A taxonomic revision of Garcinia section Xanthochymus (Clusiaceae) in Thailand

Taxonomic Treatment

Garcinia L. sect. Xanthochymus (Roxb.) Pierre, Fl. Forest. Cochinch. 1(5): 3. 1883; Vesque, Epharmosis 2: 14. 1889 et in A. DC. & C. DC., Monogr. Phan. 8: 254–255. 1893.—Xanthochymus Roxb., Pl. Coromandel 2(4): 51, t. 196. 1805. Type: Xanthochymus pictorius Roxb. = Garcinia xanthochymus Hook. f. ex T. Anderson.

Tree evergreen; latex usually white, sticky; branches decussate, horizontal; branchlets usually 4-ridged (except G. prainiana, terete), pubescent or glabrous. Leaves decussate, usually large and coriaceous, pubescent or glabrous; petiole usually transversely rugose. Inflorescences on short, leafless lateral branchlets (in axils of fallen leaves), sometimes terminal (e.g., G. prainiana), cymose, in fascicles of several to many flowers, pubescent or glabrous. Flowers unisexual, sometimes bisexual, plants dioecious, sometimes polygamo-dioecious (e.g., G. dulcis); bracteolate; sepals and petals quincuncial; sepals 5, unequal, margin ciliolate or eciliolate, pubescent or glabrous outside; petals 5, usually subequal, margin ciliolate or eciliolate. Male flowers: disk in the center of the flower, intrastaminal, 5-lobed, with lobes positioned between the stamen bundles and antesepalous (opposite sepals) (except G. prainiana, disk ring-shaped); stamens numerous, united into 5 bundles (phalanges), antepetalous (opposite petals); anthers very small; pistillode very small or absent. Female flowers: appendages (disk lobes) 5, antesepalous, alternating with staminode bundles; staminodes united into 5 bundles, antepetalous (except G. prainiana, appendages and staminodes absent); ovary 5-locular (except G. prainiana, 5–8-locular); stigma 5-lobed (except G. prainiana, weakly 5–8-lobed or indistinctly lobed). Fruit a fleshy berry, with sticky yellow latex, exocarp thin, usually sinuously wrinkled when dry.

A section of four species in Thailand; three species in Indo-China (Garcinia dulcis, G. phuongmaiensis, and G. xanthochymus).

A key to the species of Garcinia sect. Xanthochymus in Thailand

1. Garcinia dulcis (Roxb.) Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 43(2): 88. 1874 et Forest Fl. Burma 1: 92. 1877; Pierre, Fl. Forest. Cochinch. 1(5): 4. 1883; Vesque, Epharmosis 2: 14. t. 87. 1889 et in A. DC. & C. DC., Monogr. Phan. 8: 312. 1893; King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 59(2): 169. 1890; Koord. & Valeton, Bijdr. Boomsoort. Java 9: 359. 1903; Merr., Philipp. J. Sci. 3: 362. 1908; Ridl., Fl. Malay Penins. 1: 179. 1922; Merr., Enum. Philipp. Fl. Pl. 3: 84. 1923; Corner, Wayside Trees Mal. 1: 316. fig. 105. ed. 2. 1952; Backer & Bakh. f., Fl. Java 1: 386. 1963; Maheshw., Bull. Bot. Surv. India 6: 115. t. 1. fig. 4. 1964; Whitmore in Whitmore, Tree Fl. Malaya 2: 209. 1973; H. Keng, Concise Fl. Singapore: 48. 1990; E. W. M. Verheij & R. E. Coronel (eds), PROSEA 2: 175, 176. t. 176. 1992; N. P. Singh in B. D. Sharma & Sanjappa, Fl. Ind. 3: 109. 1993; S. Baruah et al., Ethnobot. Res. Appl. 21(33): 6. fig. 6. 2021; W. E. Cooper, Austrobaileya 9(1): 4. fig. 1A–1B. 2013; A. Begum, S. K. Barthakur & J. Sarma, Pleione 7(2): 546. t. 1. 2013; S. Gardner, P. Sidisunthorn & Chayam., Forest Trees S. Thailand 1: 352. fig. 541. 2015.—Xanthochymus dulcis Roxb. [Hort. Bengal.: 42. 1814, nom. nud.], Pl. Coromandel 3(3): 66. t. 270. 1820 et in Carey, Fl. Ind. 2: 631. 1832; Wight, Icon. Pl. Ind. Orient. 1(10): 10. t. 192. 1839.—Garcinia elliptica Choisy in DC., Prodr. 1: 561. 1824 [non Garcinia elliptica Wall., Numer. List. 4869. 1831, nom. nud.].—Xanthochymus javanensis Blume, Bijdr. Fl. Ned. Ind. 5: 216. 1825.—Stalagmitis dulcis Cambess., Mém. Mus. Hist. Nat. 16: 393, 426, 1828.—Stalagmitis elliptica G. Don, Gen. Hist. 1: 621. 1831.—Stalagmitis javanensis Spach, Hist. Nat. Vég. 5: 328. 1836. Type: Roxburgh’s illustration, Xanthochymus dulcis Roxb., Pl. Coromandel 3(3): 66. t. 270 (Roxburgh, 1820) (lectotype designated here)

— Garcinia vilersiana Pierre, Bull. Mens. Soc. Linn. Paris 1: 348. 1882 et Fl. Forest. Cochinch. 1(5): t. 71B–C. 1883; Vesque, Epharmosis 2: 14. t. 85. 1889 et in A. DC. & C. DC., Monogr. Phan. 8: 318. 1893; Engl. in Engl. & Prantl, Die Naturlichen Pflanzenfamilien 3(6): 234. fig. 109D–F. 1895; Pit. in Lecomte et al., Fl. Indo-Chine 1(4): 297. fig. 29: 6–8. 1910; Craib, Fl. Siam. 1(1): 118. 1925; Gagnep. in Gagnep., Fl. Indo-Chine Suppl.: 257. 1943; Pételot, Arch. Rech. Agron. Cambodge Laos Vietnam 1: 64. 1952; P. H. Hô, Câyco Vietnam 1: 568. fig. 1572. 1991. Type: Thailand, ad flumen Mekong (also spelled Khong River) in regno Siamico, June 1868, Pierre 3642 (lectotype selected here P [P04701111!]; isolectotypes P [P04701114!, P04701115!]), syn. nov. (Fig. 1).

—Garcinia cambodgiensis Vesque, Epharmosis 2: 14. t. 92, 93. 1889 et in A. DC. & C. DC., Monogr. Phan. 8: 316. 1893; Pit. in Lecomte et al., Fl. Indo-Chine 1(4): 298. 1910; Craib, Fl. Siam. 1(1): 114. 1925; Gagnep. in Gagnep., Fl. Indo-Chine Suppl.: 257. 1943.—Garcinia andersoni Pierre, Fl. Forest. Cochinch. 1(5): 1. t. 72. 1883. Type: Cambodia (originally “Cambodgiae” on the label), ad montes Kuang Repoeu, in prov. Tpong, May 1870, Pierre 775 (lectotype selected here P [P05062528!]; isolectotypes K [K000677687!], P [P05062544, P05062564]), syn. nov. (Fig. 2).

—Garcinia andamanica King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 59(2): 170. 1890; Vesque in A. DC. & C. DC., Monogr. Phan. 8: 328. 1893; Brühl & King, Ann. Roy. Bot. Gard. (Calcutta) 5(2): 141. t. 169. 1896; Brandis, Indian Trees: 49. 1906; C. E. Parkinson, Forest Fl. Andaman Isl.: 89. 1923; Maheshw., Bull. Bot. Surv. India 6: 112. 1964; N. P. Singh in B. D. Sharma & Sanjappa, Fl. Ind. 3: 103. 1993; Shameer & N. Mohanan, Rheedea 29(2): 181. 2019. Type: India, Andaman Islands, 1884, King’s Collector 224 (lectotype designated by Shameer & Mohanan (2019), CAL [CAL0000024974, photo seen], syn. nov. (Fig. 3).

—Garcinia andamanica King var. pubescens King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 59(2): 170. 1890; Maheshw., Bull. Bot. Surv. India 6: 112. 1964; N. P. Singh in B. D. Sharma & Sanjappa, Fl. Ind. 3: 104. 1993. Type: India, Andaman Islands, 1884, King’s Collector 136 (lectotype designated by Maheshwari (1964), CAL [CAL0000024973, photo seen]; isolectotypes K [K000677630!], P [P05062496!]), syn. nov. (Fig. 4).

Tree evergreen, 5–20(–25) m tall, 30–160 cm girth, sometimes buttressed near the base of the stem in large trees; latex white, turning creamish white, sticky; branches decussate, horizontal; branchlets green, 4-ridged, pubescent, slightly pubescent or glabrescent, sometimes glabrous. Bark smooth or slightly rough, dark brown or greyish brown; inner bark pale yellow. Terminal bud concealed between the bases of the uppermost pair of petioles. Leaves decussate; lamina variable in shape, lanceolate-ovate or lanceolate (broadest at the basal part, gradually narrowing towards the apex), sometimes narrowly oblong, oblong or elliptic, 12.5–32.5 × 4.5–16 cm, apex acute or acuminate, sometimes obtuse, base obtuse, cuneate or subcordate, margin repand and slightly revolute, thickly coriaceous, slightly bullate or bullate, shiny dark green above, paler below, glabrous above, pubescent, slightly pubescent or glabrescent, sometimes glabrous below (of lamina and veins), midrib and secondary veins flattened above, raised as a prominent ridge below, secondary veins 10–17 pairs, curving towards the margin connected in distinct loops and united into an intramarginal vein, conspicuous on both surfaces, with intersecondary veins, veinlets reticulate, visible on both surfaces, interrupted long wavy lines (glandular wavy lines, also called exudate containing canals) of differing lengths, running across the secondary veins to the apex or the margin, visible below; petiole green, 0.7–3.2 cm long, 2–6 mm in diam., not grooved, distinctly transversely rugose, indumentum same as in lamina, with a conspicuous basal appendage clasping the branchlet; young leaves shiny pale green; mature leaves turning greenish yellow to yellow before falling off. Inflorescences on short, leafless lateral branchlets (in axils of fallen leaves), cymose, in dense fascicles of 7–17 flowers. Flowers unisexual or bisexual, plants polygamo-dioecious, 5-merous, petals erect with overlapping edges and corolla forming a bowl shape; bracteolate; sepals and petals quincuncial, coriaceous, concave. Male flowers in fascicles of 11–17 flowers, 0.8–1.5 cm in diam.; bracteoles triangular, 0.7–2  × 0.6–2.7 mm, apex acute, pubescent; pedicel green, 0.5–1.7 cm long, 1–2 mm in diam., terete (circular in cross-section), pubescent or glabrescent, rarely glabrous; sepals 5, pale green, whitish pale green or green, broadly elliptic, semi-orbicular, suborbicular, 3–5.2 × 2.5–5 mm, unequal, apex rounded, margin ciliolate, pubescent or glabrescent, rarely glabrous outside; petals 5, whitish pale green, creamish white or pale yellow, broadly elliptic, elliptic or suborbicular, 5–8.5  × 4–8.5 mm, subequal, apex rounded, margin ciliolate; disk in the center of the flower, intrastaminal, yellow, 5-lobed, fleshy, pitted, lobes positioned between the stamen bundles, antesepalous; stamens numerous, united in 5 bundles, 6–13 in each bundle, antepetalous, 0.3–1 cm   × 0.5–1 mm each bundle, creamish white or pale yellow; filaments 0.5–0.8 mm long; anthers yellow, 2 thecae, 0.2–0.4  × 0.3–0.7 mm; pistillode small or absent. Female flowers in fascicles of 7–9 flowers, 0.8–1.6 cm in diam.; bracteoles same as in male flowers; pedicel pale green or green, 1.2–2 cm long, widened at the apical part, middle part 1.6–2.7 mm in diam., apical part 2.7–4 mm in diam., terete, pubescent or glabrescent, rarely glabrous; sepals and petals same as or slightly larger than in male flowers; sepals 3–7.2  × 2.5–7.2 mm; petals 0.5–1  × 0.4–1 cm; appendages 5, antesepalous, alternating with staminode bundles, fleshy, pitted; staminodes united in 5 bundles, 2–5 in each bundle, antepetalous, 2.2–4.7 mm long each bundle, whitish pale green or creamish white; pistil 5–7.5 mm long; ovary pale green, subglobose, 3–4.5  × 4–6.5 mm, beaked, 0.5–2  × 1.4–2 mm (ovary including beaked looks like obpyriform in outline), unlobed, glabrous, 5-locular; stigma pale green, sessile, radiate, deeply 5-lobed, 3.5–5 mm in diam., papillate. Bisexual flowers same as in male and female flowers (androecium same as in male flowers; gynoecium same as in female flowers). Fruits berries, green, turning yellow or orangish yellow when ripe, smooth, glabrous, glossy, with sticky yellow latex, then exocarp becoming dark brownish black and sinuously wrinkled when dry, subglobose, globose or broadly ovoid, sometimes depressed globose, 3.8–7.5  × 4.5–8 cm, sometimes oblique, asymmetrical, unlobed, with a short, thick beak, pericarp 0.8–1.8 cm thick, exocarp thin; persistent stigma dark brown or blackish brown, radiate, deeply 5-lobed; persistent sepals green, slightly larger than in flowering materials; fruiting stalk green, 1.3–2.1 cm long, widened at the apical part, middle part 1.3–3.8 mm in diam., apical part 2.5–4.5 mm in diam., pubescent or glabrescent, rarely glabrous. Seeds 1–5, sometimes aborted, brown mottled with irregular lines, ellipsoid, 1.7–3  × 0.9–2 cm, slightly oblique, rounded at both ends, with yellow or orangish yellow fleshy pulp (Fig. 5).

