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circaea, enchanter's nightshade, nightshade

Habit Herbs, perennial, caulescent, colonial; stolons numerous. Herbs glabrous or pubescent with at least a few hairs at nodes; stolons tipped by apical tuber or, more commonly, apex slightly dilated.
Stems

erect, unbranched or sparsely branched.

7–70 cm.

Leaves

cauline, opposite;

stipules present, soon deciduous;

petiolate;

blade margins dentate to prominently dentate.

petiole 0.8–5.5 cm;

blade narrowly to broadly ovate, 2–7(–11) × 1.3–4.2(–7) cm, margins denticulate to prom-inently serrate, base cordate, apex short-acuminate to acute.

Inflorescences

simple or branched racemes, terminal on main stem or also at apex of branches, erect.

1.5–18 cm, sparsely to densely stipitate glandular.

Flowers

bisexual, zygomorphic, buds erect;

floral tube inconspicuous, deciduous (with sepals, petals, and stamens) after anthesis, with a nectary wholly within and filling proximal portion of floral tube or elongated and projecting above opening of floral tube as a fleshy, cylindrical or ringlike disc;

sepals 2, reflexed to spreading;

petals 2, alternate sepals, white or pink, without spots, clawed, apex notched;

stamens 2, anthers basifixed, pollen shed singly;

ovary 1- or 2-locular, stigma bilobed or obpyramidal, surface wet, minutely papillate.

opening after elongation of axis, ± loosely spaced;

pedicels ascending to spreading at anthesis, 1.8–5.5 mm, with a minute, setaceous bracteole at base;

floral tube 0.4–1.2 mm, funnelform to narrowly so;

nectary projecting beyond opening of floral tube, 0.1–0.4 mm;

sepals pink or pale green with apex commonly purple tinged, narrowly oblong to oblong-obovate or ovate, 1.6–3.5 × 0.9–2 mm;

petals white or pink, narrowly obtriangular to very broadly obovate, base cuneate to rounded, 1–3.6 × (0.6–)1.5–3.6 mm;

apical notch (1/3–)1/2–3/4 length of petal;

filaments 1.3–3.7 mm, pollen highly sterile (less than 2% fertile);

style 2.7–4.7 mm.

Fruit

a capsule, spreading or slightly reflexed, globose to clavoid or obovoid, indehiscent, surface smooth or with prominent longitudinal grooves (sulci) and rounded ridges, burlike, with stiff, hooked hairs;

pedicellate, deciduous at maturity.

Capsules

sterile, usually aborting before maturity, rarely developing to 3 × 1.5 mm, clavoid to obovoid, somewhat rounded to smooth or with low ribs and shallow grooves, tapering to pedicel, 2-locular, often with one locule larger, each with infertile seed.

Seeds

1 or 2, ellipsoid, glabrous, without appendages.

xI> = 11.

2n

= 22 (9 bivalents plus a ring or chain of 4, or 11 bivalents at meiosis).

Circaea

Circaea ×sterilis

Phenology Flowering summer.
Habitat Moist places in deciduous, mixed, or coniferous forests, especially in naturally disturbed areas along streams.
Elevation 0–1000 m. (0–3300 ft.)
Distribution
from USDA
North America; Europe; Asia; n Africa
[BONAP county map]
from FNA
CT; MA; ME; MI; MN; NC; NH; NJ; NY; OH; PA; SD; VA; VT; WI; WV; NB; NS; ON; PE; QC
[BONAP county map]
Discussion

Species 8 (3, including 1 hybrid, in the flora).

