughout the range, including molecu-
Distribution.—Southern Mexico, Cen- lar work. The first work to be done with
tral America, and northern South America DNA sequence data showed that samples
(Savage 2002:522), but see Comments, from Dubulay Ranch were most closely
below. related to those from Chacachacare Is-
Vouchers for Guyana.—AMNH R- land, Trinidad, that the next closest
140974–140975 from Dubulay Ranch; relative was from the Rupununi Savanna
males. (now known as Gymnophthalmus vanzoi,
Coloration in life.—The following is see account and Comments below), and
that these might be paraphyletic with
based on the AMNH specimens from
respect to a sample from Venezuela
Dubulay Ranch (CJC field notes, 1994).
(Kizirian & Cole 1999). No samples from
Dorsum copper; tip of snout very pale tan;
Central America or Mexico were available
sides, arms, and legs very dark brown
for that study.
(sides almost black); dorsolateral stripe
light tan but posterior to midbody con-
tinuing to base of tail as dots; just above
dorsolateral stripe on neck is a short series Gymnophthalmus underwoodi Grant, 1958
of black dots; tail tan or reddish tan above Plate 22A, B
with gray iridescence (when viewed on an Type material.—The holotype is
angle), having light tan dots aligned UIMNH.42334 from Barbados.
around tail appearing as inconspicuous, Distribution.—Specimens identified with
incomplete bands; belly cream or very pale certainty are known from islands in the
tan, with tan or metallic copper irides- West Indies, Guyana, Suriname, and
cence; beneath tail tan becoming pale gray French Guiana (Cole et al. 1990). The
posteriorly. species is expected to occur in Venezuela,
but no records are known as yet. For occurred in one generation. Analyses of
Brazil, specimens known may include both DNA sequence data (more than 1800
this species and an un-named cryptic nucleotides) indicated that specimens from
species, which requires additional molecu- Guyana, Suriname, Trinidad, and the type
lar analyses to confirm (see Comments, locality are of one clone (Kizirian & Cole
below). 1999), so the name G. underwoodi is
Vouchers for Guyana.—AMNH R- correctly applied to these populations.
137971–137972, R-137987–137993, R- At least two cryptic, unisexual species
137999, R-138010–138028, and USNM are masquerading under the name G.
566468–566482 from Karanambo and Yu- underwoodi, and there are also cryptic,
pukari; AMNH R-139879 from Aishalton; bisexual (gonochoristic) species of which
AMNH R-140976, R-141867–141868, R- males (or possibly hybrids) have been
141870, R-141872–141874, and USNM misidentified as G. underwoodi (discussed
566483–566485 from Dubulay Ranch; by Cole et al. 1990). This is understandable
AMNH R-151933–151941 and USNM because the species are difficult to identify,
566486–566489 from Berbice River Camp; especially on the basis of external mor-
and AMNH R-152338 from Ceiba Biolog- phology of specimens that have been
ical Station, Madewini River, ca. 5 km preserved for a long time. Of special
(linear) E Timehri Airport (06829 0 57 00 N, interest is the Brazilian sample from Ilha
058813 0 06 00 W). da Maracá, Roraima, which Cole et al.
Coloration in life.—The following is (1990:12–15) referred to as ‘‘G. under-
based on AMNH and USNM specimens woodi?’’ On the basis of a few distinctive
from Karanambo (and Yupukari), Aishal- morphological characters, those authors
ton, Dubulay Ranch, and Berbice River stated: The lizards ‘‘could represent a
Camp (CJC field notes, 1992–1997). Dor- distinctive clone of G. underwoodi or they
sum dark copper, extending well onto tail, could be a different species.’’ Subsequent-
may become grayish tan or tan with gray ly, Yonenaga-Yassuda et al. (1995) showed
iridescence; sides, arms, and legs dark that these Brazilian lizards also have a
brown to black; light tan dorsolateral karyotype rather different from that of G.
stripe fading posterior to shoulders; belly underwoodi, so it would be appropriate to
gray or bluish gray (paler anteriorly), with consider them as a separate species after
light coppery iridescence; posterior tail all. In fact, the karyotype of the unisexual
gray or bluish gray, especially below. Brazilian sample is very similar to that of
Comments.—This is a diploid, unisexual G. cryptus presented by Cole et al. (1993).
(all-female) species that had a hybrid CJC compared 39 recently collected
origin (sympatric speciation) and of which representatives of all known populations
females reproduce by parthenogenetic from Guyana with the Brazilian ones for a
cloning (Hardy et al. 1989, Cole et al. more-or-less diagnostic morphological
1990). The trigger for embryogenesis is character (number and size of infralabial
unknown. The unisexual lineage arose scales from tip of snout to posterior edge
from a female hybrid between Gymnoph- of eye; Cole et al. 1990). Of these, 37 were
thalmus cryptus 3 G. cf. speciosus in the typical of true G. underwoodi; 1 out of 7
Guianan Region (Cole et al. 1993), and the from Dubulay Ranch (AMNH R-141874;
female parent of the hybrid was the G. mixed habitat) was like underwoodi on the
cryptus (see Kizirian & Cole 1999). Be- right side but similar to Maracá specimens
cause members of the clone possess the F1 on the left, presumably owing to individual
generation combination of parental genes, variation; and the only specimen from
the switch from gonochoristic reproduc- Aishalton (AMNH R-139879; savanna
tion to parthenogenetic cloning must have habitat) was similar to Maracá specimens
(of forest habitat) on both sides. Addi- legs darker coppery brown (not black);
tional comparative work, including more very pale copper dorsolateral stripe, in-
extensive molecular analyses and direct conspicuous or absent past midbody but
comparisons of Brazilian and Guyanan sometimes continuing to hip; belly pale
samples are needed to confirm that the copper or coppery gray, but brighter in
unisexual populations in these two coun- adult males, which may have bright orange
tries had separate and independent hybrid on chin and throat (white, copper, or pale
origins and to determine the range of the gray on some lizards); tail posterior to base
species found at Ilha da Maracá. bright orange or salmon, especially in
At Dubulay Ranch, G. underwoodi is juveniles and females; regenerated tails
sympatric with G. cf. speciosus. At Kar- and those of males were noted as being
anambo and Yupukari it is sympatric with tan or brownish gray (gray below), not as
Gymnophthalmus vanzoi (see below). At bright as in juveniles and females.
Aishalton, it is sympatric with both G. Comments.—This diploid, bisexual
leucomystax and G. vanzoi (see below). (gonochoristic) species is very similar to
Because sympatry is rare among species of G. speciosus and specimens from northern
Gymnophthalmus and because G. under- Brazil and elsewhere have been identified
woodi had an origin involving interspecific as G. speciosus in the past (discussed by
hybridization, detailed studies of ecology Carvalho 1997). The similarities between
and behavior and the potential for hybrid- G. cf. speciosus and G. vanzoi in Guyana
ization should be conducted at localities and elsewhere are so great that the species
where there are sympatric species of are not adequately diagnosed as yet.
Gymnophthalmus.
Iphisa elegans Gray, 1851

Gymnophthalmus vanzoi Carvalho, 1997 Plate 22E
Plate 22D
Type material.—The holotype is 1946.9.1.1, a male (Avila-Pires 1995) from
MZUSP 72598, a male from Brasil; within a circuit of about 300 miles of
Roraima: Fazenda Salvamento, Rio Urar- Belém, Para, Brazil (Avila-Pires 1995).
icoera (03820 0 N, 61824 0 W). Distribution.—‘‘Amazonia and some pe-
Distribution.—Known only from savan- ripheral areas, in Brazil, French Guiana,
na regions in the Roraima area of northern Suriname, Guyana, southeastern Colom-
Brazil and southern Guyana (within or bia, Ecuador, Peru, and Bolivia’’ (Avila-
near the Rupununi Savanna). Pires 1995:388).
Vouchers for Guyana.—AMNH R- Vouchers for Guyana.—AMNH R-
137973 from Karanambo; AMNH R- 21294 and R-137363 from Kartabo;
138029–138034, R-138052–138056, and AMNH R-70624 from Kuyuwini Landing;
USNM 566455–566464 from Yupukari; AMNH R-151946–151949 and USNM
and AMNH R-139867–139871, R- 566490–566491 from Berbice River Camp;
139875–139878, and USNM 566465– and USNM 531663 from Iwokrama Forest
566467 from Aishalton; both sexes are Reserve.
represented. Coloration in life.—The following is
Coloration in life.—The following is based on AMNH and USNM specimens
based on AMNH and USNM specimens from Berbice River Camp (CJC field
from Yupukari and Aishalton (CJC field notes, 1997). Brown above (darker on
notes, 1992, 1993). Dorsum light copper sides, but on some, uniform dark brown
(extending onto base of tail); sides, arms, above and on sides) with inconspicuous
dark tan markings; venter translucent male, from Ceiba Biological Station (CJC
gray, but bluish gray beneath tail. An field notes, 1994). Dorsum medium brown
adult male also had an inconspicuous tan with a light tan dorsolateral stripe, below
dorsolateral stripe anteriorly and bright which is dark brown; throat and chest
orange ventral surfaces, including the orange, abdomen orangish yellow (proba-
arms, legs, and tail. bly not so brightly colored in females and
juveniles).
Comments.—The karyotype of this dip-
Kaieteurosaurus hindsi Kok, 2005 loid, bisexual (gonochoristic) species was
described by Pellegrino et al. (1999). This
Type material.—The holotype is IRSNB is one of the cryptic species in Leposoma,
2628, an adult male from Tukeit trail, ca. of several that are extremely difficult to
1250 m NEE from the beginning of the distinguish from each other, and is a
Kaieteur National Park airstrip (from candidate for being one of the ancestors
point closest to the gorge), 420 m eleva- for unisexual forms (see Comments for
tion, Kaieteur National Park, Potaro- Leposoma percarinatum, below).
Siparuni district, Guyana.
type locality. Leposoma percarinatum (Müller, 1923)
Vouchers for Guyana.—The holotype is Plate 22F
the only known specimen.
Coloration in life.—‘‘Dark brown dor- Type material.—The original name was
sally and laterally. Chin whitish, venter Hylosaurus percarinatus and Uzzell &
and underside of limbs uniformly reddish Barry (1971) stated that the holotype, a
orange. Proximal quarter of tail reddish female, was probably destroyed during
orange, remaining distal part brownish World War II. It was from Peixeboi (a.d.
speckled with whitish orange’’ (Kok Bragançabahn), Parà, Brazil.
2005:41). Distribution.—‘‘Northern South Ameri-
Comments.—Practically nothing is ca east of the Andes, in French Guiana,
known about this species. Suriname, Guyana, Venezuela, southeast-
ern Colombia, and Brazil’’ (Avila-Pires
1995:408).
Leposoma guianense Ruibal, 1952
Plate 22G 139883 from forest 6.4 km SE Aishalton;
AMNH R-140960, R-141888–141891, and
Type material.—The holotype is USNM 566512 from Dubulay Ranch;
UMMZ 46770, adult female from Du- AMNH R-152010–152015, R-152017–
noon, Demerara River, Guyana. 152020, and USNM 566513–566518 from
Distribution.—‘‘. . .Guyana, Suriname, Berbice River Camp; AMNH R-152027–
French Guiana, and Brazil’’ (Avila-Pires 152038 and USNM 566519–566524 from
1995:394). Konawaruk Camp; also UMMZ 58240
Vouchers for Guyana.—The holotype; from Kartabo (thanks to information
AMNH R-140962 from Ceiba Biological provided by Greg Schneider); USNM
Station, Madewini River, ca. 5 km E 531665–531673 from Iwokrama Forest
T i m e h r i A i r p o r t ( 0 6 82 9 0 5 7 00 N , Reserve (various sites); USNM 535792
058813 0 06 00 W); and UMMZ 46768 from from Baramita; and USNM 566511 from
the type locality (Ruibal 1952). Ceiba Biological Station, Madewini River,
Coloration in life.—The following is ca. 5 km E Timehri Airport (06829 0 57 00 N,
based on the AMNH voucher, an adult 058813 0 06 00 W).
Coloration in life.—The following is R-61365–61366 from Shudikarwau;

based on AMNH and USNM specimens AMNH R-151983–151985 from Berbice
from Aishalton, Ceiba Biological Station, River Camp; and USNM 497802 from
Dubulay Ranch, Berbice River Camp, and Moco Moco at base of the Kanuku
Konawaruk Camp (CJC field notes, 1993– Mountains.
1998). Brown above (all dorsal surfaces), Coloration in life.—The following is
with inconspicuous yellowish tan dorsolat- based on the AMNH specimens from the
eral light stripe; some specimens with Berbice River Camp (CJC field notes,
inconspicuous light tan to yellow spots on 1997). Brown above (including arms, legs,
sides; chin translucent white; throat translu- and tail), with network of darker brown
cent very pale gray to pale yellow or orange; markings; dirty white to tan or pale yellow
chest, abdomen, beneath arms, legs, and spots on sides; head brown; upper eyelid
base of tail translucent pale yellow or gray. with thin edge of copper; lower eyelid with
Comments.—This is an all-female (uni- thin edge of white, extending upward
sexual) species, which probably includes at posterior to eye; ventral surfaces very pale
least two cryptic species, one of which tan (but chin and throat may be white),
remains un-named. See Avila-Pires with gray spots on chin and gray streaks
(1995:411), Pellegrino et al. (2003, 2011), on throat; one with yellow patch ventro-
and Laguna et al. (2010) for discussions of laterally on neck in front of arm; on some,
the taxonomic difficulties involving several most of distal tail black below (but tan
cryptic species of Leposoma that occur in
with gray or brown dots beneath base, as
Amazonia and that may be or may have
beneath legs).
been undergoing hybridization. The cryptic
species include two unisexual forms, one
diploid and one triploid (Laguna et al.
2010, Pellegrino et al. 2011). Specimens of Neusticurus rudis Boulenger, 1900
this complex are extremely difficult to Plate 23C
identify with confidence based on morphol-
ogy alone. Extensive molecular research on
1946.8.31.64 according to Avila-Pires
the entire complex has recently been
initiated by Pellegrino et al. (2011). (1995), from the foot of Mt. Roraima,
3500 ft, reported originally as being from
Guyana. However, the type specimen was
probably collected in Venezuela (Phelps
Neusticurus bicarinatus (Linnaeus, 1758) 1938). See below (Vouchers) for records
Plate 23B from Guyana.
Type material.—The original name was Distribution.—‘‘Guiana region, in Ven-
Lacerta bicarinata and Avila-Pires (1995) ezuela (Bolı́var), Guyana, Suriname,
stated that the holotype is UUZM 70. French Guiana, and Brazil’’ (Avila-Pires
Hoogmoed (1973) restricted the type lo- 1995:446).
cality to the vicinity of Paramaribo, Vouchers for Guyana.—Possibly the
Surinam. holotype; AMNH R-29927 from Kartabo;
Distribution.—‘‘. . .eastern Venezuela, AMNH R-151986–151998, R-152002–
Guyana, Suriname, French Guiana, and 152004, and USNM 566525–566526 from
Brazil’’ (Avila-Pires 1995:426). Konawaruk Camp; ROM 39497–39500
Vouchers for Guyana.—AMNH R-6138 from Mount Ayanganna; ROM 20514
from Marudi; AMNH R-21259 from and 20517 from Tukeit; and USNM
Kartabo; AMNH R-61288–61290 from 549324–549325 from the north slope of
Onora; AMNH R-61361, R-61363, and Mt. Roraima (5817 0 N, 060845 0 W; 800 m).
Coloration in life.—The following is black. Iris red. . .Tongue dark gray in its
based on AMNH and USNM specimens anterior two-thirds, whitish with gray
from Konawaruk Camp (CJC field notes, flecks on its posterior third. Palms and
1998). Dorsum dark brown, with dark soles black. Underside of head and throat
yellowish tan squares and rectangles; head, black with irregular light gray markings;
arms, and legs dark brown (arms and legs venter orangish brown, ventral scales
with few yellowish tan spots); yellowish peppered with black; underside of upper
tan spot beneath eye; eyelids may be edged legs [arms] black; underside of lower legs
with yellow; tail, above, as dorsum; chin black with irregular orangish brown mark-
and throat orange, but throat may be tan ings; underside of tail black, except prox-
or pale yellow; chest and beneath arms and imally where some scales are white’’ (Kok
legs yellow or orangish yellow; abdomen 2009b:65).
gray or grayish tan; very pale orangish tan Comments.—Essentially nothing is
beneath tail base, with numerous brown known about this species.
flecks, tail gradually becoming black pos-
teriorly; on some, ventrolateral tail with
row of tiny white dots.
Comments.—The vouchers from Guya- Riolama leucosticta (Boulenger, 1900)
na are of the morph that Avila-Pires (1995) Type material.—The original name was
discussed as having the tubercles scattered Prionodactylus leucostictus and Colin Mc-
on the body (not in regular longitudinal Carthy informed us (pers. comm.) that the
rows) and having roundish and convex holotype was first cataloged as BMNH
(not trihedral) gular scales. However, the 99.3.25.4 and later, after World War II,
Konawaruk lizards have significantly more
‘‘re-registered with the number BMNH
femoral and preanal pores (both sexes)
1946.8.2.8.’’ The specimen was noted as
than reported by Avila-Pires, which could
being from the summit of Mt. Roraima,
represent geographic variation or the
8600 ft, Guyana, for which Colin Mc-
presence of cryptic species yet to be
Carthy also informed us that ‘‘the original
diagnosed (see Remarks in Avila-Pires
[catalog] entry (in Boulenger’s handwrit-
1995).
ing) states ‘Mt. Roraima, Brit. Guiana
8600 f (summit).’’’ However, the type
specimen was probably collected on the
Pantepuisaurus rodriguesi Kok, 2009b Venezuelan part of the summit (Phelps
Type material.—The holotype is IRSNB 1938).
2650, an adult male from the summit Distribution.—This species is known
plateau of Mount Maringma, Cuyuni- only from the summit of Mount Roraima
Mazaruni District, Guyana (05812 0 57 00 N, and one or two additional tepuis in
060835 0 07 00 W; 2080 m). Venezuela (Gorzula & Se ñaris 1999
Distribution.—Known only from the [1998], MacCulloch et al. 2007). We do
type locality. not doubt that this species occurs in both
Vouchers for Guyana.—The holotype is Venezuela and Guyana on Mount Rorai-
the only specimen known. ma, as the habitats are the same within the
Coloration in life.—‘‘Dorsal and lateral summit areas of both countries. We
surfaces completely black; closer examina- emphasize, however, that voucher speci-
tion shows that black scales on body and mens having localities clearly within Guya-
neck are peppered with brown or reddish na are needed.
brown. Upper surface of head black. Vouchers for Guyana.—See above (Type
Tympanum light pink. Arms legs and tail Material and Distribution).
Coloration in life.—The following is Corytophanidae). Representatives of this

f r o m G o r z u l a & S e ña r i s ( 1 9 9 9 family occur in ‘‘tropical and subtropical
[1998]:142), based on specimens from America from the southwestern United
Cerro Kukenán, Venezuela. ‘‘In life their States and eastern Mexico south to south-
ground color was shiny black with yellow- ern Brazil and Paraguay; Galapagos Is-
ochre speckling dorsally and dorsolateral- lands; Antilles; Fiji and Tonga Islands’’
ly. The ventrals, anals and subcaudals had (Frost & Etheridge 1989:39). Although the
yellowish white speckles. Pale yellow man- occurrence of some iguanids in the Gala-
dibular stripes formed an inverted ‘V.’’’ pagos Islands undoubtedly resulted from
Comments.—See above (Distribution) over-water rafting, their occurrence in Fiji
for comments on the need to clearly and the Tonga Islands may reflect their
document the existence of this species in Gondwanan origins and ancient terrestrial
Guyana. As for most of the mountaintop connections (Noonan & Sites 2010).
fauna of Mount Roraima, practically
nothing is known about the biology of
this species. Iguana iguana (Linnaeus, 1758)
Tretioscincus agilis (Ruthven, 1916) Lacerta Iguana and there are two syntypes,
Plate 23A NRM 114 and UUZM Linnean collection
Type material.—The original name was no. 10 (Avila-Pires 1995). Hoogmoed
Calliscincopus agilis and the holotype is (1973) restricted the type locality to the
UMMZ 47798 from a sand ridge near confluence of the Cottica River and Perica
Dunoon, Guyana. Creek, Surinam.
Distribution.—‘‘Guyana, Suriname, Distribution.—‘‘From Mexico, though
French Guiana, and Brazil’’ (Avila-Pires Central America and part of the Antilles,
1995:482). to central Brazil and Paraguay’’ (Avila-
Pires 1995:47).
Vouchers for Guyana.—The holotype is
the only specimen known from Guyana.
14111–14114, R-21289, and R-21330 from
Kartabo; also CJC and CRT saw an
from Avila-Pires (1995:482). ‘‘Brown on
individual (not collected) at Aishalton in
back, with dorsolateral stripe anteriorly
1993.
salmon to pale orange, posteriorly yellow-
ish or greenish, and blue on tail; [dorso-
based on specimens from Suriname
lateral stripe]. . .may be bordered dorsally
(Hoogmoed 1973:155–156). ‘‘Grass-green
by a black stripe. Flanks black. Ventral
all over, slightly lighter on the ventral
region pearl-white or with a greenish shine,
parts. In old specimens the overall colour
with centre of most scales, especially
fades to a bluish grey. A distinct white line
posteriorly, black-peppered. Tail mostly
on the lower anterior surface of the upper
deep pastel blue, proximally with dorsal
arm, from the insertion to near the elbow.
and lateral black stripes. Iris black.’’
The upper surface of the head in adults
Hoogmoed (1973) stated that the ventral
may be bluish green or grayish. A number
area may be greenish blue.
of more or less distinct vertical white
stripes, accompanied by a black zone is
Iguanidae present on the lower part of the flanks.
Gular appendage with black and reddish
This is one of the families of iguanian brown stripes. Infra- and sublabials and
lizards (see Introduction above, before anterior margin of gular appendage with
reddish brown spots. Tail with white and Vouchers for Guyana.—AMNH R-8087,
black transverse bands on a grass-green R-60063–60066, R-60068–60070, R-
background. In old specimens the tail 60072–60074, R-60076–60086, R-60088–
shows black bands, alternating with bluish 60090, and R-60333–60337 from George-
grey bands. Iris brown with a golden rim town, identified by E. E. Williams. Also
around the pupil.’’ AMNH R-98386–98391 from near Karta-
Comments.—This is the largest species bo, identified by E. E. Williams. Also
of lizard in the Western Hemisphere. AMNH R-84156–84157 from Rockstone
Savage (2002) mentioned an individual Landing and AMNH R-38573 from De-
over 2 m in total length (body and tail), merara River, Akyma, identified by E. E.
and weight of 4 kg. Avila-Pires (1995) Williams. Also AMNH R-139786, ROM
mentioned a preserved specimen from 20571, and USNM 497793 and 566365
Brazil with a total length of about 1.7 m. from Georgetown; and USNM 163000–
The species is primarily arboreal. 163015 from Enmore Estate (6844 0 N,
057859 0 W).
based on two AMNH and USNM speci-
Polychrotidae mens from Georgetown (CJC field notes,
This is one of the families of iguanian 1993). Greenish yellow above and below,
lizards (see introduction above, before including dewlap on large one, but dewlap
Corytophanidae). The family occurs in with yellow scales and orange skin in
between scales on smaller one.
‘‘southern North America to southern
Comments.—This species is abundant in
South America; West Indies’’ (Frost &
Georgetown and Kartabo, but it is not
Etheridge 1989:43). Representatives of two
part of the original, native herpetofauna of
genera are known to occur in Guyana—
Guyana (Murphy 1996, Avila-Pires 2005).
Anolis and Polychrus. Anolis is a very large
The species is a member of the roquet
genus, which some recent authors have
species group, which has a natural distri-
tried to partition into several smaller bution on islands of the West Indies.
genera. However, the most recent phylo- Historically, three names are relevant to
genetic analyses have shown that the the taxon in Guyana: Anolis roquet
smaller genera were largely not monophy- (Lacépède 1788); Anolis aeneus Gray,
letic groups and that retention of the single 1840 (once treated as a variety of roquet);
large genus concept is most consistent with and Anolis extremus Garman, 1887 (orig-
evidence-based phylogenetic conclusions inally named as a variety of roquet). The
(Poe 2004). Townsend et al. (2011) split specific names applied in Guyana have
Anolis out to a new family, Dactyloidae. been both A. aeneus and A. extremus, if
not using names of subspecies, A. roquet if
using subspecies. All three forms are very
Anolis aeneus Gray, 1840 similar to each other, particularly aeneus
and extremus.
Type material.—The holotype is BMNH Peters & Donoso-Barros (1970) stated
1946.8.28.7 and the type locality was that both A. aeneus and A. roquet extremus
restricted to Point Saline, Grenada (Lazell occur in Guyana, citing Underwood (1964)
1972). but not any individual specimens. They
Distribution.—Petit Martinique, Grena- also gave key characters for distinguishing
dines, Trinidad and Guyana (Peters & among these species, but the characters
Donoso-Barros 1970); also, see Comments were wrong. Murphy (1997) said that both
(below). aeneus and extremus occur in Trinidad,
using a new character of the rostral scales RPR showed clearer, but not sharp,
to distinguish them, but this character also differences in axillary coloration.
fails to work on AMNH specimens that AMNH has several series of roquet
CJC examined. The roquet group was last group anoles from Guyana, mostly old
reviewed by Lazell (1972:80), who present- ones. All are identified as A. aeneus, and in
ed little morphological information and each case where the determiner is recorded,
stated about coloration of aeneus that it was Ernest E. Williams. These are the
‘‘there is so much variation that verbal majority of those cited among the vouch-
description is wasteful.’’ He also said ‘‘I ers above. Clearly, the taxonomic status of
have not seen Guyana...specimens alive, Anolis aeneus versus Anolis extremus needs
but MCZ 81283–92 (Georgetown) are further investigation, both in areas where
precisely this anole.’’ He gave no rationale they occur in the West Indies and where
for this determination, nor did he justify they have been introduced on islands and
treating these forms as separate species. the mainland of South America.
However, in a key Lazell (1972:23) said
that aeneus has the ‘‘axilla without dark
pigment,’’ extremus has the ‘‘axilla with Anolis auratus Daudin, 1802b
dark pigment.’’ Plate 23D
For A. aeneus, Lazell (1972:81) stated
that the main range is Grenada and the Type material.—The type material, from
Grenadines, ‘‘...introduced into Trinidad Surinam, is unknown (Avila-Pires 1995).
and Guyana.’’ For extremus, the range was Distribution.—‘‘. . .Brazil, French Gui-
given as Barbados and introduced into ana, Suriname, Guyana, Venezuela, and
other islands and Caracas, Venezuela (no Colombia, northward to Panama’’ (Avila-
mention of Guyana). The diagnostic char- Pires 1995:52).
acters do not work, given variation ob- Vouchers for Guyana.—AMNH R-
served on AMNH specimens that CJC 137943–137948, R-137953, R-137959–
examined, including no distinct difference 137970, R-138063, R-138087, and USNM
in axillary coloration, despite Lazell’s 566337–5663345 from Karanambo and
(1972:85) following statement: ‘‘The axil- Yupukari; AMNH R-139763–139775, R-
lary pigmentation makes a simple distinc- 139783–139784, and USNM 566346–
tion between extremus and aeneus, 566352 from Aishalton. Also USNM
especially when specimens (even preserved) 163006–163007 from Enmore Estate,
are compared directly.’’ Our comparisons 6844 0 N, 057859 0 W; USNM 291131 from
involved specimens from Barbados (ex- Kato; and USNM 291185 from Para-
tremus; AMNH R-14096–14101 and R- makatoi. In addition, ‘‘RMNH 26479’’
15105–15109), Grenada (aeneus; AMNH from ‘‘Berbice River, near Blairmont’’
R-104731–104734), and Guyana (see (Avila-Pires 1995:49), and Mike Tamessar
Guyana Vouchers, above), and there was (pers. comm.) informed us that he has seen
no clear resolution between aeneus and this species at Dubulay Ranch.
extremus using any of the characters Coloration in life.—The following is
mentioned by Peters & Donoso-Barros, based on AMNH and USNM specimens
Murphy, or Lazell. In general the armpit from Karanambo, Yupukari, and Aishal-
character looked a bit darker on most ton (CJC field notes, 1992, 1993). All dorsal
specimens from Barbados vs. Grenada, surfaces tan; cream or dark brown to black
but it was subtle and relative, and not lateral stripe; if stripe cream, lined above
something that was applied easily on and below by black flecks, but the lower
specimens from Guyana. More recently edge may be a black stripe; some lizards
collected USNM specimens examined by also with a ventrolateral black stripe; rust-
orange spots on posterior of thigh; tail tan tail; dewlap of males light gray anteriorly,
above with light gray bands, smudges, or bright pink posteriorly.
indistinct, short black stripes; ventral sur-
faces cream, but orangish tan under tail;
small dewlap of females with grayish blue Anolis ortonii Cope, 1868
skin between cream scales; larger dewlap of
males similar but with yellow scales in Type material.—The holotype is ANSP
center and at end beside base; tongue 11404 (Avila-Pires 1995) from Napo or
orange, brightly displayed when captured Upper Rio Marañon, Ecuador or Peru.
lizard gapes. On an adult male the lateral Distribution.—‘‘Northern South Ameri-
bright cream stripe was observed to change ca east of the Andes, in Brazil, French
to black following capture. Guiana, Suriname, Guyana, Colombia,
Comments.—These little anoles often Ecuador, Peru, and Bolivia’’ (Avila-Pires
sleep at night on tall grass on the 1995:94).
Rupununi Savanna. Vouchers for Guyana.—AMNH R-
21292 from Kartabo; and AMNH R-
25074 and R-25141 from Kamakusa,
Anolis fuscoauratus identified by E. E. Williams.
Duméril & Bibron, 1837 Coloration in life.—The following is
Plate 23F based on a female from Suriname
Type material.—The original name was (Hoogmoed 1973:140). ‘‘Body yellowish
Anolis fuscoauratus and the holotype is grey, black to dark-brown on the vertebral
MNHN 2420 (Avila-Pires 1995). The type region, a brown line between the eyes,
locality is Rio Mamoré, between Loreto upper surface of the head with a tinge of
and the confluence of Rio Sara, Moxos pink, flanks reticulated with dark-brown,
province, Bolivia (Avila-Pires 1995). tail grey with brownish transverse bands,
Distribution.—‘‘Northern South Ameri- belly grey, dewlap yellow-ochre with a few
ca east of the Andes, in Brazil, French horizontal rows of white scales. Iris
Guiana, Suriname, Guyana, Venezuela, golden.’’ Avila-Pires (1995:93) added that
Colombia, Ecuador, Peru and Bolivia’’ ‘‘no distinction in colour between sexes
(Avila-Pires 1995:64). was observed.’’
Vouchers for Guyana.—AMNH R-21306
and R-21324 from Kartabo; AMNH R-
151828–151841 and USNM 566358–566363 Anolis planiceps Troschel, 1848
from Konawaruk Camp; USNM 291267 Plate 23E
from Kaieteur, near guest house; and
USNM 531642–531644 from Iwokrama Type material.—The holotype is ZMB
Forest Reserve (various localities). 529 from Caracas, Venezuela.
Coloration in life.—The following is Distribution.—This species occurs in
based on AMNH and USNM specimens much of the area of the Guiana Shield in
from Konawaruk Camp (CJC field notes, northern Brazil, Guyana, and Venezuela,
1998). Dark tan dorsum with subtle dark as well as Trinidad (D’Angiolella et al.
brown spots along vertebral area; tail 2011).
similar, but with subtle dark brown bands Vouchers for Guyana.—Vanzolini &
instead of spots, and tail may become Williams (1970:21) cited a long list of
lighter distally; venter white to gray with vouchers from Guyana, using the name
extensive tan flecking; some males have Anolis chrysolepis, in the following collec-
very pale yellow iridescent ventrolateral tions: AMNH, BMNH, MCZ, and
wash on body and yellow beneath base of UMMZ. Specimens that we examined are
the following: AMNH R-15137–15142, R- these species might be the size of the scales
21307–21308, R-21310–21314, R-32273– on the upper arm; if larger than the scales
32278 from Kartabo; AMNH R-151810– in the vertebral area, A. planiceps; if the
151818, and USNM 566353–566356 from same size or smaller, A. chrysolepis.
Berbice River Camp; AMNH R-151823–
151825 and USNM 566357 from Kona-
waruk Camp; ROM 22853–22868 and Anolis punctatus Daudin, 1802b
USNM 535793–535794 from Baramita;
Plate 24A
USNM 200508 from Bartica; USNM
233871 from Kartabo Point (6823 0 N, Type material.—The holotype is
058842 0 W); USNM 497792 from CEIBA MNHN 2340, a female from ‘‘South
(6829 0 55 00 N, 058813 0 04 00 W); and USNM America’’ (Avila-Pires 1995:102).
549322 from the north slope of Mount Distribution.—‘‘Widespread in Amazo-
Roraima. nia; present probably throughout French
Coloration in life.—The following is Guiana and Suriname, and in parts of
based on AMNH and USNM specimens Guyana, ...Colombia, Ecuador, Peru, Bo-
from Berbice River Camp and Konawaruk livia (Fugler 1988), and Brazil’’ (Avila-
Camp (CJC field notes, 1997, 1998). The Pires 1995:109).
animal is covered with a pattern of browns Vouchers for Guyana.—AMNH R-
and tans; with a fine vertebral light rust 18185 and R-21293 from Kartabo;
stripe, conspicuous in sacral region; on AMNH R-61237 from Kuyuwini Landing;
some, a wide tan vertebral stripe with dark
AMNH R-61249 from Onora; AMNH R-
brown edges; head brown, with dark
98379–98380 from near Kartabo; and
brown patches, fine irregular rust-orange
USNM 566364 from Berbice River Camp.
lines; some with pale yellow patches
beneath eye; tail dark brown; venter white
based the USNM specimen from Berbice
or yellowish tan, some with black specks;
throat may be brown; dewlap orange or River Camp (CJC field notes, 1997). Bright
red, but may be purple at base. green (can change at least in large part to
Comments.—Avila-Pires (1995) treated brown), with small pale blue spots on
this taxon as A. nitens, with priority over sides; chin, throat, and chest light green;
A. chrysolepis. However, in preparing our belly and beneath base of tail pale bluish
manuscript we followed Myers & Donnelly gray, with tiny cream dots (most of tail
(1997) in accepting the name Anolis regenerated). Hoogmoed (1973:147) added
chrysolepis to represent a wide-ranging ‘‘Dewlap in males yellow-ochre with rows
species with several subspecies. Most of white scales, in females the scales on the
recently, D’Angiolella et al. (2011) con- small dewlap are yellow. Iris orange-
cluded primarily from molecular analyses brown.’’
that each of the subspecies should be
elevated to species status. Their interpre-
tation is that two species (formerly sub- Polychrus marmoratus (Linnaeus, 1758)
species) of this complex occur in Guyana, Plate 24B
A. planiceps and A. chrysolepis. The former
occurs widely in the northern and central Type material.—The original name was
part of Guyana, the latter is represented by Lacerta marmorata and the syntypes are
a few specimens from the south (D’Angio- UUZM Linnean collection nos. 14, 31
lella et al. 2011). All the specimens we (Avila-Pires 1995). Hoogmoed (1973) re-
examined are A. planiceps. The best stricted the type locality to the vicinity of
morphological character for distinguishing Paramaribo, Surinam.
Distribution.—‘‘Northern South Ameri- Mabuya nigropunctata (Spix, 1825)

ca east of the Andes (French Guiana, Plate 24D
Suriname, Guyana, Venezuela, Colombia,
Ecuador, Peru, and Amazonian Brazil), Type material.—The original name was
and in the Atlantic forest [of Brazil]’’ Scincus Nigropunctatus and the holotype
(Avila-Pires 1995:142). was lost. Avila-Pires (1995) designated
MPEG 15248 as neotype. It is from Santa
Rita, Municı́pio de Maraã, left margin of
21315, R-32279–32281, and R-137404–
Rio Japurá (Lago Paricá), Amazonas,
137409 from Kartabo; AMNH R-139787
Brasil.
from Aishalton; and AMNH R-141901
Distribution.—‘‘Throughout Brazilian
from Dubulay Ranch; USNM 84487–
Amazonia and most probably in the entire
84489 from Pomeroon; USNM 163053
Amazonian region. Outside Amazonia, it
from Enmore Estate (6844 0 N, 057859 0 W);
seems to occur in the state of Mato
and USNM 566374 from Yupukari.
Grosso..., and in part of the Atlantic
Coloration in life.—The following is Forest’’ (Avila-Pires 1995:590). Because
based on AMNH and USNM specimens of the difficulties of distinguishing among
from Karanambo or Yupukari, Aishalton, the similar species in this complex and the
and Dubulay Ranch (CJC field notes, lack of a detailed analysis of geographic
1992–1995). Dorsum (body, arms, legs, variation, the limits of the range of
tail) brown with dark brown bands and distribution remain unknown. Neverthe-
many scales green (the lizard becoming less, all specimens of Mabuya that we have
greener with thin black bands on body examined from Guyana represent this
while being handled); top of head gray, species.
with a few dark brown spots, short dark Vouchers for Guyana.—AMNH R-
brown or black lines from eyes, and pale 6785–6786 from Kaieteur; AMNH R-
green wash; undersides all pale green, but 8543 and R-37550–37552 from George-
darker on posterior abdomen, preanal town; AMNH R-15120–15122, R-18183,
area, hind legs, and base of tail; when and R-21326 from Kartabo; AMNH R-
excited and bright in color, orange patches 25070 and R-25128 from Kamakusa;
on posterior of thigh and in axilla. AMNH R-139814 and USNM 566395
from Aishalton; AMNH R-151880–
151883, R-151888–151891, and USNM
Scincidae 566396–566399 from Berbice River Camp;
AMNH R-151892 from Konawaruk
The Scincidae is a very large family. It Camp; ROM 22887–22891 and USNM
has representatives basically all over the 535801 from Baramita; ROM 20516 from
world wherever any kind of reptiles exist. Kurupukari; ROM 28349 from Parama-
Nevertheless, only one species is known to katoi; USNM 164190 from Mabaruma
occur in Guyana. This is a member of the (8812 0 N, 059847 0 W); and USNM 531658–
genus Mabuya, in which many of the 531662 from Iwokrama Forest Reserve
species are very similar to each other and (various localities).
difficult to identify. Recent research based Coloration in life.—The following is
on morphology and phylogenetic analyses based on AMNH and USNM specimens
including DNA sequence data are begin- from Aishalton, Berbice River Camp, and
ning to improve our understanding of Konawaruk Camp (CJC field notes, 1993,
these lizards (e.g., Avila-Pires 1995, Maus- 1997, 1998). Dorsum, including top of
feld et al. 2002, Miralles et al. 2009), but a head and tail, copper or bronze, often with
lot of work remains to be done. dark brown spots; pale coppery tan
dorsolateral light stripe may be present; from Marudi; USNM 570538 from Gunn’s
arms, legs, side of face, and side of body Strip (01836 0 46 00 N, 058838 0 15 00 W).
dark brown; often a ventrolateral yellow- Coloration in life.—The following is
ish tan light stripe, with dark brown below; based on specimens from Brazil (Avila-Pires
throat very pale yellow to yellowish green; 1995:248). Dorsum reddish brown to brown;
beneath arms, chest, abdomen, legs, and light spots on head and body cream, beige,
base of tail yellow to yellowish green, yellow, or pale brown; on some, beige band
brightest posteriorly; underside of hands, on posterior head; posterior thigh and base
feet, fingers, and toes black; distal tail of tail with short, light, beige to orange
yellowish copper. stripe; underside of head and belly white, but
Comments.—See above (Distribution) posterior belly pale pink or orange; under-
for comments on the need for research side of tail (and legs of some) pink to beige;
on the overall distribution of this and ‘‘iris dark grayish-brown with an orange rim
related species in South America. around pupil. Tongue grey anteriorly, white
posteriorly.’’
Sphaerodactylidae Comments.—This species is similar to the
following (C. septentrionalis), so the small
Until recently, all Guyanan genera of specimens need to be identified with extra
geckoes were included in the large family care (see key for characters). No compara-
Gekkonidae, as mentioned above (intro- tive studies have been done on the life
duction to Gekkonidae). However, recent histories and ecology of these two species.
phylogenetic analyses (Gamble et al. This species formerly was in the genus
2008), including considerable DNA se- Coleodactylus, but Gamble et al. (2011)
quence data, indicated that Guyanan named Chatogekko for this species, follow-
geckoes of the genera Chatogekko, Coleo- ing morphological and DNA sequence
dactylus, Gonatodes, Pseudogonatodes, and analyses, because it is in a very distinctive
Sphaerodactylus are appropriately consid- lineage. They also noted that there probably
ered as representatives of another family, are several cryptic species masquerading
Sphaerodactylidae. Individuals of most under the name C. amazonicus.
species of sphaerodactylid lizards are very
small, including those that are among the
smallest in Guyana. Coleodactylus septentrionalis
Vanzolini, 1980
Plate 24C
Chatogekko amazonicus (Andersson, 1918)
Type material.—The original name was Type material.—The holotype is
Sphaerodactylus amazonicus and the holo- MZUSP 52866, a male from Brasil,
type from Amazonas, Manaos, Lago Roraima: Ilha de Maracá.
Poraquecuare, Brazil is NRM 3254 (Avi- Distribution.—‘‘Northern part of Rorai-
la-Pires 1995). ma, Brazil; western Suriname. . .; Pacarai-
Distribution.—‘‘Eastern and central ma foot hills, Guyana; Delta Amacuro,
Amazonia, including Brazil, French Gui- Venezuela’’ (Avila-Pires 1995:257).
ana, Suriname, southern Guyana, and Vouchers for Guyana.—AMNH R-
Southern Venezuela (Amazonas)’’ (Avila- 132006 from Isheartun; AMNH R-
Pires 1995:250), south to northern Bolivia 137976-137977 and USNM 566300 from
(Gamble et al. 2011). Karanambo; USNM 531620 from Iwo-
Vouchers for Guyana.—AMNH R- krama Forest Reserve, Burro Burro River;
144757–144759, R-144761, and R-144763 and AMNH R-148592–148594 and
USNM 531621–531622 from Iwokrama GUYANA: Magdalen’s Creek Camp, near

