Taxonomy of Liliaceae: Difference between revisions

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History

Pre-Darwinian

The Liliaceae family was first described by Michel Adanson in 1763,^[1]^[2] but formally named by Antoine Laurent de Jussieu in 1789.^[3] Adanson described eight subfamilies with seventy eight genera. However the subfamily he described as Lis (Lilies) (p. 48) had seven (Uvularia, Mithridatium, Mendoni, Lilium, Imperialis (Fritillaria), Fritillaria and Tulipa) of which four are in the modern genus. De Jussieu placed them as the fourth Ordo (Order: Lilia) of the third class (Stamina epigyna) of Monocts, with eight genera (Tulipa, Erythronioum, Methonica, Uvularia, Fritillaria, Imperialis, Lilium, Yucca) only four of which remain in the family. However the use of the term Ordo at that time was closer to what we now understand as Family, rather than Order.^[4]^[5] Although Jussieu used the term 'Lilia' in his Genera Plantarum, elsewhere he used the French term 'Liliacées',^[6] as had Adanson before him using both French and Latin, and the word 'Liliaceae' was soon widely used by botanists such as Samuel Frederick Gray,^[7] John Lindley,^[8] and Pierre-Joseph Redouté^[9] in the early nineteenth century. Jussieu defined this grouping as having a calyx of six equal coloured parts, six stamens, a superior ovary, single style, and trilocular capsule. Gray (1821) provided the first description of Jussieu's scheme in English, identifying two genera occurring in Britain (Tulipa, Fritillaria), distinguished by the absence or presence of basal nectaries. His key used the presence of six stamens, a single style, equal stamina, a simple petaloid perianth and a trilocular capsule with flat seeds to identify the family.^[10] Although De Candolle (1813) had not explicitly described the Liliaceae, his overall classification scheme influenced many later writers including Gray.^[11] In this scheme,^[12] the Liliaceae were considered a family within the Endogenæ phanerogamæ, that is those vascular plants (Vasculares) whose vascular bundles were thought to arise from within (Endogènes, endogenous), a term he preferred to monocots (Monocotylédonés). Of these, the true monocots of Jussieu became the Phanerogams or Phanérogames (Phenogamae in Gray), hence Endogenæ phanerogamæ.^[13] Candolle also instituted the concept of ordered ranks, based on classes, subclasses, familles (Latin: ordines naturales) and tribus (tribes),^[5] subdividing the Liliaceae.

Lindley was the first English systematist, publishing his work in 1830, and following the reasoning of Jussieu. he used the term tribe to describe the Liliaceae as a division of the hexapetaloid monocots, characterised by a superior ovary, highly developed perianth, inward turning anthers, a trilocular polyspermous capsule and seeds with a soft spongy coat. He offered seven genera as examples (Erythronium, Lilium, Calochortus, Blandfordia, Polianthes, Hemerocallis and Funkia). By 1845, in his final work, he refined and greatly expanded his taxonomy further favouring the term Alliances of Endogens over monocots, as a class, with eleven alliances including the Liliales. This includes four Orders (families) including Liliaceae which he referred to as lilyworts in the vernacular. In this work he unhappily acknowledged the great diversity in the circumscription of the order, and that it had expanded vastly, with many subdivisions. As he saw it, the Liliaceae were already paraphyletic ("catch-all"), being all Liliales not included in the other orders, but hoped that the future would reveal some characteristic that would group them better. He recognised 133 genera and 1200 species.^[14] By the time of the next major British classification, that of Bentham and Hooker in 1883 (published in Latin) several of Lindley's other families had been absorbed into the Liliaceae.^[15]

Post-Darwinian

Although the appearance of Charles Darwin's Origin of Species in 1859 changed the way that taxonomists considered plant classification, incorporating evolutionary information into their schemata, this did little to further define the circumscription of Liliaceae.^[16] For instance, the first phylogenetic based system, by Bessey (1915) followed Engler in defining Liliaceae as "Pistil mostly 3-celled; stamens 6; perianth of two similar whorls, each of three similar leaves", although placing the Liliales in a novel sublass of monocots, the Strobiloideae.^[17] The Liliaceae were treated as Liliiflorae by Wettstein (1901–1935) and by Engler and Prantl (1931), and Liliales from the systems of Hutchinson (1959) onwards. Over time the Liliaceae became increasingly broadly, 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 the order Liliales in the scheme of Arthur Cronquist (1981) who merged it with the Amaryllidaceae.^[18] Later similar schemes include the system of Robert F. Thorne^[19] and that of Armen Takhtajan.^[20] They were characterised as petaloid monocots, characterized by showy flowers with tepals and without starch in the endosperm. Cronquist had placed most flowering petaloid monocots with six stamens in this very broad (and clearly polyphyletic) family.^[21] Cronquist rejected the importance of the position of the ovary and thus included the Amarylloidaceae some species of which had an inferior ovary, and which others separated into a distinct family,^[22] although this then created other problems.^[23] The Liliaceae were one of the major families in the Cronquist system which considerably broadened the circumscription to include 22 families in addition to Liliaceae s.s..^[24]

Dismembering Liliaceae

Other botanists in the twentieth century echoed Lindley's concerns about the phylogeny of the Liliaceae. The earliest of these was Johannes Paulus Lotsy in 1911,^[25] followed by Huber in 1969 who suggested two separate families.^[26] These various proposals to separate small groups of genera into more homogeneous families, made little impact till that of Dahlgren (1985) incorporating new information including synapomorphy.^[21] Where Cronquist saw one family, Dahlgren saw forty distributed over three orders (predominantly Liliales and Asparagales).^[27]^[28] Over the 1980s, in the context of a more general review of the classification of angiosperms, the Liliaceae were subjected to more intense scrutiny. By the end of that decade, the Royal Botanic Gardens at Kew, the British Museum of Natural History and the Edinburgh Botanical Gardens formed a committee to examine the possibility of separating the family at least for the organization of their herbaria. That committee finally recommended that 24 new families be created in the place of the original broad Liliaceae, largely by elevating subfamilies to the rank of separate families.^[29]

The 1990s saw considerable progress in plant phylogeny and phylogenetic theory, enabling a phylogenetic tree to be constructed for all of the flowering plants. The establishment of major new clades necessitated a departure from the older but widely used classifications such as Cronquist and Thorne based largely on morphology rather than genetic data. This complicated discussion about plant evolution and necessitated a major restructuring.^[30] rbcL gene sequencing and cladistic analysis of monocots had redefined the Liliales in 1995^[31] from four morphological orders sensu Dahlgren. The largest clade representing the Liliaceae, all previously included in Liliales, but including both the Calochortaceae and Liliaceae sensu Tamura. This redefined family, that became referred to as core Liliales, but corresponded to the emerging circumscription of the Angiosperm Phylogeny Group (1998).^[32]

Modern APG classification

To meet this need, flowering plant systematists formed the Angiosperm Phylogeny Group (APG) resulting in a new classification published in 1998^[33] largely based on the work of Kåre Bremer and colleagues at Uppsala and Stockholm universities in the late 1970s, ^[34]^[35]^[36] which became available on the internet in 1996.^[37] This scheme was ordinal (concentrating on orders rather than families), prioritising monophyly. However progress was rapid and the modern era of the taxonomy of the Liliaceae family comes from Judd et al.^[23] (second edition 2002), the APG II (2003^[30]) and APG III (2009 ^[38]), while the Linear APG III assigned it the family number 61. ^[39] While the original APG did not specifically address the issues of the polyphyly within Liliaceae, APG II did so within the two closely morphologically related orders, Liliales and Asparagales^[40] recognising the continued common use of the sensu lato (s.l.).

These studies of DNA and morphological data (particularly reproductive morphology) together with phylogenetic analyses, allowed the conclusion that the "petaloid monocots" do not really belong to one botanical family but rather are distributed across two separate different orders, the Asparagales and Liliales. The monophyly of these newly defined orders is supported by cladistic analysis based on morphology, 18S rDNA, and many other DNA sequences, particularly the plastid gene rbcL.^[41]^[42]^[43]^[44]^[45]^[46]^[47]^[48]^[49]^[50]^[51]^[52]^[53]^[54]^[55]^[56]^[57]^[58]^[59]

Together with other analyses within each of these two orders has allowed the redistribution of the original genera of Liliaceae sensu lato into a variety of families, as illustrated in the two cladograms of the Asparagales and Liliales shown here.^[60] These changes have resulted in considerable changes in the positioning of Liliaceae within the overall classification, as shown in the table below, as well as within the family.

Table 1: Evolution of placement of Liliaceae in different taxonomic schemes^[16]
Rank	Bentham and Hooker (1883)^[15]	Cronquist (1981)^[18]	Takhtajan (1980)^[61]	Dahlgren (1985)^[21]	Thorne (1992)^[19]	APG (2003-9)^[30]^[38]
Division		Magnoliophyta	Magnoliophyta
Class	Monocotyledons	Liliopsida	Liliopsida	Liliopsida	Angiospermae
Subclass		Liliidae	Liliidae	Liliidae	Liliidae
Superorder (Series)	Coronarieæ		Dioscoreanae	Liliiflorae	Lilianae	Monocots
Order		Liliales	Liliales	Liliales	Liliales	Liliales

For a comparison of the classifications of Genera from 1959 (Hutchinson)^[62] to 2000 (Wilson and Morrison)^[63] see Table 1 in Fay et al. 2006.^[64]

Cladistic analysis of Liliales and families within former Liliaceae

Within the Liliales, cladistic analysis revealed the Liliaceae as one of four main groupings.^[40] The following cladograms based on molecular phylogenetics reveal the polyphyletic nature of the original Liliaceae family, which encompassed a large number of differing groups of genera. This led to botanists increasingly adopting a stricter more narrow monophyletic concept (sensu stricto) of the family based on phylogenetic relationships, as expressed in the 2009 APG III system, rather than the older broader one (sensu lato),

Asparagales

Orchidaceae

Hypoxidaceae (traditionally placed in Amaryllidaceae)

Iridaceae

Xanthorrhoeaceae sensu lato

	Hemerocallidaceae (traditionally placed in Liliaceae)

	Xanthorrhoeaceae (traditionally placed in Liliaceae)

	Asphodelaceae (traditionally placed in Liliaceae)

Alliaceae sensu lato

	Agapanthaceae (traditionally placed in Liliaceae)

	Alliaceae (traditionally placed in Liliaceae)

	Amaryllidaceae

Laxmanniaceae (traditionally placed in Liliaceae)

	Asparagaceae (traditionally placed in Liliaceae)

	Ruscaceae (traditionally placed in Liliaceae)

	Themidaceae (traditionally placed in Liliaceae)

	Hyacinthaceae (traditionally placed in Liliaceae)

Agavaceae (traditionally placed in Amaryllidaceae or Liliaceae)

Liliales

	Alstroemeriaceae (traditionally placed in Liliaceae)

	Colchicaceae (traditionally placed in Liliaceae)

Melanthiaceae (traditionally placed in Liliaceae)

	Smilacaceae (traditionally placed in Liliaceae)

	Liliaceae sensu stricto

Other displaced groups included members of the Nartheciaceae, Tecophilaeaceae, Tofieldiaceae, and Uvulariaceae families and Dioscoreales, and Alismatales orders.

