Research Article |
Corresponding author: Chatchai Ngernsaengsaruay ( fsciccn@ku.ac.th ) Academic editor: Hanno Schaefer
© 2024 Chatchai Ngernsaengsaruay, Pichet Chanton.
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:
Ngernsaengsaruay C, Chanton P (2024) A taxonomic revision of Garcinia sections Dicrananthera and Macrostigma (Clusiaceae) in Thailand. PhytoKeys 248: 339-360. https://doi.org/10.3897/phytokeys.248.130311
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Garcinia sections Dicrananthera and Macrostigma (Clusiaceae) is revised for Thailand. Three species are enumerated, i.e., one species, G. thorelii belongs to the section Dicrananthera, and two species, G. nuntasaenii and G. prainiana, are in the section Macrostigma. Detailed morphological descriptions and illustrations are provided, together with notes on distribution, habitats and ecology, phenology, conservation assessments, etymology, vernacular names, uses, and specimens examined. An identification key to the species of section Macrostigma is presented. One name, G. thorelii, is lectotypified here. For Garcinia nuntasaenii we recommend a conservation status of Endangered [EN B2ab(v)] and for the other two species (G. prainiana and G. thorelii) a conservation status of Least concern [LC]. A number of vegetative characters and features of reproductive organs, especially the flowers, distinguish the two sections and we enumerate these herein.
Dioecy, Discostigma, edible fruits, Guttiferae, lectotypification, Malpighiales, taxonomy
Garcinia L. is a group of evergreen trees, occasionally shrubs, which are usually dioecious but sometimes polygamo-dioecious. It also has obligately and facultatively agamospermous species. The color of latex secreted from cut boles, twigs, leaves, and fruits can be yellow, pale yellow, white, cream, or clear. The genus consists of c. 400 species (
The last monograph of the genus Garcinia was published by
Garcinia section Macrostigma includes largely species that were included in
A taxonomic revision of Garcinia in Thailand has recently been undertaken by the first author as part of the Flora of Thailand project. More recently,
From these publications, the genus has a total of c. 30 accepted species in Thailand. However, identifications mostly rely on the literature and the type specimens, and this is the case for Garcinia sections Dicrananthera and Macrostigma, which have never been revised for Thailand. Therefore, in this paper, we provide here an updated account for these two sections in Thailand in order to present a taxonomic treatment that includes lectotypification, detailed morphological descriptions, and illustrations, along with notes on distributions, habitats and ecology, phenology, conservation assessments, etymology, vernacular names, uses, and specimens examined. An identification key to the species of the section Macrostigma is presented.
The collected specimens were examined by consulting taxonomic literature (e.g.,
Sectional classification sensu
Garcinia thorelii Pierre, Fl. Forest. Cochinch. 1(4): t. 62. 1882.
Garcinia section Dicrananthera is distinguished by the presence of a pair of interpetiolar stipuliform structures (rare in Clusiaceae); leaves with prominent, widely spaced secondary veins and scalariform tertiary veins; axillary or terminal cymose inflorescences with three to many flowers [i.e., G. yaatapsap (3–5-flowered), G. paucinervis (4–10-flowered), G. stipulata (4–6-flowered), G. nujiangensis (6–10-flowered), and G. thorelii (20–40-flowered thyrse)]; flowers with 4 sepals and 4 petals; male flowers with numerous stamens united into an annular mass encircling and attached to the pistillode (i.e., G. stipulata and G. yaatapsap) or united into 4 bundles surrounding the pistillode (i.e., G. nujiangensis, G. paucinervis, and G. thorelii), 2-thecous anthers; 1–2-locular ovaries; unlobed and smooth stigmas; and fruits with a smooth surface and unlobed. Distinguishing morphological characters reported here for this section were taken from
A section of five species worldwide (
Laos • Paklai, fl. & fr., Expedition Mekong 1866–1868, C. Thorel 3365 (lectotype designated here P [P04701082!]; isolectotypes [P04701076!, P04701080!, P04701081!, P04701083!], K [K000677688!].
Garcinia thorelii A branchlets, leaves, and male inflorescences B branchlets and male inflorescences with male flower buds and fully open male flowers C male inflorescences with male flower buds and early stage of open male flower D early stage of open male flower E fully open male flower showing 4 petals and numerous stamens united into 4 bundles surrounding a pistillode F infructescence G fruit. Photo: Drawn by Wanwisa Bhuchaisri.
