The order has worldwide distribution. The larger families (with more than 100 species) are roughly confined to the Northern Hemisphere, or are distributed worldwide, centering on the north. On the other hand, the smaller families (with up to 10 species) are confined to the Southern Hemisphere, or sometimes just to Australia or South America. The total number of species in the order is now about 1768.
As with any herbaceous group, the fossil record of the Liliales is rather scarce. There are several species from the Eocene, such as Petermanniopsis anglesaensis or Smilax, but their identification is not definite. Another known fossil is Ripogonum scandens from the Miocene. 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 Lower Cretaceous, over 100 million years ago. Fossil aquatic plants from the Cretaceous of northeastern Brazil and a new terrestrial species placed in the new genus Cratosmilax 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.
Description
The Liliales are a diverse order of predominantly perennial erect or twining herbaceous and climbing plants. Climbers, such as the herbaceous Gloriosa (Colchicaceae) and Bomarea (Alstroemeriaceae), are common in the Americas in temperate and tropical zones, while most species of the subtropical and tropical genus Smilax (Smilacaceae) are herbaceous or woody climbers and comprise much of the vegetation within the Liliales range. They also include woodyshrubs, which have fleshy stems and underground storage or perennating organs, mainly bulbous geophytes, sometimes rhizomatous or cormous.[6]Leaves are elliptical and straplike with parallel venation or ovate with palmate veins and reticulate minor venation (Smilacaceae). In Alstroemeria and Bomarea (Alstroemeriaceae) the leaves are resupinate (twisted).[7][8][9]
The flowers are highly variable, ranging in size from the small green actinomorphic (radially symmetric) blooms of Smilax to the large showy ones found in Lilium, Tulipa and Calochortus (Liliaceae) and Lapageria (Philesiaceae). Sepals and petals are undifferentiated from each other, and known as tepals, forming a perianth. They are usually large and pointed and may be variegated in Fritillaria (Liliaceae). Nectaries may be perigonal (at base of tepals) but not septal (on ovaries). 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 in Tricyrtis 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 and stigma capitate (pin headed). In a number of taxa there are three separate styles, particularly some Melanthiaceae s.l. (e.g. Helonias, Trillium, Veratrum) and Chionographis. The outer integument epidermis of the seed coat is cellular, and the phytomelanin pigment is lacking. The inner integument is also cellular and these features are plesiomorphic.[7][8][9]
The Liliales are characterised by (synapomorphies) the presence of nectaries at the base of the tepals (perigonal nectaries) or stamen filaments (Colchicum, Androcymbium) most taxa but the absence of septal nectaries,[10] together with extrorse (outward opening) anthers. This distinguishes them from the septal nectaries and introrse anthers that are the features of most other monocots.[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]
A number of later taxonomists, such as Endlicher (1836) substitituted the term Coronarieae for this higher order, including six subordinate taxa. Endlicher divided the Cormophyta into five sections, of which Amphibrya contained eleven classes, including Coronarieae.[18] The term Liliales was introduced by Lindley (1853),[19] referring to these higher orders as Alliances. Lindley included four families in this alliance. Lindley called the monocots class Endogenae, with eleven alliances including Liliales.[19] Although Bentham (1877) restored Coronariae as one of seven Series making up the monocotyledons,[20] it was replaced by Liliiflorae and then Liliales in subsequent publications (see Table for history).[21]
Phyletic systems
Subsequent authors, now adopting a phylogenetic (phyletic) or evolutionary 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 did Engler (1903),[24]Lotsy (1911),[25] and Wettstein 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 include Cronquist (1981),[28]Dahlgren (1985),[29]Takhtajan (1997)[30] as well as Thorne and Reveal (2007).[31]
Hutchinson (1973) derived a more elaborate hierarchy, placing order Liliales as one of 14 in division Corolliferae, one of three divisions of subphylum monocotyledons. Cronquist (1981) placed the order Liliales as one of two in subclass Liliidae, one of five in the class Liliopsida (monocotyledons) of division Magnoliophyta (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 within superorderLilianae, one of four within subclass Liliidae. Liliidae in turn was one of four subclasses in class Liliopsida (monocots).[32] In contrast Thorne and Reveal (2007) abandoned the use of monocotyledons as a distinct taxon, replacing it with 3 separate subclasses of Magnoliopsida (angiosperms), of which Liliidae consists of 3 superorders, placing Liliales in superorder Lilianae.[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 Angiosperm Phylogeny Group APG system (1998) established a structure of monocot classification with ten orders.[37] Notable was the separation of asparagids, as suggested by Dahlgren,[10] into Asparagales, with other taxa placed in Dioscoreales, resulting in a much reduced order.[9][8]
