View source for Biology:Asteraceae

{{Hatnote|Not to be confused with Astraeaceae, a family of fungi.}}
{{Short description|Large family of flowering plants}}
{{Use list-defined references|date=November 2022}}
{{Automatic taxobox
|fossil_range = {{fossil range|76|0}}[[Earth:Campanian|Campanian]]<ref name=PNAS2015/>–recent
|image = Asteracea poster 3.jpg
|image_upright = 1.3
|image_alt = refer to caption
|image_caption = Twelve species of Asteraceae from the subfamilies Asteroideae, Carduoideae, and Cichorioideae
|taxon = Asteraceae
|authority = [[Biography:Friedrich von Berchtold|Bercht.]] & J.Presl<ref name=tropicos/>
|diversity = 1,911 genera
|diversity_link = List of Asteraceae genera
|diversity_ref = <ref name=PlantList/>
|type_genus = ''[[Biology:Aster (genus)|Aster]]''
|type_genus_authority = [[Biography:Carl Linnaeus|L.]]
|subdivision_ranks = Subfamilies
|subdivision =
* [[Biology:Asteroideae|Asteroideae]] <small>[[Biography:John Lindley|Lindl.]]</small>
* [[Biology:Barnadesioideae|Barnadesioideae]] <small>[[Biography:Kåre Bremer|K.Bremer]] & R.K.Jansen</small>
* [[Biology:Carduoideae|Carduoideae]] <small>Sweet</small>
* [[Biology:Cichorioideae|Cichorioideae]] <small>Chevall.</small>
* [[Biology:Corymbioideae|Corymbioideae]] <small>Panero & Funk</small>
* [[Biology:Famatinanthoideae|Famatinanthoideae]] <small>S.E.Freire, Ariza & Panero</small>
* [[Biology:Gochnatioideae|Gochnatioideae]] <small>Panero & Funk</small>
* [[Biology:Gymnarrhenoideae|Gymnarrhenoideae]] <small>Panero & Funk</small>
* [[Biology:Hecastocleidoideae|Hecastocleidoideae]] <small>Panero & Funk</small>
* [[Biology:Mutisioideae|Mutisioideae]] <small>Lindl.</small>
* [[Biology:Pertyoideae|Pertyoideae]] <small>Panero & Funk</small>
* [[Biology:Stifftioideae|Stifftioideae]] <small>Panero</small>
* [[Biology:Wunderlichioideae|Wunderlichioideae]] <small>Panero & Funk</small>
|synonyms =
{{Collapsible list | title = List | bullets = on
| Compositae <small>Giseke</small>
| Acarnaceae <small>[[Biography:Johann Heinrich Friedrich Link|Link]]</small>
| Ambrosiaceae <small>Bercht. & J.Presl</small>
| Anthemidaceae <small>Bercht. & J.Presl</small>
| Aposeridaceae <small>[[Biography:Constantine Samuel Rafinesque|Raf.]]</small>
| Arctotidaceae <small>Bercht. & J.Presl</small>
| Artemisiaceae <small>Martinov</small>
| Athanasiaceae <small>Martinov</small>
| Calendulaceae <small>Bercht. & J.Presl</small>
| Carduaceae <small>Bercht. & J.Presl</small>
| Cassiniaceae <small>Sch.Bip.</small>
| Cichoriaceae <small>Juss.</small>
| Coreopsidaceae <small>Link</small>
| Cynaraceae <small>Spenn.</small>
| Echinopaceae <small>Bercht. & J.Presl</small>
| Eupatoriaceae <small>Bercht. & J.Presl</small>
| Helichrysaceae <small>Link</small>
| Inulaceae <small>Bercht. & J.Presl</small>
| Lactucaceae <small>Drude</small>
| Mutisiaceae <small>Burnett</small>
| Partheniaceae <small>Link</small>
| Perdiciaceae <small>Link</small>
| Senecionaceae <small>Bercht. & J.Presl</small>
| Vernoniaceae <small>Burmeist.</small>
}}
|synonyms_ref = <ref name=GRIN/>
}}

The family '''Asteraceae''' ({{IPAc-en|ˌ|æ|s|t|ə|'|r|eɪ|s|i|.|iː|,_|-|s|i|.|aɪ}}), with the original name '''Compositae''',<ref name=BarkleyEtAl/> consists of over 32,000 known species of [[Biology:Flowering plant|flowering plant]]s in over 1,900 genera within the [[Biology:Order|order]] [[Biology:Asterales|Asterales]]. Commonly referred to as the '''aster''', '''daisy''', '''composite''', or '''sunflower family''', Compositae were first described in the year 1740. The number of species in Asteraceae is rivaled only by the [[Biology:Orchidaceae|Orchidaceae]], and which is the larger family is unclear as the quantity of extant species in each family is unknown.

Most species of Asteraceae are [[Biology:Annual plant|annual]], [[Biology:Biennial plant|biennial]], or [[Biology:Perennial plant|perennial]] herbaceous plants, but there are also shrubs, vines, and trees. The family has a widespread distribution, from subpolar to tropical regions, in a wide variety of habitats. Most occur in hot desert and cold or hot [[Earth:Semi-arid climate|semi-desert climates]], and they are found on every continent but [[Earth:Antarctica|Antarctica]]. Their primary common characteristic is flower heads, technically known as capitula, consisting of sometimes hundreds of tiny individual florets enclosed by a whorl of protective [[Biology:Bract#Involucral bracts|involucral bract]]s.

The oldest known fossils are pollen grains from the [[Earth:Late Cretaceous|Late Cretaceous]] ([[Earth:Campanian|Campanian]] to [[Earth:Maastrichtian|Maastrichtian]]) of Antarctica, dated to {{Circa|76–66}} million years ago (mya). It is estimated that the [[Biology:Crown group|crown group]] of Asteraceae evolved at least 85.9 mya (Late Cretaceous, [[Earth:Santonian|Santonian]]) with a stem node age of 88–89 mya (Late Cretaceous, [[Earth:Coniacian|Coniacian]]).

Asteraceae is an economically important family, providing food staples, garden plants, and herbal medicines. Species outside of their native ranges can be considered weedy or [[Earth:Invasive species|invasive]].

