Zoosyst. Evol. 100 (3) 2024, 1075-1084 | DOI 10.3897/zse.100.130661 eee BERLIN A new species of 7ropidauchenia Lindholm, 1924 (Gastropoda, Stylommatophora, Clausiliidae, Garnieriinae) from Guangdong, China Zhong-Guang Chen’, Zhe-Yu Chen’, Ran-Xi Lin?, Yu-Ting Dai!, Xiao-Ping Wu’, Jiao Jiang*, Shan Ouyang? 1 School of Life Sciences, Nanchang University, Nanchang, Jiangxi 330031, China 2 Parasitology research group, Department of Veterinary Biosciences, Faculty of Science, The University of Melbourne, Parkville VIC 3010, Australia 3 State Key Laboratory for Conservation and Utilization of Subtropical Agro-Bioresources, Guangdong Laboratory for Lingnan Modern Agriculture, College of Agriculture, College of Life Sciences, South China Agricultural University, Guangzhou 510642, China 4 Zhejiang Museum of Natural History, Hangzhou, Zhejiang 310012, China https://zoobank. org/BBA9A2D 1-3 E5 D-4F 42-9BA 3-456957BA0706 Corresponding authors: Jiao Jiang (149152414@qq.com); Shan Ouyang (ouys1963@qq.com) Academic editor: Frank Kohler # Received 27 June 2024 # Accepted 17 July 2024 Published 1 August 2024 Abstract A molecular phylogenetic study was conducted on genus TJropidauchenia, based on CO/ and /6S 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. Key Words Door snails, karst landscape, phylogeny, taxonomy Introduction Clausiliidae Gray, 1855 is a group of small to large-sized land snails with nearly global distribution (Nordsieck 2007a). A total of three subfamilies of it have been re- corded in East Asia: Phaedusinae Wagner, 1922, Garni- eriinae Boettger, 1926 and Synprosphyminae Nordsieck, 2007 (Nordsieck 2007a, 2007b, 2012a, 2012b, 2012c; Uit de Weerd et al. 2023). Among the three subfamily, Garn- ieriinae has a relatively low species diversity and the nar- rowest distribution. It is relatively narrowly distributed from Myanmar to southern China and consists of seven genera, three of which are recorded in China: Garnieria Bourguignat, 1877, Grandinenia Minato & Chen, 1984 and Tropidauchenia Lindholm, 1924 (Nordsieck 2012a, 2012b, 2012c). The genus 7ropidauchenia is a group of medium to large-sized door snails distributed from south- ern China to central Vietnam (Nordsieck 2007a, 2007b, 2012a, 2012b, 2012c; Grego and Szekeres 2017; Qiu 2021) and defined by a furrowed neck, projected and un- attached, so-called apostrophic peristome, a lunella-type lunellar region, and the inferior lamella near to or fused with superior lamella (Nordsieck 2007a, b, 2012a). Cur- rently, it consists of 18 species, 12 of which are recorded in China: 7. orientalis (Mabille, 1887), 7: dorri (Bavay & Dautzenberg, 1899), 7? hitomiae Nordsieck, 2007, T. lucida Nordsieck, 2007, T. nakaharai Nordsieck, 2007, T! napoensis Nordsieck, 2007, 7’ ootanii Nordsieck, 2007, T. mengyuanensis Chen, Tian & Fan, 2016, T. sulcicollis Grego & Szekeres, 2017, 7? yanghaoi Grego Copyright Chen, Z.-G. et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 1076 & Szekeres, 2017, T. danjuan Qiu, 2021 and 7: parasul- cicollis Qiu, 2021 (Nordsieck 2007b, 2012c; Chen, Tian and Fan 2016; Grego and Szekeres 2017; Qiu 2021; Lin and Lin 2022). These species are mainly distributed in the west Guangxi, with only three species being recorded in Yunnan and Guangdong. Guangdong Is situated in southern China and adjacent to Guangxi. In contrast to Guangxi, which is renowned for its karst landscape and high diversity of land snails, Guang- dong is relatively understudied and lacks an understanding of its land snail fauna. Only one recently described species of Tropidauchenia, namely T: yanghaoi, has been recorded in Guangdong (Grego and Szekeres 2017). The distribu- tion of this species is significantly far from that