DOI: 10.3897/vz.75.e169790

Authors: Diogo Parrinha, Francisco M. G. Calado, Mariana P. Marques, Aaron M. Bauer, Luis M. P. Ceríaco

Abstract: Abstract As part of a nineteenth century scientific network, José Vicente Barbosa du Bocage regularly sent “duplicate” specimens from the zoological collections of the National Museum of Lisbon to natural history museums across Europe. These duplicates gained exceptional significance following the 1978 fire that destroyed the Lisbon Museum’s zoological collections, making them the last surviving representatives of its historical holdings. Despite their importance for taxonomic and nomenclatural stability, the full extent of Bocage’s duplicate specimens remains poorly documented. Here we present a comprehensive and integrative revision of the herpetological material sent by Bocage to the British Museum of Natural History. We assess its historical, taxonomic and nomenclatural value, providing an illustrated and annotated catalogue of type specimens. A total of 92 specimens representing 57 species were sent from Lisbon between 1864 and 1896, including 30 type specimens for 27 nominal taxa. We provide evidence for the correction of the type locality associated with the only surviving syntype of Agama anchietae, as well as the recognition of previously unknown types of Chioglossa lusitanica, Hylambates angolensis, Hylambates cynnamomeus, Cystignathus bocagii, Hyperolius insignis, Hyperolius huillensis, Hemidactylus cessacii and Ophirhina anchietae.

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DOI: 10.3897/vz.75.e159878

Authors: Francisco Brusquetti, Diego Bueno-Villafañe, Andrea Caballero-Gini, Paulo D. P. Pinheiro, Flavia Netto, Freddy Burgos-Gallardo, Danilo Fernández Ríos, Edgar Bernabé Cardozo, Diego Baldo

Abstract: Abstract The widely distributed Neotropical frog Leptodactylus fuscus has long been suspected to represent a species complex. Among its numerous synonyms is Leptodactylus gualambensis Gallardo, 1964, described based on specimens from the western South American Gran Chaco and regarded as a typical Chacoan species. This species was synonymized 40 years ago and has not been reassessed since. In this study, we evaluate the taxonomic validity of the Chacoan populations of L. fuscus potentially assignable to L. gualambensis, based on molecular phylogenetic analyses, lineage delimitation, bioacoustics, and morphology using extensive sampling from the South American Gran Chaco and adjacent regions. Our phylogenetic analyses revealed three clades: One distributed across northern South America and Central America, including topotypes of L. fuscus; a second primarily occurring in the South American Gran Chaco, including topotypes of L. gualambensis; and a third found in parts of Paraguay, Argentina, and Brazil. This structure is corroborated by lineage delimitation analysis, genetic differentiation, gene flow estimations, and by differences in advertisement call duration and frequency, as well as head and limb proportions. Based on these results, we support the revalidation of L. gualambensis, redefine the geographic distribution of L. fuscus sensu stricto, and discuss the status of other available synonyms. In addition, we discuss variation in certain morphological traits of L. gualambensis that may be associated with its broad geographic distribution, which encompasses both dry, highly seasonal environments and humid, relatively homogeneous habitats.

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DOI: 10.3897/vz.75.e142775

Authors: Anita Malhotra, Mrinalini Walter, Guillem Limia Russel, Roger S. Thorpe

Abstract: Abstract In this study, we present a comprehensive analysis of species in the Trimeresurus albolabris complex (comprising the white-lipped pitviper, its former subspecies, and closely related species) to identify genetic and phenotypic lines of evidence to assess whether these taxa represent independently evolving lineages or populations with wide-ranging variants. Our morphometric data set includes over 400 live and preserved specimens spanning almost the entire range of T. albolabris sensu stricto and its relatives. We compare patterns of differentiation in three mitochondrial gene fragments for over 300 specimens and conduct a population genetic analysis of nuclear NT3 sequences from over 200 specimens. This level of detail allowed us to identify instances of incongruence between morphological affinities, mitochondrial clades, and nuclear haplotype distribution in putative taxa throughout Indochina. Although recently described species from this region, including T. salazar, T. caudornatus, and T. uetzi, are related to T. septentrionalis in the mitochondrial phylogeny, they extensively share nuclear haplotypes with the purpureomaculatus group, particularly with T. erythrurus. The most common haplotype of T. albolabris sensu stricto is also present throughout Indochina. However, populations in southern Indochina and western Java are morphologically differentiated, belong to distinct mitochondrial clades, and have a high proportion of private NT3 alleles. As they appear to intergrade extensively with populations further north, we herein resurrect the Javan taxon Bothrops viridis var. fario Jan, 1859 as a nomen for this population at the subspecies level, as Trimeresurus albolabris fario comb. nov. Furthermore, we lower the rank of taxa from the extensive zone of intergradation that stretches from Myanmar to central Vietnam to that of subspecies under T. albolabris (T. a. guoi) or T. septentrionalis (T. s. salazar, T. s. caudornatus, and T. s. uetzi), pending detailed further investigations of the extent of gene flow between them and other recognised species in the complex.

