Hypericum L.

1. Hypericum L. 4

(Inch Elodes Adanson and Triadenia Spach)

Flowers hermaphrodite. Sepals (4-)5. Petals (4-)5, yellow, sometimes tinged with red. Stamens in 3 or 5 fascicles of (l-)3 to c. 125, sometimes alternating with sterile fascicles (Jasciclodes), or in 5 irregular groups; fascicles, if 5, antepetalous; if 3, one antepetalous and two (larger) antesepalous. Ovary (2-)3- to 5-locular or partly or completely 1 -locular; ovules numerous. Styles (2-)3-5, free, slender. Fruit a septicidal capsule, rarely fleshy and more or less indéhiscent.

Glands are designated marginal if they protrude sufficiently to interrupt the line of the margin of a leaf, sepal or petal, intramarginal if they abut on the margin but do not interrupt its line, superficial if they are quite clear of the margin.

In many species the ovary and capsule have glandular streaks or patches on the wall. These are referred to as vittae if flat or slightly swollen, and as vesicles if conspicuously swollen. Vittae on or near the midrib of a carpel are described as dorsal.

1 Edit. T. G. Tutin. 2 By T. G. Tutin.

3 Edit. D. A. Webb. 4 By N. K. B. Robson.

Many European species are cultivated in gardens, as also are several species and hybrids of large-flowered shrubs from Asia belonging to the section Norysca (Spach) Endl.

Literature: B. Stefanov, God. Sof. Univ. ( Agron .-Les. Fak.) 10: 19-58 (1932); 11: 139-186 (1933); 12: 69-100 (1934).

1 Plant without red or black glands on leaves, sepals, petals or anthers

2 Leaves in whorls of 3-4

3 Petalsandstamens deciduous;leaves smooth 10. amblycalyx 3 Petals and stamens persistent; leaves papillose 12. ericoides 2 Leaves opposite

4 Broad-leaved shrub; petals and stamens deciduous

5 Stems and leaves covered with glandular vesicles

6. balearicum 5 Stems and leaves smooth

6 Styles 5; anthers reddish (Sect. Eremanthe ) 1. calycinum 6 Styles 3(—4): anthers yellow (Sect. Androsaemum )

7 Petals shorter than sepals; ripe fruit black and succulent

5. androsaemum 7 Petals longer than sepals; ripe fruit red or green, scarcely succulent

8 Sepals shrivelling and falling before fruit ripens; foliage often goat-scented 3. hircinum 8 Sepals persistent at least until fruit ripens; foliage not goat-scented

9 Sepals acute or cuspidate; fruit subglobose 2. foliosum 9 Sepals obtuse or subacute; fruit ellipsoid or narrowly ovoid 4. inodorum 4 Herbormicrophyllousshrub; stamensandusuallypetals persistent

10 Microphyllous shrub; stamens in 3 fascicles (Sect.

Triadenia )

11 Leaves linear-spathulate; inflorescence usually 3-flowered

7. acifenun 11 Leaves elliptical to narrowly oblong; flowers solitary

8. aegyptìcum 10 Herb; stamens 5, or in 5 irregular groups (Sect. Brathys )

12 Leaves linear-subulate, closely appressed; branches numerous, fastigiate 61. gentianoides 12 Leaves flat, ± patent; branchesfew or none, not fastigiate

13 Leaves suborbicular to ovate-triangular or broadly oblong

14 Stem simple or nearly so; middle and upper leaves ovate-triangular, ± acute 57. gymnanthum 14 Stem usually branched; middle and upper leaves oblong or ovate to suborbicular, obtuse 58. mutilum 13 Leaves linear to lanceolate, oblanceolate or narrowly oblong

15 At least the upper leaves lanceolate, with (3—)5—7 veins at the base; sepals 4-7 mm 59. majus 15 All leaves linear to linear-lanceolate or oblanceolate,

with 1—3(—5) veins at the base; sepals 2-5 mm

60. canadense 1 Red or black glands present, at least on leaves, sepals or anthers

