DOI: 10.3897/vz.76.e189837

Authors: Amanda K. Pinion, Daemin Kim, Elizabeth P. Dolan, David S. Portnoy, Gary Voelker, Kevin W. Conway

Abstract:

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DOI: 10.3897/vz.76.e176241

Authors: Aurélien Miralles, Mark D. Scherz, Sam Hyde Roberts, Andolalao Rakotoarison, Frank Glaw, Miguel Vences

Abstract: Abstract Madascincus is a genus of quadrupedal skinks endemic to Madagascar, with 12 recognized species described between the 19th and 21st centuries, occupying diverse habitats from humid forests to arid southern regions and even high-altitude areas. Recent field expeditions uncovered two morphologically distinct forms that did not match any known species, prompting integrative taxonomic analyses that combine multilocus phylogenetics and morphology. Results revealed that each of these forms represents a highly divergent and likely ancient lineage, as evidenced by the substantial branch lengths in both mitochondrial and nuclear phylogenetic trees. These investigations led to the formal description of two new species with restricted distribution ranges: Madascincus irery sp. nov., likely endemic to the island of Nosy Be, and Madascincus minotaurus sp. nov., apparently restricted to the Tsingy de Namoroka karst system.

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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.e171834

Authors: Vagner Cavarzere, Enrico L. Breviglieri, Luís F. Silveira

Abstract: The genus Cercomacra includes six species, among them C. cinerascens, which is widespread throughout the Amazon Basin. Historically, six names have been proposed for this species complex, but only four are currently recognized. Past studies have explored relationships among Cercomacra species using either morphological or molecular data, with taxonomic assessments mainly conducted in the early 20th century. To reevaluate the validity of the currently accepted C. cinerascens taxa, we performed a comprehensive taxonomic revision that combined traditional morphological and vocal analyses. Additionally, we used BirdNET, a pre-trained deep learning algorithm developed for bird sound classification. We examined 682 specimens and 347 recordings and identified two distinct morphotypes distributed north and south of the Amazon Basin, along with four recognizable loudsong types. Using both morphological and vocal features, we identified five distinct lineages, two of which are newly described and named. Cercomacra cinerascens occurs north of the Pastaza and Amazon Rivers in northern Amazonia; C. sclateri is found in eastern Ecuador and Peru, south of the Pastaza River and east of the Andes, east to the Ucayali River; and C. iterata stat. nov. is located east of the Tapajos River. The two newly described species are found south of the Amazon and Marañon Rivers. Cercomacra mura sp. nov. occupies the Ucayali-Madeira interfluve, and Cercomacra raucisona sp. nov. inhabits the Madeira-Tapajos interfluve.

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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.e167908

Authors: Irina V. Kulikova, Philip Lavretsky, Kevin G. McCracken, Yury N. Zhuravlev, Irina L. Miroshnichenko, Andrew B. Correll, Jeffrey L. Peters

Abstract: Eurasian wigeon (Mareca penelope) and American wigeon (Mareca americana) are sister species with diagnosable differences mostly in male plumage. They breed in the Palearctic and Nearctic, respectively, but due to transoceanic migrations come in contact in North America, Western Europe, and north-eastern Asia, where they occasionally hybridize. To estimate genomic divergence and study their population structure we sequenced mitochondrial (mt) DNA control region and obtained 3092 autosomal and 189 Z chromosome loci from double-digest restriction associated DNA sequencing (ddRAD-seq). Consistent with previous work with few nuclear loci, we observed discordant patterns between mtDNA and nuclear DNA divergence. Deeply divergent species-specific mtDNA haplogroups contrasted with low autosomal differentiation and moderate Z-sex chromosome divergence. Meanwhile, Z-linked differentiation (ФST = 0.192) between taxa was five times higher than differentiation of autosomal loci (ФST = 0.0386), with four fixed and eight nearly fixed differences in SNPs discovered in three and six Z-linked outlier loci, respectively. No species-specific SNP variants were found among 83 autosomal outlier loci. This elevated Z-chromosome differentiation is most likely the result of selection that has been important in speciation. The lack of population genetic structure within Eurasian wigeon and American wigeon supports the common notion that migratory waterfowl have high dispersal ability that contributes to strong genetic connectivity between geographic populations.

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DOI: 10.3897/vz.75.e156077

Authors: Amanda K. Pinion, Daemin Kim, Elizabeth P. Dolan, David S. Portnoy, Gary Voelker, Kevin W. Conway

Abstract: Combined analyses of molecular and morphological data support a revised classification of Notropis stramineus comprising five distinct evolutionary lineages, two for which names are available, Notropis stramineus (Cope, 1865) and N. missuriensis (Cope, 1871), herein redescribed, and three for which no names are available, here formally described as new species. Four of these clades comprise a species complex, here termed the N. stramineus species complex, while these four clades, in addition to N. procne, N. topeka, N. chihuahua and the fifth clade form a monophyletic group here termed the N. stramineus species group. Notropis lucifer sp. nov., sister taxon of N. chihuahua, is distributed in the Colorado River and Rio Grande basins of Texas (USA), where it is distributed in the Devils River, and presumably also in Pinto Creek; based on distribution, populations in tributaries of the Rio Grande in Mexico are tentatively also assigned to this species. Notropis lucifer sp. nov. can be distinguished from other members of the N. stramineus species group primarily by coloration in life and pigmentation patterns which are still visible after preservation. Notropis oblitus sp. nov. is found in several Gulf Slope streams in Texas, including the upper reaches of Nueces, San Antonio, and Guadalupe River basins, though likely absent from the Colorado River basin. This species can be distinguished from other members by head shape, eye size and pigmentation patterns. Notropis multicorniculatus sp. nov. is found in the western portions of the Arkansas, Canadian and Red River basins in parts of Texas, Oklahoma, and Kansas, and likely also Arkansas and Colorado, as well as the Pecos River (New Mexico), where it is likely non-native. This species can be distinguished from other members of the species complex by comparatively larger and more abundant tubercles in males and differences in body shape. Comments are provided regarding the genus Miniellus, which has been recently suggested to comprise 21 species, including N. stramineus.

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DOI: 10.3897/vz.75.e158031

Authors: Shuo Qi, Hai Ngoc Ngo, L. Lee Grismer, Hao-Tian Wang, Han-Ming Song, Xiao-Yu Zhu, Zhu-Qing He, Zi-Chen Zhou, Pi-Peng Li, Ji-Chao Wang, Ying-Yong Wang

Abstract: Understanding the taxonomy of the Goniurosaurus lichtenfelderi group has long been obscured by limited sampling, inconsistent morphological characters, and a lack of publicly available molecular and distribution data. In this study, we conducted the most comprehensive morphological and molecular assessment of this group to date, integrating extensive sampling from Hainan Island and adjacent mainland regions. Phylogenetic analyses using mitochondrial and nuclear genes, along with SNPs delivered from ddRAD-seq, revealed that G. hainanensis is polyphyletic, consisting of two deeply divergent clades (i.e., western and eastern) on Hainan Island. The western clade, from the type locality at Mt. Wuzhi, shows small genetic divergence from G. sinensis (= G. kwanghua), supporting their synonymy. In contrast, the eastern clade is genetically more closely related to the continental G. lichtenfelderi than to its western counterpart. Divergence time estimates further indicate that the eastern and western Hainan populations have distinct evolutionary histories. Based on integrated evidence, we synonymize G. kwanghua and G. sinensis with G. hainanensis, and delimit the eastern Hainan clade as G. cf. lichtenfelderi, pending further morphological confirmation. Consequently, the number of valid species in the G. lichtenfelderi group is revised from five to four (including G. lichtenfelderi, G. hainanensis, G. bawanglingensis, and G. zhoui). In addition, a revised taxonomic account and updated distribution information for all recognized species are provided in this study.

