Biodiversity in Lilium: A Review

International Journal of Horticulture, 2018, Vol. 8, No.
8, 83-97
http://ijh.biopublisher.ca
Research Article Open Access

Biodiversity in Lilium: A review
M.R. Dhiman , Siddharth Moudgil, Chander Parkash, Raj Kumar, Sandeep Kumar
ICAR-Indian Agricultural Research Institute, Regional Station, Katrain, Kullu-Valley-175129, H.P., India
Corresponding email: mrarjun01@gmail.com
International Journal of Horticulture, 2018, Vol. 8, No. 8 doi: 10.5376/ijh.2018.08.0008
Received: 15 Feb., 2018
Accepted: 22 Feb., 2018
Published: 13 Apr., 2018
Copyright © 2018 Dhiman et al., This is an open access article published under the terms of the Creative Commons Attribution License, which permits
unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
Preferred citation for this article:
Dhiman M.R., Moudgil S., Parkash C., Kumar R., and Kumar S., 2018, Biodiversity in Lilium: a review, International Journal of Horticulture, 8(8): 83-97 (doi:
10.5376/ijh.2018.08.0008)
Abstract Lilium is a subbstantial genus administered throughout temperate and cooler regions of the Northern Hemisphere
consisting approximately 110 species. The genus possess a great genetic diversity in many valuable horticultural traits which is
manifested in flower colour, forms, shape, size, fragrance, resistance to diseases, and many physiological characteristics. Intensive
agricultural practices, climate change and industrialization are having a straight impact on biodiversity. Comprehensive
understanding of the species, including levels and form of genetic variation forms the basis for the successful management and
safeguarding of populations of rare, endangered or threatened species. The biodiversity become important components of different
ecosystems. Use of single new improved varieties of crops for large areas is a big threat for crop biodiversity. This review
concentrates to provide species-level information on biodiversity in the genus lilium and their future use in breeding programs. We
focus mainly on species used in breeding programme and grown mainly for cut flowers and pot production. For example; trumpet
shaped Lilium species showed comparative better prospective for exploitation than other species. We also present a brief summary on
research area that needs further development using biotechnological techniques like molecular assisted breeding, QTLs and
GISH/FISH and chloroplast genomes for comparative and phylogenetic analyses.
Keywords Lilium species; Biodiversity; Breeding; Domestication; Cytogenetic
Background
Considerable diversity ranging from nano sized plants to large trees found in the wild or cultivated in various
ecological habitats exists among over quarter million plant species dwelling on planet. Apart from this, a great
deal of interspecific diversity deems to exist. Also, the flowers of same plants differ morphologically to great
extent. These variations in germplasm allow plant breeders to breed novel cultivars or improve crop species for
various attributes. The term germplasm collection consists of over 6.5 million accessions stored securely in 1400
gene banks all over the world amongst which the most useful germplasm conserved are of food, industrial and
forage crops. However, despite the existence of floriculture as an important multibillion dollar industry barely less
than one percent of these are of ornamental herbaceous species and very few of bulbous ornamentals. The most
abundant temperate flora, with over 24000 species and varieties of indigenous flowering plants, has been found in
South Africa (Rourke, 1996), while Brazil, Malaysia, Colombia, Australia, Madagascar, Indonesia, Mexico, China
and Eucador are the world’s major centers of diversity for tropical ornamental plants.
Lilium is one of the majorgenus of geophytes commonly grown throughout the world for its ornamental value.
Owing to the availability of variable flower form, shape, colour and other phenotypic attributes, the genus possess
alarge number of hybrids and cultivars commercially (De Hertogh, 1996). Although a latin term Lilium, but is
derived from the Greek ‘Leirion’ for the Madona lily (Liliumcandidum) used by Theophrastus. In the language of
flower, lily symbolizes ‘purity and innocence’. In Christianity, lily is the symbol of virgin mother and in the
Semitic world, the symbol of motherhood. Apart from its ornamental value, few species are also known for
medicinal and food values, thus adding to its economic importance. Roman naturalist and writer Plinius recorded
that salvas and oils were prepared from the leaves and flowers of the madona lily. Bulbs of Liliumpolyphyllum
has been used as refrigerant, expectorant, aphrodisiac, diuretic, antipyretic and tonic (Warrier et al., 1997; Dhyani,
2007). In the customary system of medicine, this species reported to restore health immediately and works as
antioxidant in the body. Lily oil is an aid in childbirth. The white lily was also renowned as a cosmetic.
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International Journal of Horticulture, 2018, Vol. 8, No. 8, 83-97
1 Origin and Distribution

