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Flowering plants include multiple members of the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the
Greek#REDIRECT Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Europe. Its population is approximately 10.7 million as of ...
words ('container, vessel') and ('seed'), and refers to those plants that produce their
seeds A seed is an embryonic ''Embryonic'' is the twelfth studio album by experimental rock band the Flaming Lips released on October 13, 2009, on Warner Bros. Records, Warner Bros. The band's first double album, it was released to generally positiv ...

seeds
enclosed within a
fruit In botany Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the ...

fruit
. They are the most diverse group of
land plants The Embryophyta (), or land plants, are the most familiar group of green plants that comprise vegetation on Earth. Embryophyta is a clade within the Phragmoplastophyta, a larger clade that also includes several groups of green algae including the ...

land plants
with 64
orders Orders is a surname In some cultures, a surname, family name, or last name is the portion of one's personal name that indicates their family, tribe or community. Practices vary by culture. The family name may be placed at either the start of ...
, 416
families In human society, family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of families is to maintain the w ...
, approximately 13,000 known
genera Genus /ˈdʒiː.nəs/ (plural genera /ˈdʒen.ər.ə/) is a taxonomic rank Taxonomy (general) is the practice and science of classification of things or concepts, including the principles that underlie such classification. The term may also ref ...
and 300,000 known
species In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interactions, Physiology, physiological mechanis ...

species
. Angiosperms were formerly called Magnoliophyta (). Like
gymnosperm The gymnosperms ( Greek language, lit. revealed seeds) are a group of Spermatophyte, seed-producing plants that includes Pinophyta, conifers, cycads, ''Ginkgo'', and gnetophyta, gnetophytes, forming the clade Gymnospermae, the living members of ...
s, angiosperms are seed-producing plants. They are distinguished from gymnosperms by characteristics including
flower A flower, sometimes known as a bloom or blossom, is the reproduction, reproductive structure found in flowering plants (plants of the division Magnoliophyta, also called angiosperms). The biological function of a flower is to facilitate reproduct ...

flower
s,
endosperm 350px, right The endosperm is a tissue produced inside the seed A seed is an embryonic ''Embryonic'' is the twelfth studio album by experimental rock band the Flaming Lips released on October 13, 2009, on Warner Bros. Records, Warner Bros. The ...
within their seeds, and the production of fruits that contain the seeds. The ancestors of flowering plants diverged from the common ancestor of all living gymnosperms during the
Carboniferous The Carboniferous ( ) is a geologic period A geological period is one of the several subdivisions of geologic time enabling cross-referencing of rocks and geologic events from place to place. These periods form elements of a hierarchy of divisi ...
, over 300 million years ago, with the earliest record of angiosperm pollen appearing around 134 million years ago. The first remains of flowering plants are known from 125 million years ago. They diversified extensively during the
Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous ( chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous The Cretaceous (, ) is a geological period that lasted from about 145 to 66 mi ...
, became widespread by 120 million years ago, and replaced
conifers Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All exta ...

conifers
as the dominant trees from 60 to 100 million years ago.


Description


Angiosperm derived characteristics

Angiosperms differ from other
seed plants The spermatophytes (; ), also known as phanerogams (taxon Phanerogamae) or phaenogams (taxon Phaenogamae), comprise those plant Plants are predominantly photosynthetic eukaryotes of the Kingdom (biology), kingdom Plantae. Historically, the pl ...
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.


Vascular anatomy

Angiosperm stems are made up of seven layers as shown on the right. The amount and
complexity Complexity characterises the behaviour of a system or model (disambiguation), model whose components interaction, interact in multiple ways and follow local rules, meaning there is no reasonable higher instruction to define the various possible in ...
of tissue-formation in flowering plants exceeds that of gymnosperms. In the
dicotyledon The dicotyledons, also known as dicots (or more rarely dicotyls), are one of the two groups into which all the flowering plant The flowering plants, also known as Angiospermae (), or Magnoliophyta (), are the most diverse group of Embryophyte ...
s, the
vascular bundle Cross section of celery stalk, showing vascular bundles, which include both phloem and xylem Detail of vascular bundle: closed, collateral vascular bundles of the stem axis of '' Zea mays'' A vascular bundle is a part of the transport system ...
s of the stem are arranged such that the
xylem Xylem is one of the two types of transport tissue Tissue may refer to: Biology * Tissue (biology), an ensemble of similar cells that together carry out a specific function * ''Triphosa haesitata'', a species of geometer moth found in North Amer ...

xylem
and
phloem Phloem (, ) is the living tissue Tissue may refer to: Biology * Tissue (biology), an ensemble of similar cells that together carry out a specific function * ''Triphosa haesitata'', a species of geometer moth found in North America * ''Triphosa d ...

phloem
form concentric rings. 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 The meristem is a type of tissue Tissue may refer to: Biology * Tissue (biology), an ensemble of similar cells that together carry out a specific function * ''Triphosa haesitata'', a species of geometer moth found in North America * ''Triphosa dub ...
or active formative tissue known as
cambium A cambium (plural cambia or cambiums), in plants, is a tissue layer that provides partially undifferentiated cell Cell most often refers to: * Cell (biology), the functional basic unit of life Cell may also refer to: Closed spaces * Monastic ...
. By the formation of a layer of
cambium A cambium (plural cambia or cambiums), in plants, is a tissue layer that provides partially undifferentiated cell Cell most often refers to: * Cell (biology), the functional basic unit of life Cell may also refer to: Closed spaces * Monastic ...
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 A season is a division of the year based on changes in weather Weather is the state of the atmosphere, describing for example the degree to which it is hot or cold, wet or dry, calm or stormy, clear or cloud cover, cloudy. On Earth, most ...

season
of growth, called
annual rings Dendrochronology (or tree-ring dating) is the scientific method The scientific method is an Empirical evidence, empirical method of acquiring knowledge that has characterized the development of science since at least the 17th century. It in ...
. Among the
monocotyledon Monocotyledons (), commonly referred to as monocots, (Lilianae ''sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one Embryo#Plant embryos, embryonic leaf, or cotyledon. The ...
s, 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 Fertilisation or fertilization (see spelling differences Despite the various English dialects spoken from country to country and within different regions of the same country, there are only slight regional variations in English orth ...

