DOI: 10.3897/vz.76.e182505

Authors: Gerald Mayr

Abstract: Abstract A diverse array of early Eocene zygodactyl birds has been assigned to the Psittacopasseres, the clade including parrots and passerines, but the exact affinities of the fossils are controversially resolved. Here, new analyses are performed based on a revised character matrix. Concerning critical taxa, the results of the primary analysis and that of the analyses constrained to a molecular backbone phylogeny show disparate tree topologies, and probably none correctly reflects the true interrelationships of the fossil taxa. The new taxon Psittacomimidae fam. nov. is introduced for a clade formed by the taxa Psittacomimus and Parapsittacopes, which were before assigned to the Psittacopedidae. The Psittacomimidae fam. nov. are likely to be the sister taxon of the Parapasseres, that is, the clade formed by the Zygodactylidae and Passeriformes, with the Morsoravidae branching next. A clade formed by all or most Halcyornithidae and the Messelasturidae is termed Messelasturiformes. A derived morphology of the proximal tarsometatarsus is reported, which may support a clade including the Vastanavidae and Messelasturiformes. However, although psittacopasserine affinities of the Vastanavidae conform to the overall osteology of these birds, the higher-level affinities of the Messelasturiformes are more elusive.

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DOI: 10.3897/vz.76.e169219

Authors: Diego Omar Di Pietro, Julieta Sánchez, Sebastián Poljak, Leandro Alcalde

Abstract: We analysed the genetic and morphological variation in Philodryas patagoniensis, a widely distributed South American racer snake. Two well-differentiated haplogroups were identified using mitochondrial gene sequences (12S and 16S) and the nuclear gene c-mos. Genetic divergence between these haplogroups correlates strongly with morphological differences, allowing the recognition of two morphotypes within P. patagoniensis. We integrated genetic and morphological data into a total evidence analysis using parsimony. Our results support the distinction between the two haplogroups/morphotypes, consistent with recognising two species within P. patagoniensis. Accordingly, we re-describe P. patagoniensis, refining its morphological variation and geographical distribution to reflect the observed genetic differentiation, and describe a new species. Morphological characteristics can distinguish the two species, including body measurements, scale patterns, and cranial osteology. The new species differs from P. patagoniensis sensu stricto in traits associated with arboreal habits, which are strongly correlated with the distribution of the two taxa across forested and open habitats in South America. Furthermore, P. patagoniensis sensu stricto has a significant Lycosa spider component in its diet, which is absent in the newly described species.

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DOI: 10.3897/vz.76.e166419

Authors: N. Joris Fleck, Alice Petzold, Andolalao Rakotoarison, Miguel Vences, Mark D. Scherz

Abstract: The role of regional diversification versus the effect of migration in generating local species assemblages remains poorly known. Here, we contribute to the understanding of the role of colonisation and in situ diversification by studying an assemblage of miniaturised microhylid frogs of the genus Stumpffia Boettger, 1881, of which six species have been known to occur on Montagne d’Ambre, a volcanic mountain in the north of Madagascar. These six species are distributed over different, partly overlapping elevational levels. We examined this assemblage based on molecular data (16S mtDNA and Rag1 nDNA), new data on the elevational distribution among local Stumpffia species, and differences in advertisement calls. Our results revealed several genetic lineages constituting distinct species, including another species record for the mountain, S. mamitika Rakotoarison et al., 2017, as well as the new candidate species Stumpffia sp. aff. angeluci. This brings the total number of described species known to occur on the mountain to seven, four of which are micro-endemic. Our data indicate that one clade, consisting of four species, has arisen in situ as a microendemic radiation. We discuss alternative evolutionary scenarios for the biogeographic origin of the observed Stumpffia species.

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DOI: 10.3897/vz.76.e154822

Authors: Zijia Hong, M. S. Shahrul Anuar, L. Lee Grismer, Evan S. H. Quah

Abstract: Phylogenetic analyses of newly discovered populations of Hemiphyllodactylus from sky-islands across Peninsular Malaysia using the mitochondrial gene ND2, recovered two new upland species embedded within the harterti group. Hemiphyllodactylus puncak sp. nov. from Langkawi Island and H. jeraiensis sp. nov. from Gunung Jerai are sister species with an uncorrected pairwise sequence divergence of 3.6% between them. Together, they formed the sister lineage to H. cicak from Penang Hill, with an uncorrected pairwise sequence divergence of 3.9–4.8% and 4.5–5.7%, respectively. Given that these three populations occur on mountain tops isolated by lowland habitat and the Straits of Malacca, gene flow between them is highly unlikely, and there are unique combinations of characters that differentiate them from one another, as well as from all other species of the harterti group. The time-calibrated BEAST phylogeny shows that the harterti group diversified across the uplands of Peninsular Malaysia in the Oligocene and Pleistocene, which is consistent with climatic fluctuations during this period. During the Middle Oligocene, the ancestral population of the harterti group diverged into two distinct populations, one in the Banjaran Titiwangsa and another in the Banjaran Timur. These two populations then radiated independently across Peninsular Malaysia, giving rise to at least nine additional species within the harterti group.

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DOI: 10.3897/vz.75.e167366

Authors: Werner Conradie, Chad Keates, Eli Greenbaum, Javier Lobón-Rovira, Krystal A. Tolley, Max Benito, Pedro Vaz Pinto, Reuben V. van Breda, Luke Verburgt

Abstract: Abstract Ichnotropis is a genus of medium-sized lacertids endemic to sub-Saharan Africa, characterised by rough head shields. The genus currently comprises six nominal species distributed across much of southern, central, and eastern Africa. Some species are apparently active at only certain times of the year, resulting in limited specimen collections and severely hampering research. This scarcity of material has historically made comprehensive systematic reviews of the genus difficult and has led to the description of numerous regional morphological variants as distinct species or subspecies. Material collected in recent years has enabled us to provide a new phylogenetic hypothesis of Ichnotropis using two mitochondrial genes (16S and ND4) and two nuclear genes (c-mos and RAG-1). Our phylogenetic dataset includes 56 individuals representing five of the six currently recognised species (excluding I. chapini). Additionally, the broad geographical sampling of the widespread I. capensis group has allowed us to explore the taxonomic status of several species and subspecies within the group. As a result, we demonstrate the monophyly of Ichnotropis in relation to other African lacertids and present the most comprehensive phylogeny of the genus to date. We also provide the first phylogenetic placements for I. tanganicana and I. grandiceps, which allows us to validate their taxonomic statuses. Furthermore, we recovered a new cryptic species closely related to I. grandiceps, and identified several well-supported clades within the I. capensis group, all corroborated by multi-locus species delimitation analyses. One of these clades is described herein as a new species, while the remaining taxa of interest are discussed and highlighted for future investigation. Based on our findings, we recommend the following taxonomic revisions: Ichnotropis longipes and I. macrolepidota should remain synonyms of I. capensis; I. bivittata pallida and I. capensis nigrescens are treated as a junior synonyms of I. bivittata; and I. overlaeti is considered a junior synonym of I. tanganicana. Although we could not determine the phylogenetic placement of I. chapini due to the lack of genetic material, its head morphology and scalation support its reassignment to the I. bivittata group. Thus, it is retained as a valid species pending the availability of new material for further taxonomic actions. In conclusion, this study resolves several long-standing taxonomic issues within one of Africa’s most understudied lacertid genera and lays a solid foundation for future research on the genus Ichnotropis.

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DOI: 10.3897/vz.75.e148133

Authors: Bitupan Boruah, V. Deepak, Abhijit Das

Abstract: Abstract Bush frogs currently in the genera Raorchestes and Philautus are poorly documented from northeast India when compared to the Western Ghats of southwestern India. They are morphologically cryptic, but variable in their acoustic repertoire and genetic divergence. We present a long-overdue revision of the bush frogs of northeast India by sampling 81 localities in eight Indian States (including nine type localities of known species) and by comparing historical (especially, type) material. Using an integrative approach that combines levels of divergence in mitochondrial DNA, a nuclear-encoded gene and comparing morphological and bioacoustic data, we demonstrate the existence of 13 new species, which are formally described here. In addition, based on the evidence from molecular and morphological data we synonymise four previously described species from this region. We resolve the taxonomic identities of three other frog species found in northeast India. After extensive survey across northeast India, we did not find a single specimen of the genus Philautus. The only species in this genus with ambiguous generic identity is P. dubius which requires further taxonomic investigation.

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DOI: 10.3897/vz.75.e162650

Authors: Amirtha Balan, Abhijit Das, Bitupan Boruah, Frank Tillack, Samuel Lalronunga, Veerappan Deepak

Abstract: Abstract The genus Ptyctolaemus Peters, 1864 is currently represented by three species; P. gularis, P. collicristatus and P. chindwinensis. Previous molecular phylogenetic studies have highlighted additional lineages within P. gularis. Phylogenetic analysis using the ND2 mitochondrial gene recovered three distinct lineages in India. Multivariate analyses using morphological data placed the type specimen of P. gularis (ZMB 5004) with the samples collected from Meghalaya. We assign this lineage as P. gularis and describe the other two lineages as new species. Ptyctolaemus siangensis sp. nov. found west of Siang River in Arunachal Pradesh and Ptyctolaemus namdaphaensis sp. nov. found in Namdapha, Changlang district of Arunachal Pradesh. Although phenotypically similar the two new species differ from its congeners in gular colouration and subtle morphological differences.

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DOI: 10.3897/vz.75.e155717

Authors: Rajeev Raghavan, Remya L. Sundar, C.P. Arjun, Arya Sidharthan, Nithinraj Panangattu Dharmarajan, Appukuttannair Biju Kumar, Siby Philip, Anvar Ali, Mandar S. Paingankar, Glavin Thomas Rodrigues, Ralf Britz, Neelesh Dahanukar

Abstract: Abstract Subterranean biodiversity continues to be poorly known as a result of uncertainties, challenges and hazards associated with sampling in microhabitats such as aquifers and caves. Focusing on the narrow, lateritic aquifers and associated groundwater habitats in the Western Ghats freshwater ecoregion (southern peninsular India), we investigate the genetic diversity of an enigmatic group of eel-like fishes (family Synbranchidae). A maximum-likelihood phylogenetic analysis based on mitochondrial cytochrome oxidase subunit 1 (cox1) gene sequences recovered these synbranchid eels into two distinct clades comprising genera Ophichthys (fossorial eels ‘with eyes’) and Rakthamichthys (aquifer-dwelling ‘blind’ eels). Additionally, three species-delimitation approaches (based on the mitochondrial cox1 gene), revealed the presence of 11 Evolutionarily Distinct Lineages (EDLs) within Rakthamichthys separated by an inter-lineage divergence between 5.8–20.3%, and an intra-lineage divergence between 0–4.5%. Rakthamichthys in southern peninsular India exhibited a distribution pattern comprising both restricted-range and wide-ranging lineages. Fossorial eels of the genus Ophichthys, on the other hand, are widely distributed in southern peninsular India, with clear geographical boundaries separating the two known species. The genetic network of Rakthamichthys and Ophichthys revealed multiple haplotypes within various EDLs, with a large number of mutations separating the haplotypes within, and between species and/or lineages. Though represented by high levels of genetic divergence revealing the potential existence of at least 11 EDLs, their remarkable morphological uniformity combined with a complex distribution pattern makes it difficult to assign known species names to various Rakthamichthys lineages. Most subterranean habitats in southern peninsular India are under severe anthropogenic threats. Therefore, resolving the taxonomy of, and developing conservation actions for groundwater-dependent species is a priority, for which we suggest future steps.

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DOI: 10.3897/vz.75.e146618

Authors: Sergio Sánchez-Fenollosa, Alberto Cobos

Abstract: Abstract Stegosauria is an iconic clade of thyreophoran dinosaurs mainly characterized by two parasagittal rows of osteoderms that extend from the neck to the end of the tail. The fossil record of stegosaurian cranial material is remarkably fragmentary and scarce. This study describes the most complete stegosaurian skull from Europe and proposes a new hypothesis for the phylogenetic relationships of stegosaurs. This new cranial material was recovered from beds of the Villar del Arzobispo Formation (Upper Jurassic, Teruel, Spain) and is confidently referred to Dacentrurus armatus. It provides valuable insights into the anatomy of this species and enhances the understanding of skull evolution in stegosaurs. Furthermore, the diagnosis of D. armatus is updated with the identification of a new autapomorphy. Stegosaurian phylogenetic nomenclature is also revised. Maximum Parsimony has been applied to analyse a new stegosaurian data matrix. Phylogenetic analyses suggest that Stegosauria is divided into two major clades: Huayangosauridae and Stegosauridae. These analyses support Isaberrysaura mollensis as a stegosaur and place it within Huayangosauridae, a clade that also includes several Jurassic stegosaurs from Asia. For the first time, Mongolostegus exspectabilis is included in a phylogenetic analysis, the results of which suggests that a lineage of huayangosaurids or early-diverging stegosaurids persisted in Asia until at least the late Early Cretaceous. The new tree topologies challenge the synonymization of the genera Stegosaurus and Wuerhosaurus. Moreover, it is concluded that a taxonomic re-evaluation of Early Cretaceous Chinese stegosaurs is necessary. Alcovasaurus longispinus and Kentrosaurus aethiopicus are recovered as dacentrurines.

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DOI: 10.3897/vz.75.e152054

Authors: Javier Torres, Dexter Reilly, Claudia Nuñez-Penichet, R. Graham Reynolds, Richard E. Glor

Abstract: Abstract Cuba is the only landmass with more than one species in the Anolis carolinensis subgroup. We test the hypothesis that three rather than two distinct species occur on Cuba, based on substantial prior evidence of paraphyly. To test this hypothesis, we collected phenotypic data from all described species in the subgroup, including eastern and west-central Cuban populations of A. porcatus, and assessed phenotypic diagnosability using uni- and multivariate analyses. We also examined geographic isolation using all available occurrence records for Cuban lineages. Additionally, we conducted ecological niche modeling and niche overlap analyses, considering only Cuban lineages, to test for ecological differentiation. Finally, we reconstructed phylogenetic trees, incorporating all species from the subgroup for the first time. Our results support the recognition of three species in Cuba: A. allisoni and eastern and west-central A. porcatus as two distinct cryptic species, showing minimal phenotypic differentiation but clear geographic isolation, distinct ecological niches, and deep genetic divergence. We restrict the name A. porcatus to west-central Cuba, with Havana as the type locality, and formally describe the eastern Cuban populations as Anolis torresfundorai sp. nov., designating Baracoa, Guantánamo, as the type locality.

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DOI: 10.3897/vz.75.e138687

Authors: Christophe Dufresnes, Simeon Lukanov, Sven Gippner, Johanna Ambu, Ilias Strachinis, Dragan Arsovski, Benjamin Monod-Broca, Hugo Cayuela, Petros Lymberakis, Daniele Canestrelli, Dan Cogălniceanu, Nikolay A. Poyarkov, Spartak N. Litvinchuk, Tomasz Suchan, Mathieu Denoël, Daniel Jablonski

Abstract: Abstract The Balkan Peninsula hosts a great proportion of Europe’s biodiversity, and this is well illustrated by amphibian richness and endemism. Among them, the yellow-bellied toad Bombina variegata has been a model in ecology and evolution, but several aspects of its phylogeography and taxonomy remain surprisingly poorly understood. In this study, we combine cytochrome b DNA barcoding data (1238 individuals from 355 localities), mitogenome phylogenetics (17.2 kb), gene-based nuclear phylogenetics (3.7 kb from four gene fragments) and multilocus phylogenomics (4759 loci / ~554 kb obtained by double digest Restriction Associated DNA sequencing; ddRAD-seq) to re-assess the diversification of B. variegata, and revisit its nomenclatural history to assign scientific names to phylogeographic lineages. The analyses support four major lineages, one assigned to B. v. variegata (Carpathians and northwestern ranges), one assigned to B. v. pachypus (Apennine Peninsula), and two assigned to B. v. scabra (Dinarides, Hellenides and Balkanides vs. the Rhodope mountains). Spatiotemporal patterns of diversification suggest a role for a Late Miocene marine incursion in the Pannonian Plain (Paratethys) as the initial trigger of divergence, followed by a vicariance event in the Apennines and a “sky island” process of Pleistocene differentiation in the Balkan Peninsula. As it reached the Dinarides during the Late Pleistocene, B. v. variegata potentially hybridized with B. v. scabra and captured its mitochondrial DNA, which resulted in a massive cyto-nuclear discordance across all northwestern European populations. Finally, we show that the two lineages of B. v. scabra significantly differ in morphology and ventral coloration patterns, and describe the Rhodope lineage as a new subspecies.

