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1. Asteraceae

菊科 ju ke

Authors: Zhu Shi, Yilin Chen, Yousheng Chen, Yourun Lin(Ling Yuou-ruen), Shangwu Liu, Xuejun Ge, Tiangang Gao, Shixin Zhu, Ying Liu, Christopher J. Humphries, Qiner Yang, Eckhard von Raab-Straube, Michael G. Gilbert, Bertil Nordenstam, Norbert Kilian, Luc Brouillet, Irina D. Illarionova, D. J. Nicholas Hind, Charles Jeffrey, Randall J. Bayer, Jan Kirschner, Werner Greuter, Arne A. Anderberg, John C. Semple, Jan Štěpánek, Susana Edith Freire, Ludwig Martins, Hiroshige Koyama, Takayuki Kawahara, Leszek Vincent, Alexander P. Sukhorukov, Evgeny V. Mavrodiev & Günter Gottschlich

Saussurea stella

Credit: Harvard University Herbaria

Herbs, subshrubs, or shrubs, rarely trees or climbers, bisexual, monoecious, or sometimes dioecious, many with laticiferous cells or canals and/or resinous ducts. Leaves often in a basal rosette; cauline leaves usually alternate, more rarely (in China) opposite or whorled, sessile or petiolate, without stipules; petiole sometimes auriculate at base; leaf blade entire to variously lobed or divided. Florets bisexual (perfect), female (pistillate), or functionally male (functionally staminate), solitary or few to many enclosed in an involucre of 1- to many-seriate phyllaries (involucral bracts) to form a capitulum; calyculus of outer bracts sometimes present and often differing markedly from phyllaries. Capitula solitary or few to many arranged in variously formed synflorescences, homogamous or heterogamous, discoid, disciform, radiate, subradiate, radiant, or ligulate; receptacle usually flattened, sometimes slightly concave or convex, rarely conical, smooth or alveolate, paleate (scaly) or epaleate, with bristles, scales, hairs, or naked. Corolla (3-) 5-merous, gamopetalous, tubular or tubular-filiform, and regular, bilabiate, radiate, or ligulate. Stamens (4 or)5, inserted within corolla tube; filaments adnate to proximal part of corolla; anthers basifixed or dorsifixed, usually coalescent into a tube, base caudate or not, apex with (rarely without) a sterile, ovate or lanceolate appendage; pollen grains usually tricolporate, echinate or sometimes lophate or spinulate, often caveate. Style apically bifid (rarely entire); style branches variously shaped, with or without an apical appendage, hairs, or papillae. Ovary inferior, 1-loculed; ovule 1, basal, anatropous. Fruit an achene (cypsela). Pappus consisting of 1 to many rows of scales or bristles, or absent. Seed erect; endosperm scanty; embryo straight.

Between 1,600 and 1,700 genera and ca. 24,000 species: cosmopolitan (except Antarctica); 15 tribes (one introduced), 248 genera (18 endemic, 49 introduced), and 2,336 species (1,145 endemic, 109 introduced) in China.

Taxonomic discussion on the Asteraceae is included under the relevant tribes. The following notes apply to the whole family. Modes of reproduction in the Chinese Asteraceae are known only fragmentarily. As regards the world diversity of the family, only about 12% of genera have been studied using cytoembryology to identify the reproduction system. In 3%, agamospermy was found to occur. Probably the most common reproduction system is allogamy with a sporophytic self-incompatibility; occasional selfing is also possible (stigma ageing or mentor effect). More or less obligate autogamy is relatively rare. Other uncommon reproduction systems include gynodioecy, a very rare dioecy (with heterogametic females). Asexual reproduction is a widespread phenomenon in the Asteraceae, with a wide range of mechanisms. In addition to the rare adventitious embryony, there are common types of autonomous apomixis, agamospermy (apospory, diplospory) associated with various forms of embryogenesis (parthenogenesis, apogamety). All the types of reproduction are to be considered in the evaluation of population variation and for taxonomic conclusions because the prevailing type of reproduction substantially influences the variation limits of taxa.