fertilization
of the
ovule In seed plant The spermatophytes, also known as phanerogams (taxon Phanerogamae) or phaenogams (taxon Phaenogamae), comprise those plants that produce seeds, hence the alternative name seed plants. They are a subset of the embryophytes or l ...

ovule
and development of
fruit In botany Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the ...

fruit
containing
seed A seed is an embryonic ''Embryonic'' is the twelfth studio album by experimental rock band the Flaming Lips released on October 13, 2009, on Warner Bros. Records, Warner Bros. The band's first double album, it was released to generally positiv ...

seed
s. The floral apparatus may arise terminally on a shoot or from the
axil A leaf (plural leaves) is the principal lateral appendage of the vascular plant plant stem, stem, usually borne above ground and specialized for photosynthesis. The leaves and stem together form the shoot. Leaves are collectively referred ...
of a leaf (where the
petiole Petiole may refer to: *Petiole (botany), the stalk of a leaf, attaching the blade to the stem *Petiole (insect anatomy), the narrow waist of some hymenopteran insects {{disambiguation ...
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 An inflorescence is a group or cluster of flower A flower, sometimes known as a bloom or blossom, is the reproduction, reproductive structure found in flowering plants (plants of the division Magnoliophyta, also called angiosperms). The biol ...
. There are two kinds of reproductive cells produced by flowers.
Microspore Microspores are land plant The Embryophyta () or land plants are the most familiar group of green plants that form vegetation on earth. Embryophyta is a clade within the Phragmoplastophyta, a larger clade that also includes several green algae g ...
s, which will divide to become
pollen grains
pollen grains
, are the "male" cells and are borne in the
stamen The stamen (plural The plural (sometimes abbreviated An abbreviation (from Latin ''brevis'', meaning ''short'') is a shortened form of a word or phrase, by any method. It may consist of a group of letters, or words taken from the full vers ...
s (or microsporophylls). The "female" cells called
megaspore Megaspores, also called macrospores, are a type of spore )'', growing on a thinned hybrid black poplar ''(Populus x canadensis)''. The last stage of the moss lifecycle is shown, where the sporophytes are visible before dispersion of their sp ...

megaspore
s, which will divide to become the egg cell ( megagametogenesis), are contained in the
ovule In seed plant The spermatophytes, also known as phanerogams (taxon Phanerogamae) or phaenogams (taxon Phaenogamae), comprise those plants that produce seeds, hence the alternative name seed plants. They are a subset of the embryophytes or l ...

ovule
and enclosed in the
carpel Gynoecium (; ) is most commonly used as a collective term for the parts of a flower A flower, sometimes known as a bloom or blossom, is the reproduction, reproductive structure found in flowering plants (plants of the division Magnoliophyta, ...
(or megasporophyll). The flower may consist only of these parts, as in
willow Willows, also called sallows and osiers, from the genus Genus /ˈdʒiː.nəs/ (plural genera /ˈdʒen.ər.ə/) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classifica ...

willow
, where each flower comprises only a few
stamen The stamen (plural The plural (sometimes abbreviated An abbreviation (from Latin ''brevis'', meaning ''short'') is a shortened form of a word or phrase, by any method. It may consist of a group of letters, or words taken from the full vers ...
s or two carpels. Usually, other structures are present and serve to protect the
sporophyll A sporophyll is a leaf A leaf (plural leaves) is the principal lateral appendage of the vascular plant plant stem, stem, usually borne above ground and specialized for photosynthesis. The leaves and stem together form the shoot. Leaves ar ...
s and to form an envelope attractive to pollinators. The individual members of these surrounding structures are known as
sepal upright=1.4, Diagram showing the parts of a mature flower. In this example the perianth is separated into a calyx (sepals) and corolla (petals) A sepal ( or ) is a part of the flower A flower, sometimes known as a bloom or blossom, is the ...
s and
petal upright=1.4, Diagram showing the parts of a mature flower. In this example the perianth is separated into a calyx (sepals) and corolla (petals) Petals are modified leaves A leaf (plural leaves) is the principal lateral appendage of the ...

petal
s (or
tepal A tepal is one of the outer parts of a flower A flower, sometimes known as a bloom or blossom, is the reproduction, reproductive structure found in flowering plants (plants of the division Magnoliophyta, also called angiosperms). The biological ...
s in flowers such as ''
Magnolia ''Magnolia'' is a large genus of about 210The number of species in the genus ''Magnolia'' depends on the taxonomic view that one takes up. Recent molecular and morphology (biology), morphological research shows that former genera ''Talauma'', ''D ...

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 Insects (from Latin ') are pancrustacean Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, Thorax (inse ...

insect
or
bird Birds are a group of warm-blooded vertebrates constituting the class Class or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of indiv ...

bird
pollinators. Attraction is effected by color,
scent An odor (American English American English (AmE, AE, AmEng, USEng, en-US), sometimes called United States English or U.S. English, is the set of varieties of the English language native to the United States. Currently, American English ...
, and
nectar Nectar is a sugar Sugar is the generic name for Sweetness, sweet-tasting, soluble carbohydrates, many of which are used in food. Table sugar, granulated sugar, or regular sugar, refers to sucrose, a disaccharide composed of glucose and fru ...

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 In reproductive biology Reproductive biology includes both sexual and asexual reproduction. Reproductive biology includes a wide number of fields: * Reproductive systems * Endocrinology Endocrinology (from '' endocrine'' + '' -ology'') is a b ...

hermaphrodite
(having both pollen and ovule producing parts in the same flower structure), flowering plants have developed numerous morphological and
physiological Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a sub-discipline of biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, B ...
mechanisms to reduce or prevent self-fertilization. Heteromorphic flowers have short carpels and long stamens, or vice versa, so animal
pollinator A pollinator is an animal that moves pollen File:Pollen Tube.svg, Pollen Tube Diagram Pollen is a powdery substance consisting of pollen grains which are Sporophyte, microsporophytes of spermatophyta, seed plants, which produce male gametes ...

pollinator
s cannot easily transfer pollen to the pistil (receptive part of the carpel). Homomorphic flowers may employ a biochemical (physiological) mechanism called
self-incompatibilitySelf-incompatibility (SI) is a general name for several genetic mechanisms in angiosperms The flowering plants, also known as Angiospermae (), or Magnoliophyta (), are the most diverse group of land plants, with 64 orders, 416 families, approxi ...
to discriminate between self and non-self pollen grains. Alternatively, in
dioecious Dioecy (; Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Europe. Its population is approximately 10.7 million as o ...
species, the male and female parts are morphologically separated, developing on different individual flowers.


