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Charophyceae is a class (biology) of charophyte green algae, and consist of the single order Charales,[2] commonly known as "stoneworts" and "brittleworts".[3] It is a basal Phragmoplastophyta clade as sister of another unnamed clade which contains the Embryophytes (land plants). Charophyceae are a class within the Streptophyta.[4][5][6] Current consensus treats Charophyceae as a class under division Charophyta, with Chlorophyta remaining a distinct division.[7] In 2018, the first nuclear genome sequence from a species belonging to the most basal branch within the Phragmoplastophyta, the Charophyceae, has been published: that of Chara braunii[8]

Chara fragilis
Scientific classification

Smith, 1938


  • Characeae Rabenhorst, 1863[1]

Many of the complex traits related to sexual reproduction, photosynthesis, and other defining characteristics of plants evolved first in Streptophytes; analysis of cpDNA (chloroplast DNA), for instance, reveals that many characteristics of plant chloroplasts evolved first in the Streptophyte genera Staurastrum and Zygnema.[9][10][11]


The thallus is erect with regular nodes and internodes. At each node there is a whorl of branches. The whole plant is calcified and Equisetum-like. The internodes of the main axis consist of a single elongated cell, in Chara the internodes are corticated covering the main axis. In other genera these are absent. Where there is a single row of cortical cells the cortex is referred to as diplostichous, where there are two rows of cortical cells it is termed triplostichous. The intermodal cells elongate and do not divide they become many times longer than broad.[12]


Below is a consensus reconstruction of green algal relationships, mainly based on molecular data.[13][14][15][16][17][18][19][20][21][22]













Embryophytes (land plants)

Green algae


  1. ^ Rabenhorst, L. (1863). Kryptogamen-flora von Sachsen, pp. xiv, 282, 'Characeae'.
  2. ^ Ecological Biochemistry: Environmental and Interspecies Interactions
  3. ^ Charophyceae - an overview | ScienceDirect Topics
  4. ^ Hoek, C. van den, Mann, D. G. & Jahns, H. M. 1995. Algae: An Introduction to Phycology. Cambridge University Press, Cambridge. ISBN 0-521-30419-9
  5. ^ Streptophytina
  6. ^ McCourt, R. M., Chapman, R. L., Buchheim, M. & Mishler, B. D. “Green Plants”. Accessed 13 December 2007
  7. ^ Guiry, M.D. & Guiry, G.M. 2007. AlgaeBase. World-wide electronic publication, National University of Ireland, Galway.; searched on 13 December 2007
  8. ^ Nishiyama, Tomoaki; Sakayama, Hidetoshi; de Vries, Jan; Buschmann, Henrik; Saint-Marcoux, Denis; Ullrich, Kristian K.; Haas, Fabian B.; Vanderstraeten, Lisa; Becker, Dirk; Lang, Daniel; Vosolsobě, Stanislav; Rombauts, Stephane; Wilhelmsson, Per K.I.; Janitza, Philipp; Kern, Ramona; Heyl, Alexander; Rümpler, Florian; Villalobos, Luz Irina A. Calderón; Clay, John M.; Skokan, Roman; Toyoda, Atsushi; Suzuki, Yutaka; Kagoshima, Hiroshi; Schijlen, Elio; Tajeshwar, Navindra; Catarino, Bruno; Hetherington, Alexander J.; Saltykova, Assia; Bonnot, Clemence; Breuninger, Holger; Symeonidi, Aikaterini; Radhakrishnan, Guru V.; Van Nieuwerburgh, Filip; Deforce, Dieter; Chang, Caren; Karol, Kenneth G.; Hedrich, Rainer; Ulvskov, Peter; Glöckner, Gernot; Delwiche, Charles F.; Petrášek, Jan; Van de Peer, Yves; Friml, Jiri; Beilby, Mary; Dolan, Liam; Kohara, Yuji; Sugano, Sumio; Fujiyama, Asao; Delaux, Pierre-Marc; Quint, Marcel; Theißen, Günter; Hagemann, Martin; Harholt, Jesper; Dunand, Christophe; Zachgo, Sabine; Langdale, Jane; Maumus, Florian; Van Der Straeten, Dominique; Gould, Sven B.; Rensing, Stefan A. (July 2018). "The Chara Genome: Secondary Complexity and Implications for Plant Terrestrialization". Cell. 174 (2): 448–464.e24. doi:10.1016/j.cell.2018.06.033. PMID 30007417.
  9. ^ Campbell, N. A. & Reece, J. B. 2005. Biology, Seventh Edition. Benjamin Cummings, San Francisco.
  10. ^ Turmel, M., Otis, C. & Lemieux, C. 2005. The Complete Chloroplast DNA Sequences of the Charophycean Green Algae Staurastrum and Zygnema Reveal that the Chloroplast Genome Underwent Extensive Changes During the Evolution of the Zygnematales. BMC Biology 3:22
  11. ^ de Vries, Jan; Stanton, Amanda; Archibald, John M.; Gould, Sven B. (June 2016). "Streptophyte Terrestrialization in Light of Plastid Evolution". Trends in Plant Science. 21 (6): 467–476. doi:10.1016/j.tplants.2016.01.021. PMID 26895731.
  12. ^ Smith, G.M. Cryptogamic Botany. Second Edition. 1955.p 121-130. McGraw-Hill Book Company.INC.
  13. ^ Leliaert, Frederik; Smith, David R.; Moreau, Hervé; Herron, Matthew D.; Verbruggen, Heroen; Delwiche, Charles F.; De Clerck, