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Flowering plant
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Main page The flowering plants, also known as angiosperms, Angiospermae,[6][7] or Magnoliophyta,[8] are the most diverse group of land
Flowering plant
Contents plants, with 64 orders, 416 families, approximately 13,000 known genera and 300,000 known species.[9] Like gymnosperms,
Featured content Temporal range: Early Cretaceous –
angiosperms are seed-producing plants. They are distinguished from gymnosperms by characteristics including flowers, endosperm
Current events present, 130–0 Ma[1]
within the seeds, and the production of fruits that contain the seeds. Etymologically, angiosperm means a plant that produces seeds
Random article PreЄ Є O S D C P T J K PgN
within an enclosure; in other words, a fruiting plant. The term comes from the Greek words angeion ("case" or "casing") and sperma
Donate to Wikipedia Possible Early Permian record
Wikipedia store ("seed").

The ancestors of flowering plants diverged from gymnosperms in the Triassic Period, 245 to 202 million years ago (mya), and the first
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flowering plants are known from ~140 mya. They diversified extensively during the Early Cretaceous, became widespread by 120 mya,
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and replaced conifers as the dominant trees from 100 to 60 mya.
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Contents [hide]
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Contact page 1 Description
1.1 Angiosperm derived characteristics
Tools 1.2 Vascular anatomy
What links here 1.3 Reproductive anatomy
Related changes
2 Taxonomy
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2.1 History of classification
Special pages
Permanent link
2.2 Modern classification
Page information 2.3 Evolutionary history
Wikidata item 2.3.1 Paleozoic Diversity of angiosperms
Cite this page 2.3.2 Triassic and Jurassic
Scientific classification
2.3.3 Cretaceous
In other projects Kingdom: Plantae
3 Diversity
Wikimedia Commons Clade: Tracheophytes
4 Reproduction
Wikispecies
4.1 Fertilization and embryogenesis Clade: Spermatophytes
Print/export 4.2 Fruit and seed Clade: Angiosperms
Download as PDF 4.3 Meiosis Groups (APG IV)[2]
Printable version 4.4 Apomixis
5 Uses †Nanjinganthus?
Languages
6 See also Basal angiosperms
Deutsch
Español 7 Notes Amborellales
한국어 8 References Nymphaeales
ह ी 9 Bibliography Austrobaileyales
Italiano 9.1 Articles, books and chapters
⽇本語 9.2 Websites
Core angiosperms
Português Clades
10 External links
‫اردو‬
中⽂ magnoliids
monocots
105 more Description [ edit ] eudicots
Edit links
Orders
Angiosperm derived characteristics [ edit ]
Chloranthales
Angiosperms differ from other seed plants in several ways, described in the table below. These distinguishing characteristics taken Ceratophyllales
together have made the angiosperms the most diverse and numerous land plants and the most commercially important group to
Synonyms
humans.[a]
Anthophyta Cronquist[3]
Distinctive features of angiosperms
Angiospermae Lindl.
Feature Description Magnoliophyta Cronquist, Takht. &
Flowers, the reproductive organs of flowering plants, are the most remarkable feature distinguishing them from the W.Zimm.[4]
other seed plants. Flowers provided angiosperms with the means to have a more species-specific breeding system, Magnolicae Takht.[5]
Flowering
and hence a way to evolve more readily into different species without the risk of crossing back with related species.
organs
Faster speciation enabled the Angiosperms to adapt to a wider range of ecological niches. This has allowed
flowering plants to largely dominate terrestrial ecosystems.[citation needed]
Stamens are much lighter than the corresponding organs of gymnosperms and have contributed to the
Stamens with
diversification of angiosperms through time with adaptations to specialized pollination syndromes, such as particular
two pairs of
pollinators. Stamens have also become modified through time to prevent self-fertilization, which has permitted
pollen sacs
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.[10] The smaller size of the pollen reduces the amount of time between pollination — the pollen grain reaching
Reduced male the female plant — and fertilization. In gymnosperms, fertilization can occur up to a year after pollination, whereas in
parts, three cells angiosperms, fertilization begins very soon after pollination.[11] 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
enclosing the
The closed carpel of angiosperms also allows adaptations to specialized pollination syndromes and controls. This
ovules (carpel or
helps to prevent self-fertilization, thereby maintaining increased diversity. Once the ovary is fertilized, the carpel and
carpels and
some surrounding tissues develop into a fruit. This fruit often serves as an attractant to seed-dispersing animals.
accessory parts
The resulting cooperative relationship presents another advantage to angiosperms in the process of dispersal.
may become the
fruit)
Reduced female
The reduced female gametophyte, like the reduced male gametophyte, may be an adaptation allowing for more
gametophyte,
rapid seed set, eventually leading to such flowering plant adaptations as annual herbaceous life-cycles, allowing the
seven cells with
flowering plants to fill even more niches.
eight nuclei
In general, endosperm formation begins after fertilization and before the first division of the zygote. Endosperm is a
Endosperm highly nutritive tissue that can provide food for the developing embryo, the cotyledons, and sometimes the seedling
when it first appears.

