Rubus L.

9. Rubus L. View in CoL 2

Perennial herbs or shrubs. Stems usually with prickles. Leaves usually pinnate, digitate or pedate, with 3-7 dentate leaflets. Flowers solitary or in racemose or paniculate inflorescences. Flowers usually 5-merous; hypanthium flat, with a large, usually convex receptacle; epicalyx absent; petals red, purple, pink or white; stamens numerous; carpels numerous; styles subterminal, usually deciduous; ovules 2. Fruit usually a coherent head of 1-seeded drupelets.

The European species of Rubus , native and naturalized, are placed in 5 subgenera. The first four contain 9 species, and present no taxonomic difficulties. In the remaining subgenus, Rubus , some 2000 species have been described. Almost all are agamospecies, segregated from R. fruticosus L. This name, which is based on a mixture of two species (12, R. plicatus and 38, R. ulmifolius ) belonging to different subsections, is now used only in an aggregate sense, and it covers the whole of the section Rubus , except for 75 and the Corylifolii (see note following 75). Many of the species are tetraploid (In = 28) but diploids, triploids, pentaploids and hexaploids are also known. It is likely that many o f the species have arisen during the Pleistocene era as a result of hybridization and apomixis. The apomicts are all polyploid and pseudogamous; apomixis is often facultative so that new apomictic biotypes can arise at the present time by hybridization, thus increasing the number of potential taxa.

It is thus not profitable to treat fully and by conventional means the whole array of agamospecies, and no attempt at a detailed treatment is made here. The full spectrum of morphological variation has, however, been covered, and this has been done by recognizing 66 circle-species which are relatively wide­ spread and distinct. Round these the remaining species can be grouped. (This method was proposed by Gustafsson, op. cit.). The description of a circle-species applies only to the circlespecies and does not cover the related species which are listed after the appropriate circle-species. All species recognised in Basic and Standard Floras are listed. The key should enable all European species to be run down to their respective subsections and sometimes series, but below this level only those species selected as ‘circle-species’ are keyed out.

The classificatory framework is based on that of Sudre (op. cit.).

In the descriptions the stem- and leaf-characters are those of the middle part of the first-year or non-flowering stem of a wellgrown plant in a normal environment.

The armature may include large prickles, smaller pricklets, acides, stalked glands and eglandular hairs. The acides, which may be gland-tipped, are stouter and more rigid than the glands or hairs, but are always straight and never expanded at the base. Prickles which are straight but directed backwards along the stem are said to be deflexed', those which are curved as well as deflexed are said to be falcate. Sepals which are furnished with pricklets are aculeolate', sepals with the apex prolonged are appendiculate', descriptions of the indumentum of sepals referto the outer surface.

Although the inflorescence is always determinate, it resembles a raceme, corymb or panicle in appearance and is called by these terms for convenience.

Endemic signs have not been used for the ‘related species’ because of lack of information, but it is probable that the great majority of those listed are endemic to Europe.

Literature: W. O. Focke, Biblioth. Bot. (Stuttgart) 72(1-2): 1-223 (1910-11); 83 (1914). Â. Gustafsson, Lunds Univ. Ârsskr. ser. 2, 39(6) (1943). H. Sudre, Rubi Europae. Paris. 1908-1913. W. C. R. Watson, Handbook o f the Rubi o f Great Britain and Ireland. Cambridge. 1958. K. E. Weihe & C. G. Nees von Esenbeck, Rubi Germanici. Eberfeld. 1822-27.

Some of the more recent Floras treat Rubus very fully, e.g.: J. Legrain in W. Robyns, Flore Générale de Belgique 3: 10-274 Bruxelles. 1958-59. E. I. Nyärady in T. Sävulescu, Flora Republicii Populäre Romàne 4: 276-580. Bucureçti. 1956. J. Dostâl, Kvëtena CSR 572-630. Praha. 1950. Ä. Kiss in R. Soó & S. Jâvorka. A Magyar Nôvényvilâg Kézikônyve 251-270. Budapest. 1951.

