Veronica
Sp. Pl. 1: 9. 1753.
Herbs or subshrubs (V. fruticans) [shrubs], perennial, biennial (V. triphyllos), or annual; rhizomes creeping or absent. Stems creeping to erect, glabrous or hairy, glandular or eglandular. Leaves cauline, rarely in basal rosettes, opposite, sometimes alternate distally, rarely proximally (V. fruticans); petiole absent or present; blade not fleshy, not leathery (except V. fruticans), margins entire, dentate, serrate, 3–5-pinnately lobed, 3–7 pinnatifid, or ± palmatifid. Inflorescences terminal and/or axillary, racemes; bracts present, sometimes leaflike. Pedicels absent or present; bracteoles absent. Flowers bisexual; sepals 4 (or 5), basally connate, calyx bilaterally symmetric, rotate to campanulate, lobes linear to suborbiculate or triangular-cordate; corolla light to deep blue with lighter or reddish center, violet, lavender, or purple to pink with darker center, or white, bilaterally symmetric, weakly bilabiate, rotate or short-tubular, sometimes campanulate, tube base not spurred or gibbous, lobes 4, abaxial 3, adaxial 1; stamens 2, basally adnate to corolla, filaments glabrous; staminode 0; ovary 2-locular, placentation axile; stigma pointed to capitate. Fruits capsules, dehiscence loculicidal. Seeds (1–) 4–140, yellow to brown, rarely blackish (V. triphyllos), planoconvex to urn-shaped, wings absent. × = 9.
Distribution
North America, Mexico, Central America, South America, Eurasia, Africa, Atlantic Islands, Pacific Islands, Australia
Discussion
Species ca. 450 (34 in the flora).
Veronica includes many horticultural and weedy plants. Introduction and subsequent naturalization of species is often a possibility. The classification by D. C. Albach et al. (2004b) recognized 13 subgenera within Veronica; this was reduced to 12 by P. J. Garnock-Jones et al. (2007). Of these 12, ten are represented in the flora area. Species in this treatment are ordered by their placement in these subgenera as follows: subg. Pseudolysimachium (W. D. J. Koch) Buchenau (species 1 and 2, type V. spicata), subg. Veronica Linnaeus (species 3–10, type V. officinalis), subg. Beccabunga (Hill) M. M. Martínez Ortega, Albach & M. A. Fischer (species 11–17, type V. beccabunga), subg. Pellidosperma (E. B. J. Lehman) M. M. Martínez Ortega, Albach & M. A. Fischer (species 18, type V. praecox Allioni), subg. Cochlidiosperma (Reichenbach) M. M. Martínez Ortega, Albach & M. A. Fischer (species 19–22, type V. hederifolia), subg. Chamaedrys (W. D. J. Koch) M. M. Martínez Ortega, Albach & M. A. Fischer (species 23–26, type V. chamaedrys), subg. Stenocarpon (Borissova) M. M. Martínez Ortega, Albach & M. A. Fischer (species 27, type V. ciliata Fischer), subg. Pocilla (Dumortier) M. M. Martínez Ortega, Albach & M. A. Fischer (species 28–33, type V. agrestis), and subg. Pentasepalae (Bentham) M. M. Martínez Ortega, Albach & M. A. Fischer (species 34, type V. teucrium).
One of the subgenera, subg. Synthyris (Bentham) M. M. Martínez Ortega, Albach & M. A. Fischer, includes species treated here as the genus Synthyris. Other taxa included in Veronica but sometimes recognized at the generic rank are Pseudolysimachion (W. D. J. Koch) Opiz (D. C. Albach 2008) and Hebe Commerson ex Jussieu (P. J. Garnock-Jones et al. 2007).
Subgenus Pseudoveronica J. B. Armstrong includes Veronica franciscana Eastwood [= Hebe franciscana (Eastwood) Souster] and V. speciosa R. Cunningham ex A. Cunningham [= H. speciosa (R. Cunningham ex A. Cunningham) Andersen], which have been introduced to California as ornamentals that might escape.
Characters known to vary in Veronica are the indument (to densely white tomentose in V. incana Linnaeus), leaf shape, and flower color. Among many of the creeping species of the genus (for example, V. agrestis, V. filiformis, V. hederifolia, V. persica), most leaflike organs on the stems are bracts. The genus also includes a number of ornamentally grown plants, most of them more or less closely related to V. teucrium. They differ mainly in habit and leaf shape and should key out close to V. chamaedrys, V. officinalis, or V. teucrium, but none of them have been reported to persist outside their native range.
