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  • Mao Lin; Zhenzu Xu; Jiaqi Huang; Donghui Guo; Chunguang Wang
    Zoological Systematics.
    A generic diagnosis of family Eucodoniidae is revised, with the monotype genus Eucodonium. Three new species, Eucodonium bitentaculatum Xu, Huang & Guo, sp. nov., E. brevistyle Xu, Huang & Lin, sp. nov. and E. longitentaculatum Xu, Huang & Wang, sp. nov. from South China Sea are described and illustrated. A key to all species of Eucodonium is provided. The type specimens are deposited in the Third Institute of Oceanography, State Oceanic Administration.
  • Communication
    Shuqiang Li, Yejie Lin
    Zoological Systematics. 2024, 49(1): 1-3.
  • Original Article
    Yejie Lin, Shuqiang Li, Haolin Mo, Xihao Wang
    Zoological Systematics. 2024, 49(1): 4-98.
  • Original Article
    Zhihong Zhan, , Daniel K. Young
    Zoological Systematics. 2023, 48(4): 279-360.
    This study focused on and examined the traditional Lucanus fortunei group comprising more than 60% of all Lucanus taxa in China. Each species treatment includes a description, distribution, collecting methods, natural history and digital habitus and other pertinent images. A dichotomous taxonomic key was created specific to the Chinese L. fortunei group and the related groups. The Lucanus fortunei group is arguably the largest group in terms of species richness in Lucanus Scopoli; however, the group definition is unclear and needs to be critically assessed and redefined. Field work and sampling of the Chinese Lucanus fortunei species group was conducted between summer, 2017 and summer, 2022. Field sites included Anhui, Zhejiang, Fujian, Taiwan, Jiangxi, Hubei, Hunan, Guizhou, Sichuan, Chongqing, Yunnan, Xizang, Guangdong, and Guangxi Provinces. Thirty-two taxa (species and subspecies), including 473 specimens, previously considered to fall within the scope of the Lucanus fortunei group were examined by morphological study. Three new groups, the Lucanus brivioi group, the Lucanus kraatzi group, and the Lucanus parryi group are proposed and separated from the traditional Lucanus fortunei group, based on morphological characters. The subspecies, L. parryi laetus, is proposed as the synonym of the nominate subspecies.
  • Letter
    Guangchun Liu
    Zoological Systematics. 2023, 48(4): 361-364.
  • Communication
    Shiyu Du1, Yinhuan Ding2, Hu Li3, Aibing Zhang4, Arong Luo5, Chaodong Zhu4, 5, Feng Zhang1, 5
    Zoological Systematics. 2023, 48(2): 107-116.
    Phylogenomics is a new field that infers evolutionary relationships of taxa at the genome-scale level. The increment of molecular data may raise the potential bias as the limiting factor in phylogenomics. It is particularly important to explore these factors in phylogenomic analyses by simple, convenient, time-saving and (relatively) robust means. Here, we construct a set of custom scripts for USCO (universal single-copy orthologs) loci extraction, multiple sequence alignment, trimming poorly aligned regions, loci filtering and creating a concatenation matrix, prior to reconstructing the phylogenetic trees, to simplify analytical pipelines and improve the accuracy of tree estimation. These scripts employed a series of computationally efficient bioinformatic tools, and were used with a universal ‘BASH’ shell or visual interface by Windows-like ‘drag and drop’ operations in LINUX systems. Most steps in these scripts are parallelized to accelerate analyses. These new custom scripts provide a convenient analytical solution for phylogenomics data preparation, data quality control, and detection of potential analytical errors. Details and scripts usage are provided at Phylogenomics/tree/main/scripts. The virtual mirror file (.vmdk) integrates the operating system and required environment. All tools and scripts can be downloaded from 10.6084/m9.figshare.21283026. Besides, the video introduction and “walk-through” for each script are provided at
  • Communication
    Hecai Zhang, Junhua He, Changying Shi, Guangwen Chen, Dezeng Liu
    Zoological Systematics. 2023, 48(2): 117-125.
    Planarian is an excellent model for studying several scientific issues, and planarianology has become an active research field in the world. Researchers at home and abroad have conducted a lot of in-depth studies on planarians in various areas. This paper outlined the panorama of planarian research in China and summarized the advances from three aspects of taxonomy, molecular phylogeny and phylogeography, regeneration, tissue homeostasis and immunity as well as toxicology. Furthermore, the current problems and future development direction and tendency of planarian research in China are briefly analyzed and prospected. Importantly, the catalogue of planarians in China is also presented.
  • Original Article
    Xiaolei Huang1, 2, Samiran Chakrabarti3, Junjie Li1, Bradford A. Hawkins4, Gexia Qiao2
    Zoological Systematics. 2023, 48(2): 126-139.
    The history of the Qinghai-Tibetan Plateau and the Himalayas (QTPH) region makes it a ‘living laboratory’ for understanding how the Earth’s abiotic evolution has shaped regional biotas. However, no studies to date have attempted to investigate general macro-scale biogeographical patterns and biogeographical affinities of the QTPH terrestrial fauna in a global context. Based on the analyses of the global distributions of QTPH Aphidomorpha species, we examined generalized distribution patterns for them and investigated the biogeographical affinities between the QTPH and other regions. Track analysis and cluster analysis of the global distributions of 298 Aphidomorpha species in the QTPH identified seven generalized biogeographical tracks and eight general distribution patterns. Both approaches detected similarly generalized distribution patterns. Four marginal areas where the generalized tracks intersect, i.e. the eastern Himalayas, the western Himalayas, the Hengduan Mountains, and the northeastern QTPH, were identified as biogeographical nodes. The generalized distribution patterns indicate the biogeographical affinities between the QTPH and other regions and suggest the complex nature of the QTPH fauna. A spatial congruence between the nodes and diversity centres detected by previous studies indicates the four marginal areas have been key areas in fauna evolution as centres for faunal exchange as well as glacial refugia. Based on the biogeographical patterns, the geological history of the QTPH and the evolutionary history of Aphidomorpha, we propose an evolutionary scenario for the evolution of the QTPH aphid fauna in which faunal exchanges, vicariance due to the uplift of geographical barriers, environmental heterogeneity and Quarternary glaciations have shaped the diversity patterns and fauna in the QTPH.
  • Original Article
    Yejie Lin, Shuqiang Li, Dinh-Sac Pham
    Zoological Systematics. 2023, 48(1): 1-99.
