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heath family

azalea, Labrador-tea, menziesia, rhododendron

Habit Herbs, subshrubs, shrubs, or trees, (rarely vines), perennial, deciduous or evergreen, usually autotrophic, sometimes mycotrophic (subfam. Subshrubs, shrubs, or trees, (rhizomatous or not).
Stems

(absent in some Monotropoideae) erect or decumbent to prostrate, glabrous or hairy, (aerial stems sometimes produced from suckers, rhizomes, or corms), pith solid (hollow, with diaphragms in Agarista).

erect (and/or creeping, prostrate in R. groenlandicum, R. lapponicum, R. tomentosum);

twigs hairy or glabrous, (older twigs without peglike projections).

Leaves

(reduced or absent in some Monotropoideae), usually cauline, sometimes in basal rosettes (subfam.

persistent or deciduous, alternate;

petiole present;

blade coriaceous to membranous, (base cuneate, rounded, or obtuse), margins entire, (sometimes ciliate, plane to revolute, abaxial surface 1/3+ visible except sometimes in bud, surfaces hairy and/or scaled, midvein hairy or not, adaxial surface sometimes impressed; venation usually brochidodromous). (Winter buds usually large, with imbricate scales; floral buds usually larger than vegetative buds.) Inflorescences terminal (axillary in R. albiflorum), short, corymbiform or rounded racemes (fasciculate racemes in R. canadense, R. lapponicum; fasciculate in R. albiflorum), 3–35-flowered (1–2-flowered in R. albiflorum);

perulae brownish, scalelike, dry. (Pedicels horizontal to erect (recurved);

bracteoles 2, brownish, scalelike, basal.) Flowers bisexual, weakly bilaterally symmetric (radially symmetric in R. columbianum, R. groenlandicum, R. tomentosum, scented or not);

sepals 5[–9], slightly connate;

petals 5[–9], strongly to only slightly connate (sometimes nearly distinct), corolla deciduous, rotate to campanulate or funnelform;

stamens 5–12[–20], included to long-exserted; (filaments usually unequal, usually unicellular-hairy, glabrous in R. vaseyi, glabrous or proximally unicellular-hairy in R. groenlandicum, R. lapponicum);

anthers without awns, dehiscent by terminal pores;

ovary 5[–18]-locular;

style inserted in slight depression at ovary apex or smoothly intergrading, (usually curved, long, slender);

stigma capitate.

Inflorescences

terminal or axillary racemes, umbels, corymbs, panicles, fascicles, spikes, or solitary flowers.

Flowers

usually bisexual, rarely unisexual (subfam.

Fruits

capsular and dehiscent (loculicidal, septifragal, or septicidal), or drupaceous (axis fibrous or soft in some Monotropoideae) or baccate (rarely each surrounded by accrescent or fleshy calyx in Gaultheria) and indehiscent.

capsular, elongate, usually ovoid to cylindric, dehiscence usually basipetally septicidal (acropetally septicidal in R. columbianum, R. groenlandicum, R. tomentosum).

Seeds

1–10(–1000+), tan to yellowish brown or brown, ellipsoid, ovoid or spheroidal, or fusiform to flattened, or oblong (sometimes 3-sided);

testa thin (bony in subfam.

(10–)100+, flattened-ellipsoidal to fusiform, often tailed, ± winged;

testa smooth.

Monotropoideae

), usually chlorophyllous and autotrophic, sometimes achlorophyllous and heterotrophic (subfam.; monotropoideae ), aromatic compounds (e.g., methyl salicylate) sometimes present (Gaultheria).;

monotropoideae ), usually alternate or pseudoverticillate, sometimes opposite or, rarely, whorled, simple;

stipules absent;

petiole present or absent;

blade plane or acicular, often coriaceous, margins entire or toothed, plane or revolute.;

monotropoideae and subfam.

Ericoideae

), radially symmetric (sometimes slightly bilaterally symmetric in subfam.; ericoideae );

perianth and androecium hypogynous (epigynous in some Vaccinioideae);

hypanthium absent;

sepals absent or (2–)4–5(–7), distinct or connate basally;

petals (2–)4–5(–8), rarely absent or highly reduced, connate or distinct, not sticky (covered with sticky exudate in Bejaria), corolla absent or rotate to crateriform, campanulate, cylindric, globose, or urceolate (salverform in Epigaea);

intrastaminal nectary disc present or absent;

stamens (2–)5–8(–10) [14, 16, 20];

filaments distinct;

anthers inverted during development, often with awns, dehiscent by pores or short slits (at apparent apex) or slits (lateral);

pistils 1, 4–5-carpellate;

ovary superior (inferior in some Vaccinioideae), incompletely (2–)5–10-locular (1-locular in some Monotropoideae), often furrowed or lobed externally;

placentation axile or parietal;

ovules anatropous, unitegmic, tenuinucellate;

styles 1, straight or declinate (curved in Elliottia), hollow;

stigmas 1, capitate or peltate to funnelform, usually 5-lobed.

