Flowering plant: Difference between revisions

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m Added clarity to lifespan to accurately describe biennials which live for two years.
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'''Flowering plants''' are [[plant]]s that bear [[flowers]] and [[fruit]]s, and form the [[clade]] '''Angiospermae''' ({{IPAc-en|ˌ|æ|n|dʒ|i|ə|ˈ|s|p|ər|m|i:}}).<ref>{{cite book |last=Lindley |first=J. |year=1830 |url=https://www.biodiversitylibrary.org/item/31944#page/21/mode/1up |title=Introduction to the Natural System of Botany |location=London |publisher=Longman, Rees, Orme, Brown, and Green |pages=xxxvi |no-pp=true |access-date=29 January 2018 |archive-date=27 August 2017 |archive-url=https://web.archive.org/web/20170827171755/http://www.biodiversitylibrary.org/item/31944#page/21/mode/1up |url-status=live }}</ref><ref>{{cite journal |last1=Cantino |first1=Philip D. |last2=Doyle |first2=James A. |last3=Graham |first3=Sean W. |last4=Judd |first4=Walter S. |last5=Olmstead |first5=Richard G. |last6=Soltis |first6=Douglas E. |author-link6=Douglas E. Soltis |last7=Soltis |first7=Pamela S. |author-link7=Pamela S. Soltis |last8=Donoghue |first8=Michael J. |display-authors=3 |year=2007 |title= Towards a phylogenetic nomenclature of ''Tracheophyta'' |journal=Taxon |volume=56 |issue=3 |pages=E1–E44 |doi=10.2307/25065865|jstor=25065865 }}</ref> The term '''angiosperm''' is derived from the [[Ancient Greek|Greek]] words {{lang|grc|ἀγγεῖον}} ({{lang|grc-Latn|angeion}}; 'container, vessel') and {{lang|grc|σπέρμα}} ({{lang|grc-Latn|sperma}}; 'seed'), meaning that the [[seed]]s are enclosed within a fruit. The group was formerly called '''Magnoliophyta'''.{{sfn|Takhtajan|1980}}
'''Flowering plants''' are [[plant]]s that bear [[flowers]] and [[fruit]]s, and form the [[clade]] '''Angiospermae''' ({{IPAc-en|ˌ|æ|n|dʒ|i|ə|ˈ|s|p|ɜːr|m|i:}}).<ref>{{cite book |last=Lindley |first=J. |year=1830 |url=https://www.biodiversitylibrary.org/item/31944#page/21/mode/1up |title=Introduction to the Natural System of Botany |location=London |publisher=Longman, Rees, Orme, Brown, and Green |page=xxxvi |no-pp=true |access-date=29 January 2018 |archive-date=27 August 2017 |archive-url=https://web.archive.org/web/20170827171755/http://www.biodiversitylibrary.org/item/31944#page/21/mode/1up |url-status=live }}</ref><ref>{{cite journal |last1=Cantino |first1=Philip D. |last2=Doyle |first2=James A. |last3=Graham |first3=Sean W. |last4=Judd |first4=Walter S. |last5=Olmstead |first5=Richard G. |last6=Soltis |first6=Douglas E. |author-link6=Douglas E. Soltis |last7=Soltis |first7=Pamela S. |author-link7=Pamela S. Soltis |last8=Donoghue |first8=Michael J. |display-authors=3 |year=2007 |title= Towards a phylogenetic nomenclature of ''Tracheophyta'' |journal=Taxon |volume=56 |issue=3 |pages=E1–E44 |doi=10.2307/25065865|jstor=25065865 }}</ref> The term '''angiosperm''' is derived from the [[Ancient Greek|Greek]] words {{lang|grc|ἀγγεῖον}} ({{lang|grc-Latn|angeion}}; 'container, vessel') and {{lang|grc|σπέρμα}} ({{lang|grc-Latn|sperma}}; 'seed'), meaning that the [[seed]]s are enclosed within a fruit. The group was formerly called '''Magnoliophyta'''.{{sfn|Takhtajan|1980}}