of all its congeners, suggesting the potential existence of additional yet-to-be-described species. Based on the land snails’ sur- vey conducted in April to May 2024, it has been discovered that the Tropidauchenia specimens with a ribbed shell from Luoding of Guangdong represent an undescribed species. In this study, we conducted the molecular phylogenetic analysis of three subfamilies of Clausiliidae distributed in China based on partial CO/ and /6S sequences, and de- scribed a new species of 7Zropidauchenia. The discovery of the new taxon has expanded the distribution range of Tropidauchenia and confirmed the existence of yet-to-be- described species in the distribution gap. Materials and methods Samples were collected from Sichuan, Chongqing, Yun- nan, Guangxi and Guangdong, China in 2023-2024. Living specimens were initially frozen at -20 °C for 12 hours and subsequently thawed at room temperature for 12 hours to extract the soft parts. The soft parts were then fixed in 70% ethanol. Empty shells were cleaned, dried, and preserved at 4 °C. All specimens were deposited in the School of Life Sciences, Nanchang University (Nanchang, Jiangxi, China) and the mollusc collection of Museum of Hebei University (Hebei, China). Photographs were taken by Sony® Alpha a6500 Digital Camer and edited in Adobe Photoshop CC 2015 (Adobe, San Jose, US). Maps were made in ArcGIS Pro (Esri, Redlands, US). Genomic DNA was extracted from foot tissues pre- served in 70% ethanol using a TIANamp Marine Animals DNA Kit (Tiangen Biotech, China). The quality and con- centration of the DNA were checked on 1% agarose gel electrophoresis and NanoDrop 2000 (Thermo Scientific, USA). Partial cytochrome c oxidase subunit 1 (CO/) and partial 16S ribosomal RNA (/6S) gene segments were amplified and sequenced for molecular phylogenetic analyses. Polymerase chain reaction (PCR) systems, con- ditions and primer pairs are listed in Table 1. Sequences were aligned using MEGA v. 6.0 (Tamura et al. 2013) and checked manually. The accession numbers of newly ob- tained sequences and other species are given in Table 2. Phylogenies were reconstructed by the dataset combined two genes using maximum likelihood (ML) and Bayesian zse.pensoft.net Chen, Z.-G. et al.: A new species of Tropidauchenia inference (BI). Seventeen species of three subfamilies dis- tributed in east Asia were included. Three species of sub- family Alopiinae Wagner, 1913 were used as outgroups for rooting the tree. ML analyses were performed in IQ- TREE v. 1.6.12 (Minh et al. 2013) using Ultrafast fast bootstrap approach (Minh et al. 2013) with 10000 reiterations. The most appropriate model of sequence evolution (GTR+I+G) was selected under PartitionFinder2 v. 1.1 (Lanfear et al. 2017). Bayesian inference (BI) was conducted in MrBayes v. 3.2.6 (Ronquist et al. 2012). The most appropriate mod- el of sequence evolution (GTR+I+G) was selected under ModelFinder (Kalyaanamoorthy et al. 2017). Four simul- taneous runs with four independent Markov Chain Monte Carlo (MCMC) were implemented for 10 million genera- tions, and trees were sampled every 1000 generations with a burn-in of 25%. The convergence was checked with the average standard deviation of split frequencies <0.01 and the potential scale reduction factor (PSRF) ~1. Trees were visualised in FigTree v.1.4.3. Abbreviations. NCU_XPWU Laboratory of Xiao- Ping Wu, Nanchang University (Nanchang, Jiangxi, Chi- na); HBUMM mollusc collection of Museum of Hebei University, Hebei, China; cp clausilium plate; il inferior lamella; lu lunella; pp principal plica; se subcolumellar lamella; sl superior lamella; sp spiral lamella; At atrium; BC bursa copulatrix; BCD bursa copulatrix duct; D