DOI: 10.3897/zse.102.187659

Authors: Yuri Gomes Abrantes, Sergio Maia Queiroz Lima, Dalton Tavares Bressane Nielsen, Fabio Origuela de Lira, Márcio Joaquim da Silva, Waldir Miron Berbel-Filho

Abstract: Based on morphological and molecular evidence, this study describes a new species of rivulid of the genus Laimosemion from the lower Rio Amazonas Basin in the south of the state of Amapá in northern Brazil. The new species, Laimosemion laranja sp. nov., exhibits a color pattern characteristic of the L. geayi species group, consisting of longitudinal rows of spots on the anterior portion of flank and oblique chevron-shaped bars on the posterior portion of flank. It can be distinguished from its congeners by unique combination of characters, including position of dorsal-fin origin relative to anal-fin rays, frontal squamation pattern of head, counts of transverse and longitudinal scale series, fin-ray counts, vertebral count, osteology of caudal fin, and number of neuromasts in infraorbital series. Partial sequences of the mitochondrial cytochrome c oxidase subunit I (cox1) gene indicates that the new species diverges genetically by 5.3% from its closest relatives, L. agilae and L. strigatus, by 9.2% from L. dibaphus, and 12.3% from L. geayi. We further discuss the systematic relationships within the L. geayi species group and the conservation status of the new species based on IUCN criteria, including recommendations for its conservation.

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DOI: 10.3897/zse.102.200796

Authors: Ruiwen Wu, Fang Nan, Xianan Wang, Mengmeng Wu, Xiongjun Liu, Fengyue Shu, Jianmei An

Abstract: Precise species identification and classification are prerequisites for assessing and conserving the biodiversity of freshwater mussels (Bivalvia, Unionidae), which are one of the most threatened animal groups in the world. Previous classifications based on shell morphological characteristics have left Lamellidens liuovatus He & Zhuang, 2013 in a state of taxonomic uncertainty at the generic level. Especially recently, based on subjective similarities in shell morphology and the geographic proximity of their type localities, a proposed junior synonym (Pseudocuneopsis heqing) has been assigned to this species, and the species has been reclassified under Pseudocuneopsis, further exacerbating taxonomic confusion. This study re-evaluated the species and its synonymous taxon by collecting specimens from the type locality and integrating comparative morphology, geometric morphometrics, and molecular phylogenetic analysis. Morphometric analyses reveal that the shell outlines of Pseudocuneopsis heqing and Pseudocuneopsis liuovatus exhibit significant quantitative differences (MANOVA, p < 0.001). The ML and BI phylogenetic analyses based on multiple gene loci (COI, 16S rRNA, and 28S rRNA) show that P. liuovatus and P. heqing belong to different lineages, while P. liuovatus is sister to Nodularia hanensis with strong support within the genus Nodularia (PP = 0.99, BS = 98%). Based on the above evidence, we support the restoration of the valid species status of Pseudocuneopsis heqing stat. rev. and the reclassification of Nodularia liuovatus comb. nov. Although the morphological characteristics of N. liuovatus and Nodularia micheloti are extremely similar, the absence of topotypic material or molecular data from the type locality of N. micheloti prevents unequivocal confirmation of their conspecificity. Therefore, we provisionally retain N. liuovatus as a distinct species pending further study.

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DOI: 10.3897/zse.102.195486

Authors: Jianhan Cao, Zhiyong Deng, Hongmei Xiang, Yixing Xie, Cheng Wang, Hong Li, Yang Hu, Ying Wei, Wansheng Jiang

Abstract: The genus Acrossocheilus (Cyprinidae), which occurs in Southeast Asia and southern China, is of significant ecological and economic importance. A new species, Acrossocheilus zhangjiajiensis sp. nov., is described from the upper Lishui River Basin, a tributary of the Yangtze River, in Sangzhi County, Zhangjiajie City, Hunan Province, China. The description is based on an integrative taxonomic approach combining morphological examination, DNA barcoding, and mitogenomic analyses. This new species is distinguished from all congeners by the following combination of characters: a maxillary barbel extending to the midline of the eye; a slightly hardened last unbranched dorsal-fin ray with a smooth posterior margin; dorsal-fin membranes with black stripes; the second primary vertical bar (PVB2) positioned anterior to the origin of the dorsal fin; juveniles and adults possessing six distinct vertical black bars on the flank; and fewer than 40 lateral line scales. Notably, it further differs from its sister species, A. jishouensis, in several morphometric traits, as well as the texture of the last unbranched dorsal-fin ray and the persistence of vertical flank bars throughout all life stages. The complete mitogenome of A. zhangjiajiensis sp. nov. is 16,592 bp in length. Phylogenetic analyses based on the mitochondrial COI gene and the 13 protein-coding genes (PCGs) support a sister-species relationship between A. zhangjiajiensis sp. nov. and A. jishouensis, with genetic distances of 3.1%​ (COI) and 3.3% (13 PCGs). A COI-based haplotype network and species delimitation analyses further support the distinctiveness of the new species. This discovery increases the total number of valid Acrossocheilus species to 25 and provides critical data for the systematics, evolution, and conservation of this genus.

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DOI: 10.3897/zse.102.203747

Authors: George Popovici, Oana Paula Popa, Elena Iulia Iorgu

Abstract: This corrigendum addresses two errors in Popovici et al. (2026), wherein two new species of Geophilus, Geophilus captiosus and Geophilus tenuis, are described from Romania and their relationships to other congeners are investigated. In the original publication, a row of headings in Table 1, representing species names, was accidentally removed during typesetting. The correct, complete version of Table 1 is provided herein. Additionally, within Figure 12, the legend incorrectly labels records of G. tenuis as G. debilis. The corrected version of Figure 12, with an updated legend is provided herein. The authors sincerely apologize for these oversights.

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DOI: 10.3897/zse.102.177928

Authors: George Popovici, Oana Paula Popa, Elena Iulia Iorgu

Abstract: Two new species of Geophilus Leach, 1814, are identified and described from Romania. The examined material is compared with morphologically similar congeners, suggesting the presence of a complex of closely related species within the highly diverse genus Geophilus, characterized by a unique combination of diagnostic morphological and molecular characters. Two nuclear (18S, 28S) and one mitochondrial (COI) markers are sequenced for Geophilus tenuis sp. nov., and COI sequence data are provided for Geophilus captiosus sp. nov. Morphological data, phylogenetic reconstruction, and three molecular species delimitation tools (ASAP, bPTP, and GMYC) confirm their validity, with diagnostic nucleotide combinations, based on the COI barcode, additionally provided for them. Their placement in Geophilus is assessed through the most complete phylogeny of the genus hitherto constructed, using sequence data of 18S, 28S, and COI markers from all species present in public repositories. The extent of potential undescribed species diversity in Geophilus is discussed, highlighting the need for comprehensive sampling of small endogeic members of this genus.

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DOI: 10.3897/zse.102.184032

Authors: Hui-Xia Chen, Rui-Jia Yang, Shan-Shan Wu, Zhi-Jun Yu, Liang Li

Abstract: The family Isthmosacanthidae is a group of acanthocephalans mainly parasitic in fishes of Osteichthyes. A new isthmosacanthid species, Gorgorhynchus fistularius sp. nov. (Polymorphida, Isthmosacanthidae) is described based on specimens collected from the red cornetfish Fistularia petimba (Lacépède, 1803) (Syngnathiformes, Fistulariidae) using the integrated morphological characters and molecular data. The complete mitochondrial genome of G. fistularius sp. nov. is presented, which is 14,409 bp in length and represents the first mitogenomic data for the genus Gorgorhynchus and also for the family Isthmosacanthidae. Comparative mitogenomic analysis revealed that the overall A+T content (68.1%) in the mitogenome of G. fistularius sp. nov. is distinctly higher than that of the other mitogenomes of the order Polymorphida reported so far (54.5‒64.4%), but the composition and gene rearrangement of the mitogenome of G. fistularius sp. nov. are identical to the current known mitogenomes of polymorphid species except for Southwellina hispida. Mitogenomic phylogenies supported the Isthmosacanthidae as a distinct family, and suggested new insights into the interfamilial relationships occurring as Plagiorhynchidae + [Isthmosacanthidae + (Centrorhynchidae + Polymorphidae)] in the order Polymorphida.

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DOI: 10.3897/zse.102.173390

Authors: Ya-Ting Liang, Li Ding, Zheng-Jun Wu, Rui-Gang Yang, Ze-Ning Chen

Abstract: In this study, we describe a green pit viper of the genus Trimeresurus Lacépède, 1804, from Yunnan Province, China, based on both morphological and molecular evidence. Morphologically, the new species can be distinguished from other congeners by a combination of the following characteristics: (1) dorsal body grass-green, ventral body yellowish green; (2) postocular stripe absent in females; (3) ventrolateral stripe white, present on the first dorsal scale rows in females; (4) iris golden yellow in females; (5) DSR 19/20–19–15 (N = 2), VEN 145–147 (N = 2), SC 55–60 (N = 2); (6) 10–11 cephalic scales; (7) first supralabial separated from the nasal scale by a distinct suture. Phylogenetic analysis based on the mitochondrial cyt b, ND4, 12S rRNA, and 16S rRNA gene fragments indicates that the new species is genetically divergent from all congeners (BPP 1.00/UFB 100), with uncorrected genetic distances of the mitochondrial gene cyt b ranging from 3.6% (T. mayaae) to 13.1% (T. hageni, T. septentrionalis, T. guoi, and T. ayeyarwadyensis) compared with its congeners.

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DOI: 10.3897/zse.102.180695

Authors: Yuan Xiong, Huiming Chen, Luqi Huang, Xuankong Jiang, Chao Jiang

Abstract: The millipede genus Mandarinopus Verhoeff, 1933 (Sulciferini, Paradoxosomatidae), endemic to western China, currently comprises five species. Its taxonomic delimitation from Kronopolites Attems, 1914, remains controversial because of morphological similarity and overlapping distributions. Previous studies relying exclusively on morphological characters yielded ambiguous results. To clarify the taxonomic status of Mandarinopus, we conducted an integrative taxonomic study using topotypic specimens of all species of Mandarinopus and representative species of Kronopolites. Phylogenetic analyses based on four genetic markers (COI, 16S, 18S, and 28S rRNA) reveal that Mandarinopus is polyphyletic and deeply nested within Kronopolites. Notably, the type species of both genera form a strongly supported sister pair. Morphological re-examination demonstrates that the diagnostic characters previously used to distinguish the genera, specifically gonopodal features, are continuous and lack clear boundaries. Consequently, we formally propose Mandarinopus Verhoeff, 1933, as a junior synonym of Kronopolites Attems, 1914, transferring all valid Mandarinopus species to Kronopolites. Additionally, based on morphological and molecular evidence from topotypic material, we propose Kronopolites rugosus Golovatch, 2013, as a junior synonym of Kronopolites corticinus (Attems, 1936).

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DOI: 10.3897/zse.102.176342

Authors: Mao-Liang Li, Tan Van Nguyen, Jun-Jie Huang, Sabira S. Idiiatullina, Son Xuan Le, Nikolay A. Poyarkov, Patrick David, Bu-Ying Han, Wei Wu, Jia-Tang Li, Jin-Long Ren

Abstract: Hebius atemporalis (Bourret, 1934) has been reported from both northern Vietnam and southern China. Previous studies revealed deep genetic divergence between the Chinese and Vietnamese populations; however, no comprehensive taxonomic revision has examined the phylogenetic relationships and taxonomic status of these divergent lineages. Here, we present an integrative taxonomic reassessment of the H. atemporalis complex, combining detailed morphological comparisons and molecular data. As a result, we identify and describe a distinct new species, Hebius malnatei sp. nov., endemic to the Yunnan-Guizhou Plateau, China. The new species can be distinguished from H. atemporalis sensu stricto by its larger adult body size, shorter relative tail length, higher numbers of ventrals and subcaudals, and distinctive coloration characterized by the absence of a nuchal collar. Phylogenetic analyses based on cytochrome b sequences consistently recover Hebius malnatei sp. nov. as the sister taxon of H. atemporalis s. str., with uncorrected p-distances of 8.0–9.8%. According to the IUCN Red List criteria, we recommend assigning Hebius malnatei sp. nov. to the category Least Concern (LC). This study highlights the need for taxonomic revision of widely distributed Hebius species and underscores the role of the Yunnan-Guizhou Plateau uplift in promoting species diversification.

