DOI: 10.3897/vz.76.e195284

Authors: Freddy A. Grisales-Martínez, Juan M. Daza

Abstract: Abstract Phenotypic traits and genetic variation are fundamental for understanding biological diversity and evolutionary processes. In natural populations, phenotypic novelties are generally expected to align with genetic differentiation, establishing phenotype–genotype relationships. This pattern is evident in Neotropical lizards of the genus Anolis, where traits such as the dewlap and hemipenes play key roles in species recognition and sexual selection, and are often associated with reproductive isolation, genetic structure, and lineage divergence. However, in Anolis mariarum, an Andean species from northern Colombia, previous limited sampling suggested a mismatch between these traits and the mitochondrial DNA (mtDNA) lineages. Here, we analyzed specimens from 41 localities across the species’ range, examining hemipenial morphology, dewlap coloration, phylogenetic signal, and genetic structure using three mitochondrial markers. We identified seven mitochondrial clades within A. mariarum, with divergence partially shaped by Andean topography and genetic distances comparable to those reported among Anolis species and subspecies. Two hemipenial morphologies (types A and B) and three dewlap colour patterns (orange–yellow, orange–pink, reddish) were detected, reflecting notable intraspecific variation. Although both traits showed strong phylogenetic signal, they exhibited weak spatial and genetic structuring, suggesting an early stage of divergence. Because our inferences are based exclusively on mtDNA, these results should be interpreted cautiously and provide only a preliminary framework for investigating the evolutionary processes underlying phenotypic diversification. These results highlight the evolutionary relevance of dewlap and hemipenial traits and emphasize the need for extensive morphological, geographic, and genetic sampling to accurately assess lineage boundaries and uncover patterns and processes leading to enormous diversity in Neotropical fauna.

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DOI: 10.3897/vz.76.e205950

Authors: Anna B. Vassilieva, Trung Duc Nguyen

Abstract: Abstract Rhacophorid frogs, with their unique diversity of reproductive modes, are a perfect model group for studying the evolution of different forms of ontogeny. While phylogenetic studies yield controversial results, the morphological basis of the ontogenetic remodeling remains insufficiently explored. In the present research, we compare embryonic and early-larval development in eight species of Rhacophoridae with four different reproductive modes: jelly nests (Feihyla palpebralis, Theloderma vietnamense), bubble nests (Feihyla hansenae), foam nests (Chirixalus nongkhorensis, Polypedates megacephalus, Rhacophorus annamensis, R. vampyrus), and terrestrial direct-developing egg clutches (Kurixalus gryllus). The main focus is on the degree of embryonization in each species, as well as on the development of three transient early-larval features: external gills, opercular fold, and adhesive organs. The initial signs of gill reduction were observed in biphasic jelly-nesting species with a slightly prolonged embryonic period. In direct-developing species, the reduction of larval structures progressed significantly. Development varied among foam-nesting species: while early ontogeny in Chirixalus showed no signs of significant remodeling, both species of Rhacophorus exhibited a strikingly modified developmental pattern, with a severe reduction in external gills and accelerated formation of tadpole morphology within the foam nest, whereas the studied species of Polypedates displayed transitional developmental features. The morphological changes discovered showed that, in the evolution of ontogeny in Rhacophoridae, direct development and advanced foam-nesting represent two fundamentally divergent and highly derived reproductive modes.

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DOI: 10.3897/vz.76.e163421

Authors: Maria Clara El Bayeh de Andrade, Karla D. A. Soares

Abstract: Abstract Chondrichthyes, a monophyletic group comprising sharks, batoids, and chimaeras, includes approximately 1200 valid species and displays remarkable morphological diversity, both internal and external. Among soft tissues, the heart and its associated structures in cartilaginous fishes remain understudied, with most research focusing on circulatory physiology rather than comparative anatomy. This study provides a detailed description of the morphological variation of the afferent hyoid and branchial arteries across 13 of the 14 recognized orders of Chondrichthyes. A total of 16 anatomical characters associated with this complex were analyzed, seven were previously described, with two being redefined, and nine others are proposed as new. All characters were tested cladistically in order to explore evolutionary patterns. Our findings were able to expand the current knowledge of the morphological diversity of afferent branchial arteries in Chondrichthyes and provide new insights into the evolution of cardiovascular structures within this lineage.