Taxonomy


History of classification

The botanical term "angiosperm", from Greek words ( 'bottle, vessel') and ( '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 Kingdom may refer to: Monarchy * A type of monarchy * A realm ruled by: **A king, during the reign of a male monarch **A queen regnant, during the reign of a female monarch Taxonomy * Kingdom (biology), a category in biological taxonomy Arts an ...
. This included flowering plants possessing seeds enclosed in capsules, distinguished from his Gymnospermae, or flowering plants with or schizo-carpic fruits, the whole fruit or each of its pieces being here regarded as a seed and naked. Both the term and its antonym were maintained by
Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his Nobility#Ennoblement, ennoblement as Carl von Linné#Blunt, Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomia ...

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 and , 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
dicotyledon The dicotyledons, also known as dicots (or more rarely dicotyls), are one of the two groups into which all the flowering plant The flowering plants, also known as Angiospermae (), or Magnoliophyta (), are the most diverse group of Embryophyte ...
ous 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, with
Anthophyta The anthophytes are a grouping of plant taxa bearing flower-like reproductive structures. They were formerly thought to be a clade A clade (; from grc, , ''klados'', "branch"), also known as a monophyletic group or natural group, is a group ...
(lit. 'flower-plants') a second choice (both unranked). The
Wettstein systemA system of plant taxonomy, the Wettstein system recognised the following main groups, according to Richard Wettstein's ''Handbuch der Systematischen Botanik'' (1901–1924). 3rd edition (1924) Outline Synopsis * Flagellatae p. 65 * ...
and
Engler system One of the prime systems of plant taxonomy, the Engler system was devised by Adolf Engler Heinrich Gustav Adolf Engler (25 March 1844 – 10 October 1930) was a German botanist. He is notable for his work on plant taxonomy and phytogeography, ...
treated them as a subdivision (Angiospermae). The
Reveal system A 20th-century system of plant taxonomy, the Reveal system (see also the Thorne & Reveal system) of plant classification was drawn up by the American botanist James Reveal (1941-2015). The system was published online in 1997 in ten parts as lectur ...
also treated them as a subdivision (Magnoliophytina), but later split it to Magnoliopsida, Liliopsida, and Rosopsida. The
Takhtajan system Armen Leonovich Takhtajan or Takhtajian ( hy, Արմեն Լևոնի Թախտաջյան; russian: Армен Леонович Тахтаджян; surname also transliterated Takhtadjan, Takhtadzhi︠a︡n or Takhtadzhian, pronounced TAHK-tuh-jahn) ...
and
Cronquist systemCronquist can refer to: * Arthur J. Cronquist, a North American botanist (1919–1992). * The Cronquist system, a system attributed to Arthur J. Cronquist. Many authors use their own variation of this system, which they also refer to as the ...
treat them as a division (Magnoliophyta). The
Dahlgren systemOne of the modern systems of plant taxonomy, the Dahlgren system was published by monocot Monocotyledons (), commonly referred to as monocots, (Lilianae ''sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds o ...
and
Thorne system (1992)A system of plant taxonomy, the Thorne system of plant classification was devised by the American botanist Robert F. Thorne (1920–2015) in 1968, and he continued to issue revisions over many years (1968–2007). Some versions of the system are a ...
treat them as a class (Magnoliopsida). The
APG systemThe APG system (Angiosperm Phylogeny Group system) of plant classification is the first version of a modern, mostly molecular-based, system of plant taxonomy. Published in 1998 by the Angiosperm Phylogeny Group, it was replaced by the improved AP ...
of 1998, and the later 2003 and 2009 revisions, treat the flowering plants as an unranked clade without a formal Latin name (angiosperms). A formal classification was published alongside the 2009 revision in which the flowering plants rank as a subclass (Magnoliidae). The internal classification of this group has undergone considerable revision. The
Cronquist systemCronquist can refer to: * Arthur J. Cronquist, a North American botanist (1919–1992). * The Cronquist system, a system attributed to Arthur J. Cronquist. Many authors use their own variation of this system, which they also refer to as the ...
, proposed by
Arthur Cronquist Arthur John Cronquist (March 19, 1919 – March 22, 1992) was an American biologist, botanist Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytolo ...
in 1968 and published in its full form in 1981, is still widely used but is no longer believed to accurately reflect
phylogeny , based on completely sequenced genomes. A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolution ...

phylogeny
. A consensus about how the flowering plants should be arranged has recently begun to emerge through the work of the
Angiosperm Phylogeny Group The Angiosperm Phylogeny Group, or APG, is an informal international group of systematic botanists who collaborate to establish a consensus on the taxonomy of flowering plant The flowering plants, also known as Angiospermae (), or Magnoliophyt ...
(APG), which published an influential reclassification of the angiosperms in 1998. Updates incorporating more recent research were published as the APG II system in 2003, the
APG III system The APG III system of flowering plant classification is the third version of a modern, mostly molecular-based, system of plant taxonomy being developed by the Angiosperm Phylogeny Group (APG). Published in 2009, it was superseded in 2016 by a fu ...
in 2009, and the
APG IV systemAPG is an abbreviation with several different meanings: * Aberdeen Proving Ground, a United States Army installation in Aberdeen, Maryland, also ** Phillips Army Airfield, the airfield of the above, from its IATA airport code * Aboriginal Provision ...
in 2016. Traditionally, the flowering plants are divided into two groups, * Dicotyledoneae or
Magnoliopsida Magnoliopsida is a valid botanical name for a class of flowering plant The flowering plants, also known as Angiospermae (), or Magnoliophyta (), are the most diverse group of Embryophyte, land plants, with 64 Order(biology), orders, 416 Family ...
*
Monocotyledoneae Monocotyledons (), commonly referred to as monocots, (Lilianae ''sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one Embryo#Plant embryos, embryonic leaf, or cotyledon. The ...
or
Liliopsida Liliopsida Batsch (synonym (taxonomy), synonym: Liliatae) is a botanical name for the Class (biology), class containing the Family (biology), family Liliaceae (or Lily Family). It is considered synonymous (or nearly synonymous) with the name monoc ...
to which the Cronquist system ascribes the classes Magnoliopsida (from "
Magnoliaceae The Magnoliaceae () are a flowering plant The flowering plants, also known as Angiospermae (), or Magnoliophyta (), are the most diverse group of Embryophyte, land plants, with 64 Order(biology), orders, 416 Family (biology), families, approxim ...
" and Liliopsida (from "
Liliaceae The lily family In human society A society is a Social group, group of individuals involved in persistent Social relation, social interaction, or a large social group sharing the same spatial or social territory, typically subject ...