Selected References
Lower Taxa
Key
| 1 | Racemes 1–25, axillary. | > 2 |
| 2 | Stems hairy, rarely glabrate or glabrous; leaf blades 1–3 times as long as wide, rarely to 4 times as long (V. teucrium); meadows, forests, and other dry habitats. | > 3 |
| 3 | Leaf blades elliptic-ovate, ovate, ovate-oblong, or obovate, bases cuneate to attenuate or obtuse. | > 4 |
| 4 | Pedicels 8–10 mm, longer than subtending bracts; arctic. | Veronica grandiflora |
| 4 | Pedicels (0.5–)1–2(–3) mm, shorter than subtending bracts; not arctic. | Veronica officinalis |
| 3 | Leaf blades ± ovate, ovate-orbiculate, or oblong-ovate, rarely lanceolate, bases truncate to ± cordate. | > 5 |
| 5 | Stems ascending, rarely erect, (7–)10–30(–50) cm, hairs along stem in 2 prominent lines; calyces 4-lobed; styles (2.5–)4–5 mm. | Veronica chamaedrys |
| 5 | Stems erect, rarely ascending, (15–)30–70(–100) cm, hairs evenly distributed around stem; calyces 5-lobed; styles 5–6 mm. | Veronica teucrium |
| 2 | Stems glabrous or, if hairy, then only distally; leaf blades 1.5–10 times as long as wide; wet meadows, marshes, or aquatic habitats. | > 6 |
| 6 | Petioles 2–6(–10) mm. | > 7 |
| 7 | Leaf blades widest near or distal to middle; styles (1.3–)1.5–3(–3.5) mm. | Veronica beccabunga |
| 7 | Leaf blades widest proximal to middle; styles (1.7–)2.5–3.5(–4) mm. | Veronica americana |
| 6 | Petioles 0(–5) mm or proximals to 8 mm and distals 0 mm. | > 8 |
| 8 | Capsules compressed in cross section, apices emarginate by 1/3 length; leaf blades narrowly lanceolate or linear, rarely elliptic. | Veronica scutellata |
| 8 | Capsules ± compressed or ± round in cross section, apices rounded or ± emarginate; leaf blades ovate-oblong, ovate, oblong, linear-lanceolate, lanceolate, elliptic, oblong-ovate, or oblong-lanceolate. | > 9 |
| 9 | Corollas white to pale pink; calyx lobe apices obtuse; stamens 5 mm; pedicels equal to ± shorter than subtending bract; leaf margins entire or subentire. | Veronica catenata |
| 9 | Corollas pale lilac to pale blue or lavender, rarely white; calyx lobe apices acute; stamens 2–3.5 mm; pedicels equal to longer than subtending bract; leaf margins crenate, serrulate, or denticulate to subentire, sometimes serrate or ± undulate (V. undulata). | > 10 |
| 10 | Pedicels suberect or arcuate-erect to subpatent; racemes 5–10 mm diam., axes glabrous, rarely glandular-hairy. | Veronica anagallis-aquatica |
| 10 | Pedicels patent; racemes 10–15 mm diam., axes sparsely glandular-hairy, rarely glabrous. | Veronica undulata |
| 1 | Racemes 1(–20), terminal, sometimes also with 1–4 axillary racemes, flowers sometimes appearing solitary. | > 11 |
| 11 | Perennials; leaf blades (5–)7–40(–150) mm (usually at least 3 mm longer than bracts); bracts 1–9(–15) mm. | > 12 |
| 12 | Corollas campanulate; racemes (50–)70–200(–300) mm, (50–)100–300-flowered. | > 13 |
| 13 | Stems usually with 10–20 nodes, (50–)70–120(–150) cm; leaf blade margins serrate to biserrate, apices acute. | Veronica longifolia |
| 13 | Stems usually with 4–10 nodes, 5–45(–60) cm; leaf blade margins shallowly crenate to shallowly serrate to subentire, apices obtuse to rounded. | Veronica spicata |
| 12 | Corollas rotate; racemes (± loose), 5–100(–130) mm, (1–)2–30(–60)-flowered. | > 14 |
| 14 | Capsules wider than long; stems with scattered eglandular hairs only, often also with glandular hairs, sometimes glabrate. | Veronica serpyllifolia |
| 14 | Capsules ca. as long as or longer than wide; stems eglandular- or glandular-hairy. | > 15 |
| 15 | Subshrubs; stems densely branched from woody base. | Veronica fruticans |
| 15 | Herbs; stems unbranched or sparsely branched. | > 16 |
| 16 | Stamens 1–2.3 mm; corollas 2.5–11 mm diam.; styles 0.5–4(–6) mm. | > 17 |
| 17 | Raceme axes eglandular-hairy or glabrate; pedicels 1.5–2(–4) mm; styles 0.5–1.5(–2) mm. | Veronica alpina |
