Research Article
Print
Research Article
Notes on the genus Syzygium (Myrtaceae) from Cambodia, Thailand, Vietnam, China and Taiwan
expand article infoPranom Chantaranothai§
‡ Khon Kaen University, Khon Kaen, Thailand
§ Royal Botanic Gardens, Richmond, United Kingdom
Open Access

Abstract

Syzygium kampotense is a new species from Cambodia. Syzygium cerasiforme, S. foxworthianum, and S. angkae and S. thorelii are new records from Cambodia, Vietnam and China, respectively. Syzygium prainianum subsp. minor and S. densinervium var. insulare are placed under S. prainianum and S. densinervium, respectively. Eugenia cochinchinensis, E. eburnea and E. ripicola, are reduced to synonymy under S. pellucidum. Lectotypes for Eugenia densiflora var. angustifolia, E. pellucida and E. pellucida var. contracta are designated.

Key words

Lectotype, Myrtales, new record, new species, synonym, taxonomy

Introduction

Syzygium Gaertn. is the largest genus in the family Myrtaceae and many species from allied genera, namely Acmena DC. and Cleistocalyx Blume, are now included in Syzygium based on molecular studies. Thus Syzygium currently comprises ca. 1,200−1,500 species (Biffin et al. 2006; Craven and Biffin 2010), the majority of which occur in the Old World tropics and subtropics (Biffin et al. 2006). A comprehensive revision of the genus in Thailand was published by Parnell and Chantaranothai (2002). Since then, the number of taxa recorded in Thailand has increased through the discovery of new species and new country records (Chantaranothai 2014; Soh and Parnell 2015; Chantaranothai et al. 2016; Tagane et al. 2018). Similar discoveries have been made in China (Chen and Craven 2007) and Indochina (Soh and Parnell 2015).

During my visits to herbaria in Asia and Europe between 2017 and 2023, many unidentified specimens of Syzygium from Thailand and neighbouring countries were examined. Material of an unidentified taxon from Cambodia was found to represent a species new to science, which is described below. Previously unidentified specimens of S. cerasiforme (Blume) Merr. & L.M.Perry from Cambodia, S. foxworthianum (Ridl.) Merr. & L.M.Perry from Vietnam, S. angkae (Craib) Chantar. & J.Parn. and S. thorelii (Gagnep.) Merr. & L.M.Perry from China are newly recorded. Syzygium prainianum (King) Chantar. & J.Parn. subsp. minor Chantar. & J.Parn. and S. densinervium (Merr.) Merr. var. insulare C.E.Chang are placed under S. prianianum and S. densinervium, respectively. Eugenia cochinchinensis Gagnep., E. eburnea Gagnep. and E. ripicola Craib (S. ripicola (Craib) Merr. & L.M.Perry) are reduced to synonymy under S. pellucidum (Duthie) N.P.Balakr. Lectotypes are designated for E. densiflora (Blume) Duthie var. angustifolia Ridl., E. pellucida Duthie and E. pellucida var. contracta Wall. ex Duthie.

Material and methods

This study is based on both herbarium and field collections in Thailand. Herbarium material was also consulted in the following herbaria: AAU, BK, BKF, BM, K, KKU, KYO, P and QBG (herbarium acronyms following Thiers, updated continuously). Specimens were examined with a binocular microscope and via digital images on the JSTOR website (https://plants.jstor.org/). An illustration of the new species was prepared.

Taxonomic treatment

Syzygium kampotense Chantar., sp. nov.

Figs 1, 2, 3

Diagnosis

Syzygium kampotense resembles S. championii (Benth.) Merr. & L.M.Perry and S. claviflorum (Roxb.) Wall. ex Steud., in the clavate hypanthium shape, cuneate leaf base, small leaves and short petioles. The new species differs from both species in having thickly coriaceous leaves with strongly revolute leaf margins, an acute or obtuse leaf apex and fewer secondary veins (Fig. 1, Table 1).

Figure 1. 

Syzygium kampotense Chantar.: holotype, McDonal et al. 5676 (AAU). In set leaf, hypanthial cups and fruit. Photographed by B. Boonsuk.

Table 1.

Morphological comparison of S. kampotense and similar species.

