Taxonomy of Liliaceae

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Taxonomy of Liliaceae
Temporal range: 68–0 
Ma
 Late Cretaceous - Recent
Lilium candidum
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Monocots
Order: Liliales
Family:
Juss.[1]
Type genus
Lilium
Type species
Lilium candidum
Subfamilies and tribes[2]
tribe:
Medeoleae
tribe: Lilieae
Diversity
About 600 species

The

phylogenetic
principles, has it been possible to identify the many separate taxonomic groupings within the original family and redistribute them, leaving a relatively small core as the modern family Liliaceae, with fifteen genera and 600 species.

The Liliaceae emerged from the

Medeoleae arose in North America but was subsequently dispersed, as may have the Streptopoideae and Calochortoideae. Liliaceae fossils have been dated to the Paleogene and Cretaceous eras in the Antarctic
.

The Liliaceae probably arose as shade plants, with subsequent evolution to open areas including

bulbs
, to more showy flowers, the production of capsular fruit and narrower parallel-veined leaves.

While the suprageneric (above genus level) structure of the family has varied greatly with its ever-changing circumscription, as currently constituted the family consists of three

Medeoleae and Lilieae
. The three subfamilies contain fifteen genera and approximately 600 species in all.

History

Pre-Darwinian

The

calix" (perianth) of six equal coloured parts, six stamens, a superior ovary, single style, and trilocular capsule. The term ordo at that time was closer to what we now understand as family, rather than order.[9][10] Although Jussieu used the Latin "lilia" in his Genera Plantarum, elsewhere he used the French "liliacées",[11] as had Adanson. The word "Liliaceae" was soon widely used by botanists such as Samuel Frederick Gray,[12] John Lindley,[13] and Pierre-Joseph Redouté[14]
in the early nineteenth century.

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 equal stamens, a single style, a simple

Phanérogames (Phenogamae in Gray), meaning "visible seed", hence Endogenæ phanerogamæ.[18] Candolle also instituted the concept of ordered ranks, based on classes, subclasses, familles (Latin: ordines naturales) and tribus (tribes),[10]
subdividing the Liliaceae.

Lindley was the first post-Linnaean English

phyletic.[24][25][26]

Post-Darwinian

Liliiflorae.[28][29][30][31][32] In the English literature, Charles Bessey (1915) followed Adolf 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 subclass of monocots, the Strobiloideae,[33] while from John Hutchinson (1959) onwards the Liliaceae were treated as part of the Liliales (see Table 1
).

Over time the Liliaceae became increasingly broadly, and somewhat arbitrarily, defined as all species of plants with six tepals and a superior ovary. They eventually came to encompass about 300 genera and 4,500 species, within the

petaloid monocots, characterised by showy flowers with tepals and without starch in the endosperm
.

Deconstructing Liliaceae

Other botanists in the twentieth century echoed Lindley's concerns about the lack of a clearly defined grouping for Liliaceae. The earliest of these was

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.[40][47]

The 1990s saw considerable progress in plant

Minoro Tamura (sensu Tamura)[53] and (iii) Clintonia-Medeola which had been included in Liliaceae sensu Tamura[53] (see Table 3).[54]

This newly, more narrowly (

Medeoleae) Medeoloideae sensu Tamura (see Table 3).[2]

Modern classification of Liliaceae

To meet the need for a thorough revision of the taxonomy of the flowering plants (

systematists formed the Angiosperm Phylogeny Group (APG), resulting in a new classification published in 1998.[55] The scheme was largely based on the work of Kåre Bremer and colleagues at Uppsala and Stockholm universities in the late 1970s,[56][57][58] and became universally accessible on the internet in 1996.[59]
It was an ordinal system, concentrating on orders rather than families, prioritising
sensu lato
, s.l.).

These studies of

18S rDNA, the plastid gene rbcL, and other DNA sequences.[64][65][66][67][68][69][70][71][72][73][74][75][76][77][78][79][80][81][82]

These studies, together with other analyses within each of these two orders, allowed the redistribution of the original genera of Liliaceae s.l. into a variety of families across the Liliales and Asparagales, as illustrated in Cladogram 1.[83] This redistribution resulted in considerable changes both in the suprafamilial positioning of Liliaceae within the overall APG classification (as shown in Table 1 below), as well as the subfamilial structure (see Suprageneric subdivisions).[84]

Table 1: Evolution of placement of Liliaceae in different taxonomic schemes[27][85]
Rank Bentham and Hooker (1883)[23] Cronquist (1981)[34] Takhtajan (1966, 1980, 2009)[86][41][87] Dahlgren (1977, 1985)[88][35] Thorne (1992–2007)[38][89] APG (2003–9)[60][50]
Division
Magnoliophyta
 Magnoliophyta Magnoliophyta
Class
Monocotyledons
 Liliopsida Liliopsida (
monocots
) - unranked
Angiospermae
)		 (
monocots
) - unranked
Subclass
 Liliidae Liliidae Liliidae Liliidae
Superorder
(Series)		 Coronarieæ (Liliiflorae) Lilianae[90] (Lilianae)
Liliiflorae
 Lilianae Lilianae
Order Liliales Liliales Liliales Liliales Liliales
For a comparison of the classifications of genera from 1959 (Hutchinson)[91] to 2000 (Wilson and Morrison),[92] see Table 1 in Fay et al. 2006,[93] Table 1 in Peruzzi et al. 2009[94] and Table 3.

