Research Article |
Corresponding author: Siti-Munirah Mat Yunoh ( sitimunirahfrim1@gmail.com ) Academic editor: Eberhard Fischer
© 2021 Siti-Munirah Mat Yunoh.
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:
Yunoh S-MM (2021) Chroesthes (Acanthaceae) in Peninsular Malaysia, including a new species from Kelantan and a new record from Terengganu. PhytoKeys 174: 127-146. https://doi.org/10.3897/phytokeys.174.62023
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Chroesthes is a small genus that includes three species from Peninsular Malaysia: Chroesthes faizaltahiriana Siti-Munirah sp. nov., C. lanceolata (T. Anderson) B.Hansen and C. longifolia (Wight) B.Hansen. Chroesthes faizaltahiriana, recently discovered in the State of Kelantan, is described and illustrated. This species is similar to the common species C. longifolia, but is distinguished mainly by its inflorescence type, calyx shape and its flowers being bright orange instead of dark purple internally. Chroesthes lanceolata is a new record for Peninsular Malaysia and has only been collected once. Following the IUCN Red List Categories and Criteria, these three species are assessed (national scale assessment) as Critically Endangered (C. faizaltahiriana and C. lanceolata) and Least Concern (C. longifolia).
Conservation status, revision, taxonomy
The small genus Chroesthes
The Forest Research Institute Malaysia (
During a recent botanical survey at Berangkat Forest Reserve (FR), Kelantan, an upright shrub with an unusual bright orange corolla colour was encountered growing in a patch of forest, under shade beside an old logging road. Its morphological characteristics, including its stamen type, indicated that it belongs to the genus Chroesthes, but is unmatched amongst the three species currently recognised. This new species is described here as Chroesthes faizaltahiriana Siti-Munirah, which brings the total species for Chroesthes in the world to four. It is also an additional endemic species in Peninsular Malaysia. With this recent discovery, an account of the genus Chroesthes in Peninsular Malaysia is provided, including the key to Chroesthes of the world; however, only the Malaysian species are considered in the remainder of the treatment.
Specimens of Chroesthes species from Peninsular Malaysia, held in the herbaria at Kepong Herbarium (KEP) and Singapore Botanic Gardens Herbarium (
Bull. Mus. Natl. Hist. Nat. 33: 107. 1927, in Fl. Gen. I.C. 4: 684. 1935; Hansen, Nordic J. Bot. 3: 209. 1983; Hu, J.C., Deng, Y.F., Daniel, T. & Wood, J.R.I. Acanthaceae. In: Wu, Z.Y., Revan, P. & Hong, D.Y. (Eds.) Flora of China 19: 472. 2011.
Shrubs; cystoliths present. Leaves opposite, petiolate; leaf blade margin entire; subisophyllous or anisophyllous. Inflorescences terminal thyrses (the thyrses are branched (then paniculiform) or unbranched (then racemoid); bracts and bracteoles greenish. Calyx unequally five-lobed: posterior lobe largest, two lateral lobes smaller than two anterior lobes. Corolla tube basally cylindrical, expanded distally into a throat; limb two-lipped, upper lip two-lobed, lower lip three-lobed; lobes quincuncial in bud; four stamens, connate to the corolla, not connate to one another, posterior pair shorter than anterior pair, inserted at the base of the corolla throat; anthers bi-thecous; thecae parallel, inserted at different heights, dorsally pubescent, base of each theca spurred; ovary with two ovules per locule; style basally sparsely pubescent; stigma capitate. Capsule stipe absent or barely present; retinacula present. Seeds compressed, brownish, shortly pubescent.
China, Laos, Malaysia, Myanmar, Thailand, Vietnam. Three species in Malaysia.
Lowland dipterocarp forest to upper hill dipterocarp forest.
