Zoosyst. Evol. 101 (2) 2025, 779-790 | DOI 10.3897/zse.101.151083 nett BERLIN A new tribe, genus, and species of freshwater mussel from the Changjiang River Basin in China (Bivalvia, Unionidae, Unioninae) Yu-Ting Dai!*, Zhong-Guang Chen!*, Shan Ouyang', Xiao-Chen Huang!, Xiao-Ping Wu! 1 School of Life Sciences, Nanchang University, Nanchang 330031, China https://zoobank. org/219F 8CF 1-F 488-4B0C-9C65-FS&FD9833 1040 Corresponding authors: Xiao-Chen Huang (xiaochenhuang@hotmail.com); Xiao-Ping Wu (xpwu@ncu.edu.cn) Academic editor: Matthias Glaubrecht # Received 22 February 2025 Accepted 26 March 2025 Published 10 April 2025 Abstract A new tribe, genus, and species of freshwater mussels, Globunionini Dai, Chen, Huang & Wu, tribe nov. and Globunio mirificus Chen, Dai, Huang & Wu, gen. et sp. nov., is described from the Changjiang River Basin in China based on comparative morphology and molecular phylogeny. The species presence of a special small-sized and elongated globular, which distinguishes it from all other tribes. The discovery increases the diversity of freshwater mussels in the Changjiang River Basin. Key Words Biodiversity, molluscs, phylogeny, taxonomy Introduction Freshwater mussels (Mollusca: Bivalvia: Unionida) play an important role in freshwater ecosystems (Graf and Cummings 2007; Huang et al. 2019) and are one of the most threatened freshwater organisms globally (Vaughn 2018; Lopes-Lima et al. 2020; Bohm et al. 2021). China is one of the regions with the highest spe- cies diversity of freshwater mussels in the world, with about 100 accepted species recorded and new taxa con- tinuing to be discovered (Heude 1875, 1877a, 1877b, 1878, 1879, 1880a, 1880b, 1881, 1883, 1885; Simpson 1900; He and Zhuang 2013; Graf and Cummings 2021; Guo 2022; Liu et al. 2022; Wu et al. 2022a, 2022b; Chen et al. 2023; Dai et al. 2023, 2024a, 2024b, 2024c, 2024d). The freshwater mussels in China are mainly concentrated in the Changjiang River Basin (Yangtze), with about 70 accepted species recorded (Heude 1875, 1877a, 1877b, 1878, 1879, 1880a, 1880b, 1881, 1883, * These authors contributed equally to this work. 1885; Simpson 1900; He and Zhuang 2013; Graf and Cummings 2021; Guo 2022; Chen et al. 2023; Dai et al. 2024a, 2024b). Even in the well-studied middle and lower reaches of the Changjiang River, new freshwa- ter mussels have been continuously discovered in re- cent years, demonstrating the extremely high diversity in the region (Guo 2022; Chen et al. 2023; Dai et al. 2023, 2024a, 2024b; Wu et al. 2024). During the surveys in 2022, we discovered a group of freshwater mussel specimens with the special small- sized and elongated-globular shell that did not resemble any known species and were challenging to place in any genus or tribe. Based on a combination of morphology and molecular phylogeny, we describe it as a new tribe, genus, and species of subfamily Unioninae Rafinesque, 1820. The discovery improved the diversity of freshwater mussels in the Changjiang River Basin and showed that even in the central city, there are still yet-to-be-described species of freshwater mussels that exist. Copyright Dai, Y.-T. et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 780 Materials and methods Materials preparation Specimens were collected from Jiangxi, China, in 2022. Living specimens were initially frozen at -20 °C for 24 hours and subsequently thawed at room temperature for 2 hours to facilitate the extraction of soft parts. The soft parts were then fixed in 95% ethanol. Empty shells were cleaned, dried, and preserved at room temperature. Pho- tographs were taken by camera and edited in Adobe Pho- toshop CC 2015 (Adobe, San Jose, US). Maps were made in ArcGIS Pro (Esri, Redlands, US). DNA extraction, sequencing, and mitogenome assembly Genomic DNA was extracted from muscle preserved in 95% ethanol using a TIANamp Marine Animals DNA Kit (Tiangen Biotech, China). The quality and concen- tration of the DNA were checked on 1% agarose gel electrophoresis and NanoDrop 2000 (Thermo Scien- tific, USA). The qualified genomic DNA sample was sent to Novogene (Beijing, China). Two datasets were used for phylogenetic analyses: the partial cytochrome c oxidase subunit 1 (CO/) sequences and the mitochon- drial genomes. The polymerase chain reaction (PCR) systems, conditions, and primer pairs of CO/ are fol- lowed by Chen et al. (2023). After quality controls, the library was successfully prepared and sequenced in the Illumina NovaSeq 6000 platform, yielding 12 Gb of Dai, Y.