General
floral formula of the Liliacaeae: Flowers actinomorphic or slightly zygomorphic and hermaphrodite with 6 undifferentiated tepals in two whorls of three, the same number and arrangement of stamens, and a superior ovary with 3 fused carpels. Individual species and genera may have more or less derived formulas.
The lily
family, Liliaceae, consists of about 15
genera and 610
species of
flowering plants within the
orderLiliales.[2] They are
monocotyledonous,
perennial,
herbaceous, often
bulbousgeophytes. Plants in this family have evolved with a fair amount of morphological diversity despite genetic similarity. Common characteristics include large flowers with parts arranged in threes: with six colored or patterned
petaloidtepals (undifferentiated petals and sepals) arranged in two
whorls, six
stamens and a superior
ovary. The leaves are linear in shape, with their veins usually arranged parallel to the edges, single and arranged alternating on the stem, or in a rosette at the base. Most species are grown from bulbs, although some have
rhizomes. First described in 1789, the lily family became a
paraphyletic "catch-all" (
wastebasket) group of
lilioid monocots that did not fit into other families and included a great number of genera now included in other families and in some cases in other orders. Consequently, many sources and descriptions labelled "Liliaceae" deal with the broader sense of the family.
The family evolved approximately 68
million years ago during the
Late Cretaceous to
Early Paleogene epochs. Liliaceae are widely distributed, mainly in temperate regions of the Northern Hemisphere and the flowers are insect pollinated. Many Liliaceae are important
ornamental plants, widely grown for their attractive flowers and involved in a major
floriculture of
cut flowers and dry bulbs. Some species are
poisonous if eaten and can have adverse health effects in humans and household pets.
A number of Liliaceae genera are popular cultivated plants in private and public spaces. Lilies and tulips in particular have had considerable symbolic and decorative value, and appear frequently in paintings and the decorative arts. They are also an economically important product. Most of their genera, Lilium in particular, face considerable herbivory pressure from deer in some areas, both wild and domestic.[4][5]
The diversity of characteristics complicates any description of the Liliaceae morphology, and confused taxonomic classification for centuries. The diversity is also of considerable evolutionary significance, as some members emerged from shaded areas and adapted to a more open environment (see
Evolution).[6]
General
The Liliaceae are characterised as monocotyledonous, perennial, herbaceous, bulbous (or
rhizomatous in the case of
Medeoleae)[7] flowering plants with simple
trichomes (root hairs) and contractile
roots.[8] The flowers may be arranged (inflorescence) along the stem, developing from the base, or as a single flower at the tip of the stem, or as a cluster of flowers. They contain both male (androecium) and female (gynoecium) characteristics and are symmetric radially, but sometimes as a mirror image. Most flowers are large and colourful, except for Medeoleae. Both the petals and sepals are usually similar and appear as two concentric groups (whorls) of 'petals', that are often striped or multi-coloured, and produce nectar at their bases. The stamens are usually in two groups of three (trimerous) and the pollen has a single groove (monosulcate). The ovary is placed above the attachment of the other parts (superior). There are three fused carpels (syncarpus) with one to three chambers (locules), a single style and a three-lobed stigma. The embryo sac is of the Fritillaria type. The fruit is generally a wind dispersed capsule, but occasionally a berry (Medeoleae) which is dispersed by animals. The leaves are generally simple and elongated with veins parallel to the edges, arranged singly and alternating on the stem, but may form a rosette at the base of the stem.
Usually
indeterminate (lacking terminal flower) as a
raceme (Lilium); sometimes reduced to a single terminal flower (Tulipa). When
pluriflor (multiple blooms), the flowers are arranged in a cluster or rarely are
subumbellate (Gagea) or a
thyrse (spike).[9]
Hermaphroditic,
actinomorphic (radially symmetric) or slightly zygomorphic (bilaterally symmetric),[10]pedicellate (on a short secondary stem), generally large and showy but may be inconspicuous : (Medeoleae).
Bracts may (bracteate) or may not (ebracteate) be present. The
perianth is undifferentiated (
perigonium) and biseriate (two
whorled), formed from six
tepals arranged into two separate whorls of three parts (
trimerous) each, although Scoliopus has only three petals, free from the other parts, but overlapping. The tepals are usually petaloid (petal like) and
apotepalous (free) with lines (striate) or marks in other colors or shades. The
perianth is either homochlamydeous (all
tepals equal, e.g. Fritillaria) or dichlamydeous (two separate and different whorls, e.g. Calochortus) and may be united into a tube.