Arbutoideae

and subfam.

Vaccinioideae

);

embryo usually straight, fusiform, rarely minute and undifferentiated;

endosperm abundant, cellular, fleshy.

x

= 13.

Ericaceae

Rhododendron

Distribution
Nearly worldwide
[BONAP county map]
from USDA
North America; Europe; Asia; Australia
[BONAP county map]
Discussion

Genera ca. 120, species ca. 4100 (46 genera, 212 species in the flora).

The closest relatives of the broadly defined Ericaceae are Clethraceae and Cyrillaceae. Some phylogenies show Cyrillaceae as sister to Ericaceae; other analyses have Clethraceae and Cyrillaceae as closest relatives to each other, together forming the sister group to Ericaceae. Monotropa and related genera (genera 5–12 of this treatment), and Pyrola and related genera (genera 1–4 of this treatment) have been treated as families Monotropaceae and Pyrolaceae. Not all botanists agreed with this, as summarized by G. H. M. Lawrence (1951): “Many botanists (including Hutchinson) have held the view that the Pyrolaceae are not sufficiently distinct from the Ericaceae to be treated as a separate family.” Differences in habit, floral features, and pollen have helped maintain family status for Pyrolaceae and Monotropaceae in regional floras. Molecular and morphological analyses (K. A. Kron et al. 2002) show these lineages embedded within Ericaceae. Similarly, Empetraceae has been demonstrated to be nested within Ericaceae and is here included in the Ericaceae.

P. F. Stevens (2004) recognized eight subfamilies within Ericaceae; six of these are represented in the flora area. Subfamily Enkianthoideae, basal in recent phylogenies of the family, forms a sister clade to the remaining subfamilies. The subfamily includes only the single genus Enkianthus Loureiro (12 species), native to temperate eastern Asia. Enkianthus campanulatus (Miquel) G. Nicholson is cultivated occasionally in the northeastern and northwestern United States (M. A. Dirr 1998). Subfamily Styphelioideae Sweet (subfam. Epacridoideae Arnott) of the Southern Hemisphere (especially diverse in Australia with such genera as Astroloma R. Brown, Epacris Cavanilles, and Styphelia Smith), long considered a close relative of the Ericaceae, has been demonstrated as embedded within the Ericaceae. As G. H. M. Lawrence (1951) noted, distinctions between the two families are weak.

Studies in the last several decades, especially since 1990 including molecular data, have resulted in rearrangements of generic limits in the Ericaceae. These are discussed under the various genera; for the reader’s convenience they are summarized here. Ledum is included in Rhododendron; Leiophyllum and Loiseleuria are included in Kalmia; and Hypopitys is included in Monotropa. Arctous is separated from the much larger Arctostaphylos, to which it is inferred to form a sister clade. Eubotrys is segregated from Leucothoë, with which it has often been combined. Vaccinium is treated in a broad sense, to include segregates such as Oxycoccus; although Vaccinium is decidedly polymorphic, this seems a workable approach until generic limits in the Vaccinieae Reichenbach are better understood.

Most Ericaceae are evergreen shrubs. Some species are deciduous, notably in Rhododendron and Vaccinium. The propensity of members of the family to grow in acidic soils is well known. Although the family Ericaceae is generally regarded as exclusively growing on acidic substrates, some members of the family do occur in neutral or alkaline soils in North America and elsewhere.

Ericaceae are widely distributed in the Northern Hemisphere, almost ubiquitous except in desert areas. In the tropics, especially in South America, the family is diverse in upland and montane areas, and notably diverse in such genera as Bejaria and Cavendishia Lindley. Rhododendron, with centers of diversity in the Himalayas, New Guinea, and eastern North America, and Erica, diverse in southern Africa and Europe, are the largest genera in the family. The largest genus in the flora area is Arctostaphylos, with most species endemic to California and bordering states.

Species among some genera of the family enrich the human condition with edible fruits. In North America, by far the most important of these is Vaccinium. The high-bush blueberry, V. corymbosum, is cultivated in some states, notably Michigan, New Jersey, and North Carolina, and the low-bush blueberry, V. angustifolium, in Maine, Quebec, and the Canadian Maritime Provinces. The fruits of V. macrocarpon, the cranberry, are cultivated commercially in some provinces, including British Columbia, Nova Scotia, and Quebec, and some states including Massachusetts, Oregon, Rhode Island, Washington, and Wisconsin. Vaccinium vitis-idaea, lingonberry, is collected and sold in Newfoundland and Labrador as fresh fruits and preserves, and is an important addition to diet and health in the more northern areas of Canada. The leaves of Rhododendron groenlandicum (Ledum groenlandicum), Labrador tea, are used for a beverage in parts of its transcontinental range. The foliage of some genera, notably Leucothoë, Lyonia, and Rhododendron, contains andromedotoxins and is occasionally implicated in poisonings of humans, domestic pets, and livestock (J. M. Kingsbury 1964; S. D. Mancini and J. M. Edwards 1979). Kalmia also is reportedly toxic, perhaps why it is called sheep laurel, in addition to probably being allelopathic and thus detrimental to reforestation in some situations in the eastern boreal forest. Lyonia ferruginea is a valuable “forest product” in Florida. It is harvested and the stems are used in interior decoration; once silk leaves have been added, they are marketed as “artificial” plants.

