A new genus of river snails, Bakyietaia (Mollusca, Viviparidae), from South China and the Indochinese Peninsula

1
Eu opean Jou nal o Taxonomy 1005: 1–64
h ps://doi.o g/10.5852/ej .2025.1005.2985
eu opeanjou nalo axonomy.eu
ISSN 2118-9773
2025 · Zhang L.-J. e al.
This wo k is licensed unde a C ea i e Commons A ibu ion License (CC BY 4.0)
Recei ed: 29 July 2024 • Accep ed: 7 Ap il 2025 • Published: 31 July 2025
Topic edi o : Magalie Cas elin • Sec ion edi o : Thie y Backeljau • Desk edi o : Pepe Fe nández
Monog aph
u n:lsid:zoobank.o g:pub:BAD3B10F-13AB-43E3-8B52-0B613A992602
A new genus o i e snails, Bakyie aia (Mollusca, Vi ipa idae),
om Sou h China and he Indochinese Peninsula
Le-Jia ZHANG 1+ , Yu-Hsiu YEN 2,*+ , Zhe-Yu CHEN 3  , Li-Na DU 4 ,
Ting Hui NG 5   & Thomas on RINTELEN 6,*
1 Kunming Ins i u e o Zoology, Chinese Academy o Sciences, Kunming 650023, China.
1 Museum ü Na u kunde, Leibniz Ins i u ü E olu ions und Biodi e si ä s o schung,
Be lin 10115, Ge many.
1 Sou heas Asia Biodi e si y Resea ch Ins i u e, Chinese Academy o Sciences, Yezin,
Nay Pyi Taw, 05282, Myanma .
2 Depa men o Ma ine Bio echnology and Resou ces, Na ional Sun Ya -sen Uni e si y,
Kaohsiung 80424, Taiwan.
3 Pa asi ology Resea ch G oup, Depa men o Ve e ina y Biosciences, Facul y o Science,
The Uni e si y o Melbou ne, Pa k ille VIC 3010, Aus alia.
3 Depa men o Li e Sciences, Na u al His o y Museum, London SW7 5BD, UK.
4 Key Labo a o y o Ecology o Ra e and Endange ed Species and En i onmen al P o ec ion,
Guangxi No mal Uni e si y, Minis y o Educa ion, Guilin, Guangxi 541004, China.
5 Ins i u e o T opical Biology and Conse a ion, Uni e si i Malaysia Sabah, Jalan UMS,
88400 Ko a, Malaysia.
5 Lee Kong Chian Na u al His o y Museum, Na ional Uni e si y o Singapo e, 2 Conse a o y D i e,
Singapo e 117377, Republic o Singapo e.
6 Museum ü Na u kunde, Leibniz Ins i u ü E olu ions- und Biodi e si ä s o schung,
Be lin 10115, Ge many.
* Co esponding au ho s: [email p o ec ed] – [email p o ec ed]
1 Email: [email p o ec ed]
3 Email: [email p o ec ed]
4 Email: [email p o ec ed]
5 Email: [email p o ec ed]
+
These wo au ho s con ibu ed equally o he manusc ip
Abs ac . A new genus o he i e snail amily Vi ipa idae, Bakyie aia Zhang, Yen & on Rin elen
gen. no ., is desc ibed based on a comp ehensi e s udy o i s mo phology, ana omy, m DNA (COI),
ecology and dis ibu ion. This genus is widely dis ibu ed in Sou h China and he Indochinese Peninsula.
I can be dis inguished om all o he i ipa id gene a, especially Angulyag a, Sino aia and Anulo aia, by
cha ac e s o he columella lip o he shell, he ope culum and he ma ginal ee h o adula. Based on an
in eg a i e axonomic app oach, 17 species a e assigned o his new genus and desc ibed he e, including
7 new combina ions and 10 new species: Bakyie aia subcos a a gen. e comb. no ., B. polyzona a gen. e
Eu opean Jou nal o Taxonomy 1005: 1–64 (2025)
2
comb. no ., B. boe ge i gen. e comb. no ., B. duchie i gen. e comb. no ., B. guangdungensis gen. e
comb. no ., B. mu ica gen. e comb. no ., B. wilhelmi gen. e comb. no ., B. chenghuang Zhang, Yen &
on Rin elen gen. e sp. no ., B. jingweiae Yen, Zhang & on Rin elen gen. e sp. no ., B. liusanjieae
Zhang, Yen & on Rin elen gen. e sp. no ., B. a is ena o is Yen, Zhang & on Rin elen gen. e sp. no .,
B. liangzhuo um Yen, Zhang & on Rin elen gen. e sp. no ., B. luikongi Yen, Zhang & on Rin elen
gen. e sp. no ., B. on inalis Zhang, Yen & on Rin elen gen. e sp. no ., B. naiadica Zhang, Yen & on
Rin elen gen. e sp. no ., B. luuemxlong Zhang, Yen & on Rin elen gen. e sp. no . and B. ind apu a
Zhang, Yen & on Rin elen gen. e sp. no . The monophyly o all mo phospecies is suppo ed by he COI
phylogeny. Radula cha ac e s, especially he numbe o cusps o he ou e ma ginal ee h, a e conside ed
aluablein heclassi ica iono specieso Bakyie aia wi h simila shells.
Keywo ds. Angulyag a, biogeog aphy, gas opod, sys ema ics, axonomy.
Zhang L.-J., Yen Y.-H., Chen Z.-Y., Du L.-N., Ng T.H. & on Rin elen T. 2025. A new genus o i e snails,
Bakyie aia (Mollusca, Vi ipa idae), om Sou h China and he Indochinese Peninsula. Eu opean Jou nal o
Taxonomy 1005: 1–64. h ps://doi.o g/10.5852/ej .2025.1005.2985
In oduc ion
Vi ipa idae J.E. G ay, 1847 ( i e snails) is a amily o eshwa e snails comp ising 125 o 150 alid
desc ibed ex an species wi h a wo ldwide na i e dis ibu ion excep o Sou h Ame ica and An a c ica
(S ong e al. 2008). The Vi ipa idae ha e a ich ossil eco d (P ashad 1928), and many ex inc species a e
ound ou side he cu en geog aphic ange o his amily, such as in he Middle Eas (Si an e al. 2006) and
Sou h Ame ica (Ghila di e al. 2011). The oldes ossils can be aced back o he Ea ly o Middle Ju assic:
Vi ipa us sangongheensis Zhu, 1994 om Xinjiang, China (Ea ly Ju assic; Zhu 1994) and Ba honella
sco ica (Ta e, 1873) om England, UK (Middle Ju assic; T acey e al. 1993). The ea ly o igin, abundan
ossil eco d and (almos ) wo ldwide dis ibu ion o he Vi ipa idae has d awn many esea che s’ a en ion
o he sys ema ics, e olu iona y his o y and biogeog aphy o his amily (Sengup a e al. 2009; Schul heiß
e al. 2014; Zhang e al. 2015; Hi ano e al. 2019a, 3019b, 2023; S elb ink e al. 2020; Ye e al. 2020).
Sou h China and Sou heas Asia ha bou he highes biodi e si y o Vi ipa idae (S ong e al. 2008;
Zhang e al. 2015; S elb ink e al. 2020). Yunnan P o ince in Sou hwes e n China ha bou s en gene a,
including ou endemic ones (Anula ya Zhang & Chen, 2015; Dalipaludina Zhang, 2023; Ma ga ya
Ne ill, 1877; Tchangma ga ya He, 2013) and a leas 15 endemic species (Zhang e al. 2023). Sou heas
Asia ha bou s 12 gene a, including ou endemic o he Indochinese Peninsula (Anulo aia B and ,
1968; Ey iesia Fische , 1885; Taia Annandale, 1918; T ocho aia B and , 1974) and wo endemic o
insula Sou heas Asia (Cele aia Clench, 1966; To o aia F. Haas, 1939) (Molluscabase 2023). Howe e ,
Vi ipa idae om his egion a e unde s udied, wi h only a ew ecen s udies ocussing on his ho spo
o i ipa id di e si y (Zhang e al. 2015; S elb ink e al. 2019, 2020; Zhang e al. 2023).
Rao (1925) es ablished a new genus in he Vi ipa idae, Dac ylochlamys, o a single species wi h many
smoo h, s ong keels on he shell, endemic o he Manipu Valley o India, Paludina oxy opis Benson,
1836.Howe e ,Rao(1931)la e iden i ied Dac ylochlamys Rao, 1925 as a junio homonym o he cilia e
Dac ylochlamys Lau e bo n, 1901; he e o e, Angulyag a Rao, 1931 was es ablished as a eplacemen
name o his genus. Angulyag a soon became a ‘was ebaske axon’ o species om Asia possessing a
shell wi h many smoo h, s ong keels (P ashad 1928; Yen 1939, 1943; S a oboga o 1970; Liu e al. 1995).
Howe e , mos o hese species a e dis inc om he ype species o Angulyag a based on cha ac e s o he
shell, adula, ope culum, and ana omy. The e o e, a sys ema ic e ision o Angulyag a is needed.
He e, we e ise he species o Angulyag a om Sou h China and he Indochinese Peninsula based on
hei mo phology (shell, ope culum, adula), ecology, dis ibu ion and a ia ion in he mi ochond ial
ZHANG L.-J. e al., New genus o Vi ipa idae (Mollusca) om Sou h China
3
COI gene. Ou in eg a i e s udy has e ealed ha he e a e a leas 17 species wi hin his g oup, and all
o hese species o Angulyag a should ac ually be assigned o a new genus desc ibed he e: Bakyie aia
Zhang, Yen & on Rin elen gen. no .
Ma e ial and me hods
Ma e ial
Typespecimenso  en i ipa idspecies om i eins i u ions(Na u alHis o yMuseum,London
(NHMUK); Gymnasium de F anziskane , Bozen (GFBI); Na u his o isches Museum, Vienna (NHMW);
Na u museum Senckenbe g, F ank u (SMF); Muséum na ional d’His oi e na u elle, Pa is (MNHN))
ha e been examined. The ype specimens o nine new species desc ibed he e ha e been deposi ed in
he Kunming Ins i u e o Zoology, Chinese Academy o Sciences, Kunming (KIZ) and he Museum ü
Na u kunde, Be lin (ZMB), non- ypes a e also in he collec ions o Le-Jia Zhang (ZLJ) and Yu-Hsiu
Yen (YHY). In addi ion, compa a i e non- ype ma e ial held in ZMB, KIZ and and he BORNEENSIS
Collec ion, Ins i u e o T opical Biology and Conse a ion, Uni e si i Malaysia Sabah (BOR) ha e
also been examined. The specimens used o he mo phological and molecula analysis we e collec ed
du ing 2006–2022. A wo-mon h expedi ion o collec ing species o Vi ipa idae om i e s, sp ings,
lakes and we lands o Sou he n China was conduc ed in ea ly 2022. Al oge he 73 si es we e sampled
and a e illus a ed in he map (Fig. 1).
Fig. 1. Map o he main i e basins in Sou h China and he Indochinese Peninsula wi h he collec ing
si es o species o Bakyie aia Zhang, Yen & on Rin elen gen. no . Si es wi h mo e han one species a e
indica ed by an a ow and mul iple species symbols on a g ey backg ound. The symbol “?” indica es
si es wi h species o Bakyie aia onlyiden i ied ogenusle el.
Eu opean Jou nal o Taxonomy 1005: 1–64 (2025)
4
Examina ion o mo phology
The heigh (H) and wid h (W) o shells o ma u e and comple e specimens we e measu ed wi h an
elec onic callipe o a p ecision o 0.1 mm. The mass (M) o se e al selec ed shell specimens was
measu ed by an elec onic scale o a p ecision o 0.01 g. Selec ed shells comp ised nine specimens o
Angulyag a oxy opis (Benson, 1836) and 44 specimens o species o Bakyie aia gen. no . The shells
we e comple ely d y and he mass was cons an . All specimens we e pho og aphed in a consis en
o ien a ion using a Sony alpha 7R 4 came a. Radulae we e ex ac ed by dissec ion, cleaned by boiling in
1% NaOH solu ion o hal an hou and insed wi h dis illed wa e . Radulae we e coa ed wi h gold and
imaged wi h a Zeiss EVO LS10 scanning elec on mic oscope. Te ms used o desc ibing he ope culum
ollow Zhang & on Rin elen (2021).
Abb e ia ions o ea u es o he adula:
c = cen al ee h
im = inne ma ginal ee h
l = la e al ee h
om = ou e ma ginal ee h.