Habit evergreen tree, 8–20 m tall, 50–180 cm GBH; latex yellow, sticky; branches decussate, horizontal or nearly horizontal; branchlets green, 4-angular, glabrous. Bark reddish brown or brown, cracked or shallowly fissured; inner bark pale brown. Terminal bud concealed between the bases of the uppermost pair of petioles. Stipuliform structures 2, interpetiolar, caducous, triangular, 1.2–2.7 × 1–2.8 mm, apex acute. Leaves decussate; lamina elliptic, narrowly elliptic or oblanceolate-obovate, 7.5–18 × 3–7.5 cm, apex acuminate or acute, base cuneate or obtuse, margin repand, coriaceous, dark green above, paler below, glabrous on both surfaces, midrib shallowly grooved or flattened above, raised below, secondary veins 5–10 each side, curving towards the margin and connected in distinct loops and united into an intramarginal vein, flattened above, slightly raised below, intersecondary veins usually absent, tertiary veins scalariform, veinlets reticulate, visible below, interrupted long wavy lines (glandular wavy lines, also called exudate containing canals) present, of differing lengths, running across the secondary veins to the apex, faint; petiole green, 0.6–1.5 cm long, grooved above, slightly transversely rugose, glabrous, with a basal appendage clasping the branchlet. Inflorescences axillary or at leafless nodes (in axils of fallen leaves), a short thyrse of many flowers, 2–2.5 cm long; bracts caducous, triangular, 0.6–2 mm long, apex acute; peduncle 5–8 mm long, 4-angular; rachis 0.8–2 cm long, 4-angular. Flowers unisexual, plants dioecious, 4-merous, fully open flowers 0.5–1.5 cm in diam.; bracteoles caducous, triangular, 0.8–1.5 mm long, apex acute; pedicels 2–4 mm long; sepals 4 and petals 4, decussate, glabrous; sepals pale green, not concave, triangular, 1–2.5 × 1–2.5 mm, subequal, apex obtuse; petals pale yellow or creamish white, concave, suborbicular, obovate or broadly elliptic, 3–5.5 × 2–4.5 mm, subequal, apex rounded, margin irregularly dentate. Male flowers: stamens numerous united into 4 bundles surrounding a pistillode, bundles 1.3–2.8 × 0.6–2 mm; filaments very short; anthers small; pistillode fungiform (mushroom-shaped), 2–3 mm long; rudimentary ovary slender, cylindrical, 0.5–1.5 mm long; sterile stigma pale yellow, sessile, convex, unlobed, 0.8–1.5 mm in diam., papillate. Female flowers: staminodes absent; pistil fungiform, 2.5–3 mm long; ovary ellipsoid, c. 1.5 mm long, glabrous, 1–2-locular; stigma convex, unlobed, 1.5–2 mm in diam., smooth. Fruits berries, ellipsoid or broadly ellipsoid, 1.5–2.5 × 1–2 cm, green, smooth, glabrous, with persistent sepals; persistent stigma convex, unlobed, 2.5–4 mm in diam., smooth; fruiting stalks 2–5 mm long, glabrous. Seeds 1, ellipsoid, 1.5–2 × 1–1.2 cm, with a thin fleshy pulp.
Garcinia thorelii A habit and stem B slashed bark with yellow latex C, D branchlets, leaves, and male inflorescences with male flower buds and open male flowers E interpetiolar stipuliform structure F branchlets and male inflorescences with male flower buds and early stage of open male flower G branchlets and male inflorescences with male flower bud and fully open male flowers showing 4 petals and numerous stamens united into 4 bundles surrounding a pistillode H branchlets, leaves, and fruit. Photos: Chatchai Ngernsaengsaruay (A–G), Yotsawate Sirichamorn (H).
Types of Garcinia thorelii, C. Thorel 3365 from Paklai, Laos, Expedition Mekong 1866–1868 A lectotype designated here P [P04701082!] B isolectotype P [P04701076!]. Photos: Muséum National d’Histoire Naturelle, Paris, France, http://coldb.mnhn.fr/catalognumber/mnhn/p/p04701082, http://coldb.mnhn.fr/catalognumber/mnhn/p/p04701076.
Northern: Nan, Lamphun, Lampang, Phrae, Phitsanulok; North-Eastern: Phetchabun, Loei, Bueng Kan, Nakhon Phanom; Eastern: Chaiyaphum, Nakhon Ratchasima; South-Western: Kanchanaburi, Phetchaburi (Fig.
It is found in dry evergreen forests, lower montane rain forests, mixed deciduous forests, often along streams, sometimes on limestone hills, 50–1,300 m amsl.
Flowering and fruiting more than once, nearly throughout the year.
Garcinia thorelii is distributed from Vietnam to Laos and Thailand. It is known from many localities and has a large EOO of 338,548.61 km2 and a small AOO of 128 km2. In Thailand, this species is known to be naturally distributed in four floristic regions and 13 provinces and has an EOO of 145,632.54 km2 and an AOO of 96 km2. Because of its number of localities and because it does not face any threat of extinction we therefore suggest the conservation status Least Concern (LC).
