The family Xystodesmidae in the Korean Peninsula (Diplopoda: Polydesmida)

Keywords: Levizonus, North and South Korea, Parafontaria, Riukiaria, Xystodesmus

Abstract

The polydesmidan family Xystodesmidae is reviewed with respect to the millipede fauna of the Korean Peninsula. It encompasses eight species in four genera: Xystodesmus Cook, 1895; Riukiaria Attems, 1898; Levizonus Attems, 1898; and Parafontaria Verhoeff, 1936. We redescribe five species of Xystodesmus including Xystodesmus pallidus (Verhoeff, 1937) and Xystodesmus amoenus (Takakuwa, 1942). Xystodesmus shirozui (Takakuwa, 1942) is new to the fauna of Korea. The enigmatic Pachydesmus bazanensis Takakuwa, 1942 is transferred to Xystodesmus as X. bazanensis comb. nov. A new species is described, X. bifurcus sp. nov. Of the other three genera, only one species of each is known from Korea, and they are as follows: Riukiaria koreana sp. nov. (from R. semicircularis (Takakuwa, 1941)), Levizonus circularis (Takakuwa, 1942) and Parafontaria koreana (Paik, 1963). Their validity and taxonomic position, based partly on new material, is discussed. Our maximum likelihood phylogeny, estimated based on concatenated sequences of mitochondrial COI and nuclear EF1-α, supported the monophyly of these four genera. Furthermore, the presence of subclades within these clades suggests potential genetic differentiation at the population level.

References

Altschul S.F., Gish W., Miller W., Myers E.W. & Lipman D.J. 1990. Basic local alignment search tool. Journal of Molecular Biology 215 (3): 403–410. https://doi.org/10.1016/S0022-2836(05)80360-2

Attems C. 1898. System der Polydesmiden, I. Denkschriften der Kaiserlichen Akademie der Wissenschaften Mathematisch-Naturwissenschaftliche Classe 67: 221–482.

Attems C. 1931. Die Familie Leptodesmidae und andere Polydesmiden. Zoologica 30 (3/4): 1–149.

Attems C. 1938. Polydesmoidea II. Fam. Leptodesmidae, Platyrhachidae, Oxydesmidae, Gomphodesmidae. Das Tierreich 69: 1–487. Walter de Gruyter & Co., Berlin & Leipzig.

Bańkowska R. & Sterzyńska M. 1997. Faunistic investigations conducted in North Korea by researchers from the Institute of Zoology Polish Academy of Sciences from 1959 to 1990. Fragmenta faunistica 40 (20): 247–253. https://doi.org/10.3161/00159301FF1997.40.20.247

Castresana J. 2000. Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Molecular Biology and Evolution 17 (4): 540–552. https://doi.org/10.1093/oxfordjournals.molbev.a026334

Chamberlin R.V. 1943. On some genera and species of American millipeds. Bulletin of the University of Utah 34: 3–20.

Chamberlin R.V. & Wang Y.M. 1953. Records of millipeds (Diplopoda) from Japan and other oriental areas, with descriptions of new genera and species. American Museum Novitates 1621: 1–13.

Chao J-L. & Chang H-W. 2008. Neotype designation for two centipedes, Scolopocryptops curtus (Takakuwa, 1939) and Cryptops nigropictus Takakuwa, 1936, and a review of species of Scolopendromorpha (Chilopoda) in Taiwan. Collection and Research 21: 1–15. Available from https://libknowledge.nmns.edu.tw/nmns/upload/bulletin/000000177/209000c/200812-1.pdf [accessed 9 Jan. 2026].

Darriba D., Taboada G.L., Doallo R. & Posada D. 2012. jModelTest 2: more models, new heuristics and parallel computing. Nature Methods 9: 772. https://doi.org/10.1038/nmeth.2109

Folmer O., Black M., Hoeh W., Lutz R. & Vrijenhoek R. 1994. DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology 3: 294–299.

