Liliales is anorder ofmonocotyledonousflowering plants in theAngiosperm Phylogeny Group andAngiosperm Phylogeny Websystem, within thelilioid monocots. This order of necessity includes thefamilyLiliaceae. TheAPG III system (2009) places this order in themonocotclade. In APG III, the familyLuzuriagaceae is combined with the familyAlstroemeriaceae and the familyPetermanniaceae is recognized. Both the order Lililiales and the family Liliaceae have had a widely disputed history, with thecircumscription varying greatly from one taxonomist to another. Previous members of this order, which at one stage included most monocots with conspicuoustepals and lackingstarch in theendosperm are now distributed over three orders, Liliales,Dioscoreales andAsparagales, using predominantlymolecular phylogenetics. The newly delimited Liliales ismonophyletic, with ten families. Well known plants from the order includeLilium (lily),tulip, the North American wildflowerTrillium, andgreenbrier.
Thus circumscribed, this order consists mostly ofherbaceous plants, butlianas andshrubs also occur. They are mostlyperennial plants, with foodstorage organs such ascorms orrhizomes. The familyCorsiaceae is notable for beingheterotrophic.
The order has worldwide distribution. The larger families (with more than 100 species) are roughly confined to theNorthern Hemisphere, or are distributed worldwide, centering on the north. On the other hand, the smaller families (with up to 10 species) are confined to theSouthern Hemisphere, or sometimes just toAustralia orSouth America. The total number of species in the order is now about 1768.
As with anyherbaceous group, thefossil record of the Liliales is rather scarce. There are several species from theEocene, such asPetermanniopsis anglesaensis orSmilax, but their identification is not definite. Another known fossil isRipogonum scandens from theMiocene. Due to the scarcity of data, it seems impossible to determine precisely the age and the initial distribution of the order. It is assumed that the Liliales originate from the LowerCretaceous, over 100 million years ago. Fossil aquatic plants from theCretaceous of northeastern Brazil and a new terrestrial species placed in the new genusCratosmilax suggest that the first species have appeared around 120 million years ago when the continents formed Pangea, before dispersing as Asia, Africa and America.[4] The initial diversification to the current families took place between 82 and 48 million years ago.[5] The order consists of 10 families, 67 genera and about 1,768 species.
The Liliales are a diverseorder of predominantlyperennial erect or twiningherbaceous andclimbing plants. Climbers, such as the herbaceousGloriosa (Colchicaceae) andBomarea (Alstroemeriaceae), are common in the Americas intemperate andtropical zones, while most species of the subtropical and tropical genusSmilax (Smilacaceae) are herbaceous or woody climbers and comprise much of the vegetation within the Liliales range. They also includewoodyshrubs, which have fleshystems and undergroundstorage orperennating organs, mainlybulbous geophytes, sometimesrhizomatous orcormous.[6]Leaves are elliptical and straplike withparallel venation or ovate with palmate veins andreticulate minor venation (Smilacaceae). InAlstroemeria andBomarea (Alstroemeriaceae) the leaves areresupinate (twisted).[7][8][9]
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Theflowers are highly variable, ranging in size from the small greenactinomorphic (radially symmetric) blooms ofSmilax to the large showy ones found inLilium,Tulipa andCalochortus (Liliaceae) andLapageria (Philesiaceae).Sepals andpetals are undifferentiated from each other, and known astepals, forming aperianth. They are usually large and pointed and may be variegated inFritillaria (Liliaceae).Nectaries may be perigonal (at base of tepals) but not septal (onovaries). Perigonal nectaries may be a simple secretory epidermal region at the tepal bases (Lapageria) or small, depressed regions fringed with hairs, often with glandular surface protuberances, at the bases of the inner tepals (Calochortus), while inTricyrtis the tepals become bulbous or spur-like at the base, forming a nectar-containing sac. Ovaries may be inferior or superior, the style often long andstigma capitate (pin headed). In a number of taxa there are three separate styles, particularly some Melanthiaceaes.l. (e.g.Helonias,Trillium,Veratrum) andChionographis. The outerintegument epidermis of theseed coat is cellular, and thephytomelanin pigment is lacking. The inner integument is also cellular and these features areplesiomorphic.[7][8][9]