Some species of Ericaceae, both native and exotic, are cultivated and of importance to the horticultural industry (M. A. Dirr 1998). Chief among these are Kalmia and Rhododendron (including many deciduous species known as “azaleas”). Other cultivated genera include Arbutus, Elliottia, Enkianthus, Leucothoë, Menziesia, Oxydendrum, and Pieris. Rock garden plants include Arctostaphylos uva-ursi, Kalmia buxifolia, K. procumbens, and Rhododendron lapponicum, as well as species of the Old World genera Calluna and Erica. Shrubs of some genera, including Gaultheria, Gaylussacia, and Vaccinium, are prominent in the understories of deciduous and evergreen forests, especially in regions of acidic soil, such as the southeastern United States. Wetlands in much of Canada and the northern United States support dense populations of ericaceous shrubs, notably Andromeda polifolia and Chamaedaphne calyculata; Kalmia spp., Rhododendron canadense, and R. groenlandicum may be as prominent depending on the region.

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

Anthodendron Reichenbach; Azaleastrum (Planchon ex Maximowicz) Rydberg; Biltia Small; Hymenanthes Blume; Ledum Linnaeus; Rhodora Linnaeus; Tsusiophyllum Maximowicz; Tsutsusi Adanson

Species ca. 1000 (25 in the flora).

R. Good (1974) provided an excellent discussion of the phytogeography of Rhododendron, which occurs in arctic to tropical montane regions of North America, Europe, and Asia, extending southward to northeastern Australia. Its main centers of diversity are in the mountains of southern China and bordering countries, with a secondary center in New Guinea; eastern North America is a third center of diversity.

The species of Rhododendron are divided into subgenera (see references and discussion under various representative species). The subgenera are occasionally segregated; see, for example, H. F. Copeland (1943). The name Azalea Linnaeus often has been used for species with deciduous leaves; such use is misapplied.

Therorhodion has usually been included within Rhododendron, and these species are then placed in subg. Therorhodion (Maximowicz) A. Gray (W. R. Philipson and M. N. Philipson 1986). Here, Therorhodion is recognized as a genus distinct from Rhododendron (see K. A. Kron and W. S. Judd 1990; Kron 1997; Kron et al. 2002) because of its foliaceous perulae and base chromosome number of 12. Therorhodion probably is sister to the remaining genera of Rhodoreae (most of which belong within Rhododendron). Rhododendron is closely related to Menziesia, and the recognition of the latter may render Rhododendron paraphyletic (Kron 1997).

Numerous non-native species are grown for their beautiful flowers, and some of these, e.g., Rhododendron molle (Blume) G. Don, R. luteum Sweet, R. simsii Planchon, and R. obtusum (Lindley) Planchon, sometimes persist after cultivation. Probably, all species are poisonous due to the presence of andromedotoxin, a resinoid (M. D. McGee 1973).

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

Key

Synoptic Key to Subfamilies

1. Plants achlorophyllous, heterotrophic, herbaceous; leaves absent or reduced, scalelike, blade plane; inflorescences racemes; fruits not fleshy
a. Ericaceae subfam. Monotropoideae
1. Plants chlorophyllous, autotrophic, woody or herbaceous; leaves not reduced, blade plane or acicular; inflorescences usually fascicles, racemes, panicles, spikes, umbels, or corymbs, sometimes solitary flowers; fruits fleshy or not
→ 2
2. Plants herbs or subshrubs; leaves persistent, mostly basal (reduced or absent); petals distinct; fruits capsular and dehiscence loculicidal or baccate and indehiscent
a. Ericaceae subfam. Monotropoideae
2. Plants vines, subshrubs, shrubs, or trees; leaves deciduous or persistent, cauline; petals connate or distinct; fruits capsular and dehiscence loculicidal or septicidal (sometimes septifragal) or baccate or drupaceous and indehiscent
→ 3
3. Fruit pulp ± juicy or mealy; pyrenes with stonelike endocarp, seeds 1-5, sometimes connate; corollas usually urceolate, sometimes cylindric; bark smooth, not furrowed, usually flaky
b. Ericaceae subfam. Arbutoideae
3. Fruit pulp ± juicy or dry; testa thin, rarely surrounded by bony endocarp, seeds (1-)10 (-300), distinct; corollas rotate, campanulate, cylindric, or urceolate (rarely salverform), sometimes absent; bark smooth or furrowed, not flaky
→ 4
4. Subshrubs; leaves persistent; inflorescences solitary flowers; (arctic and alpine regions)
→ 5
4. Shrubs (rarely vines, subshrubs, or trees); leaves deciduous or persistent; inflorescences fascicles, racemes, panicles, or spikes (rarely solitary flowers)
→ 6
5. Inflorescences axillary; leaves opposite
c. Ericaceae subfam. Cassiopoideae
5. Inflorescences terminal; leaves alternate
e. Ericaceae subfam. Harrimanelloideae
6. Shrubs (subshrubs or trees); ovaries superior; fruits, when dehiscent, usually septicidal, sometimes loculicidal or septifragal capsules; corollas sometimes persistent; seeds winged or not
d. Ericaceae subfam. Ericoideae
6. Vines, shrubs, or trees; ovaries inferior or superior; fruits, when dehiscent, loculicidal capsules; corollas deciduous; seeds usually not winged, sometimes slightly winged
f. Ericaceae subfam. Vaccinioideae

