Viridiplantae

{{short description|Clade of archaeplastids including green algae and the land plants}} {{For|an explanation of very similar terms|Plant|Green algae}} {{Automatic taxobox | fossil_range = {{long fossil range|1000|0|earliest=1200|Stenian<ref name="Qing et al">{{cite journal |last=Tang |first=Qing |title=A one-billion-year-old multicellular chlorophyte |journal=Nature Ecology and Evolution |date=24 February 2020 |volume=4 |issue=5 |pages=543–549 |doi=10.1038/s41559-020-1122-9 |pmid=32094536 |doi-access=free |pmc=8668152 |bibcode=2020NatEE...4..543T }}</ref> – Present}} | image = Taiwan 2009 East Coast ShihTiPing Giant Stone Steps Algae FRD 6581.jpg | image_caption = An assortment of thallophyte Viridiplantae in a rock pool, Taiwan | display_parents = 3 | taxon = Viridiplantae | authority = Cavalier-Smith, 1981 | synonyms = * Plantae, Copeland, 1938, 1956<ref>{{cite journal |last=Copeland |first=Herbert F. |year=1938 |title=The kingdoms of organisms |journal=The Quarterly Review of Biology |volume=13 |issue=4 |pages=383–420 |doi=10.1086/394568 |s2cid=84634277 }}</ref><ref>{{cite book |last=Copeland |first=H.F. |year=1956 |title=The Classification of Lower Organisms |location=Palo Alto |publisher=Pacific Books |page=6 |url=https://archive.org/details/classificationof00cope }}</ref> * Euchlorophyta Whittaker, 1969<ref>{{cite journal |last=Whittaker |first=R.H. |title=New concepts of kingdoms or organisms. Evolutionary relations are better represented by new classifications than by the traditional two kingdoms |journal=Science |volume=163 |issue=3863 |pages=150–60 |date=January 1969 |pmid=5762760 |doi=10.1126/science.163.3863.150 |url=https://www.ib.usp.br/inter/0410113/downloads/Whittaker_1969.pdf |citeseerx=10.1.1.403.5430 |access-date=2015-01-31 |archive-url=https://web.archive.org/web/20171117160137/http://www.ib.usp.br/inter/0410113/downloads/Whittaker_1969.pdf |archive-date=2017-11-17 }}</ref> * Chlorophyta ''sensu'' van den Hoek & Jahns, 1978<ref>{{cite book |last1=van den Hoek |first1=C. |last2=Jahns |first2=H.M. |year=1978 |url=https://books.google.com/books?id=tfkmAAAAMAAJ |title=Einführung in die Phykologie |language=de |publisher=Georg Thieme Verlag |location=Stuttgart |isbn=978-3-13-551101-6 }}</ref> * Chlorobionta Jeffrey 1982, emend. Bremer 1985, emend. Lewis and McCourt 2004 * Chlorobiota Kendrick and Crane 1997 * Chloroplastida Adl et al., 2005 * Viridiplantae Cavalier-Smith 1981<ref name=CavalierSmith1981>{{cite journal |last=Cavalier-Smith |first=Tom |author-link=Tom Cavalier-Smith |title=Eukaryote kingdoms: seven or nine? |journal=Bio Systems |volume=14 |issue=3–4 |pages=461–481 |date=1981 |pmid=7337818 |doi=10.1016/0303-2647(81)90050-2 |bibcode=1981BiSys..14..461C }}</ref> * Phyta Barkley 1939 emed. Holt & Uidica 2007 * Cormophyta Endlicher, 1836 * Cormobionta Rothmaler, 1948 * Euplanta Barkley, 1949 * Telomobionta Takhtajan, 1964 * Embryobionta Cronquist et al., 1966 * Metaphyta Whittaker, 1969 | subdivision_ranks = Subgroups | subdivision = * Prasinodermophyta * Chlorophyta * Streptophyta ** Mesostigmatophyceae ** Chlorokybophyceae ** Klebsormidiophyceae ** Charophyceae ** Coleochaetophyceae ** Zygnematophyceae ** Embryophyta }} '''Viridiplantae''' (kingdom Plantae ''sensu stricto''), the green plants, is a natural group or clade of around half a million eukaryotes. They are green because they contain chloroplasts, cell organelles able to produce food by photosynthesis. They are major primary producers of food both in the sea and on land. The group includes both green algae and the land plants (embryophytes) that arose from them.