Nectar is produced in perigonal nectaries at the base of the tepals.[9][11][12][13]
Six
stamens in two trimerous whorls, with free filaments, usually epiphyllous (fused to tepals) and diplostemonous (outer whorl of stamens opposite outer tepals and the inner whorl opposite inner tepals), although Scoliopus has three stamens opposite the outer tepals.[14] The attachment of the anthers to the filaments may be either peltate (to the surface) or pseudo-basifixed (surrounding the filament tip, but not adnate, that is not fused) and
dehisce longitudinally and are extrorse (dehiscing away from center). The
pollen is usually monosulcate (single groove), but may be inaperturate (lacking aperture: Clintonia, some Tulipa spp.) or operculate (lidded: Fritillaria, some Tulipa spp.),[15] and reticulate (net patterned: Erythronium, Fritillaria, Gagea, Lilium, Tulipa).[9][11][14]
Superior
ovary (hypogynous), syncarpous (with fused carpels), with three connate (fused) carpels and is trilocular (three
locules, or chambers) or unilocular (single locule, as in Scoliopus and Medeola). There is a single style and a three lobed
stigma or three stigmata more or less elongated along the style. There are numerous anatropous (curved)
ovules which display axile
placentation (parietal in Scoliopus and Medeola),[14] usually with an
integument and thinner
megasporangium. The embryo sac (
megagametophyte) varies by genera, but is mainly
tetrasporic (e.g. Fritillaria).[11] Embryo sacs in which three of the four
megaspores fuse to form a
triploidnucleus, are referred to as Fritillaria-type, a characteristic shared by all the core Liliales.[6][9][11][16]
A
capsule that is usually loculicidal (splitting along the
locules) as in the Lilioideae,[6] but occasionally septicidal (splitting between them, along the separating
septa) in the Calachortoideae and wind dispersed, although the Medeoleae form
berries (baccate).[6] The
seeds may be flat, oblong, angular,
discoid,
ellipsoid or
globose (spherical), or compressed with a well developed
epidermis. The exterior may be smooth or roughened, with a wing or
raphe (ridge),
aril or one to two tails, rarely hairy, but may be dull or shiny and the lack of a black
integument distinguishes them from related taxa such as
Allioideae that were previously included in this family, and striate (parallel longitudinally ridged) in the Steptopoideae. The
hilum (scar) is generally inconspicuous. The bitegmic (separate testa and tegmen) seed coat itself may be thin, suberose (like
cork), or crustaceous (hard or brittle). The
endosperm is abundant,
cartilaginous (fleshy) or horny and contains oils and
aleurone but not
starch (non-farinaceous). Its cells are
polyploid (
triploid or
pentaploid, depending on the
embryo sac type). The
embryo is small (usually less than one quarter of seed volume), axile (radially sectioned), linear (longer than broad) or rarely
rudimentary (tiny relative to
endosperm) depending on placentation type, and straight, bent, curved or curled at the upper end.[9][11][17][18]
Simple, entire (smooth and even), linear, oval to filiform (thread-like), mostly with parallel veins, but occasionally net-veined. They are alternate (single and alternating direction) and
spiral, but may be
whorled (three or more attached at one
node, e.g. Lilium,
Fritillaria), cauline (arranged along the aerial
stem) or sheathed in a basal
rosette. They are rarely
petiolate (stem attached before apex), and lack
stipules. The aerial stem is unbranched.[9][11][12][19][20]
The Liliaceae include a species with one of the largest
genome size within the angiosperms, Fritillaria assyriaca (
1C=127.4 pg), while Tricyrtis macropoda is as small as 4.25 pg.[21]Chromosome numbers vary by genus.[11] Some genera like Calochortus (x=6-10), Prosartes (6,8,9,11), Scoliopus (7,8), Streptopus (8, 27) and Tricyrtis (12-13) have a small and variable number of chromosomes while subfamily Lilioideae have a larger and more stable chromosome number (12) as have the Medeoleae (7).[7][22][23][24]
Characteristics often vary by habitat, between
shade-dwelling genera (such as Prosartes, Tricyrtis, Cardiocrinum, Clintonia, Medeola, Prosartes, and Scoliopus) and sun loving genera. Shade-dwelling genera usually have broader leaves with smooth edges and net venation, and fleshy fruits (
berries) with animal-dispersed seeds,
rhizomes, and small, inconspicuous flowers while genera native to sunny habitats usually have narrow, parallel-veined leaves,
capsular fruits with wind-dispersed seeds, bulbs, and large, visually conspicuous flowers.[7] (See also Evolution).