Key to Genera

1. Plants achlorophyllous; leaves reduced or absent
→ 2
1. Plants chlorophyllous; leaves usually present
→ 10
2. Flowers bilaterally symmetric.
Pyrola
2. Flowers radially symmetric
→ 3
3. Ovaries 1-locular; fruits baccate (fleshy); inflorescence axes not fibrous, not persistent
→ 4
3. Ovaries (4-)5(-6)-locular; fruits capsular; inflorescence axes fibrous, persistent after seed dispersal
→ 7
4. Petals connate
→ 5
4. Petals distinct
→ 6
5. Inflorescence axes violet to purple; e United States
Monotropsis
5. Inflorescence axes pink to cream; Pacific Coast (British Columbia to California).
Hemitomes
6. Petals glabrate abaxially, pubescent adaxially; anthers horseshoe-shaped; stigmas umbilicate, subtended by ring of hairs.
Pityopus
6. Petals glabrous; anthers elongate; stigmas crownlike, without subtending ring of hairs.
Pleuricospora
7. Petals connate; fruits dehiscent acropetally or indehiscent to irregularly dehiscent; seeds ovoid
→ 8
7. Petals distinct; fruits dehiscent basipetally; seeds fusiform
→ 9
8. Inflorescence axes pink to reddish or brownish, 0.5-1.5 cm diam. at proximal flower; anthers with awns; fruits dehiscent; seeds winged (with broad, rounded, membranous wing attached at 1 end).
Pterospora
8. Inflorescence axes bright red to dark orange, 2-10 cm diam. at proximal flower; anthers without awns; fruits indehiscent to irregularly dehiscent; seeds not winged.
Sarcodes
9. Inflorescence axes white with red to maroon vertical stripes; pedicels erect; sepals usually absent, rarely 2-5; stamens exserted.
Allotropa
9. Inflorescence axes white or yellowish to orange or reddish; pedicels decurved to spreading at anthesis; sepals (3-)4-5(-6); stamens included.
Monotropa
10. Herbs or subshrubs; leaves basal (or appearing so), or cauline and alternate or pseudoverticillate; petals distinct
→ 11
10. Trees, shrubs, subshrubs, or vines; leaves cauline, usually alternate, rarely opposite, pseudoverticillate, spirally arranged, or whorled; petals usually connate, rarely distinct or absent
→ 14
11. Inflorescences solitary flowers or 2-7-flowered corymbs or subumbels; fruits with no cobwebby tissue exposed by splitting valves at dehiscence
→ 12
11. Inflorescences solitary flowers or 2-29-flowered racemes; fruits with cobwebby tissue exposed by splitting valves at dehiscence
→ 13
12. Inflorescences solitary flowers; filaments glabrous; styles exserted; stigmas 5-lobed.
Moneses
12. Inflorescences 2-7-flowered corymbs or subumbels, rarely solitary flowers; filaments ciliate or villous basally; styles included; stigmas entire or obscurely 5-ridged.
Chimaphila
13. Inflorescences symmetric, usually erect; petals without basal tubercles.
Pyrola
13. Inflorescences secund (becoming ± erect in fruit, often lax in bud or flower); petals with 2 inconspicuous, basal tubercles.
Orthilia
14. Ovaries inferior; fruits fleshy
→ 15
14. Ovaries superior; fruits fleshy or not
→ 16
15. Fruits baccate; seeds 2-40, testa thin.
Vaccinium
15. Fruits drupaceous; seeds 10, testa bony (pyrenes).
Gaylussacia
16. Petals distinct or connate to 1/4 their lengths
→ 17
16. Petals connate 1/3+ their lengths (absent in Corema)
→ 22
17. Petals 7, covered by sticky exudate; (Alabama, Florida, Georgia).
Bejaria
17. Petals 2-5, not sticky
→ 18
18. Leaves whorled; fruits dry, red; pyrenes 2.
Ceratiola
18. Leaves alternate or opposite (sometimes whorled); fruits capsules, without pyrenes
→ 19
19. Sepals 3; fruits drupaceous.
Empetrum
19. Sepals (4-)5; fruits capsules
→ 20
20. Styles usually curved, sometimes straight; (Pacific Northwest coast, Georgia, South Carolina).
Elliottia
20. Styles straight (if bent or curved, then leaves alternate)
→ 21
21. Petals pink; flowers usually slightly bilaterally symmetric; leaves deciduous.
Rhododendron
21. Petals white or pink, flowers radially symmetric; leaves persistent (K. cuneata deciduous).
Kalmia
22. Flowers unisexual or bisexual
→ 23
22. Flowers bisexual
→ 24
23. Corollas ± salverform; leaves alternate; fruits capsular; inflorescences spikes or dense racemes.
Epigaea
23. Corollas inconspicuous or absent; leaves whorled or opposite (sometimes spiral); fruits drupaceous; inflorescences usually capitula, cymes, or fascicles, sometimes solitary flowers.
Corema
24. Fruits enclosed by fleshy calyx; leaves aromatic.
