The Jepson Herbarium
		University of California, Berkeley

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Annual to perennial herb, generally erect. Stem: branches 0 or generally ascending to erect, hairy. Leaf: generally sessile; basal whorled, cauline generally alternate, reduced above (opposite below); generally strigose, rough-hairy, or bristly, largest bristles generally bulbous-based. Inflorescence: generally terminal, raceme- or generally spike-like cymes, in groups of 1–5 (> 5), generally coiled in bud, generally elongated in fruit; bracts generally 0. Flower: generally unscented, persistent or not; sepals fused at base; corolla tube generally 1–13 mm, limb 0.5–12 mm diam, generally white, appendages generally 5; anthers included; ovary generally 4-lobed. Fruit: pedicel 0 or < 12 mm in fruit; nutlets 1–4, generally gray to brown, smooth to granular, tubercled, or papillate, with abaxial, longitudinal ridge to not; margin rounded, sharp-edged, or a ± flat rim or wing; adaxially grooved above attachment scar, scar generally lateral, narrow, open to closed, raised or generally not, edges inrolled to sharp-angled, generally forked or flared open at base; central fruit axis ("axis") not reaching to extending beyond fruit.
± 200 species: western North America, western South America, northeastern Asia (1 sp.). (Greek: hidden flowers, from cleistogamous flowers of some South American species) Generally homostylous. The tissue between ovary lobes, interpreted as a receptacle and/or style (style sometimes 0, then stigma attached to top of receptacle), extends to various degrees in fruit, forming what is often called the gynobase (here "fruit axis"), to which the nutlets are laterally attached at maturity, leaving an attachment scar. Annual species without yellow corolla appendages generally self-pollinating; perennial herb species generally homostylous in California. Some species, e.g., Cryptantha angustifolia, Cryptantha ambigua, Cryptantha barbigera, Cryptantha mariposae, hybridize with co-occurring species. Observation of nutlets, hairs best at 10+× generally critical for identification. Corolla limb diam generally < at end of flower period, especially for annual species Cryptantha sobolifera Payson does not occur in California.
Unabridged references: [Johnston 1925 Contr Gray Herbarium 74:1–125; Higgins 1971 Brigham Young Sci Bull Biol Ser 13:1–63, 1979 Great Basin Naturalist 39:293–350; Simpson & Hasenstab 2009 Crossosoma 35:1–59]

Key to Cryptantha

1. Bracts present; generally annual, generally wider than tall, often rounded to cushion-like; root generally red-purple, staining

2. Perennial herb from a woody, branched caudex; root not red-purple ..... C. cinerea var. abortiva

2' Annual; root generally red-purple, staining

3. Calyx circumscissile below middle in fruit, upper tube, lobes falling as 1 unit, lower tube scarious, persistent, cup-like in fruit

4. Corolla limb 3.5–6 mm diam, appendages larger than small to minute, yellow ..... C. similis

4' Corolla limb generally 1–2 mm diam, appendages small to minute, white to pale yellow ..... C. circumscissa

5. Stem generally > 2 cm; calyx 3–4 mm in fruit; fruit axis ± = mature nutlets; nutlet beak 0; leaf margin rolled under ..... var. circumscissa

5' Stem 0.5–2 cm; calyx 2.5–3 mm in fruit; fruit axis < mature nutlets; nutlet strongly beaked; leaf margin ± flat ..... var. rosulata

3' Calyx not circumscissile in fruit ..... C. micrantha

6. Corolla limb 1.5–4 mm diam, appendages larger than minute, yellow ..... var. lepida

6' Corolla limb 0.5–1.2 mm diam, appendages minute, ± white ..... var. micrantha

1' Bracts generally 0; annual or perennial herb, generally taller than wide (rounded or cushion-like); root generally not red-purple

7. Biennial to perennial herb; leaves generally basal or tufted; nutlet wide-rounded to obtuse at tip; tip of attachment scar groove well below nutlet tip

8. Corolla tube > calyx; flowers heterostylous

9. Corolla yellow ..... C. confertiflora

9' Corolla white ..... C. flavoculata

8' Corolla tube <= calyx; flowers homostylous

10. Stem 1–2(3) cm; basal leaves < 1.5 cm ..... C. roosiorum

10' Stem generally > 2 cm; basal leaves generally > 1.5 cm

11. Nutlet abaxially ± smooth

12. Nutlet 4.5–6 mm, margin winged ..... C. crymophila

12' Nutlet generally <= 4.5 mm, margin a ± flat narrow rim or occasionally sharp-angled