The specific epithet of Garcinia thorelii honors Clovis Thorel (1833–1911), a French physician, botanist, and plant collector (
Kok (ก๊อก) (Phitsanulok, from the specimen A. F. G. Kerr 5816); Khrak khamin (ครากขมิ้น), Ma da khi non (มะดะขี้หนอน) (Northern); Mai khwak lueang (ไม้ขวากเหลือง) [Phrae, from the specimen Unknown s.n. (BKF40333)]; Khi non (ขี้หนอน) (Laos, from the specimen J. E. Vidal 5952).
The wood is used for house construction [from the specimen Unknown s.n. (BKF40333)].
Garcinia thorelii was named by
According to
According to
According to
Thailand. Northern • Nan [Nam Haeng, young fr., 14 Jul 1926, Winit 1751 (BKF, K [K003964565])] • Lamphun [along the trail to Tat Moei Waterfall, Doi Khun Tan National Park, Mae Tha District, fl., 29 Jun 1993 (as Garcinia hanburyi), J. F. Maxwell 93-728 (BKF, CMUB, L [L2403744, L2403745]) • Doi Khun Tan National Park, fl., 29 Apr 1994 (as G. hanburyi), J. F. Maxwell 94-562 (BKF, CMUB)] • Lampang [Mae Saloi, fl., 29 Oct 1925, Winit 1492 (BKF, K [K003964566]) • Chae Son National Park, Wang Nuea District, ♂ fl., 2 Jun 1996 (as G. hanburyi), J. F. Maxwell 96-792 (BKF, CMUB) • ibid., sterile, 3 Jun 2014 (as G. hanburyi), T. Riythiwigrom 1 (CMUB)] • Phrae [Locality unspecified, fl., 1912, Luang Vanpruk 279 (BKF, K [K003964569]) • Locality unspecified, fl., s.d., Luang Vanpruk 514 (E [E00839797], K [K003964567]) • Huai Ta, Mae Song, fl., 28 May 1912, Luang Vanpruk 303 (BKF, K [K003964568]) • Long District, fl., 12 Mar 1930, Winit 1909 (BKF, K [K003964570]) • Thung Laeng Subdistrict, Long District, fr., 4 Feb 1963 (Garcinia sp.), Unknown s.n. (BKF40333) • Huai Rong Waterfall, fr., 19 Jun 1996 (as Garcinia sp.), R. Pooma & P. Puudjaa 1400-1 (BKF) • Phitsanulok [Nakhon Thai District, fl., 11 Apr 1922, A. F. G. Kerr 5816 (BM, E [E00839798], K [K003964572], P [P04701077]) • Huai Sai, fl., 18 Jun 1967 (as Garcinia sp.), S. Phusomsaeng 241 (BKF, K [K003964575], P [P05062032]) • Political and Military School, Phu Hin Rong Kla National Park, Nakhon Thai District, ♂ fl., 30 Sep 2007, C. Ngernsaengsaruay G56-30092007 (BKF, spirit material)] • Province unspecified [Pa Hia, near Pang Pue, fl., 28 Mar 1914, A. F. G. Kerr 3174 (BM, E [E00160901], K [K003964571])]; North-Eastern • Phetchabun [Khao Paya Paw, fr., 4 Mar 1931, A. F. G. Kerr 20342 (BKF, BM, C, E [E00160902], K) • Nam Nao National Park, fr., 8 Apr 1976 (Garcinia sp.), Bunnak 3108 (BKF) • Wang Pong District, fr., 1 Feb 2001, T. Wongprasert 012-3 (BKF, L [L3811042]) • Huai Phrom Laeng, Nam Nao National Park, fl., 18 Jan 2003 (as Garcinia sp.), P. Chantaranothai et al. 512003 (AAU) • along stream, Nam Nao National Park, fl., 28 May 2013 (as Garcinia sp.), C. Maknoi 5589 (QBG)] • Loei [Phu Luang Wildlife Sanctuary, sterile, 17 May 1998 (as Garcinia sp.), K. Chayamarit et al. 1480 (BKF) • Huai Baeng Forest Protection Station, Phu Luang Wildlife Sanctuary, Wang Saphung District, fr., 22 Jun 2003, T. Wongprasert 036-47 (BKF)] • Bueng Kan [Bueng Khong Long, Kinnari Waterfall, Phu Langka National Park, fr., 20 May 2014, S. Sirimongkol et al. 596 (BKF, K [K003964580], L [L4367330])] • Nakhon Phanom [Tat Kham Waterfall, Phu Langka National Park, fr., 30 Oct 1998, T. Wongprasert s.n. (BKF120854)]; Eastern • Chaiyaphum [Nam Phrom, fr., 10 Dec 1971 (as Garcinia sp.), C. F. van Beusekom et al. 4085 (BKF, C, K [K003964578], P [P05062035]) • Ban Nam Phrom, fr., 24 May 1974 (as Garcinia sp.), R. Geesink et al. 6910 (BKF, K [K003964574], L [L2409515]) • Phu Khiao, young fr., 3 Aug 1972 (Garcinia sp.), K. Larsen et al. 31312 (BK, K [K003964576], L [L2409542]) • Phu Khiao Wildlife Sanctuary, fr., 8 Nov 1984 (as Garcinia sp.), G. Murata T-50251 (BKF) • ibid., fr., 