Molecula me hods
DNAwasex ac ed om10–20mg oo  issue omeachsnail,usingamollusc-speci icCTAB/
chlo o o m ex ac ion p o ocol (Winnepenninckx e al.1993). A agmen (570 bp) o he mi ochond ial
cy och omecoxidasesubuni I(COI)genewasampli ied h oughpolyme asechain eac ion(PCR)wi h
he ollowingp ime pai :LCO1490,5′-GGTCAACAAATCATAAAGATATTGG-3′andCOX-B7R,
5′-ACCACCAGCTGGATCAAAAA-3′.(Schul heiße al.2011)PCRampli ica ionswe econduc ed
in25μl olumesunde  he ollowingcyclingcondi ions:ini ialdena u ings epa 94°C o 10min,
ollowedby30cycleso 94°C o 1min,50°C o 1min,and72°C o 1min,wi ha inalex ensions ep
o 10mina 72°C.Thepu i ica ionandsequencingwe econduc edbyMac ogenEu ope,Ams e dam,
Ne he lands.
Sequence and phylogene ic analyses
140 new COI sequences o species o Vi ipa idae and 16 sequences o 13 gene a o he sub amily
Bellamyinae Roh bach, 1937, mos ly om S elb ink e al. (2020), we e included in he p esen s udy
(Supp. ile1). Vi ipa us con ec us (Mille , 1813) o he sub amily Vi ipa inae J.E. G ay, 1847 was
selec ed as ou g oup based on S elb ink e al. (2020).
Sequences we e aligned using MUSCLE as implemen ed in Geneious P ime 2020
(h ps://www.geneious.com). Gene ic dis ances we e calcula ed using MEGA X (Kuma e al. 2018).
TN93+G+Iwassugges edas hebes - i modelo sequencee olu ion o ou da ase bymeanso  he
Akaike and Bayesian in o ma ion c i e ia in MEGA X. A Bayesian in e ence (BI) analysis was pe o med
wi h M Bayes e . 3.2.6 (Ronquis e al. 2012) as implemen ed in Geneious P ime 2020 wi h ou
independen chains o 5000000gene a ions,sample eq=1000,bu nin=25%,andwecon i med
ha con e gence was eached based on he ace plo s gene a ed in Geneious P ime 2020. A maximum
likelihood (ML) analysis was conduc ed wi h RAxML as implemen ed in Geneious P ime 2020, wi h
nodal suppo es ima ed by 1000 boo s ap eplica es (Kozlo e al. 2019).
Species delimi a ion
Assemble Species by Au oma ic Pa i ioning (ASAP) (Puilland e e al. 2020) was pe o med on he
ASAP web pla o m (h ps://bioin o.mnhn. /abi/public/asap/) using he Jukes Can o (JC69) model and
simplegene icdis ancecalcula edwi hde aul se ings:p obabili y.01,seed alue−1,gene icdis ances
be ween 0.005 and 0.05.
ZHANG L.-J. e al., New genus o Vi ipa idae (Mollusca) om Sou h China
5
Fig. 2. Bayesian in e ence phylog am o he species o Bakyie aia Zhang, Yen & on Rin elen gen. no .
and some ela ed i ipa id gene a based on cy och ome c oxidase subuni I (COI) wi h ope a ional
axonomic uni (OTU) assignmen s. Numbe s a nodes indica e Bayesian pos e io p obabili ies (le )
and maximum likelihood boo s ap suppo alues ( igh ). Suppo ing alues lowe han 0.5/50 a e no
p esen ed and a e shown as ‘-’. Backg ound in colou s indica e new combina ions (blue) and new species
(o ange). ASAP: Assemble Species by Au oma ic Pa i ioning. Scale ba = 0.04 subs i u ions pe si e.

Eu opean Jou nal o Taxonomy 1005: 1–64 (2025)
6
Table 1. Shell measu emen s o species o Angulyag a Rao, 1931 and Bakyie aia Zhang, Yen & on
Rin elen gen. no . Abb e ia ions: H = heigh ; M = mass; N = measu ed specimens; n = measu ed
comple ely d y shells; W = wid h.
Species N n H (mm) W (mm) W/H M (g)
Angulyag a oxy opis 9 9 34.60 ± 3.77 27.15 ± 3.79 0.78 ± 0.03 1.01 ± 0.28
Bakyie aia polyzona a 326.60 ± 1.91 19.19 ± 1.30 0.72 ± 0.01 –
Bakyie aia subcos a a 10 27.17 ± 3.64 20.25 ± 1.75 0.75 ± 0.05 2.52 ± 0.36
Bakyie aia boe ge i 9 7 26.76 ± 6.67 18.87 ± 5.12 0.70 ± 0.04 2.16 ± 2.78
Bakyie aia duchie i 232.88 ± 1.00 24.46 ± 1.77 0.74 ± 0.03 –
Bakyie aia guangdungensis 7 3 20.87 ± 0.93 14.57 ± 0.61 0.70 ± 0.03 0.78 ± 0.18
Bakyie aia mu ica 11 222.14 ± 1.74 15.49 ± 1.06 0.70 ± 0.04 0.84 ± 0.16
Bakyie aia wilhelmi 10 10 25.53 ± 3.72 17.28 ± 2.11 0.68 ± 0.04 1.63 ± 0.97
Bakyie aia chenghuang 3 2 23.70 ± 5.37 17.67 ± 3.42 0.74 ± 0.03 1.19 ± 0.02
Bakyie aia jingweiae 9 9 26.79 ± 5.37 17.67 ± 3.42 0.66 ± 0.04 1.94 ± 1.90
Bakyie aia liusanjieae 12 1 25.69 ± 3.35 18.98 ± 2.31 0.74 ± 0.04 3.00
Bakyie aia a is ena o is 619.78 ± 1.67 14.50 ± 1.48 0.73 ± 0.03 –
Bakyie aia liangzhuo um 8 4 25.25 ± 3.07 18.13 ± 2.33 0.72 ± 0.04 0.77 ± 0.22
Bakyie aia luikongi 320.66 ± 0.70 15.44 ± 0.62 0.75 ± 0.02 –
Bakyie aia on inalis 8 1 24.25 ± 2.59 17.44 ± 1.84 0.72 ± 0.03 0.40
Bakyie aia naiadica 325.73 ± 0.62 18.02 ± 0.31 0.70 ± 0.01 –
Bakyie aia luuemxlong 423.56 ± 2.58 16.49 ± 1.29 0.70 ± 0.02 –
Bakyie aia ind apu a 3 3 26.92 ± 2.45 17.67 ± 1.40 0.66 ± 0.01 1.07 ± 0.41
Fig. 3. Compa ison o Bakyie aia Zhang, Yen & on Rin elen gen. no . and Angulyag a Rao, 1931 wi h
espec o pa ame e s H and W/H (A), and wi h espec o pa ame e s H and log (10M) (B).
ZHANG L.-J. e al., New genus o Vi ipa idae (Mollusca) om Sou h China
7
Resul s
Gene ic di e en ia ion, phylogeny and species delimi a ion
The COI p-dis ances wi hin species o Bakyie aia gen. no . a e 0–1.42% and be ween species o
Bakyie aia a e 1.68%–7.53%; he COI p-dis ances be ween Bakyie aia and o he gene a o Vi ipa idae
a e 8.45%–15.65%.
The phylogene ic ees econs uc ed by BI and ML based on COI a e cong uen , bu he ML ee
(Supp. ile2) con ains se e al subs ain ial poly omies and, he e o e, only he BI ee is displayed
(Fig. 2). The new genus Bakyie aia gen. no . is monophyle ic wi h high and low suppo in he BI and
ML ees, espec i ely, and i is dis inc om all o he gene a o Vi ipa idae included in he analysis. The
ype species o Angulyag a (Angulyag a oxy opis) is no included in he cu en ee. Angulyag a cos a a
(Quoy & Gaima d, 1834) clus e s wi h To o aia c . maini ensis (Ba sch, 1907) wi h a high deg ee o
suppo and is no closely ela ed o any species o Bakyie aia. Filopaludina Habe, 1964 (including
ype species F. bengalensis (Lama ck, 1822), F. dolia is (A. Gould, 1844), F. ja anica ( on dem Busch,
1844), F. decipiens (Tappa one Cane i, 1883)) is monophyle ic wi h mode a e o high suppo and in
he BI analysis, and is eco e ed as he en a i e sis e clade o Bakyie aia wi h low suppo . The 17
mo phologicallyiden i iedspecieso Bakyie aia a e each eco e ed as monophyle ic, mos ly wi h high
suppo alues, al hough he suppo o some species such as B. liangzhuo um gen. e sp. no . and
B. wilhelmi is weak o mode a e.
The 17 g oups/ope a ional axonomic uni s (OTUs) gi en by he ASAP analysis we e he bes pa i ion
om he lowes ASAP-sco e unde all de aul se ings. The wo ASAP esul s a e consis en wi h he
phylogenyandmo phologicaliden i ica ions.
S a is ics o shell cha ac e s
The shell measu emen s e eal ha Angulyag a oxy opis has a la ge , mo e compac bu ligh e shell
compa ed o species o Bakyie aia gen. no . The shell heigh o Angulyag a oxy opis is la ge han ha
o any species o Bakyie aia, and he la ge W/H o Angulyag a oxy opis shows ha i has a ela i ely
mo e compac shell ega dless o size (Table 1, Fig. 3A). Angulyag a oxy opis always has a la ge shell
han species o Bakyie aia when he mass o bo h is he same; co espondingly, Angulyag a oxy opis
always has a much ligh e shell han species o Bakyie aia when he heigh o bo h is he same (Fig. 3B).
Values a e a i hme ic means and s anda d de ia ions o heigh (H) and wid h (W) o N measu ed
specimens and mass (M) o n measu ed comple ely d y shells.
Sys ema ics
Class Gas opoda Cu ie , 1795
O de A chi aenioglossa Halle , 1892
Family Vi ipa idae J.E. G ay, 1847
Sub amily Bellamyinae Roh bach, 1937
Genus Angulyag a Rao, 1931
Dac ylochlamys Rao, 1925: 132 [non Dac ylochlamys Lau e bo n, 1901 (Ciliopho a), junio homonym].
Angulyag a Rao, 1931: 301.
Type species
Paludina oxy opis Benson, 1836, by o iginal designa ion.
Eu opean Jou nal o Taxonomy 1005: 1–64 (2025)
8
Diagnosis
“The shell is o la ge size, conical, hin bu o namen ed wi h p ominen smoo h spi al idges which a e
conca eon hein e nalsu ace.Thebaseissomewha  la ened.Theumbilicusisb oadly ima ewi ha
b oadchanneldescendingdownwa ds omi .Theman lehasenla gedandhighly ascula  inge -shaped
p ocesses on he ma gin” (Rao 1925).
Rema ks
The shell o Angulyag a is mos simila o ha o To o aia o Dalipaludina, all o hem possessing a
la ge hin shell wi h s ong keels, acu e apex, and simple columella lip. I can be dis inguished om he
la e wo gene a based on he ollowing combina ion o cha ac e s: eleoconch who ls wi h s ong keels
ob iously da ke han he es o he shell su ace in colou , open umbilicus bo de ed by a s ong keel;
ope culum wi h g ow h lamellae on ex e io su ace and adial s iae on in e io su ace; adula wi h
na ow ma ginal ee h, he wid h o bo h inne and ou e ma ginal ee h is only hal ha o he la e al
ee h,andou e ma ginal ee hwi h15 inycusps;highly ascula  inge -shapedp ocesseson heman le
ma gin.
Angulyag a oxy opis (Benson, 1836)
Fig. 4
Paludina oxy opis Benson, 1836: 745 ( ype locali y no s a ed in he o iginal desc ip ion).
Paludina py amida a on dem Busch, 1844 in Philippi 1842–1850:113,pl.1 igs3–4.
Vi ipa a oxy opis – P es on 1915: 84. — Annandale & Seymou Sewell 1920: 113. — Annandale 1921:
544–552, igs2–5.
Dac ylochlamys oxy opis – Rao 1925: 132.
Angulyag a oxy opis –Rao1931:301.—SubbaRao1989:51, ig.70.—P eecee al. 2022: 94,
ig.38a.
Diagnosis
Shell la ge, agile and ligh weigh . Each eleoconch who l wi h h ee s ong keels, open umbilicus
bo de ed by one s ong keel. Ope culum wi h g ow h lamellae on ex e io su ace and adial s iae on
in e io su ace. Inne and ou e ma ginal ee h o adula bo h nea ly hal o la e al oo h in wid h. Man le
edgewi hanumbe o  inge -shapedp ocesses.
Ma e ial examined
Holo ype
BENGAL•d yp ese edspecimen;NHMUK.1879.12.26.120.
O he ma e ial
INDIA • 8 specs (p ese ed d y, 6 adul s and 2 ju eniles); Manipu , Lok ak “Log ak Lake”;
NHMUK.1974.03.7.1 oNHMUK.1974.03.8•2 specs (p ese ed d y); Assam, Manipu , Lok ak Lake;
NHMUK.2019.04.31.