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DOI: 10.3897/zse.101.165370

Authors: Hiranya Sudasinghe, Kumudu Wijesooriya, Tharindu Ranasinghe, Rohan Pethiyagoda, Madhava Meegaskumbura

Abstract: The Palk Isthmus, which connected Sri Lanka to the Indian mainland during periods of low sea levels, has been the only route by which freshwater organisms could enter the island. The endemic aquatic biodiversity of Sri Lanka is dominated by diversifications stemming from founders that immigrated when the Isthmus was both emergent and possessed an aseasonal-mesic climate. The loach genus Schistura McClelland, 1838, among the most speciose of freshwater-fish genera, has three known representative species on the island. Based on two mitochondrial and two nuclear gene markers, we investigate the evolutionary relationships of Schistura in Sri Lanka, showing it to be a monophyletic diversification. Molecular species delimitation analysis recovered eight lineages (MOTUs). These exhibit strong geographical structure; each confined to one or more contiguous basins and two confined to a single basin. There is evidence for gene flow across watersheds, likely due to headwater river-capture. Schistura lineages in the central hills appear to result from multiple independent colonization events. The most widespread lineage in this region likely represents a more recent colonization, following a Late Pleistocene highland extirpation. An analysis of 39 phenetic traits across > 250 specimens revealed marked morpho-molecular incongruence. Taxonomy dependent entirely on phenetic traits is likely to be unreliable in rapidly diversifying populations such as these.

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DOI: 10.3897/zse.101.153239

Authors: Behare Rexhepi, Teo Delić, Gaj Kušar, François Lefebvre, Thierry Alezine, Xavier Chevillot, Marie-José Olivier, Florian Malard, Christophe J. Douady, Fabio Stoch, Jacques Chauvin, Aurélie Husté, Matthieu Fournier, Maja Zagmajster, Cene Fišer

Abstract: Niphargus is the most speciose amphipod genus inhabiting groundwater of the Western Palearctic. The genus is renowned for its notorious taxonomy, associated with large within- but often negligible between-species morphological variation. Results of molecular taxonomy suggested that morphological variation captured less than half of its species diversity, calling for a molecular revision of past taxonomic and faunistic works. Molecular assessment of species diversity within Niphargus has been conducted since the 2000s, but it remains spatially incomplete given the genus’ extensive geographic range. Here, we provide the first comprehensive assessment of the diversity of Niphargus in France using molecular markers. We sequenced the COI mitochondrial marker from 436 individuals collected from 234 locations. Using Assemble Species by Automated Partitioning, we identified 99 molecular operational taxonomic units (MOTU), among which 74 could represent potential new species. In turn, we selected one individual per MOTU, amplified nuclear markers 28S and Histone 3, aligned them with an additional 142 MOTUs from across the genus range, and reconstructed their phylogenetic relationships using the Maximum Likelihood method. We found that several nominal species occurring in France are polyphyletic. Furthermore, Niphargus from France comprised lineages derived from deep and terminal splits, some of which showed signs of evolutionary radiation. The phylogenetic relationships are consistent with the hypothesis that Niphargus originated in the western part of its range. This study addressed a knowledge gap in the molecular and spatial coverage of Niphargus, provided new barcodes for monitoring the diversity of the genus, and identified future directions for research.

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DOI: 10.3897/zse.101.156742

Authors: Fan Wu, Lei Wang, Ronald Sluys, Xin-Xin Sun, De-Zeng Liu, Zi-Mei Dong, Guang-Wen Chen

Abstract: Two new species of the genus Dugesia from southwest China are described using an integrative approach based on morphological, histological, and molecular data. Dugesia patula Chen & Dong, sp. nov. is characterized by the following features: symmetrical openings of the oviducts into the bursal canal; vasa deferentia opening symmetrically into the mid-lateral portion of the seminal vesicle; a large and pointed diaphragm; a short duct between the seminal vesicle and diaphragm; and a very broad, short ejaculatory duct with a very wide opening at the tip of the penis papilla. Dugesia postica Chen & Dong, sp. nov., is characterized by the following features: symmetrical openings of the oviducts into the bursal canal; a small penis bulb; a large and long penis papilla with a slightly larger dorsal lip and a small ventral lip; a long connecting duct between the seminal vesicle and diaphragm; a small diaphragm located near the tip of the penis papilla; and an extremely short ejaculatory duct. In the molecular phylogenetic tree, the two new species share a sister-group relationship and fall within an eastern Palearctic/Oriental clade of conspecifics. The distinct specific status of these two new species is also supported by their genetic distances. Through a multi-gene concatenated phylogenetic analysis, this study, for the first time, elucidates the evolutionary relationships among the Oriental, eastern Palearctic, and Australasian groups, enabling the development of a hypothesis on the potential dispersal routes of Dugesia into the eastern Palearctic region.

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DOI: 10.3897/zse.101.157834

Authors: Xiangjian Wu, Yuanqiang Pan, Wei Xiao, Ju Chen, Guohua Yu, Xinkui Wei

Abstract: Currently, the genus Brachytarsophrys contains nine species. In this study, we describe a new species of Brachytarsophrys, named Brachytarsophrys guilinensis sp. nov., from northeastern Guangxi, China, based on morphological, molecular, and acoustic evidence. This new species can be distinguished from other known congeners by the combination of the following morphological characters: (1) smaller body size (SVL 70.0‒81.9 mm in eight adult males); (2) head wider than long, with head width nearly 1.5 times the length and about half of the SVL; (3) tongue broad and round, feebly notched; (4) male with nuptial pad on the dorsal surface of the first and second fingers; (5) hind limbs relatively short and robust, heels not meeting when legs are positioned at a right angle to the body; (6) tibiotarsal articulation reaching the angle of the mouth when hind limbs are extended forward alongside the body; (7) absence of outer metatarsal tubercle, with the inner metatarsal tubercle oval and approximately equal to the first toe; (8) fingers without webbing; toes with incomplete webbing, webbing formula: I1 - 2II1 - 2III1½ - 3-IV3- - 1½V; (9) lateral fringes on toes wide; (10) several warts on the outer side of the upper eyelid, with one being relatively larger. Phylogenetic analyses based on mitochondrial 16S rRNA, COI, and Cytb genes revealed that this new species is more closely related to B. popei and B. orientalis than to other known species, with strong Bayesian posterior probability and moderate bootstrap support. More studies are necessary to clarify the taxonomy and species diversity of the genus Brachytarsophrys in Guangxi, China.

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DOI: 10.3897/zse.101.154682

Authors: Fatah Zarei, Melissa B. Martin, Paul H. Skelton, Albert Chakona

Abstract: Pseudobarbus burchelli (Smith, 1841) was previously considered to have a relatively wide distribution range spanning four isolated river systems in the Cape Fold Ecoregion of South Africa: the Breede, Duiwenhoks, Goukou, and Heuningnes. Extensive exploration and the application of molecular data led to the discovery and description of a new species, Pseudobarbus skeltoni, from the Breede River system, and the identification of three distinct lineages within P. burchelli. These lineages were provisionally named P. burchelli ‘Tradouw’ from the Tradouw catchment in the Breede River system, P. burchelli ‘Breede’, distributed across the Breede, Duiwenhoks, and Goukou river systems, and P. burchelli ‘Heuningnes’, distributed in the Heuningnes River system. Additionally, recent surveys found a new redfin population in the upper Klein River, genetically linked to the Heuningnes lineage. This study used an integrative approach to assess the taxonomic status of these three lineages. Detailed morphological examination identified a combination of characters that consistently separate the lineages, supporting their recognition as distinct species. Based on both genetic and morphological distinctiveness, we redescribe P. burchelli s.s. for the P. burchelli ‘Tradouw’ lineage, revalidate Pseudobarbus vulneratus (Castelnau, 1861) for the P. burchelli ‘Breede’ lineage, and describe the P. burchelli ‘Heuningnes’ lineage as a new species, Pseudobarbus agulhas sp. nov. These three species differ in circumpeduncular scale count, head and snout proportions, and the number of teeth in the outer pharyngeal tooth row. The biogeographic history, key threats, and conservation needs of these three species are discussed.

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DOI: 10.3897/zse.101.144081

Authors: Xuechun Su, Deyuan Yang, Xiu Wu, Yanan Sun, Jian-Wen Qiu, Yanjie Zhang

Abstract: Capitellidae is a family of marine annelids commonly found in coastal to deep-sea sediments. These annelids are characterized by capillary chaetae at the anterior and long-handled hooks at the posterior part. Although mitochondrial genomes (mtgenomes) are widely used in phylogenetic analyses of invertebrates, their application is limited in many marine annelid families, particularly in Capitellidae. In this study, we obtained complete or nearly complete (except control region) mtgenomes through high-throughput sequencing of eight species across five genera of Capitellidae: Barantolla sp., Capitella teleta, Mediomastus sp., Notodasus sp. A, Notodasus sp. B, Notodasus sp. C, and Notomastus sp. A and Notomastus sp. B. Our results indicate that species from genera with fewer capillary chaetae (Barantolla and Mediomastus) exhibit a relatively conserved mitochondrial gene order, while those from other genera show significant gene order rearrangements. Group II intron in cox1 is found in newly sequenced Notomastus sp. B and Notodasus sp. A & C. Phylogenetic analysis based on the 13 protein-coding genes (PCGs) or 37 mitochondrial genes (mtgenes) revealed three distinct clades for nine capitellid worms with the mtgenome: Clade 1 includes Mediomastus and Barantolla; Clade 2 consists of Notodasus and Capitella; and Clade 3 comprises Notomastus. Notably, Clade 2 is sister to Clade 3, and both form the sister group to Clade 1. In contrast, a phylogenetic tree constructed from nuclear genes (ncgenes; 18S, 28S, and H3) identified Capitella as an early branching clade within Capitellidae. The tree based on 37 mtgenes + ncgenes identified the Capitella as the sister taxon of Notodasus + Notomastus. Additionally, the Ka/Ks ratios of 13 PCGs in Mediomastus and Barantolla were much lower than those in Notodasus or Notomastus. Together, our results indicate different trajectories of mtgenome evolution in the Capitellidae.

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DOI: 10.3897/zse.101.151890

Authors: Ya-Zhen Zou, Jie Huang, Hai-Yang Xiang, Shi Li, Yan Tang, Zhi-Xiao Liu

Abstract: Gordius wulingensis sp. nov., a newly described horsehair worm, parasitizes Tachycines (Orthoptera, Rhaphidophoridae). Free-living specimens are mainly found in karstic cave puddles. In recent years, a total of 37 creamy white horsehair worms were collected from various caves in the Wuling Mountains. Characteristic for the species is that males exhibit concentrated bristles at the tips of the inner lobes and scattered bristles along the posterior half of the caudal lobe. The inner wall of the cloacal opening displays honeycombed areoles. The body cuticle contains dense bristles in the anterior and posterior ends; the density of bristles is decreasing in the mid-body. Distinct longitudinal sharp ridges are visible on the ventral and dorsal surfaces. Adults are present in caves year-round, while the entire life cycle is completed within the caves. In the laboratory, specimens can be kept alive for more than 3 months.

