Integrative Taxonomy Reveals Two New Dragon Millipede Species in the Genus Desmoxytes Chamberlin, 1923 from Thailand (Diplopoda: Polydesmida: Paradoxosomatidae)

Authors

  • Ruttapon Srisonchai Department of Biology, Faculty of Science, Khon Kaen University, Khon Kaen 40002, THAILAND: Animal Systematics Research Unit, Department of Biology, Faculty of Science, Chulalongkorn University, Phayathai Road, Pathumwan, Bangkok 10330, THAILAND
  • Khanet Srikampha Department of Biology, Faculty of Science, Khon Kaen University, Khon Kaen 40002, THAILAND
  • Theemaporn Benchapong Department of Biology, Faculty of Science, Khon Kaen University, Khon Kaen 40002, THAILAND
  • Somsak Panha Animal Systematics Research Unit, Department of Biology, Faculty of Science, Chulalongkorn University, Phayathai Road, Pathumwan, Bangkok 10330, THAILAND: Academy of Science, The Royal Society of Thailand, Bangkok 10300, THAILAND

DOI:

https://doi.org/10.58837/tnh.25.8.266903

Keywords:

biodiversity, evolution, karst, Southeast Asia, systematics

Abstract

Recent field surveys in northern Thailand led to the discovery of two remarkable new species of dragon millipede in the genus Desmoxytes Chamberlin, 1923. Using an integrative approach that combined morphological and molecular data, we confirmed the validity of these species, herein described as Desmoxytes chaofa Srisonchai, sp. nov. from Mae Hong Son Province and Desmoxytes sirindhornae Srisonchai, sp. nov. from Tak Province. To assess their systematic position and relationships, we conducted morphological examinations and reconstructed the phylogeny based on three gene fragments (COI, 16S rRNA, and 28S rRNA). Phylogenetic analysis supports the validity of these two new species, with genetic divergence in the COI gene clearly distinguishing them from congeners, ranging from 10.67–18.67% for D. chaofa Srisonchai, sp. nov. and 10.05–17.91% for D. sirindhornae Srisonchai, sp. nov. (mean = 14.71%). Notably, the phylogenetic tree aligns well with morphological characters, particularly those of the gonopod. Desmoxytes chaofa Srisonchai, sp. nov. is closely related to D. octoconigera Srisonchai, Enghoff & Panha, 2018, while D. sirindhornae Srisonchai, sp. nov. clusters together with D. des Srisonchai, Enghoff & Panha, 2016, both pairs exhibiting similarities in body colouration and gonopod telopodite. Based on current data, these two species are considered as narrow endemics and are restricted to limestone rock wall habitats. Complete morphological illustrations of the two new species and a distribution map of all Desmoxytes species are provided.

References

American Veterinary Medical Association. 2020. AVMA Guidelines for the Euthanasia of Animals: 2020 edition. Available at: https://www.avma.org/sites/default/files/2020-02/Guidelines-on-Euthanasia-2020.pdf.

Clements, R., Sodhi, N.S., Schilthuizen, M. and Ng, P.K.L. 2006. Limestone karsts of Southeast Asia: Imperiled arks of biodi-versity. BioScience, 56: 733–742. https://doi.org/10.1641/ 0006-3568(2006)56[733:lkosai]2.0.co;2

Colgan, D.J., Ponder, W.F. and Eggler, P.E. 2000. Gastropod evolutionary rates and phylogenetic relationships assessed using partial 28S rDNA and histone H3 sequences. Zoologica Scripta, 29: 29–63. https://doi.org/10.1046/j.1463-6409.2000. 00021.x

Decker, P. 2016. Integrative taxonomic revision of the polymorphic flat-millipede genera Oncocladosoma and Somethus in South Australia (Diplopoda: Polydesmida: Paradoxosomatidae). Invertebrate Systematics, 30: 201–218. https://doi.org/10.1071 /is15047

Decker, P. 2016b. Phylogenetic analysis of the Australian trans-Bass Strait millipede genus Pogonosternum (Carl, 1912) (Diplopoda, Polydesmida, Paradoxosomatidae) indicates mul-tiple glacial refugia in southeastern Australia. ZooKeys, 578: 15–31. https://doi.org/10.3897/zookeys.578.8052

Enghoff, H., Sutcharit, C. and Panha, S. 2007. The shocking pink dragon millipede, Desmoxytes purpurosea sp. n., a colourful aposematic millipede (Diplopoda: Polydesmida: Paradoxoso-matidae) from Thailand. Zootaxa, 1563: 31–36. https://doi.org/ 10.11646/zootaxa.1563.1.3