Characters S. championii S. claviflorum S. kampotense
Petiole (mm) 1.5–2 3–6 2–3
Leaf texture coriaceous chartaceous to coriaceous thickly coriaceous
Leaf shape elliptic to oblong elliptic, ovate, sometimes obovate elliptic or elliptic-oblong
Leaf size (cm) 4–8.5 × 1–4 6–10 (–22) × 1.5–3.5(–7.5) 4–6.5 × 1.7–3
Leaf base cuneate, slightly attenuate cuneate, slightly attenuate broadly cuneate
Leaf apex acuminate with acumen mostly acute without acumen, sometimes acuminate with distinct acumen acute or obtuse
Leaf margin flat flat strongly revolute
Secondary veins (pairs) 25–30 15–30 ca 12
Hypanthium cup (mm) 10–13 5–10 13–15
Sepal (mm) 0.5 × 2 0.5–1 × 0.8–2.5 ca. 1 × 1.5
Fruit (mm) oblongoid, clavate, 12.5 × 5 ellipsoid to obovoid, 1–15 × 5–9 globose or ellipsoid, 8–10 × 8–10

Type

Cambodia: Kampot Province, Kampot District, southern summit of Phnom Bokor, vicinity of Bokor, 1–2 km NE of artificial on near Church toward Prek Kaoh Toch, 13 May 1997, McDonal et al. 5676 (holotype AAU, isotype QBG).

Figure 2. 

Syzygium kampotense Chantar., showing leaf, hypanthial cups and fruit. Photographed by B. Boonsuk.

Description

Shrub ca. 1 m tall; branching sparingly, bark corky, whitish grey or reddish. Leaves with petiole 2–3 mm long, wrinkled; lamina thickly coriaceous, 4–6.5 × 1.7– 3 cm, elliptic or elliptic-oblong, base broadly cuneate, apex acute or obtuse, margin strongly revolute; midrib impressed on the upper surface, rounded with sparse pustules on the lower surface; secondary veins in ca. 12 pairs, indistinct on upper surface and distinct on lower surface; intramarginal vein 1. Inflorescence not seen. Hypanthial cup 13–15 mm long, clavate. Pseudostipe absent. Sepals red 4, ca. 1 × 1.5 mm, triangular. Petals not seen. Stamens numerous, white. Ovary 2-locular, ca. 9 ovules per locule. Fruit purple, 8–10 mm in diameter, globose or ellipsoid, crowned with remnant of calyx lobes.

Figure 3. 

Syzygium kampotense A habit B lower surface of leaf C calyx tube (side view) D longitudinal section of calyx tube and ovary, showing two locules E fruit (from McDonald et al. 5676; drawn by N. Triyutthachai).

Distribution

Endemic to Cambodia.

Ecology

Dense sclerophyllous, short-tree forest, canopy 2–4 m tall, some emergent to 7 m tall; associates include Ardisia, Lithocarpus, Elaeocarpus, Melastoma, Schima, Syzygium and Vaccinum.

Conservation status

The species is only known from the type locality. It should be categorised as Critically endangered [CR, B1ab (iii)] according to the IUCN Red List Criteria and Categories version 3.1 (IUCN 2012). The Extent of Occurrence is estimated to be less than 20 km2 and this species is found in a restricted area on open limestone hills which is a threatened ecosystem.

Etymology

The name of this species is given based on the location where the plant was collected.

Notes

The new species most likely belongs to Syzygium subgenus Perikion Craven & Biffin (Craven and Biffin 2010; Hatt et al. 2023, because of its clavate hypanthium shape and ellipsoid, obovoid or globose fruit. These characteristics resemble S. claviflorum, the type of this subgenus.

Additional specimens examined

Cambodia, Kampot Province, Bokor National Park, near field station, near top of plateau, 10°20.38'N, 104°01.4'E, alt. 1,045 m, 10 Mar. 2001, Midlleton & Monyrak 660 (P [P00589160]); Bokor, Dy Phon 1130 (P [P04658853]).

New Records

Syzygium cerasiforme (Blume) Merr. & L.M.Perry, Mem. Amer. Cad. Arts. 18: 187. 1939.

Myrtus cerasiformis Blume , Bijdr. Fl. Ind. Ned.: 1087. 1826. Type: Indonesia, Java, Blume s.n. (Isotype NY [NY00405548]).

Type

Based on Myrtus cerasiformis Blume.

Distribution

Thailand, Cambodia, Malaysia, Java, Borneo.

