Research Article
Research Article
Lysimachia danxiashanensis, a new species of Primulaceae from Guangdong, China
expand article infoXing-Yue Zhang, Jing-Min Dai, Qiang Fan, Zai-Xiong Chen§, Guang-Da Tang|, Wen-Bo Liao
‡ Sun Yat-sen University, Guangzhou, China
§ Administrative Commission of Danxiashan National Park, Shaoguan, China
| South China Agricultural University, Guangzhou, China
Open Access


Lysimachia danxiashanensis, a new Primulaceae species, endemic to the Danxia landscape in Guangdong Province, China, is described and illustrated. This new species is morphologically similar to L. pseudohenryi, L. phyllocephala, L. congestiflora and L. kwangtungensis, but it differs from the similar species by its purplish-red plants, petiole without wings, calyx with orange glandular and the corolla margin serrated on upper half with orange-red glandular punctates. This new species belongs to Lysimachia subgen. Lysimachia sect. Nummularia. Phylogenetic analysis confirmed that L. danxiashanensis is a distinct clade, based on the combined data of ITS and rbcL sequences. The conservation status of the new species was evaluated as Endangered (EN) according to IUCN Red List Categories and Criteria.

Key words

Danxia landscape, IUCN Red List, Lysimachia, taxonomy


Lysimachia L. is a member of the tribe Lysimachieae (Primulaceae) and is composed of over 200 species (Chen and Hu 1979; Hu and Kelso 1996; Wang et al 2018). This genus is the most widely distributed genera of Primulaceae and is mainly distributed in the temperate and subtropical regions of the Northern Hemisphere, but there are also a few species in Africa, Australia and South America (Hu 1994; Hao and Hu 2001; Hao et al. 2004; Kodela 2006). China is considered to be the origin and diversity centre of Lysimachia, with 138 native species having been recorded in Flora of China and nearly 80% of them are endemic species (Chen and Hu 1979; Hu and Kelso 1996; Hao and Hu 2001). In recent years, many new species of this genus have been described and this highlights more opportunities for discoveries in China (Wang et al 2018; Huang et al 2020; Yan et al 2022).

An unknown species of Lysimachia was discovered during a field floristic investigation from May 2022 to August 2023 in Danxiashan National Park, Renhua County, Guangdong Province. It is most similar to L. congestiflora Hemsl., but its purplish-red plants, petiole without wings, corolla lobes serrated on upper half and calyx with orange glandular punctates clearly distinguish from the latter. After careful morphological comparison by specimens and consultation with relevant literature and molecular phylogenetic analysis, we confirmed that it represented a new species, described and illustrated here. The threat status of the new species is assessed according to the IUCN Red List Categories and Criteria (IUCN Standards and Petitions Committee 2022).

Materials and methods

Morphological study

The morphological characters of the new species were observed and measured, based on fresh and dry specimens using a micrometer and a stereomicroscope and were compared with its related species, based on herbarium specimens deposited at the Herbarium of SYS and IBSC (the herbarium acronyms follow Thiers (2023)), as well as the digital images on the Chinese Virtual Herbarium ( and the China Field Herbarium ( Morphological observation and examination were conducted in the SYS.

