Morinaceae [Draft]

刺参科 ci shen ke

Hong Deyuan (洪德元); Fred Barrie, Charles D. Bell, Peter F. Stevens

Herbs perennial, caudexes usually covered with remains of old leaves. Leaves opposite or verticillate, estipulate; petioles often connate  at base, forming a sheath. Flowers hermaphroditic, zygomorphic, in verticillasters or nearly in heads of dichasia. Involucel (epicalyx) of 4 fused bracteoles, with several to numerous spiny teeth or setae, 12-nerved. Calyx slit along adaxial side, or 2-lipped. Corolla tubular; limb 5-lobed, scarcely to markedly 2-lipped. Stamens 4, all fertile, or 2 fertile while other 2 sterile, inserted in upper portion of corolla tube; anthers introrse,. (just about all plants have such stamens) Ovary of 3 carpels, 2 much reduced, inferior, 1-locular; ovule solitary, apical, pendulous; style slender, longer than stamens, stigma ?entire. Fruit an achene, enclosed in involucel and surmounted with persistent calyx; seeds with large straight embryo and scanty endosperm.

Two  genera, with about 13 species; distributed from Asia to the eastern Mediterranean with two  genera with nine (or 10) species in China.

 

1a.   Fertile stamens 4; calyx limb oblique; fertile stems always coming from below rosettes; plants not thistle-like  1. Acanthocalyx

1b.   Fertile stamens 2; calyx limb 2-lipped; fertile stems not from below rosettes (this is not clear; are the stems terminal, or what?; plants thistle-like .....................................................................................  2. Morina

 

1. ACANTHOCALYX (de Candolle) M. J. Cannon, Bull. Brit. Mus. Nat. Hist. (Bot.) 12(1): 9. 1984.

刺续断属  ci xu duan shu

Morina sect. Acanthocalyx de Candolle, Prodr. 4: 644. 1830.

Caudexes woody, often covered with remains of old leaf bases. Flowering stems always coming from below rosettes, usually with 2 lines of hairs continued down internodes. Cauline leaves opposite;. Inflorescences subcapitate, often with one or more whorls of flowers below primary head; involucrate bracts free, connate or sheathing at base; flowers sessile; involucels campanulate, very fragile and papery at anthesis, enlarging and becoming ± leathery in fruit, persistent, with several to numerous teeth. Calyx tubular, oblique at mouth, with a ventral fissure, often with 2 lateral and 3 posterior (I very much prefer the ad/abaxial pair) subspinose teeth. Corolla tubular, somewhat swollen below limb, with 2 posterior and 3 anterior spreading lobes. Stamens 4, ± equal, inserted just below swollen part of corolla tube. Style equal to, or exceeding corolla tube in length; stigma disc-shaped Achenes smooth or rugose, apex somewhat cup-shaped.

Two (or three) species: Himalayas and the Hengduan Mountains; all in western China.

 

1a.   Corolla pink or purple; leaf blades elliptic, lanceolate or linear-lanceolate; calyx 7–15 mm  1. A. nepalensis

1b.   Corolla white or yellowish white; leaf blades linear or linear-lanceolate; calyx 4–7 mm  2. A. alba

 

1. Acanthocalyx nepalensis (D. Don) M. J. Cannon in Bull. Br. Mus. nat. Hist. (Bot.) 12(1): 12. 1984.

刺续断ci xu duan

Morina nepalensis D. Don, Prodr. Fl. Nepal. 161. 1825.