-T. et al.: A new tribe, genus, and species of freshwater mussel data, with 2 x 150 bp paired-end reads. After discarding low-quality reads, clean reads were obtained and assem- bled de novo using CLC Genomic Workbench v. 12.0 (https://digitalinsights.qiagen.com). Contigs identified as mitogenome sequences were inspected manually for overlap at the beginning and end, resulting in a circular mitogenome. Geneious v. 11.0 (https://www.geneious. com) (Kearse et al. 2012) was used to check the com- plete mitogenome and analyze nucleotide composition. Strand asymmetry was calculated using the formulas: GC skew = (G-C)/(G+C) and AT skew = (A-T)/((A+T). The initial annotation of the mitogenome was carried out using the MITOS web server (http://mitos.bioinf. uni-leipzig.de/index.py) (Bernt et al. 2013). ARWEN (http://130.235.244.92/ARWEN) (Laslett and Canback 2008) was also used to identify the locations of all tRNA genes. The annotations of two rRNA genes were fur- ther refined based on the positions of neighboring genes. PCGs were initially identified using the Open Reading Frame Finder (ORF Finder) implemented on the NCBI website (http://www.ncbi.nlm.nih.gov/orffinder) and BLAST searches (http://blast.ncbi.nlm.nih. gov). Alignments, partitioning strategies The COI dataset consists of 51 sequences of the fami- ly Unionidae and two outgroup taxa (Table 1). The mi- tochondrial genome dataset consists of 81 sequences (including 12 PCGs (excluding ATP8) and two rRNA genes) of the family Unionidae and nine outgroup taxa (Table 2). The CO/ sequences were aligned using MEGA Table 1. GenBank accession numbers of the CO/ sequences used in this study. Species Unioninae Rafinesque, 1820 Globunio mirificus gen. et sp. nov. 1 Globunio mirificus gen. et sp. nov. 2 Globunio mirificus gen. et sp. nov. 3 Globunio mirificus gen. et sp. nov. 4 Globunio mirificus gen. et sp. nov. 5 Globunio mirificus gen. et sp. nov. 6 Globunio mirificus gen. et sp. nov. 7 Globunio mirificus gen. et sp. nov. 8 Globunio mirificus gen. et sp. nov. 9 Diaurora aurorea (Heude, 1883) Acuticosta chinensis (Lea, 1868) Schistodesmus lampreyanus (Baird & Adams, 1867) Schistodesmus spinosus (Simpson, 1900) Schistodesmus sp. Unio pictorum (Linnaeus, 1758) Unio crassus Philipsson, 1788 Tchangsinaia piscicula (Heude, 1874) Cuneopsis celtiformis (Heude, 1874) Cuneopsis heudei (Heude, 1874) Aculamprotula fibrosa (Heude, 1877) Nodularia douglasiae (Gray, 1833) Nodularia breviconcha Lee, Kim, Bogan & Kondo, 2020 zse.pensoft.net Locality Accession References number Honggutan, Nanchang, Jiangxi, China PV368604 This study Honggutan, Nanchang, Jiangxi, China PV368605 This study Honggutan, Nanchang, Jiangxi, China PV368606 This study Honggutan, Nanchang, Jiangxi, China PV368607 This study Honggutan, Nanchang, Jiangxi, China PV368608 This study Honggutan, Nanchang, Jiangxi, China PV368609 This study Honggutan, Nanchang, Jiangxi, China PV368610 This study Honggutan, Nanchang, Jiangxi, China PV368611 This study Honggutan, Nanchang, Jiangxi, China PV368612 This study Jian, Jiangxi, China 0Q829360* Chen et al. 2023 Jiangxi, China MG462921 Huang et al. 2019 Jiangxi, China MG463037 Huang et al. 2019 Jiangxi, China MG463046 Huang et al. 2019 Hunan, China MG463043 Huang et al. 2019 Europe KC429109 Sharma et al. 2013 Poland KY290446 Burzynski et al. 2017 Jiangxi, China MG462977 Huang et al. 2019 Jiangxi, China MG462964 Huang et al. 2019 Jiangxi, China MG462970 Huang et al. 2019 Jiangxi, China MG462909 Huang et