Asteraceae

The family Asteraceae, alternatively

Compositae, consists of over 32,000
known species of flowering plants in over
1,900 genera within the order Asterales.
Commonly referred to as the aster, daisy,
composite, or sunflower family,
Compositae were first described in 1740.
The number of species in Asteraceae is
rivaled only by the Orchidaceae, and
which is the larger family is unclear as
the quantity of extant species in each
family is unknown.
 Asteraceae
Temporal range: 76–0 Ma
PreꞒ Ꞓ O S D C P T J K PN

Campanian[1]
g
– recent

Twelve species of Asteraceae from the

subfamilies Asteroideae, Carduoideae and
Cichorioideae

Scientific classification

Kingdom: Plantae

Clade: Tracheophytes
Clade: Angiosperms

Clade: Eudicots

Clade: Asterids

Order: Asterales

Family: Asteraceae
Bercht. & J.Presl[2]

Type genus

Aster
L.

Subfamilies

Asteroideae Lindl.
Barnadesioideae K.Bremer &
R.K.Jansen
Carduoideae Sweet
Cichorioideae Chevall.
Corymbioideae Panero & Funk
Famatinanthoideae S.E.Freire, Ariza &
Panero
Gochnatioideae Panero & Funk
Gymnarrhenoideae Panero & Funk
Hecastocleidoideae Panero & Funk
Mutisioideae Lindl.
Pertyoideae Panero & Funk
Stifftioideae Panero
Wunderlichioideae Panero
[3] & Funk
Diversity

1,911 genera

Synonyms[4]

Compositae Giseke
Acarnaceae Link
Ambrosiaceae Bercht. & J.Presl
Anthemidaceae Bercht. & J.Presl
Aposeridaceae Raf.
Arctotidaceae Bercht. & J.Presl
Artemisiaceae Martinov
Athanasiaceae Martinov
Calendulaceae Bercht. & J.Presl
Carduaceae Bercht. & J.Presl
Cassiniaceae Sch.Bip.
Cichoriaceae Juss.
Coreopsidaceae Link
Cynaraceae Spenn.
Echinopaceae Bercht. & J.Presl
Eupatoriaceae Bercht. & J.Presl
 Helichrysaceae Link
Inulaceae Bercht. & J.Presl
Lactucaceae Drude
Mutisiaceae Burnett
Partheniaceae Link
Perdiciaceae Link
Senecionaceae Bercht. & J.Presl
Vernoniaceae Burmeist

Most species of Asteraceae are annual,

biennial, or 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
semi-desert climates, and they are found
on every continent but Antarctica. The
primary common characteristic is the
existence of sometimes hundreds of tiny
individual florets which are held together
by protective involucres in flower heads,
or more technically, capitula.

The oldest known fossils are pollen

grains from the Late Cretaceous
(Campanian to Maastrichtian) of
Antarctica, dated to ∼76–66 million years
(myr). It is estimated that the crown
group of Asteraceae evolved at least 85.9
myr (Late Cretaceous, Santonian) with a
stem node age of 88–89 myr (Late
Cretaceous, 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 invasive.

Description
Members of the Asteraceae are mostly
herbaceous plants, but some shrubs,
vines, and trees (such as Lachanodes
arborea) do exist. Asteraceae species are
generally easy to distinguish from other
plants because of their unique
inflorescence and other shared
characteristics.[5] However, determining
genera and species of some groups such
as Hieracium is notoriously difficult (see
"damned yellow composite" for
example).

Roots …

Members of the family Asteraceae

generally produce taproots, but
sometimes they possess fibrous root
systems. Some species have
underground stems in the form of
caudices or rhizomes. These can be
fleshy or woody depending on the
species.[6]

Stems …
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,[6] or latex
which is particularly common among the
Cichorioideae.[7]

Leaves …

Leaves can be alternate, opposite, or

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[6] or latex[7] also can be present in
the leaves.
Inflorescences …

Nearly all Asteraceae bear their flowers

in dense flower heads called capitula.
They are surrounded by 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 …
 A typical A flower head
Asteraceae showing the
flower head individual flowers
showing the opening from the
individual outside
flowers (Bidens (Argyranthemum
torta) 'Bridesmaid')

In plants of the family Asteraceae, what

appears to be a single flower is actually a
cluster of much smaller flowers. The
overall appearance of the cluster, as a
single flower, functions in attracting
pollinators in the same way as the
structure of an individual flower in some
other plant families. The older family
name, Compositae, comes from the fact
that what appears to be a single flower is
actually a composite of smaller
flowers.[8]