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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.e167008

Authors: Christian Thrane, John V. Lyakurwa, H. Christoph Liedtke, Michele Menegon, Alice Petzold, Simon P. Loader, Mark D. Scherz

Abstract: Abstract For the last century, herpetologists have referred to any Nectophrynoides Noble, 1926 toad characterized by a large, robust body, with large, distinct parotoid glands, as Nectophrynoides viviparus (Tornier, 1905). Consequently, Nectophrynoides viviparus is also considered to have the broadest distribution of all its congeners, with populations ranging from the Tanzanian Southern Highlands, close to the Tanzania-Malawi border, through the Udzungwa and Mahenge Mountains in the south to Uluguru, Rubeho, and Nguru Mountains in the central part of the Eastern Arc Mountains. However, there is underappreciated morphological diversity within what is generally referred to as N. viviparus, and various populations are isolated by large distances and geographical barriers. Recent molecular studies have shown that N. viviparus from the Southern Highlands, the type locality, is genetically distinct from all other N. viviparus populations in the Eastern Arc Mountains, suggesting the existence of a species complex warranting taxonomic revision. Here, we present an integrative taxonomic assessment of southern populations by supplementing the genetic results with the analysis of morphometric and morphological data for 257 specimens assigned to N. viviparus, including museomic data for name-bearing types. Based on the results, we describe three new species from the N. viviparus species complex, covering the southern Eastern Arc Mountains populations. Together with a revised morphological key to the genus and a gazetteer of known populations, we provide Extent of Occurrence and Area of Occupancy for N. viviparus sensu stricto and the new species to investigate their conservation status compared to other members of the genus.

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DOI: 10.3897/vz.75.e162071

Authors: Carlos M. Marín, Daniel Bocanumenth, Juan M. Daza

Abstract: Abstract The lizard genus Anolis is the second most diverse genus of terrestrial vertebrates. Within Anolis, the highly speciose clade Dactyloa comprises six species series, including the latifrons series. Despite previous efforts to reconstruct its phylogeny, earlier studies have excluded a substantial proportion of the clade’s species diversity. Here, we integrated both historical and newly generated genetic data to reconstruct the most comprehensive molecular phylogeny of the latifrons series to date including 88% of the current species diversity. We also conducted a thorough morphological examination of museum specimens representing ten species, primarily distributed in Colombia, including vouchers of A. danieli from the Central Cordillera used in previous molecular phylogenies. Our phylogeny also included genetic samples of A. danieli from several localities in the Western Cordillera, samples of Anolis limon and A. mirus (two species previously lacking genetic data), and sequences from the Central American species A. kathydayae and A. brooksi. Our results recovered topological differences for A. limon and A. mirus compared to previous hypotheses and revealed that specimens assigned to A. danieli in earlier studies were misidentified and are not phylogenetically related to this species. Instead, our results showed that the true A. danieli is sister to a green anole clade distributed across the Central Cordillera, Pacific region, and Panama. Based on our phylogenetic and genetic distance analyses, we conclude that A. kathydayae should be considered a junior synonym of A. brooksi. Lastly, we describe the taxon previously confused with A. danieli and comment on the taxonomic implications of our findings.

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DOI: 10.3897/vz.75.e149399

Authors: Nikolay A. Poyarkov, Dana D. Skorinova, Andrey M. Bragin, Veniamin V. Kolchanov, Vladislav A. Gorin, Alexey V. Trofimets, Alexander P. Yuzefovich, Dac Xuan Le, Tan Van Nguyen, Pavel P. Skutschas

Abstract: Abstract Herein we examined the cranial osteology of 15 species of Ichthyophis (I. asplenius, I. beddomei, I. glutinosus, I. kohtaoensis, I. larutensis, I. mindanaoensis, I. multicolor, I. nguyenorum, I. nigroflavus, I. sikkimensis, I. singaporensis, I. supachaii, I. tricolor, I. weberi, and Ichthyophis sp. from northern Vietnam) with a special emphasis on the temporal region. We presented the first detailed description of the cranium and the atlas of an Ichthyophis species based on micro-CT scanning data. We discuss the implications of temporal region composition for the systematics of this group and the evolution of the cranium in Gymnophiona as a whole. We further provided comments on a jaw-closing mechanism and reported on the presence of phylogenetically basal cranial features in ichthyophiids that are also found in stem caecilians. Our detailed morphological description was based on a specimen from a previously unknown population of unstriped Ichthyophis from northern Vietnam. We consequently described this population as a new species based on morphological and molecular (3967 bp from cyt b, 12S rRNA, and 16S rRNA mitochondrial DNA genes) lines of evidence. We provide comparisons of external morphological traits of the new species with its congeners and further compare its cranial osteological features with other Ichthyophis for which skull descriptions exist. The new species differs from the morphologically similar species I. yangi and I. chaloensis by a significant divergence in cyt b and 16S rRNA mitochondrial DNA gene sequences (p = 6.5%–6.9% and p = 4.5%, respectively). The new species is currently known only from evergreen forests of Xuan Lien National Park (Thanh Hoa Province) and Pu Hoat (Nghe An Province) Nature Reserve, northern Vietnam, and was recorded at elevations of 700–800 m asl. We suggest the new species be considered Data Deficient (DD), following the IUCN’s Red List categories.

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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.e153514

Authors: François S. Becker, Graham J. Alexander, Krystal A. Tolley

Abstract: Abstract Barking geckos, Ptenopus Gray, 1866 are burrowing geckos that occur across the xeric regions of southern Africa. They possess unique vocal abilities, with males producing loud advertisement calls to attract females. The taxonomy of the genus has remained stable for six decades, with three recognised species: Ptenopus garrulus (Smith, 1849), P. kochi Haacke, 1964, and P. carpi Brain, 1962. Within P. garrulus, two subspecies have been recognised since 1935: the nominotypical form (P. g. garrulus) and P. g. maculatus Gray, 1866. A recent phylogenetic analysis of the genus found that it contains eight to ten putative species. We used an integrated taxonomic approach to delimit a total of nine species, including evidence from phylogenetics, ecology, calls, and morphology. Ptenopus g. maculatus is elevated to full species, thereby restricting the geographic range of P. garrulus sensu stricto to the greater Kalahari. Additionally, four new species are named which were previously included in ‘P. g. maculatus’: Ptenopus adamanteus sp. nov. from the southern Namib Desert, P. circumsyrticus sp. nov. from the central Namib Desert, P. kenkenses sp. nov. from the northern Nama Karoo, and P. australis sp. nov. from southern Nama Karoo. As a result, the range of P. maculatus sensu stricto is restricted to the central northern Namib Desert. Furthermore, one new species previously included in P. carpi is named P. sceletus sp. nov. from the Skeleton Coast (northern coastal Namib Desert), thereby restricting the range of P. carpi sensu stricto to a small strip of coastal Namib Desert between the Swakop and Kuiseb rivers. The Namib Desert is the centre of diversity for the genus Ptenopus, containing seven of the nine species including the oldest divergent lineages. Two species-level keys are provided: a morphological key and a unique bioacoustic key to the advertisement calls.