Phylogeny

The synthesis of molecular data with

rbcL and trnL-F plastid genes revealed four main Liliales lineages:[63]

  1. Liliaceae group:
    Calochortaceae), Philesiaceae and Smilacaceae
    ;
  2. Campynemataceae;
  3. Colchicaceae group (Colchicoid lilies): Colchicaceae (including Petermannia and Uvularia), Alstroemeriaceae and Luzuriaga;
  4. Melanthiaceae (including Trilliaceae).

The original family Liliaceae in the broad sense (

photosynthesising) Corsiaceae were added to Liliales by APG III in 2009.[50][95]

rbcL and matK chloroplast genes of Lilium and related genera[96] confirmed the circumscription of the family in the sensu stricto usage of Tamura (1998).[97] Chloroplast ndhF gene sequencing also supported Liliaceae monophyly, reuniting the Liliaceae sensu Tamura and Calochortaceae sensu Tamura (see Table 2 & Table 3
below).

Cladogram I: Phylogeny of Liliaceae and related families
Liliales

Corsiaceae

Campynemataceae

MelanthiaceaeL

Ripogonaceae
L

PhilesiaceaeL

SmilacaceaeL

Liliaceae

sensu stricto

Asparagales

Orchidaceae

BoryaceaeL

Blandfordiaceae
L

Lanariaceae
L

Ixioliriaceae
A

Tecophilaeaceae

Doryanthaceae

Iridaceae

Xeronemataceae
L

Xanthorrhoeaceae
L

Amaryllidaceae

Aphyllanthaceae
L

Themidaceae
L

Hyacinthaceae
L

Agavaceae
L/A

Laxmanniaceae
L

AsparagaceaeL

Ruscaceae
L

commelinids

  1. Molecular phylogenetic relationships for Liliales and Asparagales, showing relationship between Liliaceae s.s. and families formerly placed in Liliaceae s.l.L or AmaryllidaceaeA (itself sometimes treated as part of Liliaceae),[a] and families formerly placed in other orders.[63][2]
  2. APG III (2009) absorbed Luzuriagaceae into Alstroemeriaceae and added Corsiaceae.[50][95][98]

Evolution and biogeography

The major

sister clades (crown node) occurring later at around 55–52 mya.[b] divergence within the Liliaceae appeared at about 36–34 mya, within Liliaceae sensu Tamura (Lilioideae s.l.) at 27 mya, Liliaceae sensu Dahlgren (Lilieae s.s. and Tulipeae) at 20 mya (Miocene
).

Within the Liliaceae sensu Dahlgren there developed two main evolutionary subclades (see Cladogram II and Table 3). The first of these, characterised as the Lilieae s.s. (Lilium, Fritillaria, Nomocharis), Cardiocrinum), Notholirion) diverged around 12 mya. The second subclade was the Tulipeae (Erythronium, Tulipa), (Gagea). Divergence within Calochortus is dated to 7 mya. This places the emergence of the Liliaceae at approximately the last (Maastrichtian period) of the Late Cretaceous periods (72 to 66 mya) to early (Paleocene) Paleogene periods (66 to 23 mya), formerly the Cretaceous–Tertiary boundary, 65 mya.[78][54][102]

The southern hemisphere intercontinental distributions of Liliales suggests a connection to

vicariance (separation of populations by continental division) split between Medeoleae and Lilieae involving Eurasia and North America (Cladogram II).[78]

Cladogram II: Evolution of Liliaceae
146–100 
Monocots
 — 124

(other)

Liliales  117–82 (Gondwana)

Campynemataceae

Melanthiaceae

Alstroemeriaceae + Luzuriagaceae + Colchicaceae

68–65 Smilacaceae+Liliaceae 55–52

Smilacaceae

Liliaceae sensu APG 36–34
Calochortaceae
 sensu Tamura<br />(North America)
Liliaceae sensu Tamura 27<br />=Lilioideae s.l.

Medeoleae
(North America
)

Lilaceae sensu Dahlgren 20<br />=Lilieae s.s. + Tulipeae<br />(Eurasia)
Lilieae s.s. 12 (Himalayas)

Lilium etc.

Tulipeae (East Asia)

Tulipa
etc.

Biogeographical origins and dates in mya[101][78][54]

Liliaceae sensu Dahlgren arose in

Medeoleae) may have appeared in North America but subsequently underwent intercontinental dispersal (although some evidence points to a Eurasian origin). The Calochortaceae sensu Tamura (Streptopoideae and Calochortoideae) appears to have evolved in western North America, with subsequent colonisation of East Asia by Streptopus and the ancestral Tricyrtis.[54]

Liliaceae probably arose as

convergence in similar habitats. [54]

The fossil record of Liliales is relatively poor,

fossils have been dated to the Paleogene[104] and Cretaceous periods in the Antarctic.[105][106][107]

Characteristics

Main article: Liliaceae § Description

Liliaceae

The diversity of characteristics complicates description of Liliaceae morphology, and confused taxonomic classification for centuries. The diversity is also of considerable evolutionary significance (see

trichomes (root hairs) and contractile roots.[108] The flowers may be arranged along the stem, developing from the base, or as a single flower at the tip of the stem, or as a cluster of flowers. They contain both male (androecium) and female (gynoecium) characteristics and are symmetric radially, but sometimes as a mirror image. Most flowers are large and colourful, except for Medeoleae. Both the petals and sepals are usually similar and appear as two concentric groups (whorls) of "petals", that are often striped or multi-coloured, and produce nectar at their bases. The stamens are usually in two groups of three (trimerous) and the pollen has a single groove (monosulcate). The ovary is superior, i.e. placed above the attachment of the other parts. There are three fused carpels (syncarpous) with one to three chambers (locules
), a single style and a three-lobed stigma. The embryo sac is of the Fritillaria type. The fruit is generally a wind-dispersed capsule, but occasionally a berry (Medeoleae) which is dispersed by animals. The leaves are generally simple and elongated with veins parallel to the edges, arranged singly and alternating on the stem, but may form a rosette at the base of the stem.

