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Research Article
Three new species of the genus Olios Walckenaer, 1837 (Araneae, Sparassidae) from southern China
expand article infoHe Zhang‡§, Weitao Niu‡§, Jie Liu|, Caijun Wang‡, Changhao Hu§‡|
‡ Xingtai University, Xingtai, China
§ Xingtai Innovation Center for Sweet Potato Germplasm and Rapid Propagation Technology, Xingtai, China
| Hubei University, Wuhan, China
¶ Hubei Broad Nature Technology Service Co., Ltd., Wuhan, China

Corresponding author: Changhao Hu ( changhaohu1998@gmail.com )

Academic editor: Cristina Rheims
© 2025 He Zhang, Weitao Niu, Jie Liu, Caijun Wang, Changhao Hu.
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: Zhang H, Niu W, Liu J, Wang C, Hu C (2025) Three new species of the genus Olios Walckenaer, 1837 (Araneae, Sparassidae) from southern China. ZooKeys 1245: 343-356. https://doi.org/10.3897/zookeys.1245.144552
ZooBank: urn:lsid:zoobank.org:pub:7A73A97F-AE84-4A3F-B5EC-9A6C764B177F
Open Access

Abstract

Three new species of the sparassid spider genus Olios Walckenaer, 1837 are described from China: O. biprocessus Hu, Zhang & Liu, sp. nov. (♂) from Hunan Province, O. uniprocessus Hu, Zhang & Liu, sp. nov. (♂) from Chongqing Municipality and O. lincangensis Hu, Zhang & Liu, sp. nov. (♂♀) from Yunnan Province. Diagnoses, descriptions, photos and a distribution map of these three new species are provided.

Key words:

Biodiversity, huntsman spiders, morphology, new division, sparassids, taxonomy

Introduction

The genus Olios Walckenaer, 1837 is the third-largest genus in the spider family Sparassidae Bertkau, 1872, with 166 known species (World Spider Catalog 2025). For a long time, this genus was used as a “dumping ground” for the family Sparassidae (Jäger and Kunz 2005). More than 60 species previously assigned to Olios have been transferred to other genera in past studies, particularly among Neotropical species (e.g., Caayguara Rheims, 2010, Curicaberis Rheims, 2015, Meri Rheims & Jäger, 2022, Nolavia Kammerer, 2006, and Sadala Simon, 1880) (Rheims 2010, 2015; Jäger 2020; Rheims and Jäger 2022). Jäger (2020) further revised Olios and reported eight Olios species-groups (argelasius-group, auricomis-group, coenobitus-group, correvoni-group, hirtus-group, nentwigi-group, rossettii-group, and stimulator-group), comprising 71 species. However, many species remain improperly assigned to Olios, necessitating further systematic revision (Jäger 2020).

Phylogenetic analyses of Sparassidae have consistently demonstrated that Olios is a polyphyletic assemblage (Jäger and Kunz 2005; Moradmand et al. 2014; Tong et al. 2018; Gorneau et al. 2022), which may explain the remarkable morphological diversity of copulatory organs in Olios species. Recent phylogenetic work by Gorneau et al. (2022) delimited the scope of true Olios, which is a monophyletic clade represented by four species: O. argelasius (Walckenaer, 1806) (type species) from the Mediterranean, O. rossettii (Leardi, 1901) from southern Asia, O. lamarcki (Latreille, 1806) from Sri Lanka, and an unidentified species from Singapore.

In China, seven Olios species are currently known, which are classified into three species-groups: O. menghaiensis (Wang & Zhang, 1990) within the hirtus-group; O. digitatus Sun, Li & Zhang, 2011, O. nanningensis (Hu & Ru, 1988), O. scalptor Jäger & Ono, 2001 and O. suung Jäger, 2012 within the nentwigi-group; and O. tiantongensis (Zhang & Kim, 1996), O. sericeus (Kroneberg, 1875) within the rossettii-group (Kroneberg 1875; Hu and Ru 1988; Wang and Zhang 1990; Zhang and Kim 1996; Jäger and Ono 2001; Jäger and Yin 2001; Sun et al. 2011; Jäger 2012, 2020; Hu et al. 2025). The aim of the current paper is to contribute to the taxonomic knowledge of Olios in China by describing three new species.

