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FOC | Family List | FOC Vol. 14 | Apiaceae

96. Heracleum Linnaeus, Sp. Pl. 1: 249. 1753.

独活属 du huo shu

Pu Fading (溥发鼎 Pu Fa-ting); Mark F. Watson

Sphondylium Miller; Tetrataenium (de Candolle) Mandenova.

Herbs, perennial, rarely biennial. Root fusiform or cylindrical, thickened, rarely fibrous. Stem erect, terete and often ribbed or striate, branching. Basal and lower leaves petiolate; petiole sheaths usually conspicuously broad; blade ternately or pinnately compound, hairy or glabrous. Umbels loose compound, terminal and lateral, terminal umbel with bisexual flowers, the lateral often with only staminate flowers; bracts few or absent, often caducous; rays numerous, spreading-ascending; bracteoles several, linear or lanceolate, entire; umbellules many-flowered. Calyx teeth triangular, lanceolate or obsolete. Petals white, rarely pinkish or pale yellow, obovate or obcordate, base cuneate, apex notched with a narrowly incurved lobule; outer flowers of the umbel often radiant with outer petals enlarged, broadly obovate, apex deeply 2-lobed. Stylopodium conic; styles short, erect or reflexed. Fruit obovoid, ovoid, broadly ovoid or suborbicular, strongly dorsally compressed, hairy or glabrous; dorsal and intermediate ribs filiform, sometimes raised, lateral ribs usually winged; vittae 1(–2) in each furrow, 2(–6) or absent on commissure, narrow, reaching to base of mericarp or clavate and much shorter than mericarp. Seed face plane, rarely slightly concave. Carpophore parted to base, usually persistent.

About 70 species: mainly in Asia and Europe, one species in North America, a few species in E Africa; 29 species (21 endemic) in China.

This is a widespread, taxonomically complex genus with the Hengduan Mountains forming one of two centers of diversity. Generic delimitation is problematic, both within Heracleum (recent molecular studies have shown it not to be monophyletic) and with several other genera with dorsally compressed fruit (e.g., Angelica, Peucedanum, and Semenovia). The distinctly clavate vittae, shorter than the length of the mericarp, are characteristic of many Heracleum species, but this does not hold for some of the Chinese species. Problems are compounded by the general paucity of good herbarium specimens and the inadequacy of historic type material. Many species are robust and tall plants, in which the primary inflorescences and basal leaves are large and therefore difficult to accommodate in a specimen press. Unfortunately, collectors have tended to select the smaller, lateral branches and upper leaves, which are less informative. Good fruiting material is also often lacking, and some taxa are recorded only from a very few collections. Revised classifications have been proposed for parts of the range of the genus, but as present knowledge is incomplete for Chinese taxa a traditional treatment is followed here.

Many species have reputed medicinal uses, and some are important elements in traditional Chinese medicine.

The following taxa have been described or reported from Chinese material, but are imperfectly known by the present authors because no specimens have been seen or the specimens are inadequate.

Heracleum canescens Lindley (in Royle, Ill. Bot. Himal. Mts. 232. 1839), described from NW India (“Mussooree,” J. F. Royle s.n., lectotype, K), has been doubtfully reported from Xizang and NW Yunnan. All Chinese specimens allegedly of this W Himalayan (NW India, Pakistan) species seen by us were inadequate for accurate determination.

Heracleum kansuense Diels (Repert. Spec. Nov. Regni Veg. 2: 66. 1906), described from Ningxia (“Gansu: Hsi ning fu” [Xiningpu]), W. Filchner 21, holotype, P).

Heracleum likiangense H. Wolff (Repert. Spec. Nov. Regni Veg. 33: 78. 1933), described from Yunnan (Lijiang, 13500 ft, J. F. C. Rock 4957, holotype, unlocalized).

Heracleum moellendorffii var. sageniifolium K. T. Fu (Fl. Tsinling. 1(3): 464. 1981 [“sagenifolium”]), described from Gansu and Shaanxi (K. T. Fu 17238, holotype, WNU).

Heracleum schansianum Fedde ex H. Wolff (Repert. Spec. Nov. Regni Veg. 33: 78. 1933), described from Shanxi (K. A. H. Smith 7632, holotype, GB).