Changes to family structure in APG III

The genus list above is based on the APG III system. When it was published in 2009, the families Xanthorrhoeaceae, Amaryllidaceae, and Asparagaceae, were greatly expanded.^[65] Thirteen of the families of the earlier APG II system were thereby reduced to subfamilies. The APG II families and their equivalent APG III subfamilies are as follows:

Asphodelaceae=Asphodeloideae, Xanthorrhoeaceae
Xanthorrhoeaceae=Xanthorrhoeoideae, Xanthorrhoeaceae
Hemerocallidaceae=Hemerocallidoideae, Xanthorrhoeaceae
Agapanthaceae=Agapanthoideae, Amaryllidaceae
Alliaceae =Allioideae, Amaryllidaceae
Amaryllidaceae=Amaryllidoideae, Amaryllidaceae

Aphyllanthaceae = Aphyllanthoideae, Asparagaceae
Laxmanniaceae = Lomandroideae, Asparagaceae
Asparagaceae = Asparagoideae, Asparagaceae
Ruscaceae = Nolinoideae, Asparagaceae
Agavaceae = Agavoideae, Asparagaceae
Themidaceae = Brodiaeoideae, Asparagaceae
Hyacinthaceae = Scilloideae, Asparagaceae

Phylogeny

Sequencing of the rbcL and matK regions of the chloroplast DNA of Lilium and related genera^[66] confirmed the circumscription of the family s.s. as described by Tamura (1998).^[67] Chloroplast ndhF gene sequencing also supports Liliaceae monophyly, reuniting the Liliaceae and Calochortaceae sensu Tamura.^[32]^[64]

Despite establishing this relative degree of monophyly for the Liliaceae family,^[31]^[68] morphology remains diverse^[67] and there exists within the Liliaceae clade, a number of subclades. Particularly enigmatic were Clintonia, Medeola, Scoliopus, and Tricyrtis. Clintonia (which has a disjunct distribution involviong both East Asia and North America) and the closely related Medeola form one of those subclades, and are now considered a separate tribe (Medeoleae), within the Lilioideae although at different times they have been both a separate subfamily (Medeoloideae) and family (Medeolaceae). Sequencing of the rbcL and matK chloroplast genes established monophyly for Clintonia, but with separate clades corresponding to the two distribution regions.^[69]

While the ten genera of the Lilioideae subfamily are characterised by contractile bulbs and roots, and a Fritillaria-type embryo-sac (megagametophyte with four megaspores), the five genera constituting the Streptopoideae and Calochortoideae subfamilies form another distinct group, previously characterised under the Calochortoideae alone. They are characterised by creeping rhizomes, styles which are divided at their apices, and by megagametophyte development of the Polygonum-type (a simple megaspore and triploid endosperm) embryo-sac. At times these genera were considered in a separate family (Calochortaceae) (e.g. Tamura) or even the more heterogeneous Uvulariaceae sensu Dahlgren. However most of the latter had low morphological similarity to the Liliaceae and Uvularia and Disporum are now located in the Colchicaceae. Disporum contained both Asian and North American species which had always been distinguishable. Following molecular analysis the North American species were restored to the genus Prosartes and retained in Liliaceae, and are now included in the Streptopoideae. The Asian species were moved to Colchicaceae.^[70]

Evolution

The development of a phylogenetic approach to taxonomy, starting with Charles Bessey's with The phylogenetic taxonomy of flowering plants (1915) suggested the Liliales formed some of the earliest monocots.^[17] Molecular analysis suggests that Liliaceae arose in Eurasia, around 68 million years ago during the late (Maastrichtian) Cretaceous to early (Paleocene) Paleogene eras^[71] with the development of two main evolutionary clades. The first of these, characterised as Lilileae (Lilium, Fritillaris, Nomocharis), Cardiocrinum), Notholirion) from the Himalayas, and the second, the Tulipae (Erythronium, Tulipa), (Gagea)) from East Asia. On the other hand Clintonia-Medeola may have appeared in North America but was subsequently dispersed, as may have the Calochortaceae sensu Tamura. Divergence amongst the Liliales probably occurred around 36 million years ago, with the Liliaceae (sensu Tamura) emerging at 27 million years, Liliaceae s.s. at 20 million, Lilieae 12 million and Calochortus 7 million.^[32] Liliaceae fossils have been dated to the Paleogene^[72] and Cretaceous^[73] eras in the Antarctic.

Core Liliales probably arose as shade plants, with subsequent evolution of Liliaceae s.s. and Calochortus to open areas including deciduous forest in the more open autumnal period, but then a return of some species (e.g. Cardiocrinum). This was accompanied by a shift from rhizomes to bulbs, 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) ^[32]

Subdivisions and genera

Suprageneric subdivisions

Due to the diversity of the originally broadly defined Liliaceae s.l., many attempts have been made to form suprageneric classifications, e.g. subfamilies and tribes.^[74] By 1813, Candolle recognised five subdivisions which he called tribes (Asparagées, Trilliacées, Asphodelées, Bromeliées, Tulipacées),^[12] all of which Jussieu had made separate families, with the exception of Tulipa, which was a genus within the Liliaceae. By 1845 John Lindley observed that the family had become extremely diverse, ill defined and unstable not only by its overall circumscription, but also by its subdivisions. Of the 133 genera he included, he described eleven suborders.^[14]

In 1879 a revision of the North American Liliaceae described sixteen tribes^[75] and Bentham and Hooker twenty tribes in 1883, while Engler and Prantl in their extensive description of the Liliaceae in 1899 identified 31 tribes distributed over 11 subfamilies.^[76] In 1936 Buxbaum undertook a major revision of the Liliaceae and inter alia described the subfamily Lilioideae, with three tribes: Lloydieae, Tulipeae and Lilieae.^[77]^[78]^[79] Peruzzi et al. (2009) discuss the complex rearrangement of the various genera, tribes and subfamilies over a 30 year period (Peruzzzi et al. Table1) from 1985,^[74] partly summarised here in Table 2.

Classifications published since the use of molecular methods in phylogenetics have taken a narrower view of the Liliaceae. In 1998 Tamura considered Calochortus sufficiently distinct to elevate the subfamily Calochortoideae to family status as Calochortaceae.^[67]^[80] In 2009, Takhtajan used a very narrow definition (see the table below).^[61] As of January 2014^[update] the Angiosperm Phylogeny Website (APweb)^[81] included four of Takhtajan's families in its Liliaceae, recognizing three subfamilies, one of which is divided into two tribes.^[82]

Table 2: Comparison of Four 21st Century Classifications of the Liliaceae
Tamura^[67]			Takhtajan^[61]		Taxonomicon^[83]			APweb^[82]
Family	Subfamily	Tribe	Family	Tribe	Family	Subfamily	Tribe	Family	Subfamily	Tribe
Liliaceae	Lilioideae	Lilieae	Liliaceae	Lloydieae	Liliaceae	Lilioideae	Lloydieae	Liliaceae	Lilioideae	Lilieae
		Lilieae		Lilieae			Lilieae
		Tulipeae		Tulipeae			Tulipeae
	Medeoloideae		Medeolaceae				Medeola, Clintonia			Medeoleae
Calochortaceae		Calochorteae	Scoliopaceae			Calochortoideae			Calochortoideae
Calochortaceae		Tricyrtideae	Tricyrtidaceae			Streptopoideae			Streptopoideae

Genera

History

The Cronquist system (1981) had nearly 300 genera in Liliaceae. Most of those have been reassigned to other families, as shown here, following ITIS, together with their disposition as APG III transfers to other families (including subfamilies and orders other than Liliales where applicable) in brackets (Family: Subfamily, Order). Current members of Liliaceae in bold:

Template:Multicol

Abama Adans. = Narthecium
Agapanthus L'Her. (Amaryllidaceae: Agapanthoideae, Asparagales)
Aletris L. (Nartheciaceae, Dioscoreales)
Allium L. Onion and garlic (Amaryllidaceae: Allioideae, Asparagales)
Alstroemeria L. (Alstroemeriaceae)
Amaryllis L. Amaryllis (Amaryllidaceae: Amaryllidoideae, Asparagales)
Amianthium Gray (Melanthiaceae)
Androstephium Torr. (Asparagaceae: Brodiaeoideae, Asparagales)
Anthericum L. (Asparagaceae: Agavoideae, Asparagales)
Asparagus L. (Asparagaceae: Asparagoideae, Asparagales)
Asphodelus L. (Xanthorrhoeaceae: Asphodeloideae, Asparagales)
Astelia Banks & Soland. ex R. Br. (Asteliaceae, Asparagales)
Atamosco Adans. = Zephyranthes
Bloomeria Kellogg (Asparagaceae: Brodiaeoideae, Asparagales)
Brodiaea Sm. (Asparagaceae: Brodiaeoideae, Asparagales)
Calochortus Pursh
Camassia Lindl. Quamash (Asparagaceae: Agavoideae)
Cardiocrinum (Endl.) Lindl.
Chamaelirium Willd. (Melanthiaceae)
Chionodoxa Boiss. (Asparagaceae: Scilloideae, Asparagales)
Chlorogalum Kunth (Asparagaceae: Agavoideae, Asparagales)
Chlorophytum Ker-Gawl. (Asparagaceae: Agavoideae, Asparagales)
Clintonia Raf.
Colchicum L. (Colchicaceae)
Convallaria L. Lily of the valley (Asparagaceae: Nolinoideae, Asparagales)
Cooperia Herbert (Amaryllidaceae: Amaryllidoideae, Asparagales)
Cordyline Comm. ex R. Br. (Asparagaceae: Lomandroideae, Asparagales
Crinum L. (Amaryllidaceae: Amaryllidoideae, Asparagales)
Curculigo Gaertn. (Hypoxidaceae, Asparagales)
Dasylirion Zucc. (Asparagaceae: Nolinoideae, Asparagales)
Dianella Lam. (Xanthorrhoeaceae: Hemerocallidoideae, Asparagales)
Dichelostemma Kunth (Asparagaceae: Brodiaeoideae, Asparagales)
Disporum Salisb. ex D. Don (Colchicaceae)

(Asian species. N American species restored as Prosartes D. Don)

Dracaena L. (Asparagaceae: Nolinoideae, Asparagales)
Echeandia Ortega (Asparagaceae: Agavoideae, Asparagales)
Eremocrinum M.E. Jones (Asparagaceae: Agavoideae, Asparagales)
Erythronium L.
Eucharis Planch. & Linden (Amaryllidaceae: Amaryllidoideae, Asparagales)
Fritillaria L.
Gagea Salisb.
Galanthus L. (Amaryllidaceae: Amaryllidoideae, Asparagales)
Gloriosa L. (Colchicaceae)
Habranthus Herbert (Amaryllidaceae: Amaryllidoideae, Asparagales)
Harperocallis McDaniel (Tofieldiaceae, Alismatales)
Hastingsia S. Wats. (Asparagaceae: Agavoideae, Asparagales)
Helonias L. (Melanthiaceae, Asparagales)
Hemerocallis L. Daylily (Xanthorrhoeaceae: Hemerocallidoideae, Asparagales)
Hesperanthes S.Watson = Echeandia
Hesperocallis Gray (Tofieldiaceae, Alismatales)
Hesperoscordum Lindl. = Triteleia
Hippeastrum Herbert "Amaryllis" (Amaryllidaceae: Amaryllidoideae, Asparagales)
Hosta Tratt. (Asparagaceae: Agavoideae, Asparagales)
Hyacinthoides Medik. (Asparagaceae: Scilloideae, Asparagales)

Template:Multicol-break

Hyacinthus L. (Asparagaceae: Scilloideae, Asparagales)
Hymenocallis Salisb. (Amaryllidaceae: Amaryllidoideae, Asparagales)
Ipheion Raf. (Amaryllidaceae, Allioideae, Asparagales)
Kniphofia Moench (Xanthorrhoeaceae: Asphodeloideae, Asparagales)
Leucocrinum Nutt. ex Gray (Asparagaceae, Lomandroideae, Asparagales)
Leucojum L. Snowflake (Amaryllidaceae: Amaryllidoideae, Asparagales)
Lilium L.
Liriope Lour. Lilyturf (Asparagaceae: Nolinoideae, Asparagales)
Lloydia Salisb. ex Reichenb. = Gagea
Lophiola Ker-Gawl. (Nartheciaceae, Dioscoreales)
Lycoris Herbert (Amaryllidaceae: Amaryllidoideae, Asparagales)
Maianthemum G.H. Weber ex Wiggers False Solomon's seal

(Asparagaceae: Nolinoideae, Asparagales)

