Three new rock-dwelling species of Cathaica Möllendorff, 1884 from the Taihang Mountains, northern China (Stylommatophora: Camaenidae)

CATHAICA (贝类多样性研究), 1 (5): 47–68, October 25, 2025 DOI: https://doi.org/10.5281/zenodo.17430005 ISSN 2755-9653 | ISSN 2755-9661 (Online) ©The author. This work is freely available under the Creative Commons Attribution 4.0 International licence (CC BY 4.0) Three new rock-dwelling species of Cathaica Möllendorff, 1884 from the Taihang Mountains, northern China (Stylommatophora: Camaenidae) Zhi-Yao Wang1,#,*, Quan-Yu Zhang2,#, Yue-Ming He3, Hui Chen4 & Shi-Yang Feng5 1 College of Plant Protection, China Agricultural University, Beijing 100091, China; https://orcid.org/0009-0001-3113-4058 2 College of Music and Dance, Henan Normal University, Xinxiang 453007, China; https://orcid.org/0009-0006-8534-1710 3 College of Engineering and Technology, Tianjin Agricultural University, Tianjin 300384, China; https://orcid.org/0009-0002-9849-7977 4 School of Life Sciences, Nanchang University, Nanchang 330031, China; https://orcid.org/0009-0004-5222-3975 5 Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu 610041, China; https://orcid.org/0009-0004-9290-8733 Abstract. Species of Cathaica are widespread in the temperate montane regions of China, exhibiting high diversity and a broad spectrum of shell morphologies that reflect adaptation to heterogeneous environments. Here we describe three new Cathaica species from the Taihang Mountains of Henan and Hebei, China: Cathaica sculptilis Wang, Chen, He & Zhang, sp. nov., C. zhangcunxiangi Wang, Chen, He & Zhang, sp. nov., and C. wangjiaxuni Wang, Chen, He & Zhang, sp. nov. Two of these, C. sculptilis sp. nov. and C. wangjiaxuni sp. nov., share shell features with C. multicostata G. Zhang, 2023. Integrative morphological and molecular analyses support recognition of all three new species. The genital anatomical information of C. mengi is also provided for the first time. Key words. Comparative morphology, molecular phylogeny, new species, northern China, rocky habitats Introduction Cathaica Möllendorff, 1884 is a specious genus in Camaenidae, with 53 species found in China (MolluscaBase, 2025). A subgenera system of Cathaica was proposed by Andreae (1900) and adapted by many authors (e.g., Gude, 1902; Thiele, 1931; Zilch, 1960; Richardson, 1983; Schileyko, 2004), namely Cathaica (s.str.), Cathaica (Pliocathaica) Andreae, 1900, Cathaica (Xerocathaica) Andreae, 1900, and Cathaica (Campylocathaica) Andreae, 1900. However, Cathaica is no longer considered monophyletic based on recent works, and the status of the subgenera also remains uncertain, as the type species of each requires a comprehensive study to confirm its placement (Wu et al., 2023; Wu & Zhang, 2024). Two recurrent problems were produced by previous research relying solely on shell morphology: the unrelated species have been included in Cathaica, as with Bradybaena brevispira (H. Adams, 1870) (Wu & Zhang, 2024); and distinct species with similar shells have been treated as conspecific, as in Cathaica pyrrhozona (R. A. Philippi, 1845) and C. fasciola (Draparnaud, 1801), which were separated only recently through anatomical and #These authors contributed equally to this work. *Corresponding author: wangchuan199811[email