Liliaceae
"). Other descriptive names allowed by Article 16 of the
ICBN The ''International Code of Nomenclature for algae, fungi, and plants'' (ICN) is the set of rules and recommendations dealing with the formal botanical names that are given to plants, fungi and a few other groups of organisms, all those "tradi ...
include Dicotyledones or Dicotyledoneae, and Monocotyledones or Monocotyledoneae, which have a long history of use. In plain English, their members may be called "dicotyledons" ("dicots") and "monocotyledons" ("monocots"). The Latin behind these names refers the observation that the dicots most often have two
cotyledon File:Germination-en.svg, 335px, Schematic of epigeal vs hypogeal germination A cotyledon (; ; ; , gen. (), ) is a significant part of the embryo within the seed of a plant, and is defined as "the embryonic leaf in seed-bearing plants, one o ...
s, 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 300px, A cladogram of the primates, showing a ''monophyletic'' taxon: ''the simians'' (in yellow); a ''paraphyletic'' taxon: ''the prosimians'' (in cyan, including the red patch); and a ''polyphyletic'' group: ''the night-active primates, i.e., ...

monophyletic
group (a
clade A clade (), also known as a monophyletic group or natural group, is a group of organism In biology, an organism () is any organic, life, living system that functions as an individual entity. All organisms are composed of cells (cell t ...

clade
) but that the dicots are
paraphyletic In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and all descendants of that ancestor excluding a few—typically only one or two—Monophyly, monophyletic subgroups. The group is said to be paraphyleti ...

paraphyletic
. Nevertheless, the majority of dicot species fall into a clade, the
eudicots The eudicots, Eudicotidae or eudicotyledons are a clade A clade (), also known as a monophyletic group or natural group, is a group of organism In biology, an organism () is any organic, life, living system that functions as an ind ...
or tricolpates, and most of the remaining fall into another major clade, the
magnoliids Magnoliids (or Magnoliidae or Magnolianae) are a group of flowering plant The flowering plants, also known as Angiospermae (), or Magnoliophyta (), are the most diverse group of Embryophyte, land plants, with 64 Order(biology), orders, 416 Famil ...
, containing about 9,000 species. The rest include a paraphyletic grouping of early branching taxa known collectively as the
basal angiosperms '', from the Nymphaeales The basal angiosperms are the flowering plant The flowering plants, also known as Angiospermae (), or Magnoliophyta (), are the most diverse group of Embryophyte, land plants, with 64 Order(biology), orders, 416 Family (b ...
, plus the families Ceratophyllaceae and
Chloranthaceae Chloranthaceae is a family In human society, family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of families is to maintain t ...
.


Modern classification

There are eight groups of living angiosperms: *
Basal angiosperms '', from the Nymphaeales The basal angiosperms are the flowering plant The flowering plants, also known as Angiospermae (), or Magnoliophyta (), are the most diverse group of Embryophyte, land plants, with 64 Order(biology), orders, 416 Family (b ...
(ANA: ''Amborella'', Nymphaeales, Austrobaileyales) ** ''
Amborella ''Amborella'' is a monotypic In biology, a monotypic taxon is a taxonomic group ( taxon) that contains only one immediately subordinate taxon. A monotypic species In biology, a species is the basic unit of biological classification, cla ...

Amborella
'', a single species of shrub from
New Caledonia ) , anthem = " Soyons unis, devenons frères" , image_map = New Caledonia on the globe (small islands magnified) (Polynesia centered).svg , map_alt = Location of New Caledonia , map_caption = Location of New Caledonia , mapsize = 290px , s ...

New Caledonia
; **
Nymphaeales The Nymphaeales are an order of flowering plant The flowering plants, also known as Angiospermae (), or Magnoliophyta (), are the most diverse group of Embryophyte, land plants, with 64 Order(biology), orders, 416 Family (biology), families, a ...
, about 80 species,
Figure 2
/ref>
water lilies ''Water Lilies'' (or ''Nymphéas'', ) is a Serial imagery, series of approximately 250 oil paintings by French Impressionism, Impressionist Claude Monet (1840–1926). The paintings depict his Fondation Monet in Giverny, flower garden at Fond ...

water lilies
and
Hydatellaceae Hydatellaceae are a family In human society, family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of families is to maintain th ...
; **
Austrobaileyales Austrobaileyales is an order of flowering plants consisting of about 100 species of woody plants growing as trees, shrubs and lianas. Perhaps the most familiar species is '' Illicium verum'', from which comes the spice star anise. The order be ...