Subclades

(See

mycorrhizae.[54]

The five genera constituting the Streptopoideae and Calochortoideae subfamilies form another distinct group, previously characterised under the Calochortoideae alone. These have creeping rhizomes, styles divided at their apices, and an embryo-sac of the Polygonum-type with a simple megaspore and triploid endosperm. At times, these genera were considered as a separate family (Calochortaceae; e.g. Tamura) or even placed in the more heterogeneous Uvulariaceae sensu Dahlgren. However most of the latter had low morphological similarity to the Liliaceae, and Uvularia and Disporum are now classified 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, subfamily Streptopoideae, while the Asian species were moved to Colchicaceae.[54][110][111]

Liliaceae: Morphological characteristics
  • Lilium: Perigonium of six undifferentiated tepals, in two trimerous whorls and side-connected (dorsifixed) anthers
    Lilium: Perigonium of six undifferentiated tepals, in two trimerous whorls and side-connected (dorsifixed) anthers
  • Calochortus nuttallii: Tepals in two clearly distinguished whorls of three sepals and three petals
    Calochortus nuttallii: Tepals in two clearly distinguished whorls of three sepals and three petals
  • Erythronium revolutum: Three stigmata and pseudo-basifixed anthers surrounding filament tip
    Erythronium revolutum: Three stigmata and pseudo-basifixed anthers surrounding filament tip
  • Lilium: Longitudinal dehiscence of anthers
    Lilium: Longitudinal dehiscence of anthers
  • Tulipa humilis: Multiple connate (fused) carpels surrounded by stamens
    Tulipa humilis: Multiple connate (fused) carpels surrounded by stamens
  • Fritillaria pudica: Trilocular fruit
    Fritillaria pudica: Trilocular fruit

Subdivisions

Suprageneric subdivisions

Due to the diversity of the originally broadly defined Liliaceae (s.l.), many attempts have been made to form supageneric classification systems, organizing the genera into

Tulipacées),[17] 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. For the 133 genera he included, he described eleven suborders.[20] By the 1870s, as Baker describes in his revision of the family, the taxonomy of Liliaceae had become vast and complicated. His approach was to divide the family into eight tribes.[112][113]

In 1879, a revision of the North American Liliaceae by

Korolkowia, Fritillaria, Notholirion, Cardiocrinum, Nomocharis and Lilium).[116][117][118] Hutchinson included most of these genera within the tribe Tulipeae.[91] The complex rearrangements of the various genera, tribes and subfamilies over a 30-year period from 1985, discussed by Peruzzi and colleagues (2009),[94] are partly summarised in Table 2
below.

Classifications published since the use of

Takhtajan used an even narrower definition (see Table 2 & Table 3 below).[87]

Despite now having established a taxonomic grouping for the family Liliaceae that is genetically monophyletic, compared to the prior longstanding

polyphyletic assemblages under this name,[51][119] the morphology remains diverse,[97] and there exists within the Liliaceae clade a number of subclades. There appeared to be two major clades, the first and largest of these consisted of three subclades: Clintonia—Medeola—Gagea, Lilium Fritillaria—Notholirion, and Tulipa—Erythronium. The second smaller clade, Streptopus—Tricyrtis contained elements of Dahlgren's Uvulariaceae. The position of Calochortus remained problematic, being considered a sister clade to Liliaceae, as treated by Tamura, but further analysis suggested it was in fact sister to Tricyrtis, although it is now considered separate once again (see Table 3).[63][54][93][120][121]

Also enigmatic were Clintonia, Medeola, Scoliopus, and Tricyrtis. Clintonia, with a disjunct distribution involving East Asia and North America, and the closely related Medeola form a subclades and are now considered a separate tribe (Medeoleae) within the Lilioideae, although at different times they have been considered a separate subfamily (Medeoloideae) or family (Medeolaceae). Sequencing of the rbcL and matK chloroplast genes established monophyly for Clintonia, but with separate clades corresponding to the two areas of distribution.[122] The Angiosperm Phylogeny Website (APWeb)[2] includes four of Takhtajan's families in Liliaceae, recognizing three subfamilies, one of which is divided into two tribes and referred to as Liliaceae sensu APG III.[123]

Table 2: Comparison of Four 21st Century System Subdivisions of Liliaceae
Tamura[97] Takhtajan[87] Taxonomicon[124] APWeb[123][2]
Family Subfamily Tribe Family Tribe Family Subfamily Tribe Family Subfamily Tribe
Liliaceae Lilioideae s.s. Lilieae s.s. Liliaceae Lloydieae Liliaceae Lilioideae s.l. Lloydieae Liliaceae Lilioideae s.l. Lilieae s.l.
Lilieae Lilieae
Tulipeae Tulipeae Tulipeae
Medeoloideae Medeolaceae Medeola, Clintonia Medeoleae
Calochortaceae Calochorteae Scoliopaceae Calochortoideae Calochortoideae
Tricyrtideae Tricyrtidaceae Streptopoideae Streptopoideae

