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Two new species of the genus Thereuopoda Verhoeff, 1904 (Scutigeromorpha, Scutigeridae) from Sichuan and Hainan Provinces, China

Ji, Jie-Hong; Shen, Chen-Yang; Gao, Yi-Xin; Wu, Hui-Yuan; Yu, Dan-Na; Liu, Sheng-Long; Zhang, Jia-Yong

Abstract

Scutigeromorph centipedes are morphologically distinctive myriapods that have historically received limited scientific attention, particularly in China. The taxonomy of the genus Thereuopoda Verhoeff, 1904 has remained largely unresolved since the late 1970s, and recent molecular evidence has indicated the presence of cryptic species within this group. In this study, we describe two new species, T. kaijiangensis sp. nov. Ji, Shen & Zhang from Kaijiang County, Dazhou City, Sichuan Province, China, and T. edgecombei sp. nov. Ji, Liu & Zhang from Chengmai County, Hainan Province, China. Comprehensive morphological descriptions and detailed illustrations are provided for both species, along with an updated diagnostic key to all currently recognized Thereuopoda species in China. Phylogenetic analyses conducted using both maximum-likelihood and Bayesian-inference methods consistently support the monophyly of the genus Thereuopoda and further validate the taxonomic distinctness of the two newly described species.

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351 Two new species of the genus Thereuopoda Verhoeff, 1904 (Scutigeromorpha, Scutigeridae) from Sichuan and Hainan Provinces, China Jie-Hong Ji1, Chen-Yang Shen1, Yi-Xin Gao1, Hui-Yuan Wu1, Dan-Na Yu1,2 , Sheng-Long Liu3, Jia-Yong Zhang1,2 1 College of Life Sciences, Zhejiang Normal University, Jinhua, 321004, Zhejiang Province, China 2 Key Lab of Wildlife Biotechnology, Conservation and Utilization of Zhejiang Province, Zhejiang Normal University, Jinhua, 321004, Zhejiang Province, China 3 Longquan Conservation Center of Qianjiangyuan-Baishanzu National Park, Longquan, 323000, Zhejiang Province, China Corresponding authors: Jia-Yong Zhang ([email protected], zhangjiay[email protected]); Sheng-Long Liu ([email protected]) Copyright: © Jie-Hong Ji et al. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International – CC BY 4.0). Research Article Abstract Scutigeromorph centipedes are morphologically distinctive myriapods that have historically received limited scientific attention, particularly in China. The taxonomy of the genus Thereuopoda Verhoeff, 1904 has remained largely unresolved since the late 1970s, and recent molecular evidence has indicated the presence of cryptic species within this group. In this study, we describe two new species, T. kaijiangensis sp. nov. Ji, Shen & Zhang from Kaijiang County, Dazhou City, Sichuan Province, China, and T. edgecombei sp. nov. Ji, Liu & Zhang from Chengmai County, Hainan Province, China. Comprehensive morphological descriptions and detailed illustrations are provided for both species, along with an updated diagnostic key to all currently recognized Thereuopoda species in China. Phylogenetic analyses conducted using both maximum-likelihood and Bayesian-inference methods consistently support the monophyly of the genus Thereuopoda and further validate the taxonomic distinctness of the two newly described species. Key words: Centipede, molecular phylogeny, morphology, scutigeromorph, taxonomy Introduction Scutigeromorpha, the most morphologically distinct of the five centipede orders, is characterized by a suite of unique morphological features, including multisegmented tarsi, a domed head capsule, compound eyes, and dorsally positioned spiracles located on the tergites (Perez-Gelabert and