View source for Biology:Charophyta

{{short description|Phylum of algae}}
{{Paraphyletic group
| image = Chara sp reproductive structure.JPG
| auto = yes
| taxon = Charophyta
| authority = Migula 1897,<ref name=AB_97242/> ''sensu'' Leliaert et al. 2012
| includes =
* [[Biology:Mesostigmatophyceae|Mesostigmatophyceae]]
* Chlorokybophyceae
* ''Streptofilum''
* [[Biology:Klebsormidiophyceae|Klebsormidiophyceae]]
* [[Biology:Phragmoplastophyta|Phragmoplastophyta]]
** [[Biology:Charophyceae|Charophyceae]]
** [[Biology:Coleochaetophyceae|Coleochaetophyceae]]
** [[Biology:Zygnematophyceae|Zygnematophyceae]]
| excludes = [[Biology:Embryophyta|Embryophyta]]
| synonyms = 
}}

'''Charophyta''' ({{IPAc-en|UK|k|ə|ˈ|r|ɒ|f|ᵻ|t|ə|,_|ˌ|k|ær|ə|ˈ|f|aɪ|t|ə}}) is a group of freshwater [[Biology:Green algae|green algae]], called '''charophytes''' ({{IPAc-en|ˈ|k|ær|ə|ˌ|f|aɪ|t|s}}), sometimes treated as a [[Biology:Phylum|division]],<ref name=":1">{{cite journal |last1=Lewis |first1=Louise A. |first2=Richard M. |last2=McCourt |year=2004 |title=Green algae and the origin of land plants |journal=American Journal of Botany |volume=91 |issue=10 |pages=1535–56 |doi=10.3732/ajb.91.10.1535 |pmid=21652308 }}</ref> yet also as a superdivision<ref name="Rugg15">{{cite journal |last1=Ruggiero |first1=M. A. |last2=Gordon |first2=D. P. |last3=Orrell |first3=T. M. |last4=Bailly |first4=N. |last5=Bourgoin |first5=T. |last6=Brusca |first6=R. C. |display-authors=0 |date=2015 |title=A higher level classification of all living organisms |journal=PLOS One |volume=10 | issue = 4 |pages=e0119248 |doi=10.1371/journal.pone.0119248 |pmid=25923521 |pmc=4418965 |bibcode=2015PLoSO..1019248R |doi-access=free }}</ref> or an unranked [[Biology:Clade|clade]]. The terrestrial plants, the [[Biology:Embryophyta|Embryophyta]] emerged deep within Charophyta, possibly from terrestrial unicellular charophytes,<ref>{{cite journal|last1=de Vries|first1=J|last2=Archibald|first2=JM|title=Plant evolution: landmarks on the path to terrestrial life.|journal=The New Phytologist|date=March 2018|volume=217|issue=4|pages=1428–1434|doi=10.1111/nph.14975|pmid=29318635|doi-access=free}}</ref> with the class [[Biology:Zygnematophyceae|Zygnematophyceae]] as a [[Biology:Sister group|sister group]].<ref>{{Cite journal|last=Del-Bem|first=Luiz-Eduardo|date=2018-05-31|title=Xyloglucan evolution and the terrestrialization of green plants|journal=New Phytologist|language=en|volume=219|issue=4|pages=1150–1153|doi=10.1111/nph.15191|pmid=29851097|issn=0028-646X|doi-access=free|hdl=1843/36860|hdl-access=free}}</ref><ref name=":2">{{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|date=2014-02-17|title=From algae to angiosperms–inferring the phylogeny of green plants (Viridiplantae) from 360 plastid genomes|journal=BMC Evolutionary Biology|volume=14|pages=23|doi=10.1186/1471-2148-14-23|issn=1471-2148|pmid=24533922|pmc=3933183 |doi-access=free }}</ref><ref>{{Cite journal|last1=Wickett|first1=Norman J.|last2=Mirarab|first2=Siavash|last3=Nguyen|first3=Nam|last4=Warnow|first4=Tandy|last5=Carpenter|first5=Eric|last6=Matasci|first6=Naim|last7=Ayyampalayam|first7=Saravanaraj|last8=Barker|first8=Michael S.|last9=Burleigh|first9=J. Gordon|date=2014-11-11|title=Phylotranscriptomic analysis of the origin and early diversification of land plants|journal=Proceedings of the National Academy of Sciences|language=en|volume=111|issue=45|pages=E4859–E4868|doi=10.1073/pnas.1323926111|issn=0027-8424|pmc=4234587|pmid=25355905|bibcode=2014PNAS..111E4859W|doi-access=free}}</ref><ref>{{Cite journal|last1=Vries|first1=Jan de|last2=Stanton|first2=Amanda|last3=Archibald|first3=John M.|last4=Gould|first4=Sven B.|date=2016-02-16|title=Streptophyte Terrestrialization in Light of Plastid Evolution|journal=Trends in Plant Science|volume=21|language=en|issue=6|doi=10.1016/j.tplants.2016.01.021|pmid=26895731|issn=1360-1385|pages=467–476}}</ref>

