Knautia L.

Tutin, T. G., Heywood, V. H., Burges, N. A., Moore, D. M., Valentine, D. H., Walters, S. M. & Webb, D. A., 1976, Flora Europaea. Volume 4. Plantaginaceae to Compositae (and Rubiaceae), Cambridge University Press : 60-62

6. Knautia L. 2

Annual to perennial, herbs Leaves opposite, undivided to pinnate. Capitula long-pedunculate, hemispherical to cylindrical, hermaphrodite or female. Involucral bracts numerous, herbaceous, free. Receptacle hemispherical, hairy, without scales. Involucel compressed, 4-angled, inconspicuous, entire to dentate in fruit. Calyx patelliform to cupuliform, with (6-)8-16(-24) minute apical awns or teeth, deciduous. Corolla-tube short; limb unequally 4-lobed, patent and often larger in marginal florets. Fruit ovoid, oblong or cylindrical, more or less hairy, with soft elaiosome at base. The perennial and biennial species (1-44) belong to Sect. Trichera and form an extremely polymorphic group of diploids, tetraploids and hexaploids (x=10). Hybridization is frequent and has given rise to many intermediate populations, as well as more or less autonomous races; taxonomic boundaries are, therefore, often obscured. It would be possible, for convenience, to assemble all these taxa into one species-group, but 8-15, 18-22, 23-24, 28-37, 38-41 and 42-43 form particularly closely inter-related subgroups.

The structure of the monopodial or sympodial stock and the colour of the corolla are diagnostic, and should be noted in the field. Several plants from each population should be examined, because of variation in the division of the leaves, hairiness, and presence or absence of glandular hairs on the peduncles. A double indumentum is sometimes present on stems or leaves, consisting of both pubescence and another kind of hairs.

Literature: Z. Szabó, Math. Term. Közl. 31: 1-436 (1911); Bot. Közl. 31: 115-141 (1934). F. Ehrendorfer, Österr. Bot. Zeitschr. 109: 276-343 (1962). S. Breton-Sintes, Étude Biosystématique du Genre Knautia (L.) Coult. en Auvergne, Thèse Sér. E, no. 146, Univ. Clermont-Ferrand. 1971. 1 Annual, with a slender root

2 Calyx patelliform, 8- to 10-awned; involucral bracts with rigid cilia, eglandular 45. byzantina 2 Calyx cupuliform, 12- to 24-dentate, the teeth rarely awned; involucral bracts ciliate-hirsute or shortly setose

3Capitula with 20-40 florets; involucre patelliform, the bracts with inconspicuous veins; peduncles usually eglandular

46. integrifolia 3Capitula with 5-15 florets; involucre cyathiform to cylindrical, the bracts with prominent veins; peduncles glandular

4 Involucre cyathiform; capitula with 10—15 florets; corolla lilac to violet, the tube of the marginal florets usually

5-6 mm 47. degenii 4 Involucre cylindrical; capitula with 5-10 florets; corolla purplish-red, the tube of the marginal florets usually

7-12 mm 48. orientalis 1 Perennial or biennial with a monopodial or sympodial stock,

or a taproot

5 Plant usually with a terminal, central, persistent leaf-rosette; flowering stems developed laterally from axils of previous season’s rosette-leaves; stock usually monopodial; leaves always undivided

6 Lower cauline leaves subcoriaceous, glabrous, ± ciliate

4. sarajevensis 6 Lower cauline leaves membranous, usually hairy

7 Lower cauline leaves narrowly elliptical to suborbicular,

acute; base of stems usually with soft hairs 1. drymeia 7 Lower cauline leaves narrowly elliptical to lanceolate,

acuminate; base of stems glabrous or with ±rigid hairs

8 Stems glabrous below; upper internodes with soft hairs not more than 1 mm; stock monopodial 2. gussonei 8 Stems with ±rigid hairs often more than 1 mm; stock sometimes sympodial 3. arvemensis 5 Plant without a terminal, central, persistent leaf-rosette, sometimes with lateral leaf-rosettes; flowering stems developed terminally; stock sympodial or plant with a taproot; leaves sometimes pinnate.