Material and methods

The specimens examined in this study are deposited in the Centre for Behavioral Ecology and Evolution (CBEE), College of Life Sciences, Hubei University in Wuhan. Specimens were examined using an Olympus SZX7 stereo microscope. Photographs were taken on a Leica M205C stereo microscope. The male palp was examined and photographed after dissection. The epigyne was examined after being dissected from the body and treated in a warm 0.1 mg/ml Protease K solution. Eye diameters were taken at the widest point. Legs and palpal measurements were given as total length (femur, patella, tibia, metatarsus [absent in palp], tarsus). All measurements were in millimeters (mm). Spination follows that given in Davies (1994).

The terminology used in this paper follows Jäger (2020). Abbreviations: ALE = anterior lateral eyes, AME = anterior median eyes, C = conductor, CD = copulatory duct, CH = clypeus height, CO = copulatory opening, dRTA = dorsal part of retrolateral tibial apophysis, DS = dorsal shield of prosoma, E = embolus, ES = epigynal slit, FD = fertilization duct, Fe = femur, LL = lateral lobes, Mt = metatarsus, OS = opisthosoma, Pa = patella, PLE = posterior lateral eyes, PME = posterior median eyes, Pp = palp, ptA = proximal tibial apophysis, RTA = retrolateral tibial apophysis, S = spermathecae, TA = tegular apophysis, TAP I–IV = projection I to IV of tegular apophysis, Ti = tibia, vRTA = ventral part of retrolateral tibial apophysis, I, II, III, IV = legs I to IV.

Result

Taxonomy

Family Sparassidae Bertkau, 1872

Olios Walckenaer, 1837

Type species.

Olios argelasius (Walckenaer, 1806).

Diagnosis.

See Jäger (2020).

Olios rossettii -group

Diagnosis. See Jäger (2020).

Species included. Olios baulnyi (Simon, 1874), O. bhattacharjeei (Saha & Raychaudhuri, 2007), O. biprocessus Hu, Zhang & Liu, sp. nov., O. brachycephalus Lawrence, 1938, O. floweri Lessert, 1921, O. jaldaparaensis Saha & Raychaudhuri, 2007, O. japonicus Jäger & Ono, 2000, O. kiranae Sethi & Tikader, 1988, O. kolosvaryi (Caporiacco, 1947), O. longipes (Simon, 1884), O. lutescens (Thorell, 1894), O. mahabangkawitus Barrion & Litsinger, 1995, O. obesulus (Pocock, 1901), O. rossettii (Leardi, 1901), O. rotundiceps (Pocock, 1901), O. sericeus (Kroneberg, 1875), O. sherwoodi Lessert, 1929, O. suavis (O. Pickard-Cambridge, 1876), O. tarandus (Simon, 1897), O. tener (Thorell, 1891), O. tiantongensis (Zhang & Kim, 1996), O. uniprocessus Hu, Zhang & Liu, sp. nov.

Remarks. This species-group is characterized by the presence of one or two apophyses on palpal tibia, a variable tegular apophysis, and a small U-shaped embolus arising from the tegulum at the central to sub-central position (Jäger 2020). The two newly described species, Olios biprocessus Hu, Zhang & Liu, sp. nov. and O. uniprocessus Hu, Zhang & Liu, sp. nov., exhibit all the diagnostic characteristics of this species-group. Therefore, they clearly belong to this species-group.

Olios biprocessus Hu, Zhang & Liu, sp. nov.

https://zoobank.org/6525BA3E-BE51-4564-B1DD-5EA95D1F256A
Figs 1, 2, 8

Type material.

Holotype male: China • Hunan Province: Changsha City, Yuelu District, Xihu Park, under bark, 28.20758°N, 112.94004°E, elevation 30 m, 19 December 2023 as subadult, molted to adult on 18 February 2024, Jiwei Lin leg.

Etymology.

The specific name is a combination of the Latin bi (meaning double) and processus (meaning process, projection), referring to the two projections (TAP III, TAP IV) on the embolic base; noun.

Diagnosis.

The male of Olios biprocessus Hu, Zhang & Liu, sp. nov. resembles that of O. rossettii (Leardi, 1901) (cf. Fig. 1A–C vs. figs 146, 147 in Jäger 2020) by having two tibial apophyses, and a small U-shaped embolus arising sub-centrally from the tegulum, but can be recognized by: 1) retrolateral tibial apophysis leaf-shaped in retrolateral view, with a serrated retrolateral margin; and 2) tegular apophysis complex, with a sharp projection (TAP I) prolaterally, and two sharp projections (TAP III, TAP IV) on the embolic base (vs. retrolateral tibial apophysis triangular, tegular apophysis long triangular in O. rossettii).

