Liliales

Liliales is a plant taxon belonging to the order taxonomic category, used in modern classification systems such as the 2009 APG III classification system and the APWeb, and is necessarily circumscribed to the except for the Liliaceae family. Liliales is a widely recognized order, but not necessarily delimited as in the aforementioned classification systems. Liliales was traditionally a difficult order to circumscribe and divide into families, since its characters do not occur in patterns that clearly delimit groups, and previously many monocots with showy tepals and no starch in the endosperm were located within a very broad Liliales., which today are distributed in Liliales, Dioscoreales and Asparagales. The current circumscription is supported by molecular DNA analyzes which indicate that the order as defined here is monophyletic. It is located within the class Liliopsida (=monocotyledons). The order as currently circumscribed includes 10 families.

The representative members of this order are mainly herbaceous, although there are also lianas and shrubs with organs that store food, such as corms or rhizomes, and even herbs without chlorophyll and mycoheterotrophic (Corsiaceae).

Description

Theoretical Introduction in Descriptive Terminology of Plants

The group consists mainly of geophytes bearing elliptic leaves with fine reticulate venation. Synapomorphies underlying this group include nectaries primarily at the base of the tepals or filaments, septal nectaries that are almost always missing, extrorse anthers (opening outward from the flower), and frequent dotting on the florets. tepals, which are usually large. The outer epidermis of the seed coat has a cellular structure and does not have phytomelanins (a black scab), the inner part of the seed coat (tegmen) also has a cellular structure, both are plesiomorphic (see also Stevenson et al. 2000, Rudall et al. 2000a).

The group includes some species with the largest genomes among angiosperm plants (Soltis et al. 2003b, in some Melanthiaceae, Liliaceae and Alstroemeriaceae see Leitch et al 2005).

In addition, the plants of this order have fructans in the stem, chelidonic acid, present saponins, in some species there are velamen, in others there are cuticular waxes like parallel platelets, the base of the leaves is not sheathing, the inflorescence is terminal, there are many tenuinucellated ovules per carpel, they present a nucellar cap ("nucellar cap "), the long style, the capitate stigma, the deciduous perianth, the endosperm with cells with thick hemicellulosic walls, and without the presence of the mitochondrial gene sdh3.

Melanthiaceae, Liliaceae, Colchicaceae, as well as Corsiaceae (Rudall and Eastman 2002) have tepals with three foliar traces (that is, they have three vascular bundles connected to the central vascular system of the stem). Smilacaceae, on the other hand, have tepals with a single leaf pattern.

Ecology

The numerous species that make up this order are distributed throughout the world.

Many Liliales often have Paris type mycorrhizae (F. A. Smith and Smith 1997).

Diversity

The taxonomic diversity of monocots is presented in detail by Kubitzki (1998, 2006).

Here is the list of the diversity of Liliales. The descriptions are deliberately incomplete. For more information follow the links.

Flower of Corsia ornata. Note the stamens and the label. Corsiaceae Main article: Corsiaceae Corsiaceae, distributed in southern China, South America, and in Papua to Australia, are herbs without chlorophyll, mycoheterotrophic, with closed leaves and flowers of large bilateral symmetry, terminals, solitary. The flowers have the middle part of the outer verticyl of the larger perianto than the rest of the teapals, it is called labelo and its position is adaxial. Its 6 stamens are extruded and the ovary is inferous.

Habits of Veratrum lobelianum. Melanthiaceae Main article: Melanthiaceae Melanthiaceae, distributed in temperate regions of the Northern Hemisphere, especially in East Asia and East North America (with connections to Peru), consist of 16 genders from which 5 distinctive groups can be distinguished morphologically, among which are genders such as Trillium (formerly in Trilliaceae). They have alwaysgreen leaves, with a staggering base, the cluster inflorescence, 3 or 4 teapals, separate styles and the dry stigma, the persistent perianto in the fruit, the small embryo.