di- verticulum; Ep epiphallus; P penis; PR penial retractor muscle; V vagina; VD vas deferens. Results Phylogenetic analyses A dataset consisting of 44 CO/ and 47 /6S sequences from 17 species, along with three outgroup taxa, was employed for phylogenetic analyses (Table 2). The CO/ sequence of 7. yanghaoi was unable to be amplified by the several uni- versal primers employed. The aligned lengths of CO/ and 16S genes were 669 and 492 nucleotides. Within these sequences, 330 and 266 were revealed as variable sites, while 325 and 255 were designated as parsimony infor- mative sites. Phylogenetic analyses generated ML and BI trees with congruent topologies (Fig. 1). The results sup- ported the monophyly of three subfamilies of Clausiliidae Gray, 1855 distributed in China. It demonstrated a phylo- genetic relationship of Garnieriinae + (Synprosphyminae + Phaedusinae). The genus Excussispira Lindholm, 1925, which has an undetermined systematic position, has been confirmed as belonging to Synprosphyminae. The mono- phyly of Garnieriinae is supported but with relatively low nodal support (bootstrap support = 66, posterior proba- bility = 0.70). Genus Tropidauchenia form a monophyly and further clustered into two clades, consisted of the spe- cies from Guangdong and Guangxi respectively. Tropid- auchenia jiangjilini sp. nov. was sistered with 7. yanghaoi (bootstrap support =99, posterior probability = 1) and the genetic distances of /6S sequences between them is 8.6%. Zoosyst. Evol. 100 (3) 2024, 1075-1084 Table 1. Primer pairs and PCR conditions used in the analyses of the CO/ and 16S rRNA genes. Genes CO! 16S Primer pairs LCO1490: GGTCAACAAATCATAAAGATATTGG HCO2198: TAAACTTCAGGGTGACCAAAAAATCA (Vazyme, Nanjing, China), 1 pl template 50 °C: 60s, 72 °C: 1 min, DNA, 1 ul of each pair of primers, 16SA: CGGCCGCCTGTTTATCAAAAACAT 16SB: GGAGCTCCGGTTTGAACTCAGATC 9.5 pl ddH20 9.5 pl ddH20 Table 2. GenBank accession numbers of the sequences for this study. Species Tropidauchenia jiangjilini sp. nov. T. danjuan T. parasulcicollis T. cf. lucida T. yanghaol T. orientalis Grandinenia mirifica G. gastrum G. fuchsi Excussispira fargesiana E. lunatica Synprosphyma suilla S. basilissa Miraphaedusa takagli Serriphaedusa zhengpingi Paraformosana indurata Cirrophaedusa plicilabris Agathylla goldi Alopia mariae lsabellaria praestans Locality Pingtang, Luoding, Guangdong, China, 22°43'26'N, 1t1°44-56°E Maan, Jiangzhou, Chongzuo, Guangxi, China (type locality), 22°27'16'N, 107°19'56'E Baoan, Jiangzhou, Chongzuo, Guangxi, China, 22°17'50'N, 107°21'24'E Daxin, Chongzuo, China (type locality), 22°42'41'N, 107°11'55'E Chongzuo, Guangxi, China, 22°8'15"N, 106°46'41"E Huaiji, Zhaoging, Guangxi, China (type locality), 23°55'13'N,; 112°9'59'E Chongzuo, Guangxi, China, 22°16'29"N, 107°4'14"E Lianggu, Qintang, Guigang, Guangxi, China (type locality), 23°19'1'N, 109°14'34"E Lianggu, Qintang, Guigang, Guangxi, China (type locality), 23°18'51"N, 109°15'49"E Guilin, Guangxi, China, 25°18'35'N, 110°16'19"E Sichuan, China (type locality), 32°5'2"N, 108°3'12"E Chengkou, Chongaing, China (type locality), 31°57'34'N, 108°38'31"E Honghe, Yunnan, China, 29°56'39"N, 103°02'35'E Chengdu, Sichuan, China, 30°55'26"N, 103°29'23°E Guilin, Guangxi, China (type locality), 25°18'35'N, 110°16'19"E Yaan, Sichuan, China (type locality), 29°37'28'N, 10275318 'F Wushan, Chongqing, China, 31°19'2"N, 109°47'53°E Beibei, Chongqing, China (type locality), 30°1'47'N, 106°37'24"E Europe Europe Europe Reaction systems 12.5 ul 2x Taq Plus Master Mix II 12.5 ul 2x Taq Plus Master Mix II (Vazyme, Nanjing, China), 1 pl template 50 °C: 60s, 72 °C: 1 min, DNA, 1 ul of each pair of primers, col PP945861 PP945862 PP945863 PP945864 PP945865 PP945866 