Asteraceae
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{{Taxobox
| name = Asteraceae
| image = Asteracea poster 3.jpg
| image_width = 250px
| image_caption = A poster with twelve different species of Asteracea from the Asteroideae, Cichorioideaea and Carduoideae subfamilies
| fossil_range = {{fossil range|80}}[[Eocene]] - Recent
| regnum = [[Plant]]ae
| divisio = [[Flowering plant|Magnoliophyta]]
| classis = [[Magnoliopsida]]
| ordo = [[Asterales]]
| familia = '''Asteraceae'''
| familia_authority = [[Friedrich von Berchtold|Bercht.]] & [[Jan Svatopluk Presl|J.Presl]]
| type_genus = ''[[Aster]]'' L.
| type_genus_authority=
| diversity = Genus
| diversity_link = List of Asteraceae genera
| subdivision_ranks = Subfamilies
| subdivision =
[[Asteroideae]]<br/>
[[Barnadesioideae]]<br/>
[[Carduoideae]]<br/>
[[Cichorioideae]]<br/>
[[Corymbioideae]]<br/>
[[Gochnatioideae]]<br/>
[[Gymnarrhenoideae]]<br/>
[[Hecastocleidoideae]]<br/>
[[Mutisioideae]]<br/>
[[Pertyoideae]]<br/>
[[Stifftioideae]]<br/>
[[Wunderlichioideae]]<br/>
| synonyms =
Compositae <small>Giseke</small><br/>
Acarnaceae <small>Link</small><br/>
Ambrosiaceae <small>Bercht. & J. Presl</small><br/>
Anthemidaceae <small>Bercht. & J. Presl</small><br/>
Aposeridaceae <small>Raf.</small><br/>
Arctotidaceae <small>Bercht. & J. Presl</small><br/>
Artemisiaceae <small>Martinov</small><br/>
Athanasiaceae <small>Martinov</small><br/>
Calendulaceae <small>Bercht. & J. Presl</small><br/>
Carduaceae <small>Bercht. & J. Presl</small><br/>
Cassiniaceae <small>Sch. Bip.</small><br/>
Cichoriaceae <small>Juss.</small><br/>
Coreopsidaceae <small>Link</small><br/>
Cynaraceae <small>Spenn.</small><br/>
Echinopaceae <small>Bercht. & J. Presl</small><br/>
Eupatoriaceae <small>Bercht. & J. Presl</small><br/>
Helichrysaceae <small>Link</small><br/>
Inulaceae <small>Bercht. & J. Presl</small><br/>
Lactucaceae <small>Drude</small><br/>
Mutisiaceae <small>Burnett</small><br/>
Partheniaceae <small>Link</small><br/>
Perdiciaceae <small>Link</small><br/>
Senecionaceae <small>Bercht. & J. Presl</small><br/>
Vernoniaceae <small>Burmeist.</small><br/><br/>
Sources: UniProt<ref name="UniProt">{{UniProt Taxonomy
| name = Asteraceae
| id = 4210
| accessdate = 2008-06-12
}}</ref> GRIN<ref name="GRIN">{{cite web
| url = http://www.ars-grin.gov/cgi-bin/npgs/html/family.pl?110
| title = Family: Asteraceae Bercht. & J. Presl, nom. cons.
| accessdate = 2008-06-12
| author = Germplasm Resources Information Network (GRIN)
| authorlink = Germplasm Resources Information Network
| date = 2007-04-13
| format = HTML
| work = Taxonomy for Plants
| publisher = [[United States Department of Agriculture|USDA]], [[Agricultural Research Service|ARS]], National Genetic Resources Program, National Germplasm Resources Laboratory, Beltsville, Maryland
}}</ref>
}}
The family '''Asteraceae''' or '''Compositae''' (known as the '''aster''', '''daisy''', or '''sunflower family''') is the largest [[family]] of [[flowering plant]]s, in terms of number of species.
The name 'Asteraceae' is derived from the type genus ''[[Aster (genus)|Aster]]'', while 'Compositae', an older but still valid<ref>[http://ibot.sav.sk/icbn/main.htm International Code of Botanical Nomenclature]</ref> name, means ''composite'' and refers to the characteristic [[inflorescence]], a special type of [[pseudanthium]] found in only a few other angiosperm families. The study of this family is known as [[synantherology]].
