Bangladesh J. Plant Taxon. 24(2): 173–182, 2017 (December) © 2017 Bangladesh Association of Plant Taxonomists PRIMULINA WENII (GESNERIACEAE), A NEW SPECIES FROM CHINA JIAN LI, XIN CHEN, SHU LI1 AND LI-JIAO YAN2 Institute of Ecology, College of Life Sciences, Zhejiang University, Hangzhou, Zhejiang province, China, CN-315008, P.R. China Keywords: East of China; Flora of Fujian; Gesneriaceae; New taxon; Primulina juliae Abstract Primulina wenii (Gesneriaceae) is described and illustrated here as a new species. Morphologically, this new species resembles P. juliae (Hance) Mich. Möller & A. Weber, but it is distinct from the latter by some obvious features in leaf blade, bract, indumentum, corolla, filament, stigma and pistil. Nuclear ribosomal internal transcribed spacer (ITS) region and plastid trnL-F intronspacer (trnL-F) DNA sequence data from the new species and its 22 relatives are used to determine the systematic position of the new species within Primulina. Molecular evidence suggests that P. wenii is strongly supported as a sister to a clade in which P. juliae is included. The conservation status of P. wenii is assessed as "Critically Endangered" (CR) according to IUCN Red List categories and criteria. Introduction A great number of new species of Primulina have been described since 2011 at present the number has rapidly reached more than 150 species in China (Möller et al., 2016). Undoubtedly, the limestone areas of South-west and South China and North Vietnam are considered to be of the highest biodiversity and differentiation for Primulina (Gesneriaceae) (Wei et al., 2010; Wen et al., 2015). It is easily understood that very few species of Primulina in Fujian province have been recorded. Before 1992, there were two species and one variety, namely P. pinnatifida (Hand.- Mazz.) Y.Z. Wang [former Chirita pinnatifida (Hand.-Mazz.) Burtt], Primulina fimbrisepala (Hand.-Mazz.) Y.Z. Wang (former Chirita fimbrisepala Hand.-Mazz.) and Chirita gueilinensis W.T. Wang var. brachycarpa W.T. Wang (Zhang, 1993). However, Chirita gueilinensis var. brachycarpa was brought into Chirita juliae Hance (Wang et al., 1998; Li and Wang, 2004) based on the similar morphological characters of the two congeners. But soon afterwards, Chirita along with several other associated genera, namely almost all the species and Chirita sect. Gibbosaccus C.B. Clarke, two species of Wentsaiboea D. Fang & D.H. Qin (W. renifolia D. Fang & D.H. Qin (2004) and W. luochengensis Yan Liu & W.B. Xu (2010)) and all the species of Chiritopsis W.T. Wang (1981) were shifted to Primulina Hance (Li and Wang, 2007; Wang et al., 2011; Weber et al., 2011). It was originally the monotypic genus (Hance, 1883). Subsequently, a new species of Primulina, viz., P. xiuningensis (X.L. Liu & X.H. Guo) Mich. Möller & A. Weber (former Chiritopsis xiuningensis X.L. Liu & X.H. Guo) (Liu and Guo, 1989; Weber et al., 2011), from Fujian, China was discovered and reported formally in recent years (Geng et al., 2014). To sum it up, Fujian province in East China cannot still be considered to have high biodiversity of Primulina because only three species were found in this area. 1Gesneriaceae Conservation Center of China (GCCC), Guilin Botanical Garden, Guangxi Inst. of Botany, Guangxi Zhuang Autonomous Region and Chinese Academy of Sciences, CN-541006 Guilin, P.R. China. 