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Scientific classification

Kingdom: Plantae

Subkingdom: Embryophyta

(unranked): Spermatophyta

(unranked): Angiosperms

Groups (APG IV)[1]

Basal angiosperms

Amborellales Nymphaeales Austrobaileyales

Core angiosperms

magnoliids Chloranthales monocots Ceratophyllales eudicots

Synonyms

Anthophyta Cronquist[2] Angiospermae Lindl. Magnoliophyta Cronquist, Takht.
Takht.
& W.Zimm.[3] Magnolicae Takht.[4]

The flowering plants, also known as angiosperms, Angiospermae[5][6] or Magnoliophyta,[7] are the most diverse group of land plants, with 416 families, approximately 13,164 known genera and c. 295,383 known species.[8] 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
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.

Contents

1 Description

1.1 Angiosperm derived characteristics 1.2 Vascular anatomy 1.3 Reproductive anatomy

2 Taxonomy

2.1 History of classification 2.2 Modern classification 2.3 Evolution

3 Flowering plant
Flowering plant
diversity 4 Reproduction

4.1 Fertilization
Fertilization
and embryogenesis 4.2 Fruit
Fruit
and seed 4.3 Meiosis 4.4 Apomixis

5 Uses 6 See also 7 Notes 8 References 9 Bibliography

9.1 Articles, books and chapters 9.2 Websites

10 External links

Description[edit] Angiosperm derived characteristics[edit] Angiosperms
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

Feature Description

Flowering organs 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
Angiosperms
to adapt to a wider range of ecological niches. This has allowed flowering plants to largely dominate terrestrial ecosystems.[citation needed]

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.[9] 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.[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 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.

Endosperm In general, endosperm formation begins after fertilization and before the first division of the zygote. Endosperm
Endosperm
is a highly nutritive tissue that can provide food for the developing embryo, the cotyledons, and sometimes the seedling when it first appears.

Vascular anatomy[edit]

Cross-section of a stem of the angiosperm flax: 1. Pith, 2. Protoxylem, 3. Xylem
Xylem
I, 4. Phloem
Phloem
I, 5. Sclerenchyma
Sclerenchyma
(bast 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 exceptional cases. Reproductive anatomy[edit] Main articles: Flower
Flower
and Plant
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 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
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[edit] History of classification[edit]

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 maintained by Carl Linnaeus
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.[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.

An auxanometer, a device for measuring increase or rate of growth in 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 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
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
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] 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
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
Dicotyledoneae
or Magnoliopsida Monocotyledoneae
Monocotyledoneae
or Liliopsida

which in the Cronquist system
Cronquist system
are called Magnoliopsida
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
ICBN
include Dicotyledones
Dicotyledones
or Dicotyledoneae, and Monocotyledones
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
Ceratophyllaceae
and Chloranthaceae. Modern classification[edit]

Monocot
Monocot
(left) and dicot seedlings

There are eight groups of living angiosperms:

Basal 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
Core angiosperms
(Mesangiospermae)[14]

Chloranthales, several dozen species 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; Ceratophyllum, about 6 species[16] of aquatic plants, perhaps most familiar as aquarium plants; 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.[17] 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.[18] 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.[19] Ceratophyllum
Ceratophyllum
seems to group with the eudicots rather than with the monocots. The 2016 Angiosperm Phylogeny
Phylogeny
Group revision (APG IV) retained the overall higher order relationship described in APG III.[13]