Medeola L.
Melanthium L. (Melanthiaceae)
Milla Cav. (Asparagaceae: Brodiaeoideae, Asparagales)
Muilla S. Wats. ex Benth. (Asparagaceae: Brodiaeoideae, Asparagales)
Muscari P. Mill. (Asparagaceae: Scilloideae, Asparagales)
Narcissus L. Jonquil, daffodil (Amaryllidaceae: Amaryllidoideae, Asparagales)
Narthecium Huds. (Nartheciaceae, Dioscoreales)
Nolina Michx. (Asparagaceae: Nolinoideae, Asparagales)
Nomocharis Franch.
Notholirion Wall. ex Boiss.
Nothoscordum Kunth (Amaryllidaceae: Allioideae, Asparagales)
Odontostomum Torr. (Tecophilaeaceae, Asparagales)
Ophiopogon Ker-Gawl. (Asparagaceae: Nolinoideae, Asparagales)
Ornithogalum L. (Asparagaceae: Scilloideae, Asparagales)
Pancratium L. (Amaryllidaceae: Amaryllidoideae, Asparagales)
Pleea Michx. (Tofieldiaceae, Alismatales)
Pleomele Salisb. (Asparagaceae: Nolinoideae, Asparagales)
Polygonatum P. Mill. Solomon's seal (Asparagaceae: Nolinoideae, Asparagales)
Sansevieria Thunb. (Asparagaceae: Nolinoideae, Asparagales)
Schoenocaulon Gray (Melanthiaceae)
Schoenolirion Torr. ex Dur. (Asparagaceae: Agavoideae, Asparagales)
Scilla L. (Asparagaceae: Scilloideae, Asparagales)
Scoliopus Torr.
Smilacina Desf. = Maianthemum
Stenanthium (Gray) Kunth (Melanthiaceae)
Sternbergia Waldst. & Kit. (Amaryllidaceae: Amaryllidoideae, Asparagales)
Streptopus Michx.
Tofieldia Huds. (Tofieldiaceae, Alismatales)
Tricyrtis Wallich
Trillium L. Trillium (Melanthiaceae)
Tristagma Poepp. (Amaryllidaceae: Allioideae, Asparagales)
Triteleia Dougl. ex Lindl. (Asparagaceae: Brodiaeoideae, Asparagales)
Triteleiopsis Hoover (Asparagaceae: Brodiaeoideae, Asparagales)
Tulipa L.
Urginea Steinh. (Asparagaceae: Scilloideae, Asparagales)
Uvularia L. (Colchicaceae)
Vagnera Adans. = Maianthemum
Veratrum L. (Melanthiaceae)
Xerophyllum Michx. (Melanthiaceae)
Zephyranthes Herbert (Amaryllidaceae: Amaryllidoideae, Asparagales)
Zigadenus Michx. (Melanthiaceae)

|}

The following includes other genera that have historically been classified in family Liliaceae s.l.:

Albuca L. (Asparagaceae: Scilloideae, Asparagales)
Anticlea Kunth (Melanthiaceae)
ArthropodiumR.Br. (Asparagaceae, Asparagales)
BellevaliaLapeyr. (Asparagaceae: Scilloideae, Asparagales)
BomareaMirb. (Alstroemeriaceae)
Chionographis Maxim. (Melanthiaceae)
Conanthera Ruiz & Pav. (Tecophilaeaceae, Asparagales)
Cyclobothra D.Don = Calochortus
Daiswa Raf. (Melanthiaceae)
Drimia Jacq. (Asparagaceae: Scilloideae, Asparagales)
Drimiopsis Lindl. & Paxton (Asparagaceae: Scilloideae, Asparagales)
Eremurus M.Bieb. (Xanthorrhoeaceae: Asphodeloideae, Asparagales)
Eriospermum Jacq. (Asparagaceae: Nolinoideae, Asparagales)
Eucrosia (Amaryllidaceae: Amaryllidoideae, Asparagales)
Eustephia Cav. (Amaryllidaceae: Amaryllidoideae, Asparagales)
Griffinia Ker Gawl. (Amaryllidaceae: Amaryllidoideae, Asparagales)
Heloniopsis A.Gray (Melanthiaceae)
Hemiphylacus S.Watson (Asparagaceae: Asparagoideae, Asparagales)
Herreria Ruiz & Pav. (Asparagaceae: Agavoideae, Asparagales)
Hessea Herb. (Amaryllidaceae: Amaryllidoideae, Asparagales)
Hookera Salisb. = Brodiaea
Hypoxis (Hypoxidaceae, Asparagales)
Ismene Salisb. (Amaryllidaceae: Amaryllidoideae, Asparagales)

Jaimehintonia B.L.Turner (Amaryllidaceae: Allioideae, Asparagales)
Johnsonia Mill. (Asparagaceae: Lomandroideae, Asparagales)
Laxmannia R.Br. (Asparagaceae: Lomandroideae, Asparagales)
Lepidopharynx Rusby = Hippeastrum
Metanarthecium Maxim. (Nartheciaceae, Dioscoreales)
Miersia Lindl. (Amaryllidaceae: Allioideae, Asparagales)
Molineria Colla (Hypoxidaceae, Asparagales)
Nietneria Klotzsch (Melanthiaceae)
Paradisea Mazzuc. (Anthericaceae, Asparagales)
Paris L. (Melanthiaceae)
Peliosanthes Andrews (Asparagaceae: Nolinoideae, Asparagales)
Phaedranassa Herb. (Amaryllidaceae: Amaryllidoideae, Asparagales)
Selonia Regel = Eremurus
Stropholirion Torr. = Dichelostemma
Theropogon Maxim. (Amaryllidaceae: Allioideae, Asparagales)
Tovaria Neck. (Asparagaceae: Nolinoideae, Asparagales)
Toxicoscordion Rydb. (Melanthiaceae)
Tracyanthus Small = Stenanthium
Tulbaghia Heist. Society garlic (Amaryllidaceae: Allioideae, Asparagales)
Tupistra Ker Gawl. (Asparagaceae: Nolinoideae, Asparagales)
Urceolina Rchb. (Amaryllidaceae: Amaryllidoideae, Asparagales)
Wurmbea Thunb. (Colchicaceae, Asparagales)

Modern classification

Various authorities (e.g. ITIS 16,^[84] GRIN 27,^[85] WCSP,^[86] NCBI,^[87] DELTA^[88] ) differ on the exact number of genera included in Liliaceae s.s., but generally there are about fifteen to sixteen genera, depending on whether or not Amana is included in Tulipa and Lloydia in Gagea. For instance Amana is still listed separately in WCSP.

Currently the APWeb lists fifteen genera, arranged as follows:^[81]^[89]

Subfamily: Lilioideae Eaton
- Tribe: Medeoleae Benth. (synonyms: Medeolaceae Takht., Medeoloideae Benth.)
  - Clintonia Raf.
  - Medeola Gronov. ex L.
- Tribe: Lilieae Ritgen (synonyms: Erythroniaceae Martinov, Fritillariaceae Salisb., Liriaceae Borkh., Tulipaceae Borkh.)
  - Cardiocrinum (Endl.) Lindl.
  - Erythronium L. – trout lily
  - Fritillaria Tourn. ex L.
  - Gagea Salisb. (including Lloydia Salisb. ex Rchb.)
  - Lilium Tourn. ex L. – lily
  - Nomocharis Franch.
  - Notholirion Wall. ex Boiss.
  - Tulipa L. (including Amana Honda) – tulip
Subfamily: Calochortoideae Dumort. (synonyms: Calochortaceae Dumort., Compsoaceae Horan., nom. illeg., Tricyrtidaceae Takht., nom. cons.)
- Calochortus Pursh
- Tricyrtis Wall.
Subfamily: Streptopoideae (synonym: Scoliopaceae Takht.)
- Prosartes D.Don
- Scoliopus Torr.
- Streptopus Michx.

The largest genera are Gagea (200), Fritillaria (130), Lilium (110), and Tulipa (75 species), all within the Lilieae tribe.

Etymology

From Lilium the type species which is Latin for Lily, which in turn came from the Greek λείριον (leírion).

References

^ Lobstein, Marion Blois. "Where Have All the Lillies Gone? Long-Time Changin' in the Liliaceous Families". Prince William Wildflower Society. Retrieved 9 February 2014.
^ Adanson, Michel (1763). "VIII Liliaceae". Familles des plantes. Paris: Vincent. p. Part II: 42. Retrieved 9 February 2014.
^ Jussieu, Antoine Laurent de (1789). "Lilia". Genera Plantarum, secundum ordines naturales disposita juxta methodum in Horto Regio Parisiensi exaratam. Paris. pp. 48–49. OCLC 5161409. Retrieved 4 February 2014.
^ "Names of families and subfamilies, tribes and subtribes". p. 18.2. Retrieved 1 February 2014. In ICN (2011).
^ ^a ^b "Des familles et des tribus". pp. 192–195 http://www.biodiversitylibrary.org/item/88297#page/204/mode/1up. Retrieved 5 February 2014. {{cite book}}: Missing or empty |title= (help) In A. P. de Candolle (1813).
^ de Jussieu, Antoine Laurent (1778). "Examen de la famille des Renoncules". Histoire de l'Académie royale des sciences avec les mémoires de mathématique et physique Année 1773. 1. Paris: Académie royale des sciences: 228. Retrieved 4 February 2014.
^ "Liliaceae". p. vol 2: 173 http://www.biodiversitylibrary.org/item/95185#page/185/mode/1up. Retrieved 4 February 2014. {{cite book}}: Missing or empty |title= (help) In Gray (1821),
^ Lindley, John (1830). "CCLI: Liliaceae". An introduction to the natural system of botany : or, A systematic view of the organisation, natural affinities, and geographical distribution, of the whole vegetable kingdom : together with the uses of the most important species in medicine, the arts, and rural or domestic economy. London: Longman. p. 279. Retrieved 2 February 2014.
^ Redouté, P. J. (1802–1816). Les liliacées. Paris: Redouté. Retrieved 2 February 2014.{{cite book}}: CS1 maint: date format (link)
^ "An Arrangement of the Families, and of the anomalous Genera of phenogamous Plants, according to their sexual Organs". p. vol 2: vi http://www.biodiversitylibrary.org/item/95185#page/10/mode/1up. Retrieved 4 February 2014. {{cite book}}: Missing or empty |title= (help) In Gray (1821).
^ "The genera of British plants, according to their mutual relations, with the number of species in each genus. 2. Plantae Endogenae: B Plantae Endogenae Phenogamae - 14. Liliaceae". p. vol 1: xx http://www.biodiversitylibrary.org/item/95186#page/26/mode/1up. Retrieved 5 February 2014. {{cite book}}: Missing or empty |title= (help) In Gray (1821).
^ ^a ^b "Esquisse. D'une Série linéaire et par conséquent artificielle, pour la disposition des familles naturelles du règne végetal". p. 219 http://www.biodiversitylibrary.org/item/88297#page/231/mode/1up. Retrieved 5 February 2014. {{cite book}}: Missing or empty |title= (help) In A. P. de Candolle (1813).
^ von Marilaun, Anton Kerner (1895-6). The natural history of plants, their forms, growth, reproduction, and distribution', trans. FW Oliver et al. from Pflanzenleben, 1890–1891. New York: Holt. p. vol 4: 603. Retrieved 5 February 2014. {{cite book}}: Check date values in: |year= (help)
^ ^a ^b Lindley, John (1846). "Order LXII: Liliaceae - Lilyworts". The Vegetable Kingdom: or, The structure, classification, and uses of plants, illustrated upon the natural system. London: Bradbury. pp. 200–205. Retrieved 5 February 2014.
^ ^a ^b Bentham, G. (1883). "Liliaceae". Genera plantarum ad exemplaria imprimis in herbariis kewensibus servata definita. Vol III Part II. London: L Reeve & Co. pp. 748–836. Retrieved 24 January 2014. {{cite book}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ ^a ^b "Historical background of plant classification". pp. 10–29 http://books.google.ca/books?id=In_Lv8iMt24C&pg=10. Retrieved 23 January 2014. {{cite book}}: Missing or empty |title= (help) In Singh (2004).
^ ^a ^b Bessey, Charles E. (Feb.–Apr., 1915). "The Phylogenetic Taxonomy of Flowering Plants". Annals of the Missouri Botanical Garden. 2 (1/2): 109–164. doi:10.2307/2990030. Retrieved 12 February 2014. {{cite journal}}: Check date values in: |date= (help)
^ ^a ^b Cronquist, A (1981). An integrated system of classification of flowering plants. New York: Columbia University Press.
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^ ^a ^b ^c Rasmussen, F. N. "Superorder Liliiflorae". pp. 107–274. ISBN 978-3-642-64903-5 http://link.springer.com/chapter/10.1007/978-3-642-61663-1_11. Retrieved 10 February 2014. {{cite book}}: Missing or empty |title= (help) In Dahlgren, Clifford & Yeo (1985).
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^ Huber, H (1969). "Die Samenmerkmale und Verwandtschaftsverhältnisse der Liliiflorae". Mitt. Bot. Staatssamml. Muenchen. 8: 219–538.
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^ ^a ^b ^c ^d Patterson, T. B. (2002). "Phylogeny, concerted convergence, and phylogenetic niche conservatism in the core Liliales: insights from rbcL and ndhF sequence data" (PDF). Evolution. 56: 233–252. Retrieved 14 January 2014. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
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^ Hilu, K.; Borsch, T.; Muller, K.; Soltis, D.E.; Soltis, P.S.; Savolainen, V.; Chase, M.W.; Powell, M.P.; Alice, L.A.; Evans, R.; Sauquet, H.; Neinhuis, C.; Slotta, T.A.B.; Rohwer, J.G.; Campbell, C.S.; Chatrou, L.W. (December 2003). "Angiosperm phylogeny based on <011> matK sequence information". American Journal of Botany. 90 (12): 1758–1766. doi:10.3732/ajb.90.12.1758. Retrieved 4 January 2014.
^ Pires, J.C.; Maureira, I.J.; Givnish, T.J.; Sytsma, K.J.; Seberg, O.; Petersen, G.; Davis, J.I.; Stevenson, D.W.; Rudall, P.J.; Fay, M.F.; Chase, M.W. (2006). "Phylogeny, genome size, and chromosome evolution of Asparagales". 22: 278–304. Retrieved 4 January 2014. {{cite journal}}: Cite journal requires |journal= (help)
^ Rudall, P.; Furness, C.A.; Chase, M.W.; Fay, M.F. (1997). "Microsporogenesis and pollen sulcus type in Asparagales (Lilianae)". Canad. J. Bot. 75 (3): 408–430. ISSN 0008-4026. Retrieved 4 January 2014.
^ Källersjö, M.; Farris, J.S.; Chase, M.W.; Bremer, B.; Fay, M.F.; Humphries, C.J.; Petersen, G.; Seberg, O.; Bremer, K. (1998). "Simultaneous parsimony jacknife analysis of 2538rbcL DNA sequences reveals support for major clades of green plants, land plants, and flowering plants". Pl. Syst. Evol. 213 (3–4): 259–287. doi:10.1007/BF00985205.
^ Fay, M.F. (2000). "Phylogenetic studies of Asparagales based on four plastid DNA regions". pp. 360–371 http://ninaronsted.dk/page1/files/page1_6.pdf. {{cite book}}: Missing or empty |title= (help) In Wilson & Morrison (2000).
^ Soltis, D. E.; Soltis, P.S.; Chase, M.W.; Mort, M.E.; Albach, D.C.; Zanis, M.; Savolainen, V.; Hahn, W.H.; Hoot, S.B.; Fay, M.F.; Axtell, M.; Swensen, S.M.; Prince, L.M.; Kress, W.J.; Nixon, K.C.; Farris, J.S. (2000). "Angiosperm phylogeny inferred from 18S rDNA, rbcL, and atpB sequences". Bot. J. Linn. Soc. 133 (4): 381–461. Retrieved 4 January 2014.
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^ Goldblatt, P. (1995). "The status of R. Dahlgren's orders Liliales and Melanthiales". In Rudall, P.J.; Cribb, P.J.; Cutler, D.F.; Humphries, C.J. (eds.). Monocotyledons: Systematics and evolution. Kew: Royal Botanic Gardens. pp. 181–200. Retrieved 4 January 2014. {{cite book}}: Missing |editor1= (help)
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^ Soltis, D.E.; Soltis, P.F.; Endress, P.K.; Chase, M.W. (2005). Phylogeny and evolution of angiosperms. Sunderland, MA: Sinauer.
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^ Jeffrey D. Stilwell, Rodney M. Feldmann, ed. (2000). Paleobiology and Paleoenvironments of Eocene Rocks: McMurdo Sound, East Antarctica. American Geophysical Union. p. 162. ISBN 0-87590-947-7. Retrieved 28 January 2014.
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^ Botanical authorities from WCSP.^[86]