Genera

Main article: List of Liliaceae genera

Historical treatment

Historically, the inclusion of genera within Liliaceae has been extremely broad. Of the various published systems, one of the best known and also the broadest modern circumscription is the

ITIS, together with their disposition as APG III transfers to other families and subfamilies, within Liliales and three other orders, Alismatales, Asparagales and Dioscoreales. Current members of Liliaceae are shown in bold.[125]

List of disposition of genera previously included in Liliaceae
Genus Authority Family Subfamily Order
List of genera included in Liliaceae by Cronquist
Abama=Narthecium
Adans.
Agapanthus
L'Her.
 Amaryllidaceae Agapanthoideae Asparagales
Aletris L. Nartheciaceae Dioscoreales
Allium L. Amaryllidaceae Allioideae Asparagales
Alstroemeria L. Alstroemeriaceae Liliales
Amaryllis L. Amaryllidaceae Amaryllidoideae Asparagales
Amianthium Gray Melanthiaceae Liliales
Androstephium
Torr.
 Asparagaceae Brodiaeoideae Asparagales
Anthericum L. Asparagaceae Agavoideae Asparagales
Asparagus L. Asparagaceae Asparagoideae Asparagales
Asphodelus L.
Xanthorrhoeaceae
 Asphodeloideae Asparagales
Astelia
R. Br.
 Asteliaceae Asparagales
Atamosco= Zephyranthes Adans.
Bloomeria Kellogg Asparagaceae Brodiaeoideae Asparagales
Brodiaea
Sm.
 Asparagaceae Brodiaeoideae Asparagales
Calochortus
Pursh
 Liliaceae Liliales
Camassia
Lindl.
 Asparagaceae Agavoideae Asparagales
Cardiocrinum (
Endl.
) Lindl.		 Liliaceae Liliales
Chamaelirium
Willd.
 Melanthiaceae Liliales
Chionodoxa
Boiss.
 Asparagaceae Scilloideae Asparagales
Chlorogalum
Kunth
 Asparagaceae Agavoideae Asparagales
Chlorophytum
Ker-Gawl.
 Asparagaceae Agavoideae Asparagales
Clintonia Raf. Liliaceae Liliales
Colchicum L. Colchicaceae Liliales
Convallaria L. Asparagaceae Nolinoideae Asparagales
Cooperia Herb. 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[c]
D. Don
 Colchicaceae Liliales
Dracaena L. Asparagaceae Nolinoideae Asparagales
Echeandia
Ortega
 Asparagaceae Agavoideae Asparagales
Eremocrinum
M.E.Jones
 Asparagaceae Agavoideae Asparagales
Erythronium L. Liliaceae Liliales
Eucharis
Lind.
 Amaryllidaceae Amaryllidoideae Asparagales
Fritillaria L. Liliaceae Liliales
Gagea Salisb. Liliaceae Liliales
Galanthus L. Amaryllidaceae Amaryllidoideae Asparagales
Gloriosa L. Colchicaceae Liliales
Habranthus
 Herb. Amaryllidaceae Amaryllidoideae Asparagales
Harperocallis McDaniel Tofieldiaceae Alismatales
Hastingsia
S. Wats.
 Asparagaceae Agavoideae Asparagales
Helonias L. Melanthiaceae Asparagales
Hemerocallis
 L. Xanthorrhoeaceae Hemerocallidoideae Asparagales
Hesperanthes= Echeandia S. Wats.
Hesperocallis Gray Tofieldiaceae Alismatales
Hesperoscordum= Triteleia Lindl.
Hippeastrum Herb. Amaryllidaceae Amaryllidoideae Asparagales
Hosta Tratt. Asparagaceae Agavoideae Asparagales
Hyacinthoides
Medik.
 Asparagaceae Scilloideae Asparagales
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. Amaryllidaceae Amaryllidoideae Asparagales
Lilium L. Liliaceae Liliales
Liriope
Lour.
 Asparagaceae Nolinoideae Asparagales
Lloydia= Gagea Salisb. ex
Reichenb.
Lophiola Ker-Gawl. Nartheciaceae Dioscoreales
Lycoris Herb. Amaryllidaceae Amaryllidoideae Asparagales
Maianthemum
G.H. Weber ex Wiggers
 Asparagaceae Nolinoideae Asparagales
Medeola L. Liliaceae Liliales
Melanthium L. Melanthiaceae Asparagales
Milla
Cav.
 Asparagaceae Brodiaeoideae Asparagales
Muilla S. Wats. ex
Benth.
 Asparagaceae Brodiaeoideae Asparagales
Muscari
P. Mill.
