A new genus and species o eshwa e mussel (Bi al ia, Unionidae)
om Anhui, China
Hui Chen1, Wen-Hao Wang1, Zhong-Guang Chen1, Hong-Quan Xiang2, Yue-Ming He3,
Shan Ouyang1, Ke-Lin Chu4, Xiao-Ping Wu1
1 School o Li e Sciences, Nanchang Uni e si y, Nanchang, 330031, China
2 Depa men o Animal Science, Yuxi Ag icul u e Voca ion-Technical College, Yuxi, 653106, China
3 College o Enginee ing and Technology, Tianjin Ag icul u al Uni e si y, Tianjin, 300384, China
4 Nanjing Ins i u e o En i onmen al Sciences, Minis y o Ecology and En i onmen , Nanjing 210042, China
h ps://zoobank.o g/A8F7B4D9-C570-4BDD-B2C3-00A85E67A390
Co esponding au ho s: Xiao-Ping Wu ([email p o ec ed]); Ke-Lin Chu ([email p o ec ed])
Academic edi o : Le-Jia Zhang ♦
Recei ed
4 Sep embe 2025 ♦
Accep ed
7 No embe 2025 ♦
Published
18 No embe 2025
Abs ac
The middle and lowe eaches o he Yang ze Ri e hos a highly di e se Unionidae auna. He ein, a new genus and species o
eshwa e mussel, Huizhou xie ianxiangi H. Chen, Z.-G. Chen, H.-Q. Xiang, Y.-M. He & X.-P. Wu, gen. e sp. no ., is desc ibed
om Huangshan Ci y, Anhui P o ince, based on compa a i e mo phological and molecula phylogene ic e idence. The new axon
is dis inguished by a unique se o shell cha ac e is ics, including an o al shape, a zigzag sculp u e co e ing he en i e su ace, and
equen ado nmen wi h elegan da k g een ays. These dis inc mo phological ea u es, coupled wi h signi ican gene ic di e en i-
a ion, obus ly suppo i s es ablishmen as a new genus and species.
Key Wo ds
Globunionini, Mollusca, phylogeny, axonomy, Yang ze Ri e Basin
In oduc ion
The Yang ze Ri e Basin in China ha bo s a ich and en-
demic auna o p ima y eshwa e mussels, wi h mo e
han 70 ecognized species desc ibed o da e (He and
Zhuang 2013; G a and Cummings 2021; Guo 2022; Dai
e al. 2025). Recen ex ensi e su eys and axonomic
s udies ha e led o he con inual disco e y o new axa
(Chen e al. 2023; Dai e al. 2023, 2024a, b; Chen e al.
2025). Despi e he nume ous endemic species al eady
documen ed, hei ull di e si y emains incomple ely
unde s ood, pa icula ly among small-bodied o ms, wi h
e en new ibes being iden i ied (Dai e al. 2025). These
indings indica e ha a signi ican numbe o eshwa e
mussel species in he egion ha e ye o be desc ibed.
In his s udy, esea che s collec ed eshwa e mus-
sel specimens wi h dis inc i e shell mo phology om
Huangshan Ci y, Anhui P o ince, China, which did no
ma ch any known species. Based on in eg a ed mo pho-
logical and molecula phylogene ic analyses, we desc ibe
i he e as a new genus and new species wi hin he sub-
amily Unioninae Ra inesque, 1820: Huizhou xie ianx-
iangi sp. no . This disco e y u he en iches he known
eshwa e mussel di e si y o he Yang ze Ri e Basin,
pa icula ly among he iny auna.
Ma e ials and me hods
Specimen sampling and mo phological
obse a ions
F eshwa e mussels we e collec ed in he Xinanjiang
Ri e (29.5678°N, 117.9225°E), Bing an Village, Xikou
Zoosys . E ol. 101 (4) 2025, 2243–2250|DOI 10.3897/zse.101.171070
Copy igh Chen, H. e al. This is an open access a icle dis ibu ed unde he e ms o he C ea i e Commons A ibu ion License (CC BY 4.0), which pe mi s un es ic ed use,
dis ibu ion, and ep oduc ion in any medium, p o ided he o iginal au ho and sou ce a e c edi ed.