Austrobaileyales
, about 100 species of
woody plant A woody plant is a plant that produces wood as its structural tissue and thus has a hard stem. In cold climates, woody plants further survive winter or dry season The dry season is a yearly period of low rainfall, especially in the tropics ...
s from various parts of the world *
Core angiosperms Mesangiospermae (core angiosperms) is a clade of flowering plants (angiosperms), informally called "mesangiosperms". They are one of two main groups of angiosperms. It is a name created under the rules of the ''PhyloCode'' system of phylogenetic n ...
(Mesangiospermae) **
Chloranthales Chloranthaceae is a family In human society, family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of families is to maintain t ...
, 77 known species of aromatic plants with toothed leaves; **
Magnoliids Magnoliids (or Magnoliidae or Magnolianae) are a group of flowering plant The flowering plants, also known as Angiospermae (), or Magnoliophyta (), are the most diverse group of Embryophyte, land plants, with 64 Order(biology), orders, 416 Famil ...
, about 9,000 species, characterised by
trimerousMerosity (from the greek "méros," which means "having parts") refers to the number of component parts in a distinct whorl of a plant structure. It is most commonly used in the context of flowers where it refers to the number of sepals in a whorl o ...
flowers, pollen with one pore, and usually branching-veined leaves—for example
magnolia ''Magnolia'' is a large genus of about 210The number of species in the genus ''Magnolia'' depends on the taxonomic view that one takes up. Recent molecular and morphology (biology), morphological research shows that former genera ''Talauma'', ''D ...

magnolia
s,
bay laurel ''Laurus nobilis'' is an aromatic evergreen tree or large shrub A shrub (or bush, but this is more of a gardening term) is a small- to medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems abo ...
, and
black pepper Black pepper (''Piper nigrum'') is a flowering plant, flowering vine in the family Piperaceae, cultivated for its fruit, known as a peppercorn, which is usually dried and used as a spice and seasoning. When fresh and fully mature, the fruit is ...

black pepper
; **
Monocots Monocotyledons (), commonly referred to as monocots, (Lilianae ''sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one Embryo#Plant embryos, embryonic leaf, or cotyledon. The ...

Monocots
, about 70,000 species, characterised by trimerous flowers, a single
cotyledon File:Germination-en.svg, 335px, Schematic of epigeal vs hypogeal germination A cotyledon (; ; ; , gen. (), ) is a significant part of the embryo within the seed of a plant, and is defined as "the embryonic leaf in seed-bearing plants, one o ...
, pollen with one pore, and usually parallel-veined leaves—for example
grasses Poaceae () or Gramineae () is a large and nearly ubiquitous family In human society A society is a Social group, group of individuals involved in persistent Social relation, social interaction, or a large social group sharing the ...
,
orchid Orchidaceae ( ), commonly called the orchid family, is a diverse and widespread family In human society A society is a Social group, group of individuals involved in persistent Social relation, social interaction, or a large social ...

orchid
s, and ; ** ''
Ceratophyllum ''Ceratophyllum'' is a cosmopolitan genus Genus (plural genera) is a taxonomic rank Taxonomy (general) is the practice and science of classification of things or concepts, including the principles that underlie such classification. The term ...
'', about 6 species of
aquatic plant Aquatic plants are plant Plants are predominantly photosynthetic eukaryotes of the Kingdom (biology), kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; howeve ...
s, perhaps most familiar as
aquarium An aquarium (plural: ''aquariums'' or ''aquaria'') is a vivarium of any size having at least one transparent side in which aquatic plants or animals are kept and displayed. Fishkeepers use aquaria to keep fish Fish are Aquatic animal, ...

aquarium
plants; **
Eudicots The eudicots, Eudicotidae or eudicotyledons are a clade A clade (), also known as a monophyletic group or natural group, is a group of organism In biology, an organism () is any organic, life, living system that functions as an ind ...
, about 175,000 species, characterised by 4- or 5-merous flowers, pollen with three pores, and usually branching-veined leaves—for example
sunflower ''Helianthus'' () is a genus Genus /ˈdʒiː.nəs/ (plural genera /ˈdʒen.ər.ə/) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, vi ...

sunflower
s,
petunia ''Petunia'' is genus of 20 species of flowering plants of South American origin. The popular flower of the same name derived its epithet from the French, which took the word ''petun'', meaning "tobacco," from a Tupi–Guarani language. An Annual ...

petunia
,
buttercup ''Ranunculus'' is a large genus of about 600 species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined a ...

buttercup
,
apple An apple is an edible fruit produced by an apple tree (''Malus domestica''). Apple fruit tree, trees are agriculture, cultivated worldwide and are the most widely grown species in the genus ''Malus''. The tree originated in Central Asia, wher ...

apple
s, and
oak An oak is a tree or shrub A shrub (or bush, but this is more of a gardening term) is a small- to medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be deciduous o ...

oak
s. The exact relationships among these eight groups is not yet clear, although there is agreement that the first three groups to diverge from the ancestral angiosperm were
Amborellales ''Amborella'' is a monotypic genus Genus (plural genera) is a taxonomic rank Taxonomy (general) is the practice and science of classification of things or concepts, including the principles that underlie such classification. The term may als ...
,
Nymphaeales The Nymphaeales are an order of flowering plant The flowering plants, also known as Angiospermae (), or Magnoliophyta (), are the most diverse group of Embryophyte, land plants, with 64 Order(biology), orders, 416 Family (biology), families, a ...
, and Austrobaileyales (basal angiosperms) Of the remaining five groups (core angiosperms), the relationships among the three broadest groups remains unclear (magnoliids, monocots, and eudicots). Zeng and colleagues (Fig. 1) describe four competing schemes.The eudicots and monocots are the largest and most diversified, with ~ 75% and 20% of angiosperm species, respectively. Some analyses make the magnoliids the first to diverge, others the monocots. ''Ceratophyllum'' seems to group with the eudicots rather than with the monocots. The APG IV retained the overall higher order relationship described in APG III.