13. Caudex generally woody, generally branched; basal leaves sparse-hairy ..... C. nubigena (2)

13' Caudex not woody, unbranched; basal leaves dense-tomentose — e Modoc Plateau ..... C. schoolcraftii (2)

11' Nutlet abaxially granular, tubercled, or wrinkled

14. Nutlet adaxially ± smooth, especially near attachment scar

15. Nutlet attachment scar edges gapped ..... C. subretusa

15' Nutlet attachment scar edges generally not gapped

16. Caudex often woody, generally branched; basal leaves green, sparse-soft-bristly to dense-rough-hairy ..... C. nubigena (2)

16' Caudex not woody, unbranched; basal leaves ± gray, dense-tomentose — e Modoc Plateau ..... C. schoolcraftii (2)

14' Nutlet adaxially granular, tubercled, or wrinkled

17. Pedicel in fruit 0–2(3) mm

18. Corolla limb 1–3 mm diam; 1 nutlet > 2–3 ..... C. racemosa (2)

18' Corolla limb 3–10 mm diam; nutlets ± same size

19. Corolla limb 7–10 mm diam; calyx in fruit 7.5–10 mm; nutlet attachment scar edges flat; e Warner Mountains ..... C. celosioides

19' Corolla limb 3–7 mm diam; calyx in fruit 3.5–8 mm; nutlet attachment scar edges raised; White and Inyo Mountains, n Desert Mountains ..... C. hoffmannii

17' Pedicel in fruit (2)3–12 mm

20. Corolla late-deciduous to persistent; nutlet attachment scar edges flat

21. Corolla cream, aging brown, late-deciduous; nutlet abaxial ridge low ..... C. humilis

21' Corolla white, aging white, persistent; nutlet abaxial ridge 0 ..... C. racemosa (2)

20' Corolla ± early-deciduous; nutlet attachment scar edges raised

22. Plant long-lived perennial herb; caudex woody; nutlet abaxial ridge rounded, ± indistinct near base ..... C. tumulosa

22' Plant biennial or short-lived perennial herb; caudex not woody; nutlet abaxial ridge sharp-angled ..... C. virginensis

7' Generally annual; leaves generally cauline; nutlet narrow-acute to acuminate at tip; tip of attachment scar groove ± to nutlet tip

23. Nutlets ± smooth

24. Nutlet asymmetric, attachment scar off-center

25. Nutlets 4, attachment scar strongly off-center, appearing marginal; inflorescence elongate spikes ..... C. affinis

25' Nutlet 1(2), attachment scar off-center; inflorescence spheric clusters ..... C. glomeriflora

24' Nutlet symmetric, attachment scar central

26. Calyx hairs curved or hooked and ± straight; nutlets 1(2)

27. Nutlet attachment scar edges widening into an open gap at base; fruit axis > 1/2 nutlet

28. Corolla limb 1–2 mm diam; calyx lobe midveins with many stout, hooked or curved bristles, occasionally basally encrusted with exudate; nutlet 2.3–2.8 mm ..... C. rostellata

28' Corolla limb 3–6 mm diam; calyx lobe midveins with appressed to ascending curved hairs, with hooked or curved bristles only at tip, not basally encrusted with exudate; nutlet 1.6–2 mm ..... C. spithamaea

27' Nutlet attachment scar edges generally abutted to overlapped at base; fruit axis < 1/2 nutlet

29. Nutlet lance-ovate, swollen at base, margin rounded, tip acuminate, long-beaked; corolla limb 1–5(6) mm diam; calyx hairs generally basally encrusted with exudate ..... C. flaccida

29' Nutlet ovate, not swollen at base, margin sharp-angled, tip ± acute, short-beaked; corolla limb 0.5–1(1.5) mm diam; calyx hairs not basally encrusted with exudate ..... C. sparsiflora

26' Calyx hairs not hooked, ± straight (wavy); nutlets 1–4

30. Calyx lobe tip bristles reflexed; corolla limb 0.5–1 mm diam — nutlets 1 or 2(4) ..... C. nemaclada

30' Calyx lobe tip bristles generally spreading or ascending; corolla limb 0.5–8 mm diam