al. 2019 China KX822653 Lopes-Lima et al. 2017 South Korea MT020662 ~~ Lopes-Lima et al. 2020 Zoosyst. Evol. 101 (2) 2025, 779-790 Species Inversiunio yanagawensis (Kondo, 1982) Inversiunio reinianus (Kobelt, 1879) Pseudocuneopsis sichuanensis Huang, Dai, Chen & Wu, 2022 Pseudocuneopsis capitata (Heude, 1874) Alasmidonta marginata Say, 1818 Lasmigona compressa (Lea, 1829) Anodonta anatina (Linnaeus, 1758) Pseudanodonta complanata (Rossmassler, 1835) Lanceolaria gladiola (Heude, 1877) Lanceolaria oxyrhyncha (Martens, 1861) Cristaria plicata (Leach, 1814) Lepidodesma languilati (Heude, 1874) Sinanodonta woodiana (Lea, 1834) Beringiana beringiana (Middendorff, 1851) Pletholophus tenuis (Gray, 1833) Anemina arcaeformis (Heude, 1877) Amuranodonta kijaensis Moskvicheva, 1973 Parreysiinae Henderson, 1935 Coelatura aegyptiaca (Cailliaud, 1823) Indonaia andersoniana (Nevill, 1877) Parreysia nagpoorensis (Lea, 1860) Gonideinae Ortmann, 1916 Pronodularia japanensis (Lea, 1859) Lamprotula leaii (Gray, 1833) Ambleminae Rafinesque, 1820 Lampsilis siliquoidea (Barnes, 1823) Quadrula quadrula (Rafinesque, 1820) Margaritiferidae Henderson, 1929 Margaritifera dahurica (Middendorff, 1850) Gibbosula rochechouartii (Heude, 1875) X (Kumar et al. 2018) and checked manually. MUSCLE (Edgar 2004) was used for codon alignment of all PCGs implemented in MEGA X (Kumar et al. 2018). MAFFT (Katoh et al. 2019) with the Q-INS-1i algorithm was used for the alignment of ribosomal genes (12S and 16S rR- NAs). Gblocks (Castresana 2000) were used to exclude ambiguous regions from the alignment of each gene. Par- titionFinder (Lanfear et al. 2017) was used to determine the best-fit partitioning schemes and substitution models under greedy search. Predefined data blocks for parti- tioning scheme searches were designated by gene region (rRNA gene) or codon position (PCG). Branch lengths were allowed to be unlinked, and model selection and partitioning schemes were determined using the AlCc method (Table 3). Phylogenetic analyses ML analyses were performed in IQ-TREE v. 1.6.12 (Minh et al. 2013) using the Ultrafast bootstrap approach (Minh et al. 2013) with 10,000 iterations. Bayesian in- ference (BI) analysis was conducted in MrBayes v. 3.2.6 (Ronquist et al. 2012). Four simultaneous runs with four independent Markov Chain Monte Carlo (MCMC) were implemented for 10 million generations, and trees were sampled every 10,000 generations with a burn-in 781 Locality Accession References number Japan MT020654 ~~ Lopes-Lima et al. 2020 Japan MT020657 ~—Lopes-Lima et al. 2020 Sichuan, China MZ540966 Wu et al. 2022b Anhui, China NC042469 Wu et al. 2019 U.S. AF156502 ~— Graf & O'Foighil 2000 U.S. AF156503 ~— Graf & O'Foighil 2020 Russia KX822632 Lopes-Lima et al. 2017 Ukraine KX822661 Lopes-Lima et al. 2017 Jiangxi, China KY067441 Unpublished Japan MT020648 Lopes-Lima et al. 2020 Jiangxi, China MG462956 Huang et al. 2019 Jiangxi, China MG463015 Huang et al. 2019 China KX822668 ~~ Lopes-Lima et al. 2017 Japan MT020557 ~— Lopes-Lima et al. 2020 Vietnam KX822658 ~— Lopes-Lima et al. 2016 Jiangxi, China MG462936 Huang et al. 2019 Russia MK574204 Bolotov et al. 2020 Egypt KJ081162 Graf et al. 2014 Myanmar MF352275 Bolotov et al. 2017 India JQ861229 Unpublished Japan LC505454 __s‘Fukata and ligo 2020 Jiangxi, China MG462996 Huang et al. 2019 WES: MH560773 Unpublished Urs: HM230409 Unpublished Russia KJ161516 Bolotov et al. 2015 Jiangxi, China MG463022 Huang et al. 2019 of 25%. The convergence was checked with the average standard deviation of split frequencies < 0.01 and the potential scale reduction factor (PSRF) ~ 1. Trees were visualized in FigTree v.1.4.3 (http://tree.bio.ed.ac.uk/ software/figtree/). Abbreviations NCU_XPWU: Laboratory of Xiao-Ping Wu, Nanchang University (Nanchang, Jiangxi, China). Results COI phylogeny The alignment of the CO/ gene had lengths of 606 char- acters. Within these alignments, 260 sites were variable, and 236 sites were parsimony informative. The maximum likelihood and Bayesian analyses produced consistent phylogenies (Figs 1, 2). Although a single gene marker cannot effectively address inter tribes relationships in the subfamily Unioninae, the new species formed an inde- pendent branch that does not belong to any tribe. Its sys- tematic position needs further verification through mito- chondrial genome phylogenetic analysis. zse.pensoft.net 782 Dai, Y.