Distribution. India (Assam, Andaman Islands), Vietnam, Laos, Cambodia, Thailand, Peninsular Malaysia (also called Malaya) (Perlis, Perak, Kelantan), Indonesia [Java, Lesser Sunda Islands, Sulawesi, Moluccas (also called Maluku)], Borneo (Sabah), Philippines, New Guinea, Australia (Queensland), French Polynesia (Fig. 6).

Distribution in Thailand. Eastern: Buriram, Surin, Ubon Ratchathani; South-Western: Prachuap Khiri Khan; Central (cultivated); South-Eastern: Sa Kaeo, Prachin Buri, Chachoengsao, Chon Buri, Chanthaburi, Trat; Peninsular: Surat Thani, Krabi, Stun, Songkhla, Pattani, Narathiwat (Fig. 6).

Habitat and Ecology. It is found in dry evergreen, tropical lowland evergreen rain forests, and lower montane forests, often in limestone areas, sometimes along streams, 0–1,100 m alt.

Phenology. Flowering and fruiting more than once, nearly throughout the year; flowering usually in November to June; fruiting usually in January to July.

Conservation Status. Garcinia dulcis is widely distributed from India, Andaman Islands to North Queensland and Tahiti Islands. It is known from many localities and has a large Extent of Occurrence (EOO) of 14,325,860.58 km² and a relatively large Area of Occupancy (AOO) of 824 km². In Thailand, this species is known to be naturally distributed in the eastern, the south-western, the south-eastern and the peninsular regions, and has an EOO of 422,327.01 km² and an AOO of 184 km². Because of this wide distribution and the number of localities, it is considered Least Concern (LC).

Etymology. The specific epithet of Garcinia dulcis is a Latin word meaning sweet and refers to the ripe fruits that have a sweet-sour taste (Stearn, 1992; Radcliffe-Smith, 1998; Gledhill, 2002). The specific epithet of its new synonym, G. vilersiana is in honour of the late Charles Marie Le Myre de Vilers (1833–1918), a French naval officer, then departmental administrator. He was governor of the colony of Cochinchina (1879–1882). The specific epithet of its two new synonyms, G. cambodgiensis and G. andamanica are named after Cambodia and Andaman Islands, respectively where the type specimens for these species were collected.

Vernacular Name. Khai chorakhe ( ) (Chanthaburi); Champhut ( ) (Central); Taphut ( ) (Chanthaburi); Prahot ( ), Prahut ( ) (Khmer-Surin); Pahut ( ) (Northeastern); Phahut ( ) (Buri Ram); Phawa bai yai ( ) (Chanthaburi, Chonburi); Phut ( ) (Satun); Maphut ( ) (Central, Peninsular); Sompong ( ), Sommuang ( ) (Chanthaburi); Baniti (Philippines); Cay vang nhua, Vang nhua (Vietnam); Madaw mu (Andaman Islands); Mundu (Javanese, Malay), Munu (Malay); Prahout (Cambodia); Yellow mangosteen (English).

Uses. Garcinia dulcis is often cultivated for its fruits. The fruits (pericarp and fleshy pulp) are edible and have a sour or sweet-sour taste. It is also grown in some botanical gardens as an ornamental tree to provide botanical education. It is cultivated as a fruit tree in Southeast Asia (Allen, 1965; Begum, Barthakur & Sarma, 2013). The sour fruits can be eaten raw or cooked, and they also make jams and preserves (Corner, 1952; Sastri, 1956; Maheshwari, 1964; Allen, 1965; Burkill et al., 1966; Verheij & Coronel, 1992). In Java and Singapore, pounded seeds are applied to cure swellings. In Java, the bark is used to dye mats (Sastri, 1956; Maheshwari, 1964; Burkill et al., 1966; Verheij & Coronel, 1992).

Lectotypification. Xanthochymus dulcis was named by Roxburgh (1820: 66–67. t. 270), who reported that the species is a native of the Molucca Islands and from thence introduced to the Botanic Garden at Calcutta. This species was transferred to the genus Garcinia by Kurz (1874: 88). Roxburgh’s protologue of X. dulcis implies that he based his description on a tree living in the Botanical Garden at Calcutta before he departed India in 1813. No herbarium specimens were cited. Following advice in Forman (1997), in addition to relevant specimens Roxburgh’s illustration (Plate 270) in “Plants of the Coast of Coromandel” should be considered as a lectotype.

Garcinia vilersiana was named by Pierre (1882: 348; 1883: t. 71), who cited four gatherings, Pierre 128, Pierre 773 and Pierre 3641 from Vietnam and Pierre 3642 from Thailand and Vietnam but he did not mention the herbaria in which they were present, and following Art. 9.6 of the ICN (Turland et al., 2018), they constitute syntypes. We could locate the specimens Pierre 128 (in montibus Dinh ad Baria Austro-Cochinchinae) at C [without barcode], K [K000677686], L [L0700329, L0700330, L0700331] and P [P04701129, P04899814] collected on October 1866; at BM [BM000611613] collected on November 1866; at P [P04701128, P04701131, P04701136] collected on March 1867, Pierre 773 (in planitie ad Thu Duc Austro-Cochinchinae) at K [K000742482] and P [P04450818, P04451355, P04458339, P06137125] collected on February 1870; at K [K000677684], L [U1199409] and P [P00903311, P00903312, P04701110, P04898863] collected on February 1871, Pierre 3641 at P [P04701126, P04701127] (in montibus Dinh ad Baria Austro-Cochinchinae, collected in 1866); at L [U1199410], P [P04701122] (in montibus Dinh ad Baria Austro-Cochinchinae, collected in 1867); at BM [BM000611612], K [K000742481, K000677685] and P [P04899815] (in montibus Dinh ad Baria Austro-Cochinchinae, collected on March 1868); at P [P04701123, P04701124] (in montibus Lu in prov. Bien Hoa Austro-Cochinchinae, collected on March 1877) and Pierre 3642 from Thailand at P [P04701111, P04701114, P04701115, ad flumen Mekong in regno Siamico, collected on June 1868]; and from Vietnam at P [P04701120, P04701121, in insula Phu Quoc, collected on July 1877]. Hence, the specimen Pierre 3642 at P [P04701111] is in the best condition and clearly shows the diagnostic characters for the species and is selected here as the lectotype, following Art. 9.3 and 9.12 of the ICN (Turland et al., 2018).

Garcinia cambodgiensis was named by Vesque (1889: 14 t. 92; 1893: 316), who cited the material Pierre 775 from Cambodia (ad montes Kuang Repoeu, in prov. Tpong) but did not mention the herbaria in which they were present, and following Art. 9.6 of the ICN (Turland et al., 2018), they constitute syntypes. We located the specimen Pierre 775 at K [K000677687] and P [P05062528, P05062544, P05062564], therefore the P [P05062528] material is in the best condition and clearly shows the diagnostic characters for the species and is selected here as the lectotype, following Art. 9.3 and 9.12 of the ICN (Turland et al., 2018).