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DOI: 10.3897/zse.101.139075

Authors: Yan-Rong Wang, Zhong-Li Sha

Abstract: A new stenothoid species, Metopa propoda sp. nov., is described from a seamount of the Western Pacific. The new species differs from all other congeneric species by the mature males having a large rectangular notch in the palmar corner of the gnathopod 2. One mitochondrial gene (COI) was used to analyze the validity of Metopa propoda sp. nov. An identification key of the Pacific Metopa species, including this new species, is provided.

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DOI: 10.3897/zse.101.147368

Authors: Ilya S. Turbanov, Aleksey A. Bolotovskiy, Oleg N. Artaev, Aleksandr A. Gandlin, Marina A. Levina, Ekaterina D. Vasil’eva, Boris A. Levin

Abstract: A new species of cypriniform fish was discovered during studies of the riverine fish fauna of the Caspian Sea basin, employing an integrative taxonomy approach. The Ural gudgeon is herein described as a new species from the Ural River system, which is considered a geographical boundary between Europe and Asia. The new species is distinguished from the other gudgeon species by a combination of morphological characters and genetic data. Morphologically, it is very similar to Gobio volgensis from the Volga basin. Genetically, however, it is closer to G. acutipinnatus, which occurs in the upper Irtysh River (Ob’ River system), and to G. multipunctatus from the Lake Alakol basin in Central Asia (p-distance = 0.037; COI mtDNA). Therefore, the Ural gudgeon, which is endemic to the Ural River system, is of Asian rather than European origin. The biogeography of freshwater aquatic animals in the region is discussed in relation to paleogeographic events and connections between the Caspian Sea and Ob’ drainages.

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DOI: 10.3897/zse.101.137781

Authors: Chenghao Jia, Zening Xu, Fengping Li, Liuna Chen, Aimin Wang, Fei Gao

Abstract: To explore the taxonomic status of a tropical sea cucumber species with controversy, we conducted an investigation into its morphological characteristics (external form and ossicles) and mitochondrial genome and rearranged the historical materials of the species and its confusing species. The species was first described in the middle of the 18th century but later synonymized with Bohadschia ocellata Jaeger, 1833, Holothuria hamata Pearson, 1913, or Holothuria kurti Ludwig, 1891, by different scholars. The observed ossicles encompassed multiple types, including buttons, C-shaped ossicles, rods, and tables, with a distinct subtype identified as tack-like tables. Ventral and dorsal body walls exhibited similar table and button ossicle shapes. The mitogenome contained 13 protein-coding genes (PCGs), 2 ribosomal RNAs (rRNAs), and 22 transfer RNA (tRNA) genes, with a sequence length of 15,797 bp. Subsequent construction of a phylogenetic tree using Bayesian inference (BI) and maximum likelihood (ML) methods indicated that the species was grouped into a separate branch firstly with Holothuria (Theelothuria) spinifera Théel, 1886, so suggesting its taxonomic classification under subgenus Theelothuria Deichmann, 1958. Based on the data of this study and previous literature, we suggested that the Latin name for this species should tentatively be “Holothuria ocellata Jaeger, 1833 sensu Théel 1886.” The morphologic and genomic information in the current study could be valuable in further biological studies of this sea cucumber species.

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DOI: 10.3897/zse.101.142007

Authors: Qiong Wu, Yi-Tao Lin, Jian-Wen Qiu, Mei Yu Xu, Bing Peng Xing

Abstract: Mussels of the genus Idas (Bathymodiolinae) are small bivalves inhabiting organic falls and cold seep ecosystems. Despite their ecological significance, the taxonomy and phylogeny of Idas remain understudied. In this study, 30 Idas specimens were collected from a piece of sunken wood in the East China Sea at depths of 460–550 m. DNA barcoding and phylogenetic analysis revealed three distinct clades: 23 specimens matched Idas sp. ESU D, previously reported from Japan and the Philippines, five samples clustered with Idas iwaotakii (Habe, 1958), and two samples formed a new clade related to I. iwaotakii. These findings resulted in the description of two new species, Idas sinensis sp. nov. and Idas pacificus sp. nov. We present the complete mitochondrial genomes for both species and investigate the diversity of their symbiotic microbes through full-length 16S rRNA amplicon sequencing. Phylogenetic analyses of mitochondrial genomes further support the taxonomic placement of the new species. The gill-associated microbial communities of these mussels include sulfur-oxidizing SUP05 bacteria, suggesting chemosynthetic symbiosis.

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DOI: 10.3897/zse.101.142297

Authors: Shiyang Weng, Xiaolong Liu, Jianchuan Li, Guohua Yu, Junkai Huang

Abstract: A new species of Rhacophorus (Anura, Rhacophoridae), Rhacophorus medogensis sp. nov., is described from Medog, Xizang Tibetan Autonomous Region, China. The new taxon can be distinguished from all phylogenetically closely related taxa (R. rhodopus and R. bipunctatus complexes) by the following combination of features: 1) body size medium (adult males SVL 31.6–38.7 mm, n = 17; adult females SVL 50.1–55.7 mm, n = 2); 2) pineal ocellus obvious; 3) toe webbing formula: I1‒1II1‒1.5III1‒1IV1‒1V; 4) irregularly shaped large black spots, white pattern in black spots on flanks; 5) snout pointed with appendage on tip; 6) tongue pyriform, with a deep notch at posterior tip; 7) tibiotarsal articulation reaching eye. In addition, we also discuss the distribution of R. bipunctatus, which is limited to northern India and central-western Myanmar, rather than the traditionally presumed range across South and Southeast Asia.

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DOI: 10.3897/zse.101.143073

Authors: Khanet Srikampha, Thomas Wesener, Natdanai Likhitrakarn, Chirasak Sutcharit, Ruttapon Srisonchai

Abstract: Three new species of the giant pill-millipede genus Sphaerobelum Verhoeff, 1924, from Thailand and Laos, are described based on an integrative approach that combines morphological characters and COI data; these are S. nonghinensis Srikampha & Srisonchai, sp. nov., S. onyx Srikampha & Srisonchai, sp. nov., and S. petramurum Srikampha & Srisonchai, sp. nov. Additionally, new material of previously described species, S. aesculus Rosenmejer & Wesener, 2021, S. truncatum Wongthamwanich, 2012, and S. turcosa Srisonchai & Pimvichai, 2023, were collected and examined. The new species can be distinguished from all congeners by a combination of unique characters, including body colour, endotergum, vulva, and details of the telopods. The validity of these new species is further supported by high genetic distances from their congeners in the 658 base pairs of the COI barcoding fragment, ranging from 13.24–30.12%. Preliminary phylogenetic analyses using Bayesian inference and maximum likelihood indicate that each new species forms a distinct clade, supporting their morphological uniqueness, though the interrelationships with most other congeners remain unsupported. All new species are restricted to narrow distribution areas in Thailand and Laos, suggesting they are endemic to these regions. Morphological characters of the new species are illustrated, and an identification key and a distribution map for known Sphaerobelum species in Thailand are also provided.

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DOI: 10.3897/zse.101.137392

Authors: Tao Luo, Zi-Fa Zhao, Zhong-Lian Wang, Chang-Ting Lan, Ming-Yuan Xiao, Huai-Qing Deng, Ning Xiao, Jiang Zhou

Abstract: The uplift of the Qinghai-Tibet Plateau and the Indochina extrusion are two of the most prominent consequences of the India–Asia collision. These two geologic events greatly altered topography and drainage patterns that, in turn, affected the regional climate, the landscape and the evolution of biodiversity. Despite this, little is known about how orogeny and climate affect the evolution of biodiversity, especially the dynamics of diversification, including origins, peaks and endings. Here, we performed phylogenetic and biogeographic analyses of Leptobrachella distributed in Southeast Asia and southern China, based on mitochondrial 16S ribosomal RNA. The results revealed that Leptobrachella may be roughly divided into Clade I from south of the Indo-Burma and Clade II from central and northern Indo-Burma and southern China. We then investigated the diversification of Leptobrachella over time through biogeographic meta-analyses. We showed that the speciation of Leptobrachella was dominated by in situ diversification that was most likely associated with the uplift of the Qinghai–Tibet Plateau, the Indochina extrusion and the intensification of the Asian monsoon and that diversification may have been less influenced by temperature. In situ diversification experienced three small accelerated phases and one decelerated phase initiated at ~ 32 Ma, with a sharp increase at ~ 15 Ma, a peak at ~ 8.7 Ma and a gradual decline after ~ 6 Ma and the peaks of diversification were asynchronous in Southeast Asia and southern China. Our results suggest a three-phase scenario for the diversification of Leptobrachella, with periods of acceleration and deceleration at every stage, a pattern consistent with the Indochina extrusion, the uplift of the Qinghai–Tibet Plateau and the intensification of the Asian monsoon since the Oligocene. This study highlights how biogeographic meta-analyses can be utilised to estimate diversification history in taxa lacking sufficient molecular markers to quantify the impact of orogeny and climatic shifts on diversification processes. In addition, we also identified four undescribed species and described one new species, Leptobrachella xishuiensis sp. nov., from Xishui County, Guizhou Province, China.

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DOI: 10.3897/zse.100.136428

Authors: Rodrigo Brincalepe Salvador, Maria Elina Bichuette

Abstract: The freshwater gastropod fauna in Brazil presently consists of circa 180 known species, though this is deemed an underestimate. The little-studied cavern faunas have been considered a potential source of undescribed species, particularly regarding the Truncatelloidea. Here, based on new collection efforts in caves in Bahia state, Brazil, we describe two new troglobitic species of Idiopyrgus (Tomichiidae): Idiopyrgus eowynae sp. nov. and Idiopyrgus meriadoci sp. nov. We improve upon the previous molecular phylogeny of this genus and take the opportunity to reclassify Spiripockia umbraticola Simone & Salvador, 2021, as Idiopyrgus umbraticola (Simone & Salvador, 2021), comb. nov. The genus Idiopyrgus now contains ten species, reinforcing the presence of Tomichiidae in Brazil as a relict lineage that largely radiated into cave environments. Furthermore, considering that so far only snapshots of their biology are known, we have collated all the information currently available from the literature and field observations on the cavernicolous species of Idiopyrgus. Notably, due to human action in the area where the two new species have been found, they can already be considered vulnerable.

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DOI: 10.3897/zse.100.133964

Authors: Tao Luo, Fang-Wei Luo, Chang-Ting Lan, Ming-Yuan Xiao, Jia-Jun Zhou, Mei Liao, Ning Xiao, Jiang Zhou

Abstract: The genus Karstsinnectes of blind fishes known as karst loaches from China was established in 2023 during a revision of the genus Oreonectes (Nemacheilidae). Six species are recorded in this study and some taxonomic progress has been made; however, the lack of genetic data for some species (e.g., K. anophthalmus) may have weakened our current understanding of Karstsinnectes. This study reassessed the phylogeny and evolutionary history of Karstsinnectes by integrating a combination of previously published and newly sequenced mitochondrial genomic data. A phylogenetic tree was developed that was able to divide Karstsinnectes into two clades corresponding to drainages and clarify the phylogenetic position of K. anophthalmus. Divergence times show that Karstsinnectes originated at the Oligocene/Miocene boundary (~22.37 Mya), with the most recent common ancestor occurring in the early Miocene (~18.87 Mya) and interspecific divergence occurring in the late Miocene. Ancestral area reconstruction suggests that the most recent common ancestor of Karstsinnectes most likely inhabited the Hongshui River basin and dispersed into the Zuojiang-Yujiang, Beipanjiang, and Youjiang river basins during the early Miocene (~18.87 Mya), middle Miocene (~12.78 Mya), and late Miocene (~6.71 Mya), respectively. The dispersal under the influence of orogenesis and a monsoon climate drove the speciation and diverse distribution of Karstsinnectes. Such findings are important for conservation considering that Karstsinnectes strictly inhabits deep caves. Additionally, the taxonomic status of the distributed Karstsinnectes population in Leiping Town, Daxin County, Guangxi, China was revised by combining genetic and morphological differences to describe this population as a new species, Karstsinnectes daxinensis Luo, Zhou & Zhou, sp. nov. The definition of the phylogenetic position of K. anophthalmus emphasizes the importance of using type locality material for the identification of cryptic species.