Golovatch S.I. 2014. The millipede genus Riukiaria Attems, 1938 in continental China, with descriptions of new species (Diplopoda: Polydesmida: Xystodesmidae). Zootaxa 3793 (1): 188–200. https://doi.org/10.11646/zootaxa.3793.1.9

Golovatch S.I. 2023. On the synonymy of the millipede genera Riukiaria Attems, 1938 and Parariukiaria Nguyen, 2016, with records of Riukiaria cf. davidiani Golovatch, 2014 from Sichuan, southern China (Diplopoda: Polydesmida: Xystodesmidae). Arthropoda Selecta 32 (3): 231–238. https://doi.org/10.15298/arthsel.32.3.01

Hall T.A. 1999. BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series 41: 95–98.

Hoffman R.L. 1956. Studies on some oriental xystodesmine millipeds (Polydesmida, Chelodesmidae). Proceedings of the Entomological Society of Washington 58 (2): 95–104.

Hoffman R.L. 1978a. On the classification and phylogeny of chelodesmoid Diplopoda. Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg 21/22: 129–134.

Hoffman R.L. 1978b. The taxonomic and nomenclatorial status of the milliped generic names Parafontaria Verhoeff, Cyphonaria Verhoeff, and Japonaria Attems (Polydesmida, Xystodesmidae). Spixiana 1: 215–224.

Hoffman R.L. 1980. Classification of the Diplopoda. Muséum d’Historie Naturelle, Genève.

Jeekel C.A.W. 1971. Nomenclator generum et familiarum Diplopodorum: A list of the genus and family-group names in the Class Diplopoda from the 10th edition of Linnaeus, 1758, to the end of 1957. Monografieën van de Nederlandse Entomologische Vereniging 5: 1–412.

Kim G., Adebimpe A.A., Nguyen A.D., Korsós Z. & Hwang U.W. 2023. An annotated checklist of millipedes (Myriapoda: Diplopoda) inhabiting the Korean Peninsula. Animal Systematics, Evolution and Diversity 39 (4): 133–154. https://doi.org/10.5635/ASED.2023.39.4.019

Kimura M. 1980. A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences. Journal of Molecular Evolution 16: 111–120. https://doi.org/10.1007/BF01731581

Korsós Z. & Nakamura Y. 2025. Review of the millipede genus Xystodesmus (Diplopoda: Polydesmida), with seven new species from the southwestern part of Japan. ZooKeys 1231: 191–232. https://doi.org/10.3897/zookeys.1231.141443

Korsós Z., Nakamura Y. & Tanabe T. 2011. Two new millipede species of the genus Riukiaria (Diplopoda, Polydesmida, Xystodesmidae) endemic to the Ryukyu Archipelago, Japan. Zootaxa 2877: 55–68. https://doi.org/10.11646/zootaxa.2877.1.3

Larkin M.A., Blackshields G., Brown N.P., Chenna R., McGettigan P.A. & McWilliam H. 2007. Clustal W and Clustal X version 2.0. Bioinformatics 23: 2947–2948. https://doi.org/10.1093/bioinformatics/btm404

Lim K.-Y. 2001. Taxonomy of Millipedes (Arthropoda: Diplopoda) in Korea. PhD thesis, Chonbuk National University, Jeonju.

Lokshina I.E. & Golovatch S.I. 1977. The new myriapods (Diplopoda) from the USSR. Byulleten Moskovskogo Obshchestva Ispytateley Prirody, Otdel Biol. 82 (1): 73–78. [In Russian.]

Marek P., Tanabe T. & Sierwald P. 2014. A species catalog of the millipede family Xystodesmidae (Diplopoda: Polydesmida). Virginia Museum of Natural History Special Publication 17: 1–117. Available from https://www.vmnh.net/content/vmnh/uploads/PDFs/research_and_collections/special_publications/vmnhspecialpub17.pdf [accessed 9 Jan. 2026].

