Phylogeny, taxonomy, and biogeography of the Sphaeriinae (Bivalvia: Sphaeriidae) Author Bespalaya, Yulia V. N. Laverov Federal Center for Integrated Arctic Research of the Ural Branch of the Russian Academy of Sciences, Nikolsky Avenue 20, 163020 Arkhangelsk, Russian Federation & Laboratory of Macroecology & Biogeography of Invertebrates, Saint Petersburg State University, Universitetskaya Emb. 7 - 9, 199034 Saint Petersburg, Russian Federation Author Vinarski, Maxim V. Laboratory of Macroecology & Biogeography of Invertebrates, Saint Petersburg State University, Universitetskaya Emb. 7 - 9, 199034 Saint Petersburg, Russian Federation Author Aksenova, Olga V. N. Laverov Federal Center for Integrated Arctic Research of the Ural Branch of the Russian Academy of Sciences, Nikolsky Avenue 20, 163020 Arkhangelsk, Russian Federation Author Babushkin, Evgeniy S. Laboratory of Macroecology & Biogeography of Invertebrates, Saint Petersburg State University, Universitetskaya Emb. 7 - 9, 199034 Saint Petersburg, Russian Federation & Surgut State University, Lenina Ave., 1, 628403 Surgut, Russian Federation Author Gofarov, Mikhail Yu. N. Laverov Federal Center for Integrated Arctic Research of the Ural Branch of the Russian Academy of Sciences, Nikolsky Avenue 20, 163020 Arkhangelsk, Russian Federation Author Kondakov, Alexander V. N. Laverov Federal Center for Integrated Arctic Research of the Ural Branch of the Russian Academy of Sciences, Nikolsky Avenue 20, 163020 Arkhangelsk, Russian Federation Author Konopleva, Ekaterina S. N. Laverov Federal Center for Integrated Arctic Research of the Ural Branch of the Russian Academy of Sciences, Nikolsky Avenue 20, 163020 Arkhangelsk, Russian Federation Author Kropotin, Alexander V. N. Laverov Federal Center for Integrated Arctic Research of the Ural Branch of the Russian Academy of Sciences, Nikolsky Avenue 20, 163020 Arkhangelsk, Russian Federation Author Mabrouki, Youness Biotechnology, Conservation and Valorisation of Natural Resources Laboratory, Faculty of Sciences Dhar El Mehraz, Sidi Mohamed Ben Abdellah University, Fez. B. P. 1796, 30003 Fès-Atlas, Morocco Author Ovchankova, Nadezda B. Laboratory of Macroecology & Biogeography of Invertebrates, Saint Petersburg State University, Universitetskaya Emb. 7 - 9, 199034 Saint Petersburg, Russian Federation & Zoological Institute, Russian Academy of Sciences, Universitetskaya Emb. 1, 199034 Saint Petersburg, Russian Federation Author Palatov, Dmitry M. Laboratory of Macroecology & Biogeography of Invertebrates, Saint Petersburg State University, Universitetskaya Emb. 7 - 9, 199034 Saint Petersburg, Russian Federation & A. N. Severtsov Institute of Ecology and Evolution of the Russian Academy of Sciences, Leninsky Prt., 33, 119071 Moscow, Russian Federation Author Sokolova, Svetlana E. N. Laverov Federal Center for Integrated Arctic Research of the Ural Branch of the Russian Academy of Sciences, Nikolsky Avenue 20, 163020 Arkhangelsk, Russian Federation Author Shevchenko, Alexander R. N. Laverov Federal Center for Integrated Arctic Research of the Ural Branch of the Russian Academy of Sciences, Nikolsky Avenue 20, 163020 Arkhangelsk, Russian Federation Author Travina, Oksana V. N. Laverov Federal Center for Integrated Arctic Research of the Ural Branch of the Russian Academy of Sciences, Nikolsky Avenue 20, 163020 Arkhangelsk, Russian Federation Author Taybi, Abdelkhaleq F. Mohammed Premier University, Multidisciplinary Faculty of Nador, Applied Biology and