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DOI: 10.3897/vz.75.e161536

Authors: Wolfgang Wüster

Abstract: The advent of molecular methods has revolutionised the field of species delimitation and description, one of the key tasks of systematic biology. In animal taxonomy, one marker, the mitochondrial DNA (mtDNA) molecule, has acquired and retained disproportionate influence. This is despite its uniparental, clonal mode of inheritance, as a result of which the entire molecule acts as a single locus, and that precludes its use as a test for admixture between putative lineages, a key consideration in species delimitation. To establish the extent to which the limitations of mtDNA affect present-day taxonomy in non-avian reptiles, I surveyed species descriptions and delimitations published during the years 2023–2024, determined the markers used, and whether analyses of different markers were set up to critically test or just to confirm mtDNA-inspired candidate species. Mitochondrial DNA remains the dominant molecular marker in reptile taxonomy, being used in 84% of species descriptions and delimitations, and as the sole molecular marker in 44%. Despite the immense progress in next generation sequencing (NGS) technologies and their increasing affordability, only 3.4% of descriptions used NGS approaches. In 61% of descriptions, taxa were identified primarily through mtDNA divergence, and additional data (morphology, single-copy nuclear gene sequences) were used as confirmatory evidence rather than as rigorous tests of mitochondrially inferred species limits. I reiterate the importance of truly integrative species delimitation that critically tests species limits first hypothesised from mtDNA, and suggest ways of improving the robustness of species delimitations by optimising the allocation of resources to more appropriate markers and through analytical approaches that critically test the evolutionary independence of putative species.

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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.e153115

Authors: Anna A. Bannikova, Paulina D. Jenkins, Vladimir S. Lebedev, Svetlana V. Pavlova, Vasily D. Yakushov, Alexandra A. Raspopova, Yongke Zhu, Yun Fang, Yue-Hua Sun, Boris I. Sheftel

Abstract: Abstract The taxonomy of the stripe-backed shrew complex (Sorex cylindricauda species group), distributed in mountains of western China, appears challenging due to remarkable variation in morphological traits and relatively recent times of diversification. According to classical points of view only two or three species of the stripe-backed shrews can be distinguished. However, previous molecular reconstructions revealed at least 14 genetic lineages including a number of undescribed cryptic species. In the current study we revise the taxonomic status of large-sized stripe-backed shrews occurring in high mountain areas in south Gansu, north-western Sichuan and western Qinghai that were previously treated as S. aff. cylindricauda or S. sinalis. The available molecular data place them in a separate species-level lineage of the stripe-backed shrew complex. Our morphological analysis indicate that shrews of this lineage are distinct from the two other large-sized Chinese species, S. cylindricauda and S. sinalis, based on both cranial and external traits. Therefore, we here describe it as a species new to science, the karyotype of which is characterized by 2n = 26 with an additional B chromosome and NFa = 44. Our molecular phylogenetic analysis demonstrates multiple instances of mitonuclear discordance among lineages within the S. cylindricauda complex, which is likely a result of mtDNA introgression, thus highlighting the important role of reticulation events in the evolution of the group.

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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.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.e148147

Authors: Andrey Y. Puzachenko, Viktor G. Yudin, Alexei V. Abramov

Abstract: Abstract Four species of Eurasian badgers (Meles) are widely distributed across Eurasia. Small-sized and dark-coloured Far Eastern M. leucurus amurensis is the most distinctive form among all Asian badgers, M. leucurus. Traditionally, Far Eastern badgers are treated as a subspecies of the M. leucurus. This study aims to revise the geographical variability of the M. leucurus sensu lato from the entire species range (324 adult skulls) and assess the position of the Far Eastern badgers. In addition, 551 adult skulls of the European badger, M. meles, the Southwest Asian badger, M. canescens, and the Japanese badger, M. anakuma, were used for comparison with Asian badgers. Meles leucurus amurensis from the Far East resembles the Japanese M. anakuma in many respects, but is very different from the M. l. leucurus of southern Siberia and Middle Asia. Analysis of palaeontological data suggests that the Far Eastern badgers could have descended from the ancestral “leucurus-like” badger chronospecies known from China since the mid-Early Pleistocene. It then probably dispersed westwards into southern Siberia no later than the MIS9 stage (~ 0.32 Ma), where it evolved into M. leucurus. Previous molecular data suggest that the Japanese badgers are unique, whereas the Far Eastern badgers are closer to Asian badgers from the Urals, Siberia and Tibet. We propose that the Far Eastern badger retains some craniometrics features of the ancestral form of M. leucurus sensu stricto and M. anakuma and should therefore be considered a separate species, M. amurensis Schrenck, 1859 stat. rev. The presumed range of this species is in the Far East, east of the Great Khingan Range to Russian Primorye, Manchuria and the Korean Peninsula and possibly eastern China.

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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.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.e134683

Authors: Uliana V. Gorobeyko, Denis V. Kazakov, Anastasia A. Kadetova, Irina N. Sheremetyeva, Valentin Yu. Guskov, Irina V. Kartavtseva, Nikolai E. Dokuchaev, Evgeniy S. Zakharov, Sergei V. Kruskop

Abstract: Abstract Myotis petax is a common and widespread Asian bat species, whose intraspecific sequence variability remains poorly understood. In this work we analyzed the variability of the mitochondrial control region and craniometric measurements for an extensive sample set originating from the entire species range. This made it possible to identify the main genetic lineages and to compare their distribution with the morphological groups. From our investigations, we found that the prevalent genetic lineages, namely, “Siberia,” “Amur,” and “Okhotsk,” appear to be connected to large river systems. The cohabitation of various genetic lineages occurs only in territories where different river basins are connected, such as the Primorsky Territory, Khabarovsk Territory, Transbaikalia Territory, and Mongolia. Moreover, we discovered that the five morphological groups (Siberia, Okhotsk, Amur, Kunashir, and Korea) are partially correlated with previously identified genetic lineages and subspecies. However, M. p. petax and M. p. loukashkini were the only two out of the five subspecies that could be well-defined using specific mtDNA sequences and morphological descriptions. Nonetheless, the subspecies M. p. ussuriensis does not have a distinct genetic lineage to allow for their classification. Notably, a specific mix of morphological group and a genetic lineage characterize the “Amurian morphological form,” which may support its validity as a subspecies rank. That notwithstanding, more information is needed to fully unravel the intraspecific structure of M. petax in the southern Far East and potential contact zones of diverse forms.

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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.e144189

Authors: Uwe Fritz

Abstract: Editorial Expression of Concern

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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.e133467

Authors: Arlo Hinckley, Jesús E. Maldonado, Noriko Tamura, Jennifer A. Leonard, Melissa T. R. Hawkins

Abstract: Abstract We present a comprehensive integrative taxonomic review of Callosciurus caniceps and Tamiops mcclellandii as they are currently defined. This review combines published molecular evidence, craniodental morphometrics, pelage and bacular variation, evaluations of potential hybrid zones using museum specimens and citizen science photographs, and, for C. caniceps, bioacoustic evidence. Our findings lead to the recognition of two species that had been lost in synonymy and highlight future perspectives on species delimitation in Sciuridae. By comparing phenotypic differentiation across climatic and vegetation transitions and contextualizing our results with the evolutionary history of our study systems, we provide insights into distribution, ecogeographical patterns, and speciation drivers in Southeast Asian vertebrates.

