Groups (APG IV)
Takht. & W.Zimm.
The flowering plants, also known as angiosperms, Angiospermae or
Magnoliophyta, are the most diverse group of land plants, with 416
families, approximately 13,164 known genera and c. 295,383 known
species. Like gymnosperms, angiosperms are seed-producing plants.
However, they are distinguished from gymnosperms by characteristics
including flowers, endosperm within the seeds, and the production of
fruits that contain the seeds. Etymologically, angiosperm means a
plant that produces seeds within an enclosure; in other words, a
fruiting plant. The term comes from the Greek words angeion ("case" or
"casing") and sperma ("seed").
The ancestors of flowering plants diverged from gymnosperms in the
Triassic Period, 245 to 202 million years ago (mya), and the first
flowering plants are known from 160 mya. They diversified extensively
during the Lower Cretaceous, became widespread by 120 mya, and
replaced conifers as the dominant trees from 100 to 60 mya.
1.1 Angiosperm derived characteristics
1.2 Vascular anatomy
1.3 Reproductive anatomy
2.1 History of classification
2.2 Modern classification
Flowering plant diversity
Fertilization and embryogenesis
Fruit and seed
6 See also
9.1 Articles, books and chapters
10 External links
Angiosperm derived characteristics
Angiosperms differ from other seed plants in several ways, described
in the table below. These distinguishing characteristics taken
together have made the angiosperms the most diverse and numerous land
plants and the most commercially important group to humans.[a]
Distinctive features of angiosperms
Flowers, the reproductive organs of flowering plants, are the most
remarkable feature distinguishing them from the other seed plants.
Flowers provided angiosperms with the means to have a more
species-specific breeding system, and hence a way to evolve more
readily into different species without the risk of crossing back with
related species. Faster speciation enabled the
Angiosperms to adapt to
a wider range of ecological niches. This has allowed flowering plants
to largely dominate terrestrial ecosystems.
Stamens with two pairs of pollen sacs
Stamens are much lighter than the corresponding organs of gymnosperms
and have contributed to the diversification of angiosperms through
time with adaptations to specialized pollination syndromes, such as
particular pollinators. Stamens have also become modified through time
to prevent self-fertilization, which has permitted further
diversification, allowing angiosperms eventually to fill more niches.
Reduced male parts, three cells
The male gametophyte in angiosperms is significantly reduced in size
compared to those of gymnosperm seed plants. The smaller size of
the pollen reduces the amount of time between pollination — the
pollen grain reaching the female plant — and fertilization. In
gymnosperms, fertilization can occur up to a year after pollination,
whereas in angiosperms, fertilization begins very soon after
pollination. The shorter amount of time between pollination and
fertilization allows angiosperms to produce seeds earlier after
pollination than gymnosperms, providing angiosperms a distinct
Closed carpel enclosing the ovules (carpel or carpels and accessory
parts may become the fruit)
The closed carpel of angiosperms also allows adaptations to
specialized pollination syndromes and controls. This helps to prevent
self-fertilization, thereby maintaining increased diversity. Once the
ovary is fertilized, the carpel and some surrounding tissues develop
into a fruit. This fruit often serves as an attractant to
seed-dispersing animals. The resulting cooperative relationship
presents another advantage to angiosperms in the process of dispersal.
Reduced female gametophyte, seven cells with eight nuclei
The reduced female gametophyte, like the reduced male gametophyte, may
be an adaptation allowing for more rapid seed set, eventually leading
to such flowering plant adaptations as annual herbaceous life-cycles,
allowing the flowering plants to fill even more niches.
In general, endosperm formation begins after fertilization and before
the first division of the zygote.
Endosperm is a highly nutritive
tissue that can provide food for the developing embryo, the
cotyledons, and sometimes the seedling when it first appears.
Cross-section of a stem of the angiosperm flax:
1. Pith, 2. Protoxylem, 3.
Xylem I, 4.
Phloem I, 5.
fibre), 6. Cortex, 7. Epidermis
The amount and complexity of tissue-formation in flowering plants
exceeds that of gymnosperms. The vascular bundles of the stem are
arranged such that the xylem and phloem form concentric rings.