Gaultheria
24. Fruits enclosed by nonfleshy calyx; leaves not aromatic
→ 25
25. Corollas persistent; leaf blades 2.5-3.5 mm, base auriculate.
Calluna
25. Corollas deciduous; leaf blades 3-100+ mm, base cuneate, rounded, or obtuse (not auriculate)
→ 26
26. Fruits drupaceous or baccate
→ 27
26. Fruits capsular
→ 33
27. Leaves whorled, opposite, or spirally arranged, blade ± linear (rarely narrowly elliptic to ovate or lanceolate to linear or linear-oblong or very narrowly lanceolate); fruits drupaceous
→ 28
27. Leaves alternate (rarely opposite in Xylococcus), blade narrowly elliptic, elliptic, ovate, broadly ovate, obovate, or oblanceolate; fruits baccate or drupaceous
→ 29
28. Shrubs erect, ca. 2(-5) m; leaves whorled or opposite, blade 2.5-8 cm; inflorescences terminal panicles; fruits dry, brown or reddish brown; s California.
Ornithostaphylos
28. Shrubs prostrate, 0.5-1 m; leaves whorled or spirally arranged, blade 0.2-0.7 cm; inflorescences axillary, solitary flowers; fruits fleshy, black, purple, pink, or red; Greenland, Canada, n United States.
Empetrum
29. Fruits papillate or roughened and tuberculate, ± juicy
→ 30
29. Fruits smooth, dry, mealy, or juicy
→ 31
30. Leaf blades glabrous abaxially; fruits baccate, orange-red, red, or blackish red.
Arbutus
30. Leaf blades usually densely gray-tomentose abaxially; fruits drupaceous, red.
Comarostaphylis
31. Leaf margins strongly revolute; fruits dry, pyrenes connate.
Xylococcus
31. Leaf margins plane (rarely revolute); fruits fleshy (usually mealy, sometimes juicy), pyrenes usually distinct, sometimes some or all connate
→ 32
32. Petioles winged; leaf blade margins crenate to serrulate; shrubs; stems prostrate.
Arctous
32. Petioles not winged or absent; leaf blade margins entire or serrulate (sometimes ciliate); shrubs or trees; stems prostrate to erect.
Arctostaphylos
33. Leaves whorled; corollas persistent.
Erica
33. Leaves usually alternate or opposite (if whorled, corolla sympetalous and saucer-shaped); corollas deciduous
→ 34
34. Anthers not awned; fruit dehiscence septicidal or septifragal
→ 35
34. Anthers awned or not; fruit dehiscence loculicidal
→ 40
35. Corollas with pockets holding anthers until they open.
Kalmia
35. Corollas without pockets holding anthers
→ 36
36. Leaf blade margins appearing revolute (abaxial surfaces to 1/3 visible); older twigs glabrous or puberulent, roughened peglike projections remaining after fall of leaves
Phyllodoce
36. Leaf blade margins plane to revolute (abaxial surfaces 1/3+ visible except sometimes in bud); older twigs without peglike projections
→ 37
37. Corollas cylindric-urceolate, globose, or campanulate
→ 38
37. Corollas shallowly bell- to funnel-shaped, rotate, or campanulate
→ 39
38. Stems erect, spreading, or straggling; leaves deciduous; sepals connate ca. 3/4 their lengths; stamens 8.
Menziesia
38. Stems erect or trailing; leaves persistent; sepals nearly distinct; stamens 10; (sw Oregon).
Kalmiopsis
39. Inflorescence axes ± elongate; perulae leaflike proximally, green; (sw Alaska).
Therorhodion
39. Inflorescence axes ± short; perulae scalelike, brownish.
Rhododendron
40. Plants subshrubs; stems erect to decumbent or prostrate; leaf blades 2-6 mm; inflorescences solitary flowers; corollas campanulate
→ 41
40. Plants vines, shrubs, or trees; stems erect, (sometimes arching), ascending, or spreading; leaf blades 5-100+ mm; inflorescences usually racemes, panicles, fascicles, or corymbs, sometimes solitary flowers; corollas broadly campanulate to cylindric or urceolate
→ 42
41. Leaves opposite; inflorescences axillary; flowers pendulous.
Cassiope
41. Leaves alternate; inflorescences terminal; flowers erect to horizontal or nodding.
Harrimanella
42. Inflorescences racemes of (2-)5-12-flowered corymbs or solitary flowers, borne on leafless stems; corollas broadly campanulate
Zenobia
42. Inflorescences racemes, fascicles, panicles, or umbelliform corymbs, borne on leafy stems or twigs; corollas cylindric to urceolate
→ 43
43. Inflorescences panicles on shoots of current season, terminal; corollas densely unicellular-hairy.
Oxydendrum
43. Inflorescences racemes, fascicles, panicles, or umbelliform corymbs, terminal or axillary; corollas glabrous or with various multicellular and/or unicellular hairs, never densely unicellular-hairy
