Published January 29, 2026 | Version v1
Journal article Open

The millipede genus Mandarinopus Verhoeff, 1933, is a junior synonym of Kronopolites Attems, 1914 (Polydesmida, Paradoxosomatidae), with a revision of the species based on integrated evidence

  • 1. China Academy of Chinese Medical Sciences, Beijing, China
  • 2. Guizhou Academy of Sciences, Guiyang, China

Description

The millipede genus Mandarinopus Verhoeff, 1933 (Sulciferini, Paradoxosomatidae), endemic to western China, currently comprises five species. Its taxonomic delimitation from Kronopolites Attems, 1914, remains controversial because of morphological similarity and overlapping distributions. Previous studies relying exclusively on morphological characters yielded ambiguous results. To clarify the taxonomic status of Mandarinopus, we conducted an integrative taxonomic study using topotypic specimens of all species of Mandarinopus and representative species of Kronopolites. Phylogenetic analyses based on four genetic markers (COI, 16S, 18S, and 28S rRNA) reveal that Mandarinopus is polyphyletic and deeply nested within Kronopolites. Notably, the type species of both genera form a strongly supported sister pair. Morphological re-examination demonstrates that the diagnostic characters previously used to distinguish the genera, specifically gonopodal features, are continuous and lack clear boundaries. Consequently, we formally propose Mandarinopus Verhoeff, 1933, as a junior synonym of Kronopolites Attems, 1914, transferring all valid Mandarinopus species to Kronopolites. Additionally, based on morphological and molecular evidence from topotypic material, we propose Kronopolites rugosus Golovatch, 2013, as a junior synonym of Kronopolites corticinus (Attems, 1936).

Files

ZSE_article_180695.pdf

Files (10.7 MB)

Name Size Download all
md5:00eee1efa47170f5cfcefa3487d01caa
10.6 MB Preview Download
md5:dbb2866d8c03fbd682d643540e5e1287
141.3 kB Preview Download