31. Calyx lobes 6–10 mm in fruit; longest calyx lobe bristles 3–4 mm in fruit — nutlets 1 or 2 ..... C. ganderi (2)

31' Calyx lobes 1–6 mm in fruit; longest calyx lobe bristles 0 or < 3 mm in fruit

32. Bracts present

33. Stem prostrate to decumbent, ± green in fruit; nutlets (3)4, smooth ..... C. leiocarpa

33' Stem erect to ascending, red-brown in fruit; nutlet 1, smooth or 2, 1 smooth, 1 rough ..... C. maritima (2)

32' Bracts generally 0

34. Nutlet margin a ± flat narrow rim, especially near tip, to sharp-angled; nutlets 4(3)

35. Corolla limb (4)5–7.5 mm diam; nutlets ± ovate ..... C. mohavensis

35' Corolla limb 0.5–1.5 mm diam; nutlets ± lanceolate ..... C. watsonii

34' Nutlet margin generally rounded, occasionally sharp-angled; nutlets 1–4

36. Calyx (1)1.5–3 mm in fruit

37. Nutlet 1.3–1.6 mm, abaxially ± round ..... C. microstachys

37' Nutlet 1.5–2.1 mm, abaxially ± flat

38. Calyx obconic at base; corolla limb 1–2.5 mm diam; nutlets generally 1 ..... C. gracilis

38' Calyx rounded at base; corolla limb 3–6 mm diam; nutlets generally 2 ..... C. incana (2)

36' Calyx (2.5)3–6.5(8) mm in fruit

39. Nutlet abaxially ± flat ..... C. torreyana

40. Calyx 1–1.5(2) mm, (2.5)3–4.5 mm in fruit, lobe margin sparse-strigose; nutlet 1.3–1.5 mm ..... var. pumila

40' Calyx 2–2.5 mm, 4–6.5(8) mm in fruit, lobe margin dense-strigose; nutlet (1.6)1.7–2.2(2.5) mm ..... var. torreyana

39' Nutlet abaxially ± round

41. Nutlet 1(2), adaxially ± rounded to convex, basally ± round in ×-section ..... C. hispidula

41' Nutlets 1–4, adaxially ± flat, basally not round in ×-section

42. Corolla limb 0.5–1(1.5) mm diam; nutlets generally 4; stem often stiffly branched above; White and Inyo Mountains ..... C. fendleri

42' Corolla limb 1–8 mm diam; nutlets 1–4; stem 0–few-branched ± above middle or branched throughout; California Floristic Province

43. Stem 0–few branched ± above middle ..... C. dissita

43' Stem branched throughout

44. Calyx long-soft-hairy, midveins not thickened, not bristly; nutlets 1(2) ..... C. milobakeri

44' Calyx minute-hairy, midveins thickened, bristly; nutlets 1–4 ..... C. clevelandii

45. Pedicel 0–0.5 mm; calyx ± appressed to stem in fruit, narrow at base, lobes linear; corolla limb 1–2.5 mm diam; nutlets 1–2 ..... var. clevelandii

45' Pedicel 0.5–1 mm; calyx not appressed to stem in fruit, ± wide at base, lobes ovate; corolla limb (1.5)2–5 mm diam; nutlets (1–2)3–4 ..... var. florosa

23' Nutlets, or at least 1, rough

46. At least 1 nutlet with all or part of margin a ± flat rim (occasionally seeming sharp-angled) or wing

47. Pedicel 1–4(6) mm in fruit ..... C. holoptera

47' Pedicel generally 0–1(1.5) mm in fruit

48. Nutlets 1 (2), margin a ± flat inward-bent narrow rim to basally sharp-angled, white-grainy papillate to minute-spiny; calyx narrow-urn-shaped in fruit ..... C. utahensis

48' Nutlets generally 4(3), margin occasionally seeming sharp-angled, generally narrow- to wide-winged along entirety, not white-grainy papillate to minute-spiny; calyx wide-urn-shaped or ovoid in fruit

49. At least some nutlet margins with a strongly lobed, wide wing ..... C. pterocarya

50. Nutlets similar in 1 fruit, all winged ..... var. cycloptera

50' Nutlets dissimilar in 1 fruit, 3 winged, 1 not

51. Nutlet wing wide, generally > 0.5 mm ..... var. pterocarya

51' Nutlet wing narrow, generally < 0.5 mm ..... var. purpusii (2)

