Research Article |
Corresponding author: Xuankong Jiang ( antoma93@gmail.com ) Corresponding author: Chao Jiang ( jiangchao0411@126.com ) Academic editor: Michelle Hamer
© 2025 Yuan Xiong, Huiming Chen, Xuankong Jiang, Chao Jiang.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Xiong Y, Chen H, Jiang X, 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
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The millipede genus Kronopolites Attems, 1914 was originally described by monotypy with Strongylosoma swinhoei Pocock, 1895 as the type species, which was based on a single female specimen. Although this species has been believed to be widespread in China, there have been no confirmed reports of it from its type locality, leading to uncertainty about its taxonomic status. To address this issue, we newly sampled specimens from its type locality in Zhifu, Shandong Province, China. Our morphological analysis suggests that Kronopolites swinhoei (Pocock, 1895) should be reclassified as Nedyopus swinhoei (Pocock, 1895) comb. nov. and is a senior synonym of Nedyopus patrioticus Attems, 1898, syn. nov. The results also support the recovery of the name Kronopolites svenhedini (Verhoeff, 1934) sp. reval., which was previously misidentified as a junior synonym under K. swinhoei. The former is now the genus type of Kronopolites.
New combination, revalidation, revision, taxonomy, topotypes
To address this issue, millipedes closely matching Pocock’s description were collected on Zhifu Island and identified as topotypic K. swinhoei. Subsequent morphological studies indicate that Brölemann’s identification of K. swinhoei was incorrect. Consequently, it is determined that K. swinhoei belongs to the genus Nedyopus and is a senior synonym of Nedyopus patrioticus (Attems, 1898), which is widely distributed in East Asia. This removal of K. swinhoei from Kronopolites leads to the revalidation and reassignment of Kronopolites svenhedini (Verhoeff, 1934), instead, as the type species of the genus Kronopolites.
Specimens were collected by tweezers and preserved in 75% ethanol for morphological studies. Live animals were photographed with a Sony A7R4A camera with a Sony FE 90 mm macro lens. Specimens are deposited in National Resource Center for Chinese Materia Medica, China Academy of Chinese Medical Sciences, Beijing, China (
Specimens were examined, photographed, and measured using a Leica M205 MCA microscope equipped with a Leica DMC 6200 camera and LAS software v. 4.1 (Leica, Germany). Photos were converted into hand-drawn illustrations using SKETCHBOOK v. 6.0.6. Maps were generated with ArcMap v. 10.7.1 software (Figs
Family Paradoxosomatidae Daday, 1889
Subfamily Paradoxosomatinae Daday, 1889
Orthomorpha cingulata Attems, 1898, by original designation.
Stronglosoma Swinhoei Pocock, 1895: 354–355. Type specimen: holotype female, collected from Chee Foo (= Zhifu), Yantai, Shandong Province of China, deposited at the British Museum of Natural History (
Kronopolites swinhoei:
Strongylosoma patrioticum Attems, 1898: 300, figs 12, 13. Type locality: Japan. New synonymy.
Nedyopus patrioticus:
Differs from other species of the genus by the following combination of characters. The metaterga have strong contrasting colors, which are not circularly patterned as in other Nedyopus species, and the gonopod femur suddenly widens at the base, with l’ and l” not jagged.
China – Anhui Province: • 1 ♂, Fuyang, 30.I.2020, Yihao Ge leg. (
Length ca 17.5–25.1 mm (♂), 18.2–32.7 mm (♀) with 20 segments. Live color variable (Fig.
Nedyopus swinhoei comb. nov. A anterior part of body, dorsal view B anterior part of body, lateral view C segments 10 and 11, dorsal view D segments 9–11, lateral view E sternal cones between coxae 4, anterior view F–H posterior part of body, dorsal, lateral, and ventral view, respectively. Scale bar: 1 mm.
Clypeolabral region and vertex densely setose. Epicranial suture distinct. Width of body gradually expanded from head to 5th segment, approximately equal in width from 5th to 16th segments, and tapering from 16th to telson. Caudal corner of collum broadly rounded, declined ventrad, produced behind rear tergal margin (Fig.
Cuticle shining (Fig.
Gonopods (Fig.
China: Anhui (New record), Jiangsu, Jiangxi (New record), Shandong, Taiwan (
The specimens from Zhoushan Island were initially identified as K. swinhoei by Brölemann in 1896, without providing a justification. However, our investigation reveals a distinct divergence from the original description. For instance, the specimens from Zhoushan are notably larger (47 mm vs 35 mm) and have more vivid in color on the metazonites (orange-red vs yellow).
During our research in Zhifu, we found a species that closely matches Pocock’s description, leading us to confidently identify it as K. swinhoei. On examination of the topotypes, we observed significant differences in the gonopods compared to Brölemann’s illustrations (1896). These differences, including the femorite (strongly twisted and expanded in Nedyopus vs straight in Kronopolites), the postfemoral sulcus (missing in Nedyopus vs existed in Kronopolites) and the solenophore (lamelliform in Nedyopus vs tubuliform in Kronopolites), indicate that this species belongs to Nedyopus rather than Kronopolites, and is identical to the widespread species Nedyopus patrioticus (Attems, 1898). Consequently, K. swinhoei is formally transferred to Nedyopus, and Nedyopus patrioticus is considered a junior synonym of Nedyopus swinhoei (Pocock, 1895) comb. nov. Additionally, N. patrioticus consists of two subspecies Nedyopus patrioticus patrioticus (Attems, 1898) from Japan and Nedyopus patrioticus unicolor (Carl, 1902) from Indonesia. Therefore, the subspecies unicolor should be treated as Nedyopus swinhoei unicolor (Carl, 1902) comb. nov.
