L I L I U M PYRENAICUM - A C O M P L E X SPECIES

Victoria Matthews

The writing of an account of Lilium in cultivation for the forthcom- ing European Garden Flora, combined with some years of observ- ing European lilies both in the wild and in cultivation, have led me gradually to the conclusion that the situation given in Flora Europaea 5 is not entirely satisfactory and should be emended. These observations and conclusions are based also on the exami- nation of many specimens from European herbaria.

At present five turk’s-cap lilies are recognized in Europe (Mat- thews, 1980)*. L. martagon L. is a well-known and distinct species with flowers which vary from white, through pink to dark purplish-red; its geographical range extends through N Asia. The other four are endemic to Europe: L. pyrenaicum Gouan comes from the Pyrenees and N Spain, L. Pomponium L. from France and Italy (l lari t ime Alps, W Ligurian Apennines), L. carniolicum Bernh. ex I\*. Koch (including the associated species Ll . albanicum, bosniacum andjankae) ranges from the SE Alps through Yugoslavia to the Balkans, and L. chalcedonicum L. (Syn. L. heldreichii Freyn) is native from S Albania to S and W Greece. The flowers of all these lilies produce a characteristic rather unpleasant smell. Historically there has been considerable confusion between these European endemics and this confusion is reflected in their early synonymy and classification.

Lilium chalcedonicum is easily distinguished from the other turk’s-cap lilies by its leaves, which are spreading on the’ lower part of the stem and adpressed on the upper part. The orange-red flowers are unspotted, although there is a variety, var. maculatum Stoker, with deep purple or maroon spots which is known only in cultivation. L. pomponium produces flowers of a brilliant red which set it apart from the other three lilies, as do its generally narrower leaves, 2-5 mm wide, which have only 1-3 veins. L. pyrenaicum is occasionally found with leaves as narrow as 3 mm wide, and there is a form of L. bosniacum (f. stenophyllum Beck) with leaves which vary from 3-10 mm wide, so this character, although useful, is not absolutely dependable. However, the two latter taxa always have

*I am excluding L. rhodopaeum Delip. from Greece and Bulgaria, and L. monadelphum Bieb. from the Crimea from this turk’s-cap group because they possess flowers whose perianth segments are much less tightly recurved.

36

3 or more veins per leaf, whereas the leaves of L. pomponium may have only 1 vein. In general the leaves of L. pomponium do not extend as far up the main stem as they do in the other lilies, tending to stop a little way below the inflorescence.

The distinctions between L. pyrenaicum and L. carniolicum are less marked. Both orange-red- and yellow-flowered plants occur in both species. O n the whole the leaf-veins of L. pyrenaicum are glabrous beneath (there are rare specimens which possess sparse hairs). Those of L. carniolicum are downy beneath (except for the variant with glabrous veins, L. albanicum). Just how similar these two lilies can look is demonstrated by the photographs at the bottom of p. 167 in Rix & Phillips, The Bulb Book (1981).

At this point, L. ponticum C. Koch from NE Turkey and Soviet Georgia must also be considered. L. ponticum can have yellow or orange flowers and the leaf-veins are downy beneath. It differs from both L. pyrenaicum and L. carniolicum in having stamens with papillose filaments; those of the other lilies are smooth. It differs also in lacking warts on the inside of the perianth segments. Davis & Henderson in Flora of Turkey vol. 8 (in press) and in Notes R.B.G. Edinb. 41: 53 , 1983, realizing the similarities between L. ponticum and L. carniolicum, treat L. ponticum as a subspecies of L. carniolicum. Their study of Turkish and Caucasian material in- cluded a consideration of L. carniolicum, but understandably did not extend as far as the Pyrenees. It is a drawback of writing a regional flora that there is no time to step far outside the geograph- ical limits of that flora, if it is to be completed within a sensible time. Davis & Henderson did, however, suggest that further studies on other related European lilies ‘might lead to a different circum- scription’. When they wrote this, Grey-Wilson had already implied (Curtis’s Bot. Mag. 184, 1: 9-10,. 1982) that it might be best to regard the geographically separated Ll. pyrenaicum, carniolicum and ponticum as subspecies: L. pyrenaicum is the earliest name. This suggestion coincided with my own conclusions and I now propose to divide L. pyrenaicum into three subspecies; subsp. pyrenaicum, subsp. carniolicum and subsp. ponticum. Subsp. carniolicum will be further divided into four varieties including the type variety. Within subsp. ponticum, following Davis & Henderson I recognize var. ponticum and var. artvinense; unfortunately the earlier name of L. pyrenaicum necessitates different combinations from those pub- lished for the account in the Flora of Turkey (see above).