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DOI: 10.3897/vz.75.e150370

Authors: Christian Kehlmaier, Uwe Fritz, Gerald Kuchling

Abstract: Abstract Using mitochondrial genomes and nine nuclear loci, we examined genetic variation in snake-necked turtles (Chelodina sensu lato), with a focus on northern Australian taxa. The mitochondrial phylogeny of the genus is confounded by multiple introgression events, rendering the subgenera Chelodina sensu stricto and Chelydera non-monophyletic. However, in the analyses of our nuclear dataset (6071 bp), the recognition of the subgenera is supported. The morphologically most distinct taxa (Chelodina expansa, C. longicollis, C. oblonga, C. parkeri, C. steindachneri) are well differentiated genetically. However, many other species are not or only weakly distinct, calling their validity into question. Our dataset includes sequences from historical museum material and the holotype of C. kuchlingi, a species currently listed as Critically Endangered by the Biodiversity Conservation Act of Western Australia. Resequencing its mitogenome using protocols optimized for formalin-preserved specimens provides evidence that the formerly reported mitochondrial distinction of C. kuchlingi was based on a sequencing artifact. Two historical specimens of C. kuchlingi are genetically indistinguishable from snake-necked turtles living today on the Ord River floodplain. In addition, C. walloyarrina, a geographically close taxon with introgressed mitochondria from another species, is not differentiated on the nuclear genomic level. We conclude that Chelodina walloyarrina (McCord & Joseph-Ouni, 2007) is a junior synonym of Chelodina kuchlingi Cann, 1997 and that the extant snake-necked turtles from the Ord River floodplain are conspecific. This implies that morphological traits used in the past to diagnose the involved taxa are less important than previously thought. The redefined species C. kuchlingi is distributed on the sandstone plateau and associated escarpments as well as on the lowland coastal plains of the Kimberley region of tropical northern Australia. It no longer qualifies as Critically Endangered and has to be downlisted, pending a new status evaluation. Our results underline the importance of a robust taxonomy for conservation decisions. Further research is warranted to examine the validity of the remaining weakly differentiated Chelodina taxa, which could not be resolved in our analyses.

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DOI: 10.3897/vz.75.e126067

Authors: Shing-Lai Ng, Nicolas Straube, Kwang-Ming Liu, Shoou-Jeng Joung

Abstract: Abstract The shark genus Etmopterus is the most species-rich shark genus, however, several of its species level taxa pose taxonomic challenges. Especially the Etmopterus lucifer species group is in need of a taxonomic re-evaluation. In the present study, we review the status of E. lucifer and E. molleri from the north- and southwestern Pacific applying an integrative taxonomic approach. Our dataset comprises 100 morphological characters and the mitochondrial NADH2 marker (1,010 bp) for 178 and 83 specimens, respectively. Our results show that E. lucifer and E. molleri from the Northwestern Pacific are distinct from specimens sampled in the Southwestern Pacific. We therefore (1) resurrect E. abernethyi for specimens in the southwestern Pacific hitherto assigned to E. lucifer, (2) synonymize the Northwestern Pacific E. burgessi with E. lucifer and (3) resurrect E. schmidti for specimens in the Northwestern Pacific hitherto assigned to E. molleri. A lectotype is designated herein for E. lucifer. Redescriptions of the four valid species, E. abernethyi, E. lucifer, E. molleri, and E. schmidti, are given and an updated key to all members of the E. lucifer group from the central Indo-Pacific is provided. The current division of the E. lucifer subgroups is challenged, as the key character, the relative length of flank-marking branches, shows great intraspecific variation.

