Flowering plant

Angiosperms are terrestrial vascular plants; like the gymnosperms, they have roots, stems, leaves, and seeds. They differ from other seed plants in several ways.

{{further|Plant ecology}}

File:MountainAshWithCars.jpg|Eucalyptus regnans,
a tree almost 100 m tall

File:WolffiaArrhiza2.jpg|Wolffia arrhiza, a rootless floating freshwater plant under 2 mm across

The largest angiosperms are Eucalyptus gum trees of Australia, and Shorea faguetiana, dipterocarp rainforest trees of Southeast Asia, both of which can reach almost {{convert|100|m|ft}} in height.{{cite web |title=Menara, yellow meranti, Shorea |url=https://www.guinnessworldrecords.com/world-records/572236-tallest-flowering-plant-angiosperm |website=Guinness World Records |date=6 January 2019 |access-date=8 May 2023 |quote=yellow meranti (Shorea faguetiana) ... 98.53 m (323 ft 3.1 in) tall ... swamp gum (Eucalyptus regnans) ... In 2014, it had a tape-drop height of 99.82 m (327 ft 5.9 in)}} The smallest are Wolffia duckweeds which float on freshwater, each plant less than {{convert|2|mm|in|2}} across.{{Cite web |date=2009-11-25 |title=The Charms of Duckweed |url=http://www.mobot.org/jwcross/duckweed/duckweed.htm |access-date=2022-07-05 |publisher=Missouri Botanical Garden |archive-url=https://web.archive.org/web/20091125213317/http://www.mobot.org/jwcross/duckweed/duckweed.htm |url-status=dead |archive-date=25 November 2009 }}

File:Sunlight on a gunnera leaf, 'Quarry Garden', Belsay estate - geograph.org.uk - 1384733.jpg|Gunnera captures sunlight for photosynthesis over the large surfaces of its leaves, which are supported by strong veins.

File:Orobanche purpurea.jpg|Orobanche purpurea, a parasitic broomrape with no leaves, obtains all its food from other plants.

Considering their method of obtaining energy, some 99% of flowering plants are photosynthetic autotrophs, deriving their energy from sunlight and using it to create molecules such as sugars. The remainder are parasitic, whether on fungi like the orchids for part or all of their life-cycle,{{cite journal |last=Leake |first=J.R. |year=1994 |title=The biology of myco-heterotrophic ('saprophytic') plants |journal=New Phytologist |volume=127 |issue=2 |pages=171–216 |doi=10.1111/j.1469-8137.1994.tb04272.x |pmid=33874520 |bibcode=1994NewPh.127..171L |s2cid=85142620 }} or on other plants, either wholly like the broomrapes, Orobanche, or partially like the witchweeds, Striga.{{cite journal |last1=Westwood |first1=James H. |last2=Yoder |first2=John I. |last3=Timko |first3=Michael P. |last4=dePamphilis |first4=Claude W. |title=The evolution of parasitism in plants |journal=Trends in Plant Science |volume=15 |issue=4 |year=2010 |issn=1360-1385 |doi=10.1016/j.tplants.2010.01.004 |pages=227–235|pmid=20153240 |bibcode=2010TPS....15..227W }}

File:Carnegiea gigantea Saguaro cactus plant (cropped).jpg|Carnegiea gigantea, the saguaro cactus, grows in hot dry deserts in Mexico and the southern United States.

File:Dryas octopetala (Colorado, USA).jpg|Dryas octopetala, the mountain avens, lives in cold arctic and montane habitats in the far north of America and Eurasia.

File:Nelumbo nucifera Gaertn. (47502598342).jpg|Nelumbo nucifera, the sacred lotus, grows in warm freshwater across tropical and subtropical Asia.

File:Zostera.jpg|Zostera seagrass grows on the seabed in sheltered coastal waters.

In terms of their environment, flowering plants are cosmopolitan, occupying a wide range of habitats on land, in fresh water and in the sea. On land, they are the dominant plant group in every habitat except for frigid moss-lichen tundra and coniferous forest.{{cite web |title=Angiosperms |url=http://landau.faculty.unlv.edu//angiosperms.htm |publisher=University of Nevada, Las Vegas |access-date=6 May 2023}} The seagrasses in the Alismatales grow in marine environments, spreading with rhizomes that grow through the mud in sheltered coastal waters.{{cite book |last1=Kendrick |first1=Gary A. |last2=Orth |first2=Robert J. |last3=Sinclair |first3=Elizabeth A. |last4=Statton |first4=John |title=Plant Regeneration from Seeds |chapter=Effect of climate change on regeneration of seagrasses from seeds |year=2022 |doi=10.1016/b978-0-12-823731-1.00011-1 |pages=275–283 |isbn=978-0-1282-3731-1 }}

File:Drosera anglica ne2.jpg|Drosera anglica, a sundew, lives in nutrient-poor acid bogs, deriving nutrients from trapped insects.

File:Gentiana verna.jpg|Gentiana verna, the spring gentian, flourishes in dry limestone habitats.

Some specialised angiosperms are able to flourish in extremely acid or alkaline habitats. The sundews, many of which live in nutrient-poor acid bogs, are carnivorous plants, able to derive nutrients such as nitrate from the bodies of trapped insects.{{cite journal |last1=Karlsson |first1=P. S. |last2=Pate |first2=J. S. |title=Contrasting effects of supplementary feeding of insects or mineral nutrients on the growth and nitrogen and phosphorous economy of pygmy species of Drosera. |journal=Oecologia |date=1992 |volume=92 |issue=1 |pages=8–13 |doi=10.1007/BF00317256 |pmid=28311806 |bibcode=1992Oecol..92....8K |s2cid=13038192 }} Other flowers such as Gentiana verna, the spring gentian, are adapted to the alkaline conditions found on calcium-rich chalk and limestone, which give rise to often dry topographies such as limestone pavement.{{cite book |last=Pardoe |first=H. S. |title=Mountain Plants of the British Isles |year=1995 |publisher=National Museum of Wales |page=24 |isbn=978-0-7200-0423-6 |url=https://books.google.com/books?id=bRHwm1F15S0C&pg=PA24}}

File:GT Herb Robert.jpg|Geranium robertianum, herb-Robert, is an annual or biennial herb of Europe and North America.