Forest Reserve (several localities). (ca. 275 m N) the NW bank of the
Coloration in life.—The following is Konawaruk River (ca. 40 km [linear]
based on AMNH and USNM specimens WSW Mabura Hill), 120 m elevation
from Karanambo (CJC field notes, 1992). To (5813 0 07 00 N, 59802 0 43 00 W) in WGS 84.
the naked eye, this little lizard (maximum Distribution.—Known only from Guya-
body length about 30 mm or less) may simply na and eastern Venezuela (Schargel et al.
appear to be brown to dark brown above, 2010).
with an inconspicuous light tan stripe dorso- Vouchers for Guyana.—AMNH R-2714
laterally in hip area and on base of tail; venter from Kartabo; AMNH R-151804–151809
translucent pinkish gray to gray with brown from the type locality; IRSNB 2630–2639
speckles. Avila-Pires (1995:254) stated that from several sites in Kaieteur National
‘‘the only colour description of living speci- Park (Cole & Kok 2006:3); and UTA
mens available is that of O’Shea (1989:55), 57504 from ‘‘near Imbaimadai, Pacaraima
‘brown with a series of paired white dorsal Mountains, Guyana’’ (Schargel et al.
spots and occasionally a pair of broken white 2010:327).
dorso-lateral stripes.’’’ She also noted that Coloration in life.—‘‘Dorsum dark red-
other color descriptions in the literature were dish brown [gray on some] through base of
from preserved specimens (perhaps aided by tail [grayish brown on some]; rest of tail
microscopy), and she noted that there is gray; dorsum with tiny irregular black
significant variation in color pattern, includ- markings [not seen on some]; orange wash
ing specimens with an apparent ‘‘absence of a on side of face, around neck dorsally
pattern’’ (Avila-Pires 1995:257). Neverthe- [yellowish orange on some], and on anterior
less, Avila-Pires noted that specimens from surface of arms; yellowish orange beneath
Roraima have a light band posteriorly on the arms [or bright yellow, but not seen on
head, and Vanzolini (1980:9) illustrated that some]; venter gray, but [underside of head
on the holotype, which also showed the and] throat bright orange [yellowish orange
paired dorsal white spots. on some], chest bright yellow [orangish
CJC compared four preserved speci- yellow on some]’’ (Cole & Kok 2006:6).
mens from Brazil (AMNH R-131810– Aspects of sexual dimorphism and possible
131813) with five from Guyana (AMNH seasonal variation in brightness of colors
R-132006, R-137976, and R-148592– are also mentioned by Cole & Kok
148594; see Vouchers, above). The light (2006:9). Schargel et al. (2010:327) de-
spots or stripes were subtle to absent in scribed a new color morph with a ‘‘much
Guyanan material, more evident in general darker body coloration. . .and. . .small ocelli
but also variable in appearance in Brazil- on the sides of the neck and a yellow hood.’’
ian material. Comments.—Phylogenetic analyses in-
Comments.—We would not be surprised if cluding DNA sequence data confirm that
there is more than one species of lizard G. alexandermendesi is a distinct species and
masquerading under this name. Also, see that the Venezuelan G. superciliaris may be
Comments above for Chatogekko amazonicus. a junior synonym, but more work is needed
to be certain (Schargel et al. 2010).
Gonatodes alexandermendesi
Cole & Kok, 2006 Gonatodes annularis Boulenger, 1887
Plate 25A, B Plate 24E, F
Type material.—The holotype is Type material.—The syntypes from
AMNH R-151805 an adult male from Maccasseema (on the Pomeroon River),
Guyana are BMNH 1946.8.22.97–98 (Avi- Gonatodes humeralis (Guichenot, 1855)

la-Pires 1995:258). Plate 25C, D
Distribution.—‘‘Rivero-Blanco (1979)
reported the species for the wet tropical Type material.—The original name was
forests of eastern Venezuela (south of the Gymnodactylus humeralis and the six syn-
Orinoco), Guyana, Suriname, French types are MNHN 6745 (Avila-Pires 1995)
Guyana, and Amapá, Brazil. Here speci- from the edge of the Ucayali River,
Mission de Sarayacu, Peru.
mens are reported from the southern bank
Distribution.—‘‘Widespread in Amazo-
of Rio Trombetas, near Cruz Alta, close to
nia and lowland surroundings, in Brazil,
Rio Amazonas’’ (Avila-Pires 1995:263).
French Guiana, Suriname, Guyana, Ven-
ezuela, Colombia, Ecuador, Peru, and
21251, holotype of Gonatodes beebei Noble
Bolivia. In the Guianas, including the
(1923; a junior synonym), R-2714, and R-
eastern part of Venezuela, it reaches the
coast, as well as the islands of Trinidad
from Georgetown; AMNH R-151801–
and Tobago (Rivero-Blanco 1979)’’ (Avila-
151802 and USNM 566301 from Berbice
Pires 1995:281).
River Camp; ROM 22921–22964 and Vouchers for Guyana.—AMNH R-
USNM 535784–535791 from Baramita; 18193–18194, R-29922–29924, and R-
ROM 20518–20520 from Kurupukari; 137411 from Kartabo; AMNH R-61421
and USNM 164182–164183 from Maba- and R-131999–132001 from Isheartun;
ruma Compound (08812 0 N, 059847 0 W). AMNH R-139740 from Aishalton;
Coloration in life.—The following is AMNH R-140952–140953, R-140955,
based on AMNH and USNM specimens and USNM 566302 from Dubulay Ranch;
from Berbice River Camp (CJC field notes, AMNH R-151772–151783 and USNM
1997). On one individual, dorsum was 566303–566309 from Berbice River Camp;
reticulation of dark and medium brown, AMNH R-151791–151797 and USNM
with a few inconspicuous, irregular, light 566310–566312 from Konawaruk Camp;
spots of yellow (anteriorly) or gray (pos- ROM 43203–43204 from Kaieteur; and
teriorly), their color transition being com- USNM 531623–531639 from Iwokrama
pleted around the hips; arms and legs as Forest Reserve (various localities).
body; iris dark brown with very thin inner Coloration in life.—The following is
ring of copper; ventral surfaces pale based on AMNH and USNM specimens
chestnut brown; chin, throat, and chest from Aishalton, Dubulay Ranch, Berbice
lighter, with small streaks and spots of River Camp, and Konawaruk Camp (CJC
very pale blue. Two other specimens field notes, 1993–1998). Dorsum brown
(possibly males, indicating sexual dimor- with tan or lighter brown markings that
phism), had dorsal surfaces black with become lighter posteriorly, including out
gold speckling, but hip and base of tail to tip of tail; arms and legs similar; yellow
black with tan mottling and lower part of collar anterior to arms; black dot anterior
legs mostly dark brown (but tail of one to collar; greenish yellow half-moon cres-
animal brown with black reticulation); all cent across back of head, eye to eye; pale
ventral surfaces brownish orange, with, on greenish yellow light spots on head; also
throat only, gold streaks and spots (but metallic dark reddish brown spots on
one lizard with orange patch on throat and head; tiny pale yellow dots on side of
chest). Both Hoogmoed (1973) and Avila- neck; light tan spot at hip posterior to leg;
Pires (1995) discuss sexual dichromatism undersides of body tan, but chest and
and other color and pattern variants in this abdomen may be light brown, throat may
species. be yellowish tan; chin, throat, underside of
arms, and legs with black reticulation or chin with oblique stripes.’’ The photo-
streaks; beneath tail orange or orangish graphs in Murphy (1997:pls. 84–85) show
tan. Adult males have numerous green a male with bright orange on the chin and
flecks and red spots on the body and hind throat and a female with a tan vertebral
legs; red spots atop head; green color to stripe bordered by dark brown, tan, and
head markings, spots on side of neck, and gray spots.
shoulder bar; arms speckled brown with Comments.—The vouchers listed above
green; red flecks on tail; venter of body compare favorably with specimens from
gray with green flecks; orange all along Trinidad, including AMNH R-64454, R-
underside of tail. Sexual dichromatism is 72892, R-72900, and R-72903. This spe-
discussed in more detail by Hoogmoed cies, the natural distribution of which is
(1973) and Avila-Pires (1995). primarily in the Antilles, probably arrived
in Guyana as a result of human introduc-
tion. However, its continued presence in
Gonatodes vittatus (Wiegmann, 1856) Guyana should be confirmed. There is a
question of whether authorship of the
Type material.—The original name was specific name here is Wiegmann or Lich-
Gymnodactylus vittatus based on material tenstein. We followed Kluge (1993) in
from Laguayra, Puerto-Cabello, Caracas, citing Wiegmann.
Venezuela (Wiegmann 1856:6). The Type
is unknown by us.
Distribution.—According to Avila-Pires Pseudogonatodes guianensis Parker, 1935
(2005:31) this species was introduced onto
continental South America and it occurs in Type material.—The holotype, a preg-
Colombian Guayana and Bolivar, Vene- nant female from the upper Cuyuni River,
zuela. According to Gorzula & Señaris is now BMNH 1946.8.27.8 (Avila-Pires
(1999 [1998]:113), ‘‘the distribution of this 1995), which would be the new number
species is restricted to the dry Caribbean assigned after World War II.
coastal areas of eastern Colombia, Ven- Distribution.—‘‘Widespread in a large
ezuela, Trinidad and Tobago and the part of Amazonia, in Brazil, French
Antilles (Murphy 1997, Rivero-Blanco Guiana, Suriname, Guyana, Colombia,
1979).’’ Gorzula & Señaris discuss evidence E c u a d o r , a n d P e r u ’’ ( A v i la - P i r e s
that this was introduced into El Manteco, 1995:304).
Bolı́var State, Venezuela, but the popula- Vouchers for Guyana.—‘‘BM 1946.8.
tion there has not survived. 27.9 [from the type locality]; AMNH R-
Vouchers for Guyana.—AMNH R- 61433,’’ six specimens from Shudikar-wau
61398 and R-132320–132321 from Rock- (Avila-Pires 1995:299). Also AMNH R-
stone Landing, Essequibo River (68– 0 N, 141886–141887 from Dubulay Ranch;
58835 0 W); UMMZ 82848–50 from George- AMNH R-151760–151765 and USNM
town. 566327–566329 from Berbice River Camp;
Coloration in life.—The following is and AMNH R-151769–151770 and
quoted from Murphy (1997:126). ‘‘Males USNM 566330 from Konawaruk Camp.
brown-grey or cinnamon-brown with Coloration in life.—The following is
bright white middorsal stripe bordered based on AMNH specimens from Dubulay
with black, and extending to tip of tail; Ranch and AMNH and USNM specimens
females with white vertebral stripe [not so from Berbice River Camp, and Konawa-
conspicuous as in males], and mottling on ruk Camp (CJC field notes, 1995–1998).
the sides. Venters of both sexes tend to be Dark brown above, with tan band across
white or tan; males sometimes have a white back of head and tan H across hip or base
of tail (sides of H essentially being body...Females tan above with a pair of

dorsolateral stripes); throat white with yellow, dark-bordered dorsolateral stripes
irregular gray markings; gray beneath extending from eye to tail. Venter lighter in
arms, legs, chest, abdomen, tail. Avila- color, chin may be speckled.’’ A color
Pires (1995:302) described a male from photograph (Murphy 1997:pl. 90) suggests
Brazil ‘‘with a middorsal dark stripe, more that the dorsolateral light stripes can be
evident anteriorly.’’ inconspicuous and their dark borders can
Comments.—While preserving speci- be dark brown or black spots.
mens, CJC and CRT noted that these The following is based on an adult male
lizards naturally float, apparently because and female from Kartabo (Beebe 1944:154).
structure of the surface of their scales Adult male: dorsum buff (brownish yellow)
prevents them from wetting. Upon submer- at times tinged with green; brown spots on
sion in formalin, a complete envelope of air body, becoming darker posteriorly and on
instantly forms around them, floating them base of tail; arms and legs similar but
to the surface of the liquid. This must have lighter; ‘‘top and sides of head and neck jet
survival benefits during times of heavy rains black, variegated with creamy white, chiefly
and sudden floods that might sweep these in the form of two broken lines from the
tiny lizards away in running water. This nostrils back over the eyes, forming an
phenomenon, which has been observed in indefinite ‘Y’ on the neck’’; . . .a solid broad
other lizards also, should be studied further, white line from the eye back to the abrupt
including examination of the skin with a ending of the black nuchal area in front of
scanning electron microscope. the fore limb’’; . . .tail abruptly ochracheous-
orange. . .resulting from many large spots
set close together both above and below;
Sphaerodactylus molei Boettger, 1894 rostral, all labials, lower side of head and
neck, the chin and throat yellow-green,
Type material.—The lectotype is SMF remainder of ventral surface of body
9611 and type locality is Caparo, Trinidad creamy-white; . . .iris ivory white, finely
(Mertens 1967:60). dotted with large, dark brown crescentic
Distribution.—According to Gorzula & marks.’’ Adult female: ‘‘buff with the head
Señaris (1999 [1998]:114) ‘‘this species has and neck rather indefinite gray. Apparently
been recorded from northern Colombia, unmarked except for two light flank lines,
Venezuela, Trinidad, Tobago and Guyana down each side of the dorsal pelvic region.
(Murphy 1997, Peters & Donoso-Barros In the right light, however, all the broad
1970).’’ cephalic lines of the male are visible as
Vouchers for Guyana.—AMNH R- lighter brown, i.e., the two lateral lines and
15136, R-18192, R-21255–21257, R- the central ‘Y.’ The lateral lines extend very
21297–21298, R-32284, R-36249, and R- faintly down the body and, becoming strong
137419–137420 from Kartabo. at the pelvic zone, die abruptly at the caudal
Coloration in life.—The following is change to coarser imbricated scalation,
based on Murphy (1997:131). Adult males which marks the area of regeneration.’’
usually have the top of the head conspicu-
ously marked with ‘‘black [or dark brown]
with bright white [or yellow] longitudinal Teiidae
stripes, with lower parts tan-yellow, with
head pattern ending abruptly at back of This group occurs in tropical to temper-
head; [body with] dark stripes which edge ate regions of the New World from North
lighter dorsolateral stripes which break up America to South America. These are
into spots toward the posterior of the mostly medium-sized lizards that are
nervous, common, and frequently seen ROM 20535–20544 from Amatuk; ROM
running rapidly on the ground in forest 20545–20554 from Kurupukari; ROM
or savanna during warm to hot times of 22896–22900 and USNM 535802–535805
day. One species (Tupinambis teguixin) from Baramita; and ROM 28358–28366
shares with Iguana iguana the title of being and USNM 291186–291187 and 291189–
among the largest lizards in the Western 291190 from Paramakatoi. In addition,
Hemisphere. Two genera (Cnemidophorus USNM 162905 from Atkinson (¼Timehri
and Kentropyx) have all-female (unisexual) Airport); USNM 164191–164194 from
species in the Guianan Region, one of Mabaruma (8812 0 N, 059847 0 W); USNM
which (K. borckiana) occurs in Guyana 257523–257524 and 257529–257531 from
and can be seen on warm sunny days in the Kartabo Point (6823 0 N, 058841 0 W);
Botanical Gardens in Georgetown. USNM 291138 from Kato; and USNM
531674–531683 from Iwokrama Forest
Reserve (various localities).
Ameiva ameiva (Linnaeus, 1758) Coloration in life.—Colors and patterns
Plate 26A, B on these lizards are complex and vary
according to age, sex, season, and locality
Type material.—The original name was (geographic variation). The following is
Lacerta Ameiva and there are three syn- based on AMNH and USNM specimens
types, NRM 120, NRM 124, and UUZM from Karanambo, Aishalton, Dubulay
12 (Avila-Pires 1995). Hoogmoed (1973) Ranch, Berbice River Camp, and Kona-
restricted the type locality to the conflu- waruk Camp (CJC field notes, 1992–1998).
ence of the Cottica River and Perica Adult females have a tan or light brown
Creek, Surinam. dorsum (with green iridescence anteriorly
Distribution.—‘‘Panama and most of if body sighted over from the rear), bright
South America east of the Andes (Brazil, green on some from snout to midbody, if
French Guiana, Suriname, Guyana, Ven- not to rump, often with black mottling;
ezuela, Colombia, Ecuador, Peru, Bolivia, some with bright green on posterior body,
Argentina). Southern limits in southern hind legs, base of tail, with tail becoming
Brazil and northern Argentina’’ (Avila- turquoise blue posteriorly; row of brown
Pires 1995:499). spots (with lighter centers) on each side
Vouchers for Guyana.—AMNH R- beside vertebral area; dark brown dorso-
14118–14119, R-15125–15126, R-15130– lateral stripe with scalloped edges, on some
15132, R-21295–21296, and R-46440 from blending below with brown sides, contain-
Kartabo; AMNH R-25040–25042, R- ing pale cream, yellow, tan or blue (on
25062–25063, R-25092, and R-25100– largest) spots or bars; inconspicuous pale
25102 from Kamakusa; AMNH R- greenish yellow stripe above eye along
137907, R-137910–137914, R-138095, and upper edge of dark brown dorsolateral
USNM 566401–566402 from Karanambo; stripe, fading posterior to arms; lateral
AMNH R-139818–139826 and USNM light stripe from below eye, tan or yellow,
566403–566405 from Aishalton; AMNH often becoming cream or tan posterior to
R-140882–140888, R-140895–140896, R- arms; top of head dark tan, sometimes
141853-141855, and USNM 566406– with dark brown spots; lips cream to tan;
566413 from Dubulay Ranch; AMNH R- arms and legs dark brown or tan with tan
151897–151903 and USNM 566414– or brown spots, stripes, or reticulation
566416 from Berbice River Camp; and (sometimes with few light dots of cream to
AMNH R-151904–151911 and USNM pale tan); tail brown above, with irregular
566417–566420 from Konawaruk Camp. dark brown stripes (or not), spots, possibly
Also ROM 20530–20533 from Tukeit; bands; largest individuals with blue spots
on anterior of thighs and feet; chin and mankondre, on the west bank of the
throat cream or tan; chest light tan to pale Marowijne River.
orange; underside of arms, abdomen, legs, Distribution.—‘‘The range includes
feet, base of tail orange, paler anteriorly, northern Brazil, French Guiana, Suri-
darker posteriorly but underside of tail name, Guyana, Venezuela, and Colombia,
becoming very pale orange or gray poste- extending northward through Central
riorly (turquoise blue on largest), some- America to Guatemala, plus the Republic
times with brown bands. of Trinidad and Tobago and other islands
Adult males are similar to the above but of the southern West Indies’’ (Cole &
differ as follows. Dorsum with conspicu- Dessauer 1993:20).
ous green superimposed on pattern, from Vouchers for Guyana.—AMNH R-
back of head to hind legs; colors of 46441–46447 from Kartabo; AMNH R-
scalloped dark brown dorsolateral stripe 138058–138062, R-138065–138069, R-
and brown sides blend together; all lateral 138079, R-138081, and USNM 566429–
area with pale yellow, green, or blue spots 566437 from Yupukari (on Rupununi
or bars; lateral light stripe from below eye River), 11 km (linear) SSW Karanambo,
is tan or green, extends beyond arms; tail northern Rupununi Savanna; AMNH R-
with irregular dark brown spots, no 138099–138108 from Karasabai Village, 40
stripes; throat with black specks; cream, km (airline) NW Karanambo, northern
pale greenish cream, or tan under arms, Rupununi Savanna; AMNH R-139839–
139855 and USNM 566440–566446 from
chest, anterior abdomen; abdomen on
Aishalton; and AMNH R-140862–140871,
some with blue suffusion; pale yellow or
R-140873–140874, R-140880–140881,
orange under legs; blue spots anteriorly on
USNM 497801, and USNM 566447–
legs, feet, side of tail; orange on posterior
566452 from Dubulay Ranch. Also
thigh, preanal area, undersides of feet,
USNM 162906–162912 from Atkinson
beneath base of tail; posteriorly, underside
(¼Timehri Airport); USNM 257525–
of tail very pale grayish green and orange; 257528 from Kartabo Point (6823 0 N,
largest males with green obscuring the 058841 0 W); USNM 291140–291150 from
pattern over entire body above, legs, tail; Kato; USNM 566438–566439 from Jouri,
no lateral stripe. 32 km (linear) NW Karanambo.
Coloration in life.—Colors and patterns
among these lizards vary according to
Cnemidophorus lemniscatus age, sex, season, and locality (geographic
(Linnaeus, 1758) variation). The following is based on
Plate 26C AMNH and USNM specimens from
Yupukari, Aishalton, and Dubulay
Type material.—The original name was Ranch (CJC field notes, 1992–1995).
Lacerta lemniscata. Maslin & Secoy (1986) Dark brown dorsum with light tan
designated as lectotype ZMUU Linnaean stripes; lower sides dark tan in young
Collection 15A, a male. This was over- males and females, tannish gray in adult
looked by Cole & Dessauer (1993:18), so males, with light tan and cream spots
their later designation of a different (some yellow, green, blue in adult males);
lectotype is invalid (Markezich et al. top of head and arms brown, outer lower
1997:51–54), although the invalid speci- arm with tan outer stripe or row of spots
men is in outstanding condition and the (may fade in adults); legs brown with
lectotype is not. Cole & Dessauer (1993) numerous conspicuous large light pale
restricted the type locality to Suriname: yellow to tan spots; tail brown; ventral
Marowijne: Christiaankondre and Langa- surfaces (including beneath arms) cream,
tan, or greenish yellow on abdomen, USNM 163021–163052 from Enmore

preanal area, hind legs, base of tail Estate (6844 0 N, 057859 0 W). In addition,
(becoming gray near tip). Adult males Gallagher & Dixon (1992:163) listed
much more colorful than young males or specimens from Guyana with locality
females, some with powder blue face (at data from several collections.
times including chin and throat); dark Coloration in life.—The following is
brown dorsal stripes; with bright yellow, based on AMNH and USNM specimens
tan, or green dorsal and lateral wash on from the Botanical Gardens in George-
body, turquoise blue on anterior surface town (CJC field notes, 1992), and para-
and underside of hind legs, groin (per- phrased from Cole et al. (1995:4–5).
haps greenish yellow), posterior abdo- Dorsum brown with green wash anteri-
men, and atop feet; bright bluish green orly; short green middorsal stripe atop
or green on lateral aspect of tail (becom- head and on nape of neck; cream, tan, or
ing turquoise distally); bright turquoise green (anteriorly) light stripes, one orig-
blue on midventral surface of tail (paler inating on temporal region, wavy on
yellowish green in adult females). For neck; dark brown stripe (anteriorly)
comparisons with specimens from Sur- becoming spots (posteriorly) above dor-
iname and Venezuela, see Cole & Des- solateral light stripe, dark brown area
sauer (1993) and Markezich et al. (1997). below; small light spots on dorsal sur-
Comments.—Two closely related unisex- faces of legs; dark brown spots dorsally
ual (all-female) species in the Guianan on brown tail; cream stripes laterally on
Region, one diploid and one triploid, had tail.
a hybrid origin in which C. lemniscatus was Comments.—This is an all-female spe-
one of their ancestors (reviewed by Cole & cies that had an origin resulting from
Dessauer 1993). Consequently, these spe- hybridization between K. calcarata 3 K.
cies are similar to C. lemniscatus. They striata, possibly in Guyana, and it
occur in Venezuela, Brazil, and Suriname, appears to reproduce by parthenogenetic
so it seems possible that either or both cloning (Cole et al. 1995). See Hoog-
might some day be found in Guyana. The moed (1973:292–293) and Gallagher &
diploid is Cnemidophorus cryptus and the Dixon (1992) for a discussion on the use
triploid is Cnemidophorus pseudolemnis- of the name K. borckiana.
catus.
Kentropyx calcarata Spix, 1825

Kentropyx borckiana Peters, 1869a Plate 26E, F
Plate 26D
Type material.—The original name was Kentropyx calcaratus based on material
Centropyx Borckiana and the lectotype is from flumen Itapicurú in provincia Mar-
ZMB 897. The type locality is Guiana anhao, Brazil. The type material is ‘‘pre-
(Peters 1869a). sumed lost’’ (Gallagher & Dixon 1992).
Distribution.—Guyana, Surinam, Distribution.—‘‘Mainly in central and
French Guiana, and Barbados (Gallagher eastern Amazonia (Brazil, Guyana, French
& Dixon 1992:132, Fig. 3). Guiana, and Suriname) and Atlantic For-
Vouchers for Guyana.—AMNH R- est (coastal Brazil)’’ (Avila-Pires 1995:524).
37457–37464 from Georgetown; AMNH Vouchers for Guyana.—AMNH R-8489
R-138111–138115, AMNH R-138648, from near Georgetown; AMNH R-8563
and USNM 566492–566493 from the from Georgetown; AMNH R-25065, R-
Botanical Gardens in Georgetown; and 25068, R-25090, and R-25111–25112 from
Kamakusa; AMNH R-21328, R-137358– the widespread use of the name K.

137359, and R-137362 from Kartabo; calcarata.
AMNH R-57448–57449 and R-61459
from Karanambo; AMNH R-61255–
61256 and R-61294–61295 from Kuyuwini Kentropyx striata (Daudin, 1802c)
Landing; AMNH R-60972–60973 from Plate 27A
Marudi; AMNH R-61292–61293 from
Onora; AMNH R-61378–61379 from Ish- Type material.—The original name was
eartun; AMNH R-140967–140968, R- Lacerta striata and holotype from Surinam
141857, R-141861–141865, and USNM is MNHN 4191 (Gallagher & Dixon 1992).
566494–566496 from Dubulay Ranch; Distribution.—‘‘Northern South Ameri-
AMNH R-151957–151964 and USNM ca, in Brazil. . .Suriname, Guyana, Vene-
566497–566501 from Berbice River Camp; zuela, and Colombia. Also in Trinidad’’
AMNH R-151970–151978 and USNM (Avila-Pires 1995:534).
566502–566506 from Konawaruk Camp. Vouchers for Guyana.—AMNH R-
Also ROM 20525–20528 from Amatuk; 61271–61272 from Wichabai; AMNH R-
ROM 20529 from Kurupukari; ROM 61331 from Marudi; AMNH R-61487–
22901–22920 and USNM 535806 from 61488 from Haiowa Falls, Essequibo
Baramita; and USNM 531684–531693 River; AMNH R-61377–61378 from Ish-
from Iwokrama Forest Reserve (various eartun; AMNH R-138057 from Yupukari
localities). (on Rupununi River), 11 km (airline) SSW
of Karanambo, northern Rupununi Sa-
Coloration in life.—The only notes
vanna; AMNH R-138083–138085 from the
taken in life in Guyana were based on a
Simoni area, about 16 km (by trail) E
male at Dubulay Ranch (CJC field notes,
Yupukari; AMNH R-60881–60884, R-
1994), noted to have green and cream
138088–138091, R-138096–138098, and
spots on the sides. Detailed notes were not
USNM 566508–566510 from the Kara-
taken at that time because CJC and CRT
nambo area; AMNH R-139880–139882
recognized the Guyanan individuals as from Aishalton; ROM 28353–28357 from
being like those with which they had Paramakatoi; USNM 163018 and 163041–
experience previously in Suriname, so 163044 from Enmore Estate (6844 0 N,
the following notes paraphrased from 057859 0 W); and USNM 566507 from
Cole et al. (1995:4–5) are relevant. Dor- Dubulay Ranch. In addition, ‘‘RMNH
sum of adults brown with green wash 25405–406’’ ‘‘from Ogle, just W [sic] of
anteriorly, with bright green (anteriorly) Georgetown’’ (Avila-Pires 1995:530).
or cream to tan middorsal stripe from Coloration in life.—The following is
snout or top of head to about midbody; based on AMNH and USNM specimens
cream, tan, or green (anteriorly) dorso- from the areas of Karanambo, Yupukari,
lateral light stripe; row of dark brown Simoni, Aishalton, and Dubulay Ranch
spots above the dorsolateral light stripe; (CJC field notes, 1992–1995). On large
some adults with pale cream, tan, green, adults, head and neck (to level of arms)
gray, or blue spots on sides; tail brown washed with bright green dorsally and
with dark brown, sometimes also tan or laterally, including lips; paler yellowish
cream, spots; on some individuals, tan or green on chin, throat; cream or tan light
light brown spots on dorsal surface of spots on sides. Otherwise, dorsum gray,
hind legs. grayish brown or brown, with paired
Comments.—See comments above for paravertebral series of small dark brown
the unisexual species, K. borckiana, and spots; dorsolateral light stripe dirty cream
the discussion by Avila-Pires (1995) on (may be green anteriorly, tan posteriorly);
lateral light stripe gray; brown or reddish some anteriorly, elliptical markings; arms
brown dark stripes, lower one with a black similar but legs with tan spots instead of
irregular streak or spots; brownish gray bands; on some, pale gold dots on body,
below lateral light stripe, with inconspicu- arms, legs, tail; tail black with yellowish
ous light tan stripe within; top of head tan bands; head above mottled black and
brown; lips tan (or with green wash, tan; venter yellow or orangish yellow with
perhaps bright); corner of mouth greenish black spots on chin, throat, chest; black
yellow (color absent from smallest individ- bands encircling tail.
uals); arms, legs, tail tannish gray to Comments.—This is one of the largest
brown; legs sometimes with dark brown species of lizards in the Western Hemi-
spots, tail usually with vertebral row of sphere, although it is exceeded in length by
dark brown spots; chin white or pale tan; Iguana iguana. Avila-Pires (1995) men-
throat, chest, beneath arms, anterior ab- tioned a preserved specimen from Brazil
domen tan (orange in large adults); poste- with a total length of about 1 m. This
rior abdomen, beneath legs, preanal area, species is primarily terrestrial. In some
beneath tail pale tannish orange, brightest literature in the recent past, the species was
under legs and base of tail (bright orange referred to as T. nigropunctatus, but that is
on adult males and females); posterior a junior synonym, as discussed by Avila-
underside of tail brown. Pires (1995).
Comments.—See comments above for
the unisexual species, K. borckiana.
Tropiduridae
Tupinambis teguixin (Linnaeus, 1758) This is one of the families of iguanian

Plate 27B lizards (see introduction above, before
Corytophanidae). Representatives of this
Type material.—The original name was family occur in ‘‘the Bahama Islands,
Lacerta Teguixin and the lectotype is Cuba and Hispaniola and associated
UUZM, Linnaeus Coll. No. 14 (Presch banks; Cayman Islands; South America,
1973). Presch (1973) restricted the type excluding northern Colombia and north-
locality to the vicinity of Paramaribo, ern Venezuela, southward to northern
Surinam. Tierra del Fuego; Galapagos Islands’’
Distribution.—‘‘Northern South Ameri- (Frost & Etheridge 1989:45). At one point,
ca (French Guiana, Suriname, Guyana, the genera Plica, Uracentron, and Urano-
Venezuela, Colombia, Ecuador, Peru, Bo- scodon were synonymized with Tropidurus,
livia, and Brazil)’’ (Avila-Pires 1995:563). but they have all been resurrected as they
Vouchers for Guyana.—AMNH R- appear to be monophyletic groups after all
18190 and R-107640 from Kartabo; (Frost et al. 2001).
AMNH R-25039 from Kamakusa;
AMNH R-140937–140938 from Dubulay
Ranch; AMNH field number JC 7829 Plica plica (Linnaeus, 1758)
from Konawaruk Camp (given to UG); Plate 27D
and USNM 84514–84515, 86869, 89368,
and 94412–94414 from Pomeroon. Type material.—The original name was
Coloration in life.—The following is Lacerta Plica and the holotype is NRM
based on AMNH and UG specimens from 112 (Avila-Pires 1995). Hoogmoed (1973)
Dubulay Ranch and Konawaruk Camp restricted the type locality to the conflu-
(CJC field notes, 1994, 1998). Black above ence of the Cottica River and Perica
with tan to yellow or gold bands and, on Creek, Surinam.
Distribution.—‘‘Northern South Ameri- UUZM 73 (Avila-Pires 1995). Etheridge

ca east of the Andes, throughout most of (1970) restricted the type locality to the
the Amazonian region (Brazil, French vicinity of Paramaribo, Surinam.
Guiana, Suriname, Guyana, Venezuela, Distribution.—‘‘South America north of
Colombia, Ecuador, Peru, and Bolivia) Rio Amazonas/Solimões and Rio Japurá,
and reaching the Atlantic coast in the in Brazil, French Guiana, Suriname and
Guianas and eastern Venezuela. Also in Guyana’’ (Avila-Pires 1995:177). This
Trinidad’’ (Avila-Pires 1995:170). range is for the subspecies Avila-Pires
Vouchers for Guyana.—AMNH R-8090 recognized as P. umbra umbra, which is
from Kamaria Landing, Cuyuni River; the form that occurs in Guyana.
AMNH R-8091 from Groeta Creek, Esse- Vouchers for Guyana.—AMNH R-8097
quibo River; AMNH R-14117, R-21290, from Kalacoon; AMNH R-14115–14116
and R-21331 from Kartabo; AMNH R- and R-21316 from Kartabo; AMNH R-
25071, R-25089, and R-25113–25115 from 17690–17691 from Georgetown; AMNH
Kamakusa; AMNH R-61316 from Kuyu- R-25118 from Kamakusa; AMNH R-
wini Landing; and AMNH R-151847– 61436 from Isheartun; AMNH R-141879
151849 and USNM 566366 from Kona- from Dubulay Ranch; AMNH R-151851
waruk Camp. Also ROM 22850 and from Berbice River Camp; AMNH R-
USNM 535795–535800 from Baramita; 151853–151863 and USNM 566367–
USNM 84492–84505 and 94408–94411 566373 from Konawaruk Camp. Also
from Pomeroon; and USNM 164186– ROM 22851–22852 from Baramita; ROM
164188 from Mabaruma Compound 20507–20508 from Tukeit; ROM 28350
(8812 0 N, 059847 0 W). from Paramakatoi; ROM 43206 from
Coloration in life.—The following is Kaieteur Plateau; USNM 164189 from
based on AMNH and USNM specimens M a b a r u m a C o m p o u n d ( 8 81 2 0 N ,
from Konawaruk Camp (CJC field notes, 059847 0 W); USNM 326119–326120 from
1998). Brownish green with black mark- Kwakwani (5830 0 N, 058800 0 W); and
ings on body and tail; arms, legs, hands, USNM 531645–531649 from Iwokrama
feet brownish green or gray, with black Forest Reserve (various localities).
bands (not complete across undersides); Coloration in life.—The following is
chin and throat pale yellowish green with based on AMNH specimens from Dubu-
black markings; underside of neck subtle lay Ranch and Berbice River Camp, and
orange anterior to black patch; beneath
AMNH and USNM specimens from
arms, legs, chest, posterior abdomen,
preanal area very pale yellowish green;
1995–1998). Brownish green or green
orange on posterior chest, abdomen (per-
above, including head, with dark brown
haps with orangish yellow patch posteri-
or black markings (basically bands) on
orly), beneath hips, beneath base of tail
body, arms, legs, tail (ground color of tail
(particularly bright ventrolaterally); iris
on some light tan); lips possibly bluish
copper.
green; orangish yellow beneath neck,
becoming paler on throat and chin;
orangish yellow patches on abdomen,
Plica umbra (Linnaeus, 1758) thighs, preanal area, beneath tail (bright-
Plate 27E est at base); other ventral surfaces light
tan to grayish orange. An adult male was
Type material.—The original name was noted specifically to have a bright yellow-
Lacerta Umbra and there are three syn- ish orange throat, and an adult female as
types, NRM 111 (2 specimens) and having only a trace of this color.
Tropidurus hispidus (Spix, 1825) yellow to orange; underside of base of tail

Plate 27C very pale yellow to orange, becoming
cream then gray posteriorly; lips pale
Type material.—The original name was yellow or orange with gray smudges. Adult
Agama hispida and the lectotype is RMNH males have orange in the axilla and gray
2912 (Hoogmoed & Gruber 1983). The streaks on the chest. They also have dark
type locality was ‘‘restricted by Rodrigues, patches of small, round black spots on the
1987 to Salvador, Bahia, Brazil’’ (Avila- underside of the thighs and preanal area,
Pires 1995:186). which may be seasonal black patches
Distribution.—Avila-Pires (1995:190) correlated with the reproductive cycle, as
stated that this species occurs in parts of noted for T. semitaeniatus by Ribeiro et al.
Brazil, southern French Guiana, ‘‘Suri- (2010).
name, Guyana, and Venezuela.’’ Comments.—We noted slight differences
Vouchers for Guyana.—AMNH R- in coloration of the lizards from Dubulay
137915, R-137920–137924, R-137927, and Ranch versus the Rupununi Savanna, but
USNM 566375–566378 from Karanambo; this might have reflected seasonal or
AMNH 138086 from the Simoni area, geographic variation.
about 16 km (by trail) E of Yupukari,
northern Rupununi Savanna; AMNH R-
139791–139803, R-139811–139812, and
Uracentron azureum (Linnaeus, 1758)
USNM 566379–566385 from Aishalton;
and AMNH R-140906–140915, R- Type material.—The original name was
140921–140922, and USNM 497794– Lacerta azurea and there are two syntypes,
497797 and 566386–566390 from Dubulay NRM 113 (Avila-Pires 1995). Hoogmoed
Ranch. Also, USNM 146375 from Leth- (1973) restricted the type locality to the
em; USNM 162895–162904 from Atkinson confluence of the Cottica River and Perica
(¼Timehri Airport); USNM 163054 from Creek, Surinam.
Enmore Estate (6844 0 N, 057859 0 W); Distribution.—For U. azureum azureum,
USNM 291132 and 291134–291137 from the form that occurs in Guyana, ‘‘eastern
Kato; and USNM 497798 from CEIBA Amazonia, in Guyana, Suriname, French
B i o l o g i c a l S t a t i o n ( 0 6 82 9 0 5 7 00 N , Guiana, and Brazil’’ (Avila-Pires
058813 0 06 00 W). 1995:207).
Coloration in life.—The following is Vouchers for Guyana.—BMNH
based on AMNH and USNM specimens 1905.10.21.1 and 62.12.15.36 from ‘‘British
from Karanambo, Yupukari, Aishalton, Guiana’’ (quoted from the BMNH cata-
and Dubulay Ranch (CJC field notes, log; no additional locality data). Colin
1992–1994). Dorsum gray, tan, or brown, McCarthy at the BMNH confirmed the
with indistinct dark brown to black or pale catalog data and sent us digital photo-
tan spots or bands and indistinct pale tan, graphs of both specimens for confirmation
orangish yellow, or greenish yellow wash; of the identification.
some individuals with indistinct, diffuse, Coloration in life.—The following is
very light gray dorsolateral stripe; top of from Hoogmoed (1973:196), based on a
head, arms, legs, tail similar to body; dark specimen from Suriname. ‘‘Colour in life
brown to black collar on neck; chin, of RMNH 15249, above bright grass-
throat, chest pale yellow or orange, on green with black cross-bands. Belly,
some with gray smudges or broad gray throat and ventral surface of the fore-
wash on throat; abdomen yellow to limbs yellow-green, chin green. Ventral
orange, paler posteriomedially; underside surface of the hind limbs chrome-yellow.
of arms and legs dirty cream with pale Scales on the dorsal surface of the tail
green, the outer and next to outer row from Aishalton, Dubulay Ranch, Berbice
with the tips of the spines yellow; scales River Camp, and Konawaruk Camp
under the tail yellow-ochre and green. (CJC field notes, 1993, 1994, 1996–
Iris golden brown.’’ 1998). Dorsum, including head and tail,
Comments.—Only two specimens are brown; alternating tan and black dashes
known from Guyana (see Vouchers, or dark brown bands on vertebral line,
above). Little is known about the life continuing onto tail; arms and legs
history, behavior, and ecology of this brown, with black bands and irregular
species. Avila-Pires (1995:212) stated that spots; dark brown and tan lines radiating
‘‘it is predominantly an arboreal forest from eye; pale tan, tannish yellow, or
inhabitant. Probably it mainly lives in the orange flecks low on sides; chin may be
canopy, which could explain why it is only dark grayish brown; white, dirty cream,
rarely observed.’’ or yellow on throat, chest, abdomen, and
beneath arms, legs, and tail; some scat-
tered brown to black spots or gray
Uranoscodon superciliosus smudges on chest, posterior abdomen
(Linnaeus, 1758) and beneath legs. A female had a broad
Plate 28A orange streak on her sides.
Comments.—We usually found individ-
Type material.—The original name was uals of this species in trees that are in or
Lacerta superciliosa and there are three adjacent to bodies of water. Massary &
syntypes, NRM 109–110, and UUZM Ineich (1999:168) reported on an adult
Linnean collection no. 69 (Avila-Pires male that ‘‘was captured, photographed,
1995). Hoogmoed (1973) restricted the measured, and released at 1545 h from a
type locality to the confluence of the boat ca. 9 m from shore. The specimen
Cottica River and Perica Creek, Surinam. dived just in front of us and remained at
Distribution.—‘‘Northern South Ameri- a depth of ca. 22 cm for 24 min 40 sec.
ca east of the Andes, in Brazil, French The lizard then emerged for ca. 1 sec,
Guiana, Suriname, eastern Venezuela, took a single breath, and submerged a
eastern Colombia, and northeastern Peru’’ second time’’ for an unknown duration.
(Avila-Pires 1995:228).
15133–15135 and R-21317 from Kartabo; Species Accounts: Reptilia:
AMNH R-61306 from Isheartun; AMNH Squamata: Snakes
R-613 90– 6139 1 f rom Ka ranambo;
AMNH R-139788–139790 from Aishal- Snakes of one kind or another have
ton; AMNH R-140949 from Dubulay been around for at least 165 million yr,
Ranch; AMNH R-151871–151872 and based on fossils and modern phylogenetic
USNM 566391 from Berbice River Camp; analyses using DNA sequence data (Vidal
and AMNH R-151876, R-151878–151879, et al. 2010a). The oldest families in
and USNM 566392–566394 from Kona- Guyana, those having the species among
waruk Camp. Also, ROM 20509 from the first to evolve are the Aniliidae,
Kurupukari; USNM 497799 from CEIBA Anomalepidae, Boidae, Leptotyphlopidae,
B i o l o g i c a l S t a t i o n ( 0 6 82 9 0 5 7 00 N , and Typhlopidae. The modern snakes (of
058813 0 16 00 W); and USNM 531650– most recent origin) are the Colubridae
531657 from Iwokrama Forest Reserve sensu lato, Elapidae (coral snakes), and
(various localities). Viperidae. Despite the use of such words
Coloration in life.—The following is as ‘‘oldest’’ and ‘‘modern’’ here, every
based on AMNH and USNM specimens species alive today is a modern represen-
tative of its group, no matter how ancient does not have reliable locality data, is
the group’s earliest representatives may NRM 13 (McDiarmid et al. 1999).
have been. Distribution.—‘‘Northern South Ameri-
Since 1995, several groups of investiga- ca from southern and eastern Venezuela,
tors have been analyzing phylogenetic Guyana, Suriname, French Guiana
relationships of snakes using DNA se- through the Amazon Basin of Colombia,
quence data, including Lawson et al. Ecuador, and Brazil’’ (McDiarmid et al.
(2005), Vidal et al. (2007, 2010a), Zaher 1999:157).
et al. (2009), and Pyron et al. (2011). The Vouchers for Guyana.—AMNH R-8162
studies have involved numerous and from Kalacoon; AMNH R-21286 from
varied samples of taxa among the many Kartabo; USNM 85092 from McKenzie
kinds of snakes in the world, different (¼Linden); USNM 145457 from ‘‘George-
genes, and somewhat different techniques town, 50 mi above, on the Abary River’’;
and philosophy. Consequently there are USNM 566250 from Berbice River Camp;
some differences in proposed revisions to and field number JC 8087 from Konawa-
the classification of snakes, including at ruk Camp (given to UG).
the family level, and today there is not Coloration in life.—The following notes
full agreement among specialists as to are based on the USNM specimen from
how many families of snakes there are Berbice River Camp and the one from
and what names should be applied to Konawaruk Camp (CJC field notes, 1997,
them. As this is an active area of research 1998). Dorsum of alternating black and
with certainly more important changes to reddish orange bands; similar to this
come in the near future, we adopt a beneath head and tail (but reddish orange
rather conservative use of family names is paler); ventral surfaces of body pale
here. For now, at least 8 families, 52 yellow with black bands.
genera, and 97 species of snakes are Comments.—These snakes eat primarily
known to occur in Guyana. amphisbaenians, small snakes, and caeci-
lians, most of which are burrowing; they
forage ‘‘mainly on the ground, at night, as
Aniliidae well as in aquatic environments’’ (Maschio
et al. 2010:184).
The species Anilius scytale is so
distinctive morphologically among the
snakes of the world that it has been Anomalepidae
assigned to a family of its own for
nearly 200 yr and recent analyses of These small burrowing snakes are su-
DNA sequence data indicate that this perficially similar to worm snakes (Typhlo-
species belongs to an ancient lineage pidae; see introduction to that family,
(Noonan & Sites 2010). It is not unusual below) but were recognized morphologi-
to find this colorful snake in Guyana, cally by Taylor (1939) as comprising a
although very little is known about its separate family. Only one of the few
natural history. surviving species of this clade of snakes
occurs in Guyana.
Anilius scytale (Linnaeus, 1758)