Desc ip ion
Shell (Fig. 4A–F) la ge, b oadly conical, hin, ligh weigh , agile, oli e o g eenish b own; up o six
who ls a adul hood, including one ela i ely smoo h p o oconch who l, apex acu e; each eleoconch
who l wi h h ee s ong p ima y keels o da ke colou , he s onges keel abo e su u e, one o wo
weake smoo h seconda y keels loca ed be ween p ima y keels; base o shell wi h h ee o ou spi al
keels o da ke colou , one o wo weake seconda y keel loca ed be ween s ong p ima y keels; ape u e
ZHANG L.-J. e al., New genus o Vi ipa idae (Mollusca) om Sou h China
9
o a e, inne shell su ace bluish whi e, wi h ob ious g oo es co esponding o he posi ion o he h ee
s ong keels, ou e lip hin, h ee s ong keels ex ending o ma gin o ou e lip, columella lip hin and
simple; umbilicus b oadly open, bo de ed by a s ong keel.
Ope culum (Fig. 4G–H) co neous, anspa en pale b own, a he hin; nucleus sub-cen al, close o
inne ma gin; ex e io su ace wi h egula ly dis ibu ed g ow h lamellae; in e io su ace wi h la ge
inne ope cula egion and adial s iae s a ing om nucleus; nuclea egion ela i ely la ge, smoo h
and glossy, wi h eins a ma gin.
Radula cen al oo hwi honeb oadcen alden icleand i esmallsha pcuspsonei he side;la e al
oo h wi h one b oad cen al den icle, h ee small sha p cusps on inne side and six small sha p cusps
on ou e side; inne and ou e ma ginal ee h na ow, bo h nea ly hal o la e al oo h in wid h; inne
ma ginal oo h wi h one b oad cen al den icle and h ee small sha p cusps on ei he side; ou e ma ginal
oo hwi h15 inysha pcusps(summa izedbasedon he igu eanddesc ip ion omAnnandale1921).
Man leedgewi hanumbe o  inge -shapedp ocesses, h eeo  hesep ocessesla ge ,eachla gep ocess
co esponding oag oo eo  heinne shellsu ace;gilllamellawi hab oadbaseandalong inge -
shaped pa ; abou en emb yos and en eggs in he u e us o a p egnan emale (summa ized based on
he igu eanddesc ip ion omRao(1925)andAnnandale(1921)).
Rema ks
This species can be easily dis inguished om he o he i ipa id species acco ding o he ema ks o
he genus. As he ype species o Angulyag a, A. oxy opis shows la ge mo phological and ana omical
Fig. 4. Shells and ope culum o Angulyag a oxy opis (Benson, 1836). A–D. NHMUK.2019.04.31.1,
Lok ak Lake, India. E–F. NHMUK.2019.04.31.2, Lok ak Lake, India. G–H. Ope culum o
NHMUK.2019.04.31.1. Scale ba s = 1 cm, op le ba applies o all shells, sepa a e ba o ope culum
G–H.
Eu opean Jou nal o Taxonomy 1005: 1–64 (2025)
16
Ma e ial examined
Lec o ype
CHINA?•d yp ese edspecimen;designa edbyYen(1939);NHMW68037.
Pa alec o ype
CHINA?•1spec.(p ese edd y);NHMW68038.
O he ma e ial
CHINA•3specs(p ese edine hanol);Guangxi,ChongzuoCi y,LongzhouCoun y,ShuikouTown,
Poliu Village, ou le o an unde g ound i e in a ca e; 4 Ma . 2023; KIZ.240051 o KIZ.240053.
VIETNAM•11specs(p ese edine hanol);NinhBinh,Rese oi nea Dang,CucPhuongN.P.;
ZMB.114132•1spec.(p ese edine hanol);HoaBinh,TanLacdis ic ,NgocLy;ZMB.114150•
3 specs (p ese ed in e hanol); Bac Kan, Phu Luong, Yen Ninh, Dong Dang; ZMB.114171.
Desc ip ion
Adul shell (Fig. 7A–J) medium in size (26.60 ± 1.91 mm), hin bu s u dy, conical, oli e o g eenish
b own,wi h i e osixwho ls,apexacu e; eleoconchwho lssligh lyin la edespeciallyonbodywho l.
Shoulde angula , wi h h ee smoo h p ima y keels and one o wo weake smoo h seconda y keels
be weensu u eand i s keel.Some imes,oneaddi ionalweakkeelloca edbe ween i s andsecond/
secondand hi dkeels.Shellbasewi h h ee o i eweakkeels,andusuallyda ke  hano he pa so 
shell su ace; ape u e o a e, less han hal o shell in heigh , inne shell su ace bluish whi e, ou e lip
hin, smoo h and ounded. Inne laye s o adul columella lip hickened, whi e o o ange, ecu ed
columella lip pa ially o wholly co e ing umbilicus, some imes o ming a callus.
Ope culum (Fig. 7J–K) co neous, o a e, hin, yellow o o ange; ex e io su ace smoo h; in e io su ace
wi h smal nuclea egion (a ound 1/4 o inne ope cula egion), wi h shiny g ains o eins.
Radula (Fig.8)(n=4)cen al oo hwi honeb oadcen alden icleand i esmallsha pcuspsonei he 
side;la e al oo hwi honeb oadcen alden icleand i esmallsha pcuspsonei he side;inne ma ginal
oo hna ow,abou hal o la e al oo hinwid h,wi honeb oadcen alden icleand i e osixsmall
sha p cusps on ei he side; ou e ma ginal oo h wide, almos same as la e al oo h in wid h, wi h 21 o
26 small sha p cusps.
Fig. 8. SEM pho o o he adula o Bakyie aia polyzona a (F auen eld, 1862) gen. e comb. no .,
ZMB.114171.

ZHANG L.-J. e al., New genus o Vi ipa idae (Mollusca) om Sou h China
17
Rema ks
The newly collec ed specimens om Longzhou, China ha e a shell ha is e y simila o ha o
he lec o ype. The ype locali y o his species was no clea om he o iginal desc ip ion, and i
was conside ed simply as China (Yen 1939). Bakyie aia polyzona a can be dis inguished om mos
congene s, especially Bakyie aia ind apu a gen. e sp. no ., by i s medium-sized, hin conical shell
wi hin la edwho ls,mo e oundedou e lipanddis ibu ion.The ecommendedChinesenameis
多棱越田螺.
Habi a and dis ibu ion
Ri e s and lakes o Wes e n Guangxi o China and No he n Vie nam.
Bakyie aia subcos a a (G ay, 1833) gen. e comb. no .
Figs 7–8
Paludina subcos a aG ay,1833inG i i h&Pidgeon1833–1834:599,pl.36 ig.3(“China”).
Paludomus u aii G edle , 1889: 159 (“Peho, Kuang- ung”, e e s o Bei Ri e , Guangdong P o ince,
China). Syn. no .
Sino aia? u aii –Zilch1974:64,pl.7 ig.9.
Angulyag a subcos a a – Liu e al. 1995: 14.
Diagnosis
Adul shell medium o la ge (27.17 ± 3.64 mm), ounded and hin, p ima y keels weak, eleoconch wi h
weak p ima y keels; columella lip ecu ed, o ming callus co e ing umbilicus comple ely; ope culum
wi h small nuclea egion (a ound 1/5 o inne ope cula egion) wi h shiny g ains o eins; ou e ma ginal
oo h o adula wi h 29 o 43 small sha p cusps.
Ma e ial examined
Holo ype o Paludina subcos a a
CHINA•d yp ese edspecimen;NHMUK.2007.01.7.1.
Holo ype o Paludomus u aii
CHINA•d yp ese edspecimen;GuangdongP o ince,BeiRi e ;GFBI.22.
O he ma e ial
CHINA – Guangdong P o ince•4specs(p ese edine hanol);GuangzhouCi y,ZhengguoTown,
ZengRi e ,0.7mdeepunde  ock;25Sep.2019;Da-MingShileg.;ZMB.122694•3specs(p ese ed
d y); Guangzhou Ci y, Zhengguo Town, Zeng Ri e , 0.7 m deep unde ock; 25 Sep. 2019; Da-Ming
Shileg.;YHY-BAKSUB-ZG-1,YHY-BAKSUB-ZG-18,YHY-BAKSUB-ZG-28•3specs(p ese ed
d y); Zhaoqing Ci y, Xiaoxiang Town, Xi Ri e , unde ock; Aug. 2021; Ji-Lin Jiang leg.; KIZ.240084
•2specs(p ese edine hanol);QingyuanCi y,Shi anTown,BinRi e ,0.6mdeepon ock;24Sep.
2022;M Chenleg.;ZMB.122767•2specs(p ese edd y);QingyuanCi y,Shi anTown,BinRi e ,
0.6mdeepon ock;24Sep.2022;M Chenleg.;YHY-BAKSUB-ST-1,YHY-BAKSUB-ST-4•2specs
(p ese edine hanol);ZhaoqingCi y,HuaijiCoun y,SuiRi e ;15Sep.2022;ZMB.122766•1spec.
(p ese edine hanol);QingyuanCi y,Jin anTown,BinRi e ;10Oc .2022;KIZ.240085•4specs
(p ese ed in e hanol); Zhaoqing Ci y, Fengkai Coun y, Xi Ri e ; 19 Jan. 2024; Lejia Zhang leg.;
KIZ.240086.
Eu opean Jou nal o Taxonomy 1005: 1–64 (2025)
18
Desc ip ion
Adul shell (Fig. 9A–Q) medium o la ge (27.17 ± 3.64 mm), hin bu s u dy, ounded, oli e o da k
b own;wi h i e osixwho ls,apexacu e; eleoconchwho lsin la ed,some imes o mingshoulde ,
wi h h ee smoo h weak p ima y keels, wi h one o wo weake smoo h seconda y keels loca ed be ween
Fig. 9. Shells and ope culum o Bakyie aia subcos a a (G ay, 1833) gen. e comb. no . A–D. Holo ype,
NHMUK.2007.01.7.1, China, pho o by Jona han Able . E–H. KIZ.240084, Xiaoxiang Town,
Guangdong, China. I–J. ZMB.122766, Huaiji Coun y, Guangdong, China. K. KIZ.240085, Jin an
Town, Guangdong, China. L. Holo ype o Paludomus u aii, GFBI.22, Bei Ri e , Guangdong, China.
M–O. YHY-BAKSUB-ZG-18 (M), YHY-BAKSUB-ZG-28 (N), ZMB.122694 (O), Zhengguo,
Guangdong, China. P–Q. ZMB.122767, Shi an Town, Guangdong, China. R–S. Ope culum o
KIZ.240086; scale ba s = 1 cm, op le ba applies o all shells, sepa a e ba o ope culum R–S.
ZHANG L.-J. e al., New genus o Vi ipa idae (Mollusca) om Sou h China
19
su u eand i s keel,some imesoneaddi ionalweakkeelloca edbe ween i s andsecond/secondand
hi dkeels,baseo shellwi h h ee o i eweakkeels;ape u eo a e,la ge,a oundhal o shellinheigh ,
inne shell su ace bluish whi e o pu ple, ou e lip hin, always wi h a ela i ely ounded pe iphe y,
columella lip co e ed wi h hick inne shell laye s, always whi ish o ange o pink, adul columella lip
ecu ed, o ming a e y na ow c escen -shaped callus o wi hou his s uc u e, columella lip o his
s uc u e comple ely co e ing umbilicus.
Ope culum (Fig. 9R–S) co neous, o a e, hin, yellow; ex e io su ace smoo h; in e io su ace wi h
nuclea egion small, a ound 1/5 o inne ope cula egion, wi h shiny g ains o eins.
Radula (Fig.10)(n=4)cen al oo hwi honeb oadcen alden icleand i e osixsmallsha pcusps
onei he side;la e al oo hwi honeb oadcen alden icleand i esmallsha pcuspsonei he side;
inne ma ginal oo h na ow, abou hal o la e al oo h in wid h, wi h one b oad cen al den icle and ou
small sha p cusps on ei he side; ou e ma ginal oo h wide, almos same as la e al oo h in wid h, wi h
29 o 43 small sha p cusps.
Rema ks
Bakyie aia subcos a a can be easily dis inguished om mos congene s based on he ounded hin shell
wi h weak p ima y keels. I can be di e en ia ed om he mo phologically simila species Bakyie aia
chenghuang gen. e sp. no . based on he la ge numbe o small cusps on he ou e ma ginal ee h o
he adula (29–43 s 15–16). Paludomus u aii was desc ibed based on a ju enile shell o his species,
and i is he e placed in synonymy o B. subcos a a. The ecommended Chinese name is 次肋越田螺.
Habi a and dis ibu ion
Rock subs a e in he shallow wa e o i e s o Guangdong P o ince, China.
Bakyie aia boe ge i (Möllendo , 1890) gen. e comb. no .
Figs 11–12
Paludina boe ge iMöllendo ,1890inHeude1882–1890:177,pl.10 ig.6(“Hai-nan”,Hainan
P o ince, China).
Vi ipa us (Taia) boe ge i – Zilch 1938: 131.
Angulyag a boe ge i –Yen1939:38,pl.3 ig.17.—Liue al. 1995: 14.
Fig. 10. SEM pho o o he adula o Bakyie aia subcos a a (G ay, 1833) gen. e comb. no ., ZMB.122766.