Evolutionary history


Paleozoic

Fossilised
spore )'', growing on a thinning, thinned hybrid black poplar ''(populus, Populus x canadensis)''. The last stage of the moss#Life cycle, moss lifecycle is shown, where the sporophytes are visible before dispersion of their spores: the calyptra (1) is ...
s suggest that land plants (
embryophyte The Embryophyta (), or land plants, are the most familiar group of green plant Plants are predominantly photosynthetic eukaryotes of the Kingdom (biology), kingdom Plantae. Historically, the plant kingdom encompassed all living things that w ...

embryophyte
s) have existed for at least 475 million years. Early land plants reproduced sexually with flagellated, swimming sperm, like the green algae from which they evolved. An adaptation to terrestrialization was the development of upright meiosporangia for dispersal by
spore )'', growing on a thinning, thinned hybrid black poplar ''(populus, Populus x canadensis)''. The last stage of the moss#Life cycle, moss lifecycle is shown, where the sporophytes are visible before dispersion of their spores: the calyptra (1) is ...
s to new habitats. This feature is lacking in the descendants of their nearest algal relatives, the 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 thumbnail, 180px, Wallace's Selaginella (''Selaginella wallacei'') ''Selaginella'' is the sole genus of vascular plant Vascular plants (from Latin ''vasculum'': duct), also known as Tracheophyta (the tracheophytes , from the Greek ''trācheia ...
'', the spike-moss. The result for the ancestors of angiosperms was enclosing them in a case, the seed. The apparently sudden appearance in the fossil record of nearly modern flowers, and in great diversity, initially posed such a problem for the theory of gradual
evolution Evolution is change in the heritable Heredity, also called inheritance or biological inheritance, is the passing on of Phenotypic trait, traits from parents to their offspring; either through asexual reproduction or sexual reproduction, ...

evolution
that
Charles Darwin Charles Robert Darwin (; ; 12 February 1809 – 19 April 1882) was an English natural history#Before 1900, naturalist, geologist and biologist, best known for his contributions to the science of evolution. His proposition that all spe ...

Charles Darwin
called it an "abominable mystery". However, the fossil record has considerably grown since the time of Darwin, and recently discovered angiosperm fossils such as '''', 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 fern A seed is an Plant embryogenesis, embryonic plant enclosed in a testa (botany), protective outer covering. The formation of the seed is part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosp ...
s, have been proposed as the
ancestors An ancestor, also known as a forefather, fore-elder or a forebear, is a parent A parent is a caregiver of the offspring in their own species. In humans, a parent is the caretaker of a child (where "child" refers to offspring, not necessarily ...
of flowering plants, but there is no continuous fossil evidence showing how flowers evolved, and botanists still regard it as a mystery. Some older fossils, such as the upper
Triassic The Triassic ( ) is a geologic period A geological period is one of the several subdivisions of geologic time enabling cross-referencing of rocks and geologic events from place to place. These periods form elements of a hierarchy of divisions int ...

Triassic
'' Sanmiguelia lewisi'', have been suggested. The first seed bearing plants, like the
ginkgo ''Ginkgo'' is a genus Genus /ˈdʒiː.nəs/ (plural genera /ˈdʒen.ər.ə/) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. ...

ginkgo
, and
conifer Conifers are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single ex ...
s (such as
pine A pine is any Pinophyta, conifer in the genus ''Pinus'' () of the family (biology), family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The Plant List compiled by the Royal Botanic Gardens, Kew and Missouri Botanical Garden a ...

pine
s and
fir Firs (''Abies'') are a genus Genus (plural genera) is a taxonomic rank Taxonomy (general) is the practice and science of classification of things or concepts, including the principles that underlie such classification. The term may also re ...

fir
s), 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 Oleanane is a natural triterpenoid Triterpenes are a class of chemical compounds composed of three terpene units with the molecular formula C30H48; they may also be thought of as consisting of six isoprene units. Animals, plants and fungi all pro ...

Oleanane
, a
secondary metabolite Secondary is an adjective meaning "second" or "second hand". It may refer to: * Secondary (chemistry), term used in organic chemistry to classify various types of compounds * The group of (usually at least four) defensive backs in gridiron football ...
produced by many flowering plants, has been found in
Permian The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous period million years ago (Mya), to the beginning of the Triassic period 251.902 Mya. It is the last period of the Paleo ...
deposits of that age together with fossils of gigantopterids. Gigantopterids are a group of extinct seed plants that share many morphological traits with flowering plants, although they are not known to have been flowering plants themselves.


Triassic and Jurassic

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. A close relationship between angiosperms and gnetophytes, proposed on the basis of Morphology (biology), morphological evidence, has more recently been disputed on the basis of molecular biology, molecular evidence that suggest gnetophytes are instead more closely related to other gymnosperms. The fossil plant species Nanjinganthus, ''Nanjinganthus dendrostyla'' from Early Jurassic China seems to share many exclusively angiosperm features, such as a thickened Receptacle (botany), receptacle with
ovule In seed plant The spermatophytes, also known as phanerogams (taxon Phanerogamae) or phaenogams (taxon Phaenogamae), comprise those plants that produce seeds, hence the alternative name seed plants. They are a subset of the embryophytes or l ...

ovule
s, and thus might represent a Crown group, crown-group or a stem-group angiosperm. However, these have been disputed by other researchers, who contend that the structures are misinterpreted decomposed conifer cones. The evolution of seed plants and later angiosperms appears to be the result of two distinct rounds of Paleopolyploidy, whole genome duplication events. These occurred at and . Another possible whole genome duplication event at perhaps created the ancestral line that led to all modern flowering plants. That event was studied by sequencing the genome of an ancient flowering plant, ''Amborella trichopoda'', and directly addresses Darwin's "abominable mystery". 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.


Cretaceous

Whereas the earth had previously been dominated by ferns and conifers, angiosperms appeared and quickly spread during the Cretaceous. They now comprise about 90% of all plant species including most food crops. 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. The oldest known fossils definitively attributable to angiosperms are reticulated monosulcate pollen from the late Valanginian (Early or Lower Cretaceous - 140 to 133 million years ago) of Italy and Israel, likely representative of the Basal angiosperms, basal angiosperm grade. The earliest known macrofossil confidently identified as an angiosperm, ''Archaefructus liaoningensis'', is dated to about 125 million years BP (the Cretaceous period), whereas pollen considered to be of angiosperm origin takes the fossil record back to about 130 million years BP, with ''Montsechia'' representing the earliest flower at that time. 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. 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 Flower#Evolution, 140 million years ago. A Bayesian analysis of 52 angiosperm taxa suggested that the crown group of angiosperms evolved between and . Recent DNA analysis based on molecular systematics showed that Amborella, ''Amborella trichopoda'', found on the Pacific island of
New Caledonia ) , anthem = " Soyons unis, devenons frères" , image_map = New Caledonia on the globe (small islands magnified) (Polynesia centered).svg , map_alt = Location of New Caledonia , map_caption = Location of New Caledonia , mapsize = 290px , s ...

