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Research Article
Three new species of Pseudopoda Jäger, 2000 (Araneae, Sparassidae, Heteropodinae) from Qizimeishan National Nature Reserve of Hubei, China
expand article infoJian Chang, He Zhang§, Jie Liu, Yang Zhu, Changyong Liu|, Kuai Chen|, Changhao Hu
‡ Hubei University, Wuhan, China
§ Xingtai University, Xingtai, China
| Qizimeishan National Nature Reserve Administration, Xuan’en, China
¶ Hubei Broad Nature Technology Service Co., Ltd., Wuhan, China
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Abstract

The Qizimeishan National Nature Reserve is situated in the southwestern region of Hubei Province, adjacent to the northeastern edge of the Yunnan-Guizhou Plateau. A survey of spiders of this reserve was conducted recently, leading to the discovery of three new species of the genus Pseudopoda Jäger, 2000: P. arcuata Zhang, J. Liu & Hu, sp. nov. (♀), P. qizimeishanensis Zhang, J. Liu & Hu, sp. nov. (♂, ♀) and P. weimiani Zhang, J. Liu & Hu, sp. nov. (♂, ♀). Diagnoses, descriptions, photos, and a distribution map are provided.

Key words

Biodiversity, high-altitude niche, huntsman spiders, morphology, taxonomy, Yunnan-Guizhou Plateau

Introduction

The Qizimeishan National Nature Reserve is located in Xuan’en County, southwest Hubei, with a total area of 345.5 km2 and the highest peak about 2010 m above sea level (Liu et al. 2006). It mainly protects the middle subtropical mountain evergreen broad-leaved forest and the subalpine sphagnum marshes wetland areas. The reserve is rich in wildlife resources and has been listed as a key area of biodiversity in the country by the China Biodiversity Conservation Action Plan (Xu et al. 2006).

The genus Pseudopoda Jäger, 2000 is the largest genus of the family Sparassidae Bertkau, 1872, with 256 known species (World Spider Catalog 2024). Currently, 155 Pseudopoda species are known in China and five in Hubei (Quan et al. 2014; Zhang et al. 2023; Gong et al. 2023; Wen et al. 2024). Recently, a series of taxonomic works on the genus was published: Wen et al. (2024) described one new species from Hubei; Wu et al. (2024) described three new species from China, Laos, and Thailand; Gong et al. (2023) described a new species from Hubei; Deng et al. (2023) described four new species from Chongqing; and Zhang et al. (2023) described 99 new species from East, South and Southeast Asia. Pseudopoda species are small to large spiders, living primarily in the leaf litter and less commonly on vegetation (Jäger et al. 2015).

A survey of spiders in the Qizimeishan National Nature Reserve carried out by colleagues of the Hubei University from June to July 2023 yielded three new species of Pseudopoda, which are described herein.

Material and methods

Specimens were examined using an Olympus SZX7 stereo microscope. Photographs were taken with a Leica M205C stereo microscope, and final multifocal images were produced with Helicon Focus (version 7.7.0). The male palps were examined and photographed after dissection. The epigynes were examined after being dissected from the spider’s body. All morphological measurements were calculated using a Leica M205C stereo microscope. Eye diameters were taken at the widest point. Legs and palps measurements were given as total length (femur, patella, tibia, metatarsus [absent in palp], tarsus). The terminologies used in text and figure legends follow Quan et al. (2014). Spination follows that given in Davies (1994). All measurements were in millimetres (mm). The specimens examined in this study were deposited in the Centre for Behavioural Ecology and Evolution (CBEE), College of Life Sciences, Hubei University in Wuhan.

Abbreviations in text and figures: AB, anterior bands; ALE, anterior lateral eyes; AME, anterior median eyes; C, conductor; CH, clypeus height; CO, copulatory opening; dRTA, dorsal retrolateral tibial apophysis; DS, dorsal shield of prosoma; E, embolus; EF, epigynal field; EP, embolic projection; FD, fertilization duct; Fe, femur; FW, first winding; LL, lateral lobes; Mt, metatarsus; OS, opisthosoma; Pa, patella; PLE, posterior lateral eyes; PME, posterior median eyes; Pp, palp; RTA, retrolateral tibial apophysis; S, spermathecae; Sp, spermophor; ST, subtegulum; T, tegulum; Ti, tibia; vRTA, ventral retrolateral tibial apophysis; I, II, III, IV, legs I to IV.