PP945851 PP945852 PP945853 PP945854 PP945855 PP945856 PP945857 PP945858 PP945859 PP945859 PP473359 PP473360 PP473344 PP473345 PP473346 PP473348 PP473349 PP473350 PP473351 PP473352 PP473353 PP945871 PP945872 PP945873 PP945867 PP945868 PP945869 PP945870 PP945876 PP945874 PP945875 PP945877 PP945878 PP945879 PP945880 PP945881 PP945882 PP945883 KC756080 JQ911821 AY425575 Cycling conditions 94 °C: 2 min; 94 °C: 10s, 94 °C: 2 min; 94 °C: 10s, 16S PP956568 PP956569 PP956570 PP956571 ProDoor2 PP956573 PP956558 PP956559 PP956560 PP956561 PP956562 PP956563 PP956564 PP956565 PP956566 PP956567 PP472620 PP472621 PP472622 PP472591 PP472592 PP472576 PP472577 PP472578 PP472580 PP472581 PP472582 PP472583 PP472584 PP472585 PP956578 PP936579 PP956580 PP956574 PP956575 PR956526 PP956577 PP956583 PP956581 PP956582 PP956584 PP956585 PP956586 PP956587 PP956588 PP956589 PP956590 KF601271 1077 Reference 35 cycles; 72 °C: 10 min 35 cycles; 72 °C: 10 min References This study This study This study This study This study This study This study This study This study This study This study This study This study This study This study This study Unpublished Unpublished Unpublished Unpublished Unpublished Unpublished Unpublished Unpublished Unpublished Unpublished Unpublished Unpublished Unpublished Unpublished This study This study This study This study This study This study This study This study This study This study This study This study This study This study This study This study (2004) Folmer et al. 1994 Pall-Gergely et al. 2019 Fehér et al. (201 3b), Parmakelis et al. (2013) Fehér et al. (2013a) Uit de Weerd et al. zse.pensoft.net 1078 95/0.99 98/1 98/1 92/0.99 99/1 Chen, Z.-G. et al.: A new species of Tropidauchenia Tropidauchenia danjuan Tropidauchenia danjuan Tropidauchenia danjuan Tropidauchenia danjuan Tropidauchenia parasulcicollis Tropidauchenia parasulcicollis Tropidauchenia parasulcicollis Tropidauchenia parasulcicollis Tropidauchenia orientalis Tropidauchenia orientalis Tropidauchenia cf. lucida Tropidauchenia cf. lucida Tropidauchenia jiangjilini sp. nov. Tropidauchenia jiangjilini sp. nov. Tropidauchenia jiangjilini sp. nov. Tropidauchenia jiangjilini sp. nov. Tropidauchenia jiangjilini sp. nov. Tropidauchenia jiangjilini sp. nov. Garnieriinae Tropidauchenia yanghaoi Tropidauchenia yanghaoi Tropidauchenia yanghaoi 66/0.70 100/1 100/1 100/1 100/1 84/0.73 Bee 74/0.89 93/0.99 96/1 Grandinenia gastrum Grandinenia gastrum Grandinenia gastrum Grandinenia mirifica Grandinenia mirifica Grandinenia mirifica Grandinenia fuchsi Grandinenia fuchsi Grandinenia fuchsi Excussispira lunatica Excussispira lunatica Excussispira lunatica Excussispira lunatica Excussispira fargesiana Excussispira fargesiana Excussispira fargesiana Synprosphyma basilissa Synprosphyma basilissa Synprosphyma suilla Miraphaedusa takagii Miraphaedusa takagii Serriphaedusa zhengpingi Serriphaedusa zhengpingi Paraformosana indurata Paraformosana indurata 100/1 Synprosphyminae Phaedusinae 97/0.95 Alopia mariae 100/1 Isabellaria praestans Agathylla goldi 0.1 Cirrophaedusa plicilabris outgroups Figure 1. Maximum likelihood tree and Bayesian inference tree inferred from CO/ and /6S gene sequences. Bootstrap supports/ posterior probabilities are shown on the left/right of nodes. Taxonomy Family Clausiliidae Gray, 1855 Subfamily Garnieriinae Boettger, 1926 Genus Tropidauchenia Lindholm, 1924 Type species. Clausilia bavayi Lindholm, 1924, by orig- inal designation. Tropidauchenia jiangjilini Z.-G. Chen, Z.-Y. Chen & R.-X. Lin, sp. nov. https://zoobank.org/0421ED62-C 136-4882-AF91-544D7B15821C Figs 2A, 3, 4A, B, SB-D Type material. Holotype. 