Research Article
Research Article
Primulina malipoensis (Gesneriaceae), a new species from Sino-Vietnamese border area
expand article infoLi-Hua Yang§, Jun-Lin Chen|, Fang Wen, Ming Kang
‡ South China Botanical Garden, Chinese Academy of Sciences, Guangzhou, China
§ University of Chinese Academy of Sciences, Beijing, China
| College of Humanities Sichuan Agricultural University, Ya’an, China
¶ Guangxi Institute of Botany, Guangxi Zhuang Autonomous Region and Chinese Academy of Sciences, Guilin, China
Open Access


Primulina malipoensis, a new species from limestone areas around the Sino-Vietnamese border, is described and illustrated. This new species is morphologically similar to P. maguanensis and P. lungzhouensis, but obviously differs from the latter two species by its pale greenish-yellow flowers (vs. purple, with different colour patterns). The phylogenetic affinity, illustration and photographs of this new species are provided in this paper.


Limestone flora, New taxon, Sino-Vietnamese border area, Taxonomy


The recently redefined Primulina Hance has become a species-rich genus within the subfamily Didymocarpoideae of Gesneriaceae (Wang et al. 2011, Weber et al. 2011, 2013) and its species diversity is still growing due to numerous new species being constantly discovered (e.g. Pan et al. 2013, Guo et al. 2015, Lai and Wen 2015). This group shows high levels of endemism and ecological (edaphic) specialisation (Hao et al. 2015). The majority of its species occur in karst areas of southern and southwestern China and northern Vietnam, with narrow, island distributions, often limited to a single cave or karst limestone hill system (Wang et al. 1998, Li and Wang 2004, Wei et al. 2010). Local-scale mosaics of soil type are ubiquitous features in the karst landscapes and thus, soil nutrient availability may influence diversification and speciation of Primulina via local adaptation to specific edaphic microhabitats (Hao et al. 2015). However, in the P. eburnea complex, geographical isolation has been shown to be a major driver of its diversification and speciation in Primulina (Gao et al. 2015, Wang et al. 2017).

During field explorations in 2013, one of the authors (JC) found an unknown species of Primulina near the Sino-Vietnamese border at Malipo County, southeastern Yunnan, China. Several living individuals from the population found in the field were brought to the South China Botanical Garden (SCBG) and cultivated there. These plants showed leaf blade characteristics very common in Primulina. However, when flowering, they displayed uncommon yellow flowers. Flower colour has been used as an important character for the description of new Primulina species (Pan et al. 2016, Yang et al. 2017). Therefore, these plants soon caught the authors’ attention. Checking of specimens and literature studies were undertaken immediately. When specimens were checked in KUN (by its online service), an interesting specimen was found (numbered KUN 1275938), which possesses a similar leaf to these plants and had been collected from nearly the same locality as the findings. This specimen was identified as Chirita eburnea (a synonym to P. eburnea). However, this specimen was represented by only a piece of leaf and without flowers, thus, its identification is doubtful. To further reveal the true taxonomic identity of both of these plants and the specimen, other field works were carried out by one of the authors (FW) in 2017. Fortunately, he found this species at the recoded site of the specimen (KUN 1275938) and also found other populations at a nearby location in Vietnam. At the same time, additional investigations, i.e. phylogenetic analysis and morphological comparison, were undertaken. Based on these results, all of these plants from the three populations are considered as the same new species, which is described and illustrated here.


Morphological observations were carried out using living cultivated plants (ten individuals) as well as dried specimens. All morphological characters were measured using dissecting microscopes and descriptions were made following the terminology presented by Wang et al. (1998). Literature studies included all relevant monographs (Wang et al. 1998, Li and Wang 2004, Wei et al. 2010) and recently published literature (Xu et al. 2008, 2012, Li and Möller 2009, Pan et al. 2013, 2016, Li et al. 2014, Lai and Wen 2015, Guo et al. 2015). Checking of specimens was undertaken at IBSC and IBK and with the help of web databases (Chinese Virtual Herbarium:; Herbarium, Kunming Institute of Botany, CAS:; Global Plants: A map of the species’ geographical distribution was prepared based on field records. The molecular phylogenetic analyses of the species were included in a broader study in which the most comprehensive species-level phylogeny of Primulina was reconstructed based on 20 plastid and nuclear regions (Kong et al. 2017).


Primulina malipoensis L.H. Yang & M. Kang, sp. nov.