[Lobstein-1] Lobstein, Marion Blois. "Where Have All the Lillies Gone? Long-Time Changin' in the Liliaceous Families". Prince William Wildflower Society. Retrieved 9 February 2014.

[Adanson-2] Adanson, Michel (1763). "VIII Liliaceae". Familles des plantes. Paris: Vincent. p. Part II: 42. Retrieved 9 February 2014.

[Jussieu-3] Jussieu, Antoine Laurent de (1789). "Lilia". Genera Plantarum, secundum ordines naturales disposita juxta methodum in Horto Regio Parisiensi exaratam. Paris. pp. 48–49. OCLC 5161409. Retrieved 4 February 2014.

[icn18.2-4] "Names of families and subfamilies, tribes and subtribes". p. 18.2. Retrieved 1 February 2014. In ICN (2011).

[Candolle2-5] "Des familles et des tribus". pp. 192–195 http://www.biodiversitylibrary.org/item/88297#page/204/mode/1up. Retrieved 5 February 2014. {{cite book}}: Missing or empty |title= (help) In A. P. de Candolle (1813).

[JussieuRan-6] Jussieu, Antoine Laurent (1778). "Examen de la famille des Renoncules". Histoire de l'Académie royale des sciences avec les mémoires de mathématique et physique Année 1773. 1. Paris: Académie royale des sciences: 228. Retrieved 4 February 2014.

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[Lindley2-14] Lindley, John (1846). "Order LXII: Liliaceae - Lilyworts". The Vegetable Kingdom: or, The structure, classification, and uses of plants, illustrated upon the natural system. London: Bradbury. pp. 200–205. Retrieved 5 February 2014.

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[Singh2-16] "Historical background of plant classification". pp. 10–29 http://books.google.ca/books?id=In_Lv8iMt24C&pg=10. Retrieved 23 January 2014. {{cite book}}: Missing or empty |title= (help) In Singh (2004).

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[Cronquist_1981-18] Cronquist, A (1981). An integrated system of classification of flowering plants. New York: Columbia University Press.

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[Dahlgren_et_al._1985-21] Rasmussen, F. N. "Superorder Liliiflorae". pp. 107–274. ISBN 978-3-642-64903-5 http://link.springer.com/chapter/10.1007/978-3-642-61663-1_11. Retrieved 10 February 2014. {{cite book}}: Missing or empty |title= (help) In Dahlgren, Clifford & Yeo (1985).

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[Lotsy-25] Lotsy, Johannes Paulus (1911). "Liliifloren". Vorträge über botanische Stammesgeschichte, gehalten an der Reichsuniversität zu Leiden. Ein Lehrbuch der Pflanzensystematik. vol 3: Cormophyta Siphonogamia. Jena: Fischer. p. 715. Retrieved 9 February 2014.

[Huber-26] Huber, H (1969). "Die Samenmerkmale und Verwandtschaftsverhältnisse der Liliiflorae". Mitt. Bot. Staatssamml. Muenchen. 8: 219–538.

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[Kelch-28] Kelch, D. G. (2002). "Consider the Lilies" (PDF). Fremontia. 30 (2): 23–29.

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[APG_II_2003-30] APG II (2003). "An Update of the Angiosperm Phylogeny Group Classification for the orders and families of flowering plants: APG II". Botanical Journal of the Linnean Society. 141 (141): 399–436. Retrieved 3 January 2014.

[Chase_et_al._1995a-31] Chase, M. W.; Duvall, M. R.; Hills, H. G.; Conran, J. G.; Cox, A. V.; Eguiarte, L. E.; Hartwell, J.; Fay, M. F.; Caddick, L. R.; Cameron, K. M.; Hoot, S. "Molecular phylogenetics of Lilianae". pp. 109–137. {{cite book}}: Missing or empty |title= (help) In Rudall et al. (1995).

[Patterson_2002-32] Patterson, T. B. (2002). "Phylogeny, concerted convergence, and phylogenetic niche conservatism in the core Liliales: insights from rbcL and ndhF sequence data" (PDF). Evolution. 56: 233–252. Retrieved 14 January 2014. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[APG-33] APG (1998). "An ordinal classification for the families of flowering plants". Annals of the Missouri Botanical Garden. 85: 531-553. (4): 531–553. Retrieved 5 January 2014.

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[bremer-35] Bremer, K. (1995, 1996, 1997). Introduction to Phylogeny and Systematics of Flowering Plants. 1st, 2nd, 3rd eds. Uppsala University. {{cite book}}: |first2= missing |last2= (help); |first3= missing |last3= (help); Check date values in: |year= (help); Missing pipe in: |first2= (help); Missing pipe in: |first3= (help)CS1 maint: numeric names: authors list (link)

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[bremernet-37] Bremer, K. (1996 onwards). "Classification of flowering plants". Retrieved 6 January 2014. {{cite web}}: |first2= missing |last2= (help); |first3= missing |last3= (help); Check date values in: |year= (help); Missing pipe in: |first2= (help); Missing pipe in: |first3= (help)CS1 maint: numeric names: authors list (link)

[APG_III_2009-38] APG III (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. Retrieved 3 January 2014.

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[Rudall_et_al._2000a-40] Rudall, P. J. Consider the lilies: systematics of Liliales. pp. 347–359. Retrieved 14 January 2014. {{cite book}}: Unknown parameter |coauthors= ignored (|author= suggested) (help) In Wilson & Morrison (2000)

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[Hilu_et_al._2003-45] Hilu, K.; Borsch, T.; Muller, K.; Soltis, D.E.; Soltis, P.S.; Savolainen, V.; Chase, M.W.; Powell, M.P.; Alice, L.A.; Evans, R.; Sauquet, H.; Neinhuis, C.; Slotta, T.A.B.; Rohwer, J.G.; Campbell, C.S.; Chatrou, L.W. (December 2003). "Angiosperm phylogeny based on <011> matK sequence information". American Journal of Botany. 90 (12): 1758–1766. doi:10.3732/ajb.90.12.1758. Retrieved 4 January 2014.

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[Rudall_et_al._1997a-47] Rudall, P.; Furness, C.A.; Chase, M.W.; Fay, M.F. (1997). "Microsporogenesis and pollen sulcus type in Asparagales (Lilianae)". Canad. J. Bot. 75 (3): 408–430. ISSN 0008-4026. Retrieved 4 January 2014.

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[Fay_et_al._2000-49] Fay, M.F. (2000). "Phylogenetic studies of Asparagales based on four plastid DNA regions". pp. 360–371 http://ninaronsted.dk/page1/files/page1_6.pdf. {{cite book}}: Missing or empty |title= (help) In Wilson & Morrison (2000).

[Soltis_et_al._2000-50] Soltis, D. E.; Soltis, P.S.; Chase, M.W.; Mort, M.E.; Albach, D.C.; Zanis, M.; Savolainen, V.; Hahn, W.H.; Hoot, S.B.; Fay, M.F.; Axtell, M.; Swensen, S.M.; Prince, L.M.; Kress, W.J.; Nixon, K.C.; Farris, J.S. (2000). "Angiosperm phylogeny inferred from 18S rDNA, rbcL, and atpB sequences". Bot. J. Linn. Soc. 133 (4): 381–461. Retrieved 4 January 2014.

[Stevenson_et_al._2000-51] 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". pp. 17–24. {{cite book}}: Missing or empty |title= (help) In Wilson & Morrison (2000).

[McPherson_and_Graham_2001-52] McPherson, M.A.; Graham,, S.W. (2001). "Inference of Asparagales phylogeny using a large chloroplast data set". Proceedings of the Botanical Society of America: 126. {{cite journal}}: |format= requires |url= (help)CS1 maint: extra punctuation (link)

[Goldblatt_1995-53] Goldblatt, P. (1995). "The status of R. Dahlgren's orders Liliales and Melanthiales". In Rudall, P.J.; Cribb, P.J.; Cutler, D.F.; Humphries, C.J. (eds.). Monocotyledons: Systematics and evolution. Kew: Royal Botanic Gardens. pp. 181–200. Retrieved 4 January 2014. {{cite book}}: Missing |editor1= (help)

[Stevenson_and_Loconte_1995-54] Stevenson, D.W.; Loconte, H. (1995). "Cladistic analysis of monocot families". In Rudall, P.J.; Cribb, P.J.; Cutler, D.F. (eds.). pp. 543–578. {{cite book}}: Missing |editor1= (help); Missing or empty |title= (help)

[Vinnersten_and_Bremer_2001-55] Vinnersten, A.; Bremer, K. (September 2001). "Age and biogeography of major clades in Liliales". Amer. J. Bot. 88 (9): 1695–1703. Retrieved 5 January 2014.

