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Research Article
Gastrochilus heminii (Orchidaceae, Epidendroideae), a new species from Sichuan, China, based on molecular and morphological data
expand article infoMin Liao‡§, Yue-Hong Cheng|, Jun-Yi Zhang‡, Yu Feng‡, Gui-Ying Liu|, Ping Ye|, Sen-Long Jin|, Hong-Qiang Lin|, Bo Xu‡§
‡ Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu, China
§ University of Chinese Academy of Sciences, Beijing, China
| Wolong National Natural Reserve Administration Bureau, Wenchuan, China
¶ Chongqing Normal University, Chongqing, China

Corresponding author: Bo Xu ( xubo@cib.ac.cn )

Academic editor: Timothée Le Péchon
© 2022 Min Liao, Yue-Hong Cheng, Jun-Yi Zhang, Yu Feng, Gui-Ying Liu, Ping Ye, Sen-Long Jin, Hong-Qiang Lin, Bo Xu.
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: Liao M, Cheng Y-H, Zhang J-Y, Feng Y, Liu G-Y, Ye P, Jin S-L, Lin H-Q, Xu B (2022) Gastrochilus heminii (Orchidaceae, Epidendroideae), a new species from Sichuan, China, based on molecular and morphological data. PhytoKeys 215: 95-106. https://doi.org/10.3897/phytokeys.215.91061
Open Access

Abstract

Gastrochilus heminii (Orchidaceae), a new orchid species from Sichuan Province, Southwest China, is described and illustrated. It morphologically resembles G. affinis and G. yei, but differs markedly from the former in having a thinner and slightly rolled downwards reniform epichile and the central thickened purple-red mat with irregular folds (vs. subtriangular epichile curves upwards, with 2 thick, brown to purplish-brown median ridges from base to apex), and can be clearly distinguished from the latter by having reniform epichile with lobed apex and subconical hypochile with bilobed apex that splits into two conical protrusions (vs. semi-rounded epichile not lobed and subconical hypochile not bilobed). The results of molecular phylogenetic analysis based on nuclear ribosome internal transcribed spacer (nrITS) and four chloroplast DNA fragments (matK, psbA-trnH, psbM-trnD, and trnL-F) of 36 Gastrochilus species showed that G. heminii was closely related to G. affinis and G. distichus.

Keywords

Gastrochilus, morphological, Orchidaceae, phylogeny, taxonomy

Introduction

Gastrochilus D.Don (Orchidaceae, Epidendroideae; Don 1825) is diagnosed by a subdivided labellum with a recurved epichile and a saccate hypochile attached to the column at the base and two subglobose pollinia attached to a slender and filiform stipe (Tsi 1999; Pridgeon et al. 2014). The genus comprises more than 70 accepted species, most of which are distributed in Southeast Asia (Kumar et al. 2014; Liu et al. 2016, 2019; Liu and Gao 2018; Rao et al. 2019; Govaerts et al. 2021; Nguyen et al. 2021; Zhou et al. 2021a, b; Zhang et al. 2022). Liu et al. (2019) first constructed the phylogenetic relationships of Gastrochilus and divided them into five clades. Recently, the phylogenetic studies by Liu et al. (2020), Li et al. (2022), and Zhang et al. (2022) also supported the monophyly of Gastrochilus.

An unusual arboreal miniature orchid was recently discovered for the first time in Wolong Nature Reserve (Wenchuan County, Sichuan Province, China) during a routine survey. It was tentatively identified as a species of Gastrochilus sect. Microphyllae Bentham & Hooker (Bentham and Hooker 1883), characterized by plants with extremely small flowers, distichous and alternate leaves compared to plants in the other sections (larger flowers or clustered leaves). However, a morphological examination revealed that this species shows an unusual combination of characters that does not match any known species of G. sect. Microphyllae. The phylogenetic analysis using five DNA markers (nrITS, matK, psbA-trnH, psbM-trnD, and trnL-F) confirmed the monophyly of this taxon. The objectives of this study are (1) to describe, (2) to examine both molecular and morphological affinities of this new Gastrochilus species, Gastrochilus heminii M.Liao, B.Xu & Yue.H.Cheng, sp. nov.

Materials and methods

Morphological analyses

The measurements and description of Gastrochilus heminii were based on two living plant individuals and two herbarium specimens (voucher information: Min Liao & Yue-Hong Cheng ZJY143; Min Liao & Yue-Hong Cheng ZJY167), respectively. The taxonomic description follows the terminology used by Beentje (2012). Voucher specimens and additional silica-gel dried leaves are deposited in CDBI Herbarium (herbarium follows Thiers 2021).

