FNA: Cotoneaster vs. Rosaceae tribe Maleae

Distribution

Eurasia; n Africa [Introduced in North America; introduced also in Europe, Pacific Islands, Australia]

HI; North America; Mexico; Central America; Eurasia; Africa; Atlantic Islands (Madeira) [Introduced in temperate southern hemisphere]

Discussion

Species ca. 400 (34, including 1 hybrid, in the flora).

A marked concentration of Cotoneaster species is found in the Himalaya and western China. The majority of species (ca. 90%) are apomictic, tetraploid (G. H. Kroon 1975; I. V. Bartish et al. 2001), and breed true when raised from seed.

Cotoneaster was long placed near Crataegus. Based on a reevaluation of flower and fruit characters, it is now believed to be more closely related to Heteromeles and Pyracantha (K. R. Robertson et al. 1991; J. R. Rohrer et al. 1991), but lacks thorns and toothed leaf margins. Recent molecular studies also distance Crataegus from Cotoneaster; as of yet, there is no consensus about the closest allies of Cotoneaster (C. S. Campbell et al. 1995, 2007; R. C. Evans and Campbell 2002). D. Potter et al. (2007) grouped Cotoneaster with Chamaemeles and Malus, but further study is needed to resolve its placement. The genera of Maleae are difficult to organize in a phylogeny, due in part to interfertility, lack of divergence, and reticulate evolution (Campbell et al. 2007).

Plants of Cotoneaster are used by apiarists, as the flowers have abundant nectar and are much loved by bees; the resulting honey is pale golden with a delicate flavor. In Asia, leaves and fruits are used in tea, and the wood for implements. Cotoneaster is used in India and Iran as the source of a sweet mannalike substance high in dextrose. The fruit of C. integerrimus Medikus has been used in the treatment of diarrhea. Many species are ornamentals, with brilliant fall foliage or colorful fruit persisting throughout the winter. The genus is bird-disseminated, mainly by American robins (Turdus migratorius Linnaeus) and cedar waxwings (Bombycilla cedrorum Vieillot) in North America. A few orange- or red-fruited species (C. franchetii, C. lacteus, C. pannosus, and C. simonsii) are weedy on the Pacific coast (J. M. Randall and J. Marinelli 1996; C. C. Bossard et al. 2000). Only C. divaricatus and the black-fruited C. lucidus are widely escaped in the interior.

More than 70 species of Cotoneaster are cultivated in North America, and additional escapes might be expected. Reports of several species in the wild appear to be unfounded. These include C. acutifolius Turczaninow and C. apiculatus Rehder & E. H. Wilson from Pennsylvania (A. F. Rhoads and W. M. Klein 1993; Rhoads and T. A. Block 2000), C. bullatus Bois from British Columbia (J. Pojar 1999), C. hupehensis Rehder & E. H. Wilson from New York (R. S. Mitchell and G. C. Tucker 1997), and C. multiflorus Bunge and C. racemiflorus (Desfontaines) K. Koch from Quebec (J. Cayouette et al. 1983). Without reference specimens it is often difficult to name adventives, given the lack of North American literature.

As with many plants of Maleae, Cotoneaster foliage is variable, yet mature leaf size, shape, texture, and luster on vigorous sterile shoots are usually diagnostic. The leaves of fertile shoots are usually smaller and less typical than those on sterile ones. Shaded plants have larger leaves than individuals in full sun. Deciduous species in flower have smaller leaves than when fruiting. A few species, such as C. gamblei, often hold leaves into January (at least on vigorous sheltered shoots) and key as evergreen shrubs, although naturalized populations in North America will eventually drop most leaves in late winter, especially where exposed. Several species are reported in the literature as semievergreen shrubs (for example, C. horizontalis, C. simonsii), but most of their leaves color and drop by midwinter where they are naturalized on the Pacific coast, so they are keyed as deciduous here. Shrubs with tomentose leaves can be sorted into evergreen and deciduous species by checking for the presence of darkened and weathered tomentum on the abaxial surfaces of overwintered leaves. Sunken (versus superficial) veins are mentioned in the keys and descriptions, always referring to the adaxial leaf surfaces. Collectors should observe this on fresh material and note it on herbarium labels. Sunken veins and wrinkling or bulging between the lateral veins can be obscured when pressing thin-leaved species. The number of veins in the descriptions represents the major lateral veins on one side of the leaf or midvein, most easily observed on the adaxial surface.

