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buttercup family, crowfoot family

anemone, anémone, windflower

Habit Herbs, sometimes woody or herbaceous climbers or low shrubs, perennial or annual, often rhizomatous. Herbs, perennial, from rhizomes, caudices, or tubers.
Stems

unarmed.

Leaves

blade undivided or more commonly divided or compound, base cordate, sometimes truncate or cuneate, margins entire, toothed, or incised;

venation pinnate or palmate.

blade lobed or parted or undivided, reniform to obtriangular or lanceolate, margins entire or variously toothed.

Inflorescences

terminal or axillary, racemes, cymes, umbels, panicles, or spikes, or flowers solitary, flowers pedicellate or sessile.

terminal, 2-9-flowered cymes or umbels, or flowers solitary, to 60 cm;

involucres present, often with primary involucres subtending inflorescences, and secondary and tertiary involucres subtending inflorescence branches or single flowers (primary, secondary, and tertiary involucres appearing to be in tiers), involucral bracts 2-7(-9), leaflike or sepaloid, distant from or close to flowers.

Flowers

bisexual, sometimes unisexual, inconspicuous or showy, radially or bilaterally symmetric;

sepaloid bracteoles absent;

perianth hypogynous;

sepals usually imbricate, 3-6(-20), distinct, often petaloid and colored, occasionally spurred;

petals 0-26, distinct (connate in Consolida), plane, cup-shaped, funnel-shaped, or spurred, conspicuous or greatly reduced;

nectary usually present, rarely absent;

stamens 5-many, distinct;

anthers dehiscing longitudinally;

staminodes absent (except in Aquilegia and Clematis);

pistils 1-many;

styles present or absent, often persistent in fruit as beak.

bisexual, radially symmetric;

sepals not persistent in fruit, 4-20(-27), white, purple, blue, green, yellow, pink, or red, plane, linear to oblong or ovate to obovate, 3.5-40 mm;

petals usually absent (present in A. patens), distinct, plane, obovate to elliptic, 1.5-2 mm;

nectary present;

stamens 10-200;

filaments filiform or somewhat broadened at base;

staminodes absent between stamens and pistils;

pistils many, simple;

ovule 1 per pistil;

style present.

Fruits

achenes, follicles, or rarely utricles, capsules, or berries, often aggregated into globose to cylindric heads.

achenes, aggregate, sessile or stalked, ovoid to obovoid, sides not veined;

beak (persistent style) present, sometimes rudimentary, terminal, straight or curved, to 40(-50) mm, sometimes plumose.

Seeds

1-many per ovary, never stalked, not arillate;

endosperm abundant;

embryo usually small.

x

=7 or 8.

Ranunculaceae

Anemone

Distribution
Worldwide
[BONAP county map]
from USDA
Nearly worldwide; primarily in cooler temperate and arctic regions
[BONAP county map]
Discussion

Genera ca. 60, species 1700 (22 genera, 284 species in the flora).

The flowers of many species of Ranunculaceae begin to open long before anthesis, while the floral organs are just partly expanded. Only mature flowers with open anthers should be used for determination of diagnostic characteristics (especially measurements).

The literature is inconsistent about the term for the whorl of organs between sepals and stamens; these may be conspicuous and petaloid, or reduced to stalked nectaries, or intermediate between the two states. They have been called petals, honey-leaves, or (when they are inconspicuous) staminodes or nectaries. We follow M. Tamura (1993) and treat as petals all organs between the sepals and stamens, except in Clematis and Aquilegia where they usually bear rudimentary anthers and clearly represent staminodes.

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

Species ca. 150 (25 in the flora).

