Research Article
Research Article
Lysimachia xiangxiensis (Primulaceae), a new species from limestone area in Hunan Province, central China
expand article infoCun Mou§|, Yu Wu, Liang Xiang#, Xiao-Mei Xiang¤, Dai-Gui Zhang¤
‡ Key Laboratory of Cultivation and Protection for Non-Wood Forest Trees of Ministry of Education, Changsha, China
§ Central South University of Forestry and Technology, Changsha, China
| Hunan Forest Botanical Garden, Changsha, China
¶ Hunan Normal University, Changsha, China
# Jishou Dehang Scenic Area Management Office, Jishou, China
¤ Jishou University, Jishou, China
Open Access


A new species of Lysimachia, L. xiangxiensis (Primulaceae), is described and illustrated from western Hunan, central China. The species is similar to L. melampyroides in plant densely strigillose, leaves subglabrous adaxially, and flowers usually solitary in axils of upper leaves, but differs by the succulent leaves, the creeping or ascending stems 15–25 cm long, and the suborbicular to broadly elliptic corolla lobes. This new species is also supported by a molecular phylogenetic analysis of some Lysimachia species based on ITS sequence data.


Lysimachia, L. xiangxiensis, new species, taxonomy, western Hunan


The genus Lysimachia L., a large genera of Primulaceae s. l. (APG III 2009), consists of over 180 species of annual or perennial herbs (Hu and Kelso 1996). Lysimachia has a nearly cosmopolitan distribution, mainly occurring in the temperate and subtropical parts of the northern hemisphere, with a few species in Africa, Australia and South America (Hu and Kelso 1996, Liu et al. 2014a). Southwestern China and its neighboring region of Indochina Peninsula have an extremely high species diversity with ca. 130 species and have been considered to be the diversity center of the genus (Yan et al. 2017).

During our expedition in 2017 and 2019 to the Youshui River valley in western Hunan, China, an unusual population of Lysimachia, with the plants having revolute succulent leaves, caught our attention. After consulting the relevant literature (Chen et al. 1989, Hu and Kelso 1996, Yan and Hao 2012, Liu et al. 2014a, Liu et al. 2014b, Zhou et al. 2015, Wang et al. 2018) and checking relevant specimens, we determined that the population represents a new species. Additionally, the new species is supported by a molecular phylogenetic analysis of some Lysimachia species based on ITS sequence data.

Materials and methods

Taxon sampling and morphological analysis

The type specimens and fresh materials of the new species were collected from Huayuan County and Jishou City, Hunan Province, central China. Morphological observations and measurements were randomly made on flowering and fruiting plants. We examined related specimens kept in JIU and HUN and also specimen images in the online database of Chinese Virtual Herbarium ( and JSTOR Global Plants (

A total of 39 nuclear ribosomal ITS sequences for 34 species (Appendix S1) were downloaded from GenBank, following a study of Lysimachia (Zhang et al. 2011, Zhou et al. 2015). Two accessions of the putatively new species were sequenced for this study (GenBank Acc. No.: MN647744, MN647745). Ardisia verbascifolia Mez was selected as outgroup following Zhang et al. (2011). Voucher specimens of those specimens of the new species used for sequencing were deposited in JIU.

Molecular analyses

Total genomic DNA of the two accessions of the putatively new species was isolated from silica gel-dried leaves using a modified cetyltrimethylammonium bromide procedure (Doyle and Doyle 1987). The ITS region was amplified and sequenced by method of Zhang et al. (2011).

Phylogenetic trees were constructed using maximum likelihood (ML) and Bayesian inference (BI). The models determined for the datasets using the Akaike information criterion (Burnham and Anderson 2003) as implemented in MrModeltest 2.3 (Nylander 2004). ML trees were generated in RAxML 7.2.6 (Stamatakis 2006) with 1000 bootstrap replicates. BI trees were inferred in MrBayes version 3.1.2 (Huelsenbeck and Ronquist 2001). Four chains, each starting with a random tree, were run for 1,000,000 generations with trees sampled every 1000 generations. The convergence of the two runs was accessed with the average standard deviation of split frequencies less than 0.01. After the first ca. 25% discarded as burn-in, the remaining trees were imported into PAUP* v.4.0b10 (Swofford 2002) and a 50% majority rule consensus tree was produced to obtain posterior probabilities (PP) of the clades.