1 | Bracts approximately half the length of the calyx | C. lanceolata |
– | Bracts approximately as long as the calyx | 2 |
2 | All calyx lobes narrow, linear | C. bracteata (only in Thailand) |
– | Posterior and anterior calyx lobes elliptic to elliptic-lanceolate, the lateral lobes linear | 3 |
3 | Inflorescence terminal branched; corolla entirely dark purple to purplish-red or sometimes white externally | C. longifolia |
– | Inflorescence terminal unbranched; corolla yellow to dark orange | C. faizaltahiriana |
Chroesthes faizaltahiriana most closely resembles C. longifolia; however, it differs in its inflorescence type, the presence of a terminal raceme not branching (vs. terminal raceme branching) and posterior lobe size ratio 1:4 (vs. 1:2) and corolla length 4.5–5.5 cm (vs. 2–3 cm) and in the corolla tube and lobes being entirely bright yellow to dark orange (vs. entirely dark purple, purplish-red or occasionally white externally) and other significant characters (see Table
Morphological comparison between Chroesthes faizaltahiriana and C. longifolia.
Character | C. faizaltahiriana | C. longifolia |
---|---|---|
Habit | ||
Height (m) | 0.7–1 | 1–2.5 |
Stem | Unbranched | Branching |
Inflorescence | ||
Type | Terminal raceme not branching (single) | Terminal raceme always branching (always two) |
Length (cm) | 8–13 | up to 25 |
Flowers | ||
Bract (mm) | 20–30 × 2–5 | 15–20 × 4.5–6 |
Shape | Narrowly lanceolate | Lanceolate |
Bracteoles (mm) | 10–15 × 1–2 | 10 × 3 |
Calyx length (cm) | 2–3 | 1.6–1.8 |
Posterior lobe (mm) | 30 × 7 | 18 × 8–9 |
Anterior lobe (mm) | 22 × 2–2.5 | 16 × 5–6 |
Lateral lobe (mm) | 20 × 1 | 10–12 × 0.5 |
Calyx colour | Completely always greenish | Green or green to purplish (especially purplish at apex) |
Total corolla length (cm) | 4.5–5.5 | 2–3 |
Corolla colour | Entirely yellow to dark orange: pale yellow externally, bright orangish-yellow with darker spots internally | Entirely dark purple to purplish-red or sometimes white externally |
Filament length (cm) | ||
Longer pair | 1.5 | 1 |
Shorter pair | 1.2 | 0.8 |
Chroesthes faizaltahiriana Siti-Munirah A habit of the whole plant B inflorescence (front view) C inflorescence (side view) D flower with bracteoles and calyx E perianth lobes (flower from front view) showing the anthers F calyx and bracteoles G bract H bracteoles I calyx (a) anterior, (l) lateral, (p) posterior lobes (photo of a dry specimen) J corolla (inner view) K corolla (outer view) L stamens. (All photos by Siti-Munirah MY).
Malaysia. Peninsular Malaysia: Kelantan, Gua Musang Distr., Berangkat FR, ca. 822 m alt., 26 Feb 2020, Siti-Munirah, FRI 91215 (holotype KEP!, barcode KEP 280001).
Shrubs 0.7–1 m high. Stems terete, erect, not branched, surface glabrous, diameter ca. 2 mm, swollen at nodes. Leaves opposite; petiole 0.2–1 cm long; straight or twisted (makes the leaf arrangement look decussate); leaf blades elliptic, lanceolate to oblanceolate, 4.5–19 × 1.5–5.5 cm, both surfaces glabrous, lateral veins ca. 8–10 on each side of mid-vein, base attenuate to cuneate, margin entire, apex acute to acuminate. Inflorescence a terminal raceme (unbranched), up to 13 cm long; flowers secund (one bract at each node being sterile, the other bracts subtending each flower), ca. 10-flowered; glandular-pubescent on most parts; peduncles ca. 1 cm; bracts narrowly lanceolate, 20–30 × 2–5 mm, apex acute, glandular-pubescent, conspicuously 1-nerved; bracteoles narrowly lanceolate, 10–15 × 1–2 mm; pedicel short, ca. 1–2 mm long. Calyx 2–3 cm long, posterior lobe lanceolate, 30 × 7 mm, 3–5-nerved, apex acuminate; two anterior lobes, elliptic-lanceolate, 22 × 2–2.5 mm, 1-nerved, apex acuminate; two lateral lobes, linear-lanceolate, 20 × 1 mm, 1-nerved; all glandular-pubescent on both sides; all greenish. Corolla bilabiate, orange-yellow, ca. 4.5–5.5 cm long; outer surface pale yellow, glandular-pubescent; inner surface bright orange-yellow to orange with dark orange spots (or stripes), glabrous; tube with cylindrical basal portion ca. 2 cm long, expanded throat ca. 1.5 cm long; upper lip shortly two-lobed (8–9 mm long), lower lip deeply three-lobed (8–10 mm long). Stamens 4, didynamous, included in the throat, inserted at the base of the inflated part of the corolla, longer pair with filaments ca. 1.5 cm long, shorter pair with filaments ca. 1.2 cm long, all filaments with sparse glandular trichomes on the surface; anther thecae ca. 1–2 mm long, basal spur pointed, surface cover with simple trichomes. Pistil whitish-green; ovary ovoid, 1.5–2 mm long, apex pubescent; style ca. 3.2 cm long, pilose below, glabrous above; stigma subcapitate, minutely bilobed. Fruit not known.