==Description<span class="anchor" id="Characteristics"></span>==
Members of the Asteraceae are mostly herbaceous plants, but some shrubs, vines, and trees (such as ''[[Biology:Lachanodes arborea|Lachanodes arborea]]'') do exist. Asteraceae species are generally easy to distinguish from other plants because of their unique [[Biology:Inflorescence|inflorescence]] and other shared characteristics, such as the joined anthers of the [[Biology:Stamen|stamen]]s.<ref name=Judd/> Nonetheless, determining genera and species of some groups such as ''[[Biology:Hieracium#Species|Hieracium]]'' is notoriously difficult (see "[[Biology:Damned yellow composite|damned yellow composite]]" for example).<ref name=MandelEtAl2019 />

===Roots===
Members of the family Asteraceae generally produce [[Biology:Taproot|taproot]]s, but sometimes they possess fibrous root systems. Some species have underground stems in the form of [[Biology:Caudex|caudices]] or [[Biology:Rhizome|rhizome]]s. These can be fleshy or woody depending on the species.<ref name=BarkleyEtAl/>

===Stems===
The stems are herbaceous, aerial, branched, and cylindrical with glandular hairs, generally erect, but can be prostrate to ascending. The stems can contain secretory canals with resin,<ref name=BarkleyEtAl/> or [[Chemistry:Latex|latex]], which is particularly common among the [[Biology:Cichorioideae|Cichorioideae]].<ref name=KGL/>

===Leaves===
Leaves can be alternate, opposite, or [[Biology:Whorl (botany)|whorled]]. They may be simple, but are often deeply lobed or otherwise incised, often conduplicate or revolute. The margins also can be entire or toothed. Resin<ref name=BarkleyEtAl/> or latex<ref name=KGL/> can also be present in the leaves.

===Inflorescences<span class="anchor" id="Flowers"></span>===

Nearly all Asteraceae bear their flowers in dense flower heads called ''capitula''. They are surrounded by [[Biology:Bract#Involucral bracts|involucral bracts]], and when viewed from a distance, each capitulum may appear to be a single flower. Enlarged outer (peripheral) flowers in the capitulum may resemble petals, and the involucral bracts may look like a calyx.

==== Floral heads ====
[[File:Bidens flwr.jpg|thumb|right|300px|A typical Asteraceae flower head showing the (five) individual ray florets and the (approximately 16) disk florets of a specimen of (''[[Biology:Bidens torta|Bidens torta]]'')]]

In plants of the Asteraceae, what appears to be a single "daisy"-type flower is actually a composite of several much smaller flowers, known as the ''capitulum'' or ''head''. By visually presenting as a single flower, the capitulum functions in attracting pollinators, in the same manner that other "showy" flowering plants in numerous other, older, plant families have evolved to attract pollinators. The previous name for the family, ''Compositae'', reflects the fact that what appears to be a single floral entity is in fact a ''composite'' of much smaller flowers.<ref name=CDF/>

The "petals" or "sunrays" in an "asteraceous" head are in fact individual strap-shaped<ref name=MDW/> flowers called ''ray flowers'' or ''ray florets'', and the "sun disk" is made up of smaller, radially symmetric, individual flowers called ''disc flowers'' or ''disc florets''. The word ''aster'' means "star" in Greek, referring to the appearance of most family members as a "[[Astronomy:Star|celestial body]] with rays".
The capitulum, which often appears to be a single flower, is often referred to as a ''head''.<ref name=Spellenberg2001 /> In some species, the entire head is able to pivot its floral stem in the course of the day to track the sun (like a "smart" solar panel), thus maximizing the reflectivity of the entire floral unit and further attracting flying pollinators.<ref name=CDF/>

Nearest to the flower stem lie a series of small, usually green, scale-like [[Biology:Bract|bract]]s. These are known as ''[[Biology:Phyllary|phyllaries]]''; collectively, they form the [[Biology:Involucre|involucre]], which serves to protect the immature head of florets during its development.{{r|CDF|p=29}} The individual florets are arranged atop a dome-like structure called the ''receptacle''.{{r|CDF}}

The individual florets in a head consist, developmentally, of five fused [[Biology:Petal|petal]]s (rarely four); instead of [[Biology:Sepal|sepal]]s, they have threadlike, hairy, or bristly structures,<ref name=Spellenberg2001 /> known collectively as a ''pappus'', (plural ''pappi''). The pappus surrounds the ovary and can, when mature and attached to a seed, adhere to animal fur or be carried by air currents, aiding in [[Earth:Seed dispersal|seed dispersal]]. The whitish, fluffy head of a dandelion, commonly blown on by children, consists of numerous seeds resting on the receptacle, each seed attached to its pappus. The pappi provide a parachute-like structure to help the seed travel from its point of origin to a more hospitable site.<ref name=CDF/>

[[File:Leucanthemum diagram.svg|thumb|upright=3|center|alt=refer to caption|Schemes and floral diagrams of the different floret types of the Asteraceae: ''[[Leucanthemum vulgare]]'': a = disc flower; b = ray flower.<br>
1 – [[Biology:Style (botany)|style]] with [[Biology:Stigma (botany)|stigmas]]<br>
2 – anthers
<br>3 – corolla ([[Biology:Petal|petal]]s); typically, in the ray flower, three petals are joined to form a strap (in other species, five petals can fuse to form a ''ligule'')<br>
4 – reduced calyx<br>
4’ – ''Carduus acanthoides'' (left shaded circle): pappus: in many Asteraceae species, the calyx develops as a fibrous or bristly pappus<br>
5 – inferior ovary: fused ovary consisting of two carpels, containing one abaxial [[Biology:Ovule|ovule]] (basal placentation).]]

A ''ray flower'' is a two- or three-lobed, strap-shaped, individual flower, found in the head of most members of the Asteraceae.<ref name=CDF/><ref name=MDW/> The [[Biology:Petal|corolla]] of the ray flower may have two tiny, vestigial teeth, opposite to the three-lobed strap, or tongue, indicating its evolution by fusion from an ancestral, five-part corolla. In some species, the 3:2 arrangement is reversed, with two lobes, and zero or three tiny teeth visible opposite the tongue.

