Research Article |
Corresponding author: Suman Neupane ( sneupane2@murraystate.edu ) Academic editor: Yasen Mutafchiev
© 2023 Bui Hong Quang, Khang Sinh Nguyen, Tuan Anh Le, Le Thi Mai Linh, Quoc Luan Nguyen, Duy Hoang Vu Ngo, Lei Wu, Suman Neupane.
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:
Quang BH, Nguyen KS, Le TA, Linh LTM, Nguyen QL, Ngo DHV, Wu L, Neupane S (2023) Hedyotis konhanungensis (Rubiaceae): A new species from the central highlands of Vietnam. PhytoKeys 221: 73-84. https://doi.org/10.3897/phytokeys.221.95895
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A new species of Hedyotis L. (Rubiaceae), Hedyotis konhanungensis B.H. Quang, T.A. Le, K.S. Nguyen & Neupane, is described and illustrated from the central highlands of Vietnam based on morphological and phylogenetic evidence. The new species belongs to the morphologically diverse tribe Spermacoceae (ca. 1000 species) of the family Rubiaceae, which is represented by 70–80 species in Vietnam. The phylogenetic analysis, based on four DNA regions (ITS, ETS, petD, rps 16), confirms the new species’ placement within the genus Hedyotis – one of the largest genera in the tribe, comprising ca. 180 species across Asia and the Pacific. Hedyotis konhanungensis is morphologically distinct from all southeastern Asian Hedyotis L. in its set of traits such as leaf type (shape and thickness), growth habit, and floral parts (color of inflorescence axis and the shape of calyx lobes). The new species shows similarities with Hedyotis shenzhenensis, H. shiuyingiae, and H. yangchunensis from China in its herbaceous habit, fleshy ovate leaf blades, and dark purple floral parts, but it is phylogenetically distinct and can be distinguished from them by the following combination of morphological traits: habit with slightly smaller stature (<25 cm), broadly ovate or deltoid stipules with cuspidate apex and entire margin, and ovate or nearly ovate calyx lobes.
Gia Lai Province, Hedyotis-Oldenlandia complex, Indochina, phylogenetics, Spermacoceae, taxonomy
The Asian-Pacific genus Hedyotis L. (ca. 180 species) lies within a polymorphic, mainly herbaceous tribe Spermacoceae (ca. 1000 species) of the family Rubiaceae. In Vietnam, this tribe is represented by 70–80 species belonging to the genera Dimetia (Wight & Arn.) Meisn., Debia Neupane & N.Wikstr., Exallage Bremek., Leptopetalum Hook. & Arn., Hedyotis L., Involucrella (Benth. & Hook.f.) Neupane & N.Wikstr., Neanotis W.H.Lewis, Oldenlandia L., and Spermacoce L. (
During a botanical field survey in Kon Ha Nung Biosphere Reserve (
Our collected specimens were compared with all described species from southeast Asia and southern China by studying relevant literature and examining digital herbarium images. Morphological characters were recorded using Nikon SMZ745/SMZ745T stereoscopic microscope and photographs of vegetative and floral parts were taken both in the field and from the samples preserved in 70% ethanol using Canon EOS 7D. The type specimens have been stored in the following three herbaria (acronyms follow
To establish its phylogenetic position within the tribe Spermacoceae, total genomic DNA was extracted from silica-dried material with the DNeasy Plant Kit (Qiagen, Valencia, California, U.S.A.). Four DNA regions (nuclear genome: ITS, ETS; plastid genome: petD, rps16) that were used in our earlier studies (
We added the DNA sequences of our sample to our existing DNA data matrix (Suppl. material
The Maximum likelihood tree obtained from RAxML analysis on the concatenated data confirms the position of our sample from Vietnam within Hedyotis and distinct from H. yangchunensis, H. shenzhenensis, and H. shiuyingiae (Fig.
