Editing Flowering plant (section)

== Evolution ==

=== History of classification ===
{{main|Plant taxonomy}}

[[File:Ehret-Methodus Plantarum Sexualis.jpg|thumb|upright|From 1736, an illustration of Linnaean classification]]

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 (botanist)|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 (Scottish botanist from Montrose)|Robert Brown]], when angiosperm came to mean a seed plant with enclosed ovules.<ref>{{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 }}</ref><ref name="Buggs-2021">{{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 }}</ref> In 1851, with [[Wilhelm Friedrich Benedikt Hofmeister|Wilhelm Hofmeister]]'s work on embryo-sacs, Angiosperm came to have its modern meaning of all the flowering plants including Dicotyledons and Monocotyledons.<ref name="Buggs-2021"/>{{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}}<ref>{{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}}</ref> and the [[APG IV system]] in 2016.{{sfn|APG|2016}}

=== Phylogeny ===

==== External ====

In 2019, a [[Molecular phylogenetics|molecular phylogeny]] of [[plant]]s placed the flowering plants in their evolutionary context:<ref>{{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 }}</ref>

{{clade
|label1= [[Embryophyte]]s 
|sublabel1= land plants 
|1={{clade
   |1={{clade
      |1= [[Bryophyte]]s  [[File:WWB-0265-127-Polytrichum formosum.png|20px]]
      |label2= [[Tracheophyte]]s
      |sublabel2= vascular plants 
      |2={{clade
         |1=[[Lycophyte]]s [[File:Lycopodium clavatum - Köhler–s Medizinal-Pflanzen-219 (extracted).jpg|50px]]
         |2={{clade
            |1=[[Fern]]s [[File:The ferns of Great Britain, and their allies the club-mosses, pepperworts, and horsetails (Pl. 12) (8516512808).jpg|40px]]
            |label2= [[Spermatophyte]]s 
            |sublabel2= seed plants 
            |2={{clade
               |label1= [[Gymnosperm]]s  
               |sublabel1= conifers and allies 
               |1= [[File:Pinus montesumae tab22 2.jpg|50px]]
               |label2= '''Angiosperms''' 
               |sublabel2= '''flowering plants''' 
               |2= [[File:160 Ranunculus repens.jpg|50px]]
               }}
            }}
         }}
      }}
   }}
}}

==== 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}}

{{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]] [[File:Amborella trichopoda.jpg|70px]] 1 sp. [[New Caledonia]] shrub |barbegin1=red
   |2={{clade
      |1=[[Nymphaeales]] [[File:2007 nymphaea lotus.jpg|70px]] c. 80 spp.<ref name="Palmer-2004">{{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 }}</ref> [[Nymphaeaceae|water lilies]] & allies  |bar1=red
      |2={{clade
         |1=[[Austrobaileyales]] [[File:Schisandra rubriflora.jpg|70px]] c. 100 spp.<ref name="Palmer-2004"/> woody plants  |barend1=red
         |2={{clade
            |1={{clade
               |1=[[Magnoliids]] [[File:Magnolia obovata 02.jpg|70px]] c. 10,000 spp.<ref name="Palmer-2004"/> [[merosity|3-part]] flowers, 1-pore pollen, usu. branch-veined leaves |barbegin1=green
               |2=[[Chloranthales]] [[File:Sarcandra_glabra2.jpg|70px]] 77 spp.<ref>{{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 }}</ref> Woody, apetalous |bar2=green
               }}
            |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
               |2={{clade
                  |1=[[Ceratophyllales]] [[File:CeratophyllumSubmersum.jpg|70px]] c. 6 spp.<ref name="Palmer-2004"/> [[aquatic plant]]s  |bar1=green
                  |2=[[Eudicots]] [[File:Senecio_angulatus_003.jpg|70px]] c. 175,000 spp.<ref name="Palmer-2004"/> 4- or 5-part flowers, 3-pore pollen, usu. branch-veined leaves |barend2=green
                  }}
               }}
            }}
         }}
      }}
   }}
}}
}}

