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Key to families | Table of families and genera

Specimen numbers are hyperlinked to records in the Consortium of California Herbaria data view where possible. Taxa are hyperlinked to entries in the Jepson Interchange via the "[Online Interchange]" link.

 

CASTILLEJA PAINTBRUSH, OWL'S-CLOVER

Margriet Wetherwax, T.I. Chuang & Lawrence R. Heckard


Annual to subshrub, green. Leaf: sessile, entire to dissected. Inflorescence: spike-like; bracts becoming shorter, wider, more lobed than leaves, mature tips generally cream to red or green. Flower: calyx unequally 4-lobed, colored like bract tips; corolla upper 2 lip lobes fused, beak-like, tip open, lower lip reduced, 3-toothed to -pouched; stamens 4, anther sacs 2, unequal; stigma entire to 2-lobed, generally exserted. Fruit: ± asymmetric. Seed: generally ± brown, attached at base; coat netted, net-like walls ladder-like or not.
± 200 species: especially western North America. (Domingo Castillejo, Spanish botanist, 1744–1793) [Tank & Olmstead 2008 Amer J Bot 95:608–625] Hybridization and polyploidy common. Biologically consistent taxa difficult to define. Castilleja chrymactis Pennell not in California, sole (1947) record a misidentified, incomplete specimen. {hort link}
Unabridged references: [Chuang & Heckard 1991 Syst Bot 16:644–666]

Key to Castilleja

1. Inflorescence generally green, white, yellow, ± red-gray, bright pink, or ± purple, but not bright red; corolla 1–3 cm, beak generally <= tube; generally bee-pollinated; lower corolla lip generally 3-pouched (except Castilleja grisea, Castilleja plagiotoma), teeth generally ± erect, white or green to purple-red

2. Perennial herb to subshrub

3. Herbage gray-green, except bract lobes and calyx white-woolly; calyx divided deeper on sides than abaxially and adaxially ..... C. plagiotoma (2)

3' Herbage, including bract lobes and calyx, gray-green to densely white-woolly; calyx divided deeper abaxially and adaxially than on sides, or divided ± equally

4. Calyx divided much deeper abaxially and adaxially than on sides

5. Hairs stellate; San Clemente Island ..... C. grisea (2)

5' Hairs simple or branched; Klamath Ranges, High Cascade Range, High Sierra Nevada, San Bernardino Mountains, Great Basin Floristic Province

6. Herbage glandular; corolla 16–20 mm, lower lip 5–7 mm, teeth ± green-white or ± purple, ± triangular; stem generally unbranched; bract lobes ± acute ..... C. lemmonii

6' Herbage stiff-hairy, nonglandular; corolla 13–16 mm, lower lip 2 mm, teeth dark green, incurved; stem much-branched; bract lobes truncate or rounded ..... C. praeterita

4' Calyx divided ± equally

7. Herbage ± white-woolly

8. Plant 8–30 cm; leaves 20–60 mm; hairs simple; Klamath Ranges, High North Coast Ranges, High Cascade Range, n High Sierra Nevada, Modoc Plateau ..... C. arachnoidea

8' Plant 8–15 cm; leaves 5–20 mm; hairs branched; Klamath Ranges ..... C. schizotricha

7' Herbage puberulent to coarsely hairy

9. Herbage ashy-puberulent; bract tips dusty red, ± purple, or green-yellow; hairs dense, partitioned, much-branched; seed coat walls irregularly netted — seed coat tight-fitting; San Bernardino Mountains ..... C. cinerea

9' Herbage not ashy-puberulent; bract tips generally yellow-green or ± purple; hairs ± stiff, ± unbranched; seed coat walls with ladder-like thickenings

10. Bract lobes acuminate, green-margined; seed coat loose-fitting ..... C. nana

10' Bract lobes generally truncate or rounded (acute), white-margined; seed coat tight-fitting ..... C. pilosa

2' Annual

11. Bracts green throughout

12. Leaves and bracts entire ..... C. campestris

13. Leaves and bracts linear, thin, flexible; bracts <= flowers; corolla light to bright yellow; lower anther sac 1/4–1/3 upper ..... subsp. campestris

13' Leaves and bracts lanceolate, thick, ± brittle; bracts > flowers; corolla generally yellow to orange; lower anther sac ± 1/2 upper ..... subsp. succulenta

12' Leaves entire or lobed; bracts lobed

14. Corolla beak densely white-hairy — San Bernardino Mountains, n&c Peninsular Ranges ..... C. lasiorhyncha

14' Corolla beak puberulent

15. Lower corolla lip pouches 2–4 mm wide, ± 2 mm deep; stigma included ..... C. tenuis

15' Lower corolla lip pouches 4–10 mm wide, 3–6 mm deep; stigma generally ± exserted

16. Corolla 13–22 mm, pouches 4–8 mm wide; Klamath Ranges, Outer North Coast Ranges, Inner North Coast Ranges, Cascade Range, Sierra Nevada, Modoc Plateau ..... C. lacera

16' Corolla 20–28 mm, pouches 8–10 mm wide; North Coast Ranges, Cascade Range Foothills, Sacramento Valley, Central Coast, San Francisco Bay Area ..... C. rubicundula

