Download DIVeRSITy Of THe MARTAGON LILy (LILIUM MARtAgON L.) IN

issn 1392-3714, e-issn 2345-0215
Diversity of the martagon lily
(Lilium martagon L.) in Latvia
Antra Balode
Latvia University of Agriculture, Faculty of Agriculture, Institute of Agrobiotechnology,
Liela 2, Jelgava, LV 3001, Latvia; e-mail: [email protected]
Abstract
The genus Lilium L. includes approximately 100 species distributed throughout
the cold and temperate parts of the northern hemisphere. One of the best known
of wild lilies is the martagon lily (L. martagon L.). Of all the lily species, it is
distributed across the largest growing area – from western Portugal through
Europe and Asia. To evaluate the natural habitat of the martagon lily, research
was carried out in locations at Aizkraukle, Ventspils, Kuldiga and Tukums region
in Latvia. Research results indicate that diversity in genotypes varies according
to location and distribution. The individuals of each biotope were characterized
by stable inherited traits: color, shape and flower diameter, stem length and bulb
color. The color of flowers was basically consistent, but there were variations in
intensity from pale to bright violet-pink. Spotting was also extremely variable,
ranging from a few small spots in the centre to spotting that covered the entire
flower to the tips of the tepals. Diameter of flower varied between of 3.5 to 6.0 cm.
The height of these plants varied up to 114-130 cm. The flowering time was from
mid June to the middle of July and seeds ripen in September. Wild martagon lilies
found in these regions were growing in calcareous soils among shrubs in river
valleys and semi-shaded deciduous forests. Bulbs were growing to about 10 to
20 cm under the ground.
Key words: phenological observations, variability, wild species.
Introduction
It has been discovered that the first Lilium martagon L. of the north
blossomed in Bergen, Norway, in 1597, and was introduced to Sweden
in 1795 (Lundquist, 2005). Many of the Scandinavian populations are
restricted in size and geographically isolated from one another. The
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largest population in Sweden (at the estate of Ulfåsa) and perhaps
in Scandinavia has, however, more than 25 thousand individuals
(Persson, 2001). It is mentioned that L. martagon had probably been
first brought to England as a garden plant in 1550-1565. It has been
introduced to North Germany and to Britain simply as an ornamental
plant: to Britain after 1548 but shortly before 1568 (Lundquist, 2005).
L. martagon in Latvia was discovered in 1839 (Andrušaitis, 1985).
The species of the genus Lilium is classified and the most
authoritative is the classification of Harold F. Comber. In 1949, a revised
subclassification of the genus Lilium was published. H. F. Comber
considers physical features such as flowers, seeds, the type of
germination, arrangement of the leaves and the form and growth habit
of the bulb. He also gives importance to geographical distribution and
to the evolutionary relationships of species and groups. This resulted
in seven part categorization (Comber, 1949). The martagon section
comprises such species: L. martagon L., L. distichum Nakai ex Kamib.,
L. hansonii Leichtlin ex D. D. T. Moore, L. medeoloides A. Gray and
L. tsingtauense Gilg (Comber, 1949; McRae, 1998). Despite the great
variation in form and disposition of the flower there can be no doubt
that the species are very closely related. Whorled leaves and hypogeal
seed germination indicate this relationship.
In the wild, martagons have the ability to survive for years by
being tucked under trees and using bulbs, rhizome, stolon or some
other device to protect food reserves. It is a selfincompatible diploid,
2n=24, which propagates predominantly by seed-set and only very
rarely by bulbils (Lundquist, 1991). It takes 7-8 years to flower from
seed. L. martagon is cross-pollinated and, at the end of the summer,
many light-weight seeds are produced and dispersed by wind. The
flowers have an unpleasant sweetly fragrant odour – especially at night
to attract the night-moth for pollination. The whorled leaves give the
plant the ability to secure most of the available light most efficiently;
therefore, this situation suits L. martagon admirably.