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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.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.e113347

Authors: Sabira S. Idiiatullina, Tan Van Nguyen, Parinya Pawangkhanant, Chatmongkon Suwannapoom, Lawan Chanhome, Zeeshan A. Mirza, Patrick David, Gernot Vogel, Nikolay A. Poyarkov

Abstract: Abstract Despite recent progress in our understanding of diversity within the genus Trimeresurus Lacépède, 1804, the subgenus Popeia Malhotra & Thorpe, 2004, distributed across most parts of East and Southeast Asia, remains taxonomically challenging. We applied an integrative taxonomic approach including analyses of morphological data and four mitochondrial genes (12S and 16S rRNA, cytochrome b, and NADH dehydrogenase subunit 4), along with examination of available type material, to address longstanding taxonomic questions in one clade within Popeia, the T. popeiorum group, and reveal a high level of hidden diversity of these snakes in the Indo-Burma Biodiversity Hotspot. We confirm that T. popeiorum Smith, 1937 sensu stricto is restricted to Northeast India, eastern Nepal, southern Bhutan, southeastern Bangladesh, western Yunnan Province (China), and northern and southwestern Myanmar. We further confirm that the recently described species T. yingjiangensis Chen et al., 2019 is a junior synonym of T. popeiorum. In addition, we discovered that the combination Trimesurus [sic] elegans Gray, 1853 is a valid senior synonym of T. popeiorum and threatens the stability of the latter taxon. Therefore, in order to protect the nomen popeiorum and in accordance with Article 23.9 of the International Code of Zoological Nomenclature, we regard the taxon Trimesurus elegans as a nomen oblitum and render Trimeresurus popeiorum a nomen protectum. Examination of a larger series of specimens allows us to describe two new cryptic species of Trimeresurus from the Indo-Burma Region. This study brings the total number of species in the subgenus Popeia to six and also suggests that the subspecific taxonomy of the T. sabahi complex requires further investigation. We urge adequate actions regarding the conservation of the newly discovered species and recommend further studies on their toxicology.

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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.e112782

Authors: Alex Slavenko, Stephen J. Richards, Stephen C. Donnellan, Allen Allison, Paul M. Oliver

Abstract: Abstract Skinks are the most diverse component of the reptile fauna in the mountains of New Guinea and many seemingly specialised high-elevation species remain undescribed. Here we describe two spectacular new gold-patterned skinks in the montane-specialist genus Papuascincus. Both species can be diagnosed from all congeners by their distinctive colouration, in addition to aspects of scalation and body size. One new species is mainly recorded from lower montane forest in karst habitats spanning more than five hundred kilometres along the southern edge of New Guinea’s Central Cordillera and is likely to warrant an IUCN conservation status of Least Concern. The second new species has thus far only been recorded from cloud forest on the summit of Mt. Menawa in the North Coastal Ranges and we suggest it should be considered Data Deficient. However, if further survey work confirms a restricted distribution with little scope for upslope elevational retreat under future warming climates it will likely qualify for Endangered or Critically Endangered status.

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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.e110875

Authors: Heok Hee Ng, Maurice Kottelat

Abstract: Abstract Mystus celator sp. nov. is described from the Irrawaddy River drainage in northern Myanmar. It can be distinguished from congeners in having a combination of: three equally dark longitudinal stripes separated by two pale interspaces on sides of body; round, dark tympanic spot; ovoid, dark spot on caudal peduncle; length of adipose-fin base 18.0–23.3% SL; angle of predorsal profile 21–24°; posterior cranial fontanelle not reaching base of supraoccipital process; 25–30 rakers on the first branchial arch; and 35–36 vertebrae. The identity of Mystus pulcher is fixed with the designation of a lectotype.