Figures 1, 2


Primulina malipoensis mainly differs from P. maguanensis and P. lungzhouensis by its pale greenish-yellow flowers (vs. purple, with different colour patterns). This new species can further be distinguished from P. maguanensis by its greenish bracts (vs. white) and from P. lungzhouensis by its entire bracts margin (vs. denticulate).


CHINA. Guangdong Province, Guangzhou City, voucher from a cultivated plant at South China Botanical Garden, 29 July 2016 (flowering), Li-Hua Yang, YLH369 (holotype: IBSC!), introduced from Yunnan province, Malipo county, Xiajinchang town, growing on moist limestone rocks, Alt. 1500 m, 23°10'N, 104°49'E, 31 August 2013, Jun-lin Chen.


Perennial herbs. Rhizomatous stem subterete, 20–60 mm long, 5–15 mm in diameter. Leaves 8–12, basal or clustered at apex of stem, opposite decussate. Petiole flattened, 20–40 mm long, 8–10 mm wide, pubescent. Leaf blade slightly fleshy when fresh, thickly chartaceous when dried, ovate to broadly elliptic, 7–12 × 7–10 cm, adaxially densely pubescent, abaxially glabrescent and only puberulent along veins, apex subacute to obtuse, base cuneate, margin inconspicuously serrate; lateral veins 4 on each side, abaxially conspicuous. Cymes 3–5, axillary, 2–4 branched, 8–16-flowered; peduncles 15–27 cm long, ca. 2 mm in diameter, densely pubescent; bracts 2, sometimes with bracteoles (narrowly ovate, 8–12 × 4–7 mm), green, opposite, ovate, 16–25 × 8–13 mm, margin entire, apex acute, outside densely pubescent, inside sparsely pubescent. Pedicel 10–14 mm long, ca. 1 mm in diameter, densely glandular pubescent and puberulent. Calyx 5-parted to near base, lobes narrowly lanceolate, white, 7–9 × ca. 2 mm, outside densely glandular pubescent and puberulent, inside sparsely pubescent, margin entire. Corolla pale greenish-yellow, 24–32 mm long, outside glandular-pubescent and puberulent, inside glabrous; tube infundibuliform, 21–25 mm long, ca. 8 mm in diameter at mouth, ca. 5 mm in diameter at base; limb distinctly 2-lipped, adaxial lip 2-parted, lobes broadly ovate, 7–9 × 6–7 mm, apex rounded, abaxial lip 3-lobed, lobes oblong, 11–13 × 5–7 mm, apex rounded. Stamens 2, adnate to 10–13 mm above the corolla tube base; filaments linear, 9–11 mm long, pale greenish-yellow, geniculate near middle, sparely pubescent; anthers fused by the entire adaxial surfaces, ca. 2 mm long, abaxially densely covered with glandular hairs. Staminodes 3, lateral ones 6–7 mm long, adnate to 10–12 mm above the corolla tube base, middle one ca. 1.5 mm long, adnate to 6–8 mm above the corolla tube base. Disc annular, ca. 1.5 mm in height. Pistil 22–26 mm long; ovary cylindrical, 15–18 mm long, ca. 1.5 mm in diameter, densely glandular pubescent and puberulent; style ca. 7 mm long, densely glandular-pubescent and puberulent; stigma 1, its upper lobe lacking, lower lobe obtrapeziform, shallowly 2-lobed at apex, ca. 2 mm long, ca. 1.5 mm wide. Capsule linear, ca. 30 mm long, densely pubescent.

Figure 1. 

Primulina malipoensis. A habit B flower in front view C flower in side view D opened corolla, showing stamens and staminodes E fertile stamens F pistil and stigma G staminodes H bract. Drawn by Yun-Xiao Liu based on a cultivated individual collected from type locality.

Figure 2. 

Primulina malipoensis. A flowering plant cultivated in South China Botanical Garden B plant in natural habitat C flower in side view D opened corolla, showing stamens and staminodes E flower in front view F pistil and calyx G bracts. Photographs by Li-Hua Yang.

Figure 3. 

Primulina maguanensis (A, B), P. lungzhouensis (D, E), P. beiliuensis var. fimbribracteata (C, F), P. beiliuensis var. beiliuensis (G, H) and P. maculata (I, J). (A, C, D, G, J) habit, (B, E, F, H, I) flower. Photographs by Fang Wen (A–H) and Li-Hua Yang (I, J).