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Chen, H. e al.: New genus and species o eshwa e mussels
2244
Town, Xiuning Coun y, Huangshan Ci y, Anhui P o ince,
China, in Augus 2025 (Fig. 1). The specimens we e p e-
se ed in 95% e hanol and deposi ed a he Museum o
Biology, Nanchang Uni e si y (NCUMB), China. A Ve -
nie calipe was used o measu e shell leng h (L), shell
wid h (W), and shell heigh (H).
DNA ex ac ion, PCR, sequencing, and
phylogene ic analyses
To al genomic DNA was ex ac ed om oo issues using
he TIANamp Ma ine Animals DNA Ki (Tiangen Bio-
ech) ollowing he manu ac u e ’s p o ocol. F- ype mi-
ochond ial COI sequences we e ampli ied using p ime s
LCO22me2 (GGTCAACAAAYCATAARGATATTGG)
and HCO700dy2 (TCAGGGTGACCAAAAAAYCA)
(Walke e al. 2007). Polyme ase chain eac ion (PCR)
ampli ica ions o COI we e pe o med in a inal 25 μL e-
ac ion mix u e con aining 1 μL o empla e DNA, 1 μL o
each p ime , 12.5 μL o G een Taq Mix (Vazyme, China),
and 9.5 μL o ddH2O. The mal cycling began wi h one
cycle a 98 °C o 10 s, ollowed by 35 cycles o dena-
u a ion a 94 °C o 1 min, annealing a 50 °C o 1 min,
and ex ension a 72 °C o 1 min, wi h a inal ex ension
a 72 °C o 7 min. The PCR p oduc s we e pu i ied and
sequenced using an ABI 3730XL analyze . The newly ob-
ained sequences we e deposi ed in GenBank (Table 1).
Sequences we e aligned using MAFFT .7 based on he
Q-INS-i me hod (Ka oh e al. 2019). Pai wise dis ances
be ween species we e calcula ed using MEGA X (Kuma
e al. 2018). The bes subs i u ion model was selec ed us-
Figu e 1. Collec ion si es o Huizhou xie ianxiangi gen. e sp. no . The gold s a indica es he ype locali y.
Zoosys . E ol. 101 (4) 2025, 2243–2250
zse.penso .ne
2245
ing he co ec ed Bayesian In o ma ion C i e ion (BIC) in
MODELFINDER .2.2.0 (Kalyaanamoo hy e al. 2017).
Bayesian in e ence (BI) and maximum likelihood (ML)
analyses we e pe o med using M Bayes ( e . 3.2.6;
Ronquis e al. 2012) and IQ-TREE ( e . 2.2; Minh e al.
2020), espec i ely, wi h e e ence o he selec ed model
o sequence e olu ion. Fo Bayesian analysis, wo uns
we e pe o med simul aneously wi h ou Ma ko chains
s a ing om a andom ee. Bayesian pos e io p obabil-
i ies (BPPs) o nodes we e de e mined using Me opo-
lis-coupled Ma ko chains (one cold chain) o 2,000,000
gene a ions, wi h sampling e e y 1000 gene a ions. The
i s 25% o sampled ees we e disca ded as bu n-in
when he s anda d de ia ion o spli equencies be ween
he wo uns was less han 0.01; he emaining ees we e
hen used o c ea e a 50% majo i y- ule consensus ee
and o es ima e BPPs. Node suppo o maximum likeli-
hood analysis was de e mined using 1000 apid boo s ap
(BS) eplica es.
Resul s
The GTR+F+I+R4 model was selec ed as he bes - i
model o nucleo ide subs i u ion by he AIC c i e ion.
Molecula phylogenies based on BI and ML analyses
Table 1. GenBank accession numbe s o he COI sequences used in his s udy.
