High diversity and new species of meiofaunal Nerillidae (Annelida) in subtidal sediments and anchialine caves of the East China Sea
Abstract
Nerillidae is the most species-rich, meiofaunal, annelid family, but little is known about its diversity and distribution in the Northwest Pacific. Four sampling campaigns to Jeju Island, South Korea, and the Ryukyu Islands, Japan, retrieved an unexpectedly high diversity of about 17 nerillid species in subtidal and submarine cave localities of the East China sea, with only Nipponerilla irabuensis Worsaae, Hansen & Fujita in Worsaae et al., 2021 previously described. Bayesian inference and maximum likelihood methods were performed on datasets of concatenated gene sequences (18S rRNA, 28S rRNA, COI, and H3) comprising a total of 55 terminals from 11 different genera of Nerillidae. Integrating detailed morphological assessments (light-, scanning electron-, and confocal laser scanning microscopy) with molecular analyses, we here describe seven new species and a new genus, Cirrinerilla gen. nov. None of the species show geographical overlap between South Korea and the Ryukyu Islands, possibly due to oceanographic and anchialine dispersal barriers and poor dispersal capabilities of Nerillidae. However, two congeneric terminals of Meganerilla and two of Leptonerilla from different islands of the Ryukyus show relatively high molecular similarity, suggesting recent vicariance of these populations (or species). The anchialine caves hold a remarkably high diversity of nerillids compared to coastal subtidal localities, supporting a preadaptation of Nerillidae to these isolated habitats under freshwater influence. This study underlines the need for further exploration of the unique anchialine habitats as well as additional subtidal meiofauna surveys in the East China Sea, for shedding further light on benthic diversifications and colonizations in this region.
References
Atherton S. & Jondelius U. 2020. Biodiversity between sand grains: Meiofauna composition across southern and western Sweden assessed by metabarcoding. Biodiversity Data Journal 8: e51813. https://doi.org/10.3897/BDJ.8.e51813
Boaden P.J.S. 1961. Meganerilla swedmarki nov. gen. nov. spec. an archiannelid of the family Nerillidae. Arkiv för Zoologi 13: 553–559.
Brown S., Rouse G., Hutchings P. & Colgan D. 1999. Assessing the usefulness of histone H3, U2 snRNA and 28S rDNA in analyses of polychaete relationships. Australian Journal of Zoology 47 (5): 499–516. https://doi.org/10.1071/ZO99026
Chan B.K.K., Xu G., Kim H.K., Park J. & Kim W. 2018. Living with marginal coral communities: Diversity and host-specificity in coral-associated barnacles in the northern coral distribution limit of the East China Sea. PloS ONE 13 (5): e0196309. https://doi.org/10.1371/journal.pone.0196309
Chiu R.W.T., Iwatani H., Kitamura A., Yasuhara M. & Fujita K. 2017. Response of subtropical submarine-cave ecosystem to Holocene cave development and Asian monsoon variability. Paleobiology 43: 425–434. https://doi.org/10.1017/pab.2016.53
Chough S.K., Lee H. & Yoon S.H. 2000. Marine Geology of Korean Seas. Elsevier Science B.V.
