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, Amoro´s (1998), Starace (1998), Gorzula & 1 amphisbaenian, 4 crocodilians, 33 liz- Sen˜aris (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), Sen˜aris & 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. 320 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 VOLUME 125, NUMBER 4 321

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 7.400 ; 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 4.500 ; 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, Sen˜aris & 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- 322 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 VOLUME 125, NUMBER 4 323 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 (782201400 N, 6082902800 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 (68210N, 588410W 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 & Sen˜aris (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). 324 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 VOLUME 125, NUMBER 4 325

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. 326 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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. VOLUME 125, NUMBER 4 327

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. 328 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Dubulay Ranch (584005500 N, 578510 when we stepped on them, and animal 3200 W; ca. 30 m elevation): CJC and CRT activity in general was low. collected here from 25 Feb–24 Mar 1994 Karanambo (384501000 N, 5981803300 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 (584101400 N, 5785405900 W), ca. 12 mostly savanna habitat, but there is gallery km (linear) WNW the ranch house; a forest along the waterways, especially the 0 Rupununi River. swamp along Aramatani Creek (5840 0 00 0 00 2200 N, 5785502100 W); a spring (583905300 N, Aishalton (2828 31 N, 59819 16 W; ca. 5785403100 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 (583904600 N, 5785302400 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 (580500600 N, forest along the waterways. 5881401400 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. 58230N, 598590W; 2000 m): Mount Ayan- 0 00 0 Konawaruk Camp (5813 07 N, 59802 ganna is the highest peak located entirely 00 43 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 withina1km Ayanganna (58240N, 598570W; 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 Nin˜o 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., VOLUME 125, NUMBER 4 329 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 58250N, 598580W): RDM and AL collected 58050N, 598500W; 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 580603500 N, 5984901500 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 580704600 N, 5984803700 W): RDM and AL were made by RDM, AL, and Samir collected here from 21–26 Oct 2004. Khan,withsomehelpfromguidesand Kaieteur (above Kaieteur Falls; above porters, on the northeast slope (58050N, 450 m; 58100N, 598290W): RPR collected 598510W; 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 (58050N, 598500W; 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 (58120N, 608440W): 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 Pe´rez (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: Estacio´n Biolo´gica de Don˜ana, Upland sites (between 500–1400 m).— Seville, Spain. Locations of these sites are shown on Figs. EBRG: Estacio´n Biologia de Rancho 3 and 4, although none of us spent Grande, Museo, Maracay, Vene- sufficient time at these sites to include zuela. 330 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

FMNH: Field Museum of Natural ULAM (¼ULABG in some papers): History, Chicago, Illinois. Herpetological Collection, Labora- IBSP: Instituto Butantan, Sa˜o Paulo, tory of Biogeography, Universidad Brazil. de Los Andes, Me´rida, 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 Biolog´ıa, Universi- to Rico, Mayagu¨ez. 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- Muse´um 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 Sa˜o Paulo, Brazil.

burg. NHMW (¼NMW in some papers): ZIUU: see UUZM. Naturhistorisches Museum, Wien ZMB: Museum fu¨r Naturkunde (for- (Vienna), Austria. merly Zoologische Museum), Leibniz- NMNH: National Museum of Natural Institut fu¨r Evolutions- und Biodiver- History, Smithsonian Institution, sita¨tsforschung an der Humboldt- Washington, D.C. (USNM for speci- Universita¨t, 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 fu¨r 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. maybenecessarytoseesomeofthe VOLUME 125, NUMBER 4 331 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)isfully 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- 332 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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, capsinadults...... 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 .IorIV.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) ...... seesepa- 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- ...... seeseparatekeyforAromo- 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 VOLUME 125, NUMBER 4 333

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. 334 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 malesgreaterthan30mm...... 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 VOLUME 125, NUMBER 4 335

(eachside)...... 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 & Sen˜aris 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; 336 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 Rodr´ıguez & Duellman (1994), Grant et 3a (2). Bony spine or protuberance at al. (1997, 2006), Gorzula & Sen˜aris (1999), base of thumb (Fig. 6A) and on Lescure & Marty (2000), Jungfer & Bo¨hme 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 VOLUME 125, NUMBER 4 337

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. 338 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

20mm...... 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 ontoes..... 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 VOLUME 125, NUMBER 4 339 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 mostconspicuous...... 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 sharplypointedsnout...... 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 340 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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- abouttwicewidth...... 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 Stefania tan or brown X across back...... Hemiphractidae ( ) 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) VOLUME 125, NUMBER 4 341

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,anditdirects 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 landsabove900m...... 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 offoot...... below900m...... 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 ...... 4 342 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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).Discsonfingersandtoes 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 anddowntowardupperlip) 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 VOLUME 125, NUMBER 4 343

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,andbellygray..... 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. Skin on back smooth or 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 344 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 dinalstripe ...... 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 towardvent...... 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 VOLUME 125, NUMBER 4 345

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 reddishorange...... 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 Rodr´ıguez & 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. Toes about three-fourths ...... 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 ...... 3 346 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 &Sen˜aris (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, VOLUME 125, NUMBER 4 347

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- gleofjaws...... 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 whitedots...... 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- orangebars...... ed, rounded tips; toes without ...... Leptodactylus knudseni,p.423 fringes on sides; back very rough 13a (10). A dorsolateral fold on each withwarts...... 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- 348 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 onthroat...... 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 acrossbody...... 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) VOLUME 125, NUMBER 4 349

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- sostronglykeeled...... 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 350 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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.Topofheadwithsomelarge 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 VOLUME 125, NUMBER 4 351

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. 352 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 VOLUME 125, NUMBER 4 353

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,andflat ...... (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 . . . . dinalridges...... 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. 471 354 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 VOLUME 125, NUMBER 4 355

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) transverserowsonsideoftail..... 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 notlongitudinalrows...... 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. 449 356 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 VOLUME 125, NUMBER 4 357

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 bodyscalesorslightlywidened; 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. 518 358 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 frontofeye)...... 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 VOLUME 125, NUMBER 4 359

(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), entbehindchininmiddleof 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. 495 360 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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. Wu¨ster 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 VOLUME 125, NUMBER 4 361

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...... seeseparatekey 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 ...... 68 362 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 adjacentrows...... 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 VOLUME 125, NUMBER 4 363

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. 505 364 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 on neck...... than 20 ... Atractus tamessari, p. 484 ...... 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 VOLUME 125, NUMBER 4 365

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 cornersofblotchesatventrals.. 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. 499 366 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 . . . . aboutaswideasredonesorwider ...... 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). VOLUME 125, NUMBER 4 367

1a. Arms and legs flattened as oar-like 7a (6). Webbing between toes; legs flippers (sea turtles) ...... 2 not elephantine or columnar ...... 8 1b. Notas1a...... 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(viewedfromabove)...... 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 .. 13 368 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 VOLUME 125, NUMBER 4 369

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. 370 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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,orEpipedobates, 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, VOLUME 125, NUMBER 4 371

Upper Demerara-Berbice Region, Guyana did not note the partial oblique pale lateral (58090N, 588410W, 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. 372 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 thighandshankcreamyyellow.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 (581100600 N, 5982805100 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 (0581203700 N, 06083305900 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- (058240N, 598570W, 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 VOLUME 125, NUMBER 4 373 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 (58100N, 608470W; 1950 m), stripe. A few white spots (covering small Estado Bol´ıvar, 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 (0581201600 N, 0608340 dark brown; tympanum area light cream- 3900 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- browntodarkgraywithdarkbrownto 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 maleswithgraytoverydarkgraythroat, 374 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 Bol´ıvar, 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 VOLUME 125, NUMBER 4 375 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 rivie`re Ampiyacu (Dpto Loreto, Pe´rou)’’ 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, Para´ , Amapa´ (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 Taracua´ , Rio Uaupe´s, Brazil. The website 376 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of the Go¨teborgs 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,orwasintheearlystagesofa 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 (5824.110N, 59857.410W). 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 VOLUME 125, NUMBER 4 377

14333–14336 from ‘‘Maringma tepui, Vouchers for Guyana.—USNM 118230 southeast slope, 1376 m asl (0581201600 N, is from Mount Roraima within the Guya- 06083403900 W)’’ (Kok 2009a:49); USNM na portion. In addition, Sen˜aris et al. 549306 from the north slope of Mount (2005:67) included the following: ‘‘GUYA- Roraima, 058160N, 608440W, at 1300 m NA-BRAZIL: Wei-Assipu-tepui, elevation. (581301 00 N, 6084201900 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 (0581205900 N, Comments.—Individuals on Maringma 06083500500 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 378 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 Sen˜aris, 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 (078160N, 588430W); USNM Sima de los Gua´ charos, 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 58130100 N, 6084201900 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’’ (Sen˜aris 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’’ (Sen˜aris 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- VOLUME 125, NUMBER 4 379

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 (0384500800 N, 5981803600 W); USNM 497696 from Mount Ayanganna; and USNM from Lethem, Takutu River (0382203800 N, 118071 from Kartabo. 5984801300 W); USNM 497697 from Kaie- Coloration in life.—The following is teur National Park (near paved airstrip); basedonHoogmoed(1977)andAL’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 &Ve´lez-Rodriquez, 2007a marina based on an illustration (Frost Plate 3E, F 2011). The type locality was restricted to Surinam by Mu¨ller & 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 (48560N, 558100W), 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 (68270N, 588150W); USNM 162968– 566034 from Dubulay Ranch; AMNH A- 162974 from Enmore Estate (68440N, 166096–166105 and USNM 566035– 578590W); USNM 291104 from Kato, 566037 from Berbice River Camp; AMNH 380 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 (48400N, 588390W; paratypes); USNM of warts; chin, throat and chest black with 164164 from Mabaruma Compound gray dots; anterior belly pale yellow with (088120N, 598470W); 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 (058200N, Comments.—These toads are part of a 608460W); 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 VOLUME 125, NUMBER 4 381 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- basedontheAMNHspecimensfrom 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 382 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON to change them in their paper because 58.9600 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 (Ayarzagu¨ena, 1992) Plate 4B–E Type material.—The holotype is RMNH 4463 from the Saramacca River Type material.—The original name was and vicinity (Gasso´ 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 Yuruan´ı, 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 (05809019.3000 N, 588410 la, and in the Amazonian lowlands of VOLUME 125, NUMBER 4 383

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). Maiko Creek (05809019.30000 N, Vouchers for Guyana.—SMNS 12242– 58841058.9600 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, Ya´ nez-Mun˜oz 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 Ya´ nez-Mun˜oz 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 (Sen˜aris & Ayar- parietal peritonea white; iris yellow to dull zagu¨ena 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- Ya´ nez-Mun˜oz 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) Sen˜aris & Ayarzagu¨ena, 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 Can˜o Acoima, afluente Guyana. Frost (2011) noted Hoogmoed’s del R´ıo Grande (88220N, 618320W), 15 comment in the 1985 edition of the msnm., estribaciones de la Serran´ıade amphibian checklist volume that the type Imataca, Estado Delta Amacuro, Vene- locality is within an area that is in dispute zuela’’ (Sen˜aris & Ayarzagu¨ena 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). 384 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 (Ayarzagu¨ena, 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 (58560N, 628340W; 1850 m), Bol´ıvar, Venezuela. Distribution.—‘‘Known from Auyante- pui, Atapare, and Sierra de Lema, 1800– Vitreorana helenae (Ayarzagu¨ena, 1992) 1900 m elevation, state of Bol´ıvar, 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 Yuruan´ı, Bol´ıvar, Venezuela. Vouchers for Guyana.—IRSNB 13974– Distribution.—‘‘Known from the type 13978 from Kaieteur National Park; ROM locality...and Salto Karuay, Bol´ıvar, 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 VOLUME 125, NUMBER 4 385 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 phylogenetic analyses with related frogs, Ceratophrys cornuta (Linnaeus, 1758) 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). Vouchers for Guyana.—AMNH A- 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 (0580000800 N, 598520 labial region. In brown individuals, the 4700 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 386 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 Lo¨e Creek and Sau¨l) 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 VOLUME 125, NUMBER 4 387 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 (0580502500 N, 5985001800 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). 388 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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) (05804003.300 N, 59851041.800 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 (0682905700 N, 0588130 another specimen (USNM 563633) was 0600 W). VOLUME 125, NUMBER 4 389

Coloration in life.—The following is mung Massif, near Falls Camp, Potaro- from Kok & Kalamandeen (2008:242), Siparuni District, Guyana (0580502500 N, based on observations in Kaieteur Nation- 5985001800 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, Regio´n 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 Uaire´n, 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 390 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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, VOLUME 125, NUMBER 4 391

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). Type material.—The original name was Hyla Trivittata and RMNH 1836 from Rio Tefe´, 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). Ha¨upl and Tiedemann, 1978. Vouchers for Guyana.—Silverstone Distribution.—‘‘Guianan Orinoco drain- (1976:48) listed 11 localities scattered age of Venezuela north to the R´ıo 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). 392 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 Bo¨hme 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- &Bo¨hme (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- 49301–49328 from Shudikar-wau and Jungfer & Bo¨hme, 2004 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 (58580N, 588320W) (Silverstone 1975:45–46) are for specimens ...7 m above sea level’’ (Jungfer & Bo¨hme 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 VOLUME 125, NUMBER 4 393 specimens are similar in pattern and color (Bol´ıvar), 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 Amapa´ , 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 (068160N, color) but ‘‘the tympanum is larger than in 0608300W); and USNM 549315 from the the type material of D. azureus. Provision- N slope of Mt. Roraima (58170N, ally, I include the Shudikar-wau popula- 608450W). 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 ular lineage should be split 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-Amoro´s, 2008 and V, northern slopes of Mount Rorai- Plate 6A ma, Guyana (58170N, 608460W, 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 (058060 tains, Guyana, nearby Venezuela 3500 N, 05984901500 W, 1234 m). 394 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 Jose´ (Costa Rica), Curac¸ao, 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 VOLUME 125, NUMBER 4 395 primarily on the basis of DNA sequence plateau of Mount Ayanganna, Guyana data, although Frost (2011) commented (058240N, 0598570W; 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 plateau of Mount Ayanganna, Guyana Stefania ackawaio 0 0 MacCulloch & Lathrop, 2002 (05824 N, 059857 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– 396 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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; (058240N, 0598570W; 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- VOLUME 125, NUMBER 4 397 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. 398 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 Sen˜aris 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 Uaire´n, 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 VOLUME 125, NUMBER 4 399 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, Peru´, 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 Bele´m, 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. Coloration in life are now Trachycephalus; 4) frogs of the .—‘‘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. 400 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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.). (68150N, 0598050W); 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 VOLUME 125, NUMBER 4 401

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). Vouchers for Guyana.—AMNH A- Dendropsophus minusculus (Rivero, 1971) 141033–141036, A-141038–141039, A- Plate 7C 167097–167098, and USNM 566097 from Dubulay Ranch; AMNH A-166209– Type material.—The original name was 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 Bele´m, Brazil’’ (Frost 2011). Coloration in life.—The following is Vouchers for Guyana.—AMNH A- 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 Coloration in life.—The following is subtle darker tan markings. One individual based on AMNH and USNM specimens 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 402 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 058170N. 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 Type material.—The holotype is 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 (058380N, 608440W; elevation Type material.—The original name was 1480 m), Rupununi District, Guyana. Rana boans and the types, from unspecified VOLUME 125, NUMBER 4 403 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 (68230N, 166143–166144 and USNM 566061 from 0598050W); 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 (381505200 N, 05984303400 W); USNM Coloration in life.—The following notes 497811 from Karanambo; USNM 497812– are based on AMNH and USNM speci- 497813 from Moco Moco (381801200 N, mens from Konawaruk Camp (CJC field 05981803600 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). 404 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 (88120N, 0598470W); 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 (482501200 N, 05885005800 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 (Gu¨nther, 1858) to as H. cinerascens. Type material.—The original name was 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 (482001500 N, 05884903700 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 VOLUME 125, NUMBER 4 405 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 Uaire´n, Serran´ıa 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– (580000800 N, 05985204700 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 406 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON edge. Limbs with dark brown crossbands. Hypsiboas multifasciatus (Gu¨nther, 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 Bele´m, Para´ , 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 Ceara´ orly to the nostrils’’ (MacCulloch & and Goia´ s, Brazil, limited by the Amazon, Lathrop 2005:25–26). the Orinoco, and Rio Negro’’ (Frost 2011). Vouchers for Guyana.—AMNH A- 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 (0682905700 N, 05881300600 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 (581005100 N, 598280 566099 from Konawaruk Camp; ROM 5700 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; USNM 118043 from Bartica; USNM

known from only two localities. 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 (381801200 N, 05983900000 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 VOLUME 125, NUMBER 4 407 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 Amapa´ , Brazil, to Guain´ıa, Colombia’’ (Frost Hypsiboas roraima 2011). (Duellman & Hoogmoed, 1992) Vouchers for Guyana.—The holotype. Plate 9E, F Coloration in life.—The following is 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 1979.560, an adult female from the north light green to yellowish green, with small 0 0 reddish brown to yellow spots; dark slope of Mt. Roraima (05838 N, 60844 W; reddishbrownbaredgedwithyellow elev. 1480 m), Rupununi District, Guyana. According to Warren (1973) the latitude across top of head from one eyelid to the should be 058170N, 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, Bol´ıvar, 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 408 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON stripe never present on individuals with andV,andFingerIV.Diffuselargerdark 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. Uaire´n, 1300–1400 m; Serran´ıa de Lema,’’ Distribution.—‘‘Southwestern Guyana Bol´ıvar, 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 VOLUME 125, NUMBER 4 409 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 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 (Amapa´ ), 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 410 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 Can˜o Guacaya´ , 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 (583703000 N, in adjacent Brazil and Surinam’’ (Frost 6081404200 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- VOLUME 125, NUMBER 4 411 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 (Dume´ril & Bibron, 1841) Type material.—The holotype is Plate 10F MZUSP 69852, an adult male from Reserva Florestal Adolfo Ducke (28550S, Type material.—The original name was 598590W), 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 (Guibe´ 1950). the Guiana Shield and in the Amazon Distribution.—‘‘It occurs on the Guiana Basin, from east and north of State of Para´ 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- Uaupe´s’’ (Jungfer & Ho¨dl 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 412 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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.—Theholotypeis but each of the latter usually has a white NHMW 16492 (Ha¨upl & Tiedemann spot that makes it appear more conspicu- 1978) from Mana´ us, 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- (058240N, 0598570W; 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 VOLUME 125, NUMBER 4 413

(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 0 00 0 00 Phyllomedusa bicolor (Boddaert, 1772) Airport (06829 57 N, 058813 06 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). 414 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 R´ıo 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 (68440N, Guiana, and northeastern Brazil to Co- 0578590W). 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 VOLUME 125, NUMBER 4 415 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 on the R´ıo Aguarico, Provincia Napo, the underside of the limbs; iris golden 0 0 yellow with a horizontal chestnut bar. Ecuador (0802 N, 76858 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 R´ıo Napo, R´ıo Maran˜on, and R´ıo Ucaya- gigantic tadpole’’ (Duellman 2005:261). li); reported from 70–90 km north of Manaus, Brazil; possibly in Amazonian southeastern Colombia’’ (Frost 2011). Vouchers for Guyana.—AMNH A- 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 Para´ , 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 basedontheAMNHspecimensfrom 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 416 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Brazil; also Amazonian Bolivia’’ (Frost Reserve, Siparuni River, 484501700 N, 2011). 05080102800 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 (68440N, 0578590W); and ventral surface reddish brown, but the USNM 497718 from Kumu, at base of throat is white in females, grayish in males, 0 00 0 Kanuku Mountains (3815 52 N, 059843 and the belly is white; iris gold to silver 00 34 W). with a horizontal brown band across the Coloration in life.—The following is middle. based on the AMNH and USNM speci- 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 (68440N, 0578590W); Surinam is number Z. 354 of the Zoolog- USNM 164165–164177 from Mabaruma ical Museum of Amsterdam (in the process Compound (88120N, 0598470W); USNM of being transferred to the RMNH). 284390–284391 from Takutu Mountains Distribution.—‘‘Interior of the Guianas’’ (68150N, 0598050W); 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- (68480N, 0588100W); 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 VOLUME 125, NUMBER 4 417 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. Coloration in life.—The following is 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 Vouchers for Guyana.—AMNH A- 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- 418 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 tanwithminutedarkbrownfleckson 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 Ame´rique, 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 (2008:204) provide additional details on catalog); ROM 11081 and 11090 from the vicinity of Georgetown; USNM coloration. 127888 from Akyma, Demerara River (58530N, 0598180W); and USNM 162942 from Enmore Estate (68440N, 0578 Trachycephalus coriaceus (Peters, 1867) 590W). Plate 12E Coloration in life.—The following is Type material.—The original name was 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 (058100N, 598280W), Mazaruni- brown markings (including large dorsal Potaro District, Guyana. hourglass); sides yellow; black patch on VOLUME 125, NUMBER 4 419 shoulder; chin, throat, and chest yellow; from Mission of Santo Antonio do Prata abdomen of yellow warts surrounded by at the River Maracana˜(interior of the reticulation of bright orangish red; ventral State of Para´ ). 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 basedonfrogsfromFrenchGuiana 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 Parana´ (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 (68440N, 0578590W); and USNM 164179 from Mabaruma (88120N, Trachycephalus resinifictrix (Goeldi, 1907) 0 Plate 12F 059847 W). Coloration in life.—The following is 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, 420 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 Maran˜on River, Peru. Distribution.—‘‘Amazonian forests of southeastern Colombia and Venezuela east through the Guianas and south into Adenomera andreae Mu¨ller, 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. Braganc¸abahn) Staat Para`, Vouchers for Guyana.—AMNH A- Brasilien’’ (Mu¨ller 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 VOLUME 125, NUMBER 4 421

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 blackwithnoorabarelydiscernible 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 basedonfrogsfromFrenchGuiana 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). 422 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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;wartsonanteriorpartofbody 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 Vouchers for Guyana.—AMNH A- 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 & Sen˜aris 1999; also compared with Enmore Estate (68440N, 0578590W); 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 VOLUME 125, NUMBER 4 423 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 Sa´ , 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 (484501700 N, 5980102800 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 Sa´ (Heyer & de Sa´ 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– (08240S, 768370W; 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). 424 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Coloration in life.—The following is chest, and in some individuals the anterior basedontheAMNHspecimensfrom 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 (Guain´ıa), 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), VOLUME 125, NUMBER 4 425 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- basedontheAMNHandUSNM 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 posterior surfaces of the arms; rear of of lower legs translucent gray. Lescure & 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) Type material.—The original name was 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 Solimo˜es, 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). 426 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Vouchers for Guyana.—AMNH A- 0578590W); and USNM 531542–531557 70118–20 from Shudikar-wau, a headwater from Iwokrama Forest Reserve (several stream of the Essequibo River (18150N, localities). 588350W); 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 (68440N, from Berbice River Camp (CJC field notes, VOLUME 125, NUMBER 4 427

