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FOC | Family List | FOC Vol. 20-21 | Asteraceae


菊苣族 ju ju zu

Authors: Zhu Shi, Xuejun Ge, Norbert Kilian, Jan Kirschner, Jan Štěpánek, Alexander P. Sukhorukov, Evgeny V. Mavrodiev & Günter Gottschlich

Annual to perennial, acaulescent, scapose, or caulescent herbs, more rarely subshrubs, exceptionally scandent vines, latex present. Leaves alternate, frequently rosulate. Capitulum solitary or capitula loosely to more densely aggregated, sometimes forming a secondary capitulum, ligulate, homogamous, with 3-5 to ca. 300 but mostly with a few dozen bisexual florets. Receptacle naked, or more rarely with scales or bristles. Involucre cylindric to campanulate, ± differentiated into a few imbricate outer series of phyllaries and a longer inner series, rarely uniseriate. Florets with 5-toothed ligule, pale yellow to deep orange-yellow, or of some shade of blue, including whitish or purple, rarely white; anthers basally calcarate and caudate, apical appendage elongate, smooth, filaments smooth; style slender, with long, slender branches, sweeping hairs on shaft and branches; pollen echinolophate or echinate. Achene cylindric, or fusiform to slenderly obconoidal, usually ribbed, sometimes compressed or flattened, apically truncate, attenuate, cuspidate, or beaked, often sculptured, mostly glabrous, sometimes papillose or hairy, rarely villous, sometimes heteromorphic; pappus of scabrid [to barbellate] or plumose bristles, rarely of scales or absent.

About 95 genera and ca. 2,500 species (and at least 8,000 apomictic and hybrid taxa): chiefly in the temperate to subtropical zones of the N Hemisphere, and extending to tropical and S Africa, tropical Asia, Australia and New Zealand, and South America, also on SE Pacific and mid-Atlantic archipelagos; 35 genera (two endemic, one introduced) and 388 species (190 endemic, 12 introduced) in China.

1 Pappus absent in all or in outer achenes, or in all achenes present but of minute (≤ 0.3 mm) scales or very short (≤ 1.5 mm) bristles   (2)
+ Pappus well developed in all achenes, of soft or stiff bristles, usually more than 1/2 as long as achene   (6)
2 (1) Pappus a tiny crown of fimbriate 0.1-0.3 mm scales; florets blue.   83 Cichorium
+ Pappus absent in all or in outer achenes, or of very short (≤ 1.5 mm) bristles; florets yellow   (3)
3 (2) Achene dimorphic, outer achenes apically attenuate or shortly beaked, inner ones with a long slender beak strongly exceeding involucre; pappus in all or in inner achenes present   (4)
+ Achene homomorphic, never beaked; pappus in all achenes absent   (5)
4 (3) Outer achenes columnar, curved, with appressed hairs, apically attenuate; pappus of smooth to scabrid. 2-1.5 mm bristles, shorter in outer, longer in inner achenes; inner phyllaries hardened in fruit.   70 Garhadiolus
+ Outer achenes broadly obconical, compressed, lateral ribs strongly winglike and enlarged, shortly beaked; pappus absent in outer achenes, mostly present and of 3-5 mm bristles in inner ones; inner phyllaries not hardened in fruit.   69 Heteracia
5 (3) Achene body columnar-scorpioid, ca. 1 cm, abaxially with antrorse and apically with retrorse rigid hooked needlelike spines; leaves undivided, narrowly grasslike.   53 Koelpinia
+ Achene body narrowly ellipsoid, subcompressed, to ca. 5 mm, with 0 or (1 or)2-4 main ribs apically prolonged into 0.2-2.2 mm slender hooked appendages; leaves pinnatipartite or pinnatisect.   67 Lapsanastrum
6 (1) Pappus bristles all or at least inner ones plumose but often apically scabrid   (7)
+ Pappus bristles never plumose, all smooth to scabrid   (12)
7 (6) Plumose bristles stiffly fimbriately plumose, fimbriae not intertwining; florets yellow or white; leaves never grasslike   (8)
+ Plumose bristles softly or lanately fimbriately plumose, fimbriae conspicuously intertwining; florets of some shade of yellow, white, blue, or purple; leaves often grasslike   (9)
8 (7) Receptacle with linear scarious scales ± as long as involucre and enclosing base of florets; stem, leaves, and involucre with rigid simple hairs.   81 Hypochaeris
