Abstract
Paraphlomisyingdeensis (Lamiaceae), a new species from the limestone area in northern Guangdong Province, China, is described and illustrated. Phylogenetic analyses, based on two nuclear DNA regions (ITS and ETS) and three plastid DNA regions (rpl32-trnL, rps16 and trnL-trnF), suggest that P.yingdeensis represents a distinct species in Paraphlomis. Morphologically, P.yingdeensis is similar to P.foliatasubsp.montigena and P.nana, but can be distinguished from the former by its densely villous lamina and calyx, not decurrent base of lamina and bristle-like-acuminate apex of calyx teeth, and distinguished from the latter by its significantly taller plant (15–20 cm vs. 1–5 cm) and larger lamina (6.2–16.5 × 4–11.5 vs. 2–7 × 1.5–4 cm), densely villous stem, lamina and calyx and yellow corolla.
Keywords: endemics, limestone, new taxon, Paraphlomideae, phylogeny
Introduction
As a member of tribe Paraphlomideae (Lamiaceae, Lamioideae) (Bendiksby et al. 2011; Zhao et al. 2021), the genus Paraphlomis Prain is characterised by its herbaceous habit, actinomorphic calyx with five lobes less than half as long as the tube, corolla 2-lipped (1/3) with hairy upper lip, but hardly bearded along the margin, included stamens and an apically truncate ovary (Wu and Li 1977; Bendiksby et al. 2011; Chen et al. 2021). A total of 36 species and seven varieties are recognised within Paraphlomis, most of which are distributed in southern China (Chen et al. 2022b; Yuan et al. 2022), with several species occurring in the Himalayas, Korea and Southeast Asia (Li and Hedge 1994; Ko et al. 2014; Chen et al. 2021). Many species of Paraphlomis are endemics of limestone soils, including the recently described P.kuankuoshuiensis R.B. Zhang, D. Tan & C.B. Ma (Zhang et al. 2020), P.longicalyx Y.P. Chen & C.L. Xiang (Chen et al. 2022a) and P.hsiwenii Y.P. Chen & X. Li (Chen et al. 2022b). This shows species richness of Paraphlomis has been quite underrated and more field investigations are needed to infer its diversity in limestone areas.
The botanical expedition to the Shimentai National Nature Reserve in Guangdong Province, China in October 2021, showed an unknown species of Paraphlomis. Based on other field observations (from May to August in 2022), morphological comparisons with congeneric species, as well as molecular phylogenetic studies, we confirmed that it represented a new species, here described and illustrated.
Materials and methods
Morphological study
Field observations and collections of the new species were carried out from May to August in 2022 in Boluo Town of Yingde City in northern Guangdong Province, China. Morphological comparisons of the putative new species with other Paraphlomis species were conducted firstly by consulting relevant taxonomic literature, included “Flora of China” (Li and Hedge 1994), “Flora of Guangdong” (Luo 1995) and other recently described species and infraspecies of Paraphlomis (Yan and Fang 2009; Ding et al. 2019; Zhang et al. 2020; Chen et al. 2021, 2022a, b, c; Zhao et al. 2022). We also carried out a check of herbarium specimens deposited in LBG, AU, IBK, FJFC, PE, ANUB, KUN, FJSI and SYS (herbarium acronyms following Thiers 2022). All morphological characters were measured using dissecting microscopes.
Phylogenetic analyses
Previous molecular phylogenetic study revealed genus Paraphlomis is not monophyletic, because species of Matsumurella were recovered within it (Chen et al. 2021; Chen et al. 2022b). Thus, Matsumurella was also included in our phylogenetic analyses. A total of 37 accessions, representing 20 species and four varieties/subspecies of Paraphlomis and two Matsumurella species were selected as ingroups. One species each of Phlomis L. and Phlomoides Moench were included as outgroups following Chen et al. (2022a, b). Except for the three accessions of the new species that were newly sampled here, sequences of the remaining accessions were all retrieved from our previous studies (Chen et al. 2021, 2022a, b, c). Genomic DNA of the potential new species was extracted from silica-gel-dried leaves using the modified 2× CTAB procedure of Doyle and Doyle (1987). We selected five DNA markers for the phylogenetic reconstruction, including two nuclear ribosomal regions (internal and external transcribed spacers, i.e. ITS and ETS) and three plastid DNA regions (rpl32-trnL, rps16 and trnL-trnF). Primers used for the polymerase chain reaction (PCR) amplification and sequencing were the same as those of Chen et al. (2021), while PCR procedures followed those described in Chen et al. (2016). The specimen information of samples and GenBank accession numbers for all sequences are listed in Appendix 1.
