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buttercup family, crowfoot family

aconit, aconite, monkshood, wolfsbane

Habit Herbs, sometimes woody or herbaceous climbers or low shrubs, perennial or annual, often rhizomatous. Herbs, perennial, from tubers or elongate, fascicled roots.
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 palmately divided into 3-7 segments, ultimate segments narrowly elliptic or lanceolate to linear, margins incised and toothed.

Inflorescences

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

terminal, sometimes also axillary, 1-32(-more) racemes or panicles, to 28 cm;

bracts leaflike, not forming involucre.

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, bilaterally symmetric;

sepals not persistent in fruit;

lower sepals (pendents) 2, plane, 6-20 mm;

lateral sepals 2, round-reniform;

upper sepal (hood) 1, saccate, arched, crescent-shaped or hemispheric to rounded-conic or tall and cylindric, usually beaked, 10-50 mm;

petals 2, distinct, bearing near apex a capitate to coiled spur, concealed in hood, long-clawed;

nectary present, on spur;

stamens 25-50;

filaments with base expanded;

staminodes absent between stamens and pistils;

pistils 3(-5), simple;

ovules 10-20 per pistil;

style present.

Fruits

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

follicles, aggregate, sessile, oblong, sides prominently transversely veined;

beak terminal, straight, 2-3 mm.

Seeds

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

endosperm abundant;

embryo usually small.

deltoid, usually with small, transverse, membranous lamellae.

x

=8.

Ranunculaceae

Aconitum

Distribution
Worldwide
[BONAP county map]
from USDA
Circumboreal; southward into n Mexico and n Africa
[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. 100 (5 in the flora).

The greatest concentration of species of Aconitum is in Asia, with a smaller group in Europe.

Aconitum is phylogenetically most closely related to Delphinium Linnaeus as evidenced by similarities in karyotype, production of diterpene alkaloids, and similarities in floral morphology. Distinctive and unique floral morphology clearly distinguishes Aconitum from all other genera.

The aconites have been of interest since ancient times because they contain diterpene alkaloids that range from relatively nontoxic to deadly poisonous. In various parts of the world they have been used medicinally and as a source of poisons throughout history (D. E. Brink 1982). Use of Aconitum alkaloids in modern medicine was largely discontinued by the late 1930s and early 1940s (E. E. Swanson et al. 1938; H. C. Wood and A. Osol 1943; A. Osol et al. 1960).

Aconitum is a circumboreal arctic and alpine genus that extends into lower latitudes where there is suitable mesic habitat at high elevations along the north-south chains of mountains in eastern and western North America, and also in outlying, scattered, mesic, interglacial refugia, occasionally at low elevations.

The genus Aconitum worldwide is notorious for complex patterns of morphologic intergradation that blur the lines between taxa. Aconites from different regions may be morphologically distinct but connected by a series of intermediate races. Aconitum columbianum exemplifies this in North America, and A. delphiniifolium may extend this complex of variation into Asia. Intergradation between A. columbianum and A. delphiniifolium should be more fully investigated.

Cultivated aconites with origins outside North America sometimes persist in old gardens or are encountered as garden escapes, especially in eastern Canada (New Brunswick, Newfoundland, Nova Scotia, Ontario, and Quebec). These may include Aconitum lycoctonum Linnaeus, A. napellus Linnaeus, A. variegatum Linnaeus, and A.bicolor Schultes. Aconitum lycoctonum is similar to A. reclinatum of the southeastern United States in having the tall, conic-cylindric hood that is characterisitc of species in Aconitum sect. Lycoctonum de Candolle. Aconitum reclinatum has white flowers whereas A. lycoctonum has lilac-purple flowers.

Aconitum napellus and A. variegatum are European introductions with leaves divided to the base as in A. delphiniifolium, which is native to Canada and Alaska. The introduced species have taller hoods and relatively short-petiolate leaves compared to A. delphiniifolium; in A. delphiniifolium petioles are longer, i.e., mostly as long as blades. Aconitum napellus has smooth seeds that lack the undulating membranous lamellae present in A. delpinifolium and A. variegatum.

Aconitum bicolor is a reputed hybrid between A. napellus and A. variegatum, having leaves like the former and flowers similar to the latter. It is always sterile; seeds are not viable. Flowers of A. bicolor are frequently white with purple margins.

A more complete treatment of the cultivated aconites likely to be encountered in North America can be found in H. J. Scoggan (1978-1979, part 3, pp. 718-720) and P. A. Munz (1945).

(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. Aerial stem arising from elongate, fascicled roots; hoods very tall, conic or nearly cylindric, flowers white to cream colored; e United States.
A. reclinatum
1. Aerial stem arising from tuber; hoods low, conic-hemispheric to crescent-shaped, flowers commonly blue to purple, occasionally white or yellowish; w Canada, e,w United States (including Alaska).
→ 2
2. Cauline leaf blades deeply divided to very base or with at most 2(–4) mm tissue between base and deepest sinus; hood low, crescent-shaped to conic-hemispheric; w Canada, Alaska.
A. delphiniifolium
2. Cauline leaf blades usually not divided to the very base, usually more than 2 mm between base and deepest sinus; hood conic-hemispheric, occasionally crescent-shaped; widespread.
→ 3
3. Tuber distally bulblike; terminal inflorescence often contracted, capitate; Alaska (Alaska Peninsula, and Aleutian Islands).
A. maximum
3. Tuber distally not obviously bulblike; inflorescence elongate, not capitate; sw Canada, United States except Alaska.
→ 4
4. Parent and daughter tubers separated by connecting rhizome 5–30 mm; eUnited States excluding Iowa, New York, and Wisconsin.
A. uncinatum
4. Parent and daughter tubers contiguous or nearly so; sw Canada, w United States, and Iowa, New York, Ohio, and Wisconsin.
A. columbianum
Source FNA vol. 3, p. 85. Authors: Alan T. Whittemore, Bruce D. Parfitt. FNA vol. 3. Authors: D. E. Brink, J. A. Woods.
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. columbianum, A. delphiniifolium, A. maximum, A. reclinatum, A. uncinatum
Name authority Jussieu Linnaeus: Sp. Pl. 1: 532. 175: Gen. Pl. ed. 5, 236. (1754)
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