Research Article |
Corresponding author: Dongwei Zhao ( zhaodw@csuft.edu.cn ) Academic editor: Oscar Vargas
© 2024 Dongwei Zhao.
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:
Zhao D (2024) Taxonomic revision of Camellia (Theaceae) in Thailand. PhytoKeys 239: 29-57. https://doi.org/10.3897/phytokeys.239.113878
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Natural plants of Camellia (Theaceae) in Thailand are taxonomically revised with a key, a distribution map, descriptions, specimens examined, and figures of living plants and/or dry specimens. Nine taxa comprising seven species and two varieties are recognized, including C. caudata, C. connata, C. furfuracea, C. kissi, C. kissi var. confusa, C. laotica, C. sinensis var. assamica, C. suddeeana, and C. taliensis. Camellia caudata and C. laotica are new records to Thailand, and C. connata and C. suddeeana are endemic to the country. Formerly recorded C. pleurocarpa and C. tenii are excluded from Thai flora because of misidentification, and C. oleifera and C. sinensis are merely cultivated in the country. Morphological descriptions of C. connata and C. laotica are improved based on additional collections examined.
Assam tea, Indochina, new records, oil camellias
Camellia L. (Theaceae) contains economically important plants, including tea, oil camellia and camellias. They are evergreen shrubs or trees distributed in East, South and Southeast Asia (
Morphological characters were described based on living plants, herbarium specimens or their images from BK, BKF, BM, C, CMUB, E, GXMI, HITBC, IBK, IBSC, K, KKU, KUN, L, MO, NSW, P, PE, PNH, QBG, SING, SYS, and TCD (acronyms following
A key to all nine taxa of Camellia occurring in Thailand is present below. Taxa in Thailand are subsequently listed alphabetically.
1 | New branchlet and petiole glabrous | 2 |
– | New branchlet and petiole hairy | 3 |
2 | Pedicel 1–3 mm long, sepals abaxially pubescent | C. furfuracea |
– | Pedicel 4–8 mm long, sepals abaxially glabrous | C. taliensis |
3 | Sepals caducous after flowering | 4 |
– | Sepals persistent after flowering | 5 |
4 | Flowers 1–4.5 cm in diam., styles 1.5–7 mm long, pericarp 0.5–1.5 mm thick | C. kissi |
– | Flowers 4.5–10 cm in diam., styles 8–12 mm long, pericarp 2–8 mm thick | C. kissi var. confusa |
5 | Style 1, apically (2–)3–5 lobed | 6 |
– | Styles 3, distinct | 8 |
6 | Sepals 3–4, ovary glabrous | C. laotica |
– | Sepals 5, ovary hairy | 7 |
7 | Bracteoles caducous, sepals abaxially glabrous | C. sinensis var. assamica |
– | Bracteoles persistent, sepals abaxially pubescent | C. caudata |
8 | Filaments yellowish orange, completely united into a tube, ovary pubescent | C. connata |
– | Filaments white, basally connate ca. 1/3, ovary glabrous | C. suddeeana |
= Camellia assimilis Champ. ex Benth., Hooker’s J. Bot. Kew Gard. Misc. 3: 309. 1851. Lectotype (designated by Chang & Bartholomew [1984: 205]): China. Hong Kong, J.G. Champion 65 (K 000380537!).
= Camellia gracilis Hemsl., Ann. Bot. (Oxford) 9(33): 146. 1895. Holotype: China. Taiwan: Bankinsing mountains, A. Henry 1612 (K 000380535!).
= Camellia triantha Hung T. Chang, Taxon. Gen. Camellia 144. 1981. Holotype: China. Guangxi: Yongning, 20 October 1963, F.S. Huang 17625 (SYS 00094835!).
= Camellia tubiformis Hung T. Chang & S.X. Ren, Acta Sci. Nat. Univ. Sunyatseni 31(1): 75. 1992. Holotype: China. Guangdong: Fengkai, Heishiding, 580 m, 1 January 1991, R.X. Jiang s.n. (SYS 00094839!).
Lectotype
(designated by
Trees or shrubs up to 15 m tall. New branchlets and terminal buds pubescent. Petioles 1–5 mm long, pubescent; leaf blades elliptic, oblong or lanceolate, 3.5–11.5 × 1.5–3 cm, thinly coriaceous or papery, abaxially pale green, sparsely appressed pubescent or villous, adaxially yellowish or dark green, shiny, hirsute along midrib, midrib abaxially elevated and adaxially slightly impressed, secondary veins 8–12 on each side of midrib, abaxially slightly elevated and adaxially obscure, base attenuate to obtuse, margin serrulate, apex caudate. Flowers solitary or up to 3 in a cluster, 2–4 cm in diam. Pedicel 2.5–7 mm long. Bracteoles 3–6, alternate, persistent, deltate to semi-orbicular, 1–2.5 × 1–3 mm, abaxially sparsely pubescent to pubescent, adaxially glabrous, margin ciliolate. Sepals 5, persistent, suborbicular, 2–3 × 2–4 mm, abaxially pubescent, adaxially glabrous. Petals 5–7 in 1–2 whorls, white, elliptic to obovate, 11–20 × 8–14 mm, abaxially pubescent, adaxially glabrous, apex rounded to emarginate, inner 4–5 petals basally adnate to filament whorl for 2–4 mm. Stamens numerous, 10–15 mm long; filaments white, outer filaments basally connate for 6–10 mm, distinct part villous. Ovary globose to ovoid, densely pubescent. Style 1, 8–18 mm long, basally densely pubescent and gradually becoming sparsely pubescent apically, apically 3-lobed for 1–3 mm. Capsule globose to ovoid, 11–15 mm in diam., 1-loculed with 1 seed; pericarp 0.4–1 mm thick. Seeds fuscous, globose, 1–1.5 cm in diam., glabrous Fig.
Flowering October–December, fruiting March–December.
Camellia caudata occurs in semi-evergreen, evergreen and deciduous montane forests at the elevations of 500–2000 m in China, India, Laos, Myanmar, Nepal, Thailand (Fig.
Distribution of Camellia taxa in Thailand: C. caudata (Pink square), C. connata (Purple star), C. furfuracea (Pink plus), C. kissi (Blue dot), C. kissi var. confusa (Brown diamond), C. laotica (Red star), C. sinensis var. assamica (Green triangle), C. suddeeana (Red dot) and C. taliensis (Blue plus).
