Published November 17, 2025
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Pseudocuneopsis heqing sp. nov. (Bivalvia, Unionidae) from karst rivers of Guizhou, China: integrative species delimitation and mitogenomic phylogeny
Authors/Creators
- 1. Shanxi Normal University, Taiyuan, China
- 2. Beijing NO.15 High School, Beijing, China
Description
The endemic freshwater mussel genus Pseudocuneopsis (Unionidae: Unioninae) in China currently includes six species, mainly distributed in the Yangtze River and Pearl River basins. In this study, a new species of this genus was discovered in the karst rivers of Guizhou Province, a transitional zone between the Yangtze River and Pearl River basins: Pseudocuneopsis heqing sp. nov. This new species can be distinguished from its congeners by its regularly elliptical shell shape and unique hinge teeth. The mitogenomic phylogenetic analyses strongly support the following species-level relationships: ((((P. sichuanensis + P. yemaoi) + (P. wuana + P. yangshuoensis) + P. heqing sp. nov.) + P. perflora) + P. capitata). The discovery of this new taxon contributes to the existing knowledge on freshwater mussels in China and implies that future comprehensive surveys of unexplored regions, particularly those with unique and scarce habitats, will unveil novel diversity.
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References
- Barrett RDH, Hebert PDN (2005) Identifying spiders through DNA barcodes. Canadian Journal of Zoology 83(3): 481–491. https://doi.org/10.1139/z05-024
- Böhm M, Dewhurst-Richman NI, Seddon M, Ledger SEH, Albrecht C, Allen D, Bogan AE, Cordeiro J, Cummings KS, Cuttelod A, Darrigran G, Darwall W, Fehér Z, Gibson C, Graf DL, Köhler F, Lopes-Lima M, Pastorino G, Perez KE, Smith K, van Damme D, Vinarski MV, von Proschwitz T, von Rintelen T, Aldridge DC, Aravind NA, Budha PB, Clavijo C, Van Tu D, Gargominy O, Ghamizi M, Haase M, Hilton-Taylor C, Johnson PD, Kebapçı Ü, Lajtner J, Lange CN, Lepitzki DAW, Martínez-Ortí A, Moorkens EA, Neubert E, Pollock CM, Prié V, Radea C, Ramirez R, Ramos MA, Santos SB, Slapnik R, Son MO, Stensgaard AS, Collen B (2021) The conservation status of the world's freshwater molluscs. Hydrobiologia 848(12): 3231–3254. https://doi.org/10.1007/s10750-020-04385-w
- Brauer CJ, Beheregaray LB (2020) Recent and rapid anthropogenic habitat fragmentation increases extinction risk for freshwater biodiversity. Evolutionary Applications 13(10): 2857–2869. https://doi.org/10.1111/eva.13128
- Brinkmann R, Parise M (2012) Karst environments: Problems, management, human impacts, and sustainability: An introduction to the special issue. Journal of Caves and Karst Studies 74(2): 135–136. https://doi.org/10.4311/2011JCKS0253
- Castresana J (2000) Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Molecular Biology and Evolution 17(4): 540–552. https://doi.org/10.1093/oxfordjournals.molbev.a026334