Flowers Colchicum autumnale. Colchicáceas Main article: Colchicaceae Colchicáceas, from temperate to tropical but absent regions in South America, are geófitas with cormos or rhizomes that can be recognized by their fairly long flowers with six teapals usually free, each more or less wrapping up a stamen, and with nectarios on the surface above or at the base of the teapals, and a super ovary. They are divided into 6 tribes.

Flowers Alstroemeria aurea. Alstroemeriaceae Main article: Alstroemeriaceae Main article: Luzuriageae The alstroemeriaceae, distributed in temperate and tropical zones of South America, Central America, New Zealand and Australia can be recognized by their spiral leaves that are rotated at the base so they are more or less "downside" and have fairly large sheets. The Alstroemerieae tribe has large flowers and bilateral symmetry and possess the mean part of the external verticil of the adaxial perianto (so the flowers are "bleep"), and its ovary is inferous. The Luzuriageae tribe is distributed from Peru to Tierra del Fuego, in the Falkland Islands and New Zealand and Australia, has only 2 genera and 5 species. They are climbers with more or less woody stems and can be recognized by their polysymmetric white flowers with smooth-colored and super ovary tepalos.

Habits of Lapageria rosea. Files Main article: Philesiaceae The files, distributed in the south of Chile, are quite woody or vinegar bushes, with peciolada leaves, and large and perndula flowers, the fruit is a berry.

Leaves and fruits Smilax aspera (brazaparrilla). Smilacáceas Main article: Smilacaceae The smilacáceas, distributed in pantropical regions to temperate, are often spiny climbers. The leaves are spiraled and pecioladas, have paired paired claws, and the foil has many longitudinal strong veins and refined venation. The flowers are quite small, typical of monocothylenes, in umbelled inflorescences, the fruit is a berry.

Flowers Lilium (blue or lilies). Liliaceae Main article: Liliaceae Liliaceae, as currently circumscribed, are distributed in temperate regions of the Northern Hemisphere, especially East Asia and North America. They are more or less bulbous geophytes that can be recognized by their large flowers with six free and often punctuated teapals, six extruded stamens, and a super ovary.

History of the constituency of order

Many of the families currently treated within Dioscoreales, Asparagales, and Liliales were formerly considered within a broader Liliales (Cronquist 1981, Thorne 1992), such as the petaloid monocots, a "group" characterized by flowers with showy tepals and no starch in the endosperm. Cronquist (1981) placed most of the petaloid monocots with 6-stamen flowers in a very large (and now known to be widely polyphyletic) Liliaceae. Others have divided the petaloid monocots with 6 stamens into Liliaceae, including species with a superior ovary, and Amaryllidaceae, including species with an inferior ovary (Lawrence 1951). This separation is also artificial, separating clearly related genera such as Agave and Yucca (Agavaceae) and Crinum (Amaryllidaceae) and Allium (Alliaceae), as discussed in the treatments of each family.

The concept of Liliales as a distinct order and separate from other lilioid taxa such as those placed in Asparagales and Dioscoreales, originated with Huber (1969, 1977), whose ideas were later adopted by Dahlgren et al. (1985). Almost all taxa placed in the order were considered to be closely related, perhaps even members of a single family, Liliaceae. Cronquist (1981) placed all these families together with others such as Velloziaceae and Pontederiaceae in a subclass Liliidae, but if they were not arborescent or did not have reticulate leaves then he placed them directly in Liliaceae. The reason why such broad concepts of the Liliaceae and Liliidae were formulated is that the character patterns did not indicate clearly delimited subgroups, and authors who segregated some genera into other families did not do so consistently. For example, Cronquist (1981) segregated Aloe and some of its relatives into "Aloaceae", largely because Aloe had some arborescent species and the other genera were strongly succulent (eg Haworthia and allies), but retained the highly related Bulbine in the Liliaceae because it was strictly herbaceous. However, Kniphofia, which is herbaceous and not succulent, was placed in Aloaceae because its flowers and inflorescences were "aloeoid". Dahlgren et al. (1985) left Bulbine and Kniphofia in the Asphodelaceae due to their nearly identical and strongly bimodal karyotypes, while placing the others in Aloaceae (and the two families in Asparagales due to their seeds with phytomelanins).