The pyrenes are usually pubescent at the apex, if exposed by the navel between the sepals on the summit of the pome. Extracted pyrenes are planoconvex or trigonous (like orange segments). Potentially useful taxonomic characters such as pyrene size, outline, and surface texture are not described here but warrant investigation. The style remnant is usually visible as a small peg or scar on the distal half of the inner surface or central keel of the pyrene; its location is easy to assess if fresh material is available. At least five pomes should be opened to determine the number of pyrenes present.

Identification of most naturalized species requires fruit characters and mature foliage; unfortunately, floral characters are also necessary to separate some species pairs. Floral and fruit dimensions are from fresh material and will be smaller on dried specimens. Floral diameter is provided throughout in subg. Chaenopetalum (Koehne) G. Klotz (species 1–16), which has rotate blossoms with spreading petals. For subg. Cotoneaster (species 17–34), the cupulate flowers are described by their floral length, including hypanthium and the erect petals.

Cotoneaster is properly treated as a masculine name (Yu T. T. 1954), although horticultural works still can be found with feminine endings applied to species epithets.

Key to Groups of Cotoneaster

Key to Species by Group

Group A. Shrubs, prostrate or long-trailing, rarely a few shoots to 30 cm tall; leaves deciduous or persistent, not tomentose.

Group B. Leaves deciduous, abaxial surfaces tomentose.

Group C. Shrubs or trees, erect; leaves persistent, abaxial surfaces tomentose.

Group D. Stems erect; leaves persistent, glabrous or sparsely hairy abaxially.

Group E. Stems erect; leaves deciduous, glabrous abaxially or hairy, not densely tomentose; pomes black or purple-black.

Group F. Leaves deciduous, abaxial surfaces glabrous or hairy, not densely tomentose; pomes orange or red, dark red, or red-purple.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Genera 29, species 550–840+ (18 genera, 270 species, including 18 hybrids, in the flora).

The family name Malaceae Small (1903) is a conserved name, with Malus as its type genus. In contrast, the family name Pyraceae Vest (1818), with Pyrus as its type, is not a conserved name. Although Maleae was published later than Pyreae (1869), a Rosaceae tribe that includes both Malus and Pyrus is to be called Maleae (see Melbourne Code, Article 19.5, Example 5).

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Key

Key to Groups of Cotoneaster

1. Shrubs, prostrate or long-trailing, rarely a few shoots to 30 cm tall	Group A
1. Shrubs or trees, erect, more than 30 cm tall	→ 2
2. Mature leaves tomentose abaxially (hairs obscuring veins and most or all of surfaces)	→ 3
2. Mature leaves glabrous or slightly hairy abaxially (surfaces easily visible)	→ 4
3. Leaves deciduous	Group B
3. Leaves persistent	Group C
4. Leaves persistent	Group D
4. Leaves deciduous	→ 5
5. Pomes black or dark purple; styles and pyrenes 1–3	Group E
5. Pomes orange to red, dark red, or red-purple; styles and pyrenes (1 or)2–5	Group F

Group A. Shrubs, prostrate or long-trailing, rarely a few shoots to 30 cm tall; leaves deciduous or persistent, not tomentose

1. Leaves deciduous, blades chartaceous, margins slightly undulate, not revolute; petals erect, dark red.	C. adpressus
1. Leaves persistent, blades coriaceous, margins slightly revolute, not undulate; petals spreading, white	→ 2
2. Styles and pyrenes 2(or 3); leaf blades 5–14 mm.	C. cochleatus
2. Styles and pyrenes 4 or 5; leaf blades 13–43 mm.	C. dammeri