Ling Yong, Chen Yi-ling, Shih Chu, Chen Feng-hwai, Chang Chao-chien, Tseng Yong-qian, Hu Chi-ming & Huang Xiu-lan. 1979. Compositae (2): Inuleae-[Heliantheae]-Helenieae. In: Ling Yong, ed., Fl. Reipubl. Popularis Sin. 75: i-xviii, 1-422; Shih Chu & Fu Gou-xun. 1983. Compositae (3): Anthemideae [1]. In: Ling Yong & Shih Chu, eds., Fl. Reipubl. Popularis Sin. 76(1): i-viii, 1-149; Ling Yong, Chen Yi-ling & Shih Chu. 1985. Compositae (1): Carduoideae Kitam.: Vernonieae-[Eupatorieae]-Astereae. In: Ling Yong & Chen Yi-ling, eds., Fl. Reipubl. Popularis Sin. 74: i-xiv, 1-388; Shih Chu. 1987. Compositae (7): Echinopeae ["Echinopsideae"], Cynareae. In: Ling Yong & Shih Chu, eds., Fl. Reipubl. Popularis Sin. 78(1): i-ix, [t.] 1-7, 1-226; Liu Shang-wu. 1989. Compositae (4): Senecioneae: Senecioninae [Farfugium, Ligularia, Cremanthodium]. In: Ling Yong & Liu Shang-wu, eds., Fl. Reipubl. Popularis Sin. 77(2): i-vii, 1-188; Ling Yourun (= Ling Yeou-ruenn). 1991. Compositae (3): Anthemideae (2). In: Ling Yong & Ling Yourun (= Ling Yeou-ruenn), eds., Fl. Reipubl. Popularis Sin. 76(2): i-xii, 1-316; Tseng Yung-chien. 1996. Compositae (9): Mutisieae. In: Tseng Yung-chien, ed., Fl. Reipubl. Popularis Sin. 79: i-iv, 1-113; Shih Chu. 1997. Compositae (10): Cichorioideae: Lactuceae. In: Ling Yong & Shih Chu, eds., Fl. Reipubl. Popularis Sin. 80(1): i-xiv, 1-342; Chen Yiling. 1999. Compositae (5): Senecioneae, Calenduleae. In: Chen Yiling, ed., Fl. Reipubl. Popularis Sin. 77(1): i-xiii, 1-369; Ge Xuejun, Ling Yeouruenn & Zhai Datong. 1999. Compositae (11): Cichorieae: Taraxacum. In: Ling Yeouruenn & Ge Xuejun, eds., Fl. Reipubl. Popularis Sin. 80(2): i-iv, 1-94; Shih Chu & Jin Shuying. 1999. Compositae (8): Cynareae [Saussurea]. In: Chen Yiling & Shih Chu, eds., Fl. Reipubl. Popularis Sin. 78(2): i-x, 1-243.


Key to tribes

1 Capitula of 2 kinds, either with only functionally male florets or with only female florets; flowering not precocious   (2)
+ Capitula all alike, homogamous or heterogamous, rarely plants subdioecious and then flowering precocious   (6)
       
2 (2) Plants monoecious, with male and female capitula on same plant.   14 Heliantheae (p. 852)
+ Plants dioecious, with male and female capitula on different plants   (3)
       
3 (3) Leaves spiny.   3 Carlineae (Tugarinovia; p. 39)
+ Leaves not spiny   (4)
       
4 (4) Shrubs.   1 Mutisieae (Myripnois, rarely Pertya; p. 9)
+ Herbs   (5)
       
5 (5) Phyllaries papery; female florets filiform.   10 Gnaphalieae (Antennaria; p. 774)
+ Phyllaries herbaceous; female florets tubular, apically 3- or 4-lobed.   246 Cavea (p. 892)
       
6 (6) Fertile achenes with conspicuous stipitate glands.   1 Mutisieae (Adenocaulon; p. 9)
+ Fertile achenes without conspicuous stipitate glands   (7)
       
7 (7) Capitula homogamous and all florets ligulate, i.e., corollas with an expanded abaxial, apically 5-toothed, 5-lobed, or 5-cleft limb; latex (milky juice) present.   5 Cichorieae (p. 195)
+ Capitula heterogamous, or if homogamous then corollas tubular and 5-lobed or zygomorphic and pseudoligulate (5-lobed with one deeper cleft or sinus); plants without or rarely with latex   (8)
       
8 (7) Capitula homogamous; corollas pseudoligulate, 5-lobed and zygomorphic   (9)
+ Capitula heterogamous (marginal florets with a strap-shaped, entire or 2-4-toothed or 2-4-lobed outer limb) or homogamous and corollas 3-5-lobed, actinomorphic   (10)
       
9 (9) Capitula in bracteate glomerules; phyllaries 8, decussate; style branches long.   6 Vernonieae (Elephantopus, Pseudelephantopus; p. 354)
+ Capitula not in bracteate glomerules; phyllaries not decussate; style branches very short or scarcely bilobed.   1 Mutisieae (Ainsliaea, Pertya; p. 9)
       
10 (10) Capitula with all or some florets bilabiate or pseudobilabiate (slightly zygomorphic)   1 Mutisieae (Gerbera, Leibnitzia, Nouelia, Piloselloides; p. 9)
+ Capitula lacking bilabiate florets   (11)
       
11 (11) Capitula each with only 1 floret, aggregated into a terminal globose pseudocephalium; leaves spiny.   2 Echinopeae (p. 33)
+ Capitula with more than 1 floret, or if with only 1 floret then leaves not spiny   (12)
       