3 May 1997 (as Garcinia sp.), R. Pooma 1539 (BKF, CMUB) • Tabo–Huai Yai Wildlife Sanctuary, fr., 22 Aug 2019 (as Garcinia sp.), N. Boonruang 0350 (QBG)] • Nakhon Ratchasima [Koa Lem, fl. & fr., 12 Jan 1925, A. F. G. Kerr 9978 (BM, E [E00839799], K [K003964573], P [P04701078]), 9979 (BM, P [P04701079]) • Khao Yai National Park, fl. & fr., 9 Apr 1974, T. Smitinand 12001 (BKF)]; South-Western • Kanchanaburi [Erawan Waterfall, fl., 25 Jan 1962 (as Garcinia sp.), K. Larsen & T. Smitinand 9271 (BKF, C, K [K003964577]) • Sai Yok, fr., 26 Nov 1971 (as Garcinia sp.), C. F. van Beusekom et al. 3987 (BK, C, K [K003964579], P [P05062045]) • Vajiralongkorn Dam, Tha Khanun Subdistrict, Thong Pha Phum District, fl., 11 Jan 1985 (as Garcinia sp.), H. Koyama T-49019 (BKF)] • Phetchaburi [Mae Kradang La Waterfall, Kaeng Krachan National Park, Nong Ya Plong District, female fl. & fr., Feb 2024, Y. Sirichamorn personal observation with photos].
Vietnam • Tonkin [prov. de Lauyson, fr., 19 Dec 1913 (as Garcinia sp.), Unknown 29663 (P [P04788166]) • Hoa-Binh, fr., 1 Nov 1929 (as G. bonii), M. Brillet unreadable number (P [P04899809, P04899810])].
Laos • Luang Phrabang [Locality unspecified, fl., 20 Mar 1932 (as Garcinia sp.), Poilane 20452 (P [P04899801])] • Sayaboury [Mekong River, Na Konken Village, ♂ fl., 28 Apr 2012, J. F. Maxwell 12-154 (CMUB, L [L4345648])] • Vientiane [Reservoir Nam Ngum, fr., 19 Oct 1974 (Garcinia sp.), J. E. Vidal 5952 (P [P05061722, P05061726])].
Garcinia latissima Miq., Ann. Mus. Bot. Lugduno-Batavi 1: 209. 1864.
Garcinia section Macrostigma is characterized by its axillary or terminal cymose inflorescences with two to many flowers (in a simple cyme or in fascicles of several simple cymes), or a solitary flower (in female flowers); 4(–5)-merous flowers [4-merous, e.g., G. nuntasaenii; 5-merous, e.g., G. phuongmaiensis, G. prainiana]; male flowers with numerous stamens united into a central column, sometimes lobed with lobes equaling number of petals [a single weakly 4-lobed bundle, e.g., G. nuntasaenii], or into completely separate antepetalous bundles [united into 5 bundles, e.g., G. phuongmaiensis, G. prainiana]; androecium often adnate to the petals to varying degrees; 2-thecous anthers; pistillode absent in male flowers, but rudimentary or well-developed pistillode present in some species; (–3)4–8-locular ovaries; unlobed or lobed and smooth or papillate stigmas; and fruits with a smooth surface, unlobed or faintly, shallowly or deeply lobed. Distinguishing morphological characters of this section based on
A section of 29 species worldwide (
A related species from Vietnam is included in the key.
1 | Shrub; leaves smaller (4–17 × 2.5–7.5 cm); flowers smaller (0.8–1 cm in diam.); petals pale yellow, creamish white or white; fruits lobed, turning red or bright red when ripe; distributed in Central Vietnam, Central Laos, and North-Eastern Thailand | 2 |
– | Tree; leaves larger (12.5–27.5 × 5.5–11.5 cm); flowers larger (2.5–4 cm in diam.); petals variable in color: pale yellow, yellowish pink, yellowish red, pinkish red, pink or red; fruits unlobed, turning bright yellow, orangish yellow and bright orange when ripe; distributed in Peninsular Malaysia and Peninsular Thailand | Garcinia prainiana (2.) |
2 | Flowers 4-merous; petals pale yellow or creamish white; fruits 4–6-lobed; distributed in Central Laos and North-Eastern Thailand | Garcinia nuntasaenii (1.) |
– | Flowers (4–)5-merous; petals white; fruits 3–4-lobed; distributed in Central Vietnam | Garcinia phuongmaiensis |
Thailand • Bueng Kan Province, Bung Khla District, Phu Wua Wildlife Sanctuary, fr., 13 Dec 2008, N. Nuntasaen 10 (holotype BKF!; isotype BKF!).