-T. et al.: A new tribe, genus, and species of freshwater mussel Table 2. GenBank accession numbers of the mitochondrial genome sequences used in this study. Species Accession number Unioninae Rafinesque, 1820 Globunio mirificus gen. et sp. nov. PV394654 Tchangsinaia pisciculus (Heude, 1874) NC026306 Cuneopsis heudei (Heude, 1874) NC042471 Cuneopsis rufescens (Heude, 1874) MZ571512 Cuneopsis celtiformis (Heude, 1874) MW464617 Cuneopsis demangei Haas, 1929 MZ571513 Pseudocuneopsis capitata (Heude, 1874) NC042469 Pseudocuneopsis capitata (Heude, 1874) MZ571517 Pseudocuneposis sichuanensis Huang, Dai, Chen & Wu, 2022 MZ571510 Pseudocuneopsis yemaoi Dai, Chen, Huang & Wu, 2024 OR392755 Pseudocuneopsis yangshuoensis Wu & Liu, 2023 OR392756 Schistodesmus lampreyanus (Baird & Adams, 1867) NC042470 Schistodesmus spinosus (Simpson, 1900) MZ571511 Unio crassus Philipsson, 1788 KY290446 Unio delphinus Spengler, 1793 KT326917 Unio pictorum (Linnaeus, 1 758) NCO15310 Unio tumidus Philipsson, 1788 KY021076 Nodularia breviconcha Lee, Kim, Bogan & Kondo MT955592 Nodularia douglasiae (Griffith & Pidgeon, 1833) NCO026111 Nodularia fusiformans Wu & Liu, 2024 MT764726 Nodularia nuxpersicae (Dunker, 1848) OR888962 Nodularia nuxpersicae (Dunker, 1848) OR888961 Aculamprotula coreana (Martens, 1886) NC026035 Aculamprotula polysticta (Heude, 1877) MK728823 Aculamprotula scripta (Heude, 1875) MF991456 Aculamprotula tientsinensis (Crosse & Debeaux, 1863) NC029210 Aculamprotula tortuosa (Lea, 1865) NC0O21404 Acuticosta chinensis (Lea, 1868) MF687347 Anodonta anatina (Linnaeus, 1758) NC022803 Anodonta cygnea (Linnaeus, 1758) NC036488 Sinanodonta lucida (Heude, 1877) NC026673 Sinanodonta woodiana (Lea, 1834) HQ283346 Cristaria plicata (Leach, 1814) NCO12716 Anemina arcaeformis (Heude, 1877) NC026674. Anemina euscaphys (Heude, 1879) NC026792 Utterbackia imbecillis (Say, 1829) NCO15479 Utterbackia peninsularis Bogan & Hoeh, 1995 HM856636 Pyganodon grandis (Say, 1829) NCO13661 Lasmigona compressa (Lea, 1829) NCO15481 Lanceolaria gladiola (Heude, 1877) KY067441 Lanceolaria grayili (Gray, 1833) NCO26686 Lanceolaria lanceolata (Lea, 1856) NC023955 Lepidodesma languilati (Heude, 1874) NC029491 Gonideinae Ortmann, 1916 Sinosolenaia carinata (Heude, 1877) NC023250 Sinosolenaia oleivora (Heude, 1877) NC022701 Ptychorhynchus pfisteri (Heude, 1874) KY067440 Parvasolenaia rivularis (Heude, 1877) KX966393 Inversidens rentianensis Wu & Wu, 2021 OR823224 Postolata guangxiensis Dai, Huang, Guo & Wu, 2023 OP009366 Microcondylaea bonellii (Férussac, 1827) NC044111 Monodontina vondembuschiana (Lea, 1840) NC044112 Pilsbryoconcha exilis (Lea, 1838) NC044124 Lamprotula gottscherSchistodesmus.sp NC023806 Lamprotula leaii (Griffith & Pidgeon, 1833) NC023346 Lamprotula caveata (Heude, 1877) NC030336 Pronodularia japanensis (Lea, 1859) AB055625 Potomida littoralis (Cuvier, 1798) NC030073 Sinohyriopsis cumingii (Lea, 1852) NCO11763 Sinohyriopsis schlegelii (Martens, 1861) NCO15110 Chamberlainia hainesiana (Lea, 1856) NC044110 Lens contradens (Lea, 1838) MW242812 Physunio superbus (Lea, 1843) MW242814 Hyriopsis bialata Simpson, 1900 MW242816 Rectidens sumatrensis (Dunker, 1852) MW242818 Lampsilis powellii (Lea, 1852) NC037720 zse.pensoft.net Zoosyst. Evol. 101 (2) 2025, 779-790 783 Species Accession number Lampsilis siliquoidea (Barnes, 1823) NC037721 Lampsilis cardium Rafinesque, 1820 BKO10478 Lampsilis ornata (Conrad, 1835) NC005335 Venustaconcha ellipsiformis (Conrad, 1836) FJ809753 Leaunio lienosus (Conrad, 1834) BKO10479 Potamilus leptodon (Rafinesque, 1820) NC028522 Potamilus alatus (Say, 1817) KU559010 