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DOI: 10.3897/zse.100.120872

Authors: Diego J. Santana, João Victor A. Lacerda, Priscila S. Carvalho, Manuella Folly, Bruno Bove da Costa, Iuri Ribeiro Dias, Luiz Fernando Carmo, Henrique C. Costa, Donald B. Shepard, Clarissa Canedo

Abstract: As many new evolutionary lineages are being discovered and formally named, sequencing topotypes when holotypes are not available becomes essential for taxonomy. This study uses a DNA-taxonomy approach to sequence new populations of the Ischnocnema verrucosa species complex (Brazilian Wart Frogs) from different locations, including, for the first time, individuals from the type localities. Phylogenetic analysis of the mitochondrial 16S gene recovered a monophyletic Ischnocnema verrucosa species series composed of three main clades. The most recent common ancestor was estimated to be 33.76 million years ago, and diversification within the three main clades occurred primarily during the Miocene. We delimited eight species-level lineages with high levels of sequence divergence (7% to 16%). Our study highlights the importance of DNA taxonomy and the necessity of protecting and sequencing topotypes in taxonomic studies. Our study also contributes to the conservation and understanding of the genus Ischnocnema and the biodiversity of the Brazilian Atlantic Forest region.

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DOI: 10.3897/zse.100.130676

Authors: Zhong-Guang Chen, Yu-Ting Dai, Xiao-Ping Wu, Jiao Jiang, Shan Ouyang

Abstract: Our study contains the first molecular phylogeny of Chinese enids based on the mitochondrial markers cytochrome oxidase c (COI) and 16S rRNA (16S). We have sequenced 19 species belonging to 10 out of the 12 currently accepted genera. A new species, Petraeomastus limenghuai Chen, Dai, Wu & Ouyang, sp. nov., is described from the Lancangjiang River Valley in southwest China based on comparative morphology and molecular phylogeny. The currently accepted classification of Chinese enids relies largely on shell morphology and is deemed to require systematic revision.

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DOI: 10.3897/zse.100.129455

Authors: Arthit Pholyotha, Chirasak Sutcharit, Somsak Panha, Piyoros Tongkerd

Abstract: The Malay Peninsula has traditionally been considered to harbour a diverse land-snail fauna, both in terms of a high species richness and a wide variety of conchological traits, especially within the limacoid land snails. A recent survey along the Malay Peninsula of southern Thailand discovered an overlooked limacoid taxon “Trochomorpha” sculpticarina Martens, 1883, previously assigned to genus Trochomorpha of the Trochomorphidae. This genus is herein described as Janbinmorpha gen. nov. based on comparative studies of shell morphology, external features, genital anatomy and radular morphology, as well as analyses of partial sequences of two mitochondrial markers, COI and 16S rRNA, and of one nuclear marker, 28S rRNA. This new genus is characterised by a combination of distinct morphological and anatomical features. The most distinguishing features are a depressed trochiform shell with a keeled last whorl, gametolytic organ without a duct, gametolytic sac with two lobes, and proximal epiphallus encircled with a thick sheath and attached by the penial retractor muscle. In addition, an analysis of the differentiation in mitochondrial and nuclear markers confirmed that this new genus, first recognised by morphology, is also genetically distinct. The molecular data also confirm that J. sculpticarina comb. nov. is a member of the Ariophantidae and has a close evolutionary relationship to Hemiplecta and Maelamaodiscus.

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DOI: 10.3897/zse.100.130661

Authors: Zhong-Guang Chen, Zhe-Yu Chen, Ran-Xi Lin, Yu-Ting Dai, Xiao-Ping Wu, Jiao Jiang, Shan Ouyang

Abstract: A molecular phylogenetic study was conducted on genus Tropidauchenia, based on COI and 16S sequences. A total of six out of 18 species in the genus, as well as three species of Grandinenia and eight species of subfamily Synprosphyminae and Phaedusinae were sequenced. The phylogenetic results supported the monophyly of three Clausiliid subfamilies distributed in East Asia and demonstrated that Tropidauchenia was divided into two clades that corresponded to its two main distribution areas. A new species, Tropidauchenia jiangjilini Z.-G. Chen, Z.-Y. Chen & R.-X. Lin, sp. nov., from Guangdong, southern China, has been identified and described based on morphological comparison and molecular phylogeny. The discovery expanded the distribution range of Tropidauchenia and revealed the potential species diversity of land snails in the karst region of Guangdong.

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DOI: 10.3897/zse.100.117859

Authors: Zi-Ming Yuan, Wei Jiang, Zhong-Li Sha

Abstract: A new genus and species within the family Xanthidae MacLeay, 1838, are described from coral reefs in the South China Sea. The new genus, Gothus, with its type species G. teemo sp. nov., is distinguishable from allied genera by characteristics of the carapace, chelipeds, and male pleon. Based on morphological evidence, we tentatively place this genus within the subfamily Euxanthinae Alcock, 1898. However, molecular systematic analysis based on COI, 12S, 16S, and H3 indicates that it does not form a stable monophyletic group with any related subfamily. Another species, Actaeodes consobrinus (A. Milne-Edwards, 1873), is also reclassified into this new genus, based on both morphological and molecular evidence.

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DOI: 10.3897/zse.100.124370

Authors: Jing-Chen Chen, Jia-Jia Li, Wen-Qiao Tang, Xin-Rui Pu, Hao-Tian Lei

Abstract: Two new species, Beaufortia granulopinna and Beaufortia viridis, are described from the upper Pearl River system in southwest China. Both species share the characteristics of the Beaufortia pingi species group, including prominent vertical stripes on the lateral body and pinnate-type lower lips, distinguishing them from other Beaufortia species. Beaufortia granulopinna is differentiated from other species in the group by possessing a unique set of characteristics: the presence of well-developed prominent tubercles on the first 6–9 pectoral fin rays in adults; and a significant proportion (54.76%) of individuals experiencing blurriness or absence of vertical stripes in the mid-section of the lateral body upon reaching adulthood. The minimum interspecific genetic distance within the genus based on mitochondrial cytb gene sequences is 10.80%. Beaufortia viridis is distinguished from other species in the group by consistently exhibiting vertical stripes of uniform length, width, and spacing across all stages of growth; the absence of tubercles on the branched rays of pectoral fins; and a body coloration of dark cyan to green. The minimum interspecific genetic distance within the genus based on mitochondrial cytb gene sequences is 4.60%. Molecular phylogenetic results confirm that the Beaufortia pingi species group forms a monophyletic clade, which is congruent with morphological classification findings. This study also addresses and resolves the taxonomic ambiguity surrounding Beaufortia pingi and Beaufortia zebroida, providing a redescription of these taxa.

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DOI: 10.3897/zse.100.125798

Authors: Ya-Ting Liang, Zi-Dan Huang, Li Ding, Gernot Vogel, Natalia B. Ananjeva, Nikolai L. Orlov, Sheng-Chao Shi, Zheng-Jun Wu, Ze-Ning Chen

Abstract: The reed snakes of the genus Calamaria Boie, 1827 are one of the largest groups of Asian snakes, distributed from northeast India to the Maluku Islands of east Indonesia. Recent research on the genus in China has revealed that the species diversity of the group was underestimated. In this study, morphological comparisons and mitochondrial DNA analysis revealed that a junior synonym of C. pavimentata Duméril, Bibron & Duméril, 1854 — Calamaria berezowskii Günther, 1896 is valid, hence we redescribed and recovered the validity of C. berezowskii. This species can be distinguished from other congeners by the combination of the following characters: four supralabials; one preocular; rostral shield width larger than height; mental not touching anterior chin shields; eye diameter less than the distance from eye to mouth edge; less than 1/2 of the posterior chin shield meets in the midline; dorsal scales reduced to six rows at tail; indistinct light ring present in the nuchal region or a more or less distinct yellowish collar. Phylogenetically, this species is sister to C. pavimentata, with significant genetic differences (0.190) on mitochondrial gene Cyt b.

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DOI: 10.3897/zse.100.120224

Authors: Ping-Shin Lee, Ben Liu, Meng Ouyang, Ren-Da Ai, Xiao-Long Liu, Yan-Hong He, Ping-Qian Huang, Ying-Chun Li, R. S. Naveen, Zhi-Yong Yuan, Jin-Min Chen

Abstract: The Gaoligong Mountains are characterized by large variations in elevation and topography, which support high levels of biodiversity and endemism that remain largely understudied. Herein, based on the integration of morphological comparisons and phylogenetic reconstruction, we describe a new species of Rhacophorus from the northern Gaoligong Mountains, Yunnan Province, China. The new species, Rhacophorus dulongensis sp. nov., is morphologically distinguishable from its congeners based on the differences in body size, head length, tibia length, snout and tongue shape, toe webbing formula and coloration, ventral skin texture and coloration, dorsal pattern and coloration, body macroglands, iris coloration, and pattern of markings on flanks. Phylogenetically, it differs from its congeners by uncorrected p-distances of >4.8% for the 16S rRNA gene fragment. Rhacophorus dulongensis sp. nov. is likely to be found in Myanmar, considering its type locality lies close to the China-Myanmar border. The phylogenetic analysis revealed that the “widespread” species, R. rhodopus, is a species complex and a composite of five distinct lineages. The results revealed that R. napoensis is also found in Vietnam, making it a new country record for Vietnam. Interestingly, R. dulongensis sp. nov. likely breeds in bamboo, a hidden behavioral characteristic that makes them easy to overlook. Given the ongoing habitat loss and degradation in the region, further biological exploration is urgently needed in the Gaoligong Mountains as a biodiversity reservoir.

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DOI: 10.3897/zse.100.118000

Authors: Wilson J. E. M. Costa, Caio R. M. Feltrin, José Leonardo O. Mattos, Axel M. Katz

Abstract: Colour patterns are diverse in trichomycterine catfishes and are often used to diagnose species. Here, we analyse the first case of adults of two syntopic species of Trichomycterus sharing nearly identical colour patterns: a rare new species of the subgenus Paracambeva and Trichomycterus maculosus, a distantly related species of the subgenus Trichomycterus. Both species are endemic to the upper Rio Paraíba do Sul basin (RPSB), which had a different course until the Tertiary period and is situated within the Southeastern Brazilian Continental Rift, mostly active in the Eocene-Oligocene. A time-calibrated multigene analysis, 3144 bp, supported the new species as sister to Trichomycterus itatiayae, both comprising a lineage with Middle Miocene age, when that colour pattern would have first arisen. The new species is diagnosed by characters from the latero-sensory system and bone morphology. Our results, combined with available biogeographical data, indicated the colour pattern of T. maculosus arising in the Late Pliocene, following the dispersal of its group to the upper RPSB after river course changing. Two hypotheses for the independent origin of the same colour pattern are discussed. First, a case of evolutionary convergence for adaptation to live on a similarly coloured gravel substrate, giving some cryptic advantage against predators. Second, mimetic association through anti-predation features. In the latter case, although trichomycterids lack fin spines to inoculate venom as in other catfishes, the species here studied have a supposed axillary gland above the pectoral fin, just posterior to the opercular odontodes, but with properties and functions still unknown.