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DOI: 10.3897/vz.74.e121110

Authors: Miguel Vences, Jörn Köhler, Carl R. Hutter, Michaela Preick, Alice Petzold, Andolalao Rakotoarison, Fanomezana M. Ratsoavina, Frank Glaw, Mark D. Scherz

Abstract: Abstract The Malagasy stream-breeding treefrog species Boophis marojezensis contains bioacoustically and genetically highly divergent populations. Some of these populations have been defined as candidate species and emit somewhat bizarre advertisement calls consisting of multiple whistle-notes. We here enable a long-overdue taxonomic revision of this species complex by applying a museomics approach to sequence DNA from the holotype of B. marojezensis. Based on an integrative approach that combines divergence levels in mitochondrial DNA and in three nuclear-encoded genes, morphological data, and bioacoustic comparisons, we conclude that eight different species exist in this complex, seven of which are formally described herein as new. Although morphological differences between species are small and mainly separate small-sized from larger-sized species, conclusive evidence for the new species comes from their sympatric and sometimes syntopic occurrence without haplotype sharing in three nuclear genes and under maintenance of bioacoustic differences. Uncorrected genetic divergences in the mitochondrial 16S rRNA gene are >3% in almost all cases, and in some cases up to 8%. In reference to the otherworldly sounds by which these frogs fill Malagasy rainforests, some of them reminiscent of sounds of technical equipment in the fictional “Star Trek” universe, we here name and describe the seven new species in honor of fictional captains of starships, namely B. kirki sp. nov., B. picardi sp. nov., B. siskoi sp. nov., B. janewayae sp. nov., B. archeri sp. nov., B. pikei sp. nov., and B. burnhamae sp. nov. The majority of these species occur in northern Madagascar, where up to three species can occur in immediate geographical proximity, e.g., B. marojezensis, B. burnhamae sp. nov. and B. pikei sp. nov. at different elevations in the Marojejy Massif. South of 16°S latitude, only B. janewayae sp. nov., B. picardi sp. nov., and B. kirki sp. nov. are found, with the latter extending southwards to Ranomafana National Park. Our study confirms the existence of numerous morphologically cryptic and microendemic species among Madagascar’s amphibians, some of which are known only from unprotected sites and require adequate conservation management.

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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.e130038

Authors: George M.T. Mattox, Flávio C. T. Lima, Ralf Britz, Camila S. Souza, Claudio Oliveira

Abstract: Abstract The miniature fish genus Priocharax currently comprises seven valid species: P. ariel, P. britzi, P. marupiara, P. nanus, P. pygmaeus, P. toledopizae and P. varii. Except for P. ariel and P. pygmaeus, all the species are endemic to Brazil. Priocharax is characterized by several paedomorphic features such as reductions in the laterosensory system, number of fin rays, and ossification of parts of the skull. The most striking reductive character of Priocharax is the larval rayless pectoral fin in which most of its ossified endoskeletal elements are absent. We describe herein two new species of Priocharax from the vicinity of Santarém municipality, Pará state, Brazil. Both new species are distinguished from each other and from congeners by a combination of morphological features (i.e., osteological, morphometric, and meristic data) and molecular information (i.e., DNA barcode). We also present an updated maximum likelihood tree which now includes all nine species of Priocharax.

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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.e123251

Authors: Stephen J. Richards, Paul M. Oliver

Abstract: Abstract We describe two new species of torrent-breeding Litoria Tschudi, 1838 from low-elevation hill-forest habitats on the southern fringe of Papua New Guinea’s Central Cordillera. One is currently known only from the Kikori River basin, and the other is known from the Kikori and adjacent Strickland River basins. The two new species can be distinguished from all other Litoria by aspects of morphology and advertisement call structure. Both are known only from below 500 m a.s.l. and so are considered less likely to be threatened by the devastating frog pathogen Batrachochytrium dendrobatidis Longcore, Pessier & Nichols, 1999 than torrent-breeding Melanesian pelodryadid frogs occupying higher, cooler habitats, should that pathogen be introduced to the region. One hundred and ten frog species have now been documented from the Kikori River basin, a near doubling of the total recognised when the first field guide to the region was published nearly 20 years ago, emphasising the rich anuran community of this area.

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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.e108013

Authors: Yun-He Wu, Zhong-Bin Yu, Chen-Qi Lu, Yin-Peng Zhang, Wen-Jie Dong, Xiao-Long Liu, Felista Kasyoka Kilunda, Yun Xiong, Yun-Fang Jiang, Hong Ouyang, Zhong-Xiong Fu, Yun-Biao He, Zhi-Yong Yuan, Jing Che

Abstract: Abstract The torrent frogs of the genus Amolops represent a great anuran diversification in southern China and Southeast Asia. Previous studies have shown that, the diversity of this genus still remains underestimated. During herpetological surveys from 2021 to 2022, several Amolops specimens were collected from the international border regions of southwestern Yunnan Province, China. Herein, we utilized molecular phylogenetic and morphological data to identify these specimens. Our findings indicate the presence of a separate and previously unknown lineage in the A. viridimaculatus group, which we formally describe as a new species. Furthermore, the specimen from Xishuangbanna National Nature Reserve clustered with A. vitreus from the paratype, supporting the morphological diagnosis. Therefore, we describe a new species and a new species record for China. Our study contributes to the species richness of the genus Amolops as well as the diversity of amphibians in China. Notably, our discovery brings the total number of Amolops species to 85 and the total number of torrent frog species known to occur in China to 53. In addition, our study further confirmed that Yunnan and Indochina Peninsula have similar faunal composition, implying that more studies are needed to achieve a complete understanding of the species diversity and distribution pattern.

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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.e113955

Authors: Kyoji Fujiwara, Hiroyuki Motomura, Adam P. Summers, Kevin W. Conway

Abstract: Abstract Rhinolepadichthys, a new genus of the gobiesocid subfamily Diademichthyinae, is described for the “Lepadichthys” lineatus complex (including Rhinolepadichthys geminus comb. nov., R. heemstraorum comb. nov., R. lineatus comb. nov., and R. polyastrous comb. nov.). Detailed investigation of external morphology and osteological anatomy of the new genus and related genera suggests that Rhinolepadichthys represents the sister genus to Discotrema, based on the following putative synapomorphies: (1) presence of a hardened (potentially keratinized) cap on the surface of at least some disc papillae (vs. surface of disc papillae soft, without hardened cap); and (2) the anterolateral part of the ventral postcleithrum extended anteriorly as a well-developed rod-like process, its tip close to the base of pelvic-fin soft ray 4 (vs. only weakly pointed, or irregular). Compared with Discotrema, Rhinolepadichthys gen. nov. is distinguished by the presence of a row of 8–12 large papillae on the inner surface of the upper and lower lips (vs. inner surface of lips smooth, without distinct papillae); the absence (vs. presence) of a well-developed lateral process on the pterotic immediately posterior to the opening of the otic canal; the presence (vs. absence) of gill rakers on the anterior edge of ceratobranchials 1–3; the presence (vs. absence) of gill rakers on the posterior edge of ceratobranchial 4; having the upper pharyngeal teeth arranged in a loose patch on the ventral surface of the pharyngobranchial 3 toothplate, with tooth tips directed posteroventrally (vs. arranged in a single row along posteroventral edge of the pharyngobranchial 3 toothplate, with tooth tips directed posteriorly); features of the adhesive disc, including outline of disc papillae roughly hexagonal or ovoid and with a flattened surface (vs. outline circular, at least some with raised, dome-like surface); the absence (vs. presence) of a deep cavity at the center of disc region C; the absence (vs. presence) of three paired and one median cluster of small papillae (reminiscent of bunches of grapes) across the surface of the adhesive disc; and having the ventral postcleithrum entire, not divided into two separate, articulating elements (vs. ventral postcleithrum divided into an anterior and posterior element, separated via a specialized joint). Reexamination of materials of the poorly known genus Unguitrema, considered a close relative of Discotrema, revealed no morphological differences between the two genera. Unguitrema therefore represents a junior synonym of Discotrema.