- Chen J (2020) Ecosystem of the Yangtze River Basin. In: Chen J (Ed.) Evolution and Water Resources Utilization of the Yangtze River. Changjiang Press (Wuhan) Co., Ltd. & Springer Nature Singapore Pte Ltd, Singapore, 163–220. https://doi.org/10.1007/978-981-13-7872-0_4
- Chen F, Xue G, Wang YK, Zhang HC, Clift PD, Xing YW, He J, Albert JS, Chen J, Xie P (2023) Evolution of the Yangtze River and its biodiversity. The Innovation 4(3): 100417. https://doi.org/10.1016/j.xinn.2023.100417
- Chen ZG, Dai YT, Chen H, Hu HF, Jiang J, Guo L, Ouyang S, Zhao XX, Wu XP (2025a) Gibbosula sinensis, a new species on the verge of extinction of the world's most threatened freshwater mussel family Margaritiferidae Henderson, 1929 from China (Bivalvia, Palaeoheterodonta, Unionida). Zoosystematics and Evolution 101(4): 1613–1620. https://doi.org/10.3897/zse.101.166584
- Chen H, Xiang HQ, Dai YT, Chen ZG, He YM, Guo G, Wu XP, Huang XC (2025b) A new genus and two new species of freshwater mussels (Bivalvia, Unionidae) from Sichuan, China: Overlooked cryptic endemism in the upper Yangtze River Basin. Zoosystematics and Evolution 101(4): 1459–1470. https://doi.org/10.3897/zse.101.161018
- Dai YT, Chen ZG, Peng KJ, Ouyang S, Huang XC, Wu XP (2024) Revisiting the genus Pseudocuneopsis (Bivalvia, Unionidae): Morphology, mitochondrial phylogenomics, and the description of a new species. Zoologica Scripta 53(3): 323–337. https://doi.org/10.1111/zsc.12647
- Do VT, Tuan LQ, Bogan AE (2018) Freshwater mussels (Bivalvia: Unionida) of Vietnam: diversity, distribution, and conservation status. Freshwater Mollusk Biology and Conservation 21(1): 1–18. https://doi.org/10.31931/fmbc.v21i1.2018.1-18
- Donath A, Jühling F, Al-Arab M, Bernhart SH, Reinhardt F, Stadler PF, Middendorf M, Bernt M (2019) Improved annotation of protein-coding genes boundaries in metazoan mitochondrial genomes. Nucleic Acids Research 47(20): 10543–10552. https://doi.org/10.1093/nar/gkz833
- Duan YF, Li M, Xu KW, Zhang L, Zhang LB (2021) Protect China's karst cave habitats. Science 374(6568): 699. https://doi.org/10.1126/science.abm5389
- Froufe E, Bolotov I, Aldridge DC, Bogan AE, Breton S, Gan HM, Kovitvadhi U, Kovitvadhi S, Riccardi N, Secci-Petretto G, Sousa R, Teixeira A, Varandas S, Zanatta D, Zieritz A, Fonseca MM, Lopes-Lima M (2020) Mesozoic mitogenome rearrangements and freshwater mussel (Bivalvia: Unionoidea) macroevolution. Heredity 124(1): 182–196. https://doi.org/10.1038/s41437-019-0242-y
- Guan DL, Xu SQ (2016) The complete mitogenome of the Asiatic Salamander Batrachuperus tibetanus (Amphibia: Caudata: Hynobiidae). Mitochondrial DNA. Part A, DNA Mapping, Sequencing, and Analysis 27(6): 4052–4053. https://doi.org/10.3109/19401736.2014.1003839
- Haag WR (2012) North American Freshwater Mussels: Natural History, Ecology, and Conservation. Cambridge University Press, 505–505. https://doi.org/10.1017/CBO9781139048217
- Haas F (1969) Superfamilia Unionacea. Das Tierreich 88: 1–663.
- He J, Zhuang Z (2013) The Freshwater Bivalves of China. ConchBooks, Germany, 198 pp.