With the Embryophyta now cladistically placed in the Charophyte, it is a synonym of Streptophyta.<ref name=":0">{{Cite book|title=Handbook of the Protists|last1=Cook|first1=Martha E.|last2=Graham|first2=Linda E.|chapter=Chlorokybophyceae, Klebsormidiophyceae, Coleochaetophyceae |date=2017|publisher=Springer International Publishing|isbn=9783319281476|editor-last=Archibald|editor-first=John M.|pages=185–204|language=en|doi=10.1007/978-3-319-28149-0_36|editor-last2=Simpson|editor-first2=Alastair G. B.|editor-last3=Slamovits|editor-first3=Claudio H.}}</ref><ref>{{Cite journal|last1=Delwiche|first1=Charles F.|last2=Timme|first2=Ruth E.|title=Plants|journal=Current Biology|volume=21|issue=11|pages=R417–R422|doi=10.1016/j.cub.2011.04.021|pmid=21640897|year=2011|doi-access=free}}</ref><ref>{{Cite journal|last1=Karol|first1=Kenneth G.|last2=McCourt|first2=Richard M.|last3=Cimino|first3=Matthew T.|last4=Delwiche|first4=Charles F.|date=2001-12-14|title=The Closest Living Relatives of Land Plants|journal=Science|language=en|volume=294|issue=5550|pages=2351–2353|doi=10.1126/science.1065156|issn=0036-8075|pmid=11743201|bibcode=2001Sci...294.2351K|s2cid=35983109}}</ref><ref>{{Cite journal|last1=Lewis|first1=Louise A.|last2=McCourt|first2=Richard M.|date=2004|title=Green algae and the origin of land plants|url=https://onlinelibrary.wiley.com/doi/abs/10.3732/ajb.91.10.1535|journal=American Journal of Botany|language=en|volume=91|issue=10|pages=1535–1556|doi=10.3732/ajb.91.10.1535|pmid=21652308 |issn=1537-2197}}</ref> The sister group of the charophytes are the [[Biology:Chlorophyta|Chlorophyta]]. In some charophyte groups, such as the [[Biology:Zygnematophyceae|Zygnematophyceae]] or conjugating green algae, [[Biology:Flagellum|flagella]] are absent and sexual reproduction does not involve free-swimming flagellate sperm. Flagellate sperm, however, are found in stoneworts ([[Biology:Charales|Charales]]) and [[Biology:Coleochaetales|Coleochaetales]], orders of parenchymatous charophytes that are the closest relatives of the land plants, where flagellate sperm are also present in all except the [[Biology:Pinophyta|conifers]] and [[Biology:Angiosperm|flowering plants]].<ref name=Vaughn>{{cite journal|last1=Vaughn|first1=K.C.|last2=Renzaglia|first2=K.S.|year=2006|title=Structural and immunocytochemical characterization of the ''Ginkgo biloba'' L. sperm motility apparatus|journal=Protoplasma|volume=227|issue=2–4|pages=165–73|doi=10.1007/s00709-005-0141-3|pmid=16736257|s2cid=9864200|url=https://naldc-legacy.nal.usda.gov/naldc/download.xhtml?id=32589&content=PDF}}</ref> Fossil stoneworts of early [[Earth:Devonian|Devonian]] age that are similar to those of the present day have been described from the Rhynie chert of Scotland.<ref name=Kelman2003/> Somewhat different charophytes have also been collected from the Late Devonian (Famennian) Waterloo Farm lagerstätte of South Africa. These include two species each of ''[[Biology:Octochara|Octochara]]'' and ''[[Biology:Hexachara|Hexachara]]'', which are the oldest fossils of Charophyte axes bearing in situ oogonia.