Results and discussion

Morphological comparisons

According to the key in Hu and Kelso (1996), the new species is positioned to “Key 2” by flowers 5-merous, homomorphic, corolla yellow, anthers shorter than filaments, and further to “19a” by anthers distinctly dorsifixed (1b), inflorescences not paniculate(3b), stems more than 5 cm and leaves opposite (5b), corolla subfunnelform, filaments connate 1/3–1/2 into a tube (7b), flowers axillary and solitary or in terminal clusters with bracts leaflike (12b), inflorescences not capitate (17a), leaf blade not connate-perfoliate (18b), flowers solitary and axillary or in terminal racemes, plants strigillose (19a).

Morphologically, the new species is most similar to L. melampyroides R. Knuth in Engler with which it shares such features as the plants densely strigillose, leaves subglabrous adaxially, and flowers that are usually solitary in axils of upper leaves. However, the new species differs from L. melampyroides by the succulent leaves, the creeping or drooping stems 15–25 cm long, and the suborbicular to broadly elliptic corolla lobes. A morphological comparison between the new species and L. melampyroides is presented in Table 1.

Table 1.

Morphological comparison between Lysimachia xiangxiensis sp. nov. and its similar species.

Character L. xiangxiensis sp. nov. L. melampyroides
Stems creeping or drooping. erect or ascending.
Plant height 15–25 cm 15–50 cm
Petiole not auriculate at base dilated and auriculate at base
Blades of lower leaves succulent, rhomboid-ovate to ovate, the basal 1 or 2 pairs scale-like papery, ovate to linear-lanceolate
Blades of upper leaves succulent, ovate to elliptic-lanceolate, 2–5.5 cm × 1–2.3 cm papery, ovate to linear-lanceolate,
1.5–9 × 0.3–2.5 cm
Secondary veins blurry or invisible on adaxial surface, slightly raising on abaxial surface visible on both surfaces
Glandular dots on leaves Absent transparent, sparse
Corolla lobes suborbicular to broadly elliptic, apex cuspidate or emarginated, 7–9 mm long and wide obovate-elliptic, apex rounded, 6–7 × 4–6 mm
Calyx lobes costa indistinct costa distinct

Phylogenetic position

The aligned lengths of ITS are 655 bp with gaps treated as missing data. BI and ML analyses produced similar topology and only the BI tree was presented in Figure 1. The phylogenetic results indicate that two samples of the new species were grouped together with a strong support (PP = 1.00, LP = 100%) and closely related to L. melampyroides (PP = 1.00, LP = 94%).

Figure 1. 

The phylogram of Bayesian inference (BI) tree from the ITS sequence data, showing the phylogenetic position of Lysimachia xiangxiensis sp. nov. (shown in bold). Values above the branches represent Bayesian posterior probabilities (PP) and bootstrap values (LP) for maximum likelihood, respectively; the dash (–) indicates LP < 50%.

On the basis of the classification in Handel-Mazzetti (1928), Chen and Hu (1979) divided the genus into five subgenera as well as many series, Subgen. Lysimachia, Subgen. Palladia (Moench) Hand.-Mazz, Subgen. Idiophyton Hand. -Mazz., Subgen. Naumburgia (Moench) Klatt and Subgen. Heterostylandra (Hand.-Mazz.) Chen et C. M. Hu. In this topology, all Lysimachia species form three main clades: Subgen. Lysimachia (PP = 1.00, LP = 70%), Subgen. Palladia (PP = 1.00, LP = 93%) and Subgen. Idiophyton. (PP = 1.00, LP = 80%). In addition, L. crispidens (Hance) Hemsley in F. B. Forbes & Hemsley of Subgen. Heterostylandra is close to Subgen. Palladia (PP = 1.00, LP = 91%) and L. longipes Hemsley is assigned to Subgen. Lysimachia with weak supported (PP = 0.54) in a neutral position between Subgen. Lysimachia and L. crispidens. But classification of series are not well reflected in this analysis.

Taxonomic treatment

Lysimachia xiangxiensis D.G.Zhang & C.Mou, Y.Wu, sp. nov.

Figure 2, 3, 4


CHINA. Hunan Province, Huayuan County, Buchou Town, Da-long-dong, cliff of a valley, 28°19'06.42"N, 109°30'03.22"E, alt. 295 m, 26 August 2019, D. G. Zhang 0826075 (holotype: JIU!; isotype: JIU!).


The new species differs from L. melampyroides by the succulent leaves; the creeping or drooping stems (15–25 cm long); and the suborbicular to broadly elliptic corolla lobes.


