Dipsacaceae [Draft]

川续断科  chuan xu duan ke

Hong Deyuan (洪德元), Ma Liming (马黎明); Fred Barrie

Herbs, mostly perennial, less frequently annual or biannual, very rarely subshrubs. Leaves opposite, or sometimes verticillate, simple, entire or toothed, pinnatifid to pinnatisect. Flowers bisexual, epigynous, in cymes forming a terminal, involucral head, or a panicle (Triplostegia) nearly always subtended by a cupular, apically toothed or subentire involucel (epicalyx); receptacle hairy or naked. Calyx mostly small, cupular, or 4- or 5-segmented, or with up to 10 teeth or setae. Corolla sympetalous, 5(or 4)-lobed, ± irregular. Stamens 4, rarely 2 or 3, inserted at summit of corolla tube; anthers 4-sporangiate, opening by longitudinal slits. Gynoecium of 2 carpels, but one carpel obsolete; ovary 1-loculed; ovule solitary, apical, pendulous; style slender; stigma entire or 2-fid. Fruit an achene, enclosed by involucel and crowned by persistent calyx. Seeds with a large, straight embryo and scanty endosperm.

Ten genera with about 250 species: Eurasia and Africa; four genera and about 17 species in China.

1a.     Flowers in a panicle .......................................................................................  1. Triplostegia

1b.     Flowers in a compact head.

2a.    Plants spiny; flowers uniform; involucels usually without an evident crown ........  4. Dipsacus

2b.    Plants without spines; corolla of marginal flowers larger and 2-lipped; involucels usually with a crown.

3a.     Leaves usually all basal; calyx segments 8 or more, plumose or setose, shedding  2. Pterocephalus

3b.     Leaves mostly cauline; calyx segments 5, setose, persistent .........................  3. Scabiosa

 

1. Triplostegia Wallich ex de Candolle, Prodr. 4: 642. 1830.

双参属  shuang shen shu

Herbs, perennial, erect. Tap root fusiform. Rhizomes horizontal. Basal leaves dense, rosulate; leaf blade serrate or pinnatifid; cauline leaves decussate, similar to basal ones. Flowers in dichasia, forming a large, terminal and lax panicle; all parts of inflorescence densely white villous and glandular hairy; bracts linear; involucels in two layers, 4-lobed. Flowers small, epygynous, bisexual, 5-merous, nearly actinomorphic. Calyx small, urceolate, 8-ribbed; limb extremely short, disclike, 4- or 5-dentate. Corolla tubular-infundibular, 4- or 5-lobed; lobes nearly equal in size, imbricate in bud. Stamens 4, equal in length, alternate with corolla lobes; filaments inserted at upper part of corolla; anthers introrse, versatile, 4-sporangiate, opening by longitudinal slits. Ovary inferior; style solitary; stigma capitate. Achenes enveloped in urceolate involucels. Seeds nearly globose, smooth, with 2 obscure angles, acute at both ends; endosperm copious; embryo small.

Two species: from India and Nepal northward to Gansu, China, southward to Malaysia, and eastward to Taiwan, China; both in China.

Triplostegia pinifolia H. Lιveillι and T. mairei H. Lιveillι have been identified respectively as Sedum fastigiatum J. D. Hooker & Thomson and Chrysosplenium macrophyllum Oliver by L. A. Lauener (Notes Roy. Bot. Gard. Edinburgh, 30: 284. 290. 1970).

1a.     Corolla 3–5 mm; involucels hooked at apex; leaf blade ovate-lanceolate, petiolate  1. T. glandulifera

1b.     Corolla 7–12 mm; involucels not hooked at apex; leaf blade obovate to obovate-lanceolate, sessile    2. T. grandiflora

 

1. Triplostegia glandulifera Wallich ex de Candolle, Prodr. 4: 642. 1830.

双参  shuang shen

Hoeckia aschersoniana Engler & Graebner; Triplostegia repens Hemsley.

Herbs, perennial, erect, 15–40 cm tall. Tap root 2-forked, brown, nearly fusiform, 30–50 Χ 2–3 mm, slightly fleshy. Rhizomes elongate, 4-angular, with 4–6 nodes, internodes 0.5–2 cm, nodes rooted. Stems angular, glabrous or sparsely puberulous, above glandular hairy. Leaves opposite, petiolate; petiole 1–3 cm; leaf blade obovate-lanceolate, 2–6 Χ 1.5–4 cm, abaxially puberulous on veins, adaxially sparsely puberulous, late glabrescent, pinnatifid, more deeply so at base, margin irregularly lobed or serrate; upper leaves smaller, lobed, sessile. Inflorescence a large, lax and terminal panicle of dichasia; bracts 2–4 mm, with midrib, margin sparsely ciliate; pedicels ca. 1 mm at fruiting; involucels 4-lobed; lobes lanceolate, 1.5–2 mm, abaxially densely covered with purple glandular hairs. Calyx tube urceolate, ca. 1.5 mm, 8-ribbed, apex constricted into limb; limb 8-toothed or 8-dentate. Corolla white or rose, funnel-shaped, 3–5 mm, 5-lobed; lobes nearly actinomorphic, obtuse at apex. Stamens 4, slightly exserted; filaments inserted at near throat, ca. 5 mm; anthers white. Style slightly longer than stamens; ovary enveloped by saclike, narrowly oblong involucel. Achenes enveloped by involucels; involucels 3–4 mm in fruit, glandular hairy, 4-lobed; lobes long acuminate, hooked. Fl. and fr. Jul–Oct. 2n=18.