Species Locali y Accession numbe
Huizhou xie ianxiangi gen. e sp. no . Xiuning, Huangshan, Anhui, China In submission
Huizhou xie ianxiangi gen. e sp. no . Xiuning, Huangshan, Anhui, China In submission
Globunio mi i icus Chen, Dai, Huang & Wu, 2025 Nanchang, Jiangxi, China PV368604
Globunio mi i icus Chen, Dai, Huang & Wu, 2025 Nanchang, Jiangxi, China PV368605
Diau o a au o ea (Heude, 1883) Ji’an, Jiangxi, China OQ829360
Acu icos a chinensis (Lea, 1868) Jiangxi, China MG462921
Schis odesmus lamp eyanus (Bai d & Adams, 1867) Jiangxi, China MG463037
Schis odesmus lamp eyanus (Bai d & Adams, 1867) Hunan, China MG463043
Schis odesmus spinosus (Simpson, 1900) Jiangxi, China MG463046
Unio pic o um (Linnaeus, 1758) Eu ope KC429109
Unio c assus Philipsson, 1788 Poland KY290446
Tchangsinaia piscicula (Heude, 1874) Jiangxi, China MG462977
Cuneopsis cel i o mis (Heude, 1874) Jiangxi, China MG462964
Cuneopsis heudei (Heude, 1874) Jiangxi, China MG462970
Aculamp o ula ib osa (Heude, 1877) Jiangxi, China MG462909
Nodula ia douglasiae (G ay, 1833) China KX822653
Nodula ia b e iconcha Lee, Kim, Bogan & Kondo, 2020 Sou h Ko ea MT020662
In e siunio yanagawensis (Kondo, 1982) Japan MT020654
In e siunio einianus (Kobel , 1879) Japan MT020657
Pseudocuneopsis sichuanensis Huang, Dai, Chen & Wu, 2022 Sichuan, China MZ540966
Pseudocuneopsis capi a a (Heude, 1874) Anhui, China NC042469
Alasmidon a ma gina a Say, 1818 U.S. AF156502
Lasmigona comp essa (Lea, 1829) U.S. AF156503
Anodon a ana ina (Linnaeus, 1758) Russia KX822632
Pseudanodon a complana a (Rossmässle , 1835) Uk aine KX822661
Lanceola ia gladiola (Heude, 1877) Jiangxi, China MG463007
Lanceola ia oxy hyncha (Ma ens, 1861) Japan MT020648
C is a ia plica a (Leach, 1814) Jiangxi, China MG462956
Lepidodesma languila i (Heude, 1874) Jiangxi, China MG463015
Sinanodon a woodiana (Lea, 1834) China KX822668
Be ingiana be ingiana (Middendo , 1851) Japan MT020557
Ple holophus enuis (G ay, 1833) Vie nam KX822658
Anemina a cae o mis (Heude, 1877) Jiangxi, China MG462936
Amu anodon a kijaensis Mosk iche a, 1973 Russia MK574204
Coela u a aegyp iaca (Cailliaud, 1823) Egyp KJ081162
Indonaia ande soniana (Ne ill, 1877) Myanma MF352275
Pa eysia nagpoo ensis (Lea, 1860) India JQ861229
P onodula ia japanensis (Lea, 1859 Japan LC505454
Lamp o ula leaii (G ay, 1833) Jiangxi, China MG462996
Lampsilis siliquoidea (Ba nes, 1823) U.S. MH560773
Quad ula quad ula (Ra inesque, 1820) U.S. HM230409
Ma ga i i e a dahu ica (Middendo , 1850) Russia KJ161516
Gibbosula ochechoua ii (Heude, 1875) Jiangxi, China MG463022
zse.penso .ne
Chen, H. e al.: New genus and species o eshwa e mussels
2246
Figu e 2. Maximum likelihood ee in e ed om COI gene sequences.