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DOI: 10.3897/zse.100.122293

Authors: Florencia Matusevich, Valeria Gabbanelli, Gonzalo Vulcano, Natalia Plá, Victoria M. Lenain, Diego M. Vazquez, Juan M. Díaz de Astarloa, Ezequiel Mabragaña

Abstract: Currently, the genus Zenopsis, also known as silver John Dory, comprises at least five valid species with a wide range of distribution. However, recent studies have proposed the existence of a new Zenopsis species inhabiting the Indian Ocean, and a preliminary search in the Barcode of Life Database reveals the presence of different barcode index numbers (BIN) for the nominal species Zenopsis conchifer. In the Southwest Atlantic Ocean (SWA), Z. conchifer is the only species reported so far. Therefore, the aim of this work was to evaluate, at the molecular level, the potential taxonomic diversity within the genus Zenopsis and to assess if the species occurring in the SWA corresponds with Z. conchifer. Using data available in worldwide genetic databases, a maximum likelihood tree, a BIN, and an automatic barcode gap discovery analysis were carried out. Additionally, specimens sampled from the SWA were morphologically compared with specimens from different parts of its distribution using available data. The specific identity at the molecular level of specimens occurring in the SWA was confirmed as Z. conchifer. The results of the molecular analysis highlight the existence of hidden specific diversity within the genus.

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DOI: 10.3897/zse.100.122472

Authors: Zhong-Guang Chen, Yan-Shu Guo, Yu-Ting Dai, Xiao-Chen Huang, Jun-Hao Huang, Jiao Jiang, Shan Ouyang, An-Xiang Wen, Xiao-Ping Wu

Abstract: A new catfish species, Liobagrus chenhaojuni Chen, Guo & Wu, sp. nov., is described from the Tiaoxi River, a tributary of Taihu Lake, located in Zhejiang Province, China. This description is based on morphological characteristics and phylogenetic analysis. This species belongs to a group defined by the presence of a smooth posterior edge of the pectoral-fin spine and can be distinguished from other species in the group by a unique combination of characteristics, including: an upper jaw longer than the lower jaw; maxillary barbels reaching the middle of the pectoral fin; irregular blotches present on the lateral body; a rounded caudal-fin with a length ranging from 16.5% to 19.9% of the standard length; 39 to 41 post-Weberian vertebrae; and 15 to 17 anal-fin rays. The validity of this new species is further supported by the molecular phylogenetic analysis based on Cytb sequences.

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DOI: 10.3897/zse.100.122288

Authors: Dragan Antić, Thomas Wesener, Nesrine Akkari

Abstract: Based on the study of freshly-collected material and old museum specimens, we have solved a decades-old riddle surrounding the name Onychoglomeris herzogowinensis (Verhoeff, 1898). The southern Dinaric coastal species Glomeris herzogowinensis Verhoeff, 1898 is revived, while Onychoglomeris herzogowinensis australis Attems, 1935 and O. h. media Attems, 1935, are treated here as full species after returning the specific name to Glomeris Latreille, 1902, O. australis Attems, 1935, stat. nov. and O. media Attems, 1935, stat. nov. Besides the designation of lectotypes, we provide comprehensive illustrations, diagnoses, detailed remarks and a distribution map for all three species. In addition, DNA barcoding provided COI sequences for Glomeris herzogowinensis and Onychoglomeris australis stat. nov., along with the first barcoding data of one additional species of Onychoglomeris Verhoeff, 1906, O. ferraniensis Verhoeff, 1909 and two Glomeris species, the Balkan G. balcanica Verhoeff, 1906 and the trans-Adriatic G. pulchra Koch, 1847. The significance of historical specimens from natural history museums is briefly discussed.

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DOI: 10.3897/zse.100.119915

Authors: Davut Turan, Sadi̇ Aksu, Sali̇m Serkan Güçlü, Gökhan Kalaycı

Abstract: The taxonomic status of the Oxynoemacheilus from the Karınca and Havran streams in the north Aegean Basin was evaluated, and it was concluded that these populations contain a new species. The new species, Oxynoemacheilus kottelati sp. nov., is distinguished by a body with a marbled pattern, a deeper caudal peduncle, a shorter caudal peduncle, a wider interorbital distance, and a shorter middle caudal-fin lope. Oxynoemacheilus kottelati sp. nov. is differentiated from the closest species, O. marmaraensis, in possessing 51 nucleotide substitution sites, a genetic distance of 8.40%, the presence of an axillary lobe at the base of the pelvic fin (vs. absent), and a narrower median incision in the upper lip (vs. absent). Three species delimitation tests (ASAP, ABGD, and PTP) and phylogenetic analyses reinforce the validity of O. kottelati sp. nov. as a distinct species.

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DOI: 10.3897/zse.100.114482

Authors: Hai-Long Liu, Da-Hao Lin, An-Tai Wang, Zhang-Li Hu, Yu Zhang

Abstract: A new species of the polyclad genus Prosthiostomum is described from the intertidal zone of the South China Sea, Huidong, China, based on morphological and molecular analyses. Prosthiostomum huidongense sp. nov. is characterized by i) few marginal eyes scattered between the marginal band and the cerebral eyes; ii) sucker located at two-thirds of the body length, being removed from the female gonopore by twice the distance between the male and female gonopores; iii) shallow male atrium with slightly ruffled inner wall, positioned approximately perpendicular to the body wall. Molecular phylogenetic analyses based on 28S rDNA sequence showed that the new species was nested in a clade composed of Prosthiostomum species. The uncorrected p-distance of COI between P. huidongense sp. nov. and other Prosthiostomum species ranged from 20.3 to 24.3%, and the high genetic divergence further supports P. huidongense as a new species.

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DOI: 10.3897/zse.100.113972

Authors: Yanan Yu, Yingyi Jiao, Junlong Zhang

Abstract: The present study describes a new species within the genus Cultellus Schumacher, 1817 collected from the South China Sea. An integrative taxonomic approach incorporating morphological comparisons, geometric morphometrics and genetic analyses was used to identify and differentiate the new species. Cultellus exilis sp. nov. is distinguished from its congeners by its slender, fragile and translucent valves, curved posteroventral margins and relatively large protractor scars. The geometric morphometric analyses, based on outlines data, indicated that samples of Cultellus exilis sp. nov. clustered together and were distinctly separated from other species. Multiple species delimitation results, based on the mitochondrial COI gene, support the separation of Cultellus exilis sp. nov. from its related congeners. Phylogenetic analyses of a nuclear (28S rRNA) and two mitochondrial (COI, 16S rRNA) genes using Maximum Likelihood and Bayesian Inference methods revealed that the species belongs to the genus Cultellus. The superfamily Solenoidea Lamarck, 1809, which includes the families Solenidae Lamarck, 1809 and Pharidae H. Adams & A. Adams, 1856, exhibits closer affinity to the family Hiatellidae Gray, 1824 than to Solecurtidae d’Orbigny, 1846. Furthermore, we found that the genus Siliqua was clustered alongside the genera Ensiculus and Phaxas as a sister clade, which contradicts the current systematics of the subfamily within the family Pharidae. This work highlights the utility of integrative taxonomy for species identification, recognition and phylogenetic investigation.

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DOI: 10.3897/zse.100.119297

Authors: Taro Jonishi, Takafumi Nakano

Abstract: Centipedes of the genus Scolopocryptops Newport, 1844 are blind species mostly described from the New World and East Asia. In this study, a Japanese species, S. quadristriatus (Verhoeff, 1934), which is characterised by four longitudinal keels on the tergites, is re-described, based on the likely holotype preserved in the Zoologische Staatssammlung München and specimens newly collected from near the type locality. In addition, S. longisetosus sp. nov., a new species that bears tergal keels like S. quadristriatus, is described from the Ryukyu Islands in Japan and Taiwan. Although the presence of four keels on tergites is unique to these two species, phylogenetic analyses using nuclear and mitochondrial markers showed that S. longisetosus sp. nov. is not sister to S. quadristriatus. The obtained phylogeny indicates that the tergal longitudinal keels evolved in parallel within Scolopocryptops or that the presence of keels represents a plesiomorphic character of the clade containing these species.

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DOI: 10.3897/zse.100.113850

Authors: Yuanqiang Pan, Mian Hou, Guohua Yu, Shuo Liu

Abstract: We described herein Zhangixalus yunnanensis sp. nov., a new treefrog species from central and western Yunnan, China, which had previously been confused with Z. nigropunctatus, based on morphological and molecular evidence. Our phylogenetic analyses revealed that the new species is sister to the clade of Z. nigropunctatus and Z. melanoleucus with strong support (100% and 73% for BI and ML, respectively). Our morphological analysis suggested that Z. yunnanensis sp. nov. is distinctly different from all known congeners by the combination of the following morphological characters: black blotches on body flank and hind-limb, medium body size (SVL31.3–36.0 mm in males and 47.6–48.6 mm in females), head wider than long, iris yellowish-brown, dorsum uniformly green, vocal sac external, throat black, webbing greyish and fingers webbed one third and toes webbed half. Additionally, we revealed that the specimens ROM 38011 (Sa Pa, Vietnam) and VNMN 4099 (Son La, Vietnam) are neither Z. dorsoviridis nor Z. nigropunctatus, but probably represent one or two cryptic species of Zhangixalus pending further morphological and molecular data. Including the new species described herein, the genus Zhangixalus currently comprises 42 species, 30 of which are distributed in China with 11 species known from Yunnan. Amongst different zoogeographic regions in Yunnan, south-eastern Yunnan has the highest diversity of Zhangixalus, followed by western Yunnan and southern Yunnan. More studies are required to clarify the species diversity of this genus based on multiple lines of evidence (e.g. morphological and molecular data).

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DOI: 10.3897/zse.100.115692

Authors: Matteo Garzia, Daniele Salvi

Abstract: The giant deep-sea oyster Neopycnodonte zibrowii Gofas, C. Salas & Taviani, 2009 is a keystone deep-sea habitat builder species. Discovered about fifteen years ago in the Azores, it has been described and assigned to the genus Neopycnodonte Fischer von Waldheim, 1835 based on morphological features. In this study, we generated DNA sequence data for both mitochondrial (COI and 16S) and nuclear (ITS2 and 28S) markers based on the holotype specimen of N. zibrowii to establish a molecular phylogenetic framework for the systematic assessment of this species and to provide a reliable (i.e., holotype-based) reference sequence set for multilocus DNA barcoding approaches. Molecular data provide compelling evidence that the giant deep-sea oyster is a distinct species, rather than a deep-water ecophenotype of Neopycnodonte cochlear (Poli, 1795), with extremely high genetic divergence from any other gryphaeid. Multilocus phylogenetic analyses place the giant deep-sea oyster within the clade “Neopycnodonte/Pycnodonte” with closer affinity to N. cochlear rather than to P. taniguchii Hayami & Kase, 1992, thus supporting its assignment to the genus Neopycnodonte. Relationships within this clade are not well supported because mitochondrial variation is inflated by saturation that eroded phylogenetic signal, implying an old split between taxa within this clade. Finally, the set of reference barcode sequences of N. zibrowii generated in this study will be useful for a wide plethora of barcoding applications in deep-sea biodiversity surveys. Molecular validation of recent records of deep-sea oysters from the Atlantic Ocean and the Mediterranean Sea will be crucial to clarify the distribution of N. zibrowii and assess the phenotypic variation and ecology of this enigmatic species.

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DOI: 10.3897/zse.99.102604

Authors: Aoi Tsuyuki, Yuki Oya, Hiroshi Kajihara

Abstract: In this study, we describe a new species of elongated marine flatworm, Eucestoplana ittanmomen sp. nov., collected from the intertidal zone of the Okinawa Islands, Japan. Eucestoplana ittanmomen sp. nov. is distinguished from other congeners based on the following characteristics: i) its translucent body lacking coloration, ii) its dome-shaped penis sheath, iii) the absence of cilia on the inner wall of the male atrium except outside the penis sheath, and iv) the presence of an adhesive organ at the posterior end of the body. Additionally, we report the occurrence of E. cf. cuneata (Sopott-Ehlers & Schmidt, 1975) in Japan; E. cuneata has previously been documented in the Galapagos and Fiji Islands. We conducted phylogenetic analyses to infer the positions of the two Eucestoplana species within Cestoplanidae using a concatenated dataset comprising partial 18S and 28S rDNA sequences from E. cf. cuneata and E. ittanmomen sp. nov. from Japan, as well as four known Cestoplana species with sequences available in public databases. Our phylogenetic analyses revealed that Cestoplana and Eucestoplana were reciprocally monophyletic. Furthermore, the genetic distance of the 16S rDNA sequences supported the genetic independence of the two sister species, E. cf. cuneata and E. ittanmomen sp. nov.