According to the [[Royal Botanical Gardens]] of Kew the family comprises more than 1,600 genera and 23,000 species. The largest genera are ''[[Senecio]]'' (1,500 species), ''[[Vernonia]]'' (1,000 species), ''[[Cousinia]]'' (600 species), <!-- ''[[Eupatorium]]'' (600 species), [not anymore!] -->''[[Centaurea]]'' (600 species). The circumscription of the genera is often problematic and some of these have been frequently divided into minor subgroups<ref name="Judd">Judd & al., Plant Systematics: A Phylogenetic Approach</ref>.
Asteraceae are cosmopolitan, but most common in the temperate regions and tropical mountains.
==Taxonomy==
The family has been universally recognised and placed in the order [[Asterales]].
Traditionally two subfamilies were recognised: [[Asteroideae]] (or 'Tubuliflorae') and [[Cichorioideae]] (or 'Liguliflorae'). The latter is [[paraphyletic]] and has been divided into many minor groups in most newer systems. The phylogenetic tree presented below is based on Panero & Funk (2002) and also shown in the [[APG system]].
A tentative [[cladogram]] is shown below. The diamond denotes a very poorly supported branching (<50%), the dot a poorly supported branching (<80%)<ref name="APW">Stevens, P.F., [http://www.mobot.org/MOBOT/research/APweb/orders/asteralesweb.htm'' Angiosperm Phylogeny Website'']</ref>.
{{clade|style=font-size:80%;line-height:75%
|1={{clade|1='''[[Barnadesioideae]]''': 9 genera, 93 species. [[South America]], mainly the [[Andes]].
|2={{clade|1='''[[Stifftioideae]]''': [[South America]] and [[Asia]].|label2=♦
|2={{clade|1='''[[Mutisioideae]]''': 58 genera, 750 species. [[South America]].|label2=•
|2={{clade|1='''[[Wunderlichioideae]]''':8 genera, 24 species, mostly in [[Venezuela]] and [[Guiana]]
|2={{clade|1='''[[Gochnatioideae]]''': 4 or 5 genera, 90 species.|2={{clade
|1='''Hecastocleidoideae''': Only ''[[Hecastocleis]] shockleyi''. South-western [[U.S.A.]].|2={{clade
|1='''[[Carduoideae]]''': 83 genera, 2,500 species. Worldwide.|2={{clade
|1='''[[Pertyoideae]]''': 5 or 6 genera, 70 species.
|2={{clade|1='''[[Gymnarrhenoideae]]''': Only ''[[Gymnarrhena micrantha]]''. Northern [[Africa]].|label2=•
|2={{clade|1='''[[Cichorioideae]]''': 224 genera, 3,200 species. Worldwide.|2={{clade
|1='''[[Corymbioideae]]''': Only the genus ''[[Corymbium]]'', with 7 species.|label2=•|2={{clade
|1='''[[Asteroideae]]''': 1,130 genera and 16,200 species. Worldwide.}}}}}}}}}}}}}}}}}}}}}}}}}}
It is noteworthy that the four subfamilies [[Asteroideae]], [[Cichorioideae]], [[Carduoideae]] and [[Mutisioideae]] comprise 99% of the specific diversity of the whole family (appr. 70%, 14%, 11% and 3% respectively).
== Description ==
Asteraceae are most usually [[herb]]s, but some shrubs, [[tree]]s and [[Vine|climber]]s do exist. They are generally easy to distinguish, mainly because of their characteristic [[inflorescence]] and many shared [[apomorphy|apomorphies]]<ref name="Judd"/>.
===Leaves and stems===
The [[leaf|leaves]] and the stems very often contain secretory canals with [[resin]] or [[latex]] (particularly common among the Cichorioideae). The leaves can be [[phyllotaxis|alternate]], [[phyllotaxis|opposite]], or [[phyllotaxis|whorled]]. They may be simple, but are often deeply [[leaf#morphology|lobed]] or otherwise [[leaf#morphology|incised]], often [[leaf#morphology|conduplicate]] or [[leaf#morphology|revolute]]. The margins can be entire or dentate.
===Flowers===
[[Image:Bidens flwr.jpg|thumb|A typical Asteraceae flower head (here ''[[Bidens torta]]'') showing the individual flowers.]]