Taxon sampling and molecular analysis

Leaf tissue of the putative new species and related species was collected from one population and silica dried in zip-lock plastic bags until use for comparisons and taxonomic treatment. Total DNA was extracted with a modified CTAB method (Doyle and Doyle 1987). Regions of the partial internal transcribed spacer 1, the 5.8S ribosomal RNA gene and partial internal transcribed spacer 2 were amplified using the previously-reported primers ITS1 and ITS4 (White et al. 1990) and the ribulose-1,5-bisphosphate carboxylase/oxygenase large subunit (rbcL) gene was amplified using the primers rbcLa-f (Kress and Erickson 2007) and 724R (Fay et al. 1997). PCR amplifications were performed following Fan et al. (2015). Following the studies of Lysimachia (Zhang et al. 2011; Yan et al. 2018), we retrieved 67 ITS and rbcL accession of 30 species from GenBank, which belong to subgenus Lysimachia L., subgenus Palladia (Moench) Hand.-Mazz., subgenus Heterostylandra (Hand. -Mazz.) Chen et C.M.Hu. and subgenus Idiophyton Hand.-Mazz. Two accessions of the putative new species (GenBank Acc. ITS No.: OR665389, OR665390; rbcL No: PP025352, PP035354) and one accession of Lysimachia kwangtungensis (GenBank Acc. ITS No.: OR941025; rbcL No: PP025355) were sequenced for this study. Ardisia verbascifolia was selected as outgroup. The sequences were aligned using MAFFT v.7 (Katoh and Standley 2013) and subsequently manually adjusted. Phylogenetic constructions were based on Maximum Likelihood (ML) and Bayesian Inference (BI) and were respectively run by IQ-TREE v. 2.0.3 (Minh et al. 2020) and MrBayes version 3.1.2 (Huelsenbeck and Ronquist 2001), selecting best-fit model as SYM+I+G4 with 2000 bootstraps (BS) for ML analysis. ModelFinder v.2.2.0 (Kalyaanamoorthy et al. 2017) was used to select the best-fit partition model (Edge-linked) using the BIC criterion. The best-fit models according to BIC were SYM+G4 for ITS and K2P+I+G4 for rbcL. BI analysis employed random starting trees and four Markov Chain Monte Carlo (MCMC) simulations were run simultaneously and sampled every 1000 generations for 10 million generations. The average standard deviation of split frequencies (< 0.01) was used to assess the convergence of the two runs. Bayesian posterior probabilities (PP) were calculated as the majority consensus of all sampled trees with the first 25% discarded as burn-in.

Results and discussion

Morphological comparison

According to the classification of Chen and Hu (1979), L. danxiashanensis is a member of subgenus Lysimachia sect. Nummularia, which is characterised by stems prostrate to erect on the upper part, leaves opposite, racemes shortened to subcapitate, filaments longer than anthers, lower part connate into a tube, corolla and calyx with coloured glandular punctates (Fig. 2). In China, there are over 50 species of sect. Nummularia and it widely distributed from southwest to the east and south China. Morphologically, Lysimachia danxiashanensis is similar to L. phyllocephala Hand.-Mazz., L. pseudohenryi Pamp., L. congestiflora Hemsl. and L. kwangtungensis (Hand.-Mazz.) C.M.Hu by sharing the following morphological features: stems with multicellular hairs, leaves opposite and racemes terminal. However, the new species can be easily distinguished from similar species by combination characters including its purplish-red plants (vs. green), petiole without wings (vs. narrowly winged in L. pseudohenryi and L. congestiflora, narrowly margined and auriculate at base in L. kwangtungensis and absent in L. phyllocephala) and the corolla lobes serrated on upper half (vs. entire margin in all four species). A more detailed morphological comparison of these species is summarised in Table 1.

Table 1.

Morphological comparison of Lysimachia danxiashanensis with its four closely-related species.

Characters L. danxiashanensis L. phyllocephala L. pseudohenryi L. congestiflora L. kwangtungensis
Stems upper erect, creeping at base erect to ascending-erect, prostrate at base erect or arcuate at base prostrate and branches ascending erect
Colour of plants purplish-red green green green green
Leaf shape ovate to broadly ovate or oval ovate to ovate-lanceolate rhomboid-ovate to ovate, rarely ovate-lanceolate ovate to broadly ovate or suborbicular ovate-lanceolate to lanceolate
Petiole wings absent absent narrowly winged narrowly winged narrowly margined and auriculate at base
Corolla lobes margin serrated on upper half with orange-red glandular punctates margin entire, with sparsely transparent glandular punctates margin entire, with transparent glandular punctates margin entire, with dull red or black glandular punctates margin entire, with red to dark purple glandular punctates
Style 6–8 mm; glabrous c. 8 mm; puberulous 5–6 mm; lower part with pubescent 5–7 mm; glabrous 5–6 mm; glabrous
Glandular dots on Calyx orange, dense absent absent absent orange, sparse