Eu opean Jou nal o Taxonomy 1005: 1–64 (2025)
20
Diagnosis
Adul shellmedium ola ge(26.76±6.67mm),s u dy,conical,who lsno in la ed,p ominen keel
abo e su u e; columella lip ecu ed, o ming callus co e ing umbilicus comple ely; ope culum wi h
small nuclea egion (a ound 1/5 o inne ope cula egion) wi h shiny g ains o eins; ou e ma ginal
oo h o adula wi h 23 o 24 small sha p cusps.
Ma e ial examined
Pa a ypes
CHINA•5specs(p ese edd y);HainanIsland; O.Bö ge leg.;SMF.7651•2specs(p ese edd y);
Hainan Island; O. . Möllendo leg.; SMF.7653.
O he ma e ial
CHINA – Guangxi•1spec.(p ese edine hanol);NanningCi y,Dawang anRese oi ;Aug.2019;
KIZ.240087•5specs(p ese edine hanol);BaiseCi y,YoujiangDis ic ,WulaVillage,Sanglu,0.5m
deepon ockso c eeks;17Oc .2019;Jian-ZhuoLileg.;ZMB.122687•3specs(p ese edd y);same
da aas o p eceding;YHY-BAKBOE-SL-1,YHY-BAKBOE-SL-2,YHY-BAKBOE-SL-18•5specs
(p ese ed in e hanol); Guigang Ci y, Gangnan Dis ic , Ba ang Village, a pond nea Yapengling, 0.7 m
deepon ocks;31Oc .2020;Xia-MingLileg.,ZMB.122759•1spec.(p ese edd y);sameda aas o 
p eceding;YHY-BAKBOE-BT-1•1spec.(p ese edd y);ChongzuoCi y,DaxinCoun y,LeipingTown,
HeishuiRi e ,1mdeepon ocks;27Ma .2021;Yi-WenLiangleg.;ZMB.122690•4specs(p ese ed
d y); same da a as o p eceding; YHY-BAKBOE-HSH-21, YHY-BAKBOE-HSH-15, YHY-BAKBOE-
HSH-9,YHY-BAKBOE-HSH-3•4specs(p ese edine hanol);ChongzuoCi y,JiangzhouDis ic ,
Tuozao Village, Zuo Ri e , 0.5 o 1.5 m deep on ocks; 27 Ma . 2021; Yi-Wen Liang leg.; ZMB.122758
•3specs(p ese edd y);sameda aas o p eceding;YHY-BAKBOE-JZ-15,YHY-BAKBOE-JZ-14,
YHY-BAKBOE-JZ-1•2specs(p ese edine hanol);NanningCi y,MashanCoun y,Shi angVillage,
0.3mdeeponsand/mudsubs a ein hewe land;19Ma .2022;Le-JiaZhangleg.;ZMB.122734•
3 specs (p ese ed in e hanol); Pingguo Ci y, Yihao Village, Bujian Ri e , 0.5 o 1 m deep on mud
subs a e; 19 Ma . 2022; Le-Jia Zhang leg.; ZMB.122720. – Hainan P o ince •2specs(p ese ed
d y);HainanIsland,Je schinleg.;SMF.88400•5specs(p ese edine hanol);Haikou,we landsnea 
Yangshan Rese oi ; 20 Oc . 2019; bough by Sheng-Zhuo Huang on local ma ke ; ZMB.122710.
Desc ip ion
Adul shell (Fig. 11A–R) medium o la ge (26.76 ± 6.67 mm), hin o hick, s u dy, conical, oli e b own
oda kb own,wi h i e osixwho ls,apexacu e; eleoconchwho lsno in la ed,wi h h eesmoo h
p ima ykeels,wi hone o woweake smoo hseconda ykeelsloca edbe weensu u eand i s keel,
some imesoneaddi ionalweakkeelloca edbe ween i s andsecond/secondand hi dkeels,baseo 
shellwi hone o i eweakkeels,allkeelsusuallyda ke  han es o shellsu ace,p ominen keelabo e
su u e; ape u e o a e, less han hal o shell in heigh , inne shell su ace bluish whi e, o ange- ed o
da k pu ple, ou e lip hin, always wi h an ob ious angula pe iphe y, columella lip co e ed wi h hick
inne shell laye s, whi e o o ange- ed, adul columella lip ecu ed, o ming a ela i ely wide c escen -
shaped callus, comple ely co e ing umbilicus.
Ope culum (Fig. 11S–T) co neous, o a e, ela i ely hick, da k ed; ex e io su ace smoo h; in e io
su ace wi h small nuclea egion, a ound 1/5 o inne ope cula egion, wi h shiny g ains o eins o
smoo h glossy su ace.
Radula (Fig.12)(n=3)cen al oo hwi honeb oadcen alden icleand i e osixsmallsha pcusps
onei he side;la e al oo hwi honeb oadcen alden icleand ou  o i esmallsha pcuspsonei he 
side; inne ma ginal oo h na ow, abou hal o la e al oo h in wid h, wi h one b oad cen al den icle and
ZHANG L.-J. e al., New genus o Vi ipa idae (Mollusca) om Sou h China
21
Fig. 11. Shells and ope culum o Bakyie aia boe ge i (Möllendo , 1890) gen. e comb. no .
A–C. Pa a ype, SMF.7651, Hainan, China. D. Pa a ype, SMF7653, Hainan, China. E. SMF.88400,
Hainan, China. F–I. ZMB.122710, Haikou, Hainan, China. J. KIZ.240087, Nanning, Guangxi,
China. K. ZMB.122687, Sanglu, Guangxi, China. L. YHY-BAKBOE-JZ-1, Jiangzhou, Guangxi,
China. M. ZMB.122720, Pingguo, Guangxi, China. N. ZMB.122758, Jiangzhou, Guangxi, China.
O–Q. ZMB.122690 (O), YHY-BAKBOE-HSH-9 (P–Q), Leiping, Guangxi, China. R. ZMB.122734,
Mashan Coun y, Guangxi, China. S–T. Ope culum o KIZ.240087. Scale ba s = 1 cm, op le ba applies
o all shells, sepa a e ba o ope culum K–L.

Eu opean Jou nal o Taxonomy 1005: 1–64 (2025)
22
ou small sha p cusps on ei he side; ou e ma ginal oo h wide, almos same as la e al oo h in wid h,
wi h 23 o 24 small sha p cusps.
Rema ks
The newly collec ed specimens om Haikou Ci y o Hainan ha e a shell ha is e y simila o ha o he
ype specimens. Bakyie aia boe ge i can be dis inguished om mos congene s based on i s medium-
sized ola geconicalshellwi hnon-in la edwho ls,ands onglykeeledsu u e.Somesmallindi iduals
o his species may be con used wi h Bakyie aia jingweiae Yen, Zhang & on Rin elen gen. e sp. no .
based on shell mo phology, bu di e by possessing ewe cusps on he ou e ma ginal oo h (23–24 s
26–28). The COI ee suppo s B. boe ge i and B. jingweiae as dis inc species, and he wo species a e
allopa ic (Fig. 1). The ecommended Chinese name is 伯氏越田螺.
Habi a and dis ibu ion
Mud, sand and ock subs a e in he shallow wa e o i e s, lakes and we lands o no he n Hainan and
sou he n Guangxi, China.
Bakyie aia duchie i (Fische , 1898) gen. e comb. no .
Fig. 13
Paludina duchie iFische ,1898:330–331,pl.18 igs48–49(“Ri .Song-KI-Kong,àDéo-Ca ”,KỳCùng
Ri e ,a ĐèoKhách,LạngSơn,Vie nam).
Angulyag a duchie i – S a oboga o 1970: 202. — Thanh e al.2004:3–4, ig.5.
Angulyag a boe ge i –Madsen&Hung2014:109, igs2(10le ),(10middle)[nonPaludina duchie i
Fische , 1898].
Diagnosis
Shellla ge(32.88±1.00mm), hickands u dy,who lsin la ed,wi hna owangula shoulde ;
columella lip ecu ed, o ming callus co e ing umbilicus comple ely; ope culum wi h small nuclea
egion (a ound 1/5 o inne ope cula egion) wi h shiny g ains o eins.
Fig. 12. SEM pho o o he adula o Bakyie aia boe ge i (Möllendo , 1890) gen. e comb. no .,
ZMB.122710.
ZHANG L.-J. e al., New genus o Vi ipa idae (Mollusca) om Sou h China
23
Ma e ial examined
Holo ype
VIETNAM•d yp ese edspec.;LạngSơn,ĐèoKhách,KỳCùngRi e ;Juille Bille leg.;MNHN-
IM-2000-4486.
Fig. 13. Shells and ope culum o Bakyie aia duchie i (Fische , 1898) gen. e comb. no . A–B. Holo ype,
MNHN-IM-2000-4486 (h p://coldb.mnhn. /ca alognumbe /mnhn/im/2000-4486, c edi :
MNHN-B aban D.-2005),ĐèoKhách,LạngSơn,Vie nam.C–D. SMF.48146, Tonkin, Vie nam.
E–L. KIZ.240049–240050, Longzhou, Guangxi, China. M–N. Ope culum o KIZ.240049. Scale ba s =
1 cm, op le ba applies o all shells, sepa a e ba o ope culum M–N.
Eu opean Jou nal o Taxonomy 1005: 1–64 (2025)
24
O he ma e ial
CHINA•2specs(p ese edine hanol);Guangxi,ChongzuoCi y,LongzhouCoun y,ShuikouTown,
Poliu Village, Shuikou Ri e , 0.5 m deep on mud subs a e; 4 Ma . 2023; KIZ.240049, KIZ.240050.
VIETNAM•1spec.(p ese edd y);“Tonkin”(Hanoi);O. .Möllendo leg.;SMF.48146
Desc ip ion
Adul shell (Fig. 13A–K) la ge (32.88 ± 1.00 mm), hick, s u dy, conical, oli e o oli e b own, wi h
sixwho ls,apexacu e; eleoconchwho lssligh lyin la ed,wi h h eesmoo hp ima ykeels,wi hone
o woweake smoo hseconda ykeelsloca edbe weensu u eand i s keel,some imesoneaddi ional
weakkeelloca edbe ween i s andsecond/secondand hi dkeels,baseo shellwi hone o h eeweak
keels, all keels usually da ke han es o shell su ace; ape u e o a e, less han hal o shell in heigh ,
inne shell su ace bluish whi e, ou e lip hin, always wi h an ob ious angula pe iphe y, columella
lip co e ed wi h hick inne shell laye s, whi e o o ange ed, adul columella lip ecu ed, some imes
o ming a na ow o wide c escen -shaped callus, his s uc u e o columella lip comple ely co e ing
umbilicus.
Ope culum (Fig. 13M–N) co neous, o a e, ela i ely hick, da k ed; ex e io su ace smoo h; in e io
su ace wi h small nuclea egion, a ound 1/5 o inne ope cula egion, wi h shiny g ains o eins o
smoo h glossy su ace.
Radula cu en ly unknown.
Rema ks
The newly collec ed specimens in Longzhou o China ha e a shell ha is e y simila o ha o he
holo ype, and he collec ing si e is close o he ype locali y on he same i e basin. Bakyie aia duchie i
can be easily dis inguished om he o he species o Bakyie aia gen. no . based on i s la ge s u dy shell
andin la edbodywho lwi hshoulde .The ecommendedChinesenameis杜氏越田螺.
Habi a and dis ibu ion
Sand subs a e and ocks in he shallow wa e o i e s and lakes o Wes e n Guangxi, China and
No he n Vie nam.
Bakyie aia guangdungensis (Kobel , 1906) gen. e comb. no .
Figs 14–15
Vi ipa a heudei a . guangdungensisKobel ,1906inKobel 1906–1909:124–125,pl.21 igs12,14
(“P o inz Guang-dung”, Guangdong P o ince, China).
Dac ylochalamys guangdungensis – P ashad 1928: 169.
Vi ipa a polyzona a–Yen1939:37,pl.3 igs14–15[nonVi ipa a polyzona a F auen eld, 1862].
Angulyag a guangdungensis – Yen 1943: 128.
Sino aia polyzona a–Zilch1955:77, ig.52.—Qiane al. 2014 [non Vi ipa a polyzona a F auen eld,
1862].
Taia polyzona a – Johnson 1973: 120, 122. — Palmie i e al. 1980: 40. — Ng 1992: 125. — Chan 1996:
184–185, ig.2.—Maassen2001:44.—Clemen se al. 2006: 144. — Tan & Woo 2010: 26. —
Yeo & Chia 2010: 32. — Tan e al. 2012: 133 [non F auen eld 1862].
Vi ipa us polyzona a – Ng e al. 1993: 20 [non Vi ipa a polyzona a F auen eld, 1862].
Bellamya heudei guangdungensis – Shea 1994: 3–11.
Sino aia guangdungensis – Ng e al. 2014: 401–408
ZHANG L.-J. e al., New genus o Vi ipa idae (Mollusca) om Sou h China
25
Angulyag a polyzona a –Madsen&Hung2014:110, ig.2(11)[nonVi ipa a polyzona a F auen eld,
1862].