File:Betula_pendula_001.jpg|Betula pendula, the silver birch, is a perennial deciduous tree of Eurasia.

File:Lianas.jpg|Lianas Austrosteenisia, Parsonsia, and Sarcopetalum climbing trees in Australia

As for their growth habit, the flowering plants range from small, soft herbaceous plants, often living as annuals or biennials that set seed and die after one growing season,{{cite journal |last1=Hart |first1=Robin |title=Why are Biennials so Few? |journal=The American Naturalist |date=1977 |volume=111 |issue=980 |pages=792–799 |doi=10.1086/283209 |jstor=2460334 |bibcode=1977ANat..111..792H |s2cid=85343835 }} to large perennial woody trees that may live for many centuries and grow to many metres in height. Some species grow tall without being self-supporting like trees by climbing on other plants in the manner of vines or lianas.{{cite journal |last1=Rowe |first1=Nick |last2=Speck |first2=Thomas |title=Plant growth forms: an ecological and evolutionary perspective |journal=New Phytologist |volume=166 |issue=1 |date=2005-01-12 |issn=0028-646X |doi=10.1111/j.1469-8137.2004.01309.x |pages=61–72 |pmid=15760351 |doi-access=free |bibcode=2005NewPh.166...61R }}

The number of species of flowering plants is estimated to be in the range of 250,000 to 400,000.{{cite journal |last=Thorne |first=R.F. |title=How many species of seed plants are there?|journal=Taxon|volume=51 |pages=511–522 |year=2002|doi=10.2307/1554864 |issue=3 |jstor=1554864|bibcode=2002Taxon..51..511T }}{{cite journal |last1=Scotland |first1=R. W. |last2=Wortley |first2=A. H. |title=How many species of seed plants are there? |journal=Taxon |volume=52 |pages=101–104|year=2003 |doi=10.2307/3647306 |issue=1 |jstor=3647306|bibcode=2003Taxon..52..101S }}{{cite journal |last=Govaerts |first=R. |title=How many species of seed plants are there? – a response |journal=Taxon |volume=52 |issue=3 |pages=583–584 |year=2003 |doi=10.2307/3647457 |jstor=3647457|doi-access=free |bibcode=2003Taxon..52..583G }} This compares to around 12,000 species of moss{{cite journal |last1=Goffinet |first1=Bernard |last2=Buck |first2=William R. |year=2004 |title=Systematics of the Bryophyta (Mosses): From molecules to a revised classification |journal=Monographs in Systematic Botany |volume=98 |pages=205–239 }} and 11,000 species of pteridophytes.{{cite book |last1=Raven |first1=Peter H. |last2=Evert |first2=Ray F. |last3=Eichhorn |first3=Susan E. |date=2005 |title=Biology of Plants |edition=7th |location=New York |publisher=W. H. Freeman and Company |isbn=0-7167-1007-2 }} The APG system seeks to determine the number of families, mostly by molecular phylogenetics. In the 2009 APG III there were 415 families.{{sfn|APG|2009}} The 2016 APG IV added five new orders (Boraginales, Dilleniales, Icacinales, Metteniusales and Vahliales), along with some new families, for a total of 64 angiosperm orders and 416 families.{{sfn|APG|2016}}

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 25 most species-rich of 443 families,{{cite web |last=Stevens |first=P. F. |year=2011 |url=https://www.mobot.org/MOBOT/Research/APweb/welcome.html |title=Angiosperm Phylogeny Website (at Missouri Botanical Garden) |access-date=21 February 2022 |archive-date=20 January 2022 |archive-url=https://web.archive.org/web/20220120065914/http://www.mobot.org/mobot/research/apweb/welcome.html |url-status=live}} containing over 166,000 species between them in their APG circumscriptions, are:

{{main|Plant taxonomy}}

File:Ehret-Methodus Plantarum Sexualis.jpg

The botanical term "angiosperm", from Greek words {{lang|grc-Latn|angeíon}} ({{wikt-lang|grc|ἀγγεῖον}} 'bottle, vessel') and {{lang|grc-Latn|spérma}} ({{wikt-lang|grc|σπέρμα}} 'seed'), was coined in the form "Angiospermae" by Paul Hermann in 1690, including only flowering plants whose seeds were enclosed in capsules.{{sfn|Balfour|Rendle|1911|p=9}} The term angiosperm fundamentally changed in meaning in 1827 with Robert Brown, when angiosperm came to mean a seed plant with enclosed ovules.{{cite book |last=Brown |first=Robert |chapter=Character and description of Kingia, a new genus of plants found on the southwest coast of New Holland: with observations on the structure of its unimpregnated ovulum; and on the female flower of Cycadeae and Coniferae |pages=534–565 |chapter-url={{Google books|RjdCAAAAIAAJ|page=534|plainurl=yes}} |editor=King, Philip Parker |title=Narrative of a Survey of the Intertropical and Western Coasts of Australia: Performed Between the Years 1818 and 1822 |date=1827 |publisher=J. Murray |oclc=185517977 }}{{Cite journal |last=Buggs |first=Richard J.A. |date=January 2021 |title=The origin of Darwin's "abominable mystery" |journal=American Journal of Botany |volume=108 |issue=1 |pages=22–36 |doi=10.1002/ajb2.1592 |pmid=33482683 |s2cid=231689158 |doi-access=free }} In 1851, with Wilhelm Hofmeister's work on embryo-sacs, Angiosperm came to have its modern meaning of all the flowering plants including Dicotyledons and Monocotyledons.{{sfn|Balfour|Rendle|1911|p=10}} The APG system{{sfn|APG|2009}} treats 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 the subclass Magnoliidae.{{sfn|Chase|Reveal|2009}} From 1998, the Angiosperm Phylogeny Group (APG) has reclassified the angiosperms, with updates in the APG II system in 2003,{{sfn|APG|2003}} the APG III system in 2009,{{sfn|APG|2009}}{{cite press release |title=As easy as APG III – Scientists revise the system of classifying flowering plants |publisher=The Linnean Society of London |url=https://www.linnean.org/index.php?id=448 |access-date=2 October 2009 |date=8 October 2009 |archive-url=https://web.archive.org/web/20101126092206/https://www.linnean.org/index.php?id=448 |archive-date=26 November 2010}} and the APG IV system in 2016.{{sfn|APG|2016}}