Plate 28B Typhlophis squamosus (Schlegel, 1839)
Anguis Scytale and the holotype, which Typhlops squamosus based on material
from Cayenne. Hahn (1980) stated the Boa constrictor Linnaeus, 1758
holotype was at the MNHN, now lost. Plate 28C
Distribution.—Atlantic lowlands of
South America from French Guiana, Type material.—According to McDiar-
mid et al. (1999:185) two syntypes exist:
Guyana, and Suriname south to northern
NRM 10 and NRM 20001, and a third is
Brazil (McDiarmid et al. 1999).
presumed lost. Credible locality data is
lacking for the types.
25051 from Kamakusa.
Distribution.—‘‘From northern México,
Coloration in life.—The following is through Central America . . . to South
from Starace (1998:68–69) based on spec- America North of latitude 358S. . .; Dom-
imens from French Guiana. ‘‘Dorsally, the inica and St. Lucia in the Lesser Antilles;
body is toned from black to dark brown. San Andrés and Providencia Islands (Co-
The head is pinky-white, the eyes are lombia), and many other continental
barely visible. The belly is uniformly islands along the Atlantic and Pacific
white.’’ He also stated that his specimens coasts of Mexico, Central and South
had some small white spots toward the America’’ (McDiarmid et al. 1999:185).
swollen rear of the snakes. Vouchers for Guyana.—AMNH R-
Comments.—Starace (1998:69) stated 65568 from Kartabo; USNM 164195–
that this species is nocturnal and fossorial, 164196 from Mabaruma Compound
‘‘found in primary rainforests. . ., inside ant ( 8 81 2 0 0 0 00 N , 0 5 9 84 7 0 0 0 00 W ) ; U S N M
and termite nests. Feeds on ants, ant eggs 497803 from Dubulay Ranch; and USNM
and ant larva as well as on termites. 566251 from 24 km NE Aishalton. Also,
Oviparous.’’ RDM photographed and released an
individual of this species at Baramita,
and CJC and CRT photographed and
released several at Dubulay Ranch and
Boidae Berbice River Camp.
These morphologically distinctive
from Starace (1998:92), based on speci-
snakes have been recognized as a family
mens from French Guiana. ‘‘The dorsal
separate from other snakes for nearly 200
[surface] is light brown, covered with dark
yr. They use constriction to subdue their [brown] geometric patterns while the tail
prey, give birth to living young, often are is red-brick coloured [with cream areas in
found in trees, can become quite large between].’’ There is a longitudinal mid-
(including the largest snake in the Western dorsal brown stripe on top of the head.
Hemisphere, the anaconda, Eunectes mur- Comments.—The specimens seen at Ber-
inus), and are fairly abundant in Guyana. bice River Camp included a female more
The common ancestor of the Boidae than 3 m in length. USNM 497803 was
existed approximately 70 million yr ago, captured at 1815 h, ca. 6 m high in a tree
so the modern distribution of the group as while constricting/ingesting a kiskadee
a whole (Western Hemisphere, Africa, (Pitangus sulphuratus) it had just caught.
Madagascar, Asia, southwestern Pacific
islands) may reflect primarily a history of
over-land dispersal within and on drifting
Corallus caninus (Linnaeus, 1758)
parts of Gondwanaland, rather than more Plate 28D
recent over-water dispersal or rafting
across the Pacific Ocean (Noonan & Sites Type material.—The original name was
2010). Boa canina and the holotype, known only
to be from America, is NRM ‘‘no. Lin. 8’’ variation of several variable characters in
(McDiarmid et al. 1999:190). the absence of geographically-correlated
Distribution.—‘‘Guyana, Suriname, analyses of clinal variation. Consequent-
French Guiana, eastern and southern ly, we personally still treat C. caninus as
Venezuela (states of Bolı́var and Amazo- one widely distributed species, awaiting
nas), and northeastern Brazil north of the more convincing evidence that it should
Rio Amazonas and north and east of the be partitioned into two. This species is
Rio Negro (in the states of Amapá, Pará, arboreal.
Roraima, and Amazonas),’’ as restricted
by Henderson et al. (2009:575; see Com-
ments, below). For C. caninus sensu lato,
Corallus hortulanus (Linnaeus, 1758)
include the broader Amazon Basin, south
Plate 28E, F
of the Amazon also, and ‘‘Amazonian
Colombia, Ecuador, Peru, and Bolivia, as Type material.—The original name was
well as in northwestern Colombia north Boa Hortulana and the holotype, from
of the Andes’’ (Henderson et al. ‘‘America,’’ is NRM ‘‘no. Lin. 7’’
2009:578). (McDiarmid et al. 1999:192).
Vouchers for Guyana.—AMNH R-8368 Distribution.—‘‘South America: south-
from Kalakun (Mazaruni River); AMNH ern Colombia east of the Andes, southern
R-14133 from Kartabo; AMNH R-25029 Venezuela, Guyana, Suriname, French
from Kamakusa (Mazaruni River); Guiana, and Amazonian Brazil, Ecuador,
AMNH R-60835 without data additional Peru and Bolivia’’ (McDiarmid et al.
to ‘‘British Guiana’’ (collected by R.
1999:193).
Snedigar, Terry-Holden Expedition);
BMNH 1929.7.13.5 from Demerara River;
from Kalacoon; AMNH R-67869 from
MCZ 33384 from Georgetown; and field
Kartabo; AMNH R-140234 and USNM
number JC 8066 from Konawaruk Camp
566252 from Dubulay Ranch; ROM 22849
(given to UG). In addition, Alexander
from Baramita; USNM 141763 and
Mendes reports that this species occurs at
Dubulay Ranch. 145462–145472 from ‘‘Georgetown, 50 mi
Coloration in life.—The following is above, on the Abary River’’; USNM
based on the UG specimen from Kona- 166830 from Atkinson Airport (¼Timehri
waruk Camp (CJC field notes, 1998). Airport); and USNM 531694–531696 from
Dorsum green (paler laterally) with light Iwokrama Forest Reserve (various locali-
crossbars; anteriorly on body, crossbars ties). In addition, CJC and CRT saw two
gray with yellow spots at lateralmost ends; individuals at their Berbice River Camp
posteriorly on body, crossbars white or (not collected).
cream with yellow edging here and there; Coloration in life.—The following is
venter yellow; green and yellow colors of based on the AMNH and USNM speci-
dorsum and venter interdigitate ventrolat- mens from Dubulay Ranch (CJC field
erally; iris tan. notes, 1994). Pale tan or gray above with
Comments.—Henderson et al. (2009) brown markings that are darker brown
confirmed that C. caninus occurs in dorsally than laterally; head yellowish tan
Guyana. They also named as a new or grayish tan with dark brown markings;
species specimens from populations to iris rust; chin very pale yellow; anterior
the south and west, which previously venter cream with tan or brown markings,
had been included in C. caninus. The posteriorly becoming dark tan or grayish
new species was diagnosed primarily on tan. Some have a touch of orange among
the basis of partitioning the range of the facial pits and on the chin.
Comments.—This species frequently has cenchria maurus to specific status. They

been referred to as Corallus enydris, but C. stated that E. cenchria occurs in humid
hortulanus has precedence (McDiarmid et forests and E. maurus occurs in drier
al. 1996). The species is arboreal. savannas, but their data were limited, they
studied no specimens from Guyana, they
performed no geographically-correlated
Epicrates cenchria (Linnaeus, 1758) analyses of variation, and the meristic
Plate 29A characters used to distinguish the forms
were part of a continuum divided in the
Type material.—The original name was middle. We consider this to be an interest-
Boa Cenchria and the holotype, from ing working hypothesis rather than a
Surinam, is NRM ‘‘no. Lin. 6’’ (McDiar- matter that has been settled. Whether
mid et al. 1999:196). these authors are correct could be tested
Distribution.—‘‘Lower Central America by obtaining samples along a north to
in Costa Rica and Panama; South America south transect through Guyana, including
east of the Andes in Colombia, Venezuela, humid forest and drier savanna habitats
Guayana [sic], Suriname, French Guiana, with gallery forests, and similar ecological
Brazil, Ecuador, Peru, Bolivia, Paraguay, transects in other areas of the Guianas and
and northern Argentina. . .; Trinidad and analyzing geographic variation. It also
Tobago’’ (McDiarmid et al. 1999:196). would be of interest to include molecular
data.
from Kalacoon; AMNH R-21285 and R-
We examined each of the Guyanan
voucher specimens listed above for the
from near Kartabo; and AMNH R-28956
characters used by Chippaux (1986), Star-
and R-60851 from ‘‘British Guiana’’ but
ace (1998), and Passos & Fernandes
with no additional locality data. Also,
UMMZ 46676–46677 from Dunoon, De- (2008). The number of ventral scales
merara River; UMMZ 66809 from Araka- ranged from 252–273 and subcaudals from
ka; USNM 566253 from Dubulay Ranch; 57–62, which is consistent with E. c.
and field number JC 7757 from Konawa- cenchria, as was the color pattern of each
ruk Camp (given to UG). specimen. The loreal scales of these spec-
Coloration in life.—The following is imens included one elongate upper loreal
based on the USNM specimen from (but 2 on the left side of one specimen) and
Dubulay Ranch (CJC field notes, 1994). 2–4 (usually 3) smaller ones below, which
Brown with dorsal circles, ellipses, and is more like E. c. maurus. This also
irregular ovals of tan edged in black; indicates that more research is needed on
lateral spots black, lined above in light these animals.
tan then black; ventrolaterally, spots very
dark brown to black on light brown
ground color; head brown with black Eunectes murinus (Linnaeus, 1758)
stripes; chin and throat cream; rest of Plate 29B
venter pale gray posteriorly through anal
plate; mottled light gray and black beneath Type material.—The original name was
tail. Boa murina and the holotype, only known
Comments.—Chippaux (1986), Starace to be from America, is NRM ‘‘no. Lin. 9’’
(1998), and Passos & Fernandes (2008) (McDiarmid et al. 1999:201).
treated the rainbow boas of the Guianan Distribution.—‘‘South America east of
Region as two species, not two subspecies the Andes in Colombia, Venezuela, the
as per prior authors, elevating Epicrates Guianas, Ecuador, Peru, Bolivia, and
Brazil; Trinidad’’ (McDiarmid et al. the ventrals. Chin pale pinkish-cinnamon

1999:201). flecked laterally on the throat with gray
Vouchers for Guyana.—AMNH R- and black. Ventrals warm buff with
14266 from Kartabo; AMNH R-25032 exceedingly variegated markings com-
from Kamakusa; USNM 85089–85091, posed of small, black, geometrical patches,
86640–86642, 89388–89389, and 90900 which frequently form two broken, longi-
from McKenzie (¼Linden); USNM tudinal lines.’’
141764 from ‘‘Georgetown, 50 mi above, Comments.—This is the largest species
on the Abary River’’; and USNM 497804 of snake in the Western Hemisphere, and
from Dubulay Ranch. Gerald and Wesley one of the largest in the world; adults can
King (Amerindian field assistants of CJC attain 10.5 m in length and have a heavy
and CRT) saw a large one (not collected) a body. They are ‘‘generally found in water-
short walking distance from our Konawa- courses and in marshlands’’ (Starace
ruk Camp. 1998:113). De Freitas (2009:98) reported
Coloration in life.—The following is that an individual of 5 m total length was
based on the juvenile USNM specimen swallowing an adult Boa constrictor of 1.8
from Dubulay Ranch (RPR field notes, m total length when found, which in turn
1994). ‘‘Dorsum olive green with dark had swallowed an Iguana iguana, ‘‘in the
brown spotting; venter pink cream with South Rupununi savannas of Guyana
lateral dark spotting forming lines on about 7 km N of Dadanawa Ranch
lateral edge of ventrals.’’ The following is (2.888N, 59.538W).’’
from Starace (1998:111–113) based on
specimens from French Guiana. ‘‘The
dominant dorsal colouring is light Colubridae, sensu lato
greeny-brown with black oscillated pat-
terns distributed on either side along the Colubridae sensu lato is a very large
spinal axe.’’ On each side of the head in the family of snakes (more than 2400 species),
temporal region is a light orange elongate including some representatives on nearly
triangular area. The ventral surface is all of the continents that support snakes.
generally light green with some black Many recent papers on snake taxonomy
spots. and phylogeny, including those based on
The following is from Beebe (1946:20) modern molecular and phylogenetic anal-
based on specimens from Kartabo, Guya- yses, have suggested ways to subdivide the
na, which he referred to as Eunectes gigas. traditional Colubridae into smaller possi-
‘‘Top of head dark olive, almost black on bly monophyletic groups involving multi-
orbits; side of face in front of eye, upper ple smaller families and often more
labials and post oculars dark olive; A restricted geographic distributions (see
broad cinnamon band equal in width to introductory paragraph for the snakes,
eye extends from eye to posterior point of above). One partition uses the family
jaw, becoming darker and narrower on Dipsadidae as a large family of primarily
side of neck. This cinnamon band is neotropical snakes. Two recent papers
bounded below by a narrowed band of (Zaher et al. 2009, Vidal et al. 2010a)
black which extends along the neck. Back reviewed the earlier analyses and proposed
olive green with many alternately placed, different generic taxonomic conclusions of
irregular, large, round and oval black their own, based on extensive new analyses
spots. Sides buffy brown with smaller, focused on South American species. These
more crowded spots of orange yellow, included 19 species of ‘‘colubrids’’ that
edged with black. The lowermost of these occur in Guyana, and for which the
are much broken and extend in laces over authors supported revising the familial
classification to place most of them in the 3. Zaher et al. (2009) suggested changing
family Dipsadidae. Nevertheless, in the the name Waglerophis merremii to
most recent paper with detailed phyloge- Xenodon merremii. This seems reason-
netic analyses, Pyron et al. (2011:340) used able and has not been opposed by other
the family Colubridae for the relevant authors, so we use Xenodon merremii
South American snakes, consistent with here.
‘‘a conservative approach to taxonomic 4. Zaher et al. (2009) suggested changing
changes,’’ which we follow here. the name Xenoxybelis argenteus to
Zaher et al. (2009) also argued for Philodryas argenteus. However, Vidal
changing the generic names applied to et al. (2010a:52) stated that ‘‘Philodryas
certain South American species, to be appears to be paraphyletic with respect
consistent with their understanding of to Xenoxybelis, but with low support,
monophyly and priority of authorship. and we recommend the recognition of
This included four species that occur in the morphologically distinctive genus
Guyana. Consistent with Curcio et al. Xenoxybelis pending further studies
(2009) and Vidal et al. (2010a) we followed with denser taxonomic and character
only one of these four suggestions, as sampling.’’ We retain use of Xenoxybe-
discussed below. Interestingly, three of lis argenteus here.
these four species have been jumping
among genera for some time, which
underscores the difficulties with colubrid
Apostolepis nigrolineata (Peters, 1869b)
classification.
1. Zaher et al. (2009) suggested changing Elapomorphus nigrolineatus and the holo-
the former name Erythrolamprus aescu- type is ZMB 6447 (Bauer et al. 1995:71).
lapii to Liophis aesculapii. However, The type locality was given as Guinea,
Curcio et al. (2009) and Vidal et al. ‘‘but assumed by Peters to be South
(2010a) proposed that Zaher et al. had America’’ (Peters & Orejas-Miranda
priority of authorship incorrect, that 1970:22).
Erythrolamprus has priority over Lio- Distribution.—This species is known
phis, that type species are available and only from widely scattered localities in
should be included in the analyses, and Guyana, French Guiana, and western and
that such a generic change should await eastern Amazonian Brazil (Lema & Albu-
‘‘more analyses...with greater taxon and querque 2010, Vidal et al. 1999 [1998]).
character sampling’’ (Vidal et al. Voucher for Guyana.—Colin McCarthy
2010a:52), although there is apparently of the BMNH provided us with the
a problem with nonmonophyly of the following details on the holotype for the
traditional Liophis, Erythrolamprus, junior synonym Apostolepis quinquelineata
and Umbrivaga. We retain use of (see comment below). The specimen is
Erythrolamprus aesculapii and the ge- BMNH 1946.1.9.59 (formerly
nus Liophis here. 1889.9.30.12) from ‘‘Demerara,’’ presented
2. Zaher et al. (2009) suggested changing by J. Quelch. This is the only specimen
the name Liophis lineatus to Lygophis from Guyana of which we are aware.
lineatus. Curcio et al. (2009) argued Coloration.—Lema & Albuquerque
similar to the case with Erythrolamprus, (2010:345), without specifying whether
that this change of generic name would their notes were based on living individu-
be premature. We retain use of Liophis als, stated the following: ‘‘dorsum reddish
lineatus here. brown...; a blackish brown head...lacking a
pair of cream blotches on preocular brown, separating the spots. A light spot
scales...a pair of cream blotches [present] on supralabials five and six. Belly...black
on neck covering the 3rd and 4th dorsal cross-bars...frequently only extend to the
scales on 5th scale row on each side...The middle [of the belly] and alternate with
caudal band is dark brown...and covers their counterparts.’’
only the dorsal surface of the end of the
tail.’’ Starace (1998:268), based on pre-
served specimens, added that there are five Atractus schach (Boie, 1827)
dark brown stripes on the body, the
ventral surface is yellow, and the end of Type material.—The original name was
the tail can be black instead of dark Brachyorrhos schach and Hoogmoed
brown. (1980) designated RMNH 119a as the
Comments.—The genus Apostolepis is lectotype. He also restricted the type
poorly known taxonomically, as well as in locality to Mamadam, Saramacca River,
other aspects of its biology. In the last few Brokopondo, Surinam.
years, the one species that occurs in Distribution.—Known from western and
Guyana has been referred to with three central Surinam and Manaus, Brazil
different names: A. quinquelineata, A. (Hoogmoed 1980); now widely in the
flavotorquata, and A. nigrolineata. Amazon Basin (Hoogmoed, pers. comm.).
Vouchers for Guyana.—BMNH
1939.1.1.95–1939.1.1.96 from ‘‘New River,
Atractus favae (Filippi, 1840) Guiana, 750 0 ,’’ identified by F. Irish
(quoted from catalog records of the
Type material.—The original name was BMNH; no additional data).
Calamaria Favae and type(s) and type Coloration in life.—Hoogmoed
locality are unknown to us. (1980:34) described the coloration of a
Distribution.—Hoogmoed (1980:19) living individual from Surinam, based on
stated that the only known localities for color slides as: ‘‘The back is orange brown
this species are in Guyana and Surinam with black blotches. The belly is white with
and that ‘‘probably this species is endemic a median row of black spots, underside of
to the Guiana Shield, possibly only to a tail grey.’’ He also described (p. 33) the
small part of it in northern Suriname and pattern of preserved specimens: ‘‘Snout
Guyana.’’ and anterior part of head black, a light
Vouchers for Guyana.—AMNH R- brown, transverse band on the back of the
17678 from Georgetown; AMNH R- head, followed by a black band on the
61557 from Wismar, both identified by neck. Body light brown with rectangular
M. S. Hoogmoed (1980) and confirmed black blotches, those of both flanks either
recently by CJC; BMNH 1934.11.1.132 alternating or confluent. A vague but
from ‘‘Demerara near Mackenzie’’ distinct black vertebral line. Blotches on
(Hoogmoed 1980:16). flanks often bifurcating near the ventrals.
Coloration.—Hoogmoed (1980:19) stat- Belly immaculate or with a single row of
ed that ‘‘according to Parker (1935) it is brown or black spots down the middle,
‘black above and banded black and coral- posteriorly accompanied by other brown
red beneath with a white spot around the or black spots. Underside of tail either
vent.’’’ Hoogmoed (1980:18) described completely grey or heavily mottled with
preserved specimens as being ‘‘brown with brown.’’ In life, the ground color of the
an indistinct dark-brown vertebral stripe. belly is cream to yellow (Starace 1998:136).
Light spots on snout...Rostral and median Comments.—Colin McCarthy kindly
suture of internasals and prefrontals dark sent photographs of the two BMNH
voucher specimens to CJC. They compare descriptions of A. steyermarki in the

favorably with this species and with each literature (Roze 1958, Myers & Donnelly
other, so we accept this identification. 2008). All important characters of scala-
tion and color pattern visible on the
photographs fit reasonably closely with
Atractus steyermarki Roze, 1958 A. steyermarki and not A. zidoki or any
other species of the genus. This suggests
Type material.—The holotype is CNHM that A. steyermarki has a wider geographic
69920, a male from Chimantá Tepui, distribution than had been known and is
Bolı́var, Venezuela; 1430 m (Roze consistent with Roze’s (1961) comment
1958:301). about its possible occurrence on the Gran
Distribution.—This species is known Sabana of eastern Venezuela, not far from
only from Guyana and Venezuela, from Mt. Roraima. UF 150240 originally had
very few specimens. been reported (Means 2007) incorrectly as
Vouchers for Guyana.—BMNH 1976. representing Atractus zidoki.
348 from ‘‘Roraima Guyana, 7500–8000 0 ’’
(quoted from the BMNH records) and UF
150240 (six photographs of UG number Atractus tamessari Kok, 2006a
HR 701) from ‘‘GUYANA: Potaro-Sipa- Plate 29C
runi District, talus slope of Mt. Roraima
beneath its northernmost promontory Type material.—The holotype is IRSNB
called ‘The Prow,’ (05815 0 30 00 N, 60843 0 2640, an adult male from along a tributary
30 00 W), ca. 2090 m elev.’’ (Means of Elinkwa River, within the ESE area of
2007:484). Kaieteur National Park, ca. 500 m eleva-
Coloration.—Roze (1958) described the tion, Potaro-Siparuni District, Guyana
type material from Venezuela as being (5808 0 09 00 N, 59825 0 28 00 W).
dark gray above, the head similar to the Distribution.—Known only from the
body; white on the supralabials and lower type locality (Kok 2006a:25) and eastern
secondary temporals (absent on a melanis- Venezuela (Rivas et al. 2012).
tic specimen); lateral ends of ventrals with Vouchers for Guyana.—The three spec-
black spots forming a ventrolateral row; imens known are all from Kaieteur Na-
light beneath head and anterior ventral tional Park, Guyana, the type locality (see
surface, but soon with black spots on above). This includes the holotype (see
midventral line (with considerably more above) and two paratypes (IRSNB 2641–
black on melanistic individual); midventral 2642).
spots gradually increase in size posteriorly Coloration in life.—Kok (2006a:22–25)
to occupy most of middle of ventrals; described coloration of the holotype as
subcaudals black with light ends. follows. ‘‘The dorsal ground color is
Comments.—Colin McCarthy of the brownish black with numerous irregular
BMNH and Kenneth L. Krysko of the bright red and paler red markings. The tail
UF provided CJC with nine digital pho- is brownish black with a few bright red
tographs of the two Guyanan vouchers lateral markings. The top of the head is
cited above. These were compared with brownish black. The snout is lighter, with
AMNH specimens of various species of a slight yellow wash. The supralabials are
Atractus from the Guianan Region, in- mostly yellow with the upper third dark
cluding A. zidoki (AMNH R-131799 from brown, except for the eighth, which is
Brazil, identified by M. Rodriguez, and almost completely dark brown with a
AMNH R-108792 from Surinam, identi- small yellow spot anteriorly, and the first
fied by M. S. Hoogmoed), and with and the second, which are speckled with
dark brown. The infralabials are mostly Underside of tail orange. Iris chestnut
yellow with the lower third and posterior brown.’’
portion of each scale dark brown. The
chinshields are mostly yellow with dark
brown anteriorly and laterally. The mental Atractus trilineatus Wagler, 1828
groove area has a few dark brown Plate 29E
speckles. The venter is yellowish cream
with many irregular brownish black Type material.—Hoogmoed (1982) des-
blotches, fewer blotches anteriorly, heavily ignated RMNH 48 as the lectotype. There
mottled posteriorly. Underside of tail is are no reliable data for the type locality.
brownish black with a few small irregular Distribution.—‘‘A lowland Guiana en-
yellowish cream markings. The iris is demic, ranging from eastern Venezuela to
brown.’’ The paratypes showed variation the Guianas; and occurs on Trinidad,
in dorsal color tones and one specimen had Tobago and Little Tobago’’ (Murphy
two ventrolateral black stripes. 1997:166).
Comments.—Practically nothing is Vouchers for Guyana.—AMNH R-
known about the biology of this snake. 137334 from Kartabo, identified by A. L.
Markezich; ROM 19197 from ‘‘Demerara’’
(quoted from catalog; no additional local-
Atractus torquatus ity data); and USNM 566255 from Dubu-
(Duméril, Bibron, & Duméril, 1854) lay Ranch.
Plate 29D Coloration in life.—The following is
Rabdosoma torquatum and Hoogmoed
Brown above, with dark brown stripes;
(1980) designated RMNH 114, a male, as
chin and anterior throat pinkish tan; rest
the lectotype. Hoogmoed (1980) restricted
the type locality to Paramaribo, Surinam. of ventral surface lemon yellow. Beebe
Distribution.—This species is known (1946:21) described a specimen from Kar-
from Surinam, Guyana, French Guiana, tabo as having the ‘‘head above cinnamon
Peru, and Bolivia according to Hoogmoed brown with distinct asymmetrical mot-
(1980:38); ‘‘Amazonian Colombia and tlings of mummy brown. Back hazel
Bolivia; Amazonas Brazil’’ according to brown with three longitudinal stripes of
Peters & Orejas-Miranda (1970:35). dark clove brown. Upper labials citrine
Vouchers for Guyana.—AMNH R- yellow. The upper side of body below the
60788 from Kuyuwini Landing; USNM lateral dorsal stripes, tawny olive and
566254 from Dubulay Ranch; BMNH below this a faint, narrow line of brown.
72.10.16.72–72.10.16.73 from Demerara Lower labials, chin and anterior ventrals
Falls (Hoogmoed 1980:36). lemon yellow. Remaining ventrals mustard
Coloration in life.—The following is yellow with pinkish tinge. Iris dark cinna-
based on the USNM specimen from mon brown.’’
Dark brown above, with black markings
that tend to form bands; ventral surface Chironius carinatus
yellow, with black spots. Hoogmoed (Linnaeus, 1758)
(1980:38) stated that specimens from Sur-
inam had ‘‘upper parts iridescent dark Type material.—The original name was
greyish brown to reddish brown. Ventral Coluber carinatus and the holotype is
parts creamish to yellow with brown spots. NRM 33, which lacks reliable locality data
(http://linnaeus.nrm.se/zool/herp/ follows. ‘‘Dorsolateral body ground colour

madserp.html.en). light brown, darkening posteriorly, with a
Distribution.—According to Dixon et al. series of ca. 31 ill-defined greyish brown
(1993: 73) this species occurs in ‘‘north- transverse bands (1, rarely 2 scales wide)
eastern Brazil. . .French Guiana, Surinam, outlined in black. Tail darker brown
Guyana, eastern Venezuela, and Trini- without conspicuous transverse bands.
dad.’’ These authors recognized some Dorsal scales with black edges, sometimes
subspecies, but it is the nominate form slightly peppered with dark brown or
that occurs in Guyana, and the quote black; some dorsal scales yellow anteriorly,
covers only that form (see Comments, especially on anterior body and on lower
below). flank. Skin between scales sky blue. Head
Vouchers for Guyana.—AMNH R-8705 light brown, supralabials yellow, with
from Kalacoon and AMNH R-60781 from labials 2–3 having dark posterior edges;
Karanambo, identified by J. A. Wiest; and black postocular stripe from eye across
USNM 84520 from Pomeroon. lower part of primary temporal and upper
Coloration.—According to Dixon et al. part of ultimate supralabial reaching post-
(1993:75) ‘‘an ontogenetic change occurs, cephalic scales then breaking into blotches
in which light brown or olive juveniles on the lateral side of ventrals; iris dark
(some with faint flecking on dorsal scales copper; tongue black. Underside of head
or faint crossbands) become dark olive immaculate yellowish white; ventrals im-
adults with distinct yellow spots on the maculate yellow until ca. the 36th, then
first scale row of tail.’’ Starace (1998:144) posteriorly bordered by black, mottled
stated that this species is ‘‘olive green to with brown and black spots increasing in
brown, the sides are bluish and the belly is number posteriorly, becoming almost
orange-yellow. Often light brown, the head completely black near the vent; subcaudals
is slightly lighter than the body.’’ similar in colour and pattern, but less
Comments.—Dixon et al. (1993) recog- heavily mottled by black than the posteri-
nized three subspecies, but Hollis (2006) ormost ventral scales.’’
elevated each of them to specific status. Comments.—Kok (2010b) pointed out
Chironius carinatus is the form that occurs that this species is superficially similar to
in Guyana. Chironius fuscus. Some specimens that
were identified as C. fuscus before 2010
may actually be C. challenger.
Chironius challenger Kok, 2010b
Plate 29F
Chironius exoletus (Linnaeus, 1758)
Type material.—The holotype is IRSNB Plate 30A
2659, a subadult female from the south-
eastern slope of Maringma Tepui, Cuyuni- Type material.—The original name was
Mazaruni District, Guyana (05812 0 N, Coluber exoletus and the holotype is
060835 0 W; 1500 m) (Kok 2010b:32). ZMUU 150 or 135 as discussed by Dixon
Distribution.—The four known speci- et al. (1993:92–93). Data for the type
mens are from tepuis in Guyana and locality are not reliable.
Venezuela (Kok 2010b). Distribution.—‘‘Costa Rica south
Vouchers for Guyana.—The holotype; through western South America to Ecua-
and ROM 42603 from Mount Wokomung, dor, northern South America, the Gui-
1400 m elevation (Kok 2010b). anas, and the Amazon basin to Peru,
Coloration in life.—Kok (2010b:36–37) southern Brazil, and extreme northeastern
described the holotype from Guyana as Argentina’’ (Savage 2002:650).
Vouchers for Guyana.—AMNH R-6801 from Mount Wokomung; ROM 22845–

from Rockstone; AMNH R-14261–14262, 22846 from Baramita; and USNM 549326
R-15150, R-18169, and R-44907 from from E bank of the Waruma River
Kartabo; AMNH R-25048 from Kamaku- (5823 0 00 00 N, 060846 0 00 00 W).
sa; AMNH R-36107 from Lama Creek, Coloration in life.—The following is
Demerara River; AMNH R-57791 from based on the AMNH and USNM speci-
Wismar; and AMNH R-60823 from mens from Berbice River Camp (CJC field
Onora, all identified by J. A. Wiest. Also, notes, 1997). Dorsum chestnut brown with
AMNH R-137974 from Karanambo and tan bands; chin and throat white; venter
USNM 566256 from Konawaruk Camp. toward posterior becomes tan, then gray-
Coloration in life.—The following is ish tan, then tannish gray; iris copper.
based on the AMNH and USNM vouch- Starace (1998:150) presented a color pho-
ers from Karanambo and Konawaruk tograph showing a light tan vertebral
Camp (CJC field notes, 1992, 1998). stripe on the anterior part of the body
Dorsum brown or greenish brown; sides that is edged with dark brown.
same or brownish green; dark brown
dorsolateral stripe may be present anteri-
orly, disappearing on neck and midbody; Chironius multiventris
keels on scales of midbody with thin dark Schmidt & Walker, 1943
brown line; snout tan; top of head brown; Plate 30C
lips and neck yellow; white or yellow under
head; yellow or orangish tan belly; yellow- Type material.—The holotype, from the
ish tan or orangish yellow under tail, Department of Madre de Dios, Peru, is
perhaps with brown stripe on underside; FMNH 38250.
iris brown, with tan in uppermost and Distribution.—The map of Klaczko et
lowermost parts. al. (2010:477) and Starace (1998) give
records for Amazonian Brazil; eastern
Peru, Ecuador, and Colombia; then east-
Chironius fuscus (Linnaeus, 1758) ward through southeastern Venezuela,
Plate 30B Guyana, Suriname, and French Guiana.
Type material.—The original name was and R-8991 from Kalacoon; AMNH R-
Coluber fuscus and the holotype is NRM 15151, R-18174, and R-18175 from Karta-
34 (http://linnaeus.nrm.se/zool/herp/ bo; AMNH R-25030 from Kamakusa;
madserp.html.en), without reliable locality AMNH R-36137 from Georgetown;
data. AMNH R-60849 from lower Kuyuwini
Distribution.—‘‘Amazonian Brazil, River; USNM 164212 from Mabaruma
northern Bolivia, eastern Peru and Ecua- Compound (8812 0 00 00 N, 059847 0 00 00 W);
dor, southeastern Colombia, southern and USNM 566258 from Berbice River
Venezuela, Guyana, Surinam, and French Camp.
Guiana’’ (Dixon et al. 1993:115). Coloration.—The following is based on
Vouchers for Guyana.—AMNH R-6080 the USNM specimen from Berbice River
from Shudikarwau; AMNH R-18171 and Camp (CJC field notes, 1997). Dorsum
R-65561 from Kartabo; AMNH R-25033, dark greenish brown; ventral surfaces
R-25047, and R-25054 from Kamakusa; yellow; iris mostly brown. In describing
and AMNH R-152260–152261 and preserved specimens, Dixon et al.
USNM 566257 from Berbice River Camp; (1993:163) mentioned ‘‘a pale vertebral
ROM 39442–39444 from the base of stripe.’’ Klaczko et al. (2010:483) stated
Mount Ayanganna; ROM 42601–42602 that there are ‘‘two dark brown lines
following the keels of the paravertebral tween the scales); no stripe on face; venter
scales.’’ lighter green, bluish green anteriorly; lower
Comments.—In the past, several subspe- lips, chin, and throat pale pastel blue or
cies were recognized. Recently, however, turquoise, some of which extends up to the
Hollis (2006), elevated the northern one supralabials below the eye. Dixon et al.
that occurs in Guyana to a full species, (1993:192) stated that ‘‘AMNH 104608, a
Chironius cochranae, rather than Chironius subadult from Surinam, had the following
multiventris cochranae. Most recently, description on its attached field tag: above
Klaczko et al. (2010) concluded that dark green, sides brown; below light green;
cochranae is a junior synonym of Chironius eye pupil gray-green.’’ Starace (1998:157)
multiventris, and there is no justification mentioned that individuals change onto-
for treating it as a subspecies. genetically from being green juveniles to
become ‘‘adults...usually uniformly dark
brick-red, while the belly is of a lighter
Chironius scurrulus (Wagler, 1824) orange in colour.’’
Plate 30D
Type material.—The original name was Clelia clelia (Daudin, 1803a)
Natrix scurrula and the lectotype is ZSM
2628/0, a male (Hoogmoed & Gruber Type material.—The original name was
1983). The type locality is the Japura Coluber clelia based on MNHN material
River, Brazil or Colombia (Franzen & (now lost; Duellman 2005) from Sur-
Glaw 2007). inam.
Distribution.—‘‘Amazonian Brazil, Distribution.—‘‘Atlantic lowlands of
northern Bolivia, eastern Peru and Ecua- Guatemala south on the Atlantic versant
dor, southeastern Colombia, Trinidad, throughout Central America through Co-
eastern Venezuela, Guyana, Surinam and lombia, northern Venezuela, Trinidad and
French Guiana’’ (Dixon et al. 1993:190, Grenada, and south from the Guianas to
map on p. 193). central Bolivia, and on the Pacific slope
Vouchers for Guyana.—AMNH R-8360 from Costa Rica to northern Peru’’ (Sav-
from Kalacoon; AMNH R-14132 and R- age 2002:574).
15154 from Kartabo; AMNH R-17677 Vouchers for Guyana.—AMNH R-
from Georgetown; AMNH R-25037 from 14257–14258 and R-137337 from Karta-
Kamakusa; AMNH R-60752 from Shu- bo.
dikarwau; AMNH R-60794 from Para- Coloration in life.—The following is
bam; AMNH R-152263 and USNM from Starace (1998:273), based on spec-
566259 from Berbice River Camp; imens from French Guiana. ‘‘Along the
AMNH R-152265 from Konawaruk dorsum adults bear a uniformly charcoal
Camp; ROM 28376–28377 from Para- grey to black colouring. The belly is blue
makatoi; USNM 84519 from Pomeroon; grey...Newborns [and young] are red [on
USNM 531697 from Iwokrama Forest the dorsum] with black heads from the
Reserve, Kurupukari Base Camp; and snout up to the [anterior] edges of the
USNM 573506 from SW part of Woko- parietals. The rest of the head is white
mung Massif (5800 0 08 00 N, 059852 0 47 00 W). but becomes black again along 3 to 5
Coloration in life.—The following is mm of the neck...[so that young with red
based on the AMNH and USNM vouch- bodies] closely resemble Pseudoboa neu-
ers from Berbice River Camp and Kona- wiedii and Pseudoboa coronata.’’ The
waruk Camp (CJC field notes, 1997, 1998). young also resemble Drepanoides anom-
Dorsum leaf green (with black skin be- alus. Beebe (1946:23–24) described indi-
viduals from Kartabo as ‘‘white or body have faint blue edges. The chin and
yellowish-white or rarely salmon colored throat are white, and the belly is lemon
below.’’ Duellman (1978:235), describing yellow. The tongue is gray.’’
specimens from Ecuador, mentioned that
half-grown individuals have a dorsum of
‘‘dull reddish brown to brownish black’’ Dipsas catesbyi (Sentzen, 1796)
and ‘‘the ventrals and subcaudals...are Plate 30E
immaculate cream throughout develop-
ment.’’ Type material.—The original name was
Coluber Catesbyi based on material from
America, but ‘‘present location [of type
Dendrophidion dendrophis (Schlegel, 1837) material is] unknown’’ (Peters 1960:56).
Distribution.—‘‘Amazonas region of
Type material.—The original name was South America, from Andean slopes of
Herpetodryas dendrophis and the lectotype, Bolivia, Peru, Ecuador, and Colombia to
from Cayenne, French Guiana is MNHN coast of Venezuela and British Guiana,
41 (Lieb 1988). and through northern half of Brazil’’
Distribution.—As illustrated by Lieb’s (Peters 1960:56).
map (1988:167), this species occurs in far Vouchers for Guyana.—AMNH R-
eastern Venezuela, Guyana, Suriname, 17679 from Georgetown; AMNH R-
western French Guiana, Amazonian Bra- 18152, R-18153, R-21283, and R-98193
zil, Amazonian Colombia, and Ecuador from Kartabo; AMNH R-25034 and R-
and Peru east of the Andes. 25036 from Kamakusa; AMNH R-152267
Vouchers for Guyana.—AMNH R- and USNM 566260 from Berbice River
60820 and R-60833 from Parabam; Camp; ROM 39440 from Ayanganna;
AMNH R-60832 from Kuyuwini River. ROM 42597 from Potaro River; USNM
Also FMNH 30958 from ‘‘Demerara: 164203 from Mabaruma Compound
Itabu Creek Head, Boundary Camp’’ (Lieb ( 8 81 2 0 0 0 00 N , 0 5 9 84 7 0 0 0 00 W ) ; U S N M
1988:174). 200509–200510 from 24-Mile Forest Re-
Coloration in life.—The following is serve south of Bartica.
from Starace (1998:160), based on speci- Coloration in life.—The following is
mens from French Guiana. Dorsum ‘‘grey based on the AMNH and USNM speci-
in colour, fading to dark grey towards the mens from Berbice River Camp (CJC field
tail, the body is covered with...[dark] notes, 1997). Dorsum with alternating
coloured spots along the dorsum and sides. black and brown bands (but white ventro-
The belly is distinctly...[pale].’’ The follow- laterally); belly white with black markings
ing is from Duellman (1978:237), based on that increase in number posteriorly; iris
specimens from Ecuador. ‘‘The top of the brown. The following is from Duellman
head is olive-brown. The body has 49–55 (1978:237), based on specimens from
dark brown blotches 3 scales in width, Ecuador. ‘‘The head is black with a white
separated by narrow (½ scale in width) line across the snout; usually the line
creamy tan interspaces on the anterior half continues across the chin. The dorsum is
of the body. The anterior edge of each marked by paired elliptical dark brown
blotch is dark brown or black[;] posteriorly blotches, meeting or not, middorsally. The
the pale interspaces do not extend across dark chocolate brown blotches are bor-
the body, so the blotches are delimited dered with white, especially anteriorly, and
only by their dark anterior borders. stand out in marked contrast with the
Laterally the interspaces are orange. The reddish brown ground color. The venter is
dorsal scales on the anterior part of the white with longitudinal black marks and
flecks, especially posteriorly. The iris is across the dorsal... [surface]...[mostly on]
reddish brown.’’ the sides...The belly is light-coloured.’’
Dipsas copei (Günther, 1872) Dipsas pakaraima