The taxonomy of the Liliaceae has a very complex history. The family was first described in the eighteenth century, and over time many other genera were added until it became one of the largest of the monocotyledon families, and also extremely diverse. Modern taxonomic systems, such as the
APG which is based on
phylogenetic principles using
molecular biology, have redistributed many of these genera resulting in the relatively small family that is currently recognised. Consequently, there are many different accounts of the Liliaceae in the literature and older uses of the term occur commonly. To distinguish between them, the
Latin terms sensu lato and sensu stricto are frequently used (together with their abbreviations, s.l. and s.s.) to denote the broader or stricter sense of the circumscription respectively, e.g. Liliaceae s.s..
History
The family Liliaceae was described by
Michel Adanson in 1763 and formally named by
Antoine Laurent de Jussieu in 1789. Jussieu defined this grouping as having a
calyx of six equal colored parts, six
stamens, a superior
ovary, single
style, and a trilocular (three-chambered)
capsule. By 1845,
John Lindley, the first English
systematist, unhappily acknowledged the great diversity in the
circumscription of the family, and that it had expanded vastly, with many subdivisions. As he saw it, the Liliaceae were already
paraphyletic ("catch-all"), being all
Liliales not included in the other orders, but hoped that the future would reveal some characteristic that would group them better. He recognized 133 genera and 1200 species.[28] By the time of the next major British classification – that of
Bentham and Hooker in 1883 (published in Latin) – several of Lindley's other families had already been absorbed into the Liliaceae.[29] Over time the family became increasingly broad and somewhat arbitrarily defined as all species of plants with six tepals and a superior ovary, eventually coming to encompass about 300 genera and 4,500 species within the
orderLiliales under the
Cronquist system (1981). Cronquist merged the Liliaceae with the
Amaryllidaceae, making this one of the largest monocotyledon families.[30]
Many other botanists echoed Lindley's earlier concerns about the
phylogeny of the Liliaceae, but various schemes to divide the family gained little traction.
Dahlgren (1985) suggested there were in fact forty – not one – families distributed over three orders (predominantly
Liliales and
Asparagales).[31][32] In the context of a general review of the classification of
angiosperms, the Liliaceae were subjected to more intense scrutiny. Considerable progress in plant phylogeny and
phylogenetic theory enabled a
phylogenetic tree to be constructed for all of the flowering plants, as elaborated by the
Angiosperm Phylogeny Group (1998).[6]
Modern APG classification and phylogeny
The Angiosperm Phylogeny Group (
APG) made rapid progress in establishing a modern
monophyletic classification of the flowering plants by 2009.[33] Despite establishing this relative degree of
monophyly (genetic homogeneity) for the family Liliaceae,[34][35] their morphology remains diverse[7] and there exists within the Liliaceae
clade or grouping, a number of subclades (subgroups). Particularly enigmatic were Clintonia, Medeola, Scoliopus, and Tricyrtis.
Of the fifteen genera within the Liliaceae, the ten genera of the
Lilioideaesubfamily form one morphological group that is characterised by contractile bulbs and roots, and a Fritillaria-type
embryo-sac (megagametophyte with four megaspores). Within the Lilioideae, Clintonia and the closely related Medeola form a subclade, and are now considered a separate tribe (
Medeoleae).[36] The other major grouping consists of the five genera constituting the
Streptopoideae (including Scoliopus) and
Calochortoideae (including Tricyrtis) subfamilies characterised by creeping rhizomes, styles which are divided at their apices, and by megagametophyte development of the Polygonum-type (a simple megaspore and triploid endosperm) embryo-sac.
