Symplocos subgenus Hopea

Leaf and infructescence of Symplocos crassipes var. penangiana, from Nooteboom (1975).

Belongs within: Symplocaceae.
Contains: Symplocos cochinchinensis ssp. leptophylla.

Tropical sweetness
Published 18 November 2025

It is no secret that our awareness of the world’s biota is not proportionally distributed. The greatest concentration of biodiversity is in the tropics but, for various reasons (many to do with the distribution of wealth), popular representation and understanding is skewed towards the temperate zones. As a result, there are some major groups of organisms that one might struggle to label with so much as an English vernacular name. One such example is the tree genus Symplocos.

Symplocos wikstroemiifolia, copyright Jody Hsieh.

Symplocos is a genus of over three hundred trees and shrubs found in tropical montane regions of Asia and the Americas, with a very small number of species extending into temperate climes. Characteristic features of the genus include leaves without stipules, actinomorphic (radially symmetrical) flowers with numerous, epipetalous stamens and an inferior ovary, and drupaceous fruits with a hard endocarp (Wang et al. 2004). The affinities of Symplocos have long been uncertain, leading to its assignment to its own family, though recent authorities agree that it belongs within the order Ericales. Though no longer found west of India, fossils indicate that the genus was found in Europe for most of the Tertiary. Indeed, it dominated forests there during the Eocene; coal deposits have been identified from that period almost entirely composed of Symplocos endocarps (Nooteboom 2004). Symplocos disappeared from Europe with cooling climates at the beginning of the Pleistocene.

Common sweetleaf Symplocos tinctoria, copyright Amy Hill.

In a revision based on the Asian species, Nooteboom (1975) divided Symplocos between two subgenera: subgenus Symplocos had flowers with petals connate for at least half their length, whereas Hopea had petals connate only at the base. Leaves of Hopea species also tended to become yellow as they dried in herbarium specimens whereas leaves of subgenus Symplocos usually remained green. Both subgenera were found in both Asia and the Americas. However, more recent phylogenetic studies have lead to the genus being reorganised. Fritsch et al. (2008) separated the Asian representatives of subgenus Symplocos into a separate genus Cordyloblaste from the remaining Symplocaceae. Subgenus Hopea as recognised by Nooteboom turned out to be paraphyletic to the New World subgenus Symplocos, and was divided by Fritsch et al. (2008) between a series of subgenera and sections. The name Hopea (now a section of subgenus Symplocos) should now be applied to a smaller assemblage (about 35 species) with non-monadelphous stamens (i.e. filaments not fused), usually single-celled endocarps, and mostly glabrous nectary disks. Nevertheless, it remains applicable to species found in both Asia and the Americas.

Symplocos tinctoria in fruit, copyright Evan M. Raskin.

For the most part, Symplocos species are not of much economic significance to humans. A North American species of section Hopea, S. tinctoria, has historically been used to produce a yellow dye from the bark and leaves (hence the botanical name). Some Asian species are similarly used as dye mordants. The leaves of S. tinctoria are supposed to have a sweet taste, giving it the vernacular names of ‘sweetleaf’ or ‘horse-sugar’. It is the only Symplocos species found in more temperate climes of North America (the southeast United States), a lone hold-out while its relatives have retreated to the tropics.