    Three new genera of spiders are described: Jishiyu Lin & Li, gen. nov., of Agelenidae C. L. Koch, 1837, with J. songjiang Lin & Li, sp. nov. (♂, China: Yunnan) as the type species, Marusik Lin & Li, gen. nov., of Gnaphosidae Banks, 1892, with M. yurii Lin & Li, sp. nov. (♂, Vietnam) as the type species and Xiaoliguang Lin & Li, gen. nov., of Phrurolithidae Banks, 1892, with X. huarong Lin & Li, sp. nov. (♂♀, Vietnam) as the type species. Thirty-five new species are described in 12 families: Corinnidae Karsch, 1880: Spinirta qishuoi Lin & Li, sp. nov. (♀, China: Guangxi); Hahniidae Bertkau, 1878: Cicurina miniae Lin & Li, sp. nov. (♀, Vietnam); Halonoproctidae Pocock, 1901: Latouchia wenruni Lin & Li, sp. nov. (♂♀, China: Hainan); Liocranidae Simon, 1897: Oedignatha wuyong Lin & Li, sp. nov. (♂♀, Vietnam) and Sphingius gongsunsheng Lin & Li, sp. nov. (♀, Vietnam); Nemesiidae Simon, 1889: Sinopesa guansheng Lin & Li, sp. nov. (♂, China: Yunnan); Phrurolithidae Banks, 1892: Edelithus huyanzhuo Lin & Li, sp. nov. (♂♀, Vietnam), E. linchong Lin & Li, sp. nov. (♂♀, Vietnam), E. qinming Lin & Li, sp. nov. (♂♀, Vietnam), Grandilithus linglingae Lin & Li, sp. nov. (♂♀, Vietnam), G. yangzhi Lin & Li, sp. nov. (♀, Vietnam), Lingulatus chaijin Lin & Li, sp. nov. (♀, Vietnam), L. dongping Lin & Li, sp. nov. (♂♀, Vietnam), L. liying Lin & Li, sp. nov. (♂♀, Vietnam), L. longfeiae Lin & Li, sp. nov. (♂♀, Vietnam), L. luzhishen Lin & Li, sp. nov. (♂♀, Vietnam), L. zhangqing Lin & Li, sp. nov. (♂♀, Vietnam), L. zhutong Lin & Li, sp. nov. (♂♀, Vietnam), Otacilia liuxinyei Lin & Li, sp. nov. (♂♀, Vietnam), O. wusong Lin & Li, sp. nov. (♂, Vietnam) and O. xuning Lin & Li, sp. nov. (♀, Vietnam); Pimoidae Wunderlich, 1986: Pimoa zeluni Lin & Li, sp. nov. (♂, China: Hebei); Salticidae Blackwall, 1841: Epeus covid Lin & Li, sp. nov. (♂♀, China: Yunnan); Theridiidae Sundevall, 1833: Phoroncidia minlangi Lin & Li, sp. nov. (♀, China: Fujian); Thomisidae Sundevall, 1833: Borboropactus nanda Lin & Li, sp. nov. (♂♀, China: Yunnan), Cozyptila haocongi Lin & Li, sp. nov. (♂♀, Vietnam), Lysiteles huyilinae Lin & Li, sp. nov. (♂♀, Vietnam), Massuria daizong Lin & Li, sp. nov. (♂♀, China: Yunnan) and Zametopina wanliae Lin & Li, sp. nov. (♂, Vietnam); Trachelidae Simon, 1897: Utivarachna yumaoi Lin & Li, sp. nov. (♂, Vietnam); Zodariidae Thorell, 1881: Asceua shuangreni Lin & Li, sp. nov. (♂♀, Vietnam), Cydrela likui Lin & Li, sp. nov. (♂♀, Vietnam), Mallinella lisongi Lin & Li, sp. nov. (♂♀, Vietnam), M. pingzhoui Lin & Li, sp. nov. (♂♀, Vietnam) and M. sena Lin & Li, sp. nov. (♂♀, Vietnam). One genus name and one species name are detected as homonyms and thereby replacement names for the preoccupied names are proposed, in accordance with Article 60 of the International Code of Zoological Nomenclature (ICZN, 1999), i.e., Xilithus Liu & Li, nom. nov., ex Acrolithus Liu & Li, 2022 (preoccupied nomen); Araneus xiang Mi & Li, nom. nov., ex Araneus arcuatus Mi & Li, 2022 (preoccupied nomen). Therefore, all 16 species of Acrolithus Liu & Li, 2022 are transferred to Xilithus Liu & Li, nom. nov.
  • Editorial
    Deyuan Hong, Wenying Zhuang, Min Zhu, Keping Ma, Xiaoquan Wang, Dawei Huang, Yalin Zhang, Guodong Ren, Wenjun Bu, Wanzhi Cai, Dong Ren, Ding Yang, Aiping Liang, Fengyan Bai, Runzhi Zhang, Fumin Lei, Shuqiang Li, Hongzhi Kong, Lei Cai, Yucheng Dai, Chaodong Zhu, Qisen Yang, Jun Chen, Zhongli Sha, Jianping Jiang, Jing Che, Donghui Wu, Jiatang Li, Qiang Wang, Xinli Wei, Ming Bai, Xingyue Liu, Xuexin Chen, Gexia Qiao
    Zoological Systematics. 2022, 47(3): 185-187.
  • Communication
    Chaodong Zhu, Arong Luo, Ming Bai, Michael C. Orr, Zhonge Hou, Siqin Ge, Jun Chen, Yibo Hu, Xuming Zhou, Gexia Qiao, Hongzhi Kong, Limin Lu, Xiaohua Jin, Lei Cai, Xinli Wei, Ruilin Zhao, Wei Miao, Qingfeng Wang, Zhongli Sha, Qiang Lin, Meng Qu, Jianping Jiang, Jiatang Li, Jing Che, Xuelong Jiang, Xiaoyong Chen, Lianming Gao, Zongxin Ren, Chunlei Xiang, Shixiao Luo, Donghui Wu, Dong Liu, Yanqiong Peng, Tao Su, Chenyang Cai, Tianqi Zhu, Wanzhi Cai, Xingyue Liu, Hu Li, Huaijun Xue, Zhen Ye, Xuexin Chen, Pu Tang, Shujun Wei, Hong Pang, Qiang Xie, Feng Zhang, Feng Zhang, Xianjin Peng, Aibing Zhang, Taiping Gao, Changfa Zhou, Chen Shao, Libin Ma, Zhaoming Wei, Yunxia Luan, Ziwei Yin, Wu Dai, Cong Wei, Xiaolei Huang, Jingxian Liu, Xiangsheng Chen, Tianci Yi, Zhisheng Zhang, Zhulidezi Aishan, Qin Li, Hongying Hu
    Zoological Systematics. 2022, 47(3): 188-197.
