五福花科 wu fu hua ke
Hong Deyuan (洪德元), Qin-er Yang (杨亲二); David E. Boufford [non-Viburnum], Valéry Malécot [Viburnum]
Shrubs, less frequently perennial herbs, or small trees. Leaves opposite, simple or compound: ternate, biternate, ternate-pinnate, biternate-pinnate, or odd-pinnate. Inflorescences terminal panicles or umbellate, spicate, or capitate cymes. Flowers bisexual; calyx and corolla both connate, (3–)5-merous. Stamens 5 (Sambucus, Viburnum) or 5, 4, or 3 (Adoxa, Sinadoxa, Tetradoxa), alternate corolla lobes, inserted on corolla tube, filaments cleft into 2 semi-stamens (in Adoxa, Sinadoxa, Tetradoxa); anthers 1-celled, peltate, extrorse, longitudinally dehiscent; staminodes 5, 4, or 3, in inner whorl, opposite corolla lobes. Ovary semi-inferior to inferior, 1- or 3–5-loculed; styles 5, 4, or 3, connate or free, or absent; stigmas capitate or 2- or 3-fid. Fruit a drupe; seeds 1 or 3–5.
Four genera with about 220 species: mainly in the Northern Hemisphere; four genera and ca. 80 species in China (??#endemic).
1a. Leaves simple; ovary 1-loculed; drupes 1-seeded; shrubs or small trees ................ 1. Viburnum
1b. Leaves compound; ovary 3–5-loculed; drupes usually 3–5-seeded; shrubs, perennial herbs, or small trees.
2a. Shrubs or small trees, rarely coarse perennial herbs more than 0.5 m tall; stamens 5, undivided 2. Sambucus
2b. Perennial herbs less than 30 cm tall; fertile stamens 5, 4, or 3, cleft into 2 semi-stamens, anthers 2-celled.
3a. Basal and cauline leaves pinnately compound; inflorescence of several condensed cymes in an interrupted spike ........................................................................................................ 4. Sinadoxa
3b. Basal leaves ternately or biternately compound; cauline leaves simple and 3-cleft or ternate; inflorescences condensed, capitate, several flowered cymes or racemes.
4a... Basal leaves simple, 3-cleft; cauline leaves simple, 3-cleft; flowers 3–5, 4-merous; pedicels 5–10 mm; filaments 2-fid to middle ...................................................... 3. Tetradoxa
4b... Basal leaves ternate or biternate; cauline leaves simple and 3-cleft or ternate; flowers 3–9, 4- or 5-merous, nearly sessile or shortly pedicellate; filaments bifid to base .. 5. Adoxa
接骨木属 jie gu mu shu
Qin-er Yang (杨亲二); David E. Boufford
Shrubs, small trees, or perennial herbs, gynodioecious or hermaphroditic, deciduous, whole plant sometimes with extrafloral nectariferous glands. Branches smooth, striate, or warty, with stout pith. Leaves with or without stipules, imparipinnate, or incompletely bipinnate, rarely laciniate; leaflets opposite or alternate, serrate or divided. Inflorescences terminal, flat or convex corymbs or panicles, pedunculate or sessile. Flowers actinomorphic or sometimes dimorphic, sometimes with glandular nectaries, articulate with pedicel; bracts mostly absent; bracteoles 1 or absent. Calyx tube: limb 3–5-parted; corolla rotate, white, lobes 3–5. Stamens 5, inserted at base of corolla; filaments erect, filiform; anthers 2-celled, oblong, cells free, attached at middle. Ovary locules 3–5, ovules 1 per locule; style cushionlike; stigmas 3 or 5. Fruit berry-like, 3–5-seeded; seeds triquetrous or ellipsoid; embryo ca. as long as seed.
About 10 species: temperate to subtropical regions and tropical mountains; four species in China (one endemic); plus one (Sambucus nigra Linnaeus) occasionally cultivated.
