Research Article
Research Article
Hemiboea kaiyangensis, a new species of Gesneriaceae endemic to Guizhou, China
expand article infoTao Peng, Shun-Zhi He§, Shun-Li Wang, Dan Huang, Xu-Ping Zhou
‡ Guizhou Normal University, Guiyang, China
§ Guizhou University of Traditional Chinese Medicine, Guiyang, China
Open Access


A new species of Gesneriaceae from Guizhou, China, Hemiboea kaiyangensis sp. nov., is described and illustrated. We investigated its phylogenetic position and relationships with 13 other species of Hemiboea C.B.Clarke, which present large morphological diversity in the genus, based on molecular analyses of the nuclear ribosomal internal transcribed spacer (ITS) and the chloroplast trnL-F intron-spacer sequences. The molecular phylogenetic analyses revealed that the new species is most closely related to H. ovalifolia. A diagnostic table and discussion of morphological characters are provided to differentiate the new species from H. longisepala, H. flaccida and H. ovalifolia.


Gesneriad, ITS, morphology, phylogeny, taxonomy, trnL-F


The genus Hemiboea C.B.Clarke has traditionally been divided into two sections, sect. Subcapitatae C.B.Clarke and sect. Hemiboea (Li 1987a, 1987b). Many species of this genus can be found on limestone hills in tropical and subtropical evergreen broadleaved forests. Most species of Hemiboea are located in southern and southwestern China, while only a few are located outside China. For instance, H. subcapitata C.B.Clarke and H. cavaleriei Lévl. var. paucinervis W.T.Wang & Z.Y.Li ex Z.Y.Li are also found in northern Vietnam; H. bicornuta (Hayata) Ohwi is distributed in Taiwan Island of China and the Ryukyu Islands of Japan (Li and Wang 2005).

So far, a total of 38 species and six varieties have been described in Hemiboea (Li and Liu 2004; Xu et al. 2010; Huang et al. 2011; Wen et al. 2011; Pan et al. 2012; Xu et al. 2012; Wen et al. 2013; Zhou et al. 2013; Zhang et al. 2014; Huang et al. 2017; Chen et al. 2018; Li et al. 2018; Wei 2018; Li et al. 2019; Nguyen et al. 2019; Wu et al. 2019; Huang et al. 2020; Nguyen et al. 2021). The small genus Metabriggsia W.T.Wang was established based on two species, M. ovalifolia W.T.Wang and M. purpureotincta W.T.Wang from Guangxi, China (Wang 1983). Some similarities in corolla morphology with certain species of the former Briggsia Craib are reflected in the generic name (Wang 1983), though Metabriggsia is morphologically similar to Hemiboea, and in recent molecular-based taxonomic studies, the two species of Metabriggsia were sunk into Hemiboea as H. ovalifolia (W.T.Wang) A.Weber & Mich.Möller and H. purpureotincta (W.T.Wang) A.Weber & Mich.Möller (Weber et al. 2011).

During the course of a 2009 floristic study in Guizhou Province, China, we collected specimens of an unidentified Hemiboea from a limestone area in Kaiyang County. It seemed a species of the sect. Subcapitatae of the genus Hemiboea because of the free calyx lobes and elongate corolla. However, it differed from all known species in this genus. We also observed that its vegetative morphology was similar to that of H. ovalifolia because of the stems covered with long white pubescent hairs, and the leaf blades herbaceous and ovate or ovate-oblong. The inflorescence and flower morphology indicated that this species belongs to Hemiboea as previously circumscribed; for example, it possesses involucre, its corolla is infundibular-tubular, and has a ring of hairs on the interior above the corolla base. After consulting national floras and the relevant literature (Li 1983, 1987a, b; Wei and Wen 1995; Weitzman et al. 1997; Wang et al. 1998; Li and Liu 2004; Li and Wang 2005; Wei et al. 2010; Xu et al. 2010), as well as herbarium specimens, we confirmed that it is an undescribed species. We described and illustrated the new species herein. We used molecular data of the nuclear ribosomal internal transcribed spacers (ITS) and the chloroplast trnL-F intron-spacer (trnL-F) to confirm the placement of the newly collected material in Hemiboea and to infer its phylogenetic relationships with other species in the genus.


Taxon sampling

To investigate the phylogenetic placement of the newly collected taxon, our sampling focused on species of Hemiboea and its closest related genera according to Möller et al. (2011) and Weber et al. (2011). The ITS and trnL-F sequences used in Weber et al. (2011) included 13 species of Hemiboea and four outgroup samples (one species of Ornithoboea, one of Paraboea and two of Glabrella) were downloaded from Genbank (Table 1). Leaf material of the new species was collected from living plants in the type locality and rapidly dried in silica gel.