In 2019, a molecular phylogeny of plants placed the flowering plants in their evolutionary context:{{cite journal |last1=Leebens-Mack |first1=M. |last2=Barker |first2=M. |last3=Carpenter |first3=E. |author4-link=Michael Deyholos |last4=Deyholos |first4=M. K. |last5=Gitzendammer |first5=M. A. |last6=Graham |first6=S.W. |last7=Grosse |first7=I. |last8=Li |first8=Zheng |display-authors=3 |title=One thousand plant transcriptomes and the phylogenomics of green plants |journal=Nature |volume=574 |issue=7780 |year=2019 |pages=679–685 |doi=10.1038/s41586-019-1693-2 |pmid=31645766 |pmc=6872490 |doi-access=free }}

{{clade

|label1= Embryophytes

|sublabel1= land plants

|1={{clade

|1={{clade

|1= Bryophytes 20px

|label2= Tracheophytes

|sublabel2= vascular plants

|2={{clade

|1=Lycophytes 50px

|2={{clade

|1=Ferns 40px

|label2= Spermatophytes

|sublabel2= seed plants

|2={{clade

|label1= Gymnosperms

|sublabel1= conifers and allies

|1= 50px

|label2= Angiosperms

|sublabel2= flowering plants

|2= 50px

}}

}}

}}

}}

}}

}}

The main groups of living angiosperms are:{{cite journal |last=Guo |first=Xing |title=Chloranthus genome provides insights into the early diversification of angiosperms |journal=Nature Communications |date=26 November 2021 |volume=12 |issue=1 |page=6930 |doi=10.1038/s41467-021-26922-4 |pmid=34836973 |pmc=8626473 |bibcode=2021NatCo..12.6930G |doi-access=free }}{{sfn|APG|2016}}

{{barlabel |size=20

|at1=4 |label1=Basal angiosperms

|at2=14 |label2=Core angiosperms

|cladogram={{clade|style=font-size:100%;line-height:100%

|label1= Angiosperms

|1={{clade

|1=Amborellales 70px 1 sp. New Caledonia shrub |barbegin1=red

|2={{clade

|1=Nymphaeales 70px c. 80 spp.{{cite journal |last1=Palmer |first1=Jeffrey D. |last2=Soltis |first2=Douglas E. |last3=Chase |first3=Mark W. |title=The plant tree of life: an overview and some points of view |journal=American Journal of Botany |volume=91 |issue=10 |pages=1437–45 |date=October 2004 |pmid=21652302 |doi=10.3732/ajb.91.10.1437 |doi-access=free }}, [https://www.amjbot.org/cgi/content/full/91/10/1437/F2 Figure 2] {{Webarchive|url=https://web.archive.org/web/20110202073937/http://www.amjbot.org/cgi/content/full/91/10/1437/F2 |date=2 February 2011 }} water lilies & allies |bar1=red

|2={{clade

|1=Austrobaileyales 70px c. 100 spp. woody plants |barend1=red

|2={{clade

|1={{clade

|1=Magnoliids 70px c. 10,000 spp. 3-part flowers, 1-pore pollen, usu. branch-veined leaves |barbegin1=green

|2=Chloranthales 70px 77 spp.{{cite book |last1=Christenhusz |first1=Maarten J. M. |last2=Fay |first2=Michael F. |last3=Chase |first3=Mark W. |url=https://books.google.com/books?id=LLo7DwAAQBAJ&pg=PA114 |title=Plants of the World: An Illustrated Encyclopedia of Vascular Plants |isbn=978-0-226-52292-0 |page=114 |date=2017 |publisher=University of Chicago Press }} Woody, apetalous |bar2=green

}}

|2={{clade

|1=Monocots 45px c. 70,000 spp.{{Cite journal |last1=Massoni |first1=Julien |last2=Couvreur |first2=Thomas L.P. |last3=Sauquet |first3=Hervé |date=2015-03-18 |title=Five major shifts of diversification through the long evolutionary history of Magnoliidae (angiosperms) |journal=BMC Evolutionary Biology |volume=15 |issue=1 |pages=49 |doi=10.1186/s12862-015-0320-6 |pmc=4377182 |pmid=25887386 |bibcode=2015BMCEE..15...49M |doi-access=free }} 3-part flowers, 1 cotyledon, 1-pore pollen, usu. parallel-veined leaves   |bar1=green

|2={{clade

|1=Ceratophyllales 70px c. 6 spp. aquatic plants |bar1=green

|2=Eudicots 70px c. 175,000 spp. 4- or 5-part flowers, 3-pore pollen, usu. branch-veined leaves |barend2=green

}}

}}

}}

}}

}}

}}

}}

}}

In 2024, Alexandre R. Zuntini and colleagues constructed a tree of some 6,000 flowering plant genera, representing some 60% of the existing genera, on the basis of analysis of 353 nuclear genes in each specimen. Much of the existing phylogeny is confirmed; the rosid phylogeny is revised.{{cite journal |last1=Zuntini |first1=Alexandre R. |last2=Carruthers |first2=Tom |last3=Maurin |first3=Olivier |last4=Bailey |first4=Paul C. |last5=Leempoel |first5=Kevin |last6=Brewer |first6=Grace E. |display-authors=etal |title=Phylogenomics and the rise of the angiosperms |journal=Nature |date=24 April 2024 |volume=629 |issue=8013 |pages=843–850 |issn=0028-0836 |doi=10.1038/s41586-024-07324-0|pmid=38658746 |pmc=11111409 |bibcode=2024Natur.629..843Z }}

File:Tree of Angiosperm Phylogeny 2024.jpg

{{main|Fossil history of flowering plants}}

File:Sagaria cilentana (cropped).jpg in the Cretaceous created many flowering plants, such as Sagaria in the Ranunculaceae.]]