The name comes from the [[Biology:Genus|genus]] ''[[Biology:Chara (alga)|Chara]]'', but the finding that the Embryophyta actually emerged in them has ''not'' resulted in a much more restricted meaning of the Charophyta, namely to a much smaller side branch. This more restricted group corresponds to the [[Biology:Charophyceae|Charophyceae]].

==Description==
The ''[[Biology:Zygnematophyceae|Zygnematophyceae]]'' or, as they used to be called, Conjugatophyceae, generally possess two fairly elaborate chloroplasts in each cell, rather than many discoid ones. They reproduce asexually by the development of a septum between the two cell-halves or semi-cells (in unicellular forms, each daughter-cell develops the other semi-cell afresh) and sexually by conjugation, or the fusion of the entire cell-contents of the two conjugating cells. The saccoderm desmids and the placoderm or true desmids, unicellular or filamentous members of the Zygnematophyceae, are dominant in non-calcareous, acid waters of oligotrophic or primitive lakes (e.g. Wastwater), or in lochans, tarns and bogs, as in the West of Scotland, Eire, parts of Wales and of the Lake District.<ref name=West>{{cite book|last1=West |first1=G.S |last2=Fritsch|first2=F.E.|title=A Treatise of the British Freshwater Algae|date=1927|publisher=Cambridge University Press|location=Cambridge}}</ref>

''[[Biology:Klebsormidium|Klebsormidium]]'', the type of the [[Biology:Klebsormidiophyceae|Klebsormidiophyceae]], is a simple filamentous form with circular, plate-like chloroplasts, reproducing by fragmentation, by dorsiventral, biciliate swarmers and, according to Wille, a twentieth-century algologist, by aplanospores.<ref name=Fritsch>{{cite book|last1=Fritsch|first1=F.E.|title=The Structure and Reproduction of the Algae, vol I|date=1935|publisher=Cambridge University Press|pages=205–206}}</ref> Sexual reproduction is simple and [[Biology:Isogamy|isogamous]] (the male and female gametes are outwardly indistinguishable).<ref name=Fritsch/>

The ''[[Biology:Charales|Charales]]'' (Charophyceae), or stoneworts, are freshwater and [[Earth:Brackish water|brackish]] algae with slender green or grey stems; the grey colour of many species results from the deposition of lime on the walls, masking the green colour of the chlorophyll. The main stems are slender and branch occasionally. Lateral branchlets occur in [[Biology:Whorl (botany)|whorls]] at regular intervals up the stem, they are attached by rhizoids to the substrate.<ref name="Bryant 2007">Bryant 2007, J. The Stoneworts (Chlorophyta, Charales) in Guiry, M.D., John, D.M., Rindi, F. and McCarthy, T.K (Ed) ''New Survey of Clare Island Volume 6: The Freshwater and Terrestrial Algae''.  Royal Irish Academy. {{ISBN|9781904890317}}</ref> The reproductive organs consist of [[Biology:Antheridium|antheridia]] and [[Biology:Oogonium|oogonia]], though the structures of these organs differ considerably from the corresponding organs in other algae. As a result of fertilization, a [[Biology:Protonema|protonema]] is formed, from which the sexually reproducing algae develops.