Both the order Liliales and the family Liliaceae are here closely delimited, following Dahlgren et al. (1985) and recent cladistic analyses. Huber (1977) and Dahlgren et al. (1985) included Iridaceae and Orchidaceae in the order, while in molecular DNA analyzes these two families appear as part of Asparagales. They also excluded Melanthiaceae and Campynemataceae (as Melanthiales), which consistently appear in DNA analysis as members of Liliales.

DNA analysis on the rbcL sequence by Chase et al. (1993) and Duvall et al. (1993) identified a Liliales clade containing Alstroemeriaceae, Colchicaceae, Liliaceae, Melanthiaceae, and Smilacaceae. Chase et al. (1995a) added Luzuriagaceae, Philesiaceae, Smilacaceae, and Rhipogonaceae, and also placed in order genera that had previously been placed in the families Calochortaceae (now Liliaceae), Trilliaceae (today in Melanthiaceae) and Uvulariaceae (today in Colchicaceae or Liliaceae).

In cladistic analyzes of morphology (not DNA) by Chase et al. (1995b) and Stevenson and Loconte (1995), Alstroemeriaceae, Colchicaceae, Liliaceae, and Melanthiaceae formed a clade, but the other families were located further away from this clade. Iridaceae (today in Asparagales) was also included in Liliales in the two morphological studies.

Knowledge of relationships within Liliales has greatly improved, but delineation of some families is still problematic.

Phylogeny

Theoretical Introduction in Philogenia

Liliales (sensu APG II 2003) now seems to occupy the position of sister clade to (Asparagales + Commelinidae).

The monophyly of Liliales is supported by cladistic analyzes based on morphology and different DNA sequences (Chase et al. 1995a, b, 2000, Davis et al. 2004, Fay et al. 2006, Goldblatt 1995, Graham et al. 2006, Hilu et al. 2003, Källersjö et al. 1998, Soltis et al. 2000, Stevenson and Loconte 1995, Vinnersten and Bremer 2001, see also Chen et al. 2007 for a Bayesian analysis, but support from many branches is weak).

Putative phylogenetic relationships within the order are as follows:

Colchicaceae, Melanthiaceae and Smilacaceae were traditionally placed in Liliaceae.

The full cladogram is given below (APWeb, updated as of July 2008, based primarily on analysis by Fay et al. 2006, relationships suggested by 's study rbcL from Janssen and Bremer 2004 are quite different, but did not include Petermanniaceae or Corsiaceae):

Alstroemeriaceae (3 genera, 165 species, tropical America) is sister to Luzuriagaceae (2 genera, 5 species), native to South America (Luzuriaga ) and Australia and New Zealand (Drymophila). The two families share vegetative characters such as being vines with resupinate leaves in such a way that the upper surface during development becomes lower during maturity, although the ovary is superior in Alstroemeriaceae. The two may need to be combined into a single family (but the APWeb still, as of July 2008, keeps them separate).

Sister to (Alstroemeriaceae + Luzuriagaceae) is Colchicaceae, which is native mainly to the Old World, the only exception being Uvularia (from temperate regions of the Northern Hemisphere). Some genera of Colchicaceae have twisted leaves (as do Alstroemeriaceae and Luzuriagaceae). Colchicine (an alkaloid used to inhibit spindle formation and cause chromosome nondisjunction during mitosis, generating polyploid offspring) is found in all members of the family.