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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.e102098

Authors: Sarah Mângia, Diego José Santana, Leandro de Oliveira Drummond, Leandro Talione Sabagh, Luiz Ugioni, Paulo Nogueira Costa, Milena Wachlevski

Abstract: The number of described species of Brachycephalus has rapidly increased in the last decade (n = 22, which represents 56% of the total). Species of the genus Brachycephalus are mostly distributed in isolated mountaintops from Bahia (northeastern Brazil) to Santa Catarina states (southern Brazil), each one occupying only one or a few adjacent mountaintops. Herein, we described a new species of Brachycephalus of the B. pernix group, from Serra do Tabuleiro in Santa Catarina state, which also represents the southernmost known species. The new species can be distinguished from its congeners by a combination of characters, including the following: (1) “bufoniform” body; (2) small adult SVL: 9.57–11.10 mm for males and 10.88–12.70 mm for females; (3) head proportionally small (HL/SVL 19–28%) and eye proportionally large (ED/HL 36–56%); (4) dorsum texture rough; (5) snout shape rounded in dorsal and lateral views; (6) general dorsal body color olive green with head, arms and legs yellow-orangish scattered with olive green, and an orangish vertebral stripe spotted with white and brown colors; (7) skull and skeleton without hyperossification; (8) frontoparietal and sphenethmoid not fused; (9) advertisement with one or two high-frequency notes (6,115–6,562 Hz), and 2–4 pulses per note. The type locality is adjacent to Parque Estadual da Serra do Tabuleiro, a protected area, but we observed various agricultural activities in this locality, including the presence of exotic plants, which can change the amount and the quality of leaf litter, somehow compromising the population of the new species. Another aggravating factor is that the municipality of São Bonifácio has conflicts over land use with irregular occupation and unfinished expropriation processes in Parque Estadual da Serra do Tabuleiro. Considering that Brachycephalus sp. nov. is probably a mountaintop microendemic species, it is paramount that future studies quantifying the new species’ full distribution and evaluating population trends to accurately assess its conservation status.

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DOI: 10.3897/vz.73.e101487

Authors: Vuong Tan Tu, Neil M. Furey, Tamás Görföl, Alexandre Hassanin, Satoru Arai, Daisuke Koyabu, Bounsavane Douangboubpha, Gábor Csorba

Abstract: This study integrates analyses of mitochondrial DNA sequences and morphological and acoustic data to re-evaluate the taxonomic status of Rhinolophus rex rex, R. r. paradoxolophus and R. schnitzleri throughout their distribution ranges. Based on a dense geographic sampling of specimens hitherto referred to these taxa and contrary to the current taxonomic view, our results indicate that all examined specimens of these taxa are representatives of a single, widely distributed and morphologically variable species, R. rex. The recognition of its geographic populations as different subspecies (R. r. rex and R. r. paradoxolophus) or distinct species (R. schnitzleri) based on morphological and acoustic data should be regarded as invalid. In the light of this revision, we also reassess the conservation status of R. rex against IUCN Red List criteria as Near Threatened.

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DOI: 10.3897/vz.73.e98170

Authors: Angele Martins, Manuella Folly, Guilherme Nunes Ferreira, Antônio Samuel Garcia da Silva, Claudia Koch, Antoine Fouquet, Alessandra Machado, Ricardo Tadeu Lopes, Roberta Pinto, Miguel Trefaut Rodrigues, Paulo Passos

Abstract: Threadsnakes are known for their conserved external morphology and historically controversial systematics, challenging taxonomic, biogeographic and evolutionary researches in these fields. Recent morphological studies—mostly based on µCT data of the skull and lower jaw—have resolved systematic issues within the group, for instance leading to the description of new taxa or re-positioning little-known scolecophidian taxa in the tree of life. Herein we describe a new polymorphic species of the genus Siagonodon from Amazonia based on morphological (external, osteology and hemipenis) and molecular data, and provide the first hemipenial description for the genus. We also reassign Siagonodon acutirostris to the genus Trilepida based on osteological data in combination with molecular evidence. The new species described represents an evolutionary paradox for scolecophidians because the species displays a remarkable variation in the shape of the snout region that is otherwise always highly conserved in this clade. Finally, this study reinforces the importance of protected areas as essential in maintaining vertebrate populations, including those that are not yet formally described.