The Liliaceae probably arose as
shade plants, with subsequent evolution to open areas including
deciduous forest in the more open autumnal period, but then a return of some species (e.g. Cardiocrinum). This was accompanied by a shift from
rhizomes to
bulbs, to more showy flowers, the production of capsular fruit and narrower parallel-veined leaves. Again, some reversal to the broader reticulate-veined leaves occurred (e.g. Cardiocrinum).[6]
Due to the diversity of the originally broadly defined Liliaceae s.l., many attempts have been made to form suprageneric classifications, e.g.
subfamilies and
tribes.[23] Classifications published since the use of
molecular methods in phylogenetics have taken a narrower view of the Liliaceae (Liliaceae s.s.). The
Angiosperm Phylogeny Website (APweb)[2] recognizes three subfamilies, one of which is divided into two tribes.[11]
Various authorities (e.g.
ITIS 16,[43]GRIN 27,[44]WCSP,[45]NCBI,[46] DELTA[19] ) differ on the exact number of genera included in Liliaceae s.s., but generally there are about fifteen to sixteen genera, depending on whether or not Amana is included in Tulipa and Lloydia in Gagea.
Currently the APWeb lists fifteen genera, arranged as shown in this table:[2][11]
APweb Distribution of subfamilies, tribes and genera of Liliaceae
The Liliaceae are widely distributed, but mainly in the
temperate regions of the
Northern Hemisphere. The
centre of diversity is from
southwest Asia to
China. Their distribution is diverse, mainly in
plains,
steppes, and
alpine meadows, but also in
deciduous forests,
Mediterranean scrub and
arctic tundra.[6][9]Tulipa and Gagea provide examples of ornamental geophyte biomorphological types representing continental thermoperiodic zones (
Irano-Turanian region), characterised by cessation of underground growth at high temperatures in early summer and requiring low winter temperatures for spring flowering.[40] While some genera are shade-dwelling, such as the Medeoleae, and Streptopoideae, Tricyrtis, and Cardiocrinum, others prefer a more open habitat.
Ecology
The Liliaceae are ecologically diverse.[6] Species of Liliaceae bloom at various times from spring to late summer. The colorful flowers produce large amounts of
nectar and
pollen that attract
insects which
pollinate them (entomophily), particularly
bees and
wasps (hymenopterophily),
butterflies (psychophily) and
moths (phalaenophily).[20] The
seeds are
dispersed by wind and water. Some species (e.g. Scoliopus, Erythronium and Gagea) have seeds with an
aril structure that are dispersed by
ants (
myrmecochory).[6]
The proliferation of deer populations in many areas, due to human factors such as the elimination of their animal predators and introduction to alien environments, is placing considerable herbivory pressure on many of the family's species.[4] Fences as high as 8 feet may be required to prevent them from consuming the plants, an impractical solution for most wild areas.[5] Those of the genus Lilium are particularly palatable, while species in Fritillaria are repellant.
Both Lilium and Tulipa are susceptible to a group of five viruses of the family Potyviridae, specifically the
potyvirus (named for potato virus Y) group, which includes the tulip-breaking virus (TBV) and the lily streak virus (lily mottle virus, LMoV) resulting in 'breaking' of the color of the flowers. The viruses are transmitted by
aphids. This breaking effect was of economic importance during the
tulip mania of the seventeenth century, because it appeared to be producing new varieties.[51][52] In modern times tulip breeders have produced varieties that mimic the effect of the virus, without being infected. One of these varieties is known as 'Rembrandt', after the
Dutch artist of that name. Contemporary tulip owners commonly had Rembrandt and other artists paint their flowers to preserve them for posterity, hence the 'broken' tulips were known as Rembrandt tulips at that time. Another modern variety is 'Princess Irene'.[53][54][55] One of the tulip breaking viruses is also named the Rembrandt tulip-breaking virus (ReTBV).[56][57]
Cultivation
Cultivation
Tulipa varieties are popular ornamental flowers in public and private spaces in the spring
Tulips have been cultivated since at least the tenth century in
Persia.[58] Tulip production has two main markets:
cut flowers and
bulbs. The latter are used, in turn, to meet the demand for bulbs for parks, gardens, and home use and, secondly, to provide the necessary bulbs for cut flower production.
