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FOC | Family List | FOC Vol. 9 | Rosaceae

10. COTONEASTER Medikus, Philos. Bot. 1: 154. 1789.

栒子属 xun zi shu

Lu Lingdi (Lu Ling-ti); Anthony R. Brach

Cotoneaster multiflorus

Credit: Harvard University Herbaria

Shrubs, rarely small trees, erect, decumbent, or prostrate, deciduous, semievergreen, or evergreen. Branchlets mostly terete, rarely slightly angulate, unarmed. Winter buds small; scales several, imbricate, exposed. Leaves alternate, simple, shortly petiolate; stipules caducous, usually subulate, small; margin of leaf blade entire, venation camptodromous. Inflorescences terminal or axillary, cymose or corymbose, sometimes flowers several fascicled or solitary. Hypanthium turbinate or campanulate, rarely cylindric, adnate to ovary. Sepals 5, persistent, short. Petals 5, erect or spreading, imbricate in bud, white, pink, or red. Stamens 10–20(–22), inserted in mouth of hypanthium. Ovary inferior or semi-inferior, 2–5-loculed; carpels 2–5, connate abaxially, free adaxially; ovules 2 per carpel, erect; styles 2–5, free; stigmas dilated. Fruit a drupe-like pome, red, brownish red, or orange to black, with persistent, incurved, fleshy sepals, containing pyrenes; pyrenes (1 or)2–5, bony, 1-seeded; seeds compressed; cotyledons plano-convex.

About 90 species in the broad sense: widespread in temperate N Africa, Asia (except Japan), Central America (Mexico), and Europe, most abundant in SW China; 59 species (37 endemic) in China.

The shrubs are widely planted as ornamentals for their attractive fruits and flowers, and as borders, hedges, and ground cover. Further studies are necessary to clarify a taxonomy complicated by hybridization and apomixis.