Garcinia nuntasaenii A habit and habitat B branchlets and young leaves C terminal bud concealed between the bases of the uppermost pair of petioles D terminal inflorescences with flower buds E branchlets and male inflorescences in axils of fallen leaves with male flower buds and open male flowers F female flower (top view) G branchlets, leaves, and fruit. Photos: Chatchai Ngernsaengsaruay (A–D, F), Manop Poopath (E, G).
Habit
shrub, 1–2 m tall; latex white, turning pale yellow, sticky; branches decussate, horizontal or nearly horizontal; branchlets green, 4-angular, glabrous. Bark green when young, turning dark brown when mature, smooth, usually lenticellate; inner bark pale yellow. Terminal bud concealed between the bases of the uppermost pair of petioles. Leaves decussate; lamina lanceolate-ovate, ovate or elliptic, 6–17 × 3–7.5 cm, apex acuminate or acute and rigid, base subcordate, margin thick, entire and slightly undulate (repand), coriaceous, slightly bullate or bullate, apical part of leaves conduplicate, twisted and recurved, glossy dark green above, paler below, glabrous on both surfaces, midrib shallowly grooved above, raised as a prominent ridge below, secondary veins 12–20 each side, curving towards the margin and connected in distinct loops and united into an intramarginal vein, flattened above, slightly raised below, conspicuous on both surfaces, intersecondary veins conspicuous, veinlets reticulate, visible on both surfaces, interrupted long wavy lines present, of differing lengths, nearly parallel to the midrib, running across the secondary veins to the apex, visible below; petiole green, 0.2–1 cm long, not grooved, transversely rugose, glabrous, with a basal appendage clasping the branchlet; young leaves red. Inflorescences terminal or at leafless nodes (in axils of fallen leaves), cymose. Flowers unisexual, plants dioecious, 4-merous, fully open flowers 0.8–1 cm in diam.; bracteoles narrowly triangular or triangular, 2–10 × 1.5–2.5 mm, apex acute or acuminate, somewhat thick; pedicels short; sepals 4 and petals 4, decussate, pale yellow or creamish white, somewhat thick; sepals obovate or elliptic, 3–6 × 2.5–3.5 mm, concave, apex rounded; petals broadly obovate, 5.5–8.5 × 3.5–7 mm, apex rounded, apical part recurved. Flower buds: sepals pink or pink-pale yellow. Male flowers in fascicles of several simple cymes; stamens numerous, united into a single weakly 4-lobed bundle; filaments very short; anthers 0.3–0.5 mm long; pistillode usually absent. Female flowers usually in a cluster of 3 flowers (a simple cyme); staminodes numerous, united into a single weakly 4-lobed bundle, surrounding the ovary; pistil fungiform; ovary depressed globose, 1.5–2 × 2–3 mm, shallowly 4–6-lobed, 4–6-locular; stigma convex, hemispherical, 1.5–2 × 2.5–3.5 mm, papillate. Fruits berries, depressed globose, 0.5–0.7 × 1–2 cm, 4–6-lobed, green with white dots, turning red when ripe, smooth, glabrous, glossy, with persistent sepals; persistent stigma circular, flat, radiately lobed or unlobed; fruiting stalk 2–4 mm. long. Seeds 4–6, with fleshy pulp. The description of this species is based on
Central Laos and North-Eastern Thailand.
North-Eastern: Bueng Kan, Nakhon Phanom.
It is found in dry evergreen forests, 150–220 m amsl.
Flowering and fruiting more than once; flowering in December to July; fruiting in December to April.
Garcinia nuntasaenii is known only from four localities in Bueng Kan and Nakhon Phanom Provinces, North-Eastern Thailand and Nam Kading National Protected Area, Laos. All occurrences are in small populations. It has a small EOO of 796.87 km2 and a relatively small AOO of 16 km2. Because of its number of localities, and because the roots of this species are used for medicinal purpose by nearby villagers in Thailand, the population is suspected to be declining. We therefore suggest the conservation status Endangered [EN B2ab(v)].
The specific epithet of Garcinia nuntasaenii honours Mr Narong Nuntasaen, a staff member and a plant collector of BKF, who collected the type specimen.
Chang nga ek (ช้างงาเอก) (Bueng Kan, Nakhon Phanom).
The fruits are edible (from the specimen M. Norsaengsri & N. Tathana 8630). The roots are locally used for medicinal purposes (Ngernsaengsaruay personal observation).
