Flora of North America species comparison

The green links below add additional plants to the comparison table. Blue links lead to other Web sites.
enable glossary links

Gratiola

Plantaginaceae

hedge-hyssop, hyssop

plantain family
Habit Herbs, annual or perennial. Herbs, subshrubs, or shrubs, annual or perennial, sometimes biennial, rarely fleshy (Plantago maritima), autotrophic.
Stems

decumbent to ascending or erect, glabrous or glandular-hairy.

prostrate, decumbent, ascending, or erect, sometimes arching, creeping, sprawling, spreading, scrambling, pendent, climbing, or reclining, sometimes absent.
Leaves

cauline (also basal and clustered in G. amphiantha), opposite, monomorphic (dimorphic in G. amphiantha);

petiole absent;

blade not fleshy, not leathery, margins entire or toothed, surfaces often glandular-punctate.

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, flowers 1 or 2 per node;

bracts present.

axillary or terminal, spikes, racemes, cymes, corymbs, thyrses, whorls, panicles, spiciform, or flowers 1–4.
Pedicels

present, sometimes short;

bracteoles present or absent, sepal-like.

Flowers

bisexual;

sepals 5, distinct or proximally connate, calyx ± bilaterally symmetric, campanulate, lobes (sepals when distinct) linear to lanceolate, obovate, or oblong;

corolla white, cream, or yellow, sometimes pinkish, tube yellowish, brownish, or purplish, often with dark veins, bilaterally symmetric, bilabiate, tubular, tube base not spurred or gibbous, lobes 5, abaxial 3, adaxial 2, nearly completely connate and appearing as 1 emarginate lobe, throat glabrous or pubescent;

stamens 2, medially adnate to corolla, filaments glabrous, pollen sacs perpendicular to filaments, connective dilated;

staminodes 0 or 2, minute;

ovary 2-locular, placentation axile;

stigma capitate or lamelliform.

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 septicidal, sometimes also loculicidal.

capsules, dehiscence loculicidal, septicidal, or poricidal, or nutlets (Littorella), pyxides (Plantago), schizocarps (Callitriche), drupes (Hippuris), or drupelike (Lagotis).
Seeds

ca. 200, yellow to brown or black, oblong to obpyramidal or short-cylindric, wings absent.

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.
× = 9.

Gratiola

Plantaginaceae
Distribution
from USDA
North America; Mexico; South America; Central America (Guatemala); Europe; e Asia (Japan); Africa (Morocco); Pacific Islands (New Zealand); Australia
[BONAP county map]
nearly worldwide; apparently absent in much of tropical South America; and arid northern Africa and northern Australia
[BONAP county map]
Discussion

Species ca. 30 (13 in the flora).

Members of Gratiola sometimes are confused with Lindernia and Mecardonia. Sophronanthe often is included in Gratiola; molecular and morphological data support separating the two as sister genera (D. Estes 2008).

This treatment was prepared in part from information contained in a partial manuscript submitted by D. Estes.

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

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.