Terrestrial, perennial herbs. Rhizome brown, reduced to a small tuber or rarely creeping, with sparse fibrous roots. Stems creeping or drooping on cliffs, 15–25 cm long, clustered, branched at base, unbranched or rarely branched from the middle, terete, purple-red, densely strigillose, the internodes usually 3–7 cm long. Leaves petiolate, opposite. Petioles 5–7 mm long, with a furrow on adaxial side, green or purple-red, strigillose. Leaf blade succulent; blade of lower leaves rhomboid-ovate to ovate, with 1 or 2 pairs of basal leaves scalelike (much smaller); blade of upper leaves ovate to elliptic-lanceolate, 2–5.5 cm × 1–2.3 cm, base cuneate, apex acuminate or acute to subobtuse, margin entire and revolute, adaxially dark green, shiny, subglabrous, abaxially purple-red (in arid places) or light green (in moist places), densely strigillose along the midrib, not glandular on both surfaces; secondary veins 3–4 pairs, blurry or invisible adaxially, slightly raising abaxially, veinlets invisible. Flowers bisexually, solitary in axils of upper leaves, occasionally in terminal racemes with bractlike leaves. Pedicels 1.5–3 cm long, gradually reduced toward stem apex, purple-red or light purple-red, densely strigillose, recurved in fruit. Calyx lobes 5, rarely 6, persistent, lanceolate with indistinct costa, 6–8 mm × 1.5–2 mm, apex acuminate-subulate, inside glabrous and with 3–4 veins, outside purple-red or green, densely strigillose. Corolla yellow, tube 1–2 mm long, actinomorphic, contorted; lobes 5, 7–9 mm × 7–9 mm, suborbicular to broadly elliptic, apex cuspidate or rounded, erose above the middle. Stamens 5, yellow, opposite to corolla lobes; filaments connate basally into a tube ca. 2.5 mm high, free parts 3.5–4.5 mm; anthers ca. 2 mm long, dorsifixed, opening by lateral slits. Style ca. 6 mm long, apex slightly expanded, strigillose on lower part. Ovary cylindrical, ca. 1.5 cm in diam., strigillose on apex, superior. Capsule brown, subglobose, 3–4 mm in diam., densely strigillose, dehiscing by valves. Seeds small, black, angular, papillate.


Flowering May–June, fruiting July–August.

Distribution and habitat

This new species is currently known from Huayuan County and Jishou City in western Hunan Province, central China. It usually grows on limestone cliffs in valleys (Figure 2), and is associated with e.g. Eriophorum comosum (Wallich) Nees in Wight, Pteris vittata Linnaeus, Pteris deltodon Baker, and Dryopteris sp.

Figure 2. 

Lysimachia xiangxiensis sp. nov. in the wild A habitat (dry limestone cliff) B stems drooping on the cliff.

Figure 3. 

Lysimachia xiangxiensis sp. nov. in the field A corolla opened, showing the suborbicular lobes B flower (lateral view), showing the lanceolate calyx lobes indistinctly costate C proximal stems and underground part, showing stems clustered, rhizome, sparse fibrous roots, and 1 or 2 pairs of scalelike basal leaves D longitudinal section of flower, showing filaments connate basally into a tube E pistil, showing strigillose hairs on apex of ovary and base of style F dehiscent capsule G plant in fruiting, showing the recurved pedicels H plant in flowering, showing the solitary flowers in axils of upper leaves I plant in flowering, showing the reduced basal leaves J papillate seeds

Figure 4. 

Lysimachia xiangxiensis sp. nov. A upper stem leaf (abaxial surface), showing revolute margins B lower stem leaf (adaxial surface) C portion of a leaf (abaxial surface), showing the revolute margins and strigillose midrib D plant in flowering E corolla lobes F pistil.


The specific epithet “xiangxiensis”, literally meaning western Hunan, refers to the Xiangxi Tujia and Miao Autonomous Prefecture in central China, to which Huayuan County and Jishou City belong. The Chinese name of the Lysimachia xiangxiensis is xiang xi guo lu huang in Pinyin.

Conservation status

Lysimachia xiangxiensis usually grows on limestone cliffs in valleys so we suggest its placement in the Data Deficient category of IUCN (2017)

Additional collection

CHINA. Hunan Province, Jishou City, Aizhai Town, National Forest Park, cliff of a valley, 31 May 2019, Y. Wu 0531001(paratype, JIU!).


