Flowering plant

The flowering plants, also known as angiosperms, Angiospermae[5][6] or Magnoliophyta,[7]

Flowering plants
are the most diverse group of land plants, with 64 orders, 416 families, approximately 13,000
known genera and 300,000 known species.[8] Like gymnosperms, angiosperms are seed- Temporal range: Early Cretaceous –
producing plants. However, they are distinguished from gymnosperms by characteristics present, 130–0 Ma
including flowers, endosperm within the seeds, and the production of fruits that contain the PreЄ Є O S D C P T J K PgN
seeds. Etymologically, angiosperm means a plant that produces seeds within an enclosure; in Possible Early Jurassic record
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 ~140 mya. They
diversified extensively during the Early Cretaceous, became widespread by 120 mya, and
replaced conifers as the dominant trees from 100 to 60 mya.

Contents
Description
Angiosperm derived characteristics
Vascular anatomy
Reproductive anatomy
Taxonomy
Diversity of angiosperms
History of classification
Modern classification Scientific classification
Evolution
Kingdom: Plantae
Diversity
Clade: Spermatophytes
Reproduction
Fertilization and embryogenesis Clade: Angiosperms
Fruit and seed
Meiosis Groups (APG IV)[1]
Apomixis
†Nanjinganthus?
Uses
See also Basal angiosperms

Notes
Amborellales
References
Nymphaeales
Bibliography
Articles, books and chapters Austrobaileyales
Websites
Core angiosperms
External links
Clades

Description magnoliids
monocots

Angiosperm derived characteristics eudicots

Angiosperms differ from other seed plants in several ways, described in the table below. These Orders
distinguishing characteristics taken together have made the angiosperms the most diverse and
numerous land plants and the most commercially important group to humans.[a]
Chloranthales
Ceratophyllales
 Distinctive features of angiosperms Synonyms
Feature Description
Anthophyta Cronquist[2]
Flowers, the reproductive organs of flowering plants, are the most
remarkable feature distinguishing them from the other seed plants. Angiospermae Lindl.
Flowers provided angiosperms with the means to have a more
Flowering species-specific breeding system, and hence a way to evolve more Magnoliophyta Cronquist, Takht.
organs readily into different species without the risk of crossing back with
related species. Faster speciation enabled the Angiosperms to & W.Zimm.[3]
adapt to a wider range of ecological niches. This has allowed
flowering plants to largely dominate terrestrial ecosystems. Magnolicae Takht.[4]
Stamens are much lighter than the corresponding organs of
gymnosperms and have contributed to the diversification of
Stamens with angiosperms through time with adaptations to specialized pollination
two pairs of syndromes, such as particular pollinators. Stamens have also
pollen sacs become modified through time to prevent self-fertilization, which has
permitted further diversification, allowing angiosperms eventually to
fill more niches.
The male gametophyte in angiosperms is significantly reduced in
size compared to those of gymnosperm seed plants.[9] The smaller
size of the pollen reduces the amount of time between pollination —
the pollen grain reaching the female plant — and fertilization. In
Reduced
gymnosperms, fertilization can occur up to a year after pollination,
male parts,
whereas in angiosperms, fertilization begins very soon after
three cells
pollination.[10] The shorter amount of time between pollination and
fertilization allows angiosperms to produce seeds earlier after
pollination than gymnosperms, providing angiosperms a distinct
evolutionary advantage.
Closed
carpel The closed carpel of angiosperms also allows adaptations to
enclosing the specialized pollination syndromes and controls. This helps to
ovules prevent self-fertilization, thereby maintaining increased diversity.
(carpel or Once the ovary is fertilized, the carpel and some surrounding
carpels and tissues develop into a fruit. This fruit often serves as an attractant to
accessory seed-dispersing animals. The resulting cooperative relationship
parts may presents another advantage to angiosperms in the process of
become the dispersal.
fruit)
Reduced
The reduced female gametophyte, like the reduced male
female
gametophyte, may be an adaptation allowing for more rapid seed
gametophyte,
set, eventually leading to such flowering plant adaptations as annual
seven cells
herbaceous life-cycles, allowing the flowering plants to fill even more
with eight
niches.
nuclei
In general, endosperm formation begins after fertilization and before
the first division of the zygote. Endosperm is a highly nutritive tissue
Endosperm
that can provide food for the developing embryo, the cotyledons,
and sometimes the seedling when it first appears.