protected] http://zoobank.org/urn:lsid:zoobank.org:pub:F837079A-8AAE-4221-AA22-DA01CEC34B08 phylogenetic evidence (Suzuki, 1939; Chen & Zhang, 2004; Wu, 2004; Zhang & Wade, 2023). These issues underscore the need for integrative evidence beyond shells. Here we combine shell morphology, reproductive anatomy, and molecular phylogenetics to describe three new species of Cathaica from the Taihang Mountains of northern China, namely C. sculptilis sp. nov., C. zhangcunxiangi sp. nov., and C. wangjiaxuni sp. nov. All three exhibit a prominent keel, with C. sculptilis sp. nov. and C. wangjiaxuni sp. nov. also possess fine ribs. Among described congeners, only C. multicostata G. Zhang, 2023 and C. mengi Yen, 1935 present a distinct keel with strong ribs (Yen, 1939; Zhang et al., 2020b; Zhang & Wade, 2023). To clarify relationships, we surveyed the type localities and adjacent areas of C. multicostata and C. mengi and provide additional conchological and anatomical information for C. mengi. Materials and methods Materials and morphological examination All specimens were collected in the Taihang Mountains (Hebei, Henan, Shanxi) and Taishan Mountain (Shandong) in the years 2023 and 2024. Specimens were preserved in 95% ethanol and were deposited in the Mollusc collection of the Museum of Hebei University, Baoding, China (HBUMM), Yu-Xuan Huan private collection (HYX), Zhi-Yao Wang private collection (WZY) and Quan-Yu Zhang private collection (ZQY). Photographs were taken with a SOPTOP SZX12 stereomicroscope and a OD500F camera (Sunny Optical Technology, China). DNA extraction, PCR amplification, and phylogenetic analyses Total genomic DNA was isolated from a small piece of tissue taken from the foot of each ethanol-preserved specimen using a TreliefTM Animal Genomic DNA kit (Tsingke®). Partial sequences of 16S rDNA were amplified using the universal primer set 16Sar and 16Sbr (Palumbi et al., 1991). Partial sequences of COI were amplified using LCO1490 and HCO2198 (Folmer et al., 1994). A partial fragment of the ITS2 gene was amplified using the primer pair LSU-1 and LSU-3 (Wade & Mordan, 2000). Both ends of the sequences were obtained by automated sequencing on an Applied Biosystems 3730 platform at Sangon Biotech Co. Ltd. (Shanghai, China). Eighteen specimens representing eight species (based on COI, 16S and ITS2 data) were used in this study (Table 1). Sequences were aligned using MAFFT v. 7.505 based on the L-INS-i method (Katoh & Toh, 2008). Pairwise distances between species were calculated using MEGA X (Kumar et al., 2018). Aligned sequences were concatenated in PhyloSuite v.2.3 (Zhang et al., 2020a). The best substitution model was selected using the corrected Bayesian Information Criterion (BIC) in ModelFinder v.2.2.0 (Kalyaanamoorthy et al., 2017). For Bayesian analysis, two runs were performed simultaneously with four Markov chains starting from a random tree. Bayesian inference and maximum likelihood analysis were performed using MrBayes v.3.2.7 (Ronquist et al., 2012) and IQ-TREE v.2.2 (Minh et al., 2013), respectively, with reference to the selected model of sequence evolution. Bayesian posterior probabilities (BPPs) of nodes were determined using Metropolis-coupled Markov chains (one cold chain) for 2,000,000 generations, sampled every 1,000 generations. The first 25% of sampled trees were discarded as burn-in when the