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DOI: 10.3897/vz.76.e172567

Authors: Mark D. Scherz, Frank Glaw, Alice Petzold, Franco Andreone, Jörn Köhler, Achille P. Raselimanana, Carl Hutter, Francesco Belluardo, Andolalao Rakotoarison, Michael Hofreiter, Miguel Vences, Angelica Crottini

Abstract: Abstract The diamond frogs of the genus Rhombophryne are a poorly understood clade of morphologically diverse frogs native to the humid forests of Madagascar. We here present a monographic revision of the taxonomy and natural history of these frogs. We first provide morphological diagnoses and contents of the established species groups in Rhombophryne. We then integrate molecular phylogenetic data from mitochondrial and nuclear genetic markers, with morphometrics, bioacoustics, and micro-CT-based osteology, to describe seven new species from across Madagascar. This brings the number of species in the genus to 27. We have obtained DNA sequences from name-bearing specimens of 23 of these species, plus topotypical specimens of two further species; we used museomics methods to sequence name-bearing types of three species collected before 1976, as well as several other specimens of interest, allowing us to clarify taxonomic assignment of those individuals. In our treatments of all existing and new species, we provide new insights into their anatomy, diet, ecology, and distribution. We provide the first published key to the genus, which we hope will make their identification more straightforward. Finally, we suggest a revised conservation assessment in light of the data that we present. We estimate 24 of the 27 species to be threatened, with two species (R. kilonjy sp. nov. and R. matavy) probably worthy of being classified as ‘Critically Endangered’. Several candidate species remain taxonomically unaddressed, due to the poor representation of specimens in museum collections. We therefore emphasise the need for increased survey efforts to fill knowledge gaps in both the taxonomy and the natural history of these frogs.

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DOI: 10.3897/vz.76.e187462

Authors: Pablo Teta, Agustina A. Ojeda, Andrea P. Tarquino Carbonell, J. Raquel Alvarado-Larios, Pablo Cuello, Paula Cornejo, Julián Mignino, Ricardo A. Ojeda, Diego H. Verzi

Abstract: Abstract Octodontidae is a moderately speciose family (seven genera and 16 species) of South American caviomorph rodents; however, from an ecological point of view, it is a diverse clade that includes terrestrial, cursorial, scansorial, and fossorial to strictly subterranean forms. Its distribution is mostly restricted to the western portion of southern South America, on both sides of the Andes. Here we describe a new living genus and species of Octodontidae, whose distribution is restricted to the Sierra de Guasapampa (central Argentina). Phylogenetic analysis of mitochondrial molecular markers (cytochrome b) recovers this taxon as sister to Octomys; the genetic distance observed between them is similar to the range of distances found between other genera of Octodontidae (>9%). This, together with its morphological divergence from Octomys in the context of Octodontidae, supports its treatment as a separate genus. This rodent is characterized by a unique combination of morphological features, including a long and very hairy tail; grayish brown dorsal coloration, which contrasts against the whitish belly; robust skull, with a proportionally short rostrum; wide interorbital region, with slightly divergent supraorbital edges; proportionally wide incisive foramina; orbitosphenoid with its origin displaced to the basisphenoid; inflated but small auditory bullae; median lacerate foramen not covered by the bulla; and upper molariform lobes with acuminate labial tips. Specimens of the new genus and species were captured on rocky slopes with abundant bromeliad vegetation in the understory. This finding highlights the importance of continuing field research, especially in areas that have not previously been surveyed.

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DOI: 10.3897/vz.76.e173627

Authors: Diego A. Caraballo, Denise H. Campo, Pablo Teta, Cecilia Lanzone

Abstract: Abstract Species delimitation in recently evolving taxa presents unique challenges. Such taxa may not have accumulated enough genetic or morphological differences to be easily distinguished using traditional methods, while processes such as incomplete lineage sorting or gene flow may obscure these distinctions. One paradigmatic recently evolved taxon is the Corrientes group of the subterranean genus Ctenomys (Rodentia, Ctenomyidae), a set of populations with different chromosomal forms occurring in the homonymous province of Argentina. Previously, three nominal species and several independently evolving lineages had been proposed; however, a comprehensive integrative approach—including qualitative and quantitative morphological analyses, as well as additional molecular proxies for lineage delimitation—was still lacking, and these results had not been evaluated in a taxonomic context. In this study, we added new lines of evidence, including Bayesian phylogenetics and divergence time estimation, haplotype networks, and genetic distances. We integrated this information with karyotypic and microsatellite data to discuss previously proposed lineages, as well as to analyze the evolutionary processes that shaped the history of this group. Finally, we analyzed the qualitative and quantitative morphology of individuals from the nominal species and a chromosomally derived lineage, and we propose a revised species-level and infraspecific classification for this complex group of rodents. This resulted in the delimitation of two species, C. roigi and C. perrensi, the latter being subdivided into three subspecies: Ctenomys perrensi perrensi, C. perrensi dorbignyi, and C. perrensi iberaensis subsp. nov. In addition, other identified lineages, such as Sarandicito, Manantiales, and Santa Rosa, deserve future taxonomic investigation.