Chiang Rai: Wiang Pa Pao, Doi Luang, Doi Mok Mountain, close to source of Man Mae Nga Stream, 19°15'36"N, 99°20'24"E, 950 m, 23 July 1998, Morci 1365.0 (CMUB).
Nan: Doi Tiu, 1100 m, 8 March 1921, Kerr 5041 (BK 203924, BM, K, L.2399739); Tha Wang Pha, Pa Kha, Doi Wao, 1400–1700 m, 10 September 1995, Larsen et al. 46319 (BKF SN147973) & 46292 (IBSC 0256482).
Uttaradit: Phu Soi Dao, 1531 m, 17 November 2009, Norsaengsri & Intamusik 6104 (QBG 42621), 1570 m, 18 November 2009, Norsaengsri & Intamusik 6202 (QBG 42737).
Camellia caudata is a new record to Thailand. It generally occurs in the montane forests at elevations of 900–1700 m of Northern Thailand. The species is one of several camellias that are widely distributed in Indochina, along with Assam tea, C. furfuracea and C. kissi. Kerr A.F.G. collected specimens of C. caudata in 1921 (Kerr 5041 at BK, BM, K and L). The specimens were, however, misidentified as C. connata by Keng H. in 1970 as shown on the sheet conserved at BM. Camellia caudata can be distinguished from C. connata by its basally connate styles whereas the latter bears distinct styles.
≡ Thea connata Craib, Bull. Misc. Inform. Kew (1): 6. 1914. Lectotype (first-step designated by
Shrubs or trees up to 8 m tall. New branchlets puberulous; terminal buds pubescent. Petioles 2.5–7 mm long, sparsely puberulous; leaf blades elliptic to oblong, 5.5–12 × 3–5 cm, thinly coriaceous, abaxially yellowish green and brown punctate, glabrous or sparsely puberulous along midrib, adaxially yellowish or dark green, shiny, hirsute along midrib, midrib abaxially elevated and adaxially slightly raised, secondary veins 9–12 on each side of midrib, slightly raised on both surfaces, base attenuate to rounded, margin serrulate, apex acute to attenuate. Flowers solitary or paired, ca. 2 cm in diam., subsessile. Bracteoles 3–6, alternate, persistent, gradually transitioning to sepals, deltate to semi-orbicular, 1.5–3 × 1–2.5 mm, abaxially glabrous or sparsely puberulous, adaxially glabrous. Sepals 4–5, persistent, sub-orbicular or ovate, 3.5–5 × 4–6 mm, abaxially glabrous or sparsely puberulous, adaxially glabrous, margin ciliolate and usually lacerate. Petals 6–7 in 2 whorls, elliptic or ovate, glabrous, outer 2 petals 5–7 × 7–9 mm, greenish white, inner 4–5 petals basally adnate to filament whorl for ca. 3 mm. Stamens 14–20; filaments yellowish orange, glabrous, outer 9–14 filaments completely united into a 7–12 mm long tube, ca. 5 mm in diam., apex irregularly dentate, each tooth bearing an anther, remaining filaments 6–9 mm long, borne on the inside of filament tube. Ovary globose to ovoid, pubescent. Styles 3, distinct, ca. 1.5 mm long, glabrous or sparsely puberulous at base. Capsule ovoid or globose, 10–25 mm in diam., 1–3-loculed with 1–3 seeds; pericarp 0.5–1 mm thick. Seeds black to brown, globose or hemispherical, 1–1.5 cm in diam., glabrous Fig.
Flowering April–August, fruiting July–November.
Camellia connata is endemic to northern Thailand (Fig.
Chiang Mai: 610 m, 16 July 1923, Winit 1134 (BKF SN060818, K); Chiang Dao, Doi Chiang Dao, east side of Sop Huay Pah Dahng-Huay Nah Lao Station, 575 m, 18 August 1995, Bella & Nanny 5 (CMUB 06549); Chom Thong, Doi Inthanon, Mae Uam Station, ca. 1700 m, 2 April 2008, Watthana 2735 (QBG 37391); Fang, Doi Ang Khang, ca. 1500 m, 14 July 1922, Kerr 6294 (BK 203711, BM, K), 1400–1800 m, 8 June 1976, Charoensorn s.n. (BK 203714); Mae Chaem, ca. 800 m, 22 October 1922, Kerr 6428 (BK 203710, K), Doi Inthanon, between 34 km and 35 km, on the road to Mae Aum Watershed Management Station, 1590–1640 m, 12 November 2015, Zhao et al. 84 (BKF, TCD), 85–88 (TCD), 89 (BKF, TCD), 90 (BKF, KUN, TCD), 91 (KUN, TCD), 92 (BKF, KUN, TCD), 93 (1), (2) (TCD) & 94 (BKF, KUN, TCD); Mueang Chiang Mai, Doi Suthep-Pui, 18°49'42"N, 98°53'26"E, 1636 m, 13 November 2015, Zhao et al. 97 (1) & (2) (TCD). Mae Hong Son: Pai, Doi Chang, 2000 m, 31 May 1977, Santisuk 1131 (BKF SN178498).
≡ Thea furfuracea Merr., Philipp. J. Sci., C 13: 149. 1918. Holotype: China. Guangdong: Huizhou, Boluo, Luofu Mountain, 9–27 August 1917, E.D. Merrill 10681 (PNH 87432, image!).
= Thea bolovenensis Gagnep., Notul. Syst. (Paris) 10: 124. 1942. Lectotype (designated by
= Camellia suaveolens C.X. Ye, X.J. Wang & X.G. Shi, Acta Sci. Nat. Univ. Sunyatseni 43(3): 129. 2004. Holotype: China. Guangdong: Yingde, cultivated, introduced from Lechang, 27 October 2002, C.X. Ye 5919 (SYS 00142796!).
= Camellia maiana Orel, Novon 20(2): 198. 2010. Holotype: Vietnam. Lam Dong: Dalat Plateau, 19 November 2002, G. Orel et al. 21149 (NSW 901884, image!).
= Camellia curryana Orel & Luu, Nordic J. Bot. 32(1): 42. 2014. Holotype: Vietnam. Lam Dong: Dalat Plateau, 27 February 2002, G. Orel et al. 21147 (NSW 901031, image!).
= Camellia duyana Orel, Curry & Luu, Novon 23(3): 308. 2014. Holotype: Vietnam. Lam Dong: Dalat Plateau, ca. 1400 m, 27 November 2010, G. Orel & N.V. Duy 0719 (NSW 901883, image!).