2Corresponding author.Email: yanljzju@163.com mailto:yanljzju@163.com 174 LI et al. In 2012, one of the authors found a population of the peculiar Gesneriaceae plants growing on a rocky slope near a stream in a limestone gorge close to Fuzhou city, Fujian province, China. Firstly, we considered it to be Prmulina juliae because it is recorded in Ninghua County and Songxi County of Fujian. The next year we re-visited the same location to collect specimens with flowers, and we found that the two similar congeners are actually different. We sent some specimens with flowers and fruits to IBK (Dr. Fang Wen, FW) for identification. He thought they should be an unknown species and have never been seen before. After consulting relevant literature (Wang, 1985; Wang et al., 1990, 1998; Li and Wang, 2004; Wei et al., 2010; Wang et al., 2011; Weber et al., 2011; Möller et al., 2011, 2016;Xu et al., 2012; Möller and Clark, 2013) and checking herbarium specimens of Primulina (Some former Chirita), ‘World checklist of Gesneriaceae’ (Skog and Boggan, 2007) and ‘The genera of Gesneriaceae’ (Weber and Skog, 2007), and consulting Dr. Fang Wen based on his rich experience and knowledge about the identification and taxonomy of Primulina, we found the morphology of these newly discovered plants to be similar to P. juliae (Hance) Mich. Mich. & A. Weber (Wei et al., 2010). However, they differ significantly in some characters. We have determined that they belong to a new species of Primulina viz. P. wenii, sp. nov.The new species Primulina wenii is described and illustrated below and compared with the morphologically similar P. juliae and their conservation status is evaluated. Material and Methods An overview of the genus Primulina from South, South-west and East China was made. All the available specimens of Primulina stored in the following herbaria in China (ANU, BJFC, CDBI, CTC, HGAS, HIB, IBK, IBSC, KUN, PE, SZ) were used as material of Primulina from recent fieldwork by the re-searching team of authors in South, South-west and East China (Thiers, 2015). All the morphological characters were studied under dissecting microscopes, and described using the terminology presented by Wang et al. (1990, 1998). Leaf material of the new species was collected in the field and dried by silica gel for DNA extraction (Chase and Hills, 1991). The nuclear ribosomal internal transcribed spacer (ITS) region and the plastid trnL-F intron spacer (trnL-F) were used as molecular markers. The molecular methods and protocols followed Möller et al. (2009, 2011). GenBank accession numbers for ITS and trnL-F of the new species are KX985576 and KX985577, respectively (Table 1). To elucidate the phylogenetic affinities of the new species, ingroups (22 species of Primulina) and outgroups [Ornithoboea wildeana Craib, Paraboea rufescens (Franch.) B.L.Burtt] selection were chosen based on recent phylogenetic analyses (Li et al., 2007; Mölleret al., 2011; Weber et al., 2011; Kang et al., 2014), and from which sequences were available from GenBank (Table 1). Sequence data were edited and assembled using Lasergene Navigator 7.1 (DNAStar, Madison, Wisconsin, USA) and then aligned with the MEGA 5.1 (Tamura et al., 2011) with additional manual refinements where necessary. Phylogenetic analyses were performed using maximum parsimony (MP) method implemented in PAUP* 4.0b10. Heuristic searches were performed using a starting tree built from stepwise in addition with TBR branch swapping and 1,000 random addition replicates. To assess confidence in clades, bootstrap analyses based on 1,000 replicates with 10 random