Chroesthes faizaltahiriana Siti-Munirah A habit B a flower with bracteoles and calyx front side view C perianth lobes (flower from front view) D bract E bracteole F calyx G corolla (inner view) showing the stamens and pistil H pistil I cross-section of the ovary J stigma K1 anthers front view K2 anthers side view. All from FRI 91215, drawn by Mohamad Aidil Noordin).
Chroesthes faizaltahiriana is found in upper hill dipterocarp forests under shade at 822 m elevation. It was found flowering in February in patches of unlogged forest beside a logging road (Fig.
Chroesthes faizaltahiriana is dedicated to Mohd Faizal Mat Tahir (known as Faizal Tahir), the husband of Siti-Munirah for his strong support in many ways towards the author’s botanical work.
Critically Endangered B2 ab(ii,iii). Following the IUCN Red List Categories and Criteria (
Based on the general morphology of this plant, C. faizaltahiriana is close to C. longifolia, which was previously the only known species of Chroesthes in Peninsular Malaysia. However, a detailed comparison has shown that its inflorescence type is entirely different (Fig.
Chroesthes lanceolata (T. Anderson) B. Hansen, Nordic J. Bot. 3: 209. 1983; Hu, J.C., Deng, Y.F., Daniel, T. & Wood, J.R.I. Acanthaceae. In: Wu, Z.Y., Revan, P. & Hong, D.Y. (Eds.) Flora of China 19: 472. 2011. Basionym: Asystasia lanceolata T. And., J. Linn. Soc., Bot. 9: 524. 1867. Type: Myanmar, Pegu, Thaungyin, Brandis s.n. (holotype CAL).
Shrubs 0.5–3 m tall, anisophyllous. Stems terete, slender, rarely branched, glabrous. Leaves petiole 1–2.5 cm; leaf blade elliptic to oblanceolate to lanceolate, 10–16 × 3–7 cm, both surfaces glabrous, secondary veins 6–9 on each side of mid-vein, base cuneate, margin entire or sub-sinuate, apex acuminate. Inflorescence thyrses 3–7 cm; cymes sessile, 1–3-flowered; peduncles ca.2 cm; bracts elliptic to broadly lanceolate, 3–9 × 1–3 mm, apex acute, glandular-pubescent; bracteoles narrowly elliptic to broadly lanceolate, 4–9 × 0.7–1.2 mm; pedicel 1–5 mm long. Calyx 1–1.6 cm, outside glandular-pubescent; posterior lobe lanceolate, ovate or subelliptic; anterior lobes connate to two-thirds of their length. lateral lobes linear-lanceolate. Corolla white with pink or purple spots, ca. 2.5 cm, outside pubescent; tube basal portion ca. 9 mm, throat ca. 1.5 cm; upper lip two-lobed; lower lip three-lobed. Stamens 4, included in throat; filaments 1–1.2 cm, glabrous; anther thecae 2.1–2.3 mm, pubescent at the apex and along sides, basal spur pointed; ovary apex pubescent; style ca. 2.5 cm. Capsule subellipsoid to obovoid, 1.2–1.6 cm, glabrous or only at apex pubescent, four-seeded. Seeds subcircular in outline.
Myanmar, N Thailand, N Laos, N Vietnam, SW China (Yunnan), Malaysia. In Peninsular Malaysia, recorded from one specimen collected from the trail to Gunung Padang, Ulu Brang, Terengganu in 2010 (FRI 66129) (Map
In Peninsular Malaysia, found in a lowland dipterocarp forest at 473 m a.s.l., under a canopy near a small river. (Trail to Gunung Padang).