Means J.C., Hennen D.A., Tanabe T. & Marek P.E. 2021. Phylogenetic systematics of the millipede family Xystodesmidae. Insect Systematics and Diversity 5 (2): 1–26. https://doi.org/10.1093/isd/ixab003

Mikhaljova E.V. 2021. Review of the millipede genus Levizonus Attems, 1898, with description of a new species from the Far East of Russia (Diplopoda, Polydesmida, Xystodesmidae). European Journal of Taxonomy 751: 159–184. https://doi.org/10.5852/ejt.2021.751.1387

Mikhaljova E.V., Golovatch S. I. & Wytwer J. 2000. On some new or poorly-known millipedes (Diplopoda) from North Korea. Fragmenta Faunistica 43 (10): 109–122. https://doi.org/10.3161/00159301FF2000.43.10.109

Miyosi Y. 1952. Beiträge zur Kenntniss japanischer Myriopoden. 4. Aufsatz: Ueber eine neue Art und eine neue Unterart von Diplopoda. Zoological Magazine 61 (9): 281–282.

Miyosi Y. 1959. Über japanische Diplopoden. Arachnological Society of East Asia, Spec. Number. Osaka. [In Japanese with German summary.]

National Library of Medicine 1988. National Center for Biotechnology Information (NCBI ). Bethesda, MD: National Library of Medicine (US). Available from https://www.ncbi.nlm.nih.gov/ [accessed 12 Oct. 2024].

Nguyen A.D. 2016. Discovery of a new millipede species in northern Vietnam, and the proposal of a new genus, Parariukiaria (Diplopoda, Polydesmida, Xystodesmidae). Zootaxa 4121 (3): 331–336. https://doi.org/10.11646/zootaxa.4121.3.7

Nguyen A.D., Jang K.H., Hyun J.S. & Hwang U.W. 2016. An updated checklist and perspective study of millipedes (Arthropoda: Myriapoda: Diplopoda) in the Korean Peninsula. Animal Systematics, Evolution and Diversity 32: 44–48. https://doi.org/10.5635/ASED.2016.32.1.044

Paik K.-Y. 1963. Survey of the myriapods of Mt. Sokkri, Chungcheung-Pookto, Korea. Kyungpook University Theses Collection 7: 33–42.

Park J.O., Kimura M. & Taira A. 1996. Late Pleistocene unconformity of the Tsushima and Korea Straits revealed by seismic reflection profiles. Journal of Geography (Chigaku Zasshi) 105 (3): 297–305. https://doi.org/10.5026/jgeography.105.3_297

Rambaut A. 2012. FigTree ver. 1.4. Available from https://github.com/rambaut/figtree [accessed 9 Jan. 2026].

Reip H.S. & Wesener T. 2018. Intraspecific variation and phylogeography of the millipede model organism, the Black Pill Millipede Glomeris marginata (Villers, 1789) (Diplopoda, Glomeridae, Glomeridae). ZooKeys 741: 93–131. https://doi.org/10.3897/zookeys.741.21917

Ronquist F., Teslenko M., Van Der Mark P., Ayres D.L., Darling A., Höhna 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 61: e539542. https://doi.org/10.1093/sysbio/sys029

Schwarz G. 1978. Estimating the dimension of a model. The Annals of Statistics 6 (2): 461–464. https://doi.org/10.1214/aos/1176344136

Shinohara K. 1977. Revaluation on Riukiaria (Diplopoda). Acta Arachnologica 27: 115–119. [In Japanese with English synopsis.] https://doi.org/10.2476/asjaa.27.Specialnumber_115

Sierwald P., Decker P. & Spelda J. 2025. MilliBase. https://doi.org/10.14284/370

Sota T. & Tanabe T. 2010. Multiple speciation events in an arthropod with divergent evolution in sexual morphology. Proceedings of the Royal Society of London B: Biological Sciences 277: 689–696. https://doi.org/10.1098/rspb.2009.1822

Takakuwa Y. 1941a. Weitere japanische Lithobius-Arten und zwei neue Diplopoden. Transactions of the Sapporo Natural History Society 17 (1): 1–9.

Takakuwa Y. 1941b. Rhysodesmus-Arten aus Japan. Transactions of the Natural History Society of Formosa 31: 413–415. [In German with Japanese summary.]