Diagnosis
Shell ela i elysmall(20.87±0.93mm),who lsin la ed;columella lip ecu ed, o mingcallus
co e ing umbilicus comple ely; ope culum wi h small nuclea egion (a ound 1/5 o inne ope cula
egion) wi h shiny g ains o eins; ou e ma ginal oo h o adula wi h 28 o 29 small sha p cusps.
Ma e ial examined
Lec o ype
CHINA•d yp ese edspec.;GuangdongP o ince;O. .Möllendo leg.;designa edbyYen(1939);
SMF.38890.
Pa alec o ypes
CHINA•4specs(p ese edd y);GuangdongP o ince;O. .Möllendo leg.;SMF.38822.
O he ma e ial
CHINA – Guangdong P o ince•1spec.(p ese edd y);GuangzhouCi y,BaiyunMoun ain,c eeks
inTaohuajianValley,mudsubs a e,onbank;25Feb.2022;Le-JiaZhangleg.;KIZ.240054•1spec.
(p ese ed in e hanol); Qingyuan Ci y, Lianzhou Coun y, Xian Town; 18 No . 2019; bough by Hai-Ling
Zhangonlocalma ke ;ZMB.122693-1•3specs(p ese edine hanol);ZhaoqingCi y,BohaiPa k,
Qixing Lake, 0.1–0.2 m deep unde ocks; 19 Jan. 2020; Ji-Lin Jiang leg.; ZMB.122760. – Guangxi•
4 specs (p ese ed in e hanol); Guilin Ci y, Yangshuo Town; 14 Ma . 2022; bough on local ma ke by
Le-JiaZhang;ZMB.122727•2specs(p ese edine hanol);LaibinCi y,HeshanCoun y,RenyiVillage,
Huaiji Rese oi , 0.5 m deep mud subs a e; 7 Jan. 2020; Ri-Bao Huang leg.; KIZ.240088.
SINGAPORE•3specs(p ese edine hanol);Canala Sp inglea Rd;ZMB.192077.
Desc ip ion
Adul shell (Fig. 14A–H, K–R) ela i ely small o he genus (20.87 ± 0.93 mm), hin bu s u dy, conical,
oli e ooli eb own,wi h i e osixwho ls,apexacu e; eleoconchwho lsin la ed,some imeswi h
shoulde , wi h h ee smoo h p ima y keels, wi h one o wo weake smoo h seconda y keels loca ed
be weensu u eand i s keel,some imesoneaddi ionalweakkeelloca edbe ween i s andsecond/
secondand hi dkeels,baseo shellwi h h ee o i eweakkeels,allkeelsusuallyda ke  han es o 
shell su ace; ape u e o a e, less han hal o shell in heigh , inne shell su ace bluish whi e, ou e lip
hin, always wi h an ob ious angula pe iphe y, columella lip co e ed wi h hick inne shell laye s, whi e
o o ange, adul columella lip ecu ed, some imes o ming a ela i ely na ow c escen -shaped callus,
his s uc u e o columella lip comple ely co e ing umbilicus.
Ope culum (Fig. 14I–J) co neous, o a e, ela i ely hin, yellow o o ange; ex e io su ace smoo h;
in e io su ace wi h small nuclea egion, a ound 1/5 o inne ope cula egion, wi h shiny g ains o eins
o smoo h glossy su ace.
Radula (Fig.15)(n=3)cen al oo hwi honeb oadcen alden icleand ou  o i esmallsha pcusps
on ei he side; la e al oo h wi h one b oad cen al den icle and ou small sha p cusps on ei he side;
inne ma ginal oo h na ow, abou hal o la e al oo h in wid h, wi h one b oad cen al den icle and h ee
o ou small sha p cusps on ei he side; ou e ma ginal oo h wide, almos same wid h as la e al oo h,
wi h 28 o 30 small sha p cusps.
Eu opean Jou nal o Taxonomy 1005: 1–64 (2025)
32

ZHANG L.-J. e al., New genus o Vi ipa idae (Mollusca) om Sou h China
33
Desc ip ion
Adul shell (Fig. 18A–AA) small o medium (25.53 ± 3.72 mm), hin o hick, s u dy, conical, oli e o
da koli eb own,wi h i e osixwho ls,apexacu e; eleoconchwho lssome imeswi hou shoulde ,
some imesin la edandwi hshoulde ,wi h h eesmoo hp ima ykeels,wi hone o woweake smoo h
seconda ykeelsloca edbe weensu u eand i s keel,some imesoneaddi ionalweakkeelloca ed
be ween i s andsecond/secondand hi dkeels,baseo shellwi h wo o i eweakkeels,allkeels
usually da ke han es o shell su ace; ape u e o a e, less han hal o shell in heigh , inne shell
su ace bluish whi e, ou e lip hin, always wi h an angula pe iphe y, columella lip co e ed wi h hick
inne shell laye s, whi e o o ange, adul columella lip ecu ed, some imes o ming a a he wide
c escen -shaped callus, his s uc u e o columella lip comple ely co e ing umbilicus.
Ope culum (Fig. 18AB–AG) co neous, o a e, ela i ely hin, yellow o o ange; ex e io su ace smoo h;
in e io su ace wi h small nuclea egion, a ound 1/4 o 1/5 o inne ope cula egion, wi h shiny g ains
o eins o smoo h glossy su ace.
Radula (Fig.19)(n=6)cen al oo hwi honeb oadcen alden icleand ou  o i esmallsha pcusps
on ei he side; la e al oo h wi h one b oad cen al den icle and ou small sha p cusps on ei he side;
inne ma ginal oo h na ow, abou hal o la e al oo h in wid h, wi h one b oad cen al den icle and h ee
o i esmallsha pcuspsonei he side;ou e ma ginal oo hwide,almos samewid hasla e al oo h,
wi h 18 o 25 small sha p cusps.
Fig. 18 (p eceding page). Shells and ope cula o Bakyie aia wilhelmi (Yen, 1939) gen. e comb. no .
A–C. Holo ype, SMF.40812, Guangdong, China. D. ZMB.122686-2, Jiangxikong, Guangdong,
China. E. ZMB.122696, Heyuan, Guangdong, China. F–I. ZMB.122686-1, Jiangxikong, Guangdong,
China. J. ZMB.122689, Fogang, Guangdong, China. K–L. ZMB.122688, Zhuhai, Guangdong,
China. M. ZMB.122692, Shenzhen, Guangdong, China. N–O. ZMB.122685, Liuzhou, Guangxi,
China. P. ZMB.122751, Zhanjiang, Guangdong, China. Q–R. ZMB.122729, Gejiu, Yunnan, China.
S. ZMB.122737-3, Ledong, Hainan, China. T. ZMB.122709, Yangshan, Hainan, China. U. ZMB.114206,
Ha Quang, Vie nam. V. ZMB.122770, Niuyun, Guangxi, China. W. P1010042, Niuyun, Guangxi,
China. X. ZMB.122760-1, Zhaoqing, Guangdong, China. Y. YHY-BAKWIL-HS-2, Heshan, Guangxi,
China. Z. 122693-2, Lianzhou, Guangdong, China. AA. ZMB.122704, Jieyang, Guangdong, China.
AB–AC. Ope culum o ZMB.122685. AD–AE. Ope culum o ZMB.122686-1. AF–AG. Ope culum
o ZMB.114206. Scale ba s = 1 cm, op le ba applies o all shells, sepa a e ba o ope cula AB–AG.
Fig. 19. SEM pho o o he adula o Bakyie aia wilhelmi (Yen, 1939) gen. e comb. no ., ZMB.122686.
Eu opean Jou nal o Taxonomy 1005: 1–64 (2025)
34
Rema ks
The newly collec ed specimens om Qingyuan Ci y in Guangdong ha e a shell ha is e y simila o
ha o he holo ype. Bakyie aia wilhelmi is he mos common species o his genus. I has he wides
dis ibu ion ange, and displays ex emely high polymo phism o shell cha ac e s. This species was
also ound sympa ically wi h o he species o Bakyie aia gen. no . (B. boe ge i, B. duchie i and
B. guangdungensis) in some locali ies (such as Qixing Lake in Zhaoqing Ci y in Guangdong and Honghu
Lake in Ledong Coun y o Hainan). The mo pho ype ha is simila o he holo ype can be dis inguished
ommos congene sbasedoni scompa a i elysmallconicalshellwi hnon-in la edwho lsand
mode a ely keeled su u e. Howe e , some mo pho ypes canno be dis inguished om Bakyie aia
guangdungensis based on hei shell mo phology. The COI ee suppo s he s a us o B. wilhelmi and
B. guangdungensis as dis inc species ha a e qui e dis an ly ela ed. In addi ion, B. wilhelmi always
has ewe small cusps on he ou e ma ginal ee h han B. guangdungensis (18–25 s 28–30). DNA
ba coding is conside ed necessa y o unambiguously di e en ia e B. wilhelmi and B. guangdungensis.
The ecommended Chinese name is 威廉越田螺.
Habi a and dis ibu ion
Mud, sand and ock subs a e in he shallow wa e o i e s, we lands and lakes in Guangdong, Hainan,
Guangxi and sou he n Yunnan P o ince, China and no he n Vie nam. In oduced in o Chengdu Ci y,
Sichuan P o ince, China and no he n Bo neo, Malaysia.
Bakyie aia chenghuang Zhang, Yen & on Rin elen gen. e sp. no .
u n:lsid:zoobank.o g:ac :24C0929E-836B-4AC5-9829-294C4C6345B4
Figs 20–21
Diagnosis
Shell medium in size (23.70 ± 5.37 mm), ounded and hin, eleoconch wi h weak p ima y keels;
columella lip ecu ed, o ming callus co e ing umbilicus comple ely; ope culum wi h small nuclea
egion (a ound 1/5 o inne ope cula egion) wi h shiny g ains o eins; ou e ma ginal oo h o adula
wi h 15 o 16 small sha p cusps.
E ymology
‘Cheng Huang’ means ‘ci y god’ in Chinese, e e ing o he ype locali y in Chenghuang Town. Fo med
as a noun in apposi ion. The ecommended Chinese name is 城隍越田螺.
Ma e ial examined
Holo ype
CHINA•spec.p ese edine hanol;Guangxi,YulinCi y,ChenghuangTown,a i e nea Shi en ou
Moun ain, 0.5 m deep on ock; 13 Ma . 2022; Le-Jia Zhang leg.; KIZ.240001.
Pa a ypes
CHINA•4specs(p ese edine hanol);sameda aas o holo ype;KIZ.240002 oKIZ.240005•2specs
(p ese edine hanol);sameda aas o holo ype;ZMB.122722-1•4specs(p ese edine hanol);same
da a as o holo ype; KIZ.240089.
O he ma e ial
1 spec. (p ese ed d y); same da a as o holo ype; YHY-BAKCHE-CH-1.
Desc ip ion
Adul shell (Fig. 20A–E, H–J) medium in size (23.70 ± 5.37 mm), hin bu s u dy, b oadly conical o
ounded,oli e ooli eb own,wi h i ewho ls,apexacu e; eleoconchwho lsin la ed,wi h h eeweak,
ZHANG L.-J. e al., New genus o Vi ipa idae (Mollusca) om Sou h China
35
smoo hp ima ykeels,wi hone o woweake smoo hseconda ykeelloca edbe weensu u eand i s 
keel,some imesoneaddi ionalweakkeelloca edbe ween i s andsecond/secondand hi dkeels,
baseo shellwi h h ee o i eweakkeels;ape u eo a e,la ge,a oundhal o shellinheigh ,inne 
Fig. 20. Shells and ope culum o Bakyie aia chenghuang Zhang, Yen & on Rin elen gen. e sp. no .
A–D. Holo ype, KIZ.240001, Chenghuang Town, Guangxi, China. E. Pa a ype, ZMB.122722-1,
Chenghuang Town, Guangxi, China. F–G. Ope culum o KIZ.240002. H–J. YHY-BAKCHE-CH-1,
Chenghuang Town, Guangxi, China. Scale ba s = 1 cm, op le ba applies o all shells, sepa a e ba o
ope culum F–G.
Eu opean Jou nal o Taxonomy 1005: 1–64 (2025)
36
shell su ace bluish whi e o pu ple whi e, ou e lip hin, always wi h a ela i ely ounded pe iphe y,
columella lip co e ed wi h hick inne shell laye s, always whi e, adul columella lip ecu ed, o ming
a e y na ow c escen -shaped callus o wi hou his s uc u e, columella lip o his s uc u e comple ely
co e ing umbilicus.
Ope culum (Fig. 20F–G) co neous, o a e, hin, yellow; ex e io su ace smoo h; in e io su ace wi h
small nuclea egion, a ound 1/5 o inne ope cula egion, wi h shiny g ains o eins.