Garcinia nuntasaenii is similar to G. phuongmaiensis in its habit (shrubs); sticky white latex, turning pale yellow when exposed to the air; 4-angular branchlets, especially when young; coriaceous, bullate, shiny dark green, subcordate leaves with a short petiole; flower size, c. 1 cm in diam.; and the color of fruits, turning red when ripe, but differs in having 4-merous flowers [vs (4–)5-merous flowers]; pale yellow or creamish white petals (vs white petals); 4–6-lobed fruits (vs 3–4-lobed fruits); and is also distributed in Central Laos and North-Eastern Thailand (vs distributed in Central Vietnam). The morphological characteristics and distribution of G. phuongmaiensis were taken from
Thailand. North-Eastern • Bueng Kan [Phu Wua Wildlife Sanctuary, Bung Khla District, fl., 8 Nov 1996 (Garcinia sp.), C. Niyomdham 4910 (BKF) • ibid., fl., 27 Aug. 2001 (Garcinia sp.), R. Pooma et al. 2791 (BKF) • ibid., ♂ fl., 1 Jan 2008 (spirit specimen), N. Nuntasaen 11 (BKF) • ibid., fr., 27 Dec 2011 (Garcinia sp.), M. Norsaengsri & N. Tathana 8630 (BKF, QBG) • ibid., 12 Feb 2015, M. Poopath 981-1, 981-2 (BKF) • ibid., ♀ fl., 24 Jul 2015, C. Ngernsaengsaruay & N. Meeprom 754 (BKF) • The Upper Northeast Wild Plants Conservation Center, Bung Khla District, ♂ fl., 24 Jul 2015, C. Ngernsaengsaruay & N. Meeprom 750, 751 (BKF) • ibid., ♀ fl., 24 Jul 2015, C. Ngernsaengsaruay & N. Meeprom 752, 753 (BKF)] • Nakhon Phanom [Ban Phaeng District, Phu Langka National Park (Narong Nuntasaen own observation)].
Laos • Bolikhamxay [Nam Kading National Protected Area, ♂ fl., L8 • ibid., fl. & fr., L431 • ibid., L1034 (
Peninsular Malaysia • Perak, Kuala Dipang (originally ‘‘Kwala Dipang’’ on the label; originally published ‘‘Kwala Dynong’’), fl., February 1885, B. Scortechini 1796 (lectotype designated by
Garcinia prainiana A, B branchlets, young leaves, and mature leaves C inflorescences with male flower buds and open male flower D branches and branchlets with mature and ripe fruits E branchlets, leaves, and mature and ripe fruits F ripe fruits (pericarp removed and in transverse section showing seeds with fleshy pulp). Photos: Chatchai Ngernsaengsaruay (A–C, E, F), G Rawit Sichaikhan (D).
Habit
evergreen trees, 3–12 m tall, 15–75 cm GBH; latex white, sticky; branches decussate, horizontal or nearly horizontal; branchlets 4-ridged, glabrous. Bark pale brown, grayish brown or blackish brown, smooth or slightly rough; inner bark pale yellow. Terminal bud concealed between the bases of the uppermost pair of petioles. Leaves decussate; lamina elliptic, oblong or elliptic-oblong, sometimes narrowly oblong, 12.5–27.5 × 5.5–11.5 cm, apex acute or obtuse, base subcordate, often subamplexicaul, margin repand and slightly revolute, coriaceous, bullate or slightly bullate, shiny dark green above, paler below, glabrous on both surfaces, midrib flattened above, raised as a prominent ridge below, secondary veins 9–20 each side, curving towards the margin and connected in distinct loops and united into an intramarginal vein, flattened above, raised below, conspicuous on both surfaces, intersecondary veins conspicuous, veinlets reticulate, visible on both surfaces, interrupted long wavy lines present, of differing lengths, nearly parallel to the midrib, running across the secondary veins to the apex, visible below; petiole green, 1.5–6 mm long, 2–5 mm in diam., not grooved, transversely rugose, glabrous, with a basal appendage clasping the branchlet; young leaves shiny pale green. Inflorescences terminal, sometimes on short, leafless lateral branchlets, cymose, usually in dense fascicles of several to many flowers. Flowers unisexual, plants dioecious, 5-merous, fully open flowers with spreading petals; bracteolate; sepals 5 and petals 