→ 44
44. Leaf blades with multicellular (stalked-glandular and/or elongate and eglandular) hairs, with or without unicellular hairs
→ 45
44. Leaf blades with unicellular hairs or peltate or lepidote scales, or glabrous
→ 49
45. Capsules with pale, decidedly thickened, whitish sutures; corollas sparsely stipitate-glandular.
Lyonia
45. Capsules without thickened sutures; corollas without multicellular hairs
→ 46
46. Pith chambered; leaf blade venation reticulodromous (reticulum rather dense and with all orders ± equally prominent).
Agarista
46. Pith solid; leaf blade venation brochidodromous to reticulodromous (of varied thickness and conspicuousness)
→ 47
47. Pedicels with 2 proximal bracteoles; anthers without awns or 2-awned.
Leucothoe
47. Pedicels with distal or medial bracteoles; anthers with or without awns
→ 48
48. Leaves persistent; pedicels with 2 medial bracteoles; anthers with 2 papillose, downward-pointing spurs at anther- filament junction.
Pieris
48. Leaves deciduous; pedicels with 2 distal bracteoles; anthers with 2 or 4 ± erect awns at anther apex.
Eubotrys
49. Leaf blades linear to narrowly elliptic or oblong, without peltate scales; stamens without awns.
Andromeda
49. Leaf blades elliptic, lanceolate, oblanceolate, oblong, or obovate, with silvery, brownish, stramineous, or ferrugineous scales (sometimes also hairy)
→ 50
50. Inflorescences fascicles, axillary, developing and blooming in spring, without leaflike bracts; corollas lepidote on outside; anthers without hollow "awns"; capsules with pale, thickened sutures.
Lyonia
50. Inflorescences racemes, terminal, developing in late summer and overwintering in exposed condition, with leaflike bracts; corolla glabrous on outside; anthers apically narrowed, forming hollow "awns" (tubules); capsules without thickened sutures.
Chamaedaphne
1. Leaves persistent, blade margins entire
→ 2
1. Leaves deciduous; leaf blade margins serrulate (serrate), or entire or undulate
→ 9
2. Leaf blade surfaces eglandular-hairy (hairs branched) and/or stipitate-glandular-hairy abaxially, hairs usually quickly deciduous to becoming matted and obscure, but not lepidote
→ 3
2. Leaf blade surfaces with glandular-peltate scales (lepidote) abaxially, scales sometimes obscured by various eglandular hairs
→ 5
3. Pedicels glabrous; petioles glabrous.
R. macrophyllum
3. Pedicels multicellular, stipitate-glandular- or eglandular-hairy; petioles multicellular-hairy (hairs ± branched), often glabrescent
→ 4
4. Calyx lobes 2-6 mm; pedicels and ovaries stipitate-glandular-hairy; leaf blades: apex acuminate to acute, length/width ratio (2.4-)2.6-5.5(-8), abaxial surface smooth to slightly roughened.
R. maximum
4. Calyx lobes 0.5-1.7 mm; pedicels and ovaries multicellular eglandular-hairy (i.e., hirsute); leaf blades: apex rounded/mucronate to obtuse or acute, length/width ratio (1.3-)1.5-3(-3.5); abaxial surface minutely and obscurely papillose.
R. catawbiense
5. Corollas white or pale to dark pink to rose or purple, petals connate 3/4+ their lengths; capsules basipetally dehiscent; leaf scales broad-rimmed
→ 6
5. Corollas white to cream, petals slightly connate basally; capsules acropetally dehiscent; leaf scales not broadly rimmed
→ 7
6. Shrubs to 3(-5) m; leaf blades (1-)5-8(-13) × (1-)2-3.5(-5.5) cm; corollas 15-37 mm, tube 8-22 mm, upper lobe usually green-spotted; stamens 10; capsules 6-14 mm.
R. minus
6. Shrubs 0.5(-0.7) m; leaf blades 0.4-2(-2.5) × 0.2-0.7(-0.9) cm; corollas (6.5-) 7.5-14(-15) mm, tube 1.5-6.5 mm, upper lobe not spotted; stamens 5-10; capsules 4-7 mm.
R. lapponicum
7. Twigs unicellular-hairy, papillate, and with flattened, glandular scales; petioles with glandular-peltate scales without broad rim and few ferruginous, long-crisped hairs; abaxial leaf surfaces sparsely to densely papillate, with few ferruginous, long-crisped hairs intermixed with conspicuous glandular- peltate scales.
R. columbianum
7. Twigs and petioles with ± dense covering of ferruginous, multicellular, long-crisped hairs; abaxial leaf surfaces (and especially midvein) with usually dense, ferruginous, multicellular, crisped hairs, these often intermixed with white, short, straight to curved hairs, glandular-peltate scales thus ± inconspicuous