Figure 1. 

Olios biprocessus Hu, Zhang & Liu, sp. nov., left male palp. A. Prolateral; B. Ventral; C. Retrolateral. Abbreviations: C = conductor, E = embolus, ptA = proximal tibial apophysis, RTA = retrolateral tibial apophysis, TA = tegular apophysis, TAP I–IV = projection I to IV of tegular apophysis. Scale bars: 0.5 mm.

The male of Olios biprocessus Hu, Zhang & Liu, sp. nov. also resembles that of O. uniprocessus Hu, Zhang & Liu, sp. nov. (cf. Fig. 1A–C vs. fig. 3A–C) by having a horn-shaped proximal tibial apophysis, a leaf-shaped retrolateral tibial apophysis with a serrated retrolateral margin, sharp TAP I, lamellar TAP II, and a small U-shaped embolus arising sub-centrally from the tegulum, but can be recognized by the tegular apophysis with two sharp projections (TAP III, TAP IV) on the embolic base (vs. one projection (TAP III) in O. uniprocessus Hu, Zhang & Liu, sp. nov.).

Description.

Male (holotype): Total length 8.5; DS 4.0 long, 4.1 wide; OS 4.6 long, 3.2 wide. Eyes: AME 0.24; ALE 0.19; PME 0.19; PLE 0.20; AME–AME 0.17; AME–ALE 0.17; PME–PME 0.42; PME–PLE 0.31; AME–PME 0.15; ALE–PLE 0.13; CH AME 0.18; CH ALE 0.25. Measurements of palp and legs: Pp: 5.3 (1.5, 0.6, 1.1, -, 2.1); I: 19.8 (5.4, 1.6, 5.2, 6.0, 1.6); II: 20.3 (5.4, 1.9, 6.6, 3.1, 3.3); III: 14.9 (4.7, 1.3, 3.6, 3.6, 1.7); IV: 17.3 (5.3, 1.2, 4.6, 4.9, 1.3). Leg formula: II-I-IV-III. Spination: Pp: 131, 000, 2000; legs: Fe I–II 223, III 323, IV 321; Pa I–IV 000; Ti I 2124, II 2024, III 2124, IV 1204; Mt I–IV 2024. Cheliceral furrow with 2 anterior and 5 posterior teeth.

Palp (Fig. 1A–C): As in diagnosis, ptA arising submedially from Ti, RTA arising distally from Ti. Cymbium almost 1.5 times longer than Ti. TA complicated, with a sharp projection (TAP I) on prolateral margin, a lamellar projection (TAP II) on dorsal TA, and two sharp projections (TAP III, TAP IV) on embolic base. C tiny. E arising sub-centrally from tegulum in 9-o’clock-position.

Colouration (Fig. 2A, B): DS pale yellow, with brown patterns. Sternum light yellow. Legs yellow to light brown, with brown spots. OS dorsally yellow, with yellowish-brown patterns. OS ventrally brownish purple, with four yellow longitudinal lines; anterior part yellow.

Figure 2. 

Olios biprocessus Hu, Zhang & Liu, sp. nov., male habitus. A. Dorsal; B. Ventral. Scale bars: 2 mm.

Female: Unknown.

Distribution.

China (Hunan Province) (Fig. 8).

Olios uniprocessus Hu, Zhang & Liu, sp. nov.

https://zoobank.org/5F787A9C-980C-46CF-9ADB-0E9D9B5E9434
Figs 3, 4, 8

Type material.

Holotype male: China • Chongqing Municipality: Beibei District, Southwest University, next to the international student apartment, 29.82852°N, 106.42972°E, elevation 253 m, 30 June 2015, Mingxin Liu leg.

Etymology.

The specific name is a combination of the Latin uni (meaning single) and processus (meaning process, projection), referring to the single projection (TAP III) on the embolic base, noun.

Diagnosis.

The male of Olios uniprocessus Hu, Zhang & Liu, sp. nov. resembles that of O. rossettii (Leardi, 1901) (cf. Fig. 3A–C vs. figs 146–147 in Jäger 2020) by having two tibial apophyses, and small U-shaped embolus arising sub-centrally from the tegulum, but can be recognized by: 1) retrolateral tibial apophysis leaf-shaped in retrolateral view, with a serrated retrolateral margin; and 2) tegular apophysis complex, with a sharp projection (TAP I) prolaterally and a sharp projection (TAP III) on the embolic base (vs. retrolateral tibial apophysis triangular, tegular apophysis long triangular in O. rossettii).