Cohen B.L., Améziane N., Eleaume M. & de Forges B.R. 2004. Crinoid phylogeny: a preliminary analysis (Echinodermata: Crinoidea). Marine Biology 144: 605–617. https://doi.org/10.1007/s00227-003- 1212-7
Colgan D.J., McLauchlan A., Wilson G.D., Livingston S.P., Edgecombe G.D., Macaranas J., Cassis G. & Gray M.R. 1998. Histone H3 and U2 snRNA DNA sequences and arthropod molecular evolution. Australian Journal of Zoology 46 (5): 419–437. https://doi.org/10.1071/ZO98048
Colgan D.J., Hutchings P.A. & Brown S. 2001. Phylogenetic relationships within the Terebellomorpha. Journal of the Marine Biological Association of United Kingdom 81 (5): 765–773. https://doi.org/10.1017/S002531540100457X
Curini-Galletti M., Artois T., Delogu V., De Smet W.H., Fontaneto D., Jondelius U., Leasi F., Martínez A., Meyer-Wachsmuth I., Nilsson K.S., Tongiorgi P., Worsaae K. & Todaro M.A. 2012. Patterns of diversity in soft-bodied meiofauna: dispersal ability and body size matter. PloS ONE 7 (3): 1–13. https://doi.org/10.1371/journal.pone.0033801
Di Domenico M., Lana P., Martínez A. & Worsaae K. 2014. Molecular and morphological phylogeny of Saccocirridae (Annelida) reveals two cosmopolitan clades with specific habitat preferences. Molecular Phylogenetics and Evolution 75: 202–218. https://doi.org/10.1016/j.ympev.2014.02.003
Eisma D. 2010. South Korea. In: Bird E.C.F. (ed.) Encyclopedia of the World’s Coastal Landforms: 1207–1212. Springer Netherlands, Dordrecht. https://doi.org/10.1007/978-1-4020-8639-7_222
Fujikura K., Kitazato H., Lindsay D., Nishida S. & Shirayama Y. 2010. Marine biodiversity in Japanese waters. PloS ONE 5: e11836. https://doi.org/10.1371/journal.pone.0011836
Fujimoto S. 2015. Quisarctus yasumurai gen. et sp. nov. (Arthrotardigrada: Halechiniscidae) from a submarine cave, off Iejima, Ryukyu Islands, Japan. Zootaxa 3948: 145–150. https://doi.org/10.11646/zootaxa.3948.1.10
Fujimoto S. & Miyazaki K. 2013. Neostygarctus lovedeluxe n. sp. from the Miyako Islands, Japan: The first record of Neostygarctidae (Heterotardigrada: Arthrotardigrada) from the Pacific. Zoological Science 30: 414–419. https://doi.org/10.2108/zsj.30.414
Fujita K., Arakaki T., Denda T., Hidaka M., Hirose E. & Reimer J. 2015. Nature in the Ryukyu Archipelago: Coral Reefs, Biodiversity, and the Natural Environment. International Research Hub Project for Climate Change and Coral Reef/Island Dynamics, Faculty of Science, University of the Ryukyus.
Fujita Y. & Naruse T. 2024. A new genus and species of a submarine cave crab of the family Plagusiidae Dana, 1851 (Crustacea: Brachyura: Grapsoidea) from Okinawa Island, Ryukyu Islands, southwestern Japan. Zootaxa 5410 (3): 408–418. https://doi.org/10.11646/zootaxa.5410.3.8
Gallagher S.J., Iryu Y., Itaki T., Kitamura A., Koizumi I. & Hoiles P.W. 2015. The Pliocene to recent history of the Kuroshio and Tsushima Currents: a multi-proxy approach. Progress in Earth and Planetary Science 2 (1): 17. https://doi.org/10.1186/s40645-015-0045-6
Giere O. 2009. Meiobenthology: The Microscopic Motile Fauna of Aquatic Sediments. Springer-Verlag, Berlin Heidelberg. https://doi.org/10.1007/b106489
Giribet G., Carranza S., Baguña J., Riutort M. & Ribera C. 1996. First molecular evidence for the existence of a Tardigrada + Arthropoda clade. Molecular Biology and Evolution 13 (1): 76–84. https://doi.org/10.1093/oxfordjournals.molbev.a025573
Gonzalez B.C., Martínez A., Worsaae K. & Osborn K.J. 2021. Morphological convergence and adaptation in cave and pelagic scale worms (Polynoidae, Annelida). Scientific Reports 11: 10718. https://doi.org/10.1038/s41598-021-89459-y
Hayami I. & Kase T. 1993. Submarine cave Bivalvia from the Ryukyu Islands: Systematics and evolutionary significance. The University Museum, The University of Tokyo, Bulletin 35: 1–133.
He L., Mukai T., Chu K.H., Ma Q. & Zhang J. 2015. Biogeographical role of the Kuroshio current in the amphibious mudskipper Periophthalmus modestus indicated by mitochondrial DNA data. Scientific Reports 5: 15645. https://doi.org/10.1038/srep15645
Higgins R. & Thiel H. 1988. Introduction to the Study of Meiofauna. Smithsonian Institution Press, Washington DC.
