scieee AI-readable full text Open interactive document viewer

Clematis danxiacola (Ranunculaceae), a new species from the Danxia landform area in Zhejiang Province, China

Li, Jun-Ping; Ma, Qing; Ma, Dan-Dan; Wang, Jun-Feng; Xie, Wen-Yuan; Wu, Dong-Hao; Ying, Shang-Jiao; Chen, Zheng-Hai; Li, Pan

Abstract

Clematis danxiacola, a novel species of Ranunculaceae discovered in the Danxia landform area of Zhejiang, China, is described and illustrated herein. This species exhibits morphological similarities to C. terniflora and C. chinensis, but can be distinguished by specific characteristics, including the indumentum of leaflet blades, sepal apex morphology, stamen and pistil quantities, and achene morphology and dimensions. The plastome of C. danxiacola sp. nov. extends 159,506 bp and contains two inverted repeats of 31,040 bp, separated by a large single-copy region of 82,836 bp and a small single-copy region of 18,407 bp. The analysis revealed 136 functional genes, including 92 protein-coding genes, 36 tRNA genes, and 8 rRNA genes. Phylogenetic analyses demonstrated that C. danxiacola shares close evolutionary relationship with C. terniflora and C. chinensis.

Full text

97 Clematis danxiacola (Ranunculaceae), a new species from the Danxia landform area in Zhejiang Province, China Jun-Ping Li1* , Qing Ma2* , Dan-Dan Ma3, Jun-Feng Wang4, Wen-Yuan Xie5, Dong-Hao Wu6, Shang-Jiao Ying1, Zheng-Hai Chen5,7 , Pan Li8,9 1 Zhejiang Yongkang Forestry Bureau, Yongkang 321300, China 2 College of Biology and Environmental Engineering, Zhejiang Shuren University, Hangzhou 310015, China 3 Jiyang College, Zhejiang A&F University, Zhuji 311800, China 4 ScientificResearchManagementCenterofEastChinaMedicalBotanicalGarden,Lishui323000,China 5 ZhejiangForestResourcesMonitoringCentre,Hangzhou310020,China 6 LishuiMunicipalAdministrationofMarketSupervisionofZhejiang,Lishui323000,China 7 ZhejiangForestrySurveyPlanningandDesignCo.Ltd.,Hangzhou310020,China 8 KeyLaboratoryofBiodiversityandEnvironmentontheQinghaiTibetanPlateau,MinistryofEducation,SchoolofEcologyandEnvironment,XizangUniversity, Lhasa850000,China 9 MotuoBiodiversityObservationandResearchStationofXizangAutonomousRegion,Motuo860700,China Correspondingauthors:Zheng-HaiChen([email protected]);PanLi([email protected]) Copyright: © Jun-Ping Li 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 Clematis danxiacola, a novel species of Ranunculaceae discovered in the Danxia landform area of Zhejiang, China, is described and illustrated herein. This species exhibits morphological similarities to C. terniflora and C. chinensis, but can be distinguished by specific characteristics, including the indumentum of leaflet blades, sepal apex morphology, stamen and pistil quantities, and achene morphology and dimensions. The plastome of C. danxiacola sp. nov. extends 159,506 bp and contains two inverted repeats of 31,040 bp, separated by a large single-copy region of 82,836 bp and a small single-copy region of 18,407 bp. The analysis revealed 136 functional genes, including 92 protein-coding genes, 36 tRNA genes, and 8 rRNA genes. Phylogenetic analyses demonstrated that C. danxiacola shares close evolutionary relationship with C. terniflora and C. chinensis. Key words: China, Clematis, Danxia landform, new species Introduction Clematis L. (1753: 543) is one of the few cosmopolitan genera in the buttercup family (Ranunculaceae), comprising approximately 300 species predominantly distributed across north temperate zones (Wang 1980; Ziman and Keener 1989; Tamura 1995; Johnson 1997; Grey-Wilson 2000; He et al. 2021). China hosts approximately 147 Clematis species, with 93 being endemic (Wang and Bartholomew 2001). The taxonomy of Clematis has been notoriously difficult due to high species-level morphological variation (Johnson 1997; Brandenburg 2000; Grey-Wilson 2000; Wang and Li 2005). Multiple taxonomic schemes have emerged in recent years addressing genus delineation, infrageneric classification, and Academic editor: Andrey Erst Received: 3 April 2025 Accepted: 14 October 2025 Published: 7 November 2025 Citation: Li J-P, Ma Q, Ma D-D, Wang J-F, Xie W-Y, Wu D-H, Ying S-J, Chen Z-H, Li P (2025) Clematis danxiacola (Ranunculaceae), a new species from the Danxia landform area in Zhejiang Province, China. PhytoKeys 266: 97–115. https://doi.org/10.3897/ phytokeys.266.154626 PhytoKeys 266: 97 –115 (2025) DOI: 10.3897/phytokeys.266.154626 * The authors contributed equally to this paper. 