Additional specimens examined. Thailand. Eastern: Buri Ram [Nang Rong District, March 1922 [as Garcinia sp.], Luang Anuwat Wanarak 11 (BK, K)], Surin [Cambodia Boundary, Sangkha District, 15 January 1924 [as G. vilersiana], Kerr 8295 (BM, E [E00839801], K, P [P04701109])], Ubon Ratchathani [Huai Don, Buntharik District, 12 May 2005 [as G. vilersiana], Puudjaa 1410 (BKF); Phu Chong Na Yoi National Park, Na Chaluai District, August 2007, Ngernsaengsaruay G45-10082007 (BKF, dry and spirit materials)]; South-Western: Prachuap Khiri Khan [Kui Buri National Park, Kui Buri District, 23 January 2004 [as G. griffithii, G. xanthochymus], Middleton et al. 2426 (BKF, E [E00348153]); Pran Buri District, 14 April 2012 [as G. vilersiana], Phengklai 16300 (BKF)]; Central: Ang Thong [Wat Klang, Mueang District, 3 April 1972 [as Garcinia sp., G. cf. xanthochymus], cultivated, Maxwell 72-180 (AAU, BK); ibid., 7 March 1976 [as Garcinia sp., G. xanthochymus, G. vilersiana], cultivated, Maxwell 76-135 (AAU, BK, L [L2409257, L2409258])], Saraburi [Phu Khae, 2 April 1948 [as G. vilersiana], cultivated, Chamrueangsri 14 (BKF)], Pathum Thani [locality not specified, 23 July 1979 [as G. vilersiana], cultivated, Muangnoicharoen s.n. (BKF68140); Suan Pathum Palace, 1 February 2010 [as G. xanthochymus], cultivated, Krajangvuthi s.n. (BKF163988)], Nonthaburi [Bang Yai District, 1 December 1987 [as G. vilersiana], cultivated, Paisooksantivatana Y1-12-87(1) (BK)], Bangkok [locality not specified, 13 June 1920, cultivated, Kerr s.n. (BM); locality not specified, 29 January 1922 [as G. vilersiana], cultivated, Marcan 662 (BM, SING); locality not specified, 21 January 1923 [as G. vilersiana], cultivated, Collins 1144 (BM); locality not specified, 4 March 1923 [as G. vilersiana], cultivated, Kerr s.n. (BM); Rat Burana, 1 December 1987 [as G. vilersiana], cultivated, Paisooksantivatana Y1-12-87(2) (BK); Faculty of Forestry, Kasetsart University, Chatuchak District, 15 March 1994 [as G. vilersiana], cultivated, Santi 33 (BK); ibid., 4 June 2023, cultivated, Ngernsaengsaruay & Chanton G50-04062023 (BK, BKF, dry and spirit materials); locality not specified, 27 January 1992 [as G. vilersiana], cultivated, Niyomdham s.n. (BKF126881); Nong Chok District, 2 July 2007, cultivated, Ngernsaengsaruay G44-02072007 (BKF, spirit material); Bangkok Herbarium, Department of Agriculture, Chatuchak District, 12 June 2023, cultivated, Ngernsaengsaruay & Chanton G51-12062023 (BK, BKF)], Samut Prakan [Song Khanong Subdistrict, Phra Pradaeng District, 25 March 2012, cultivated, Ngernsaengsaruay G47-25032012 (BK, BKF, dry and spirit materials), Samut Songkhram [Kradangnga Subdistrict, Bang Khonti District, 1 December 1987 [as G. vilersiana], cultivated, Paisooksantivatana Y1-12-87(4) (BK); Amphawa District, 16 December 2003, cultivated, Bamrungsri 01 (PSU)]; South-Eastern: Sa Kaeo [Ban Dong Yang, Aranyaprathet District, 22 March 1962 [as G. vilersiana, G. cf. xanthochymus], Chantanamuck 95 (BK); Watthana Nakhon District, 5 March 1977 [as G. vilersiana], Unknown s.n. (BKF110455); Khao Ang Rue Nai Wildlife Sanctuary, 18 January 1997 [as G. vilersiana], Santisuk et al. s.n. (BKF201290)], Prachin Buri [Kabin Buri District, 22 December 1924 [as Garcinia sp., G. cambodgiensis, G. xanthochymus], Kerr 9738 (AAU, BK, BM, E [E00839759], K)], Chachoengsao [Khao Takrup, 6 November 1993 [as Garcinia sp., G. xanthochymus], Larsen et al. 44255 (AAU, K)], Chon Buri [Si Racha Forest, 1916 [as G. cambodgiensis], Collins 609 (K); Nong Nam Khiao, Si Racha Forest, 1 December 1927 [as Garcinia sp., G. cambodgiensis, G. cf. xanthochymus], Collins 1856 (BK, BM, K); Phan Sadet Nok, Si Racha District, 13 January 1946 [as Garcinia sp.], Din 155 (BKF); Khao Khiao, Si Racha District, 20 March 1970 [as Garcinia sp., G. cambodgiensis, G. xanthochymus], van Beusekom & Santisuk 3266 (AAU, BKF, C, E [E00772041], K, L [L2409500, L2409501], P [P05062012]); ibid., 5 January 1975 [as G. cambodgiensis, G. xanthochymus], Maxwell 75-2 (AAU, BK, L [L2408878]); ibid., 18 May 1975 [as Garcinia sp., G. xanthochymus], Maxwell 75-529 (AAU, BK, [L (L2409260)]); Khao Khiao Open Zoo, 9 July 2000, Phengklai et al. 12659 (BKF)], Chanthaburi [Makham District, 7 August 1933 [as Garcinia sp.], Winit 418 (BKF); Khao Sa Bap, Laem Sing District, 11 November 1945 [as Garcinia sp.], Wit 117 (BKF); ibid., 20 June 1946 [as Garcinia sp.], Wit 279 (BKF); Laem Sing District, 22 December 1961 [as G. cambodgiensis, G vilersiana], Nicolson 1645 (K, L [L409326]); Khao Soi Dao, 18 December 1974 [as Garcinia sp., G. cambodgiensis, G. xanthochymus], Geesink, Hiepko & Phengklai 7904 (BKF, C, K, P [P05062000]); Khao Soi Dao, Pong Nam Ron District, 5 May 1975 [as Garcinia sp., G. cambodgiensis, G. xanthochymus], Maxwell 75-487 (AAU, BK, L [L2408876, L2408877]); Khao Soi Dao Wildlife Sanctuary, Khao Soi Dao District, 14 March 1995 [as G. cf. vilersiana], Santisuk s.n. (BKF100190); Khao Soi Dao Wildlife Sanctuary, 11 January 1999 [as G. cf. vilersiana], Wongprasert et al. s.n. (BKF124721); Khao Soi Dao Wildlife Sanctuary, 1 March 2007 [as G. nervosa, G. xanthochymus], Mantharanon s.n. (BKF146453); Khao Soi Dao Wildlife Sanctuary, 15 March 2008, Ngernsaengsaruay G46-15032008 (BKF, spirit material); Khao Soi Dao Wildlife Sanctuary, 17 May 2013 [as G. vilersiana], Tagane et al. T1553 (BKF); ibid., 17 May 2013 [as G. vilersiana], Tagane et al. T1596 (BKF); Khao Soi Dao Wildlife Sanctuary, 8 March 2014 [as Garcinia sp.], Tagane et al. T2688 (BKF); Khao Phra Bat, Khao Khitchakut National Park, Makham District, 29 November 1979 [as Garcinia sp., G. xanthochymus], Shimizu et al. T-23989 (BKF, L [L2409479]); Thung Phen Forest Protection Unit, Khao Khitchakut National Park, 27 December 1993 [as G. vilersiana], Niyomdham 3485 (BKF); Khlong Saba, Khao Sip Ha Chan, Kaeng Hang Maeo, 11 February 2007 [as G. xanthochymus], Watthana 2221 (QBG)], Trat [Bo Rai, 28 November 1924 [as G. cambodgiensis], Kerr 9461 (BM, K); Dan Chumphon, 21 December 1929 [as Garcinia sp., G. cambodgiensis, G. xanthochymus], Kerr 17657 (AAU, BK, BM, K); Ban Salak Phet, Ko Chang, 7 March 2003 [as G. mangostana], cultivated, Phengklai et al. 14528 (BKF)]; Peninsular: Surat Thani [Ko Tao, 18 April 1927 [as Garcinia sp., G. vilersiana], Kerr 12797 (BM, K); locality not specified, 1 December 1987 [as G. vilersiana], Paisooksantivatana Y1-12-87(5) (BK); Thung Khai Han, Khao Sok National Park, 14 June 1994 [as G. vilersiana], Niyomdham & Puudjaa 3786 (BKF); Giant Bamboo nature trail, Khlong Phanom National Park, Phanom District, 20 June 2004 [as Garcinia sp., G. vilersiana], Gardner & Sidisunthorn ST0804 (K); ibid., 20 March 2005 [as Garcinia sp., G. vilersiana], Gardner, Sidisunthorn & Tippayasri ST1707 (BKF, K)], Krabi [locality not specified, 4 April 1930 [as G. vilersiana, G. cf. xanthochymus], Kerr 18845 (BK, BM, C, K, L [L2409325]); Ko Pu, 14 April 1930 [as G. vilersiana, G. cf. xanthochymus], Kerr 18972 (BK, BM, K, L [L2409324]); Ao Luek, 23 July 1972 [as Garcinia sp., G. vilersiana], Larsen et al. 31282 (AAU, BKF, K); Than Bok Korani National Park, 9 May 1973 [as Garcinia sp., G. vilersiana], Geesink & Santisuk 5303 (AAU, BKF, C, E [E00839800]); Than Bok Korani National Park, Ao Luek District, 7 March 2022, Ngernsaengsaruay, Meeprom & Boonthasak G49-07032022 (BK, BKF, dry and spirit materials), Trang [Khao Chong Botanical Garden, Chong Subdistrict, Na Yong District, 17 March 2018, cultivated, Ngernsaengsaruay, Wessapak, Meeprom & Boonthasak G48-17032018 (BK, BKF, dry and spirit materials)], Stun [Ban Ton, 15 March 1928 [as Garcinia sp.], Kerr 14598 (BM, K); Tarutao National Park, trail from Malacca Creek to Ao Talo Wao, 24 March 1980 [as Garcinia sp., G. xanthochymus], Congdon 501 (AAU, PSU); near Nam Ra Village, Thung Nui Subdistrict, Khuan Kalong District, 6 January 1985, Maxwell 85-43 (BKF, PSU); Thale Ban National Park, Khuan Don District, 25 May 2004 [as Garcinia sp.], Gardner & Setsin ST0604 (K)], Songkhla [Namtok Boriphat Forest Park, Rattaphum District, 24 October 1985, Maxwell 85-997 (AAU, BKF, L [L2402922, L2402923], PSU); Rattaphum District, 15 May 2002, Upho UBON981 (QBG); Hat Yai District, 6 April 2004, Deachathai 02 (PSU)]; Hat Yai District, 6 April 2004, Mahabusarakam 02 (PSU); Khao Nam Khang National Park, Na Thawi District, 6 October 2004, Tippayasri & Sidisunthorn ST1064 (K); Hat Yai, Khuan Mot Daeng, PSU Campus, 30 May 2005 [as G. nervosa ], Panwiriyarat 01 (PSU)], Pattani [Mueang Pattani District, October 1937, Unknown 32 (BKF221); Tugong Village, Nong Chik District, 14 January 1985 [as G. nervosa var. pubescens], Saree 1 (P [P04701559], PSU), Narathiwat [Ao Manao, 8 August 1999, Niyomdham & Puudjaa 5753 (BKF); Ao Manao, Khao Tanyong National Park, 27 March 2002, Ngernsaengsaruay G43-27032002 (BKF, spirit material); Chon Thara Singhe, Tak Bai District, 17 March 2002, cultivated, Ngernsaengsaruay G42-17032002 (BKF, spirit material)].

India. Bengal, s.d., [as G. xanthochymus], Unknown 257 (E [E00772050]); locality not specified, s.d., [as G. xanthochymus], Stewart’s Collection 311 (E [E00839674]); South Andaman, 1867 [as X. pictorius], Kurz 241 (P [P04701158]); South Andaman, 1867 [as X. pictorius], Kurz s.n. (G [G00458419]); Flora of the Andamans, 1884 [as G. andamanica], King’s Collector s.n. (G [G00458474], L [L2408492], P [P05062493]); South Andaman, 1890 [as G. andamanica], King’s Collector s.n. (E [E00438019], L [L2408491]); South Andaman, 7 April 1894 [as G. andamanica], King’s Collector s.n. (P [P05062497]); Little Andaman, 10 January 1976 [as G. andamanica], Bhargava 3382 (L [L2408490]); Middle Andaman, 4 November 1977 [as G. xanthochymus], Bhargawa & Nooteboom 6343 (L [L2409259]).

Vietnam. Unreadable, April 1863 [as G. xanthochymus], Pierre 3385 (P [P04701165]); Phu Quoc in Gallicae Austro-Cochinchina, May 1865 [as G. xanthochymus ], Pierre 3385 (L [L2409238]); in insula Condor, October 1876 [as G. vilersiana], Harmand 920 (P [P04701105, P04701106, P04701107]); Tonkin, December 1889 [as G. cambodgiensis], Balansa 4345 (P [P05062548, P05062562]); Bieu Hieu, Thanh Hoa, 10 June 1892 [as G. cambodgiensis], l’abbé Bon 5398 (P [P05062536]); Bien Hoa, 28 February 1914 [as G. vilersiana], Fleury 31300 (P [P04701116]); Bien Hoa, 28 March 1914 [as G. vilersiana], Fleury 32046 (P [P04701119]); Phu Tho, 30 May 1918 [as G. cambodgiensis], Fleury 37545 (P [P05062534]); Annam, Dac Kiet, prov. de Thanh Hoa, 10 September 1920 [as G. cambodgiensis], Poilane 1815 (K, P [P05062532]); Tonkin, Lao Cai, 4 January 1931 [as G. cambodgiensis], Poilane 18764 (P [P05062541]); Lai Chau et Muong, 19 April 1936 [as G. cambodgiensis], Poilane 25747 (P [P05062527]); Tonkin, Lai Chau, 12 January 1938 [as G. cambodgiensis], Poilane 27115 (K, P [P05062542]); Plantes du Tonkin occidental, s.d. [as G. cambodgiensis], l’abbé Bon 5362 (P [P05062533, P05062566]).