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 and the Guianas, and north-central Ro- Plate 15A 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 (38200N, na; USNM 118063–118064 and 129537– 0598390W); 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. (68440N, 0578590W). 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 428 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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. VOLUME 125, NUMBER 4 429

Comments.—The vouchers for Guyana Distribution.—‘‘Disjunct regions of east- compare favorably with characters given ern Colombia, northern Brazil (Roraima), by Gorzula & Sen˜aris (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). Vouchers for Guyana.—AMNH A- 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 (381801200 N, River, New Grenada (Malnate 1971). 05983900000 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 Curac¸ao, 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. creamtoorangishtan,thinvertebral 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 430 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 Humaita´ , Igarape do Banheiro (078310S, 638000W). Distribution.—Known only from the 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. Vouchers for Guyana.—AMNH A- 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 whitestripeontheposteriorfemoral 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). VOLUME 125, NUMBER 4 431

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. Coloration in life.—The following is 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– 432 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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: VOLUME 125, NUMBER 4 433

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. Type material.—The holotype is BMNH 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 Wacara´ , Vaupe´s Colombia. This site is Distribution.—The region of the Paka- east of Mitu´ and close to the Brazilian raima Mountains of western Guyana and border at approximately 018090N, southeastern Venezuela, at elevations of 698550W. approximately 400-1200 m; probably also Distribution.—‘‘From Vaupe´s and Am- in adjacent Brazil (Campbell & Clarke azonas, Colombia, across the lowlands of 1998). southern Venezuela and through the Vouchers for Guyana.—Possibly the GuiananregiontoAmapa´ s, 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 434 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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.—Theholotypeis 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). Vouchers for Guyana.—The holotype; 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 R´ıo 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 Timbo´, 44778–44779, and 45271 from Mount Vaupe´s, 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 VOLUME 125, NUMBER 4 435 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 Type material.—The original name was 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 Vouchers for Guyana.—AMNH A- 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 Amazoˆnia, Munic´ıpio 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). Vouchers for Guyana.—AMNH A- 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- 436 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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. Type material.—The original name was 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 (0581801000 N, 05984905500 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 (381505200 N, The most recent taxonomic revision, 05984303400 W); and USNM 531617– consistent with the phylogeny reconstruc- 531619 from Iwokrama Forest Reserve, tion, concluded that globally there are nine Sipuruni River (484501700 N, 05980102800 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. VOLUME 125, NUMBER 4 437

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; Para´ and Maranha˜o, 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, 28420N 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 Type material.—The holotype is 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. 438 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Rhinatrematidae Vouchers for Guyana.—The holotype; BMNH 1976.102 from Winiperu, (68100N, Frost et al. (2006) and Wilkinson et al. 588340W); 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 Konawaruk Camp (CJC field notes, 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 (48200N, 588480W; 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 (58080N, 598250W) (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 VOLUME 125, NUMBER 4 439 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, R´ıo Chicana´ n, Bol´ıvar, Venezuela. na. Distribution.—‘‘State of Bol´ıvar, 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 trees’’ (Nussbaum & Hoogmoed [tentacular apertures] and nares lie within 1979:225). 440 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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; (Dume´ril & 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 VOLUME 125, NUMBER 4 441 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) Type material.—The original name was 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. Coloration in life.—The following is 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 442 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 Solimoe¨ns [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.Fromthree to five large dark blotches Plate 19C on the sides of the jaws... Ear coverlets black, Type material.—The original name was 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.’’ Gu¨nther 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. VOLUME 125, NUMBER 4 443

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].’’ 444 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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- (88120N, 0598470W). stricted the type locality to the confluence Coloration in life.—The following is 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 notes, 1994). They are black and white, South America...Venezuela, Trinidad, 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.—Theholotypeis 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.—Theholotypeis 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 2850 North and 598 East’’’ (Gans AMNH R-60907 from Marudi River; 1963:13). VOLUME 125, NUMBER 4 445

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 al. 2003), with the eight groups of Frost &

RMNH 15191 (from Sipaliwini) was found 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 & Sen˜aris 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- 446 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 Jonne`s, 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 (68440N, 0578590W); and USNM photograph of the specimen was sent to 291268 and 497791 from Georgetown. VOLUME 125, NUMBER 4 447

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 Mabaruma Compound (88120N, R-138064 from Yupukari; AMNH R- 0598470W); 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 (0682905700 N, 05881300600 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 448 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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- Vouchers for Guyana.—AMNH R- 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 Coloration in life.—The following is hoogmoedi, Echinosaura sulcarostrum, based on the USNM specimen from Kaieteurosaurus hindsi,andPantepuisau- 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- (058240N, 0598570W; 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 VOLUME 125, NUMBER 4 449 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 (58170N, 0608450W; 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 (581206000 N, Vouchers for Guyana.—AMNH R- 6083500600 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 450 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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.’’ Type material.—The original name was Cercosaura (Pantodactylus) argulus and the holotype is ZMB 4555 from ‘‘moun- Bachia flavescens (Bonnaterre, 1789) tainous regions around Santa Fe´ de Plate 20F Bogota´ , 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 Vouchers for Guyana.—AMNH R- 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. Vouchers for Guyana.—AMNH R- 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. VOLUME 125, NUMBER 4 451

Cercosaura ocellata Wagler, 1830 Distribution.—Known only from Guya- Plate 21D na. Vouchers for Guyana.—The holotype; 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; 0682905700 N, 0588130 al. 2006:400). 0600 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

based on AMNH and USNM specimens 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 (078220N, 0608290W; 100 m). ventrolateral) light stripe, on some pres- 452 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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. asinglespecies,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 basedontheAMNHspecimensfrom 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.—Theholotypeis 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, VOLUME 125, NUMBER 4 453 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 (0682905700 N, interest is the Brazilian sample from Ilha 05881300600 W). da Maraca´ , 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 Maraca´ 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 Maraca´ specimens 454 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(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 Maraca´ . 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 wheretherearesympatricspeciesof are not adequately diagnosed as yet. Gymnophthalmus.

Iphisa elegans Gray, 1851 Gymnophthalmus vanzoi Carvalho, 1997 Plate 22E Plate 22D Type material.—The holotype is BMNH 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- Bele´m, Para, Brazil (Avila-Pires 1995). icoera (038200N, 618240W). 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 basedonAMNHandUSNMspecimens 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 VOLUME 125, NUMBER 4 455 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. Distribution.—Known only from the type locality. Leposoma percarinatum (Mu¨ller, 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 Braganc¸abahn), Para`, 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). Vouchers for Guyana.—AMNH R- 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 Timehri Airport (0682905700 N, Reserve (various sites); USNM 535792 05881300600 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 (0682905700 N, based on the AMNH voucher, an adult 05881300600 W). 456 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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- Type material.—The holotype is BMNH 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 (Bol´ıvar), 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 (58170N, 0608450W; 800 m). VOLUME 125, NUMBER 4 457

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 (0581205700 N, and one or two additional tepuis in 06083500700 W; 2080 m). Venezuela (Gorzula & Sen˜aris 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). 458 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Coloration in life.—The following is Corytophanidae). Representatives of this from Gorzula & Sen˜aris (1999 family occur in ‘‘tropical and subtropical [1998]:142), based on specimens from America from the southwestern United Cerro Kukena´ n, 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) Type material.—The original name was 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 Creek, Surinam. Dunoon, Guyana. 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 Vouchers for Guyana.—AMNH R- the only specimen known from Guyana. 14111–14114, R-21289, and R-21330 from Coloration in life.—The following is 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- Coloration in life.—The following is 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 VOLUME 125, NUMBER 4 459 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– 0 The species is primarily arboreal. 163015 from Enmore Estate (6844 N, 0578590W). Coloration in life.—The following is 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- (Lace´pe`de 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, 460 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 68440N, 0578590W; 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- VOLUME 125, NUMBER 4 461 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 Maran˜on, 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. Dume´ril & 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 Mamore´, 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 462 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 (68230N, Type material.—Theholotypeis 0588420W); USNM 497792 from CEIBA MNHN 2340, a female from ‘‘South 0 00 0 00 (6829 55 N, 058813 04 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 Coloration in life.—The following is 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. VOLUME 125, NUMBER 4 463

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 Vouchers for Guyana.—AMNH R- Rita, Munic´ıpio de Maraa˜, left margin of 21315, R-32279–32281, and R-137404– Rio Japura´ (Lago Parica´ ), 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 (68440N, 0578590W); 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 (88120N, 0598470W); 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 464 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON dorsolateral light stripe may be present; from Marudi; USNM 570538 from Gunn’s arms, legs, side of face, and side of body Strip (0183604600 N, 05883801500 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.—Theholotypeis Sphaerodactylus amazonicus and the holo- MZUSP 52866, a male from Brasil, type from Amazonas, Manaos, Lago Roraima: Ilha de Maraca´ . 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 VOLUME 125, NUMBER 4 465

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] basedonAMNHandUSNMspecimens WSW Mabura Hill), 120 m elevation from Karanambo (CJC field notes, 1992). To (581300700 N, 5980204300 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- onthesidesoftheneckandayellowhood.’’ 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), 466 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 Amapa´ , 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- Vouchers for Guyana.—AMNH R- 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 137411 from Kartabo; AMNH R-17687 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- 0 0 and R-131999–132001 from Isheartun; ruma Compound (08812 N, 059847 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 VOLUME 125, NUMBER 4 467 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 & Sen˜aris 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 & Sen˜aris discuss evidence Ecuador, and Peru’’ (Avila-Pires that this was introduced into El Manteco, 1995:304). Bol´ıvar 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 (68–0N, 141886–141887 from Dubulay Ranch; 588350W); 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 468 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 Sen˜aris (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 VOLUME 125, NUMBER 4 469 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 (88120N, 0598470W); 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 (68230N, 0588410W); 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 470 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 (68230N, largest males with green obscuring the 0588410W); 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, VOLUME 125, NUMBER 4 471 tan, or greenish yellow on abdomen, USNM 163021–163052 from Enmore preanal area, hind legs, base of tail Estate (68440N, 0578590W). In addition, (becominggrayneartip).Adultmales 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.—Thisisanall-femalespe- one of their ancestors (reviewed by Cole & ciesthathadanoriginresultingfrom 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 Type material.—The original name was Kentropyx calcaratus based on material Centropyx Borckiana and the lectotype is from flumen Itapicuru´ 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 472 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 (68440N, the following notes paraphrased from 0578590W); 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); VOLUME 125, NUMBER 4 473 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 This is one of the families of iguanian Tupinambis teguixin (Linnaeus, 1758) 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. 474 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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/Solimo˜es and Rio Japura´ , 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 0 0 (8812 N, 059847 W). from Paramakatoi; ROM 43206 from

Coloration in life.—The following is Kaieteur Plateau; USNM 164189 from based on AMNH and USNM specimens Mabaruma Compound (88120N, from Konawaruk Camp (CJC field notes, 0598470W); USNM 326119–326120 from 1998). Brownish green with black mark- Kwakwani (58300N, 0588000W); 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, Konawaruk Camp (CJC field notes, 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. VOLUME 125, NUMBER 4 475

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 (68440N, 0578590W); 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 Biological Station (0682905700 N, Guiana, and Brazil’’ (Avila-Pires 05881300600 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 476 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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). Vouchers for Guyana.—AMNH R- 15133–15135 and R-21317 from Kartabo; Species Accounts: Reptilia: AMNH R-61306 from Isheartun; AMNH Squamata: Snakes R-61390–61391 from Karanambo; 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, Biological Station (0682905700 N, and Typhlopidae. The modern snakes (of 05881301600 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- VOLUME 125, NUMBER 4 477 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) Type material.—The original name was Type material.—The original name was Anguis Scytale and the holotype, which Typhlops squamosus based on material 478 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON from Cayenne. Hahn (1980) stated the Boa constrictor Linnaeus, 1758 holotype was at the MNHN, now lost. Plate 28C Distribution.—Atlantic lowlands of Type material South America from French Guiana, .—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 Vouchers for Guyana.—AMNH R- lacking for the types. 25051 from Kamakusa. Distribution.—‘‘From northern Me´xico, 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 Andre´s 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 (881200000 N, 05984700000 W); USNM 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. Coloration in life.—The following is 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 VOLUME 125, NUMBER 4 479 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 Bol´ıvar 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 Amapa´ ,Para´ , 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); Vouchers for Guyana.—AMNH R-8102 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. 480 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 Vouchers for Guyana.—AMNH R-8102 data. from Kalacoon; AMNH R-21285 and R- We examined each of the Guyanan 31523 from Kartabo; AMNH R-98186 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.Thisalso 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 VOLUME 125, NUMBER 4 481

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 482 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 amonggeneraforsometime,which underscores the difficulties with colubrid Apostolepis nigrolineata (Peters, 1869b) classification. Type material.—The original name was 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 oftheBMNHprovideduswiththe a problem with nonmonophyly of the following details on the holotype for the traditional Liophis, Erythrolamprus, junior synonym Apostolepis quinquelineata and Umbrivaga.Weretainuseof (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 VOLUME 125, NUMBER 4 483 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, 0 Atractus favae (Filippi, 1840) Guiana, 750 ,’’ 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 484 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 Chimanta´ Tepui, distribution than had been known and is Bol´ıvar, 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, 7500–80000’’ (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,’ (0581503000 N, 608430 2640, an adult male from along a tributary 3000 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 (580800900 N, 5982502800 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 VOLUME 125, NUMBER 4 485 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- (Dume´ril, Bibron, & Dume´ril, 1854) lay Ranch. Plate 29D Coloration in life.—The following is based on the USNM specimen from Type material.—The original name was Dubulay Ranch (CJC field notes, 1994). 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.’’ Dubulay Ranch (CJC field notes, 1994). 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 486 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(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 (058120N, Coluber exoletus and the holotype is 0608350W; 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). VOLUME 125, NUMBER 4 487

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- (582300000 N, 06084600000 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. Vouchers for Guyana.—AMNH R-8216 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 (881200000 N, 05984700000 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 488 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 basedonMNHNmaterial 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 (580000800 N, 05985204700 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- VOLUME 125, NUMBER 4 489 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 (881200000 N, 05984700000 W); USNM 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 490 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

flecks, especially posteriorly. The iris is across the dorsal... [surface]...[mostly on] reddish brown.’’ the sides...The belly is light-coloured.’’

Dipsas copei (Gu¨nther, 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 (058240N, 0598570W; 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 (28070N, 598010W) 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... VOLUME 125, NUMBER 4 491

‘‘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 (Dume´ril, Bibron, & Dume´ril, 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 basedontwo 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 Para´ , 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. ‘‘Muse´e des Pay-Bas [¼ RMNH]’’ (Peters Coloration in life.—The following is 1960:61). from Starace (1998:176), based on speci- Distribution.—‘‘Guianas to Para´ , 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 492 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 each side, never meeting in the middle. The

Coloration in life.—The following is 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) Type material.—The original name was Coluber corais based on a specimen from Drymobius rhombifer (Gu¨nther, 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 VOLUME 125, NUMBER 4 493 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 Vouchers for Guyana.—AMNH R- ‘‘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 (0582000000 N, However, Starace (1998) does not include 06084600000 W; 550 m); and USNM 566261 this species among the snakes known from from Berbice River Camp. French Guiana. Considering the uncer- Coloration in life.—The following is 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- 494 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 (0881200000 N, USNM 84528 from Pomeroon. 05984700000 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. VOLUME 125, NUMBER 4 495

Hydrops triangularis (Wagler, 1824) Imantodes cenchoa (Linnaeus, 1758) Plate 31E Type material.—The original name was 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 Tefe´, at (038210S, 648420W), on to us. the inner mouth of the canal...that con- Distribution.—‘‘...from southern Tam- nects the lake of Tefe´ with the Solimo˜es’’ 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- from Berbice River Camp. site, sometimes alternate. These narrow

Coloration in life.—The following is suddenly and extend upward as scale- based on the AMNH and USNM speci- 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 496 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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: (0682905700 N, 05881300600 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. Type material.—The original name was 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 Type material.—The original name was Amazon, Para´ , 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. VOLUME 125, NUMBER 4 497

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 flanks of the Andes from near Villavicen- Liophis breviceps Cope, 1860a 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 Peru´, 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, R´ıo Mamore´ (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 (881200000 N, 0598470 566269 from Dubulay Ranch; AMNH R- 0000 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 498 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 & Sen˜aris 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 Vouchers for Guyana.—AMNH R- de Joucou´, assez pre`s de la rivie`re 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. VOLUME 125, NUMBER 4 499

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 (881200000 N, 05984700000 W); USNM 549328– speckling, especially on the neck. 549329 from N slope of Mt. Roraima (581700000 N, 6084500000 W); USNM 561439 from Washikura River (881500000 N, 0598440 Mastigodryas boddaerti (Sentzen, 1796) 0000 W); and USNM 573507 from Woko- Plate 32E 0 00 0 00 mung Massif (5800 08 N, 059852 47 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. (881200000 N, 05984700000 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 500 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 Tefe´, and ‘‘the locality brown with grayish tan bands; white spots (038210S, 648420W) 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 Tefe´ with the Sol- with dark brown irregular spots; venter imo˜es’’ (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 & Sen˜aris 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. (383604200 N, 05984003000 W); and USNM Vouchers for Guyana.—The holotype 531701 from Iwokrama Forest Reserve, and the paratype (BMNH 1946.1.15.20), ThreeMileCamp(483705900 N, 0588420 both from the type locality, are the only 5200 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 VOLUME 125, NUMBER 4 501 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 Itza´ , Yucata´ n, 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 Yucata´ n 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 502 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 Type material.—The original name was 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 Solimo¨ens 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 Amapa´ , 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 (58090N, 588420W). 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 VOLUME 125, NUMBER 4 503 from Mabaruma Compound (881200000 N, (very thin) black line below eye, from 05984700000 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. basedontheUSNMspecimenfrom Dubulay Ranch (CJC field notes, 1994). 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 basedonmaterial 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 504 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 (0682905700 N, 0588130 (1946:39) described a specimen from 0600 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); (Dume´ril, Bibron, & Dume´ril, 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 Cumana´ , 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, VOLUME 125, NUMBER 4 505 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 (583000000 N, 05880000000 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 (Gu¨nther, 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 Potos´ı, 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 506 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON pupil, extending out to the external rim Rhinobothryum lentiginosum of the iris.’’ (Scopoli, 1788) Plate 34F Type material.—The original name was 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 R´ıos 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 Japura´ from its mouth (038080S, Coloration in life.—The following is 648660W), to the falls of Araracuara based on Starace’s (1998) description of (008240S, 728170W), 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. VOLUME 125, NUMBER 4 507

Distribution.—‘‘Northern South Amer- specimens from French Guiana. Body ica...lowlandCentralAmericatoMi- dark gray with numerous narrow yellow choaca´ n...and Veracruz, Mexico, bands; irregular row of red scales along including Yucata´ n 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. (881200000 N, 05984700000 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 (482001500 N, 05884903700 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). Maranha˜o [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. 508 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 (881200000 N, 05984700000 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 (881200000 N, 05984700000 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 is the most strongly marked.’’

yellow bands issuing from the belly which is also yellow. [Labials yellow and black.] The tail is often black...[without] the yellow markings.’’ Some individuals are Thamnodynastes pallidus (Linnaeus, 1758) melanistic. Type material.—The original name was 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. Para´ , 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 VOLUME 125, NUMBER 4 509 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 & Sa´ nchez, 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 0 00 0 00 edge of parietals outlines the darker crown (10801 27 N, 64807 47 O[¼W]) macizo del and joins a dark paravertebral line bor- Turimiquire, municipio Freites, Anzoa´ te- dering a light nuchal stripe on the gui, Venezuela, 2130 m (Manzanilla & nape;...the five series of dark dorsal-spots Sa´ nchez 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 & Sen˜aris 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 510 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(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 Type material.—The original name was 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 (138000S, white areas;...head brown above, usually 388300W) (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 Vouchers for Guyana.—AMNH R-8103 light brown, edged with dark; throat cream from Kalacoon. with very little dark pigment, which is usually Coloration in life.—The following is 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) Type material.—The original name was Type material.—The original name was 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); VOLUME 125, NUMBER 4 511

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 (881200000 N, 0598470 location of the type specimen are unknown 0000 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 Vouchers for Guyana.—AMNH R- 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 Coloration in life.—The following is 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 Can˜avieras, Matta de S. Joa˜o, 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, 512 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON above). Often when disturbed, an individual ian border (18400N, 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 VOLUME 125, NUMBER 4 513

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 Amapa´ (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 2 = 10 triads on the body and 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). 514 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 Parana´ , Mato Grosso do Sul and Minas and 16575 from ‘‘Region 9, Dadanawa Gerais...’’ (Campbell & Lamar 2004:223). Ranch (N 0284903000 W, 5983103400 ), 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 (881200000 N, composed of 4 black bands. The first triad 05984700000 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 Bele´m, Vipera psyches and the holotype, from Para´ , Brazil, by Schmidt & Walker (1943) Surinam, is MNHN 7654 (Campbell & does not correspond to the known distri- Lamar 2004). VOLUME 125, NUMBER 4 515

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 Amapa´ ’’ (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 Type material.—The original name was 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 ha˜o northward including Rondoˆnia, Acre, appear to be bicolored. In some cases the Amazonas, Roraima, Para´ , and Amapa´ ; 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 basedontheAMNHandUSNMspec- 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,occursinVen- longest and stoutest of all the coral snakes ezuela and Trinidad (Campbell & Lamar (Campbell & Lamar 2004:231). 516 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 (682300000 N, 05884100000 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. Bele´m, Para´ , 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 Para´ , Pernambuco, and Rio Anguis Septemstriatus and the holotype, VOLUME 125, NUMBER 4 517 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 Sa˜oMarcos the breakup of Gondwanaland, based on (038050N, 608250W; 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 Vouchers for Guyana.—AMNH R-137979 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, 518 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 (881200000 N, 05984700000 W). Coloration in life.—The following is Typhlops minuisquamus based on the AMNH and USNM speci- 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 ‘‘spine’’ at the tip of the tail is dark brown

