Paraphlomis leigongshanensis (Lamiaceae), a new species from Guizhou, China

23 Paraphlomis leigongshanensis (Lamiaceae), a new species from Guizhou, China Wei-Hao Yao1, Guo-Bin Jiang1, De-Hui Yu2, Sheng Chen3, Hong-Mei Chen1, Li Dai1, Bai-Qiu He4, He Li1 1 Key Laboratory of National Forestry and Grassland Administration on Biodiversity Conservation in Karst Mountainous Areas of Southwestern China, Guizhou Academy of Forestry, Guiyang, 550005, China 2 Guizhou Leigong Mountain National Nature Reserve Administration, Leishan, 557100, China 3 Institute of Biology, Guizhou Academy of Sciences, Guiyang, 550009, China 4 College of Life Sciences, University of Chinese Academy of Sciences, Beijing, 100049, China Corresponding author: Guo-Bin Jiang ([email protected]) Copyright: © Wei-Hao Yao 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 Paraphlomis leigongshanensis G.B. Jiang & W.H. Yao (Lamiaceae), a new species endemic to Leigong Mountain National Nature Reserve (Guizhou, China), is described and illustrated. Morphologically, it is most similar to P. jiangyongensis, but can easy be distinguished by its lamina shape, calyx teeth shape, and corolla color. A close relationship between the new species and P. gracilis var. lutienensis was revealed by molecular phylogenetic analyses based on ETS and ITS sequences, but they are morphologically distinct from each other. With only three known populations (ca. 200 mature individuals) in subtropical evergreen forests, the new species is assessed as Endangered (EN) under IUCN criteria B2ab(iii); D. Key words: Guizhou, Lamioideae, new taxon, phylogenetic analyses Introduction Paraphlomis (Prain) Prain (Lamiaceae, Lamioideae) is a member of the tribe Paraphlomideae Bendiksby, which was established to accommodate Paraphlomis, Matsumurella Makino, and Ajugoides Makino based on molecular phylogenetic evidence (Bendiksby et al. 2011; Zhao et al. 2021). Most species of Paraphlomis are distributed in southern China, with several taxa extending into the Himalayas, Korea, and Southeast Asia (Wu and Li 1977; Li and Hedge 1994a; Ko et al. 2014; Chen et al. 2021). Previous molecular phylogenetic studies revealed that Paraphlomis was not monophyletic, as species of Matsumurella were nested within it (Chen et al. 2021, 2022a; Guo et al. 2023). Recently, Yuan et al. (2024) treated Matsumurella and Ajugoides as synonyms of Paraphlomis based on plastome data and nuclear ribosomal DNA (nrDNA) sequences, as well as morphological evidence. Morphologically, Paraphlomis is characterized by its rhizomatous or stoloniferous habit, plants with simple hairs, nutlets with a truncate apex, and actinomorphic calyces (Wu and Li 1977; Bendiksby et al. 2011; Chen et al. 2021; Yuan et al. 2024). According to the Catalogue of Life China 2025 Annual Checklist and recent references, Paraphlomis Academic editor: Chun-Lei Xiang Received: 4 May 2025 Accepted: 13 October 2025 Published: 2 December 2025 Citation: Yao W-H, Jiang G-B, Yu D-H, Chen S, Chen H-M, Dai L, He B-Q, Li H (2025) Paraphlomis leigongshanensis (Lamiaceae), a new species from Guizhou, China. PhytoKeys 267: 23–35. https://doi.org/10.3897/ phytokeys.267.157814 PhytoKeys 267: 23–35 (2025) DOI: 10.3897/phytokeys.267.157814 24 PhytoKeys 267: 23–35 (2025), DOI: 10.3897/phytokeys.267.157814 Wei-Hao Yao et al.: A new species from Guizhou Province, China comprises 51 species and six varieties (Li and Hedge 1994a; Chen et al. 2021; Yuan et al. 2024, 2025; Liu et al. 2025). During a botanical field survey in Leigong Mountain National Nature Reserve, Guizhou Province, in 2024, we discovered