Angiosperms are by far the most diverse group of [[Embryophyte|land plants]] with 64 [[Order (biology)|orders]], 416 [[Family (biology)|families]], approximately 13,000 known [[Genus|genera]] and 300,000 known [[species]].<ref>{{cite journal |last1=Christenhusz |first1=M. J. M. |last2=Byng |first2=J. W. |year=2016 |title=The number of known plants species in the world and its annual increase |journal=Phytotaxa |volume=261 |pages=201–217 |url=https://biotaxa.org/Phytotaxa/article/download/phytotaxa.261.3.1/20598 |doi=10.11646/phytotaxa.261.3.1 |issue=3 |doi-access=free |bibcode=2016Phytx.261..201C |access-date=21 February 2022 |archive-date=6 April 2017 |archive-url=https://web.archive.org/web/20170406045346/http://biotaxa.org/Phytotaxa/article/download/phytotaxa.261.3.1/20598 |url-status=live }}</ref> They include all [[forb]]s (flowering plants without a [[wood]]y [[Plant stem|stem]]), [[grass]]es and grass-like plants, a vast majority of [[broad-leaved tree]]s, [[shrub]]s and [[vine]]s, and most [[aquatic plant]]s. Angiosperms are distinguished from the other major [[seed plant]] clade, the [[gymnosperm]]s, by having [[flower]]s, [[xylem]] consisting of [[vessel element]]s instead of [[tracheids]], [[endosperm]] within their seeds, and fruits that completely envelop the seeds. The ancestors of flowering plants diverged from the common ancestor of all living gymnosperms before the end of the [[Carboniferous]], over 300 million years ago. In the [[Cretaceous]], angiosperms [[Cretaceous Terrestrial Revolution|diversified explosively]], becoming the dominant group of plants across the planet.
Angiosperms are by far the most diverse group of [[Embryophyte|land plants]] with 64 [[Order (biology)|orders]], 416 [[Family (biology)|families]], approximately 13,000 known [[Genus|genera]] and 300,000 known [[species]].<ref>{{cite journal |last1=Christenhusz |first1=M. J. M. |last2=Byng |first2=J. W. |year=2016 |title=The number of known plants species in the world and its annual increase |journal=Phytotaxa |volume=261 |pages=201–217 |url=https://biotaxa.org/Phytotaxa/article/download/phytotaxa.261.3.1/20598 |doi=10.11646/phytotaxa.261.3.1 |issue=3 |doi-access=free |bibcode=2016Phytx.261..201C |access-date=21 February 2022 |archive-date=6 April 2017 |archive-url=https://web.archive.org/web/20170406045346/http://biotaxa.org/Phytotaxa/article/download/phytotaxa.261.3.1/20598 |url-status=live }}</ref> They include all [[forb]]s (flowering plants without a [[wood]]y [[Plant stem|stem]]), [[grass]]es and grass-like plants, a vast majority of [[broad-leaved tree]]s, [[shrub]]s and [[vine]]s, and most [[aquatic plant]]s. Angiosperms are distinguished from the other major [[seed plant]] clade, the [[gymnosperm]]s, by having [[flower]]s, [[xylem]] consisting of [[vessel element]]s instead of [[tracheids]], [[endosperm]] within their seeds, and fruits that completely envelop the seeds. The ancestors of flowering plants diverged from the common ancestor of all living gymnosperms before the end of the [[Carboniferous]], over 300 million years ago. In the [[Cretaceous]], angiosperms [[Cretaceous Terrestrial Revolution|diversified explosively]], becoming the dominant group of plants across the planet.
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|[[Flower]]s ||The [[reproductive organ]]s of flowering plants, not found in any other [[seed plants]].<ref>{{cite web |title=Angiosperms {{!}} OpenStax Biology 2e |url=https://courses.lumenlearning.com/suny-osbiology2e/chapter/angiosperms/ |access-date=19 July 2021 |website=courses.lumenlearning.com |archive-date=19 July 2021|archive-url=https://web.archive.org/web/20210719225359/https://courses.lumenlearning.com/suny-osbiology2e/chapter/angiosperms/ |url-status=live}}</ref> || [[File:Daffodil flower in section, labelled.svg|thumb|none|A ''[[Narcissus (plant)|Narcissus]]'' flower in section. [[Petal]]s and [[sepal]]s are replaced here by a fused tube, the corona, and tepals.]]
|[[Flower]]s ||The [[reproductive organ]]s of flowering plants, not found in any other [[seed plants]].<ref>{{cite web |title=Angiosperms {{!}} OpenStax Biology 2e |url=https://courses.lumenlearning.com/suny-osbiology2e/chapter/angiosperms/ |access-date=19 July 2021 |website=courses.lumenlearning.com |archive-date=19 July 2021|archive-url=https://web.archive.org/web/20210719225359/https://courses.lumenlearning.com/suny-osbiology2e/chapter/angiosperms/ |url-status=live}}</ref> || [[File:Daffodil flower in section, labelled.svg|thumb|none|A ''[[Narcissus (plant)|Narcissus]]'' flower in section. [[Petal]]s and [[sepal]]s are replaced here by a fused tube, the corona, and tepals.]]
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|Reduced [[Gametophyte#Seed plants|gametophytes]], three [[cell (biology)|cells]] in male, seven cells with eight nuclei in female (except for basal angiosperms)<ref>{{cite journal |last1=Friedman |first1=William E. |last2=Ryerson |first2=Kirsten C. |title=Reconstructing the ancestral female gametophyte of angiosperms: Insights from Amborella and other ancient lineages of flowering plants |journal=American Journal of Botany |volume=96 |issue=1 |date=2009 |doi=10.3732/ajb.0800311 |pages=129–143|pmid=21628180 |bibcode=2009AmJB...96..129F }}</ref> ||The gametophytes are smaller than those of gymnosperms.<ref>{{cite book |author1=Raven, Peter H. |author2=Evert, Ray F. |author3=Eichhorn, Susan E. |title=Biology of Plants |url=https://archive.org/details/biologyofplants00rave_0 |url-access=registration |year=2005 |publisher=W. H. Freeman |isbn=978-0-7167-1007-3 |pages=[https://archive.org/details/biologyofplants00rave_0/page/376 376]–}}</ref> The smaller size of the [[pollen]] reduces the time between pollination and [[fertilization]], which in gymnosperms is up to a year.<ref>{{cite journal |last=Williams |first=Joseph H. |title=The evolution of pollen germination timing in flowering plants: Austrobaileya scandens (Austrobaileyaceae) |journal=AoB Plants |volume=2012 |pages=pls010 |date=2012 |pmid=22567221 |pmc=3345124 |doi=10.1093/aobpla/pls010 }}</ref>   
|Reduced [[Gametophyte#Seed plants|gametophytes]], three [[cell (biology)|cells]] in male, seven cells with eight nuclei in female (except for basal angiosperms)<ref>{{cite journal |last1=Friedman |first1=William E. |last2=Ryerson |first2=Kirsten C. |title=Reconstructing the ancestral female gametophyte of angiosperms: Insights from Amborella and other ancient lineages of flowering plants |journal=American Journal of Botany |volume=96 |issue=1 |date=2009 |doi=10.3732/ajb.0800311 |pages=129–143|pmid=21628180 |bibcode=2009AmJB...96..129F }}</ref> ||The gametophytes are smaller than those of gymnosperms.<ref>{{cite book |author1=Raven, Peter H. |author2=Evert, Ray F. |author3=Eichhorn, Susan E. |title=Biology of Plants |url=https://archive.org/details/biologyofplants00rave_0 |url-access=registration |year=2005 |publisher=W. H. Freeman |isbn=978-0-7167-1007-3 |pages=[https://archive.org/details/biologyofplants00rave_0/page/376 376]–}}</ref> The smaller size of the [[pollen]] reduces the time between pollination and [[fertilization]], which in gymnosperms is up to a year.<ref>{{cite journal |last=Williams |first=Joseph H. |title=The evolution of pollen germination timing in flowering plants: Austrobaileya scandens (Austrobaileyaceae) |journal=AoB Plants |volume=2012 |article-number=pls010 |date=2012 |pmid=22567221 |pmc=3345124 |doi=10.1093/aobpla/pls010 }}</ref>   
|rowspan=2 | [[File:Angiosperm embryo sac with female gametophyte.JPG|thumb|none|[[Embryo sac]] is a reduced female [[gametophyte]].]]
|rowspan=2 | [[File:Angiosperm embryo sac with female gametophyte.JPG|thumb|none|[[Embryo sac]] is a reduced female [[gametophyte]].]]
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|[[Endosperm]] ||Endosperm forms after fertilization but before the [[zygote]] divides. It provides food for the developing [[embryo]], the [[cotyledon]]s, and sometimes the [[seedling]].<ref>{{cite journal |last1=Baroux |first1=C. |last2=Spillane |first2=C. |last3=Grossniklaus |first3=U. |title=Evolutionary origins of the endosperm in flowering plants |journal=Genome Biology |volume=3 |article-number=reviews1026.1 |year=2002 |issue=9 |pages=reviews1026.1 |doi=10.1186/gb-2002-3-9-reviews1026 |pmid=12225592 |pmc=139410 |doi-access=free }}</ref> <!--cell/image shared with row above-->
|[[Endosperm]] ||Endosperm forms after fertilization but before the [[zygote]] divides. It provides food for the developing [[embryo]], the [[cotyledon]]s, and sometimes the [[seedling]].<ref>{{cite journal |last1=Baroux |first1=C. |last2=Spillane |first2=C. |last3=Grossniklaus |first3=U. |title=Evolutionary origins of the endosperm in flowering plants |journal=Genome Biology |volume=3 |article-number=reviews1026.1 |year=2002 |issue=9 |pages=reviews1026.1 |doi=10.1186/gb-2002-3-9-reviews1026 |pmid=12225592 |pmc=139410 |doi-access=free }}</ref> <!--cell/image shared with row above-->
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|Closed [[carpel]] enclosing the [[ovule]]s. ||Once the ovules are fertilised, the carpels, often with surrounding tissues, develop into fruits. Gymnosperms have unenclosed seeds.<ref>{{cite journal |last=Gonçalves |first=Beatriz |title=Case not closed: the mystery of the origin of the carpel |journal=EvoDevo |volume=12 |issue=1 |date=2021-12-15 |page=14 |issn=2041-9139 |doi=10.1186/s13227-021-00184-z |pmid=34911578 |pmc=8672599 |doi-access=free }}</ref> || [[File:Alternating peas in peapod (cropped).jpg|thumb|none|Peas (seeds, from ovules) inside pod (fruit, from fertilised carpel).]]
|Closed [[carpel]] enclosing the [[ovule]]s. ||Once the ovules are fertilised, the carpels, often with surrounding tissues, develop into fruits. Gymnosperms have unenclosed seeds.<ref>{{cite journal |last=Gonçalves |first=Beatriz |title=Case not closed: the mystery of the origin of the carpel |journal=EvoDevo |volume=12 |issue=1 |date=2021-12-15 |article-number=14 |issn=2041-9139 |doi=10.1186/s13227-021-00184-z |pmid=34911578 |pmc=8672599 |doi-access=free }}</ref> || [[File:Alternating peas in peapod (cropped).jpg|thumb|none|Peas (seeds, from ovules) inside pod (fruit, from fertilised carpel).]]
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| [[Xylem]] made of [[vessel element]]s || Open vessel elements are stacked end to end to form continuous tubes, whereas gymnosperm xylem is made of tapered [[tracheids]] connected by small [[Pit (botany)|pits]].<ref>{{cite book | last=Baas | first=Pieter | title=New Perspectives in Wood Anatomy | chapter=Systematic, phylogenetic, and ecological wood anatomy — History and perspectives | series=Forestry Sciences | publisher=Springer Netherlands | publication-place=Dordrecht | year=1982 | volume=1 | isbn=978-90-481-8269-5 | issn=0924-5480 | doi=10.1007/978-94-017-2418-0_2 | pages=23–58}}</ref> || [[File:Herbaceous Dicot Stem Xylem Vessels Cucurbita (35463815631).jpg|thumb|none| [[Xylem]] vessels (long tubes).]]
| [[Xylem]] made of [[vessel element]]s || Open vessel elements are stacked end to end to form continuous tubes, whereas gymnosperm xylem is made of tapered [[tracheids]] connected by small [[Pit (botany)|pits]].<ref>{{cite book | last=Baas | first=Pieter | title=New Perspectives in Wood Anatomy | chapter=Systematic, phylogenetic, and ecological wood anatomy — History and perspectives | series=Forestry Sciences | publisher=Springer Netherlands | publication-place=Dordrecht | year=1982 | volume=1 | isbn=978-90-481-8269-5 | issn=0924-5480 | doi=10.1007/978-94-017-2418-0_2 | pages=23–58}}</ref> || [[File:Herbaceous Dicot Stem Xylem Vessels Cucurbita (35463815631).jpg|thumb|none| [[Xylem]] vessels (long tubes).]]
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</gallery>