A ''ligulate flower'' is a five-lobed, strap-shaped, individual flower found in the heads of certain other asteraceous species.<ref name=CDF/> A ''ligule'' is the strap-shaped tongue of the corolla of either a ray flower or of a ligulate flower.{{clarify|date=April 2023}}{{example needed|date=April 2023}}<ref name=MDW/> A ''disk flower'' (or ''disc flower'') is a radially symmetric individual flower in the head, which is ringed by the ray flowers when both are present.<ref name=CDF/><ref name=MDW/> In some species, ray flowers may be arranged around the disc in irregular symmetry, or with a weakly bilaterally symmetric arrangement.<ref name=CDF/>

== Variations ==
A ''radiate head'' has disc flowers surrounded by ray flowers. A ''ligulate head'' has all ligulate flowers and no disc flowers. When an Asteraceae flower head has only disc flowers that are either sterile, male, or [[Biology:Plant reproductive morphology#Bisexual|bisexual]] (but not female and fertile), it is a ''discoid head''.

''Disciform heads'' possess only disc flowers in their heads, but may produce two different sex types (male or female) within their disciform head.

Some other species produce two different head types: ''staminate'' (all-male), or ''pistillate'' (all-female). In a few unusual species, the "head" will consist of one single disc flower; alternatively, a few species will produce both single-flowered female heads, along with multi-flowered male heads, in their "pollination strategy".<ref name=CDF/>

=== Floral structures<span class="anchor" id="Floral structures"></span> ===
[[File:Carduus floral diagram.svg|thumb|Flower diagram of ''[[Biology:Carduus|Carduus]]'' (Carduoideae) shows (outermost to innermost): subtending bract and stem axis; calyx forming a pappus; fused corolla; stamens fused to corolla; gynoecium with two carpels and one locule]]
[[File:Senecio mikanioides (delairea odorata).jpg|thumb|Discoid flowerheads of ''[[Biology:Delairea odorata|Delairea odorata]]''.]]
The distinguishing characteristic of Asteraceae is their [[Biology:Inflorescence|inflorescence]], a type of specialised, composite flower head or ''[[Biology:Pseudanthium|pseudanthium]]'', technically called a calathium or ''[[Biology:Pseudanthium|capitulum]]'',<ref name=Beentje/><ref name=Usher/> that may look superficially like a single flower. The ''capitulum'' is a contracted [[Biology:Raceme|raceme]] composed of numerous individual [[Biology:Sessility (botany)|sessile]] [[Biology:Flower|flower]]s, called ''florets'', all sharing the same [[Biology:Receptacle (botany)|receptacle]].

A set of [[Biology:Bract|bract]]s forms an involucre surrounding the base of the capitulum. These are called "phyllaries", or "involucral bracts". They may simulate the sepals of the pseudanthium. These are mostly herbaceous but can also be brightly coloured (e.g. ''[[Biology:Helichrysum|Helichrysum]]'') or have a scarious (dry and membranous) texture. The phyllaries can be free or fused, and arranged in one to many rows, overlapping like the tiles of a roof (''imbricate'') or not (this variation is important in identification of tribes and genera).

Each floret may be subtended by a bract, called a "palea" or "receptacular bract". These bracts are often called "[[Biology:Chaff|chaff]]". The presence or absence of these bracts, their distribution on the receptacle, and their size and shape are all important diagnostic characteristics for genera and tribes.

The florets have five petals fused at the base to form a corolla tube and they may be either actinomorphic or zygomorphic. ''Disc florets'' are usually actinomorphic, with five petal lips on the rim of the corolla tube. The petal lips may be either very short, or long, in which case they form deeply lobed petals. The latter is the only kind of floret in the [[Biology:Carduoideae|Carduoideae]], while the first kind is more widespread. ''Ray florets'' are always highly zygomorphic and are characterised by the presence of a ''ligule'', a strap-shaped structure on the edge of the corolla tube consisting of fused petals. In the Asteroideae and other minor subfamilies these are usually borne only on florets at the circumference of the capitulum and have a 3+2 scheme – above the fused corolla tube, three very long fused petals form the ligule, with the other two petals being inconspicuously small. The Cichorioideae has only ray florets, with a 5+0 scheme – all five petals form the ligule. A 4+1 scheme is found in the Barnadesioideae. The tip of the ligule is often divided into teeth, each one representing a petal. Some marginal florets may have no petals at all (filiform floret).

The calyx of the florets may be absent, but when present is always modified into a [[Biology:Pappus (botany)|pappus]] of two or more teeth, scales or bristles and this is often involved in the dispersion of the seeds. As with the bracts, the nature of the pappus is an important diagnostic feature.

There are usually four or five [[Biology:Stamen|stamen]]s.<ref name=Spellenberg2001 /> The filaments are fused to the corolla, while the anthers are generally [[Biology:Connation|connate]] (''syngenesious'' anthers), thus forming a sort of tube around the style (''theca''). They commonly have basal and/or apical appendages. Pollen is released inside the tube and is collected around the growing style, and then, as the style elongates, is pushed out of the tube (''nüdelspritze'').

The pistil consists of two connate carpels. The style has two lobes. Stigmatic tissue may be located in the interior surface or form two lateral lines. The ovary is inferior and has only one [[Biology:Ovule|ovule]], with basal [[Biology:Placentation|placentation]].

=== Fruits and seeds ===
In members of the Asteraceae the fruit is [[Biology:Achene|achene]]-like, and is called a ''cypsela'' (plural ''cypselae''). Although there are two fused carpels, there is only one locule, and only one [[Biology:Seed|seed]] per fruit is formed.<ref name=Spellenberg2001 /> It may sometimes be winged or spiny because the pappus, which is derived from calyx tissue often remains on the fruit (for example in dandelion). In some species, however, the pappus falls off (for example in ''[[Biology:Helianthus|Helianthus]]''). Cypsela [[Biology:Morphology|morphology]] is often used to help determine plant relationships at the genus and species level.<ref name=McKenzieEtAl/> The mature [[Biology:Seed|seed]]s usually have little [[Biology:Endosperm|endosperm]] or none.<ref name=Judd/>

=== Pollen ===
The pollen of composites is typically echinolophate, a [[Biology:Morphology|morphological]] term meaning "with elaborate systems of ridges and spines dispersed around and between the apertures."<ref name=Tomb1974/>