<!------------------- collapsible element containing large cladogram of APG IV phylogeny --------------->
{|class="collapsible collapsed" style="width:100%; border:solid 1px #aaa"
|-
! style="background:#F0F2F5" |Detailed [[cladogram]] of the 2016 [[Angiosperm Phylogeny Group]] (APG) IV classification.{{sfn|APG|2016}}
|-
|
{{clade|style=font-size:80%;line-height:80%
|label1='''Angiosperms'''
|1={{clade
   |1=[[Amborellales]] <small>Melikyan, Bobrov & Zaytzeva 1999</small>
   |2={{clade
      |1=[[Nymphaeales]] <small>Salisbury ex von Berchtold & Presl 1820</small>
      |2={{clade
         |1=[[Austrobaileyales]] <small>Takhtajan ex Reveal 1992</small>
         |label2=[[Mesangiosperms]]
         |2={{clade
            |1=[[Chloranthales]] <small>Mart. 1835</small>
            |2={{clade
               |label1=[[Magnoliids]] <!--insert dummy clade for spacing-->
               |1={{clade
                  |1={{clade
                     |1=[[Canellales]] <small>Cronquist 1957</small>
                     |2=[[Piperales]] <small>von Berchtold & Presl 1820</small>
                     }}
                  |2={{clade
                     |1=[[Magnoliales]] <small>de Jussieu ex von Berchtold & Presl 1820</small>
                     |2=[[Laurales]] <small>de Jussieu ex von Berchtold & Presl 1820</small>
                     }}
                  }}
               }}
            |3={{clade
               |1={{clade 
                  |label1=[[Monocots]] <!--insert dummy clade-->
                  |1={{clade
                     |1=[[Acorales]] <small>Link 1835</small>
                     |2={{clade
                        |1=[[Alismatales]] <small>Brown ex von Berchtold & Presl 1820</small>
                        |2={{clade
                           |1=[[Petrosaviales]] <small>Takhtajan 1997</small>
                           |2={{clade
                              |1={{clade
                                 |1=[[Dioscoreales]] <small>Brown 1835</small>
                                 |2=[[Pandanales]] <small>Brown ex von Berchtold & Presl 1820</small>
                                 }}
                              |2={{clade
                                 |1=[[Liliales]] <small>Perleb 1826</small>
                                 |2={{clade
                                    |1=[[Asparagales]] <small>Link 1829</small>
                                    |label2=[[Commelinids]]
                                    |2={{clade
                                       |1=[[Arecales]] <small>Bromhead 1840</small>
                                       |2={{clade
                                          |1=[[Poales]] <small>Small 1903</small>
                                          |2={{clade
                                             |1=[[Zingiberales]] <small>Grisebach 1854</small>
                                             |2=[[Commelinales]] <small>de Mirbel ex von Berchtold & Presl 1820</small>
                                             }}
                                          }}
                                       }}
                                    }}
                                 }}
                              }}
                           }}
                        }}
                     }}
                  }}
               |2={{clade
                  |1=[[Ceratophyllales]] <small>Link 1829</small>
                  |label2=[[Eudicots]]
                  |2=EUDICOTS <!--target keyword that will be substituted by Eudicot subtree (see below)-->
                  }}
               }}
            }}
         }}
      }}
   }}

|targetA=EUDICOTS
|subcladeA=   {{clade
              |1=[[Ranunculales]] <small>de Jussieu ex von Berchtold & Presl 1820</small>
              |2={{clade
                 |1=[[Proteales]] <small>de Jussieu ex von Berchtold & Presl 1820</small>
                 |2={{clade
                    |1=[[Trochodendrales]] <small>Takhtajan ex Cronquist 1981</small>
                    |2={{clade
                       |1=[[Buxales]] <small>Takhtajan ex Reveal 1996</small>
                       |label2=[[Core eudicots]]
                       |2={{clade
                          |1=[[Gunnerales]] <small>Takhtajan ex Reveal 1992</small>
                          |2={{clade
                             |1=[[Dilleniales]] <small>de Candolle ex von Berchtold & Presl 1820</small>
                             |label2=[[Superrosids]]
                             |2=SUPERROSIDS
                             |label3=[[Superasterids]]
                             |3=SUPERASTERIDS
                             }}
                          }}
                       }}
                    }}
                 }}
              }}