Vietnam. Central Highlands of Vietnam, Kon Ha Nung Biosphere Reserve, Gia Lai province: K’Bang District, Kon Pne Commune, 14°20'53"N, 108°20'48"E, primary evergreen forest slopes at elevation 1150 m a.s.l., 28 November 2021, Nguyen Quoc Luan, Ngo Duy Hoang Vu & Le Tuan Anh LTA 531 (holotype:
Hedyotis konhanungensis is similar to H. shenzhenensis, H. shiuyingiae and H. yangchunensis from southeastern China (Guangdong and Hongkong) in the morphology of the leaf blades, floral bracts, dichasial cymes, and fruits, but differs from them by its broadly ovate or deltoid (vs. triangular or broadly triangular) stipules with entire (vs. hairy or lacerated) margins and cuspidate (vs. acute) apex, suborbicular or broadly oval (vs. subovate or ovate to lanceolate) lowest floral bracts, ovate or nearly oval (vs. triangular or subulate to lanceolate) persistent calyx lobes on fruits, and stamens in long-styled flowers inserted in lower ¼ or near the base (vs. at the middle or near the mouth) of the corolla tube (Table
Morphological comparison of Hedyotis konhanungensis with its putative closest allies. The characters of H. shenzhenensis T.Chen, H. shiuyingiae T.Chen and H. yangchunensis W.C.Ko & Zhang are taken from
Morphological characters | H. konhanungensis | H. shenzhenensis | H. shiuyingiae | H. yangchunensis |
---|---|---|---|---|
Plant height, including the inflorescences (cm) | 15–25 | 20–40 | 15–35 | 30–39 |
Stem | terete or subterete, 8–18 cm long | terete or subterete,1–2 cm long | slightly tetragonal, 10–22 cm long | tetragonal or subterete, 15–30 cm long |
Leaf arrangement | evenly spaced along the stem | fascicled on short rosette-like stem | evenly spaced along the stem | evenly spaced along the stem |
petiole length (mm) | 4–6 | 0.5 | 5–10 | 15–20 |
blade shape and size (cm) | obovate to oval, 6–10 × 2–4 | elliptic, oblong-elliptic or obovate, 8.5–15 × 5–9 | broadl, lanceolate or obovate-lanceolate, 2–19 × 1.4–8 | narrowly elliptic, oblanceolate or elliptic-oblong, 3–12 × 1–4.5 |
Stipule shape and size (mm) | broadly ovate or deltoid, apex cuspidate, margin entire, 2.5–3 × 5–6 | triangular, apex acute, margin shortly hairy, 3–5 × 5–10 | triangular, apex acute, margin dense trichomes, 4–8 × 5–10 | triangular, apex and margin lacerate or deeply divided into several linear lobes, ca. 16 × 8 |
Inflorescence | Longistylous flowers lax or not congested into a head-like inflorescence | Longistylous flowers lax or not congested into a head-like inflorescence | Longistylous flowers congested into a head-like inflorescence | Longistylous flowers congested into cymose or capitate inflorescence |
Peduncle length (cm) | 2–4 | 10–18 | 10 | 3–4 |
Lowest floral bracts | suborbicular or broadly oval, 2–2.5 × 1.5–2 cm | subovate, 2.5–3 × 1.8–2 cm or sometimes larger | ovate to lanceolate, 0.6–2.5 × 0.2–1 cm | ovate, ca. 1.5 × 0.5–0.6 cm |
Calyx lobes | ovate or nearly oval, 1.7–2.2 × 0.8–1.4 mm | subulate, ca. 1.5 mm long | subulate, ca. 3 mm long | unknown |
Corolla | ||||
outer colour | bluish purple | white | purplish white | unknown |
tube dimension and shape | 6–7 × 1–1.5 mm, slightly enlarged both ends | ca.3 × 1.5–1.8 mm, slightly narrower near mouth | tube ca. 4.5 × 2–2.5 mm, enlarged at mouth | unknown |
Stamens of long-styled flowers | inserted in lower ¼ (or near the base) of corolla tube | inserted in upper ¼ (or near the throat) of corolla tube | inserted in ½ (or middle) of corolla tube | unknown |
Stamens of short-styled flowers | inserted in the upper ¼ (or near the throat) of corolla tube; | inserted near the throat of corolla tube; | inserted near base of corolla tube; | unknown |
filaments ca. 0.2 mm long, not exserted beyond corolla mouth | filaments ca. 0.8 mm long, exserted beyond corolla mouth | filaments ca. 0.3 mm long, not exserted beyond corolla mouth | ||
Persistent calyx lobes on fruits | ovate or nearly oval, 1.8–2.3 × 0.8–1.5 mm, unveined | narrowly triangular or subulate, 1.2–2 × 0.5–0.7 mm, unveined | narrowly triangular or subulate, 3–4 × ca.1 mm, unveined | lanceolate, 3–6 × 1.2–3 mm, veined |
Perennial herbs, erect, 15–25 cm tall, completely glabrous. Stem simple, rarely branched, terete, 8–18 cm long, 5–7 mm in diam.; internodes 1–1.5 cm long. Leaves 5–7 pairs per stem, decussate, evenly spaced along the stem, abaxial side dark green, adaxial side glossy, dark purple to purplish black; petioles 4–6 mm long; blade flattened, thick, fleshy (subcoriaceous when dried), obovate-lanceolate to nearly oval, 6–10 × 2–4 cm; base decurrent or cuneate; apex broadly acute or obtuse; midrib depressed adaxially and prominent abaxially; 4–5 secondary veins on each side of the midrib, inconspicuous on adaxial side. Stipule interpetiolar, fused to leaf bases or very shortly around stem, broadly ovate or deltoid, 2.5–3 mm long, 5–6 mm wide at base, flattened, dark purple outside; apex cuspidate or aristate, with