{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,5,14]],"date-time":"2025-05-14T03:45:15Z","timestamp":1747194315879,"version":"3.40.5"},"reference-count":1,"publisher":"Magnolia Press","issue":"4","content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Zootaxa"],"abstract":"The taxon Koinocystididae is the third most species-rich family within Eukalyptorhynchia. However, its diversity and phylogeny have been largely neglected in former studies. We introduce three new genera and twelve new species of Koinocystididae including Simplexcystis asymmetrica gen. n. sp. n., Galapagetula cubensis sp. n., eight species of Reinhardorhynchus gen. n. and two species of Itaipusa. This raises the total number of species within Koinocystididae from 51 to 63. We also report on new distribution records for six known species: I. divae (Cuba, Panama and New Caledonia), I. karlingi (Sardinia and Lanzarote), Reinhardorhynchus riegeri comb. n. (Cuba), R. ruffinjonesi comb. n. (Cuba and Panama), Utelga heinckei (Cuba and Lanzarote), and U. pseudoheinckei (Sardinia). Simplexcystis asymmetrica gen. n. sp. n. is characterised by a male duct running eccentrically through the copulatory bulb, lack of any hard structures in the male system, lack of a bursa, and the fact that the epithelia of the female, the male, and part of the common atrium are covered by a brush border. Galapagetula cubensis sp. n. has a caudal gonopore, a divisa-type copulatory bulb with an unarmed penis papilla, and a female duct without a sphincter. The new species of Itaipusa and Reinhardorhynchus gen. n. differ from their congeners in the detailed structure of the copulatory bulb and especially the hard structures associated with it. In a molecular phylogenetic analysis based on all available 18S and 28S rDNA sequences of koinocystidids, we found support for the monophyly of the family and the genus Utelga Marcus, 1949. The genus Itaipusa is not monophyletic in that I. sinensis forms a clade with Rhinolasius dillonicus, while other species of Itaipusa that have a copulatory bulb armed with hooks form a clade together with Sekerana stolzi. As the type species of Itaipusa (I. divae) is in neither of these clades, we erected a new genus for I. sinensis (Koinogladius gen. n.) and one for species of Itaipusa having a hook-bearing copulatory bulb (Reinhardorhynchus gen. n.), respectively. Whether the remaining species of Itaipusa form a monophylum remains uncertain.\u00a0<\/jats:p>","DOI":"10.11646\/zootaxa.4948.4.1","type":"journal-article","created":{"date-parts":[[2021,3,22]],"date-time":"2021-03-22T23:44:27Z","timestamp":1616456667000},"source":"Crossref","is-referenced-by-count":4,"title":["Taxonomy and phylogeny of Koinocystididae (Platyhelminthes, Kalyptorhynchia), with the description of three new genera and twelve new species"],"prefix":"10.11646","volume":"4948","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-8741-4799","authenticated-orcid":false,"given":"YANDER L.","family":"DIEZ","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"MARLIES","family":"MONNENS","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7311-2308","authenticated-orcid":false,"given":"ROSA ISABEL","family":"AGUIRRE","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2047-5375","authenticated-orcid":false,"given":"RANA","family":"YURDUSEVEN","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6436-2401","authenticated-orcid":false,"given":"PHILIPPE","family":"JOUK","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2676-7862","authenticated-orcid":false,"given":"NIELS W. L.","family":"VAN STEENKISTE","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0798-0470","authenticated-orcid":false,"given":"BRIAN S.","family":"LEANDER","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0666-5604","authenticated-orcid":false,"given":"ERNEST","family":"SCHOCKAERT","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8721-6290","authenticated-orcid":false,"given":"PATRICK","family":"REYGEL","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9673-8824","authenticated-orcid":false,"given":"KAREN","family":"SMEETS","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2491-7273","authenticated-orcid":false,"given":"TOM","family":"ARTOIS","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"4913","published-online":{"date-parts":[[2021,3,23]]},"reference":[{"key":"50704","unstructured":"