+ Receptacle naked; stem, leaves, and involucre with simple and 2-hooked hairs.   82 Picris
9 (7) Phyllaries in 1(or 2) series, outer series if present never leaflike.   55 Tragopogon
+ Phyllaries in several series, or in 2 series but then outer series leaflike and longer than inner series   (10)
10 (9) Phyllaries in 2 series, outer phyllaries herbaceous, leaflike, usually longer than inner phyllaries, inner usually 5.   54 Epilasia
+ Phyllaries in several series, outer phyllaries never leaflike and longer than inner ones   (11)
11 (10) Leaves undivided, parallel-veined, grasslike or more rarely also lanceolate to ovate.   51 Scorzonera
+ Leaves pinnately divided.   52 Podospermum
12 (6) Pappus white, of numerous fine cottony outer bristles intermixed with some thicker inner ones.   63 Sonchus
+ Pappus white or grayish, yellowish, straw-colored, brownish, or reddish brown, of bristles ± equal in diam. and stiffness   (13)
13 (12) Leaves, axes, or involucres with various types of hairs, but including stellate hairs and/or multiseriate hairs with projecting cell apices   (14)
+ Leaves, axes, or involucres glabrous or hairy, but never with stellate hairs or multiseriate hairs with projecting cell apices   (15)
14 (13) Achene 2.5-5 mm, with 8-10 equal ribs apically confluent in an obscure ring.   84 Hieracium
+ Achene 1-2 mm, with apically distinct ribs not confluent in an obscure ring.   85 Pilosella
15 (13) Capitulum solitary on a hollow scape (without nodes, leaves, or bracts) single or few from a leaf rosette; achene beak usually longer than achene body, body at least apically mostly spinulose, scaly, and/or tuberculate and usually with an apical cone.   71 Taraxacum
+ Capitula usually few to numerous, either fascicled or aggregated amid a leaf rosette, or on a branched stem; if capitulum rarely solitary then not on a hollow scape and achene not as above   (16)
16 (15) Rosulate herbs with several to many capitula either on short, wiry unbranched axes (peduncles) directly from and fasciculate amid leaf rosette, or sessile to very shortly pedunculate and densely aggregated on an apically ± inflated, flat, convex, or more rarely cylindrically elongate rosette shoot   (17)
+ Capitula few to numerous on a branched stem a few cm to more than 2 m tall; rarely capitulum solitary on an unbranched stem   (22)
17 (16) Capitula with 3-6 florets, sessile or on peduncle shorter than involucre; phyllaries in 1 series, hardened in fruit, outer phyllaries absent, inner ones connate in basal part   (18)
+ Capitula with 4-30 florets, peduncle shorter or longer than involucre; phyllaries in more than 1 series, herbaceous in fruit, outer phyllaries 2 or more, inner ones basally connate or free   (19)
18 (17) Florets yellow or pale to medium purplish; achene obconical, compressed, with 5 ribs, apex truncate; pappus bristles caducous.   77 Syncalathium
+ Florets purplish red to blue; achene obovoid, compressed, with winglike lateral ribs and 1 slender rib on either side, apex constricted into a ca. 0.5 mm fragile thin beak; pappus caducous with pappus disk (M. souliei).   58 Melanoseris
19 (17) Florets blue to bluish purple (D. amoena and D. gombalana).   76 Dubyaea
+ Florets yellow or rarely white   (20)
20 (19) Capitula with 4 or 5 florets, with a peduncle shorter than to ± as along as involucre and from an apically ± inflated, flat, convex, or more rarely cylindrically elongate rosette shoot (Soroseris spp. 3-7).   80 Soroseris
+ Capitula with 12-30 florets, with a peduncle usually longer than involucre, from a non-inflated rosette shoot   (21)
21 (20) Synflorescence hemispheric, of many densely crowded capitula surrounded by a rosette of orbicular to ovate leaves abruptly contracted into an unwinged petiole-like basal portion; florets yellow or white (Soroseris spp. 1 and 2).   80 Soroseris
+ Synflorescence loosely corymbiform with few to several capitula amid a rosette of ± spatulate leaves very gradually attenuate toward base; florets yellow.   66 Youngia
22 (16) Achene isodiametric and with ribs of ± equal shape and size; pappus white, rarely pale yellowish   (23)