New Caledonia
, belongs to a sister group of the other flowering plants, and morphological studies suggest that it has features that may have been characteristic of the earliest flowering plants. The orders
Amborellales ''Amborella'' is a monotypic genus Genus (plural genera) is a taxonomic rank Taxonomy (general) is the practice and science of classification of things or concepts, including the principles that underlie such classification. The term may als ...
,
Nymphaeales The Nymphaeales are an order of flowering plant The flowering plants, also known as Angiospermae (), or Magnoliophyta (), are the most diverse group of Embryophyte, land plants, with 64 Order(biology), orders, 416 Family (biology), families, a ...
, and
Austrobaileyales Austrobaileyales is an order of flowering plants consisting of about 100 species of woody plants growing as trees, shrubs and lianas. Perhaps the most familiar species is '' Illicium verum'', from which comes the spice star anise. The order be ...

Austrobaileyales
diverged as separate lineages from the remaining angiosperm clade at a very early stage in flowering plant evolution. The great angiosperm Adaptive radiation, 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 of the eight main groups occurred around 140 million years ago. (the genus ''
Ceratophyllum ''Ceratophyllum'' is a cosmopolitan genus Genus (plural genera) is a taxonomic rank Taxonomy (general) is the practice and science of classification of things or concepts, including the principles that underlie such classification. The term ...
'', the family
Chloranthaceae Chloranthaceae is a family In human society, family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of families is to maintain t ...
, the
eudicots The eudicots, Eudicotidae or eudicotyledons are a clade A clade (), also known as a monophyletic group or natural group, is a group of organism In biology, an organism () is any organic, life, living system that functions as an ind ...
, the
magnoliids Magnoliids (or Magnoliidae or Magnolianae) are a group of flowering plant The flowering plants, also known as Angiospermae (), or Magnoliophyta (), are the most diverse group of Embryophyte, land plants, with 64 Order(biology), orders, 416 Famil ...
, and the Monocotyledon, monocots) . It is generally assumed that the function (biology), 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 specialised 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 (biology), specialisation in both the plant(s) and their partners. Note that the wasp example is not incidental; bees, which, it is postulated, evolved specifically due to mutualistic plant relationships, are descended from wasps. Animals are also involved in the distribution of seeds. Fruit, which is formed by the enlargement of flower parts, is frequently a seed-dispersal tool that attracts animals to eat or otherwise disturb it, incidentally scattering the seeds it contains (see frugivory). Although many such Mutualism (biology), mutualistic relationships remain too fragile to survive competition (biology), competition and to spread widely, flowering proved to be an unusually effective means of reproduction, spreading (whatever its origin) to become the dominant form of land plant life. Flower ontogeny uses a combination of genes normally responsible for forming new shoots. The most primitive flowers probably had a variable number of flower parts, often separate from (but in contact with) each other. The flowers tended to grow in a spiral pattern, to be bisexual (in plants, this means both male and female parts on the same flower), and to be dominated by the ovary (plants), ovary (female part). As flowers evolved, some variations developed parts fused together, with a much more specific number and design, and with either specific sexes per flower or plant or at least "ovary-inferior". Flower evolution continues to the present day; modern flowers have been so profoundly influenced by humans that some of them cannot be pollinated in nature. Many modern domesticated flower species were formerly simple weeds, which sprouted only when the ground was disturbed. Some of them tended to grow with human crops, perhaps already having symbiotic companion plant relationships with them, and the prettiest did not get plucked because of their beauty, developing a dependence upon and special adaptation to human affection. A few Paleontology, 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 Herbivore, 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. By the late Cretaceous, angiosperms appear to have dominated environments formerly occupied by ferns and cycadophytes, but large canopy-forming trees replaced
conifer Conifers are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single ex ...
s as the dominant trees only close to the end of the Cretaceous 66 million years ago or even later, at the beginning of the Paleogene. The radiation of herbaceous angiosperms occurred much later. Yet, many fossil plants recognisable as belonging to modern families (including beech,
oak An oak is a tree or shrub A shrub (or bush, but this is more of a gardening term) is a small- to medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be deciduous o ...

oak
, maple, and
magnolia ''Magnolia'' is a large genus of about 210The number of species in the genus ''Magnolia'' depends on the taxonomic view that one takes up. Recent molecular and morphology (biology), morphological research shows that former genera ''Talauma'', ''D ...

magnolia
) had already appeared by the late Cretaceous. Flowering plants appeared in Australia about 126 million years ago. This also pushed the age of South Polar region of the Cretaceous, ancient Australian vertebrates, in what was then a Polar forests of the Cretaceous, south polar continent, to 126-110 million years old.


Gallery of photos

Asteracea poster 3.jpg, A poster of twelve different species of flowers of the family Asteraceae Lupinus-pilosus-2015-Zachi-Evenor-cropped01.jpg, ''Lupinus pilosus'' Rose bud.jpg, Bud of a pink rose