9 Biennial with a thick taproot; cauline leaves subamplexicaul; flowers whitish 44. tatarica 9 Perennial and with a sympodial stock, rarely biennial; leaves not subamplexicaul; flowers not whitish

10 Basal rosette-leaves + pubescent and hispid with yellowish setae

11 Peduncles eglandular; corolla bluish-lilac 8. basaltica 11 Peduncles glandular; corolla purple

12 Perennial not more than 80 cm; cauline leaves broadly or narrowly lanceolate, undivided or sublyrate with

1-4 lateral lobes 16. dinarica 12 Biennial or short-lived perennial up to 150 cm; cauline leaves ovate-lanceolate, undivided or pinnate with up to 8 lateral lobes 17. lucana 10 Basal rosette-leaves not pubescent and without yellowish setae

13 Allcaulineleavesundivided(anduniformlysoinallplants of a population), usually widest at or below the middle

14 Lowerinternodeswith numeroushairsmore than 1 mm (often also with shorter hairs)

15 Leaves green

16 Stem with soft hairs 7. nevadensis 16 Stem hispid, at least below

17 Upper cauline leaves attenuate or rounded towards the base; calyx 8- to 16-awned 3. arvemensis 17 Upper cauline leaves cuneate, subauriculate or cordate-amplexicaul; calyx usually 8-awned

5. dipsacifolia 15 Leaves ± greyish-white, especially beneath

18 Lower internodes densely long-villous, sometimes also sparsely pubescent; leaves ± entire 15. magnifica 18 Lower internodes short-villous or tomentose, and densely pubescent

19 Leaves narrowly lanceolate, entire to subserrate, appressed-hirsute above 19. baldensis 19 Leaves ovate to broadly lanceolate, crenate-serrate, sparsely pubescent above

20 Upper cauline leaves subcordate at base; peduncles usually glandular 18. subcanescens 20 Upper cauline leaves cuneate to rounded at base; peduncles usually eglandular 24. norica 14 Lower internodes glabrous, or(rarely) with hairs less than 1 mm

21 Lower cauline leaves ± hispid, mostly ± dentate, usually less than 5 times as long as wide

22 Fruit 5-6 x 2-2-5 mm, oblong-ovoid; lower internodes usually hispid 5. dipsacifolia 22 Fruit c. 4-5 x 1 -5-2 mm, cylindrical; lower internodes glabrous 6. ressmannii 21 Lower cauline leaves glabrous or subglabrous, often subentire, usually more than 5 times as long as wide

23 Peduncles glandular

24 Upper cauline leaves broadly cuneate to rounded at base; lower cauline leaves usually 6-9 times as long as wide; calyx usually 8-awned 12. longifolia 24 Upper cauline leaves often cordate and amplexicaul at base; lower cauline leaves usually 4-6 times as long as wide; calyx usually 8- to 12-awned

14. midzorensis 23 Peduncles eglandular

25 Capitula 2-2-5 cm in diameter, 5-9 on each stem 13. pancicii 25 Capitula 2-5-5 cm in diameter, fewer than 5 on each stem

26 Corolla pink 11. salvadoris 26 Corolla bluish-lilac or -violet

27 Lower cauline leaves linear-lanceolate, 6-10 times as long as wide; upper cauline leaves with narrowly cuneate base 10. godetii 27 Lower cauline leaves elliptic-lanceolate, 4^-7 times as long as wide

28 Middle cauline leaves broadly cuneate to rounded at base 9. foreziensis 28 Middle cauline leaves ± amplexicaul and subcordate at base 8. basaltica 13 Upper cauline leaves lyrate or pinnate (at least in some plants of each population); undivided leaves usually widest at or above the middle

29 Leaves± greyish-tomentose, or rarely densely hirsute beneath; terminal lobe of divided cauline leaves usually as long as the divided part

30 Peduncles usually eglandular

31 Upper cauline leaves narrowly lanceolate, usually sublyrate with 1-4 lateral lobes 25. albanica 31 Upper cauline leaves broadly lanceolate, usually lyrate-pinnate with (2-)4-12 lateral lobes

32 Corolla pale yellow 30. kitaibelii 32 Corolla pink, red, violet or lilac

33 Leaves densely hirsute; upper cauline leaves narrow at base 26. velebitica 33 Leaves tomentose-villous; upper cauline leaves broadly cuneate to subcordate at base

34 Corolla bright red; upper cauline leaves often subcordate at base 22. persiana 34 Corolla pale lilac to violet; upper cauline leaves broadly cuneate to rounded at base

35 Lower leaves lanceolate-acuminate; corolla pale lilac 23. carinthiaca 35 Lower leaves oblong-ovate; corolla violet 24. x norica 30 Peduncles usually glandular