Forests, grassy slopes, meadows, by streams; 1500–4000 m. Chongqing, SE Gansu (Xigu), Hubei (Enshi, Jianshi), Shaanxi (Yangxian), W Sichuan, Taiwan (Yushan), SE Xizang, NW Yunnan [Bhutan, India, Malaysia, Myanmar, Nepal].

2. Triplostegia grandiflora Gagnepain, Bull. Soc. Bot. France 47: 333. 1901 [‘1900’].

大花双参  da hua shuang shen

Triplostegia delavayi Franchet ex Diels.

Herbs perennial, erect, 20–45 cm tall. Tap root often dichotomous, nearly fusiform, 30–40 Χ ca. 5 mm, slightly fleshy. Rhizomes 2–4-noded, internodes ca. 1 cm. Stems solitary, simple, slightly 4-angular, white villous, sometimes glandular hairy above. Leaves opposite, sessile, slightly amplexicaul, obovate to obovate-lanceolate, 3–8 cm, both surfaces villous, base cuneate, apex rounded, pinnatifid or pinnatilobate, terminal lobe largest, all lobes serrate; upper leaves gradually smaller and becoming bract-like upward on stems. Inflorescences a large, lax, terminal panicle of dichasia, inflorescence branches elongate, densely white villous and glandular hairy; bracts leafy to linear, 5–20 mm, 1-veined, white villous and glandular hairy, margin shallowly serrate or entire, apex obtuse; pedicels 2–3 mm; involucels calyxlike, 4-lobed; lobes lanceolate, 2–3 mm, densely covered with black glandular hairs, apex acute. Calyx tube ovoid, 8-ribbed; limb 5-dentate, teeth acute at base, hirsute. Corolla whitish rose, funnel-shaped, nearly actinomorphic, 1–1.2 cm, outside sparsely puberulous, 5-lobed; lobes ca. 1/3 as long as total length of corolla, apex obtuse. Stamens 4, inserted at upper part of corolla tube, slightly exserted; anthers yellow, introrse. Ovary enveloped by urceolate involucel; style shorter than stamens; stigma capitate. Achenes enveloped in involucels. Fl. and fr. Jul–Oct.

● Forests, forest margins, grassy slopes; 2000–3000 m. SW Sichuan, N Yunnan.

2. PTEROCEPHALUS Vaillant ex Adanson, Fam. 2: 152. 1763.

翼首花属  yi shou hua shu

Scabiosa sect. Pterocephalus (Vaillant ex Adanson) Bentham & J. D. Hooker.

Herbs, perennial or annual, or sometimes subshrubs. Leaves basal, rosulate, entire, pinnatifid to pinnatisect. Capitula solitary, terminal; involucre of two, rarely three whorls of 4–6 involucral bracts; receptacles covered with hirsute hairs or bracts; involucels 4–8-ribbed, apex obscurely dentate, or corona-like, or membranous and dentate. Calyx segmented and 8–24-setose or pinnate. Corolla 4- or 5-lobed, that of marginal flowers nearly 2-lipped, upper lip with one lobe, entire or 2-lobed; lower lip usually 3-lobed. Stamens 4, rarely 2 or 3, usually inserted at upper part of corolla. Ovary inferior, enveloped in involucels. Achenes smooth or ribbed.

About 25 species: from the Mediterranean to Central Asia and tropical Africa; two species in southwest China.

1a.     Leaves spatulate or linear-spatulate, entire or narrowly segmented; inflorescences globose in fruit; involucral bracts of outer layer narrowly ovate; persistent calyx segmented into 20 pinnate hairs; corolla 5-lobed, obovoid, ca. 4–5 mm ..........................................................................................................  1. P. hookeri

1b.     Leaves oblong-lanceolate or oblanceolate, pinnatifid to bipinnatisect; inflorescences compressed-globose in fruit; involucral bracts of outer layer lanceolate; persistent calyx segmented into 8 brown setae; corolla 4-lobed …………………………………………………… 2. P. bretschneideri

 

1. Pterocephalus hookeri (C. B. Clarke) Hφck in Engler & Prantl, Nat. Pflanzenfam. 4(4): 189. 1897.

匙叶翼首花  chi ye yi shou hua

Scabiosa hookeri C. B. Clarke in J. D. Hooker, Fl. Brit. India 3: 218. 1881; Pterocephalus batangensis Pax ex K. Hoffmann.

Herbs, perennial, 10–50 cm tall, white villous throughout. Tap root robust, columnar, 8–15 Χ 1.5–3.5 cm, lignified. Leaves all basal, rosulate, oblanceolate, 5–18 Χ 1–2.5 cm, apex obtuse or acute, base attenuate into winged petiole, entire, sparsely large serrate, or pinnatifid, with 3–5 pairs of segments, segments obliquely ovate or lanceolate, 1–2 cm, terminal one larger, lanceolate, abaxially white, midvein conspicuous, adaxially green, white puberulous. Scapes 10–40 cm, 2–4 mm in diam., white villous, more densely so near inflorescences. Capitula solitary, terminal, erect or slightly pendulous, globose, 3–4 cm in diam.; involucral bracts 2- or 3-layered, narrowly ovate to ovate-lanceolate, 12–18 Χ 5–7 mm, hairy, margin villous, apex acute; bracts linear-oblanceolate, 1–1.2 cm, densely white villous, midvein conspicuous, base becoming a slender claw; involucels tubular, 4–5 Χ ca. 1.5 mm, outside white hirsute, attenuate toward base, sinuate-dentate, apex slightly spreading. Calyx segmented to base into 20 soft pinnate hairs. Corolla yellowish white or pale purple, tubular-infundibular, 1–1.2 cm, outside rather densely villous, shallowly 5-lobed; lobes obtuse, ca. 3.5 mm. Stamens 4, exserted; anthers dark purple, ca. 3 mm. Ovary enveloped in involucel; style ca. 1.5 cm, obviously exserted; stigma pale brown, compressed-globose. Achenes pale brown, obovoid, 3–5 mm, longitudinally 8-ribbed, sparsely adpressed-hairy; pinnate hairs of persistent calyx ca. 1 cm, white. Fl. and fr. Jul–Oct.