Linked records

Additional details

References

  • Anisimova M, Gil M, Dufayard JF, Dessimoz C, Gascuel O (2011) Survey of branch support methods demonstrates accuracy, power, and robustness of fast likelihood-based approximation schemes. Systematic Biology 60: 685–699. https://doi.org/10.1093/sysbio/syr041
  • Attems C (1914) Die indo-australischen Myriapoden. Archiv für Naturgeschichte 80A: 1–398.
  • Attems C (1929) Diplopoden des Belgischen Congo. I. Polydesmoidea. Revue de Zoologie et de Botanique Africaines 17(3): 253–378.
  • Attems C (1931) Die Familie Leptodesmidae und andere Polydesmiden. Zoologica (Stuttgart) 79: 1–150.
  • Attems C (1936) Diplopoda of India. Memoirs of the Indian Museum 11(4): 133–353.
  • Attems C (1937) Myriapoda 3. Polydesmoidea I. Fam. Strongylosomidae. Das Tierreich 68: 1–300. https://doi.org/10.1515/9783111567099
  • Bernt M, Donath A, Jühling F, Externbrink F, Florentz C, Fritzsch G, Pütz J, Middendorf M, Stadler PF (2013) MITOS: Improved de Novo Metazoan Mitochondrial Genome Annotation. Molecular Phylogenetics and Evolution 2013(69): 313–319. https://doi.org/10.1016/j.ympev.2012.08.023
  • Chen CC, Golovatch SI, Chang HW (2006) The millipede tribe Sulciferini in Taiwan (Diplopoda: Polydesmida: Paradoxosomatidae). In: Meidell N, Hansen LO, Sømme L (Eds) Proceedings of the 13th International Congress of Myriapodology, Bergen, Norway, 15–29 July 2005. Norwegian Journal of Entomology 53(2): 249–270.
  • Golovatch SI (2009) On several new or poorly-known Oriental Paradoxosomatidae (Diplopoda: Polydesmida), IX. Arthropoda Selecta 18(3–4): 119–124.
  • Golovatch SI (2013a) On several new or poorly-known oriental paradoxosomatidae (Diplopoda: Polydesmida), XIII. Arthropoda Selecta 22(1): 1–31. https://doi.org/10.15298/arthsel.22.1.01
  • Golovatch SI (2013b) On several new or poorly-known Oriental Paradoxosomatidae (Diplopoda: Polydesmida), XIV. Arthropoda Selecta 22(4): 307–332. https://doi.org/10.15298/arthsel.22.4.01
  • Golovatch SI (2015) On several new or poorly-known Oriental Paradoxosomatidae (Diplopoda: Polydesmida), XVII. Arthropoda Selecta 24(2): 127–168. https://doi.org/10.15298/arthsel.24.2.01
  • Golovatch SI (2016) On several new or poorly-known Oriental Paradoxosomatidae (Diplopoda: Polydesmida), XVIII. Arthropoda Selecta 25(1): 1–18. https://doi.org/10.15298/arthsel.25.1.01
  • Golovatch SI (2019) On several new or poorly-known Oriental Paradoxosomatidae (Diplopoda: Polydesmida), XXVI. Arthropoda Selecta 28(3): 347–367. https://doi.org/10.15298/arthsel.28.3.01
  • Golovatch SI (2020) On several new or poorly-known Oriental Paradoxosomatidae (Diplopoda: Polydesmida), XXIX. Arthropoda Selecta 29(3): 297–308. https://doi.org/10.15298/arthsel.29.3.01
  • Golovatch SI, Liu W (2020) Diversity, distribution patterns, and faunogenesis of the millipedes (Diplopoda) of mainland China. ZooKeys 930: 153–198. https://doi.org/10.3897/zookeys.930.47513
  • Golovatch SI, Semenyuk II (2021) On several new or poorly-known Oriental Paradoxosomatidae (Diplopoda: Polydesmida), XXX. Arthropoda Selecta 30(4): 473–496. https://doi.org/10.15298/arthsel.30.4.04
  • Hoffman RL (1962) A contribution to the knowledge of Asiatic Strongylosomoid Diplopoda (Polydesmida: Strongylosomatidae). Annals and Magazine of Natural History Series 13(5): 577–593. https://doi.org/10.1080/00222936208651289
  • Hoffman RL (1980) Classification of the Diplopoda. Muséum d'histoire naturelle, Genève, 237 pp.
  • Jeekel CAW (1968) On the classification and geographical distribution of the family Paradoxosomatidae (Diplopoda, Polydesmida). Academisch Proefschrift, Rotterdam, 162 pp. [privately published].
  • Jeekel CAW (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, I–XII, 1–412.
  • Jeekel CAW (1982) New records and descriptions of southeast Asian Paradoxosomatidae (Diplopoda, Polydesmida). Bollettino del Museo civico di Storia naturale di Verona 9: 225–253.