Vascular anatomy
Angiosperm stems are made up of seven layers as shown on the right. 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 exceptional cases.
Reproductive anatomy
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- Cross-section of a stem of the
bearing or vegetative portion, and forms a more or less elaborate branch- angiosperm flax:
system called an inflorescence. 1. pith, 2. protoxylem, 3. xylem, 4.
phloem, 5. sclerenchyma (bast fibre),
There are two kinds of reproductive cells produced by flowers. 6. cortex, 7. epidermis
Microspores, which will divide to become pollen grains, are the "male"
A collection of
flowers forming cells and are borne in the stamens (or microsporophylls). The "female"
an cells called megaspores, which will divide to become the egg cell (megagametogenesis), are contained in the ovule and
inflorescence. 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 among humans.

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.

Taxonomy

History of classification
The botanical term "Angiosperm", from the Ancient Greek ἀγγεῖον, angeíon (bottle, vessel) and σπέρμα,
sperma (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
maintained by 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,[11] 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 plants.

In 1851, Hofmeister discovered the changes occurring in the embryo-sac of flowering plants, and determined From 1736, an illustration of
the correct relationships of these to the Cryptogamia. This fixed the position of Gymnosperms as a class Linnaean classification
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 (Angiosperms), with 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 subdivision. The 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. The Takhtajan system and Cronquist system treat this group at the rank of division, leading to the
name Magnoliophyta (from the family name Magnoliaceae). The 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[12] and 2009[13] 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 Subclass Magnoliidae.[14]
An auxanometer, a device
The internal classification of this group has undergone considerable revision. The Cronquist system,
for measuring increase or
proposed by Arthur Cronquist in 1968 and published in its full form in 1981, is still widely used but is no rate of growth in plants
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,[12] the APG III system in 2009,[13][15] 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 Liliopsida (at the
rank of class, formed from the family name Liliaceae). Other descriptive names allowed by Article 16 of the ICBN include Dicotyledones or
Dicotyledoneae, and Monocotyledones or 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 Ceratophyllaceae and Chloranthaceae.

Modern classification
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,[16] water lilies and Hydatellaceae;
Austrobaileyales, about 100 species[16] of woody plants from various parts of the world
Core angiosperms (Mesangiospermae)[14]

Chloranthales, 77 known species[17] of aromatic plants with toothed leaves;

Magnoliids, about 9,000 species,[16] 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,[16] characterized by trimerous flowers, a single
cotyledon, pollen with one pore, and usually parallel-veined leaves—for example grasses,
orchids, and palms;
Monocot (left) and dicot
Ceratophyllum, about 6 species[16] of aquatic plants, perhaps most familiar as aquarium
plants; seedlings

Eudicots, about 175,000 species,[16] 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.[18] 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.[19] 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.[20]
Ceratophyllum seems to group with the eudicots rather than with the monocots. The 2016 Angiosperm Phylogeny Group revision (APG IV) retained
the overall higher order relationship described in APG III.[13]

Amborella

Nymphaeales

Austrobaileyales

magnoliids
angiosperms
Chloranthales

monocots

Ceratophyllum

eudicots

1. Phylogeny of the flowering plants, as of APG III (2009).[13]

Amborella

Nymphaeales

Austrobaileyales

monocots
angiosperms
Chloranthales

magnoliids

Ceratophyllum

eudicots

2. Example of alternative phylogeny (2010)[20]

angiosperms
Amborellales

Nymphaeales basal angiosperms

Austrobaileyales

magnoliids

Chloranthales
core angiosperms
 monocots

Ceratophyllales

eudicots

3. APG IV (2016)[1]

Detailed Cladogram of the Angiosperm Phylogeny Group (APG) IV classification.[1]