Fossilised spores suggest that land plants (embryophytes) have existed for at least 475 million years.{{cite journal |last=Edwards |first=D. |title=The role of mid-palaeozoic mesofossils in the detection of early bryophytes |journal=Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences |volume=355 |issue=1398 |pages=733–54; discussion 754–5 |date=June 2000 |pmid=10905607 |pmc=1692787 |doi=10.1098/rstb.2000.0613 }} However, angiosperms appear suddenly and in great diversity in the fossil record in the Early Cretaceous (~130 mya).{{Cite journal |last1=Herendeen |first1=Patrick S. |last2=Friis |first2=Else Marie |last3=Pedersen |first3=Kaj Raunsgaard |last4=Crane |first4=Peter R. |date=2017-03-03 |title=Palaeobotanical redux: revisiting the age of the angiosperms |url=https://rdcu.be/c0Zhm |journal=Nature Plants |volume=3 |issue=3 |pages=17015 |doi=10.1038/nplants.2017.15 |pmid=28260783 |bibcode=2017NatPl...317015H |s2cid=205458714 |issn=2055-0278}}{{Cite journal |last=Friedman |first=William E. |date=January 2009 |title=The meaning of Darwin's "abominable mystery" |url=https://onlinelibrary.wiley.com/doi/10.3732/ajb.0800150 |journal=American Journal of Botany |volume=96 |issue=1 |pages=5–21 |doi=10.3732/ajb.0800150 |pmid=21628174}} Claimed records of flowering plants prior to this are not widely accepted.{{Cite journal |last=Bateman |first=Richard M |date=2020-01-01 |editor-last=Ort |editor-first=Donald |title=Hunting the Snark: the flawed search for mythical Jurassic angiosperms |url=https://academic.oup.com/jxb/article/71/1/22/5571867 |journal=Journal of Experimental Botany |language=en |volume=71 |issue=1 |pages=22–35 |doi=10.1093/jxb/erz411 |pmid=31538196 |issn=0022-0957}} Molecular evidence suggests that the ancestors of angiosperms diverged from the gymnosperms during the late Devonian, about 365 million years ago.{{Cite journal |last1=Stull |first1=Gregory W. |last2=Qu |first2=Xiao-Jian |last3=Parins-Fukuchi |first3=Caroline |last4=Yang |first4=Ying-Ying |last5=Yang |first5=Jun-Bo |last6=Yang |first6=Zhi-Yun |last7=Hu |first7=Yi |last8=Ma |first8=Hong |last9=Soltis |first9=Pamela S. |last10=Soltis |first10=Douglas E. |last11=Li |first11=De-Zhu |display-authors=3 |date=19 July 2021 |title=Gene duplications and phylogenomic conflict underlie major pulses of phenotypic evolution in gymnosperms |url=https://www.nature.com/articles/s41477-021-00964-4 |journal=Nature Plants |volume=7|issue=8 |pages=1015–1025 |doi=10.1038/s41477-021-00964-4 |pmid=34282286 |bibcode=2021NatPl...7.1015S |s2cid=236141481 |access-date=10 January 2022 |archive-date=10 January 2022 |archive-url=https://web.archive.org/web/20220110174725/https://www.nature.com/articles/s41477-021-00964-4/ |url-status=live}} The origin time of the crown group of flowering plants remains contentious.{{Cite journal |last1=Sauquet |first1=Hervé |last2=Ramírez-Barahona |first2=Santiago |last3=Magallón |first3=Susana |date=2022-06-24 |editor-last=Melzer |editor-first=Rainer |title=What is the age of flowering plants? |url=https://academic.oup.com/jxb/article/73/12/3840/6570702 |journal=Journal of Experimental Botany |language=en |volume=73 |issue=12 |pages=3840–3853 |doi=10.1093/jxb/erac130 |pmid=35438718 |issn=0022-0957}} By the Late Cretaceous, angiosperms appear to have dominated environments formerly occupied by ferns and gymnosperms. Large canopy-forming trees replaced conifers as the dominant trees close to the end of the Cretaceous, 66 million years ago.{{cite book |last1=Sadava |first1=David |last2=Heller |first2=H. Craig |last3=Orians |first3=Gordon H. |last4=Purves |first4=William K. |last5=Hillis |first5=David M. |display-authors=3 |title=Life: the science of biology |url=https://books.google.com/books?id=1m0_FLEjd-cC&pg=PA477 |access-date=4 August 2010 |date=December 2006 |publisher=Macmillan |isbn=978-0-7167-7674-1 |pages=477– |archive-date=23 December 2011 |archive-url=https://web.archive.org/web/20111223082952/http://books.google.com/books?id=1m0_FLEjd-cC&pg=PA477|url-status=live}} The radiation of herbaceous angiosperms occurred much later.{{cite book |last1=Stewart |first1=Wilson Nichols |last2=Rothwell |first2=Gar W. |title=Paleobotany and the evolution of plants |edition=2nd |publisher=Cambridge University Press |year=1993 |page=498 |isbn=978-0-521-23315-6 }}

{{main|Flower|Plant reproductive morphology}}

File:Angiosperm life cycle diagram-en.svg showing reproductive parts and life cycle]]