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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.e112446

Authors: Vagner Cavarzere, Luís F. Silveira

Abstract: Abstract Cercomacroides serva is widely distributed in northern South America. However, this species has never been thoroughly evaluated taxonomically. We conducted a taxonomic study of three taxa currently classified under Cercomacroides serva, based on a study of 307 skins and 145 recordings of male loudsongs. Females from the northwest Amazonian population differ qualitatively from populations from the southwest Amazon in the coloration of the upperparts, primary, and tail. Male loudsongs are superficially similar between these two populations, but the shape of the notes differs significantly. There is no evidence of intergradation or clinal variation in female plumage or male loudsongs. Based on differences in plumage coloration, loudsongs, and lack of evidence of intergradation, we suggest that two species are best recognized. The name Cercomacroides serva is here restricted to populations north of the Marañon River in Peru, to the north in Ecuador, and in southwestern Colombia. The name Cercomacroides hypomelaena comb. nov. is revived. This species is found south of the Marañon River, and on both banks of the Ucayali River in Peru, to the eastern left bank of the Madeira River in Brazil, and in northern Bolivia.

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DOI: 10.3897/vz.74.e116992

Authors: Sang Ngoc Nguyen, Manh Van Le, Amy Lathrop, Thi-Dieu-Hien Vo, Robert W. Murphy, Jing Che

Abstract: Abstract Homalopsid snakes of the genus Myrrophis include only two species distributed in southern China and northern Vietnam. Here, we describe a third species from southern Vietnam based on morphological data and nucleotide sequences from the mitochondrial gene cyt b. Myrrophis dakkrongensis sp. nov. is diagnosed by the following morphological characters: Medium-sized mud snake (largest total length 452 mm); internasal single and distinctly separated from loreals; dorsal scales smooth, in 23 rows at midbody, reduced to 19 or 20 rows before vent; ventrals 133–138; subcaudals 34–42, paired; cloacal plate divided; supralabials 8, fourth entering orbit; second pair of chin-shields small and oblique; maxillary teeth 17 or 18; gland-like tubercles present in the cloacal region; hemipenis short, forked and spinose, reaching 7th subcaudal; dorsum dark brown to black; and a white or yellow to orange lateroventral stripe present. The new species differs from its congeners by an uncorrected p distance in cyt b sequences of at least 10.5%.

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DOI: 10.3897/vz.74.e114112

Authors: M. Alejandra Sosa, Carolina Acosta Hospitaleche

Abstract: Abstract The vertebral column in tetrapods consists of several constant regions, namely the cervical, thoracic, lumbar, sacral, and caudal regions. Each of these regions is characterized by a specific number of vertebrae, contributing to the overall vertebral formula. Supernumerary and/or missing vertebrae have only been sporadically mentioned for penguins, and the specific vertebral formula is only determined for some non-passeres orders. Variations in the anatomy and vertebral number of South American and Antarctic penguin species are evaluated here. Sixty-six specimens of Aptenodytes forsteri, Pygoscelis adeliae, P. antarcticus, P. papua, Spheniscus magellanicus, and Eudyptes chrysocome were examined to establish the vertebral formula for six South American and Antarctic species, reporting the type and frequency of the variations found in the generalized configuration. We found no intraspecific variation in respect of the number of cervical as well as cervicothoracic vertebrae in all penguin species studied. Intra- and interspecific variation occur in the thoracic, synsacral, and caudal regions comprising 6–7, 13–14 and 5–8 vertebrae, respectively. Particularly, the variations were found in the transitional zones between one region and another and/or between synsacral segments.

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DOI: 10.3897/vz.74.e115242

Authors: Raquel Alvarado-Larios, Pablo Teta, Pablo Cuello, J. Pablo Jayat, Andrea P. Tarquino-Carbonell, Guillermo D’Elía, Paula Cornejo, Agustina A. Ojeda

Abstract: Abstract The genus Ctenomys Blainville, 1826 includes 68 living species of small to medium-sized (100–1200 g) caviomorph rodents of subterranean habits. During the last decade, this genus has been the subject of numerous taxonomic studies, including the description of new species and the proposal of novel synonyms. Based on phylogenetic analysis of mitochondrial DNA sequences and qualitative and quantitative morphological traits, here we review the species boundaries of the tuco-tucos of the species group of C. mendocinus and describe a new species. The new species is morphologically distinct from other phylogenetically and geographically close species of Ctenomys (e.g., C. fochi, C. mendocinus), showing several differences in their craniodental traits (e.g., proportionally longer nasals and less globose tympanic bullae). The new species occurs in montane grasslands and shrublands of northwestern Mendoza (ca. 2710 m a.s.l.) and in lowlands (ca. 1000 m a.s.l.) of the Monte Desert ecoregion in an area highly impacted by accelerated processes associated with the wine industry.

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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.e113084

Authors: Sandeep Das, Saunak Pal, Surya Narayanan, K. Subin, Muhamed Jafer Palot, K. P. Rajkumar, V. Deepak

Abstract: Abstract We describe a new species of Agasthyagama from Idukki district, Kerala, India. The new species is distinguished from its sister species Agasthyagama beddomii by a combination of scale characters and are also genetically different from each other with 11% uncorrected pairwise difference in ND2 gene and 3% in 16S gene. The two species are also geographically separated, the closest distributional records are approximately 80 km apart.

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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.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.e103935

Authors: Ninda L. Baptista, Pedro Vaz Pinto, Chad Keates, Javier Lobón-Rovira, Shelley Edwards, Mark-Oliver Rödel

Abstract: Abstract The pygmy toad genus Poyntonophrynus is endemic to southern Africa. The morphology of these small toads is conserved. They are usually dully colored, and are predominately adapted to arid conditions. During recent surveys in Angola we found Poyntonophrynus specimens that were not assignable to known species. Using an integrative approach, based on mitochondrial and nuclear DNA, morphology, osteology, biogeography and ecology, we identified three new lineages, and describe two of them as new species. All three lineages are closely related to P. pachnodes, an Angolan endemic species, but they are geographically isolated from it. The new species are morphologically distinguishable, and are associated with two of the most important Angolan centers of endemism: the western escarpment and the central highlands. In order to get a more comprehensive understanding of the osteology of the genus, we also provide an osteological characterization of P. dombensis, which was not available to date. Our findings i) increase the number of earless species in the genus Poyntonophrynus, ii) emphasize southwestern Africa as the cradle of diversification in this genus, iii) report the occurrence of Poyntonophrynus in humid environments, thus showing that these toads are ecologically more variable than previously thought, and iv) underline the importance of further biodiversity studies in Angolan centers of endemism.

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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.e102475

Authors: Jennifer A. Sheridan, Somphouthone Phimmachak, Niane Sivongxay, Bryan L. Stuart

Abstract: Abstract The Lao torrent frog Amolops cremnobatus Inger & Kottelat, 1998 was recently hypothesized, based on mitochondrial DNA, to consist of more than a single species across its range in Laos and flanking regions of Vietnam and Thailand. We tested this hypothesis using mitochondrial DNA, nuclear DNA, and quantitative and qualitative morphological data from adults and larvae. We found corroborating lines of evidence for five distinct evolutionary lineages that we hypothesize to be species. Amolops cremnobatus sensu stricto is restricted to the southeastern portion of its previous range, and remaining populations are described as four new species. Some of the new species are easier to diagnose with morphology as larvae than as adults. Further sampling in northern Thailand may reveal an additional species of this torrent frog complex.