17. Corolla yellow ..... subsp. lithospermoides

17' Corolla white, turning ± pink ..... subsp. rubicundula

11' Bracts and calyx lobes tipped ± purple, ± yellow, or ± white

18. Corolla beak tip hooked, densely shaggy-hairy; filaments puberulent ..... C. exserta

19. Distal bracts generally 5–7 mm wide, lobes < 2 mm, tipped pale lavender; inflorescence appearing alternately light- and dark-banded — North Coast, n&c Central Coast ..... subsp. latifolia

19' Distal bracts generally < 5 mm wide, lobes > 2 mm; inflorescence not appearing alternately light- and dark-banded

20. Corolla white, pale yellow or rose to ± purple; widespread ..... subsp. exserta

20' Corolla bright rose-red except yellow- to orange-tipped pouch; w Mojave Desert ..... subsp. venusta

18' Corolla beak straight, puberulent; filaments glabrous

21. Inflorescence 10–20 mm wide; corolla linear, pouches ± 2 mm wide; stigma including

22. Bract lobes 3, tipped white or pale yellow; seed < 1 mm, coat shallowly netted ..... C. attenuata

22' Bract lobes 3–5, tipped pink or purple-red; seed 1–1.5 mm, coat ± deeply netted ..... C. brevistyla

21' Inflorescence 20–40 mm wide; corolla wider distally, pouches generally 3–7 mm wide; stigma generally ± exserted

23. Stem ± stiffly spreading-hairy, glandular; bract tips cream to pale yellow — Sierra Nevada Foothills ..... C. lineariloba

23' Stem glabrous to puberulent, not glandular; bract tips yellow (cream) to pink- or rose-purple

24. Branches generally many; leaf lanceolate to oblong, > 3 mm wide; seed coat tight-fitting ..... C. ambigua

25. Bracts tipped yellow ..... subsp. ambigua

25' Bracts tipped ± yellow-pink to rose-purple

26. Plant ± fleshy, branches 0–few from mid-stem; salt marshes ..... subsp. humboldtiensis

26' Plant not fleshy, much-branched from base; grassy coastal bluffs ..... subsp. insalutata

24' Branches 0 or few from mid-stem; leaf lance-linear, < 3 mm wide; seed coat loose-fitting ..... C. densiflora

27. Calyx 5–10 mm; lower corolla lip widened abruptly, pouches generally shorter than deep ..... subsp. gracilis

27' Calyx 8–20 mm; lower corolla lip widened gradually, pouches longer than deep

28. Inflorescence generally rose-purple (cream) ..... subsp. densiflora

28' Inflorescence cream to pale yellow ..... subsp. obispoensis

1' Inflorescence generally red, orange, or yellow; corolla 1.2–5 cm, beak often >= tube; generally hummingbird-pollinated; lower corolla lip of small, incurved, generally dark green, red, black, ± yellow, or ± purple teeth

29. Annual; stem ± simple; wet places — leaves and bracts entire, lance-linear ..... C. minor

30. Corolla 15–20(30) mm; plant shaggy-hairy, most hairs glandular ..... subsp. minor

30' Corolla 25–35 mm; plant puberulent, many hairs nonglandular ..... subsp. spiralis

29' Perennial herb to subshrub; stems generally branched; generally drier places (except Castilleja miniata)

31. Herbage gray-green to densely white-woolly, hairs branched

32. Herbage ± gray-green; calyx white-woolly, divided deeper on sides than abaxially and adaxially ..... C. plagiotoma (2)

32' Entire plant densely white-woolly or ash-gray; calyx sinuses deeper abaxially and adaxially than on sides

33. Leaves oblong to obovate, entire, tip rounded; bract tips rounded or 0–3-lobed, yellow(-green) — n Channel Islands ..... C. mollis

33' Leaves linear to lanceolate, distal sometimes 3-lobed, tip acute to obtuse; inflorescence red, orange, or yellow; bracts 0–7-lobed, tips colored like calyx

34. Herbage white-felty, with long, intertwined, ± branched hairs; leaves entire — n Channel Islands ..... C. hololeuca

34' Herbage white-woolly or ash-gray with shorter, much-branched or stellate hairs; upper leaves sometimes 3-lobed

35. Inflorescence pale yellow-green; plant ash-gray; hairs stellate; bracts 5–7-lobed — s Channel Islands ..... C. grisea (2)

35' Inflorescence bright red to bright yellow (yellow-green); plant white-woolly or ash-gray; hairs much-branched; bracts 0–5-lobed

36. Herbage ± white to ± gray; calyx entire or barely notched on sides ..... C. foliolosa

36' Herbage ash-gray or green; calyx divided 1/5–1/4 on sides

37. Calyx lobes ovate, obtuse; lower corolla lip dark green to ± black; leaf lobes 0–3; San Gabriel Mountains ..... C. gleasoni

37' Calyx lobes lanceolate, acute; lower corolla lip green; leaf lobes 0–5; Northwestern California, Cascade Range, n&c Sierra Nevada, Modoc Plateau ..... C. pruinosa (2)