In the dicotyledons, the bundles in the very young stem are arranged
in an open ring, separating a central pith from an outer cortex. In
each bundle, separating the xylem and phloem, is a layer of meristem
or active formative tissue known as cambium. By the formation of a
layer of cambium between the bundles (interfascicular cambium), a
complete ring is formed, and a regular periodical increase in
thickness results from the development of xylem on the inside and
phloem on the outside. The soft phloem becomes crushed, but the hard
wood persists and forms the bulk of the stem and branches of the woody
perennial. Owing to differences in the character of the elements
produced at the beginning and end of the season, the wood is marked
out in transverse section into concentric rings, one for each season
of growth, called annual rings.
Among the monocotyledons, the bundles are more numerous in the young
stem and are scattered through the ground tissue. They contain no
cambium and once formed the stem increases in diameter only in
Plant reproductive morphology
A collection of flowers forming an inflorescence.
The characteristic feature of angiosperms is the flower. Flowers show
remarkable variation in form and elaboration, and provide the most
trustworthy external characteristics for establishing relationships
among angiosperm species. The function of the flower is to ensure
fertilization of the ovule and development of fruit containing seeds.
The floral apparatus may arise terminally on a shoot or from the axil
of a leaf (where the petiole attaches to the stem). Occasionally, as
in violets, a flower arises singly in the axil of an ordinary
foliage-leaf. More typically, the flower-bearing portion of the plant
is sharply distinguished from the foliage-bearing or vegetative
portion, and forms a more or less elaborate branch-system called an
There are two kinds of reproductive cells produced by flowers.
Microspores, which will divide to become pollen grains, are the "male"
cells and are borne in the stamens (or microsporophylls). The "female"
cells called megaspores, which will divide to become the egg cell
(megagametogenesis), are contained in the ovule and enclosed in the
carpel (or megasporophyll).
The flower may consist only of these parts, as in willow, where each
flower comprises only a few stamens or two carpels. Usually, other
structures are present and serve to protect the sporophylls and to
form an envelope attractive to pollinators. The individual members of
these surrounding structures are known as sepals and petals (or tepals
in flowers such as
Magnolia where sepals and petals are not
distinguishable from each other). The outer series (calyx of sepals)
is usually green and leaf-like, and functions to protect the rest of
the flower, especially the bud. The inner series (corolla of petals)
is, in general, white or brightly colored, and is more delicate in
structure. It functions to attract insect or bird pollinators.
Attraction is effected by color, scent, and nectar, which may be
secreted in some part of the flower. The characteristics that attract
pollinators account for the popularity of flowers and flowering plants
While the majority of flowers are perfect or hermaphrodite (having
both pollen and ovule producing parts in the same flower structure),
flowering plants have developed numerous morphological and
physiological mechanisms to reduce or prevent self-fertilization.
Heteromorphic flowers have short carpels and long stamens, or vice
versa, so animal pollinators cannot easily transfer pollen to the
pistil (receptive part of the carpel). Homomorphic flowers may employ
a biochemical (physiological) mechanism called self-incompatibility to
discriminate between self and non-self pollen grains. In other
species, the male and female parts are morphologically separated,
developing on different flowers.
History of classification
From 1736, an illustration of Linnaean classification
The botanical term "Angiosperm", from the Ancient Greek
αγγείον, angeíon (bottle, vessel) and σπέρμα, (seed),
was coined in the form Angiospermae by Paul Hermann in 1690, as the
name of one of his primary divisions of the plant kingdom. This
included flowering plants possessing seeds enclosed in capsules,
distinguished from his Gymnospermae, or flowering plants with achenial
or schizo-carpic fruits, the whole fruit or each of its pieces being
here regarded as a seed and naked. The term and its antonym were
Carl Linnaeus with the same sense, but with restricted
application, in the names of the orders of his class Didynamia. Its
use with any approach to its modern scope became possible only after
1827, when Robert Brown established the existence of truly naked
ovules in the Cycadeae and Coniferae, and applied to them the name
Gymnosperms. From that time onward, as long as these
Gymnosperms were, as was usual, reckoned as dicotyledonous flowering
plants, the term Angiosperm was used antithetically by botanical
writers, with varying scope, as a group-name for other dicotyledonous
An auxanometer, a device for measuring increase or rate of growth in
In 1851, Hofmeister discovered the changes occurring in the embryo-sac
of flowering plants, and determined the correct relationships of these
to the Cryptogamia. This fixed the position of Gymnosperms as a class
distinct from Dicotyledons, and the term Angiosperm then gradually
came to be accepted as the suitable designation for the whole of the
flowering plants other than Gymnosperms, including the classes of
Dicotyledons and Monocotyledons. This is the sense in which the term
is used today.