→ 8
8. Pedicels with at least a few ferruginous, elongated, eglandular hairs, covered with ferruginous, long-crisped, unicellular and/or peltate scales, sometimes also with long-stalked, multicellular, glandular hairs; stamens 10; adaxial leaf surfaces with veins deeply impressed, abaxial with ferruginous, crisped hairs (usually more abundant on midrib, which is usually visible), sometimes forming dense, ± uniform mat; leaf blades ± linear (often much longer than wide); fruits borne on sharply recurved pedicels.
R. tomentosum
8. Pedicels with unicellular and/or glandular-peltate scales and sometimes ferruginous, long-crisped, multicellular hairs; stamens usually 5-10; adaxial leaf surfaces with veins usually not impressed, abaxial with ferruginous, crisped hairs forming usually dense, ± even covering, (usually concealing midvein); leaf blades ovate-lanceolate, sometimes narrowly elliptic to linear; fruits borne on broadly recurved pedicels.
R. groenlandicum
9. Inflorescences lateral, 1-2-flowered; flowers pendulous, ± radially symmetric.
R. albiflorum
9. Inflorescences terminal, 3-24-flowered; flowers erect to horizontal, bilaterally symmetric
→ 10
10. Corollas glabrous (or sparsely stipitate-glandular-hairy) on outside, tube to 1/4 as long as lobes or absent; stamens 5-7 or 10; seeds with testa tightly appressed
→ 11
10. Corollas often unicellular-hairy on outside, tube usually equaling or much longer than lobes; stamens 5; seeds with testa usually ± loose
→ 12
11. Corolla tubes absent due to deep division between 2 lower lobes and between lateral and lower lobes; upper corolla lobes 3-8 mm; stamens 10; filaments unicellular-hairy proximally; capsules moderately multicellular, stipitate-glandular- and eglandular-hairy; seed tails flattened; leaf margins usually revolute, sometimes plane, apex acute to rounded.
R. canadense
11. Corolla tubes 3-8 mm; upper corolla lobes 10-18 mm; stamens (5-)7; filaments glabrous; capsules very sparsely to moderately multicellular, stipitate-glandular-hairy; seed tails ± stellate-globular; leaf margins plane, apex acuminate.
R. vaseyi
12. Flowers opening after leaves (i.e., essentially all leaves unfolded, and vegetative bud scales absent)
→ 13
12. Flowers opening before or with (or sometimes right after in R. occidentale) leaves (i.e., at least some leaves still folded or vegetative bud scales still present)
→ 17
13. Twigs usually glabrous; abaxial surface of leaves glabrous
→ 14
13. Twigs multicellular-hairy (glandular or eglandular) and/or unicellular-hairy; abaxial leaf surface glabrous or sparsely or densely eglandular-hairy
→ 15
14. Corollas red to orange-red or orange; capsules eglandular-hairy and sparsely unicellular-hairy; seed testa expanded, dorsiventrally flattened.
R. prunifolium
14. Corollas white or sometimes light pink (filaments and style dark pink to red); capsules multicellular, stipitate-glandular-hairy, sometimes also sparsely unicellular-hairy; seed testa not dorsiventrally flattened.
R. arborescens
15. Corollas red, upper lobe with blotch or darker-colored spot, tube abruptly expanding into lobes; floral bud-scale margins glandular proximally (usually ciliate distally), abaxial surface glabrous.
R. cumberlandense
15. Corollas white (or sometimes pink), upper lobe with or without yellow or orange blotch, tube gradually expanding into lobes; floral bud-scale margins usually ciliate, glandular along proximal 2/3 or with glands and unicellular hairs mixed proximally, abaxial surface glabrous or densely unicellular-hairy
→ 16
16. Corollas densely glandular-hairy on outer surface (i.e., tube densely scattered, conspicuous, stipitate-glandular-hairy, hairs extending in line up corolla lobes), upper lobe without blotch; flowers strongly scented at night; styles white or greenish to pinkish.
R. viscosum
16. Corollas weakly glandular-hairy on outer surface, upper lobe with yellow to orange blotch; flowers strongly scented during mid day; styles white.
R. eastmanii
17. Upper corolla lobes with contrasting blotch often appearing as darker-colored area
→ 18
17. Upper corolla lobes without contrasting or darker-colored blotch
→ 22
18. Corolla lobes spreading nearly as broadly as tube is long, tube usually abruptly expanding into lobes, corolla yellow to orange, red-orange, or red
