Research Article |
Corresponding author: Tao Peng ( pengtao@gznu.edu.cn ) Academic editor: Laura Clavijo
© 2022 Tao Peng, Shun-Zhi He, Shun-Li Wang, Dan Huang, Xu-Ping Zhou.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Peng T, He S-Z, Wang S-L, Huang D, Zhou X-P (2022) Hemiboea kaiyangensis, a new species of Gesneriaceae endemic to Guizhou, China. PhytoKeys 211: 63-73. https://doi.org/10.3897/phytokeys.211.85630
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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 (
So far, a total of 38 species and six varieties have been described in Hemiboea (
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 (
To investigate the phylogenetic placement of the newly collected taxon, our sampling focused on species of Hemiboea and its closest related genera according to
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 |
Sequences of ITS and trnL-F of the new species were aligned with the existing matrices of
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.
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
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 |
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
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.
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.
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.
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.
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 (
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
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.