aristae 3–4 mm long and 0.7–0.9 mm wide; margins entire. Inflorescence terminal, a compound dichasial cyme, 5–7 cm long, with 3–4 orders of branching, purplish, 30–50-flowered. Peduncle terete, 2–4 cm long; bracts subtending the basal branches of the inflorescence leaf-like, suborbicular or broadly oval, slightly concave, 2–2.5 × 1.5–2 cm, with apex rounded to broadly acute or obtuse, abaxially dark purple to dark bluish purple, pale green adaxially; bracts subtending the upper inflorescence branches smaller, ovate or lanceolate, 0.8–1.5 × 0.5–1 cm, with acute apex. Pedicels terete, 3–6 mm long, usually bluish purple or purplish, bracteolate or ebracteolate; bracteoles narrowly ovate or lanceolate, somewhat concave, ca. 2.5 mm × 1.5 mm, with apex broadly acute or obtuse. Flowers 4-merous, distylous. Calyx hypanthium, bluish purple, cupular, 0.8–1.2 mm long and wide, glossy and glabrous; lobes 4, sub-equal, ovate or nearly oval, somewhat longitudinally concave, 1.7–2.2 mm long, 0.8–1.4 mm wide, broadly acute to obtuse at apex, adaxially white or purplish white, abaxially bluish purple. Corolla narrowly infundibuliform or tubular, bluish purple outside, white pubescent inside; tube 6–7 mm long, 1–1.5 mm wide, slightly enlarged at both ends; lobes 4, strongly reflexed, triangular-ovate, 1.5–1.8 × 1.2–1.5 mm, acute, adaxially white or purplish white, sparsely puberulent near base on adaxial surface. Longistylous flowers: stamens inserted in the lower ¼ of corolla tube, 1.5–2 mm above the base of tube; filaments very short, ca. 0.2 mm long; anthers linear, 1–1.2 × 0.2 mm, dorsifixed, introrse; style filiform, 5.5–6.5 mm long, white, glabrous, 2-parted; lobes oblong, 0.6–0.7 mm long, abaxially minutely papillose, pure white, slightly exserted beyond the corolla mouth; ovary inferior, 2-locular, with numerous ovules, placentation axile. Brevistylous flowers: stamens inserted on the upper ¼ of corolla tube, 1–1.5 mm below the corollas mouth; filaments and anthers similar to those in the long-styled flowers; style filiform, 1.5–1.8 mm long, 2-parted; lobes oblong, 0.6–0.7 mm long, densely papillose abaxially. Disks fleshy, ring-shaped, concave in the center, glabrous, white. Fruits capsular, nearly cupuliform with slightly concave top, 2.5–3 mm in diam., glabrous, purplish, crowned by the persistent, unveined, oval or nearly oval calyx lobes. Seeds many, black, irregularly angular, reticulate, minute, 0.3–0.5 mm long.
Hedyotis konhanungensis A habit B stipule C inflorescence D floral bracts E bracteoles and fruits F, G calyx top and side views H–K long-styled flower, entire and open showing pistil, corolla and anthers L–P short-styled flower, dissection showing calyx, corolla, anthers and pistil Q–T fruit, entire and dissection U seeds. Photos and design by B.H. Quang from Luan et al. LTA 531.
(Paratypes). Vietnam. Central Highlands of Vietnam. Kon Ha Nung Biosphere Reserve, Central Highlands of Vietnam, Gia Lai province: K’Bang District, Kon Pne Commune, 14°20'43.06"N, 108°20'38.33"E, 906 m a.s.l., 26 March 2022, Bui Hong Quang et al. BHQ 453 (HN, and herbarium of Kon Ka Kinh National Park).
This species is named after the “Kon Ha Nung Biosphere Reserve” where it was discovered.
Vietnamese: An điền Kon Hà Nừng
Flowering in October to November, fruiting from November to December.
Hedyotis konhanungensis is recorded only from the type locality in the Kon Pne Commune of the Central Highlands of Vietnam, which is part of the Annamite Range. This range is known for its rich biodiversity and high number of endemic species (
The genus Hedyotis, as currently circumscribed, corresponds to a primarily woody genus characterized by “diplophragmous” capsules (septicidally dehiscent capsules that separate into two distinct valves, as described by Wight and Arnott in 1834) and “fruiticosa type” seeds (dorsiventrally flattened seeds with a ventral hilar ridge topped by a punctiform apical hilum, as described by Terrell and Robinson in 2003). The members of this hyperdiverse genus (approximately 180 species) primarily occupy the mountains of Asia and the Micronesian Islands (in the northwestern Pacific). The molecular dating analysis suggests that Hedyotis split from its sister lineages of approximately 10 species (the African/Malagasy Agathisanthemum group and the African-tropical Asian-North American Edrastima group) nearly 27 million years ago (
The authors gratefully acknowledge the help of the directors, curators and staff of the following herbaria: HN, P, VNM, and
Taxa and NCBI GenBank accessions used to estimate phylogeny of Spermacoceae for this study
Data type: Excel datasheet
A concatenated nexus alignment of four DNA regions (ITS, ETS, petD, rps16) used to estimate phylogeny of Spermacoceae for this study
Data type: Nexus alignment
Output of RAxML tree analysis with bootstrap values (1000 replicates)
Data type: Newick tree file