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.<ref>{{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)}}</ref> The smallest are ''[[Wolffia]]'' duckweeds which float on freshwater, each plant less than {{convert|2|mm|in|2}} across.<ref>{{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 }}</ref>
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.<ref>{{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)}}</ref> The smallest are ''[[Wolffia]]'' duckweeds which float on freshwater, each plant less than {{convert|2|mm|in|2}} across.<ref>{{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 |archive-date=25 November 2009 }}</ref>


<gallery class=center mode=nolines widths=225px heights=225px caption="Photosynthetic and parasitic">
<gallery class=center mode=nolines widths=225px heights=225px caption="Photosynthetic and parasitic">
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</gallery>
</gallery>


Considering their method of obtaining energy, some 99% of flowering plants are [[photosynthetic]] [[autotroph]]s, deriving their energy from sunlight and using it to create molecules such as [[sugar]]s. The remainder are [[parasitic]], whether [[myco-heterotrophy|on fungi]] like the [[orchids]] for part or all of their life-cycle,<ref>{{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 }}</ref> or [[Parasitic plant|on other plants]], either wholly like the broomrapes, ''[[Orobanche]]'', or partially like the witchweeds, ''[[Striga]]''.<ref>{{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 }}</ref>
Considering their method of obtaining energy, some 99% of flowering plants are [[photosynthetic]] [[autotroph]]s, deriving their energy from sunlight and using it to create molecules such as [[sugar]]s. The remainder are [[parasitic]], whether [[myco-heterotrophy|on fungi]] (myco-heterotrophic, formerly thought to be [[saprophytic]]) like the [[orchids]] for part or all of their life-cycle,<ref>{{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 }}</ref> or [[Parasitic plant|on other plants]], either wholly like the broomrapes, ''[[Orobanche]]'', or partially like the witchweeds, ''[[Striga]]''.<ref>{{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 }}</ref>


