Editing Liliaceae (section)

== Description ==
{{multiple image
| header    = Liliaceae floral morphology
| align     = left
| direction = vertical
| width     = 180

| image1    = Kievitsbloem.jpg
| caption1  = Section through flower of ''[[Fritillaria meleagris]]''

| image2    = Zanlophator1a.UME.jpg
| caption2  = ''[[Lilium]]'' flower with perigonium of six undifferentiated tepals, arranged in two trimerous whorls and side-connected (''dorsifixed'') anthers.

| image3    = Sego lily cm.jpg
| caption3  = Sego lily (''[[Calochortus nuttallii]]'') with tepals in two clearly distinguished whorls of three sepals and three petals.

| image4    = Tulip Tulipa clusiana 'Lady Jane' Rock Ledge Flower 2000px.jpg
| caption4  = ''[[Tulipa clusiana]]'' with three sepals resembling petals

| image5    = 2006-10-18Tricyrtis11.jpg
| caption5  = ''[[Tricyrtis]]'' flower with patterned tepals

| image6   = Lillium Stamens.jpg
| caption6 = ''[[Lilium longiflorum]]'': 1.&nbsp;[[Stigma (botany)|Stigma]], 2.&nbsp;[[Gynoecium|style]], 3.&nbsp;[[stamen]]s, 4.&nbsp;filament, 5.&nbsp;[[tepal]]

| image7    = Erythronium revolutum 8.jpg
| caption7  = ''[[Erythronium revolutum]]'' flower with three stigmata and pseudo-basifixed anthers surrounding the filament tip

| image8    = Anther dehiscence in Lilium.jpg
| caption8  = ''[[Lilium]]'' with longitudinal dehiscence of anthers

| image9    = Antera Lilium.jpg
| caption9  = ''Lilium'' anthers in cross section

| image10    = Lilium auratum - pollen.jpg
| caption10  = ''[[Lilium auratum]]'' pollen with typical single-grooved (monosulcate) pattern
   }}

{{multiple image
| align     = left
| direction = vertical
| width     = 180

| image1    = Tulipa aucheriana 240408 stamens and pistil.jpg
| caption1  = ''[[Tulipa humilis]]'' flower showing multiple connate (fused) carpels surrounded by stamens.

| image2   = Lilyfruit.jpg
| caption2 = ''Lillium'' fruit capsule
}}

The diversity of characteristics complicates any description of the Liliaceae morphology, and confused taxonomic classification for centuries. The diversity is also of considerable evolutionary significance, as some members emerged from shaded areas and adapted to a more open environment (see [[#Evolution|Evolution]]).<ref name="Patterson 2002"/>

=== General ===
The Liliaceae are characterised as monocotyledonous, perennial, herbaceous, bulbous (or [[rhizomatous]] in the case of [[Medeoleae]])<ref name="Tamura 1998b"/> flowering plants with simple [[trichomes]] (root hairs) and contractile [[root]]s.<ref name=Spichiger/> The flowers may be arranged (inflorescence) 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 placed above the attachment of the other parts (superior). There are three fused carpels (syncarpus) 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.

=== Specific ===
; [[Inflorescence]] : Usually [[Indeterminate growth|indeterminate]] (lacking terminal flower) as a [[raceme]] (''Lilium''); sometimes reduced to a single terminal flower (''Tulipa''). When [[pluriflor]] (multiple blooms), the flowers are arranged in a cluster or rarely are [[umbel|subumbellate]] (''Gagea'') or a [[thyrse]] (spike).<ref name=Simp11/>

; [[Flowers]] : [[Hermaphroditic]], [[actinomorphic]] (radially symmetric) or slightly zygomorphic (bilaterally symmetric),<ref name="Simpson2006">{{cite book|first=Michael G. |last=Simpson|title=Plant Systematics|url=https://books.google.com/books?id=Enuaxb9FymEC&pg=PA180|year=2006|publisher=Academic Press|isbn=978-0-12-644460-5|pages=180–}}</ref> [[pedicellate]] (on a short secondary stem), generally large and showy but may be inconspicuous : (Medeoleae). [[Bracts]] may (bracteate) or may not (ebracteate) be present. The [[perianth]] is undifferentiated ([[perigonium]]) and biseriate (two [[Whorl (botany)|whorled]]), formed from six [[tepals]] arranged into two separate whorls of three parts ([[Merosity|trimerous]]) each, although ''[[Scoliopus]]'' has only three petals, free from the other parts, but overlapping. The tepals are usually petaloid (petal like) and [[apotepalous]] (free) with lines (striate) or marks in other colors or shades. The [[perianth]] is either homochlamydeous (all [[tepals]] equal, e.g. ''[[Fritillaria]]'') or dichlamydeous (two separate and different whorls, e.g. ''[[Calochortus]]'') and may be united into a tube. [[Nectar]] is produced in perigonal nectaries at the base of the tepals.<ref name=Simp11/><ref name=Mabberley/><ref name=Singh1/><ref name=Weberling/>

