The Jepson Herbarium
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Annual, perennial herb, occasionally ± shrubby, glabrous to hairy, generally scabrous; dioecious, bisexual, or flowers unisexual and bisexual.
Stem: 4-angled, occasionally ridged lengthwise.
Leaf: generally in whorls of >= 4, including leaf-like stipules.
Inflorescence: panicles, axillary clusters ( cymes), or occasionally 1 in axils.
Flower: bisexual, or unisexual with sterile stamens or pistils; calyx 0; corolla generally rotate, occasionally ± bell-shaped, generally ± green, yellow to white, occasionally pink or red, lobes generally 4; ovary 2-lobed, styles 2, bases ± fused.
Fruit: 2 nutlets or berry.
± 650 species: worldwide, especially temperate. (Greek: milk, from use of some species for curdling) [Dempster 1978 Univ Calif Publ Bot 73:1–33; Soza & Olmstead 2010 Amer J Bot 97:1630–1646] Ovary and fruit generally ± equally hairy on a plant; staminate plants generally identified by vestigial ovaries, pistillate plants generally by vestigial anthers. Galium saxatile L., Galium schultesii Vest, and Galium verum L. are lawn weeds in CA.
Unabridged references: [Dempster 1973 Univ Calif Publ Bot 64:1–36; Dempster & Ehrendorfer 1965 Brittonia 17:289–334; Dempster & Stebbins 1968 Univ Calif Publ Bot 46:1–52; Dempster & Stebbins 1971 Madroño 21:71–95]

Key to Galium

1. Annual

2. Corolla lobes generally 3

3. Fr hairs hooked; leaves in whorls of 4, in 2 unequal pairs, or distal-most generally opposite; flowers generally 1 in axils ..... G. bifolium

3' Fr glabrous; leaves in whorls of 4–6; flowers several in axils ..... G. trifidum (2)

4. Plant weak, sprawling; stems generally >= 10 cm ..... subsp. columbianum (2)

4' Plant tufted or matted; stems generally <= 15 cm ..... subsp. subbiflorum (2)

2' Corolla lobes generally 4

5. Nutlets sausage-shaped, hooked hairs unevenly distributed ..... G. murale

5' Nutlets ± spheric, glabrous or hooked hairs evenly distributed, occasionally with hair-like tubercles

6. Lvs in whorls of 3–4 and some opposite; fruit hairs hooked ..... G. proliferum

6' Lvs in whorls of 5–8; fruit hairs hooked or not, or 0

7. Plant erect, very slender; leaf <= 9 mm

8. Fr glabrous; leaf generally weakly reflexed ..... G. divaricatum

8' Fr hairs generally hooked, or 0; leaf generally reflexed in age ..... G. parisiense

7' Plant generally climbing to trailing, coarse; leaf 12–31 mm

9. Fr hairs hooked; nutlet pair < 4 mm wide; pedicel slender, ascending to spreading ..... G. aparine

9' Fr with acute tubercles only; nutlet pair > 4 mm wide; pedicel stout, recurved ..... G. tricornutum

1' Perennial or shrub

10. Corolla generally 3-lobed; fruit 2 nutlets, spheric, glabrous, hard ..... G. trifidum (2)

11. Plant weak, sprawling; stem generally >= 10 cm ..... subsp. columbianum (2)

11' Plant tufted or matted; stem generally <= 15 cm ..... subsp. subbiflorum (2)

10' Corolla generally 4-lobed; fruit 2 nutlets, variously shaped, glabrous to hairy, or berry

12. Fr hairs hooked

13. Lvs in whorls of 4 ..... G. oreganum

13' Lvs in whorls of 6 ..... G. triflorum

12' Fr hairs 0 or not hooked

14. Fls always bisexual

15. Lvs in whorls of 4 ..... G. boreale

15' Lvs in whorls of 5–12

16. Corolla bell-shaped; ovary with short, upcurved hairs ..... G. mexicanum subsp. asperulum

16' Corolla rotate; ovary glabrous or small-tubercled ..... G. mollugo

14' Fls generally unisexual, either anthers or ovaries small, sterile, or sometimes flowers unisexual and bisexual

17. Fr a berry, glabrous to short-hairy

18. Lvs in whorls of 6 ..... G. hardhamiae

18' Lvs generally in whorls of 4

19. Plant low, generally cushion-like, generally not woody above ground

20. Lf often ± bristle- or awl-like, sometimes linear to lanceolate, sharp to touch, ± thick ..... G. andrewsii

21. Plant dense, internode < leaf, main stem obscure, leaf generally awl-like, ± keeled, ascending; plant glabrous ..... subsp. andrewsii