Hillis D.M. & Dixon M.T. 1991. Ribosomal DNA: molecular evolution and phylogenetic inference. The Quarterly Review of Biology 66 (4): 411–453. https://doi.org/10.1086/417338
Hoang D.T., Chernomor O., von Haeseler A., Minh B.Q. & Vinh L.S. 2018. UFBoot2: Improving the ultrafast bootstrap approximation. Molecular Biology and Evolution 35 (2): 518–522. https://doi.org/10.1093/molbev/msx281
Ibrahim A.K., Gamil I.S., Abd-El baky A.A., Hussein M.M. & Tohamy A.A. 2011. Comparative molecular and conventional detection methods of Babesia equi (B. Equi) in Egyptian equine. Global Veterinaria 7 (2): 201–210.
Ise Y., Vacelet J., Mizuyama M. & Fujita Y. 2023. New lithistid sponge of the genus Sollasipelta (Porifera, Demospongiae, Tetractinellida, Neopeltidae) from submarine caves of the Ryukyu Islands, southwestern Japan, with redescription of S. sollasi. Zootaxa 5285 (2): 293–310. https://doi.org/10.11646/zootaxa.5285.2.4
Jimi N. 2024. The polychaetous annelids of Japan: Updated checklist of known species. Species Diversity 29 (2): 337–377. https://doi.org/10.12782/specdiv.29.337
Jouin C. 1968. Sexualité et biologie de la reproduction chez Mesonerilla Remane et Meganerilla Boaden (Archiannélides Nerillidae). Cahiers de Biologie Marine 9: 31–52.
Jouin C. 1971. Status of the knowledge of the systematics and ecology of Archiannelida. Smithsonian Contributions to Zoology 76: 47–56.
Jouin C. & Swedmark B. 1965. Paranerilla limicola n. g., n. sp., Archiannélide Nerillidae du benthos vaseux marin. Cahiers de Biologie Marine 6: 201–218
Kakui K. & Fujita Y. 2018. Haimormus shimojiensis, a new genus and species of Pseudozeuxidae (Crustacea: Tanaidacea) from a submarine limestone cave in Northwestern Pacific. PeerJ 6: e4720. https://doi.org/10.7717/peerj.4720
Kakui K. & Fujita Y. 2023. New sea spider species (Pycnogonida: Austrodecidae) from a submarine cave in Japan. Journal of the Marine Biological Association of the United Kingdom 103: e44. https://doi.org/10.1017/S002531542 10.1017/S002531542
Kang T. & Kim D. 2020. Meiobenthic community structure on the Northeast coastal area of Jeju Island, Korea. Ocean and Polar Research 42 (1): 1–13. https://doi.org/10.4217/OPR.2020.42.1.001
Katoh K. & Standley D.M. 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
Katoh K., Kuma K., Misawa K. & Miyata T. 2002. MAFFT: A novel method for rapid multiple sequence alignment based on fast Fourier transform. Nucleic Acids Research 30 (14): 3059–3066. https://doi.org/10.1093/nar/gkf436
Kim D., Je J. & Lee J. 2000. The community structure and spatial distribution of meiobenthos in the Kanghwa tidal flat, west coast of Korea. Ocean and Polar Research 22: 15–23.