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DOI: 10.3897/vz.74.e127269

Authors: Dorottya Győrössy, Vuong Tan Tu, Gábor Csorba, Sanjan Thapa, Péter Estók, Gábor Földvári, Tamás Görföl

Abstract: Abstract In taxonomic works, the weight to be given to morphological, mitochondrial, or nuclear signals, and the assessment of differences as species or subspecies distinctions has also varied considerably over the past decades and is largely a subjective research decision. This apparent example of the “grey zone of taxonomy” underpins the need of critical studies of as many specimens as possible and of using both mitochondrial and nuclear genes in taxonomic-systematic studies, as phylogeny based on uniparentally inherited genes alone may not represent true evolutionary scenarios. Myotis sicarius, a species occurring thorough the Himalayan foothills was found for the first time out of South Asia, in North Vietnam. Analysis of topotypical and Vietnamese specimens revealed high mitochondrial heterogeneity – at the upper limit of the usual threshold of intraspecific difference – but only minute nuclear sequence and negligible morphological differences. Albeit the large geographic distance between the two records might suggest the existence of two putative reproductively isolated taxonomic units, based on the incongruent results we concluded that the split of geographic populations of M. sicarius into different taxa is unsupported. As a morphologically closely resembling species, we also reviewed the taxonomic status of the two morphological forms of M. annectans and synonymizing M. primula with M. annectans was also corroborated by our phylogenetic analyses.

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DOI: 10.3897/vz.74.e128528

Authors: Jorge Brito, Rubí García, Francisco X. Castellanos, Gabriela Gavilanes, Jenny Curay, Julio C. Carrión-Olmedo, Daniela Reyes-Barriga, Juan M. Guayasamin, Jorge Salazar-Bravo, C. Miguel Pinto

Abstract: Abstract The Andean cloud forests of Ecuador are home to numerous unique mammals. Rodents of the tribe Thomasomyini are particularly abundant in many Andean localities, with Thomasomys – the largest genus in the subfamily Sigmodontinae (51 species) – especially species-rich and diverse. Despite recent advances on the systematics of the genus, where seven species have been described in the last five years, there is tantalizing evidence that its true diversity remains completely understood. Over the course of approximately ten years of fieldwork in Ecuador, a significant number of Thomasomys specimens were collected from various localities in both, the eastern and western Andean ranges. Through an extensive genetic study of these specimens, augmented with what is available in public databases, we argue that there exist at least 20 undescribed species in the genus, with no less that twelve potential new species in Ecuador alone. In this paper, we describe two of these species belonging to the group cinereus, one recently collected and the other previously referred to as Thomasomys sp. 1; further, we present an updated cyt b gene tree of the genus. The gene tree includes at least 56 valid and putative species and supports the monophyly of the genus, while at the same time suggest a paraphyletic “aureus” group. Our findings suggest that the genus likely exhibits additional hidden diversity in significant portions of Colombia, Peru, and Bolivia which calls for the need for a comprehensive reassessment of the entire genus. The recognition of these two new species brings the total number of known Thomasomys to 53 species, 19 of which occur in Ecuador, including 17 that are endemic to this country.

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DOI: 10.3897/vz.74.e127937

Authors: Alexei V. Trofimets, Christophe Dufresnes, Parinya Pawangkhanant, Andrey M. Bragin, Vladislav A. Gorin, Mahmudul Hasan, Hmar Tlawmte Lalremsanga, Mohd Abdul Muin, Dac Xuan Le, Tan Van Nguyen, Chatmongkon Suwannapoom, Nikolay A. Poyarkov

Abstract: Abstract Berdmore’s narrow-mouthed frog, Microhyla berdmorei (Blyth, 1856), is the largest member of the genus Microhyla and is distributed all over Southeast Asia, from Northeast India and southern China to Sumatra and Borneo in Indonesia. Here we demonstrate that M. berdmorei represents a complex of four species that are morphologically, acoustically, and genetically distinct from each other, and we implement taxonomic revisions. Phylogenetic analyses of three mitochondrial DNA (hereafter mtDNA, including COI, 12S, and 16S rRNA; 3119 bp) and one nuclear (BDNF; 716 bp) gene sequences are corroborated by phylogenomic analyses of 2700 ddRAD-seq loci (387,270 bp). All support that the M. berdmorei complex, which originates from the early Miocene (ca. 19.7 mya), consists of two clades that we date to the late Miocene (ca. 7.5 mya). The first clade, which regroups populations of large-sized individuals, is distributed in Indo-Burma and includes a lineage from Northeast India, Bangladesh, and Myanmar (M. berdmorei sensu stricto) and a lineage from West Malaysia, Thailand, Laos, Cambodia, and Vietnam (for which we propose to resurrect the species M. malcolmi Cochran, 1927). The second clade, which regroups populations of small-sized individuals, occurs in Sundaland and also includes two lineages for which we provide taxonomic descriptions on species rank. The first new species is widely distributed from Peninsular Malaysia to the islands of Borneo and Sumatra (described herein as M. sundaica sp. nov.), while the second one is restricted to the Malay Peninsula and occurs in extreme southern Thailand and adjacent Malaysia (M. peninsularis sp. nov.). We further provide evidence for the synonymy of Callula natatrix Cope, 1867 with M. berdmorei sensu stricto, and M. fowleri Taylor, 1934 with M. malcolmi. Our study illustrates the high diversity of Southeast Asian amphibians, especially in the genus Microhyla, which presently totals 54 species.

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DOI: 10.3897/vz.74.e128775

Authors: Peter J. McDonald, Aaron L. Fenner, Janne Torkkola, Paul M. Oliver

Abstract: Abstract Stable upland habitats in arid zone biomes are often characterised by locally endemic lineages. Explanations for this pattern include habitat or substrate specialisation (ecological specialisation) or intensifying aridity driving retreat into climatically buffered habitats (climatic refugia). Here we present an analysis of these alternative models using genetic, morphological and climate data for Diplodactylus galeatus, a gecko from central Australia that occurs in a series of isolated populations associated with dissected tablelands and mountain ranges. Analyses of mtDNA and SNP data support four distinct lineages, and dating analyses suggest divergence through the Pliocene. Morphological data show slight differences across lineages. Investigation of climate niche shows that two lineages are restricted to areas more arid than the intervening uninhabited region. These data suggest that specialisation to rocky substrates, potentially with subsequent paleo-erosion of dissected tablelands after a Pliocene wet pulse, was the key driver of divergence in this clade. Based on their deep genetic divergence, and differences in morphology and pattern, we recognise two isolated populations as new species.

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DOI: 10.3897/vz.74.e130147

Authors: Omar Torres-Carvajal, Camila Sandoval, Diego A. Paucar

Abstract: The taxonomic status of the skinks from Ecuador has never been carefully addressed. In this paper we examine populations of Mabuya lizards across Amazonian Ecuador in an attempt to establish their taxonomic identity and phylogenetic affinities. We confirm the presence of both M. altamazonica and M. nigropunctata and describe a new species from Yasuní National Park, one of the most biodiverse places on Earth. The new species differs from its congeners in lepidosis and color patterns. For the first time, we include samples from Ecuador in a molecular phylogenetic analysis of Mabuya, which confirms the monophyly of the new species and the taxonomic identity of both M. altamazonica and M. nigropunctata from Ecuador. The new species is closely related to M. bistriata. Finally, we present an identification key for species of Ecuadorian Mabuya.

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DOI: 10.3897/vz.74.e123824

Authors: Simona Papežíková, Martin Ivanov, Petr Papežík, Adam Javorčík, Konrad Mebert, Daniel Jablonski

Abstract: Abstract The Western Palearctic harbours a diverse snake fauna, including numerous endemic species and yet unnamed clades, identified through molecular analyses. However, morphological characteristics of these clades, even of common species, often remain relatively unexplored. In this study, we provide an examination of the morphology and cranial anatomy of the semi-aquatic snake species Natrix tessellata (Laurenti, 1768), with a focus on populations of the so-called ‘Europe’ and ‘Greece’ clades. Utilising both museum collections and field data, we first morphologically examined 541 individuals of N. tessellata, categorising them according to previously established clades and lineages that resulted in relatively low morphometric and meristic variation across the species’ range. When assessing the 448 specimens from the ‘Europe’ and the ‘Greece’ clades separately, we similarly observed little variation in meristic characteristics. On the other hand, individuals of the ‘Greece’ clade displayed smaller and more slender body and head proportions compared to those of the ‘Europe’ clade and the pigmentation of the labial scales is distinctively paler in the ‘Greece’ clade, whereas the overall body colouration remains largely similar between the two. Our osteological analysis of 47 N. tessellata skulls also indicated slight differences in the frontoparietal portion of the braincase between the ‘Europe’ and the ‘Greece’ clades, warranting further examination with a larger dataset and extending to other skull components. These findings hold significance for ongoing enquiries into the species’ biogeography, morphology and ecological adaptations. In summary, the integration of morphological and osteological data with genetic information offers a promising avenue for potential taxonomic revisions of N. tessellata in the future.

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DOI: 10.3897/vz.74.e126944

Authors: Franziska Wagner, Valerie Burke DeLeon, Christopher J. Bonar, Timothy D. Smith

Abstract: Abstract Comparative studies rely on the identification of homologous traits, which is challenging especially when adult stages alone are available. Inferring homology from developmental series represents the most reliable approach to recognize similar phenotypes. The primate nasal cavity exhibits a plastic morphology (shape) and topology (structure) which challenge the terminology of turbinals. Turbinal development largely corresponds to the therian template: turbinals emerge from the cartilaginous nasal capsule, ossify endochondrally, and increase their size through appositional bone growth. We studied histological serial sections and µCT data of eleven primate species in six genera representing four to five age stages (fetal to adult), and the neonate and adult stage of another primate species. We reconstructed cartilaginous precursors and followed their growth patterns until adulthood to inform the identification of structures. The developmental stages were transformed to character states for better comparison across the sample. Strepsirrhines conserved the plesiomorphic condition, with turbinal morphology similar to other placentals. In contrast, haplorhines showed a reduced turbinal number. Most strikingly, some cartilaginous turbinals are absent in the ossified nasal cavity (Saguinus); others seem to emerge as appositional bone without a cartilaginous precursor (Aotus, Pithecia). Our observation that successive developmental sequences differ from the established placental template emphasizes the significance of ontogenetic series for comparative anatomy. Structures which exhibit analogous growth patterns might be falsely considered as being homologous in adults, resulting in biased phenotypic data that strongly affects comparative analyses (e.g., phylogenetic reconstructions).

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DOI: 10.3897/vz.74.e124752

Authors: Bitupan Boruah, Surya Narayanan, Neelavar Ananthram Aravind, Samuel Lalronunga, V. Deepak, Abhijit Das

Abstract: Abstract We describe six new species of Cyrtodactylus from the khasiensis group using morphological characteristics, supported by the molecular analyses based on the mitochondrial ND2 gene. We used four different molecular species delimitation analyses that recovered six distinct undescribed lineages distributed across four states in northeastern India. Our phylogenetic analyses using ML and Bayesian approaches recovered a clade where the recently described C. arunachalensis and C. cayuensis align together with our other samples from Arunachal Pradesh, north of Brahmaputra River. Based on these results and overlapping morphological characteristics we synonymize C. arunachalensis with C. cayuensis. We provide updated comparative morphological characters for species in the khasiensis group and where available these characters are tabulated for males and females separately. Including the six new species the khasiensis group now contains 35 species, of which 26 are endemic to India.

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DOI: 10.3897/vz.74.e125958

Authors: Uwe Fritz, Hans-Werner Herrmann, Philip C. Rosen, Markus Auer, Mario Vargas-Ramírez, Christian Kehlmaier

Abstract: Abstract Trachemys is a speciose genus of freshwater turtles distributed from the Great Lakes in North America across the southeastern USA, Mexico and Central America to the Rio de la Plata in South America, with up to 13 continental American species and 11 additional subspecies. Another four species with three additional subspecies occur on the West Indies. In the present study, we examine all continental Trachemys taxa except for Trachemys hartwegi using mitochondrial and nuclear DNA sequences (3221 and 3396 bp, respectively) representing four mitochondrial genes and five nuclear loci. We also include representatives of all four West Indian species and discuss our results in the light of putative species-diagnostic traits in coloration and pattern. We provide evidence that one Mexican species, T. nebulosa, has captured a deeply divergent foreign mitochondrial genome that renders the mitochondrial phylogeny of Trachemys paraphyletic. Using nuclear markers, Trachemys including T. nebulosa represents a well-supported monophylum. Besides the mitochondrial lineage of T. nebulosa, there are six additional mitochondrial Trachemys lineages: (1) T. venusta, (2) T. ornata + T. yaquia, (3) T. grayi, (4) T. dorbigni + T. medemi, (5) T. gaigeae + T. scripta, and (6) West Indian Trachemys. These six mitochondrial lineages constitute a well-supported clade. Each mitochondrial Trachemys lineage is corroborated by our nuclear markers. For T. gaigeae another mitochondrial capture event is likely because its mitochondrial genome is sister to T. scripta, although T. gaigeae is deeply divergent in nuclear markers and resembles Mexican, Central and South American Trachemys species in morphology, sexual dimorphism and courtship behavior. The two subspecies of T. nebulosa and many Mexican and Central American subspecies of T. venusta are not clearly distinct in our studied genetic markers. Also, the putatively diagnostic coloration and pattern traits of the T. venusta subspecies are more variable than previously reported, challenging their validity. Our analyses fail to identify T. taylori as a lineage distinct from T. venusta and we propose to assign it as a subspecies to the latter species (Trachemys venusta taylori nov. comb.).

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DOI: 10.3897/vz.74.e122822

Authors: Micaela A. Chambi Velasquez, Romina Pavé, María A. Argoitia, Pablo Schierloh, María G. Piccirilli, Valeria C. Colombo, Fernando J. Beltrán, Daniel M. Cisterna, Diego A. Caraballo

Abstract: Abstract Understanding species diversity and delineating their boundaries are crucial for effective management and conservation efforts. In the case of bats, species identification holds particular importance from an epidemiological standpoint. The genus Molossus (Chiroptera: Molossidae) encompasses 15 species distributed across the Neotropics, ranging from the southeastern United States to Argentina. This genus exhibits two contrasting patterns of variation: some species are cryptic, while others are morphologically distinct yet genetically similar. This study explores the diversity of Molossus in Argentina through a molecular phylogenetic approach. We analyzed sequences from three molecular markers (cyt b, COI, and FGB) along with morphology data obtained from a sample of 64 individuals. Uni- and multivariate analyses of external and cranial measurements were conducted, alongside comparisons of external and cranial characteristics among species. Based on molecular and morphological differences, we describe a new species within the Molossus genus. This newly discovered species exhibits a broad distribution spanning the Paraná River basin across three distinct ecoregions. It is noteworthy that this species is pseudo-cryptic with respect to similar-sized species such as M. molossus and M. melini. Additionally, it is important to mention that all species in Argentina have overlapping distribution ranges. In summary, this study provides valuable insights into the diversity and distribution of Molossus bats in Argentina, employing molecular and morphological analyses. The discovery of a new species underscores the ongoing importance of comprehensive research efforts in understanding and conserving bat populations in the Neotropics.

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DOI: 10.3897/vz.74.e122418

Authors: Jason R. Ali, Uwe Fritz

Abstract: Abstract Our focus is the colonization history of the Galápagos’ vertebrate clades: 11 land-bound groups (eight reptiles, three rodents) and 13 taxa of flyers and swimmers (ten winged birds, two pinnipeds, one penguin). Using ‘colonization intervals’ and ‘colonization profiles’, it is clear that the two sets of taxa assembled very differently. The former includes older clades with between one, and potentially eight, predating the emergence of the oldest island (4 Mya). For the origin of some lineages, now-sunken landmasses associated with the Galápagos mantle-plume hotspot must have been involved, but for others it could reflect taxonomic uncertainties. In contrast, the taxa of flyers and swimmers are markedly younger, indicating either higher rates of colonization and extirpation for these sorts of animal, or continued genetic influx from mainland populations, or some combination of both factors. Concerning the first, possible drivers are the environmental stressors associated with the El Niño–La Niña climate system; the recent clades may be vulnerable to extreme events within the oscillation sequence, perhaps on ≥104-year timescales. Therefore, loose temporal thresholds might exist for the archipelago’s vertebrate groups beyond which selection fortifies them from the most challenging of seasonal states. Moreover, in a world of climate uncertainty, the findings appear relevant to conservation initiatives suggesting a focusing on the younger elements within the Galápagos’ biota.