12 (11) Leaves at least below synflorescence opposite or mostly so   (13)
+ Leaves all alternate   (14)
       
13 (13) Style branch tips longer than stigmatic lines, prominent; capitula discoid; corollas never yellow.   15 Eupatorieae (p. 879)
+ Style branch tips shorter than stigmatic lines, or absent; capitula radiate, disciform, or discoid; corollas often yellow.   14 Heliantheae (p. 852)
       
14 (14) Style shaft and branches glabrous.   1 Mutisieae (Leucomeris; p. 9)
+ Style shaft glabrous or with hairs or evident papillae in distal part and/or style branches with hairs or evident papillae abaxially or apically or on an apical appendage   (15)
       
15 (14) Disk corollas 3- or 4-merous   (16)
+ Disk corollas or all corollas 5-merous   (18)
       
16 (15) Receptacle with scales.   248 Symphyllocarpus (p. 893)
+ Receptacle without scales   (17)
       
17 (17) Achenes dorsiventrally compressed, elliptic in cross section   9 Anthemideae (p. 653)
+ Achenes subterete, broadly ribbed.   247 Centipeda (p. 892)
       
18 (15) Style shaft with a papillose-pilose thickening below branches; leaves spiny, at margin finely serrulate-spinulose, or not spiny   (19)
+ Style shaft without a papillose-pilose thickening below branches; leaves neither spiny nor at margin finely serrulate-spinulose   (20)
       
19 (19) Achene densely hairy, capitula homogamous, plants biennial or perennial, and leaves never grasslike; pappus inserted directly on upper edge of achene, not surrounded by a rim.   3 Carlineae (p. 39)
+ Achene glabrous or subglabrous, rarely densely hairy but then capitula radiant, plants annual, or leaves grasslike; pappus insertion surrounded by a ± conspicuous rim or crown.   4 Cardueae (p. 42)
       
20 (20) Achenes large, heteromorphic, outer cymbiform or curved, inner strongly curved to circular; receptacle epaleate; pappus absent.   11 Calenduleae (p. 819)
+ Achenes small, columnar or compressed, straight or only slightly curved, homomorphic or only slightly heteromorphic, if large then receptacle paleate; pappus present or absent   (21)
       
21 (20) Phyllaries rather dry, either with distinct scarious margin or papery, brownish, yellowish, or whitish, never herbaceous and green throughout   (22)
+ Phyllaries herbaceous, all or at least outer ones green throughout, except at very apex   (23)
       
22 (22) Anthers rounded at base; phyllaries with distinct pale or brownish scarious margin.   9 Anthemideae (p. 653)
+ Anthers tailed at base; phyllaries papery, whitish, brownish, or yellowish, especially in distal part.   10 Gnaphalieae (p. 774)
       
23 (23) Style branches long, slender, subulate, without an apical appendage, hairy abaxially and with stigmatic papillae over entire adaxial surface; capitula homogamous.   6 Vernonieae (p. 354)
+ Style branches not with above combination of characters; capitula homogamous or heterogamous   (24)
       
24 (23) Receptacle paleate   (25)
+ Receptacle epaleate   (27)
       
25 (25) Achenes with a carbonized layer in pericarp, thereby black or streaked with black.   14 Heliantheae (p. 852)
+ Achenes without a carbonized layer in pericarp, thereby usually not black   (26)
       
26 (26) Phyllaries linear.   12 Inuleae (Buphthalmum; p. 820)
+ Phyllaries broadly elliptic to oblanceolate.   13 Athroismeae (p. 851)
       
27 (24) Style branches terminating in a triangular to subulate appendage distal to stigmatic lines   (28)
+ Style branches without an apical appendage distal to stigmatic lines, apically rounded or truncate   (29)
       
28 (28) Phyllaries uniseriate, but sometimes with an outer series of much shorter bracts (calyculus); involucre cylindric.   7 Senecioneae (p. 371)
+ Phyllaries 2- to several seriate, if subuniseriate then involucre saucer-shaped to subglobose.   8 Astereae (p. 545)
       
29 (29) Phyllaries uniseriate, but sometimes with an outer series of much smaller bracts (calyculus); capitula never aggregated into a compact synflorescence.   7 Senecioneae (p. 371)
+ Phyllaries 2- or 3-seriate or imbricate in several series; capitula sometimes aggregated into a compact synflorescence   (30)
       
30 (29) Corolla bearing numerous robust uniseriate multicellular hairs.   246 Cavea (p. 892)
+ Corolla without such hairs   (31)
       
31 (31) Stigmatic papillae confined to 2 submarginal lines often confluent at apex.   12 Inuleae (p. 820)
+ Stigmatic papillae covering entire adaxial surface of style branches.   7 Senecioneae (Doronicum; p. 371)

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