; [[Androecium]] : Six [[stamens]]  in two trimerous whorls, with free filaments, usually epiphyllous (fused to tepals) and diplostemonous (outer whorl of stamens opposite outer tepals and the inner whorl opposite inner tepals), although ''Scoliopus'' has three stamens opposite the outer tepals.<ref name="Rudall et al. 2000a"/> The attachment of the anthers to the filaments may be either peltate (to the surface) or pseudo-basifixed (surrounding the filament tip, but not adnate, that is not fused) and [[dehisce]] longitudinally and are extrorse (dehiscing away from center). The [[pollen]] is usually monosulcate (single groove), but may be inaperturate (lacking aperture: ''Clintonia'', some ''Tulipa'' spp.) or operculate (lidded: ''Fritillaria'', some ''Tulipa'' spp.),<ref name=Furness/> and reticulate (net patterned: ''Erythronium'', ''Fritillaria'', ''Gagea'', ''Lilium'', ''Tulipa'').<ref name=Simp11/><ref name=Mabberley/><ref name="Rudall et al. 2000a"/>

; [[Gynoecium]] : Superior [[Ovary (botany)|ovary]] (hypogynous), syncarpous (with fused carpels), with three connate (fused) carpels and is trilocular (three [[locule]]s, or chambers) or unilocular (single locule, as in ''Scoliopus'' and ''Medeola''). There is a single style and a three lobed [[Stigma (botany)|stigma]] or three stigmata more or less elongated along the style. There are numerous anatropous (curved) [[ovules]] which display axile [[placentation]] (parietal in ''Scoliopus'' and ''Medeola''),<ref name="Rudall et al. 2000a"/> usually with an [[integument]] and thinner [[megasporangium]]. The embryo sac ([[megagametophyte]]) varies by genera, but is mainly [[tetraspore|tetrasporic]] (e.g. ''[[Fritillaria]]'').<ref name=Mabberley/> Embryo sacs in which three of the four [[megaspores]] fuse to form a [[triploid]] [[Nucleus (cell)|nucleus]], are referred to as ''Fritillaria-type'', a characteristic shared by all the core Liliales.<ref name="Patterson 2002"/><ref name=Simp11/><ref name=Mabberley/><ref name=Ownbey/>

; [[Fruit]] : A [[Capsule (botany)|capsule]] that is usually loculicidal (splitting along the [[locule]]s) as in the Lilioideae,<ref name="Patterson 2002"/> but occasionally septicidal (splitting between them, along the separating [[Septum|septa]]) in the Calachortoideae and wind dispersed, although the Medeoleae form [[berries]] (baccate).<ref name="Patterson 2002"/> The [[seeds]] may be flat, oblong, angular, [[wikt:discoid|discoid]], [[ellipsoid]] or [[globose]] (spherical), or compressed with a well developed [[Epidermis (botany)|epidermis]]. The exterior may be smooth or roughened, with a wing or [[raphe]] (ridge), [[aril]] or one to two tails, rarely hairy, but may be dull or shiny and the lack of a black [[integument]] distinguishes them from related taxa such as [[Allioideae]] that were previously included in this family, and striate (parallel longitudinally ridged) in the Steptopoideae. The [[Hilum (biology)|hilum]] (scar) is generally inconspicuous. The bitegmic (separate testa and tegmen) seed coat itself may be thin, suberose (like [[Cork (material)|cork]]), or crustaceous (hard or brittle). The [[endosperm]] is abundant, [[cartilaginous]] (fleshy) or horny and contains oils and [[aleurone]] but not [[starch]] (non-farinaceous). Its cells are [[polyploid]] ([[triploid]] or [[pentaploid]], depending on the [[embryo sac]] type). The [[Ovule|embryo]] is small (usually less than one quarter of seed volume), axile (radially sectioned), linear (longer than broad) or rarely [[Rudiment (biology)|rudimentary]] (tiny relative to [[endosperm]]) depending on placentation type, and straight, bent, curved or curled at the upper end.<ref name=Simp11/><ref name=Mabberley/><ref name=Kozlowski/><ref name=Baskin/>

