Dysosma xishuiensis (Berberidaceae), a new species from Guizhou, China, based on morphological and molecular evidence

59 Dysosma xishuiensis (Berberidaceae), a new species from Guizhou, China, based on morphological and molecular evidence Lang Huang1,2 , Wei-Hao Yao1,2 , Yan-Bing Yang1,2 , Ming-Tai An3,4 , Yuan-Lin Yu5, Mei Zhou5, He Li1,2 1 Guizhou Academy of Forestry, Guiyang 550005, Guizhou, China 2 Key Laboratory for Biodiversity Conservation in Karst Mountain Area of Southwestern China, National Forestry and Grassland Administration, Guiyang 550005, Guizhou, China 3 College of Forestry, Guizhou University, Guiyang 550025, Guizhou, China 4 Center for Biodiversity and Natural Conservation, Guizhou University, Guiyang 550025, Guizhou, China 5 Guizhou Xishui National Nature Reserve Management Bureau, Xishui 564600, Guizhou, China Corresponding author: Ming-Tai An ([email protected]) Copyright: © Lang Huang 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 Dysosma xishuiensis Y. B. Yang, M. T. An & C. H. Yang is described and illustrated as a new species from Xishui County in northern Guizhou Province, China. This species is morphologically similar to D. versipellis, but differs by its glabrous abaxial leaf surface, pedicels, and sepals; inflorescences bearing more flowers (8–20 vs. 5–8); larger obovate–oblong petals (2.8–3.4 × 1.5–2.0 cm vs. 2.5 × 0.8 cm); and an obpyriform ovary (vs. ellipsoid). The phylogenetic relationships reconstructed using ITS, matK, and rbcL sequences further confirm that it is a new species within Dysosma. This species is currently known only from the Xishui National Nature Reserve in Guizhou. According to IUCN criteria (B2, D), due to its limited distribution and low population numbers, D. xishuiensis is assessed as Endangered (EN). Key words: Dysosma, molecular phylogeny, morphology, new taxon, taxonomy Introduction Dysosma Woodson (Podophylloideae, Berberidaceae) is a small genus comprising nine species, primarily distributed in China and Vietnam. China is considered the center of diversity and origin for Dysosma (Ma and Hu 1997), with the eastern Yunnan–Guizhou Plateau to the Three Gorges region being a key center for the distribution and diversification of Dysosma species (Ying 1979). Dysosma is closely related to Sinopodophyllum (Royle) T.S. Ying, Podophyllum L., and Diphylleia Michx., all of which belong to the Podophylloideae subfamily, but Dysosma can be distinguished by its perennial growth habit, creeping rhizomes, fibrous roots, 3–9-lobed or peltate leaves, umbellate inflorescences, and berries containing numerous seeds (Stähelin and Von 1991; Ying et al. 2011). Except for Podophyllum, the other three genera are distributed in China. Recent phylogenetic studies have resolved the relationships among Dysosma, Sinopodophyllum, Podophyllum, and Diphylleia, supporting that Dysosma is sister to a clade comprising Sinopodophyllum and Podophyllum (Loconte and Estes 1989; Nickol Academic editor: Andrey Erst Received: 7 March 2025 Accepted: 22 November 2025 Published: 15 December 2025 Citation: Huang L, Yao W-H, Yang Y-B, An M-T, Yu Y-L, Zhou M, Li H (2025) Dysosma xishuiensis (Berberidaceae), a new species from Guizhou, China, based on morphological and molecular evidence. PhytoKeys 268: 59–67. https://doi.org/10.3897/ phytokeys.268.152287 PhytoKeys 268: 59–67 (2025) DOI: 10.3897/phytokeys.268.152287 60 PhytoKeys 268: 59–67 (2025), DOI: 10.3897/phytokeys.268.152287 Lang Huang et al.: Dysosma xishuiensis, a new species from China 1995; Kim and Jansen 1998; Liu et al. 2002; Wang et al. 2007; Mao et al. 2014; Ye et al. 2018; Hsieh et al. 2022). Currently, nine species are recognized in Dysosma, including seven species described in the “Flora of China”. Additionally, D. villosa Z.W. Wang & H.C. Xi was recently described (Wang et al. 2019), and D. tonkinense (Gagnep.) M. Hiroe has been proposed as a distinct species (Pham et al. 2020). It is noteworthy