37

The other closely related species frorp NE Turkey, L. ciliatum Davis, is distinct enough for its specific status to be maintained. The leaves are glabrous except for a long-ciliate margin and there are long hairs on the tips of the cream to pale yellow perianth segments. The segments, as in L. ponticum are marked and spotted with purplish-brown and lack warts, but the filaments are smooth.

SYNOPSIS OF THE INFRASPECIFIC DIVISION OF LILIUM PYRENAICUM

L. pyrenaicum Gouan subsp. pyrenaicum subsp. carniolicum (Bernh. ex W. Koch) Matthews

var. carniolicum var. albanicum (Griseb.) Matthews var. bosniacum (Beck) Matthews var. jankae (Kerner) Matthews

subsp. ponticum (C. Koch) Matthews var. ponticum var. artvinense (Misch.) Matthews

Lilium pyrenaicum Gouan, Obs. Bot. 25 (1773). Stem roots often present. Stem 15-135 cm, green, sometimes with sparse

purple spots, glabrous. Leaves scattered, 3-15 cm long, 0.3-2 cm wide, linear to lanceolate or narrowly elliptic with 3-15 veins, margin very minutely ciliate, veins glabrous to densely downy beneath. Flowers up to 12, turk’s-cap or martagon type (with the perianth segments rolled strongly back), nodding, faintly to strongly and unpleasantly scented. Perianth segments 3-7 cm long, 0.4-1.3 cm wide, yellow, orange or red with dark purplish or reddish-brown spots and lines inside especially towards the base, and usually with purplish warts. Filaments greenish or yellowish, smooth or papillose. Anthers 4-10 mm, with orange, red or brown pollen.

DISTRIBUTION: Pyrenees and N Spain, SE Alps, Balkans, NE Twkey and Soviet Georgia.

KEY TO THE SUBSPECIES OF LILIUM PYRENAICUM

1. Leaf-veins glabrous beneath . . 2 Leaf-veins with at least some hairs beneath . . 3

2. Flowers spotted . . A. subsp. pyrenaicum Flowers unspotted . . B. subsp. carniolicum

3. Perianth segments bearing warts towards the base inside; filaments smooth . B. subsp. carniolicum

Perianth segments lacking warts; filaments papillose C. subsp. ponticum

A. subsp. pyrenaicum Slem 30-135 cm, green with sparse purple spots. Leaves 7-15 cm long,

0.3-2 cm wide, with 3-15 veins which are glabrous beneath. Flowers up to 12. Perianth segments 4-6.5 cm long, yellow to greenish-yellow with dark purple spots and lines inside and purplish warts. Filaments smooth.

38

DISTRIBUTION: SW France, N Spain. Occasional plants have been found in the wild in which the leaf-veins

are very slightly downy beneath. A form with orange-red flowers is found in N Spain and is known as

f. rubrum (Stoker, 1938).

B. subsp. carniolicum (Bernh. ex Mi. Koch) Matthews, comb. et stat. nov. L . carniolicum Bernh. ex W. Koch, Syn. F1. Germ. 708 (1837); L . pomponium

L. var. carniolicum (Bernh. ex W. Koch) Fiori in Fiori & Paoletti, F1. Anal. Ital. 1: 183 (1896). Stem to 120 cm, green. Leaves 3-1 1 cm long, 0.4-1.7 cm wide with 3-9

veins which are glabrous or downy beneath. Flowers up to 6 (sometimes

to 12 in cultivation). Perianth segmenls 3-7 cm, red, orange or yellow, often spotted with brownish-purple, and with purplish or rarely yellow warts. Filaments smooth.

DISTRIBUTION: SE Europe.