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[Zomlefer_2001-59] Zomlefer, W.B.; Williams, N.H.; Whitten, W.M.; Judd, W.S. (2001). "Generic circumscription and relationships in the tribe Melanthieae (Liliales, Melanthiaceae), with emphasis on Zigadenus: evidence from ITS and trnL-F sequence data". American Journal of Botany. 88: 1657–1669. Retrieved 14 January 2014.

[Soltis_et_al._2005-60] Soltis, D.E.; Soltis, P.F.; Endress, P.K.; Chase, M.W. (2005). Phylogeny and evolution of angiosperms. Sunderland, MA: Sinauer.

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[89] Botanical authorities from WCSP.^[86]

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@@ Line 16: / Line 16: @@
 == Modern APG classification ==
 To meet this need, flowering plant [[systematist]]s formed the Angiosperm Phylogeny Group ([[Angiosperm Phylogeny Group|APG]]) resulting in a new classification published in 1998<ref name=APG/> largely based on the work of [[Kåre Bremer]] and colleagues at [[Uppsala University|Uppsala]] and [[University of Stockholm|Stockholm]] universities in the late 1970s,
-<ref name = "Bremer and Wanntorp78"/><ref name=bremer/><ref name=Williams/> which became available on the internet in 1996.<ref name=bremernet/>
+<ref name = "Bremer and Wanntorp78"/><ref name=bremer/><ref name=Williams/> which became available on the internet in 1996.<ref name=bremernet>{{cite web |last=Bremer |first=K. |first2=B. last2=Bremer |first3=M. last3=Thulin |title=Classification of flowering plants |url=http://www.egs.uu.se/sysbiol/classification/classification.htm |accessdate=6 January 2014 |year=1996 onwards}}</ref>
 This scheme was ordinal (concentrating on orders rather than families), prioritising [[monophyly]]. However progress was rapid and the modern era of the taxonomy of the Liliaceae family comes from [[Walter S Judd|Judd]] et al.<ref name="Judd 2007"/> (second edition 2002), the [[APG II]] (2003<ref name="APG II 2003"/>) and [[APG III]] (2009
+<ref name="APG III 2009">{{cite journal |last = APG III |title=An Update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. |year = 2009 |journal=Botanical Journal of the Linnean Society |volume=161 |issue=2 |pages=105–121 |url=http://www3.interscience.wiley.com/journal/122630309/abstract |accessdate=3 January 2014}}</ref>), while the Linear APG III assigned it the family number 61.
-<ref name="APG III 2009"/>), while the Linear APG III assigned it the family number 61.
-<ref name=lapg/> While the original APG did not specifically address the issues of the [[polyphyly]] within Liliaceae, APG II did so within the two closely morphologically related orders, Liliales and Asparagales<ref name="Rudall et al. 2000a"/> recognising the continued common use of the ''[[sensu lato]]'' (''s.l.'').
+<ref name=lapg>{{cite journal |last=LAPG III |title=The Linear Angiosperm Phylogeny Group (LAPG) III: a linear sequence of the families in APG III |journal=Botanical Journal of the Linnean Society |year=2009 |volume=161 |issue=2 |pages=128–131 |url=http://onlinelibrary.wiley.com/doi/10.1111/j.1095-8339.2009.01000.x/abstract |accessdate=3 January 2014}}</ref> While the original APG did not specifically address the issues of the [[polyphyly]] within Liliaceae, APG II did so within the two closely morphologically related orders, Liliales and Asparagales<ref name="Rudall et al. 2000a">{{cite book| last =Rudall| first =P. J.| coauthors =Stobart, K. L., Hong, W-P., Conran, J. G., Furness, C. A., Kite, G. C., and Chase, M. W.| pages =347–359
+| title=Consider the lilies: systematics of Liliales. | url=http://books.google.ca/books?id=YzQBUQqLS0YC&pg=PT833&lpg=PT833|accessdate=14 January 2014}} In {{Harvtxt|Wilson|Morrison|2000}}</ref> recognising the continued common use of the ''[[sensu lato]]'' (''s.l.'').
+These studies of [[DNA]] and [[morphology (biology)|morphological]] data (particularly reproductive morphology) together with [[Phylogenetics|phylogenetic analyses]], allowed the conclusion that the "[[lilioid monocot|petaloid monocots]]" do not really belong to one botanical family but rather are distributed across two separate different orders, the [[Asparagales]] and [[Liliales]]. The [[monophyly]] of these newly defined orders is supported by [[cladistic]] analysis based on morphology, [[18S rDNA]], and many other DNA sequences, particularly the [[plastid]] [[gene]] [[rbcL|''rbc''L]].<ref name="Chase et al. 2000">{{cite book |last=Chase |first=M.W. |last2=Soltis |first2=D. E. |last3=Soltis |first3=P. S. |last4=Rudall |first4=P. J. |last5=Fay |first5=M. F. |last6=Hahn |first6=W. H. |last7=Sullivan |first7=S. |last8=Joseph |first8=J. |last9=Molvray |first9=M. |last10=Kores |first10=P. J. |last11=Givnish |first11=T. J. |last12=Sytsma |first12=K. J. |last13=Pires |first13=J. C. |title=Monocots: Systematics and evolution |year=2000 |publisher=[[CSIRO]] |location=Collingwood, Australia |pages=3–16 |editor2-last=Wilson |editor2-first=K.L. |editor3-last=Morrison |editor3-first=D.A. |chapter=Higher-level systematics of the monocotyledons: An assessment of current knowledge and a new classification}}</ref><ref name="Chase et al. 2006">{{cite journal |last=Chase |first=M. W. |last2=Fay |first2=M. F. |last3=Devey |first3=D. S. |last4=Maurin |first4=O |last5=Rønsted |first5=N |last6=Davies |first6=T. J |last7=Pillon |first7=Y |last8=Petersen |first8=G |last9=Seberg |first9=O |last10=Tamura |first10=M. N. |last11=Asmussen |first11=C. B. |last12=Hilu |first12=K |last13=Borsch |first13=T |last14=Davis |first14=J. I |last15=Stevenson |first15=D. W. |last16=Pires |first16=J. C. |last17=Givnish |first17=T. J. |last18=Sytsma |first18=K. J. |last19=McPherson |first19=M. A. |last20=Graham |first20=S. W. |last21=Rai |first21=H. S. |title=Multigene analyses of monocot relationships : a summary |pages=63–75 |url=http://www.ninaronsted.dk/page1/files/page1_5.pdf |accessdate=4 January 2014 |issn=00656275}} In {{Harvtxt|Columbus|Friar|Porter|Prince|2006}}</ref><ref name="Davis et al. 2004">{{cite journal |last=Davis |first=J.I. |last2=Stevenson |first2=D.W. |last3=Petersen |first3=G. |last4=Seberg |first4=O. |last5=Campbell |first5=L.M. |last6=Freudenstein |first6=J.V. |last7=Goldman |first7=D.H. |last8=Hardy |first8=C.R. |last9=Michelangeli |first9=F.A. |last10=Simmons |first10=M.P. |last11=Specht |first11=C.D. |last12=Vergara-Silva |first12=F. |last13=Gandolfo |first13=M. |title=A phylogeny of the monocots, as inferred from ''rbcL'' and ''atpA'' sequence variation, and a comparison of methods for calculating jacknife and bootstrap values |journal=[[Syst. Bot.]] |date=July–September 2004 |volume=29 |issue=3 |pages=467–510 |url=http://www.ingentaconnect.com/content/aspt/sb/2004/00000029/00000003/art00001 |accessdate=4 January 2014}}</ref><ref name="Graham et al. 2006">{{cite journal |last=Graham |first=S.W. |last2=Zgurski |first2=J.M. |last3=McPherson |first3=M.A. |last4=Cherniawsky |first4=D.M. |last5=Saarela |first5=J.M. |last6=Horne |first6=E.S.C. |last7=Smith |first7=S.Y. |last8=Wong |first8=W.A. |last9=O'Brien |first9=H.E. |last10=Biron |first10=V.L. |last11=Pires |first11=J.C. |last12=Olmstead |first12=R.G. |last13=Chase |first13=M.W. |last14=Rai |first14=H.S. |title=Robust inference of monocot deep phylogeny using an expanded multigene plastid data set |pages=3–21 |url=http://depts.washington.edu/phylo/OlmsteadPubs/Graham.2006.Aliso.pdf |accessdate=4 January 2014}} In {{Harvtxt|Columbus|Friar|Porter|Prince|2006}}</ref><ref name="Hilu et al. 2003">{{cite journal |last=Hilu |first=K. |last2=Borsch |first2=T. |last3=Muller |first3=K. |last4=Soltis |first4=D.E. |last5=Soltis |first5=P.S. |last6=Savolainen |first6=V. |last7=Chase |first7=M.W. |last8=Powell |first8=M.P. |last9=Alice |first9=L.A. |last10=Evans |first10=R. |last11=Sauquet |first11=H. |last12=Neinhuis |first12=C. |last13=Slotta |first13=T.A.B. |last14=Rohwer |first14=J.G. |last15=Campbell |first15=C.S. |last16=Chatrou |first16=L.W. |title=Angiosperm phylogeny based on ''<011>'' ''matK'' sequence information |journal=[[American Journal of Botany]] |date=December 2003 |volume=90 |issue=12 |pages=1758–1766 |doi=10.3732/ajb.90.12.1758 |url=http://www.amjbot.org/cgi/content/full/90/12/1758 |accessdate=4 January 2014}}</ref><ref name="Pires et al. 2006">{{cite journal |last=Pires |first=J.C. |last2=Maureira |first2=I.J. |last3=Givnish |first3=T.J. |last4=Sytsma |first4=K.J. |last5=Seberg |first5=O. |last6=Petersen |first6=G. |last7=Davis |first7=J.I. |last8=Stevenson |first8=D.W. |last9=Rudall |first9=P.J. |last10=Fay |first10=M.F. |last11=Chase |first11=M.W. |title=Phylogeny, genome size, and chromosome evolution of Asparagales |year=2006 |volume=22 |pages=278–304 |url=http://cat.inist.fr/?aModele=afficheN&cpsidt=18269727 |accessdate=4 January 2014}}</ref><ref name="Rudall et al. 1997a">{{cite journal |last=Rudall |first=P. |last2=Furness |first2=C.A. |last3=Chase |first3=M.W. |last4=Fay |first4=M.F. |title=Microsporogenesis and pollen sulcus type in Asparagales (Lilianae) |journal=[[Canad. J. Bot.]] |year=1997 |volume=75 |issue=3 |pages=408–430 |url=http://cat.inist.fr/?aModele=afficheN&cpsidt=10769694 |accessdate=4 January 2014 |issn=0008-4026}}</ref><ref name="Källersjö et al. 1998">{{cite journal |last=Källersjö |first=M. |last2=Farris |first2=J.S. |last3=Chase |first3=M.W. |last4=Bremer |first4=B. |last5=Fay |first5=M.F. |last6=Humphries |first6=C.J. |last7=Petersen |first7=G. |last8=Seberg |first8=O. |last9=Bremer |first9=K. |title=Simultaneous parsimony jacknife analysis of 2538''rbc''L DNA sequences reveals support for major clades of green plants, land plants, and flowering plants |journal=[[Pl. Syst. Evol.]] |year=1998 |volume=213 |issue=3-4 |pages=259–287 |doi=10.1007/BF00985205 |url=http://www.springerlink.com/content/u864187n02112656/}}</ref><ref name="Fay et al. 2000">{{cite book |last=Fay |first=M.F. |year=2000 |pages=360–371 |chapter=Phylogenetic studies of Asparagales based on four plastid DNA regions|url=http://ninaronsted.dk/page1/files/page1_6.pdf}} In {{Harvtxt|Wilson|Morrison|2000}}.</ref><ref name="Soltis et al. 2000">{{cite journal |last=Soltis |first=D. E. |last2=Soltis |first2=P.S. |last3=Chase |first3=M.W. |last4=Mort |first4=M.E. |last5=Albach |first5=D.C. |last6=Zanis |first6=M. |last7=Savolainen |first7=V. |last8=Hahn |first8=W.H. |last9=Hoot |first9=S.B. |last10=Fay |first10=M.F. |last11=Axtell |first11=M. |last12=Swensen |first12=S.M. |last13=Prince |first13=L.M. |last14=Kress |first14=W.J. |last15=Nixon |first15=K.C. |last16=Farris |first16=J.S. |title=Angiosperm phylogeny inferred from 18S rDNA, rbcL, and ''atpB'' sequences |journal=Bot. J. Linn. Soc. |year=2000 |volume=133 |issue=4 |pages=381–461 |url=http://www.ingentaconnect.com/content/ap/bt/2000/00000133/00000004/art00380 |accessdate=4 January 2014}}</ref><ref name="Stevenson et al. 2000">{{cite book |last=Stevenson |first=D.W. |last2=Davis |first2=J.I. |last3=Freudenstein |first3=J.V. |last4=Hardy |first4=C.R. |last5=Simmons |first5=M.P. |last6=Specht |first6=C.D. |year=2000 |pages=17–24 |chapter=A phylogenetic analysis of the monocotyledons based on morphological and molecular character sets, with comments on the placement of ''Acorus'' and Hydatellaceae}} In {{Harvtxt|Wilson|Morrison|2000}}.