Diversity

The number of species of flowering plants is estimated to be in the range of 250,000 to 400,000. This compares to around 12,000 species of moss and 11,000 species of Pteridophyta, pteridophytes, showing that flowering plants are much more diverse. The number of family (biology), families in APG system, APG (1998) was 462. In APG II (2003) it is not settled; at maximum it is 457, but within this number there are 55 optional segregates, so that the minimum number of families in this system is 402. In APG III (2009) there are 415 families. Compared to the APG III system, the APG IV system recognizes five new orders (Boraginales, Dilleniales, Icacinales, Metteniusales and Vahliales), along with some new families, making a total of 64 angiosperm orders and 416 families. The diversity of flowering plants is not evenly distributed. Nearly all species belong to the eudicot (75%), monocot (23%), and magnoliid (2%) clades. The remaining five clades contain a little over 250 species in total; i.e. less than 0.1% of flowering plant diversity, divided among nine families. The 43 most diverse of 443 families of flowering plants by species, in their APG circumscriptions, are # Asteraceae or Compositae (Bellis perennis, daisy family): 22,750 species; # Orchidaceae (orchid family): 21,950; # Fabaceae or Leguminosae (bean family): 19,400; # Rubiaceae (Rubia, madder family): 13,150; # Poaceae or Gramineae (grass family): 10,035; # Lamiaceae or Labiatae (mentha, mint family): 7,175; # Euphorbiaceae (spurge family): 5,735; # Melastomataceae or Melastomaceae (Melastoma, melastome family): 5,005; # Myrtaceae (Myrtus, myrtle family): 4,625; # Apocynaceae (dogbane family): 4,555; # Cyperaceae (sedge family): 4,350; # Malvaceae (Malva, mallow family): 4,225; # Araceae (arum family): 4,025; # Ericaceae (Erica (plant), heath family): 3,995; # Gesneriaceae (gesneriad family): 3,870; # Apiaceae or Umbelliferae (parsley family): 3,780; # Brassicaceae or Cruciferae (cabbage family): 3,710: # Piperaceae (Piper (genus), pepper family): 3,600; # Bromeliaceae (bromeliad family): 3,540; # Acanthaceae (Acanthus (plant), acanthus family): 3,500; # Rosaceae (rose family): 2,830; # Boraginaceae (borage family): 2,740; # Urticaceae (nettle family): 2,625; # Ranunculaceae (
buttercup ''Ranunculus'' is a large genus of about 600 species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined a ...

buttercup
family): 2,525; # Lauraceae (Laurus, laurel family): 2,500; # Solanaceae (nightshade family): 2,460; # Campanulaceae (Campanula, bellflower family): 2,380; # Arecaceae (palm (plant), palm family): 2,361; # Annonaceae (custard apple family): 2,220; # Caryophyllaceae (dianthus, pink family): 2,200; # Orobanchaceae (broomrape family): 2,060; # Amaranthaceae (amaranth family): 2,050; # Iridaceae (Iris (plant), iris family): 2,025; # Aizoaceae or Ficoidaceae (Aizoaceae, ice plant family): 2,020; # Rutaceae (rue family): 1,815; # Phyllanthaceae (phyllanthus family): 1,745; # Scrophulariaceae (figwort family): 1,700; # Gentianaceae (gentian family): 1,650; # Convolvulaceae (Convolvulus, bindweed family): 1,600; # Proteaceae (protea family): 1,600; # Sapindaceae (Sapindus, soapberry family): 1,580; # Cactaceae (Mammillaria, cactus family): 1,500; # 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


Fertilisation and embryogenesis

Double fertilization refers to a process in which two sperm cells fertilise 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 fertilise the egg cell, producing a diploid (2''n'') zygote. The second sperm cell fuses with both central cell nuclei, producing a triploid (3''n'') 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 the embryo and endosperm proceeds within the embryo sac, the sac wall enlarges and combines with the nucellus (which is likewise enlarging) and the integument to form the ''seed coat''. The ovary wall develops to form the fruit or pericarp, whose form is closely associated with type of seed dispersal system. Frequently, the influence of fertilisation 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 An apple is an edible fruit produced by an apple tree (''Malus domestica''). Apple fruit tree, trees are agriculture, cultivated worldwide and are the most widely grown species in the genus ''Malus''. The tree originated in Central Asia, wher ...

apple
, Fragaria, 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 Dehiscence (botany), dehiscent and the seed is exposed, in general, the seed-coat is well developed and must discharge the functions otherwise executed by the fruit. In some cases, like in the Asteraceae family, species have evolved to exhibit heterocarpy, or the production of different fruit morphs.Gardocki, M. E., Zablocki, H., El-Keblawy, A., & Freeman, D. C. (2000). Heterocarpy in Calendula micrantha (Asteraceae): The effects of competition and availability of water on the performance of offspring from different fruit morphs. Evolutionary Ecology Research. 2(6):701-718 These fruit morphs, produced from one plant, are different in size and shape, which influence dispersal range and germination rate. These fruit morphs are adapted to different environments, increasing chances for survival.


Meiosis

Like all diploid multicellular organisms that use sexual reproduction, flowering plants generate gametes using a specialised type of cell division called meiosis. Meiosis takes place in the
ovule In seed plant The spermatophytes, also known as phanerogams (taxon Phanerogamae) or phaenogams (taxon Phaenogamae), comprise those plants that produce seeds, hence the alternative name seed plants. They are a subset of the embryophytes or l ...

ovule
—a structure within the ovary that is located within the pistil at the centre 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. It is thought that the basal chromosome number in angiosperms is n = 7. 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 produce three antipodal cells, two 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 (microsporangia, microsporangium). During meiosis, a diploid microspore mother cell undergoes two successive meiotic divisions to produce four 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 fertilisation, 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 DNA damage (naturally occurring), damages in the DNA to be passed on to progeny. To explain the adaptive function of meiosis in flowering plants, some authors emphasise diversity and others emphasise DNA repair.


Apomixis

Apomixis (reproduction via asexually formed seeds) is found naturally in about 2.2% of angiosperm genera. One type of apomixis, gametophytic apomixis found in a dandelion species involves formation of an unreduced embryo sac due to incomplete meiosis (apomeiosis) and development of an embryo from the unreduced egg inside the embryo sac, without fertilisation (parthenogenesis). Some angiosperms, including many citrus varieties, are able to produce fruits through a type of apomixis called nucellar embryony.


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 capsicum, 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 Orange (fruit), oranges, lemons, grapefruits, etc.), and the Rosaceae, or rose family (including
apple An apple is an edible fruit produced by an apple tree (''Malus domestica''). Apple fruit tree, trees are agriculture, cultivated worldwide and are the most widely grown species in the genus ''Malus''. The tree originated in Central Asia, wher ...

apple
s, pears, cherry, 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 (''coconut, Cocos nucifera'') on Pacific atolls, and the olive (''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. Coffee and Hot chocolate, cocoa are the common beverages obtained from the flowering plants. 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.


See also

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


Notes


References


Bibliography


Articles, books and chapters

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1st edition published by Oxford University Press in 1991
* * * Cromie, William J. (December 16, 1999)

Harvard University Gazette. * * * * * * * * * * * * * * * * *


Websites

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External links

* * * {{DEFAULTSORT:Flowering Plant Angiosperms, Plant sexuality Plants Pollination