Biotechnology Research Team, B. P 300, 62700 Selouane, Morocco Author Soboleva, Alena A. N. Laverov Federal Center for Integrated Arctic Research of the Ural Branch of the Russian Academy of Sciences, Nikolsky Avenue 20, 163020 Arkhangelsk, Russian Federation Author Zubrii, Natalia A. N. Laverov Federal Center for Integrated Arctic Research of the Ural Branch of the Russian Academy of Sciences, Nikolsky Avenue 20, 163020 Arkhangelsk, Russian Federation Author Bolotov, Ivan N. N. Laverov Federal Center for Integrated Arctic Research of the Ural Branch of the Russian Academy of Sciences, Nikolsky Avenue 20, 163020 Arkhangelsk, Russian Federation text Zoological Journal of the Linnean Society 2024 2023-11-04 201 2 305 338 http://dx.doi.org/10.1093/zoolinnean/zlad139 journal article 10.1093/zoolinnean/zlad139 0024-4082 13220074 DC94015-12D0-42CB-B21E-F7C950E94EFDEuglesa Genus Sphaerium Scopoli, 1777 Continued. 1 Table Type species: Tellina cornea Linnaeus, 1758 . Diagnosis: Shell large-sized (shell length in adults varies between 6 and 25 mm ), rounded to ovate in shape, typically thick walled ( Fig. 5E ). The beaks rounded and slightly prominent or narrow, pore density varies among species ( Fig. 5E , 6E ). Cardinal and lateral Figure 2. The phylogeny of the subfamily Sphaeriinae recovered from ML analysis (W-IQ-TREE) and obtained for the complete dataset of mitochondrial and nuclear sequences (16S rRNA+28S rRNA). Black numbers near nodes are bootstrap support values (BS)/ Bayesian posterior probabilities (BPP). The new taxa are highlighted in red. Subgenera are marked by curly brackets. 318 • Bespalaya et al. Figure 3. Maps of approximate geographic distribution of six genera of the Sphaeriinae in the world based on available 16S rRNA sequences: (A) Sphaerium ; (B) Euglesa ; (C) Pisidium s.s. ; (D) Odhneripisiduim ; (E) Hindupisidium ; (F) Conventus . teeth well developed ( Fig. 6E ). Nephridia moderately elongated. Two siphons (exhalant and inhalant) are present ( Fig. 7E ). The outer and inner demibranch relatively well developed. Reproductive biology: The species of the genus are asynchronous brooders (they incubate their progeny in several separate brood sacs). Distribution: Subglobal. This genus is known from Eurasia, Africa ( Sphaerium stuhlmanni ), North and South America ( Sphaerium argentinum ), East Africa, New Zealand ( Sphaerium novaezealandiae ), and Australia ( Sphaerium tasmanicum ) ( Fig. 3A ). Sphaerium nitidum Clessin in Westerlund, 1876 ( Fig. 8 ) Sphaerium nitidum Westerlund, 1876: 97–104 . Type series: The lectotype [pair of valves, designated by Starobogatov in 1968] and eight paralectotypes are deposited in National Museum of Natural History , Stockholm , Sweden Figure 4. Fossil-calibrated phylogeny of the Sphaeriinae calculated under a lognormal relaxed clock model and a Yule process speciation implemented in BEAST 1.10.4 and obtained for the complete dataset of mitochondrial and nuclear sequences (16S rRNA+28S rRNA). Bars indicate 95% HPD of the estimated divergence times between lineages (Mya). Red numbers above the nodes are mean ages (Mya). Black numbers under the nodes are Bayesian posterior probabilities (BPP). Stratigraphic chart according to the International Commission on Stratigraphy (2022) (URL: http://www.stratigraphy.org/ICSchart/ChronostratChart2022-02.pdf). The node pies indicate ancestral area reconstructions of the Sphaeriinae clades (probability of each area combination) in accordance with the generalized