Research Article
Research Article
A new species of freshwater snail of Fenouilia (Gastropoda, Pomatiopsidae) from northern Guangxi, China, based on morphological and DNA evidence
expand article infoHui Chen, Yue Ming He§, Chong Rui Wang§, Da Pan
‡ Nanjing Normal University, Nanjing, China
§ Hunan Fisheries Science Institute, Changsha, China
Open Access


A new species of pomatiopsid freshwater snail, Fenouilia undata Chen & He, sp. nov., is described from Guangxi, China, based on morphological and molecular evidence. The new species can be distinguished from its congeners by the following combination of characters: shell with low, prosocline, rounded axial ribs and fine spiral striae, broader than high; aperture broader than shell height; radula with lateral teeth have only two or three faint, wavy ridges on inner side. A molecular analysis of partial mitochondrial COI and 16S DNA sequences supports the systematic position of the new taxon.

Key words

Diversity, southern China, taxonomy, Triculinae


Pomatiopsidae Stimpson, 1865 is a family of minute snails with shells usually 1–10 mm high. However, a few species can reach up to 20 mm in high. Typically, pomatiopsids inhabit rivers and streams, while some species also occur in brackish water or even in damp places on land (Liu et al. 1993; Lydeard and Cummings 2019). Shells of pomatiopsids vary in shape from spherical to oval, conical, or tower-shaped (Liu et al. 1979a). The Pomatiopsidae is widely distributed in Asia, South America, North America, Africa, and Australia. It is one of the most species-rich freshwater gastropod families, with approximately 36 recognized genera (Gerlach 2003; Ho and Dang 2010; Lydeard and Cummings 2019). There two main hypotheses about the origin of the Pomatiopsidae are that they originated in either Gondwanaland origin (Davis 1979) or Australia (Attwood 2009); neither of these hypotheses has been completely rejected yet by multilocus phylogenetic analyses (Kameda and Kato 2011; Wilke et al. 2013). The genus and species-level classification are poorly understood (Lydeard and Cummings 2019).

The tropical hills and rivers of China have rich biodiversity and are home to many species of freshwater snails. With at least 18 genera, China has the highest species richness of Pomatiopsidae in the world (Fulton 1904; Kang 1983; Davis et al. 1984; Davis et al. 1990; Davis et al. 1992; Zhang et al. 1997; Xiong and Li 2008; Lydeard and Cummings 2019).

In China, Pomatiopsidae are mainly distributed in the southwestern region (Shu et al. 2014), but the biodiversity of these freshwater snails is likely underestimated, especially in remote regions. Guangxi Province is in southwestern China, and 44 species of freshwater gastropods have been recorded from there (Liang 2023). However, only two genera of Pomatiopsidae have been recorded so far; Oncomelania (Gredler, 1881) and Tricula (Benson, 1843) occur in mountainous streams in the north of the province (Liu et al. 1979b).

In a recent survey in the Longjiang River, Hechi City, Guangxi, China, a new species of freshwater snail belonging to the genus Fenouilia Heude, 1889 was discovered. On comparison of its morphological traits with those of other freshwater snails known from this area, we conclude that this species is indeed undescribed. The new species can be distinguished from its congener Fenouilia kreitneri (Neumayr, 1880) by having low, rounded axial ribs on its shell, which is unique to this species. Molecular phylogenetic analyses, based on partial sequences of the mitochondrial 16S rRNA (16S) and COI genes, provide additional evidence supporting the novelty of this species. Our study contributes to a better understanding of pomatiopsid diversity in China and encourages the further exploration of freshwater gastropods in the region.

Materials and methods

Materials and morphological examination

All specimens were collected by hand in August 2022 and March 2023 on the Longjiang River, Yizhou District, Hechi City, Guangxi Province, China (Fig. 1). They were preserved in 95% ethanol and have been deposited in the Jiangsu Key Laboratory for Biodiversity and Biotechnology, Nanjing Normal University (NNU), Nanjing, China.

Figure 1. 

Known distribution of Fenouilia undata sp. nov. (Hechi City), and collection site (Longjiang River).