|targetB=SUPERROSIDS
|subcladeB=   {{clade
              |1=[[Saxifragales]] <small>von Berchtold & Presl 1820</small>
              |label2=[[Rosids]]
              |2={{clade
                 |1=[[Vitales]] <small>de Jussieu ex von Berchtold & Presl 1820</small>
                 |2={{clade
                    |label1=Fabids |sublabel1=(eurosids I)
                    |1={{clade
                       |1=[[Zygophyllales]] <small>Link 1829</small>
                       |2={{clade
                          |1={{clade
                             |1=[[Celastrales]] <small>Link 1829</small>
                             |2={{clade
                                |1=[[Oxalidales]] <small>von Berchtold & Presl 1820</small>
                                |2=[[Malpighiales]] <small>de Jussieu ex von Berchtold & Presl 1820</small>
                                }}
                             }}
                          |2={{clade
                             |1=[[Fabales]] <small>Bromhead 1838</small>
                             |2={{clade
                                |1=[[Rosales]] <small>von Berchtold & Presl 1820</small>
                                |2={{clade
                                   |1=[[Cucurbitales]] <small>de Jussieu ex von Berchtold & Presl 1820</small>
                                   |2=[[Fagales]] <small>Engler 1892</small>
                                   }}
                                }}
                             }}
                          }}
                       }}
                    |label2=Malvids |sublabel2=(eurosids II)
                    |2={{clade
                       |1={{clade
                          |1=[[Geraniales]] <small>de Jussieu ex von Berchtold & Presl 1820</small>
                          |2=[[Myrtales]] <small>de Jussieu ex von Berchtold & Presl 1820</small>
                          }}
                       |2={{clade
                          |1=[[Crossosomatales]] <small>Takhtajan ex Reveal 1993</small>
                          |2={{clade
                             |1=[[Picramniales]] <small>Doweld 2001</small>
                             |2={{clade
                                |1=[[Sapindales]] <small>de Jussieu ex von Berchtold & Presl 1820</small>
                                |2={{clade
                                   |1=[[Huerteales]] <small>Doweld 2001</small>
                                   |2={{clade
                                      |1=[[Malvales]] <small>de Jussieu ex von Berchtold & Presl 1820</small>
                                      |2=[[Brassicales]] <small>Bromhead 1838</small>
                                      }}
                                   }}
                                }}
                             }}
                          }}
                       }}
                    }}
                 }}
              }}

|targetC=SUPERASTERIDS
|subcladeC=   {{clade
              |1=[[Berberidopsidales]] <small>Doweld 2001</small>
              |2={{clade
                 |1=[[Santalales]] <small>Brown ex von Berchtold & Presl 1820</small>
                 |2={{clade
                    |1=[[Caryophyllales]]
                    |label2=[[Asterids]]
                    |2={{clade
                       |1=[[Cornales]] <small>Link 1829</small>
                       |2={{clade
                          |1=[[Ericales]] <small>von Berchtold & Presl 1820</small>
                          |2={{clade
                             |label1=Lamiids |sublabel1=(euasterids I)
                             |1={{clade
                                |1=[[Icacinales]] <small>Van Tieghem 1900</small>
                                |2={{clade
                                   |1=[[Metteniusales]] <small>Takhtajan 1997</small>
                                   |2={{clade
                                      |1=[[Garryales]] <small>Mart. 1835</small>
                                      |2={{clade
                                         |1=[[Gentianales]] <small>de Jussieu ex von Berchtold & Presl 1820</small>
                                         |2=[[Solanales]] <small>de Jussieu ex von Berchtold & Presl 1820</small>
                                         |3=[[Boraginales]] <small>de Jussieu ex von Berchtold & Presl 1820</small>
                                         |4=[[Vahliales]] <small>Doweld 2001</small>
                                         |5=[[Lamiales]] <small>Bromhead 1838</small>
                                         }}
                                      }}
                                   }}
                                }}
                             |label2=Campanulids |sublabel2=(euasterids II)
                             |2={{clade
                                |1=[[Aquifoliales]] <small>Senft 1856</small>
                                |2={{clade
                                   |1=[[Escalloniales]] <small>Mart. 1835</small>
                                   |2=[[Asterales]] <small>Link 1829</small>
                                   |3={{clade
                                      |1=[[Bruniales]] <small>Dumortier 1829</small>
                                      |2={{clade
                                         |1=[[Apiales]] <small>Nakai 1930</small>
                                         |2={{clade
                                            |1=[[Paracryphiales]] <small>Takhtajan ex Reveal 1992</small>
                                            |2=[[Dipsacales]] <small>de Jussieu ex von Berchtold & Presl 1820</small>
                                            }}
                                         }}
                                      }}
                                   }}
                                }}
                             }}
                          }}
                       }}
                    }}
                 }}
              }}
}}
|}

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 [[Rosidae|rosid]] phylogeny is revised.<ref name="Zuntini-2024">{{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 <!-- 279 authors--> |title=Phylogenomics and the rise of the angiosperms |journal=[[Nature (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 }}<!--images are CC-by-SA--></ref>

[[File:Tree of Angiosperm Phylogeny 2024.jpg|thumb|center|upright=3|Tree of Angiosperm phylogeny 2024]]

=== Fossil history ===
{{main|Fossil history of flowering plants}}

[[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}}</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}}</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}}</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>