an unknown species of Paraphlomis. This species is distinguished by its dwarf habit, a white corolla with pink stripes, and hairy mericarps. After careful field observations, morphological comparisons with other members of Paraphlomis, and molecular phylogenetic analyses, we confirmed its specific status and taxonomic placement within the genus. We hereby named it as P. leigongshanensis G.B. Jiang & W.H. Yao. Materials and methods Morphological study During February 2024 and July 2025, four field surveys were carried out at the type locality of the new species. Specimens from several herbaria were examined, including GYBG, GZTM, HGAS, HIB, IBK, IBSC, KUN, NAS, and PE (herbarium abbreviations follow the conventions outlined by Thiers (2025)). Additionally, we consulted several online databases, including the China Plant Photography Database (http://ppbc.iplant.cn/), the Chinese Field Herbarium (http://www.cfh.ac.cn/), and Global Plants (http://plants.jstor.org/), to obtain supplementary information on species distributions and morphological data. To distinguish the new species from other species of Paraphlomis, we conducted detailed morphological comparisons, focusing on key characteristics such as stems, leaves, calyx, corolla, mericarps, and trichomes. All measurements were made using a dissecting microscope. The morphological description followed relevant taxonomic and floristic literature on Paraphlomis, with terminology standardized according to Li and Hedge (1994b), ensuring consistent and accurate morphological interpretation. Molecular phylogenetic analyses For the molecular phylogenetic reconstruction of Paraphlomis, we included a total of 53 samples: 38 representing Paraphlomis taxa (34 species and four varieties) and two outgroup taxa. The outgroups comprised Phlomoides dentosa var. glabrescens (Danguy) C.L. Xiang & H. Peng and Phlomis fruticosa L. Only one accession of the new species was sampled and sequenced in this study, while sequence data for other species were obtained from GenBank and previous studies (Chen et al. 2021, 2022a, 2022b). Genomic DNA was extracted from dried leaf tissue using a modified CTAB method (Doyle and Doyle 1987). Previous studies have shown that phylogenetic trees based on nuclear ribosomal DNA internal and external transcribed spacers (nrITS and nrETS) offer more effective insights into the phylogenetic relationships of Paraphlomis than those constructed using chloroplast DNA (Chen et al. 2021; Yuan et al. 2024). Thus, only the nrDNA data was used here for phylogenetic reconstruction. Voucher information and GenBank accession numbers for all sequences are provided in Appendix 1. The best-fitting substitution model was selected using the Akaike Information Criterion (AIC) based on sequence data in jModelTest v.2.1.7 (Darriba et al. 2012). Phylogenetic analyses were performed using Bayesian Inference (BI) 25 PhytoKeys 267: 23–35 (2025), DOI: 10.3897/phytokeys.267.157814 Wei-Hao Yao et al.: A new species from Guizhou Province, China and Maximum Likelihood (ML) methods, employing MrBayes (Ronquist et al. 2012) and RAxML v.8.2.9 (Stamatakis 2014) on the CIPRES Science Gateway v.3.3 (Miller et al. 2010), respectively. The Markov chain Monte Carlo algorithm was run for 20 million generations with four incrementally heated chains starting from random trees and sampling one out of every 1,000 generations. The initial 25% of sampled trees were discarded as burn-in and the 50% majority-rule consensus tree was calculated from remaining trees with nodal support summarized as posterior probabilities (PP). All