Molecular analysis

The combined aligned matrix had a length of 1268 bp (ITS: 650, rbcL: 615), including 373 variable sites, of which 291 were parsimony-informative. The two accessions of the new species were from the same population and formed a separate monophyletic lineage (Fig. 1: BS = 93%, PP = 0.72), the sister group of L. rubiginosa. Although L. danxiashanensis and L. rubiginosa both belong to subgen. Lysimachia sect. Nummularia, the new species can be easily distinguished from the latter by its shorter plants (10–28 cm vs. 30–60 (100) cm), orange glandular punctates on corolla lobes and calyx (vs. black or brown glandular striate on leaves, corolla lobes and calyx), 5–9 flowered on branches and stems axis (vs. 3–5 flowered on branches, seldom on main axis).

Figure 1. 

Phylogenetic tree of Lysimachia danxiashanensis and related species generated by Bayesian Inference (BI) of the combined dataset (ITS, rbcL). Bootstrap values of the ML and BI posterior probabilities are shown along the branches. The new species in yellow shaded area, green indicates subgen. Lysimachia, orange is subgen. Palladia, purple is subgen. Heterostylandra and grey is subgen. Idiophyton. Blue indicates outgroup, Ardisia verbascifolia.

Geographically, L. rubiginosa is distributed in Guangxi, Guizhou, Hubei, Hunan, Sichuan, Yunnan and Zhejiang and it mainly grows in limestone. In contrast, the new species is distributed in Danxia landscape, Guangdong. The geographical distribution of these two taxa does not overlap.

Although the infrageneric phylogenetic relationships within Chinese Lysimachia remain controversial (Zhen and Chen 2012; Liu et al. 2023), the phylogenetic tree placed L. danxiashanensis distant from other species in this genus (Fig. 1). Based on the morphological and molecular evidence, we confirmed that L. danxiashanensis is a distinct species. Therefore, we describe and provide illustrations for the new species below.

Taxonomic treatment

Lysimachia danxiashanensis W.B.Liao, Q.Fan & G.D.Tang, sp. nov.

Figs 2, 3


Lysimachia danxiashanensis can be distinguished from L. congestiflora by its purple-red plants (vs. green), petiole without wings (vs. narrowly winged), corolla lobes yellow with serrations on upper half (vs. dull red at base with entire margin) and calyx with orange glandular (vs. without glandular) (Fig. 4).

Figure 2. 

Lysimachia danxiashanensis W.B.Liao, Q.Fan & G.D.Tang, sp. nov. A habit B flowering branch C inflorescence D abaxial and adaxial views of leave E lateral view of flower F dorsal view of flower G adaxial side of corolla lobes H stamens I immature capsule J abaxial (1st, 2nd, 5th) and adaxial (3rd, 4th) views of calyx lobes K pistil and densely pilose pedicel L stigma M cross-section of ovary (Photographers: A, C, D, I by Xing-Yue Zhang; B, E, F by Qiang Fan; G, H, J–M by Jing-Min Dai).

Figure 3. 

Lysimachia danxiashanensis W.B.Liao, Q.Fan & G.D.Tang, sp. nov. A habit B abaxial (left) and adaxial (right) views of leaf C adaxial side of corolla lobes D lateral view of flower E dorsal view of flower F stamens G pistil and stigma H fruit I adaxial (left) and abaxial (right) views of calyx (Drawn by Rong-En Wu).

Figure 4. 

Morphological differences between L. congestiflora and L. danxiashanensis A L. congestiflora B L. danxiashanensis 1 plants 2 corolla lobes 3 calyx lobes 4 petiole (Photographers: A–1 by Wan-Yi Zhao A2–4 by Xin-Xin Zhu B1, 2, 4 by Xing-Yue Zhang B–3 by Qiang Fan).