Ecology

In evergreen swamp forest and a drainage along river.

Notes

POWO (2023) accepts Syzygium lineatum as a synonym of S. cerasiforme and also shows the distribution of S. cerasiforme in Cambodia, Laos, Vietnam and China. I agree with Merrill and Perry (1939) that both species are not conspecific. Moreover, Parnell and Chantaranothai (2002), Chen and Craven (2007), and especially Soh and Parnell (2015) and Tagane et al. (2015, 2018) working on Cambodian taxa, indicated that Syzygium cerasiforme had never been recorded in Cambodia. Syzygium cerasiforme is similar to S. lineatum in having terminal and axillary inflorescences, leaf shape and 14 or more pairs of secondary veins. It differs by smaller leaves, sepals and petals and the outer stamens and style are shorter (4.3–6.7 mm vs 10–15 mm and 5.2–7.7 mm vs 11–13 mm, respectively). However, the Cambodian specimens were collected from Kampong Thom and Stung Treng provinces in Cambodia and I identified them as S. cerasiforme. Therefore, these two specimens are the first new record for Cambodia.

Specimens examined

Cambodia, Kampong Thom Province, Sandan District, Rey Long, ca. 13 km south-eastern of Spong, ‘Cheum Takong’ (Takong swam; 3 × 4 km), a drainage of O Long River, Base camp 13°20.27'N, 105°36.077'E, 7 Apr. 2008, McDonald et al. 7901 (AAU) & Stung Treng Province, Prey Long Forest, Cheum Takong and O Long River, 13°20.359'N, 105°36.32'E, 7–10 April 2009, McDonald et al. PL8 (AAU).

Syzygium foxworthianum (Ridl.) Merr. & L.M.Perry, Mem. Mer. Acad. Arts 18: 168. 1939.

Eugenia foxworthyi Ridl., Fl. Mal. Penins. 1: 728. 1922, non Elmer, 1912.

Eugenia foxworthiana Ridl., Fl. Mal. Penins. 5: 308. 1925. Type: Peninsular Malaysia, Pahang, Bukit Goh Reserve, 12 Jan. 1920, Foxworthy Field No. 3624 (lectotype, designated by Chantaranothai and Parnell 1994, p. 63: K [K001005521]).

Syzygium foxworthianum (Ridl.) Masam., Enum. Phan. Born.: 528. 1942.

Eugenia densiflora Miq. var. angustifolia Ridl., Fl. Mal. Penins. 1: 729. 1922. Type: Peninsular Malaysia, Perak, Ulu Temengoh, Ridley s.n. (lectotype, designated here: K [without barcode]).

Syzygium pycnanthum Merr. & L.M.Perry var. angustifolium (Ridl.) P.S.Ashton, Tree Fl Sabah & Sarawak 7: 270. 2011.

Type

Based on Eugena foxworthiana Ridl.

Distribution

Thailand, Peninsular Malaysia, Sumatra, Borneo, Vietnam (Lao Cai (Laokhay), Pakha, 10 Dec. 1935, Poilane 25052 (K).

Ecology

In evergreen forest, 50–200 m alt.

Notes

Ashton (2011) placed S. foxworthianum under S. pycnanthum (Eugenia densiflora Miq.) but I do not agree with this placement because the former differs from the latter by having a long and lax inflorescence (vs short and dense), slightly smaller and narrower leaves, 16–18.5 × 4–6.5 cm (vs larger and broader, 17.5–26 × 4.5–8.5 cm), two intramarginal veins, sometimes without the intramarginal vein and secondary veins ascending to a shallow loop (vs 2–3 intramarginal veins). Although the Vietnamese specimen Poilane 25052 (K), has young fruits, I found that it belongs to S. foxworthianum. Therefore, it is a new record of this species for Vietnam.

Eugenia densiflora var. angustifolia Ridl. was described based on two specimens from Peninsular Malaysia, Peak (Ridley s.n.) and Kelantan (Yapp s.n.). Ridley s.n. (K) is available and is designated here as the lectotype.

Syzygium angkae (Craib) Chantar. & J.Parn., Kew Bull. 48(3): 592. 1993.

Eugenia angkae Craib, Bull. Misc. Inform. Kew 1929: 115. 1929.

Type

Thailand, Doi Inthanon (Doi Angka), 30 April 1921, Kerr 5287 (lectotype, designated by Chantaranothai and Parnell 1994, p. 35: BK; isolectotypes: BM, K, TCD).

