The
Mayweeds and Chamomiles encompass several genera of superficially
similar plants which all belong to the
daisy family (Asteraceae). They are common weeds,
but closer study reveals them to be an interesting group of
plants. The most used species to make chamomile tea are Matricaria chamomilla and Chamaemelum
nobile
(sometimes described as the 'true chamomile'). Above: Tripleurospermum
inodorum.
These genera are (according to Stace 3rd edition): Tripleurospermum, Matricaria, Chamaemelum and Anthemis.
Which species is shown in the above photograph? It is very hard to
be certain without looking at the achenes (fruit)
under the microscope, but I suspect that this plant is either Chamaemelum nobile or Matricaria
chamomilla,
and
probably the former. The receptacle appears too conical for Tripleurospermum. After seeing hundreds
of
specimens, it is often possible to tell by looking at the overall
appearance of the plant, or by close examination of the
leaves, though this can lead to errors and microscopical
examination of the achenes, along with the receptacle
scales is the only way to be certain. Consideration of the habitat
can also help.
The following are the main species normally found in the British
Isles:
Tripleurospermum
inodorm
(Scentless Mayweed)
Tripleurospermum
maritimum
(Sea Mayweed)
Matricaria
chamomilla
(Scented Mayweed)
Matricaria
discoidea
(Pineappleweed)
Chamaemelum
nobile
('true' chamomile)
(Chamaemelum
mixtum
is a scarce alien)
Anthemis
cotula
(Stinking Chamomile)
Anthemis
arvensis
(Corn Chamomile)
Anthemis
tinctoria
(Yellow Chamomile)
Anthemis
punctata
(Sicilian Chamomile)
Anthemis
austriaca
(Austrian Chamomile)
There
are no receptacle scales in Tripleurospermum and Matricaria. In
Chamaemelum a receptacle scale
closely ensheaths each floret. In Anthemis
the morphology of the receptacle scales help distinguish the
various species.
The receptacle is hollow in Matricaria, and solid to
indistinctly hollow in
Tripleurospermum and much more conical in
Matricaria and Chamaemelum
than in Tripleurospermum.
Left: a ray floret with the two-lobed stigma. The ray florets
appear to be
female, as is generally the case in Asteraceae, and produce
full-looking (and
hence presumably fertile) achenes (these characteristics need
verifying for
each individual species). The ligules are white, except in Anthemis tinctoria in
which they are yellow, and the disc florets yellow. The ligule
typically has two
or three teeth at its tip and in Asteraceae is often derived from
three of the
corolla lobes, the other two being suppressed. The ligule
functions to attract
pollinating insects.
Above: a
tubular / disc floret. The corolla is yellow and when mature the
five lobes
open out (the petals being fused further down). On the right-hand
flower the style
bearing the bilobed stigma can be seen emerging from the cylinder
of the five anthers.
Above: a
capitulum of Tripleurospermum
inodorum.
The florets are arranged in a spiral on the disc-like
receptacle. The disc florets mature from the base towards the
apex. Here, the basal florets are already
opening their corolla lobes (December, Kent, N. Hemisphere -
unusually late flowering for this species
due to a mild winter and urban location).
Above: a
ray floret of Tripleurospermum
inodorum.
Note the
two stigma lobes borne on the emerging style. Right: a
phyllary from the same plant: note the central green region
and the typical brown scarious (thin, dry and membranous)
margin.
Above:
detail of disc florets of Tripleurospermum
inodorum.
The style bearing the bilobed stigmas can be
seen emerging in some of the open florets, carrying pollen with
it. This makes the pollen more easily
accessible to pollinating insects. Mayweeds are probably largely
self-incompatible, and the pollen is abaxial
(on the outer or underside of the spreading stigma lobes) not on
the receptive upper (adaxial) surface.
Furthermore, the stigma will not become receptive until this
pollen is removed. These mechanisms help
ensure cross-pollination.
Left:
close-up view of part of a leaf of
Tripleurospermum
inodorum.
The leaves are
compound, greatly divided up into narrow
branching leaflets, giving the leaves a
variously feathery appearance. Note the
central gutters which allow rainwater to drain,
keeping the leaves clean and dry so as not to
impede photosynthesis, as well as funneling
water down the stem to the roots. Although
the leaves of Tripleurospermum
maritimum
are sometimes similar, they are usually
fleshier and more succulent, being more
xerophytic and tolerant to dehydration in line
with Tripleurospermum
maritimum's
preferred
habitat on beaches and coastlines.
Tripleurospermum
inodorum
can occur
coastally, but is usually on more arable fields
slightly further from the shoreline, however,
their are places where both species occur
and hybridisation is known. Hybrids have
'intermediate' leaf characteristics.