angiosperms

Amborella

Nymphaeales

Austrobaileyales

magnoliids

Chloranthales

monocots

Ceratophyllum

eudicots

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

angiosperms

Amborella

Nymphaeales

Austrobaileyales

monocots

Chloranthales

magnoliids

Ceratophyllum

eudicots

2. Example of alternative phylogeny (2010)[19]

angiosperms

 Amborellales 

 Nymphaeales 

 Austrobaileyales 

 magnoliids 

 Chloranthales 

 monocots 

 Ceratophyllales  

 eudicots 

basal angiosperms core angiosperms

3. APG IV (2016)[1]

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

Amborellales
Amborellales
Melikyan, Bobrov & Zaytzeva 1999

Nymphaeales
Nymphaeales
Salisbury ex von Berchtold & Presl 1820

Austrobaileyales
Austrobaileyales
Takhtajan
Takhtajan
ex Reveal 1992

Mesangiosperms

Chloranthales
Chloranthales
Mart. 1835

Magnoliids

Canellales
Canellales
Cronquist 1957

Piperales
Piperales
von Berchtold & Presl 1820

Magnoliales
Magnoliales
de Jussieu ex von Berchtold & Presl 1820

Laurales
Laurales
de Jussieu ex von Berchtold & Presl 1820

Monocots

Acorales
Acorales
Link 1835

Alismatales
Alismatales
Brown ex von Berchtold & Presl 1820

Petrosaviales
Petrosaviales
Takhtajan
Takhtajan
1997

Dioscoreales
Dioscoreales
Brown 1835

Pandanales
Pandanales
Brown ex von Berchtold & Presl 1820

Liliales
Liliales
Perleb 1826

Asparagales
Asparagales
Link 1829

Commelinids

Arecales
Arecales
Bromhead 1840

Poales
Poales
Small 1903

Zingiberales
Zingiberales
Grisebach 1854

Commelinales
Commelinales
de Mirbel ex von Berchtold & Presl 1820

Ceratophyllales
Ceratophyllales
Link 1829

Eudicots

Ranunculales
Ranunculales
de Jussieu ex von Berchtold & Presl 1820

Proteales
Proteales
de Jussieu ex von Berchtold & Presl 1820

Trochodendrales
Trochodendrales
Takhtajan
Takhtajan
ex Cronquist 1981

Buxales
Buxales
Takhtajan
Takhtajan
ex Reveal 1996

Core eudicots

Gunnerales
Gunnerales
Takhtajan
Takhtajan
ex Reveal 1992

Dilleniales
Dilleniales
de Candolle ex von Berchtold & Presl 1820

Superrosids

Saxifragales
Saxifragales
von Berchtold & Presl 1820

Rosids

Vitales
Vitales
de Jussieu ex von Berchtold & Presl 1820

Fabids

Zygophyllales
Zygophyllales
Link 1829

Celastrales
Celastrales
Link 1829

Oxalidales
Oxalidales
von Berchtold & Presl 1820

Malpighiales
Malpighiales
de Jussieu ex von Berchtold & Presl 1820

Fabales
Fabales
Bromhead 1838

Rosales
Rosales
von Berchtold & Presl 1820

Cucurbitales
Cucurbitales
de Jussieu ex von Berchtold & Presl 1820

Fagales
Fagales
Engler 1892

(eurosids I)

Malvids

Geraniales
Geraniales
de Jussieu ex von Berchtold & Presl 1820

Myrtales
Myrtales
de Jussieu ex von Berchtold & Presl 1820

Crossosomatales
Crossosomatales
Takhtajan
Takhtajan
ex Reveal 1993

Picramniales
Picramniales
Doweld 2001

Sapindales
Sapindales
de Jussieu ex von Berchtold & Presl 1820

Huerteales
Huerteales
Doweld 2001

Malvales
Malvales
de Jussieu ex von Berchtold & Presl 1820

Brassicales
Brassicales
Bromhead 1838

(eurosids II)

Superasterids

Berberidopsidales
Berberidopsidales
Doweld 2001

Santalales
Santalales
Brown ex von Berchtold & Presl 1820

Caryophyllales

Asterids

Cornales
Cornales
Link 1829

Ericales
Ericales
von Berchtold & Presl 1820

Lamiids

Icacinales
Icacinales
Van Tieghem 1900

Metteniusales
Metteniusales
Takhtajan
Takhtajan
1997

Garryales
Garryales
Mart. 1835

Gentianales
Gentianales
de Jussieu ex von Berchtold & Presl 1820

Solanales
Solanales
de Jussieu ex von Berchtold & Presl 1820

Boraginales
Boraginales
de Jussieu ex von Berchtold & Presl 1820

Vahliales Doweld 2001

Lamiales
Lamiales
Bromhead 1838

(euasterids I)

Campanulids

Aquifoliales
Aquifoliales
Senft 1856

Escalloniales
Escalloniales
Mart. 1835

Asterales
Asterales
Link 1829

Bruniales
Bruniales
Dumortier 1829

Apiales
Apiales
Nakai 1930

Paracryphiales
Paracryphiales
Takhtajan
Takhtajan
ex Reveal 1992

Dipsacales
Dipsacales
de Jussieu ex von Berchtold & Presl 1820

(euasterids II)

Evolution[edit] Further information: 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.[20] 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
Ginkgo
and cycads produce a pair of flagellated, mobile sperm cells that "swim" down the developing pollen tube to the female and her eggs.