    Taxonomy plays an important role in understanding the origin, evolution, and ecological functionality of biodiversity. There are large number of unknown species yet to be described by taxonomists, which together with their ecosystem services cannot be effectively protected prior to description. Despite this, taxonomy has been increasingly underrated insufficient funds and permanent positions to retain young talents. Further, the impact factor-driven evaluation systems in China exacerbate this downward trend, so alternative evaluation metrics are urgently necessary. When the current generation of outstanding taxonomists retires, there will be too few remaining taxonomists left to train the next generation. In light of these challenges, all co-authors worked together on this paper to analyze the current situation of taxonomy and put out a joint call for immediate actions to advance taxonomy in China.
  • Original Article
    Yejie Lin, Huifeng Zhao, Joseph K H Koh, Shuqiang Li
    Zoological Systematics. 2022, 47(3): 198-270.
    Five new genera are reported, namely Rathalos Lin & Li, gen. nov., of Anyphaenidae Simon, 1892, with Anyphaena xiushanensis Song & Zhu, 1991 (♂♀, China: Hubei, Sichuan) as the type species, Funny Lin & Li, gen. nov., of Dictynidae O. Pickard-Cambridge, 1871, with Funny valentine Lin & Li, sp. nov. (♂♀, China: Sichuan) as the type species, Pararana Lin & Li, gen. nov., of Leptonetidae Simon, 1890, with Pararana gaofani Lin & Li, sp. nov. (♂, China: Jiangsu) as the type species, and Burmalema Zhao & Li, gen. nov. and Milema Zhao & Li, gen. nov. of Telemidae Fage, 1913, with Burmalema shan Zhao & Li, sp. nov. (♂♀, Myanmar) and Milema nuichua Zhao & Li, sp. nov. (♂♀, Vietnam) as the type species. Another twenty-two new species are reported in the current paper, including Araneidae Clerck, 1757: Ordgarius bo Lin & Li, sp. nov. (♀, China: Yunnan), O. liyuanba Lin & Li, sp. nov. (♀, China: Hainan) and Paraplectana magnaculata Lin & Li, sp. nov. (♀, China: Hunan); Ctenidae Keyserling, 1877: Amauropelma gui Lin & Li, sp. nov. (♂, China: Guangxi); Filistatidae Simon, 1864: Pholcoides erebus Lin & Li, sp. nov. (♂, China: Tibet); Gnaphosidae Banks, 1892: Cryptodrassus beijing Lin & Li, sp. nov. (♂♀, China: Beijing); Halonoproctidae Pocock, 1901: Latouchia yuanjingae Lin & Li, sp. nov. (♂♀, China: Hainan); Liocranidae Simon, 1897: Paratus meiyingae Lin & Li, sp. nov. (♂, China: Yunnan) and P. nanling Lin & Li, sp. nov. (♂♀, China: Guangdong); Ochyroceratidae Fage, 1912: Speocera tongyaoi Lin & Li, sp. nov. (♂♀, China: Guangdong) and S. wangi Lin & Li, sp. nov. (♂, China: Hainan); Oxyopidae Thorell, 1869: Hamataliwa wangi Lin & Li, sp. nov. (♂♀, China: Fujian); Salticidae Blackwall, 1841: Hyllus shanhonghani Lin & Li, sp. nov. (♂♀, China: Jiangxi); Selenopidae Simon, 1897: Siamspinops banna Lin & Li, sp. nov. (♂♀, China: Yunnan); Telemidae Fage, 1913: Mekonglema chiangmai Zhao & Li, sp. nov. (♂♀, Thailand), Milema lorkor Zhao & Li, sp. nov. (♂♀, Thailand) and M. sai Zhao & Li, sp. nov. (♂♀, Thailand); Tetrablemmidae O. Pickard-Cambridge, 1873: Singaporemma shenzhen Lin & Li, sp. nov. (♂♀, China: Guangdong); Tetragnathidae Menge, 1866: Dolichognatha yue Lin & Li, sp. nov. (♂♀, China: Guangdong); Theridiidae Sundevall, 1833: Spheropistha pingshan Lin & Li, sp. nov. (♂♀, China: Guangdong); Theridiosomatidae Simon, 1881: Wendilgarda housaiyuae Lin & Li, sp. nov. (♂♀, China: Yunnan) and Titanoecidae Lehtinen, 1967: Pandava banna Lin & Li, sp. nov. (♂♀, China: Yunnan). Two new combinations in the family Anyphaenidae are proposed: Rathalos treecko (Lin & Li, 2021), comb. nov., R. xiushanensis (Song & Zhu, 1991), comb. nov., both transferred from Anyphaena Sundevall, 1833.
  • Letter
    Qiang Li, Xiaoyan Luo, Qian Liu, Zhentao Cheng, Xiaolei Huang
    Zoological Systematics. 2022, 47(3): 271-273.
  • Communication
    Xinzheng Li
    Zoological Systematics. 2022, 47(2): 95-108.
    The researches on taxonomy and phylogeny of the subphylum Crustacea in China, mainly the marine species, are reviewed in the present paper. The researches involved taxonomy, morphology, fauna and zoogeography, phylogeny and systematics. The mentioned groups almost cover all groups of the Crustacea which have been studied by Chinese scholars. This review is of great significance to summarize the achievements in the field of taxonomy and phylogeny on marine crustaceans in China.
  • Original Paper
    Yejie Lin, Shuqiang Li
    Zoological Systematics. 2022, 47(2): 132-145.
    Seven species of the spider family Hersiliidae Thorell, 1869 are included in this study. Four new species from two genera are described: Hersilia aoqin sp. nov. (♂♀), H. long sp. nov. (♀), H. sagada sp. nov. (♂♀), and Murricia caishen sp. nov. (♀). The genus Murricia, 1882 and two species, H. lelabah Rheims & Brescovit, 2004 and H. martensi Baehr & Baehr, 1993, are reported from China for the first time. H. yunnanensis Wang, Song & Qiu, 1993, syn. nov. is regarded as a junior synonym of H. sumatrana (Thorell, 1890).