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DOI: 10.3897/vz.74.e112132

Authors: Parinya Pawangkhanant, Nikolay A. Poyarkov, Harry Ward-Smith, Rupert Grassby-Lewis, Montri Sumontha, Nikita S. Kliukin, Sabira S. Idiiatullina, Alexei V. Trofimets, Chatmongkon Suwannapoom, Justin L. Lee

Abstract: Abstract We describe a new species of kukri snake (Oligodon Fitzinger, 1826) from the limestone karst formations of Satun and Trang Provinces in southern Thailand. Phylogenetic analyses based on three mitochondrial DNA fragments (12S–16S ribosomal rRNA and cytochrome b) recover the new species within the Oligodon cinereus species complex, where it forms a deeply divergent yet poorly supported clade sister to Oligodon saiyok Sumontha et al., 2017 and another unnamed lineage currently referred to Oligodon cinereus (Günther, 1864) from southwest Myanmar. Morphologically, the new species is distinguished from all other members of the genus by the following combination of characters: ventral scales 189–193 with distinct lateral keeling; subcaudal scales 47–54, paired; anterior dorsal scale rows 17–19, with the reduction from 19 to 17 rows occurring above the 28th–30th ventral scale when present; maxillary teeth 8, blade-like and laterally compressed; dorsum olive–gray, plain; ventral surface white anteriorly, dark gray posteriorly; underside of tail dark gray, smeared with white. We briefly discuss the natural history and conservation status of this new species and provide observations of other kukri snakes inhabiting limestone karst habitats. Our study also incorporates genetic samples of four recently described Oligodon endemic to Thailand, all of which are recovered in the O. cinereus species complex. In agreement with previous studies, we demonstrate that species-level diversity within the O. cinereus species complex is underestimated, and additional sampling is necessary to revise this taxonomically challenging clade.

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DOI: 10.3897/vz.74.e114285

Authors: Stephen Mahony, Rachunliu G. Kamei, Rafe M. Brown, Kin Onn Chan

Abstract: Abstract Although the differentiation of clades at the species level is usually based on a justifiable and testable conceptual framework, the demarcation of supraspecific boundaries is less objective and often subject to differences of opinion. The increased availability of large-scale phylogenies has in part promulgated a practice of what we consider excessively splitting clades at the “genus” level. Many of these new genus-level splits are predicated on untenable supporting evidence (e.g., weakly supported phylogenies and purportedly “diagnostic” but actually variable, non-exclusive, or otherwise problematic opposing character state differences) without careful consideration of the effects on downstream applications. As case studies, we critically evaluate several recent examples of splitting established monophyletic genera in four amphibian families that resulted in the creation/elevation of 20 genus-level names (Dicroglossidae: Phrynoglossus, Oreobatrachus, Frethia split from Occidozyga; Microhylidae: Nanohyla split from Microhyla; Ranidae: Abavorana, Amnirana, Chalcorana, Humerana, Hydrophylax, Indosylvirana, Papurana, Pulchrana, Sylvirana split from Hylarana; Rhacophoridae: Tamixalus, Vampyrius, Leptomantis, Zhangixalus split from Rhacophorus, Rohanixalus split from Feihyla, Orixalus split from Gracixalus, and Taruga split from Polypedates), and also address the taxonomic status of the monotypic genus Pterorana relative to Hylarana. We reassess the original claims of diagnosability and justifications for splitting and argue that in many cases, the generic splitting of clades is not only unnecessary but also destabilizes amphibian taxonomy, leading to a host of downstream issues that affect categories of the user community (stakeholders such as taxonomists, conservationists, evolutionary biologists, biogeographers, museum curators, educators, and the lay public). As an alternative, we advocate for the use of the subgenus rank in some cases, which can be implemented to establish informative partitions for future research without compromising on information content, while avoiding gratuitous (and often transient) large-scale binomial (genus-species couplet) rearrangements. We encourage taxonomists to consider the actual needs and interests of the larger non-taxonomic end-user community who fund the majority of taxonomic research, and who require a system that remains reasonably stable and is relatively intuitive, without the need for inaccessible laboratory equipment or advanced technical scientific knowledge to identify amphibian species to the genus level.

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DOI: 10.3897/vz.74.e109088

Authors: Kai Wang, V. Deepak, Abhijit Das, L. Lee Grismer, Shuo Liu, Jing Che

Abstract: Abstract Owing to the harsh terrain, few biodiversity surveys have been carried out in the Pan-Himalaya Region. Among the understudied taxa from this region, Jerdon’s forest lizard, Calotes jerdoni, is believed to have a wide distribution, from northeast India to southwestern China. However, given the heterogeneous environment across its range and the lack of studies on this species, its taxonomy remains questionable. Using integrative taxonomic methods, we combined both morphological and genetic data from the type and topotypic specimens and examined the current taxonomic hypothesis of C. jerdoni across its range. Molecular data reveal that C. jerdoni as currently recognized, contains three deeply diverged lineages: one from the type locality in Northeast India, one from Western Myanmar, and another one from Southwestern China. The uncorrected genetic distances of mitochondrial coding gene ND2 among these three clades ranged over 10%. The Chinese population is sister to C. medogensis and paraphyletic to the remaining two clades of C. jerdoni. Morphological analyses confirm the results of the molecular analyses, where the Myanmar and Chinese populations can be diagnosed statistically in both univariate and multivariate space from the true C. jerdoni, as well as by a suite of reliable categorical morphological characters, including the size and shape of gular scales and ventral scales. To resolve the current taxonomic confusion, we resurrect the junior synonym, C. yunnanensis, for the Chinese population and expand its distribution to Myanmar, redescribe the elusive C. maria and C. medogensis based on its type material, and describe the remaining western Myanmar population as a new species. We further discuss the possibility of additional cryptic species within the complex in the Pan-Himalaya Region and provide a diagnostic key to all recognized members of the C. jerdoni complex.

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DOI: 10.3897/vz.74.e113899

Authors: Hao-Tian Wang, Shuo Qi, Dan-Yang Zhou, Ying-Yong Wang

Abstract: Abstract A new species of the genus Gekko Laurenti, 1768, Gekko paucituberculatus sp. nov., is described here, based on two specimens from Tianyang District, Baise City, Guangxi Zhuang Autonomous Region, China. It was placed in subgenus Japonigekko on the basis of morphological and molecular phylogenetic analysis and can be distinguished from all congeners in this subgenus by significant divergences in the mitochondrial 16S and ND2 genes and by a combination of the following morphological characters: Moderate body size, SVL 77.2 mm in the adult male and SVL 85.9 mm in the adult female; tubercles only present along dorsolateral trunk and absent on other regions; fingers and toes with weak webbing; continuous precloacal pores 12 in the male, absent in the female; a single postcloacal tubercle on each side; a light-coloured vertebral line from nape to tip of tail; dorsum greyish-brown, with 7–8 dirty-white bands between nape and sacrum. Meanwhile, the distribution of G. palmatus in China has been confirmed as occurring in Guangxi and Guangdong Provinces. This study brings the total species of the subgenus Japonigekko in China to 19.

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DOI: 10.3897/vz.74.e106238

Authors: Vinícius Sudré, Albedi Andrade-Junior, Manuella Folly, Josué A. R. Azevedo, Robson Waldemar Ávila, Felipe Franco Curcio, Pedro M. Sales Nunes, Paulo Passos

Abstract: Abstract Currently, the proposed diagnoses for the Chironius bicarinatus complex reflect a wide variation in color pattern and pholidosis. Herein, we review the Chironius bicarinatus complex based on morphological and molecular data from a sample of 485 specimens covering the species distribution. Our results corroborate the recognition of C. bicarinatus and C. gouveai, and diagnose a distinct lineage without an available name. Thus, here we describe this new species restricted to the Baturité Massif, a relictual rainforest isolated in the Caatinga xerophytic domain, in the state of Ceará, northeastern Brazil. The new species can be distinguished from its congeners by its unique combination of qualitative and quantitative morphological characters (scale counts, morphometric, color pattern), and is also supported by molecular and ecological evidence. Additionally, we rectify data on the distribution and morphological variability of C. gouveai to accurately infer the boundaries between this taxon and C. bicarinatus, which was not properly addressed. Finally, we discuss our results in the light of previous studies that suggest diversification hypotheses in the Atlantic Forest already detected for other taxa, highlighting the importance of conserving the areas of “Brejos de Altitude”, in northeastern Brazil, and the southern limit of Serra do Mar up to Serra do Tabuleiro, in southern Brazil.

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DOI: 10.3897/vz.74.e110674

Authors: Ishan Agarwal, Tejas Thackeray, Akshay Khandekar

Abstract: Abstract Subdoluseps is a recently described genus of Lygosomine skinks distributed in peninsular India and Southeast Asia. We conduct the first revision of Indian Subdoluseps based on range-wide sampling including 89 specimens from 33 localities. We use two mitochondrial and three nuclear markers, 58 morphological characters, and ecological data to reconstruct the evolutionary history of Indian Subdoluseps and assess their diversity and distribution, providing insights into lygosominin biogeography. We formally describe the Indian clade as a new genus, Dravidoseps gen. nov. and name five new species from Tamil Nadu, India in an integrative taxonomic framework – D. gingeeensis sp. nov., D. jawadhuensis sp. nov., D. kalakadensis sp. nov., D. srivilliputhurensis sp. nov., and D. tamilnaduensis sp. nov.. We transfer Riopa goaensis, Subdoluseps pruthi and S. nilgiriensis to the new genus and designate neotypes for the former two. Members of Dravidoseps gen. nov. are the first known viviparous skinks from peninsular India and the only known viviparous lygosominins apart from a few species of east African Mochlus. The Lygosomini have a Southeast Asian origin and began diversifying in the Eocene with three dispersals between India and Southeast Asia. Species level diversification in Dravidoseps gen. nov. was likely driven by a combination of niche conservatism, paleoclimate and past forest distribution. The discovery of a new genus and five new species reiterates the high levels of diversity and endemism present in peninsular India and how much more remains to be discovered.

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DOI: 10.3897/vz.74.e103573

Authors: Manuella Folly, Thais H. Condez, Davor Vrcibradic, Carlos F. D. Rocha, Alessandra S. Machado, Ricardo T. Lopes, José P. Pombal Jr.

Abstract: Abstract Brachycephalus is a genus of small ground-dwelling anurans, endemic to the Brazilian Atlantic Forest. Recent molecular analyses have corroborated the monophyly of three species groups within this genus (B. ephippium, B. ephippium, and B. ephippium). In the meantime, the genus has been targeted as a group with recent taxonomic issues owing to its interspecific morphological similarity and genetic conservatism. Herein, we describe a new species of Brachycephalus from the northern portion of Serra do Mar mountain range, in the state of Rio de Janeiro, Brazil. It belongs to the B. ephippium species group, exhibiting moderate hyperossification of the skull and vertebral column. The new species can be distinguished from all other congeners based on morphological, acoustic, and molecular data. Furthermore, we provide information on osteology and natural history of the new species.

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DOI: 10.3897/vz.73.e109309

Authors: Ernesto Castillo-Urbina, Miguel Vences, César Aguilar-Puntriano, Frank Glaw, Jörn Köhler

Abstract: Abstract We studied the taxonomic status of a population of Pristimantis from the southern Cordillera Azul, Departamento Huánuco, central Peru. A phylogenetic analysis based on the mitochondrial 16S rRNA gene revealed that it represents a lineage within the Pristimantis lacrimosus species group, being the closest relative of a species of uncertain taxonomic status from a lowland rainforest in central Peru (Panguana), and P. pulchridormientes from the Tingo Maria National Park. However, the focal lineage is divergent from all nominal species in the P. lacrimosus group for which respective data are available by >7.9% uncorrected pairwise distance in the 16S rRNA gene fragment. An integrative taxonomic approach, including morphological and bioacoustic analyses, provided multiple lines of evidence for the focal specimens belonging to an unnamed evolutionary lineage at the species level that we describe and name herein. The systematics of Peruvian populations associated with the P. lacrimosus group are discussed, particularly highlighting problematic taxa with uncertain taxonomic status and unknown relationships. We point to scientific challenges and actions needed to achieve a better taxonomic resolution of this species-rich clade of frogs.

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DOI: 10.3897/vz.73.e110626

Authors: Karin Tamar, Marek Uvizl, Mohammed Shobrak, Mohammed Almutairi, Salem Busais, Al Faqih Ali Salim, Raed Hamoud M. AlGethami, Abdulaziz Raqi AlGethami, Abdulkarim Saleh K. Alanazi, Saad Dasman Alsubaie, Laurent Chirio, Salvador Carranza, Jiří Šmíd

Abstract: Abstract A recent molecular phylogeny of the agamid genus Pseudotrapelus, distributed in the rocky areas of North Africa and the Arabian Peninsula, revealed the presence of a genetically distinct lineage around the city of Riyadh in central Saudi Arabia. With the inclusion of additional specimens, we were able to describe this lineage as a new species, P. tuwaiqensis sp. nov., confined to the Tuwaiq Escarpment, thus endemic to central Saudi Arabia. Our results of morphological examinations and molecular analyses, using three mitochondrial (COI, 16S, ND4-tRNAs) and two nuclear (c-mos, MC1R) gene fragments, show the new species is genetically differentiated and phylogenetically close to P. sinaitus and P. chlodnickii.

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DOI: 10.3897/vz.73.e102645

Authors: Juan Fernando Vélez-García, Diego Alejandro Carrión Blanco, Gabriela Moreno Gómez, Stephanie San Martín Cañas

Abstract: Abstract The kinkajou (Potos flavus) is a carnivoran of the suborder Caniformia and the family Procyonidae, inhabiting regions throughout Central and South America. Potos flavus has arboreal preferences and exhibits unique anatomical adaptations that facilitate movement within trees. Its pelvic limbs enable hindfoot reversal, while its thoracic limbs possess remarkable prehensile capabilities. Previous anatomical studies in Potos flavus have presented discrepancies in the description of the intrinsic shoulder and brachial muscles. Therefore, this study aims to provide a comprehensive anatomical description of these muscles in five specimens. The findings are compared with descriptions reported for other caniforms. The application of the Density-Based Spatial Clustering of Application with Noise (DBSCAN) algorithm aids in identifying relationships among caniforms based on the presence or absence of specific muscles. Our analysis reveals several key differences, including the presence of a biceps brachii with two capita (longum and breve), two coracobrachiales muscles (longus and brevis), a tensor fasciae antebrachii with two distinct parts (cranialis and caudalis), and an anconeus medialis. The caput breve of the biceps brachii and coracobrachialis longus muscles are absent in some individuals, with prevalence rates of 10% and 20%, respectively. One specimen exhibited an accessory caput laterale of the m. triceps brachii bilaterally. The comparative analysis suggests that the shoulder and brachial muscles of Potos flavus share more similarities with those of Ailurus fulgens and ursids of the genera Ursus and Tremarctos. These findings suggest the retention of muscles that may have been present in the common ancestor of the infraorder Arctoidea.

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DOI: 10.3897/vz.73.e95639

Authors: Michael F. Bates, Javier Lobón-Rovira, Edward L. Stanley, William R. Branch, Pedro Vaz Pinto

Abstract: Cordylus angolensis (Bocage, 1895) was described 128 years ago on the basis of a single specimen collected at Caconda in the west-central highlands of Angola. Additional specimens referred to this species were collected at ‘Mombolo’ (also in the central highlands) during the Vernay Angola Expedition in 1925. As the holotype was apparently destroyed in the fire of 1978 at the Museu Bocage in Lisbon and no additional specimens have been collected, its taxonomic status and phylogenetic relationships has remained uncertain. The species has eluded all efforts aimed at its re-discovery in the vicinity of the type locality, with a single specimen from near Condé, north of Mombolo—collected in 1970 by Wulf Haacke—the only other specimen of Cordylus known from west-central Angola. Recent field work in the Angolan highlands resulted in the collection of a series of specimens from Taqueta Mountain (west of Caconda), Monte Verde (Sandula, ‘Mombolo’) and Uassamba (Vondo). A phylogenetic analysis, using three mitochondrial and six nuclear genes, indicated the existence of two distinct species-level lineages in the Angolan highlands. These two species are allopatric and morphologically distinct, differing especially in terms of their colour patterns, eye colour and certain scalation characteristics. We therefore confirm that C. angolensis is a valid species and designate a neotype, and describe a new species, Cordylus momboloensis sp. nov.

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DOI: 10.3897/vz.73.e98444

Authors: Bitupan Boruah, V. Deepak, Naitik G. Patel, Vijayan Jithin, Tajum Yomcha, Abhijit Das

Abstract: Abstract We describe a new species of rhacophorid frog of the genus Gracixalus from northeast India based on molecular, morphological and acoustic evidence. The new species, formally described herein as Gracixalus patkaiensis sp. nov., is morphologically distinct from other congeners by a suite of morphological characters such as snout-vent length 23.6–26.5 mm in adult males; green dorsum with irregular brown spots; dorsal skin shagreened with numerous spinules; snout shape nearly acuminate in dorsal and ventral view; a prominent dark streak along the cranial margins; white reticulations along lateral side and ventrum distinct in life. Genetically, the new taxon is found to differ from all the recognized Gracixalus species by 4–14.8% divergence in the 16S mitochondrial gene. The discovery confirms the presence of genus Gracixalus from the Indian state of Arunachal Pradesh underlines the importance of biological exploration even in well-known protected areas of India.

