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FOC | Family List | FOC Vol. 19 | Campanulaceae

10. Adenophora Fischer, Mém. Soc. Imp. Naturalistes Moscou. 6: 165. 1823.

沙参属 sha shen shu

Authors: Deyuan Hong, Thomas G. Lammers & Laura L. Klein

Herbs, perennial. Roots usually thickened, various in shape, carrot-shaped, mostly fleshy. Caudexes often short, sometimes elongated and branched. Stems erect, less frequently ascending. Basal leaves often rosulate, usually long petiolate, cordate; cauline leaves mostly alternate, rarely opposite or verticillate. Flowers in a cyme; cymes sometimes reduced to a branch with 1 terminal flower and 1 to several bracts, sometimes glomerate into a panicle. Calyx epigynous, rarely semi-epigynous, tube various in shape; lobes 5, entire or dentate. Corolla always epigynous, campanulate, funnelform, or tubular, 5-lobed or 5-fid to middle. Stamens 5; filaments much dilated into a lamella at base; lamellae densely ciliate-villous, valvate, enveloping disk; anthers elongate. Disk usually tubular, rarely annular. Ovary inferior, 3-locular; ovules numerous; style with pollen-receiving hairs; stigma 3-lobed, lobes narrow, recurved. Capsule 3-poricidal below persistent calyx lobes. Seeds numerous, ellipsoid, 1-angular.

Sixty-two species: E Asia, south to India and Vietnam, with one species extending into Europe and another endemic to Crimea; 38 species (23 endemic) in China.


1 Cauline leaves all or mostly verticillate   (2)
+ Cauline leaves all alternate   (5)
       
2 (1) Corolla small, less than 11 mm; calyx lobes filiform.   38 A. tetraphylla
+ Corolla larger than 13 mm; calyx lobes lanceolate or subulate   (3)
       
3 (2) Calyx lobes toothed; corolla tubular-campanulate, 18-25 mm; style slightly shorter than corolla.   33 A. wulingshanica
+ Calyx lobes entire; corolla campanulate, 13-20 mm; style exserted or included   (4)
       
4 (3) Cauline leaves partially verticillate; inflorescence branches not verticillate; calyx lobes lanceolate, 3-6 mm.   31 A. pereskiifolia
+ Cauline leaves all verticillate; inflorescence branches often verticillate; calyx lobes elliptic-lanceolate, 5-10 mm.   32 A. divaricata
       
5 (1) Cauline leaves (at least lower ones) petiolate, very rarely subsessile   (6)
+ Cauline leaves sessile, only some individuals of some species (e.g., A. stricta) with lower cauline leaves very shortly winged-petiolate   (17)
       
6 (5) Calyx lobes toothed   (7)
+ Calyx lobes entire, usually ovate to lanceolate, rarely linear-lanceolate, (1-)1.5-4 mm wide   (10)
       
7 (6) Cauline leaves pinnatifid up to over half; capsule columnar, base rounded, only ca. 3 mm wide.   34 A. lobophylla
+ Cauline leaves toothed; capsule ellipsoid, base obtuse, more than 3 mm wide   (8)
       
8 (7) Cauline leaves with petioles 2-4 cm, cordate at base; disk ca. 0.7 mm high; style equal to corolla in length.   7 A. cordifolia
+ Cauline leaves with petioles less than 2.5 cm, cuneate or shallowly cordate at base; disk 1.5-2.5 mm; style ± exserted   (9)
       
9 (8) Inflorescence a pseudoraceme or narrow panicle, without secondary branches; calyx lobes with verrucose teeth, or even entire; leaves glabrous adaxially.   8 A. liliifolia
+ Inflorescence a large panicle, with secondary branches; calyx lobes often with 1-4 pairs of long teeth, rarely verrucose teeth; leaves sparsely scaberulose adaxially.   20 A. potaninii
       
10 (6) Cauline leaves all obviously petiolate, base cordate or rounded, not decurrent or shortly decurrent; calyx lobes obtuse at apex   (11)
+ Cauline leaves only lower ones petiolate, upper ones sessile or shortly cuneate-petiolate; calyx lobes acute or acuminate at apex   (12)
       
11 (10) Cauline leaves all cordate, papery; calyx tube (hypanthium) obconic at least at anthesis.   1 A. trachelioides
+ Cauline leaves rounded or broadly cuneate, only some lower ones shallowly cordate, membranous; calyx tube obovoid or obovoid-conical.   2 A. remotiflora
       