  • Original Paper
    Yafei Xin, Tongyao Jiang, Zhiyuan Yao, Shuqiang Li
    Zoological Systematics. 2022, 47(1): 1-65.
    Twenty new species of extinct spiders belonging to 11 genera in five families from Late Cretaceous Kachin amber are reported. One new genus, Triangulum Jiang & Li, gen. nov., of the family Theridiidae Sundevall, 1833 was established with T. thutai Jiang & Li, sp. nov. as the type species. The remaining 19 new species are: Retrooecobius lwini Jiang & Li, sp. nov., Zamilia arkari Jiang & Li, sp. nov., Z. sheini Jiang & Li, sp. nov., Z. shwayi Jiang & Li, sp. nov. from Oecobiidae Blackwall, 1862; Furcembolus inzaliae Jiang & Li, sp. nov. from Pacullidae Simon, 1894; Priscaleclercera chimei Jiang & Li, sp. nov., P. foshou Jiang & Li, sp. nov., P. hlaingi Jiang & Li, sp. nov., P. kani Jiang & Li, sp. nov., P. kyawae Jiang & Li, sp. nov., P. thanae Jiang & Li, sp. nov., P. thaungi Jiang & Li, sp. nov. from Psilodercidae Machado, 1951; Bicornoculus aungi Jiang & Li, sp. nov., B. wunnai Jiang & Li, sp. nov., B. yarzari Jiang & Li, sp. nov., Cymbioblemma ohnmari Jiang & Li, sp. nov., Palpalpaculla phyui Jiang & Li, sp. nov. from Tetrablemmidae O. Pickard-Cambridge, 1873; Burmatheridon cetani Jiang & Li, sp. nov., Cretotheridion champoi Jiang & Li, sp. nov., from Theridiidae. All species are described based on males. All types are deposited in the Institute of Zoology, Chinese Academy of Sciences (IZCAS) in Beijing, China.
  • Original Paper
    Tao Luo, Shasha Yan, Ning Xiao, Wei Li, Huaiqing Deng, Jiang Zhou
    Zoological Systematics. 2022, 47(1): 66-88.
    A new newt species, Tylototriton daloushanensis Zhou, Xiao & Luo, sp. nov., is described from Guizhou, China, based on phylogenetic and morphological analyses. Phylogenetic analyses based on mitochondrial 16S and ND2 genes show that the new species is an independent lineage closely related to T. maolanensis, T. broadoridgus, T. dabienicus, T. anhuiensis, and T. wenxianensis. Morphologically, the new species is distinguished from its congeners by a combination of the following morphological characters: (1) large body size (SVL 64.7–83.6 mm in male and 70.5–100.3 mm in female); (2) head longer than wide, prominent bony ridges present; (3) snout rounded in dorsal view; (4) tail length less than snout-vent length in both males and females; (5) dark black body coloration, except for bright orange cranial region, palms and soles, vent region, and ventral ridge of tail; (6) relative length of toes III > IV > II > I > V; (7) fingertips reaching to between the eyes and nostrils when the forelimbs are stretched forward; (8) distal tip of the limbs greatly overlapping when the fore and hind limbs are pressed along the trunk; (9) presence of gular fold; (10) vertebral ridge slightly segmented, slightly flattened rib nodules, indistinct interspaces between rib nodules; and (11) lacking orange color markings on posterior parotoids and rib nodules. The found of the new species suggests that the species diversity of the genus Tylototriton may be underestimated.
  • Letter
    Arong Luo, Chaodong Zhu
    Zoological Systematics. 2022, 47(1): 89-92.
  • Original Paper
    Yejie Lin, Yuri M. Marusik, Caixia Gao, Hao Xu, Xiaoqing Zhang, Ziyi Wang, Wenhui Zhu, Shuqiang Li
    Zoological Systematics. 2021, 46(2): 91-152.
    A new genus, Sinophaena Lin & Li, gen. nov., of the spider family Anyphaenidae Bertkau, 1878 is reported, with S. xiweni Lin & Li, sp. nov. (♀) from Sichuan, China as the type species. Sinophaena bivalva (Zhang & Song, 2004) comb. nov. is transferred from Anyphaena Sundevall, 1833. Further new species reported in the current paper include: Anyphaena grovyle Lin & Li, sp. nov. (♂♀, Hainan), A. sceptile Lin & Li, sp. nov. (♂♀, Hainan), A. shenzhen Lin & Li, sp. nov. (♂♀, Guangdong), A. tibet Lin & Li, sp. nov. (♂♀, Tibet), A. treecko Lin & Li, sp. nov. (♂♀, Hainan), Cybaeus fushun Lin & Li, sp. nov. (♀, Liaoning), C. huadian Lin & Li, sp. nov. (♀, Jilin), Eresus lishizheni Lin, Marusik & Li, sp. nov. (♂, Xinjiang), Alistra pikachu Lin & Li, sp. nov. (♂♀, Guangdong), Sinopesa gollum Lin & Li, sp. nov. (♂, Guangdong), Pimoa gagna Zhang & Li, sp. nov. (♀, India), P. nainital Zhang & Li, sp. nov. (♂♀, India), P. shoja Zhang & Li, sp. nov. (♂♀, India), Selenops crewsae Lin & Li, sp. nov. (♀, Sichuan), Sinopoda helii Wang & Li, sp. nov. (♂♀, Sichuan), Phlogiellus jiaxiangi Lin & Li, sp. nov. (♂♀, Guangxi), Episinus bonjovi Lin & Li, sp. nov. (♂♀, Yunnan), E. jiangweni Lin & Li, sp. nov. (♂♀, Yunnan), E. tongyani Lin & Li, sp. nov. (♂♀, Yunnan), Meotipa luoqiae Lin & Li, sp. nov. (♂, Yunnan), M. menglun Lin & Li, sp. nov. (♂, Yunnan), and M. zhengguoi Lin & Li, sp. nov. (♂, Yunnan). The female of Belisana yuexiu Yao & Li, 2020 is reported for the first time. All types are deposited in the Institute of Zoology, Chinese Academy of Sciences (IZCAS) in Beijing, China.
  • Original Paper
    Tao Chen, Xiaoning Chen, Biao Wang, Jianzhen Nie, Ping You
    Zoological Systematics. 2020, 45(4): 243-258.