Figure 3. 

Olios uniprocessus Hu, Zhang & Liu, sp. nov., left male palp. A. Prolateral; B. Ventral; C. Retrolateral. Abbreviations: C = conductor, E = embolus, ptA = proximal tibial apophysis, RTA = retrolateral tibial apophysis, TA = tegular apophysis, TAP I–III = projection I to III of tegular apophysis. Scale bars: 0.5 mm.

Description.

Male (holotype): Total length 9.0; DS 4.3 long, 4.2 wide; OS 4.8 long, 3.1 wide. Eyes: AME 0.31; ALE 0.22; PME 0.20; PLE 0.23; AME–AME 0.17; AME–ALE 0.06; PME–PME 0.31; PME–PLE 0.28; AME–PME 0.19; ALE–PLE 0.09; CH AME 0.17; CH ALE 0.21. Measurements of palp and legs: Pp: 5.4 (1.7, 0.5, 1.2, -, 2.0); I: 23.0 (6.2, 2.1, 6.5, 6.4, 1.8); II: 26.4 (7.4, 2.0, 7.8, 7.3, 1.9); III: 17.8 (5.3, 1.7, 4.8, 4.5, 1.5); IV: 19.9 (5.9, 1.6, 5.3, 5.6, 1.5). Leg formula: II-I-IV-III. Spination: Pp: 131, 000, 3000; legs: Fe I 223, II–III 323, IV 220; Pa I–IV 000; Ti I 2134, II–III 2124, IV 2014; Mt I–III 2024, IV 3034. Cheliceral furrow with 2 anterior and 5 posterior teeth.

Palp (Fig. 3A–C): As in diagnosis, ptA arising submedially from Ti, RTA arising distally from Ti. Cymbium almost 1.5 times longer than Ti. TA complicated, with a sharp projection (TAP I) on prolateral margin, a lamellar projection (TAP II) on dorsal TA, and a sharp projection (TAP III) on embolic base. C tiny. E arising sub-centrally from tegulum in 10-o’clock-position.

Colouration (Fig. 4A, B): DS orange, with thick hairs. Sternum and legs yellow. OS light yellow, dorsal with several brown marks.

Figure 4. 

Olios uniprocessus Hu, Zhang & Liu, sp. nov., male habitus. A. Dorsal; B. Ventral. Scale bars: 2 mm.

Female: Unknown.

Distribution.

China (Chongqing Municipality) (Fig. 8).

Olios species without affiliation to a species-group

Remarks. The following species do not exhibit clear affiliation with any of the species-groups proposed to date.

Olios lincangensis Hu, Zhang & Liu, sp. nov.

https://zoobank.org/2F2871EE-9705-455D-8510-1B39551AEDE0
Figs 5, 6, 7, 8

Type material.

Holotype male: China • Yunnan Province: Lincang City, Yongde County, Daxueshan Yi, Lahu, Dai Township, Daxueshan Mountain, 24.11470°N, 99.64802°E, elevation 3428 m, 13 November 2021, Huachang Li leg. Paratype: 1 female, with same data as for holotype.

Etymology.

The specific name is derived from the type locality: Lincang City; adjective.

Diagnosis.

The male of Olios lincangensis Hu, Zhang & Liu, sp. nov. resembles that of O. feldmanni Strand, 1915 (cf. Fig. 5A–C vs. figs 294–295 in Jäger 2020) by having a spike-like dorsal part of retrolateral tibial apophysis, complex ventral part of the retrolateral tibial apophysis, well-developed conductor arising sub-centrally from the tegulum in an 11-o’clock-position, and an embolus arising from the tegulum in a 6-o’clock-position, but can be recognized by: 1) main part of ventral part of retrolateral tibial apophysis irregularly thick forked, 2) tegulum with a small nipple-shaped tegular apophysis, and 3) conductor with two apophyses (see arrows in Fig. 5A) (vs. main part of ventral part of retrolateral tibial apophysis clavate, tegular apophysis absent, conductor without apophysis in O. feldmanni).

Figure 5. 