- Hebert PDN, Cywinska A, Ball SL, deWaard JR (2003) Biological identifications through DNA barcodes. Proceedings. Biological Sciences 270(1512): 313–321. https://doi.org/10.1098/rspb.2002.2218
- Heude RP (1874) Conchyliologie fluviatile de la Province de Nanking et de la Chine centrale. Librairie F. Savy, Paris, France. https://doi.org/10.5962/bhl.title.16067
- Howard JK, Cuffey KM (2006) The functional role of native freshwater mussels in the fluvial benthic environment. Freshwater Biology 51(3): 460–474. https://doi.org/10.1111/j.1365-2427.2005.01507.x
- Inoue K, Hayes DM, Harris JL, Christian AD (2013) Phylogenetic and morphometric analyses reveal ecophenotypic plasticity in freshwater mussels Obovaria jacksoniana and Villosa arkansasensis (Bivalvia: Unionidae). Ecology and Evolution 3(8): 2670–2683. https://doi.org/10.1002/ece3.649
- Inoue K, Mcqueen AL, Harris JL, Berg DJ (2014) Molecular phylogenetics and morphological variation reveal recent speciation in freshwater mussels of the genera Arcidens and Arkansia (Bivalvia: Unionidae). Biological Journal of the Linnean Society. Linnean Society of London 112(3): 535–545. https://doi.org/10.1111/bij.12282
- Kalyaanamoorthy S, Minh BQ, Wong TKF, von Haeseler A, Jermiin LS (2017) ModelFinder: Fast model selection for accurate phylogenetic estimates. Nature Methods 14(6): 587–589. https://doi.org/10.1038/nmeth.4285
- Katoh K, Standley DM (2013) MAFFT multiple sequence alignment software version 7: Improvements in performance and usability. Molecular Biology and Evolution 30(4): 772–780. https://doi.org/10.1093/molbev/mst010
- Kumar S, Stecher G, Tamura K (2016) MEGA7: Molecular evolutionary genetics analysis version 7.0 for bigger datasets. Molecular Biology and Evolution 33(7): 1870–1874. https://doi.org/10.1093/molbev/msw054
- Lanfear R, Frandsen PB, Wright AM, Senfeld T, Calcott B (2017) Partitionfinder 2: New methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analyses. Molecular Biology and Evolution 34(3): 772–773.
- Letunic I, Bork P (2007) Interactive tree of life (iTOL): An online tool for phylogenetic tree display and annotation. Bioinformatics 23(1): 127–128. https://doi.org/10.1093/bioinformatics/btl529
- Li SL, Liu CQ, Chen JA, Wang SJ (2021) Karst ecosystem and environment: Characteristics, evolution processes, and sustainable development. Agriculture, Ecosystems & Environment 306: 107173. https://doi.org/10.1016/j.agee.2020.107173
- Liu YY, Zhang WZ, Wang QX, Wang EY (1979) Economic fauna of China: Freshwater Mollusks. Science Press, 134 pp.
- Liu XJ, Liu YY, Wu RW, Zanatta DT, Lopes-Lima M, Gonçalves DV, Bogan AE, Ouyang S, Wu XP (2022) Systematics, distribution, biology, and conservation of freshwater mussels (Bivalvia: Unionida) in China. Aquatic Conservation 32(5): 859–895. https://doi.org/10.1002/aqc.3799
- Liu LL, Zhang LP, Jin DD, Wang HT, Liu XJ, Wu RW (2023) Molecular and morphological evidence reveals a hidden new taxon in the endemic genus Pseudocuneopsis (Bivalvia, Unionidae) from China. ZooKeys 1179: 219–229. https://doi.org/10.3897/zookeys.1179.109817
- Long J, Wu QS, Li J, Liao HK, Liu LF, Huang BC, Zhang JM (2021) Effects of different microhabitat types on rock dissolution in Maolan Karst Forest, Southwest China. Turang Xuebao 58(1): 151–161.