The "petals" or "sunrays" in a sunflower

head are actually individual strap-
shaped[9] flowers called ray flowers, and
the "sun disk" is made of smaller circular
shaped individual flowers called disc
flowers. The word "aster" means "star" in
Greek, referring to the appearance of
some family members, as a "star"
surrounded by "rays". The cluster of
flowers that may appear to be a single
flower, is called a head. The entire head
may move tracking the sun, like a "smart"
solar panel, which maximizes reflectivity
of the whole unit and can thereby attract
more pollinators.[8]

On the outside the flower heads are

small bracts that look like scales. These
are called phyllaries, and together they
form the involucre that protects the
individual flowers in the head before they
open.[8]:29 The individual heads have the
smaller individual flowers arranged on a
round or dome-like structure called the
receptacle. The flowers mature first at
the outside, moving toward the center,
with the youngest in the middle.[8]
The individual flowers in a head have 5
fused petals (rarely 4), but instead of
sepals, have threadlike, hairy, or bristly
structures called pappus, which surround
the fruit and can stick to animal fur or be
lifted by wind, aiding in seed dispersal.
The whitish fluffy head of a dandelion,
commonly blown on by children, is made
of the pappus, with tiny seeds attached
at the ends, whereby the pappus provides
a parachute like structure to help the
seed be carried away in the wind.[8]
Schemes and floral diagrams of the different flower
types (Leucanthemum vulgare agg.): a – disc
flower; b – ray flower. 1 – style with stigmas; 2 –
anthers; 3 – corolla (petals), in the ray flower three
petals are joined to form a strap (in other species, 5
petals can form a ligule); 4 – reduced calyx; 4’ – in
many other species (like Carduus acanthoides
shown in the circle), it forms a pappus; 5 – inferior
ovary fused of two carpels containing one abaxial
ovule (basal placentation).

A ray flower is a 3-tipped (3-lobed), strap-

shaped, individual flower in the head of
some members of the family
Asteraceae.[8][9] Sometimes a ray flower
is 2-tipped (2-lobed). The corolla of the
ray flower may have 2 tiny teeth opposite
the 3-lobed strap, or tongue, indicating
evolution by fusion from an originally 5-
part corolla. Sometimes, the 3:2
arrangement is reversed, with 2 tips on
the tongue, and 0 or 3 tiny teeth opposite
the tongue. A ligulate flower is a 5-tipped,
strap-shaped, individual flower in the
heads of other members.[8] A ligule is the
strap-shaped tongue of the corolla of
either a ray flower or of a ligulate
flower.[9] A disk flower (or disc flower) is
a radially symmetric (i.e., with identical
shaped petals arranged in circle around
the center) individual flower in the head,
which is ringed by ray flowers when both
are present.[8][9] Sometimes ray flowers
may be slightly off from radial symmetry,
or weakly bilaterally symmetric, as in the
case of desert pincushions Chaenactis
fremontii.[8]

A radiate head has disc flowers

surrounded by ray flowers. A ligulate
head has all ligulate flowers. When a
sunflower family flower head has only
disc flowers that are sterile, male, or have
both male and female parts, it is a
discoid head. Disciform heads have only
disc flowers, but may have two kinds
(male flowers and female flowers) in one
head, or may have different heads of two
kinds (all male, or all female). Pistillate
heads have all female flowers. Staminate
heads have all male flowers. Sometimes,
but rarely, the head contains only a single
flower, or has a single flowered pistillate
(female) head, and a multi-flowered male
staminate (male) head.[8]

Floral structures …

Flower diagram of 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

The distinguishing characteristic of

Asteraceae is their inflorescence, a type
of specialised, composite flower head or
pseudanthium, technically called a
calathium or capitulum,[10][11] that may
look superficially like a single flower. The
capitulum is a contracted raceme
composed of numerous individual
sessile flowers, called florets, all sharing
the same receptacle.

A set of bracts 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. 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 "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 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 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 stamens. The

filaments are fused to the corolla, while
the anthers are generally 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
ovule, with basal placentation.