Tap roots fleshy. Caudexes short, woodyFertile stems single or 2–3 in number, , 10–50 cm tall, sparsely pubescent above. Rosulate leaves entire, lanceolate or linear-lanceolate, less frequently linear, 10–25 × 0.5–2.6 cm, both surfaces glabrous, base attenuate, margin entire and usually setose, apex acute or acuminate. Leaves on flowering stems 2–4-paired, elliptic to linear-lanceolate, shorter upward on stems; lowest pair of leaves often much smaller, nearly ovate or spatulate, spineless; uppermost pair of leaves more spiny at base, overlapping but not sheathing. Inflorescence a head, sometimes with one or two separate clusters of flowers below primary head; involucrate bracts ovate, spinose, concave or folded; involucels sessile, tubular-campanulate, 8–10 mm, outside sparsely villose, apex truncate, villose, with more than 15 spines unequal in length. Calyx tubular, green below, purple above, or entirely purple, 7–15 mm; tube ca. 1/2 as long as total length of calyx; fissure large, edges villous and dentate-spiny, spines usually 5, irregularly arranged. Corolla pink or purple, slightly zygomorphic, straight or slightly curved; tube 10–25 × 1.5–3 mm, villous; lobes 5, obcordate, 3–4 mm. Stamens 4, included; filaments sparsely villous, inserted at corolla throat.. Achenes columnar, 4–6 mm, glabrous or puberulous. Fl. Jun–Aug; fr. Jul–Sep.

Grassy slopes, alpine meadows: 28004200 m. W Sichuan, S and SE Xizang, N Yunnan [Bhutan, N India, Nepal].

1a.     Ovary and achene usually glabrous .......................................................  1a. subsp. nepalensis

1b.     Ovary and achene usually puberulous .......................................................  1b. subsp. delavayi

 

1a. Acanthocalyx nepalensis subsp. nepalensis

细叶刺续断(原亚种)  xi ye ci xu duan (yuan ya zhong)

Morina betonicoides Bentham; M. nana Wallich ex Bentham.

Ovary and achene usually glabrous.

Grassy slopes: 2800–4200 m. S Xizang (Cona, Damxung, Gyirong, Tingri, Nyalam, Yadong) [Bhutan, N India, Nepal].

1b. Acanthocalyx nepalensis subsp. delavayi (Franchet) D. Y. Hong, comb. nov.

刺续断(亚种)ci xu duan

Morina delavayi Franchet in Bull. Soc. Bot. France 32:8. 1885; Acanthocalyx delavayi (Franchet) M. J. Cannon; M. bulleyana Forrest & Diels; Barleria crotalaria H. Léveillé; M. nepalensis D. Don subsp. delavayi (Franchet) D. Y. Hong & L. M. Ma; M. nepalensis var. delavayi (Franchet) C. H. Hsing.

Ovary and achene usually puberulous.

Alpine meadows; 3000–4200 m. W Sichuan (Baoxing, Barkam, Daocheng, Emei, Hanyuan, Heishui, Jinchuan, Jiulong, Kangding, Muli, Xiangcheng, Xichang, Yanyuan, Yuexi), SE Xizang (Bomi, Gongbo’gyamda, Mainling, Markam, Nyingchi, Riwoqê, Zayü), N Yunnan (Dali, Dêqên, Dongchuan, Eryuan, Gongshan, Lijiang, Luquan, Qiaojia, Shangri-La, Yangbi, Weixi).

2. Acanthocalyx alba (Handel-Mazzetti) M. J. Cannon, Bull. Brit. Mus. (Nat. Hist.), Bot. 12(1): 14. 1984.

白花刺续断 bai hua ci xu duan

Morina alba Handel-Mazzetti, Anz. Akad. Wiss. Wien, Math.-Naturwiss. Kl. 62: 68. 1925; M. leucoblephara Handel-Mazzetti; M. nepalensis var. alba Y. C. Tang ex C. H. Hsing.

Caudex short, woody. Fertile stems 1–3, 10–40 cm tall,. Rosulate leaves linear or linear-lanceolate, 10–20 × 0.5–1 cm, both surfaces glabrous, base attenuate, margin entire. Leaves on flowering stems 2–4-paired, similar to rosulate leaves but shorter; petioles folded; lowest pair of leaves often much smaller, nearly ovate, usually spineless, with sheaths ca. as long as or longer than leaf blades; uppermost pair of leaves more spiny and much wider at base. Inflorescences a head, sometimes with one or two separate clusters of flowers below primary head; involucral bracts ovate, spinose, concave or folded; involucels sessile, tubular-campanulate, 4–7 mm, apex truncate, villous, with 12–16 spines unequal in length. Calyx green throughout, tubular, 5–8 mm; tube 1–2.5 mm; limb usually with 2 lateral and 3 posterior teeth, margin villous. Corolla white, yellowish, or yellowish green; tube markedly curved, 12–20 mm, densely puberulous above; limb 5-lobed; lobes obcordate, ca. 3 mm... Achenes columnar, 4–6 mm, glabrous to densely puberulous. Fl. Jun–Aug, fr. Jul–Sep.

Subalpine or alpine meadows, forests: 2500–4100 m. SE Gansu (Taohe basin), S Qinghai (Henan, Maqên, Nangqên, Yushu, Zadoi), W Sichuan (W of Songpan, Lixian and Butuo), SE Xizang (Baxoi, Jamda, Mainling, Qamdo, Riwoqê, Sogxian, Yadong, Zayü), NW Yunnan (Dêqên, Eryuan, Heqing, Lanping, Lijiang, Shangri-La, Weixi) [N India].

2. Morina Linnaeus, Sp. Pl. 1: 28. 1753.

  ci shen shu

Asaphes Sprengel; Cryptothladia (Blume) M. J. Cannon; Morina sect. Cryptothladia Blume.

Roots usually thickened, branched. Caudex short, woody, usually covered with remaining leaf bases. Leaves verticillate, 3 or 4(–6), rarely opposite, linear to oblong-lanceolate, entire to pinnatipartite, spinose. Flowers in a terminal head or verticillaster,; involucels campanulate, with 816 spiny teeth unequal in length. Calyx-tube oblique, campanulate; limb 2-lipped, lips 2- or 3-lobed. Corolla tube elongate; limb spreading, 2-lipped; upper lip 2-lobed, lower one 3-lobed. Fertile stamens 2, inserted at corolla throat (is this not a family character?); staminodes 2, at base of corolla tube, cordate. Nectary 1, anterior, at base of corolla tube, 3-lobed. Ovary ;; stigma disc-likeAchenes rugose, columnar, slightly or markedly obliquely truncate.

About 10 species: from the Balkan to Central Asia and E Himalayas; 7 or 8 species in W China.

Cannon, M. J. [1984 in Bull. Br. Mus. nat. Hist (Bot.) 12(1): 1–35] stated: Morina parviflora Karelin & Kirilow [=Cryptothladia parviflora (Karelin & Kirilow) M. J. Cannon] occurred on the border of Kazakhstan with China (Dzungarian Ala-Tau). However, we have not seen any specimen so far which could be identified as this species.

1a.    Involucral bracts connate, forming shallow cups; teeth of involucels much shorter than tube (1/2 or less as long as tube) ..............................................................................................................  5. M. polyphylla

1b.   Involucral bracts free; teeth of involucels at least 2/3 as long as tube.

2a.    Leaves long-petioled, petioles 57 cm; leaf margin spinose, rarely sinuate-dentate  1. M. chlorantha

2b.    Leaves sessile or nearly sessile; leaf margin mostly lobed, less frequently entire.

3a. Leaves entire or slightly toothed; flowers in 20 or more closely packed whorls of 812  4. M. ludlowii

3b. Leaves lobed; flowers in less and usually interrupted whorls.

4a.     Corolla 2–3 cm, strongly exserted from calyx, obviously 5-lobed; staminodes inserted at middle part of corolla-tube.

5a.    Involucrate bracts broad-ovate or orbicular, as long as wide excluding pointed apex; calyx entirely covered by involucrate bracts .....................................................  6. M. longifolia

5b.    Involucrate bracts broadly ovate-lanceolate, much longer than wide; calyx exposed  7. M. coulteriana

4b.     Corolla less than 1 cm, almost hidden by calyx, weakly lobed with 2 or 4 major lobes; staminodes inserted at base of corolla tube.

6a.    Calyx-lobes 4, ovate, rounded at apex; leaves shallowly lobed ........  2. M. chinensis

6b.    Calyx-lobes 46, long-ovate or ovate-lanceolate, usually spiny-pointed at apex; leaves deeply cleft, almost to midrib ...........................................................................  3. M. kokonorica

 

1. Morina chlorantha Diels, Notes Roy. Bot. Gard. Edinburgh 5: 208. 1912.

绿花刺参 hua ci shen

Cryptothladia chlorantha (Diels) M. J. Cannon; Morina chlorantha var. subintegra Pax & K. Hoffmann ex H. Limpricht.

