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Research Article
Paraphlomis yingdeensis (Lamiaceae), a new species from Guangdong (China)
expand article infoGuo-Xin Guo, Wan-Yi Zhao§, Ya-Ping Chen|, Jin-Hai Xiao, Yuan-Qiu Li, Qiang Fan§
‡ Guangdong Shimentai National Nature Reserve, Yingde, China
§ Sun Yan-Sen University, Guangzhou, China
| Kunming Institute of Botany, Chinese Academy of Science, Kunming, China
Open Access

Abstract

Paraphlomis yingdeensis (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. foliata subsp. 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, Paraphlomis albiflora (Hemsl.) Hand.-Mazz., P. nana Y.P. Chen, C. Xiong & C.L. Xiang and P. javanica var. 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 23) 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. foliata subsp. 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. Paraphlomis foliata subsp. 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 23) indicated that P. foliata subsp. montigena might represent an independent species within the genus as it is distantly related to P. foliata subsp. foliata. The new species can be distinguished from P. foliata subsp. 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. foliata subsp. montigena; the base of lamina is broadly cuneate and not decurrent in the new species, but is cuneate and decurrent in P. foliata subsp. montigena; P. yingdeensis has bristle-like-acuminate apex of calyx teeth, in contrast, the apex of calyx teeth of P. foliata subsp. 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 23). 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 Paraphlomis yingdeensis, P. foliata subsp. montigena and P. nana.

Characters P. yingdeensis P. foliata subsp. 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. foliata subsp. 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.

Figs 2, 3, 4 Chinese name: 英德假糙苏

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!)

Figure 2. 

Paraphlomis yingdeensis from the type locality A habitat B, C plants D stem. (Photographs: A, C, D by W.-Y. Zhao; B by Y.-Q. Li).

Diagnosis

Paraphlomis yingdeensis is morphologically similar to P. foliata subsp. 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).

Figure 3. 

Floral traits of Paraphlomis yingdeensis A, 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).

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.

Figure 4. 

Line drawing of Paraphlomis yingdeensis A 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).

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. Paraphlomis yingdeensis usually grows on moist limestone cliffs in evergreen broad-leaved forests in association with Tectaria devexa Copel., Primulina yingdeensis Z.L. Ning, M. Kang & X.Y. Zhuang, Begonia leprosa 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. foliata subsp. 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).

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).