36 Corolla, at least in some plants of each population, pale yellow

37 Robust; capitula usually 3-4 cm in diameter 30. kitaibelii 37 Slender; capitula usually 2-3 cm in diameter 31. ambigua 36 Corolla purple,violetorlilac

38 Cauline leaves usually with 4-10 lateral lobes and a terminal lobe usually shorter than the divided part

39 Terminal lobe of cauline leaves ovate-lanceolate; corolla reddish-purple 27. mollis 39 Terminal lobe of cauline leaves narrowly lanceolate; corolla purplish-violet 28. calycina 38 Cauline leaves usually with 2-6 lateral lobes and a terminal lobe usually about as long as the divided part

40 Basal leaf-rosettes usually absent at anthesis

24. x norica 40 Basal leaf-rosettes present at anthesis

41 Leaves densely velutinous with rather short hairs above; capitula mostly 2-3 cm in diameter; corolla purple 20. velutina 41 Leaves sparsely velutinous with long hairs above; capitula mostly 3-4 cm in diameter; corolla pinkish-lilac 21. transalpina 29 Leaves green, hirsute or pubescent beneath; terminal lobe of divided cauline leaves usually shorter than the divided part

42 Calyx(7-)9-to 18 -awned or- dentate; divided lower cauline leaves with lanceolate terminal lobe scarcely longer than lateral lobes; rosette-leaves divided

43 Peduncleseglandular

44 Leaves uniformly pubescent, the lateral lobes usually

3-2-5 mm wide 38. pectinata 44 Leaves pubescent on margin and veins, hirsute or subglabrous elsewhere, the lateral lobes usually

2-3-5 mm wide 39. clementii 43 Peduncles glandular

45 Lateral leaf-lobes linear-lanceolate, usually 2-3-5

mm wide 40. adriatica 45 Lateral leaf-lobes linear, 0-5-1 -5 mm wide

41. dalmatica 42 Calyx (6-)8- to 10-awned; divided lower cauline leaves with ovate, ovate-lanceolate or rhombic terminal lobe usually ± longer than lateral lobes; rosetteleaves usually undivided

46 Leavessubcoriaceous,sparselysetoseorglabrescent, shiny; peduncles usually eglandular

47 Stem hispid or subglabrous; lower leaves ovatelanceolate; calyx up to ll(-15)-awned

42. travnicensis 47 Stem usually subglabrous; lower leaves narrowly lanceolate; calyx 8-awned 43. fleischmannii 46 Leaves membranous and ± densely hirsute, if± sub- coriaceousand glabrescentthen peduncles glandular

48 Corolla, at least in some plants of each population, dark red, pale pink or pale yellow CLXVII DIPSACACEAE 49 Corolla, at least in some plants of each population,

dark red; flowering stems usually without green basal leaves at anthesis, and with evenly distributed cauline leaves 32. macedonica

49 Corolla pale yellow or pale pink; flowering stems usually with green basal leaves at anthesis, and with cauline leaves confined to the lower half of stem

50 Robust; corolla usually pale yellow; capitula usually 3-4 cm in diameter 30. kitaibelii

50 Slender; corolla pale yellow or pale pink; capitula usually 2-3 cm in diameter 31. ambigua 48 Corolla in all plants of a population bluish-violet,

lilac or purple

51 Leaves finely papillose, nearly glabrous, sub-

coriacous; densely caespitose chasmophyte

35. rupicola

51 Leaves hirsute, pubescent or glabrescent, not papillose, usually membranous; not caespitose

52 Corolla predominantly bluish-violet to lilac;

underground stolons usually present 29. arvensis

52 Corolla predominantly purple; underground stolons absent

53 Usually biennial; capitula often 1-8-2-5 cm in diameter 33. visianii

53 Usually perennial; capitula usually more than 2-5

cm in diameter

54 Upper cauline leaves usually with 4-6 lateral lobes and a cuspidate terminal lobe about as long as the divided part 37. illyrica

54 Upper cauline leaves usually with more than 6

lateral lobes and a non-cuspidate terminal lobe shorter than the divided part

55 Terminal lobe of upper cauline leaves lanceolate-rhombic; stem usually more than 20

cm, finely hirsute 34. purpurea

55 Terminal lobe of upper cauline leaves ovatesuborbicular; stems usually not more than

20 cm, coarsely hirsute 36. subscaposa