Vascular anatomy [ edit ]

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
Cross-section of a stem of the
each season of growth, called annual rings. angiosperm flax:
1. pith, 2. protoxylem, 3. xylem, 4.
Among the monocotyledons, the bundles are more numerous in the young stem and are scattered through the ground tissue. They
phloem, 5. sclerenchyma (bast fibre),
contain no cambium and once formed the stem increases in diameter only in exceptional cases. 6. cortex, 7. epidermis

Reproductive anatomy [ edit ]

Main articles: Flower and Plant reproductive morphology

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 inflorescence.

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,
A collection of 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,
flowers forming
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
an inflorescence.
humans.[citation needed]

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.[citation needed]

Taxonomy [ edit ]

History of classification [ edit ]

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,[12] and
applied to them the name Gymnosperms.[citation needed] 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 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 From 1736, an illustration
of Linnaean classification
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,[13] 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[14] and 2009[15] revisions,
treat the flowering plants as a clade called angiosperms without a formal botanical name. A formal classification was published alongside the
2009 revision in which the flowering plants form the Subclass Magnoliidae.[16]

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
An auxanometer, a device
published an influential reclassification of the angiosperms in 1998. Updates incorporating more recent research were published as the APG II
for measuring increase or rate
system in 2003,[14] the APG III system in 2009,[15][17] and the APG IV system in 2016. of growth in plants

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.[citation needed]

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.[citation needed]

Modern classification [ edit ]

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,[18] water lilies and Hydatellaceae;
Austrobaileyales, about 100 species[18] of woody plants from various parts of the world
Core angiosperms (Mesangiospermae)[16]
Chloranthales, 77 known species[19] of aromatic plants with toothed leaves;
Magnoliids, about 9,000 species,[18] 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,[18] 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[18] of aquatic plants, perhaps most familiar as aquarium plants; Monocot (left) and dicot
seedlings
Eudicots, about 175,000 species,[18] 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.[20] 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.[21] 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.[22] 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.[15]

1. Phylogeny of the flowering plants, as of APG III 2. Example of alternative phylogeny (2010)[22]
(2009).[15]
  Amborella
     
  Amborella
      Nymphaeales
 
 
  Nymphaeales
  Austrobaileyales
 
 
  Austrobaileyales
  monocots
 
angiosperms  
  magnoliids
    Chloranthales
angiosperms    
   
        Chloranthales   magnoliids
     
         
        monocots
      Ceratophyllum
     
    Ceratophyllum      
    eudicots
     
 
    eudicots
 
3. APG IV (2016)[2]

 
  Amborellales
  Nymphaeales
  basal angiosperms
  Austrobaileyales    (ANA group)
 
 
angiosperms   magnoliids
      Chloranthales
     
        monocots
     
  Ceratophyllales  core angiosperms
 
     
    eudicots
 

Detailed Cladogram of the Angiosperm Phylogeny Group (APG) IV classification.[2] [show]

Evolutionary history [ edit ]

Further information: Evolutionary history of plants § Flowers

Paleozoic [ edit ]

Fossilized spores suggest that land plants (embryophytes) have existed for at least 475 million years.[23] 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 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".[24] 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.

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.

Oleanane, a secondary metabolite produced by many flowering plants, has been found in Permian deposits of that age together with fossils of gigantopterids.[25][26]
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.[citation needed]

Triassic and Jurassic [ edit ]

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.[27] 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.[28][29]

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.[30][31] However, the
Fluffy flowers of Tetradenia riparia
interpretation of the structures in this fossils are highly contested.[32][33]
(misty plume bush)
The evolution of seed plants and later angiosperms appears to be the result of two distinct rounds of whole genome duplication
events.[34] 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.[35] That event was studied by sequencing
the genome of an ancient flowering plant, Amborella trichopoda,[36] and directly addresses Darwin's "abominable mystery".