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DOI: 10.3897/zse.99.101943

Authors: Jiří Šmíd, Sergio Matilla Fernández, Hassan Sh Abdirahman Elmi, Tomáš Mazuch

Abstract: The biological diversity of the Horn of Africa is one of the least studied in the world. Yet the Horn supports rich communities of species that are mostly endemic to the region. Here we study the diversity of Sand Snakes (Psammophis) in East Africa, their phylogeny and systematics. Previous studies have unveiled several cryptic and potentially undescribed species of Psammophis that occur in the Horn and their taxonomic status has remained unclear to this day. We used sequence data from two mitochondrial and one nuclear genes to reconstruct the phylogeny of the genus, in which we included newly obtained samples of six different Psammophis species from Somalia, Ethiopia, Eritrea, Sudan, and Egypt. Our aim was to assess the status of some of the undescribed species, examine the level of intraspecific genetic variation within individual species, improve our understanding of the species distributions, and contribute to the taxonomy of the genus. Our results confirm the existence of two undescribed species, one in eastern Somalia, which we formally describe as new, and one in southern Ethiopia that we refer to as Psammophis cf. sudanensis in accordance with previous studies. Further, we provide first genetic data for the nominotypical subspecies of P. punctulatus and confirm the species status for its subspecies P. trivirgatus. In addition, we provide new genetic data for P. tanganicus from Ethiopia and Somalia, and range extension records for P. rukwae from Eritrea and Ethiopia and for P. aegyptius from Somalia. Our findings contribute considerably to our understanding of the diversity and distribution of Psammophis in East Africa.

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DOI: 10.3897/zse.99.99285

Authors: John T. D. Caleb, Ayush Parag, Aniruddha Datta-Roy

Abstract: A new chrysilline jumping spider species belonging to the genus Siler Simon, 1889 is described from Odisha, India. Detailed morphological descriptions, illustrations of the male palp and female genitalia and phylogenetic relationships of the new Siler species are presented. Phylogenetic analysis reveals that the new species is sister to a clade of predominantly Southeast Asian Siler species. Furthermore, the results indicate the presence of multiple cryptic species masquerading as S. semiglaucus sensu lato. We also briefly discuss some unique behavioural observations on the newly-described species.

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DOI: 10.3897/zse.99.99030

Authors: Tuangthong Boonmachai, Elizabeth A. Bergey, Nattawadee Nantarat

Abstract: The micro land snail genus Diplommatina (family Diplommatinidae) is widely distributed in Southeast Asia and includes many endemic species. Three new species of Diplommatina are described from Satun Province in southern Thailand. Diplommatina bulonensis sp. nov., D. laemsonensis sp. nov. and D. prakaiphetensis sp. nov. are distinguished from other species in the genus by their shell size and shape, the number of radial ribs on the penultimate whorl, the number of whorls, and features of the peristome. The agreement between phylogenetic tree based on analyses of COI and 16S sequences and comparative morphology support the delineation of these new species which, when compared to related species, belong to well-differentiated clades. The K2P distance between any of the three new species and other Diplommatina species included in the molecular phylogenetic analysis was at least 5.5% in COI and 3.9% in 16S. Two of the three new species (D. prakaiphetensis sp. nov. and D. bulonensis sp. nov.) are apparently endemic to Prakaiphet Hill and Bulon Pai Island, respectively. Additionally, we documented a new regional record for D. naiyanetri in Satun Province. These new species and records contribute to the knowledge of Thailand’s land snail biodiversity and highlight the need of conservation protections for regional karst habitats.

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DOI: 10.3897/zse.99.96386

Authors: Ehsan Damadi, Faezeh Yazdani Moghaddam, Mehdi Ghanbarifardi

Abstract: The subfamily Plectorhinchinae (sweetlips) is composed of poorly-known species with high commercially and ecologically values that exhibit phenotypic plasticity and various morphologies. Few studies have assessed the validity of sweetlips, intergeneric relationships and evolutionary survey in this subfamily, which have not yet been resolved. This study investigated the DNA sequences of (1) the mitochondrial COI gene to delimit species, and (2) two mitochondrial (COI and Cyt b), and one nuclear (RAG1) markers to infer phylogenetic relationships and evolutionary and biogeographic history. The molecular results could differentiate Diagramma punctatum from the other species, but failed to distinguish D. labiosum as a distinct species with considerably lower genetic distances for the COI (0.53%) and Cyt b (0.51%) markers. However, additional taxonomic investigations are required to shed light on this issue. All previously described nominal species of sweetlips in the northwest Indian Ocean were found to be well supported. The monophyly of Plectorhinchus is not supported and Diagramma pictum and D. punctatum should be assigned to the genus Plectorhinchus. The biogeographic history of Plectorhinchinae likely originated in the Indo-Pacific ca. 34 Ma (30–39 Ma; late Eocene/ middle Oligocene) and subsequently colonised the Western Indian Ocean and the Central Indo-Pacific. Maximum diversification within the subfamily occurred from the middle Miocene to Pliocene, coinciding with dispersal and vicariance events. Diversification was probably driven by both biological and geographical factors.

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DOI: 10.3897/zse.99.90784

Authors: Ekaterina S. Konopleva, Ivan N. Bolotov, Ilya V. Vikhrev, Khamla Inkhavilay, Mikhail Yu. Gofarov, Alexander V. Kondakov, Alena A. Tomilova, Yulia E. Chapurina, Tu Van Do, John M. Pfeiffer, Manuel Lopes-Lima, Arthur E. Bogan

Abstract: Freshwater mussels belonging to the genus Cristaria Schumacher, 1817 (Bivalvia: Unionidae) are widespread from Mongolia to Indochina while the range of one species, C. plicata (Leach, 1814), covers two biogeographic subregions, i.e., East Asian (Amur River to Vietnam) and Sundaland (Mekong River basin). We present here a taxonomic revision of the nominal taxon Anodonta bellua Morelet, 1866 which was described from the Mekong (Lake Tonle-Sap, Cambodia) but is currently considered a synonym of C. plicata. We obtained molecular data for newly collected Cristaria representatives from the Mekong’s tributaries in Laos, which were found as a divergent species-level phylogenetic clade within the genus that is distant from C. plicata. Nevertheless, comparative morphological and morphometric studies did not reveal any significant differences between these two congeners. Our time-calibrated biogeographic modeling reveals that the split between Cristaria bellua (Mekong) and C. clessini (East Asia) probably occurred in the mid-Miocene (15.8 Ma) and may reflect an ancient stream capture between the Mekong Basin and East Asian rivers.

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DOI: 10.3897/zse.98.90032

Authors: Piyatida Pimvichai, Henrik Enghoff, Somsak Panha, Thierry Backeljau

Abstract: A new genus of the millipede family Pseudospirobolellidae, Siliquobolellus gen. nov., is described from limestone mountains in Thailand, based on three new species, viz. Siliquobolellus amicusdraconis gen. et sp. nov. from Uthaithani Province, Siliquobolellus constrictus gen. et sp. nov. from Prachuap Khiri Khan Province and Siliquobolellus prasankokae gen. et sp. nov. (type species) from Lampang Province. The descriptions are based on gonopod morphology and mitochondrial DNA data (COI barcodes). The COI barcodes grouped the three new species in a well-supported Siliquobolellus gen. nov. clade. The mean interspecific COI sequence divergence among the three new species was 12% (range: 8–15%). The mean intergeneric COI sequence divergence between Siliquobolellus gen. nov., Coxobolellus Pimvichai, Enghoff, Panha & Backeljau, 2020, and Pseudospirobolellus Carl, 1912 was 19% (range: 14–23%). Three conspicuous gonopodal synapomorphies differentiate Siliquobolellus gen. nov. from other pseudospirobollellid genera: (1) the telopodital part of the posterior gonopod forms a deep concavity, (2) the telopodite of the anterior gonopod is directed distad and does not reach the tip of the coxal part of the anterior gonopod, and (3) the tip of the anterior gonopod coxa is narrowed, curving mesad. As such, the monophyly of the new genus is well supported by both morphological and mitochondrial DNA data. A distribution map and an identification key to the species are provided.

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DOI: 10.3897/zse.98.79213

Authors: Daniel Jablonski, Rafaqat Masroor, Sylvia Hofmann

Abstract: The common Asian toad Duttaphrynus melanostictus (Schneider, 1799) complex has a wide distribution ranging from western foothills of the Himalaya to the easternmost range of the Wallacea, with the evidence of human-mediated introductions to some other areas. In the entire distribution range, the complex is formed by several evolutionary clades, distributed mostly in South-East Asia with unresolved taxonomy. In the northwestern edge of its distribution (Pakistan), the name D. melanostictus hazarensis (Khan, 2001) has been assigned to local populations but its biological basis remained, so far, understudied and unvalidated. Therefore, we re-evaluated the available genetic data (mitochondrial and nuclear) to show the relationships between Pakistani populations (including the type locality of D. m. hazarensis) and others from across the range. Our results showed that Pakistani populations are associated with one, deeply diverged, well-supported and widely distributed clade (so-called Duttaphrynus sp. 1 according to 16S, or clade B based on tRNAGly-ND3), that has already been detected in previous studies. This clade is further distributed in India, Nepal, Bangladesh, Malaysia, Singapore, and Indonesia and is characterized by a low level of genetic variability. This further suggests that both natural, as well as potential human-mediated dispersal, might have played an important role in setting up the current phylogeographic and distribution pattern of this clade. The clade is deeply divergent from other clades of the complex and represents a taxonomically unresolved entity. We here argue that the clade Duttaphrynus sp. 1/B represents a distinct species for which the name Duttaphrynus bengalensis (Daudin, 1802) comb. nov. is applicable, while the description of D. m. hazarensis does not satisfy the rules of the International Code of Zoological Nomenclature.

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DOI: 10.3897/zse.98.84963

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

Abstract: The taxonomic status of the strabomantid frog species Pristimantis nebulosus (Henle, 1992), originating from the southern Cordillera Azul in central Peru, is investigated based on examination of the holotype and its comparison with freshly collected topotypic material. Following current standards, we provide a redescription of the holotype. It is in a rather poor state and exhibits certain damages and preservation artifacts, conditions that have hampered an allocation of this nominal taxon to any known living population of Pristimantis in the past. Our detailed specimen-to-specimen comparison provided morphological evidence for the conspecifity of the holotype and freshly collected topotypes. Molecular phylogenetic analysis, based on the mitochondrial 16S gene fragment places P. nebulosus in the P. conspicillatus species group, being closely related to P. bipunctatus and an undescribed candidate species from the Cordillera de Carpish. From both, P. nebulosus differs by considerable divergence in the 16S gene (p-distance 4.1–6.2%). Based on the specimens available, we provide an updated diagnosis for P. nebulosus, compare it to other species in the P. conspicillatus group and describe its advertisement call. In addition, we describe and name the closely related candidate species from the Cordillera de Carpish. It is sister to P. bipunctatus and P. nebulosus, but differs from both mainly by its tuberculate dorsal skin (versus shagreen) and divergence in the 16S gene (3.3–4.1%). We briefly discuss cryptic species diversity in the P. conspicillatus and P. danae species groups and provide justification for the description of a singleton species.