0.05
Schis odesmus lamp eyanus
Cuneopsis heudei
Aculamp o ula ib osa
Lepidodesma languila i
Pseudocuneopsis sichuanensis
Amu anodon a kijaensis
Pseudanodon a complana a
Acu icos a chinensis
Lasmigona comp essa
Cuneopsis cel i o mis
In e siunio yanagawensis
Sinanodon a woodiana
Huizhou xie ianxiangi gen. & sp. no .
Globunio mi i icus
Nodula ia b e iconcha
Lamp o ula leaii
Ple holophus enuis
Tchangsinaia piscicula
Anemina a cae o mis
Huizhou xie ianxiangi gen. & sp. no .
Quad ula quad ula
Ma ga i i e a dahu ica
Pseudocuneopsis capi a a
Nodula ia douglasiae
P onodula ia japanensis
Indonaia ande soniana
Unio pic o um
Lanceola ia gladiola
In e siunio einianus
Schis odesmus lamp eyanus
Pa eysia co uga a
Be ingiana be ingiana
Alasmidon a ma gina a
Globunio mi i icus
Schis odesmus spinosus
Lampsilis siliquoidea
C is a ia plica a
Anodon a ana ina
Lanceola ia oxy hyncha
Gibbosula ochechoua ii
Diau o a au o ea
Unio c assus
Coela u a aegyp iaca
97
99
100
100
60
100
100
50
100
44
100
99
66
55
38
100
99
57
67
100
100
100
93
90
95
61
99
92
74
99
78
99
49
85
86
79
77
42
92
46
100
Unioninae
Pa eysiinae
Ambleminae
Gonideinae
ou g oups
Unionidae
Ma ga i i e idae
showed ha he sub amily Unioninae o med a monophy-
le ic g oup (BS = 85%, BPP = 0.94). The species collec -
ed om Huangshan, Anhui, China, oge he wi h Globu-
nio mi i icus Chen, Dai, Huang & Wu, 2025, o med a
s ongly suppo ed monophyle ic clade (BS = 86%, BPP
= 0.92) (Figs 2, 3). Pai wise unco ec ed COI p-dis ance
analysis demons a ed a gene ic dis ance o 11% be ween
his species and Globunio mi i icus. These dis ances, as
well as he dis inc mo phological cha ac e is ics, p o-
ide compelling e idence o i s classi ica ion as a new
genus and species, Huizhou xie ianxiangi sp. no ., in he
ibe Globunionini.
Zoosys . E ol. 101 (4) 2025, 2243–2250
zse.penso .ne
2247
Figu e 3. Bayesian in e ence ee in e ed om COI gene sequences.
0.2
Schis odesmus lamp eyanus
Nodula ia b e iconcha
Indonaia ande soniana
Schis odesmus lamp eyanus
Ple holophus enuis
Acu icos a chinensis
Globunio mi i icus
Schis odesmus spinosus
Gibbosula ochechoua ii
Cuneopsis cel i o mis
Anodon a ana ina
Be ingiana be ingiana
Ma ga i i e a dahu ica
Sinanodon a woodiana
Pa eysia co uga a
Nodula ia douglasiae
Pseudocuneopsis sichuanensis
Coela u a aegyp iaca
In e siunio einianus
Huizhou xie ianxiangi gen. & sp. no .
Unio c assus
Aculamp o ula ib osa
C is a ia plica a
Diau o a au o ea
Tchangsinaia piscicula
Globunio mi i icus
Lepidodesma languila i
In e siunio yanagawensis
Lasmigona comp essa
Cuneopsis heudei
Alasmidon a ma gina a
Lamp o ula leaii
Pseudanodon a complana a
Quad ula quad ula
Lampsilis siliquoidea
P onodula ia japanensis
Unio pic o um
Amu anodon a kijaensis
Lanceola ia oxy hyncha
Anemina a cae o mis
Lanceola ia gladiola
Huizhou xie ianxiangi gen. & sp. no .
