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23. Pleurospermum Hoffmann, Gen. Pl. Umbell. viii. 1814.

棱子芹属 leng zi qin shu

Pan Zehui (潘泽惠); Mark F. Watson

Pleurospermum amabile

Credit: Harvard University Herbaria

Aulacospermum Ledebour; Hymenidium de Candolle; Hymenolaena de Candolle; Pterocyclus Klotzsch.

Herbs perennial, rarely biennial. Root crown often surrounded with fibrous remnant sheaths. Stems erect, sometimes shortened. Leaves 1–4-pinnate or ternate-pinnate; ultimate segments serrate to incised or pinnate. Umbels terminal and lateral; bracts several, entire or pinnate, margin usually white scarious; rays often extending in fruit; bracteoles numerous, scarious, sometimes white margined. Calyx teeth conspicuous or obsolete. Petals oblong to broad-ovate, white or purple-red, base clawed, apex narrow, inflexed. Stylopodium conic or short-conic. Fruit oblong to broad-ovoid, slightly flattened laterally, glabrous, often with numerous, shining tubercules; ribs prominent and acute, sometimes undulate, cristate or narrowly winged; vittae 1(–3) in each furrow, 2(or 4 or 6) on commissure. Seed face concave. Carpophore 2-parted.

About 50 species: N Asia, E Europe, and especially diverse in the Himalayan region and W China; 39 species (22 endemic) in China.

This is a widespread, heterogeneous genus of complex and controversial taxonomy. Russian authors delimit Pleurospermum sensu stricto by only two species (the type, P. austriacum Linnaeus, and P. uralense), referring the other species to Aulacospermum, Hymenidium, Hymenolaena, Physospermopsis, and Pterocyclus. Other morphologically similar genera, where generic boundaries become indistinct, include Trachydium and Pseudotrachydium (Kljuykov et al.) Pimenov & Kljuykov. A full discussion of the Russian classification of Pleurospermum is presented by Pimenov and Kljuykov (Feddes Repert. 111: 499–515, 517–534, 535–552. 2000). As yet, this rather radical classification has not gained widespread acceptance, and the proponents admit that this is a taxonomic hypothesis, and (particularly for some groups) a more natural classification will only be possible following critical revision in the field and herbarium. A traditional treatment is adopted for the following account, with due influence from the work of the Russian taxonomists.

The following species have been described from Chinese material, but are imperfectly known as no specimens have been seen or the specimens are inadequate.

Hymenidium pachycaule Pimenov & Kljuykov (Edinburgh J. Bot. 53: 275. 1996), described from NW Gansu (“Lienhuo Shan,” J. F. C. Rock 12703, holotype, E; isotype, NAS). It is possibly referable to Pleurospermum, but further research is required.
Pleurospermum albimarginatum H. Wolff (Repert. Spec. Nov. Regni Veg. 21: 243. 1925), described from W Sichuan (“Tongolo,” J. A. Soulié 2618, holotype, P).
Pleurospermum grandifolium H. Wolff (Repert. Spec. Nov. Regni Veg. 21: 244. 1925), described from W Sichuan (“Tatsienlu” [Kangding], J. A. Soulié 2182, holotype, P). It has possible affinities with P. benthamii.
Pleurospermum microphyllum H. Wolff (Repert. Spec. Nov. Regni Veg. 21: 242. 1925), described from W Sichuan (“Tongolo,” J. A. Soulié 2605, holotype, P).
Pleurospermum microsciadium H. Wolff (Repert. Spec. Nov. Regni Veg. 21: 241. 1925), described from W Sichuan (“Tongolo,” J. A. Soulié 2614, holotype, P).
Pleurospermum souliei H. Wolff (Repert. Spec. Nov. Regni Veg. 19: 309. 1924), described from W Sichuan (Kangding, J. A. Soulié 2196, holotype, P).