Herbs perennial, 2070 cm tall. Caudex robust, base covered with brown fibrous remaining leaves. Basal leaves rosulate; petiole 57 cm; leaf blade lanceolate to ovate-lanceolate, 1535 × 22.5 cm, base decurrent, apex acuminate, margin spinose, rarely sinuate-dentate, tooth margin spinose. Cauline leaves in 2 or 3 whorls of 4, rarely opposite, similar to rosulate ones, but shorter, shortly petiolate, gradually becoming sessile upward, glabrous. Flowers in 6–8 whorls, whorls compact but separated after anthesis, each whorl with 4 involucral bracts, involucral bracts narrowly ovate, 2.5–3 cm, glabrous, apex acuminate, margin hard-spinose, base even denser; involucels tubular, 1.2–1.5 cm, stalked, outside pubescent, ca. 10-spinose at apex, spines ca. 5 mm, unequal in length. Calyx green, 8–10 mm, both sides pubescent, 2-lipped, each lip 2-lobed, lobes lanceolate, 5–6 mm, long ciliate, apex obtuse or 2 of them spinose. Corolla green-yellow, slightly shorter than calyx, 7–9 mm, outside villous; limb 2-lipped, with 5 lobes. Stamens 4, fertile 2 inserted at upper part of corolla tube; filaments short, villous; staminodes 2, nearly sessile, inserted at base of corolla tube. Gynoecium slightly longer than stamens, but included; stigma capitate, slightly compressed. Achenes purple-brown, oblong, ca. 5 mm, angular and furrowed, apex obliquely truncate. Fl. MayAug, fr. AugSep.

Grassy slopes, forest margins; 2800–4000 m. W Sichuan (Baoxing, Dawo, Jiulong, Kangding), NW Yunnan (Lijiang, Shangri-La).

2. Morina chinensis Y. Y. Pai, Repert. Spec. Nov. Regni Veg. 44: 122. 1938.

刺参  ci shen

Morina parviflora Karelin & Kirilow var. chinensis Batalin ex Diels, Notes Roy. Bot. Gard. Edinburgh 5: 208. 1912, nom. nud.; Cryptothladia chinensis M. J. Cannon; M. lorifolia C. Y. Cheng & H. B. Chen.

Herbs perennial. Caudex short, covered with old, nearly complete leaf bases, simple or sometimes multi-branched. Roots robust, usually not branched. Leaves on sterile stems 6–8, linear-lanceolate, 10–25 × 1–2 cm, glabrous, midrib marked, base decurrent and amplexicaul, apex acuminate, margin irregularly and shallowly lobed, lobes nearly triangular, with 3–9 rigid spines. Fertile stems 15–70 cm tall, glabrous and purple below, purplish and white villous above. Cauline leaves similar to those on sterile stems, but shorter, 5–15 cm, 2- or 3-whorled, each whorl of 4–6 leaves, gradually smaller upward, margin of lobes hard-spiny. Flowers in 6–9 whorls compact at anthesis but separated in fruit, each whorl with 4 involucral bracts and up to 20 flowers; involucral bracts leafy, narrowly ovate, 2.5–3.5 cm, margin densely spiny, apex acuminate; involucels hidden in involucral bracts, campanulate, 1–1.4 cm, apex truncate, margin with 10 spines unequal in length, usually 2 longer, to 6 mm, stalked, outside sparsely villous. Calyx ca. 3 mm above involucel, 2-lipped, 6–10 mm, each lip 2-lobed; lobes rounded, obviously veined, outside glabrous, inside lanate, base barbate. Corolla pale green, shorter than calyx, 6–7 mm; limb 2-lipped, upper lip 2-lobed, lower lip 3-lobed, sparsely villous. Stamens 4; posterior 2 fertile, adnate to upper part of corolla tube; anterior 2 sterile, inserted at base of corolla tube; anthers included. Style slightly longer than stamens; stigma capitate. Achenes brown, oblong, 2–3 mm, rugose, apex obliquely truncate, with persistent calyx, hidden in involucels. Fl. Jul–Aug, fr. Sep.