Raw sequences were assembled and edited using Sequencher 4.1.4 (Gene Codes, Ann Arbor, MI, USA) and then aligned using MUSCLE (Edgar 2004) and manually adjusted in MEGA 6.0 (Tamura et al. 2013). Bayesian Inference (BI) (Ronquist et al. 2012) and Maximum Likelihood (ML) (Stamatakis 2014) analyses were used for phylogenetic reconstruction and detailed settings for the two analyses followed those described in Chen et al. (2021). The resulting trees with posterior probabilities (PP) and Bootstrap support (BS) values were visualised and annotated in TreeGraph 2 (Stöver and Müller 2010). The combined nuclear dataset and the combined plastid dataset were initially analysed separately. Topological incongruence between the two reconstructions was visually inspected, based on the thresholds of PP ≥ 0.95 and/or BS ≥ 70%. After excluding the taxa that exhibited strong conflicts between the nuclear tree and the plastid tree, the combined nuclear dataset and the combined plastid dataset were then concatenated for phylogenetic analyses.
Results and discussion
The aligned length of the combined nuclear dataset was 1254 bp (810 bp for ITS, 444 bp for ETS) and that of the combined plastid dataset was 2479 bp (850 bp for rpl32-trnL, 812 bp for rps16, 817 bp for trnL-trnF). Since the placements of three taxa, Paraphlomisalbiflora (Hemsl.) Hand.-Mazz., P.nana Y.P. Chen, C. Xiong & C.L. Xiang and P.javanicavar.pteropoda D. Fang & K.J. Yan, showed hard incongruences in the nuclear tree (Appendix 2) and the plastid tree (Appendix 3), these taxa were excluded prior to the combination of the nuclear and plastid datasets. All the resulting trees (Fig. 1; Appendices 2–3) were topologically consistent with those in previous study (Chen et al. 2021). With the two species of Matsumurella deeply nested within Paraphlomis, both genera were shown to be non-monophyletic. The three individuals of the putative new species formed a strongly supported clade (Fig. 1: PP = 1.00 / BS = 100%), but its relationship with other species of Matsumurella-Paraphlomis was not resolved.
Figure 1.
Optimal Maximum Likelihood tree of Paraphlomis inferred from combined nuclear (ETS and ITS) and plastid (rpl32-trnL, rps16 and trnL-trnF) dataset. Support value ≥ 50% BS or 0.50 PP are displayed above the branches (“-” indicates a support value < 0.50 PP).
Our morphological study revealed that the new species P.yingdeensis W.Y.Zhao, Y.Q.Li & Q.Fan is most similar to P.foliatasubsp.montigena X.H. Guo & S.B. Zhou and P.nana for some morphological characters as they have short habits and triangular-laceolate calyx teeth with apices acuminate or bristle-like-acuminate. Paraphlomisfoliatasubsp.montigena was classified by Guo (1993) as a subspecies of P.foliata (Dunn) C.Y. Wu & H.W. Li. However, previous molecular phylogenetic studies (Chen et al. 2022b, c) and our present analyses (Fig. 1; Appendices 2–3) indicated that P.foliatasubsp.montigena might represent an independent species within the genus as it is distantly related to P.foliatasubsp.foliata. The new species can be distinguished from P.foliatasubsp.montigena in the morphology and indumentum of laminae and calyces. Both the laminae and calyces are densely villous in P.yingdeensis, but are sparsely strigose in P.foliatasubsp.montigena; the base of lamina is broadly cuneate and not decurrent in the new species, but is cuneate and decurrent in P.foliatasubsp.montigena; P.yingdeensis has bristle-like-acuminate apex of calyx teeth, in contrast, the apex of calyx teeth of P.foliatasubsp.montigena is acuminate. The phylogenetic placement of P.nana was conflicting in the nuclear tree and plastid tree, but it was consistently sister to P.albiflora (Appendices 2–3). Both P.nana and P.yingdeensis have translucent and membranous calyces with bristle-like-acuminate apex of calyx teeth. The two species mainly differ in the height of plants, size and indumentum of laminae, as well as colour of corollae. Specifically, plants of P.nana are 1–5 cm tall, whereas those of P.yingdeensis are 10–20 cm tall. The stems and laminae are densely villous in P.nana, but are densely strigose in P.yingdeensis. Moreover, P.yingdeensis has significantly larger laminae than P.nana (6.2–16.5 × 4–11.5 cm vs. 2–7 × 1.5–4 cm) and the corollae of P.yingdeensis are yellow, differing from the white corollae of P.nana. Detailed morphological comparisons amongst the three taxa were summarised in Table 1.