 Asparagaceae Scilloideae Asparagales
Narcissus L. Amaryllidaceae Amaryllidoideae Asparagales
Narthecium
Huds.
 Nartheciaceae Dioscoreales
Nolina
Michx.
 Asparagaceae Nolinoideae Asparagales
Nomocharis
Franch.
 Liliaceae Liliales
Notholirion
Wall.
ex Boiss.		 Liliaceae Liliales
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
Polygonatum P. Mill. Asparagaceae Nolinoideae Asparagales
Sansevieria
Thunb.
 Asparagaceae Nolinoideae Asparagales
Schoenocaulon Gray Melanthiaceae Liliales
Schoenolirion Torr. ex Dur. Asparagaceae Agavoideae Asparagales
Scilla
 L. Asparagaceae Scilloideae Asparagales
Scoliopus Torr. Liliaceae Liliales
Smilacina=Maianthemum
Desf.
Stenanthium (Gray) Kunth Melanthiaceae LIliales
Sternbergia
Kit.
 Amaryllidaceae Amaryllidoideae Asparagales
Streptopus Michx. Liliaceae Liliales
Tofieldia Huds. Tofieldiaceae Alismatales
Tricyrtis Wall. Liliaceae Liliales
Trillium L. Melanthiaceae Liliales
Tristagma
Poepp.
 Amaryllidaceae Allioideae Asparagales
Triteleia
Dougl.
ex Lindl.		 Asparagaceae Brodiaeoideae Asparagales
Triteleiopsis Hoover Asparagaceae Brodiaeoideae Asparagales
Tulipa L. Liliaceae Liliales
Urginea
 Steinh. Asparagaceae Scilloideae Asparagales
Uvularia L. Colchicaceae Liliales
Vagnera=Maianthemum Adans.
Veratrum L. Melanthiaceae Liliales
Xerophyllum Michx. Melanthiaceae Liliales
Zephyranthes Herb. Amaryllidaceae Amaryllidoideae Asparagales
Zigadenus Michx. Melanthiaceae Liliales
List of other genera historically included in Liliaceae s.l.
Albuca L. Asparagaceae Scilloideae Asparagales
Anticlea Kunth Melanthiaceae Liliales
Arthropodium R.Br. Asparagaceae Lomandroideae Asparagales
Bellevalia
Lapeyr.
 Asparagaceae Scilloideae Asparagales
Bomarea
Mirb.
 Alstroemeriaceae Liliales
Chionographis
Maxim.
 Melanthiaceae Liliales
Conanthera
Ruiz & Pav.
 Tecophilaeaceae Asparagales
Cyclobothra= Calochortus D.Don
Daiswa
 Raf. Melanthiaceae Liliales
Drimia
Jacq.
 Asparagaceae Scilloideae Asparagales
Drimiopsis Lindl. & Paxton Asparagaceae Scilloideae Asparagales
Eremurus
M.Bieb.
 Xanthorrhoeaceae Asphodeloideae Asparagales
Eriospermum Jacq. Asparagaceae Nolinoideae Asparagales
Eucrosia Ker Gawl. Amaryllidaceae Amaryllidoideae Asparagales
Eustephia Cav. Amaryllidaceae Amaryllidoideae Asparagales
Griffinia Ker Gawl. Amaryllidaceae Amaryllidoideae Asparagales
Heloniopsis
A.Gray
 Melanthiaceae Liliales
Hemiphylacus S.Wats. Asparagaceae Asparagoideae Asparagales
Herreria Ruiz & Pav. Asparagaceae Agavoideae Asparagales
Hessea Herb. Amaryllidaceae Amaryllidoideae Asparagales
Hookera= Brodiaea Salisb.
Hypoxis L. 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= Hippeastrum Rusby
Metanarthecium Maxim. Nartheciaceae Dioscoreales
Miersia Lindl. Amaryllidaceae Allioideae Asparagales
Molineria
 Colla Hypoxidaceae Asparagales
Nietneria
Klotzsch
 Melanthiaceae Liliales
Paradisea Mazzuc. Anthericaceae Asparagales
Paris L. Melanthiaceae Liliales
Peliosanthes Andrews Asparagaceae Nolinoideae Asparagales
Phaedranassa Herb. Amaryllidaceae Amaryllidoideae Asparagales
Selonia= Eremurus
Regel
Stropholirion= Dichelostemma Torr.
Theropogon Maxim. Amaryllidaceae Allioideae Asparagales
Tovaria= Maianthemum
Neck.
 Asparagaceae Nolinoideae Asparagales
Toxicoscordion
Rydb.
 Melanthiaceae Liliales
Tracyanthus= Stenanthium Small
Tulbaghia
Heist.
 Amaryllidaceae Allioideae Asparagales
Tupistra Ker Gawl. Asparagaceae Nolinoideae Asparagales
Urceolina Rchb. Amaryllidaceae Amaryllidoideae Asparagales
Wurmbea Thunb. Colchicaceae Liliales

