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evergreen plantain

plantain

Habit Perennials, sometimes woody; roots taproots, slender. Herbs, annual or perennial, sometimes biennial [rarely suffrutescent or arborescent]; caudex usually present when perennial.
Stems

100–400 mm, freely branched.

present or absent, if present, erect, glabrous or hairy.

Leaves

cauline, opposite, 30–60 × 0.75–1 mm;

blade linear to linear-lanceolate, margins entire, veins inconspicuous, surfaces hairy.

usually basal, usually alternate, (cauline and opposite in P. afra, P. indica, P. sempervirens);

petiole absent or present;

blade fleshy, leathery or not, margins entire or toothed.

Scapes

30–80 mm, hairy.

erect or ascending, rarely decumbent (P. coronopus), surpassing leaves, sometimes slightly so (P. tweedyi) or not (P. major).

Inflorescences

axillary, spikes or spiciform, dull, sometimes shiny (P. canescens, P. lanceolata, P. media);

bracts present.

Spikes

greenish or brownish, 40–85 mm, densely flowered;

bracts broadly ovate, 5–6 mm, lengths equal to sepals.

Pedicels

absent or present;

bracteoles absent.

Flowers

sepals 5–6 mm;

corolla radially symmetric, lobes reflexed, 3–3.5 mm, base obtuse;

stamens 4.

bisexual;

sepals 3 or 4, nearly distinct (abaxials connate in P. lanceolata), oblong, calyx radially, rarely bilaterally, symmetric, cuplike;

corolla semitransparent, radially or weakly bilaterally symmetric, lateral lobes smaller, ± tubular to ± funnelform, tube base not spurred or gibbous, tube glabrous, rarely hairy (P. coronopus, P. maritima), lobes 4;

stamens 2 or 4, free, equal, filaments glabrous;

staminode 0;

ovary 2-locular, placentation free-central, sometimes axile;

stigma elongate.

Fruits

pyxides, lanceoloid, rarely ovoid (P. macrocarpa), dehiscence circumscissile (indehiscent or dehiscence not circumscissile in P. macrocarpa).

Seeds

1 or 2, 2 mm.

(1 or)2–35, black or brown, sometimes dark red (P. rhodosperma) or yellowish brown (P. virginica), oblong, wings absent.

× = 4, 5, 6.

2n

= 12.

Plantago sempervirens

Plantago

Phenology Flowering summer–fall.
Habitat Disturbed habitats.
Elevation 0–200 m. (0–700 ft.)
Distribution
from FNA
CA; s Europe [Introduced in North America]
from USDA
North America; Mexico; Central America; South America; Europe; Asia; Africa; Pacific Islands (New Zealand); Australia
[BONAP county map]
Discussion

Plantago sempervirens is known from San Diego County; one specimen (JEPS) was collected in 2008 in Torrey Pines State Reserve.

Plantago cynops Linnaeus (1762, not 1753), a rejected name, has been misapplied to P. sempervirens.

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

Species ca. 210 (32 in the flora).

Plantago lanceolata and P. major have become established on all continents except Antarctica. A specimen of P. asiatica Linnaeus (New York City, US 295731) is ambiguous as to locality, and there is no evidence that it is established outside of cultivation in the flora area. Among North American Plantago, several native species have been introduced to states or provinces outside their native range.

For species with bilaterally symmetric calyces, sepal lengths in the descriptions are for the adaxial sepals.