Systematics of Symplocos subgenus Hopea
Symplocos subg. Hopea Clarke 1882 (see below for synonymy)N77
    |--S. adenophylla Wall. ex Don 1837 (see below for synonymy)N77
    |--S. anomala Brand 1900 [incl. S. concolor Brand 1901]N77
    |--S. atjehensis Noot. 1975N77
    |--S. barisanica Noot. 1975N77
    |--S. barringtoniifolia Brand 1904 (see below for synonymy)N77
    |--S. batakensis Noot. 1975N77
    |--S. borneensis Brand 1901N77
    |--S. brachybotrys Merr. 1917N77
    |--S. brandisii K. & V. 1900N77
    |    |--S. b. var. brandisii [incl. S. koordersiana Brand 1906]N77
    |    `--S. b. var. pseudoclethra (Hall. f.) Noot. 1975 [=S. pseudoclethra Hall. f. 1912]N77
    |--S. buxifolia Stapf 1894N77
    |--S. calycodactylos Brand 1901N77
    |--S. celastrifolia Griff. ex Clarke 1882 (see below for synonymy)N77
    |--S. cerasifolia Wall. ex DC. 1844 [=Bobua cerasifolia Miers 1879, Eugeniodes cerasifolium O. K. 1891]N77
    |    |--S. c. var. cerasifoliaN77
    |    `--S. c. var. grandifolia Noot. 1975N77
    |--S. cochinchinensis (Lour.) Moore 1914 [=Dicalix cochinchinensis Lour. 1790]N77
    |    |--S. c. ssp. cochinchinensisUB06
    |    |    |--S. c. ssp. c. var. cochinchinensis (see below for synonymy)N77
    |    |    |--S. c. ssp. c. var. imbricata (Brand) Noot. 1975 [=S. imbricata Brand 1909]N77
    |    |    |--S. c. ssp. c. var. philippinensis (Brand) Noot. 1975 (see below for synonymy)N77
    |    |    `--S. c. ssp. c. var. sessifolia (Bl.) Noot. 1975 (see below for synonymy)N77
    |    |--S. c. ssp. laurina (Retz.) Noot. 1876 (see below for synonymy)N77
    |    |--S. c. ssp. leptophyllaN77
    |    `--S. c. ssp. thwaitesii (F.v.M.) Noot. 1975 [=S. thwaitesii F.v.M. 1862]N77
    |         |--S. c. ssp. t. var. thwaitesiiN77
    |         |--S. c. ssp. t. var. montana nec S. montana Brongn. & Gris 1866 nec S. montana Vidal 1886 nec S. adenophylla var. montana Ridl. 1923N77
    |         `--S. c. ssp. t. var. stawellii (F.v.M.) Noot. 1975 (see below for synonymy)N77
    |--S. colombonensis Noot. 1975N77
    |--S. composiracemosa Noot. 1975N77
    |--S. costata (Bl.) Choisy 1854 (see below for synonymy)N77
    |--S. crassipes Clarke 1882N77
    |    |--S. c. var. crassipesN77
    |    |--S. c. var. brandiana (King & Gamble) Noot. 1975 [=S. brandiana King & Gamble 1906]N77
    |    |--S. c. var. curtisii (Oliv.) Noot. 1975 [=S. curtisii Oliv. 1888; incl. S. monticola King & Gamble 1906]N77
    |    |--S. c. var. ernae (Brand) Noot. 1975 [=S. ernae Brand 1901]N77
    |    |--S. c. var. havilandii (Brand) Noot. 1975 [=S. havilandi Brand 1901]N77
    |    |--S. c. var. penangiana (King & Gamble 1906) [=S. penangiana King & Gamble 1906]N77
    |    `--S. c. var. rufomarginata Noot. 1975N77
    |--S. cylindracea Noot. 1975N77
    |--S. deflexa Stapf 1894N77
    |--S. fasciculata Zoll. 1854 (see below for synonymy)N77
    |--S. filipes Noot. 1975N77
    |--S. gambliana Brand 1921 [incl. S. havilandii King & Gamble 1906 non Brand 1901]N77
    |--S. gigantifolia Noot. 1975N77
    |--S. glabriramifera Noot. 1975N77
    |--S. glomerata King ex Clarke 1882N77
    |--S. goodeniacea Noot. 1975N77
    |--S. herzogii Sleum. 1937N77
    |--S. johniana Stapf 1894N77
    |--S. junghuhnii Koord. 1908N77
    |--S. laeteviridis Stapf 1894N77
    |    |--S. l. var. laeteviridis [incl S. forbesii Brand 1901]N77
    |    |--S. l. var. alternifolia Noot. 1975N77
    |    |--S. l. var. basirotunda Noot. 1975N77
    |    |--S. l. var. kinabaluensis (Heine) Noot. 1975 [=S. kinabaluensis Heine 1953]N77
    |    |--S. l. var. mjobergii (Merr.) Noot. 1975 [=S. mjobergii Merr. 1928]N77
    |    |--S. l. var. pauciflora Noot. 1975N77
    |    `--S. l. var. velutinosa Noot. 1975N77
    |--S. lancifolia S. & Z. 1846 (see below for synonymy)N77
    |--S. lucida (Thunb.) S. & Z. 1835 (see below for synonymy)N77
    |--S. malieliensis Noot. 1975N77
    |--S. multibracteata Noot. 1975N77
    |--S. nivea Brand 1901N77
    |--S. obovatifolia Merr. 1917N77
    |--S. odoratissima (Bl.) Choisy ex Zoll. 1854 (see below for synonymy)N77
    |    |--S. o. var. odoratissima (see below for synonymy)N77
    |    `--S. o. var. wenzelii (Merr.) Noot. 1975 [=S. wenzelii Merr. 1915; incl. S. trichophlebia Merr. 1929]N77
    |--S. ophirensis Clarke 1882 [=Eugeniodes ophirense O. K. 1891]N77
    |    |--S. o. ssp. ophirensisN77
    |    |    |--S. o. ssp. o. var. ophirensisN77
    |    |    `--S. o. ssp. o. var. densireticulata Noot. 1975N77
    |    |--S. o. ssp. cumingiana (Brand) Noot. 1975 [=S. cumingiana Brand 1901]N77
    |    |    |--S. o. ssp. c. var. cumingiana (see below for synonymy)N77
    |    |    `--S. o. ssp. c. var. pachyphylla (Merr.) Noot. 1975 [=S. pachyphylla Merr. 1915]N77
    |    `--S. o. ssp. perakensis (King & Gamble) Noot. 1975 [=S. perakensis King & Gamble 1906]N77
    |         |--S. o. ssp. p. var. perakensis [incl. S. fragrans Elmer 1908]N77
    |         |--S. o. ssp. p. var. lingaensis Noot. 1975N77
    |         `--S. o. ssp. p. var. sumatrana Noot. 1975 non S. sumatrana Brand 1901N77
    |--S. paucistaminea F.v.M. & Bailey 1890N77
    |--S. polyandra (Blanco) Brand 1901 (see below for synonymy)N77
    |--S. pulchraN77
    |--S. pulvinata Noot. 1975N77
    |--S. pyriflora Ridl. 1915 [incl. S. bakeri Symington 1936]N77
    |--S. robinsonii Ridl. 1917N77
    |    |--S. r. var. robinsoniiN77
    |    |--S. r. var. angustifolia Noot. 1975N77
    |    |--S. r. var. latifolia Noot. 1975 non S. latifolia Krug & Urb. 1892N77
    |    `--S. r. var. pilosa Noot. 1975N77
    |--S. rubiginosa Wall. ex DC. 1844 [=Lodhra rubiginosa Miers 1879]N77
    |--S. salicoides Noot. 1975N77
    |--S. sumatrana Brand 1901N77
    |--S. sumuntia Buch.-Ham. ex Don 1825N77
    |--S. trichomarginalis Noot. 1975N77
    |--S. tricoccata Noot. 1975N77
    |--S. trisepala Merr. 1917N77
    |--S. verticillifolia Noot. 1975N77
    |--S. vidalii Rolfe 1912 [=S. cagayanensis Brand 1912]N77
    |--S. whitfordii Brand 1908N77
    |--S. wikstroemifolia Hayata 1915 [incl. S. microtricha Hand.-Mazz. 1943]N77
    `--S. zizyphoides Stapf 1894 [incl. S. clementis Merr. 1917]N77