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DOI: 10.3897/vz.73.e104438

Authors: Uwe Fritz, Christian Kehlmaier, Rodney J. Scott, Raúl Fournier, James R. McCranie, Natalia Gallego-García

Abstract: Using 3226-bp-long mtDNA sequences and five nuclear loci (Cmos, ODC, R35, Rag1, Rag2, together 3409 bp), we examine genetic differentiation and relationships of Central American slider turtles (Trachemys grayi, T. venusta). Our investigation also included samples from taxa endemic to North America (T. gaigeae, T. scripta), the Antilles (T. decorata, T. decussata, T. stejnegeri, T. terrapen), and South America (T. dorbigni, T. medemi plus the two T. venusta subspecies endemic to northern South America). Our mitochondrial phylogeny retrieves all studied species as distinct, with three well-supported clades in a polytomy: (1) the Central and South American species (T. grayi + T. venusta) + (T. dorbigni + T. medemi), (2) the Antillean species, and (3) T. gaigeae + T. scripta. Our nuclear DNA analyses also suggest three distinct but conflicting clusters: (1) T. scripta plus the Antillean species, (2) T. gaigeae, and (3) the Central and South American species T. dorbigni, T. grayi, T. medemi, and T. venusta. However, in the mitochondrial phylogeny, T. gaigeae is the little divergent sister taxon of T. scripta. This conflicting placement of T. gaigeae suggests a distinct evolutionary trajectory and old hybridization with T. scripta and mitochondrial capture. Despite prominent color pattern differences, genetic divergences within T. grayi and T. venusta are shallow and the taxonomic diversity of each species with several currently recognized subspecies could be overestimated. Finally, we provide for the first time evidence for the occurrence of T. grayi along the Caribbean versant of Costa Rica.

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DOI: 10.3897/vz.73.e104494

Authors: Surya Narayanan, Peter Christopher, Kothandapani Raman, Nilanjan Mukherjee, Ponmudi Prabhu, Maniezhilan Lenin, Sivangnanaboopathidoss Vimalraj, V. Deepak

Abstract: A new large-bodied (SVL 101–109 mm) gecko of the genus Hemidactylus is described from the Gingee Hills in the Eastern Ghats of India. The new species is closely related to H. graniticolus and the recently described H. easai, from which it can be distinguished by its lower femoral pores count. The new species described here was previously identified as H. cf. graniticolus based only on the molecular data, pending its formal description. Our findings were consistent with the results from the molecular DNA analyses, showing that this population is morphologically distinct from other closely related species. As a result, we formally describe this lineage as a new species, providing a comprehensive description of its morphological characteristics based on a type series of five specimens and compare it with its congenerics.

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DOI: 10.3897/vz.73.e94063

Authors: Thore Koppetsch, Maciej Pabijan, Carl R. Hutter, Jörn Köhler, Philip-Sebastian Gehring, Andolalao Rakotoarison, Fanomezana M. Ratsoavina, Mark D. Scherz, David R. Vieites, Frank Glaw, Miguel Vences

Abstract: The small arboreal frog Guibemantis liber (Anura: Mantellidae) has served as an example for the existence of deep conspecific lineages that differ by a substantial amount in mitochondrial DNA but are similar in morphology and bioacoustics and thus are assigned to the same nominal species. During fieldwork in northern Madagascar, we identified additional such lineages and surprisingly, observed close syntopy of two of these at various sites. In-depth study based on DNA sequences of the mitochondrial cytochrome b gene from 338 specimens of G. liber sensu lato from across its range, sequences of four nuclear-encoded markers for 154‒257 of these specimens, a phylogenomic dataset obtained by the FrogCap target capture approach, and additional mitochondrial genes for representatives of most mitochondrial lineages, as well as bioacoustic and morphological comparisons, revealed concordant differentiation among several lineages of the G. liber complex. We identify nine lineages differing by 5.3‒15.5% in cytochrome b and 2.4‒10.1% in the 16S rRNA gene, and find that several of these lack or have only limited allele sharing in the nuclear-encoded genes. Based on sympatric or parapatric occurrence without genetic admixture, combined with differences in bioacoustic and morphological characters, we scientifically name three lineages from northern Madagascar as new species: G. razoky sp. nov., G. razandry sp. nov., and G. fotsitenda sp. nov. Of these new species, G. razoky sp. nov. and G. razandry sp. nov. show widespread syntopy across northern Madagascar and differ in body size and advertisement calls. Guibemantis fotsitenda sp. nov. is sister to G. razandry sp. nov., but appears to occur at lower elevations, including in close geographic proximity on the Marojejy Massif. We also detected subtle differences in advertisement calls among various other mitochondrial lineages distributed in the Northern Central East and Southern Central East of Madagascar, but the status and nomenclatural identity of these lineages require further morphological and bioacoustic study of reliably genotyped individuals, and assignment of the three available names in the complex: Rhacophorus liber Peracca, 1893, Gephyromantis albogularis Guibé, 1947, and Gephyromantis variabilis Millot and Guibé, 1951. We discuss the identity and type material of these three nomina, designate a lectotype for Gephyromantis variabilis from Itremo, and flag the collection of new material from their type localities, Andrangoloaka and Itremo, as paramount for a comprehensive revision of the G. liber complex.

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DOI: 10.3897/vz.73.e98618

Authors: Anna B. Vassilieva, Thi Van Nguyen

Abstract: We compare reproductive features, development, and larval morphology in three closely related species of sticky frogs (Kalophrynus Tschudi, 1838) inhabiting the lowland and mountain forests of Vietnam and displaying a variety of reproductive modes. While K. interlineatus breeds in open temporary ponds, K. honbaensis and K. cryptophonus are phytotelm-breeders using tree hollows and bamboo stems for reproduction. Their tadpoles also differ in trophic specialization: larval K. interlineatus are typical suspension-feeders, whereas K. honbaensis and K. cryptophonus are obligatorily oophagous. All three species differ in egg and clutch sizes, duration of embryonal period and hatching stage, and the structure of the larval digestive tract and skeleton. Based on external and internal morphology, we conclude that tadpoles of K. interlineatus and K. cryptophonus represent two “extremes” of the adaptive spectrum of microhylid larvae, while K. honbaensis displays a set of transitory traits. Relying on these new findings in anuran biology, we discuss reproductive, ontogenetic, and morphological rearrangements during the transition from pond breeding to phytotelm breeding and from microphagy to macrophagy as well as the significance of the revealed adaptations to different habitats and larval life modes.

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DOI: 10.3897/vz.73.e101329

Authors: Mariana P. Marques, Diogo Parrinha, Luis M. P. Ceríaco, Ian G. Brennan, Matthew P. Heinicke, Aaron M. Bauer

Abstract: Several specimens of Pachydactylus angolensis, a poorly known Angolan endemic gecko, have recently been collected in southern Angola, considerably increasing its known distribution range. Previous observations led to the hypothesis that two different morphological forms exist in the country—a coastal form and an inland form. Based on the morphological examination of historical and recently collected specimens, as well as on newly generated molecular data, we conducted a taxonomic revision of this putative species complex. The results support the separation of these two forms as two different species. The coastal form belongs to the nominotypic population, while the inland form is here described as a new species, Pachydactylus maiatoi sp. nov.. A brief comment on the biogeographical implications of this discovery is also provided.

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DOI: 10.3897/vz.73.e96656

Authors: Diego H. Verzi, Nahuel A. De Santi, A. Itatí Olivares, Cecilia C. Morgan, Néstor G. Basso, Federico Brook

Abstract: The subterranean rodent Ctenomys is the most polytypic South American mammal genus and one of the most speciose and rapidly diversifying mammal genera in the world. Its systematics is unstable due to the underlying accelerated diversification processes that give rise to evolutionary lineages at different stages of differentiation and to remarkable morphological homogeneity even among long-differentiated species. As a result, species boundaries are often difficult to define. Diversity of this genus in the coastal area of central Argentina has been extensively studied, with two independent lineages currently recognized while a distinct third population had not been previously detected. Through a phylogenetic analysis based on combined morphological and molecular evidence, Bayesian estimates of divergence times, and morphometric and morphological assessments, we recognize this third population as an independently evolving lineage. The new species, Ctenomys pulcer sp. nov., is here described for both the living fauna and the fossil record of the Pampean region of central Argentina. According to phylogenetic results, Ctenomys pulcer sp. nov. belongs to the essentially Patagonian magellanicus clade, and would have diverged from its sister species, Ctenomys bidaui, during the middle Pleistocene (ca. 0.4 Ma). Its current distribution in the fixed and semifixed dunes of the coastal Pampean region is assumed to represent a relict of a wider and continuous distribution of potentially suitable environments during the late Pleistocene. Ctenomys pulcer sp. nov. occurs in a particularly fragile natural system subjected to profound disturbances caused by diverse anthropic actions and therefore measures for the conservation of its habitat will be indispensable.

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DOI: 10.3897/vz.73.e94111

Authors: Willem G. Coetzer

Abstract: The greater cane rat (Thryonomys swinderianus) is an African rodent with a wide Sub-Saharan distribution range. This species is viewed as an important protein source in many African countries. These rodents are also regularly viewed as a pest species who frequently raid croplands in agricultural settings. No phylogenetic work has to date been published on T. swinderianus from southern Africa. This paper therefore reports the first phylogenetic assessment on the species across the South African distribution range. Thirty samples were sourced from local museum collections, with one direct submission by a member of the public who found a rodent carcass identified as T. swinderianus west of its known distribution range in the Eastern Cape Province of South Africa. Two mitochondrial loci previously used in West African studies of this species were used in the current study to asses T. swinderianus population genetic diversity and phylogenetic structure across the South African distribution. A comparison to sequence data from West Africa was also performed. A divergence time estimation was conducted to further investigate the evolutionary history of the South African sub-population. Similar genetic diversity estimates were observed for the South African sub-population when compared to the West African datasets. Specimens from the eastern parts of South Africa showed higher genetic diversity estimates, possibly indicative of an initial colonisation site from eastern Africa. Two distinct phylogenetic clades were identified by Bayesian inference, forming distinct West African and South African groups. The divergence estimates showed similar ages for the T. swinderianus most recent common ancestor (MRCA) as previously reported. The MRCA estimates for the South African group identified a possible middle to late Pleistocene migratory event from eastern African into southern Africa. Further fine scale sampling across the African distribution range is however needed to provide more accurate assessments for future conservation management planning for the different sub-populations, as needed.

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DOI: 10.3897/vz.73.e90979

Authors: Suranjan Karunarathna, Kanishka D. B. Ukuwela, Anslem De Silva, Aaron M. Bauer, Majintha Madawala, Nikolay A. Poyarkov, Madhava Botejue, Dinesh Gabadage, L. Lee Grismer, Vladislav A. Gorin

Abstract: Sri Lanka is a local hotspot for Cnemaspis day geckos with 40 currently known species with 100% endemism. In this paper, we evaluate the phylogenetic relationships of Cnemaspis species belonging to the alwisi group of the podihuna clade and describe two additional new species of Cnemaspis from Sri Lanka; one from Galgiriya mountain, Kurunegala District, and another from Ethagala mountain, Ampara District. These new species were recorded from granite caves within forested areas in isolated mountains in the dry bioclimatic zone (point-endemics). Both new species are microhabitat specialists with narrow niches limited to humid, cool, canopy-shaded granite caves and old buildings associated with granite caves, where they are camouflaged by their cryptic morphology and body colouration. Furthermore, both species prefer narrow (~ 6–12 mm), long (~ 120–450 mm) and deep (~ 80–260 mm) crevices as refugia. The regions in which these habitats are located receive relatively low annual rainfall (1,000–1,500 mm). These new species are medium in size (28.5–36.8 mm SVL) and can be differentiated from all other Sri Lankan Cnemaspis by the presence of clearly enlarged, subhexagonal subcaudal scales and the absence of precloacal pores in males. Both species described here are categorised herein as Critically Endangered (CR) under the IUCN Red List criteria. The major threats for these new species are habitat loss due to expansion of commercial-scale agriculture, illicit forest encroachments, and forest fires. Therefore, we recommend that relevant authorities take immediate conservation action to ensure the protection of these forest areas with their buffer zones in the near future.

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DOI: 10.3897/vz.73.e101311

Authors: Surya Narayanan, Saunak Pal, L. Lee Grismer, N. A. Aravind

Abstract: We describe a new species of rupicolous Cnemaspis from the Male Mahadeshwara Wildlife Sanctuary in the Eastern Ghats. The current description is based on a type series of eight specimens that bear a unique combination of morphological and colour pattern characteristics that do not occur in other closely related species. Maximum likelihood and Bayesian analyses based on the mitochondrial gene ND2 recovered the new species to be nested within the gracilis clade and the sister species to the recently described C. agarwali from the Eastern Ghats.

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DOI: 10.3897/vz.73.e94265

Authors: Rafael A. Moreno-Arias, Miguel A. Méndez-Galeano, Iván Beltrán, Mario Vargas-Ramírez

Abstract: The Anolis heterodermus group comprises eight big-headed and short-legged lizard species from the highlands of the northernmost South American Andes. Recent studies revealed unknown lineages within this group that had previously been categorized as a species complex. By widely sampling and applying an integrative taxonomic framework, we (1) assessed the species diversity of the group using a molecular dataset (two mitochondrial and one nuclear markers) along with an inclusive morphological study (scalation, scale configuration and ornamentation, morphometrics, and dewlap and body colour patterns); and (2) we inferred the evolutionary relationships within this species group. Our analyses confirmed the formerly reported differentiation between populations of those high-altitude lizards, and we identified several unknown evolutionary lineages. Our results provided evidence for the existence of nine distinct, independently evolving evolutionary lineages in the heterodermus group. As a result, we described two morphologically and genetically highly distinct lineages as species new to science (A. quimbaya sp. nov. and A. tequendama sp. nov.). We redescribed A. heterodermus and erected as a valid species Anolis richteri, a previously described synonym of A. heterodermus. A taxonomic key for the identification of species of the Phenacosaurus clade was presented. The identification of two additional poorly-known lineages suggested that the diversity of this group of lizards is still unknown; therefore, it is necessary to establish measurements for the group´s conservation, as well as to perform fieldwork and revision of herpetological collections to identify possible hidden diversity within the group.

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DOI: 10.3897/vz.73.e96958

Authors: Platon V. Yushchenko, Justin L. Lee, Hieu Minh Pham, Peter Geissler, Elena V. Syromyatnikova, Nikolay A. Poyarkov Jr.

Abstract: We investigated the taxonomic status of the recently described kukri snake Oligodon arenarius Vassilieva, 2015 and the morphologically similar Oligodon macrurus (Angel, 1927), two species endemic to the southern coast of Vietnam. Based on phylogenetic analyses using three mitochondrial genes (12S–16S rRNA, cytochrome b), we recovered O. arenarius and O. macrurus in a clade within the O. cyclurus-taeniatus species group, agreeing with previous intrageneric classifications. Genetic distances between O. arenarius and O. macrurus are extremely low (less than 0.5% based on 12S–16S) and render O. arenarius paraphyletic. All preserved specimens of O. arenarius and O. macrurus convey little to no differences in color pattern, hemipenial morphology and osteological features; the latter of which is based on three dimensional micro computer tomography (µCT) scans of one specimen per species. Contrasting these results, univariate and multivariate analyses revealed significant differences in relative tail length, and the number of ventral and subcaudal scales between both species. Although the molecular and morphological datasets present conflicting results, integrating the evidence leads us to synonymize O. arenarius with O. macrurus. We provide a formal redescription of O. macrurus, designate a neotype specimen to avoid future taxonomic confusion, and provide the first detailed osteological description of this species. Oligodon macrurus sensu stricto is endemic to coastal dunefields and adjacent forest habitats in southern Vietnam, where ongoing human development, tourism and road mortality pose significant threats to its conservation. Consequently, we suggest that O. macrurus should be listed as “Vulnerable” based on the assessment criteria of the International Union for Conservation of Nature (IUCN).

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DOI: 10.3897/vz.73.e91230

Authors: Platon V. Yushchenko, Justin L. Lee, Thy Neang, Hun Seiha, Nguyen Van Tan, Gernot Vogel, Nikolay A. Poyarkov Jr.

