Liliales

Liliales Temporal range: Early Cretaceous- Recent
Lilium martagon (Martagon lily)
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Angiosperms
Clade:	Monocots
Order:	Liliales Perleb (1826)
Type species
Lilium candidum L.
Families
Alstroemeriaceae Campynemataceae Colchicaceae Corsiaceae Liliaceae Melanthiaceae Petermanniaceae Philesiaceae Ripogonaceae Smilacaceae

Liliales (older name: Lilia) is an order of monocotyledonous flowering plants in the Angiosperm Phylogeny Group and Angiosperm Phylogeny Web system, within the lilioid monocots. This order of necessity includes the family Liliaceae. The APG III system (2009) places this order in the monocot clade. In APG III, the family Luzuriagaceae is combined with the family Alstroemeriaceae and the family Petermanniaceae is recognized. Both the order Lililiales and the family Liliaceae have had a widely disputed history, with the circumscription varying greatly from one taxonomist to another. Previous members of this order, which at one stage included most monocots with conspicuous tepals and lacking starch in the endosperm are now distributed over three orders, Liliales, Dioscoreales and Asparagales, using predominantly molecular phylogenetics. The newly delimited Liliales is monophyletic, with ten families. Well known plants from the order include Lilium (lily), tulip, the North American wildflower Trillium, and greenbrier.

Thus circumscribed, this order consists mostly of herbaceous plants, but lianas and shrubs also occur. They are mostly perennial plants, with food storage organs such as corms or rhizomes. The family Corsiaceae is notable for being heterotrophic.

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. The initial diversification to the current families took place between 82 and 48 million years ago. The order consists of 10 families, 67 genera and about 1,768 species.

Description

The Liliales are 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 woody shrubs, which have fleshy stems and underground storage or perennating organs. They are mainly geophytes with elliptical straplike leaves with parallel venation or ovate with palmate veins and reticulate minor venation (Smilacaceae). In Alstroemeria and Bomarea (Alstroemeriaceae) the leaves are resupinate (twisted). 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. These are usually large and pointed and may be variegated in Fritillaria (Liliaceae). Nectaries may be perigonal (on tepals) or septal (on ovaries). Ovaries may be inferior or superior. 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.

The Liliales are characterised by (synapomorphies) the presence of nectaries at the base of the tepals or stamen filaments, together with extrorse (outward opening) anthers. This distinguishes them from the septal nectaries and introrse anthers that are the features of most other monocots.

Phytochemistry

The stems contain fructans, the plants also contain Chelidonic acid, saponins, while some species contain Velamen. The epicuticular wax is of the Convallaria type, consisting of parallel orientated platelets.

Genome

The order includes taxa with some of the largest genomes among Angiosperms, particularly Melanthiaceae, Alstroemeriaceae and Liliaceae.

Taxonomy

With 11 families, about 67 genera and about 1,558 species, Liliales is a relatively small angiosperm order, but a large group within the monocotyledons.

History

Origins

The botanical authority for Liliales is given to Perleb (1826), who grouped eleven families (Asparageae, Pontederiaceae, Asphodeleae, Coronariae, Colchicaceae, Dioscoreaceae, Hypoxideae, Amaryllideae, Haemodoraceae, Burmanniaceae, Irideae) into an order he called Liliaceae. Historically this taxonomic grouping has included many diverse monocot taxa.

A number of later taxonomists, such as Endlicher (1836) substitituted the term Coronarieae for this higher order, including eight subordinate taxa. The term Liliales was introduced by Lindley (1853), referring to these higher orders as Alliances. Lindley included four families in this alliance. Although Bentham (1877) restored Coronariae as one of seven Series making up the monocotyledons, it was replaced by Liliiflorae and then Liliales in subsequent publications.

Phyletic systems

Subsequent authors, now adopting a phylogenetic (phyletic) or evolutionary approach over the natural method, did not follow Bentham's nomenclature. Eichler (1886) used Liliiflorae for the higher order including Liliaceae, as did Engler (1903). Lotsy (1911) restored Liliales for the higher rank, an approach that has been adopted by most major classification systems since.

Although Wettstein (1924) used Liliiflorae (in class Monocotyledones, subdivision Angiospermae), from Hutchinson (1973) onwards, including Cronquist (1981), Takhtajan (1997) and Thorne and Reveal (2007) Liliales has been used. This is also the nomenclature of the molecular phylogenetic based modern system of the Angiosperm Phylogeny Group (1998–2016).