Table 1.

Species names and accession numbers of ITS and trnL-F sequences used for phylogenetic analysis.

Species Voucher number trnL-F ITS
Outroup samples
Briggsia longipes (Hemsl. ex Oliv.) Craib MMO 01-122 FJ501545 AF055052/AF055053
Briggsia mihieri (Franch.) Craib Y.Z.Wang 11315B FJ501544 FJ501363
Ornithoboea wildeana Craib Middleton & al. 4531 JN934710 JN934752
Paraboea rufescens (Franchet) B.L.Burtt Möller MMO 01-108/3 JN934730 JN934772
Ingroup samples
Hemiboea ovalifolia W.T.Wang (W.T.Wang) A.Weber & Mich.Möller B.M.Nong 06-1 HQ632883 HQ632980
Hemiboea purpureotincta (W.T.Wang) A.Weber & Mich.Möller MMO 06-813 HQ632884 HQ632981
Hemiboea bicornuta (Hayata) Ohwi RBGE cult. 19951207 FJ501534 FJ501356
Hemiboea cavaleriei Lévl. Z.J.Gu G3 FJ501533 FJ501355
Hemiboea fangii Chun ex Z.Yu Li MMO 08-1284 HQ632882 HQ632979
Hemiboea follicularis C.B.Clarke Y.G.Wei G03 HQ632885 HQ632982
Hemiboea gracilis Franchet Y.Z.Wang 11317 FJ501536 *
Hemiboea longgangensis Z.Yu Li Y.G.Wei 07-550 HQ632889 HQ632986
Hemiboea longzhouensis W.T.Wang MMO 07-1127 HQ632888 HQ632985
Hemiboea omeiense W.T. Wang MMO 08-1271 HQ632886 HQ632983
Hemiboea kaiyangensis T. Peng & S.Z.He, sp. nov. Shun-Zhi He, 090819 JN644339 JN644335
Hemiboea rubribracteata Z.Yu Li & Yan Liu MMO 07-1093 HQ632890 HQ632987
Hemiboea subcapitata C.B.Clarke Y.Z.Wang 11306 FJ501535 FJ501357

DNA extraction, PCR and direct sequencing

Molecular methods and protocols followed Möller et al. (2009) and Weber et al. (2011). The Genbank accession numbers for ITS and trnL-F of the new species are JN644339 and JN644335, respectively (Table 1).

Phylogenetic analysis

Sequences of ITS and trnL-F of the new species were aligned with the existing matrices of Weber et al. (2011), and the combined data analyzed by Möller et al. (2009, 2011) and Weber et al. (2011). All characters were unordered and equally weighted. Heuristic searches were implemented as 1000 random taxon-addition sequences, with tree bisection–reconnection (TBR) branch swapping, the MULTRESS and STEEPEST DESCENT option in effect. Branch support was obtained by bootstrapping (Felsenstein, 1985ab) with 1000 random resamplings and TBR on and MULTREES off (Möller et al. 2009).


The combined dataset included 1490 characters and contained 1125 (75.5%) constant sites, 333 (22.3%) variable sites and 154 (10.3%) parsimony-informative sites. The heuristic analysis resulted in three most parsimonious trees with a length of 553 steps, a consistency index (CI) of 0.7776, and a retention index (RI) of 0.6295. The strict consensus tree supports that the new species nests in a strongly supported clade of Hemiboea (BS = 100%), and it is the sister taxon to H. ovalifolia (BS = 99%) (Fig. 3).


The molecular phylogenetic analysis revealed that the new collection Hemiboea kaiyangensis fell into the clade of Hemiboea, and its most closely related species was H. ovalifolia, which is congruent to the morphological evidence. Morphologically, this species is most similar to H. longisepala, H. flaccida, and H. ovalifolia, and it can be easily distinguished by the characters summarized in Table 2.

Table 2.

Diagnostic characters used to differentiate Hemiboea kaiyangensis from most similar taxa.