1 Stems herbaceous or nearly so; stipules free from petiole

2 Leaves simple

3 Dioecious; ripe fruit orange 1. chamaemorus 3 Flowers hermaphrodite; ripe fruit red 2. humulifolius 2 Leaves ternate

4 All stems flower-bearing, without prickles; petals pink

3. arcticus 4 Some stems vegetative, with prickles; petals white 4. saxatilis 1 Stems woody; stipules united with petiole

5 Leaves simple; receptacle flat 5. odoratus 5 Leaves compound; receptacle convex

6 Ripe fruit red or orange, separating from receptacle; leaves ternate or pinnate

7 Leaves subglabrous beneath; flowers ±solitary; petals bright purple 6. spectabilis 7 Leaves white-tomentose beneath; flowers in racemes; petals white or pink

8 Stems densely covered with reddish bristles; petals pink, curved inwards 9. phocnicolasius 8 Stems not densely covered with bristles; petals white, erect

9 Inflorescence-axis glandular,with abundant acides; leaves all ternate 8. sachalinensis 9 Inflorescence-axis eglandular, with sparse acides; leaves usually pinnate 7. idaeus 6 Ripe fruit blackish, adhering to receptacle; leaves ternate, digitate or pedate

10 Stemspruinose; stipuleslanceolate; leaflets3; drupelets 2-20, pruinose (Subsect. Caesii ) 15. caesius 10 Stemsusuallynotpruinose; stipuleslinear-lanceolateto filiform; leaflets 3-7; drupelets usually more than 20, rarely pruinose (vide also p. 25 Corylifolii )

11 Pricklesnearlyalwaysveryunequal, oftenscattered onthe stem-faces; stem and inflorescence usually with stalked glands (Subsect. Appendiculatï)

12 Stems with sparse glands; upper leaves usually covered with stellate hairs above; petals yellowishwhite 44. canescens 12 Stems usually with many glands; leaves without stellate hairs above; petals never yellowish

13 Stems weak and terete, often procumbent, pruinose; prickles weak, or broad-based and curved; leaves normally with 3-5 leaflets; inflorescence usually with weak prickles; sepals appressed to the young fruit; petals often small; stamens usually equalling or exceeding styles (Ser. Glandulosï )

14 Prickles compressed at base

15 Prickles on any one part of stem all similar; inflorescence with most glands shorter than the diameter of the axis; petals erect 70. scaber 15 Prickles on stem distinctly unequal; inflorescence with most glands longer than the diameter of the axis; petals ± patent 71. schleicheri 14 Prickles scarcely compressed at base

16 Terminalleaflet rounded and ± entire at base, shortly acuminate 72. glandulosus 16 Terminal leaflet cordate or subcordate at base

17 Glands and acides pale yellow 73. serpens 17 Glands and acides brown or purple 74. hirtus 13 Plant not possessing this combination of characters

18 Inflorescence-glands often longer than the diameter of the axis; prickles very unequal and merging into numerous stalked glands, acides, and pricklets of varying lengths

19 Stems robust, pruinose, with numerous, long, often glandular hairs and acides; prickles straight or falcate; leaves pedate with 3-5 leaflets, grey-white tomentose beneath; inflorescence large, with long lower branches and pedicels 69. incanescens 19 Plant without this combination of characters (Ser.

Histrices )

20 Flowers usually less than 2 cm in diameter

21 Stem-prickles short, not confluent, not hairy; sepals without prickles; petals 5—7(— 11)

66. rosaceus 21 Stem-prickles large, confluent and hairy; sepals with prickles; petals 5 64. pilocarpus 20 Flowers usually more than 2 cm in diameter

22 Flowers white or pale pink 68. koehleri 22 Flowers bright or deep pink

23 Stems glabrous or glabrescent 67. histrix 23 Stems sparsely or densely hairy

24 Stems densely hairy; terminal leaflet ovate to suborbicular, acute, cordate at base 63. fuscater 24 Stems sparsely hairy; terminal leaflet obovate, acuminate, subcuneate at base 65. lejeunei 18 Inflorescence-glands shorter than the diameter of the axis; stem-prickles slightly or moderately unequal,

but distinct from the pricklets

25 Stem-prickles only slightly unequal, mainly on the angles of the stem, with a few tuberculate pricklets on the faces and a few stalked glands (Ser. Vestiti )