A useful
(though by no means
foolproof) diagnostic feature is
the tip of the leaf segments,
which often have a hyaline
(translucent) spine at the tip in
Tripleurospermum
inodorum.
These are usually absent in
Tripleurospermum
maritimum
or
less frequent and less prominent
(shorter).
Tripleurospermum
inodorum
also typically has a very different habit (growth form) than Tripleurospermum
maritimum.
T. inodorum is annual and so often
retains the light-weight primary stem, whilst T.
maritimum
is
perennial (or occasionally bienniel) and so, except when in its
first year, acquires a woody stem (and can even
resemble a small bonsai tree!). T.
inodorum
is more upright, whereas T.
maritimum
is often at least partly
procumbent (lying horizontal). Together with its fleshier leaves,
this gives T.
maritimum
a more scrambling and
more compact appearance whereas T.
inodorum
tends to be more upright and open with more delicate leaves.
The stems of T.
maritimum
are also often darker and redder stems (though T.
inodorum
can have reddish
stems), especially when growing along the southern coast of
Britain. Those with obviousluy red stems are
considered to be the subspecies vinicaule. More northern
populations give way to the classic form (subspecies
maritimum) and then to subspecies nigriceps (Northern Scotland and
the Shetland isles) which has broader
and darker scarious margins on its phyllaries. There are
gradations, however, for example many
Tripleurospermum
maritimum
specimens have brown or dark brown stems which grade into the very
red stems
characteristic of subspecies vinicaule.
Despite
the aforementioned differences, mistakes can happen when
determining Tripleurospermum
maritimum
from Tripleurospermum
inodorum.
A more definite character is found by looking at the
achenes. In the photomicrograph above the achenes are still
immature and a very pale apple-green
colour. However, each bears a pair of circular oil glands which
determines this plant as
Tripleurospermum
inodorum.
The oil glands of Tripleurospermum
maritimum
are more than about 1.5
times as long as wide, and are often greatly elongated.
Above
and below: immature achenes of Tripleurospermum
inodorum
(with the flower tubes still attached).
Below:
another key difference in the achenes is seen when looking at the
back (adaxial
surface) of the achenes. The achenes of Tripleurospermum have three prominent
ribs at the
back, and in Tripleurospermum
maritimum
these ribs are very strongly developed with little
other surface tissue in-between and they join at the bottom of the
achene (generally). These
are achenes of Tripleurospermum
inodorum
which do not join at the base and leave wide
open strips of surface tissue between them. The achenes of T. maritimum are also generally
quite a nit larger when mature.
Above:
from the picture my guess is that this is Tripleurospermum
inodorum.
However, in order to check one could see whether the receptacles
are hollow, as they would be in Matricaria
chamomilla
(which is also usually more strongly and sweetly scented than T. inodorum). M. chamomilla
has a distinctively different look about it, but this is only
picked up with practice and may also depend on local populations.
The achenes of M.
chamomilla
also lack oil glands and have 4-5 weak ribs (not 3 strong ribs as
in Tripleurospermum). Chamaemelum nobile
is also typically strongly scented, but it has prominent
receptacle scales (visible under the microscope or with a hand
lens). Anthemis will also have
receptacle scales and the shape of the scales can distinguish Anthemis from Chamaemelum and also distinguish the
species in each case. At first the conical flower head
receptacles made me think of C. nobile, but the shape also
depends on maturity: mature receptacles are more conical. My final
determination for this specimen, taken into consideration
where it is growing, is Tripleurospermum inodorum.
The
capitula (flower-heads or pseudoflowers) are typical of the
Asteraceae: consisting of an enlarged receptacle,
subtended by phyllaries (small bracts) and
bearing a number of florets (small true flowers). The florets are
arranged in a helix around the receptacle; the angular positioning
of the florets follows a mathematical sequence
called a Fibonacci series. The phyllaries form an involucre (cup). The basal
(outermost) ray florets usually bear
ligules (a ligule is a
petal-like extensions of the corolla lobe) except in Matricaria discoidea and occasional ligueless
forms of the other species. The remaining disc florets have
cylindrical (radially symmetrical) corollas (floral tubes)
formed by the fusion of the 5 petals which are still separate as 5
lobes at the corolla tip, except in Matricaria
discoidea
in which it has only 4 lobes. Typical of Asteraceae the ovule in
each ovary is erect (vertically upright) and
anatropous (inverted or upside-down with the micropyle pointing
downwards). The ovary of each floret develops into
an achene fruit. In the Mayweeds
and Chamomiles, however, there is no pappus of sepals (except
where the
pappus remains as merely a short rim, as in Tripleurospermum). As in other Asteraceae
the 5 anthers are introrse
(turned inwards) and united via their cuticles such that they form
a cylinder into which the anthers open to shed their
pollen. As in most Asteraceae, they are protandrous (the male organs ripen
before the female organs). The pollen
is then pushed out by the piston-like style which later grows up
slowly inside the anther tube. This secondarily
presents pollen to the outside, where it is more accessible to
insect pollinators. When this pollen has gone, the two
stigma lobes expand and become receptive. Some Asteracea may later
elf-fertilise (if cross-pollination has been
unsuccessful) as the floret wilts. However, Anthemis
cotula
and Anthemis
arvensis,
at least, are known to be
self-incompatible.