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DOI: 10.3897/vz.76.e180973

Authors: Oleg V. Brandler, Andrey R. Tukhbatullin, Svetlana Y. Kapustina, Sergey N. Matveevsky, Fatimat A. Tembotova, Andrey Y. Puzachenko

Abstract: Abstract The systematics of blind mole rats (Spalacinae), a group of highly specialized subterranean rodents, remain a subject of debate. Within the genus Spalax, the greater blind mole rat (Spalax microphthalmus) is distinguished by its unique diploid chromosome number (2n = 60 versus 2n = 62 in others) and the most extensive geographic range. However, its intraspecific variation has been insufficiently studied. Previous finding of specimens with 2n = 62 in the North Caucasus were attributed to chromosomal polymorphism within S. microphthalmus. In this study, we conducted a comprehensive analysis of morphometric, morphological, molecular, and chromosomal variation across the entire range of S. microphthalmus, integrated with a comparative analysis of the genus Spalax. Our results demonstrate that the North Caucasian populations (2n = 62) exhibit consistent species-level differences in molecular genetics (from 8% to 12% divergence in cyt b), karyotype, and cranial morphology, distinguishing them from S. microphthalmus and all other congeners. This lineage also possesses a unique combination of morphological traits, including features that bring it closer to the hypothetical common ancestor of all modern Spalax species. Based on this integrative evidence, we describe this lineage as a new species: Spalax lyapunovae sp. nov., increasing the number of extant Spalax species to nine. This newly recognized species, endemic to the central North Caucasus, requires further ecological and distributional studies. Given its presumably limited range, an urgent assessment of its conservation status is warranted.

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DOI: 10.3897/vz.76.e182515

Authors: Kun-Hao Chen, Xiao-Yun Wang, Ze-Feng Huang, Yun-Zheng Mo, Yi Wu, Yi-Bo Hu, Yang Yue, Wen-Hua Yu

Abstract: Abstract The taxonomy of genus Myotis (Chiroptera: Vespertilionidae) has long posed considerable challenges, with numerous species in China remaining poorly defined. To address long-standing taxonomic ambiguities in Chinese Myotis, this study integrates over 15 years of fieldwork and conducts a comprehensive assessment of 197 specimens collected primarily in eastern China, which represent approximately 70% of the country’s known species. Molecular species delimitation, phylogenetic reconstruction, and multivariate analyses of morphological data were jointly employed to reassess species diagnostic traits. Phylogenetic and molecular delimitation supported the validity of 30 Myotis species in China, and resolved several long-debated complexes, including M. davidii, M. siligorensis, and M. frater. Principal component and hierarchical clustering analyses revealed mixed and overlapping patterns among species, particularly within small to medium size taxa. These results highlighted the limitations of traditional morphometric traits for distinguishing closely related Myotis. Initial classification accuracy using morphological traits alone was modest. However, when categorical phenotypic data was added into the dataset, model performance improved markedly: Random forest accuracy increased from 77.9% to 90.5%, and the decision tree model successfully discriminated 16 taxonomic units. These suggested that categorical phenotypic data can substantially enhance identification within morphologically conservative groups. Based on integrative evidence, we established an updated identification key. In addition, high-resolution 3D digital models of craniodental structures were generated to facilitate open access for future research. This study provided a foundation for subsequent phylogeny, ecology, and conservation biology studies on this taxonomically difficult genus.

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DOI: 10.3897/vz.76.e180962

Authors: Nikoleta Dubjelová, Tereza Hadravová, Martin Ivanov, Ivan Horáček

Abstract: The European Pleistocene populations of the narrow-headed vole (Stenocranius gregalis), an index species of the Palearctic glacial communities, were recently found to differ from the extant Asian species by a deep genetic divergence and are to be considered a separate species, Stenocranius anglicus, which had to persist through the interglacial stages in local European refugia. Here, we analyze over 2000 first lower molars from 14 stratified localities in the Czech Republic and Slovakia, spanning the Middle Pleistocene to Holocene, employing geometric morphometrics, biometric measurements, and morphotype classifications to assess molar shape variation. Our results demonstrate persistent morphological variability, with particularly high morphotype diversity during MIS 5–3, followed by simplification and reduced variance in post–LGM populations. Morphological divergence was greater among geographic localities than stratigraphic stages, suggesting strong regional and ecological influences. Stratified sequences reveal diverse evolutionary trajectories from long-term morphological stability in refugia to gradual simplification preceding extinction in the early Holocene. These patterns align with broader Eurasian trends but also highlight regionally specific responses to climatic and ecological change accompanying the species’ extinction dynamics during the early to middle Holocene. The paper underscores the importance of integrating detailed morphometrics with stratigraphic and ecological evidence to shed light on these topics.