Abstract: The ashy kukri snake Oligodon cinereus (Günther, 1864) is a widely distributed and morphologically variable species found throughout mainland Southeast Asia. In this paper, we re-assessed the taxonomic status of O. cinereus populations found in southern Indochina (southern Vietnam, Cambodia, and southern Laos), including the recently described Cat Tien kukri snake Oligodon cattienensis Vassilieva et al., 2013, which was previously confused with this species. Phylogenetic analyses using mitochondrial DNA from the 12S–16S ribosomal subunit and cytochrome b gene revealed that O. cattienensis is embedded in a mixed clade containing samples of the subspecies O. cinereus pallidocinctus, which bears a dorsal color pattern with white crossbars and black edges. This clade forms a strongly supported sister group with a topotypic sample of O. cinereus cinereus, representing populations bearing a uniform dorsal color pattern and slight reticulate markings, however the genetic divergence between the two clades is very low. The morphological characters used to distinguish O. cattienensis from O. cinereus sensu lato broadly overlap and supposed differences in hemipenial morphology between the two taxa are due to outdated terminologies used to describe the organ. We relegate both O. cattienensis and O. cinereus pallidocinctus to the junior synonymy of O. cinereus and consider all color patterns of this species found near the type locality in Cambodia, southern Laos, and southern Vietnam to represent O. cinereus sensu stricto. Future integrative investigations across the range of O. cinereus sensu lato are needed to resolve the status of the remaining subspecies and synonyms associated with this taxon. Problems associated with hemipenial morphology and Oligodon systematics are also discussed.

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DOI: 10.3897/vz.73.e98367

Authors: Rajeev Raghavan, Remya L. Sundar, C.P. Arjun, Ralf Britz, Neelesh Dahanukar

Abstract: The lateritic aquifers of the southern Indian state of Kerala harbour a unique assemblage of enigmatic stygobitic fishes which are encountered very rarely, only when they surface during the digging and cleaning of homestead wells. Here, we focus on one of the most unusual members of this group, the catfish Horaglanis, a genus of rarely-collected, tiny, blind, pigment less, and strictly aquifer-residing species. A six-year exploratory and citizen-science backed survey supported by molecular phylogenetic analysis reveals novel insights into the diversity, distribution and population structure of Horaglanis. The genus is characterized by high levels of intraspecific and interspecific genetic divergence, with phylogenetically distinct species recovered above a 7.0% genetic-distance threshold in the mitochondrial cytochrome oxidase subunit 1 gene. Contrasting with this deep genetic divergence, however, is a remarkable stasis in external morphology. We identify and describe a new cryptic species, Horaglanis populi, a lineage that is the sister group of all currently known species. All four species are represented by multiple haplotypes. Mismatch distribution reveals that populations have not experienced recent expansions.

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DOI: 10.3897/vz.73.e97002

Authors: Surya Narayanan, Sandeep Das, Y. Muhammed Anvar, Frank Tillack, Pratyush P. Mohapatra, David J. Gower, K. P. Rajkumar, V. Deepak

Abstract: Colour polymorphism has been previously reported in several colubrid snakes including Boiga spp. In this paper, we report colour variations within the poorly known southern Indian Boiga dightoni, provide the first molecular data for this species, from two localities (including the type locality) and compare them with data from other congeners. Additionally, we provide detailed dentition and hemipenis descriptions for B. dightoni. Molecular data for B. dightoni show very little difference (0.2–0.4% 16S; 0.9–1.2% cyt b) to the recently described Boiga whitakeri, also from southern India. We have re-examined and present new information on the pholidosis of the type specimens of B. whitakeri and reconsider its taxonomic status. On the basis of molecular data and overlapping morphological characteristics, we argue that Boiga whitakeri and Boiga dightoni are conspecific, and place B. whitakeri under the subjective synonymy of the latter. Furthermore, we show that colour polymorphism in B. dightoni is a gender-independent character and that both colour morphs are found in high as well as low elevations and partly in sympatry. A revised key to the Boiga ceylonensis complex is provided.

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DOI: 10.3897/vz.72.e94799

Authors: Ishan Agarwal, Tejas Thackeray, Akshay Khandekar

Abstract: South Asian Cnemaspis are one of the most diverse clades of gekkonids in South Asia with their highest diversity in the Western Ghats and Sri Lanka. These geckos include only a few nocturnal species and are largely diurnal or cathemeral and restricted to relatively cool habitats. One of the prominently diurnal subgroups in South Asian Cnemaspis is the bangara clade, which includes six species distributed in southern India on the eastern slopes of the Western Ghats, the southern Eastern Ghats and Palghat Gap. In this paper, we describe five more species of the bangara clade from the Shevaroyan landscape, including three from Kollimalai and one each from Yercaud and Pachaimalai, all in Tamil Nadu. These new species show 4.6–19.7 % uncorrected sequence divergence on the mitochondrial ND2 gene from each other and known species of the bangara clade and are morphologically diagnosable in body size, the number of paravertebral tubercles between limb insertions, the number of dorsal tubercle rows, the number of ventral scale rows across the belly, the number of femoral and precloacal pores and poreless scales separating these series, and aspects of colouration. The discovery of these five new species adds to the growing discoveries of cool-adapted species in southern India outside the Western Ghats and highlights the role of sky-islands in diversification. The Shevaroyan landscape shows high levels of microendemism with eight species distributed in an area of < 2000 km2, and all these species restricted to much smaller areas of actual distribution. With an area of < 500 km2 respectively, the massif of Pachaimalai has a single endemic and the massifs of Yercaud and Kollimalai have three endemic Cnemaspis species each.

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DOI: 10.3897/vz.72.e95681

Authors: Uwe Fritz, Krystal A. Tolley, Melita Vamberger, Flora Ihlow

Abstract: Species-level phylogeny and especially phylogeography of African chelonians is a comparatively under-studied field of research. We review the current knowledge of phylogeny and phylogeography, highlight congruence of spatial phylogeographic patterns amongst chelonians and other taxa and suggest future research directions to address gaps in knowledge. Our review shows that phylogeographic and phylogenetic investigations have led to unexpected findings. For example, for Pelomedusa, a putatively wide-ranging monotypic terrapin genus, cryptic diversity was revealed, with more than ten species being uncovered. The formerly recognized tortoise genus Homopus sensu lato was found to be paraphyletic with respect to Chersina. To resolve this situation, Homopus was restricted to the four-toed species H. areolatus and H. femoralis and the genus Chersobius was resurrected for the five-toed species C. boulengeri, C. signatus, and C. solus. Three previously recognized taxa were shown to be invalid, viz. the putatively extinct terrapin species Pelusios seychellensis and the tortoise subspecies Chersobius signatus cafer and Stigmochelys pardalis babcocki. Together with taxonomy, the knowledge of phylogeographic structuring sets a solid foundation for conservation measures and allows the identification of Management and Conservation Units. However, the current legislation, in particular the enforcement of the Nagoya Protocol under the Convention of Biological Diversity (CBD), has largely halted research on widely distributed taxa and turned the well-intended concept of Access and Benefit Sharing into a major impediment for conservation and research. The current situation leads for many species to a continued usage of outdated and incorrect taxonomic classifications resulting in an error cascade of conservation decisions. This is counterproductive to the aims of the CBD, that is, the protection of biodiversity. Sequencing historical DNA from museum specimens using aDNA approaches could be a short-term approach to mitigate, but not solve, this impediment.

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DOI: 10.3897/vz.72.e87275

Authors: Erasmo Cázares-Hernández, H. David Jimeno-Sevilla, Sean M. Rovito, Marco Antonio López-Luna, Luis Canseco-Márquez

Abstract: We describe a new species of plethodontid salamander of the genus Pseudoeurycea from the Sierra de Zongolica, Veracruz, Mexico. The new species is distinguished from all other species in the genus by morphological and genetic features and by coloration. Based on a mtDNA phylogeny, the new species belongs to the Pseudoeurycea juarezi group and is most closely related to P. ruficauda from the Sierra Mazateca in northern Oaxaca. The newly described salamander increases the number of species of plethodontid salamanders from Veracruz to 43 and those recognized from Mexico to 140.

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DOI: 10.3897/vz.72.e82874

Authors: Mathias Wirkner, Katharina Heyder, Irina Ruf

Abstract: The bony labyrinth (inner ear) of mammals reveals systematic as well as morphofunctional information. However, detailed knowledge of bony labyrinth morphology and ontogeny in Pantherinae, that comprise some of the most iconic mammals, is still pending. Hence, we present the first comparative description of the bony labyrinth in all extant species of Panthera and Neofelis some of which are represented by several postnatal stages; particular focus is set on Panthera leo. Our study is based on µCT scans and virtual 3D reconstructions and accompanied by selected morphometric measurements. Even though quite similar in morphology, both genera as well as their species can be distinguished by several features, e.g., shape and relative size of the semicircular canals and presence or absence of an osseous secondary crus commune. In case of the latter, P. pardus shows some intraspecific variation. We also traced the reduction of the fossa subarcuata during ontogeny in P. leo which conforms with previous studies. Negative allometry of the bony labyrinth in relation to skull basal length can be observed during ontogeny as demonstrated by P. leo as well as between different sized species. Although not correlated with the length of the cochlear canal, the number of cochlear turns is higher in captive non-adult P. leo and P. tigris, but lower in adult captive P. pardus. If these intraspecific differences are related to captivity or represent an ontogenetic pattern, needs to be evaluated in future studies based on larger samples.

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DOI: 10.3897/vz.72.e89324

Authors: Surya Narayanan, Aravind NA

Abstract: We describe a new species of small-sized (SVL 24.6–28.8 mm) rupicolous Cnemaspis from the Biligirirangan Hills (BR Hills), Karnataka based on morphology and molecular (mitochondrial 16S and ND2) data. The new species is nested within the monticola, mysoriensis and gracilis clades, where it is basal to monticola and gracilis clades in the Maximum Likelihood analysis and is basal to the monticola clade in the Bayesian phylogenetic analysis.

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DOI: 10.3897/vz.72.e86348

Authors: Omar Rafael Regalado Fernández, Ingmar Werneburg

Abstract: A literature review showed that there is not a defined consensus on what specimens belong to Plateosaurus in current phylogenetic analyses, and after the assignation of SMNS 13200 as the neotype for Plateosaurus, the specimen composition of Plateosaurus as an operational taxonomic unit (OTU) needs to be addressed in further iterations of phylogenetic analyses. At least one of the specimens used to illustrate plateosaurian anatomy contains several characters identified in more derived sauropodomorphs commonly referred to as massopodans. This partial skeleton, traditionally known as specimen ‘GPIT IV’, was found in the lower dinosaur bone bed of the Obere Mühle, a Trossingen Formation outcrop, during an excavation in 1922 near the city of Tübingen, Germany. The holotype of Plateosaurus trossingensis and several other specimens referred to as this species were found in this level, which was initially interpreted as a synchronic deposit of animals. However, the current understanding of the Trossingen Formation indicates that this bed was probably a constant accumulation of carcasses through miring and transport down a river for hundreds of years. In this work, a framework to compare phylogenetic signals with morphological and histological data is provided to help in the species delineation of Plateosaurus, and support is found to refer the historic specimen ‘GPIT IV’ as a new genus and a new species.

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DOI: 10.3897/vz.72.e85269

Authors: Mariana P. Marques, Luis M. P. Ceríaco, Matthew P. Heinicke, Rachal M. Chehouri, Werner Conradie, Krystal A. Tolley, Aaron M. Bauer

Abstract: The genus Heliobolus comprises four recognized species, all endemic to sub-Saharan Africa. Of these, only Heliobolus lugubris occurs in southern Africa, its distribution extending from Angola in the west to Mozambique in the east and reaching as far south as parts of northern South Africa. Like many of the reptile species that occur in southern Africa, Heliobolus lugubris is poorly studied, and preliminary investigation suggested that it may contain cryptic diversity. The present work focusses on the Angolan population of H. lugubris and uses an integrative taxonomic approach based on morphological, coloration and DNA sequence data. The results indicate that some of the current and historical specimens of H. lugubris from Angola do not correspond to the nominotypical form, and that differences between specimens suggest the presence of two additional species, described here as Heliobolus bivari sp. nov. from the southernmost xeric/desertic regions and plateau of Namibe Province, southwestern Angola and H. crawfordi sp. nov. from the Serra da Neve inselberg north through the sub-desert coastal regions of northern Namibe, Benguela, and Kwanza Sul provinces. Nominotypical Heliobolus lugubris is confirmed to occur in Cuando Cubango Province, southeastern Angola.

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DOI: 10.3897/vz.72.e89660

Authors: Surya Narayanan, Sandeep Das, Amirtha Balan, Roshin Tom, Nitin Divakar, Rajkumar Kp, P. Hopeland, V. Deepak

Abstract: A new species of Cyrtodactylus is described from the southern Western Ghats of India. It is distinguished from all species of the ­Cyrtodactylus collegalensis species complex in colour pattern and resembles the Srilankan endemic C. yakhuna in overall colouration. Phylogenetic analysis of mitochondrial ND2 DNA sequences indicates that the new species is most closely related to species in the C. collegalensis complex and differs from them by an uncorrected pairwise genetic distance of 10.5–12.9%.

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DOI: 10.3897/vz.72.e80019

Authors: Zhi-Tong Lyu, Jian Wang, Zhao-Chi Zeng, Lin Luo, Yan-Wu Zhang, Chun-Peng Guo, Jin-Long Ren, Shuo Qi, Yun-Ming Mo, Ying-Yong Wang

Abstract: The recognition for the floating frogs’ genus Occidozyga is in controversy for decades, and the species diversity of these frogs has recently been considered to be underestimated. In southeastern China, two floating frog species are currently recorded, namely Occidozyga lima and Occidozyga martensii. However, their current taxonomic statuses are unresolved after a series of recent taxonomic revisions. In this work, we perform morphological examinations and phylogenetic analyses on these two recorded floating frogs from southeastern China, to clarify their current taxonomic placements. The population previously recorded as Occidozyga lima should be re-assigned to the nomenclature Occidozyga obscura comb. nov., and the population previously recorded as Occidozyga martensii should be an undescribed species which is erected as Occidozyga lingnanica sp. nov. in this work.

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DOI: 10.3897/vz.72.e83544

Authors: Sirpa Nummela, Gabriel Aguirre-Fernández, Kathleen K. Smith, Marcelo R. Sánchez-Villagra

Abstract: We studied the mass growth trajectories of middle ear ossicles and tympanic membrane and oval window area in 19 specimens of postnatal ages 30–180 days of the gray short-tailed opossum Monodelphis domestica. We weighed the skull mass and the mass of the three middle ear ossicles with appropriate balances. Using a binocular microscope provided with a grid, we measured the length of malleus and incus, as well as the longest axis and the one perpendicular to it on both the tympanic membrane and the stapes footplate. The size variation was studied with least squares regression analyses between various measurements. The incus and stapes change little in mass after 40 days of postnatal life, while the malleus does, reaching maximum mass at around 100 PND (postnatal days). This modularity in growth trajectory is in contrast with the shared evolutionary origin of malleus and incus from branchial arch 1. The maturation of the middle – and as indicated by previous work, that of the inner ear – is coupled with the improvement of hearing sensitivity at low and high frequencies after the initial onset of hearing at 29 PND.

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DOI: 10.3897/vz.72.e82709

Authors: M. Judith Babot, Guillermo W. Rougier, Daniel A. García-López, Sara B. Bertelli, Claudia M. Herrera, M. Virginia Deraco, Norberto P. Giannini

Abstract: We present a detailed description of the anatomy of the dentary and lower teeth of a new specimen of Callistoe vincei, a large carnivorous metatherian from the Eocene (?Ypresian) of northwestern Argentina. The recently collected specimen is a young adult represented by a partial right dentary with the canine, p1, roots of p3, and very well-preserved m1 to m4. The description includes a comparison with the holotype specimen, a much older individual, and other closely related large sparassodonts (e.g., Arminiheringia). The analysis of this new material allowed identifying plesiomorphic molar features in Callistoe, such as the presence of a reduced metaconid on the m3 and a tricuspated, basined talonid on m1–m3. We also described the mesowear facets in the lower dentition, showing that the self-sharpening facet typically present in extinct and extant placental and some marsupial carnivorous forms, was absent in Callistoe. The presence of a short-term cutting edge in the trigonid related to the thinness of the enamel layer, and the associated tooth wear susceptibility, were likely compensated by a dental mechanism (overeruption) to maintain occlusal contact among antagonist teeth. This process could explain the marked extrusion of the tooth roots observed in Callistoe as well as in other large closely related members of the group.

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DOI: 10.3897/vz.72.e85020

Authors: Ananthanarayanan Sankar, Ingg Thong Law, Ing Sind Law, Rasu Shivaram, Robin K. Abraham, Kin Onn Chan

Abstract: The genus Micryletta, also known as paddy frogs, ranges across much of south, east, and southeast Asia. Due to their relatively broad distribution and overall morphological similarities, many species have gone undetected until recently, largely owing to the use of molecular data. Consequently, the species diversity within this genus has quadrupled in just three years from three species prior to 2018, to 12 species in 2021, indicating that the systematics of this genus is still poorly understood. As such, we assembled the most comprehensive molecular phylogeny of Micryletta hitherto including novel sequences from a previously unsampled population from Singapore to assess the species diversity within this genus. In particular, we investigate the population from Singapore whose specific identity remains in question due to the lack of voucher specimens and genetic material. Our results show that the Singapore population represents a strongly supported and distinct lineage that is most closely related to M. inornata sensu stricto from Sumatra, Indonesia. Morphological and species delimitation analyses corroborate its distinction as a new species, which we describe herein as M. subaraji sp. nov. This and recent new taxon discoveries in Singapore demonstrate that the biodiversity of the highly urbanized island-state is still far from being fully realized and underscores the need for continued systematic surveys and protection of remaining habitats.

