Research Article
Research Article
Viola shiweii, a new species of Viola (Violaceae) from karst forest in Guizhou, China
expand article infoXiao-Chen Li§, Zheng-Wei Wang§, Qi Wang§, Bin-Jie Ge§, Bin Chen§, Ping Yu|, Xin Zhong§
‡ Eastern China Conservation Center for Wild Endangered Plant Resources, Shanghai, China
§ Shanghai Chenshan Botanical Garden, Shanghai, China
| Guizhou Maolan Karst Forest, Libo, China
Open Access


Viola shiweii Xiao C. Li & Z. W. Wang (Violaceae), a new species from Guizhou, China, is described, based on morphological and molecular evidence. The new species is morphologically most similar to V. kwangtungensis Melchior in its glabrous lateral petals and stoloniferous habit, but can be distinguished by its obtuse teeth along the leaf margin, its abaxially greyish-green leaf blade and its broader and entire sepals with a distinct basal appendage.


Morphology, phylogeny, sect. Plagiostigma, subg. Viola


Viola L. is the largest genus amongst the Violaceae, comprising approximately 580–620 species (Wahlert et al. 2014; Marcussen et al. 2015), which are widely distributed in temperate regions and tropical high mountain regions worldwide, with south-western China as one of its current centres of diversity (Wahlert et al. 2014). The diversity and high number of species has resulted in an extremely complex interspecific relationship in this genus due to hybridisation and horizontal evolution amongst sections and species (Marcussen et al. 2015). Since Becker (1925) provided the first infrageneric classification for Viola, several infrageneric classifications of the genus have been proposed (Clausen 1927, 1929, 1931, 1964; Gershoy 1934). In the latest taxonomical revision of Viola of China, 96 species were recognised as native (Chen et al. 2007). However, delimitation of the species with stolons distributed in southern and south-western China remains highly problematic and new species are still being discovered (Zhou and Xing 2007; Chen and Yang 2008, 2009; Zhou et al. 2008a; Dong et al. 2009; Ning et al. 2012; Huang et al. 2021).

During an expedition to Guizhou Province in November 2019, an unfamiliar violet whose habit was somewhat similar to that of Viola kwangtungensis Melchior caught the authors’ attention on the karst rock outcrops. Several specimens with cleistogamous flowers were collected from the field and living material was transplanted and cultivated in Chenshan Botanical Garden for further observation.

Materials and methods

In this study, molecular phylogenetic analysis, based on the ITS dataset, was firstly conducted to resolve the phylogenetic position of the unfamiliar violet and its relationship with V. kwangtungensis Melchior. Subsequently, morphological characters of this unfamiliar violet and its related species were compared, based on living plants and herbarium specimens, including the digital resource of the Chinese Virtual Herbarium ( and the China Field Herbarium ( Herbarium specimens were examined in IBK and CSH. Original protologues and relevant literature were also investigated. Leaf material of the putative new species and its related species was collected and stored with silica. Six species, represented by eight individuals, were newly sampled. Voucher specimens were deposited in Chenshan Herbarium (CSH). Total genomic DNA was extracted with the modified CTAB method (Doyle and Doyle 1987) for library construction at Benagen ( Paired-end sequencing of the whole sequences from both ends of 150 bp fragments was performed on the DNBSEQ T7, about 2 Gb clean data for every sample were produced. The nrDNA were de novo assembled using the GetOrganelle pipeline (Jin et al. 2020) and sequences of ITS1-5.8s-ITS2 were extracted with ITSx 1.1.3 (Bengtsson-Palme et al. 2013). Another 31 sample sequences were obtained from NCBI (Gong et al. 2010; Liang and Xing 2010). The sequences of the species and related ones were aligned in Geneious Prime 2021.2.2 ( using MAFFT (Katoh and Standley 2013) by default setting. Phylogenetic construction was conducted by Maximum Likelihood with IQ-Tree 2 software (Minh et al. 2020), selecting the best-fit model of GTR+F+G4 with 2000 bootstraps. The tree file was visualised and annotated on iTOL ( (Ivica and Peer 2021). All the sequences accession numbers were listed in Table 1.

Table 1.