Edgecombe 2013). The family Scutigeridae constitutes the largest family in this order, comprising approximately 18 genera and 90 species (Porta and Giribet 2024). Two subfamilies are currently recognized in Scutigeridae: Scutigerinae Leach, 1814 and Thereuoneminae Verhoeff, 1905. The genus Thereuopoda Verhoeff, 1904, distributed across East and Southeast Asia, is classified within the subfamily Thereuoneminae (Bonato et al. 2016). The earliest Thereuopoda species discovered in Hong Kong is Thereuopoda clunifera (Wood, 1862), which was originally placed in the genus Cermatia Academic editor: Pavel Stoev Received: 15 July 2025 Accepted: 23 November 2025 Published: 19 December 2025 ZooBank: https://zoobank. org/9000110B-1588-461A-9DC50D4A8B43CED9 Citation: Ji J-H, Shen C-Y, Gao Y-X, Wu H-Y, Yu D-N, Liu S-L, Zhang J-Y (2025) Two new species of the genus Thereuopoda Verhoeff, 1904 (Scutigeromorpha, Scutigeridae) from Sichuan and Hainan Provinces, China. ZooKeys 1264: 351–376. https://doi. org/10.3897/zookeys.1264.165241 ZooKeys 1264: 351–376 (2025) DOI: 10.3897/zookeys.1264.165241 352 ZooKeys 1264: 351–376 (2025), DOI: 10.3897/zookeys.1264.165241 Jie-Hong Ji et al.: Two new species of Thereuopoda from China (Wood 1862). Subsequently, further taxonomic and faunistic studies were conducted by Meinert (1886), Haase (1887), and Pocock (1895). The genus Thereuopoda was originally established as a subgenus by Verhoeff (1904) and comprised four species, T. rubrolineata Verhoeff, 1904, T. longicornis (Fabricius, 1793), T. clunifera (Wood, 1862), and T. amokiana Verhoeff, 1904. Subsequently, Verhoeff (1905) elevated Thereuopoda to generic rank and subdivided it into three subgenera: Orthothereua Verhoeff, 1905, Thereuopoda sensu stricto Verhoeff, 1904, and Microthereua Verhoeff, 1905. During the first half of 20th century, numerous new species of Thereuopoda were described by Chamberlin (1944) and Verhoeff (1925, 1936, 1937, 1939, 1942, 1943, 1944), including Thereuopoda nivicomes Verhoeff, 1942 from China (“Quellgebiet des Yangtsekiang bei Dji-tu im Hzifan-Bergland”). Verhoeff (1936) also established a new genus Teleotelson Verhoeff, 1936 for two species previously misassigned to Thereuonema by Silvestri (1924). In the 1950s, Chamberlin and Wang reported T. clunifera from Taiwan Island (Chamberlin and Wang 1952; Wang 1955, 1956, 1959). In 1979, the taxonomy of Thereuopoda underwent comprehensive revision (Würmli 1979). The genus Teletelson and the subgenus Orthothereua were recognized as junior synonyms of Thereuopoda and many species described by early taxonomists were treated as junior synonyms of T. longicornis and T. clunifera. The taxonomic status of Thereuopoda chinensis Verhoeff, 1905 and its subgenus Microthereua remains uncertain due to the absence of examined type material. Since 1979, only a few publications have reported records of Thereuopoda species (Stoev 2002; Stoev and Geoffroy 2004; Li 2007; Xu et al. 2013; Dunlop et al. 2017; Lei et al. 2021; Huang et al. 2023). To date, three valid species of Thereuopoda have been documented in China, T. (M.) chinensis, T. clunifera, and T. longicornis. The generic placement of two poorly known taxa—Scutigera complanata Haase, 1887 and Scutigera sinuata Haase, 1887—remains uncertain (Haase 1887). The remarkable diversity and complexity of China’s zoogeographical and geological characteristics have contributed to an exceptionally rich faunal assemblage. Recent studies have revealed cryptic species within the genus Thereuopoda and high diversity in Thereuonema tuberculata (Wood, 1862) through molecular data analyses (Yang et al. 2022; Manivannan