Grassy slopes, meadows, by fields; 1800–4800 m. S Qinghai, W Sichuan (W of Heishui and Kangding), Xizang (west to Burang and north to Riwoqκ), N Yunnan [India, Nepal].

2. Pterocephalus bretschneideri (Batalin) Pritzel ex Diels, Bot. Jahrb. Syst. 29: 601. 1900.

裂叶翼首花  lie ye yi shou hua

Scabiosa bretschneideri Batalin, Acta Hort. Petrop. 14: 184. 1895.

Herbs, perennial, 8–40 cm tall. Tap root columnar, 3–20 mm in diam. Leaves basal, rosulate; petiole 3–10 cm; leaf blade oblong to oblanceolate, 5–20 cm, both surfaces glabrous or puberulous, pinnatifid to bipinnatisect, segments linear. Scapes white puberulous, more densely so near inflorescence. Capitula terminal, solitary, oblate, 2.5–3 cm in diam.; involucral bracts 2-layered, 10–14, broadly linear, outside white puberulous; receptacles disclike, densely white hairy; bracts brown, linear-lanceolate, small, 4–6 mm, pubescent, apex acuminate; involucels ellipsoid-obovoid, 4–5 mm, 8-ribbed, densely white hirsute, apex membranous, dentate. Calyx segmented into 8 brown setae, setae 1–1.2 cm, hispidulous above. Corolla pale rose to purple-red, tubular, 12–15 mm; tube 7–10 mm, densely white villous; limb with 4 obtuse lobes, slightly 2-lipped; upper lip slightly larger, ca. 2.5 mm; 3 lobes of lower lip ca. 2 mm, with several brown veins. Stamens 4, inserted at upper part of corolla; filaments ca. 5 mm, obviously exserted. Stigma capitate. Achenes ellipsoid, ca. 4 mm, apex tapering into a rostrum, 8-veined, sparsely villous; persistent calyx 8-setose. Fl. Jul–Aug, fr. Sep–Oct.

* Mountainous rocky crevices, grassy slopes in forests; 1600–3400 m. W Sichuan, SE Xizang (Nangxian), N Yunnan.

3. SCABIOSA Linnaeus, Sp. Pl. 1: 98. 1753.

蓝盆花属  lan pen hua shu

Herbs, perennial, sometimes semishrubs due to lignification at base, or biannual, rarely annual. Leaves opposite, cauline leaves amplexicaul; leaf blade pinnatipartite or pinnatisect, rarely entire. Capitula terminal, oblate-globose, ovoid, or ovoid-conical, long pedunculate; involucral bracts herbaceous, in 1 or 2 series; receptacles arched to semi-globose, sometimes columnar, with bracts; bracts linear-lanceolate, 1-veined, back often carinate; involucels (epicalyx) broadly infundibular or angular-columnar, 8-ribbed in fruit, furrowed in entire length, or furrowed only above and rounded at base, above often fissured into 2–8 pits, top becoming a membranous crown, crown campanulate or rotate, 16–30-veined, margin dentate. Calyx stalked, disc-like, 5-segmented into radial setae. Corolla tubular, blue, purple, purple-red, yellow, or white, 4- or 5-lobed, that of marginal ray flowers usually larger, 2-lipped; upper lip usually 2-lobed, shorter; lower lip 3-lobed; that of central flowers usually tubular, lobes nearly equal in length. Stamens 4. Ovary inferior, enveloped in involucels; style slender; stigma peltate or capitate. Achenes enveloped in involucels, crowned with persistent setose calyx.

About 100 species: Europe, Asia, S and W Africa, mainly in the Mediterranean; six species in China.

1a.     Annuals; capitula ca. 5 mm broad at anthesis; calyx setae to 1 cm ........................  3. S. olivieri

1b.     Perennials; capitula over 30 mm broad at anthesis; calyx setae shorter.

2a.    Involucels nearly cylindrical, not angular below, furrowed, or pitted above.

3a.     Leaf blades of basal leaves lanceolate or elliptic, entire .............................  1. S. alpestris

3b.     Leaf blades of basal leaves pinnatifid; leaf segments 1.5–3 mm wide  2. S. austro-altaica

2b.    Involucels 8-ribbed, or with 4 obvious and 4 obscure ribs.

4a.     Corolla yellow; involucels obviously 8-ribbed .......................................  4. S. ochroleuca

4b.     Corolla blue or purple; involucels 8-ribbed, but only 4 ribs marked, while others obscure.

5a.     Leaves pinnatifid to pinnatisect; segments linear or lanceolate ................  5. S. comosa

5b.     Leaves linear-lanceolate, margins serrate or lacerate ........................  6. S. lacerifolia

 