standard deviation of split frequencies from the two runs was less than 0.01; the remaining trees were then used to create a 50% majority-rule consensus tree and to estimate BPPs. Node support for the maximum likelihood analysis was determined using 1000 rapid bootstrap (BS) replicates. WANG ET AL.48 49Three new species of Cathaica from northern China Species Voucher number COI 16S ITS2 Location Cathaica mengi HBUMM10083a PV446475 PV446368 PV446359 Yuncheng City, Yongji City, Shanxi Province, China Cathaica mengi HBUMM10083b PV446476 PV446369 PV446360 Yuncheng City, Yongji City, Shanxi Province, China Cathaica mengi HBUMM10083c PV446370 Yuncheng City, Yongji City, Shanxi Province, China Cathaica multicostata WZY20240424A/1 PV446477 PV446371 PV446361 Zaozhuang City, Shandong Province, China Cathaica multicostata WZY20240424A/2 PV446478 PV446372 PX453178 Zaozhuang City, Shandong Province, China Cathaica sculptilis sp. nov. HBUMM10084 PV446479 PV446373 PV446363 Gongyi City, Zhengzhou City, Henan Province, China Cathaica sculptilis sp. nov. HBUMM10085a PV446480 PV446374 PV446364 Gongyi City, Zhengzhou City, Henan Province, China Cathaica zhangcunxiangi sp. nov. HBUMM10086 PV446473 PV446366 Huixian City, Xinxiang City, Henan Province, China Cathaica zhangcunxiangi sp. nov. HBUMM10087a PV446474 PV446367 PV446358 Huixian City, Xinxiang City, Henan Province, China Cathaica wangjiaxuni sp. nov. HBUMM10088 PV446471 see Appendix 1 PV446357 Wuan City, Handan City, Hebei Province, China Cathaica wangjiaxuni sp. nov. HBUMM10089a PV446472 PV446365 PX453179 Wuan City, Handan City, Hebei Province, China Cathaica sp1 WZY20240422A/1 N/A N/A N/A Wuan City, Handan City, Hebei Province, China Cathaica sp1 WZY20240422A/2 N/A N/A N/A Wuan City, Handan City, Hebei Province, China Cathaica sp1 WZY20240422A/3 N/A N/A N/A Wuan City, Handan City, Hebei Province, China Cathaica sp2 HYX20240714A/1 N/A N/A N/A Zibo City, Shandong Province, China Cathaica sp2 HYX20240714A/2 N/A N/A N/A Zibo City, Shandong Province, China Pseudiberus tectumsinense PV391143 PV391021 Liantaishan Mout, Jinan City, Shandong Province, China Ponsadenia duplocincta MN369704 PP668898 N/A TABLE 1.TABLE 1. Vouchers, localities, and GenBank accession numbers for all samples used in phylogenetic analysis of this study. Vouchers, localities, and GenBank accession numbers for all samples used in phylogenetic analysis of this study. Colour scheme for anatomical illustrations To present the reproductive anatomy more clearly, we use a simple, consistent colour scheme in the line drawings that minimises confusion from overlapping structures. In genital overviews, the mucous glands are blue, the bursa copulatrix and its duct are red, and all remaining parts are yellow. In detailed dissected views, blue again denotes the mucous glands; red marks the opening of the proximal accessory sac, the entrances of the mucous glands, and the penial opening; yellow indicates the remaining structures. Abbreviations At, atrium; BC, bursa copulatrix; BCD, bursa copulatrix duct; DS, dart sac; Dt, love dart; DtC, love-dart chamber, the chamber secreting and containing the love dart; FO, free oviduct; MG, mucous glands; MGE, entrance(s) of mucous glands; P, penis; PAS, proximal accessory sac, a blind sac on WANG ET AL.50 ID Species 1 2 3 4 5 6 7 8 1Cathaica sp1 2Cathaica