Flowers of Malus sylvestris
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 evolution that Charles Darwin
Charles Darwin
called it an "abominable mystery".[21] 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
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
Triassic
period (245–202 million years ago). Fossil
Fossil
angiosperm-like pollen from the Middle Triassic
Triassic
(247.2–242.0 Ma) suggests an older date for their origin.[22] 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.[citation needed] The evolution of seed plants and later angiosperms appears to be the result of two distinct rounds of whole genome duplication events.[23] 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.[24] That event was studied by sequencing the genome of an ancient flowering plant, Amborella trichopoda,[25] and directly addresses Darwin's "abominable mystery."

Flowers and leaves of Oxalis pes-caprae
Oxalis pes-caprae
(Bermuda buttercup)

The earliest known macrofossil confidently identified as an angiosperm, Archaefructus
Archaefructus
liaoningensis, is dated to about 125 million years BP (the Cretaceous
Cretaceous
period),[26] 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
Jurassic
plant Schmeissneria, traditionally considered a type of ginkgo, may be the earliest known angiosperm, or at least a close relative.[27] 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
Permian
deposits of that age together with fossils of gigantopterids.[28][29] 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, suggesting it was carried by insects.[30] Recent DNA
DNA
analysis based on molecular systematics[31][32] showed that Amborella
Amborella
trichopoda, found on the Pacific island of New Caledonia, belongs to a sister group of the other flowering plants, and morphological studies[33] suggest that it has features that may have been characteristic of the earliest flowering plants. The orders Amborellales, Nymphaeales, and Austrobaileyales
Austrobaileyales
diverged as separate lineages from the remaining angiosperm clade at a very early stage in flowering plant evolution.[34] 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.[35] 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
Cretaceous
66 million years ago or even later, at the beginning of the Tertiary.[36] The radiation of herbaceous angiosperms occurred much later.[37] 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
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]

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
Island genetics
provides one proposed explanation for the sudden, fully developed appearance of flowering plants. Island genetics
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.[39] 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
Flower
ontogeny uses a combination of genes normally responsible for forming new shoots.[40] 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
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.[41] 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.[42] 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.[43][44] 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.[45] Flowering plant
Flowering plant
diversity[edit]

A poster of twelve different species of flowers of the Asteraceae family

Lupinus pilosus

Bud
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.[46][47][48] This compares to around 12,000 species of moss[49] or 11,000 species of pteridophytes,[50] 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][51] 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,[52] in their APG circumscriptions, are

Asteraceae
Asteraceae
or Compositae (daisy family): 22,750 species; Orchidaceae
Orchidaceae
(orchid family): 21,950; Fabaceae
Fabaceae
or Leguminosae (bean family): 19,400; Rubiaceae
Rubiaceae
(madder family): 13,150;[53] Poaceae
Poaceae
or Gramineae (grass family): 10,035; Lamiaceae
Lamiaceae
or Labiatae (mint family): 7,175; Euphorbiaceae
Euphorbiaceae
(spurge family): 5,735; Melastomataceae
Melastomataceae
or Melastomaceae (melastome family): 5,005; Myrtaceae
Myrtaceae
(myrtle family): 4,625; Apocynaceae
Apocynaceae
(dogbane family): 4,555; Cyperaceae
Cyperaceae
(sedge family): 4,350; Malvaceae
Malvaceae
(mallow family): 4,225; Araceae
Araceae
(arum family): 4,025; Ericaceae
Ericaceae
(heath family): 3,995; Gesneriaceae
Gesneriaceae
(gesneriad family): 3,870; Apiaceae
Apiaceae
or Umbelliferae (parsley family): 3,780; Brassicaceae
Brassicaceae
or Cruciferae (cabbage family): 3,710: Piperaceae
Piperaceae
(pepper family): 3,600; Bromeliaceae
Bromeliaceae
(bromeliad family): 3,540; Acanthaceae
Acanthaceae
(acanthus family): 3,500; Rosaceae
Rosaceae
(rose family): 2,830; Boraginaceae
Boraginaceae
(borage family): 2,740; Urticaceae
Urticaceae
(nettle family): 2,625; Ranunculaceae
Ranunculaceae
(buttercup family): 2,525; Lauraceae
Lauraceae
(laurel family): 2,500; Solanaceae
Solanaceae
(nightshade family): 2,460; Campanulaceae
Campanulaceae
(bellflower family): 2,380; Arecaceae
Arecaceae
(palm family): 2,361; Annonaceae
Annonaceae
(custard apple family): 2,220; Caryophyllaceae
Caryophyllaceae
(pink family): 2,200; Orobanchaceae
Orobanchaceae
(broomrape family): 2,060; Amaranthaceae
Amaranthaceae
(amaranth family): 2,050; Iridaceae
Iridaceae
(iris family): 2,025; Aizoaceae
Aizoaceae
or Ficoidaceae (ice plant family): 2,020; Rutaceae
Rutaceae
(rue family): 1,815; Phyllanthaceae
Phyllanthaceae
(phyllanthus family): 1,745; Scrophulariaceae
Scrophulariaceae
(figwort family): 1,700; Gentianaceae
Gentianaceae
(gentian family): 1,650; Convolvulaceae
Convolvulaceae
(bindweed family): 1,600; Proteaceae
Proteaceae
(protea family): 1,600; Sapindaceae
Sapindaceae
(soapberry family): 1,580; Cactaceae
Cactaceae
(cactus family): 1,500; Araliaceae
Araliaceae
( Aralia
Aralia
or ivy family): 1,450.