24 NCU_XPWU_YG01, Ju- longdong Scenic Spot [28 Jia MX], Pingtang Town [243344], Luoding City [272 1], Guangdong Province UA], China, 22°43'26"N, 111°44'56"E, leg. Ji-Lin Jiang, May 2024. zse.pensoft.net Paratypes. 11 specimens. 24 NCU_XPWU_YG02- 09, HBUMN 10073-—74, other information same as holo- type; 24 NCU _XPWU_YGI1O0, Longji village [4A], Luoding City [2’7€TH], Guangdong Province, China, 22°44'40"N, 111°48'01"E, leg. Ji-Lin Jiang & Ran-Xi Lin, May 2024. Diagnosis. Shell entire (vs. decollated or fragile to almost decollated in 7. bavayi, T: dorri, T: dong- giaensis Nordsieck, 2002, 7: giardi and T. proctosto- ma (Mabille, 1889)), slender-fusiform (vs. broader in T. danjuan, T: hitomiae, T: lucida, T: mengyuanensis, T. messageri (Bavay & Dautzenberg, 1899), 7) nakaharai, T. napoensis, T: orientalis, T. palatalis, T. parasulcicollis and 7: sulcicollis), fragile; teleoconch with very thin and dense ribs (vs. smooth in 7? yanghaoi; ribs stronger but sparser in 7’ danjuan, T. messageri, T. orientalis, T: oot- anii, T. parasulcicollis and T: sulcicollis; ribs weaker and sparser in 7’ hitomiae, T. lucida, T. mengyuanensis, T. na- kaharai, T: napoensis and T: palatalis); aperture colored, lower part reddish-brown; penial pilasters transversal, relatively sparse, irregular zigzag. Zoosyst. Evol. 100 (3) 2024, 1075-1084 1079 Figure 2. Tropidauchenia jiangjilini sp. nov. and its most similar congener. A. Tropidauchenia jiangjilini sp. nov., holotype (24 __ NCU_XPWU_YG01); B. 7. yanghaoi. Description. Shell (n=12). Entire, with 10.5—-11.5 whorls, hardly decollated, slender-fusiform, thin, fragile, semitranslucent, light brown, with indistinct darkish-red ribbon beneath the suture; body whorl in front of lunella darker; apical part conical to slowly attenuated. Suture shallow. Protoconch smooth with 2.5—3.0 whorls. Ribs on the teleoconch very thin and dense, extending across the whole whorl, rather evenly distributed and narrowly spaced; on the neck riblets white, broader, stronger, more widely spaced and undulate. Aperture vastly extended, oval. Peristome expanded, thickened, slightly reflected, lower part reddish-brown. Only the superior lamella vis- ible through aperture. Superior lamella bent, fused with inferior lamella but separated with spiral lamell. The end of superior lamella longer than that of spiral lamella. Subcolumellar lamella invisible in oblique view, strong, bent, its end shorter than the end of superior lamella. Lunella invisible in oblique view, vertical. Principal plica short, initiates ventrolaterally and extending laterally, not reaching peristome. Clausilium plate invisible in oblique view, semitranslucent; overall slender; stalk thin; plate relatively broad. Genitalia (n=4). Atrium short and relatively narrow. Penis relatively slender, almost cylindrical. Penial pi- lasters transversal, relatively sparse, irregular zigzag. Penial caecum strong, long, tubular. Epiphallus slender, zse.pensoft.net 1080 Chen, Z.-G. et al.: A new species of Tropidauchenia Figure 3. Detailed shell morphology of Tropidauchenia jiangjilini sp. nov. Abbreviations: cp clausilium plate; il inferior lamella; lu lunella; pp principal plica; sc subcolumellar lamella; sl superior lamella; sp spiral lamella. shorter and thinner than penis. Penial retractor relatively thick and long, inserted at the middle part of penis. Vas deferens relatively slender and short. Vagina relatively slender, cylindrical. Diverticulum slender, unexpanded. Bursa copulatrix duct slender and long. Bursa copulatrix small, oval. Measurements. Holotype: shell height 27.7 mm, width 5.0 mm; aperture height 5.6 mm, width 5.3 mm. Paratypes: shell height 23.6—28.8 mm, width 4.6— 5.2 mm; aperture height 4.9-5.9 mm, width 4.6— 5.2mm (n= 11). zse.pensoft.net Etymology. The species is named after Ji-Lin Jiang who first discovered the new species and assisted with the field survey. Vernacular name. {L[042e 8 (Pinyin: jiang shi san guan luo). Distribution and ecology. 