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87. Vernonia Schreber, Gen. Pl. 2: 541. 1791.

斑鸠菊属 ban jiu ju shu

Vernonia esculenta

Credit: Harvard University Herbaria

Herbs, shrubs, or trees, sometimes climbing; indumentum of simple or T-shaped hairs and sessile globose glands. Leaves alternate, rarely opposite, petiolate or subsessile, often ill-defined, margin entire, serrate, or dentate, pinnately veined. Synflorescence terminal or axillary, paniculate, densely clustered to laxly corymbose, sometimes thyrsoid, racemelike, or reduced to solitary capitulum. Involucre bowl-shaped to narrowly cylindric; phyllaries imbricate, to 6-seriate, often with scarious margins, apex obtuse, acute, or acuminate into spinescent or threadlike tip, outer phyllaries shorter, inner occasionally caducous. Receptacle flat, areolate, naked or sometimes shortly hairy. Florets (1-)3-100+; corolla pink, purplish, rarely white, narrowly tubular, often glandular, with campanulate or campanulate-funnelform limb; lobes 5. Anthers hastate, obtusely auricled. Style branches subulate, puberulent. Achenes cylindric or obconical, mostly with (7-)10 well-defined ribs, less often 4- or 5-ribbed or angled, rarely terete, usually glandular between ribs, glabrous or shortly hairy, apex truncate, often callose at base. Pappus usually double, outer pappus of short bristles or narrow scales, occasionally absent, inner pappus of many scabrid-barbellate-plumose bristles, deciduous or persistent, often colored.

About 1,000 species in the wide sense: tropical Asia and Africa, North and South America; 31 species (eight endemic, one introduced) in China.

Robinson (Proc. Biol. Soc. Washington 112: 220-247. 1999; in Kadereit & C. Jeffrey, Fam. Gen. Vasc. Pl. 8: 149-175. 2007) has proposed the segregation of many smaller genera from Vernonia, with Vernonia s.s. restricted to ca. 17 species in North America. Most of his segregates can be rather difficult to delimit, and the placement of individual species is not always easy; so Vernonia continues to be used mostly in its old, very wide, sense. However, there are some segregates that are quite widely accepted. One of these, Distephanus, occurs in China and is recognized in this Flora. The Chinese species are divided below into informal groups corresponding to these potential segregate genera.

Vernonia andersonii C. B. Clarke (Compos. Ind. 27. 1876; Decaneuropsis andersonii (C. B. Clarke) H. Robinson & Skvarla) is included in the following key on the basis of Robinson and Skvarla (Taiwania 55: 262. 2010). All the material so named seen so far by the present authors has proved to be V. cumingiana.