Before taking any action, preserved samples were soaked overnight in a saline solution. The tissue was extracted using anatomical needles. The shell and operculum were cleaned with a fine brush and then flushed with distilled water and photographed under a Nikon SMZ645 stereomicroscope. For traditional morphometrics, we measured five shell characteristics to the nearest 0.01 mm for each individual using digital calipers as follows: (1) shell height (H), which is the maximum dimension parallel to the axis of coiling; (2) shell width (W), the maximum dimension perpendicular to H; (3) length of aperture (LA), the maximum dimension from the junction of the outer lip with the penultimate whorl to the anterior lip; (4) width of aperture (WA), the maximum dimension perpendicular to LA; and (5) height of the body whorl (BW), the dimension from the lower margin of the aperture to the upper suture delimiting the first whorl. Other terminology used follows Liu et al. (1979a), Strong (2011), and Du et al. (2019).

Radulae were dissected from the buccal masses of three specimens and cleaned enzymatically with proteinase K following Du and Yang (2023); they were sonicated, mounted on aluminum specimen stubs with adhesive pads, and then observed using a JEOL JSM5610LV scanning electron microscope (SEM).

DNA extraction, PCR amplification, and phylogenetic analyses

Total genomic DNA was extracted from foot tissue of three ethanol-preserved specimens using a Trelief TM Animal Genomic DNA kit (Tsingke®). Partial sequences of 16S rDNA were amplified using the universal primer set 16Sar CGCCTGTTTATCAAAAACAT and 16Sbr CCGGTCTGAACTCAGATCACGT (Kessing et al. 1989). Partial sequences of COI were amplified using LCO1490 GCTCAACAAATCATAAAGATATT (Folmer et al. 1994) and HCO2198 TAWACTTCTGGGTGKCCAAARAAAT (Glaubrecht and Rintelen von 2003).

Each PCR reaction was performed in a total volume of 20 μL, including 9 μL of PCR mix, 8 μL of double distilled water, 1 μL of each primer and 1 μL of the DNA template. The PCR conditions were as follows: initial denaturation at 95 °C for 3 min; 35 cycles of denaturation at 95 °C for 40 s, annealing at 55 °C for 30 s and extension at 72 °C for 30 s; and final extension at 72 °C for 7 min. Both ends of sequences were obtained by automated sequencing using Applied Biosystems 3730 in Sangon Biotech Co. Ltd (Shanghai, China). In addition, two individuals of Lithoglyphopsis modesta (Gredler, 1886) and one of Fenouilia kreitneri (Neumayr, 1880) had their 16S rDNA extracted in this study.

To clarify the generic relationship of the new species, we included 16 sequences generated in this study with addition of 16S and COI gene sequences of 49 specimens representing 26 genera and 41 species, which were downloaded from GenBank (Table 1); Lithoglyphus naticoides (L. Pfeiffer, 1828) was used as the outgroup. Sequences obtained in the present study have been deposited in GenBank (for accession numbers, see Table 1). Sequences were aligned using MAFFT v. 7.505 based on the L-INS-i method (Katoh and Toh 2008). Pairwise distances between species were calculated using MEGA X (Kumar et al. 2018). The 16S rDNA and COI were concatenated in PHYLOSUITE v. 2.3 (Zhang et al. 2020).

Table 1.

Nucleotide compositions of partial 16S rDNA and COI sequences of specimens investigated in this study.