The literature gives the following countries where L. martagon may
be found: Portugal, Spain, France, Switzerland, Germany, Austria,
Italy, Czech, Hungary, Albania, Greece, Poland, Romania, Bulgaria,
Turkey, Britain, Belgium, Holland, Denmark, Norway, Sweden,
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Antra Balode
Finland, Russia – central, southwestern and southeastern regions,
and the Baltics. It is mentioned that in the Baltic countries its most
northerly site is in Latvia (Fox, 1987; Andrušaitis, 1985). There are
conflicting views that the northern latitude is in Estonia (Lundquist,
2005). L. martagon is found in Lithuania near Vilnius and Trakai in
the forest as well as the area of Kėdainiai, Ramygala. L. martagon is an
endangered species in Poland (Lundquist, 2005). East of the Urals in
Asia the lily stretches to 124°E on the Rivers Vilyui and Lena. Its most
northerly station in Siberia is 68°40ʹN on the lower reaches of the River
Yenisei. On the south, it is found in northern parts of Mongolia, in the
Caucasus and the western area of northern Turkey in Asia (Fox, 1987).
Over all this vast range it is never far from woodlands. M. V. Baranova
says the martagon lily is thriving within forest glades, on the edge of
the forest and among bushes on herb-covered meadows (Баранова,
1990). It is found that L. martagon is a more polymorphic plant in the
wild comprised with other lily species (McRae, 1998).
Over recent decades, under the impact of human activity the
L. martagon has decreased (Kedra, Bach, 2005) – despite strict
protection. Currently a certain role in the protection of endangered
species has been providing by law in the Czech Republik (Štrůtka, 2009).
It establishes certain ways of how to safeguard the future of endangered
species. L. martagon has been included in the previous – but not the last
issue of the Red Data Book of Latvia (Andrušaitis, 1985, 2003).
Ranging over vast area, L. martagon includes several distinct forms
which differ from the typical species. L. martagon var. albiflorum Vuk.
grows in the wild in Yugoslavia and Germany, has white flowers spotted
in carmine pink. L. martagon var. album Weston has pure white flowers.
L. martagon var. cattaniae Vis. are more common in Balkans: Albania,
Bulgaria, Greece, Rumania, Turkey, Serbia and Croatia (Fox, 2006). It
is extraordinarily stately with deep wine-colored unspotted flowers
in a large inflorescence. In 1934, L. martagon var. martagon has been
described from Latvian botanist N. Malta, as to where it occurs on the
dolomite hills near the mouth of the river Daugava. It is distinctive by its
height – up to 180 cm and by short hair on the stem. The leaves are 5 cm
wide and the 3 to 10 flowers have strongly reflexed tepals of pale purplered with dark spots and red hair on the tepals (Fox, 1987; McRae, 1998).
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Methods
During the growing seasons of different years (1994, 2001, 2003
and 2006), in locations of natural habitat the following phenological
observations were made: flowering time (days from the beginning of
vegetation till first flower open), plant height (cm), number of flowers
per stem (count), diameter (cm) and form and coloring of the flowers.
From the Institute of Biology of the Latvia University, the initial dates
of natural habitats were obtained (Fig. 1).
Kuksas
Padure
Rīva
Plani
Klintaine
Staburags
Vīgante
Fig. 1. Natural habitats of L. martagon in Latvia
Plants were estimated individually by the following traits: plant
height (cm), number of flowers per stem (count) and diameter of a
flower (cm). The flowering time was when first flower opens.
Results and Discussion
Lily breeders are very interested to find bulbs in the wild, collect
them and use them in breeding. The goal of this research was to
estimate L. martagon distribution in the locations of west of Latvia −
in the regions of Ventspils, Kuldiga and Tukums and in the east – in
Aizkraukle.
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Antra Balode
Significant differences were observed between the L. martagon
found at different habitats – such as variability in some morphological
traits (start of growth, plant height, flower diameter and color, flowering
time and number of flowers per stem) (Table 1). The environment of the
habitat differs regionally. Taking into account the morphological traits,
these martagon lilies were classified into groups. The wild martagon
lilies found on the left bank of the River Daugava near Vigante Park
and Staburags (areas which belong to the region of Aizkraukle) are
considered as one group. Significantly different are the martagon lilies
in Aizkraukle region from those found on other sites. Their flowers are
pale grayish pink with small dots and their start of growth is late April
to early May – with flowering late in June (57 days from the beginning
of vegetation till first flower open) (Table 2). The height of the plant is
up to 105-110 cm. Variability in some morphological traits could be
explained by the impact of genotype and the environment.