Vouchers of specimens and GenBank accession number.

Taxon Voucher Accession no.
Ingroup taxon
sect. Diffusae (W.Becker) C.J.Wang
ser. Australasiaticae Okamoto
V. mucronulifera Hand.-Mazz. Lingyun, Guangxi, Zhou J S 311 (IBSC) FJ002910
V. sumatrana Miq. Lvchun, Yunnan, Wang Zheng-wei et al.WZW04206 (CSH) OM406231
V. kwantungensis Melchior Guidong, Hunan, Huang Cun-zhong LXC01887 (CSH) OM406227
V. kwantungensis Melchior Jinyunshan, Chongqing, Huang Yan-shuang HYS210206 OM406230
V. kwangtungensis Melchior Malipo, Yunnan, Wang Zheng-wei et al. WZW04187 (CSH) OM618008
V. austrosinensis Y.S.Chen & Q.E.Yang Tianlin, Guangxi, Li Xiao-chen et al. LXC02318 (CSH) OM406228
V. davidii Franch. Mt. Maoershan, Guangxi, Zhou J S 273 (IBSC) FJ002902
V. davidii Franch. Mt. Jiulongshan, Zhejiang, Zhong Xin et al. ZX01824 (CSH) OM406229
V. grandisepala W.Becker Mt. Emeishan, Sichuan, Zhou J S 425 (IBSC) FJ002903
V. fargesii H.Boissieu originally published as V. principis Ruyuan, Guangdong, Zhou J S 103 (IBSC) FJ002904
Viola sp. nov. Maolan, Guizhou, Li Xiao-chen et al. LXC00927 (CSH) OM406226
Viola sp. nov. Maolan, Guizhou, Li Xiao-chen et al. LXC00323 (CSH) OM406225
Viola sp. nov. Maolan, Guizhou, Li Xiao-chen et al. LXC00324 (CSH) OM406224
ser. Diffusae (W.Becker) Steenis
V. nanlingensis J.S.Zhou & F.W.Xing Mt. Nankunshan, Guangdong, Liang G. X. 0185 (IBSC) FJ002916
V. yunnanensis W.Becker & H.Boiss. Mt. Diaoluoshan, Hainan, Zhou J. S. s.n. (IBSC) FJ002915
V. diffusa Ging Huaiji, Guangdong, Gong Q. 00043 (IBSC) FJ002917
V. lucens W.Becker Tanziyan, Guizhou, Zhou J. S. 348 (IBSC) FJ002913
V. guangzhouensis A.Q.Dong Conghua, Guangdong, Dong A. Q. 1104 (IBSC) FJ002918
sect. Chamaemelanium Ging
V. biflora L. FJ002905
V. orientalis (Maximowicz) W.Becker FJ002909
V. delavayi Franch. Diqing, Yunnan, Zhou J. S. Xing F. W. 487 (IBSC) FJ002908
sect. Viola L.
V. collina Bess. FJ002880
V. mirabilis L. MK828568
V. rupestris F.W.Schmidt HM851448
V. grypoceras A.Gray Mt. Lushan, Jianghxi, Liang G. X. 0002 (IBSC) FJ002881
V. acuminata Ledeb. FJ002884
sect. Violidium (K. Koch) Juz.
V. inconspicua Blume SCBG, Guangdong, Liang G. X. 0187 (IBSC) FJ002897
V. japonica Langsdorff ex Candolle EU591965
V. prionantha Bunge Jinan,Shandong, Zhang R. J., Xing F. W. 17955 (IBSC) FJ002893
V. hancokii W.Becker FJ002890
V. pekinensis (Regel) W.Becker FJ002892
V. chaerophylloides (Regel) W.Becker FJ002898
V. dissecta Ledeb. FJ002891
V. magnifica C. J. Wang & X.D.Wang Mt. Lushan, Jiangxi, Liang G. X. 0038 (IBSC) FJ002899
sect. Bilobatae (W.Becker) Juz.
V. verecunda A.Gray Mt. Nanling, Guangdong, Zhou J. S. 1553 (IBSC) FJ002911
V. triangulifolia W.Becker Mt. Jinggangshan, Jiangxi, Zhou J. S. 140 (IBSC) FJ002912
outgroup taxon
Afrohybanthus enneaspermus (L.) Flicker HM483598