Result

Taxonomy

Family Sparassidae Bertkau, 1872

Subfamily Heteropodinae Thorell, 1873

Pseudopoda Jäger, 2000

Type species

Pseudopoda prompta (O. Pickard-Cambridge, 1885).

Diagnosis

Male Pseudopoda species can be diagnosed by the following combination of characters: 1) embolus at least in its basal part broadened and flattened; 2) conductor membranous (absent in some species); and 3) retrolateral tibial apophysis arising proximally or mesially from the tibia. Females can be diagnosed by the following combination of characters: 1) epigyne with lateral lobes extending distinctly beyond epigastric furrow, and covering median septum in most species; 2) first winding membranous, and with bent margins in most species; and 3) first winding or first winding and lateral lobes covering the internal duct system (Jäger, 2000; Jiang et al. 2018; Zhang et al. 2023; Wu et al. 2024).

Distribution

East, South and Southeast Asia.

Pseudopoda arcuata Zhang, J. Liu & Hu, sp. nov.

Figs 2, 3, 10

Type material

Holotype • female: China, Hubei Province: Enshi Tujia and Miao Autonomous Prefecture, Xuan’en County, Qizimeishan National Nature Reserve, Chunmuying Town, Shaiping Village; 29°57'47.52"N, 109°45'20.60"E; elev. 1822 m; 31 July 2023; Changhao Hu & Mian Wei leg. (CBEE, QZMS00602). Paratype • 1 female, Enshi Tujia and Miao Autonomous Prefecture, Xuan’en County, Qizimeishan National Nature Reserve, Chunmuying Town, Huoshaobao; 30°1'27.03"N, 109°45'23.23"E; elev. 1919 m; 13 July 2023; Changhao Hu & Mian Wei leg. (CBEE, QZMS00582).

Etymology

The specific epithet is a Latin word meaning “arc-shaped”, referring to the arc-shaped LL; adjective.

Diagnosis

The female of P. arcuata Zhang, J. Liu & Hu, sp. nov. resembles that of P. allantoides Zhang, Jäger & Liu, 2023 (cf. fig. 2A–C vs. fig. 8A–C in Zhang et al. 2023) by having curved anterior and posterior margins of LL, but can be recognised by: 1) S extending horizontally; and 2) FD arising anteriorly from S (vs. S extending longitudinally, FD arising medially from S in P. allantoides).

Female: Measurements: Small-sized. Body length 6.0, DS length 3.0, width 2.8; OS length 2.9, width 2.0. Eyes: AME 0.13, ALE 0.19, PME 0.22, PLE 0.23, AMEAME 0.11, AMEALE 0.07, PMEPME 0.21, PMEPLE 0.28, AMEPME 0.19, ALEPLE 0.26, CH AME 0.23, CH ALE 0.19. Spination: Pp 131, 101, 2121, 1014; Fe I–III 323, IV 321; Pa I–IV 000; Ti I–II 2221, III–IV 2126; Mt I–II 2024, III–IV 2026. Measurements of palp and legs: Pp 3.4 (1.1, 0.8, 0.4, –, 1.1), I 9.3 (2.5, 1.1, 2.6, 2.2, 0.9), II 12.1 (3.2, 1.4, 3.6, 2.8, 1.1), III 7.3 (2.0, 0.7, 2.1, 1.7, 0.8), IV 9.2 (2.9, 0.9, 2.2, 2.3, 0.9). Leg formula: II-I-IV-III. Promargin of chelicerae with three teeth, retromargin with four teeth, cheliceral furrow with c. 28 denticles.

Epigyne (Fig. 2A–C): As in diagnosis. EF 2 times wider than long, without obvious AB. Anterior and posterior margins of LL almost parallel and strongly curved. FW covering the whole S. S with enlarged terminal. FD narrow and long, laterad.

Colouration (Fig. 3A, B): DS yellow, with black marks. Fovea black. Legs with black spots. Dorsal OS brown, with black spots, ventral OS dark yellow, with several black spots, with a brown patch in front of spinnerets.

Male: Unknown.

Distribution

China (Hubei Province) (Fig. 10).

Pseudopoda qizimeishanensis Zhang, J. Liu & Hu, sp. nov.