KEY T O THE VARIETIES OF SUBSP. CARNIOLICUM

1. Leaf-veins glabrous beneath; flowers unspotted var. albanicum Leaf-veins downy beneath, sometimes irregularly so; flowers usually

. var. carniolicum

Leaf-veins less densely downy beneath, sometimes irregularly SO;

3. Leaf-veins evenly downy beneath; flowers spotted or unspotted var. jankae

Leaf-veins irregularly downy beneath; flowers spotted . . var. bosniacum

var. carniolicum Stem to 120 cm. Leaf-veins densely downy beneath. Flowers red or

DISTRIBUTION: SE Alps (Austria and Italy), N Yugoslavia.

var. albanicum (Griseb.) Matthews, comb. nov. L. albanicum.Griseb., Spicil. F1. Rumel. 2: 385 (1846); L. carniolicurn Bernh.

ex W. Koch var. albanicum (Griseb.) Baker in J. Roy. Hort. SOC. 4: 46 (1873); L. chalcedonicum L. var. albanicum (Griseb.) Ascherson & Graeb- ner, Syn. Mitteleur. F1. 3: 183 (1905); L. carniolicum Bernh. ex W. Koch subsp. albanicum (Griseb.) Hayek, Prodr. F1. Balcan. 3: 62 (1932). Stem to 40 cm. Leaf-veins glabrous beneath. Perianth segments usually

DISTRIBUTION: SW Yugoslavia, Albania, NW Greece. Specimens have occasionally been found which, like subsp. ponticum,

var. bosniacum (Beck) Matthews, comb. nov.

Wien 2: 47 (1887); L. bosniacum (Beck) Beck ex Fritsch in Mitt. Naturw. Ver. Steierm. 45: 163 (1908). Stem to 80 cm. Leaf-veins irregularly downy beneath. Perianth segments

DISTRIBUTION: C Yugoslavia. As mentioned previously, there is a narrow-leaved form which was

published as L. carniolicum var. bosniacum f. stenophyllum by Beck in Glasn. Zemaljsk. Muz. Bosni Hercegovini 15: 206 (1903), and which occurs locally within the range of var. bosniacum. In view of the variation in leaf width within other parts of the ‘Ryrenaicum’ complex, it does not seem to be worthy of recognition.

spotted . . 2 2. Leaf-veins densely downy beneath; flowers red or orange .

flowers usually yellow . . 3

orange, spotted.

yellow, occasionally orange, unspotted.

have papillose, rather than smooth filaments.

L. carniolicum Bernh. ex W. Koch var. bosniacum Beck in Ann. Nat. Mus.

usually yellow, occasionally orange-red, spotted.

Var. bosniacum has not been introduced into European gardens.

40

Map showing the disrribution of the varieties of subsp. carniolicum Smmm var. carniolicum 0-0-0- var. albanicum

- - - - var. jankae w v w w I L var. bosniacum

var. jankae (Kerner-) Matthews, comb. et stat. nov. L. jankae Kerner in Ost. Bot. Zeitschr. 27: 402 (1877); L. carniolicum

Bernh. ex W. Koch subsp. jankae (Kerner) Ascherson & Graebner, Syn. Mitteleur. F1. 3: 181 (1905); L. carniolicum Benh. ex W. Koch f. sulphureum Pamp. in Nuovo Giorn. Bot. Ital. 12: 151 (1905). Stem to 80 cm. Leafveins evenly downy beneath. Perianth segments yellow,

spotted or unspotted. DISTRIBUTION: NW Italy, NW Yugoslavia, W Bulgaria, C - SC

Romania. This division of subsp. carniolicum into four varities is by no means

clear-cut. Plants which combine characters of var. albanicum and var. bosniacum in having yellow, unspotted flowers but leaf-veins which are slightly downy, have been occasionally found in C Yugoslavia and in the Pindus Mountains of N Greece. Other intermediate plants with yellow, spotted flowers (typical of var. bosniacum or var. jankae) but glabrous leaves are known from C Yugoslavia and C Albania.

The relatively uncommon plants with orange or orange-red flowers which have been assigned to both var. albanicum and var. bosniacum have

41

never been given names, and there seems to be little point in separating them taxonomically from the yellow-flowered plants. They are not known to be in cultivation.

It was mentioned in Flora Europaea 5: 35 that the perianth segments of typical L. carniolicum were 5-6.5 cm long, whereas those of L. albanicum were only c. 3 cm long. Subsequent examination of more specimens has shown that this is not the case; there is a range of size (3-7 cm) within subsp. carniolicum, showing correlation neither with the varieties as delim- ited above, nor with geographical distribution.

C. subsp. ponticum (C. Koch) Matthews, comb. et stat. nov. L. ponticum C. Koch in Linnaea 22: 234 (1849); L. carniolicum Bernh. ex W. Koch subsp. ponticum (C. Koch) Davis & Henderson in Notes R.B.G. Edinb. 41: 53 (1983).