Takakuwa Y. 1942a. Die Myriopoden von Formosa, Philippinien, u. s. w. Transactions of the Natural History Society of Formosa 32 (231): 359–367.

Takakuwa Y. 1942b. Einige neue Arten von Diplopoda aus Nippon. Zoological Magazine 54: 237–239.

Takakuwa Y. 1942c. Zur Kenntnis der Japanischen Diplopoden. Annotationes Zoologicae Japonenses 21: 39–47.

Tamura K., Glen S. & Sudhir K. 2021. MEGA11: Molecular evolutionary genetics analysis version 11. Molecular Biology and Evolution 38: 3022–3027. https://doi.org/10.1093/molbev/msab120

Tanabe T. 1994. The millipede genus Levizonus (Polydesmida, Xystodesmidae) in Japan. Japanese Journal of Entomology 62 (1): 101–113.

Tanabe T. 2002. Revision of the millipede genus Parafontaria Verhoeff, 1936 (Diplopoda, Xystodesmidae). Journal of Natural History 36: 2139–2183. https://doi.org/10.1080/00222930110085610

Tanabe T. & Shinohara K. 1996. Revision of the millipede genus Xystodesmus, with reference to the status of the tribe Xystodesmini (Diplopoda: Xystodesmidae). Journal of Natural History 30: 1459–1494. https://doi.org/10.1080/00222939600770831

Tanabe T. & Sota T. 2008. Complex copulatory behavior and the proximate effect of genital and body size differences on mechanical reproductive isolation in the millipede genus Parafontaria. American Naturalist 171: 692–699. https://doi.org/10.1086/587075

Tanabe T. & Sota T. 2014. Both male and female novel traits promote the correlated evolution of genitalia between the sexes in an arthropod. Evolution 68 (2): 441–452. https://doi.org/10.1111/evo.12288

Tanabe T., Ishii K. & Yin W.Y. 1996. Two new xystodesmid millipeds [Arthropoda] from the Tian-mu mountains, Zhejiang province, China. Edaphologia 57: 13–19.

Trifinopoulos J., Nguyen L.T., von Haeseler A. & Minh B.Q. 2016. W-IQ-TREE: a fast online phylogenetic tool for maximum likelihood analysis. Nucleic Acids Reseach 44: W232–W235. https://doi.org/10.1093/nar/gkw256

Verhoeff K.W. 1936a. Über Diplopoden aus Japan, Gesammelt von Herrn Y. Takakuwa, Transactions of the Sapporo Natural History Society 54: 148–172.

Verhoeff K.W. 1936b. Zur Kenntnis ostasiatischer Strongylosomiden and Fontariiden. 149. Diplopoden-Aufsatz. Zoologischer Anzeiger 115 (11–12): 297–311.

Verhoeff K.W. 1937. Zur Kenntnis ostasiatischer Diplopoden. Zoologischer Anzeiger 117 (11/12): 309–321.

Verhoeff K.W. 1941. Über Gruppen der Leptodesmiden und neues System der Ordo Polydesmoidea. Archiv für Naturgeschichte 10 (3): 399–415.

Walker M.J., Stockman A.K., Marek P.E. & Bond J.E. 2009. Pleistocene glacial refugia across the Appalachian Mountains and coastal plain in the millipede genus Narceus: Evidence from population genetic, phylogeographic, and paleoclimatic data. BMC Evolutionary Biology 9: 1–11. https://doi.org/10.1186/1471-2148-9-25

Xu R., Chen J., Pan Y., Wang J., Chen L., Ruan H., Wu Y., Xu H., Wang G. & Liu H. 2022. Genetic diversity and population structure of Spirobolus bungii as revealed by mitochondrial DNA sequences. Insects 13: 729. https://doi.org/10.3390/insects13080729

Published
2026-02-05
How to Cite
Kim, G., Korsós, Z., & Hwang, U. W. (2026). The family Xystodesmidae in the Korean Peninsula (Diplopoda: Polydesmida). European Journal of Taxonomy, 1039(1), 70-115. https://doi.org/10.5852/ejt.2026.1039.3181
Section
Research article