Table 1.
Morphological comparisons amongst Paraphlomisyingdeensis, P.foliatasubsp.montigena and P.nana.
Characters | P.yingdeensis | P.foliatasubsp.montigena | P.nana |
---|---|---|---|
Stem | 10–20 cm tall, densely villous | 15–20 cm tall, densely villous | 1–5 cm tall, densely retrorse strigose |
Lamina | 6.2–16.5 × 4–11.5 cm, base broadly cuneate, not decurrent, densely villous | 5–16 × 2.5–6.5 cm, base cuneate, decurrent, sparsely strigose | 2–7 × 1.5–4 cm, base cuneate to broadly cuneate, decurrent, densely to sparsely strigose |
Calyx | Densely villous outside, teeth 3–4 mm long, apex bristle-like-acuminate | Sparsely strigose outside, teeth ca. 2.5 mm long, apex acuminate | Appressed strigose outside, teeth ca. 3 mm long, apex bristle-like-acuminate |
Corolla | yellow | yellow | white |
Geographically, P.foliatasubsp.montigena is restricted to the Qingliangfeng Nature Reserve at the border area of Zhejiang and Anhui Provinces in eastern China (Guo 1993) and P.nana is now only known from Chongqing City in central China (Chen et al. 2022c). Both the two species are not karst-adapted. In contrast, the new species is distributed in the limestone area in Guangdong Province, southern China.
Taxonomic treatment
. Paraphlomis yingdeensis
W.Y.Zhao, Y.Q.Li & Q.Fan sp. nov.
2B9BE82B-19BE-5389-B061-A4A45A2AFD06
urn:lsid:ipni.org:names:77313387-1
Figs 2 , 3 , 4 Chinese name: 英德假糙苏
Figure 2.
Paraphlomisyingdeensis from the type locality A habitat B, C plants D stem. (Photographs: A, C, D by W.-Y. Zhao; B by Y.-Q. Li).
Figure 3.
Floral traits of ParaphlomisyingdeensisA, B inflorescences C frontal view of flower D lateral view of flower E corolla and dissected calyx (inner view) F pistil and stamens G anthers H ovary. (Photographs: A, B by Y.-Q. Li; C–H by W.-Y. Zhao).
Figure 4.
Line drawing of ParaphlomisyingdeensisA plant B transverse section of stem C pistil D frontal view of flower E dissected calyx (outside view) F dissected corolla G lateral view of flower. (Drawn by Rong-En Wu).
Type.
China. Guangdong Province: Yingde City, Boluo Town, on the way from Xinzhai Village to Changshan Village, on the limestone cliff at the roadside, 24°24'N, 113°0'E, alt. 61 m, 29 May 2021, Zhao Wan-Yi, Li Yuan-Qiu, Pan Jia-Wen & Yang Ling-Han ZWY-2092 (holotype: SYS00236856! isotypes: KUN!, SYS00236857!, SYS00236858!, SYS00236859!)
Diagnosis.
Paraphlomisyingdeensis is morphologically similar to P.foliatasubsp.montigena and P.nana, but differs from the former in its lamina and calyx densely villous (vs. sparsely strigose), base of lamina not decurrent (vs. decurrent) and apex of calyx teeth bristle-like-acuminate (vs. acuminate) and from the latter in its plants 10–20 cm tall (vs. 1–5 cm tall), lamina 6.2–16.5 × 4–11.5 cm and densely villous (vs. 2–7 × 1.5–4 cm and densely strigose) and corolla yellow (vs. white).