that Shaw (2002) and Ying et al. (2011) indicated that three insufficiently studied taxa originally placed in Podophyllum, namely P. glaucescens J.M.H. Shaw, P. hemsleyi J.M.H. Shaw & Stearn, and P. trilobulum J.M.H. Shaw, may actually belong to Dysosma. Due to insufficient specimens for detailed study, the taxonomic status of these three taxa remains controversial. In medicine, plants of the genus Dysosma have been widely used as herbal remedies primarily for treating throat swelling and pain, venomous snake bites, bruises and sprains, lymph node inflammation, and related disorders (Woodson 1928; Ying et al. 2011; Mao et al. 2014; Pham et al. 2020). During our field investigation in Xishui National Nature Reserve, Xishui County, Guizhou Province, China (November 2021 and May 2022), we found an unusual population of Dysosma species. Initially, the plant seemed similar to D. versipellis in its stems (erect, unbranched, glabrous), leaves (alternate, blade suborbicular, to 25–30 cm in diameter, palmately lobed), and flowers (attached near the base of the blade, red). However, further examination revealed several key diagnostic features that differed from those of D. versipellis, such as a glabrous abaxial leaf surface, pedicels, and sepals; inflorescences bearing more flowers (8–20 vs. 5–8); larger obovate–oblong petals (2.8–3.4 × 1.5–2.0 cm vs. 2.5 × 0.8 cm); and an obpyriform ovary (vs. ellipsoid). To determine the taxonomic status of this taxon, we conducted systematic molecular studies using nuclear ITS and plastid markers (matK, rbcL) and inferred its position within Berberidaceae. Both morphological and molecular evidence support that this distinct population represents a novel species; thus, we formally describe it here. Materials and methods Morphological studies The morphological study of the new species was based on field observations of over 10 living individuals in Xishui National Nature Reserve. Comparison of morphological characteristics with four other closely related species was supplemented by high-resolution digitized herbarium specimens accessed via the Chinese Virtual Herbarium (CVH; https://www.cvh.ac.cn/) and JSTOR Global Plants (https://plants.jstor.org/). Detailed observations and measurements were performed to document key diagnostic features. Voucher specimens of the new species have been deposited in the Herbaria of the Forestry College, Guizhou University (GZAC); the Guizhou Academy of Forestry Science (GF); and the Kunming Institute of Botany, Chinese Academy of Sciences (KUN). Taxon sampling, DNA sequencing, and molecular analysis We collected one individual from each of the two natural populations within the protected area and dried the fresh leaf samples using silica gel. Genomic DNA was extracted from the dried leaves using a modified CTAB protocol 61 PhytoKeys 268: 59–67 (2025), DOI: 10.3897/phytokeys.268.152287 Lang Huang et al.: Dysosma xishuiensis, a new species from China (Doyle and Doyle 1987). DNA sequencing, primer design, and PCR amplification followed protocols from Mao et al. (2014). We retrieved ITS, matK, and rbcL sequences of 13 species (totaling 39 sequences) from GenBank to resolve the phylogenetic position of the new species. These encompassed representatives from Podophylloideae (Berberidaceae): Dysosma (seven species), Diphylleia (three species), Podophyllum (one species), and Sinopodophyllum (one species), with Achlys triphylla (Berberidaceae) serving as the outgroup. Corresponding GenBank accession numbers are presented in Table 1. All sequences were aligned using MAFFT v7.505 (Katoh and Standley 2013), trimmed with Gblocks 0.91b (Talavera and Castresana 2007), and concatenated to generate the final alignment matrix. Phylogenies were subsequently reconstructed using maximum likelihood (ML) and Bayesian inference (BI). ML analysis was implemented in IQ-TREE v1.6.12 (Nguyen et al. 2015) with the GTR+I+G4 substitution model