DNA extraction, amplification and sequencing

The sequences of the two individuals of the new species newly generated in this study, and the sequences of the remaining 42 species used in the molecular phylogenetic analysis, were retrieved from GenBank. The information on the DNA fragments and four complete plastid genomes were listed in Appendix 1. Total DNA was extracted exclusively from silica-gel dried leaves via a Plant DNA Isolation Kit (Cat.No.DE-06111). We used the same primers as Liu et al. (2019) to amplify the nuclear ribosome internal transcribed spacer (nrITS) and the four chloroplast DNA fragments (i.e., matK, psbA-trnH, psbM-trnD, and trnL-F) through polymerase chain reaction (PCR). All DNA samples were sent to TSINGKE Biotech Co. Ltd (Chengdu, China) for sequencing. The sequences were then deposited with GenBank, with the following accession numbers: G. heminii, nrITS (ON286752, ON286753), matK (ON331126, ON331127), psbA-trnH (ON331128, ON331129), psbM-trnD (ON331130, ON331131), and trnL-F (ON331132, ON331133).

Phylogenetic analyses

All sequences were edited via Sequencher v4.1.4 (Gene Codes, Ann Arbor, Michigan, USA) and aligned via MAFFT v7.475 (Katoh and Standley 2013) with default parameters. We performed phylogenetic analyses based on combined nuclear ribosome internal transcribed spacer (nrITS) and the four chloroplast DNA fragments. The nucleotide substitution model for the data matric was estimated using jModeltest v2.1.6 (Posada 2008) software and the best-fit model (GTR+I+G) was selected using the corrected Akaike Information Criterion (AICc). Two different methods, Maximum likelihood (ML) and Bayesian inference (BI), were employed. The ML analysis was performed using IQ-TREE v1.4.2 (Nguyen et al. 2014) with branch support estimated by 2,000 replicates of ultrafast bootstrapping algorithm (UFboot) (Minh et al. 2013). The BI analysis was conducted using MrBayes v3.2.7a (Ronquist and Huelsenbeck 2003), with two separate Markov-chain Monte Carlo (MCMC) chains (1,000,000 generations and sampled every 1,000 generations). The first 25% of the trees were discarded as burn-in, and the remaining trees were used to generate a majority-rule consensus tree.

Results

The molecular phylogenetic tree showed that the 36 taxa of Gastrochilus formed a well-supported monophyletic group (BI/ML = 1/97, Fig. 1). The two individuals of G. heminii were resolved as sisters to each other (BI/ML = 1/99, Fig. 1). Our data recovered a sister relationship between G. affinis (King & Pantl.) Schltr. (King and Pantling 1898; Schlechter 1913) and G. distichus (Lindl.) Kuntze (Lindley 1859; Kuntze 1891); these two species formed a monophyletic group with G. heminii (BI/ML = 0.90/89, Fig. 1), which formed a subclade of section Microphyllae together with G. fargesii (Kraenzl.) Schltr. (Kraenzlin 1903; Schlechter 1919), G. alatus X.H.Jin & S.C.Chen (Jin et al. 2007), G. fuscopunctatus (Hayata) Hayata (Hayata 1912, 1917), and G. pseudodistichus (King & Pantl.) Schltr. (King and Pantling 1895; Schlechter 1913).

Figure 1. 

Maximum likelihood tree of Gastrochilus from phylogenetic analysis of combined nrITS and plastid DNA markers. Numbers before and after the slash indicate Bayesian posterior probabilities and ML bootstrap supports, respectively. The position of Gastrochilus heminii is indicated in red.

Morphologically, G. heminii is closest to G. affinis and G. yei JianW.Li & X.H.Jin (Li et al. 2021). A thorough morphological comparison among G. heminii, G. affinis, G. yei and G. distichus is summarized in Table 1 and Fig. 2.

Table 1.
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Morphological comparison of Gastrochilus heminii with three related species, G. affinis, G. distichus and G. yei.