Distribution

Myanmar, Thailand, Laos, Cambodia, Vietnam and China (Yunnan province, Menghai county, Xiding country, Mt. Dahei, Hesong, 2 May 2011, Li-Jianwu 625 (HITBC no. 136852, QBG no. 64320, fruiting specimen).

Ecology

In evergreen broad-leaf forest, ca. 1,960 m alt.

Notes

Syzygium angkae is characterized by having axillary or terminal inflorescences ca. 2 cm long and numerous secondary veins. It is found on mountains at an altitude of at least 1,500 m or more. The unidentified fruiting specimen at QBG belongs to S. angkae. The distribution of the species was mainly in Myanmar, Thailand, Laos, and Vietnam; it is now extended to China.

Syzygium thorelii (Gagnep.) Merr. & L.M.Perry, J. Arnold Arbor. 19: 107. 1938.

Eugenia thorelii Gagnep. in Lecomte., Notul. Syst. (Paris) 3: 333. 1918.

Type

Thailand, Ubon Ratchathani, Kemmarat, Thorel 3010 (lectotype, designated by Soh and Parnell 2015, p. 261: P [P00589178]; isolectotypes: A [A00069448], P [P00589179], K [K000276196]).

Distribution

Thailand, Laos, Cambodia, Vietnam, China (Yunnan Province, Mengla County, Mt. Gongbeng, Luosuo River Estuary, 500 m alt., 3 Jan. 2011, Li-Jianwu 239 (HITBG, no. 135842, QBG).

Ecology

Open rock crevices in the Mekong River, partly submerged at high water or in sandy soils along the side of the river.

Notes

Syzygium thorelii is distinctive in having an obtuse or acute leaf apex and alternate leaves in the lower parts of the plant. A previously unidentified specimen seen at QBG undoubtedly belongs to S. thorelii and is a new record for China. The Chinese specimen extends the range of the species from Thailand, Laos, Cambodia and Vietnam to Yunnan.

New synonymy

Syzygium densinervium (Merr.) Merr., Phillip. J. Sci. 79: 387. 1951.

Eugenia densinevia Merr., Philipp. J. Sci. 1(Suppl.): 105. 1905. Type: Philippines, Luzon, Laguna province, Los Banos, Mt. Maquiling, Jun. 1917, Elmer 18011, (holotype K [K000800201]).

Eugenia silvestrei Elm, Leafl. Philipp. Bot. 8: 3095. 1919. Type: Philippines, Luzon Island, Los Baños, Mount Maquiling, July 1917, Elmer 18011 (holotype A [A00069784]).

Syzygium densinervium var. insulare C.E.Chang, Bull. Taiwan Prov. Pingtung Inst. Agri. 5: 52. 1964. Type: Taiwan, Botel Tobago, 17 Apr. 1962, Chang 2846, (isotype L [L0009615]), syn. nov.

Type

Based on Eugenia densinevia Merr.

Distribution

Philippines (Luzon) and Taiwan (South Cape).

Ecology

In evergreen forest, mountain slope.

Notes

Syzygium densinervium resembles S. fastigiatum (Blume) Merr. & L.M.Perry in having paniculate inflorescence, funnel-shaped hypanthium and persistent bracts and bracteoles. It differs by its rugulose hypanthial cup. I have examined an unidentified specimen, Henry 1998 at K [K001003761] and found that it belongs to S. densinervium. I have also examined the isotype type of S. densinervium var. insulare from Taiwan and then placed it under S. densinervium. This species is distributed in the Philippines and Taiwan.

Specimens examined

Taiwan, Botel Tobago, 7 Feb. 1980, Chang 14691 (KYO); ibid., 6 Sept. 1980, Chang 14694 (KYO).

Syzygium pellucidum (Duthie) N.P.Balakr., Bull. Bot. Surv. India 22(1–4): 14. 1982.

Eugenia pellucida Duthie, Fl. Brit. India 25(4): 485. 1878. Type: Tenasserim & Andamans, Helfer 2406 (lectotype, designated here: K [K000821333]; isolectotype A [GH00069438].