Phylogeny
The position of Liliales within the monocots (Lilianae) is shown in the following cladogram. The monocot orders form three grades, the alismatid monocots, lilioid monocots and the commelinid 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 the basal 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) is pectinate, that is diverging in succession from the line that leads to the commelinids.[41] The lilioid monocot orders constitute a paraphyletic assemblage, that is groups with a common ancestor that do not include all direct descendants (in this case commelinids which are a sister group to Asparagales); to form a clade, all the groups joined by thick lines would need to be included. In the cladogram the numbers indicate crown group (most recent common ancestor of the sampled species of the clade of interest) divergence times in mya (million years ago).[42]
Cladogram 1: The phylogenetic composition of the monocots[38][43]
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 in Asparagales, with the remainder in commelinids and Dioscoreales, as shown in this table.
Liliales families in progressive taxonomic schemes
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]
The exact phylogenetic relationship between the families of Liliales has been subject to revision. This cladogram shows that of the Angiosperm Phylogeny Website (2020):[47][11]
The bulk of the Liliales species are found in the very diverse 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]
The Corsiaceae (ghost-flower family) are a very small family of 3 mycoheterotrophic genera, lacking chlorophyll, 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 genus Corsia. The name commemorates the Florentine plant collector Marquis Bardo Corsi Salviati (1844–1907).[48][49]
The Campynemataceae (Green-mountainlily family) are a very small family of two genera and four species of rhizomatous herbaceous plants found in Tasmania and New Caledonia. The name is derived from the Greek words kampylos (curved) and nema (thread).[48][49]
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 and boreal 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 is Trillium (44 species) but many genera are monotypic. A number of genera, including Trillium are used as garden ornamentals, especially for woodland gardens. Paris japonica is noted for having the largest genome known to date. The family name is derived from the Greek words melas (black) and anthos (flower) in reference to the dark colour of the petals. [48][49]
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 the botanical garden at Leipzig.[48][49]
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 of Kuntheria 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 include Colchicum and Gloriosa. The family is named after Colchis on the eastern Black Sea.[48][49][9]
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 erect Alstroemeria (S America 125 spp.) and twining Bomarea (Central & S America 122 spp.) and two very small 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 of Bomarea edulis are directly consumed. Luzuriaga radicans, also from S America, produces fibre used in rope making. Alstroemeria cultivars are popular ornamentals and widely used as cut flowers (Peruvian lilies). The family is named for Baron Clas Alströmer (1736–1794), a student of Linnaeus.[48][49]
The Ripogonaceae (Supplejack family) is a very small 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) and gony (node) in reference to their node bearing shoots.[48][49]
The Philesiaceae (Chilean-bellflower family) are a very small family consisting of two monotypic genera, the two species being Philesia magellanica and the similar Lapageria rosea. They grow from a short woody rhizome, forming shrubs and vines respectively. They are found in the cool temperate forest of central and southern Chile, Magellan straits and adjacent Argentina, among the southern beech (Nothofagus) trees. Lapageria is the national flower of Chile and a popular ornamental with edible fruit. The name is thought to be related to the Greek word phileo (love), because of the attractiveness of its flowers.[48]
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 treat gout. S. aristolochiifolia was used to treat syphilis (but later as sarsaparilla to flavor root beer and confectionary). The fruit of S. megacarpa is consumed in conserves. The young shoots of many species are also edible. The family is named after the Greek myth of the affair between the mortal Krokos (or Crocus) and the nymph Smilax, whose punishment was to be turned into the prickly vine Smilax aspera.[48]