The Cronquist system (1981) placed the order in subclass Liliidae in the class Liliopsida [= monocotyledons] of division Magnoliophyta [= angiosperms]. It used a much wider circumscription (many of the plants here are assigned to Asparagales and Dioscoreales by APG II):

• order Liliales

  family Agavaceae

  family Aloaceae

  family Cyanastraceae

  family Dioscoreaceae

  family Haemodoraceae

  family Hanguanaceae

  family Iridaceae

  family Liliaceae

  family Philydraceae

  family Pontederiaceae

  family Smilacaceae

  family Stemonaceae

  family Taccaceae

  family Velloziaceae

  family Xanthorrhoeaceae

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

• order Liliales

  family Alstroemeriaceae

  family Calochortaceae

  family Colchicaceae

  family Iridaceae

  family Liliaceae

  family Uvulariaceae

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

• order Liliales

  family Alstroemeriaceae

  family Campynemataceae

  family Colchicaceae

  family Iridaceae

  family Liliaceae

  family Melanthiaceae

  family Trilliaceae

Angiosperm Phylogeny Group

The APG system (1998) also placed the order in the clade monocots, but with a slightly different circumscription (missing the family Corsiaceae):

• order Liliales

  family Alstroemeriaceae

  family Campynemataceae

  family Colchicaceae

  family Liliaceae

  family Luzuriagaceae

  family Melanthiaceae

  family Philesiaceae

  family Ripogonaceae [sic]

  family Smilacaceae

The APG II system (2003) places this order in the clade monocots and uses this circumscription:

• order Liliales

  family Alstroemeriaceae

  family Campynemataceae

  family Colchicaceae

  family Corsiaceae

  family Liliaceae

  family Luzuriagaceae

  family Melanthiaceae

  family Philesiaceae

  family Ripogonaceae

  family Smilacaceae

APG III (2009) uses this circumscription:

• order Liliales

  family Alstroemeriaceae

  family Campynemataceae

  family Colchicaceae

  family Corsiaceae

  family Liliaceae

  family Melanthiaceae

  family Petermanniaceae

  family Philesiaceae

  family Ripogonaceae

  family Smilacaceae

Phylogeny

According to the APWeb, the families of Liliales are related as follows:

Liliales	Corsiaceae Campynemataceae branch with 50‑80% support Melanthiaceae branch with 50‑80% support Petermanniaceae Colchicaceae Luzuriagaceae Alstroemeriaceae Ripogonaceae Philesiaceae Smilacaceae Liliaceae

Distribution and habitat

Widely distributed but most commonly found in subtropical and temperate regions, especially the Northern Hemisphere. Since many species are cultivated they have been introduced in many regions and consequently worldwide, and a number have subsequently escaped and naturalised.

Uses

Liliales form important sources of food and pharmaceuticals as well as playing a significant role in horticulture and floriculture as ornamental plants. Pharmaceutical products include colchicine from Colchicum and Gloriosa (Colchicaceae) and veratrine and related compounds from Veratrum (Melanthiaceae) and Zigadenus (Melanthiaceae).