<gallery class=center mode=nolines widths=225px heights=225px caption="Hot, cold, wet, dry, fresh, salt">
<gallery class=center mode=nolines widths=225px heights=225px caption="Hot, cold, wet, dry, fresh, salt">
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==== Internal ====
==== Internal ====


The main groups of living angiosperms are:<ref>{{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 }}</ref>{{sfn|APG|2016}}
The main groups of living angiosperms are:<ref>{{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 |article-number=6930 |doi=10.1038/s41467-021-26922-4 |pmid=34836973 |pmc=8626473 |bibcode=2021NatCo..12.6930G |doi-access=free }}</ref>{{sfn|APG|2016}}


{{barlabel |size=20
{{barlabel |size=20
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               }}
               }}
             |2={{clade
             |2={{clade
               |1=[[Monocots]] [[File:White orchid in Clara bog. 03.jpg|45px]] c. 70,000 spp.<ref>{{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 }}</ref> 3-part flowers, 1 [[cotyledon]], 1-pore pollen, usu. parallel-veined leaves  &nbsp; |bar1=green
               |1=[[Monocots]] [[File:White orchid in Clara bog. 03.jpg|45px]] c. 70,000 spp.<ref>{{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 |page=49 |doi=10.1186/s12862-015-0320-6 |pmc=4377182 |pmid=25887386 |bibcode=2015BMCEE..15...49M |doi-access=free }}</ref> 3-part flowers, 1 [[cotyledon]], 1-pore pollen, usu. parallel-veined leaves  &nbsp; |bar1=green
               |2={{clade
               |2={{clade
                   |1=[[Ceratophyllales]] [[File:CeratophyllumSubmersum.jpg|70px]] c. 6 spp.<ref name="Palmer-2004"/> [[aquatic plant]]s  |bar1=green
                   |1=[[Ceratophyllales]] [[File:CeratophyllumSubmersum.jpg|70px]] c. 6 spp.<ref name="Palmer-2004"/> [[aquatic plant]]s  |bar1=green
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[[File:Sagaria cilentana (cropped).jpg|thumb|upright|[[Adaptive radiation]] in the [[Cretaceous]] created many flowering plants, such as ''[[Sagaria]]'' in the [[Ranunculaceae]].]]
[[File:Sagaria cilentana (cropped).jpg|thumb|upright|[[Adaptive radiation]] in the [[Cretaceous]] created many flowering plants, such as ''[[Sagaria]]'' in the [[Ranunculaceae]].]]


Fossilised [[spore]]s suggest that land plants ([[embryophyte]]s) have existed for at least 475&nbsp;million years.<ref>{{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 }}</ref> However, angiosperms [[abominable mystery|appear suddenly]] and in great diversity in the fossil record in the [[Early Cretaceous]] (~130 mya).<ref>{{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}}</ref><ref>{{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|bibcode=2009AmJB...96....5F |url-access=subscription }}</ref> Claimed records of flowering plants prior to this are not widely accepted,<ref>{{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|url-access=subscription }}</ref> as all supposed pre-Cretaceous “flowers” can be explained through being misidentifications of other seed plants. Furthermore, almost all of these controversial fossils are described in papers co-authored by the researcher Xin Wang, such as the particularly debated ''[[Nanjinganthus]]''.<ref>{{cite journal |last1=Sokoloff |first1=Dmitry D. |last2=Remizowa |first2=Margarita V. |last3=El |first3=Elena S. |last4=Rudall |first4=Paula J. |last5=Bateman |first5=Richard M. |title=Supposed Jurassic angiosperms lack pentamery, an important angiosperm-specific feature |journal=New Phytologist |date=October 2020 |volume=228 |issue=2 |pages=420–426 |doi=10.1111/nph.15974|pmid=31418869 |bibcode=2020NewPh.228..420S }}</ref> Molecular evidence suggests that the ancestors of angiosperms diverged from the [[gymnosperms]] during the late [[Devonian]], about 365 million years ago.<ref>{{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}}</ref> The origin time of the [[crown group]] of flowering plants remains contentious.<ref>{{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|url-access=subscription }}</ref> By the Late Cretaceous, angiosperms appear to have dominated environments formerly occupied by [[fern]]s and gymnosperms. Large [[Canopy (biology)|canopy]]-forming trees replaced [[conifer]]s as the dominant trees close to the end of the Cretaceous, 66 million years ago.<ref>{{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}}</ref> The radiation of herbaceous angiosperms occurred much later.<ref>{{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 }}</ref>
Fossilised [[spore]]s suggest that land plants ([[embryophyte]]s) have existed for at least 475&nbsp;million years.<ref>{{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 }}</ref> However, angiosperms [[abominable mystery|appear suddenly]] and in great diversity in the fossil record in the [[Early Cretaceous]] (~130 mya).<ref>{{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 |page=17015 |doi=10.1038/nplants.2017.15 |pmid=28260783 |bibcode=2017NatPl...317015H |s2cid=205458714 |issn=2055-0278}}</ref><ref>{{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|bibcode=2009AmJB...96....5F |url-access=subscription }}</ref> Claimed records of flowering plants prior to this are not widely accepted,<ref>{{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|url-access=subscription }}</ref> as all supposed pre-Cretaceous "flowers" can be explained through being misidentifications of other seed plants. Furthermore, almost all of these controversial fossils are described in papers co-authored by the researcher Xin Wang, such as the particularly debated ''[[Nanjinganthus]]''.<ref>{{cite journal |last1=Sokoloff |first1=Dmitry D. |last2=Remizowa |first2=Margarita V. |last3=El |first3=Elena S. |last4=Rudall |first4=Paula J. |last5=Bateman |first5=Richard M. |title=Supposed Jurassic angiosperms lack pentamery, an important angiosperm-specific feature |journal=New Phytologist |date=October 2020 |volume=228 |issue=2 |pages=420–426 |doi=10.1111/nph.15974|pmid=31418869 |bibcode=2020NewPh.228..420S }}</ref> Molecular evidence suggests that the ancestors of angiosperms diverged from the [[gymnosperms]] during the late [[Devonian]], about 365 million years ago.<ref>{{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|url-access=subscription }}</ref> The origin time of the [[crown group]] of flowering plants remains contentious.<ref>{{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|url-access=subscription }}</ref> By the Late Cretaceous, angiosperms appear to have dominated environments formerly occupied by [[fern]]s and gymnosperms. Large [[Canopy (biology)|canopy]]-forming trees replaced [[conifer]]s as the dominant trees close to the end of the Cretaceous, 66 million years ago.<ref>{{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}}</ref> The radiation of herbaceous angiosperms occurred much later.<ref>{{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 }}</ref>