1. Scabiosa alpestris Karelin & Kirilow, Bull. Soc. Imp. Naturalistes Moscou 15: 536. 1842.

高山蓝盆花  gao shan lan pen hua

Herbs, perennial. Caudex lignified, 2–5-branched, outside brown-black. Stems 20–50 cm tall, with 2–4 nodes, usually pubescent, or glabrous. Basal leaves and first pair of lower cauline leaves petiolate; petiole 5–7 cm; leaf blade lanceolate, 5–8 Χ 0.5–1.1 cm, glabrous or sparsely puberulous, base attenuate into petiole, margin usually entire, margin with white verrucose hairs, apex acuminate. Cauline leaves 1–3-paired, second and third pairs of leaves pinnatisect; lateral segments linear-lanceolate; terminal one larger, lanceolate, 4–7 Χ 0.6–0.8 cm; all segments glabrous, but with white verrucose hairs on margins and beneath. Peduncles ca. 10 cm; capitula terminal, 3–4 cm broad at anthesis, to 5–6 cm including edges of flowers, globose and 3 cm broad in fruit; involucral bracts linear-lanceolate, 1.2–1.5 cm, densely white hirsute; involucels 8–10 mm, rounded and sparsely white-pilose below, with 8 pits above, membranous crown 5–6 mm, 16–18-veined, edges sinuous-dentate. Calyx 5-setose, setae brown, radial, 1–3 mm, longer than crown. Corolla pale purple or purple-red, outside lanate; limb ± lipped, 5-lobed; lobes unequal in size. Stamens 4; anthers yellow. Style purple-red; stigma capitate, exserted. Fl. May–Aug, fr. Aug–Sep.

Alpine meadows; 3000–3200 m. Xinjiang (Huocheng, Wuqia, Zhaosu) [Kazakhstan, Kirghizia].

2. Scabiosa austro-altaica Bobrov in Komarov, Fl. URSS 24: 457. 1957.

阿尔泰蓝盆花  a er tai lan pen hua

Scabiosa xinjiangensis Y. K. Yang, G. J. Liu & J. K. Wu.

Subshrubs, 20–60 cm tall. Caudex robust, lignified, several branched. Stem erect, with 1–4 nodes, densely puberulous. Leaves mostly basal; basal leaves petiolate; petiole 1–2.5 cm; leaf blade oblong, 3–10 Χ 0.7–3 cm, pinnatifid, segments linear-lanceolate, incised. Cauline leaves smaller, 2- or 3-paired, sessile, sparsely puberulous, ± setose at axils of veins, pinnatisect; segments lanceolate, 5–20 Χ 1.5–3 mm. Capitula 2–3 cm broad at anthesis, globose and 1.5–2 cm broad in fruit; involucral bracts narrowly ovate, apex acuminate, 1/3–1/2 as long as marginal ray flowers, sparsely puberulous; bracts linear-lanceolate, upper 1/3 carinate, lower part linear, sparsely villous; involucels 3–4 mm, above with 8 pits, base densely white hirsute, crown 2–3 mm broad. Calyx 5-setose. Corolla blue-purple, outside adpressed-puberulous. Fl. Jun–Jul, fr. Aug.

Grassy slopes; ca. 1200 m. Xinjiang (Altai, Burgin, Habahe) [E Kazakhstan].

3. Scabiosa olivieri T. Coulter, Mιm. Dipsac. 36. 1823.

小花蓝盆花  xiao hua lan pen hua

Lomelosia olivieri (T. Coulter) Greuter & Burdet; Scabiosa olivieri var. longinvolucra Y. K. Yang, N. R. Cui & Y. Hazit.

Herbs, annual, 10–38 cm tall. Tap root solitary, 40–50 Χ 3–4 mm. Stems terete, slender, branched from base and dichotomous from lower middle part, white puberulous. Leaves opposite, sessile; leaf blade oblong or linear-lanceolate, 20–50 Χ 3–10 mm, both surfaces densely villous, margin often entire, not segmented, or with 1 or 2 pairs of leaflets, sometimes pinnatifid. Peduncles 3–10 cm, slender, sparsely villous; capitula globose, 3–5 mm broad at anthesis, 5–15-flowered, marginal ray flowers larger than central ones; involucral bracts 3–5, oblong-ovate or ovate-orbicular, 2–3 Χ 1.5–2 mm, shorter than flowers, rather densely villous, apex acuminate; involucels broadly infundibular, ca. 2 mm, base white hispidulous, above half with 8 deep pits; crown spreading, 3–6 mm broad, membranous, with 20–24 ribs, sparsely villous along ribs outside. Calyx radially 5-setose, shortly stalked, brown, setae sometimes to 1 cm, 4–5 Χ longer than involucel crown. Corolla pale purple or white, outside puberulous. Achenes white hairy. Fl. May–Jun, fr. Jun–Jul. 2n=18

Sandy places, deserts of plains. N Xinjiang (Manas River basin, άrόmqi) [from E Mediterranean to C Asia and India].

4. Scabiosa ochroleuca Linnaeus, Sp. Pl. 1: 101. 1753.

黄盆花  huang pen hua

Herbs, perennial. Caudex columnar, outside brown, often multibranched. Stems caespitose or solitary, erect, 25–80 cm tall, shallowly furrowed, white villous at upper part. Basal leaves petiolate; petiole 1–5 cm; leaf blade pinnatifid with 2–4 pairs of segments, or undivided, 5–10 cm. Cauline leaves 2–5-paired, almost sessile or lower ones shortly petiolate; leaf blade 4–10 cm, abaxially densely pubescent, adaxially white villous, pinnatifid to pinnatisect or bipinnatisect, segments unequal in length, ultimate segments lanceolate or linear-lanceolate, 1.5–2 mm broad, 1-veined, apex acuminate. Peduncle 18–30 cm, white villous; capitulum solitary, terminal, oblate-spheroidal, 2–2.5 cm broad at anthesis, but becoming oblong and ca. 1.5 cm long in fruit; involucral bracts linear-lanceolate, both surfaces sparsely pubescent, apex acuminate; bracts oblanceolate, sparsely puberulous, apex acute; involucels infundibular, 3–4 mm, 8-ribbed and 8-furrowed, sparsely white villous on ribs and sparsely puberulous in furrows, crown 1.5–2 mm broad, sparsely puberulous outside, dentate on edges. Calyx lobes 5, brown-yellow, ca. 4 mm, setose. Corolla yellow or white, 7–10 mm, marginal ray flowers nearly equal to central ones in size. Corolla tube 6–7 mm, outside densely white villous; lobes 5, unequal in size, middle one of lower lip to 3 mm, others 1.5–2 Χ ca. 1.5 mm. Stamens 4, inserted at middle part of corolla tube, exserted. Achenes yellowish, ellipsoid, ca. 2.5 mm, setae to 7 mm; receptacles visible after achenes shed, fusiform, 10–15 Χ ca. 4 mm, densely puberulous. Fl. Jul–Aug, fr. Aug–Sep. 2n=16