= Camellia albata Orel & Curry, Pursuit Hidden Camellias Vietnam China 239. 2015. Holotype: Vietnam. Quang Ninh, 2 December 1999, G. Orel et al. 991202c (NSW 901898 [sheet 1 of 2, image!] and NSW 849513 [sheet 2 of 2, image!]).
= Camellia reflexa Orel & Curry, Pursuit Hidden Camellias Vietnam China 209. 2015. Holotype: Vietnam. Vinh Phuc: Tam Dao National Park, 2 April 2001, G. Orel et al. 1240 (NSW 901749, image!).
= Camellia viscosa Orel & Curry, Pursuit Hidden Camellias Vietnam China 214. 2015. Holotype: Vietnam. Lam Dong: Dalat Plateau, 19 November 2002, G. Orel et al. 21148G (NSW 901821, image!).
Shrubs or trees up to 15 m tall. New branchlets glabrous; terminal buds glabrous or puberulous. Petioles 3–12 mm long, glabrous; leaf blades elliptic or oblong, 7.5–17 × 3–7 cm, coriaceous, abaxially yellowish or pale green, brown punctate or glabrous, adaxially dark green, glabrous and shiny, midrib abaxially elevated and adaxially channelled, secondary veins 7–11 on each side of midrib, abaxially raised and adaxially slightly impressed, base attenuate, cuneate or obtuse, margin sparsely serrulate, apex attenuate, acute or obtuse. Flowers solitary or paired, 4–6 cm in diam. Pedicels 1–3 mm long. Bracteoles and sepals 6–9, caducous, crescent or ovate, 2–13 × 3.5–20 mm, abaxially puberulous or pubescent, adaxially glabrous, margin membranous. Petals 6–9 in 1–2 whorls, white, elliptic or obovate, 17–25 × 9–15 mm, abaxially puberulous or glabrous, adaxially glabrous, basally connate for 1–2 mm. Stamens numerous, 10–15 mm long; filaments glabrous, basally adnate to petals for ca. 2 mm. Ovary ovoid or globose, pubescent. Styles 3–4, distinct, 6–10 mm long, pubescent and gradually becoming glabrous apically. Capsule oblate or globose, 3–5 cm in diam., 2–3 cm in height; usually 3-loculed with 1–3 seed per locule; pericarp 1–10 mm thick, furfuraceous. Seeds brown or fuscous, hemispherical or polyhedral, 1.5–2 cm in diam., glabrous Fig.
Flowering October–February, fruiting December–April.
Camellia furfuracea is distributed in the evergreen forest, on slopes or along streams at elevations of 450–1800 m in China, Laos, Thailand (Fig.
Chanthaburi: Pong Nam Ron, Khao Soi Dao, ca. 1600 m, 23 January 1956, Smitinand 3242 (BKF SN060898), 1100 m, 5 April 1974, Smitinand 11994 (BKF SN060894), 1650 m, 9 February 1975, Smitinand 12023 (BKF SN060895, BKF SN060896, P 04500286; Fig.
Camellia furfuracea is absent in
= Camellia keina Buch.-Ham. ex D. Don, Prodr. Fl. Nepal. 224. 1825. Holotype: Nepal. Narain hetty, 9 September 1802, F. Buchanan-Hamilton s.n. (BM 000521691!).
= Camellia symplocifolia Griff., Itin. Pl. Khasyah Mts. 40, No. 652. 1848. Lectotype (designated by
= Thea bachmaensis Gagnep., Notul. Syst. (Paris) 10: 124. 1942. Lectotype (first-step designated by
= Thea brachystemon Gagnep., Notul. Syst. (Paris) 10: 125. 1942. Lectotype (designated by
= Theopsis euonymifolia Hu, Acta Phytotax. Sin. 10(2): 140. 1965. Holotype: China. Yunnan: Pu’er, Jingdong, 1050 m, 13 December 1939, M.G. Li 1506 (KUN 1206027!).
= Camellia thailandica Hung T. Chang & S.X. Ren, Acta Sci. Nat. Univ. Sunyatseni 30(1): 67. 1991. Holotype: Thailand. Khao Yai National Park, Khao Khieo, 14°21'N, 101°22'E, 1200–1300 m, 29 October 1970, C. Charoenphol et al. 4205 (MO; isotypes BKF!, C!, K!).
= Camellia ligustrina Orel, Curry & Luu, Novon 23(3): 310. 2014. Holotype: Vietnam. Lam Dong: Mount Lang Biang, 1850 m, 16 December 2011, G. Orel & A.S. Curry 0734 (NSW 900397, image!).
= Camellia cuongiana Orel & Curry, Pursuit Hidden Camellias Vietnam China 180. 2015. Holotype: Vietnam. Lam Dong: Bidoup-Nui Ba National Park, 5 November 2012, G. Orel et al. 0721 (NSW 901040, image!).
Holotype : NEPAL. 1818, Gardner s.n. (BM 000948697!).
Shrubs or trees up to 9 m tall. New branchlets puberulous; terminal buds puberulous or glabrous. Petioles 2–7 mm long, puberulous; leaf blades elliptic, oblong or obovate, 3–13.5 × 1.5–5 cm, coriaceous, abaxially yellowish green and sparsely puberulous or glabrous, adaxially dark green, hirsute along midrib, midrib abaxially elevated and adaxially slightly impressed or flat, secondary veins 6–11 on each side of midrib, base attenuate, cuneate or rounded, margin serrate, apex attenuate, acuminate or caudate. Flowers 2–4.5 cm in diam., subsessile. Bracteoles and sepals 5–12, caducous, crescent to broadly ovate, 1.5–7 × 2.5–9 mm, abaxially glabrous, puberulous or pubescent, adaxially glabrous, margin ciliolate. Petals 5–6, white, elliptic or obovate, 8–25 × 4–17 mm, abaxially glabrous, puberulous or pubescent at apex, adaxially glabrous. Stamens numerous, 4–9 mm long; filaments yellow, glabrous, basally connate for 1–2 mm. Ovary globose, pubescent. Styles 3, basally connate, 2–7 mm long, basally pubescent and gradually becoming glabrous apically. Capsule globose or ovoid, 1–3 cm in diam., 1.5–3.5 cm in height, 1–3-loculed with 1–3 seeds; pericarp 0.5–1.5 mm thick. Seeds brown or black, hemispherical, polyhedral or globose, 1–2 cm in diam., glabrous Fig.