Description.
Herbs perennial, 10–20 cm tall. Rhizomes short; roots fibrous, yellowish-brown, glabrous. Stems erect or prostrate, 4-angled, green (young branch) to purplish-red, densely villous. Leaves opposite, leafless towards base, upper two pairs crowded and rosulate; petiole 0.3–2.5 cm long, densely villous; lamina obovate, papery, 6.2–16.5 cm long, 4–11.5 cm wide, apex obtuse, base broadly cuneate, margin crenate-serrate; adaxially green, abaxially light green, densely villous on both sides; lateral veins 5–7-paired, obviously raised abaxially. Verticillasters in compact, sometimes capitate-like inflorescences, 8–16-flowered, 2.2–3.0 cm in diam.; bracteoles lanceolate to linear, 7–8 mm long, densely villous. Calyx light green, translucent, membranous, campanulate, 6–7 mm long, densely villous outside, glabrous inside, conspicuously 10-veined; teeth 5, subequal, triangular lanceolate, 3–4 mm long, apex bristle-like-acuminate. Corolla yellow, 1.5–1.8 cm long; tube 1.0–1.1 cm long, ca. 1.5 mm in diam., straight, pubescent annulate in throat inside; 2-lipped, villous outside, upper lip oblong, margin entire, erect, ca. 6 mm long, ca. 3.5 mm wide; lower lip spreading or reflexed, 4–5 mm long, 3-lobed, medium lobe suborbicular, apex emarginate, lateral lobes oblong, apex obtuse. Stamens 4, inserted above middle and upper of corolla tube, straight, included, filaments flat, sparsely puberulent-villous; anther cells 2, ovoid, glabrous. Style filiform, included, glabrous, apex subequally 2-lobed. Ovary 4-loculed, glabrous. Nutlets not seen.
Distribution and habitat.
Currently, only one population of P.yingdeensis was found in Boluo Town, Yingde City, in northern Guangdong Province. This town was located in the subtropical monsoon climate region, with development of a large area of karst landform. Paraphlomisyingdeensis usually grows on moist limestone cliffs in evergreen broad-leaved forests in association with Tectariadevexa Copel., Primulinayingdeensis Z.L. Ning, M. Kang & X.Y. Zhuang, Begonialeprosa Hance and Ficus spp.
Phenology.
Flowering from May to June and fruiting from June to August.
Etymology.
The specific epithet “yingdeensis” is derived from the type locality of the new species, i.e. Yingde City in Guangdong, China.
Additional specimens examined
(paratypes). China. Guangdong Province: Yingde City, Boluo Town, on the way from Xinzhai Village to Changshan Village, 24°24'N, 113°0'E, alt. 61 m, 9 June 2021, Q. Fan 19013 (SYS); ibid., 5 June 2022, Li Yuan-Qiu ZWY-2020 (SYS); ibid., 14 August 2022, Ye Fan ZWY-2032 (SYS).
Specimens of P.foliatasubsp.montigena examined.
China. Anhui Province: Xi County, Qingliangfeng, alt. 1300 m, 29 October 1980, Guo Xin-Hu 800023 (ANUB 13030926); ibid., 16 July 1989, Guo Xin-Hu & Zhou Shou-Biao 89011 (KUN 778733).
Specimens of P.nana examined.
China. Chongqing: Chongkou County, Mingzhong Town, Jinchi Village, Longmenxi, Dabashan National Natural Reserve, on the moist cliff, alt. 996 m, 7 July 2021, Chi Xiong XC21097 (holotype: KUN; isotypes: CQNM, IBK); Wushan County, Zhuxian Town, Shizhuzi Village, Daguling, Wulipo National Natural Reserve, in the moist valley, alt. 1310 m, 18 July 2021, Chi Xiong & Hou-Lin Zhou XC21126 (KUN); ibid., 11 September 2021, Hou-Lin Zhou s.n. (KUN).
Supplementary Material
Acknowledgements
We thank Fan Ye and Ling-Han Yang for their help in the fieldwork, and Rong-En Wu for the line drawing. This work was supported by the Guangdong Provincial Special Research Grant for the Creation of National Parks (2021GJGY034).
Appendix 1
Table A1.
