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sand dune wallflower, Sierra wallflower

wallflower

Habit Perennials or, rarely, biennials; (caudex slender). Plants not scapose; pubescent, trichomes sessile, medifixed, appressed, 2-rayed (malpighiaceous) or 3–5(–8)-rayed (stellate), rays (when 2) parallel to long axis of stems, leaves, sepals, and fruits.
Stems

erect, unbranched or branched (few to several) basally, 0.4–6.5 dm.

erect or ascending [decumbent], unbranched or branched basally and/or distally.

Leaves

basal and cauline; petiolate or sessile;

basal rosulate or not, petiolate, blade margins usually entire, dentate, sinuate-dentate, or denticulate, rarely pinnatifid or pinnatisect;

cauline petiolate or sessile, blade (base cuneate or attenuate [auriculate]), margins entire, dentate, denticulate, dentate-sinuate, or repand.

Basal leaves

blade spatulate to broadly oblanceolate, 2.5–7 cm × 3–10 mm, base attenuate, margins dentate or subentire, apex often obtuse, (surfaces pubescent adaxially, trichomes 2 or 3–5-rayed).

Cauline leaves

(distal) sessile;

blade margins often entire.

Racemes

considerably elongated in fruit.

(densely flowered, E. pallasii bracteate basally).

Flowers

sepals linear-oblong to oblong, 8–12 mm, lateral pair saccate basally;

petals yellow, broadly obovate to suborbicular, 15–22 × 3.5–6 mm, claw 8–14 mm, apex rounded;

median filaments 7–14 mm;

anthers linear, 3–4 mm.

sepals oblong or linear, lateral pair saccate or not basally (pubescent);

petals suborbicular, obovate, or spatulate, claw differentiated from blade (subequaling or longer than sepals, apex rounded [emarginate]);

stamens (erect), tetradynamous;

filaments not dilated basally;

anthers oblong or linear;

nectar glands (1, 2, or 4), distinct or confluent, subtending bases of stamens, median glands present or absent.

Fruiting pedicels

divaricate-ascending, slender, narrower than fruit, 4–12 mm.

erect, ascending, divaricate, reflexed, horizontal, or spreading, slender or stout (nearly as wide as fruit).

Fruits

erect to ascending, narrowly linear, straight, torulose, 3.8–14 cm × 1.2–3 mm, latiseptate, not striped;

valves with prominent midvein, pubescent outside, trichomes 2 or 3 (or 4)-rayed, glabrous inside;

ovules 26–44 per ovary;

style cylindrical, slender, (1.5–)2–5.5 mm, sparsely pubescent;

stigma subentire to slightly 2-lobed, lobes as long as wide.

usually sessile, rarely shortly stipitate (gynophore to 4 mm), usually linear or narrowly so [oblong], smooth or torulose, (keeled or not);

valves each with obscure to prominent midvein, pubescent outside, usually glabrous inside;

replum rounded;

septum complete, (not veined);

ovules [15–]20–120 per ovary; (style relatively short, rarely 1/2 as long as or subequaling fruit, often pubescent);

stigma capitate.

Seeds

ovoid, 2–3.4 × 1–2 mm;

not winged or, rarely, winged distally.

plump or flattened, winged, margined, or not winged, oblong, ovoid, obovate, or suborbicular;

seed coat (minutely reticulate), mucilaginous when wetted;

cotyledons incumbent, rarely accumbent.

Trichomes

of leaves 2–5-rayed.

x

= (6) 7, 8 (9–17).

2n

= 36.

Erysimum perenne

Erysimum

Phenology Flowering Jun–Sep.
Habitat Alpine fellfields, decomposing marble, gravelly ground and knolls, rocky slopes, talus, granitic sand
Elevation 2000-4000 m (6600-13100 ft)
Distribution
from FNA
CA; NV; OR
[WildflowerSearch map]
[BONAP county map]
from USDA
North America; n Mexico; Central America; Europe; Asia; n Africa; Atlantic Islands (Macaronesia) [Introduced in South America, Australia]
[BONAP county map]
Discussion

Erysimum perenne is a high alpine species of the western sierras in California from Fresno, Inyo, and Madera counties northward into Plumas, Siskiyou, and Trinity counties. Its range in Nevada appears to be restricted to Douglas and Washoe counties.