additions per replicate were used. PRIMULINA WENII (GESNERIACEAE), A NEW SPECIES 175 Table 1. List of species along with accession number of trnL-F and ITS sequences used in this study. Species name Voucher number trnL-F ITS Ornithoboea wildeana Craib Jia-Mei Li Ljm-04-44 DQ872824 DQ865197 Paraboea rufescens( Franch.) Burtt. Jia-Mei Li 0185 DQ872825 DQ865196 Primulina bipinnatifida (W.T. Wang) Y.Z. Wang Guangxi NA/Li J.M.NA DQ872806 DQ872842 Primulina danxiaensis DXS04 KF498157 KF498050 Primulina dryas (Dunn) Mich. Möller & A. Weber T.C. Godfrey 369 FJ501524 FJ501348 Primulina glandulosa (D. Fang, L. Zeng & D.H. Qin) Y.Z. Wang Jia-Mei Li 054291 DQ872804 DQ872841 Primulina glandulosa var. yangshuoensis (Fang Wen, Yue Wang & Q.X. Zhang) Mich. Möller & A. Weber M. Möller MMO 06-912 HQ632948 HQ633045 Primulina heterotricha (Merr.) Y.Z. Wang Yin-Zheng Wang 067311 DQ872816 DQ872826 Primulina juliae JXGF01 KF498228 KF498107 Primulina linearifolia (W.T. Wang) Y.Z. Wang Jia-Mei Li 11121 DQ872810 DQ872834 Primulina longgangensis (W.T. Wang) Y.Z. Wang A. Takhtajan & N. Aruzytov 1975 AJ492290 FJ501347 Primulina minutimaculata (D. Fang & W.T. Wang) Y.Z. Wang Jia-Mei Li 067134 DQ872815 DQ872828 Primulina mollifolia (D. Fang & W.T. Wang) Y.Z. Wang Jia-Mei Li 054281 DQ872802 DQ872847 Primulina ophiopogoides (D. Fang & W.T. Wang) Y.Z. Wang Yin-Zheng Wang 067134 DQ872814 DQ872829 Primulina pinnata (W.T. Wang) Y.Z. Wang Expedition Beijing 896526 FJ501526 FJ501349 Primulinapinnatifida (Hand.-Mazz) Y.Z. Wang Q.J. Xie J-037 FJ501527 FJ501350 Primulina pteropoda (W.T. Wang) Y.Z. Wang Yin-Zheng Wang 067312 DQ872817 DQ872827 Primulina repanda var. guilinensis ex Smithsonian Institute 94-083 AJ492292 FJ501351 Primulina spadiciformis (W.T. Wang) Mich. Möller & A. Weber ex Smithsonian Institute 94-087 AJ492291 FJ501346 Primulina spinulosa (D. Fang & W.T. Wang) Y.Z. Wang Yin-Zheng Wang 067133 DQ872813 DQ872830 Primulina tabacum Hance Q.J. Xie & C.X. Ye AJ492300 FJ501352 Primulina weii Mich. Möller & A. Weber Jia-Mei Li, Ljm-04-42 DQ872811 DQ872832 Primulina wenii Jian Li & L.J. Yan sp. nov. Jian Li & F. Wen 20130412-01 KX985577 KX985576 Primulina wentsaii (D. Fang& L. Zeng) Y.Z. Wang Jia-Mei Li 11630 DQ872812 DQ872831 Primulina xiuningensis (X.L. Liu & X.H. Guo) Mich. Möller & A. Weber ZJJS01 KF498252 KF498124 176 LI et al. Results and Discussion Taxonomical treatment Primulina wenii Jian Li & L. J. Yan, sp. nov. (Figs 1 & 2). Diagnosis: Primulina wenii Jian Li & L. J. Yan differs from its congener, P. juliae (Hance) Mich. Möller & A. Weber, by the combination of the following characters: the lateral veins of leaf blade 3 or 4; peduncle densely covered with erectly spreading white eglandular pubescent and villous hairs; bracts 3, outside pubescent and villous, inside pubescent; corolla outside densely white glandular-pubescent, inside sparsely pubescent; filaments glabrous; stigma trapeziform, 2- lobed to irregularly lobed (Table 2) Type: CHINA. Fujian province, Fuzhou city, Rixi town, 26.35139°N, 119.27056°E, 130 m, on moist tufa and the rocky surface of limestone cliff, 12 April 2013, Jian Li & F. Wen 20130412- 01 (Holotype: IBK!; Isotype: IBK!). Table 2. Morphological comparison of Primulina wenii sp. nov. and P. juliae. Characters P. wenii P. juliae Leaf blade indumentum Lower surface densely appressed pubescent and veins villous, upper surface densely pubescent and villous Both surfaces appressedpuberulent Leaf blade margin Irregularly serrated in number