Laos. Luang Phrabang, expedition du Mekong, 1866–1868 [as G. cambodgiensis], Thorel 9189 (P [P05062546]); Haut-Mékong, 19 May 1936 [as G. cambodgiensis], Poilane 26214 (K, P [P05062535]); Env. de Pakse, Sedone, 18 November 1965 [as G. aff. vilersiana], Vidal 4477 (P [P04701130]).

Cambodia. Expedition du Mekong, 1866–1868 [as G. cambodgiensis], Thorel 3183 (P [P05062529, P05062530, P05062543]); expedition du Mekong, 1866–1868 [as G. vilersiana], Thorel s.n. (P [P04701133]); locality not specified, November 1881 [as G. cambodgiensis], Pierre 4171 (P [P04700185]); locality not specified, 1896 [as G. vilersiana], Hahn s.n. (P [04701134]); Khsach Kandal District, Kandal Province, 7 March 1914 [as G. vilersiana], Unknown 31838 (P [P04701132]); locality not specified, 24 June 1929 [as G. vilersiana], Bejeaud 532 (P [P04701135]); Krapoeu, 26 June 1930 [as G. cambodgiensis], Poilane 17711 (P [P05062545]); Ko Kong, 28 February 2000 [as G. cambodgiensis], Meng 94 (K); Keo Seima District, Mondulkiri Province, 21 May 2001 [as G. vilersiana], Eanghourt & Phirun 851 (K); Siem Reap, Phnom Kulen, 2 July 2006 [as G. vilersiana], Long, Cheng & Leti CL265 (P [P00626101]); Ko Kong, Thma Baing, Veal Kandevech, 19 December 2008 [as G. cambodgiensis], Newman et al. 2069 (E [E00319084]); Stung Treng, Thala Borivat District, 26 March 2012 [as G. cambodgiensis], Maxwell 12-52 (L [L4311881, L4311882]).

Cochinchine. Country and locality not specified, 1862–1866 [as G. vilersiana], Thorel 9189 (P [P04701125]); Country and locality not specified, 1862–1866 [as G. vilersiana], Thorel s.n. (P [P04701108]).

Indonesia. Java, Ujung Kulon National Park, 28 November 1960, Kostermans 241 (L [L2403308]); Java, 30 April 1974, Wiriadinata 80 (AAU); Lesser Sunda Islands, Sumbawa, 4 October 1982, Danimihardja SD2217 (L [L2403235]); Lesser Sunda Islands, East Sumba, Ngallu, 5 September 1994, McDonald & Sunaryo 4402 (E [E00037289]); Sulawesi, 4 July 1976, Meijer 10817 (L [L2403229]); Moluccas, Ceram, 13 December 1990, Burley, Tukirin & Ismail 4393 (E [E00160942]).

Borneo. Malaysia: Sabah, Jesselton District, 6 May 1963, Hashim 33874 (L [L2403302]); Sabah, Kampung Payas, Pitas District, 11 May 1987, Amin et al. SAN121250 (E [E00160943]).

Philippines. Palawan, 12 June 1994, Soejarto et al. 8258 (L [L3813146]); Luzon, 22 February 1991, Loher 74 (US [US351445]); Malapackun, 15 April 1984, Ridsdale SMHI442 (L [L2403249]); Camarines Sur, Lupi, s.d., Vidal 639 (AAU); Angat, Bulacan, s.d., Vidal 640 (AAU); Buenavista, Marinduque, s.d., Vidal 1155 (AAU).

New Guinea. Mt. Klangal, Kandrain Subdistrict, West New Britain District, 16 May 1973, Croft & Katik NGF15591 (US); New Ireland, Ugana, Songmum, West Coast, January 1938, Peekel 30 (L2403148); Kombi Subdistrict, West New Britain District, 28 May 1973, Isles & Katik NGF32248 (L2403218).

Australia. Queensland, Cook District, 11 October 1962, Smith 11719 (L2403506); Queensland, Cook District, 12 October 1962, Smith 11854 (L2403507).

French Polynesia. Papeari, Tahiti Islands, 29 November 1963, Maclet 18 (US).

2. Garcinia nervosa (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavi 1: 208. 1864; Pierre, Fl. Forest. Cochinch. 1(5): 5. 1883; King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 59(2): 169. 1890; Vesque in A. DC. & C. DC., Monogr. Phan. 8: 327. 1893; Merr., Philipp. J. Sci., C 10(5): 325. 1915; Ridl., Fl. Malay Penins. 1: 179. 1922; Merr., Enum. Philipp. Fl. Pl. 3: 86. 1923; Corner, Wayside Trees Mal. 1: 318. ed. 2, 1952; Whitmore in Whitmore, Tree Fl. Malaya 2: 217. 1973; H. Keng, Concise Fl. Singapore: 49. 1990; N. P. Singh in B. D. Sharma & Sanjappa, Fl. Ind. 3: 121. 1993; I. M. Turner, Gard. Bull. Singapore 47(1): 262. 1995; S. Gardner, P. Sidisunthorn & Chayam., Forest Trees S. Thailand 1: 358. fig. 548. 2015.—Stalagmitis nervosa Miq., Fl. Ned. Ind., Eerste Bijv. 3: 496. 1861. Type: Indonesia, Sumatra, Pariaman (originally “Priaman” on the label), s.d., Diepenhorst HB647 (lectotype selected here L [U0002403, photo seen]; isolectotype L [U0002404, photo seen]) (Fig. 7).

— Garcinia andersonii [as andersoni] Hook. f. ex T. Anderson in Hook. f., Fl. Brit. India 1: 270. 1874; Vesque in A. DC. & C. DC., Monogr. Phan. 8: 318. 1893. Type: Peninsular Malaysia, Malacca, s.d., Maingay 157 (distributed at K in 1871) (lectotype selected here CAL [CAL0000005828, photo seen]; isolectotype K [K000677676!]) (Fig. 8).

— Garcinia macrophylla T. Anderson in Hook. f., Fl. Brit. India 1: 270. 1874, nom. inval.

—Garcinia spectabilis Pierre, Fl. Forest. Cochinch. 1(5): 3. 1883; Vesque, Epharmosis 2: 15 t. 91. 1889. Type: Borneo, 1865, Beccari 2966 (lectotype selected here P [P04700284!]; isolectotype K [K000677704!]), syn. nov. (Fig. 9).

—Garcinia nervosa (Miq.) Miq. var. pubescens King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 59(2): 169. 1890; Vesque in A. DC. & C. DC., Monogr. Phan. 8: 327. 1893; Whitmore in Whitmore, Tree Fl. Malaya 2: 218. 1973; I. M. Turner, Gard. Bull. Singapore 47(1): 262. 1995. Type: Peninsular Malaysia, Perak, Larut, 1882, Kunstler 3197 (lectotype selected here CAL [CAL0000005834, photo seen]; isolectotypes G [G00458441, photo seen], K [K000677677!], P [P05062500!], SING [SING00636112!, SING00636113!]), syn. nov. (Fig. 10).

Tree evergreen, 10–35 m tall, 60–160 cm girth; latex white, turning creamish white, very sticky; branches decussate, horizontal; branchlets green, 4-ridged, two of these with a strongly ridged or a narrowly winged, pubescent, slightly pubescent or glabrescent. Bark smooth, rough or fine scaly, brown, dark brown or greyish brown; inner bark pale brown. Terminal bud concealed between the bases of the uppermost pair of petioles. Leaves decussate; lamina narrowly elliptic, narrowly oblong, oblong, lanceolate, lanceolate-ovate or elliptic-oblong, 33.5–80  × 8–27 cm, apex acute, base subcordate, margin repand and slightly revolute, very thickly coriaceous, slightly bullate or bullate, shiny dark green above, paler below, glabrous above, pubescent, slightly pubescent or glabrescent below (of lamina and veins), midrib and secondary veins slightly raised above, strongly ridged below, secondary veins 11–24 pairs, curving towards the margin connected in distinct loops and united into an intramarginal vein, sometimes forked, with intersecondary veins, veinlets reticulate, all veins conspicuous on both surfaces, interrupted wavy lines absent; petiole green, the uppermost pair of petioles reddish purple, turning reddish purple tinged with green to green with age, 2.3–7.2 cm long, 0.4–1.1 cm in diam., not grooved, distinctly transversely rugose, indumentum same as in lamina, with a conspicuous basal appendage clasping the branchlet; young leaves shiny pale green; mature leaves turning greenish yellow to yellow before falling off; dry leaves yellowish brown. Inflorescences on short, leafless lateral branchlets, cymose, in fascicles of 5–15 flowers. Flowers unisexual, plants dioecious, 5-merous, petals erect with overlapping edges and corolla forming a bowl-shaped; bracteolate; sepals and petals quincuncial, coriaceous, concave. Male flowers 0.8–1.4 cm in diam.; bracteoles triangular, 2–5.2 × 1.8–5 mm, apex acute, pubescent; pedicel green, 1.8–3.2 cm long, widened at the apical part, middle part 3.5–4 mm in diam., apical part 4.5–7 mm in diam., 4-angular, thick, pubescent; sepals 5, reddish green, greenish red, red or green, semi-orbicular or broadly ovate, 3.5–8.5 × 4–9 mm, unequal, apex rounded, pubescent outside; petals 5, whitish pale green, creamish white or pale yellow, suborbicular, obovate or broadly obovate, 0.8–1.4 × 0.5–1 cm, subequal, apex rounded, margin ciliolate; disk in the center of the flower, intrastaminal, yellow, 5-lobed, fleshy, pitted, lobes positioned between the stamen bundles, antesepalous; stamens numerous, united in 5 bundles, 11–20 in each bundle, antepetalous, 0.8–1 cm × 2–3 mm each bundle, creamish white or pale yellow; filaments 0.5–0.6 mm long; anthers yellow, 2 thecae, 0.3–0.5 × 0.3–0.7 mm; pistillode absent. Female flowers 1–1.7 cm in diam.; bracteoles and pedicel same as in male flowers; sepals and petals same as or slightly larger than in male flowers; appendages 5, antesepalous, alternating with staminode bundles, fleshy, pitted; staminodes united in 5 bundles, 4–6 in each bundle, antepetalous, 3–4 mm long, 1–1.5 mm wide each bundle, creamish white or pale yellow; pistil 0.7–1 cm long; ovary reddish green or green, broadly ovoid, subglobose or globose, 3.5–6  × 3.2–5.5 mm, beaked, 1.5–3.5 × 1.2–2 mm (ovary including beaked looks like obpyriform in outline), unlobed, glabrous, 5-locular; stigma red, sessile, radiate, deeply 5-lobed, 3–4 mm in diam., papillate. Fruits berries, green or reddish green, turning yellow or yellow blotched with red when ripe, slightly rough, glabrous, not glossy, with sticky yellow latex, then exocarp becoming dark brownish black and sinuously wrinkled when dry, subglobose, globose or broadly ovoid, 5–7.5 × 4.5–7 cm, sometimes oblique, asymmetrical, unlobed, with a short, thick beak; persistent stigma red or greenish red, radiate, deeply 5-lobed, erect; persistent sepals slightly larger than in flowering materials; fruiting stalk green, 3–5.2 cm long, widened at the apical part, middle part 4–6.5 mm in diam., apical part 0.5–1 cm in diam., 4-angular, thick, pubescent. Seeds 2–5, sometimes aborted, brown mottled with irregular pale brown lines, semi-ellipsoid, 2.8–3.2 × 1.5–2 cm, rounded at both ends, with a thin white fleshy pulp (Fig. 11).