  • Jeekel CAW (1988) The generic position of Orthomorpha bucharensis Lohmander and O. muminabadensis Gulicka, and the taxonomic status of Hedinomorpha Verhoeff (Diplopoda, Polydesmida, Paradoxosomatidae). Bulletin Zoölogisch Museum. Universiteit van Amsterdam 11(11): 97–104.
  • Jin JJ, Yu WB, Yang JB, Yu S, dePamphilis CW, Yi TS, Li DZ (2020) GetOrganelle: A fast and versatile toolkit for accurate de novo assembly of organelle genomes. Genome Biology 21: 241. https://doi.org/10.1186/s13059-020-02154-5
  • Johnson MG, Gardner EM, Liu Y, Medina R, Goffinet B, Shaw AJ, Zerega NJ, Wickett NJ (2016) HybPiper: Extracting coding sequence and introns for phylogenetics from high-throughput sequencing reads using target enrichment. Applications in Plant Sciences 4(7): apps.1600016. https://doi.org/10.3732/apps.1600016
  • Kalyaanamoorthy S, Minh BQ, Wong TKF, von Haeseler A, Jermiin LS (2017) ModelFinder: Fast model selection for accurate phylogenetic estimates. Nature Methods 14(6): 587–589. https://doi.org/10.1038/nmeth.4285
  • Kumar S, Stecher G, Li M, Knyaz C, 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
  • Likhitrakarn N, Golovatch SI, Panha S (2015) Review of the millipede genus Kronopolites Attems, 1914 (Diplopoda, Polydesmida, Paradoxosomatidae), with the description of a new species from Laos. ZooKeys (472): 27–41. https://doi.org/10.3897/zookeys.472.9001
  • Nguyen AD (2017) A second species of the genus Vietnamorpha Golovatch, 1984 (Polydesmida: Paradoxosomatidae) and notes on the generic relationship. Journal of Natural History 51(39–40): 2331–2343. https://doi.org/10.1080/00222933.2017.1355491
  • Nguyen AD, Sierwald P (2013) A worldwide catalog of the family Paradoxosomatidae Daday, 1889 (Diplopoda: Polydesmida). Check List 9(6): 1132–1353. https://doi.org/10.15560/9.6.1132
  • Nguyen LT, Schmidt HA, von Haeseler A, Minh BQ (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Phylogenetics and Evolution 32: 268–274. https://doi.org/10.1093/molbev/msu300
  • Nguyen AD, Korsós Z, Jang KH, Hwang UW (2017) A revision and phylogenetic analysis of the millipede genus Oxidus Cook, 1911 (Polydesmida: Paradoxosomatidae). European Journal of Taxonomy 293: 1–22. https://doi.org/10.5852/ejt.2017.293
  • Nguyen AD, Nguyen S, Le XS (2018) On the millipede genus Antheromorpha Jeekel, 1968 (Diplopoda: Polydesmida: Paradoxosomatidae) from Vietnam, with a description of a new species. The Raffles Bulletin of Zoology 66: 96–109.
  • Nguyen AD, Vu TTT, Phung HLT, Tran DL, Le HA (2025) Mountainous millipedes in Vietnam. V. The millipede genus Kronopolites Attems, 1914 (Diplopoda, Polydesmida, Paradoxosomatidae), with descriptions of two new species. ZooKeys 1249: 223–245. https://doi.org/10.3897/zookeys.1249.155280
  • Ronquist F, Teslenko M, van der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61: 539–542. https://doi.org/10.1093/sysbio/sys029
  • Takakuwa Y (1954) A Study of Japanese Diplopoda. A Monograph of Diplopoda of Japan, 241 pp.
  • Takashima H (1950) A general view of Japanese myriapods (II). Acta Arachnologica 11(1): 36–38. [in Japanese] https://doi.org/10.2476/asjaa.12.36
  • Verhoeff KW (1933) Schwedisch-chinesische wissenschaftliche Expedition nach den nordwestli Geoffroy Provinzen Chinas unter Leitung von Dr. Sven Hedin und Prof. Sü Ping-Chang. Myriapoda gesammelt vom schwedischen Arzt der Expedition Dr. David Hummel 1927–1930. Arkiv för Zoologi 26A(10): 1–41.
  • Verhoeff KW (1939) Zur Kenntnis ostasiatischer Diplopoden IV. Zoologischer Anzeiger 127(11–12): 273–285.
  • Wang DQ, Mauriès JP (1996) Review and perspective of study on myriapodology of China. In: Geoffroy JJ, Mauries JP, Nguyen Duy-Jacquemin M (Eds) Acta Myriapodologica, Mémoires du Museum National d'Histoire Naturelle 169: 81–99.
  • Xiong Y, Chen HM, Jiang XK, Jiang C (2025) Topotypes of the millipede species Kronopolites swinhoei (Pocock, 1895) reveal a new synonym with revalidation of Kronopolites svenhedini (Verhoeff, 1934) (Diplopoda, Polydesmida, Paradoxosomatidae). ZooKeys 1231: 85–98. https://doi.org/10.3897/zookeys.1231.137769