Bibliography

Books

• Cronquist, A (1981). An integrated system of classification of flowering plants. New York: Columbia University Press. ISBN 0-231-03880-1. {{cite book}}: Invalid |ref=harv (help)
• Hutchinson, John (1973). The families of flowering plants, arranged according to a new system based on their probable phylogeny. 2 vols (3rd ed.). Oxford University Press. {{cite book}}: Invalid |ref=harv (help)
• Judd, Walter S. (2002). Plant Systematics: A Phylogenetic Approach (2nd ed.). W.H. Freeman. ISBN 978-0-87893-403-4. {{cite book}}: Invalid |ref=harv (help)
• Kubitzki, Klaus; Huber, Herbert, eds. (1998). The families and genera of vascular plants. Vol. 3. Flowering plants. Monocotyledons: Lilianae (except Orchidaceae). Berlin, Germany: Springer-Verlag. ISBN 978-3-540-64060-8. {{cite book}}: Invalid |ref=harv (help); Unknown parameter |editorlink1= ignored (|editor-link1= suggested) (help); Unknown parameter |editorlink2= ignored (|editor-link2= suggested) (help)
• Simpson, Michael G. (2011). Plant Systematics. Academic Press. ISBN 978-0-08-051404-8. {{cite book}}: Invalid |ref=harv (help)
• Stevenson, D. W.; Davis, J. I.; Freudenstein, J. V.; Hardy, C. R.; Simmons, M. P.; Specht, C. D. (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
• Stuessy, Tod F. (2009). Plant taxonomy: the systematic evaluation of comparative data (2nd ed.). New York: Columbia University Press. ISBN 0-231-51864-1. {{cite book}}: Invalid |ref=harv (help)
• Takhtajan, Armen Leonovich (1997). Diversity and Classification of Flowering Plants. Columbia University Press. ISBN 978-0-231-10098-4. {{cite book}}: Invalid |ref=harv (help)
• Wilson, K. L.; Morrison, D. A., eds. (2000). Monocots: Systematics and evolution (Proceedings of the Second International Conference on the Comparative Biology of the Monocotyledons, Sydney, Australia 1998). Collingwood, Australia: CSIRO. ISBN 0-643-06437-0. Retrieved 14 January 2014. {{cite book}}: Invalid |ref=harv (help) Excerpts

Historical sources

• Agardh, Carl Adolph (1825). "XI. Liliiflorae". Classes Plantarum (in Latin). Lund: Literis Berlingianis. p. 8. {{cite book}}: Invalid |ref=harv (help)
• Bentham, G.; Hooker, J.D. (1883). "Coronariae". Genera plantarum ad exemplaria imprimis in herbariis kewensibus servata definita. vol. 3 (in Latin). London: L Reeve & Co. pp. ix–x. {{cite book}}: Invalid |ref=harv (help)
• Eichler, August W. (1886) [1876]. Syllabus der Vorlesungen über specielle und medicinisch-pharmaceutische Botanik (4th ed.). Berlin: Borntraeger. {{cite book}}: Invalid |ref=harv (help)
• Endlicher, Stephanus (1836). "Coronariae". Genera plantarum secundum ordines naturales disposita (in Latin). Vindobonae: Apud Fr.Beck. pp. 129–157. {{cite book}}: Invalid |ref=harv (help)
• Engler, Adolf (1903) [1892]. Syllabus der Pflanzenfamilien: eine Übersicht über das gesamte Pflanzensystem mit Berücksichtigung der Medicinal- und Nutzpflanzen nebst einer Übersicht über die Florenreiche und Florengebiete der Erde zum Gebrauch bei Vorlesungen und Studien über specielle und medicinisch-pharmaceutische Botanik (3rd ed.). Berlin: Gebrüder Borntraeger Verlag. p. 233. {{cite book}}: Invalid |ref=harv (help)
• Lindley, John (1853) [1846]. "Liliales". The Vegetable Kingdom: or, The structure, classification, and uses of plants, illustrated upon the natural system (3rd. ed.). London: Bradbury & Evans. pp. 195–206. {{cite book}}: Invalid |ref=harv (help)
• Lotsy, J. P. (1911). Vorträge über botanische stammesgeschichte, gehalten an der Reichsuniversität zu Leiden. Ein lehrbuch der pflanzensystematick. III Cormophyta Siphonogamia. Jena: G. Fischer. {{cite book}}: Invalid |ref=harv (help)
• Perleb, Karl Julius (1826). Lehrbuch der Naturgeschichte des Pflanzenreichs. Freiburg im Breisgau: Friedrich Wagner. p. 129. {{cite book}}: Invalid |ref=harv (help)
• Wettstein, Richard (1924). Handbuch der Systematischen Botanik 2 vols (3rd ed.). Retrieved 11 January 2020. {{cite book}}: Invalid |ref=harv (help)