Distribution and habitat

Primulina malipoensis is a narrowly endemic species restricted to a small area at both sides of the Sino-Vietnamese border (Xiajinchang Town, Malipo County, Yunnan Province, China. Khau La Village, Quyet Tien Community, Quan Ba District, Ha Qiang province, Vietnam.) (Figure 4). It grows on moist and shady limestone rocks, at ca. 1000–1500 m altitude.

Figure 4. 

Geographical distribution of Primulina malipoensis (triangle), P. lungzhouensis (cross), P. maguanensis (dot), P. maculata (pentagon), P. beiliuensis var. beiliuensis (square) and P. beiliuensis var. fimbribracteata (star).

Conservation status

Based on the field investigations, Primulina malipoensis is currently only known from three sites around the Sino-Vietnamese boundary. Each population possesses no more than 150 mature individuals. However, the type population, which grew close to a road, had disappeared in 2017 and thus, the primary reason why it disappeared is probably due to its destruction by human activities. Based on currently available information, P. malipoensis should be considered as Endangered (EN): B1b(iii,v)c(iv)+2b(iii,v)c(iv); C2b, following the IUCN Categories and Criteria (IUCN 2016).


This new species was observed flowering from June to July and fruiting from August to September.


The specific epithet is derived from the place, Malipo County in Yunnan province, China, where the new species was first found.


Primulina malipoensis (Figures 1 and 2) can be morphologically connected to P. maguanensis (Z. Yu Li, H. Jiang & H. Xu) Mich. Möller & A. Weber (Figure 3A–B) and P. lungzhouensis (W.T. Wang) Mich. Möller & A. Weber (Figure 3D–E) by its ovate or broadly elliptic leaf blade, with inconspicuously (or conspicuously) serrate margin, obvious bracts, white calyx lobes and infundibuliform corolla tube. However, it can easily be distinguished from the latter two species by the characters summarised in the diagnosis.

The authors’ molecular phylogenetic analyses illustrate that P. malipoensis, P. lungzhouensis, P. beiliuensis B. Pan & S.X. Huang (Pan et al. 2013) and P. beiliuensis B. Pan & S.X. Huang var. fimbribracteata. F. Wen & B.D. Lai (Lai and Wen 2015) form a monophyletic clade (Kong et al. 2017). However, their morphology and geographical distribution allow the assumption that P. maguanensis and P. maculata W.B. Xu & J. Guo (Guo et al. 2015) are also closely related to this group. Both P. maguanensis and P. maculata were compared to P. eburnea in the original protologue (Xu et al. 2008, Guo et al. 2015). Nevertheless, based on the observation of living plants, P. maguanensis seems most similar to P. lungzhouensis and P. malipoensis; P. maculata (Figure 3I–J) seems most similar to P. beiliuensis var. beiliuensis (Figure 3G–H) and P. beiliuensis var. fimbribracteata (Figure 3C–F). Further, the geographical distribution of P. maguanensis is adjacent to P. lungzhouensis and P. malipoensis (Figure 4) and the geographical distribution of P. maculata is adjacent to P. beiliuensis (Figure 4). Moreover, the results of the phylogenetical analysis in Guo et al. (2015) show that P. maculata is more closely related to P. lungzhouensis than P. eburnea. All of the above five species occur in nearly the same latitude zone of karst limestone areas from Southern China (from S-Yunnan to S-Guangdong), but with a disjunctive distribution (Figure 4). Therefore, these species perhaps represents a complex of longitudinal speciation, which may be caused by geographical isolation. Further studies are needed to confirm the phylogenetic relationship of this species complex and to determine its evolutionary mechanism of speciation.

Primulina malipoensis could also be related to other species by its yellow flowers. However, the phylogenetic results illustrate that P. malipoensis has a distant relationship with all yellow flowering species, such as P. lutea (Yan Liu & Y. G. Wei) Mich. Möller & A. Weber, P. alutacea F. Wen, B. Pan & B.M. Wang (Pan & al. 2016), P. pteropoda (W.T. Wang) Yan Liu, P. leprosa (Yan Liu & W.B. Xu) W.B. Xu & K.F. Chung and P. jiangyongensis X.L. Yu & Ming Li (Li et al. 2014) (cf. Kong et al. 2017). These yellow flowering species are distributed across different clades (Kong et al. 2017), which means that yellow flowers have independently evolved in different species. This result also suggests that flower colour can be used as an important character to differentiate species in Primulina.