Bolli, R. 1994. Revision of the genus Sambucus. Dissertationes Botanicae Bd 223. 1–227 p., 29 p. of plates: ill.
In some species, the vegetative parts of the plant when bruised and the flowers have a fetid odor.
1a. Perennial herbs or shrubs; lenticels absent or inconspicuous; young branches striate; inflorescences flat topped umbellate cymes.
2a. All flowers hermaphroditic, rotate; lenticels inconspicuous or absent; lateral leaflets without glandular teeth; terminal leaflet narrowly cuneate, often decurrent and connected to next lower leaflet pair; pith of roots red or white; pyrenes rugose or smooth .................................................................... 1. S. adnata
2b. Some flowers sterile, urceolate; lenticels inconspicuous; lateral leaflets with 2–4 glandular teeth below middle and at base; terminal leaflet rounded to cuneate at base, not or only slightly decurrent; pith of roots white; pyrenes tuberculate .................................................................................... 2. S. javanica
1b. Shrubs or small trees; lenticels conspicuous; young branches round; inflorescences and infructescences pyramidal paniculate cymes.
3a. Petiolules, foliar axis, and abaxial surface of leaflets glabrous ....................... 3. S. williamsii
3b. Petiolules, foliar axis, and veins at base of abaxial surface of leaflets hairy ....... 4. S. sibirica
血满草 xue man cao
Sambucus schweriniana Rehder.
Herbs, suffrutescent, or low shrubs, 1–2 m tall, rhizomatous. Pith of roots and rhizomes white or red. Stem herbaceous, obviously striate; pith white or red; lenticels absent. Leaves imparipinnate; stipules blade-like or linear; leaflets 3–5 pairs, narrowly elliptic to narrowly ovate or lanceolate, 4–15 × 1.5–2.5 cm, abaxially sparsely pubescent, more densely pubescent on veins, base obtuse and oblique, margin serrate, apex acuminate; terminal pair of leaflets often connate at base along rachis, sometimes connected also with terminal leaflet, remaining leaflets alternate or sometimes nearly opposite; stipules of leaflets reduced to urceolate glands. Inflorescences terminal, pedunculate, with 3–5 rays, umbellate cymes, 12–15 cm broad and tall, with dense yellow pubescence when young and ± glandular hairy. Flowers all normal, not reduced to glands, 4-5 mm in diam.; calyx urceolate, pubescent; corolla white. Filaments dilated at base, anthers yellow. Ovary locules 3; styles short or nearly absent; stigma 3-lobed. Fruit orange or red, becoming black on drying, globose, 3-4 mm in diam.; pyrenes ovoid, rugose or smooth, ca. 2 × 1.5 mm. Fl. May–Jul(-Sep), fr. Sep–Oct. 2n = 36*.
Forests, thickets, streamsides, moist shaded mountain slopes, alpine grasslands; 1600–3600 m. Gansu, Guizhou, Hubei, Ningxia, Qinghai, Shaanxi, Sichuan, Xizang, Yunnan [Bhutan, India (Darjeeling, Sikkim)].
接骨草 jie gu cao
Ebulus chinensis (Lindley) Nakai; E. formosana (Nakai) Nakai; Sambucus argyi H. Léveillé; S. chinensis Lindley; S. chinensis var. formosana (Nakai) H. Hara; S. chinensis var. pinnatilobatus G. W. Hu; S. formosana Nakai; S. formosana var. arborescens Kanehira & Sasaki; S. henriana Samutina; S. hookeri Rehder; S. javanica var. argyi (H. Léveillé) Rehder; S. javanica subsp. chinensis (Lindley) Fukuoka.
