Plantaginaceae

Herbs, subshrubs, or shrubs, annual or perennial, sometimes biennial, rarely fleshy (Plantago maritima), autotrophic. Stems prostrate, decumbent, ascending, or erect, sometimes arching, creeping, sprawling, spreading, scrambling, pendent, climbing, or reclining, sometimes absent. Leaves deciduous or persistent, basal, basal and cauline, or cauline, opposite, subopposite, alternate, helical, or whorled, sometimes opposite proximally, alternate distally, simple; stipules absent; petiole present or absent; blade fleshy or not, leathery or not, margins entire to subentire, toothed, or lobed. Inflorescences axillary or terminal, spikes, racemes, cymes, corymbs, thyrses, whorls, panicles, spiciform, or flowers 1–4. Flowers bisexual, rarely unisexual (Callitriche, Hippuris, Littorella), perianth and androecium hypogynous (epigynous in Hippuris); sepals (2–)4 or 5, basally or proximally connate, sometimes distinct, rarely a minute rim (Hippuris) or 0 (Callitriche), calyx radially or bilaterally symmetric; petals 0 (Callitriche, Hippuris) or (3 or)4 or 5, connate, corolla radially or bilaterally symmetric, ± bilabiate, bilabiate and personate, or regular, rarely unilabiate (Synthyris), ± rotate to salverform, ± cylindric, tubular, funnelform, urceolate, ellipsoid, globular, ovoid, or ligulate; stamens (1 or)2–4 or 5, adnate to corolla or free, didynamous or equal, staminodes 0 or 1(–3); pistil 1, 2-carpellate (1 in Hippuris), ovary superior (inferior in Hippuris), 2-locular, sometimes 1-locular or 4-locular (Callitriche), placentation axile, sometimes parietal (Dopatrium), parietal centrally and axile distally and proximally (Synthyris), apical (Hippuris), basal (Callitriche, Littorella), or free-central (Plantago); ovules anatropous, hemitropous, hemianatropous, or campylotropous, unitegmic, tenuinucellate; styles [0]1 (2 in Callitriche); stigma 1 (2 in Callitriche). Fruits capsules, dehiscence loculicidal, septicidal, or poricidal, or nutlets (Littorella), pyxides (Plantago), schizocarps (Callitriche), drupes (Hippuris), or drupelike (Lagotis). Seeds 1–300, white, tan, brown, yellow, gray, black, maroon, or red, ovoid to ellipsoid, cylindric, globular, oblong, conic, disciform, patelliform, or angled; embryo straight, rarely curved, endosperm present.

Distribution

nearly worldwide, apparently absent in much of tropical South America, and arid northern Africa and northern Australia.

Discussion

Genera ca. 100, species ca. 1900 (45 genera, 460 species in the flora).

The concept of Plantaginaceae has expanded considerably since the molecular study by R. G. Olmstead and P. A. Reeves (1995) reported that many of the genera at that time placed in the Scrophulariaceae did not cluster with Scrophularia and instead formed a clade with genera that usually had been considered to be distantly related and placed in different families, including Callitriche, Hippuris, and Plantago. As treated here, Plantaginaceae include Callitrichaceae Link and Hippuridaceae Vest. Subsequent studies (for example, D. C. Albach et al. 2005; B. Oxelman et al. 2005) have confirmed this basic alignment; as yet, there is no comprehensive phylogenetic classification of the family. Albach et al. suggested that the genera of Plantaginaceae might be placed in 12 tribes, noting molecular support for at least six of them. Species in the flora area are distributed among nine of the 12 tribes: Angelonieae Pennell (genus 1), Antirrhineae Dumortier (genera 2–18), Callitricheae Dumortier (genera 19 and 20), Cheloneae D. Don (genera 21–27), Digitalideae Dumortier (genus 28), Gratioleae D. Don (genera 29–38), Plantagineae Dumortier (genera 39 and 40), Russelieae Pennell (genus 41), and Veroniceae Duby (genera 42–45). Angelonieae, Digitalideae, and Russelieae are represented in the flora area only by introduced species. Within the family, genera are arranged alphabetically within tribes.

In the flora area, 20 of the 45 genera contain one species each; Penstemon contains nearly half of all North American Plantaginaceae species. Chelone, Chionophila, Holmgrenanthe, Howelliella, Nothochelone, Sophronanthe, and Tonella are endemic. Genera that include popular ornamental plants are Angelonia, Antirrhinum, Bacopa, Chelone, Digitalis, Linaria, Penstemon, Russelia, Veronica, and Veronicastrum. Ten genera are represented only by introduced species: Angelonia, Antirrhinum, Chaenorhinum, Cymbalaria, Digitalis, Kickxia, Limnophila, Linaria, Misopates, and Russelia. Linaria, Plantago, Scoparia, and Veronica contain globally important pest species, some of which are listed as noxious weeds in parts of the flora area.