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.[37]

Cretaceous [ edit ]

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.[38]
Flowers of Malus sylvestris (crab
The earliest known macrofossil confidently identified as an angiosperm, Archaefructus liaoningensis, is dated to about 125 million years apple)
BP (the Cretaceous period),[39] whereas pollen considered to be of angiosperm origin takes the fossil record back to about 130 million
years BP,[40] with Montsechia representing the earliest flower at that time.[41] In 2018, scientists reported that the earliest flowers began
about 180 million years ago, 50 million years earlier than thought earlier.[42] Nonetheless, circumstantial chemical evidence has been
found for the existence of angiosperms as early as 250 million years ago (see the above Paleozoic section)

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.[43] 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.[44][45]
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.[46] Flowers and leaves of Senecio
angulatus (Cape Ivy)
Recent DNA analysis based on molecular systematics[47][48] 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[49] 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.[50]

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.[51] 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 Two bees on the composite flower
relationship, with a hypothetical wasp carrying pollen from one plant to another much the way fig wasps do today, could result in the head of creeping thistle, Cirsium
arvense
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.[52]

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.[citation needed]

Flower ontogeny uses a combination of genes normally responsible for forming new shoots.[53] 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.[54]

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.[55]

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.[56] The radiation of herbaceous angiosperms
occurred much later.[57] Yet, many fossil plants recognizable as belonging to modern families (including beech, oak, maple, and magnolia) had already appeared by the late
Cretaceous. Flowering plants appeared in Australia about 126 million years ago. This also pushed the age of ancient Australian vertebrates, in what was then a south polar
continent, to 126-110 million years old.[41]

Diversity [ edit ]

The number of species of flowering plants is estimated to be in the range of 250,000 to 400,000.[58][59][60] This compares to around
12,000 species of moss[61] or 11,000 species of pteridophytes,[62] showing that the flowering plants are much more diverse. The number
of families in APG (1998) was 462. In APG II[14] (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.[15][63]

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,[64] 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;[65]
5. Poaceae or Gramineae (grass family): 10,035;
6. Lamiaceae or Labiatae (mint family): 7,175;
A poster of twelve different species
7. Euphorbiaceae (spurge family): 5,735;
of flowers of the family Asteraceae
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:
18. Piperaceae (pepper family): 3,600;
19. Bromeliaceae (bromeliad family): 3,540;
20. Acanthaceae (acanthus family): 3,500;
21. Rosaceae (rose family): 2,830; Lupinus pilosus
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;
36. Phyllanthaceae (phyllanthus family): 1,745;
37. Scrophulariaceae (figwort family): 1,700;
Bud of a pink rose
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 [ edit ]

Fertilization and embryogenesis [ edit ]

Main articles: Fertilization and Plant 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 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 [ edit ]

Main articles: Seed and Fruit

As the development of embryo and endosperm proceeds within the embryo sac, the sac wall Angiosperm life cycle
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.[66]

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.[citation needed]

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
The fruit of the Aesculus or horse
chestnut tree 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.[citation needed]

Meiosis [ edit ]

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.[67] It is thought that the basal chromosome number in angiosperms is n = 7.[68] 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.
[citation needed]

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.[citation needed]

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[69] and others emphasize DNA repair.[70]

Apomixis [ edit ]

Apomixis (reproduction via asexually formed seeds) is found naturally in about 2.2% of angiosperm genera.[71] One type of apomixis, gametophytic apomixis found in a
dandelion species[72] 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).[citation needed]

Uses [ edit ]

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.).[citation needed]

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.[73]

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.[74]

See also [ edit ]

List of garden plants

List of plant orders
List of plants by common name
List of systems of plant taxonomy

Notes [ edit ]

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

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Websites [ edit ]
Cole, Theodor C.H.; Hilger, Harmut H.; Stevens, Peter F. (2017). "Angiosperm Phylogeny Watson, L.; Dallwitz, M.J. (1992). "The Families of Flowering Plants: Descriptions,
Poster – Flowering Plant Systematics" (PDF). Illustrations, Identification, and Information Retrieval" . 14 December 2000. Archived from
the original on 2014-08-02.
"Flowering plant" at the Encyclopedia of Life

External links [ edit ]

Media related to Magnoliophyta at Wikimedia Commons

Data related to Magnoliophyta at Wikispecies
Magnoliophyta at Wikibooks

V·T·E Classification of Archaeplastida / Plantae sensu lato [show]

V·T·E Botany [show]

V·T·E Extant life phyla/divisions by domain [show]

Taxon identifiers Wikidata: Q25314 · Wikispecies: Magnoliopsida · EPPO: 1ANGC · ITIS: 18061 · PPE: angiosperma-parasites

Authority control GND: 4144254-4 · LCCN: sh85005034 · NDL: 00563270

Categories: Angiosperms Plant sexuality Plants Pollination

This page was last edited on 14 April 2020, at 01:26 (UTC).

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