Petermania was listed in Colchicaceae in APG (1998) and APG II (2003), but it is now known that the DNA used to do the analysis actually came from a Tripladenia cunninghamii that had been misidentified as Petermania (M. W. Chase, unpublished data, cited in Soltis et al. 2005: p.104). The true Petermania turned out to be sister to the 3 families already mentioned (Colchicaceae, Alstroemeriaceae and Luzuriagaceae), so it may be appropriate to reinstate the family Petermanniaceae (as suggested by Soltis et al. 2005 and in fact it is done by APWeb).

Melanthiaceae (16 genera, 170 species) was studied in detail by Zomlefer et al. (2001), the family now includes members of what was formerly known as Trilliaceae. All genera have a distribution in temperate regions of the Northern Hemisphere, except for a single genus, Schoenocaulon, which is present in Peru, but may have been brought there by humans (they are medicinal plants). The family contains powerful alkaloids. Xerophyllum (two species in North America) is sister to the earlier Trilliaceae genera (Chase et al. 1995a, 1995b, Rudall et al. 2000a). These species are adapted to xerophytic environments (they have thin, graminiform leaves, and the habit is in a tight rosette), which contrasts with the habit of Trillium and allies, which are adapted to living below ground. forest canopy and have reticulate leaves. Some authors (Thorne 1992) considered Veratrum (Melanthiaceae) one of the more primitive monocots due to its plicate reticulate leaves, and mostly unfused carpels, but it is deeply embedded within Liliales and not it is not related to the base of the monocots at all, which is to say that it is unlikely that it retained these characters as primarily primitive.

Liliaceae sensu APG is composed of far fewer genera than in most previous constituencies (eg Cronquist 1981). However, as circumscribed by the APG it is broader than that of others who would have limited the family to only the core genera related to Lilium (Tamura 1998), which excluded Calochortus, Prosartes, Tricyrtis, and several others, placing them in Calochortaceae or Tricyrtidaceae. Liliaceae is exclusive to temperate regions of the Northern Hemisphere and is composed of geophytes with often large and dotted flowers, extrorse dehiscence anthers, and a superior ovary.

Related to the Liliaceae are Smilacaceae (monogeneric, 315 species), nearly cosmopolitan, Philesiaceae (2 monospecific genera), from southern South America, and Rhipogonaceae (monogeneric, 6 species), from Australasia. There are no known characters outside of DNA that unite all these families. Smilacaceae, Philesiaceae, and Rhipogonaceae have unique spiny pollen (Rudall et al. 2000a), but never formed a clade in molecular analyses. Rhipogonum is often listed as a sister to Philesia /Lapageria, so it could be combined with them, and Smilax is generally a sister to Liliaceae (but never more than 80% support). Fay et al. (2006: low support), Givnish et al. (2006: high support), and Chase et al. (2006) found Philesiaceae and Rhipogonaceae as sister taxa, and Smilacaceae as sister to Liliaceae.

Neyland (2002), analyzing variation in 26S rDNA, suggested that Corsiaceae (3 genera, 30 species, China, South America, and Australasia) were associated with the Liliales. While this position has only weak molecular support, it is largely consistent with the molecular evidence; Davis et al. (2004) also found Corsiaceae associated with this order.

Melanthiaceae, Campynemataceae (2 genera, 4 species, Australasia), and Corsiaceae have an unclear pattern of relationships with other members of the Liliales. Campynemataceae was previously considered to be related to Melanthiaceae (Dahlgren et al. 1985) due to their largely unfused carpels, while Corsiaceae was considered to be related to Burmanniaceae because of their shared mycoparasitic history. However, the two sets of characters are unreliable: the free carpels are potentially a simplesiomorphy, while the syndrome of characters associated with mycoheterotrophy is convergent even between eudicots and monocots with the same life history. Campynemataceae and Corsiaceae generally fit the pattern of characters seen among Liliales families.

Taxonomy

Theoretical Introduction in Taxonomy

The order was recognized by APG III (2009), Linear APG III (2009) assigned family numbers 52 to 61. The order had already been recognized by APG II (2003).

Liliales includes 10 families and about 1300 species, the largest families being Alstroemeriaceae, Liliaceae, Colchicaceae, Smilacaceae, and Melanthiaceae.