    Gyrodactylus konovalovi is an ectoparasite on the Amur minnow (Rhynchocypris lagowskii) that is widely distributed in the cold fresh waters of East Asia. In the present study, the phylogeography of G. konovalovi and the distribution of its host in the Qinling Mountains are examined. A total of 102 parasite specimens was sequenced for the mitochondrial cytochrome oxidase subunit 1 (cox1) gene, and 43 haplotypes were obtained. The ratio of substitution sites (dN/dS) was 0.016 and indicated strongly purifying selection. Haplotype diversity (h) and nucleotide diversity (π) of suprapopulations of G. konovalovi varied widely between distinct localities from the Qinling Mountains. Phylogenetic trees based on Bayesian inference (BI), maximum likelihood (ML) and maximum parsimony (MP) methods and network analysis revealed that all haplotypes were consistently well-supported in three different lineages A, B, and C, indicating a significant geographic distribution pattern. There was a significant positive correlation between genetic differentiation (Fst) and geographic distance (P < 0.001). The results of mismatch distribution, neutrality test and Bayesian skyline plot analyses showed that lineages A and B underwent population expansion after the Last Glacier Maximum (LGM) during the Late Pleistocene, while the lineage C underwent population contraction during the Middle to Late Pleistocene. Based on the molecular clock calibration, the most common ancestor was estimated to have emerged in the Middle to Late Pleistocene. Our study suggests that a clearly phylogeography of G. konovalovi was shaped by climatic oscillation and geological events, such as orogenesis, drainage capture changes and vicariance, during the Pleistocene in the Qinling Mountains in central China.
  • Communication
    Junxia Zhang, Jiaxing Lai
    Zoological Systematics. 2020, 45(3): 151-162.
    Phylogenomic approaches are growingly applied in the systematic studies of animals with the rapid advancement of sequencing and analytical tools. Aiming to accelerate the systematic studies using genome-scale data in China, this paper outlines the commonly used techniques for generating phylogenomic datasets: RNA-seq based transcriptome, target-enrichment based UCEs and AHE, RAD-seq, whole genome sequencing, and briefly discusses the pros and cons of each technique. Major analytical procedures for the 1KITE transcriptome pipeline, the PHYLUCE (UCEs) and AHE pipelines and the PLWS (low-coverage whole genome sequencing) pipeline are summarized, and recent achievements valuable for phylogenomic analyses are introduced.
  • Communication
    Shuqiang Li
    Zoological Systematics. 2020, 45(2): 73-77.
    This paper summarizes the advances and challenges in spider taxonomy of China. A table comparing the number of valid genera and species in China, Europe and the world is provided. To date, the Chinese spider fauna is represented by 5,084 species, under 809 genera in 69 spider families. An “All Species Inventory” of spiders in Xishuangbanna Tropical Botanical Garden provides an example of China’s spider species richness, with more than 782 species recorded within a small area. Intensified research in recent years has grown the national catalog of Chinese spiders from 1,050 species in 1983 to 5,084 species today. There are reasons to believe that spider species richness in China will far exceed this grand total. With enhanced support and investment in spider taxonomic research, many more new species will be discovered and described in China
  • Communication
    Michael C. Orr, Arong Luo, Douglas Chesters, Qingsong Zhou, Feng Zhang, Chaodong Zhu
    Zoological Systematics. 2020, 45(2): 78-80.
    The 4th Systematic Biology Forum, China took place in December 2019, covering a vast array of topics across animals, bacteria, fungi, plants, and viruses in the fields of systematics and evolutionary biology. Here, we discuss the emergent themes of this meeting and how these and other interdisciplinary research programs can be synthesized to better understand evolution in a systematic framework.
  • Original Paper
    Feng Zhang, Bruno C. Bellini2, Felipe N. Soto-Adames
    Zoological Systematics. 2019, 44(4): 249-278.
    The largest superfamily Entomobryoidea is one of the key groups in Collembola. However, incorrect recognition of chaetotaxic homology within Entomobryoidea severely impedes the accurate species comparison of adults and phylogenetic reconstruction. Traditional classification of the superfamily at suprageneric level is disputable in the light of recent advances. Transformational homology of tergal chaetotaxy was traced and revised based on 38 species of first instar and partial early instars. Morphological phylogenetic reconstructions were reconstructed mainly relying on first instar characters using both parsimony and likelihood-based algorithms. Outgroup selection and several rogue taxa impacted on resolution and support of otherwise well-supported clades. Integrating published molecular phylogeny, a revised classification of three families and nine subfamilies was presented: Orchesellidae, Entomobryidae and Paronellidae. Orchesellidae includes all basal taxa having a short fourth abdominal segment. Cyphoderidae and Microfalculidae taxa were sunk into Paronellidae. New Paronellidae was divided into two subfamilies: Paronellinae (Paronellini + Cyphoderini + Bromacanthini) and Salininae (Cremastocephalini + Callyntrurini). Microfalculidae (Microfalcula) was closer to Salina and Akabosia and thus transferred into Cremastocephalini. This study erected a new classification framework for Entomobryoidea based upon comprehensive phylogenies. Chaetotaxic homologization across a wide range provided a standardized, comparable, powerful tool for taxonomy.
  • Original Paper
    Heng Zhang, Guiyou Wu, Yanqing Wu, Jianfei Yao, Shuo You, Chencheng Wang, Feng Cheng, Jingjing Chen, Mingxia Tang, Chunlin Li, Baowei Zhang
    Zoological Systematics. 2019, 44(4): 279-293.
    A new species of white-toothed shrew, Crocidura anhuiensis Zhang, Zhang & Li, sp. nov. is described from Mount Huang, China. Genetic sequence (mtDNA Cyt b gene) and morphological (external and skull) data are used to distinguish this newly discovered species. The phylogenetic analysis shows that the materials of this work are monophyletic and allied to C. attenuata by the uncorrected genetic distance 4.9–5.1%, which suggests a species-level divergence. Morphologically, the materials are different from C. attenuata by presenting a greater proportion of tail-to-body ratio and the presence of an obvious tooth root in the mandible. Based on those data, the unnamed species is described as a new species which is currently known only from the Wild Monkey Valley, Mount Huang, Anhui Province, China.
  • Original Paper
    Yuanyuan Lu, Haidong Yang, Ming Bai
    Zoological Systematics. 2019, 44(4): 294-303.
