Veronica

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

Albach, D. C., P. Schönswetter, and A. Tribsch. 2006. Comparative phylogeography of the Veronica alpina complex in Europe and North America. Molec. Ecol. 15: 3269–3286.

Albach, D. C. and H. M. Meudt. 2010. Phylogeny of Veronica in the southern and northern hemispheres based on plastid, nuclear ribosomal and nuclear low-copy DNA. Molec. Phylogen. Evol. 54: 457–471.

Lower Taxa

Veronica agrestis, Veronica alpina, Veronica americana, Veronica anagallis-aquatica, Veronica argute-serrata, Veronica arvensis, Veronica beccabunga, Veronica biloba, Veronica catenata, Veronica chamaedrys, Veronica copelandii, Veronica cusickii, Veronica cymbalaria, Veronica dillenii, Veronica filiformis, Veronica fruticans, Veronica grandiflora, Veronica hederifolia, Veronica longifolia, Veronica officinalis, Veronica peregrina, Veronica persica, Veronica polita, Veronica scutellata, Veronica serpyllifolia, Veronica spicata, Veronica stelleri, Veronica sublobata, Veronica teucrium, Veronica triloba, Veronica triphyllos, Veronica undulata, Veronica verna, Veronica wormskjoldii

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