The complete list of families, according to the Angiosperm Phylogeny Website (as of January 2011) that coincides with that of the APG III, the family numbers assigned according to the LAPG III (2009):

• Campynemataceae (family No. 52)
• Melanthiaceae (family n. 53)
• Petermanniaceae (family No. 54)
• Alstroemeriaceae (family No. 55)
• Colchicaceae (family No. 56)
• Philesiaceae (family no. 57)
• Rhipogonaceae (family no. 58)
• Smilacaceae (family n.o 59)
• Corsiaceae (family no. 60)
• Liliaceae (family n. 61)

The APG II classification system (2003) used this circumscription:

• Alstroemeriaceae
• Campynemataceae
• Colchicaceae
• Corsiaceae
• Liliaceae
• Luzuriagaceae
• Melanthiaceae
• Philesiaceae
• Rhipogonaceae
• Smilacaceae

Petermania was included in Colchicaceae, today it is known that it was due to an error, for this reason the APWeb and the APG III give it a separate family. Furthermore, the APG III nests Luzuriagaceae within Alstroemeriaceae.

In the APG (1998) classification system, the constituency was as follows:

• Alstroemeriaceae
• Campynemataceae
• Colchicaceae
• Liliaceae
• Luzuriagaceae
• Melanthiaceae
• Philesiaceae
• Ripogonaceae [sic]
• Smilacaceae

The family Corsiaceae was missing, later determined to be a basal lilial.

Cronquist's system (1981) placed the order in the subclass Liliidae in the class Liliopsida [= monocots] of the division Magnoliophyta [= angiosperms]. The circumscription was much broader (many of these plants are now located in Asparagales and Dioscoreales):

• Agavaceae
• Aloaceae
• Cyanastraceae
• Godcoreaceae
• Haemodoraceae
• Hanguanaceae
• Iridaceae
• Liliaceae
• Philydraceae
• Pontederiaceae
• Smilacaceae
• Stemonaceae
• Taccaceae
• Velloziaceae
• Xanthorrhoeaceae

The Thorne (1992) system placed the order in the superorder Lilianae of the subclass Liliidae [= monocots ] of the class Magnoliopsida [= angiosperms] and used this circumscription:

• Alstroemeriaceae
• Campynemataceae
• Colchicaceae
• Iridaceae
• Liliaceae
• Melanthiaceae
• Trilliaceae

The Dahlgren (1985) system placed the order in the superorder Lilianae in the subclass Liliidae [= monocots] of the class Magnoliopsida [= angiosperms] and used this circumscription:

• Alstroemeriaceae
• Calochortaceae
• Colchicaceae
• Iridaceae
• Liliaceae
• Uvulariaceae

In Engler's classification system (last updated in 1964) a similar order was called Liliiflorae, located in the Monocotyledoneae case of the Angiospermae subdivision. Today it is prohibited by the International Code of Botanical Nomenclature to use that name for an order (because its suffix does not comply with the rules), but it could be used for a superorder.

The Wettstein system, with its last revision in 1935, used names similar to those of the Engler system: the order was called Liliiflorae and placed in the class Monocotyledones of the subdivision Angiospermae. In this constituency the order was similar to that of Cronquist.

Another previously used name for the order was Coronarieae in the Bentham and Hooker system.

Synonyms: Alstroemeriales Hutchinson, Campynematales Doweld, Colchicales Dumortier, Liriales K. Koch, Melanthiales Reveal, Paridales Dumortier, Smilacales Lindley, Veratrales Dumortier - Lilianae Takhtajan, Melanthianae Doweld - Liliidae J. H. Schnaffner - Liliopsida Batsch, Liriopsida Brongniart.

Evolution

The Liliales stem group is dated at about 124 million years to present, the Liliales crown group at about 117 million years to present (Janssen and Bremer 2004), these estimates are quite different from Bremer's (2000) who considered them younger.