wangjiaxuni sp. nov. 0.46–0.92 3Cathaica zhangcunxiangi sp. nov. 5.86–6.93 5.39–6.33 4Cathaica mengi 8.17–8.78 8.01–8.47 8.47–9.55 5Cathaica sp2 6.63–7.55 6.63–6.78 5.7–5.86 7.55–7.86 6Cathaica multicostata 6.47–7.24 6.16–6.47 6.16–6.78 8.47–8.94 4.93–5.08 7Cathaica sculptilis sp. nov. 4.62–5.39 4.16–4.78 5.86–6.63 7.4–7.7 6.78–6.93 6.01–6.32 8Pseudiberus tectumsinense 17.41 16.95–17.1 17.87 17.26–17.41 18.34 18.34–18.49 16.95 9Ponsadenia duplocincta 18.06–18.23 18.06–18.23 18.58–18.75 17.19 17.88 18.92–19.1 17.36–17.71 14.24 TABLE 2.TABLE 2. Genetic differentiation of Genetic differentiation of COCOI by means of I by means of pp distances. distances. FIGURE 1.FIGURE 1. Bayesian inference (BI) tree for the Cathaica species. Values (BPP/BS) at nodes Bayesian inference (BI) tree for the Cathaica species. Values (BPP/BS) at nodes represent Bayesian posterior probabilities and Bootstrap values.represent Bayesian posterior probabilities and Bootstrap values. proximal dart sac and opening into dart sac chamber or elsewhere; PE, entrance of penis; PO, opening of proximal accessory sac; PR, penial-retractor muscle; PS, penis sheath; SD, dart-sac septum, a fleshy septum between the atrial opening and the opening of the DtC; VD, vas deferens. Directions used in descriptions of genitalia: proximal, towards the genital atrium; distal, away from the genital atrium; left and right side of dart sac (see Wu et al., 2023: fig. 1A). Results The alignments of the COI, 16S and ITS2 sequences are 649, 359 and 891 base pairs, respectively. The HKY+I+F model was selected as the best-fit model of nucleotide substitution based on the BIC criterion. Ponsadenia duplocincta (Martens, 1879) was used as the outgroup to root the phylogenetic tree. Both Bayesian Inference (BI) and Maximum Likelihood (ML) methods produced identical topologies (Fig. 1). Three new species are supported: Cathaica zhangcunxiangi sp. nov. (BPP=1, BS=100), C. sculptilis sp. nov. (BPP=1, BS=91) and C. wangjiaxuni sp. nov. (BPP=0.96, BS=80). The species Cathaica zhangcunxiangi sp. nov., C. sp2, C. multicostata, C. sculptilis sp. nov., C. wangjiaxuni sp. nov. and C. sp1 are nested in a monophyletic group with good support (BPP=1, BS=85), and form a sister lineage to the C. mengi (BPP=1, BS=100). Cathaica sp2 is the sister lineage of C. multicostata (BPP=1, BS=92). C. wangjiaxuni sp. nov. and C. sp2 form a sister groups in the tree (BPP=1, BS=99), with C. sculptilis sp. nov. falling outside of this grouping (BPP=1, BS=84). The genetic distances among Cathaica species based on COI sequences range from 4.16% to 9.55% (Table 2). 51Three new species of Cathaica from northern China FIGURE 2.FIGURE 2. Shell of Shell of Cathaica multicostataCathaica multicostata. . A.A. WZY20240424A/1. WZY20240424A/1. B. B. WZY20240424A/2. WZY20240424A/2. C. C. WZY20240424A/3.. Systematics Family Camaenidae Pilsbry, 1895 Subfamily Bradybaeninae Pilsbry, 1934 (1898) Genus Cathaica Möllendorff, 1884 Type species. Helix pyrrhozona Philippi, 1845, by original designation. Cathaica multicostata G. Zhang, 2023 多肋华蜗牛 (Fig. 2) Cathaica multicostata G. Zhang in Zhang & Wade, 2023: 573, figs 1, 3E, 4D, 5, 6. Type locality. “Baodu, Zaozhuang, Shandong; 34.984°N, 117.721°E, 520 m a.s.l.” Material examined. WZY20240424A (10 specimens). Mount Baodugu [抱犊崮], Zaozhuang City, Shandong