Genus Species GenBank References
Lithoglyphus Lithoglyphus naticoides AF445341 AF445332 Hausdorf et al. 2003
Bythinella Bythinella austriaca FJ028832 FJ028942 Benke et al. 2009
Bythinella austriaca FJ028831 FJ028943
Bythinella carinulata FJ028884 FJ029100
Paludinella Paludinella minima AB822685 AB822663 Wada et al. 2013
Paludinella minima AB822686 AB822664
Paludinella kuzuuensis AB822695 AB822675
Erhaia Erhaia jianouensis AF212894 AF213340 Wilke et al. 2000
Erhaia wangchuki KY798003 MT237715 Gittenberger et al. 2020
Akiyoshia Akiyoshia kobayashii AB611822 AB611823 Kameda and Kato 2011
Bithynia Bithynia tentaculata FJ160288 JX970605 Wilke et al. 2013
Pomatiopsis Pomatiopsis lapidaria AY676118 AF367636 Wilke et al. 2001
Robertsiella Robertsiella sp. AF531548 AF531550 Attwood et al. 2003
Pachydrobia Pachydrobia munensis KC832721 KC832700 Liu et al. 2014
Pachydrobia sp. KC832711 KC832690
Jullienia Jullienia rolfbrandti KC832718 KC832697
Hubendickia Hubendickia schuetti KC832709 KC832688
Hubendickia spiralis KC832710 KC832689
Jinghongia Jinghongia jinghongensis KC832728 KC832707
Manningiella Manningiella velimirovici KC832716 KC832695
Manningiella conica KC832719 KC832698
Manningiella polita KC832715 KC832694
Tricula Tricula bambooensis KC832720 KC832699
Tricula ludongbini KC832717 KC832696
Tricula hudiequanensis KC832712 KC832691
Tricula hongshanensis EF394876 EF394896 Guan et al. 2008
Oncomelania Oncomelania hupensis robertsoni DQ212900 DQ212855 Wilke et al. 2006
Oncomelania hupensis robertsoni DQ212901 DQ212856
Oncomelania minima AB611790 AB611795 Kameda and Kato 2011
Blanfordia Blanfordia integra AB611722 AB611723
Blanfordia japonica japonica AB611726 AB611727
Cecina Cecina manchurica AB611746 AB611747
Cecina manchurica AB611742 AB611743
Neotricula Neotricula burchi AF531542 AF531544 Attwood et al. 2003
Neotricula aperta MF663277 MF663265 Attwood et al. 2019
Gammatricula Gammatricula fujianensis AF212896 AF213342 Wilke et al. 2000
Gammatricula shini AB611798 AB611799 Kameda and Kato 2011
Gammatricula chinensis EU573993 AF253067 Wilke et al. 2000
Lacunopsis Lacunopsis munensis KC832726 KC832705 Liu et al. 2014
Delavaya Delavaya dianchiensis KC832713 KC832692
Paraprososthenia Paraprososthenia levayi KC832708 KC832687
Lithoglyphopsis Lithoglyphopsis modesta OR515659 PP327217 This study
Lithoglyphopsis modesta OR515660 PP327222
Fenouilia Fenouilia kreitneri OR515658 PP340173
Fenouilia undata sp. nov. OR515661 PP333612
Fenouilia undata sp. nov. OR515662 PP333613
Fenouilia undata sp. nov. OR515663 PP333614
Kunmingia Kunmingia kunmingensis OR784230 OR780554
Kunmingia kunmingensis OR784231 OR780555

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 analyses were performed using MrBayes v. 3.2.7 (Ronquist et al. 2012) and IQTREE v. 2.2 (Minh et al. 2020), 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 million generations, with sampling every 1,000 generations. The first 25% of sampled trees were discarded as burn-in when the standard deviation of split frequencies of 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.

Furthermore, to investigate the behavior of the new species, six individuals were maintained in an artificial field environment within the laboratory of Hunan Fisheries Science Institute for one year.


Family Pomatiopsidae Stimpson, 1865

Subfamily Pomatiopsinae Stimpson, 1865

Genus Fenouilia Heude, 1889

Fenouilia undata Chen & He, sp. nov.

Materials examined

Holotype : China • Guangxi Province, Hechi City, Yizhou District, Longjiang River; 24.4927°N, 108.6851°E; August 2022; Xu Cheng Wei & Yue Ming He leg.; NNU230701 (Fig. 2A–D), shell height 3.39 mm. Paratypes: China • 2 specimens; same locality data as holotype; August 2022; NNU230702–03 • _2_ specimens; same locality data as holotype; March 2023; NNU230704–05. Shell height of all paratypes: 3.04–3.44 mm (Fig. 2E–P, Table 2).

Table 2.

Measurements of Fenouilia undata sp. nov. (in mm). Abbreviations: W, shell width; BW, height of the body whorl; H, shell height; LA, length of aperture; WA, width of aperture.