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

Key
1. Leaves cauline.
→ 2
2. Perennials, sometimes woody.
P. sempervirens
2. Annuals.
→ 3
3. Spikes glandular-hairy; all bracts similar.
P. afra
3. Spikes eglandular; proximal bracts strongly differing from distal bracts.
P. indica
1. Leaves basal.
→ 4
4. Leaf margins usually 1- or 2-pinnatifid; scapes decumbent, sometimes erect; corolla tubes hairy.
P. coronopus
4. Leaf margins entire, toothed, or lobed; scapes erect or ascending; corolla tubes glabrous, rarely hairy (P. maritima).
→ 5
5. Annuals; leaf blades linear, narrowly lanceolate, narrowly elliptic, or almost filiform; roots taproots.
→ 6
6. Seeds (3 or)4–25(–30); corollas radially symmetric, lobes 0.5–1 mm; leaf blade surfaces glabrous or hairy.
→ 7
7. Seeds 10–25(–30), 0.5–0.8 mm.
P. heterophylla
7. Seeds (3 or)4–9(–12), 0.8–2.5 mm.
→ 8
8. Corolla lobes spreading or reflexed, not forming a beak; seeds (3 or)4–9(–12), 1.5–2.5 mm.
P. elongata
8. Corolla lobes erect, forming a beak; seeds 4, 0.8–1.3 mm.
P. pusilla
6. Seeds 2; corollas bilaterally or radially symmetric, lobes 1.3–3.6 mm; leaf blade surfaces lanate, sericeous, or villous, rarely glabrate or glabrous.
→ 9
9. Spikes: flowers in spirals; scapes without antrorse hairs; bracts ovate or elliptic.
P. ovata
9. Spikes: flowers in whorls or pairs; scapes with some antrorse hairs; bracts ovate, triangular, or almost linear.
→ 10
10. Corollas radially symmetric, lobe bases obtuse or slightly cordate.
→ 11
11. Scapes with antrorse, long and short hairs; bract lengths 0.3–0.8 times sepals; corolla lobes 2–2.7 mm; California, Oregon.
P. erecta
11. Scapes with patent, long and antrorse, short hairs; bract lengths 0.6–2.2 times sepals; corolla lobes 3–3.6 mm; New Mexico, Texas.
P. helleri
10. Corollas bilaterally symmetric, lobe bases slightly to deeply cordate.
→ 12
12. Leaf blades: adaxial surfaces glabrous or sparsely villous, margins entire, rarely toothed; stems 10–40 mm.
→ 13
13. Bract lengths 2–12 times sepals; corolla lobes: adaxials 1.4–2.3 mm, laterals symmetric; flowering spring–fall.
P. aristata
13. Bract lengths 0.4–0.8 times sepals; corolla lobes: adaxials 2.4–3 mm, laterals asymmetric; flowering summer.
P. wrightiana
12. Leaf blades: adaxial surfaces sericeous or villous, rarely lanate, margins entire or toothed; stems 0–20 mm.
→ 14
14. Bracts ovate, lengths 0.4–0.7 times sepals; leaves 1.5–4 mm wide; anther connectives slightly elongated, apices obtuse.
P. argyrea
14. Bracts triangular or ovate, lengths 0.6–2 times sepals; leaves 1–4 or 4–10 mm wide; anther connectives elongated to significantly elongated, apices acute.
→ 15
15. Bract lengths 0.6–1.4 times sepals; corolla lobes 2.2–2.5 mm; leaves 4–10 mm wide, blade margins toothed, rarely entire; flowering spring.
P. hookeriana
15. Bract lengths 1–2 times sepals; corolla lobes 1.6–2.1 mm; leaves 1–4 mm wide, blade margins entire, rarely toothed; flowering early summer.
P. patagonica
5. Perennials or annuals; leaf blades ovate, elliptic, or lanceolate, sometimes cordate-ovate, lanceolate-spatulate, linear, oblanceolate, obovate, or oval; roots taproots or fibrous.
→ 16
16. Corolla lobes usually forming a beak, erect or patent; annuals or perennials (usually without caudex).
→ 17
17. Annuals; roots taproots.
→ 18
18. Seeds: adaxial faces flat; bracts triangular, 2–3.1 mm; sepals 1.8–2.8 mm.
P. firma
18. Seeds: adaxial faces concave; bracts ovate or triangular, 1.6–3.2 mm; sepals 1.5–3.6 mm.
→ 19
19. Sepals 2.7–3.6 mm, apices acuminate; bracts 2.5–3.2 mm, narrowly triangular or triangular; seeds dark red.