Taxon Hemiboea kaiyangensis H. longisepala H. flaccida H. ovalifolia
Stem densely pubescent Glabrous densely brown puberulent to villous brown villous
Leaf blade Texture herbaceous Papery slightly fleshy herbaceous;
Shape ovate or ovate-oblong ovate-lanceolate to elliptic-lanceolate elliptic to ovate ovate
Hairs appressed pubescent on adaxially and abaxially surface adaxially sparsely pubescent, abaxially glabrous pubescent to densely on adaxially and abaxially surface appressed pubescent on adaxially and abaxially surface
Lateral veins 7–10 on each side 10–12 on each side 5–8 on each side 5–10 on each side
Cyme Peduncle(cm) 10–18 3–3.6 0.4–1.9 7.5–12.5
Hairs on peduncle densely white long glandular pubescent glabrous sparsely glandular puberulent to pilose brown glandular pubescent
Involucre cordate, early deciduous spheroidal, ca. 1.7 cm in diam. nearly spheroidal, 1–2.5 cm in diam., outside sparsely glandular puberulent nearly spheroidal, early deciduous
Calyx lobes oblong-lanceolate, apex obtuse or slightly obtuse, 12–13 × 2.5–3 mm, outside densely glandular pubescent, glabrous inner, 3 veins on one lobe linear-lanceolate, 19–20× ca. 2.5 mm, outside and margin glabrous linear, 5–9 × 2.5–3 mm, outside and margin sparsely glandular puberulent lanceolate-linear, 9–10 × 1.5–2 mm, pubescent outer, glabrous inner, 3–5 veins on one lobe
Flower Colour outside pale yellowish-green to pale yellowish-white outside white outside white, inside purple spotted white, suffused yellow-green
Hairs outside densely glandular puberulent, inside glabrous outside glabrous outside sparsely glandular puberulent, inside glabrous outside sparsely pubescent near apex, inside glabrous
Size 4.5–5 cm long; tube 3.5–4 cm; adaxial lip 6–7 mm, abaxial lip 6–7 mm ca. 3.4 cm long, tube ca. 2.6 cm; adaxial lip ca. 6 mm; abaxial lip ca. 8 mm 3–3.4 cm long, tube 2.3–2.5 cm; adaxial lip 4–5 mm; abaxial lip 7–9 mm ca. 3.6 cm long, tube ca. 2.7 cm; adaxial lip ca. 2.8 mm; abaxial lip ca. 1 cm
Staminodes 2, ca. 7 mm long 3, central 1 ca 1.5 mm long, lateral 5.5 mm 2, 6–8 mm long 3, central 1 ca. 1.5 mm long, lateral 2, 9–10 mm
Pistil glabrous glabrous sparsely glandular puberulent sparsely puberulent

Taxonomic treatment

Hemiboea kaiyangensis T.Peng & S.Z.He, sp. nov.

Figs 1, 2, 3


China; center of Guizhou Province, Zijiang gorge, Kaiyang County, grows on cliffs under forests along the road; alt. 1000–1020 m. 2009-08-18, Shun-Zhi He 90819 (Holotype: HGCM!, isotype: GNUB!, IBK!) (Figs 1, 2).

Figure 1. 

Hemiboea kaiyangensis T.Peng & S.Z.He A habit, showing flowering branch B opened corolla, showing stamens, staminodes and ring of hairs at base of tube C calyx, pistil and disc D stigma E stamens F infructescence G cross section of ovary, showing parietal placentation. Drawing by S.Q.He and Y.X.Zhu.


Hemiboea kaiyangensis is most similar to H. longisepala Z.Y.Li, H. flaccida Chun ex Z.Y.Li and H. ovalifolia (W.T.Wang) A.Weber & Mich.Möller in the glabrous pistil, but it differs in the cordate involucre bracts that are early deciduous, the corolla densely glandular puberulent outside and glabrous inside, and pale yellowish-green to pale yellowish-white outside.

Figure 2. 

Hemiboea kaiyangensis T.Peng & S.Z.He A habitat B plant with flowering branches C involucrum, early deciduous, before flower opening D pair-flowered cymes E calyx, disc and pistil F opened corolla, showing stamens, staminodes and ring of hairs at base of tube G cross section of capsule, showing two parietal placentation H seeds. Scale bar: 0.5 mm (Based on the holotype Shun-Zhi He 90819).