Fruits and seeds …

In members of the Asteraceae the fruit is

achene-like, and is called a cypsela
(plural cypselae). Although there are two
fused carpels, there is only one locule,
and only one seed per fruit is formed. 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 Helianthus). Cypsela
morphology is often used to help
determine plant relationships at the
genus and species level.[12] The mature
seeds usually have little endosperm or
none.[5]

Pollen …

The pollen of composites is typically

echinolophate, a morphological term
meaning "with elaborate systems of
ridges and spines dispersed around and
between the apertures."[13]

Metabolites …

In Asteraceae, the energy store is

generally in the form of inulin rather than
starch. They produce iso/chlorogenic
acid, sesquiterpene lactones, pentacyclic
triterpene alcohols, various alkaloids,
acetylenes (cyclic, aromatic, with vinyl
end groups), tannins. They have
terpenoid essential oils which never
contain iridoids.[14]

Asteraceae produce secondary

metabolites, such as flavonoids and
terpenoids. Some of these molecules
can inhibit protozoan parasites such as
Plasmodium, Trypanosoma, Leishmania
and parasitic intestinal worms, and thus
have potential in medicine.[15]

Taxonomy

History …

Compositae, the original name for

Asteraceae, were first described in 1740
by Dutch botanist Adriaan van
Royen.[16]:117–118 Traditionally, two
subfamilies were recognised:
Asteroideae (or Tubuliflorae) and
Cichorioideae (or Liguliflorae). The latter
has been shown to be extensively
paraphyletic, and has now been divided
into 12 subfamilies, but the former still
stands.[17] The study of this family is
known as synantherology.

Phylogeny …

The phylogenetic tree presented below is

based on Panero & Funk (2002)[17]
updated in 2014,[18] and now also
includes the monotypic
Famatinanthoideae.[18][19][20] The
diamond ( ♦) denotes a very poorly
supported node (<50% bootstrap
support), the dot (•) a poorly supported
node (<80%).[14]
    Barnadesioideae: 9 genera, 93 species.
    South America, mainly the Andes.
  Famatinanthoideae: South America, 1
    genus, 1 species Famatinanthus
  decussatus.

  Mutisioideae: 58 genera, 750

    species. Absent from Europe,
  mostly in South America.

  Stifftioideae: 10 genera. South

  America.
♦    Wunderlichioideae: 8 genera, 24
species. Mostly in Venezuela and
  Guyana.

  Gochnatioideae: 4 or 5 genera,
    90 species. Latin America and
  southern United States.

  Hecastocleidoideae: Only
    Hecastocleis shockleyi.
  Southwestern United States.

  Carduoideae: 83 genera,
    2,500 species. Worldwide.
 
   
    Pertyoideae: 5 or 6
genera, 70 species.
Asia.

Gymnarrhenoideae:
Two genera/species,
Gymnarrhena
  micrantha (Northern
  Africa, Middle East)
and Cavea tanguensis
(Eastern Himalayas).

Cichorioideae: 224
  genera, 3,200
  species.
Worldwide.
 
 
Corymbioideae:
Only the genus
•   Corymbium, with
    9 species. Cape
provinces, South
  Africa.
 
Asteroideae:
• 1,130 genera and
  16,200 species.
Worldwide.
The family includes over 32,000 currently
accepted species, in over 1,900 genera
(list) in 13 subfamilies.[3] The number of
species in the family Asteraceae is
rivaled only by Orchidaceae.[14][21] Which
is the larger family is unclear, because of
the uncertainty about how many extant
species each family includes. The four
subfamilies Asteroideae, Cichorioideae,
Carduoideae and Mutisioideae contain
99% of the species diversity of the whole
family (approximately 70%, 14%, 11% and
3% respectively).

Because of the 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.[5]

Evolutionary processes …

The oldest known fossils of members of

Asteraceae are pollen grains from the
Late Cretaceous of Antarctica, dated to
∼76–66 myr (Campanian to
Maastrichtian) and assigned to the
extant genus Dasyphyllum.[1] Barreda, et
al. (2015) estimated that the crown
group of Asteraceae evolved at least 85.9
myr (Late Cretaceous, Santonian) with a
stem node age of 88–89 myr (Late
Cretaceous, Coniacian).[1]
It is still unknown whether the precise
cause of their great success was the
development of the highly specialised
capitulum, their ability to store energy as
fructans (mainly inulin), which is an
advantage in relatively dry zones, or
some combination of these and possibly
other factors.[14] 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 are adapted to different
environments, increasing chances of
survival.[22]

Etymology and pronunciation …

The name Asteraceae
(English: /ˌæstəˈreɪsi, -siˌaɪ, -siˌeɪ, -siˌiː/)
comes to international scientific
vocabulary from New Latin, from Aster,
the type genus, + -aceae,[23] a
standardized suffix for plant family
names in modern taxonomy. The genus
name comes from the Classical Latin
word aster, "star", which came from
Ancient Greek ἀστήρ (astḗr), "star".[23] It
refers to the star-like form of the
inflorescence.