Nedyopus swinhoei (Pocock, 1895) comb. nov. has a wide distribution across Asia, from Indonesia to China, Korea, and Japan (
Kronopolites
Kronopolites:
Kansupus
Kansupus:
Parakansupus
Parakansupus:
Kronopolites svenhedini (Verhoeff, 1934) sp. reval., by present designation.
See
Stronglosoma Swinhoei:
Kansupus svenhedini
Verhoeff, 1934: 17, figs.4–8, synonymized by
Kronopolites swinhoei:
Kronopolites svenhedini:
Kansupus svenhedini var. dentiger
Verhoeff, 1934: 19, fig. 9;
China – • Gansu Province: 5 ♂♂ and 40 ♀♀ (20230922044, -45, -46, 20230922048–20230922051, 20230922054–20230922057, -60, -61, -63, -66, -67, -68, -70, -72, -73, -75, -76, 20230922078–20230922090, 20230922092–20230922098, -101, -102, -104), Lintao County, Fenghuangshan Forest Park (35.4009°N, 103.8901°E), 1960 m a.s.l., 22.IX.2023, Tianyun Chen, Jiabo Fan & Yiying Zhao leg., (
Differs from other species of the genus by the following combination of characters: metazonae have two shapes, either as a transverse band or a median oval spot, and also have two color variations, ranging from pale yellow to orange-red; paraterga relatively poorly developed, set lower (mostly at about 1/3 height of segments), caudal corners usually not surpassing rear tergal contours, at most narrowly rounded; ♂ sternal cones present; processes a and b of gonopod on a broad common stem, neither slender nor long (
Length ca 26.0–50.0 mm (♂), 27.0–60.0 mm (♀) with 20 segments. Live color variable (Fig.
Head densely setose. Antennae moderately long (Fig.
Collum of different specimens with one or two transverse rows of setae, one row with 1+1 anterior, two rows with 1+1 at both anterior and intermediate. Caudal corner of collum very broadly rounded, declined ventrad, produced behind rear tergal margin (Fig.
Cuticle shining, prozonae finely shagreened, metaterga finely rugulose (Fig.
Kronopolites svenhedini sp. reval A anterior part of body, dorsal view B anterior part of body, lateral view C segments 10 and 11, dorsal view D segments 9–11, lateral view E sternal cones between coxae 4, ventral view F–H posterior part of body, dorsal, lateral and ventral views, respectively. Scale bars: 1 mm.
Coxite of gonopods (Fig.
China: Chongqing, Qinghai, Gansu, Guizhou, Henan (new record), Shaanxi, Sichuan, Yunnan and Zhejiang (
Kronopolites svenhedini (Verhoeff, 1934) sp. reval. is widely distributed in China, with the westernmost occurrence in Qinghai, the easternmost in Zhejiang, the southernmost in Yunnan and the northernmost in Gansu. It shows variation in color, body size, and subtle differences in gonopod shape among different populations (
The authors have declared that no competing interests exist.
No ethical statement was reported.
This research was supported by grants from the CACMS Innovation Fund (nos. CI2023E002), the National Natural Science Foundation of China (32100365), the Doctoral Foundation of Guizhou Academy of Sciences (R [2021]1) and the Forestry and Grassland Ecological Protection and Restoration Fund of Guizhou Province (2022-2024) and Key project at central government level: The ability establishment of sustainable use for valuable Chinese medicine resources (nos. 2060302).
Yuan Xiong: Resources, Methodology, Software, Data Curation, Writing − Original Draft, Writing − Review & Editing. Huiming Chen: Resources, Supervision, Writing − Review & Editing, Project administration. Xuankong Jiang: Conceptualization, Resources, Methodology, Supervision, Writing − Review & Editing, Project administration, Funding acquisition. Chao Jiang: Conceptualization, Resources, Methodology, Supervision, Writing − Review & Editing, Project administration, Funding acquisition.
Yuan Xiong https://orcid.org/0009-0000-3812-9932
Huiming Chen https://orcid.org/0000-0002-2449-3036
Xuankong Jiang https://orcid.org/0000-0003-3506-5894
Chao Jiang https://orcid.org/0000-0003-1841-1169
All of the data that support the findings of this study are available in the main text.
We sincerely thank Paul Marek (Virginia Polytechnic Institute and State University, USA), Dr Cathy Car for reviewing the manuscript and offering critical comments. We appreciate the help of Dr Sergei I. Golovatch (Institute for Problems of Ecology and Evolution, Russian), Petra Sierwald, and Dr Peter Decker for providing important references and their generous help. We thank Dr Lu Tian (Shandong Jianzhu University, China), Shi Leilei (Henan University, China) and Mr Lu Chongwei (Guizhou Institute of Biology, China) for the assistance during fieldwork. We are grateful to Di Zhiyong (Hebei University, China), Dr Ge Yihao (Anhui Normal University, China) and Ms Fu Rong for providing the valuable specimens.