Kimura M. 1980. A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences. Journal of Molecular Evolution 16 (2): 111–120. https://doi.org/10.1007/BF01731581
Koike K. 2010. Japan. In: Bird E.C.F. (ed.) Encyclopedia of the World’s Coastal Landforms: 1213–1224. Springer Netherlands, Dordrecht. https://doi.org/10.1007/978-1-4020-8639-7_223
Kojima S., Iijima A., Kamimura S., Kimura T., Kurozumi T. & Furota T. 2006. Molecular phylogeny and population structure of tideland snails in the genus Cerithidea around Japan. Marine Biology 149 (3): 525–535. https://doi.org/10.1007/s00227-005-0183-2
Komai T. & Fujita Y. 2018. A new genus and new species of alpheid shrimp from a marine cave in the Ryukyu Islands, Japan, with additional record of Salmoneus antricola Komai, Yamada & Yunokawa, 2015 (Crustacea: Decapoda: Caridea). Zootaxa 4369 (4): 575–586. https://doi.org/10.11646/zootaxa.4369.4.7
Kumar S., Knyaz C., Li M., Stecher G. & Tamura K. 2018. MEGA X: Molecular evolutionary genetics analysis across computing platforms. Molecular Biology and Evolution 35 (6): 1547–1549. https://doi.org/10.1093/molbev/msy096
Lee H., Kang T.W., Rho H.S. & Kim D. 2019. Seasonal distribution characteristics of meiobenthos at Gwangyang Bay, Korea. The Sea Journal of the Korean Society of Oceanography 24 (3): 400–421. https://doi.org/10.7850/JKSO.2019.24.3.400
Lévi C. 1953. Archiannélides Nerillidae de la région de Roscoff. Archives de Zoologie expérimentale et générale 90 (2): 64–70.
Liu J.Y. 2013. Status of marine biodiversity of the China seas. PLoS ONE 8 (1): e50719. https://doi.org/10.1371/journal.pone.0050719
Magagnini G. 1966. Un nouvel Archiannélide Nerillidae des côtes de Roscoff: Meganerilla clavata n.sp. Cahiers de Biologie Marine 7: 331–335.
Martínez A., Artois T., Curini-Galletti M., Di Domenico M., Fontaneto D., Gobert S., Jörger K.M., Leasi F., Norenburg J., Núñez J., Todaro M.A., Zotto M.D. & Worsaae K. 2019. Patterns of diversity and endemism of soft-bodied meiofauna in an oceanic island, Lanzarote, Canary Islands. Marine Biodiversity 49 (5): 2033–2055. https://doi.org/10.1007/s12526-019-01007-0
Mendes S.L. da S.D., Rodrigues J.C. & Rizzo A.E. 2024. A new species of Paranerilla Jouin & Swedmark, 1965 (Annelida: Polychaeta) from Brazil. European Journal of Taxonomy 943: 144–153. https://doi.org/10.5852/ejt.2024.943.2591
Meyer C.P. 2003. Molecular systematics of cowries (Gastropoda: Cypraeidae) and diversification patterns in the tropics. Biological Journal of the Linnean Society 79: 401–459. https://doi.org/10.1046/j.1095-8312.2003.00197.x
Miller M., Pfeiffer W.T. & Schwartz T. 2010. Creating the CIPRES Science Gateway for inference of large phylogenetic trees. 2010 Gateway Computing Environments Workshop (GCE), New Orleans: 1–8. https://doi.org/10.1109/GCE.2010.5676129
Minh B.Q., Schmidt H.A., Chernomor O., Schrempf D., Woodhams M.D., von Haeseler A. & Lanfear R. 2020. IQ-TREE2: 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
Motokawa M. & Kajihara H. 2017. Species Diversity of Animals in Japan. Springer Japan. https://doi.org/10.1007/978-4-431-56432-4
Müller M.C. 2002. Aristonerilla: A new nerillid genus (Annelida: Polychaeta) with description of Aristonerilla (Micronerilla) brevis comb. nov. from a seawater aquarium. Cahiers de Biologie Marine 43: 131–139. https://doi.org/10.21411/CBM.A.137553DE
Müller M.C. & Worsaae K. 2006. CLSM analysis of the phalloidin-stained muscle system in Nerilla antennata, Nerillidium sp. and Trochonerilla mobilis (Polychaeta; Nerillidae). Journal of Morphology 267 (8): 885–896. https://doi.org/10.1002/jmor.10292
Müller M.C., Bernhard J.M. & Jouin-Toulmond C. 2001. A new member of Nerillidae (Annelida: Polychaeta), Xenonerilla bactericola gen. et sp. nov., collected off California, USA. Cahiers de Biologie Marine 42 (3): 203–217. https://doi.org/10.21411/CBM.A.64923395
Nakae M., Aizawa M., Fujiwara K., Fukui Y., Hagiwara K., Hata H., Jeong B., Koeda K., Senou H., Shinohara G., Motomura H., Tashiro S., Yamakawa T., Yoshida T. & Matsuura, K. 2018. An annotated checklist of fishes of Amami-oshima Island, the Ryukyu Islands, Japan. Memoirs of the National Museum of Nature and Science 52: 205–361.