In most taxonomies, the flowering plants are treated as a coherent
group. The most popular descriptive name has been Angiospermae
Anthophyta ("flowering plants") a second choice.
These names are not linked to any rank. The
Wettstein system and the
Engler system use the name Angiospermae, at the assigned rank of
Reveal system treated flowering plants as subdivision
Magnoliophytina (Frohne & U. Jensen ex Reveal, Phytologia 79: 70
1996), but later split it to Magnoliopsida, Liliopsida, and Rosopsida.
Takhtajan system and
Cronquist system treat this group at the rank
of division, leading to the name Magnoliophyta (from the family name
Dahlgren system and
Thorne system (1992) treat this
group at the rank of class, leading to the name Magnoliopsida. The APG
system of 1998, and the later 2003 and 2009 revisions, treat
the flowering plants as a clade called angiosperms without a formal
botanical name. However, a formal classification was published
alongside the 2009 revision in which the flowering plants form the
The internal classification of this group has undergone considerable
revision. The Cronquist system, proposed by
Arthur Cronquist in 1968
and published in its full form in 1981, is still widely used but is no
longer believed to accurately reflect phylogeny. A consensus about how
the flowering plants should be arranged has recently begun to emerge
through the work of the Angiosperm
Phylogeny Group (APG), which
published an influential reclassification of the angiosperms in 1998.
Updates incorporating more recent research were published as the APG
II system in 2003, the
APG III system in 2009, and the APG
IV system in 2016.
Traditionally, the flowering plants are divided into two groups,
Dicotyledoneae or Magnoliopsida
Monocotyledoneae or Liliopsida
which in the
Cronquist system are called
Magnoliopsida (at the rank of
class, formed from the family name Magnoliaceae) and
the rank of class, formed from the family name Liliaceae). Other
descriptive names allowed by Article 16 of the
Dicotyledones or Dicotyledoneae, and
Monocotyledoneae, which have a long history of use. In English a
member of either group may be called a dicotyledon (plural
dicotyledons) and monocotyledon (plural monocotyledons), or
abbreviated, as dicot (plural dicots) and monocot (plural monocots).
These names derive from the observation that the dicots most often
have two cotyledons, or embryonic leaves, within each seed. The
monocots usually have only one, but the rule is not absolute either
way. From a broad diagnostic point of view, the number of cotyledons
is neither a particularly handy nor a reliable character.
Recent studies, as by the APG, show that the monocots form a
monophyletic group (clade) but that the dicots do not (they are
paraphyletic). Nevertheless, the majority of dicot species do form a
monophyletic group, called the eudicots or tricolpates. Of the
remaining dicot species, most belong to a third major clade known as
the magnoliids, containing about 9,000 species. The rest include a
paraphyletic grouping of early branching taxa known collectively as
the basal angiosperms, plus the families
Monocot (left) and dicot seedlings
There are eight groups of living angiosperms:
Basal angiosperms (ANA: Amborella, Nymphaeales, Austrobaileyales)
Amborella, a single species of shrub from New Caledonia;
Nymphaeales, about 80 species, water lilies and Hydatellaceae;
Austrobaileyales, about 100 species of woody plants from various
parts of the world
Core angiosperms (Mesangiospermae)
Chloranthales, several dozen species of aromatic plants with toothed
Magnoliids, about 9,000 species, characterized by trimerous
flowers, pollen with one pore, and usually branching-veined
leaves—for example magnolias, bay laurel, and black pepper;
Monocots, about 70,000 species, characterized by trimerous
flowers, a single cotyledon, pollen with one pore, and usually
parallel-veined leaves—for example grasses, orchids, and palms;
Ceratophyllum, about 6 species of aquatic plants, perhaps most
familiar as aquarium plants;
Eudicots, about 175,000 species, characterized by 4- or 5-merous
flowers, pollen with three pores, and usually branching-veined
leaves—for example sunflowers, petunia, buttercup, apples, and oaks.
The exact relationship between these eight groups is not yet clear,
although there is agreement that the first three groups to diverge
from the ancestral angiosperm were Amborellales, Nymphaeales, and
Austrobaileyales. The term basal angiosperms refers to these three
groups. Among the remaining five groups (core angiosperms), the
relationship between the three broadest of these groups (magnoliids,
monocots, and eudicots) remains unclear. Zeng and colleagues (Fig. 1)
describe four competing schemes. Of these, eudicots and monocots
are the largest and most diversified, with ~ 75% and 20% of
angiosperm species, respectively. Some analyses make the magnoliids
the first to diverge, others the monocots.