=== Metabolites ===
In Asteraceae, the energy store is generally in the form of [[Chemistry:Inulin|inulin]] rather than starch. They produce iso/[[Chemistry:Chlorogenic acid|chlorogenic acid]], [[Chemistry:Sesquiterpene|sesquiterpene]] lactones, pentacyclic triterpene alcohols, various [[Chemistry:Alkaloid|alkaloid]]s, [[Chemistry:Acetylene|acetylene]]s (cyclic, aromatic, with vinyl end groups), [[Chemistry:Tannin|tannin]]s. They have [[Biology:Terpene|terpenoid]] [[Chemistry:Essential oil|essential oil]]s that never contain [[Chemistry:Iridoid|iridoid]]s.<ref name=Stevens/>

Asteraceae produce secondary metabolites, such as [[Chemistry:Flavonoid|flavonoid]]s and [[Biology:Terpenoid|terpenoid]]s. Some of these molecules can inhibit protozoan parasites such as ''[[Biology:Plasmodium|Plasmodium]]'', ''[[Biology:Trypanosoma|Trypanosoma]]'', ''[[Biology:Leishmania|Leishmania]]'' and parasitic intestinal worms, and thus have potential in medicine.<ref name=PandaLuyten2018/>

== Taxonomy ==

===History===
Compositae, the original name for Asteraceae, were first described in 1740 by Dutch botanist Adriaan van Royen.{{r|vanRoyen1740|p=117–118}} Traditionally, two subfamilies were recognised: [[Biology:Asteroideae|Asteroideae]] (or Tubuliflorae) and [[Biology:Cichorioideae|Cichorioideae]] (or Liguliflorae).{{r|BayerStarr1998|p=242}} The latter has been shown to be extensively paraphyletic, and has now been divided into 12 subfamilies, but the former still stands.<ref name=PaneroFunk/>{{Update inline|reason=Possibly changed since 2002; see Mandel, et al (2019)|date=February 2021}} The study of this family is known as [[Biology:Synantherology|synantherology]].

===Phylogeny===

The [[Biology:Phylogenetic tree|phylogenetic tree]] of subfamilies presented below is based on Panero & Funk (2002)<ref name=PaneroFunk/> updated in 2014,<ref name=Deep/> and now also includes the monotypic [[Biology:Famatinanthoideae|Famatinanthoideae]].<ref name=Deep/><ref name=FJNG/><ref name=FunkFragman-Sapir/>{{Update inline|reason=Possibly changed since 2014|date=February 2021}}
The diamond (♦) denotes a very poorly supported [[Biology:Cladistics|node]] (<50% bootstrap support), the dot (•) a poorly supported node (<80%).<ref name=Stevens/>

{{clade|style=font-size:90%;line-height:90%
|1={{clade|1='''[[Biology:Barnadesioideae|Barnadesioideae]]''': 9 genera, 93 species. [[Earth:South America|South America]], mainly the [[Place:Andes|Andes]].
|2={{clade|1='''[[Biology:Famatinanthoideae|Famatinanthoideae]]''': South America, 1 genus, 1 species ''Famatinanthus decussatus''.
|2={{clade|1='''[[Biology:Mutisioideae|Mutisioideae]]''': 58 genera, 750 species. Absent from Europe, mostly in South America.
|2='''[[Biology:Stifftioideae|Stifftioideae]]''': 10 genera. South America.
|label3=♦|3={{clade|1='''[[Biology:Wunderlichioideae|Wunderlichioideae]]''': 8 genera, 24 species. Mostly in Venezuela and Guyana.
|2={{clade|1='''[[Biology:Gochnatioideae|Gochnatioideae]]''': 4 or 5 genera, 90 species. Latin America and southern United States.
|2={{clade
|1='''[[Biology:Hecastocleidoideae|Hecastocleidoideae]]''': Only ''Hecastocleis shockleyi''. Southwestern United States.
|2={{clade
|1='''[[Biology:Carduoideae|Carduoideae]]''': 83 genera, 2,500 species. Worldwide.
|2={{clade
|1='''[[Biology:Pertyoideae|Pertyoideae]]''': 5 or 6 genera, 70 species. Asia.
|2={{clade|1='''[[Biology:Gymnarrhenoideae|Gymnarrhenoideae]]''': Two genera/species, ''Gymnarrhena micrantha'' (Northern Africa, [[Place:Middle East|Middle East]]) and ''[[Biology:Cavea tanguensis|Cavea tanguensis]]'' ([[Place:Eastern Himalayas|Eastern Himalayas]]).
|label2=•|2={{clade|1='''[[Biology:Cichorioideae|Cichorioideae]]''': 224 genera, 3,200 species. Worldwide.
|2={{clade
|1='''[[Biology:Corymbioideae|Corymbioideae]]''': Only the genus ''[[Biology:Corymbium|Corymbium]]'', with 9 species. Cape provinces, South Africa.
|label2=•|2='''[[Biology:Asteroideae|Asteroideae]]''': 1,130 genera and 16,200 species. Worldwide.}}}}}}}}}}}}}}}}}}}}}}}}

The family includes over 32,000 currently accepted species, in over 1,900 genera ([[Biology:List of Asteraceae genera|list]]) in 13 subfamilies.<ref name=PlantList/>{{Update inline|reason=Possibly changed since 2016|date=February 2021}} The number of species in the family Asteraceae is rivaled only by Orchidaceae.<ref name=Stevens/><ref name=PaneroCrozier/> Which is the larger family is unclear, because of the uncertainty about how many extant species each family includes.{{Citation needed|date=February 2021}} The four subfamilies [[Biology:Asteroideae|Asteroideae]], [[Biology:Cichorioideae|Cichorioideae]], [[Biology:Carduoideae|Carduoideae]] and [[Biology:Mutisioideae|Mutisioideae]] contain 99% of the species diversity of the whole family (approximately 70%, 14%, 11% and 3% respectively).{{Citation needed|date=February 2021}}

Because of the [[Biology:Morphology|morphological]] complexity exhibited by this family, agreeing on generic circumscriptions has often been difficult for taxonomists. As a result, several of these genera have required multiple revisions.<ref name=Judd/>