Critically Endangered B2 ab(ii). Following the IUCN Red List Categories and Criteria (
Chroesthes longifolia (Wight) B. Hansen Nordic J. Bot. 3: 210. 1983. Basionym: Lepidagathis longifolia Wight, Icones Plantarum Indiae Orientalis 4 (4): 8–9, Pl: t.1564. 1850; Ridley, The Flora of the Malay Peninsula 2: 587. 1923. Type: Malaysia, Malacca, Griffith s.n. (lectotype K).
Shrub 1–2.5 m high. Stem branches terete, glabrous. Leaves with petiole 0.5–2 cm long; lamina lanceolate, 16–24 × 3.4–8 cm, glabrous, lateral nerves up to 14 pairs, base attenuate, margin entire, apex acuminate. Inflorescence terminal raceme branching, up to 25 cm long; flowers secund, peduncles up to ca. 2.5 cm; one bract at each node being sterile, the other bract subtending one flower; bracts lanceolate, 15–20 × 4.5–6 mm, base obtuse, apex acuminate, glandular-pubescent, nerves conspicuous; bracteoles lanceolate, 10 × 3 mm, as bracts; pedicel very short. Calyx 1.6–1.8 cm long, posterior and anterior lobes elliptic-lanceolate, acuminate, conspicuously nerved, one posterior lobe, elliptic-lanceolate, 18 × 8–9 mm, 3–5 nerved/conspicuously nerved, apex acuminate; two anterior lobes, elliptic-lanceolate, 16 × 5–6 mm, 1-nerved/conspicuously nerved, apex acute; two lateral lobes, linear-lanceolate, 10–12 × 0.5 mm, one-nerved; all glandular-pubescent on both sides; lateral segments linear, one-nerved, glandular-pubescent on both sides; calyx greenish to purplish on the upper part or at the apex. Corolla bilabiate, dark purple-maroon (or claret), 2–3 cm long; outer surface dark purple, dark maroon, claret, sometimes turning white, glandular-pubescent outside; inner surface, dark purple, dark maroon, claret, sometimes whitish on nerves and base, glabrous; narrow part of tube 0.6 cm long, inflated part 1 cm; upper lip shortly emarginate, ca. 3–4 mm long, lower lip deeply trifid, ca. 5 mm long. Stamens 4, inserted at the base of the inflated part of the corolla; longer pair filament 1 cm, shorter pair filament 0.8 cm, filaments 0.8–1 cm long, glabrous, sometimes with glandular trichomes; anthers thecae ca. 1 mm long, glandular-pubescent along the back, bicalcarate at the base. Pistil whitish-green, stigma capitate; ovary glabrous; style ca. 1.4 cm long, pubescent. Capsule ca. 1.5 cm long, glabrous.
In primary lowland forest up to upper hill dipterocarp forest at 60 to 700 m a.s.l. Sometimes also found in logged and disturbed forests.
Least concern (LC). This is a widespread species throughout Peninsular Malaysia. However, many old collections are from places that have already changed its habitat. Development of these areas has led to population declines. However, its occurrence still has a wide range of distribution and many are still in totally protected areas.
Peninsular Malaysia. Johor: Batu Pahat: 6 November 1892, Lake HW s.n. (
Chroesthes longifolia (Wight) B. Hansen, collected from Taman Negara, Pahang (Kuala Keniam) A habit B–D inflorescence B front view C side view D back view E bract F–I flower with bracteoles and calyx F from the top view G lower view H–I side view J corolla (inner view) showing the stamens and pistil K corolla lips (flower from front view) showing anthers (All photos by Siti-Munirah MY).
This research was conducted as part of the Flora of Peninsular Malaysia Project (FPM) at the Forest Research Institute Malaysia, Kepong, funded by the Ministry of Energy and Natural Resources of Malaysia under the 11th Malaysian Plans (SPPII No. P23085100018003). I should like to thank the State Forestry Department of Kelantan, Malaysia for permission to make botanical collections at Berangkat FR, to the Department of Wildlife and National Parks (PERHILITAN) and Universiti Teknologi Mara (UiTM) for organising the scientific expedition to Taman Negara (Kuala Keniam) and to Serena Lee for providing images of the