Amborellales Melikyan, Bobrov & Zaytzeva 1999

Nymphaeales Salisbury ex von Berchtold & Presl 1820

Austrobaileyales Takhtajan ex Reveal 1992

Mesangiosperms
Chloranthales Mart. 1835

Canellales Cronquist 1957

Piperales von Berchtold & Presl 1820

Magnoliids
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

Petrosaviales Takhtajan 1997

Dioscoreales Brown 1835

Pandanales Brown ex von Berchtold & Presl 1820

Monocots Liliales Perleb 1826

Asparagales Link 1829

Arecales Bromhead 1840

Poales Small 1903

Commelinids
Zingiberales Grisebach 1854

Commelinales de Mirbel ex von Berchtold & Presl 1820

Ceratophyllales Link 1829

Eudicots
Ranunculales de Jussieu ex von Berchtold & Presl 1820

Proteales de Jussieu ex von Berchtold & Presl 1820

Trochodendrales Takhtajan ex Cronquist 1981

Buxales Takhtajan ex Reveal 1996

Core eudicots
Gunnerales 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

Fabids
(eurosids I) Fabales Bromhead 1838

Rosales von Berchtold & Presl

1820

Cucurbitales de Jussieu ex
von Berchtold & Presl 1820

Superrosids Fagales Engler 1892

Rosids
Geraniales de Jussieu ex von Berchtold
& Presl 1820

Myrtales de Jussieu ex von Berchtold &

Presl 1820

Crossosomatales Takhtajan ex Reveal

1993

Picramniales Doweld 2001

Malvids
(eurosids II)
Sapindales de Jussieu ex von
Berchtold & Presl 1820

Huerteales Doweld 2001

Malvales de Jussieu ex von

Berchtold & Presl 1820

Brassicales Bromhead 1838

Superasterids
Berberidopsidales Doweld 2001

Santalales Brown ex von Berchtold & Presl 1820

Caryophyllales

Asterids
 Cornales Link 1829

Ericales von Berchtold & Presl 1820

Icacinales Van Tieghem 1900

Metteniusales Takhtajan 1997

Garryales Mart. 1835

Gentianales de Jussieu ex
von Berchtold & Presl 1820

Lamiids Solanales de Jussieu ex von

(euasterids I)
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

Campanulids
(euasterids II) Apiales Nakai 1930

Paracryphiales Takhtajan ex
Reveal 1992

Dipsacales de Jussieu ex von

Berchtold & Presl 1820

Evolution
Fossilized spores suggest that land plants (embryophytes) have existed for at least 475 million years.[21] 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 gametophytes) of Ginkgo and cycads produce a pair of flagellated, mobile sperm cells that "swim" down
the developing pollen tube to the female and her eggs.

The apparently sudden appearance of nearly modern flowers in the fossil record initially posed such a problem for the theory of evolution that
Charles Darwin called it an "abominable mystery".[22] 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 Triassic Sanmiguelia, 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 Triassic (247.2–242.0 Ma) suggests an older date for their origin.[23] 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.[24][25] The fossil plant species
Nanjinganthus dendrostyla from Early Jurassic China seems to share many exclusively angiosperm
features, such as a thickened receptacle with ovules, and thus might represent a crown-group or a
stem-group angiosperm.[26][27] However, the interpretation of the structures in this fossils are
highly contested.[28][29]

The evolution of seed plants and later angiosperms appears to be the result of two distinct rounds of
whole genome duplication events.[30] These occurred at 319 million years ago and
192 million years ago. Another possible whole genome duplication event at 160 million years ago Flowers of Malus sylvestris (crab
perhaps created the ancestral line that led to all modern flowering plants.[31] That event was studied apple)
by sequencing the genome of an ancient flowering plant, Amborella trichopoda,[32] and directly
addresses Darwin's "abominable mystery."

The earliest known macrofossil confidently identified as an angiosperm, Archaefructus

liaoningensis, is dated to about 125 million years BP (the Cretaceous period),[33] whereas pollen
considered to be of angiosperm origin takes the fossil record back to about 130 million years BP.[34]
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.[35]
In 2018, scientists reported that the earliest flowers began about 180 million years ago, 50 million
years earlier than thought earlier.[36] Nonetheless, 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.[37][38] 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 plants themselves.

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, Flowers and leaves of Oxalis pes-
suggesting it was carried by insects.[39] caprae (Bermuda buttercup)

Recent DNA analysis based on molecular systematics[40][41] 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[42] 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.[43]

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.[44] 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.[45]
The radiation of herbaceous angiosperms occurred much later.[46] Yet, many fossil plants recognizable as belonging to modern families (including
beech, oak, maple, and magnolia) had already appeared by the late Cretaceous.