Sequence information for all samples used in present study. A “/” indicates a missing sequence. Herbarium abbreviations are listed after the vouchers. The accession numbers marked in bold represent sequences newly generated.
Taxon | Voucher | Country | ITS | ETS | rpl32-trnL | rps16 | trnL-trnF |
---|---|---|---|---|---|---|---|
Matsumurellachinensis (Benth.) Bendiksby 1 | Y. Yang OYY00316 (KUN) | Pingxiang, Jiangxi, China | MW602147 | MW602117 | MW602021 | MW602053 | MW602084 |
Matsumurellachinensis (Benth.) Bendiksby 2 | Y. Yang OYY00131 (KUN) | Guilin, Guangxi, China | MW602148 | MW602118 | MW602022 | MW602054 | MW602085 |
Matsumurellayangsoensis (Y.Z. Sun) Bendiksby | L. Wu & W.B. Xu 10965 (IBK) | Yangshuo, Guangxi, China | MW602142 | MW602112 | / | / | / |
ParaphlomisalbidaHand.-Mazz.var.albida | A. Liu et al. LK0841 (CSFI) | Ningyuan, Hunan, China | MW602124 | MW602091 | MW601996 | MW602028 | MW602060 |
Paraphlomisalbidavar.brevidens Hand.-Mazz. | Y.P. Chen EM312 (KUN) | Hezhou, Guangxi, China | MW602130 | MW602098 | MW602003 | MW602035 | MW602067 |
Paraphlomisalbiflora (Hemsl.) Hand.-Mazz. | C.M. Tan et al. 1806393 (JJF) | Jiujiang, Jiangxi, China | / | MW602101 | MW602006 | MW602038 | MW602069 |
Paraphlomiscoronata (Vaniot) Y.P. Chen & C.L. Xiang 1 | E.D. Liu et al. 3043 (KUN) | Emeishan, Sichuan, China | MW602137 | MW602107 | MW602012 | MW602044 | MW602075 |
Paraphlomiscoronata (Vaniot) Y.P. Chen & C.L. Xiang 2 | C.L. Xiang 358 (KUN) | Jiangkou, Guizhou, China | MW602123 | MW602090 | MW601995 | MW602027 | MW602059 |
Paraphlomisfoliata(Dunn)C.Y. Wu & H.W. Lisubsp.foliata | S.P. Chen s.n. (KUN) | Jiangle, Fujian, China | / | MW602097 | MW602002 | MW602034 | MW602066 |
Paraphlomisfoliatasubsp.montigena X.H. Guo & S.B. Zhou | Y.C. Dai s.n. (KUN) | Hangzhou, Zhejiang, China | OM836064 | OM884453 | OM884456 | OM884459 | OM884462 |
Paraphlomisgracilis(Hemsl.) Kudôvar.gracilis 1 | A. Liu LK0931 (CSFI) | Changsha, Hunan, China | MW602134 | MW602104 | MW602009 | MW602041 | MW602072 |
Paraphlomisgracilis(Hemsl.) Kudôvar.gracilis 2 | C.L. Xiang XCL1315 (KUN) | Chongqing, China | MW602141 | MW602111 | MW602016 | MW602048 | MW602079 |
Paraphlomisgracilisvar.lutienensis (Y.Z. Sun) C.Y. Wu | C.L. Xiang XCL881 (KUN) | Shibing, Guizhou, China | MW602131 | MW602099 | MW602004 | MW602036 | MW602068 |
Paraphlomishispida C.Y. Wu | X. Li LX200702 (GXF) | Napo, Guangxi, China | MW602132 | MW602102 | MW602007 | MW602039 | MW602070 |
Paraphlomishsiwenii Y.P. Chen & Xiong Li 1 | W.H. Wu et al. DD426 (KUN) | Jingxi, Guangxi, China | OP605346 | OP609841 | OP609848 | OP609855 | OP609862 |
Paraphlomishsiwenii Y.P. Chen & Xiong Li 2 | W.H. Wu et al. DD426 (KUN) | Jingxi, Guangxi, China | OP605347 | OP609842 | OP609849 | OP609856 | OP609863 |
Paraphlomisintermedia C.Y. Wu & H.W. Li | X. Zhong et al. ZX16823 (CSH) | Suichang, Zhejiang, China | MW602135 | MW602105 | MW602010 | MW602042 | MW602073 |
Paraphlomisjavanica(Blume)Prainvar.javanica 1 | Y.P. Chen s.n. (KUN) | Kunming, Yunnan, China | MW602121 | MW602088 | MW601993 | MW602025 | MW602057 |