China. Guangdong Province, Danxiashan National Park, 25°0'N, 113°37'E, 311 m a.s.l., 12 May 2023 (fl.), Xing-Yue Zhang, Zai-Xiong Chen DNPC 3801 (holotype SYS!; isotypes CANT! SYS!).


Perennial herb, 10–28 cm tall. Stems prostrate, rooting at nodes, upper part ascending, purplish-red, terete, with dense white multicellular hairs. Leaves opposite, upper 2 or 3 pairs usually crowded, papery, ovate to broadly ovate or oval, wavy margin, 1.6–3.8 × 1.2–2.4 cm, apex acute, base broadly cuneate; adaxial surface dark green with antrorse strigose, abaxial surface purple-red, with strigose and densely pilose along the mid-rib vein; lateral veins 2–4-paired; petiole without wings, 0.5–2.7 cm, densely villous. Racemes terminal, abbreviated, capitate, 5–9 flowered; pedicel 3–5 mm long, densely pilose. Calyx 5, parted nearly to base; lobes lanceolate, 2–2.5 × 7–8.5 mm long, sparsely orange glandular on both surfaces, pilose outside. Corolla yellow; tube 1.5–2 mm; lobes 5, obovate-elliptical, 3–5 × 9–10 mm, serrate on upper half, apex acute to obtuse, abaxially glabrous, sparsely orange-red glandular, adaxially glabrous with orange-reddish glandular. Stamens 5, filaments glabrous, connate basally into a 2–3 mm high tube, free parts 2.8–4.5 mm; anthers ovate-lanceolate, dorsifixed, opening by lateral slits, ca. 2.8 mm long. Ovary white, terete, apex puberulous, glabrous at lower part; style glabrous, 6–8 mm long, stigma obtuse with papillae. Capsule subglobose, green, apex puberulous, 3–5 mm in diam.


The flowering of Lysimachia danxiashanensis is from May to June; and the fruiting in June.


Lysimachia danxiashanensis is currently known only from the type locality, Danxiashan National Nature Reserve, Guangdong, China.


Lysimachia danxiashanensis was observed to grow on wet rocks of Danxia landform at elevations 270 to 320 m.


The specific epithet refers to the type locality, Danxiashan National Nature Reserve in Guangdong Province, China.

Local name

The Chinese name of the new species is here given as 丹霞山过路黄 (Dān xiá Shān Guò Lù Huáng).

Provisional conservation status

Endangered (EN). In the past two years, we have conducted several field investigations on the Danxia landscapes in Guangdong Province, with only four populations of Lysimachia danxiashanensis being found in Danxiashan National Nature Reserve and the number of mature individuals is stable (total < 200 individuals); However, its habitat is on the side of the road, which is vulnerable to human disturbance. L. danxiashanensis is regarded as Endangered (EN) according to D (the number of mature individuals in the population < 250) (IUCN Standards and Petitions Committee 2022).

Additional specimens examined

(paratypes). China, Guangdong: Danxiashan National Park, 25°0'N, 113°38'E, 298 m a.s.l., 12 June 2023 (fr.), Jie-Hao Jin DNPC 3803 (SYS!); Danxiashan National Park, 25°0'N, 113°38'E, 14 August 2023, Qiang Fan, Jie-Hao Jin & Li-Juan Liao DNPC 3845 (SYS!).


We are grateful to Jie-Hao Jin and Li-Juan Liao for their assistance during field collections. We are deeply grateful to Mrs Rong-En Wu for her excellent illustrations in the manuscript. We are grateful for the photos provided by Xin-Xin Zhu and Wan-Yi Zhao. We appreciate Dr Wan-Yi Zhao and Dr Cui-Ying Huang for their help and suggestions for this article.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.