    Three dimensional morphology of Melanopopillia hainanensis Lin, 1980 is reported firstly by using Micro CT technology, including internal and external structures. The male of M. hainanensis is described for the first time. A key to the species of Melanopopillia is given. This study shows an example of the potential application on the 3D morphology in invertebrate taxonomy, especially in the comparison of important structures with complicated 3D morphology, e.g. aedeagus, muscles.
  • Original Paper
    Yao Ji, Xin Yu
    Zoological Systematics. 2019, 44(2): 91-99.
    A molecular study based on COI, 16S and 28S genes reveals that a batch of specimens (7 males and 4 females) of Dysphaea Selys, 1853 collected from central Vietnam, which include different color patterns of wings and body, and were originally identified as three different species, are all the same species. This study implies that, in some group of Odonata, identification only depending on color pattern may be unreliable, no matter what huge variations there are.
  • Original Paper
    Yongfu Gao, Yajun Gong, Ling Ma, Lijun Cao, Jincui Chen, Min Chen, Shujun Wei
    Zoological Systematics. 2019, 44(2): 100-110.
    The melon thrips, Thrips palmi Karny, is an economically important pest on many vegetables from Solanaceae and Cucurbitaceae. In this study, we developed novel microsatellite markers for Thrips palmi. First, we obtained randomly sequenced regions from the genome using next-generation sequencing and assembled 356 Mb genomic sequences. In total, 155,789 microsatellites were identified from the genomic sequences, of which, 64.02% were loci of dinucleotide repeat. Sixty primer pairs were initially validated in seven individuals. Thirty-five polymorphic markers were retained according to the amplification efficiency and polymorphism. These 35 microsatellite loci were then assessed in 96 individuals in four geographical populations collected from China. The allele numbers ranged from 2 to 19 with an average value of 6.9 per locus, while the polymorphism information content ranged from 0.2219 to 0.8490. The observed and expected heterozygosity varied from 0.5252 to 0.5367 and 0.5934 to 0.6148, respectively. Genetic structure was found between populations from southern and northern China. These novel microsatellite markers developed in our study provide abundant molecular markers for the genetic study of T. palmi.
  • Original Paper
    Yanfeng Tong, Fengyuan Li, Yang Song, Haifeng Chen, Shuqiang Li
    Zoological Systematics. 2019, 44(1): 1-75.
    Thirty-two new species of the spider genus Speocera Berland, 1914 from the spider family Ochyroceratidae Fage, 1912 are described from China, Madagascar, and Southeast Asia, namely S. amber Li & Li sp. nov. (♂♀), S. ankalana Li & Li sp. nov. (♂♀), S. bachma Li & Li sp. nov. (♂♀), S. balikpapan Li & Li sp. nov. (♀), S. ballarini Li & Li sp. nov. (♂), S. batang Li & Li sp. nov. (♀), S. bawangling Li & Li sp. nov. (♂♀), S. bontoc Li & Li sp. nov. (♀), S. bukittinggi Li & Li sp. nov. (♂♀), S. cattien Li & Li sp. nov. (♂♀), S. cucphuong Li & Li sp. nov. (♂♀), S. cuyapo Li & Li sp. nov. (♀), S. dongjing Li & Li sp. nov. (♀), S. gexuejuni Li & Li sp. nov. (♂♀), S. griswoldi Li & Li sp. nov. (♀), S. heilan Li & Li sp. nov. (♂♀), S. huifengi Li & Li sp. nov. (♀), S. huisun Li & Li sp. nov. (♀), S. lahrak Li & Li sp. nov. (♂♀), S. longyan Li & Li sp. nov. (♂♀), S. manhao Li & Li sp. nov. (♂♀), S. melinh Li & Li sp. nov. (♀), S. nuichua Li & Li sp. nov. (♂♀), S. payakumbuh Li & Li sp. nov. (♂♀), S. suea Li & Li sp. nov. (♂♀), S. tabuk Li & Li sp. nov. (♀), S. trapezialis Li & Li sp. nov. (♂♀), S. trusmadi Li & Li sp. nov. (♂♀), S. tubularis Li & Li sp. nov. (♂♀), S. xiaoxiaoae Li & Li sp. nov. (♂♀), S. xuanson Li & Li sp. nov. (♂♀), S. zhigangi Li & Li sp. nov. (♂♀). Types of all new species are deposited in the Institute of Zoology, Chinese Academy of Sciences (IZCAS) in Beijing, China and the California Academy of Sciences (CAS) in San Francisco, USA.
  • Original Paper
    Bingyan Li, Qiao Gao, Lijun Cao, Ary Anthony Hoffmann, Qiong Yang, Jiaying Zhu, Shujun Wei
    Zoological Systematics. 2018, 43(4): 341-355.
    Double-digested Restriction Site Associated DNA Sequencing (ddRAD) through next-generation sequencing (NGS) generates large numbers of loci for characterizing genome-wide variation among multiple samples using next-generation sequencing. Different combinations of restriction endonucleases (REs) may produce varying size distributions of digested fragments, which affect the number of genotyped loci. Understanding digestion profiles across different species will help in selecting REs for digestion in a particular organism. In this study, we use of genome sequences to compare the in silico digestion profile of 26 combinations of REs in 131 insect species with two simulation programs. The number of digested fragments in the 300–450 bp range increases linearly with the size of the genome. Different species and insect orders showed similar profiles when digested by different combinations of REs in silico, indicating the conservation of digestion by double enzymes in insect genomes. Combinations with NlaIII or TaqαI usually produced higher number of fragments in the range 300–450 bp, while combinations with EcoRI or MluCI produced fewer fragments. The proportion of fragments with the same overhangs at the two ends of digested DNA was higher than those with different overhangs. The two four-base enzyme pairs produced more fragments in the 300–450 bp range than pairs of four-base + six-base enzymes. Experimental digestion of three species from Hymenoptera, Lepidoptera and Thysanoptera showed profiles congruent with in silico expectations. Our results shed light on understanding the digestion profiles of insect genomes and provide guidance on selecting REs for ddRAD projects.
  • Original Paper
    Xiaoya Sun, Yanhui Wang, Pingping Chen, Hesheng Wang, Lixiang Lu, Zhen Ye, Yanzhuo Wu, Teng Li, Wenjun Bu, Qiang Xie
    Zoological Systematics. 2018, 43(4): 356-386.