1 Inflorescence a compact compound corymb, more than 20-flowered; petals white, spreading; leaves large, more than 2.5 cm.   (2)
+ Inflorescence a lax corymb; less than 20-flowered, sometimes reduced to a solitary flower; petals pink and erect, or white and spreading; leaves often smaller.   (10)
2 (1) Leaf blade abaxially initially pilose, glabrate.   (3)
+ Leaf blade abaxially persistently densely tomentose.   (4)
3 (2) Leaf blade elliptic to ovate, apex obtuse or acute; fruit reddish yellow, ovoid to obovoid, 6–7 mm in diam.   8 C. glaucophyllus
+ Leaf blade oblong-lanceolate to oblong-oblanceolate, apex acuminate or acute; fruit red, globose, 4–5 mm in diam.   9 C. glabratus
4 (2) Deciduous shrub to small tree.   (5)
+ Evergreen or semievergreen shrub.   (6)
5 (4) Leaf blade narrowly elliptic to ovate-lanceolate; fruit bright red, ellipsoid, 4–5 mm in diam.   6 C. frigidus
+ Leaf blade ovate or elliptic-ovate; fruit purplish brown to black, ovoid, 6–8 mm in diam.   7 C. affinis
6 (4) Leaf blade abaxially persistently densely tomentose, apex obtuse or acute.   (7)
+ Leaf blade abaxially initially tomentose, glabrescent, apex acute to acuminate.   (8)
7 (6) Inflorescences densely yellow tomentose; fruit obovoid; leaf blade obovate to elliptic.   4 C. coriaceus
+ Inflorescences densely gray tomentose; fruit turbinate; leaf blade obovate-lanceolate to oblong- lanceolate.   5 C. turbinatus
8 (6) Pedicel very short or nearly absent; inflorescences 2–3.5 cm; fruit ellipsoid; pyrenes always 2.   3 C. harrovianus
+ Pedicel 2–4 mm; inflorescences 3–5 cm; fruit subglobose or pyriform; pyrenes 2–4.   (9)
9 (8) Leaf blade adaxially shallowly rugose, abaxially glaucous and tomentose; fruit subglobose.   1 C. salicifolius
+ Leaf blade adaxially deeply rugose, abaxially yellow tomentose; fruit pyriform.   2 C. rhytidophyllus
10 (1) Corymb 3–15(–20)-flowered, sometimes reduced to a solitary flower; leaf blade longer than 2 cm, rarely slightly shorter.   (11)
+ Inflorescence a solitary flower, rarely 2 or 3(–7)-flowered (but 3–9(–11)-flowered in C. sherriffii); leaf blade often less than 2 cm, rarely slightly longer.   (38)
11 (10) Petals white, spreading; fruit red.   (12)
+ Petals pink, very rarely white, erect; fruit red, rarely black (C. melanocarpus, C. tenuipes).   (18)
12 (11) Leaf blade abaxially glabrous or slightly pilose.   (13)
+ Leaf blade abaxially tomentose, rarely tomentose-villous.   (14)
13 (12) Pedicel and abaxial surfaces of leaf blade and hypanthium sparsely pilose.   15 C. submultiflorus
+ Pedicel and abaxial surfaces of leaf blade and hypanthium glabrous.   16 C. multiflorus
14 (12) Hypanthium abaxially glabrous.   (15)
+ Hypanthium abaxially tomentose or villous.   (16)
15 (14) Leaf blade oblong-elliptic, abaxially gray tomentose, sometimes adaxially initially pubescent; petals 3.5–4 mm; fruit purplish red, with slightly glaucous bloom, obovoid.   13 C. mongolicus
+ Leaf blade elliptic to broadly ovate, abaxially tomentose-villous, with glaucous bloom, always adaxially glabrous; petals 5–6 mm; fruit dark red, ovoid, rarely oblong.   14 C. hebephyllus
16 (14) Semievergreen shrub; anthers purplish red; leaf blade leathery; fruit with 2 pyrenes.   10 C. pannosus
+ Deciduous shrub; anthers yellow; leaf blade herbaceous; fruit with 1 or 2 pyrenes.   (17)
17 (16) Leaf blade apically obtuse, abaxially white tomentose; hypanthium abaxially tomentose; petals 2.5–3.5 mm; fruit ovoid to ellipsoid.   11 C. soongoricus
+ Leaf blade apically acute, rarely obtuse, abaxially gray tomentose; hypanthium abaxially villous; petals 4–5 mm; fruit subglobose.   12 C. silvestrii
18 (11) Leaf blade abaxially tomentose; fruit red, rarely black.   (19)
+ Leaf blade abaxially glabrous or pilose.   (31)
19 (18) Hypanthium abaxially glabrous or slightly puberulous.   (20)
+ Hypanthium abaxially densely tomentose or pilose.   (23)
20 (19) Fruit black; inflorescences 3–15-flowered, rachis tomentose.   29 C. melanocarpus
+ Fruit red; inflorescences 2–7-flowered, rachis glabrous to pilose.   (21)
21 (20) Hypanthium abaxially puberulous; flowers 2–4 in short fascicles, nearly 1/2 as long as leaves.   26 C. oliganthus
+ Hypanthium abaxially glabrous; flowers 2–7 in corymbs.   (22)
22 (21) Inflorescence nearly 1/2 as long as leaves; rachis and pedicels glabrous, rarely puberulous; leaf blade broadly elliptic, broadly ovate, or suborbicular; fruit subglobose, rarely ovoid, glabrous.   27 C. integerrimus