The genus Lilium L. (family Liliaceae) is constituted of about 110 species dispersed around the world that are
distributed between the 100 and 600N in Asia (50 to 60 species), North America (approximately 24 species), and
Europe (approximately 12 species) (Bryan, 1989; McRae, 1998; Liang and Tamura, 2000; Wang et al., 2015). All
species of Lilium are indigenous to the hilly region of the Northern Hemisphere up to Siberia and South Canada
up to Nilgirimountains of India and Florida in the South (Figure 1). Although no fossil record available, Lilium,
an important genus of the core Liliales, appears to have evolved approximately 12 million years ago inthe
Himalayas (Patterson and Givnish, 2002; Gao et al., 2015). Southwest and north of China is considered natural
home for 55 speciesof wild lily and is believed to be its centre of genetic diversity (De Jong, 1974). The East coast
of Asia, the West Coast of North America and the Mediterranean region are the three most richly garnished places.
Far East is the home of almost half of the world’s lilies and majority of our finest garden lilies originated there. A
few species have been found in Northeast India (Liliumwallichianum, L. napalense and L. polyphyllum) also. A
species may be either confined like, Lilium regale found only in one Chinese valley; or can be globe-trotter like
Liliummartagonwhich is recognized from Siberia across to Poland and down to the Balkans. The trumpet
lily(Liliumformosanum) indigenous to Taiwan, grows from sea-level to 3600 m. The worldwide distribution of
total lily cultivars and species and their distribution is registered in the lily Register from Wisley, Great Britain
from 1960-1993 (Table 1 and Table 2) (Mynett, 1996).
Figure 1 The geographical distribution of native lily species

Note: (a) Eurasian species range from the Atlantic through Mediterranean and Central Europe to the Caucasus and Ural mountains. (b)
Most American species are found in an area ranging from the Atlantic to the Midwest. The distribution of the Western species is
limited by the Rocky Mountains and the Pacific. (c) Most native lilies originate from the Eastern Asia. Their distribution ranges from
the Pacific to Ural and Caucasus mountains, to the Northern India and Burma in the south and Siberia in the north
2 History
With a history across thousands of years, the lily has been recognized in almost every sphere of human existence.
Going back through the ages, various descriptions have been given referring to lilies. The early European writer,
Pliny the Elder (23-73AD) expressed this flower as being ‘as far and noble as the rose’; an opinion that has been
shared by others including Wahlafrid Strabo, Bartholomew, Mattioli and Holland. Dioscorides, a Greek herbalist
in the 1st Century, observed the medicinal virtues of lilies, namely that of Liliumcandidum and healing qualities of
other types were also recognized by Asian and American civilizations. The lily was favoured by both the Greek
and Romans, and later became a sacred symbol in the monastery gardens of the Middle Ages, featuring in many
herbals and illuminated manuscripts. By the 17th century, several different lilies were known in the European
regions such as L. chalcedonicum, L. martagon, L. bulbiferum and L. candidum, and these were depicted in the
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works of Parkinson and Gerard. However, it was the botanical expeditions that various plant collectors undertook
to Asia in the 1800s and 1900s, that led to the discovery of many species of lily in China and Japan. Bulbs were
shipped to Europe and England in vast numbers and introduced into gardens by plant nurseries. Since that time,
lily species have spread throughout the world and many still retain their status as popular garden subjects, despite
being sidelined by the ever growing number of hybrids. Lilium regale, L. longiflorum, L. auratum, L. candidum,
and L. lancifolium are several examples of these. Meanwhile, for some native species existing in the wild, it is a
matter of trying to survive the often harsh results that human intrusion brings.
Table 1 The worldwide distribution of lily cultivars and species registered in the lily Register from Wisley, Great Britain from
1960-1993
Parts of world Number of cultivars and species % age
Europe 2259 44.0
U.S.A 2115 41.2
Australia 596 11.6
Asia 153 3.0
Africa 100 0.2
Table 2 The distribution of lily cultivars and species registered in the Lily Register from Wisley, Great Britain from 1960-1993
Countries Number of cultivars and species % age
Netherlands 782 34.6
Great Britain 383 17.6
Germany 347 15.4
Former Czechoslovakia 245 10.7
Latvia 239 10.4
Former Soviet Union 233 10.0
Poland 30 1.3
3 Botany
Lilium is an herbaceous perennial from family Liliaceae and subclass Monocotyledonae. These are generally
propagated by bulbs, composed of fleshy scales which act as store for food reserves, helping in growth during the
subsequent season. The scaled are attached to the short stems also known as axis or basal plate. It is the most
important segment from which roots emerges; also scales, and new growth buds arise from this region. Lilium
specie can be identified from its scale’s colour but this may vary on exposure to light. Lilium bulb may be either
concentric or rhizomatous, while most European and Asiatic lilies have concentric bulbs. Some species such
asLiliumlankongense, L. nepalense, and L. wilsonii, have stoloniferous habit wherethe stem bear bulblets away
from the mother bulb as it travels underground for some distance before emerging. The Eastern American species
(L. canadense, L. michiganense, and L. superbum) shows best development of rhizomatous type bulb while the
Western American species, L.pardalinum and L. parryiare some typical examples of modification of rhizomatous
type of bulbs.
Length of mature lilies’ flowering stem may vary from few inches in Liliumnanum or other high alpine species, to
as tall as 250 cm, in Liliumleucanthum var. centifolium or L. superbum. The stems may arise straight from the bulb,
as in L. regale or may travel some distance horizontally undergroundbefore emerging, as in L. lankongense and L.
nepalense. The leaves may be eitherlanceolate (broad) as in L. auratum var. platyphyllum or may be narrow grass
like, as in L.pumilum. In most of the oriental or trumpet species the leaves are alternately arranged or they might
be arranged in whorl, as some North American species while in some species, such as L. taliense, a naked,
asparagus-like stem is produced that rises 30 cm or more before the leaves expand. Whorling of leaves is also
noticed in L. candidum and L. x testaceum, but their leaves size reduces from the base upward. Small purple
bulbils in the axils of leaves are also observed in some species e.g. L. lancifolium.
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The lily inflorescence may be a raceme, an umbel, or a single terminal flower. A great deal of vividness in form
and colour is observed which adds on to the attractiveness of the genus. Trumpet (Lilium longiflorum and L.
regale), Turk’s-cap (Liliummartagon), and bowl shaped (Liliumauratum) are the basic forms. Lilies are also
distinguished as per the carriage of the flower being upright, pendant or outfacing. Flowers possess six stamens
consisting of slender filaments which bears anther containing the pollens. Lily pollen may be soft yellow to dark
brown in colour depending on the species or hybrid. Stigma is tri-lobed borne at the tip of long slender style, with
an ovary at the base. Nectar is secreted from the nectar furrow present in most species, as a groove at the base of
each tepal;to attract pollinating insects and birds.
Seeds are flat contained in structures called, capsules, which may be either short as in Liliumcandidum or long,
slender capsule as in L. formosanum. The capsule is divided into three sections each containing seeds stacked in
two rows; which are separated by thin papery divisions. A fertile seed can be distinguished by the presence of a
dark mass at the centre of the seed, which is marked as the line through the endosperm. Seeds of few species have
papery extensions or wings for facilitating easy dispersal by wind such as those of Liliumauratum or the wing
tissue might be very little as those of L. polyphyllum.
4 Classification and Species