selected by RAxML-NG v1.1 under the Bayesian Information Criterion (BIC). Branch support was assessed with 1,000 standard bootstrap replicates. BI analysis was performed in MrBayes v3.2.7a (Ronquist et al. 2012) under the SYM+G model. Two independent runs of 10,000,000 generations each were executed, sampling trees every 1,000 generations after a 25% burn-in. Four Markov chain Monte Carlo (MCMC) chains were employed, starting from random trees. Results and discussion Morphological comparison Through examination of type specimen descriptions and high-resolution images from JSTOR Global Plants (http://plants.jstor.org), Dysosma xishuiensis is morphologically distinguished from congeners by the following diagnostic Table 1. Species names and GenBank accession numbers of ITS, matK, and rbcL sequences used for analysis. Superscripts denote two distinct populations of the new species. Species ITS matK rbcL Achlys triphylla MG235275 MG593050 MG593050 Diphylleia cymosa KC494675 KC539368 KC539405 Diphylleia grayi KC494679 KC539373 KC539409 Diphylleia sinensis KC494674 KC539367 KC539403 Dysosma aurantiocaulis KC494665 KC539356 KC539395 Dysosma delavayi KC494672 KC539365 KC539402 Dysosma difformis KC494660 KC539359 KC539390 Dysosma majoensis KC494662 KC539353 KC539392 Dysosma pleiantha KC494652 KC539345 KC539382 Dysosma tsayuensis KC494668 KC539361 KC539398 Dysosma versipellis KC494658 KC539351 KC539388 Dysosma xishuiensis1PX138794 PX229892 PX229894 Dysosma xishuiensis2PX138795 PX229893 PX229895 Podophyllum peltatum KC494685 KC539378 KC539415 Sinopodophyllum hexandrum KC494684 KC539377 KC539413 62 PhytoKeys 268: 59–67 (2025), DOI: 10.3897/phytokeys.268.152287 Lang Huang et al.: Dysosma xishuiensis, a new species from China characters. D. xishuiensis shares similarities with D. versipellis in leaf shape, phyllotaxy, and floral attachment position. However, it is readily distinguishable by the following characters: glabrous abaxial leaf surfaces, pedicels, and sepals (vs. pubescent); 8–20-flowered inflorescences (vs. 5–8); petals obovate–oblong (vs. spatulate–obovate) and larger (2.8–3.4 × 1.5–2.0 cm vs. 2.5 × 0.8 cm); and an obpyriform ovary (vs. ellipsoid). Compared to D. majoensis, it is glabrous throughout (vs. puberulent on stems, abaxial leaf surfaces, and pedicels), with uniformly green leaf surfaces (vs. dark green adaxial and grey– purple abaxial surfaces) and obovate–oblong petals (vs. elliptic–lanceolate). Relative to D. pleiantha, it displays alternate leaves (vs. opposite), deeply divided leaves (vs. lobed), and flowers attached near the base of the blade (vs. attached near the base of the petiole) (Table 2). Phylogenetic relationships The combined ITS, matK, and rbcL datasets yielded a concatenated alignment of 2,051 bp. The phylogeny placed Dysosma xishuiensis in a strongly supported clade (PP = 1.00, BP = 99) with other Dysosma species (Fig. 1A, B), while its two geographically isolated populations comprised a monophyletic lineage, affirming taxonomic distinctiveness. It is noteworthy that the phylogenetic analysis revealed D. xishuiensis and D. difformis formed a sister-group relationship, collectively constituting a strongly supported clade (PP = 1.00, BP = 88) with D. pleiantha and D. versipellis. Morphologically, D. xishuiensis differs from D. pleiantha and D. versipellis as previously described and from D. difformis in leaf shape (suborbicular vs. obliquely peltate), pedicels (glabrous vs. sparsely white pubescent), and sepals (glabrous vs. pubescent externally). Table 2. Morphological comparison of Dysosma xishuiensis with four congeners. Data of the four Dysosma species are sourced from the “Flora of China”. Characters D. xishuiensis D. majoensis D. pleiantha D. versipellis D. difformis Plant height 80–120 cm tall ca. 50 cm tall 20–60 (–80) cm tall 40–150 cm tall 15–30 cm tall Leaf blade glabrous; palmately 5–8 deeply divided abaxially puberulent, deeply 4–6-divided, lobes 3-fid at apex glabrous; 5–9-lobed abaxially pubescent; palmately 4–9-lobed glabrous; entire or lobed Stem and Petiole glabrous puberulent glabrous glabrous glabrous Inflorescence attached near base of blade; 8–20-fascicled flowers attached near base of blade; 2–5 flowers attached near base of petiole; 5–8-fascicled flowers attached near base of blade, 5–8-fascicled flowers attached near base of blade. 