Alpine meadows with shrubs; 2800–4300 m. SE Gansu (Labrang, Minxian, Xiahe, Yuzhong), Nei Mongol (Mt. Helan), Qinghai (Darlag, Datong, Gangca, Henan, Huangyuan, Maqên, Guide, Mingke, Ping’an, W bank of Qinghai Lake, Qilian, Tianjun, Tongde, Tongren, Yushu), NW Sichuan (Aba, Barkam, Dêgê, Kangding, Litang, Lixian, Sêrtar, Songpan, Taining, Xiaojin, Zoigê), Xizang (Qamdo).

3. Morina kokonorica K. S. Hao, Repert. Spec. Nov. Regni Veg. 40: 215. 1936.

青海刺参 qing hai ci shen

Cryptothladia kokonorica (K. S. Hao) M. J. Cannon; Morina parviflora auct. non Karelin & Kirilow: Anonymous, Icon. Cormophyt. Sin. 4: f. 6089. 1975.

Herbs perennial, 20–70 cm tall. Roots robust, ca. 40 cm, to 3 cm in diam., simple or below branched. Caudexes always covered with brown-black remains of old leaves, usually branched. Stems usually caespitose, simple or with 2 or 3 branches, below ridged, glabrous, above villous. Leaves on sterile shoots 5 or 6, rosulate, linear-lanceolate, 7–20 × 1–1.5 cm, both surfaces glabrous, base decurrent into petiole, margin coarsely and irregularly pinnatifid, with triangular, spinose lobes, apex acuminate. Leaves on fertile stems similar to rosulate ones, often in 2 or 3 whorls each of 4, narrowly lanceolate, smaller upward on stems, base amplexicaul. Fertile stems with up to 8 whorls of 16–20 flowers, whorls compact at anthesis, but separated after anthesis, each whorl with 4 involucral bracts; involucral bracts narrowly ovate, 2–3 cm, margin yellow-spinose, apex acuminate; involucels campanulate, hidden in involucral bracts, 1.2–1.5 cm, obviously veined, stalked, edge with more than 10 hard spines, spines unequal in length, usually 1 or 2 longer, to 7 mm. Calyx cup-shaped, 8–15 mm, ca. 3 mm exserted above involucral bracts, outside glabrous, inside villous, base barbate, 2-segmented, each segment 2- or 3-lobed, thus with 4–6 lobes; lobes lanceolate, apex often spine-pointed. Corolla pale green, 6–8 mm, outside glabrous, shorter than calyx; limb 2-lipped, 5-lobed. Stamens 4; fertile 2 inserted at upper part of corolla tube; filaments short, villous; sterile 2 inserted at base of corolla tube, nearly sessile. Style included, slightly longer than stamens; stigma capitate. Achenes brown, columnar, 6–7 mm, smooth, angular, apex obliquely truncate. Fl. Jun–Aug, fr. Aug–Sep.

● Stony slopes, meadows, floodplains; 3000–4500 m. S Gansu, Qinghai (Mengyuan, W bank of Qinghai Lake, Yushu, Zadoi), W Sichuan (Dêgê, Garzê, Luhuo, Xiangcheng), E and S Xizang (westward to Burang, northwards to Baingoin, Sogxian).

4. Morina ludlowii (M. J. Cannon) D. Y. Hong in Novon

藏南刺参  zang nan ci shen

Cryptothladia ludlowii M. J. Cannon, Bull. Brit. Mus. (Nat. Hist.), Bot. 12: 22. 1984.