Molecular Analysis

The ITS dataset comprises 37 accessions representing 32 species, including Afrohybanthus enneaspermus (L.) Flicker selected as an outgroup. The aligned matrix of ITS sequences was 696 bp in total. The result of ML is shown in Fig. 1. The samples of the putative new species (pink clade) clustered into a strongly supported monophyletic lineage (clade 1), forming a weak sister relationship with a clade formed by V. mucronulifera and V. sumatrana. The morphologically most similar V. kwangtungensis was resolved on a more distant phylogenetic position (clade 2, blue clade). Based on morphological characters and phylogenetic results, we recognise this unfamiliar violet as a distinct species and described it here as Viola shiweii Xiao C. Li & Z.W.Wang.

Figure 1. 

ML tree of the new species Viola shiweii sp. nov. and its related species inferred by IQ-Tree 2, based on ITS dataset. Bootstrap values of the Maximum Likelihood are shown along the branches. Outgroup taxon: Afrohybanthus enneaspermus.

Taxonomic treatment

Viola shiweii Xiao C. Li & Z. W. Wang, sp. nov.

Figs 2, 3, Appendix 1: Figs A1, A2, A3, A4, A5, A6, A7, A8, A9, A10, A11, A12


China. Shanghai Chenshan Botanical Garden, cultivated plants collected from Guizhou, Qiannan Buyi and Miao Autonomous Prefecture (黔南布依族苗族自治区), Libo county (荔波县), Maolan National Nature Reserve (茂兰国家级自然保护区), on the rocks along the karst forest margin, 25°16'39.1039"N, 107°55'2.7598"E, 867 m elevation, 9 Nov 2019, Wang Zheng-wei and Li Xiao-chen, LXC00927 Holotype: CSH0182173 (CSH!); isotypes: ZJFC!, CSFI!, IBSC!.

Figure 2. 

Viola shiweii sp. nov. A flower side view B flower front view C habit D stipule E capsule F stamens and pistil.


The new species is morphologically most similar to V. kwangtungensis Melchior in its glabrous lateral petals and stoloniferous habit, but can be distinguished by its obtuse teeth along the leaf margin, its abaxially pale green leaf blade and its broader and entire sepals with a distinct basal appendage.


Perennial herb, acaulescent, with stolons. Rhizome short, straight or oblique, densely noded, usually covered by brown remains of stipules. Stipules free, brown, broadly lanceolate, 5–10 mm long, margin long fimbriate-dentate, lobes remotely dentate. Basal leaves glabrous, slightly grooved, with petioles stout, petioles unequal in length; blade thick leathery, ovate or suborbicular, 15–30 × 15–20 mm, base deep cordate, apex usually obtuse, abaxially green, scabrous, abaxially greyish-green, mid-vein distinct above, glabrous on both surfaces, margin glandular-serrate or shallowly glandular-crenate, slightly wavy, teeth obtuse at the apex; stolon leaves scattered, smaller. Pedicel equal to or longer than petiole, two bracts narrowly lanceolate, at the middle or lower part of the pedicel. Sepals 5, ca. 6 mm long and 2 mm wide, lanceolate, glabrous, margin narrowly membranous, apex somewhat acute, base distinctly decurrent, apex obtuse or shallowly dentate. Flower 1.5–2.5 cm in diameter, petals 5, white, posterior and lateral ones obovate, ca. 1.2 cm × 5 mm, narrow at the base, lateral petals purplish near the middle, glabrous, anterior petal shorter, ca. 10 mm (spur included) long, oblong, purple-veined, apex rounded, obtuse, spur saccate, 2–3 mm long and 1.5 mm wide. Style clavate, base geniculate, stigmas flattened on top, narrowly margined on lateral sides and abaxially, shortly beaked ventrally. Capsule ellipsoid, valves carinate, ca. 10 mm long and 2.5 mm wide, glabrous. Seeds black, ca. 2 mm long and 1 mm in diameter.

Figure 3. 