In 2005, Sina Adl and colleagues proposed the name Chloroplastida for the group. In 2012, Frederik Leliaert and colleagues suggested a revised taxonomy of the Viridiplantae. In 2019, M. Leebens-Mack and colleagues proposed a phylogeny based on analysis of over a thousand plant genomes. It renders the former "chlorophyte algae" and "streptophyte algae" paraphyletic, as the land plants arose from within them.

== Definition ==

'''Viridiplantae''' ({{lit|green plants}})<ref name=CavalierSmith1981/> is a clade of around 450,000–500,000 species of chloroplast-bearing eukaryotes. Most of them are autotrophs that obtain their energy by photosynthesis and play important primary production roles in both terrestrial and aquatic ecosystems.<ref>{{cite journal |last1=Leebens-Mack |first1=J.H. |display-authors=etal |collaboration=One Thousand Plant Transcriptomes Initiative |title=One thousand plant transcriptomes and the phylogenomics of green plants |journal=Nature |volume=574 |issue=7780 |pages=679–685 |date=October 2019 |pmid=31645766 |pmc=6872490 |doi=10.1038/s41586-019-1693-2 }}</ref> The clade includes all green algae, which are primarily aquatic; many are microscopic unicellular phytoplankton. It also includes the macroscopic, multicellular, generally complex-structured land plants (embryophytes, i.e. Plantae ''sensu strictissimo''), which emerged from within the freshwater green algae clade Streptophyta<ref name="pmid19216746">{{cite journal | last1=Cocquyt | first1=Ellen | last2=Verbruggen | first2=Heroen | last3=Leliaert | first3=Frederik | last4=Zechman | first4=Frederick W | last5=Sabbe | first5=Koen | last6=De Clerck | first6=Olivier |title=Gain and loss of elongation factor genes in green algae |journal=BMC Evolutionary Biology |volume=9 |page=39 |date=February 2009 | issue=1 |pmid=19216746 |pmc=2652445 |doi=10.1186/1471-2148-9-39 | bibcode=2009BMCEE...9...39C |doi-access=free }}</ref><ref name="pmid17964920">{{cite book |last=Becker |first=B. |title=Function and Evolution of the Vacuolar Compartment in Green Algae and Land Plants (Viridiplantae) |volume=264 |pages=[https://archive.org/details/internationalrev0000unse_w6f2/page/1 1–24] |year=2007 |pmid=17964920 |doi=10.1016/S0074-7696(07)64001-7 |isbn=978-0-12-374263-6 |series=International Review of Cytology |url-access=registration |url=https://archive.org/details/internationalrev0000unse_w6f2/page/1 }}</ref><ref name="pmid18612431">{{cite journal |last1=Kim |first1=E. |last2=Graham |first2=L.E. |title=EEF2 analysis challenges the monophyly of Archaeplastida and Chromalveolata |journal=PLOS One |volume=3 |issue=7 |article-number=e2621 |date=July 2008 |pmid=18612431 |pmc=2440802 |doi=10.1371/journal.pone.0002621 |bibcode=2008PLoSO...3.2621K |editor1-last=Redfield |editor1-first=Rosemary Jeanne |doi-access=free }}</ref> during the Ordivician.<ref>{{Cite journal |last1=Su |first1=Danyan |last2=Yang |first2=Lingxiao |last3=Shi |first3=Xuan |last4=Ma |first4=Xiaoya |last5=Zhou |first5=Xiaofan |last6=Hedges |first6=S Blair |last7=Zhong |first7=Bojian |date=2021-07-29 |editor-last=Battistuzzi |editor-first=Fabia Ursula |title=Large-Scale Phylogenomic Analyses Reveal the Monophyly of Bryophytes and Neoproterozoic Origin of Land Plants |url=https://academic.oup.com/mbe/article/38/8/3332/6237914 |journal=Molecular Biology and Evolution |volume=38 |issue=8 |pages=3332–3344 |doi=10.1093/molbev/msab106 |pmc=8321542 |pmid=33871608}}</ref><ref name="Becker-2009">{{Citation |last1=Becker |first1=B. |last2=Marin |first2=B. |year=2009 |title=Streptophyte algae and the origin of embryophytes |journal=Annals of Botany |volume=103 |issue=7 |pages=999–1004 |doi=10.1093/aob/mcp044 |name-list-style=amp|pmid=19273476 |pmc=2707909}}</ref>