Paraphlomisjavanica(Blume)Prainvar.javanica 2 | L.B. Jia et al. JLB0029 (KUN) | Maguan, Yunnan, China | MW602143 | MW602113 | MW602017 | MW602049 | MW602080 |
Paraphlomisjavanicavar.pteropoda D. Fang & K.J. Yan | X. Li 2020090501 (GXF) | Jingxi, Guangxi, China | MW602140 | MW602110 | MW602015 | MW602047 | MW602078 |
Paraphlomisjiangyongensis X.L. Yu & A. Liu 1 | A. Liu et al. LK1104 (CSFI) | Jiangyong, Hunan, China | MW602128 | MW602095 | MW602000 | MW602032 | MW602064 |
Paraphlomisjiangyongensis X.L. Yu & A. Liu 2 | A. Liu et al. LK1104 (CSFI) | Jiangyong, Hunan, China | MW602129 | MW602096 | MW602001 | MW602033 | MW602065 |
Paraphlomiskwangtungensis C.Y. Wu & H.W. Li | Y.P. Chen & Y. Zhao EM1391 (KUN) | Huaiji, Guangdong, China | MW602126 | MW602093 | MW601998 | MW602030 | MW602062 |
Paraphlomislanceolata Hand.-Mazz. 1 | C.Z. Huang s.n. (KUN) | Guidong, Hunan, China | MW602145 | MW602115 | MW602019 | MW602051 | MW602082 |
Paraphlomislanceolata Hand.-Mazz. 2 | A. Liu et al. LK0825 (CSFI) | Ningyuan, Hunan, China | MW602146 | MW602116 | MW602020 | MW602052 | MW602083 |
Paraphlomislancidentata Y.Z. Sun | X. Zhong et al. ZX16824 (CSH) | Suichang, Zhejiang, China | MW602136 | MW602106 | MW602011 | MW602043 | MW602074 |
Paraphlomislongicalyx Y.P. Chen & C.L. Xiang | Y.P. Chen et al. EM583 (KUN) | Huanjiang, Guangxi, China | OK104771 | OK104774 | OK104778 | OK104780 | OK104783 |
Paraphlomismembranacea C.Y. Wu & H.W. Li | M.S. Nuraliev 1057 (MW) | Thanh Son, Phu Tho, Vietnam | / | MW602100 | MW602005 | MW602037 | / |
Paraphlomisnana Y.P. Chen, C. Xiong & C.L. Xiang 1 | C. Xiong XC21097 (KUN) | Chengkou, Chongqing, China | OM836062 | OM884451 | OM884454 | OM884457 | OM884460 |
Paraphlomisnana Y.P. Chen, C. Xiong & C.L. Xiang 2 | C. Xiong & H.L. Zhou XC21126 (KUN) | Wushan, Chongqing, China | OM836063 | OM884452 | OM884455 | OM884458 | OM884461 |
Paraphlomispagantha Dunn | L.X. Yuan et al. s.n. (KUN) | Qionghai, Hainan, China | OP605345 | OP609840 | OP609847 | OP609854 | OP609861 |
Paraphlomispaucisetosa C.Y. Wu 1 | X.X. Zhu s.n. (KUN) | Malipo, Yunnan, China | MW602125 | MW602092 | MW601997 | MW602029 | MW602061 |
Paraphlomispaucisetosa C.Y. Wu 2 | X. Li LX200704 (GXF) | Napo, Guangxi, China | MW602133 | MW602103 | MW602008 | MW602040 | MW602071 |
Paraphlomisreflexa C.Y. Wu & H.W. Li | Z.Z. Yang et al. s.n. (HIB) | Tongshan, Hubei, China | MW602122 | MW602089 | MW601994 | MW602026 | MW602058 |
Paraphlomisyingdeensis W.Y.Zhao, Y.Q.Li & Q.Fan 1 | Q. Fan et al. 19013 (SYS) | Yingde, Guangdong, China | OP605348 | OP609843 | OP609850 | OP609857 | OP609864 |
Paraphlomisyingdeensis W.Y.Zhao, Y.Q.Li & Q.Fan 2 | Q. Fan et al. 19013 (SYS) | Yingde, Guangdong, China | OP605349 | OP609844 | OP609851 | OP609858 | OP609865 |
Paraphlomisyingdeensis W.Y.Zhao, Y.Q.Li & Q.Fan 3 | Q. Fan et al. 19013 (SYS) | Yingde, Guangdong, China | / | OP609845 | OP609852 | OP609859 | OP609866 |
Phlomisfruticosa L. | Y. Tong s.n. (KUN) | Shanghai, China (cultivated) | MW602119 | MW602086 | MW601991 | MW602023 | MW602055 |
Phlomoidesdentosavar.glabrescens (Danguy) C.L. Xiang & H. Peng | Y.P. Chen EM360 (KUN) | Beijing, China (cultivated) | MW602120 | MW602087 | MW601992 | MW602024 | MW602056 |
Appendix 2
Figure A1.