The Liliales are characterised by (synapomorphies) the presence of nectaries at the base of the tepals (perigonal nectaries) orstamen filaments (Colchicum,Androcymbium) most taxa but the absence of septal nectaries,[10] together withextrorse (outward opening) anthers. This distinguishes them from the septal nectaries and introrse anthers that are the features of most othermonocots.[5][8] Exceptions are some Melanthiaceae in which nectaries are absent or septal and anthers that are introrse (dehiscence directed inwards) in Campynemataceae, Colchicaceae, and some Alstroemeriaceae, Melanthiaceae, Philesiaceae, Ripogonaceae and Smilacaceae. Tepals are largely three-traced in net-veined taxa of Liliales (e.g.Clintonia,Disporum), distinguishing them from the single-traced Asparagales, and is associated with the presence of tepal nectaries, presumably to supply them. The presence of separate styles is also a distinguishing feature from Asparagales, where it is rare.Phytomelan is completely absent in Liliales seed coats, unlike Asparagales, which nearly all contain it.[11][8]
The stems containfructans, the plants also containchelidonic acid,saponins, while some species containvelamen. Theepicuticular wax is of the Convallaria type, consisting of parallel orientated platelets.[12]
The order includestaxa with some of the largestgenomes amongAngiosperms,[13] particularlyMelanthiaceae,Alstroemeriaceae andLiliaceae.[14]
With 11families, about 67genera and about 1,558species, Liliales is a relatively smallangiospermorder, but a large group within the monocotyledons.[9][15]
Thebotanical authority for Liliales is given toPerleb (1826), who grouped eleven families (Asparageae,Pontederiaceae,Asphodeleae,Coronariae,[a]Colchicaceae,Dioscoreaceae,Hypoxideae,Amaryllideae,Haemodoraceae,Burmanniaceae,Irideae) into an order he called Liliaceae.[17] In Perleb's system, he divided thevascular plants into seven classes, of which thePhanerogamicae or seed plants he called his class IV, or Ternariae. The latter, he divided into five orders (ordo), including the Liliaceae.[17]
A number of latertaxonomists, such as Endlicher (1836) substitituted the term Coronarieae for this higher order, including six subordinate taxa. Endlicher divided theCormophyta into five sections, of which Amphibrya contained eleven classes, including Coronarieae.[18] The term Liliales was introduced byLindley (1853),[19] referring to these higher orders asalliances. Lindley included four families in this alliance. Lindley called the monocots class Endogenae, with eleven alliances including Liliales.[19] AlthoughBentham (1877) restored Coronariae as one of sevenSeries making up themonocotyledons,[20] it was replaced by Liliiflorae and then Liliales in subsequent publications (seeTable for history).[21]
Subsequent authors, now adopting aphylogenetic (phyletic) orevolutionary approach over the natural method,[22] did not follow Bentham's nomenclature.Eichler (1886) used Liliiflorae for the higher order including Liliaceae, placing it as the first order (Reihe) in his class monocotyledons,[23] as didEngler (1903),[24]Lotsy (1911),[25] andWettstein in 1924, in class Monocotyledones, subdivision Angiospermae.[26]
Hutchinson (1973)[27] restored Liliales for the higher rank, an approach that has been adopted by most major classification systems onwards, reserving Liliiflorae for higher ranks. These includeCronquist (1981),[28]Dahlgren (1985),[29]Takhtajan (1997)[30] as well asThorne andReveal (2007).[31]