Erixon, P., Svennblad, B., Britton, T. and Oxelman, B. 2003. Reliability of Bayesian posterior probabilities and bootstrap frequencies in phylogenetics. Systematic Biology, 52: 665–673. https://doi.org/10.1080/10635150390235485

Folmer, O., Back, M., Hoeh, W., Lutz, R. and 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., Li, Y., Liu, W. and Geoffroy, J.J. 2012. Three new cavernicolous species of dragon millipedes, genus Desmoxytes Chamberlin, 1923, from southern China, with notes on a formal congener from the Philippines (Diplopoda, Polydesmida, Paradoxosomatidae). ZooKeys, 185: 1–17. https://doi.org/10. 3897/zookeys.185.3082

Huelsenbeck, J.P., Hillis, D.M. 1993. Success of phylogenetic methods in the four-taxon case. Systematic Biology, 42: 247–264. https://doi.org/10.2307/2992463

Jeekel, C.A.W. 1964. Two new species of Pratinus Attems, with taxonomic notes on the genus and a redescription of its type-species (Diplopoda, Polydesmida). Beaufortia, 11: 61–73.

Kalyaanamoorthy, S., Minh, B.Q., Wong, T.K.F., von Haeseler, A. and Jermiin, L.S. 2017. ModelFinder: fast model selection for accurate phylogenetic estimates. Nature Methods, 14: 587–589. https://doi.org/10.1038/nmeth.4285

Kumar, S., Stecher, G. and Tamura, K. 2016. MEGA7: Molecular evolutionary genetics analysis version 7.0 for bigger datasets. Molecular Biology and Evolution, 33: 1870–1874. https://doi. org 10.1093/molbev/msw054

Lanfear, R., Calcott, B., Ho, S.Y. and Guindon, S. 2012. PartitionFinder: combined selection of partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology and Evolution, 29: 1695–1701. https://doi.org/10.1093/ molbev/mss020

Larget, B., Simon, D.L. 1999. Markov Chain Monte Carlo algorithms for the Bayesian analysis of phylogenetic trees. Molecular Biology and Evolution, 16: 750–750. https://doi.org/ 10.1093/oxfordjournals.molbev.a026160

Likhitrakarn, N., Golovatch, S., Panha, S. 2011. Revision of the Southeast Asian millipede genus Orthomorpha Bollman, 1893, with the proposal of a new genus (Diplopoda, Polydesmida, Paradoxosomatidae). ZooKeys, 131: 1–161. https://doi.org/ 10.3897/zookeys.131.1921

Likhitrakarn, N., Jeratthitikul, E., Jirapatrasilp, P. and Wesener, T. 2024. Integrated taxonomy of three new species of Glyphiulus Gervais, 1847 (Diplopoda, Spirostreptida, Cambalopsidae) from Laos. Raffles Bulletin of Zoology, 72: 42–61.

Liu, W., Golovatch, S. and Tian, M. 2014. A review of the dragon millipede genus Desmoxytes Chamberlin, 1923 in China, with descriptions of four new species (Diplopoda, Polydesmida, Paradoxosomatidae). ZooKeys, 448: 9–26. https://doi.org/ 10.3897/zookeys.448.8081

Liu, W., Golovatch, S. and Tian, M. 2016. Six new species of dragon millipedes, genus Desmoxytes Chamberlin, 1923, mostly from caves in China (Diplopoda, Polydesmida, Parado-xosomatidae). ZooKeys, 577: 1–24. https://doi.org/10.3897/ zookeys.577.7825

Means, J.C., Hennen, D.A. and Marek, P.E. 2021a. A revision of the minor species group in the millipede genus Nannaria Chamberlin, 1918 (Diplopoda, Polydesmida, Xystodesmidae). ZooKeys, 1030: 1–180. https://doi.org/10.3897/zookeys.1030. 62544

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

Minh, B.Q., Schmidt, H.A., Chernomor, O., Schrempf, D., Wood-hams, M.D., von Haeseler and A. and Lanfear, R. 2020 IQ-TREE 2: New models and efficient methods for phylogenetic inference in the genomic era. Molecular Biology and Evolution, 37: 1530–1534. https://doi.org/10.1093/molbev /msaa015

Nguyen, A.D., Nguyen, M.-H., Nguyen, T.-A.T. and Phung, H.-L.T. 2019. Review of dragon millipedes (Diplopoda, Polydes-mida, Paradoxosomatidae) in the fauna of Vietnam, with descriptions of three new specie. Zoological Studies, 58: 14. https://doi.org/10.6620/ZS.2019.58-14

Pocock, R.I. 1895. The Myriopoda of Burma, Pt IV Report upon the Polydesmoidea collected by Sig L Fea, Mr E. W. Oates and others, by R.I. Pocock of the British Museum of Natural History. Annali del Museo Civico di Storia Naturale Giacomo Doria, 14: 787–834.