98 PhytoKeys 266: 97 –115 (2025), DOI: 10.3897/phytokeys.266.154626 Jun-Ping Li et al.: Clematis danxiacola, a new species from Danxia landform Areas in Eastern China species delimitation in Clematis (Tamura 1995; Johnson 1997; Wang 1998, 2003, Grey-Wilson 2000; Wang and Li 2005; Xiao et al. 2022). Recent molecular phylogenetic studies suggest substantial convergence in floral and trophic characteristics in Clematis, challenging previous infrageneric classifications based on morphological characteristics (Miikeda et al. 2006; Xie et al. 2011), but conforms to Johnson’s broad concept that Clematis should encompass all previously recognized genera of subtribe Clematidinae (Johnson, 1997). Currently, ten clades within Clematis have been resolved as regional geographic groups (Xie et al. 2011). During a field survey of the Danxia landform area in Zhejiang, China, we identified an unknown species of Clematis. The species displayed morphological similarities to C. terniflora DC. (1818: 137), but was notably distinct in habitat, and leaf and floral characteristics, prompting investigation of its taxonomic status. Materials and methods Morphological observation From 2021 to 2024, we conducted field excursions to the Danxia landform area in Zhejiang Province, China, including Yongkang County, Wuyi County, and Wucheng District (Jinhua); Songyang County, Jinyun County, Qingtian County, and Liandu District (Lishui); Jiangshan County and Kecheng District (Quzhou); Zhuji County and Xinchang County (Shaoxin); and Yuyao County and Fenghau District (Ningbo), to observe and investigate this unknown species. Specimens in the principal herbaria in Zhejiang Province (HHBG, HTC, HZU, and ZM) (Thiers 2024) and specimen photographs from the Chinese Virtual Herbarium (CVH; www.cvh.ac.cn/) were examined to identify specimens similar to the unknown species. High-definition images of similar specimens from international herbaria (A, BM, E, FI, G, K, LD, LINN, MPU, and P) were also consulted. Based on field investigation and herbarium specimen evaluation, the morphology of this new species was documented through comparison with protologues from Greuter (1965) and de Candolle (1818). Sampling and sequencing Material sampling From August 10 to August 18, 2023, we collected one leaf sample each from 7 populations of the unknown taxon in Jinhua and Lishui cities of the Danxia landform area in Zhejiang Province (Table 1). Leaves were initially placed in non-woven bags, then stored in sealing bags containing allochroic silica gel for rapid drying. Herbarium specimens were preserved in the herbaria of Zhejiang Museum of Natural History (ZM); Institute of Botany, Chinese Academy of Sciences (PE); and Kunming Institute of Botany, Chinese Academy of Sciences (KUN) (Table 1). DNA extraction, PCR amplification, and sequencing Whole-genomic DNA from silica-dried leaf tissue was extracted using a modified CTAB protocol (Chen et al. 2014). For plastome sequencing, high-quality DNA was fragmented into lengths of ≤800 bp. Fragment quality was verified 99 PhytoKeys 266: 97 –115 (2025), DOI: 10.3897/phytokeys.266.154626 Jun-Ping Li et al.: Clematis danxiacola, a new species from Danxia landform Areas in Eastern China using an Agilent Bioanalyzer 2100 (Agilent Technologies, Palo Alto, CA, USA). A short-insert (500 bp) paired-end library was constructed and sequenced by Nanopore Technology (Wuhan, China) on a DNBSEQ-T7 sequencer with 150 bp paired-end reads. For amplification and sequencing of ribosomal DNA (rDNA) internal transcribed spacer (ITS) regions, primers were designed using ITS sequences of closely related species: forward, 5′-ATGCGATACTTGGTGTGAAT-3′; reverse, 5′-GACGCTTCTCCAGACTACA-3′. The amplification