Toxolasma parvum (Barnes, 1823) NC015483 Popenaias popeii (Lea, 1857) NC050058 Amblema plicata (Say, 1817) NC050056 Elliptio complanata (Lightfoot, 1786) BKO10477 Pleurobema oviforme (Conrad, 1834) NC050057 Quadrula quadrula (Rafinesque, 1820) NC013658 Uniomerus tetralasmus (Say, 1831) BKO10480 Margaritiferidae Henderson, 1929 Margaritifera dahurica (Middendorff, 1850) NC023942 Margaritifera margaritifera (Linnaeus, 1758) MK421958 Margaritifera falcata (Gould, 1850) NC015476 Pseudunio marocanus (Pallary, 1918) KY131953 Cumberlandia monodonta (Say, 1829) KU873123 Gibbosula rochechouartii (Heude, 1875) KX378172 Hyriidae Swainson, 1840 Echyridella menziesii (Gray, 1843) KU873121 lridinidae Swainson, 1840 Mutela dubia (Gmelin, 1791) KU873120 Mycetopodidae Gray, 1840 Anodontites trapesialis (Lamarck, 1819) KU873119 99) Inversiunio yanagawensis 63 Inversiunio reinianus 76 Tchangsinaia pisciculus 78 100 Unio pictorum Unio crassus 95 Pseudocuneopsis Sichuanensis Pseudocuneopsis capitata 69;-- Schistodesmus sp. Unionini 82 Schistodesmus spinosus 68 Schistodesmus lampreyanus Diaurora aurorea 63 98 Cuneopsis celtiformis Cuneopsis heudei as 99 Nodularia douglasiae Nodularia breviconcha 73 Middendorffinaia mongolica 68 Aculamprotula fibrosa 99 Aculamprotula coreana 100 100; Aculamprotula scripta Aculamprotulini Gibbosula polysticta Cc 82 Aculamprotula tortuosa 5 Globunio mirificus gen. & sp. nov. 1 roy Globunio mirificus gen. & sp. nov. 2 5 Globunio mirificus gen. & sp. nov. 3 =) Globunio mirificus gen. & sp. nov. 4 rab) 66 Globunio mirificus gen. & sp. nov. 6 Globunionini trib. nov. © Globunio mirificus gen. & sp. nov. 5 Globunio mirificus gen. & sp. nov. 7 78 66 97 73 70 90 Globunio mirificus gen. & sp. Globunio mirificus gen. & sp. Acuticosta chinensis Lepidodesma languilati Lanceolaria gladiola Lanceolaria oxyrhyncha Cristaria plicata Alasmidonta marginata Lasmigona compressa Anodonta anatina nov. 9 nov. 8 Acuticostini Lepidodesmini Lanceolariini sepiuoiun Pseudanodonta complanata 94 Sinanodonta woodiana 97 Beringiana beringiana Pletholophus tenuis Anemina arcaeformis Amuranodonta kijaensis 88 Coelatura aegyptiaca Indonaia andersoniana Parreysia corrugata Lampsilis siliquoidea Quadrula quadrula 94 Pronodularia japanensis Lamprotula leaii Gibbosula rochechouartii Margaritifera dahurica 0.2 Anodontini 92 98 77 pie 100 Parreysiinae 100 . ee Ambleminae Gonideinae 100 | outgroups Figure 1. Maximum likelihood tree inferred from CO/ gene sequences. Bootstrap supports are shown on the left of nodes on the tree if greater than 50%. zse.pensoft.net 784 0.98 0.98 0.98 0.53 Lanceolaria gladiola Lanceolaria oxyrhyncha Pseudanodonta complanata In Lamprotula leaii Lampsilis siliquoidea Quadrula quadrula Dai, Y.-T. et al.: A new tribe, genus, and species of freshwater mussel 1 Unio pictorum Unio crassus Cuneopsis pisciculus 1; Inversiunio yanagawensis Inversiunio reinianus Pseudocuneopsis Sichuanensis Pseudocuneopsis capitata Schistodesmus lampreyanus 1 Schistodesmus sp. Schistodesmus spinosus Diaurora aurorea 1 Cuneopsis celtiformis Cuneopsis heudei 1 Nodularia douglasiae Nodularia breviconcha Middendorffinaia mongolica 0.62 Aculamprotula fibrosa 1 Aculamprotula coreana 1 Aculamprotula scripta Gibbosula polysticta Aculamprotula tortuosa Globunio mirificus gen. & sp. nov. 1 Globunio mirificus gen. & sp. nov. 2 Globunio mirificus gen. & sp. nov. 3 Globunio mirificus gen. & sp. nov. 7 Globunio mirificus gen. & sp. nov. 8 Globunio mirificus gen. & sp. nov. 9 Globunio mirificus gen. & sp. nov. 4 Globunio mirificus gen. & sp. nov. 5 Globunio mirificus gen. & sp. nov. 6 Acuticosta chinensis Lepidodesma languilati Unionini Aculamprotulini Globunionini trib. nov. seuluoIun sepiuoiun Acuticostini Lepidodesmini Lanceolariini Cristaria plicata Alasmidonta marginata Lasmigona compressa Anodonta anatina Sinanodonta woodiana Anodontini Beringiana beringiana Pletholophus tenuis Anemina arcaeformis Amuranodonta kijaensis Coelatura aegyptiaca donaia andersoniana Parreysia corrugata