This study was supported by the Guangdong Provincial Special Research Grant for the Creation of National Parks (2021GJGY034) and the Foundation of Administrative Committee of Danxiashan National Park (K22-33000-060, 2016-0293).

Author contributions

Xing-Yue Zhang participated field investigation, taxonomic confirmation and wrote the original draft. Jing-Min Dai performed the molecular analysis. Zai-Xiong Chen and Qiang Fan participated field investigation and collected the plants. Wen-Bo Liao and Guang-Da Tang are project leaders and contributed to paper writing.

Author ORCIDs

Xing-Yue Zhang

Jing-Min Dai

Qiang Fan

Zai-Xiong Chen

Guang-Da Tang

Wen-Bo Liao

Data availability

The newly-obtained sequences of Lysimachia danxiashanensis and L. kwangtungensis have been submitted to the NCBI website. The ITS and rbcL sequence data supporting the findings of this study are available within Appendix 1. The morphological data used in the study are included in this paper.


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Appendix 1

Table A1.

List of the GenBank accession numbers of the rbcL and ITS sequences of sampled species in this study.

Taxon Voucher Locality rbcL ITS
Lysimachia alfredii Hance Hao394 Lianping, Guangdong, China JF942344 JN638405
Y2009279 Ruyuan, Guangdong, China JF942343 JN638406
Lysimachia candida Lindl. Ge2010001 Yangchun, Guangdong, China JF942346 JF976885
Y2010016 Tongbai, Henan, China JF942345 JF976884
Lysimachia chapaensis Merrill GBOWS704 Maguan, Yunan, China JF942350 JF976889
GBOWS878 Hekou, Yunnan, China JF942349 JF976888
Hao209 Wuhan, Hubei, China JF942392 AF547691
Lysimachia chekiangensis C.C.Wu Y2009263-1 Longquan, Zhejiang, China JF942352 JF976891
Y2009263-2 Longquan, Zhejiang, China JF942351 JF976890
Lysimachia christinae Hance Y2009209 Jiujiang, Jiangxi, China JF942357 JF976896
Y2009235 Shucheng, Anhui, China JF942356 JF976895
Y2009272 Jiangle, Fujian, China JF942354 JF976893
Lysimachia clethroides Duby Y2009157 Tongbai, Henan, China JF942362 JF976899
Y2009248 Lin’an, Zhejiang, China JF942360 JF976898
Hao955 Wuxi, Chongqing, China JF942359 JF976897
Lysimachia congestiflora Hemsl. Y2009196 Xinjian, Jiangxi, China JF942367 JF976904
Y2009266 Longquan, Zhejiang, China JF942366 JF976903
GBOWS262 Malipo, Yunnan, China JF942365 JF976902
Lysimachia crispidens Hemsl. Hao212 Yichang, Hubei, China JF942369 JF976906
Y2010029 Xinhua, Hubei, China JF942368 JF976905
Lysimachia decurrens Forst.F. GBOWS1234 Hekou, Yunnan, China JF942371 JF976908
Ye et al. 3980 Lianshan, Guangdong, China JF942370 JF976907
Lysimachia deltoidea var. cinerascens Franch. Hao & Yan1033 Dali, Yunnan, China JF942374 JF976911
Hao731 Yongsheng, Yunnan, China JF942373 JF976910
GLM081121 Zhongdian, Yunnan, China JF942372 JF976909
Lysimachia dextrorsiflora X.P.Zhang, X.H.Guo & J.W.Shao Y2009265-1 Longquan, Zhejiang, China JF942376 JF976913
Y2009265-2 Longquan, Zhejiang, China JF942375 JF976912
Lysimachia erosipetala Chen et C.M.Hu Y2010037-1 Emeishan, Sichuan, China JF942378 JF976915
