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toadflax

Habit Herbs, perennial or annual; caudex woody or herbaceous, shoot buds arising from roots.
Stems

prostrate to erect, glabrous or glandular-pubescent in inflorescence.

Leaves

cauline, usually whorled proximally, alternate distally;

petiole absent;

blade sometimes ± fleshy, not leathery, margins entire.

Inflorescences

terminal, racemes, compact to elongate;

bracts present.

Pedicels

present;

bracteoles absent.

Flowers

bisexual;

sepals 5, proximally connate, calyx ± bilaterally symmetric to nearly radially symmetric, 5-lobed, lobes linear to linear-lanceolate, lanceolate, ovate-lanceolate, or ovate;

corolla violet, blue, purple, red, yellow, or white, bilaterally symmetric, bilabiate, subcylindric, tube base not gibbous, spurred abaxially, lobes 5, abaxial 2, adaxial 3, abaxial lip as long as or slightly longer than adaxial, adaxial erect-patent to patent, with palate ± occluding mouth of tube;

stamens 4, basally adnate to corolla, didynamous, filaments usually hairy proximally;

staminode 1, scalelike;

ovary 2-locular, placentation axile;

styles simple or 2-fid;

stigma capitate, clavate, linear, 2-lobed, or emarginate.

Fruits

capsules, 9–12 mm, dehiscence poricidal, each loculus wall later splitting into valves.

Seeds

3–122, gray, brown, or black, globular, reniform, trigonous, tetrahedral, or disciform, wings present or absent.

x

= 6.

Linaria

Distribution
from USDA
Europe (especially Mediterranean region); w Asia; n Africa [Introduced in North America; introduced also in temperate regions of s South America (Argentina), Pacific Islands (New Zealand), Australia]
[BONAP county map]
Discussion

Species 150 (10 in the flora).

In this treatment, the species are arranged following the infrageneric classification proposed by D. A. Sutton (1988), with 150 species in seven sections; our ten belong to four of these sections.

North American species of Linaria canadensis (Linnaeus) Dumont de Courset, L. floridana Chapman, and L. texana Scheele included in sect. Leptoplectron Pennell, an invalidly published sectional name, have been included in Nuttallanthus, which differs from Linaria in having the abaxial lip of the corolla greatly exceeding the adaxial lip, a weakly developed palate, and prismatic seeds usually with prominent longitudinal ridges. Phylogenetic studies of Linaria and Nuttallanthus based on nuclear ribosomal DNA ITS sequences (M. Fernández-Mazuecos et al. 2013) concluded that on the basis of morphological traits and phylogenetic results, it would probably be appropriate to circumscribe Nuttallanthus species as a section (sect. Lectoplectron) of Linaria.

Linaria species often are cultivated and some (such as L. bipartita, L. dalmatica, L. triornithophora, and L. vulgaris) become naturalized; how long they may persist is unknown.

Although Linaria supina (Linnaeus) Chazelles de Prizy subsp. supina (lesser butter-and-eggs) has been collected in several states (New Jersey, New York, and Pennsylvania), all of the collections were made before 1900, and L. supina has yet to become established in the flora area. It would key closest to L. vulgaris, having shorter fertile stems [5–24 cm versus (25–)35–90(–120) cm], smaller corollas (15–25 mm versus 27–33 mm), and globular (versus oblong-globular or ovoid) capsules. The report from Massachusetts by C. H. Knowlton and W. Deane (1923) was based on a cultivated plant (10 July 1863, Boott s.n., GH), and the report from California is an error (http://ucjeps.berkeley.edu/cgi-bin/get_IJM.pl?tid=11372).

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

Key
1. Styles 2-fid; stigmas with 2 discrete areas; seed wings absent; annuals [sect. Versicolores].
→ 2
2. Racemes: rachis glandular-hairy, hairs 0.1–0.7 mm; capsule loculi subequal to unequal.
L. maroccana
2. Racemes: rachis glabrous or sparsely glandular-hairy, hairs 0.1–0.4 mm; capsule loculi equal, sometimes subequal.
→ 3
3. Corollas purple, mauve, or violet, usually with yellow palate.
L. bipartita
3. Corollas yellow.
L. spartea
1. Styles simple; stigmas entire; seed wings present or absent; perennials (some plants may bloom first year).
→ 4
4. Seeds disciform, subglobular, elliptic-globular, or deltate-globular, with prominent encircling wing.
→ 5
5. Corollas 27–32(–33) mm, white, pale yellow, or bright yellow, sometimes with orange palate [sect. Vulgaris].
L. vulgaris
5. Corollas 35–57 mm, violet, blue-violet, reddish purple, or pink, rarely white, with yellow, white, or lilac palate [sect. Pelisserianae].
L. triornithophora
4. Seeds trigonous, subtrigonous, or ± tetrahedral, wings absent [sect. Speciosae].
→ 6
6. Spurs 2–5 mm; corollas white or pale lilac.
L. repens
6. Spurs 5–30 mm; corollas yellow or violet to purple.
→ 7
7. Corollas violet to purple, with yellow or lilac palate.
L. purpurea
7. Corollas pale to bright yellow.
→ 8
8. Corollas 16–20(–23) mm; spurs 7–9(–10) mm.
L. genistifolia
8. Corollas 27–52 mm; spurs 11–30 mm.
→ 9
9. Corollas: adaxial lip sinuses 2–3.5(–4) mm; calyx lobes (1.9–)3–9.5 × (0.9–)1–3.5 mm in flower.
L. dalmatica
9. Corollas: adaxial lip sinuses 4–6 mm; calyx lobes 7–14 × 3.5–7 mm in flower.
L. grandiflora
Source FNA vol. 17, p. 27. Author: Llorenç Sáez Goñalons.
Parent taxa Plantaginaceae
Subordinate taxa
L. bipartita, L. dalmatica, L. genistifolia, L. grandiflora, L. maroccana, L. purpurea, L. repens, L. spartea, L. triornithophora, L. vulgaris
Name authority Miller: Gard. Dict. Abr. ed. 4, vol. 2. (1754)
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