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9. Ericaceae A. L. Jussieu

杜鹃花科 du juan hua ke

Authors: Mingyuan Fang, Ruizheng Fang, Mingyou He, Linzheng Hu, Hanbi Yang, Haining Qin, Tianlu Min, David F. Chamberlain, Peter Stevens, Gary D. Wallace & Arne Anderberg

Agapetes mannii

Credit: Harvard University Herbaria

Plants usually woody, or herbs, sometimes lacking chlorophyll. Leaves spiral, sometimes decussate, margin often toothed. Inflorescence racemose; bracteoles paired, basal. Flowers (4 or)5-merous. Calyx imbricate. Corolla connate, imbricate. Stamens 10, sometimes with spurs or awns, dehiscing by pores, pollen in tetrads, rarely single. Ovary superior or inferior, placentation axile, rarely parietal, often many ovules per locule. Style ± as long as corolla, slender. Fruit a capsule or berry, rarely a drupe; calyx persistent.

About 125 genera and 4000 species: widely distributed in temperate and subarctic regions, also at high elevations in tropical regions; 22 genera and 826 species (524 endemic) in China.

The Monotropoideae are here included in the Ericaceae; previously in FRPS (56: 157–216. 1990), they were treated as the Pyrolaceae. Chiogenes, recorded from China in FRPS (57(3): 69–71. 1991), is here included in Gaultheria. Over the last half century, the Empetraceae have usually been separated from, but closely associated with, the Ericaceae. In their ecology, leaf morphology and insertion, rusts, embryology, stamen anatomy, etc., they largely agree with that family. Molecular data place Empetrum and its relatives firmly within the Ericaceae, and in particular within the subfamily Ericoideae, in agreement with phytochemical and palynological data, and there they are best recognized as a separate tribe. There are distinctive features of the Empetreae that were responsible for their past familial status, e.g., reduced perianth with separate members, low ovule number, enlarged stigmas, etc. However, these are likely to be derived features associated with wind pollination. See Kron et al. (Bot. Rev. 68: 335–423. 2002) and the recent treatment of the Ericaceae by Stevens et al. (in Kubitzki, Fam. Gen. Vasc. Pl. 6: 145–194. 2004).

Several genera and many species are ornamentals. Some fruits of Vaccinium in N China are sweet and edible, but of no particular value to humans. Some species of Chamaedaphne, Craibiodendron, Leucothoë, Lyonia, Pieris, and Rhododendron contain more or less toxic diterpenes, which are harmful to humans or domestic animals.

Fang Rhuicheng, Yang Hanbi & Chin Tsenli. 1999. Ericaceae (1). In: Fang Rhuicheng, ed., Fl. Reipubl. Popularis Sin. 57(1): 1–213; Hu Wenkuang, Hu Lincheng, Fang Mingyuan & He Mingyou. 1994. Ericaceae (2). In: Hu Lincheng, Fang Mingyuan, ed., Fl. Reipubl. Popularis Sin. 57(2): 1–438; Hsu Tingzhi, Gao Baochun, Fang Rhuicheng & Huang Shuhua. 1991. Ericaceae (3). In: Fang Rhuicheng, ed., Fl. Reipubl. Popularis Sin. 57(3): 1–207; Chou Yiliang & Zhou Ruichang. 1990. Pyrolaceae. In: Fang Wenpei & Hu Wenkuang, eds., Fl. Reipubl. Popularis Sin. 56: 157–216; Ming Tien lu. 1980. Empetraceae. In: Cheng Mien & Ming Tien lu, eds., Fl. Reipubl. Popularis Sin. 45(1): 60–62.


1 Herbs to subshrubs; petals free.   (2)
+ Plants woody; petals nearly always fused (absent and replaced by petaloid sepals in Empetrum).   (7)
       
2 (1) Achlorophyllous plant without green leaves; anthers without pores.   (3)
+ Plant usually with green leaves, usually autotrophic; anthers opening by pores.   (4)
       
3 (2) Fruit a capsule; ovary 5-loculed, with axile placentation.   6 Monotropa
+ Fruit a berry; ovary 1-loculed, with parietal placentation.   7 Monotropastrum
       
4 (2) Subshrub; leaves along stem; flowers in corymbs or umbels, or solitary; filaments ciliate on lower half; style thick, short.   2 Chimaphila
+ Herb; leaves generally basal; flowers in racemes, or solitary; filaments not ciliate; style slender.   (5)
       