==== Paleontology and evolutionary processes ====
The oldest known fossils of members of Asteraceae are pollen grains from the [[Earth:Late Cretaceous|Late Cretaceous]] of [[Earth:Antarctica|Antarctica]], dated to ~76–66 mya ([[Earth:Campanian|Campanian]] to [[Earth:Maastrichtian|Maastrichtian]]) and assigned to the extant genus ''[[Biology:Dasyphyllum|Dasyphyllum]]''. Barreda, ''et al.'' (2015) estimated that the [[Biology:Crown group|crown group]] of Asteraceae evolved at least 85.9 mya (Late Cretaceous, [[Earth:Santonian|Santonian]]) with a stem node age of 88–89 mya (Late Cretaceous, [[Earth:Coniacian|Coniacian]]).<ref name=PNAS2015/>

It is not known whether the precise cause of their great success was the development of the highly specialised capitulum, their ability to store energy as [[Chemistry:Fructan|fructan]]s (mainly [[Chemistry:Inulin|inulin]]), which is an advantage in relatively dry zones, or some combination of these and possibly other factors.<ref name=Stevens/> Heterocarpy, or the ability to produce different fruit morphs, has evolved and is common in Asteraceae. It allows seeds to be dispersed over varying distances and each is adapted to different environments, increasing chances of survival.<ref name=Gardocki/>

=== Etymology and pronunciation ===
The original name Compositae is still valid under the International Code of Nomenclature for algae, fungi, and plants.<ref name=ICN185/> It refers to the "composite" nature of the capitula, which consist of a few or many individual flowers.{{Citation needed|date=February 2021}}

The alternative (as it came later) name Asteraceae ({{IPAc-en|lang|ˌ|æ|s|t|ə|ˈ|r|eɪ|s|i|,_|-|s|i|ˌ|aɪ|,_|-|s|i|ˌ|eɪ|,_|-|s|i|ˌ|iː}}) comes to international scientific vocabulary from Neo-Latin, from ''[[Biology:Aster (genus)|Aster]]'', the [[Biology:Type genus|type genus]], + ''-aceae'',<ref name=MWU/> a standardized [[Social:Suffix|suffix]] for plant family names in modern taxonomy. This genus name comes from the Classical Latin word {{wikt-lang|la|aster}}, "star", which came from [[Social:Ancient Greek|Ancient Greek]] {{lang|grc|ἀστήρ}} ({{transl|grc|astḗr}}), "star".<ref name=MWU/> It refers to the star-like form of the inflorescence.{{Citation needed|date=February 2021}}

The vernacular name ''daisy'', widely applied to members of this family, is derived from the [[Social:Old English|Old English]] name of the daisy (''[[Biology:Bellis perennis|Bellis perennis]]''): {{wikt-lang|ang|dæġes ēaġe}}, meaning "day's eye". This is because the petals open at dawn and close at dusk.<ref name=daisy />

== Distribution and habitat ==
Asteraceae species have a widespread distribution, from subpolar to tropical regions in a wide variety of habitats. Most occur in hot desert and cold or hot semi-desert climates, and they are found on every continent but Antarctica. They are especially numerous in tropical and subtropical regions (notably Central America, eastern {{wipe|Brazil}}, the Mediterranean, the Levant, [[Place:Southern Africa|southern Africa]], [[Place:Central Asia|central Asia]], and southwestern China).<ref name=PaneroCrozier/> The largest proportion of the species occur in the arid and semi-arid regions of subtropical and lower temperate latitudes.<ref name=BarkleyEtAl/> The Asteraceae family comprises 10% of all flowering plant species.<ref name=MandelEtAl2019/>

== Ecology ==
{{multiple image
| align     = right
| direction = horizontal
| image1    = Kih-silberdistel.jpg
| width1    = {{#expr: trunc (150 * 800 / 532)}}
| caption1  = Anemochory in ''[[Biology:Carlina|Carlina]]''
| image2    = Epizoochoria NRM.jpg
| width2    = {{#expr: trunc (150 * 800 / 600)}}
| caption2  = Epizoochory in ''[[Biology:Bidens tripartita|Bidens tripartita]]''
}}
Asteraceae are especially common in open and dry environments.<ref name=Judd/> Many members of Asteraceae are [[Biology:Pollination|pollinated]] by insects, which explains their value in attracting beneficial insects, but [[Biology:Pollination syndrome|anemophily]] is also present (e.g. ''[[Biology:Ragweed|Ambrosia]]'', ''[[Biology:Artemisia (genus)|Artemisia]]''). There are many [[Biology:Apomixis|apomictic]] species in the family.

Seeds are ordinarily dispersed intact with the fruiting body, the cypsela. ''[[Earth:Seed dispersal#Wind|Anemochory]]'' (wind dispersal) is common, assisted by a hairy pappus. ''[[Earth:Seed dispersal#By animals|Epizoochory]]'' is another common method, in which the dispersal unit, a single cypsela (e.g. ''[[Biology:Bidens|Bidens]]'') or entire capitulum (e.g. ''[[Biology:Arctium|Arctium]]'') has hooks, spines or some structure to attach to the fur or plumage (or even clothes, as in the photo) of an animal just to fall off later far from its mother plant.

Some members of Asteraceae are economically important as weeds. Notable in the United States are ''[[Biology:Senecio jacobaea|Senecio jacobaea]]'' (ragwort),<ref name=WSNWCB/> ''[[Biology:Senecio vulgaris|Senecio vulgaris]]'' (groundsel),<ref name=WSNWCB2/> and ''[[Biology:Taraxacum|Taraxacum]]'' (dandelion).<ref name=Gonzalez/> Some are [[Earth:Invasive species|invasive species]] in particular regions, often having been introduced by human agency. Examples include various [[Biology:Tumbleweed|tumbleweed]]s, ''[[Biology:Bidens|Bidens]]'', [[Biology:Ambrosia (plant)|ragweeds]], [[Thistle|thistle]]s, and [[Biology:Taraxacum officinale|dandelion]].<ref name=MartinFores2018 /> Dandelion was introduced into [[Earth:North America|North America]] by European settlers who used the young leaves as a salad green.<ref name=dandelion/> A number of species are toxic to grazing animals.<ref name=Spellenberg2001 />