Number H W LA WA BW
Holotype NNU230701 3.39 4.95 3.63 2.64 2.84
Paratype 1 NNU230702 3.25 4.67 3.44 2.43 3.07
Paratype 2 NNU230703 3.44 4.68 3.49 2.61 2.92
Paratype 3 NNU230704 3.07 4.37 3.22 2.49 2.81
Paratype 4 NNU230705 3.04 4.01 3.06 2.26 2.74
Figure 2. 

Fenouilia undata sp. nov. shells and operculum A–D holotype, NNU230701 E–H paratype, NNU230702 I–L paratype, NNU230703 M paratype, NNU230704 N paratype, NNU230705 O, P operculum, holotype, NNU230701. Scale bars: 2 mm (A–N); 1 mm (O–P).


Shell small, thin but solid, with rounded, rather flattened shape, and width greater than height; sculptured with low, rounded axial ribs and fine spiral striae; whorls 4–5; body whorl swollen and large. Suture shallow; umbilicus narrow, crescent-shaped or closed. Aperture large, its length greater than shell height. Operculum ovate, corneous, slightly transparent, yellowish.


Shell small, 3.04–3.44 mm high, thin but solid, with a rounded, rather flattened shape; whorls 4–5; body whorl swollen and large, taking up most (about 84–94%) of shell; whorls of spire rapidly expanding. Shell width longer than shell height (Fig. 2). Apex obtuse, usually eroded. Suture low. Shell amber-yellow, with low, prosocline, rounded axial ribs and fine spiral striae. Aperture round, large, broader than shell height. Lip slightly thickened; inner lip smooth, white; outer lip white or yellowish and slightly rolled outward. Umbilicus narrowly crescent-shaped or closed; base white (Fig. 2A–N).

Operculum ovate, smaller than aperture, corneous, thin, slightly transparent, yellowish, length 1.86–2.12 mm, width 1.53–1.72 mm; surface, including nucleus, of operculum smooth; nucleus located at bottom left third (Fig. 2O, P).

Radula small; ribbon approximately 0.88 mm long. Central tooth with one large, triangular, pointed major cusp without serrations, with two small, sharp cusps on either side at base. Inner side of lateral teeth with two or three faint, wavy ridges; outer side smooth. Inner marginal teeth with five or six small cusps. Outer marginal teeth with 6–8 small cusps (Fig. 3).

Figure 3. 

Radula of Fenouilia undata sp. nov. A frontal view of radula B, C magnified view of radula.

Tentacles short, white; snout stubby, white, black pigmented. Mantle smooth, light gray, with small black spots. Intestine wider than base of tentacle; digestive gland milky white. Penis translucent white, thin, coiled, located behind right tentacle in neck area (Fig. 4).

Figure 4. 

Fenouilia undata sp. nov. A dissection with labelled structures of female B head of male. Abbreviations: e, eye; t, tentacle; sn, snout; f, foot; op, operculum; dg, digestive gland; int, intestine; pe, penis. Scale bars: 1 mm (A); 0,5 mm (B).

Habitat and distribution

The new species was discovered in the Longjiang River, where the depth of the water was less than 5 m, water flow is variable, and the substrate is composed of large stones (Fig. 5).

Figure 5. 

Fenouilia undata sp. nov. A color in life B natural habitat. Photographs by Xu Cheng Wei and Yue Ming He.


In the laboratory aquarium, the new species fed on algae present on the surface of stones or watergrass. Snails reproduced many times during their year in captivity. Each brownish egg was laid 1.5 mm from the next. Eggs were affixed to the surfaces of rocks with a secretion. In some months, some individuals were observed to occasionally perform a “dance” in which they repeatedly twisted their shells clockwise or counterclockwise. They were more active at night.


