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Research Article
A new species of Tropidauchenia Lindholm, 1924 (Gastropoda, Stylommatophora, Clausiliidae, Garnieriinae) from Guangdong, China
expand article infoZhong-Guang Chen, Zhe-Yu Chen§, Ran-Xi Lin|, Yu-Ting Dai, Xiao-Ping Wu, Jiao Jiang, Shan Ouyang
‡ Nanchang University, Nanchang, China
§ The University of Melbourne, Parkville, Australia
| South China Agricultural University, Guangzhou, China
¶ Zhejiang Museum of Natural History, Hangzhou, China
Open Access

Abstract

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

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 recorded in East Asia: Phaedusinae Wagner, 1922, Garnieriinae Boettger, 1926 and Synprosphyminae Nordsieck, 2007 (Nordsieck 2007a, 2007b, 2012a, 2012b, 2012c; Uit de Weerd et al. 2023). Among the three subfamily, Garnieriinae has a relatively low species diversity and the narrowest 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 Tropidauchenia is a group of medium to large-sized door snails distributed from southern China to central Vietnam (Nordsieck 2007a, 2007b, 2012a, 2012b, 2012c; Grego and Szekeres 2017; Qiu 2021) and defined by a furrowed neck, projected and unattached, so-called apostrophic peristome, a lunella-type lunellar region, and the inferior lamella near to or fused with superior lamella (Nordsieck 2007a, b, 2012a). Currently, it consists of 18 species, 12 of which are recorded in China: T. orientalis (Mabille, 1887), T. dorri (Bavay & Dautzenberg, 1899), T. hitomiae Nordsieck, 2007, T. lucida Nordsieck, 2007, T. nakaharai Nordsieck, 2007, T. napoensis Nordsieck, 2007, T. ootanii Nordsieck, 2007, T. mengyuanensis Chen, Tian & Fan, 2016, T. sulcicollis Grego & Szekeres, 2017, T. yanghaoi Grego & Szekeres, 2017, T. danjuan Qiu, 2021 and T. parasulcicollis 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, Guangdong 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 distribution 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’ survey 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 COI and 16S sequences, and described a new species of Tropidauchenia. 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, Yunnan, 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 preserved in 70% ethanol using a TIANamp Marine Animals DNA Kit (Tiangen Biotech, China). The quality and concentration of the DNA were checked on 1% agarose gel electrophoresis and NanoDrop 2000 (Thermo Scientific, USA). Partial cytochrome c oxidase subunit 1 (COI) and partial 16S ribosomal RNA (16S) gene segments were amplified and sequenced for molecular phylogenetic analyses. Polymerase chain reaction (PCR) systems, conditions 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 obtained sequences and other species are given in Table 2.

Table 1.

Primer pairs and PCR conditions used in the analyses of the COI and 16S rRNA genes.

Genes Primer pairs Reaction systems Cycling conditions Reference
COI LCO1490: GGTCAACAAATCATAAAGATATTGG HCO2198: TAAACTTCAGGGTGACCAAAAAATCA 12.5 μl 2× Taq Plus Master Mix II (Vazyme, Nanjing, China), 1 μl template DNA, 1 μl of each pair of primers, 9.5 μl ddH2O 94 °C: 2 min; 94 °C: 10s, 50 °C: 60s, 72 °C: 1 min, 35 cycles; 72 °C: 10 min Folmer et al. 1994
16S 16SA: CGGCCGCCTGTTTATCAAAAACAT 16SB: GGAGCTCCGGTTTGAACTCAGATC 12.5 μl 2× Taq Plus Master Mix II (Vazyme, Nanjing, China), 1 μl template DNA, 1 μl of each pair of primers, 9.5 μl ddH2O 94 °C: 2 min; 94 °C: 10s, 50 °C: 60s, 72 °C: 1 min, 35 cycles; 72 °C: 10 min Páll-Gergely et al. 2019
Table 2.

GenBank accession numbers of the sequences for this study.