Ceratophyllum seems to
group with the eudicots rather than with the monocots. The 2016
Phylogeny Group revision (APG IV) retained the overall
higher order relationship described in APG III.
Phylogeny of the flowering plants, as of
APG III (2009).
2. Example of alternative phylogeny (2010)
3. APG IV (2016)
Cladogram of the Angiosperm
Phylogeny Group (APG) IV
Amborellales Melikyan, Bobrov & Zaytzeva 1999
Nymphaeales Salisbury ex von Berchtold & Presl 1820
Takhtajan ex Reveal 1992
Chloranthales Mart. 1835
Canellales Cronquist 1957
Piperales von Berchtold & Presl 1820
Magnoliales de Jussieu ex von Berchtold & Presl 1820
Laurales de Jussieu ex von Berchtold & Presl 1820
Acorales Link 1835
Alismatales Brown ex von Berchtold & Presl 1820
Dioscoreales Brown 1835
Pandanales Brown ex von Berchtold & Presl 1820
Liliales Perleb 1826
Asparagales Link 1829
Arecales Bromhead 1840
Poales Small 1903
Zingiberales Grisebach 1854
Commelinales de Mirbel ex von Berchtold & Presl 1820
Ceratophyllales Link 1829
Ranunculales de Jussieu ex von Berchtold & Presl 1820
Proteales de Jussieu ex von Berchtold & Presl 1820
Takhtajan ex Cronquist 1981
Takhtajan ex Reveal 1996
Takhtajan ex Reveal 1992
Dilleniales de Candolle ex von Berchtold & Presl 1820
Saxifragales von Berchtold & Presl 1820
Vitales de Jussieu ex von Berchtold & Presl 1820
Zygophyllales Link 1829
Celastrales Link 1829
Oxalidales von Berchtold & Presl 1820
Malpighiales de Jussieu ex von Berchtold & Presl 1820
Fabales Bromhead 1838
Rosales von Berchtold & Presl 1820
Cucurbitales de Jussieu ex von Berchtold & Presl 1820
Fagales Engler 1892
Geraniales de Jussieu ex von Berchtold & Presl 1820
Myrtales de Jussieu ex von Berchtold & Presl 1820
Takhtajan ex Reveal 1993
Picramniales Doweld 2001
Sapindales de Jussieu ex von Berchtold & Presl 1820
Huerteales Doweld 2001
Malvales de Jussieu ex von Berchtold & Presl 1820
Brassicales Bromhead 1838
Berberidopsidales Doweld 2001
Santalales Brown ex von Berchtold & Presl 1820
Cornales Link 1829
Ericales von Berchtold & Presl 1820
Icacinales Van Tieghem 1900
Garryales Mart. 1835
Gentianales de Jussieu ex von Berchtold & Presl 1820
Solanales de Jussieu ex von Berchtold & Presl 1820
Boraginales de Jussieu ex von Berchtold & Presl 1820
Vahliales Doweld 2001
Lamiales Bromhead 1838
Aquifoliales Senft 1856
Escalloniales Mart. 1835
Asterales Link 1829
Bruniales Dumortier 1829
Apiales Nakai 1930
Takhtajan ex Reveal 1992
Dipsacales de Jussieu ex von Berchtold & Presl 1820
Evolutionary history of plants
Evolutionary history of plants § Flowers
Fossilized spores suggest that higher plants (embryophytes) have lived
on land for at least 475 million years. Early land plants
reproduced sexually with flagellated, swimming sperm, like the green
algae from which they evolved. An adaptation to terrestrialization was
the development of upright meiosporangia for dispersal by spores to
new habitats. This feature is lacking in the descendants of their
nearest algal relatives, the Charophycean green algae. A later
terrestrial adaptation took place with retention of the delicate,
avascular sexual stage, the gametophyte, within the tissues of the
vascular sporophyte. This occurred by spore germination within
sporangia rather than spore release, as in non-seed plants. A current
example of how this might have happened can be seen in the precocious
spore germination in Selaginella, the spike-moss. The result for the
ancestors of angiosperms was enclosing them in a case, the seed. The
first seed bearing plants, like the ginkgo, and conifers (such as
pines and firs), did not produce flowers. The pollen grains (male
Ginkgo and cycads produce a pair of flagellated,
mobile sperm cells that "swim" down the developing pollen tube to the
female and her eggs.