Y2010037-2 Emeishan, Sichuan, China JF942377 JF976914
Lysimachia fistulosa var. wulingensis Chen et C.M.Hu Ye et al. 3561 Lianshan, Guangdong, China JF942381 JF976917
Ning20101 Jinggangshan, Jiangxi, China JF942380 JF976916
Lysimachia fordiana Oliv. Ye et al. 3940 Lianshan, Guangdong, China JF942384 JF976920
Lysimachia fortunei Maxim Y2009285 Ruyuan, Guangdong, China JF942383 JF976919
Lysimachia hemsleyana Maxim Guo20001 Ningguo, Anhui, China JF942398 JF976932
Y2009245 Lin’an, Zhejiang, China JF942395 JF976929
Y2010008 Tongbai, Henan, China JF942394 JF976928
Lysimachia hemsleyi Franch. Hao713 Huili, Sichuan, China JF942402 JF976935
Hao730 Yongsheng, Yunnan, China JF942401 JF976934
Lysimachia heterogenea Klatt Y2009199 Jiujiang, Jiangxi, China JF942407 JF976939
Y2010009 Tongbai, Henan, China JF942405 JF976938
Lysimachia klattiana Hance Y2010014-1 Tongbai, Henan, China JF942415 JF976947
Y2010014-2 Tongbai, Henan, China JF942414 JF976946
Lysimachia lobelioides Wall. Hao303 Menglian, Yunnan, China JF942419 JF976951
Y2010001 Jingping, Yunan, China JF942418 JF976950
Lysimachia longipes Hemsl. Y2009255-1 Kaihua, Zhejiang, China JF942422 JF976954
Y2009255-2 Kaihua, Zhejiang, China JF942421 JF976953
Guo xinhu200012 Shitai, Anhui, China JF942420 JF976952
Lysimachia melampyroides R.Knuth Lichanghan8174 Shangzhi, Hunan, China JF942424 JF976956
Dengyunfei15945 Xinning, Hunan, China JF942423 JF976955
Lysimachia omeiensis Hemsl. Y2010033 Emeishan, Sichuan, China JF942426 JF976958
Hao224 Emeishan, Sichuan, China JF942425 JF976957
Lysimachia paridiformis var. paridiformis Franch. Chen s.n. Enshi, Hubei, China JF942429 JF976961
Lysimachia paridiformis var. stenophylla Franch. Deng15921 Xinning, Hunan, China JF942431 JF976963
Y2010044 Emeishan, Sichuan, China JF942430 JF976962
GLM07658 Zhenxiong, Yunnan, China JF942428 JF976960
Lysimachia patungensis Hand.-Mazz. Y2009187 Jinggangshan, Jiangxi, China JF942435 JF976967
Y2009258 Kaihua, Zhejiang, China JF942434 JF976966
Y2009280 Ruyuan, Guangdong, China JF942433 JF976965
Ye et al. 3851 Lianshan, Guangdong, China JF942432 JF976964
Lysimachia pentapetala Bunge Y2010013-1 Tongbai, Henan, China JF942437 JN638407
Lysimachia phyllocephala Hand.-Mazz. Y2010030 Emeishan, Sichuan, China JF942439 JF976969
Y2010048 Nanchuan, Chongqing, China JF942438 JF976968
GLM07662 Yanjin, Yunnan, China JF942399 JF976933
Lysimachia pseudohenryi Pamp. Guo XH 20007 East Asia MG950600 MG877828
Lysimachia rubiginosa Hemsl. Hao704 Hongya, Sichuan, China JF942444 JF976974
Y2010036 Emeishan, Sichuan, China JF942443 JF976973
Hao419 Dujiangyan, Sichuan, China JF942442 JF976972
Lysimachia danxiashanensis DNPC-3711 Danxiashan, Guangdong, China PP025352 OR665389
DNPC-3711 Danxiashan, Guangdong, China PP025354 OR665390
Lysimachia kwangtungensis (Handel-Mazzetti) C.M.Hu DNPC-3743 Danxiashan, Guangdong, China PP025355 OR941025
Ardisia verbascifolia Mez. GBOWS1216 Hekou, Yunnan, China JN638410 JN638408
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