5, quincuncial, coriaceous, glabrous. Male flowers 2.5–3.5 cm in diam.; bracteoles pale green, triangular 2.3–4.5 × 1.8–3.7 mm, apex acute, conduplicate with a central keel; pedicel pinkish green, reddish green or greenish red, 3–6 mm long, 2.5–4 mm in diam., widened at the apical part, terete, glabrous; sepals pinkish green, reddish green or greenish red, concave, broadly ovate or suborbicular 4.8–8 × 5–7.8 mm, unequal, apex rounded; petals variable in color: pale yellow, yellowish pink, yellowish red, pinkish red, pink or red, broadly obovate or obovate, 0.8–1.4 × 0.6–1.1 cm, subequal, sometimes unequal, apex rounded; stamens numerous, united into 5 bundles surrounding a pistillode, antepetalous, 1.7–4.2 mm long, each bundle 1.2–4 mm wide, pale yellow, pink or red; filaments fused throughout their entire length; anthers yellow, 0.3–0.6 mm long; pistillode fungiform, 5.5–7.5 mm long; sterile stigma pale yellow, pink or red, sessile, convex, indistinctly lobed, 5–6 mm in diam., papillate. Female flowers 2.5–4 cm in diam.; bracteoles and pedicel same as in male flowers; sepals and petals same as or slightly larger than in male flowers; staminodes absent; pistil fungiform, 6–8.5 mm long; ovary pale green, depressed globose 4–6 × 4.5–6.5 mm, unlobed, glabrous, 5–8-locular; stigma pale yellow, pink or red, sessile, convex, weakly 5–8-lobed or indistinctly lobed, 5–7 mm in diam., papillate. Fruits berries, depressed globose or depressed subglobose, sometimes globose, 2–3.5 × 2–5.3 cm, sometimes oblique, asymmetrical, unlobed, slightly concave or flattened at the apex, green, turning greenish yellow, bright yellow, orangish yellow and bright orange when ripe, smooth, glabrous, glossy, then exocarp becoming dark brownish black and slightly sinuously wrinkled when dry, pericarp 3.5–8 mm thick, exocarp thin; persistent stigma dark brown or blackish brown, circular, button-like, 0.6–1.1 cm in diam., slightly concave or flattened, weakly 5–8-lobed or indistinctly lobed, papillate; persistent sepals pale green, turning yellowish green and orangish green, larger than in flowering material; fruiting stalk green, thick, 0.4–1.5 cm long, 3–6 mm in diam., Seeds 1–6, often aborted, brown, broadly ellipsoid, ellipsoid or subglobose, 0.9–1.6 × 0.7–1.4 cm, with pale orange fleshy pulp. The description of this species was taken from
Known only from Peninsular Thailand and Peninsular Malaysia. It is widely distributed in Peninsular Malaysia (Perlis, Kedah, Penang, Perak, Kelantan, Terengganu, Pahang, Selangor, Negeri Sembilan, Malacca, Johor) (
Peninsular: Yala (Than To), Narathiwat (Waeng, Su-ngai Kolok).
It is found in tropical lowland evergreen rain forests, occasionally along streams, 30–200 m amsl. It is also cultivated in villages and botanical gardens.
In Peninsular Malaysia, it occurs in lowland and hill forests, on hillsides and ridges up to elevations of 1,000 m amsl. It is also cultivated in villages (
Flowering and fruiting more than once; flowering nearly throughout the year, usually in February to May; fruiting April to June and September to December.
LC (
The specific epithet of Garcinia prainiana refers to Sir David Prain (1857–1944), a British botanist, a herbarium curator of the Royal Botanic Garden, Calcutta (1887–1898), and a director of the Royal Botanic Gardens, Kew (1905–1922) (
Chupu (จูปู) (Malay-Narathiwat); Cerapu, Chekau, Chepu, Cherapu, Cherpu, Cherupu, Chupak, Chupu, Kechupu, Kecupu, Menchepu, Menchupu (Malay); Button mangosteen (English).
Garcinia prainiana is locally cultivated for its fruits in peninsular Thailand. The fruits (pericarp and fleshy pulp surrounding the seeds) are edible and have a sour or sweet-sour taste. It is also grown in some botanical gardens as an ornamental plant to provide botanical education.