12 (10) Cauline leaves only 2, lower one cordate, upper one much smaller.   5 A. brevidiscifera
+ Cauline leaves many, not cordate   (13)
       
13 (12) Calyx tube globose, calyx lobes subulate-lanceolate, 8-14 mm; pedicels 1.5-3 cm.   6 A. longipedicellata
+ Calyx tube obovoid or obconical, calyx lobes usually shorter; pedicels shorter   (14)
       
14 (13) Corolla 1.5-2.7 cm, lobes 5-11 mm; calyx lobes ovate to linear-lanceolate, 1.5-4 mm wide.   3 A. petiolata
+ Corolla less than 1.8 cm, lobes less than 5 mm; calyx lobes linear-lanceolate or subulate, 1-2 mm wide   (15)
       
15 (14) Leaves usually sparsely hispidulous, 7-13 × 1.5-3 cm; calyx mostly hairy, occasionally glabrous, lobes 1-2 mm wide; corolla ca. 17 mm.   4 A. rupincola
+ Leaves glabrous or hairy along veins, 3-8 cm; calyx glabrous, occasionally scaberulose; corolla 13-15 mm   (16)
       
16 (15) Leaves often hairy on margins and along veins; calyx tube obovoid or obconic; inflorescence a narrow panicle or pseudoraceme.   8 A. liliifolia
+ Leaves glabrous throughout; calyx tube usually globose, rarely obovoid-globose; inflorescence a panicle.   11 A. sinensis
       
17 (5) Corolla small, less than 17 mm, constricted at throat and thus corolla urceolate; style strongly exserted, more than 1.5 × as long as corolla; disk narrowly tubular, 2-7 mm, less than 1 mm wide   (18)
+ Corolla usually larger, campanulate; style slightly exserted or included, less than 1.5 × as long as corolla; disk not much longer than broad   (20)
       
18 (17) Calyx lobes filiform, 3-20 mm, sometimes with 1 or 2 pairs of verrucose teeth at lower part.   37 A. capillaris
+ Calyx lobes subulate, entire   (19)
       
19 (18) Stems and leaves densely scaberulose; corolla 10-17 mm.   35 A. stenanthina
+ Stems and leaves glabrous or villous; corolla 8-12 mm.   36 A. liliifolioides
       
20 (17) Calyx lobes entire   (21)
+ Calyx lobes toothed   (32)
       
21 (20) Calyx scaberulose; calyx lobes subulate, broadest at base, 6-8 mm; style nearly as long as corolla.   10 A. stricta
+ Calyx mostly glabrous, if hairy then corolla ± constricted at throat; calyx lobes triangular-lanceolate to linear-lanceolate, 2-6 mm, if more than 6 mm then calyx tube never globose and glabrous; style obviously exserted or not   (22)
       
22 (21) Calyx lobes very short, shorter than 2.5 mm   (23)
+ Calyx lobes longer than 3 mm, broadest at middle   (24)
       
23 (22) Cauline leaf blade linear, ca. 5 cm × 0.8 mm; corolla 18-19 mm.   14 A. pinifolia
+ Cauline leaf blade linear to oblong, 1.5-4 × 0.3-1 cm; corolla 12-14 mm.   15 A. micrantha
       
24 (22) Calyx variously hairy; corolla shallowly lobed, ± constricted at throat, lobes less than 1/4 as long as total length, often reflexed   (25)
+ Calyx glabrous; corolla more deeply lobed, not constricted at throat, lobes more than 1/4 as long as total length   (26)
       
25 (24) Corolla throat slightly constricted, 9-15 mm wide; corolla lobes 4-6 mm.   12 A. polyantha
+ Corolla throat strongly constricted, 4-7 mm wide; corolla lobes up to 3 mm.   13 A. contracta
       
26 (24) Calyx lobes elliptic, ca. 2 mm wide, middle part broadest, nearly 2 × as broad as base.   16 A. palustris
+ Calyx lobes linear-lanceolate or subulate, base broadest or middle part broadest but not 2 × as broad as base   (27)
       
27 (26) Calyx lobes subulate, base broadest   (28)
+ Calyx lobes linear-lanceolate, middle part broadest, or at least broadest part not at base   (30)
       