Radula (Fig.21)(n=2)cen al oo hwi honeb oadcen alden icleand ou  o i esmallsha pcusps
on ei he side; la e al oo h wi h one b oad cen al den icle and ou small sha p cusps on ei he side;
inne ma ginal oo h na ow, abou hal wid h o la e al oo h, wi h one b oad cen al den icle and h ee
small sha p cusps on ei he side; ou e ma ginal oo h wide, almos same wid h as la e al oo h, wi h 15
o 16 small sha p cusps.
Rema ks
Bakyie aia chenghuang gen. e sp. no . can be easily dis inguished om mos congene s based on i s
hin, ounded shell wi h a wide ape u e and a e y small ope culum. I can be di e en ia ed om he
mo phologically simila species Bakyie aia subcos a a based on he smalle numbe o small cusps on
ou e ma ginal ee h o he adula (15–16 s 29–43).
Habi a and dis ibu ion
Only known om he ype locali y in Guangxi, China, whe e i occu s on ocky subs a es in he shallow
wa e .
Bakyie aia jingweiae Yen, Zhang & on Rin elen gen. e sp. no .
u n:lsid:zoobank.o g:ac :7D19E72D-8C68-4AB8-A41B-70CA77A28C0C
Figs 22–23
Diagnosis
Shellsmall omediuminsize(26.79±5.37mm),s u dy,conical,who lsno in la ed,p ominen keel
abo e su u e; columella lip ecu ed, o ming callus co e ing umbilicus comple ely; ope culum wi h
small nuclea egion (a ound 1/4 o inne ope cula egion) wi h shiny g ains o eins; ou e ma ginal
oo h o adula wi h 26–28 small sha p cusps; dis ibu ed in no he nVie nam, ac oss he bo de in
wes e n Guangxi and in wes e n Hainan Island, China.
Fig. 21. SEM pho o o he adula o Bakyie aia chenghuang Zhang, Yen & on Rin elen gen. e sp. no .,
pa a ype, ZMB.122722-1.
ZHANG L.-J. e al., New genus o Vi ipa idae (Mollusca) om Sou h China
37
E ymology
Named a e ‘Jingwei’, a Chinese my hical bi d in o which he Flame Empe o Yandi’s daugh e Nüwa,
whod ownedin hesea,was ans o med,andwhich ied o illup hesea.The ecommendedChinese
name is 精卫越田螺.
Ma e ial examined
Holo ype
CHINA•spec.p ese edine hanol;Guangxi,BaiseCi y,JingxiCoun y,XinjingTown,Long an
We land, 0.1 m on ock; 30 Dec. 2019; Hua-Pu Huang leg.; KIZ.240006.
Pa a ypes
CHINA – Guangxi•4specs(p ese edine hanol);sameda aas o holo ype;KIZ.240056 o
KIZ.240059•5specs.(p ese edine hanol);sameda aas o holo ype;ZMB.122701•1spec.
(p ese ed in e hanol); Guangxi, Baise Ci y, c eek nea Nawang Village, 0.1 m deep on sand o ock
subs a e; 20 Ma . 2022; Le-Jia Zhang leg.; KIZ.240007. – Hainan P o ince•3specs(p ese edin
e hanol); Ledong Li Au onomous Coun y, Honghu Village, Honghu Lake, 0.1 m deep sand subs a e;
9Ma .2022;Le-JiaZhangleg.;ZMB.122737-1,ZMB.122737-2,ZMB.122737-4•1spec.(p ese edin
e hanol); Changjiang Li Au onomous Coun y, Changhua Town, Sanjia Village, Changhua Ri e , 0.5 m
deep on ock; 9 Ma . 2022; Le-Jia Zhang leg.; KIZ.240008.
VIETNAM • 4 specs (p ese ed in e hanol); Cao Bang, i e  Sông Bằng; 2000; IEBR leg.;
MZMB.114431•3specs(p ese edine hanol);CaRi e d ainage,Ho-Chi-Minh-Highwaybe ween
YenCa andTháiHòa,NghêAn;21–31Oc .2006;F ankKöhle leg.;ZMB.114407•2specs(p ese ed
ine hanol);BaBeLakes,BacKan;2001;IEBRleg.;ZMB.114430•10specs(p ese edine hanol);Ba
Be Lakes, Bac Kan, e y po ; 21 Ma . 2006; F ank Köhle leg.; ZMB.114189.
O he ma e ial
CHINA•3specs(p ese edd y);sameda aas o holo ype;YHY-BAKDUC-JX-1,YHY-BAKDUC-
JX-10, YHY-BAKDUC-JX-15.
Desc ip ion
Adul shell (Fig. 22A–N) small o medium (26.79 ± 5.37 mm), hin o hick, s u dy, conical, oli e o
oli eb own,wi h i e osixwho ls,apexacu e; eleoconchwho lssligh lyin la ed,wi h h eesmoo h
p ima ykeels,wi hone o woweake smoo hseconda ykeelsloca edbe weensu u eand i s keel,
some imesoneaddi ionalweakkeelloca edbe ween i s andsecond/secondand hi dkeels,baseo 
shell wi h one o h ee weak keels, all keels usually da ke han es o shell su ace; ape u e o a e, less
han hal o shell in heigh , inne shell su ace bluish whi e, ou e lip hin, always wi h a s ong angula
pe iphe y, columella lip co e ed wi h hick inne shell laye s, whi e o o ange- ed, adul columella lip
ecu ed, some imes o ming a na ow o wide c escen -shaped callus, his s uc u e o columella lip
comple ely co e ing umbilicus.
Ope culum (Fig. 22O–P) co neous, o a e, ela i ely hick, da k ed; ex e io su ace smoo h; in e io
su ace wi h small nuclea egion, a ound 1/4 o inne ope cula egion, wi h shiny g ains o eins o
smoo h glossy su ace.
Radula (Fig. 23) (n = 4) cen al oo h wi h one b oad cen al den icle and ou o six small sha p cusps
onei he side;la e al oo hwi honeb oadcen alden icleand i e osixsmallsha pcuspsonei he 
side; inne ma ginal oo h na ow, abou hal o la e al oo h in wid h, wi h one b oad cen al den icle
and ou  o i esmallsha pcuspsonei he side;ou e ma ginal oo hwide,almos samewid hasla e al
oo h, wi h 26 o 28 small sha p cusps.

Eu opean Jou nal o Taxonomy 1005: 1–64 (2025)
38
Fig. 22. Shells and ope culum o Bakyie aia jingweiae Yen, Zhang & on Rin elen gen. e sp. no . A–D.
Holo ype, KIZ.240006, Jingxi, Guangxi, China. E. Pa a ype, KIZ.240007, Nawang, Guangxi, China.
F–H. Pa a ypes, ZMB.122737-1, ZMB.122737-2, ZMB.122737-4, Ledong, Hainan, China. I. Pa a ype,
ZMB.114407,NghêAn,Vie nam.J. Pa a ype, KIZ.240008, Changhua, Hainan, China. K–L. Pa a ype,
ZMB.114189, Ba Be Lakes, Bac Kan, Vie nam. M–N. Pa a ypes, ZMB.114430, Ba Be Lakes, Bac Kan,
Vie nam. O–P. Ope culum o ZMB.122701. Scale ba s = 1 cm, op le ba applies o all shells, sepa a e
ba o ope culum O–P.
ZHANG L.-J. e al., New genus o Vi ipa idae (Mollusca) om Sou h China
39
Rema ks
Bakyie aia jingweiae gen. e sp. no . is mos simila o Bakyie aia boe ge i based on shell mo phology,
bu di e s in possessing a smalle shell and g ea e numbe o small cusps on ou e ma ginal ee h (26–28
s 23–24). The dis ibu ion a eas o hese wo species a e close o each o he bu no o e lapping nea
he bo de be ween Guangxi, China and Vie nam.
Habi a and dis ibu ion
Sand subs a e and ocks in he shallow wa e o i e s and lakes o wes e n Guangxi and wes e n Hainan
P o ince, China and no he n Vie nam.
Bakyie aia liusanjieae Zhang, Yen & on Rin elen gen. e sp. no .
u n:lsid:zoobank.o g:ac :16B296D0-9634-4460-B0EB-770DB94244DD
Figs 24–25
Diagnosis
Shellmediuminsize(25.69±3.35mm),s u dy,conicalshellwi hlowe spi e,who lsno in la ed;
columella lip ecu ed, o ming callus co e ing umbilicus comple ely; ope culum wi h small nuclea
egion (a ound 1/4 o inne ope cula egion) wi h shiny g ains o eins; ou e ma ginal oo h o adula
wi h 26–27 small sha p cusps; dis ibu ed in no he n Guangxi, China.
E ymology
Named a e ‘Liu Sanjie’ (刘三姐, Liu amily’s hi d daugh e ), a legenda y olk singe o he Zhuang
people in Guangxi. The ype locali y o his species is close o he home own. The ecommended Chinese
name is 歌仙越田螺.
Ma e ial examined
Holo ype
CHINA•spec.p ese edine hanol;Guangxi,HechiCi y,LuochengMulaoAu onomousCoun y,nea 
Hemu Town, i e in Xinhui Village, 0.5 m deep on dam; 16 Ma . 2022; Le-Jia Zhang leg.; KIZ.240009.
Pa a ypes
CHINA – Guangxi•2specs(p ese edine hanol);sameda aas o holo ype;KIZ.240010,KIZ.240011
•2specs(p ese edine hanol);sameda aas o holo ype;ZMB.122722-2•2specs(p ese edd y);
Fig. 23. SEM pho o o he adula o Bakyie aia jingweiae Yen, Zhang & on Rin elen gen. e sp. no .,
ZMB.122701.
Eu opean Jou nal o Taxonomy 1005: 1–64 (2025)
40
Fig. 24. Shells and ope culum o Bakyie aia liusanjieae Zhang, Yen & on Rin elen gen. e sp. no .
A–D. Holo ype, KIZ.240009, Xinhui, Guangxi, China. E. Pa a ype, KIZ.240092, Liusanjie Town,
Guangxi, China. F–G. Pa a ypes, KIZ.240010, KIZ.240011, Xinhui, Guangxi, China. H. YHY-BAKLIU-
XH-1 Xinhui, Guangxi, China. I–J. Pa a ype, ZMB.122722-2, Baishou, Guangxi, China. K. Pa a ype,
ZMB.122763, Hezhou, Guangxi, China. L–M. Ope culum o pa a ype KIZ.240010. N–O. Ope culum
o ZMB.122722-2. Scale ba s = 1 cm, op le ba applies o all shells, sepa a e ba o ope cula L–O.
ZHANG L.-J. e al., New genus o Vi ipa idae (Mollusca) om Sou h China
41
sameda aas o holo ype;KIZ.240090•2specs(p ese edine hanol);GuilinCi y,BaishouTown,
Baishou Ri e , 0.5 m deep on mud and sand subs a e; 15 Ma . 2022; Le-Jia Zhang leg.; KIZ.240012,
KIZ.240013•2specs(p ese edine hanol);sameda aas o p eceding;ZMB.122738•1spec.;same
da aas o p eceding;KIZ.240091•1spec.(p ese edd y);HechiCi y,YizhouDis ic ,Liusanjie
Town,onbanko XiajianRi e ;16Ma .2022;Le-JiaZhangleg.;KIZ.240092•2specs(p ese edd y);
sameda aas o p eceding;17Ma .2022;KIZ.240093•2specs(p ese edine hanol);HezhouCi y,
Zhongshan Coun y, Tonggu Town, Fengzhen Village, Shangpu, 0.5 m deep on mud subs a e in a pond;
9Ma .2020;Shui-YangHuangleg.;KIZ.240014,KIZ.240015•1spec.(p ese edine hanol);same
da a as o p eceding; ZMB.122763.
O he ma e ial
CHINA•1spec.(p ese edd y);sameda aas o holo ype;YHY-BAKLIU-XH-1.
Desc ip ion
Adul shell (Fig. 24A–K) medium in size (25.69 ± 3.35 mm), hin bu s u dy, conical shell wi h low
spi e,oli e ooli eb own,wi h i e osixwho ls,apexacu e; eleoconchwho lsno in la ed,wi h h ee
smoo h ela i ely weak p ima y keels, wi h one o wo e en weake smoo h seconda y keels loca ed
be weensu u eand i s keel,some imesoneaddi ionalweakkeelloca edbe ween i s andsecond/
secondand hi dkeels,baseo shellwi hone o i eweakkeels,allkeelsusuallyda ke  han es o 
shell su ace; ape u e o a e, less han hal o shell in heigh , inne shell su ace bluish whi e, o ange-
ed o da k pu ple, ou e lip hin, always wi h a s ong basal angula pe iphe y, columella lip co e ed
wi h hick inne shell laye s, whi e o o ange ed, adul columella lip ecu ed, some imes o ming a
ela i ely wide c escen -shaped callus, his s uc u e o columella lip comple ely co e ing umbilicus.
Ope culum (Fig. 24L–O) co neous, o a e, ela i ely hick, da k o ange o da k ed; ex e io su ace
smoo h; in e io su ace wi h small nuclea egion, a ound 1/4 o inne ope cula egion, wi h shiny g ains
o eins o smoo h glossy su ace.