26 Leaves green beneath

27 Terminal leaflet orbicular or obovate, with a ± abrupt, short or long point; sepals patent, green, with white margins 48. mucronulatus 27 Terminal leaflet ovate or ovate-oblong, gradually narrowed to a long point; sepals deflexed, tomentose 49. gremiti 26 At least the upper leaves white-tomentose beneath

28 Petals suborbicular 45. vestitus 28 Petals ovate or elliptical, longer than wide

29 Stems with sparse, tufted hairs and a few glands; sepals patent to erect 46. boraeanus 29 Stems pubescent, with long and short hairs and sunken glands; sepals deflexed 7. adscitus 25 Stem-prickles unequal, scattered over the stem surface making it rough and tuberculate, with many stalked glands

30 Inflorescence-axis glandular, tomentose or glabrous (Ser. Rudes )

31 Stems ± terete, becoming white-pruinose; leaflets usually 3 62. vallisparsus 31 Stems angled, not white-pruinose; leaflets usually 5

32 Cauline leaves cordate at base; inflorescence flexuous and lax 61. melanoxylon 32 Cauline leaves subcuneate at base; inflorescence short, subcorymbiform 60. rudis 30 Inflorescence-axis glandular and hairy, the glands mainly not exceeding the hairs (Ser. Radulae )

33 At least the upper leaves white-tomentose beneath

34 Inflorescence with few, or weak, slender prickles; stamens about equalling styles 52. apiculatus 34 Inflorescence with numerous strong prickles; stamens exceeding styles

35 Sepals not appendiculate; carpels glabrous

50. radula 35 Sepals appendiculate; carpels pubescent

51. genevieri 33 Leaves all green beneath, though sometimes greentomentose

36 Sepals usually deflexed after anthesis, or variable in the same plant

37 Stems densely hairy; inflorescence with many prickles; bracts long, linear-lanceolate 53. fuscus 37 Stems sparsely hairy; inflorescence with very few or no prickles; bracts leaf-like 54. foliosus 36 Sepals patent or becoming erect after anthesis

38 Stems with few or no hairs

39 Sepals green, with white margin, acute, without long apex; inflorescence short, few-flowered

55. infestas 39 Sepals grey-tomentose, without white margin, with long, linear apex; inflorescence usually long, dense and tapered 56. thyrsiflorus 38 Stems hairy

40 Flowers deep pink 58. obscurus 40 Flowers white or pale pink

41 Inflorescence with very few prickles 57. pallidus 41 Inflorescence with numerous prickles 59. menkei 11 Prickles ± equal, mainly on the stem-angles; stem and inflorescence ± eglandular, or with sessile glands only

42 Stems glabrous, often suckering from base, rarely rooting apically; inflorescence a raceme or panicle (Subsect. Suberecti)

43 Stems high-arching; inflorescence a panicle

44 Leaflets 5-7, those of upper leaves persistently pubescent or almost tomentose beneath; receptacle densely pubescent 15. affinis 44 Leaflets 5, often glabrous at maturity; receptacle glabrous 14. divaricatus 43 Stems ± erect; inflorescence usually a raceme

45 Leaflets 5 or 7; prickles weak or short; ripe fruit dark red

46 Stems up to 300 cm; prickles short, conical, often purplish-black; leaflets plane, glabrescent beneath

10. nessensis 46 Stems not more than 150 cm; prickles slender, subulate, yellowish; leaflets plicate, pubescent beneath

11. scissus 45 Leafletsusually5; pricklesstrong; fruitblack

47 Stems with rather few, mainly patent, strong prickles; terminal leaflet rather long-acuminate, evenly serrate 13. sulcatus 47 Stems with numerous, falcate or deflexed prickles; terminal leaflet shortly acuminate, unevenly serrate

12. plicatus 42 Stems often hairy, usually rooting apically, not suckering; inflorescence a panicle