Of these, the Orchidaceae, Poaceae, Cyperaceae, Araceae, Bromeliaceae, Arecaceae, and Iridaceae
Iridaceae
are monocot families; Piperaceae, Lauraceae, and Annonaceae
Annonaceae
are magnoliid dicots; the rest of the families are eudicots. Reproduction[edit] Fertilization
Fertilization
and embryogenesis[edit] Main articles: Fertilization
Fertilization
and Plant
Plant
embryogenesis

Angiosperm life cycle

Double fertilization
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
Fruit
and seed[edit] Main articles: Seed
Seed
and Fruit

The fruit of the Aesculus
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.[54] 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.

Meiosis[edit] Flowering plants generate gametes using a specialized cell division called meiosis. 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.[55] 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
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
DNA
to be passed on to progeny. To explain the adaptive function of meiosis in flowering plants, some authors emphasize diversity[56] and others emphasize DNA repair.[57] Apomixis[edit] Apomixis
Apomixis
(reproduction via asexually formed seeds) is found naturally in about 2.2% of angiosperm genera [58] One type of apomixis, gametophytic apomixis found in a dandelion species [59] 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[edit] Agriculture
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.[60] 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]

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

References[edit]

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Biology. 30: 159–165. doi:10.1016/j.pbi.2016.03.015.  Takhtajan, A. (June 1964). "The Taxa of the Higher Plants above the Rank of Order". Taxon. 13 (5): 160–164. doi:10.2307/1216134. JSTOR 10.2307/1216134.  Takhtajan, A. (July–September 1980). "Outline of the Classification of Flowering Plants (Magnoliophyta)". Botanical Review. 46 (3): 225–359. doi:10.1007/bf02861558. JSTOR 10.2307/4353970.  Zeng, Liping; Zhang, Qiang; Sun, Renran; Kong, Hongzhi; Zhang, Ning; Ma, Hong (24 September 2014). "Resolution of deep angiosperm phylogeny using conserved nuclear genes and estimates of early divergence times". Nature Communications. 5 (4956). doi:10.1038/ncomms5956. 

Websites[edit]

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

External links[edit]

Wikimedia Commons has media related to Magnoliophyta.

Wikispecies
Wikispecies
has information related to Magnoliophyta

The Wikibook Dichotomous Key has a page on the topic of: Magnoliophyta

v t e

Classification of Archaeplastida
Archaeplastida
/ Plantae
Plantae
sensu lato

Domain Archaea Bacteria Eukaryota (Supergroup Plant Hacrobia Heterokont Alveolata Rhizaria Excavata Amoebozoa Opisthokonta

Animal Fungi)

Rhodophyta (red algae)

Cyanidiophyceae Porphyridiophyceae Compsopogonophyceae Stylonematophyceae Rhodellophyceae Bangiophyceae Florideophyceae

Glaucocystophyta (glaucophytes)

Glaucocystophyceae

Glaucocystis Cyanophora Gloeochaete

Viridiplantae (green algae, & land plants)

Mesostigmatophyceae Chlorokybophyceae

C+S

Chlorophyta

Palmophyllales Nephrophyceae Prasinophyceae Pseudoscourfieldiales Pyramimonadophyceae Scourfieldiales Pedinophyceae Chlorodendrophyceae UTC clade