The taxonomy of Anemone continues to be problematic. Anemone occidentalis and A. patens var. multifida (the first two taxa in this treatment) are frequently placed in the genus Pulsatilla Miller on the basis of the long plumose achene beaks, and A. acutiloba and A. americana (the last two taxa in this treatment) in the genus Hepatica Miller, primarily on the basis of the involucre immediately subtending the flower and the lobed, persistent leaves. Recent phylogenetic analyses of Anemone in the broad sense, however, indicate that both Pulsatilla and Hepatica should be subsumed within Anemone. While traditional morphologic characters are useful in distinguishing between Pulsatilla and Hepatica species, respectively, many other morphologic and molecular attributes are shared with Anemone, strongly suggesting that these genera should be united (S. B. Hoot et al. 1994). In addition, a number of genera that have been recognized primarily on a cytotaxonomic basis (e.g., Anemonastrum, Anemonidium, Anemonoides, and Jurtsevia) are reduced to synonymy here. Some North American species of Anemone are closely related to plants in Europe, Asia, and South America and continue to be recognized at different ranks. For example, Anemone patens Linnaeus var. multifida (a species included in this treatment) was called Pulsatilla multifida (Pritzel) Juzepczuk for the former Soviet Union by S. V. Juzepczuk (1970) and Pulsatilla patens (Linnaeus) Miller var. multifida (Pritzel) Li S.H. & Huang Y. H. for China by Wang W.-T. (1980). Moreover, interspecific hybridization among some sympatric or nearly sympatric North American species also contributes to the confusion (see N. L. Britton 1891; C. L. Hitchcock et al. 1955-1969, vol. 2; R. S. Mitchell and J. K. Dean 1982). Additional analyses (e.g., G. Boraiah and M. Heimburger 1964; M. Heimburger 1959; C. Joseph and M. Heimburger 1966; and C. S. Keener et al. 1995) may prove to be helpful in resolving the taxonomy within this morphologically diverse genus.

Anemone nemorosa Linnaeus, A. ranunculoides Linnaeus, and A. blanda Schott & Kotschy, all native to Europe, are cultivated and may persist in the flora. Although apparently they rarely become naturalized, A. nemorosa is established at two sites in Newfoundland and Quebec, and A. ranunculoides in Quebec. Both are close relatives of A. quinquefolia and its allies.

Anemone ranunculoides is the only species in North America combining yellow sepals with rhizomes and 1-2-ternate leaves. Anemone blanda will key to A. caroliniana or A. berlandieri in this treatment. It can be distinguished by its short-pilose achenes, in contrast to the densely woolly achenes of A. caroliniana and A. berlandieri. Anemone nemorosa will key to A. quinquefolia; it differs in having 6-8 sepals and brown or black (never white) rhizomes with a 3-5 mm diameter in contrast to the 5 sepals and white or black rhizomes with 1-3 mmdiameter of A. quinquefolia.

Protoanemonin, an irritating acrid oil, is an enzymatic breakdown product of the glycoside ranunculin and is found in many species of Anemone. While protoanemonin can cause severe topical and gastrointestinal irritation, it is unstable and changes into harmless anemonin when plants are dried (N. J. Turner and A. F. Szczawinski 1991).

A caudex, as the term is used here, is the "woody," perennating base of an aerial shoot (inflorescences and basal leaves). The word tuber refers to a swollen, more or less vertical underground stem. The aerial shoots arise from the apex of either of those persistent structures. Rhizome, as the term is used here, refers to an underground, usually horizontal stem (more or less vertical in Anemone piperi), that is nearly uniform in diameter (about 1-4 mm diam., depending on the species) along its length. Aerial shoots arise directly from nodes at or near the apex of the rhizome.

Many species of Anemone have only one type of underground stem. Some species, however, have both rhizomes and caudices. In such cases the aerial shoots arise from the apex of a caudex attached to the rhizome. Some other species sometimes have both tubers and rhizomes. In those, one or more horizontal rhizomes arise near the apex of the tuber; the aerial shoots arise from the apex of the tuber.

Proportions given in the key for the middle lobes of basal leaves are calculated as follows: measure length of lobe from apex to a line connecting bases of sinuses; and measure total length of blade from leaf apex to summit of petiole.