Hutchinson (1973) derived a more elaborate hierarchy, placing order Liliales as one of 14 in division Corolliferae, one of three divisions ofsubphylum monocotyledons.Cronquist (1981) placed the order Liliales as one of two in subclassLiliidae, one of five in the classLiliopsida (monocotyledons) of divisionMagnoliophyta (angiosperms).Dahlgren (1985) made Liliales one of six orders in Superorder Liliiflorae, one of ten divisions of the monocots.Takhtajan (1997) had a more complex system of higher taxonomic ranks, placing Liliales as one of 15 orders withinsuperorderLilianae, one of four within subclassLiliidae. Liliidae in turn was one of four subclasses in classLiliopsida (monocots).[32] In contrastThorne and Reveal (2007) abandoned the use of monocotyledons as a distinct taxon, replacing it with 3 separate subclasses of Magnoliopsida (angiosperms), of whichLiliidae consists of 3 superorders, placing Liliales in superorderLilianae.[31]
In all these systems, Liliales (or Liliiflorae) were visualised as either a direct division of the monocots (or equivalent) or were placed in an intermediate division of the monocots, such as superorder Lilianae.[33]
The development ofmolecular phylogenetic methods for determiningtaxonomiccircumscription andphylogeny led to considerable revision of angiosperm classification,[34] and establishment of Liliales as amonophyletic group.[35][36][8] It was clear by 1996, that the most useful system to date, that of Dahlgren, required urgent revision.[34] The new classification was formalised with the creation of theAngiosperm Phylogeny Group (APG) system (1998–2016),[37][38] based onmonophyleticclades, which continued the use of Liliales as the name for the taxon.[39]
The Angiosperm Phylogeny GroupAPG system (1998) established a structure of monocot classification with ten orders.[37] Notable was the separation of asparagids, as suggested by Dahlgren,[10] intoAsparagales, with other taxa placed inDioscoreales, resulting in a much reduced order.[9][8]
The position of Liliales within the monocots (Lilianae) is shown in the followingcladogram. The monocot orders form threegrades, thealismatid monocots,lilioid monocots and thecommelinid monocots by order of branching, from early to late. These have alternatively been referred to as Alismatanae, Lilianae and Commelinanae.[10] The alismatid monocots form thebasal group, while the remaining grades (lilioid and commelinid monocots) have been referred to as the "core monocots".[40] The relationship between the orders (with the exception of the two sister orders) ispectinate, that is diverging in succession from the line that leads to the commelinids.[41] The lilioid monocot orders constitute aparaphyletic assemblage, that is groups with a common ancestor that do not include all direct descendants (in this case commelinids which are asister group to Asparagales); to form a clade, all the groups joined by thick lines would need to be included. In the cladogram the numbers indicatecrown group (most recent common ancestor of the sampled species of the clade of interest)divergence times inmya (million years ago).[42]
Cladogram 1: The phylogenetic composition of the monocots[38][43]
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Thecrown group of Liliales has been dated to ca. 117 Myr (million years ago) in theEarly Cretaceous period of theMesozoic era.[45][40]
The circumscription of Liliales has varied greatly since Perleb's original construction with 11 families in 1826.[8] Many of these families are now considered to be inAsparagales, with the remainder incommelinids andDioscoreales, as shown in this table.
| Perleb (1826)[17] | Endlicher (1836)[18] | Lindley (1853)[19] | Bentham & Hooker (1883)[20] | Eichler (1886)[23] | Engler (1903)[24] | Lotsy (1911)[25] | Wettstein (1924)[26] | Hutchinson (1973)[27] | Cronquist (1981)[28] | Dahlgren (1985)[29] | Takhtajan (1997)[30] | Thorne & Reveal (2007)[31] | APG IV (2016)[38] |
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| Liliaceae | Coronarieae | Liliales | Coronariae | Liliiflorae | Liliiflorae | Liliiflorae | Liliiflorae | Liliales | Liliales | Liliales | Liliales | Liliales | Liliales |