→ 19
18. Corolla lobes shorter than or equaling tube, tube gradually expanding into lobes, corolla white, sometimes pink tinged, or yellow to orange
→ 20
19. Floral bud scales with glandular margins, abaxial surface glabrous; corollas scattered, multicellular, stipitate-glandular-hairy on outside.
R. calendulaceum
19. Floral bud scales with ciliate margins, abaxial surface glabrous or densely unicellular-hairy; corolla tubes multicellular eglandular-hairy abaxially, sometimes very weakly glandular.
R. flammeum
20. Corollas yellow to orange (tube often orange-red or red), upper lobe with indistinct, darker yellow, orange, or red blotch; floral bud-scale margins glandular.
R. austrinum
20. Corollas white, sometimes pink tinged, upper lobe with contrasting yellow blotch; floral bud-scale margins unicellular, sometimes with unicellular hairs mixed with glands, or glandular
→ 21
21. Capsules moderately to densely unicellular-hairy and multicellular eglandular-hairy; floral bud scales glabrous or glabrate abaxially, margins unicellular-ciliate.
R. alabamense
21. Capsules sparsely unicellular-hairy (unicellular hairs rarely absent) and multicellular eglandular- and stipitate-glandular-hairy, or without eglandular hairs; floral bud scales sparsely to densely unicellular-hairy abaxially, margins unicellular-ciliate or with glands and cilia mixed, or with only glands.
R. occidentale
22. Corolla lobes multicellular stipitate-glandular-hairy (hairs forming lines that continue along outer surface), corolla white, usually flushed pink or rose; plants to 0.5-1 m, strongly rhizomatous
R. atlanticum
22. Corolla lobes scattered, multicellular stipitate-glandular- or eglandular-hairy on outer surface (hairs not forming distinct lines), corolla pink to white; plants to 3-6 m, not strongly rhizomatous
→ 23
23. Corollas pink, scattered, multicellular eglandular-hairy on outer surface; floral bud scales glabrous abaxially, margins moderately unicellular-ciliate; leaf blades glabrous or nearly so.
R. periclymenoides
23. Corollas pink or white with pink tube, multicellular stipitate-glandular-hairy; floral bud scales glabrous or sparsely to densely unicellular-hairy abaxially; leaf blades sparsely to densely unicellular-hairy
→ 24
24. Floral bud scales glabrous abaxially; pedicels (and sepal margins) usually eglandular, often not unicellular-hairy or only sparsely unicellular-hairy; leaf blades glabrous or only sparsely unicellular-hairy
R. periclymenoides
24. Floral bud scales sparsely to densely unicellular-hairy abaxially; pedicels (and sepal margins) eglandular or glandular, (sparsely) moderately to densely unicellular-hairy; leaf blades sparsely to densely unicellular-hairy abaxially
→ 25
25. Pedicels usually eglandular, 4-17 mm; leaf blade margins inconspicuously ciliate (cilia appressed to margins); capsules densely unicellular-hairy.
R. canescens
25. Pedicels usually glandular, 5-26 mm; leaf blade margins conspicuously ciliate (cilia ascending away from margins); capsules sparsely unicellular-hairy
R. prinophyllum
Source FNA vol. 8, p. 370. Author: Gordon C. Tucker. FNA vol. 8, p. 455. Authors: Walter S. Judd, Kathleen A. Kron.
Parent taxa Ericaceae > subfam. Ericoideae
Subordinate taxa
Agarista, Allotropa, Andromeda, Arbutus, Arctostaphylos, Arctous, Bejaria, Calluna, Cassiope, Ceratiola, Chamaedaphne, Chimaphila, Comarostaphylis, Corema, Elliottia, Empetrum, Epigaea, Erica, Eubotrys, Gaultheria, Gaylussacia, Harrimanella, Hemitomes, Kalmia, Kalmiopsis, Leucothoe, Lyonia, Menziesia, Moneses, Monotropa, Monotropsis, Ornithostaphylos, Orthilia, Oxydendrum, Phyllodoce, Pieris, Pityopus, Pleuricospora, Pterospora, Pyrola, Rhododendron, Sarcodes, Therorhodion, Vaccinium, Xylococcus, Zenobia
R. alabamense, R. albiflorum, R. arborescens, R. atlanticum, R. austrinum, R. calendulaceum, R. canadense, R. canescens, R. catawbiense, R. columbianum, R. cumberlandense, R. eastmanii, R. flammeum, R. groenlandicum, R. lapponicum, R. macrophyllum, R. maximum, R. minus, R. occidentale, R. periclymenoides, R. prinophyllum, R. prunifolium, R. tomentosum, R. vaseyi, R. viscosum
Name authority Jussieu Linnaeus: Sp. Pl. 1: 392. (1753): Gen. Pl. ed. 5, 185. 1754 ,
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