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DOI: 10.3897/zse.98.83109

Authors: Wilson J. E. M. Costa, José Leonardo O. Mattos, Wagner M. S. Sampaio, Patrícia Giongo, Frederico B. de Almeida, Axel M. Katz

Abstract: The Brazilian Cerrado highlands shelter the headwaters of the three largest South American hydrographic basins, where a great species diversity is concentrated, but some biological groups are still insufficiently known. The focal taxa of this study are trichomycterid catfishes of the subgenus Cryptocambeva, genus Trichomycterus, endemic to mountain areas of south-eastern Brazil. The primary objective of this study is to test through a molecular phylogeny if a new species collected in streams of the upper Rio Paraná basin draining the Serra da Canastra is sister to T. macrotrichopterus, endemic to the upper Rio São Francisco at another facet of the Serra da Canastra, as suggested by morphological data. The analysis corroborated sister group relationships between these two species, besides supporting four main clades in Cryptocambeva, each of them endemic to distinct mountain regions. A time-calibrated analysis supported the divergence timing between the new species and T. macrotrichopterus at the Pliocene, which is chronologically compatible with the final period of intense fluvial configuration re-arrangement, when São Francisco headwater streams were captured by the Paraná basin. The new species herein described is similar to T. macrotrichopterus and distinguished from all other species of Cryptocambeva by having a long pectoral-fin filament. These two species are distinguished from each other by characteristics of the latero-sensory system, colour pattern and bone morphology.

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DOI: 10.3897/zse.97.61351

Authors: Jackie L. Childers, Sebastian Kirchhof, Aaron M. Bauer

Abstract: The lacertid genus Pedioplanis is a moderately speciose group of small-bodied, cryptically-colored lizards found in arid habitats throughout southern Africa. Previous phylogenetic work on Pedioplanis has determined its placement within the broader context of the Lacertidae, but interspecific relations within the genus remain unsettled, particularly within the P. undata species complex, a group largely endemic to Namibia. We greatly expanded taxon sampling for members of the P. undata complex and other Pedioplanis, and generated molecular sequence data from 1,937 bp of mtDNA (ND2 and cyt b) and 2,015 bp of nDNA (KIF24, PRLR, RAG-1) which were combined with sequences from GenBank resulting in a final dataset of 455 individuals. Both maximum likelihood and Bayesian analyses recover similar phylogenetic results and reveal the polyphyly of P. undata and P. inornata as presently construed. We confirm that P. husabensis is sister to the group comprising the P. undata complex plus the Angolan sister species P. huntleyi + P. haackei and demonstrate that P. benguelensis lies outside of this clade in its entirety. The complex itself comprises six species including P. undata, P. inornata, P. rubens, P. gaerdesi and two previously undescribed entities. Based on divergence date estimates, the P. undata species complex began diversifying in the late Miocene (5.3 ± 1.6 MYA) with the most recent cladogenetic events dating to the Pliocene (2.6 ± 1.0 MYA), making this assemblage relatively young compared to the genus Pedioplanis as a whole, the origin of which dates back to the mid-Miocene (13.5 ± 1.8 MYA). Using an integrative approach, we here describe Pedioplanis branchi sp. nov. and Pedioplanis mayeri sp. nov. representing northern populations previously assigned to P. inornata and P. undata, respectively. These entities were first flagged as possible new species by Berger-Dell’mour and Mayer over thirty years ago but were never formally described. The new species are supported chiefly by differences in coloration and by unique amino acid substitutions. We provide comprehensive maps depicting historical records based on museum specimens plus new records from this study for all members of the P. undata complex and P. husabensis. We suggest that climatic oscillations of the Upper Miocene and Pliocene-Pleistocene era in concert with the formation of biogeographic barriers have led to population isolation, gene flow restrictions and ultimately cladogenesis in the P. undata complex.

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DOI: 10.3897/zse.97.60099

Authors: Suranjan Karunarathna, Anslem De Silva, Dinesh Gabadage, Madhava Botejue, Majintha Madawala, Kanishka D.B. Ukuwela

Abstract: A new day gecko of the genus Cnemaspis Strauch, 1887 is described from the intermediate bioclimatic zone (Haputale Forest and Idalgashinna Forest in Badulla District) of Sri Lanka. The new species belongs to the Cnemaspis kandiana clade and was recorded from granite caves and abandoned buildings within forested areas. The region in which these habitats are located, receives relatively high annual rainfall (2500–3500 mm) and has fairly cool, moist and well-shaded conditions. The new species is medium in size (30.2–32.9 mm SVL) and can be differentiated from all other Sri Lankan Cnemaspis by the presence of small subcaudals, heterogenous dorsal scales, smooth pectoral and ventral scales, 7 or 8 supralabials and infralabials, 143–159 ventral scales, 15–17 belly scales, 95–103 mid-body scales, 122–132 paravertebrals, 3 pre-anal pores, 4 or 5 femoral pores and 17 or 18 lamellae on 4th toe. The species described herein is categorised as Critically Endangered (CR) under the IUCN Red List Criteria. The major threats for the new species are habitat loss due to expansion of commercial-scale agriculture and illicit forest encroachments. Therefore, we recommend relevant authorities to take immediate conservation action to ensure the protection of these forest areas in Haputale and Idalgashinna along with the buffer zone in the near future.

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DOI: 10.3897/zse.96.52049

Authors: Witold Morek, Bartłomiej Surmacz, Łukasz Michalczyk

Abstract: Tardigrada are a phylum of microscopic animals inhabiting a variety of ecosystems, both aquatic and terrestrial, being recognised for their remarkable abilities to withstand tough environmental conditions. The order Apochela groups exclusively carnivorous species, with the vast majority representing the genus Milnesium Doyère, 1840. Representatives of this genus are characterised by simplified morphology, therefore possessing an extremely limited set of taxonomically meaningful morphological traits. Nevertheless, the taxonomy of Milnesium is mostly based on classical data: observations and measurements in light microscopy with the majority of descriptions lacking integrative data, most importantly DNA barcodes, but also scanning electron microscopy photographs and developmental variability analysis. Hence, re-descriptions that include novel integrative data are urgently needed. In this contribution, we provide new taxonomic data for two species described from Central Asia, Milnesium almatyense (a single population) and Milnesium reductum Tumanov, 2006 (five populations): morphometrics, DNA barcodes, SEM observations and description of developmental variability. As a result, we amend the description of both species and reveal phylogenetic relationships of those species and other sequenced congeners. The integrative data confirm the validity of the two species and include them in the growing set of Milnesium species associated with DNA sequences.

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DOI: 10.3897/zse.96.51350

Authors: Sébastien Lavoué, Siti Zafirah Ghazali, Jamsari Amirul Firdaus Jamaluddin, Siti Azizah Mohd Nor, Khaironizam Md. Zain

Abstract: The fish genus Notopterus Lacepède, 1800 (Notopteridae) currently includes only one species, the Asian bronze featherback Notopterus notopterus (Pallas, 1769). This common freshwater species is widely distributed in the Oriental region, from the Indus basin in the west, the Mekong basin in the east and Java Island in the south. To examine the phylogeographic structure of N. notopterus across its range, we analysed 74 publicly available cytochrome oxidase I (COI) sequences, 72 of them determined from known-origin specimens, along with four newly-determined sequences from Peninsular Malaysian specimens. We found that N. notopterus is a complex of two allopatric species that diverge from each other by 7.5% mean p-distance. The first species is endemic to South Asia (from Indus basin to Ganga-Brahmaputra system), whereas the distribution of the second species is restricted to Southeast Asia. The exact limit between the distributions of these two species is not known, but it should fall somewhere between the Ganga-Brahmaputra and Salween basins, a region already identified as a major faunal boundary in the Oriental region. The name N. notopterus is retained for the Southeast Asian species, while the name Notopterus synurus (Bloch & Schneider, 1801) should be applied to the South Asian species. A comparative morphological study is needed to reveal the degree of morphological differentiation between the two species.

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DOI: 10.3897/zse.96.53660

Authors: Rodrigo B. Salvador, Fred J. Brook, Lara D. Shepherd, Martyn Kennedy

Abstract: A phylogenetic analysis using a combination of mitochondrial (COI, 16S) and nuclear markers (ITS2, 28S) indicated that Punctoidea, as previously interpreted, is polyphyletic. It comprises two main groups, containing northern hemisphere (Laurasian) and predominantly southern hemisphere (Gondwanan) taxa respectively, treated here as separate superfamilies. Within Punctoidea sensu stricto, Punctidae, Cystopeltidae and Endodontidae form separate monophyletic clades, but Charopidae, as currently interpreted, is paraphyletic. Most of the charopid taxa that we sequenced, including Charopa coma (Gray, 1843) and other Charopinae, grouped in a clade with Punctidae but some charopid taxa from Australia and South America grouped with Cystopeltidae. Cystopeltidae previously contained a single Australia-endemic genus, Cystopelta Tate, 1881, but our analysis suggests that it is considerably more diverse taxonomically and has a much wider distribution. For taxonomic stability, we suggest that Charopidae be retained as a family-level group for now, pending further study of the systematic relationships of its constituent taxa. A new superfamily, Discoidea, is erected here for two Northern Hemisphere families, Discidae and Oreohelicidae, which were previously assigned to Punctoidea. The North American species Radiodomus abietum, previously in Charopidae, is also here assigned to Discoidea. The phylogenetic relationships of Helicodiscidae, previously assigned to Punctoidea, were not fully resolved in our analysis, but the family is apparently closely related to Arionoidea Gray, 1840 and infraorder Limacoidei.

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DOI: 10.3897/zse.96.50821

Authors: Piotr Gąsiorek, Katarzyna Vončina, Peter Degma, Łukasz Michalczyk

Abstract: The phyletic relationships both between and within many of tardigrade genera have been barely studied and they remain obscure. Amongst them is the cosmopolitan Bryodelphax, one of the smallest in terms of body size echiniscid genera. The analysis of newly-found populations and species from the Mediterranean region and from South-East Asia gave us an opportunity to present the first phylogeny of this genus, which showed that phenotypic traits used in classical Bryodelphax taxonomy do not correlate with their phyletic relationships. In contrast, geographic distribution of the analysed species suggests their limited dispersal abilities and seems to be a reliable predictor of phylogenetic affinities within the genus. Moreover, we describe three new species of the genus. Bryodelphax australasiaticus sp. nov., by having the ventral plate configuration VII:4-4-2-4-2-2-1, is a new member of the weglarskae group with a wide geographic range extending from the Malay Peninsula through the Malay Archipelago to Australia. Bryodelphax decoratus sp. nov. from Central Sulawesi (Celebes) also belongs to the weglarskae group (poorly visible ventral plates VII:4-2-2-4-2-2-1) and is closely related to the recently described Bryodelphax arenosus Gąsiorek, 2018, but is differentiated from the latter by well-developed epicuticular granules on the dorsum. Finally, a new dioecious species, Bryodelphax nigripunctatus sp. nov., is described from Mallorca and, by the reduced ventral armature (II/III:2-2-(1)), it resembles Bryodelphax maculatus Gąsiorek et al., 2017. The latter species, known so far only from northern Africa, is recorded from Europe for the first time. A taxonomic key to the genus members is also presented.

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DOI: 10.3897/zse.96.50360

Authors: Jessica Colavite, Amanda M. Windsor, William Santana

Abstract: A new genus of majoid spider crab, Pohleus gen. nov. is established for Pericera septemspinosa Stimpson, 1871 and Pericera heptacantha Bell, 1836, based on morphology and molecular data from the partial sequences of the 12S and 16S mitochondrial genes and the 18S small subunit rRNA nuclear locus. The species are re-described and illustrated, based on material from several localities of the western Atlantic and eastern Pacific oceans. The carapace, antennal and pterygostomial spines, male thoracic sternum and first gonopods are distinctive characters, distinguishing Pohleus gen. nov. from species assigned to Macrocoeloma Miers, 1879, where P. septemspinosus and P. heptacanthus are currently included.