Loreto, Peru. 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). Vouchers for Guyana.—AMNH R- 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- VOLUME 125, NUMBER 4 519 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? Vic¸osa [Maroba´ ], 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 520 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON neotype. It is from Goia´ s, 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. 05804003.300 by long. Type material.—The original name was 59851044.800 , 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- VOLUME 125, NUMBER 4 521 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 Tome´ Assu´, Acara´ Mirim River, Para´ , 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 522 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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. 58330N, 558540W). Middorsally, blotches are separated by 1 Distribution.—‘‘Mexico...to Costa or2palescales.Thebodyblotches,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 (¼Guyana)’’ (Harris & Simmons 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 VOLUME 125, NUMBER 4 523 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 Territo´rio 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- Type material.—The original name was 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 524 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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. Coloration in life.—The following is 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 & Havasˇ (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, VOLUME 125, NUMBER 4 525

Guyanas, Surinam, northeastern Brazil)’’ and the Guianas and the Amazon drain- (Fritz & Havasˇ 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 (0482501200 N, 05885005800 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 extendingoneachsideofthemedial 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, 526 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 occasional straggler’’ (Pritchard 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 VOLUME 125, NUMBER 4 527 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 Type material.—The original name was 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 dorsally. The carapace and upper surfaces Waini Point; PCHP 2158–2162, 2501, 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 (0684400000 N, than the shell. The plastron, bridge, and 05785900000 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 528 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 & Havasˇ (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). Type material.—The original name was 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,betweenLidodiOstiaandTor (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 andJapansouthtoArgentina,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 (0881200000 N, found dead on the beach. PCHP 1021 from 05984700000 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 VOLUME 125, NUMBER 4 529 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 ‘‘Caribbean drainages of Guyana and Point; PCHP 3709 from NW District, 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 530 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 038080S, 598550W, 2775 and 2777 from Rupununi River, and the mouth of the Furo do Tajapuru, at Massala; PCHP 2776 from Rupununi 018020S, 51820W.’’ 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 Choco´ of Colombia, but its ni River, mouth of Rewa River; and PCHP mainrangeiseastoftheAndesineastern 5202–5203 from the Ireng River. Colombia, Venezuela, and the Guianas VOLUME 125, NUMBER 4 531 toeasternBrazil,southtoRiode 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 0 PCHP 2650–2651 from NW District, USNM 291194 from Wandapa (04832 Mabaruma; PCHP 2832–2833 from Bar- 0000 N, 05984300000 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 Coloration in life.—The following is (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.’’ 532 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 (Dume´ril, 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 & Sen˜aris 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: 68210N588410 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 VOLUME 125, NUMBER 4 533

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 (Dume´ril & 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).Ofthese,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,ofwhichcircinalis 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 534 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 & Bo¨hme 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 mate this, and we found a good model in Mesoclemmys nasuta recent works on the mammals of Paracou, VOLUME 125, NUMBER 4 535

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). 536 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 XX – – – – – Allobates spumaponens ––X –X–– Anomaloglossus kaiei ––X – ––– Atelopus spumarius –XX – – –– Dendrophryniscus minutus ––– –X–– Rhaebo guttatus –XX XXX– Rhaebo nasicus –X– – – –– Rhinella merianae –X– X – –X Rhinella marina XX – X XXX Rhinella martyi XXX X X – – Ceratophrys cornuta –X– – – –– Pristimantis zeuctotylus ––– –X–– Ameerega hahneli XX – – – – – Ameerega trivittata XX – – – – – Dendrobates leucomelas XX – – – – – Stefania evansi –XX – – –– Dendropsophus brevifrons ––X – ––– Dendropsophus marmoratus XXX – – – – Dendropsophus minusculus X– – X –XX Dendropsophus minutus ––XX––– Hypsiboas boans XXX X XX– Hypsiboas calcaratus ––X – –––

Hypsiboas cinerascens XXX – X – – Hypsiboas crepitans ––– X––X Hypsiboas geographicus XXX X X – – Hypsiboas multifasciatus XXX X – – – Lysapsus laevis ––– – –XX Osteocephalus buckleyi –X– – – –– Osteocephalus cabrerai ––X – ––– Osteocephalus leprieurii –XX – X–– Osteocephalus oophagus ––X –X–– Osteocephalus taurinus XXX X X – – Phyllomedusa bicolor XX – – – – – Phyllomedusa hypochondrialis –XXXX–– Phyllomedusa vaillantii X– – – X–– Pseudis paradoxa –X– – – –– Scinax boesemani ––– X––– Scinax nebulosus ––– X––– Scinax ruber XX – X –X– Scinax cf. x-signatus X– – X – –X Scinax sp. – – – – X – – Trachycephalus coriaceus ––– –X–– Trachycephalus typhonius –X– X – –– Adenomera andreae ––– –X–– Adenomera hylaedactyla XX – X XX– Leptodactylus fuscus –X– X –XX Leptodactylus guianensis –XX XXX– Leptodactylus knudseni XXX X X – – Leptodactylus leptodactyloides –X– – – –– Leptodactylus longirostris –X– – – –X Leptodactylus macrosternum ––– – –XX Leptodactylus mystaceus XXX X X – – VOLUME 125, NUMBER 4 537

Table 1.—Continued.

Taxon BaF KtF KoF DuM BeF KaS AiS

Leptodactylus petersii –XXXXX– Leptodactylus rhodomystax –XX – X–– Leptodactylus rugosus –X– – ––– Leptodactylus validus XX – X –X– Lithodytes lineatus XXX X X– – Physalaemus cuvieri X– – X ––– Pleurodema brachyops ––– – –XX Pseudopaludicola boliviana ––– – –XX Chiasmocleis jimi ––– –X–– Ctenophryne geayi XX – X X– – Elachistocleis surinamensis –––X––X Otophryne pyburni –X– – ––– Synapturanus salseri –XX – ––– Pipa arrabali XX – – – – – Pipa pipa –X– – X–– Lithobates palmipes –XX – ––X Caecilia tentaculata –––X––– Rhinatrema shiv ––X– ––– Microcaecilia sp. – – X – X – – Caiman crocodilus –X– XX–– Melanosuchus niger ––– – –X– Paleosuchus trigonatus –XX – ––– Amphisbaena alba –X– – ––– Amphisbaena fuliginosa –X– X ––– Hemidactylus mabouia –X– X –––

Hemidactylus palaichthus –––X–XX Thecadactylus rapicauda –XXXX–– Alopoglossus angulatus ––– –X–– Arthrosaura reticulata ––– –X–– Bachia flavescens –XXXX–– Cercosaura ocellata –X– X ––X Echinosaura sulcarostrum X– – – ––– Gymnophthalmus leucomystax ––– – ––X Gymnophthalmus speciosus –––X––– Gymnophthalmus underwoodi –––XXXX Gymnophthalmus vanzoi ––– – –XX Iphisa elegans –X– – X–– Leposoma percarinatum XXX X X– – Neusticurus bicarinatus –X– – X–– Neusticurus rudis –XX – ––– Iguana iguana –X– X –XX Anolis aeneus –X– – ––– Anolis auratus –––X–XX Anolis fuscoauratus XXX – – – – Anolis ortonii –X– – ––– Anolis planiceps XXX – X– – Anolis punctatus –X– – X–– Polychrus marmoratus –X– X –XX Mabuya nigropunctata XXX – X–X Coleodactylus septentrionalis ––– – –X– Gonatodes alexandermendesi –XX – ––– Gonatodes annularis XX – – X– – Gonatodes humeralis –XXXX–X Pseudogonatodes guianensis ––XX––– Sphaerodactylus molei –X– – ––– 538 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Continued.

Taxon BaF KtF KoF DuM BeF KaS AiS

Ameiva ameiva XXX X XXX Cnemidophorus lemniscatus –X– X –XX Kentropyx calcarata XXX X XX– Kentropyx striata –––X–XX Tupinambis teguixin –XX X ––– Plica plica XXX – – – – Plica umbra XXX X X – – Tropidurus hispidus –––X–XX Uranoscodon superciliosus –XX XXXX Anilius scytale –XX – X–– Boa constrictor XX – X X – – Corallus caninus –XX X ––– Corallus hortulanus XX – X X – – Epicrates cenchria –XX X ––– Eunectes murinus –XX X ––– Atractus torquatus –––X––– Atractus trilineatus –X– X ––– Chironius carinatus ––– – –X– Chironius exoletus –XX – ––– Chironius fuscus XX – – X – – Chironius multiventris –X– – X–– Chironius scurrulus –XX – X–– Clelia clelia –X– – ––– Dipsas catesbyi –X– – X–– Dipsas indica –X– – –––

Dipsas pavonina –X– – ––– Dipsas variegata XX – – – – – Drymoluber dichrous –X– – X–– Erythrolamprus aesculapii –X– X ––– Helicops angulatus –X– – X–– Hydrops triangularis –X– – ––– Imantodes cenchoa –XX – X–– Imantodes lentiferus X– – – ––– Leptodeira annulata –X– X –X– Leptophis ahaetulla XXX X –X– Liophis breviceps –XX X ––– Liophis cobella –X– – ––– Liophis lineatus –X– – ––– Liophis miliaris X– – – ––– Liophis poecilogyrus ––– – –X– Liophis reginae –X– – ––– Liophis typhlus XX – X – – – Mastigodryas boddaerti XX – X XX– Oxybelis aeneus –X– – ––– Oxybelis fulgidus –XX – X–– Oxyrhopus melanogenys –X– XX–– Oxyrhopus petolarius XX – X – – – Philodryas olfersii –––X––X Philodryas viridissimus –X– – ––– Phimophis guianensis –––X––– Pseudoboa coronata ––– –X–– Pseudoboa neuwiedii –––X––X Pseustes poecilonotus –XX – ––– Pseustes sulphureus –X– – ––– Rhinobothryum lentiginosum X– – – ––– VOLUME 125, NUMBER 4 539

Table 1.—Continued.

Taxon BaF KtF KoF DuM BeF KaS AiS

Siphlophis compressus XXX X – –– Tantilla melanocephala XX – XX–X Thamnodynastes ramonriveroi –––X––– Xenodon rabdocephalus –X– – ––– Xenodon severus XX – – – –– Micrurus hemprichii X– – – ––– Micrurus lemniscatus XX – – –X– Micrurus psyches –X– – ––– Micrurus surinamensis –––XX–– Epictia albifrons XX – XX–– Siagonodon septemstriatus –X– – ––– Tricheilostoma dimidiatum ––– ––X– Typhlops brongersmianus –––X––– Typhlops minuisquamus –X– X––– Typhlops reticulatus –X– X––– Bothriopsis bilineata –X– – ––– Bothrops atrox XX – XX–– 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–XX

Chelonoidis denticulata XXX – 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=N þ N C 3 100; How similar (or dissimilar) are the 1 2 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). 540 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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=N 3 100; 1 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 VOLUME 125, NUMBER 4 541

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 Nin˜o 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 an index of 68–72% compared with the Ba F Kt F Ko F Du M Be F Ka S Ai S 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]-R´ıo 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-R´ıo 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-R´ıoBran- 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-R´ıo Branco Depression in both comparisons, one site is in savanna also served as a route for lowland Amazo- 542 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 rainyseasonconnectionbetweentheR´ıo 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 Sen˜aris & 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. VOLUME 125, NUMBER 4 543

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 Xremarkably 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 –– Xduced 23–26 species, the highest number of Vitreorana gorzulae –– Xspecies 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 XMounts Ayanganna and Wokomung, Pristimantis marmoratus XX Xwhich are 37 km apart, have the greatest Pristimantis pulvinatus –X – Pristimantis saltissimus –– Xfaunal resemblance (61%), while signifi- Adelophryne patamona –– Xcantly fewer species are shared between Stefania ackawaio –X Xthese two sites and the more distant Stefania ayangannae –X XRoraima (both less than 40%). The num- Stefania coxi –X X Stefania roraimae XX Xber of species found at only one of the Stefania scalae XX – highland sites is especially striking, indi- ‘‘Hyla’’ warreni XX – cating a strong distinctiveness of each site. Hypsiboas roraima XX X Hypsiboas sibleszi XX X Table 4.—Comparisons of number of species of Myersiohyla kanaima XX 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 XX 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/N 3 100, comparing sites where rows Neusticurus rudis XX – 1 and columns meet. Riolama leucosticta X– – Anolis planiceps X– – Roraima Ayanganna Wokomung Mabuya nigropunctata X– – Tropidurus hispidus X– –Roraima 23 96 Atractus steyermarki X– –Ayanganna 39 26 14 Chironius challenger –– XWokomung 26 61 23 544 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 VOLUME 125, NUMBER 4 545

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- fauna of Guyana includes 324 species Roraima 2 5 4 4 2 3 3 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%)and25 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 546 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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. þ¼yes; – 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 VOLUME 125, NUMBER 4 547

Table 6.—Continued.

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 548 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 tus, listed from the least to greatest conservation, with the exception of species listed as 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 ened species are included in the categories Anomaloglossus roraima DD Ukn 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 impacted hardest by the lethal amphibian Osteocephalus phasmatus DD Ukn 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 rest of the world, threat levels are com- Microcaecilia rabei DD Ukn paratively low in the Guiana Shield (Bolan˜os 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, TheIUCNhasyettocompletea 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% VOLUME 125, NUMBER 4 549

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. 550 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 VOLUME 125, NUMBER 4 551 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 (Andre´s 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 Sen˜aris (MHNLS); Jonathan Losos and for all this effort also. Line drawings Jose´ 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. Wu¨ster 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 552 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 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 Para´ , Brazil: a forgotten part of the University of Guyana. Guianan region, 1. Herpetofauna. Boletim do This report is number 179 in the Museu Paraense Em´ılio Goeldi Cieˆncias Naturais, Bele´m 5(1):13–112. Smithsonian’s BDG publication series Ayarzagu¨ena, J. 1992. Los Centrole´nidos de la and contribution number 346 of the Guayana venezolana. Publicaciones de la ROM Centre for Biodiversity and Conser- Asociacio´n de Amigos de Don˜ana 1:1–48. vation Biology. We are especially grateful Bailey, J. R., & R. A. Thomas. 2007 [2006]. A to the Biological Society of Washington, revision of the South American snake genus 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- important financial contributions toward botensis Pe´rez-Santos and Moreno and Tham- nodynastes ramonriveroi Manzanilla and producing this volume. Any use of trade, Sa´ nchez. Memoria de la Fundacio´n 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. ment by the U.S. Government. A revision of the South American snake genus Thamnodynastes Wagler, 1830 (Serpentes, Colubridae, Tachymenini). I. Two new species Literature Cited of Thamnodynastes from central Brazil and adjacent areas, with a redefinition of and Adalsteinsson, S. A., W. R. Branch, S. Trape, L. J. neotype designation for Thamnodynastes pal- Vitt, & S. B. Hedges. 2009. Molecular lidus (Linnaeus, 1758). Phyllomedusa 4(2):83–

phylogeny, classification, and biogeography 101. of snakes of the family Leptotyphlopidae Barbour, T. 1914. A contribution to the zoo¨geog- (Reptilia, Squamata). Zootaxa 2244:1–50. raphy of the West Indies, with especial Albuquerque, N. R. de, & T. de Lema. 2008. reference to amphibians and reptiles. Memoirs Taxonomic revision of the neotropical water of the Museum of Comparative Zoo¨logy at snake Hydrops triangularis (Serpentes, Colu- Harvard College 44(2):209–359. bridae). Zootaxa 1685:55–66. Barbour, T. 1920. Comments on a recent check-list. Anderson, J. S., R. R. Reisz, D. Scott, N. B. The American Naturalist 54(632):284–288. Fro¨bisch, & S. S. Sumida. 2008. A stem Barrio-Amoro´s, C. L. 1998. Sistema´ tica y bio- batrachian from the Early Permian of Texas geograf´ıa de los anfibios (Amphibia) de and the origin of frogs and salamanders. Venezuela. Acta Biologica Venezuelica 18(2):1–93. Nature 453:515–518. Bauer, A. M., & R. Gu¨nther. 2006. An annotated Andersson, L. G. 1918. New lizards from South catalogue of the types of crocodilians (Repti- America. Collected by Nils Holmgren and A. lia: Crocodylia) in the collection of the Roman. Arkiv fo¨r Zoologi 11(16):1–9. Museum fu¨r Naturkunde der Humboldt-Uni- Andersson, L. G. 1945. Batrachians from east versita¨t zu Berlin (ZMB). Zoosystematics and Ecuador collected 1937, 1938 by Wm. Evolution 82(2):243–247. Clarke-Macintyre and Rolf Blomberg. Arkiv Bauer, A. M., R. Gu¨nther, & M. Klipfel. 1995. The fo¨r Zoologi, Stockholm 37A(2):1–88. herpetological contributions of Wilhelm C. H. Angulo, A., R. B. Cocroft, & S. Reichle. 2003. Peters (1815–1883); with an introduction, Species identity in the genus Adenomera annotated bibliography, and synopsis of taxa. (Anura: Leptodactylidae) in southeastern Pe- Society for the Study of Amphibians and ru. Herpetologica 59(4):490–504. Reptiles, in cooperation with Deutsche Ge- Avila-Pires, T. C. S. 1995. Lizards of Brazilian sellschaft fu¨r Herpetologie und Terrarien- Amazonia (Reptilia: Squamata). Zoo¨logische kunde, Ithaca, New York, 714 pp. Verhandelingen, Leiden 299:1–706. Bauer, A. M., V. Wallach, & R. Gu¨nther. 2002. An A´ vila-Pires, T. C. S. 2005. Reptiles. Pp. 25–40 in T. annotated type catalogue of the scolecophi- Hollowell, & R. P. Reynolds, eds., Checklist dian, alethinophidian, and macrostomatan of the Terrestrial Vertebrates of the Guiana snakes in the collection of the Museum fu¨r VOLUME 125, NUMBER 4 553

Naturkunde der Humboldt-Universita¨tzu Berridge, P. Ramani, & B. E. Young, eds., Berlin. Zoosystematics and Evolution Threatened amphibians of the world. Lynx 78:157–176. Edicions, Barcelona, Spain; IUCN—The Beebe, W. 1919. Higher vertebrates of British World Conservation Union, Gland, Switzer- Guiana, with special reference to the fauna land; Conservation International, Arlington, of Bartica District. Zoologica, Scientific Con- Virginia, and NatureServe, 758 pp. tributions of the New York Zoological Bonnaterre, P.-J. 1789. Tableau encyclope´dique et Society 2(7):205–238. me´thodique des trois re`gnes de la nature, Beebe, W. 1944. Field notes on the lizards of de´die´ et pre´sente´ a M. Necker, ministre d’E´ tat, Kartabo, British Guiana, and Caripito, Ven- & directeur ge´ne´ral des Finances: Erpe´tologie. ezuela. Part 1. Gekkonidae. Zoologica, Scien- Chez Panckoucke, Paris, 70 pp. tific Contributions of the New York Boulenger, G. A. 1882. Catalogue of the Batrachia Zoological Society 29:145–160. Salientia s. Ecaudata in the collection of the Beebe, W. 1945. Field notes on the lizards of British Museum. Second Edition. Taylor and Kartabo, British Guiana, and Caripito, Ven- Francis, London, 503 pp. þ 30 pls. ezuela. Part 3. Teiidae, Amphisbaenidae and Boulenger, G. A. 1884 [1883]. On a collection of Scincidae. Zoologica, Scientific Contributions frogs from Yurimaguas, Huallaga River, of the New York Zoological Society 30:7–32. northern Peru. Proceedings of the Zoological Beebe, W. 1946. Field notes on the snakes of Society of London 1883:635–638. Kartabo, British Guiana, and Caripito, Ven- Boulenger, G. A. 1887. On a new geckoid lizard from ezuela. Zoologica, Scientific Contributions of British Guiana. Proceedings of the Zoological the New York Zoological Society 31:11–51. Society of London 1887:153–154. Beireis, G. C. 1783. Beschreibung eines bisher Boulenger, G. A. 1895. Description of a new unbekannt gewesenen amerikanischen batrachian (Oreophryne Quelchii) discovered Frosches. Schriften der Berlinischen Gesell- by Messrs. J. J. Quelch and F. McConnell on schaft naturforschender Freunde 4:178–182. the summit of Mount Roraima. Annals and Berry, J. F., & J. B. Iverson. 2001. Kinosternon Magazine of Natural History, Sixth Series, scorpioides. Catalogue of American Amphib- 15(90):521–522. ians and Reptiles 725:1–11. Boulenger, G. A. 1900a. Batrachians. Pp. 55–56 in