Province, China, 34°59’N, 117°43’E, 500 m above sea level, leg. Zhi-Yao Wang, 24 April 2024. Measurements (n = 3): Shell width: 16.6–17.7 mm; shell height: 6–6.7 mm. Cathaica mengi Yen, 1935 孟氏华蜗牛 (Figs 3, 4, 12A, 14D–F) Cathaica mengi Yen 1935: 35, pl. 2, figs 12–12b; Zhang et al., 2020b: 62, figs 5A–C. Type locality. “In the mountain near Yu-hsiang Hsien, Shansi” Material examined. HBUMM10083 (5 specimens), Yuxiang County [虞乡镇], Yongji City [永 济市], Yuncheng City [运城市], Shanxi Province, China; 34°50’N, 110°38’E, 400–1000 m above sea level, leg. Shi-Yang Feng, Quan-Yu Zhang and Yue-Ming He, 24 April 2024; WZY20250621A (3 specimens), same location as above, leg. Zhi-Yao Wang, 21 June 2024; HYX20250905A (2 specimens), Shifosicun [石佛寺村], Yuxiang County, Yuncheng City, Yongji City, Shanxi Province, China, 34°49’N, 110°35’E, 500 m above sea level, leg. Yu-Xuan Huan, 5 September 2024. Redescription. Shell (Fig. 3) slightly depressed to conoid, thin, dextral. Spire conoid, elevated. Whorls slightly convex. Suture superficial, slightly deeper on the last whorl. Umbilicus a slit to narrow; transition from shell to umbilicus gradual. . Columella oblique; columellar lip slightly expanded, slightly or almost completely covering umbilicus. Protoconch radially and densely granulate (Fig. 11A). Spiral furrows absent. Shell surface with low ribs, approximately 60–100 on body whorl, forming crenulations at periphery. Growth lines between ribs indistinct. Young shell carinate. Adult shell smooth; last whorl carinate at periphery, with base convex. Aperture roundedly quadrate, slightly expanded below, oblique, with a well-developed basal tooth. Peristome sometimes weakly sinuate, not continuous, thick. Parietal callus thin and indistinct. Shell glossy. Protoconch and conchial whorls show no obvious color difference, apically light brown to cream yellow. Ventral surface milk white, with a dark brown peripheral band under keel. Measurements: High altitude WANG ET AL.52 population (n = 3): Shell width: 17.6–18.4 mm; shell height: 9.4–9.4 mm. Low altitude population (n = 3): Shell width: 17.2–18.9 mm; shell height: 10.6–11.7 mm. Genitalia (Fig. 4). Membranous sac surrounding terminal genitalia absent. Penial sheath approximately 1/3 penis length. Penis slender, simple outside. Flagellum absent. Vas deferens thickened near penial-retractor muscle, thickened portion accounts for about 2/5 of vas deferens. Mucous glands containing 10–15 tubules, similar in length to dart sac, inflated, straight and usually unbranched, if branched, then only simply so. Each tubule with a distinct peduncle, not attached to vagina by connective tissue, opening into dart-sac chamber. Proximal part of dart sac neither elongate nor forming a neck-like structure. Dart sac containing a single dart, curved and possessing chaeta, cross-section fusiform, surface covered with dense villi except at the distal end, approximately 8 mm in length. Proximal accessory sac absent. Vagina entering atrium. Measurements (average of three individuals): DS: 9.1 mm in length, 2.4 mm in width; MG: 6.0–9.0 mm; PS+P: 14.8 mm; VD: 19.3 53Three new species of Cathaica from northern China FIGURE 3.FIGURE 3. Shell of Shell of Cathaica mengiCathaica mengi. . A.A. HBUMM10083a. HBUMM10083a. B.B. HBUMM10083b. HBUMM10083b. C. C. HBU-HBUMM10083c. MM10083c. D.D. HBUMM10083d. HBUMM10083d. mm; PR: 