Herbs, suffrutescent, or low shrubs, 1–2 m tall. Stem obviously striate; pith white; lenticels inconspicuous. Leaves imparipinnate; stipules leaflike or sometimes reduced to blue glands; lateral leaflets 2 or 3 pairs, alternate or opposite, narrowly ovate, 6–13 × 2–3 cm, adaxially sparsely pubescent when young, base obtuse and oblique, margin serrulate, often with 1–several glandular teeth from below middle nearly to base, apex long acuminate; terminal leaflet ovate or obovate, base cuneate, sometimes connected with next lower leaflets, basal pair of leaflets shortly petiolulate; stipules of leaflets absent. Inflorescences terminal, compound umbellate cymes, lax, pedunculate, with leaflike bracts at base of peduncle, with 3–5 slender rays, with sparse yellow pubescence. Some flowers modified into persistent urceolate nectaries. Calyx tube urceolate, lobes triangular; corolla white, connate at base. Anthers yellow or purple. Ovary locules 3, styles short or nearly absent; stigma 3-lobed. Fruit red, nearly globose, 3–4 mm in diam.; pyrenes 3 or 4, ovoid, ca. 2.5 mm, verrucate. Fl. Apr–May(–Aug), fr. Aug–Sep.
Mountain slopes, forests, streamsides, grasslands; 300–2600 m. Anhui, Fujian, Gansu, Guangdong, Guangxi, Guizhou, Hainan, Henan, Hubei, Hunan, Jiangsu, Jiangxi, Shaanxi, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [Cambodia, India, Indonesia, Japan, Laos, Malaysia, Myanmar, Philippines, Thailand, Vietnam].
Sambucus javanica often has abundant extrafloral nectaries on the petioles, branchlets, young stems, peduncle, and inflorescence branches.
接骨木 jie gu mu
Sambucus barbinerivis Nakai; S. buergeriana (Nakai) Blume ex Nakai var. buergeriana; S. buergeriana f. cordifoliata Skvortsov & W. Wang; S. foetidissima Nakai; S. junnanica J. J. Vassiljev; S. latipinna Nakai var. pendula Skvortsov; S. manshurica Kitagawa; S. racemosa Linnaeus subsp. manshurica (Kitagawa) Voroshilov; S. peninsularis Kitagawa; S. potaninii J. J. Vassiljev.
Shrubs or small trees, 5–6 m tall. Old branches reddish brown, with conspicuous, narrowly elliptic lenticels; pith brownish. Leaves imparipinnate; leaflets (1 or)2- or 3(–5)-jugate, lateral leaflets ovate-orbiculate or narrowly elliptic to oblong-oblanceolate, 5–15 × 1.2–7 cm, base cuneate or rounded, sometimes cordate, asymmetrical, margin irregularly serrate, sometimes with 1 to several glandular teeth at base or below middle, apex acute to acuminate or caudate; lowest pair of leaflets sessile or petioles to ca. 0.5 cm; terminal leaflet ovate or obovate, adaxially sparsely pubescent when young, glabrescent, petiolule ca. 2 cm, base cuneate, apex acuminate or caudate; stipules narrowly linear or reduced to bluish protrusions. Inflorescences terminal cymose panicles, 5–11 × 4–14 cm, pedunculate, sometimes sparsely pubescent, soon glabrescent. Flowers appearing simultaneously with leaves, dense; calyx tube urceolate, ca. 1 mm, lobes triangular-lanceolate, slightly shorter than tube; corolla pinkish in bud, white or yellowish when open; tube short; lobes oblong or narrowly ovate-orbiculate, ca. 2 mm. Stamens spreading, ca. as long as corolla lobes; filaments slightly dilated at base; anthers yellow. Ovary 3-loculed; styles short; stigmas 3-lobed. Fruit red, rarely bluish or purplish black, ovoid or nearly globose, 3–5 mm in diam.; pyrenes 2 or 3, ovoid to ellipsoid, 2.5–3.5 mm, slightly rugose. Fl. Apr–May, fr. Sep–Oct. 2n = 36*.
* Mountain slopes, scrub, streamsides, roadsides, beside houses; 500–1600 m. Anhui, Fujian, Gansu, Guangdong, Guangxi, Guizhou, Hebei, Heilongjiang, Henan, Hubei, Hunan, Jiangsu, Jilin, Liaoning, Shaanxi, Shandong, Shanxi, Sichuan, Yunnan, Zhejiang.