Malus sylvestris (crab apple)
The apparently sudden appearance of nearly modern flowers in the
fossil record initially posed such a problem for the theory of
Charles Darwin called it an "abominable mystery".
However, the fossil record has considerably grown since the time of
Darwin, and recently discovered angiosperm fossils such as
Archaefructus, along with further discoveries of fossil gymnosperms,
suggest how angiosperm characteristics may have been acquired in a
series of steps. Several groups of extinct gymnosperms, in particular
seed ferns, have been proposed as the ancestors of flowering plants,
but there is no continuous fossil evidence showing exactly how flowers
evolved. Some older fossils, such as the upper
have been suggested. Based on current evidence, some propose that the
ancestors of the angiosperms diverged from an unknown group of
gymnosperms in the
Triassic period (245–202 million years ago).
Fossil angiosperm-like pollen from the Middle
Ma) suggests an older date for their origin. A close relationship
between angiosperms and gnetophytes, proposed on the basis of
morphological evidence, has more recently been disputed on the basis
of molecular evidence that suggest gnetophytes are instead more
closely related to other gymnosperms.
The evolution of seed plants and later angiosperms appears to be the
result of two distinct rounds of whole genome duplication events.
These occurred at 319 million years ago and 192 million
years ago. Another possible whole genome duplication event at
160 million years ago perhaps created the ancestral line that led
to all modern flowering plants. That event was studied by
sequencing the genome of an ancient flowering plant, Amborella
trichopoda, and directly addresses Darwin's "abominable mystery."
Flowers and leaves of
Oxalis pes-caprae (Bermuda buttercup)
The earliest known macrofossil confidently identified as an
Archaefructus liaoningensis, is dated to about 125 million
years BP (the
Cretaceous period), whereas pollen considered to be
of angiosperm origin takes the fossil record back to about 130 million
years BP. However, one study has suggested that the early-middle
Jurassic plant Schmeissneria, traditionally considered a type of
ginkgo, may be the earliest known angiosperm, or at least a close
relative. In addition, circumstantial chemical evidence has been
found for the existence of angiosperms as early as 250 million years
ago. Oleanane, a secondary metabolite produced by many flowering
plants, has been found in
Permian deposits of that age together with
fossils of gigantopterids. Gigantopterids are a group of
extinct seed plants that share many morphological traits with
flowering plants, although they are not known to have been flowering
In 2013 flowers encased in amber were found and dated 100 million
years before present. The amber had frozen the act of sexual
reproduction in the process of taking place. Microscopic images showed
tubes growing out of pollen and penetrating the flower's stigma. The
pollen was sticky, suggesting it was carried by insects.
DNA analysis based on molecular systematics showed that
Amborella trichopoda, found on the Pacific island of New Caledonia,
belongs to a sister group of the other flowering plants, and
morphological studies suggest that it has features that may have
been characteristic of the earliest flowering plants.
The orders Amborellales, Nymphaeales, and
Austrobaileyales diverged as
separate lineages from the remaining angiosperm clade at a very early
stage in flowering plant evolution.
The great angiosperm radiation, when a great diversity of angiosperms
appears in the fossil record, occurred in the mid-Cretaceous
(approximately 100 million years ago). However, a study in 2007
estimated that the division of the five most recent (the genus
Ceratophyllum, the family Chloranthaceae, the eudicots, the
magnoliids, and the monocots) of the eight main groups occurred around
140 million years ago. By the late Cretaceous, angiosperms appear
to have dominated environments formerly occupied by ferns and
cycadophytes, but large canopy-forming trees replaced conifers as the
dominant trees only close to the end of the
Cretaceous 66 million
years ago or even later, at the beginning of the Tertiary. The
radiation of herbaceous angiosperms occurred much later. Yet, many
fossil plants recognizable as belonging to modern families (including
beech, oak, maple, and magnolia) had already appeared by the late
It has been proposed that the swift rise of angiosperms to dominance
was facilitated by a reduction in their genome size. During the early
Cretaceous period, only angiosperms underwent rapid genome downsizing,
while genome sizes of ferns and gymnosperms remained unchanged.