Altschul, S.F., Gish, W., Miller, W., Myers, E.W. & Lipman, D.J. (1990) Basic local alignment search tool. Journal of Molecular Biology<\/em>, 215, 403\u2013410.<\/p>

https:\/\/doi.org\/10.1016\/S0022-2836(05)80360-2<\/a><\/p>

Attems, C.G. (1897) Beitrag zur Kenntnis der rhabdocoelen Turbellarien Helgolands. Wissenschaftliche Meeresuntersuchungen der Kommission Abteilung Kiel<\/em>, 2 (1), 219\u2013232.<\/p>

Benson, D.A., Karsch-Mizrachi, I., Clark, K., Lipman, D.J., Ostell, J. & Sayers, E.W. (2012) GenBank. Nucleic Acids Research<\/em>, 40, D48\u2013D53.<\/p>

https:\/\/doi.org\/10.1093\/nar\/gkr1202<\/a><\/p>

Brunet, M. (1965) Turbellari\u00e9s calyptorhynques de substrats meubles de la r\u00e9gion de Marseille. Recueil des Travaux de la Station Marine d\u2019Endoume<\/em>, 39 (55), 128\u2013218.<\/p>

Brunet, M. (1972) Koinocystididae de la r\u00e9gion de Marseille. Zoologica Scripta<\/em>, 1, 157\u2013174.<\/p>

https:\/\/doi.org\/10.1111\/j.1463-6409.1972.tb00673.x<\/a><\/p>

Castresana, J. (2000) Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Molecular Biology and Evolution<\/em>, 17, 540\u2013552.<\/p>

https:\/\/doi.org\/10.1093\/oxfordjournals.molbev.a026334<\/a><\/p>

Claus, C. (1887) Lehrbuch der Zoologie<\/em>. 4<\/em>. Auflage<\/em>. N. G. Elwertsche Verlagsbuchhandlung, Marburg, Leipzig, 1279 pp.<\/p>

Ehlers, U. (1985) Cytoskelette bei freilebenden Plathelminthen. Verhandlungen der Deutschen Zoologischen Gesellschaft<\/em>, 78, 161.<\/p>

Ehrenberg, C.G. (1831) Animalia evertebrata exclusis insectis recensuit C.G. Ehrenberg. Series prima cum Tabularum decade prima. In<\/em>: Hemprich, F.G. & Ehrenberg, C.G. (Eds.), Symbolae Physicae 2. Phytozoa Turbellaria<\/em>. Officina Academica, Berolina, pp. 1\u201315.<\/p>

Fuhrmann, O. (1904) Ein neuer Vertreter eines marinen Turbellariengenus in S\u00fcsswasser. Zoologischer Anzeiger<\/em>, 27, 381\u2013384.<\/p>

Graff, L. von (1905) Marine turbellarien Orottavas und der K\u00fcsten Europas. Zeitschrift f\u00fcr wissenschaftliche Zoologie<\/em>, 83, 68\u2013148.<\/p>

Guindon, S., Dufayard, J.-F., Lefort, V., Anisimova, M., Hordijk, W. & Gascuel, O. (2010) New Algorithms and Methods to Estimate Maximum-Likelihood Phylogenies: Assessing the Performance of PhyML 3.0. Systematic Biology<\/em>, 59 (3), 307\u2013321.<\/p>

https:\/\/doi.org\/10.1093\/sysbio\/syq010<\/a><\/p>

Hoang, D.T., Chernomor, O., Haeseler, Arndt von, Minh, B.Q. & Vinh, L.S. (2013) UFBoot2: Improving the Ultrafast Bootstrap Approximation. Molecular Biology and Evolution<\/em>, 35 (2), 518\u2013522.<\/p>

https:\/\/doi.org\/10.1093\/molbev\/msx281<\/a><\/p>

Kalyaanamoorthy, S., Minh, B.Q., Wong, T.K.F., Haeseler, A. von & Jermiin, L.S. (2017) ModelFinder: Fast Model Selection for Accurate Phylogenetic Estimates. Nature Methods<\/em>, 14 (6), 587\u2013589.<\/p>

https:\/\/doi.org\/10.1038\/nmeth.4285<\/a><\/p>

Karling, T.G. (1952a) Kalyptorhynchia (Turbellaria). Further Zoological Results of the Swedish Antarctic Expedition<\/em>, 4 (9), 1\u201350.<\/p>