The more modern phylogenetically based treatment of the genera, including the major systems of the 1980s of Dahlgren and Tamura, are shown in Table 3.

Table 3: Historical distribution of Liliaceae sensu APWeb/APG ("core Liliales") genera[123][2] by family and author with subsequent subfamilial divisions
Genera sensu
Dahlgren (1985)
(tribes)[35]		 sensu
Tamura (1998)
subfamilies
(tribes)[97][53]		 sensu
APG		 Tribes Subfamilies
Streptopus Uvulariaceae
(Uvularieae)
Calochortaceae

(Tricyrtideae)		 Liliaceae5 Streptopoideae
syn. Scoliopaceae Takht.
Scoliopus Trilliaceae1 Calochortaceae
(Tricyrtideae)		 Liliaceae
Prosartes Uvulariaceae
(Uvularieae)		 Calochortaceae
(Tricyrtideae)		 Liliaceae
Calochortus Calochortaceae Calochortaceae
(Calochorteae)		 Liliaceae
Dumort., Compsoaceae Horan.
, nom. illeg.,
Tricyrtidaceae Takht., nom. cons.
Tricyrtis6 Uvulariaceae
(Tricyrtideae)		 Calochortaceae
(Tricyrtideae)		 Liliaceae
Medeola Trilliaceae1 Liliaceae
Medeolioideae		 Liliaceae
Benth.
 Lilioideae s.l.3
Clintonia Uvulariaceae
(Uvularieae)		 Liliaceae
Medeolioideae		 Liliaceae
Cardiocrinum Liliaceae Liliaceae
Lilioideae
(Lilieae)		 Liliaceae Lilieae s.s. (Tamura) Lilieae s.l.
syn. Lilioideae s.s. (Tamura),3 Erythroniaceae
Borkh.
, Tulipaceae Borkh.
Notholirion Liliaceae Liliaceae
Lilioideae
(Liliae)		 Liliaceae
Nomocharis2 Liliaceae Liliaceae
Lilioideae
(Lilieae)		 Liliaceae
Fritillaria Liliaceae Liliaceae
Lilioideae
(Lilieae)		 Liliaceae
Lilium Liliaceae Liliaceae
Lilioideae
(Lilieae)		 Liliaceae
Gagea Liliaceae Liliaceae
Lilioideae
(Tulipeae)		 Liliaceae Lloydieae Tulipeae s.l. (Tamura)4
Lloydia2 Liliaceae Liliaceae
Lilioideae
(Tulipeae)		 Liliaceae
Amana2 Liliaceae Liliaceae
Lilioideae
(Tulipeae)		 Liliaceae Tulipeae s.s.
Tulipa Liliaceae Liliaceae
Lilioideae
(Tulipeae)		 Liliaceae
Erythronium Liliaceae Liliaceae
Lilioideae
(Tulipeae)		 Liliaceae
  1. Dahlgren expressed some uncertainty as to whether to include these genera in Trilliaceae or Uvulariaceae, tribus Uvularieae[35]
  2. Some authorities embed Nomocharis within Lilium,[126][81] and Lloydia within Gagea.[127] Amana had been embedded in Tulipa but was subsequently restored as a separate genus.[128][129]
  3. In Tamura's classification Lilioideae is used s.s. (Tulipeae s.l. and Lilieae s.s.), whereas in APWeb it is used s.l. to also include Medeoleae. Thus Lilioideae s.s. is equivalent to Lilieae s.l.
  4. While Tulipae s.l. is embedded in Lilieae s.l. in APWeb, many authors support it as a separate taxon[120][121][130]
  5. "Core Liliales" corresponds to Liliaceae sensu APG, incorporating all genera shown here[54]
  6. Tricyrtis is probably not a sister clade to Calochortus, and may represent a further subfamily separate from Calochortoideae[120][121]

Modern subfamilial divisions within Liliaceae

The evolutionary and phylogenetic relationships between the genera currently included in Liliaceae are shown in Cladogram III.

Cladogram III:
Phylogeny and biogeography
of the genera of the Liliaceae
Liliaceae
West NA

Calochortus

Lilioideae* East NA EA
Medeoleae
 East NA
Lilieae s.l. EA
Tulipeae East Asia

Lloydia
)

Lilieae s.s. Himalayas
Phylogenetic tree reflecting relationships based on molecular phylogenetic evidence.[54][78][94][102][95][131][120][121]
*=Liliaceae sensu Tamura; EA=Eurasia NA=North America

The largest genera are Gagea (200),

Tulipa and Lloydia in Gagea
. For instance Amana is still listed separately in WCSP.

The exact subdivision of Liliaceae differs between authors. In 2002 Patterson and Givnish identified two major clades corresponding to Tamura's Calochortaceae and Liliaceae,[97] but preferred to retain his original division into two separate families rather than the overarching "core Liliales" (Liliaceae sensu APG). Within Liliaceae sensu Tamura they confirmed his decision to include Medeola-Clintonia as a separate subfamily, Medeolioideae, with the remaining genera as subfamily Lilioideae. Liliodeae was then divided into two tribes, Lilieae and Tulipeae (Tulipa, Erythronium, Gagea, Lloydia). Within Calochortaceae sensu Tamura, they proposed erecting a second subfamily, Streptopoideae (Prosartes, Scoliopus, Streptopus), with the remaining genera in subfamily Calochortoideae.[54] Subsequent work by Rønsted et al. (2005)[81] and by Fay et al. (2006) confirmed the overall phylogenetic relationships of Patterson and Givnish and their subdivisions, and further elucidated the position of Gagea within the tribe Tulipae, but the latter authors restored the broader circumscription of Liliaceae sensu APG .[93] In 2013, Kim et al. proposed further subdivision, placing the two genera of Calochortoideae (Calochortus and Tricyrtis) into subfamilies of their own and splitting off Gagea from the rest of Tulipeae by resurrecting the tribe Lloydieae.[120][121] (see Table 3)

The best known schema, the

APWeb, lists fifteen genera, arranged as follows, and illustrated in Table 4, with three subfamilies, Lilioideae representing Liliaceae sensu Tamura and the two subfamilies of Calochortaceae sensu Tamura (Streptopoideae and Calochortoideae) as proposed by Patterson and Givnish now included within Lilaceae sensu APG.[2]

Table 4: APWeb/APG Distribution of Subfamilies, Tribes and Genera of Liliaceae[2]
with illustration of morphological diversity
Subfamily Tribe Genus
Lilioideae Eaton
Medeoleae Benth.
Clintonia borealis
Clintonia borealis
 Clintonia Raf. - bead lilies
Gronov. ex L.
- Indian cucumber-root
Ritgen
Cardiocrinum giganteum
Cardiocrinum giganteum

