Amy Lathrop

The dazed and confused identity of Agassiz’s land tortoise, Gopherus agassizii (Testudines, Testudinidae) with the description of a new species, and its consequences for conservation

Abstract We investigate a cornucopia of problems associated with the identity of the desert tortoise, Gopherus agassizii (Cooper). The date of publication is found to be 1861, rather than 1863. Only one of the three original cotypes exists, and it is designated as the lectotype of the species. Another cotype is found to have been destroyed in the 1906 San Francisco earthquake and subsequent fire. The third is lost. The lectotype is genetically confirmed to be from California, and not Arizona, USA as sometimes reported. Maternally, the holotype of Gopherus lepidocephalus (Ottley & Velázques Solis. 1989) from the Cape Region of Baja California Sur, Mexico is also from the Mojavian population of the desert tortoise, and not from Tiburon Island, Sonora, Mexico as previously proposed. A suite of characters serve to diagnose tortoises west and north of the Colorado River, the Mojavian population, from those east and south of the river in Arizona, USA, and Sonora and Sinaloa, Mexico, the Sonoran population. Species recognition is warranted and because Gopherus lepidocephalus is from the Mojavian population, no names are available for the Sonoran species. Thus, a new species, Gopherus morafkai sp. n., is named and this action reduces the distribution of Gopherus agassizii to only 30% of its former range. This reduction has important implications for the conservation and protection of Gopherus agassizii, which may deserve a higher level of protection.

Cryptic Species of a Cascade Frog from Southeast Asia: Taxonomic Revisions and Descriptions of Six New Species

Abstract The Southeast Asian cascade frog, Rana livida (Blyth, 1856), has long been suspected to be a complex of species. Several different forms are described from across its vast range. The loss of type material and disparate sampling efforts are challenges. Is variability in this species due to geographic variation or to the presence of multiple species? We use concordant evidence from morphology, morphometrics, cellular DNA content, and allozyme electrophoresis to investigate diversity in R. livida from Vietnam. Three distinct species are recognized on the basis of morphology, as are four other suspect groups (morphotypes). Discriminant function analyses of morphometric data detect patterns of morphological variation among all seven groups. Pairwise comparison of cellular DNA content using t-tests shows significant differences among sympatric morphotypes, suggesting they represent distinct species. This hypothesis is supported by an analysis of 14 allozymic loci, in which fixed allelic differences are found among specimens in sympatry and allopatry. Examination of available type material of four junior synonyms of R. livida results in their recognition as species. One of these species, R. chloronota, is a wide-ranging species erroneously referred to as R. livida. Seven species occur in Vietnam. We describe six new cryptic species belonging to the Rana chloronota complex, redescribe R. chloronota, R. livida, R. sinica, and R. graminea, and give comments on R. leporipes. Three of these new species (R. bacboensis, new species, R. hmongorum, new species, and R. daorum, new species) occur in montane forests in northern Vietnam, and two (R. banaorum, new species and R. morafkai, new species) are known only from the Tay Nguyen Plateau of Vietnams Central Highlands. One species, R. megatympanum, new species, occurs in portions of both northern Vietnam and the Central Highlands. An identification key for the Rana chloronota complex from Vietnam is provided. The finding of six cryptic species within a small portion of the geographic region of R. chloronota suggests that many more cascade ranids await discovery. This documentation has serious implications for conservation; each of the new species occurs in sympatry with at least one other member of the complex. Consequently, far more species are being affected by habitat loss than was previously thought.

EXCEPTIONAL DIVERSITY OF STEFANIA (ANURA: HYLIDAE) ON MOUNT AYANGANNA, GUYANA: THREE NEW SPECIES AND NEW DISTRIBUTION RECORDS

Herpetological survey on Mt. Ayanganna in west-central Guyana resulted in the discovery of five species of Stefania occurring syntopically at about 1500 m. Two additional species occur at the base of the mountain. We describe three new species of Stefania and provide redescriptions and range extensions for four other Stefania. The number of Stefania known from Guyana increases from three to seven species. Resumen Cinco nuevas especies de Stefania han sido descubiertas tras un reconocimiento herpetológico llevado a cabo en el Monte Ayanganna en la Guyana centro occidental. Las cinco especies han sido halladas en torno a los 1500 m de altitud. Dos especies mas han sido halladas en la base de la montaña. Se describen aquí tres especies nuevas de Stefania halladas. Para las restantes cuatro especies se recoge su redescripción así como el área de distribución. El número de especies conocidas de Stefania aumenta de este modo de tres a siete especies.

The Phylogenetic Relationships of the Chinese and Vietnamese Waterfall frogs of the genus Amolops

Ranid frogs of the genus Amolops occur in Southeast Asia and are typically found near waterfalls. Their phylogenetic relationships have not been resolved. We include 2,213 aligned nucleotide sites of the 12S, 16S and tRNA(val) gene regions of the mitochondrial DNA genome from 43 individuals of Chinese and Vietnamese Amotops, Huia, Hylarana, Meristogenys, Odorrana and Rana. The outgroup species were from the genera Chaparana, Limnonectes, Nanorana, and Paa. The data were analyzed within the framework of a refutationist philosophy using maximum parsimony. Four clades of waterfall frogs were resolved. Meristogenys was not resolved as the sister group to either Huia nor Amolops. The hypothesis Of evolutionary relationships placed Amolops chapaensis and Huia nasica in the genus Odorrana.