Flowering October–February, fruiting March–August.
Camellia kissi occurs in evergreen and mixed deciduous forests, usually by streams, at elevations of 50–2150 m in Bhutan, Cambodia, China, India, Laos, Myanmar, Nepal, Thailand (Fig.
Chiang Mai: Chom Thong, Doi Inthanon, 3 December 1964, Bunchuai 1420 (BKF SN060820), from 23 km point on the main road to Bang Nong Lum, ca. 1100 m, 16 February 1998, Konta et al. 4312 (BKF); Mae Chaem, Doi Inthanon, 1700 m, 24 December 1996, Hara A178 (BKF SN173304).
Chanthaburi: Soi Dao, Khao Soi Dao, ca. 1400 m, 13 December 1924, Kerr 9651 (BK 203727, BM, C).
Kamphaeng Phet: Pang Sila Thong, Mae Wong, 900–1140 m, 14 June 1995, Niyomdham et al. 4379 (BKF).
Kanchanaburi: Sai Yok, Chongkhaosoong, ca. 900 m, 12 August 1995, Wongprasert s.n. (BKF SN112557); Tong-pha-phum, E-tong, 28 January 2001, Veesommai 1.38 (BKF SN145916).
Khon Kean: Phu Wiang, Tap Phaya Suea, 1 December 2003, Mattapha 472 (KKU), 16°37'58"N, 102°12'59"E, 430 m, 22 December 2015, Zhao et al. 137 (BKF, KKU, KUN, TCD).
Loei: Dan Sai, Phu Lom Lo, 1500–1600 m, 8 April 1922, Kerr 5782 (BK 203732, BM, C); Phu Kradueng, ca. 1200 m, 12 March 1924, Kerr 8695 (BK 203922, BM), 16°53'–54'N, 101°47'–49'E, 31 October 1984, Mitsuta et al. T-40368 (BKF SN107567) & T-42270 (BKF SN060860), Pha Nok En, 1 September 1988, Fukuoka T-63791 (BKF SN103457), summit plateau, trail from Than Sawan to Tham Sor Nue waterfall, 1100–1200 m, 12 September 1990, Chantaranothai et al. 90/143 (K); Phu Ruea, 1300 m, 4 May 1997, Pooma 1550 (BKF), Phu Luang, 16 June 2004, Bunwong 296 (KKU), Khoknokkraba, ca. 950 m, 17 November 2000, Wongprasert 11-13 (BKF SN133293); Wang Saphung, Phu Luang, 15 April 1968, Chermsirivathana 899 (BK 203742).
Nakhon Nayok: Mueang Nakhon Nayok, Khao Yai, 14°26'10"N, 101°22'28"E, 730 m, 8 May 2003, Charoenchai & Phomphuang 399 (BK 263128, CMUB 26064); Pak Phli, Khao Yai, 14°21'N, 101°22'E, 1200–1300 m, 29 October 1970, Charoenphol et al. 4205 (BKF SN060889).
Nakhon Ratchasima: Pak Chong, Khao Yai, ca. 600 m, 11 August 1974, Maxwell 74-786 (BK 203724), Orchid waterfall, 600–750 m, 18 November 1982, Koyama et al. T-30008 (BKF SN060882), Haew Suwat waterfall, 700–730 m, 19 November 1982, Koyama et al. T-30110 (BKF SN060881), 14°26'10"N, 101°22'29"E, 730 m, 7 October 2002, Charoenchai & Phomphuang 343 (BK 263207, CMUB 26356).
Nakon Si Thammarat: Lan Saka, Khao Luang, 950–1000 m, 25 May 1968, Beusekom & Phengkhlai 1002 (BKF SN060906, C).
Nan: Pua, Doi Phukha, 1700 m, 26 June 2008, Pooma & Tamura 7113 (BKF SN188552).
Phang-Nga: Khao Pawta Luang Keow, 900–1000 m, Geesink et al. 7690 (BKF SN060902).
Phetchaburi: Kaeng Krachan, Panoen Thung Ranger Substation, 12°52'12"N, 99°22'12"E, 123 m, 26 January 2005, Williams et al. 1112 (K), summit of Khao Phanoen Thung, 12°52'5"N, 99°22'20"E, 1240 m, 8 May 2005, Middleton et al. 3271 (BKF SN168267, E 00226065).
Phitsanulok: Chat Trakan, Phu Soi Dao, 1600 m, 14 August 2000, Suksanthan 2684 (QBG 19436); Nakhon Thai, on the way to the headquarters of Phu Hin Rong Kla National Park, 16°57'5"N, 101°1'24"E, 1600 m, 9 November 2015, Zhao et al. 71 (BKF, TCD).
Prachin Buri: ca. 1000 m, 4 July 1924, Kerr 10824 (BK 203739); Khao Yai, at the Heo Suwat Falls, 750 m, 8 July 1966, Larsen et al. 99 (C).
Ranong: 50 m, 17 November 1973, Santisuk 597 (C); Muang Len, 160 m, 11 January 1966, Hansen & Smitinand 11909 (BKF SN060905, C); Suk Samran, Khlong Na kha, ca. 9°45'N, 98°40'E, 50 m, 22 June 1974, Geesink et al. 7381 (K) & 7404 (C, K).
Sakon Nakhon: Phu Phan, 19 November 1962, Suwanakoset 1916 (BKF, K), ca. 30 km SW of Sakonnakhon city, ca. 380 m, 13 November 1984, Murata et al. T-48771 (BKF SN060859), Huay Yai waterfall, 22 June 2003, Chantaranothai et al. s.n. (KKU); Sawang Daen Din, Ban Thon, 1 December 1962, Adisai 194 (BK 203726).
Si Sa Ket: Kantharalak, Phanom Dongrak, 200 m, 25 November 2005, Suddee et al. 2632 (BKF SN181662, QBG 29057), Sao Thongchai Phulaor Falls, Phanom Dongrak WS, trail to waterfalls, 200 m, 22 December 2005, Pooma et al. 6068 (BKF SN183620, K).
Surin: Mueang Surin, Arloor-Doonban Community Forest, 124 m, 3 February 2008, Petrmitr 840 (CMUB 29794).
Udon Thani: Na Yung, Phu Luang, 1050–1300 m, 8 January 1966, Hennipman 3556 (BKF SN060870, C, K).
Uttaradit: Nam Pad, Phu Soi Dao, 1613 m, 17 November 2009, Norsaengsri & Intamusik 6145 (QBG 42661).