Character G. heminii G. affinis G. distichus G. yei
Plant length 3.0–6.5 cm 4.0–15.0 cm 1.5–20.0 cm 3.0–8.0 cm
Leaf shape narrowly oblong or falcate oblong oblong-lanceolate to subspathulate lanceolate or falcate lanceolate blade lanceolate
No. of flowers per inflorescence 1 or 2 flowers 1–4 flowers 2–4 flowers 2–6 flowers
Peduncle length 0.4–0.7 cm 1.5–2.0 cm 1.0–1.5 cm 0.7–1.0 cm
Dorsal sepal elliptic-oblong, ca. 2.4 × 1.5 mm, concave, apex obtuse elliptic-oblong, 3–5 × 1–1.3 mm, concave, apex obtuse oblong-elliptic, 4.5–0.5 × 2.5–3 mm, concave, apex obtuse oblong, 3.3 × 1.9–2.0 mm, apex rounded
Lateral sepals similar to dorsal sepal, equal in size elliptic-ovate, 3.5–4 × 0.7–1.3 mm, slightly oblique and incurved, apex obtuse similar to dorsal sepal, equal in size oblong, 3.9–4.0 × 1.8–1.9 mm, apex obtuse
Petals narrowly oblong, ca. 2.6 × 1.3 mm, apex acute, base narrowed ovate-elliptic to elliptic, 3–4 × 1–1.3 mm, apex obtuse subobovate, slightly smaller than sepals, apex obtuse oblong, 3.5 × 1.8 mm, apex rounded
Epichile reniform, 4.2–6.5 × 2.0–3.0 mm, margin erose, smooth and glabrous above, central thickened purple-red mat with irregular folds subtriangular, decurved, subacute at apex, margin finely erose at base, disk with 2 thick, brown to purplish-brown median ridges from base to apex. subcircular, 5 × 3 mm, apex obtuse, margin entire, smooth and glabrous above and thickened cushion-like centrally, with 2 conical callosities near base semi-rounded, 2.0–2.2 × 4.0–4.2 mm, glabrous, with a thicken central, rugose cushion, tint with purple, margin irregularly denticulate
Hypochile subconical or helmet-shaped, ca. 2–2.4 × 1.6–2 mm, dorsally compressed, slightly bent outward, the end splits into two conical protrusions obconical, 3–4 × 2–3 mm, dorsally compressed, slightly bent outward, subacute to obtuse and shortly bifid at apex subcupular, 4 × 2–3 mm, rounded at end, dorsally compressed, slightly bent outward subconical, 3 mm tall, 3 mm in diameter, apex rounded
Figure 2. 

Comparison of three taxa of Gastrochilus A, B Gastrochilus heminii C, D G. affinis E, F G. distichus G, H G. yei. [Images C, D cited from Jalal et al. 2020; image E reproduced from website (http://www.orchidspecies.com/gastdistichus.htm); image F cited from Kumar et al. 2014, which was photographed by Xiao-Hua Jin; images G and H cited from Li et al. 2021].

Taxonomic treatment

Gastrochilus heminii M.Liao, B.Xu & Yue H.Cheng, sp. nov.

urn:lsid:ipni.org:names:77310051-1
Figs 2A, B, 3

Type

China. Sichuan: Wenchuan, coniferous and broadleaf mixed forest, on tree trunk, elev. ca. 2640 m, in flowering and fruiting, 15 March 2022, Min Liao & Yue-Hong Cheng ZJY143 (holotype CDBI!).

Diagnosis

Gastrochilus heminii is morphologically related to G. affinis and G. yei based on vegetative and floral characteristics such as similar habit, distichous and alternate leaves, epichile surface smooth and glabrous, sepals and petals with purplish-red patches. However, it can be differentiated from G. affinis on the basis of flower numbers (1–2 in the former vs. 1–4 in the latter), peduncle length (0.4–0.7 cm in the former vs. 1.5–2.0 cm in the latter) and an additional morphological characteristic: young leaves are densely covered with purple-red spots and old leaves have hardly any purple-red spots in the former (both have purple-red spots in the latter); the reniform epichile is rolled downwards, smooth and glabrous above, and central thickened purple-red mat with irregular folds in the former (subtriangular epichile curves upwards, with 2 thick, brown to purplish-brown median ridges from base to apex in the latter). It differs from G. yei by having reniform and lobed epichile (not lobed in the latter), apex of hypochile bilobed and splits into two conical protrusions (not bilobed in the latter), apex of the leaf with 1–2 lobules, lobes setaceous (apex of leaf with a tine in the latter).

Figure 3. 

Gastrochilus heminii A flowering plant B young leaves (B1: front view; B2: reverse view) C old leaves (C1: front view; C2: reverse view) D raceme and capsule (side view) E–I flowers (different views) J anatomy of flower (J1: dorsal sepals; J2 and J3: lateral sepals; J4 and J5: petals; J6: labellum) K hypochile (dorsal view) L stamens.