In Peninsular Malaysia, it is commonly cultivated in village gardens. The ripe fruits are edible and are sometimes used fresh in beverages (
According to
Garcinia prainiana is also similar to G. phuongmaiensis in having coriaceous, bullate, shiny dark green, subcordate, subamplexicaul leaves with a short petiole; 5-merous flowers; and numerous stamens, united into 5 antepetalous bundles surrounding a pistillode, but differs in relatively larger habit as a 3–12 m tall tree (vs smaller habit, shrubs, 1–3 m tall); larger leaves, 12.5–27.5 × 5.5–11.5 cm (vs smaller leaves, 4–11 × 2.5–5 cm); larger flowers, 2.5–4 cm in diam. (vs smaller flowers, c. 1 cm in diam.); variable in color of petals: pale yellow, yellowish pink, yellowish red, pinkish red, pink or red (vs white petals); staminodes absent (vs present); unlobed fruits, turning bright yellow, orangish yellow and bright orange when ripe (vs shallowly 3–4-lobed fruits, turning bright red when ripe); seeds with pale orange fleshy pulp (vs seeds with white fleshy pulp); and is distributed in Peninsular Malaysia and Peninsular Thailand (vs Central Vietnam). The morphological characteristics and distribution of G. phuongmaiensis were taken from
Thailand. Central • Nakhon Nayok [Phrueksaphan Thepparat Botanicical Garden, Chulachomklao Royal Military Academy, cultivated, 31 May 2019, C. Ngernsaengsaruay & W. Boonthasak G30-31052019 (BKF); Peninsular • Trang [Khao Chong Botanical Garden, Chong Subdistrict, Na Yong District, cultivated, 16 Feb 2022, C. Ngernsaengsaruay et al. G32-16022022 (BKF)] • Yala [Chulabhorn Phatthana 7 Project, Than To District, near waterfall, 27 Nov 2019, C. Ngernsaengsaruay & G. Sichaikhan G31-27112019 (BKF)] • Narathiwat [Hala-BalaWildlife Sanctuary, Ban Bala, Lo Chut Subdistrict, Waeng District, ♂ fl., 13 May 2005 (as Garcinia sp.), M. Poopath 274 (BKF) • Hala-Bala Wildlife Sanctuary, Waeng District, fr., 22 Sep 2005, C. Niyomdham & P. Puudjaa 7593 (BKF) • Su-ngai Kolok District, fr., 20 Apr 2002, U. Upho 556 (QBG) • Su-ngai Kolok District, fl., cultivated, 20 May 2003, U. Upho 550 (BKF)].
Peninsular Malaysia • Perak [Kwala Dipang, ♂ fl., Dec 1896, C. Curtis 3273 (K [K000677679]) • Kg Kepayang near Ipoh, fr., 30 Oct 1971, Syed Abu Bakar Kep. FRI20440 (L [L2417220])] • Pahang [Su-ngai Bertam at Kuala Mensum, fl., 2 Jun 1971, T. C. Whitmore Kep. FRI20091 (L [L2417222]) • Path leading to Kuala Mensum from Boh Tea, Cameron Highlands, fr. 24 Sep 1971, H. S. Loh Kep. FRI19187 (L [L2417221]) • Cameron Highlands Road, fr., 18 Jan 1982, K. M. Kochummen Kep. FRI29377 (L [L2417225]) • Kelantan [0.5 mile east of Gua Musang, fr., 14 Jul 1967, T. C. Whitmore Kep. FRI4018 (L [L2417226]) • Su-ngai Lebir, below Kuala Relai at Jentah, fl., 24 Apr 1976, B. C. Stone & M. Sidek 12426 (BKF, L [L2417224], US [US02961246]) • Su-ngai Long off Su-ngai Pergau, Jeli, fr., 26 Sep 1986, A. Latiff et al. ALM1856 (L [L3806490], PSU) • Ketam, Cicar Tinggi, Kampung Bata, Pasir Mas, ♀ fl. & fr., 1 Aug 1992, A. Noorsiha et al. Kep. FRI39214 (L [L3878683]) • Pasir Putih, fl., 23 Oct 1992, H. Husmady et al. Kep. FRI39551 (L [L3806959]) • near Brooke Camp, Gua Musang, fl., 2 Jun 1994, H. Husmady et al. Kep. FRI41841 (L [L2417223])].
We would like to thank the curators and staff of the following herbaria AAU, BK, BKF, BM, C, CMUB, K, P, PSU, and QBG for their assistance during visits and allowing access to the herbarium specimens, and those included in the digital herbarium databases of AAU, CAL, E, K (including K-W), L, P, and US. We are grateful to the plant collectors of Garcinia sections Dicrananthera and Macrostigma. We also would like to thank Wanwisa Bhuchaisri for the line drawings, Dr Nattanon Meeprom and Weereesa Boonthasak for their kind help with field work, Manop Poopath for two photos of G. nuntasaenii, G Rawit Sichaikhan for a photo of G. prainiana, and Asst. Prof. Dr Yotsawate Sirichamorn for a photo and information of G. thorelii from Kaeng Krachan National Park, Phetchaburi Province.
The authors have declared that no competing interests exist.
No ethical statement was reported.
This research was funded by the Basic Research Fund (BRF) and the International SciKU Branding (ISB), Faculty of Science, Kasetsart University.
Conceptualization: CN. Data curation: CN, PC. Formal analysis: CN. Funding acquisition: CN. Investigation: CN. Methodology: CN, PC. Project administration: CN. Resources: CN, PC. Writing - original draft: CN, PC. Writing - review and editing: CN.
Chatchai Ngernsaengsaruay https://orcid.org/0000-0002-7131-976X
Pichet Chanton https://orcid.org/0009-0001-7325-6109
All of the data that support the findings of this study are available in the main text.