- Lopes-Lima M, Froufe E, Do VT, Ghamizi M, Mock KE, Kebapçi U, Klishko O, Kovitvadhi S, Kovitvadhi U, Paulo OS, Pfeiffer JM III, Raley M, Riccardi N, Şereflişan H, Sousa R, Teixeira A, Varandas S, Wu X, Zanatta DT, Zieritz A, Bogan AE (2017) Phylogeny of the most species-rich freshwater bivalve family (Bivalvia: Unionida: Unionidae): Defining modern subfamilies and tribes. Molecular Phylogenetics and Evolution 106: 174–191. https://doi.org/10.1016/j.ympev.2016.08.021
- Lopes-Lima M, Burlakova LE, Karatayev AY, Mehler K, Seddon M, Sousa R (2018) Conservation of freshwater bivalves at the global scale: Diversity, threats and research needs. Hydrobiologia 810(1): 1–14. https://doi.org/10.1007/s10750-017-3486-7
- Lu XL (2024) A review of Karst environment and biodiversity in China. Advances in Environmental Protection 14(2): 348–353. https://doi.org/10.12677/aep.2024.142047
- Minh BQ, Nguyen MAT, von Haeseler A (2013) Ultrafast approximation for phylogenetic bootstrap. Molecular Biology and Evolution 30(5): 1188–1195. https://doi.org/10.1093/molbev/mst024
- Minh BQ, Schmidt HA, Chernomor O, Schrempf D, Woodhams MD, von Haeseler A, Lanfear R (2020) IQ-TREE 2: New models and efficient methods for phylogenetic inference in the genomic era. Molecular Biology and Evolution 37(5): 1530–1534. https://doi.org/10.1093/molbev/msaa015
- Reid AJ, Carlson AK, Creed IF, Eliason EJ, Gell PA, Johnson PTJ, Kidd KA, MacCormack TJ, Olden JD, Ormerod SJ, Smol JP, Taylor WW, Tockner K, Vermaire JC, Dudgeon D, Cooke SJ (2018) Emerging threats and persistent conservation challenges for freshwater biodiversity. Biological Reviews of the Cambridge Philosophical Society 94(3): 849–873. https://doi.org/10.1111/brv.12480
- Ronquist F, Teslenko M, van der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61(3): 539–542. https://doi.org/10.1093/sysbio/sys029
- Simpson CT (1900) Synopsis of the naiades, or pearly fresh-water mussels. Proceedings of the United States National Museum 22(1205): 501–1044. https://doi.org/10.5479/si.00963801.22-1205.501
- Simpson CT (1914) A Descriptive Catalogue of the Naiades, or Pearly Fresh-Water Mussels. Parts I–III. 1540 pp. https://doi.org/10.5962/bhl.title.10910
- Tickner D, Opperman JJ, Abell R, Acreman M, Arthington AH, Bunn SE, Cooke SJ, Dalton J, Darwall W, Edwards G, Harrison I, Hughes K, Jones T, Leclère D, Lynch AJ, Leonard P, McClain ME, Muruven D, Olden JD, Ormerod SJ, Robinson J, Tharme RE, Thieme M, Tockner K, Wright M, Young L (2020) Bending the curve of global freshwater biodiversity loss: An emergency recovery plan. Bioscience 70(4): 330–342. https://doi.org/10.1093/biosci/biaa002
- Vaughn CC (2018) Ecosystem services provided by freshwater mussels. Hydrobiologia 810(1): 15–27. https://doi.org/10.1007/s10750-017-3139-x
- Vaughn CC, Nichols SJ, Spooner DE (2008) Community and foodweb ecology of freshwater mussels. Journal of the North American Benthological Society 27(2): 409–423. https://doi.org/10.1899/07-058.1
- Walker JM, Bogan AE, Bonfiglio EA, Campbell DC, Christian AD, Curole JP, Harris JL, Wojtecki RJ, Hoeh WR (2007) Primers for amplifying the hypervariable, male-transmitted COII-COI junction region in amblemine freshwater mussels (Bivalvia: Unionoidea: Ambleminae). Molecular Ecology Notes 7(3): 489–491. https://doi.org/10.1111/j.1471-8286.2006.01630.x
- Wang RQ, Jiang XH, Nie T (2022) Spatiotemporal variation of NDVI and its driving forces in the Pearl River Basin based on geographical detector. People's. Renmin Zhujiang 43(7): 61–73.