; [[Leaves]] : [[Leaf shape|Simple]], entire (smooth and even), linear, oval to filiform (thread-like), mostly with parallel veins, but occasionally net-veined. They are alternate (single and alternating direction) and [[spiral]], but may be [[Whorl (botany)|whorled]] (three or more attached at one [[Node (botany)|node]], e.g. ''[[Lilium]], [[Fritillaria]]),'' cauline (arranged along the aerial [[Plant stem|stem]]) or sheathed in a basal [[Rosette (botany)|rosette]]. They are rarely [[Petiole (botany)|petiolate]] (stem attached before apex), and lack [[stipules]]. The aerial stem is unbranched.<ref name=Simp11/><ref name=Mabberley/><ref name=Singh1/><ref name=WatsonL/><ref name=Sharma/>

; [[Genome]] : The Liliaceae include a species with one of the largest [[genome size]] within the angiosperms, ''[[Fritillaria assyriaca]]'' ([[C-value|1C]]=127.4&nbsp;pg), while ''[[Tricyrtis macropoda]]'' is as small as 4.25&nbsp;pg.<ref name="Leitch et al 2007">{{cite journal |last1=Leitch |first1=I.J. |last2=Beaulieu |first2=J.M. |last3=Cheung |first3=K. |last4=Hanson |first4=L. |last5=Lysak |first5=M.A. |last6=Fay |first6=M.F. |author-link6=Michael Francis Fay |title=Punctuated genome size evolution in Liliaceae |journal=Journal of Evolutionary Biology |date=November 2007 |volume=20 |issue=6 |pages=2296–308 |pmid=17956392 |doi= 10.1111/j.1420-9101.2007.01416.x|doi-access=free }}</ref> [[Chromosome]] numbers vary by genus.<ref name=Mabberley/> Some genera like ''Calochortus'' (x=6-10), ''Prosartes'' (6,8,9,11), ''Scoliopus'' (7,8), ''Streptopus'' (8, 27) and ''Tricyrtis'' (12–13) have a small and variable number of chromosomes while subfamily Lilioideae have a larger and more stable chromosome number (12) as have the Medeoleae (7).<ref name="Tamura 1998b"/><ref name="Tamura 1995"/><ref name="Peruzzi 2009"/><ref name="Tamura 1998C"/>

; [[Phytochemistry]] : The seeds contain [[saponins]] but no [[calcium oxalate]] [[raphide]] crystals,<ref name="Rudall et al. 2000a"/> [[chelidonic acid]]<ref name="Patterson 2002"/> (unlike [[Asparagales]]) or [[cysteine]] derived [[sulphur]] compounds (allyl sulphides), another distinguishing feature from the characteristic [[Garlic breath|alliaceous odour]] of the [[Allioideae]]. ''[[Fritillaria]]'' in particular contains [[steroidal alkaloid]]s of the cevanine and [[solanum]] type.<ref name=Simp11/> [[Solanidine]] and solanthrene<ref name=solanthrene/> alkaloids have been isolated from some ''[[Fritillaria]]'' species.<ref name=Turner83/><ref name=Li2006/> ''[[Tulipa]] '' contains [[tulipanin]], an [[anthocyanin]]. (see also: [[#Toxicology|Toxicology]])

Characteristics often vary by habitat, between [[Shade tolerance|shade-dwelling]] genera (such as ''[[Prosartes]]'', ''[[Tricyrtis]]'', ''[[Cardiocrinum]]'', ''[[Clintonia]]'', ''[[Medeola virginiana|Medeola]]'', ''[[Prosartes]]'', and ''[[Scoliopus]]'') and sun loving genera. Shade-dwelling genera usually have broader leaves with smooth edges and net venation, and fleshy fruits ([[berries]]) with animal-dispersed seeds, [[rhizomes]], and small, inconspicuous flowers while genera native to sunny habitats usually have narrow, parallel-veined leaves, [[Capsule (botany)|capsular]] fruits with wind-dispersed seeds, bulbs, and large, visually conspicuous flowers.<ref name="Tamura 1998b"/> (See also ''[[#Evolution|Evolution]]'').