Sambucus williamsii is treated as a synonym of a highly variable, circumboreal S. racemosa Linnaeus by Bolli (1994). The black fruit and dark green leaves (Bolli, 1994) distinguish S. williamsii and S. melanocarpa A. Gray of western North America from other members of the S. racemosa complex. Sambucus williamsii also has a more pronounced fetid odor. Because of the high degree of variation and the pronouncement by Bolli that the variation within his circumscription of S. racemosa should be further evaluated in a geographical context through field, cytological, and biochemical methods, for the time being, we prefer to maintain S. williamsii, and, with less certainty, the following species, S. sibirica Nakai, as distinct.
西伯利亚接骨木 xi bo li ya jie gu mu
Sambucus buergeriana (Nakai) Blume ex Nakai var. miquelii (Nakai) Nakai; S. racemosa Linnaeus subsp. sibirica H. Hara; S. williamsii Hance var. miquelii (Nakai) Y. C. Tang ex J. Q. Hu.
Shrubs, 2–4 m tall, densely branched. Bark reddish brown, longitudinally sulcate, with elliptic lenticels; pith brownish; young branches yellowish white hirsute, sometimes papillose. Leaflets 2-jugate, abaxially pale, adaxially green, ovate-lanceolate to lanceolate, 5–14 × 1.6–5.5 cm, midvein long hirsute, base cordate and oblique, margin irregularly acutely toothed, apex long acuminate; petiolule and leaf rachis yellow hirsute; stipules glandular. Inflorescence an erect cymose panicle, 3.5–5 cm; peduncle papillose. Flowers opening with leaves; calyx tube urceolate, ca. 1 mm; lobes triangular-lanceolate, slightly shorter than tube; corolla greenish or yellowish; lobes oblong. Stamens yellowish brown; anthers yellow. Ovary 3-loculed; styles short; stigmas 3-lobed. Fruit red, ovoid or nearly globose, 3–5 mm in diam.; pyrenes 2 or 3, ovoid to ellipsoid, 2.5–3.5 mm, slightly rugose. Fr. Jul–Aug. 2n = 18, 36, 38*.
Rocky mountain slopes, rock crevices along rivers. Heilongjiang, Jilin, Liaoning, Xinjiang [Mongolia, Russia].
Sambucus sibiricus should perhaps be included within S. williamsii. The differences separating them, young branches, petiolule, and leaf rachis slightly pubescent versus glabrous in S. williamsii, are minor.
四福花 shi fu hua
Hong Deyuan (洪德元); David E. Boufford
Adoxa sect. Tetradoxa (C. Y. Wu) H. Hara; Adoxa subgen. Tetradoxa (C. Y. Wu) S. Y. Li & Z. H. Ning
Herbs, perennial. Rhizomes creeping. Stems 1 or 2. Basal leaves 1–3; cauline leaves 2, opposite; all leaves petiolate, compound. Inflorescences cymes, terminal, headlike clusters. Flowers yellowish green, sessile, 4- or 5-merous. Calyx shallowly cup shaped. Corolla rotate; tube extremely short; lobes papillate adaxially. Fertile stamens 4 or 5, inserted on corolla tube; filaments 2-fid to middle; anthers 1-celled, peltate, extrorse, longitudinally dehiscent; inner stamens reduced to glandular papillae. Carpels 4, nearly free, nearly superior; locules 4 or 5; styles 4 or 5, connate at base; stigmas capitate. Fruit a drupe. x=9, 2n=36, 54, 108.
* One species: endemic to Sichuan, China.
四福花 shi fu hua
Adoxa omeiensis H. Hara.