fragments encompass the ITS1, ITS2, 5.8 rDNA, and 26S rDNA regions. DNA amplification was conducted in 50 μL reactions containing 1 μL of total DNA (20 ng/μL), 5 μL of 10× buffer with 2.5 mM Mg2+, 1 μL of dNTP (10 mM), 1.5 μL of each primer (10 μM), and 1 μL of Taq polymerase (5 u/μL) in an ABI-2720 thermocycler (Applied Biosystems, Waltham, MA, USA). The PCR protocol consisted of initial denaturation at 95 °C for 5 min, followed by 35 cycles of denaturation at 95 °C for 30 s, annealing at 58 °C for 30 s, and extension at 72 °C for 90 s, with a final extension at 72 °C for 7 min. PCR products were purified by ethanol precipitation. The purified products were analyzed by electrophoresis on a 1% (w/v) agarose gel and spectrophotometric analysis (NanoDrop 2000; Thermo Fisher Scientific, Waltham, MA, USA) before Sanger sequencing using an ABI3730-XL sequencer (Applied Biosystems). Data assembly and annotation For plastome assembly and annotation, raw sequencing data was filtered using FastQC (www.bioinformatics.babraham.ac.uk/projects/fastqc/). Clean paired-end reads were de novo assembled using the GetOrganelle pipeline (Jin et al. 2020) using SPAdes 3.10.1 as the assembler (Bankevich et al. 2012). The plastome of C. terniflora (NC028000) served as the reference. The assembled plastome was imported into Geneious Prime (https://www. geneious.com) for annotation and identification of putative starts, stops, and intron positions through comparison with homologous genes of published Clematis plastomes. The circular plastome map of the new species was generated using OGDRAW (http://ogdraw.mpimp-golm.mpg.de/) (Greiner et al. 2019). Table 1. List of analyzed samples of Clematis danxiacola sp. nov. Molecular specimen ID Herbarium specimen ID Locality Coordinates Altitude (a.s.l.) YK-03 ZMNH0067437 Feilongshan, Xicheng Subdistrict, Yongkang County, Jinhua City 120˚0'41.40"E, 28˚59'23.88"N 181 YK-04 ZMNH0067438 Longqingkeng, Wangxitian Village, Xiangzhu Town, Yongkang County, Jinhua City 120˚2'33.06"E, 29˚0'8.40"N 246 YK-07 ZMNH0067439 Qianlang Village, Shizhu Town, Yongkang County, Jinhua City 120˚7'14.50"E, 28˚49'59"N 175 YK-08 ZMNH0067436 PE02621603 KUN1644125 Chiyantang Reservoir, Xianling Village, Zhiying Town, Yongkang County, Jinhua City 120˚8'7.86"E, 28˚54'29.28"N 172 LD-01 ZMNH0067440 Nanbenjis, Baiyunshan, Baiyun Subdistrict, Liandu District, Lishui City 119˚55'4.71"E, 28˚28'54.4"N 176 LD-02 ZMNH0067441 Huangnidan Village, Zijin Subdistrict, Liandu District, Lishui City 119˚57'47.73"E, 28˚27'37.23"N 98 LD-03 ZMNH0067442 Huangcun Village, Huangcun Town, Liandu District, Lishui City 120˚2'48.74"E, 28˚30'40.02"N 129 100 PhytoKeys 266: 97 –115 (2025), DOI: 10.3897/phytokeys.266.154626 Jun-Ping Li et al.: Clematis danxiacola, a new species from Danxia landform Areas in Eastern China Phylogenetic analyses For phylogenetic analyses, complete plastome sequences of 85 Clematis species and ITS sequences of 80 Clematis species were obtained from GenBank (Suppl. material 1: table S1). The sampling strategy encompassed all major clades within Clematis previously identified (Xie et al. 2011; Xiao et al. 2022). The plastome and ITS dataset were aligned independently using multiple alignment using fast Fourier transform (MAFFT) v7.490 implemented in Geneious Prime (Katoh and Standley 2013). Phylogenetic analyses based on ITS and plastome sequences (excluding one copy of the inverted repeat) were conducted to explore the evolutionary relationship among the new species and other Clematis species using the maximum likelihood method with RAxML-HPC v8.2.12 (Stamatakis 2014) on the CIPRES Science Gateway (http://www.phylo.org/). According to Jiang et al. (2017) and Xiao et al. (2022), Anemoclema glaucifolium (Franch.) W.T. Wang (Ranunculaceae) was chosen as an outgroup. Morphological Analysis Morphological characteristics were documented from 10 wild individuals of C. danxiacola sp. nov. during field surveys. For comparative analysis, 10 herbarium specimens each of C. terniflora, C. chinensis, and C. chinensis var. vestita were examined from the Chinese Virtual