Comber (1949) divided the genus Lilium into seven taxonomic sections Martagons (5 species), Americans or
Pseudolirium (21 species), Lilium/Candidum/Liriotypus (15 species), Orientals or Archelirion (7 species), Asiatics
or Sinomartagon (46 species), Trumpets or Leucolirion (11 species), and Daurolirion or Oxypetala (2 species),
based upon 15 morphological and physiological attributes, which was later revised by Lighty (1968) and De Jong
(1974). Origin and distribution of each species within each section is shown in Table 3.
Table 3 Origin and Native distribution of Lilium hybrids and species

Section Species Native distribution
Martagon L. distichum Amur and Vladivostok regions in Siberia, and to Manchuria and Korea
Ullung-Do and Takeshima Islands off the coast of Korea and the Diamond and
L. hansonii
Negita mountains in mainland Korea
L. martagon Eurasia
Honshu and Hokkaido in Japan, north to Sakhalin, the Kurile Island off Kamchatka,
L. medeoloides
and Cheju Island off southern Korea
L. tsingtauense China and Korea
American
a: L. bolanderi Southern Oregon and northern California
L. columbianum Western North America
L. humboldtii United States
L. kelloggii United States
L. rubescens Pacific Coast of the United States
L.washingtonianum Western American mountains
b: L. maritimum Coastal California in the United States
L. occidentale Pacific Coast of the United States
L. pardalinum Pacific Coast of the United States
L. parryi South-western United States
L. parvum Western United States
c: L. canadense Eastern North America
L. grayi North Carolina, Tennessee, and Virginia in the
United States
L. iridollae Southern Alabama and northwestern Florida in
the southeastern United States
L. michauxii Southeastern United States
L. michiganense United States
L. superbum United States
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Continued Table 3
Section Species Native distribution
d: L. catesbaei Southeastern United States
L. philadelphicum North America
Candidum L. bulbiferum Europe
L. candidum Eastern Mediterranean
L. carniolicum Balkans
L. chalcedonicum Albania and Greece
L. monadelphum Northern Caucasus
L. polyphyllum Himalayas (India)
L. pomponium Alps-Maritimes of France
L. pyrenaicum Europe, Turkey, and the Caucasus
L. alexandrae Japan.
Oriental L. auratum Japan
L. brownie
L. japonicum Southern China
L. nobilissimum Japan
L. rubellum Japan
L. speciosum Japan
L. davidii Japan, Taiwan, China
Asiatic
a: L. duchartrei China
L. henryi China
L. lancifolium China
L. lankongense Japan, Korea, China
L. leichtlinii China
L.papilliferum Japan
L. amabile Korea
b: L. callosum Eastern Asia
L. cernuum Korea, Russia, China
L. concolor China, Japan, Russia, Korea
L. pumilum South and North Korea, Russia, Mongolia, northern China
L. bakerianum Burma, Nepal, China
c: L. mackliniae Burma
L. nepalense Himalayas of Nepal, Bhutan, and Kumaon
L. ochraceum Nepal
L. sempervivoideum China
L. taliense China
L. wardii Southeastern Tibet
Trumpet section
a: L. leucanthum China
L. regale China
L. sargentiae China
L. sulphureum China
b: L. formosanum Taiwan
L. longiflorum Japan
L. neilgherrense India
L. philippinense Philippines
L. wallichianum Himalayas
Dauricum section L. dauricum Asia
L. maculatum Japan
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5 Hybrid Lilies
All lilies species and their hybrids belong to the genus ilium. There are nine groups under which they are defined
and maintained by the International Lily Register. The name of these divisions gives a broad indication of the
attributes of the hybrids (Table 4).
Table 4 Summary of horticultural divisions of genus Lilium