== Uses ==
[[File:Senecio angulatus flowers.jpg|thumb|upright=1.5|left|The [[Biology:Vine|twining]] succulent, ''Senecio angulatus'', is used for its cut flowers,<ref name=senecioang/> despite being an invasive weed in some places, such as Victoria, Australia and New Zealand.<ref name=senecioang2/>]]
Asteraceae is an economically important family, providing products such as cooking oils, leaf vegetables like [[Biology:Lettuce|lettuce]], sunflower seeds, [[Biology:Artichoke|artichoke]]s, sweetening agents, coffee substitutes and herbal teas. Several genera are of [[Earth:Horticulture|horticultural]] importance, including pot marigold (''[[Biology:Calendula officinalis|Calendula officinalis]]''), ''[[Biology:Echinacea|Echinacea]]'' (coneflowers), various daisies, [[Biology:Erigeron|fleabane]], [[Biology:Chrysanthemum|chrysanthemum]]s, [[Biology:Dahlia|dahlia]]s, [[Biology:Zinnia|zinnia]]s, and [[Biology:Helenium|helenium]]s. Asteraceae are important in herbal medicine, including ''[[Biology:Grindelia|Grindelia]]'', [[Biology:Achillea millefolium|yarrow]], and many others.<ref name=Michel2020 />

Commercially important plants in Asteraceae include the food crops ''[[Biology:Lactuca sativa|Lactuca sativa]]'' (lettuce), ''[[Biology:Cichorium|Cichorium]]'' (chicory), ''[[Biology:Cynara scolymus|Cynara scolymus]]'' (globe artichoke), ''[[Biology:Helianthus annuus|Helianthus annuus]]'' (sunflower), ''[[Biology:Smallanthus sonchifolius|Smallanthus sonchifolius]]'' (yacón), ''[[Biology:Carthamus tinctorius|Carthamus tinctorius]]'' (safflower) and ''[[Biology:Helianthus tuberosus|Helianthus tuberosus]]'' (Jerusalem artichoke).<ref name=ezcurra />

Plants are used as herbs and in herbal teas and other beverages. Chamomile, for example, comes from two different species: the annual ''[[Biology:Matricaria chamomilla|Matricaria chamomilla]]'' (German chamomile) and the perennial ''[[Biology:Chamaemelum nobile|Chamaemelum nobile]]'' (Roman chamomile). ''[[Biology:Calendula|Calendula]]'' (known as pot marigold) is grown commercially for herbal teas and potpourri. ''[[Biology:Echinacea purpurea|Echinacea]]'' is used as a medicinal tea. The wormwood genus ''[[Biology:Artemisia (genus)|Artemisia]]'' includes [[Biology:Absinthe|absinthe]] (''A. absinthium'') and [[Biology:Tarragon|tarragon]] (''A. dracunculus''). Winter tarragon (''[[Biology:Tagetes lucida|Tagetes lucida]]''), is commonly grown and used as a tarragon substitute in climates where tarragon will not survive.<ref name=jekkas />

Many members of the family are grown as ornamental plants for their flowers, and some are important ornamental crops for the cut flower industry. Some examples are ''[[Biology:Chrysanthemum|Chrysanthemum]]'', ''[[Biology:Gerbera|Gerbera]]'', ''[[Biology:Calendula|Calendula]]'', ''Dendranthema'', ''[[Biology:Argyranthemum|Argyranthemum]]'', ''[[Biology:Dahlia|Dahlia]]'', ''[[Biology:Tagetes|Tagetes]]'', ''[[Biology:Zinnia|Zinnia]]'', and many others.<ref name=Aphididae/>
[[File:Senecio madagascariensis plant13 (16143326805).jpg|thumb|upright=1.2|''[[Biology:Senecio madagascariensis|Senecio madagascariensis]]'' (Fireweed) is an environmental weed in Australia, growing in wastelands, grasslands and suburban bushland.<ref name=AuldMeld1992/>]]
Many species of this family possess medicinal properties and are used as traditional [[Biology:Parasite|antiparasitic]] medicine.<ref name=PandaLuyten2018/>

Members of the family are also commonly featured in medical and phytochemical journals because the [[Chemistry:Sesquiterpene|sesquiterpene]] [[Chemistry:Lactone|lactone]] compounds contained within them are an important cause of allergic contact dermatitis. Allergy to these compounds is the leading cause of allergic contact dermatitis in florists in the US.<ref name=OdomJamesBerger2000/> [[Biology:Pollen|Pollen]] from ragweed ''[[Biology:Ragweed|Ambrosia]]'' is among the main causes of so-called hay fever in the {{wipe|United States}}.<ref name=AsthmaAllergy/>

Asteraceae are also used for some industrial purposes. French Marigold (''[[Biology:Tagetes patula|Tagetes patula]]'') is common in commercial poultry feeds and its oil is extracted for uses in cola and the cigarette industry. The genera ''[[Biology:Chrysanthemum|Chrysanthemum]]'', ''[[Biology:Pulicaria|Pulicaria]]'', ''[[Biology:Tagetes|Tagetes]]'', and ''[[Biology:Tanacetum|Tanacetum]]'' contain species with useful insecticidal properties. ''[[Biology:Parthenium argentatum|Parthenium argentatum]]'' (guayule) is a source of hypoallergenic [[Chemistry:Latex|latex]].<ref name=Aphididae/>

Several members of the family are copious nectar producers<ref name=Aphididae/> and are useful for evaluating [[Biology:Pollinator|pollinator]] populations during their bloom.{{Citation needed|date=January 2017}} ''[[Biology:Centaurea|Centaurea]]'' (knapweed), ''Helianthus annuus'' (domestic sunflower), and some species of ''[[Biology:Solidago|Solidago]]'' (goldenrod) are major "honey plants" for beekeepers. ''Solidago'' produces relatively high protein [[Biology:Pollen|pollen]], which helps [[Biology:Honey bee|honey bee]]s over winter.<ref name=SolidagoWinter/>
{{clear}}