Karling, T.G. (1952b) Studien \u00fcber Kalyptorhynchien (Turbellaria) IV. Einige Eukalyptorhynchia. Acta Zoologica Fennica<\/em>, 69, 1\u201349.<\/p>

Karling, T.G. (1954) Einige marine Vertreter der Kalyptorhynchien-Familie Koinocystididae. Arkiv f\u00f6r Zoologi<\/em>, 7, 165\u2212183.<\/p>

Karling, T.G. (1956) Morphologisch-histologische Untersuchungen an den m\u00e4nnlichen Atrialorganen der Kalyptorhynchia (Turbellaria). Arkiv f\u00f6r Zoologi<\/em>, Serie 2, <\/em>9 (7), 187\u2013279.<\/p>

Karling, T.G. (1964) \u00dcber einige neue und ungen\u00fcgend bekannte Turbellaria Eukalyptorhynchia. Zoologischer Anzeiger<\/em>, 172, 159\u2212183.<\/p>

Karling, T.G. (1978) Anatomy and Systematics of Marine Turbellaria from Bermuda. Zoologica Scripta<\/em>, 7, 25\u2013248.<\/p>

Karling, T.G. (1980) Revision of Koinocystididae (Turbellaria). Zoologica Scripta<\/em>, 9, 241\u2013269.<\/p>

https:\/\/doi.org\/10.1111\/j.1463-6409.1980.tb00666.x<\/a><\/p>

Karling, T.G., Mack-Fira, V. & D\u00f6rjes, J. (1972) First report on marine microturbellarians from Hawaii. Zoologica Scripta<\/em>, <\/em>1, 251\u2013269.<\/p>

https:\/\/doi.org\/10.1111\/j.1463-6409.1972.tb00575.x<\/a><\/p>

Katoh, K. & Standley, D.M. (2013) MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology and Evolution<\/em>, 30 (4), 772\u2013780.<\/p>

https:\/\/doi.org\/10.1093\/molbev\/mst010<\/a><\/p>

Katoh, K., Rozewicki, J., Yamada, K.D. (2017) MAFFT online service: multiple sequence alignment, interactive sequence choice and visualization. Briefings in Bioinformatics<\/em>, 20 (4), 1160\u20131166.<\/p>

https:\/\/doi.org\/10.1093\/bib\/bbx108<\/a><\/p>

Kearse, M., Moir, R., Wilson, A., Stones-Havas, S., Cheung, M., Sturrock, S., Buxton, S., Cooper, A., Markowitz, S., Duran, C., Thierer, T., Ashton, B., Meintjes, P. & Drummond, A.A. (2012) Geneious Basic: An integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics<\/em>, 28 (12), 1647\u2013164.<\/p>

https:\/\/doi.org\/10.1093\/bioinformatics\/bts199<\/a><\/p>

Lanfear, R., Calcott, B., Ho, S.Y.W. & Guindon, S. (2012) Partition Finder: Combined selection of partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology and Evolution<\/em>, 29, 1695\u20131701.<\/p>

https:\/\/doi.org\/10.1093\/molbev\/mss020<\/a><\/p>

Lin, Y.-T., Feng, W.-T., Zhuang, J.-Y., Zhang, Y. & Wang, A.-T. (2017) Two new species of Kalyptorhynchia (Koinocystididae and Gnathorhynchidae) from China. Zootaxa<\/em>, 4337 (4), 573-583.<\/p>

https:\/\/doi.org\/10.11646\/zootaxa.4337.4.8<\/a><\/p>

Mack-Fira, V. (1968) Turbellari\u00e9s de la Mer Noire. Rapport de la Commission pour la Mer M\u00e9diterran\u00e9e<\/em>, 9 (2), 179\u2013182.<\/p>