Species Locality CO1 16S References
Tropidauchenia jiangjilini sp. nov. Pingtang, Luoding, Guangdong, China, 22°43'26"N, 111°44'56"E PP945861 PP956568 This study
PP945862 PP956569 This study
PP945863 PP956570 This study
PP945864 PP956571 This study
PP945865 PP956572 This study
PP945866 PP956573 This study
T. danjuan Maan, Jiangzhou, Chongzuo, Guangxi, China (type locality), 22°27'16"N, 107°19'56"E PP945851 PP956558 This study
PP945852 PP956559 This study
Baoan, Jiangzhou, Chongzuo, Guangxi, China, 22°17'50"N, 107°21'24"E PP945853 PP956560 This study
PP945854 PP956561 This study
T. parasulcicollis Daxin, Chongzuo, China (type locality), 22°42'41"N, 107°11'55"E PP945855 PP956562 This study
PP945856 PP956563 This study
PP945857 PP956564 This study
PP945858 PP956565 This study
T. cf. lucida Chongzuo, Guangxi, China, 22°8'15"N, 106°46'41"E PP945859 PP956566 This study
PP945859 PP956567 This study
yanghaoi Huaiji, Zhaoqing, Guangxi, China (type locality), 23°55'18"N, 112°9'59"E PP472620 Unpublished
PP472621 Unpublished
PP472622 Unpublished
T. orientalis Chongzuo, Guangxi, China, 22°16'29"N, 107°4'14"E PP473359 PP472591 Unpublished
PP473360 PP472592 Unpublished
Grandinenia mirifica Lianggu, Qintang, Guigang, Guangxi, China (type locality), 23°19'1"N, 109°14'34"E PP473344 PP472576 Unpublished
PP473345 PP472577 Unpublished
PP473346 PP472578 Unpublished
G. gastrum Lianggu, Qintang, Guigang, Guangxi, China (type locality), 23°18'51"N, 109°15'49"E PP473348 PP472580 Unpublished
PP473349 PP472581 Unpublished
PP473350 PP472582 Unpublished
G. fuchsi Guilin, Guangxi, China, 25°18'35"N, 110°16'19"E PP473351 PP472583 Unpublished
PP473352 PP472584 Unpublished
PP473353 PP472585 Unpublished
Excussispira fargesiana Sichuan, China (type locality), 32°5'2"N, 108°3'12"E PP945871 PP956578 This study
PP945872 PP956579 This study
PP945873 PP956580 This study
E. lunatica Chengkou, Chongqing, China (type locality), 31°57'34"N, 108°38'31"E PP945867 PP956574 This study
PP945868 PP956575 This study
PP945869 PP956576 This study
PP945870 PP956577
Synprosphyma suilla Honghe, Yunnan, China, 29°56'39"N, 103°02'35"E PP945876 PP956583 This study
S. basilissa Chengdu, Sichuan, China, 30°55'26"N, 103°29'23"E PP945874 PP956581 This study
PP945875 PP956582 This study
Miraphaedusa takagii Guilin, Guangxi, China (type locality), 25°18'35"N, 110°16'19"E PP945877 PP956584 This study
PP945878 PP956585 This study
Serriphaedusa zhengpingi Yaan, Sichuan, China (type locality), 29°37'28"N, 102°53'18"E PP945879 PP956586 This study
PP945880 PP956587 This study
Paraformosana indurata Wushan, Chongqing, China, 31°19'2"N, 109°47'53"E PP945881 PP956588 This study
PP945882 PP956589 This study
Cirrophaedusa plicilabris Beibei, Chongqing, China (type locality), 30°1'47"N, 106°37'24"E PP945883 PP956590 This study
Agathylla goldi Europe KC756080 KF601271 Fehér et al. (2013b), Parmakelis et al. (2013)
Alopia mariae Europe JQ911821 Fehér et al. (2013a)
Isabellaria praestans Europe AY425575 Uit de Weerd et al. (2004)