Boddaert, P. 1772. Epistola ad virum celeberrimum Report on a collection made by Messrs. F. V. Johannem Oosterdyk Schacht ... de rana McConnell and J. J. Quelch at Mount bicolore, descripta atque accuratissima icone Roraima in British Guiana. (Communicated illustrata ex museo ... Johannis Alberti by Prof. E. Ray Lankester, D.C.L., F.R.S., Schlosseri ... ¼ Brief van Pieter Boddaert aan Director of the Natural History Museum.) den zeer weledelen hooggeleerden heere Jo- Transactions of the Linnean Society of hannes Oosterdyk Schacht ... behelzende eene London, Second series, Zoology 8(2):51–76. naauwkeurige beschrijving, en naar het leven Boulenger, G. A. 1900b. Reptiles. Pp. 53–54 in vervaardigde afbeelding, van den twee-ko- Report on a collection made by Messrs. F. V. leurigen kikvorsch, uit de verzameling van ... McConnell and J. J. Quelch at Mount Johannes Albertus Schlosser. M. Mage´rum, Roraima in British Guiana. (Communicated Amsterdam, pp. 1–48. by Prof. E. Ray Lankester, D.C.L., F.R.S., Boettger, O. 1894. A preliminary list of the reptiles Director of the Natural History Museum.) and batrachians of the island of Trinidad. Transactions of the Linnean Society of With descriptions of two new species by London, Second series, Zoology 8(2):51–76. Professor Dr. O. Boettger, p. 80 in R. R. Boulenger, G. A. 1904. Description of a new tree- Mole & F. W. Urich. Trinidad Field Natu- frog of the genus Hyla, from British Guiana, ralists’ Club 2:77–90. carrying eggs on the back. Proceedings of the Boie, F. 1827. Bemerkungen u¨ber Merrem’s Versuch Zoological Society of London 2:106. eines Systems der Amphibien. Isis von Oken Bour, R., & A. Dubois. 1983. Nomenclatural 20(6):508–566. availability of Testudo coriacea Vandelli, Bolan˜os, F., F. Castro, C. Cortez, I. De la Riva, T. 1761: a case against a rigid application of Grant, B. Hedges, R. Heyer, R. Iba´ n˜ez, E. La the rules to old, well-known zoological works. Marca, E. Lavilla, D. L. Silvano, S. Lo¨tters, Journal of Herpetology 17:356–361. G. P. Olea, S. Reichle, R. Reynolds, L. Bourne, G. R. 1997. Reproductive behavior of Rodriguez, G. Santos Barrera, N. Scott, C. terrestrial breeding frogs Eleutherodactylus Ubeda, A. Veloso, M. Wilkinson, & B. johnstonei in Guyana. Journal of Herpetology Young. 2008. Amphibians of the neotropical 31(2):221–229. realm. Pp. 92–105 in S. N. Stuart, M. Bourne,G.R.2001.Colorpattern,predator Hoffmann, J. S. Chanson, N. A. Cox, R. J. avoidance, and foraging behavior in the 554 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

golden frog Colostethus beebei (Anura: Den- Cannatella, D. C., & W. E. Duellman. 1984. drobatidae). Herpetological Review Leptodactylid frogs of the Physalaemus pus- 32(4):225–228. tulosus group. Copeia 1984(4):902–921. Bourne, G. R., A. C. Collins, A. M. Holder, & C. L. Caramaschi, U. 2010. Notes on the taxonomic status McCarthy. 2001. Vocal communication and of Elachistocleis ovalis (Schneider, 1799) and reproductive behavior of the frog Colostethus description of five new species of Elachistocleis beebei in Guyana. Journal of Herpetology Parker, 1927 (Amphibia, Anura, Microhyli- 35:272–281. dae). Boletim do Museu Nacional, Nova Se´rie Brazaitis, P. 1973. The identification of living Zoologia 527:1–30. crocodilians. Zoologica, Scientific Contribu- Caramaschi, U., & C. A. G. Cruz. 2001. A new tions of the New York Zoological Society species of Chiasmocleis Me´hely¨ , 1904 from 58:59–101. Brazilian Amazonia (Amphibia, Anura, Mi- Brochu, C. A., J. R. Wagner, S. Jouve, C. D. crohylidae). Boletim do Museu Nacional, Sumrall, & L. D. Densmore. 2009. A correc- Nova Se´rie Zoologia 469:1–8. tion corrected: consensus over the meaning of Carvalho, C. M. de. 1997. Uma nova espe´cie de Crocodylia and why it matters. Systematic microteiideo do geˆnero Gymnophthalmus do Biology 58(5):537–543. estado de Roraima, Brasil (Sauria, Gymnoph- Brongersma, L. D. 1933. Ein neuer Laubfrosch aus thalmidae). Pape´is Avulsos de Zoologia, Sa˜o Surinam. Zoologischer Anzeiger 103(9/ Paulo 40(10):161–174. 10):267–270. Carvalho, V. T., R. D. MacCulloch, L. Bonora, & R. Brongersma, L. D. 1966. Note on Leptomicrurus C. Vogt. 2010. New species of Stefania collaris (Schlegel) (Reptilia, Serpentes). Zoo- (Anura: Cryptobatrachidae) from northern logische Mededelingen 41(17):243–254. Amazonas, Brazil. Journal of Herpetology Brown, J. L., E. Twomey, A. Ame´zquita, M. B. de 44(2):229–235. Souza, J.-L. P. Caldwell, S. Lo¨tters, R. von Castoe, T. A., T. M. Doan, & C. L. Parkinson. 2004. May, P. R. Melo-Sampaio, D. Mej´ıa-Vargas, Data partitions and complex models in P. Perez-Pen˜a, M. Pepper, E. H. Poelman, M. Bayesian analysis: the phylogeny of gymnoph- Sanchez-Rodriguez, & K. Summers. 2011. A thalmid lizards. Systematic Biology 53:448– taxonomic revision of the Neotropical poison 469.

frog genus Ranitomeya (Amphibia: Dendro- Castroviejo-Fisher, S., J. M. Guayasamin, & P. J. R. batidae). Zootaxa 3083:1–120. Kok. 2009. Species status of Centrolene lema Brust, D. G. 1993. Maternal brood care by Duellman and Sen˜aris, 2003 (Amphibia: Dendrobates pumilio: a frog that feeds its Centrolenidae) revealed by integrative taxon- young. Journal of Herpetology 27:96–98. omy. Zootaxa 1980:16–28. Bugoni, L., & P. Welff-Neto. 2008. Hemidactylus Castroviejo-Fisher, S., C. Vila`, J. Ayarzagu¨ena, M. mabouia. Human-induced introduction. Her- Blanc, & R. Ernst. 2011. Species diversity of petological Review 39(2):226–227. Hyalinobatrachium glassfrogs (Amphibia: Cadle, J. E., & C. W. Myers. 2003. Systematics of Centrolenidae) from the Guiana Shield, with snakes referred to Dipsas variegata in Panama the description of two new species. Zootaxa and western South America, with revalidation 3132:1–55. of two species and notes on defensive behav- Chanson, J., S. Stuart, N. Cox, B. Young, & M. iors in the Dipsadini (Colubridae). American Hoffman. 2008a. The global amphibian as- Museum Novitates 3409:1–47. sessment (GAA): history, objectives and Camargo, A., R. O. de Sa´ , & W. R. Heyer. 2006. methodology. Pp. 30–32 in S. N. Stuart, M. Phylogenetic analyses of mtDNA sequences Hoffmann, J. S. Chanson, N. A. Cox, R. J. reveal three cryptic lineages in the widespread Berridge, P. Ramani, & B. E. Young, eds., neotropical frog Leptodactylus fuscus Threatened amphibians of the world. Lynx (Schneider, 1799) (Anura, Leptodactylidae). Edicions, Barcelona, Spain; IUCN—The Biological Journal of the Linnean Society World Conservation Union, Gland, Switzer- 87:325–341. land; Conservation International, Arlington, Campbell, J. A., & B. T. Clarke. 1998. A review of Virginia, and NatureServe, 758 pp. frogs of the genus Otophryne (Microhylidae) Chanson, J., M. Hoffman, N. Cox, & S. Stuart. with the description of a new species. Herpe- 2008b. The state of the world’s amphibians. tologica 54:301–317. Pp. 33–52 in S. N. Stuart, M. Hoffmann, J. S. Campbell, J. A., & W. W. Lamar. 2004. The Chanson, N. A. Cox, R. J. Berridge, P. venomous reptiles of the Western Hemisphere Ramani, & B. E. Young, eds., Threatened (Two volume set). Comstock Publishing amphibians of the world. Lynx Edicions, Associates, Cornell University Press, Ithaca, Barcelona, Spain; IUCN—The World Con- New York, 976 pp. servation Union, Gland, Switzerland; Conser- VOLUME 125, NUMBER 4 555

vation International, Arlington, Virginia, and Cope, E. D. 1860b. Supplement to ‘‘A catalogue of NatureServe, 758 pp. the venomous serpents in the Museum of the Chao, A. 1984. Nonparametric estimation of the Academy,’’ etc. Proceedings of the Academy number of classes in a population. Scandina- of Natural Sciences of Philadelphia 12:72–74. vian Journal of Statistics 11:265–270. Cope, E. D. 1864. Contributions to the herpetology Chaparro, J. C., J. B. Pramuk, & A. G. Gluesen- of tropical America. Proceedings of the kamp. 2007. A new species of arboreal Academy of Natural Sciences, Philadelphia Rhinella (Anura; Bufonidae) from cloud forest 16:166–181. of southeastern Peru. Herpetologica 63:203– Cope, E. D. 1865. Sketch of primary groups of 212. Batrachia salientia. The Natural History Chippaux, J.-P. 1986. Les Serpents de la Guyane Review, New Series 5:97–120. franc¸aise. Collection Faune Tropicale 27, Cope, E. D. 1868. An examination of the Reptilia E´ ditions de l’ORSTOM, Institut Franc¸ais de and Batrachia obtained by the Orton Expedi- Recherche Scientifique pour le De´velop- tion to Ecuador and the Upper Amazon, with pement en Coope´ration, Paris, 165 pp. notes on other species. Proceedings of the Cochran, D. M., & C. J. Goin. 1970. Frogs of Academy of Natural Sciences, Philadelphia Colombia. United States National Museum 20:96–140. Bulletin 288:1–655. Cope, E. D. 1869 [1868]. Sixth contribution to the Cole, C. J., & H. C. Dessauer. 1993. Unisexual and herpetology of tropical America. Proceedings bisexual whiptail lizards of the Cnemidophorus of the Academy of Natural Sciences, Phila- lemniscatus complex (Squamata: Teiidae) of delphia 20:305–313. the Guiana Region, South America, with Cope, E. D. 1871. Ninth contribution to the descriptions of new species. American Muse- herpetology of tropical America. Proceedings um Novitates 3081:1–30. of the Academy of Natural Sciences, Phila- Cole, C. J., & P. J. R. Kok. 2006. A new species of delphia 23:200–224. gekkonid lizard (Sphaerodactylinae: Gonato- Cope, E. D. 1879. Eleventh contribution to the des) from Guyana, South America. American herpetology of tropical America. Proceedings Museum Novitates 3524:1–13. of the American Philosophical Society 18:261–

Cole, C. J., H. C. Dessauer, & A. L. Markezich. 277. 1993. Missing link found: the second ancestor Cope, E. D. 1894. On the species of Himantodes D. & of Gymnophthalmus underwoodi (Squamata: B. The American Naturalist 28(331):612–614. Teiidae), a South American unisexual lizard of Curcio,F.F.,V.deQ.Piacentini,&D.S. hybrid origin. American Museum Novitates Fernandes. 2009. On the status of the snake 3055:1–13. genera Erythrolamprus Boie, Liophis Wagler Cole, C. J., H. C. Dessauer, C. R. Townsend, & M. and Lygophis Fitzinger (Serpentes, Xenodon- G. Arnold. 1990. Unisexual lizards of the tinae). Zootaxa 2173:66–68. genus Gymnophthalmus (Reptilia: Teiidae) in Cuvier, G. 1797. Tableau ele´me´ntaire de l’histoire the neotropics: genetics, origin, and systemat- naturelle des animaux. Baudouin, Paris, 710 ics. American Museum Novitates 2994:1–29. pp. Cole, C. J., H. C. Dessauer, C. R. Townsend, & M. Cuvier, G. 1807. Sur les diffe´rentes espe`ces de G. Arnold. 1995. Kentropyx borckiana (Squa- crocodiles vivans et sur leurs caracte`res dis- mata: Teiidae): a unisexual lizard of hybrid tinctifs. Annales du Muse´um d’Histoire Na- origin in the Guiana Region, South America. turelle, Paris 10:8–66. American Museum Novitates 3145:1–23. Cuvier, G. 1817. Le re`gne animal distribue´ d’apre`s Condit, J. M. 1964. A list of the types of hylid frogs son organisation, pour servir de base a` in the collection of the British Museum l’histoire naturelle des animaux et d’introduc- (Natural History). Journal of the Ohio Her- tion a` l’anatomie compare´e. Tome II, les petological Society 4:85–98. reptiles, les poissons, les mollusques et les Conrad, J. L. 2008. Phylogeny and systematics of anne´lides. Chez De´terville, Paris, 532 pp. Squamata (Reptilia) based on morphology. D’Angiolella, A. B., T. Gamble, T. C. S. Avila-Pires, Bulletin of the American Museum of Natural G. R. Colli, B. P. Noonan, & L. J. Vitt. 2011. History 310:1–182. Anolis chrysolepis Dume´ril and Bibron, 1837 Cope, E. D. 1860a. Catalogue of the Colubridae in (Squamata: Iguanidae), revisited: molecular the Museum of the Academy of Natural phylogeny and taxonomy of the Anolis chrys- Sciences of Philadelphia, with notes and olepis species group. Bulletin of the Museum descriptions of new species. Part 2. Proceed- of Comparative Zoology 160(2):35–63. ings of the Academy of Natural Sciences of Daly, J. W., T. F. Spande, & H. M. Garraffo. 2005. Philadelphia 12:241–266. Alkaloids from amphibian skin: a tabulation 556 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

of over eight-hundred compounds. Journal of Daudin, F. M. 1803b. Histoire naturelle, ge´ne´rale et Natural Products 68:1556–1575. particulie`re des reptiles; ouvrage faisant suite a` Daly, J. W., F. Gusovsky, C. W. Myers, M. Yotsu- l’Histoire naturelle ge´ne´rale et particulie`re, Yamashita, & T. Yasumoto. 1994. First compose´e par Leclerc de Buffon, et re´dige´e occurrence of tetrodotoxin in a dendrobatid par C. S. Sonnini, membre de plusieurs frog (Colostethus inguinalis), with further Socie´te´s savantes. Tome 8. F. Dufart, Paris, reports for the bufonid genus Atelopus. 439 pp. Toxicon 32:279–285. De Freitas, D. 2009. Eunectes murinus diet. Herpe- Daly, J. W., H. M. Garraffo, P. Jain, T. F. Spande, tological Review 40:98. R. R. Snelling, C. Jaramillo, & A. S. Rand. Dixon, J. R. 1981. The neotropical colubrid snake 2000. Arthropod-frog connection: decahydro- genus Liophis: the eastern Caribbean complex. quinoline and pyrrolizidine alkaloids common Copeia 1981(2):296–304. to microsympatric myrmicine ants and den- Dixon, J. R. 1983a. The Liophis cobella group of the drobatid frogs. Journal of Chemical Ecology neotropical colubrid snake genus Liophis. 26:73–85. Journal of Herpetology 17:149–165. Daly, J. W., T. Kaneko, J. Wilham, H. M. Garraffo, Dixon, J. R. 1983b. Taxonomic status of the South T. F. Spande, A. Espinosa, & M. A. Donnelly. American snakes Liophis miliaris, L amazoni- 2002. Bioactive alkaloids of frog skin: combi- cus, L. chrysostomus, L. mossoroensis and L. natorial bioprospecting reveals that pumilio- purpurans (Colubridae: Serpentes). Copeia toxins have an arthropod source. Proceedings 1983(3):791–802. of the National Academy of Sciences Dixon, J. R. 1983c. Systematics of Liophis reginae 99(22):13996–14001. and L. williamsi (Serpentes, Colubridae), with Daudin, F. M. 1800. Histoire naturelle des quad- a description of a new species. Annals of the rupe`des ovipares; avec des gravures faites Carnegie Museum of Natural History 52:113– enlumine´e sur les dessins d’apre`s nature par 138. J. Barraband. Fuchs et Delalain fils, Paris, 24 Dixon, J. R. 1987. Taxonomy and geographic pp. þ 12 pls. variation of Liophis typhlus and related Daudin, F. M. 1802a. Histoire naturelle des rain- ‘‘green’’ species of South America (Serpentes: ettes, des grenouilles et des crapauds; de´die´e a` Colubridae). Annals of Carnegie Museum

B. G. E. L. Lace´pe`de. Bertrandet, Paris, 108 56:173–191. pp. þ 38 pls. Dixon, J. R. 1989. A key and checklist to the Daudin, F. M. 1802b. Histoire naturelle, ge´ne´rale et neotropical snake genus Liophis with country particulie`re des reptiles; ouvrage faisant suite a` lists and maps. Smithsonian Herpetological l’Histoire naturelle ge´ne´rale et particulie`re, Information Service 79:1–28. compose´e par Leclerc de Buffon, et re´dige´e Dixon, J. R., & F. S. Hendricks. 1979. The worm- par C. S. Sonnini, membre de plusieurs snakes (Family Typhlopidae) of the neotrop- Socie´te´s savantes. Tome 4. F. Dufart, Paris, ics, exclusive of the Antilles. Zoologische 397 pp. Verhandelingen 173:1–39. Daudin, F. M. 1802c. Histoire naturelle, ge´ne´rale et Dixon, J. R., & A. L. Markezich. 1992. Taxonomy particulie`re des reptiles; ouvrage faisant suite a` and geographic variation of Liophis poecilo- l’Histoire naturelle ge´ne´rale et particulie`re, gyrus (Wied) from South America (Serpentes: compose´e par Leclerc de Buffon, et re´dige´e Colubridae). Texas Journal of Science 44:131– par C. S. Sonnini, membre de plusieurs 166. Socie´te´s savantes. Tome 3. F. Dufart, Paris, Dixon, J. R., J. A. Wiest, Jr., & J. M. Cei. 1993. 452 pp. Revision of the neotropical snake genus Daudin, F. M. 1802d. Histoire naturelle, ge´ne´rale et Chironius Fitzinger: (Serpentes, Colubridae). particulie`re des reptiles; ouvrage faisant suite a` Monografie di Museo Regionale di Scienze l’histoire naturelle ge´ne´rale et particulie`re, Naturali, Torino 13:1–280. compose´e par Leclerc de Buffon, et re´dige´e Doan, T. M. 2003. A new phylogenetic classification par C. S. Sonnini, membre de plusieurs for the gymnophthalmid genera Cercosaura, Socie´te´s savantes. Tome 2. F. Dufart, Paris, Pantodactylus and Prionodactylus (Reptilia: 432 pp. Squamata). Zoological Journal of the Linnean Daudin, F. M. 1803a. Histoire naturelle, ge´ne´rale et Society 137:101–115. particulie`re des reptiles; ouvrage faisant suite a` Donnelly, M. A., M. H. Chen, & G. G. Watkins. l’Histoire naturelle ge´ne´rale et particulie`re, 2005. The Iwokrama herpetofauna: an explo- compose´e par Leclerc de Buffon, et re´dige´e ration of diversity in a Guyanan rainforest. par C. S. Sonnini, membre de plusieurs Pp. 428–460 in M. A. Donnelly, B. I. Crother, Socie´te´s savantes. Tome 6. F. Dufart, Paris, C. Guyer, M. H. Wake, & M. E. White, eds., 447 pp. Ecology and evolution in the tropics: a VOLUME 125, NUMBER 4 557

herpetological perspective. The University of Duellman, W. E., & M. Yoshpa. 1996. A new species Chicago Press, Chicago, 675 pp. of Tepuihyla (Anura: Hylidae) from Guyana. Donnelly, M. A., R. D. MacCulloch, C. A. Ugarte, Herptetologica 52:275–281. & D. Kizirian. 2006. A new riparian gym- Dume´ril, A. M. C., & G. Bibron. 1837. Erpe´tologie nophthalmid (Squamata) from Guyana. Co- ge´ne´rale, ou histoire naturelle comple`te des peia 2006(3):396–403. reptiles. Tome 4. Librairie Encyclope´dique de Dubois, A. 2007. Genitives of species and subspecies Roret, Paris, 571 pp. nomina derived from personal names should Dume´ril, A. M. C., & G. Bibron. 1841. Erpe´tologie not be emended. Zootaxa 1550:49–68. ge´ne´rale ou histoire naturelle comple`te des Duellman, W. E. 1958. A monographic study of the reptiles. Tome 8. Librairie Encyclope´dique de colubrid snake genus Leptodeira. Bulletin of Roret, Paris, 792 pp. the American Museum of Natural History Dume´ril, M. C., & M. A. Dume´ril. 1851. Catalogue 114:1–152. me´thodique de la collection des reptiles du Duellman, W. E. 1965. A biogeographic account of Muse´um d’Histoire Naturelle de Paris. Gide the herpetofauna of Michoaca´ n, Me´xico. et Baudry, Paris, 222 pp. University of Kansas Publications, Museum Dume´ril, A. M. C., G. Bibron, & A. Dume´ril. 1854. of Natural History 15(14):627–709. Erpe´tologie ge´ne´rale ou histoire naturelle Duellman, W. E. 1972. A new species of Hyla from comple`te des reptiles. Tome 7, Parties 1 & 2. Amazonian Ecuador. Copeia 1972(2):265– Librairie Encyclope´dique de Roret, Paris; 271. partie 1, pp. 1–780; partie 2, pp. 781–1536. Duellman, W. E. 1974. Taxonomic notes on Phyllo- Dunn, E. R. 1942. The American caecilians. Bulletin medusa (Anura: Hylidae) from the Upper of the Museum of Comparative Zoo¨logy at Amazon Basin. Herpetologica 30:105–112. Harvard College 91(6):439–540. Duellman, W. E. 1978. The biology of an equatorial Dunn, E. R. 1949. Notes on South American frogs of herpetofauna in Amazonian Ecuador. Uni- the family Microhylidae. American Museum versity of Kansas Museum of Natural History Novitates 1419:1–21. Miscellaneous Publication 65:1–352. Ernst, C. H. 1981. Phrynops gibbus. Catalogue of Duellman, W. E. (ed.). 1979. The South American American Amphibians and Reptiles 279:1–2. herpetofauna: its origin, evolution, and dis- Ernst, C. H. 1983. Geographic variation in the

persal. Monograph of the Museum of Natural neotropical turtle, Platemys platycephala. History, The University of Kansas, Lawrence Journal of Herpetology 17(4):345–355. 7:1–485. Ernst, C. H., & R. W. Barbour. 1989. Turtles of the Duellman, W. E. 1997. Amphibians of La Escalera World. Smithsonian Institution Press, Wash- region, southeastern Venezuela: taxonomy, ington, D.C., 313 pp. ecology, and biogeography. Scientific Papers, Ernst, C. H., & T. E. J. Leuteritz. 1999a. Geochelone Natural History Museum, The University of carbonaria. Catalogue of American Amphib- Kansas, Lawrence 2:1–52. ians and Reptiles 690:1–7. Duellman, W. E. 2005. Cusco Amazo´nico: the lives Ernst, C. H., & T. E. J. Leuteritz. 1999b. Geochelone of amphibians and reptiles in an Amazonian denticulata. Catalogue of American Amphib- rainforest. Comstock Publishing Associates, ians and Reptiles 691:1–6. Cornell University Press, Ithaca, New York, Ernst, R., M.-O. Ro¨del, & D. Arjoon. 2005. On the 488 pp. cutting edge—the anuran fauna of the Ma- Duellman, W. E., & M. L. Crump. 1974. Speciation bura Hill Forest Reserve, central Guyana. in frogs of the Hyla parviceps group in the Salamandra 41(4):179–194. Upper Amazon Basin. Occasional Papers of Eschscholtz, F. 1829. Zoologischer Atlas, enthaltend the Museum of Natural History, The Univer- Abbildungen und Beschreibungen neuer sity of Kansas, Lawrence 23:1–40. Thierarten, wa¨hrend des Flottcapitains von Duellman, W. E., & M. S. Hoogmoed. 1984. The Kotzebue zweiter Reise um die Welt, auf der taxonomy and phylogenetic relationships of Russisch-Kaiserlichen Kriegsschlupp Pred- the hylid frog genus Stefania. The University priaetie¨ in den Jahren 1823–1826. Part 1. G. of Kansas Museum of Natural History Reimer, Berlin. Miscellaneous Publications 75:1–39. Etheridge, R. 1970. A review of the South American Duellman, W. E., & M. S Hoogmoed. 1992. Some iguanid lizard genus Plica. Bulletin of the hylid frogs from the Guiana Highlands, British Museum (Natural History), Zoology northeastern South America: new species, 19(7):237–256. distributional records, and a generic realloca- Faivovich, J., C. F. B. Haddad, P. C. A. Garcia, D. tion. Occasional Papers of the Museum of R. Frost, J. A. Campbell, & W. C. Wheeler. Natural History, The University of Kansas, 2005. Systematic review of the frog family Lawrence 147:1–21. Hylidae, with special reference to Hylinae: 558 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

phylogenetic analysis and taxonomic revision. Fritz, U., & P. Havasˇ. 2006. Checklist of Chelonians Bulletin of the American Museum of Natural of the World. An online reference at http:// History 294:1–240. www.cites.org/common/com/NC/tax_ref/ Faivovich, J., C. F. B. Haddad, D. Baeˆta, K.-H. Chelonians_Checklist_2006.pdf (accessed on Jungfer, G. F. R. A´ lvares, R. A. Branda˜o, C. 19 August 2012). Sheil, L. S. Barrientos, C. L. Barrio-Amoro´s, Frost, D. R. (ed.). 1985. Amphibian species of the C. A. G. Cruz, & W. C. Wheeler. 2010. The world. A taxonomic and geographical refer- phylogenetic relationships of the charismatic ence. Allen Press and The Association of poster frogs, Phyllomedusinae (Anura, Hyli- Systematics Collections, Lawrence, Kansas, dae). Cladistics 26:227–261. 732 pp. Fernandes, D. S., F. L. Franco, & R. Fernandes. Frost, D. R. 2011. Amphibian species of the world: 2004. Systematic revision of the genus Lach- an online reference. Version 5.5 (31 January esis Daudin, 1803 (Serpentes, Viperidae). 2011). Electronic database accessible at http:// Herpetologica 60:245–260. research.amnh.org/vz/herpetology/amphibia/ Filippi, F. De. 1840. Catalogo ragionato e descrittivo (accessed on 27 May 2012). American Muse- della raccolta de’serpenti del Museo dell’I. R. um of Natural History, New York, U.S.A. Universita` di Pavia. Biblioteca Italiana (accessed on 19 August 2012). 99:163–187 & 306–343. Frost, D. R., & R. Etheridge. 1989. A phylogenetic Fitzinger, L. J. 1826. Neue Classification der analysis and taxonomy of iguanian lizards Reptilien nach ihren natu¨rlichen Verwandt- (Reptilia: Squamata). The University of Kan- schaften: Nebst einer Verwandtschafts-Tafel sas Museum of Natural History Miscella- und einem Verzeichnisse der Reptilien-Samm- neous Publication 81:1–65. lung des K. K. Zoologischen Museum’s zu Frost, D. R., M. T. Rodrigues, T. Grant, & T. A. Wien. J. G. Hu¨bner, Wien, 66 pp. Titus. 2001. Phylogenetics of the lizard genus Forlani, M. C., P. H. Bernardo, & H. Zaher. 2012. Tropidurus (Squamata: Tropiduridae: Tropi- Amphibia, Anura, Hylidae, Phyllomedusa durinae): direct optimization, descriptive effi- tarsius Cope, 1868: distribution extension, ciency, and sensitivity analysis of congruence new country record and geographic distribu- between molecular data and morphology. tion map. Check List 8:155–157. Molecular Phylogenetics and Evolution