In traditional taxonomy, the classification of green algae typically exclude the land plants, rendering them a paraphyletic group; however it is cladistically accurate to regard land plants as a specialized clade of green algae that had evolved to thrive on dry land,<ref name="Delwiche-2011"/> thus making Viridiplantae a monophyletic group. Since the realization that the embryophytes emerged from green algae, some authors are starting to include them.<ref name="Delwiche-2011">{{cite journal |last1=Delwiche |first1=C.F. |last2=Timme |first2=R.E. |title=Plants |journal=Current Biology |volume=21 |issue=11 |pages=R417–22 |date=June 2011 |pmid=21640897 |doi=10.1016/j.cub.2011.04.021 |bibcode=2011CBio...21.R417D |doi-access=free }}</ref><ref>{{Cite web|url=http://www.life.umd.edu/labs/delwiche/Charophyte.html |title=Charophycean Green Algae Home Page |website=www.life.umd.edu |access-date=2018-02-24}}</ref><ref name="Ruhfel_2014">{{cite journal |last1=Ruhfel |first1=Brad R. |last2=Gitzendanner |first2=Matthew A. |last3=Soltis |first3=Pamela S. |last4=Soltis |first4=Douglas E. |last5=Burleigh |first5=J. Gordon |title=From algae to angiosperms-inferring the phylogeny of green plants (Viridiplantae) from 360 plastid genomes |journal=BMC Evolutionary Biology |volume=14 |page=23 |date=February 2014 |issue=1 |pmid=24533922 |pmc=3933183 |doi=10.1186/1471-2148-14-23 |bibcode=2014BMCEE..14...23R |doi-access=free }}</ref><ref>{{cite journal |last1=Delwiche |first1=Charles F. |last2=Cooper |first2=E.D. |title=The Evolutionary Origin of a Terrestrial Flora |journal=Current Biology |volume=25 |issue=19 |pages=R899–910 |date=October 2015 |pmid=26439353 |doi=10.1016/j.cub.2015.08.029 |bibcode=2015CBio...25.R899D |doi-access=free }}</ref><ref>{{cite journal |last1=Parfrey |first1=Laura Wegener |last2=Lahr |first2=Daniel J. G. |last3=Knoll |first3=Andrew H. |last4=Katz |first4=Laura A. |title=Estimating the timing of early eukaryotic diversification with multigene molecular clocks |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=108 |issue=33 |pages=13624–9 |date=August 2011 |pmid=21810989 |pmc=3158185 |doi=10.1073/pnas.1110633108 |bibcode=2011PNAS..10813624P |doi-access=free }}</ref>

Viridiplantae species all have cells with cellulose in their cell walls, and primary chloroplasts derived from endosymbiosis with cyanobacteria that contain chlorophylls a and b and lack phycobilins. In some classification systems, the group has been treated as a kingdom<ref name="urlViridiplantae">{{cite web |url=https://www.uniprot.org/taxonomy/33090 |title=Viridiplantae |access-date=2009-03-08}}</ref> under various names such as Viridiplantae, '''Chlorobionta''' or simply the kingdom '''Plantae''' (''sensu stricto''), the lattermost expanding upon the traditional grouping of (land) plants to include all green algae closely and distantly related to Embryophyta. Adl'' et al.'', who produced a classification for all eukaryotes in 2005, introduced the name '''Chloroplastida''' for this group, reflecting the group having primary chloroplasts, and they rejected the name Viridiplantae on the grounds that some of the species are not plants as understood traditionally.<ref name=Adl2005>{{cite journal | last1=Adl | first1=Sina M. | last2=Simpson | first2=Alastair G.B. | last3=Farmer | first3=Mark A. |display-authors=etal |title=The new higher level classification of eukaryotes with emphasis on the taxonomy of protists |journal=The Journal of Eukaryotic Microbiology |volume=52 |issue=5 |pages=399–451 |year=2005 |pmid=16248873 |doi=10.1111/j.1550-7408.2005.00053.x |s2cid=8060916 |doi-access=free }}</ref> Together with Rhodophyta (red algae), Glaucophyta (grey algae) and other basal groups such as the phagotrophic Rhodelphidia<ref>{{Cite journal |last1=Bowles |first1=Alexander M. C. |last2=Williamson |first2=Christopher J. |last3=Williams |first3=Tom A. |last4=Lenton |first4=Timothy M. |last5=Donoghue |first5=Philip C. J. |date=2022-10-31 |title=The origin and early evolution of plants |url=https://www.cell.com/trends/plant-science/abstract/S1360-1385(22)00271-0 |journal=Trends in Plant Science |volume=28 |issue=3 |pages=312–329 |doi=10.1016/j.tplants.2022.09.009 |pmid=36328872|hdl=10871/131900 |s2cid=253303816 |hdl-access=free }}</ref> and the picoplanktonic Picozoa (both considered sister to red algae), Viridiplantae belong to the larger primary algae clade Archaeplastida, which in itself is sometimes described as "Plantae ''sensu lato''".{{cn|date=March 2026}}