Meadows, grasslands, grassy slopes: 1300–2200 m. NW Xinjiang [Kazakhstan, Mongolia, Russia (Siberia); C Europe].

5. Scabiosa comosa Fischer ex Roemer & Schultes, Syst. Veg. 3: 84. 1818.

蓝盆花  lan pen hua

Scabiosa austro-mongolica Hurusawa; S. comosa var. lachnophylla (Kitagawa) Kitagawa; S. fischeri de Candolle; S. fischeri f. breviseta Handel-Mazzetti; S. hairalensis Nakai; S. henanensis Y. K. Yang & J. K. Wu; S. hopeiensis Nakai; S. japonica Miquel var. acutiloba Hara; S. japonica subsp. tschiliensis Hurusawa; S. lachnophylla Kitagawa; S. liaoningensis Y. K. Yang & J. K. Wu; S. mansenensis Nakai; S. superba Grόning; S. superba f. elatior Grόning; S. superba f. nana Grόning; S. togashiana Hurusawa; S. tschiliensis Grόning; S. tschiliensis var. brevisecta Hurusawa; S. tschiliensis var. longiseta Hurusawa; S. tschiliensis var. superba (Grόning) S. Y. He.

Herbs, perennial, 30–80 cm tall. Roots 2- or 3-branched or simple, 12–25 Χ 0.3–0.5 cm. Stems erect, angular, internodes 6–12 cm, with adpressed white puberulous hairs more dense at base and below inflorescences. Basal leaves rosulate, petiolate; petiole 3–10 cm; leaf blade narrowly elliptic to oblong, 6–10 Χ 1–2 cm, both surfaces sparsely white pubescent, pinnatisect, rarely dentate-lobed; segments linear, 1–1.5 mm broad, often wilted at anthesis. Cauline leaves opposite, amplexicaul, forming a short sheath, sessile or with petiole 1–1.2 cm; leaf blade oblong, 8–15 Χ 4–5 cm, pinnatisect or bipinnatisect; segments linear, 1–4 mm wide, both surfaces glabrous or sparsely white puberulous, apex acuminate. Peduncles 10–25 cm, with dense, curved, white puberulous hairs; capitula 1 or 3, oblate-globose, 2.5–4(–7) cm in diam. at anthesis, globose and 1–1.5 cm in diam. in fruit; involucral bracts 6–14, lanceolate, 10–12(–28) Χ 1–3 mm, glabrous or sparsely pubescent; involucels yellowish, obconical, 2.5–5 mm (excluding crown), ca. 1.7 mm broad, 8-ribbed, but only 4 marked, ribs densely white hirsute, with 8 pits at apex, crown ca. 1.2 mm, 18–20-veined, densely white villous on veins, edge dentate. Calyx 5-divided; segments needle-like, 2.5–3 mm, sparsely hispidulous. Corolla pale blue or light purple, outside densely puberulous, those of central flowers tubular, 4–10 mm; limb 5-lobed, lobes equal in length; those of marginal ray flowers 2-lipped, to 2 cm, upper lip 2-lobed, lower lip 3-lobed, longer, middle lobe to 1 cm. Stamens 4, exserted. Style ca. 1 cm, exserted; stigma capitate. Achenes oblong, with 5 brown veins and persistent calyx setae. Fl. Jul–Aug, fr. Sep. 2n=16, 18.

Sandy dunes, dry mountain slopes, steppes; 300–1600(–3000) m. E Gansu, Hebei, W Henan, Heilongjiang, Jilin, Liaoning, Nei Mongol, S Ningxia (Guyuan), N Shaanxi, Shanxi [Korea, Mongolia, Russia (Far East, Siberia)].

6. Scabiosa lacerifolia Hayata, Bot. Mag. (Tokyo) 20:16. 1906.

台湾蓝盆花  tai wan lan pen hua

Herbs, perennial, glabrous or puberulous. Stems erect, 10–20 cm tall, sparsely branched. Leaves sessile, semi-amplexicaul; leaf blade linear-lanceolate, pinnatisect or lacerate, segments irregularly dentate; basal leaves 5–12 Χ 5–6 cm. Cauline leaves opposite, smaller, 4–5 cm, apex acute. Peduncles 5–6 cm or even longer; capitula terminal, oblate-globose, 3.5–4 cm broad; involucral bracts 3-whorled, lanceolate, outermost ones ca. 1.5 cm, inner ones shorter; bracts spatulate, ca. 5 mm; involucels 4-ribbed, with 8 pits at apex, crown membranous, 4-lobed, lobes obtuse. Calyx 5-setose, radial. Corolla blue-purple; marginal ray flowers usually larger, corolla ca. 1.8 cm, 5-lobed, 2-lipped. Stamens 4, all fertile. Style filiform. Achenes surrounded by involucels, and crowned with persistent involucels and calyx setae, obovoid, 3–3.5 mm; setae ca. 1 mm.

* Grassy slopes, rocky ridges; 2000–3600 m. Taiwan (Zhongyang Shan).