et al. 2024; Ji et al. 2025). In this study, two new species—Thereuopoda kaijiangensis sp. nov. Ji, Shen & Zhang and Thereuopoda edgecombei sp. nov. Ji, Liu & Zhang—are described, and their phylogenetic relationships are analyzed. The distribution of the five known Thereuopoda species in China is shown in Fig. 1. Materials and methods Specimens All samples were collected using the button tube method and preserved in 80% ethanol in 2023 and 2024. Specimens were examined under an Olympus SZX16 stereomicroscope (Olympus Corporation, Tokyo, Japan) and a NOVEL DN-401 biological microscope (Ningbo Yongxin Optics Co. Ltd, Ningbo, China). Images were captured with an Olympus DP73 digital camera (Olympus Corporation, Tokyo, Japan). The raw photographs were processed through image alignment and focus-stacking in Affinity Photo v. 2.6.0 (Serif Ltd, Nottingham, UK) to generate composite images, which were subsequently refined with Adobe Photoshop CC 353 ZooKeys 1264: 351–376 (2025), DOI: 10.3897/zookeys.1264.165241 Jie-Hong Ji et al.: Two new species of Thereuopoda from China 2019 (Adobe Systems Inc., Los Angeles, CA, USA). Type specimens are deposited in the Animal Herbarium at Zhejiang Normal University, Jinhua, China (ZJNU). Morphological terminology follows that of Edgecombe and Giribet (2006) and Edgecombe (2011). The measurement protocol for the female gonopod follows Würmli (1973). Tergites are abbreviated as TT1–TT8. Figure 1. The distribution of the five known Thereuopoda species in China. T. chinensis: Macao. T. clunifera: Taitung County, Haulien County, Nantou County, Changhua County and Taipei City, Taiwan Province; Hong Kong; Nanjing City, Jiangsu Province; Hangzhou City, Shaoxing City and Ningbo City, Zhejiang Province; Guangzhou City, Foshan City and Shanwei City, Guangdong Province; Guigang City and Laibin City, Guangxi Province; Pingdingshan City, Henan Province; Beijing Municipality; Shanghai Municipality; Jiujiang City, Jiangxi Province; Bijie City and Libo County, Guizhou Province; Longshan County, Hunan Province. Suizhou City, Hubei Province. T. kaijiangensis sp. nov.: Dazhou city, Sichuang Province. T. edgecombei sp. nov.: Chengmai County, Hainan Province. T. longicornis: Guangzhou City, Guangdong Province; Qamdo City, Tibet; Menzi County, Yunnan Province; Qinshui County, Shanxi Province. 354 ZooKeys 1264: 351–376 (2025), DOI: 10.3897/zookeys.1264.165241 Jie-Hong Ji et al.: Two new species of Thereuopoda from China Molecular analyses Six molecular markers (18S rRNA, 28S rRNA, H3, 16S rRNA, 12S rRNA, and COX1), previously used in scutigeromorph phylogenetic analyses (Giribet and Edgecombe 2013; Manivannan et al. 2024), were sequenced to assess the phylogenetic position of the two new species. Two legs of the holotype were used to extract genomic DNA with the Ezup Column Animal Genomic DNA Purification Kit (Sangon Biotech Co., Shanghai, China). Genomic DNA extraction and subsequent library preparation were performed by BerryGenomics (Beijing, China) for next-generation sequencing (NGS). Raw NGS data were quality-filtered and de novo assembled using MEGAHIT v. 1.2.9 (Li et al. 2015). The nuclear ribosomal genes 18S rRNA and 28S rRNA were obtained using Barrnap v. 0.9 (Seemann 2013). The histone H3 gene was identified using HMMER v. 3.4 (Eddy 2011). The three mitochondrial genes (16S rRNA, 12S rRNA, and COX1) were obtained following the protocol described in Ji et al. (2025). All newly generated sequences have been deposited in GenBank (Suppl. material 1). The final data used for phylogenetic analyses are summarized in Suppl. material 1. Six genes were aligned using MAFFT