Ulvophyceae Trebouxiophyceae Chlorophyceae

Streptophyta (charophytes, & land plants)

Klebsormidiophyceae

Phragmo- plastophyta

Charophyceae Coleochaetophyceae Zygnematophyceae

Embryophyta (land plants)

Bryophytes (non-vascular)

Marchantiophyta Anthocerotophyta Bryophyta "Moss" †Horneophytopsida

Tracheophyta (vascular)

Lycopodiophyta (microphylls)

†Zosterophyllopsida †Sawdoniales Isoetopsida Lycopodiopsida

Euphyllophyta (megaphylls)

Moniliformopses (ferns)

†Cladoxylopsida †Stauropteridales †Zygopteridales Equisetopsida Psilotopsida Marattiopsida Polypodiopsida

Spermatophyta (seed plants)

Seed
Seed
ferns Gymnosperms

Gnetopsida Pinopsida Cycadopsida Ginkgoopsida

Angiosperms
Angiosperms
or flowering plants

Amborellales Nymphaeales Austrobaileyales Magnoliids Monocots Eudicots

Other

†Trimerophytopsida †Progymnosperm

Other

†Rhyniopsida

† = extinct. See also the list of plant orders.

v t e

Botany

History of botany

Subdisciplines

Plant
Plant
systematics Ethnobotany Paleobotany Plant
Plant
anatomy Plant
Plant
ecology Phytogeography

Geobotany Flora

Phytochemistry Plant
Plant
pathology Bryology Phycology Floristics Dendrology

Plant
Plant
groups

Algae Archaeplastida Bryophyte Non-vascular plants Vascular plants Spermatophytes Pteridophyte Gymnosperm Angiosperm

Plant
Plant
morphology (glossary)

Plant
Plant
cells

Cell wall Phragmoplast Plastid Plasmodesma Vacuole

Tissues

Meristem Vascular tissue

Vascular bundle

Ground tissue

Mesophyll

Cork Wood Storage organs

Vegetative

Root Rhizoid Bulb Rhizome Shoot

Stem Leaf

Petiole Cataphyll

Bud Sessility

Reproductive (Flower)

Flower
Flower
development Inflorescence

Umbel Raceme Bract Pedicellate

Flower

Whorl Floral symmetry Floral diagram Floral formula

Receptacle Hypanthium
Hypanthium
(Floral cup) Perianth

Tepal Petal Sepal

Sporophyll Gynoecium

Ovary

Ovule

Stigma

Archegonium Androecium

Stamen Staminode Pollen Tapetum

Gynandrium Gametophyte Sporophyte Plant
Plant
embryo Fruit

Fruit
Fruit
anatomy Berry Capsule Seed

Seed
Seed
dispersal Endosperm

Surface structures

Epicuticular wax Plant
Plant
cuticle Epidermis Stoma Nectary Trichome Prickle

Plant
Plant
physiology Materials

Nutrition Photosynthesis

Chlorophyll

Plant
Plant
hormone Transpiration Turgor pressure Bulk flow Aleurone Phytomelanin Sugar Sap Starch Cellulose

Plant
Plant
growth and habit

Secondary growth Woody plants Herbaceous plants Habit

Vines

Lianas

Shrubs

Subshrubs

Trees Succulent plants

Reproduction

Evolution Ecology

Alternation of generations Sporangium

Spore Microsporangia

Microspore

Megasporangium

Megaspore

Pollination

Pollinators Pollen
Pollen
tube

Double fertilization Germination Evolutionary development Evolutionary history

timeline

Hardiness zone

Plant
Plant
taxonomy

History of plant systematics Herbarium Biological classification Botanical nomenclature

Botanical name Correct name Author citation International Code of Nomenclature for algae, fungi, and plants
International Code of Nomenclature for algae, fungi, and plants
(ICN) - for Cultivated Plants (ICNCP)

Taxonomic rank International Association for Plant
Plant
Taxonomy (IAPT) Plant
Plant
taxonomy systems Cultivated plant taxonomy

Citrus taxonomy cultigen

cultivar Group grex

Practice

Agronomy Floriculture Forestry Horticulture

Lists Related topics

Botanical terms Botanists

by author abbreviation

Botanical expedition

Category Portal WikiProject

Taxon identifiers

Wd: Q25314 EoL: 282 EPPO: 1ANGC

Authority control

GND: 41442

.