- Wu RW, Liu XJ, Guo L, Zhou CH, Ouyang S, Wu XP (2022a) DNA barcoding, multilocus phylogeny, and morphometry reveal phenotypic plasticity in the Chinese freshwater mussel Lamprotula caveata (Bivalvia: Unionidae). Ecology and Evolution 12(7): e9035. https://doi.org/10.1002/ece3.9035
- Wu XP, Dai YT, Yin N, Shu FY, Chen ZG, Guo L, Zhou CH, Ouyang S, Huang XC (2022b) Mitogenomic phylogeny resolves Cuneopsis (Bivalvia: Unionidae) as polyphyletic: The description of two new genera and a new species. Zoologica Scripta 51(2): 173–184. https://doi.org/10.1111/zsc.12527
- Wu RW, Liu LL, Zhang LP, Jia JL, Jin DD, Wu XP, Liu XJ (2023a) New species of the genus Pseudocuneopsis Huang, Dai, Chen & Wu, 2022 (Bivalvia, Unionidae) from Guangxi Province, China. ZooKeys 1166: 261–270. https://doi.org/10.3897/zookeys.1166.104150
- Wu RW, Liu LL, Liu XJ, Ye YY, Wu XP, Xie ZC, Liu ZY, Li ZF (2023b) Towards a systematic revision of the superfamily Cyrenoidea (Bivalvia, Imparidentia): Species delimitation, multi-locus phylogeny and mitochondrial phylogenomics. Invertebrate Systematics 37(9): 607–622. https://doi.org/10.1071/IS23015
- Wu RW, Liu LL, Zhang LP, Liu XJ, Hu ZK, Jin DD, Zhang ZP, Wu XP, Xie ZC, Li ZF, Lopes-Lima M (2024a) Diversity, morphology, and phylogeny of freshwater mussels of the genus Nodularia (Bivalvia: Unionidae) from China, with descriptions of four new species. Zoologica Scripta 53(5): 594–613. https://doi.org/10.1111/zsc.12677
- Wu RW, Liu LL, Zhang LP, Bogan AE, Niu GY, Jin DD, Wu XP, Liu XJ (2024b) Taxonomic revision of two species in the genus Ptychorhynchus Simpson, 1900 (Bivalvia, Unionidae, Gonideinae), with description of a new species. Invertebrate Systematics 38(7): IS24014. https://doi.org/10.1071/IS24014
- Wu RW, Zhang LP, Liu LL, Jia JL, Liu XJ (2024c) Unraveling the Phylogenetic Relationships and Taxonomic Status of a Puzzling Freshwater Mussel Genus Inversidens (Bivalvia, Unionidae) through Multilocus Phylogeny and Mitochondrial Phylogenomics. Journal of Zoological Systematics and Evolutionary Research 2024(1): 1499508. https://doi.org/10.1155/2024/1499508
- Zhang D, Gao F, Jakovlić I, Zhou H, Zhang J, Li WX, Wang GT (2020) PhyloSuite: An integrated and scalable desktop platform for streamlined molecular sequence data management and evolutionary phylogenetics studies. Molecular Ecology Resources 20(1): 348–355. https://doi.org/10.1111/1755-0998.13096
- Zheng HB, Clift PD, Wang P, Tada R, Jia JT, He MY, Jourdan F (2013) Pre-Miocene birth of the Yangtze River. Proceedings of the National Academy of Sciences of the United States of America 110(19): 7556–7561. https://doi.org/10.1073/pnas.1216241110
- Zheng S, Poczai P, Hyvönen J, Tang J, Amiryousefi A (2020) Chloroplot: An online program for the versatile plotting of organelle genomes. Frontiers in Genetics 11: 576124. https://doi.org/10.3389/fgene.2020.576124
- Zieritz A, Aldridge DC (2009) Identification of ecophenotypic trends within three European freshwater mussel species (Bivalvia: Unionoida) using traditional and modern morphometric techniques. Biological Journal of the Linnean Society. Linnean Society of London 98(4): 814–825. https://doi.org/10.1111/j.1095-8312.2009.01329.x
- Zieritz A, Bogan AE, Froufe E, Klishko O, Kondo T, Kovitvadhi U, Kovitvadhi S, Lee JH, Lopes-Lima M, Pfeiffer JM, Sousa R, Do VT, Vikhrev I, Zanatta DT (2018) Diversity, biogeography and conservation of freshwater mussels (Bivalvia: Unionida) in East and Southeast Asia. Hydrobiologia 810(1): 29–44. https://doi.org/10.1007/s10750-017-3104-8
- Zieritz A, Froufe E, Bolotov I, Gonçalves DV, Aldridge DC, Bogan AE, Gan HM, Gomes-Dos-Santos A, Sousa R, Teixeira A, Varandas S, Zanatta D, Lopes-Lima M (2021) Mitogenomic phylogeny and fossil-calibrated mutation rates for all F-and M-type mtDNA genes of the largest freshwater mussel family, the Unionidae (Bivalvia). Zoological Journal of the Linnean Society 193(3): 1088–1107. https://doi.org/10.1093/zoolinnean/zlaa153