Viola shiweii sp. nov. A petals, sepals, stamens and pistils B, G flower in front view and detail of a longitudinal section in side C, H basal leaf adaxially and abaxially D cleistogamous flowers E capsule and seeds F margin teeth I stipule J leaves on stolons.

Distribution and habitat

The species was observed to grow on dry and partially shaded limestone, around the karst forest edge, at 700–900 m elevation.

Additional specimens examined

China, Guizhou, Qiannan Autonomous Prefecture, Libo County, Maolan National Nature Reserve, karst forest, 24 Jul 2008, Zhang Dai-Gui 080724077 (JIU!); China, Guizhou, Qiannan Buyi and Miao Autonomous Prefecture, Libo County, Maolan National Nature Reserve, 21 Nov 2021, Li Xiao-chen, Wang Zheng-wei & Wei Hong-jin, LXC02320 (CSH!), LXC02322 (CSH!), LXC02323 (CSH!), LXC02324 (CSH!), LXC02325 (CSH!).


Cultivated plants flower in September-March, fruiting in September.


The specific name epithet “shiweii” was proposed in memory of Deng Shi-wei (191?-1936), who dedicated his life to the exploration of the flora of Guizhou. The Chinese name is given as “世纬堇菜”.

Conservation status

Only two populations of V. shiweii are currently known from Maolan National Nature Reserve, Libo County, in an area of the karst formation across Guizhou and Guangxi (Fig. 4). This species is represented by no more than 200 large and mature individuals. Due to its rarity, the low number of individuals and habitat vulnerability, V. shiweii is considered to be Critically Endangered (CR, B1), according to the IUCN (2019).

Figure 4. 

Habitat and distribution of Viola shiweii.


Although our phylogenetic analysis, based on ITS sequences, did not fully clarify the infrageneric relationships within Viola, it produced informative evidence for differentiation amongst lower taxa. V. shiweii can be placed in Viola ser. Australasiaticae (Okamoto et al. 1993), which is characterised by the stolons with scattered leaves, absent aerial stems, short spur of anterior petal and stigma beaked ventrally. The phylogenetic analysis in this study (Fig. 1) also confirmed this conjecture (Fig. 1); however, the monophyly of Viola ser. Australasiaticae was not supported, which was consistent with a previous study (Gong et al. 2010). Viola ser. Australasiaticae was proved to be nested in the subg. Viola sect. Plagiostigma Godr. (Marcussen et al. 2012).

Viola ser. Australasiaticae comprises ca. 27 species, widely distributed in the Himalayan Region, southern China, south-eastern Asia and Ryukyu Island of Japan, with 14 species occurring in China (Chen 2006), in which Viola davidii, V. schneideri W.Becker, V. kwangtungensis, V. mucronulifera and V. austrosinensis form a complex in this series and caused mass misidentification due to their high degree of morphological similarity.

Viola davidii Franchet was published by Adrien René Franchet (1885), based on the collection of David from Moupine (Baoxing County, Sichuan, China) (isotype: David#s.n., K000254222) in 1869 [1870]. It is a morphologically variable and widespread species characterised by its ovate or ovate-orbicular leaf blade with 6–8 rounded teeth along each side, bearded lateral petals and short spurred anterior petal. It was originally regarded as species similar to V. biflora L., but its beaked stigma (vs. bilobed), white and purple petals (vs. yellow) indicated a distinctly different affiliation amongst the genus. Later, Becker (1921) described a strikingly similar violet with ovate leaves, V. schneideri W.Becker, based on the collection of Schneider (isotype: Schneider C.K. #739, G00343327) from Te-chang (De-chang County, Sichuan, China), but the diagnostic leaf shape falls within the morphological variation of V. davidii Franchet and, for this reason, it was recently treated as its synonym, which is further supported by overlapping distributions (Chen 2006). The only collection of V. shiweii before this study was misidentified as V. davidii. Viola shiweii shares a similar leaf shape with V. davidii, but can be differed by its glabrous lateral petals and obtuse teeth along the leaf margin.