Stem 15-90 cm, green. Leaves 3-8 cm long, 0.8-2 cm wide, with 7-15 veins which are downy beneath. Flowers up to 5, occasionally to 12. Perianth segments 3.5-5.5 cm, deep yellow or deep orange with dense reddish-brown or purplish stripes and spots, but lacking warts. Filaments papillose.

DISTRIBUTION: NE Turkey, USSR (Georgia).

KEY T O THE VARIETIES OF SUBSP. PONTICUM

Perianth segments deep yellow, 7-10 mm wide . var. ponticum Perianth segments deep orange, 5-6 mm wide . .var. artvinense

.

var. ponticum. Stem usually stout. Perianth segments deep yellow, 7-10 mm wide. DISTRIBUTION: NE Turkey.

var. artvinense (Misch.) Matthews, comb. nov. L . artvinense Misch. in Trudy Bot. Mus. Imp. Akad. Nauk 8: 191 (1911);

L. ponticum C. Koch var. artvinense (Misch.) Davis & Henderson in RHS Lily Year Book 33: 21 7 91969); L. carniolicum Bernh. ex W. Koch subkp. ponticum (C. Koch) Davis & Henderson var. artvinense (Misch.) Davis & Henderson in Notes R.B.G. Edinb. 41: 53 (1983). Stem more slender than in var. ponticum and with fewer, less crowded

leaves. Perianth segments deep orange, 5-6 mm wide. DISTRIBUTION: NE Turkey, USSR (Georgia). The distinctions given above between the varieties of subsp. ponticum

apply to cultivated material and to almost all the specimens found in the wild. However, plants with intermediate characters have been collected in NE Turkey.

I am aware that the above name changes may well upset some gardeners and horticulturists and will probably annoy nurserymen. As grown in the gardens of Europe, these lilies are separable and form stable recognizable entities, but it must be realized that the cultivated plants represent only

42

parts of the range of variation present in the wild. When the wild plants are studied it becomes obvious that the species, as previously accepted, are less distinct than the ‘species’ grown in gardens, and overlap in many of the characters which have traditionally been used to separate them. In the classification set out above, familiar names have not been lost; there has merely been a change in status to reflect more closely the relationships within this group of lilies.

Subsp. pyrenaicum is the most commonly seen subspecies in gardens and has been cultivated for many years; it was known to have been grown in Paris by 1598. In Britain it has escaped from gardens and become naturalized in Devon, in N Wales near Port Merion, and in an increasing number of localities. It is easy to grow and requires little attention, its only drawback being its rather unpleasant smell. The form with orange-red flowers is not so frequently seen despite being commercially available.

Subsp. carniolicum has also been in cultivation for many years, at least since 1629 and probably earlier. It is easy to grow but is not seen as often as subsp. pyrenaicum. Var. albanicum and var.jankae are grown less widely, being more or less confined to specialist collections. Var. bosniacum is not in cultivation.

Subsp. ponticum is not difficult to grow but is not widespread in culti- vation. I t was first introduced in 1933 by the plant collector E. K. Balls.

All these lilies thrive in a sunny or partly shaded position and appre- ciate a well-drained soil. They are tolerant of limy conditions.

REFERENCES

Davis, P. H. & Henderson, D. M. (1969). A Revision of Turkish Lilies.

Davis, P. H. & Henderson, D. M. Lilium in Davis, P.H. (ed.), Flora of

Elwes, H. J. (1877-1880). A Monograph of the Genus Lilium. London Grey-Wilson, C. ( 1982). Lilium carniolicum. Curtis’s Botanical Magazine

Grove, A. & Cotton, A.D. (1933-1940). A supplement to Elwes’ Mono-

Matthew, V. A. (1980). Lilium in Tutin, T.G. et al. (ed.), Flora Europaea

Stoker, F. (1938). The Carniolicum group of Lilies. RHS Lily Year Book

Synge, P. M. (1980) Lilies - A Revision of Elwes’ Monograph of the

Turrill. W. B. (1953). The Lilies of the Balkan Peninsula. RHS Lily Year

Woodcock, H. B. D. & Stearn, W.T. (1950). Lilies of the World. London.

RHS Lily Year Book 33: 212-223:

Turkey 8 (in press.)

184: 9-11 and N.S. t. 839.

graph of the Genus Lilium. London.

5 : 34-35.

7: 86-107.

Genus Lilium and its Supplements. London

Book 17: 30-39.

43