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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.e101011

Authors: Neelesh Dahanukar, Remya L. Sundar, Duwaki Rangad, Graham Proudlove, Rajeev Raghavan

Abstract: The world’s largest subterranean fish was discovered in 2019, and was tentatively identified as a troglomorphic form of the golden mahseer, Tor putitora. Detailed analyses of its morphometric and meristic data, and results from molecular analyses now reveal that it is a new species of the genus Neolissochilus, the sister taxon of Tor. We formally describe the new species as Neolissochilus pnar, honouring the tribal communities of East Jaintia hills in Meghalaya, Northeast India, from where it was discovered. Neolissochilus pnar possesses a number of characters unique among species of Neolissochilus, with the exception of the similarly subterranean N. subterraneus from Thailand. The unique characters that diagnose N. pnar from all epigean congeners comprise highly reduced eye size to complete absence of externally visible eyes, complete lack of pigmentation, long maxillary barbels, long pectoral-fin rays, and scalation pattern. Neolissochilus pnar is distinguished from the hypogean N. subterraneus, the type locality of which is a limestone cave ~2000 kms away in Central Thailand, by a lesser pre-pelvic length (47.8–49.4 vs. 50.5–55.3 %SL), a shorter caudal peduncle (16.1–16.8 vs. 17.8–23.7 %SL), and shorter dorsal fin (17.4–20.8 vs. 21.5–26.3 %SL). In addition, Neolissochilus pnar is also genetically and morphologically distinct from its close congeners with a raw genetic divergence of 1.1–2.7% in the COI gene with putative topotype of N. hexastichus and 2.1–2.6% with putative topotype of N. hexagonolepis.

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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.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.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.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.e80508

Authors: John R. Wible

Abstract: Abstract The os paradoxum or dumb-bell-shaped bone is a paired bone occurring in the middle of the specialized bill of the platypus Ornithorhynchus anatinus. It has been variously considered as a neomorph of the platypus, as the homologue of the paired vomer of sauropsids, or as a part of the paired premaxillae. A review of the near 200-year history of this element strongly supports the os paradoxum as a remnant of the medial palatine processes of the premaxillae given its ontogenetic continuity with the premaxillae and association with the vomeronasal organ and cartilage, incisive foramen, and cartilaginous nasal septum. In conjunction with this hypothesis, homologies of the unpaired vomer of extant mammals and the paired vomer of extant sauropsids are also supported. These views are reinforced with observations from CT scans of O. anatinus, the Miocene ornithorhynchid Obdurodon dicksoni, and the extant didelphid marsupial Didelphis marsupialis. At the choanae, Obdurodon has what appears to be a separate parasphenoid bone unknown in extant monotremes.

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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.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.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.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.e67501

Authors: George Sangster, Guy M. Kirwan, Jérôme Fuchs, Edward C. Dickinson, Andy Elliott, Steven M. S. Gregory

Abstract: Abstract Multiple molecular phylogenetic studies have demonstrated that two Neotropical raptors, tiny hawk Accipiter superciliosus and its sister species semicollared hawk A. collaris, are not closely related to core Accipiter, and that A. superciliosus, at least, possesses osteological characters not replicated in the remainder of the genus. Based on these data, there is a need to recognise their distinctiveness at generic level. However, as recently noted in two global bird checklists, no name is available to accommodate them, so we provide a new nomen here. Furthermore, two Asian accipitrids, crested goshawk A. trivirgatus and its presumed closest relative Sulawesi goshawk A. griseiceps, are also phylogenetically distinctive; in this case the genus-group name Lophospiza is applicable. We also designate type species for two genus-group names (Hieraspiza and Eusparvius) currently in the synonymy of Accipiter, and, as an aid to future workers, we provide a synonymy of the genus Accipiter and a list of species currently included in Accipiter for which published molecular phylogenetic data are apparently lacking.

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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.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.e65097

Authors: Omar M. Entiauspe-Neto, Claudia Koch, Michael B. Harvey, Guarino R. Colli, Thaís B. Guedes

Abstract: Apostolepis is a diverse genus of dipsadid snakes, currently comprising 34 species occurring in most cis-Andean South America. The taxonomy of the group is highly unstable. Upon discovering its type series, we redescribe the rare species A. ambiniger (Peters, 1869) and provide an account of its geographic distribution and morphological variation in pholidosis, osteology, and hemipenial characters. We also discuss some aspects of the taxonomy of Apostolepis.

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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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