A new terrestrial genus found in sandy soil in the [[Place:Czech Republic|Czech Republic]], ''Streptofilum'', may belong in its own class due its unique phylogenetic position. A [[Biology:Cell wall|cell wall]] is absent, instead the [[Biology:Cell membrane|cell membrane]] consist of many layers of specific scales. It is a short, filamentous and unbranched algae surrounded by a mucilaginous sheath, which often disintegrates to diads and unicells.<ref name=":3" />

===Reproduction===
The cells in Charophyta algae are all [[Biology:Haploid|haploid]], except during sexual reproduction, where a diploid unicellular zygote is produced. The zygote turns into four new haploid cells through meiosis, which will develop into new algae. In multicellular forms these haploid cells will grow into a [[Biology:Gametophyte|gametophyte]]. In embryophytes (land plants) the zygote will instead give rise to a multicellular [[Biology:Sporophyte|sporophyte]].<ref>[https://gtr.ukri.org/projects?ref=BB%2FM020517%2F1 Evolution and development of land plant embryos - GtR - UKRI]</ref><ref name=Becker2009>{{Cite journal
 | last1 = Becker| first1 = B.
 | last2 = Marin | first2 = B.
 | year = 2009
 | title = Streptophyte algae and the origin of embryophytes
 | journal = Annals of Botany
 | volume = 103
 | pages = 999–1004
 | url= | doi = 10.1093/aob/mcp044
 | pmid = 19273476
 | issue = 7
 | pmc = 2707909
}}</ref>

==Classification==
Charophyta are complex green algae that form a sister group to the [[Biology:Chlorophyta|Chlorophyta]] and within which the [[Biology:Embryophyta|Embryophyta]] emerged. The [[Biology:Chlorophyta|chlorophyte]] and charophyte green algae and the embryophytes or land plants form a clade called the green plants or [[Viridiplantae]], that is united among other things by the absence of [[Biology:Phycobilin|phycobilin]]s, the presence of [[Chemistry:Chlorophyll a|chlorophyll a]] and [[Chemistry:Chlorophyll b|chlorophyll b]], cellulose in the cell wall and the use of [[Chemistry:Starch|starch]], stored in the plastids, as a storage polysaccharide. The charophytes and embryophytes share several traits that distinguish them from the chlorophytes, such as the presence of certain enzymes (class I aldolase, Cu/Zn [[Biology:Superoxide dismutase|superoxide dismutase]], glycolate oxidase, flagellar [[Biology:Peroxidase|peroxidase]]), lateral flagella (when present), and, in many species, the use of [[Biology:Phragmoplast|phragmoplast]]s in [[Biology:Mitosis|mitosis]].<ref name="leliaert">{{cite journal |year=2012 |title=Phylogeny and molecular evolution of the green algae |url=http://images.algaebase.org/pdf/5628E58F0ecc431F0CsJm2B04CAD/49951.pdf |doi=10.1080/07352689.2011.615705 |journal=Critical Reviews in Plant Sciences |volume=31 |pages=1–46 |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 |s2cid=17603352 |access-date=2016-10-04 |archive-date=2015-06-26 |archive-url=https://web.archive.org/web/20150626102452/http://images.algaebase.org/pdf/5628E58F0ecc431F0CsJm2B04CAD/49951.pdf |url-status=dead }}</ref> Thus Charophyta and Embryophyta together form the clade [[Biology:Streptophyta|Streptophyta]], excluding the Chlorophyta.

Charophytes such as ''Palaeonitella cranii'' and possibly the yet unassigned ''[[Biology:Parka decipiens|Parka decipiens]]''<ref name=Hemsley>{{cite journal|last=Hemsley|first=A.R.|title=The ultrastructure of the spores of the Devonian plant ''Parka decipiens''|journal=Annals of Botany|year=1989|volume=64|issue=3|pages=359–367|doi=10.1093/oxfordjournals.aob.a087852}}</ref> are present in the fossil record of the [[Earth:Devonian|Devonian]].<ref name=Kelman2003>{{cite journal|last1=Kelman|first1=R.|last2=Feist|first2=M.|last3=Trewin|first3=N.H.|last4=Hass|first4=H.|title= Charophyte algae from the Rhynie chert|journal=Transactions of the Royal Society of Edinburgh: Earth Sciences|volume=94|issue=4|year=2003|pages=445–455|doi=10.1017/s0263593300000808|s2cid=128869547}}</ref> ''Palaeonitella'' differed little from some present-day stoneworts.