Figs 1, 4, 5, 6, 10

Type material

Holotype • male: China, Hubei Province: Enshi Tujia and Miao Autonomous Prefecture, Xuan’en County, Changtanhe Dong Autonomous Township, Qizimeishan National Nature Reserve, Qizimeishan Mountain; 30°1'45.19"N, 109°43'45.42"E; elev. 1270 m; 6–11 July 2023; Changhao Hu & Mian Wei leg. (CBEE, QZMS00902). Paratypes • 8 males and 10 females, with same data as for holotype (CBEE, QZMS02441–QZMS02458).

Figure 1. 

Pseudopoda qizimeishanensis Zhang, J. Liu & Hu, sp. nov. (photos by Mian Wei) A male B female.

Figure 2. 

Pseudopoda arcuata Zhang, J. Liu & Hu, sp. nov., female A epigyne, ventral B vulva, dorsal C vulva, dorsal; red line represents schematic course of internal duct system. Abbreviations: CO, copulatory opening; FD, fertilization duct; FW, first winding; LL, lateral lobes; S, spermathecae. Scale bars: 0.2 mm.

Figure 3. 

Pseudopoda arcuata Zhang, J. Liu & Hu, sp. nov., female habitus (A dorsal B ventral). Scale bars: 2 mm.

Etymology

The specific epithet is derived from the type locality, the Qizimeishan Mountain; adjective.

Diagnosis

The male of P. qizimeishanensis Zhang, J. Liu & Hu, sp. nov. resembles that of P. baoshanensis Zhang, Jäger & Liu, 2023 (cf. fig. 4A–C vs. fig. 26A–C in Zhang et al. 2023) by having the expanded E, but can be recognised by: 1) RTA long, arising basally from Ti; 2) T without prolaterad outgrowth; and 3) tip of E pointing 11 o’clock (vs. RTA short, arising medially from Ti, T with prolaterad outgrowth, tip of E pointing 7 o’clock in P. baoshanensis). The female of P. qizimeishanensis Zhang, J. Liu & Hu, sp. nov. resembles that of P. nanyueensis Tang & Yin, 2000 (cf. fig. 5A–C vs. figs 2, 3 in Tang and Yin 2000) by: 1) anterior margins of LL V-shaped; 2) anterior margins of LL parallel to posterior margins of LL, but can be recognised by: S long, with wrinkles, almost parallel to anterior margins of LL (vs. S without wrinkles, extending horizontal in P. nanyueensis).

Male: Measurements: Medium-sized. Body length 14.9, DS length 7.7, width 6.6; OS length 6.8, width 4.8. Eyes: AME 0.29, ALE 0.42, PME 0.25, PLE 0.39, AMEAME 0.26, AMEALE 0.17, PMEPME 0.46, PMEPLE 0.59, AMEPME 0.37, ALEPLE 0.39, CH AME 0.72, CH ALE 0.66. Spination: Pp 131, 101, 2111; Fe I–II 323, III 322, IV 321; Pa I–III 101, IV 000; Ti I–II 2226, III–IV 2126; Mt I–II 2024, III 3025, IV 3036. Measurements of palp and legs: Pp 10.1 (3.3, 1.7, 1.9, –, 3.2), I 33.2 (9.3, 3.0, 9.0, 8.9, 3.0), II 34.8 (9.9, 2.8, 9.5, 9.5, 3.1), III 26.6 (7.7, 2.2, 7.3, 7.0, 2.4), IV 29.5 (8.5, 2.2, 7.7, 8.6, 2.5). Leg formula: II-I-IV-III. Promargin of chelicerae with three teeth, retromargin with four teeth, cheliceral furrow with c. 26 denticles.

Figure 4. 

Pseudopoda qizimeishanensis Zhang, J. Liu & Hu, sp. nov., left male palp (A prolateral B ventral C retrolateral). Abbreviations: C, conductor; dRTA, dorsal retrolateral tibial apophysis; E, embolus; Sp, spermophor; ST, subtegulum; T, tegulum; vRTA, ventral retrolateral tibial apophysis. Scale bars: 0.5 mm.

Palp (Fig. 4A–C): As in diagnosis. C membranous, arising from T at 11 o’clock position. E expanded and plate-like, arising from T at 9 o’clock position; embolic tip curved. RTA arising basally from Ti; vRTA triangular; dRTA long, with two thin teeth in retrolateral view.