Herbs, 10–16 cm tall, slender, glabrous. Rhizomes white, filiform, with dense white fleshy scales; stolons filiform, from axils of scales. Stems 4-angled. Basal leaves 1–3; petiole 3–12 cm; leaf blade ovate or deltoid-ovate, 1–3 × 1–3 cm, 3-cleft; segments narrowly ovate, 2–5-lobed, apex of lobes mucronate. Cauline leaves 2, opposite, similar to those of basal leaves; petiole 0.4–2 cm. Cymes condensed in a pseudo-raceme, flowers 3–5 together; pedicels 0.5–1 cm; flowers yellowish green, 5–8 mm in diam. Calyx discoid, 4-lobed; lobes persistent, narrowly triangular-lanceolate, 2–3 mm. Corolla nearly rotate, 4-cleft; lobes narrowly ovate, 2.5–4 mm, inside densely glandular papillate, with papillate glands in center at base, apex acuminate. Outer (fertile) stamens 4, 0.4–1 mm; filaments incurved, divergent and stamens T-shaped apically; anthers ca. 0.6 mm. Styles slightly recurved; stigmas punctiform. Fl. May–Jun, fr. Aug.
* Wet places in forests, sometimes on rocks; ca. 2300 m. Sichuan (Emei Shan, Ya’an).
华福花属 hua fu hua shu
Hong Deyuan (洪德元); David E. Boufford
Herbs, perennial. Rhizomes erect. Roots fibrous. Stems 2–4, caespitose, glabrous. Basal leaves ca. 10; cauline leaves 2, opposite, both basal and cauline leaves ternate or biternate. Cymes with 3–5 flowers in an interrupted spike; lowest cymes long pedunculate, in axils of cauline leaves; flowers small. Calyx cup shaped, fleshy, often 3-lobed; lobes saccate and closed, narrowly winged on ridge. Corolla rotate, 3- or 4-lobed; tube short. Fertile stamens equal to corolla lobes in number, inserted at throat of corolla, alternate with corolla lobes, 2-fid to base; anthers 1-celled, extrorse; staminodes equal to fertile ones in number, opposite to corolla lobes. Carpels 2; ovary ovoid-globose, semi-inferior, 1-loculed; ovules solitary, pendulous; styles absent; stigmas solitary, dotted.
Monospecific: endemic to Qinghai, China.
华福花 hua fu hua
Herbs, perennial, caespitose, glabrous throughout. Stems 1–4, erect or ascending, green, 10–25 cm tall, 3–5 mm in diam. Basal leaves ca. 10, ternate-pinnate or biternate-pinnate; petiole to 10 cm; terminal leaflet ovate or ovate-oblong, 3–10 cm, irregularly lobed or pinnatipartite to pinnatisect; segments 3-lobed to multi-lobed or cleft; lateral leaflets ovate, 1.5–2 cm, 3-lobed. Cauline leaves 2, opposite, ternate, 2.5–5 cm; petiole 1–2 cm; leaf blade ovate-deltoid; terminal leaflet ovate, 1.5–3.5 cm, irregularly lobed; lateral leaflets ovate-elliptic or ovate, 0.8–1.8 cm, 3-, or 5–10-lobed. Spikes to 8 cm; flowers yellowish green. Calyx cup-shaped, fleshy, (2 or)3(or 4)-lobed. Corolla yellow-brown, 2–3 mm in diam., 3- or 4-lobed; lobes oblong-ovate, glandular-punctate near top inside, base with a nectary. Fertile stamens 3 or 4; filaments filiform, ca. 1.5 mm; anthers yellow, spherical, ca. 0.5 mm in diam. Stigma solitary. Fl. Jun–Jul; fruit not seen.
* Rock shelters, moist ravines, alpine debris slopes; 3900–4800 m. Qinghai (Nangqên, Yushu).