Naruse T. & Fujita Y. 2015. Lipkemera iejima, a new cavernicolous crab (Brachyura: Xanthidae) from a submarine cave at Ie Island, central Ryukyu Islands, Japan. Crustacean Research 44: 21–27. https://doi.org/10.18353/crustacea.44.0_21
Núñez J., Ocaña O. & Brito M.C. 1997. Two new species (Polychaeta: Fauveliopsidae and Nerillidae) and other polychaetes from the marine lagoon cave of Jameos del Agua, Lanzarote (Canary Islands). Bulletin of Marine Science 60 (2): 252–260.
Odani S., Uchiyama Y., Kashima M., Kamidaira Y. & Mitarai S. 2017. Influence of the Kuroshio current on dispersal of coral spawn and larvae around Ryukyu Islands. Journal of Japan Society of Civil Engineers 73 (2): 1315–1320. https://doi.org/10.2208/kaigan.73.I_1315
Okanishi M. & Fujita Y. 2018. First finding of anchialine and submarine cave dwelling brittle stars from the Pacific Ocean, with descriptions of new species of Ophiolepis and Ophiozonella (Echinodermata: Ophiuroidea: Amphilepidida). Zootaxa 4377 (1): 1–20. https://doi.org/10.11646/zootaxa.4377.1.1
Osawa M. & Fujita Y. 2019. Submarine cave hermit crabs (Crustacea: Decapoda: Anomura: Paguroidea) from three islands of the Ryukyu Islands, southwestern Japan. Zootaxa 4560 (3): 463–482. https://doi.org/10.11646/zootaxa.4560.3.3
Osawa M. & Fujita Y. 2022. A new species of the genus Eutrichopagurus Komai, 2015 (Crustacea: Decapoda: Anomura: Paguridae) from a submarine cave in the Ryukyu Islands, southwestern Japan. Zootaxa 5214 (3): 440–450. https://doi.org/10.11646/zootaxa.5214.3.6
Park J. 2023. Systematic Studies on the Korean Interstitial Polychaetes (Annelida) Based on Morphological and Molecular Data. PhD Thesis, Ewha Womans University, Seoul, Korea.
Pavlyuk O.N. & Trebukhova Y.A. 2011. Intertidal meiofauna of Jeju Island, Korea. Ocean Science Journal 46 (1): 1–11. https://doi.org/10.1007/s12601-011-0001-3
Posada D. 2008. jModelTest: Phylogenetic model averaging. Molecular Biology and Evolution 25: 1253–1256. https://doi.org/10.1093/molbev/msn083
Rambaut A., Baele G., Drummond A.J., Xie D. & Suchard M.A. 2018. Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Systematic Biology 67 (5): 901–904. https://doi.org/10.1093/sysbio/syy032
Reimer J., Biondi P., Lau Y.W., Masucci G.D., Nguyen X., Santos M. & Wee H.B. 2019. Marine biodiversity research in the Ryukyu Islands, Japan: Current status and trends. PeerJ Preprints 7: e27029v2. https://doi.org/10.7717/peerj.6532
Riser N.W. 1988. Morphology of a new species of nerillid polychaete from the north shore of Massachusetts Bay, U.S.A. Transactions of the American Microscopical Society 107 (2): 171–179. https://doi.org/10.2307/3226459
Ronquist F. & Huelsenbeck J. 2003. MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19: 1572–1574. https://doi.org/10.1093/bioinformatics/btg180
Saito T. & Fujita Y. 2022. A new shrimp of the genus Odontozona Holthuis, 1946 (Decapoda: Stenopodidea: Stenopodidae) from a submarine cave of the Ryukyu Islands, Indo-West Pacific. Zootaxa 5175 (4): 439–452. https://doi.org/10.11646/zootaxa.5175.4.2
Saito Y. & Alino P.M. 2008. Regional Conditions. In: Mimura N. (ed.) Asia-Pacific Coasts and their Management: States of Environment: 255–331. Springer, Dordrecht. https://doi.org/10.1007/978-1-4020-3625-5_5
Saphonov M.V. & Tzetlin A.B. 1988. Nerilla jouini sp. n. (Polychaeta, Nerillidae) – a new archiannelid from the Japan Sea. Zoologicheskii Zhurnal 67 (6): 839–845.