== Reproduction ==
== Reproduction ==
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[[File:Angiosperm life cycle diagram-en.svg|thumb|upright=2|Angiosperm [[flower]] showing [[Plant reproductive morphology|reproductive parts]] and life cycle]]
[[File:Angiosperm life cycle diagram-en.svg|thumb|upright=2|Angiosperm [[flower]] showing [[Plant reproductive morphology|reproductive parts]] and life cycle]]


The characteristic feature of angiosperms is the flower. Its function is to ensure [[Fertilisation|fertilization]] of the [[ovule]] and development of [[fruit]] containing [[seed]]s.<ref>{{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|url-access=subscription }}</ref> It may arise terminally on a shoot or from the [[axil]] of a leaf.<ref>{{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 }}</ref> 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}}
The characteristic feature of angiosperms is the flower. Its function is to ensure [[Fertilisation|fertilization]] of the [[ovule]] and development of [[fruit]] containing [[seed]]s.<ref>{{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|url-access=subscription }}</ref> It may arise terminally on a shoot or from the [[axil]] of a leaf.<ref>{{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=978-0-12-227620-0 }}</ref> 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. [[Microspore]]s, which divide to become [[pollen|pollen grains]], are the male cells; they are borne in the [[stamen]]s.<ref name="Salisbury-1970">{{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.}}</ref> The female cells, [[megaspore]]s, [[megagametogenesis|divide to become the egg cell]]. They are contained in the [[ovule]] and enclosed in the [[carpel]]; one or more carpels form the [[pistil]].<ref name="Salisbury-1970" />
Flowers produce two kinds of reproductive cells. [[Microspore]]s, which divide to become [[pollen|pollen grains]], are the male cells; they are borne in the [[stamen]]s.<ref name="Salisbury-1970">{{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=1 July 2025 |editor2-last=Parke |editor2-first=Robert V.}}</ref> The female cells, [[megaspore]]s, [[megagametogenesis|divide to become the egg cell]]. They are contained in the [[ovule]] and enclosed in the [[carpel]]; one or more carpels form the [[pistil]].<ref name="Salisbury-1970" />


The flower may consist only of these parts, as in [[Anemophily|wind-pollinated]] plants like the [[willow]], where each flower comprises only a few [[stamen]]s or two carpels.{{sfn|Balfour|Rendle|1911|p=10}} In [[Entomophily|insect-]] or [[Ornithophily|bird-pollinated]] plants, other structures protect the [[sporophyll]]s and attract pollinators. The individual members of these surrounding structures are known as [[sepal]]s and [[petal]]s (or [[tepal]]s 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, [[Floral scent|scent]], and [[nectar]].{{Sfn|De Craene|P.|2010|p=8}}{{Sfn|D. Mauseth|2016|p=226}}
The flower may consist only of these parts, as in [[Anemophily|wind-pollinated]] plants like the [[willow]], where each flower comprises only a few [[stamen]]s or two carpels.{{sfn|Balfour|Rendle|1911|p=10}} In [[Entomophily|insect-]] or [[Ornithophily|bird-pollinated]] plants, other structures protect the [[sporophyll]]s and attract pollinators. The individual members of these surrounding structures are known as [[sepal]]s and [[petal]]s (or [[tepal]]s 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, [[Floral scent|scent]], and [[nectar]].{{Sfn|De Craene|P.|2010|p=8}}{{Sfn|D. Mauseth|2016|p=226}}
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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.<ref>{{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 }}</ref>
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.<ref>{{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 }}</ref>


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.<ref>{{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}}</ref>
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.<ref>{{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 |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}}</ref>


[[Apomixis]], setting seed without fertilization, is found naturally in about 2.2% of angiosperm genera.<ref>{{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 }}</ref> Some angiosperms, including many [[citrus]] varieties, are able to produce fruits through a type of apomixis called [[nucellar embryony]].<ref>{{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}}</ref>
[[Apomixis]], setting seed without fertilization, is found naturally in about 2.2% of angiosperm genera.<ref>{{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 }}</ref> Some angiosperms, including many [[citrus]] varieties, are able to produce fruits through a type of apomixis called [[nucellar embryony]].<ref>{{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}}</ref>
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[[File:Kingfisher-iris.jpg|thumb|upright| [[Bird-and-flower painting]]: ''Kingfisher and iris'' ''kachō-e'' woodblock print by Ohara Koson (late 19th century)]]
[[File:Kingfisher-iris.jpg|thumb|upright| [[Bird-and-flower painting]]: ''Kingfisher and iris'' ''kachō-e'' woodblock print by Ohara Koson (late 19th century)]]