Eugenia pellucida var. contracta Wall. ex Duthie, Fl. Brit. India 25(4): 485. 1878. —Syzygium contractum Wall., nom.nud., non Eugenia contracta Poir., 1828. Type: Myanmar, Ataran [Attran] river, 1827, Wallich 3602 (lectotype, designated here: K-W [K001119797]; isolectotypes: K [K000821331, K000821332], A [A01143296]).

Eugenia ripicola Craib, Bull. Misc. Inform. Kew 1915(10): 428. 1915.— Syzygium ripicola (Craib) Merr. & L.M.Perry, Britt. 4: 127. 1941. Type: Thailand, Mae [Mê] Ping Rapids, Keng Soi, 16 Mar. 1913, Kerr 2944 (lectotype, designated by Chantaranothai and Parnell 1994, p. 104: ABD; isolectotypes: BM [BM000944095], E [E00284095], K [K000800078]), syn. nov.

Eugenia cochinchinensis Gagnep. in Lecomte, Notul. Syst. (Paris) 3: 324. 1918.— Syzygium cochinchinense (Gagnep.) Merr. & L.M.Perry, J. Arnold Arb. 19:107. 1938. Type: Cambodia, Kompong Speu, Samroang Tong, April 1870, Pierre 527 (lectotype, designated by Soh and Parnell 2015, p. 254: P [P00589286]; isolectotypes: P [P00589287, P00589288, P00589350], syn. nov.

Eugenia eburnea Gagnep. in Lecomte, Notul. Syst. (Paris) 3: 324. 1918.— Syzygium eburneum (Gagnep.) Merr. & L.M.Perry, J. Arnold Arb. 19:107. 1938. Type: Cambodia, plain of Pen-lovier, May 1870, Pierre 991 (lectotype, designated by Soh and Parnell 2015, p. 255: P [P00589209]; isolectotypes: P [P00589210, P00589211], K [K000276209], E [E00284602], syn. nov.

Type

Based on Eugenia pellucida Duthie

Distribution

India, Myanmar, Thailand, Vietnam.

Ecology

Along rivers or streams.

Notes

Based on three collections, Helfer 2406 (K000821333, GH00069438), Helfer 2407 and Kurz s.n., the last two have not been seen. Therefore, K000821333 is designated here as the lectotype of E. pellucida. The original description of E. pellucida var. contracta is based on Wallich 3602 which has four sheets. The specimen, K001119797 is designated here to be the lectotype because it has more leaves and inflorescences. Syzygium pellucidum was considered to be endemic to Myanmar (POWO 2023) but its distribution is now extended to Thailand, Laos and Vietnam.

Syzygium prainianum (King) Chantar. & J.Parn., Kew Bull. 48: 608. 1993.

Eugenia prainiana King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 70(1): 116. 1901. Type: Peninsular Malaysia, Perak, Blanta Mabok, Apr. 1890, Wray 3990 (lectotype, designated by Chantaranothai and Parnell 1994, p. 97: K).

Syzygium prainianum subsp. minor Chantar. & J.Parn., Kew Bull. 48: 608. 1993. Type: Thailand, Phangnga, Khao (Kao) Kata Kwam, 9 Mar. 1930, Kerr 18481 (holotype BM, isotypes BKF, K [K001007999]), syn. nov.

Type

Based on Eugenia prainiana King.

Distribution

Thailand, Peninsular Malaysia, Borneo.

Ecology

In evergreen forest, 900 m alt.

Notes

Chantaranothai and Parnell (1993) proposed subsp. minor for Thai material with a slightly shorter hypanthial cup (3–4 mm long vs 5 mm long for the typical variety), stamens and style. After examination of Gardner & Chamchumroon ST2467 K [001007729], a second new specimen collected from Trang, Thailand, I found that these characteristics are variable. Therefore, subsp. minor is here placed into synonymy of S. prainianum. This species is uncommon in Thailand but widespread in Peninsular Malaysia and Borneo.

Specimens examined

Trang, Yanta Khao, Khao Banthat Wildlife Sanctuary, valley above Sai Rung Waterfall, Camp 2, 800 m alt., 14 Mar. 2006, Gardner & Chamchumroon ST2467 (K).

Acknowledgements

The directors, curators and staff of the herbaria at AAU, BKF, BM, E, K, KKU, KYO, PSU and QBG are thanked for allowing access to their collections. Finally, I thank Wuu Kuang Soh and an anonymous reviewer for helpful comments, David Simpson for language revision, Boonchuang Boonsuk for the photograph and Natthawut Triyutthachai for the line drawing.