4. DIPSACUS Linnaeus, Sp. Pl. 1: 97. 1753

川续断属  chuan xu duan shu

Herbs, biannual or perennial. Stems erect, angulate and sulcate, usually echinulate or hispid along ridges. Basal leaves long petiolate, entire, 3-cleft or pinnatipartite, usually dentate or lobed. Cauline leaves opposite, sessile or petiolate, usually 3–5-lobed, sometimes pinnatipartite or entire, both surfaces usually hispid, but glabrous or papillate-echinulate in some species. Capitula terminal, oblong, globose, or ovoid-globose; involucral bracts erect or spreading, leafy, 1- or 2-layered; bracts rostrate; involucels 4–8-ribbed totally adnate to ovary; flowers bisexual. Calyx actinomorphic, shallowly disciform, 4-lobed, white pubescent. Corolla tubular, 4-lobed; lobes unequal in size. Stamens 4, inserted at corolla tube, alternate with corolla lobes. Gynoecium of 2 carpels; ovary inferior, enveloped by involucel; ovule solitary, pendulous, anatropous; style filiform; stigma oblique or lateral. Achenes surmounted with persistent calyx, enveloped by leathery involucel. Seed coat membranous; embryo imbedded in endosperm.

About 20 species: Eurasia and northern Africa; seven species in China, mainly in southwest provinces.

1a.     Stems glabrous; bract rostra 1.5–3 mm .............................................................  1. D. azureus

1b.     Stems ± spiny; bract rostra 3–11 mm.

2a.    Heads oblong-spherical, 2.5–7 Χ 2–4.9 cm; bract rostra 5–11 mm; lower thinly tubular part of corolla 3.5–5 mm.

3a.     Corolla black-purple; corolla thin-tube ca. 3.5 mm; bract rostra ca. 5 mm ....  4. D. atratus

3b.     Corolla white, or yellowish; corolla thin-tube 3.5–5 mm; bract rostra 5–11 mm.

4a.     Segments of cauline leaves 1–3-paired; heads 2–3.5 cm broad; leaves glabrous abaxially  2. D. inermis

4b.     Segments of cauline leaves more than 3-paired, heads usually 3.5–4.9 cm broad; leaves spiny abaxially .......................................................................................................  3. D. chinensis

2b.    Heads spherical, 1.5–3 Χ 1.5–3.2 cm; bract rostra 3–5 mm; lower thinly tubular part of corolla 2–2.5 mm.

5a.     Corolla black-purple .....................................................................  7. D. atropurpureus

5b.     Corolla white, yellow, or pink.

6a.     Corolla white or yellow; plants perennial, roots fleshy and fusiform ...........  5. D. asper

6b.     Corolla pink; plants biannual, roots ± lignified, not fleshy ....................  6. D. japonicus

 

1. Dipsacus azureus Schrenk in Fischer & C. A. Meyer, Enum. Pl. Nov. 1: 53. 1841.

天蓝续断  tian lan xu duan

Cephalaria beijiangensis Y. K. Yang, J. K. Wu & A. Sayit; C. dipsacoides Karelin & Kirilow; Dipsacus dipsacoides (Karelin & Kirilow) V. I. Bochantsev; D. xinjiangensis Y. K. Yang, J. K. Wu & T. Abdulla.

Herbs, perennial, 0.6–1.5 m tall. Stems glabrous, nearly spineless. Basal leaves long petiolate, undivided; cauline leaves opposite, ca. 6-paired, glabrous, from long petiolate with petiole to 10 cm to sessile; leaf blade 6–26 Χ 3–10 cm, pinnatifid to pinnatisect; terminal segment lanceolate, margin large serrate, lateral segments 4–7-paired, lanceolate to elliptic, 1.2–7 Χ 1–3 mm. Heads 3 or 4, subspherical, terminal one 2–2.5 Χ 3–4.5 cm; peduncles 30–60 cm, glabrous or pubescent above; involucral bracts 2-layered, 5 or 6 in outer layer, ovate, 7–9 in inner layer, elliptic or lanceolate; bracts obovate-lanceolate or oblanceolate, 5–14 Χ 4–5 mm, rostra 1.5–3 mm; involucels 4-ribbed, 3–4 mm. Calyx disciform, 0.5–1 mm, irregularly lobed. Corolla purple-blue, infundibular, 8–15 mm, outside villous, 4-lobed; lobes elliptic or oblong, 3–4 Χ 1.5–2.5 mm. Stamens 4, inserted at upper part of corolla tube; filaments 4–5 mm; anthers ca. 2 mm. Ovary included in involucel; style longer than stamens; stigma clavate. Achenes 4-angular, ca. 3.5 mm, totally enveloped in involucel. Fl. and fr. Aug–Sep.

1900–2000 m. Xinjiang (Huocheng, Tekes) [Kazakhstan, Kirghizia].

2. Dipsacus inermis Wallich in Roxburgh, Fl. Ind. 1: 367. 1820.

续断  zang xu duan

Cephalaria cachemiria Decaisne; Dipsacus inermis var. mitis (D. Don) Y. Nasir; D. mitis D. Don; D. strictus D. Don.