Both real and [[Fictional plants|fictitious plants]] play a wide variety of [[List of fictional plants|roles in literature and film]].<ref>{{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. }}</ref> Flowers are the subjects of many poems by poets such as [[William Blake]], [[Robert Frost]], and [[Rabindranath Tagore]].<ref>{{cite web |title=Flower Poems |url=http://www.poemhunter.com/poems/flower/ |website=Poem Hunter |access-date=21 June 2016}}</ref> [[Bird-and-flower painting]] ({{Transliteration|zh|Huaniaohua}}) is a kind of [[Chinese painting]] that celebrates the beauty of flowering plants.<ref>{{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 }}</ref> Flowers have been [[language of flowers|used in literature to convey meaning]] by authors including [[William Shakespeare]].<ref>{{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}}</ref>  
Both real and [[Fictional plants|fictitious plants]] play a wide variety of [[List of fictional plants|roles in literature and film]].<ref>{{cite journal |title=Literary Plants |journal=Nature Plants |date=2015 |volume=1 |issue=11 |article-number=15181 |doi=10.1038/nplants.2015.181 |pmid=27251545 |doi-access=free |bibcode=2015NatPl...115181. }}</ref> Flowers are the subjects of many poems by poets such as [[William Blake]], [[Robert Frost]], and [[Rabindranath Tagore]].<ref>{{cite web |title=Flower Poems |url=http://www.poemhunter.com/poems/flower/ |website=Poem Hunter |access-date=21 June 2016}}</ref> [[Bird-and-flower painting]] ({{Transliteration|zh|Huaniaohua}}) is a kind of [[Chinese painting]] that celebrates the beauty of flowering plants.<ref>{{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-/ }}</ref> Flowers have been [[language of flowers|used in literature to convey meaning]] by authors including [[William Shakespeare]].<ref>{{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 }}</ref>  
Flowers are used in a variety of art forms which arrange cut or living plants, such as [[bonsai]], [[ikebana]], and flower arranging. [[Ornamental plant]]s have sometimes changed the course of history, as in [[tulip mania|tulipomania]].<ref>{{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}}</ref> Many countries and regions have [[floral emblem]]s; a survey of 70 of these found that the most popular flowering plant family<!--Figure 9 in the source--> 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).<ref>{{cite book |last1=Lim |first1=Reuben |last2=Tan |first2=Heok |last3=Tan |first3=Hugh |year=2013 |title=Official Biological Emblems of the World <!--online--> |publisher=[[Raffles Museum of Biodiversity Research]] |location=Singapore |isbn=978-9-8107-4147-1 |url=https://www.researchgate.net/publication/283008038}}</ref>
Flowers are used in a variety of art forms which arrange cut or living plants, such as [[bonsai]], [[ikebana]], and flower arranging. [[Ornamental plant]]s have sometimes changed the course of history, as in [[tulip mania|tulipomania]].<ref>{{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}}</ref> Many countries and regions have [[floral emblem]]s; a survey of 70 of these found that the most popular flowering plant family<!--Figure 9 in the source--> 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).<ref>{{cite book |last1=Lim |first1=Reuben |last2=Tan |first2=Heok |last3=Tan |first3=Hugh |year=2013 |title=Official Biological Emblems of the World <!--online--> |publisher=[[Raffles Museum of Biodiversity Research]] |location=Singapore |isbn=978-9-8107-4147-1 |url=https://www.researchgate.net/publication/283008038}}</ref>


Line 574: Line 574:
[[File:Viola_calcarata20052002fleur2.JPG|thumb|left|upright=0.9|''[[Viola calcarata]]'', a species highly vulnerable to climate change.<ref>{{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 }}</ref>]]
[[File:Viola_calcarata20052002fleur2.JPG|thumb|left|upright=0.9|''[[Viola calcarata]]'', a species highly vulnerable to climate change.<ref>{{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 }}</ref>]]


[[Human impact on the environment]] has driven a range of species extinct and [[Holocene extinction|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.<ref name="Lughadha-2020">{{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 }}</ref> The majority are threatened by [[Habitat destruction|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.<ref name="BGCI-2023"/><ref>{{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 }}</ref><ref>{{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}}</ref>
[[Human impact on the environment]] has driven a range of species extinct and [[Holocene extinction|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.<ref name="Lughadha-2020">{{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 }}</ref> The majority are threatened by [[Habitat destruction|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.<ref name="BGCI-2023"/><ref>{{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 |article-number=e2001104 |doi=10.1371/journal.pbio.2001104 |doi-access=free |pmid=27930674 |pmc=5147797 |hdl=10150/622757 |hdl-access=free }}</ref><ref>{{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}}</ref>




Relatively few plant diversity assessments currently consider [[climate change]],<ref name="Lughadha-2020" /> yet it is [[Effects of climate change on plant biodiversity|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}}.<ref>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/}}</ref> In worst-case scenarios, half of all tree species may be driven extinct by climate change over that timeframe.<ref name="Lughadha-2020" />
Relatively few plant diversity assessments currently consider [[climate change]],<ref name="Lughadha-2020" /> yet it is [[Effects of climate change on plant biodiversity|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}}.<ref>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/}}</ref> In worst-case scenarios, half of all tree species may be driven extinct by climate change over that timeframe.<ref name="Lughadha-2020" />


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 conservation|ex situ]]'' in [[seed bank]]s or as living plants.<ref name="BGCI-2023">{{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}}</ref> 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."<ref>{{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}}</ref> The [[United Nations]]' [[Global Strategy for Plant Conservation]] asserts that "without plants, there is no life".<ref name="CBD-2023"/> It aims to "halt the continuing loss of plant diversity" throughout the world.<ref name="CBD-2023">{{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}}</ref>{{Clear}}
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 conservation|ex situ]]'' in [[seed bank]]s or as living plants.<ref name="BGCI-2023">{{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}}</ref> 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."<ref>{{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 |archive-date=18 April 2024 |archive-url=https://web.archive.org/web/20240418081054/https://www.botanic.cam.ac.uk/festival-of-plants-2020/fop-day3-conservation/plant-conservation-around-the-world/ |url-status=dead }}</ref> The [[United Nations]]' [[Global Strategy for Plant Conservation]] asserts that "without plants, there is no life".<ref name="CBD-2023"/> It aims to "halt the continuing loss of plant diversity" throughout the world.<ref name="CBD-2023">{{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}}</ref>{{Clear}}


== References ==
== References ==
Line 617: Line 617:
* {{cite journal |last1=Takhtajan |first1=A. |author-link=Takhtajan |title=The Taxa of the Higher Plants above the Rank of Order |journal=[[Taxon (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=The Taxa of the Higher Plants above the Rank of Order |journal=[[Taxon (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=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}}
* {{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 |page=4956 |doi=10.1038/ncomms5956 |pmid=25249442 |pmc=4200517 |bibcode=2014NatCo...5.4956Z |ref=none}}
{{Refend}}
{{Refend}}



Latest revision as of 13:09, 6 November 2025

Template:Short description Script error: No such module "redirect hatnote". Template:Good article Template:Use dmy dates Template:Automatic taxobox

Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (Template:IPAc-en).[1][2] The term angiosperm is derived from the Greek words Script error: No such module "Lang". (Script error: No such module "Lang".; 'container, vessel') and Script error: No such module "Lang". (Script error: No such module "Lang".; 'seed'), meaning that the seeds are enclosed within a fruit. The group was formerly called Magnoliophyta.Template:Sfn

Angiosperms are by far the most diverse group of land plants with 64 orders, 416 families, approximately 13,000 known genera and 300,000 known species.[3] They include all forbs (flowering plants without a woody stem), grasses and grass-like plants, a vast majority of broad-leaved trees, shrubs and vines, and most aquatic plants. Angiosperms are distinguished from the other major seed plant clade, the gymnosperms, by having flowers, xylem consisting of vessel elements instead of tracheids, endosperm within their seeds, and fruits that completely envelop the seeds. The ancestors of flowering plants diverged from the common ancestor of all living gymnosperms before the end of the Carboniferous, over 300 million years ago. In the Cretaceous, angiosperms diversified explosively, becoming the dominant group of plants across the planet.

Agriculture is almost entirely dependent on angiosperms, and a small number of flowering plant families supply nearly all plant-based food and livestock feed. Rice, maize and wheat provide half of the world's staple calorie intake, and all three plants are cereals from the Poaceae family (colloquially known as grasses). Other families provide important industrial plant products such as wood, paper and cotton, and supply numerous ingredients for drinks, sugar production, traditional medicine and modern pharmaceuticals. Flowering plants are also commonly grown for decorative purposes, with certain flowers playing significant cultural roles in many societies.

Out of the "Big Five" extinction events in Earth's history, only the Cretaceous–Paleogene extinction event occurred while angiosperms dominated plant life on the planet. Today, the Holocene extinction affects all kingdoms of complex life on Earth, and conservation measures are necessary to protect plants in their habitats in the wild (in situ), or failing that, ex situ in seed banks or artificial habitats like botanic gardens. Otherwise, around 40% of plant species may become extinct due to human actions such as habitat destruction, introduction of invasive species, unsustainable logging, land clearing and overharvesting of medicinal or ornamental plants. Further, climate change is starting to impact plants and is likely to cause many species to become extinct by 2100.

Distinguishing features

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

Feature Description Image
Flowers The reproductive organs of flowering plants, not found in any other seed plants.[4]
File:Daffodil flower in section, labelled.svg
A Narcissus flower in section. Petals and sepals are replaced here by a fused tube, the corona, and tepals.
Reduced gametophytes, three cells in male, seven cells with eight nuclei in female (except for basal angiosperms)[5] The gametophytes are smaller than those of gymnosperms.[6] The smaller size of the pollen reduces the time between pollination and fertilization, which in gymnosperms is up to a year.[7]
File:Angiosperm embryo sac with female gametophyte.JPG
Embryo sac is a reduced female gametophyte.
Endosperm Endosperm forms after fertilization but before the zygote divides. It provides food for the developing embryo, the cotyledons, and sometimes the seedling.[8]
Closed carpel enclosing the ovules. Once the ovules are fertilised, the carpels, often with surrounding tissues, develop into fruits. Gymnosperms have unenclosed seeds.[9]
File:Alternating peas in peapod (cropped).jpg
Peas (seeds, from ovules) inside pod (fruit, from fertilised carpel).
Xylem made of vessel elements Open vessel elements are stacked end to end to form continuous tubes, whereas gymnosperm xylem is made of tapered tracheids connected by small pits.[10]
File:Herbaceous Dicot Stem Xylem Vessels Cucurbita (35463815631).jpg
Xylem vessels (long tubes).

Diversity

Ecological diversity

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The largest angiosperms are Eucalyptus gum trees of Australia, and Shorea faguetiana, dipterocarp rainforest trees of Southeast Asia, both of which can reach almost Template:Convert in height.[11] The smallest are Wolffia duckweeds which float on freshwater, each plant less than Template:Convert across.[12]

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 (myco-heterotrophic, formerly thought to be saprophytic) like the orchids for part or all of their life-cycle,[13] or on other plants, either wholly like the broomrapes, Orobanche, or partially like the witchweeds, Striga.[14]

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.[15] The seagrasses in the Alismatales grow in marine environments, spreading with rhizomes that grow through the mud in sheltered coastal waters.[16]

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.[17] 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.[18]

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 or two growing seasons,[19] 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.[20]

Taxonomic diversity

The number of species of flowering plants is estimated to be in the range of 250,000 to 400,000.[21][22][23] This compares to around 12,000 species of moss[24] and 11,000 species of pteridophytes.[25] The APG system seeks to determine the number of families, mostly by molecular phylogenetics. In the 2009 APG III there were 415 families.Template:Sfn 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.Template:Sfn

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,[26] containing over 166,000 species between them in their APG circumscriptions, are:

The 25 largest angiosperm families[26]
Group Family English name No. of spp.
1 Eudicot Asteraceae or Compositae daisy 22,750
2 Monocot Orchidaceae orchid 21,950
3 Eudicot Fabaceae or Leguminosae pea, legume 19,400
4 Eudicot Rubiaceae madder 13,150[27]
5 Monocot Poaceae or Gramineae grass 10,035
6 Eudicot Lamiaceae or Labiatae mint 7,175
7 Eudicot Euphorbiaceae spurge 5,735
8 Eudicot Melastomataceae melastome 5,005
9 Eudicot Myrtaceae myrtle 4,625
10 Eudicot Apocynaceae dogbane 4,555
11 Monocot Cyperaceae sedge 4,350
12 Eudicot Malvaceae mallow 4,225
13 Monocot Araceae arum 4,025
14 Eudicot Ericaceae heath 3,995
15 Eudicot Gesneriaceae gesneriad 3,870
16 Eudicot Apiaceae or Umbelliferae parsley 3,780
17 Eudicot Brassicaceae or Cruciferae cabbage 3,710
18 Magnoliid dicot Piperaceae pepper 3,600
19 Monocot Bromeliaceae bromeliad 3,540
20 Eudicot Acanthaceae acanthus 3,500
21 Eudicot Rosaceae rose 2,830
22 Eudicot Boraginaceae borage 2,740
23 Eudicot Urticaceae nettle 2,625
24 Eudicot Ranunculaceae buttercup 2,525
25 Magnoliid dicot Lauraceae laurel 2,500