</ref><ref name="McPherson and Graham 2001">{{cite journal |last=McPherson |first=M.A. |last2=Graham, |first2=S.W. |title=Inference of Asparagales phylogeny using a large chloroplast data set |journal=Proceedings of the Botanical Society of America |year=2001 |page=126 |format=Abstract No. 120}}</ref><ref name="Goldblatt 1995">{{cite book |last=Goldblatt |first=P. |title=Monocotyledons: Systematics and evolution |year=1995 |publisher=Royal Botanic Gardens |location=Kew |pages=181–200 |editor2-last=Rudall |editor2-first=P.J. |editor3-last=Cribb |editor3-first=P.J. |editor4-last=Cutler |editor4-first=D.F. |editor5-last=Humphries |editor5-first=C.J. |chapter=The status of R. Dahlgren's orders Liliales and Melanthiales |url=http://www.press.uchicago.edu/ucp/books/book/distributed/M/bo9856357.html |accessdate=4 January 2014}}</ref><ref name="Stevenson and Loconte 1995">{{cite book |last=Stevenson |first=D.W. |last2=Loconte |first2=H. |year=1995 |pages=543–578 |editor2-last=Rudall |editor2-first=P.J. |editor3-last=Cribb |editor3-first=P.J. |editor4-last=Cutler |editor4-first=D.F.  |chapter=Cladistic analysis of monocot families}}</ref><ref name="Vinnersten and Bremer 2001">{{cite journal |last=Vinnersten |first=A. |last2=Bremer |first2=K. |title=Age and biogeography of major clades in Liliales |journal=Amer. J. Bot. |date=September 2001 |volume=88 |issue=9 |pages=1695–1703 |url=http://www.amjbot.org/cgi/content/abstract/88/9/1695 |accessdate=5 January 2014}}</ref><ref name="Givnish 2005"/><ref name="Givnish 2006"/><ref name="Rønsted 2005">{{cite journal|last=Rønsted|first=N., S.|coauthors=Law, H. Thornton, M. F. Fay, and M. W. Chase|title=Molecular phylogenetic evidence for the monophyly of ''Fritillaria'' and ''Lilium'' (Liliaceae; Liliales) and the infrageneric classification of ''Fritillaria''|journal=Molecular Phylogenetics and Evolution|year=2005|volume=35|pages=509–527|url=http://www.ncbi.nlm.nih.gov/pubmed/15878122|accessdate=14 January 2014}}</ref><ref name="Zomlefer 2001">{{cite journal|last=Zomlefer|first=W.B. |last2=Williams |first2=N.H. |last3=Whitten |first3=W.M. |last4=Judd |first4=W.S. |title=Generic circumscription and relationships in the tribe Melanthieae (Liliales, Melanthiaceae), with emphasis on ''Zigadenus'': evidence from ITS and ''trnL-F'' sequence data |journal=American Journal of Botany |year=2001 |volume=88 |pages=1657–1669 |url=http://www.ncbi.nlm.nih.gov/pubmed/21669700 |accessdate=14 January 2014}}</ref>
-These studies of [[DNA]] and [[morphology (biology)|morphological]] data (particularly reproductive morphology) together with [[Phylogenetics|phylogenetic analyses]], allowed the conclusion that the "[[lilioid monocot|petaloid monocots]]" do not really belong to one botanical family but rather are distributed across two separate different orders, the [[Asparagales]] and [[Liliales]]. The [[monophyly]] of these newly defined orders is supported by [[cladistic]] analysis based on morphology, [[18S rDNA]], and many other DNA sequences, particularly the [[plastid]] [[gene]] [[rbcL|''rbc''L]].<ref name="Chase et al. 2000"/><ref name="Chase et al. 2006"/><ref name="Davis et al. 2004"/><ref name="Graham et al. 2006"/><ref name="Hilu et al. 2003"/><ref name="Pires et al. 2006"/><ref name="Rudall et al. 1997a"/><ref name="Källersjö et al. 1998"/><ref name="Fay et al. 2000"/><ref name="Soltis et al. 2000"/><ref name="Stevenson et al. 2000"/><ref name="McPherson and Graham 2001"/><ref name="Goldblatt 1995"/><ref name="Stevenson and Loconte 1995"/><ref name="Vinnersten and Bremer 2001"/><ref name="Givnish 2005"/><ref name="Givnish 2006"/><ref name="Rønsted 2005"/><ref name="Zomlefer 2001"/>
-Together with other analyses within each of these two orders has allowed the redistribution of the original genera of Liliaceae ''sensu lato'' into a variety of families, as illustrated in the two cladograms of the Asparagales and Liliales shown here.<ref name="Soltis et al. 2005"/> These changes have resulted in considerable changes in the positioning of Liliaceae within the overall classification, as shown in the table below, as well as within the family.
+Together with other analyses within each of these two orders has allowed the redistribution of the original genera of Liliaceae ''sensu lato'' into a variety of families, as illustrated in the two cladograms of the Asparagales and Liliales shown here.<ref name="Soltis et al. 2005">{{cite book |last=Soltis |first=D.E. |last2=Soltis |first2=P.F. |last3=Endress |first3=P.K. |last4=Chase |first4=M.W. |title=Phylogeny and evolution of angiosperms |year=2005 |publisher=Sinauer |location=Sunderland, MA }}</ref> These changes have resulted in considerable changes in the positioning of Liliaceae within the overall classification, as shown in the table below, as well as within the family.
 {| class="wikitable" border="1"
 |+ <big>Table 1: Evolution of placement of Liliaceae in different taxonomic schemes</big><ref name=Singh2/>
-! style="border-right:solid black 2px;  background: #ccf"|Rank !! style="border-right:solid black 2px;  background: #ccf"|[[Bentham & Hooker system|Bentham and Hooker]] (1883)<ref name=BandH/>!! style="border-right:solid black 2px;  background: #ccf"|[[Cronquist system|Cronquist]] (1981)<ref name="Cronquist 1981"/>!! style="border-right:solid black 2px;  background: #ccf"|[[Takhtajan system|Takhtajan]] (1980)<ref name=Takh09/> !! style="border-right:solid black 2px;  background: #ccf"|[[Dahlgren system|Dahlgren]] (1985)<ref name="Dahlgren et al. 1985"/>!! style="border-right:solid black 2px;  background: #ccf"|[[Thorne system|Thorne]] (1992)<ref name="Thorne 1992"/> !! style= "background: #ccf"|[[Angiosperm Phylogeny Group|APG]] (2003-9)<ref name="APG II 2003"/><ref name="APG III 2009"/>
+! style="border-right:solid black 2px;  background: #ccf"|Rank !! style="border-right:solid black 2px;  background: #ccf"|[[Bentham & Hooker system|Bentham and Hooker]] (1883)<ref name=BandH/>!! style="border-right:solid black 2px;  background: #ccf"|[[Cronquist system|Cronquist]] (1981)<ref name="Cronquist 1981"/>!! style="border-right:solid black 2px;  background: #ccf"|[[Takhtajan system|Takhtajan]] (1980)<ref name=Takh09>{{cite book |last=Takhtadzhi︠a︡n |first=Armen Leonovich |title=Flowering Plants |year=2009 |publisher=Springer |isbn=1-4020-9609-7 |url=http://books.google.ca/books?id=oumyfO-NHuUC&pg=PA634&lpg=PA634 |accessdate=7 January 2014}}</ref> !! style="border-right:solid black 2px;  background: #ccf"|[[Dahlgren system|Dahlgren]] (1985)<ref name="Dahlgren et al. 1985"/>!! style="border-right:solid black 2px;  background: #ccf"|[[Thorne system|Thorne]] (1992)<ref name="Thorne 1992"/> !! style= "background: #ccf"|[[Angiosperm Phylogeny Group|APG]] (2003-9)<ref name="APG II 2003"/><ref name="APG III 2009"/>
 |-
 | style="border-right:solid black 2px; background:#F2CEEC"  |'''[[Phylum|Division]]''' ||style="border-right:solid black 2px"| ||style="border-right:solid black 2px"|[[Magnoliophyta]]||style="border-right:solid black 2px"|Magnoliophyta||style="border-right:solid black 2px"  | ||style="border-right:solid black 2px"  | ||
@@ Line 40: / Line 41: @@
 |}
-For a comparison of the classifications of Genera from 1959 ([[John Hutchinson (botanist)|Hutchinson]])<ref name=Hutchinson/> to 2000 (Wilson and Morrison)<ref name="Wilson and Morrison"/> see Table 1 in Fay et al. 2006.<ref name="Fay et al. 2006"/>
+For a comparison of the classifications of Genera from 1959 ([[John Hutchinson (botanist)|Hutchinson]])<ref name=Hutchinson>{{cite book|last=Hutchinson|first=John|title=The families of flowering plants, arranged according to a new system based on their probable phylogeny. 2 vols|year=1959|publisher=Macmillan}}</ref> to 2000 (Wilson and Morrison)<ref name="Wilson and Morrison">{{cite book|editor1-last= Wilson|editor1-first= K. L. |editor2-last=Morrison |editor2-first=D. A.|title=Monocots: Systematics and evolution (Proceedings of the Second International Conference on the Comparative Biology of the Monocotyledons, Sydney, Australia 1998)|year=2000|publisher=[[CSIRO]]|location=Collingwood, Australia|url=http://www.publish.csiro.au/pid/2424.htm|isbn=0-643-06437-0|accessdate=14 January 2014 }}</ref> see Table 1 in Fay et al. 2006.<ref name="Fay et al. 2006">{{cite journal|last=Fay|first=M. F.|coauthors=Chase, M. W., Ronsted, N., Devey, D. S., Pillon, Y., Pires, J. C., Petersen, G., Seberg, O., and Davis, J. I.
+|year=2006|title=Phylogenetics of Liliales: summarized evidence from combined analyses of five plastid and one mitochondrial loci.|pages=559–565|url=
+http://ninaronsted.dk/page1/files/page1_6.pdf|accessdate=18 January 2014}} In {{Harvtxt|Columbus|Friar|Porter|Prince|2006}}</ref>
 == Cladistic analysis of Liliales and families within former Liliaceae ==
@@ Line 129: / Line 132: @@
 == Phylogeny ==
-[[Sequencing]] of the [[rbcL|''rbc''L]] and [[Maturase K|''mat''K]] regions of the [[chloroplast]] [[DNA]] of ''Lilium'' and related genera<ref name="Hayashi and Kawano 2000"/> confirmed the [[Circumscription (taxonomy)|circumscription]] of the family ''s.s.'' as described by [[Minoru N. Tamura|Tamura]] (1998).<ref name="Tamura 1998b"/> Chloroplast [[ndhF|''ndh''F]] gene sequencing also supports Liliaceae monophyly, reuniting the Liliaceae and Calochortaceae ''sensu'' Tamura.<ref name="Patterson 2002"/><ref name="Fay et al. 2006"/>
+[[Sequencing]] of the [[rbcL|''rbc''L]] and [[Maturase K|''mat''K]] regions of the [[chloroplast]] [[DNA]] of ''Lilium'' and related genera<ref name="Hayashi and Kawano 2000">{{cite journal|author =Hayashi, K.|coauthor=Kawano, S.|year=2000|title=Molecular systematics of ''Lilium'' and allied genera (Liliaceae): Phylogenetic relationships among ''Lilium'' and related genera based on the rbcL and matK gene sequence data.|journal= Plant Species Biology|volume=15|issue=|pages=73–79|url=http://www.researchgate.net/publication/228911730__systematics_of_Lilium_and_allied_genera_%28Liliaceae%29_phylogenetic_relationships_among_Lilium_and_related_genera_based_on_the_rbcL_and_matK_gene_sequence_data|accessdate= 14 January 2013}}</ref> confirmed the [[Circumscription (taxonomy)|circumscription]] of the family ''s.s.'' as described by [[Minoru N. Tamura|Tamura]] (1998).<ref name="Tamura 1998b">{{cite book| last =Tamura| first =M. N.|  year =1998|  pages =343–353| chapter =Liliaceae|url=http://link.springer.com/chapter/10.1007%2F978-3-662-03533-7_41}} In {{Harvtxt|Kubitzki|1998}}. [http://books.google.ca/books?id=FyPVYzL76sMC&pg=PA164 additional except]</ref> Chloroplast [[ndhF|''ndh''F]] gene sequencing also supports Liliaceae monophyly, reuniting the Liliaceae and Calochortaceae ''sensu'' Tamura.<ref name="Patterson 2002"/><ref name="Fay et al. 2006"/>