The original name Compositae is still

valid under the International Code of
Nomenclature for algae, fungi, and
plants.[24] It refers to the "composite"
nature of the capitula, which consist of a
few or many individual flowers.

The vernacular name daisy, widely

applied to members of this family, is
derived from the Old English name of the
daisy (Bellis perennis): dæġes ēaġe,
meaning "day's eye". This is because the
petals open at dawn and close at dusk.

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 Brazil, the
Mediterranean, the Levant, southern
Africa, central Asia, and southwestern
China).[21] The largest proportion of the
species occur in the arid and semi-arid
regions of subtropical and lower
temperate latitudes.[6] The Asteraceae
may represent as much as 10% of
indigenous flora in many regions of the
world.

Ecology
 Anemochory in Epizoochory in
Carlina Bidens
tripartita

Asteraceae are especially common in

open and dry environments.[5] Many
members of Asteraceae are pollinated by
insects, which explains their value in
attracting beneficial insects, but
anemophily is also present (e.g.
Ambrosia, Artemisia). There are many
apomictic species in the family.

Seeds are ordinarily dispersed intact with

the fruiting body, the cypsela.
Anemochory (wind dispersal) is common,
assisted by a hairy pappus. Epizoochory
is another common method, in which the
dispersal unit, a single cypsela (e.g.
Bidens) or entire capitulum (e.g. 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 Senecio
jacobaea (ragwort),[25] Senecio vulgaris
(groundsel),[26] and Taraxacum
(dandelion).[27] Some are invasive
species in particular regions, often
having been introduced by human
agency. Examples include various
tumbleweeds, Bidens, ragweeds, thistles,
and dandelion. Dandelion was introduced
into North America by European settlers
who used the young leaves as a salad
green.[28]

Uses

The twining succulent, Senecio angulatus, is used

for its cut flowers,[29] despite being an invasive
weed in some places, such as Victoria, Australia
and New Zealand.[30]
Asteraceae is an economically important
family, providing products such as
cooking oils, leaf vegetables like lettuce,
sunflower seeds, artichokes, sweetening
agents, coffee substitutes and herbal
teas. Several genera are of horticultural
importance, including pot marigold
(Calendula officinalis), Echinacea
(coneflowers), various daisies, fleabane,
chrysanthemums, dahlias, zinnias, and
heleniums. Asteraceae are important in
herbal medicine, including Grindelia,
yarrow, and many others.

Commercially important plants in

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).

Plants are used as herbs and in herbal

teas and other beverages. Chamomile,
for example, comes from two different
species: the annual Matricaria
chamomilla (German chamomile) and the
perennial Chamaemelum nobile (Roman
chamomile). Calendula (known as pot
marigold) is grown commercially for
herbal teas and potpourri. Echinacea is
used as a medicinal tea. The wormwood
genus Artemisia includes absinthe (A.
absinthium) and tarragon (A.
dracunculus). Winter tarragon (Tagetes
lucida), is commonly grown and used as
a tarragon substitute in climates where
tarragon will not survive.

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.[31]
Senecio madagascariensis (Fireweed) is an
environmental weed in Australia, growing in
wastelands, grasslands and suburban bushland.[32]

Many species of this family possess

medicinal properties and are used as
traditional antiparasitic medicine.[15]

Members of the family 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.[33] Pollen
from ragweed Ambrosia is among the
main causes of so-called hay fever in the
United States.[34]

Asteraceae are also used for some

industrial purposes. French Marigold
(Tagetes patula) is common in
commercial poultry feeds and its oil is
extracted for uses in cola and the
cigarette industry.[31] The genera
Chrysanthemum, Pulicaria, Tagetes, and
Tanacetum contain species with useful
insecticidal properties.[31] Parthenium
argentatum (guayule) is a source of
hypoallergenic latex.[31]

Several members of the family are

copious nectar producers[31] 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 beekeepers.
Solidago produces relatively high protein
pollen, which helps honey bees over
winter.[35]

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External links
 Media related to Asteraceae at
Wikimedia Commons
Data related to Asteraceae at
Wikispecies
Asteraceae at the Angiosperm
Phylogeny Website
Compositae.org – Compositae
Working Group (CWG) and Global
Compositae Database (GCD)

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