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, with Medeola 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 and Luzon, but are absent from the southern hemisphere. The bulbs have been used as foodstuffs or in traditional medicine. Cardiocrinum cordatum and Erythronium japonicum are sources of starch. Many Liliaceae are important in the floriculture and horticulture industries, particularly Tulipa and Lilium, but also Fritillaria. Many are also important ornamentals, such as Calochortus, Cardiocrinum, Clintonia, Erythronium and Tricyrtis. The name is derived from the Latin word for lily, lilium, which in turn is derived from the Greek leirion, a white lily.[48][49][40]
Distribution and habitat
Widely distributed but most commonly found in subtropical and temperate regions, especially herbaceous taxa in temperate regions of the Northern Hemisphere, and subtropical regions of the Southern hemisphere, including vines.[8] Since many species are cultivated they have been introduced in many regions and consequently worldwide, and a number have subsequently escaped and naturalised.[9]
↑ Coronariae used sensuAgardh (1825),[16] that is corresponding to Linnaeus' Liliaceae
Related Research Articles
Asparagales is an order of plants in modern classification systems such as the Angiosperm Phylogeny Group (APG) and the Angiosperm Phylogeny Web. The order takes its name from the type family Asparagaceae and is placed in the monocots amongst the lilioid monocots. The order has only recently been recognized in classification systems. It was first put forward by Huber in 1977 and later taken up in the Dahlgren system of 1985 and then the APG in 1998, 2003 and 2009. Before this, many of its families were assigned to the old order Liliales, a very large order containing almost all monocots with colorful tepals and lacking starch in their endosperm. DNA sequence analysis indicated that many of the taxa previously included in Liliales should actually be redistributed over three orders, Liliales, Asparagales, and Dioscoreales. The boundaries of the Asparagales and of its families have undergone a series of changes in recent years; future research may lead to further changes and ultimately greater stability. In the APG circumscription, Asparagales is the largest order of monocots with 14 families, 1,122 genera, and about 36,000 species.
The Alismatales (alismatids) are an order of flowering plants including about 4,500 species. Plants assigned to this order are mostly tropical or aquatic. Some grow in fresh water, some in marine habitats. Perhaps the most important food crop in the order is the corm of the taro plant, Colocasia esculenta.
Arecales is an order of flowering plants. The order has been widely recognised only for the past few decades; until then, the accepted name for the order including these plants was Principes.
The Dioscoreales are an order of monocotyledonous flowering plants, organized under modern classification systems, such as the Angiosperm Phylogeny Group or the Angiosperm Phylogeny Web. Among monocot plants, Dioscoreales are grouped with the lilioid monocots, wherein they are a sister group to the Pandanales. In total, the order Dioscoreales comprises three families, 22 genera and about 850 species.
LiliopsidaBatsch is a botanical name for the class containing the family Liliaceae. It is considered synonymous with the name monocotyledon. Publication of the name is credited to Scopoli : see author citation (botany). This name is formed by replacing the termination -aceae in the name Liliaceae by the termination -opsida.
Smilacaceae, the greenbriers, is a family of flowering plants. While they were often assigned to a more broadly defined family Liliaceae, most recent botanists have accepted the two as distinct families, diverging around 55 million years ago during the Early Paleogene. One characteristic that distinguishes Smilacaceae from most of the other members of the Liliaceae-like Liliales is that it has true vessels in its conducting tissue. Another is that the veins of the leaves, between major veins, are reticulate (net-shaped), rather than parallel as in most monocots.
Colchicaceae is a family of flowering plants that includes 15 genera with a total of about 285 known species according to Christenhusz and Byng in 2016.
A system of plant taxonomy, the Thorne system of plant classification was devised by the American botanist Robert F. Thorne (1920–2015) in 1968, and he continued to issue revisions over many years (1968–2007).
One of the modern systems of plant taxonomy, the Dahlgren system was published by monocot specialist Rolf Dahlgren in 1975 and revised in 1977, and 1980. However, he is best known for his two treatises on monocotyledons in 1982 and revised in 1985. His wife Gertrud Dahlgren continued the work after his death.
A system of plant taxonomy, the Takhtajan system of plant classification was published by Armen Takhtajan, in several versions from the 1950s onwards. It is usually compared to the Cronquist system. It admits paraphyletic groups.