2–5-fascicled flowers Pedicel ca. 5 cm, glabrous 1–3 cm, long puberulent 2–4 cm, glabrous length unknown, with pubescent 1–2 cm, sparsely white pubescen Sepals long elliptic, 1.5–2.0 cm × 5.0– 8.0 mm, glabrous elliptic, 7–15 mm, glabrous elliptic-oblong or ovate-oblong, 1–2 × ca. 0.8 cm oblong-elliptic, 0.6–1.8 cm × 3.0–8.0 mm, outside puberulent oblong-lanceolate, 2–2.5 cm × 2–5 mm, outside pubescent, inside glabrous Petal obovate-oblong, 2.8–3.4 × 1.5–2.0 cm elliptic-lanceolate, ca. 9 × 1.5 cm obovate-oblong, 3–4 × 1–1.3 cm spatulate-obovate, 2.5 × 0.8 cm oblong-loriform, 4–5 × 0.8–1 cm Pistil ovary obpyriform, ca.1.5 cm long; style ca. 3 mm long, stigma shield-shaped Ovary oblong; stigmas shield-shaped ovary oblong, ca. 1.3 cm; style ca. 3 mm ovary ellipsoid, glabrous; style short; stigma shieldshaped ovary urceolate; ca. 0.9 cm; style ca. 2 mm; stigma shield-shaped 63 PhytoKeys 268: 59–67 (2025), DOI: 10.3897/phytokeys.268.152287 Lang Huang et al.: Dysosma xishuiensis, a new species from China Taxonomic treatment Dysosma xishuiensis Y. B. Yang, M. T. An & C. H. Yang, sp. nov. urn:lsid:ipni.org:names:77373457-1 Fig. 2 Diagnosis. Dysosma xishuiensis is morphologically most similar to D. versipellis in its stems (erect, unbranched, glabrous), leaves (alternate, blade suborbicular, to 25–30 cm in diam, palmately lobed), flowers (attached near base of blade, Figure 1. The Bayesian inference phylogeny of Dysosma xishuiensis and its close relatives based on the nuclear DNA (ITS) and plastid gene regions (matK and rbcL). Bayesian inference posterior probability (PP ≥ 0.5; Fig. 1A) and maximum likelihood bootstrap percentages (BP ≥ 50%; Fig. 1B) are shown above the main branches. The samples of the new species are displayed in bold, with superscripts designating two distinct populations. 64 PhytoKeys 268: 59–67 (2025), DOI: 10.3897/phytokeys.268.152287 Lang Huang et al.: Dysosma xishuiensis, a new species from China red), but differs by the following characters: leaf abaxial surface, pedicels, and sepals glabrous (vs. pubescent); inflorescences bearing more flowers (8–20 vs. 5–8); larger obovate-oblong petals (2.8–3.4 × 1.5–2.0 cm vs. 2.5 × 0.8 cm); and ovary obpyriform (vs. ellipsoid). Type. China • Guizhou, Xishui County, Sanchahe Town, Xishui National Nature Reserve, alt. 975 m, under broadleaf forest beside the stream of Danxia landform, 20 May 2022, Yan-Bing Yang, Cheng-Hua Yang & He Li (holotype: GF!; isotype: GZAC!). • Xishui County, Tucheng Town, Tongyi Village, Xishui National Nature Reserve, alt. 1168 m, in the middle of a mountain under broadleaf forest of Danxia landform, 22 May 2022, Cheng-Hua Yang, He Li & Mao Li C1031 (isotype: KUN!). Figure 2. Images of living plants of Dysosma xishuiensis Y. B. Yang, M. T. An & C. H. Yang. A. Individual; B. Root; C. An adaxially leaf blade; D. Abaxial leaf blade; E. Inflorescence anatomy of a flower; F. Stamens and pistils; G. Plant habit and leaf morphology (D. versipellis); H. Inflorescence (D. versipellis); I. Leaf morphology (D. majoensis). Photos credit: A, F, G, I by C. H. Yang; C, D by L. Huang; B, E by J. G. Wang; G, H by Z. Wei. 65 PhytoKeys 268: 59–67 (2025), DOI: 10.3897/phytokeys.268.152287 Lang Huang et al.: Dysosma xishuiensis, a new species from China Description. Herbs, 80–120 cm tall. Rhizomes stout, densely fibrous. Aerial stems erect, pale green, unbranched, glabrous. Leaves alternate; petioles of lower leaves 10–20 cm, upper leaves ca. 3 cm; leaf blade suborbicular, up to 25 cm in diameter, thinly papery, glabrous