The most evident characteristic of Asteraceae is perhaps their [[inflorescence]]: a specialised ''[[capitulum]]'', technically called a ''calathid'' or ''calathidium'', but generally referred to as ''flower head'' or, alternatively, simply ''capitulum''<ref>Usher, 1966</ref>. The ''capitulum'' is a contracted [[raceme]] composed of numerous individual sessile [[flower]]s, called the ''florets'', all sharing the same [[receptacle]].
The capitulum of the Asteraceae has evolved many characteristics that make it look superficially like a big single flower. This kind of flower-like inflorescences are quite widespread amongst plants and have been given the name of ''[[pseudanthium|pseudanthia]]''.
Many [[bract]]s form an [[involucre]] under the basis of the capitulum; these are called "phyllaries", or "involucral bracts". They may simulate the [[sepal]]s of the pseudanthium. These are mostly herbaceous but can also be brightly coloured (e.g. ''[[Helichrysum]]'') or have a scarious texture. The bracts 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 itself be subtended by a bract, called a "palea" or "receptacular bract". These bracts as a group are often called "[[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.
[[Image:Carduus flowerdiagram.png|thumb|left|Flower diagram of ''[[Carduus]]'' (Carduoideae)]]The florets have five [[petal]]s fused at the base to form a [[Corolla (flower)|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 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 [[petal]]s form the ligule, with the other two petals being inconspicuously small. The Cichorioidea 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 (botany)|calyx]] of the florets may be absent, but when present it is always modified into a [[pappus (flower structure)|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 five [[stamen]]s. The filaments are fused to the corolla, while the anthers are generally [[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, expelled with a sort of pump mechanism (''nüdelspritze'') or a brush.
The [[pistil]] is made of two connate [[carpel]]s. The [[carpel|style]] has two lobes; stigmatic tissue may be located in the interior surface or form two lateral lines. The [[ovary (plants)|ovary]] is inferior and has only one [[ovule]], with basal [[placentation]].
=== Fruits and seeds ===
[[Image:Kih-silberdistel.jpg|thumb|250px|right|Seeds are dispersed by the wind in ''[[Carlina]]'']]
The fruit of the Asteraceae is a specialised type of [[achene]], sometimes called ''cypsela'' (plural ''cypselae''). One [[seed]] per fruit is formed. It may sometimes be flat, winged or spiny and it adheres to the persistent pappus. Its [[morphology (biology)|morphology]] is often used to help determine plant relationships at the genus and species level<ref>{{cite journal | journal = Annals of the Missouri Botanical Garden | pages = 569–594 | url = http://apt.allenpress.com/aptonline/?request=get-abstract&issn=0026-6493&volume=092&issue=04&page=0569 | title = Morphology Of Cypselae In Subtribe Arctotidinae (Compositae–Arctotideae) And Its Taxonomic Implications | author = R. J. McKenzie, J. Samuel, E. M. Muller, A. K. W. Skinner, and N. P. Barker | volume = 92 | issue = 4 | date = December 2005 }}</ref>. The [[seed]]s usually have little or lack [[endosperm]]<ref name="Judd"/>.
=== Metabolites ===
Asteraceae generally store energy in the form of [[inulin]].
They produce iso/chlorogenic acid, [[sesquiterpene]] [[lactones]], pentacyclic triterpene alcohols, various [[alkaloid]]s, [[acetylene]]s (cyclic, aromatic, with vinyl end groups), [[tannin]]s. They have [[terpene|terpenoid]] [[essential oils]] which never contain [[iridoid]]s<ref name="APW"/>.
== Ecology ==
[[Image:Epizoochoria NRM.jpg|thumb|200px|Epizoochory in ''[[Bidens tripartita]]'']]
Asteraceae are especially common in open and dry environments<ref name="Judd"/>.
Many members of the Asteraceae are [[pollination|pollinated]] by insects, which explains their value in attracting [[beneficial insects]], but [[pollination syndrome|anemophyly]] is also present (e.g. ''[[Ragweed|Ambrosia]]'', ''[[Artemisia (genus)|Artemisia]]''). There are many [[apomixis|apomictic]] species in the family.