66
64
62
Days
60
58
56
54
52
50
48
Vīgante
Staburags
Klintaine
Riva
Padure
Plani
Kuksas
Habitats of L.martagon
Fig. 2. The flowering time of different species and cultivars of lilies on average, in
years 2008 and 2009 (in days)
Wild martagon lilies found in the regions of Ventspils, Kuldiga
and Tukums are similar by phenotypes and are characterized by bright
violet-pink flowers. The height of the plants is up to 114-130 cm. The
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XVII ❧ Botanikos sodo raštai / Scripta Horti Botanici
beginning of their season’s growth is late April and the time of flowering
is late June (54 days) (Fig. 2). If a population is confined to the vicinity
of some old or abandoned site, its habitat is questionable. It may have
originated from domesticated species. The habitats in Padure (in the
region of Kuldiga), Plani and Kuksas (in the region of Tukums) would
probably be situated in areas of old parks. Their soils are generally
clayey. The martagon lilies grows on grasslands and among shrubs, but
cannot be found in deep forests, because sunlight is necessary for its
growth. The environment of the habitat differs regionally. In Kuksas
(in the region of Tukums) L. martagon habitat is found in grassland
in the vicinity of a old manor house. The habitat is abundant with
weeds, ferns and climbing weeds (Aegopodium podagraria L., Dactylis
glomerata L., Vicia sepium L.) and surrounded by shrubs and trees:
maple – Acer platanoides L., oak – Quercus robur L., linden – Tilia
cordata Mill., hazelnut – Corylus avellana L., alder – Alnus glutinosa
(L.) Gaertn.). It is all a surrounded area – a private park. In Kuksas
there were found variable plants – most heavily spotted clone; some
with a few spots.
The habitat in Padure (Kuldiga) is situated in an area of an old palace
and a park. Significant differences between L. martagon in Padure
habitat and other sites (Kuksas, Plani and Riva) were also observed.
The inflorescence form of L. martagon is typical in the raceme.
Flowering stems range in height from 114 to 130 cm. This factor seems
to depend upon the habitat. In an open, sunny location, the stems
are fairly consistent 110-120 cm in height. The extremes in height
seem to appear in more protected and somewhat shaded locations or
where they must compete with vigorous vegetation. In Padure habitat,
L. martagon varies significantly in flower diameter and color, and also
in plant height. In Padure location, generally the color of the flowers is
more intensely pink and with stronger spotting than at any other site
in Latvia.
The valley of the small River Riva form the Riva L. martagon habitat
of shrubs and deciduous forest (Ulmus laevis Pall. and Aegopodium
podagraria L., Gagea lutea L., Dryopteris filix-mas (L.) Schott, Pteridium
aquilinum (L.) Kuhn., Urtica dioica L.). The leaves of this martagon
lily are formed in whorls ranging from 3 to 4 in each whorl of 5 to
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Antra Balode
14 leaves. The width of the individual leaf varies from 2.5 to 4 cm. In
the habitat of Riva, martagon lilies were found with wider leaves than
elsewhere.
Wild martagon lilies in the region of Aizkraukle on the right bank
of the River Daugava, near Klintaine, are having distinct characteristics
when compared with other L. martagon specimens – distinct parameters;
lighter violet-pink flowers with exceedingly fine and scattered dots; up
to 90 cm stems and blossoms of greater size. They also are late starters –
with growth period from early May and blossoming through early July
(63 days from the beginning of vegetation till first flower open).
It has been found that martagon lilies grow mostly in semi-shade
(light woodland) and in calcareous soils – preferably with roots in the
shade. Generally flowering spans from mid June to the middle of July
and seeds ripen throughout September. Table 1 shows the comparison
of characteristics of L. martagon from different habitats. The results
concur with findings in the literature that L. martagon can be very
variable in morphological characters like color of flowers, the number
of flowers per stem, the intensity of spotting on the tepals and the
pubescence of flower buds, also within a single population (Feldmaier,
McRae, 1982).