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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.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.e173317

Authors: Sebastián Lyons, Sergio D. Rosset, Mariana Picasso, Carolina Acosta Hospitaleche

Abstract: Woodpeckers possess specialised cranial and cervical skeletal adaptations that enable them to excavate wood, yet how these structures form and integrate during development is still largely unknown. While several cranial skeletal traits have been described in this context, their postnatal development has received little attention. This study examines the postnatal skeletal development of four species, from nestlings to juveniles, using cleared and stained specimens to assess ossification and bone fusion. Woodpeckers show delayed cranial and cervical skeletal ossification typical of altricial birds, with many elements remaining cartilaginous in the first post-hatching days. Lineage-specific features—including the dorsal process of the pterygoid and the rostral process of the paroccipital process—ossify later. The rostral process originates in the exoccipital, with minor contribution from the squamosal, supporting its reinterpretation as processus rostralis paroccipitalis (new term). The lacrimal is consistently absent, indicating a true secondary loss probably linked to cranial kinesis. The mesethmoid may contribute to the frontal overhang in species where present. The epiotic forms part of the external cranial vault, also reported in other birds, suggesting it is more widespread than previously assumed. Cervical vertebrae, in turn, follow the altricial pattern, with late ossification of the atlas and unfused neural arches at early stages, showing considerable heterogeneity among altricial birds. Collectively, these findings show how postnatal skeletal development integrates functional, mechanical, and evolutionary constraints, offering new insights into the ontogeny and specialisation of the woodpecker skull and neck.

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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.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.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.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.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.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.e139103

Authors: Anna B. Vassilieva, Trung Duc Nguyen, Pavel A. Sorokin

Abstract: Abstract Dicroglossid tadpoles are an important element of pond and stream communities in Southeast Asia. They are often found in syntopy but belong to different trophic guilds, from generalized detritophages to specialized predators. In the present research, the external and internal larval morphologies of seven anuran species of the family Dicroglossidae distributed in Vietnam (Fejervarya limnocharis, Fejervarya moodiei, Hoplobatrachus rugulosus, Limnonectes dabanus, Occidozyga lima, Occidozyga martensii, and Quasipaa verrucospinosa) are described and compared. An integrated approach is followed in the study by describing external and mouthparts morphologies, gross structure of the digestive tract, and the structure of chondrocranium and hyobranchium, which revealed the main trophic adaptations of tadpoles. Different trends in the evolution of larval macrophagy in Dicroglossidae led to the formation of fundamentally different larval forms. In the first case, the tadpoles are opportunistic predators that retain the ability to use a wide range of food resources (Hoplobatrachus); they have extremely specialized mouthparts but moderately transformed digestive tract and larval skeleton. In the other case, tadpoles are highly specialized obligate carnivores (Occidozyga) that have completely lost omnivory, with a radically transformed digestive tract, chondrocranium and hyobranchium. A comparison with generalized tadpoles revealed that in dicroglossids, the transformations of the larval skeleton due to carnivory mainly affect the labial cartilages and the palatoquadrate cartilage, as well as the hyobranchium. These transformations are common to macrophagous tadpoles of different taxonomic groups and result in greater larval morphological diversity.

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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.68.e176207

Authors: Mariana Beatriz Julieta Picasso, María Clelia Mosto

Abstract: Caracaras (Aves, Falconiformes, Falconidae) are Neotropical diurnal raptors that belong to the subfamily Polyborinae. The forelimb myology of this group has not been comprehensively studied or compared with that of other Falconidae. Thus, the aims of this study were i) to describe the forelimb muscles of two species of Polyborinae (Caracara plancus and Milvago chimango), ii) to explore the possible relationship between muscular features and their function in flight behavior and iii) to compare the muscular features of these species with those of species of the subfamily Falconinae. To this end, the forelimb muscles of C. plancus (n = 4) and M. chimango (n = 4) were dissected. Additonally, to complement this data set, two specimens of M. chimachima were dissected. The mass of each muscle of one wing and its percentage with respect to the body mass were obtained. A total of 45 muscles were identified, and differences with respect to Falconinae were related to the presence of single or additional bellies. The total forelimb muscle mass represented between 7.68 and 10.26 % of the body mass. The muscle pectoralis represented ̴ 5% of the body mass, followed by the muscles scapulohumeralis caudalis (0.64 – 0.79%), deltoideus major (0.43 – 0.53%), supracoracoideus (0.34 – 0.38%) and biceps brachii (0.26 – 0.39%). The high values of these muscles are in agreement with their important function: they are involved in the downstroke and upstroke phases of the flapping flight. On the other hand, the muscles that seemed to contribute little to the mechanical power for flight presented low values that ranged between 0.01 and 0.25%. Comparison of the forelimb muscles of caracaras with published data on Falconinae species suggests that their muscular features might be associated with their type of flight, which is more erratic and less powerful than that of Falconinae.

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