Pronodularia japanensis Parreysiinae Gonideinae Ambleminae 1 Gibbosula rochechouartii Margaritifera dahurica 0.1 outgroups Figure 2. Bayesian inference tree inferred from CO/ gene sequences. Posterior probabilities are shown on the left of nodes on the tree if greater than 50%. Table 3. Partitioning strategies for the mitogenome dataset (from PartitionFinder2) according to AICc. Subset Best Model Partition scheme 1 GTR+l+G COX1 codonl, ATP6 codonl, ND4 codonl1, ND3 codon1, ND5 codon1, ND4L codonl, CO3 codonl 2 GTR+I+G ND5 codon2, ND3 codon2, ND4L codon2, COX1 codon2, CO3 codon2, ND4 codon2, ATP6 codon2 3 GTR+I+G CO2 codon3, ND4L codon3, ND3 codon3, ATP6 codon3, ND4 codon3, ND5 codon3, CO3 codon3, COX1 codon3 4 SYM+I+G ND1 codonl, CYTB codonl, CO2 codon2, CO2 codonl 5 GTR+I+G ND1 codon2, ND6 codon2, ND2 codon2, CYTB codon2 6 GTR+G ND1 codon3, ND6 codon3, CYTB codon3, ND2 codon3 f GTR+I+G 16S, 12S, ND2 codonl1, ND6 codon1l Mitogenome characteristics The length of the mitogenome is 16,240 bp. The mater- nal mitogenomes of the new species contain the 13 PCGs typically found in metazoan mitochondrial genomes, the type-specific F-orf described for all Unionida mitoge- nomes with the DUI system, 22 transfer RNA (tRNA), and two ribosomal RNA (rRNA) genes (Fig. 3A). Most genes zse.pensoft.net were encoded on the light strand (L-strand), whereas 11 genes (COI, COI, COMI, ND3, ND4, ND4L, NDS5, ATP6, ATP8, tRNA“, and tRNA") were located on the heavy strand (H-strand). The location of the F-orf was between ND2 and tRNA°", and the gene order of the mitogenomes was consistent with other Unioninae and Ambleminae spe- cies. The rrnS gene of it was located between tRNA“" and tRNA’, while the rmL gene was located between tRNA™ Zoosyst. Evol. 101 (2) 2025, 779-790 1001 Globunio mirificus 16,240 bp 92/1 7310.99 100/11 Hicomplex | (NADH dehydrogenase) Ecomplex III (ubichinol cytochrome c reductase) EI complex IV (cytochrome c oxidase) ribosomal RNA transfer RNA MIF-ORF BIATP synthase 100/4 100/11 a_ BI topology Cuneopsis 51/0.84 Tchangsinaia 100/ Schistodesmus 100/1 Nodularia 8710.95 96/1 : 100/1 Pseudocuneopsis 100/1 100/1 1001 100/1 1001 100/1 100/11 100/1 100/1 100/1 100/11 100/1 100/11 75/0.92 Elliptio co. Q 0.2 Figure 3. A. Gene map of the F-type mitochondrial genome of 100/1} 4100/1 Pronodularia japanensis Potomida littoralis 100/ Popenaias popeii Amblema plicata Pleurobema oviforme Gibbosula rochechouarti Hyridella menziesii 785 100/1 76/1 100/ Nodularia nuxpersicae Nodularia nipponensis Nodularia douglasiae Nodularia fusiformans Nodularia breviconcha Pseudocuneposis sichuanensis Pseudocuneopsis yemaoi Pseudocuneopsis yangshuoensis Pseudocuneopsis capitata 100/1- Cuneopisis heudei Cuneopsis demangei Unionini Cuneopsis celtiformis Cuneopsis rufescens Tchangsinaia pisciculus Schistodesmus sp. Schistodesmus lampreyanus Schistodesmus spinosus 100/1 100/0.69 Unio elongatulus Unio mancus Unio pictorum Unio delphinus Unio crassus Unio tumidus Aculamprotula polysticta Aculamprotula scripta Aculamprotula coreana Aculamprotula tientsinensis Aculamprotula tortuosa Globunio mirificus gen. & sp. nov. Utterbackia imbecillis 100/1 Aculamprotulini seuIuoIup Globunionini trib. nov. Utterbackia peninsularis Pyganodon grandis Lasmigona compressa Anodonta anatina Anodonta cygnea 100/- Anemina arcaeformis Anemina euscaphys Sinanodonta lucida Sinanodonta woodiana Cristaria plicata Lanceolaria lanceolata Lanceolaria grayana Lanceolaria gladiola Acuticosta chinensis Anodontini 100/1 Lanceolariini Acuticostini Lepidodesma languilati Lepidodesmini Postolata guangxiensis Ptychorhynchus pfisteri Parvasolenaia rivularis Sinosolenaia carinata Sinosolenaia oleivora Microcondylaea bonellii Inversidens rentianensis Monodontina vondembuschiana Pilsbryoconcha exilis Lamprotula caveata Lamprotula leaii sepiuoiun Gonideini Pseudodontini Lamprotulini Contradens contradens Physunio superbus Hyriopsis bialata Rectidens sumatrensis Contradentini Rectidentini SeUISPIUOS 100/1 Sinohyriopsis cumingii