Fouquet, A., P. Gaucher, M. Blanc, & C. M. Ve´lez- 21:352–371. Rodriguez. 2007a. Description of two new Frost, D. R., T. Grant, J. Faivovich, R. H. Bain, A. species of Rhinella (Anura: Bufonidae) from Haas, C. F. B. Haddad, R. O. de Sa´ ,A. the lowlands of the Guiana shield. Zootaxa Channing, M. Wilkinson, S. C. Donnellan, C. 1663:17–32. J. Raxworthy, J. A. Campbell, B. L. Blotto, P. Fouquet, A., M. Vences, M.-D. Salducci, A. Meyer, Moler, R. C. Drewes, R. A. Nussbaum, J. D. C. Marty, M. Blanc, & A. Gilles. 2007b. Lynch, D. M. Green, & W. C. Wheeler. 2006. Revealing cryptic diversity using molecular The amphibian tree of life. Bulletin of the phylogenetics and phylogeography in frogs of American Museum of Natural History 297:1– the Scinax ruber and Rhinella margaritifera 370. species groups. Molecular Phylogenetics and Fugler,C.M.1988.Anolis punctatus Daudin Evolution 43:567–582. (Reptilia: Iguanidae), un nuevo saurio regis- Franco, F. L., & R. R. Pinto. 2009. Stenostoma trado en Bolivia. Museo Nacional de Historia albifrons Wagler in Spix, 1824 as nomen Natural (Bolivia) Comunicacio´n 8:5–6. dubium and recognition of the name Leptoty- Gaige, H. T. 1926. A new frog from British Guiana. phlops tenellus Klauber, 1939 (Serpentes: Occasional Papers of the Museum of Zoology, Leptotyphlopidae). Salamandra 45(4):239– University of Michigan 176:1–3. 244. Gallagher, D. S., Jr., & J. R. Dixon. 1992. Franzen, M., & F. Glaw. 2007. Type catalogue of Taxonomic revision of the South American reptiles in the Zoologische Staatssammlung lizard genus Kentropyx Spix (Sauria: Teiidae). Mu¨nchen. Spixiana 30(2):201–274. Bollettino del Museo Regionale di Scienze Fretey, J., & R. Bour. 1980. Rede´couverte du type de Naturali, Torino 10:125–171. Dermochelys coriacea (Vandelli) (Testudinata, Gallardo, J. M. 1965. The species Bufo granulosus Dermochelyidae). Bolletino di zoologia 47(1– Spix (Salientia: Bufonidae) and its geographic 2):193–205. variation. Bulletin of the Museum of Com- Fretey, J., M. S. Hoogmoed, & J. Lescure. 1977. parative Zoology 134:107–138. Etude taxinomique de Rhinoclemmys punctu- Gamble, T., A. M. Bauer, E. Greenbaum, & T. R. laria punctularia (Daudin) (Testudinata, Emy- Jackman. 2008. Evidence for Gondwanan didae). Zoologische Mededelingen 52(6):63– vicariance in an ancient clade of gecko lizards. 80. Journal of Biogeography 35:88–104. VOLUME 125, NUMBER 4 559

Gamble, T., J. D. Daza, G. R. Colli, L. J. Vitt, & A. Goin, C. J. 1968 [1967]. A new Centrolenid frog from M. Bauer. 2011. A new genus of miniaturized Guyana. Quarterly Journal of the Florida and pug-nosed gecko from South America Academy of Science 30:115–118. (Sphaerodactylidae: Gekkota). Zoological Goin, C. J., & J. D. Woodley. 1969. A new tree-frog Journal of the Linnean Society 163:1244– from Guyana. Zoological Journal of the 1266. Linnean Society 48(1):135–140. Gans, C. 1962. Notes on amphisbaenids (Amphis- Gorzula, S., & J. C. Sen˜aris. 1999 [1998]. Contribu- baenia, Reptilia). 5. A redefinition and a tion to the herpetofauna of the Venezuelan bibliography of Amphisbaena alba Linne´. Guayana I: a data base. Scientia Guaianae American Museum Novitates 2105:1–31. 8:1–269. Gans, C. 1963. Notes on amphisbaenids (Amphis- Gower, D. J., M. Wilkinson, E. Sherratt, & P. J. R. baenia, Reptilia). 8. A redescription of Am- Kok. 2010. A new species of Rhinatrema ´ phisbaena stejnegeri and the description of a Dumeril & Bibron (Amphibia: Gymnophiona: Rhinatrematidae) from Guyana. Zootaxa new species of Amphisbaena from British 2391:47–60. Guiana. American Museum Novitates Grant, C. 1958. A new Gymnophthalmus (Reptilia, 2128:1–18. Teiidae) from Barbados, B.W.I. Herpetolog- Gans, C. 1964. A redescription of, and geographic ica 14:227–228. ´ variation in, Liophis miliaris Linne,the Grant, T., E. C. Humphrey, & C. W. Myers. 1997. common water snake of southeastern South The median lingual process of frogs: a bizarre America. American Museum Novitates character of Old World ranoids discovered in 2178:1–58. South American dendrobatids. American Mu- Gans, C. 1967. A check list of Recent amphisbae- seum Novitates 3212:1–40. nians (Amphisbaenia, Reptilia). Bulletin of Grant, T., D. R. Frost, J. P. Caldwell, R. Gagliardo, the American Museum of Natural History C. F. B. Haddad, P. J. R. Kok, D. B. Means, 135(2):63–105. B. P. Noonan, W. E. Schargel, & W. C. Gans, C. (ed.). 1969. Biology of the Reptilia, vol. 1, Wheeler. 2006. Phylogenetic systematics of Morphology A. (A. d’A. Bellairs & T. S. dart-poison frogs and their relatives (Amphib- Parsons, co-editors.) Academic Press, New ia: Athesphatanura: Dendrobatidae). Bulletin

York, 353 pp. of the American Museum of Natural History Garman, S. 1888 [1887]. West Indian Batrachia in 299:1–262. the Museum of Comparative Zoology. Bulle- Gray, J. E. 1825. A synopsis of the genera of reptiles tin of the Essex Institute 19:13–16. and Amphibia, with a description of some Garman, S. 1888 [1887]. On West Indian reptiles. new species. The Annals of Philosophy, New Iguanidae. Bulletin of the Essex Institute Series 10:193–217. 19:25–50. Gray, J. E. 1840. Catalogue of the species of reptiles Gasso´ Miracle, M. E., L. W. van den Hoek Ostende, collected in Cuba by W. S. MacLeay, Esq.;— & J. W. Arntzen. 2007. Type specimens of with some notes of their habits extracted from amphibians in the National Museum of his MS. Annals of Natural History; or, Natural History, Leiden, The Netherlands. Magazine of Zoology, Botany, and Geology Zootaxa 1482:25–68. 5:108–115. Gray, J. E. 1851. Description of a new genus and Gatesy, J., G. Amato, M. Norell, R. DeSalle, & C. family of Cyclosaurian lizards, from Para. Hayashi. 2003. Combined support for whole- Proceedings of the Zoological Society of sale taxic atavism in gavialine crocodylians. London 19:38–39. Systematic Biology 52:403–422. Guayasamin, J. M., S. Castroviejo-Fisher, J. Ayar- Glaw, F., & M. Franzen. 2006. Type catalogue of zagu¨ena, L. Trueb, & C. Vila`. 2008. Phyloge- amphibians in the Zoologische Staatssamm- netic relationships of glassfrogs (Centro- lung Mu¨nchen. Spixiana 29(2):153–192. lenidae) based on mitochondrial and nuclear Goeldi, E. A. 1907. Description of Hyla resinifictrix genes. Molecular Phylogenetics and Evolution Goeldi, a new Amazonian tree-frog peculiar 48(2):574–595. for its breeding-habits. Proceedings of the Guayasamin, J. M., S. Castroviejo-Fisher, L. Trueb, Zoological Society of London 1907:135–140. J. Ayarzagu¨ena, M. Rada, & C. Vila`. 2009. Goin, C. J. 1966a. A new frog of the genus Hyla Phylogenetic systematics of glassfrogs (Am- from British Guiana. Quarterly Journal of the phibia: Centrolenidae) and their sister taxon Florida Academy of Science 29:39–42. Allophryne ruthveni. Zootaxa 2100:1–97. Goin, C. J. 1966b. Description of a new frog of the Guibe´, J. 1950 [1948]. Catalogue des types d’am- genus Hyla from Suriname. Zoologische phibiens du Muse´um national d’histoire na- Mededelingen 41(15):229–232. turelle. Imprimerie Nationale, Paris, 71 pp. 560 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Guichenot, A. 1855. Expe´dition dans les parties Harvey, M. B. 2009 [2008]. New and poorly known centrales de l’Ame´rique du Sud. Partie 7: Dipsas (Serpentes: Colubridae) from northern Zoologie. Animaux nouveaux ou rares re- South America. Herpetologica 64(4):422–451. cueillis pendant l’expe´dition dans les parties Ha¨upl, M., & F. Tiedemann. 1978. Typenkatalog der centrales de l’Ame´rique du Sud, de Rio de Herpetologischen Sammlung. Amphibia. Ka- Janeiro a` Lima, et de Lima au Para; exe´cute´e taloge der Wissenschaftlichen Sammlungen par ordre du gouvernement franc¸ais pendant des Naturhistorischen Museums Wien 2, les anne´es 1843 a` 1847, sous la direction du Vertebrata 1, 34 pp. Comte Francis de Castelnau. Reptiles. Partie Hawkins, M. A., J. W. Sites, Jr., & B. P. Noonan. 7, tome 2, partie 2 P. Bertrand, Paris, 95 pp. 2007. Dendropsophus minutus (Anura: Hyli- Gu¨nther, A. 1858a. Neue Batrachier in der Samm- dae) of the Guiana Shield: using DNA lung des Britischen Museums. Archiv fu¨r barcodes to assess identity and diversity. Naturgeschichte 24(1):319–328. Zootaxa 1540:61–67. Gu¨nther, A. 1858b. Catalogue of colubrine snakes in Hedges, S. B., W. E. Duellman, & M. P. Heinicke. the collection of the British Museum. Taylor 2008. New World direct-developing frogs & Francis, London, 281 pp. (Anura: Terrarana): molecular phylogeny, Gu¨nther, A. 1859 [1858]. Catalogue of the Batrachia classification, biogeography, and conserva- salientia in the collection of the British tion. Zootaxa 1737:1–182. Museum. Taylor & Francis, London, 160 pp. Heinicke, M. P., W. E. Duellman, & S. B. Hedges. Gu¨nther, A. 1860. Third list of cold-blooded 2007. Major Caribbean and Central American Vertebrata collected by Mr. Fraser in Ecua- frog faunas originated by ancient oceanic dor. Proceedings of the Zoological Society of dispersal. Proceedings of the National Acad- London 28:233–240. emy of Sciences 104(24):10092–10097. Gu¨nther, A. 1872. Seventh account of new species of Heinicke, M. P., W. E. Duellman, L. Trueb, D. B. snakes in the collection of the British Muse- Means, R. D. MacCulloch, & S. B. Hedges. um. The Annals and Magazine of Natural 2009. A new frog family (Anura: Terrarana) History, Including Zoology, Botany, and from South America and an expanded direct- Geology, Fourth Series 9:13–37. developing clade revealed by molecular phy- Hahn, D. E. 1980. Liste der rezenten Amphibien und logeny. Zootaxa 2211:1–35. Reptilien, Anomalepididae, Leptotyphlopi- Henderson, R. W., P. Passos, & D. Feitosa. 2009. dae, Typhlopidae. Das Tierreich 101. Walter Geographic variation in the emerald treeboa, de Gruyter, Berlin, 93 pp. Corallus caninus (Squamata: Boidae). Copeia Hallowell, E. 1861. Report upon the Reptilia of the 2009:572–582. North Pacific Exploring Expedition, under command of Capt. John Rodgers, U. S. N. Hermann, J. 1804. Observationes zoologicae, quibus Proceedings of the Academy of Natural novae complures, aliaeque animalium species Sciences of Philadelphia (1860) 12:480–510. describuntur et illustrantur. Opus posthumum Hardy, D. L., Sr. 1994. Bothrops asper (Viperidae) edidit Fridericus Ludovicus Hammer. Argen- snakebite and field researchers in Middle torati apud Amandum Koenig, Parisiis, 332 America. Biotropica 26:198–207. pp. Hardy, J. D., Jr., & H. S. Harris, Jr. 1979. Heyer, W. R. 1972. The status of Leptodactylus Occurrence of the West Indian frog, Eleuther- pumilio Boulenger (Amphibia, Leptodactyli- odactylus johnstonei, in South America and on dae) and the description of a new species of the island of Curac¸ao. Bulletin of the Mary- Leptodactylus from Ecuador. Contributions in land Herpetological Society 15:124–133. Science, Natural History Museum of Los Hardy, L. M., C. J. Cole, & C. R. Townsend. 1989. Angeles County 231:1–8. Parthenogenetic reproduction in the neotrop- Heyer, W. R. 1973. Systematics of the marmoratus ical unisexual lizard, Gymnophthalmus under- group of the frog genus Leptodactylus (Am- woodi (Reptilia: Teiidae). Journal of phibia, Leptodactylidae). Contributions in Morphology 201:215–234. Science, Natural History Museum of Los Harris, H. S., Jr., & R. S. Simmons. 1977. A new Angeles County 251:1–50. subspecies of Crotalus durissus (Serpentes: Heyer, W. R. 1975. Adenomera lutzi (Amphibia: Crotalidae) from the Rupununi Savanna of Leptodactylidae), a new species of frog from southwestern Guyana. Memo´rias do Instituto Guyana. Proceedings of the Biological Society de Butantan 40/41:305–311. of Washington 88:315–318. Harshman, J., C. J. Huddleston, J. P. Bollback, T. J. Heyer, W. R. 1979. Systematics of the pentadactylus Parsons, & M. J. Braun. 2003. True and false species group of the frog genus Leptodactylus gharials: a nuclear gene phylogeny of Croc- (Amphibia: Leptodactylidae). Smithsonian odylia. Systematic Biology 52(3):386–402. Contributions to Zoology 301:1–43. VOLUME 125, NUMBER 4 561

Heyer, W. R. 2005. Variation and taxonomic Hoogmoed, M. S. 1979. The herpetofauna of the clarification of the large species of the Guianan Region. Pp. 241–279 in W. E. Leptodactylus pentadactylus species group Duellman, ed., The South American Herpe- (Amphibia: Leptodactylidae) from Middle tofauna: its origin, evolution, and dispersal. America, northern South America, and Am- Monograph No. 7 of the Museum of Natural azonia. Arquivos de Zoologia, Sa˜o Paulo History, The University of Kansas, Lawrence, 37(3):269–348. 485 pp. Heyer, W. R., & R. O. de Sa´ . 2011. Variation, Hoogmoed, M. S. 1980. Revision of the genus systematics, and relationships of the Lepto- Atractus in Surinam, with the resurrection of dactylus bolivianus complex (Amphibia: Anu- two species (Colubridae, Reptilia). Notes on ra: Leptodactylidae). Smithsonian Contribu- the Herpetofauna of Surinam VII. Zoologi- tions to Zoology 635:1–58. sche Verhandelingen 175:1–47. Heyer, W. R., & M. M. Heyer. 2006. Leptodactylus Hoogmoed, M. S. 1982. Nomenclatural problems knudseni. Catalogue of American Amphibians relating to Atractus trilineatus Wagler, 1828. and Reptiles 807:1–12. Zoologische Mededelingen 56(10):131–138. Heyer, W. R., & A. S. Thompson. 2000. Leptodac- Hoogmoed, M. S. 1983 [1982]. Snakes of the tylus rugosus. Catalogue of American Am- Guianan region. Memo´rias do Instituto Bu- phibians and Reptiles 743:1–5. tantan 46:219–254. Hillis, D. M., & T. P. Wilcox. 2005. Phylogeny of the Hoogmoed, M. S., & T. C. S. Avila-Pires. 1992. New World true frogs (Rana). Molecular Studies on the species of the South American Phylogenetics and Evolution 34(2):299–314. lizard genus Arthrosaura Boulenger (Reptilia: Hoffmann, M., et al. 2010. The impact of conserva- Sauria: Teiidae), with the resurrection of two tion on the status of the world’s vertebrates. species. Zoologische Mededelingen 66:453– Science 330:1503–1509. 484. Hoogmoed, M. S., & S. J. Gorzula. 1979. Checklist Hoge, A. R. 1954 [1953]. A new Bothrops from of the savanna inhabiting frogs of the El Brazil. Bothrops brazili, sp. nov. Memo´rias do Manteco region with notes on their ecology Instituto Butantan 25:15–21. and the description of a new species of Hoge, A. R. 1958 [1959]. E´ tude sur Apostolepis treefrog (Hylidae, Anura). Zoologische Me- coronata (Sauvage 1877) et Apostolepis quin- dedelingen 54(13):183–216. quelineata Boulenger 1896. Memo´rias do Hoogmoed, M. S., & U. Gruber. 1983. Spix and Instituto Butantan 28:73–78. Wagler type specimens of reptiles and am- Hoge, A. R., & A. R. Lancini V. 1962. Sinopsis de phibians in the Natural History Musea in las serpientes venenosas de Venezuela. Pub- Munich (Germany) and Leiden (The Nether- licaciones Ocasionales del Museo de Ciencias lands). Spixania, Supplement 9:319–415. Naturales, Caracas, Venezuela, Zoologia 1:1– Hoogmoed, M. S., & J. Lescure. 1984. A new genus 24. and two new species of minute leptodactylid Hoge, A. R., & S. A. R. W. D. L. Romano. 1966 frogs from northern South America, with [1965]. Leptomicrurus in Brasil (Serpentes: comments upon Phyzelaphryne (Amphibia: Elapidae). Memo´rias do Instituto Butantan Anura: Leptodactylidae). Zoologische Mede- 32:1–8. delingen 58(6):85–115. Hoge, A. R., N. P. Santos, C. Heitor, L. A. Lopes, & Hoogmoed, M. S., & T. Mott. 2003. On the identity I. M. de Souza. 1973 [1972]. Serpentes of Amphisbaena hugoi Vanzolini, 1990 (Repti- coletadas pelo projeto Rondon VII em lia: Squamata: Amphisbaenidae). Zoologische Iauareteˆ, Brasil. Memo´rias do Instituto Bu- Mededelingen 77:455–457. tantan 36:221–232. Houttuyn, M. 1782. Het onderscheid der salaman- Hollis, J. L. 2006. Phylogenetics of the genus deren van de haagdissen in ’t algemeen, en van Chironius Fitzinger, 1826 (Serpentes, Colubri- de gekkoos in ’t byzonder. Verhandelingen dae) based on morphology. Herpetologica uitgegeven door het Zeeuwsch Genootschap 62(4):435–453. der Wetenschappen te Vlissingen 9:305–336. Hoogmoed, M. S. 1973. Notes on the herpetofauna Huber, O. 1995. Vegetation. Pp. 97–160 in J. A. of Surinam IV: the lizards and amphisbae- Steyermark, P. E. Berry, & B. K. Holst, eds., nians of Surinam. Biogeographica, vol. 4. Dr. Flora of the Venezuelan Guayana, vol. 1, W. Junk b.v. Publishers, The Hague, 419 pp. Introduction. Missouri Botanical Garden Hoogmoed, M. S. 1977. On the presence of Bufo Press, St. Louis, 363 pp. nasicus Werner in Guiana, with a redescrip- Huber, O., G. Gharbarran, & V. Funk. 1995. tion of the species on the basis of recently Preliminary vegetation map of Guyana. Bio- collected material. Zoologische Mededelingen logical Diversity of the Guianas Programme, 51(16):265–275. Smithsonian Institution, Washington, D.C. 562 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

