Research Article |
Corresponding author: Xi-Qiang Song ( songstrong@hainanu.edu.cn ) Academic editor: Murielle Simo-Droissart
© 2023 Zhi-Heng Chen, Zhong-Yang Zhang, Xi-Qiang Song, Zhe Zhang.
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:
Chen Z-H, Zhang Z-Y, Song X-Q, Zhang Z (2023) Gastrodia bawanglingensis (Orchidaceae, Epidendroideae), a new species from Hainan Island, China. PhytoKeys 220: 39-50. https://doi.org/10.3897/phytokeys.220.95137
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Gastrodia bawanglingensis, a new species of Orchidaceae from Hainan Island, China, is described and illustrated. It is morphologically similar to G. theana, G. albidoides and G. albida with dwarf habits, scarcely opening flowers, elongated fruit stems, curved and fleshy perianth tubes and similar columns and lips, but can be easily distinguished from them by having a pair of lateral wings bent outwards at the apex of the column and lateral wings with acuminate tips lower than the anther. According to the IUCN Red List Categories and Criteria, the new species is assessed as Endangered (EN). The plastome of G. bawanglingensis is greatly reduced and reconfigured with approximately 30876 bp in size and 25.36% in GC content. Morphological characteristics and molecular phylogenetic results based on chloroplast gene sequences support the recognition of G. bawanglingensis as a new species within Gastrodia.
Gastrodieae, Hainan Tropical Rainforest National Park, holomycotrophic orchids, taxonomy, tropical rainforest
Gastrodia
During our field investigation in April 2021, Gastrodia specimens with significantly different floral morphology from all the known species in China were collected in the forests of Bawangling, Hainan Tropical Rainforest National Park. Further studies, based on examination of specimens and literature of Gastrodia (
The next generation sequencing technology (high-throughput sequencing) was applied to extract the total genomic DNA of plant materials and chloroplast splicing software GetOrganelle was used to assemble the plant genome (
To estimate the phylogenetic position of the Gastrodia sp. nov. within Gastrodia, phylogenies were reconstructed by Maximum likelihood (ML) and Bayesian Inference (BI) analyses using the coding sequences (CDSs). All plastomes were downloaded from the NCBI database except Gastrodia sp. nov. (
Phylogenetic tree reconstruction of Gastrodia using the maximum likelihood (ML) method based on chloroplast gene sequences of Gastrodia sp. nov. and 11 other species. Only the ML tree is shown, because its topology is nearly identical to that of the obtained BI tree. Numbers associated with the branches are BI posterior probabilities (PP) and ML bootstrap value (BS). The species name is followed by the accession number of the GenBank accession. D, Didymoplexis; E, Epipogium; G, Gastrodia.
Morphological observations of Gastrodia sp. nov. were based on living plants (four individuals) and dried herbarium specimens all belonging to the type specimen, which is kept in the
The plastome of the novelty is 30876 bp in length with its GC content approximately 25.36% (GenBank accession number: OP219766) (Fig.
Information on the chloroplast genomes of Gastrodia sp. nov. and other 11 species of Gastrodia.
Species | Length of chloroplast genome (bp) | GC content (%) | Number of genes | ||
---|---|---|---|---|---|
Protein coding genes | tRNA genes | rRNA genes | |||
Gastrodia angusta | 36.812 | 25.4 | 19 | 5 | 4 |
Gastrodia crispa | 30.582 | 25.7 | 19 | 5 | 4 |
Gastrodia elata | 35.304 | 26.8 | 20 | 5 | 3 |
Gastrodia flexistyla | 30.797 | 25.4 | 19 | 5 | 4 |
Gastrodia javanica | 31.896 | 24.8 | 18 | 4 | 4 |
Gastrodia longistyla | 30.464 | 24.8 | 18 | 5 | 3 |
Gastrodia menghaiensis | 30.118 | 24.9 | 19 | 4 | 3 |
Gastrodia peichatieniana | 29.696 | 25.9 | 18 | 5 | 4 |
Gastrodia shimizuana | 30.019 | 25.5 | 18 | 5 | 4 |
Gastrodia sp. | 29.944 | 25.8 | 18 | 5 | 4 |
Gastrodia sp. nov. | 30.876 | 25.4 | 19 | 5 | 3 |
Gastrodia uraiensis | 30.746 | 24.9 | 19 | 5 | 4 |
Our ML and BI phylogenetic trees constructed from the chloroplast gene sequences showed that the novelty belongs to the genus Gastrodia, and is related to G. uraiensis, G. flexistyla and G. crispa.