7ropidauchenia jiangjili- ni sp. nov. is found from two adjacent hills in Luoding (Figs 6, 7). Other surrounding hills are also potential distributions but require further survey. It inhabits the vertical limestone cliff together with Gyliotrachela sp. (Figs SB—-D). Zoosyst. Evol. 100 (3) 2024, 1075-1084 1081 At A C At : = Ep V Ep P vD = PR ip PC ) Bc «BCD a RR OFS D D 2mm 2mm 1mm 1mm Figure 4. Genital anatomy. A. 7ropidauchenia jiangjilini sp. nov., B. Penial pilasters of Tropidauchenia jiangjilini sp. nov., C. 7. yanghaoi, D. Penial pilasters of 7’ yanghaoi. Abbreviations: At atrium; BC bursa copulatrix; BCD bursa copulatrix duct; D diverticulum; Ep epiphallus; P penis; PR penial retractor muscle; V vagina; VD vas deferens. Me i Big a viehsy 3 ae bre “’ i eg Po Mee ae fe a, uf ' ee wh ok wd g ORE ais mr me Figure 5. Living specimens. A. 7? yanghaoi;, B—D. Tropidauchenia jiangjilini sp. nov. zse.pensoft.net 1082 Chen, Z.-G. et al.: A new species of Tropidauchenia 102°E 104°E 106°E 108°E 110°E 112°E 26°N 26°N fo %y Guizhou ~ - \ # a 24°N 24°N Guang 22°N 22°N ‘ Lj = se a ed Fae be PRE . Vietnam 20°N 20°N 102°E 104°E 106°E 108°E 110°E 112°E Figure 6. Distribution of Zropidauchenia in China. Red star. Tropidauchenia jiangjilini sp. nov., orange star. 7: yanghaoi, red square. T. mengyuanensis, orange square. 7: ootanii, purple dot. 7’ napoensis, black dot. 7: nakaharai, green dot. T dorri, yellow dot. T: hitomiae, orange dot. T: lucida, brown dot. 7: sulcicollis, pink dot. 7! parasulcicollis, blue dot. 7: danjuan, grey dot. 7: orientalis. Pag ( aoe) A le Figure 7. Type locality of Tropidauchenia jiangjilini sp. nov. A. overall environment; B. sampling locality. Discussion microenvironmental preferences. Phaedusinae exhibits the widest adaptability to the environment, with a pref- It is possible that different environmental choices may _ erence for well-ventilated and relatively humid habitats. promote the differentiation of the Clausiliidae in East Synprosphyminae inhabits the extremely humid environ- Asia. Although the three subfamilies are to some ex- ments at relatively high altitudes. It is highly dependent tent sympatric in the region, they display different on the extremely humid and low temperature, with the zse.pensoft.net Zoosyst. Evol. 100 (3) 2024, 1075-1084 majority of species being found in the seepage points of groundwater or even by the river. Some species of it have evolved specialized respiration channels to facilitate survival in frequently inundated habitats. Garnieriinae 1s a strict rock-dwelling group and rarely descends to the ground except for spawning and dormancy. Within the subfamily, the genera Grandinenia and Tropidauchenia may demonstrate differential environmental selectivity. It has been observed that 7ropidauchenia is predominantly found on the shaded, humid rock walls of dense forests, whereas Grandinenia is more commonly encountered in the more exposed and drier environment. The placement of the new species within Tropidauche- nia is supported by both morphology (inferior lamella fused with superior lamella) and molecular phylogeny. The absence of description of the genitalia, as well as the dearth of illustrations of lamellae and genitalia in the most original descriptions of Tropidauchenia species, precludes the possibility of a detailed comparison of the new species with most other congeners for these two characters. How- ever, the distinctive shell morphology can distinguish it from all its congeners. Tropidauchenia jiangjilini sp. nov. can be