16 Leaves in whorls of 3-4

17 Petals and stamens deciduous; leaves in whorls of 3

9. empetrifolium 17 Petals and stamens persistent; leaves usually in whorls of 4

18 Leaves smooth; sepals with sessile marginal glands 11. coris 18 Leaves papillose; sepals eglandular or glandular-ciliate

12. ericoides 16 Leavesopposite

19 Leaves without intramarginal black glands (apical black glands or intramarginal translucent glands sometimes present)

20 Sepalsentire,withoutmarginalorintramarginalglands

21 Sepals broadly imbricate; seeds reticulate-pitted

22 Glabrous 21. olympicum 22 Pubescent 22. cerastoides 21 Sepals not or only slightly imbricate; seeds papillose

23 Leaves 2-6 mm, ovate or ovate-elliptical, glaucous; flowers solitary 19. taygeteum 23 Leaves (5—)8—30 mm, linear to narrowly elliptical, not glaucous; inflorescence many-flowered

24 Stems 30-70 cm, scarcely rooting at the base; petals not red-tinged 20. hyssopifolium 24 Stems 5—20(—35) cm, rooting at the base; petals usually red-tinged 15. linarioides 20 Sepals ciliate, denticulate or fimbriate, with marginal or intramarginal glands

25 Dwarf shrub, with stems conspicuously 4-angled

13. haplophylloides 25 Herb, with stems not or scarcely 4-angled

26 Lower leaves glandular-denticulate; capsule with orange vesicles 33. vesiculosum 26 Leaves entire; capsule with vittae, but without vesicles

27 Glands on sepals red; filaments united for $ of their length 32. elodes 27 Glands on sepals black; filaments free almost to the base

28 Leaves pubescent 14. hirsutum 28 Leaves glabrous

29 Inflorescence subcorymbose; petioles articulated at base; stems diffuse

30 Leaves glaucous only beneath; petals 10-16 mm

17. nummularium 30 Leaves glaucous on both sides; petals 6-9 mm

18. fragile 29 Inflorescence cylindrical or narrowly pyramidal;

petioles, if present, not articulated; stems erect or ascending

31 Anthers pink or orange; leaves ovate-cordate to oblong 16. pulchrum 31 Anthers yellow; leaves usually linear to linearlanceolate 32 Stems 30-70 cm, scarcely-rooting at the base;

petals not red-tinged 20. hyssopifolium

32 Stems 5-17(-32) cm, rooting at the base; petals usually red-tinged 15. linarioides 19 At least some of the leaves with intramarginal black glands (sometimes very few in 29 and 30)

33 Sepalswithoutmarginalorintramarginalblackglands

34 Sepals broadly imbricate; flowerless axillary shoots usually absent

35 Flowerslessthan2 cmin diameter;vittaeoncapsule conspicuous 50. humifusum 35 Flowersat least 2 cmin diameter;vittaeon capsulefaint or absent

36 Glabrous 21. olympicum 36 Pubescent 22. cerastoides 34 Sepals not, or only slightly imbricate; flowerless axillary shoots present (Sect. Hypericum )

37 Stems with2 raisedlines

38 Leaves neither undulate nor amplexicaul; branches of inflorescence ascending 54. perforatum 38 Leaves undulate, amplexicaul; branches of inflorescence patent 55. triquetrifolium 37 Stems with4 raisedlinesor wings

39 Sepals obtuse; stems not winged 53. maculatum 39 Sepals acute; stems winged

40 Petals not more than 7- 5 mm, pure yellow or rarely red-veined; leaves usually flat 51. tetrapterum 40 Petals at least 7 -5 mm, usually tinged with red; leaves undulate 52. undulatum 33 Sepalswithmarginalorintramarginalblackglands

41 Leaves hairy or papillose on both sides

42 Capsule withvesiclesas wellas vittae;stems glabrousor finely papillose

43 Stems 1 -5(-10) cm, procumbent, glabrous 44. kelleri 43 Stems 15-45 cm, erect or decumbent, usually minutely papillose 45. aviculariifolium 42 Capsule without vesicles; stem hairy

44 Leaves connate at base 31. caprifolium 44 Leaves free

45 Bracts auriculate; sepals with long teeth or cilia

46 Plant puberulent to velutinous; stems ± erect, not rooting at the nodes 24. annulatum 46 Plant hirsute; stems decumbent or procumbent, rooting at the nodes