Distribution. India (Andaman and Nicobar Islands), Peninsular Thailand, Peninsular Malaysia [Perak, Terengganu (also called Trengganu), Pahang, Selangor, Malacca (also called Melaka)], Singapore, Indonesia (Sumatra), Borneo (Sabah, Sarawak, Brunei, East Kalimantan, West Kalimantan, South Kalimantan), Philippines (Palawan, Luzon) (Fig. 12).

Distribution in Thailand. Peninsular: Nakhon Si Thammarat, Phatthalung, Trang, Satun, Pattani, Yala, Narathiwat (Fig. 12).

Habitat and Ecology. It is usually found in tropical lowland evergreen rain forests, sometimes in limestone areas, often near or along streams, 50–250(–600) m alt. In Peninsular Malaysia and Singapore, it occurs in lowland, lowland dipterocarp, hill dipterocarp and freshwater swamp forests, sometimes along streams or near the sea up to elevations of 750 m amsl. (Corner, 1952; Whitmore, 1973; the specimens from Peninsular Malaysia and Singapore).

Phenology. Flowering and fruiting more than once; flowering nearly throughout the year, usually in January to April; fruiting February to August.

Conservation Status. Garcinia nervosa is widely distributed from Andaman and Nicobar Islands to Philippines (POWO, 2023) and is very rare in Singapore (Keng, 1990); the specimen Sinclair 10915). It has a large EOO of 3,215,117.07 km² and a relatively large AOO of 292 km². In Thailand, this species is known to be naturally distributed in the peninsular region, and has an EOO of 25,909.66 km² and an AOO of 40 km². There doesn’t appear to be an imminent threat to the plants or their habitats. Therefore, we consider the conservation assessment here as LC.

Etymology. The specific epithet of Garcinia nervosa is a Latin word, referring to the strongly or prominently nerved (veined) leaves (Stearn, 1992; Gledhill, 2002). The specific epithet of its new synonym G. spectabilis is a Latin word meaning admirable, spectacular or good-looking, refers to the character of the leaves (Gledhill, 2002).

Vernacular Name. Chamuang nam ( ) (Yala); Phut ( ) (Nakhon Si Thammarat, Phatthalung, Trang); Maphut pa ( ) (Pattani); Mu-lu ( ) (Malay-Pattani); Asam garam, Kandis gajah, Pakok lapan taun (Malay); Buradgis, Kabal, Gatatán (Philippines); Pear mangosteen (English).

Uses. The fruits (pericarp and fleshy pulp) are edible and have a sour or sweet-sour taste. In Malaysia, the sour pulp is eaten cooked with sugar (Bircher & Bircher, 2000). In Andaman and Nicobar Islands, the fruits have been used by the Nicobarese for food and medicinal purposes and the wood is used by the Nicobarese and the Shompen for making canoe paddles (Waman, Bohra & Mane, 2018). Leaves are pounded into paste, boiled in coconut oil, and rubbed onto body and joints for pain relief (National Parks, Flora and Fauna Web, 2023). The leaves and bark contain high antioxidative and anti-inflammation properties, which have great potential in the development of pharmaceutical and dermatological products (Seruji, Khong & Kutoi, 2013). It is a tree suitable for gardens, parks and roadsides (National Parks, Flora and Fauna Web, 2023).

Lectotypification. Stalagmitis nervosa was named by Miquel (1861: 496), who cited two gatherings from Sumatra: D. (Diepenhorst) in “Priaman” and T. (Teysman) in “Lubu-alang” but did not mention the collector number and the herbaria in which they were present. This species was transferred to the genus Garcinia by Miquel (1864: 208). We only found two sheets of the specimen Diepenhorst HB647 collected from “Priaman” at L [U0002403, U0002404], and following Art. 9.6 of the ICN (Turland et al., 2018), they constitute syntypes. Friedrich Anton Wilhelm Miquel (Miq.) (1811–1871) was a Dutch botanist, a professor of botany and a director of the Amsterdam botanical garden (1846–1859), then a director of the Utrecht Botanical Garden (1859–1871), and from 1862 was a director of the Leiden Rijksherbarium. Miquel’s private herbarium, containing many of his types, is the basis of the general herbarium of U. Other Miquel types are, however, in several herbaria because Miquel described many new taxa based on material obtained on loan. Most of these types are at L; others are at G, P, and K. The plants in many herbaria labelled “Ex Herbario Miquel”, were not collected by Miquel himself but by various other collectors (Stafleu & Cowan, 1981). Therefore, the specimen Diepenhorst HB647 at L [U0002403] is selected here as the lectotype, following Art. 9.3 and 9.12 of the ICN (Turland et al., 2018).

Garcinia andersonii (as G. andersoni) was named by Joseph Dalton Hooker (Hook. f.) and then described by Anderson (1874: 270), who cited the collection Maingay 157 from Malacca but did not mention the herbaria in which it was present. The authors located two sheets of this specimen at CAL [CAL0000005828] and K [K000677676], and following Art. 9.6 of the ICN (Turland et al., 2018), they constitute syntypes. Thomas Anderson (1832–1870) was a Scottish botanist and a superintendent of the Calcutta Botanical Gardens (1860–1868) (Stafleu & Mennega, 1992). We therefore selected the CAL specimen as the lectotype, following Art. 9.3 and 9.12 of the ICN (Turland et al., 2018).

Garcinia spectabilis was named by Pierre (1883: 3) based on the specimen Beccari 2966 from Borneo but he did not mention the herbaria in which it was present. We could locate two sheets of this specimen at K [K000677704] and P [P04700284], and following Art. 9.6 of the ICN (Turland et al., 2018), they constitute syntypes. Jean Baptiste Louis Pierre (1833–1905) was a French botanist, a director at the Saigon botanical garden, and explored Cambodia, Cochinchina, and southern Thailand (1865–1877), and returned to France in 1877 (Stafleu & Cowan, 1983)). Hence, the P [P04700284] material is selected here as the lectotype, following Art. 9.3 and 9.12 of the ICN (Turland et al., 2018).

Garcinia nervosa var. pubescens was named by King (1890: 169), based on the cited specimen King’s Collector 3197 from Larut, Perak but he did not mention the herbaria in which it was present. We traced six sheets of the specimen Kunstler 3197 at CAL [CAL0000005834], G [G00458441], K [K000677677], P [P05062500] and SING [SING00636112, SING00636113] but only the specimens at K, P and SING were labeled as King’s Collector 3197. However, all materials are of Hermann H. Kunstler as many of his collections are labelled “King’s Collector” (van Steenis-Kruseman & van Steenis, 1950), and were collected from the same locality. Following Art. 9.6 of the ICN (Turland et al., 2018), they constitute syntypes. Sir George King (1840–1909) was a British botanist, a superintendent of the Calcutta Botanic Gardens (1871–1898), and a director Botanical Survey of India (1891–1898) (Stafleu & Cowan, 1979). Therefore, the CAL [CAL0000005834] specimen is selected here as the lectotype, following Art. 9.3 and 9.12 of the ICN (Turland et al., 2018).

Additional specimens examined. Thailand. Peninsular: Nakhon Si Thammarat [National Park Protection Unit, Krung Ching Waterfall, Khao Luang National Park, Nop Pitam District (originally “Tha Sala District” on the label), 17 March 2005 [as Garcinia sp.], Sidisunthorn & Tippayasri ST1688 (K); ibid., 27 February 2006 [as Garcinia sp.], Gardner & Sidisunthorn ST1688a (BKF, K)], Phatthalung [Lan Mom Chui Waterfall, Ban Tamot Wildlife Protection Unit, Tamot District, 23 August 1996, BGO. Staff 7317 (QBG); Tamot Waterfall (also called Lan Mom Chui Waterfall), Tamot Subdistrict, Tamot District, 8 March 2022, Ngernsaengsaruay & Boonthasak G35-08032022 (BK, BKF); locality not specified, 24 November 2004, Watcharin 01 (PSU)], Trang [16-ha plot, Khao Chong, February 2001, Sinbumroong & Davies AS286 (BKF); 24-ha long-term ecological research plot, Khao Chong, Chong Subdistrict, Na Yong District, 16 February 2022, Ngernsaengsaruay, Wessapak, Meeprom & Boonthasak G34-16022022 (BK, BKF); Ton Te Waterfall, Palian District, February 2001 [as Garcinia sp.], Sinbumroong & Davies AS259 (BKF); Ban Tha Khao, Palian District, 10 July 2004, Maneenoon & Srimok 698 (PSU); Namtok Phan Forest Park, Palian Subdistrict, Palian District, 17 March 2018, Ngernsaengsaruay, Wessapak, Meeprom & Boonthasak G33-17032018 (BK, BKF)], Satun [Ya Roi Waterfall, Thaleban National Park, Khuan Don District, 18 March 2004, Gardner & Sidisunthorn ST0247 (BKF)], Pattani [reported by Gardner, Sidisunthorn & Chayamarit (2015)], Yala [Bang Lang National Park, Aiyoeweng Subdistrict, Betong District, 22 May 2022 (Ngernsaengsaruay own observation, with photos); reported by Gardner, Sidisunthorn & Chayamarit (2015)], Narathiwat [Chat Warin Waterfall, To Deng Subdistrict, Su-ngai Padi District, 22 April 1995, Niyomdham 4080 (BKF); Pacho Waterfall, Budo–Su-ngai Padi National Park, Bacho District, 27 December1999 [as Garcinia sp.], Wongprasert 9912-77 (BKF)]; Peninsular, Province not specified [locality not specified, s.d., Winit 35 (BKF)].

Peninsular Malaysia. Perak [Larut, July 1886, King’s Collector 10491 (CAL [CAL0000005829]); Gerik, 3 February 1994, Penomot & Teo 4334 (P [P04701558])], Terengganu [Tasik Kenyir, Hulu Terengganu, 31 July 2007, Imin et al. Kep. Forest Research Institute, FRI58501 (L [L3806836])], Pahang [Ulu Keniyam, 3 March 1968, Shah 1531 (C); Krau Wildlife Reserve, Kuala Lompat, 15 February 1983, Daveson s.n. (E [E00839740])], Malacca [Air Kroh, 26 February 1989, Khairuddin Kep. FRI33158 (L [L3806892]).

Singapore. MacRitchie Reservoir, 31 August 1964, Sinclair 10746 (E [E00839741, E00839742]); Nee Soon, Seletar Forest, 6 September 1966, Sinclair 10915 (E [E00839738, E00839739]).

Indonesia. Sumatra [Sibolangit, 29 March 1908, Lörzing 5601 (L [L2408483, U1199434]); Eil. Simaloer, 3 August 1918, Achmad 554 (L [L2408480, U1199435]); ibid., 26 December 1918, Achmad 813 (L [L2408503, L2408504, U1199431]); ibid., 9 June 1919, Achmad 1165 (L [L2408488, L2408489, U1199433]); Res. Palembang, 17 April 1920, Endert 165EIP 857 (L [U1199432])].