Herbs, perennial, 0.5–2 m tall. Tap root carrot-shaped, to 3.5 cm in diam. Stems branched, usually 6-ridged, ridges with sparse spines. Basal leaves rosulate, long petiolate; leaf blade elliptic, 5–13 Χ 3.5 cm, 3- or 5-segmented, or undivided but margin serrate, often wilted at anthesis, terminal segment much larger than lateral ones; cauline leaves petiolate to nearly sessile; petiole to 7 cm; leaf blade broadly lanceolate, 10–22 Χ 3.5–7 cm, 3-segmented to pinnatifid, segments 2-, rarely 3-paired, terminal segment largest, ovate, all segments yellowish hirsute on both surfaces, serrate on margins. Heads globose or oblong, solitary or 3 in number, 2–3.5 cm broad at anthesis and in fruit; peduncles to 40 cm; involucrate bracts linear, white hirsute; bracts lanceolate or obovate-lanceolate, 14–20 mm at anthesis, rostrum 5–9 mm, setose and hirsute on two lateral sides; involucels ovoid-columnar, 5–8 mm. Corolla white or yellowish, infundibular, 11–15 mm, lower thinly tubular part, 3.5–4 mm; limb 4-lobed, lobes unequal in size. Stamens 4, inserted at corolla tube, exserted. Style strongly exserted. Achenes narrowly ellipsoid, 4-angular, glabrous, ca. 4.5 Χ 1.5 mm. Fl. Jul–Aug, fr. Aug–Oct.

Forests, grassy slopes, by streams; 2100–3900 m. Xizang (Nyalam) [Afghanistan, Bhutan, India, Myanmar, Nepal, Pakistan].

3. Dipsacus chinensis Batalin, Trudy Imp. S.-Petersburgsk. Bot. Sada 13: 375. 1894.

大头续断 da tou xu duan

Dipsacus lijiangensis T. M. Ai & H. B. Chen.

Herbs, perennial, to 2 m tall. Tap root robust, to 2 cm in diam. Stems branched, 8-ridged, ridges sparsely spiny. Cauline leaves opposite, from long petiolate with petiole ca. 5 cm, to nearly sessile upward on stems; leaf blade broadly lanceolate, to 25 Χ 7 cm, lyrate, terminal segment much larger than lateral ones; lateral segments 3–8-paired, both adaxially and abaxially setose. Heads solitary or 3, globose or oblong, 3.5–4.9 cm broad; peduncles to 25 cm; involucrate bracts linear, setose; bracts lanceolate or obovate-lanceolate 14–15 mm, rostrum 5–11 mm, both spiny and pilose on two lateral sides; involucels ovoid-columnar, 5–8 mm. Corolla 9–14 mm, lower thinly tubular part 3.5–5 mm, 4-lobed, lobes unequal in size. Stamens 4, inserted at corolla tube, exserted. Style exserted. Achenes cuneate, 4-angular, densely pubescent, ca. 6 Χ 1.5 mm, apex exposed. Fl. Jul–Aug, fr. Sep–Oct.

* Forests, grassy slopes, by streams; 2100–3900 m. W Sichuan, SE Xizang, Yunnan (Dκqκn, Lijiang, Xianggelila).

4. Dipsacus atratus J. D. Hooker & Thomson ex C. B. Clarke in J. D. Hooker, Fl. Brit. India 3: 218. 1881.

紫花续断  zi hua xu duan

Virga atrata (J. D. Hooker & Thomson ex C. B. Clarke) Holub.

Herbs, perennial ca. 1 m tall. Stems branched, ridged, retrorse-spiny, to 1 cm in diam. Basal leaves and lower cauline leaves wilted at anthesis; upper cauline leaves sessile, 10–15 Χ 4–5 cm, both surfaces hispid, pinnatisect; terminal segment much larger than lateral ones, elliptic, apex acuminate, lateral segments 2–4-paired, all segments large serrate on margins. Heads globose, 3–3.5 cm at anthesis; peduncles to 40 cm; involucral bracts 5 or 6, ovate-elliptic, undivided, abaxially hirsute; bracts obovate, 9–11 mm, rostrum triangular, ca. 5 mm, base ca. 2 mm broad, spiny-hirsute on two lateral sides; involucels ca. 3 mm, glabrous, 4-ribbed and 4-dentate, with 4 minute teeth and 4 obscure ribs between large teeth and ribs. Calyx shallowly cup-shaped, 4-lobed, ca. 1 mm, edge spiny-hirsute. Corolla black-purple, infundibular, ca. 1.1 cm, base narrowly tubular, ca. 3.5 mm. Stamens exserted. Fl. Sep.

Forests on southern slopes; ca. 3600 m. S Xizang (Cona) [Bhutan, N India, Nepal].

5. Dipsacus asper Wallich ex C. B. Clarke in J. D. Hooker, Fl. Brit. India 3: 218. 1881.

川续断  chuan xu duan

Dipsacus asperoides C. Y. Cheng & T. M. Ai; D. asperoides var. emeiensis Z. T. Yin; D. cyanocapitatus C. Y. Cheng & T. M. Ai; D. daliensis T. M. Ai; D. daliensis var. multifidus H. B. Chen; D. enshiensis C. Y. Cheng & T. M. Ai; D. kangdingensis T. M. Ai & F. X. Feng; D. simaoensis Y. Y. Qian; D. yulongensis T. M. Ai & L. J. Yang.

