Generalfloral formula of the Liliacaeae:Flowers actinomorphic or slightly zygomorphic and hermaphrodite with 6undifferentiated tepals intwo whorls of three, the samenumber and arrangement ofstamens, and asuperior ovary with3 fused carpels. Individual species and genera may have more or less derived formulas.
The lilyfamily,Liliaceae, consists of about 15genera and 610species offlowering plants within theorderLiliales.[2] They aremonocotyledonous,perennial,herbaceous, oftenbulbousgeophytes. 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 patternedpetaloidtepals (undifferentiated petals and sepals) arranged in twowhorls, sixstamens and a superiorovary. 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 haverhizomes. First described in 1789, the lily family became aparaphyletic "catch-all" (wastebasket) group oflilioid 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 68million years ago during theLate Cretaceous toEarly Paleogene epochs. Liliaceae are widely distributed, mainly in temperate regions of the Northern Hemisphere and the flowers are insect pollinated. Many Liliaceae are importantornamental plants, widely grown for their attractive flowers and involved in a majorfloriculture ofcut flowers and dry bulbs. Some species arepoisonous 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 (seeEvolution).[6]
The Liliaceae are characterised as monocotyledonous, perennial, herbaceous, bulbous (orrhizomatous in the case ofMedeoleae)[7] flowering plants with simpletrichomes (root hairs) and contractileroots.[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 theFritillaria 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.
Usuallyindeterminate (lacking terminal flower) as araceme (Lilium); sometimes reduced to a single terminal flower (Tulipa). Whenpluriflor (multiple blooms), the flowers are arranged in a cluster or rarely aresubumbellate (Gagea) or athyrse (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. Theperianth is undifferentiated (perigonium) and biseriate (twowhorled), formed from sixtepals arranged into two separate whorls of three parts (trimerous) each, althoughScoliopus has only three petals, free from the other parts, but overlapping. The tepals are usually petaloid (petal like) andapotepalous (free) with lines (striate) or marks in other colors or shades. Theperianth is either homochlamydeous (alltepals 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]
Sixstamens 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), althoughScoliopus 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) anddehisce longitudinally and are extrorse (dehiscing away from center). Thepollen is usually monosulcate (single groove), but may be inaperturate (lacking aperture:Clintonia, someTulipa spp.) or operculate (lidded:Fritillaria, someTulipa spp.),[15] and reticulate (net patterned:Erythronium,Fritillaria,Gagea,Lilium,Tulipa).[9][11][14]
Superiorovary (hypogynous), syncarpous (with fused carpels), with three connate (fused) carpels and is trilocular (threelocules, or chambers) or unilocular (single locule, as inScoliopus andMedeola). There is a single style and a three lobedstigma or three stigmata more or less elongated along the style. There are numerous anatropous (curved)ovules which display axileplacentation (parietal inScoliopus andMedeola),[14] usually with anintegument and thinnermegasporangium. The embryo sac (megagametophyte) varies by genera, but is mainlytetrasporic (e.g.Fritillaria).[11] Embryo sacs in which three of the fourmegaspores fuse to form atriploidnucleus, are referred to asFritillaria-type, a characteristic shared by all the core Liliales.[6][9][11][16]
Acapsule that is usually loculicidal (splitting along thelocules) as in the Lilioideae,[6] but occasionally septicidal (splitting between them, along the separatingsepta) in the Calachortoideae and wind dispersed, although the Medeoleae formberries (baccate).[6] Theseeds may be flat, oblong, angular,discoid,ellipsoid orglobose (spherical), or compressed with a well developedepidermis. The exterior may be smooth or roughened, with a wing orraphe (ridge),aril or one to two tails, rarely hairy, but may be dull or shiny and the lack of a blackintegument distinguishes them from related taxa such asAllioideae that were previously included in this family, and striate (parallel longitudinally ridged) in the Steptopoideae. Thehilum (scar) is generally inconspicuous. The bitegmic (separate testa and tegmen) seed coat itself may be thin, suberose (likecork), or crustaceous (hard or brittle). Theendosperm is abundant,cartilaginous (fleshy) or horny and contains oils andaleurone but notstarch (non-farinaceous). Its cells arepolyploid (triploid orpentaploid, depending on theembryo sac type). Theembryo is small (usually less than one quarter of seed volume), axile (radially sectioned), linear (longer than broad) or rarelyrudimentary (tiny relative toendosperm) 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) andspiral, but may bewhorled (three or more attached at onenode, e.g.Lilium,Fritillaria), cauline (arranged along the aerialstem) or sheathed in a basalrosette. They are rarelypetiolate (stem attached before apex), and lackstipules. The aerial stem is unbranched.[9][11][12][19][20]
The Liliaceae include a species with one of the largestgenome size within the angiosperms,Fritillaria assyriaca (1C=127.4 pg), whileTricyrtis macropoda is as small as 4.25 pg.[21]Chromosome numbers vary by genus.[11] Some genera likeCalochortus (x=6-10),Prosartes (6,8,9,11),Scoliopus (7,8),Streptopus (8, 27) andTricyrtis (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, betweenshade-dwelling genera (such asProsartes,Tricyrtis,Cardiocrinum,Clintonia,Medeola,Prosartes, andScoliopus) 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 alsoEvolution).