v. 7.475 (Katoh and Standley 2013), and the resulting alignments were trimmed with Gblocks v. 0.91b (Castresana 2000). The individual gene alignments were concatenated using PhyloSutie v. 1.2.2 (Zhang et al. 2020). Partitioning schemes and best-fitting models for each partition were determined using PartitionFinder v. 2.2.1 (Lanfear et al. 2012) (Suppl. material 2). The maximum-likelihood (ML) tree was inferred using RaxML v. 8.2.0 (Stamatakis 2014) with 1,000 bootstrap replicates. The Bayesian-inference (BI) tree was conducted in MrBayes v. 3.2.7a (Ronquist et al. 2012) with 10 million MCMC generations, sampling every 1,000 generations and applying a burn-in of 25%. The resulting phylogenetic trees were visualized and edited using Affinity Photo v. 2.6.0 (Serif Ltd, Nottingham, UK). Genetic distances for COX1 were calculated in MEGA v. 11 (Tamura et al. 2021) using the Kimura 2-parameter model. Results Order Scutigeromorpha Pocock, 1895 Family Scutigeridae Gervais, 1837 Subfamily Thereuoneminae Verhoeff, 1925 Genus Thereuopoda Verhoeff, 1904 Thereuopoda kaijiangensis Ji, Shen & Zhang, sp. nov. https://zoobank.org/41807141-54CA-4320-B932-DFA2BC1656C8 Figs 2–7; Tables 1, 2 Type material. Holotype. China • Male; Sichuan Province, Dazhou City, Kaijiang County, Yongxing Town; 31°08'N, 107°53'E; 11 May 2024; Jiehong Ji leg.; ZJNU SCKJ102. Paratype. China • 3 females; same data as for holotype; ZJNU SCKJ129–131. Diagnosis. Thereuopoda with body length up to 40 mm (Fig. 2). Head capsule yellow with a reddish-brown medial patch surrounded by dark-brown pigment; radiating network of dark-brown pigment on the posterolateral part 355 ZooKeys 1264: 351–376 (2025), DOI: 10.3897/zookeys.1264.165241 Jie-Hong Ji et al.: Two new species of Thereuopoda from China Figure 2. Thereuopoda kaijiangensis sp. nov., paratype female. Scale bars: 6.0 mm. of head capsule (Fig. 3A, B). Tergites pale yellow with three longitudinal darkbrown stripes; transverse projections between central and lateral stripes; a narrow yellow stripe in the middle of central stripe (Figs 3A, 4C). Stoma saddles blue-gray; reddish-purple pigment present laterally (Fig. 4C). Posterior margin of tergites unevenly rounded. Spiracle on T6 about 75–85% length of the stoma saddle. Posterior margin of T8 with slight median concavity (Fig. 4E). Leg 15 2.85–3.14× body length. Tibial spine-bristles 0/2 on leg 1. Maximum length of female gonopod 2.25–2.6× maximum width (Fig. 6B, C). Hairs absent in the central area of proarthron; angle at median distal end of proarthron about 130° (Fig. 6C). Sinus between mesarthron broadly parabolic (Fig. 6B). Posterior termination of female subanal plate pointed or with process (Fig. 5G). Length of first male gonopod 1.5× second male gonopod (Figs 5H, 6A). 356 ZooKeys 1264: 351–376 (2025), DOI: 10.3897/zookeys.1264.165241 Jie-Hong Ji et al.: Two new species of Thereuopoda from China Description. Length up to 39 mm in largest female, 40 mm in largest male. Colour: head capsule yellow with a reddish-brown patch medially surrounded by dark brown pigment; radiating network of dark brown pigment in the posterolateral part of head capsule (Fig. 3A, B). Compound eyes bright red. Tergites pale yellow with three longitudinal dark-brown stripes; transverse projections between central and lateral stripes; a narrow yellow stripe in the middle of central stripe (Figs 3A, 4C). Stoma saddles blue-gray; reddish-purple pigment present laterally. Leg yellow with blackish-green pigment on prefemur and femur (Fig. 4H). Sternites and female gonopod yellow (Figs 4F, 6B). Head capsule: anterior projection of sutures with divergent