Seeds are ordinarily dispersed intact with the fruiting body, the cypsela. Wind dispersal is common (''[[Biological dispersal#wind|anemochory]]'') assisted by a hairy pappus. Another common variation is (''[[Biological dispersal#By animals|epizoochory]]''), in which the dispersal unit, a single cypsela (e.g. ''[[Bidens]]'') or entire capitulum (e.g. ''[[Arctium]]'') provided with hooks, spines or some equivalent structure, sticks to the fur or plumage of an animal (or even to clothes, like in the photo) just to fall off later far from its mother plant.
== Evolution ==
Diversification of Asteraceae may have been within 42-36 million years, the stem group perhaps being up to 49 million years old<ref name="APW"/>.
It is still unknown whether the precise cause of their great success was the development of the calathid, their ability to store energy as [[fructan]]s (mainly [[inulin]]), which is an advantage in relatively dry zones, or some combination of these and possibly other factors<ref name="APW"/>.
==Uses==
[[Image:Field of sunflowers.JPG|thumb|left|200px|[[Sunflower]]s are a commonly cultivated member of Asteraceae]]
Commercially important plants in the Asteraceae include the food crops ''[[Lactuca sativa]]'' (lettuce), ''[[Cichorium]]'' (chicory), ''[[Cynara scolymus]]'' (globe artichoke), ''[[Helianthus annuus]]'' (sunflower), ''[[Smallanthus sonchifolius]]'' (yacón), ''[[Carthamus tinctorius]]'' (safflower) and ''[[Helianthus tuberosus]]'' (Jerusalem artichoke).
Other commercially important species include Compositae used as herbs and in herbal teas and other beverages. Chamomile which comes from two different species, the annual ''[[Matricaria recutita]]'' or German chamomile, and the perennial ''[[Chamaemelum nobile]]'', also called Roman chamomile. ''[[Calendula]]'', also called the pot marigold is grown commercially for herbal teas and the potpourri industry. Echinacea (''[[Echinacea purpurea]]''), used as a medicinal tea. Winter tarragon, also called Mexican mint marigold, ''[[Tagetes lucida]]'' is commonly grown and used as a tarragon substitute in climates where tarragon will not survive. Finally, the wormwood genus ''[[Artemisia]]'' includes [[absinthe]] (''A. absinthium'') and [[tarragon]] (''A. dracunculus'').
Industrial use of Compositae is also known. Common in all commercial poultry feed, marigold (''[[Tagetes patula]]'') is grown primarily in Mexico. Marigold oil, extracted from ''[[Tagetes minuta]]'' is used by the metric ton in the cola and cigarette industry.
Plants in Asteraceae are medically important in areas that don't have access to Western medicine. They are also commonly featured in medical and phytochemical journals because the [[sesquiterpene]] [[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>Odom & al., 2000</ref> [[Pollen]] from Ragweed ''[[Ambrosia]]'' is among the main causes of so called [[hay fever]] in the United States<ref>[http://www.aafa.org/display.cfm?id=9&sub=19&cont=267 Ragweed Allergy]</ref>.
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 ''[[Chrysanthemum]]'', ''[[Gerbera]]'', ''[[Calendula]]'', ''[[Dendranthema]]'', ''[[Argyranthemum]]'', ''[[Dahlia]]'', ''[[Tagetes]]'', ''[[Zinnia]]'' and many others.
[[Image:Unidentified Pink Flower Closeup 2048px.jpg|thumb|right|200px|A ''[[Dahlia]] cultivar]]
Many members of Asteraceae are copious [[nectar (plant)|nectar]] producers and are useful for evaluating [[pollinator]] populations during their bloom. ''[[Centaurea]]'' (knapweed), ''[[Helianthus annuus]]'' (domestic sunflower), and some species of ''[[Solidago]]'' (goldenrod) are major "[[honey]] plants" for [[beekeeper]]s. ''Solidago'' produces relatively high protein [[pollen]], which helps [[honey bee]]s overwinter {{Fact|date=November 2007}}.
Some members of the Asteraceae are economically important as weeds. Notably in the United States are the ragwort, ''[[Senecio jacobaea]]'', groundsel ''[[Senecio vulgaris]]'' and ''[[Taraxacum]]'' (dandelion).
The genera ''[[Tanacetum]]'', ''[[Chrysanthemum]]'' and ''[[Pulicaria]]'' contain species with insecticidal properties.
''[[Parthenium argentatum]]'' (Guayule) is a source of [[hypoallergenic]] [[latex]].