Optimal Maximum Likelihood tree of Paraphlomis inferred from combined nuclear (ITS and ETS) dataset. Support value ≥ 50% BS or 0.50 PP are displayed above the branches (“-” indicates a support value < 0.50 PP).
Appendix 3
Figure A2.
Optimal Maximum Likelihood tree of Paraphlomis inferred from combined plastid (rpl32-trnL, rps16 and trnL-trnF) dataset. Support value ≥ 50% BS or 0.50 PP are displayed above the branches (“-” indicates a support value < 0.50 PP).
Citation
Guo G-X, Zhao W-Y, Chen Y-P, Xiao J-H, Li Y-Q, Fan Q (2023) Paraphlomis yingdeensis (Lamiaceae), a new species from Guangdong (China). PhytoKeys 219: 107–120. https://doi.org/10.3897/phytokeys.219.97547
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. 10.1002/tax.602015 [DOI] [Google Scholar]
- Chen YP, Drew BT, Li B, Soltis DE, Soltis PS, Xiang CL. (2016) Resolving the phylogenetic position of Ombrocharis (Lamiaceae), with reference to the molecular phylogeny of tribe Elsholtzieae. Taxon 65(1): 123–136. 10.12705/651.8 [DOI] [Google Scholar]
- Chen YP, Liu A, Yu XL, Xiang CL. (2021) A preliminary phylogenetic study of Paraphlomis (Lamiaceae) based on molecular and morphological evidence. Plant Diversity 43(3): 206–215. 10.1016/j.pld.2021.03.002 [DOI] [PMC free article] [PubMed] [Google Scholar]
- Chen YP, Sun ZP, Xiao JF, Yan KJ, Xiang CL. (2022a) Paraphlomislongicalyx (Lamiaceae), a new species from the limestone area of Guangxi and Guizhou Provinces, southern China. Systematic Botany 47(1): 251–258. 10.1600/036364422X16442668423572 [DOI] [Google Scholar]
- Chen YP, Xiao JF, Xiang CL, Li X. (2022b) Paraphlomishsiwenii (Lamiaceae), a new species from the limestone area of Guangxi, China. PhytoKeys 212: 85–96. 10.3897/phytokeys.212.91174 [DOI] [PMC free article] [PubMed] [Google Scholar]
- Chen YP, Xiong C, Zhou HL, Chen F, Xiang CL. (2022c) Paraphlomisnana (Lamiaceae), a new species from Chongqing, China. Turkish Journal of Botany 46(2): 176–182. 10.55730/1300-008X.2680 [DOI] [Google Scholar]
- Ding BY, Chen ZH, Xu YL, Jin XF, Wu DF, Chen JB, Wu WJ. (2019) New species and combination of Lamiaceae from Zhejiang, China. Guihaia 39(1): 10–15. [Google Scholar]
- Doyle JJ, Doyle JL. (1987) A rapid DNA isolation procedure for small quantities of fresh leaf tissue. Phytochemical Bulletin 19: 11–15. [Google Scholar]
- Edgar RC. (2004) MUSCLE: Multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research 32(5): 1792–1797. 10.1093/nar/gkh340 [DOI] [PMC free article] [PubMed] [Google Scholar]
- Guo XH. (1993) New taxa of Labiatae from China. Acta Phytotaxonomica Sinica 31(3): 266–272. [Google Scholar]
- Ko SC, Lee YM, Chung KS, Son DC, Nam BM, Chung GY. (2014) A new species of Paraphlomis (Lamiaceae) from Korea: An additional genus to the Korean flora. Phytotaxa 175(1): 51–54. 10.11646/phytotaxa.175.1.6 [DOI] [Google Scholar]
- Li HW, Hedge IC. (1994) Lamiaceae. In: Wu CY, Raven PH. (Eds) Flora of China (Vol.17). Science Press, Beijing & Missouri Botanical Garden Press, St. Louis, 50–299. http://www.iplant.cn/info/Lamiaceae?t=foc
- Luo XR. (1995) Lamiaceae. In: Wu DL. (Ed.) Flora of Guangdong (Vol.3). Guangdong Science and Technology Press, Guangzhou, 386–4779.