-Despite establishing this relative degree of monophyly for the Liliaceae family,<ref name="Chase et al. 1995a"/><ref name="Chase et al. 1995b"/> morphology remains diverse<ref name="Tamura 1998b"/> and there exists within the Liliaceae clade, a number of subclades. Particularly enigmatic were ''Clintonia, Medeola, Scoliopus, and Tricyrtis''. ''Clintonia'' (which has a disjunct distribution involviong both East Asia and North America) and the closely related ''Medeola''  form one of those subclades, and are now considered a separate tribe (Medeoleae), within the Lilioideae although at different times they have been both a separate subfamily (Medeoloideae) and family (Medeolaceae). Sequencing of the ''rbc''L and ''mat''K chloroplast genes established monophyly for ''Clintonia'', but with separate clades corresponding to the two distribution regions.<ref name="Hayashi 2001"/>
+Despite establishing this relative degree of monophyly for the Liliaceae family,<ref name="Chase et al. 1995a"/><ref name="Chase et al. 1995b">{{cite book |last=Chase |first=M. W. |last2=Stevenson |first2=D. W. |last3=Wilkin |first3=P. |last4=Rudall |first4=P. J. |chapter=Monocotyledons: systematics and evolution |year=1995 |publisher=Royal Botanic Gardens |location=Kew | pages=685–730 |chapter=Monocot systematics: A combined analysis}} In {{Harvtxt|Rudall|Cribb|Cutler|Humphries|1995}}.</ref> morphology remains diverse<ref name="Tamura 1998b"/> and there exists within the Liliaceae clade, a number of subclades. Particularly enigmatic were ''Clintonia, Medeola, Scoliopus, and Tricyrtis''. ''Clintonia'' (which has a disjunct distribution involviong both East Asia and North America) and the closely related ''Medeola''  form one of those subclades, and are now considered a separate tribe (Medeoleae), within the Lilioideae although at different times they have been both a separate subfamily (Medeoloideae) and family (Medeolaceae). Sequencing of the ''rbc''L and ''mat''K chloroplast genes established monophyly for ''Clintonia'', but with separate clades corresponding to the two distribution regions.<ref name="Hayashi 2001">{{cite journal|last = Hayashi|first=Kazuhiko|coauthors= Seiji Yoshida, Frederick H. Utech, Shoichi Kawano|year=2001|title=Molecular systematics in the genus Clintonia and related taxa based on rbcL and matK gene sequence data|journal=Plant Species Biology|volume=16|issue=2|pages=119–137|doi=10.1046/j.1442-1984.2001.00057.x|url=http://www.blackwell-synergy.com/doi/abs/10.1046/j.1442-1984.2001.00057.x|accessdate= 18 January 2014}}</ref>
-While the ten genera of the Lilioideae subfamily are characterised by contractile bulbs and roots, and a Fritillaria-type embryo-sac (megagametophyte with four megaspores), the five genera constituting the Streptopoideae and Calochortoideae subfamilies form another distinct group, previously characterised under the Calochortoideae alone. They are characterised by creeping rhizomes, styles which are divided at their apices, and by megagametophyte development of the Polygonum-type (a simple megaspore and triploid endosperm) embryo-sac. At times these genera were considered in a separate family (Calochortaceae) (e.g. Tamura) or even the more heterogeneous [[Uvulariaceae]] ''sensu'' Dahlgren. However most of the latter had low morphological similarity to the Liliaceae and ''[[Uvularia]]'' and ''[[Disporum]]'' are now located in the [[Colchicaceae]]. ''Disporum '' contained both Asian and North American species which had always been distinguishable. Following molecular analysis the North American species were restored to the genus ''Prosartes'' and retained in Liliaceae, and are now included in the Streptopoideae. The Asian species were moved to Colchicaceae.<ref name="Shinwari et al. 1994"/>
+While the ten genera of the Lilioideae subfamily are characterised by contractile bulbs and roots, and a Fritillaria-type embryo-sac (megagametophyte with four megaspores), the five genera constituting the Streptopoideae and Calochortoideae subfamilies form another distinct group, previously characterised under the Calochortoideae alone. They are characterised by creeping rhizomes, styles which are divided at their apices, and by megagametophyte development of the Polygonum-type (a simple megaspore and triploid endosperm) embryo-sac. At times these genera were considered in a separate family (Calochortaceae) (e.g. Tamura) or even the more heterogeneous [[Uvulariaceae]] ''sensu'' Dahlgren. However most of the latter had low morphological similarity to the Liliaceae and ''[[Uvularia]]'' and ''[[Disporum]]'' are now located in the [[Colchicaceae]]. ''Disporum '' contained both Asian and North American species which had always been distinguishable. Following molecular analysis the North American species were restored to the genus ''Prosartes'' and retained in Liliaceae, and are now included in the Streptopoideae. The Asian species were moved to Colchicaceae.<ref name="Shinwari et al. 1994">{{cite journal|last =Shinwari|first= Z. K.|coauthors=Terauchi, R., Hteck, F. H., and Kawano, S.|year=1994|title=Recognition of the New World ''Disporum'' section ''Prosartes'' (Liliaceae) based on the sequence data of the ''rbcL'' gene.|journal=Taxon|volume=43|issue=3|pages=353–366|url=http://links.jstor.org/sici?sici=0040-0262(199408)43%3A3%3C353%3AROTNWD%3E2.0.CO%3B2-U|doi=10.2307/1222713| accessdate= 18 January 2014}}</ref>
 == Evolution ==
-The development of a phylogenetic approach to taxonomy, starting with [[Charles Bessey]]'s  with ''[[Bessey system|The phylogenetic taxonomy of flowering plants]]'' (1915) suggested the Liliales formed some of the earliest monocots.<ref name=Bessey/> Molecular analysis suggests that Liliaceae arose in [[Eurasia]], around 68&nbsp;million years ago during the late ([[Maastrichtian]]) [[Late Cretaceous|Cretaceous]] to early ([[Paleocene]]) [[Paleogene]] eras<ref name=Leitch/> with the development of two main evolutionary clades. The first of these, characterised as Lilileae (''Lilium, Fritillaris, Nomocharis), Cardiocrinum), Notholirion'') from the [[Himalayas]], and the second, the Tulipae (''Erythronium, Tulipa''), (''Gagea'')) from [[East Asia]]. On the other hand ''Clintonia-Medeola'' may have appeared in North America but was subsequently dispersed, as may have the Calochortaceae ''sensu'' Tamura. Divergence amongst the Liliales probably occurred around 36&nbsp;million years ago, with the Liliaceae (''sensu'' Tamura) emerging at 27&nbsp;million years, Liliaceae ''s.s.'' at 20&nbsp;million, Lilieae 12&nbsp;million and ''Calochortus'' 7&nbsp;million.<ref name="Patterson 2002"/> Liliaceae [[fossils]] have been dated to the Paleogene<ref name=Stillwell/> and Cretaceous<ref name=Goin/> eras in the [[Antarctic]].
+The development of a phylogenetic approach to taxonomy, starting with [[Charles Bessey]]'s  with ''[[Bessey system|The phylogenetic taxonomy of flowering plants]]'' (1915) suggested the Liliales formed some of the earliest monocots.<ref name=Bessey/> Molecular analysis suggests that Liliaceae arose in [[Eurasia]], around 68&nbsp;million years ago during the late ([[Maastrichtian]]) [[Late Cretaceous|Cretaceous]] to early ([[Paleocene]]) [[Paleogene]] eras<ref name=Leitch>{{cite journal|last=Leitch|first=I. J.|coauthors=IJ, Beaulieu JM, Cheung K, Hanson L, Lysak MA, Fay MF|title=Punctuated genome size evolution in Liliaceae|journal=Journal of evolutionary biology|date=November 2007|volume=20|issue=6|pages=2296–308|pmid=17956392|url=http://www.unboundmedicine.com/medline/citation/17956392/Punctuated_genome_size_evolution_in_Liliaceae_|accessdate=26 January 2014}}</ref> with the development of two main evolutionary clades. The first of these, characterised as Lilileae (''Lilium, Fritillaris, Nomocharis), Cardiocrinum), Notholirion'') from the [[Himalayas]], and the second, the Tulipae (''Erythronium, Tulipa''), (''Gagea'')) from [[East Asia]]. On the other hand ''Clintonia-Medeola'' may have appeared in North America but was subsequently dispersed, as may have the Calochortaceae ''sensu'' Tamura. Divergence amongst the Liliales probably occurred around 36&nbsp;million years ago, with the Liliaceae (''sensu'' Tamura) emerging at 27&nbsp;million years, Liliaceae ''s.s.'' at 20&nbsp;million, Lilieae 12&nbsp;million and ''Calochortus'' 7&nbsp;million.<ref name="Patterson 2002"/> Liliaceae [[fossils]] have been dated to the Paleogene<ref name=Stillwell>{{cite book|title=Paleobiology and Paleoenvironments of Eocene Rocks: McMurdo Sound, East Antarctica|year=2000|publisher=American Geophysical Union|isbn=0-87590-947-7|page=162|url=http://books.google.ca/books?id=vNdk7dBWdKkC&pg=pa162&lpg=pa162#v=onepage&q&f=false|accessdate=28 January 2014|editor=Jeffrey D. Stilwell, Rodney M. Feldmann}}</ref> and Cretaceous<ref name=Goin>{{cite book|last=Goin|first=Francisco|title=Late Cretaceous/Paleogene West Antarctica Terrestrial Biota and its Intercontinental Affinities|year=2012|publisher=Springer|isbn=94-007-5491-4|url=http://books.google.ca/books?id=D0jDXAWyylMC&pg=PA58&lpg=PA58|coauthors=Carolina Acosta Hospitaleche, Tania Dutra, Sergio Marenssi|accessdate=28 January 2014}}</ref> eras in the [[Antarctic]].
 Core Liliales probably arose as shade plants, with subsequent evolution of Liliaceae s.s. and ''Calochortus'' to open areas including deciduous forest in the more open autumnal period, but then a return of some species (e.g. ''Cardiocrinum''). This was accompanied by a shift from rhizomes to bulbs, 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'') <ref name="Patterson 2002"/>
@@ Line 147: / Line 150: @@
 === Suprageneric subdivisions ===
-Due to the diversity of the originally broadly defined Liliaceae ''s.l.'', many attempts have been made to form suprageneric classifications, e.g. [[subfamilies]] and [[Tribe (biology)|tribes]].<ref name="Peruzzi 2009"/> By 1813, Candolle recognised five subdivisions which he called tribes ([[Asparagaceae|Asparagées]], [[Trilliaceae|Trilliacées]], [[Asphodeloideae|Asphodelées]], [[Bromeliaceae|Bromeliées]], [[Tulipa]]cées),<ref name=Candolle1/> all of which Jussieu had made separate families, with the exception of ''Tulipa'', which was a genus within the Liliaceae. By 1845 John Lindley observed that the family had become extremely diverse, ill defined and unstable not only by its overall circumscription, but also by its subdivisions. Of the 133 genera he included, he described eleven ''suborders''.<ref name=Lindley2/>