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DOI: 10.3897/vz.73.e100889

Authors: Tianyu Qian, Ke Hu, Xiaoyang Mo, Zhiwei Gao, Na Zhang, Daode Yang

Abstract: Abstract We re-examined the megophryinid population from Mt. Hengshan, Hunan Province, China previously identified as Boulenophrys brachykolos (under the name Megophrys brachykolos). Based on newly obtained molecular data, this population appears to be an independent lineage with a relatively distant phylogenetic relationship to B. brachykolos sensu stricto. Furthermore, this population exhibits distinct morphological characteristics that distinguish it from all its congeners. Therefore, we propose to recognize the Hengshan population of B. brachykolos as a new species, Boulenophrys hengshanensis sp. nov. described herein.

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DOI: 10.3897/vz.73.e110512

Authors: Akshay Khandekar, Tejas Thackeray, Ayuthavel Kalaimani, Ishan Agarwal

Abstract: Abstract We describe two new small-bodied, sympatric species of south Asian Cnemaspis belonging to the mysoriensis + adii clade from the Gingee Hills in Tamil Nadu, peninsular India. The two new species can be easily distinguished from the other eight described members of the mysoriensis + adii clade by their dorsal pholidosis, the configuration of femoral and precloacal pores in males, a number of meristic characters and subtle differences in colouration, beside 6.7–20.8 % uncorrected pairwise ND2 sequence divergence. The two species represent different ecomorphs, one a stouter, microhabitat generalist and the other a more slender, elongate rock specialist. The discovery of two new species from granite boulder habitats and Tropical Dry Evergreen Forests is indicative of the importance of these areas for biodiversity. It is likely that similar rocky habitats across southern peninsular India will harbour many more undescribed species.

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DOI: 10.3897/vz.73.e101372

Authors: Zbigniew Szyndlar, Georgios L. Georgalis

Abstract: Abstract We here present a thorough documentation of the vertebral morphology and intracolumnar variation across non-caenophidian snakes. Our studied sample of multiple individuals covers a large number of genera (67) and species (120), pertaining to almost all extant non-caenophidian families. Detailed figuring of multiple vertebrae across the trunk, cloacal, and caudal series for many different individuals / taxa documents the intracolumnar, intraspecific, and interspecific variation. An emphasis is given in the trunk-to-caudal transition and the pattern of the subcentral structures in that region of the column. Extant non-caenophidian snakes show an astonishing diversity of vertebral morphologies. Diagnostic vertebral features for extant families and many genera are given, though admittedly vertebral distinction among genera in certain groups remains a difficult task. A massive compilation of vertebral counts for 270 species, pertaining to 78 different genera (i.e., almost all known valid genera) and encompassing all extant non-caenophidian families, is provided based on our observations as well as an extensive literature overview. More particularly, for many taxa, detailed vertebral counts are explicitly given for the trunk, cloacal, and caudal portions of the column. Extant non-caenophidian snakes witness an extremely wide range of counts of vertebrae, ranging from 115 up to 546. A discussion on the diagnostic taxonomic utility and potential phylogenetic value of certain vertebral structures is provided. Comparisons of the subcentral structures of the cloacal and caudal vertebral series are also made with caenophidian lineages. We anticipate that this illustrative guide will set the stage for more vertebral descriptions in herpetological works but will also be of significant aid for taxonomic identifications in ophidian palaeontology and archaeozoology.

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DOI: 10.3897/vz.73.e109854

Authors: Sabira S. Idiiatullina, Parinya Pawangkhanant, Tanapong Tawan, Thanawut Worranuch, Bunyarit Dechochai, Chatmongkon Suwannapoom, Tan Van Nguyen, Lawan Chanhome, Nikolay A. Poyarkov

Abstract: Abstract We describe a new species of pitvipers from Trang Province of Thailand, near the Thailand–Malaysian border, based on morphological and molecular (2427 bp from cyt b, ND4, and 16S rRNA mitochondrial DNA genes) lines of evidence. Morphologically, Trimeresurus ciliaris sp. nov. is distinguished from its congeners by the following combination of morphological characters: a long papillose hemipenis; first supralabial and nasal scale fused; three to four small supraocular scales; internasals not in contact; small scale between nasal and the scale formed by the fused second supralabial and loreal present; dorsal scales in 17–17–15 rows across the body; ventral scales 172–175 in males, 171 in female; subcaudal scales 59–63 in males, 61 in female, all paired; in life an emerald-green dorsum with reddish-brown bands; creamy-white venter lacking dark dots or stripes on the lateral sides of the ventrals; white vertebral spots present in both sexes on every two or three dorsal scales; dark brown spots forming discontinuous pattern present on 1–3 lateral dorsal scale rows; males with reddish-brown postocular stripe. The new species forms a distinct clade on the phylogenetic tree of the genus Trimeresurus and differs from the morphologically similar species T. venustus by a significant divergence in cytochrome b mitochondrial DNA gene sequences (p = 12.5%). The new species is currently known from a small karstic area in the Nakawan Range spanning the border of Thailand and Malaysia, in particular in limestone forests in Trang and Satun provinces (Thailand); it likely also occurs in the adjacent parts of Perlis State (Malaysia). Our study also suggests that the taxonomy of T. kanburiensis species complex requires further studies; in particular our study suggests that the status of populations from Chumphon Province of Thailand and Pulau Langkawi Island of Malaysia should be re-assessed.

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DOI: 10.3897/vz.73.e108402

Authors: Mariana Chuliver, Agustín Scanferla

Abstract: Abstract Snakes lay their eggs in clutches of different size, which are usually attached to each other forming a cluster. Egg-clustering is a widespread phenomenon across alethinophidian snakes, mostly recorded in Pythonoidea and caenophidian clades. Here we report a new type of egg-clustering for threadsnakes (Leptotyphlopidae) that departs from the alethinophidian type. We found that females of Epictia australis and Leptotyphlops sylvicolus lay their eggs connected to each other through a filament, and we dubbed it ‘string-egg clustering’. The histomorphology of the filament linking the eggs in E. australis showed an outer calcareous layer underlain by a thick layer of collagen fibers, demonstrating that it is an integral part of the eggshell formed during its deposition process in the oviduct. String egg-clustering seems to be present only among species belonging to both subfamilies of threadsnakes, Epictinae and Leptotyphlopinae. Egg-clustering in alethinophidians has been demonstrated to have several advantages for embryo development and post-hatching survival, including fixing the position of the embryo within the egg, protection against predators, and embryo-to-embryo communication. The presence of a filament connecting the eggs in leptotyphlopid species might be relevant for maintaining the position of the embryo in the egg, to avoid the dispersion of the egg in the nesting site, and potentially for the transmission of physical cues. Thus, we hypothesize that the string-egg clustering constitutes an advantageous reproductive trait among threadsnakes.