Viola kwangtungensis Melchior, which shows the highest resemblance to V. shiweii (Table 2, Figs 5, 6) is an overlooked species frequently being confounded with V. davidii or V. mucronulifera, but considered as a distinct taxon by Flora of China (Chen et al. 2007). Viola mucronulifera Hand.-Mazz. was published by Handel-Mazzetti (1931), based on the collection of R. C. Ching #7016 from Guangxi (holotype: PE00025463, isotypes: NY00097644 & A00067198) and is characterised by the distinctly stipitate tooth glands. Later, V. kwangtungensis Melchior (1933) was published, based on a collection of Woon-Young Chun and his assistants (isotype: P. Ko #50326, A00067196) from Guangdong, which can be easily recognised by its characteristic leaf crenation of the leaves, but it was subsequently reduced to a synonym of V. mucronulifera in Flora Reipublicae Popularis Sinicae (Wang 1991). Viola kwangtungensis has spinules at the apex of the teeth, as the horizontal extension of the teeth, while the spinules of V. mucronulifera are perpendicular to the leaf blade and placed between the teeth, which can be distinguished in field observation. Viola kwangtungensis also used to be considered conspecific with V. schneideri due to morphology transition in the spinose, based only on the specimen observation (Zhou et al. 2008b).

Table 2.

Morphology and distribution comparison between Viola shiweii sp. nov., V. kwangtungensis and Viola davidii.

Viola shiweii Viola kwangtungensis Viola davidii
Leaf blade Ovate or orbicular, apex usually obtuse, base deep cordate, greyish-green abaxially. Ovate to triangularly ovate, base shallowly cordate, apex usually acute, purple abaxially. Ovate or ovate-orbicular, glaucous abaxially, base deeply coedate, apex rounded or acute.
Leaf margin Serrate Crenate Shallowly 6–8-crenate on each side.
Stipule Long fimbriate Fimbriate Remotely fimbriate-dentate
Sepals Lanceolate, ca. 6 mm × 2 mm, entire, green, glabrous, basal distinctly decurrent. Lanceolate, 3–5 mm × 1.5–2 mm, sparsely shallowly dentate, purplish-red, sparsely pubescent, base not decurrent. Lanceolate or ovate-lanceolate, 5–6 mm × 1.5–2 mm, brown, glabrous, base shortly decurrent, margin narrowly membranous, apex truncate
Posterior and lateral petals Obovate, base constricted Obovate-oblong Oblong-obovate
Seed Black Brown Brown
Habitat Dry and partially shaded limestone Humid and shaded stream valley Shaded place under forest, stream valley, or grassy slope.
Distribution Guizhou, Guangxi. Fujian, N Guangdong, Guizhou, Hunan, Jiangxi, Sichuan, Yunnan, and Taiwan (Lu et al. 2019) Chongqing, Fujian, Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangxi, Sichuan, SE Xizang, Yunnan, Zhejiang.
Figure 5. 

Flowers and leaves of Viola shiweii sp. nov. and Viola kwangtungensis A, B V. kwangtungensis, lower, front and side views C, D V. kwangtungensis, adaxial and abaxial leaf surfaces E, F Viola shiweii, flower, front and side views G, H V. kwangtungensis, adaxial and abaxial leaf surfaces.

More recently, as the latest supplement of this complex, a new species, V. austrosinensis, distinguished from V. kosanensis Hayata (ser. Rosulantes Borbas (Y.S.Chen)), was described, of which the leaves were coriaceous, glabrous, not glandular-dotted on the abaxial surface (Chen and Yang 2008). Viola austrosinensis is different from V. shiweii in its ovate leaf blade and acute anterior petal’s apex.

In China, V. mucronulifera was found to occur only in the Province of Yunnan and its type locality in Guangxi; its occurrence in Guizhou was a mistake caused by the misidentification of V. kwangtungensis in Flora of Guizhou (Yao 1989), as we personally observed in the locality cited in this work (Fanjingshan, Jiangkou). The morphology and distribution differences between V. shiweii, V. kwangtungensis and V. davidii are listed in Table 2. Comparision of V. shiweii and V. kwangtungensi was visualised in Figs 5, 6. Keys to V. shiweii and its allies were also presented.

Figure 6. 

Leaf margin of V. shiweii sp. nov. and V. kwangtungensis A V. shiweii, holotype B V. kwangtungensis, isotype: P. Ko #50326, A00067196.