Division Species Remarks/traits/hybrids
Division 1:Asiatic hybrids L.amabile, L. bulbiferum, Bred mainly from the earlier flowering Asian species
L.callosum, L.cernuum, L.concolor,
L. dauricum, L. davidii, L.
lankongense, L.leichtlinii,
L.pumilum, L. lancifolium, L.
wilsonii, L.maculatum
Division 2: Martagon L. hansonii, L. martagon, L. (1) Took longer time to flowering from seed (5-7 years)
hybrids medeoloides, L. tsingtauense. ‘Marhan’ first hybrid;
(2) ‘Backhouse’ hybrid between L. martagon and L. hansonii;
(3) ‘Paisley’ hybrids appeared from a cross between L. martagon
var. album and L. hansonii;
(4) Resistant to viruses and cold hardy.
Divison 3:Candidum L. candidum, L. chalcedonicum, L. (1) Oldest known hybridsLiliumx testaceum;
hybrids monadelphum. (2) ‘Nankeen’ a cross between L. candidum and L.
chalcedonicum;
(3) ‘June Fragrance’ is a cross between L. candidumsalonikae
with L. monadelphum.
Division 4: American L. bolanderi, L. humboldtii, L. (1) Turk’s cap-shaped flowers;
hybrids kelloggii, L. pardalinum, L. parryi. (2) ‘Bellingham’ hybrids bred from L. humboldtii var. ocellatum,
L. pardalinum and L. parryi.
Division 5: Longiflorum L. longiflorum (1) Known as ‘Easter lily’;
hybrids (2) Vigrous growth and pure white trumpet shaped flowers.
Division 6: Chinese trumpet L. leucanthum, L. regale, L. (1) Grown in temperate zones;
and Aurelian hybrids sargentiae, L. sulphureum, L. (2) ‘Olympic hybrids’ derived from the crosses between L.
henryi, L. brownie. sargentiae, L. sulphureumand L. brownii;
(3) ‘Black Dragon’ is a selection from black magic strain;
(4) ‘Pink Perfection strain’ is a progeny of L. regale x L.
sargentiae and L. leucanthumcentifolium;
(5) Split into four classes separated by flower forms.
Division 7. Oriental hybrids L. alexandrae, L. auratum, L. (1) Large scented, showy, bowl shaped, flat or reflexed flower
japonicum, L. nobilissimum, L. with broad alternate leaves, late flowering;
rubellum, L. speciosum. (2) Vigor, disease resistance, virus tolerance;
(3) ‘Empress of Japan’ a huge flowered, red spotted gold band
derived from crossing L.xparkmannii with ‘Jillian Wallace’;
(4) ‘Potomac hybrids’ originated from the cross (L.auratum x L.
speciosum var. punctatum) x L. speciosum.
Division 8. Orienpet Derived from the crossing of (1) Hybrids between Oriental and Trumpet or Aurelians
hybrids species and hybrids from division 6 Suitable for warm climate;
and division 7. (2) Bloom 3-4 weeks later after Asiatics;
(3) Mostly sterile;
(4) Form embryos without endosperm, suitable for embryo
culture;
(5) Examples : ‘Scheherazade’, ‘Northern Carillon’, ‘Silk Road’,
‘Starburst Sensation’, ‘Northern Sensation’, and ‘Leslie
Woodriff’.
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Continued Table 4
Division Species Remarks/traits/hybrids
Division 9: Species All true species and their botanical -
varieties.
Division 10: Miscellaneous All hybrids not covered in the above -
hybrids divisions
The Archelirion, Leucolirion and Sinomartagon are considered to be the major economic groups due to
availability of greater genetic diversity and vivid morphological characteristics. The wild species (2n=2x=24)
yields fertile hybrids within each sections and are more or less crossable (McRae, 1990; Van Tuyl et al., 2002).
Thehybridization within these three sections has led to development of five prime lily cultivar groups viz; Asiatic,
Longiflorum, Oriental, Longiflorum x Asiatic and Longiflorum x Oriental (Figure 2 and Table 5).
Figure 2 Figures from different lily hybrids: (a) Longiflorum hybrids, (b) Oriental hybrids and (c) Asiatic hybrids
(a) The white trumpet-shaped flowers of longiflorum hybrids possesing distinctive fragrance, which can be
cultivated all round the year originated as a resultof intra or interspecific hybridization among the Leucolirion
sections i.e. LiliumlongiflorumThunb. × Liliumformosanum Wallace (McRae, 1990).
(b) The cultivation of Asiatic hybrids can be tracked down to the early 1800s in Japan, which are obtained as a
result of inter or intra-specific hybridization among at least 12 species of the Sinomartagon section (Shimizu,
1987).During the 1930s and early 1940s, the Mid-Century hybrids developed at Oregon bulb Farms, the United
States, were a great achievement in Asiatic hybrids (McRae, 1998). Besides possessing a wide array of colours
with diverse flowering periods (Woodcock and Stearn, 1950) few of the species of this section showed resistance
to Fusarium and viruses (McRae, 1998).
(c) The fragrant Oriental hybridsare cultivated and used in breeding since 1950s. These were derived frominter or
intra-specific crosses among five species of Archelirion (McRae, 1998). Majority of oriental hybrids are resistant
to Botrytis.
6 Breeding and Domestication