Perennial herb. Rhizomatous. Stems 25–60 cm long, 5–7 mm in diam., densely pubescent. Petiole 0.5–4.5 cm long, densely pubescent. Leaf blade herbaceous, oblique, iso- to distinctly anisophyllous, ovate or ovate-oblong, 13–26 × 5–8 cm, apex acute, base oblique cuneate, margin nearly entire or unapparent sinuous dentate, appressed pubescent on both sides, lateral veins 7–10 on each side. Cymes 2–3, terminal or subterminal, 6–12 flowers per cyme; peduncle 10–18 cm long, densely pubescent with white long glandular hairs; involucre cordate, apex cuspidate, early deciduous; pedicel 0.7–1.5 cm long, pubescent with white long glandular hairs. Calyx lobes 5; lobes oblong-lanceolate, apex obtuse or slightly obtuse, 12–13 × 2.5–3 mm, outside densely pubescent with glandular hairs of 3–4 mm long, inside glabrous, 3 veins per lobe. Corolla pale yellowish-green to pale yellowish-white outside, small purplish-brown spotted inside, 4.5–5 cm long, densely glandular-puberulent outside, glabrous with a ring of white hairs ca. 4 mm above the corolla base inside; tube 3.5–4 cm long, mouth 1–1.3 cm in diam.; limb distinctly 2-lipped, adaxial lip 2-lobed, lobes obliquely semicircular, apex obtuse to rounded, 6–7 mm long, 8–9 mm in diam. at the base of lobes; abaxial lip tripartite, lobes margin ciliolate, the central broadly ovate to ovate-elliptic, 6–7 mm long, two lateral oblique triangle, 7–8 cm long. Stamens 2, glabrous, adnate to 1.8 cm above the corolla base, filament ca. 1.3 cm long, geniculate at the middle; anthers slightly oblong, dark purple, apex coherent. Staminodes 2, glabrous, adnate to 1.7 cm above the corolla base, ca. 7 mm long. Nectary disc ring-like, 1.1–1.2 mm high, atop with ca. 6 obviously erose crena. Pistil 2–2.8 cm long, ovary 7–9 mm long, glabrous, style 1.4–2 cm long, glabrous, stigma 1, terminal truncate, sightly 2-lobed. Capsule obliquely linear-lanceolate, 2–2.4 cm long, 3–3.3 mm in diam., glabrous, slightly curved.

Figure 3. 

Strict consensus tree of three parsimony trees based on combined ITS and trnL-F data. Numbers above branches are bootstrap values. Underline indicates the new species.

Pollen description

Pollen grains of Hemiboea kaiyangensis are prolate-spheroidal, long or oblate, 3-colporoidate grains. In polar view, the outline is close to triangular-circular. The ectocolpi measures 33.05–33.57 × 12.15–14.23 μm and the endoapertures are laterally fused to form an endocingulum. Exine reticulate, muri smooth. The width of muri is unequal in size. The sizes and shapes of perforations are irregular, and vary in size from 0.14–0.67 × 0.11–0.61 μm.

Distribution and ecology

Known only from a single limestone gorge in Kaiyang county, Guizhou Province, China. Only five populations were found, growing on the mouth of caves in shady and damp forests, close to a road, between 900 and 930 m in elevation.


The name of the new species, kaiyangensis, refers to the type locality, Kaiyang County, Guiyang, Guizhou Province, China.

Conservation status

The populations of Hemiboea kaiyangensis are endemic to Kaiyang county, center of Guizhou Province, China, and the species only known from the type locality at present. The five detected populations grow dispersed in a limestone gorge, and cover only an area of about 1.25 km2 and include a total of 75–120 individuals. However, until further investigation, the species should be designated as “Data Deficient” (DD) according to the IUCN standards (IUCN 2019).


As previously mentioned, Hemiboea kaiyangensis is most morphologically similar to H. longisepala, H. flaccida and H. ovalifolia in their glabrous pistil, but some characters, such as the early deciduous and cordate involucre bracts, the corolla indumentum (outside densely glandular puberulent and inside glabrous), and the corolla color (outside pale yellowish-green to pale yellowish-white), help us to easily distinguish them. Specifically, H. kaiyangensis is similar to H. flaccida in having two staminodes, but it differs in the longer peduncle (10–18 cm), the involucre cordate but early deciduous, the bigger calyx lobes, oblong-lanceolate (12–13 × 2.5–3 mm), and the glabrous pistil. This new species is also similar to H. ovalifolia in the texture and shape of leaf blade, but it can be distinguished in the peduncle indument, with dense white long glandular hairs, the involucre cordate, the calyx lobes outside densely glandular pubescent, the corolla outside densely glandular puberulent, 2 staminodes, and the pistil glabrous. Lastly, H. kaiyangensis is similar to H. longisepala, but it differ from the latter in the stem densely pubescent, the longer peduncle (10–18 cm), the peduncle with dense white long glandular hairs, the corolla outside densely glandular puberulent and 2 staminodes. All compared details of four congeners were listed in Table 2.


We are grateful to Mr. S.Q.He and Mr. Y.X.Zhu for their excellent drawings. This research was supported by the Guizhou Natural Science Foundation [Qiankehe LH (2016)7206]. We thank Laura Clavijo and two reviewers for their efforts to help us improving our manuscript. We also thank the National Gesneriaceae Resources Bank of GXIB, Gesneriad Committee of China Wild Plant Conservation Association and Gesneriad Conservation Center of China for their invaluable help.


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