≡ Thea confusa Craib, Bull. Misc. Inform. Kew (1): 5 1914. Lectotype (first-step designated by
Trees or shrubs up to 12 m tall. New branchlets puberulous; terminal buds puberulous or glabrous. Petioles 5–15 mm long, puberulous; leaf blades elliptic, obovate, oblong or ligulate, 5.5–13 × 2–4.5 cm, coriaceous, abaxially yellowish green and glabrous, adaxially dark green, hirsute along midrib, midrib abaxially elevated and adaxially slightly impressed or flat, secondary veins 7–11 on each side of midrib, flat or obscure on both surfaces, base attenuate, cuneate or rounded, margin sparsely serrate, apex attenuate, acute, acuminate or caudate. Flowers solitary, 4.5–10 cm in diam., subsessile. Bracteoles and sepals 6–8, caducous, crescent to broadly ovate, 1.5–20 × 3–10 mm, abaxially glabrous, puberulous or pubescent, adaxially glabrous, margin ciliolate. Petals 6–8 in 1–2 whorls, white, elliptic or obovate, 2.5–5.5 × 1.5–4 cm, abaxially glabrous, puberulous or pubescent at apex, adaxially glabrous, apex bilobed. Stamens numerous, 1–1.5 cm long; filaments yellow, glabrous, basally connate for 2–3 mm. Ovary globose, pubescent. Styles 3(–4), basally connate or distinct, 8–12 mm long, basally pubescent and gradually becoming glabrous apically. Capsule globose or oblate, 2.5–6 cm in diam., 2–5 cm in height, 1–3-loculed with 1–3 seeds; pericarp 2–8 mm thick. Seeds brown or fuscous, hemispherical, polyhedral or globose, 1.5–2.5 cm in diam., glabrous Fig.
Flowering October–January, fruiting December–September.
Camellia kissi var. confusa is distributed in the evergreen forest at elevations of 350–1700 m in China, Laos, Myanmar, Thailand (Fig.
Chiang Mai: Chom Thong, Ban Pha Mon, 900 m, 31 May 1979, Vidal & Vidal 6247C (K); Doi Inthanon, 18°40'N, 98°25'E, 1400 m, 3 January 1975, Geesink et al. 8045 (BKF SN060804, C), 1680 m, 19 December 1983, Fukuoka & Ito T-35316 (BKF SN060813), 1700 m, 18 November 1998, Hara et al. C002 (BKF SN173305) & C044 (BKF SN173303), road to Khun Wang, 1310 m, 24 October 2000, Chayamarit et al. 2204 (BKF SN173163); Fang, Doi Ang Khang, 1490 m, 8 December 1934, Garrett 905 (BKF SN060834, E 00681092, K); Mae Cham, Mae Sanga, 1300 m, 6 May 2000, Sangnin & Sillapasuwan 3346 (BKF SN138882), between 34 and 35 km, on the road to Mae Aum Watershed Management Station, 18°30'30"N, 98°30'19"E, 1636 m, 12 November 2015, Zhao et al. 83 (BKF, KUN, TCD); Mueang Chiang Mai, Doi Suthep, ca. 1600 m, 22 December 1920, Kerr 4680 (BK 203745), 1300 m, 20 February 1959, Sørensen et al. 6967 (BKF SN060827, C, K), ca. 900 m, 17 November 1922, Kerr 6676 (BK 203730, BM, K) & 6676A (BK 203741, BM, E 00681089, K), 1050 m, 7 November 1987, Maxwell 87-1379 (BKF SN060825); east side of Doi Pui at the Chang Kian Agricultural Station, 1350 m, 24 November 1988, Maxwell 87-1351 (BKF SN060851); Samoeng, Samoeng Tai, Ban Pa Kar, 1424 m, 24 June 2008, Jatupol 08-233 (QBG 35892).
Loei: Phu Kradueng, top area of Phu Kradueng, 1200–1300 m, 19 December 1982, Koyama et al. T-31384 (BKF SN060872); Phu Ruea, 13 December 1966, Umpai 311 (BK 203736); Phu Luang, 17°17'2"N, 101°31'8"E, 1440 m, 3 November 2015, Zhao et al. 26, 27, 32 (BKF, KUN, TCD) & 33 (TCD); Wang Saphung, Phu Luang, 900–1000 m, 15 November 1968, Chermsirivathana 1057 (BK 203722); 1410 m, 13 May 1998, Chayamarit et al. 1357 (BKF).
Nan: Pua, Doi Phu Kha, 1100 m, 17 August 1995, Pooma 1104 (BKF SN167083), 19°13'N, 101°5'E, 1500 m, 8 October 1998, Srisanga 293 (QBG 12268) & 294 (QBG 12269).
Sakon Nakhon: Phu Phan, ca. 17°N, 104°E, 380–450 m, 12 November 1984, Murata et al. T-51155 (BKF SN115022); Huai Lub Num Lud, 9 March 1996, Puudjaa 195 (BKF SN091534).
Sukhothai: Khiri Mat, Kao Luang, ca. 1000 m, 2 May 1922, Kerr 5917 (BK 203747).
The variety usually bears a larger flower and a thicker pericarp than C. kissi var. kissi (Figs
≡ Thea laotica Gagnep., Notul. Syst. (Paris) 10: 128. 1942. Lectotype (first-step designated by
Shrubs up to 5 m tall. New branchlets puberulous; terminal buds pubescent. Petioles 2–5 mm long, puberulous; leaf blades elliptic to oblong, 4–9 × 1–4.3 cm, thinly coriaceous, abaxially sparsely appressed pubescent, especially along midrib, adaxially glabrous, midrib and secondary veins abaxially elevated and adaxially channelled, secondary veins 6–8 on each side of midrib, base attenuate, margin sparsely serrulate, apex attenuate to shortly caudate. Flowers solitary or up to 3 in a cluster. Pedicels 7–14 mm long, glabrous or puberulous at base. Bracteoles 2, opposite or alternate, caducous. Sepals 3–4, persistent, sub-orbicular or broadly ovate, 3–8 × 6–9 mm, abaxially glabrous, adaxially sericeous, margin ciliolate. Petals 5, white, broadly ovate to obovate, 8–12 × 5–7 mm, glabrous on both surfaces, margin ciliolate. Stamens numerous, 5–7 mm long; filaments glabrous. Ovary ovoid, glabrous. Style 1, 5–7 mm long, glabrous, apically 3–5-lobed for 1–2 mm. Capsule globose, 12–20 mm in diam.; pericarp 2–3 mm thick. Seeds glabrous when immature Fig.