==References==
{{Reflist|refs=

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| date      = 1998
| title     = Tribal phylogeny of the Asteraceae based on two non-coding chloroplast sequences, the trnL Intron and trnL/trnF Intergenic Spacer
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<ref name=jekkas>
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| last5      = del Pozo     | first5 = Alejandro
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| archive-date     = 15 January 2023
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| title        = Tansy ragwort ''Jacobaea vulgaris''
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| url          = http://www.ars-grin.gov/cgi-bin/npgs/html/family.pl?110
| website      = [[Biology:Germplasm Resources Information Network|Germplasm Resources Information Network]] (GRIN) (www.ars-grin.gov)
| publisher    = USDA, [[Organization:Agricultural Research Service|ARS]], National Genetic Resources Program, National Germplasm Resources Laboratory
| location     = Beltsville, Maryland
| archive-url  = https://web.archive.org/web/20080917105707/http://www.ars-grin.gov/cgi-bin/npgs/html/family.pl?110
| archive-date = 17 September 2008
| url-status   = dead <!-- may not be dead, but when I (Eewilson) tried to access it February 2021, it timed out more than once -->
| language     = en
| access-date  = 12 June 2008
}}
</ref>

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<ref name = GBW>
{{Cite book
| last1     = Blackwell | first1 = Laird R.
| year      = 2006
| title     = Great Basin Wildflowers: A Guide to Common Wildflowers of the High Deserts of Nevada, Utah, and Oregon
| page      = 275
| isbn      = 978-0762738052
| publisher = FalconGuides
| location  = Guilford, Connecticut
| language  = en
}}
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<ref name = PNAS2015>
{{Cite journal
| last1      = Barreda   | first1 = Viviana D.
| last2      = Palazzesi | first2 = Luis
| last3      = Tellería  | first3 = Maria C.
| last4      = Olivero   | first4 = Eduardo B.
| last5      = Raine     | first5 = J. Ian
| last6      = Forest    | first6 = Félix
| year       = 2015
| title      = Early evolution of the angiosperm clade Asteraceae in the Cretaceous of Antarctica
| journal    = Proceedings of the National Academy of Sciences of the United States of America
| volume     = 112
| issue      = 35
| pages      = 10989–10994
| pmid       = 26261324
| pmc        = 4568267
| bibcode    = 2015PNAS..11210989B
| doi        = 10.1073/pnas.1423653112
| doi-access = free
| language   = en
}}
</ref>

<ref name = tropicos>
{{Tropicos
| 50307371
| Asteraceae
| Bercht. & J.Presl
| language    = en
| access-date = 14 July 2017
}}
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<ref name = Stevens>
{{Cite web
 |year         = 2022
 |orig-year    = 2001
 |title        = Angiosperm Phylogeny Website: Asterales
 |url          = http://www.mobot.org/mobot/research/apweb/welcome.html#Asterales
 |website      = Angiosperm Phylogeny Website
 |archive-date = 29 November 2022
 |archive-url  = https://web.archive.org/web/20221129174143fw_/https://www.mobot.org/MOBOT/research/APweb/orders/Asteralesweb.htm#Asterales
 |url-status   = dead
 |language     = en
 |access-date  = 15 January 2023
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<ref name = PlantList>
{{Cite web
 |author       = <!--Not stated-->
 |year         = 2013
 |title        = Compositae
 |url          = http://www.theplantlist.org/1.1/browse/A/Compositae/
 |website      = The Plant List (www.theplantlist.org)
 |publisher    = [[Organization:Royal Botanic Gardens, Kew|Royal Botanic Gardens, Kew]] & [[Organization:Missouri Botanical Garden|Missouri Botanical Garden]]
 |archive-date = 11 February 2012
 |archive-url  = https://web.archive.org/web/20120211132122/http://www.theplantlist.org/browse/A/Compositae/
 |url-status   = live
 |language     = en
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}}
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<ref name = PaneroCrozier>
{{Cite web
 |last1        = Panero
 |last2        = Crozier
 |date         = 27 January 2012
 |title        = Asteraceae. Sunflowers, daisies
 |url          = http://tolweb.org/Asteraceae/20780/2012.01.27
 |website      = The Tree of Life Web Project (tolweb.org)
 |archive-date = 21 January 2022
 |archive-url  = https://web.archive.org/web/20220121040917/http://tolweb.org/Asteraceae/20780/2012.01.27
 |url-status   = live
 |language     = en
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}}
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<ref name = ICN185>
{{Cite web
| url          = http://www.iapt-taxon.org/nomen/main.php?page=art18
| title        = International Code of Nomenclature for algae, fungi, and plants – Article 18.5
| website      = iapt-taxon.org
| archive-date = 27 September 2013
| archive-url  = https://web.archive.org/web/20130927152429/http://www.iapt-taxon.org/nomen/main.php?page=art18
| url-status   = live
| language     = en
| access-date  = 2 June 2015
}}
</ref>