posterior part; lateral edge blunt and rounded (Fig. 3B). Posteromedian impression moderately deep. Antenna 1.57–1.8× body length. First flagellum with 73–78 articles; ring-like articles with dense hairs and a few setae forming a single whorl surrounding distal end of article (Fig. 3C). Epipharynx: labral median arc 1.94–2.28× depth of labral mid-piece tooth (Fig. 3D). Mandible: three teeth, all smooth-surfaced and tricuspid (Fig. 3F). Nearly 23 pectinate lamellae on gnathal edge. Table 2. Number of segments in Tarsus I and Tarsus II of Thereuopoda kaijiangensis sp. nov. Leg Tarsus I Tarsus II 124–26 54–64 220–21 54–63 317–18 52–54 4 17–18 51–54 5 16–21 45–50 6 14–16 46–49 7 13–16 45–50 8 14–19 40–51 9 15–16 43–49 10 14–16 48–49 11 11–15 49–52 12 14–16 46–54 13 13–14 54–55 14 15–23 56–60 Table 1. Number of spines on stoma saddles and whole tergites of Thereuopoda kaijiangensis sp. nov. Tergites Stoma saddles Whole tergites 10 0 20–18 13–39 350–79 120–157 4 56–94 220–267 5 70–90 193–197 6 58–86 149–186 7 24–45 81–114 8 – 3–4 357 ZooKeys 1264: 351–376 (2025), DOI: 10.3897/zookeys.1264.165241 Jie-Hong Ji et al.: Two new species of Thereuopoda from China Figure 3. Thereuopoda kaijiangensis sp. nov., holotype male. A. Habitus, dorsal view; B. Head capsule; C. First flagellum of antenna; D. Epipharynx; E. First maxillae; F. Mandible. Scale bars: 4.0 mm (A); 1.0 mm (B); 0.2 mm (C); 0.4 mm (D–F). 358 ZooKeys 1264: 351–376 (2025), DOI: 10.3897/zookeys.1264.165241 Jie-Hong Ji et al.: Two new species of Thereuopoda from China Figure 4. Thereuopoda kaijiangensis sp. nov., holotype male. A. Forcipular segment; B. Second maxilla; C. Tergite 6; D. Details of tergite 6; E. Tergite 8; F. Sternite 15; G. Details of sternite 15; H. Leg 10. Scale bars: 1.0 mm (A–C); 0.2 mm (D, G); 0.4 mm (E, F); 4.0 mm (H). 359 ZooKeys 1264: 351–376 (2025), DOI: 10.3897/zookeys.1264.165241 Jie-Hong Ji et al.: Two new species of Thereuopoda from China Figure 5. Thereuopoda kaijiangensis sp. nov., holotype male. A. Leg 10 prefemur; B. Leg 10 femur; C. Leg 10 tibia; D. Leg 10 tarsus I; E. Leg 10 tarsus II, ventral view, showing resilient sole hairs (r.s.h.) and tarsal papillae (t.p.); F. Male subanal plate; H. First male gonopod. Paratype female. G. Female subanal plate. Scale bars: 0.4 mm (A, B, G); 0.2 mm (C–F, H). 366 ZooKeys 1264: 351–376 (2025), DOI: 10.3897/zookeys.1264.165241 Jie-Hong Ji et al.: Two new species of Thereuopoda from China (Figs 10D, 13A, B). Isolated Tastborsten with short, paired spines at bases on all tergites; these spines joined at a common base. Basal spines 10–20% as long as associated bristle. All tergites with spiculae and spinulae. Short, triangular spiculae separated by several polygonal scales that lack spiculae. Legs: tarsus I and II segmentation as shown in Table 4. Leg 15 length 1.79– 1.94× that of body. Prefemoral spine-bristles in a 2/1 pattern on legs 1–15; femoral spine-bristles 1/2 on all legs; tibial spine-bristles 0/1 on leg 1, 0/1 or 0/2 on leg 2, 1/2 on leg 3–15. Spine rows on prefemur, femur and tibia with spine– setae pairing (Fig. 11A–C). Tarsus I of leg 6–14 with spines (Fig. 11D). Tarsus II of legs 1–14 with paired and equally sized tarsal papillae on successive segments except for first and last few segments on each leg (Fig. 11E). Resilient sole hairs originating near posteromedial edge of tarsal papilla, extending to the succeeding segment. Dense tuft of setae on ventrolateral side of tarsus II. Sternites: median embayment in posterior margin of sternites lacking or shallow (Fig. 10F). Longitudinal median furrow observable passing through entire length of the