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

Key
1. Flowers bilaterally symmetric; sepals showy; petals smaller than sepals.
→ 2
1. Flowers radially symmetric; sepals showy or not; petals present or absent, smaller to larger than sepals.
→ 4
2. Upper (adaxial) sepal (hood) saccate or helmet-shaped; petals completely hidden by sepals.
Aconitum
2. Upper (adaxial) sepal spurred; petals at least partly exserted from calyx.
→ 3
3. Perennials; pistils 3(-5); petals 4, distinct.
Delphinium
3. Annuals; pistil 1; petals 2, connate.
Consolida
4. Fruits achenes or utricles; ovule 1 per pistil.
→ 5
4. Fruits follicles, capsules, or berries; ovules 2 or more per pistil (1 of 2 aborting in Xanthorhiza, leaving 1 seed at maturity).
→ 12
5. Sepals spurred; leaves all basal, blade linear or narrowly oblanceolate.
Myosurus
5. Sepals plane; leaves either not all basal, or blade not linear or narrowly oblanceolate.
→ 6
6. Leaves all cauline and opposite; stems ±woody, at least at base.
Clematis
6. Leaves cauline and alternate (rarely opposite), or basal, or plants with basal leaves and opposite or whorled involucral bracts; stems herbaceous.
→ 7
7. Plants with 1 or more pairs (opposite) or whorls of involucral bracts, these leaflike or calyxlike.
→ 8
7. Plants without involucral bracts (inconspicuous, linear-lanceolate involucral bracts in Trautvetteria), cauline leaves if present alternate (rarely a pair of opposite, unlobed leaves in Ranunculus sect. Flammula).
→ 9
8. Achenes with conspicuous veins or ribs on lateral surfaces; style absent.
T. thalictroides
8. Achenes without veins on lateral surfaces; style present.
Anemone
9. Petals absent; inflorescences panicles, racemes, or corymbs (umbels in Thalictrum thalictroides); filaments filiform or dilated distally.
→ 10
9. Petals present (rarely absent in Ranunculus pedatifidus); inflorescences simple or compound cymes or flowers solitary; filaments filiform.
→ 11
10. Leaves simple, blade lobed; flowers bisexual; inflorescences corymbs.
Trautvetteria
10. Leaves compound; flowers unisexual or bisexual; inflorescences panicles, racemes, corymbs, or umbels.
Thalictrum
11. Petals without nectaries; sepals 5(-8).
Adonis
11. Petals with basal nectaries; sepals 3-5(-6).
Ranunculus
12. Leaves dissected into linear, threadlike segments; pistils compound; fruits capsules.
Nigella
12. Leaves not dissected, if parted or compound the segments not linear; pistils simple; fruits aggregates of follicles or solitary or aggregate berries.
→ 13
13. Shrubs; beak of follicle lateral, strongly incurved against abaxial surface of follicle.
Xanthorhiza
13. Herbs; beak of follicle, if present, terminal or nearly so, straight or slightly curved, sometimes hooked at tip.
→ 14
14. Petals prominent, spurred.
Aquilegia
14. Petals if present inconspicuous, plane or funnel-shaped.
→ 15
15. Flowers 12-50, in racemes or racemelike panicles.
→ 16
15. Flowers 1-10, in leafy cymes or solitary.
→ 17
16. Pistils 1-8; fruits follicles, usually aggregate; petals 2-cleft or absent.
Cimicifuga
16. Pistil 1; fruits berries; petals unlobed.
Actaea
17. Leaves simple, blade often lobed 1/2-3/4 its length, margins entire, crenate, or toothed; petals absent.
→ 18
17. Leaves compound or divided to base; petals usually inconspicuous (absent in Enemion).
→ 19
18. Leaf blades unlobed, margins entire, dentate, or crenate; fruits follicles.
Caltha
18. Leaf blades lobed, margins serrate; fruits berries.
Hydrastis
19. Leaves ternately 1-2× compound.
→ 20