Sinohyriopsis schlegelii Chamberlainia hainesiana Chamberlainiini Lampsilis powellii Lampsilis siliquoidea Venustaconcha ellipsiformis Lampsilis cardium Lampsilis ornata Villosa lienosa Potamilus streckersoni Potamilusalatus Leptodea leptodon Toxolasma parvum Lampsilini Popenaiadini Amblemini Pleurobemini Quadrulini SeUIWO|GUIYY mplanata uadrula quadrula Uniomerus tetralasmus Margaritifera dahurica Margaritifera margaritifera Margaritifera falcata Margaritifera marocana Cumberlandia monodonta (e) utg roups Anodontites trapesialis Mutela dubia Globunio mirificus gen. et sp. nov.; B. Maximum likelihood and Bayesian inference tree inferred from mitochondrial genome sequences. Bootstrap supports/posterior probabilities are shown on the left/right of nodes on the tree if they are greater than 50%. and tRNA‘. The overall base composition was A (25.7%), T (38.1%), C (12.2%), and G (24.0%), with the A + T con- tent of 63.8%. Nucleotide asymmetry of the mitochondrial strands was assessed by AT skew and GC skew that were —0.19 and 0.33, respectively. Mitochondrial phylogeny The alignment of the mitogenomes had lengths of 11,547 characters. Within these alignments, 7,300 sites were variable, and 6,604 sites were parsimony informative. The maximum likelihood and Bayesian analyses pro- duced largely consistent phylogenies (Fig. 3B). The new species formed an independent branch in the subfamily Unioninae and was sistered with the clade consisting of Unionini + Aculamprotulini with a relatively well support rate (BS/PP = 70/0.99). Systematics Family Unionidae Rafinesque, 1820 Subfamily Unioninae Rafinesque, 1820 Tribe Globunionini Dai, Chen, Huang & Wu, tribe nov. https://zoobank.org/C3A C7 1B4-5393-4B57-B8C9-5D067E23B3E1 Type genus. G/lobunio Dai, Chen, Huang & Wu, 2024, gen. nov. zse.pensoft.net 786 Diagnosis. Shell small-sized, elongated-globular, inflated, thick, solid; very wide, with some individu- als even wider than height. Anterior extremely short, inflated, and round: posterior long and flat. Anterior margin rounded, dorsal margin straightened, and slope downward at an obtuse angle, usually covered in fine upward wrinkles; ventral margin weakly curved. Umbo inflated, under dorsal margin, almost at the very front of the shell, often eroded. Vernacular name. PREETI (qid bang zu). Remarks. The new tribe has a special small, thick, and elongated-globular shell, which is completely dissimilar to all other tribes of the subfamily. Its independence was also supported by the molecular phylogeny. Genus Globunio Dai, Chen, Huang & Wu, gen. nov. https://zoobank.org/88F4B05A-65A8-49E6-A339-DAE71D4DC071 Type species. Globunio mirificus Chen, Dai, Huang & Wu, 2024, sp. nov. Dai, Y.-T. et al.: A new tribe, genus, and species of freshwater mussel Diagnosis. Same as the tribe. Etymology. The species is made from the Latin glob for globular, and unio for the unionid type genus. Vernacular name. SE) (qit bang shit). Globunio mirificus Chen, Dai, Huang & Wu, sp. nov. https://zoobank.org/C2C368AA-A510-4FDC-B58C-C037009B42CB Figs 4, 6C, D Holotype. 24 NCU_XPWU_GM601, 9, Honggutan Dis- trict [ZL 7 HE X], Nanchang City [Fg STH], Jiangxi Prov- ince [YLPH4], China, 24.68146°N, 109.69794°E, leg. Zhong-Guang Chen & Yu-Ting Dai, September 2022. Paratypes. n = 10, 24 NCU _XPWU_GMO2, Honggutan District [ZL 4X ], Nanchang City [Fa & TH], Jiangxi Province [YLPH4], China, 24.68146°N, 109.69794°E, leg. Zhong-Guang Chen, December 2021; 24 NCU_XPWU_GMO03-11, other information same as holotype. Diagnosis. Same as the tribe. Figure 4. Globunio mirificus gen. et sp. nov. A. Holotype; B-K. Paratypes. zse.pensoft.net Zoosyst. Evol. 101 (2) 2025, 779-790 Description. Shell small-sized, elongated-globular, inflated, thick, solid. Anterior extremely short, inflated, and round: posterior long and flat. Anterior margin rounded, dorsal margin straightened, and slope downward at an obtuse angle, usually covered in fine upward wrinkles; ventral margin weakly curved. Umbo inflated, under dorsal margin, almost