Symplocos subg. Hopea Clarke 1882 [=Hopea Linné 1767 (nom. rej.); incl. Baranda Llanos 1857, Carlea Pr. 1851, Dicalix Lour. 1790, Dicalyx, Eugeniodes O. K. 1891 (nom. illeg.), Sariava Reinw. 1825]N77

Symplocos adenophylla Wall. ex Don 1837 [=Eugeniodes adenophyllum O. K. 1891; incl. S. adenophylla var. atrata Brand 1906, S. bancana Miq. 1861, S. beccarii Brand 1901, S. brandii Elmer 1912, S. constricta Brand 1901, S. iteophylla var. elliptica Miq. 1861, S. fulvosa King & Gamble 1906, S. iteophylla Miq. 1861, S. adenophylla var. merrittii Brand 1908, S. adenophylla var. montana Ridl. 1923 nec S. montana Brongn. & Gris 1866 nec Vidal 1886, S. pahangensis Brand in Fedde 1916, S. palawanensis Brand 1908, S. pruniflora Ridl. 1909, S. iteophylla var. rostrata Miq. 1861, S. adenophylla var. virgata Wall. ex Brand 1901]N77

Symplocos barringtoniifolia Brand 1904 [incl. Doxomma rigidum Miers 1875, Barringtonia rigida Clarke 1879, Eugeniodes rigidum O. K. 1891, Symplocos rigida Clarke 1882 non Don 1837]N77