47 Leaves 12-45 mm, sessile 25. delphiciun 47 Leaves 8-15 mm, shortly petiolate 26. athoum 45 Bracts not auriculate; sepals subentire or with short teeth or cilia

48 Sepals glabrous, with superficial black glands

49 Petals at least 8 mm; leaves at least 15 mm, sessile

27. atomarium 49 Petals not more than 8 mm; leaves not more than

15 mm, usually petiolate 28. cuisinii 48 Sepals hairy, without superficial black glands

50 Sepals 5-10 mm, aristate, entire; petals 9-15 mm

30. pubescens 50 Sepals 3-6 mm, rarely aristate, shortly glandularciliate; petals 6-11 mm 29. tomentosum 41 Leaves smooth and glabrous, at least above

51 Capsule with longitudinal vittae only; seeds reticulatepitted

52 Styles 5 46. thasium 52 Styles (2-)3(-4)

53 Flowerless axillary shoots present; stems with black glands

54 Sepals entire, with intramarginal black glands; leaves undulate 52. undulatum 54 Sepals glandular-denticulate or with sessile marginal glands; leaves flat 56. elegans 53 Flowerless axillary shoots absent; stems usually without black glands

55 Sepals equal, regularly dentate or ciliate

56 Stems erect; leaves 20-70 mm 23. montanum

56 Stems diffuse or decumbent; leaves2-15mm 28. cuisinii

55 Sepals often unequal, subentire or irregularly ciliate or fimbriate (Sect. Oligostema)

57 Sepals fimbriate or denticulate 47. aucberi

57 Sepals entire or ciliate

58 Sepals unequal, with few or no superficial black glands; petals 4-6(-8) mm 50. humifusum

58 Sepals ± equal, with numerous superficial black glands; petals (5—)7—15 mm

59 Leaves on flowering stems linear to linearlanceolate; sepals glandular-ciliate 48. linarifolium

59 Leaves on flowering stems ovate to lanceolate or oblong; marginal glands of sepals usually sessile 49. australe

51 Capsule with some vesicles or oblique vittae, or almost smooth; seeds usually longitudinally ribbed (Sect.

Drosocarpium)

60 Leaves with dense, conspicuous, reticulate venation

61 Capsule with black vesicles 38. richeri

61 Capsule without black vesicles

62 Capsule with interrupted vittae; leaves with numerous superficial glands 36. umbellatum

62 Capsule with orange vesicles; leaves with few or no superficial glands 37. bithynicum

60 Leaves with lax or indistinct reticulate venation

63 Black glands on petals absent, or confined to margin and apex

64 Glands on capsule prominent

65 Capsule narrowly pyramidal; sepals patent or deflexed in fruit 35. montbretii

65 Capsule ovoid; sepals erect in fruit

66 Capsule with dorsal vittae and lateral vesicles;

leaves with numerous translucent glands

34. perfoliatum

66 Capsule with vesicles only; leaves usually without translucent glands 40. rochelii

64 Glands on capsule not prominent

67 Sepals eglandular-fimbriate 41. barbatimi

67 Sepals entire or glandular-denticulate to -fimbriate

68 Leaves 5-14 mm, smooth; seeds longitudinally grooved 43. trichocaulon

68 Leaves 2-6 mm, papillose or undulate; seeds reticulate-pitted 44. kelleri

63 Black glands dispersed over whole surface of petals

69 Capsule with prominent vesicles 39. spruneri

69 Capsule smooth, or with slightly prominent glands

70 Sepals eglandular-fimbriate 41. barbatum

70 Sepals entire or glandular-denticulate to -fimbriate

71 Petals 13-20 mm; sepals glandular-ciliate to -fimbriate 42. rumeliacum

71 Petals 6-12 mm; sepals entire to glandulardenticulate

72 Leaves 5-14 mm, smooth; seeds longitudinally grooved 43. trichocaulon

72 Leaves 2-6 mm, papillose or undulate; seeds reticulate-pitted 44. kelleri