==Genera==
{{main|List of Asteraceae genera}}
==Image gallery==
<gallery>
Image:Monticellogardenflower.jpg
Image:Aster-alpinus.JPG|''[[Aster alpinus]]''
Image:Golden Ragwort Senecio aureus Flowers 2616px.jpg|''[[Senecio aureus]]''
Image:Vernonia fasciculata.jpg|''[[Vernonia fasciculata]]''
Image:Koninginnenkruid A.jpg|''[[Eupatorium cannabinum]]''
Image:Chuquiraga oppositifolia.jpg|''[[Chuquiraga oppostifolia]]'' (Barnadesioideae)
Image:Carduus tenuiflorus1.jpg|''[[Carduus tenuiflorus]]'' (Carduoideae)
Image:Big yellow.jpg|''[[Gerbera]] sp.'' (Mutisoideae). Note the zygomorphic disc flowers.
Image:Pertya scandens2.jpg|''[[Pertya scandens]]'' (Pertyoideae)
Image:Cichorium intybus-alvesgaspar1.jpg|''[[Cichorium intybus]]'' (Cichorioideae). Note the dark blue anther tubes and the bilabiate styles.
Image:399px-IMG 5243aw.jpg|''[[Aster novae-angliae]] (Asteroideae)
Image:Ray-floret.svg|Ray floret (as in Chichorioideae, and the outer florets in Asteroideae)
Image:Disc floret01.jpg|Disc floret (as in Asteroideae)
Image:Artichaut2.jpg|The involucral leaves of the capitula of the [[globe artichoke]] (here flowering) are eaten as vegetable
Image:Tragopogon pratensis-Ejdzej-2006-B.jpg|The large seed head of ''[[Tragopogon pratensis]]'' is composed of many single achenes, each with its own pappus
Image:Bidens_dentes_NRM.jpg|Sticking teeth of a ''Bidens tripartita'' achene under stereomicroscope (30X).
Image:Taraxacum sect Ruderalia10 ies.jpg|The stem of the [[dandelion]] (''Taraxacum'', Cichorioidea) contains a bitter [[latex]].
</gallery>
== See also ==
*[[Terminology for Asteraceae]]
== Footnotes ==
{{reflist}}
== References ==
*Panero J. L. and V. A. Funk. 2002. [http://www.mnh.si.edu/biodiversity/bdg/Panero&Funk2002.pdf Toward a phylogenetic subfamilial classification for the Compositae (Asteraceae)]. Proc. Biol. Soc. Wash. 115: 909-922.
*Stevens, P. F. ''Angiosperm Phylogeny Website''
*{{cite book |last= Usher|first= George|title= A dictionary of botany, including terms used in bio-chemistry, soil science, and statistics|year= 1966|publisher= Van Nostrand|location= Princeton|id= {{LCCN|66|0|25447}}}}
* Judd & al. ''Plant Systematics: A Phylogenetic Approach
*http://ibot.sav.sk/icbn/main.htm International Code of Botanical Nomenclature (VIENNA CODE)
* [http://www.itis.usda.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=35420 ITIS report 2002-09-10]
* Walters, Dirk R. and David J. Keil (1996). ''Vascular plant taxonomy''. 4th ed. Kendall/Hunt Publishing Company. Dubuque, Iowa.
* Wagner,W.L., D.R. Herbst, and S.H. Sohmer. 1990. ''Manual of the Flowering Plants of Hawai‘i'', Vol. I. University of Hawaii Press, Honolulu. 988 pp.
* D. J. N. Hind, C. Jeffrey & G. V. Pope (eds.), Advances in Compositae systematics. - Royal Bot. Gardens, Kew, 469 pp., 1995
*{{cite book | title=Andrews' Diseases of the Skin: Clinical Dermatology| last=Odom| first=Richard B.| coauthors=William D. James, Timothy G. Berger| date=2000| pages=1135 pages| publisher=W.B. Saunders Company| id=ISBN 0721658326}}
==External links==
{{commonscat-inline}}
{{Wikispecies-inline}}
* {{dmoz|/Science/Biology/Flora_and_Fauna/Plantae/Magnoliophyta/Magnoliopsida/Asteraceae/}}
[[Category:Plant families]]
[[Category:Asteraceae]]
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