phylogenetic trees were visualized and edited for clarity using FigTree v.1.4.2 (Rambaut 2014), with PP and bootstrap support (BS) values annotated on the branches. Results and discussion Morphological comparison Morphological comparisons were based on an examination of 196 specimens representing 14 species of Paraphlomis. A key diagnostic character of the potential new species P. leigongshanensis is the hairy mericarps, which serves as an important taxonomic feature for the infrageneric classification (Chen et al. 2021; Yuan et al. 2024). Morphologically, P. leigongshanensis is most similar to P. jiangyongensis in the dwarf habit ascending stems, and hairy mericarps (Table 1). However, it differs from P. jiangyongensis by its abaxially purple (vs. green) laminae, calyx teeth acuminate (vs. acute) at apex, and longer corolla tubes. Additionally, the corolla of P. leigongshanensis is white with pink stripes and spots, with the markings concentrated on the lower lip, differing from the pale yellow corolla lips with red spots in P. jiangyongensis. This species is distributed in the mid-subtropical evergreen broad-leaved forests in Leishan County, Guizhou, exhibiting geographical isolation from P. jiangyongensis (Appendix 2: Fig. A1). Molecular phylogenetic analyses The concatenated nrDNA dataset had a total length of 1259 bp, comprising 814 bp from the ITS region and 445 bp from the ETS region. The dataset contained 356 variable sites (168 from ITS, 188 from ETS) and 205 parsimony-informative characters (89 from ITS, 116 from ETS). The resulting phylogenetic tree of Paraphlomis was consistent with previous studies (Chen et al. 2021; Yuan et al. 2024). The potential new species P. leigongshanensis was resolved as sister to P. gracilis var. lutienensis (Y. Z. Sun) C. Y. Wu (Fig. 1: BS = 100%/PP = 1.00), and the P. leigongshanensis–P. gracilis var. lutienensis clade was further sister to P. gracilis var. gracilis Kudô (Fig. 1: BS = 100% / PP = 1.00). The three taxa were all nested within the “Clade III” proposed by Chen et al. (2021) and Yuan et al. (2024) (Fig. 1: BS = 83%/PP = 1.00). Species in this clade (highlighted in blue in Fig. 1) are all characterized by hairy mericarps (Chen et al. 2021; Yuan et al. 2024). Despite the close relationships and geographical affinities (Appendix 2: Fig. A1) among P. leigongshanensis, P. gracilis var. gracilis, and P. gracilis var. lutienensis, P. leigongshanensis is morphologically distinct from the latter two taxa. Specifically, P. leigongshanensis exhibits a creeping habit with stems 5–10 cm tall, ovate to elliptic leaves, reddish-purple calyces, sessile flowers, and white corollas with pink stripes and spots. In contrast, P. gracilis var. gracilis 26 PhytoKeys 267: 23–35 (2025), DOI: 10.3897/phytokeys.267.157814 Wei-Hao Yao et al.: A new species from Guizhou Province, China * / * 0.54 / <50 0.98 / 51 0.94 / 52 * / 66 0.55 / <50 * / 83 0.96 / 63 0.70 / 51 0.65 / <50 * / 78 * / 95 * / 94 * / 98 Paraphlomis octopus 2 Paraphlomis octopus 1 Paraphlomis octopus 3 * / 98 Paraphlomis pagantha 1 Paraphlomis pagantha 2 Paraphlomis kwangtungensis * / * Paraphlomis jiangyongensis 1 Paraphlomis jiangyongensis 2 * / * Paraphlomis hsiwenii 1 Paraphlomis hsiwenii 2 * / * * / * Paraphlomis subcoriacea 1 Paraphlomis subcoriacea 2 Paraphlomis baiwanensis 1 Paraphlomis baiwanensis 2 * / * Paraphlomis qingyuanensis 1 Paraphlomis qingyuanensis 2 * / 99 Paraphlomis albida var. albida Paraphlomis albida var. brevidens Paraphlomis yangsoensis * / * * / * Paraphlomis leigongshanensis sp. nov. Paraphlomis gracilis var. lutienensis * / * Paraphlomis gracilis var. gracilis 1 Paraphlomis gracilis var. gracilis 2 Paraphlomis longicalyx * / 84 * / 93 * / * * / * Paraphlomis intermedia Paraphlomis lancidentata Paraphlomis reflexa * / 99 * / 98 Paraphlomis nana 1 Paraphlomis nana 2 Paraphlomis albiflora * / 92 * / 99 Paraphlomis lanceolata 1 Paraphlomis lanceolata 2 Paraphlomis montigena 0.57 / <50 * / * Paraphlomis chinensis 1 Paraphlomis chinensis 2 Paraphlomis strictiflora Paraphlomis youyangensis Paraphlomis foliata Paraphlomis yingdeensis Paraphlomis membranacea * / 79 * / * * / 94 Paraphlomis javanica var. javanica 1 Paraphlomis javanica var. pteropoda Paraphlomis javanica var. javanica 2 * / * Paraphlomis paucisetosa 1 Paraphlomis paucisetosa 2 * / 99 Paraphlomis hispida Paraphlomis patentisetulosa * / 87 Paraphlomis coronata 2 Paraphlomis coronata 1 * / 97 Paraphlomis hirsutissima Paraphlomis caloneura Phlomis fruticosa Phlomoides dentosa var. glabrescens 0.01 Figure 1. Bayesian 50% majority-rule consensus tree of Paraphlomis based on combined nuclear (ITS and ETS) data set. Support values ≥ 0.50 PP or 50% BS are displayed above the branches (an “*” indicates a support value of 1.00 PP or 100% BS). Multiple accessions of the same species are numbered according to Appendix 1 27 PhytoKeys 267: 23–35 (2025), DOI: 10.3897/phytokeys.267.157814 Wei-Hao Yao et al.: A new species from Guizhou Province, China has erect and much taller stems (up to 1 m tall), lanceolate leaves, green calyces, subsessile flowers, and yellowish green to white corollas with purple spots (Table 1). Paraphlomis gracilis var. lutienensis differs from both species by its subsessile and narrowly lanceolate laminae and yellow corollas. Moreover, a recent phylogenetic study based on plastome data (Yuan et al. 2024) indicated that the phylogenetic relationship between P. gracilis var. gracilis and P. gracilis var. lutienensis is consistent with the results of our study. Based on our phylogenetic results, although P. leigongshanensis is grouped with P. gracilis var. gracilis and P. gracilis var. lutienensis, there is sufficient evidence to support its recognition as a Table 1. Morphological comparisons among P. leigongshanensis, P. jiangyongensis, P. gracilis var. gracilis, and P. gracilis var. lutienensis. Character P. leigongshanensis P. jiangyongensis P. gracilis var. gracilis P. gracilis var. lutienensis Stem Prostrate to ascending, 5–10 cm tall Ascending, 5–30 cm tall Erect, 30–100 cm tall Erect, 30–100 cm tall Lamina Ovate to long elliptic, 2.5–7 × 2–5 cm, abaxially purple, densely villose Ovate to subcircular, 3–8 × 2–5 cm, abaxially light green, densely strigose Lanceolate, 5–10 × 1.7– 3.3 cm, abaxially green, densely strigose and glandular Lanceolate, 6.5–10.5 × 1.7–2.5 cm, abaxially green, densely strigose and glandular, subsessile and narrowly lanceolate Verticillaster 2–10-flowered 2–8-flowered (2–) 4–8 (–12)-flowered (2–) 4–8 (–12)-flowered Pedicel Sessile Sessile Subsessile Ca. 4 mm long Calyx Purplish–red, densely villose Green, densely strigose Green, densely retrorse strigose Green, densely retrorse strigose Calyx teeth 4–8 mm long, triangular, apex acuminate Ca. 2 mm long, triangular, apex acute Ca. 6 mm long, triangular, apex acuminate or subulate Ca. 4 mm long, triangular, apex acuminate Corolla 12–17 mm long, white lower lip with pink stripes and spots 7.5–10 mm long, pale yellow, lower lip with red spots 15 mm long, upper yellow, white or yellowish green, lower lip white with purple spots or yellow with red spots 15 mm long, yellow, lower lip with pink spots distinct species. The morphological