28 (27) Disk broadly tubular, 3-8 mm, up to 3 mm wide; calyx lobes 5-10 mm; inflorescence a pseudoraceme.   17 A. himalayana
+ Disk less than 1.5 mm; calyx lobes 3-5 mm; inflorescence a panicle, rarely a pseudoraceme   (29)
       
29 (28) Cauline leaves ovate to lanceolate, 5-25 mm wide; corolla 12-21 mm.   18 A. amurica
+ Cauline leaves mostly linear to linear-elliptic, less than 5 mm wide; corolla 10-13 mm.   23 A. stenophylla
       
30 (27) Corolla 16-28 mm, lobes 6-8 mm; leaves entire or sparsely serrate, 2-13 mm wide; style usually shorter than or nearly as long as corolla.   19 A. gmelinii
+ Corolla 13-22 mm, lobes 4-6 mm; leaves always crenulate, 5-40 mm wide; calyx lobes 3-6 mm; style exserted or not   (31)
       
31 (30) Style included or as long as corolla; leaves 2.5-7 cm; corolla lobes 4-6 mm.   9 A. lamarckii
+ Style exserted; leaves 6-13 cm; corolla lobes ca. 4 mm.   31 A. pereskiifolia
       
32 (20) Calyx lobes ovate-triangular, overlapping at lower part, recurved toward lateral side; capsule subglobose; style included.   21 A. tricuspidata
+ Calyx lobes narrower, never overlapping, not recurved; capsule subglobose or not; style exserted or included   (33)
       
33 (32) Cauline leaves aggregated at middle part of stems, glabrous, lanceolate or linear-elliptic.   22 A. wilsonii
+ Cauline leaves evenly distributed or aggregated at lower part of stems, glabrous or hairy   (34)
       
34 (33) Cauline leaves linear or linear-elliptic, less than 5 mm wide; calyx lobes subulate, 3-4 mm; corolla 10-13 mm.   23 A. stenophylla
+ Cauline leaves wider; calyx lobes mostly larger; corolla mostly longer   (35)
       
35 (34) Flowers solitary and terminal, or several in a pseudoraceme, very occasionally in a narrow panicle; corolla 2-3.4 cm, narrowly campanulate.   24 A. elata
+ Flowers several to numerous, in a pseudoraceme or panicle; corolla rarely (except in A. morrisonensis) more than 2.4 cm, campanulate or broadly campanulate   (36)
       
36 (35) Leaves mostly glabrous; stems mostly glabrous, or puberulent   (37)
+ Leaves ± hairy; stems usually hairy   (40)
       
37 (36) Calyx lobes narrowly subulate, ca. 0.5 mm wide; style exserted 4-7 mm above corolla.   28 A. hubeiensis
+ Calyx lobes 1-2 mm wide; style included, or slightly exserted, less than 4 mm above corolla   (38)
       
38 (37) Corolla 28-35 mm; disk less than 1 mm.   29 A. morrisonensis
+ Corolla much shorter; disk longer   (39)
       
39 (38) Capsule ellipsoid or oblong, more than 3 mm in diam.; calyx lobes 5-7 mm.   8 A. liliifolia
+ Capsule narrowly ellipsoid, ca. 3 mm in diam.; calyx lobes 2-4(-6) mm.   30 A. ningxianica
       
40 (36) Leaves mostly aggregated at lower half of stems, upper ones absent or much smaller; calyx lobes 5-20 mm, obviously toothed   (41)
+ Leaves evenly distributed; calyx lobes 3-7 mm   (42)
       
41 (40) Calyx lobes 5-8(-10) mm, often gray; corolla lobed to middle or near middle; disk 0.5-1 mm.   26 A. jasionifolia
+ Calyx lobes 8-20 mm, green; corolla lobed to 1/3; disk 1.2-3.5 mm.   27 A. coelestis
       
42 (40) Cauline leaves with 2 to several large teeth on each side, puberulent on both surfaces; disk 1.5-3 mm.   20 A. potaninii
+ Cauline leaves with numerous teeth, adaxially sparsely scaberulose, hispidulous, or glabrous, abaxially hirsute or hispidulous; disk less than 1.4 mm   (43)
       
43 (42) Cauline leaves narrowly elliptic or lanceolate, 0.5-2.5 cm; style mostly as long as corolla.   18 A. amurica
+ Cauline leaves ovate, 1.5-6.5 cm; style ± exserted.   25 A. khasiana

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