China. Hainan Province: Bawangling, Hainan Tropical Rainforest National Park, in tropical rainforest, 850–950 m elevation, 25 April 2022, Z.Y. Zhang 006 (Holotype,
Gastrodia bawanglingensis is similar to G. albidoides with dwarf habits, scarcely opening flowers, elongated fruit stems, curved and fleshy perianth tubes and similar columns and lips, but can be easily distinguished from the latter by having lateral sepals adnate to 4/5 of total length (vs. lateral sepals adnate to 1/2 of total length), lip with four ridges (vs. lip with two ridges), the absence of a column foot (vs. the presence of a column foot) and a pair of lateral wings bent outwards (vs. lateral wings upright) at the column apex (Table
Differences between Gastrodia bawanglingensis, G. albidoides, G. theana and G. albida.
Character | G. bawanglingensis | G. albidoides | G. theana | G. albida |
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Perianth tube | slightly verrucous in the middle and upper part, distinct verrucose apically | slightly verrucose towards apex, otherwise smooth | distinctly striate and verrucose throughout | distinctly verrucose throughout |
Lateral sepals | adnate, to 4/5 of their length | adnate, to 1/2 their length | adnate, 1/3–1/4 their length | adnate, 1/5–1/6 their length |
Petals | brownish, fleshy, petals whitish on both sides, triangular-ovate, ca. 1.5 × 1.0 mm | whitish, thin in texture, triangular-ovate, 0.8–1.0 × 0.6–0.8 mm | salmon-pink, thin in texture, narrowly triangular, 0.4–0.8 × 0.2–0.3 mm | whitish outside, orange inside, fleshy, oblong-ovate, ca. 1.5 × 1.0 mm |
Lip | red at the base, light green at the middle, reddish-brown apically and marginally, epichile rhombic-ovate, 5–7 nerved, disc thickened with four ridges, a pair of low ridges outside the two main ridges. truncate at base, hypochile with two whitish, globose, subsessile, nectarless calli, ca. 0.5 mm in diameter | pale green, epichile rhombic-ovate, 6–7-nerved, disc thickened with two ridges, rounded at base, hypochile with two whitish, globose, subsessile, nectarless calli, ca.1 mm in diameter | green, epichile ovate, 5-nerved, disc slightly elevate longitudinally at middle, with four ridges four ridges, arranged one behind the other. cordate at base, hypochile with two whitish, globose, subsessile, nectarless calli, ca.0.8 mm in diameter | white, epichile triangular, disc thickened with two ridges, truncate at base; hypochile with two whitish, globose, subsessile, nectarless calli, ca.1 mm in diameter |
Column | apex with a pair of lateral wings bent outwards; lateral wings with acuminate tips lower than anther | apex with a pair of lateral wings; lateral wings with acuminate tips superior to anther | apex with a pair of lateral wings bent inwards; lateral wings with acuminate tips superior to anther | with a pair of lateral wings distally; edges of lateral wings parallel to column |
Column foot | Absent | 1.5–1.8 mm | 1.5–1.8 mm | column foot very short |
Rostellum | 0.2 × 1.0 mm | 0.2 × 1.5 mm | 0.2 × 1.5 mm | Absent |
Terrestrial, leafless, achlorophyllous herbs. Roots few, slender, 1–7 cm long, ca. 0.5–0.7 mm in diameter. Rhizome fleshy, tuberous, fusiform, 3–4 cm long, 5–7 mm in diameter, dark brown, covered with numerous scales. Scales verticillate, lanceolate, dark brown,1–2 mm long. Inflorescence erect, terminal, 2.0–6.5 cm long, ca. 2.2 mm in diameter, white to orange-brown, peduncle 3–4 noded, ovate to broadly ovate, sheath membranous, 3–5 × 2–3 mm; rachis often less than 5 mm long. Bracts membranous, ovate to ovate-oblong, apex pointed, pale yellowish-brown, 4–6 mm long, 1.5–3 mm wide. Ovary 3–6 mm long, 2–3 mm in diameter. Flowers (1–) 2–4 (–6), erect, bell-shaped, slightly curved, not opening widely, 8–10 mm long, 4–5 mm in diameter. Flowers whitish on both surfaces, apex brownish, lip red at the base, light green at the middle, reddish-brown apically and marginally; column white. Sepals and petals united, forming a 5–lobed perianth tube, 8–10 mm long, slightly verrucous in the middle and upper part, distinctly verrucose apically. Sepals fleshy, thickened, similar. Lateral sepals fused to 4/5 of their length, whitish on both surfaces, apex is brownish; free lobe of dorsal sepal triangle, ca. 2.5 × 2.0 mm; free lobes of lateral sepals ovate, ca. 2.0 × 2.0 mm. Petals connate with sepals, free portions brownish, whitish on both sides, triangular-ovate, ca. 1.5 mm long, 1 mm wide, connate portions distinctly thickened and the inside is obviously reddish-brown, forming a pair of ridge-like structures inside the perianth tube and the other side of the ridge-like structures is flesh-coloured. Lip rhombic-ovate, base adnate to perianth tube, 3.5–4.5 × 2.0–2.2 mm; hypochile with two whitish, globose, subsessile, nectarless calli, ca. 0.5 mm in diameter; epichile 5–7 nerved, truncate at base, entire, disc thickened with four ridges, a pair of low ridges outside the two main ridges; the two main ridges fused into one before reaching the tip, main ridges much raised and tinged orange near apex. Column 4.2–4.5 × 1.6–1.8 mm, apex with a pair of lateral wings bent outwards; lateral wings with acuminate tips lower than anther; column foot absent; rostellum 0.2 × 1 mm; stigma located near base. Anther hemispherical, 0.6–0.7 mm in diameter; pollinia 2. Capsule ellipsoid, 1.2–1.8 cm long, 0.5–0.8 cm in diameter; pedicel elongating to 10–25 cm in fruit. Seeds fusiform, 1.6–2.2 mm long.