The characteristic feature of angiosperms is the flower. Its function is to ensure fertilization of the ovule and development of fruit containing seeds.{{Cite journal |last=Willson |first=Mary F. |date=1 June 1979 |title=Sexual Selection in Plants |url=https://www.journals.uchicago.edu/doi/10.1086/283437 |journal=The American Naturalist |volume=113 |issue=6 |pages=777–790 |doi=10.1086/283437 |bibcode=1979ANat..113..777W |s2cid=84970789 |access-date=9 November 2021 |archive-date=9 November 2021 |archive-url=https://web.archive.org/web/20211109164204/https://www.journals.uchicago.edu/doi/10.1086/283437|url-status=live}} It may arise terminally on a shoot or from the axil of a leaf.{{cite book |last=Bredmose |first=N. |title=Encyclopedia of Rose Science |chapter=Growth Regulation: Axillary Bud Growth |publisher=Elsevier |year=2003 |doi=10.1016/b0-12-227620-5/00017-3 |pages=374–381|isbn=9780122276200 }} The flower-bearing part of the plant is usually sharply distinguished from the leaf-bearing part, and forms a branch-system called an inflorescence.{{sfn|Balfour|Rendle|1911|p=10}}

Flowers produce two kinds of reproductive cells. Microspores, which divide to become pollen grains, are the male cells; they are borne in the stamens.{{cite book |last1=Salisbury |first1=Frank B. |chapter=Sexual Reproduction |date=1970 |title=Vascular Plants: Form and Function |pages=185–195 |editor-last=Salisbury |editor-first=Frank B. |series=Fundamentals of Botany Series |location=London |publisher=Macmillan Education |doi=10.1007/978-1-349-00364-8_13 |isbn=978-1-349-00364-8 |last2=Parke |first2=Robert V. |doi-broken-date=22 January 2025 |editor2-last=Parke |editor2-first=Robert V.}} The female cells, megaspores, divide to become the egg cell. They are contained in the ovule and enclosed in the carpel; one or more carpels form the pistil.

The flower may consist only of these parts, as in wind-pollinated plants like the willow, where each flower comprises only a few stamens or two carpels.{{sfn|Balfour|Rendle|1911|p=10}} In insect- or bird-pollinated plants, other structures protect the sporophylls and attract pollinators. The individual members of these surrounding structures are known as sepals and petals (or tepals in flowers such as Magnolia where sepals and petals are not distinguishable from each other). The outer series (calyx of sepals) is usually green and leaf-like, and functions to protect the rest of the flower, especially the bud.{{Sfn|De Craene|P.|2010|p=7}}{{Sfn|D. Mauseth|2016|p=225}} The inner series (corolla of petals) is, in general, white or brightly colored, is more delicate in structure, and attracts pollinators by colour, scent, and nectar.{{Sfn|De Craene|P.|2010|p=8}}{{Sfn|D. Mauseth|2016|p=226}}

Most flowers are hermaphroditic, producing both pollen and ovules in the same flower, but some use other devices to reduce self-fertilization. Heteromorphic flowers have carpels and stamens of differing lengths, so animal pollinators cannot easily transfer pollen between them. Homomorphic flowers may use a biochemical self-incompatibility to discriminate between self and non-self pollen grains. Dioecious plants such as holly have male and female flowers on separate plants.{{Cite journal |last=Ainsworth |first=C. |date=August 2000 |title=Boys and Girls Come Out to Play: The Molecular Biology of Dioecious Plants |journal=Annals of Botany |volume=86 |issue=2 |pages=211–221 |doi=10.1006/anbo.2000.1201 |doi-access=free|bibcode=2000AnBot..86..211A }} Monoecious plants have separate male and female flowers on the same plant; these are often wind-pollinated,{{Cite book |last=Batygina |first=T.B. |url=https://books.google.com/books?id=4VOWDwAAQBAJ&q=monoecy&pg=PA43 |title=Embryology of Flowering Plants: Terminology and Concepts, Vol. 3: Reproductive Systems |date=2019 |publisher=CRC Press |isbn=978-1-4398-4436-6 |page=43}} as in maize,{{cite journal | last1=Bortiri | first1=E. | last2=Hake | first2=S. | title=Flowering and determinacy in maize | journal=Journal of Experimental Botany | publisher=Oxford University Press (OUP) | volume=58 | issue=5 | date=2007-01-13 | issn=0022-0957 | doi=10.1093/jxb/erm015 | pages=909–916| pmid=17337752 }} but include some insect-pollinated plants such as Cucurbita squashes.{{cite book |last=Mabberley |first=D. J. |year=2008 |title=The Plant Book: A Portable Dictionary of the Vascular Plants |publisher=Cambridge University Press |location=Cambridge |isbn=978-0-521-82071-4 |page=235}}{{efloras|2|108644 |access-date=21 February 2015}}

{{main|Fertilization|Plant embryogenesis}}

Double fertilization requires two sperm cells to fertilise cells in the ovule. A pollen grain sticks to the stigma at the top of the pistil, germinates, and grows a long pollen tube. 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. The pollen tube grows from the stigma, down the style and into the ovary. When it reaches the micropyle of the ovule, it digests its way into one of the synergids, releasing its contents including the sperm cells. The synergid that the cells were released into degenerates; one sperm makes its way to fertilise the egg cell, producing a diploid (2n) zygote. The second sperm cell fuses with both central cell nuclei, producing a triploid (3n) cell. The zygote develops into an embryo; the triploid cell develops into the endosperm, the embryo's food supply. The ovary develops into a fruit and each ovule into a seed.{{cite journal |last=Berger |first=F. |date=January 2008 |title=Double-fertilization, from myths to reality |journal=Sexual Plant Reproduction |volume=21 |issue=1 |pages=3–5 |doi=10.1007/s00497-007-0066-4 |s2cid=8928640 }}

File:Aesculus hippocastanum fruit.jpg tree, showing the large seed inside the fruit, which is dehiscing or splitting open. ]]