and size Dentate to pinnately lobed basally or crenate, apex acute to rounded Lateral veins of leaf blade 3 or 4, abaxially obviously prominent, adaxially conspicuously sunken 4 or 5, abaxially inconspicuously prominent, adaxially inconspicuous Peduncle indumentum Densely covered with erectly spreading white eglandular pubescent and villous hairs Densely covered with spreading puberulent hairs Bract number and indumentum 3; outside pubescent and villous, inside pubescent 2; outside puberulent, inside glabrous Corolla indumentum Outside densely white glandular- pubescent, inside sparsely pubescent Outside sparsely puberulent, inside nearly glabrous Filaments Glabrous Glabrous to glandular puberulent near apex Pistil Densely covered with white glandular puberulent hairs Puberulent Stigma Trapeziform, 2-lobed to irregularly lobed Narrowly oblong, 2-lobed Herbs, perennial. Rhizome cylindric, 2–4 cm long, 0.6–1.0 cm in diam. Leaves 4–6, basal; petiole compressed, 1.9–2.5 × c. 0.8 cm, densely villous; leaf blade pale green to green, oblong to broadly elliptic, 10–20 × 7–14 cm, herbaceous but chartaceous when dried, lower surface densely appressed pubescent and along veins villous, upper surface densely pubescent and villous, base attenuate to cuneate, margin irregularly serrate, ciliate, apex obtuse; lateral veins 3 or 4 on each side of midrib, abaxially obviously prominent, adaxially conspicuously sunken. Cymes 3–4, axillary, 3(5)–7-flowered or more; peduncle green, 8–10 cm long, 2.5–3.0 mm in diam., densely covered with erectly spreading white eglandular pubescent and villous hairs; bracts 3, lateral ones PRIMULINA WENII (GESNERIACEAE), A NEW SPECIES 177 Fig. 1. Primulina wenii sp. nov. A. Habitat; B. Habit in flowering; C. Leaf blade – adaxial surface; D. Leaf blade – abaxial surface; E. Cyme, buds and bracts; F. bracts; G. Cymes and flowers; H. Peduncle and its indumentum; I. Front view of corolla; J. Upward view of corolla; K. Lateral view of corolla; L. Stigma with dissected lobes; M. Curved immature capsules. A and M: collected from type locality, 22 Jul 2013; B–L: photographed in nursery of Gesneriad Conservation Center of China, Guilin, 12 Apr 2014, by F. Wen. 178 LI et al. Fig. 2. Primulina juliae. A. Habit; B. Leaf blade – adaxial surface; C. Front view of corolla; D. Lateral view of corolla; E. Cyme, calyx lobes and indumentum; F. Young straight capsules. A– B and E– F: photographed in type location by J. Li; C– D: photographed in nursery of Gesneriad Conservation Center of China by F. Wen. PRIMULINA WENII (GESNERIACEAE), A NEW SPECIES 179 conspicuously opposite, green, linear to lanceolate, 14–16 × 2.5–3.0 mm, the central one smaller, 10–12 × 1.3–1.6 mm, persistent at flowering; outside white pubescent and villous, inside white pubescent, margin entire and ciliate, apex acute. Calyx 5-partite to base; lobes lanceolate, 14–15 × c. 2.5 mm, nearly equal, outside densely erectly white villous and pubescent, inside sparsely shortly pubescent to nearly glabrous, margin entire, apex acute. Corolla outside pale bluish purple, throat with 2 purple stripes, upper portion of inside corolla surface with 2 dark purplish brown swollen spots, 2 swollen spots sparsely covered with short glandular hairs, inside of lobes pale purplish blue with slightly darker purple longitudinal stripes, corolla c. 3.2 cm long, outside densely white glandular-pubescent, inside sparsely pubescent; tube nearly tubular, c. 2.8 cm long, Fig. 3. The strict