Two new species of Leptolalax (Anura: Megophryidae) from northern Vietnam

Two new species of Leptolalax are described from two mountain ranges in northern Vietnam (Song Gam and Tam Dao) that are less than 150 km apart. Currently, only two species of Leptolalax, L. pelodytoides and L. bourreti, are described from Vietnam. The two new species are distinguished from other Leptolalax by a combination of characters including skin texture, ventral color pattern, presence or absence of spots on the flanks, and lateral fringes on the toes. Furthermore, the new species can be distinguished from the widely distributed and sympatric L. pelodytoides by their large size, indistinct color pattern, and an absence of granules under the chin. In addition, one of the new species has a higher cellular DNA content than sympatric L. pelodytoides.

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

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. 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.

Polyandry and multiple paternities in the threatened Agassiz's desert tortoise, Gopherus agassizii

We used data from 17 to 20 microsatellite markers to investigate the incidence of multiple paternities in wild Agassiz’s desert tortoises, Gopherus agassizii. Neonates were sampled from clutches of eggs laid by wild mothers in nesting enclosures at Edwards Air Force Base and at the Marine Corps Air Ground Combat Center, California. We genotyped 28 clutches from 26 females sampling an average of six neonates per clutch. The number of paternal alleles was used to determine the minimum number of sires for each clutch. Based on conservative criteria requiring evidence from at least two loci to determine multiple paternity, a minimum of 64% of females were polyandrous, while a minimum of 57% of clutches were sired by multiple males. This formed one of the highest incidences of multiple paternities recorded to date in any species of tortoise. The high number of microsatellite loci involved in the analyses allowed detection of multiple paternities in clutches where this may have been missed if fewer loci were used. Our results highlighted the potential pitfalls of quantitatively comparing paternity studies based on differing sampling strategies. Finally, we summarized the conservation implications of the high rate of multiple paternities in this threatened species.

Phylogeny of the Cyrtodactylus irregularis species complex (Squamata: Gekkonidae) from Vietnam with the description of two new species

The number of described species of bent-toed geckos of the Cyrtodactylus irregularis species complex in Vietnam has increased from one to eight in the last six years. We combined morphological and molecular analyses to explore phylogenetic relationships among all described species in the group. The phylogeny required the description of two new species, Cyrtodactylus phuocbinhensis sp. nov. and Cyrtodactylus taynguyenensis sp. nov. Further, the tree resolved two additional undescribed clades that may also be new species. The species C. bugiamapensis and C. ziegleri were found to require redefinition. Cyrtodactylus phuocbinhensis sp. nov. is characterized by a series of enlarged femoral scales separated from preanal scales while Cyrtodactylus taynguyenensis sp. nov. does not possess enlarged femoral scales. Both new species are distinguished from other congeners by a combination of the following characters: small subcaudal scales, not transversely enlarged; presence (C. phuocbinhensis sp. nov.) or absence (C. taynguyenensis sp. nov.) of enlarged femoral scales; number of preanal pores; and dorsal pattern. Genetic distances between described species and new species were 16.5% and 2.0% in COI and RPL35, respectively, for C. phuocbinhensis sp. nov., and these distances were 18.8% and 2.2% for C. taynguyenensis sp. nov., respectively.

Multilocus species delimitation in the Crotalus triseriatus species group (Serpentes: Viperidae: Crotalinae), with the description of two new species

Members of the Crotalus triseriatus species group of montane rattlesnakes are widely distributed across the highlands of Mexico and southwestern USA. Although five species are currently recognized within the group, species limits remain to be tested. Genetic studies suggest that species may be paraphyletic and that at least one cryptic species may be present. We generate 3,346 base pairs of DNA sequence data from seven nuclear loci to test competing models of species delimitation in the C. triseriatus group using Bayes factor delimitation. We also examine museum specimens from the Trans-Mexican Volcanic Belt for evidence of cryptic species. We find strong support for a nine-species model and genetic and morphological evidence for recognizing two new species within the group, which we formally describe here. Our results suggest that the current taxonomy of the C. triseriatus species group does not reflect evolutionary history. We suggest several conservative taxonomic changes to the group, but future studies are needed to better clarify relationships among species and examine genetic patterns and structure within wide-ranging lineages.

The genus Oxyrhopus (Serpentes: Dipsadidae: Xenodontinae) in Guyana: morphology, distributions and comments on taxonomy

Existe uma consideravel confusao sobre a taxonomia e distribuicoes de Oxyrhopus no norte da America do Sul. Os especimes e registros de Oxyrhopus da Guyana foram examinados. Oxyrhopus melanogenys, O. occipitalis e O. petola ocorre na Guyana. Os dados de morfologia e coloracao destas especies sao apresentados. As distribuicoes destas especies e caracteres para distinguir essas especies sao discutidos.