(Discussion copyrighted by Flora of North America; reprinted with permission.)
Key
1. Capsules bilaterally symmetric; leaves dimorphic, submersed blades usually narrower than floating blades.
G. amphiantha
1. Capsules ± radially symmetric; leaves monomorphic, submersed blades absent or, if present, similar to emersed blades.
→ 2
2. Perennials.
→ 3
3. Bracteoles 0 or 1; leaf blades linear-lanceolate to lanceolate-ovate, 1–3 mm wide, margins entire or with 1 or 2(or 3) pairs of blunt teeth.
G. ramosa
3. Bracteoles (1 or)2; if 1, leaf blades lanceolate to lanceolate-ovate, ovate, or oblong, rarely linear-lanceolate, (1–)3–8(–13) mm wide, margins with (1 or)2–4 pairs of sharp teeth.
→ 4
4. Corolla limbs yellow, rarely white or cream, veins yellow to orangish yellow, rarely white; pedicels sparsely glandular-puberulent.
G. lutea
4. Corolla limbs white, veins yellow, brownish yellow, blue, or lavender; pedicels sparsely to densely glandular-puberulent to glandular-pubescent.
→ 5
5. Leaf blade margins with (1 or)2–4 pairs of teeth, blades lanceolate to lanceolate-ovate or oblong; corolla veins yellow or brownish yellow; sepals linear-lanceolate.
G. brevifolia
5. Leaf blade margins with (3–)5–10(–12) pairs of teeth, blades lanceolate-ovate to ovate or oblong, rarely linear-lanceolate in submersed forms; corolla veins blue or lavender; sepals lanceolate to elliptic or ovate-lanceolate.
G. viscidula
2. Annuals.
→ 6
6. Bracteoles 0; w North America.
→ 7
7. Calyces slightly bilaterally symmetric, sepals distinct, (4–)7–11 mm, lanceolate; leaf blade apices acuminate to attenuate.
G. ebracteata
7. Calyces distinctly bilaterally symmetric, sepals connate proximally, lobes 3.5–6 mm, elliptic-oblanceolate; leaf blade apices obtuse to rounded.
G. heterosepala
6. Bracteoles 1 or 2; e, c, or w North America.
→ 8
8. Corolla limbs yellow; capsules ovoid.
G. torreyi
8. Corolla limbs white, sometimes tinged pink, lavender, or purple; capsules ovoid to short-cylindric or subglobular.
→ 9
9. Pedicels stout, 1–12 mm, lengths 0.1–0.3 times bracts.
G. virginiana
9. Pedicels slender, (5–)7–45(–55) mm, lengths (0.3–)0.4–2(–2.3) times bracts.
→ 10
10. Corollas 14–25 mm; pedicels 20–45(–55) mm.
G. floridana
10. Corollas 7–12 mm; pedicels (5–)7–35 mm.
→ 11
11. Leaf blades 6–13(–18) mm; pedicel lengths 0.9–2(–2.3) times bracts; granite outcrops.
G. graniticola
11. Leaf blades 10–65 mm; pedicel lengths (0.3–)0.4–1(–1.6) times bracts; wet meadows, stream banks, shores of ponds, mudflats, salt marshes, crop fields, seeps, pools, and streams in limestone and dolomite glades, calcareous grasslands.
→ 12
12. Leaf blades linear to narrowly elliptic, oblanceolate, or elliptic-obovate, rarely falcate, 3–11(–18) mm wide, margins with (1 or)2–5(–7) pairs of blunt to sharp teeth, rarely entire; wet meadows, stream banks, shores of ponds, mudflats, salt marshes, crop fields.
G. neglecta
12. Leaf blades linear to linear-lanceolate or lanceolate-elliptic, often falcate, (1–)2.5–5 mm wide, margins entire or with 1 or 2(or 3) pairs of blunt teeth distally; seeps, pools, and streams in limestone and dolomite glades, calcareous grasslands.
G. quartermaniae
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
Source FNA vol. 17, p. 264. Author: Craig C. Freeman. FNA vol. 17, p. 11. Authors: Craig C. Freeman, Richard K. Rabeler, Wayne J. Elisens.
Parent taxa Plantaginaceae
Subordinate taxa
G. amphiantha, G. brevifolia, G. ebracteata, G. floridana, G. graniticola, G. heterosepala, G. lutea, G. neglecta, G. quartermaniae, G. ramosa, G. torreyi, G. virginiana, G. viscidula
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
Synonyms Amphianthus
Name authority Linnaeus: Sp. Pl. 1: 17. (1753): Gen. Pl. ed. 5, 11. (1754) Jussieu
Web links 
Unless otherwise noted, all data comes from the online version of the Flora of North America, who have made most of their content available via a Creative Commons license (CC BY 4.0).



Type/paste one or more Latin names into the boxes above; matching taxa will show up below the box as you type. You can use the up/down arrow keys to move through the list.

(If the plant you're looking for doesn't show up in the auto-complete list, that means it either isn't described in Flora of North America yet, or it's listed under a synonym. And please note that Flora of North America is still missing several plant families, including Apiaceae, Boraginaceae, and Polemoniaceae.)

Choose a data source: Flora of North America, OregonFlora, UW Burke Herbarium

The species comparison tool is brought to you by:
Northwest Wildflowers