consensus tree of Primulina taxa based on a maximum parsimony (MP) analysis of combined ITS and trnL-F. Numbers of the branches indicate bootstrap values > 50% by MP analysis. * indicates the new species. orifice c. 1.7 cm in diam.; limb distinctly 2-lipped; upper lip 2-lobed, lobes triangular to semicircle, 1.0–1.2 cm long; lower lip 3-lobed, lobes oblong to triangle, 1.0–1.3 cm long. Stamens 2, adnate to c. 1.2 cm above corolla base; filaments white, c. 1 cm long, geniculate c. 3 mm above insertion, glabrous; anthers reniform, slightly constricted at middle, c. 3.5 mm long; staminodes 3, lateral ones adnate to c. 1 cm above corolla base, white, linear, c. 7.5 mm long, glabrous, apex small capitate, central staminode adnate to c. 0.5 cm above corolla base, punctiform, inconspicuous, c. 0.5 mm long. Disk annular, c. 10 mm high; pistil c. 2.3 cm long, densely 180 LI et al. covered with white eglandular hairs; style linear, c. 1.3 cm, pubescent; stigma trapeziform, 2-lobed to irregularly lobed, c. 2.5 mm long. Fruit c. 5 cm long, straight to slightly curved, hairy. Flowering and fruiting period: April – June. Etymology: The species epithet ‘wenii’ is used to commemorate a Chinese botanist, Fang Wen, who studied Gesneriaceae of China for many years, and is devoted to the conservation and taxonomy of Chinese Gesneriaceae plants. Vernacular name: Chinese: Wēn Shì Bào Chūn JǜTái. Habitat: Primulina wenii grows on shaded moist turf of limestone cliff in subtropical limestone evergreen broad-leaf forest on a North-facing slope of a limestone hill at an altitude of about 130 meters. The climate of Fuzhou, Fujian province is the monsoon of subtropical moist marine climate zone. The average annual temperature of Fuzhou is 20.1°C, the average annual precipitation is c. 1718.1 mm. Notes: East China is not the area with high biodiversity of Gesneriaceae in China. For example, only six new species of Gesneriaceae, namely Primulina xiziae Fang Wen, Yue Wang & G. J. Hua (Li et al., 2012), P. chizhouensis Xin Hong, S.B. Zhou & F.Wen (Hong et al., 2012), P. suichuanensis X.L. Yu & J.J. Zhou (Zhou et al., 2016), Didymocarpus dissectus Fang Wen, Y. L. Qiu, Jie Huang & Y.G. Wei (Wen et al., 2013), Oreocharis striata Fang Wen & C.Z. Yang (Yang et al., 2015) and Beccarinda baolianis Q.W. Lin (Lin, 2016) were discovered and described in the past decade. Further, Fujian province, belongs to E China, only possesses three species of Primulina before this new one was discovered. This also implies that more field investigations will help more discoveries. Molecular analysis The combined matrix had a length of 1,670 characters, 672 for ITS and 998 for trnL-F. Of the 254 (15.2%) varied, 257 (15.4%) were parsimoniously informative, including the indels. The maximum parsimony analysis on the combined matrix resulted in two trees of 937 steps in length, a consistency index (CI) of 0.7311, retention index (RI) of 0.6644 and homoplasy index (HI) of 0.2689. The strict consensus tree (Fig. 3) was highly resolved and tree topology was consistent with the previous phylogenetic study by Möller et al. (2011) and Kang et al. (2014). DNA analyses of sequence data show that the new species is sister to a clade (BS = 100) comprising of Primulina tabacum, P. danxiaensis, P. juliae and P. xiuningensis. Of these species the undescribed species is morphologically most similar to P. juliae from which it can be distinguished by certain qualitative and quantitative characters in leaf