Phylogenies were reconstructed by the dataset combined two genes using maximum likelihood (ML) and Bayesian inference (BI). Seventeen species of three subfamilies distributed in east Asia were included. Three species of subfamily 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 model of sequence evolution (GTR+I+G) was selected under ModelFinder (Kalyaanamoorthy et al. 2017). Four simultaneous runs with four independent Markov Chain Monte Carlo (MCMC) were implemented for 10 million generations, 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, China); HBUMM mollusc collection of Museum of Hebei University, Hebei, China; cp clausilium plate; il inferior lamella; lu lunella; pp principal plica; sc subcolumellar lamella; sl superior lamella; sp spiral lamella; At atrium; BC bursa copulatrix; BCD bursa copulatrix duct; D diverticulum; Ep epiphallus; P penis; PR penial retractor muscle; V vagina; VD vas deferens.

Results

Phylogenetic analyses

A dataset consisting of 44 COI and 47 16S sequences from 17 species, along with three outgroup taxa, was employed for phylogenetic analyses (Table 2). The COI sequence of T. yanghaoi was unable to be amplified by the several universal primers employed. The aligned lengths of COI 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 informative sites. Phylogenetic analyses generated ML and BI trees with congruent topologies (Fig. 1). The results supported the monophyly of three subfamilies of Clausiliidae Gray, 1855 distributed in China. It demonstrated a phylogenetic relationship of Garnieriinae + (Synprosphyminae + Phaedusinae). The genus Excussispira Lindholm, 1925, which has an undetermined systematic position, has been confirmed as belonging to Synprosphyminae. The monophyly of Garnieriinae is supported but with relatively low nodal support (bootstrap support = 66, posterior probability = 0.70). Genus Tropidauchenia form a monophyly and further clustered into two clades, consisted of the species from Guangdong and Guangxi respectively. Tropidauchenia jiangjilini sp. nov. was sistered with T. yanghaoi (bootstrap support =99, posterior probability = 1) and the genetic distances of 16S sequences between them is 8.6%.

Figure 1. 

Maximum likelihood tree and Bayesian inference tree inferred from COI and 16S gene sequences. Bootstrap supports/posterior probabilities are shown on the left/right of nodes.

Taxonomy

Family Clausiliidae Gray, 1855

Subfamily Garnieriinae Boettger, 1926

Tropidauchenia Lindholm, 1924

Type species

Clausilia bavayi Lindholm, 1924, by original designation.

Tropidauchenia jiangjilini Z.-G. Chen, Z.-Y. Chen & R.-X. Lin, sp. nov.

Figs 2A, 3, 4A, B, 5B–D

Type material

Holotype. 24_NCU_XPWU_YG01, Julongdong Scenic Spot [聚龙洞风景区], Pingtang Town [苹塘镇], Luoding City [罗定市], Guangdong Province [广东省], China, 22°43'26"N, 111°44'56"E, leg. Ji-Lin Jiang, May 2024.

Paratypes. 11 specimens. 24_NCU_XPWU_YG02–09, HBUMN 10073–74, other information same as holotype; 24_NCU_XPWU_YG10, Longji village [龙吉村], Luoding City [罗定市], 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 T. bavayi, T. dorri, T. donggiaensis Nordsieck, 2002, T. giardi and T. proctostoma (Mabille, 1889)), slender-fusiform (vs. broader in T. danjuan, T. hitomiae, T. lucida, T. mengyuanensis, T. messageri (Bavay & Dautzenberg, 1899), T. nakaharai, T. napoensis, T. orientalis, T. palatalis, T. parasulcicollis and T. sulcicollis), fragile; teleoconch with very thin and dense ribs (vs. smooth in T. yanghaoi; ribs stronger but sparser in T. danjuan, T. messageri, T. orientalis, T. ootanii, T. parasulcicollis and T. sulcicollis; ribs weaker and sparser in T. hitomiae, T. lucida, T. mengyuanensis, T. nakaharai, T. napoensis and T. palatalis); aperture colored, lower part reddish-brown; penial pilasters transversal, relatively sparse, irregular zigzag.