1 Plants of high altitudes, (3500–)4000–5000 m, short and squat 5–20(–40) cm; stems usually much reduced, often thickened, unbranched.   (2)
+ Plants from varying altitudes and habitats, usually much larger (0.3–2 m), smaller specimens slender-stemmed.   (10)
2 (1) Bracteole apex acute, acuminate or obtuse.   (3)
+ Bracteole apex 1–2-pinnate/pinnatifid.   (5)
3 (2) Fruit wings narrow, crisped; calyx teeth lanceolate; rays thickened, particularly at the distal ends; pedicels flattened and winged.   1 Pleurospermum nubigenum
+ Fruit wings narrow or broad, plane; calyx teeth triangular or obsolete; rays uniform thickness, pedicels terete.   (4)
4 (3) Petals white; fruit thinly winged; calyx teeth triangular; vittae 3 per furrow.   2 Pleurospermum pulszkyi
+ Petals purple-red; fruit broadly winged; calyx teeth obsolete; vittae 2 per furrow.   3 Pleurospermum lindleyanum
5 (2) Bracteole apex 2-pinnate/pinnatifid; ultimate leaf segments linear, 0.3–0.5 mm broad; pedicels flattened.   4 Pleurospermum heterosciadium
+ Bracteole apex 1-pinnate/pinnatifid; ultimate leaf segments lanceolate, 0.5–5 mm broad; pedicels terete.   (6)
6 (5) Compact rosette, stemless; inflorescence densely capitate, subsessile; rays reduced, crowded; fruit wings broad and sinuate, not dentate.   5 Pleurospermum hedinii
+ Stems usually distinct; inflorescences not densely capitate, usually distinctly pedunculate; rays not reduced, lax; fruit ribs prominent or irregularly dentate or sinuate winged.   (7)
7 (6) Umbels usually subsessile; rays very unequal; fruit tuberculate, wings irregularly dentate.   8 Pleurospermum astrantioideum
+ Umbels distinctly pedunculate; rays subequal; if fruit winged then not tuberculate.   (8)
8 (7) Plants with strong unpleasant odor; rays 14–30; fruit wings irregularly dentate.   9 Pleurospermum foetens
+ Plant without unpleasant odor; rays 5–15(–20); fruit ribs prominent to narrowly winged, wing margin entire.   (9)
9 (8) Leaves 1–2-pinnate, ultimate segments ovate or orbicular; fruit ribs all broadly winged.   6 Pleurospermum stellatum
+ Leaves 2–3-pinnate, ultimate segments linear-lanceolate; fruit ribs prominent or narrowly sinuate- winged.   7 Pleurospermum nanum
10 (1) Large, often robust plants, (0.5–)0.8–2 m, rarely shorter.   (11)
+ Small, slender plants, 20–40(–50) cm rarely taller.   (23)
11 (10) Leaflets of lower leaves broadly ovate, regularly serrate, base truncate or cordate.   (12)
+ Leaflets of lower leaves variously lobed and dissected, often pinnatifid, not broadly ovate, bases usually cuneate to decurrent.   (13)
12 (11) Umbels 8–10 cm across; rays 15–20, 4–7 cm; fruit 8–10 × 4–6 mm.   10 Pleurospermum rivulorum
+ Umbels 2–4 cm across; rays 6–8(–15), 0.6–2 cm; fruit 7–8 × 3–4 mm.   11 Pleurospermum rotundatum
13 (11) Plants usually very robust, thick-stemmed (except P. stylosum); umbels (7–)12–28 cm across; rays (15–)25–40.   (14)
+ Plants less robust, slender-stemmed; umbels 3–15 cm across; rays 7–28.   (16)
14 (13) Rays 15–35; fruit 4–6 mm; ribs sinuate-winged (Xinjiang).   12 Pleurospermum stylosum
+ Rays 20–40(–60); fruit 6–10 mm; ribs cristate- or plane-winged.   (15)
15 (14) Ultimate leaf segments pinnatifid, lobes linear; calyx teeth conspicuous, triangular-ovate; fruit wings thick, cristate (NW China).   13 Pleurospermum uralense
+ Ultimate leaf segments coarsely dentate, lobes ovate; calyx teeth obsolete; fruit wings thin, broad and plane (SW China).   14 Pleurospermum aromaticum
16 (13) Petioles of upper leaves expanded into auriculate sheaths; fruit oblong, 8–15 mm.   (17)
+ Petioles of upper leaves expanded or not, but not auriculate; fruit ovate to oblong-ovate, 3.5–8(–10) mm.   (18)
17 (16) Leaves irregularly serrate; bracteoles ca. 5 mm broad; seed face deeply concave.   15 Pleurospermum angelicoides
+ Leaves regularly crenate; bracteoles ca. 1 mm broad; seed face slightly concave.   16 Pleurospermum longicarpum
18 (16) Leaves 2–4-ternate/pinnate.   (19)
+ Leaves 1–2-ternate/pinnate (3–5-partite to 2-ternate).   (20)
19 (18) Leaves with narrow ultimate segments 2–5 mm broad, hirtellous along the margins, lobes round, obtuse; primary umbel overtopped by laterals; bracteoles oblong to obovate, entire to 3-lobed at apex, lobes obtuse; petals white; fruit 4–5 mm.   19 Pleurospermum franchetianum