31' Herbage not both with branched hairs and gray-green to white-woolly

38. Calyx divided ± 2/3 abaxially, 1/6–1/3 adaxially, lobes generally curved upward; corolla (including lower lip) generally curved out through abaxial calyx sinus

39. Herbage glabrous to slightly puberulent; leaves linear, margins rolled upward; bracts generally narrowly 3-lobed, < calyx; corolla beak sparsely puberulent; Cascade Range, e slope High Sierra Nevada, Transverse Ranges, Great Basin Floristic Province, Desert Mountains ..... C. linariifolia

39' Herbage puberulent to hairy; leaves lanceolate, not rolled upward; bracts generally entire, >= calyx; corolla beak densely shaggy-puberulent; mostly c&s California Floristic Province ..... C. subinclusa

40. Inflorescence 2-colored, red and yellow; corolla yellow-orange; calyx lobes clearly curved upward, sides ± entire to divided < 1/5 ..... subsp. franciscana

40' Inflorescence ± red throughout; corolla ± red; calyx lobes barely curved upward, sides divided 1/5–1/3 ..... subsp. subinclusa

38' Calyx divided generally 1/4–1/2 (except Castilleja miniata, Castilleja peirsonii) abaxially and adaxially, lobes not curved upward; corolla not curved out through abaxial side of calyx sinus

41. Hairs dense, branched (puberulent) ..... C. pruinosa (2)

41' Hairs generally unbranched (or ± 0), plant puberulent to long-soft-hairy or bristly (Castilleja affinis subsp. affinis occasionally with some branched hairs)

42. Plant glandular-puberulent to -sticky-hairy proximal to inflorescence

43. Calyx divided >= 1/2 abaxially; corolla beak ± 1/2 tube; stigma clearly 2-lobed ..... C. peirsonii

43' Calyx generally divided 1/4–1/2 abaxially; corolla beak 1–2 × tube; stigma generally entire or slightly 2-lobed

44. Branches generally crowded proximal to inflorescence; leaves generally crowded; seed coat deeply netted; coastal ..... C. wightii

44' Branches few proximal to inflorescence; leaves generally ± well spaced; seed coat shallowly netted; inland

45. Plant glandular but not sticky; leaves entire or shallowly lobed; calyx lobes obtuse to rounded or acute

46. Leaf lobes 0–7; calyx lobes obtuse; nw Klamath Ranges ..... C. brevilobata

46' Leaf lobes 0–3; calyx lobes acute; e San Bernardino Mountains ..... C. montigena

45' Plant densely glandular-sticky; leaves 0–3-lobed; calyx lobes generally acute ..... C. applegatei

47. Stem 10–25 cm; leaf lobes generally 3; calyx 13–15 mm, divided ± 1/4 on sides; red fir forest to alpine barrens — High Sierra Nevada, Great Basin Floristic Province ..... subsp. pallida

47' Stem 30–80 cm; most leaves entire; calyx 12–25 mm, generally divided <= 1/5 on sides; generally below subalpine

48. Calyx lobes obtuse ..... subsp. martinii

48' Calyx lobes acute

49. Calyx 12–18 mm ..... subsp. disticha

49' Calyx 16–25 mm ..... subsp. pinetorum

42' Plant generally not glandular proximal to inflorescence (except occasionally Castilleja affinis or Castilleja latifolia)

50. Leaves ± fleshy, oblong to rounded, lobes 0–3, truncate-rounded; bract lobes 0–3, truncate-rounded; hairs generally bristly or shaggy

51. Corolla 20–30 mm, beak 10–15 mm; c&s Central Coast ..... C. latifolia

51' Corolla 30–45 mm, beak 15–25 mm; c North Coast ..... C. mendocinensis

50' Leaves generally not fleshy, linear to lanceolate; leaves and bracts entire or lobed, tips acute to obtuse; generally puberulent or glabrous (bristly)

52. Calyx divided 1/4–1/3 abaxially and adaxially — dry sagebrush scrub, pinyon/juniper woodland ..... C. chromosa

52' Calyx divided 1/3–2/3 or more abaxially and adaxially

53. Herbage generally ± bristly-puberulent; leaves 0–5-lobed ..... C. affinis (in part)

54. Inflorescence generally bright red to orange-red (yellow), 30–50 mm wide ..... subsp. affinis

54' Inflorescence generally yellow (pink or ± red-orange), 15–25 mm wide — s Inner North Coast Ranges, San Francisco Bay Area, open serpentine slopes ..... subsp. neglecta

53' Herbage glabrous to sparsely puberulent, to long-soft-hairy distally; leaves generally entire

55. Bract and calyx lobes obtuse; seed coat deeply netted, tight-fitting; generally dry sea bluffs — n&c North Coast ..... C. affinis subsp. litoralis

55' Bract and calyx lobes acute; seed coat shallowly netted, loose-fitting; moist places ..... C. miniata

56. Stem slender; inflorescence ± pink to yellow-orange; serpentine bogs; nw Klamath Ranges ..... subsp. elata

56' Stem stout; inflorescence generally bright red to orange-red; widespread ..... subsp. miniata

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