| 17 | Raceme axes glandular-hairy; pedicels 2–10(–15) mm; styles 0.8–4(–6) mm. | Veronica wormskjoldii |
| 16 | Stamens 4–8 mm; corollas (8–)10–13 mm diam.; styles 3–10 mm. | > 18 |
| 18 | Leaf blade surfaces glabrous or glabrate. | Veronica cusickii |
| 18 | Leaf blade surfaces hairy. | > 19 |
| 19 | Leaf blade margins entire; stems glandular-hairy. | Veronica copelandii |
| 19 | Leaf blade margins dentate or serrate; stems eglandular-hairy. | Veronica stelleri |
| 11 | Mostly annuals (V. triphyllos sometimes biennial, V. filiformis perennial); leaf blades (2–)5–28(–35) mm; bracts 3–25 mm (if bracts less than 9 mm, leaf blades usually less than 10 mm). | > 20 |
| 20 | Pedicels 0–4(–5) mm. | > 21 |
| 21 | Corollas white or pale pink; leaf blades 3–10 times as long as wide, margins entire or dentate distally. | Veronica peregrina |
| 21 | Corollas ± blue; leaf blades 1–2.5 times as long as wide, margins crenate-serrate, pinnatifid, or subpalmatifid. | > 22 |
| 22 | Leaf margins crenate-serrate. | Veronica arvensis |
| 22 | Leaf margins, at least distal leaves, pinnatifid or ± palmatifid. | > 23 |
| 23 | Corollas 1.5–3 mm diam.; styles 0.2–0.6 mm, stigmas white. | Veronica verna |
| 23 | Corollas 4–6 mm diam.; styles 0.8–1.5 mm, stigmas violet. | Veronica dillenii |
| 20 | Pedicels (2–)4–30 mm. | > 24 |
| 24 | Leaf blade margins 3–5(–7)-palmatifid. | Veronica triphyllos |
| 24 | Leaf blade margins serrate, dentate, crenate-serrate, or 3–9-lobed. | > 25 |
| 25 | Styles (1.5–)2–4 mm; pedicels (12–)15–30(–38) mm; corollas ± blue, 8–14 mm diam. | > 26 |
| 26 | Capsules reticulate with prominent veins, sinus angle (80–)90–120(–150)°; pedicels (12–)15–27(–38) mm, length 1–2(–3) times subtending bracts. | Veronica persica |
| 26 | Capsules absent or almost smooth, sinus angle 50–90°; pedicels (15–)20–30 mm, length 2–5 times subtending bracts. | Veronica filiformis |
| 25 | Styles 0.2–2 mm; pedicels (2–)3–15(–30) mm; corollas ± blue, white, pale lilac, pinkish, or pale violet, 2–12 mm diam. | > 27 |
| 27 | Stems erect. | > 28 |
| 28 | Bract margins: proximals serrate, distals sometimes entire; leaf blades (6–)12–28(–35) mm; styles (0.7–)1–1.3(–1.7) mm. | Veronica argute-serrata |
| 28 | Bract margins entire; leaf blades (3–)4–12(–20) mm; styles 0.4–0.8(–2) mm. | Veronica biloba |
| 27 | Stems creeping, decumbent, or ascending, rarely erect in young plants (V. sublobata). | > 29 |
| 29 | Leaf blade margins dentate to serrate; capsules ± compressed or ± round in cross section, apices emarginate. | > 30 |
| 30 | Calyx lobes broadly ovate, 2.6–3.8 mm wide; capsules densely eglandular-hairy (hairs less than 0.1 mm), sometimes also glandular-puberulent (hairs 0.2–0.3 mm), rarely glabrous; corollas intense dark to bright blue, rarely pale lilac or white. | Veronica polita |
| 30 | Calyx lobes linear-lanceolate, 2.2–2.6 mm wide; capsules sparsely glandular-hairy; corollas white or pale pinkish or pale blue. | Veronica agrestis |
| 29 | Leaf blade margins 3–5(–9)-lobed; capsules ± round in cross section, apices ± emarginate. | > 31 |
| 31 | Corollas white; calyx lobes not ciliate. | Veronica cymbalaria |
| 31 | Corollas blue or pale violet to whitish; calyx lobes ciliate. | > 32 |
| 32 | Corollas deep blue with white center; calyx lobe abaxial surfaces puberulent, rarely glabrous; pedicel length 1–2 times calyx. | Veronica triloba |
| 32 | Corollas bright blue with bright white center or pale violet to whitish; calyx lobe abaxial surfaces sparsely hairy or glabrous; pedicel length 2–4(–6) times calyx. | > 33 |
| 33 | Corollas bright blue with bright white center, 5–7(–9) mm diam.; styles (0.6–)0.7–0.9(–1.1) mm; pedicel length 2–3 times calyx, eglandular-hairy in single line adaxially. | Veronica hederifolia |
| 33 | Corollas pale violet to whitish, 4–5(–6) mm diam.; styles 0.2–0.7 mm; pedicel length 3–4(–6) times calyx, densely eglandular-hairy adaxially. | Veronica sublobata |