Article
created: 1 Jan 2016
Article updated:
12 Aug 2017
11 Jun 2019
Tripleurospermum
maritimum
gallery
Note the
sprawling prostrate habit which is typical of T.
maritimum,
especially when growing on
exposed coastlines. Individual plants can be quite large and are
almost subshrubs.
A very
prostrate individual with the flowering stems at the periphery.
This is an adaptation to
resist high salty sea winds which can easily desiccate exposed
plants.
Notice
that each floret on the capitulum has five corolla-lobes (click to
enlarge).
Note the
succulent leaflets that make up the highly-divided leaves. This is
an adaptation to the
drying effects of sea salt: the leaves store water and are
thickened and so less easily
dehydrated by sea salt and sea winds.
Note the
woody perennial stem. The stems of Tripleurospermum
maritimum
growing on the
southern coasts of the British Isles tend to develop more red
anthocyanin pigment, which has
been shown to be both a genetic trait and an adaptation to the
growing conditions, probably to
high irradiance, since anthocyanins have a protective effect
against UV light. Those with the
reddest stems are designated subspecies vinicaule, those with the usual
brownish stems are
subspecies maritimum. Plants fitting the
description of both these subspecies often co-exist in
the same locale.
The sea
breeze was ruffling the ligules on these capitula.
These
attractive plants were growing on a shingle bank about 15 m behind
the seaweed line on the
south-western coast of the UK (VC 15).
Tripleurospermum
inodorum
(Scentless Mayweed) gallery
Tripleurospermum inodorum, the Scentless Mayweed, is an
annual found flowering mainly in the Summer, but sometimes
continuing over Winter. Sometimes they overwinter as a basal rosette
in some Scottish populations, and some may continue flowering over
winter in Kent, England, particularly during mild winters. The
habitat is distinct from that of the Sea Mayweed, T. inodorum
is often found growing on roadsides, the edges of arable fields. It
is a ruderal plant, being one of the first to colonise
disturbed ground. Those nera the coast may hybridise with Sea
Mayweed. Since the latter is perennial, this may produce
overwintering hybrids (which may be biennial or perennial?)
according to Kay (1994). However, winter flowering specimens I have
examined near coastal salt marshes in Kent were pure T. inodorum.
These probably represent late stragglers surviving a mild winter.The
habit is usually more upright than in Sea Mayweed and the stems are
not woody (though they do toughen as the plant matures).
Matricaria
discoidea
(Pineapple Weed) gallery
Pineappleweed is a rayless mayweed: the outer florets lack
the petal-like ligules and so are discoid rather than rayed. This is
subspecies discoidea. This plant is a native of North
America, but introduced in the British Isles. In both locations,
however, its a ruderal plant growing on disturbed ground. It is
often found in the British Isles on trampled ground such as along
tracks and paths. When crushed the plant gives off the pleasant
scent of pineapple, hence its name.
Specimens growing out from pavements, where free soil surface is
limited, tend to be less sprawling and more upright, rather like
miniature trees. They are, however, annuals.
Achenes of Matricaria discoidea subsp. discoidea.
Note the resinous glandular stripes (of which there is usually one
lateral pair) visible as brownish-green lines (these appear red in
fully matured achenes). These glands contain essential oil.
Matricaria
chamomilla
(Scented Mayweed) gallery
Matricaria chamomilla is probably the mayweed most
often used in chamomile tea. Note that the capitula are very
distinctly different in appearance from those of Tripleurospermum
inodorum as the ligules are shorter and blunter and the
capitula (flower heads) are smaller. This difference has proven
consistent, at least in the populations I have seen in Kent,
although I always check by close inspection of the achenes which
lack the oil sacs and thick ribs of Tripleurospermum. This
plant often grows on the edges of arable fields, alongside T.
inodorum.
Above: Matricharia chamomilla achenes. These lack the
glandular stripes of Pineappleweed achenes, but they do have rows of
enlarged mucilage secreting cells also called myxogenic
cells. These cells release their mucilage when moist, the
mucilage swelling as it absorbs water and this moisture collecting
ability helps the seed inside each achene to acquire enough water
for germination in dry habitats.
Above: A sample of chamomile tea as seen under the microscope. The
tea contains mainly fragments of chamomile flowers, including this
achene.