Mack-Fira, V. (1974) The turbellarian fauna of the Romanian littoral waters of the Black Sea and its annexes. In<\/em>: Riser, N.W. & Morse, M.P. (Eds.), Biology of Turbellaria<\/em>. McGraw-Hill, New York. pp. 248\u2013290.<\/p>

Marcus, E. (1949) Turbellaria Brasileiros (7). Boletins da Faculdade de Filosofia Ciencias e Letras, Universidade de S\u00e3o Paulo Zoologia<\/em>, 7 (14), 7\u2212155.<\/p>

https:\/\/doi.org\/10.11606\/issn.2526-4877.bsffclzoologia.1949.129106<\/a><\/p>

Marcus, E. (1951) Turbellaria Brasileiros (9). Boletins da Faculdade de Filosofia, Ciencias e Letras, Universidade de S\u00e3o Paulo Zoologia<\/em>, 16, 1\u2212215.<\/p>

https:\/\/doi.org\/10.11606\/issn.2526-4877.bsffclzoologia.1951.125221<\/a><\/p>

Marcus, E. (1954) Turbellaria Brasileiros (11). Pap\u00e9is Avulsos<\/em>, 11, 419\u2212489.<\/p>

https:\/\/doi.org\/10.11606\/issn.2526-4877.bsffclzoologia.1946.125301<\/a><\/p>

Meixner, J. (1924) Studien zu einer Monographie der Kalyptorhynchia und zum System der Turbellaria <\/strong>Rhabdocoela. Zoologischer Anzeiger<\/em>, 60, 89\u2212105.<\/strong><\/p>

Meixner, J. (1925) Beitrag zur Morphologie und zum System der Turbellaria-Rhabdocoela. I. Die Kalyptorhynchia. Zeitschrift f\u00fcr Morphologie und \u00d6kologie der Tiere<\/em>, 3, 255\u2013343.<\/p>

https:\/\/doi.org\/10.1007\/BF00408146<\/a><\/strong><\/p>

Meixner, J. (1928) Der Genitalapparat der Tricladen und seine Beziehungen zu ihrer allgemeinen Morphologie, Phylogenie, \u00d6kologie und Verbreitung. Zeitschrift f\u00fcr Morphologie und \u00d6kologie der Tiere<\/em>, 11 <\/em>(5), 570\u2013612.<\/p>

https:\/\/doi.org\/10.1007\/BF02424587<\/a><\/p>

Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. Proceedings of the Gateway Computing Environments Workshop<\/em>, 2010, 1\u20138.<\/p>

https:\/\/doi.org\/10.1109\/GCE.2010.5676129<\/a><\/p>

Nguyen, L.-T., Schmidt, H.A., Haeseler, A. von & Minh, B.Q. (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution<\/em>, 32 (1), 268\u2013274.<\/p>

https:\/\/doi.org\/10.1093\/molbev\/msu300<\/a><\/p>

Rambaut, A. (2006\u20132019) FigTree: Tree Figure Drawing Tool. Computer program distributed by the author. Available from: http:\/\/tree.bio.ed.ac.uk\/software\/figtree\/<\/a> (accessed 14 August 2020)<\/p>

Reygel, P.C., Willems, W.R. & Artois, T.J. (2011) Koinocystididae and Gnathorhynchidae (Platyhelminthes: Rhabdocoela: Kalyptorhynchia) from the Galapagos, with the description of three new species. Zootaxa<\/em>, 3096 (1), 27\u201340.<\/p>

https:\/\/doi.org\/10.11646\/zootaxa.3096.1.3<\/a><\/p>

Ronquist, F., Teslenko, M., Mark, P. van der, Ayres, D.L., Darling, A., H\u00f6hna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2 efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology<\/em>, 61 (3), 539\u2013542.<\/p>

https:\/\/doi.org\/10.1093\/sysbio\/sys029<\/a><\/p>

Schockaert, E.R. (1996) Turbellarians. In<\/em>: Hall, G.S. (Ed.), Methods for the Examination of Organismal Diversity in Soils and Sediments<\/em>. CAB International Wallingford, pp. 211\u2013225.<\/p>