Pseudocuneopsis capi a a
1
1
0.41
1
0.88
0.32
0.68
0.39
0.92
0.63
1
1
0.91
0.51
0.64
1
1
1
0.29
0.99
0.14
0.43
1
0.26
1
0.75
0.98
0.66
0.93
0.6
1
0.64
0.9
0.98
0.94
0.77
0.56
0.94
1
0.35
1
Ma ga i i e idae
Ambleminae
Gonideinae
Pa eysiinae
Unioninae
Unionidae
ou g oups
Sys ema ics
Family Unionidae Ra inesque, 1820
Sub amily Unioninae Ra inesque, 1820
T ibe Globunionini Y.-T. Dai, Z.-G. Chen, X.-
C. Huang & X.-P. Wu, 2025
Genus Huizhou H. Chen, Z.-G. Chen, H.-Q. Xiang,
Y.-M. He & X.-P. Wu, gen. no .
h ps://zoobank.o g/0486F794-ADEB-46B5-8A00-94DBD10A0E5A
Type species. Huizhou xie ianxiangi H. Chen, Z.-G.
Chen, H.-Q. Xiang, Y.-M. He & X.-P. Wu, gen. e sp. no .
(Fig. 4).
Diagnosis. Shell small-sized, o al, hick, solid; wi h
a zigzag sculp u e co e ing he en i e su ace, o en
ado ned wi h elegan da k g een ays. Umbo in la ed,
o en e oded. Incu en ape u e and excu en ape u e
ob ious papillae a anged in one ow, excu en ape -
u e papillae a e mo e han wice he size o he incu -
en ape u e.
E ymology. The name o his genus o igina es om
he ancien Chinese place name Huizhou, which was an
adminis a i e egion in Chinese his o y. The ancien
name o he place whe e his species was disco e ed.
Ve nacula name. 徽州蚌 (hui zhou bang). English
common name: Huizhou Mussel.
Rema ks. The es ablishmen o he new genus Huizhou
is jus i ied by cong uen e idence om mo phology and
molecula phylogeny. I di e s mo phologically om he
ype genus Globunio o he ibe Globunionini by lacking
he diagnos ic elonga ed-globula shell. Phylogene ically,
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Chen, H. e al.: New genus and species o eshwa e mussels
2248
Figu e 4. Huizhou xie ianxiangi gen. e sp. no . A–E. Holo ype, NCUAH250801; F–H. Pa a ype, NCUAH250802; I–L. Pa a ype,
NCUAH250803; M, N. Hinge s uc u e o le and igh al es; O. Ana omical ea u es, p ia, papillae in incu en ape u e; p exa,
papillae in excu en ape u e. Scale ba s: 10 mm (A–L), 1 mm (M, N), 1 mm (O).
he wo gene a o m dis inc , s ongly suppo ed lineag-
es wi h a conside able gene ic dis ance (11% o COI),
which alls wi hin he ange o in e gene ic di e gence
obse ed in o he unionid ibes. This combina ion o
ma ked mo phological discon inui y and signi ican ge-
ne ic di e en ia ion p o ides compelling e idence o
hei sepa a ion a he gene ic le el.
Huizhou xie ianxiangi H. Chen, Z.-G. Chen, H.-Q.
Xiang, Y.-M. He & X.-P. Wu, sp. no .
h ps://zoobank.o g/2DF41693-B63D-44BD-BE42-6AF3A2625A02
Ma e ial examined. Holo ype: NCUAH250801, shell
leng h 33.7 mm, shell wid h 14.7 mm, shell heigh
25.7 mm (Fig. 4A), Xinanjiang Ri e , Bing an Village,
Xikou Town, Xiuning Coun y, Huangshan Ci y, An-
hui P o ince, China, 29.5678°N, 117.9225°E, in Au-
gus 2025.
Pa a ypes: 2 specimens, NCUAH250802, 03; shell
leng h 35.2, 33.5 mm; shell wid h 27.5, 25.1 mm; shell
heigh 15.1, 13.8 mm; locali y and habi a same as holo ype.