    Few studies have explored the differences between Sanger and HTS methods in the results of mitogenome sequencing. We used a single individual of insect to study the differences between the sequences given by Sanger and PCR-free HTS methods. Here we provided evidence for biased results of sequencing due to different methods in the mitochondrial genes of atp6, atp8, cox1, cox2, cox3, Cytb, nad2, nad3, nad4, nad5, rrnS, rrnL, trnH, trnI, and control region at various degrees. Especially, in cox1, the differently sequenced nucleotides account for 2.6% of the complete length. Furthermore, the highest value of the intraspecific genetic distance based on K2P accounts for 2.5% using a barcode fragment size of cox1 (651 bp, Sanger), while the maximum distance of the corresponding cox1 fragment obtained by the two sequencing methods was 5.0%. We revealed that the methods of Sanger and HTS may give different sequencing results of mitochondrial genes, which may reflect the heteroplasmy of mitogenomes within an insect individual. Therefore, researchers should be very cautious in using the mixed data of a gene given by different methods of sequencing.
  • Original Paper
    Lan Nie, Yunliang Wang, Dunyuan Huang, Ruisong Tao, Chengyong Su, Jiasheng Hao , Chaodong Zhu
    Zoological Systematics. 2018, 43(4): 387-409.
    In this study, the complete mitochondrial genomes of four pierid butterfly species, namely Batltia butleri, Talbotia naganum, Pontia callidice and P. daplidice, were newly sequenced and characterized. Meanwhile the phylogenetic relationships of the main Pieridae lineages covering 22 pierid butterfly species, were reconstructed with Bayesian inference and maximum likelihood methods based on different mitogenomic datasets, including the concatenated 13 PCGs, 13 PCGs + 2 rRNAs, 2 rRNAs, 2rRNAs + 22 tRNAs and 22 tRNAs sequences, respectively. Our results of mitogenomic analysis showed that the four mitogenomes were 15,124, 15,155, 15,109, 15,124 bp in size, with the gene orders and arangements identical to all other butterflies determined. Our results of phylogenetic analyses upon protein coding genes, rRNA genes or their sequence combinations indicated that the three pierid subfamilies of this study were all monophyletic, with their relationships of being (Dismorphiinae, (Pierinae, Coliadinae)); the phylogenetic relationship of Pierini of this study was (Pieris, (Baltia, (Talbotia, Pontia))) + (Prioneris, (Delias, Aporia)). Additionally, our analyses suggested that only mitogenomic tRNA sequence datasets were not ready to be utilized in resolving the deeper phylogeny, whereas somewhat suitable to be applied in clarifying the phylogenies of closely related species, of the pierid butterflies.
  • Original Paper
    Kahong Cheung, Ming Bai, Manhin Leung, Yingming Lee, Kinfung Chan, Xingke Yang
    Zoological Systematics. 2018, 43(3): 233-267.
    Scarabaeinae is a globally distributed coprophagous coleopteran group with southeast Asia and Africa as the key areas of distribution. Located in the southern part of China, Hong Kong has subtropical climate and diverse habitats supporting a high diversity of scarabaeines. This paper gives a detailed review on Scarabaeinae from Hong Kong, including the new records to Hong Kong, photographic records of type specimens from Hong Kong, and identification keys to genera and species. A total of 13 genera and 51 species of scarabaeines were recorded in Hong Kong. Among them, 14 species were new records to Hong Kong found in the authors’ recent surveys, viz. Caccobius (Caccophilus) brevis Waterhouse, 1875, Copris (s. str.) punctatus Gillet, 1910, C. (s. str.) szechouanicus Balthasar, 1958, Microcopris propinquus Felsche, 1910, Onitis excavatus Arrow, 1931, Oni. intermedius Frivaldszky, 1892, Oni. subopacus Arrow, 1931, Onthophagus brutus Arrow, 1931, Ont. tricolor Boucomont, 1914, Ont. (Matashia) lenzi Harold, 1874, Ont. (s. str.) roubali Balthasar, 1935, Ont. (Parascatonomus) horni Balthasar, 1935, Scarabaeus (Kheper) erichsoni Harold, 1867 and Sisyphus (s. str.) indicus Hope, 1831. Onthophagus (Paraphanaeomorphus) trituber jacobsoni Kabakov, 2006 was found to be a new synonym of the nominate subspecies. Catharsius davidis Deyrolle & Fairmaire, 1878 and Copris davidis (Deyrolle & Fairmaire, 1878) were found to be the new synonyms of Catharsius molossus (Linnaeus, 1758). Hong Kong was also the only locality where Liatongus pugionatus and Onthophagus convexicollis were recorded over the world.
  • Original Paper
    Xi Zhang, Hongzhang Zhou
    Zoological Systematics. 2018, 43(2): 125-138.
    Global species diversity is a historical result of speciation minus extinction and can be exhibited by phylogenetic patterns, whereas speciation is a process that may concern reproductive isolation and relating, at least in most sexual reproductive insects or other similar animals, to genital morphological evolution. The aedeagus is male genital organ that determines valid mating and reproductive isolation. However, definite correlation between aedeagus variation and species richness has not yet been clearly demonstrated. Here the phylogenetic tree of the rove beetle subfamily Staphylininae is built up based on 3085 bp DNA sequences of cytochrome oxidase subunit I (COI), nuclear 28S rDNA, nuclear genes wingless (Wg) and topoisomerase I (TP). Branching times and confidence intervals are calculated with molecular clock calibrations based on rove beetle fossil records. Different types of aedeagus, namely, median lobe plus a single paramere (Asp), a pair of parameres (App), or a single but bifurcated paramere (Abp), are marked on the phylogenetic tree and the genital morphological variations are compared in different genus-level taxa. The result shows that active cladogeneses occurred during late Cretaceous to late Paleogene, with those clades of Staphylininae evolved to be a more species-rich ones accompanied by larger aedeagus modifications. This implies that male genital morphological evolution might promote rove beetle speciation.
  • Original Paper
    Xiaoyue Zhu, Zihui Wang, Shun Yao, Jiasheng Hao
    Zoological Systematics. 2018, 43(2): 139-155.