The new species is named after Bawangling, the mid-west State of Hainan Island where it was discovered in a vast area of primitive montane rainforest.
霸王岭天麻 (Chinese pinyin: bà wáng lǐng tiān má).
Gastrodia bawanglingensis is a terrestrial mycoheterotrophic species that grows in montane rainforests which are dominated by Dysoxylum gotadhora (Buch.-Ham.) Mabb., Livistona saribus (Lour.) Merr. and A.Chev., Hancea hookeriana Seem. and Engelhardia roxburghiana Lindl. at elevations from 850 m to 950 m and associated with other orchids, such as Anoectochilus hainanensis H.Z.Tian, F.W.Xing & L.Li, A. roxburghii (Wall.) Lindl., Oxystophyllum changjiangense (S.J.Cheng & C.Z.Tang) M.A.Clem., Dendrobium hainanense Rolfe, Cymbidium kanran Makino and Micropera poilanei (Guillaumin) Garay. So far, only the type subpopulation has been found in the tropical rainforest of Bawangling, in Hainan.
Endangered [EN D1]. Gastrodia bawanglingensis was discovered in the mountain rainforest of Bawangling in Hainan Tropical Rainforest National Park. Until now, only the type subpopulation, consisting of ca. 100 individuals, has been discovered in Bawangling. Since its number of mature individuals is fewer than 250, we assess it as Endangered (EN) using criterion D1 (
Gastrodia bawanglingensis was observed flowering and fruiting in April and May.
Flowers of Gastrodia bawanglingensis barely open and pollen massulae were observed on the stigma when flowers were dissected. Through field observation, it was found that the fruiting rate is very high. We bagged buds on 3 plants with 10 flowers in total prior to the anthesis, and found that each of them has evolved into fruit after 15 days. These observations indicate that the new species probably self-pollinates. Gastrodia is probably the only genus that contains species with completely cleistogamous flowers as confirmed by intensive monitoring. Self-pollination might be an adaptation to ensure reproduction, compensating for the defi- ciency of pollinators in the habitat (
Gastrodia bawanglingensis is most similar to G. albidoides (
1 | Sepals adnate to 4/5 of their length; lip light green at the middle, reddish-brown apically and marginally; lip disc with two ridges ranging from base to apex | G. bawanglingensis |
– | Sepals adnate up to 1/2 of their length; lip green or white at the middle, uniform coloured or orange-red towards apex; lip disc without distinct ridges, but with lamellae or keel | 2 |
2 | Flowers white, sub-erect; petals margin wrinkled; column foot very short, pedicel elongated in fruit | G. menghaiensis |
– | Flowers grey-brownish, horizontal or slightly bending; petals margin entire, column foot distinct; pedicel not elongated in fruit | 3 |
3 | Tepal tube without white spots; column cylindrical and thick; lip disc with a pair of longitudinal lamellae near apex | G. longitubularis |
– | Tepal tube with white spots; column flat and thin; lip disc with four keels | G. punctata |
We acknowledge the support from the Bawangling Region of the Hainan Tropical Rainforest National Park. We are very grateful to Prof. Yi-Bo Luo and Prof. Xiao-Hua Jin for their guidance. Mrs. Ling-Yi Cao is thanked for preparing the illustrations. We also thank Mrs. Min-Ting Jin and Mrs. Meng-Xue Wang for facilitating the checking of specimens and Mr. Jin-Qiang Wang from Hainan Tropical Rainforest National Park for his kind help with fieldwork. This study was supported by the Project for Orchidaceae Plant Resources Special Investigation of National Forestry and Grassland Administration (Grant No.2020070708).