Additional information

Conflict of interest

The author has declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This research was supported by a grant (RP66-4-Research Center-001) from Thailand’s Research and Graduates Studies at Khon Kaen University. The Carlsberg Foundation, Denmark, is thanked for additional financial support.

Author contributions

The author solely contributed to this work.

Author ORCIDs

Pranom Chantaranothai https://orcid.org/0000-0002-5065-6169

Data availability

All of the data that support the findings of this study are available in the main text.

References

  • Ashton PS (2011) Myrtaceae s.l. In: Soepadmo E, Saw LG, Chung RCK, Kiew R (Eds) Tree flora of Sabah and Sarawak (Vol. 7). Sabah Forestry Department, Forest Research Institute Malaysia (FRIM), Sarawak Forestry Department, Malaysia, 87–423. https://doi.org/10.26525/TFSS7002
  • Biffin E, Craven LA, Crisp MD, Gadek PA (2006) Molecular systematics of Syzygium and allied genera (Myrtaceae): Evidence from the chloroplast genome. Taxon 55(1): 79–94. https://doi.org/10.2307/25065530
  • Chantaranothai P (2014) Three new records of Syzygium Gaertn. in Thailand and lectotypification of 19 taxa of Eugenia L. (Myrtaceae). Thai Forest Bulletin. Botany 42: 75–80.
  • Chantaranothai P, Parnell JAN (1993) New Taxa and Combinations in Cleistocalyx and Syzygium (Myrtaceae) in Thailand. Kew Bulletin 48(3): 589–610. https://doi.org/10.2307/4118723
  • Chantaranothai P, Parnell JAN (1994) A revision of Acmena, Cleistocalyx, Eugenia s.s. and Syzygium (Myrtaceae) in Thailand. Thai Forest Bulletin. Botany 21: 1–123.
  • Chen J, Craven LA (2007) Myrtaceae. In: Wu ZY, Raven PH, Hong DY (Eds) Flora of China (Vol. 13). Science Press, Beijing, and Missouri Botanical Garden Press, St. Louis, 321–359.
  • Elmer ADE (1912) Notes and descriptions of Eugenia. Leaflets of the Philippine Botany 4: 1399–1444.
  • Hatt S, Low YW, Burslem DFRP, Middleton DJ, Biffin E, Maurin O, Lucas EJ (2023) A morphological analysis of Syzygium, with a focus on fibre bundles and description of a new subgenus. Botanical Journal of the Linnean Society 20(1): 1–7. https://doi.org/10.1093/botlinnean/boac065
  • IUCN (2012) IUCN Red List Categories and Criteria: Version 3.1 (2nd edn). Gland and Cambridge, 32 pp.
  • Merrill ED, Perry LM (1939) The myrtaceous genus Syzygium Gaertner in Borneo. Memoirs of the American Academy of Arts and Sciences 18(3): 135–202. https://doi.org/10.2307/25058505
  • Parnell JAN, Chantaranothai P (2002) Myrtaceae. In: Santisuk T, Larsen K (Eds) Flora of Thailand (Vol. 7). The Forest Herbarium, Royal Forest Department, Bangkok, 778–914.
  • Poiret JLM (1828) Eugenia contracta. In: Lamarck JBAPM, Poitet JLM (Eds) Encyclopédie méthodique. Botanique. Supplement. A. Paris, Chez H. Agasse, Impimeur- Libraire, 125 pp.
  • Soh WK, Parnell JAN (2015) A revision of Syzygium Gaertn. (Myrtaceae) in Indochina (Cambodia, Laos and Vietnam). Adansonia 37(2): 179–275. https://doi.org/10.5252/a2015n2a1
  • Tagane S Van-son Dang, Souladeth P, Nagamasu H, Toyama H, Naiki A, Tran H, Yang Ceng-Jui, Prajaksood A, Yahara T (2018) Five new species of Syzygium (Myrtaceae) from Indochina and Thailand. Phytotaxa 375(4): 247–260. https://doi.org/10.11646/phytotaxa.375.4.1
  • Tagane S, Toyama H, Chhang P, Nagamasu H, Yahara T (2015) Flora of Bokor National Park, Cambodia I: Thirteen new species and one change in status. Acta Phytotaxonomica et Geobotanica 66(2): 95–135.
login to comment