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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
Matsumurella chinensis (Benth.) Bendiksby 1 Y. Yang OYY00316 (KUN) Pingxiang, Jiangxi, China MW602147 MW602117 MW602021 MW602053 MW602084
Matsumurella chinensis (Benth.) Bendiksby 2 Y. Yang OYY00131 (KUN) Guilin, Guangxi, China MW602148 MW602118 MW602022 MW602054 MW602085
Matsumurella yangsoensis (Y.Z. Sun) Bendiksby L. Wu & W.B. Xu 10965 (IBK) Yangshuo, Guangxi, China MW602142 MW602112 / / /
Paraphlomis albida Hand.-Mazz. var. albida A. Liu et al. LK0841 (CSFI) Ningyuan, Hunan, China MW602124 MW602091 MW601996 MW602028 MW602060
Paraphlomis albida var. brevidens Hand.-Mazz. Y.P. Chen EM312 (KUN) Hezhou, Guangxi, China MW602130 MW602098 MW602003 MW602035 MW602067
Paraphlomis albiflora (Hemsl.) Hand.-Mazz. C.M. Tan et al. 1806393 (JJF) Jiujiang, Jiangxi, China / MW602101 MW602006 MW602038 MW602069
Paraphlomis coronata (Vaniot) Y.P. Chen & C.L. Xiang 1 E.D. Liu et al. 3043 (KUN) Emeishan, Sichuan, China MW602137 MW602107 MW602012 MW602044 MW602075
Paraphlomis coronata (Vaniot) Y.P. Chen & C.L. Xiang 2 C.L. Xiang 358 (KUN) Jiangkou, Guizhou, China MW602123 MW602090 MW601995 MW602027 MW602059
Paraphlomis foliata (Dunn) C.Y. Wu & H.W. Li subsp. foliata S.P. Chen s.n. (KUN) Jiangle, Fujian, China / MW602097 MW602002 MW602034 MW602066
Paraphlomis foliata subsp. montigena X.H. Guo & S.B. Zhou Y.C. Dai s.n. (KUN) Hangzhou, Zhejiang, China OM836064 OM884453 OM884456 OM884459 OM884462
Paraphlomis gracilis (Hemsl.) Kudô var. gracilis 1 A. Liu LK0931 (CSFI) Changsha, Hunan, China MW602134 MW602104 MW602009 MW602041 MW602072
Paraphlomis gracilis (Hemsl.) Kudô var. gracilis 2 C.L. Xiang XCL1315 (KUN) Chongqing, China MW602141 MW602111 MW602016 MW602048 MW602079
Paraphlomis gracilis var. lutienensis (Y.Z. Sun) C.Y. Wu C.L. Xiang XCL881 (KUN) Shibing, Guizhou, China MW602131 MW602099 MW602004 MW602036 MW602068
Paraphlomis hispida C.Y. Wu X. Li LX200702 (GXF) Napo, Guangxi, China MW602132 MW602102 MW602007 MW602039 MW602070
Paraphlomis hsiwenii Y.P. Chen & Xiong Li 1 W.H. Wu et al. DD426 (KUN) Jingxi, Guangxi, China OP605346 OP609841 OP609848 OP609855 OP609862
Paraphlomis hsiwenii Y.P. Chen & Xiong Li 2 W.H. Wu et al. DD426 (KUN) Jingxi, Guangxi, China OP605347 OP609842 OP609849 OP609856 OP609863
Paraphlomis intermedia C.Y. Wu & H.W. Li X. Zhong et al. ZX16823 (CSH) Suichang, Zhejiang, China MW602135 MW602105 MW602010 MW602042 MW602073
Paraphlomis javanica (Blume) Prain var. javanica 1 Y.P. Chen s.n. (KUN) Kunming, Yunnan, China MW602121 MW602088 MW601993 MW602025 MW602057
Paraphlomis javanica (Blume) Prain var. javanica 2 L.B. Jia et al. JLB0029 (KUN) Maguan, Yunnan, China MW602143 MW602113 MW602017 MW602049 MW602080
Paraphlomis javanica var. pteropoda D. Fang & K.J. Yan X. Li 2020090501 (GXF) Jingxi, Guangxi, China MW602140 MW602110 MW602015 MW602047 MW602078
Paraphlomis jiangyongensis X.L. Yu & A. Liu 1 A. Liu et al. LK1104 (CSFI) Jiangyong, Hunan, China MW602128 MW602095 MW602000 MW602032 MW602064
Paraphlomis jiangyongensis X.L. Yu & A. Liu 2 A. Liu et al. LK1104 (CSFI) Jiangyong, Hunan, China MW602129 MW602096 MW602001 MW602033 MW602065
Paraphlomis kwangtungensis C.Y. Wu & H.W. Li Y.P. Chen & Y. Zhao EM1391 (KUN) Huaiji, Guangdong, China MW602126 MW602093 MW601998 MW602030 MW602062
Paraphlomis lanceolata Hand.-Mazz. 1 C.Z. Huang s.n. (KUN) Guidong, Hunan, China MW602145 MW602115 MW602019 MW602051 MW602082
Paraphlomis lanceolata Hand.-Mazz. 2 A. Liu et al. LK0825 (CSFI) Ningyuan, Hunan, China MW602146 MW602116 MW602020 MW602052 MW602083
Paraphlomis lancidentata Y.Z. Sun X. Zhong et al. ZX16824 (CSH) Suichang, Zhejiang, China MW602136 MW602106 MW602011 MW602043 MW602074
Paraphlomis longicalyx Y.P. Chen & C.L. Xiang Y.P. Chen et al. EM583 (KUN) Huanjiang, Guangxi, China OK104771 OK104774 OK104778 OK104780 OK104783
Paraphlomis membranacea C.Y. Wu & H.W. Li M.S. Nuraliev 1057 (MW) Thanh Son, Phu Tho, Vietnam / MW602100 MW602005 MW602037 /
Paraphlomis nana Y.P. Chen, C. Xiong & C.L. Xiang 1 C. Xiong XC21097 (KUN) Chengkou, Chongqing, China OM836062 OM884451 OM884454 OM884457 OM884460
Paraphlomis nana Y.P. Chen, C. Xiong & C.L. Xiang 2 C. Xiong & H.L. Zhou XC21126 (KUN) Wushan, Chongqing, China OM836063 OM884452 OM884455 OM884458 OM884461
Paraphlomis pagantha Dunn L.X. Yuan et al. s.n. (KUN) Qionghai, Hainan, China OP605345 OP609840 OP609847 OP609854 OP609861
Paraphlomis paucisetosa C.Y. Wu 1 X.X. Zhu s.n. (KUN) Malipo, Yunnan, China MW602125 MW602092 MW601997 MW602029 MW602061
Paraphlomis paucisetosa C.Y. Wu 2 X. Li LX200704 (GXF) Napo, Guangxi, China MW602133 MW602103 MW602008 MW602040 MW602071
Paraphlomis reflexa C.Y. Wu & H.W. Li Z.Z. Yang et al. s.n. (HIB) Tongshan, Hubei, China MW602122 MW602089 MW601994 MW602026 MW602058
Paraphlomis yingdeensis W.Y.Zhao, Y.Q.Li & Q.Fan 1 Q. Fan et al. 19013 (SYS) Yingde, Guangdong, China OP605348 OP609843 OP609850 OP609857 OP609864
Paraphlomis yingdeensis W.Y.Zhao, Y.Q.Li & Q.Fan 2 Q. Fan et al. 19013 (SYS) Yingde, Guangdong, China OP605349 OP609844 OP609851 OP609858 OP609865
Paraphlomis yingdeensis W.Y.Zhao, Y.Q.Li & Q.Fan 3 Q. Fan et al. 19013 (SYS) Yingde, Guangdong, China / OP609845 OP609852 OP609859 OP609866
Phlomis fruticosa L. Y. Tong s.n. (KUN) Shanghai, China (cultivated) MW602119 MW602086 MW601991 MW602023 MW602055
Phlomoides dentosa var. 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).

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