International Commission of Zoological Nomencla- rapid dispersal through human interactions. ture (ICZN). 1999. ‘‘The Code’’ International Herpetological Review 33:290–294. code of zoological nomenclature, 4th Edition. Kearney, M. 2003. Systematics of the Amphisbaenia International Trust for Zoological Nomencla- (Lepidosauria: Squamata) based on morpho- ture, London, 306 pp. logical evidence from recent and fossil forms. IUCN. 2010. The IUCN red list of threatened Herpetological Monographs 17:1–74. species. (Ver. 2010.4). Available at http:// Kelloff, C. L. 2003. The use of biodiversity data in www.iucnredlist.org/ (accessed 27 October developing Kaieteur National Park, Guyana 2010). for ecotourism and conservation. Contribu- Izecksohn, E. 1976. Uma nova espe´cie de Pipa,do tions to the Study of Biological Diversity 1:1– estado do Amazonas, Brasil (Amphibia, 44. Anura, Pipidae). Revista Brasileira de Biol- Kizirian, D. A., & C. J. Cole. 1999. Origin of the ogia 36:507–510. unisexual lizard Gymnophthalmus underwoodi Jan, G. 1858. Plan d’une Iconographie descriptive des (Gymnophthalmidae) inferred from mito- Ophidiens, et description sommaire de nou- chondrial DNA nucleotide sequences. Molec- velles espe`ces de Serpents. Revue et Magasin ular Phylogenetics and Evolution 11(3):394– de Zoologie Pure et Applique´e, se´rie 2, 400. 10:514–527. Klaczko, J., F. A. Machado, G. Scrocchi, & H. Jan, G. 1861 [1862]. Note sulla famiglia dei Tiflopidi Zaher. 2010. Taxonomic status of Chironius sui loro generi e sulle specie del genere multiventris and Chironius cochranae (Ser- Stenostoma. Archivio per la Zoologia l’Anat- pentes). Herpetologica 66(4):476–484. omia e la Fisiologia 1:178–199. Klauber, L. M. 1939. Three new worm snakes of the Jan, G. 1863. Elenco sistematico degli ofidi descritti e genus Leptotyphlops. Transactions of the San disegnati per l’iconografia generale. A. Lom- Diego Society of Natural History 9:59–65. bardi, Milano, 143 pp. Kluge, A. G. 1969. The evolution and geographical Jorge da Silva, N., Jr. 1993. The snakes from Samuel origin of the New World Hemidactylus hydroelectric power plant and vicinity, mabouia-brookii complex (Gekkonidae, Sau- Rondoˆnia, Brazil. Herpetological Natural ria). Miscellaneous Publications Museum of

History 1:37–86. Zoology, University of Michigan 138:1–78. Jungfer, K.-H. 2010. The taxonomic status of some Kluge, A. G. 1993. Gekkonoid lizard taxonomy. spiny-backed treefrogs, genus Osteocephalus International Gecko Society, San Diego, (Amphibia: Anura: Hylidae). Zootaxa California, 245 pp. 2407:28–50. Kok, P. J. R. 2005. A new genus and species of Jungfer, K.-H., & W. Bo¨hme. 2004. A new poison- gymnophthalmid lizard (Squamata: Gym- dart frog (Dendrobates) from northern central nophthalmidae) from Kaieteur National Park, Guyana (Amphibia: Anura: Dendrobatidae). Guyana. Bulletin de l’Institut Royal des Salamandra 40(2):99–104. Sciences Naturelles de Belgique, Biologie Jungfer, K.-H., & W. Ho¨dl. 2002. A new species of 75:35–45. Osteocephalus from Ecuador and a redescrip- Kok, P. J. R. 2006a. A new snake of the genus tion of O. leprieurii (Dume´ril & Bibron, 1841) Atractus Wagler, 1828 (Reptilia: Squamata: (Anura: Hylidae). Amphibia-Reptilia 23:21– Colubridae) from Kaieteur National Park, 46. Guyana, northeastern South America. Zoo- Jungfer, K.-H., & L. C. Schiesari. 1995. Description taxa 1378:19–35. of a central Amazonian and Guianan tree Kok, P. J. R. 2006b. A new species of Hypsiboas frog, genus Osteocephalus (Anura, Hylidae), (Amphibia: Anura: Hylidae) from Kaieteur with oophagous tadpoles. Alytes 13:1–13. National Park, eastern edge of the Pakaraima Jungfer, K.-H., & P. Weygoldt. 1999. Biparental care Mountains, Guyana. Bulletin de l’Institut in the tadpole-feeding Amazonian treefrog Royal des Sciences Naturelles de Belgique, Osteocephalus oophagus. Amphibia-Reptilia Biologie 76:191–200. 20(3):235–249. Kok, P. J. R. 2008. A new highland species of Kaiser, H., J. D. Hardy, Jr., & D. M. Green. 1994. Arthrosaura Boulenger, 1885 (Squamata: Taxonomic status of Caribbean and South Gymnophthalmidae) from Maringma tepui American frogs currently ascribed to Eleuther- on the border of Guyana and Brazil. Zootaxa odactylus urichi (Anura: Leptodactylidae). 1909:1–15. Copeia 1994(3):780–796. Kok, P. J. R. 2009a. A new species of Oreophrynella Kaiser, H., C. L. Barrio-Amoro´s, J. D. Trujillo, & J. (Anura: Bufonidae) from the Pantepui region D. Lynch. 2002. Expansion of Eleutherodac- of Guyana, with notes on O. macconnelli tylus johnstonei in northern South America: Boulenger, 1900. Zootaxa 2071:35–49. VOLUME 125, NUMBER 4 563

Kok, P. J. R. 2009b. Lizard in the clouds: a new carrying frog): reproduction. Herpetological highland genus and species of Gymnophthal- Review 37(2):212–213. midae (Reptilia: Squamata) from Maringma Kok, P. J. R., M. N. C. Kokubum, R. D. tepui, western Guyana. Zootaxa 1992:53–67. MacCulloch, & A. Lathrop. 2007. Morpho- Kok, P. J. R. 2010a. A redescription of Anomalo- logical variation in Leptodactylus lutzi (Anu- glossus praderioi (La Marca, 1998) (Anura: ra, Leptodactylidae) with description of its Aromobatidae: Anomaloglossinae), with de- advertisement call and notes on its courtship scription of its tadpole and call. Pape´is behaviour. Phyllomedusa 6:45–60. Avulsos de Zoologia, Sa˜o Paulo 50(4):51–68. Kok, P. J. R., R. D. MacCulloch, A. Lathrop, B. Kok, P. J. R. 2010b. A new species of Chironius Willaert, & F. Bossuyt. 2010. A new species of Fitzinger, 1826 (Squamata: Colubridae) from Anomaloglossus (Anura: Aromobatidae) from the Pantepui region, northeastern South the Pakaraima Mountains of Guyana. Zoo- America. Zootaxa 2611:31–44. taxa 2660:18–32. Kok, P. J. R. 2011. A new species of the genus Kok, P. J. R., D. B. Means, & F. Bossuyt. 2011. A Gonatodes Fitzinger, 1843 (Reptilia: Sphaer- new highland species of Pristimantis Jime´nez odactylidae) from central Guyana, northern de la Espada, 1871 (Anura: Strabomantidae) South America. Zootaxa 3018:1–12. from the Pantepui region, northern South Kok, P. J. R., & P. Benjamin. 2007. Stefania evansi America. Zootaxa 2934:1–19. (Groete Creek carrying frog): froglet carrying. Lace´pe`de, B. G. E. 1788. Histoire naturelle des Herpetological Bulletin 100:38–39. quadrupe`des des ovipares et des serpens, vol. Kok, P. J. R., & S. Castroviejo-Fisher. 2008. 1. Imprierie du Roi, Hoˆtel de Thou, Paris, xvii Glassfrogs (Anura: Centrolenidae) of Kaie- þ 651 pp. teur National Park, Guyana, with notes on La Marca, E. 1997 [1996]. Ranas del ge´nero the distribution and taxonomy of some species Colostethus (Amphibia: Anura: Dendrobati- of the family in the Guiana Shield. Zootaxa dae) de la Guayana Venezolana con la 1680:25–53. descripcion de siete especies nuevas. Publica- Kok, P. J. R., & R. Ernst. 2007. A new species of ciones de la Asociacio´n Amigos de Don˜ana Allobates (Anura: Aromobatidae: Allobati- 9:1–64. nae) exhibiting a novel reproductive behav- Laguna, M. M., M. T. Rodrigues, R. M. L. dos

iour. Zootaxa 1555:21–38. Santos, Y. Yonenaga-Yassuda, T. C. S. A´ vila- Kok, P. J. R., & M. Kalamandeen. 2008. Introduc- Pires, M. S. Hoogmoed, & K. C. M. tion to the taxonomy of the amphibians of Pellegrino. 2010. Karyotypes of a cryptic Kaieteur National Park, Guyana. Abc Taxa diploid form of the unisexual Leposoma 5:1–278. percarinatum (Squamata, Gymnophthalmi- Kok, P. J. R., J. A. Roze, G. L. Lenglet, H. Sambhu, dae) and the bisexual Leposoma ferreirai from & D. Arjoon. 2003. Micrurus isozonus (Cope, the lower Rio Negro, Amazonian Brazil. 1860) (Serpentes, Elapidae): an addition to the Journal of Herpetology 44(1):153–157. herpetofauna of Guyana, with comments on Lara-Ruiz, P., G. G. Lopez, F. R. Santos, & L. S. other species of coral snakes from Guyana. Soares. 2006. Extensive hybridization in Bulletin de l’Institut Royal des Sciences hawksbill turtles (Eretmochelys imbricata) Naturelles de Belgique, Biologie 73:73–79. nesting in Brazil revealed by mtDNA analy- Kok, P. J. R., G. R. Bourne, H. Sambhu, & G. L. ses. Conservation Genetics 7(5):773–781. Lenglet. 2005. Phrynohyas coriacea (Suriname Lathrop, A., & R. D. MacCulloch. 2007. A new golden-eyed treefrog). Herpetological Review species of Oreophrynella (Anura: Bufonidae) 36(4):462–463. from Mount Ayanganna, Guyana. Herpeto- Kok, P. J. R., R. D. MacCulloch, P. Gaucher, E. H. logica 63:87–93. Poelman, G. R. Bourne, A. Lathrop, & G. L. Laurenti, J. N. 1768. Specimen medicum, exhibens Lenglet. 2006a. A new species of Colostethus synopsin reptilium emendatam cum experi- (Anura, Dendrobatidae) from French Guiana mentis circa venena et antidota reptilium with a redescription of Colostethus beebei Austriacorum. Joan. Thomae nob. de Tratt- (Noble, 1923) from its type locality. Phyllo- nern, Viennae, 214 pp. medusa 5(1):43–66. Lavilla, E. O., M. Vaira, & L. Ferrari. 2003. A new Kok, P. J. R., H. Sambhu, I. Roopsind, G. L. species of Elachistocleis (Anura: Microhyli- Lenglet, & G. R. Bourne. 2006b. A new dae) from the Andean Yungas of Argentina, species of Colostethus (Anura: Dendrobati- with comments on the Elachistocleis ovalis–E. dae) with maternal care from Kaieteur Na- bicolor controversy. Amphibia-Reptilia tional Park, Guyana. Zootaxa 1238:35–61. 24:269–284. Kok, P. J. R., G. R. Bourne, P. Benjamin, & G. L. Lavilla, E. O., J. A. Langone, J. M. Padial, & R. O. Lenglet. 2006c. Stefania evansi (Groete Creek de Sa´ . 2010. The identity of the crackling, 564 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

luminescent frog of Suriname (Rana typhonia nymis, locis, 12th edition, vol. 1. Holmiae, Linnaeus, 1758) (Amphibia, Anura). Zootaxa Laurentii Salvii, 532 pp. 2671:17–30. Lynch, J. D. 1970. Systematic status of the American Lawson, R., J. B. Slowinski, B. I. Crother, & F. T. leptodactylid frog genera Engystomops, Eu- Burbrink. 2005. Phylogeny of the Colubroidea pemphix,andPhysalaemus. Copeia (Serpentes): new evidence from mitochondrial 1970(3):488–496. and nuclear genes. Molecular Phylogenetics Lynch, J. D. 1989. A review of the leptodactylid and Evolution 37:581–601. frogs of the genus Pseudopaludicola in north- Lazell, J. D., Jr. 1972. The Anoles (Sauria, Iguani- ern South America. Copeia 1989(3):577–588. dae) of the Lesser Antilles. Bulletin of the Lynch, J. D., & M. S. Hoogmoed. 1977. Two new Museum of Comparative Zoology 143:1–115. species of Eleutherodactylus (Amphibia: Lep- Lema, T. de. 1984. Sobre o geˆnero Elapomorphus todactylidae) from northeastern South Amer- Wiegmann, 1843 (Serpentes, Colubridae, Ela- ica. Proceedings of the Biological Society of pomorphinae). Iheringia, Se´rie Zoologia Washington 90:424–439. 64:53–86. MacCulloch, R. D., & A. Lathrop. 2001. A new Lema, T. de, & N. R. de Albuquerque. 2010. The species of Arthrosaura (Sauria: Teiidae) from identity of Apostolepis pymi and placement of the highlands of Guyana. Caribbean Journal A. quinquelineata in the synonymy of A. of Science 37:174–181. nigrolineata (Serpentes, Xenodontinae). Biota MacCulloch, R. D., & A. Lathrop. 2002. Exception- Neotropica 10:343–346. al diversity of Stefania (Anura: Hylidae) on Lescure, J. 1981. Contribution a` l’e´tude des amphib- Mount Ayanganna, Guyana: three new spe- iens de Guyane franc¸aise. VIII. Validation cies and new distribution records. Herpeto- d’Atelopus spumarius Cope, 1871, et logica 58:327–346. de´signation d’un ne´otype. Description d’Ate- MacCulloch, R. D., & A. Lathrop. 2004a. A new lopus spumarius barbotini nov. ssp. Donne´es species of Dipsas (Squamata: Colubridae) e´tho-e´cologiques et bioge´ographiques sur les from Guyana. Revista de Biolog´ıa Tropical Atelopus du groupe flavescens (Anoures, 52:239–247. Bufonide´s).BulletinduMuse´um national MacCulloch, R. D., & A. Lathrop. 2004b. Micrurus

d’Histoire naturelle, Paris, 4e se´r., section A, ibiboboca (Serpentes, Elapidae) is not a 3:893–910. Guiana Shield species. Phyllomedusa 3:141– Lescure, J., & C. Marty. 2000. Atlas des amphibiens 144. de Guyane. Publications Scientifiques du MacCulloch, R. D., & A. Lathrop. 2005. Hylid frogs Muse´um national d’Histoire naturelle, Paris, from Mount Ayanganna, Guyana: new spe- 388 pp. cies, redescriptions, and distributional re- Lescure, J., V. Marty, C. Marty, F. Starace, M. cords. Phyllomedusa 4:17–37. Auber-Thomay, & F. Letellier. 1995. Contri- MacCulloch, R. D., & A. Lathrop. 2006a. Stefania bution a` l’e´tude des amphibiens de Guyana ackawaio. Catalogue of American Amphibi- franc¸aise. X. Les Phyllomedusa (Anura, Hyli- ans and Reptiles 824:1–2. dae). Revue franc¸aise d’Aquariologie 22:35– MacCulloch, R. D., & A. Lathrop. 2006b. Stefania 50. ayangannae. Catalogue of American Amphib- Lichtenstein, H. 1823. Verzeichniss der Doubletten ians and Reptiles 825:1–2. des zoologischen Museums der Ko¨nigl. Uni- MacCulloch, R. D., & A. Lathrop. 2006c. Stefania versita¨t zu Berlin nebst Beschreibung vieler coxi. Catalogue of American Amphibians and bisher unbekannter Arten von Sa¨ugethieren, Reptiles 826:1–2. Vo¨geln, Amphibien und Fischen. T. Traut- MacCulloch, R. D., & A. Lathrop. 2006d. Stefania wein, Berlin, 118 pp. evansi. Catalogue of American Amphibians Lieb, C. S. 1988. Systematic status of the neotropical and Reptiles 827:1–3. snakes Dendrophidion dendrophis and D. MacCulloch, R. D., & A. Lathrop. 2006e. Stefania nuchalis (Colubridae). Herpetologica 44:162– roraimae. Catalogue of American Amphibians 175. and Reptiles 828:1–2. Linnaei, C. 1939. Caroli Linnaei Systema naturae; a MacCulloch, R. D., & A. Lathrop. 2006f. Stefania photographic facsimile of the first volume of woodleyi. Catalogue of American Amphibians the tenth edition (1758): Regnum animale. and Reptiles 829:1–2. Printed by order of the Trustees, British MacCulloch, R. D., & A. Lathrop. 2009. Herpeto- Museum (Natural History), London, 823 pp. fauna of Mount Ayanganna, Guyana: results Linne´, C. 1766. Systema naturae per regna tria of the Royal Ontario Museum Ayanganna naturae, secundum classes, ordines, genera, Expedition 2000. Royal Ontario Museum species, cum characteribus, differentiis, syno- Contributions in Science 4:1–35. VOLUME 125, NUMBER 4 565

MacCulloch, R. D., A. Lathrop, & S. Z. Khan. 2006. Mausfeld, P., A. Schmitz, W. Bo¨hme, B. Misof, D. Exceptional diversity of Stefania (Anura, Vrcibradic, & C. F. D. Rocha. 2002. Phylo- Cryptobatrachidae) II: six species from genetic affinities of Mabuya atlantica Schmidt, Mount Wokomung, Guyana. Phyllomedusa 1945, endemic to the Atlantic Ocean archipel- 5:31–41. ago of Fernando de Noronha (Brazil): neces- MacCulloch, R. D., A. Lathrop, R. P. Reynolds, J. sity of partitioning the genus Mabuya C. Sen˜aris, & G. E. Schneider. 2007. Herpe- Fitzinger, 1826 (Scincidae: Lygosominae). tofauna of Mount Roraima, Guiana Shield Zoologischer Anzeiger 241:281–293. region, northeastern South America. Herpe- McCord, W. P., M. Joseph-Ouni, & W. W. Lamar. tological Review 38:24–30. 2001. A taxonomic reevaluation of Phrynops MacCulloch, R. D., A. Lathrop, P. J. R. Kok, L. R. (Testudines: Chelidae) with the description of Minter, S. Z. Khan, & C. L. Barrio-Amoro´s. two new genera and a new species of 2008a. A new species of Adelophryne (Anura: Batrachemys. Revista de Biolog´ıa Tropical, Eleutherodactylidae) from Guyana, with ad- San Jose´, Costa Rica 49(2):715–764. ditional data on A. gutturosa. Zootaxa McDiarmid, R. W., & M. A. Donnelly. 2005. The 1884:36–50. herpetofauna of the Guayana Highlands: MacCulloch, R. D., A. Lathrop, L. R. Minter, & S. amphibians and reptiles of the Lost World. Z. Khan. 2008b. Otophryne (Anura: Micro- Pp. 461–560 in M. A. Donnelly, B. I. Crother, hylidae) from the highlands of Guyana: C. Guyer, M. H. Wake, & M. E. White, eds., redescriptions, vocalisations, tadpoles and Ecology and evolution in the tropics: a new distributions. Pape´is Avulsos de Zoolog- herpetological perspective. The University of ia, Sa˜o Paulo 48(22):247–261. Chicago Press, Chicago, 675 pp. MacCulloch, R. D., A. Lathrop, P. J. R. Kok, R. McDiarmid, R. W., T’S. A. Toure´, & J. M. Savage. Ernst, & M. Kalamandeen. 2009. The genus 1996. The proper name of the neotropical tree Oxyrhopus (Serpentes: Dipsadidae: Xenodon- boa often referred to as Corallus enydris tinae) in Guyana: morphology, distributions (Serpentes: Boidae). Journal of Herpetology and comments on taxonomy. Pape´is Avulsos 30(3):320–326. de Zoologia, Sa˜o Paulo 49(36):487–495. McDiarmid, R. W., J. A. Campbell, & T’S. A.