1 Trees, shrubs, or climbers; florets (1 or)2-12 per involucre, rarely more and then plant a climber with axillary synflorescences   (2)
+ Erect annual or perennial herbs; florets 19 or more per involucre; synflorescences terminal   (16)
2 (1) Trees or self-supporting shrubs   (3)
+ Scandent shrubs   (9)
3 (2) Leaves obovate to oblanceolate, clearly widest above middle, basal secondary veins substraight toward leaf margins; petiole short and broad or expanded into a sheath at base; synflorescence conically thyrsoid (3. Monosis group)   (4)
+ Leaves ovate to elliptic or oblong, widest at or below middle, basal secondary veins curved and often becoming subparallel to basal leaf margins; petiole slender; synflorescence flat-topped to rounded   (5)
4 (3) Trees 3-8 m tall; leaf blade to 50 × 20 cm, abaxially glabrous or sparsely pubescent, lateral veins 12-17-paired; achenes 3-4 mm.   8 V. volkameriifolia
+ Shrubs 2-3 m tall; leaf blade 15-30 × 4-9 cm, abaxially densely pubescent to fulvous tomentose, lateral veins 7-12-paired; body of mature achenes ca. 2.5 mm.   9 V. parishii
5 (3) Corolla 8-10 mm, uniformly slender from base before widening into limb; achenes strongly 10-ribbed; style base without node; sweeping hairs of style branches pointed (4. Gymnanthemum group).   10 V. extensa
+ Corollas 5-7 mm, widening gradually from near base; achenes 5-ribbed to weakly 10-ribbed; style base with large distinct node; sweeping hairs of style branches blunt (1. Strobocalyx group p.p.)   (6)
6 (5) Trees; branches fulvous or brown tomentose; inner phyllaries persistent   (7)
+ Shrubs, rarely small trees; branches gray or gray-brown tomentose; innermost phyllaries ± deciduous   (8)
7 (6) Florets 5 or 6 per capitulum; involucres 6-8 mm in diam.; phyllaries ovate-oblong, shortly pubescent or subglabrous; achenes subtriangular in section, 8-10-ribbed, glabrous, glandular; leaf blade thinly leathery, glabrous or sparsely puberulent along veins, margin entire.   1 V. arborea
+ Florets ca. 10 per capitulum; involucres 4-5 mm in diam.; phyllaries ovate, abaxially densely tomentose; achenes 4- or 5-angled, glabrous; leaf blade papery, abaxially hispidulous, adaxially tomentose, margin repand or laxly dentate.   2 V. solanifolia
8 (6) Capitula 6-8 mm in diam., florets 8-12; involucres subglobose or hemispheric with rounded base, 4-6 mm in diam.; phyllaries obtuse or rounded at apex, white sericeous; achenes 3.5-4 mm, 3-ribbed ventrally, glabrous; leaves membranous or thinly papery.   3 V. bockiana
+ Capitula 2-3 mm in diam., florets 5 or 6; involucres obconical with cuneate base, 2-3 mm in diam.; phyllaries few, acute or slightly acute at apex, sparsely shortly pubescent; achenes ca. 3 mm, slightly ribbed, sparsely puberulent, glandular; leaves hardly papery.   4 V. esculenta
9 (2) Synflorescences terminal and subterminal, forming broad panicle; pappus white or dirty white; branches densely reddish- or yellowish brown pubescent (1. Strobocalyx group p.p.)   (10)
+ Synflorescences mostly axillary, forming long thyrse; pappus reddish, yellow, or brown; branches grayish brown pubescent   (11)
10 (9) Branches densely reddish brown pubescent, eglandular; lateral veins 7- or 8-paired; capitula 7-12 mm in diam., with ca. 10 florets, peduncle up to 15 mm; achenes 10-ribbed, eglandular.   5 V. sylvatica
+ Branches densely yellowish brown pubescent, glandular; lateral veins 3-6-paired; capitula 3-5 mm in diam., with 2 or 3 florets, peduncle less than 2 mm; achenes 5-ribbed, glandular.   6 V. chunii
11 (9) Hairs of stems and leaves T-shaped with long arms, shortly stalked and pressed against stems or leaf surface, silky; corollas without long slender bases; style with distinct basal node; achenes 5-angled (2. Tarlmounia group).   7 V. elliptica
+ Hairs of stems and leaves simple, not T-shaped; corollas with long and filiform bases; bases of styles without nodes; achenes strongly 10-ribbed (5. Decaneuropsis group)   (12)
12 (11) Capitula in terminal or subterminal panicles; receptacle ferruginous pubescent; pappus reddish or reddish brown; leaves papery; branches reddish brown or grayish brown tomentose.   11 V. cumingiana
+ Capitula mostly axillary or on short lateral branches, solitary or in groups of 3-5, forming leafy thyrse; receptacle glabrous; pappus yellowish brown; leaves leathery or thinly leathery; branches sparsely, shortly pubescent or glabrescent   (13)
13 (12) Leaf blade oblong or oblong-lanceolate, lateral veins 7-9-paired, adaxially densely gray-green tomentose; capitula with ca. 10 florets; achenes 3-3.5 mm   (14)
+ Leaf blade ovate-oblong or oblong-elliptic, lateral veins 5-7-paired, adaxially sparsely pubescent; capitula with 18 or more florets; achenes 4-4.5 mm   (15)
14 (13) Phyllaries linear or linear-lanceolate, acute at apex, with dense short pubescence on margins and tips.   12 V. gratiosa
+ Phyllaries oblong, with short acute tips, glabrous.  