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DOI: 10.3897/vz.73.e98558

Authors: Shuo Qi, Zhi-Tong Lyu, Han-Ming Song, Shi-Chao Wei, Qi-Feng Zhong, Ying-Yong Wang

Abstract: Abstract In this work, we describe a new species of genus Bufo, Bufo exiguus sp. nov. from Mt. Nankun, Guangdong Province, China. This new species can be distinguished from all congeners by significant divergences in the mitochondrial 16S rRNA and CO1 genes and by a combination of morphological characters: small body size, tympanum absent, parotoid glands small and olive-shaped, tarsal fold absent, dorsal body with a fine vertebral line and white nuptial spinules present on dorsal and inner surfaces of fingers I and II in males. At present, Bufo exiguus sp. nov. is only known from the slow-flowing montane streams from its type locality and its conservation status should be carefully addressed.

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DOI: 10.3897/vz.73.e103534

Authors: Jesus R. D. Souza, Miquéias Ferrão, Igor Luis Kaefer, Antonio Saulo Cunha-Machado, Paulo Roberto Melo-Sampaio, James Hanken, Albertina Pimentel Lima

Abstract: Abstract We use integrative taxonomy to formally describe a candidate species of nurse frog of the genus Allobates from southwestern Brazilian Amazonia. The new species nests within a clade that has been defined historically as A. gasconi, but it has an 8.8–11.0% genetic distance for 16S to samples from the type locality of A. gasconi. The new species differs from congeners mainly by males having a translucent white throat and vocal sac; advertisement calls with a duration of 42–60 ms, two notes separated by an inter-note interval of 8–23 ms, and a dominant frequency of 4,953–6,331 Hz; and exotrophic tadpoles with 2 pyramidal papillae on each end of the upper lip and 10–13 pyramidal and cylindrical papillae surrounding the lower lip. Phylogenetic analyses based on mitochondrial DNA suggest that A. gasconi sensu lato as defined previously represents a complex of as many as seven species, corroborating studies that have shown high levels of cryptic diversity within Allobates.

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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.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.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.e95220

Authors: Petr Papežík, Peter Mikulíček, Michal Benovics, Monika Balogová, Lukáš Choleva, Marie Doležálková-Kaštánková, Petros Lymberakis, Edvárd Mizsei, Simona Papežíková, Nikos Poulakakis, Enerit Saçdanaku, Márton Szabolcs, Radek Šanda, Marcel Uhrin, Jasna Vukić, Daniel Jablonski

Abstract: The genus Pelophylax (water frogs) includes relatively common, widely distributed, and even invasive species, but also endemic taxa with small ranges and limited knowledge concerning their ecology and evolution. Among poorly studied species belong endemics of the southwestern Balkans, namely Pelophylax shqipericus, P. epeiroticus and P. kurtmuelleri. In this study, we focused on the genetic variability of these species aiming to reveal their phylogeographic patterns and Quaternary history. We used 1,088 published and newly obtained sequences of the mitochondrial ND2 gene and a variety of analyses, including molecular phylogenetics and dating, historical demography, and species distribution modeling (SDM). We revelated the existence of two mitochondrial lineages within P. epeiroticus and P. shqipericus that diverged at ~ 0.9 Mya and ~ 0.8 Mya, respectively. Contrarily, no deeply diverged lineages were found in P. kurtmuelleri. Pelophylax kurtmuelleri also shows a close phylogenetic relationship with widely distributed P. ridibundus, suggesting that both represent one evolutionary clade called here P. ridibundus/kurtmuelleri. The estimated split between both lineages in the clade P. ridibundus/kurtmuelleri date back to ~ 0.6 Mya. The divergence between the ridibundus and kurtmuelleri lineages on the ND2 gene is thus lower than the divergence between the two lineages found in P. epeiroticus and P. shqipericus. According to haplotype networks, demographic analyses, and SDM, endemic water frogs survived the last glacial maximum (LGM) in Balkan microrefugia, and their distribution has not changed significantly or even retracted since the LGM. Haplotypes of the kurtmuelleri lineage were also found in northern parts of Europe, where haplotype diversity is however much lower than in the Balkans, suggesting the possible hypothesis of their postglacial expansion to the north.

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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.e102602

Authors: Ishan Agarwal, Tejas Thackeray, Akshay Khandekar

Abstract: We describe two new spotted species of ground-dwelling gecko of the genus Cyrtodactylus (Geckoella) from southeastern India in an integrative taxonomic framework. The new species are recovered as sister taxa within the C. collegalensis species complex, with 13.0–16.7% uncorrected mitochondrial sequence divergence from the other eight members of the C. collegalensis complex and 10.4% from one another. The new species are morphologically diagnosed by a spotted dorsal pattern of four pairs of spots (occasionally fused into figure 8-shaped markings) from the banded species C. aravindi, C. speciosus, C. rishivalleyensis and C. yakhuna; and from the spotted species with three or fewer pairs of spots in C. collegalensis and C. srilekhae; and from C. chengodumalaensis by the absence of any enlarged dorsal scales and from C. varadgirii by the absence of a patch of enlarged roughly hexagonal scales on the canthus rostralis and beneath the angle of the lower jaw. The two new species can only be differentiated from each other based on slight differences in body size, relative body width and other statistically significant, size-corrected morphometric characters. These are among the first endemic lizards from Tropical Dry Evergreen habitats along the southeast coast of India.

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DOI: 10.3897/vz.73.e101065

Authors: Pablo Teta, J. Pablo Jayat, Raquel Alvarado-Larios, Agustina A. Ojeda, Pablo Cuello, Guillermo D’Elía

Abstract: The genus Ctenomys of subterranean rodents is one of the most species-rich genera of Mammalia, with 66 living species currently recognized. However, the taxonomy of the genus is dynamic with several new species and new synonymies proposed during the last decade. One of the species groups that have undergone more changes in contents in the last years is the Ctenomys mendocinus species group. Here, based on phylogenetic analysis of mitochondrial DNA sequences and qualitative and quantitative morphological evidence, we conducted an appraisal of the species richness of tuco-tucos of the C. mendocinus species group, describing two new species from west-central Argentina. The new taxa are morphometrically distinctive when compared with other geographically or phylogenetically close species of the genus, showing qualitative differences in their craniodental anatomy. One of the new species is known from the eastern Andean slopes of La Rioja and San Juan provinces, occurring on montane grasslands and shrublands above 3,500 m a.s.l., while the other is endemic of southwestern Mendoza province, occurring on montane grasslands and shrublands between 2,400–2,700 m a.s.l. In addition, we include for the first time the nominal forms C. fochi and C. validus in a phylogenetic analysis of the genus Ctenomys, showing that both correspond to the C. mendocinus species group, being the second a junior synonym of C. mendocinus. Finally, we made some comments about other candidate species within this species group as well as highlight issues that need to be addressed to gain a robust picture of the specific richness of Ctenomys.

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DOI: 10.3897/vz.73.e101871

Authors: Akshay Khandekar, Tejas Thackeray, Rameshwaran Mariappan, Satpal Gangalmale, Vivek Waghe, Swapnil Pawar, Ishan Agarwal

Abstract: We describe a new species of Hemidactylus based on an integrative taxonomic framework from scrub habitats at the southern tip of India, in Thoothukudi District, Tamil Nadu. The new species has the most densely packed tubercles among Indian Hemidactylus, almost resembling the most tuberculate Indian Cyrtopodion. Hemidactylus quartziticolus sp. nov. is phylogenetically placed within the brookii group of Indian Hemidactylus, where it is sister to the H. gleadowi complex from western-central India. The new species is 14.5–23.7% divergent in ND2 mitochondrial sequence data from other brookii group members, and can be easily diagnosed from regional congeners by its unique dorsal scalation, the number and arrangement of precloacal-femoral pores, the number of dorsal tubercle rows at midbody, number of lamellae under digit I and IV of manus and pes. The new species is currently known only from two isolated, low quartzite hillocks 45 km apart with scrubby, thorn forests and loose, stony soil.