Description

Miniature trunk epiphyte. Roots vermiform, 2.0–7.0 cm long and ca. 1.5 mm thick. Stem pendulous, slender, purplish-red spots, 3.0–6.5 cm long and ca. 1.8 mm thick. Leaves alternate, distichous, narrowly oblong or falcate oblong, 0.9–2.3 × 0.3–0.5 cm, apex acute and with 1–2 lobules, lobes setaceous, young leaves with conspicuous purplish-red spots, old leaves with hardly any purplish-red spots. Raceme with 1 or 2 flowers; inflorescence stalk curved upward and thickened, 4.0–9.0 mm long, proximally covered with two sheaths; floral bracts ovate-lanceolate, 0.7–1.0 mm long, apex acute; pedicel and ovary connate, 4.0–5.5 mm long; flowers spreading, ca. 6.0 × 5.0 mm; sepals and petals heterochromatic on both surfaces, outside yellow-green with purplish-red spots, inside purplish-red with yellow-green margin; dorsal sepals and lateral sepals similar and equal in size, elliptic-oblong, ca. 2.4 × 1.5 mm, apex obtuse; petals narrowly oblong, ca. 2.6 × 1.3 mm, apex acute, base narrowed; epichile reniform, yellow-green with purplish-red spots, 4.2–6.5 × 2.0–3.0 mm, margin erose, smooth and glabrous above, central thickened purple-red mat with irregular folds; hypochile subconical, yellow-green with purple-red spots, 2–2.4 × 1.6–2 mm, dorsally compressed, slightly bent outward, the apex splits into two conical protrusions; column cylindrical, ca. 1.0 mm; anther cap subhemispheric, with two chambers, 0.7 × 0.4 mm, hanging from both ends of the stipe; pollinia 2, 0.4 × 0.3 mm, yellow, full and nearly spherical, with a depression in the center; stigma deeply sunken, inverted V-shaped, ca. 0.6 mm long, yellow, apically forked, forked in a subtriangular outline. Capsule shuttle-shaped with six ribs, green with sparse purplish-red spots, ca. 1.1 cm long, inflated to ca. 0.6 cm in the middle, persistent and growing for one year until maturity.

Additional specimens examined

—China. Sichuan: Wenchuan, coniferous and broadleaf mixed forest, on tree trunk, elev. ca. 2640 m, 18 April 2022, Min Liao & Yue-Hong Cheng ZJY167 (CDBI).

Distribution, habitat and phenology

The new species is currently known only from Wenchuan County, Sichuan Province, Southwest China (Fig. 4). It is found epiphytic on the trunk of Tsuga chinensis (Franch.) Pritz. in a subalpine mixed coniferous forest at elevation ca. 2640 m. Gastrochilus heminii flowers from March to April.

Figure 4. 

Distribution map of Gastrochilus heminii.

Etymology

Named in honor of Mr. He-Min Zhang, the advocate of the panda “Release to the Forest”, one of the pioneers of panda research in China, renowned as the “father” of pandas, in recognition of his contribution to the conservation of flora and fauna in the region which was one of the earliest reserves dedicated to preserving the habitat of wild pandas. A Chinese name, he min peng ju lan (和民盆距兰), is suggested here.

Conservation status

Three populations with a total of ca. 200 individuals of Gastrochilus heminii have been documented during our investigation. However, similar habitat of this new species is widespread in the Wolong Natural Reserve. Therefore, we assessed the conservation status of Gastrochilus heminii as DD (Data Deficient) according to the IUCN (2022).

Acknowledgements

Our work was funded by the National Key Research and Development Program of China (Grant No. 2020YFE0203200), the Second Tibetan Plateau Scientific Expedition and Research (STEP) program (Grant No. 2019QZKK0502), 2022 Central Finance Forestry Grassland Ecological Protection and Restoration National Park Subsidy Project, 2022 Subsidy Projects of Prohibited Developmental Areas from the Transfer Payment of the National Key Ecological Functional Areas, and Wild Plants Sharing and Service Platform of Sichuan Province.

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Appendix I

Table A1.
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The GenBank accession numbers for DNA sequences used in this study.