V. andersonii (see note under V. cumingiana)

15 (13) Capitula 7-8 mm in diam., 3-5 in small axillary panicles; florets 20-25, reddish purple; achenes 3-3.5 mm, 10-ribbed, with short patent hairs and glands; leaves thinly leathery, elliptic or ovate-elliptic, entire.   13 V. blanda
+ Capitula ca. 20 mm in diam., solitary in leaf axils, racemose; florets many, white; achenes 7-9 mm, 7- or 8-ribbed, glabrous; leaves leathery, obovate-oblong or elliptic-oblong, margin often revolute, sparsely denticulate at upper margins.   14 V. chingiana
16 (1) Achenes 4- or 5-angled to almost terete; phyllary apex acute or aristate-acuminate; plants often annual herbs (7. Cyanthillium group)   (17)
+ Achenes (7-)10-ribbed or striate   (20)
17 (16) Achenes unribbed or rarely inconspicuously ribbed, ± compressed, densely white pilose; pappus 2-seriate, outer setae many, short, and persistent; capitula small, ca. 6 mm in diam., many in relatively broad panicles; florets 19-28   (18)
+ Achenes 4- or 5-angled, glabrous and glandular; pappus 1-seriate, deciduous; capitula larger, 6-10 mm in diam., few or 2 or 3 terminal on branches; florets more than 30   (19)
18 (17) Corolla ca. as long as pappus; leaf blade rhombic to ovate.   26 V. cinerea
+ Corolla much longer than pappus at anthesis; leaf blade linear-lanceolate to lanceolate.   27 V. benguetensis
19 (17) Capitula several, laxly corymbose, 5-7 mm in diam.; stem leaves crowded, spatulate, apex rounded or obtuse, margin subentire, abaxially densely white hispidulous; small subshrubs, ca. 10 cm tall.   28 V. maritima
+ Capitula usually 2 or 3 terminal on branches or paired, 8-10 mm in diam.; stem leaves widely separated, ovate, ovate-elliptic, rarely orbicular, apex obtuse, slightly acute, margin crenate-shallowly dentate, abaxially sparsely shortly hairy or glabrous, adaxially gray sericeous-pubescent; robust annual herbs, to 1 m tall.   29 V. patula
20 (16) Capitula subtended by foliose bracts as long as phyllaries; phyllary apex ± leaflike; corolla with slender base more than 2 × as long as abruptly wider narrowly campanulate limb; pappus bristles flattened on outer surface (9. Baccharoides group).   31 V. anthelmintica
+ Capitula without foliose or membranous appendaged bracts at base of involucre; phyllary apex not leaflike; corolla with slender base up to as long as narrowly funnel-shaped limb; pappus segments rounded and/or scabrid on outer surface   (21)
21 (20) Lower part of leaf blade with mostly spreading secondary veins; slender base of corolla tube ca. as long as limb; base of style without node (8. Khasianthus group).   30 V. subsessilis
+ Lower part of leaf blade with secondary veins ascending at 45° angles or more; slender base of corolla tube shorter than limb; base of style with distinct node (6. Acilepis group)   (22)
22 (21) Capitula ca. 30 mm in diam.; phyllaries abaxially pale yellowish white tomentose.   25 V. albosquama
+ Capitula 5-25 mm in diam.; phyllaries glabrous to densely pubescent, green, often with darker apex   (23)
23 (22) Outer pappus bristles many, short   (24)
+ Outer pappus bristles few to none   (27)
24 (23) Capitula usually more than 5 in terminal, often somewhat elongated, panicle or thyrse, 5-8 mm in diam.; achenes glabrous; pappus white or dirty white.   15 V. saligna
+ Capitula few, usually 2 or 3(-5) in ± flat-topped cyme, rarely solitary in leaf axils, 8-15 mm in diam.; achenes hairy and glandular; pappus sordid white or rubescent, outer bristles deciduous or persistent   (25)
25 (24) Stems hispidulous; capitula 10-15 mm in diam., peduncle very short to almost absent; florets ca. 30; phyllary apex apiculate; leaves oblanceolate or obovate-elliptic, sharply serrate, hispidulous on both surfaces.   16 V. aspera
+ Stems shortly pubescent or adpressed hairy; capitula 6-8(-10) mm in diam., peduncle 7-15 mm; florets 15-20; phyllary apex acute or acuminate; leaves oblong, oblong-lanceolate, obovate, or obovate-elliptic, denticulate or sparsely serrate toward apex, subglabrous or shortly pubescent along veins only   (26)
26 (25) Involucres hemispheric; phyllaries with apex acute, upper part arachnoid pubescent and glandular; achenes hispidulous; leaves papery, laxly on stems, oblong or oblong-lanceolate, sparsely serrate on upper margins.   17 V. clivorum
+ Involucres obconical; phyllaries with apex acuminate, subscarious, margin pubescent; achenes patently villous; leaves leathery, often conferted below flowering branches, obovate or obovate-elliptic, upper leaves linear or linear-lanceolate, denticulate.   18 V. attenuata
27 (23) Capitula 4-6 mm in diam.; florets 8-12; shrubs or shrubby herbs 1-3 m tall; phyllary apex obtuse or bluntly acute, adpressed to form smooth involucre.   19 V. divergens
+ Capitula 12-25 mm in diam.; florets many; herbs 15-80(-200) cm tall; phyllary apex acute to acuminate, often conspicuous or colored spinescent, often recurved   (28)
28 (27) Capitula sessile or nearly so, solitary, often axillary and forming spikelike synflorescence; outermost phyllaries almost subulate.   24 V. squarrosa
+ Capitula mostly clearly pedunculate, often in flat-topped cymes, never axillary; outer phyllaries lanceolate to ovate   (29)
29 (28) Achenes glabrous; [inner phyllaries ca. 2.5 mm wide; stem hairs pigmented, clearly multicellular]   (30)
+ Achenes adpressed hairy; inner phyllaries 1.5-2.2 mm wide; stem hairs pale, slender, obscurely multicellular   (31)
30 (29) Panicles relatively broad, ± flat-topped; phyllaries often recurved at tip.   21 V. silhetensis
+ Panicles relatively narrow, ± cylindric; phyllaries straight to slightly incurved.   22 V. doichangensis
31 (29) Involucre 12-15 mm wide; leaf blade thinly papery; achenes eglandular.   20 V. nantcianensis
+ Involucre 15-20 mm wide; leaf blade more stiffly papery; achenes glandular.   23 V. spirei

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