+ Inflorescence nearly as long as leaves; rachis and pedicels pilose; leaf blade ovate to oblong- ovate; fruit obovoid, puberulous.   28 C. gracilis
23 (19) Leaf blade apically obtuse, rarely acute.   (24)
+ Leaf blade apically acute to acuminate.   (26)
24 (23) Fruit black, ovoid, with 1 or 2 pyrenes; inflorescence 2–4-flowered.   19 C. tenuipes
+ Fruit red, globose to ovoid or obovoid, with 2 or 3 pyrenes; inflorescence 3–15-flowered.   (25)
25 (24) Leaf blade elliptic to ovate; inflorescence 3- to more than 10-flowered; fruit obovoid or ovoid- globose, with 2 pyrenes.   17 C. zabelii
+ Leaf blade broadly ovate to suborbicular; inflorescence 10–15-flowered; fruit oblong, with 3 pyrenes.   18 C. fangianus
26 (23) Rachis, pedicels, and hypanthium abaxially glabrous; fruit purplish black, subglobose, with 5 pyrenes.   25 C. reticulatus
+ Rachis, pedicels, and hypanthium abaxially pubescent; fruit red or orangish red, obovoid, ovoid, or subglobose, with 2–5 pyrenes.   (27)
27 (26) Rachis, pedicels, and hypanthium abaxially densely white tomentose; petals white; fruit obovoid, pyrenes 2.   20 C. wardii
+ Rachis, pedicels, and hypanthium abaxially pilose; petals usually pink, sometimes white or red; fruit subglobose to obovoid, pyrenes 3–5 or if pyrenes 2, then fruit ovoid.   (28)
28 (27) Inflorescences 5–11-flowered; fruit orangish red, with 3–5 pyrenes.   24 C. franchetii
+ Inflorescences 3–7(–11)-flowered; fruit dark or bright red, with 2–5 pyrenes.   (29)
29 (28) Fruit ovoid, dark red, with 2 or 3 pyrenes; petals red.   23 C. obscurus
+ Fruit subglobose or obovoid, bright red, with 3–5 pyrenes; petals white or pink.   (30)
30 (29) Inflorescence 2-flowered; petals pink; leaf blade elliptic to ovate, abaxially densely tomentose; fruit subglobose or obovoid, with 3–5 pyrenes.   21 C. dielsianus
+ Inflorescence 3–11-flowered; petals white; leaf blade ovate-lanceolate to oblong-ovate, abaxially sparsely tomentose; fruit subglobose, with 5 pyrenes.   22 C. glomerulatus
31 (18) Fruit red.   (32)
+ Fruit black.   (35)
32 (31) Hypanthium abaxially glabrous when old; flowers 5–13 in corymbs; leaf blade adaxially conspicuously rugose, veins prominently raised abaxially; styles 4 or 5; fruit with 4 or 5 pyrenes.   33 C. bullatus
+ Hypanthium abaxially slightly puberulous; flowers 1–9; leaf blade adaxially not rugose, veins raised but not prominently; styles 2; fruit with 2 pyrenes.   (33)
33 (32) Flowers (1 or)2 or 3(–5); leaf blade elliptic-ovate to ovate-lanceolate, adaxially pilose; fruit ellipsoid.   30 C. acuminatus
+ Flowers 3–9; leaf blade elliptic-lanceolate, adaxially glabrous or initially pilose, glabrescent; fruit subglobose or obovoid.   (34)
34 (33) Leaf blade elliptic-lanceolate; fruit subglobose, glabrous.   31 C. nitidifolius
+ Leaf blade broadly elliptic or broadly ovate, sometimes obovate or suborbicular; fruit obovoid, pilose or subglabrous.   32 C. schantungensis
35 (31) Hypanthium abaxially glabrous or subglabrous; inflorescences 5–10-flowered; fruit ovoid or subglobose.   37 C. ambiguus
+ Hypanthium abaxially pilose; inflorescences 2–25-flowered; fruit ellipsoid, obovoid, or subglobose.   (36)
36 (35) Leaf blade apically acute; inflorescence 2–5-flowered; fruit ellipsoid or obovoid, with 2 or 3 pyrenes.   34 C. acutifolius
+ Leaf blade apically acuminate; inflorescence 3–25-flowered; fruit subglobose or obovoid, with 3–5 pyrenes.   (37)
37 (36) Inflorescences 9–25-flowered; leaf blade adaxially rugose and conspicuously bullate; pyrenes 4 or 5, smooth.   35 C. moupinensis
+ Inflorescences 3–7-flowered; leaf blade adaxially not bullate, veins slightly impressed; pyrenes 3 or 4, furrowed and shallowly pitted on back.   36 C. foveolatus
38 (10) Petals white, spreading; fruit red, with 2 or 3(–5) pyrenes; prostrate or low evergreen shrub.   (39)
+ Petals red, rarely pink, erect; fruit red, rarely purplish black, with (1 or)2 or 3(or 4) pyrenes; spreading or erect, deciduous or semievergreen shrub.   (46)
39 (38) Abaxial surfaces of leaf blade and hypanthium tomentose.   38 C. buxifolius
+ Abaxial surfaces of leaf blade and hypanthium pilose.   (40)
40 (39) Leaf blade ovate or oblong-ovate to elliptic, rarely obovate; fruit with 2–5 pyrenes.   (41)
+ Leaf blade suborbicular, ovate, obovate, or elliptic to oblanceolate; fruit with 1 or 2 pyrenes.   (42)