=== Cladogram ===
There is an emerging consensus on green algal relationships, mainly based on molecular data.<ref name="leliaert" /><ref name="Marin20122">{{cite journal|last1=Marin|first1=Birger|year=2012|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|doi=10.1016/j.protis.2011.11.004|pmid=22192529}}</ref><ref name="Laurin-LemayBrinkmann20122">{{cite journal|last1=Laurin-Lemay|first1=Simon|last2=Brinkmann|first2=Henner|last3=Philippe|first3=Hervé|year=2012|title=Origin of land plants revisited in the light of sequence contamination and missing data|journal=Current Biology|volume=22|issue=15|pages=R593–R594|doi=10.1016/j.cub.2012.06.013|pmid=22877776|doi-access=free}}</ref><ref>{{Cite journal|last1=Leliaert|first1=Frederik|last2=Tronholm|first2=Ana|last3=Lemieux|first3=Claude|last4=Turmel|first4=Monique|last5=DePriest|first5=Michael S.|last6=Bhattacharya|first6=Debashish|last7=Karol|first7=Kenneth G.|last8=Fredericq|first8=Suzanne|last9=Zechman|first9=Frederick W.|date=2016-05-09|title=Chloroplast phylogenomic analyses reveal the deepest-branching lineage of the Chlorophyta, Palmophyllophyceae class. nov.|journal=Scientific Reports|language=en|volume=6|pages=25367|doi=10.1038/srep25367|issn=2045-2322|pmc=4860620|pmid=27157793|bibcode=2016NatSR...625367L}}</ref><ref name=":0" /><ref name=":1" /><ref name=":2" /><ref>{{Cite journal|last1=Adl|first1=Sina M.|last2=Simpson|first2=Alastair G. B.|last3=Lane|first3=Christopher E.|last4=Lukeš|first4=Julius|last5=Bass|first5=David|last6=Bowser|first6=Samuel S.|last7=Brown|first7=Matthew W.|last8=Burki|first8=Fabien|last9=Dunthorn|first9=Micah|date=2012-09-01|title=The Revised Classification of Eukaryotes|journal=Journal of Eukaryotic Microbiology|language=en|volume=59|issue=5|pages=429–514|doi=10.1111/j.1550-7408.2012.00644.x|issn=1550-7408|pmid=23020233|pmc=3483872}}</ref><ref>{{Cite journal|last1=Lemieux|first1=Claude|last2=Otis|first2=Christian|last3=Turmel|first3=Monique|date=2007-01-12|title=A clade uniting the green algae Mesostigma viride and Chlorokybus atmophyticus represents the deepest branch of the Streptophyta in chloroplast genome-based phylogenies|journal=BMC Biology|volume=5|pages=2|doi=10.1186/1741-7007-5-2|issn=1741-7007|pmid=17222354|pmc=1781420 |doi-access=free }}</ref><ref name=":4">{{Cite journal|last=Umen|first=James G.|date=2014-11-01|title=Green Algae and the Origins of Multicellularity in the Plant Kingdom|journal=Cold Spring Harbor Perspectives in Biology|language=en|volume=6|issue=11|pages=a016170|doi=10.1101/cshperspect.a016170|issn=1943-0264|pmid=25324214|pmc=4413236}}</ref><ref>{{Cite journal|last1=Sánchez-Baracaldo|first1=Patricia|last2=Raven|first2=John A.|last3=Pisani|first3=Davide|last4=Knoll|first4=Andrew H.