48 All leavesgrey-whitetomentosebeneath; sepalsgrey- white tomentose externally, deflexed in fruit (Subsect. Discolores )

49 Stemspruinose

50 Leaves coriaceous, dark green above, whitetomentose beneath; stamens scarcely exceeding the styles 38. ulmifolius 50 Leaves not coriaceous or dark green, tomentose and pubescent beneath; stamens greatly exceeding the styles 39. godronii 49 Stems not or scarcely pruinose

51 Basal leaflets subsessile; inflorescence often with few prickles above; peduncles elongated 43. candicans 51 Basal leaflets shortly stalked; inflorescence with many prickles; peduncles not distinctly elongated

52 Inflorescence-prickles patent, mostly straight.

40. bifrons 52 Inflorescence-prickles falcate

53 Stems glabrescent, angled but not sulcate

41. discolor 53 Stems hairy and sulcate 42. chloocladus 48 Leavesgreen,oronlytheuppergrey-whitetomentose beneath; sepals green or grey-tomentose externally, their posture in fruit variable (Subsect. Silvatici)

54 Leaveslaciniate 27. laciniatus 54 Leaves not laciniate

55 At least some of the lower leaves grey- or whitetomentose beneath

56 Petals 3 cm or more (Açores) 36. hochstetterorum 56 Petals less than 3 cm

57 Upper leaves white-tomentose beneath

35. rhamnifolius 57 Upper leaves grey-tomentose beneath, or not tomentose

58 Panicle narrow, long; rhachis-prickles falcate;

sepals white-tomentose 37. lindebergii 58 Panicle not long and narrow; rhachis-prickles deflexed; sepals grey-white tomentose

59 Leaflets of cauline leaves sometimes 7, becoming distinctly convex; terminal leaflet cuspidate;

sepals not long-pointed 34. polyanthemus 59 Leaflets of cauline leaves always 5, not distinctly convex; terminal leaflet acuminate; sepals usually long-pointed 33. villicaulis 55 At least the lower leaves green beneath

60 Petals fimbriate 18. pedatifolius 60 Petals not fimbriate

61 Sepals erect or patent

62 Stems angled; leaflets usually 5; sepals never appendiculate; stamens usually exceeding styles

63 Inflorescence with stalked glands

64 Stem-prickles few, long-subulate 21. bypomalacus 64 Stem-prickles numerous, some of them strong,

broad-based, falcate 20. chaerophyllus 63 Inflorescence with subsessile glands, or ±

eglandular

65 Inflorescence nearly unarmed or with weak prickles 19. gratus 65 Inflorescence with numerous strong prickles

66 Terminal leaflet elliptical or obovate, rounded or subcordate at base, coarsely serrate;

inflorescence-prickles falcate 17. vulgaris

66 Terminal leaflet broadly ovate to elliptical,

subcordate at base, finely serrate; inflorescence-prickles usually straight 16. lentiginosus

62 Stems usually terete; leaflets usually 3; sepals usually appendiculate; stamens shorter than,

or just equalling the styles

67 Stems angled, at least above; leaflets 5

25. chlorothyrsos

67 Stems terete or weakly angled; leaflets usually 3

68 Sepals green, with white margins 22. arrhenii

68 Sepals grey-tomentose, without white margins

69 Stems and inflorescence-axis hairy; petals bright pink 23. sprengelii

69 Stems and inflorescence-axis ± glabrous;

petals white 24. myricae

61 Sepals deflexed in fruit

70 Stems ± weak

71 Stems hairy; leaflets 5 31. silvaticus

71 Stems ± glabrous; leaflets usually 3 32. egregius

70 Stems strong, angled; leaflets usually 5

72 Leaves glabrescent beneath; petals emarginate

26. questieri

72 Leaves pubescent or tomentose beneath; petals entire

73 Stems angled, with plane faces; terminal leaflet usually subcordate 29. pyramidalis

73 Stems angled, with sulcate faces; terminal leaflet usually rounded or cuneate at base

74 Leaves green and pubescent, or slightly greytomentose beneath 30. macrophyllus

74 Leaves white-tomentose beneath 28. rhombifolius