Flowering October, fruiting April–August.
Camellia laotica is distributed in the evergreen or mixed deciduous forests at elevations of 400–750 m in Laos and Thailand (Fig.
Ubon Ratchathani: Khong Chiam, Ban Na Klang Pho, Dong Na Tham forest, ca. 400 m, 24 October 1998, Wongprasert s.n. (BKF SN121590, BKF SN121591; Fig.
Camellia laotica is a new record to Thailand. It is represented by a single collection, Wongprasert s.n. at BKF (Fig.
≡ Thea assamica Royle ex Hook., Kew Gardens 28. 1847. Neotype (designated by
= Thea cochinchinensis Lour., Fl. Cochinch. 1: 338. 1790. Neotype (designated by
= Camellia theifera Griff., Trans. Agric. Soc. India. 5: t. C. 1838. Lectotype (designated by
= Thea yersinii A. Chev. ex Gagnep., Fl. Indo-Chine [P.H. Lecomte et al.], Suppl. 1: 310. 1943 (“Thea yersini”). Lectotype (designated by
= Camellia multisepala Hung T. Chang & Y.J. Tan, Acta Sci. Nat. Univ. Sunyatseni 23(1): 11. 1984. Holotype: China. Yunnan: Mengla, Xiangming, cultivated, 1050 m, 3 December 1982, Y.J. Tan & S.C. Ma A31002 (SYS 00095167!).
= Camellia polyneura Hung T. Chang & Y.J. Tan, Acta Sci. Nat. Univ. Sunyatseni 23(1): 10. 1984. Holotype: China. Yunnan: Luchun, Qimaba, in tea garden, 1400 m, 18 November 1982, Y.J. Tan & S.C. Ma A26001 (SYS 00090671!).
= Camellia sinensis var. kucha Hung T. Chang & Ping S. Wang, Acta Sci. Nat. Univ. Sunyatseni 23(1): 10. 1984. Holotype: China. Yunnan: Jinping, Tongchang, 1371 m, 11 November 1982, B.H. Chen & Y.J. Yang A22003 (SYS 00095188!).
= Camellia tenuistipa Orel, Curry & Luu, Pursuit Hidden Camellias Vietnam China: 263. 2015. Holotype: Vietnam. Gia Lai: Kon Ka Kinh National Park, 22 Jan 2011, H.T. Luu & Q.D. Nguyen KKK 221 (NSW 901734, image!).
Trees or shrubs up to 20 m tall. New branchlets puberulous or pubescent, terminal buds pubescent or puberulous. Petioles 2–9 mm long, puberulous; leaf blades elliptic, oblong or obovate, 8–29 × 3.5–10 cm, coriaceous, abaxially yellowish green, puberulous, especially along midrib, adaxially dark green, shiny, midrib abaxially elevated and adaxially slightly raised or flat, secondary veins 7–15 on each side of midrib, abaxially slightly elevated and adaxially slightly impressed or flat, base cuneate or attenuate, margin serrulate, apex attenuate or acuminate. Flowers solitary or up to 4 in a cluster, 2–3.5 cm in diam. Pedicels 4–14 mm long, slender or gradually swollen towards the top, bracteoles 2–3, alternate, caducous. Sepals 5, persistent, sub-orbicular or broadly ovate, 3–4.5 × 3–5 mm, abaxially glabrous, adaxially sericeous, margin ciliolate. Petals 5–7 in 1–2 whorls, white or outer 1–3 petals with a tinge of green at apex, obovate, elliptic or rounded, 1–2.5 × 1–2 cm, glabrous on both surfaces. Stamens numerous in 3–5 whorls, 7–16 mm long, filaments pale yellow, glabrous, outer filaments basally adnate to petals for 1–3 mm. Ovary oblate or globose, pubescent. Style 1, 6–15 mm long, glabrous or gradually becoming glabrous upwards, apically (2–)3(–4)-lobed for 1–3 mm. Capsule oblate, bi-coccal or globose, 1.5–4.5 cm in diam., 1–2 cm in height, 1–3-loculed with 1–3 seeds; pericarp 0.5–1.5 mm thick. Seeds brown or fuscous, globose, hemispherical or polyhedral, 1.2–2 cm in diam., glabrous Fig.
Camellia sinensis var. assamica A habitat B branchlets with flowers and flower buds, adaxial surface of leaves C branchlets with flower and fruit, abaxial surface of leaves D a dissected flower E a flower without petals, androecium and a half sepal. Scale bars: 3 cm (B); 5 mm (E). The minimum graduation of the rulers in C, D represents 1 mm.
Flowering September–January, fruiting February–December.
Camellia sinensis var. assamica occurs in the evergreen and semi-deciduous forests at the elevations of 200–2700 m in China, India, Laos, Myanmar, Thailand (Fig.
Chiang Mai: Chiang Dao, 25 November 1963, Bunchuai 1359 (BKF SN060849, C, K); Khun Awn, 900 m, 30 January 1921, Kerr 4726 (BK 203758, BM, K); Mae On, Huai Kaeo, trail along Mae Kampong Waterfall, 1100 m, 7 September 2011, Pooma et al. 7791 (BKF SN196689); Mae On, Jae Son, 18°51'20"N, 99°22'1"E, 1500 m, 28 May 2011, Pooma & Pattharahirantricin 7729 (BKF SN196803); Mae Taeng, Mae Taman, 1300 m, 27 September 1994, Nanakorn et al. 1808 (QBG 1808); Mueang Chiang Mai, Doi Suthep, 1050 m, 7 November 1987, Maxwell 87-1378 (BKF SN060836), 15 July 1988, Maxwell 88-875 (BKF SN209981), 1 June 1993, Maxwell 93-522 (BKF SN169943, BKF SN192202, CMUB 02747), 18 September 1995, Kopachon s169b1 (BM, CMUB 07853).
Chiang Rai: Wiang Kaen, 600 m, 26 December 2009, Pongamornkul 2806 (QBG 51200).