Herbarium (CVH; https://www.cvh.ac.cn/). Twenty quantitative traits (Table 2) were selected to construct a morphological matrix. Principal component analysis (PCA) was performed for dimensionality reduction, and a multidimensional morphospace was established based on the first two principal components. All computations were conducted using GraphPad Prism v10.1.2 (Build 324). Results Characteristics of the plastome The plastome size of Clematis danxiacola sp. nov. is 159,506 bp and comprises two 31,040 bp inverted repeats, a 18,407 bp small single-copy region, and a 82,836 bp large single-copy region. The overall GC content is 38%, and the GC contents of the large single-copy, small single-copy, and inverted repeat regions are 36.3%, 31.4%, and 42.0%, respectively. The complete plastome contains 136 functional genes, including 92 protein-coding genes, 36 tRNA genes, and 8 rRNA genes (Fig. 1, Suppl. material 1: table S2). The complete plastome sequence of C. danxiacola after annotation was deposited in GenBank (PQ246280). Molecular analysis Phylogenetic analyses based on both plastome and ITS datasets revealed that individuals of C. danxiacola were clustered with high bootstrap support values (Figs 2–3). The new species is closely related to C. terniflora and C. chinensis Osbeck (1757: 205) and belongs to clade V (sensu Xie et al. 2011) or clade II (sensu Xiao et al. 2022) of Clematis, which comprises species of sect. Flammula DC., sect. Viticella DC., and sect. Viorna (sensu Tamura 1995). 101 PhytoKeys 266: 97 –115 (2025), DOI: 10.3897/phytokeys.266.154626 Jun-Ping Li et al.: Clematis danxiacola, a new species from Danxia landform Areas in Eastern China Morphological comparison Morphological comparisons between C. danxiacola sp. nov., C. terniflora, C. chinensis, and C. chinensis var. vestita (Rehder & E.H.Wilson) W.T. Wang (1998: 158) are presented in Table 2. This new species shares characteristics with C. terniflora and C. chinensis in having climbing stems, entire leaflet, cymes, small flowers, white and spreading sepals, and glabrous stamens; however, it is distinctly different from them in habitat, texture and indumentum of leaflet blades, and sepal apex shape (Figs 5–7). PCA Multivariate analysis provided a quantitative assessment of morphological differentiation among the four species. The analysis revealed distinct variation patterns along the first two axes (Fig. 4). Two clearly separated clusters emerged: one comprising all specimens of C. danxiacola sp. nov., and another containing its allied species. C. danxiacola demonstrated a discontinuous distribution in the principal component (PC) 1–PC2 morphospace compared with the other three taxa, forming statistically distinct clusters (Fig. 4). PCA revealed that the first seven principal components derived from the 20 morphological traits collectively explained 85.16% of the total variation. PC1 accounted for 31.72% of the variance, while PC2 explained 13.93%. The primary traits associated with PC1 included specimen color after drying, sepal apex characteristics, habitat preferences, fresh leaf taste, and sepal Table 2. PCA factor loadings based on 20 morphological characteristics. Variable Data Type PC1 PC2 PC3 Specimen color after drying (blackened 1 / unblackened 0) DV 0.808 −0.366 0.121 Compound leaf length (cm) CV −0.46 0.185 0.564 Compound leaf width (cm) CV −0.605 0.162 0.539 Compound leaf length/width ratio CV 0.42 −0.093 −0.105 Indumentum on leaf abaxial surface (densely pubescent 1 / glabrous or glabrescent 0) DV −0.643 −0.612 −0.187 Terminal leaflet length (cm) CV −0.693 −0.249 0.47 Terminal leaflet width (cm) CV −0.215 −0.064 0.367 Terminal leaflet length/width ratio CV −0.548 −0.256 0.303 Petiolule length (cm) CV −0.024 0.563 0.361 Pedicel length (cm) CV 0.158 0.128 0.256 Calyx apex morphology (truncate 1 / mucronate 0) DV −0.879 −0.399 −0.048 Calyx length (mm) CV −0.4 0.596 0.254 Calyx width (mm) CV −0.764 0.194 −0.3 Calyx length/width ratio CV 0.403 0.346 0.56 Stamen number CV 0.379 0.262 −0.048 Fruit length (mm) CV −0.289 0.609 −0.312 Persistent style length (cm) CV 0.502 −0.279 0.386 Fruit length/persistent style ratio CV −0.296 0.512 −0.508 Taste of fresh leaves (pungent 1 / non-pungent 0) DV −0.808 0.366 −0.121 Habitat (ridge, hillside 1 / valley, plain or wetland 0) DV −0.879 −0.399 −0.048 DV: dummy variable; CV: continuous variable 102 PhytoKeys 266: 97 –115 (2025), DOI: 10.3897/phytokeys.266.154626 Jun-Ping Li et al.: Clematis danxiacola, a new species from Danxia landform Areas in Eastern China width. PC2 was predominantly influenced by leaf abaxial surface indumentum, fruit length, sepal length, and petiolule length (Table 2). Taxonomic treatment Clematis danxiacola J.P.Li, P.Li & Z.H.Chen, sp. nov. urn:lsid:ipni.org:names:77371642-1 Figs 5–7 Chinese name: 丹霞铁线莲 Type. China • Zhejiang Province, Jinhua City, Yongkang County, Zhiying Town, Xianling Village, Chiyantang Reservoir, purple glutenite in the Danxia landform Figure 1. Plastome map of Clematis danxiacola sp. nov. Genes inside the circle are transcribed clockwise, while those outside are transcribed counterclockwise. In the inner circle, light gray represents the AT content, and dark gray represents the GC content. Different colors indicate genes belonging to distinct functional groups. 103 PhytoKeys 266: 97 –115 (2025), DOI: 10.3897/phytokeys.266.154626 Jun-Ping Li et al.: Clematis danxiacola, a new species from Danxia landform Areas in Eastern China area, in bushes at the forest margin at foothills, alt. 172 m a.s.l., 28˚54'29.28"N, 120˚8'7.86"E, August 18, 2023, He-Ping Chen, Jun-Ping Li et Zheng-Hai Chen YK23081807 (holotype: ZM barcode ZMNH0067436! isotypes: HZU!, KUN!, PE!). Diagnosis. This new species resembles C. terniflora but differs in several characteristics including leaflet blades abaxially densely persistently appressed-puberulous (vs. sparsely puberulous and glabrescent), sepals cuneate or oblanceolate, apex subtruncate and premorse (vs. obovate–oblong to oblong, apex ± acute to obtuse and entire), pistils (5–) 8–12 (vs. 4–7), achene ovate, compressed but slightly swollen in the middle, margin not or slightly thickened (vs. broadly elliptic to obovate, strongly compressed, margin distinctly thickened). Morphological description. Evergreen woody vine. Stem with shallowly 6–12-grooved; branchlets densely gray appressed-puberulous, glabrescent except at nodes; axillary buds triangular–ovate, densely gray puberulous. Leaves opposite, 1-pinnate, usually 5-foliolate, sometimes 2-pinnate on new shoots in Figure 2. Maximum likelihood tree inferred from Clematis plastomes to elucidate the phylogenetic position of C. danxiacola sp. nov. Numbers above the branches indicate the bootstrap values (≥50%) of the maximum likelihood analysis. Branches and names in red indicate the new species (C. danxiacola sp. nov). Light red shading highlights the clade containing the new species, as revealed by phylogenetic analyses in Xie et al. (2011) and Xiao et al. (2022). Numerals on the right indicate the ten Clematis clades recognized in Xie et al. (2011) and Xiao et al. (2022): yellow, Xiao et al. (2022); blue, Xie et al. (2011). 104 PhytoKeys 266: 97 –115 (2025), DOI: 10.3897/phytokeys.266.154626 Jun-Ping Li et al.: Clematis danxiacola, a new species from Danxia landform Areas in Eastern China summer–fall; petiole 3–7 cm long, base with nodes usually dark purple or gray, puberulous; leaflet blades thick-papery or subleathery, ovate, 3.5–8.0 (–9.5) × 2.0–5.0 (–6.2) cm, apex acuminate, base broadly cuneate, subrounded to shallow cordate, margin entire, slightly revolute when drying, adaxially appressed-puberulous when young, glabrescent except for basal veins, lustrous, basal veins slightly prominent, abaxially densely persistently appressed-puberulous, basal veins prominent when dry; leaflets of summer–fall shoots thick leathery, ovate– lanceolate or lanceolate, 2.5–10.0 × 1–3 cm, apex long acuminate, base cuneate; petiolules 0.7–2.0 cm long, base usually dark purple, sometimes tendrilous. PaFigure 3. Maximum likelihood tree inferred from ITS sequences to elucidate the phylogenetic position of C. danxiacola sp. nov. Numbers above the branches indicate the bootstrap values (≥50%) of the maximum likelihood analysis. Branches and names in red indicate the new species (C. danxiacola sp. nov). Light red shading highlights the clade containing the new species, as revealed by phylogenetic analyses in Xie et al. (2011) and Xiao et al. (2022). The GenBank accession numbers used for the analysis are indicated after the species names. Numerals on the right indicate the ten Clematis clades recognized in Xie et al. (2011) and Xiao et al. (2022): yellow, Xiao et al. (2022); blue, Xie et al. (2011). 