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DOI: 10.3897/zse.96.49946

Authors: Marc Domènech, Luís C. Crespo, Alba Enguídanos, Miquel A. Arnedo

Abstract: The incorporation of molecular data into current taxonomic practise has unravelled instances of incongruence among different data sets. Here we report a case of mitochondrial discordance in cobweb spiders of the genus Theridion Walckenaer, 1805 from the Iberian Peninsula. Morphological examination of samples from a country-wide bioinventory initiative revealed the existence of a putative new species and two nominal species belonging to the Theridion melanurum species group. The morphological delineation was supported by the molecular analysis of a nuclear marker but was at odds with the groups circumscribed by a mitochondrial marker. The causes of this discordance remained uncertain, once sample and sequencing errors and the existence of pseudogenes were discarded. The full sorting observed in the alleles of the more slowly evolving nuclear marker ruled out incomplete lineage sorting, while the geographic patterns recovered were difficult to reconciliate with ongoing hybridization. We propose that the apparent incongruence observed is most likely the result of old introgression events in a group with high dispersal abilities. We further speculate that endosymbiont-driven cytoplasmatic incompatibility could be involved in the fixation of mitochondrial haplotypes across species barriers. Additionally, we describe the new species T. promiscuum sp. nov., based on the presence of diagnostic morphological traits, backed up by the nuclear data delimitation. Our study contributes yet another example of the perils of relying on single methods or data sources to summarise the variation generated by multiple processes acting through thousands of years of evolution and supports the key role of biological inventories in improving our knowledge of invertebrate biodiversity.

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DOI: 10.3897/zse.96.48312

Authors: Arthur F. Sands, Peter Glöer, Mustafa E. Gürlek, Christian Albrecht, Thomas A. Neubauer

Abstract: Asia contains a high species diversity of the freshwater gastropod genus Theodoxus. Recent molecular and morphological reviews of this diversity have uncovered a number of yet undescribed species while suggesting the urgent revision of several others. Moreover, some of these studies have indicated a number of species previously not recorded for this continent. Despite the advancements, a taxonomic revision and an update on the distribution of Theodoxus spp. in Asia is still pending. Here, we construct the most robust phylogeny of Theodoxus up to now and review original descriptions, type material, recent taxonomic revisions, compendia, and species lists to provide a comprehensive checklist of all known extant Asian Theodoxus spp. Our checklist also provides descriptions for three recently discovered and yet undescribed species (Theodoxus gurur Sands & Glöer, sp. nov., Theodoxus wesselinghi Sands & Glöer, sp. nov., and Theodoxus wilkei Sands & Glöer, sp. nov.), as well as shows the need to synonymise several previously described morphospecies. The present revision recognizes 14 extant Theodoxus spp. for Asia. Some of these species are widespread, while others are endemic to just a single location. Based on the revised and new distribution data, we provide updates and new assessments of species conservation statuses.

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DOI: 10.3897/zse.95.36788

Authors: Pâmella Silva de Brito, Erick Cristofore Guimarães, Luis Fernando Carvalho-Costa, Felipe Polivanov Ottoni

Abstract: A new species of Aphyocharax is described from the Maracaçumé river basin, eastern Amazon, based on morphological and molecular data. The new species differs from all its congeners, mainly by possessing the upper caudal-fin lobe longer than the lower one in mature males, and other characters related to teeth counts, colour pattern, and body depth at dorsal-fin origin. In addition, the new species is corroborated by a haplotype phylogenetic analyses based on the Cytochrome B (Cytb) mitochondrial gene, where its haplotypes are grouped into an exclusive lineage, supported by maximum posterior probability value, a species delimitation method termed the Wiens and Penkrot analysis (WP).

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DOI: 10.3897/zse.94.28981

Authors: Nuanpan Veeravechsukij, Duangduen Krailas, Suluck Namchote, Benedikt Wiggering, Marco T. Neiber, Matthias Glaubrecht

Abstract: The freshwater thiarid gastropod Tarebia granifera (Lamarck, 1816), including taxa considered either congeneric or conspecific by earlier authors, is widespread and abundant in various lentic and lotic water bodies in mainland and insular Southeast Asia, with its range extending onto islands in the Indo-West-Pacific. This snail is, as one of the most frequent and major first intermediate host, an important vector for digenic trematodes causing several human diseases. As a typical thiarid T. granifera is viviparous and parthenogenetic, with various embryonic stages up to larger shelled juveniles developing within the female’s subhemocoelic (i.e non-uterine) brood pouch. Despite the known conchological disparity in other thiarids as well as this taxon, in Thailand Tarebia has been reported with the occurrence of one species only. In light of the polytypic variations found in shell morphology of freshwater snails in general and this taxon in particular, the lack of a modern taxonomic-systematic revision, using molecular genetics, has hampered more detailed insights to date, for example, into the locally varying trematode infection rates found in populations of Tarebia from across its range in Thailand as well as neighboring countries and areas. Here, we integrate evidence from phylogeographical analyses based on phenotypic variation (shell morphology, using biometry and geometric morphometrics) with highly informative and heterogeneous mtDNA sequence data (from the gene fragments cytochrome c oxidase subunit 1 and 16 S rRNA). We evaluate both the morphological and molecular genetic variation (using several phylogenetic analyses, including haplotype networks and a dated molecular tree), in correlation with differences in the reproductive biology among populations of Tarebia from various water bodies in the north, northwest, central, and south of Thailand, supplementing our respective analyses of parasite infections of this thiarid by cercaria of 15 trematode species, reported in a parallel study. Based on the comparison of topotypical material from the island of Timor, with specimens from 12 locations as reference, we found significant, albeit not congruent variation of both phenotype and genotype in Tarebia granifera, based on 1,154 specimens from 95 Thai samples, representing a geographically wide-ranging, river-based cross-section of this country. Our analyses indicate the existence of two genetically distinct clades and hint at possible species differentiation within what has been traditionally considered as T. granifera. These two lineages started to split about 5 mya, possibly related to marine transgressions forming what became known as biogeographical barrier north of the Isthmus of Kra. Grounded on the site-by-site analysis of individual Tarebia populations, our country-wide chorological approach focussing on the conchologically distinct and genetically diverse lineages of Tarebia allows to discuss questions of this either reflecting subspecific forms versus being distinct species within a narrowly delimited species complex. Our results, therefore, provide the ground for new perspectives on the phylogeography, evolution and parasitology of Thai freshwater gastropods, exemplified here by these highly important thiarids.

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DOI: 10.3897/zse.94.28085

Authors: Wilson J. E. M. Costa, Pedro F. Amorim

Abstract: The miniature seasonal killifish Spectrolebias costae, first described for the middle Araguaia River basin, has been also recorded from two areas in the middle Tocantins River basin, from where male specimens exhibit some differences in their colour pattern. Analyses directed to species delineation (GMYC and bPTP), using a fragment of the mitochondrial gene COI, strongly support two species, S. costae from the Araguaia River basin and a new species from the Tocantins River basin. Spectrolebias gracilis sp. n. is described on the basis of specimens collected from two localities separated by about 530 km, Canabrava River floodplains near Alvorada do Tocantins and Tocantins River floodplains near Palmeirante. Field inventories were unsuccessful in finding additional populations in the region, which is attributed to the high environmental degradation, including several large dams that have permanently inundated typical killifish habitats. Spectrolebias gracilis is member of a clade also including S. costae, S. inaequipinnatus, and S. semiocellatus, diagnosed by having the dorsal and anal fins in males with iridescent dots restricted to their basal portion, caudal fin in males hyaline, and caudal-fin base with two pairs of neuromasts. Within this clade, a single miniaturisation event is supported for the most recent common ancestor of the subclade comprising S. costae and S. gracilis, which differ from other congeners by reaching only about 20 mm standard length as maximum adult size.

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DOI: 10.3897/zse.94.14698

Authors: Adriana Bellati, Mark D. Scherz, Steven Megson, Sam Hyde Roberts, Franco Andreone, Gonçalo M. Rosa, Jean Noël, Jasmin E. Randrianirina, Mauro Fasola, Frank Glaw, Angelica Crottini

Abstract: The systematics of the cophyline microhylid frog genera Plethodontohyla and Rhombophryne have long been intertwined, and their relationships have only recently started to become clear. While Rhombophryne has received a lot of recent taxonomic attention, Plethodontohyla has been largely neglected. Our study is a showcase of just how complex the taxonomic situation between these two genera is, and the care that must be taken to resolve taxonomic conundrums where old material, multiple genus transitions, and misattribution of new material obfuscate the picture. We assessed the identity of the historic names Dyscophus alluaudi (currently in the genus Rhombophryne), Phrynocara laeve and Plethodontohyla laevis tsianovohensis (both synonyms of Rhombophryne alluaudi) based on an integrative taxonomic approach harnessing genetics, external morphology, osteological data obtained via micro-Computed Tomography (micro-CT) and bioacoustics. We show that (1) the holotype of Dyscophus alluaudi is a member of the genus Plethodontohyla; (2) the Rhombophryne specimens from central Madagascar currently assigned to Rhombophryne alluaudi have no affinity with that species, and are instead an undescribed species; and (3) Phrynocara laeve and Dyscophus alluaudi are not synonymous, but represent closely related species, whereas Plethodontohyla laevis tsianovohensis is tentatively confirmed as synonym of D. alluaudi. We resurrect and re-describe Plethodontohyla laevis, and re-allocate and re-describe Plethodontohyla alluaudi on the basis of new and historic material.

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DOI: 10.3897/zse.93.14906

Authors: Mark D. Scherz, Miguel Vences, James Borrell, Lawrence Ball, Denise Herizo Nomenjanahary, Duncan Parker, Marius Rakotondratsima, Elidiot Razafimandimby, Thomas Starnes, Jeanneney Rabearivony, Frank Glaw

Abstract: A recent study on a group of rough-skinned Gephyromantis frogs from Madagascar (Anura: Mantellidae: Mantellinae) established a new subgenus, Asperomantis, with five described species and one undescribed candidate species. Based on newly collected material from the Bealanana District, we address the taxonomy of this candidate species, and reveal that it consists of two populations with low genetic and morphological divergence but considerable bioacoustic differences that are obvious to the human ear. As a result, we describe some of the specimens formerly assigned to Gephyromantis sp. Ca28 as G. angano sp. n. and assign the remaining specimens from a locality between Bealanana and Antsohihy to a new Unconfirmed Candidate Species, G. sp. Ca29. Gephyromantis angano sp. n. is a small species that strongly resembles G. asper and G. ceratophrys, but it differs from these and all other Gephyromantis species by a unique, clinking advertisement call. The new species may be highly threatened by habitat fragmentation, but at present we recommend it be treated as Data Deficient until more data are available to assess its distribution. We discuss the curious relationship between G. angano sp. n. and G. sp. Ca29, which we suspect may represent a case of incipient speciation. We also identify two additional new Unconfirmed Candidate Species of Gephyromantis based on sequence data from other specimens collected during our surveys in the Bealanana District.

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DOI: 10.3897/zse.91.4979

Authors: Mark D. Scherz, Andolalao Rakotoarison, Oliver Hawlitschek, Miguel Vences, Frank Glaw

Abstract: The Madagascar-endemic microhylid genus Rhombophryne consists of a range of partly or completely fossorial frog species. They lead a poorly known, secretive lifestyle, and may be more diverse than previously thought. We describe a new species from the high altitude forests of the Sorata massif in north Madagascar with unusual characteristics for this genus; R. longicrus sp. n. has long, slender legs, unlike most of its fossorial or semi-fossorial congeners. The new species is closely related to R. minuta, a much smaller frog from the Marojejy massif to the southeast of Sorata with similarly long legs. We discuss the morphology of these species relative to the rest of the genus, and argue that it suggests adaptation away from burrowing and toward a more saltatorial locomotion and an accordingly more terrestrial lifestyle. If this is the case, then these frogs represent yet more ecological diversity within the already diverse Cophylinae. We recommend an IUCN Red List status of Endangered B1ab(iii) for R. longicrus sp. n., because it is known only from a single site in a forested area of roughly 250 km2, which is not yet incorporated into any protected area.

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