+ Achene (sometimes except inner ones) somewhat to distinctly compressed and/or with ± unequal ribs; pappus white, yellowish, or brownish   (25)
23 (22) Achene with 10 very prominent ± winglike ribs.   74 Ixeris
+ Achene with 10-20 ± prominent but never winglike ribs   (24)
24 (23) Plants delicate; involucre narrowly cylindric, longest outer phyllary ≤ 1/4(-1/3) of inner ones, inner phyllaries abaxially glabrous; capitula with 5-15 florets; achene pale brown, with 10 ribs.   72 Askellia
+ Plants usually robust; involucre cylindric to campanulate, longest outer phyllary 1/4-2/3 or more of inner ones, inner phyllaries abaxially usually with simple and/or glandular hairs along midvein; capitula with (8-)20-70 (and more) florets; achene of some shade of brown, with 10-20 ribs.   65 Crepis
25 (22) Pappus yellowish, straw-colored, brownish, or grayish and achene never strongly compressed with dominant lateral ribs and/or with capillaceous beak   (26)
+ Pappus white, exceptionally faintly yellowish or brownish but then achene strongly compressed with dominant lateral ribs and/or with capillaceous beak   (33)
26 (25) Florets reddish to bluish purple, or blue   (27)
+ Florets yellow, or rarely whitish to pale purplish or pale greenish   (28)
27 (26) Involucre broadly campanulate; capitula with many (usually 50-70) florets; stem in upper half and peduncles often with long stiff blackish hairs.   76 Dubyaea
+ Involucre narrowly cylindric to narrowly campanulate; capitula with 5-30 florets; stem in upper half and peduncles glabrous.   56 Faberia
28 (26) Involucre 4.5-8 mm; capitula erect   (29)
+ Involucre 10-20 mm; capitula often nodding at anthesis   (30)
29 (28) Achene with short slender beak; pappus yellowish to yellowish brown.   73 Ixeridium
+ Achene apically attenuate but not beaked; pappus ash-gray or yellowish brown.   66 Youngia
30 (28) Synflorescence secundly racemiform (Y. racemifera).   66 Youngia
+ Synflorescence corymbiform or paniculiform, or capitula 1 or 2   (31)
31 (30) Stem, branches, and phyllaries with conspicuous stiff yellowish brown, reddish, purplish brown, or blackish mostly glandular hairs, or glabrous and stem with 1 or 2 nodding capitula only.   76 Dubyaea
+ Stem, branches, and phyllaries glabrous and stem with some to many capitula, or ± hairy but never with conspicuous stiff mostly glandular hairs   (32)
32 (31) Leaves grasslike.   78 Hololeion
+ Leaves pinnately lobed or, if undivided, ovate to triangular-ovate.   79 Nabalus
33 (25) Plants broomlike and/or achene with body apically scaly and/or tuberculate and beaked (sometimes very shortly).   64 Chondrilla
+ Plants never broomlike, achene beaked or not but body never scaly or tuberculate   (34)
34 (33) One of following features applies: 1) achene ± compressed with ± dominating or broadened lateral ribs; 2) pappus double and additionally with an outer series of minute hairs (usually ≤ 0.3 mm); 3) florets purplish, bluish, or exceptionally white   (35)
+ Achene ± isodiametric or ± compressed but not with ± dominating or broadened lateral ribs; pappus always single; florets always of some shade of yellow   (39)
35 (34) Achene black or reddish-purplish (if rarely pale then plant scandent), fusiform to cylindric, without dominating lateral ribs, and apex truncate, attenuate, or shortly (ca. 1 mm) beaked; florets some shade of purple or blue; pappus simple; involucre narrowly cylindric   (36)
+ Achene pale to dark brown, more rarely blackish or reddish brown, gray, or olive green, subcylindric, ± ellipsoid to obovoid, or ± fusiform, usually with, rarely without, dominating or broadened (sometimes winglike) lateral ribs and apex truncate to long filiform beaked; pappus simple or double; florets yellow, bluish, or purplish, rarely white; involucre narrowly cylindric, cylindric, campanulate, or broadly campanulate   (37)
36 (35) Achene black, fusiform, weakly compressed, apex attenuate to shortly beaked and often pale.   59 Paraprenanthes
+ Achene reddish-purplish (rarely pale and plant scandent), fusiform to cylindric, compressed, apex truncate.   60 Notoseris