International trade in cut flowers has an approximate total value of 11 billion
euros, which provides an indication of the economic importance of this activity. The main producer of tulip bulbs is the
Netherlands, a country that accounts for 87% of the global cultivated area, with approximately 12,000
hectares. Other leading producers include
Japan,
France and
Poland. Approximately ten other countries produce commercial tulips, largely for the
domestic market. By contrast, the Netherlands is the leading international producer, to the extent of 4 billion bulbs per annum. Of these, 53% are used for the cut flower market and the remainder for the dry bulb market. Of the cut flowers, 57% are used for the domestic market in the Netherlands and the remainder exported.[59][60]
Original Tulipa species can be obtained for ornamental purposes, such as Tulipa tarda and Tulipa turkestanica. These are referred to as
species, or botanical, tulips, and tend to be smaller plants but better at
naturalising than the cultivated forms. Breeding programs have produced a wide range of tulip types, enabling blooming through a much longer season by creating early, mid- and late spring varieties. Fourteen distinct types are available in addition to botanical tulips, including Lily-flowered, Fringed, Viridiflora, and Rembrandt. In addition to blooming season, tulip varieties differ in shape and height, and exhibit a wide range of colours, both pure and in combination.[61]
Lilies
The largest area of production is also the Netherlands, with 76% of the global cultivated area, followed by
France,
Chile,
Japan, the
United States,
New Zealand and
Australia. Approximately ten countries produce lilies commercially altogether. About half of the commercial production is for cut flowers. Many of these countries export bulbs as well as supplying the domestic market. The Netherlands produces about 2,200 million lily bulbs annually, of which 96% is used domestically and the remainder exported, principally within the
European Union. One particularly important crop is the production of Lilium longiflorum, whose white flowers are associated with purity and Easter.[59]
Although many Lilium species such as Lilium martagon and Lilium candidum can be obtained commercially, the majority of commercially available lilies represent the products of a very diverse hybridisation program, which has resulted in a separate horticultural classification, including such groupings as Asian, Oriental and Orienpet. In addition to a very wide variety of heights, lilies can be obtained in many colours and combinations of colours, and if properly selected can produce an extensive blooming season from early summer to autumn.[62] Because of the history of Liliaceae, many species such as
Watsonia (bugle lily) that were previously classified in this family bear the name 'lily' but are neither part of the genus Lilium, or the family Liliaceae.
Other
A variety of Fritillaria species are used as early spring ornamental flowers. These vary from the large Fritillaria imperialis (crown imperial) available in a number of colours such as yellow or orange, to much smaller species such as Fritillaria meleagris or Fritillaria uva-vulpis with their chequered patterns.[63]Erythronium is less common but a popular cultivar is 'Pagoda' with its sulphur yellow flowers.[64]Calochortus (mariposa lily) may be sold as a mixture or as cultivars.[65]
Seeds can be used for propagation of the plant or to create
hybrids and can take five to eight years to produce flowering plants. Since
interspecificcross-pollination occurs, overlapping wild populations can create natural hybrids.[66]
Asexual reproduction
Bulb
offsets: Daughter bulbs that form on the mother bulb and can be detached.
Scaling and
twin-scaling, used to increase production in slower-growing varieties, in which multiple whole scales are detached from a single bulb.
Bulb
offsets and
tissue culture produce
geneticclones of the parent plant and thus maintaining genetic integrity of the
cultivars. Bulb offsets usually require at least a year before flowering. Commercially, plants may be propagated
in vitro and then planted out to grow into plants large enough to sell.[69][70]
Toxicology
While members of the Liliaceae s.s. have been used as food sources in humans, the bulbs of some species are
poisonous to household pets (bulb toxicosis) if eaten and may cause serious complications, such as
kidney failure in
cats from Lilies, particularly Lilium longiflorum (Easter Lily).[71][72][73] Dogs may develop less serious effects such as gastrointestinal problems and central nervous system depression.[74] Most
Fritillaria (e.g. Fritillaria imperialis, Fritillaria meleagris) bulbs contain poisonous
neurotoxicalkaloids such as
imperialin (peiminine),[75][76][77] which may be deadly if ingested in quantity, while other species such as Fritillaria camschatcensis and Fritillaria affinis are edible.[26][27] Tulips can cause skin irritation due to the presence of tuliposides and tulipalins,[78][79][80] which are also found in the flowers, leaves and stems of Fritillaria.[76] These are also toxic to a variety of animals.[81]
The type genus, Lily (Lilium), has a long history in literature and art, and a tradition of
symbolism as well as becoming a
popular female name, and a
floral emblem, particularly of
France (
fleur-de-lis). The cultivation of lilies has been described since at least the ninth century, when
Charlemagne ordered it to be grown at his imperial palaces.[85] However, the name 'lily' has historically been applied to a wide variety of plants other than the genus Lilium.