+ Leaves with ultimate segments 0.3–1 mm broad, margin glabrous, lobes lacerate, acute; primary umbel not overtopped by laterals; bracteoles oblanceolate, 3-lobed to pinnate/pinnatifid at apex, apex acute; petals white to pink; fruit 6–10 mm.   20 Pleurospermum benthamii
20 (18) Leaves 3–5-partite (sometimes to 2-ternate).   (21)
+ Leaves 2-ternate/pinnate.   (22)
21 (20) Leaves 3–5-partite (sometimes to 2-ternate); leaf bases and veins with coarse white hairs reminiscent of those of Heracleum; rays 2–4 cm; petals white.   17 Pleurospermum heracleifolium
+ Leaves always 3-partite, without white hairs; rays 5–6 cm; petals pink.   18 Pleurospermum macrochlaenum
22 (20) Rays 10–18, 2–7 cm; bracteoles lanceolate, acute or 3-lobed; fruit smooth, wings sinuate but not irregularly dentate.   21 Pleurospermum decurrens
+ Rays 7–13. 1–2.5 cm; bracteoles linear-lanceolate, entire; fruits tuberculate with irregularly dentate wings.   22 Pleurospermum cristatum
23 (10) Petiole wings greatly expanded, white with purple veining, upper leaves and bracts enveloping the inflorescence.   23 Pleurospermum amabile
+ Petiole wings not greatly expanded, not purple veined.   (24)
24 (23) Leaves 1-pinnate to 2-pinnatifid; rays 2–4; bracteoles broadly ovate, entire.   24 Pleurospermum bicolor
+ Leaves 2–4-ternate/pinnate; rays (4–)6–25; bracteoles entire or more usually pinnatifid at apex.   (25)
25 (24) Stems, petioles and leaf rachis conspicuously white pubescent.   26 Pleurospermum pilosum
+ Plants essentially glabrous, bases of umbels and/or petioles sometimes puberulent.   (26)
26 (25) Calyx teeth conspicuous, triangular to lanceolate.   (27)
+ Calyx teeth obsolete or minute.   (34)
27 (26) Ultimate leaf segments 5–20 mm broad, irregularly serrate or incised.   25 Pleurospermum yunnanense
+ Ultimate leaf segments finely dissected, 1–8(–10) mm broad.   (28)
28 (27) Calyx teeth linear-lanceolate; sheaths of upper leaves and bracts broadly auriculate; bracteoles silvery white with a narrow green midrib, apex acuminate to 3-lobed.   27 Pleurospermum album
+ Calyx teeth triangular or rounded; leaf sheaths somewhat expanded but not auriculate; bracteoles usually white margined, but not conspicuously silvery white, apex usually pinnatifid.   (29)
29 (28) Rays 4–7, very unequal; bracteoles entire, apex acuminate.   29 Pleurospermum rupestre
+ Rays (6–)8–20(–30), subequal (but see P. szechenyii); bracteole apex pinnatifid (rarely acuminate).   (30)
30 (29) Rays (15–)20–30.   (31)
+ Rays 6–12(–15).   (32)
31 (30) Plant without unpleasant odor when crushed; rays very unequal; fruit wings broad, sinuate but not dentate.   30 Pleurospermum szechenyii
+ Plant with a strong unpleasant odor when crushed; rays subequal; fruit wings broad, irregularly dentate.   9 Pleurospermum foetens
32 (30) Leaf blades oblong-lanceolate in outline, approaching 2–3-ternate/pinnate; fruit wings broad, sinuate.   31 Pleurospermum wilsonii
+ Leaf blades broadly ovate in outline, 3–4-ternate/pinnate; fruit ribs prominent to winged, but not sinuate.   (33)
33 (32) Ultimate leaf segments ca. 2 mm; calyx teeth ca. 1 mm, green.   32 Pleurospermum hookeri
+ Ultimate leaf segments 3–5 mm; calyx teeth ca. 0.3 mm, dark purple.   33 Pleurospermum tsekuense
34 (26) Rays 5–9(–11); bracts and bracteoles entire, acuminate.   (35)
+ Rays (9–)11–25; bracts and bracteoles pinnate/pinnatifid at apex, rarely acuminate.   (36)
35 (34) Stems and leaf sheaths purple-red, rays subequal; bracteoles broad, longer than the flowers.   35 Pleurospermum apiolens
+ Stems and leaf sheaths green, rays very unequal; bracteoles linear-lanceolate, shorter than the pedicels.   28 Pleurospermum simplex
36 (34) Larger plants, 30–60 cm; rays 10–25; fruit wings broad, sinuate or irregularly cristate/dentate.   (37)
+ Small slender plants, 20–35(–45) cm; rays 9–15; fruit ribs prominent to narrowly winged, but not sinuate or dentate.   (38)
37 (36) Rays subequal; fruit wings sinuate.   37 Pleurospermum linearilobum
+ Rays very unequal; fruit wings irregularly cristate/dentate.   39 Pleurospermum wrightianum
38 (36) Rays 1.5–2.5 cm.   34 Pleurospermum giraldii
+ Rays 4–12 cm.   (39)
39 (38) Stem branches often opposite or whorled; leaf rachis glabrous; ovary smooth.   36 Pleurospermum handelii
+ Stem branches usually alternate; leaf rachis tuberculate on abaxial surface; ovary tuberculate.   38 Pleurospermum calcareum

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