== Evolution ==

=== Taxonomy ===

Leliaert ''et al'', 2012 propose the following simplified taxonomy of the Viridiplantae.<ref name="Leliaert_2012">{{cite journal | last1=Leliaert | first1=Frederik | last2=Smith | first2=David R. | last3=Moreau | first3=Hervé | last4=Herron | first4=Matthew D. | last5=Verbruggen | first5=Heroen | last6=Delwiche | first6=Charles F. | last7=De Clerck | first7=Olivier |year=2012 |title=Phylogeny and molecular evolution of the green algae |doi=10.1080/07352689.2011.615705 |journal=Critical Reviews in Plant Sciences |volume=31 |issue=1 |pages=1–46 |bibcode=2012CRvPS..31....1L |s2cid=17603352 |url=https://hal.archives-ouvertes.fr/hal-01590252/file/article.pdf }}</ref> * Viridiplantae ** Chlorophyta *** core chlorophytes **** Ulvophyceae ***** Cladophorales ***** Dasycladales ***** Bryopsidales ***** Trentepohliales ***** Ulvales-Ulotrichales ***** Oltmannsiellopsidales **** Chlorophyceae ***** Oedogoniales ***** Chaetophorales ***** Chaetopeltidales ***** Chlamydomonadales ***** Sphaeropleales **** Trebouxiophyceae ***** Chlorellales ***** Oocystaceae ***** Microthamniales ***** Trebouxiales ***** ''Prasiola'' clade **** Chlorodendrophyceae ***** Chlorodendrales **** Pedinophyceae<ref name="Marin2012">{{cite journal |last=Marin |first=B. |title=Nested in the Chlorellales or independent class? Phylogeny and classification of the Pedinophyceae (Viridiplantae) revealed by molecular phylogenetic analyses of complete nuclear and plastid-encoded rRNA operons |journal=Protist |volume=163 |issue=5 |pages=778–805 |date=September 2012 |pmid=22192529 |doi=10.1016/j.protis.2011.11.004 }}</ref> *** prasinophytes (paraphyletic) **** Pyramimonadales **** Mamiellophyceae **** Pycnococcaceae **** Nephroselmidophyceae **** Prasinococcales **** Palmophyllales ** Streptophyta *** Charophytes **** Mesostigmatophyceae ***** Mesostigmatales ***** Chlorokybales **** Klebsormidiophyceae **** Phragmoplastophyta ***** Charophyceae ***** Coleochaetophyceae ***** Zygnematophyceae ***** Embryophyta (land plants)