1 Spur 5–7 mm, anterior petal 2-lobed at apex V. formosana
Spur shorter than 5 mm, anterior petal rounded, obtuse or acute at apex 2
2 Stipules usually entire, sepals broad ovate, ca. 5 mm wide V. grandisepala
Stipules fimbriate, sepals lanceolate, much narrower, not more than 5 mm 3
3 Leaf blade spinulose along margin 4
Leaf blade without spinules along margin 5
4 Leaves conspicuously spinose between teeth V. mucronulifera
Leaves shortly spinose at apex of teeth V. kwangtungensis
5 Leaves ovate, orbicular or nearly orbicular, apex obtuse 6
Leaves cordate or oblong-ovate, apex acuminate 10
6 Lateral petals beard at base V. davidii
Lateral petals glabrous at base 7
7 Leaves serrata, teeth have obtuse apices, apex of anterior petal obtuse V. shiweii
Leaves crenate, without obtuse teeth along margin 8
8 Leaves coriaceous, base shallowly cordate, anterior petal acute V. austrosinensis
Leaves chartaceous, orbicular or nearly orbicular, base deeply cordate, anterior rounded 9
9 Leaves adaxially scabrous, sparsely pubescent V. duclouxii
Leaves adaxially shiny, glabrous V. sikkimensis
10 Rhizome short, densely noded 11
Rhizome nodes elongated and stout 12
11 Leaves glabrous, shiny adaxially V. nitida
Leaves densely pubescent V. fargesii
12 Plant densely pubescent V. yunnanensis
Plant glabrous or sparsely pubescent 13
13 Leaves blade glabrous, sepals ovate V. nuda
Leaves more or less pubescent, sepals lanceolate 14
14 Leaves and capsules dot-like brown glandular, lateral petals glabrous V. sumatrana
Leaves not glandular, lateral petals bearded V. thomsonii


Our deepest gratitude goes to three reviewers Thomas Marcussen, Juliana de Paula-Souza, and Chen You-sheng and Subject editor Yasen Mutafchiev, for their careful work and thoughtful suggestions that have helped improve this paper substantially. The authors are grateful to Mr. Yu Tian-yi for his excellent illustration in the manuscript, Dr. Jiang Kai from Chenshan Botanical Garden for his help in data analysis and Dr. Huang Yan-shuang from Sun Yat-Sen University, Dr. Zhu Xin-xin from Xinyang Normal University for their generous offer of samples and collection information. The first author is also indebted to Dr. Huang Yu-song and Huang Jin-quan from IBK for hosting our visit and Miss Pi for her company during the epidemic of COVID-19. This study was supported by the project of the National Wild Plant Germplasm Resource Center for Shanghai Chenshan Botanical Garden (ZWGX2102), the project of the Special Fund for Scientific Research of Shanghai Landscaping & City Appearance Administrative Bureau (G212416, G222404).