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DOI: 10.3897/vz.72.e84516

Authors: Ho Yuen Yeung, Michael W. N. Lau, Jian-Huan Yang

Abstract: A new species of the genus Calamaria Boie, 1827, Calamaria arcana sp. nov., is described based on a single male specimen collected from Mt. Dadongshan, Guangdong, southern China. The new species can be distinguished from all known congeners by the significant genetic divergence in the mitochondrial cytochrome-b gene fragment (p-distance ≥ 13.9%), and morphologically by the combination of the following characters: (1) ten modified maxillary teeth; (2) four supralabials, second and third supralabials entering orbit; (3) preocular present; (4) mental not touching anterior chin shields; (5) six scales and shields surrounding the paraparietal; (6) 170 ventral scales; (7) 22 paired subcaudals; (8) tail not gradually tapering, abruptly tapering at the tip; (9) dorsal scales reduced to five rows above last subcaudal at tail; (10) dorsum of body and tail brownish; (11) dark collar on nuchal region absent; (12) two outermost dorsal scale rows light yellow with upper margins partly dark pigmented; (13) ventral scales immaculate, without dark outermost corners and pigmentation anteriorly; and (14) absence of distinct dark longitudinal line or scattered spots on the underside of tail. Calamaria arcana sp. nov., represents the fifth species of the genus recorded in China. Following the IUCN Red List Categories and Criteria, we propose the new species to be listed as Data Deficient.

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DOI: 10.3897/vz.72.e83324

Authors: Irina Ruf

Abstract: The turbinal skeleton inside the nasal cavity supports the respiratory and olfactory epithelia of the mammalian nose and can provide systematic and morphofunctional information. For the first time, the turbinal skeleton of Pentalagus furnessi (Amami rabbit) from Japan is described based on µCT scans and virtual 3D reconstructions of two specimens. In general, the turbinal skeleton of Pentalagus furnessi resembles the pattern and characters observed in other Leporidae. The maxilloturbinal is highly dendritic, nasoturbinal and crista semicircularis are in close contact and form a common recess, the frontoturbinal recess houses two frontoturbinals and one interturbinal between them, the ethmoturbinal recess houses three ethmoturbinals and one interturbinal between ethmoturbinal I and II. Pentalagus furnessi is derived from the leporid grundplan in having a lamina semicircularis with almost straight posterior margin and ventral lamella and in showing a single-scrolled and relatively short interturbinal between frontoturbinal 1 and 2. These characters can be regarded as autapomorphic for the Amami rabbit. Furthermore, the two specimens have an additional small and short interturbinal between frontoturbinal 2 and ethmoturbinal I that shows some variation. This pattern supports previous observations of intraspecific variation of certain interturbinals in Oryctolagus cuniculus and some Sylvilagus and Lepus species. The comparison of the turbinal skeleton of Pentalagus furnessi and its possible sister taxon (e.g., Pronolagus, Poelagus or Caprolagus) reveals a puzzling pattern which is discussed.

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DOI: 10.3897/vz.72.e82201

Authors: Anh The Nguyen, Tang Van Duong, Perry L. Wood Jr., L. Lee Grismer

Abstract: An integrative taxonomic analysis of species in the colubrid genus Lycodon Fitzinger, 1826 recovered two new syntopic species of the L. rufozonatus complex from the imperiled Song Giang River valley in Khan Hoa Province, of Southern Vietnam. Although L. truongi sp. nov. and L. anakradaya sp. nov. are syntopic, they are not particularly closely related and can be differentiated from each other and all other species in the L. rufozonatus complex on the basis of meristics, morphometrics, color pattern, and uncorrected pairwise genetic distance based on the mitochondrial gene cytochrome b. The discovery of these two new range-restricted species and a previously described range-restricted gekkonid in the genus Cyrtodactylus Gray, 1828 from the same valley, underscores the necessity of continued field work in the Song Giang River valley so as to catalog the unrealized herpetological diversity in this area and establish research-based conservation programs.

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DOI: 10.3897/vz.72.e80615

Authors: L. Lee Grismer, Attapol Rujirawan, Siriporn Yodthong, Bryan L. Stuart, Minh Duc Le, Dzung Trung Le, Yodchaiy Chuaynkern, Perry L. Wood, Jr., Anchalee Aowphol

Abstract: Abstract Convergent morphological specializations for an arboreal lifestyle in most species of the Cyrtodactylus brevipalmatus group have been a confounding factor for establishing a stable taxonomy among its species. Recent references to C. interdigitalis from throughout Thailand and Laos were made without comparisons to the type material from Tham Yai Nam Nao, Nam Nao National Park, Phetchabun Province, Thailand, but instead, were based on general morphological similarity and distribution. The taxonomy of C. interdigitalis is stabilized here by comparing the paratypes to other specimens from Thailand and Laos and recovering their phylogenetic relationships based on newly acquired genetic data, including those from the type locality. The phylogeny recovered all specimens outside the type locality to be either C. ngati from Vietnam or new species closely related to C. ngati. Cyrtodactylus interdigitalis is shown here to be a range-restricted upland endemic on the Phetchabun massif of northern Thailand. The phylogeny also indicates that C. ngati extends hundreds of kilometers farther south into northern Thailand and central Laos. We hypothesize that the significant morphological divergence in body shape of the types of C. ngati, compared to that of the Lao and Thai populations, may be due to local adaptions for utilizing karst (C. ngati) rather than vegetation (Lao and Thai populations). Additionally, phylogenetic and multivariate analyses identified a potentially new species from Phu Hin Rong Kla National Park, Phitsanulok Province, in northern Thailand and another from the Khlong Naka Wildlife Sanctuary, Ranong Province, in southern Thailand. A series of newly examined specimens from Kaeng Krachan National Park, Phetchaburi Province, Thailand represents a possible ~82 km range extension to the southeast of C. rukhadeva. This research continues to underscore the high diversity of range-restricted upland endemics in Thailand and the importance of examining type material (if possible) in the context of a phylogeny so as to construct proper taxonomies that reveal, rather than obscure, diversity.

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DOI: 10.3897/vz.72.e78219

Authors: Thomas E. Lee Jr., Nicolás Tinoco, Jorge Brito

Abstract: We name and describe a new species of Andean mouse from the eastern slope of the Andes of central Ecuador (Sangay National Park). This rodent is large-bodied (head-body length 167–184 mm) inhabiting the wet montane forest between 3,400–3,900 m in elevation. A molecular phylogeny based on mitochondrial genes resolved the new species as a member of the “aureus” group, closely related to an undescribed species from north Ecuador. This finding increases the diversity of Thomasomys to 48 species, of which 18 species inhabit Ecuador. In addition, the species described herein is the largest species of the genus described in Ecuador.

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DOI: 10.3897/vz.72.e79496

Authors: Sonali Garg, Sivaperuman Chandrakasan, G. Gokulakrishnan, C. Gopika, Indraneil Das, S. D. Biju

Abstract: Since the description of Charles Darwin’s frog as Rana charlesdarwini in 1998, its generic placement has been a taxonomic enigma. Subsequent studies first transferred this species to the dicroglossid genus Limnonectes, and then considered it as a ceratobatrachid of the genus Ingerana, which has since been moved to the family Dicroglossidae. However, recent works have doubted this generic placement and also suggested the possibility of its sister relationship with the genus Liurana, within Ceratobatrachidae. Nonetheless, there have been no detailed investigations to ascertain the generic placement of this taxon by confirming its phylogenetic position or using integrative taxonomic approaches. Here, we provide the first molecular assessment of Ingerana charlesdarwini based on mitochondrial and nuclear DNA and reveal that it is nested in the dicroglossid genus Minervarya. A member of the Minervarya andamanensis species group, Minervarya charlesdarwini comb. nov. is sister taxon to M. andamanensis and shows relatively shallow genetic distances (2.8–3.6%) in the 16S gene. Both species are widely distributed, occur sympatrically, and exhibit high morphological variations, leading to long-standing confusions with other dicroglossid frogs reported from the region. Our combined morphological and molecular studies on dicroglossid frogs sampled across the known ranges of these species suggest that reports of Limnonectes doriae (Boulenger, 1887) and L. hascheanus (Stoliczka, 1870) from the Andamans are misidentifications of the former two, pointing to the absence of genus Limnonectes from the Andaman Islands. Our study also reveals the novel record of Minervarya agricola from the Andamans, a species that appears to have been confused with Fejervarya limnocharis and Minervarya keralensis in the literature and misidentified museum specimens, and is found to be widely distributed across these islands. We further find another congener from the Nicobar group of Islands, M. nicobariensis, to be closely related to M. charlesdarwini. Similar to the case of Andaman dicroglossids, our work emphasises on the need for further studies to ascertain the taxonomic identities and generic placement of Minervarya and Limnonectes species reported from the Nicobars.

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DOI: 10.3897/vz.72.e76308

Authors: Akshay Khandekar, Tejas Thackeray, Ishan Agarwal

Abstract: We describe three new small-bodied, cryptic species of south Asian Cnemaspis belonging to the mysoriensis and goaensis clades from the Mysore Plateau and the Western Ghats biodiversity hotspot in Karnataka, peninsular India; and provide a key to members of each clade, besides providing the first ND2 sequence data for C. ranganaensis. Cnemaspis tigris sp. nov. from Kaiwara in Karnataka is a member of the mysoriensis clade and can be morphologically distinguished from all six described members of the clade in a number of meristic characters and subtle differences in colouration, beside ≥ 12.1–23.4 % uncorrected pairwise ND2 sequence divergence. Cnemaspis sakleshpurensis sp. nov. from Sakleshpur and C. vijayae sp. nov. from Coorg, both in the Western Ghats of Karnataka, are members of the goaensis clade and are easily diagnosed from all three described members of the clade in meristic characters beside 5.2–14.8 % divergence from described members of the clade and 14.6 % from each other in uncorrected pairwise ND2 sequence divergence. The discovery of these new species from two different clades and biogeographic regions is not surprising, given the steep rise in the number of species of Cnemaspis known from peninsular India. Comprehensive geographic sampling in conjunction with molecular and morphological data is essential to understand the true diversity and distributional ranges of species within this ancient clade of gekkonid lizards.

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DOI: 10.3897/vz.72.e76047

Authors: Timothy D. Smith, Christopher J. Bonar

Abstract: Abstract Nasal anatomy in rodents is well-studied, but most current knowledge is based on small-bodied muroid species. Nasal anatomy and histology of hystricognaths, the largest living rodents, remains poorly understood. Here, we describe the nasal cavity of agoutis (Dasyprocta spp.), the first large-bodied South American rodents to be studied histologically throughout the nasal cavity. Two adult agoutis were studied using microcomputed tomography, and in one of these, half the snout was serially sectioned and stained for microscopic study. Certain features are notable in Dasyprocta. The frontal recess has five turbinals within it, the most in this space compared to other rodents that have been studied. The nasoturbinal is particularly large in dorsoventral and rostrocaudal dimensions and is entirely non-olfactory in function, in apparent contrast to known muroids. Whether this relates solely to body size scaling or perhaps also relates to directing airflow or conditioning inspired air requires further study. In addition, olfactory epithelium appears more restricted to the olfactory and frontal recesses compared to muroids. At the same time, the rostral tips of the olfactory turbinals bear at least some non-olfactory epithelium. The findings of this study support the hypothesis that turbinals are multifunctional structures, indicating investigators should use caution when categorizing turbinals as specialized for one function (e.g., olfaction or respiratory air-conditioning). Caution may be especially appropriate in the case of large-bodied mammals, in which the different scaling characteristics of respiratory and olfactory mucosa result in relative more of the former type as body size increases.

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DOI: 10.3897/vz.72.e76046

Authors: Sandeep Das, Saunak Pal, Sasidharan Siddharth, Muhamed Jafer Palot, Veerappan Deepak, Surya Narayanan

Abstract: Abstract A new large (94–107mm SVL) species of gecko of the genus Hemidactylus is described from the drier parts of the Western Ghats of India. The new species is closely related to H. graniticolus, from which it can be distinguished based on dorsal pholidosis at mid-body, the structure of tubercles on the dorsum, dorsal pholidosis on the tail. The new species is also 6.6–7.2% divergent from H. graniticolus in the ND2 mitochondrial gene.

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DOI: 10.3897/vz.72.e77702

Authors: L. Lee Grismer, Lelani del Pinto, Evan S. H. Quah, Shahrul Anuar, Michael Cota, Jimmy A. McGuire, Djoko T. Iskandar, Perry L. Wood Jr, Jesse L. Grismer

Abstract: Abstract An integrative taxonomic analysis of Sundaic populations of Gekko smithii from the Thai-Malaya Peninsula, Sumatra, and Borneo recovered four deeply divergent mitochondrial lineages that are separated by major geographic barriers (mountains and seaways). Furthermore, they bear a number of concordant statistically significant differences in meristic and morphometric features, morphospatial separation in multivariate space, and discrete differences in color pattern. Gekko smithii sensu stricto is restricted to southern Thailand south of the Isthmus of Kra and Peninsular Malaysia west of the Banjaran (mountain range) Titiwangsa, being that the type locality is on Penang Island, Penang. Gekko hulk sp. nov. is a new species from extreme southern Thailand and Peninsular Malaysia east of the Banjaran Titiwangsa and five east coast islands—the type locality being Pulau (island) Tioman, Pahang. Gekko cf. albofasciolatus is tentatively used to include Bornean populations west of the Iran Mountains in Sabah and Sarawak which, in the absence of molecular data, cannot unequivocally be separated morphologically from G. albofasciolatus from the type locality at Banjarmasin, Kalimantan, Indonesia east of the Iran Mountains. In the absence of molecular data, G. albomaculatus is resurrected to include mainland Sumatran, Nias Island, and Banyak Islands populations which, based on their morphology, cannot be separated from descriptions of G. albomaculatus from the type locality of Bangka Island, 15 km off the southeast coast of mainland Sumatra. Further integrative analyses of all Sumatran and Bornean populations are currently underway as well as the enigmatic Wallacean populations from Sulawesi. Data are presented that strongly suggest all references to G. smithii from Java stem from a 151 year-old misidentification of a specimen of G. gecko of unknown provenance. Additionally, there are no vouchered records of G. smithii from Myanmar. The phylogeographic patterns of Sundaic populations of the G. smithii complex are concordant with those of a plethora of other Sundaic lineages.

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DOI: 10.3897/vz.71.e76453

Authors: Marika Asztalos, Dinçer Ayaz, Yusuf Bayrakcı, Murat Afsar, Cemal Varol Tok, Carolin Kindler, Daniel Jablonski, Uwe Fritz

Abstract: Using two mitochondrial DNA fragments and 13 microsatellite loci, we examined the phylogeographic structure and taxonomy of two codistributed snake species (Natrix natrix, N. tessellata) in their eastern distribution area, with a focus on Turkey. We found evidence for frequent interspecific hybridization, previously thought to be extremely rare, and for backcrosses. This underscores that closely related sympatric species should be studied together because otherwise the signal of hybridization will be missed. Furthermore, the phylogeographic patterns of the two species show many parallels, suggestive of a shared biogeographic history. In general, the phylogeographies follow the paradigm of southern richness to northern purity, but the dice snake has some additional lineages in the south and east in regions where grass snakes do not occur. For both species, the Balkan Peninsula and the Caucasus region served as glacial refugia, with several mitochondrial lineages occurring in close proximity. Our results show that the mitochondrial divergences in both species match nuclear genomic differentiation. Yet, in the former glacial refugia of grass snakes there are fewer nuclear clusters than mitochondrial lineages, suggesting that Holocene range expansions transformed the glacial hotspots in melting pots where only the mitochondrial lineages persisted, bearing witness of former diversity. On the other hand, the deep mitochondrial divergences in N. tessellata across its entire range indicate that more than one species could be involved, even though lacking microsatellite data outside of Turkey prevent firm conclusions. On the contrary, our microsatellite and mitochondrial data corroborate that N. megalocephala is invalid and not differentiated from sympatric populations of N. natrix. For Cypriot grass snakes, our analyses yielded conflicting results. A critical assessment of the available evidence suggests that N. natrix is a genetically impoverished recent invader on Cyprus and taxonomically not distinct from a subspecies also occurring in western Anatolia and the southern Balkans. Based on combined mitochondrial and nuclear genomic evidence we propose that for grass snakes the following subspecies should be recognized in our study region: (1) Natrix natrix vulgaris Laurenti, 1768, southeastern Central Europe and northern Balkans; (2) Natrix natrix moreotica (Bedriaga, 1882), southern Balkans, western Anatolia, and Cyprus; and (3) Natrix natrix scutata (Pallas, 1771), eastern Anatolia, Caucasus region, Iran, northeastern distribution range (from eastern Poland and Finland to Kazakhstan and the Lake Baikal region). Thus, Natrix natrix cypriaca (Hecht, 1930) becomes a junior synonym of N. n. moreotica and Natrix natrix persa (Pallas, 1814) becomes a junior synonym of N. n. scutata. Due to insufficient material, we could not resolve the status of Natrix natrix syriaca (Hecht, 1930) from the Gulf of İskenderun, southeastern Turkey.

