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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.

 

GALIUM BEDSTRAW

Valerie Soza


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

3' Fruit 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. Leaves in whorls of 3–4 and some opposite; fruit hairs hooked ..... G. proliferum

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

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

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

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

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

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

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

1' Perennial herb 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. Fruit hairs hooked

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

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

12' Fruit hairs 0 or not hooked

14. Flowers always bisexual

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

15' Leaves 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' Flowers generally unisexual, either anthers or ovaries small, sterile, or sometimes flowers unisexual and bisexual

17. Fruit a berry, glabrous to short-hairy

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

18' Leaves generally in whorls of 4

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

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

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

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

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

23' Leaf 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. Leaf linear, margin strongly rolled under; Outer South Coast Ranges ..... G. clementis

25' Leaf generally ovate (obovate) to elliptic, margin not or ± rolled under; Northwestern California, Central Western California, Southwestern California, Sierra Nevada

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

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

27. Leaf length 5–8 × width; n Sierra Nevada Foothills ..... subsp. sierrae (2)

27' Leaf length 2–4 × width; Northwestern California, Central Western California, Southwestern California

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

28' Flower, fruit hairy to glabrous; plant not woody, generally open; Northwestern California, Central Western California, Southwestern California

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

29' Plant generally not low, stem generally > 16 cm; leaf generally > 6 mm; North Coast Ranges, Central Western California, Southwestern California

30. Hair short, fine; South Coast, San Jacinto Mountains ..... subsp. primum (2)

30' Hair long, coarse to fine (0); not South Coast or San Jacinto Mountains

31. Hairs generally coarse, sparse; Outer North Coast Ranges, Inner North Coast Ranges, Central Western California ..... subsp. californicum (2)

31' Hairs generally fine, dense (0); Outer South Coast Ranges, n Channel Islands, Western Transverse Ranges, San Gabriel Mountains ..... 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. Leaf tip acute to obtuse to rounded, generally short-pointed, generally not sharp to touch, terminal hair generally not persistent ..... G. porrigens

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

34' Leaf ± linear ..... var. tenue

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

35. Leaf elliptic to ovate, acute to acuminate; plant not densely tangled, green; Outer South Coast Ranges, Western Transverse Ranges

36. Plant not shiny; stem wiry; leaf faces hairy; generally < 200 m, Central Coast, Outer South Coast Ranges ..... G. californicum subsp. maritimum

36' Plant shiny; stem stout; leaf faces ± glabrous; >= 200 m, Outer South Coast Ranges, Western Transverse Ranges ..... G. cliftonsmithii

35' Leaf ± linear to narrowly ovate, acute; plant densely tangled, red in age; South Coast, Channel Islands, Western Transverse Ranges, Peninsular Ranges ..... G. nuttallii

37. Plant hairs generally 0; Channel Islands ..... subsp. insulare

37' Plant hairs minute, sharp, curved; South Coast, Western Transverse Ranges, Peninsular Ranges ..... subsp. nuttallii

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

38. Leaf generally ± glabrous to puberulent or scabrous

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

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

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

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

38' Leaf generally hairy

41. Stem ± stout, ± stiff; montane

42. Plant sprawling or climbing, forming dense clumps; flower yellow, hairy externally, throughout; fruit hairy; leaf generally widely elliptic; San Gabriel Mountains ..... 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 San Gabriel Mountains ..... G. bolanderi (2)

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

43. Leaf length 5–8 × width; n Sierra Nevada Foothills ..... subsp. sierrae (2)

43' Leaf length 2–4 × width; Northwestern California, Central Western California, Southwestern California

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

44' Flower, fruit hairy to glabrous; plant not woody, generally open; Northwestern California, Central Western California, Southwestern California

45. Hair short, fine; South Coast, San Jacinto Mountains ..... subsp. primum (2)

45' Hair long, coarse to fine (0); not South Coast or San Jacinto Mountains

46. Hairs generally coarse, sparse; Outer North Coast Ranges, Inner North Coast Ranges, Central Western California ..... subsp. californicum (2)