Group B. Leaves deciduous, abaxial surfaces tomentose

1. Pomes purple-black, dark red, or ruby to maroon	→ 2
1. Pomes bright red or orange-red	→ 3
2. Pomes dark red or ruby to maroon; petals spreading, white; anthers purple to blackish purple; styles and pyrenes 1(or 2) .	C. monopyrenus
2. Pomes purple-black; petals erect, greenish white with pink or red; anthers white; styles and pyrenes 2–4.	C. melanocarpus
3. Filaments pink or pale pink; leaf blades: abaxial surfaces villose-strigose or sparsely to moderately tomentose.	C. tengyuehensis
3. Filaments red or dark red; leaf blades: abaxial surfaces densely tomentose	→ 4
4. Pomes orange-red.	C. qungbixiensis
4. Pomes bright red.	C. dielsianus

Group C. Shrubs or trees, erect; leaves persistent, abaxial surfaces tomentose

1. Shrubs, 0.5(–1.5) m; leaf blades 5–11 mm.	C. hodjingensis
1. Shrubs or trees, 1–9 m; leaf blades 15–120 mm	→ 2
2. Leaves: lateral veins ± superficial, rarely faintly sunken, adaxial surfaces flat between lateral veins; styles and pyrenes (1 or)2	→ 3
2. Leaves: lateral veins sunken, adaxial surfaces bulging or flat between lateral veins; styles and pyrenes 2–5	→ 5
3. Pomes slightly glaucous, sepals flat to incurved; flowers (7–)10–11 mm diam	C. crispii
3. Pomes not glaucous, sepals suberect; flowers 6–10 mm diam	→ 4
4. Leaf blades 15–35(–39) x 10–19(–26) mm; flowers 8.5–10 mm diam.	C. pannosus
4. Leaf blades 35–77 × 15–35 mm; flowers 6–6.5(–9) mm diam.	C. vestitus
5. Petals spreading, creamy white; anthers red-purple; styles and pyrenes 2; leaf blades obovate to elliptic, rarely broadly elliptic.	C. lacteus
5. Petals erect, pink or red; anthers white, pink, or purplish pink; styles and pyrenes (2 or)3–5 (always some with more than 2); leaf blades ovate to elliptic	→ 6
6. Anthers white	→ 7
6. Anthers pink to purplish pink or mostly white with pink-tinged sutures	→ 8
7. Filaments pink or pale pink, whitish distally; pomes bright red; leaves sparsely to moderately tomentose abaxially.	C. tengyuehensis
7. Filaments dark red, at least proximally; pomes orange-red; leaves densely tomentose abaxially.	C. qungbixiensis
8. Filaments pink, distally white; anthers pink to purplish pink; leaves usually flat adaxially between lateral veins; pomes obovoid, bases rounded or cuneate to obconic, straight-sided.	C. franchetii
8. Filaments red; anthers white with pink-tinged sutures; leaves usually bulging slightly adaxially between lateral veins (less obvious on shaded shoots); pomes subglobose to depressed-globose or globose, bases rounded.	C. sternianus