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DOI: 10.3897/vz.71.e73529

Authors: Chatmongkon Suwannapoom, L. Lee Grismer, Parinya Pawangkhanant, Mali Naiduangchan, Platon V. Yushchenko, Dmitriy V. Arkhipov, Jeffery A. Wilkinson, Nikolay A. Poyarkov

Abstract: Abstract The integrated results of morphological and molecular phylogenetic analyses confirmed the new species status of a recently discovered population of Ansonia from Suan Phueng District, Ratchaburi Province, Thailand. Ansonia karen sp. nov. is separated from all other species of Ansonia by a unique combination of mensural, discrete morphological, and color pattern characteristics and is the sister species of A. thinthinae from Tanintharyi Division, Myanmar. This discovery fills a geographic hiatus of 350 km between it and A. kraensis from Ranong Province, Thailand. Ansonia karen sp. nov. is the newest member of a long list of range-restricted endemics having been recently discovered in the northern Tenasserim Mountain region of western Thailand and continues to underscore the unexplored nature of this region and its need for conservation.

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DOI: 10.3897/vz.71.e71995

Authors: Gustavo Campillo-García, Oscar Flores-Villela, Brett Oliver Butler, Julián Andrés Velasco Vinasco, Fabiola Ramírez Corona

Abstract: The spiny lizard genus Sceloporus was described by Wiegmann in 1828, with S. torquatus posteriorly designated as the type species. The taxonomic history of S. torquatus is complicated, as it has been confused with other taxa by numerous authors. Many modern systematics works have been published on Sceloporus, but none have included all five recognized S. torquatus subspecies: S. t. torquatus, S. t. melanogaster, S. t. binocularis, S. t. mikeprestoni, and S. t. madrensis. Additionally, there is previous evidence for at least one unnamed taxon. The present study is the first taxonomic revision of the enigmatic S. torquatus based on molecular phylogenies using combined molecular data from 12S, ND4 and RAG1 genes, and Maximum Likelihood and Bayesian inference phylogenetic methods. This work includes the most extensive sampling across the entire distribution, as well as divergence time estimates and environmental niche modelling, which combined offer a spatio-temporal framework for understanding the evolution of the species. Additionally, a series of morphological characters are analyzed to identify significant differences between lineages consistently recovered in the molecular phylogenies. Using this integrative approach, evidence is presented for eight lineages within the S. torquatus complex, five of which correspond to previously recognized subspecies and three represent unnamed taxa masked by morphological conservatism. Finally, to maintain taxonomic stability a lectotype and paralectoype are designated for S. torquatus, and certain taxonomic changes are suggested in order to reflect the phylogenetic relationships within the S. torquatus complex.

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DOI: 10.3897/vz.71.e75967

Authors: V. Deepak, Samuel Lalronunga, Esther Lalhmingliani, Abhijit Das, Surya Narayanan, Indraneil Das, David J. Gower

Abstract: Xenodermidae is a generally poorly known lineage of caenophidian snakes found in South, East and Southeast Asia. We report molecular phylogenetic analyses for a multilocus data set comprising all five currently recognised genera and including new mitochondrial and nuclear gene sequence data for the recently described Stoliczkia vanhnuailianai. Our phylogenetic results provide very strong support for the non-monophyly of Stoliczkia, as presently constituted, with S. borneensis being more closely related to Xenodermus than to the Northeast Indian S. vanhnuailianai. Based on phylogenetic relationships and morphological distinctiveness, we transfer Stoliczkia borneensis to a new monotypic genus endemic to Borneo, Paraxenodermus gen. nov. We also present new morphological data for P. borneensis.

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DOI: 10.3897/vz.71.e76069

Authors: L. Lee Grismer, Chatmongkon Suwannapoom, Parinya Pawangkhanant, Roman A. Nazarov, Platon V. Yushchenko, Mali Naiduangchan, Minh Duc Le, Vinh Quang Luu, Nikolay A. Poyarkov

Abstract: The first integrative taxonomic analysis of the Cyrtodactylus brevipalmatus group of Southeast Asia recovered two newly discovered populations from the Tenasserim Mountains in Suan Phueng District, Ratchaburi Province, Thailand as a new species described here as C. rukhadeva sp. nov. Based on 1397 base pairs of the mitochondrial gene NADH dehydrogenase subunit 2 (ND2), C. rukhadeva sp. nov. is the well-supported sister species to a clade containing three undescribed species, C. ngati, and C. cf. interdigitalis with a large uncorrected pairwise sequence divergence from other species in the brevipalmatus group ranging from 15.4–22.1%. Cyrtodactylus elok and C. brevipalmatus are recovered as poorly supported sister species and the well-supported sister lineage to the remainder of the brevipalmatus group. Cyrtodactylus rukhadeva sp. nov. is putatively diagnosable on the basis of a number of meristic characters and easily separated from the remaining species of the brevipalmatus group by a number of discrete morphological characters as well as its statistically significant wide separation in multivariate morphospace. The discovery of C. rukhadeva sp. nov. continues to underscore the unrealized herpetological diversity in the upland forests of the Tenasserim Mountains and that additional field work will undoubtedly result in the discovery of additional new species.

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DOI: 10.3897/vz.71.e73563

Authors: Zhi-Tong Lyu, Jian Wang, Zhao-Chi Zeng, Jia-Jun Zhou, Shuo Qi, Han Wan, You-Yu Li, Ying-Yong Wang

Abstract: In this work, a new species of the genus Tylototriton is described from Guangdong, southern China. Tylototriton sini sp. nov. was recorded as T. asperrimus for decades, and was indicated to represent an independent lineage based on recent molecular phylogenetic analyses. After detailed molecular analysis and morphological comparisons, Tylototriton sini sp. nov. is recognized as a distinct species which can be clearly distinguished from all known congeners by a combination of morphological characteristics and the significant divergence in the mitochondrial gene. Because the genus Tylototriton is of high conservation concern and all formally described members are protected by law, we also provide first data on the conservation status and recommendations for IUCN categorization for Tylototriton sini sp. nov. A suggestion on the species groups division of the genus Tylototriton is also provided based on their morphological differences and phylogenetic relationships.

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DOI: 10.3897/vz.71.e62787

Authors: Gaurang Gowande, Saunak Pal, Daniel Jablonski, Rafaqat Masroor, Pushkar U. Phansalkar, Princia Dsouza, Aditi Jayarajan, Kartik Shanker

Abstract: Abstract The genus Calotes Cuvier, 1817 (Agamidae: Draconinae) is highly diverse, with species occurring in South and Southeast Asia, and Oceania. Most species of the subfamily except C. versicolor have narrow geographic distributions. Calotes versicolor is distributed from western Iran in the west to south China and Indonesia in the east and has been introduced to parts of Africa and North America. The species has had a complicated taxonomic history; multiple species and subspecies related to C. versicolor were described from India and adjoining regions, which were synonymized in subsequent revisions. However, a study of Burmese C. versicolor yielded two new species, C. htunwini and C. irawadi, indicating that C. versicolor is a species complex. Such integrative taxonomic studies have not been carried out in India, the supposed type locality of C. versicolor. Hence, we studied C. versicolor sensu lato from the Indian subcontinent and generated sequences of mitochondrial 16S and COI fragments from tissues sampled from multiple localities in the region, including the type localities of its synonyms. Phylogenetic analyses revealed four well-supported, deeply-divergent lineages, supported by morphological data. These lineages represent (i) C. versicolor sensu stricto, from South India and parts of the east coast, (ii) C. irawadi sensu lato from northeast India and Southeast Asia, (iii) a synonym from the eastern Indo-Gangetic Plains which we resurrect here, and (iv) a subspecies from Pakistan which we elevate to species level. We provide re-descriptions for the resurrected or elevated species, and a diagnostic key to the species of the C. versicolor complex. The study shows that C. versicolor sensu stricto is endemic to parts of southern and eastern India, and not widely distributed, though it may have been introduced to other parts of the world.

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DOI: 10.3897/vz.71.e71886

Authors: George M. T. Mattox, Kevin W. Conway

Abstract: Miniaturization, the evolution of extremely small adult body size, is a common phenomenon across the lineages of freshwater fishes, especially in the Neotropics where over 200 species are considered miniature (≤26 mm in standard length [SL]). Close to 30% of all miniature Neotropical freshwater fishes belong to the family Characidae, several of which are of uncertain phylogenetic placement within the family. We investigate the skeletal anatomy of Tucanoichthys tucano, a species of uncertain phylogenetic position from the upper Rio Negro basin, reaching a maximum known size of 16.6 mm SL. The skeleton of Tucanoichthys is characterized by the complete absence of ten skeletal elements and marked reduction in size and/or complexity of others, especially those elements associated with the cephalic latero-sensory canal system. Missing elements in the skeleton of Tucanoichthys include those that develop relatively late in the ossification sequence of the non-miniature characiform Salminus brasiliensis, suggesting that their absence in Tucanoichthys can be explained by a simple scenario of developmental truncation. A number of the reductions in the skeleton of Tucanoichthys are shared with other miniature characiforms, most notably species of Priocharax and Tyttobrycon, the latter a putative close relative of Tucanoichthys based on molecular data.

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DOI: 10.3897/vz.71.e66239

Authors: Ashok Kumar Mallik, Achyuthan Needamangalam Srikanthan, Sumaithangi Rajagopalan Ganesh, Seenapuram Palaniswamy Vijayakumar, Patrick D. Campbell, Anita Malhotra, Kartik Shanker

Abstract: Abstract Asian pit vipers belonging to the genus Craspedocephalus are a complex group of vipers, distributed in South and Southeast Asia. Their taxonomy is unresolved in many lineages across their distributional range. Here, we reassess the taxonomy and systematics of pit vipers of the genus Craspedocephalus in Peninsular India based on extensive field sampling, in particular in the Western Ghats. We build and expand on the previous findings of genetic relatedness between the peninsular Indian lineages with the Sundaic clade (C. puniceus complex) with greater evidence, based on additional taxa sequenced herein. We reconstruct the phylogeny of the group using three mitochondrial genes and delineated lineages using coalescent species delimitation methods. We then used multiple criteria including genetic divergence and separation in morphological and geographic space to designate taxonomic units. Our work revealed the presence of a South Asian radiation of the clade Craspedocephalus, with a few Sundaic members. Our study reveals the systematic relationships of four Peninsular Indian species of Craspedocephalus, including Peltopelor macrolepis and C. strigatus, sequenced here for the first time, that are classified or confirmed as members of Craspedocephalus. Hence, we place the genus Peltopelor in the synonymy of Craspedocephalus. Using our multi-criteria approach, we delimit four new cryptic evolutionary lineages within the Western Ghats escarpment of Peninsular India. These cryptic lineages belong to the C. malabaricus, C. gramineus and C. macrolepis complexes and are geographically and/or ecologically (in terms of habitat association) distinct from their sister lineages across their distributional range, while others are separated in morphological space. Our new phylogenetic tree and delimitation analysis thus reveals the presence of multiple clades with several cryptic lineages separated by geographical barriers or habitat association.

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DOI: 10.3897/vz.71.e62814

Authors: Pradeep Kumkar, Manoj Pise, Pankaj A. Gorule, Chandani R. Verma, Lukáš Kalous

Abstract: The hill stream loach genus Indoreonectes is endemic to peninsular India south of the Satpura hill ranges and is represented by three species I. evezardi, I. keralensis and I. telanganaensis. Indoreonectes evezardi has been suggested as a species complex based on recent genetic studies; however, due to lack of type material the species delimitation has been difficult. Here we redescribe I. evezardi collected from its type locality and describe two new species from the northern Western Ghats of India. Indoreonectes neeleshi, described from Mula River tributary of Godavari river system, can be diagnosed from all its congeners based on a combination of characters: inner rostral barbel reaching middle of nostril; maxillary barbel reaching midway between eye and posterior border of operculum; dorsal hump behind nape; bars on lateral side of the body wider than inter-bar space; total vertebrae 35 and dorsal fin insertion between 13th and 14th abdominal vertebrae. Indoreonectes rajeevi, described from Hiranyakeshi River of the Krishna river system, differs from all its congeners based on a combination of characters: inner rostral barbel reaching anterior margin of eye; maxillary barbel reaching posterior border of operculum; conspicuous black markings on lower lip, dorsal hump absent; total vertebrae 36 and dorsal fin insertion between 12th and 13th abdominal vertebrae. Further, I. neeleshi differs from its congeners by the raw genetic distance of 6.8–14.4% for the cox1 gene and 5.7–16.2% for the cytb gene, while I. rajeevi differs from its congeners by the raw genetic distance of 10.9–14.0% for the cox1 gene and 11.8–15.8% for the cytb gene.

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DOI: 10.3897/vz.71.e68897

Authors: Emmanuel Matamba, Leigh R. Richards, Michael I. Cherry, Ramugondo V. Rambau

Abstract: Abstract South African small mammals are under-represented in DNA barcoding efforts, particularly from the eastern forested regions of the country. This study reports DNA barcoding of Rhabdomys taxa from previously unsampled parts of the Eastern Cape and KwaZulu-Natal provinces of South Africa. The complete mitochondrial DNA cytochrome oxidase I (COI) gene was sequenced for 101 Rhabdomys sp. individuals from 16 localities from all three main forest groups (coastal, mistbelt, and scarp forests). Molecular data were supplemented with external morphological measurements, including those deemed potential taxonomically diagnostic characters. Findings indicate the area to be inhabited solely by Rhabdomys dilectus chakae. Haplotypes distributed across the three forest groups were separated by shallow sequence divergences ranging from 0.001–0.015 (Kimura 2-parameter model) and displayed very little population genetic structure (FST= 0.071787). Morphological data revealed some regional metric differences in external morphology, but all the head-and-body to tail (HB: tail) ratios match that of R. d. chakae, and consequently, molecular and morphological data are congruent. These data confirm a range extension of R. d. chakae, supporting the utility of COI barcodes in the identification of small mammalian species.

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DOI: 10.3897/vz.71.e64781

Authors: Javier Lobón-Rovira, Werner Conradie, David Buckley Iglesias, Raffael Ernst, Luis Veríssimo, Ninda Baptista, Pedro Vaz Pinto

Abstract: Abstract The taxonomy of Angolan Hemidactylus has recently been revised. However, the lack of fresh material for some groups and regions, has led to the misidentification of some taxa and an underestimation of actual diversity in others. To shed light on the evolutionary history and systematics of Angolan Hemidactylus, we generated a new phylogenetic hypothesis for the group, and updated the taxonomy following an integrative approach. This resulted in the description of four new species (H. pfindaensis sp. nov., H. faustus sp. nov., H. carivoensis sp. nov. and H. cinganji sp. nov.), the reevaluation of two recently described species (H. vernayi and H. paivae) and the synonymization of a recently described species (H. hannahsabinnae). We estimate divergence times for these lineages, providing a preliminary interpretation of their speciation process. Moreover, we suggest and outline 13 Angolan Main Biogeographic Units (AMBUs) in the area, defining a new biogeographic context for future works on Angolan herpetofauna. We now recognize eleven Hemidactylus species in Angola, and we provide here a new morphological key for Angolan Hemidactylus to help with identifications and species assignments in this group.

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DOI: 10.3897/vz.71.e71268

Authors: Gerald Mayr

Abstract: Abstract The occurrence of lateral openings and pleurocoels (lateral fossae) in the corpus of the thoracic vertebrae of extant and fossil neornithine birds is reviewed, with both features having been identified as osteological correlates of the avian pulmonary system. Openings mainly occur in larger species with a high overall bone pneumatization but do not seem to serve for the passage of lung or air sac diverticula. Pleurocoels, on the other hand, are not directly related to pneumatic features and constitute a plesiomorphic trait that was widespread in Mesozoic non-neornithine birds. It is noted that an inverse correlation exists between the occurrence of pleurocoels and the pneumatization of the humerus, with pleurocoels being mainly found in extant and fossil taxa, in which the humerus is not pneumatized by diverticula of the clavicular air sac. Here it is hypothesized that pleurocoels primarily serve to increase the structural resistance of the vertebral body and were reduced multiple times in neornithine birds. In some taxa, their reduction may be related to the development of the furcula, which assists ventilation of the clavicular and cervical air sacs and may thereby contribute to the pneumatization of both, the humerus and the thoracic vertebrae. If so, Mesozoic non-neornithine birds, which had a rigid furcula with massive shafts as well as non-pneumatic humeri and pronounced pleurocoels, are likely to have differed in functional aspects of their air sac system from extant birds.