{{main|Fruit|Seed}}

As the embryo and endosperm develop, the wall of the embryo sac enlarges and combines with the nucellus and 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.{{cite journal |last=Eriksson |first=O. |title=Evolution of Seed Size and Biotic Seed Dispersal in Angiosperms: Paleoecological and Neoecological Evidence |journal=International Journal of Plant Sciences |volume=169 |issue=7 |pages=863–870 |year=2008 |doi=10.1086/589888 |bibcode=2008IJPlS.169..863E |s2cid=52905335 }}

Other parts of the flower often contribute to forming the fruit. For example, in the apple, the hypanthium forms the edible flesh, surrounding the ovaries which form the tough cases around the seeds.{{cite web |title=Fruit Anatomy |url=https://ucanr.edu/sites/btfnp/generaltopics/AnatomyPollination/Fruit_Anatomy/ |publisher=University of California |work=Fruit & Nut Research & Information Center |access-date=|url-status = live |archive-url=https://web.archive.org/web/20230502175636/https://ucanr.edu/sites/btfnp/generaltopics/AnatomyPollination/Fruit_Anatomy/ |archive-date=2 May 2023}}

Apomixis, setting seed without fertilization, is found naturally in about 2.2% of angiosperm genera.{{cite journal |last1=Hojsgaard |first1=D. |last2=Klatt |first2=S. |last3=Baier |first3=R. |last4=Carman |first4=J.G. |last5=Hörandl |first5=E. |display-authors=3 |title=Taxonomy and Biogeography of Apomixis in Angiosperms and Associated Biodiversity Characteristics |journal=Critical Reviews in Plant Sciences |volume=33 |issue=5 |pages=414–427 |date=September 2014 |pmid=27019547 |pmc=4786830 |doi=10.1080/07352689.2014.898488 |bibcode=2014CRvPS..33..414H }} Some angiosperms, including many citrus varieties, are able to produce fruits through a type of apomixis called nucellar embryony.{{cite book |last=Gentile |first=Alessandra |title=The Citrus Genome |url=https://books.google.com/books?id=hsPXDwAAQBAJ |date=18 March 2020 |publisher=Springer Nature |isbn=978-3-030-15308-3 |page=171 |access-date=13 December 2020 |archive-date=14 April 2021 |archive-url=https://web.archive.org/web/20210414224058/https://books.google.com/books?id=hsPXDwAAQBAJ |url-status=live}}

{{Excerpt|Sexual selection in flowering plants}}

Charles Darwin in his 1878 book The Effects of Cross and Self-Fertilization in the Vegetable Kingdom{{cite book |last=Darwin |first=Charles R. |author-link=Charles Darwin |year=1878 |title=The effects of cross and self fertilisation in the vegetable kingdom |location=London |publisher=John Murray |url=https://darwin-online.org.uk/converted/pdf/1878_Fertilisation_F1251.pdf}} in the initial paragraph of chapter XII noted "The first and most important of the conclusions which may be drawn from the observations given in this volume, is that generally cross-fertilisation is beneficial and self-fertilisation often injurious, at least with the plants on which I experimented." Flowers emerged in plant evolution as an adaptation for the promotion of cross-fertilisation (outcrossing), a process that allows the masking of deleterious mutations in the genome of progeny. The masking effect is known as genetic complementation.{{cite journal |last1=Bernstein |first1=Harris |last2=Byerly |first2=Henry C. |last3=Hopf |first3=Frederic A. |last4=Michod |first4=Richard E. |title=Genetic Damage, Mutation, and the Evolution of Sex |journal=Science |volume=229 |issue=4719 |date=20 September 1985 |doi=10.1126/science.3898363 |pages=1277–1281|pmid=3898363 |bibcode=1985Sci...229.1277B }} Meiosis in flowering plants provides a direct mechanism for repairing DNA through genetic recombination in reproductive tissues.{{cite journal |last=Hörandl |first=Elvira |title=Apomixis and the paradox of sex in plants |journal=Annals of Botany |volume=134 |issue=1 |date=7 June 2024 |pmid=38497809 |pmc=11161571 |doi=10.1093/aob/mcae044 |doi-access=free |pages=1–18 |url=https://academic.oup.com/aob/advance-article-pdf/doi/10.1093/aob/mcae044/57132588/mcae044.pdf |access-date=17 January 2025}} Sexual reproduction appears to be required for maintaining long-term genomic integrity and only infrequent combinations of extrinsic and intrinsic factors permit shifts to asexuality. Thus the two fundamental aspects of sexual reproduction in flowering plants, cross-fertilization (outcrossing) and meiosis appear to be maintained respectively by the advantages of genetic complementation and recombinational repair.

{{main|Human uses of plants}}

File:Rice Harvest 2020 - 50248478521.jpg in Arkansas, 2020]]

File:Daal_after_Tadka_Pulse_Soup_India.jpg, Indian lentil soup]]

Agriculture is almost entirely dependent on angiosperms, which provide virtually all plant-based food and fodder for livestock. Much of this food derives from a small number of flowering plant families.{{cite web |last1=Dilcher |first1=David L. |last2=Cronquist |first2=Arthur |last3=Zimmermann |first3=Martin Huldrych |last4=Stevens |first4=Peter |last5=Stevenson |first5=Dennis William |last6=Berry |first6=Paul E. |author-link2=Arthur Cronquist |author-link4=Peter F. Stevens |title=Angiosperm: Significance to Humans |url=https://www.britannica.com/plant/angiosperm/Significance-to-humans |website=Encyclopedia Britannica |date=8 March 2016}} For instance, half of the world's calorie intake is supplied by just three plants – wheat, rice and maize.{{Cite news |last=McKie |first=Robin |date=16 July 2017 |title=Maize, rice, wheat: alarm at rising climate risk to vital crops |newspaper=The Observer |url=https://www.theguardian.com/environment/2017/jul/15/climate-change-food-famine-study |access-date=30 July 2023 }}

Flowering plants provide a diverse range of materials in the form of wood, paper, fibers such as cotton, flax, and hemp, medicines such as digoxin and opioids, and decorative and landscaping plants. Coffee and hot chocolate are beverages from flowering plants (in the Rubiaceae and Malvaceae respectively).