differences further reinforce its taxonomic distinctiveness and provide additional support for its designation as a new species. Taxonomic treatment Paraphlomis leigongshanensis G.B.Jiang & W.H.Yao, sp. nov. urn:lsid:ipni.org:names:77365161-1 Figs 2, 3; Appendix 2: Figs A1, A2 雷公山假糙苏 Type. China • Guizhou Province: Qiandongnan Miao and Dong Autonomous Prefecture, Leishan County, Leigong Mountain National Nature Reserve, 26°24'30.91"N, 108°15'53.77"E, alt. 1104 m, 23 Aug. 2024, G.B. Jiang, S. Chen, H.M. Chen & D.H. Yu GB2024156 (holotype: GF 09040011!; isotypes: KUN 1644009!, HGAS 0128505!). Diagnosis. Paraphlomis leigongshanensis is morphologically most similar to P. jiangyongensis but can be distinguished by its calyx purplish red (vs. green), apex of calyx teeth lanceolate (vs. acute), corolla white with pink stripes (vs. pale yellow with red spots), and corolla tube 12–17 mm long (vs. 7.5–10 mm long). Description. Perennial herb, 5–10 cm tall, stoloniferous. Stems simple, prostrate to ascending, slightly 4-angled, densely villose. Leaves opposite, ovate to elliptical, papery, 2.5–8 × 2–5 cm; margin shallowly crenate, apex obtuse 28 PhytoKeys 267: 23–35 (2025), DOI: 10.3897/phytokeys.267.157814 Wei-Hao Yao et al.: A new species from Guizhou Province, China Figure 2. Paraphlomis leigongshanensis from the type locality. A. Habitat; B-D. Habit; E. Inflorescence; F. Flowering condition; G. Adaxial view of leaf; H. Abaxial view of leaf; I. Stem; J. Calyces; K. Dissected corolla and pistil; L. Corollas; M. Dissected calyx and mericarps; N. Dried mericarps. (Photographed by Guo-Bin Jiang). 29 PhytoKeys 267: 23–35 (2025), DOI: 10.3897/phytokeys.267.157814 Wei-Hao Yao et al.: A new species from Guizhou Province, China to subrounded, base broadly cuneate to rounded; adaxial surface green, abaxial purple, densely villose on both surfaces, particularly along veins, lateral veins 3–5-paired; petioles 0.5–3.5 cm long, densely villose. Verticillasters 2–10-flowered, sessile; bracteoles minute, early deciduous. Calyx purplish red, tubular-campanulate, 10–15 mm long, densely villose outside, glabrous inside; teeth 5, subequal, triangular, erect, 4–8 mm long, apex acuminate. Corolla white with pink stripes and spots, 2–3 cm long, densely pubescent outside; tube 1.2– 1.7 cm long, slightly dilated at throat, pubescent annulate inside at 1/3 distance from base; bilabiate, upper lip pink, oblong, entire, erect, concave, 8–10 mm long, ca. 5 mm wide, lower lip pink, 3-lobed, medium lobe subcircular with pink stripes and spots, apex slightly emarginate, 6–7 mm wide, lateral lobes ovate, ca. 3.5 mm wide. Stamens 4, straight, included; filaments flat, pubescent at base; anther cells 2, parallel. Style included, glabrous, apex subequally 2-lobed, lobes subulate. Ovary densely pubescent and glandular at apex. Mericarps 4, dark brown, triquetrous-oblong, ca. 3.6 mm long, densely pubescent at apex. Distribution, habitat and conservation status. Paraphlomis leigongshanensis is currently known from only three populations in adjacent valleys approximately 100 m apart, within Leigong Mountain National Nature Reserve, Guizhou, China (Appendix 2: Fig. A1). The total population comprises about 200 mature individuals. The species occurs in the subtropical monsoon climate zone, with populations confined to the area near Getou Village, and is vulnerable to anthropogenic disturbances. Based on the IUCN Red List criteria B2(a, b(iii)) (extent of occurrence Figure 3. Line drawing of Paraphlomis leigongshanensis. A. Plant; B. Flower; C. Dissected corolla and pistil; D. Calyx; E. Dissected calyx; F. Nutlets. (Drawn by Hong-Mei Chen). 