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 theAPG which is based onphylogenetic principles usingmolecular 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, theLatin termssensu lato andsensu stricto are frequently used (together with their abbreviations,s.l. ands.s.) to denote the broader or stricter sense of the circumscription respectively, e.g. Liliaceaes.s..
The family Liliaceae was described byMichel Adanson in 1763 and formally named byAntoine Laurent de Jussieu in 1789. Jussieu defined this grouping as having acalyx of six equal colored parts, sixstamens, a superiorovary, singlestyle, and a trilocular (three-chambered)capsule. By 1845,John Lindley, the first Englishsystematist, unhappily acknowledged the great diversity in thecircumscription of the family, and that it had expanded vastly, with many subdivisions. As he saw it, the Liliaceae were alreadyparaphyletic ("catch-all"), being allLiliales 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 ofBentham 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 theorderLiliales under theCronquist system (1981). Cronquist merged the Liliaceae with theAmaryllidaceae, making this one of the largest monocotyledon families.[30]
Many other botanists echoed Lindley's earlier concerns about thephylogeny 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 (predominantlyLiliales andAsparagales).[31][32] In the context of a general review of the classification ofangiosperms, the Liliaceae were subjected to more intense scrutiny. Considerable progress in plant phylogeny andphylogenetic theory enabled aphylogenetic tree to be constructed for all of the flowering plants, as elaborated by theAngiosperm Phylogeny Group (1998).[6]
The Angiosperm Phylogeny Group (APG) made rapid progress in establishing a modernmonophyletic classification of the flowering plants by 2009.[33] Despite establishing this relative degree ofmonophyly (genetic homogeneity) for the family Liliaceae,[34][35] their morphology remains diverse[7] and there exists within the Liliaceaeclade or grouping, a number of subclades (subgroups). Particularly enigmatic wereClintonia,Medeola,Scoliopus, andTricyrtis.
Of the fifteen genera within the Liliaceae, the ten genera of theLilioideaesubfamily form one morphological group that is characterised by contractile bulbs and roots, and a Fritillaria-typeembryo-sac (megagametophyte with four megaspores). Within the Lilioideae,Clintonia and the closely relatedMedeola form a subclade, and are now considered a separate tribe (Medeoleae).[36] The other major grouping consists of the five genera constituting theStreptopoideae (includingScoliopus) andCalochortoideae (includingTricyrtis) subfamilies characterised by creeping rhizomes, styles which are divided at their apices, and by megagametophyte development of thePolygonum-type (a simple megaspore and triploid endosperm) embryo-sac.
The Liliaceae probably arose asshade plants, with subsequent evolution to open areas includingdeciduous forest in the more open autumnal period, but then a return of some species (e.g.Cardiocrinum). This was accompanied by a shift fromrhizomes tobulbs, 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 Liliaceaes.l., many attempts have been made to form suprageneric classifications, e.g.subfamilies andtribes.[23] Classifications published since the use ofmolecular methods in phylogenetics have taken a narrower view of the Liliaceae (Liliaceaes.s.). TheAngiosperm 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 Liliaceaes.s., but generally there are about fifteen to sixteen genera, depending on whether or notAmana is included inTulipa andLloydia inGagea.
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 thetemperate regions of theNorthern Hemisphere. Thecentre of diversity is fromsouthwest Asia toChina. Their distribution is diverse, mainly inplains,steppes, andalpine meadows, but also indeciduous forests,Mediterranean scrub andarctic tundra.[6][9]Tulipa andGagea 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, andCardiocrinum, others prefer a more open habitat.
The Liliaceae are ecologically diverse.[6] Species of Liliaceae bloom at various times from spring to late summer. The colorful flowers produce large amounts ofnectar andpollen that attractinsects whichpollinate them (entomophily), particularlybees andwasps (hymenopterophily),butterflies (psychophily) andmoths (phalaenophily).[20] Theseeds aredispersed by wind and water. Some species (e.g.Scoliopus,Erythronium and Gagea) have seeds with anaril structure that are dispersed byants (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 genusLilium are particularly palatable, while species inFritillaria are repellant.