Borneo. Malaysia: Sabah [Bengkoka Forest Reserve, Kudat, 7 September 1972 [as G. mangostana], Shea & Minjulu SAN75989 (L [L2416697]); TM 1, Ranau, Sabah, 24 February 1990, Unknown SAN128804 (L [L3806446])]; Indonesia: East Kalimantan [PT. ICTI, road Kenangan to Sepaku, 25 April 1995, Kessler et al. 896 (L [L2416702]); Kaltim Prima Coal (KPC area), Bengalon, Sebongkok Utara, 8 April 1996, Arifin et al. AA1699 (L [L2416700, L2416701]); KPC area, Sangatta, 6 April 1997, Kessler et al. 2386 (L [L2416698])], West Kalimantan [Serawai, 13 October 1995, Church et al. 2465 (L [L3811160]); Gunung Palung National Park, Ketapang, Kalimantan Barat, 8 October 1997, Laman et al. 1148 (L [L3813083, L3813084])], South Kalimantan [PT. Aya Yayang, Kabupaten Tabalong, 17 July 2000, Sidiyasa & Arifin 2160 (L [L2416699])]; Brunei [Labi, Bukit Teraja, Belait, 11 November 1990, Kirkup 267 (L [L3806485]; Labi, Teraja, Belait, 23 May 1996, Joffre et al. 17472 (L [L3878579])].

Philippines. Palawan [Municipality, San Vicente, 1 April 1990, Soejarto & Madulid 7215 (L [U1199430]); Taytay, 1913, Merrill 9387 (P [P04700489, P04700490])].

3. Garcinia prainiana King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 59(2): 171. 1890; Vesque in A. DC. & C. DC., Monogr. Phan. 8: 329. 1893; Ridl., Fl. Malay Penins. 1: 180. 1922; Corner, Wayside Trees Mal. 1: 320. fig. 112. ed. 2. 1952; Corner & Watan., Ill. Guide Trop. Pl.: t. 193. 1969; Whitmore in Whitmore, Tree Fl. Malaya 2: 220. 1973; I. M. Turner, Gard. Bull. Singapore 47(1): 263. 1995. Type: Peninsular Malaysia, Perak, Kuala Dipang (originally “Kwala Dipang” on the label; originally published “Kwala Dynong”), February 1885, Scortechini 1796 (lectotype selected here CAL [CAL0000005844, photo seen]; isolectotypes K [K000677678!], P [P04701324, photo seen]) (Fig. 13).

Tree evergreen, 3–12 m tall, 15–75 cm girth; latex white, sticky; branches decussate, horizontal; branchlets green, terete, glabrous. Bark smooth or slightly rough, pale brown, greyish brown or blackish brown; 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 pairs, curving towards the margin connected in distinct loops and united into an intramarginal vein, flattened above, raised below, conspicuous on both surfaces, with intersecondary veins, veinlets reticulate, visible on both surfaces, interrupted long wavy lines of differing lengths, nearly parallel to the midrib, running across the secondary veins to the apex, visible below; petiole green, short, 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 opened flowers with spreading petals; bracteolate; sepals and petals 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 5, pinkish green, reddish green or greenish red, concave, broadly ovate or suborbicular 4.8–8 × 5–7.8 mm, unequal, apex rounded; petals 5, 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; a small ring-shaped disk surrounding the base of the pistillode; stamens numerous, united in 5 bundles surrounding the pistillode, antepetalous, 1.7–4.2 mm long, 1.2–4 mm wide each bundle, pale yellow, pink or red; filaments fused throughout their entire length; anthers yellow, 2 thecae, 0.3–0.6 mm long; pistillode mushroom-shaped (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 mushroom-shaped, 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, 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, depressed globose or depressed subglobose, sometimes globose, 2–3.5 × 2–4.5 cm, sometimes oblique, asymmetrical, unlobed, slightly concave or flattened at the apex, pericarp 3.5–8 mm thick, exocarp thin; persistent stigma dark brown or blackish brown, circular, button-like, 6–9 mm 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, 0.4–1 cm long, 3–5 mm in diam., thick. Seeds 1–6, often aborted, brown, broadly ellipsoid, ellipsoid or subglobose, 0.9–1.5 × 0.7–1.2 cm, with pale orange fleshy pulp. (Figs. 14 and 15).

Distribution. Known only from Peninsular Thailand and Peninsular Malaysia. It is widely distributed in Peninsular Malaysia: Perlis, Kedah, Penang (also called Pulau Pinang), Perak, Kelantan, Terengganu, Pahang, Selangor, Negeri Sembilan (also called Negri Sembilan), Malacca (also called Melaka) and Johor (also spelled Johore) (Corner, 1952; Whitmore, 1973; Turner, 1995; Azuan & Salma, 2018). It can be found mainly in Pahang, Perak, and Negeri Sembilan (Syazwani, 2020) (Fig. 16).

Distribution in Thailand. It is uncommon in Peninsular Thailand: Yala (Than To) and Narathiwat (Waeng and Su-ngai Kolok) (Fig. 16).

Habitat and Ecology. It is found in tropical lowland evergreen rain forests, occasionally along streams, 30–200 m alt. 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 (Whitmore, 1973; Syazwani, 2020; from the specimen Whitmore Kep. FRI4018).

Phenology. Flowering and fruiting more than once; flowering nearly throughout the year, usually in February to May; fruiting April to June and September to December. According to Syazwani (2020), in Peninsular Malaysia, the fruits are borne once a year, from July to September.

Conservation Status. Garcinia prainiana is widely distributed in Peninsular Malaysia and less common in southern Thailand. It has an EOO of 359,696.7 km² and an AOO of 72 km² and does not face any threat of extinction. We therefore consider the conservation assessment as LC in agreement with Kochummen (1998).

Etymology. The specific epithet of Garcinia prainiana refers to Sir David Prain (1857–1944), a British botanist, an herbarium curator of the Royal Botanic Garden, Calcutta (1887–1898), and a director of the Royal Botanic Gardens, Kew (1905–1922) (Stafleu & Cowan, 1983)).

Vernacular Name. Chupu ( ) (Malay-Narathiwat); Cerapu, Chekau, Chepu, Cherapu, Cherpu, Cherupu, Chupak, Chupu, Kechupu, Kecupu, Menchepu, Menchupu (Malay); Button mangosteen (English).

Uses. Garcinia prainiana is locally cultivated for its fruits in southern Thailand. The fruits (pericarp and fleshy pulp) 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 (fleshy pulp) are edible and are sometimes used fresh in beverages (Allen, 1965; Burkill et al., 1966). The pulp of fruits of has high antioxidant content of about 91.9% and vitamin C content of about 27.3 mg per 100 g fresh weight (Azuan & Salma, 2018). In a traditional Malay recipe, the raw fruits are described as being cooked with dried fish (Zawiah & Othaman, 2012). The wood is used for house building (Allen, 1965; Burkill et al., 1966). It is an excellent ornamental plant for use in landscape gardens in parks (National Parks, Flora and Fauna Web, 2023).

Lectotypification. Garcinia prainiana was named by King (1890: 171–172), who cited the specimen Scortechini 1796 collected from Perak, “Kwala Dynong” but he did not mention the herbaria in which it was present. We located three sheets of the specimen Scortechini 1796: one sheet at CAL [CAL0000005844], one sheet at K [K000677678] and one sheet at P [P04701324], which were collected from the same locality, and following Art. 9.6 of the ICN (Turland et al., 2018), they constitute syntypes. Therefore, the CAL [CAL0000005844] specimen is selected here as the lectotype, following Art. 9.3 and 9.12 of the ICN (Turland et al., 2018).

Additional specimens examined. Thailand. Central: Nakhon Nayok [Phrueksaphan Thepparat Botanicical Garden, Chulachomklao Royal Military Academy, cultivated, 31 May 2019, Ngernsaengsaruay & Boonthasak G30-31052019 (BK, BKF); Peninsular: Trang [Khao Chong Botanical Garden, Chong Subdistrict, Na Yong District, cultivated, 16 February 2022, Ngernsaengsaruay, Meeprom & Boonthasak G32-16022022 (BK, BKF)], Yala [Chulabhorn Phatthana 7 Project, Than To District, near waterfall, 27 November 2019, Ngernsaengsaruay & Sichaikhan G31-27112019 (BK, BKF)], Narathiwat [Hala-Bala Wildlife Sanctuary, Ban Bala, Lo Chut Subdistrict, Waeng District, 13 May 2005 [as Garcinia sp.], Poopath 274 (BKF); Hala-Bala Wildlife Sanctuary, Waeng District, 22 September 2005, Niyomdham & Puudjaa 7593 (BKF); Su-ngai Kolok District, 20 April 2002, Upho 556 (QBG); Su-ngai Kolok District, cultivated, 20 May 2003, Upho 550 (BKF)].

Peninsular Malaysia. Perak [Kwala Dipang, December 1896, Curtis 3273 (K [K000677679]); Kg Kepayang near Ipoh, 30 October 1971, Syed Abu Bakar Kep. FRI20440 (L [L2417220])], Pahang [Su-ngai Bertam at Kuala Mensum, 2 June 1971, Whitmore Kep. FRI20091 (L [L2417222]); Path leading to Kuala Mensum from Boh Tea, Cameron Highlands, 24 September 1971, Loh Kep. FRI19187 (L [L2417221]); Cameron Highlands Road, 18 January 1982, Kochummen Kep. FRI29377 (L [L2417225]), Kelantan [0.5 mile east of Gua Musang, 14 July 1967, Whitmore Kep. FRI4018 (L [L2417226]); Su-ngai Lebir, below Kuala Relai at Jentah, 24 April 1976, Stone & Sidek 12426 (L [L2417224], BKF); Su-ngai Long off Su-ngai Pergau, Jeli, 26 September 1986, Latiff et al. ALM1856 (L [L3806490], PSU); Ketam, Cicar Tinggi, Kampung Bata, Pasir Mas, 1 August 1992, Noorsiha et al. Kep. FRI39214 (L [L3878683]); Pasir Putih, 23 October 1992, Husmady et al. Kep. FRI39551 (L [L3806959]); near Brooke Camp, Gua Musang, 2 June 1994, Husmady et al. Kep. FRI41841 (L [L2417223])].