The genus Fenouilia was established by Heude (1889) for Fenouilia bicingulata (Heude, 1889) from Dali, Yunnan, China; this species has a trochoidal shell, with rough, raised prosocline growth lines and no umbilicus. Subsequently, Davis et al. (1983) considered F. bicingulata to be a synonym of F. kreitneri (Neumayr, 1880); thus, the genus was thought to contain only a single species, until now. With prosocline axial ribs, triangular central tooth, and narrowly crescent-shaped or absent umbilicus, the new species is similar to F. kreitneri. However, the new species can be distinguished by its broader shell. In addition, F. undata sp. nov. has shorter tentacles (vs longer tentacles in F. kreitneri), and there are three ridges only on the inner side (vs. lateral teeth with obvious ridges on both sides). The adult shell of F. undata sp. nov. is similar to that of Lacunopsis munensis (Brandt, 1968) and Lithoglyphopsis modesta (Gredler, 1886). These species differ in the relative length of the aperture to shell height (the aperture is longer than shell height in F. undata, but shorter in L. munensis) and in relative shell width (the shell is broader than height in F. undata, but narrower in L. modesta).

Molecular results

The concatenation of COI and 16S rDNA yielded 1229 sites. The GTR+F+R5 model was selected as the best-fit of nucleotide substitution by BIC. Phylogenetic analyses revealed BI and ML trees with largely consistent topologies (Figs 6, 7). The average 16S genetic distance (uncorrected p-distance) between the F. undata sp. nov. and F. kreitneri is 1.04%; the COI p-distance is 6.96%.

Figure 6. 

Bayesian-inference (BI) tree inferred from concatenated 16S and COI gene sequences. Posterior probabilities are shown on the left of nodes on the tree.

Figure 7. 

Maximum-likelihood (ML) tree inferred from concatenated 16S and COI gene sequences. Bootstrap supports are shown on the left of nodes on the tree.


From the Latin adjective undata (wavy or wave-like form). We suggest the Chinese common name 波浪龙骨螺.


The phylogenetic relationships and morphological traits found in this study support the placement of the new species in the genus Fenouilia. The prosocline axial ribs on the shell of the new species resemble the prominent, rough, raised prosocline growth lines present in F. kreitneri, and the radular of both species has a triangular central tooth without serrated cusps. The molecular phylogenies based on ML and BI analyses show that Fenouilia undata sp. nov. and F. kreitneri are nested in a monophyletic group with strong support (BS = 94%, BPP = 0.93) and sister to Lithoglyphopsis modesta.

Our phylogenetic tree includes all the genera in China except Guoia Davis & Chen, 1992, Wuconchona Kang, 1983, and Parapyrgula Annandale & Prashad, 1919. The new species can be distinguished from Wuconchona niuzhuangensis (Kang, 1983) and Parapyrgula cogginiii (Annandale & Prashad, 1919) by the rounded, flattened shell with its width greater than height. The new species can be distinguished from Guoia viridulula (Möllendorff, 1888) by the presence of axial ribs on the shell and the aperture being longer than the shell height.

The hydrological environment is complex and heterogeneous in southern China. There are still many gaps in surveys for freshwater snails, and more new species have to be discovered. However, the freshwater-snail fauna has been given little attention, especially the small species. With the environmental destruction and habitat modifications, many freshwater snails are gradually disappearing or becoming extinct (Du et al. 2011). Our study highlights the necessity and importance of further field surveys of freshwater snails which will help promote the conservation of freshwater ecosystems. We suggest that further intensified survey efforts are urgently required for accurate understanding of the freshwater snail diversity in southern China.


We acknowledge Xu Cheng Wei from Guangxi for assistance in obtaining samples for this study. We are grateful to Kai Rui Jiao and Long Jie Xia from Nanjing Normal University for their help in image processing. We also extend our thanks to Yang Yang Li from Nanjing Normal University for assistance in DNA extraction.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.


This research was funded by the Jiangsu Key Laboratory for Biodiversity and Biotechnology, College of Life Sciences, Nanjing Normal University, and the Hunan Fisheries Science Institute. Financial support from the National Natural Science Foundation of China (No. 32200356 to PD) and from the Natural Science Foundation of the Jiangsu Higher Education Institutions (No. 22KJB180018 to PD) is acknowledged.

Author contributions

All authors have contributed equally.

Author ORCIDs

Hui Chen

Yue Ming He

Chong Rui Wang

Da Pan

Data availability

All of the data that support the findings of this study are available in the main text.


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