Selected References

Albach, D. C. et al. 2004. Evolution of Veroniceae: A phylogenetic perspective. Ann. Missouri Bot. Gard. 91: 275–302.

Albach, D. C. et al. 2004b. A new classification of the tribe Veroniceae—Problems and a possible solution. Taxon 53: 429–452.

Albach, D. C., H. M. Meudt, and B. Oxelman. 2005. Piecing together the “new” Plantaginaceae. Amer. J. Bot. 92: 297–315.

Erbar, C. and P. Leins. 2004. Callitrichaceae. In: K. Kubitzki et al., eds. 1990+. The Families and Genera of Vascular Plants. 14+ vols. Berlin etc. Vol. 7, pp. 50–56.

Ghebrehiwet, M., B. Bremer, and M. Thulin. 2000. Phylogeny of the tribe Antirrhineae (Scrophulariaceae) based on morphological and ndhF sequence data. Pl. Syst. Evol. 220: 223–239.

Leins, P. and C. Erbar. 2004. Hippuridaceae. In: K. Kubitzki et al., eds. 1990+. The Families and Genera of Vascular Plants. 14+ vols. Berlin etc. Vol. 7, pp. 163–166.

Pennell, F. W. 1946. Reconsideration of the Bacopa-Herpestis problem of the Scrophulariaceae. Proc. Acad. Nat. Sci. Philadelphia 98: 83–98.

Sutton, D. A. 1988. A Revision of the Tribe Antirrhineae. London.

Wolfe, A. D. et al. 1997. Using restriction-site variation of PCR-amplified cpDNA genes for phylogenetic analysis of tribe Cheloneae (Scrophulariaceae). Amer. J. Bot. 84: 555–564.

Wolfe, A. D. et al. 2002. A phylogenetic and biogeographic analysis of the Cheloneae (Scrophulariaceae) based on ITS and matK sequence data. Syst. Bot. 27: 138–148.

Lower Taxa

Angelonia, Antirrhinum, Bacopa, Callitriche, Chaenorhinum, Chelone, Chionophila, Collinsia, Cymbalaria, Digitalis, Dopatrium, Epixiphium, Gambelia, Gratiola, Hippuris, Holmgrenanthe, Howelliella, Keckiella, Kickxia, Lagotis, Leucospora, Limnophila, Linaria, Littorella, Mabrya, Maurandella, Mecardonia, Misopates, Mohavea, Neogaerrhinum, Nothochelone, Nuttallanthus, Penstemon, Plantago, Pseudorontium, Russelia, Sairocarpus, Schistophragma, Scoparia, Sophronanthe, Stemodia, Synthyris, Tonella, Veronica, Veronicastrum

Illustrations

	Family ⠉	Plate of taxon	Illustrator ⠉
	Tetrachondraceae Plantaginaceae Plantaginaceae	Polypremum procumbens Angelonia gardneri Antirrhinum majus	Barbara Alongi Barbara Alongi Barbara Alongi
	Plantaginaceae Plantaginaceae Plantaginaceae	Chaenorhinum minus subsp. minus Cymbalaria muralis subsp. muralis Epixiphium wislizeni	Yevonn Wilson-Ramsey Barbara Alongi Barbara Alongi
	Plantaginaceae Plantaginaceae Plantaginaceae	Gambelia speciosa Holmgrenanthe petrophila Howelliella ovata	Barbara Alongi Barbara Alongi Barbara Alongi
... further results