Colouration (Fig. 6A, B): DS yellow, with black spots. Ventral legs with black spots. Dorsal OS brown, ventral OS with black spots, spinnerets yellow, with two parallel longitudinal lines of lighter dots.

Female: Measurements: Medium-sized. Body length 16.9, DS length 7.9, width 7.1; OS length 8.6, width 6.6. Eyes: AME 0.33, ALE 0.51, PME 0.31, PLE 0.42, AMEAME 0.32, AMEALE 0.27, PMEPME 0.58, PMEPLE 0.64, AMEPME 0.44, ALEPLE 0.41, CH AME 0.60, CH ALE 0.58. Spination: Pp 131, 101, 2121, 1014; Fe I–II 323, III 322, IV 321; Pa I–IV 101; Ti I–IV 2026; Mt I 1014, II–III 2024, IV 3025. Measurements of palp and legs: Pp 9.2 (2.8, 1.1, 2.1, –, 3.2), I 25.6 (7.0, 2.9, 7.0, 6.4, 2.3), II 26.9 (7.9, 2.8, 7.2, 6.8, 2.2), III 22.3 (7.0, 2.7, 5.4, 5.2, 2.0), IV 24.6 (7.6, 2.3, 5.9, 6.5, 2.3). Leg formula: II-I-IV-III. Promargin of chelicerae with three teeth, retromargin with four teeth, cheliceral furrow with c. 26 denticles.

Figure 5. 

Pseudopoda qizimeishanensis Zhang, J. Liu & Hu, sp. nov., female A epigyne, ventral B vulva, dorsal C vulva, dorsal; red line represents schematic course of internal duct system. Abbreviations: AB, anterior bands; CO, copulatory opening; FD, fertilization duct; FW, first winding; LL, lateral lobes; S, spermathecae. Scale bars: 0.5 mm.

Figure 6. 

Pseudopoda qizimeishanensis Zhang, J. Liu & Hu, sp. nov. A, B male habitus (A dorsal B ventral) C, D female habitus (C dorsal D ventral). Scale bars: 2 mm.

Epigyne (Fig. 5A–C): As in diagnosis. EF as wide as long, with obvious AB. Anterior margins of LL V-shaped. S long and anterolaterally pointed, with wrinkles. FW covering entire S. FD narrow.

Colouration (Fig. 6C–D): As in males, but darker and with a transverse white patch in posterior part of dorsal OS.

Distribution

China (Hubei Province) (Fig. 10).

Pseudopoda weimiani Zhang, J. Liu & Hu, sp. nov.

Figs 7, 8, 9, 10

Type material

Holotype • male: China, Hubei Province: Enshi Tujia and Miao Autonomous Prefecture, Xuan’en County, Qizimeishan National Nature Reserve, Changtanhe Dong Autonomous Town Qizimeishan Mountain; 30°1'45.19"N, 109°43'45.42"E; elev. 1270 m; 6–11 July 2023; Changhao Hu & Mian Wei leg. (CBEE, QZMS00732). Paratypes • 2 males and 11 females, with same data as for holotype (CBEE, QZMS02459–QZMS02466, QZMS02779, QZMS04305–QZMS04308) • 1 male, Enshi Tujia and Miao Autonomous Prefecture, Xuan’en County, Qizimeishan National Nature Reserve, Changtanhe Dong Autonomous Town, Liangxihe Village; 29°58'46.33"N, 109°42'11.27"E; elev. 827 m; 29 June 2023; Changhao Hu & Mian Wei leg. (CBEE, QZMS00632) • 1 female, Enshi Tujia and Miao Autonomous Prefecture, Xuan’en County, Qizimeishan National Nature Reserve, Shadaogou Town, Longtan Village; 29°41'42.96"N, 109°39'46.03"E; elev. 637 m; 17 July 2023; Changhao Hu & Mian Wei leg. (CBEE, QZMS00622) • 1 female, Enshi Tujia and Miao Autonomous Prefecture, Xuan’en County, Qizimeishan National Nature Reserve, Shadaogou Town, Baishuihe Village; 29°55'25.78"N, 109°44'9.49"E; elev. 843 m; 23–24 July 2023; Changhao Hu & Mian Wei leg. (CBEE, QZMS03877).

Etymology

This species is named after one of the collectors: Mian Wei; noun in genitive case.