blade, bract, indumentum, corolla, filament, stigma and pistil. For instance, it is obviously different that the peduncles of P. wenii are densely covered with erectly spreading white eglandular pubescent and villous hairs, the number of bracts is three, filaments are glabrous, and pistil is densely covered with white glandular puberulent hairs. The morphological differences between the two species (P. wenii and P. juliae) are depicted in Table 2. Acknowledgements The authors would like to thank Dr. Fang Wen and Prof. Yi-Gang Wei for identifying specimens and cultivating plants. We also thank some anonymous reviewers for their comments on an earlier draft of this paper. This study was supported by the Guangxi Natural Science Foundation (2015GXNSFBB139004) and the Key Research and Development Project of Guangxi (Guike AB16380053). PRIMULINA WENII (GESNERIACEAE), A NEW SPECIES 181 References Chase, M.W. and Hills, H. 1991. Silica gel: an ideal material for field preservation of leaf samples for DNA studies. Taxon 40: 215–220. Fang, D. and Qin, D.H. 2004. Wentsaiboea D. Fang. & D.H. Qin, a new genus of the Gesneriaceae from Guangxi, China.ActaPhytotax. Sin. 42: 533–536. Geng, H.Q., Ma, Q.X., Yan, Y.H., Li, Y.F., Jiang, Q.Y., Lǘ, L.L., Wu, W.H., Ding, X. and Hou, X.L. 2014. New materials of plants in Fujian province (II).Subtrop. Plant Sci. 43: 24–28. Hance, H.F. 1883. New Chinese Cyrtandreae. J. Bot. 21: 165–170. Hong, X., Zhou, S.B. and Wen, F. 2012. Primulina chizhouensis sp. nov. (Gesneriaceae), a new species from a limestone cave in Anhui, China. Phytotaxa 50: 13–18. IUCN 2012. IUCN Red List categories and criteria. Version 3.1, ed. 2. Gland & Cambridge, IUCN. Kang, M., Tao, J., Wang, J., Ren, C., Qi, Q., Xiang, Q.Y. and Huang, H. W. 2014. Adaptive and non- adaptive genome size evolution in Karst endemic flora of China. New Phytol. 202: 1371–1381. Li, J., Wang, Y., Hua, G.J. and Wen, F. 2012. Primulina xiziae sp. nov. (Gesneriaceae) from Zhejiang Province, China. Nordic J. Bot.30: 77–81. Li, J.M. and Wang, Y.Z. 2007. Phylogenetic reconstruction among species of Chiritopsisand Chirita Sect. Gibbosaccus (Gesneriaceae) based on nrDNA ITS and cpDNAtrn L-F sequences. Syst. Bot.32: 888–898. Li, Z.Y. and Wang, Y.Z. 2004. Primulina, Chirita and Chiritopsis. In: Li, Z.Y. and Wang, Y.Z. (eds), Plants of Gesneriaceae in China. Henan Sci. & Technol. Publ. House, Zhengzhou, pp. 170–282. Lin, Q.W. 2016.Beccarinda baolianis, a new species of Gesneriaceae from Fujian Province. Bull. Bot. Res. 36: 650–652. Liu, X.L. and Guo, X.H. 1989. A new species of Chiritopsis from Anhui. Bull. Bot. Res. 9: 51–54. Liu, Y., Xu, W.B. and Pan, B. 2010. Wentsaiboea tiandengensis sp. nov.and W.luochengensis sp. nov. (Gesneriaceae) from Karst caves in Guangxi, southern China. Nordic J. Bot 28: 739–745. Möller, M., Pfosser, M., Jang, C.G., Mayer, V., Clark, A., Hollingsworth, M.L., Barfuss, M.H.J., Wang, Y.Z., Kiehn, M.and Weber, A. 2009. A preliminary phylogeny of the ‘Didymocarpoid Gesneriaceae’ based on three molecular data sets: incongruence with available tribal classifications. Am. J. Bot. 96: 989–1010. Möller, M. and Clark, J.L. 2013. The state of molecular studies in the family Gesneriaceae: A review. Selbyana 31: 95–125. Möller, M., Forrest, A., Wei, Y.G. and Weber, A. 2011. A molecular phylogenetic assessment of the advanced Asiatic and Malesian didymocarpoid Gesneriaceae with focus on non-monophyletic and monotypic genera. Plant Syst. Evol. 292: 