4. Garcinia xanthochymus Hook. f. ex T. Anderson in Hook. f., Fl. Brit. India 1: 269. 1874; Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 43(2): 88. 1874 et Forest Fl. Burma 1: 93. 1877; Pierre, Fl. Forest. Cochinch. 1(5): 3. t. 71A. 1883; King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 59(2): 168. 1890; Vesque, Epharmosis 2: 14. t. 82–84. 1889 et in A. DC. & C. DC., Monogr. Phan. 8: 315. 1893; Brandis, Indian Trees: 49. 1906; C. E. Parkinson, Forest Fl. Andaman Isl.: 89. 1923; Craib, Fl. Siam. 1(1): 118. 1925; Kanjilal, P. C. Kanjilal & A. Das, Fl. Assam 1(1): 104. 1934; Gagnep. in Gagnep., Fl. Indo-Chine Suppl.: 257. 1943; Maheshw., Bull. Bot. Surv. India 6: 114. t. 1. fig. 3. 1964; Whitmore in Whitmore, Tree Fl. Malaya 2: 222. 1973; C. J. Saldanha & Nicolson, Fl. Hassan Dist.: 127. 1976; Kosterm. in Dassan. & F. R. Forsberg, Revis. Handb. Fl. Ceylon 1: 87. 1980; D. G. Long in Grierson & D. G. Long, Fl. Bhutan 1(2): 370. fig. 30k–o. 1984; C. J. Saldanha & E. Rao, Fl. Karnataka 1: 207. 1984; P. H. Hô, Câyco Vietnam 1: 568. fig. 1573. 1991; E. W. M. Verheij & R. E. Coronel (eds.), PROSEA 2: 175, 176. 1992; N. P. Singh in B. D. Sharma & Sanjappa, Fl. Ind. 3: 129. 1993; S. Gardner, P. Sidisunthorn & V. Anusarnsunthorn, Field Guide Forest Trees of N. Thailand: 50. fig. 54. 2000; X. W. Li, J. Li, N. Robson & P. F. Stevens in C. Y. Wu, P. H. Raven & D. Y. Hong, Fl. China 13: 42. 2007; W. E. Cooper, Austrobaileya 9(1): 8. 2013; R. Tabassum, Angiospermic Fl. Gazipur Distr. Bangladesh (Dissertation): 110. 2015.—Xanthochymus pictorius Roxb., Pl. Coromandel 2(4): 51, t. 196. 1805, Hort. Bengal.: 42. 1814 et in Carey, Fl. Ind. 2: 633. 1832 [non Garcinia pictoria Roxb. (Roxburgh, 1832: 627–629)]; Dalzell & A. Gibson, Bombay Fl.: 31. 1861.—Stalagmitis pictoria (Roxb.) G. Don, Gen. Hist. 1: 620. 1831.—Garcinia pictoria (Roxb.) Engl. in Engl. & Prantl, Die Naturlichen Pflanzenfamilien 3(6): 234. fig. 114 C–F. 1895, nom. illeg.—Garcinia pictoria (Roxb.) D’Arcy, Ann. Missouri Bot. Gard. 67: 998. fig. 4A. 1980, nom. illeg.—Xanthochymus tinctorius DC., Prodr. 1: 562. 1824, orth. var.—Garcinia tinctoria (DC.) W. Wight, Bull. Bur. Pl. Industr. U.S.D.A. 137: 50. 1909, orth. var.—Garcinia tinctoria Dunn in Gamble, Fl. Madras 1: 74. 1915, orth. var.—Garcinia roxburghii Kurz, Prelim. Rep. Forest Pegu App. A: xiii. 1875, nom. illeg. Type: Roxburgh’s Herbarium, Wallich Cat. 4837A (lectotype designated here (K-W [K001104026, photo seen]) (Fig. 17).

Tree evergreen, 10–30 m tall, 40–160 cm girth; latex white, turning creamish white, sticky; branches decussate, horizontal; branchlets green, 4-ridged, glabrous. Bark smooth or slightly rough, brown, dark brown or greyish brown; inner bark pale yellow. Terminal bud concealed between the bases of the uppermost pair of petioles. Leaves decussate; lamina variable in shape, oblong, elliptic, oblong-elliptic, narrowly oblong, narrowly elliptic or narrowly oblong-elliptic, sometimes ovate, 19–32.5 × 6–18 cm, apex acute or acuminate, sometimes obtuse, base cuneate, margin repand and slightly revolute, thickly coriaceous, smooth (not bullate), shiny dark green above, paler below, glabrous on both surfaces, midrib and secondary veins flattened above, raised as a prominent ridge below, secondary veins 12–16 pairs, curving towards the margin connected in distinct loops and united into an intramarginal vein, conspicuous on both surfaces, with intersecondary veins, veinlets reticulate, visible on both surfaces, interrupted long wavy lines of differing lengths absent, sometimes indistinct below; petiole green, the uppermost pair of petioles reddish purple, turning green with age, 1.2–3.2 cm long, 2.5–7 mm in diam., not grooved, distinctly transversely rugose, glabrous, with a conspicuous basal appendage clasping the branchlet; young leaves shiny pale green; mature leaves turning greenish yellow to yellow before falling off. Inflorescences on short, leafless lateral branchlets, cymose, in lax fascicles of 3–8 flowers. Flowers unisexual, plants dioecious, 5-merous, petals erect with overlapping edges and corolla forming a bowl shape; bracteolate; sepals and petals quincuncial, coriaceous, concave. Male flowers the same size as in female flowers; bracteoles and pedicel same as in female flowers; sepals and petals same as or slightly smaller than in female flowers; disk in the center of the flower, intrastaminal, yellow, 5-lobed, fleshy, pitted, lobes positioned between the stamen bundles, antesepalous; stamens numerous, united in 5 bundles, 9–13 in each bundle, antepetalous, 0.8–1 cm long each bundle; filaments 0.7–2 mm long; anthers 2 thecae, 0.3–0.7 mm long; pistillode absent. Female flowers 1–1.3 cm in diam.; bracteoles triangular, 1.5–3 × 1.8–2.7 mm, apex acute, glabrous; pedicel green or reddish green, 1.5–5 cm long, widened at the apical part, middle part 1–2.5 mm in diam., apical part 2.5–4.5 mm in diam., terete, glabrous; sepals 5, pale green or greenish pale yellow, semi-orbicular or broadly obovate, 2.7–7 × 2.8–6.5 mm, unequal, apex rounded, margin ciliolate, glabrous outside; petals 5, whitish pale green or creamish white, suborbicular or broadly obovate, 0.7–1 × 0.7–1.2 cm, subequal, apex rounded, margin ciliolate; appendages 5, antesepalous, alternating with staminode bundles, fleshy, pitted; staminodes united in 5 bundles, 2–5 in each bundle, antepetalous, 2.2–4.2 mm long each bundle, whitish pale green or creamish white; pistil 0.5–1 cm long; ovary pale green, subglobose, globose or broadly ellipsoid, 3.2–7.5 × 3–6 mm, beaked, 1–3.5 × 1.3–2 mm (ovary including beaked looks like obpyriform in outline), unlobed, glabrous, 5-locular; stigma pale green, sessile, radiate, deeply 5-lobed, 4–7 mm in diam., papillate. Fruits berries, green, turning yellow when ripe, smooth, glabrous, glossy, with sticky yellow latex, then exocarp becoming dark brownish black and sinuously wrinkled when dry, subglobose, globose or broadly ovoid, 4.7–6 × 4–7.5 cm, sometimes oblique, asymmetrical, unlobed, with a short, thick beak, pericarp 1–1.5 cm thick, exocarp thin; persistent stigma dark brown or blackish brown, radiate, deeply 5-lobed; persistent sepals green, slightly larger than in flowering materials; fruiting stalk green or reddish green, 2.5–5 cm long, widened at the apical part, middle part 2.5–3.7 mm in diam., apical part 4–6 mm in diam., glabrous. Seeds 1–5, sometimes aborted, brown mottled with irregular lines, semi-ellipsoid, 1.8–3 × 1.2–2.7 cm, rounded at both ends, with yellow fleshy pulp. (Fig. 18).

Distribution. India, Andaman Islands, Nepal, Bhutan, Bangladesh, Myanmar, China, Vietnam, Thailand. This species has been introduced and is cultivated in many countries (e.g., Panama, Africa, Sri Lanka, Peninsular Malaysia, Singapore, Queensland and French Polynesia) (Fig. 19). It has been naturalized in Sri Lanka (Kostermans, 1980); from the specimens Kostermans 24019; Jayasuriya 1626; Jayasuriya 1628) and a few locations in Queensland (Cooper, 2013).

Distribution in Thailand. Northern: Mae Hong Son, Chiang Mai, Chiang Rai, Payao, Nan, Lampang, Phrae, Tak, Kamphaeng Phet; North-Eastern: Loei; South-Western: Uthai Thani, Kanchanaburi (Fig. 19).

Habitat and Ecology. It is found in dry evergreen, mixed deciduous and lower montane forests, sometimes in limestone areas, often along or near streams, 100–1300 m alt.

Phenology. Flowering and fruiting more than once, nearly throughout the year.

Conservation Status. Garcinia xanthochymus is widely distributed from Indian subcontinent to China and Indo-China. It is known from many localities and has a large EOO of 3,579,093.97 km² and a relatively large AOO of 272 km². In Thailand, this species is known to be naturally distributed in the northern, the north-eastern and the south-western regions, and has an EOO of 165,710.42 km² and an AOO of 112 km². Because of this wide distribution and the number of localities, we consider the conservation assessment here as LC.

Etymology. The specific epithet of Garcinia xanthochymus is derived from the Greek compound words, xantho- meaning yellow, and chymo, meaning sap, referring to the plant with yellow latex (Stearn, 1992; Radcliffe-Smith, 1998; Gledhill, 2002).

Vernacular Name. Cha-kha-sa ( ) (Karen-Mae Hong Son); Mada ( ) (Northern); Mada luang ( ) (Chiang Mai); Mapong ( ) (Phrae); Asam kandis (Malay); Dampel, Tamal (Hindi); Madaw (Myanmar); Rata goraka (Ceylon); Assam mangosteen, Egg tree, False mangosteen, Himalayan garcinia, Mysore gamboge, Sour mangosteen (English).

Uses. The fruits (pericarp and fleshy pulp) are edible and have a sour or sweet-sour taste and are used in the same way as Garcinia dulcis. G. xanthochymus is cultivated in Southeast Asia where the sour fruits can be used for making preserves, jams, sherbets, curries and vinegar. The latex from the bark and fruits provides a dye that is used in watercolor paints (Maheshwari, 1964; Burkill et al., 1966; D’Arcy, 1980; Kostermans, 1980; Verheij & Coronel, 1992; Singh, 1993). It is cultivated as an ornamental tree (Maheshwari, 1964; D’Arcy, 1980; Singh, 1993) and is suitable for streetscapes, parks and gardens (National Parks, Flora and Fauna Web, 2023). In India, the fruits are used for making medicaments (Maheshwari, 1964; Burkill et al., 1966) and the rootstocks are sometimes used for grafting on mangosteen (G. mangostana L.) (Maheshwari, 1964; D’Arcy, 1980; Kostermans, 1980; Singh, 1993). It contains many phytochemicals that can be extracted from its constituent parts: the bark, fruits, leaves, roots, twigs and seeds. The predominant extracted phytochemicals are xanthones, benzophenones, flavonoids, depsidones and isocoumarins. These phytochemicals contribute to the pharmacological activities of this plant as an antioxidant, antidiabetic and as having Nerve Growth Factor-potentiating, antimicrobial and cytotoxic activities. This species contains a broad range of phytochemicals with curative properties that can be greatly beneficial to man (Che Hassan, Taher & Susanti, 2018).