Symplocos celastrifolia Griff. ex Clarke 1882 [=Eugeniodes celastrifolius O. K. 1891; incl. S. candicans Brand 1901, S. hutchinsonii Brand 1909, S. nigricans Brand 1901, S. peninsularis Brand 1909]N77

Symplocos cochinchinensis (Lour.) Moore 1914 ssp. cochinchinensis var. cochinchinensis [incl. Dicalyx aluminosus Bl. 1826, S. aluminosa Brand 1901, S. odoratissima var. aluminosa K. & V. 1900, S. delectans Brand 1916, S. ferruginea var. delectans Kaneh. & Hatus. 1942, S. ferruginea Roxb. 1832, Eugeniodes ferrugineum O. K. 1891, Lodhra ferruginea Miers 1879, Symplocos lachnobotrys var. glabrior Miq. 1861, S. horsfieldiana Miq. 1861, Dicalyx javanicus Bl. 1826, Lodhra javanica Miers 1879, Symplocos javanica Kurz 1871, S. lachnobotrys Miq. 1861, Eugeniodes lachnobotryum O. K. 1891, S. mollis Wall. ex Don 1837, S. spicata var. platystachya Don 1837, S. polystachya Wall. ex DC. 1844, Lodhra polystachya Miers 1879, S. ferruginea var. polystachya Clarke 1882, S. verhuellii Jungh. & De Vr. 1845, Lodhra verhuellii Miers 1879]N77

Symplocos cochinchinensis ssp. cochinchinensis var. philippinensis (Brand) Noot. 1975 [=S. ferruginea var. philippinensis Brand 1908; incl. S. ahernii Brand 1908, S. ramosii Merr. 1917, S. syringoides Brand 1901, S. ferruginea var. syringoides Hall. f. 1923]N77

Symplocos cochinchinensis ssp. cochinchinensis var. sessifolia (Bl.) Noot. 1975 [=Dicalyx sessifolius Bl. 1826, Eugeniodes sessilifolius O. K. 1891, Symplocos cochinchinensis ssp. sessifolia Noot. ex Steen. 1972, S. sessilifolia Gürke 1890; incl. Eugeniodes diengense O. K. 1891, Dicalyx salaccensis Bl. 1826, E. salaccense O. K. 1891, Symplocos subsessilis Choisy ex Miq. 1859, S. spicata f. subsessilis K. & V. 1900]N77

Symplocos cochinchinensis ssp. laurina (Retz.) Noot. 1876 [=Myrtus laurinus Retz. 1786 non S. laurina Wall. ex Don 1837, Eugenia laurina Willd. 1799, Symplocos spicata var. laurina Clarke 1882; incl. Dicalyx acuminatus Bl. 1826, S. acuminata Miq. 1859, S. spicata f. acuminata K. & V. 1900, S. spicata var. acuminata Brad 1901, Decadia aluminosa Lour. 1790, Drupatris cochinchinensis Lour. 1790, Symplocos flavida Miq. ex Clarke 1882, S. spicata f. javanica K. & V. 1900 non S. javanica Kurz 1871, S. laurina Wall. ex Don 1837, S. spicata var. malasica Clarke 1882, S. polycarpa Wall. ex Don 1837, S. ribes Jungh. & De V. 1845, Eugeniodes ribes O. K. 1891, Lodhra ribes Miers 1879, Dicalyx spicatus Bl. 1826, S. spicata Roxb. 1832, S. spicata f. xanthophylla K. & V. 1900, Lodhra xanthophylla Miers 1879]N77

Symplocos cochinchinensis ssp. thwaitesii var. stawellii (F.v.M.) Noot. 1975 [=S. stawellii F.v.M. 1865; incl. S. spicata var. australis Bth. 1869]N77

Symplocos costata (Bl.) Choisy 1854 [=Dicalyx costatus Bl. 1826, Eugeniodes costatum O. K. 1891; incl. Symplocos arcuata Brand 1901, S. caryophylloides Zool. 1857, S. sericea Brand 1901]N77

Symplocos fasciculata Zoll. 1854 [=Eugeniodes fasciculatum O. K. 1891; incl. S. fasciculata var. blumeana K. & V. 1900, S. fasciculata var. minor Miq. 1861, S. phanerophlebia Merr. 1914, Dicalyx tinctorius Bl. 1826 non S. tinctoria L’Hérit. 1791]N77