Lily is grown throughout the world, as one of the most significant flowering crop from past fifty years. Within the
first decade of the 20th century, interspecific cross among the section Sinomartagon including Liliummaculatum, L.
davidii, L. dauricum, L. bulbiferum and L. tigrinum was performed by Japan and the US (De Graaff, 1970). The
importance of lily crop was discovered in 1970 and thereafter several hybrid groups, such as, the ‘Harlequin
hybrids’, ‘the Mid-Century hybrids’, the ‘Preston hybrids’, and the ‘Patterson hybrids’, etc. were developed (De
Graaff, 1970; McRae, 1998; Rockwell et al., 1961). In 1944, Jan de Graaffbred a Mid-Century Hybrid
‘Enchantment’ which proved to be a turning point and revolutionized lily breeding and cultivation. Little later,
after the World War II the Dutch too began with their Lily breeding program. Presently, most of the lily cultivars
grown are polyploids.
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Table 5 An overview of the wild species involved in the origin of their groups of cultivars together with their main characteristics
Cultivars Section Species Descriptions of main traits
Longiflorum Leucolirion (1) Lilium longiflorum; (1) White trumpet shaped flower, very fragrance, year round forcing
cultivation;
(2) Liliumformosanum; (2) Short stem, deliciously fragrant, precocious flowering, red-purple
trumpet flower, heat tolerance;
(3) Lilium regale; (3) Horizontal white trumpet flower with a golden heart;
(4) Liliumnepalense (4) Down facing, flared and scented, pea-green flower with dark purple
throat; late flowering.
Asiatic Sinomartagon (1) L. amable (1) Bright orange down facing flower;
(2) L. bulbiferum (2) Orange upright facing flower;
(3) L. cornuum (3) Early flowering, small, sugar-pink, turk’s-cap flower;
(4) L. concolor (4) Small upright-facing, intense lacquer-red flower, thick and waxy
tepals;
(5) L.dauricum (5) Gold and vermilion, upright flowers, Fusarium resistant;
(6) L.davidii (6) Orange flower with spots; virus tolerance;
(7) L.tigrinum (7) Vigorous, strong stem, dark purple spotted orange turk’s-cap flower,
late flowering;
(8) L.lankongense (8) Spicily fragrant, pink to violet with spots of violet-rose flower;
(9) L. leichtlinii (9) Red-orageturk’s-cap flower;
(10) L.maculatum (10) Upfacing, maroon spotted apricot flowers;
(11) L.pumilum (11) Sweety scented, shiny-red, pendent, small turk’s-cap flowers.
Oriental Archelirion (1) L.alexandrae (1) White-green horizontal trumpet flower;
(2) L. auratum (2) Large flower, waxy leaves and tepals with a few or no spot, sweet
fragrance, resistance to diseases;
(3) L.nobilissimum (3) Scented, pure white and upright flowers, late flowering;
(4) L.rubellum (4) Deliciously fragrant, with rose pink and slightly recurved trumpet
flower, very early flowering;
(5) L.speciosum (5) Recurved, spicily fragrant, pale pink to cerise with darker spots
flower, late flowering;
(6) L.henryi (6) Orange spotted turk’s-cap flower with dark red/black spots, virus
resistance.
For over 30 years, Asiatic hybrids were the prominent cultivars. During this period the interspecific hybridization
of predominantly L. auratum and L. speciosumof the section Archelirion, gave rise to thy oriental hybrids. Large
sized lily flowers, predominantly pink and white in colour were introduced, from which, a Minnesota breeder
Leslie Wood riff bred the upward facing Oriental hybrids, which predominated for over 25 years.
Assortments of intersectional hybrids were produced using various pollination techniques such as cut styles,
embryo rescue and polyploidization (Asano, 1978, 1980; Asano and Myodo, 1977a, b; Van Tuyl et al., 1991; Lim,
2000; Barba-Gonzales, 2005; Barba-Gonzalez et al., 2006). Thestarting array of intersectional hybrids
Longiflorum x Asiatic hybrids or LA hybrids were produced as a result of interspecific hybridization among
various sections; about 20 years later than the Oriental hybrids. Repeated backcrossing of chromosome doubled
F1 LA hybrid with an Asiatic hybrid has rendered them, triploid. Recently these hybrids have replaced the Asiatic
hybrid group attaining an imperative position in the market.
The breeding of other intersectional crosses, primarily triploid hybrids like OA (Oriental x Asiatic hybrids), OT