19. Leaves palmately or pedately compound or divided.
→ 21
20. Leaves all basal; leaf blade deeply divided, ternately or pinnately 1-2× compound; petals present.
Coptis
20. Leaves basal and cauline; leaf blade ternately 2× compound; petals absent.
Enemion
21. Leaf segments lobed, margins sharply toothed; sepals persistent in fruit.
Helleborus
21. Leaf segments cleft or parted, margins entire or toothed; sepals not persistent in fruit.
→ 22
22. Cauline leaves absent except for whorl of 3 involucral bracts immediately subtending flower; follicles stipitate.
Eranthis
22. Cauline leaves alternate, (0.8-)1 cm or more from flower, involucral whorl absent; follicles sessile.
Trollius
1. Achene beak 20 mm or more, plumose.
→ 2
1. Achene beak 6 mm or less, glabrous or pubescent, not plumose.
→ 3
2. Involucral bracts compound, ultimate segments of lateral leaflets 2-3 mm wide; leaflets of basal leaves pinnatifid to dissected, lateral leaflets 2×-parted; petals absent; sepals white, purple tinged (rarely abaxially blue proximally, white distally, and adaxially white).
A. occidentalis
2. Involucral bracts simple, each leaf deeply divided into 4-6 segments, segments 1-2(-3) mm wide; leaflets of basal leaves dichotomously dissected, lateral leaflets 3-4×-parted; petals present; sepals blue or purple (rarely nearly white).
A. patens
3. Basal leaves simple (often deeply divided).
→ 4
3. Basal leaves compound.
→ 7
4. Involucral bracts remotely subtending flowers, margins crenate or sharply and irregularly serrate, ±similar to basal leaves.
→ 5
4. Involucral bracts closely subtending flowers, margins entire, not resembling basal leaves.
→ 6
5. Achenes winged, beak straight or indistinct; tiers of involucral bracts usually 2.
A. canadensis
5. Achenes not winged, beak recurved; tier of involucral bracts 1.
A. richardsonii
6. Leaf lobes acute or acuminate, middle lobe 70-90% of total blade length; involucral bracts ±acute.
A. acutiloba
6. Leaf lobes rounded, middle lobe 50-70(-75%) of total blade length; involucral bracts obtuse.
A. americana
7. Involucral bracts of distalmost tier simple (sometimes deeply lobed and sessile, occasionally appearing compound in primary involucres of A.tuberosa and A.okennonii).
→ 8
7. Involucral bracts of distalmost tier compound (petiole sometimes short and flat).
→ 16
8. Achene body winged, glabrous; inflorescences umbels or flowers solitary.
A. narcissiflora
8. Achene body not winged, hairy; inflorescences cymes or flowers solitary (sometimes umbelliform in A.cylindrica).
→ 9
9. Involucral bracts in 2 or more tiers.
→ 10
9. Involucral bracts in 1 tier.
→ 12
10. Involucral bracts ±similar to basal leaves.
A. tuberosa
10. Involucral bracts dissimilar to basal leaves.
→ 11
11. Basal leaves usually ternate; sepals (8-)10-20, 2-3 mm wide.
A. edwardsiana
11. Basal leaves 2-3-ternate; sepals 7-11, 3-4.5 mm wide.
A. okennonii
12. Aerial shoots from tubers (tuber caudex-like in A.tuberosa), tubers bearing rhizomes in some species; stamens 70 or fewer; mostly s United States.
→ 13
12. Aerial shoots from rhizomes (from short caudices on rhizomes in A.parviflora); stamens 70 or more; nw United States and Canada.
→ 15
13. Sepals 8-10; aerial shoots from caudex-like tuber, rhizomes absent.
A. tuberosa
13. Sepals (7-)10-20(-30); aerial shoots from tubers, sometimes with rhizomes arising near apex of tubers (tubers rarely bearing rhizomes in Anemone berlandieri).
→ 14
14. All basal leaf blades lobed or dissected differently from involucral bracts; involucres borne above middle of scape at anthesis.