MacCulloch & Lathrop, 2004a
Type material.—The original name was Plate 30F
Leptognathus Copei and the holotype is a
male, probably from Surinam, in the Type material.—The holotype is ROM
BMNH (Peters 1960:58). 41233, an adult male from the northeast
Distribution.—‘‘Guianas and southern plateau of Mount Ayanganna, Guyana
Venezuela’’ (Peters & Orejas-Miranda (05824 0 N, 059857 0 W; 1490 m).
1970:86). Distribution.—Known only from the
Vouchers for Guyana.—BMNH type locality (MacCulloch & Lathrop
1920.1.13.2 from Georgetown, cited by 2004a).
Peters (1960). Vouchers for Guyana.—In addition to
Coloration in life.—The following is the holotype, the paratypes (ROM 41234,
from Starace (1998:169), based on speci- ROM 41236, and USNM 561837) are all
mens from French Guiana. ‘‘The upper from the vicinity of the type locality
[anterior] quarter of the body is bright (MacCulloch & Lathrop 2004a:240).
yellow fading into light brown in the lower Coloration in life.—‘‘Dorsolateral
[posterior] regions. The dorsal regions are ground colour light brown anteriorly,
covered with dark brown spots, [border- darkening posteriorly. A series of 22 pairs
ed]...with a very [narrow]...white [line]... of medium brown dorso-lateral blotches,
Apart from a black band [connecting]...the outlined in black, between neck and vent.
eyes, the head is orange in colour...The Anteriorly the blotches are each about five
belly is light brown.’’ scales long middorsally, lengthening to 8–9
scales dorsolaterally, then tapering to five
scales on the first dorsal scale row.
Dipsas indica Laurenti, 1768 Posteriorly the blotches become shorter.
Anteriorly the blotches are separated by
Type material.—‘‘Unknown’’ (Peters one or two lateral scales; toward midbody
1960:68). The type locality was restricted this separation diminishes, and the black
by Peters (1960) to the Amazon region of borders of adjacent blotches are in contact.
South America. In the five posteriormost blotches the
Distribution.—‘‘Amazon drainage in black borders are reduced laterally, present
Brazil, Colombia, British Guiana, Ecuador only middorsally and on the ventrals. This
and Peru’’ (Peters & Orejas-Miranda results in a continuous uniform medium
1970:87). Also in French Guiana and brown lateral colouration.
Surinam (M. S. Hoogmoed, pers. comm.). ‘‘Many pairs of blotches are not exactly
Vouchers for Guyana.—AMNH R- opposite or symmetrical but rather alter-
60857 from Marudi (2807 0 N, 59801 0 W) nating, creating a pattern of staggered
and AMNH R-137335 from Kartabo. blotches along the back. This asymmetry
Coloration in life.—The following is is caused by a blotch being a different
from Starace (1998:171), based on speci- length from its corresponding opposite
mens from French Guiana. ‘‘The [light]...- blotch. In the holotype the blotch pairs
brown body is patterned with darker change from opposite to alternating to
brown irregular saddle-shaped spots. Ir- opposite twice along the length of the
regular white spots are also scattered body...
‘‘Within the dorsal blotches the scales UMMZ 47747 from ‘‘Dunoon, Demerara
appear to have dark tips, the result of River’’ (cited by Harvey 2009 [2008]:449).
overlapping the next posterior scale. How- Coloration in life.—The following is
ever, the lighter-coloured scales outside the from Starace (1998:173), based on speci-
blotches do not appear dark-tipped where mens from French Guiana. ‘‘Closely re-
they overlap the next scale. Between the sembles Dipsas catesbyi. The [anterior]...
blotches the light brown background third of the body is an off-white colour,
creates a middorsal pattern of diamond fading [to]...beige [or]...brown [posterior-
shapes (or half-diamond where the blotch- ly]... Round black spots, with white-edges
es are not symmetrical): these diamonds ...[mid-dorsally]. The black head is marked
contain small dark brown spots. The by a white crescent-shaped spot...[anterior
dorsal background colour becomes pro- to] the neck. A thin white line runs
gressively darker posteriorly, converging [across]...the snout to the supralabials.
with the blotch colour. The light brown [or white] belly is covered
‘‘Ventral background colour pale with very fine dark brown spots.’’
brown, the dorsolateral blotches extending
to the midventral point, each covering two
or three ventrals. Ventrals otherwise im- Dipsas variegata
maculate anteriorly, posteriorly with small (Duméril, Bibron, & Duméril, 1854)
black patches consisting of several juxta- Plate 31A
posed or narrowly separated squarish to
half-moon shaped marks. The ventral dark Type material.—The original name was
markings become more intense posterior- Leptognathus variegatus based on two
ly... specimens from Surinam; one in the
‘‘Tail with a narrow cream middorsal MNHN, one in the RMNH (Peters
stripe flanked by narrow black stripes. 1960:132).
Background colour is dark brown laterally Distribution.—‘‘Distributed from...Ve-
and ventrally, with a few small cream-and- nezuela. . .to French Guiana,. . .thence
black lateral blotches proximally. southward to the mouth of the Amazon
‘‘Head dark brown dorsally and lateral- River in the state of Pará, Brazil’’ (Cadle &
ly. Lower labials with a narrow white band Myers 2003:5).
at lip. Chin medium brown with light Vouchers for Guyana.—AMNH R-
brown mottling increasing posteriorly’’ 21275, R-81425, and R-98194 from Karta-
(MacCulloch & Lathrop 2004a:241–242). bo (as cited in Cadle & Myers 2003:44);
ROM 28371 from Paramakatoi; ROM
42596 from Wokomung; and USNM
Dipsas pavonina Schlegel, 1837 535807 from Baramita. Harvey (2009)
cited UMMZ 47757–47758 from Dunoon;
Type material.—Types, ‘‘apparently UMMZ 53900 from Demerara River; and
from Guyanes’’ are in the MNHN and UMMZ 76990 and 77504 from Wismar.
‘‘Musée des Pay-Bas [¼ RMNH]’’ (Peters Coloration in life.—The following is
1960:61). from Starace (1998:176), based on speci-
Distribution.—‘‘Guianas to Pará, Brazil, mens from French Guiana. ‘‘Light brown
and to Amazonian slopes of Andes; body along which are [irregularly alter-
Colombia to Bolivia, on eastern slope’’ nating]...dark brown spots. On the [ante-
(Peters 1960:61). rior]...third of the body the spots tend to
Vouchers for Guyana.—AMNH R- be saddle-shaped, [becoming irregular ven-
18154 and R-81424 from Kartabo; trally]...as they join the white to off-yellow
coloured belly. [Small yellow spots]...cover southeastern Georgia, Florida and south-
the entire body.’’ ern Alabama [USA]’’ (Savage 2002:659).
Voucher for Guyana.—AMNH R-60821
from Parabam and USNM 84523 from
Pomeroon.
Drepanoides anomalus (Jan, 1863) Coloration in life.—The following is from
Beebe (1946:27), who reported seeing the
Type material.—The original name was species at Kartabo, but whose color notes
Cloelia anomala and the lectotype is were based on a specimen from Caripito,
MNHN 4 (Cat. 100) (Duellman 2005). Venezuela. ‘‘Above blue black as far back as
See Peters & Orejas-Miranda (1970) for a 600 mm. before the tail, when the dorsal
discussion of the uncertain type locality. ground color changes to empire yellow and
Distribution.—‘‘Central Bolivia north to then to warm orange. The black is continued
Southern Colombia along Andean front’’ in this yellow area as paired bands. After four
(Peters & Orejas-Miranda 1970:93). Avila- of these pairs, the bands begin to degenerate,
Pires (2005:36) listed this species as wide- diminishing in width and purity, but they
spread in the Guiana Shield and Starace continue in a succession of about 30 to the tip
(1998:267) presented a map of its occur- of the tail. The rostral, nasals, internasals,
rence in French Guiana. loreals and all the upper labials are ivory
Vouchers for Guyana.—AMNH R- white, with a flecking of black on the loreals
137569 from Guyana, with no additional and [some]. . .labials. This white color con-
data, collected by W. Beebe and identified tinues on the chin, throat and ventrals, each
by Roy W. McDiarmid. of the latter with an invasion of black from
Coloration in life.—The following is each side, never meeting in the middle. The
paraphrased from Starace (1998:267– subcaudals are yellow orange like the upper
268), based on material from French side. The iris is dragon’s blood red with a
Guiana. The dorsum is basically uniform gold pupil ring.’’ The photo in Starace
red, each scale with a dark border; sides of (1998:181) shows the posterior body and tail
body generally lighter than uppermost and the side of the face as being basically
body; head black; white collar in occipital bright yellow.
Comments.—Although Beebe (1946)
region on rear of head; belly white. These
mentioned seeing this species at Kartabo,
snakes resemble juveniles of Clelia clelia
and it is expected to occur there, the Beebe
and of both species of Pseudoboa.
material at the AMNH does not include any
specimens from this locality. Consequently,
we have not listed this species in Table 1.
Drymarchon corais (Boie, 1827)
Coluber corais based on a specimen from Drymobius rhombifer (Günther, 1860)
‘‘America.’’ The type is unknown to us. Type material.—The original name was
Distribution.—‘‘. . .southern Sonora, Coryphodon rhombifer based on a speci-
Mexico, and southern Texas to extreme men (unknown to us) from Esmeraldas,
northwestern Peru on the Pacific versant, Ecuador.
to southern Brazil, Paraguay, and extreme Distribution.—‘‘. . .Nicaragua south
northern Argentina, including most of through Central America, Colombia, Ven-
South America east of the Andes and Isla ezuela, and the Guianas and over much of
Margarita, Trinidad and Tobago on the the upper Amazon basin to southern Peru
Atlantic slope; disjunct populations in and northern Bolivia on the Atlantic
versant and from central Costa Rica to Elapomorphus quinquelineatus

western Ecuador on the pacific slope’’ (Raddi, 1820)
(Savage 2002:663).
Vouchers for Guyana.—AMNH R- Type material.—The original name was
60783 from Kuyuwini Landing and Coluber 5-lineatus based on two specimens
AMNH R-60811 from Onora. from near Rio de Janeiro, Brazil. Accord-
Coloration in life.—The following is ing to Lema (1984:58), Hoge (1958) stated
from Starace (1998:185), based on speci- that MNHN 3673 is one of the types.
mens from French Guiana. ‘‘Colour varies Distribution.—‘‘Eastern and central Bra-
from light brown to green with a series of zil’’ (Peters & Orejas-Miranda 1970:105),
distinct [dark brown] diamond shaped and possibly the Guianas (see Voucher,
[markings]...with black edging distributed below).
along the dorsum. The belly is off-yellow Voucher for Guyana.—USNM 6180
from ‘‘Guiana’’ (no additional data).
with black lateral markings.’’
Coloration.—The following is based on
a preserved specimen (AMNH R-131801)
from Brazil. Dorsum light tan with five
Drymoluber dichrous (Peters, 1863b) darker tan to brown stripes (one being
Plate 31B vertebral); top of head brown; white patch
Type material.—The original name was on snout; supralabials tan; neck with wide
Herpetodryas dichrous and the syntypes, tan collar, the two halves not quite
one from Brazil, one from Surinam, are meeting at nape of neck; venter tan
ZMB 1661–2 (Bauer et al. 1995). anteriorly, unmarked, becoming lighter
posteriorly, yellow beneath tail (which is
Distribution.—Widespread in the Gui-
very short).
ana shield region (Avila-Pires 2005). Also
Comments.—USNM 6180 is the only
‘‘Colombia, Ecuador, eastern Peru, north-
specimen reported as being from the
ern Brazil, Amazonian Venezuela’’ (Peters
Guianas. It was received by the NMNH
& Orejas-Miranda 1970:100).
in 1863 as part of an exchange with the
‘‘Paris Museum,’’ so it is possible that the
18160 from Kartabo; USNM 549327 from
specimen was found in French Guiana.
East bank of Waruma River (05820 0 00 00 N,
However, Starace (1998) does not include
060846 0 00 00 W; 550 m); and USNM 566261
this species among the snakes known from
French Guiana. Considering the uncer-
tainties, the presence of this species in
based on the adult USNM specimen from
Guyana needs to be confirmed.
Berbice River Camp (CJC field notes,
1997). Greenish black above (but head
black); green laterally on neck; dark
blackish green ventrolaterally on body; Erythrolamprus aesculapii (Linnaeus, 1758)
lips, chin, and anterior throat white; other Plate 31C
ventral surfaces lemon yellow; iris mostly
deep chestnut brown; anteriorly on dor- Type material.—The original name was
sum very inconspicuous traces of dark Coluber Aesculapii and the two types are
brown bands; skin between scales gray. NRM 95 and 84 (http://linnaeus.nrm.se/
Starace (1998) points out that the young, zool/herp/madserp.html.en). There are no
being basically brown with numerous gray reliable locality data for the types.
to tan bands, are very different in color Distribution.—Peters & Orejas-Miranda
from the adults. (1970:111) stated that this inhabits ‘‘Ama-
zonian South America to central Brazil and Vouchers for Guyana.—AMNH R-8359
Bolivia.’’ It also has been described as from Kalacoon; AMNH R-18150, R-
occurring ‘‘from Trinidad southward to 36132, and R-36144 from Georgetown;
Argentina and Bolivia’’ (Murphy 1997:173). AMNH R-65556 from Kartabo; MCZ R-
Vouchers for Guyana.—AMNH R- 24939 from Mahaica Creek; MCZ R-
98185 from near Kartabo; AMNH R- 33360–33361 from Georgetown; USNM
137333 from Kartabo; AMNH R-140941 85077–85084 from Pomeroon; USNM
and USNM 566262 from Dubulay Ranch; 145458 from 80 km above Georgetown
ROM 20504 from Kurupukari; USNM on the Abary River; USNM 164205 from
55960 from Demerara (¼Georgetown); and Mabaruma Compound (08812 0 00 00 N,
USNM 84528 from Pomeroon. 059847 0 00 00 W); USNM 531698–531699
Coloration in life.—The following is from Iwokrama Forest Reserve, Three
based on the AMNH and USNM vouch- Mile Camp; and USNM 566263 from
ers from Dubulay Ranch (CJC field notes, Berbice River Camp.
1994). Body encircled with black, red, and Coloration in life.—The following is
white rings; white rings very narrow, based on the USNM specimen from
bordered on both sides by black; snout Berbice River Camp (CJC field notes,
tan; chin yellowish tan; throat orange; 1997). Dorsum grayish tan (but yellow
sixth infralabial each side white. On some ventrolaterally), with dark brown bands;
individuals, the ‘‘white’’ bands are actually head greenish brown; chin white; venter
gray above, cream below, and the sixth
yellow, becoming paler posteriorly; dirty
infralabial is not so distinctive, although
cream beneath tail; body bands black on
the lips are white, cream, or pale yellow.
ventral surface. Starace (1998) reported
Starace (1998) presented several color
that some specimens from French Guiana
photographs of specimens from French
have a bright red belly.
Guiana that showed a wide range of color
and pattern variation.
Comments.—USNM 566262 (see
Vouchers, above) was consuming an Oxy-
Hydrodynastes bicinctus (Hermann, 1804)
rhopus melanogenys (AMNH R-140898)
when captured. The anterior 9 cm of the Type material.—The original name was
prey was yet to be swallowed. Coluber bicinctus and the holotype and
type locality are unknown to us.
Distribution.—‘‘Guianas, Amazon re-
gion of Brazil, Colombia and Venezuela’’
Helicops angulatus (Linnaeus, 1758) (Peters & Orejas-Miranda 1970:127).
Plate 31D Voucher for Guyana.—AMNH R-60798
Type material.—The original name was from Onora Falls.
Coluber angulatus and Duellman (2005) Coloration in life.—On individuals from
stated that NRM 17 is a syntype. The same French Guiana, ‘‘the body varies from
specimen is cited on the following web site: light brown to yellow with 12 to 17
http://linnaeus.nrm.se/zool/herp/madserp. brown-black complete, but irregular
html.en. There are no reliable locality data rings. The head is yellow-beige [on the
associated with this. sides] with a longitudinal black band
Distribution.—‘‘Venezuela and Colom- behind each eye’’ (Starace 1998:201).
bia throughout South America to Bolivia; Starace’s photograph (1998:200) shows
Trinidad, Ecuador and Peru’’ (Peters & dark brown as the predominant color on
Orejas-Miranda 1970:123). top of the head.
Hydrops triangularis (Wagler, 1824) Imantodes cenchoa (Linnaeus, 1758)

Plate 31E
Elaps triangularis and holotype originally Type material.—The original name was
was in the ‘‘Musee royal de Munic.’’ The Coluber Cenchoa based on material from
type locality, in the vicinity of ‘‘Ega,’’ is America and the type locality is unknown
presently Tefé, at (03821 0 S, 64842 0 W), on to us.
the inner mouth of the canal...that con- Distribution.—‘‘. . .from southern Tam-
nects the lake of Tefé with the Solimões’’ aulipas (Atlantic) and Oaxaca (Pacific),
(Vanzolini 1981:xxvi). Mexico, south through Central America
Distribution.—‘‘From Venezuela, Gui- (exclusive of Pacific slope El Salvador,
anas and Trinidad to eastern Peru and Honduras, and northwestern Costa Rica)
northern Bolivia’’ (Peters & Orejas-Miran- to central Ecuador (Pacific) and Bolivia,
da 1970:129). Paraguay, and northeast Argentina (At-
Vouchers for Guyana.—AMNH R- lantic)’’ (Savage 2002:608). The species
25035, R-25053, and R-25056 from Ka- also occurs on Trinidad and Tobago
makusa; and AMNH R-98196, R-14134– (Murphy 1997:177).
14141, and R-67874 from Kartabo. Vouchers for Guyana.—AMNH R-67875
Coloration in life.—Beebe (1946:28) de- and R-81432 from Kartabo; AMNH R-
scribed a specimen from Kartabo as 152271 from Konawaruk Camp; ROM
follows. ‘‘Below, the ground color is pure
20502 from Kurupukari; ROM 42600 from
white, with two series of dark brown
Mount Wokomung; and USNM 566264
blotches close together, sometimes oppo-
site, sometimes alternate. These narrow
suddenly and extend upward as scale-
width vertical bands, not quite meeting
on the mid-back. The back is red brown, mens from Berbice River Camp and
while between each of the lateral, vertical Konawaruk Camp (CJC field notes, 1997,
lines is a conspicuous spot of bright coral 1998). Rusty tan with brown bands or
red, suffusing from three to five scales over spots; chin and throat white with tan
two rows. flecks; venter light tan with brown flecks;
‘‘The head has four irregular cross iris copper. Beebe (1946:29) noted for a
bands of black, with inter-spaces, counting specimen from Kartabo that the iris is
from the snout backward, of pale brown, ‘‘golden, with irregular indistinct fleckings
dark brown, dark red and bright red. The and streaks of orange and golden brown.
succeeding nuchal band of black almost Two narrow lines of light yellow along
meets above, and is the broadest on the both sides of the pupil, and a small patch
whole snake. . .the iris reddish. The general of dark brown at top and bottom.’’
impression of the lateral pattern is of
successive layers of red-brown, bright red
and white, cut by numerous vertical black Imantodes lentiferus (Cope, 1894)
bands.’’
Comments.—Albuquerque & Lema Type material.—The original name was
(2008) examined a large number of spec- Himantodes lentiferus and the holotype is
imens from widespread localities and ANSP 11459 (Duellman 2005) from Pebas,
concluded that the extensive variability of Peru (Peters & Orejas-Miranda 1970).
these snakes provides no justification for Distribution.—‘‘Amazonas, Brazil, Am-
recognizing subspecies, although six had azonian Colombia, Ecuador and Peru’’
been described in the past. (Peters & Orejas-Miranda 1970:135). The
species is also known from the Guianas Isheartun; AMNH R-137981–137984 and
(Starace 1998). USNM 566265–566266 from Karanambo;
Vouchers for Guyana.—USNM 535808 AMNH R-140902–140904 and R-141893
from Baramita. from Dubulay Ranch; USNM 531700
Coloration in life.—Starace (1998:284), from Iwokrama Forest Reserve, Burro
based on specimens from French Guiana, Burro River; and USNM 566267 from
noted that ‘‘the pattern closely resembles Ceiba Biological Station, Madewini River,
that of Imantodes cenchoa but is lighter ca. 5 km E (linear) Timehri Airport
(light beige to yellow).’’ Duellman (2005: (06829 0 57 00 N, 058813 0 06 00 W).
374) noted that on specimens from Peru Coloration in life.—The following is
there is ‘‘a Y-shaped mark on the top of based on AMNH and USNM specimens
the head, with the arms of the Y disjunct from Karanambo and Dubulay Ranch
from the base on the suture of the (CJC field notes, 1992, 1994, 1995). Dor-
parietals.’’ sum light brown with darker brown
Comments.—The USNM specimen blotches, but vertebral area tan; ventral
from Baramita (see Vouchers, above) was surfaces white anteriorly, becoming tan
found at night on a branch 1.5 m above posteriorly, with grayish brown speckling
ground in primary forest. beneath tail.
Comments.—Duellman (1958) consid-
ered most specimens from Guyana as
Leptodeira annulata (Linnaeus, 1758) being intergrades between L. a. annulata
Plate 31F and L. a. ashmeadi.
Coluber annulatus and the holotype is
‘‘number 9 in the Zoologicka Institutionen Leptophis ahaetulla (Linnaeus, 1758)
Uppsala’’ (Duellman 1958:47). The type Plate 32A
locality was restricted to the lower Rio
Amazon, Pará, Brazil (Duellman 1958).
Distribution.—‘‘The range of Leptodeira Coluber Ahaetulla and syntypes are NRM
annulata extends from southern Tamauli- ‘‘37, 40, 41?’’ (Duellman 2005), from
pas and Guerrero southward at low and America.
moderate elevations along both coasts of Distribution.—‘‘Lowlands and premon-
Mexico, excluding the wet forests of tane areas from central Veracruz and
southern Veracruz and the Yucatan Pen- Oaxaca, Mexico, on the Atlantic slope
insula, through the central plateaus and through Central and South America to
along the Pacific coast of Central America northern Argentina; on the Pacific versant
to Panama, thence through the interior from Costa Rica to southwestern Ecua-
valleys of Darien to northern Colombia, dor’’ (Savage 2002:670).
Venezuela, and the Guianas, including Isla Vouchers for Guyana.—AMNH R-
de Margarita, Tobago, and Trinidad, and 21281 from Kartabo; AMNH R-60792
throughout the Amazon Basin, southward from Karanambo; AMNH R-140899 from
into the Chaco of Paraguay and northern Dubulay Ranch; USNM 84518 and
Argentina, and along the Atlantic coast of USNM 85087 from Pomeroon; USNM
Brazil southward to Sao Paulo’’ (Duell- 141762 from 50 miles above Georgetown
man 1958:28). on the Abary River; USNM 164218 from
Vouchers for Guyana.—AMNH R- Kumaka (¼Takama); USNM 535809–
18155–18157 and R-67879–67880 from 535810 from Baramita; and USNM
Kartabo; AMNH R-60784–60787 from 566268 from Konawaruk Camp.
Coloration in life.—The following is disappear past midbody (bands may be

based on AMNH and USNM specimens nearly white laterally or on neck, may
from Dubulay Ranch and Konawaruk persist posteriorly after dorsal part is
Camp (CJC field notes, 1994, 1998). Head obscure); chin cream, grayish white, or
green above, with black stripe through eye; tan with dark gray to black spots; rest of
lips cream; neck largely green, the color venter (through anal plate) checkered with
becoming dorsolateral stripes separated by large squares of orange and black (often
yellowish tan vertebral area; green stripes irregular pattern; banded in places); mostly
becoming turquoise ca. midbody, green mottled black and white beneath tail,
again posteriorly; lateral stripe gold ante- where few pale orange spots may persist.
riorly, yellowish tan posteriorly; tail most- Beebe (1946) and Starace (1998) referred
ly tan; chin cream; rest of venter tan, to the colorful ventral areas as being red
becoming more-so gradually posteriorly; rather than orange.
iris yellow. Beebe (1946) and Starace
(1998) stated that anteriorly the ventral
surface is white. Liophis cobella (Linnaeus, 1758)
Comments.—The specimen from Kona-
waruk Camp (see Vouchers, above) was Type material.—The original name was
found while it was consuming a tree frog, Coluber Cobella based on material from
Dendropsophus minutus (AMNH A- America. Dixon (1983a) discussed the
166210). confusion involved with the Linnaean
specimens of this species.
Distribution.—‘‘. . .from the eastern
Liophis breviceps Cope, 1860a flanks of the Andes from near Villavicen-
Plate 32B cio, Colombia, southward to Buenavista,
Bolivia; from Caripito, Venezuela, south-
Type material.—The holotype, from eastward to Bahia, Brasil’’ (Dixon
Surinam, is ANSP 3697 (Malnate 1971). 1983a:155, on p. 158 Dixon mapped the
Distribution.—‘‘Eastern flanks of the species in Trinidad also).
Andes of Ecuador and Perú, eastward to Vouchers for Guyana.—AMNH R-
Obidos, Brasil; on the north from central 14263, R-18158, R-21336, and R-98204
Colombia and northern Guyana south to from Kartabo; AMNH R-36135–36136,
Rı́o Mamoré (Trinidad) Bolivia, and Posto R-36142–36143, and MCZ R-33355–33359
Diuarum Brasil’’ (Dixon 1989:9). from Georgetown; ROM 28372 from
Vouchers for Guyana.—AMNH R- Paramakatoi; and USNM 145455 from
18159 and R-98203 from Kartabo; 50 miles above Georgetown on the Abary
AMNH R-25052 from Kamakusa; River; and USNM 164198 from Maba-
AMNH R-140969–140970 and USNM ruma Compound (8812 0 00 00 N, 059847 0
566269 from Dubulay Ranch; AMNH R- 00 00 W). See Dixon (1983a) for many
152273 from Konawaruk Camp; and additional specimens.
BMNH 1976.1154 from Takama Village, Coloration in life.—The following is
Berbice River. See Dixon (1983a) for many from Starace (1998:215), based on speci-
additional specimens. mens from French Guiana. ‘‘Body varies
Coloration in life.—The following is from dark grey to olive-green, speckled
based on AMNH and USNM specimens with small black marks and fine yellow-
from Dubulay Ranch and Konawaruk beige spots. The belly is off-white to yellow
Camp (CJC field notes, 1994, 1998). Dark along which irregular dark markings form
brown to black above, with subtle tan or a chequered pattern.’’ Beebe (1946:35)
gray bands (not across belly) that fade and described specimens from Kartabo as
being brown or black above ‘‘with narrow Distribution.—‘‘Eastern South America,

white dorsal bands, scarlet below with from Guyana south to Buenos Aires,
broad bands of black; head and tail below Argentina, with scattered records in the
whitish, the latter with imperfect black Amazon Basin and Cerrado of Brasil’’
bands.’’ (Dixon 1989:17).
Comments.—Sometimes this species is Vouchers for Guyana.—AMNH R-8098
referred to as Liophis cobellus, but Darrel from Kalacoon; ROM 20502 from Kuru-
Frost confirmed (pers. comm.) that our pukari; and ROM 20505 from Tukeit.
current spelling of the specific epithet, See Dixon (1983b) for additional speci-
consistent with the original name and with mens.
some recent usage (e.g., Gorzula & Señaris Coloration in life.—The following is
1999), is reasonable. from Starace (1998:217), based on speci-
mens from French Guiana. ‘‘The body is
bottle-green to black, splashed with faint
Liophis lineatus (Linnaeus, 1758) yellow spots. The head is dark and reveals
a small number of off-white fine spots. The
Type material.—The original name was belly is bright yellow.’’
Coluber lineatus and the type material is
unknown to us. The type locality informa-
tion is not reliable.
Distribution.—‘‘Central Panama east- Liophis poecilogyrus (Wied, 1824)
ward through Colombia, Venezuela, Type material.—The original name was
Guyana, Suriname, French Guiana, to Coluber poecilogyrus and there are two
the mouth of the Amazon River in Brazil’’
female syntypes, AMNH R-3593–3594
(Michaud & Dixon 1987:3).
(Dixon & Markezich 1992), from Barra
de Joucoú, assez près de la rivière Espirito
14265 and R-98206 from Kartabo;
santo [Brazil].
AMNH R-2664 and R-137647 from
Distribution.—Eastern Venezuela east-
Georgetown; AMNH R-60789 from Para-
bam; and AMNH R-60837–60838 from ward through the Guianas, south through
Isheartun. the Amazon Basin to Bolivia, Argentina,
Coloration in life.—The following is and Uruguay (Dixon & Markezich 1992).
from Starace (1998:225), based on speci- Vouchers for Guyana.—AMNH R-
mens from French Guiana. Light ‘‘brown 60803 from Kuyuwini, identified by J. R.
to grey body with three characteristic dark Dixon; USNM 146376 from Lethem; and
brown lines running along the length of the USNM 497805 from Karanambo.
dorsum. The two lateral lines are more Coloration in life.—Dixon & Markezich
distinct on the upper part [meaning ante- (1992:153–154) described extensive varia-
rior] of the body.’’ tion in coloration of this species, but
without specifying whether the notes were
based on specimens in life. Many speci-
mens from the northern part of the range
Liophis miliaris (Linnaeus, 1758)
had a gray dorsum with or without dark
Type material.—The original name was brown or black markings. The ground
Coluber miliaris and the holotype is ‘‘N.R. color of the sides of the body was similar
[Naturhistoriska Riksmuseum, Stockholm, to that of the middorsal region or lighter.
Sweden] No. 56’’ (Gans 1964:39). Dixon Middorsal scales were with or without
(1983b) restricted the type locality to white edges. Mottling of the venter was
Surinam. poorly defined.
Liophis reginae (Linnaeus, 1758) Vouchers for Guyana.—AMNH R-

Plate 32C 21335 and R-98197–98198 from Kartabo;
AMNH R-36106 from Lama Creek, De-
Type material.—The original name was merara River; ROM 20506 from Tukeit;
Coluber Reginae. Dixon (1983c) discussed ROM 22838 from Baramita; ROM 28375
issues involved with designation of a from Paramakatoi; USNM 291151 from
lectotype at the Museum Regis Adolphi along trail from Kato to Paramakatoi; and
Friderici and he restricted the type locality USNM 566270 from Dubulay Ranch.
to Suriname. Coloration in life.—The following is
Distribution.—‘‘Cis-Andean South Amer- based on the USNM specimen from Dubu-
ica, from Colombia to northern Argentina; lay Ranch (CJC field notes, 1995). Body
also Trinidad and Tobago’’ (Dixon 1989:21). green above; subtle black markings visible
Vouchers for Guyana.—AMNH R-8095 at times on sides; lips greenish yellow; chin
from Kalacoon; AMNH R-17680 from and throat yellow, then venter gradually
Georgetown; AMNH R-18170, R-67876, becoming pale orange, then yellow again
and R-137348 from Kartabo; ROM 39441 beneath tail. Starace (1998) stated that the
from Mt. Ayanganna; USNM 164207– dorsal color may be green, brown, or gray-
164210 from Mabaruma Compound blue, and there may be orange or blue
(8812 0 00 00 N, 0598470 00 00 W); USNM 549328– speckling, especially on the neck.
549329 from N slope of Mt. Roraima
(5817 0 00 00 N, 60845 0 00 00 W); USNM 561439
from Washikura River (8815 0 00 00 N, 059844 0 Mastigodryas boddaerti (Sentzen, 1796)
00 00 W); and USNM 573507 from Woko- Plate 32E
mung Massif (5800 0 08 00 N, 059852 0 47 00 W).
Coloration in life.—The following is from Type material.—The original name was
Starace (1998:219), based on specimens Coluber Boddaerti. Location of the type
specimen is unknown. Stuart (1941) and
from French Guiana. ‘‘The body varies
Merrem (1820) suggested that Surinam
from brown to olive green and is spotted
was the type locality.
with yellow scales. A dark brown line passes
Distribution.—‘‘Tropical south America,
over the supralabials. Just behind the eyes
from Colombia and Venezuela to Bolivia
run two light green triangular patterns, the and western Brazil’’ (Peters & Orejas-
apex pointing towards the neck. The belly is Miranda 1970:192).
yellow with sundry [more-or-less rectangu- Vouchers for Guyana.—AMNH R-8092
lar] black markings.’’ from Georgetown; AMNH R-44920, R-
14252, and R-18166–18167 from Kartabo;
AMNH R-60791 from Karanambo;
Liophis typhlus (Linnaeus, 1758) AMNH R-60840 from Isheartun; AMNH
Plate 32D R-60852 from Kuyuwini River; AMNH
R-152277 and USNM 566271 from Ber-
Type material.—The original name was bice River Camp; ROM 22840–22841 from
Coluber Typhlus and the syntypes are in Baramita, the last two identified by H.
the ‘‘Royal Museum, Stockholm’’ (Dixon Zaher; and USNM 164199, 164206, and
1987:173). There are no reliable locality 164211 from Mabaruma Compound
data for them. (8812 0 00 00 N, 059847 0 00 00 W).
Distribution.—‘‘Rainforests of the Gui- Coloration in life.—The following is
ana Shield and Amazon Basin, also the based on the AMNH and USNM speci-
Chaco and Cerrado of Bolivia, Brasil, and mens from Berbice River Camp (CJC field
Paraguay’’ (Dixon 1989:23). notes, 1997). This species has ontogenetic
change in coloration. The juveniles from 2645/0, female (Franzen & Glaw
Berbice River Camp (see Vouchers, above) 2007:250). The type locality is the area of
were described in life as follows. Dorsum Ega, now known as Tefé, and ‘‘the locality
brown with grayish tan bands; white spots (03821 0 S, 64842 0 W) is on the inner mouth
ventrolaterally on ends of tan bands of the canal. . ., a few kilometers long, that
anteriorly on body; chin and throat white connects the lake of Tefé with the Sol-
with dark brown irregular spots; venter imões’’ (Vanzolini 1981:xxvi).
tan. A larger individual from Berbice Distribution.—‘‘Lowland and premon-
River Camp was described as follows. tane zones from Arizona and Coahuila,
Nearly uniform brown dorsally, with Mexico, south to northern Peru on the
traces of bands anteriorly; a lateral light Pacific versant and Bolivia east of the
tan stripe on anterior half of body; venter Andes; also on the islands of Trinidad and
light gray with darker gray smudges on Tobago’’ (Savage 2002:677). The species
throat. also extends on the eastern part of the
range to southeastern Brazil (Gorzula &
Señaris 1999 [1998]).
Ninia hudsoni Parker, 1940 Vouchers for Guyana.—AMNH R-8170
from Kalacoon; AMNH R-14260, R-
Type material.—The holotype is BMNH 18179–18180, R-98207–98209, and R-
1946.1.15.38, a male from the New River, 137326–137329 from Kartabo; AMNH
Guyana, within an area that is in dispute R-60758–60766 and R-60770–60776 from
with Suriname. Isheartun; AMNH R-60815 from Kuyu-
Distribution.—‘‘British Guiana; Amazo- wini Landing; AMNH R-60839 from
nian Ecuador’’ (Peters & Orejas-Miranda Wichabai; AMNH R-148587 from Iwo-
1970:222). Duellman (1978) described a krama, Muri Scrub Camp; ROM 28374
specimen from Amazonian Ecuador and from Paramakatoi; USNM 497820 from
later (Duellman 2005:145) reported one Pirara Ranch, ca. 25 mi N of Lethem
from Amazonian Peru. (3836 0 42 00 N, 059840 0 30 00 W); and USNM
Vouchers for Guyana.—The holotype 531701 from Iwokrama Forest Reserve,
and the paratype (BMNH 1946.1.15.20), Three Mile Camp (4837 0 59 00 N, 058842 0
both from the type locality, are the only 52 00 W).
specimens known from Guyana. Coloration in life.—Beebe (1946:36) de-
Coloration in life.—The following is scribed AMNH R-18179 from Kartabo, in
from Duellman (1978:250) based on a life, as follows. ‘‘Top of head sepia. An
specimen from Ecuador. The individual indefinite streak along side of head which
‘‘has a dark gray dorsum with a pale gray deepens to a narrow band of black just
nape band and a creamy white venter.’’ above the upper labials, and continues
The following is from Parker (1940:270), across the iris in two areas of dark green.
but might be based on the preserved The second to fifth upper labials with
specimens. ‘‘Very dark purple-brown narrow shading of black along upper edge.
above, with a white nuchal collar. . .; lower A streak of pale vinaceous buff extends
surfaces white except the posterior half of across the top of the preocular, tinging the
the tail, which is brown-dusted.’’ lower portion of the supra ocular and
continuing in a horizontal line a little back
of eye, where it deepens to umber. Upper
Oxybelis aeneus (Wagler, 1824) labials pale yellow, shading downward to
bluish-white. Throat bluish-white with
Type material.—The original name was faint tinge of pink. Body above light ashy,
Dryinus aeneus and the lectotype is ZSM ventral surface dark vinaceous brown. A
narrow rim of picric yellow extends on living individuals from Kartabo that
around the lower four-fifths of the pupil, when spreading the scales apart anteriorly
widening toward the top and shading to on the body, one sees broad bands of
silver white, which extends over all the orange and pale yellow on the skin
upper part of the iris. Large irregular between the scales.
patches of parrot green lie on either side
of pupil flecked with black. The lower part
of iris is silvery white flecked with maize Oxyrhopus melanogenys (Tschudi, 1845)
yellow.’’ Starace (1998:292) added that Plate 33A, B
‘‘the body is covered with very fine dark
coloured patterns [markings].’’ Type material.—The original name was
Sphenocephalus melanogenys and the holo-
type is ‘‘MHNN 14 (Cat. 99)’’ (Duellman
Oxybelis fulgidus (Daudin, 1803a) 2005:379). The type locality is the Chan-
Plate 32F chamayo region, Peru (Peters & Orejas-
Miranda 1970:232).
Type material.—The original name was Distribution.—All records that we have
Coluber fulgidus and the type material is seen for this species are from the Guianas
unknown to us. The type locality was and Amazon Basin.
restricted to Chichen Itzá, Yucatán, Mex- Vouchers for Guyana.—AMNH R-8094
ico by Smith & Taylor (1950a). from Kalacoon; AMNH R-98211 from
Distribution.—‘‘Lowlands and adjacent Kartabo; AMNH R-140898 from Dubulay
slopes on the Pacific versant from the Ranch; AMNH R-152279–152280 and
Isthmus of Tehuantepec to eastern Pana- USNM 566273 from Berbice River Camp;
ma and on the Atlantic versant in southern IRSNB 17912 and 17940–17941 from
Veracruz, Mexico, and then south from Kaieteur National Park; and ROM 39439
the Yucatán Peninsula through Central from Mt. Ayanganna. Additional details
America and throughout South America, and localities are listed by MacCulloch et
east of the Andes to northern Bolivia and al. (2009:495).
extreme northeastern Argentina’’ (Savage Coloration in life.—The following is
2002:679). based on AMNH and USNM specimens
Vouchers for Guyana.—AMNH R- from Dubulay Ranch and Berbice River
14259, R-16049, R-65553–65554, and R- Camp (CJC field notes, 1994, 1997). One
137325 from Kartabo; AMNH R-25049 specimen had the dorsum with alternating
from Kamakusa; AMNH R-60847 from black (broadest), white (narrowest), and red
Isheartun; JC 8039 (AMNH field number) bands; usually two white bands within a
from Konawaruk Camp (given to UG); black one (triad pattern); red bands with
ROM 20501 from Kurupukari; USNM black dots; bands not crossing belly; venter
84527 from McKenzie (¼Linden); and white (but gray on chin and throat), with
USNM 566272 from Berbice River Camp. gray marks beneath tail; top of head black;
Coloration in life.—The following is iris orange. Colors on two other individuals
based on UG and USNM specimens from were similar, but with differences in the
Berbice River Camp and Konawaruk pattern. Anteriorly, these appeared to be
Camp (CJC field notes, 1997, 1998). Leaf predominantly black snakes with narrow
green above, with greenish yellow to white white bands, but the middle of the broad
ventrolateral stripe; lips, chin, throat, and black areas had a red triangle ventrolater-
belly chartreuse; iris copper, black, tan on ally, and those posteriorly on the body
one specimen, brown, gold, and pale extended upward to become red bands with
copper on another. Beebe (1946:37) noted black spots. On the posterior belly there
were red and black mixed with the light bands equal to the length of 2–4 dorsal
coloration. Another specimen was noted as scales. Bands are usually symmetrical, but in
being similar to the last two, but with a one specimen there is a half-band, on one
conspicuous red band on the neck; without side only, between otherwise symmetrical
a triad pattern; and with the mix of red and bands. . .Underside of head yellowish
black on the venter only on the posterior orange. Venter white. . .Underside of tail
25% of the body but more conspicuous banded orange and white, corresponding to
beneath the tail. MacCulloch et al. dorsal pale orange bands and red ground
(2009:491) gave more details and pointed respectively. . .Bands are prominent in juve-
out that older adults may be melanistic. niles but [usually] become inconspicuous in
Comments.—When found on the forest adults’’ (MacCulloch et al. 2009:489). Star-
floor at 1530 h, AMNH R-140898 (see ace (1998:302) stated that ‘‘a dark band
Vouchers, above) was being consumed by covering the [anterior]...half of the parietals
another snake, an Erythrolamprus aescula- crosses over the eyes.’’
pii (AMNH R-140942), tail first, with Comments.—Formerly, snakes of this
about 9 cm protruding from its mouth. coloration and pattern were considered to
In the recent past (see Zaher & Caramaschi be color pattern variants of a different
1992), Guyanan specimens of this species species, Oxyrhopus formosus, which is a
were misidentified as Oxyrhopus trigemi- valid species also but not known from
nus, a species not known to occur in Guyana (Hoge et al. 1973, Jorge da Silva
Guyana (MacCulloch et al. 2009). 1993, MacCulloch et al. 2009).
Oxyrhopus occipitalis (Wagler, 1824)

Plate 33C Oxyrhopus petolarius (Linnaeus, 1758)
Plate 33D
Natrix occipitalis and the lectotype is ZSM Type material.—The original name was
2053/0, a male (Franzen & Glaw 2007: Coluber Petolarius. The type material is
256), from the banks of the Solimöens unknown to us and the type locality lacks
River, Brazil. reliable data. Savage (2011; see Comments
Distribution.—This species occurs in below) discussed why this specific epithet is
Amazonas and Amapá, Brazil, Amazonas preferred over O. petola, which was used
and Bolivar, Venezuela, Surinam, French previously.
Guiana, and Guyana (MacCulloch et al. Distribution.—‘‘Tropical lowlands
2009:490). from Veracruz, Mexico, on the Atlantic
Vouchers for Guyana.—AMNH R-36102 versant and Costa Rica on the Pacific
from Lama Creek, Demerara River; slope through Central America to west-
IRSNB 17913 and ROM 42995 from ern Ecuador and throughout northern
Kaieteur National Park; and SMNS 12111 South America and the Amazon basin to
from Mabura Hill (5809 0 N, 588420 W). Brazil, Peru, and Bolivia’’ (Savage
Coloration in life.—‘‘Dorsal ground col- 2002:576).
our red or reddish orange; snout and Vouchers for Guyana.—AMNH R-
supralabials yellow; top of head medium 18164 from Maripa; AMNH R-36099
to dark brown. A white anterior body band and R-36524 from Lama Creek; AMNH
(sometimes obscure), 3–4 dorsal scales in 36140 from Georgetown; AMNH R-98210
width, approximately 2 cm behind the head. from Kartabo; ROM 22843 (presently
Faint, pale orange body bands with very CSBD HR 89 at UG) and USNM
narrow dark borders; width of pale orange 535811 from Baramita; USNM 164204
from Mabaruma Compound (8812 0 00 00 N, (very thin) black line below eye, from
059847 0 00 00 W); and USNM 566274 from midway between nostril and eye to tempo-
Dubulay Ranch. ral area, along upper sutures of supra-
Coloration in life.—The following is labials.
Dorsum with alternating bands (but not Philodryas viridissimus
across belly) of black and red, the latter (Linnaeus, 1758)
with black dots; top of head black;
ventral surfaces white (gray near tip of Type material.—The original name was
tail). Beebe (1946:38) described living Coluber viridissimus based on material
specimens from Kartabo as having the from Surinam, but the type(s) are un-
light bands anteriorly on the body white, known to us.
transitioning through pink to red poste- Distribution.—‘‘Amazonas and Para-
riorly. Additional details were presented guay valleys, from southern Venezuela
by MacCulloch et al. (2009:493). and Guianas to Argentina’’ (Peters &
Comments.—Savage (2011) explained Orejas-Miranda 1970:245).
his conclusion that the appropriate name Vouchers for Guyana.—AMNH R-
is Oxyrhopus petolarius for snakes that 60846 from Marudi; AMNH R-61540
previously were called O. petola. The from Matali; AMNH R-98212 from Kar-
confusion stems from the fact that tabo; and USNM 84525 and 85088 from
Linnaeus (1758) named and described McKenzie (¼Linden).
both on the same page as if they were Coloration in life.—Beebe (1946:38) de-
different species. Subsequent authors syn- scribed one of two specimens from Karta-
onymized the two using one or the other bo as follows. ‘‘General body color lettuce
name, and which name applies today green above, shading laterally to greenish-
hinges on who was determined to be the yellow on ventral scales. Top of head
first reviser. parrot green, two black spots on scales in
front of eye and three behind, forming a
broken facial band. Upper labials emerald
Philodryas olfersii (Lichtenstein, 1823) green, shading posteriorly to light yellow
Plate 33E green. Lower labials, chin and throat pale
greenish-gray touched with pale cinnamon
Type material.—The original name was pink. Scales back of gape, along lateral
Coluber Olfersii and the type specimen, neck and forming anterior border of the
from Brazil, is unknown to us. first fifty ventrals. . .a most delicate blue.
Distribution.—‘‘Western Brazil and east- This color occurs sporadically on other
ern Peru through Bolivia and Paraguay to ventral scales and on all the caudal
Uruguay and Argentina’’ (Peters & Orejas- ventrals [¼ subcaudals]. . .Iris a tangled
Miranda 1970:244); also known from the mesh of dark cinnamon brown with a
Guianas (e.g., Starace 1998). sparse but conspicuous flecking of gold
Vouchers for Guyana.—USNM 566275 dots across the upper portion.’’
from Aishalton; and AMNH R-140939–
140940 from Dubulay Ranch.
Coloration in life.—The following is Phimophis guianensis (Troschel, 1848)
based on AMNH and USNM specimens Plate 34A
from Aishalton and Dubulay Ranch (CJC
field notes, 1993, 1994). Dark leaf green Type material.—The original name was
above; pale yellowish green below; fine Heterodon guianensis based on a specimen
from the immediate vicinity of Pirara, neuwiedii (see below), and in the field the
Guyana. Location of the type is unknown vouchers (see above) from Berbice River
to us. Camp were misidentified as such and no
Distribution.—‘‘Cocle Province, Pana- particular color differences were noted.
ma, to Surinam, in savannah and scrub However, color photographs were taken,
areas’’ (Peters & Orejas-Miranda 1970: and the following is based on them.
246); also known from French Guiana Dorsum brownish red; top and sides of
(Starace 1998). head black; conspicuous white collar on
Vouchers for Guyana.—AMNH R- posterior head and neck, followed on
140943 from Ceiba Biological Station, neck by broad black area, soon fading on
Madewini River, ca. 5 km (linear) east of anterior body; venter light. Beebe
Timehri Airport (06829 0 57 00 N, 058813 0 (1946:39) described a specimen from
06 00 W); and AMNH R-140944–140946, Kartabo as having the light band around
R-141896, and USNM 497806 and the neck ‘‘yellowish-white’’ and having
566276–566277 from Dubulay Ranch. ‘‘ventrals yellowish-white, tail grayish.
Coloration in life.—The following is Iris dark,’’ but no voucher specimen
based on the AMNH and USNM speci- from Kartabo occurs at the AMNH.
mens from Ceiba Biological Station and Starace (1998:323) stated that not all
Dubulay Ranch (CJC field notes, 1994, individuals have the ‘‘white-yellow’’ col-
1995). Pale yellowish tan above (light lar on the neck and that ‘‘the belly is off-
orange in a juvenile) with dark brown to white.’’
black speckling, sometimes extensive;
snout often tan, but most of head brown;
dark brown to black band on back of head Pseudoboa neuwiedii
to neck (but not across ventral aspect); (Duméril, Bibron, & Duméril, 1854)
venter translucent pinkish white or cream, Plate 34C, D
becoming cream posteriorly, pale yellow
out on tail. A juvenile had the black band Type material.—The original name was
on the back of the head and neck preceded Scytale Neuwiedii based on several speci-
by a contrasting white band. mens. The types are unknown to us, but
the type locality was restricted to Cumaná,
Venezuela (see Peters & Orejas-Miranda
Pseudoboa coronata Schneider, 1801 1970:254).
Plate 34B Distribution.—‘‘Pacific Panama from
just west of Canal Zone through northern
Type material.—The type specimen was and interior Colombia to Surinam; south
in the ‘‘Musei Heyeri’’ and no locality data to Brazil, along Amazon River; Trinidad
were given. and Tobago Islands’’ (Peters & Orejas-
Distribution.—‘‘Guianas, Amazonian Miranda 1970:254).
watershed in Brazil, Colombia, Ecuador, Vouchers for Guyana.—AMNH R-
Peru and Bolivia’’ (Peters & Orejas-Mi- 139737 from Aishalton; and AMNH R-
randa 1970:253). 140900–140901, R-141897, and USNM
Vouchers for Guyana.—AMNH R- 566279 from Dubulay Ranch.
152283 and USNM 566278 from Berbice Coloration in life.—The following is
River Camp; USNM 56167–56168 from based on the AMNH and USNM speci-
‘‘Guyane’’ (¼Guyana, with no additional mens from Dubulay Ranch (CJC field
locality data). notes, 1994, 1995). In young individuals,
Coloration in life.—Appearance of this top of head black; pale yellow band on
species is similar to that of Pseudoboa neck and back of head, present or not,
followed by a black band on neck; rest of Vouchers for Guyana.—AMNH R-