Herbs perennial, to 50 cm tall. Caudex thickly covered with fibrous remaining leaf bases. Basal leaves and lower cauline leaves in whorls of 3 or 4, narrowly elliptic, 5–9 × 1–2 cm, adaxially pubescent, base broad, fused together into loose sheaths, margin entire or slightly toothed, margin ciliate, spinose; upper cauline leaves similar to lower ones, petioles fused, forming shorter sheaths, uppermost whorl often barely connate. Flowering stems short at first, 2–3 cm, elongating rapidly, terete, or slightly grooved, densely covered with whitish, mostly retrorse hairs. Flowers in 20 or more closely packed whorls of 8–12; involucral bracts leafy, regularly decussate, lowest whorl of involucral bracts similar to leaves but smaller, usually 3-lobed, lobes irregularly spinose-dentate, acute or cuspidate, central lobe larger than lateral ones, often reddish or deeply coloured; involucels tubular-campanulate, villous, often with stalked or sessile glands, 4–12-dentate, one tooth often larger than others, 4–6 mm, ca. 2 mm broad at base, laminate, becoming larger and rigid in fruit. Calyx tube tubular-campanulate, 2.5–4 mm at anthesis; limb 2-fid, lobes shallowly divided, lobules rounded to apiculate, glabrous outside, villous and glandular inside. Corolla included, ± tubular, outside glabrous, 4-lobed, posterior lobe ± fimbriate, anterior lobe entire, exceeding posterior, inside hispid, lateral 2 lobes very small. Stamens 4, fertile 2 with short filaments, inserted at middle part of corolla tube; filaments with a tuft of hairs below anthers; staminodes 2, at base of corolla tube, shortly stalked. Style slightly shorter than corolla tube; stigma disc-shaped. Achenes subglobose, 3–5 mm, abaxially with a shallow furrow, apex slightly obliquely truncate, obtusely pointed, somewhat rugose. Fl. May–Aug; fr. Aug–Sep.

Thickets, stony hillsides; 3700–4300 m. S Xizang (Cona) [Bhutan, NE India].

5. Morina polyphylla Wallich ex de Candolle, Prodr. 4: 644. 1830.

多叶刺参  duo ye ci shen

Cryptothladia polyphylla (Wallich ex de Candolle) M. J. Cannon, Bull. Brit. Mus. Nat. Hist. Bot. 12(1): 20. 1984.

Herbs perennial, up to 50 cm tall. Roots ± lignified, up to 2 cm in diam. Caudexes thickly covered with brown fibrous remains of old leaves, up to 3 cm in diam. Leaves on sterile shoots  rosulate, with long and narrow petioles, linear to linear-lanceolate, up to 40 cm, 1.5–4 cm broad, pinnatifid, segments usually 3–5-fid, lobules with an apical spine, glabrous or sparsely pubescent; cauline leaves on flowering shoots in whorls of (3–)4–6, similar to rosulate ones but smaller. Flowering stems terete, or furrowed, glabrous below, white-hairy above, often purple. Verticillasters 8–10 cm, of many whorls, compact; involucrate bracts in whorls of (3–)4–6, connate at base, forming a cup-shaped structure, free tips linear or linear-lanceolate, with lateral spines, tapered into a robust spine; involucels 6–13 mm, cylindrical, toothed, usually pilose, rarely nearly glabrous, with many sessile and a few stalked glands, teeth usually 8–10, apex tapered to a terminal bristle or spine, 1 or 2 longer than rest. Calyx tubular or campanulate, 7–12 mm; tube 4–5 mm, usually densely pilose and glandular inside; limb 2-lobed, lobes patent, white or lavender, deeply divided, 4 lobules apically rounded or spiny-tipped. Corolla pink or white, as long as involucel, nearly tubular; limb 4-lobed. Fertile stamens 2, inserted at middle of corolla-tube; sterile 2 inserted at base of corolla-tube. Style as long as corolla-tube; stigma disc-shaped. Achenes 4.8–5.5 × 2.8–3 mm, plano-convex, obliquely truncate, surmounted with persistent and enlarged calyx. Fl. May–Jul; fr. Aug–Sep.

Grassy slopes, alpine meadows, or rarely forests; 2600–4700 m. S Xizang (Yadong) [Bhutan, N India (Sikkim), Nepal].