37 (35) Achene always strongly compressed, ellipsoid to obovoid, dominating lateral ribs sometimes even winglike, apex with a short stout or with a long filiform beak; pappus always simple; involucre narrowly cylindric at anthesis, inner phyllaries 5 or 8, or if 3 or 4 then plant a subshrub with whitish, rigid, intricately and divaricately branched stems; stems and branches usually glabrous, at least in upper half; capitula always erect; florets some shade of yellow, or blue, bluish purple, purple, or white.   61 Lactuca
+ Achene ± compressed, dominating lateral ribs never winglike, apex truncate, attenuate, or with a short stout or slender but never long filiform beak; pappus usually double, more rarely outer series with minute hairs indistinct or missing; involucre narrowly cylindric to broadly campanulate, inner phyllaries 3 to many; capitula erect or nodding; stems and branches glabrous, or hispid, hirsute, or otherwise hairy; florets mostly some shade of blue or purple, exceptionally white, very rarely yellow   (38)
38 (37) Involucre with 5-10 inner phyllaries, 6-10(-12) mm and narrowly cylindric, or cylindric, 11-13 mm, and with longest outer phyllary 1/2-3/4 of inner ones, or to 15 mm, broadly cylindric to campanulate, and herb tall with many-capitellate narrowly racemiform synflorescence to 50 cm; florets always bluish or purplish; achene 4-5 mm.   57 Cicerbita
+ Involucre broadly cylindric to broadly campanulate, mostly exceeding 15 mm and never with many-capitellate narrowly racemiform synflorescence to 50 cm, or rarely narrowly cylindric but then with either only 3 or 4 inner phyllaries, or > 13 mm, or strongly hirsute, or florets yellow, or achene ≥ 6 mm, or, if rather cylindric, longest outer phyllary to ca. 1/2 of inner ones.   58 Melanoseris
39 (34) Achene apex ± truncate (at least before shedding of pappus disk if pappus disk caducous)   (40)
+ Achene apex distinctly attenuate to beaked   (41)
40 (39) Longest outer phyllary ≤ 1/3 of inner ones; stem stiff erect; synflorescence narrowly racemiform or paniculiform; pappus caducous (without pappus disk).   75 Sonchella
+ Longest outer phyllary 1/2-3/4 of inner ones; stem weak and procumbent to ascending, or long creeping, or if stiff erect then hardened and densely divaricately and intricately branched; synflorescence corymbose to divaricately paniculiform, or capitulum solitary; pappus persistent or caducous together with pappus disk.   62 Launaea
41 (39) Stem leaves well developed and at least upper ones conspicuously clasping stem, or plant stoloniferous (Crepidiastrum spp. 1-6).   68 Crepidiastrum
+ Stem leaves absent or present but not clasping stem and plant not stoloniferous   (42)
42 (41) Stems herbaceous to hardened, several to numerous, more rarely only 1, erect, with some to many heads, from a stout woody caudex and plant often caespitose; basal leaves and stem leaves (if present) pinnate to bipinnate with linear, lanceolate, or filiform lobes; involucre 8-12 mm, most phyllaries subapically crested or corniculate (Crepidiastrum spp. 7-9).   68 Crepidiastrum
+ Stems usually 1 or 2, of various kind, either from an inconspicuous caudex and plant never caespitose, or plants annual; involucre ≤ 7 mm and phyllaries subapically all strictly plane, or involucre to 13 mm and either some phyllaries crested or corniculate but then leaves never pinnate with linear, lanceolate, or filiform lobes, or all strictly plane   (43)
43 (42) Involucre narrowly cylindric, 7-8 mm, all phyllaries subapically plane; perennial rosulate herbs, stem solitary; either stem, branches, and phyllaries white echinulate, or rosette leaves with a pentagonal or triangular lamina and a petiole as long as or longer than lamina (Ixeridium spp. 7 and 8).   73 Ixeridium
+ Involucre 4-13 mm, phyllaries all subapically plane or some crested or corniculate; annual herbs, or if perennial herbs and rosulate then stem, branches, and phyllaries never white echinulate and leaves never as above.   66 Youngia

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