Group D. Stems erect; leaves persistent, glabrous or sparsely hairy abaxially

1. Shrubs, usually less than 0.8 m; leaf blade margins revolute	→ 2
1. Shrubs or trees, 1–10 m; leaf blade margins flat or revolute	→ 4
2. Leaf blades usually oblanceolate or oblong; pomes dull or slightly shiny, glaucous.	C. integrifolius
2. Leaf blades usually elliptic to narrowly elliptic; pomes shiny, not glaucous	→ 3
3. Leaf blades 6–12(–20) x 2–6(–8) mm, adaxial surfaces grayish green, dull to slightly shiny; styles and pyrenes 2(or 3).	C. conspicuus
3. Leaf blades 10–27 × 4–12 mm, adaxial surfaces dark green, shiny; styles and pyrenes 2–4(or 5) (always some with more than 2).	C. ×suecicus
4. Leaf blades: adaxial surfaces flat between lateral veins, veins superficial or essentially so; styles and pyrenes (1 or)2	→ 5
4. Leaf blades: adaxial surfaces wrinkled or bulging between lateral veins, or veins deeply sunken; styles and pyrenes 2–5	→ 6
5. Leaf blades 25–46 mm, adaxial surfaces dull, slightly glaucous; pomes 6–8 mm, globose or subglobose.	C. crispii
5. Leaf blades (25–)50–127 mm, adaxial surfaces shiny, not glaucous; pomes 8.5–12.8 mm, obovoid, rarely subglobose.	C. gamblei
6. Styles and pyrenes usually 3–5; petals erect, pink to red; anthers white.	C. tengyuehensis
6. Styles and pyrenes usually 2 or 3; petals spreading, white or creamy white; anthers red-purple	→ 7
7. Leaf blades shiny adaxially, narrowly elliptic to elliptic-lanceolate, 3–4 times as long as wide, margins strongly revolute, apices gradually tapered, acute or acuminate; styles and pyrenes 2 or 3(or 4); pomes maturing in October.	C. salicifolius
7. Leaf blades dull or slightly shiny adaxially, obovate to elliptic, rarely broadly elliptic, 2 times as long as wide, margins flat or slightly revolute, apices short-tapered, acute, acuminate, or obtuse; styles and pyrenes 2; pomes maturing in November.	C. lacteus

Group E. Stems erect; leaves deciduous, glabrous abaxially or hairy, not densely tomentose; pomes black or purple-black

1. Leaves usually bulging between lateral veins, veins sunken (easily obscured on pressed plants)	→ 2
1. Leaf blades: adaxial surfaces flat between lateral veins, veins superficial or slightly sunken	→ 3
2. Vigorous shoots with leaf tips short-acuminate or acute; pomes glaucous.	C. lucidus
2. Vigorous shoots with leaf tips attenuate to long-acuminate; pomes not glaucous.	C. villosulus
3. Styles and pyrenes 1 or 2; petals white, spreading, usually with hair tuft	→ 4
3. Styles and pyrenes 2–4; petals pink to red or purple, erect, glabrous	→ 5
4. Leaves of vigorous shoots 1.5 times as long as wide; pomes: sepal margins sparsely hairy.	C. monopyrenus
4. Leaves of vigorous shoots 2 times as long as wide; pomes: sepal margins glabrous.	C. transens
5. Leaf blades 8–28 mm; fall leaves turning bright red and gold; styles and pyrenes 2(or 3).	C. nitens
5. Leaf blades 33–45 mm; fall leaves lacking notable color; styles and pyrenes 2–4.	C. melanocarpus

Group F. Leaves deciduous, abaxial surfaces glabrous or hairy, not densely tomentose; pomes orange or red, dark red, or red-purple