Gagea lutea
Gagea lutea1,2

Erythronium sibiricum
Erythronium sibiricum1
Lindl.
- giant lilies
Tourn.
ex L. – fritillary or mission bells
Salisb. (including Lloydia Salisb. ex Rchb.
)
– yellow star-of-Bethlehem1,2
Lilium Tourn. ex L. – lily
Franch.
Boiss.
Tulipa L. (including Amana Honda
) – tulip1
Erythronium L. – trout lily1
Calochortoideae Dumort.3
Calochortus catalinae
Calochortus catalinae

Tricyrtis hirta
Tricyrtis hirta
Pursh
- mariposa, globe lilies
Wall.
– toad lily
Streptopoideae
Prosartes hookeri
Prosartes hookeri
D.Don
– drops of gold
Torr.
– Fetid Adder's Tongue
Michx.
– twistedstalk
  1. Some classifications place Tulipa, Erythronium and Gagea into a separate tribe, Tulipeae with the remaining genera in Lilieae s.s.[94][120][121]
  2. Other authorities place Gagea within its own tribe, Lloydieae[120][121]
  3. The situation with respect to Calochortoideae remains uncertain. Originally Calochortus and Tricyrtis were considered to be sister clades and placed together in subfamily Calochortoideae.
    Further study has not confirmed this (see Cladogram III, below) and it has been proposed that Tricyrtis be placed in a separate subfamily.[120][121]

Etymology

The name Liliaceae was coined by Michel Adanson in 1763.[6] The name was derived from Lilium and the family suffix -aceae. Lilium is the type genus of the family, which is the Latin for Lily, which in turn came from the Greek name for it, λείριον (leírion).[138][139]

See also

Notes

  1. ^ Agavaceae has been included in both families at various times
  2. ^ Janssen and Bremer (2004) have somewhat earlier dates of 91 and 80 mya respectively but consider these similar within methodological limits[101]
  3. ^ Asian species. N American species restored as Prosartes D. Don