Sekera, E. (1912) Monographie der Gruppe Olisthanellini. Sitzungsberichte der K\u00f6niglichen B\u00f6hmischen Gesellschaft der Wissenschaften. Classe f\u00fcr Mathematik und Naturwissenschaften<\/em>, 24, 1\u201393.<\/p>

Strand, E. (1914) Neue Namen verschiedener Tiere. Archiv f\u00fcr Naturgeschichte<\/em>, <\/em>80, 163\u2013164. <\/p>

Tessens, B., Janssen, T. & Artois, T. (2014) Molecular phylogeny of Kalyptorhynchia (Rhabdocoela, Platyhelminthes) inferred from ribosomal sequence data. Zoologica Scripta<\/em>, 43 (5), 519\u2013530.<\/p>

https:\/\/doi.org\/10.1111\/zsc.12066<\/a><\/p>

Trifinopoulos, J., Nguyen, L.-T., Haeseler, Arndt von & Minh, B.Q. (2017) W-IQ-TREE: a fast online phylogenetic tool for maximum likelihood analysis. Nucleic Acids Research<\/em>, 44, W232\u2013W235.<\/p>

https:\/\/doi.org\/10.1093\/nar\/gkw256<\/a><\/p>

Tyler, S., Artois, T., Schilling, S., Hooge, M. & Bush, L.F. (Eds.) (2006\u20132019) World List of turbellarian worms: Acoelomorpha, Catenulida, Rhabditophora. Koinocystididae Meixner, 1924. World Register of Marine Species. Available from: http:\/\/www.marinespecies.org\/aphia.php?p=taxdetails&id=142395<\/a> (accessed 28 November 2018)<\/p>

Van Steenkiste, N., Tessens, B., Willems, W., Backeljau, T., Jondelius, U. & Artois, T. (2013) Comprehensive Molecular Phylogeny of Dalytyphloplanida (Platyhelminthes: Rhabdocoela) Reveals Multiple Escapes from the Marine Environment and Origins of Symbiotic Relationships. PLoS ONE<\/em>, 8 (3), e59917.<\/p>

https:\/\/doi.org\/10.1371\/journal.pone.0059917<\/a><\/p>

Van Steenkiste, N.W.L. & Leander, B.S. (2018) Species diversity of eukalyptorhynch flatworms (Platyhelminthes, Rhabdocoela) from the coastal margin of British Columbia: Polycystididae, Koinocystididae and Gnathorhynchidae. Marine Biology Research<\/em>, 14 (9\u201310), 899\u2013923.<\/p>

https:\/\/doi.org\/10.1080\/17451000.2019.1575514<\/a><\/p>

Van Steenkiste, N.W.L., Herbert, E.R. & Leander, B.S. (2018) Species diversity in the marine microturbellarian Astrotorhynchus bifidus<\/em> sensu lato (Platyhelminthes: Rhabdocoela) from the Northeast Pacific Ocean. Molecular Phylogenetics and Evolution<\/em>, 120, 259\u2013273.<\/p>

https:\/\/doi.org\/10.1016\/j.ympev.2017.12.012<\/a><\/p>

Willems, W., Reygel, P., Van Steenkiste, N., Tessens, B. & Artois, T. (2017) Kalyptorhynchia (Platyhelminthes: Rhabdocoela) from KwaZulu-Natal (South Africa), with the description of six new species. Zootaxa<\/em>, 4242 (3), 441\u2013466.<\/p>

https:\/\/doi.org\/10.11646\/zootaxa.4242.3.2<\/a><\/p>

Willems, W.R., Sandberg, M.I. & Jondelius, U. (2007) First report on Rhabdocoela (Rhabditophora) from deep parts of Skagerrak, with the description of four new species. Zootaxa<\/em>, 1616 (1), 1\u201321.<\/p>

https:\/\/doi.org\/10.11646\/zootaxa.1616.1.1<\/a><\/p>

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