The limits of Erysimum perenne have been controversial, and it is with some hesitation that I recognize it as a species. G. B. Rossbach (1958) accepted it as a distinct species, R. A. Price (1993) transferred it (invalidly) to a subspecies of E. capitatum, R. C. Rollins (1993) treated it as a variety of E. capitatum, and N. H. Holmgren (2005b) treated the name as a synonym of E. capitatum. It is readily distinguished from E. capitatum by having torulose (versus not torulose) and flattened (versus 4-angled or flattened) fruits, slender (versus stout or, rarely, slender) and longer styles (1.5–)2–5.5 mm (versus 0.2–2.5(–3) mm), and yellow (versus orange to, rarely, yellow) petals. Where the two species are allopatric, they remain consistently distinct, but at lower elevations, where their ranges overlap, the distinction becomes blurred. In such areas of overlap, one finds fruit variation ranging from distinctly torulose to non-torulose, as well as continuity in the other characters above.

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

Species ca. 150 (19 in the flora).

Erysimum is found in the northern hemisphere, primarily Asia and Europe, with eight species in northern Africa and Macaronesia, and one each endemic to Baja California (E. moranii Rollins) and Costa Rica and Guatemala (E. ghiesbreghtii J. D. Smith). Of the 21 species found in North America, four are naturalized. Most of the native species have x = 9 and are believed to represent a monophyletic group (R. A. Price 1987).

Erysimum is a taxonomically difficult genus much in need of comprehensive phylogenetic and systematic studies covering its entire range. The principal sources of difficulty are the inflation in the number of species described, the heavy reliance on vegetative morphological characters in the delimitation of species, and the inadequacy of most herbarium specimens. In order to reliably identify a given sample, one often needs a complete specimen that has basal leaves, flowers, mature fruits, and seeds. Unfortunately, plants of most species shed their basal leaves or have no flowers when at full fruit maturity. Another complicating factor in North America is that almost all of the native species readily hybridize in areas of overlap to produce wide arrays of intermediates that backcross with the parents and blur species boundaries.