30 PhytoKeys 267: 23–35 (2025), DOI: 10.3897/phytokeys.267.157814 Wei-Hao Yao et al.: A new species from Guizhou Province, China < 500 km2; < 5 locations; continuing habitat degradation), D (Estimated population size < 250 mature individuals), we proposed an Endangered (EN) assessment (IUCN 2024). Since similar adjacent habitats may harbor undiscovered populations, available evidence did not support Critically Endangered (CR) status. Phenology. Flowering was observed from May to August, and fruiting from August to October. Etymology. The specific epithet “leigongshanensis” is derived from the type locality of the new species, i.e., Leigong Mountain National Nature Reserve, Leishan County, Guizhou Province, China. Chinese name (assigned here). léi gōng shān jiă cāo sū (雷公山假糙苏). The name means a species of Paraphlomis only found in Leigong Mountain National Nature Reserve, Guizhou Province, China. Additional specimens examined (paratypes). China • Guizhou Province: Qiandongnan Miao and Dong Autonomous Prefecture, Leishan County, Leigong Mountain National Nature Reserve, 26°24'19.92"N, 108°15'19.20"E, alt. 1100 m, 1 Jul. 2025, S. Chen & L. Hao GB2025015 (GF 09040012!, KUN 1644008!). Acknowledgements We are grateful to Dr. Ya-Ping Chen of the Kunming Institute of Botany, Chinese Academy of Sciences, for conducting the phylogenetic analysis. We thank TaoHua Yuan (PhD candidate, Tibet University), Xiao-Kai Xiong (Master’s candidate, Guizhou University), and Jiang-Miao Gu (Master’s candidate, Guizhou University) for their suggestions on the 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 Survey and Assessment of Newly Added National Key Protected Wild Plant Resources in Guizhou Province (Three stage) (MCHCZD20242057), Guizhou Provincial Basic Research Program (Qiankehe Fundamentals-ZK [2023] General 234), and Guizhou Provincial Basic Research Program (Qiankehe Fundamentals-MS [2025] 327). Author contributions Guo-Bin Jiang conceptualized and led the project. Field investigation and sample collection were conducted by Guo-Bin Jiang, Sheng Chen, De-Hui Yu, Hong-Mei Chen, Li Dai, and Wei-Hao Yao. Wei-Hao Yao, Guo-Bin Jiang, Hong-Mei Chen, and Bai-Qiu He examined and detailed the fine structure of the species represented in the study. The initial 31 PhytoKeys 267: 23–35 (2025), DOI: 10.3897/phytokeys.267.157814 Wei-Hao Yao et al.: A new species from Guizhou Province, China draft of the manuscript was prepared by Wei-Hao Yao and Guo-Bin Jiang. All authors approved the final manuscript. Author ORCIDs Wei-Hao Yao https://orcid.org/0009-0001-8231-0867 Guo-Bin Jiang https://orcid.org/0000-0002-4631-7906 De-Hui Yu https://orcid.org/0009-0008-4264-8995 Sheng Chen https://orcid.org/0000-0002-2727-6504 Hong-Mei Chen https://orcid.org/0009-0002-2368-7813 Li Dai https://orcid.org/0009-0004-3907-8582 Bai-Qiu He https://orcid.org/0009-0009-7491-3586 He Li https://orcid.org/0009-0006-0597-3209 Data availability The newly-obtained ITS and ETS sequences have been submitted to the NCBI (accession numbers: PV476761; PV478008). All of the other data that support the findings of this study are available in the main text. References Bendiksby M, Thorbek L, Scheen AC, Lindqvist C, Ryding O (2011) An updated phylogeny and classification of Lamiaceae subfamily Lamioideae. Taxon 60(2): 471–484. https://doi.org/10.1002/tax.602015 Chen YP, Liu A, Yu XL, Xiang CL (2021) A preliminary phylogenetic study of Paraphlomis (Lamiaceae) based on molecular and morphological evidence. 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