at the very front of the shell, often eroded. Central of shell with two rows of posterior dorsal spines (usually detached with only attachment marks), the anterior row longer and the posterior row shorter. Periostracum yellowish-green with olive-green rays and several thick growth lines. Hinge short and strong. Hinge developed. Left valve with two pseudocardinal teeth, anterior tooth small, triangular-shaped, posterior tooth well-developed, rectangular-shaped; right valve with a single well- developed, pyramidal pseudocardinal tooth. Both valves with two lateral teeth: left valve external weak, internal well-developed; right valve external well- developed, internal weak. Mantle attachment scars on the pallial line obvious. Anterior adductor muscle scars deep, samll; posterior adductor muscle scars shallow, orbicular-shaped. Umbo cavity open, deep. Nacre milky white. Measurements. Shell length 15.07—32.37 mm, width 9.83—21.54 mm, height 13.06—19.30 mm. Etymology. The species is named after the Latin mir- ificus for remarkable, referring to the remarkable shell morphology of this species. Vernacular name. 4 PEK (qi yi git bang). Distribution and ecology. Known from three localities of the Changjiang River Basin: the Ganjiang River at Nanchang, the Dongtinghu Lake at Yueyang, and the 112°E 113°E 116°E 112°E 115°E 787 Qingyijiang River at Wuhu (Fig. 5). Living in the slow- flowing rivers and lakes with muddy and sandy substrates alongside dozens of other freshwater mussels (Fig. 6). Discussion Of all freshwater mussels of subfamily Unioninae, only partial species of Aculamprotulini have a somewhat simi- lar elongated-globular shell with the new species. However, the new species has a much smaller (shell length < 35 mm), more expanded, and symmetric shell, while Aculamprotuli- ni has a larger (shell length > 100 mm), flatter, and asymmet- rical shell. A thorough examination of type specimens and more than 100 live animals of the new species has revealed that individuals over 15 mm frequently exhibit severe cor- rosion. This finding serves as a reliable indicator of the ani- mals’ advanced age, thereby confirming their status as adults rather than juveniles of other species. The new species is one of the smallest freshwater mussels in China. The type locality of the new species was located in the central city of Nanchang, a region that has been exten- sively surveyed for freshwater mussels in China. How- ever, due to the general lack of sensitivity amongst re- searchers to freshwater mussel classification, this species was overlooked for a considerable period. Its distribution in densely populated areas poses a significant threat to its survival. The preference of microhabitats with flowing and fine sediment, resulting in a very narrow distribution of the new species in the Ganjiang River. However, all the sediment at its type locality was completely excavated in 2023 for the construction of a water plant (Fig. 6B). Sur- veys conducted between 2023 and 2025, following the 117°E 117°E 118°E 119°E 120°E Figure 5. Distribution of Globunio mirificus gen. et sp. nov. Solid star: type locality; hollow stars: other localities. zse.pensoft.net 788 Dai, Y.-T. et al.: A new tribe, genus, and species of freshwater mussel Figure 6. Habitats and live animals of Globunio mirificus gen. et sp. nov. A. Type locality, Ganjiang River at Honggutan of Nanchang in 2022; B. Type locality, Ganjiang River at Honggutan of Nanchang in 2023; C, D. Live animals. completion of the water plant, failed to recover any live animals. Furthermore, the construction of multiple dams (called the Ganfuweilyu Project) in the lower reaches of the Ganjiang River in 2024, resulting in the storage of water, has led to an increase in the average water level by 10 meters, thereby transforming the river into a static res- ervoir. This has led to concerns regarding the survival of new species that are dependent on flowing microhabitats. 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