Saphonov M.V. & Tzetlin A.B. 1997. Nerillidae (Annelida: Polychaeta) from the White Sea, with description of a new species of Micronerilla Jouin. Ophelia 47 (3): 215–226. https://doi.org/10.1080/00785236.1997.10428672
Sasaki S. 1981. A new species of the genus Saccocirrus (Archiannelida) from Hokkaido, Northern Japan. Annotationes Zoologicae Japonenses 54 (4): 259–266
Schindelin J., Arganda-Carreras I., Eliceiri K., Frise E., Hartenstein V., Kaynig V., Longair M., Pietzsch T., Preibisch S., Rueden C., Saalfeld S., Schmid B., Tinevez J., Tomancak P., White D.J. & Cardona A. 2012. Fiji: An open-source platform for biological-image analysis. Nature Methods 9 (7): 676–682. https://doi.org/10.1038/nmeth.2019
Schmidt H. & Westheide W. 1998. RAPD-PCR experiments confirm the distinction between three morphologically similar species of Nerilla (Polychaeta: Nerillidae). Zoologischer Anzeiger 236: 277–285.
Shimada D. & Kajihara H. 2014. Two new species of free-living marine nematodes of Adoncholaimus Filipjev, 1918 (Oncholaimida: Oncholaimidae: Adoncholaiminae) from Hokkaido, northern Japan, with a key to species and discussion of the genus. Nematology 16: 437–451. https://doi.org/10.1163/15685411-00002776
Shimada D., Kajihara H. & Mawatari S. 2009. Three new species of free-living marine nematodes (Nematoda: Enoplida) from Northern Japan. Species Diversity 14: 137–150. https://doi.org/10.12782/specdiv.14.137
Shimada D., Kakui K. & Fujita Y. 2023. A new species of free-living marine nematode, Fotolaimus cavus sp. nov. (Nematoda, Oncholaimida, Oncholaimidae), isolated from a submarine anchialine cave in the Ryukyu Islands, southwestern Japan. Zoosystematics and Evolution 99: 519–533. https://doi.org/10.3897/zse.99.109097
Shimomura M. & Fujita Y. 2017. First description of the female of Heteromysoides simplex Hanamura & Kase, 2001 (Mysidacea: Mysidae) collected from a submarine cave on Ie-jima Island, Ryukyu Islands, southwestern Japan. Proceedings of the Biological Society of Washington 130: 75–83. https://doi.org/10.2988/17-00001
Shimomura M. & Fujita Y. 2021. Seborgia cavernicola sp. nov. from a submarine cave on Okinawa Island, Ryukyu Islands, southwestern Japan (Crustacea, Amphipoda, Seborgiidae). Zootaxa 4927 (1): 133–142. https://doi.org/10.11646/zootaxa.4927.1.9
Shimomura M., Fujita Y. & Naruse T. 2012. First record of the genus Thetispelecaris Gutu & Illife, 1998 (Crustacea: Peracarida: Bochusacea) from a submarine cave in the Pacific Ocean. Zootaxa 3367 (1): 69–78. https://doi.org/10.11646/zootaxa.3367.1.7
Sterrer W. & Iliffe T. 1982. Mesonerilla prospera, a new Archiannelid from marine caves in Bermuda. Proceedings of the Biological Society of Washington 95: 509–514. Available from https://www.biodiversitylibrary.org/page/34964823 [accessed 17 Jul. 2025].
Swedmark B. 1959. Archiannélides Nerillidae des côtes du Finistère. Archives de Zoologie expérimentale et générale 98: 26–42.