Key

1	Corolla lobes 0; stamens 1; sepals 0 or minute rims at summits of ovaries.	> 2
2	Leaves opposite; fruits schizocarps; ovaries superior; styles 2.	Callitriche
2	Leaves whorled; fruits drupes; ovaries inferior; styles 1.	Hippuris
1	Corolla lobes 3–5, rarely 0; stamens 2–4(or 5); sepals 2–5.	> 3
3	Corolla tubes spurred, ± saccate, or gibbous, or with rounded sacs at bases of median lobes.	> 4
4	Corolla tubes with rounded sacs at bases of median lobes.	Angelonia
4	Corolla tube bases spurred, ± saccate, or gibbous, sometimes obscurely so.	> 5
5	Corolla tube bases ± saccate or gibbous adaxially, sometimes obscurely so.	> 6
6	Corollas: middle lobes of abaxial lips folded lengthwise, enclosing stamens and styles; stamens included.	Collinsia
6	Corollas: middle lobes of abaxial lips not folded lengthwise, not enclosing stamens and styles; stamens exserted.	Tonella
5	Corolla tube bases spurred or gibbous abaxially.	> 7
7	Corolla tube bases spurred.	> 8
8	Inflorescences terminal, racemes.	> 9
9	Corolla abaxial lips as long as or slightly longer than adaxials; filaments usually hairy proximally; capsules 9–12 mm.	Linaria
9	Corolla abaxial lips much longer than adaxials; filaments glabrous; capsules 2–4.8 mm.	Nuttallanthus
8	Inflorescences axillary or terminal, flowers solitary.	> 10
10	Stems ascending or erect; leaf blades linear, lanceolate, or oblanceolate; calyx lobes glandular-pubescent.	> 11
11	Pedicels 5–20 mm.	Chaenorhinum
11	Pedicels 1–4 mm.	Sairocarpus
10	Stems pendent, decumbent, prostrate, or erect; leaf blades oblong-ovate, orbiculate, cordate, or reniform; calyx lobes glabrous or villous.	> 12
12	Perennials; stems glabrous; calyx lobes glabrous.	Cymbalaria
12	Annuals; stems villous to glandular-hairy; calyx lobes villous.	Kickxia
7	Corolla tube bases gibbous.	> 13
13	Inflorescences terminal, racemes.	> 14
14	Corollas strongly bilabiate, not personate.	Howelliella
14	Corollas bilabiate and personate.	> 15
15	Corollas 25–45 mm; capsules 10–15 mm.	Antirrhinum
15	Corollas 9–18 mm; capsules 4–11 mm.	> 16
16	Calyx lobes linear, usually longer than corolla tubes in flower; seed wings present.	Misopates
16	Calyx lobes ovate to lanceolate, shorter than or as long as corolla tubes in flower; seed wings absent.	Sairocarpus
13	Inflorescences axillary, flowers solitary.	> 17
17	Corollas bilabiate; stamens 2; staminodes 2 or 3.	Mohavea
17	Corollas bilabiate and personate; stamens 4; staminodes 0 or 1.	> 18
18	Locules unequal.	Sairocarpus
18	Locules equal.	> 19
19	Stems erect, ascending, or sprawling, glabrous or hairy; bracts absent; pedicels twining; pollen sacs 2.	Neogaerrhinum
19	Stems erect, glandular-hairy; bracts present; pedicels not twining, recurved in fruit; pollen sacs 1.	Pseudorontium
3	Corolla tubes not spurred, gibbous, or saccate.	> 20
20	Shrubs or subshrubs.	> 21
21	Stamens 2.	Veronica
21	Stamens 4.	> 22
22	Distal leaf blades needlelike or scalelike; capsules densely packed with white, membranous hairs; inflorescences axillary, cymes.	Russelia
22	Distal leaf blades not needlelike or scalelike; capsules not densely packed with white, membranous hairs; inflorescences terminal, panicles, thyrses, corymbs, racemes, or spikelike racemes.	> 23
23	Staminodes 0; bracteoles absent; leaves whorled.	Gambelia
23	Staminodes 1; bracteoles usually present; leaves opposite, subopposite, or whorled, distals rarely alternate.	> 24
24	Stamen filament bases eglandular-hairy; nectaries hypogynous discs.	Keckiella
24	Stamen filament bases glabrous or glandular-puberulent proximally, rarely pubescent distally; nectaries epistaminal.	Penstemon
20	Herbs.	> 25
25	Fruits pyxides or nutlets; leaves basal only, rarely cauline; corolla lobes 4; stamens free.	> 26
26	Flowers unisexual; fruits nutlets; leaf blade margins entire.	Littorella
26	Flowers bisexual; fruits pyxides; leaf blade margins entire or toothed.	Plantago
25	Fruits capsules, rarely drupelike; leaves basal and cauline or cauline only, rarely basal only; corolla lobes 0 or 3–5; stamens adnate to corolla, rarely inserted on receptacle.	> 27
27	Stamens 2 or 3.	> 28
28	Ovaries 1-locular; leaves basal or basal and cauline.	> 29
29	Leaf blade margins entire; petioles absent; sepals 5; inflorescences of solitary flowers; annuals.	Dopatrium