Olios lincangensis Hu, Zhang & Liu, sp. nov., left male palp. A. Prolateral, arrows point to the two apophysis of prolateral conductor; B. Ventral; C. Retrolateral, arrows point to the three apices of vRTA. Abbreviations: C = conductor, dRTA = dorsal part of retrolateral tibial apophysis, E = embolus, TA = tegular apophysis, vRTA = ventral part of retrolateral tibial apophysis. Scale bars: 0.5 mm.

The female of Olios lincangensis Hu, Zhang & Liu, sp. nov. resembles that of O. croseiceps (Pocock, 1898) (cf. Fig. 6A–D vs. figs 236–240 in Jäger 2020) by having an ungulate-shaped epigynal field, medially located copulatory ducts and large spermathecae, but can be recognized by the fertilization ducts laterally located, the distance between the basal fertilization ducts almost equal to the width of the epigynal field (vs. fertilization ducts medially located, the distance between basal fertilization ducts almost 1/3 of the width of the epigynal field in O. croseiceps).

Figure 6. 

Olios lincangensis Hu, Zhang & Liu, sp. nov., female. A. Epigyne, ventral; B. Vulva, dorsal; C. Vulva, dorsal, the membrane between FD removed; D. Schematic course of right internal duct system, dorsal. Abbreviations: CD = copulatory duct, CO = copulatory opening, ES = epigynal slit, FD = fertilization duct, LL = lateral lobes, S = spermathecae. Scale bars: 0.5 mm.

Description.

Male (holotype): Total length 12.5; DS 5.7 long, 5.8 wide; OS 7.1 long, 5.0 wide. Eyes: AME 0.31; ALE 0.33; PME 0.21; PLE 0.35; AME–AME 0.24; AME–ALE 0.29; PME–PME 0.54; PME–PLE 0.49; AME–PME 0.28; ALE–PLE 0.24; CH AME 0.17; CH ALE 0.22. Measurements of palp and legs [I: missing]: Pp: 8.5 (2.6, 0.9, 1.4, -, 3.6); II: 28.0 (7.8, 2.2, 8.1, 7.7, 2.2); III: 21.3 (6.6, 2.0, 5.7, 5.2, 1.8); IV: 23.6 (7.1, 2.0, 6.4, 6.3, 1.8). Spination: Pp: 131, 000, 0001; legs: Fe II–IV 323; Pa II–IV 000; Ti II–IV 2124; Mt II–III 2024, IV 3025. Cheliceral furrow with 2 anterior and 5 posterior teeth.

Palp (Fig. 5A–C): As in diagnosis. Ti short, length of Ti almost 1/3 of cymbium. RTA well developed, vRTA with three apices in retrolateral view (see arrows in Fig. 5C). Tegulum with a small nipple-shaped TA. C tubular, with two aphophyses on prolateral part (see arrows in Fig. 5A). E thick and filiform.

Colouration (Fig. 7A, B): DS and sternum orange. Legs yellow to orange. OS dorsally yellow, with some brown marks, ventrally brown, with two yellow longitudinal lines.

Figure 7. 

Olios lincangensis Hu, Zhang & Liu, sp. nov., habitus. A, B. Male (A. Dorsal; B. ventral); C, D. Female (C. Dorsal; D. Ventral). Scale bars: 2 mm.

Female (paratype): Total length 14.1; DS 6.1 long, 6.7 wide; OS 8.3 long, 5.0 wide. Eyes: AME 0.35; ALE 0.32; PME 0.17; PLE 0.29; AME–AME 0.34; AME–ALE 0.29; PME–PME 0.67; PME–PLE 0.65; AME–PME 0.38; ALE–PLE 0.30; CH AME 0.14; CH ALE 0.23. Measurements of palp and legs: Pp: 7.4 (2.2, 0.9, 1.5, -, 2.8); I: 23.5 (6.8, 2.3, 6.2, 6.2, 2.0); II: 25.9 (7.7, 2.4, 7.0, 6.8, 2.0); III: 18.7 (5.9, 1.8, 4.9, 4.4, 1.7); IV: 20.2 (6.1, 1.7, 5.3, 5.2, 1.9). Leg formula: II-I-IV-III. Spination: Pp: 131, 020, 1111, 1012; legs: Fe I 223, II–III 323, IV 321; Pa I–IV 000; Ti I–III 2024, IV 2004; Mt I–III 2024, IV 3034. Cheliceral furrow with 2 anterior and 5 posterior teeth.

Epigyne (Fig. 6A–D): As in diagnosis. Epigynal field rounded rectangular, longer than wide. Length of ES almost 2/3 of the length of epigynal field. Internal duct system obviously sclerotized. Length of CD almost half of the length of S. S large, with rough anterior part. FD long and narrow.