sternites. Minute, multifurcating spinulae and scattered setae with paired short spines in all sternites (Figs 10G, 13C). Female: gonopod with maximum length 2.03–2.37× maximum width. Longitudinal median suture in proarthron complete (Fig. 12B). Lateral margins of Table 4. Number of segments in Tarsus I and Tarsus II of Thereuopoda edgecombei sp. nov. Leg Tarsus I Tarsus II 114–20 42–47 213–16 41–44 312–14 40–45 4 11–14 35–41 5 12–14 32–37 6 10–18 28–37 7 9–11 32–36 8 10–13 31–39 9 10–12 34–37 10 11–12 37–39 11 10–14 34–44 12 12–18 30–44 13 11–13 36–47 14 13–15 40–51 Table 3. Number of spines on stoma saddle and whole tergites of Thereuopoda edgecombei sp. nov. Tergites Stoma saddles Whole tergites 10 0 21–9 9–14 325–39 71–132 4 43–66 260–401 5 48–72 185–303 6 46–54 160–282 7 26–31 89–184 8 – 2–9 367 ZooKeys 1264: 351–376 (2025), DOI: 10.3897/zookeys.1264.165241 Jie-Hong Ji et al.: Two new species of Thereuopoda from China Figure 11. Thereuopoda edgecombei sp. nov., holotype male. A. Leg 10 prefemur; B. Leg 10 femur; C. Leg 10 tibia; D. Leg 10 tarsus I; E. Leg 10 tarsus II, ventral view, showing resilient sole hairs (r.s.h.) and tarsal papillae (t.p.); F. Male subanal plate. Paratype female. G. Female subanal plate; H. Female subanal plate. Scale bars: 0.4 mm (A, D); 0.2 mm (B, C, E–H). 368 ZooKeys 1264: 351–376 (2025), DOI: 10.3897/zookeys.1264.165241 Jie-Hong Ji et al.: Two new species of Thereuopoda from China proarthron and mesarthron divergent posteriorly. Subtriangular depression on proarthron without setae. Hairs and setae on middle area of proarthron (Fig. 12C). Angle at median distal end of proarthron about 100°. Proarthron 1.18–1.42× length of mesarthron. Distomedial corner of mesarthron with a cluster of 12–15 setae. Sinus between mesarthron parabolic, broad with apex distinctly or weakly pointed. Width of mesarthron 0.54–0.66× maximum width of sinus. Proarthron + mesarthron 1.77–1.87× length of metarthron. Outer margin of metarthron uniformly curved. Subanal plate with ventral margin straighter than curved dorsal margin, posterior termination pointed or with process; maximum length 2.69–3.31× maximum height; smooth, non-setose band along Figure 12. Thereuopoda edgecombei sp. nov., holotype male. A. Male gonopod; B. Female gonopod; C. Proarthron of female gonopod; D. Male telson. Paratype male. E. Female telson. Scale bars: 0.2 mm (A, B, D); 0.1 mm (C); 0.3 mm (E). 369 ZooKeys 1264: 351–376 (2025), DOI: 10.3897/zookeys.1264.165241 Jie-Hong Ji et al.: Two new species of Thereuopoda from China Figure 13. Thereuopoda edgecombei sp. nov., paratype female. A. Tergite 6. B. Tergite 8. C. Sternite 15. Scale bars: 20 μm. middle of subanal plate; setae of varied size on outer surface of subanal plate between which are slender, curved hairs (Fig. 11G, H). Telson elongate, triangular, with rounded posterior apex in both sexes, bearing abundant setae and slender, curved hairs as on subanal plate (Fig. 12D, E). Male: typical scutigerid gonopod styles on first and second genital segment (Fig. 12A). Length of first gonopod and male second gonopods subequal. Both pairs of gonopods covered with setae and dense hairs. Subanal plate with parabolic outline, relatively shorter than in female, non-setose band along middle of subanal plate (Fig. 11F). Etymology. The species is dedicated to Dr Gregory D. Edgecombe, in recognition of his pioneering contributions to the taxonomy and phylogeny of Scutigeromorpha and for his generous provision of critical literature on this group. Genetic distance and phylogenetic relationships of two new species The genetic distances among the five Thereuopoda species based