<ref name = BarkleyEtAl>
{{Cite eFloras
| 1
| 10074
| Asteraceae
| volume       = 19
| date         = 2006
| last1        = Barkley   | first1 = T.M.
| last3        = Strother  | first3 = J.L.
| language     = en
| access-date  = 20 March 2022
}}
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<ref name = dandelion>
{{Cite web
| title        = ''Taraxacum officinale'': dandelion
| url          = http://www.invasiveplantatlas.org/subject.html?sub=3887
| website      = Invasive Plant Atlas of the United States (www.invasiveplantatlas.org)
| archive-date = 23 October 2012
| archive-url  = https://web.archive.org/web/20121023040754/http://www.invasiveplantatlas.org/subject.html?sub=3887
| url-status   = live
| language     = en
| access-date  = 10 September 2012
}}
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<ref name = MWU>
{{Cite book
| author       = Merriam-Webster
| title        = Merriam-Webster's Unabridged Dictionary
| publisher    = Merriam-Webster
| url          = http://unabridged.merriam-webster.com/unabridged/
| archive-date = 25 May 2020
| archive-url  = https://web.archive.org/web/20200525084504/https://unabridged.merriam-webster.com/subscriber/login?redirect_to=%2Funabridged%2F
| url-status   = dead
| language     = en
| access-date  = 27 July 2016
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<ref name = PaneroFunk>
{{Cite journal
 |last1        = Panero
 |last2        = Funk
 |first2       = V.A.
 |year         = 2002
 |title        = Toward a phylogenetic subfamilial classification for the Compositae (Asteraceae)
 |url          = https://www.biodiversitylibrary.org/page/35519280/
 |journal      = Proceedings of the Biological Society of Washington
 |volume       = 115
 |pages        = 909–922
 |archive-date = 7 May 2021
 |archive-url  = https://web.archive.org/web/20210507141607/https://www.biodiversitylibrary.org/page/35519280/
 |url-status   = live
 |language     = en
 |access-date  = 16 February 2021
 |via          = [[Social:Biodiversity Heritage Library|Biodiversity Heritage Library]]
}}
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<ref name = Deep>
{{Cite journal
| last2    = Freire        | first2  = Susana E.
| last3    = Ariza Espinar | first3  = Luis
| last4    = Crozier       | first4  = Bonnie S.
| last5    = Barboza       | first5  = Gloria E.
| last6    = Cantero       | first6  = Juan J.
| year     = 2014
| title    = Resolution of deep nodes yields an improved backbone phylogeny and a new basal lineage to study early evolution of Asteraceae
| url      = https://www.researchgate.net/publication/264432229
| journal  = Molecular Phylogenetics and Evolution
| volume   = 80
| issue    = 1
| pages    = 43–53
| doi      = 10.1016/j.ympev.2014.07.012
| pmid     = 25083940
| hdl = 11336/36507
| s2cid    = 6745704
| language = en
| via      = [[Company:ResearchGate|ResearchGate]] ''(abstract and introduction)''
| hdl-access= free
}}
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<ref name = FJNG>
{{Cite journal
| last1       = Fu    | first1 = Zhi-Xi
| last2       = Jiao  | first2 = Bo-Han
| last3       = Nie   | first3 = Bao
| last4       = Zhang | first4 = Guo-Jin
| last5       = Gao   | first5 = Tian-Gang
| date        = 22 July 2016
| title       = A comprehensive generic-level phylogeny of the sunflower family: Implications for the systematics of Chinese Asteraceae
| url         = https://www.researchgate.net/publication/304357265
| journal     = Journal of Systematics and Evolution
| volume      = 54
| issue       = 4
| pages       = 416–437
| doi         = 10.1111/jse.12216
| s2cid       = 89115861
| language    = en
| via         = [[Company:ResearchGate|ResearchGate]]
| doi-access= free
}}
</ref>

<ref name = FunkFragman-Sapir>
{{Cite book
 |last1        = Funk
 |first1       = Vicki A.
 |last2        = Fragman-Sapir
 |first2       = Ori
 |year         = 2009
 |chapter      = 22. Gymnarrheneae (Gymnarrhenoideae)
 |title        = Systematics, Evolution, and Biogeography of Compositae
 |publisher    = [[Organization:International Association for Plant Taxonomy|International Association for Plant Taxonomy]]
 |location     = Vienna
 |editor1      = V.A. Funk
 |editor2      = A. Susanna
 |editor3      = T. Stuessy
 |editor4      = R. Bayer
 |pages        = 327–332
 |isbn         = 978-3950175431
 |chapter-url  = https://repository.si.edu/bitstream/handle/10088/11411/bot_2009_pr_Funk_Fragman-Sapir_Fymnarrheneae.pdf
 |archive-date = 6 May 2021
 |archive-url  = https://web.archive.org/web/20210506171048/https://repository.si.edu/bitstream/handle/10088/11411/bot_2009_pr_Funk_Fragman-Sapir_Fymnarrheneae.pdf
 |url-status   = live
 |language     = en
 |access-date  = 16 February 2021
}}
</ref>

<ref name = Judd>
{{Cite book
| last1     = Judd     | first1 = W.S.
| last2     = Campbell | first2 = C.S.
| last3     = Kellogg  | first3 = E.A.
| last5     = Donaghue | first5 = M.J.
| date      = 2007
| title     = Plant Systematics: A Phylogenetic Approach
| edition   = 3rd
| publisher = [[Company:Sinauer Associates|Sinauer Associates]]
| location  = Sunderland, Massachusetts
| isbn      = 978-0878934072
| language  = en
}}
</ref>

<ref name = CDF>
{{Cite book
| last1        = Morhardt | first1 = Sia
| last2        = Morhardt | first2 = Emil
| year         = 2004
| title        = California desert flowers: an introductions to families, genera, and species
| publisher    = University of California Press
| location     = Berkeley, Los Angeles, and London
| isbn         = 978-0520240032
| pages        = 29–32
| language     = en
}}
</ref>

}}

== Bibliography ==
{{refbegin}}
* {{cite book
| editor-last1 = Funk    | editor-first1 = Vicki A.
| editor-last2 = Susanna | editor-first2 = A.
| editor-last3 = Stuessy | editor-first3 = T.F.
| editor-last4 = Bayer   | editor-first4 = R.J.
| date         = 2009
| title        = Systematics, Evolution, and Biogeography of Compositae
| publisher    = [[Organization:International Association for Plant Taxonomy|International Association for Plant Taxonomy]]
| location     = Vienna
| isbn         = 978-3-9501754-3-1
| url          = https://books.google.com/books?id=oj4lAQAAMAAJ
| archive-date = 15 January 2023
| archive-url  = https://web.archive.org/web/20230115175957/https://books.google.com/books?id=oj4lAQAAMAAJ
| url-status   = live
| access-date  = 12 November 2021
| via          = [[Google Books]]
}} {{link note|note=Also available [https://archive.org/details/systematicsevolu0000unse here] at [[Internet Archive]]}}
{{refend}}

== External links ==
* [http://www.mobot.org/MOBOT/Research/APweb/orders/asteralesweb.htm#Asteraceae Asteraceae] at the [http://www.mobot.org/mobot/research/apweb/ ''Angiosperm Phylogeny Website'']
* [http://compositae.org/ Compositae.org] – Compositae Working Group (CWG) and Global Compositae Database (GCD)

{{Taxonbar|from1=Q25400|from2=Q14847254}}

[[Category:Asteraceae| ]]
[[Category:Asterales families]]
[[Category:Extant Campanian first appearances]]

{{Sourceattribution|Asteraceae}}