|date=2017-09-12|title=Early photosynthetic eukaryotes inhabited low-salinity habitats|journal=Proceedings of the National Academy of Sciences|volume=114|issue=37|pages=E7737–E7745|doi=10.1073/pnas.1620089114|pmid=28808007|pmc=5603991|bibcode=2017PNAS..114E7737S |doi-access=free}}</ref><ref>{{Cite journal|last1=Gitzendanner|first1=Matthew A.|last2=Soltis|first2=Pamela S.|last3=Wong|first3=Gane K.-S.|last4=Ruhfel|first4=Brad R.|last5=Soltis|first5=Douglas E.|date=2018|title=Plastid phylogenomic analysis of green plants: A billion years of evolutionary history|journal=American Journal of Botany|language=en|volume=105|issue=3|pages=291–301|doi=10.1002/ajb2.1048|pmid=29603143|issn=0002-9122|doi-access=free}}</ref><ref name=":3">{{Cite journal|last1=Mikhailyuk|first1=Tatiana|last2=Lukešová|first2=Alena|last3=Glaser|first3=Karin|last4=Holzinger|first4=Andreas|last5=Obwegeser|first5=Sabrina|last6=Nyporko|first6=Svetlana|last7=Friedl|first7=Thomas|last8=Karsten|first8=Ulf|date=2018|title=New Taxa of Streptophyte Algae (Streptophyta) from Terrestrial Habitats Revealed Using an Integrative Approach|journal=Protist|volume=169|issue=3|pages=406–431|doi=10.1016/j.protis.2018.03.002|pmid=29860113|pmc=6071840|issn=1434-4610}}</ref><ref>{{Cite thesis|last=Glass|first=Sarah|date=2021|title=Chloroplast Genome Evolution in the Klebsormidiophyceae and Streptofilum|url=https://academicworks.cuny.edu/le_etds/17|type=MS thesis |publisher=Lehman College}}</ref> The Mesostigmatophyceae (including ''Spirotaenia'', and Chlorokybophyceae) are at the base of charophytes (streptophytes). The cladograms below show consensus phylogenetic relationships based on plastid genomes<ref name=Turmel-Lemieux-2018>{{Citation|last1=Turmel|first1=Monique|title=Evolution of the Plastid Genome in Green Algae|date=2018|work=Advances in Botanical Research|pages=157–193|publisher=Elsevier|doi=10.1016/bs.abr.2017.11.010|isbn=9780128134573|last2=Lemieux|first2=Claude |url=https://www.researchgate.net/publication/321734296}}</ref> and a new proposal for a third phylum of green plants based on analysis of nuclear genomes.<ref name=Li-2020>{{Cite journal|last1=Li|first1=Linzhou|last2=Wang|first2=Sibo|last3=Wang|first3=Hongli|last4=Sahu|first4=Sunil Kumar|last5=Marin|first5=Birger|last6=Li|first6=Haoyuan|last7=Xu|first7=Yan|last8=Liang|first8=Hongping|last9=Li|first9=Zhen|last10=Cheng|first10=Shifeng|last11=Reder|first11=Tanja 
|year=2020 |title=The genome of Prasinoderma coloniale unveils the existence of a third phylum within green plants|journal=Nature Ecology & Evolution|language=en|volume=4|issue=9|pages=1220–1231 |doi=10.1038/s41559-020-1221-7|issn=2397-334X|pmc=7455551|pmid=32572216}}</ref>