on COX1 ranged from 13% to 18% (Suppl. material 3). The lowest interspecific distance was observed between T. edgecombei sp. nov. and T. kaijiangensis sp. nov., 370 ZooKeys 1264: 351–376 (2025), DOI: 10.3897/zookeys.1264.165241 Jie-Hong Ji et al.: Two new species of Thereuopoda from China amounting to 13%. The phylogenetic analyses revealed an initial divergence between Pselliodidae and the remaining taxa, followed by a major split separating Scutigerinidae from Scutigeridae. Within Scutigeridae, further resolution indicated a primary division between Scutigerinae and Thereuoneminae, although monopoly of these two subfamilies was not supported. Within Thereuopoda, the ML and BI trees showed identical topologies (Fig. 14, Suppl. material 4): (((T. edgecombei sp. nov. + T. kaijiangensis sp. nov.) + T. clunifera) + (T. longicornis + T. sp.)). Discussion Two new species, Thereuopoda kaijiangensis sp. nov. and T. edgecombei sp. nov., can be distinguished from each other based on diagnostic characters listed in Table 5. Both of these new species can be easily distinguished from the morphologically similar T. longicornis by the relatively greater length of the female gonopod (2.25–2.6× in T. kaijiangensis, 2.03–2.37× in T. edgecombei versus 1.3× in T. longicornis) (Krishnan and Prasad 2022). Additionally, T. kaijiangensis and T. edgecombei exhibit a radiating network of dark-brown pigment on the posterolateral part of the head capsule, as opposed to dark greenish-brown patches in T. longicornis. There are three longitudinal stripes on central and Figure 14. Phylogenetic relationships of Scutigeromorpha were inferred from ML analyses based on six genes (18S rRNA, 28S rRNA, H3, 16S rRNA, 12S rRNA, and COX1). 371 ZooKeys 1264: 351–376 (2025), DOI: 10.3897/zookeys.1264.165241 Jie-Hong Ji et al.: Two new species of Thereuopoda from China lateral tergites in T. kaijiangensis and T. edgecombei, which are in contrast to a single brownish median longitudinal stripe in T. longicornis. Both ML and BI analyses recovered the monophyly of Thereuopoda, and the congruent topology, branch lengths, and the genetic distance provide robust evidence for the establishment of the two new species. Three poorly known taxa—T. (P. ) chinensis, Scutigera complanata Haase, 1887 and Scutigera sinuata Haase, 1887—require comparison with the two new species. Thereuopoda (P. ) chinensis was originally described based on a single female specimen collected from Macao, and its taxonomic validity remains uncertain (Verhoeff 1905; Würmli 1979). According to Verhoeff’s original description, T. (P. ) chinensis exhibits a greyish-yellow median longitudinal stripe (“Rücken nur mäßig gewölbt, grün, mit graugelblicher, auch über die Sättel ausgedehnter Mittelbinde”) and lacks spines on femora of legs 1–4 (“Femur des 1.–4. B. unbedornt”). The generic placement of S. complanata and S. sinuata remains uncertain, and both species lack longitudinal stripes on the tergites (“Ohne deutliche Längsstreifen auf den Schilden”) (Haase 1887). In contrast, the two new species possess three median longitudinal stripes and have spines on femora of all legs. They are clearly distinct from these three previously described taxa and are not considered conspecific with any of them. A major challenge in advancing the taxonomy and phylogeny of scutigeromorph centipedes is that numerous species were originally described based on limited material, with no additional data available since their initial descriptions (Butler et al. 2010). It is anticipated that this group will receive increased taxonomic and systematic attention in the future. Key to known