(Oriental x Trumpet) and LO (Longiflorum x Oriental), began. The replacement of Orientals by the recent OTs
can be predicted from the earlier experiences of the LA-hybrids largely taking the place of the Asiatic hybrids and
the Longiflorums replaced by LO-hybrids. Apart from the major seven groups of lily hybrids (O, A, L, T, LA, LO
and OT), various other successful breeding of species occurred. The interspecific crosses of L. candidum with L.
longiflorum and L. henryii were excluded from this assortment. Several other crosses were made using
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Longiflorum as the female and species like L. auratum, L. bakerianum, L. canadense, L. concolor, L. dauricum,
L. henryii, L. kelloggii, L. lankongense, L. lophophorum, L. monadelpum, L. martagon, L. hansonii, L. nepalense,
L. pardinum, L. pumilum and L. semper vivoideum as male parent. Similarly, L. dauricum, L. nepalense and L.
pardalinum were used as male with Oriental as the female parent.
Cultivar ‘Stargazer’ was the topmost grown cultivar for over 25 years. Lily varieties are protected by Plant
Breeder’s Rights, while only Star Gazer was excluded from this right. Tissue culture techniques have come into
vogue for the development of fresh lily cultivars. The assortment shifts, as witnessed from the past trends, for the
future development of several other groups, breeding attempts have major contribution. The OT and the
Longiflorum hybrids may most likely be replaced by the Orientals and the LO-hybrids in the forthcoming years.
Several Crossed species: L. hansonii and L. martagon (section Martagon), L. candidum and L. monadelpum
(section Lilium), L. canadense and L. pardalinum (section Pseudolirium), L. bakerianum, L. nepalense and L.
henryi are potential species to be used in developing new groups. Transfer of desirable traits from wild species to
the array is being carried with the assistance of Molecular cytogenetics (Lim, 2000; Barba-Gonzalez, 2005; and
Khan, 2009) and molecular assisted breeding (Shanin et al., 2009).
Genetic transformation is another tool being sought after in lily for over 25 years (Cohen, 2011). It is widely used
in Genetic studies although no cultivar has been developed till date using genetic transformation. Detailed genetic
maps including 6 QTLs for Fusarium-resistance, of two lily populations, were published (Shain et al., 2010). The
wide genetic variation already accessible to breeder can be manifested for mixing of different desirable traits into
one, through the practical understanding of DNA-sequences linked to horticultural traits.
7 Cytogenetics
Lily has been long used as a model plant by various cytogeneticsts since Strasburger (1880), for chromosomal
studies including the description of meiotic stages, crossing over (Fogwill, 1958; Stern and Hotta, 1977) and
duration of female meiotic stages (Bennett and Stern, 1975), chromosome banding (Holm, 1976; Son, 1977; Song,
1987; 1991), fertility of interspecific hybrids (Brock, 1954; Asano, 1984), Synaptonemal complexes (Holm and
Rasmussen, 1984; Anderson et al., 1994) and chromosome morphology (Sato, 1932; Stewart, 1947; Noda, 1978;
Stack, 1991 and others). The basic chromosome number of all the lilium species is x=12 having two metacentric
whereas 10 acro-centric chromosomes. The genome size of Lilium species varies from 69.5pg/2C (Lilium henryi)
up to 96 pg/2C (Liliumparryi) (Bennet and Smith, 1991; Van Tuyl and Boon, 1997). The basic nomenclature of
Lilium karyotypes is based on the order of chromosomes in sequences of decreasing short-arm length; with 1
representing the longest short arm and 12 the shortest (Stewart, 1947; Lim et al., 2000). Both diploids and
triploids are found in L. lancifolium alone while all the other species are diploid. Van Tuyl and Boon (1997)
studied the variation in DNA content within the genus Lilium. It was discovered that interspecific hybrids
generally have intermediate DNA content to that of parents. Lilium longiflorum has been a model crop for
studying gametophytic self incompatability due to presence of strong S.I. mechanism.
8 Molecular Breeding and Physical Mapping in Lilium