Malnate, E. V. 1971. A catalog of primary types in Toure´. 1999. Snake species of the world: a the herpetological collections of the Academy taxonomic and geographic reference, vol. 1. of Natural Sciences, Philadelphia (ANSP). The Herpetologists’ League, Washington, Proceedings of the Academy of Natural D.C., 511 pp. Sciences of Philadelphia 123:345–375. Means, D. B. 2004. Geographic distribution: Bo- Manzanilla, J., & D. Sa´ nchez. 2005 [2004]. Una thriopsis taeniata. Herpetological Review nueva especie de Thamnodynastes (Serpentes: 35(4):410. Colubridae) del macizo del Turimiquire, Means, D. B. 2007. Geographic distribution: Atrac- noreste de Venezuela. Memoria de la tus zidoki. Herpetological Review 38:484–485. Fundacio´n La Salle de Ciencias Naturales Means, D. B., & J. M. Savage. 2007. Three new 161–162:61–75. malodorous rainfrogs of the genus Pristiman- Markezich, A. L., C. J. Cole, & H. C. Dessauer. tis (Anura: Brachycephalidae) from the Wo- 1997. The blue and green whiptail lizards komung Massif in west-central Guyana, (Squamata: Teiidae: Cnemidophorus) of the South America. Zootaxa 1658:39–55. Peninsula de Paraguana, Venezuela: system- Means, D. B., W. E. Duellman, & V. C. Clark. 2008. atics, ecology, descriptions of two new taxa, Ovipositing behavior in the egg-brooding frog and relationships to whiptails of the Guianas. Stefania ayangannae (Anura, Hemiphracti- American Museum Novitates 3207:1–60. dae). Phyllomedusa 7(2):143–148. Maschio, G. F., A. L. C. Prudente, F. S. Rodrigues, Medem, F. 1983. Los Crocodylia de Sur Ame´rica, & M. S. Hoogmoed. 2010. Food habits of vol. II, Venezuela, Trinidad, Tobago, Guya- Anilius scytale (Serpentes: Aniliidae) in the na, Suriname, Guayana francesa, Ecuador, Brazilian Amazonia. Zoologia (Curitiba, Im- Peru´, Bolivia, Brasil, Paraguay, Argentina, presso) 27:184–190. Uruguay. Universidad Nacional de Colombia Maslin, T. P., & D. M. Secoy. 1986. A checklist of y Fondo Colombiano de Investigaciones the lizard genus Cnemidophorus (Teiidae). Cientificas y Proyectos Especiales ‘‘Francisco Contributions in Zoology, University of Jose´ de Caldas,’’ Bogota´ , 270 pp. Colorado Museum 1:1–60. Melin, D. 1941. Contributions to the knowledge of Massary, J-C. D., & I. Ineich. 1999. Uranoscodon the amphibia of South America. Go¨teborgs superciliosus: diving. Herpetological Review Kungliga Vetenskaps och Vitterhets 30(3):168. Samha¨lles Handlingar, Ser. B, Band 1, 4:1–71. 566 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Merrem, B. 1820. Versuch eines Systems der Murphy, J. C. 1997. Amphibians and reptiles of Amphibien (Tentamen systematis amphibio- Trinidad and Tobago. Krieger Publishing rum). J. C. Kreiger, Marburg, Germany, 194 Company, Malabar, Florida, 245 pp. pp. Myers, C. W., & M. A. Donnelly. 1997. A tepui Mertens, R. 1967. Die herpetologische Sektion des herpetofauna on a granitic mountain (Tam- Natur-Museums und Forschungs-Institutes acuari) in the borderland between Venezuela Senckenberg in Frankfurt a. M. nebst einem and Brazil: report from the Phipps Tapirapeco´ Verzeichnis ihrer Typen. Senckenbergiana Expedition. American Museum Novitates Biologica 48A:1–106. 3213:1–71. Michaud, E. J., & J. R. Dixon. 1987. Taxonomic Myers, C. W., & M. A. Donnelly. 2001. Herpeto- revision of the Liophis lineatus complex fauna of the Yutaje´-Corocoro massif, Ven- (Reptilia: Colubridae) of Central and South ezuela: second report from the Robert G. America. Milwaukee Public Museum Contri- Goelet American Museum-Terramar expedi- butions in Biology and Geology 71:1–26. tion to the northwestern tepuis. Bulletin of the Miralles, A., G. R. Fuenmayor, C. Bonillo, W. E. American Museum of Natural History 261:1– Schargel, T. Barros, J. E. Garc´ıa-Perez, & C. 85. L. Barrio-Amoro´s. 2009. Molecular systemat- Myers, C. W., & M. A. Donnelly. 2008. The summit ics of Caribbean skinks of the genus Mabuya herpetofauna of Auyantepui, Venezuela: re- (Reptilia, Scincidae), with descriptions of two port from the Robert G. Goelet American new species from Venezuela. Zoological Jour- Museum-Terramar expedition. Bulletin of the nal of the Linnean Society 156(3):598–616. American Museum of Natural History 308:1– Miranda-Ribeiro, A. de. 1926. Notas para servirem 147. ao´ estudo dos Gymnobatrachios (Anura) Narvaes, P., & M. T. Rodrigues. 2009. Taxonomic Brasileiros. Archivos do Museu Nacional do revision of Rhinella granulosa species group Rio de Janeiro 27:1–227. (Amphibia, Anura, Bufonidae), with a de- Mocquard, F. 1904. Description de quelques Rep- scription of a new species. Arquivos de tiles et d’un Batracien nouveaux de la Zoologia, Sa˜o Paulo 40:1–73. collection du Muse´um. Bulletin du Muse´um Nelson, C. E., & J. Lescure. 1975. The taxonomy and d’Histoire Naturelle 10:301–309. distribution of Myersiella and Synapturanus

Montingelli, G. G., & H. Zaher. 2011. New species (Anura: Microhylidae). Herpetologica of Mastigodryas Amaral, 1934 from Brazilian 31(4):389–397. Amazonia and Guyana (Serpentes: Colubri- Noble, G. K. 1923. New batrachians from the dae). Journal of Herpetology 45(1):111–119. Tropical Research Station British Guiana. Moreau de Jonne`s, A. 1818. Monographie du Zoologica, Scientific Contributions of the Mabouia des murailles, ou Gecko Mabouia New York Zoological Society 3(14):289–299. des Antilles. Bulletin des Sciences, par La Noonan, B. P., & R. M. Bonett. 2003. A new species Socie´te´ Philomatique de Paris 3:138–139. of Hyalinobatrachium (Anura: Centrolenidae) Mott, T., & D. R. Vieites. 2009. Molecular from the highlands of Guyana. Journal of phylogenetics reveals extreme morphological Herpetology 37:91–97. homoplasy in Brazilian worm lizards chal- Noonan, B. P., & P. Gaucher. 2005. Phylogeography lenging current taxonomy. Molecular Phylo- and demography of Guianan harlequin toads genetics and Evolution 51:190–200. (Atelopus): diversification within a refuge. Mu¨ller, L. 1923. Neue oder seltene Reptilien und Molecular Ecology 14:3017–3031. Batrachier der Zoologischen Sammlung des Noonan, B. P., & M. B. Harvey. 2000. A new species bayrischen Staates. Zoologischer Anzeiger of glass frog (Anura: Centrolenidae) from the 57:38–42. highlands of Guyana. Herpetologica Mu¨ller, L., & W. Hellmich. 1936. Amphibien und 56(3):294–302. Reptilien. I. Teil: Amphibia, Chelonia, Lor- Noonan, B. P., & J. W. Sites, Jr. 2010. Tracing the icata. Wissenschaftliche Ergebnisse der Deut- origins of iguanid lizards and boine snakes of schen Gran Chaco-Expedition. Band 4. the Pacific. The American Naturalist 175:61– Strecker und Schro¨der, Stuttgart, 120 pp. 72. Murphy, J. C. 1996. Crossing Bond’s Line: the Nussbaum, R. A., & M. S. Hoogmoed. 1979. herpetofaunal exchange between the eastern Surinam caecilians, with notes on Rhinatrema Caribbean and mainland South America. Pp. bivittatum and the description of a new species 207–216 in R. Powell & R. W. Henderson, of Microcaecilia (Amphibia, Gymnophiona). eds., Contributions to West Indian herpetol- Zoologische Mededelingen 54(14):217–235. ogy: a tribute to Albert Schwartz. Society for O’Shaughnessy, A. W. E. 1881. An account of the the Study of Amphibians and Reptiles, collection of lizards made by Mr. Buckley in Contributions to Herpetology, vol. 12, 457 pp. Ecuador, and now in the British Museum, VOLUME 125, NUMBER 4 567

with descriptions of the new species. Proceed- Pellegrino, K. C. M., M. T. Rodrigues, D. J. Harris, ings of the Zoological Society of London Y. Yonenaga-Yassuda, & J. W. Sites, Jr. 2011. 49:227–245. Molecular phylogeny, biogeography and in- O’Shea, M. 1989. The herpetofauna of Ilha de sights into the origin of parthenogenesis in the Maraca´ , state of Roraima, northern Brazil. neotropical genus Leposoma (Squamata: Pp. 51–72 in J. Coote, ed., Reptiles: Proceed- Gymnophthalmidae): ancient links between ings of the 1988 U. K. Herpetological the Atlantic Forest and Amazonia. Molecular Societies, Symposium on Captive Breeding. Phylogenetics and Evolution 61:446–459. The British Herpetological Society, London, Peloso, P. L. V., & T. C. S. Avila-Pires. 2010. 108 pp. Morphological variation in Ptychoglossus Orejas-Miranda, B. R. 1967. El ge´nero ‘‘Leptoty- brevifrontalis Boulenger, 1912 and the status phlops’’ en la region Amazoˆnica. Atas do of Ptychoglossus nicefori (Loveridge, 1929) Simpo´sio so˜bre a Biota Amazoˆnica 5:421–442. (Squamata, Gymnophthalmidae). Herpeto- Parker, H. W. 1927a. A revision of the frogs of the logica 66:357–372. genera Pseudopaludicola, Physalaemus, and Peters, J. A. 1960. The snakes of the subfamily Pleurodema. The Annals and Magazine of Dipsadinae. Miscellaneous Publications, Mu- Natural History, Series 9, 20:450–478. seum of Zoology, University of Michigan Parker, H. W. 1927b. The brevicipitid frogs allied to 114:1–224. the genus Gastrophryne. Occasional Papers of Peters, J. A., & R. Donoso-Barros. 1970. Catalogue the Museum of Zoology, University of Mich- of the neotropical Squamata, Part II. Lizards igan 187:1–6. and amphisbaenians. United States National Parker, H. W. 1935. The frogs, lizards, and snakes of Museum Bulletin 297, Smithsonian Institution British Guiana. Proceedings of the Zoological Press, Washington, D.C., 293 pp. Society of London 105(3):505–530. Peters, J. A., & B. Orejas-Miranda. 1970. Catalogue of the neotropical Squamata: Part I. Snakes. Parker, H. W. 1940. Undescribed anatomical struc- United States National Museum Bulletin 297, tures and new species of reptiles and amphib- Smithsonian Institution Press, Washington, ians. The Annals and Magazine of Natural D.C., 347 pp. History, Series 11, 5:257–274. Peters, W. C. H. 1857. Vier neue amerikanische Passos, P., & R. Fernandes. 2008. Revision of the Schlangen aus der Familie der Typhlopinen Epicrates cenchria complex (Serpentes: Boi- vor und machte daru¨ber einige vorla¨ufige dae). Herpetological Monographs 22:1–30. Mittheilungen. Monatsbericht der Ko¨nig- Pearse, D. E., A. D. Arndt, N. Valenzuela, B. A. lichen Preussischen Akademie der Wissen- Miller, V. Cantarelli, & J. W. Sites, Jr. 2006. schaften zu Berlin 1857:402–403. Estimating population structure under non- Peters, W. C. H. 1863a. U¨ ber Cercosaura und die mit equilibrium conditions in a conservation dieser Gattung verwandten Eidechsen aus context: continent-wide population genetics Su¨damerica. Abhandlungen der Ko¨niglichen of the giant Amazon river [sic] turtle, Podoc- Akademie der Wissenschaften zu Berlin nemis expansa (Chelonia; Podocnemididae). 1862:165–225. Molecular Ecology 15:985–1006. Peters, W. C. H. 1863b [1864]. U¨ ber einige neue oder Pellegrino, K. C. M., M. T. Rodrigues, & Y. weniger bekannte Schlangenarten des zoolog- Yonenaga-Yassuda. 1999. Chromosomal evo- ischen Museums zu Berlin. Monatsbericht der lution in the Brazilian lizards of genus Ko¨niglichen Preussischen Akademie der Wis- Leposoma (Squamata, Gymnophthalmidae) senschaften zu Berlin 1863:272–289. from Amazon and Atlantic rain forests: Peters, W. C. H. 1867. U¨ ber Flederthiere (Pteropus banding patterns and FISH of telomeric Gouldii, Rhinolophus Deckenii, Vespertilio sequences. Hereditas 131:15–21. lobipes, Vesperugo Temminckii) und Amphib- Pellegrino, K. C. M., M. T. Rodrigues, Y. Yonena- ien (Hypsilurus Godeffroyi, Lygosoma scuta- ga-Yassuda, & J. W. Sites, Jr. 2001. A tum, Stenostoma narirostre, Onychocephalus molecular perspective on the evolution of unguirostris, Ahaetulla polylepis, Pseudechis microteiid lizards (Squamata, Gymnophthal- scutellatus, Hoplobatrachus Reinhardtii, Hyla midae), and a new classification for the family. coriacea). Monatsberichte der Ko¨niglichen Biological Journal of the Linnean Society Preussischen Akademie der Wissenschaften 74:315–338. zu Berlin 1867:703–712. Pellegrino, K. C. M., M. T. Rodrigues, & Y. Peters, W. C. H. 1869a. Eine Mittheilung u¨ber neue Yonenaga-Yassuda. 2003. Triploid karyotype Gattungen und Arten von Eidechsen. Mon- of Leposoma percarinatum (Squamata, Gym- atsberichte der Ko¨niglichen Preussischen Aka- nophthalmidae). Journal of Herpetology demie der Wissenschaften zu Berlin 1869:57– 37:197–199. 66. 568 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Peters, W. C. H. 1869b. U¨ ber neue Gattungen und Pritchard, P. C. H. 1995. Paleosuchus trigonatus neue oder weniger bekannte Arten von (Schneider’s smooth-fronted caiman). Herpe- Amphibien (Eremias, Dicrodon, Euprepes, tological Review 26(1):43. Lygosoma, Typhlops, Eryx, Rhynchonyx, Ela- Pritchard, P. C. H. 2005. Long-term sea turtle pomorphus, Achalinus, Coronella, Dromicus, nesting cycles in the Guianas. Pp. 66–68 in Xenopholis, Anoplodipsas, Spilotes, Tropidono- M. S. Coyne & R. D. Clark, compilers, tus). Monatsberichte der Ko¨niglichen Preussi- Proceedings of the twenty-first annual sympo- schen Akademie der Wissenschaften zu Berlin sium on sea turtle biology and conservation. 1869:432–445. National Oceanic and Atmospheric Adminis- Peters, W. C. H. 1870 [1871]. U¨ ber Platemys tuber- tration Technical Memorandum NMFS- osa, eine neue Art von Schildkro¨ten aus SEFSC-528, 368 pp. British-Guiana. Monatsberichte der Ko¨niglich Pyburn, W. F. 1975. A new species of microhylid Preussischen Akademie der Wissenschaften zu frog of the genus Synapturanus from south- Berlin 1870:311–313. eastern Colombia. Herpetologica 31(4):439– Peters, W. C. H. 1872 [1873]. Eine Mittheilung U¨ ber 443. eine Sammlung von Batrachiern aus Neu- Pyron, R. A., & J. J. Wiens. 2011. A large-scale Freiburg in Brasilien. Monatsberichte der phylogeny of Amphibia including over 2800 Ko¨niglich Preussischen Akademie der Wissen- species, and a revised classification of extant schaften zu Berlin 1872:680–684. frogs, salamanders, and caecilians. Molecular Phelps, W. H. 1938. The geographical status of the Phylogenetics and Evolution 61:543–583. birds collected at Mount Roraima. Bolet´ınde Pyron, R. A., F. T. Burbrink, G. R. Colli, A. N. la Sociedad Venezolana de Ciencias Naturales Montes de Oca, L. J. Vitt, C. A. Kuczynski, & 36:83–95. J. J. Wiens. 2011. The phylogeny of advanced Pinto, R. R., & R. Fernandes. 2012. A new blind snakes (Colubroidea), with discovery of a new snake species of the genus Tricheilostoma subfamily and comparison of support meth- from Espinhac¸o Range, Brazil and taxonomic ods for likelihood trees. Molecular Phyloge- status of Rena dimidiata (Jan, 1861) (Ser- netics and Evolution 58:329–342. pentes: Epictinae: Leptotyphlopidae). Copeia Raddi, G. 1820. Di alcune specie nuove di rettili, e 2012:37–48. piante Brasiliane. Memorie di Matematica e di Poe, S. 2004. Phylogeny of anoles. Herpetological Fisica della Societa` Italiana delle Scienze Monographs 18:37–89. (Fisica) 18(2):313–349. Pough, F. H., R. M. Andrews, J. E. Cadle, M. L. Rafinesque, C. S. 1814. Fine del prodromo d’erpe- Crump, A. H. Savitzky, & K. D. Wells. 2004. tologia siciliana. Specchio delle Scienze, o, Herpetology, 3rd edition. Pearson, Upper Saddle River, New Jersey, 736 pp. Giornale Enciclopedico di Sicilia 2:102–104. Pramuk, J. B. 2006. Phylogeny of South American Rafinesque, C. S. 1815. Analyse de la nature ou Bufo (Anura: Bufonidae) inferred from com- tableau de l’universe et des corps organise´s. bined evidence. Zoological Journal of the Jean Barravecchia, Palermo, Italy. Linnean Society 146:407–452. Reynolds, R. P. 1990. Phrynops geoffroanus tuber- Pramuk, J. B., T. Robertson, J. W. Sites, Jr., & B. P. osus (Geoffroy’s Sideneck Turtle). Herpeto- Noonan. 2008. Around the world in 10 logical Review 21(2):40. million years: biogeography of the nearly Reynolds, R. P., R. MacCulloch, M. Tamessar, C. cosmopolitan true toads (Anura: Bufonidae). Watson, C. J. Cole, & C. Townsend. 2002. Global Ecology and Biogeography 17:72–83. Preliminary checklist of the herpetofauna of Presch, W. 1973. A review of the tegus, lizard genus Guyana. Compiled by The Biological Diver- Tupinambis (Sauria: Teiidae) from South sity of the Guiana Shield Program (BDG), America. Copeia 1973(4):740–746. Smithsonian Institution, Washington, D.C. Pritchard, P. C. H. 1964. Turtles of British Guiana. http://botany.si.edu/bdg/guyherps.html (ac- Journal of the British Guiana Museum and cessed 27 May 2012). Zoo of the Royal Agricultural and Commer- Ribeiro, L. B., M. F. Kolodiuk, & E. M. X. Freire. cial Society 39:19–32. 2010. Ventral colored patches in Tropidurus Pritchard, P. C. H. 1966. Sea turtles of Shell Beach, semitaeniatus (Squamata, Tropiduridae): sex- British Guiana. Copeia 1966:123–125. ual dimorphism and association with repro- Pritchard, P. C. H. 1969. Sea turtles of the Guianas. ductive cycle. Journal of Herpetology 44:177– Bulletin of the Florida State Museum 182. 13(2):85–140. Rivas, G. A., C. R. Molina, G. N. Ugueto, T. R. Pritchard, P. C. H. 1980. Dermochelys coriacea. Barros, C. L. Barrio-Amoro´s, & P. J. R. Kok. Catalogue of American Amphibians and 2012. Reptiles of Venezuela: an updated and Reptiles 238:1–4. commented checklist. Zootaxa 3211:1–64. VOLUME 125, NUMBER 4 569

Rivero, J. A. 1961. Salientia of Venezuela. Bulletin of na) de Venezuela. Acta Biologica Venezuelica the Museum of Comparative Zoology at 3:287–300. Harvard College 126(1):1–207. Ruibal, R. 1952. Revisionary studies on some South Rivero, J. A. 1968a. A new species of Eleutherodac- American Teiidae. Bulletin of the Museum of tylus (Amphibia, Salientia) from the Guayana Comparative Zoo¨logy at Harvard College Region, Edo. Bolivar, Venezuela. Brevoria, 106(11):477–529. Museum of Comparative Zoology 306:1–11. Rull, V. 2005. Biotic diversification in the Guayana Rivero, J. A. 1968b [1966]. Notes on the genus Highlands: a proposal. Journal of Biogeogra- Cryptobatrachus (Amphibia: Salientia) with phy 32:921–927. the description of a new race and four new Ruthven, A. G. 1916. Description of a new genus species of a new genus of hylid frogs. and species of lizard from British Guiana. Caribbean Journal of Science 6:137–149. Occasional Papers of the Museum of Zoology, Rivero, J. A. 1968c [1967]. A new race of Otophryne University of Michigan 22:1–4. robusta Boulenger (Amphibia, Salientia) from Ruthven, A. G. 1919. The amphibians of the the Chimanta-tepui of Venezuela. Caribbean University of Michigan-Walker Expedition Journal of Science 7:155–158. to British Guiana. Occasional Papers of the Rivero, J. A. 1970. On the origin, endemism and Museum of Zoology, University of Michigan distribution of the genus Stefania Rivero 69:1–14. (Amphibia, Salientia) with a description of a Ruthven, A. G. 1922. A new species of Amphisbaena new species from southeastern Venezuela. from British Guiana. Occasional Papers of the Bolet´ın de la Sociedad Venezolana de Ciencias Museum of Zoology, University of Michigan Naturales 28:456–481. 122:1–2. Rivero, J. A. 1971. Tres nuevos records y una nueva Sabaj Pe´rez, M. H. (ed.). 2012. Standard symbolic especie de anfibios de Venezuela. Caribbean codes for institutional resource collections in Journal of Science 11:1–9. herpetology and ichthyology: an online refer- Rivero, J. A. 1972 [1971]. Notas sobre los anfibios de ence. Version 3.0 (23 February 2012). Acces- Venezuela. I. Sobre los hilidos de la Guayana sible at http://www.asih.org/node/204 Venezolana. Caribbean Journal of Science American Society of Ichthyologists and Her- 11:181–193. petologists, Washington, D.C. (accessed on 19