1. Adaxial surfaces of fresh leaves strongly wrinkled or bulging between lateral veins (easily obscured when pressed); styles and pyrenes (4 or)5.	C. rehderi
1. Adaxial surfaces of fresh leaves flat between lateral veins, sometimes faintly bulging (some taxa with faintly sunken midvein or wavy margins); styles and pyrenes (1 or)2–5	→ 2
2. Leaf blades 4–30(–36) mm	→ 3
2. Leaf blades (21–)30–150 mm	→ 7
3. Shrubs 1.5–4(–6) m; pomes usually cylindric to obovoid (proportionally narrower)	→ 4
3. Shrubs 0.5–1 m; pomes usually broadly obovoid	→ 5
4. Plants ± strict or ascending, lateral branches mostly suppressed, short, ± straight; stamens 20 (found under calyx in fruit, with careful dissection); pomes bright orange to orange-red, usually obovoid; styles and pyrenes (2 or)3–5; leaf blades on vigorous shoots usually broadly elliptic to ovate.	C. simonsii
4. Plants spreading with rounded shape, lateral branches well developed, usually long, arching; stamens 10–15; pomes dark red to ruby (finally blackish red), cylindric, oblong-ellipsoid, or narrowly obovoid; styles and pyrenes (1 or)2(or 3); leaf blades on vigorous shoots usually elliptic or broadly elliptic.	C. divaricatus
5. Leaf blades on vigorous shoots usually broadly elliptic.	C. horizontalis
5. Leaf blades on vigorous shoots usually broadly obovate to orbiculate	→ 6
6. Leaf blades on vigorous shoots obovate-orbiculate; pomes bright red; sepals sparsely pilose.	C. atropurpureus
6. Leaf blades on vigorous shoots orbiculate, sometimes broadly obovate; pomes orange-red; sepals glabrous.	C. hjelmqvistii
7. Leaf blades (25–)50–150 mm; anthers purple, red-purple, or black; pomes red to purplish red or maroon to purple-black	→ 8
7. Leaf blades usually less than 50 mm; anthers white; pomes orange to dark red	→ 10
8. Pomes maroon to purple-black, sepals sparsely villous, margins glabrous.	C. transens
8. Pomes red, sepals densely villous, margins tomentose	→ 9
9. Pomes 4–6 × 4–5 mm, sepals erect or ascending; pedicels (and peduncles) densely tomentose.	C. frigidus
9. Pomes 8.5–12.8 × 8–12 mm, sepals flat; pedicels (and peduncles) sparsely villose-strigose.	C. gamblei
10. Petals spreading; flowers 12–14 mm diam.; petioles 5–8 mm.	C. magnificus
10. Petals erect or erect-incurved; flowers 4–10 mm diam.; petioles 1–5 mm	→ 11
11. Styles and pyrenes (2 or)3–5	→ 12
11. Styles and pyrenes 2(or 3)	→ 13
12. Pomes orange to orange-red; sepals erect or ascending, rarely nearly flat (not meeting over and obscuring pome summit), navel open; flowers pendent.	C. simonsii
12. Pomes bright red; sepals flat (meeting over and obscuring pome summit), navel closed; flowers erect or ascending.	C. tengyuehensis
13. Pomes bright to dark red; leaves on vigorous shoots elliptic, broadly elliptic, or ovate, sometimes suborbiculate.	C. fangianus
13. Pomes bright orange-red; leaves on vigorous shoots elliptic to narrowly ovate (usually proportionally narrower).	C. miniatus