Diagnosis

The male of P. weimiani Zhang, J. Liu & Hu, sp. nov. resembles that of P. hongqi Deng, Zhong, Irfan & Wang, 2023 (cf. fig. 7A–C vs. figs 5–10 in Deng et al. 2023) by: 1) RTA unbranched; 2) E wide; and 3) EP distinct, but can be recognised by: 1) RTA short and smooth; and 2) EP twisted (vs. RTA long and gradually narrowing toward the tip, EP without a twist, terminal not exceeding E in P. hongqi). The female of P. weimiani Zhang, J. Liu & Hu, sp. nov. resembles that of P. taipingensis Zhang, Jäger & Liu, 2023 (cf. fig. 8A–C vs. fig. 237A–C in Zhang et al. 2023) by: 1) anterior margins of LL almost straight; and 2) S “八”-shaped, but can be recognised by: 1) posterior margins of LL almost straight and parallel to anterior margins; and 2) S with an obvious turning in ventral view (vs. posterior margins of LL W-shaped, S with distinct tube-like structures in P. taipingensis).

Male: Measurements: Small-sized. Body length 7.1, DS length 3.6, width 3.2; OS length 3.3, width 2.1. Eyes: AME 0.15, ALE 0.24, PME 0.17, PLE 0.26, AMEAME 0.13, AMEALE 0.07, PMEPME 0.22, PMEPLE 0.28, AMEPME 0.24, ALEPLE 0.22, CH AME 0.34, CH ALE 0.32. Spination: Pp 131, 101, 2111; Fe I–II 323, III–IV 322; Pa I–III 101, IV 100; Ti I–II 2228, III–IV 2126; Mt I–II 2024, III 3025, IV 3036. Measurements of palp and legs: Pp 5.1 (1.7, 0.6, 1.1, –, 1.7), I 18.8 (4.9, 1.4, 5.9, 4.8, 1.8), II 19.8 (5.5, 1.5, 6.2, 5.1, 1.5), III 14.6 (4.3, 1.0, 4.4, 3.6, 1.3), IV 17.5 (5.0, 1.1, 4.8, 4.8, 1.8). Leg formula: II-I-IV-III. Promargin of chelicerae with three teeth, retromargin with four teeth, cheliceral furrow with c. 26 denticles.

Figure 7. 

Pseudopoda weimiani Zhang, J. Liu & Hu, sp. nov., left male palp (A prolateral B ventral C retrolateral). Abbreviations: C, conductor; E, embolus; EP, embolic projection; RTA, retrolateral tibial apophysis; Sp, spermophor; ST, subtegulum; T, tegulum. Scale bars: 0.5 mm.

Palp (Fig. 7A–C): As in diagnosis. C membranous, arising from 12 o’clock position of T. E wide, arising from 9 o’clock position of T; EP plate-like and twisted; embolic tip with a small sharp tooth. RTA short, with smooth margin, arising medially from Ti.

Figure 8. 

Pseudopoda weimiani Zhang, J. Liu & Hu, sp. nov., female A epigyne, ventral B vulva, dorsal C vulva, dorsal; red line represents schematic course of internal duct system. Abbreviations: AB, anterior bands; CO, copulatory opening; FD, fertilization duct; FW, first winding; LL, lateral lobes; S, spermathecae. Scale bars: 0.2 mm.

Figure 9. 

Pseudopoda weimiani Zhang, J. Liu & Hu, sp. nov. A, B male habitus (A dorsal B ventral) C, D female habitus (C dorsal D ventral). Scale bars: 2 mm.

Figure 10. 

Distribution map of the three new species of Pseudopoda. Black circles: P. arcuata Zhang, J. Liu & Hu, sp. nov.; green square: P. qizimeishanensis Zhang, J. Liu & Hu, sp. nov.; red triangles: P. weimiani Zhang, J. Liu & Hu, sp. nov.

Colouration (Fig. 9A, B): DS yellow, with black marked and black margin. Fovea black. Legs with black spots. OS light brown, with several reddish-brown marks, regularly arranged.