6. Morina longifolia Wallich ex de Candolle, Prodr. 4: 644. 1830.

长叶刺参  chang ye ci shen

Herbs perennial, to 80 cm tall. Caudex woody, top often covered with fibrous remains of old leaves. Leaves on sterile shoots rosulate, rosulate leaves and lower cauline leaves with petioles fused, forming long sheaths 2.5–4.5 cm; leaf blade linear to linear-lanceolate, to 40 × 4 cm, both surfaces glabrous, regularly coarsely lobed to nearly half, each lobe with 2 or 3 spines; upper cauline leaves similar, usually 3 per whorl, smaller, sheaths to 8 mm, or petioles barely connate. Flowering stems erect, to 90 cm, glabrous, slightly ridged to terete below, pubescent to villous above. Flowers in (4–)7–9(–11) whorls, pedicellate or sessile; upper whorls confluent but more separated in fruit; involucral bracts broadly ovate to orbicular, with spiny tip, bases markedly overlapping, adaxially sparsely hairy, abaxially villous, margins often with numerous short spiny teeth and ciliate; involucels tubular, 7–10 × 2.5–4 mm, glandular hairy, with 5–13 teeth, 2 much exceeding others. Calyx glabrous, or with short adpressed hairs on tips; lobes shallowly 2-lobed; lobules rounded or rarely slightly apiculate; tube 4.5–6 × 2.5–4 mm at anthesis. Corolla white, becoming pink to deep red; tube 20–30 × 1–2 mm, with many glandular and a few simple hairs outside; limb 2-lipped, lobes of lower lip ± spreading, central lobe ca. 7 × 3 mm, upper lip ± spreading. Stamens 4; filaments of fertile two 1–3 mm, with a tuft of hairs below anthers, inserted at throat of corolla; staminodes 2, minute, inserted 2–3.5 mm below mouth of corolla under anterior lip. Style slightly longer than stamens; stigma disc-shaped. Achenes rugose on abaxial side, with a deep furrow and prominent longitudinal veins on adaxial side, apex slightly obliquely truncate and crenulate. Fl. Jun–Sep, fr. Sep–Oct. 2n=16, 34

Grassy slopes, thickets; 3000–4300 m. S Xizang (Gyirong) [Bhutan, India, Nepal, Pakistan].

7. Morina coulteriana Royle, Illus. Bot. Himal. 245. 1835.

黄花剌参  huang hua ci shen

Herbs perennial, robust. Rhizomes woody, often covered with remains of old leaf base. Shoots of two types; sterile and fertile. Sterile shoots dwalf; fertile shoots (flowering stems) up to over 1 m tall, ridged and glabrous below, sparsely villose and purple above. Leaves on sterile shoots rosulate, 20–25 × 2.5 cm, glabrous, pinnatilobate, with spiny, simple or compound teeth; petioles fusing to form a deep sheath of 8–15 mm; lower cauline leaves similar, in whorls of 3–4 (5); upper cauline leaves also similar but smaller. Flowers in 6–8 whorls; lower 3–5 whorls separate while upper 3 or 4 confluent; flowers numerous per whorls; involucral bracts in whorls of 3 or 4, broadly ovate-lanceolate, up to 8 × 2 cm, spiny, obviously net-veined, upper ones smaller; involucels nearly cylindrical, villose, 4–9 × 2.5–4 mm, with 9–12 teeth, 2 much longer than others. Calyx glabrous to villose, with a large tuft of hairs at base, tubular-campanulate, 3–6.5 × 2–4 mm; limb 2-lipped, lips deeply bifid for 2–5 mm, apex acute or apiculate, often extended beyond tip to form a spine. Corolla yellow or greenish yellow, tube villose, 25–30 × 1–2 mm; limb 2-lipped, upper lip 2-lobed, lower one 3-lobed, median lobe of lower lip 5.5–10 × 3.5–6 mm. Fertile stamens 2; filaments 2–3 mm, with a tuft of hairs below anthers; staminodes minute, heart-shaped. Style slightly longer than stamens; stigma disc-shaped. Achenes convex on adaxial side, rugose on abaxial side, with very deep longitudinal furrows. Fl. Jun–Aug; fr. Aug–Sep.

Grassy or shrubby slopes; 3000–3700 m. S Xinjiang,, Pakistan, Tajikistan, Uzbekistan].