- Ronquist F, Teslenko M, van der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP. (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61(3): 539–542. 10.1093/sysbio/sys029 [DOI] [PMC free article] [PubMed] [Google Scholar]
- Stamatakis A. (2014) RAxML version 8: A tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics 30(9): 1312–1313. 10.1093/bioinformatics/btu033 [DOI] [PMC free article] [PubMed] [Google Scholar]
- Stöver BK, Müller KF. (2010) TreeGraph 2: Combining and visualizing evidence from different phylogenetic analyses. BMC Bioinformatics 11(1): 1–7. 10.1186/1471-2105-11-7 [DOI] [PMC free article] [PubMed] [Google Scholar]
- Tamura K, Stecher G, Peterson D, Filipski A, Kumar S. (2013) MEGA6: Molecular evolutionary genetics analysis version 6.0. Molecular Biology and Evolution 30(12): 2725–2729. 10.1093/molbev/mst197 [DOI] [PMC free article] [PubMed] [Google Scholar]
- Thiers BM. (2022) [continuously updated] Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. http://sweetgum.nybg.org/science/ih/ [accessed 04.11.2022]
- Wu CY, Li HW. (1977) Paraphlomis Prain. In: Wu CY, Li HW. (Eds) Flora Reipublicae Popularis Sinicae (Vol.65(2)). Science Press, Beijing, 545–572.
- Yan KJ, Fang D. (2009) A supplement to the Paraphlomis (Lamiaceae) from Guangxi, China. Journal of Tropical and Subtropical Botany 17(7): 91–92. http://jtsb.ijournals.cn/jtsb_cn/article/issue/2009_17_1 [Google Scholar]
- Yuan JC, Chen YP, Zhao Y, Li HB, Chen B, Xiang CL. (2022) Paraphlomisstrictiflora (Lamioideae, Lamiaceae), a new species from Guizhou, China. Phytotaxa 575(3): 276–286. 10.11646/phytotaxa.575.3.5 [DOI] [Google Scholar]
- Zhang RB, Deng T, Dou QL, Wei RX, He L, Ma CB, Zhao S, Hu S. (2020) Paraphlomiskuankuoshuiensis (Lamiaceae), a new species from the limestone areas of northern Guizhou, China. PhytoKeys 139: 13–20. 10.3897/phytokeys.139.47055 [DOI] [PMC free article] [PubMed] [Google Scholar]
- Zhao F, Chen YP, Salmaki Y, Drew BT, Wilson TC, Scheen AC, Celep F, Bräuchler C, Bendiksby M, Wang Q, Min DZ, Peng H, Olmstead RG, Li B, Xiang CL. (2021) An updated tribal classification of Lamiaceae based on plastome phylogenomics. BMC Biology 19(1): 1–2. 10.1186/s12915-020-00931-z [DOI] [PMC free article] [PubMed] [Google Scholar]
- Zhao WY, Zhang Z, Fan Q, Chen CQ, Liao WB, Boufford DE. (2022) Paraphlomisjinggangshanensis (Lamiaceae), a new species from Jiangxi, China. PhytoKeys 204: 1–8. 10.3897/phytokeys.204.87654 [DOI] [PMC free article] [PubMed] [Google Scholar]
Associated Data
This section collects any data citations, data availability statements, or supplementary materials included in this article.