10.6 mm; Va: 6.1 mm; FO: 1.1 mm; BC+BCD: 14.0 mm. Distribution. China: Shanxi. Ecology. Cathaica mengi occurs across a wide elevational gradient, from lowland farmland at the mountain base to exposed rocky habitats at higher elevations. (Fig. 14A–C). Remarks. Cathaica mengi can be readily distinguished from other Cathaica species by its carinate periphery and densely spaced, low ribs. Among all Cathaica examined to date, C. mengi is the only species known to lack a proximal accessory sac. The species shows ecological adaptability. Foothill populations of C. mengi tend to have a higher spire (Fig. 3D), whereas populations on rocky cliffs have a lower spire with sparser, more sharply defined ribs (Fig. 3A–C). This pattern indicates a strong environmental influence on shell form, and a lower spire is likely advantageous in exposed rocky habitats. Cathaica sculptilis Z.-Y. Wang, Y.-M. He, H. Chen & Q.-Y. Zhang, sp. nov. 雕刻华蜗牛 (Figs 5, 6, 12B, 16) Type materials. Holotype. HBUMM10084, Qinglongshan Mount [青龙山], Dayugou County [大峪 沟镇], Gongyi City [巩义市], Zhengzhou City [郑州市], Henan Province, China, 34°40’N, 113°04’E, 500 m above sea level, leg. Zhi-Yao Wang, 17 September 2023. Paratypes. HBUMM10085 (9 specimens), WZY20230917A (5 specimens), same data as holotype; ZQY20230126A (5 specimens), same location as holotype, leg. Quan-Yu Zhang, 26 January 2023. Etymology. This new species is named for its prominent ribs and remarkable surface sculpture. Description. Shell (Fig. 5) low-conic, thin, dextral; spire conoid. Whorls convex. Suture superficial. Umbilicus narrow, with a gradual transition from the shell base. Columella oblique; columellar lip dilated, partially covering umbilicus. Protoconch radially granulate (Fig. 11B). Spiral furrows densely distributed on spire. Body whorl clearly deflexed behind aperture. Shell surface bearing 60–70 strong ribs, each rib thickened and protruding at keel, forming crenulations at periphery. Growth lines between ribs indistinct. Both young and adult shells smooth. Young shell WANG ET AL.54 FIGURE 4.FIGURE 4. Genitalia of Genitalia of Cathaica mengiCathaica mengi. . A–B.A–B. both sides of the genitalia. both sides of the genitalia. C.C. love dart. love dart. D.D. exposed exposed dart sac apparatus.dart sac apparatus. 55Three new species of Cathaica from northern China FIGURE 6.FIGURE 6. Genitalia of Genitalia of Cathaica sculptilisCathaica sculptilis sp. nov.sp. nov. A–B.A–B. both sides of the genitalia. both sides of the genitalia. C.C. love dart. love dart. D.D. exposed dart sac apparatus. exposed dart sac apparatus. FIGURE 5.FIGURE 5. Shell of Shell of Cathaica sculptilisCathaica sculptilis sp. nov.sp. nov. A.A. HBUMM10084, holotype. HBUMM10084, holotype. B.B. HBUMM10085a, HBUMM10085a, paratype. paratype. C.C. HBUMM10085b, paratype. HBUMM10085b, paratype. Cathaica sp2 (Figs 12C, 14) Material examined. HYX20240714A (3 specimens), East of Zibo City [淄博市], Shandong Province, China, leg. Huan-Yu Xuan, 14 July 2024. Measurement (n = 1): Shell width: 17.9 mm; shell height: 10.3 mm. Remarks. The genital anatomy of C. sp2 closely matches the description of C. fasciola provided WANG ET AL.62 FIGURE 14.FIGURE 14. Genitalia of Genitalia of Cathaica Cathaica sp2. sp2. A–B.A–B. both sides of the genitalia. both sides of the genitalia. C.C. love dart. love dart. D.D. exposed exposed dart sac apparatus.dart sac apparatus. FIGURE 13.FIGURE 13. Genitalia of Genitalia of Cathaica Cathaica sp1. sp1. A–B.A–B. both sides of the genitalia. both sides of the genitalia. C.C. love dart. love dart. D.D. exposed exposed dart sac apparatus. The positions indicated by the red arrows were originally connected.dart sac apparatus. The positions indicated by the red arrows were originally connected. 