biogeographic model (combination of the S-DIVA+DEC+S-DEC models). ( SMNH ) nos 4725–4728 (see Fig. 8A ). The lectotype was collected in Dudinka. Type locality: Russia , Krasnoyarsk Territory , the Yenisei River near Dudinka (‘Jenissei, Dudino … (lat. 69°15 ʹ ) … norr om Lusino … (lat. 68°40 ʹ ); Westerlund, 1877: 67’) . Material examined: Russia : Vaygach Island , Talatinskoe Lake , VIII.2010 , Aksenova , Bespalaya leg., five specimens (lot nos INEP-615, INEP-619); Yamal Peninsula , Seyaha River Basin , VIII.2014 , Zubrii leg., 151 specimens (lot nos RMBH MSph-0011, MSph-0014, MSph-0015); Gydan Peninsula VII.2017 , Sokolova & Shevchenko leg., 23 specimens (lot nos RMBH MSph 372), Yugorsky Peninsula VIII.2015 , Spitsyn leg., two specimens (lot nos RMBH MSph-0087, MSph-0037), Krasnoyarsk Region, Taymyr Peninsula, Dudinka River , VII.2018 , Aksenova, Sokolova, Shevchenko, Travina leg., three specimens (lot no. RMBH MSph-0415), Chukotka Peninsula, Amguema River Basin, VII.2019 , Aksenova leg., four specimens (lot nos RMBH MSph-0494/2, MSph-0497/1) . Figure 5. Representatives of valid genera in the subfamily Sphaeriinae : (A) Euglesa casertana , Russia, Yamalo-Nenets Autonomus Okrug, Gydan Peninsula; (B) Hindupisidium nevillianum (Myanmar, Inle Lake); (C) Conventus conventus , Russia (Arkhangelsk Region, Novaya Zemlya); (D) Pisidium amnicum (Russia, Nenets Autonomus Okrug, Vashutkiny Lakes); (E) Sphaerium corneum (Russia, Arkhangelsk Region, Bolshoy Solovetsky Island); (F) Sphaerium lacustre (Russia, Irkutsk Region, Angara River). Photos: O.V. Aksenova. Scale bar: 1 mm. Description: Shell medium sized (mean length ± SD 3.9 ± 1.1 mm , min–max 1.9–6.5 mm , N = 151), rectangular, with a characteristic concentric sculpture around the beaks ( Fig. 8A–C ). The beaks are rounded and slightly prominent. The cardinal tooth of the right valve is slightly curved, and the weaker split on the posterior end has divergent branches. The inner cardinal tooth of the left valve is thin, curved, and slightly S shaped ( Fig. 8C ). The outer cardinal tooth of the left valve is thin, slightly curved, and shorter than the inner cardinal tooth ( Fig. 8C ). The inner side of the valves is characterized by a high pore density. The nephridia are elongated ( Fig. 8C, D ). Distribution: Kolguev Island, Vaygach Island, Yamal Peninsula, Gydan Peninsula, Yugorsky Peninsula, the Lower Yenisei Basin, Taymyr Peninsula, Chukotka Peninsula. According to some authors, S. nitidum was recorded in North America ( Mackie et al. 1980, Mackie 2007 ). However, their records are not confirmed by genetic data. Comments: The systematic position of S. nitidum has remained controversial to date ( Korniushin 2001 , Bespalaya et al. 2015a ). The specimens studied by us correspond to the morphological and anatomical diagnosis given in the original description of S. nitidum ( Fig. 8A ; Westerlund 1876) and the re-description of this species published by Odhner (1921) ( Fig. 8D ). In addition, we obtained molecular-genetic data on S. nitidum from the Arctic region ( Bespalaya et al. 2015a , 2022 ). The sequenced individuals from Vaygach, Yugorsky Peninsula, Yamal Peninsula, and Gydan Peninsula share the same 16S rRNA haplotype, which is identical to S. nitidum sequences from its type locality (Yenisei River, Dudinka Town, Krasnoyarsk Region ) ( Fig. 2 ), and is separated by three nucleotide substitutions from haplotypes of this species from Chukchi Peninsula and Kolguev Island ( Bespalaya et al . 2022 ).