File:Kingfisher-iris.jpg: Kingfisher and iris kachō-e woodblock print by Ohara Koson (late 19th century)]]

Both real and fictitious plants play a wide variety of roles in literature and film.{{cite journal |title=Literary Plants |journal=Nature Plants |date=2015 |volume=1 |issue=11 |page=15181 |doi=10.1038/nplants.2015.181 |pmid=27251545 |doi-access=free |bibcode=2015NatPl...115181. }} Flowers are the subjects of many poems by poets such as William Blake, Robert Frost, and Rabindranath Tagore.{{cite web |title=Flower Poems |url=http://www.poemhunter.com/poems/flower/ |website=Poem Hunter |access-date=21 June 2016}} Bird-and-flower painting ({{Transliteration|zh|Huaniaohua}}) is a kind of Chinese painting that celebrates the beauty of flowering plants.{{Cite web |title=Nature's Song: Chinese Bird and Flower Paintings |url=https://museum.wales/cardiff/whatson/9245/Natures-Song-Traditional-Chinese-Bird-and-Flower-Paintings-/ |access-date=2022-08-04 |website=Museum Wales |language=en |archive-date=4 August 2022 |archive-url=https://web.archive.org/web/20220804231344/https://museum.wales/cardiff/whatson/9245/Natures-Song-Traditional-Chinese-Bird-and-Flower-Paintings-/ |url-status=dead }} Flowers have been used in literature to convey meaning by authors including William Shakespeare.{{cite web |title=The Language of Flowers |url=http://www.folger.edu/template.cfm?cid=3192 |publisher=Folger Shakespeare Library |access-date=2013-05-31 |archive-url=https://web.archive.org/web/20140919185926/http://www.folger.edu/template.cfm?cid=3192 |archive-date=2014-09-19 |url-status=dead}}

Flowers are used in a variety of art forms which arrange cut or living plants, such as bonsai, ikebana, and flower arranging. Ornamental plants have sometimes changed the course of history, as in tulipomania.{{cite web |last=Lambert |first=Tim |title=A Brief History of Gardening |url=http://www.localhistories.org/gardening.html |publisher=British Broadcasting Corporation |access-date=21 June 2016 |date=2014}} Many countries and regions have floral emblems; a survey of 70 of these found that the most popular flowering plant family for such emblems is Orchidaceae at 15.7% (11 emblems), followed by Fabaceae at 10% (7 emblems), and Asparagaceae, Asteraceae, and Rosaceae all at 5.7% (4 emblems each).{{cite book |last1=Lim |first1=Reuben |last2=Tan |first2=Heok |last3=Tan |first3=Hugh |year=2013 |title=Official Biological Emblems of the World |publisher=Raffles Museum of Biodiversity Research |location=Singapore |isbn=978-9-8107-4147-1 |url=https://www.researchgate.net/publication/283008038}}

{{further|Conservation biology|Effects of climate change on plant biodiversity}}

File:Viola_calcarata20052002fleur2.JPG, a species highly vulnerable to climate change.{{Cite journal |last1=Block |first1=Sebastián |last2=Maechler |first2=Marc-Jacques |last3=Levine |first3=Jacob I. |last4=Alexander |first4=Jake M. |last5=Pellissier |first5=Loïc |last6=Levine |first6=Jonathan M. |date=26 August 2022 |title=Ecological lags govern the pace and outcome of plant community responses to 21st-century climate change |journal=Ecology Letters |volume=25 |issue=10 |pages=2156–2166 |doi=10.1111/ele.14087 |pmid=36028464 |pmc=9804264 |bibcode=2022EcolL..25.2156B }}]]

Human impact on the environment has driven a range of species extinct and is threatening even more today. Multiple organizations such as IUCN and Royal Botanic Gardens, Kew suggest that around 40% of plant species are threatened with extinction.{{Cite journal |last1=Lughadha |first1=Eimear Nic|last2=Bachman |first2=Steven P. |last3=Leão |first3=Tarciso C. C. |last4=Forest |first4=Félix |last5=Halley |first5=John M. |last6=Moat |first6=Justin |last7=Acedo|first7=Carmen |last8=Bacon |first8=Karen L. |last9=Brewer |first9=Ryan F. A. |last10=Gâteblé |first10=Gildas |last11=Gonçalves |first11=Susana C.|last12=Govaerts |first12=Rafaël |last13=Hollingsworth |first13=Peter M. |last14=Krisai-Greilhuber |first14=Irmgard |last15=de Lirio |first15=Elton J. |last16=Moore |first16=Paloma G. P. |last17=Negrão |first17=Raquel |last18=Onana |first18=Jean Michel |last19=Rajaovelona |first19=Landy R. |last20=Razanajatovo |first20=Henintsoa |last21=Reich |first21=Peter B. |last22=Richards |first22=Sophie L. |last23=Rivers |first23=Malin C. |last24=Cooper |first24=Amanda |last25=Iganci |first25=João |last26=Lewis |first26=Gwilym P. |last27=Smidt |first27=Eric C. |last28=Antonelli |first28=Alexandre |last29=Mueller |first29=Gregory M. |last30=Walker |first30=Barnaby E. |date=29 September 2020 |title=Extinction risk and threats to plants and fungi |journal=Plants People Planet |volume=2 |issue=5 |pages=389–408 |doi=10.1002/ppp3.10146 |s2cid=225274409 |doi-access=free |bibcode=2020PlPP....2..389N |hdl=10316/101227 |hdl-access=free }} The majority are threatened by habitat loss, but activities such as logging of wild timber trees and collection of medicinal plants, or the introduction of non-native invasive species, also play a role.{{cite journal |last1=Wiens |first1=John J. |title=Climate-Related Local Extinctions Are Already Widespread among Plant and Animal Species |journal=PLOS Biology |date=2016 |volume=14 |issue=12 |pages=e2001104 |doi=10.1371/journal.pbio.2001104 |doi-access=free |pmid=27930674 |pmc=5147797 |hdl=10150/622757 |hdl-access=free }}{{cite book |last=Shivanna |first=K. R. |title=Biodiversity and Chemotaxonomy |chapter=The 'Sixth Mass Extinction Crisis' and Its Impact on Flowering Plants |series=Sustainable Development and Biodiversity |publisher=Springer International Publishing |publication-place=Cham |volume=24 |date=2019 |pages=15–42 |isbn=978-3-030-30745-5 |doi=10.1007/978-3-030-30746-2_2}}