49' Nutlet margin occasionally seeming sharp-angled, generally a ± flat narrow, occasionally uneven or minute-toothed rim, not strongly lobed, not a wide wing

52. Nutlets similar in 1 fruit, not differing in persistence, size, or margin shape

53. Corolla limb 1–2(2.5) mm diam; calyx 4–6 mm in fruit; nutlets 1.2–1.8(2) mm, not compressed, abaxially strongly convex, dome-like; bracts present ..... C. costata

53' Corolla limb 4–9 mm diam; calyx 2.5–4 mm in fruit; nutlets 1.8–2.5 mm, compressed, abaxially low-rounded; bracts 0 ..... C. oxygona

52' Nutlets dissimilar in 1 fruit, 1 more persistent, also differing in size or margin shape

54. Nutlet 1.3–1.7 mm, in 1 fruit 1 > others, margins ± same; basal leaf rosette generally 0 at flower; calyx lobe midvein dense-spreading-bristly ..... C. inaequata

54' Nutlet 2–2.5 mm, in 1 fruit 1 not > others but with margin ± sharp-angled, (2)3 with margin a ± flat narrow rim or wing; basal leaf rosette generally well developed at flower; calyx lobe midvein with 0–few bristles ..... C. pterocarya var. purpusii (2)

46' All nutlets with margin rounded or sharp-angled, not a ± flat rim or wing

55. Nutlets 2–4, dissimilar in 1 fruit, 1 more persistent, > other(s), of similar textures or not

56. Nutlets similar in texture

57. Plant generally erect, not sprawling; inflorescence dense, flowers ± 2-rowed, axis not ± flat; corolla limb 1–4 mm diam; nutlets 0.9–1.7 mm ..... C. angustifolia

57' Plant generally decumbent, sprawling; inflorescence not dense, flowers not 2-rowed, axis ± flat; corolla limb 0.5–1 mm diam; nutlets 1.5–2.5 mm ..... C. dumetorum

56' Nutlets dissimilar in texture, largest ± smooth, with smaller papillae to tubercles

58. Nutlets 2, 1.5–2 mm, lanceolate; bracts present; calyx 2–3 mm in fruit, oblong, lobe midvein hairs straight ..... C. maritima (2)

58' Nutlets 4, 0.7–1 mm, triangular-ovate; bracts 0; calyx 1–1.5 mm in fruit, ± spheric, lobe midvein hairs occasionally hooked ..... C. micromeres

55' Nutlets 1–4, generally of similar persistence, size, texture

59. Nutlets generally lanceolate to lance-ovate

60. Nutlets generally 3–4

61. Nutlets fine-papillate; spikes in 1s or 2s; ne East of Sierra Nevada, White and Inyo Mountains, ne Mojave Desert ..... C. scoparia

61' Nutlets coarsely tubercled to papillate; spikes in 2s or 3s; widespread, except ne East of Sierra Nevada, including White and Inyo Mountains, ne Mojave Desert ..... C. nevadensis

62. Plant ± sprawling; nutlets lanceolate; spikes not elongated in fruit; calyx lobes recurved at tips; corolla limb 1–2 mm diam ..... var. nevadensis

62' Plant ± erect; nutlets lance-ovate; spikes elongated in fruit; calyx lobes erect to ± recurved at tips; corolla limb 2–5 mm diam ..... var. rigida

60' Nutlets 1(2–3)

63. Calyx in fruit strongly recurved from stem; nutlet incurved ..... C. recurvata

63' Calyx in fruit not recurved from stem; nutlet(s) straight

64. Nutlet attachment scar edges a large, wide, deep-concave gap; calyx ± not constricted above nutlet(s) ..... C. excavata

64' Nutlet attachment scar edges a small, narrow, shallow groove; calyx ± tightly constricted above nutlet(s)

65. Corolla limb 1–2.5 mm diam

66. Corolla limb 1–1.5 mm diam, appendages 0; calyx lobe tips recurved to spreading ..... C. decipiens

66' Corolla limb 1.5–2.5 mm diam, appendages minute, pale yellow; calyx lobe tips spreading to erect ..... C. ganderi (2)

65' Corolla limb 3–6 mm diam

67. Calyx 1.5–2 mm, 3–3.5 mm in fruit, not appressed to stem in fruit, lobes oblong, not spreading-fine-bristly; nutlets 2.2–2.5 mm ..... C. corollata