dorsum orangish red; ventral surfaces 21337 from Kartabo; AMNH R-60848
translucent white, but tail very pale orang- from the upper Rupununi River (collected
ish red below, perhaps distally. Adults are by R. Snedigar); AMNH R-152285,
brown above, greenish yellow below. AMNH field number JC 8057 (now at
UG), and USNM 566280 from Konawa-
ruk Camp; AMNH R-152286 from about
Pseudoeryx plicatilis (Linnaeus, 1758) 40 km (linear) southeast of Linden on
Arakabusa Creek; USNM 85086 from
Type material.—The original name was Pomeroon; USNM 300656 from Bartica;
Coluber plicatilis and the holotype, from and USNM 326121 from Kwakwani
Ternataeis, is NRM 57 (Duellman (5830 0 00 00 N, 058800 0 00 00 W).
2005:382). Coloration in life.—The following is
Distribution.—Colombia and Guianas based on the AMNH, UG, and USNM
to Bolivia, Paraguay and northern Argen- vouchers (see above) from Konawaruk
tina (Peters & Orejas-Miranda 1970:255). Camp (CJC field notes, 1998). On young
Vouchers for Guyana.—ROM 5270 from snakes, dorsal ground color gray or tannish
‘‘Guyana’’ (no additional data); and gray; vertebral area light tan; crossbands
USNM 85075–85076 from Pomeroon. brown (gray or tannish gray within); venter
Coloration in life.—The following is mostly gray or grayish tan, anteriorly with
from Starace (1998:234), based on speci- cream showing through extensive gray
mens from French Guiana. ‘‘The dorsum flecking; lips mostly cream; cream lines on
is [dark]...brown along which run two dark snout, nostril to lip; iris tan or pale copper.
[yellow] longitudinal lines from the head A larger adult from Arakabusa Creek,
along the sides...The belly is beige spotted provided by Alexander Mendes (AMNH
with fine black marks on the sides.’’ 152286; see Vouchers), was noted in the
Starace’s photo (p. 233) also shows a same field notes as follows: brownish green
broad black stripe on the side of the head above; lips, chin, throat, and anterior venter
(also mentioned on his p. 234), which orangish yellow; posterior venter greenish
continues as a broad black stripe on the brown; iris brown.
side of the body. Beebe (1946:39) reported seeing no
fewer than 24 individuals at Kartabo.
Adults were ‘‘monochrome above and
Pseustes poecilonotus (Günther, 1858b) below in two phases, one a deep olive
Plate 34E green with lemon yellow lower head, chin
and throat, and the other a rich red
Type material.—The original name was brown with yellowish-orange anterior
Spilotes poecilonotus and the holotype, lower parts. In the young. . .the variation
from Honduras, is BMNH 1946.1.7.39 is still more extreme, the basic colors
(Colin McCarthy, pers. comm.). being red brown, or gray or olive above,
Distribution.—‘‘Lowlands and premon- banded with darker shades of the ground
tane slopes of the Atlantic versant from color, with face and throat black-marked
San Luis Potosı́, Mexico to Amazonian white or green-marked yellow. The iris
Peru, Bolivia, and Brazil; also on the is. . .mottled silvery, with dark pigmented
Pacific versant in Oaxaca, Mexico, central areas fore and aft, which are crossed with
and southern Costa Rica, Panama, and about three, distinct, white, vein-like
western Colombia and Ecuador’’ (Savage lines, radiating from a narrow area on
2002:680). the anterior and posterior equator of the
pupil, extending out to the external rim Rhinobothryum lentiginosum

of the iris.’’ (Scopoli, 1788)
Plate 34F
Pseustes sulphureus (Wagler, 1824)
Coluber lentiginosus. The holotype and
Type material.—The original name was type locality are not known to us.
Natrix sulphureus and the lectotype is ZSM Distribution.—‘‘Basins of Rı́os Amazon
1681/0, a male (Franzen & Glaw and Paraguay in tropical South America’’
2007:257). The type locality was along (Peters & Orejas-Miranda 1970:269).
the Japura River, Brazil. According to Vouchers for Guyana.—USNM 535812
Vanzolini (1981:xxvi), ‘‘Martius followed from Baramita.
the Japurá from its mouth (03808 0 S, Coloration in life.—The following is
64866 0 W), to the falls of Araracuara based on Starace’s (1998) description of
(00824 0 S, 72817 0 W), then in disputed terri- specimens from French Guiana, and one
tory, now in Colombia.’’ (AMNH 119433) collected and photo-
Distribution.—‘‘Peru; Ecuador; Brazil; graphed by CJC and CRT in Suriname,
Guianas; Trinidad’’ (Peters & Orejas-Mi- in 1980. The dorsum is a series of colorful
randa 1970:259). rings of black (broadest), white (narrow-
Vouchers for Guyana.—AMNH R-8349 est) and red (containing black spots).
from Kalacoon; AMNH R-60817 from There is a narrow white ring on the neck,
Kuyuwini River; AMNH R-137349 from after which the sequence of rings is as
Kartabo, the last identified by R. W. follows, repeated along the body: black,
McDiarmid; and UMMZ 53928 from white, red, white, etc. The venter is dark,
Dunoon, Demerara River. except in areas of light rings. ‘‘The red [or
Coloration in life.—Beebe (1946:41) de- light-colored] edging of each cephalic scale
scribed specimens from Kartabo as follows. is a characteristic feature of this species’’
‘‘While the dorsal coloring may be mono- (Starace 1998:328), which otherwise basi-
chrome, olive, brown or yellow, this is not cally has a dark head.
an adult pattern but may occur in an
Comments.—In the past, two neotropi-
individual only one-third grown. The usual
cal species were recognized in Rhinobo-
pattern shows a series of conspicuous
thryum, the second of which was R.
dorsal cross markings, crescents or bands,
bovallii, which today is recognized as a
single or double, or even more intricate
junior synonym of R. lentiginosum. Con-
figures. The general tendency is a gradual
change from yellow background with black sequently, the genus is monotypic.
markings on the head and anterior body, to
black with corresponding yellow markings
posteriorly. . .The iris has a remarkably Sibon nebulatus (Linnaeus, 1758)
consistent pattern, the background brown-
black, with a fine mist of paler brown lines Type material.—The original name was
radiating outward from the bright silver Coluber nebulatus. According to Peters
pupil ring to the outermost borders of the (1960:199), the type is ‘‘one specimen in
visible eyeball. It thus differs radically from the Museum Regis Adolphi Friderici,
the iris of the congeneric species.’’ Starace Stockholm.’’ The website of Museum
(1998:243) stated that the belly is yellow Adolphi Friderici lists NRM numbers 60,
anteriorly, black posteriorly, and his photos 64, and 65 for this species at http://
(on his pp. 240–241) show distinct black linnaeus.nrm.se/zool/herp/madserp.html.
bars on the upper lips. en. The type locality is America.
Distribution.—‘‘Northern South Amer- specimens from French Guiana. Body

ica. . .lowland Central America to Mi- dark gray with numerous narrow yellow
c h o a c á n . . .a n d V e r a c r u z , M e x i c o , bands; irregular row of red scales along
including Yucatán Peninsula’’ (Peters back, often extending onto sides; head
1960:199). scales dark with light borders; belly light
Vouchers for Guyana.—AMNH R- with dark spots.
36092 from Lama Creek, Demerara River;
AMNH R-137340 from Kartabo (this one
identified by R. W. McDiarmid); UMMZ Siphlophis compressus (Daudin, 1803a)
56463 from ‘‘Maruca River St Rosa’’ Plate 35A
(quoted from UMMZ records); ROM
28373 from Paramakatoi; USNM 84524 Type material.—The original name was
from McKenzie (¼Linden); and USNM Coluber compressus based on material (not
164202 from Mabaruma Compound known to us) from Surinam.
(8812 0 00 00 N, 059847 0 00 00 W). Distribution.—‘‘Discontinuous. Coastal
Coloration in life.—The following is strip of Brazil from Rio de Janeiro (city) to
from Starace (1998:179), based on speci- Sergipe. Mouth of Amazon and central
mens from French Guiana. ‘‘Closely Bolivia to Trinidad and Panama’’ (Peters
resembles Dipsas variegata. The body & Orejas-Miranda 1970:311).
colour varies from grey to brown and is Vouchers for Guyana.—AMNH R-
covered with dark brown irregular bands, 18151 from Kartabo; AMNH R-25057
ring-like. Each band is edged by fine, from Kamakusa; AMNH field number
irregular, beige spots. [Top of head brown JC 7894 from Konawaruk Camp (given
with darker brown markings.] The belly to UG); USNM 531702 from Iwokrama
ranges from white to beige, speckled with Forest Reserve (4820 0 15 00 N, 058849 0 37 00 W);
dark brown miniscule points [; darker USNM 535813 from Baramita; and
under tail].’’ USNM 566283 from Dubulay Ranch.
Comments.—Often the spelling of the Coloration in life.—The following is
specific epithet for this species is nebulata, based on the UG and USNM specimens
but recent authors (Harvey 2009, Zaher et from Dubulay Ranch and Konawaruk
al. 2009) have used nebulatus consistent Camp (CJC field notes, 1994, 1998). Dusty
with Linnaeus. pinkish red above, with black bands and
spots; head orange; neck black; chin area
white with orangish tinge on lips; rest of
Siphlophis cervinus (Laurenti, 1768) venter cream, with pink tinge posteriorly,
but gray distally beneath tail; iris red.
Type material.—The original name was Comments.—Until recently this species
Coronella cervina based on material (un- was considered to represent the genus
known to us) from America. Tripanurgos, which was monotypic (Zaher
Distribution.—‘‘Central Bolivia and & Prudente 1999).
Maranhão [northward] to the Canal Zone
and Trinidad’’ (Peters & Orejas-Miranda
1970:281). Spilotes pullatus (Linnaeus, 1758)
Vouchers for Guyana.—UMMZ 53967
from ‘‘Dunoon, Demerara River’’ (quoted Type material.—The original name was
from the UMMZ records). Coluber pullatas and the holotype is listed
Coloration in life.—The following is as ‘‘NRM cat no 30’’ on the website of
adapted from the description and photo- Museum Adolphi Friderici at http://
graph in Starace (1998:331), based on linnaeus.nrm.se/zool/herp/madserp.html.
en. There are no reliable data for the type actually based on two other species,
locality. Tantilla armillata and Tantilla ruficeps.
Distribution.—‘‘Atlantic lowlands and Vouchers for Guyana.—AMNH R-
premontane slopes from Tamaulipas, 15147 and R-98216–98217 from Kartabo;
Mexico, south through Central America AMNH R-139739 from Aishalton;
and northern South America to Bolivia, AMNH R-140950–140951 from Dubulay
Paraguay, and extreme northeastern Ar- Ranch; ROM 22839 from Baramita;
gentina; on the Pacific versant from the USNM 164197 from Mabaruma Com-
Isthmus of Tehuantepec to western Ecua- pound (8812 0 00 00 N, 059847 0 00 00 W); and
dor’’ (Savage 2002:687). USNM 566281 from Berbice River Camp.
Vouchers for Guyana.—AMNH R- Coloration in life.—The following is
60804 from Parabam; AMNH R-60824 based on AMNH and USNM specimens
from Wichabai; USNM 84522 from from Aishalton, Dubulay Ranch, and Ber-
McKenzie (¼Linden); USNM 145460 from bice River Camp (CJC field notes, 1992,
50 miles above Georgetown on the Abary 1994, 1997). Dorsum reddish brown; lower
River; USNM 164214 from Mabaruma sides orangish brown; head and neck black,
Compound (8812 0 00 00 N, 059847 0 00 00 W); with tan spot on snout and tan parietal
and USNM 291152 found on trail from spots; white markings on face (side of head);
Kato to Paramakatoi. chin white; white or cream beneath throat
Coloration in life.—The following is and neck; rest of ventral surfaces pale
from Starace (1998:250), based on speci- yellow. Starace (1998:334) added that there
mens from French Guiana. ‘‘The body is are ‘‘three to five black longitudinal lines
shiny black, decorated with irregular running down the body of which the median
yellow bands issuing from the belly which is the most strongly marked.’’
is also yellow. [Labials yellow and black.]
The tail is often black. . .[without] the
yellow markings.’’ Some individuals are Thamnodynastes pallidus (Linnaeus, 1758)
melanistic.
Coluber pallidus and the neotype is ZMUU
Tantilla melanocephala (Linnaeus, 1758) 292, an adult male (Bailey et al. 2005). The
Plate 35B type locality is unknown (Bailey et al.
2005).
Type material.—The original name was Distribution.—‘‘The range of T. pallidus
Coluber melanocephalus based on material seems to be the Amazon and upper
from America, and the types are listed as Orinoco watersheds from central Bolivia
‘‘NRM cat no 15 and 86’’ on the website of to eastern Colombia and Southern Ven-
Museum Adolphi Friderici at http:// ezuela and the Guianas, and to eastern
linnaeus.nrm.se/zool/herp/madserp.html. Pará, Brazil’’ (Bailey et al. 2005:89).
en. Vouchers for Guyana.—AMNH R-2665
Distribution.—‘‘Low to high elevations and R-36134 from Georgetown. Addition-
along both versants from Guatemala al specimens from Guyana were listed by
throughout Central America into South Bailey et al. (2005:101).
America as far south as southern Peru, Coloration.—Based in part on the two
Bolivia, northern Argentina, and Uru- preserved AMNH vouchers from George-
guay; also on the islands of Trinidad and town listed above, Bailey et al. (2005:88–
Tobago’’ (Wilson 1999:11). However, Sav- 89) stated the following: ‘‘chin and throat
age (2002:693) indicated that the Central unmarked except for a few tiny dark
American records noted by Wilson are dots. . .; on anterior ventrals a fine streak
on either side becomes a double (hollow) Guiana. ‘‘Colouring ranges from yellow to
streak at about heart level; before midbody light brown orange with two darker, ill-
this has become a pair of double streaks defined longitudinal bands running down
which continue about to the cloaca where the body. The scales are [speckled]...with
it becomes once more a double streak on copious fine black marks. Around the eyes
each side of the tail venter. . .; in AMNH are light brown markings which [extend
36134 the double streak never becomes posteriorly]...along the body. The belly is
paired doubles; a variable amount of light yellow.’’
accessory pigment develops including an- Comments.—Bailey et al. (2005) placed
other pair of double streaks medially in the Thamnodynastes strigilis into the synonymy
most extreme specimens. . .; some general of T. pallidus, which was reported to be
sprinkling of fine dots is sparse in AMNH semi-aquatic (Starace 1998) and sympatric
36134 to rather profuse in AMNH 101960; with Thamnodynastes ramonriveroi in
head pale, tan to brown on top, lighter Georgetown (Bailey & Thomas 2007). Com-
elsewhere or with scattered dark dots; a parative studies of sympatric populations of
double (dark-edged, pale-centered) dark these two species may be worthwhile.
streak extending from margins of the
postoculars at rear center of eye through
corner of mouth. . .disappearing on side of Thamnodynastes ramonriveroi
neck; this postocular bar slender. . .; [in Manzanilla & Sánchez, 2005
some specimens] a similar anterior bar Plate 35C
from center of orbit to top of nasal;
also. . .[in some specimens] a single or Type material.—The holotype is EBRG
double streak from top of eye across outer 3957, an adult male from Cerro La Laguna
edge of parietals outlines the darker crown (10801 0 27 00 N, 64807 0 47 00 O [¼W]) macizo del
and joins a dark paravertebral line bor- Turimiquire, municipio Freites, Anzoáte-
dering a light nuchal stripe on the gui, Venezuela, 2130 m (Manzanilla &
nape;. . .the five series of dark dorsal-spots Sánchez 2005).
are lost at about midbody and the lateral- Distribution.—This species occurs in
most row becomes transformed, along northern and eastern Venezuela, Trinidad,
with a double stripe on rows 1–2, into a Guyana, Suriname, and northern Brazil
series of lateral streaks resembling the (Bailey & Thomas 2007).
ventral ones; the paravertebral spots be- Vouchers for Guyana.—‘‘No [specific]
come reduced to dark dots on every two or locality: (AMNH 36117–20); Demerara
three scales of the 6th rows; the vertebral (BMNH 1967.2201–03)’’ (Bailey & Thom-
series is completely lost.’’ as 2007 [2006]:26); USNM 566282 from
The following is from Gorzula & Señaris Dubulay Ranch.
(1999:180) based on a living specimen from Coloration in life.—The following is
Venezuela. ‘‘Ventral surfaces lemon yellow based on the USNM specimen from
in the anterior ¼, grading to pink in the Dubulay Ranch (CJC field notes, 1994).
posterior ł; ventrals bordered laterally on Tan above with small black spots and
each side with a double brown line, single dorsolateral brown stripe; a darker tan
anteriorly; hidden skin between the dorsals stripe low on side; head tan with brown
bright yellow; dark brown dorso-lateral markings; venter yellow with brown stripes
line along dorsal scales 3 and lower 4; and spots, the lateralmost stripe of which
upper dorsal 4 and 5 and 6 are lighter; becomes gray posteriorly.
distinct post ocular stripe.’’ Comments.—This species was reported
The following is from Starace (1998:337) to be semi-aquatic and sympatric with
based on living specimens from French Thamnodynastes pallidus in Georgetown
(Bailey & Thomas 2007); comparative French Guiana. ‘‘The body. . .is usually
studies of sympatric populations may be brown to grey along which are distributed
worthwhile. fine, dark, complete cross-patterns. . .Closely
resembles Bothrops atrox. The belly is light
coloured.’’ The following is from Savage
Xenodon merremii (Wagler, 1824) (2002:582) based on specimens from Costa
Rica. ‘‘Dorsum brown, usually with a series
of 11 to 16 hourglass-shaped blotches [which
Ophis merremii, the type material disap-
may be broken up]; each blotch dark brown
peared (Hoogmoed & Gruber 1983), and
laterally and grayish brown medially, out-
the type locality is Bahia, Brazil, which is
lined by a pale brown border with some
now the city of Salvador (13800 0 S,
white areas;. . .head brown above, usually
38830 0 W) (Vanzolini 1981).
with a dark-outlined light lyre-shaped figure
Distribution.—‘‘Guianas, Brazil, Bolivia,
originating at the internasals with a branch
Paraguay, central and northern Argentina’’
running posteriorly above the eye and
(Peters & Orejas-Miranda 1970:324).
downward to the angle of jaw;. . .supralabials
light brown, edged with dark; throat cream
from Kalacoon.
with very little dark pigment, which is usually
restricted to scale margins; venter cream to
from Starace (1998:253) based on speci-
beige with much light brown stippling and
mens from French Guiana. ‘‘The body
often with light brown irregular blotches as
colouring is light brown to beige with
well; underside of tail cream, with little or no
irregular dark brown markings [usually a
dark scale margins, contrasting sharply with
pattern of alternating dark and light
venter; iris gray brown.’’
brown bands, with lighter borders]. The
Comments.—It is likely that the speci-
belly is light coloured.’’
mens from Kartabo that Beebe (1946)
Comments.—Until recently (Zaher et al.
described under the name Xenodon colu-
2009), this species was treated as a repre-
brinus were actually this species.
sentative of a different genus, Waglerophis.
Xenodon rabdocephalus (Wied, 1824) Xenodon severus (Linnaeus, 1758)

Coluber rabdocephalus based on material Coluber severus the holotype is NRM 48
from Bahia, Brazil (see Peters & Orejas- (http://linnaeus.nrm.se/zool/herp/
Miranda 1970:324). The type material is madserp.html.en). The type locality was
unknown to us. restricted to South America (see Duellman
Distribution.—‘‘Lowlands of tropical 2005).
Mexico from Veracruz and Guerrero, south Distribution.—‘‘Amazonian South
through Central America to northwestern America’’ (Peters & Orejas-Miranda
Ecuador, west of the Andes, Amazonian 1970:325).
Colombia, and Ecuador, the Guianas and Vouchers for Guyana.—AMNH R-8161
the upper Amazon portion of Brazil, Peru, from Kalacoon; AMNH R-21284, R-
and Bolivia. . .’’ (Savage 2002:583). 98222–98223, and R-137332 from Karta-
Vouchers for Guyana.—AMNH R- bo; BMNH 1930.10.10.184-1930.10.10.185
18173, R-21338, and R-98219–98221 from and 1930.10.10.187 from Morabelli Creek,
Kartabo. Essequibo River; BMNH 1934.11.1.29
Coloration in life.—The following is from from ‘‘Camaria, R. Cuyani’’ (Colin Mc-
Starace (1998:257) based on specimens from Carthy, pers. comm., 20 October 2008);
BMNH 1971.1858 from Kamarang, Ma- Vouchers for Guyana.—AMNH R-

zaruni District; ROM 22842 from Barami- 60799 from Shudikar-wau.
ta; UMMZ 47743–47745 from Dunoon, Coloration in life.—The following is
Demerara River; UMMZ 55860 from adapted from Starace (1998:265) based
Santa Rosa Island, Moruca River; on specimens from French Guiana. Body
UMMZ 65161 from Kartabo; UMMZ brick-red to light brown with a dark brown
77507 from Wismar; UMMZ 77820 from to black middorsal line; dark brown
Malalli, Demerara River; UMMZ 83644– triangular markings extend laterally from
83645 from Kurupung Up, Mazaruni the middorsal line; belly pale yellow.
District; UMMZ 85284 from junction of
Kamarang and Parvima Rivers, Parvima
Mission; UMMZ 85285 from Kurupung Xenoxybelis argenteus (Daudin, 1803a)
River, Mackreba Falls; USNM 84521
from Pomeroon; and USNM 164200– Type material.—The original name was
164201 and USNM 164213 from Maba- Coluber argenteus and the type locality and
ruma Compound (8812 0 00 00 N, 059847 0 location of the type specimen are unknown
00 00 W). to us.
Coloration in life.—Beebe (1946:44) sum- Distribution.—‘‘Lowlands of northern
marized variation in specimens from Kar- South America east of Andes’’ (Peters &
Orejas-Miranda 1970:227).
tabo as follows. ‘‘The variety of pattern and
coloring is almost indescribable. The adult
60768 from Isheartun; AMNH R-60779
snake tends to be monochrome, green,
from Marudi; and AMNH R-60800 from
brown, black or rufous, and yellowish or
Kuyuwini Landing.
gray below. But dorsal bands or spots may
persist in large specimens, black-edged
based on an individual collected by CJC
scales or more definite bands. The young
and CRT in Suriname (AMNH R-133353;
usually have a complex angular pattern of CJC field notes, 1986). Dorsum tan, but
bands and hourglass figures. One fairly dark greenish tan on posterior half of tail;
consistent mark is a large central nuchal green lateral stripe from tip of snout,
patch of pink, red, orange or rufous, through eye, along body to fade on
surrounded by one or more concentric posterior half of tail; lighter green verte-
bands of the same color. This becomes very bral stripe on body, gradually fading past
conspicuous when the snake flattens its neck, terminating abruptly on base of tail;
neck, cobra-like,’’ which other species of chin, throat, and ventral neck bright
this genus do as well. chartreuse with black spots; venter yellow-
ish green posterior to neck, changing to
cream with a midventral green stripe and
Xenopholis scalaris (Wucherer, 1861) lateral green stripe on each side; underside
of tail cream, becoming yellowish, then
Type material.—The original name was chartreuse, with a green lateral stripe on
Elapomorphus scalaris based on two spec- each side; iris gold, peppered with copper
imens from Cañavieras, Matta de S. João, but with a dark brown bar aligned with the
Bahia, Brazil. One is in the BMNH and lateral green body stripe; interior of mouth
location of the other is unknown to us. flesh pink; tongue yellow above, grayish
Distribution.—‘‘Amazonian Bolivia, Pe- tan with black flecks below.
ru, Ecuador, and Brazil’’ (Peters & Orejas- Comments.—Previously, this species was
Miranda 1970:326). More recently known assigned to two other genera, Oxybelis and
from the Guianas (e.g., Starace 1998). Philodryas (see introduction to Colubridae,
above). Often when disturbed, an individual ian border (1840 0 N, 588W). ‘‘The type
holds the tongue sticking straight forward locality may be further defined as the
from the mouth, motionless. Boundary Commission’s ‘Boundary
Camp’ at about 2000 ft above sea level,
at head of Itabu Creek. This creek enters
Elapidae the New River, an affluent of the Cour-
antyne, at New River Depot’’ (Schmidt
The families of snakes are discussed 1939:46).
above in the introduction to the snakes. Distribution.—‘‘Known from the type
The Elapidae are usually quite colorful, locality in southern Guyana, southern
mostly brightly ringed (black, yellow/ Surinam, and the region around Manaus,
white, red), highly venomous snakes, with Amazonas, Brazil’’ (Campbell & Lamar
relatively short fangs in a set or fixed 2004:150).
position near the front of the upper jaw. Vouchers for Guyana.—The holotype is
The venoms of these and other venomous the only specimen known from Guyana.
snakes are used and continue to have Coloration in life.—The following is
strong potential for pharmacological re- from Campbell & Lamar (2004:150). ‘‘This
search, both for treatment of snakebite snake has rings of black, white, and intense
and development of new medicines. red. The pattern does not consist of triads.
In the recent past, some specialists (e.g., The head is almost entirely black (some-
Campbell & Lamar 2004) put most species times with faint red markings on the snout)
of South American coral snakes in the with a short white marking that partially
genus Micrurus and a few other species in covers the last 2–4 supralabials and the
another genus, Leptomicrurus, one species temporals, and borders the parietals. This
of which occurs in Guyana (‘‘L. collaris’’). pale marking may be red in some speci-
However, Slowinski (1995:335) presented a mens; occasionally it may be interrupted by
phylogenetic analysis that suggested that black. The black head coloration is present
Leptomicrurus clustered within Micrurus, after the pale head ring but does not
so he concluded as follows: ‘‘the best constitute a nuchal ring, an arrangement
course at present is to return Leptomicru- unique for the genus. Black is the predom-
rus to the synonymy of Micrurus.’’ We inant color on the venter of the head...
follow Slowinski and recognize only one There are 8–13 black body rings, 2 dorsal
genus of coral snakes in the Guyanan scales in length, on the body. These are
herpetofauna, including the species Micru- bordered by very short white rings that are
rus collaris. interrupted so frequently as to form series
Not all colorfully ringed snakes that of spots. There are 4–7 black tail rings. The
superficially look similar to coral snakes long red body rings (12–28 ventrals in
are venomous, as several non-venomous length) are very intense in color and usually
species of Colubridae sensu lato have lack all traces of black stippling. On the tail
evolved color patterns that mimic those of the red rings are heavily melanic and
coral snakes. Also, not every individual of shorter than the black rings; the yellow or
venomous coral snake is colorfully ringed. white rings usually remain sharply demar-
cated but may be obscured.’’
Micrurus averyi Schmidt, 1939

Micrurus collaris (Schlegel, 1837)
Type material.—The holotype is FMNH
30956, an adult female from the Cour- Type material.—The original name was
antyne District of Guyana, near the Brazil- Elaps collaris and the lectotype is RMNH
1444 (Brongersma 1966). The type locality including Guyana, Suriname, and French
was restricted to the Guianas by Hoge & Guiana, as well as all of Brazil north of the
Romano (1966). Amazon River in Amazonas, and proba-
Distribution.—‘‘Southeastern Venezuela bly Roraima and Amapá (it has not been
through Guyana and Suriname to French recorded from the latter two regions)...and
Guiana and undoubtedly northern Brazil undoubtedly has a wider distribution in
in the Guianan region’’ (Campbell & the Amazon Basin than has been record-
Lamar 2004:128). ed’’ (Campbell & Lamar 2004:218).
Vouchers for Guyana.—Campbell & Vouchers for Guyana.—ROM 22836
Lamar (2004:128) stated that the holotype from Baramita; UMMZ 80423 from Ker-
for the subspecies M. c. breviventris is muri Creek, Demerara River; and UMMZ
‘‘FMNH 26658...[from] Oko Mountains, 82867 from Bartica.
Essequibo, Guyana, 400 ms.’’ Coloration in life.—The following is
Coloration in life.—The following is from Campbell & Lamar (2004:218–219).
from Campbell & Lamar (2004:129). ‘‘The snout and top of head to the
‘‘The head and dorsum are black or very posterior edge of the parietals, the mental,
dark brown. Usually there is a white, and first pair of infralabials are black. This
yellow, or red ring around the neck, is followed by an orange occipital ring that
passing just distal to the parietals or covers the postoculars, temporals, last 2–3
occasionally touching them; sometimes labials, first few dorsals, and gular region,
this ring is narrowly interrupted dorsally,
and occasionally encroaches on the parie-
with the black pigment of the head being
tals themselves. The ring is followed by 5–
continuous with that of the body.The
10 triads on the body and 2 3 –2 on the tail.
=
[underside of the] body has a distinctive
The first triad on the body is incomplete,
pattern of 35–50 oval whitish or yellowish
and the primary and accessory black rings
blotches on a black or dark brown venter.
are subequal. These triads are composed of
These blotches are 2–4 ventrals in length
and usually extend laterally to the first or long black rings (4–10 dorsals...), of which
second dorsal scale row but are not known the accessory rings are as long as or longer
to meet middorsally. The ventral blotches than the primary black rings; the black
on the tail are usually reddish and often rings are separated by relatively short (2–5
form 2–3 rings on the proximal portion of dorsals) orange (sometimes brownish
the tail.’’ orange) rings. The triads contain short
(1–3 dorsals) white rings. The orange
scales are outlined faintly in black, as are
the white scales of many specimens. An
Micrurus hemprichii (Jan, 1858)
orange ring traverses the cloacal region,
Type material.—The original name was and it is sometimes interrupted by a black
Elaps Hemprichii and the holotype was blotch. Occasionally an orange ring is
destroyed during World War II (Campbell melanized, resulting in the coalescing of
& Lamar 2004). The type locality was two triads into a quincad.’’
restricted to Venezuela by Hoge & Lancini
(1962). Micrurus isozonus (Cope, 1860b)
Distribution.—‘‘The Amazonian slopes
of the Andes in Colombia, Ecuador, Peru, Type material.—The original name was
and Bolivia; the upper Amazon Basin, Elaps isozonus and the cotypes are ANSP
Guiana highlands, and upper Orinoco 6804–6805 (Campbell & Lamar 2004). The
region from Vichada in eastern Colombia type locality was restricted to Caracas,
through southern Venezuela eastward, Venezuela (see Campbell & Lamar 2004).
Distribution.—‘‘Semi-arid and seasonal- bution of M. l. lemniscatus and, therefore,

ly dry regions in Colombia from east of the should be considered invalid.’’
Andes throughout the Llanos Orientales Distribution.—‘‘The Amazonian slopes
eastward, the Llanos of central and south- of the Andes in Bolivia, Peru, Ecuador; the
eastern Venezuela, humid forests of the Amazonian and Orinocoan watersheds of
Cordillera de la Costa in northern and Colombia, southern and eastern Venezue-
central Venezuela, Isla Margarita and the la; Trinidad; Guyana; Suriname, and
Rio Cotinga region in Roraima, Brazil... French Guiana; and the entire Amazon
[and] Guyana’’ (Kok et al. 2003:77). Basin south to the Brazilian states of
Vouchers for Guyana.—IRSNB 16573 Paraná, Mato Grosso do Sul and Minas
and 16575 from ‘‘Region 9, Dadanawa Gerais...’’ (Campbell & Lamar 2004:223).
Ranch (N 02849 0 30 00 W, 59831 0 34 00 ), South Vouchers for Guyana.—AMNH R-
Rupununi Savannahs’’ (Kok et al. 2003:78). 14142–14150 from Kartabo; ROM 11702
Coloration in life.—The following is from ‘‘Rupununi: Ruawau River’’ (Mac-
from Kok et al. (2003:73–74). ‘‘The two Culloch & Lathrop 2004b:143); ROM
Guyanese specimens are similar...Both 22834–22835 from Baramita; USNM
have 10þ1 black triads on body and tail 145461 from ‘‘Georgetown, 50 mi above,
(10 triads on the body and one on the tail). on the Abary River;’’ USNM 164215 from
The 10th body ‘triad’ of IRSNB 16573 is Mabaruma Compound (8812 0 00 00 N,
composed of 4 black bands. The first triad 059847 0 00 00 W); and USNM 566289 from
is complete and the pale yellow bands Karanambo.
(white in preservative) are equal to or a Coloration in life.—The following is
little longer than the black bands. The based on the USNM specimen from
black bands are 3–4 (generally 3) dorsal Karanambo (CJC field notes, 1992). Body
scales long; the yellow bands are 3–4 pattern consists of triads of solid black
(generally 4) dorsals long, and the red separated by narrow bands of cream (with
bands are 2–6 (generally 5 or 6) dorsals. black flecks); triads separated from each
Red and yellow bands have black apical other by bands of red (with black flecks);
tips. The snout is predominantly black, tip of tail black; tip of snout black; cream
spotted with white, followed by an irreg- band between nostrils and eye (but not
ular white band and an interorbital black around lower jaw—dorsally only); black
band. The latter is followed by a red band at eyes (dorsally only); red parietal
frontal band and a red parietal band.’’ band (completely around and forward
below head so almost whole lower jaw
Micrurus lemniscatus (Linnaeus, 1758) and chin area is red, with a couple of black
Plate 35D spots).
Comments.—ROM 11702 (see above)
Type material.—The original name was originally had been identified as an exam-
Coluber lemniscatus and the lectotype is ple of Micrurus ibiboboca but was recently
NRM 93 (Roze 1989; http://linnaeus.nrm. re-identified (see MacCulloch & Lathrop
se/zool/herp/madserp.html.en). Roze 2004b).
(1967) stated: ‘‘The high number of black
triads combined with ventral and subcau-
dal counts and the general coloration of Micrurus psyches (Daudin, 1803b)
the Linnaean syntypes conform to the
populations of the northern Guianas. The Type material.—The original name was
restriction of the type locality to Belém, Vipera psyches and the holotype, from
Pará, Brazil, by Schmidt & Walker (1943) Surinam, is MNHN 7654 (Campbell &
does not correspond to the known distri- Lamar 2004).
Distribution.—‘‘Northern South Ameri- 2004). Donnelly et al. (2005:457) reported

ca from eastern Venezuela through north- that both M. circinalis and M. psyches were
ern Guyana to Suriname, French Guiana, found at Iwokrama, but they did not cite
and Brazil in the state of Amapá’’ (Camp- specific specimens and our requests to
bell & Lamar 2004:189). examine vouchers were not answered (see
Vouchers for Guyana.—AMNH R- Species Reported. . .Probably in Error, be-
21333, R-110160–110162, and R-137321, low).
all from Kartabo; ROM 28378 from
Paramakatoi.
Coloration in life.—Campbell & Lamar
(2004:189) stated that ‘‘this is a tricolored Micrurus surinamensis (Cuvier, 1817)
coralsnake with black rings arranged Plate 35E
singly. There are 22–41 black body rings
that are 3–4 dorsal and ventral scales long.
The white or yellowish rings are 1 dorsal Elaps Surinamensis and MNHN 3926 was
long or less; these rings are also sometimes designated as the lectotype by Schmidt
obliterated on the dorsum owing to (1952). The type locality is Surinam (see
melanization, but they are evident across Campbell & Lamar 2004:230).
the venter. Red (melanic) body rings are Distribution.—‘‘The Orinocoan and
usually about 3–7 dorsals and ventrals Amazonian drainages of Colombia; Ama-
long, and are generally slightly longer than zonian drainages of Ecuador, Peru, and
the primary black body rings. The red Bolivia; the entire Amazon Basin of Brazil
rings are so heavily melanized that they from northern Mato Grosso and Maran-
appear uniformly black and specimens hão northward including Rondônia, Acre,
appear to be bicolored. In some cases the Amazonas, Roraima, Pará, and Amapá;
red rings are only partially melanized and Guyana, Suriname, and French Guiana;
the rings are blackish red or purplish. Red and southern Venezuela’’ (Campbell &
rings are in evidence on the belly even in Lamar 2004:230–231).
dark snakes on which the dorsal parts of Vouchers for Guyana.—AMNH R-
the red rings have been totally obfuscated 152304–152305 and USNM 497807 from
[by black]. The tail is patterned with black Berbice River Camp; and CJC and CRT
and white rings or with black, red, and saw a specimen (not collected) at Dubulay
white rings...The head is black, including Ranch.
all supracephalic scales; this black head Coloration in life.—The following is
cap usually coalesces with the black nuchal
based on the AMNH and USNM spec-
ring...The side of the head including the
imens from Berbice River Camp (CJC
temporals and last 1–2 supralabials is
field notes, 1997). This species is distinc-
white with considerable black markings.
The white occipital ring is usually inter- tive in that the head is red but with the
rupted behind the parietals owing to the cephalic scales being edged with black.
fusion of the black head coloration with The pattern on the body is of triads of
the nuchal ring...All white scales [on the black. Red, black, and pale yellow bands
head] are tinged and edged with black. The completely encircle the body, the pale
chin is mostly black, including the infrala- yellow ones being white on the belly and
bials and chinshields.’’ the red ones having black spots dorsally;
Comments.—A closely related species, tail tip black.
Micrurus circinalis, previously treated as a Comments.—This species is one of the
subspecies of M. psyches, occurs in Ven- longest and stoutest of all the coral snakes
ezuela and Trinidad (Campbell & Lamar (Campbell & Lamar 2004:231).
Leptotyphlopidae Grande do Norte, and Argentina’’

(McDiarmid et al. 1999:20), but see
Leptotyphlopids have been recognized as Comments, below.
a distinct family of snakes for well over 100 Vouchers for Guyana.—AMNH R-
yr, based on morphology. Recent phyloge- 14270 and R-137351 from Kartabo;
netic analyses of DNA sequence data AMNH R-98188 from near Kartabo;
indicate that these snakes have existed for AMNH R-102569 from Camaria; AMNH
perhaps 130 million yr (before the breakup R-141884–141885 from Dubulay Ranch;
of Gondwanaland, although the group has AMNH R-152310–152317 and USNM
experienced over-water dispersal also), and 566292–566295 from Berbice River Camp;
that their closest living relatives may be the ROM 20503 from Kurupukari; ROM
superficially similar Typhlopidae, which 22847 and USNM 535814 from Baramita;
share a burrowing way of life, reduced eyes, and USNM 304357 from Kartabo Point
and ecological association with ants and (6823 0 00 00 N, 058841 0 00 00 W).
termites (Adalsteinsson et al. 2009, Vidal et Coloration in life.—The following is
al. 2010b). The Leptotyphlopidae includes based on AMNH and USNM specimens
about 116 species and the family has a from Dubulay Ranch and Berbice River
pantropical to subtropical distribution. This Camp (CJC field notes, 1995, 1997). Dark
family includes species that are among the brown above and below; tip of tail
world’s ‘‘smallest and thinnest species of (including ‘‘spine’’) bright yellow; less
snakes, often called threadsnakes (or worm- conspicuous yellow spot on snout.
snakes)’’ (Adalsteinsson et al. 2009:2), as Comments.—In the past, Guyanan spec-
does the family Typhlopidae. Until recently, imens of this snake were referred to the
the four species that occur in Guyana were species Leptotyphlops tenellus Klauber,
all considered to be members of the very 1939, with the type locality of Kartabo,
large genus Leptotyphlops, but phylogenetic Guyana. After discussing taxonomic prob-
analyses have resulted in taxonomic changes lems of these small snakes, Hoogmoed &
and now each of these four species is Gruber (1983:340) stated ‘‘that L. tenella
included in a different genus (Adalsteinsson Klauber, 1939 is nothing but a synonym of
et al. 2009). ‘‘The family is one of the most S. albifrons Wagler, 1824. The correct
poorly known of all terrestrial vertebrates name for the taxon...actually is L. albifrons
from the standpoint of systematics and (Wagler, 1824).’’ However, McDiarmid et
ecology’’ (Adalsteinsson et al. 2009:2). al. (1999:20) were not so certain that this
matter was clearly resolved, and Franco &
Pinto (2009) later decided that the appro-
Epictia albifrons (Wagler, 1824) priate name is Leptotyphlops tenellus after
Plate 36A, B all, but this was not supported by Avila-
Pires et al. (2010). Clearly, a thorough
Type material.—The original name was review of all known specimens, accompa-
Stenostoma albifrons and the holotype was nied by molecular data from specimens
destroyed during World War II (Hahn representing the entire geographic range,
1980). The type locality is the vicinity of would be appropriate.
Belém, Pará, Brazil (see Peters & Orejas-
Miranda 1970).
Distribution.—‘‘. . .Trinidad, Guyana, Siagonodon septemstriatus
Suriname, French Guiana, southeastern (Schneider, 1801)
Venezuela, northeastern Peru and possibly
Ecuador, Amazonian Brazil south to Mato Type material.—The original name was
Grosso and Pará, Pernambuco, and Rio Anguis Septemstriatus and the holotype,
from an unknown locality, is ZMB 3876 ZMB 1434 (Orejas-Miranda 1967). The
(Bauer et al. 2002). type locality was restricted to Puerto
Distribution.—‘‘Southeastern Ven- Cabello, Venezuela (Orejas-Miranda
ezuela. . ., Guyana, Suriname, French Gui- 1967).
ana, and northern Brazil’’ (McDiarmid et Distribution.—‘‘Panama, Colombia,
al. 1999:43). Venezuela, Guyana, Suriname, French
Vouchers for Guyana.—AMNH R- Guiana, and northern Brazil’’ (McDiarmid
98187 from Kartabo. et al. 1999:35).
Coloration in life.—Beebe (1946:13) stat- Vouchers for Guyana.—ROM 28367 and
ed the following based on specimens from USNM 291191 from Paramakatoi, the first
Kartabo. ‘‘The body color varies from pale of which was identified by Van Wallach.
lilac to rich golden orange with seven to Coloration in life.—The following is
nine dark lines down back and sides. In from Starace (1998:78) based on specimens
most specimens the lowermost line is from French Guiana. ‘‘The dorsum is grey-
broken or reduced to a fine thread of brown. Each dark-brown dorsal scale has
pigment.’’ The following is from Starace a cream edging, forming a regular overall
(1998:80) based on specimens from French pattern. The belly is of a lighter brown.’’
Guiana. ‘‘The dorsum is light yellow,
striped with seven brown longitudinal lines,
the belly is a shade lighter [yellow] in color.’’ Typhlopidae
There are about 260 species of typhlopid
Tricheilostoma dimidiatum (Jan, 1861) snakes, whose closest living relatives are
the Leptotyphlopidae (see discussion of
Type material.—The original name was the family, above) and Anomalepididae.
Stenostoma dimidiatum and the holotype The common ancestor of these groups
was destroyed. The neotype is IBSP 24011, existed about 160 million yr ago, before
an adult female from São Marcos the breakup of Gondwanaland, based on
(03805 0 N, 60825 0 W; 75 m), Boa Vista molecular phylogenetic analyses of DNA
Municipality, Roraima, Brazil (Pinto & sequence data (Vidal et al. 2010b). Three
Fernandes 2012). species of this rather ancient family are
Distribution.—‘‘Guyana, Suriname, known to occur in Guyana.
southeastern Venezuela, and northern
Brazil in the Rupununi Savanna and its
extensions, usually in only slightly elevated Typhlops brongersmianus Vanzolini, 1976
coastal areas’’ (McDiarmid et al. 1999:27). Plate 35F
and USNM 566291 from Karanambo. Type material.—The original name was
Coloration in life.—The following is Typhlops brongersmai (Vanzolini, 1972) but
based on AMNH and USNM specimens it was preoccupied by the name of a different
from Karanambo (CJC field notes, 1992). species, Typhlops florensis brongersmai. The
Brown dorsum; translucent pinkish gray replacement name was published by Vanzo-
ventral surface. lini in 1976. The holotype is MZUSP 5218
and type locality is Barra de Itaipe, Ilheus,
Bahia, Brazil (Vanzolini 1972).
Tricheilostoma macrolepis (Peters, 1857) Distribution.—‘‘Trinidad south through
South America (east of the Andes) to
Type material.—The original name was Buenos Aires Province, Argentina (Co-
Stenostoma macrolepis and the lectotype is lombia, Trinidad, Venezuela, Guyana,
Suriname, Peru, Brazil, Bolivia, Paraguay, Paramaribo, Surinam, is RMNH 7660, a