Circaea occurs throughout the temperate and boreal northern hemisphere, but is most diverse in eastern Asia, where all but one species occur. Reproductive features include: self-compatible; flowers diurnal, outcrossing, and pollinated by syrphid flies and small bees, or, sometimes, autogamous. It is found in rich, moist soils in deciduous forests and thickets, forest margins, and in moss or soil in mixed, coniferous-broadleaved deciduous, boreal forests. Circaea alpina subsp. alpina and C. canadensis subsp. canadensis often grow in close proximity and hybridize in eastern North America to produce C. ×sterilis. The unilocular C. alpina, with petals less than 2 mm, is self-pollinating under adverse weather conditions, but outcrosses on warm, sunny days. Because of its shorter style and much smaller pollen grains, it is probably the pollen recipient during hybridization events. Artificial hybridization experiments in England using C. alpina as the pollen donor and C. lutetiana as the pollen recipient failed to result in offspring, although hybrids were easily produced in the other direction (P. M. Benoit 1966). Recent molecular phylogenetic analysis supported the separation of the C. canadensis complex into two species; C. alpina subsp. pacifica was found to be sister to the remainder of the genus rather than being nested with other members of C. alpina (Xie L. et al. 2009). Thus, despite the strong morphological similarities of taxa within the C. canadensis and C. alpina complexes, these North American taxa may be better treated as separate species. Further detailed molecular studies are underway to examine this in more detail (Xie et al., unpubl.).

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

Circaea ×sterilis is a sterile hybrid intermediate in morphology between C. alpina × C. canadensis. Although sexually sterile, like other hybrids in Circaea, it reproduces more vigorously vegetatively than either of the parents. Its occurrence in naturally disturbed places along streams no doubt results in portions of stolons being broken off and carried away during high water to establish new colonies some distance away; colonies often extend for great distances along streams. The hybrids persist many years and colonies discovered 100 or more years ago can still be found in the same locality, often in the absence of one or both parents. The hybrid also occurs outside the range of C. canadensis subsp. canadensis, most conspicuously on the Gaspé Peninsula, Quebec, and in northern New Brunswick, but also elsewhere. Those hybrids may indicate a once more northerly range for C. canadensis subsp. canadensis, or that seeds resulting from hybridization events can be transported outside the range of the parents by birds or mammals.

M. L. Fernald (1950) misapplied the name Circaea canadensis Hill to plants now known to be hybrids (C. ×sterilis) between C. canadensis and C. alpina in North America and placed C. intermedia Ehrhart, a name published later, in its synonymy. Circaea ×intermedia is the hybrid between the European C. lutetiana Linnaeus and C. alpina Linnaeus. Hill used the name C. canadensis for plants in North America, to distinguish them from the European C. lutetiana.

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

Key
1. Flowers opening before elongation of raceme axis, clustered and corymbiform at apex of raceme, on ascending to erect pedicels; capsules clavoid, without corky ribs or grooves; stolons terminated by a tuber.
C. alpina
1. Flowers opening after elongation of raceme axis, more or less loosely spaced, borne on spreading pedicels; capsules usually obovoid to pyriform or subglobose, rarely clavoid, with corky, thickened ribs with deep grooves, or fruit sterile and aborting shortly after anthesis; stolons without or with a terminal tuber.
→ 2
2. Ovaries all or nearly all developing to maturity; capsules with corky thickened ribs separated by deep grooves; pollen highly fertile (greater than 80%); stolons without a tuber.
C. canadensis
2. Ovaries aborting shortly after anthesis, very rarely a few persistent, but easily detached, after anthesis; capsules, when somewhat persistent, smooth or with only low ribs and with shallow grooves; pollen highly sterile (less than 2% fertile); stolons terminated by a tuber or, more commonly, apex sligltly dilated.
C. ×sterilis
Source FNA vol. 10. Author: David E. Boufford. FNA vol. 10.
Parent taxa Onagraceae > subfam. Onagroideae > tribe Circaeeae Onagraceae > subfam. Onagroideae > tribe Circaeeae > Circaea
Sibling taxa
C. alpina, C. canadensis
Subordinate taxa
C. alpina, C. canadensis, C. ×sterilis
Name authority Linnaeus: Sp. Pl. 1: 8. (1753): Gen. Pl. ed. 5, 10. (1754) Boufford: Harvard Pap. Bot. 9: 256. (2005)
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