21' Plant elongate or ± dense, internode generally > leaf, main stem obvious or not, leaf wider than awl-like, flat, ascending or spreading; plant hairy or not

22. Plant open, occasionally hairy, main stem obvious; leaf generally flat, linear to lanceolate, generally spreading ..... subsp. gatense

22' Plant ± dense, generally glabrous, main stem ± obscure; leaf ± flat to ± keeled, ± linear or lance-linear, generally ascending ..... subsp. intermedium

20' Lf linear or elliptic to (ob)ovate, not sharp to touch, generally thin

23. Lf ± linear, margins ± flat or convex, generally ciliate ..... G. ambiguum

24. Lf generally thin, ± fine-hairy, margins ± flat; ovary, fruit generally glabrous to velvety ..... subsp. ambiguum

24' Lf generally thick, generally glabrous or margin bristly, margins generally convex; ovary, fruit densely hairy ..... subsp. siskiyouense

23' Lf ovate (obovate) to elliptic or linear, edges rolled under (strongly so, if leaf linear) or not, not ciliate or hairs not restricted to margin

25. Lf linear, margin strongly rolled under; SCoRO ..... G. clementis

25' Lf generally ovate (obovate) to elliptic, margin not or ± rolled under; NW, CW, SW, SN

26. Lf shiny, sparsely hairy, 6–10 mm; stem generally glabrous ..... G. muricatum

26' Lf ± hairy, not shiny, 2–25 mm; stem generally hairy ..... G. californicum (2)

27. Lf length 5–8 × width; n SNF ..... subsp. sierrae (2)

27' Lf length 2–4 × width; NW, CW, SW

28. Fl, fruit glabrous; plant congested, woody at base, generally cushion-like; n ChI ..... subsp. miguelense (2)

28' Fl, fruit hairy to glabrous; plant not woody, generally open; NW, CW, SW

29. Plant low, stem <= 16 cm; leaf generally <= 6 mm; n Santa Lucia Range, SCoRO ..... subsp. luciense

29' Plant generally not low, stem generally > 16 cm; leaf generally > 6 mm; NCoR, CW, SW

30. Hair short, fine; SCo, SnJt ..... subsp. primum (2)

30' Hair long, coarse to fine (0); not SCo or SnJt

31. Hairs generally coarse, sparse; NCoRO, NCoRI, CW ..... subsp. californicum (2)

31' Hairs generally fine, dense (0); SCoRO, n ChI, WTR, SnGb ..... subsp. flaccidum (2)

19' Plant often climbing, ± woody above ground

32. Plant generally with recurved prickles; woody stem generally long, slender, climbing or sprawling

33. Lf tip acute to obtuse to rounded, generally short-pointed, generally not sharp to touch, terminal hair generally not persistent ..... G. porrigens

34. Lf widely oblong to ovate ..... var. porrigens

34' Lf ± linear ..... var. tenue

33' Lf tip generally tapered, generally sharp to touch, terminal hair persistent

35. Lf elliptic to ovate, acute to acuminate; plant not densely tangled, green; SCoRO, WTR

36. Plant not shiny; stem wiry; leaf faces hairy; generally < 200 m, CCo, SCoRO ..... G. californicum subsp. maritimum

36' Plant shiny; stem stout; leaf faces ± glabrous; >= 200 m, SCoRO, WTR ..... G. cliftonsmithii

35' Lf ± linear to narrowly ovate, acute; plant densely tangled, red in age; SCo, ChI, WTR, PR ..... G. nuttallii

37. Plant hairs generally 0; ChI ..... subsp. insulare

37' Plant hairs minute, sharp, curved; SCo, WTR, PR ..... subsp. nuttallii

32' Plant prickles generally 0; woody stem generally short, climbing or not

38. Lf generally ± glabrous to puberulent or scabrous

39. Lf narrow to elliptic, ± sessile; inflorescence branchlets ± stout; corolla generally red ..... G. bolanderi (2)

39' Lf widely elliptic to ovate, ± petioled; inflorescence branchlets thread-like; corolla generally ± yellow ..... G. sparsiflorum