A. berlandieri
14. One or more basal leaf blades dissected similarly to involucral bracts; involucres borne below middle of scape at anthesis.
A. caroliniana
15. Leaflets of basal leaves (0.5-)0.7-1.8(-2.2) × 0.5-1.3 cm; sepals abaxially hairy; stamens 70-80; achenes ca. 1 mm wide, beak 1-2.5 mm.
A. parviflora
15. Leaflets of basal leaves (2.5-)3-5(-6) × (2-)2.5-3.5 cm; sepals abaxially glabrous; stamens 100-120; achenes 2-3 mm wide, beak ca. 0.5 mm.
A. deltoidea
16. Stamens 80 or more; involucral bracts 2-ternate; involucral bracts in 1 tier.
→ 17
16. Stamens 80(-90) or fewer; involucral bracts 1(-2)-ternate; involucral bracts in 1 or more tiers.
→ 18
17. Sepals white or adaxially white and abaxially bluish; stamens whitish; style white.
A. drummondii
17. Sepals uniformly dark blue to purple; stamens purple; style red.
A. multiceps
18. Aerial shoots from caudices or from caudices on rhizomes; involucral bracts in 1-2(-3) tiers.
→ 19
18. Aerial shoots always from rhizomes; tier of involucral bracts 1.
→ 21
19. Ultimate lobes of leaflets of involucral bracts 1.5-3(-4.3) mm wide or less.
A. multifida
19. Ultimate lobes of leaflets of involucral bracts (4-)6 mm wide or more.
→ 20
20. Heads of achenes usually spheric or oblong-ellipsoid; achene beak 1-1.5 mm; involucral bracts of primary involucre 3(-5).
A. virginiana
20. Heads of achenes cylindric; achene beak 0.5-1 mm; involucral bracts 3-7(-9).
A. cylindrica
21. Rhizomes mostly vertical.
A. piperi
21. Rhizomes mostly horizontal.
→ 22
22. Lateral leaflets of basal leaves and involucral bracts mostly 1×-lobed or -parted.
quinquefolia var. quinquefolia
22. Lateral leaflets of basal leaves and involucral bracts only occasionally lobed.
→ 23
23. Achene body 2.5-3 mm.
quinquefolia var. minima
23. Achene body 3-5 mm.
→ 24
24. Achene beak (0.5-)1-1.5 mm; sepals 20 mm or less; stamens 75 or fewer.
→ 25
24. Achene beak 0.5-1 mm; sepals 15 mm or less; stamens 40 or fewer.
→ 26
25. Leaflets of basal leaves 0.7-2.5(-3.5) cm wide; peduncle distally villous to pilose; w North America.
A. oregana
25. Leaflets of basal leaves 2.5-4(-6) cm wide; peduncle ±glabrous; e North America.
A. lancifolia
26. Sepals 7-15 × 4-8 mm; pedicel (0.5-)3-10 cm in fruit; achene beak 0.6-1 mm; stamens 25-40.
A. grayi
26. Sepals 3.5-8(-10) × 1.5-3(-3.5) mm; pedicel 1-3(-4) cm in fruit; achene beak ca. 0.5 mm; stamens 10-30(-35).
A. lyallii
Source FNA vol. 3, p. 85. Treatment authors: Alan T. Whittemore, Bruce D. Parfitt. FNA vol. 3. Treatment authors: Bryan E. Dutton, Carl S. Keener, Bruce A. Ford.
Parent taxa Ranunculaceae
Subordinate taxa
Aconitum, Actaea, Adonis, Anemone, Aquilegia, Caltha, Cimicifuga, Clematis, Consolida, Coptis, Delphinium, Enemion, Eranthis, Helleborus, Hydrastis, Myosurus, Nigella, Ranunculus, Thalictrum, T. thalictroides, Trautvetteria, Trollius, Xanthorhiza
A. acutiloba, A. americana, A. berlandieri, A. canadensis, A. caroliniana, A. cylindrica, A. deltoidea, A. drummondii, A. edwardsiana, A. grayi, A. lancifolia, A. lyallii, A. multiceps, A. multifida, A. narcissiflora, A. occidentalis, A. okennonii, A. oregana, A. parviflora, A. patens, A. piperi, A. quinquefolia var. minima, A. quinquefolia var. quinquefolia, A. richardsonii, A. tuberosa, A. virginiana
Synonyms Anemonastrum, Anemonidium, Anemonoides, Hepatica, Jurtsevia, Pulsatilla
Name authority Jussieu Linnaeus: Sp. Pl. 1: 538. 175: Gen. Pl. ed. 5, 241. (1754) — name conserved
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