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

Key
1. Petals 3-10(-15) × 1.5-3 mm; median filaments 2-7(-10) mm
→ 2
1. Petals (10-)13-30(-35) × 3-10(-15) mm; median filaments (6-)7-15 mm
→ 6
2. Fruit valves densely pubescent inside; sepals 1.8-3.2 mm; petal claws 1.5-3.5 mm.
E. cheiranthoides
2. Fruit valves usually glabrous inside, rarely sparsely pubescent; sepals 4-7(-8) mm; petal claws 3-8 mm
→ 3
3. Annuals; fruiting pedicels as wide as fruit.
E. repandum
3. Biennials or perennials (short-lived); fruiting pedicels narrower than fruit
→ 4
4. Stigmas entire; fruits appressed to rachises; leaf blade surfaces and fruit valves with 3- and 4-rayed trichomes.
E. hieraciifolium
4. Stigmas strongly 2-lobed; fruits not appressed or subappressed to rachises; leaf blade surfaces and fruit valves with 2- or 3-rayed trichomes
→ 5
5. Petals narrowly obovate to spatulate, (8-)10-15 × (2-)2.5-4 mm; fruits 1.8-2.5 mm wide; seeds 1.5-2 mm.
E. coarctatum
5. Petals oblanceolate, 6-9(-11) × 1-2 mm; fruits 1.2-1.7 mm wide; seeds 1.2-1.7 × 0.8-1 mm.
E. inconspicuum
6. Subshrubs (stems woody at base)
→ 7
6. Biennials or perennials (stems not woody at base)
→ 10
7. Fruit valve trichomes 2-rayed; stigmas strongly 2-lobed, lobes much longer than wide; petals orange, yellow, brown, red, purple, violet, or white.
E. cheiri
7. Fruit valve trichomes 2-4-rayed; stigmas not strongly 2-lobed, lobes as long as wide; petals yellow or cream
→ 8
8. Fruits angustiseptate; leaf trichomes 2- or 3-rayed.
E. insulare
8. Fruits latispetate or 4-angled; leaf trichomes 2-5-rayed
→ 9
9. Distal cauline leaves petiolate; fruits latiseptate, not 4-angled; fruiting pedicels stout, 5-17(-22) mm.
E. franciscanum
9. Distal cauline leaves sessile; fruits slightly latiseptate or 4-angled; fruiting pedicels slender, (3-)5-10 mm.
E. suffrutescens
10. Petals purple or lilac; leaf trichomes 2-rayed; sepals 5-9 mm; anthers 1-1.5 mm.
E. pallasii
10. Petals usually yellow or orange, rarely lavender or purplish; leaf trichomes (at least some) 3-7-rayed; sepals 7-14 mm; anthers 2-4 mm
→ 11
11. Basal leaf blades filiform to narrowly linear, (somewhat revolute, appearing terete).
E. teretifolium
11. Basal leaf blades not filiform or narrowly linear
→ 12
12. Fruits 4-angled, longitudinally 4-striped; valves densely pubescent between midvein and replum with 2-rayed trichomes; ovules 72-120 per ovary.
E. asperum
12. Fruits latiseptate, rarely 4-angled, not longitudinally striped; valves pubescent with 2-6-rayed trichomes; ovules 24-86 per ovary
→ 13
13. Leaf blades: surfaces with 2 (or 3)-rayed trichomes
→ 14
13. Leaf blades: surfaces with 2-5(-7)-rayed trichomes
→ 16
14. Fruits usually 4-angled, rarely latiseptate; ovules (40-)54-82 per ovary; seeds not winged, 1.5-2(-2.4) mm.
E. capitatum
14. Fruits strongly latiseptate; ovules 24-46 per ovary; seeds usually broadly winged all around or apically, rarely not winged, 2-3.5 mm
→ 15
15. Perennials; seeds not winged or winged distally; fruits 1.5-2.7 mm wide.
E. arenicola
15. Biennials; seeds winged all around; fruits (2-)2.4-3.7 mm wide.
E. occidentale
16. Fruits torulose; ovules 26-44 per ovary; petals 3.5-6 mm wide.
E. perenne
16. Fruits not torulose; ovules (32-)42-86 per ovary; petals (5-)6-16 mm wide
→ 17
17. Fruiting pedicels 2-4(-6) mm; seeds broadly obovate to suborbicular, 1.5-3 mm wide.
E. concinnum
17. Fruiting pedicels 4-17(-25) mm; seeds oblong, 1-2 mm wide
→ 18
18. Fruits divaricate, ascending, or erect; petals orange or orange-yellow to yellow; seeds winged distally; Midwestern, Mountain, Pacific states.
E. capitatum
18. Fruits spreading; petals yellow; seeds winged all around; California (Humboldt, Mendocino, Monterrey, Santa Cruz counties)
→ 19
19. Basal leaf blades linear-oblanceolate, 2-9 mm wide; stems 0.4-9(-13) dm; fruit valves each with prominent midvein; ovules 50-86 per ovary.
E. ammophilum
19. Basal leaf blades spatulate, 5-15 mm wide; stems 0.2-2.5(-3.5) dm; fruit valves each with obscure midvein; ovules 32-74 per ovary.
E. menziesii
Source FNA vol. 7, p. 543. FNA vol. 7, p. 534. Author: Ihsan A. Al-Shehbaz.
Parent taxa Brassicaceae > tribe Erysimeae > Erysimum Brassicaceae > tribe Erysimeae
Sibling taxa
E. ammophilum, E. arenicola, E. asperum, E. capitatum, E. cheiranthoides, E. cheiri, E. coarctatum, E. concinnum, E. franciscanum, E. hieraciifolium, E. inconspicuum, E. insulare, E. menziesii, E. occidentale, E. pallasii, E. repandum, E. suffrutescens, E. teretifolium
Subordinate taxa
E. ammophilum, E. arenicola, E. asperum, E. capitatum, E. cheiranthoides, E. cheiri, E. coarctatum, E. concinnum, E. franciscanum, E. hieraciifolium, E. inconspicuum, E. insulare, E. menziesii, E. occidentale, E. pallasii, E. perenne, E. repandum, E. suffrutescens, E. teretifolium
Synonyms E. asperum var. perenne, Cheiranthus perennis, Cheirinia nevadensis, E. capitatum var. perenne, E. nevadense Cheiranthus, Cheirinia, Cuspidaria, Syrenia
Name authority (S. Watson ex Coville) Abrams: in L. Abrams and R. S. Ferris, Ill. Fl. Pacific States 2: 318. (1944) Linnaeus: Sp. Pl. 2: 660. (1753): Gen. Pl. ed. 5, 296. (1754)
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