46' Hairs generally fine, dense (0); Outer South Coast Ranges, n Channel Islands, Western Transverse Ranges, San Gabriel Mountains ..... subsp. flaccidum (2)

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

47. Flowers unisexual and bisexual

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

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

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

48' Erect shrub; stem rigid, stout, woody; Channel Islands

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

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

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

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

47' Flowers 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. Leaf ± 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 Channel Islands ..... subsp. foliosum

56' Woody stem internode generally > leaf; plant hairy or not; s Sierra Nevada, South Coast Ranges, Southwestern California (except n Channel Islands), Desert Mountains

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

57' Plant tall, slender, generally glabrous except leaves scabrous; leaves deciduous; San Bernardino Mountains, e San Jacinto Mountains, Desert Mountains, Sonoran Desert ..... subsp. gracillimum

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

58. Stem generally hairy, occasionally scabrous

59. Stem, leaf hairs ± equal; stem ridges generally narrower than grooves; San Gabriel Mountains ..... subsp. gabrielense

59' Stem hairs < leaf hairs; stem ridges generally wider than grooves; s High Sierra Nevada ..... subsp. onycense

58' Stem ± glabrous

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

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

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

61' Plant 6–20 cm; leaf generally 2–15 mm; internodes < 2 × leaves; San Gabriel Mountains, San Bernardino Mountains ..... subsp. nudicaule

53' Leaf 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. Stem flexible, few-branched; leaf ovate to elliptic, tip obtuse to acute, not sharp at tip; s High Sierra Nevada, Transverse Ranges ..... G. hallii

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

62' Plant woody near base only

64. Stem glabrous

65. Fruit <= 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' Fruit generally > 4 mm wide, including hairs

67. Inflorescence few-flowered, axillary, little exserted from leaves; fruit 7–10 mm; leaf tip acute to obtuse, not sharp to touch; Klamath Ranges, High Cascade Range ..... G. glabrescens subsp. glabrescens (2)

67' Inflorescence many-flowered, terminal, much exserted from leaves; fruit generally 4–7 mm; leaf tip obtuse to acuminate, sharp; n High Sierra Nevada, Great Basin Floristic Province, Desert Mountains

68. Leaves at a node ± equal, lanceolate to ovate, tapered to tip, flat to arched; Desert Mountains ..... G. argense

68' Leaves at a node ± unequal, generally ± arched, larger 2 ovate to round, abruptly rounded to acuminate tip; n High Sierra Nevada, Great Basin Floristic Province, Desert Mountains ..... G. multiflorum (2)

64' Stem puberulent to hairy

69. Leaves 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. Leaf flat, larger leaves lanceolate to ovate; Desert Mountains (Panamint Range) ..... G. hilendiae subsp. carneum

71' Leaf ± arched, larger leaves ovate to round; High Sierra Nevada, Great Basin Floristic Province, Desert Mountains ..... G. multiflorum (2)

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

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

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

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

72' Leaf lanceolate to (ob)ovate or round

74. Fruit >= 6 mm, including 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. Leaf generally fleshy; stem <= 9 cm; Desert Mountains (Panamint Range) ..... G. hypotrichium subsp. tomentellum (2)

76' Leaf not fleshy; stem <= 27 cm; Klamath Ranges, High North Coast Ranges, High Cascade Range, n High Sierra Nevada ..... G. grayanum

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

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

74' Fruit <= 6 mm, including 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; Mojave Desert ..... subsp. hilendiae

79' Corolla throat narrow; leaf lanceolate to ovate; Desert Mountains (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. Leaf thin; corolla rotate; Warner Mountains ..... G. glabrescens subsp. modocense

80' Leaf generally fleshy; corolla rotate to bell-shaped; High Sierra Nevada, East of Sierra Nevada, Desert Mountains ..... 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 High Sierra Nevada ..... subsp. subalpinum

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

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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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