1 Commissural vittae obscure, apparently absent.   (2)
+ Commissural vittae evident, 2–6.   (4)
2 (1) Calyx teeth obsolete.   1 Heracleum xiaojinense
+ Calyx teeth present.   (3)
3 (2) Rays 14–17; leaves ternate.   2 Heracleum fargesii
+ Rays 6–13; leaves pinnate, pinnae 3–4 pairs.   3 Heracleum subtomentellum
4 (1) Dorsal and intermediate ribs closely spaced; lateral wings broader than body of mericarp; vittae filiform.   (5)
+ Dorsal and intermediate ribs widely spaced; lateral wings narrower than body of mericarp; vittae filiform or clavate.   (7)
5 (4) Lower leaves 2-pinnate; rays 35–45; vittae solitary in each furrow, 2 on commissure.   6 Heracleum nyalamense
+ Lower leaves 2–3-pinnate; rays 6–25; vittae 1–2 in each furrow, 2–6 on commissure.   (6)
6 (5) Vittae 1–2 in dorsal furrows, extending to 1/2 length of mericarp.   4 Heracleum kingdonii
+ Vittae solitary in dorsal furrows, extending almost to base of mericarp.   5 Heracleum bivittatum
7 (4) Vittae filiform, extending at least 3/4 the length of mericarp.   (8)
+ Vittae clavate, extending to 1/2(–3/4) length of mericarp.   (10)
8 (7) Plants slender, to 60 cm high; rays 6–10; fruit ca. 4 × 4 mm.   9 Heracleum yunnanense
+ Plants stout, 80–200 cm high; rays 10–40(or more); fruit 8–17 × 7–12 mm.   (9)
9 (8) Petals yellowish, outer petals slightly radiant, hardly enlarged; vittae solitary in each furrow, 2 on commissure (Xinjiang).   7 Heracleum olgae
+ Petals white or pinkish, outer petals conspicuously radiant, greatly enlarged; vittae 1–2 in each furrow, 2–4 on commissure (Yunnan).   8 Heracleum nepalense
10 (7) Basal leaves ternate or 1–3-ternate/pinnate.   (11)
+ Basal leaves 1–4-pinnate.   (16)
11 (10) Calyx teeth obsolete.   (12)
+ Calyx teeth conspicuous.   (13)
12 (11) Basal leaves ternate; fruit glabrous.   10 Heracleum tiliifolium
+ Basal leaves ternate-pinnate; fruit sparsely hispidulous.   11 Heracleum moellendorffii
13 (11) Terminal leaflets cordate at base; fruit suborbicular; vittae 1–2 in each furrow.   12 Heracleum wenchuanense
+ Terminal leaflets truncate at base; fruit obovate; vittae solitary in each furrow.   13 Heracleum vicinum
14 Calyx teeth prominent, lanceolate; ultimate segments of leaves ovate-lanceolate or obovate-lanceolate.   22 Heracleum henryi
+ Calyx teeth minute or obsolete; ultimate segments of leaves ovate, broadly ovate, or ovate-rhombic.   (15)
15 (14) Calyx teeth minute; fruit obovoid; vittae extending to 3/4 the mericarp.   23 Heracleum rapula
+ Calyx teeth obsolete; fruit suborbicular; vittae extending to 1/2 the mericarp.   24 Heracleum wolongense
16 (10) Basal leaves 1-pinnate.   (17)
+ Basal leaves 2–4-pinnate.   (22)
17 (16) Leaf blade densely grayish or silvery-white hairy, tomentose.   28 Heracleum candicans
+ Leaf blade glabrous, sparsely pubescent or hispid.   (18)
18 (17) Robust, thick-stemmed plants, umbels 20–30-rayed.   18 Heracleum dissectum
+ Slender, thin-stemmed plants, umbels 10–25-rayed.   (19)
19 (18) Leaflets shallowly lobed, lobes broadly ovate.   15 Heracleum hemsleyanum
+ Leaflets deeply lobed to pinnatifid, lobes narrowly ovate to lanceolate.   (20)
20 (19) Adaxial surface of leaf segments apparently bullate.   16 Heracleum scabridum
+ Adaxial surface of leaf segments not bullate.   (21)
21 (20) Rays 10–25; bracteoles linear, acuminate at apex, distinctly shorter than pedicels.   14 Heracleum forrestii
+ Rays 20–25; bracteoles lanceolate, caudate at apex, equal to or slightly longer than pedicels.   17 Heracleum oreocharis
22 (16) Basal leaves 3–4-pinnate, ultimate segments less than 10 mm, linear.   29 Heracleum millefolium
+ Basal leaves 2–3-pinnate, ultimate segments more than 10 mm, ovate or lanceolate.   (23)
23 (22) Basal leaves 2-pinnate; bracts absent.   (24)
+ Basal leaves 2–3-pinnate; bracts 1–3.   (26)
24 (23) Leaflets lacerate-pinnatifid, segments lanceolate, terminal leaflets without decurrent wings at base; calyx teeth minute; fruit suborbicular, 4–6 mm.   19 Heracleum dissectifolium
+ Leaflets serrate, segments ovate, terminal leaflets with decurrent wings at base; calyx teeth triangular; fruit ovate or obovate, 6–9 mm.   (25)
25 (24) Rays 12–22; fruit ovoid, 8–9 mm; vittae 2–4 on commissure.   20 Heracleum franchetii
+ Rays 30–35; fruit obovoid, 6–7 mm; vittae 2 on commissure.   21 Heracleum souliei
26 (23) Ultimate segments of leaf ovate, terminal leaflets cuneate or cordate at base.   25 Heracleum stenopteroides
+ Ultimate segments of leaf lanceolate; terminal leaflets decurrent along rachis at base.   (27)
27 (26) Calyx teeth lanceolate; fruit obovoid, sparsely pilose.   26 Heracleum stenopterum
+ Calyx teeth triangular; fruit ovoid, almost glabrous.   27 Heracleum yungningense

  • List of lower taxa


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