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DOI: 10.3897/vz.71.e66756

Authors: Ingmar Werneburg, Serjoscha W. Evers, Gabriel Ferreira

Abstract: Abstract In recent years, paleoneurology became a very popular research field and hundreds of brain-endocasts were described. The interpretation of a dorsal protuberance of the brain-endocast puzzled researchers for a long time, the so-called (cartilaginous) rider. This is mainly because of technical limitations in the past and due to non-accessibility of comparative material. Using turtles as a case-study, we conducted a literature review and studied embryological data in addition to fossil and extant species’ endocasts. We assessed three hypotheses on the origin of the rider as relating to 1) the pineal gland, to 2) the blood vessel system, and to 3) skull roof elements. Based on our integrated anatomical observations, we refute the pineal gland hypothesis (1) and an exclusive blood vessel explanation (2). However, we show that, in most cases, the cartilaginous origin applies (3). The related cartilages, mainly the anterior process of the chondrocranial tectum synoticum, can persist until adulthood. Its diversity is interpreted in regard to the mechanical support for the temporal skull region, the shape of which has been shown to be in turn related to neck retraction and jaw mechanics. Finally, we highlight the value of embryological data to provide profound hypotheses for evolutionary research despite its low quantitative evaluability. We argue that it should be studied in conjunction with modern computer-aided data acquisition whenever possible.

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DOI: 10.3897/vz.71.e66871

Authors: Amanda Kaatz, Jesse L. Grismer, L. Lee Grismer

Abstract: Abstract By studying ecomorophology in the context of phylogeny, researchers can parse out similarity due to common ancestry versus that due to convergence. This is especially true among relatively closely related species where both phylogenetic and environmental constraints may be operating simultaneously. We explored these issues among three karst-associated species from two lineages of Cyrtodactylus—the sworderi group from Peninsular Malaysia and the swamp clade from Peninsular Malaysia and western Indonesia of the agamensis group. A stochastic character mapping analysis using five different habitat preferences corroborated a larger previous analysis in recovering a general habitat preference as an ancestral condition for all habitat preferences and a karst habitat preference in C. guakanthanensis and C. gunungsenyumensis of the sworderi group and C. metropolis of the swamp clade as convergently evolved. Multivariate and univariate analyses of 10 morphometric characters revealed that the ecomorphological similarity between C. guakanthanensis and C. gunungsenyumensis of the sworderi group was also convergent. The ecomorphology of C. metropolis of the swamp clade was intermediate between a karst-adapted ecomorphology and a swamp-generalists ecomorphology. Of the 10 morphometric characters employed in this analysis, only three—head length, head width, and forelimb width—showed any signs of phylogenetic signal. Cyrtodactylus metropolis is hypothesized to be a recently refuged swamp-dwelling species that frequented the Batu Caves environments prior to urbanization of the surrounding swamp habitat to which it is now confined.

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DOI: 10.3897/vz.71.e65952

Authors: Martin Päckert, Jens Hering, Abdelkrim Ait Belkacem, Yue-Hua Sun, Sabine Hille, Davaa Lkhagvasuren, Safiqul Islam, Jochen Martens

Abstract: Abstract The Old World sparrows include some of the best-studied passerine species, such as the cosmopolitan human commensal, the house sparrow (Passer domesticus) as well as poorly studied narrow-range endemics like the Iago sparrow (P. iagoensis) from the Cape Verde Archipelago or specialists from extreme environments like the desert sparrow (P. simplex). It is therefore notable that to date the most complete phylogenetic hypothesis for the Old World sparrows comprised only ten of 43 currently accepted species. With this study we provide an updated phylogeny of Passeridae covering about two third of the family’s species richness. Though still being far from taxon-complete, this new phylogenetic hypothesis provides firm evidence to clarify some open taxonomic questions. All genus-level taxa were reciprocally monophyletic with strong support. Contrary to previous classifications, bush sparrows and rock sparrows were not sister taxa, and therefore their classification in separate genera Gymnoris and Petronia is justified. Plumage color traits like the yellow throat patch of the latter two genera or head color pattern in Passer species do not provide reliable phylogenetic information, except for the large-sized African grey-headed sparrows that resulted as a monophyletic group (P. diffusus, P. griseus, P. gongoensis). Unexpectedly, two small-sized species, P. eminibey and P. luteus that to date are regarded as close relatives were firmly nested in two separate clades of Passer sparrows. Therefore, their separate generic treatment under Sorella eminibey and Auripasser luteus (together with A. euchlorus) does not seem justified.

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DOI: 10.3897/vz.71.e65969

Authors: L. Lee Grismer, Hai Ngoc Ngo, Shuo Qi, Ying-Yong Wang, Minh Duc Le, Thomas Ziegler

Abstract: Abstract Maximum likelihood (ML) and Bayesian inference (BI) analyses using two mitochondrial (16S and cyt b) and two nuclear (CMOS and RAG1) genes and 103 specimens recovered the first phylogenies of all 23 extant species of Goniurosaurus. The analyses strongly supported the recognition of four monophyletic species groups with identical inter-specific relationships within the kuroiwae, lichtenfelderi, and yingdeensis groups but discordant topologies at some nodes within the luii group. Both analyses recovered a polyphyletic G. luii with respect to G. kadoorieorum, and owing to the lack of diagnostic characters in the latter, it is considered a junior synonym of G. luii. A stochastic character mapping analysis of karst versus non-karst habitat preference suggested that karstic landscapes may have played a major role in the evolution and diversification of Goniurosaurus. A karst habitat preference is marginally supported as the most probable ancestral condition for Goniurosaurus as well as for the kuroiwae, luii, and yingdeensis groups. However, a non-karst habitat preference is marginally supported as the most probable ancestral condition for the lichtenfelderi group. Multivariate and univariate ecomorphological analyses of the karst-adapted G. catbaensis, G. huuliensis, and G. luii of the luii group and the granite-stream-adapted G. lichtenfelderii of the lichtenfelderi group demonstrated that their markedly statistically different body shapes may be an adaptive response that contributes to habitat partitioning in areas of northern Vietnam where they are nearly sympatric.

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DOI: 10.3897/vz.71.e64345

Authors: V. Deepak, Surya Narayanan, Pratyush P. Mohapatra, Sushil K. Dutta, Gnanaselvan Melvinselvan, Ashaharraza Khan, Kristin Mahlow, Frank Tillack

Abstract: Coluber fasciolatus Shaw, 1802 (today Argyrogena fasciolata auctt.) is the name used for a widespread terrestrial colubrid snake species inhabiting subtropical and tropical dry deciduous/thorn forests of South Asia from Pakistan to India, with unconfirmed records of distribution in Nepal, Bangladesh and Myanmar and a single doubtful record from the northern tip of Sri Lanka. During the past 200 years, A. fasciolata (common name Banded racer) has been placed in different genera, i.e. Tyria Fitzinger, 1826, Zamenis Wagler, 1830, Coryphodon Duméril, Bibron and Duméril, 1854, and Coluber Linnaeus, 1758 where it primarly remained until the mid 1960s and exceptionally until the year 2011. Three subsequently introduced names, viz. Coluber hebe Daudin, 1803, Coluber curvirostris Cantor, 1839, and Argyrogena rostrata Werner, 1924 were synonymized with C. fasciolatus shortly after its description. Based on a combination of characters including body pattern, external morphology and osteological differences Wilson (1967) reviewed the taxon fasciolatus Shaw and considered it as generically distinguishable, removed it from the then heterogeneous and undefined collective genus Coluber and assigned it to the resurrected genus Argyrogena Werner, 1924. Shaw’s (1802) description of C. fasciolatus was based exclusively on the information of Russell’s “Nooni Paragoodoo” published in 1796 in his “Account of Indian Serpents, collected on the coast of Coromandel; […]”. Our analysis of the original data and the depicted type specimen in Russell (1796) revealed that the name fasciolata was initially established for a species distinct from that currently known as the “Banded racer”, and that Russell’s data have been used simultaneously but unwittingly, for more than 150 years, as original source for two valid species from two different genera. Specimens of Banded racer found in the southeastern part of peninsular India are morphologically and genetically distinct from populations of the rest of the distribution area. These populations from central and southern Tamil Nadu state represent a different species, consequently described as a new species herein. Furthermore, examination of specimens of the Banded racer from different populations across its entire range, including the type specimen of the genus Argyrogena (A. rostrata Werner), reveals a similarity in morphology with the genus Platyceps Blyth, 1860. This was further supported by molecular data which demonstrates that the genus Argyrogena is nested within Platyceps.

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DOI: 10.3897/vz.71.e60800

Authors: Daniel Jablonski, Marco Antônio Ribeiro-Júnior, Shai Meiri, Erez Maza, Oleg V. Kukushkin, Marina Chirikova, Angelika Pirosová, Dušan Jelić, Peter Mikulíček, David Jandzik

Abstract: The Levant represents one of the most important reptile diversity hotspots and centers of endemism in the Western Palearctic. The region harbored numerous taxa in glacial refugia during the Pleistocene climatic oscillations. Due to the hostile arid conditions in the warmer periods they were not always able to spread or come into contact with populations from more distant regions. One large and conspicuous member of the Levantine herpetofauna is the legless anguid lizard Pseudopus apodus. This species is distributed from the Balkans to Central Asia with a portion of its range running along the eastern Mediterranean coast. Mitochondrial and nuclear DNA sequences, microsatellite genotypes, and morphology show that populations in this region differ from the two named subspecies and presumably had a long independent evolutionary history during the Quaternary. Here we describe the Levantine population as a new subspecies and present biogeographic scenarios for its origin and diversification. The new subspecies is genetically highly diverse, and it forms a sister lineage to Pseudopus from the remaining parts of the range according to mtDNA. It is the largest-bodied of the three subspecies, but occupies the smallest range.

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DOI: 10.3897/vz.71.e60225

Authors: Anh The Nguyen, Tang Van Duong, L. Lee Grismer, Nikolay A. Poyarkov

Abstract: An integrative analysis of genetic, morphological, and ecological data recovered a new granite cave-adapted species, Cyrtodactylus raglai sp. nov., from the Song Giang River Valley, Khanh Hoa Province, Vietnam. Cyrtodactylus raglai sp. nov. is nested within one of two major clades within the irregularis species group where it forms a monophyletic group with C. cryptus and its sister species C. kingsadai. It differs from its sister species by an uncorrected pairwise sequence divergence of 16.5% and 16.8% based on the mitochondrial genes NADH dehydrogenase subunit 2 and its flanking tRNAs and the first subunit of cytochrome oxidase c (CO1), respectively. Cyrtodactylus raglai sp. nov. is a narrow-range endemic restricted to a riparian, granite cave microhabitat and its overall morphology bears that of other granite cave-dwelling ecomorphs in the genus. The Song Giang River Valley and its associated waterways are currently threatened by the construction of a hydropower station which will negatively impact the surrounding ecosystem. Urgent field surveys in this region are necessary in order to obtain critical data on its biodiversity and importance to conservation efforts in southern Vietnam.

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DOI: 10.3897/vz.71.e59307

Authors: L. Lee Grismer, Perry L. Wood, Jr., Nikolay A. Poyarkov, Minh D. Le, Fred Kraus, Ishan Agarwal, Paul M. Oliver, Sang N. Nguyen, Truong Q. Nguyen, Suranjan Karunarathna, Luke J. Welton, Bryan L. Stuart, Vinh Q. Luu, Aaron M. Bauer, Kyle A. O’Connell, Evan S. H. Quah, Kin O. Chan, Thomas Ziegler, Hanh Ngo, Roman A. Nazarov, Anchalee Aowphol, Siriwadee Chomdej, Chatmongkon Suwannapoom, Cameron D. Siler, Shahrul Anuar, Ngo V. Tri, Jesse L. Grismer

Abstract: The gekkonid genus Cyrtodactylus is the third most speciose vertebrate genus in the world, containing well over 300 species that collectively range from South Asia to Melanesia across some of the most diverse landscapes and imperiled habitats on the planet. A genus-wide phylogeny of the group has never been presented because researchers working on different groups were using different genetic markers to construct phylogenies that could not be integrated. We present here Maximum likelihood and Bayesian inference mitochondrial and mito-nuclear phylogenies incorporating of 310 species that include dozens of species that had never been included in a genus-wide analysis. Based on the mitochondrial phylogeny, we partition Cyrtodactylus into 31 well-supported monophyletic species groups which, if used as recommended herein, will increase the information content of future integrative taxonomic analyses that continue to add new species to this genus at an ever-increasing annual rate. Data presented here reiterate the outcome of several previous studies indicating that Cyrtodactylus comprises an unprecedented number of narrow-range endemics restricted to single mountain tops, small islands, or karst formations that still remain unprotected. This phylogeny can provide a platform for various comparative ecological studies that can be integrated with conservation management programs across the broad diversity of landscapes and habitats occupied by this genus. Additionally, these data indicate that the true number of Cyrtodactylus remains substantially underrepresented.

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DOI: 10.3897/vz.71.e60921

Authors: Evan S. H. Quah, L. Lee Grismer, Perry L. Wood Jr., Kelvin K. P. Lim, Paul Y. Imbun, M. S. Shahrul Anuar

Abstract: The taxonomic status of the ranid frog Abavorana luctuosa (Peters, 1871) was investigated using a combination of molecular and morphological data. The analyses revealed that A. luctuosa sensu lato is composed of two species in Borneo. One of these species agrees with the description of Rana decorata Mocquard, 1890 which is resurrected in the combination Abavorana decorata comb. nov. (Mocquard, 1890). Abavorana decorata is recovered as the sister lineage to the remainder of Abavorana and differs by a 16.0–17.0 % uncorrected pairwise sequence divergence from its congeners A. nazgul and A. luctuosa, respectively. It is distinguishable morphologically from A. luctuosa and A. nazgul by its ventral pattern (bold, black and white reticulations on its venter along with bold banding on the underside of hind limbs vs. generally immaculate and spotted in the latter two species), and a prominent white streak beneath the eye and/or tympanum extending to the corner of the jaw. Abavorana decorata further differs from A. luctuosa by having a significantly wider head and snout, larger interorbital and tympanum diameters, longer femur in both sexes, and various combinations of other mensural characters. Both species are sympatric in Borneo and this discovery adds to a growing number of widespread Sundaic species shown to be species complexes with distinct forms in Borneo.

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DOI: 10.3897/vz.71.e61728

Authors: George Sangster, Gerald Mayr

Abstract: Recent genomic data sets have resolved many aspects of higher-level phylogenetic relationships of birds. Eleven phylogenomic studies provide congruent support for a clade formed by Procellariiformes, Sphenisciformes, Ciconiiformes, Suliformes and Pelecaniformes. This clade is here named ‘Feraequornithes’ following the rules and requirements of the PhyloCode.

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DOI: 10.3897/vz.71.e59495

Authors: Aurélien Miralles, Teddy Bruy, Angelica Crottini, Andolalao Rakotoarison, Fanomezana M. Ratsoavina, Mark D. Scherz, Robin Schmidt, Jörn Köhler, Frank Glaw, Miguel Vences

Abstract: The Paroedura bastardi clade, a subgroup of the Madagascan gecko genus Paroedura, currently comprises four nominal species: P. bastardi, supposedly widely distributed in southern and western Madagascar, P. ibityensis, a montane endemic, and P. tanjaka and P. neglecta, both restricted to the central west region of the island. Previous work has shown that Paroedura bastardi is a species complex with several strongly divergent mitochondrial lineages. Based on one mitochondrial and two nuclear markers, plus detailed morphological data, we undertake an integrative revision of this species complex. Using a representative sampling for seven nuclear and five mitochondrial genes we furthermore propose a phylogenetic hypothesis of relationships among the species in this clade. Our analyses reveal at least three distinct and independent evolutionary lineages currently referred to P. bastardi. Conclusive evidence for the species status of these lineages comes from multiple cases of syntopic occurrence without genetic admixture or morphological intermediates, suggesting reproductive isolation. We discuss the relevance of this line of evidence and the conditions under which concordant differentiation in unlinked loci under sympatry provides a powerful approach to species delimitation, and taxonomically implement our findings by (1) designating a lectotype for Paroedura bastardi, now restricted to the extreme South-East of Madagascar, (2) resurrecting of the binomen Paroedura guibeae Dixon & Kroll, 1974, which is applied to the species predominantly distributed in the South-West, and (3) describing a third species, Paroedura rennerae sp. nov., which has the northernmost distribution within the species complex.

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