=== Phylogeny ===

In 2019, a phylogeny based on genomes and transcriptomes from 1,153 plant species was proposed.<ref name="1000plants">{{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> The placing of algal groups is supported by phylogenies based on genomes from the Mesostigmatophyceae and Chlorokybophyceae that have since been sequenced. Both the "chlorophyte algae" and the "streptophyte algae" are treated as paraphyletic (vertical bars beside phylogenetic tree diagram) in this analysis.<ref name="MvirideGenome1">{{cite journal |last1=Liang |first1=Zhe |display-authors=etal |title=Mesostigma viride Genome and Transcriptome Provide Insights into the Origin and Evolution of Streptophyta |journal=Advanced Science |volume=7 |issue=1 |year=2019 |article-number=1901850 |doi=10.1002/advs.201901850 |pmid=31921561 |pmc=6947507 |doi-access=free }}</ref><ref name="Mv_and_Ca_Genomes">{{cite journal |last1=Wang |first1=Sibo |display-authors=etal |title=Genomes of early-diverging streptophyte algae shed light on plant terrestrialization |journal=Nature Plants |volume=6 |issue=2 |year=2020 |pages=95–106 |doi=10.1038/s41477-019-0560-3 |pmid=31844283 |pmc=7027972 |bibcode=2020NatPl...6...95W |doi-access=free }}</ref> The classification of Bryophyta is supported both by Puttick ''et al.'' 2018,<ref name="Puttick2018">{{cite journal |last1=Puttick |first1=Mark |display-authors=etal |title=The Interrelationships of Land Plants and the Nature of the Ancestral Embryophyte |journal=Current Biology |volume=28 |issue=5 |year=2018 |pages=733–745 |doi=10.1016/j.cub.2018.01.063 |pmid=29456145|bibcode=2018CBio...28E.733P |doi-access=free |hdl=10400.1/11601 |hdl-access=free }}</ref> and by phylogenies involving the hornwort genomes that have also since been sequenced.<ref name="HornwortGenome1">{{cite journal |last1=Zhang |first1=Jian |display-authors=etal |title=The hornwort genome and early land plant evolution |journal=Nature Plants |volume=6 |issue=2 |year=2020 |pages=107–118 |doi=10.1038/s41477-019-0588-4|pmid=32042158 |pmc=7027989 |bibcode=2020NatPl...6..107Z |doi-access=free }}</ref><ref name="HornwortGenome2">{{cite journal |last1=Li |first1=Fay Wei |display-authors=etal |title=Anthoceros genomes illuminate the origin of land plants and the unique biology of hornworts |journal=Nature Plants |volume=6 |issue=3 |year=2020 |pages=259–272 |doi=10.1038/s41477-020-0618-2|pmid=32170292 |pmc=8075897 |bibcode=2020NatPl...6..259L |doi-access=free }}</ref>

{{clade|style=font-size:90%;line-height:80%; |grouplabel1={{clade labels |label1="chlorophyte algae"|top1=15% |label2="streptophyte algae"|top2=40%}} |label1=Archaeplastida |1={{clade |1=Rhodophyta 50px |2={{clade |1=Glaucophyta 50px |label2='''Viridiplantae''' |sublabel2= (green plants) |2={{clade |1=Prasinodermophyta|bar1=darkorange |2={{clade |1=Chlorophyta 50px|bar1=darkgreen |2=  |state2=none |style2=font-size:50%;line-height:50%;  |3={{clade |1= {{clade |1=Mesostigmatophyceae|bar1=blue |2={{clade |1=Chlorokybophyceae|bar1=blue |2=''Spirotaenia'' 30px |bar2=blue }} }} |2={{clade |1=Klebsormidiales 50px |bar1=blue |2={{clade |1=''Chara'' 40px |bar1=blue |2={{clade |1=Coleochaetales|bar1=blue |2={{clade |1=Zygnematophyceae 50px |bar1=blue |label2=Embryophytes |sublabel2= (land plants) |2={{clade |1={{clade |label1=Bryophytes |1={{clade |1=Hornworts 40px |2={{clade |1=Liverworts 50px |2=Mosses 20px }} }} }} |2={{clade |1=Lycophytes 50px |2={{clade |1=Ferns 50px |label2=Spermatophytes |sublabel2= (seed plants) |2={{clade |1=Gymnosperms 40px |2=Angiosperms 50px }} }} }} }} }} }} }} }} }} }} }} }} }} }}

Ancestrally, the green algae were flagellates.<ref name="Leliaert_2012" />

== References == {{Reflist}} {{Wikispecies}}

Category:Biological classification Category:Subkingdoms Category:Taxa named by Thomas Cavalier-Smith