Additional specimens examined. Thailand. Northern: Mae Hong Son [Muang Soi Waterfall, Pai District, 17 January 1983, Koyama et al. T-32653 (BKF); Kuet Luang Waterfall, Namtok Mae Surin National Park, Khun Yuam District, 23 May 2003 [as Garcinia sp.], Wongprasert 035-50 (BKF, K, P [P06899613]); Huai Mae Sae, Salawin Wildlife Sanctuary, Mae Sariang District, 22 February 2007 [as Garcinia sp.], Watthana 2302 (QBG)], Chiang Mai [Doi Suthep-Pui National Park, 23 May 1910, Kerr 1201 (BM, K); ibid., 26 April 1958 [as Garcinia sp.], Sørensen et al. 3059 (C); ibid., 29 April 1958 [as Garcinia sp.], Sørensen et al. 3128 (C); ibid., 10 October 1958 [as Garcinia sp.], Sørensen et al. 5568 (C); ibid., 28 November 1994, Kopachon s003b1 (CMUB, L [L3813373]); ibid., 17 December 1995 [as Garcinia sp.], BGO Staff 5455 (QBG); Ru Si Cave, Doi Suthep-Pui National Park, 5 January 1988, Maxwell 88-1 (AAU, L [L2409261]); ibid., 13 April 2003, Maxwell & Canines 2 (CMUB); Doi Suthep-Pui National Park, 2 December 2002, Ngernsaengsaruay G38-02122002 (BKF, dry and spirit materials); Mae Sa, Mae Rim District, 11 February 1923, Kerr s.n. (BM); Queen Sirikit Botanic Garden (originally called “Mae Sa Botanic Garden”), Mae Rim District, 9 December 1987, Santisuk s.n. (BKF); ibid., 3 July 1992, Maxwell 92-361 (AAU, CMUB, E [E00160934], L [L3806732, L3806733], P [P04701167]); ibid., 15 October 1993, Pooma 720 (BKF); ibid., 19 October 1993, BGO Staff 033 (QBG); ibid., 5 March 1994, BGO Staff 488 (QBG); ibid., 4 December 1998, Prachit et al. 13 (QBG); ibid., 16 March 1999, Panich 18 (PSU); ibid., 10 April 2007, Glamwaewwong 1291 (QBG); Queen Sirikit Botanic Garden, Mae Rim District, 9 October 2003, Ngernsaengsaruay G39-09102003 (BK, BKF, dry and spirit materials); Doi Chiang Dao, 10 January 1975 [as Garcinia sp.], Geesink et al. 8213 (AAU, C, K, L [L2409576, L2409577], P [P05061711]); Chiang Dao District, 25 June 1989, Maxwell 89-802 (L [L2409264]); Mae Pan, Doi Inthanon, 4 December 1969, van Beusekom & Phengklai 2283 (AAU, BKF, E [E00839802], P [P05062058]); Ban Mai Phatthana, Nong Pa Ko Subdistrict, Mae Chaem District, 30 July 1988, Maxwell 88-945 (BKF, L [L2409262, L2409263]); Ban Mae Mu, Mae Chaem District, 9 February 2017, Pongamornkul 5951 (QBG); Pong Duet Hot Spring, Pa Pae Subdistrict, Mae Taeng District, 19 November 1992, Maxwell 92-745 (CMUB, L (L3806731), P [P04701166])], Chiang Rai [Mae Chan District, 27 January 1970 [as G. cambodgiensis], Sutheesorn 1550 (BK); Ban Dong Mada, Mae Lao District, 11 August 2007 [as G. mckeaniana], Klomsakul 5 (BKF); Khun Chae National Park, 19 January 1998 [as Garcinia sp.], Wongchai WU067 (BKF); Tham Luang-Khun Nam Nang Non Forest Park, Mae Sai District, 14 January 2011, Norsaengsri & Tathana 7559 (QBG); ibid., 25 May 2011, Norsaengsri & Tathana 7864 (BKF, QBG); Ban San Pa Sak, Tham Luang-Khun Nam Nang Non Forest Park, Mae Sai District, 29 March 2012, Norsaengsri & Tathana 9297 (QBG)], Payao [Champathong Waterfall, Doi Luang National Park, 7 April 1999 [as Garcinia sp.], Srisanga & Watthana 611 (QBG)], Nan [Sapan Waterfall, Khun Nan National Park, Bo Kluea District, 17 November 1993 [as Garcinia sp.], Larsen et al. 44509 (AAU, BKF)], Lampang [Doi Khun Tan National Park, Hang Chat District, 2 October 1994, Maxwell 94-1079 (BKF, CMUB, L [L3811034, L3811035]); Chae Son National Park, Mueang Pan District, 22 April 1996, Maxwell 96-548 (BKF, CMUB, L [L3813120, L3813121]); Chae Son National Park, Mueang Pan District, 28 June 2002 [as Garcinia sp.], Koonkhunthod et al. 204 (BKF); ibid., 28 June 2002, Ngernsaengsaruay G36-28062002 (BK, BKF, dry and spirit materials); Wang Kaeo Waterfall (originally “Wahng Gayo Falls” on the label), Doi Luang National Park, Wang Nuea District, 27 March 1997, Maxwell 97-274 (BKF, CMUB, L [L3883203])], Phrae [Huai Pu, 30 August 1982 [as Garcinia sp.], Suvarnasuddhi 376 (BKF)], Tak [Doi Umphang, Umphang District, 9 May 1976 [as Garcinia sp.], Sutheesorn 3756 (BK); Ti Lo Su Waterfall, Umphang Wildlife Sanctuary, Umphang District, 9 September 2002, Ngernsaengsaruay 37-09092002 (BK, BKF, dry and spirit materials); Umphang Wildlife Sanctuary, Umphang District, 25 July 2008 [as Garcinia sp.], Promhitathorn s.n. (BKF); Tha Song Yang District, 29 May 2008, Pooma et al. 7069 (BKF, L [L3811067])], Kamphaeng Phet [Huai Nam Lai, 8 November 1968, Smitinand 10484 (BKF); ibid., 8 November 1968, Smitinand 10486 (BKF); Khlong Lan Waterfall, Khlong Land District, 25 November 1997 [as Garcinia sp.], Phengklai et al. 3918 (BKF, PSU)]; North-Eastern: Loei [Phu Luang Wildlife Sanctuary, 17 May 1998 [as Garcinia sp.], Chayamarit et al. 1481 (BKF); Enroute from Lon Tae to Wang Saphung, Phu Luang Wildlife Sanctuary, 17 May 1998 [as Garcinia sp.], Wongprasert et al. s.n. (BKF SN121739, the leaves belong to G. xanthochymus but the fruits belong to Syzygium; SN121740); Suan Hin Pha Ngam Park, Nong Hin District, 8 December 2014 [as Garcinia sp.], Tagane et al. T3686 (BKF)]; South-Western: Uthai Thani [Huai Kha Khaeng Wildlife Sanctuary, Ban Rai District, 21 February 1970 [as Garcinia sp.], van Beusekom & Santisuk 2895 (AAU, BKF, C, E [E00772042], L [L2409527, L2409528, L2409529], P [P05062005]); Huai Kha Khaeng Wildlife Sanctuary, 3 May 2023, Ngernsaengsaruay G40-03052023 (BK, BKF); ibid., 3 May 2023, Ngernsaengsaruay G40-03052023 (BK, BKF)], Kanchanaburi [Si Sawat District, 12 July 1962 [as G. vilersiana], Suvarnakoses 2092 (BKF); Erawan National Park, between Khwae Noi and Mae Klong River, 17 April 1968, van Beusekom & Phengklai 480 (AAU, C, E [E00839803], K, L [L2409581], P [P05062006]); Erawan Waterfall, Si Sawat District, 4 March 1975 [as Garcinia sp.], Indrapong et al. 122 (BKF, C, K, L [L2409511, L2409512]); Khao Ko Kae, Erawan National Park, Si Sawat District, 3 November 1979 [as G. nervosa], Shimizu et al. T-21656 (BKF); Erawan National Park, 7th waterfall, 2 September 1995 [as Garcinia sp.], Parnell et al. 95-684 (K); Trail to Takhian Thong Waterfall, Sangkhla Buri District, 24 August 2010 [as Garcinia sp.], Chamchumroon et al. 4786 (BKF)].

India. Sikkim, s.d. [as X. pictorius], Hooker s.n. (E [E00839669], P [P04701144]); Assam, s.d. [as X. pictorius], Masters s.n. (L [L2409240], P [P04701146, P04701149]); Khasia, s.d. [as X. pictorius ], Hooker & Thomson s.n. (P [P04701148, P04701153], G [G00458423]); Shillong, Meghalaya, 20 August 1886, Clarke 44626 (G [G00458424]); Neilgherries, 1857 [as X. pictorius], Cleghorn s.n. (E [E00839668]); Malabar Concan, s.d. [as X. pictorius], Stocks & Law s.n. (P [P04701151], G [G00458422]); East Bengal, s.d. [as X. pictorius], Herb. Griffith 872 (P [P04701147]); Peninsula Indiae Orientalis, distributed at the Royal Gardens, Kew, 1866–1867 [as X. pictorius], Herb. Wight 139 (L [L2409239]); Flora of the Andaman Islands, 1884, King’s Collector 242 (L [L2409235], P [P04701159]); Guitar Island, Andaman Islands, February–March 1934, Ram 3689 (E [E00839676]); cultivated in Hort. Bot. Calcuttensis, 1864, Pierre 3385 (P [P04701152, P04701160, P04701161]); ibid., s.d. [as X. pictorius], Unknown s.n. (L [L2409247], P [P04701154]); H.B.C. (Calcutta Botanical Garden), 1832 [as X. pictorius], Unknown Cat. 4837D (EICH 4837D) (K-W [K001104029], G [G00458427]).

Sri Lanka. Udawatte Kele above Kandy, naturalized, 7 May 1971, Kostermans 24019 (C, L [L2409230, L2409231, L2409232], P [P04701162, P04701163]); Kandy town, June 1971, Kostermans s.n. (L [L2409241]); Udawatte Kele Sanctuary, Kandy District, naturalized, 30 April 1974, Jayasuriya 1626 (L [L2409548]); ibid., naturalized, Jayasuriya 1628 (L [L2409549]); locality not specified, s.d., Walker 128 (E [E00839683]).

Bhutan. Longa Khola near Phipsoo, Sarbhang District, 17 March 1982, Grierson & Long 3804 (E [E00170193]).

Bangladesh. Sylhet (originally “Sillet” on the label), April 1824 [as X. pictorius], de Silva Cat. 4837E [East India Company Herbarium (EICH) 4837E] (G [G00458431], K-W [K001104030], P [P04701157]); Flora of the Chittagong Hill Tracts, 1886, King’s Collector 421 (E [E00839677], L [L2409237], P [P04701150]); Chittagong, s.d. [as X. pictorius], Hooker & Thomson s.n. (P [P04701145]); E. Bengal, 10 May 1945, Sinclair 4273 (E [E00839671]); Kaptai, Sitapahar west, Jarultala (Silchari), Rangamati District, 24 April 1997, Huq & A. I. 10466 (L [L3883042, L4449015]).

Myanmar. Phanoe hills, Salween River (originally “Phanoe fluminis Saluen” on the label), 13 March 1827 [as X. pictorius], Wallich Cat. 4837F (EICH 4837F) (G [G00458466], K-W [K001104031]); Nidaun, Ataran River (originally “Nidaun fluminis Attran” on the label), 28 January 1827 [as X. pictorius], Wallich Cat. 4837G (EICH 4837G) (G [G00458433], K-W [K001104032]); ibid. (originally “Nidong on the Attran” on the label), s.d., Wallich 1903 (G [G00458428]); Shan, October 1891, Huk 57 (P [P04701156]); Phanse on the Saluen, s.d. [as X. tinctorius], Wallich 1902 (G [G00458455]); Gamon Reserve, Tharrawaddy District, Bago, 9 May 1912, Smales 46 (E [E00839670]); Myingyan District, 10 August 1914, cultivated, Rogers 410 (E [E00839684]); Pozaungdaung Reserve Compart. 8, Yamethin District, Mandalay, 9 January 1915, Rogers 556 (E [E00839672]); Yangon (originally “Rangoon” on the label), March 1938, Dickason 7142 (E [E00839682]; L [L2409234]).

China. Mienning, Mayetui, Yunnan, 24 September 1938, Yu 17709 (E [E00839463]); Guangzhou, 1982 [as G. tinctoria], Xing et al. 166 (E [E00839462]); Chia I Agriculture Research Institute, Taiwan, 18 May 1966, Liao 10585 (L [L2409233, L2409243]); Fujian, Xiamen, 27 May 2020, Wang Tao 175 (AU [AU080688]).

Country not specified. Locality not specified, s.d. [as X. pictorius], Herb. Roxburgh, Unknown Cat. 4837A (EICH 4837A) (K-W [K001104026]); locality not specified, 1832 [as X. pictorius], Herb. Madras, Unknown Cat. 4837B (EICH 4837B) (K-W [K001104028], G [G00458430]).