Thereuopoda species in China 1 Posterior border of tergites evenly rounded ............................... T. clunifera – Posterior border of tergites unevenly rounded ............................................2 2 One median longitudinal stripe on tergites ............................. T. longicornis – Three longitudinal stripes on tergites ..........................................................3 3 Hairs absent on proarthron of female gonopod ......T. kaijiangensis sp. nov. – Hairs present on proarthron of female gonopod ......T. edgecombei sp. nov. Table 5. Comparison of Thereuopoda species from China. Characters T. kaijiangensis sp. nov.T. edgecombei sp. nov. T. longicornis T. clunifera Pigment in the posterolateral part of head capsule Radiating network Radiating network Patch – Stripes on tergites Three longitudinal stripes Three longitudinal stripes One median longitudinal stripe Cross–shaped stripe Posterior margin of tergites Unevenly rounded Unevenly rounded Unevenly rounded Evenly rounded Median embayment in posterior margin of T8 Present Absent – – Tibial spine-bristles on leg 1 0/2 0/1 – – Length of Leg 15 / body 2.85–3.14 1.79–1.94 – – A/B of female gonopod 2.25–2.6 2.03–2.37 1.3 – Hairs on proarthron of female gonopod Absent Present – – Shape of posterior termination of female subanal plate Pointed or with process Pointed or with process Pointed or with process Blunt and rounded Length of first male gonopod / second male gonopod 1.5 Subequal – – 372 ZooKeys 1264: 351–376 (2025), DOI: 10.3897/zookeys.1264.165241 Jie-Hong Ji et al.: Two new species of Thereuopoda from China Acknowledgements The authors gratefully acknowledge Dr Gregory D. Edgecombe and Dr Jörg Spelda for providing literature on Scutigeromorpha. We are indebted to ZooKeys editors and the reviewers who assisted us in improving the manuscript. Additional information Conflict of interest The authors have declared that no competing interests exist. Ethical statement No ethical statement was reported. Use of AI No use of AI was reported. Funding This work was supported by the Natural Science Foundation of Zhejiang Province (LY23C040002) and the Natural Science Foundation of China (32470475). Author contributions Conceptualization, D.-N.Y., J.-Y.Z., and S.-L.L.; methodology, J.-H.J., C.-Y.S., and H.-Y.W.; software, J.-H.J., Y.-X.G and H.-Y.W.; investigation, J.-H.J., H.-Y.W. and C.-Y.S.; data curation, J.-H.J. and C.-Y.S.; writing – original draft preparation, J.-H.J., C.-Y.S., and H.-Y.W.; writing – review and editing, J.-H.J., C.-Y.S., D.-N.Y., J.-Y.Z., and S.-L.L.; visualization, J.-H.J., Y.-X.G, C.-Y.S. and S.-L.L.; project administration, D.-N.Y., J.-Y.Z. and S.-L.L.; funding acquisition, J.-Y.Z. All authors have read and agreed to the published version of the manuscript. Author ORCIDs Jie-Hong Ji https://orcid.org/0009-0001-9381-0306 Hui-Yuan Wu https://orcid.org/0009-0007-6064-4819 Dan-Na Yu https://orcid.org/0000-0002-9870-1926 Jia-Yong Zhang https://orcid.org/0000-0002-7679-2548 Data availability All of the data that support the findings of this study are available in the main text or Supplementary Information. 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Molecular Ecology Resources 20(1): 348–355. https://doi.org/10.1111/1755-0998.13096 Supplementary material 1 List for each sequenced locus Authors: Jie-Hong Ji, Chen-Yang Shen, Yi-Xin Gao, Hui-Yuan Wu, Dan-Na Yu, Sheng-Long Liu, Jia-Yong Zhang Data type: pdf Explanation note: List of taxa, numbers, localities, and GenBank accession numbers for each sequenced locus. Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/zookeys.1264.165241.suppl1