1. Leaf margins usually horny; carpels free; flowers: perianth and androecium perigynous; fruits woody capsules surrounded by a hypanthium, splitting into 5 follicles; seeds winged	Vauquelinia
1. Leaf margins not horny; carpels ± adnate to hypanthium; flowers: perianth and androecium epigynous; fruits pomes; seeds not winged or pyrenes.	→ 2
2. Fruiting carpels woody or bony.	→ 3
3. Leaf margins entire; stipules short-adnate to petiole; stems unarmed; sepals erect in flower; petal base clawed.	Cotoneaster
3. Leaf margins ± serrate, crenate, serrulate, or crenulate, sometimes entire; stipules free; stems usually armed (sometimes with compound thorns), sometimes unarmed; sepals spreading in flower; petal base slightly or barely clawed.	→ 4
4. Leaves persistent or late-deciduous; flowers 3–10(–12) mm diam., hypanthium campanulate; pomes 3–8 mm diam.	Pyracantha
4. Leaves deciduous (sometimes winter-persistent in south); flowers 8–35 mm diam., hypanthium ± obconic; pomes 6–40 mm diam.	→ 5
5. Flowers 8–25 mm diam., stamens 5–20 (rarely 30–45); pomes yellow to red or purplish to black mature, 6–20(–25) mm diam.; pyrenes 1–5; short shoots present; inflorescences 1–50-flowered, domed panicles, corymbose, or flowers solitary.	Crataegus
5. Flowers 25–35 mm diam., stamens 25–35(–40); pomes brownish, 15–40 mm diam.; pyrenes 5; short shoots rare or absent; inflorescences 1(or 2) flowered	Mespilus
2. Fruiting carpels cartilaginous.	→ 6
6. Stems armed (thorns present).>	→ 7
7. Stipules persistent; pedicels short or absent; styles basally connate 1/3 of length; pome flesh without stones; stamens 40–60; fruiting sepals deciduous.	Chaenomeles
7. Stipules usually deciduous or caducous; pedicels present; styles distinct or basally connate; pome flesh with stones (at least near carpels and epidermis); stamens 15–50; fruiting sepals persistent or deciduous.	→ 8
8. Pome flesh with stone cells adjacent to carpels and epidermis; styles basally connate.	Malus
8. Pome flesh with abundant stone or grit cells; styles distinct.	Pyrus
6. Stems unarmed.	→ 9
9. Inflorescences: flowers solitary or 1–5-flowered, corymbs or cymes.	→ 10
10. Pomes yellow; ovules (seeds) many; inflorescences: flowers solitary.	→ 11
11. Leaf margins entire, abaxial surfaces densely villous; buds ovoid, apices obtuse or acuminate, tomentose; young branches tomentose, glabrescent; stipules caducous; flowers 40–50 mm diam., petals white or light pink, suborbiculate, ovate, or obovate, stamens equal to or slightly longer than petals; pomes pyriform or subglobose, 30–50 mm.	Cydonia
10. Pomes pink, yellow-orange, purple, purplish or bluish black, brownish, or nearly black; ovules (seeds) (1 or)2; inflorescences 1–5-flowered, cymes or corymbs.	→ 12
12. Pomes yellow-orange; stipules adnate to petiole and base of blade; petioles short or absent; leaf blades elliptic to oblanceolate or linear.	Peraphyllum
12. Pomes pink, bluish or purplish black, purple, brownish, or nearly black; stipules free; petioles present; leaf blades elliptic, elliptic-oblong, or oblong-ovate to orbiculate.	→ 13
13. Leaves leathery, drought-deciduous or persistent; sepals nearly orbiculate (inner broadly deltate), petals round or kidney-shaped; carpels barely connate or distinct, styles lateral; pomes translucent, vivid pink, drying purplish black.	Malacomeles
13. Leaves membranous to coriaceous (not leathery), deciduous; sepals triangular to lanceolate, petals linear to orbiculate; carpels connate, styles terminal; pomes bluish or purplish to nearly black, pinkish or maroon-purple, dark purple-blue, or brownish.	Amelanchier
9. Inflorescences (4 or)5–400+-flowered, panicles, sometimes racemes, corymbs, or subumbellate.	→ 14
14. Leaves persistent, leathery; carpels basally adnate to hypanthium.	→ 15
15. Leaf margins flat; flowers 15–20 mm diam.; pedicels short or nearly absent; hypanthia usually tomentose; stamens 20; carpels connate, styles (2–)5; pomes soft apricot yellow, 20–30 mm (diam.).	Eriobotrya
15. Leaf margins revolute; flowers 10 mm diam.; pedicels present; hypanthia glabrous or weakly floccose; stamens 10; carpels distinct, styles 2 or 3; pomes usually bright red, sometimes yellow, 5–10 mm	Heteromeles
14. Leaves usually deciduous, sometimes semipersistent or persistent (then margin entire), membranous to ± leathery; carpels adnate to all or 1/3–1/2 of hypanthium.	→ 16
16. Pome flesh with stones or sclereids.	→ 17
17. Inflorescences terminal, 6–400+-flowered flat-topped or rounded panicles; flowers opening after leaf expansion, 5–17 mm diam.; sepals erect or ascending; leaves pinnately divided, sometimes simple or lobed.	Sorbus
17. Inflorescences terminal on short shoots, 4–9-flowered racemes or simple corymbs, umbel-like; flowers developing with or before leaves, 15–45 mm diam.; sepals reflexed; leaves simple.	Pyrus
16. Pome flesh without stones.	→ 18
18. Stipules adnate to petiole, persistent	Aronia
18. Stipules free, caducous or deciduous.	→ 19
19. Leaves persistent or deciduous; inflorescences corymbose or subumbellate; pomes red or black.	Photinia
19. Leaves deciduous; inflorescences racemes; pomes bluish or purplish to nearly black, pinkish or maroon-purple, dark purple-blue, or brownish	Amelanchier