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.[47]

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.[48] Two bees on a flower head of
creeping thistle, Cirsium arvense
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.[49] 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.[50]

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.[51]

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 years ago.[52][53]

A Bayesian analysis of 52 angiosperm taxa suggested that the crown group of angiosperms evolved between 178 million years ago and
198 million years ago.[54]

Diversity
The number of species of flowering plants is estimated to be in the range of 250,000 to 400,000.[55][56][57] This compares to around 12,000 species
of moss[58] or 11,000 species of pteridophytes,[59] showing that the flowering plants are much more diverse. The number of families in APG (1998)
was 462. In APG II[12] (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.[13][60]

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,[61] in their APG circumscriptions, are
 1. Asteraceae or Compositae (daisy family): 22,750 species;
2. Orchidaceae (orchid family): 21,950;
3. Fabaceae or Leguminosae (bean family): 19,400;
4. Rubiaceae (madder family): 13,150;[62]
5. Poaceae or Gramineae (grass family): 10,035;
6. Lamiaceae or Labiatae (mint family): 7,175;
7. Euphorbiaceae (spurge family): 5,735;
8. Melastomataceae or Melastomaceae (melastome family): 5,005;
9. Myrtaceae (myrtle family): 4,625;
10. Apocynaceae (dogbane family): 4,555;
11. Cyperaceae (sedge family): 4,350;
12. Malvaceae (mallow family): 4,225;
13. Araceae (arum family): 4,025;
14. Ericaceae (heath family): 3,995;
15. Gesneriaceae (gesneriad family): 3,870;
16. Apiaceae or Umbelliferae (parsley family): 3,780;
17. Brassicaceae or Cruciferae (cabbage family): 3,710: A poster of twelve different species of
18. Piperaceae (pepper family): 3,600; flowers of the family Asteraceae
19. Bromeliaceae (bromeliad family): 3,540;
20. Acanthaceae (acanthus family): 3,500;
21. Rosaceae (rose family): 2,830;
22. Boraginaceae (borage family): 2,740;
23. Urticaceae (nettle family): 2,625;
24. Ranunculaceae (buttercup family): 2,525;
25. Lauraceae (laurel family): 2,500;
26. Solanaceae (nightshade family): 2,460;
27. Campanulaceae (bellflower family): 2,380;
28. Arecaceae (palm family): 2,361;
29. Annonaceae (custard apple family): 2,220;
30. Caryophyllaceae (pink family): 2,200;
31. Orobanchaceae (broomrape family): 2,060;
32. Amaranthaceae (amaranth family): 2,050;
33. Iridaceae (iris family): 2,025;
34. Aizoaceae or Ficoidaceae (ice plant family): 2,020;
35. Rutaceae (rue family): 1,815; Lupinus pilosus
36. Phyllanthaceae (phyllanthus family): 1,745;
37. Scrophulariaceae (figwort family): 1,700;
38. Gentianaceae (gentian family): 1,650;
39. Convolvulaceae (bindweed family): 1,600;
40. Proteaceae (protea family): 1,600;
41. Sapindaceae (soapberry family): 1,580;
42. Cactaceae (cactus family): 1,500;
43. Araliaceae (Aralia or ivy family): 1,450.
Of these, the Orchidaceae, Poaceae, Cyperaceae, Araceae, Bromeliaceae, Arecaceae, and Iridaceae
are monocot families; Piperaceae, Lauraceae, and Annonaceae are magnoliid dicots; the rest of the
families are eudicots.

Reproduction

Fertilization and embryogenesis

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 Bud of a pink rose

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

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 Angiosperm life cycle
with type of seed dispersal system.[63]

Frequently, the influence of fertilization is felt beyond the ovary, and other parts of the flower take
The fruit of the Aesculus or horse
part in the formation of the fruit, e.g., the floral receptacle in the apple, strawberry, and others.
chestnut tree
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.

Meiosis
Flowering plants generate gametes using a specialized cell division called meiosis. 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.[64] 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[65] and others emphasize DNA repair.[66]

Apomixis
Apomixis (reproduction via asexually formed seeds) is found naturally in about 2.2% of angiosperm genera.[67] One type of apomixis, gametophytic
apomixis found in a dandelion species[68] 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).

Uses
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.[69]

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.[70]

See also
List of garden plants
List of plant orders
List of plants by common name
List of systems of plant taxonomy

Notes
a. The major exception to the dominance of terrestrial ecosystems by flowering plants is the coniferous forest.

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Websites
Cole, Theodor C.H.; Hilger, Harmut H.; Stevens, Peter F. (2017). "Angiosperm Phylogeny Poster – Flowering Plant Systematics"
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External links
Media related to Magnoliophyta at Wikimedia Commons
Data related to Magnoliophyta at Wikispecies
Magnoliophyta at Wikibooks

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