Argentina)’’ (McDiarmid et al. 1999:94). subadult male (Dixon & Hendricks 1979).
Vouchers for Guyana.—AMNH R- Distribution.—‘‘Tropical South America
140972 and USNM 566296 from Dubulay east of the Andes between 128N and 148S
Ranch. (Colombia, Venezuela, Guyana, Suriname,
Coloration in life.—The following is French Guiana, Brazil, Ecuador, Peru, and
based on the AMNH and USNM speci- Bolivia); one specimen reported from
mens from Dubulay Ranch (CJC field coastal Ecuador’’ (McDiarmid et al.
notes, 1994, 1995). Dark brown above, 1999:117).
but very light tan on snout and on ‘‘spine’’ Vouchers for Guyana.—AMNH R-8110
at tip of tail (or ‘‘spine’’ dark brown with from Kalacoon; AMNH R-21280, R-
tan band anterior to it); body at times with 21332, R-67882, and R-137350 from Kar-
a few small irregular tan spots posteriorly; tabo; AMNH R-141881, AMNH R-
yellowish tan or translucent tan below, 141882, and USNM 566298 from Dubulay
with increasing yellow posteriorly. Ranch; ROM 28368 from Paramakatoi;
and USNM 164216–164217 from Maba-
ruma (8812 0 00 00 N, 059847 0 00 00 W).
Typhlops minuisquamus
Dixon & Hendricks, 1979
mens from Dubulay Ranch (CJC field
Plate 36E
notes, 1994, 1995). The colors are similar
Type material.—The holotype is TCWC to those of T. brongersmianus, with the
39130, an adult female from Mishana, dorsum black or very dark brown, but the
Loreto, Peru. ‘‘spine’’ at the tip of the tail is dark brown
Distribution.—‘‘Southeastern Colom- and there are two dorsal light tan patches;
bia. . ., southern Venezuela. . ., northeastern one posteriorly on the body; one just
Peru. . ., Brazil. . ., and Guyana (Kartabo, anterior to the ‘‘spine’’ on the tail. The
Kamakusa, and Tacoba)’’ (McDiarmid et ventral surface is light tan (darker anteri-
orly).
al. 1999:111).
25050 from Kamakusa; AMNH R-25087
from Tacoba; AMNH R-98189 from near Viperidae
Kartabo; and USNM 566297 from Dubu- The families of snakes are discussed
lay Ranch. above in the introduction to Serpentes.
Coloration in life.—The following is Among these, the Viperidae are usually
based on the USNM specimen from cryptically colored sit-and-wait predators
Dubulay Ranch (CJC field notes, 1995). that blend into their environment incon-
Dorsum (including tail and ‘‘spine’’) very spicuously, whether on the ground or in
dark brown; snout pale tan but tip with vegetation. These are venomous snakes,
narrow, vertical, dark brown stripe; ven- some of which have venom that is
tral surface translucent tan, lighter poste- dangerous to humans. The venom is
riorly than anteriorly. delivered through relatively long, hollow
fangs that can be swung forward into
Typhlops reticulatus (Linnaeus, 1758) biting position near the front of the upper
Plate 36C, D jaw; otherwise they lie back on the roof of
the mouth. However, if one is bitten by a
Type material.—The original name was viper, that does not necessarily mean that
Anguis reticulata and the neotype, from one has been envenomated; many defen-
sive bites by pit vipers are not accompa- southeastern Brazil’’ (McDiarmid et al.
nied by injection of venom (so-called ‘‘dry 1999:248).
bites’’). Vouchers for Guyana.—AMNH R-
The main function of the venom is to 60825 from ‘‘British Guiana’’ but without
subdue prey. Occasionally it will be used additional locality data, collected by R.
for defense, particularly if a person steps Snedigar, Terry-Holden Expedition;
on a snake or accidently hits it while AMNH R-98433 from near Kartabo; and
working in vegetation. Otherwise, pit UMMZ 65272 from Kartabo.
vipers usually allow very close approach Coloration in life.—This is a colorful
by humans without revealing their pres- arboreal snake. ‘‘It is pale leaf green above,
ence and without striking, frequently uniform or sparsely spotted with darker
allowing people to walk past them within [areas], and yellow below’’ (Beebe 1946:50,
inches, and without being aware of the summarizing three specimens from Karta-
snake (Hardy 1994). In fact, Hardy bo). In following paragraphs on the same
(1994:202) stated that in Costa Rica page, Beebe elaborated on one specimen as
‘‘biologists spent more than 1.5 million p- follows. ‘‘General color above lichen
hr [¼person hours] in the field with B. asper green, shading to grass green toward tail,
resulting in just three bites.’’ The venoms which changes abruptly to pale pinkish-
of these and other venomous snakes have buff near the tip. Lateral line on side of
been successfully used for, and still have head and about 40 sets of alternating,
strong potential for, pharmacological re-
irregular, short, transverse dorsal marks
search.
light cadmium yellow, edged with black.
The six species of viperids that occur in
Side of head greenish-yellow, shading into
Guyana all have an extremely sensitive
a wide patch of picric yellow on the side of
heat-detecting pit on each side of the face
the neck, which in turn merges into the
between they eye and nostril (although
lateral body line of straw deepening to
often on a lower plane). Together with its
interpretation center in the brain, this citron toward the tail. Ventrals pale
organ can accurately sense the presence, lumiere green lighter toward tail, touched
size, and location of nearby warm objects, with lighter greenish-yellow on the lateral
such as warm-blooded prey, under condi- edges of the ventrals. Iris light yellow green
tions of absolute darkness (e.g., at night or flecked with darker green, paler toward
in burrows). pupil.’’ Beebe described other individuals
as having black spots and orange colora-
tion in places on the dorsum, including one
with ‘‘blue green’’ on the belly.
Bothriopsis bilineata
Comments.—This is an arboreal pit
(Wied-Neuwied, 1821)
viper with cryptic coloration and a pre-
Type material.—The original name was hensile tail, which is relatively longer than
Cophias bilineatus and the type specimen is in other pit vipers. Is the distinctly colored
‘‘unlocated’’ (McDiarmid et al. 1999:248). end of the tail used as a caudal lure to
The type locality was restricted to Vila attract prey, as in some other pit vipers?
Viçosa [Marobá], on Rio Peruhybe, Bahia,
Brazil (see McDiarmid et al. 1999).
Distribution.—‘‘Amazonian South Bothriopsis taeniata (Wagler, 1824)
America: Colombia, Venezuela, Guyana,
Suriname, French Guiana, Brazil, Ecua- Type material.—The original name was
dor, Peru, and Bolivia. An isolated Atlan- Bothrops taeniatus. Hoogmoed & Gruber
tic slope population is known from (1983) designated MNHN 1582 as the
neotype. It is from Goiás, Brazil (see hensile tail and cryptic coloration. Is the
Campbell & Lamar 2004). distinctly colored end of the tail used as a
Distribution.—‘‘Colombia, Venezuela, caudal lure to attract prey, as in some
Guyana, Suriname, French Guiana, Bra- other vipers?
zil, Ecuador, Peru, and Bolivia’’ (McDiar-
mid et al. 1999:252).
Vouchers for Guyana.—UF 142004, two Bothrops atrox (Linnaeus, 1758)
photographs that document a specimen Plate 36F
from ‘‘Mazaruni-Potaro District, Woko-
mung Massif, lat. 05804 0 03.3 00 by long. Type material.—The original name was
59851 0 44.8 00 , ca. 1135 m elev.’’ (Means Coluber atrox and the syntypes are NRM
2004:410). 100–101 (McDiarmid et al. 1999). The type
Coloration in life.—The following is locality was restricted to Surinam (Schmidt
from Campbell & Lamar (2004:322). & Walker 1943).
‘‘The pattern [of adults] is exceedingly Distribution.—‘‘Tropical lowlands of
complex, and the overall color varies from South America east of the Andes;. . .
lavender gray to yellow-green. The head is southeastern Colombia, southern and east-
the same shade as the rest of the ground ern Venezuela, Guyana, Suriname, French
color, but with several black blotches on Guiana, eastern Ecuador, eastern Peru,
top and profuse black-and-yellow speck- northern Bolivia, and the northern half of
ling overall. A black postocular stripe Brazil’’ (McDiarmid et al. 1999:257).
terminates at the angle of the mouth. The Vouchers for Guyana.—AMNH R-6803
iris is speckled black and yellow. The chin from Libarria Savannah, Kaieteur;
is pale yellow, the dark speckling around AMNH R-14254–14256, AMNH R-
the labials becoming heavier posteriorly 21287, and USNM 257527 from Kartabo;
such that the entire underside becomes AMNH R-152293–152301 and USNM
predominantly maroon-brown with the 566284–566288 from Berbice River Camp;
yellow ground color showing through as AMNH 152302 from Dubulay Ranch;
specks. Dorsally the pattern consists of 26– ROM 22848 from Baramita; ROM 43202
40 black bands that are speckled through- from Menzies Landing, Kaieteur; and
out, often fail to meet middorsally, and USNM 531703–531705 from Iwokrama
may be divided on the sides, thus appear- Forest Reserve (various localities).
ing to be divided into 4 parts. There are Coloration in life.—The following is
areas of concentrated black-and-yellow based on AMNH and USNM specimens
speckling midway between the dorsal from Berbice River Camp and Dubulay
bands. A series of 50–90 white or yellow Ranch (CJC field notes, 1997). Dorsal
spots is situated at the juncture of the colors of browns and tans; ventral surface
dorsals and ventrals; each spot covers 1–2 brown with light gray spots; dark brown
dorsal scales and part of the adjacent 1–2 face mask; posterior 40% or so of tail
ventrals with an interspace of 1–4 ventrals yellow on juveniles. Colors and patterns
between them. The distal third to half of can be variable, so we quote Beebe’s
the tail is often pink, and the tongue is (1946:49) descriptions of several speci-
black.’’ Considerable ontogenetic change mens. A young snake was ‘‘reddish-brown
in colors and pattern exist. above with cross bands of dark olive, each
Comments.—The UF voucher specimen constricted at mid-back. Whitish below,
was ‘‘collected about 1 m off the ground in thickly checkered with pinkish-brown. Iris
cloud forest vegetation’’ (Means 2004:410). golden yellow, covered with a vertically
As the other species of Bothriopsis in striated mesh of black.’’ A larger specimen
Guyana, this arboreal species has a pre- had ‘‘a series of transverse, broad, hour-
glass figures separated by narrow diamond surface similar to specimens of B. atrox in

interspaces. The boundaries are formed by the series of AMNH R-152293–302, so we
narrow, yellowish-white lines. The anterior prefer this identification. In addition,
half of the body is of a general pale brown, AMNH R-14254–14256 and 21287 are
the posterior half being much darker with Beebe specimens from Kartabo, identified
a thick scattering of round, blackish- as B. atrox, which show variation in the
brown spots. Black, lateral head line and anterior dorsal pattern. Combined modern
pale yellow tail as usual. Ventrals domi- morphological and genetic research on a
nately black with many yellow spots.’’ A large series of Bothrops from Kartabo
larger breeding female had ‘‘head purplish might be worthwhile, to clarify the status
gray, paler at the posterior sides and of the population there. Is there one
pinkish at snout. Sides of head also purple, species, are there two, and are two taxa
all labials thickly mottled with dark interbreeding in the vicinity of Kartabo?
brown, and the usual lateral dark line.
Body purplish-gray with dark, transverse
triangular markings of dark lilac, edged Bothrops brazili Hoge, 1954
with smoke gray. Chin light buff tinged
with purplish, ventrals naples yellow, Type material.—The holotype, from
upper edges with touches of pink, fading Tomé Assú, Acará Mirim River, Pará,
to cream color, and mottled more and Brazil, is 14.721 in the Butantan Institute,
more thickly with bluish-gray toward tail. Brazil.
Iris warm yellow with dense flecking of Distribution.—‘‘Equatorial forests of
dark gray and brown all over, except for southern Colombia, eastern Ecuador,
pupil rim.’’ southern and eastern Venezuela, Guyana,
Comments.—AMNH R-137323, cata- Suriname, French Guiana, Brazil, eastern
loged only recently, was said to have been Peru, and northern Bolivia (McDiarmid et
collected by W. Beebe at ‘‘Kartabo,’’ al. 1999:258).
decades prior to cataloging. Aspects of Vouchers for Guyana.—AMNH R-8992
the anterior dorsal pattern of this specimen from Kalacoon, identified by M. S.
compare favorably with specimens identi- Hoogmoed.
fied as B. venezuelensis from Rancho Coloration in life.—The following is
Grande, Venezuela (AMNH R-137298– from Campbell & Lamar (2004:383).
137299), where Beebe also collected. This ‘‘The dorsum of the head is pinkish tan
Kartabo specimen is in poor condition, to pinkish or reddish gray, usually uni-
however, so not all characters are clear. form. In a few specimens a pale brown,
Perhaps Beebe sent or took it to the New poorly defined postocular stripe bordered
York Zoological Park alive, and, as for below by a dark brown line extends from
other specimens, the AMNH received it the eye to the angle of the mouth. The
some time after it died and was retained in rostral and inferior edges of the supra-
a freezer for an unknown period of time. labials are paler than the rest of the head.
This could have allowed for a mix-up in A lateral series of 9–19 dark ash gray
locality data, on which Barbour (1920) has bands or triangular Cs, alternating or
commented, and which we question for opposite each other, crosses the dorsum
some other Beebe records (e.g., Tretioscin- of the body. The dorsal ground color is
cus bifasciatus, see below). Nevertheless, similar to that of the head but is usually
this specimen has dark spots on the labial darker with a pinkish or reddish cast. It
region, virtual absence of light stripes on appears that specimens from the northern
the head, and a mix of pale and dark portion of the range more frequently [than
brown markings on the anterior ventral specimens from the south] have a pattern
of rectangular or slightly triangular cross- Alexander Mendes has seen the species
bands...The vertebral region often has a at Dubulay Ranch and Duane De
rusty tinge. The dorsum of the tail is Freitas (pers. comm.) has seen it at
similar in color to that of the body but Dadanawa Ranch (Plate 37A).
may be predominantly black. The venter is Coloration in life.—Beebe (1946:51)
usually yellow or pinkish cream with stated the following about a specimen
slightly darker mottlings. Conspicuous found at Kartabo. ‘‘Like the other
gray splotches are present at the juncture Kartabo specimens this rattler was green
of the ventral and dorsal scales, roughly on in general coloring, with variegated
alternate ventrals. The first row of dorsals shades from pale to deep forest green,
is usually the color of the ventrals. The iris with irregular dorsal markings of black.
is usually reddish, pinkish copper, or The eyes were connected by a black
bronze. The tongue is brown with pink band, and from the eyes back, two
flecks.’’ broad bands of dark brown extended
along the body for some distance. The
iris was green with two large vertical
Crotalus durissus Linnaeus, 1758 patches of brown mottling on the central
Plate 37A three-fifths...Below, the ventrals were
creamy white, anteriorly deepening into
Type material.—Savage et al. (2005:370) greenish posteriorly.’’
designated RMNH 12857, an adult male Campbell & Lamar (2004:544–545)
as the neotype. This specimen is from near stated that ‘‘there are 18–32 rhombic or
2.5 km east of Tibiti, Saramacca District, diamond-shaped dorsal body blotches...
Suriname (ca. 5833 0 N, 55854 0 W). Middorsally, blotches are separated by 1
Distribution.—‘‘Mexico. . .to Costa or 2 pale scales. The body blotches, at
Rica. . .; many disjunct populations in least anteriorly, are reddish-brown, dark
northern South America (Colombia, Ven- brown, or black with paler interiors,
ezuela, Guyana, Surinam, French Guiana, usually tan to pale gray-brown except in
and northern Brazil); Colombia and east- some southern South American snakes.
ern Brazil to southeastern Peru, Bolivia, Posteriorly, dorsal blotches tend to
Paraguay, Uruguay and northern Argen- become shorter and wider, in some
tina. . .’’ (McDiarmid et al. 1999:283). The populations coalescing with lateral body
species also occurs in the West Indies blotches and in others remaining dis-
(Campbell & Lamar 2004). crete. The lateral blotches are below the
Vouchers for Guyana.—AMNH R- lateral points of the dorsal series and
60754–60755 from Isheartun; AMNH usually comprise groups of 3–7 dark
R-60796 from Naju Creek, near Takutu scales bordered by pale scales...Another
River; AMNH R-60813 from Charwair secondary series of lateral blotches
Creek; AMNH R-60827 from Wichabai, alternates between the primary lateral
16 km N Dadanawa; UF 16157 and series; this series is usually more diffuse
16159–16161 from ‘‘the McTurk Home- or less dark than the primary series and
stead on the Rupununi River, about 3 involves the outer edges of the ven-
mi E Karanambu, British Guiana trals...The chin and gular region is
(¼G u y a n a ) ’’ ( H a r r i s & S i m m o n s usually mostly white or yellowish, al-
1977:306); USNM 291153 from ca. 5 though some populations have dark
km N of Kato; and USNM 497808 from flecking or mottling, especially on the
Karanambo Ranch. In addition, Beebe infralabials. The belly is usually whitish,
(1946:51) reported that the species is yellowish, or buff, and varies from
‘‘very rare in the jungle at Kartabo’’; nearly immaculate to white with pale
gray blotches or mottling that becomes and the dorsal markings became very
progressively darker posteriorly.’’ irregular and whitish on the tail.’’
Comments.—Harris & Simmons (1977) Campbell & Lamar (2004:447) stated
named a new subspecies as Crotalus that ‘‘the ground color is reddish brown,
durissus trigonicus, for which the UF pinkish tan, orangish tan, yellow, or grayish
vouchers cited above are paratypes. This yellow, and often obscurely mottled, espe-
subspecies was recognized by Campbell cially posteriorly. There are 28–35 dark
& Lamar (2004:546) who stated that it ‘‘is brown or blackish dorsal diamond-shaped
known with certainty only from the blotches (forming triangles laterally)...The
Rupununi savannas in southwestern dark dorsal markings are often narrowly
Guyana, and probably from the adjacent bordered by yellow or cream...The venter is
Território Federal de Roraima, Brazil.’’ white or ivory, with this color overlapping
the first 1 or 2 rows of dorsals anteriorly but
invaded by short brown intrusions (ends of
Lachesis muta (Linnaeus, 1766) dorsal bands) posteriorly.’’
Comments.—Three species of bushmas-
ters are recognized today, two in Central
Crotalus mutus and type material, from
America and one (L. muta) in South
Surinam, is in the NRM (see McDiarmid
America (Zamudio & Greene 1997, Fer-
et al. 1999).
nandes et al. 2004).
Distribution.—‘‘South America: L. muta
muta occurs in equatorial forests east of
the Andes (eastern Colombia, eastern Species Accounts: Reptilia: Testudinata
Ecuador, Peru, northern Bolivia, eastern (Turtles and Tortoises)
and southern Venezuela, Guyana, Suri-
name, French Guiana, and much of These uniquely shelled vertebrates, which
northern Brazil); L. muta rhombeata oc- are toothless and have their shoulder and
curs in coastal forests of southeastern hip girdles inside their rib-cage, have a
Brazil’’ (McDiarmid et al. 1999:313). history that stretches back more than 200
Vouchers for Guyana.—AMNH R-8172 million yr. In many species, as in crocody-
from Kalacoon; AMNH R-60850 from lians, the sex of a developing embryo is
Onora; AMNH R-98434–98435 from near determined by the temperature of the egg’s
Kartabo; and USNM 291192 from Para- immediate environment in the nest at a
makatoi. In addition, Alexander Mendes critical point of development. We do not
(pers. comm.) has seen this species at know what molecular phenomena control
Dubulay Ranch. this.
Coloration in life.—Beebe (1946:48) de- All turtles reproduce by laying eggs,
scribed an individual from Kartabo as usually buried on land (few in open or
follows. ‘‘This was essentially a yellow simply covered nests), and several have
brown phase and complete with the become endangered owing to over-con-
pattern of jungle debris when coiled as sumption of meat, eggs, tortoiseshell, and
we first found it. The eye was scarlet, the oil by humans. A modern phylogenetic
head and much of the sides of the body analysis of the major groups of turtles of
pale wood brown. The postocular black the world was provided by Shaffer et al.
band was very strongly marked. The (1997), based on morphological (largely
dorsal diamonds were almost solid black, skeletal) and DNA sequence data.
with a few rufous scales at their center, and Fifteen species representing seven fam-
each banded with a narrow frame of pale ilies are known to occur in Guyana, and
buff. The ventrals were yellowish-brown, these include a spectrum of diversity from
sea turtles to freshwater aquatic turtles to Mesoclemmys gibba (Schweigger, 1812)

terrestrial tortoises, as well as the remark- Plate 37B–D
able turtle (leatherback sea turtle) that
attains the greatest weight and longest Type material.—The original name was
Emys gibba and the holotype is MNHN
carapace in the world today. Most of the
8756, an adult female (Ernst 1981). The type
literature concerning turtles specifically
locality was restricted to near Cayenne,
from Guyana has been published by
French Guiana (see McCord et al. 2001).
Pritchard (e.g., 1964, 1966, 1969, 2005),
Distribution.—Trinidad and northern
with emphasis on sea turtles. South America, primarily in the Amazon
and Orinoco drainage systems (Brazil,
Colombia, Ecuador, the Guianas, Peru,
Chelidae and Venezuela; McCord et al. 2001).
These are aquatic turtles that live in Vouchers for Guyana.—AMNH R-
fresh water. They are called side-necked 64721 from Kartabo and PCHP 1820 from
turtles because they bend the neck to the Mabura Pond, 10 mi SW Linden. In
side when hiding the head. Representatives addition, CJC and CRT photographed
occur only in South America and the and released a specimen at Dubulay Ranch
(Plate 37B–D).
Australia-New Guinea areas.
from Ernst & Barbour (1989:53). ‘‘The
carapace is chestnut brown to dark gray
Chelus fimbriatus (Schneider, 1783)
or black. . .The plastron is red brown to
Plate 37E
yellow with a brown blotch on each scute,
Type material.—The original name was and a narrow yellow border may occur
Testudo Fimbriata, the type material is anteriorly and posteriorly. Bridge and
unknown to us, but Fritz & Havaš (2006) undersides of the marginals are brown to
gave the type locality as Surinam. yellow. . .Head and neck are red brown to
Distribution.—This species occurs in dark gray dorsolaterally and grayish to pale
‘‘. . .most major drainages in northern yellow ventrally; the jaw may contain dark
Bolivia, eastern Peru, Ecuador, eastern spots and the two small chin barbells are
yellow. The upper jaw is often yellow to
Colombia, Venezuela, the Guianas, and
white with black bars. . .Limbs and tail are
northern and central Brazil. It has also
gray black, [areas of] limb sockets yellow.’’
been found on Trinidad, apparently
Comments.—This species formerly was
washed there by mainland floods’’ (Ernst
included in the genus Phrynops, but
& Barbour 1989:66). McCord et al. (2001) treat Mesoclemmys
Vouchers for Guyana.—PCHP 1823 as a separate monotypic genus.
from Essequibo, Rockstone; PCHP 2828
from Karanambo; PCHP 3909, 3933–
3934, 3985, 5245–5246, 5460, and 5728
Phrynops tuberosus (Peters, 1870)
from Yupakari; and UTA R-37828–37829
Plate 37F
from Guyana (no additional locality data).
Coloration in life.—The following is Type material.—The original name was
from Ernst & Barbour (1989:66). Carapace Platemys tuberose, and the holotype, from
brown to black, often carrying aquatic Cotinga River, Mt. Roraima, Guyana, is
algae; plastron and bridge cream to yellow, ZMB 166 (McCord et al. 2001).
possibly brown. ‘‘Head, neck, limbs, and Distribution.—‘‘Northeastern South
tail of adults are grayish brown.’’ America (eastern and southern Venezuela,
Guyanas, Surinam, northeastern Brazil)’’ and the Guianas and the Amazon drain-
(Fritz & Havaš 2006). ages from northeastern Bolivia, eastern
Vouchers for Guyana.—PCHP 4485 Ecuador and Peru and southeastern Co-
from the Ireng River; PCHP 4590, 4645, lombia eastward to the vicinity of Belem,
4685, and 4917 from near Imbaimadai; Brazil’’ (Ernst & Barbour 1989:62).
and USNM 291154–291156 from Kato. Vouchers for Guyana.—AMNH R-
Coloration in life.—The following is 15145 from Kartabo; AMNH R-61523
from Ernst & Barbour (1989:59), in their and AMNH R-61531 from head of Rupu-
account for Phrynops geoffroanus (see nuni River; AMNH R-61528–61529 from
below). ‘‘The carapace is brown to black Shudikar-wau River; PCHP 0443 from
with gray mottlings and has a yellow NW District, near Mathews Ridge; and
border. . .In older adults, the undersides USNM 531706 from the Iwokrama Forest
of the marginals, bridge, and plastron may Reserve (04825 0 12 00 N, 058850 0 58 00 W).
be uniformly yellow to light brown; Coloration in life.—The following is
juveniles and young adults have an exten- from Ernst & Barbour (1989:61–62).
sive red and black plastral pattern. . .the ‘‘The carapace is. . .yellow with dark-
head is gray to olive dorsally, often with brown or black pigment covering varying
black vermiculations; a broad black stripe amounts of the surface [see next para-
runs backward on each side from the graph]. . .The plastron is dark brown or
nostril through or over the orbit and black with a yellow border; the bridge is
tympanum to the side of the neck, and a yellow with a dark transverse bar. . .The
second black stripe runs along the upper head is orange to yellow brown dorsally,
jaw to the side of the neck; between these but dark brown to black laterally and
stripes is a yellow or cream-colored band. ventrally; the light dorsal pigment extends
On the yellow chin and underside of the downward on the sides to the midpoint of
throat is a series of black streaks or stripes. the orbit and tympanum. The unnotched
The jaws are yellow. . .Limbs are gray to jaws are dark brown. . .Two small brown
olive on the outside, but have cream- chin barbells are present, and the iris is
colored areas beneath; soles and palms of brown. The neck is colored similarly to the
the feet may be black.’’ The two chin head. . .Anterior surfaces of the black limbs
barbells are yellow. are covered with large scales. . .The black
Comments.—Previously, these turtles tail is short.’’
were referred to as Phrynops geoffroanus The subspecies that occurs in the
and P. geoffroanus tuberosus, but McCord Guianas has ‘‘the dark pigment on the
et al. (2001) concluded that the name yellow carapace. . .restricted to the bor-
Phrynops tuberosus applies. der of the seam separating the vertebrals
and pleurals and to an incomplete band
extending on each side of the medial
Platemys platycephala (Schneider, 1792) groove downward through the 2d and
Plate 38A 3d pleurals to the lateral carapacial rim
(the medial groove remains yellow); the
Type material.—The original name was dark bar crosses less than 80% of the
Testudo platycephala and type material is bridge.’’
unknown to us. The type locality was
restricted to Cayenne, French Guiana by
Ernst (1983). Cheloniidae
Distribution.—This species is ‘‘restricted
to northern South America where it occurs These are sea turtles, with flipper-like
in the Caribbean drainages of Venezuela arms and legs (see also Dermochelyidae,
below). Three of the species in this family western Africa and Argentina. It also
are listed by the IUCN (2010) as being occurs in the Caribbean and Mediterra-
globally Endangered or Critically Endan- nean seas’’ (Ernst & Barbour 1989:125).
gered, owing to overharvesting by humans, Vouchers for Guyana.—PCHP 5697
and the fourth is listed as Vulnerable– from Guyana (no further data).
Decreasing (see below, Endangered Spe- Coloration in life.—The following is
cies). Analyses of sequence data of mito- from Ernst & Barbour (1989:124). ‘‘The
chondrial DNA in Brazil (Lara-Ruiz et al. carapace is reddish brown but may be
2006) have shown that reasonably frequent tinged with olive; the scutes are often
hybridization occurs between Eret- bordered with yellow. Bridge and plastron
mochelys imbricata and two other species, are yellow to cream colored. . .The head
Caretta caretta and Lepidochelys olivacea. varies from reddish or yellow chestnut to
Perhaps, if females of a species become olive brown, often with yellow-bordered
depleted while nesting, the more abundant scales. . .Limbs and tail are dark medially
males offshore will have proportionally and yellow laterally and below.’’
more females of other species with which Comments.—This species is not known
to mate; this phenomenon of unusual to nest in Guyana, but occurs ‘‘as an
hybridization may be a byproduct of o c c a s i o n a l s t r a g g l e r ’’ ( P r i t c h a r d
human predation. 1964:26). The other four species of sea
These turtles spend nearly 100% of their turtles documented for Guyana are
lives of many decades using vast areas of known to nest on the beaches northwest
the world’s oceans, mostly in the tropics, of Georgetown (Pritchard 1964, 1966,
at times moving from one continent to 1969, 2005).
another, although we still have much to
learn about their navigation mechanisms
and life histories. Four of the five species Chelonia mydas (Linnaeus, 1758)
of sea turtles known to occur in Guyana Plate 38B
are known to nest in very limited areas on
and near Shell Beach, to the northwest of Type material.—The original name was
Georgetown (Pritchard 1964, 1966, 1969, Testudo Mydas based on type material
2005). After a hatchling emerges on the (unknown to us) from Ascension Island
beach in Guyana and scrambles to the sea, (Smith & Taylor 1950b).
where does it spend the rest of its life, Distribution.—This species occurs in the
including the 20 yr or so that may be Atlantic, Pacific, and Indian oceans, pri-
required simply to reach maturity? marily in the tropics (Ernst & Barbour
1989).
Vouchers for Guyana.—The following
Caretta caretta (Linnaeus, 1758) specimens are primarily skeletal remains
found dead on the beach. PCHP 0921 and
Type material.—The original name was 0923 from Shell Beach; PCHP 2027 from
Testudo Caretta, the type material is NW District, 8 mi E Waini Point; PCHP
unknown to us, and the type locality was 2460–2462, 2469, 2504–2505, and many
restricted Bimini, British Bahamas by other PCHP specimens from NW District,
Schmidt (1953). Almond Beach.
Distribution.—This species occurs in the Coloration in life.—The following is
‘‘Pacific, Indian, and Atlantic oceans from from Ernst & Barbour (1989:120). ‘‘Car-
Washington, Japan, India, Kenya, the apacial scutes are olive to brown and may
British Isles, and Newfoundland south to contain a mottled, radiating, or wavy
Chile, Australia, South Africa, tropical pattern. . .The [plastron is] immaculate
white or yellow. . .All skin is brown or, Distribution.—These turtles occur in

sometimes, gray to black, and many head ‘‘. . .tropical waters of the Pacific and
scales may have yellow margins.’’ Indian Oceans from Micronesia, Japan,
India, and Arabia south to northern
Australia and southern Africa; in the
Eretmochelys imbricata (Linnaeus, 1766) Atlantic Ocean off the western coast of
Plate 38C Africa and the coasts of northern Brazil,
French Guiana, Surinam, Guyana, and
Venezuela in South America; and, occa-
Testudo imbricata and the type material is
sionally, in the Caribbean Sea as far north
unknown to us. The type locality was
as Puerto Rico. In the eastern Pacific it is
restricted to Belize by Schmidt (1953:106).
found from the Galapagos northward to
Distribution.—This species occurs in the
California’’ (Ernst & Barbour 1989:127).
‘‘Atlantic, Pacific, and Indian oceans from
Vouchers for Guyana.—The following
California, Japan, the Red Sea, the British
specimens are primarily skeletal remains
Isles, and Massachusetts south to Peru,
found dead on the beach. PCHP 0078,
Australia, Madagascar, northwestern Afri-
0961–0977, 1005, and other PCHP speci-
ca, and southern Brazil’’ (Ernst & Barbour
1989:123). mens from Shell Beach; PCHP 2325–2328,
Vouchers for Guyana.—The following 2503, 3182–3185, and other PCHP speci-
specimens are primarily skeletal remains mens from NW District, Almond Beach.
found dead on the beach. PCHP 0817, Coloration in life.—The following is
0951–0957, and 3880 from Shell Beach; from Zug et al. (1998:653.1). ‘‘General
PCHP 2025 from NW District, 10 mi E coloration is grayish-green to olive-brown
Waini Point; PCHP 2158–2162, 2501, dorsally. The carapace and upper surfaces
2506–2507, and many other PCHP speci- of the limbs and head are uniformly
mens from Almond Beach; and USNM pigmented, with the skin somewhat grayer
163069 from Enmore Beach (06844 0 00 00 N, than the shell. The plastron, bridge, and
057859 0 00 00 W). undersides of the limbs and head are
Coloration in life.—The following is creamy white.’’ Ernst & Barbour
from Ernst & Barbour (1989:123). ‘‘The (1989:127) referred to the bridge and
carapace is dark greenish brown; in the plastron as being ‘‘greenish white or
young it shows a tortoise-shell pat- greenish yellow.’’
tern. . .[the bridge and plastron are] yel-
low. . .Head scales are black to chestnut
brown at the center and lighter at their Dermochelyidae
margins; the jaws are yellow with some
brown streaks or bars. . .The chin and As the Cheloniidae (see above), these
throat are yellow, and the neck dark are sea turtles with flipper-like limbs that
above.’’ Juveniles may have a few dark come ashore in limited areas of coastal
spots on the plastron. Guyana to lay eggs. The single species of
this family is listed by the IUCN (2010) as
being globally Critically Endangered, ow-
Lepidochelys olivacea (Eschscholtz, 1829) ing to overharvesting by humans. These
Plate 38D turtles use vast areas of ocean, at times
moving from one continent to another,
Type material.—The original name was although we still have much to learn about
Chelonia olivacea and the type material, their navigation mechanisms and life
from Manila Bay, Philippines, is lost (see histories. The species of this family is
Zug et al. 1998). called the leatherback, because the outer
surface of its shell is not bone-hard, but Bataguridae (see Shaffer et al. 1997) and
leather-like to the touch. This is the turtle Emydidae, but Fritz & Havaš (2006)
that attains the greatest weight (ca. 867 kg) referred to these and related turtles as the
and longest carapace (ca. 2.4 m) in the Geoemydidae; all other genera of this
world (Ernst & Barbour 1989). family occur in ‘‘northwest Africa, Europe
to western Asia and the Middle East,
across southern Asia to China, Japan, the
Dermochelys coriacea (Vandelli, 1761) Philippines, and islands of the Sunda
Plate 38E Shelf’’ (Pough et al. 2004:109).

Testudine coriacea and ‘‘the specimen in Rhinoclemmys punctularia (Daudin, 1802d)
the Museum of the Padova University, Plate 38F
redescribed and figured by Fretey & Bour
(1980), is the holotype and only type- Type material.—The original name was
specimen of this species’’ (Bour & Dubois Testudo punctularia and the holotype, from
1983:358). The type locality is ‘‘Lauren- Cayenne, French Guiana, is MNHN 9130
tum, between Lido di Ostia and Tor (Fretey et al. 1977).
Paterno, shore of the Tyrrhenian Sea, Distribution.—This species occurs
Italy’’ (Bour & Dubois 1983:359). ‘‘. . .from eastern Colombia, the Orinoco
Distribution.—This species ‘‘ranges drainage of Venezuela, and Trinidad Island
throughout the waters of the Atlantic, eastward through the Guianas and north-
Pacific, and Indian Oceans from Labrador, eastern Brazil’’ (Ernst & Barbour 1989:182).
Iceland, the British Isles, Norway, Alaska, Vouchers for Guyana.—AMNH R-8080
and Japan south to Argentina, Chile, from Kalacoon; AMNH R-61530 from
Australia, and the Cape of Good Hope Shudikar-wau River; PCHP 2878 from
(Pritchard, 1980). It also enters the Med- Barama River, Cariaco Mission; PCHP
iterranean Sea’’ (Ernst & Barbour 4802 from Almond Beach (near Waini
1989:117). Point); USNM 85022 and 86861 from
Vouchers for Guyana.—The following Pomeroon; and USNM 164180 from
specimens are primarily skeletal remains Mabaruma Compound (08812 0 00 00 N,
found dead on the beach. PCHP 1021 from 059847 0 00 00 W).
Shell Beach; PCHP 2007 from 8 mi W Coloration in life.—The following is from
Waini Point; PCHP 2467–2468, 3055– Ernst & Barbour (1989:182). Adults dark
3058, and 3876 from NW District, Almond brown or black above, ‘‘but juveniles may
Beach; PCHP 4792 from Gwennie Beach; have yellow to bronze radiations on each
and PCHP 4793 from Kanwalla Beach. pleural [plate]...The plastron is red brown to
Coloration in life.—The following is black with a yellow border and seams. The
from Ernst & Barbour (1989:117). Cara- bridge is yellow with two large dark
pace brown to black; plastron cream to blotches. . .The head is. . .black with. . .two
white; head and neck dark brown to black longitudinal red or yellow stripes. . .or. . .a
with white to yellow blotches; upper jaw broad horseshoelike mark posterior to the
gray; limbs black with some light blotches. orbit. Two light spots may occur on the
nape. . .and stripes usually run between the
Geoemydidae orbit and tympanum and from the snout
along the upper jaw to the tympanum. The
In the past, turtles of the genus Rhino- iris is green to bronze. Forelimbs have large,
clemmys were included in the families yellow or red, black-spotted scales, and the
hind limbs are gray laterally and yellow with Comments.—‘‘This species lives in
black spotting medially.’’ streams, rivers, lakes, and ponds. If its
waterway dries up, it will bury itself in the
mud bottom until the next rain. . .[It] is
Kinosternidae
omnivorous’’ (Ernst & Barbour 1989:84).
This family of aquatic turtles is restrict-
ed to North and South America. Podocnemididae
These are large, aquatic, side-necked turtles
Kinosternon scorpioides (Linnaeus, 1766)
that swim well, feed mostly on vegetation, live
Plate 39A, B
in rivers and lakes, and nest on sand banks,
Type material.—The original name was usually at night. The family occurs only in
Testudo scorpioides and the holotype, from South America and Madagascar. Two species
Surinam, has apparently been lost (see occur in Guyana, and both need conservation
Berry & Iverson 2001). management activities.
Distribution.—This species ‘‘ranges at
low elevations from southern Tamaulipas, Podocnemis expansa (Schweigger, 1812)
Mexico, southward to northern Argentina, Plate 39C
Bolivia, and northern Peru’’ (Ernst &
Barbour 1989:83). Type material.—The original name was
Vouchers for Guyana.—PCHP 0222 and Emys expansa and there are three types in
0431 from the Uitvlugt Sugar Estate; PCHP the MNHN, from ‘‘America meridionali.’’
2669 from NW District, 5 km SE Waini Distribution.—This species occurs in
Point; PCHP 3709 from NW District, ‘‘Caribbean drainages of Guyana and
Almond Beach; PCHP 4221 from NW Venezuela and in the upper Amazon
District, behind Almond Beach; and PCHP tributaries in Bolivia, Peru, Colombia,
Venezuela, and Brazil. It is also occasion-
5083–5113 and many other PCHP speci-
ally found on Trinidad, especially after
mens discovered dead in the NW District,
floods of the adjacent mainland Orinoco
Avicennia forest behind Almond Beach, 6
River’’ (Ernst & Barbour 1989:27) and is
km E Waini Point (died in a fire); and
found widely on the Amazon River, Brazil
USNM 84582–84585 from Pomeroon.
(M. S. Hoogmoed, pers. comm.).
Coloration in life.—The following is Vouchers for Guyana.—The following
from Berry & Iverson (2001:725.3). ‘‘The are primarily skeletal remains of specimens
carapace is highly variable in color, found dead. PCHP 2779 from Rupununi
ranging from light brown to olive to black, River, Massala; PCHP 2827 from Rupu-
with darker seams in all but the darkest nuni River, Karanambo; PCHP 2884–
individuals. . .The color of the plastron 2886, 3133–3134, and 4132 from Rupunu-
may be gray, yellow, orange, brown or ni River, Apoteri; PCHP 4128 and 4148
black, usually with darker seams. . .The from Rupununi River, Rewa mouth; and
color pattern on the head is extremely PCHP 4129 from Kwattanmang (near
variable, with a brown, gray, or black Annai), ex. Rewa River.
background and a reticulated or spotted Coloration in life.—The following is
pattern of cream, yellow, orange, pink, or from Ernst & Barbour (1989:26–27).
red. The jaw sheaths are cream to yellow ‘‘The carapace is olive to dark gray or
with darker vertical streaks most conspic- brown and may have some dark spots and
uous in older males. The skin of other soft a light border in younger individuals. The
parts is gray or brown, usually with many plastron. . .bridge, and undersides of the
small, darker spots.’’ marginals are yellow. . .The head is gray
brown with yellow markings. Two yellow Coloration in life.—The following is

spots occur on the interparietal scutes, and from Ernst & Barbour (1989:29). ‘‘The
one on each side of the head; these spots carapace of juveniles is brown to greenish
fade with age. Jaws are tan; the chin gray with a narrow yellow border, that of
yellow. The neck is gray dorsally and adults is olive to dark gray or brown. . .The
yellow ventrally. . .; limbs are gray.’’ yellow plastron and bridge may develop
Comments.—These giant river turtles dark blotches with age. . .The head is gray
have been overharvested by humans for to olive or brown with yellow spots–one on
consumption of eggs, meat, and oil. top of the snout, one on each side of the
Consequently, populations have experi- snout extending to the upper jaw rim,
enced dramatic declines. Analyses of mi- another on each side of the head extending
tochondrial DNA sequence data and from the lower posterior edge of the orbit
nuclear microsatellites indicate that fe- to the corner of the mouth, and one on
males practice natal-river homing and each tympanum. Jaws are dark brown or
‘‘that each major tributary [sampled] cur- black, but the chin has a transverse yellow
rently forms a semi-isolated reproductive bar and a yellow spot on each side below
population and should be managed ac- the corner of the mouth. Limbs are gray to
cordingly’’ (Pearse et al. 2006:985). olive brown.’’
Podocnemis unifilis Troschel, 1848 Testudinidae

Plate 39D These are the terrestrial tortoises with
hard, high-domed shells and elephantine
Type material.—The holotype, from
arms and legs. They are frequently preyed
Guyana, was ZMB 142 but is now lost
upon by humans. Until recently, the two
(Duellman 2005).
species known to occur in Guyana were
Distribution.—This species occurs in
referred to the genus Geochelone, but now
‘‘Caribbean drainages of the Guianas,
they are referred to Chelonoidis.
Venezuela and Colombia, and the upper
tributaries of the Amazon River in Co-
lombia, Ecuador, Peru, northern Bolivia,
southern Venezuela, and Brazil’’ (Ernst & Chelonoidis carbonaria (Spix, 1824)
Barbour 1989:29). It is also found widely Plate 40A
on the Amazon River, Brazil (M. S. Type material.—The original name was
Hoogmoed, pers. comm.). Testudo carbonaria based on material from
Vouchers for Guyana.—The following Brazil. Hoogmoed & Gruber (1983:354)
are primarily skeletal remains of specimens designated ‘‘pl. XVI in Spix (1824) as the
found dead. AMNH R-8707 from Kala- lectotype.’’ Vanzolini (1994:1–2) stated
coon; AMNH R-15144 from Kartabo; that the type locality probably is ‘‘the
AMNH R-61524–61525 from Wichibei; stretch of the Rio Amazonas between the
AMNH R-61526 from Yupicari; PCHP mouth of the Negro, at 03808 0 S, 59855 0 W,
2775 and 2777 from Rupununi River, and the mouth of the Furo do Tajapuru, at
Massala; PCHP 2776 from Rupununi 01802 0 S, 5182 0 W.’’
River, Karanambo; PCHP 2882 and 4131 Distribution.—This species occurs in
from Rupununi River, Apoteri; PCHP ‘‘southeastern Panama and west of the
2883, 4134, and 4137–4138 from Rupunu- Andes in Chocó of Colombia, but its
ni River, mouth of Rewa River; and PCHP main range is east of the Andes in eastern
5202–5203 from the Ireng River. Colombia, Venezuela, and the Guianas
to eastern Brazil, south to Rio de ribbean lowlands of the Guianas to Brazil,