Relatively few plant diversity assessments currently consider climate change, yet it is starting to impact plants as well. About 3% of flowering plants are very likely to be driven extinct within a century at {{convert|2|C-change|F-change}} of global warming, and 10% at {{convert|3.2|C-change|F-change}}.Parmesan, C., M.D. Morecroft, Y. Trisurat et al. (2022) [https://www.ipcc.ch/report/ar6/wg2/downloads/report/IPCC_AR6_WGII_Chapter02.pdf Chapter 2: Terrestrial and Freshwater Ecosystems and Their Services] in {{cite book | title=Climate Change 2022 – Impacts, Adaptation and Vulnerability | chapter=Terrestrial and Freshwater Ecosystems and Their Services | publisher=Cambridge University Press | date=2023 | isbn=978-1-009-32584-4 | doi=10.1017/9781009325844.004 | pages=197–378 |url=https://www.ipcc.ch/report/ar6/wg2/}} In worst-case scenarios, half of all tree species may be driven extinct by climate change over that timeframe.

Conservation in this context is the attempt to prevent extinction, whether in situ by protecting plants and their habitats in the wild, or ex situ in seed banks or as living plants.{{cite web |title=Botanic Gardens and Plant Conservation |url=https://www.bgci.org/about/botanic-gardens-and-plant-conservation/ |website=Botanic Gardens Conservation International |access-date=19 July 2023}} Some 3000 botanic gardens around the world maintain living plants, including over 40% of the species known to be threatened, as an "insurance policy against extinction in the wild."{{cite web |title=Plant Conservation Around the World |url=https://www.botanic.cam.ac.uk/festival-of-plants-2020/fop-day3-conservation/plant-conservation-around-the-world/ |website=Cambridge University Botanic Garden |access-date=19 July 2023 |date=2020}} The United Nations' Global Strategy for Plant Conservation asserts that "without plants, there is no life". It aims to "halt the continuing loss of plant diversity" throughout the world.{{cite web |title=Updated Global Strategy for Plant Conservation 2011–2020 |url=https://www.cbd.int/gspc/ |website=Convention on Biological Diversity |access-date=19 July 2023 |date=3 July 2023}}{{Clear}}

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• {{cite journal |last1=Takhtajan |first1=A. |author-link=Takhtajan |title=The Taxa of the Higher Plants above the Rank of Order |journal=Taxon |date=June 1964 |volume=13 |issue=5 |pages=160–164 |doi=10.2307/1216134 |jstor=1216134|bibcode=1964Taxon..13..160T }}
• {{cite journal |last1=Takhtajan |first1=A. |author-link=Takhtajan |title=Outline of the Classification of Flowering Plants (Magnoliophyta) |journal=Botanical Review |date=July–September 1980 |volume=46 |issue=3 |pages=225–359 |jstor=4353970 |doi=10.1007/bf02861558 |bibcode=1980BotRv..46..225T |s2cid=30764910}}
• {{cite journal |last1=Zeng |first1=Liping |last2=Zhang |first2=Qiang |last3=Sun |first3=Renran |last4=Kong |first4=Hongzhi |last5=Zhang |first5=Ning |last6=Ma |first6=Hong |title=Resolution of deep angiosperm phylogeny using conserved nuclear genes and estimates of early divergence times |journal=Nature Communications |date=24 September 2014 |volume=5 |issue=4956 |pages=4956 |doi=10.1038/ncomms5956 |pmid=25249442 |pmc=4200517 |bibcode=2014NatCo...5.4956Z |ref=none}}

{{Refend}}

{{Refbegin|30em}}

• {{cite web |last1=Cole |first1=Theodor C.H. |last2=Hilger |first2=Harmut H. |first3=Peter F. |last3=Stevens |date=2017 |url=http://www.biologie.fu-berlin.de/sysbot/poster/poster1.pdf |title=Angiosperm Phylogeny Poster{{spaced ndash}} Flowering Plant Systematics |ref=none}}
• {{cite web |last1=Watson |first1=L. |last2=Dallwitz |first2=M.J. |date=1992 |url=http://www.biologie.uni-hamburg.de/b-online/delta/angio |title=The Families of Flowering Plants: Descriptions, Illustrations, Identification, and Information Retrieval |version=14 December 2000 |archive-url=https://web.archive.org/web/20140802080838/http://www.biologie.uni-hamburg.de/b-online/delta/angio |archive-date=2014-08-02 |ref=none}}
• {{EOL|282}}

{{Refend}}

{{Plant classification}}

{{Botany}}

{{Angiosperm orders}}

{{Lists of angiosperm families}}

{{Life on Earth}}

{{Subject bar|Plants|c=Angiosperms|commonscat=y|species=Magnoliophyta|b=Magnoliophyta|auto=1}}

{{Taxonbar|from=Q25314}}

{{Authority control}}

{{DEFAULTSORT:Flowering Plant}}

Angiosperms

Category:Plant sexuality

Category:Plants

Category:Pollination