Conclusions
1. It has been found that in Latvia L. martagon varies by genotype
in diverse natural habitats. If a population is confined to the vicinity of
a village, the habitat is questionable. It may have originated from some
cultivated species.
2. The individuals of each biotope were characterized by stable
inherited traits: color, shape and flower diameter, stem length and bulb
color. The color of flowers is basically consistent, but there are variations
as well as intensity in color. Spotting is also extremely variable, ranging
from a few small spots in the centre to spotting that covers the entire
flower to the tips of the tepals.
3. In the natural growing area the flowering started in 54 to 63 days
from the beginning of vegetation.
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Shape
Stem: Shade
Color
Bulb: Whole
shape
Shape of scale
oblong-ovate
light green
Direction
Leaves: Color
fragrant
turk-capshaped
pendulous
dark green
fragrant
turk-capshaped
pendulous
dark green
Klintaine
light
violet-pink
fragrant
turk-capshaped
pendulous
medium
green
elliptic
light green
Region
Ventspils
Riva
bright
violet-pink
narrowlanceolate
yellow
lanceolateovate
yellow
lanceolateovate
yellow
narrowlanceolate
yellow
oblon-ovate
ovate
purplish
light brown
brown
elongate-ovate elongate-ovate elongate-ovate globose
fragrant
turk-capshaped
pendulous
dark green
Scent
Shape
Pollen
Spotting
Aizkraukle
Staburags
pale greyish
pink
strong
violet-red
orange
small
reddish-brown
bright orangebrown
fragrant
turk-capshaped
pendulous
light green
Tukums
Plani
Kuksas
bright
medium
violet-pink violet-pink
elongateovate
lanceolate-ovate lanceolateovate
yellow
yellow
elongate-ovate
narrowlanceolate
yellow
elongate-ovate
fragrant
turk-capshaped
pendulous
medium
green
oblong-ovate
elliptic-ovate lanceolate
purplish brown light brown light green
fragrant
turk-capshaped
sub-pendulous
dark green
Kuldiga
Padure
vary from
brighter to paler
violet-pink
small
strong
small
strong
strong reddishreddish-brown reddish-brown reddish-brown violet-red
brown
yellow
bright orange bright orange orange-brown light-orange
Vīgante
Flowers: Color pale greyish
pink
Habitats
Characteristics
Table 1. Results of qualitative parametres of L. martagon in different habitats
XVII ❧ Botanikos sodo raštai / Scripta Horti Botanici
7.0
4.0±0.3
3.5±0.2
6.0±0.6
L. martagon (Padure)
L. martagon (Plani)
L. martagon (Kuksas)
21.1
7.0
9.3
9.3
3.5±0.2
L. martagon (Riva)
9.3
L. martagon (Klintaine) 4.0±0.3
7.0
V%
3.5±0.2
Mean ±Sx,
cm
Diameter of
flower
L. martagon (Staburags) 4.0±0.3
L. martagon (Vīgante )
Habitats
10.0
13.0
7.5
V%
115.0±7.5
114.0±6.9
8.5
14.0
130.0±10.2 20.4
130.0±10.2 20.4
90.0±4.8
110.0±7.8
105.0±5.5
Mean ±Sx,
cm
Plant height
16.0±1.5
15.0±1.3
23.0±3.7
10.0±1.7
5.0±1.6
10.0±1.7
5.0±1.6
Mean ±Sx,
number
30.2
38.3
61.5
20.2
58.5
20.2
58.5
V%
Number of flowers
Mean
±Sx, cm
10.0±0.4
15.0±0.4
15.0±0.4
18.0±0.3
10.5±0.4
12.0±1.7
17.0
28.4
28.4
36.9
19.0
16.0
16.0
V%
Leaves length,
cm
12.0±1.7
Table 2. Results of quantitative parametres cross of Lilium martagon in different habitats
2..5±0.6
3.5±0.4
3.5±0.4
4.0±0.3
2.0±0.3
3.0±0.2
3.5±0.4
Mean ±Sx,
cm
18.0
14.0
14.0
21.9
17.0
24.6
14.0
V%
Leaves width,
cm
Antra Balode
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XVII ❧ Botanikos sodo raštai / Scripta Horti Botanici
References
Andrušaitis, G. 1985. Sarkanā grāmata. Retās un iznīkstošās dzīvnieku un augu
sugas. Zinātne, Rīga, 525 p.