Takayasu K. 1978. Ryukyu Limestone of Okinawa-Jima, South Japan: A stratigraphical and sedimentological study. Memoirs of the Faculty of Science, Kyoto University. Series of Geology and Mineralogy 45 (1): 133–175.
Tzetlin A.B. & Larionov V.V. 1988. Morphology of a new Archiannelid Akessoniella orientalis gen. et sp. n. (Nerillidae). Zoologicheskii Zhurnal 67: 846–857.
Tzetlin A.B. & Saphonov M.A.V. 1992. Trochonerilla mobilis gen. et sp. n., a meiofaunal nerillid (Annelida, Polychaeta) from a marine aquarium in Moscow. Zoologica Scripta 21 (3): 251–254. https://doi.org/10.1111/j.1463-6409.1992.tb00329.x
Uchida T. 1935. Eine neue Urannelidenart, Polygordius pacificus n. sp. Proceedings of the Imperial Academy 11: 119–120. https://doi.org/10.2183/pjab1912.11.119
Veron J.E.N. 1992. Hermatypic corals of Japan. Australian Institute of Marine Science, Monograph Series 9: 1–234. https://doi.org/10.5962/bhl.title.60634
Vonnemann V., Schrödl M., Klussmann-Kolb A. & Wägele H. 2005. Reconstruction of the phylogeny of the Opisthobranchia (Mollusca: Gastropoda) by means of 18S and 28S rRNA gene sequences. Journal of Molluscan Studies 71 (2): 113–125. https://doi.org/10.1093/mollus/eyi014
Wang P., Li Q. & Li C. 2014. Geology of the China Seas. Developments in Marine Geology 6. Elsevier.
Weese D.A., Fujita Y. & Santos S.R. 2016. Looking for needles in a haystack: Molecular identification of anchialine crustacean larvae (Decapoda: Caridea) from the Shiokawa Spring, Okinawa Island, Ryukyu Islands, Japan. Journal of Crustacean Biology 36 (1): 61–67.
Westheide W. 2008. Polychaetes: Interstitial Families, Keys and Notes for the Identification of the Species. Synopses of the British Fauna (New Series) 44, 2nd Edition. Linnean Society, London.
Westheide W. & Purschke G. 1996. Leptonerilla diplocirrata, a new genus and species of interstitial polychaetes from the island of Hainan, South China (Nerillidae). Proceedings of the Biological Society of Washington 109 (3): 586–590. Available from https://www.biodiversitylibrary.org/page/34645193 [accessed 17 Jul. 2025].
Wilke U. 1953. Mesonerilla intermedia nov. sp., ein neuer Archiannelide aus dem Golf von Neapel. Zoologischer Anzeiger 150: 211–215.
Worsaae K. 2005a. Systematics of Nerillidae (Polychaeta, Annelida). Meiofauna Marina: 49–74.
Worsaae K. 2005b. Phylogeny of Nerillidae (Polychaeta, Annelida) as inferred from combined 18S rDNA and morphological data. Cladistics 21 (2): 143–162. https://doi.org/10.1111/j.1096-0031.2005.00058.x
Worsaae K. 2020. Annelida (excluding Clitellata and Sipuncula). In: Schmidt-Rhaesa A. (ed.) Guide to the Identification of Marine Meiofauna: 239–270. Verlag Dr. Friedrich Pfeil.