105 PhytoKeys 266: 97 –115 (2025), DOI: 10.3897/phytokeys.266.154626 Jun-Ping Li et al.: Clematis danxiacola, a new species from Danxia landform Areas in Eastern China niculate cymes axillary or terminal, usually many-flowered, 5–15 (–30) cm long; bracts usually narrowly lanceolate to linear, 0.7–1.5 (–2.3) × 1.5–4.0 mm, petiolate, rarely leaflike, both surfaces appressed-puberulous. Flowers 2–3 cm in diameter, pungent aromatic, bisexual, erect; buds ovoid, apex acute. Pedicel 1–2 cm long, together with peduncle and rachis densely persistently appressed-puberulous. Sepals 4, white, spreading, cuneate or oblanceolate, 10–14 × 4–6 mm, apex subtruncate and premorse, adaxially glabrous, abaxially densely puberulous, margin velutinous. Stamens 26–49, white, glabrous, 6–11 mm long; filaments 3–7 mm long; anthers linear, 2–4 mm long, apex obtuse or minutely apiculate. Pistils (5–) 8–12; ovaries pubescent; style densely villous. Achenes usually 3–7, brown, ovate, compressed but slightly swollen in the middle, 5.4–6.3 × 3.0–3.7 mm, densely ascending pubescent, margin not or slightly thickened; persistent styles 2.5–3.0 cm long, plumose, whitish (Figs 5–7). Phenology. Flowering occurs from May to November, with fruiting from September to January of the following year. Etymology. The specific epithet refers to the Danxia landform where the species occurs. Distribution and ecology. C. danxiacola occurs in Yongkang County of Jinhua City, and Qingtian County, Jinyun County, and Liandu District of Lishui City, Zhejiang Province, China (Fig. 8). The species inhabits sunny slopes or ridges of low hills of the Danxia landform within the elevation range 50–550 m a.s.l, typically climbing on tree canopies or shrubs at forest edges or roadsides. The substrate consists of soil derived from weathered purple sandy conglomerate. Conservation assessment. The currently known 19 populations of C. danxiacola are distributed across two prefecture-level cities, four counties, and 14 towns. The total distribution area is approximately 100 km2 with an estimated population exceeding 1000 mature individuals. Despite occurring in low-hill areas characterized by frequent human activities and habitat fragmentation, the species demonstrates resilience due to its drought tolerance, arid soil Figure 4. Principal component axes 1 and 2 representing morphological data of C. danxiacola sp. nov. and three other similar species. 112 PhytoKeys 266: 97 –115 (2025), DOI: 10.3897/phytokeys.266.154626 Jun-Ping Li et al.: Clematis danxiacola, a new species from Danxia landform Areas in Eastern China Conclusion This study presents a new species of Clematis based on morphological and molecular evidence. The documented plastome provides valuable data for future research on the systematics, evolution, and conservation of the genus. Acknowledgements We would like to thank the herbaria A, BM, E, FI, G, K, LD, LINN, MPU, P and PE for providing the high-resolution images of specimens. The authors are also grateful to Mr. Jun Liu from Zhejiang University Library for accessing the literature; to Prof. Gen-You Li, Prof. Jian-Sheng Wang, Mr. Jian-Ping Zhong, Mr. Yi-Rong Zhu, Mr. Wei-Qi Lin, Mr. Liang-Dong Xu, Mr. Xia-Shuo Lei, Mr. He-Ping Chen, Mr. Yong-Jun Chen as well as kindly colleagues for their assistance in the field work; to Mrs. Yi-Chen Wang for drawing the illustrations. Zhe-Yu Xu for working on the PCA calculations. We also acknowledge TopEdit LLC for the linguistic editing and proofreading during the preparation of this 