References

  1. ^ a b Jussieu 1789, Lilia pp. 48–49.
  2. ^ a b c d e f g h Stevens 2015, Liliaceae.
  3. ^ Linnaeus 1753, Lilium vol. i p. 302.
  4. ^ Linnaeus 1753, Hexandria monogynia vol. i pp.  285–352.
  5. ^ Lobstein 2013.
  6. ^ a b Adanson 1763, VIII Liliaceae p. 42–60.
  7. ^ Stevens 2013.
  8. ^ Adanson 1763, Liliaceae Section II Les Lis Lilia p. 48.
  9. ^ ICN 2012, 18.2 Names of families and subfamilies, tribes and subtribes.
  10. ^ a b Candolle 1813, Des familles et des tribus pp. 192–195.
  11. ^ de Jussieu 1778, p. 228.
  12. ^ Gray 1821, Fam IX Liliaceae Jussieu. vol 2: p. 173.
  13. ^ Lindley 1830, CCLI: Liliaceae. p. 279.
  14. ^ Redouté 1802–1816.
  15. ^ Gray 1821, An Arrangement of the Families, and of the anomalous Genera of phenogamous Plants, according to their sexual Organs. vol 2: p. vi.
  16. ^ 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. vol 1: p. xx.
  17. ^ a b Candolle 1813, Esquisse. D'une Série linéaire et par conséquent artificielle, pour la disposition des familles naturelles du règne végetal p. 219.
  18. ^ Marilaun 1890–1891.
  19. ^ Lindley 1830.
  20. ^ a b Lindley 1846, Order LXII: Liliaceae - Lilyworts. pp. 200–205.
  21. ^ Kubitzki, Rudall & Chase 1998.
  22. ^ Lindley 1846, p .201.
  23. ^ a b Bentham & Hooker 1862–1883, Vol III Part II Liliaceae pp. 748–836.
  24. ^ a b Stuessy 2009, Natural classification p. 47.
  25. ^ Datta 1988, Systems of classification p. 21.
  26. ^ Stace 1989, The development of plant taxonomy p. 17.
  27. ^ a b Singh 2004, Historical background of plant classification. pp. 10–29.
  28. ^ Eichler 1886, Liliiflorae p. 34.
  29. ^ Engler 1900–1968.
  30. ^ Engler 1903, Liliiflorae p. 93.
  31. ^ Carter 1913.
  32. ^ Wettstein 1924, Liliiflorae p. 862.
  33. ^ a b Bessey 1915.
  34. ^ a b c Cronquist 1981.
  35. ^ a b c d e Rasmussen 1985.
  36. ^ Lawrence 1951.
  37. ^ a b Judd et al. 2007.
  38. ^ a b c Thorne 1992.
  39. ^ Reveal 1997.
  40. ^ a b Datta 1988.
  41. ^ a b Takhtajan 1980.
  42. ^ Lotsy 1907–1911, 3: Cormophyta Siphonogamia. Liliifloren p. 715.
  43. ^ Huber 1969.
  44. ^ Meerow 2002.
  45. ^ Walters & Keil 1996, Liliaceae p. 418.
  46. ^ Kelch 2002.
  47. ^ Mathew 1989.
  48. ^ Chase et al 1993.
  49. ^ Utech 2003.
  50. ^ a b c d e APG III 2009.
  51. ^ a b Chase et al. 1995a.
  52. ^ Rasmussen 1985, Liliaceae pp.233–238.
  53. ^ a b c d Tamura 1998b.
  54. ^ a b c d e f g h i j k l m n o p q Patterson & Givnish 2002.
  55. ^ a b c APG 1998.
  56. ^ Bremer & Wanntorp 1978.
  57. ^ Bremer, Bremer & Thulin 1997.
  58. ^ Williams et al. 2010.
  59. ^ Bremer & Bremer 2005.
  60. ^ a b APG II 2003.
  61. ^ Chase & Reveal 2009.
  62. ^ LAPGIII 2009.
  63. ^ a b c d Rudall et al. 2000.
  64. ^ Chase et al. 2000.
  65. ^ Chase et al. 2006.
  66. ^ Davis et al. 2004.
  67. ^ Graham et al. 2006.
  68. ^ Hilu et al. 2003.
  69. ^ Pires et al. 2006.
  70. ^ Rudall et al. 1997a.
  71. ^ Källersjö et al. 1998.
  72. ^ Fay 2000.
  73. ^ Soltis et al. 2000.
  74. ^ Stevenson et al. 2000.
  75. ^ McPherson & Graham 2001.
  76. ^ Goldblatt 1995.
  77. ^ Stevenson & Loconte 1995.
  78. ^ a b c d e f g h Vinnersten & Bremer 2001.
  79. ^ Givnish et al. 2005.
  80. ^ Givnish et al. 2006.
  81. ^ a b c Rønsted et al. 2005.
  82. ^ Zomlefer et al. 2001.
  83. ^ Soltis et al. 2005.
  84. ^ Reveal 2010.
  85. ^ Jeffrey 1982, p. 123.
  86. ^ Takhtajan 1966.
  87. ^ a b c Takhtadzhi︠a︡n 2009, Liliaceae p. 634 .
  88. ^ Dahlgren 1977.
  89. ^ Thorne & Reveal 2007.
  90. ^ Naikh 1984, Post-Darwinian systems of classification, p. 111, at Google Books.
  91. ^ a b Hutchinson 1959.
  92. ^ Wilson & Morrison 2000.
  93. ^ a b c Fay et al. 2006.
  94. ^ a b c d e Peruzzi, Leitch & Caparelli 2009.
  95. ^ a b c d Meerow 2012.
  96. ^ Hayashi & Kawano 2000.
  97. ^ a b c d e f g Tamura 1998a.
  98. ^ Leadlay & Jury 2006.
  99. ^ Bremer 2000.
  100. ^ Wikström, Savolainen & Chase 2001.
  101. ^ a b c d Janssen & Bremer 2004.
  102. ^ a b c Leitch et al. 2007.
  103. ^ Wilkin & Mayo 2013, p. 43.
  104. ^ Stilwell & Feldmann 2000, p. 162.
  105. ^ Goin et al. 2012, West Antarctic Paleoflora p. 58.
  106. ^ Sambamurty 2005, p. 172.
  107. ^ Friis et al. 2011.
  108. ^ Spichiger & Perret 2002.
  109. ^ Gao et al. 2012.
  110. ^ Shinwari et al. 1994.
  111. ^ Simpson 2011.
  112. ^ Baker 1871.
  113. ^ Baker 1873.
  114. ^ Watson 1879.
  115. ^ Engler & Prantl 1888, Engler Liliaceae pp. 10–91 .
  116. ^ Buxbaum 1936.
  117. ^ Buxbaum 1937.
  118. ^ Vinnersten & Reeves 2003.
  119. ^ Chase et al. 1995b.
  120. ^ a b c d e f g h Kim et al. 2013a.
  121. ^ a b c d e f g h Kim et al. 2013b.
  122. ^ Hayashi et al. 2001.
  123. ^ a b c Mabberley 2013, Liliaceae p. 489.
  124. ^ Brands 2015, Liliaceae.
  125. ^ Chase et al. 2009.
  126. ^ Gao et al. 2011.
  127. ^ Peterson et al. 2008.
  128. ^ Tan et al. 2005.
  129. ^ Clennett et al. 2012.
  130. ^ Peruzzi 2016.
  131. ^ Stevens 2015, Liliales.
  132. ^ Tison et al. 2012.
  133. ^ ITIS 2015, Liliaceae.
  134. ^ GRIN 2015, Liliaceae.
  135. ^ WCLSPF 2015, Liliaceae.
  136. ^ NCBI 2015, Liliaceae.
  137. ^ Watson & Dallwitz 2015, Liliaceae Juss..
  138. ^ Gledhill 2006, Family names p. 16.
  139. ^ OED 2005.

Bibliography

Books

  • Kamenetsky, Rina; Okubo, Hiroshi, eds. (2012). Ornamental Geophytes: From Basic Science to Sustainable Production.
    ISBN 978-1-4398-4924-8
    .
  • Mabberley, David J (2013). Mabberley's Plant-Book (3 ed.).
    ISBN 978-1-107-78259-4
    . Retrieved 8 January 2014.
  • Redouté, P. J. (1802–1816). Les liliacées
    . Paris: Redouté. Retrieved 2 February 2014.

Paleobotany

Systematics and taxonomy

Taxonomic classifications (chronological)
Table of 58 families, Part II: Page 1
Table of 1615 genera, Part II: Page 8

Chapters

Articles

Phylogenetics

Phylogenetics: Angiosperms and monocots

Phylogenetics: Liliales

APG

Symposia

Websites

Databases

  • Media related to Liliaceae at Wikimedia Commons
  • The dictionary definition of Liliaceae at Wiktionary
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