5 (4) Flower solitary; capsule opening from tip to base, valves without fibers at margins.   3 Moneses
+ Flowers in racemes; capsule opening from base to tip, valves with fibers at margins.   (6)
       
6 (5) Leaves cauline; racemes with flowers on one side; upper portion of scape papillate; disk present.   4 Orthilia
+ Leaves appearing basal; racemes with flowers on all sides; scape not papillate; disk absent.   5 Pyrola
       
7 (1) Fruit fleshy.   (8)
+ Fruit a capsule.   (11)
       
8 (7) Ovary superior, free from calyx; seeds 4–9, enclosed in stones.   (9)
+ Ovary inferior, completely or rarely mostly adnate to hypanthium; seeds separate, numerous.   (10)
       
9 (8) Leaf blade linear, margin entire, slightly reflexed; corolla absent, sepals petaloid, free.   13 Empetrum
+ Leaf blade broad, margin serrate or crenate, flat; corolla urceolate, shortly lobed.   8 Arctous
       
10 (8) Corolla usually short, to 1 cm, urceolate or campanulate, rarely tubular; anthers free; pedicels usually not expanded toward apex; plants usually terrestrial, rarely epiphytic.   21 Vaccinium
+ Corolla usually long, (0.5–)2–6 cm, tubular or cylindric, rarely urceolate; anthers weakly connate or free; pedicels usually expanded toward apex, sometimes forming a cup below calyx; plants often epiphytic.   22 Agapetes
       
11 (7) Persistent calyx ± accrescent and succulent; capsule loculicidal, enclosed in fleshy calyx; anthers with 2–4 awns or minute projections.   20 Gaultheria
+ Persistent calyx withered.   (12)
       
12 (11) Capsule septicidal; anthers without appendages.   (13)
+ Capsule loculicidal; anthers with or without appendages.   (16)
       
13 (12) Corolla lobes free.   11 Ledum
+ Corolla sympetalous.   (14)
       
14 (13) Corolla actinomorphic or somewhat zygomorphic, funnel-shaped, campanulate, rarely rotate or cylindric, more than 10 mm; stamens exserted, rarely included; leaf blade not linear.   12 Rhododendron
+ Corolla actinomorphic, cylindric or urceolate, 4–7(–11) mm; stamens included; leaf blade linear to linear-oblong.   (15)
       
15 (14) Inflorescence shortly racemose to subcapitate; pedicels very short; corolla cylindric; anthers opening by slits.   9 Diplarche
+ Inflorescence umbellate; pedicels elongate, 2–2.5(–4) cm; corolla urceolate; anthers opening by terminal pores.   10 Phyllodoce
       
16 (12) Inflorescence a solitary flower; leaves decussate, imbricate, sessile, scalelike, 5–8 mm.   14 Cassiope
+ Inflorescences racemose, paniculate, or umbellate, many flowered, or flowers rarely solitary in Enkianthus; leaves spirally arranged, petiolate, leaf blade never scalelike, more than 3 cm.   (17)
       
17 (16) Anthers tipped with 2 awns or with dorsal spurs; filaments straight; leaf margin usually serrate.   (18)
+ Anthers usually without appendages; filaments geniculate, rarely straight; leaf margin entire.   (20)
       
18 (17) Anther awns reflexed; inflorescences paniculate.   17 Pieris
+ Anther awns erect or spreading; inflorescences racemose, umbellate, or corymbose.   (19)
       
19 (18) Inflorescences umbellate or corymbose, rachis abbreviated or nearly absent; corolla broadly campanulate to urceolate; seeds winged.   1 Enkianthus
+ Inflorescences racemose, rachis elongate; corolla suburceolate to tubular; seeds angled or rounded.   15 Leucothoe
       
20 (17) Young shoots and both surfaces of leaves lepidote.   19 Chamaedaphne
+ Young shoots and both surfaces of leaves not lepidote.   (21)
       
21 (20) Corolla tubular or urceolate; capsule subglobose, less than 5 mm in diam., sutures thickened; seeds wingless.   18 Lyonia
+ Corolla shortly campanulate; capsule depressed globose, more than 10 mm in diam., sutures not thickened; seeds winged on 1 side.   16 Craibiodendron

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