Loei: Dan Sai, Pu Lom Lo, 1500 m, 8 April 1922, Kerr 5780 (BK 203759, BM, K); Phu Kradueng, 16°53'–54'N, 101°47'–49'E, 1150–1250 m, November 1984, Murata et al. T-42519 (BKF SN060857) & T-42796 (BKF SN060856), 1160–1180 m, 2 September 1988, Fukuoka T-63807 (BKF SN103447); Phurea, Phuluang, 17°18'N, 101°30'E, 1070–1100 m, 4 November 2015, Zhao et al. 37–41 (BKF, TCD); Na Haeo, Phu Suan Sai, 10 February 2004, Pornpongrungrueng 441 (KKU), 1300 m,16 May 2006, Maknoi 850 (QBG 27664), 14 May 2008, Maknoi & Srisanga 2185 (BKF SN190334, QBG 38413), 2 September 2008, Maknoi 2738 (BKF SN191010, QBG 40367), 3 September 2008, Maknoi 2806 (BKF SN188351, QBG 40435), 17°30'N, 100°56'E, 1300–1330 m, 6 November 2015, Zhao et al. 49, 50, 52 & 54 (BKF, TCD), Zhao et al. 51 & 53 (TCD).
Mae Hong Son: Mae Sariang, Wat Chan, Ban Den, 988 m, 29 April 2014, Norsaengsri 10930 (QBG 77549).
Nakhon Sawan: Khao Pado, 4 June 1922, Kerr s.n. (BK 203756).
Nan: Tha Wang Pha, 19°04'N, 100°40'E, 1100 m, 23 July 1992, Larsen et al. 43513 (BKF); Song Khwae, Tham Sokoen, 19°23'3"N, 100°31'5"E, 1210 m, 30 November 2011, Laongsri et al. 1957 (QBG 56767).
Phayao: Mueang Phayao, Doi Luang, 625 m, 19 November 1997, Petrmitr 148 (CMUB 13047).
Phitsanulok: Chat Trakan, Phu Miang-Phu Thong, 800 m, 18 February 2010, Romklao Botanical Garden 0005/2553 (QBG 59399); Nakhon Thai, Phu Hin Rong Kla, 16°59'38"N, 101°0'9"E, 1300 m, Zhao et al. 72 (BKF, TCD); Phu Rom Rot, one of the peaks of Phu Miang, 1200–1600 m, 3 October 1967, Shimizu et al. T-11514 (BKF SN060844, K) & T-11515 (BKF SN060845), 1200–1650 m, 4 October 1967, Shimizu et al. T-11653 (BKF SN060843, K).
The nomenclature of C. sinensis var. assamica has been clarified in recent studies (
As an important resource of commercial tea, Assam tea is widely cultivated in the tropical areas of the world. Local Thai people collected natural seeds or seedlings and planted them around houses to use the new branchlets or leaves as a vegetable or fermented beverage (
Holotype : Thailand. Nakhon Phanom: Ban Phaeng, Phu Lang Ka NP, trail to hill top, 17°58'58"N, 104°7'38"E, 373 m, 23 October 2015, S. Suddee et al. 4981 (BKF SN235114!).
Shrubs or trees up to 10 m tall. New branchlets pubescent to villous, terminal buds puberulous. Petioles 2–10 mm long, pubescent or puberulous; leaf blades elliptic or oblong, 3.5–14 × 1.3–5.5 cm, coriaceous, abaxially dull green, sparsely appressed puberulous especially along midrib, or glabrous, adaxially dark green, glabrous or hirsute along midrib, midrib and secondary veins abaxially elevated and adaxially impressed, secondary veins 5–8 pairs, base attenuate, margin serrulate, apex acute, attenuate to caudate. Flowers axillary, solitary or paired, 1.5–3.5 cm in diam. Pedicels ca. 2 mm long. Bracteoles 3–6, persistent, deltate to semi-orbicular, 1–3 × 1.5–4 mm, abaxially glabrous or puberulous at apex, adaxially glabrous to slightly sericeous, margin ciliolate. Sepals 5–7, persistent, suborbicular, 2–5 × 4–5 mm, abaxially glabrous, adaxially sericeous. Petals 6–8 in 1–2 whorls, white, elliptic to obovate, 7–18 × 4–9 mm, abaxially glabrous, adaxially glabrous or sericeous at apex, apex obtuse to rounded, inner 4–5 petals basally adnate to filament whorl for ca. 2 mm. Stamens numerous, 6–10 mm long; filaments white, glabrous, outer filaments basally connate for 2–3 mm. Ovary globose to ovoid, glabrous. Styles 3(–4), distinct, 3–9 mm long, glabrous. Capsule globose or bi-coccal, 2.5–4.5 cm in diam., 1–2-loculed with 1–2 seeds; pericarp 0.5–2 mm thick, smooth or furfuraceous. Seeds light brown to fuscous, globose, 1.5–2.5 cm in diam., glabrous Fig.
Flowering October–November, fruiting April–July.
Camellia suddeeana is native to Thailand (Fig.
Loei: Ban Na Luang, 300 m, 20 November 1968, Chermsirivathana 1185 (BK 203761, BKF SN060858). Phu Luang, ca. 1200 m, 23 November 1957, Dee 1025 (BKF SN060875).
Mae Hong Son: Khun Youm, 8 April 1977, Nimanong & Phusomsaeng 1816 (BKF).
Nong Khai: Bueng Khla, Phu Wua Wildlife Sanctuary, trail from Pha Nang Khoi to Tham Noi, 18°14'14"N, 103°57'38"E, 350 m, 30 July 2008, Pooma et al. 7333 (BKF SN191784; Fig.
Phetchabun: Wang Thong, Thung Salaeng Luang National Park, 700 m, Koyama et al. T-31930 (BKF SN060873).
Pitsanulok: Nahaew to Romklao Rd, 700 m, Pooma 1237 (BKF SN090320).
Uttaradit: Nampad, Thud Phra Kiet National Park, ca. 650 m, 9 December 1994, Santisuk et al. s.n. (BKF SN109856).
A specimen of C. suddeeana, Chermsirivathana 1185 at BKF, were misidentified as C. tenii by Keng in 1970. Camellia tenii bears a smaller leaf (2.5–4.2 × 1.2–2.3 cm), villous ovary and pubescent seeds, whereas C. suddeeana has a larger leaf (3.5–14 × 1.3–5.5 cm), glabrous ovary and seeds (Fig.