Figure 2. 

Tropidauchenia jiangjilini sp. nov. and its most similar congener. A. Tropidauchenia jiangjilini sp. nov., holotype (24_NCU_XPWU_YG01); B. T. 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 visible 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.

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.

Genitalia (n=4). Atrium short and relatively narrow. Penis relatively slender, almost cylindrical. Penial pilasters transversal, relatively sparse, irregular zigzag. Penial caecum strong, long, tubular. Epiphallus slender, 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.

Figure 4. 

Genital anatomy. A. Tropidauchenia jiangjilini sp. nov., B. Penial pilasters of Tropidauchenia jiangjilini sp. nov., C. T. yanghaoi, D. Penial pilasters of T. 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.

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 4.6–5.2 mm (n = 11).

Figure 5. 

Living specimens. A. T. yanghaoi; B–D. Tropidauchenia jiangjilini sp. nov.

Etymology

The species is named after Ji-Lin Jiang who first discovered the new species and assisted with the field survey.

Vernacular name

江氏伞管螺 (Pinyin: jiāng shì sǎn guǎn luó).

Distribution and ecology

Tropidauchenia jiangjilini 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 5B–D).

Figure 6. 

Distribution of Tropidauchenia in China. Red star. Tropidauchenia jiangjilini sp. nov., orange star. T. yanghaoi, red square. T. mengyuanensis, orange square. T. ootanii, purple dot. T. napoensis, black dot. T. nakaharai, green dot. T. dorri, yellow dot. T. hitomiae, orange dot. T. lucida, brown dot. T. sulcicollis, pink dot. T. parasulcicollis, blue dot. T. danjuan, grey dot. T. orientalis.

Figure 7. 

Type locality of Tropidauchenia jiangjilini sp. nov. A. overall environment; B. sampling locality.

Discussion

It is possible that different environmental choices may promote the differentiation of the Clausiliidae in East Asia. Although the three subfamilies are to some extent sympatric in the region, they display different microenvironmental preferences. Phaedusinae exhibits the widest adaptability to the environment, with a preference for well-ventilated and relatively humid habitats. Synprosphyminae inhabits the extremely humid environments at relatively high altitudes. It is highly dependent on the extremely humid and low temperature, with the 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 is 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 Tropidauchenia 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 Tropidauchenia 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. However, the distinctive shell morphology can distinguish it from all its congeners. Tropidauchenia jiangjilini sp. nov. can be easily distinguished from T. bavayi, T. dorri, T. donggiaensis, T. giardi and T. proctostoma by the protoconch solid and preserve (vs. decollated or fragile to almost decollated) and the thinner shell. Tropidauchenia jiangjilini sp. nov. is similar to T. danjuan, T. hitomiae, T. lucida, T. mengyuanensis, T. messageri, T. nakaharai, T. napoensis, T. orientalis, T. palatalis, T. parasulcicollis and T. sulcicollis in its entire shell, but differs from them by the more slender shell. It is further distinguished from T. danjuan, T. messageri, T. orientalis, T. parasulcicollis and T. sulcicollis 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. is relatively similar to T. 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). Tropidauchenia jiangjilini sp. nov. is most similar with T. 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 T. 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 Tropidauchenia and Grandinenia species, the speculation remain further study and verification. The validity of Tropidauchenia jiangjilini sp. nov. was also supported by the molecular phylogeny. It was sistered with T. yanghaoi and forms a monophyly with the species from Guangxi.

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 species 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 (Sichuan Agriculture University), Chen-Yu Fei (Guangzhou) and Shi-Yang Feng (Sichuan Agriculture University) for assistance in collecting specimens, Frank Köhler for assistance in processing the manuscript, Chih-Wei Huang for reviewing the manuscript. This study was supported by the National Natural Science Foundation of China 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 Xixi National Wetland Park of Hangzhou.

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