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 study was financially supported by the Project of Survey, File and Codification of Wild Plants and the Second Edition of Flora of Zhejiang (335010-2015-0005), the Research and Promotion of Natural Science Popularization Education in Yongkang (202153), the Research on the Background Investigation of Plant Resources in Yongkang City (DTCGYK2023-014), the Research on the Classification of Doubtful Plants in Zhejiang Province (ZFSPLD2025-01), and Science and Technology Projects of Xizang Autonomous Region, China (XZ202402ZD0005). Author contributions Conceptualization and Methodology: ZHC, PL. Investigation: JPL, WYX, JFW, DHW, SJY, ZHC. Data Curation: WYX, JFW. Software and Visualization: QM, DDM. Writing-Original draft: ZHC, JPL, QM. Writing-Review and Editing: PL, ZHC. All authors have read and approved the final manuscript. Author ORCIDs Jun-Ping Li https://orcid.org/0009-0000-7655-3849 Qing Ma https://orcid.org/0000-0002-2322-8847 Dan-Dan Ma https://orcid.org/0000-0002-7273-9141 Jun-Feng Wang https://orcid.org/0000-0002-8110-9045 113 PhytoKeys 266: 97 –115 (2025), DOI: 10.3897/phytokeys.266.154626 Jun-Ping Li et al.: Clematis danxiacola, a new species from Danxia landform Areas in Eastern China Wen-Yuan Xie https://orcid.org/0000-0003-1529-0787 Dong-Hao Wu https://orcid.org/0009-0003-7113-6554 Shang-Jiao Ying https://orcid.org/0009-0005-7201-5775 Zheng-Hai Chen https://orcid.org/0000-0002-8852-7127 Pan Li https://orcid.org/0000-0002-9407-7740 Data availability All of the data that support the findings of this study are available in the main text or Supplementary Information. References Bankevich A, Nurk S, Antipov D, Gurevich AA, Dvorkin M, Kulikov AS, Lesin VM, Nikolenko SI, Pham S, Prjibelski AD (2012) SPAdes: A new genome assembly algorithm and its applications to single-cell sequencing. Journal of Computational Biology 19: 455–477. https://doi.org/10.1089/cmb.2012.0021 Bentham G (1861) Flora Hongkongensis: A Description of the Flowering Plants and Ferns of the Island of Hongkong. Lovell Reeve, London, 7 pp. https://doi.org/10.5962/ bhl.title.21052 Brandenburg WA (2000) Meclatis in Clematis: Yellow flowering Clematis species. Systematic studies in Clematis I. (Ranunculaceae), inclusive of cultonomic aspects. Wageningen Universiteit, Wageningen. Chen LY, Song MS, Zha HG, Li ZM (2014) A modified protocol for plant genome DNA extraction. Plant Diversity and Resources 36: 375–380. de Candolle AP (1818) Regni vegetabilis systema naturale I. Treuttel & Wurtz, Paris, 137 pp. https://doi.org/10.5962/bhl.title.59874 Franchet A (1885) Plantae Davidianae ex Sinarum Imperio: Plantes du Thibet Oriental, Province de Moupine. Nouvelles Archives du Muséum d’Histoire Naturelle, sér. 2, 8: 184. Greiner S, Lehwark P, Bock R (2019) OrganellarGenomeDRAW (OGDRAW) version 1.3.1: Expanded toolkit for the graphical visualization of organellar genomes. Nucleic Acids Research 47: W59–W64. https://doi.org/10.1093/nar/gkz238 Greuter W (1965) Beiträge zur Flora der Südägäis 1–7. Candollea 20: 212–213. https:// doi.org/10.5169/seals-880361 Grey-Wilson C (2000) Clematis the genus. Timber Press, Portland, 223 pp. He J, Lyu RD, Luo YK, Lin LL, Yao M, Xiao JM, Xie L, Wen J, Pei LY, Yan SX, Cheng J, Li JY, Li LQ (2021) An updated phylogenetic and biogeographic analysis based on genome skimming data reveals convergent evolution of shrubby habit in Clematis in the Pliocene and Pleistocene. Molecular Phylogenetics and Evolution 164: 1–14. https://doi. org/10.1016/j.ympev.2021.107259 IUCN (2024) Guidelines for Using the IUCN Red List Categories and Criteria. Version 16. Prepared by the Standards and Petitions Committee, Gland, Switzerland. http://www. iucnredlist.org/documents/RedListGuidelines.pdf [accessed 15 January 2025] Jiang N, Zhou Z, Yang JB, Zhang SD, Guan KY, Tan YH, Yu WB (2017) Phylogenetic reassessment of tribe anemoneae Ranunculaceae): Non-monophyly of Anemone s. l. revealed by plastid datasets. PLOS ONE 12: 1–17. https://doi.org/10.1371/journal. pone.0174792 Jin JJ, Yu WB, Yang JB, Song Y, Yi TH, Li DZ (2020) GetOrganelle: A fast and versatile toolkit for accurate de novo assembly of organelle genomes. Genome Biology 21: 241. https://doi.org/10.1186/s13059-020-02154-5 114 PhytoKeys 266: 97 –115 (2025), DOI: 10.3897/phytokeys.266.154626 Jun-Ping Li et al.: Clematis danxiacola, a new species from Danxia landform Areas in Eastern China Johnson M (1997) Släktet Klematis. Magnus Johnson Plantskola AB, Södertälje, 881 pp. Katoh K, Standley DM (2013) MAFFT multiple sequence alignment software version 7: Improvements in performance and usability. Molecular Biology and Evolution 30(4): 772–780. https://doi.org/10.1093/molbev/mst010 Linnaeus C (1753) Species Plantarum. Laurentius Salvius, Stockholm, 1200 pp. Maximowicz CJ (1877) Diagnoses plantarum novarum Japoniae et Mandshuriae. Bulletin de l’Académie Impériale des Sciences de Saint-Pétersbourg 22: 221. https:// doi.org/10.5962/bhl.title.46308 Miikeda O, Kita K, Handa T, Yukawa T (2006) Phylogenetic relationships of Clematis (Ranunculaceae) based on chloroplast and nuclear DNA sequences. Botanical Journal of the Linnean Society 152: 153–168. https://doi.org/10.1111/j.10958339.2006.00551.x Osbeck P (1757) Dagbok öfwer en Ostindisk resa åren 1750, 1751, 1752: Med anmårkningar uti naturkunnigheten, fråmmande folkslags språk, seder, hushållning, m. m. Stockholm, Lor. Ludv. Grefing, 205 pp. https://doi.org/10.5962/bhl.title.112527 Pallas PS (1776) Reise durch verschiedene Provinzen des russischen Reichs 3. St. Petersburg, Kaiserliche Akademie der Wissenschaften, 735 pp. Rehder A, Wilson EH (1913) Clematis. In: Sargent CS (Ed.) Plantae Wilsonianae 1. The University Press, Cambridge, 330 pp. Ruprecht FJ (1857) Die ersten botanischen Nachrichten über das Amurland. Zweite Abthkilung: Bäume und Sträucher, Bulletin de la Classe Physico-Mathematique de l’Academie Imperiale des Sciences de Saint-Pétersbourg. St. Petersburg ser. 2, 15: 258. Stamatakis A (2014) RAxML Version 8: A tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics (Oxford, England) 30(9): 1312–1313. https:// doi.org/10.1093/bioinformatics/btu033 Tamura M (1995) Clematis. In: Hiepko P (Ed.) Die Natürlichen Pflanzenfamilien. 2nd ed. 17a. Duncker & Humblot, Berlin, 368–387. Thiers B (2024 onwards) Index Herbariorum. http://sweetgum.nybg.org/science/ih/ [accessed on 13 December 2024] Wang WT (1980) Flora Reipublicae Popularis Sinicae 28. Science Press, Beijing, 75–235. Wang WT (1998) Notulae de Ranunculaceis Sinensibus (XXII). Zhiwu Fenlei Xuebao 36: 150–172. https://www.jse.ac.cn/EN/Y1998/V36/I2/150 Wang WT (2003) A revision of Clematis sect. Clematis (Ranunculaceae). Zhiwu Fenlei Xuebao 41(1): 1–62. https://doi.org/10.1360/aps050049 Wang WT, Bartholomew B (2001) Clematis L. In: Wu CY, Raven P (Eds) Flora of China 6. Science Press, Beijing & Missouri Botanical Garden Press, St. Louis, 333–386. Wang WT, Li LQ (2005) A new system of classification of the genus Clematis (Ranunculaceae). Zhiwu Fenlei Xuebao 43(5): 431–488. https://doi.org/10.1360/aps040091 Xiao JM, Lyu RD, He J, Li MY, Ji JX, Cheng J, Xie L (2022) Genome-partitioning strategy, plastid and nuclear phylogenomic discordance, and its evolutionary implications of Clematis (Ranunculaceae). Frontiers in Plant Science 13: 1–14. https://doi. org/10.3389/fpls.2022.1059379 Xie L, Wen J, Li LQ (2011) Phylogenetic analyses of Clematis (Ranunculaceae) based on sequences of nuclear ribosomal ITS and three plastid regions. Systematic Botany 36(4): 907–921. https://doi.org/10.1600/036364411X604921 Ziman SN, Keener CS (1989) A geographical analysis of the family Ranunculaceae. Annals of the Missouri Botanical Garden 76(4): 1012–1049. https://doi. org/10.2307/2399690 115 PhytoKeys 266: 97 –115 (2025), DOI: 10.3897/phytokeys.266.154626 Jun-Ping Li et al.: Clematis danxiacola, a new species from Danxia landform Areas in Eastern China Supplementary material 1 Supplementary tables S1, S2 Authors: Jun-Ping Li, Qing Ma, Dan-Dan Ma, Jun-Feng Wang, Wen-Yuan Xie, Dong-Hao Wu, Shang-Jiao Ying, Zheng-Hai Chen, Pan Li Data type: docx 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/phytokeys.266.154626.suppl1