| AsparageaeA | Asparagaceae | ||||||||||||
| AsphodeleaeA | Asphodelaceae | ||||||||||||
| Colchicaceae | Colchicaceae | Colchicaceae | Colchicaceae | ||||||||||
| Coronariae | Liliaceae | Liliaceae | Liliaceae | Liliaceae | Liliaceae | Liliaceae | Liliaceae | Liliaceae | Liliaceae | Liliaceae | Liliaceae | Liliaceae | Liliaceae |
| AmaryllideaeA | Amaryllidaceae | Amaryllidaceae | Amaryllidaceae | ||||||||||
| Pontederiaceaec | Pontederiaceae | Pontederiaceae | Pontederiaceae | Pontederiaceae | Pontederiaceae | Pontederiaceae | |||||||
| DioscoreaceaeD | Dioscoreaceae | Dioscoreaceae | Dioscoreaceae | Dioscoreaceae | Dioscoreaceae | ||||||||
| HypoxideaeA | Hypoxidaceae | ||||||||||||
| Haemodoraceaec | Haemodoraceae | Haemodoraceae | Haemodoraceae | Haemodoraceae | Haemodoraceae | ||||||||
| BurmanniaceaeD | Burmanniaceae | Burmanniaceae | |||||||||||
| IrideaeA | Iridaceae | Iridaceae | Iridaceae | Iridaceae | Iridaceae | Iridaceae | |||||||
| Juncaceaec | Juncaceae | Juncaceae | Juncaceae | Juncaceae | |||||||||
| Philydriaec | Philydraceae | Philydraceae | Philydraceae | ||||||||||
| Melanthaceae | Melanthaceae | Melanthaceae | Melanthiaceae | Melanthiaceae | Melanthiaceae | ||||||||
| Smilaceae | Smilaceae | Smilacaceae | Smilacaceae | Smilacaceae | Smilacaceae | ||||||||
| GilliesiaceaeA | Gilliesiaceae | ||||||||||||
| RoxburghiaceaeP | Stemonaceae | Stemonaceae | Stemonaceae | Stemonaceae | |||||||||
| Xyrideaec | |||||||||||||
| Mayaceaec | |||||||||||||
| Commelinaceaec | |||||||||||||
| Rapateaceaec | Rapateaceae | ||||||||||||
| Bromeliaceaec | Bromeliaceae| | Bromeliaceae | |||||||||||
| VelloziaceaeP | Vellosiaceae | Velloziaceae | Velloziaceae | ||||||||||
| TaccaceaeD | Taccaceae | Taccaceae | Taccaceae | ||||||||||
| AloinaceaeA | |||||||||||||
| EriospermaceaeA | |||||||||||||
| JohnsoniaceaeA | |||||||||||||
| AgapanthaceaeA | |||||||||||||
| AlliaceaeA | |||||||||||||
| Tulipaceae | |||||||||||||
| ScillaceaeA | |||||||||||||
| DracaenaceaeA | |||||||||||||
| Luzuriagaceae | Luzuriagaceae | ||||||||||||
| OphiopogonaceaeA | |||||||||||||
| LomandraceaeA | |||||||||||||
| Dasypogonaceaec | |||||||||||||
| Calectasiaceaec | |||||||||||||
| Flagellariaceaec | |||||||||||||
| Cyanastraceaec | Cyanastraceae | Cyanastraceae | |||||||||||
| AgavaceaeA | Agavaceae | ||||||||||||
| TecophilaeaceaeA | |||||||||||||
| Trilliaceae | Trilliaceae | (in Melanthiaceae) | |||||||||||
| RuscaceaeA | |||||||||||||
| XanthorrhoeaceaeA | |||||||||||||
| Alstroemeriaceae | Alstroemeriaceae | Alstroemeriaceae | |||||||||||
| Uvulariaceae | (in Colchicaceae Liliaceae) | ||||||||||||
| Calochortaceae | (in Liliaceae) | ||||||||||||
| GeosiridaceaeA | |||||||||||||
| Medeolaceae | (in Liliaceae) | ||||||||||||
| Corsiaceae | Corsiaceae | ||||||||||||
| Campynemataceae | Campynemataceae | ||||||||||||
| Petermanniaceae | Petermanniaceae | ||||||||||||
| Rhipogonaceae | Ripogonaceae | ||||||||||||
| Philesiaceae | Philesiaceae | ||||||||||||
Treatment of families in modern taxonomy (APG), remaining families included in Liliales:
| |||||||||||||
The availability of molecular phylogenetic methods suggested four main lineages within Liliales, and seven families;[8]
The first Angiosperm Phylogeny Group classification (APG I) in 1998 had the following circumscription, with 9 families, having separated Philesiaceae and Ripogonaceae from Smilacaceae:[37]
TheAPG II system (2003) addedCorsiaceae to the Liliales,[46] whileAPG III (2009) addedPetermanniaceae and merged Luzuriagaceae into Alstroemeriaceae.[2] The subsequent revision ofAPG IV (2016) left this unchanged, with 10 families.[38]
The exact phylogenetic relationship between the families of Liliales has been subject to revision. This cladogram shows that of theAngiosperm Phylogeny Website (2020):[47][11]
| Liliales |
| |||||||||
The bulk of the Liliales species are found in the family Liliaceae (16 genera, 610 species). Of the remaining nine families, three are referred to as the vine families (Ripogonaceae, Philesiaceae and Smilacaceae) and form a cluster.[11]