40. Lf glabrous, generally ovate, ± leathery ..... subsp. glabrius

40' Lf puberulent, elliptic to ± ovate, generally thin ..... subsp. sparsiflorum

38' Lf generally hairy

41. St ± stout, ± stiff; montane

42. Plant sprawling or climbing, forming dense clumps; flower yellow, hairy externally, throughout; fruit hairy; leaf generally widely elliptic; SnGb ..... G. grande

42' Plant tufted or ± climbing, not forming dense clumps; flower generally red, glabrous to sparsely hairy at base; fruit glabrous to hairy; leaf generally elliptic; n of SnGb ..... G. bolanderi (2)

41' St wiry, slender; often coastal, insular, sometimes montane ..... G. californicum (2)

43. Lf length 5–8 × width; n SNF ..... subsp. sierrae (2)

43' Lf length 2–4 × width; NW, CW, SW

44. Fl, fruit glabrous; plant congested, woody at base, generally cushion-like; n ChI ..... subsp. miguelense (2)

44' Fl, fruit hairy to glabrous; plant not woody, generally open; NW, CW, SW

45. Hair short, fine; SCo, SnJt ..... subsp. primum (2)

45' Hair long, coarse to fine (0); not SCo or SnJt

46. Hairs generally coarse, sparse; NCoRO, NCoRI, CW ..... subsp. californicum (2)

46' Hairs generally fine, dense (0); SCoRO, n ChI, WTR, SnGb ..... subsp. flaccidum (2)

17' Fr 2 nutlets, hair generally long, straight to strongly curved

47. Fls unisexual and bisexual

48. Herb; stem generally slender, flexible, base woody; mainland

49. Lf ovate to round, acute to obtuse; inflorescence narrow, branches short, ± erect ..... G. parishii

49' Lf linear to oblanceolate, acute to acuminate; inflorescence open, branches longer, spreading ..... G. wrightii

48' Erect shrub; stem rigid, stout, woody; ChI

50. Ovary hairs short, curved, ascending; n ChI ..... G. buxifolium

50' Ovary hairs long, straight, spreading (or ± 0); s ChI ..... G. catalinense

51. St, leaf hairs sparse, generally very short, straight to curved, not curly; San Clemente Island ..... subsp. acrispum

51' St, leaf hairs ± dense, short, curly; Santa Catalina Island ..... subsp. catalinense

47' Fls unisexual only

52. Ovary, fruit hairs all short, strongly curved, ascending; corolla bell-shaped, divided <= 1/2 to base ..... G. jepsonii

52' Ovary, fruit hairs generally long, straight, spreading; corolla generally rotate, or bell-shaped and generally divided > 1/2 to base

53. Lf ± linear to strap-shaped to linear-oblong, generally widest at or distal to middle

54. Distal internodes >> proximal; pedicel generally > fruit; fruit hairs < fruit ..... G. johnstonii

54' Distal internodes ± = proximal; pedicel generally < fruit; fruit hairs generally >= fruit ..... G. angustifolium

55. Corolla generally glabrous, or not more hairy than stem or leaf

56. Woody stem internode generally < leaf; plant generally glabrous except leaf margin; n ChI ..... subsp. foliosum

56' Woody stem internode generally > leaf; plant hairy or not; s SN, SCoR, SW (except n ChI), DMtns

57. Plant tall to low, ± stout, generally glabrous to white-hairy; leaves persistent; hills, mtns, not in D ..... subsp. angustifolium

57' Plant tall, slender, generally glabrous except leaves scabrous; leaves deciduous; SnBr, e SnJt, DMtns, DSon ..... subsp. gracillimum

55' Corolla generally hairy (sometimes glabrous in subsp. jacinticum)

58. St generally hairy, occasionally scabrous

59. St, leaf hairs ± equal; stem ridges generally narrower than grooves; SnGb ..... subsp. gabrielense

59' St hairs < leaf hairs; stem ridges generally wider than grooves; s SNH ..... subsp. onycense

58' St ± glabrous

60. Plant 35–60 cm, inflorescence pyramid-shaped, many-flowered, much-branched; DSon (Borrego Desert) ..... subsp. borregoense

60' Plant <= 35 cm; inflorescence narrow, few-flowered, few-branched; SW

61. Plant 17–35 cm; leaf generally 11–26 mm; internodes ± = leaves; SnJt ..... subsp. jacinticum

61' Plant 6–20 cm; leaf generally 2–15 mm; internodes < 2 × leaves; SnGb, SnBr ..... subsp. nudicaule

53' Lf generally (ob)ovate to lanceolate, elliptic, or ± round (to needle-like in Galium stellatum), generally widest proximal to middle