Worsaae K. 2021 (2014, Online version). 7.9 Errantia incertae sedis: Nerillidae Levinsen, 1883. In: Purschke G., Böggemann M. & Westheide W. (eds) Handbook of Zoology. Annelida Volume 3: Pleistoannelida, Sedentaria III and Errantia I: 215–227. DeGruyter, Berlin. https://doi.org/10.1515/9783110291704-011
Worsaae K. & Kristensen R.M. 2005. Evolution of interstitial Polychaeta (Annelida). In: Bartolomaeus T. & Purschke G. (eds) Morphology, Molecules, Evolution and Phylogeny in Polychaeta and Related Taxa. Developments in Hydrobiology 179: 319–340. Springer Netherlands, Dordrecht. https://doi.org/10.1007/1-4020-3240-4_18
Worsaae K. & Müller M.C.M. 2004. Nephridial and gonoduct distribution patterns in Nerillidae (Annelida: Polychaeta) examined by tubulin staining and cLSM. Journal of Morphology 261 (3): 259–269. https://doi.org/10.1002/jmor.10153
Worsaae K. & Rouse G. 2009. Mesonerilla neridae n. sp. (Nerillidae): First meiofaunal annelid from deep-sea hydrothermal vents. Zoosymposia 2: 297–303. https://doi.org/10.11646/ZOOSYMPOSIA.2.1.20
Worsaae K., Sterrer W. & Iliffe T. 2004. Longipalpa saltatrix, a new genus and species of the meiofaunal family Nerillidae (Annelida: Polychaeta) from an anchialine cave in Bermuda. Proceedings of the Biological Society of Washington 117 (3): 346–362. Available from https://www.biodiversitylibrary.org/page/35515799 [accessed 17 Jul. 2025].
Worsaae K., Martínez A. & Núñez J. 2009. Nerillidae (Annelida) from the Corona lava tube, Lanzarote, with description of Meganerilla cesari n. sp. Marine Biodiversity 39 (3): 195–207. https://doi.org/10.1007/s12526-009-0027-2
Worsaae K., Mikkelsen M.D. & Martínez A. 2019a. Description of six new species of Mesonerilla (Nerillidae, Annelida) and an emended description of M. intermedia Wilke, 1953, from marine and cave environments. Marine Biodiversity 49 (5): 2141–2165. https://doi.org/10.1007/s12526-019-00984-6
Worsaae K., Armenteros M., Gonzalez B.C., Iliffe T.M., Jørgensen J.T., Kerbl A., Nielsen S.H. & Martínez A. 2019b. Diversity and evolution of the stygobitic Speleonerilla nom. nov. (Nerillidae, Annelida) with description of three new species from anchialine caves in the Caribbean and Lanzarote. Marine Biodiversity 49 (5): 2167–2192. https://doi.org/10.1007/s12526-018-0906-5
Worsaae K., Kerbl A., Di Domenico M., Gonzalez B.C., Bekkouche N. & Martínez A. 2021a. Interstitial Annelida. Diversity 13 (2): 77. https://doi.org/10.3390/d13020077
Worsaae K., Hansen M.J., Axelsen O., Kakui K., Møller P.R., Osborn K.J., Martínez A., Gonzalez B.C., Miyamoto N. & Fujita Y. 2021b. A new cave-dwelling genus and species of Nerillidae (Annelida) from the Ryukyu Islands, Japan. Marine Biodiversity 51: 67. https://doi.org/10.1007/s12526-021-01199-4
Xia X. 2018. DAMBE7: New and improved tools for data analysis in molecular biology and evolution. Molecular Biology and Evolution 35 (6): 1550–1552. https://doi.org/10.1093/molbev/msy073
Yamanishi R. 1983. Preliminary report on the fauna of interstitial polychaetes (Annelida) of Japan. Benthos Research 24: 41–48. https://doi.org/10.5179/benthos1970.1983.41
Copyright (c) 2025 Katrine Worsaae, Malte J. Hansen, Éloïse Defourneaux, Jørgen Olesen, Jiseon Park, Taeseo Park, Yoshihisa Fujita
This work is licensed under a Creative Commons Attribution 4.0 International License.
Creative Commons Copyright Notices
Authors who publish with this journal agree to the following terms:
- Authors retain copyright and grant the journal right of first publication with the work simultaneously licensed under a Creative Commons Attribution License (CC BY 4.0) that allows others to share the work with an acknowledgement of the work's authorship and initial publication in this journal.
- Authors are able to enter into separate, additional contractual arrangements for the non-exclusive distribution of the journal's published version of the work (e.g., post it to an institutional repository or publish it in a book), with an acknowledgement of its initial publication in this journal.
- Authors are NOT PERMITTED to post their submitted work online (e.g., in institutional repositories or on personal websites) prior to publication, as it may lead to nomenclatural issues.