63Three new species of Cathaica from northern China FIGURE 15.FIGURE 15. Natural habitats of Natural habitats of CathaicaCathaica spp. spp. A–C.A–C. Cathaica multicostataCathaica multicostata. . D–F. D–F. Cathaica mengiCathaica mengi.. FIGURE 16.FIGURE 16. Natural habitats of Natural habitats of Cathaica sculptilisCathaica sculptilis sp. nov. sp. nov. White arrow in White arrow in CC shows the shows the Pseudiberus shanheicusPseudiberus shanheicus Zhang, Liu, Feng & Zhang, 2024 living in the same habitat. Zhang, Liu, Feng & Zhang, 2024 living in the same habitat. WANG ET AL.64 FIGURE 17. FIGURE 17. Natural habitats of Natural habitats of Cathaica zhangcunxiangi Cathaica zhangcunxiangi sp. nov.sp. nov. FIGURE 18.FIGURE 18. Natural habitats of Natural habitats of Cathaica wangjiaxuni Cathaica wangjiaxuni sp. nov.sp. nov. by Zhang & Wade (2023), but its shell morphology is markedly different. Cathaica sp2 shows more strongly developed ribs, a conspicuously expanded lip, no colour bands, and an overall grayish-white appearance. Because reliable molecular data for C. fasciola are not yet available, the phylogenetic relationship between C. sp2 and C. fasciola remains unresolved. Acknowledgements Thanks go to Yu-Xuan Huan [郇宇轩] (China Agricultural University) for providing materials for this study and Ms Lisa Angela Orcutt (Beijing) for language polishing. We are grateful to Dr Barna Páll-Gergely (HUN-REN Centre for Agricultural Research) and Prof. Min Wu (Nanjing University) for their constructive comments and suggestions on the manuscript. References Adams, H. 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(HBUMM10088): CTGCTCATGATTTATTTTAATAGCCGCAGTACCCTGACTGTGCTAAGGTAGCATAATCAA TTGGCTCATGATTGGAGTCTCGTATGAAAGAATTCATGGGGGTTGGCTGTTTCATATTA ATATTATTAAATTACTTATTAAGTGAAAATACTTAAAAATAAAAAATAGACGAGAAGACCCT AGAAATTTTAATTTAATTATACCTTTTTGTTGGGGCGACAAAGTAGCAAATAACCTACTTA AGTTTTACTTGAATATTATATTATGAATGAATAAATTACTCTAGGGATAACAGCATAATATTT AAAAGTTTGTGACCTCGATGTTGGATTAGGAAAACTATACCTAGAAGGTTAAT 67Three new species of Cathaica from northern China 中国中部太行山脉岩栖性华蜗牛属三新种 （腹足纲：柄眼目：坚螺科） 王志遥1,#,* 张权瑀2,# 何岳铭3 陈 辉4 冯世旸5 1中国农业大学植物保护学院 北京 100091 中国 2河南师范大学音乐舞蹈学院 新乡 453007 中国 3天津农业大学工程技术学院 天津 300384 中国 4南昌大学生命科学学院 南昌 330031 中国 5中国科学院成都生物研究所 成都 610000 中国 （#共同第一作者; *通讯作者） 摘 要 华蜗牛属 Cathaica Möllendorff, 1884 的物种广泛分布于中国温带山地地区，它们表现出 高度的多样性，并演化出多种形态以适应不同的环境。其中一些物种具有特化的外壳形态以 适应特殊的环境，并且在其分布区及周边区域未发现形态相似的物种。例如，多肋华蜗牛 C. multicostata Zhang, 2023 和孟氏华蜗牛 C. mengi Yen, 1939，具有粗壮的肋和明显突出的龙 骨。在本研究中，我们基于采自太行山区的标本，描述了华蜗牛属的三个新物种：雕刻华蜗 牛C. sculptilis Wang, Chen, He & Zhang, sp. nov.、张氏华蜗牛 C. zhangcunxiangi Wang, Chen, He & Zhang, sp. nov. 和王氏华蜗牛 C. wangjiaxuni Wang, Chen, He & Zhang, sp. nov.。这些新 种在外壳上相似，都具有比较低矮的螺塔和突出的龙骨，但生殖器解剖与分子系统学的分析 均支持这些新物种的独立性。同时为了更好地理解这些形态相似的物种之间的亲缘关系，我 们还采集了周边地区的华蜗牛标本，对它们进行了形态学和分子系统学的研究，并对孟氏华 蜗牛 C. mengi 进行了形态重新描述和生殖器补充描述。 关键词：比较形态学，分子系统发育，华北地区，新物种，岩石生境 WANG ET AL.68