41 (40) Leaf blade elliptic to oblong-elliptic, 1–2(–3) cm; fruit subglobose, with 4 or 5 pyrenes.   39 C. dammeri
+ Leaf blade ovate or oblong-ovate, rarely obovate, 0.9–1.5 cm; fruit ellipsoid-ovoid or ellipsoid, with 2 or 3 pyrenes.   40 C. morrisonensis
42 (40) Leaf blade elliptic, oblong or obovate to oblong-obovate, rarely oblanceolate, base cuneate.   (43)
+ Leaf blade suborbicular or broadly ovate, base broadly cuneate to rounded.   (45)
43 (42) Flowers solitary, rarely 2 or 3; shrub prostrate or suberect; leaf blade obovate to oblong-obovate, margin slightly revolute.   (44)
+ Flowers 3–9(–11); shrub erect; leaf blade oblong-obovate, rarely oblanceolate, margin not revolute.   43 C. sherriffii
44 (43) Prostrate shrubs; leaf blade obovate to oblong-obovate, 4–10 mm; fruit scarlet-red, 5–7 mm in diam.   41 C. microphyllus
+ Suberect shrubs; branches spreading to 40 cm; leaf blade elliptic to oblanceolate, rarely oblong, 6–16(–20) mm; fruit bright scarlet, 8–10 mm in diam.   42 C. conspicuus
45 (42) Leaf blade adaxially glabrous, abaxially pilose; flowers 1–3, petals white or tinged reddish, spreading; fruit red, obovoid, with 2 or 3 pyrenes.   44 C. rotundifolius
+ Leaf blade villous on both surfaces; flowers solitary, petals pink, erect; fruit dark red, ovoid- globose, with 2 pyrenes.   51 C. langei
46 (38) Leaf blade abaxially densely tomentose; flowers usually solitary.   (47)
+ Leaf blade abaxially glabrous or pilose; flowers 1–3(or 4).   (48)
47 (46) Petals red; hypanthium abaxially pilose; leaf blade suborbicular or broadly elliptic, apex obtuse, abaxially persistently yellow tomentose.   45 C. rubens
+ Petals pink; hypanthium abaxially glabrous; leaf blade ovate, rarely ovate-elliptic, apex acute or obtuse, abaxially tomentose, gradually glabrescent, subglabrous when old.   46 C. uniflorus
48 (46) Hypanthium abaxially glabrous; erect shrub.   (49)
+ Hypanthium abaxially pubescent; prostrate or erect shrub.   (53)
49 (48) Stems irregularly branched; fruit subglobose, sessile, erect.   (50)
+ Stems ± distichously branched; fruit obovoid or globose, shortly pedicellate, pendent.   (51)
50 (49) Leaf apically apiculate, rarely emarginate, blade suborbicular or orbicular-ovate, rarely broadly obovate, glabrous on both surfaces or slightly puberulous only along veins abaxially; petals pinkish.   58 C. apiculatus
+ Leaf apically acute, sometimes obtuse, blade ovate or elliptic-ovate, pilose on both surfaces; petals red.   59 C. sanguineus
51 (49) Branchlets conspicuously verruculose; leaf apically emarginate or partly mucronulate; flowers solitary.   57 C. verruculosus
+ Branchlets strigose, not verruculose; leaf apically acute or obtuse; flowers 1–3.   (52)
52 (51) Flowers solitary; stamens 20; leaf blade broadly ovate or broadly obovate, apex acute, rarely obtuse.   55 C. nitidus
+ Flowers 2 or 3; stamens 10; leaf blade suborbicular or broadly ovate, apex obtuse or acute.   56 C. taylorii
53 (48) Prostrate low shrub; flowers solitary or in pairs.   (54)
+ Erect shrub; flowers 2 or 3(or 4).   (56)
54 (53) Stems horizontally spreading, regularly distichously branched.   49 C. horizontalis
+ Stems tufted on ground, irregularly branched.   (55)
55 (54) Leaf blade thinly papery, margin undulate; fruit subglobose, 7–9 mm in diam., pyrenes 2 or 3.   47 C. adpressus
+ Leaf blade leathery, margin thick, not undulate; fruit ovoid, 5–6 mm in diam., pyrenes 2.   48 C. subadpressus
56 (53) Fruit black.   (57)
+ Fruit red.   (58)
57 (56) Leaf apically obtuse, rarely acute, blade ovate, rarely elliptic, adaxially glabrous, abaxially villous or subglabrous; fruit ellipsoid, pyrenes 1 or 2.   53 C. nitens
+ Leaf apically acute to acuminate, blade elliptic to ovate-elliptic, both surfaces appressed strigose-villous; fruit ovoid or subglobose, pyrenes 2 or 3.   54 C. harrysmithii
58 (56) Branchlets verruculose; leaf blade appressed pilose on both surfaces; fruit with 3 pyrenes.   52 C. chengkangensis
+ Branchlets not verruculose; leaf blade strigose-villous or subglabrous, fruit with 2 pyrenes.   (59)
59 (58) Leaf blade elliptic or broadly elliptic, rarely obovate, abaxially slightly puberulous, adaxially subglabrous, apex acute, rarely ± obtuse; fruit ellipsoid, shortly stalked.   50 C. divaricatus
+ Leaf blade broadly ovate to suborbicular, both surfaces strigose-villous, apex obtuse, rarely acute; fruit ovoid-globose, subsessile.   51 C. langei

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