Colouration (Fig. 7C–D): As in male, but generally darker.

Distribution.

China (Yunnan Province) (Fig. 8).

Figure 8. 

Distribution map of the three new Olios species 1 O. biprocessus Hu, Zhang & Liu, sp. nov. 2 O. uniprocessus Hu, Zhang & Liu, sp. nov. 3 O. lincangensis Hu, Zhang & Liu, sp. nov.

Discussion

The current paper reports three new Olios species from China: O. biprocessus Hu, Zhang & Liu, sp. nov., from Hunan Province; O. uniprocessus Hu, Zhang & Liu, sp. nov., from Chongqing Municipality; and O. lincangensis Hu, Zhang & Liu, sp. nov., from Yunnan Province. Two of these three new species (O. biprocessus and O. uniprocessus) belong to the rossettii-group, which is widely distributed across Asia and Africa (Jäger 2020). Jäger and Otto (2007) divided this species-group into two sub-groups based on the morphology of male palp: sub-group A with a simple RTA, sub-group B with both an RTA and a ptA. However, the position of the subtegulum in these sub-groups is inconsistent. Therefore, we consider proposing a new division of the sub-groups based on the position of subtegulum: group A with the subtegulum being located at a 6 o’clock position of the tegulum [e.g. O. japonicus Jäger & Ono, 2000, O. kiranae Sethi & Tikader, 1988, O. kolosvaryi (Caporiacco, 1947), O. mahabangkawitus Barrion & Litsinger, 1995, O. sericeus (Kroneberg, 1875), O. suavis (O. Pickard-Cambridge, 1876)]; group B with the subtegulum being invisible in ventral view [e.g. O. biprocessus Hu, Zhang & Liu, sp. nov., O. rossettii (Leardi, 1901), O. tiantongensis (Zhang & Kim, 1996), and O. uniprocessus Hu, Zhang & Liu, sp. nov.] (Jäger and Ono 2000; Jäger et al. 2002; Jäger and Kunz 2005; Jäger et al. 2022; Hu et al. 2025).

The copulatory organs of Olios lincangensis Hu, Zhang & Liu, sp. nov. show strong similarity to those of Olios feldmanni Strand, 1915 from Cameroon and O. croseiceps (Pocock, 1898) from Malawi, including a spike-like dRTA, complex vRTA, similar elongations of the conductor and embolus in the male palp, and similarly shaped lateral lobes and internal duct system in females. These three species probably belong to the same undefined species-group. The similar palpal conformation, especially RTA of O. feldmanni and O. lincangensis, suggests that this undefined group may be closely related to Curicaberis Rheims, 2015 (Sparassinae) from North and Central America (Rheims 2015).

The discovery of these new species in our research further enriches the species diversity of the genus Olios in China and provides new information on its geographical distribution in Asia. However, the females of two of the three new species are unknown. The low population density and docile nature of Olios spiders likely make them difficult to collect, resulting in less research attention compared to other Chinese sparassid genera such as Pseudopoda Jäger, 2000 and Sinopoda Jäger, 1999 (Zhang et al. 2025).

Acknowledgements

We are very grateful to Huachang Li (Lincang, China), Jiwei Lin (Changsha, China) and Mingxin Liu (Southwest University, China) for collecting specimens. We also thank Yejie Lin (Imperial College London, United Kingdom) for his useful suggestions on species identification, and Suping Hao for her support. This manuscript benefited greatly from the subject editor Cristina Rheims, and two reviewers, Alireza Zamani and John T.D. Caleb.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This study was supported in part by grants from National Natural Science Foundation of China (NSFC-32300378), the Science & Technology Fundamental Resources Investigation Program of China (2023FY100200), the Hebei Natural Science Foundation (C2025108010), the Introduction of active ingredient extraction technology and research on the antioxidant properties, development, and application of Ziziphus jujuba var. spinosa (C20230302), and Key Research and Development Project of Hebei Province, Scientific and Technological Innovation Team of Modern Seed Industry of Traditional Chinese Medicine (21326312D).

Author contributions

Writing – original draft: HZ, CH. Writing – review and editing: HZ, WN, JL, CW, CH.

Author ORCIDs

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

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

Caijun Wang https://orcid.org/0009-0005-2613-8117

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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He Zhang and Weitao Niu contributed equally to this work.
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