  • Becker W (1925) Viola. In: Engler A, Prantl K (Eds) Die Natürlichen Pflanzenfamilien (2nd edn. ) 21. Duncker & Humblot, Berlin, 363–376.
  • Bengtsson‐Palme J, Ryberg M, Hartmann M, Branco S, Wang Z, Godhe A, De Wit P, Sánchez-García M, Ebersberger I, de Sousa F, Amend AS, Jumpponen A, Unterseher M, Kristiansson E, Abarenkov K, Bertrand YJK, Sanli K, Eriksson KM, Vik U, Veldre V, Nilsson RH (2013) Improved software detection and extraction of ITS1 and ITS 2 from ribosomal ITS sequences of fungi and other eukaryotes for analysis of environmental sequencing data. Methods in Ecology and Evolution 4(10): 914–919.–210X.12073
  • Chen YS, Yang QE, Ohba H, Nikitin VV (2007) Violaceae. In: Wu ZY, Raven PH (Eds) Flora of China, vol. 13. Science Press, Beijing & Missouri Botanical Garden Press, St. Louis, 72–111.
  • Gershoy A (1934) Studies in North American violets. III. Chromosome numbers and species characters. Vermont Agricultural Experiment Station Bulletin 367: 1–91.
  • Gong Q, Zhou JS, Zhang YX, Liang GX, Chen HF, Xing FW (2010) Molecular Systematics of Genus Viola L. in China. Redai Yaredai Zhiwu Xuebao 18(6): 633–642.
  • Ivica L, Peer B (2021) Interactive Tree Of Life (iTOL) v5: An online tool for phylogenetic tree display and annotation. Nucleic Acids Research 49(W1): W293–W296.
  • Jin JJ, Yu WB, Yang JB, Song Y, dePamphilis CW, Yi TS, Li DZ (2020) GetOrganelle: A fast and versatile toolkit for accurate de novo assembly of organelle genomes. Genome Biology 21(1): 1–31.
  • Katoh K, Standley DM (2013) MAFFT multiple sequence alignment software version 7: Improvements in performance and usability. Molecular Biology and Evolution 30(4): 772–780.
  • Liang GX, Xing FW (2010) Infrageneric phylogeny of the genus Viola (Violaceae) Based on trnL-trnF, psbA-trnH, rpL16, ITS Sequences, Cytological and Morphological Data. Yunnan Zhi Wu Yan Jiu 32(6): 477–488.
  • Marcussen T, Jakobsen KS, Danihelka J, Ballard HE, Blaxland K, Brysting AK, Oxelman B (2012) Inferring species networks from gene trees in high-polyploid North American and Hawaiian violets (Viola, Violaceae). Systematic Biology 61(1): 107–126.
  • Marcussen T, Heier L, Brysting A, Oxelman B, Jakobsen K (2015) From gene trees to a dated allopolyploid network: Insights from the angiosperm genus Viola (Violaceae). Systematic Biology 64(1): 84–101.
  • Melchior H (1933) Viola kwangtungensis, a new violet from China. Sunyatsenia 1(203): 124–126.
  • Minh BQ, Schmidt HA, Chernomor O, Schrempf D, Woodhams MD, Haeseler A, Lanfear A (2020) IQ-TREE 2: New models and efficient methods for phylogenetic inference in the genomic era. Molecular Biology and Evolution 37(5): 1530–1534.
  • Ning ZL, Zeng ZX, Chen L, Xu BQ, Liao JP (2012) Viola jinggangshanensis (Violaceae), a new species from Jiangxi, China. Annales Botanici Fennici 49(5): 383–386.
  • Okamoto M, Okada H, Ueda K (1993) Morphology and chromosome number of Viola pilosa, and its systematic position. Taxon 42(4): 781–787.
  • Wahlert GA, Marcussen T, de Paula-Souza J, Feng M, Ballard Jr HE (2014) A phylogeny of the Violaceae (Malpighiales) inferred from plastid DNA Sequences: Implications for generic diversity and intrafamilial classification. Systematic Botany 39(1): 239–252.
  • Yao LZ (1989) Violaceae. In: Editorial Committee of Flora Guizhouensis (Eds) Flora Guizhouensis, Vol. 4. Sichuan Minorities Press, Chengdu, 228.
  • Zhou JS, Gong Q, Xing FW (2008a) Viola nanlingensis (Violaceae), a new species from Guangdong, southern China. Annales Botanici Fennici 45(3): 233–236.

Appendix 1. Type specimens of V. shiweii

Figure A1. 

Holotype: LXC00927 (CSH0182173).

Figure A2. 

Isotype: LXC00927 (CSH0182170).

Figure A3. 

Isotype: LXC00927 (CSH0182171).

Figure A4. 

Isotype: LXC00927 (CSH0182172).

Figure A5. 

Paratype: LXC02320 (CSH0189181).

Figure A6. 

Paratype: LXC02322 (CSH0189180).

Figure A7. 

Paratype: LXC02323 (CSH0189177).

Figure A8. 

Paratype: LXC02323 (CSH0189178).

Figure A9. 

Paratype: LXC02323 (CSH0189179).

Figure A10. 

Paratype: LXC02324 (CSH0189176).

Figure A11. 

Paratype: LXC02325 (CSH0189174).

Figure A12. 

Paratype: LXC02325 (CSH0189175).

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