easily distinguished from 7 bavayi, T: dorri, T: dong- giaensis, T. giardi and T. proctostoma by the protoconch solid and preserve (vs. decollated or fragile to almost de- collated) and the thinner shell. Tropidauchenia jiangjilini Sp. nov. is similar to 7’ danjuan, T: hitomiae, T: lucida, T. mengyuanensis, T. messageri, T: nakaharai, T? napoensis, 7. orientalis, T: palatalis, T. parasulcicollis and T. sulci- collis in its entire shell, but differs from them by the more slender shell. It is further distinguished from 7? danjuan, T. messageri, T. orientalis, T: parasulcicollis and T: sulci- collis by the weaker but denser ribs on teleoconch, from T. hitomiae, T. lucida, T: mengyuanensis, T. nakaharai, T. napoensis and T: palatalis by the stronger and denser ribs on teleoconch. Tropidauchenia jiangjilini sp. nov. 1s relatively similar to 7. ootanii by the similar slender and ribbed shell, but differs by the weaker but denser ribs on teleoconch, the ribs’ uniform thickness and height (vs. ribs beneath the suture stronger) and the thicker and darker colored shell. The unique and distant distribution can also distinguish the new species from all the above congeners (Guangdong vs. Guangxi or Yunnan). 7ropidauchenia ji- angjilini sp. nov. is most similar with 7. yanghaoi by the similar slender shell and the near distribution, but differs by the ribbed teleoconch (vs. smooth), the stronger and denser ribs on neck, the thinner diverticulum, the stronger penial caecum and the thinner and sparser penial pilasters. The new species and 7’? yanghaoi presence of a very long and strong penial caecum, while the known penial caecum in Grandinenia species are very small. For a long time, Tropidauchenia and Grandinenia can only distinguished by the lamella, the difference in penial caecum may be a diagnostic character of the two genera. However, due to the lack of information on the genitalia of most Trop- idauchenia and Grandinenia species, the speculation re- main further study and verification. The validity of Trop- idauchenia jiangjilini sp. nov. was also supported by the 1083 molecular phylogeny. It was sistered with 7’ yanghaoi and forms a monophyly with the species from Guangx1. The discovery of new species indicates that the species diversity of Garnieriinae in western Guangdong remains to be fully explored. As a group of rock-dwelling land snails, the species diversity of Garnieriinae is particularly high in the karst region. In comparison to the diverse spe- cies found in Guangxi, the known species of Garnieriinae in the karst region of Guangdong remain relatively few. Further comprehensive surveys in the future may reveal additional yet-to-be-described species. Acknowledgments We thank Ji-Lin Jiang (Zhaoqing), Meng-Hua Li (Sich- uan Agriculture University), Chen-Yu Fei (Guangzhou) and Shi-Yang Feng (Sichuan Agriculture University) for assistance in collecting specimens, Frank Kohler for as- sistance in processing the manuscript, Chih-Wei Huang for reviewing the manuscript. This study was support- ed by the National Natural Science Foundation of Chi- na under Grant No.32360132, No.31772412, research grants from the Malacological Society of London, the Melbourne Research Scholarship for ZYC, the research project of Zhejiang Natural History Museum under Grant No.2024001 and the Biodiversity Monitoring Project of X1x1 National Wetland Park of Hangzhou. References Chen YX, Tian M, Fan B (2016) Terrestrial molluscs in Yunnan. Science Press, China, 259 pp. Fehér Z, Németh L, Nicoara A, Szekeres M (2013a) Molecular phylog- eny of the land snail genus A/opia (Gastropoda: Clausiliidae) reveals multiple inversions of chirality. 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