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