29	Leaf blade margins toothed or deeply incised to pinnatifid; petioles present; sepals 2–4(or 5); inflorescences racemes; perennials.	Synthyris
28	Ovaries 2-locular; leaves cauline, sometimes basal and cauline.	> 30
30	Sepals 2; corolla lobes 3; fruits drupelike.	Lagotis
30	Sepals 4 or 5; corolla lobes 4 or 5; fruits capsules.	> 31
31	Bracts absent; stems prostrate; corollas radially symmetric, rarely bilaterally symmetric, regular.	Bacopa
31	Bracts present; stems creeping to decumbent, ascending, or erect; corollas bilaterally symmetric, weakly bilabiate, bilabiate, or bilabiate and personate, sometimes regular.	> 32
32	Corolla lobes 5.	> 33
33	Leaf blades not leathery; pollen sacs perpendicular to filaments, connectives dilated.	Gratiola
33	Leaf blades leathery; pollen sacs parallel to filaments, connectives not dilated.	Sophronanthe
32	Corolla lobes 4.	> 34
34	Leaves opposite, distals sometimes alternate; stamen filaments glabrous.	Veronica
34	Leaves whorled, rarely opposite; stamen filaments hairy proximally.	Veronicastrum
27	Stamens 4(or 5).	> 35
35	Leaves alternate.	> 36
36	Staminodes 0; inflorescences terminal, racemes; bracts present.	Digitalis
36	Staminodes 1; inflorescences axillary, flowers solitary; bracts absent.	> 37
37	Leaf blade margins entire; corollas blue to violet, pink, or red.	> 38
38	Annuals; seed wings present.	Epixiphium
38	Perennials; seed wings absent.	Maurandella
37	Leaf blade margins dentate or spinulose; corollas pale yellow to yellow or ochroleucous.	> 39
39	Leaf blade margins spinulose; stems erect; ovaries 1-locular.	Holmgrenanthe
39	Leaf blade margins dentate; stems pendent; ovaries 2-locular.	Mabrya
35	Leaves opposite or whorled, distals sometimes alternate.	> 40
40	Plants paludal or aquatic; leaves dimorphic, margins of submerged leaves pinnatifid, margins of aerial leaves entire, serrate, or pinnatifid.	Limnophila
40	Plants terrestrial or, if paludal or aquatic, leaves monomorphic and margins of leaves rarely pinnatifid.	> 41
41	Corollas radially symmetric.	> 42
42	Corolla lobes 5, rarely 4, throats not densely pilose internally; bracteoles present.	Bacopa
42	Corolla lobes 4, throats densely pilose internally; bracteoles absent.	Scoparia
41	Corollas bilaterally symmetric, rarely nearly radially symmetric.	> 43
43	Seeds 2 or 4; stamens equal; annuals; stigmas linear.	Tonella
43	Seeds (2–)5–150; stamens didynamous, rarely equal; perennials, sometimes annuals; stigmas capitate, 2-lobed, or cuneate.	> 44
44	Staminodes 1.	> 45
45	Bracteoles absent; leaf blade margins entire; seed wings present.	Chionophila
45	Bracteoles present; leaf blade margins toothed or entire; seed wings absent or present.	> 46
46	Bracteoles ± as large as calyx lobes, nearly surrounding calyces of flowers they subtend.	Chelone
46	Bracteoles smaller than calyx lobes, not surrounding calyces of flowers they subtend.	> 47
47	Inflorescences spikes, racemes, or flowers solitary; calyces bilaterally symmetric; capsule dehiscence loculicidal.	Stemodia
47	Inflorescences cymes or thyrses; calyces radially symmetric or nearly so; capsule dehiscence septicidal.	> 48
48	Stamen filaments hirsute proximally, puberulent distally; seed wings present; nectaries hypogynous discs.	Nothochelone
48	Stamen filaments glabrous or glandular-puberulent proximally, rarely pubescent distally; seed wings absent, sometimes narrow; nectaries epistaminal.	Penstemon
44	Staminodes 0.	> 49
49	Leaf blade margins pinnatifid to bipinnatifid; bracteoles absent.	> 50
50	Corollas pale blue to pale lavender; seeds white, parallel-ridged.	Leucospora
50	Corollas pink or purple; seeds yellow or brown, spirally ridged.	Schistophragma
49	Leaf blade margins entire, crenate, serrate, dentate, or denticulate; bracteoles present.	> 51
51	Leaf blade margins entire; sepals distinct, outer wider than inner.	Bacopa
51	Leaf blade margins toothed; sepals basally or proximally connate, outer calyx lobes ± as wide as inner.	> 52
52	Inflorescences spikelike cymes; bracteoles ± as large as calyx lobes, nearly surrounding calyces of flowers they subtend; calyces radially symmetric.	Chelone
52	Inflorescences spikes, racemes, or flowers solitary; bracteoles smaller than calyx lobes, not surrounding calyces of flowers they subtend; calyces bilaterally symmetric.	> 53
53	Stems glabrous; petioles present.	Mecardonia
53	Stems hairy; petioles absent.	Stemodia