This work was supported by STS Program of the Chinese Academy of Sciences (No. KFJ-3W-No1) and the Traditional Chinese Medicine Public Health Special Project ([2017] 66). We thank Dr Wannachai Chatan for their thoughtful review of our submitted paper .


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Appendix S1.

Accessions of the genus Lysimachia L. examined in this study.

Taxon GenBank Acc. No. Voucher Locality
L. alfredii Hance JN638405 Hao394 Lianping, Guangdong, China
L. candida Lindley JF976885 Ge2010001 Yangchun, Guangdong, China
L. capillipes Hemsley in F. B. Forbes & Hemsley JF976886 Y2009200 Jiujiang, Jiangxi, China
L. chapaensis Merrill JF976888 GBOWS878 Hekou, Yunnan, China
L. chekiangensis C. C. Wu JF976891 Y2009263-1 Longquan, Zhejiang, China
L. christiniae Hance JF976894 Y2009244 Lin'an, Zhejiang, China
JF976896 Y2009209 Jiujiang, Jiangxi, China
L. clethroides Duby in A. de Candolle JF976899 Y2009157 Tongbai, Henan, China
L. congestiflora Hemsley in F. B. Forbes & Hemsley JF976902 GBOWS262 Malipo, Yunnan, China
JF976903 Y2009266 Longquan, Zhejiang, China
L. crispidens (Hance) Hemsley in F. B. Forbes & Hemsley JF976906 Hao212 Yichang, Hubei, China
L. decurrens G. Forster JF976908 GBOWS1234 Hekou, Yunnan, China
L. deltoidea Wight JF976909 GLM081121 Zhongdian, Yunnan, China
L. dextrosiflora X. P. Zhang, X. H. Guo & J. W. Shao JF976913 Y2009265-1 Longquan, Zhejiang, China
L. erosipetala F. H. Chen & C. M. Hu JF976914 Y2010037-2 Emeishan, Sichuan, China
L. fistulosa Handel-Mazzetti JF976916 Ning20101 Jinggangshan, Jiangxi, China
JF976917 Ye et al. 3561 Lianshan, Guangdong, China
JF976919 Y2009285 Ruyuan, Guangdong, China
JF976920 Ye et al. 3940 Lianshan, Guangdong, China
L. fortune Maximowicz JF976925 Y2009195 Jinggangshan, Jiangxi, China
L. hemsleyana Maximowicz ex Oliver JF976932 Guo20001 Ningguo, Anhui, China
L. hemsleyi Franchet JF976935 Hao713 Huili, Sichuan, China
L. heterobotrys F. H. Chen & C. M. Hu JF976936 Y2010053-2 Ningming, Guangxi, China
L. heterogenea Klatt JF976939 Y2009199 Jiujiang, Jiangxi, China
L. insignis Hemsley JF976945 Hao245 Napo, Guangxi, China
L. klattiana Hance JF976947 Y2010014-1 Tongbai, Henan, China
L. laxa Baudo JF976949 Han longran6 Puer, Yunnan, China
L. lobelioides Wallich in Roxburgh JF976951 Hao303 Menglian, Yunnan, China
L. longipes Hemsley JF976952 Guo xinhu200012 Shitai, Anhui, China
L. melampyroides R. Knuth in Engler JF976955 Dengyunfei15945 Xinning, Hunan, China
JF976956 Lichanghan8174 Shangzhi, Hunan, China
L. omeiensis Hemsley JF976958 Y2010033 Emeishan, Sichuan, China
L. paridiformis Franchet JF976962 Y2010044 Emeishan, Sichuan, China
L. patungensis Handel-Mazzetti JF976964 Ye et al. 3851 Lianshan, Guangdong, China
L. pentapetala Bunge JN638407 Y2010013-1 Tongbai, Henan, China
L. phyllocephala Handel-Mazzetti JF976969 Y2010030 Emeishan, Sichuan, China
L. pittosporoides C. Y. Wu JF976970 Hao248 Malipo, Yunnan, China
L. rubiginosa Hemsley in F. B. Forbes & Hemsley JF976972 Hao419 Dujiangyan, Sichuan, China
Lysimachia xiangxiensis D.G.Zhang & C.Mou, Y.Wu, sp. nov. MN647745 Y. Wu 0531001 Jishou, Hunan, China
MN647744 D. G. Zhang 0826075 Huayuan, Hunan, China
Ardisia verbascifolia Mez JN638408 GBOWS1216 Hekou, Yunnan, China