Species of the genus Lilium have the largest genomes among plants except Fritillaria (Bennett and Smith 1991)
and have been extensively used for cytological analysis in the past (Mather, 1940; Stewart, 1947; Brown and
Zohary, 1955; Fogwill, 1958; Bennett and Stern, 1975; Noda, 1991). Nevertheless, no reliable molecular or
cytogenetic maps are available for lily so far. At present, breeding programs aim at combining desirable traits
within different plant species complexes and are mainly focused on introgression procedures (Kosmala et al., 2006
and 2007). The main thrustarea is the transfer of genes conferring to a particular trait from a wild species into a
cultivar. Markers associated with genes conferring particular characters can facilitate the marker-assisted selection
program to obtain new cultivars with desired traits. For this purpose, genetic or molecular maps have been
constructed in several crops and integrated in cytogenetic maps of respective genomes (Khrustaleva et al., 2005;
Humphreys et al., 2005).
91
Multiple approaches have been used to develop integrated maps, primarily by combining the information from
molecular maps and physical distribution of these markers in the genome through cytogenetic techniques. Such
maps are constructed by insertion of large DNA clones like yeast artificial chromosomes (YAC) and bacterial
artificial chromosomes (BAC) and comparison of these maps with genetic maps (Wu et al., 2003). The other
techniques used are in situ hybridization of BACs and YACs on plant chromosomes with recombination maps
(Kulikova et al., 2001). One of the current developments in lily breeding is the introduction of molecular breeding
techniques for the incorporation of disease resistances in commercially successful lily cultivars. Fusarium,
Botrytis and virus (LMoV) resistance are the most important goals in lily breeding. Screening methods to
determine resistance against these pathogens in several lily cultivars have been developed (Straathof and Van Tuyl,
1994; Straathof and Löffler, 1994). However, it takes a long time before resistance symptoms become visible, and
therefore, faster selection methods are needed.
In lily molecular marker techniques have been used for the identification of cultivars and interspecific hybrids
(Yamagishi, 1995; Yamagishi et al., 2002), identifying genetic diversity (Arzate-Fernández et al., 2005), genetic
analysis of anthocyanin pigmentation (Abe et al., 2002) and for finding RAPD-markerslinked to
Fusariumoxysporumresistance in Asiatic hybrids (Straathofet al. 1996). Van Heusdenet al. (2002) used AFLPs and
a QTL approach to locate markers linked to LMoV virus on Asiatic genome. A total of 251 AFLP markers based
on 100 descendents of a lily backcross population were used for the construction of a genetic map. Based on these
approaches it was found that LMoV (TBV) resistance was clearly a monogenic trait and could reliably be mapped
on linkage group 9. Despite the difficult screening for Fusarium resistance, four significant QTLs were mapped to
linkage groups 1, 5, 13 and 16 respectively.
Insertion of genes that are absent in lily can be achieved through techniques genetic transformation. Several
researchers have successfully accomplished both Agrobacterium and microprojectile mediated transformation in
lily (Bino et al., 1992; Van der Leede-Plegt et al., 1992; Watad et al., 1998). The GUS gene was successfully
transmitted into next generation. For future applications, a marker-free system is required.
Du et al., (2017) sequenced nine Lilium chloroplast genomes and retrieved seven published chloroplast genomes
for comparative and phylogenetic analyses. The genomes ranged from 151,655 bp to 153,235 bp in length and had
a typical quadripartite structure with a conserved genome arrangement and moderate divergence. A comparison of
sixteen Lilium chloroplast genomes revealed ten mutation hotspots. Single nucleotide polymorphisms (SNPs) for
any two Lilium chloroplast genomes ranged from 8 to 1,178 and provided robust data for phylogeny. Except for
some of the shortest internodes, phylogenetic relationships of the Lilium species inferred from the chloroplast
genome obtained high support, indicating that chloroplast genome data will be useful to help resolve the deeper
branches of phylogeny.
9 Utilization of Genetic Diversity