≡ Thea taliensis W.W. Sm., Notes Roy. Bot. Gard. Edinburgh 10: 73. 1917. Lectotype (first-step designated by
= Polyspora yunnanensis Hu, Bull. Fan Mem. Inst. Biol. Bot. 8: 135. 1938. Lectotype (designated by
= Camellia irrawadiensis Barua, Camellian 7(4): 18. 1956. Holotype: Myanmar. Raised from seed collected by L.O. Wilson, 1917, presumably in the region 26°–27°N, 98°–99°E (valley of Irrawadi in North Burma), January 1956, Ex. Herb. I.T.A. 3253 (consisting of 2 sheets: K 000704313! & K 000704314!).
= Camellia pentastyla Hung T. Chang, Acta Sci. Nat. Univ. Sunyatseni 20(1): 92. 1981. Lectotype (designated by
= Camellia quinquebracteata Hung T. Chang & C.X. Ye, Acta Sci. Nat. Univ. Sunyatseni 26(1): 20. 1987. Holotype: China. Yunnan: Lianghe, Dachang, 4 January 1983, P. Zeng & Q.J. Xie 17055 (SYS, isotype: KUN 1206063!).
Trees or shrubs up to 15 m tall. New branchlets glabrous, terminal buds glabrous or sparsely puberulous. Petioles 4–8 mm long, glabrous; leaf blades elliptic to oblong, 7.5–15.5 × 3–6.5 cm, coriaceous, abaxially yellowish green and adaxially dark or yellowish green, shiny and glabrous on both surfaces, midrib abaxially elevated and adaxially slightly raised, secondary veins 8–14 on each side of midrib, slightly elevated on both surfaces, base attenuate to obtuse, margin sparsely serrulate to nearly entire, apex attenuate or acute. Flowers solitary or up to 3 in a cluster, 3–5 cm in diam. Pedicels 8–15 mm long. Bracteoles 2–4, alternate, 2-ranked, caducous. Sepals 5, persistent, sub-orbicular or broadly ovate, 4–6.5 × 5.5–9 mm, abaxially glabrous, adaxially sericeous, margin ciliolate. Petals 7–11 in 2–3 whorls, white, elliptic to obovate, 16–30 × 10–21 mm, glabrous on both surfaces or inside sericeous at apex, inner petals basally adnate to filaments for 2–4 mm. Stamens numerous, 1–2.5 cm long; filaments pale yellow, glabrous. Ovary oblate, pubescent. Style 1, 11–20 mm long, basally sparsely pubescent and gradually becoming glabrous apically, apically (3–)5-lobed for 2–5 mm. Capsule oblate, 2.5–5 cm in diam., 2–3 cm in height,1–5-loculed with 1–5 seeds; pericarp 1.5–4 mm thick. Seeds brown, hemispherical or polyhedral, 1.5–2 cm in diam., glabrous Fig.
Flowering October–February, fruiting April–November.
Camellia taliensis occurs in the montane evergreen forest at the elevations of 1100–2750 m in China, Myanmar and Thailand (Fig.
Chiang Mai: Chiang Dao, Doi Chiang Dao, Den Ya Khat, 1500 m, 4 August 2007, Watthana 2420 (QBG 30728); Chom Thong, Doi Inthanon, 1900 m, 1 May 1921, Kerr 5298 (BK 203709, BM, K), 18°40'N, 98°25'E, 1700 m, 11 December 1969, Beusekom & Phengklai 2462 (BKF SN061415, P 06838120), 1900–2025 m, 14 January 1994, Fukuoka T-62168 (BKF SN102755); Fang, 1250 m, 24 February 1958, Smitinand 4331 (BKF SN060803) & 4331A ((BKF SN060831; Fig.
Kampaeng Phet: Klong Lam, Mae Wong, 1320 m, 30 October 2002, Bult 609 (CMUB 20961).
Mae Hong Son: Khun Yuam, 20 November 1998, Chusie KY308 (QBG 47303); Mae La Noi, Ban Dong, 1300 m, 15 December 2002, Pongamornkul 1871 (QBG 35067), 2 May 2011, Pongamornkul 2899 (QBG 64350); Mueang Mae Hong Son, along the trail to the peak of Doi Pui, 1100–1750 m, 16 December 2007, Tanaka et al. HN8479 (QBG 35641); Doi Khun Huai Pong, 18°58'N, 98°10'E, 1850 m, 3 March 1968, Hansen & Smitinand 12770 (BKF SN060846, C, E 00681068, K, P 04511721); Pai, en route to Doi Chang, 1800 m, 31 May 1972, Santisuk 149 (BKF SN060809); Doi Mae Ya, 1800 m, 3 November 1999, Suksathan 2030 (QBG 16229).
Nan: Pua, Doi Phu Kha, 19°17'N, 101°7'E, 1680 m, 10 April 1999, Srisanga & Watthana 685 (QBG 14019), 19°10'N, 101°7'E, 1700 m, 26 May 2000, Srisanga 1429 (QBG 17650), 1600 m, 22 August 2001, Srisanga & Maknoi 2027 (QBG 21153).
Phayao: Muang, Doi Luang, 19°5'N, 99°27'E, 1500 m, 22 April 1998, Sidisunthorn & Gardner 2568.0 (CMUB 13844); Phu Sang, Doi Pha Mon Noi, 19°44'8"N, 100°24'21"E, 1482 m, 30 April 2013, Laongsri et al. 2849 (QBG 66589); Pong, Phu Lanka, 1500 m, 4 September 2006, Watthana & Pumicong 2141 (QBG 28151).
Tak: Umpang, Umpang, near top of Khao Kheeo, 2150 m, 25 April 2001, Bult 419 (CMUB 18159).
Besides Assam tea, C. taliensis is another tea source plant that occurs in Thailand.
Among nine taxa–seven species and two varieties–of Camellia in Thailand, only two species are endemic to the country: C. connata and C. suddeeana. As stated above, C. pleurocarpa and C. tenii in
Most taxa of Camellia occur in northern and north-eastern Thailand (Fig.
I thank the staff of herbaria listed in the Materials and methods section for making available their specimens. A special thanks goes to the staff of herbaria BKF and KKU, whose help was vital to my investigations in Thailand.
The author has declared that no competing interests exist.
No ethical statement was reported.
This work is supported by the Hunan Provincial Natural Science Foundation of China (2022JJ40870), the Guangxi Key Laboratory of Special Non-wood Forest Cultivation & Utilization (JB-23-01-01), and the Scientific Research Foundation of the Central South University of Forestry and Technology (2019YJ023).
The author solely contributed to this work.
Dongwei Zhao https://orcid.org/0000-0002-7761-7127
All of the data that support the findings of this study are available in the main text.