Andrušaitis, G. 2003. Latvijas Sarkanā grāmata. 3. daļa. Vaskulārie augi.
Zinātne, Rīga, 691 p.
Comber, H. F. 1949. A New Classification of the Genus Lilium. In: The Lily
Yearbook of the North American Lily Society, Vol. 13, p. 86–105.
Feldmaier, C.; McRae, J. 1982. Lilies. Eugen Ulmer, Stuttgart, 246 p.
Fox, D. 1987. Growing Lilies. London: Christopher Helm Publishers Ltd. 264 p.
Fox, E. E. 2006. Martagon Lilies: Old World, Whorled-Leaf Lilies. Millet, Alberta,
Canada, 174 p.
Lundquist, K. 1991. Four-Locus S-gene Control of Self-Incompatibility Made
Probable in Lilium Martagon (Liliaceae). Hereditas, Vol. 114, p. 57−63.
Lundquist, K. 2005. Lilium martagon L. Doctoral dissertation, Swedish
University of Agricultural Sciences, Alnarp. Acta Universitatis Agriculturae Sueciae,
Vol. 19, p. 292−294.
Kedra, M.; Bach, A. 2005. Morphogenesis of Lilium martagon L. Explants in
Callus Culture. Acta Biologica Cracoviensia. Series Botanica, Vol. 47/1, p. 65−73.
McRae, E. A. 1998. Lilies. Portland, Oregon: Timber Press, 391 p.
Persson, H. 2001. Estimating Genetic Variability in Horticultural Crop Species at
Different Stages of Domestication. Doctoral thesis. Swedish University of Agricultural
Sciences, Alnarp, 290 p.
Štrůtka, P. 2009. News from the Old Martagon Country. In: Mitchell, A. (ed.).
Lilies and Related Plants 2009-2010. The Royal Horticultural Society Lily Group.
London, p. 54−59.
Баранова, М. В. 1990. Лилии. Ленинград: Агропромиздат, 383 с.
Miškinės lelijos (Lilium martagon L.) įvairovė latvijoje
Santrauka
Lilium L. genties augalai, kurių yra apie 100 rūšių, yra paplitę šiaurinio pusrutulio šaltojo ir vidutinio klimato zonose. Viena žinomiausių laukinių lelijų yra
miškinė lelija (Lilium martagon L.). Jos arealas didžiausias iš visų lelijų rūšių –
nuo vakarų Portugalijos per visą Europą ir Aziją. Miškinės lelijos natūralioms augavietėms įvertinti buvo atlikti tyrimai šiuose Latvijos regionuose: Aizkrauklėje,
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Antra Balode
Ventspilyje, Kuldygoje ir Tukume. Tyrimo rezultatai parodė, kad genotipo įvairovė priklauso nuo augimo vietos. Kiekvieno biotopo augalai pasižymi šiomis paveldimomis savybėmis: žiedo spalva, forma ir dydis; stiebo aukštis ir svogūno spalva.
Žiedų spalva buvo gana pastovi, bet skyrėsi jos intensyvumas – nuo šviesios iki
ryškios violetiškai rožinės. Žiedo taškuotumas taip pat varijavo – nuo kelių mažų
dėmelių centre iki daugybės dėmių ant viso žiedo. Žiedo skersmuo – nuo 3,5 iki
6,0 cm. Šie augalai gali būti 114–130 cm aukščio. Žydėjimo trukmė – nuo birželio
vidurio iki liepos vidurio; sėklos sunoksta spalį. Miškinė lelija auga kalkingoje
dirvoje, tarp krūmų, upių slėniuose, lapuočių miškuose pusiau pavėsyje. Jos svogūnai auga 10–20 cm gylyje.
Reikšminiai žodžiai: fenologiniai stebėjimai, įvairovė, laukinės rūšys.
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