Janeiro, and west to eastern Bolivia, where it occurs throughout the Amazon
Paraguay, and northern Argentina. It Basin to eastern Ecuador and Colombia,
seems absent from almost all but the northeastern Peru, and northern and
eastern parts of the Amazon Basin. This eastern Bolivia and on Trinidad’’ (Ernst
tortoise may occur naturally on Trinidad, & Barbour 1989:254).
and has been introduced on quite a few Vouchers for Guyana.—The following
Caribbean Islands, including St. Croix in are primarily skeletal remains of specimens
the Virgin Islands’’ (Ernst & Barbour found dead. AMNH R-15143 from Kar-
1989:253). tabo; AMNH R-25046 and R-44617–
Vouchers for Guyana.—The following 44618 from Kamakusa; AMNH R-
are primarily skeletal remains of specimens 128195 from Essequibo, Takutu Mts. (ca.
found dead. AMNH R-61522 from Head 45 mi SE Itabali on W bank of Mazuruni
of Rupununi River; PCHP 2790–2801 River); PCHP 2108–2116 from NW Dis-
from Rupununi, Yupakari; PCHP 2802 trict, Morawhanna (reportedly from lower
and 4146 from St. Cuthbert’s Mission; Waini); PCHP 2181–2182 and 2306 from
PCHP 3052 from Mabaruma; USNM NW District, Mabaruma area; PCHP 2491
85024 from ‘‘Georgetown, 70 mi from’’; from Moruka River, Santa Rosa Mission;
USNM 291193 from Paramakatoi; and PCHP 2650–2651 from NW District,
USNM 291194 from Wandapa (04832 0 Mabaruma; PCHP 2832–2833 from Bar-
00 00 N, 059843 0 00 00 W). In addition, CJC ama River, Cariaco (¼Kariakau) Mission;
and CRT saw specimens (not collected)
PCHP 2834–2835 from NW District,
at Dubulay Ranch, Karanambo, and
Baramanni (Waini/R. Baramanni junc-
Aishalton.
tion); PCHP 2886–2889 from Apoteri
(Rupununi/Essequibo river junction);
from Ernst & Leuteritz (1999a:690.1).
PCHP 2892 from Barama River, Iroma;
‘‘The carapace is black, with the vertebral
PCHP 2979 from Barama, Towaka; PCHP
and pleural areolae yellow to reddish
orange, and a light spot of the same color 3053 from Rewa Mouth; PCHP 5333–5338
occurs at the base of each marginal. . .The from Santa Rosa, Moruka River; and
plastron is yellowish brown with some USNM 535818 from Baramita. In addi-
dark pigment along the mid- and trans- tion, CJC and CRT saw specimens (not
verse seams. . .The head. . .scales are. . .yel- collected) at Berbice River Camp and
low, red, or orange; the jaws are dark. The Konawaruk Camp.
anterior surface of each forelimb is cov- Coloration in life.—The following is
ered with large, red, and slightly or non- from Ernst & Leuteritz (1999b:691.1).
overlapping scales.’’ ‘‘The carapace is brown with yellow to
orange vertebral and pleural areolae;
yellowish or orange pigment also occurs
at the lower edge of each marginal. . .The
Chelonoidis denticulata (Linnaeus, 1766)
plastron is yellowish brown with darker
Plate 40B
pigment along the seams. The head is
Type material.—The original name was brown with yellow dorsal scales. . .
Testudo denticulata and the holotype is The. . .head scales are. . .yellow to orange
NRM De Greer 21 (see Ernst & Leuteritz with dark borders; the jaws are dark
1999b). There are no reliable data on the brown. The anterior surface of each
type locality. forelimb is covered with large yellow or
Distribution.—This species occurs in orange, non-or only slightly overlapping
‘‘southeastern Venezuela through the Ca- scales.’’
Species Reported from Guyana but Kartabo and the same number at
Probably in Error Caripito [Venezuela].’’ The more de-
tailed notes he presented in the publica-
Tretioscincus bifasciatus (Duméril, 1851) tion were specifically based on
specimens from Caripito, and he gave
This species was described on the basis
no indication that he was aware of the
of a specimen from the vicinity of Made-
possibility that a different species, but
leine (Nouvelle Grenade), Colombia (Pe-
similar congener, Tretioscincus agilis,
ters & Donoso-Barros 1970:262). The does occur in Guyana and probably at
general area of distribution is in arid Kartabo, although voucher specimens
habitats on the Caribbean coasts of are needed to confirm this. Future
Colombia and Venezuela, and on Isla workers should not list T. bifasciatus
Margarita and Dutch Leeward Islands as a species that occurs in Guyana until
(Gorzula & Señaris 1999). it is confirmed and documented. Simi-
We do not believe that this species larly, the presence of T. agilis at
occurs in Guyana despite the existence of Kartabo should be confirmed.
one published report (Beebe 1945:23) and
one AMNH specimen (R-137364), for
which the catalog states ‘‘Guyana: [Ma-
zaruni-Potaro]: Kartabo: 6821 0 N 58841 0 Liophis juliae (Cope, 1879)
W.’’ A type-written note inside the jar The AMNH has two specimens of
states: ‘‘Near Kartabo, British Guiana.’’ Liophis juliae (R-6804–6805) that are noted
Another type-written note in the jar states: as being from ‘‘Kaieteur.’’ These were
‘‘Tretioscincus bifasciatus 3116. Kartabo.’’ identified as L. juliae by J. R. Dixon, who
However, there is no field tag or label stated ‘‘I recently examined two specimens
attached to the specimen itself. We suspect from the American Museum of Natural
that the locality data associated with the History that were labeled from Guyana. I
specimen are incorrect. believe these to be mislabeled specimens’’
The AMNH specimen was recorded (Dixon 1981:303). No other specimens of
as having been collected by W. Beebe in this species are known from Guyana, and
1924, but it was not cataloged at the all other known specimens are from islands
AMNH until 1991. The identification in the West Indies (type locality, Domin-
was confirmed by CJC as correct on 9 ica). The collector of the AMNH specimens
May 2007. However, there could be a is given in the catalog as ‘‘Crampton.’’
problem with the association of the The AMNH specimens cataloged as
specimen and the locality data. We being from Guyana compare favorably
think that the occurrence of this species with specimens of this species from Dom-
in Guyana is doubtful for the following inica (AMNH R-32889, R-44899–44900,
reasons: no other specimens are known and R-135273), as recently determined by
for Guyana; the species is present in CJC, which agrees with Dixon’s earlier
Venezuela where Beebe collected also; identification. Additional herpetological
the 67 yr delay in cataloging from the specimens at the AMNH were collected
date of collection may reflect problems by ‘‘Crampton’’ in 1911, including speci-
in transfer of data; and there are other mens from Kaieteur that are credible. This
problems associated with some of collector presumably was Henry E. Cramp-
Beebe’s specimens (Barbour 1920). Fur- ton, who was at the time, among other
thermore, Beebe (1945:23) stated the things, a curator of invertebrates at the
following: ‘‘captured or saw only five AMNH. Prior to collecting in Guyana, his
or six [individuals of T. bifasciatus] at field party stopped and collected in the
West Indies, apparently in transit to Guya- some individuals appearing to be of the

na, and they collected in Dominica, which is circinalis type after all.
the type locality for L. juliae. In addition,
Philippe Kok (pers. comm.) stated that
Typhlops lumbricalis (Linnaeus, 1766)
despite his extensive recent field work in
Kaieteur National Park, he has seen no Although Hoogmoed (1983:235) said
specimens that appear similar to L. juliae. that this is the only species of snake to
We conclude that it is likely that there was a have been successfully introduced ‘‘into
mix-up in locality data for AMNH R-6804– Guyana from the Antilles,’’ Dixon &
6805 and they probably were collected in Hendricks (1979) cite Thomas’ dissertation
Dominica, not at Kaieteur. (1976) as being doubtful of the only known
record for Guyana. The only reference
specimen is AMNH R-67881, cataloged as
Micrurus circinalis having been collected by W. Beebe at
(Duméril & Bibron, 1854) Kartabo, and CJC recently confirmed the
identification of the specimen. Considering
Donnelly et al. (2005:457) reported that that occasionally problems exist for spec-
this species was found at Iwokrama, but imens collected by Beebe (Barbour 1920;
we could not obtain voucher specimens to and see above discussion for Tretioscincus
examine for confirmation. This is of bifasciatus), and considering the lack of
interest as M. circinalis is known only additional specimens from Guyana, we
from Trinidad and Venezuela (Campbell & suggest that the record needs to be
Lamar 2004:157). It is intriguing also, as confirmed before concluding that the
AMNH R-110163 appears to be an species occurs in Guyana. It seems more
example of M. circinalis, and it is cata- likely to us that this is another case of
loged as being from Kartabo (collected by mislabeling of specimens.
W. Beebe). As cited several times in this
volume (and see account above for Tretio-
scincus bifasciatus), we are reluctant to Additional Species Possible
accept records of Beebe that are open to (As Yet Undocumented)
question, especially for taxa that occur in
Venezuela, where he collected also, and for New Species
which there was a long delay before his Considerable research remains to be
specimens were received or cataloged at done on the amphibians and reptiles of
the AMNH. Interestingly, the Beebe spec- Guyana. To date, the herpetofauna of
imen had a paper tag tied to it saying Guyana includes at least 324 species (148
‘‘Kartabo,’’ number ‘‘3263,’’ and ‘‘Dept. amphibians and 176 reptiles; Appendix
Tropical Research of the N.Y. Zoological 2). Of these, 39 (12%) were described and
Society.’’ As an alternative suggestion, one named since the year 2000. Most of the
might question the status of our knowl- species described since 2000 are amphib-
edge about intraspecific variation in M. ians (17% of the amphibian fauna), most
psyches, of which circinalis was once are from remote, isolated highland sites
treated as a subspecies. New studies based (see Biogeography, below), and most are
on larger collections, especially including endemic species that occur nowhere else
specimens from strategically important in the world, except possibly in immedi-
localities, may show that M. psyches is ately adjacent uplands and isolated high-
more variable than we recognize today, lands of small areas of Venezuela and
perhaps dimorphic in coloration, with Brazil. Considering that 12% of the
herpetofauna (17% of the amphibians, Discussion

6% of the reptiles) were described be- Biogeography
tween the years 2000–2011, undoubtedly
additional undescribed species, including Comparisons of Faunas at Different Sites
endemics, are yet to be discovered. We
There are several reasons to compare
expect that most of the new species will
the lists of species that occur at different
still be found in the uplands and isolated
localities. Academics often use patterns of
highlands of the Pakaraima Mountains,
faunal similarities and differences to dis-
but discoveries are to be made also in the
cover regions of historical significance in
lowlands (e.g., Jungfer & Böhme 2004,
evolution and biogeography, and entrepre-
Kok 2005, Cole & Kok 2006, Donnelly et
neurs often use unique biotas for ecotour-
al. 2006, Kok & Ernst 2007).
ism. The Guianan Region is recognized as
having a fauna that is distinctive, large,
and diverse among those of other regions
Other Species in South America or Central America.
This results from the geographic position
There are species known to occur in
of Guyana in Neotropica and on the
countries near Guyana but which are not
Guiana Shield, including the isolated
yet known to occur in Guyana. Candidate
highlands or tepuis of the eastern part of
species that may well be found in Guyana
the Pantepui Region, which are surround-
in the future are listed below in the same
ed by lowland rainforest and savannas.
sequence as representatives of their fam-
Consequently, there is a mixture of local
ilies are listed in the species accounts
endemic species and widespread species
above. The species that might some day
characteristic of low elevation habitats of
be found in Guyana, however, are not
Amazonia and the Guianan Region. In the
strictly limited to those on these lists. It is
vicinity of the Pakaraima Mountains, the
not surprising that the majority of the
surrounding lowland fauna usually is quite
species listed are snakes, as these can be
different from adjacent isolated highland
most difficult to find and capture in
fauna. Comparing faunas is an important
tropical environments. Consequently,
step in learning what patterns exist, which
snakes usually are the most poorly repre-
can be helpful in designating national
sented group in scientific collections from parks or other preserves. Prior to compar-
the tropics. ing faunal lists, however, it helps to know
whether the lists of local species are
Amphibians Reptiles
reasonably complete.
Vitreorana oyampiensis Amapasaurus tetradactylus

Dendropsophus Arthrosaura kockii
luteoocellatus Estimating Effectiveness and
D. melanargyreus Atractus elaps Completeness of Sampling
Hypsiboas tepuianus A. flammigerus
Phyllomedusa tomopterna A. latifrons How do investigators know whether the
Teratohyla midas A. zidoki specimens they have studied are sufficient
Leptodactylus myersi Cercophis auratus
Microcaecilia trombetas Helicops leopardinus to accurately represent the biodiversity
Mastigodryas bifossatus that occurs at a particular locality? Is the
Taeniophallus brevirostris faunal list complete or not? Various
T. nicagus authors have discussed methods to esti-
Xenodon werneri
Mesoclemmys nasuta
mate this, and we found a good model in
recent works on the mammals of Paracou,
French Guiana (Simmons & Voss 1998, only two field workers (CJC and CRT).
Voss et al. 2001). For Berbice River Camp (1997), we spent
One method is to estimate effectiveness 28 nights and the other conditions were
of sampling effort by keeping track of how similar, except that we also had the
many people hours (or days, weeks, assistance of two American Indians, Ger-
months) were spent in the field and how ald and Wesley King, who were outstand-
many species were collected for the first ing in the field. At Dubulay Ranch in 1994,
time at the site during each period of time we recorded 46 species, of which 19 were
(also see Ernst et al. 2005). The resulting singletons and 8 were doubletons, which
cumulative number of species (vertical resulted in an estimated fauna of 69 species
axis) is plotted against the period of time of amphibians and reptiles. By this esti-
(horizontal axis), and over time the plotted mate, we recorded only 67% of the fauna
line approaches an asymptote. The results around the ranch house. At Berbice River
of this approach with mammals varied Camp, we recorded 65 species, of which 14
widely among analyses of six different were singletons and 4 were doubletons,
collecting techniques at Paracou (Voss et which resulted in an estimated fauna of 90
al. 2001:167–177). We could not use this species of amphibians and reptiles. By this
method because we did not track observa- estimate, we recorded only 72% of the
tions in ways that were consistent with fauna. The results for the first sampling at
drawing such plots (e.g., how many people Dubulay Ranch and Berbice River Camp
hours or days were spent collecting, as indicate that our sampling was far from
opposed to writing notes and photogra- complete. This conclusion is supported by
phy?). Furthermore, we had very unequal simply scanning Table 1. For example,
expenditure of efforts in different micro- incompleteness of sampling probably ex-
habitats (e.g., canopy, aquatic, subterra- plains the following: apparent absence of
nean) and in some areas we used pit-fall Allobates spumaponens at Baramita and
traps with drift fences, but not everywhere. Kartabo; apparent absence of Rhinella
Another method for estimating com- marina at Konawaruk Camp; presence of
pleteness of the local sample (e.g., Voss et Ceratophrys cornuta at only one of the
al. 2001:177–179), is to apply ‘‘Chao’s sites; absence of Hypsiboas boans at
(1984) estimator, which is based on the Aishalton; presence of Hypsiboas crepitans
total number of observed species, Sobs, the at only two sites; presence of Pipa pipa at
number of singletons (species recorded only two sites; and relatively few caecil-
only once), a, and the number of double- ians, amphisbaenians or worm lizards,
tons (species recorded only twice), b. The snakes, and turtles in general. We conclude
expected total number of species, S*, that considerably more effort is needed to
is. . .given by the expression S* ¼ Sobs þ accurately document the faunas at all the
(a2/2b).’’ sites we visited.
We performed this exercise for the two Although the known herpetofauna for
lowland sites that seemed most appropri- Kartabo is much larger than that for either
ate: Dubulay Ranch, and Berbice River Dubulay Ranch or Berbice River Camp,
Camp (see Methods for details on the we could not calculate S* because we lack
localities). For Dubulay Ranch, we used data on the numbers of singletons and
observations for only the first sampling doubletons observed by Beebe and his
period, 1994, when we spent 26 nights, associates. However, we think the appar-
used drift fences and pitfall traps, did most ently larger fauna at Kartabo may be
of our work within about 1 km (linear) of misleading, reflecting the more extensive
the ranch house, and during which there time over eight field trips that Beebe and
was good rainfall, although there were his colleagues spent there (Beebe 1946).
Table 1.—Herpetofaunal comparisons of seven lowland sites in Guyana. X indicates presence of the species, –
indicates not documented. The sites are as follows: Ai, Aishalton; Ba, Baramita; Be, Berbice River Camp; Du,
Dubulay Ranch; Ka, Karanambo; Ko, Konawaruk Camp; Kt, Kartabo. The habitats are as follows: F, forest;
M, patchwork of forest, savanna, and gallery forest; S, savanna (but with gallery forest).
Taxon BaF KtF KoF DuM BeF KaS AiS
Allobates femoralis X X – – – – –
Allobates spumaponens – – X – X – –
Anomaloglossus kaiei – – X – – – –
Atelopus spumarius – X X – – – –
Dendrophryniscus minutus – – – – X – –
Rhaebo guttatus – X X X X X –
Rhaebo nasicus – X – – – – –
Rhinella merianae – X – X – – X
Rhinella marina X X – X X X X
Rhinella martyi X X X X X – –
Ceratophrys cornuta – X – – – – –
Pristimantis zeuctotylus – – – – X – –
Ameerega hahneli X X – – – – –
Ameerega trivittata X X – – – – –
Dendrobates leucomelas X X – – – – –
Stefania evansi – X X – – – –
Dendropsophus brevifrons – – X – – – –
Dendropsophus marmoratus X X X – – – –
Dendropsophus minusculus X – – X – X X
Dendropsophus minutus – – X X – – –
Hypsiboas boans X X X X X X –
Hypsiboas calcaratus – – X – – – –
Hypsiboas cinerascens X X X – X – –
Hypsiboas crepitans – – – X – – X
Hypsiboas geographicus X X X X X – –
Hypsiboas multifasciatus X X X X – – –
Lysapsus laevis – – – – – X X
Osteocephalus buckleyi – X – – – – –
Osteocephalus cabrerai – – X – – – –
Osteocephalus leprieurii – X X – X – –
Osteocephalus oophagus – – X – X – –
Osteocephalus taurinus X X X X X – –
Phyllomedusa bicolor X X – – – – –
Phyllomedusa hypochondrialis – X X X X – –
Phyllomedusa vaillantii X – – – X – –
Pseudis paradoxa – X – – – – –
Scinax boesemani – – – X – – –
Scinax nebulosus – – – X – – –
Scinax ruber X X – X – X –
Scinax cf. x-signatus X – – X – – X
Scinax sp. – – – – X – –
Trachycephalus coriaceus – – – – X – –
Trachycephalus typhonius – X – X – – –
Adenomera andreae – – – – X – –
Adenomera hylaedactyla X X – X X X –
Leptodactylus fuscus – X – X – X X
Leptodactylus guianensis – X X X X X –
Leptodactylus knudseni X X X X X – –
Leptodactylus leptodactyloides – X – – – – –
Leptodactylus longirostris – X – – – – X
Leptodactylus macrosternum – – – – – X X
Leptodactylus mystaceus X X X X X – –
Table 1.—Continued.
Leptodactylus petersii – X X X X X –
Leptodactylus rhodomystax – X X – X – –
Leptodactylus rugosus – X – – – – –
Leptodactylus validus X X – X – X –
Lithodytes lineatus X X X X X – –
Physalaemus cuvieri X – – X – – –
Pleurodema brachyops – – – – – X X
Pseudopaludicola boliviana – – – – – X X
Chiasmocleis jimi – – – – X – –
Ctenophryne geayi X X – X X – –
Elachistocleis surinamensis – – – X – – X
Otophryne pyburni – X – – – – –
Synapturanus salseri – X X – – – –
Pipa arrabali X X – – – – –
Pipa pipa – X – – X – –
Lithobates palmipes – X X – – – X
Caecilia tentaculata – – – X – – –
Rhinatrema shiv – – X – – – –
Microcaecilia sp. – – X – X – –
Caiman crocodilus – X – X X – –
Melanosuchus niger – – – – – X –
Paleosuchus trigonatus – X X – – – –
Amphisbaena alba – X – – – – –
Amphisbaena fuliginosa – X – X – – –
Hemidactylus mabouia – X – X – – –
Hemidactylus palaichthus – – – X – X X
Thecadactylus rapicauda – X X X X – –
Alopoglossus angulatus – – – – X – –
Arthrosaura reticulata – – – – X – –
Bachia flavescens – X X X X – –
Cercosaura ocellata – X – X – – X
Echinosaura sulcarostrum X – – – – – –
Gymnophthalmus leucomystax – – – – – – X
Gymnophthalmus speciosus – – – X – – –
Gymnophthalmus underwoodi – – – X X X X
Gymnophthalmus vanzoi – – – – – X X
Iphisa elegans – X – – X – –
Leposoma percarinatum X X X X X – –
Neusticurus bicarinatus – X – – X – –
Neusticurus rudis – X X – – – –
Iguana iguana – X – X – X X
Anolis aeneus – X – – – – –
Anolis auratus – – – X – X X
Anolis fuscoauratus X X X – – – –
Anolis ortonii – X – – – – –
Anolis planiceps X X X – X – –
Anolis punctatus – X – – X – –
Polychrus marmoratus – X – X – X X
Mabuya nigropunctata X X X – X – X
Coleodactylus septentrionalis – – – – – X –
Gonatodes alexandermendesi – X X – – – –
Gonatodes annularis X X – – X – –
Gonatodes humeralis – X X X X – X
Pseudogonatodes guianensis – – X X – – –
Sphaerodactylus molei – X – – – – –
Ameiva ameiva X X X X X X X
Cnemidophorus lemniscatus – X – X – X X
Kentropyx calcarata X X X X X X –
Kentropyx striata – – – X – X X
Tupinambis teguixin – X X X – – –
Plica plica X X X – – – –
Plica umbra X X X X X – –
Tropidurus hispidus – – – X – X X
Uranoscodon superciliosus – X X X X X X
Anilius scytale – X X – X – –
Boa constrictor X X – X X – –
Corallus caninus – X X X – – –
Corallus hortulanus X X – X X – –
Epicrates cenchria – X X X – – –
Eunectes murinus – X X X – – –
Atractus torquatus – – – X – – –
Atractus trilineatus – X – X – – –
Chironius carinatus – – – – – X –
Chironius exoletus – X X – – – –
Chironius fuscus X X – – X – –
Chironius multiventris – X – – X – –
Chironius scurrulus – X X – X – –
Clelia clelia – X – – – – –
Dipsas catesbyi – X – – X – –
Dipsas indica – X – – – – –
Dipsas pavonina – X – – – – –
Dipsas variegata X X – – – – –
Drymoluber dichrous – X – – X – –
Erythrolamprus aesculapii – X – X – – –
Helicops angulatus – X – – X – –
Hydrops triangularis – X – – – – –
Imantodes cenchoa – X X – X – –
Imantodes lentiferus X – – – – – –
Leptodeira annulata – X – X – X –
Leptophis ahaetulla X X X X – X –
Liophis breviceps – X X X – – –
Liophis cobella – X – – – – –
Liophis lineatus – X – – – – –
Liophis miliaris X – – – – – –
Liophis poecilogyrus – – – – – X –
Liophis reginae – X – – – – –
Liophis typhlus X X – X – – –
Mastigodryas boddaerti X X – X X X –
Oxybelis aeneus – X – – – – –
Oxybelis fulgidus – X X – X – –
Oxyrhopus melanogenys – X – X X – –
Oxyrhopus petolarius X X – X – – –
Philodryas olfersii – – – X – – X
Philodryas viridissimus – X – – – – –
Phimophis guianensis – – – X – – –
Pseudoboa coronata – – – – X – –
Pseudoboa neuwiedii – – – X – – X
Pseustes poecilonotus – X X – – – –
Pseustes sulphureus – X – – – – –
Rhinobothryum lentiginosum X – – – – – –
Siphlophis compressus X X X X – – –
Tantilla melanocephala X X – X X – X
Thamnodynastes ramonriveroi – – – X – – –
Xenodon rabdocephalus – X – – – – –
Xenodon severus X X – – – – –
Micrurus hemprichii X – – – – – –
Micrurus lemniscatus X X – – – X –
Micrurus psyches – X – – – – –
Micrurus surinamensis – – – X X – –
Epictia albifrons X X – X X – –
Siagonodon septemstriatus – X – – – – –
Tricheilostoma dimidiatum – – – – – X –
Typhlops brongersmianus – – – X – – –
Typhlops minuisquamus – X – X – – –
Typhlops reticulatus – X – X – – –
Bothriopsis bilineata – X – – – – –
Bothrops atrox X X – X X – –
Crotalus durissus – – – X – X –
Lachesis muta – X – X – – –
Chelus fimbriatus – – – – – X –
Mesoclemmys gibba – X – X – – –
Platemys platycephala – X – – – – –
Podocnemis expansa – – – – – X –
Podocnemis unifilis – X – – – X –
Chelonoidis carbonaria – – – X – X X
Chelonoidis denticulata X X X – X – –
Total (n ¼ 188 species) 54 127 59 83 65 40 32
The Kartabo fauna appears to be closer in As Simpson (1960) pointed out, no one
biodiversity to that of Iwokrama (Don- equation would apply necessarily as the
nelly et al. 2005), but for Iwokrama many most appropriate for all comparisons. For
species remain unidentified and not avail- example, in ideal cases where the two
able for examination by us. faunas were reasonably well known (i.e.,
with little to no sampling error) and of
similar size, the appropriate faunal resem-
Estimating Faunal Resemblance Between blance index would be calculated as
Various Localities follows:
C=N1 þ N2 C 3 100;
How similar (or dissimilar) are the
faunas at two or more sites, such as where C ¼ the number of species found in
Dubulay Ranch and Berbice River Camp? common at both sites, N1 ¼ the number of
Simpson (1960) discussed alternative equa- species at one site, and N2 ¼ the number of
tions for calculating how similar two species at the other site. In this case,
faunas are. The calculations were based Simpson’s equation (1), the resemblance
on the total number of species known to index is ‘‘the percentage of taxa in com-
occur in each fauna and the number of mon among the total taxa of the two
species known to be shared in common faunas or samples in question,’’ with
between those faunas, and comparisons possibilities ranging from 0–100% (Simp-
were made on a pair-wise basis. son 1960:300).
However, in many cases, as in Guyana, tween isolated highland sites. Differences

sampling of localities may be so inade- in environments and habitats are so
quate that we do not have, for most sites, a extensive that there are very few species
complete inventory of the fauna, as dis- shared between the lowlands and isolated
cussed above. Sampling at one site or both highlands (see below), and this can be
may be so incomplete as to lead to very discussed effectively without calculating
misleading indices with the above equa- indices, especially based on the incomplete
tion. To correct for this, at least in part, data that prevail today. When sites of
Simpson (1960) suggested an alternative intermediate elevations (upland sites) are
index, calculated as follows: involved, overlaps in shared fauna appear
with both highland and lowland sites, but
C=N1 3 100;
none of these intermediate sites in Guyana
where C ¼ the number of species found in has been sampled sufficiently for such
common at both sites and N1 ¼ the number comparisons now. One such site, Kaieteur
of species at the site with the smallest National Park, is currently receiving con-
fauna known, or presumably most incom- siderable important and careful work,
plete sample. In this case, Simpson’s thanks to the efforts of Kok, Kala-
equation (2), the index is a percentage of mandeen, and their colleagues, but these
the smallest fauna sampled that is shared studies are still in progress (see Kok &
by both faunas, with possibilities ranging Kalamandeen 2008).
from 0–100%.
We used Simpson’s equation (2), the one
with correction for incomplete sampling, The Lowland Faunal Comparisons
to compare samples of faunas within
Guyana, despite the fact that many herpe- We selected seven lowland sites to
tologists follow Duellman (1965, 1979, and compare, although none has its herpeto-
elsewhere), who used a different equation. fauna completely known (see above). Each
The equation Duellman used drops off the site is discussed above in Methods (Major
% to give a fraction based on 2C/N1 þ N2, Collecting Sites), its location is shown on a
which is an alternative discussed by map (Fig. 4), and the herpetofauna known
Simpson as his formula (4), and for which for each is listed in Table 1. Note that this
Simpson (1960:303) stated ‘‘it is hard to see table does not list all of the lowland species
a meaningful use for (4).’’ For faunal known for Guyana, but only those species
comparisons, Voss et al. (2001:182) used known for the seven sites included in the
‘‘Jaccard’s coefficient,’’ as did Silva & Sites table. The total number of species found at
(1995), which is the same as Simpson’s the various sites ranges from 32–127.
equation (1), which can be used with or Table 2 shows the number of species
without expression as a percentage, but known for each site (on the diagonal) and
our incomplete samples did not lend the number of species shared in common
themselves to this. Consequently, we also by each pair of sites (to upper right of the
do not take the extra step taken by the diagonal). Kartabo, with 127 known spe-
other authors cited, to use UPGMA to cies, is the site where by far the most effort
produce a dendrogram of the sites and was expended, and yet the absence of
their similarities (e.g., Voss et al. certain species of small frogs that probably
2001:182). This would be an appropriate occur there indicates that even the Karta-
future endeavor when more complete bo inventory is incomplete. The second
biodiversity data are available. largest herpetofauna known is for Dubu-
We made faunal comparisons between lay Ranch, including the samples for both
lowland sites separately from those be- years we sampled there, which has mixed
Table 2.—Comparisons of number of species of habitat and the other in rainforest, and it is
amphibians and reptiles found at seven lowland well known that these different habitats
localities in Guyana. Numbers in diagonal row (in
bold italics) are numbers of species found at each
have significant differences in faunas (e.g.,
site. Numbers to upper right of diagonal are Hoogmoed 1979). Comparing any two
numbers of species found in common at sites where pairs of rainforest sites produced indices
rows and columns meet. Numbers to lower left of of 50% or more, except for one (Baramita
diagonal are faunal resemblance indices with vs. Konawaruk Camp, 39%), which is
correction for small samples (% of species in the
smallest sample found in common between the two understandable because we worked at
samples), or C/N1 3 100, comparing sites where rows Konawaruk Camp (1998) during a major
and columns meet. The sites are as follows: Ai, El Niño event and accompanying drought,
Aishalton; Ba, Baramita; Be, Berbice River Camp; which must have had a negative impact on
Du, Dubulay Ranch; Ka, Karanambo; Ko, our collecting. Comparison of the two
Konawaruk Camp; Kt, Kartabo. The habitats are
as follows: F, forest; M, patchwork of forest, savanna sites (Aishalton and Karanambo)
savanna, and gallery forest; S, savanna (but with produced an index of 59%. The site with
gallery forest). mixed habitats (Dubulay Ranch) produced
Ba F Kt F Ko F Du M Be F Ka S Ai S
an index of 68–72% compared with the
savanna sites and 47–70% compared with
Ba F 54 45 21 30 27 11 6 the rainforest sites. Indices for rainforest
Kt F 83 127 49 58 50 20 14 sites occurring to the east of the Essequibo
Ko F 39 83 59 28 31 8 5
Du M 56 70 47 83 32 27 23 River vs. those occurring to the west varied
Be F 50 77 53 49 65 12 7 from 50–77%, demonstrating that many
Ka S 29 50 21 68 32 40 19 species are shared by faunas on both sides
Ai S 19 44 16 72 22 59 32 of this major river.
For information on general biogeogra-
phy of the herpetofauna of the Guianan
habitats. The two least diverse lowland
Region and for faunal differences among
faunas are the two sites on the Rupununi
species occurring in rainforest vs. savanna
Savanna. What is interesting in this habitats, see Hoogmoed (1979). Three very
context, though, is that there are savanna important details pertinent to the biogeog-
species that do not occur in the rainforest, raphy of Guyana are: occurrence of upland
and vice-versa (see below). and isolated highland endemics in the
Table 2 also shows the faunal resem- unique habitats of the Pakaraima Moun-
blance indices (to the lower left of the tains in western Guyana (see below),
diagonal) showing the percentage of the occupants of lowland rainforests vs. savan-
smallest sample that is shared by both nas, and the Essequibo [Rupununi]-Rı́o
faunas where rows and columns meet Branco Depression as possibly both a
(equation with correction for small sam- historical barrier to terrestrial fauna (east/
ples). The highest index is 83%, for two west) and a corridor for aquatic fauna
comparisons: Baramita vs. Kartabo and (north/south). ‘‘The Essequibo-Rı́o Branco
Konawaruk Camp vs. Kartabo. This is Depression seems to have been a barrier to
interesting because these three sites are in the eastward distribution of a number of
rainforest habitat and occur on the same species, mainly Pantepui species’’ (Hoog-
side (west) of the Essequibo River, which moed 1979:257). ‘‘The Essequibo-Rı́o Bran-
may be a significant biogeographical item co Depression forms a barrier for the
(Hoogmoed 1979). The lowest index is distribution of a number of eastern forest
16% (Aishalton vs. Konawaruk Camp) species to the west and of western forest
and close second lowest is 19% (Aishalton species to the east’’ (Hoogmoed 1979:264).
vs. Baramita). This is interesting because ‘‘The Essequibo-Rı́o Branco Depression
in both comparisons, one site is in savanna also served as a route for lowland Amazo-
nian species invading the northern part of elevation, slope, soil depth, moisture,
Guiana. . .The occurrence in Guyana of the drainage, and exposure. Herpetofaunal
aquatic Melanosuchus niger (the black collections were not made uniformly in
caiman) and Chelus fimbriatus (the mata- all microhabitats; rather, most collections
mata [turtle]) apparently is the result of the were made at or near locations suitable for
rainy season connection between the Rı́o camping, and visits to the high peaks were
Branco and the Essequibo River via the of relatively short duration. This uneven
flooded Rupununi Savanna’’ (Hoogmoed sampling is a probable cause for some of
1979:257–258). In addition, the extensive the apparent differences in faunal compo-
tables provided by Señaris & MacCulloch sition among the three mountains.
(2005), on Guianan amphibians, and Avila- For purposes of comparison, we in-
Pires (2005), on Guianan reptiles, are clude only those species collected above
annotated with brief comments about 1400 m. In previous studies, highland
endemism and general distribution of most locations have been considered as those
of the species. above the arbitrarily chosen elevation of
1500 m (McDiarmid & Donnelly 2005,
MacCulloch et al. 2007). However, in
Isolated Highland Sites some cases major collecting sites of ours
(Above 1400 m Elevation) were located between 1400–1500 m, and
these collections are included here as
The isolated highlands are located in being from isolated highland sites. Species
west-central Guyana, and constitute the
collected on the three mountains are
highest parts of the Pakaraima Mountains,
shown in Table 3.
which stand on a base of uplands of
Composition and origin of tepui herpe-
intermediate elevations. Most highlands
tofaunas can be complex. Isolated tepuis
are sandstone mountains, often called
often support endemic species, some spe-
tepuis, typical of the Guiana Shield. The
highest peaks mark the borders with cies may occur on two or more tepuis, and
Venezuela and Brazil. Mount Roraima, some lowland taxa can ‘‘invade’’ higher-
the tallest peak in the Pakaraimas within elevation habitats. In addition, now isolat-
Guyana, at 2810 m elevation, is at the ed but adjacent peaks may once have been
point where Guyana, Brazil, and Venezue- part of one high plateau from which the
la meet. The highlands provide the head- intervening material has eroded away. The
waters of the Cuyuni, Ireng, Mazaruni, degree of species overlap among moun-
and Potaro rivers. Many of the higher tains can depend on factors such as
peaks support cloud forest habitat, which proximity, microhabitats, and local extir-
is not found elsewhere in Guyana. pations. Tepui herpetofaunal composition
Herpetological collecting has been car- has been examined, for examples, by
ried out primarily at three isolated high- Myers & Donnelly (2001, 2008), McDiar-
land sites (Fig. 4): Mounts Roraima, mid & Donnelly (2005), MacCulloch et al.
Ayanganna, and Wokomung. Mount (2007), and MacCulloch & Lathrop (2009).
Ayanganna is the highest peak that is Calculations of estimated total number
entirely within Guyana, at 2000 m. Mount of species (S*) were made for Mounts
Wokomung, 37 km south of Ayanganna, Ayanganna and Wokomung, using Chao’s
has its summit at 1700 m. Ayanganna and estimator as for the lowland sites. Because
Wokomung are some 80 km east of we have no data on the numbers of
Roraima. singletons and doubletons collected on
Habitats on the three tepuis are quite Roraima, we were unable to calculate S*
variable, depending on factors such as for this location.
Table 3.—Herpetofaunal comparisons of three Table 3.—Continued.

isolated highland sites in Guyana. X indicates
presence of the species, – indicates not documented. Taxon Roraima Ayanganna Wokomung
The sites are Mounts Roraima, Ayanganna, and