Female: Measurements: Small-sized. Body length 8.2, DS length 3.6, width 3.4; OS length 4.5, width 3.9. Eyes: AME 0.15, ALE 0.18, PME 0.23, PLE 0.25, AMEAME 0.15, AMEALE 0.11, PMEPME 0.22, PMEPLE 0.31, AMEPME 0.25, ALEPLE 0.30, CH AME 0.32, CH ALE 0.26. Spination: Pp 131, 101, 2121, 1014; Fe I–II 323, III–IV 322; Pa I–III 101, IV 100; Ti I–II 2228, IV 2126; Mt I–II 2024, III 3025, IV 3036. Measurements of palp and legs: Pp 4.8 (1.5, 0.7, 1.0, –, 1.6), I 12.0 (3.2, 1.2, 3.4, 3.0, 1.2), II 13.9 (3.9, 1.0, 4.1, 3.6, 1.3), III 10.5 (3.2, 0.9, 2.8, 2.5, 1.1), IV 12.3 (3.8, 0.9, 3.1, 3.3, 1.2). Leg formula: II-IV-I-III. Promargin of chelicerae with three teeth, retromargin with four teeth, cheliceral furrow with c. 26 denticles.

Epigyne (Fig. 8A–C): As in diagnosis. EF wider than long, with indistinct AB. LL much wider than long, anterior margins of LL almost straight, posterior margins almost parallel to anterior margins of LL. Terminal of S twisted, posterolaterally pointed. FW covering anterior part of S. FD long and narrow.

Colouration (Fig. 9C, D): As in males, but with some bright dots in anterior part of dorsal OS.

Distribution

China (Hubei Province) (Fig. 10).

Discussion

The genus Pseudopoda is widely distributed across China, with 155 species accounting for approximately 61% of the world’s total. The majority of these species are found in the southwestern regions, particularly on the Yunnan-Guizhou Plateau, which harbors 84 species – more than 54% of the Chinese species – highlighting the region’s rich biodiversity and varied topography (Jäger 2001; Jäger and Vedel 2007; Yang et al. 2009; Zhang et al. 2013, 2017; Jiang et al. 2018; Zhang et al. 2019; Yang et al. 2022; Zhang et al. 2023). The distribution of Pseudopoda spiders in China is closely linked to the country’s mountainous areas, where species have adapted to specific altitudinal niches. The Qizimeishan National Nature Reserve, part of the northeastern extension of the Yunnan-Guizhou Plateau, features a diverse landscape with elevations ranging from 650 m to over 2010 m (Liu et al. 2006). This variation in altitude creates a variety of microclimates and habitats that support a wide array of species. However, research on the spiders of this area remains limited.

Furthermore, Pseudopoda is particularly diverse in high-altitude regions (Jäger, 2001, 2015; Zhang et al. 2023), with approximately 79% of the world’s species found at elevations above 1000 m. In the current study, we report three new Pseudopoda species, all of which were collected at elevations above 1000 m, except for P. weimiani Zhang, J. Liu & Hu, sp. nov., which was also found at elevations between 600 and 900 m. Finally, the limited dispersal ability of these spiders, likely due to the absence of ballooning behavior, results in small, localized populations, making these species highly susceptible to habitat changes (Bell et al. 2005; Zhang et al. 2021). Consequently, conservation efforts and taxonomic studies should prioritize the Qizimeishan National Nature Reserve to protect its unique and valuable spider fauna.

Acknowledgements

We thank Mian Wei (CBEE) for helping collecting specimens and taking photos. The manuscript benefitted greatly from comments by Francesco Ballarin, Danniella Sherwood, and Alireza Zamani.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This research was funded by the National Natural Sciences Foundation of China (NSFC-32300378), the Foundation of Hubei Key Laboratory of Regional Development and Environmental Response (2022(B)004), the China Postdoctoral Science Foundation (2023M731035), the Qizimeishan National Nature Reserve, Hubei, China (ESTXZB2023-062) and the Science & Technology Fundamental Resources Investigation Program of China (2023FY100200).

Author contributions

Writing – original draft: JC, CH. Writing – review and editing: HZ, JL, YZ, CL, KC, CH.

Author ORCIDs

Jian Chang https://orcid.org/0009-0001-9840-7348

He Zhang https://orcid.org/0000-0002-1478-9837

Jie Liu https://orcid.org/0000-0001-7744-9744

Yang Zhu https://orcid.org/0009-0002-2934-6809

Changhao Hu https://orcid.org/0009-0007-5591-3121

Data availability

All of the data that support the findings of this study are available in the main text.

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