    In this study, the first complete mitochondrial genome (mitogenome) of lepidopteran family Drepanidae (superfamily Drepanoidea) was reported with the notes about its phylogenetic implications. The Oreta fuscopurpurea mitogenome is 15,564 bp in length, including 13 protein-coding genes, two ribosomal RNAs (lrRNA and srRNA), 22 transfer RNAs and a non-coding control region (AT-rich region), with a 79.6% A+T content. The gene orientation and arrangement of the mitogenome are the same as other sequenced lepidopterans. All protein-coding genes usually start with the common ATN codon except for COI gene, which used CGA as the initial codon; eight PCGs use a typical stop codon of TAA, whereas the remaining PCGs use incomplete stop codon of T or TA. All tRNAs have the typical clover-leaf structure with the exception of tRNASer (AGN). The lrRNA and srRNA genes are 1,409 bp and 778 bp in size respectively, with the former harboring one (TA)13 microsatellite-like repeat and an 17 bp insertion. The 20 intergenic spacers totaling of 184 bp and 8 overlapping sequences totaling of 25 bp are scattered throughout the whole mitogenome. The 526 bp AT-rich region contains some structures characteristic of lepidopterans, such as the motif ATAGA preceded by an 19 bp poly-T stretch, a tRNA-like and a sterm-loop structures. Phylogenetic analysis of the Oreta fuscopurpurea with other 47 insect species covering 20 lepidopteran families were conducted based on the sequence data of the 13 mitogenomic protein coding genes with maximum likelihood (ML) and Bayesian inference (BI) methods, and the results showed distinctly that the superfamily Drepanoidea was sister to the clade of (Bombycoidea, Lasiocampoidea) + (Noctuoidea, Geometroidea).
  • Original Paper
    Yongxia Chen, Hao Chen, Dekui He, Yifeng Chen
    Zoological Systematics. 2018, 43(2): 156-168.
    Two spined loaches, Cobitis oxycephala Chen YX & Chen YF, sp. nov. and C. brachysoma Chen YX & Chen YF, sp. nov., are described based on specimens collected from Guangdong and Hainan Provinces, South China. Morphology features and sequences of mitochondrial cytochrome b gene of these two new species were analyzed. Morphological and molecular data show that C. oxycephala Chen YX & Chen YF, sp. nov. is closely related to C. dolichorhynchus Nichols, and C. brachysoma Chen YX & Chen YF, sp. nov. is closely related to C. microcephala Chen & Chen. These two new species can be distinguished from their congeners by the combination of characteristics of body shape, color pattern, lamina circularis, mental lobes, suborbital spine, subdorsal scales, and caudal peduncle.
  • Original Paper
    Bin Zheng, Yunliang Wang, Chenchen Xia, Dunyuan Huang, Yan Cao, Jiasheng Hao, Chaodong Zhu
    Zoological Systematics. 2018, 43(1): 1-17.
    In this study, the complete mitochondrial genome of Parnassius actius (Lepidoptera: Papilionidae: Parnassinae) was reported. Meanwhile, the phylogenetic position of this Apollo species was inferred from this sequence combined with other available related sequence data. The results showed that the whole mitogenome is 15,386 bp in length, containing 13 typical protein-coding genes (PCGs), 22 transfer RNA genes (tRNAs), two ribosomal RNA genes (rRNAs) and a noncoding control region (A+T-rich region). The mitogenomic gene orientation and arrangement are the same with other sequenced Parnassius species. Thirteen protein coding genes is 11,195 bp in size, encoding 3,720 amino acids totally; the lrRNA and srRNA genes are 1,351 bp and 779 bp, respectively. Eleven intergenic spacers ranging from 1 to 40 bp (129 bp in total), 10 overlapping regions from 1 to 8 bp (26 bp in total), and a 504 bp A+T-rich region harboring some Parnassius-specific sequences are scattered throughout the whole mitogenome. The maximum likelihood and Bayesian phylogenetic analyses showed that the genus Parnassius of this study contained five major clades which can be categorized into the eight subgenera reported by the previous molecular studies; additionally, the P. actius was shown to be more closely related to P. nomion than other Parnassius species, suggesting that the P. actius stands as an effective species with its phylogenetic position awaiting further studies.
  • Original Paper
    Ruixin Leng, Kai Heller, Junhao Huang, Lan Ye, Hong Wu
    Zoological Systematics. 2018, 43(1): 18-26.
    Species of the genus Dichopygina Vilkamaa, Hippa & Komarova, 2004 has been morphologically cryptic. In this paper, eight species of the genus were clearly defined by DNA barcodes. Based on both molecular and morphological evidences, D. bernhardi Vilkamaa, Hippa & Komarova, 2004, stat. rev. is separated from D. perfecta (Pettey, 1918); a closely allied new species, D. tibetana Leng, Heller & Huang, sp. nov., is described from Tibet, China. Detailed descriptions and figures of the two species are presented.
  • Original Paper
    Jingyoh Zaelor, Sangvorn Kitthawee
    Zoological Systematics. 2018, 43(1): 27-36.
    Fruit flies in Bactrocera tau complex are important pests, but their morphological taxonomy is still unsettled. In this paper, specimens from gac fruits (Momordica cochinchinensis) are attempted to classify by geometric morphometric and genetic analyses. Geometric morphometric study of wing venation and aculeus shape among samples suggested a new species in this taxon. Sequences in the cytochrome oxidase I region were compared with reference material in this complex, as well as other taxa within the same subgenus. Phylogenetic analysis based on COI revealed a new species of the B. tau complex, B. tau form J. Therefore, the criteria of body color patterns proved to be taxonomically valid for species classification and identification in this species complex.
  • Original Paper
    Chang Liu, Fengyuan Li, Shuqiang Li, Guo Zheng
    Zoological Systematics. 2017, 42(4): 395-417.
    Five new genera of the spider family Ochyroceratidae with remarkable palpal and epigynal characters are described: Luzonacera Li & Li, gen. nov., Qiongocera Li & Li, gen. nov., Relictocera Li & Li, gen. nov., Sinoderces Li & Li, gen. nov., and Thaiderces Li & Li,gen. nov. Six new species are described based on specimens collected in China, Philippines, Vietnam, and Thailand: Luzonacera chang Li & Li, sp. nov., Luzonacera duanLi & Li, sp. nov.,Qiongocera hongjunensisLi & Li, sp. nov., Relictocera qiyiLi & Li, sp. nov., Sinoderces nawanensisLi & Li, sp. nov., and Thaiderces jianLi & Li, sp. nov. In addition, two Psiloderces species are transferred to other genera: Sinoderces exilis (Wang & Li, 2013) comb. nov. and Thaiderces vulgaris (Deeleman-Reinhold, 1995) comb. nov. All type specimens are deposited in the Institute of Zoology, Chinese Academy of Sciences (IZCAS) in Beijing.
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