{{clade gallery |width=400px |height=275px
|header1=Consensus plastid phylogeny
|footer1=Consensus relationships among major green algal lineages inferred in recent plastid phylogenomic studies<ref name=Turmel-Lemieux-2018/>
|cladogram1=
{{clade |style=font-size:95%;line-height:95%
|label1='''[[Viridiplantae]]'''
|1={{clade
   |1=[[Biology:Chlorophyta|Chlorophyta]]
   |2={{clade
      |label1=[[Biology:Streptophyta|Streptophyta]]
      |sublabel1=(Charophyta ''s.l.'')
      |1={{clade
         |1=[[Biology:Mesostigmatophyceae|Mesostigmatophyceae]] ''s.l.''
         |2={{clade
            |1=[[Biology:Klebsormidiophyceae|Klebsormidiophyceae]]
            |label2x=[[Biology:Phragmoplastophyta|Phragmoplastophyta]]
            |2={{clade
               |1=[[Biology:Charophyceae|Charophyceae]]
               |2={{clade
                  |1=[[Biology:Coleochaetophyceae|Coleochaetophyceae]]
                  |2={{clade
                     |1=[[Biology:Zygnematophyceae|Zygnematophyceae]]
                     |2=[[Biology:Embryophyte|Embryophyte]]s <br/>(land plants)
                     }}
                  }}
               }}
            }}
         }}
      }}
   }}
}}
|header2=Prasinodermophyta hypothesis
|footer2=Relationships among major green algal lineages based on a recent nuclear phylogenomic study<ref name=Li-2020/>
|cladogram2=
{{clade|style=font-size:95%;line-height:95%
|label1='''[[Viridiplantae]]'''/|sublabel1='''[[Biology:Green algae|green algae]]'''
|1={{clade
   |1={{clade
      |label1=[[Biology:Prasinodermophyta|Prasinodermophyta]]
      |1={{clade
         |1=[[Biology:Palmophyllophyceae|Palmophyllophyceae]]
         |2=[[Biology:Prasinoderma|Prasinodermophyceae]]
         }}
      }}
   |2={{clade
      |1=[[Biology:Chlorophyta|Chlorophyta]]
      |2={{clade
         |label1=[[Biology:Streptophyta|Streptophyta]]
         |sublabel1=(Charophyta ''s.l.'')
         |1={{clade
            |1=[[Biology:Mesostigmatophyceae|Mesostigmatophyceae]] ''s.l.''
            |2={{clade
               |1=[[Biology:Klebsormidiophyceae|Klebsormidiophyceae]]
               |label2x=[[Biology:Phragmoplastophyta|Phragmoplastophyta]]
               |2={{clade
                  |1=[[Biology:Charophyceae|Charophyceae]]
                  |2={{clade
                     |1=[[Biology:Coleochaetophyceae|Coleochaetophyceae]]
                     |2={{clade
                        |1=[[Biology:Zygnematophyceae|Zygnematophyceae]]
                        |2=[[Biology:Embryophyte|Embryophyte]]s <br/>(land plants)
                        }}
                     }}
                  }}
               }}
            }}
         }}
      }}
   }}
}}
}}
Mesostigmatophyceae s.l. in the cladograms corresponds to a clade of a narrower circumscription, Mesostigmatophyceae s.s., and a separate class Chlorokybophyceae, as used by [[Biology:AlgaeBase|AlgaeBase]].<ref name=AB_97242>{{AlgaeBase taxon |name=Charophytes |id=97242 |access-date=2022-02-21}}</ref>

The Mesostigmatophyceae are not filamentous, but the other basal charophytes (streptophytes) are.<ref>{{Cite journal|last1=Nishiyama|first1=Tomoaki|last2=Sakayama|first2=Hidetoshi|last3=de Vries|first3=Jan|last4=Buschmann|first4=Henrik|last5=Saint-Marcoux|first5=Denis|last6=Ullrich|first6=Kristian K.|last7=Haas|first7=Fabian B.|last8=Vanderstraeten|first8=Lisa|last9=Becker|first9=Dirk|date=2018|title=The Chara Genome: Secondary Complexity and Implications for Plant Terrestrialization|journal=Cell|volume=174|issue=2|pages=448–464.e24|doi=10.1016/j.cell.2018.06.033|pmid=30007417|issn=0092-8674|doi-access=free}}</ref><ref name=":3" /><ref name=":4" />

==References==
{{reflist}}

==External links==

{{Taxonbar|from=Q133219}}
{{Plant classification}}
{{Life on Earth}}

[[Category:Charophyta| ]]
[[Category:Plant divisions]]

{{Sourceattribution|Charophyta}}