Taxa nrITS matK psbA-trnH trnL-F psbM-trnD
G. acaulis (Lindl.) Kuntze KM583455 KM583465 — — —
G. acinacifolius Z.H.Tsi KJ733412 KJ733569 KJ733492 KJ733649 MK357216
G. acutifolius (Lindl.) Kuntze MT225573 MW433889 — — MK357230
G. affinis (King & Pantl.) Schltr. — MK357141 — — MK357141
G. alatus X.H.Jin & S.C.Chen — — — — MK357228
G. bellinus (Rchb.f.) Kuntze KY966597 KY966884 MK357164 MK357202 MK357240
G. bigibbus (Rchb.f. ex Hook.f.) Kuntze — MN124439 MN124439 MN124439 MN124439
G. calceolaris (Buch.-Ham. ex Smith) D.Don MN517123 MK357144 MK357169 MK357204 MK357232
G. changjiangensis Q.Liu & M.Z.Huang MK357124 — MK357166 — MK357236
G. ciliaris F.Maekawa — MK357148 MK357173 — MK357225
G. dasypogon (Lindl.) Kuntze DQ091685 MK357149 MK357181 MK357197 MK357219
G. deminutus J.M.H.Shaw KY966600 KY966887 — — —
G. distichus (Lindl.) Kuntze KJ733414 KJ733571 KJ733494 KJ733651 —
G. fargesii (Kraenzl.) Schltr. KJ733415 KJ733572 — KJ733652 —
G. formosanus (Hayata) Hayata KJ733416 KJ733573 KJ733495 KJ733653 MK357226
G. fuscopunctatus (Hayata) Hayata MK317970 MK357150 MK357171 MK357192 MK357231
G. gongshanensis Z.H.Tsi — MN124438 MN124438 MN124438 MN124438
G. guangtungensis Z.H.Tsi KJ733417 KJ733574 KJ733496 KJ733654 KJ733654
G. heminii M.Liao, B.Xu & Yue.H.Cheng ON286752 ON331126 ON331128 ON331130 ON331132
G. heminii M.Liao, B.Xu & Yue.H.Cheng ON286753 ON331127 ON331129 ON331131 ON331133
G. intermedius (Griff. ex Lindl.) Kuntze MK357121 MK357151 MK357172 MK357190 MK357213
G. japonicus (Makino) Schltr. KJ733418 KF545886 KF545866 KF545897 KX871236
G. linearifolius Z.H.Tsi & Garay MK357133 MK357136 MK357187 MK357194 MK357229
G. linii Ormerod — MK357152 MK357176 MK357198 MK357224
G. malipoensis X.H.Jin & S.C.Chen — MK357147 MK357177 MK357200 MK357235
G. minutiflorus Aver. — MK357153 MK357179 — MK357215
G. obliquus (Lindl.) Kuntze MK357131 MK357137 KJ733498 KJ733656 MK357211
G. platycalcaratus (Rolfe) Schltr. MK357122 — MK357175 — MK357222
G. pseudodistichus (King & Pantl.) Schltr. MK357132 — MK357170 — MK357221
G. rantabunensis C.Chow ex T.P.Lin — MK357155 MK357184 MK357193 MK357223
G. raraensis Fukuyama KJ733420 KJ733577 KJ733499 KJ733657 MK357239
G. sinensis Z.H.Tsi OM985813 OK042953 OK172399 OK172401 —
G. somai (Hayata) Hayata MK357128 MN124436 MK357180 MN124436 MK357220
G. tianbaoensis Q.Liu & Y.H.Tan MK357120 MK357157 MK357186 MK357207 MK357214
G. wolongensis Jun.Y.Zhang, B.Xu & Yue.H.Cheng OM985810 OK172400 OK172402 OK172404 OK172403
G. wolongensis Jun.Y.Zhang, B.Xu & Yue.H.Cheng OM985811 OM974209 OM974211 OM974210 —
G. yunnanensis Schltr. MK165469 MK357158 MK357185 — MK357212
G. zhenyuanensis Q.Liu & D.P.Ye MK357127 MK357146 MK357168 MK357199 MK357237
Holcoglossum amesianum (Rchb.f.) Christenson HQ404389 JF763779 HQ404439 — —
Holcoglossum kimballianum (Rchb.f.) Garay HQ452901 JF763787 HQ404452 — —
Luisia magniflora Z.H.Tsi & S.C.Chen KJ733426 KJ733583 KJ733505 — KJ733663
Pomatocalpa diffusum Breda AB217576 AB217752 — — EF670432
Pomatocalpa spicatum Breda DQ091706 KJ733595 KJ733518 — KJ733675
Saccolabium pusillum Blume AB217580 AB217756 — — —
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