BothLilium andTulipa are susceptible to a group of five viruses of the familyPotyviridae, specifically thepotyvirus (named forpotato virus Y) group, which includes thetulip-breaking virus (TBV) and thelily streak virus (lily mottle virus,LMoV) resulting in 'breaking' of the color of the flowers. The viruses are transmitted byaphids. This breaking effect was of economic importance during thetulip 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 theDutch 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 theRembrandt tulip-breaking virus (ReTBV).[56][57]
Tulips have been cultivated since at least the tenth century inPersia.[58] Tulip production has two main markets:cut flowers andbulbs. 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 billioneuros, which provides an indication of the economic importance of this activity. The main producer of tulip bulbs is theNetherlands, a country that accounts for 87% of the global cultivated area, with approximately 12,000hectares. Other leading producers includeJapan,France andPoland. Approximately ten other countries produce commercial tulips, largely for thedomestic 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]
OriginalTulipa species can be obtained for ornamental purposes, such asTulipa tarda andTulipa turkestanica. These are referred to asspecies, or botanical, tulips, and tend to be smaller plants but better atnaturalising 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]
The largest area of production is also the Netherlands, with 76% of the global cultivated area, followed byFrance,Chile,Japan, theUnited States,New Zealand andAustralia. 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 theEuropean Union. One particularly important crop is the production ofLilium longiflorum, whose white flowers are associated with purity and Easter.[59]
Although manyLilium species such asLilium martagon andLilium 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 asWatsonia (bugle lily) that were previously classified in this family bear the name 'lily' but are neither part of the genusLilium, or the family Liliaceae.
A variety ofFritillaria species are used as early spring ornamental flowers. These vary from the largeFritillaria imperialis (crown imperial) available in a number of colours such as yellow or orange, to much smaller species such asFritillaria meleagris orFritillaria 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 createhybrids and can take five to eight years to produce flowering plants. Sinceinterspecificcross-pollination occurs, overlapping wild populations can create natural hybrids.[66]
Scaling andtwin-scaling, used to increase production in slower-growing varieties, in which multiple whole scales are detached from a single bulb.
Bulboffsets andtissue culture producegeneticclones of the parent plant and thus maintaining genetic integrity of thecultivars. Bulb offsets usually require at least a year before flowering. Commercially, plants may be propagatedin vitro and then planted out to grow into plants large enough to sell.[69][70]
While members of the Liliaceaes.s. have been used as food sources in humans, the bulbs of some species arepoisonous to household pets (bulb toxicosis) if eaten and may cause serious complications, such askidney failure incats from Lilies, particularlyLilium longiflorum (Easter Lily).[71][72][73] Dogs may develop less serious effects such as gastrointestinal problems and central nervous system depression.[74] MostFritillaria (e.g.Fritillaria imperialis,Fritillaria meleagris) bulbs contain poisonousneurotoxicalkaloids such asimperialin (peiminine),[75][76][77] which may be deadly if ingested in quantity, while other species such asFritillaria camschatcensis andFritillaria 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 ofFritillaria.[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 ofsymbolism as well as becoming apopular female name, and afloral emblem, particularly ofFrance (fleur-de-lis). The cultivation of lilies has been described since at least the ninth century, whenCharlemagne 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 genusLilium.
The lily appears in ancient literature associated with both sovereignty and virginal innocence, and is mentioned on a number of occasions in theBible, such as the description in Solomon'sSong 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 theGospel 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 asfrescos from the second century BC, atAmnisos andKnossos. Early Christian churches, such as that of theBasilica of Sant' Apollinare in Classe were sometimes decorated with lilies. While predominantly depicted as white, those seen atAkrotiri are red lilies. The white lily has long been seen as a symbol of purity, coming to be associated with theVirgin Mary in theMiddle Ages, from which came the name ‘Madonna lily’ (Lilium candidum). A well-known example isLeonardo da Vinci'sAnnunciation (1472–1475) in which the archangelGabriel bears a Madonna lily. Other symbolic meanings include glory, love and birth.
The stylised lily, orfleur-de-lis (lily flower) has long been associated with royalty, although it may originally have been derived from the form of aniris. 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 earliestFrankish kings. Consequently, it became incorporated into not only Frenchheraldry but also into many heraldic devices in jurisdictions where there had been historic French influence, such asQuebec andNew Orleans. In modern times it appears in many forms, symbolic and decorative, and can be used oncompasses to indicate the direction north, as well as theScouting movement.
Tulips were an essential part of the gardens ofPersia, having been cultivated from theSteppes. As theSeljuks –Oghuz-Turkic leaders originated from the steppes – moved to Persia, and then west, they took tulips with them toTurkey, where many escaped cultivation and becamenaturalised. Today there are a number of places in Turkey calledLaleli (‘with tulips’). Tulips first appeared in the decorative arts in Turkey in the thirteenth century and flourished under the Ottomans, in particular in theroyal palaces, and was adopted by theOsmans 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 toAntwerp,[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 theNetherlands 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 afinancial bubble which eventually collapsed, a period known astulip 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 thetulipiere, aceramic vase for growing tulips indoors was devised, and theGolden Age of Dutch Painting was replete with images of tulips. Althoughtulip festivals are held around the world in the spring, the most famous of these is the display atKeukenhof. One of the better-known novels on tulips isThe Black Tulip byAlexandre Dumas, père in 1850, dealing with a contest to grow a black tulip in late seventeenth-centuryHaarlem.
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