P. rhodosperma
19. Sepals 1.5–2.4 mm, apices obtuse; bracts 1.6–2.4 mm, narrowly ovate or ovate; seeds brown or yellowish brown.
P. virginica
17. Perennials; roots taproots or fibrous.
→ 20
20. Adaxial surfaces of leaves: hairs floccose, slender, 4–6 × 0.01–0.03 mm.
P. floccosa
20. Adaxial surfaces of leaves: hairs not floccose, less than 2 mm long, more than 0.03+ mm wide.
→ 21
21. Roots fibrous; sepals 2–2.5 mm.
P. australis
21. Roots taproots; sepals 2.6–3.1 mm.
P. subnuda
16. Corolla lobes not forming a beak, spreading or reflexed; perennials (sometimes with caudex), rarely annuals.
→ 22
22. Fruits ovoid, indehiscent or dehiscence not circumscissile.
P. macrocarpa
22. Fruits lanceoloid, dehiscence circumscissile.
→ 23
23. Leaf blades linear to lanceolate, veins not conspicuous; corolla tubes hairy.
P. maritima
23. Leaf blades lanceolate, linear, oblanceolate, oval, cordate-ovate, lanceolate-spatulate, ovate, or elliptic, veins conspicuous; corolla tubes glabrous.
→ 24
24. Spikes grayish, whitish, or yellowish, shiny, corolla lobes of neighboring flowers often overlapping.
→ 25
25. Sepals: adaxial 2 connate; scapes groove-angled.
P. lanceolata
25. Sepals: adaxial 2 nearly distinct; scapes not groove-angled.
→ 26
26. Leaves ascending, 6–20 mm wide, blades linear to lanceolate or oblanceolate, surfaces hairy (hairs 1 mm) or glabrate; seeds 3–7, 1–1.8 mm.
P. canescens
26. Leaves prostrate, sometimes ascending, 30–70 mm wide, blades elliptic to ovate, surfaces hairy (hairs 0.5 mm); seeds 2–4, 2 mm.
P. media
24. Spikes brownish or greenish, dull, corolla lobes of neighboring flowers not overlapping.
→ 27
27. Caudices absent.
→ 28
28. Fruits (2–)4–5 mm, dehiscing at middle; seeds 5–35, 0.5–1 mm; bracts 0.5–1 mm.
P. major
28. Fruits 4–6(–8) mm, dehiscing proximal to middle; seeds 4 or 5(–8), 1.5–2 mm; bracts 2 mm.
P. rugelii
27. Caudices well developed, conspicuous.
→ 29
29. Spikes densely flowered, rachises not clearly visible between flowers; scapes slightly surpassing leaves.
P. tweedyi
29. Spikes loosely flowered, rachises visible between flowers; scapes surpassing leaves.
→ 30
30. Leaf blades: lateral veins branching from midvein distal to base.
P. cordata
30. Leaf blades: lateral veins branching from base.
→ 31
31. Caudices brown-woolly; c, w United States and Canada.
P. eriopoda
31. Caudices glabrous; se United States.
P. sparsiflora
Source FNA vol. 17, p. 292. FNA vol. 17, p. 281. Author: Alexey Shipunov.
Parent taxa Plantaginaceae > Plantago Plantaginaceae
Sibling taxa
P. afra, P. argyrea, P. aristata, P. australis, P. canescens, P. cordata, P. coronopus, P. elongata, P. erecta, P. eriopoda, P. firma, P. floccosa, P. helleri, P. heterophylla, P. hookeriana, P. indica, P. lanceolata, P. macrocarpa, P. major, P. maritima, P. media, P. ovata, P. patagonica, P. pusilla, P. rhodosperma, P. rugelii, P. sparsiflora, P. subnuda, P. tweedyi, P. virginica, P. wrightiana
Subordinate taxa
P. afra, P. argyrea, P. aristata, P. australis, P. canescens, P. cordata, P. coronopus, P. elongata, P. erecta, P. eriopoda, P. firma, P. floccosa, P. helleri, P. heterophylla, P. hookeriana, P. indica, P. lanceolata, P. macrocarpa, P. major, P. maritima, P. media, P. ovata, P. patagonica, P. pusilla, P. rhodosperma, P. rugelii, P. sempervirens, P. sparsiflora, P. subnuda, P. tweedyi, P. virginica, P. wrightiana
Synonyms Psyllium
Name authority Crantz: Inst. Rei Herb. 2: 331. (1766) Linnaeus: Sp. Pl. 1: 112. (1753): Gen. Pl. ed. 5, 52. (1754)
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