 Corsiaceae[edit]Main article:Corsiaceae The Corsiaceae (ghost-flower family) are a family of 3mycoheterotrophic genera, lackingchlorophyll, with 27 species of perennial herbaceous plants. They are found in montane forests in South America (one genus) and from southern China to northern Australia in areas with high rainfall, and among dense leaf litter. The majority of species occur in the type genusCorsia. The name commemorates the Florentine plant collector Marquis Bardo Corsi Salviati (1844–1907).[48][49] |
 Campynemataceae[edit]Main article:Campynemataceae The Campynemataceae (Green-mountainlily family) are a family of two genera and four species of rhizomatous herbaceous plants found in Tasmania andNew Caledonia. The name is derived from theGreek wordskampylos (curved) and nema (thread).[48][49] |
 Melanthiaceae[edit]Main article:Melanthiaceae The Melanthiaceae (Wake Robin family) is a family of perennial herbaceous plants, whose storage organs include bulbs, rhizomes and corms (rarely, e.g.Schoenocaulon). Their distribution is temperate andboreal Northern hemisphere, in the Americas extending south to the Andes and in Asia to the Himalayas and Taiwan. Melanthiaceae consists of 17 genera and 173 species distributed in a number of subdivisions. The largest genus isTrillium (44 species) but many genera are monotypic. A number of genera, includingTrillium are used asgarden ornamentals, especially for woodland gardens.Paris japonica is noted for having the largestgenome known to date. The family name is derived from the Greek wordsmelas (black) andanthos (flower) in reference to the dark colour of the petals.[48][49] |
 Petermanniaceae[edit]Main article:Melanthiaceae The Petermanniaceae (Petermann's vine family) consists of a single species,Petermannia cirrosa, a perennial woody vine with underground rhizomes.Petermannia is restricted to Queensland and New South Wales, in temperate rainforests between Brisbane and Sydney. The family was named for Wilhelm Ludwwig Petermann (1806–1855), director of thebotanical garden at Leipzig.[48][49] |
 Colchicaceae[edit]Main article:Colchicaceae The Colchicaceae (Naked-ladies or Colchicum family) are perennial erect and climbing plants with underground corms, tubers and rhizomes. They are herbaceous with the exception ofKuntheria which has a somewhat woody stem. Their distribution is widespread including in temperate zones in North America, Europe, North Africa and the Middle east and tropical zones in Africa, Asia and Australasia. They are absent from South America. The family is of medium size with 15 genera and about 285 species. The largest genus is the type genus,Colchicum, with 159spp. Although the alkaloids, which characterise them, they contain are toxic to animals and humans, Colchicine has usage medicinally and in botanical laboratories. They are also include popular garden and indoor ornamentals. These includeColchicum andGloriosa. The family is named afterColchis on the eastern Black Sea.[48][49][9] |
.JPG) Alstroemeriaceae[edit]Main article:Alstroemerieae The Alstroemeriaceae (Inca-lily family) are erect or creeping perennial (rarely annual) herbaceous plants with occasional shrubby vines, some of which have evergreen stems. They are occasionally epiphytic and form often swollen rhizomes. They are found in tropical and temperate Central and South America, as well as Australasia. There are two large genera (Alstroemerieae), the erectAlstroemeria (S America 125 spp.) and twiningBomarea (Central & S America 122 spp.) and two genera (Luzuriageae) with 2 and 4 species each, for a total of 253 species in the family. Two species are widely used for food in S America,Alstroemeria ligtu is used for a flour (Chuño) that is extracted from its roots, while the tubers ofBomarea edulis are directly consumed.Luzuriaga radicans, also from S America, produces fibre used in rope making.Alstroemeria cultivars are popular ornamentals and widely used ascut flowers (Peruvian lilies). The family is named forBaron Clas Alströmer (1736–1794), a student of Linnaeus.[48][49] |