Rivero-Blanco, C. 1979. The neotropical lizard genus August 2012). Gonatodes Fitzinger (Sauria: Sphaerodactyli- Savage, J. M. 2002. The amphibians and reptiles of nae). Ph.D. dissertation, Texas A&M Univer- Costa Rica: a herpetofauna between two sity, College Station, 224 pp. continents, between two seas. The University Rodrigues, M. T. 1987. Sistema´ tica, ecologia e of Chicago Press, Chicago, 934 pp. zoogeografia dos Tropidurus do grupo torqua- Savage, J. M. 2011. The correct species-group name tus ao sul do Rio Amazonas (Sauria, Iguani- for an Oxyrhopus (Squamata: Dipsadidae) dae). Arquivos de Zoologia 31(3):105–230. variously called Coluber petalarius, C. pethola, Rodr´ıguez, L. O., & W. E. Duellman. 1994. Guide to C. petola,orC. petolarius by early authors. the frogs of the Iquitos Region, Amazonian Proceedings of the Biological Society of Peru. The University of Kansas Natural Washington 124:223–225. History Museum, Special Publication 22:1–80. Savage, J. M., J. A. Campbell, & W. W. Lamar. Roze, J. A. 1958. Los reptiles del Chimanta´ Tepui 2005. On names for neotropical rattlesnakes (Estado Bolivar, Venezuela) colectados por la (Reptilia: Serpentes: Viperidae: Crotalus). expedicio´n bota´ nica del Chicago Natural Herpetological Review 36(4):369–371. History Museum. Acta Biologica Venezuelica Schargel, W. E., G. A. Rivas, R. Makowsky, J. C. 2:299–314. Sen˜aris, M. A. Natera, T. R. Barros, C. R. Roze, J. A. 1961. El ge´nero Atractus (Serpentes: Molina, & C. L. Barrio-Amoro´s. 2010. Phy- Colubridae) en Venezuela. Acta Biolo´gica logenetic systematics of the genus Gonatodes Venezuelica 3(7):103–119. (Squamata: Sphaerodactylidae) in the Guaya- Roze, J. A. 1967. A check list of the New World na region, with description of a new species venomous coral snakes (Elapidae), with de- from Venezuela. Systematics and Biodiversity scriptions of new forms. American Museum 8(3):321–339. Novitates 2287:1–60. Schlegel, H. 1837. Essai sur la physionomie des Roze, J. A. 1989. New species and subspecies of coral serpens. Partie ge´ne´rale: 1–251 þ Partie snakes, genus Micrurus (Elapidae), with notes descriptive: 1–606. M. H. Schonekat, Amster- on type specimens of several species. Ameri- dam. can Museum Novitates 2932:1–15. Schlegel, H. 1837–1844. Abbildungen neuer oder Roze, J. A., & H. Solano. 1963. Resumen de la unvollsta¨ndig bekannter Amphibien, nach der familia Caeciliidae (Amphibia: Gymnophio- Natur oder dem Leben entworfen, herausge- 570 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

geben und mit einem erla¨uternden Texte curavit, vol. 1. Janssonio-Waesbergios, & J. begleitet. Arnz & Comp., Du¨sseldorf, 141 pp. Wetstenium, & Gul. Smith, Amsterdam. Schmidt, K. P. 1939. A new coral snake from British Sen˜aris, J. C., & J. Ayarzagu¨ena. 2001. Una nueva Guiana. Zoological Series of Field Museum of especie de rana de cristal del ge´nero Hyalino- Natural History, Chicago 24(6):45–47. batrachium (Anura: Centrolenidae) del Delta Schmidt, K. P. 1952. The Surinam coral snake del R´ıo Orinoco, Venezuela. Revista de Micrurus surinamensis. Fieldiana: Zoology Biolog´ıa Tropical 49(3–4):1083–1093. 34(4):25–34. Sen˜aris, J. C., & R. MacCulloch. 2005. Amphibians. Schmidt, K. P. 1953. A check list of North American Pp. 9–23 in T. Hollowell & R. P. Reynolds, amphibians and reptiles. Sixth edition. Amer- eds., Checklist of the terrestrial vertebrates of ican Society of Ichthyologists and Herpetolo- the Guiana Shield. Bulletin of the Biological gists, Chicago, 280 pp. Society of Washington 13:1–98. Schmidt, K. P., & W. F. Walker, Jr. 1943. Peruvian Sen˜aris, J. C., J. Ayarzagu¨ena, & S. Gorzula. 1997 snakes from the University of Arequipa. [1996]. Revisio´n taxono´mica del ge´nero Stefa- Zoological Series of Field Museum of Natural nia (Anura; Hylidae) en Venezuela con la History 24(26):279–296. descripcio´n de cinco nuevas especies. Pub- Schneider, J. G. 1783. Allgemeine Naturgeschichte licacio´nes de la Asociacio´n de Amigos de der Schildkro¨ten, nebst einem systemischen Don˜ana 7:1–57. Verzeichnisse der einzelnen Arten und zwey Sen˜aris, J. C., C. DoNacscimiento, & O. Villarreal. Kupfern. Johan Gotfried Mu¨ller, Leipzig, 364 2005. A new species of the genus Oreophry- pp. nella (Anura; Bufonidae) from the Guiana Schneider, J. G. 1792 [1791]. Beschreibung und Highlands. Pape´is Avulsos de Zoologia, Sa˜o Abbildung einer neuen Art von Wasser- Paulo 45(6):61–67. schildkro¨te nebst Bestimmungen einiger bisher Sentzen, U. J. 1796. Ophiologische Fragmente. wenig bekannten fremden Arten. Schriften der Zoologisches Archiv herausgegeben von Frie- Berlinischen Gesellschaft naturforschender drich Albrecht Anton Meyer 2:49–74. Freunde 10(3):259–284. Shaffer, H. B., P. Meylan, & M. L. McKnight. 1997. Schneider, J. G. 1799. Historiae amphibiorum Tests of turtle phylogeny: molecular, morpho- naturalis et literariae. Fasciculus primus con- logical, and paleontological approaches. Sys-

tinens ranas, calamitas, bufones, salamandras tematic Biology 46:235–268. et hydros. F. Frommanni, Jena, 264 pp. Shaw, G. 1802. General zoology or systematic Schneider, J. G. 1801. Historiae Amphibiorum natural history, vol. III, Part II, Amphibia. naturalis et literariae. Fasciculus secundus: Printed by Thomas Davison, White-Friars, continens crocodilos, scincos, chamaesauras, London, pp. 313–615. boas, pseudoboas, elapes, angues, amphisbae- Silva, N. J., Jr., & J. W. Sites, Jr. 1995. Patterns of nas et caecilias. Fried. Frommann, Jenae, 374 diversity of neotropical squamate reptile pp. species with emphasis on the Brazilian Ama- Schulte, J. A., II, J. P. Valladares, & A. Larson. zon and the conservation potential of indig- 2003. Phylogenetic relationships within Igua- enous reserves. Conservation Biology 9:873– nidae inferred using molecular and morpho- 901. logical data and a phylogenetic taxonomy of Silverstone, P. A. 1975. A revision of the poison- iguanian lizards. Herpetologica 59(3):399– arrow frogs of the genus Dendrobates Wagler. 419. Natural History Museum of Los Angeles Schweigger, A. F. 1812. Prodromus monographiae County Science Bulletin 21:1–55. Cheloniorum. Ko¨nigsberger Archiv fu¨r Na- Silverstone, P. A. 1976. A revision of the poison- turwissenschaft und Mathematik 1:271–458. arrow frogs of the genus Phyllobates Bibron in Scopoli, J. A. 1788. Deliciae florae et faunae Sagra (Family Dendrobatidae). Natural His- Insubricae seu novae, aut minus cognitae tory Museum of Los Angeles County Science species plantarum et animalium quas in Bulletin 27:1–53. Insubria Austriaca tam spontaneas, quam Simmons, N. B., & R. S. Voss. 1998. The mammals exoticas vidit, descripsit, et aeri incidi curavit of Paracou, French Guiana: a Neotropical Joannes Antonius Scopoli. Pars III. Monas- lowland rainforest fauna. Part 1. Bats. Bulle- terii S. Salvatoris, Ticini, 87 pp. tin of the American Museum of Natural Seba, A. 1734. Locupletissimi rerum naturalium History 237:1–219. thesauri accurata descriptio, et iconibus arti- Simpson, G. G. 1960. Notes on the measurement of ficiosissimus expressio, per universam physices faunal resemblance. American Journal of historiam. Opus, cui, in hoc rerum genere, Science, Bradley Volume, 258-A:300–311. nullum par exstitit. Ex toto terrarum orbe Slowinski, J. B. 1995. A phylogenetic analysis of the collegit, difessit, descripsit, et depingendum New World coral snakes (Elapidae: Leptomi- VOLUME 125, NUMBER 4 571

crurus, Micruroides, and Micrurus) based on evidence from mitochondrial DNA sequences. allozymic and morphological characters. Herpetologica 55:254–270. Journal of Herpetology 29:325–338. Taylor, E. H. 1939. Two new species of snakes of the Smith, E. N., & B. P. Noonan. 2001. A new species genus Anomalepis Jan, with a proposal of a of Osteocephalus (Anura: Hylidae) from new family of snakes. Proceedings of the New Guyana. Revista de Biolog´ıaTropical England Zoological Club 17:87–96. 49:347–357. Taylor, E. H. 1968. The caecilians of the world: a Smith, H. M., & R. F. Laurent. 1950. Further notes taxonomic review. University of Kansas Press, upon the enigmatical Bufo nasicus Werner. Lawrence, 848 pp. Bulletin du Muse´e Royal d’Histoire Naturelle Taylor, E. H. 1969. Miscellaneous notes and de Belgique 26(21):1–3. descriptions of new forms of caecilians. The Smith, H. M., & E. H. Taylor. 1950a. Type localties University of Kansas Science Bulletin of Mexican reptiles and amphibiens. The 48(9):281–296. University of Kansas Science Bulletin Thomas, R., 1976. Systematics of Antillean snakes of 33(2):313–380. the genus Typhlops (Serpentes: Typhlopidae). Smith, H. M., & E. H. Taylor. 1950b. An annotated Ph.D. dissertation, Louisiana State Universi- checklist and key to the reptiles of Mexico ty, Baton Rouge, 288 pp. exclusive of the snakes. Smithsonian Institu- Townsend, T. M., A. Larson, E. Louis, & J. R. tion, United States National Museum Bulletin Macey. 2004. Molecular phylogenetics of 199:1–253. Squamata: the position of snakes, amphisbae- Smithe, F. B. 1975. Naturalist’s color guide. Amer- nians, and dibamids, and the root of the ican Museum of Natural History, New York. squamate tree. Systematic Biology 53(5):735– Spix, J. B. 1824. Animalia nova sive Species novae 757. Testudinum et Ranarum, quas in itinere per Townsend, T. M., D. G. Mulcahy, B. P. Noonan, J. Brasiliam annis MDCCCXVII–MDCCCXX W. Sites, Jr., C. A. Kuczynski, J. J. Wiens, & jussu et auspiciis Maximiliani Josephi I. T. W. Reeder. 2011. Phylogeny of iguanian Bavariae Regis. F. S. Hu¨bschmann, Monachii, lizards inferred from 29 nuclear loci, and a Vol. 3:1–53. comparison of concatenated and species-tree

Spix, J. B. 1825. Animalia nova sive Species novae approaches for an ancient, rapid radiation. Lacertarum, quas in itinere per Brasiliam Molecular Phylogenetics and Evolution annis MDCCCXVII–MDCCCXX jussu et 61:363–380. auspiciis Maximiliani Josephi I. Bavariae The Code [International Commission of Zoological Regis. F. S. Hu¨bschmann, Monachii, Vol. Nomenclature]. 1999. International code of 1:1–26. zoological nomenclature, 4th Edition. Inter- Starace, F. 1998. Guide des serpents et amphisbe`nes national Trust for Zoological Nomenclature, de Guyane franc¸aise. Ibis Rouge Editions, London, 306 pp. Guadeloupe, Guyane, 450 pp. Troschel, F. H. 1848. Amphibien. Pp. 645–661 in R. Steindachner, F. 1862. U¨ ber Zwei noch unbeschrie- Schomburgk, ed., Reisen in Britisch-Guiana bene Batrachier aus den Sammlungen des K. in den Jahren 1840–1844. Im auftrag Sr. K. Zoologischen. Archivio per la Zoologia Ma¨jestat des Ko¨nigs von Preussen. Dritter l’Anatomia e la Fisiologia, Genova, II(1):77– Theil. Versuch einer Fauna und Flora von 82. Britisch-Guiana. Verlagbuschhandlung von J. Steindachner, F. 1864. Batrachologische Mittheilun- J. Weber, Leipzig, 1260 pp. gen. Verhandlungen der kaiserlich-ko¨nig- Trueb, L., & D. C. Cannatella. 1986. Systematics, lichen zoologisch-botanischen Gesellschaft in morphology, and phylogeny of genus Pipa Wien 14:239–288. (Anura: Pipidae). Herpetologica 42:412–449. Stephens, L., & M. A. Traylor, Jr. 1985. Ornitho- Trueb, L., & D. Massemin. 2001. The osteology and logical gazetteer of the Guianas. Harvard relationships of Pipa aspera (Amphibia: Anu- University, Cambridge, Massachusetts, 121 ra: Pipidae), with notes on its natural history pp. in French Guiana. Amphibia-Reptilia Stuart, L. C. 1941. Studies of Neotropical Colubri- 22(1):33–54. nae. VIII. A revision of the genus Dryadophis Tschudi, J. J. 1845. Reptilium conspectus quae in Stuart, 1939. Miscellaneous Publications Mu- Republica Peruana reperiuntur et pleraque seum of Zoology, University of Michigan observata vel collecta sunt in itinere. Archiv 49:1–106. fu¨r Naturgeschichte, Berlin 11(1):150–170. Summers, K., L. A. Weigt, P. Boag, & E. Berming- Underwood, G. 1964. Reptiles of the Eastern ham. 1999. The evolution of female parental Caribbean, 1st supplement June 1964. Carib- care in poison frogs of the genus Dendrobates: bean Affairs, new ser., 4 pp. 572 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Uzzell, T., & J. C. Barry. 1971. Leposoma percar- Vidal, N., & S. B. Hedges. 2005. The phylogeny of inatum, a unisexual species related to L. squamate reptiles (lizards, snakes, and am- guianense;andLeposoma ioanna,anew phisbaenians) inferred from nine nuclear species from Pacific coastal Colombia (Sauria, protein-coding genes. Comptes Rendus Biol- Teiidae). Postilla, Peabody Museum, Yale ogies 328:1000–1008. University 154:1–39. Vidal, N., J.-C. de Massary, & C. Marty. 1999 Van der Meijden, A., M. Vences, S. Hoegg, R. [1998]. Nouvelles espe`ces de serpents pour la Boistel, A. Channing, & A. Meyer. 2007. Guyane franc¸aise. Revue franc¸aise d’Aqua- Nuclear gene phylogeny of narrow-mouthed riologie 25(3–4):131–134. toads (Family: Microhylidae) and a discussion Vidal, N., A.-S. Delmas, P. David, C. Cruaud, A. of competing hypotheses concerning their Couloux, & S. B. Hedges. 2007. The phylog- biogeographical origins. Molecular Phyloge- eny and classification of caenophidian snakes netics and Evolution 44:1017–1030. inferred from seven nuclear protein-coding Van Lidth de Jeude, T. W. 1904. Note II. Reptiles genes. Comptes Rendus Biologies 330:182– and batrachians from Surinam. Notes from 187. the Leyden Museum 25:83–94. Vidal, N., M. Dewynter, & D. J. Gower. 2010a. Vandelli, D. 1761. Epistola de Holothurio, et Dissecting the major American snake radia- Testudine Coriacea ad celeberrimum Carolum tion: a molecular phylogeny of the Dipsadidae Linnaeum Equitem Naturae curiosorum dio- Bonaparte (Serpentes, Caenophidia). Comptes scoridem II. Conzatti, Patavii (Padua), 12 pp. Rendus Biologies 333(1):48–55. Vanzolini, P. E. 1972. Typhlops brongersmai spec. Vidal, N., J. Marin, M. Morini, S. Donnellan, W. R. nov. from the coast of Bahia, Brazil (Ser- Branch, R. Thomas, M. Vences, A. Wynn, C. pentes, Typhlopidae). Zoologische Medede- Cruaud, & S. B. Hedges. 2010b. Blindsnake lingen 47:27–29. evolutionary tree reveals long history on Vanzolini, P. E. 1976. Typhlops brongersmianus,a Gondwana. Biology Letters 6(4):558–561. new name for Typhlops brongersmai Vanzoli- Vockenhuber, E. A., W. Ho¨dl, & A. Ame´zquita. ni, 1972, preoccupied (Serpentes, Typhlopi- 2009. Glassy fathers do matter: egg atten- dae). Pape´is Avulsos de Zoologia, Sa˜o Paulo dance enhances embryonic survivorship in the 29:247. glass frog Hyalinobatrachium valerioi. Journal

Vanzolini, P. E. 1980. Coleodactylus septentrionalis, of Herpetology 43:340–344. sp. n., with notes on the distribution of the Voss, R. S., D. P. Lunde, & N. B. Simmons. 2001. genus (Sauria, Gekkonidae). Pape´is Avulsos The mammals of Paracou, French Guiana: a de Zoologia, Sa˜o Paulo 34:1–9. Neotropical lowland rainforest fauna. Part 2. Vanzolini, P. E. 1981. The scientific and political Nonvolant species. Bulletin of the American contexts of the Bavarian Expedition to Brasil. Museum of Natural History 263:1–236. Pp. ix–xxix in J. B. von Spix & J. G. Wagler, Wagler, J. 1824. Serpentum brasiliensium species eds., Herpetology of Brazil. Facsimile Re- novae ou histoire naturelle des espe`ces nou- prints in Herpetology, Society for the Study of velles de serpens, recueillies et observe´es Amphibians and Reptiles, 400 pp. pendant le voyage dans l’inte´rieur du Bre´sil Vanzolini, P. E. 1994. On the distribution of certain dans les anne´es 1817, 1818, 1819, 1820, South American turtles (Testudines: Testudi- exe´cute´ par ordre de sa majeste´ le roi de nidae & Chelidae). Smithsonian Herpetologi- Bavie`re, publie´e par Jean de Spix. Franc. cal Information Service 97:1–10. Seraph. Hu¨bschmanni, Monachii, 75 pp. Vanzolini, P. E. 2002. A second note on the Wagler, J. 1828. Auszu¨ge aus seinem Systema geographical differentiation of Amphisbaena Amphibiorum. Isis von Oken 21:740–744. fuliginosa L., 1758 (Squamata, Amphisbaeni- Wagler, J. 1830. Natu¨rliches System der Amphibien, dae), with a consideration of the forest refuge mit vorangehender Classification der Sa¨ug- model of speciation. Anais da Academia thiere und Vo¨gel. Ein Beitrag zur vergleichen- Brasileira de Cieˆncias 74(4):609–648. den Zoologie. J. G. Cotta, Mu¨nchen, 354 pp. Vanzolini, P. E., & Carvalho, C. M. de. 1991. Two Wake, M. H., & M. A. Donnelly. 2010. A new sibling and sympatric species of Gymnoph- lungless caecilian (Amphibia: Gymnophiona) thalmus in Roraima, Brasil (Sauria, Teiidae). from Guyana. Proceedings of the Royal Pape´is Avulsos de Zoologia, Sa˜oPaulo Society B 277:915–922. 37(12):173–226. Warren, A. 1973. Roraima: report of the 1971 British Vanzolini, P. E., & E. E. Williams. 1970. South expedition to Mount Roraima in Guyana, American anoles: the geographic differentia- South America. Seacourt Press, Oxford, 152 tion and evolution of the Anolis chrysolepis pp. species group (Sauria, Iguanidae). Arquivos Werner, F. 1896. Beitra¨ge zur Kenntniss der Repti- de Zoologia, Sa˜o Paulo 19(1–2):1–124. lien und Batrachier von Centralamerika und VOLUME 125, NUMBER 4 573

Chile, sowie einiger seltenerer Schlangenarten. scientific meetings of the Zoological Society of Verhandlungen der kaiserlich-ko¨niglichen zo- London 1861:325–326. ologisch-botanischen Gesellschaft in Wien Wynn, A., & W. R. Heyer. 2001. Do geographically 46:344–365. widespread species of tropical amphibians Werner, F. 1903. Neue Reptilien und Batrachier aus exist? An estimate of genetic relatedness dem naturhistorischen Museum in Bru¨ssel. within the neotropical frog Leptodactylus Nebst Bemerkungen u¨ber einige andere Arten. fuscus (Schneider 1799) (Anura Leptodactyli- Zoologischer Anzeiger 26(693):246–253. dae). Tropical Zoology 14:255–285. Wied-Neuwied, M. P. 1821. Reise nach Brasilien in Yanek, K., W. R. Heyer, & R. O. de Sa´ . 2006. den Jahren 1815 bis 1817. Zweiter Band. H. L. Genetic resolution of the enigmatic Lesser Bro¨nner, Frankfurt am Main, 345 pp. Antillean distribution of the frog Leptodacty- Wied-Neuwied, M. P. 1824. Verzeichniss der Am- lus validus (Anura, Leptodactylidae). South phibien, welche im zweiten Bande der Natur- American Journal of Herpetology 1:192–201. geschichte Brasiliens von Prinz Max von Ya´ nez-Mun˜oz, M., P. Pe´rez-Pen˜a, & D. Cisneros- Neuwied werden beschrieben werden. Isis Heredia. 2009. New country records of von Oken 1824(6):661–673. Hyalinobatrachium iaspidiense (Amphibia, Wiegmann, A. F. A. 1856. Pp. 6 in H. Lichtenstein, Anura, Centrolenidae) from the Amazonian Nomenclator reptilium et amphibiorum Mu- lowlands of Ecuador and Peru. Herpetology sei zoologici berolinensis. Namenverzeichniss Notes 2:49–52. der in der zoologischen Sammlung der Yonenaga-Yassuda, Y., P. E. Vanzolini, M. T. Ko¨niglichen Universita¨t zu Berlin aufgestell- Rodrigues, & C. M. de Carvalho. 1995. ten Arten von Reptilien und Amphibien nach Chromosome banding patterns in the unisex- ihren Ordnungen, Familien und Gattungen. ual microteiid Gymnophthalmus underwoodi Buchdruckerei der Ko¨niglichen Akademie der and in two related sibling species (Gymnoph- Wissenschaften, Berlin, 48 pp. thalmidae, Sauria). Cytogenetics and Cell Wiens, J. J., C. A. Kuczynski, T. Townsend, T. W. Genetics 70:29–34. Reeder, D. G. Mulcahy, & J. W. Sites, Jr. Zaher, H., & U. Caramaschi. 1992. Sur le statut 2010. Combining phylogenomics and fossils in taxinomique d’Oxyrhopus trigeminus et O. higher-level squamate reptile phylogeny: mo- guibei (Serpentes, Xenodontinae). Bulletin du

lecular data change the placement of fossil Muse´um national d’Histoire naturelle, Section taxa. Systematic Biology 59(6):674–688. A, 14:805–827. Wilkinson, M., & P. J. R. Kok. 2010. A new species of Zaher, H., & A. L. C. Prudente. 1999. Intraspecific Microcaecilia (Amphibia: Gymnophiona: Cae- variation of the hemipenis in Siphlophis and ciliidae) from Guyana. Zootaxa 2719:35–40. Tripanurgos. Journal of Herpetology 33:698–702. Wilkinson, M., R. Nussbaum, & M. Hoogmoed. Zaher, H., F. G. Grazziotin, J. E. Cadle, R. W. 2009. A new species of Microcaecilia (Am- Murphy, J. C. de Moura-Leite, & S. L. phibia: Gymnophona [sic]: Caeciliidae) from Bonatto. 2009. Molecular phylogeny of ad- Suriname. Herpetologica 65(4):413–418. vanced snakes (Serpentes, Caenophidia) with Wilkinson, M., D. San Mauro, E. Sherratt, & D. J. an emphasis on South American Xenodon- Gower. 2011. A nine-family classification of tines: a revised classification and descriptions caecilians (Amphibia: Gymnophiona). Zoo- of new taxa. Pape´is Avulsos de Zoologia, Sa˜o taxa 2874:41–64. Paulo 49(11):115–153. Wilson, L. D. 1999. Checklist and key to the species Zamudio, K. R., & H. W. Greene. 1997. Phylogeog- of the genus Tantilla (Serpentes: Colubridae), raphy of the bushmaster (Lachesis muta: with some commentary on distribution. Viperidae): implications for neotropical bio- Smithsonian Herpetological Information Ser- geography, systematics, and conservation. vice 122:1–34. Biological Journal of the Linnean Society Wollenberg, K. C., M. Veith, B. P. Noonan, & S. 62:421–442. Lo¨tters. 2006. Polymorphism versus species Zug, G. R., C. H. Ernst, & R. V. Wilson. 1998. richness—systematics of large Dendrobates Lepidochelys olivacea. Catalogue of American from the eastern Guiana Shield (Amphibia: Amphibians and Reptiles 653:1–13. Dendrobatidae). Copeia 2006(4):623–629. Wucherer, O. 1861. Description of a new species of Elapomorphus from Brazil. Proceedings of the Associate Editor: W. Ronald Heyer. 574 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 (481905200 N, 5884705800 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 (58370N, 578530W), 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 VOLUME 125, NUMBER 4 575 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 576 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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 VOLUME 125, NUMBER 4 577

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 578 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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