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We examined the muscle arrangement in the penial complex (penis and penis sheath) of most major groups of freshwater pulmonates to explore to which extent the copulatory musculature can serve as a source of phylogenetic information for Hygrophila. The penises of <jats:italic>Acroloxus lacustris<\/jats:italic> (Acroloxidae), <jats:styled-content style=\"fixed-case\"><jats:italic>Radix auricularia<\/jats:italic><\/jats:styled-content> (Lymnaeidae), and <jats:styled-content style=\"fixed-case\"><jats:italic>Physella acuta<\/jats:italic><\/jats:styled-content> (Physidae) posses inner and outer layers of circular muscles and an intermediate layer of longitudinal muscles. The inner and outer muscle layers in the penis of <jats:styled-content style=\"fixed-case\"><jats:italic>Biomphalaria glabrata<\/jats:italic><\/jats:styled-content> consist of circular muscles, but this species has two intermediate longitudinal layers separated by a lacunar space, which is crossed by radial and transverse fibers. The muscular wall of the penis of <jats:styled-content style=\"fixed-case\"><jats:italic>Planorbella duryi<\/jats:italic><\/jats:styled-content> is composed of transverse and longitudinal fibers, with circular muscles as the outer layer. In Planorbidae, the penial musculature consists of inner and outer layers of longitudinal muscles and an intermediate layer of radial muscles. The penis sheath shows more variation in muscle patterns: its muscular wall has two layers in <jats:styled-content style=\"fixed-case\"><jats:italic>A. lacustris<\/jats:italic><\/jats:styled-content>, <jats:styled-content style=\"fixed-case\"><jats:italic>P. acuta<\/jats:italic><\/jats:styled-content>, and <jats:styled-content style=\"fixed-case\"><jats:italic>P. duryi<\/jats:italic><\/jats:styled-content>, three layers in <jats:styled-content style=\"fixed-case\"><jats:italic>R. auricularia<\/jats:italic><\/jats:styled-content> and Planorbinae and four layers in <jats:styled-content style=\"fixed-case\"><jats:italic>B. glabrata<\/jats:italic><\/jats:styled-content>. To trace the evolution of the penial musculature, we mapped the muscle characters on a molecular phylogeny constructed from the concatenated 18S and mtCOI data set. The most convincing synapomorphies were found for Planorbinae (inner and outer penis layers of longitudinal muscles, three\u2010layered wall of the penis sheath). A larger clade coinciding with Planorbidae is defined by the presence of radial muscles and two longitudinal layers in the penis. The comparative analysis of the penial musculature appears to be a promising tool in unraveling the phylogeny of Hygrophila.<\/jats:p>","DOI":"10.1002\/jmor.20960","type":"journal-article","created":{"date-parts":[[2019,2,14]],"date-time":"2019-02-14T10:30:20Z","timestamp":1550140220000},"page":"508-525","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":11,"title":["Musculature of the penial complex: A new criterion in unravelling the phylogeny of Hygrophila (Gastropoda: Pulmonata)"],"prefix":"10.1002","volume":"280","author":[{"given":"Elena V.","family":"Soldatenko","sequence":"first","affiliation":[{"name":"Zoological Institute of Russian Academy of Sciences, Laboratory of Marine Research Smolensk State University  Smolensk Russia"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3010-8341","authenticated-orcid":false,"given":"Anatoly A.","family":"Petrov","sequence":"additional","affiliation":[{"name":"Zoological Institute of Russian Academy of Sciences Laboratory of Evolutionary Morphology, Universitetskaya nab. 1  Saint\u2010Petersburg Russia"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"311","published-online":{"date-parts":[[2019,2,14]]},"reference":[{"key":"e_1_2_6_2_1","doi-asserted-by":"publisher","DOI":"10.2307\/3223749"},{"key":"e_1_2_6_3_1","first-page":"318","article-title":"Descriptions of thirteen new species of shells of New England","volume":"3","author":"Adams C. 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