Evolution

History of classification

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File:Ehret-Methodus Plantarum Sexualis.jpg
From 1736, an illustration of Linnaean classification

The botanical term "angiosperm", from Greek words Script error: No such module "Lang". (Template:Wikt-lang 'bottle, vessel') and Script error: No such module "Lang". (Template:Wikt-lang 'seed'), was coined in the form "Angiospermae" by Paul Hermann in 1690, including only flowering plants whose seeds were enclosed in capsules.Template:Sfn The term angiosperm fundamentally changed in meaning in 1827 with Robert Brown, when angiosperm came to mean a seed plant with enclosed ovules.[28][29] 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.[29]Template:Sfn The APG systemTemplate:Sfn 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.Template:Sfn From 1998, the Angiosperm Phylogeny Group (APG) has reclassified the angiosperms, with updates in the APG II system in 2003,Template:Sfn the APG III system in 2009,Template:Sfn[30] and the APG IV system in 2016.Template:Sfn

Phylogeny

External

In 2019, a molecular phylogeny of plants placed the flowering plants in their evolutionary context:[31]

Template:Clade

Internal

The main groups of living angiosperms are:[32]Template:Sfn

Template:Barlabel

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.[33]

File:Tree of Angiosperm Phylogeny 2024.jpg
Tree of Angiosperm phylogeny 2024

Fossil history

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File:Sagaria cilentana (cropped).jpg
Adaptive radiation 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.[34] However, angiosperms appear suddenly and in great diversity in the fossil record in the Early Cretaceous (~130 mya).[35][36] Claimed records of flowering plants prior to this are not widely accepted,[37] as all supposed pre-Cretaceous "flowers" can be explained through being misidentifications of other seed plants. Furthermore, almost all of these controversial fossils are described in papers co-authored by the researcher Xin Wang, such as the particularly debated Nanjinganthus.[38] Molecular evidence suggests that the ancestors of angiosperms diverged from the gymnosperms during the late Devonian, about 365 million years ago.[39] The origin time of the crown group of flowering plants remains contentious.[40] 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.[41] The radiation of herbaceous angiosperms occurred much later.[42]

Reproduction

Flowers

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File:Angiosperm life cycle diagram-en.svg
Angiosperm flower 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.[43] It may arise terminally on a shoot or from the axil of a leaf.[44] The flower-bearing part of the plant is usually sharply distinguished from the leaf-bearing part, and forms a branch-system called an inflorescence.Template:Sfn

Flowers produce two kinds of reproductive cells. Microspores, which divide to become pollen grains, are the male cells; they are borne in the stamens.[45] 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.[45]

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.Template:Sfn 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.Template:SfnTemplate:Sfn 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.Template:SfnTemplate:Sfn

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.[46] Monoecious plants have separate male and female flowers on the same plant; these are often wind-pollinated,[47] as in maize,[48] but include some insect-pollinated plants such as Cucurbita squashes.[49][50]

Fertilisation and embryogenesis

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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.[51]

Fruit and seed

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

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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.[52]

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.[53]

Apomixis, setting seed without fertilization, is found naturally in about 2.2% of angiosperm genera.[54] Some angiosperms, including many citrus varieties, are able to produce fruits through a type of apomixis called nucellar embryony.[55]

Sexual selection

Template:Excerpt

Adaptive function of flowers

Charles Darwin in his 1878 book The Effects of Cross and Self-Fertilization in the Vegetable Kingdom[56] 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.[57] Meiosis in flowering plants provides a direct mechanism for repairing DNA through genetic recombination in reproductive tissues.[58] 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.[58] 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.[57]

Human uses

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Practical uses

File:Rice Harvest 2020 - 50248478521.jpg
Harvesting rice in Arkansas, 2020
File:Daal after Tadka Pulse Soup India.jpg
Food from plants: a dish of Dal tadka, 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.[59] For instance, half of the world's calorie intake is supplied by just three plants – wheat, rice and maize.[60]

Major food-providing families[59]
Family English Example foods from that family
Poaceae Grasses, cereals Most feedstocks, inc. rice, maize, wheat, barley, rye, oats, pearl millet, sugar cane, sorghum
Fabaceae Legumes, pea family Peas, beans, lentils; for animal feed, clover, alfalfa
Solanaceae Nightshade family Potatoes, tomatoes, peppers, aubergines
Cucurbitaceae Gourd family Squashes, cucumbers, pumpkins, melons
Brassicaceae Cabbage family Cabbage and its varieties, e.g. Brussels sprout, broccoli; mustard; oilseed rape
Apiaceae Parsley family Parsnip, carrot, parsley, coriander, fennel, cumin, caraway
Rutaceae Rue family[61] Oranges, lemons, grapefruits
Rosaceae Rose family[62] Apples, pears, cherries, apricots, plums, peaches

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).[59]

Cultural uses

File:Kingfisher-iris.jpg
Bird-and-flower painting: 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.[63] Flowers are the subjects of many poems by poets such as William Blake, Robert Frost, and Rabindranath Tagore.[64] Bird-and-flower painting (Template:Transliteration) is a kind of Chinese painting that celebrates the beauty of flowering plants.[65] Flowers have been used in literature to convey meaning by authors including William Shakespeare.[66] 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.[67] 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).[68]

Conservation

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File:Viola calcarata20052002fleur2.JPG
Viola calcarata, a species highly vulnerable to climate change.[69]

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.[70] 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.[71][72][73]


Relatively few plant diversity assessments currently consider climate change,[70] 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 Template:Convert of global warming, and 10% at Template:Convert.[74] In worst-case scenarios, half of all tree species may be driven extinct by climate change over that timeframe.[70]

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.[71] 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."[75] The United Nations' Global Strategy for Plant Conservation asserts that "without plants, there is no life".[76] It aims to "halt the continuing loss of plant diversity" throughout the world.[76]

References

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Bibliography

Articles, books and chapters

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  • Cromie, William J. (16 December 1999). "Oldest Known Flowering Plants Identified By Genes". Harvard University Gazette.
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Websites

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Template:Plant classification Template:Botany Template:Angiosperm orders Template:Lists of angiosperm families Template:Life on Earth Template:Subject bar Template:Taxonbar Template:Authority control

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