Rambaut, A. 2010. FigTree v1.3.1. Available at http://tree.bio.ed. ac.uk/software/figtree/

Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. and Huelsen-beck, J.P. 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

Srisonchai, R., Enghoff, H., Likhitrakarn, N. and Panha, S. 2016. Four colorful new species of dragon millipedes, genus Desmoxytes Chamberlin, 1923, from northern Thailand (Diplo-poda: Polydesmida: Paradoxosomatidae). Zootaxa, 4710: 93–113. http://dx.doi.org/10.11646/ zootaxa.4170.1.4

Srisonchai, R., Enghoff, H., Likhitrakarn, N. and Panha, S. 2018a. A revision of dragon millipedes I: genus Desmoxytes Chamberlin, 1923, with the description of eight new species (Diplopoda, Polydesmida, Paradoxosomatidae). ZooKeys, 761: 1–177. https://doi.org/10.3897/zookeys.761.24214

Srisonchai, R., Enghoff, H., Likhitrakarn, N. and Panha, S. 2018b. A revision of dragon millipedes II: the new genus Nagaxytes gen. nov., with the description of three new species (Diplopoda, Polydesmida, Paradoxosomatidae). European Journal of Taxonomy, 462: 1–44. https://doi.org/10.5852/ejt.2018.462

Srisonchai, R., Enghoff, H., Likhitrakarn, N. and Panha, S. 2018c. A revision of dragon millipedes III: the new genus Gigaxytes gen. nov., with the description of three new species (Diplopoda, Polydesmida, Paradoxosomatidae). European Journal of Taxonomy, 463: 1–43. https://doi.org/10.5852/ejt.2018.463

Srisonchai, R., Enghoff, H., Likhitrakarn, N. and Panha, S. 2018d. A revision of dragon millipedes IV: the new genus Spinaxytes, with the description of nine new species (Diplopoda, Polydes-mida, Paradoxosomatidae). ZooKeys, 797: 19–69. https://doi. org/10.3897/zookeys.797.29510

Srisonchai, R., Lin, A. and Panha, S. 2020. A revision of dragon millipedes V. The new genus Burmaxytes for two new species from Myanmar (Diplopoda: Polydesmida: Paradoxosomatidae). Raffles Bulletin of Zoology, 35: 88–100.

Srisonchai, R., Enghoff, H., Likhitrakarn, N., Jeratthitikul, E., Jirapatrasilp, P., Panha, S. and Sutcharit, C. 2024. Molecular phylogeny of dragon millipedes (Diplopoda: Polydesmida: Paradoxosomatidae) from mainland South-East Asia, with description of a new genus and species. Zoological Journal of the Linnean Society, 202: 1–33. https://doi.org/10.1093/ zoolinnean/zlad164

Srisonchai, R., Enghoff, H., Likhitrakarn, Panha, S. and Sutcharit, C. 2025. Molecular and morphological data uncover a striking new genus of dragon millipedes in Thailand, with alternately long and short legs (Diplopoda, Polydesmida, Paradoxoso-matidae). Invertebrate Systematics, 39: IS25007. https://doi.org /10.1071/IS25007

Simon, C., Frati, F., Beckenbach, A., Crespi, B., Liu, H. and Flook, P. 1994. Evolution, weighting, and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved Polymerase Chain Reaction primers. Annals of the Entomo-logical Society of America, 87: 651–701. https://doi.org/10. 1093/aesa/87.6.651

Willett, W., Jenny, B., Isenberg, T. and Dragicevic, P. 2015. Lightweight relief shearing for enhanced terrain perception on interactive maps. In: Proceedings of the 33rd ACM Conference on Human Factors in Computing Systems (CHI 2015), Seoul, South Korea, pp. 3563–3572. Available at: http://elasticterrain. xyz/map/ [Accessed 10 March 2025].

Zuo, Q., Zhang, Z. and Shen, Y. 2022. Novel mitochondrial gene rearrangements pattern in the millipede Polydesmus sp. GZCS-2019 and phylogenetic analysis of the Myriapoda. Ecology and Evolution, 12(3): e8764. https://doi.org/10.1002/ece3.8764

Published

2025-10-14

How to Cite

[1]
Srisonchai, R., Srikampha, K., Benchapong, T. and Panha, S. 2025. Integrative Taxonomy Reveals Two New Dragon Millipede Species in the Genus Desmoxytes Chamberlin, 1923 from Thailand (Diplopoda: Polydesmida: Paradoxosomatidae). Tropical Natural History. 25, 8 (Oct. 2025), 79–107. DOI:https://doi.org/10.58837/tnh.25.8.266903.