Germplasm conservation is necessary to prevent genetic erosion or narrowing of genetic base to protect
endangered species from getting extinct and availability of genetic material for developing new superior varieties.
Conservation is done keeping in view the requirements for future generations aiding in survival. A great deal of
genetic variation and novel horticultural traits, are present in lily with approximately 7000 cultivars available in
the market or wild. Commercially important characters include:
Resistance against various pests and pathogens such as Fusarium, Botrytis and several viruses (TBV, LSV and
LVX).
Novel attributes such as vibrant colour, flower shape, size, sturdy stem, form and fragrance.
Physiological characteristics such as low-light intensity and heat tolerance, precocious flowering, leaf scorch,
year-round forcing ability, long shelf life, and bulb growth speed. Some well-known examples of valuable
characters among species and different sections shown in Table 5.
92
Blending or introgression of different desirable traits from an alien species into a single cultivar is done through
interspecific hybridization or recurrent back crossing. Presence of pre and post fertilization barriers in lilium
species is a major challenge for its interspecific hybridization. Although interspecific hybridization has
successfully developed hybrids using methods, such as cut style pollination, embryo rescue and ovule culture (Van
Tuyl et al., 1991; 2000). Utilization of novel techniques like mentor pollination ovary and ovule culture and in
vitro pollination has helped in producing numerous interspecific hybrids in 80s and 90s (Van Tuyl et al., 1982;
1988; 1991; Wolf and Van Tuyl, 1984).
Skim (1942) used embryo culture to develop plants from L.henryi x L.regale. Immature hybrid embryos between
L. ‘Shikayama’ x L. henryi and L. longiflorum x L. ‘Sugehime’ were reported by Asano and Myodo (1977b).
Ascher (1973a, b) successfully obtained the plants of L. ‘damson’ x L. longiflorum. Asano (1980) developed
several hybrids between L. longiflorum x L. candidum, L. longiflorum x L. dauricum, L. longiflorum x L.
candidum, L. longiflorum x L. amabile, and L. longiflorum x L. pumilum, x L. ‘Sasatame’ x L. henryi, L. ‘Royal
Gold’ x L. speciosum and L. regale x L. leichtliniimaximowiczii through interspecific hybridization. Later, Van
Tuyl and coworkers (1991 and 2002) obtained numerous intersectional hybrids successfully. Examples include L.
longiflorum x L. canadense (Pseudolirium section); L. longiflorum (Leucolirion section) x L. monadelphum
(Lilium section), L. longiflorum x L. martagon (Martagon section), L. longiflorum x L. lankongense
(Sinomartagon section), L. longiflorum x L. rubellum (Archelirion section), L. longiflorum x Oriental hybrids,
Oriental x Asiatic hybrids.
The cultivar ‘Black Beauty’ was developed by crossing L. speciosum var. rubrum x L. henryi which turned out to
be infertile and tetraploid. Lilium cv. ‘Arc Besque’ developed by crossing ‘Black Beauty’ (4n) x ‘Journey End’
(4n), which is six feet tall has intense red flat spreading tepals about 8 inches from tip to tip and is usually disease
resistant. The first pink coloured Easter lily ‘Rote Horn’ was developed by hybridizing L .longiflorum and L.
elegans (Asano and Nyoda, 1978). ‘Yukinohikari’ developed by Takizawa (1977) is a hybrid of L. formolongi x L.
auratum and consequently displays vigorous growth and multiple flowers from a small bulb. ‘Elegant lady’
(triploid LLR), developed as a result of interspecific hybridization among L. longiflorum and a LLRR F1
interspecific hybrid is a successful example. It is an early flowering; fragrant hybrid which bears a pink coloured,
tubular flower. GISH confirms the presence of several recombinant chromosomes between the L and A genomes
as in the LA hybrid ‘Fangio’(2n=3x=36; triploid, A genome=24, L genome=12). Techniques such have embryo
culture and cut style pollination have been utilized for developing Asiatic hybrid lilies such as ‘Li-9’ (‘Mona’ x L.
concolor var pulchellum) which produces multiple scapes per bulb (Oomiya et al., 2005). Mynett (2007) obtained
a yellow and orange late flowering cultivar in Asiatic lilies by crossing ‘Yellow Blaze’ very late flowering cultivar
and L. wilsonii. The Akita Perfectural Agricultural Experiment Station, Japan has developed a series of Asiatic
hybrids that are male sterile (Shibata, 2002). “Akita Petit White’ was the first male sterile cultivars released
commercially by this station followed by ‘Akita Petit Lemon’, Akita Petit Gold’ and ‘Akita Petit Cream’ having
yellow perianth were released (Asari et al., 2005). These were the selection from the progeny of reciprocal crosses
between two Asiatic lilies, ‘Connecticut King’ and ‘Menton’. Korea released new FA hybrid ‘Supia’ and FAA
hybrid ‘Pink Pearl’. Few new FO hybrid lines, ‘FO 03-16’ line, OA hybrid ‘OA 05-1’ line, and OH hybrid ‘OH
02-1’ line were also developed at Korea (Rhee and Kim, 2008).
10 Conclusions
Lilium is one of the important genus of geophytes widely grown throughout the world. The genus possess a great
genetic diversity in many valuable horticultural traits which is manifested in flower colour, forms, shape,
fragrance, resistance to diseases, and many physiological characteristics. It is important to conserve available
diversity existing among the genus, to breed better and superior quality new cultivars. From the past trend the
popularity of large flowered Orientals taking over the Asiatic hybrids in 1970s and 1980s, it can be assumed that
there is need for more complex hybrids as demonstrated by The LA-hybrids. Recently, various FO, LO, OT and
OA hybrids are emerging. With the same pace, an indistinguishable group of hybrid lilies shall developed
possessing nearly similar characters for various species and hybrids group. Several advanced techniques like
93
GISH/FISH-techniques and molecular assisted breeding shall be utilized for breeding for novel characteristicssuch
as resistance to virus, Botrytis, Fusarium etc. Furthermore, chloroplast genome data may help to better understand
the concept of evolution and adaption further assisting in development and resolving the deeper phylogeny.
Authors’ contributions
All the authors have equally contributed in the compilation and checking of the manuscript.
Acknowledgement
The authors are highly thankful to the Indian Council of Agricultural Research, New Delhi for supporting of this research
programme.
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Biodiversity in Lilium: A Review

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Biodiversity in Lilium: A Review

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International Journal of Horticulture, 2018, Vol. 8, No.

Research Article Open Access

1 Origin and Distribution

Figure 1 The geographical distribution of native lily species

4 Classification and Species

Table 3 Origin and Native distribution of Lilium hybrids and species

Table 4 Summary of horticultural divisions of genus Lilium

6 Breeding and Domestication

8 Molecular Breeding and Physical Mapping in Lilium

9 Utilization of Genetic Diversity

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