Diagnosis. Same as he genus.
Desc ip ion. Shell small-sized, o al, hick, solid; wi h
a zigzag sculp u e co e ing he en i e su ace bu s eng h-
ening om umbo o edge, o en ado ned wi h elegan
da k g een ays. Umbo in la ed, o en e oded. Ligamen
sho and s ong. Man le muscle sca s ob ious. An e i-
o adduc o muscle sca s o al, deep; pos e io adduc o
muscle sca s o al, shallow and smoo h (Fig. 4A–L). Le
al e wi h wo pseudoca dinal ee h, equal heigh ; an e i-
o pseudoca dinal oo h o he igh al e well de eloped,
pos e io pseudoca dinal oo h educed. La e al ee h o
bo h al es hick (Fig. 4M, N). Nac e whi e bluish-whi e.
Incu en ape u e and excu en ape u es ob ious papil-
lae a anged in one ow, excu en ape u e papillae a e
mo e han wice he size o he incu en ape u e, bo h
ape u e wi h b own pigmen a ion (Fig. 4O).
Zoosys . E ol. 101 (4) 2025, 2243–2250
zse.penso .ne
2249
Figu e 5. Habi a s and li e animals o Huizhou xie ianxiangi gen. e sp. no .
E ymology. The name o his species is in hono o
M . Xie Tianxiang, a conchologis who collec ed he
specimens.
Ve nacula name. 谢氏徽州蚌 (Xie shi hui zhou
bang). English common name: Xie’s Huizhou Mussel.
Habi a and dis ibu ion. This species is cu en ly
only known o be dis ibu ed in he Xin’an Ri e in Xiun-
ing Coun y, Huangshan Ci y, Anhui P o ince (Figs 1, 5).
Discussion
The ibe Globunionini was ecen ly es ablished and p e-
iously included only Globunio mi i icus (see Dai e al.
2025). In his s udy, h ough a combina ion o mo pholog-
ical and molecula phylogene ic analyses, we desc ibe a
new genus and species, Huizhou xie ianxiangi gen. e sp.
no ., om he Xinanjiang Ri e in Anhui P o ince. The
disco e y o his second species p o ides c i ical inde-
penden suppo o he alidi y o he ibe Globunionini
and con i ms ha i is no a mono ypic lineage. This ind-
ing signi ican ly en iches species di e si y and enhances
ou unde s anding o he ibe’s e olu iona y his o y.
Simila o G. mi i icus, H. xie ianxiangi gen. e sp. no .
possesses a small and hick shell and occu s wi hin he
Yang ze Ri e Basin (Dai e al. 2025). Howe e , ou ind-
ings e eal ha H. xie ianxiangi exhibi s a conside ably
na owe dis ibu ion and is cu en ly endemic o Anhui
P o ince. This es ic ed ange is pa icula ly conce ning
gi en ha nume ous eshwa e mussel species ha e been
epo ed as endange ed o ex inc a he ime o hei dis-
co e y (Xiang e al. 2024, 2025). Al hough Anhui P o ince
is ecognized o i s ich unionid di e si y, he p esen s udy
con i ms ha undesc ibed species, pa icula ly small-bod-
ied o c yp ic axa, likely emain o e looked. The e o e,
we u ge mo e de ailed and sys ema ic su eys in his e-
gion o iden i y such hidden di e si y, ensu ing ha hese
species can be s udied and a o ded conse a ion p io i y
be o e hei habi a s expe ience i e e sible deg ada ion.
Acknowledgmen s
We hank Tianxiang Xie and Boyu Wang o hei alu-
able con ibu ions o specimen collec ion in his s udy.
We a e g a e ul o he edi o , Lejia Zhang, and he e iew-
e s, I an N. Bolo o and A hu Bogan, o hei insigh -
ul sugges ions and cons uc i e commen s, which ha e
g ea ly imp o ed he quali y o his manusc ip .
Re e ences
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