Dipsas catesbyi – X –
Wokomung. Only those specimens reported from
Dipsas pakaraima – X –
above 1400 m are included. Locations are a mix of
Liophis breviceps X – –
forest, low vegetation, and in some cases bare rock.
Liophis reginae – X –
Some data are from MacCulloch & Lathrop (2009),
Bothriopsis taeniata X – –
MacCulloch et al. (2007), and Means (2007).
Crotalus durissus X – –
Taxon Roraima Ayanganna Wokomung Total (n ¼ 49 species) 23 26 23
Anomaloglossus beebei – X –
Anomaloglossus – X – The Isolated Highland Faunal
megacephalus
Anomaloglossus praderioi X – –
Comparisons
Anomaloglossus cf. – – X
praderioi Similarities and differences among the
Anomaloglossus roraimae X – – faunas at the three locations are shown in
Anomaloglossus sp. – – X Tables 3 and 4. The three sites are
Oreophrynella – X X remarkably similar in numbers of species
dendronastes
Oreophrynella macconnelli X – – collected, despite the differences in collect-
Oreophrynella quelchii X – – ing effort. Although all three sites pro-
Rhaebo nasicus – – X duced 23–26 species, the highest number of
Vitreorana gorzulae – – X species shared among sites is only 14.
Pristimantis – – X
dendrobatoides Species overlap between sites is strongly
Pristimantis inguinalis – X – correlated with distance between the sites.
Pristimantis jester – X X Mounts Ayanganna and Wokomung,
Pristimantis marmoratus X X X which are 37 km apart, have the greatest
Pristimantis pulvinatus – X –
Pristimantis saltissimus – – X faunal resemblance (61%), while signifi-
Adelophryne patamona – – X cantly fewer species are shared between
Stefania ackawaio – X X these two sites and the more distant
Stefania ayangannae – X X Roraima (both less than 40%). The num-
Stefania coxi – X X
Stefania roraimae X X X ber of species found at only one of the
Stefania scalae X X – highland sites is especially striking, indi-
‘‘Hyla’’ warreni X X – cating a strong distinctiveness of each site.
Hypsiboas roraima X X X
Hypsiboas sibleszi X X X
Table 4.—Comparisons of number of species of
Myersiohyla kanaima X X X
amphibians and reptiles found at three isolated
Osteocephalus phasmatus – X X
highland localities in Guyana. Numbers in diagonal
Osteocephalus sp. – X X
row (in bold italics) are numbers of species found at
Adenomera lutzi – X X
each site. Numbers to upper right of diagonal are
Leptodactylus fuscus X – –
numbers of species found in common at sites where
Pseudopaludicola sp. X – –
rows and columns meet. Numbers to lower left of
Otophryne steyermarki X X X
diagonal are faunal resemblance indices with
Epicrionops niger – X –
correction for small samples (% of species in the
Arthrosaura guianensis – X –
smallest sample found in common between the two
Arthrosaura reticulata – – X
samples), or C/N1 3 100, comparing sites where rows
Neusticurus rudis X X –
and columns meet.
Riolama leucosticta X – –
Anolis planiceps X – – Roraima Ayanganna Wokomung
Mabuya nigropunctata X – –
Tropidurus hispidus X – – Roraima 23 9 6
Atractus steyermarki X – – Ayanganna 39 26 14
Chironius challenger – – X Wokomung 26 61 23
For Mount Roraima, 14 of 23 species were Camp, and Dubulay Ranch), Liophis
found only there (61%); for Mount Ayan- reginae (Ayanganna and Kartabo), and
ganna, 8 of 26 species (31%); and for Crotalus durissus (Roraima, Dubulay
Mount Wokomung, 9 of 23 species (39%). Ranch, and Karanambo). Many of these
Just as the highlands are important as species apparently are widely distributed
significant watersheds for Guyana, they in the Guianan Region and they appear
also support unique biotas. to have a wide range of tolerance for
living in different environments, especial-
ly with different temperature regimes.
Comparisons of Lowland vs. Isolated However, one must bear in mind the
Highland Sites state of our taxonomic knowledge and
realize that some of the apparently wide-
At this point in time, only inadequate ranging taxa, such as Leptodactylus
comparisons of the lowland and isolated fuscus, may actually represent two or
highland faunas can be made because the more cryptic species that have not yet
faunal lists are incomplete and more sites been recognized as such (e.g., Wynn &
need sampling in lowland, upland (inter- Heyer 2001).
mediate elevations), and isolated highland All 11 species shared between any
areas. Consequently, we provide only a lowland and isolated highland sites are
few general comments, hoping that future species that are wide-ranging at low
work will lead to rigorous comparisons elevations. It appears as if these lowland
and a better understanding of the biolog- species historically dispersed upward on-
ical phenomena involved. to the mountains from below. Note also
For the seven lowland sites compared, that of these 11 species only one,
a total of 188 species were listed (Table Neusticurus rudis, was found at more
1). For the three isolated highland sites, than one highland site. This suggests a
49 species were listed (Table 3). Of these, haphazard, perhaps opportunistic, nature
only 11 species out of the total of 224 of lowland species successfully colonizing
(5%) were shared between any lowland the highlands. None of the 11 shared
and highland sites, again showing how species appears to be a high elevation
distinctive the isolated highland sites are. form that has invaded the lowlands from
Those 11 species were Rhaebo nasicus above, although highland taxa did shift
(Wokomung and Kartabo), Leptodacty- ranges downward in elevation during
lus fuscus (Roraima, Kartabo, Dubulay glacial maxima, which affected distribu-
Ranch, Karanambo, and Aishalton), tion of biota even in areas that lacked
Arthrosaura reticulata (Wokomung and glaciers (e.g., Rull 2005). This also
Berbice River Camp), Neusticurus rudis demonstrates the importance of each
(Roraima, Ayanganna, Kartabo, and isolated highland as a unique area in
Konawaruk Camp), Anolis planiceps which evolution has produced endemic
(Roraima, Baramita, Kartabo, Konawa- novelties, although, paradoxically, the
ruk Camp, and Berbice River Camp), lineages of some highland endemics may
Mabuya nigropunctata (Roraima, Bara- be ancient, reflecting the deep history of
mita, Kartabo, Konawaruk Camp, Ber- the tepuis of the Guiana Shield, which
bice River Camp, and Aishalton), has been above sea level for longer than
Tropidurus hispidus (Roraima, Dubulay the entire evolutionary history of the
Ranch, Karanambo, and Aishalton), vertebrates. The Pakaraima Mountains
Dipsas catesbyi (Ayanganna, Kartabo, have high peaks of distinctive habitats,
and Berbice River Camp), Liophis brevi- including considerable bare rock, sur-
ceps (Roraima, Kartabo, Konawaruk rounded below by lowlands of tropical
Table 5.—Numbers of species shared in common better than reptiles at tolerating and
between seven lowland and three isolated highland adapting to relatively low temperature
sites in Guyana. A total of 224 different species
were found at these ten sites, showing that few
regimes within the tropics.
species are shared between the lowlands and
highlands. The sites are as follows: Ai, Aishalton;
Ba, Baramita; Be, Berbice River Camp; Du,
Dubulay Ranch; Ka, Karanambo; Ko,
Areas of Endemism and Other
Konawaruk Camp; Kt, Kartabo. Special Areas
Site Ba Kt Ko Du Be Ka Ai The currently documented herpeto-
Roraima 2 5 4 4 2 3 3
fauna of Guyana includes 324 species
Ayanganna 1 2 1 0 1 0 0 (148 amphibians, 176 reptiles, including
Wokomung 0 1 0 0 1 0 0 those in Appendix 1). Of these, 49
species (15%) are endemic or near-
endemic species, found nowhere else in
rainforest, like islands in the sea. As the world (or only marginally, in similar
habitat of immediately adjacent areas,
erosion of the highlands progresses, new
for example, the Pakaraima highlands of
species evolve on newly isolated peaks
Brazil and Venezuela). For amphibians,
while the surrounding lowland species
the number of endemic species (27%) is
may remain relatively stable. In addition,
much higher than for reptiles (5%). This
the peaks are very small geographic areas
is consistent with the observation that
and consequently, while their floras and
most of the endemic species are from
faunas are distinctive and unique, the list
isolated highlands, where amphibians
of their species is relatively short.
apparently are better than reptiles at
Table 5 shows the number of species adapting to low temperatures (see Bio-
shared in the isolated highland vs. lowland geography, above).
sites. For 13 of the 21 comparisons (62%), 0 Table 6 lists the endemic species, their
or only 1 species was shared, again showing type localities, and also the type localities
how distinctive the isolated highlands are. within Guyana for more wide-ranging
The highest degree of shared highland and species that were originally described
lowland species was between Roraima and primarily or only based on samples from
Kartabo, with 22% of Roraima’s fauna Guyana. Type localities are within Guya-
shared. The next highest were Roraima vs. na for 78 species of amphibians and
Konawaruk Camp (17% of Roraima’s) and reptiles, or 24% of the herpetofauna. This
Roraima vs. Dubulay Ranch (17% of includes 53 amphibians (36%) and 25
Roraima’s). These sites are all forested or reptiles (14%). The fact that one-third of
with mixed habitats. Few species were the amphibians have their type localities
shared between isolated highlands and the in Guyana and more than one-quarter of
more distant sites on the low-elevation the amphibians are endemic to Guyana
Rupununi Savanna. attests to how special this fauna is in
The list of species for the seven lowland global terms.
sites compared (Table 1) included 71 Two of the species in Table 6 have yet
amphibians and 117 reptiles; or amphibi- to be described and named. For the 78
ans comprised 38%. In strong contrast, the species with type localities in Guyana, 35
list of species for the three isolated (45%) are from 6 areas in the Pakaraima
highland sites (Table 3) included 34 uplands and isolated highlands, as fol-
amphibians and only 15 reptiles; or am- lows: Kaieteur National Park (9 species,
phibians comprised 69%. In general, this or 12%); Mt. Roraima (8 species, or
probably indicates that amphibians are 11%); Mt. Ayanganna (8 species, or
Table 6.—Endemic species (although these may occur in limited areas of adjacent countries in the
Pakaraima Mountains) and type localities of amphibians and reptiles within Guyana. Type localities are
abbreviated; see species accounts (in same sequence as given here) for details. þ ¼ yes; – ¼ no.
Species Endemic Type locality
Amphibians
Allophryne ruthveni – Tukeit Hill, below Kaieteur Falls
Allobates spumaponens þ Mabura Hill Forest Reserve
Anomaloglossus beebei þ Kaieteur National Park
Anomaloglossus kaiei þ Kaieteur National Park
Anomaloglossus megacephalus þ Mt. Ayanganna
Anomaloglossus roraima þ Mt. Roraima
Oreophrynella dendronastes þ Mt. Ayanganna
Oreophrynella macconnelli þ Mt. Roraima
Oreophrynella quelchii þ Mt. Roraima
Oreophrynella seegobini þ Mt. Maringma
Oreophrynella weiassipuensis þ Wei-Assipu-Tepui
Hyalinobatrachium taylori – New River area
Ceuthomantis smaragdinus þ Mt. Kopinang, Wokomung Massif
Pristimantis aureoventris þ Wei Assipu Tepui
Pristimantis dendrobatoides þ Wokomung Massif
Pristimantis inguinalis – New River area
Pristimantis jester þ Mt. Wokomung
Pristimantis saltissimus þ Wokomung Massif
Dendrobates nubeculosus þ Rockstone, Essequibo River
Adelophryne gutturosa – Mt. Roraima
Adelophryne patamona þ Mt. Wokomung
Stefania ackawaio þ Mt. Ayanganna
Stefania ayangannae þ Mt. Ayanganna
Stefania coxi þ Mt. Ayanganna
Stefania evansi þ Groete Creek, Essequibo
Stefania roraimae þ Mt. Roraima
Stefania woodleyi þ Mt. Kanaima, Pakaraima Mts.
Dendropsophus grandisonae þ Forest at Mazaruni
‘‘Hyla’’ helenae þ Dunoon, valley of Demerara River
‘‘Hyla’’ warreni þ Mt. Roraima
Hypsiboas calcaratus – ‘‘British Guiana’’
Hypsiboas liliae þ Kaieteur National Park
Hypsiboas ornatissimus – Meamu, Mazaruni River
Hypsiboas roraima þ Mt. Roraima
Lysapsus laevis – Rupununi Savanna
Myersiohyla kanaima þ Mt. Kanaima, Pakaraima Mts.
Osteocephalus exophthalmus þ Pakaraima uplands
Osteocephalus phasmatus þ Mt. Ayanganna
Scinax sp. ‘‘þ’’ Undescribed, Berbice River Camp
Tepuihyla talbergae þ Kaieteur National Park
Trachycephalus hadroceps – Acarai Mts., W of New River
Adenomera lutzi þ Pakaraima uplands
Leptodactylus guianensis – Iwokrama Forest Reserve
Leptodactylus rugosus – Kaieteur National Park
Chiasmocleis shudikarensis – Shudikar-wau, upper Essequibo River
Synapturanus mirandaribeiroi – Kanashen, upper Essequibo River
Caecilia pressula þ Marudi Mts.
Oscaecilia zweifeli – Tributary of Mazaruni River
Epicrionops niger – Arundabara uplands
Rhinatrema shiv þ Kaieteur National Park
Caecilita iwokramae þ Iwokrama uplands
Microcaecilia iyob þ Oko River at Cuyuni River
Microcaecilia cf. rabei ‘‘þ’’ Undescribed
Species Endemic Type locality
Reptiles
Amphisbaena stejnegeri þ Vreeden Rust, Demerara River
Amphisbaena vanzolinii – Marudi Mts.
Hemidactylus palaichthus – Kurupukari
Arthrosaura guianensis þ Mt. Ayanganna
Arthrosaura hoogmoedi þ Mt. Maringma
Echinosaura sulcarostrum þ Baramita
Kaieteurosaurus hindsi þ Kaieteur National Park
Leposoma guianense – Dunoon, Demerara River
Neusticurus rudis – Foot of Mt. Roraima
Pantepuisaurus rodriguesi þ Mt. Maringma
Ptychoglossus brevifrontalis – New River area
Riolama leucosticta þ Mt. Roraima
Tretioscincus agilis – Near Dunoon
Gonatodes alexandermendesi – Konawaruk Camp
Gonatodes annularis – Maccasseema, Pomeroon River
Gonatodes timidus þ Iwokrama Forest Reserve
Pseudogonatodes guianensis – Upper Cuyuni River
Atractus tamessari þ Kaieteur National Park
Chironius challenger – Maringma Tepui
Dipsas pakaraima þ Mt. Ayanganna
Ninia hudsoni – New River area
Phimophis guianensis – Savanna, vicinity of Pirara
Micrurus averyi – Head, Itabu Creek, New River area
Phrynops tuberosus – Cotinga River, Mt. Roraima
Podocnemis unifilis – Rupununi and Takutu rivers
Totals 49 78
11%); the Wokomung Massif (5 species, River and Konawaruk Camps; Groete
or 7%); Mt. Maringma (4 species, or Creek and Rockstone, Essequibo River;
5%); and Wei Assipu Tepui (1 species, or Kanashen and Shudikar-wau, upper Es-
1%). It is noteworthy that the site sequibo River; the Rupununi Savanna;
representing the highest number of spe- Marudi Mountains; and the New River
cies in this context, Kaieteur National area.
Park, is both a site of intermediate
elevations and the one where the most
extensive studies have been made (e.g., Endangered Species
Kok & Kalamandeen 2008).
The numbers cited above show clearly The Red List of the International Union
that the Pakaraima uplands and isolated for the Conservation of Nature (IUCN)
highlands are special for the biodiversity includes a comprehensive assessment of
of Guyana. Nevertheless, the type local- the conservation status of the known
ities (Table 6) also show that areas of species of amphibians of the world. The
special significance to biodiversity are first assessment of all amphibians was
scattered throughout the country, even completed as the Global Amphibian As-
in the lowlands, including the following sessment in 2004 (Chanson et al. 2008a),
sites: Baramita; Dunoon and Vreeden and the process is ongoing with periodic
Rust, Demerara River; Iwokrama and updates that incorporate new information,
Mabura Hill forest reserves; Berbice including newly described species. The
current Red List (IUCN 2010) uses the Table 7.—Amphibians listed by the IUCN Red
following categories of conservation sta- List (IUCN 2010) as species of concern for
conservation, with the exception of species listed as
tus, listed from the least to greatest Least Concern. DD ¼ Data Deficient; VU ¼
concern: Least Concern; Near Threatened; Vulnerable. Population Trend is indicated by
Vulnerable; Endangered; Critically Endan- Decreasing, Stable, or Unknown (Ukn).
gered; Extinct in the Wild; and Extinct.
Species of concern Status Population Trend
Another category, Data Deficient, is for
species for which there is not yet sufficient Anomaloglossus beebei VU Stable
information to make a judgment. Threat- Anomaloglossus praderioi DD Ukn
Anomaloglossus roraima DD Ukn
ened species are included in the categories Atelopus spumarius VU Decreasing
of Vulnerable, Endangered, and Critically Oreophrynella dendronastes DD Ukn
Endangered. Oreophrynella macconnelli VU Stable
The Neotropical Realm, which stretches Oreophrynella quelchii VU Stable
from central Mexico southward through- Oreophrynella weiassipuensis DD Ukn
Vitreorana gorzulae DD Ukn
out South America and includes all of the Vitreorana helenae DD Ukn
Caribbean Islands, has by far the greatest Hyalinobatrachium iaspidiense DD Ukn
species richness of amphibians. This in- Dendrobates nubeculosus DD Ukn
cludes nearly half of the amphibians of the Adelophryne patamona DD Stable
world, 96% of which are endemic to the Stefania roraimae DD Ukn
Dendropsophus grandisonae DD Ukn
Realm (Chanson et al. 2008b). This Realm ‘‘Hyla’’ helenae DD Ukn
also has the greatest proportion of threat- ‘‘Hyla’’ warreni DD Ukn
ened and extinct species of amphibians in Hypsiboas roraima DD Ukn
the world, and it is the region that has been Osteocephalus exophthalmus DD Ukn
Osteocephalus phasmatus DD Ukn
impacted hardest by the lethal amphibian
Tepuihyla talbergae DD Ukn
fungal disease, chytridiomycosis. Whereas Adenomera lutzi DD Ukn
the overall threat level is high in Neo- Chiasmocleis jimi DD Ukn
tropica (60% of all globally threatened Caecilia pressula DD Ukn
amphibians occur here) compared to the Oscaecilia zweifeli DD Ukn
Microcaecilia rabei DD Ukn
rest of the world, threat levels are com-
paratively low in the Guiana Shield
(Bolaños et al. 2008), and in general,
populations of amphibians in Guyana lus, Oreophrynella seegobini, Ceuthomantis
appear to be in good condition. smaragdinus, Scinax sp., Caecilita iwokra-
None of the modern species of amphib- mae, Microcaecilia iyob, Microcaecilia cf.
ians that occur in Guyana is listed as rabei, and Rhinatrema shiv.
Extinct or Extinct in the Wild. Similarly, The IUCN has yet to complete a
none is listed as Critically Endangered, modern assessment of the conservation
Endangered, or Near Threatened. Consid- status of the reptiles of the world, compa-
ering the species that occur in Guyana rable to that of the amphibians. The
(and surrounding areas), 81% are listed as reptiles are the last group to be completed
Least Concern, 16% are listed as Data for the terrestrial vertebrates, and it has
Deficient, and only 3% are listed in the been initiated through a series of regional
lowest threatened category of Vulnerable projects (IUCN 2010). Nevertheless, an
(Table 7, in which we did not include the earlier assessment had listed only the five
species listed as of Least Concern). The sea turtles of Guyana (100% of them) as
following eight species that were described being of significant conservation concern
recently or have yet to be described were among all the reptiles (excluding birds),
not included in the latest assessment with 40% of the sea turtles Critically
(IUCN 2010): Anomaloglossus megacepha- Endangered, 40% Endangered, and 20%
Vulnerable–Decreasing. The status of We present dichotomous keys for iden-

these species is also under review. tifying representatives of all the species
known in the herpetofauna of Guyana,
and we present brief annotated species
accounts. However, some species are listed
Summary and Conclusions and discussed in Appendix 1, as they
became known to us after we completed
We confirm that at least 324 species of
nearly all of our text in 2011. For the
amphibians and reptiles occur in Guya-
species accounts, we have examined and
na. This includes 148 amphibians (137
verified all the original literature references
frogs and toads, 11 caecilians) and 176
in which the species were named, but we
non-avian reptiles (4 crocodylians, 4
have not tried to survey all scientific
amphisbaenians, 56 lizards, 97 snakes,
collections in the world to confirm every
and 15 turtles). These are listed in
single specimen collected in Guyana in the
Appendix 2.
past.
Documentation of these species was Aspects of biogeography are discussed
accomplished through field work and based on collections made at seven low-
identification of specimens that we and land sites (4 in rainforest, 2 in savanna, 1
field associates including UG students in mixed habitats; all below 500 m
collected; reidentifying old museum speci- elevation) and three sites in isolated
mens collected by others in the past (e.g., highlands (in montane forest and ever-
William Beebe); and reviewing document- green high-tepui forest; all above 1400 m
ed publications and collection records of elevation). This discussion presents pre-
colleagues (e.g., Pritchard 1969, Kok & liminary comparisons because the sam-
Kalamandeen 2008; collection of Peter C. pling of each local fauna is incomplete;
H. Pritchard’s Chelonian Research Insti- more field effort is necessary.
tute). Voucher specimens are cited in all Although much remains to be learned, it
our species accounts and references in is clear that an area of about 2.5 km2 of
Appendix 1. To date, no other large group lowland rainforest in Guyana probably
of animals of Guyana has received such supports more than 130 species of am-
detailed treatment. phibians and reptiles, whereas many fewer
Undoubtedly, Guyana has additional species (fewer than 30 documented) live in
species that remain to be discovered, a comparable area of the isolated high-
named, and described, and perhaps 20 lands. Comparisons of the faunal lists of
additional species known from adjacent these 10 sites show that very few species
countries will ultimately be found to occur occur in both the lowlands and isolated
in Guyana. Furthermore, future research, highlands (all are widespread lowland
especially involving molecular genetics, forms); that many endemic species occur
undoubtedly will reveal that some popula- in the isolated highlands (mostly amphib-
tions interpreted today as representing ians); and that each of the isolated
wide-ranging species are actually com- highlands, lowland savannas, and lowland
prised of multiple cryptic species yet to rainforests at 10 sites have distinctive
be recognized. faunal elements. No two sites were identi-
Consequently, we are confident that cal in species composition. Much more
within a few decades the herpetofauna of work is needed to compare a variety of
Guyana will be confirmed at more than sites, and especially to incorporate upland
350 species and this report will need sites of intermediate elevations in future
updating. comparisons.
Endemic species, which occur nowhere Acknowledgments

else in the world, comprise 15% of the
herpetofauna of Guyana. Although some While visiting Guyana, each author
of the endemic species are lowland rain- received considerable advice and other
forest species, most are amphibians of the assistance from Mike Tamessar at the
isolated highlands and uplands of the UG. In addition, our efforts through the
Pakaraima Mountains, where they live in BDG were greatly assisted by Malcolm
headwaters of major watersheds. In fact, Rodrigues and Indarjit (Charles) Ramdass
about 27% of the amphibian fauna is of UG, as well as, Vicki Funk and Carol
endemic to Guyana, or nearly so. Kelloff of the NMNH, without whose help
Another gauge of the distinctiveness of a this project would not have been complet-
fauna is to review the type localities in an ed. Critically important logistical support
area. These are the localities from which for our field work in Guyana was provided
the population samples were collected that by the following outstanding individuals:
included the type specimens, which are the Deokie (Jackie) Arjoon, Malcolm and
permanent, formal name-bearers for the Margaret Chan-A-Sue, Philip DaSilva,
species. For 24% of the herpetofauna (36% Bunny Fernandes, Karen Pilgrim, and
of amphibians, 13% of reptiles), the type the late Diante Narine.
locality of the species is within Guyana, CJC and CRT conducted all their field
and we discuss several specimens of trips (1992–1998) with participation in the
amphibians in the species accounts that BDG, a collaboration between the UG
probably represent new species, as yet to and the NMNH, which included funding
be described and named. our expenses while in Guyana. Our field
Five species of sea turtles utilize the efforts were significantly assisted by the
limited areas of Atlantic coastal beaches in following, in addition to all the others
Guyana, primarily to the northwest of named above: Harold Ameer, Gerald and
Georgetown. All of these, (100%) are listed Wesley King, Walter Lachman, Clifton
by the International Union for the Con- and Maureen Laurindo, Compton
servation of Nature as being of global McKenzie, Diane McTurk, Alfred Mekde-
concern for long-term survival, mostly ci, Alexander Mendes and Adriana Man-
owing to predation by humans (IUCN dal, Ronnie and Petronella Michaels, and
2010): 2 species (40%) are listed as being many additional citizens of Aishalton and
Critically Endangered (Eretmochelys im- Yupukari. CJC and CRT also received
bricata and Dermochelys coriacea); 2 (40%) funding from the American Museum of
are listed as Endangered (Caretta caretta Natural History (AMNH) when necessary,
and Chelonia mydas); and 1 (20%) is listed with special thanks to Michael J. Novacek
as Vulnerable–Decreasing (Lepidochelys and Darrel Frost.
olivacea). Clearly, it is important to protect In addition to the individuals cited
the nesting beaches of the sea turtles. above, RPR extends special thanks to
It appears to us that Guyana is fortu- Harold Ameer, Deokie (Jackie) Arjoon,
nate in having its native amphibian and Oliver Flint, Lynne Gillespie, Bill Hahn,
reptilian faunas in very good shape in Traci Hartsell, Wayne Mathis, Suzanne
comparison to many other countries, Peurach, Eustace Smith, Ruby Smith, Paul
particularly those elsewhere in which Spangler, Barrington Williams, Don Wil-
certain amphibians have become extinct son, and the late Harry Persaud for greatly
and others are suffering dramatic losses. assisting his field work in Guyana from
We hope that preservation of the original 1989 to 1999. RPR especially appreciates
fauna and flora will continue to be an the funding provided through Vicki Funk
important goal within Guyana. and Carol Kelloff of the BDG, and
through W. Ronald Heyer of the Smithso- We are very grateful to several colleagues
nian Neotropical Lowlands Program, and great friends who allowed us to publish
without which this work could not have some of their color photographs on the
been completed. color plates herein. The legends provide the
In addition to individuals cited above, initials of the photographer for each image,
RDM and AL received extraordinary except where no initials are given, those
assistance from Calvin Bernard, Godfrey were taken by CJC and CRT (AMNH).
Bourne, Michelle Kalamandeen, and Fr. For all the other images, we thank DDF,
C. L. Roland, and field assistance from Duane De Freitas, Dadanawa Ranch,
Carter Cox, Samir Khan, Brad Hubley, Guyana; SCF, Santiago Castroviejo-Fisher,
and guides and porters from Chinowieng, AMNH; PJRK, Philippe J. R. Kok,
Kaibarapai, and Paramakatoi. Their field IRSNB, Belgium; AL, Amy Lathrop,
work received enthusiastic travel and ROM, Toronto; JM, John Mitchell, Earth
logistical support in 1990 and 1992 from Images Foundation (photo donated to
Youth Challenge International, Toronto, ROM); and Peter C. H. Pritchard, Chelo-
and was funded by the Royal Ontario nian Research Institute, Oviedo, Florida. In
Museum Governors, the ROM Depart- addition, we are grateful for the use of three
ment of Natural History, and a BDG on-line photographs of animals for AL to
fellowship. adapt to line-drawings of key characters, as
For significant assistance in examining follows: Fig. 6E upper, from http://www.
biotransitando.blogspot.com/2009/01/
specimens or obtaining information or
biologia-fascinante-mesmo.html; Fig. 6E
digital photographs from other museum
lower, from http://www.biodiversitygroup.
collections, we thank Jonathan Campbell
org/topics/amphibians.html (Paul S. Ham-
and Carl Franklin (UTA); Darrel Frost,
ilton, The Biodiversity Group); and Fig.
Iris Calderon, David Dickey, Bartek Ja-
6F, from http://calphotos.berkeley.edu/cgi/
blonski, David Kizirian, Robert Pascocel-
img_query?enlarge¼0000þ0000þ0410þ1169
lo, Christopher Raxworthy, and Thomas (Andrés Acosta).
Trombone (AMNH); Steve Gotte, Ronald The basic maps in Figs. 2 and 3 were
Heyer, Roy McDiarmid, James Poin- provided by Ray Sterner, The Johns
dexter, and George Zug (NMNH); Phil- Hopkins University Applied Physics Lab-
ippe Kok (IRSNB); Kenneth Krysko oratory, Laurel, Maryland, who worked
(UF); Arnold Kluge, Ronald Nussbaum, with satellite images upon which AL
and Gregory Schneider (UMMZ); Celsa applied final details. We are most grateful
Señaris (MHNLS); Jonathan Losos and for all this effort also. Line drawings
José Rosado (MCZ); Colin McCarthy illustrating the key characters for identify-
(BMNH); Robert Murphy (ROM); and ing specimens (Figs. 5–9) and the outline
Annemarie Ohler (MNHN). In addition, map (Fig. 2) were executed by AL. Figure
Godfrey Bourne provided some specimens, 7G was adapted from an internet photo by
mostly from CEIBA, for this work. We are Philippe Macquet (http://www.flickr.com/
also extremely indebted to Thomas photos/26642064@N06/3989708345/) and
Baione, Director of the AMNH Library, Fig. 8E from internet photos of W. Wüster
and the many AMNH Reference Librari- (http://calphotos.berkeley.edu/).
ans who provided extensive help during This work was conducted in accordance
the 10 yr of preparation of this manuscript. with the national and international regu-
In addition, we thank Julian Faivovich, lations that apply to travel in Guyana,
Ronald Heyer, Marinus Hoogmoed, and wildlife protection, and humane euthana-
Robert Voss for comments on an earlier sia of samples with approved protocols of
draft of the manuscript. the various institutions involved. We also
are grateful for assistance in these matters Shield. Bulletin of the Biological Society of
from the following in Guyana: Environ- Washington 13:1–98 pp.
Avila-Pires, T. C. S., M. S. Hoogmoed, & W. Alves
mental Protection Agency, Department of da Rocha. 2010. Notes on the vertebrates of
Agriculture (Wildlife division), and the northern Pará, Brazil: a forgotten part of the
University of Guyana. Guianan region, 1. Herpetofauna. Boletim do
This report is number 179 in the Museu Paraense Emı́lio Goeldi Ciências
Naturais, Belém 5(1):13–112.
Smithsonian’s BDG publication series Ayarzagüena, J. 1992. Los Centrolénidos de la
and contribution number 346 of the Guayana venezolana. Publicaciones de la
ROM Centre for Biodiversity and Conser- Asociación de Amigos de Doñana 1:1–48.
vation Biology. We are especially grateful Bailey, J. R., & R. A. Thomas. 2007 [2006]. A
revision of the South American snake genus
to the Biological Society of Washington, Thamnodynastes Wagler, 1830 (Serpentes:
the Biodiversity of the Guiana Shield Colubridae, Tachymenini). II. Three new
Program, and the U.S. Geological Survey species from northern South America, with
Patuxent Wildlife Research Center for further descriptions of Thamnodynastes gam-
botensis Pérez-Santos and Moreno and Tham-
important financial contributions toward
nodynastes ramonriveroi Manzanilla and
producing this volume. Any use of trade, Sánchez. Memoria de la Fundación La Salle
product, or firm names is for descriptive de Ciencias Naturales 166:7–27.
purposes only and does not imply endorse- Bailey, J. R., R. A. Thomas, & N. J. Silva, Jr. 2005.
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Thamnodynastes Wagler, 1830 (Serpentes,
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Appendix 1 extent Alopoglossus angulatus, but there are consid-

erable differences in scalation.
Species Added to the Guyana Herpetofauna after
Completion of Our Text Reptilia, Lizards, Polychrotidae
Amphibia, Anura, Craugastoridae Anolis chrysolepis.—See comments in the species
account for Anolis planiceps. In the past, these two forms
Pristimantis aureoventris.—This recently de-
were treated as subspecies of one wide-ranging species,
scribed species is known from only two isolated
but recent molecular analyses indicated that the
highland localities, from 2210–2305 m elevation. The
subspecies should be elevated to the species level,
type locality is on Wei Assipu Tepui on the border of
although some forms are not well-differentiated mor-
Guyana and Brazil, and the other locality is on the
phologically (D’Angiolella et al. 2011). These authors
Guyana portion of Mount Roraima. The description
cited two specimens of A. chrysolepis from southern
is very thorough, lavishly illustrated, and compares
Guyana, and one was recently received at the ROM.
the new species with congeners from relevant
Anolis (sensu lato).—Townsend et al. (2011)
localities (Kok et al. 2011).
performed extensive phylogenetic analyses of DNA
Amphibia, Anura, Dendrobatidae sequence data and concluded that Anolis and Polychrus
are not supported as each others’ closest relative. As
Ranitomeya amazonica.—Species of Ranitomeya representatives of these genera clustered in different
are colorful, diminutive frogs that were previously monophyletic groups, these authors proposed a new
included in the genus Dendrobates. In particular, family, Dactyloidae, to accommodate the genus Anolis.
Ranitomeya is a taxonomically difficult genus that
Reptilia, Lizards, Sphaerodactylidae
has representatives basically throughout Amazonia,
with many cryptic species. The most recent generic Gonatodes timidus.—This recently described spe-
revision (Brown et al. 2011) shows that specimens are cies is known only from the base of the Iwokrama
identified to species with the greatest confidence by Mountains in the Iwokrama Forest Reserve, central
using molecular data, but in many cases one can Guyana (4819 0 52 00 N, 588470 58 00 W; 209 m elev.). The
estimate an identification based only on old museum description is thorough, with extensive illustrations
specimens or photographs. Brown et al. (2011:85) and detailed comparisons with congeners (Kok 2011).
stated that ‘‘based on morphology’’ of a museum
specimen, this species occurs at the following Reptilia, Snakes, Colubridae
locality: ‘‘Guyana (Region: Upper Takutu-Upper
Essequibo)’’ (Brown et al. 2011:86). On pages 117– Mastigodryas moratoi.—Montingelli & Zaher
120 they listed the museum specimens examined but (2011) described and named this new species, in part
did not give the locality data for them. based on AMNH R-141798 from 3.2 km by road
Ranitomeya uakarii.—Brown et al. (2011) on their WNW Dubulay Ranch house (5837 0 N, 57853 0 W),
page 51 showed (Fig. 17H) a frog photographed ‘‘in Guyana. All known specimens are from Brazil and
life, Iwokrama, Guyana (unknown photographer).’’ Guyana. On their page 119 and on map (p. 112) the
Apparently there is no voucher specimen. In their authors included Kuyuwini Landing, Guyana in the
Fig. 22 (p. 56), this was mapped as R. uakarii ‘‘based range, but they did not cite any specimens from this
on morphology’’ and on page 57 they stated that ‘‘we locality. AMNH has specimens of Mastigodryas from
provisionally consider the Guyana population as R. Kuyuwini Landing, including R-60782 and R-60810,
uakarii.’’ This needs to be confirmed, especially with both or either of which might be the basis of the
molecular data, as all other specimens known are comment. CJC examined both of these specimens and
from southern Colombia, eastern Peru, or extreme confirmed that AMNH R-60810 is M. moratoi;
western Brazil. This species is very similar to AMNH R-60782 was not identified, as neither the
Ranitomeya ventrimaculata and names previously light stripes nor hemipenes were visible (see below).
applied to it include Dendrobates quinquevittatus and Another AMNH specimen from Kuyuwini Landing,
D. ventrimaculatus. R-60852, is M. boddaerti, so the two species are
sympatric at this locality. Mastigodryas moratoi is a
Reptilia, Lizards, Gymnophthalmidae member of the pleei species group, which ‘‘is
distinguished from both [the] melanolomus and
Ptychoglossus brevifrontalis.—Peloso & Avila- boddaerti groups by the length and position of the
Pires (2010:372) reported that BMNH 139.1.1.75 is upper light lateral stripes. . .on the dorsum. In the
a representative of this previously described species former group, the larger upper light lateral stripes are
from ‘‘Suriname/Guyana: New River, 750 feet.’’ formed by three dorsal scale rows that include the
Their determination was made from photographs. upper portion of row 3, [all of] row 4, and the inner [¼
Very superficially (color, pattern, size), this species lower] portion of row 5, whereas in the latter two
resembles Arthrosaura reticulata and to a lesser groups the upper light lateral stripes are only two
dorsal scale rows wide, being formed by rows number Rhinella martyi (formerly margaritifera, ty-
4 and 5. . .’’ (Montingelli & Zaher 2011:111). The phonius)
adult from near Dubulay Ranch house (AMNH R- Rhinella merianae
141798) was described in life by CJC (field notes, Centrolenidae (genera formerly Centrolene, Co-
1995) as follows. Anteriorly, dorsum light tan with chranella):
brown and black contrasting stripes (the black only Hyalinobatrachium cappellei (sr. syn. crurifas-
anteriorly on the dorsolateral stripe). Posteriorly, the ciatum in Guyana)
pattern soon gradually becomes a broad brown stripe Hyalinobatrachium iaspidiense
on a tan ground color. Further posteriorly, the Hyalinobatrachium mondolfii
dorsum becomes uniform dark tan, continuing Hyalinobatrachium taylori
through the tail. Top of head brown; lips white, with Vitreorana gorzulae (sr. syn. papillahallicum)
dark brown to black line from snout through eye, Vitreorana helenae (sr. syn. oyampiensis in
along top of labials; underside of head dark gray with Guyana)
white spots; throat and anterior ventral surfaces pale Ceratophryidae (formerly in Leptodactylidae):
tan with extensive gray markings; rest of ventral Ceratophrys cornuta
surfaces orangish tan with gray markings becoming Ceuthomantidae:
fewer posteriorly while orange becomes bolder, Ceuthomantis smaragdinus
especially beneath base of tail, but turning yellow Craugastoridae: (formerly in Leptodactylidae;
further posteriorly beneath tail. genus Eleutherodactylus):
Pristimantis aureoventris
Pristimantis chiastonotus
Pristimantis dendrobatoides
Pristimantis inguinalis
Appendix 2 Pristimantis jester
Pristimantis marmoratus
List of Currently Known Herpetofauna of Guyana Pristimantis pulvinatus
(324 Species) Pristimantis saltissimus
Pristimantis zeuctotylus
The following list includes all the species in our
Dendrobatidae:
keys and species accounts, as well as those most Ameerega hahneli (formerly Dendrobates,
recently discovered after our basic text was complet- Epipedobates pictus in Guyana)
ed (Appendix 1). Ameerega trivittata
Dendrobates leucomelas
Amphibia: 148 species:
Dendrobates nubeculosus
Anura (137 species of frogs and toads)
Dendrobates tinctorius
Allophrynidae:
Ranitomeya amazonica
Allophryne ruthveni
Ranitomeya uakarii
Aromobatidae (formerly in Dendrobatidae;
Eleutherodactylidae (formerly in Leptodactyli-
genera formerly Colostethus, Dendrobates,
dae):
Epipedobates):
Adelophryne gutturosa
Allobates femoralis Adelophryne patamona
Allobates spumaponens (was C. brunneus in Eleutherodactylus johnstonei
Guyana) Hemiphractidae (formerly in Hylidae):
Anomaloglossus beebei Stefania ackawaio
Anomaloglossus kaiei Stefania ayangannae
Anomaloglossus megacephalus Stefania coxi
Anomaloglossus praderioi Stefania evansi
Anomaloglossus roraima Stefania roraimae
Bufonidae: Stefania scalae
Atelopus spumarius Stefania woodleyi
Dendrophryniscus minutus Hylidae:
Oreophrynella dendronastes Dendropsophus brevifrons (formerly Hyla)
Oreophrynella macconnelli Dendropsophus grandisonae
Oreophrynella quelchii Dendropsophus leucophyllatus
Oreophrynella seegobini Dendropsophus marmoratus
Oreophrynella weiassipuensis Dendropsophus minusculus
Rhaebo guttatus (formerly Bufo) Dendropsophus minutus
Rhaebo nasicus ‘‘Hyla’’ helenae
Rhinella marina (formerly Bufo) ‘‘Hyla’’ warreni
Hypsiboas boans (formerly Hyla) Leptodactylus rugosus

Hypsiboas calcaratus Leptodactylus validus (formerly pallidirostris)
Hypsiboas cinerascens (formerly granosa in Lithodytes lineatus
Guyana) Physalaemus cuvieri
Hypsiboas crepitans Pleurodema brachyops
Hypsiboas fasciatus Pseudopaludicola boliviana
Hypsiboas geographicus Microhylidae:
Hypsiboas lemai Chiasmocleis hudsoni
Hypsiboas liliae Chiasmocleis jimi
Hypsiboas multifasciatus Chiasmocleis shudikarensis
Hypsiboas ornatissimus Ctenophryne geayi
Hypsiboas punctatus Elachistocleis surinamensis (formerly ovalis)
Hypsiboas roraima Hamptophryne boliviana
Hypsiboas sibleszi Otophryne pyburni
Lysapsus laevis (formerly limellus) Otophryne robusta
Myersiohyla kanaima (formerly Hyla) Otophryne steyermarki
Osteocephalus buckleyi Synapturanus mirandaribeiroi
Osteocephalus cabrerai Synapturanus salseri
Osteocephalus exophthalmus Pipidae:
Osteocephalus leprieurii Pipa arrabali
Osteocephalus oophagus Pipa pipa
Osteocephalus phasmatus Ranidae:
Osteocephalus taurinus Lithobates palmipes (formerly Rana)
Phyllomedusa bicolor Gymnophiona (11 species of caecilians):
Phyllomedusa hypochondrialis Caeciliidae:
Phyllomedusa tarsius Caecilia gracilis
Phyllomedusa vaillantii Caecilia pressula
Pseudis paradoxa Caecilia tentaculata
Scinax boesemani (formerly Hyla, Ololygon) Oscaecilia zweifeli
Scinax cruentommus Rhinatrematidae:
Scinax nebulosus Epicrionops niger
Scinax proboscideus Rhinatrema shiv
Scinax ruber Siphonopidae:
Scinax trilineatus Caecilita iwokramae
Scinax cf. x-signatus Microcaecilia iyob
Scinax sp. Microcaecilia rabei
Sphaenorhynchus lacteus Microcaecilia cf. rabei
Tepuihyla talbergae Typhlonectidae:
Trachycephalus coriaceus (formerly Phryno- Typhlonectes compressicauda
hyas)
Trachycephalus hadroceps Reptilia: 176 species:
Trachycephalus resinifictrix Crocodylia (4 species):
Trachycephalus typhonius (formerly venulosus) Alligatoridae:
Leptodactylidae: Caiman crocodilus
Adenomera andreae Melanosuchus niger
Adenomera hylaedactyla Paleosuchus palpebrosus
Adenomera lutzi Paleosuchus trigonatus
Engystomops cf. pustulosus Squamata: Amphisbaenians or worm lizards (4
Leptodactylus fuscus species)
Leptodactylus guianensis (formerly bolivianus) Amphisbaenidae:
Leptodactylus knudseni Amphisbaena alba
Leptodactylus leptodactyloides Amphisbaena fuliginosa
Leptodactylus longirostris Amphisbaena stejnegeri
Leptodactylus macrosternum (formerly ocella- Amphisbaena vanzolinii
tus) Squamata: lizards (56 species)
Leptodactylus mystaceus Corytophanidae:
Leptodactylus pentadactylus Basiliscus basiliscus
Leptodactylus petersii (formerly podicipinus) Gekkonidae:
Leptodactylus rhodomystax Hemidactylus mabouia
Hemidactylus palaichthus Tropidurus hispidus

Thecadactylus rapicauda Uracentron azureum
Gymnophthalmidae (formerly within Teiidae): Uranoscodon superciliosus
Alopoglossus angulatus Squamata: snakes (97 species):
Arthrosaura guianensis Aniliidae:
Arthrosaura hoogmoedi Anilius scytale
Arthrosaura reticulata Anomalepidae:
Bachia flavescens (formerly monodactylus) Typhlophis squamosus
Cercosaura argulus (formerly Prionodactylus) Boidae:
Cercosaura ocellata Boa constrictor
Echinosaura sulcarostrum Corallus caninus
Gymnophthalmus leucomystax Corallus hortulanus
Gymnophthalmus cf. speciosus Epicrates cenchria
Gymnophthalmus underwoodi Eunectes murinus
Gymnophthalmus vanzoi Colubridae sensu lato:
Iphisa elegans Apostolepis nigrolineata
Kaieteurosaurus hindsi Atractus favae
Leposoma guianense Atractus schach
Leposoma percarinatum Atractus steyermarki
Neusticurus bicarinatus Atractus tamessari
Neusticurus rudis Atractus torquatus
Pantepuisaurus rodriguesi Atractus trilineatus
Ptychoglossus brevifrontalis Chironius carinatus
Riolama leucosticta Chironius challenger
Tretioscincus agilis Chironius exoletus
Iguanidae: Chironius fuscus
Iguana iguana Chironius multiventris
Polychrotidae: Chironius scurrulus
Anolis aeneus Clelia clelia
Anolis auratus Dendrophidion dendrophis
Anolis chrysolepis Dipsas catesbyi
Anolis fuscoauratus Dipsas copei
Anolis ortonii Dipsas indica
Anolis planiceps (formerly nitens, chrysolepis) Dipsas pakaraima
Anolis punctatus Dipsas pavonina
Polychrus marmoratus Dipsas variegata
Scincidae: Drepanoides anomalus
Mabuya nigropunctata (formerly mabouia) Drymarchon corais
Sphaerodactylidae (formerly in Gekkonidae): Drymobius rhombifer
Chatogekko amazonicus (formerly Coleodac- Drymoluber dichrous
tylus) Elapomorphus quinquelineatus
Coleodactylus septentrionalis Erythrolamprus aesculapii
Gonatodes alexandermendesi Helicops angulatus
Gonatodes annularis Hydrodynastes bicinctus
Gonatodes humeralis Hydrops triangularis
Gonatodes timidus Imantodes cenchoa
Gonatodes vittatus Imantodes lentiferus
Pseudogonatodes guianensis Leptodeira annulata
Sphaerodactylus molei Leptophis ahaetulla
Teiidae: Liophis breviceps
Ameiva ameiva Liophis cobella
Cnemidophorus lemniscatus Liophis lineatus
Kentropyx borckiana Liophis miliaris
Kentropyx calcarata Liophis poecilogyrus
Kentropyx striata Liophis reginae
Tupinambis teguixin Liophis typhlus
Tropiduridae: Mastigodryas boddaerti
Plica plica Mastigodryas moratoi
Plica umbra Ninia hudsoni
Oxybelis aeneus Siagonodon septemstriatus

Oxybelis fulgidus Tricheilostoma dimidiatum
Oxyrhopus melanogenys Tricheilostoma macrolepis
Oxyrhopus occipitalis (formerly formosus in Typhlopidae:
Guyana) Typhlops brongersmianus
Oxyrhopus petolarius Typhlops minuisquamus
Philodryas olfersii Typhlops reticulatus
Philodryas viridissimus Viperidae:
Phimophis guianensis Bothriopsis bilineata
Pseudoboa coronata Bothriopsis taeniata
Pseudoboa neuwiedii Bothrops atrox
Pseudoeryx plicatilis Bothrops brazili
Pseustes poecilonotus Crotalus durissus
Pseustes sulphureus Lachesis muta
Rhinobothryum lentiginosum Testudinata (15 species of turtles and tortoises):
Sibon nebulatus Chelidae:
Siphlophis cervinus
Chelus fimbriatus
Siphlophis compressus (formerly Tripanurgos)
Mesoclemmys gibba (formerly in Phrynops)
Spilotes pullatus
Phrynops tuberosus
Tantilla melanocephala
Platemys platycephala
Thamnodynastes pallidus
Cheloniidae:
Thamnodynastes ramonriveroi
Xenodon merremii (formerly Waglerophis) Caretta caretta
Xenodon rabdocephalus Chelonia mydas
Xenodon severus Eretmochelys imbricata
Xenopholis scalaris Lepidochelys olivacea
Xenoxybelis argenteus Dermochelyidae:
Elapidae: Dermochelys coriacea
Micrurus averyi Emydidae:
Micrurus collaris (formerly Leptomicrurus) Rhinoclemmys punctularia
Micrurus hemprichii Kinosternidae:
Micrurus isozonus Kinosternon scorpioides
Micrurus lemniscatus Podocnemididae (formerly in Pelomedusidae):
Micrurus psyches Podocnemis expansa
Micrurus surinamensis Podocnemis unifilis
Leptotyphlopidae (genera formerly Leptoty- Testudinidae:
phlops): Chelonoidis carbonaria (formerly Geochelone)
Epictia albifrons (formerly tenellus) Chelonoidis denticulata

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Amphibians and reptiles of Guyana, South America: illustrated

keys, annotated species accounts, and a biogeographic synopsis

BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the

Amphibians and reptiles of Guyana, South America: illustrated keys,

Abstract.—Guyana has a very distinctive herpetofauna. In this ﬁrst ever

Keywords: amphibians, biogeography, Guyana, herpetology, keys, rep-

Introduction 1–40). A total of 78 of these species were

sus, Typhlops lumbricalis, and others. the caecilian Typhlonectes compressicauda

is the largest fauna known from a single examination. Nevertheless, we examined

Dubulay Ranch (5840 0 55 00 N, 57851 0 when we stepped on them, and animal

FMNH: Field Museum of Natural ULAM (¼ULABG in some papers):

(each side) . . . . . . . . . . . . . . . . . . . . . 25b. No dorsolateral fold or ridge . . . . 26

20 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . 8b. Not as 8a; adults larger; toes not

very prominent . . . . . . . . . . . . . . . . . . no chevrons on back; 6–9 vomerine

3b. Skin of usual thinness of most brown . . . . . . . . . . . . . . . . . . . . . . . . . . .

14b. Small to moderate tubercles or and inner side of tarsus . . . . . . . . . .

3a (2). The largest Phyllomedusa, 2b. Absence (usually) of bright ﬂash-

snout to groin; bright red spots in yellowish, marbled with black . . . .

on chin fused . . . . . . . . . . . . . . . . . . . . 2a (1). Pupil vertical with lobed mar-

19b. Relatively large (maximum 24b. Vertebral crest not extending

19b. No white snout or tail ring; 27b. Not as 27a . . . . . . . . . . . . . . . . . . . . .

The following key is based largely on

Allophryne ruthveni Gaige, 1926 Allobates femoralis (Boulenger, 1884)

Type material.—The holotype is Type material.—Originally named Pros-

almost solid black, with black blotches, Anomaloglossus roraima

probably originated in South America (irregular circles, ellipses, reticulations);

of the Göteborgs Naturhistoriska Museum 46409–46413, and 46417–46418 from

14333–14336 from ‘‘Maringma tepui, Vouchers for Guyana.—USNM 118230

specimens known to science. One was Vouchers for Guyana.—AMNH A-

Rhaebo guttatus (Schneider, 1799) Type material.—Originally named Bufo

A-166109–166116 and USNM 566038– Camp (AMNH A-166100) was noted

names on the other specimens of similar Comments.—After we had reviewed

to change them in their paper because 58.96 00 W; identiﬁcation conﬁrmed by Cas-

Hyalinobatrachium mondolﬁi Hyalinobatrachium taylori (Goin, 1968)

Vouchers for Guyana.—The holotype Castroviejo-Fisher et al. (2009) for addi-

Vouchers for Guyana.—The holotype; mostly yellow-green dorsally with four

Pristimantis jester Means & Savage, 2007 Pristimantis marmoratus

Coloration in life.—The following is mung Massif, near Falls Camp, Potaro-

stripe also have a mid-ventral, off-white Dendrobatidae

Vouchers for Guyana.—AMNH A- from preservative for a few minutes and

Distribution.—This species is known Distribution.—‘‘Lesser Antilles; likely

Eleutherodactylus johnstonei Barbour, 1914 Hemiphractidae

primarily on the basis of DNA sequence plateau of Mount Ayanganna, Guyana

561835 from Mount Ayanganna; and Vouchers for Guyana.—ROM 39477–

Coloration in life.—‘‘The dorsum is dark brown mottling. This species is similar in

Dendropsophus grandisonae (Goin, 1966a) Vouchers for Guyana.—AMNH A-

Vouchers for Guyana.—AMNH A- USNM 497705–497714, and USNM

Hypsiboas sibleszi (Rivero, 1972) Lysapsus laevis Parker, 1935

Phyllomedusa hypochondrialis Phyllomedusa tarsius (Cope, 1869)

Vouchers for Guyana.—AMNH A- Forest Reserve (various localities); and

Brazil; also Amazonian Bolivia’’ (Frost Reserve, Siparuni River, 4845 0 17 00 N,

tan, with dark brown stripes (two each Distribution.—‘‘Non-forest habitats of

Scinax cf. x-signatus (Spix, 1824) Voucher for Guyana.—AMNH A-

with tan or yellowish tan markings (body, Vouchers for Guyana.—AMNH A-

Coloration in life.—The following is Leptodactylus fuscus (Schneider, 1799)

Coloration in life.—The following is chest, and in some individuals the anterior

probably from Salvador, Bahia, Brazil Ranch; AMNH A-166384–166393 and

Leptodactylus mystaceus (Spix, 1824)

Vouchers for Guyana.—AMNH A- 057859 0 W); and USNM 531542–531557