The lily appears in ancient literature associated with both sovereignty and virginal innocence, and is mentioned on a number of occasions in the
Bible, such as the description in Solomon's
Song of Songs (2, 1–2) "I am a rose of Sharon, a lily of the valleys. Like a lily among thorns is my darling among the young women" or the
Gospel of Mathew (6, 28) "Consider the lilies of the field, how they grow; they toil not, neither do they spin" to represent beauty. Artistic representations can be found as far back as
frescos from the second century BC, at
Amnisos and
Knossos. Early Christian churches, such as that of the
Basilica of Sant' Apollinare in Classe were sometimes decorated with lilies. While predominantly depicted as white, those seen at
Akrotiri are red lilies. The white lily has long been seen as a symbol of purity, coming to be associated with the
Virgin Mary in the
Middle Ages, from which came the name ‘Madonna lily’ (Lilium candidum). A well-known example is
Leonardo da Vinci's Annunciation (1472–1475) in which the archangel
Gabriel bears a Madonna lily. Other symbolic meanings include glory, love and birth.
The stylised lily, or fleur-de-lis (lily flower) has long been associated with royalty, although it may originally have been derived from the form of an
iris. It has also been associated with the head of a spear.[86] Its three parts have been associated with the three classes of mediaeval society,[87] or alternatively faith, wisdom and chivalry.[88] Whatever its exact derivation, it has come to be associated with France and the French monarchy since the earliest
Frankish kings. Consequently, it became incorporated into not only French
heraldry but also into many heraldic devices in jurisdictions where there had been historic French influence, such as
Quebec and
New Orleans. In modern times it appears in many forms, symbolic and decorative, and can be used on
compasses to indicate the direction north, as well as the
Scouting movement.
Tulips (Tulipa) also have a long cultural tradition, particularly in the
Islamic world. The
Persian word for tulips, lâleh (لاله), was borrowed into
Turkish and
Arabic as lale. In
Arabic letters, lale is written with the same letters as Allah, and is used to denote God symbolically. Tulips became widely used in decorative
motifs on
tiles,
fabrics, and
ceramics in
Islamic art and the
Ottoman Empire in particular,[58] and were revered in poetry, such as that of
Omar Khayam in the twelfth century.[58][89]
Tulips were an essential part of the gardens of
Persia, having been cultivated from the
Steppes. As the
Seljuks –
Oghuz-Turkic leaders originated from the steppes – moved to Persia, and then west, they took tulips with them to
Turkey, where many escaped cultivation and became
naturalised. Today there are a number of places in Turkey called Laleli (‘with tulips’). Tulips first appeared in the decorative arts in Turkey in the thirteenth century and flourished under the Ottomans, in particular in the
royal palaces, and was adopted by the
Osmans as their symbol. Further species were collected from Persia and the spreading Ottoman Empire saw to it that tulip culture also spread. By the sixteenth century it was a national symbol, hence the designation "
Tulip era", by which time they were becoming of economic importance.[58]
By 1562 the tulip trade had reached Europe with the first shipment to
Antwerp,[90] where they were mistaken for vegetables,[91] although they had been cultivated in Portugal since 1530, and first appeared in illustration in 1559, and the first tulip flowers were seen in the
Netherlands in 1625. Tulips spread rapidly across Europe in the seventeenth century, and became an important trading item, initially in France before concentrating in the Netherlands.[58][92] Eventually speculative trading in tulips became so intense as to cause a
financial bubble which eventually collapsed, a period known as
tulip mania (tulipomania), from 1634 to 1637,[93][94] similar to the Ottoman Empire's Tulip era.
Nevertheless, since then the tulip has become indelibly associated with the Netherlands and all things Dutch. it was during this period that the
tulipiere, a
ceramic vase for growing tulips indoors was devised, and the
Golden Age of Dutch Painting was replete with images of tulips. Although
tulip festivals are held around the world in the spring, the most famous of these is the display at
Keukenhof. One of the better-known novels on tulips is The Black Tulip by
Alexandre Dumas, père in 1850, dealing with a contest to grow a black tulip in late seventeenth-century
Haarlem.
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