Articles

• Angiosperm Phylogeny Group (2009), "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III", Botanical Journal of the Linnean Society, 161 (2): 105–121, doi:10.1111/j.1095-8339.2009.00996.x, archived from the original on 2017-05-25, retrieved 2010-12-10
• Balkrishna, Acharya; Joshi, Bhasker; Srivastava, Anupam; et al. (July–September 2019). "A comparison Of the Angiosperm Phylogeny Group Classification and other prominent classifications of angiosperms" (PDF). Indian Journal of Plant Sciences. 8 (3): 40–75.{{cite journal}}: CS1 maint: date format (link)
• Flaviana J. De Lima; Antônio A.F. Saraiva; Maria A.P. Da Silva; Renan A.M. Bantim; Juliana M. Sayão (2014), "A new angiosperm from the Crato Formation (Araripe Basin, Brazil) and comments on the Early Cretaceous Monocotyledons" (PDF), Anais da Academia Brasileira de Ciências
• Givnish, Thomas J.; Zuluaga, Alejandro; Marques, Isabel; Lam, Vivienne K. Y.; Gomez, Marybel Soto; Iles, William J. D.; Ames, Mercedes; Spalink, Daniel; Moeller, Jackson R.; Briggs, Barbara G.; Lyon, Stephanie P.; Stevenson, Dennis W.; Zomlefer, Wendy; Graham, Sean W. (December 2016). "Phylogenomics and historical biogeography of the monocot order Liliales: out of Australia and through Antarctica". Cladistics. 32 (6): 581–605. doi:10.1111/cla.12153.
• Kim, Jung Sung; Hong, Jeong-Ki; Chase, Mark W.; Fay, Michael F.; Kim, Joo-Hwan (May 2013). "Familial relationships of the monocot order Liliales based on a molecular phylogenetic analysis using four plastid loci: matK, rbcL, atpB and atpF-H". Botanical Journal of the Linnean Society. 172 (1): 5–21. doi:10.1111/boj.12039.
• Kim, Jung Sung; Kim, Joo-Hwan; Robinson-Rechavi, Marc (14 June 2013). "Comparative Genome Analysis and Phylogenetic Relationship of Order Liliales Insight from the Complete Plastid Genome Sequences of Two Lilies (Lilium longiflorum and Alstroemeria aurea)". PLoS ONE. 8 (6): e68180. doi:10.1371/journal.pone.0068180. PMC 3688979. PMID 23950788.{{cite journal}}: CS1 maint: unflagged free DOI (link)
• Leitch, I. J.; Soltis, D. E.; Soltis, P. S.; Bennett, M. D. (1 January 2005). "Evolution of DNA Amounts Across Land Plants (Embryophyta)". Annals of Botany. 95 (1): 207–217. doi:10.1093/aob/mci014. PMC 4246719. {{cite journal}}: Cite has empty unknown parameter: |1= (help)
• Rudall, P.; Stobart, K. L.; Hong, . -P.; Conran, J. G.; Furness, C. A.; Kite, G. C.; Chase, M. W., Consider the lilies: systematics of Liliales., pp. 347–359, in Wilson & Morrison (2000)
• Sen, Sumitra (1975). "Cytotaxonomy of Liliales". Feddes Repertorium. 86 (5): 255–305. doi:10.1002/fedr.19750860502.
• Soltis, Douglas E.; Soltis, Pamela S.; Bennett, Michael D.; Leitch, Ilia J. (November 2003). "Evolution of genome size in the angiosperms". American Journal of Botany. 90 (11): 1596–1603. doi:10.3732/ajb.90.11.1596. {{cite journal}}: Cite has empty unknown parameter: |1= (help)
• Thorne, Robert F.; Reveal, James L. (April 2007). "An Updated Classification of the Class Magnoliopsida ("Angiospermae")". The Botanical Review. 73 (2): 67–181. doi:10.1663/0006-8101(2007)73[67:AUCOTC]2.0.CO;2. {{cite journal}}: Invalid |ref=harv (help)
• Traub, Hamilton P; Kress, WJ (2016). "Liliales". Encyclopædia Britannica. {{cite encyclopedia}}: Invalid |ref=harv (help)
• Vinnersten, A.; Bremer, K. (2001). "Age and biogeography of major clades in Liliales". American Journal of Botany. 88 (9): 1695–1703. doi:10.2307/3558415. JSTOR 3558415. PMID 21669704. {{cite journal}}: Invalid |ref=harv (help) (Available online: [1])

Websites

• Stevens, P.F. (2016) [2001], Angiosperm Phylogeny Website, Missouri Botanical Garden, retrieved 1 February 2016 {{citation}}: Invalid |ref=harv (help)
• WFO (2019). "Liliales Perleb". World Flora Online. Retrieved 27 December 2019. {{cite web}}: Invalid |ref=harv (help)
• "Liliales Perleb". Tropicos. Missouri Botanical Garden. 2019. Retrieved 27 December 2019.