Radula (Fig. 25) (n = 3) cen al oo h wi h one b oad cen al den icle and h ee o ou small sha p cusps
on ei he side; la e al oo h wi h one b oad cen al den icle and wo o h ee small sha p cusps on ei he
side; inne ma ginal oo h na ow, abou hal o la e al oo h in wid h, wi h one b oad cen al den icle and
i esmallsha pcuspsonei he side;ou e ma ginal oo hwide,almos sameasla e al oo hinwid h,
wi h 26 o 27 small sha p cusps.
Fig. 25. SEM pho o o he adula o Bakyie aia liusanjieae Zhang, Yen & on Rin elen gen. e sp. no .,
ZMB.122722-2.
Eu opean Jou nal o Taxonomy 1005: 1–64 (2025)
48
E ymology
Named a e ‘Lui Kong’, he god o hunde in he local Min Nan Chinese dialec , popula in he olklo e
o Leizhou Coun y which includes he ype locali y o his species. The ecommended Chinese name is
雷神越田螺.
Ma e ial examined
Holo ype
CHINA•spec.p ese edine hanol;GuangdongP o ince,ZhanjiangCi y,LeizhouCoun y,Li en
Village, nea Nanguang Fa m Second Team, 0.1–0.5 m deep on mud and ock subs a e in ponds; 6 Ap .
2020; Yu-Xuan Zhou leg.; KIZ.240037.
Pa a ypes
CHINA•4specs(p ese edine hanol);sameda aas o holo ype;KIZ.240038 oKIZ.240041•2specs
(p ese edine hanol);sameda aas o holo ype;ZMB.122764•3specs(p ese edine hanol);same
da a as o holo ype; KIZ.240101.
O he ma e ial
CHINA•1spec.(p ese edd y);sameda aas o holo ype;YHY-BAKLK-LR-1.
Fig. 30. Shells and ope culum o Bakyie aia luikongi Yen, Zhang & on Rin elen gen. e sp. no .
A–D. Holo ype, KIZ.240037, Leizhou, Guangdong, China. E–F. Pa a ype, ZMB.122764, Leizhou,
Guangdong, China. G. YHY-BAKLK-LR-1, Leizhou, Guangdong, China. H–I. Ope culum o holo ype,
KIZ.240037. Scale ba s = 1 cm, op le ba applies o all shells, sepa a e ba o ope culum G–H.

ZHANG L.-J. e al., New genus o Vi ipa idae (Mollusca) om Sou h China
49
Desc ip ion
Adul shell (Fig. 30A–G) ela i ely small o he genus (20.66 ± 0.70 mm), hin bu s u dy, b oad conical
oconical,da koli eb own,wi h i ewho ls,apexacu e; eleoconchwho lssome imeswi hshoulde ,
wi h h ee weak bu ob ious smoo h p ima y keels, wi h one o wo weake smoo h seconda y keels
loca edbe weensu u eand i s keel,alwayswi honeweakkeelloca edbe ween i s andsecond/second
and hi d keels, base o shell wi h h ee o ou weak keels, all keels usually da ke han es o shell
su ace; ape u e o a e, lnea ly hal o shell in heigh , inne shell su ace bluish whi e, ou e lip hin,
always wi h an ob ious angula pe iphe y, columella lip co e ed wi h hin inne shell laye s, whi e o
o ange, columella lip olded ou wa ds, comple ely co e ing umbilicus.
Ope culum (Fig. 30H–I) co neous, o a e, hin, da k o ange o da k ed; ex e io su ace smoo h; in e io
su ace wi h small nuclea egion, a ound 1/5 o inne ope cula egion, wi h shiny g ains o eins o
smoo h glossy su ace.
Radula (Fig. 31) (n = 2) cen al oo h wi h one b oad cen al den icle and ou small sha p cusps on
ei he side; la e al oo h wi h one b oad cen al den icle and ou small sha p cusps on ei he side; inne
ma ginal oo h na ow, abou hal o la e al oo h in wid h, wi h one b oad cen al den icle and h ee o
ou small sha p cusps on ei he side; ou e ma ginal oo h wide, almos same wid h as la e al oo h, wi h
19 o 20 small sha p cusps.
Rema ks
Bakyie aia luikongi gen. e sp. no . is he only species o Bakyie aia gen. no . wi h a conical shell and
weak p ima y keels dis ibu ed ou side Hainan Island and is he sis e species o he clade including all
species wi h weak p ima y keels om Hainan. I can be easily dis inguished om all congene s based on
he small hin conical shell wi h h ee e y weak p ima y keels and lowe spi e. I is he smalles species
among hose wi h weak p ima y keels.
Habi a and dis ibu ion
Only known om mud and ock subs a e in he shallow wa e o ponds in Leizhou Peninsula,
Guangdong P o ince, China.
Fig. 31. SEM pho o o he adula o Bakyie aia luikongi Yen, Zhang & on Rin elen gen. e sp. no .,
pa a ype, ZMB.122764.
Eu opean Jou nal o Taxonomy 1005: 1–64 (2025)
50
Bakyie aia on inalis Zhang, Yen & on Rin elen gen. e sp. no .
u n:lsid:zoobank.o g:ac :8DFECB0B-656C-42B6-B3FE-51C2BC907619
Figs 32–33
Diagnosis
Shellmediuminsize(24.25±2.59mm),s u dy,conical,who lsno in la ed,wi hweakp ima ykeels,
shoulde ounded; columella lip ecu ed, o ming callus co e ing umbilicus comple ely; ope culum
wi h small nuclea egion (a ound 1/5 o inne ope cula egion) wi h shiny g ains o eins; ou e ma ginal
oo h o adula wi h 26–29 small sha p cusps.
E ymology
Re e s o he cold sp ing habi a o he ype locali y. The ecommended Chinese name is 冷泉越田螺.
Ma e ial examined
Holo ype
CHINA•spec.p ese edine hanol;HainanP o ince,DinganCoun y,Jiuwen angTown,Jiuwen ang
Cold Sp ing, 0.5 m deep on mud and sand subs a e; 6 Ma . 2022; Le-Jia Zhang leg.; KIZ.240042.
Fig. 32. Shells and ope culum o Bakyie aia on inalis Zhang, Yen & on Rin elen gen. e sp. no .
A–D. Holo ype, KIZ.240042, Jiuwen ang, Hainan, China. E. YHY-BAKFON-LQ-1, Jiuwen ang, Hainan,
China. F. Pa a ype, ZMB.122764, Jiuwen ang, Hainan, China. G. Pa a ype, KIZ.240104, Wanquan
Ri e , Hainan, China. H–I. Ope culum o holo ype, KIZ.240042. Scale ba s = 1 cm, op le ba applies
o all shells, sepa a e ba o ope culum H–I.
ZHANG L.-J. e al., New genus o Vi ipa idae (Mollusca) om Sou h China
51
Pa a ypes
CHINA•1spec.(p ese edine hanol);sameda aas o holo ype;KIZ.240043•2specs(p ese ed
ine hanol);sameda aas o holo ype;ZMB.122733•1spec.(p ese edine hanol);sameda aas o 
holo ype;KIZ.240102•2specs(p ese edine hanol);HainanP o ince,DinganCoun y, i e nea 
LeimingTown;2018;KIZ.240103•1spec.(p ese edine hanol);HainanP o ince,QionghaiCi y,
Shibi Town, Wanquan Ri e , 0.1 m deep on ock; 7 Ma . 2022; Le-Jia Zhang leg., KIZ.240104.
O he ma e ial
CHINA•1spec.(p ese edd y);sameda aas o holo ype;YHY-BAKFON-LQ-1.
Desc ip ion
Adul shell (Fig. 32A–G) medium in size (24.25 ± 2.59 mm), hin o hick, s u dy, conical, oli e o da k
oli eb own,wi h i ewho ls,apexacu e; eleoconchwho lsno in la ed,wi h h eeweakbu ob ious
smoo hp ima ykeels,wi hone o woweake smoo hseconda ykeelloca edbe weensu u eand i s 
keel,alwayswi honeaddi ionalweakkeelloca edbe ween i s andsecond/secondand hi dkeels,base
o shell wi h wo o ou weak keels, all keels usually da ke han he o he pa o shell su ace in colou ;
ape u e o a e, less han hal o shell in heigh , inne shell su ace bluish whi e, ou e lip hin, always
wi h a s ong angula pe iphe y, columella lip co e ed wi h hick inne shell laye s, whi e o o ange,
adul columella lip ecu ed, some imes o ming a c escen -shaped callus, his s uc u e o columella
lip comple ely co e ing umbilicus.
Ope culum (Fig. 32H–I) co neous, o a e, hin, o ange o da k ed; ex e io su ace smoo h; in e io
su ace wi h small nuclea egion, a ound 1/5 o inne ope cula egion, wi h shiny g ains o eins o
smoo h glossy su ace.
Radula (Fig.33)(n=2)cen al oo hwi honeb oadcen alden icleand ou  o i esmallsha pcusps
on ei he side; la e al oo h wi h one b oad cen al den icle and ou small sha p cusps on ei he side;
inne ma ginal oo h na ow, abou hal wid h o la e al oo h, wi h one b oad cen al den icle and ou
o i esmallsha pcuspsonei he side;ou e ma ginal oo hwide,almos sameasla e al oo hinwid h,
wi h 26 o 29 small sha p cusps.
Rema ks
Bakyie aia on inalis gen. e sp. no . is one o he species o Bakyie aia gen. no . wi h weak p ima y
keels. This species can be easily dis inguished om all congene s, especially he mo phologically simila
Fig. 33. SEM pho o o he adula o Bakyie aia on inalis Zhang, Yen & on Rin elen gen. e sp. no .,
pa a ype, ZMB.122733.
Eu opean Jou nal o Taxonomy 1005: 1–64 (2025)
52
species B. mu ica,basedoni smedium-sizedconicalshellwi hwho lsno in la ed, oundedshoulde 
and mo e cusps on he ou e ma ginal ee h (26–29 s 19–24).
Habi a and dis ibu ion
Mud and ock subs a e in he shallow wa e o sp ings and i e s o cen al-eas e n Hainan P o ince,
China.
Bakyie aia naiadica Zhang, Yen & on Rin elen gen. e sp. no .
u n:lsid:zoobank.o g:ac :1E34D07F-05C1-4819-ADC6-D02399C18B8B
Figs 34–35
Diagnosis
Shell medium in size (25.73 ± 0.62 mm), s u dy, cylind ical ou line, wi h e y weak p ima y keels,
weak keel abo e su u e, ou e lip angula ; columella lip ecu ed, o ming callus co e ing umbilicus
comple ely; ope culum wi h small nuclea egion (a ound 1/5 o inne ope cula egion) wi h shiny g ains
o eins; ou e ma ginal oo h o adula wi h 19–22 small sha p cusps.
E ymology
Named a e Naiads, he nymphs o eshwa e bodies such as sp ings in G eek my hology. The
ecommended Chinese name is 龙婆越田螺.
Fig. 34. Shells and ope culum o Bakyie aia naiadica Zhang, Yen & on Rin elen gen. e sp. no .
A–D. Holo ype, KIZ.240044, Longpo Temple, Hainan, China. E. Pa a ype, KIZ.240045, Xue Village,
Hainan, China. F–G. YHY-BAKNAI-XC-1, Xue Village, Hainan, China. H–I. Ope culum o holo ype,
KIZ.240044. Scale ba s = 1 cm, op le ba applies o all shells, sepa a e ba o ope culum F–G.
ZHANG L.-J. e al., New genus o Vi ipa idae (Mollusca) om Sou h China
53
Ma e ial examined
Holo ype
CHINA•spec.p ese edine hanol;HainanP o ince,HaikouCi y,sp ingnea LongpoTemple,0.1m
deep on ock; 5 Ma . 2022; Le-Jia Zhang leg.; KIZ.240044.
Pa a ypes
CHINA•1spec.(p ese edine hanol);sameda aas o holo ype;ZMB.122743•1spec.(p ese edin
e hanol); Hainan P o ince, we land in Xue Village, Haikou Ci y, 0.1 m deep on ock subs a e; 3 May
2022; Le-Jia Zhang leg.; KIZ.240045.
O he ma e ial
CHINA•1spec.(p ese edd y);HainanP o ince,we landinXueVillage,HaikouCi y,0.1mdeepon
ock subs a e; 3 May 2022; Le-Jia Zhang leg.; YHY-BAKNAI-XC-1.
Desc ip ion
Adul shell (Fig. 34A–G) medium (25.73 ± 0.62 mm), hin bu s u dy, conical, oli e o da k oli e b own,
wi h i ewho ls; eleoconchwho lsin la ed,wi hshoulde ,wi h h ee e yweaksmoo hp ima ykeels,
wi hone o woe enweake smoo hseconda ykeelsloca edbe weensu u eand i s keel,some imes
wi honeaddi ionalweakkeelloca edbe ween i s andsecond/secondand hi dkeels,baseo shellwi h
wo o ou weak keels, all keels usually da ke han es o shell su ace; ape u e o a e, less han hal o
shell in heigh , inne shell su ace bluish whi e, ou e lip hin, always wi h an ob ious angula pe iphe y,
columella lip co e ed wi h hick inne shell laye s, whi e o pale o ange, adul columella lip ecu ed,
some imes o ming a c escen -shaped callus, his s uc u e comple ely co e ing umbilicus.