Smaller genomes–and smaller nuclei–allow for faster rates of cell
division and smaller cells. Thus, species with smaller genomes can
pack more, smaller cells–in particular veins and stomata–into a
given leaf volume. Genome downsizing therefore facilitated higher
rates of leaf gas exchange (transpiration and photosynthesis) and
faster rates of growth. This would have countered some of the negative
physiological effects of genome duplications, facilitated increased
uptake of carbon dioxide despite concurrent declines in atmospheric
CO2 concentrations, and allowed the flowering plants to outcompete
other land plants.
Two bees on a flower head of Creeping Thistle, Cirsium arvense
It is generally assumed that the function of flowers, from the start,
was to involve mobile animals in their reproduction processes. That
is, pollen can be scattered even if the flower is not brightly colored
or oddly shaped in a way that attracts animals; however, by expending
the energy required to create such traits, angiosperms can enlist the
aid of animals and, thus, reproduce more efficiently.
Island genetics provides one proposed explanation for the sudden,
fully developed appearance of flowering plants.
Island genetics is
believed to be a common source of speciation in general, especially
when it comes to radical adaptations that seem to have required
inferior transitional forms. Flowering plants may have evolved in an
isolated setting like an island or island chain, where the plants
bearing them were able to develop a highly specialized relationship
with some specific animal (a wasp, for example). Such a relationship,
with a hypothetical wasp carrying pollen from one plant to another
much the way fig wasps do today, could result in the development of a
high degree of specialization in both the plant(s) and their partners.
Note that the wasp example is not incidental; bees, which, it is
postulated, evolved specifically due to mutualistic plant
relationships, are descended from wasps.
Animals are also involved in the distribution of seeds. Fruit, which
is formed by the enlargement of flower parts, is frequently a
seed-dispersal tool that attracts animals to eat or otherwise disturb
it, incidentally scattering the seeds it contains (see frugivory).
Although many such mutualistic relationships remain too fragile to
survive competition and to spread widely, flowering proved to be an
unusually effective means of reproduction, spreading (whatever its
origin) to become the dominant form of land plant life.
Flower ontogeny uses a combination of genes normally responsible for
forming new shoots. The most primitive flowers probably had a
variable number of flower parts, often separate from (but in contact
with) each other. The flowers tended to grow in a spiral pattern, to
be bisexual (in plants, this means both male and female parts on the
same flower), and to be dominated by the ovary (female part). As
flowers evolved, some variations developed parts fused together, with
a much more specific number and design, and with either specific sexes
per flower or plant or at least "ovary-inferior".
Flower evolution continues to the present day; modern flowers have
been so profoundly influenced by humans that some of them cannot be
pollinated in nature. Many modern domesticated flower species were
formerly simple weeds, which sprouted only when the ground was
disturbed. Some of them tended to grow with human crops, perhaps
already having symbiotic companion plant relationships with them, and
the prettiest did not get plucked because of their beauty, developing
a dependence upon and special adaptation to human affection.
A few paleontologists have also proposed that flowering plants, or
angiosperms, might have evolved due to interactions with dinosaurs.
One of the idea's strongest proponents is Robert T. Bakker. He
proposes that herbivorous dinosaurs, with their eating habits,
provided a selective pressure on plants, for which adaptations either
succeeded in deterring or coping with predation by herbivores.
In August 2017, scientists presented a detailed description and 3D
model image of what the first flower possibly looked like, and
presented the hypothesis that it may have lived about 140 million
A Bayesian analysis of 52 angiosperm taxa suggested that the crown
group of angisperms evolved between 178 million years ago and
198 million years ago.
Flowering plant diversity
A poster of twelve different species of flowers of the Asteraceae
Bud of a pink rose
The number of species of flowering plants is estimated to be in the
range of 250,000 to 400,000. This compares to around
12,000 species of moss or 11,000 species of pteridophytes,
showing that the flowering plants are much more diverse. The number of
families in APG (1998) was 462. In APG II (2003) it is not
settled; at maximum it is 457, but within this number there are 55
optional segregates, so that the minimum number of families in this
system is 402. In
APG III (2009) there are 415 families.