Herbs, perennial, to 2 m tall. Tap root simple or branched, fusiform, fleshy, to 1.2 cm in diam. Stems erect, branched, 4–8-ridged, ridges sparsely covered with retrorse spines. Basal leaves rosulate; petiole to 25 cm; leaf blade pinnatisect, rarely entire, 15–25 Χ 5–20 cm, adaxially white setose, or papillate-setose, abaxially densely setose on veins, segments 5–9 in number, terminal one largest, to 15 Χ 9 cm, lateral ones usually obovate or spatulate; lower cauline leaves similarly long petiolate, leaf blade entire, but mostly pinnatifid, with 5–9 segments, terminal one lanceolate, to 11 Χ 5 cm, margin sparsely large serrate, apex acuminate; upper cauline leaves with shorter petioles, lanceolate, undivided or 3-segmented. Heads globose, 1.5–3.2 cm broad; peduncles to 50 cm; involucrate bracts 5–7, leafy, lanceolate or linear, hispid; bracts obovate, 7–11 mm, hispidulous, rostrate, rostrum 3–5 mm, setose on two lateral sides; involucels 4-ribbed, obovoid-columnar, 5–8 mm, white puberulous, apex 8-dentate. Calyx 4-ribbed, disc-shaped, ca. 1 mm, undivided or 4-lobed, hispidulous outside. Corolla yellowish or white, infundibular, 9–11 mm, base thinly tubular, ca. 2.5 mm; limb 4-lobed; lobes unequal, puberulous outside. Stamens 4, inserted at corolla tube, exserted; anthers purple. Achenes obovoid-columnar, ca. 4 mm. Fl. Jul–Sep, fr. Sep–Nov. 2n=18

Forest margins, thickets, among herbs, by streams or roadsides; 1500–3400 m. Chongqing, N Guangdong (Lianxian), N and W Guangxi (Longshen, Napo, Xing’an), Guizhou, W Hubei (Xingshan), Sichuan, S Xizang (Cona, Nyalam), Yunnan [Bhutan, India, Myanmar, Nepal].

6. Dipsacus atropurpureus C. Y. Cheng & Z. T. Yin, Acta Phytotax. Sin. 23: 302. 1. 1985.

深紫续断  shen zi xu duan

Dipsacus fulingensis C. Y. Cheng & T. M. Ai.

Herbs, perennial, 1–1.8 m tall. Tap root columnar, slightly fleshy. Stems erect, 6–8-ridged, ridges sparsely covered with retrorse spines. Basal leaves rosulate; petiole to 30 cm; leaf blade broadly lanceolate, 10–18 Χ 6–12 cm, pinnatifid or pinnatisect; segments 5–7; terminal one larger, oblong or broadly ovate, 6–12 Χ 4–8 cm; lateral ones smaller downward, lanceolate to ovate, all segments abaxially glabrous, adaxially puberulous or glabrous. Lower and middle cauline leaves with shorter petioles; leaf blade pinnatisect, terminal segment larger, lateral segments 2- or 3-paired, ovate to lanceolate; upper cauline leaves undivided or 3-segmented, segments lanceolate, both surfaces glabrous, sometimes adaxially sparsely white puberulous, margin entire. Heads globose, 2–2.5 cm broad; peduncles to 30 cm; involucral bracts 7 or 8, leafy, lanceolate, white puberulous; bracts oblong-obovate, 6–8 mm, rostrum 1–3 mm, both lateral sides glabrous or sparsely setose, only base white villous when glabrous; involucels obovoid-columnar; limb 4-lobed; lobes acute at apex. Calyx disc-shaped, 4-angular, outside glabrous, inside and apex villous. Corolla black-purple; tube 6–7 mm, base narrowly tubular, 1–1.5 mm; limb 4-lobed, one of lobes larger. Stamens inserted at corolla tube, obviously exserted. Achenes pale brown, 2.5–4 mm, 4-angular, cuneate. Fl. Jul–Sep, fr. Sep–Nov.

* Among herbs by streams, grassy slopes by fields. Chongqing (Fuling, Nanchuan, Qianjiang).

Dipsacus atropurpureus appears to differ from D. asper only in having a black-purple corolla (vs. yellowish or white) and a shorter rostrum (1–2 vs. 3–5 mm). We question whether they are two different species or conspecific. Thus, D. atropurpureus is now considered as a doubtful species. Field observations and population sampling are needed to clarify this issue.

7. Dipsacus japonicus Miquel, Verslagen Meded. Afd. Natuurk. Kon. Akad. Wetensch., ser. 2, 2: 83. 1868.

日本续断  ri ben xu duan

Dipascus lushanensis C. Y. Cheng & T. M. Ai; D. tianmuensis C. Y. Cheng & Z. T. Yin.

Herbs, biannual or (?)perennial, to 1.5 m tall. Roots not fleshy. Stems often branched, 4–6-ridged, with retrorse spines on ridges. Basal leaves rosulate, long petiolate; leaf blade narrowly elliptic, pinnatisect or undivided. Cauline leaves opposite, from long petiolate to nearly sessile; leaf blade elliptic-ovate to elliptic, 8–25 Χ 3–10 cm, abaxially spiny-hispid, adaxially hispidulous, pinnatisect or pinnatifid, rarely undivided, segments 2- or 3-paired, terminal one much larger than lateral ones. Capitula terminal, globose, 1.5–3.2 cm at anthesis and in fruit; involucral bracts linear, spiny-hispid; bracts obovate, 9–11 Χ 4–5 mm, hispidulous, rostrum 3–5 mm, lateral sides spiny-hispid; involucels 4-ribbed, 5–6 mm, white villous, 8-dentate at apex. Calyx cup-shaped, ca. 1 mm, 4-lobed, white villous. Corolla pink, purple-red, or purplish, 5–8 mm, outside villous. Stamens slightly exserted. Achenes oblong-cuneate, 4-angular, ca. 6 mm, glabrous. Fl. Aug–Sep, fr. Sep–Nov.

Grassy slopes, roadsides; below 2600 m. Anhui (Qingyang, Shucheng), Chongqing (Chengkou, Wushan, Wuxi), E Gansu (Pingliang, Yuzhong), Hebei, Henan, Hubei (Xianfeng), Hunan (Cili), Jiangsu, Jiangxi, Liaoning, Shaanxi, Shandong, Shanxi, N Sichuan (Barkam, Songpan), W Zhejiang [Japan, Korea].