Ope culum (Fig. 34H–I) co neous, o a e, hin, da k ed; ex e io su ace smoo h; in e io su ace wi h
small nuclea egion, a ound 1/5 o inne ope cula egion, wi h shiny g ains o eins o smoo h glossy
su ace.
Radula (Fig.35)(n=1)cen al oo hwi honeb oadcen alden icleand ou  o i esmallsha pcusps
onei he side;la e al oo hwi honeb oadcen alden icleand ou  o i esmallsha pcuspsonei he 
side; inne ma ginal oo h na ow, abou hal wid h o la e al oo h, wi h one b oad cen al den icle and
i e osixsmallsha pcuspsonei he side;ou e ma ginal oo hwide,almos sameasla e al oo hin
wid h, wi h 19 o 22 small sha p cusps.
Fig. 35. SEM pho o o he adula o Bakyie aia naiadica Zhang, Yen & on Rin elen gen. e sp. no .,
pa a ype, ZMB.122743.

Eu opean Jou nal o Taxonomy 1005: 1–64 (2025)
54
Rema ks
Bakyie aia naiadica gen. e sp. no . can be easily dis inguished om all congene s, especially he
mo phologically simila B. mu ica, based on i s mo e cylind ical shell wi h weake p ima y keels and
less p ominen keel a su u e.
Habi a and dis ibu ion
Rock subs a e in he shallow wa e o sp ings and we lands o no he n Hainan P o ince, China.
Bakyie aia luuemxlong Zhang, Yen & on Rin elen gen. e sp. no .
u n:lsid:zoobank.o g:ac :3C33CDFF-1DD8-4AC2-A4D6-A28D0535B1A4
Figs 36–37
Diagnosis
Shellsmall omediuminsize(23.56±2.58mm),s u dy,who lsin la ed,wi hweakp ima ykeels,
mode a e keel abo e su u e; columella lip ecu ed, o ming callus co e ing umbilicus comple ely;
ope culum wi h small nuclea egion (a ound 1/5 o inne ope cula egion) wi h shiny g ains o eins;
ou e ma ginal oo h o adula wi h 20 small sha p cusps; dis ibu ed in sou he n Hainan Island.
E ymology
Acco ding o he my hology o he Hlai people o Hainan Island, ‘luuemx long’, which means ‘ o ge o
g ow up’ in he Hlai language, is a ace o dwa es li ing in he o es . Fo med as a noun in apposi ion.
The ecommended Chinese name is 阑咙越田螺.
Ma e ial examined
Holo ype
CHINA•spec.p ese edine hanol;HainanP o ince,QionghaiCi y,HuishanTown,c eeksnea 
a mland o Mu ang Village, 0.1 m deep on mud subs a e; 7 Ma . 2022; Le-Jia Zhang leg.; KIZ.240046.
Pa a ypes
CHINA•2specs(p ese edine hanol);sameda aas o holo ype;KIZ.240047–240048•1spec.
(p ese ed in e hanol); same da a as o holo ype; ZMB.122739.
O he ma e ial
CHINA•1spec.(p ese edd y);sameda aas o holo ype;YHY-BAKLUU-MT-1.
Desc ip ion
Adul shell (Fig. 36A–E, H–J) small o medium (23.56 ± 2.58 mm), hin bu s u dy, conical, da k oli e
b own,wi h i ewho ls; eleoconchwho lsin la ed,wi h h eeweakbu ob iousp ima ykeels,wi h
one o woe enweake smoo hseconda ykeelsloca edbe weensu u eand i s keel,alwayswi hone
addi ionalweakkeelloca edbe ween i s andsecond/secondand hi dkeels,baseo shellwi h wo o
ou weak keels, all keels usually da ke han es o shell su ace; ape u e o a e, less han hal o shell
in heigh , inne shell su ace bluish whi e, ou e lip hin, always wi h an ob ious angula pe iphe y,
columella lip co e ed wi h hick inne shell laye s, whi e, columella lip olded ou wa ds, co e ing
umbilicus.
Ope culum (Fig. 36F–G) co neous, o a e, hin, da k ed; ex e io su ace smoo h; in e io su ace wi h
small nuclea egion, a ound 1/5 o inne ope cula egion, wi h shiny g ains o eins o smoo h glossy
su ace.
ZHANG L.-J. e al., New genus o Vi ipa idae (Mollusca) om Sou h China
55
Fig. 36. Shells and ope culum o Bakyie aia luuemxlong Zhang, Yen & on Rin elen gen. e sp. no .
A–D. Holo ype, KIZ.240046, Mu ang, Hainan, China. E. Pa a ype, ZMB.122739, Mu ang, Hainan,
China. F–G. Ope culum o holo ype, KIZ.240046. H–J. YHY-BAKLUU-MT-1, Mu ang, Hainan, China.
Scale ba s = 1 cm, op le ba applies o all shells, sepa a e ba o ope culum F–G.
Eu opean Jou nal o Taxonomy 1005: 1–64 (2025)
56
Radula (Fig. 37) (n = 1) cen al oo h wi h one b oad cen al den icle and ou small sha p cusps on
ei he side; la e al oo h wi h one b oad cen al den icle and h ee small sha p cusps on ei he side; inne
ma ginal oo h na ow, abou hal o la e al oo h in wid h, wi h one b oad cen al den icle and unknown
numbe o small sha p cusps on ei he side (all hea ily wo n); ou e ma ginal oo h wide, almos same
as la e al oo h in wid h, wi h a ound 20 small sha p cusps.
Rema ks
Bakyie aia luuemxlong gen. e sp. no . is one o he species o Bakyie aia gen. no . wi h weak p ima y
keels. This species can be easily dis inguished om all congene s based on he small- o medium-sized
conicalshellwi hin la edwho lsandmode a ekeelabo esu u e.
Habi a and dis ibu ion
Only known om he ype locali y on mud subs a e in he shallow wa e o c eeks in sou he n Hainan
P o ince, China.
Bakyie aia ind apu a Zhang, Yen & on Rin elen gen. e sp. no .
u n:lsid:zoobank.o g:ac :D7B758FD-46BD-4DED-8C15-AD9BED3528AA
Figs 38–39
Diagnosis
Shellmediuminsize(26.92±2.45mm), hin,s u dy,who lsin la ed,wi hp ominen p ima ykeelsa 
su u e; columella lip ecu ed, o ming callus co e ing umbilicus comple ely; ope culum wi h small
nuclea egion (a ound 1/5 o inne ope cula egion) wi h shiny g ains o eins; ou e ma ginal oo h o
adula wi h 21–26 small sha p cusps; dis ibu ed in cen al Vie nam.
E ymology
Named a e Ind apu a, capi al o he ancien kingdom o Champa, which is close o he ype locali y.
Fo med as a noun in apposi ion. The ecommended Chinese name is 占婆越田螺.
Ma e ial examined
Holo ype
VIETNAM•1spec.(p ese edine hanol);PhuocSon,KhamDuc,CaiRi e d ainage;21–31Oc .
2006; F ank Köhle leg.; ZMB.114403.1.
Fig. 37. SEM pho o o he adula o Bakyie aia luuemxlong Zhang, Yen & on Rin elen gen. e sp. no .,
pa a ype, ZMB.122739.
ZHANG L.-J. e al., New genus o Vi ipa idae (Mollusca) om Sou h China
57
Pa a ypes
VIETNAM•2specs(p ese edine hanol);sameda aas o holo ype;ZMB.114403•5specs(p ese ed
ine hanol);NghêAn,TânDuongDis ic ,TamQuang,T uongSonM s,moun ainc eek;21–31Oc .
2006;F ankKöhle leg.;ZMB.114402.1,ZMB.114402•3specs(p ese edine hanol); same da a as
o p eceding; ZMB.114406.
Desc ip ion
Adul shell (Fig. 38A–F) medium in size (26.92 ± 2.45 mm), hin bu s u dy, conical, oli e o g eenish
b own,wi h i e osixwho ls,apexacu e; eleoconchwho ls,especiallybodywho l,sligh lyin la ed,
some imes o ming shoulde , wi h h ee smoo h p ima y keels, wi h one o wo weake smoo h seconda y
keelsloca edbe weensu u eand i s keel,some imesoneaddi ionalweakkeelloca edbe ween i s and
second/secondand hi dkeels,baseo shellwi h h ee o i eweakkeels,keelsusuallyda ke  han es 
o shell su ace; ape u e o a e, less han hal o shell in heigh , inne shell su ace bluish whi e, ou e
lip hin, always wi h a s ong angula pe iphe y, columella lip co e ed wi h hick inne shell laye s,
whi e o o ange, adul columella lip ecu ed, o ming a e y na ow c escen -shaped callus o wi hou
his s uc u e, columella lip o his s uc u e usually co e ing umbilicus, umbilicus some imes na owly
open, bo de ed by a keel.
Ope culum (Fig. 38G–H) co neous, o a e, hin, yellow o o ange; ex e io su ace smoo h; in e io
su ace wi h small nuclea egion, a ound 1/5 o inne ope cula egion, wi h shiny g ains o eins.
Fig. 38. Shells and ope culum o Bakyie aia ind apu a Zhang, Yen & on Rin elen gen. e sp. no .
A–D. Pa a ype, ZMB.114402.1, Nghe An, Vie nam. E–F. Holo ype, ZMB.114403, Phuoc Son, Vie nam.
K–L. Ope culum o pa a ype ZMB.114402. Scale ba s = 1 cm, op le ba applies o all shells, sepa a e
ba o ope culum G–H.
Eu opean Jou nal o Taxonomy 1005: 1–64 (2025)
64
Zhang L.J. & on Rin elen T. 2021. The neglec ed ope culum: A e ision o he ope cula cha ac e s in
i e snails (Caenogas opoda: Vi ipa idae). Jou nal o Molluscan S udies 87: eyab008.
h ps://doi.o g/10.1093/mollus/eyab008
Zhang L.J., Chen S.C., Yang L.T., Jin L. & Köhle F. 2015. Sys ema ic e ision o he eshwa e snail
Ma ga ya Ne ill, 1877 (Mollusca: Vi ipa idae) endemic o he ancien lakes o Yunnan, China, wi h
desc ip ion o new axa. Zoological Jou nal o he Linnean Socie y 174: 760–800.
h ps://doi.o g/10.1111/zoj.12260
Zhang L.J., Du L.N. & on Rin elen T. 2023. A new genus o i e snails, Dalipaludina (Gas opoda,
Vi ipa idae), endemic o he Yunnan Pla eau o SW China. Zoosys ema ics and E olu ion 99: 285–297.
h ps://doi.o g/10.3897/zse.99.102586
Zhu X.G. 1994. Ju assic-C e aceous nonma ine gas opods om no he n Xinjiang. Ac a Palaeon ologica
Sinica 33: 85–105.
Zilch A. 1938. Das Vo kommen on Buliminopsis ince a (P .) au Fo mosa und in China. A chi ü
Molluskenkunde 7: 3.
Zilch A. 1955. Die Typen und Typoide des Na u -Museums Senckenbe g, 14: Mollusca, Vi ipa idae.
A chi ü Molluskenkunde 84: 45–87.
Zilch A. 1974. Zu Geschich e de deu schen Malakozoologie, XIV. Vinzenz G edle und die E o schung
de Weich ie e Chinas du ch F anziskane aus Ti ol. A chi ü Molluskenkunde 104: 171–228.
P in ed e sions o all pape s a e deposi ed in he lib a ies o ou o he ins i u es ha a e membe s o he
EJTconso ium:Muséumna ionaldʼHis oi ena u elle,Pa is,F ance;MeiseBo anicGa den,Belgium;
Royal Museum o Cen al A ica, Te u en, Belgium; Royal Belgian Ins i u e o Na u al Sciences,
B ussels, Belgium. The o he membe s o he conso ium a e: Na u al His o y Museum o Denma k,
Copenhagen, Denma k; Na u alis Biodi e si y Cen e , Leiden, he Ne he lands; Museo Nacional de
Ciencias Na u ales-CSIC, Mad id, Spain; Leibniz Ins i u e o he Analysis o Biodi e si y Change,
Bonn – Hambu g, Ge many; Na ional Museum o he Czech Republic, P ague, Czech Republic; The
S einha d Museum o Na u al His o y, Tel A i , Is aël.
Supplemen a y iles
Supp. ile 1. The alignmen o he COI sequences in he p esen s udy.
h ps://doi.o g/10.5852/ej .2025.1005.2985.13431
Supp. ile 2. ML ee based on he COI sequences in he p esen s udy
h ps://doi.o g/10.5852/ej .2025.1005.2985.13433