The diversity of flowering plants is not evenly distributed. Nearly
all species belong to the eudicot (75%), monocot (23%), and magnoliid
(2%) clades. The remaining 5 clades contain a little over 250 species
in total; i.e. less than 0.1% of flowering plant diversity, divided
among 9 families. The 43 most-diverse of 443 families of flowering
plants by species, in their APG circumscriptions, are
Asteraceae or Compositae (daisy family): 22,750 species;
Orchidaceae (orchid family): 21,950;
Fabaceae or Leguminosae (bean family): 19,400;
Rubiaceae (madder family): 13,150;
Poaceae or Gramineae (grass family): 10,035;
Lamiaceae or Labiatae (mint family): 7,175;
Euphorbiaceae (spurge family): 5,735;
Melastomataceae or Melastomaceae (melastome family): 5,005;
Myrtaceae (myrtle family): 4,625;
Apocynaceae (dogbane family): 4,555;
Cyperaceae (sedge family): 4,350;
Malvaceae (mallow family): 4,225;
Araceae (arum family): 4,025;
Ericaceae (heath family): 3,995;
Gesneriaceae (gesneriad family): 3,870;
Apiaceae or Umbelliferae (parsley family): 3,780;
Brassicaceae or Cruciferae (cabbage family): 3,710:
Piperaceae (pepper family): 3,600;
Bromeliaceae (bromeliad family): 3,540;
Acanthaceae (acanthus family): 3,500;
Rosaceae (rose family): 2,830;
Boraginaceae (borage family): 2,740;
Urticaceae (nettle family): 2,625;
Ranunculaceae (buttercup family): 2,525;
Lauraceae (laurel family): 2,500;
Solanaceae (nightshade family): 2,460;
Campanulaceae (bellflower family): 2,380;
Arecaceae (palm family): 2,361;
Annonaceae (custard apple family): 2,220;
Caryophyllaceae (pink family): 2,200;
Orobanchaceae (broomrape family): 2,060;
Amaranthaceae (amaranth family): 2,050;
Iridaceae (iris family): 2,025;
Aizoaceae or Ficoidaceae (ice plant family): 2,020;
Rutaceae (rue family): 1,815;
Phyllanthaceae (phyllanthus family): 1,745;
Scrophulariaceae (figwort family): 1,700;
Gentianaceae (gentian family): 1,650;
Convolvulaceae (bindweed family): 1,600;
Proteaceae (protea family): 1,600;
Sapindaceae (soapberry family): 1,580;
Cactaceae (cactus family): 1,500;
Aralia or ivy family): 1,450.
Of these, the Orchidaceae, Poaceae, Cyperaceae, Araceae, Bromeliaceae,
Iridaceae are monocot families; Piperaceae, Lauraceae,
Annonaceae are magnoliid dicots; the rest of the families are
Fertilization and embryogenesis
Angiosperm life cycle
Double fertilization refers to a process in which two sperm cells
fertilize cells in the ovule. This process begins when a pollen grain
adheres to the stigma of the pistil (female reproductive structure),
germinates, and grows a long pollen tube. While this pollen tube is
growing, a haploid generative cell travels down the tube behind the
tube nucleus. The generative cell divides by mitosis to produce two
haploid (n) sperm cells. As the pollen tube grows, it makes its way
from the stigma, down the style and into the ovary. Here the pollen
tube reaches the micropyle of the ovule and digests its way into one
of the synergids, releasing its contents (which include the sperm
cells). The synergid that the cells were released into degenerates and
one sperm makes its way to fertilize the egg cell, producing a diploid
(2n) zygote. The second sperm cell fuses with both central cell
nuclei, producing a triploid (3n) cell. As the zygote develops into an
embryo, the triploid cell develops into the endosperm, which serves as
the embryo's food supply. The ovary will now develop into a fruit and
the ovule will develop into a seed.
Fruit and seed
Seed and Fruit
The fruit of the
Aesculus or Horse Chestnut tree
As the development of embryo and endosperm proceeds within the embryo
sac, the sac wall enlarges and combines with the nucellus (which is
likewise enlarging) and the integument to form the seed coat. The
ovary wall develops to form the fruit or pericarp, whose form is
closely associated with type of seed dispersal system.
Frequently, the influence of fertilization is felt beyond the ovary,
and other parts of the flower take part in the formation of the fruit,
e.g., the floral receptacle in the apple, strawberry, and others.
The character of the seed coat bears a definite relation to that of
the fruit. They protect the embryo and aid in dissemination; they may
also directly promote germination. Among plants with indehiscent
fruits, in general, the fruit provides protection for the embryo and
secures dissemination. In this case, the seed coat is only slightly
developed. If the fruit is dehiscent and the seed is exposed, in
general, the seed-coat is well developed, and must discharge the
functions otherwise executed by the fruit.