Symplocos lancifolia S. & Z. 1846 [incl. S. depauperata var. angustissima Brand 1912, S. betula Brand 1908, S. depauperata Merr. 1905, S. inconspicua Brand 1909, S. luzoniensis Rolfe 1886, S. merrilliana Brand 1908, S. montana Vidal 1886 non Brongn. & Gris 1866, S. depauperata var. sordida Brand 1908, S. zamboangensis Brand 1916]N77

Symplocos lucida (Thunb.) S. & Z. 1835 [=Laurus lucida Thunb. 1784, Hopea lucida Thunb. 1800; incl. Dicalyx ciliatus Bl. 1826, Symplocos ciliata Miq. 1859 non Presl 1831, S. laeviramulosa Elmer 1914, S. loheri Brand 1912, S. ridleyi King & Gamble 1906, S. theaefolia Don 1825, S. theifolia]N77

Symplocos odoratissima (Bl.) Choisy ex Zoll. 1854 [=Dicalyx odoratissimus Bl. 1826, Eugeniodes odoratissima O. K. 1891, incl. S. odoratissima var. divaricata Brand 1901]N77

Symplocos odoratissima (Bl.) Choisy ex Zoll. 1854 var. odoratissima [incl. S. acuminatissima Merr. 1916, S. apoensis Elmer 1914, S. bulusanensis Elmer 1939 (nom. illeg.), S. ciliata Presl 1831, S. patens f. ciliata Brand 1908, S. patens var. ciliata Brand 1901, S. dagamensis Brand 1916, S. elmeri Brand 1904, S. patens f. elmeri Brand 1908, S. floridissima Brand 1908, Pygeum grandiflorum King 1897, S. megabotrys Merr. 1914, S. patens Presl 1831, S. pseudospicata Vidal 1886, S. pulgarensis Elmer 1913, S. pulverulenta King & Gamble 1906, S. repandula Miq. 1861, S. salix Brand 1916, S. floridissima var. serrata Brand 1909, S. verdifolia Elmer 1939 (nom. illeg.), S. villarii Vidal 1886, Pygeum viride Baker 1924]N77

Symplocos ophirensis ssp. cumingiana (Brand) Noot. 1975 var. cumingiana [incl. S. agusanensis Elmer 1914, S. angularis Elmer 1908, S. curtiflora Elmer 1908, S. elliptifolia Merr. 1917, S. ilocana Merr. 1928, S. minutiflora Elmer 1914, S. purpurascens Brand 1912]N77

Symplocos polyandra (Blanco) Brand 1901 [=Guettarda polyandra Blanco 1845; incl. Baranda angatensis Llanos 1857, Carlea oblongifolia Presl 1851, Symplocos oblongifolia Rolfe 1885, S. superba Brand 1901]N77

*Type species of generic name indicated

References

Fritsch, P. W., L. M. Kelly, Y. Wang, F. Almeda & R. Kriebel. 2008. Revised infrafamilial classification of Symplocaceae based on phylogenetic data from DNA sequences and morphology. Taxon 57 (3): 823–852.

Nooteboom, H. P. 1975. Revision of the Symplocaceae of the Old World, New Caledonia excepted. Universitaire pers Leiden.

[N77] Nooteboom, H. P. 1977. Symplocaceae. Flora Malesiana, Series I—Spermatophyta, Flowering Plants 8 (2): 205–274.

Nooteboom, H. P. 2004. Symplocaceae. In: Kubitzki, K. (ed.) The Families and Genera of Vascular Plants vol. 6. Flowering Plants. Dicotyledons. Celastrales, Oxalidales, Rosales, Cornales, Ericales pp. 443–449. Springer.

[UB06] Udayan, P. S. & I. Balachandran. 2006. Cleistanthus sankunnianus Sivar. & Indu Balach.—a rare and little known endemic plant rediscovered from wild populations in Kollam district of Kerala state, India. Bulletin of the Botanical Survey of India 48 (1-4): 217–218.

Wang, Y., P. W. Fritsch, S. Shi, F. Almeda, B. C. Cruz & L. M. Kelly. 2004. Phylogeny and infrageneric classification of Symplocos (Symplocaceae) inferred from DNA sequence data. American Journal of Botany 91 (11): 1901–1914.

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