 Ripogonaceae[edit]Main article:Ripogonaceae The Ripogonaceae (Supplejack family) is a family with a single genus,Ripogonum, and six species. They are woody evergreen shrubs and vines arising from a horizontal rhizome, swollen at its base to form a tuber. They are confined to Eastern Australasia, with the type species,Ripogonum scandens as the sole New Zealand species. The stems have a use in basketry and building and the young shoots are edible. The name is derived from two Greek words,ripos (wicker) andgony (node) in reference to their node bearing shoots.[48][49] |
 Philesiaceae[edit]Main article:Philesiaceae The Philesiaceae (Chilean-bellflower family) are a family consisting of two monotypic genera, the two species beingPhilesia magellanica and the similarLapageria rosea. They grow from a short woody rhizome, forming shrubs and vines respectively. They are found in the cooltemperate forest of central and southern Chile,Magellan straits and adjacent Argentina, among the southern beech (Nothofagus) trees.Lapageria is thenational flower of Chile and a popular ornamental with edible fruit. The name is thought to be related to the Greek wordphileo (love), because of the attractiveness of its flowers.[48] |
 Smilacaceae[edit]Main article:Smilacaceae The Smilacaceae (Catbrier family) consist of a single large genus,Smilax, with about 210 species,[50] making it the second largest family of the order, after Liliaceae. They are perennial vines, shrubs or herbaceous, sometimes woody, plants with short fibrous woody (sometimes tuberous) rhizomes. Smilacaceae are pantropical with extension into temperate zones north (N America, Mediterranean, Russian Far East) and south (Eastern Australia). A number of species have been used in traditional medicine and as foodstuffs.Smilax china was used to treatgout.S. aristolochiifolia was used to treatsyphilis (but later as sarsaparilla to flavorroot beer and confectionary). The fruit ofS. megacarpa is consumed inconserves. The young shoots of many species are also edible. The family is named after the Greek myth of the affair between the mortal Krokos (orCrocus) and the nymph Smilax, whose punishment was to be turned into the prickly vineSmilax aspera.[48] |
 Liliaceae[edit]Main article:Liliaceae The lily family, Liliaceae, are the largest Liliales family, with 15 genera and about 700 species, though much reduced from earlier circumscriptions, in four subfamilies. Of these genera,Gagea is the largest (204 spp.), but some are quite small, withMedeola being monotypic. They are perennial herbaceous plants, growing from bulbs or corms (rarely creeping rhizomes), with actinomorphic hypogynous flowers that are often coloured and patterned. They are predominantly northern temperate in distribution, with extension to subtropical areas of N Africa, India, China andLuzon, but are absent from the southern hemisphere. The bulbs have been used as foodstuffs or in traditional medicine.Cardiocrinum cordatum andErythronium japonicum are sources of starch. Many Liliaceae are important in the floriculture and horticulture industries, particularlyTulipa andLilium, but alsoFritillaria. Many are also important ornamentals, such asCalochortus,Cardiocrinum,Clintonia,Erythronium andTricyrtis. The name is derived from the Latin word for lily,lilium, which in turn is derived from the Greekleirion, a white lily.[48][49][40] |
Widely distributed but most commonly found insubtropical andtemperate regions, especially herbaceous taxa in temperate regions of theNorthern Hemisphere, and subtropical regions of the Southern hemisphere, including vines.[8] Since many species arecultivated they have beenintroduced in many regions and consequently worldwide, and a number have subsequently escaped andnaturalised.[9]
Liliales form important sources of food andpharmaceuticals as well as playing a significant role inhorticulture andfloriculture asornamental plants. Pharmaceutical products includecolchicine fromColchicum andGloriosa (Colchicaceae) andveratrine and related compounds fromVeratrum (Melanthiaceae) andZigadenus (Melanthiaceae).[9]
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