62. Plant woody at base and extending distally to >= 50 cm

63. St flexible, few-branched; leaf ovate to elliptic, tip obtuse to acute, not sharp at tip; s SNH, TR ..... G. hallii

63' St rigid, many-branched; leaf lanceolate to needle-like, tip acute to acuminate, sharp at tip; D ..... G. stellatum

62' Plant woody near base only

64. St glabrous

65. Fr <= 4 mm wide, including hairs

66. Plant flexible; leaf 8–23 mm, flat, tip tapered to acute, not sharp to touch ..... G. magnifolium

66' Plant stiff; leaf 2–10 mm, generally arched, tip acuminate, sharp to touch ..... G. matthewsii

65' Fr generally > 4 mm wide, incl hairs

67. Infl few-flowered, axillary, little exserted from leaves; fruit 7–10 mm; leaf tip acute to obtuse, not sharp to touch; KR, CaRH ..... G. glabrescens subsp. glabrescens (2)

67' Infl many-flowered, terminal, much exserted from leaves; fruit generally 4–7 mm; leaf tip obtuse to acuminate, sharp; n SNH, GB, DMtns

68. Lvs at a node ± equal, lanceolate to ovate, tapered to tip, flat to arched; DMtns ..... G. argense

68' Lvs at a node ± unequal, generally ± arched, larger 2 ovate to round, abruptly rounded to acuminate tip; n SNH, GB, DMtns ..... G. multiflorum (2)

64' St puberulent to hairy

69. Lvs at a node unequal, leaf tip generally sharp to touch, abruptly rounded to tapered to sharp tip

70. Corolla rotate ..... G. munzii

70' Corolla ± bell-shaped

71. Lf flat, larger leaves lanceolate to ovate; DMtns (Panamint Range) ..... G. hilendiae subsp. carneum

71' Lf ± arched, larger leaves ovate to round; SNH, GB, DMtns ..... G. multiflorum (2)

69' Lvs at a node equal or leaf tip not sharp to touch, acute to obtuse

72. Lf lanceolate to narrowly elliptic ..... G. serpenticum

73. Corolla generally ± cup-shaped; leaf tip generally flat; KR ..... subsp. scotticum

73' Corolla rotate; leaf tip generally reflexed; Wrn ..... subsp. warnerense

72' Lf lanceolate to (ob)ovate or round

74. Fr >= 6 mm, incl hairs

75. Plant generally green, minute-puberulent; leaf ovate to obovate ..... G. glabrescens subsp. glabrescens (2)

75' Plant ± gray, densely short-gray hairy; leaf widely ovate to round

76. Lf generally fleshy; stem <= 9 cm; DMtns (Panamint Range) ..... G. hypotrichium subsp. tomentellum (2)

76' Lf not fleshy; stem <= 27 cm; KR, NCoRH, CaRH, n SNH ..... G. grayanum

77. Plant 5–27 cm, ± open ..... var. grayanum

77' Plant 2.5–14 cm, ± compact ..... var. nanum

74' Fr <= 6 mm, incl hairs

78. Plant woody generally for 3–4 cm above base; inflorescence ± terminal, diffuse, branches many, long, spreading, many-flowered, leaves reduced; corolla bell-shaped ..... G. hilendiae

79. Corolla throat wide; leaf ovate to ± round; DMoj ..... subsp. hilendiae

79' Corolla throat narrow; leaf lanceolate to ovate; DMtns (Kingston Range) ..... subsp. kingstonense

78' Plant not woody above base; inflorescence axillary, narrow, dense, branches few, short, ascending, few-flowered, leaves often not reduced; corolla rotate to bell-shaped

80. Lf thin; corolla rotate; Wrn ..... G. glabrescens subsp. modocense

80' Lf generally fleshy; corolla rotate to bell-shaped; SNH, SNE, DMtns ..... G. hypotrichium

81. Plant generally open, tall, 12–30 cm, scabrous, base woody ..... subsp. inyoense

81' Plant generally dense or ± open, short, generally 2–15 cm, scabrous to hairy, base not woody

82. Plant minutely velvety ..... subsp. hypotrichium

82' Plant scabrous to hairy, not velvety

83. Plant green, scabrous to hairy; plant ± open; stem <= 15 cm; s SNH ..... subsp. subalpinum

83' Plant gray, ± densely fine-hairy; plant matted; stem <= 9 cm; DMtns (Panamint Range) ..... subsp. tomentellum (2)

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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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