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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.e103319

Authors: Uwe Fritz, L. Lee Grismer, Marika Asztalos

Abstract: Using georeferenced photographic records of 2944 grass snakes from Germany, Austria, and northern Italy as well as previously published mtDNA sequences (n = 1062) and microsatellite data (n = 952) for grass snakes from the same regions, we examined whether or not coloration and pattern reliably allow to differentiate between Natrix natrix and N. helvetica and if so, whether the distribution patterns revealed by phenotypes and genetics are congruent. Furthermore, we used cline analyses across hybrid zones to test whether the phenotypic transition from one species to the other parallels the steep clines unveiled by genetics. Our results suggest that the two species can be reliably differentiated using coloration and pattern. The most powerful diagnostic traits are the presence/absence of side bars on the body flanks, the number of occipital spots, and the shape of the posterior dark occipital spot. The distributions of morphologically identified N. natrix and N. helvetica match their genetically confirmed ranges. Single conflicting individuals morphologically identified as N. natrix or hybrids within the distribution range of N. helvetica either represent misidentifications or translocated snakes. For the genetic markers and phenotypes, our cline analyses revealed concordant steep clines across hybrid zones. However, the southern part of the hybrid zone in Italy, for which no sufficient genetic data are available, should be studied in more detail because the phenotypic data suggest a smooth cline in this region. The unexpected high percentage of putative hybrids with dorsal stripes in this region also calls for further research. For northwestern Germany, another region for which no genetically verified records are available, iNaturalist data suggest that the contact zone of N. natrix and N. helvetica is near the Ems River and extends from there southeastwards to the region of Höxter, North Rhine-Westphalia.

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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.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.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.e97515

Authors: Harald Ahnelt, Oliver Macek, Vanessa Robitzch

Abstract: Species of the gobiid genus Schindleria are among the smallest and fastest reproducing vertebrates of the oceans. We describe a new species, Schindleria qizma, from the Red Sea, Saudi Arabia. It is an extreme example of progenesis, within the already paedomorphic genus, with morphological traits clearly differentiating it from its congeners. Schindleria qizma has a unique, unflexed notochord with a straight urostyle of which the tip is inserted into the hypural cartilage, rather than the typical flexed notochord with an upturned urostyle of the other species of Schindleria. Schindleria qizma belongs to the short dorsal-fin type of Schindleria. It is further characterized by an elongated but relatively deep body; a short dorsal fin originating just slightly anterior to the anal fin (predorsal-fin length 59.4% of SL vs. preanal-fin length 60.2% of SL); a head continuously increasing in depth posteriorly with a straight dorsal profile; a short snout (18.6% of head length); large eyes (34.4% of head length); a short pectoral-radial plate (6.3% of SL); 13 dorsal-fin rays; 11 anal-fin rays; 0–2 procurrent rays (where the last procurrent ray is short, if present); an anal fin with the first anal-fin ray situated opposite the second dorsal-fin ray; toothless oral jaws; females with few (10–11, total) but very large (4.6% of SL) eggs and with a conspicuous urogenital papilla characterized by a wide urogenital opening flanked by two long, bilobed projections; a dorsally pigmented swim-bladder; blackish, iridescent eyes, capped by a silvery layer with irregular rows of black dots or blotches; and no additional external pigmentation on its body, at least in preserved specimens.

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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.e89339

Authors: Napat Ratnarathorn, Bartosz Nadolski, Montri Sumontha, Sjon Hauser, Sunutcha Suntrarachun, Suchitra Khunsap, Panithi Laoungbua, Curtis Andrew Radcliffe, Taksa Vasaruchapong, Tanapong Tawan, Lawan Chanhome

Abstract: The morphological variation, extended distribution, and sequence divergence of a recently described of cobra Naja fuxi Shi et al., 2022 captured from mountainous areas in Thailand are evaluated by using molecular and morphological analyses. We investigated the genetic variation and affinities of 72 specimens in the genus Naja by using mitochondrial DNA (cytochrome b and control region) and the nuclear DNA gene, C-mos. Morphological examination was conducted for 33 cobra specimens obtained from the northern, western, and north-eastern regions, and data on their natural history were gathered during field surveys. A high degree of genetic differentiation was shown to exist between the cobras collected from lowlands and those from mountainous areas. N. fuxi occurs in uplands bordering Thailand’s Central Basin, whereas the similar looking N. kaouthia Lesson, 1831 is more or less restricted to the lowlands. All phylogenetic and network analyses supported a distinct clade of N. fuxi from north, west, and, north-east regions. In addition, N. fuxi seems to exhibit a split between the north-eastern population and those from the north and west. The range of N. fuxi probably extends far into the mountainous areas of the neighbouring countries Myanmar, Laos, and Vietnam. Morphologically, N. fuxi in Thailand can be distinguished from all other cobra species in the adjacent Oriental Region. The speciation of cobras in Thailand likely reflects key events in the region’s geographical, climate and environmental history.

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DOI: 10.3897/vz.73.e99495

Authors: Scott A. Thomson, Natália R. Friol, Arthur White, Dion Wedd, Arthur Georges

Abstract: Disagreement exists on the taxonomic identity of the extant populations of the Australian Elseya referred to in 1992 as the gulf Elseya (= Elseya sp. aff. dentata [Nicholson]). The extant form has since 1997 been considered conspecific with the late Pleistocene fossil Elseya lavarackorum (White and Archer, 1994). Recently it has been considered a new species, Elseya oneiros Joseph-Ouni et al., 2020, conspecific with another fossil found in the same site and stratum as Elseya lavarackorum. Here we re-examine the fossil material and reassess the characters used by previous authors in an attempt to decide the issue. We find that the anterior bridge suture with the carapace of the fossil Elseya lavarackorum is associated with extensive and prominent plastral elements, which has led to misinterpretation of characters associated with this structure. We furthermore show that interindividual variation in sulci patterns is so great as to render them of little taxonomic value. On the basis of (a) deviation of the anterior shape of the carapace from ovoid such that, in aged individuals, the most anterior point of the carapace occurs at marginal scutes M2 (a resultant nuchal bay occurs in such individuals); (b) the typical absence of a cervical scute; (c) no evidence of a medial constriction in the anterior bridge strut suture; and (d) absence of evidence of any other informative variation of taxonomic value; we conclude that the decision to consider the late Pleistocene (ca 23 kyr old) fossil and the extant Elseya sp. aff. dentata [Nicholson] as Elseya lavarackorum (White and Archer, 1994) as conspecific should stand.

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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.e94207

Authors: Rainer Günther, Djoko T. Iskandar, Stephen J. Richards

Abstract: The microhylid genus Oreophryne reaches its greatest diversity in the New Guinea region, where more than 60 species have been documented to date. Most Oreophryne are small (<30 mm SVL) and only three species, O. anthonyi, O. idenburgensis and O. inornata, exceed 40 mm SVL adult body size. Here we describe a fourth large species of Oreophryne that was collected in 1998 from the mountains of Papua Province in western New Guinea. In having a cartilaginous connection between the procoracoid and scapula it is most similar to O. idenburgensis, a species known only from the mountains of central-western New Guinea but differs from that species in a suite of morphological characters including a broader head, a hidden (vs. visible) tympanum and a more pointed snout.

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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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