五福花属 wu fu hua shu
Hong Deyuan (洪德元); David E. Boufford
Herbs, perennial. Rhizomes creeping, filiform, terminated by a tuber; tuber white, giving rise to following year’s plant. Stems 1 or 2. Basal leaves 1–3, ternate to biternate; cauline leaves 2, opposite, rarely alternate; all leaves petiolate, 3-cleft or compound. Inflorescences cymes, terminal, headlike clusters. Flowers yellowish green, sessile, 4- or 5-merous. Calyx shallowly cup-shaped; lobes of terminal flower 2, lobes of lateral flowers 3. Corolla rotate; tube short; lobes 3–6 in terminal flowers, 4–6 in lateral flowers, lobes papillate adaxially. Fertile stamens 4 or 5, inserted on corolla tube; filaments 2-fid to base; anthers 1-celled, peltate, extrorse, longitudinally dehiscent; inner stamens reduced to glandular papillae. Ovary inferior to semi-inferior, locules 4 or 5; styles 4 in terminal flowers, 5 in lateral flowers, connate at base; stigmas capitate. Fruit fleshy, berry-like. x=9, 2n=36, 54, 108.
Two or three species: circumpolar; two species in China (one endemic).
1a. Peduncle in flower and fruit straight, erect to ascending, held above leaves; corolla lobes broadly ovate to orbicular 1. A. moschatellina
1b. Peduncle in flower and fruit recurved or coiled, nearly always held below leaves; corolla lobes lanceolate to ovate 2. A. xizangensis
五福花 wu fu hua
Adoxa inodora (Falconer ex C. B. Clarke) Nepomnjasczaja; A. moschatellina var. inodora Falconer ex C. B. Clarke.
Rhizomes to 2.5–25 cm long. Stems 1 or 2, 8–15 cm tall, glabrous. Basal leaves 1–3; petiole 4–9 cm; leaflets broadly ovate or orbicular, 1–2 cm, 3-cleft; petiolules 0.6–1.2 cm. Cauline leaves 2, opposite, 3-cleft or ternate, segments 3-lobed; petiole ca. 1 cm. Peduncle erect in flowers and fruit, inflorescence held above leaves. Inflorescences compact headlike cymes of 5–9 flowers. Flowers sessile or pedicels 0.5–1.5 mm, yellowish green, 4–7 mm in diam. Calyx lobes rectangular or broadly elliptic to ovate, 1.5–3 × ca. 1 mm, apex obtuse. Corolla lobes broadly ovate to orbicular or rectangular, 1.75–3.25 × 2–2.5 mm, apex rounded to obtuse. Ovary semi-inferior to inferior, 4- or 5-loculed. Fruit not seen. Fl. Apr–Jul, fr. Jul–Aug. 2n=36.
Forests, margins of forests, meadows; below 4000 m. Hebei, Heilongjiang, Liaoning, Nei Mongol, Qinghai, Shanxi, Sichuan, Xinjiang, Xizang, Yunnan [India, Japan, Korea, Nepal, Pakistan, Russia; NW Africa, Europe, North America].
??vernacular name
Rhizomes to 4–15 cm long. Stems 1 or 2, 5–10 cm tall, glabrous. Basal leaves 1–3, ternate to biternate; petiole of basal leaves 2–12 cm; leaflets elliptic, ovate, orbicular, or obovate, 1–2.5 × 1–3 cm, 3-cleft; petiolules 0.6–1.2 cm. Cauline leaves 2, opposite, ternate or rarely biternate, 3-cleft, segments 3-lobed, lobes apiculate; petiole 0.5-2 cm. Peduncle coiled in flower and fruit. Inflorescences often held at or below level of leaves, compact headlike cymes of 5–9 flowers. Flowers sessile or subsessile, yellowish green, 2.5–5 mm in diam. Calyx lobes lanceolate to narrowly rectangular, 1.2–2 × ca. 1 mm, apex obtuse. Corolla lobes lanceolate to ovate, ca. 2 × 0.9–1.2 mm. Fruit 5–8 mm in diam. Fl. Jun–Aug, fr. Jul–Sep.
* Stony or rocky limestone soils; forests, forest margins, meadows, shaded slopes; 3400–3900 m. Sichuan, Xizang, Yunnan.