223–248. Möller, M., Wei, Y.G., Wen, F., Clark, J.L. and Weber, A. 2016. You win some you lose some: updated generic delineations and classification of Gesneriaceae-implications for the family in China. Guihaia36: 44–60. Skog, L. E. and Boggan, J.K. 2007. World checklist of Gesneriaceae. Department of Botany, Smithsonian Institution, Washington, DC. http://botany.si.edu/Gesneriaceae/Checklist [accessed 28 April 2010]. Tamura, K., Peterson, D., Peterson, N., Stecher, G., Nei, M. and Kumar, S. 2011. MEGA5: molecular evolutionary genetics analysis using maximum likelihood, evolutionary distance, and maximum parsimony methods. Mol. Biol. Evol. 28: 2731–2739. The International Plant Names Index. 2015+. [continuously updated]: Primulina. Published at http://www.ipni.org/ [accessed 12 Feb 2015] Thiers, B. 2015. Index Herbariorum: a global directory of public herbaria and associated staff. New York Botanical Garden: published at http://sweetgum.nybg.org/ih/ [accessed 3 Feb 2015]. Wang, W.T. 1981. Quinque genera nova gesneriacearum E Sina. Bull. Bot. Res. 1: 21–51. Wang, W.T. 1985. A revision of the genus Chirita (Gesneriaceae) in China (II). Bull. Bot. Res., Harbin 5: 37–86. http://botany.si.edu/Gesneriaceae/Checklist http://www.ipni.org/ http://sweetgum.nybg.org/ih/ 182 LI et al. Wang, W.T., Pan, K.Y. and Li, Z.Y. 1990. Gesneriaceae. In: Wang, W.T. (Ed.), Flora Reipublicae Popularis Sinicae69. Science Press, Beijing, pp. 125–581. Wang W.T., Pan, K.Y. and Li, Z.Y. 1998. Gesneriaceae. In: Wu, Z.Y. and Raven, P.H. (eds), Flora of China Vol. 18. Science Press, Beijing & Missouri Bot. Garden Press, St. Louis, 322 pp. Wang, Y.Z., Mao, R.B., Liu, Y., LI, J.M., Dong, Y., LI, Z.Y. and Smith, J. F. 2011. Phylogenetic reconstruction of Chiritaand allies (Gesneriaceae) with taxonomic treatments. J. Syst. Evol. 49: 50–64. Weber, A., Middleton, D.J., Forrest, A., Kiew, R., Lim, C.L., Rafidah, A.R., Sontag, S., Triboun, P., Wei, Y.G., .Yao, T.L. and Möller, M. 2011. Molecular systematics and remodelling of Chirita and associated genera (Gesneriaceae). Taxon 60: 767–790. Weber, A. and Skog, L.E. 2007. The genera of Gesneriaceae. Basic information with illustration of selected species. Ed. 2. Available from http://www.genera-gesneriaceae.at/. Wei, Y.G., Wen, F., Möller, M., Monro, A., Zhang, Q., Gao, Q., Mou, H.F., Zhong, S.H. and Cui, C. 2010. Gesneriaceae of South China. Guangxi Science and Technology Publishing House, Nanning, 777 pp. Wen, F., Qiu, Y.L., Huang, J., Zhao, B. and Wei, Y.G. 2013. Didymocarpus dissectu ssp. nov. (Gesneriaceae) from Fujian, eastern China. Nordic J. Bot. 31: 316–320. Wen, F., Lai, B.D., Zhao, Z.G., He, J.Y. and Jiang, B.S. 2015. Primulina heterochroa (Gesneriaceae), a new species from a tropical limestone area in Guangxi, China. Willdenowia 45: 45–51. Yang, C.Z., Cai, D.L. and Wen, F. 2015. Oreocharis striata (Gesneriaceae), a new species from Fujan, China. Ann. Bot. Fennici 52: 369–372. Xu, W.B., Zhang, Q., Wen, F., Liao, W.B., Pan, B., Chang, H. and Chung, K.F. 2012. Nine new combinations and one new name of Primulina. Phytotaxa 64: 1–8. Zhang, Y.T. 1993. Gesneriaceae. In: Lin, L.G. (ed.), Flora of Fujian 3. Fuzhou: Fujian Science and Technology Publishing House. Zhou, D.S., Zhou, J.J., Li, M. and Yu, X.L. 2016: Primulina suichuanensis sp. nov. (Gesneriaceae) from Danxia landform in Jiangxi, China. Nordic J. Bot. 34: 148–151. (Manuscript received on 14 October 2016; revised on 28 August 2017) http://www.genera-gesneriaceae.at/.