Flowering plants generate gametes using a specialized cell division
Meiosis takes place in the ovule (a structure within
the ovary that is located within the pistil at the center of the
flower) (see diagram labeled "Angiosperm lifecycle"). A diploid cell
(megaspore mother cell) in the ovule undergoes meiosis (involving two
successive cell divisions) to produce four cells (megaspores) with
haploid nuclei. One of these four cells (megaspore) then undergoes
three successive mitotic divisions to produce an immature embryo sac
(megagametophyte) with eight haploid nuclei. Next, these nuclei are
segregated into separate cells by cytokinesis to producing 3 antipodal
cells, 2 synergid cells and an egg cell. Two polar nuclei are left in
the central cell of the embryo sac.
Pollen is also produced by meiosis in the male anther
(microsporangium). During meiosis, a diploid microspore mother cell
undergoes two successive meiotic divisions to produce 4 haploid cells
(microspores or male gametes). Each of these microspores, after
further mitoses, becomes a pollen grain (microgametophyte) containing
two haploid generative (sperm) cells and a tube nucleus. When a pollen
grain makes contact with the female stigma, the pollen grain forms a
pollen tube that grows down the style into the ovary. In the act of
fertilization, a male sperm nucleus fuses with the female egg nucleus
to form a diploid zygote that can then develop into an embryo within
the newly forming seed. Upon germination of the seed, a new plant can
grow and mature.
The adaptive function of meiosis is currently a matter of debate. A
key event during meiosis in a diploid cell is the pairing of
homologous chromosomes and homologous recombination (the exchange of
genetic information) between homologous chromosomes. This process
promotes the production of increased genetic diversity among progeny
and the recombinational repair of damages in the
DNA to be passed on
to progeny. To explain the adaptive function of meiosis in flowering
plants, some authors emphasize diversity and others emphasize DNA
Apomixis (reproduction via asexually formed seeds) is found naturally
in about 2.2% of angiosperm genera  One type of apomixis,
gametophytic apomixis found in a dandelion species  involves
formation of an unreduced embryo sac due to incomplete meiosis
(apomeiosis) and development of an embryo from the unreduced egg
inside the embryo sac, without fertilization (parthenogenesis).
Agriculture is almost entirely dependent on angiosperms, which provide
virtually all plant-based food, and also provide a significant amount
of livestock feed. Of all the families of plants, the Poaceae, or
grass family (providing grains), is by far the most important,
providing the bulk of all feedstocks (rice, maize, wheat, barley, rye,
oats, pearl millet, sugar cane, sorghum). The Fabaceae, or legume
family, comes in second place. Also of high importance are the
Solanaceae, or nightshade family (potatoes, tomatoes, and peppers,
among others); the Cucurbitaceae, or gourd family (including pumpkins
and melons); the Brassicaceae, or mustard plant family (including
rapeseed and the innumerable varieties of the cabbage species Brassica
oleracea); and the Apiaceae, or parsley family. Many of our fruits
come from the Rutaceae, or rue family (including oranges, lemons,
grapefruits, etc.), and the Rosaceae, or rose family (including
apples, pears, cherries, apricots, plums, etc.).
In some parts of the world, certain single species assume paramount
importance because of their variety of uses, for example the coconut
(Cocos nucifera) on Pacific atolls, and the olive (Olea europaea) in
the Mediterranean region.
Flowering plants also provide economic resources in the form of wood,
paper, fiber (cotton, flax, and hemp, among others), medicines
(digitalis, camphor), decorative and landscaping plants, and many
other uses. The main area in which they are surpassed by other plants
— namely, coniferous trees (Pinales), which are non-flowering
(gymnosperms) — is timber and paper production.
List of garden plants
List of plant orders
List of plants by common name
List of systems of plant taxonomy
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Wikimedia Commons has media related to Magnoliophyta.
Wikispecies has information related to Magnoliophyta
The Wikibook Dichotomous Key has a page on the topic of: Magnoliophyta
Plantae sensu lato
& land plants)
& land plants)
Angiosperms or flowering plants
† = extinct. See also the list of plant orders.
History of botany
Hypanthium (Floral cup)
Plant growth and habit
Alternation of generations
History of plant systematics
International Code of Nomenclature for algae, fungi, and plants
International Code of Nomenclature for algae, fungi, and plants (ICN)
- for Cultivated Plants (ICNCP)
International Association for
Plant Taxonomy (IAPT)
Plant taxonomy systems
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by author abbreviation