In plant taxonomy, commelinids is a clade of flowering plants within the monocots, distinguished by having cell walls containing ferulic acid.
Liliidae is a botanical name at the rank of subclass. Circumscription of the subclass will vary with the taxonomic system being used ; the only requirement being that it includes the family Liliaceae.
Campynemataceae (Campynemaceae) is a family of flowering plants. The family consists of two genera and four species of perennial herbaceous plants endemic to New Caledonia and Tasmania.
Uvulariaceae is a family of flowering plants. While seldom recognised, the family is accepted by the Dahlgren system, which places it in order Liliales, superorder Lilianae, and the subclass Liliidae [=monocotyledons] of class Magnoliopsida [=angiosperms].
Lilianae is a botanical name for a superorder of flowering plants. Such a superorder of necessity includes the type family Liliaceae. Terminations at the rank of superorder are not standardized by the International Code of Nomenclature for algae, fungi, and plants (ICN), although the suffix -anae has been proposed.
Lilioid monocots is an informal name used for a grade of five monocot orders in which the majority of species have flowers with relatively large, coloured tepals. This characteristic is similar to that found in lilies ("lily-like"). Petaloid monocots refers to the flowers having tepals which all resemble petals (petaloid). The taxonomic terms Lilianae or Liliiflorae have also been applied to this assemblage at various times. From the early nineteenth century many of the species in this group of plants were put into a very broadly defined family, Liliaceae sensu lato or s.l.. These classification systems are still found in many books and other sources. Within the monocots the Liliaceae s.l. were distinguished from the Glumaceae.
The taxonomy of the plant family Liliaceae has had a complex history since its first description in the mid-eighteenth century. Originally, the Liliaceae were defined as having a "calix" (perianth) of six equal-coloured parts, six stamens, a single style, and a superior, three-chambered (trilocular) ovary turning into a capsule fruit at maturity. The taxonomic circumscription of the family Liliaceae progressively expanded until it became the largest plant family and also extremely diverse, being somewhat arbitrarily defined as all species of plants with six tepals and a superior ovary. It eventually came to encompass about 300 genera and 4,500 species, and was thus a "catch-all" and hence paraphyletic. Only since the more modern taxonomic systems developed by the Angiosperm Phylogeny Group (APG) and based on phylogenetic principles, has it been possible to identify the many separate taxonomic groupings within the original family and redistribute them, leaving a relatively small core as the modern family Liliaceae, with fifteen genera and 600 species.
MelanthialesLink was an order of monocotyledons, whose name and botanical authority is derived by typification from the description of the type family, Melanthiaceae by Johann Heinrich Friedrich Link in 1829.
Coronariae is a term used historically to refer to a group of flowering plants, generally including the lilies (Liliaceae), and later replaced by the order Liliales. First used in the 17th century by John Ray, it referred to flowers used to insert in garlands. Coronariae soon came to be associated with Liliaceae in the Linnaean system. The term was abandoned at the end of the 19th century, being replaced with Liliiflorae and then Liliales.
Stevenson, D. W.; Davis, J. I.; Freudenstein, J. V.; Hardy, C. R.; Simmons, M. P.; etal. (2000). "A phylogenetic analysis of the monocotyledons based on morphological and molecular character sets, with comments on the placement of Acorus and Hydatellaceae". In Wilson, K. L.; Morrison, D. A. (eds.). Monocots: Systematics and evolution. Collingwood, Australia: CSIRO Publ. pp.17–24.
Hutchinson, John (1973). The families of flowering plants, arranged according to a new system based on their probable phylogeny. 2 vols (3rded.). Oxford University Press.
Wettstein, Richard (1924). "Liliiflorae". Handbuch der Systematischen Botanik 2 vols (3rded.). p.862. Archived from the original on 18 February 2015. Retrieved 11 January 2020.
Davis, Jerrold I.; Mcneal, Joel R.; Barrett, Craig F.; Chase, Mark W.; Cohen, James I.; etal. (2013), "Contrasting patterns of support among plastid genes and genomes for major clades of the monocotyledons", in Wilkin, Paul; Mayo, Simon J (eds.), Early Events in Monocot Evolution, pp.315–349, doi:10.1017/CBO9781139002950.015, ISBN978-1-139-00295-0
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