+Due to the diversity of the originally broadly defined Liliaceae ''s.l.'', many attempts have been made to form suprageneric classifications, e.g. [[subfamilies]] and [[Tribe (biology)|tribes]].<ref name="Peruzzi 2009">{{cite journal |last=Peruzzi |first=L. |last2=Leitch |first2=I.J. |last3=Caparelli |first3=K.F.  |title=Chromosome diversity and evolution in Liliaceae |journal=[[Annals of Botany]] |year=2009 |volume=103 |issue=3 |pages=459–475 |doi=10.1093/aob/mcn230 |url=http://aob.oxfordjournals.org/content/103/3/459.full |accessdate=8 January 2014}}</ref> By 1813, Candolle recognised five subdivisions which he called tribes ([[Asparagaceae|Asparagées]], [[Trilliaceae|Trilliacées]], [[Asphodeloideae|Asphodelées]], [[Bromeliaceae|Bromeliées]], [[Tulipa]]cées),<ref name=Candolle1/> all of which Jussieu had made separate families, with the exception of ''Tulipa'', which was a genus within the Liliaceae. By 1845 John Lindley observed that the family had become extremely diverse, ill defined and unstable not only by its overall circumscription, but also by its subdivisions. Of the 133 genera he included, he described eleven ''suborders''.<ref name=Lindley2/>
-In 1879 a revision of the North American Liliaceae described sixteen tribes<ref name=WatsonS/> and [[Bentham and Hooker]] twenty tribes in 1883, while [[Adolf Engler|Engler]] and [[Karl Anton Eugen Prantl|Prantl]] in their [[Engler system|extensive description]] of the Liliaceae in 1899 identified 31 tribes distributed over 11 subfamilies.<ref name = "Engler and Prantl"/> In 1936 Buxbaum undertook a major revision of the Liliaceae and ''inter alia'' described the subfamily [[Lilioideae]], with three tribes: [[Lloydieae]], [[Tulipeae]] and [[Lilieae]].<ref name=buxbaum1/><ref name=buxbaum2/><ref name="Vinnersten and Reeves"/> Peruzzi et al. (2009) discuss the complex rearrangement of the various genera, tribes and subfamilies over a 30&nbsp;year period (Peruzzzi et al. Table1) from 1985,<ref name="Peruzzi 2009"/> partly summarised here in Table 2.
+In 1879 a revision of the North American Liliaceae described sixteen tribes<ref name=WatsonS>{{cite journal |last=Watson |first=Sereno |title=Revision of the North American Liliaceae: Descriptions of Some New Species of North American Plants |year=1879 |volume=XIV |pages=213–312 |url=http://books.google.ca/books?id=N8EwAQAAMAAJ |accessdate=6 January 2014 |journal=Proceedings of the American Academy of Arts and Sciences}}</ref> and [[Bentham and Hooker]] twenty tribes in 1883, while [[Adolf Engler|Engler]] and [[Karl Anton Eugen Prantl|Prantl]] in their [[Engler system|extensive description]] of the Liliaceae in 1899 identified 31 tribes distributed over 11 subfamilies.<ref name = "Engler and Prantl">{{cite book|last=Engler|first=Adolf|title=Die Natürlichen Pflanzenfamilien nebst ihren Gattungen und wichtigeren Arten, insbesondere den Nutzpflanzen, unter Mitwirkung zahlreicher hervorragender Fachgelehrten. II(5)|year=1899|publisher=W. Engelmann|location=Leipzig|url=http://www.biodiversitylibrary.org/item/56456#page/672/mode/1up|editor=Engler, Adolf and Prantl, Karl|pages = 10–91|chapter = Liliaceae|accessdate=6 February 2014}}</ref> In 1936 Buxbaum undertook a major revision of the Liliaceae and ''inter alia'' described the subfamily [[Lilioideae]], with three tribes: [[Lloydieae]], [[Tulipeae]] and [[Lilieae]].<ref name=buxbaum1>{{cite journal |first=F. |last=Buxbaum |title=Die Entwicklungslinien der Lilioideae. I. Die systematische Stellung der gattung ''Gagea'' |journal=Botanisches Archiv |volume=38 |year=1936 |pages=213–293 |language=German}}</ref><ref name=buxbaum2>{{cite journal |first=F. |last=Buxbaum |title=Die Entwicklungslinien der Lilioideae. II. Die Wurmbaeoideae |journal=Botanisches Archiv |volume=38 |year=1937 |pages=305–398 |language=German}}</ref><ref name="Vinnersten and Reeves">{{cite journal|title=Phylogenetic relationships within Colchicaceae|journal=American Journal of Botany|date=October|volume=90|issue=10): 1455–1462. 2003.|pages=1455–1462. 2003.|doi=10.3732/ajb.90.10.1455|url=http://www.amjbot.org/content/90/10/1455.full|accessdate=5 February 2014|year=2003}}</ref> Peruzzi et al. (2009) discuss the complex rearrangement of the various genera, tribes and subfamilies over a 30&nbsp;year period (Peruzzzi et al. Table1) from 1985,<ref name="Peruzzi 2009"/> partly summarised here in Table 2.
-Classifications published since the use of [[Molecular phylogenetics|molecular methods in phylogenetics]] have taken a narrower view of the Liliaceae. In 1998 Tamura considered ''[[Calochortus]]'' sufficiently distinct to elevate the subfamily [[Calochortoideae]] to family status as Calochortaceae.<ref name="Tamura 1998b"/><ref name="Tamura 1998C"/> In 2009, Takhtajan used a very narrow definition (see the table below).<ref name=Takh09/> {{As of|2014|January}} the [[Angiosperm Phylogeny Website]] (APweb)<ref name="Stevens 2001"/> included four of Takhtajan's families in its Liliaceae, recognizing three subfamilies, one of which is divided into two tribes.<ref name=Mabberley/>
+Classifications published since the use of [[Molecular phylogenetics|molecular methods in phylogenetics]] have taken a narrower view of the Liliaceae. In 1998 Tamura considered ''[[Calochortus]]'' sufficiently distinct to elevate the subfamily [[Calochortoideae]] to family status as Calochortaceae.<ref name="Tamura 1998b"/><ref name="Tamura 1998C">{{cite book| last =Tamura| first =M. N.|  year =1998|  pages =164–172| chapter =Calochortaceae|url=http://books.google.ca/books?id=FyPVYzL76sMC&pg=PA164#v=onepage&q&f=false}} In {{Harvtxt|Kubitzki|1998}}. [http://link.springer.com/chapter/10.1007/978-3-662-03533-7_22 additional excerpt]</ref> In 2009, Takhtajan used a very narrow definition (see the table below).<ref name=Takh09/> {{As of|2014|January}} the [[Angiosperm Phylogeny Website]] (APweb)<ref name = "Stevens 2001">{{cite web |last=Stevens |first=P. F. |title=Liliaceae |url=http://www.mobot.org/MOBOT/Research/APweb/orders/lilialesweb.htm#Liliaceae |work=Angiosperm Phylogeny Website. Version 12, September 28, 2013 |accessdate=2 January 2014 |year=2001 onwards}}</ref> included four of Takhtajan's families in its Liliaceae, recognizing three subfamilies, one of which is divided into two tribes.<ref name=Mabberley/>
 {| class="wikitable" border="1"
 |+ <big>Table 2: Comparison of Four 21st Century Classifications of the Liliaceae</big>
-! colspan=3  style="border-right:solid black 2px;  background: #ccf"|Tamura<ref name="Tamura 1998b"/>  !! colspan=2  style="border-right:solid black 2px" | Takhtajan<ref name=Takh09/> !! colspan=3  style="border-right:solid black 2px;  background: #ccf" | Taxonomicon<ref name=brands/> !! colspan=3 | APweb<ref name=Mabberley/>
+! colspan=3  style="border-right:solid black 2px;  background: #ccf"|Tamura<ref name="Tamura 1998b"/>  !! colspan=2  style="border-right:solid black 2px" | Takhtajan<ref name=Takh09/> !! colspan=3  style="border-right:solid black 2px;  background: #ccf" | Taxonomicon<ref name=brands>{{cite web |last=Brands |first=S.J. |title=The Taxonomicon |url=http://taxonomicon.taxonomy.nl/TaxonTree.aspx?id=10608 |publisher=Universal Taxonomic Services, Zwaag, The Netherlands |accessdate=8 January 2014 |year=1989 onwards}}</ref> !! colspan=3 | APweb<ref name=Mabberley/>
 |-
 ! style="background: #ccf;"|Family !! style="background: #ccf;"|Subfamily !! style="border-right:solid black 2px;  background: #ccf"| Tribe!! Family !! style="border-right:solid black 2px"| Tribe !! style="background: #ccf;"|Family !! style="background: #ccf;"|Subfamily !! style="border-right:solid black 2px;  background: #ccf"| Tribe !! Family !! Subfamily !! Tribe
@@ Line 344: / Line 347: @@
 ==== Modern classification ====
-Various authorities (e.g. [[ITIS]] 16,<ref name=ITIS_Liliaceae/> [[Germplasm Resources Information Network|GRIN]] 27,<ref name=GRIN_Liliaceae/> [[World Checklist of Selected Plant Families|WCSP]],<ref name=WCSP_Liliaceae/> [[National Center for Biotechnology Information|NCBI]],<ref name=NCBI/> DELTA<ref name=WatsonL/> ) differ on the exact number of genera included in Liliaceae ''s.s.'', but generally there are about fifteen to sixteen genera, depending on whether or not ''[[Amana (genus)|Amana]]'' is included in ''[[Tulipa]]'' and ''Lloydia'' in ''[[Gagea]]''. For instance ''Amana'' is still listed separately in WCSP.
+Various authorities (e.g. [[ITIS]] 16,<ref name=ITIS_Liliaceae>{{cite web |last=Interagency Taxonomic Information System ([[ITIS]]) |title=Liliaceae |url=http://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=42633&source=from_print |accessdate=6 January 2014}}</ref> [[Germplasm Resources Information Network|GRIN]] 27,<ref name=GRIN_Liliaceae>{{cite web |last=Germplasm Resources Information Network ([[Germplasm Resources Information Network|GRIN]]) |title=Liliaceae |url=http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?652 |accessdate=6 January 2014}}</ref> [[World Checklist of Selected Plant Families|WCSP]],<ref name=WCSP_Liliaceae>{{Citation|title=Liliaceae |work=[[World Checklist of Selected Plant Families]] |publisher=[[Royal Botanic Gardens, Kew]] |url=http://apps.kew.org/wcsp/qsearch.do?plantName=Liliaceae |accessdate=14 January 2014}} In {{Harvtxt|WCSP|2001}}.</ref> [[National Center for Biotechnology Information|NCBI]],<ref name=NCBI>{{cite web|title=Liliaceae|url=http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?id=4677|publisher=National Center for Biotechnology Information|accessdate=22 January 2014}}</ref> DELTA<ref name=WatsonL>{{cite web |last=Watson |first=L. |last2=Dallwitz |first2=M.J. |title=The families of flowering plants: descriptions, illustrations, identification, and information retrieval |url=http://delta-intkey.com/angio/www/liliacea.htm |accessdate=5 January 2014 |date=1992 onwards |date=19 October 2013|publisher=DELTA – DEscription Language for TAxonomy}}</ref> ) differ on the exact number of genera included in Liliaceae ''s.s.'', but generally there are about fifteen to sixteen genera, depending on whether or not ''[[Amana (genus)|Amana]]'' is included in ''[[Tulipa]]'' and ''Lloydia'' in ''[[Gagea]]''. For instance ''Amana'' is still listed separately in WCSP.
 Currently the APWeb lists fifteen genera, arranged as follows:<ref name = "Stevens 2001"/>{{refn|Botanical authorities from WCSP.<ref name="WCSP_Liliaceae"/>}}