67' Calyx 1–1.5 mm, 2.5–3(3.5) mm in fruit, appressed to stem in fruit, lobes linear, outer 3 spreading-fine-bristly; nutlets 1.5–2 mm ..... C. rattanii

59' Nutlets lance-ovate, ovate, or triangular-ovate

68. Nutlet abaxial ridge ± present

69. Corolla limb 3–8 mm diam

70. Calyx 1–1.5, 2–2.5 mm in fruit, ± round; nutlet ± papillate ..... C. incana (2)

70' Calyx 1.5–2, 3–4 mm in fruit, ovate; nutlet dense-sharp-papillate to tubercled ..... C. muricata var. muricata

69' Corolla limb 1–3.5 mm diam

71. Stem few-branched, without an evident central axis; nutlets 1.8–2.2 mm, ± tubercled ..... C. muricata var. denticulata

71' Stem many-branched, with an evident central axis; nutlets 1.1–1.3(1.9) mm, sharp-papillate ..... C. muricata var. jonesii

68' Nutlet abaxial ridge generally 0

72. Corolla limb (2)3–9 mm diam

73. Pedicel 2–3 mm in fruit; calyx silky-hairy ..... C. crinita

73' Pedicel 0–1.5(2) mm in fruit; calyx rough-hairy to bristly

74. Nutlets ovate, tip abruptly beaked; spikes generally in 1s to occasionally 3s, bracts generally sparse; leaves 1–2 cm, generally lowest 1–3 pairs opposite ..... C. mariposae

74' Nutlets lance-ovate to ovate, tip gradually tapered or acute; spikes generally in 2–4(5)s, bracts generally 0; leaves 1–5(8) cm, alternate

75. Calyx lobes narrow-oblong, midvein without coarse, spreading bristles; nutlets generally 1–2, ± ovate, sparse-low-tubercled (smooth) ..... C. intermedia var. hendersonii

75' Calyx lobes lance-linear to lanceolate, midvein with few to many coarse, spreading bristles; nutlets generally 3–4, lance-ovate, dense-tubercled to warty

76. Calyx 4.5–9 mm in fruit; leaves oblong to lanceolate; inflorescence generally in 2s; nutlet 1.9–2.5 mm ..... C. barbigera var. fergusoniae

76' Calyx 4–5.5 mm in fruit; leaves generally linear to lanceolate; inflorescence generally in 3s; nutlet 1.4–2.1(2.2) mm ..... C. intermedia var. intermedia

72' Corolla limb 0.5–2.5(3) mm diam

77. Bracts present; nutlets 1.2–1.4(1.5) mm

78. Calyx 4–6 mm in fruit, lanceolate; flower persistent; lowest stem leaves opposite; corolla limb generally < 1 mm diam; n&c San Joaquin Valley ..... C. hooveri

78' Calyx 3–4 mm in fruit, oblanceolate; flower not persistent; lowest stem leaves alternate; corolla limb 1.5–2 mm diam; s Channel Islands ..... C. traskiae

77' Bracts generally 0; nutlets 1.4–2.8 mm

79. Nutlet attachment scar edges triangular-flare-gapped at base

80. Nutlets 1.5–2 mm, tubercle tips not translucent; nutlets 1–4, lance-ovate to ovate ..... C. barbigera var. barbigera

80' Nutlets ± 3 mm, tubercle tips translucent; nutlets 3–4, triangular-ovate ..... C. clokeyi

79' Nutlet attachment scar edges wide-forked, but generally abutted, not gapped, at base

81. Nutlet tubercles very numerous, narrow, ± longer than broad; pedicel < 0.5 mm in fruit ..... C. echinella

81' Nutlet tubercles fewer, wider than broad, rounded; pedicel 0.5–1 mm in fruit

82. Spikes generally in 1s; stem branches 1–many; plant generally ± without main stem ..... C. ambigua

82' Spikes generally in 2s or 3s; stem branches few; plant with main stem ..... C. simulans

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Citation for the whole project: Jepson Flora Project (eds.) [year] Jepson eFlora, http://ucjeps.berkeley.edu/IJM.html [accessed on month, day, year]
Citation for an individual treatment: [Author of taxon treatment] [year]. [Taxon name] in Jepson Flora Project (eds.) Jepson eFlora, [URL for treatment]. Accessed on [month, day, year].
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