on both surfaces, abaxially with prominent venation, palmately 5–8-lobed, lobes deeply divided (ca. 2/3 of radius); lobes obovate to obovate-oblong, margins remotely serrate, apex shallowly 3-lobed or bearing 2–3 mucronate teeth. Inflorescence an 8–20-flowered fascicle. Pedicels pendulous, slender, glabrous. Flowers dark red, attached near leaf base. Sepals long-elliptic, 1.5–2.0 × 0.5–0.8 cm, glabrous, apex acute. Petals oblong-obovate, 2.8–3.4 × 1.5–2.0 cm, glabrous. Stamens ca. 2 cm; filaments shorter than anthers; anther connective slightly prolonged, glabrous, acute. Pistil ca. 1.5 cm; ovary obpyriform, glabrous; style ca. 0.3 cm; stigma shieldshaped. Berry ellipsoid. Seeds numerous. Phenology. Flowering from April to June, fruiting from June to September. Distribution and habitat. Dysosma xishuiensis is only known from two localities in Xishui County, Guizhou, China, where it grows under the broadleaf forest by the stream or in the middle of a mountain of Danxia landform. Preliminary conservation status. Currently, Dysosma xishuiensis is documented exclusively in two discrete populations within Xishui National Nature Reserve. These populations occur in mid-subtropical to warm temperate zones characterized by a humid monsoon climate. The species exhibits a restricted distribution range, with both populations each comprising fewer than 100 mature individuals. In accordance with the International Union for Conservation of Nature (IUCN) Red List criteria B2 (a, b(iii)) (extent of occurrence < 500 km2, fewer than five locations, continuing decline in habitat quality) and D (population size < 250 mature individuals), we propose that D. xishuiensis be designated as Endangered (EN) (IUCN 2022). Etymology. The species epithet “xishuiensis” refers to the type locality of the new species. Vernacular name. The Chinese name is “xí shuĬ bā jiăo lián” (习水八角莲). Acknowledgements We thank the staff of the Guizhou Xishui Nature Reserve Management Bureau for assistance in fieldwork. We also thank Prof. Guo-Xiong Hu, Ph.D. candidate Chao Ye, and Rong-Rong Yan for their assistance with molecular data analysis. 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. 66 PhytoKeys 268: 59–67 (2025), DOI: 10.3897/phytokeys.268.152287 Lang Huang et al.: Dysosma xishuiensis, a new species from China Funding This study was funded by the Survey and Evaluation of Newly Added National Key Protected Wild Plant Resources in Guizhou Province (first phase) (MCHC-ZC20222009), the Guizhou Forestry Scientific Research Project (QLKHJZ [2024]07), and the Second Comprehensive Scientific Survey of the Guizhou Xishui National Nature Reserve. Author contributions A.M.T. conceptualized and led the project. Field investigation and sample collection were conducted by H.L., Y.Y.B., Y.Y.L., Z.M., and L.H. Y.Y.B. and A.M.T. examined and detailed the fine structure of the species represented in the study. The initial draft of the manuscript was prepared by H.L., Y.W.H., and Y.Y.B. All authors approved the final manuscript. Author ORCIDs Lang Huang https://orcid.org/0009-0004-8038-7432 Wei-Hao Yao https://orcid.org/0000-0002-5941-676X Yan-Bing Yang https://orcid.org/0009-0007-9014-1817 Ming-Tai An https://orcid.org/0000-0003-3886-0287 Yuan-Lin Yu https://orcid.org/0009-0005-5875-511X Mei Zhou https://orcid.org/0009-0000-3806-1333 He Li https://orcid.org/0009-0006-0597-3209 Data availability The newly obtained ITS, matK, and rbcL sequences have been submitted to NCBI. All other data that support the findings of this study are available in the main text. References Doyle JJ, Doyle JL (1987) A rapid DNA isolation procedure for small quantities of fresh leaf tissue. Phytochemical Bulletin 19: 11–15. Hsieh CL, Yu CC, Huang YL, Chung KF (2022) Mahonia vs. Berberis unloaded: Generic delimitation and infrafamilial classification of Berberidaceae based on plastid phylogenomics. 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