Other specimen examined

CHINA. Yunnan Province, Malipo county, Xiajinchang town, Aotang, 23°07'45.41"N, 104°51'29.25"E, Alt. 1400 m, growing on moist limestone rocks near a road, 8 January 2010, Southeast Yunnan investigation team of DNA barcoding, GBOWS189 (KUN!). CHINA. Guangdong Province, Guangzhou City, voucher from a cultivated plant at South China Botanical Garden, 12 June 2016 (flowering), Li-Hua Yang, YLH350 (IBSC!), introduced from same locality and by the same people as the type. VIETNAM. Ha Qiang province, Quan Ba District, Quyet Tien Community, Khau La Village, Alt. 1100 m, growing on moist limestone rocks, 17 October 2017, Fang Wen et al. VMN-CN 874 (IBK!, VMN!).


This work was supported by grants from the NSFC- Guangdong Natural Science Foundation Joint Project (U1501211), Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01), the Key Research and Development Project of Guangxi (Guike AB16380053). We thank Yun-Xiao Liu for the illustration and Dr. Annemarie Heiduk for the modification of English.


  • Gao Y, Ai B, Kong HH, Kang M, Huang HW (2015) Geographical pattern of isolation and diversification in karst habitat islands: a case study in the Primulina eburnea complex. Journal of Biogeography 42: 2131–2144.
  • Guo J, Pan B, Liu J, Xu WB, Chung KF (2015) Three new species of Primulina (Gesneriaceae) from limestone karsts of China based on morphological and molecular evidence. Botanical Studies 56: 34.
  • Hao Z, Kuang YW, Kang M (2015) Untangling the influence of phylogeny, soil and climate on leaf element concentrations in a biodiversity hotspot. Functional Ecology 29: 165–176.
  • IUCN (2016) Guidelines for using the IUCN Red List categories and criteria. Version 12. Prepared by the Standards and Petitions Subcommittee.
  • Kong HH, Condamine FL, Harris AJ, Chen JL, Pan B, Möller M, Hoang VS, Kang M (2017) Both temperature fluctuations and East Asian monsoons have driven plant diversification in the karst ecosystems from southern China. Molecular Ecology 26: 6414–6429.
  • Lai BD, Wen F (2015) Primulina beiliuensis var. fimbribracteata (Gesneriaceae), a new variety in a limestone cave from Northern Guangdong, China. Taiwania 60: 43‒48.
  • Li ZY, Wang YZ (2004) Plants of Gesneriaceae in China. Henan Science and Technology Publishing House, Zhengzhou, 1–721.
  • Pan B, Wang BM, He JY, Wen F (2016) Primulina versicolor and P. alutacea spp. Nov. (Gesneriaceae), two new species with yellow flowers from northern Guangdong, China. Edinburgh Journal of Botany 73: 25–37.
  • Wang WT, Pan KY, Li ZY, Weitzman AL, Skog LE (1998) Gesneriaceae. In: Wu ZY, Raven PH (Eds) Flora of China.Vol.18. Science Press, Beijing, & Missouri Botanical Garden Press, St. Louis, 244–401.
  • Wang YZ, Mao RB, Liu Y, Li JM, Dong Y, Li ZY, Smith JF (2011) Phylogenetic reconstruction of Chirita and allies (Gesneriaceae) with taxonomic treatments. Journal of Systematics and evolution 49: 50–64.
  • Wang J, Ai B, Kong H, Kang M (2017) Speciation history of a species complex of Primulina eburnea (Gesneriaceae) from limestone karsts of southern China, a biodiversity hot spot. Evolutionary Applications 10: 919–934.
  • Wei YG, Wen F, Möller M, Monro A, Zhang Q, Gao Q, Mou HF, Zhong SH, Cui C (2010) Gesneriaceae of South China. Guangxi Science and Technology Publishing House, Guilin, 1–777.
  • Weber A, Middleton DJ, Forrest A, Kiew R, Lim CL, Rafidah AR, Sontag S, Triboun P, Wei YG, Yao TL, Möller M (2011) Molecular systematics and remodeling of Chirita and associated genera (Gesneriaceae). Taxon 60: 767–790.
  • Weber A, Clark JL, Möller M (2013) A new formal classification of Gesneriaceae. Selbyana 31: 68–94.
  • Xu WB, Zhang Q, Wen F, Liao WB, Pan B, Chang H, Chung KF (2012) Nine new combinations and one new name of Primulina (Gesneriaceae) from South China. Phytotaxa 64: 1–8.
  • Yang LH, Ning ZL, Kang M (2017) Primulina rubella sp. nov. (Gesneriaceae) from a limestone area in Guangdong, China. Nordic Journal of Botany 35: 506–512. https://dx.doi: 10.1111/njb.01374
login to comment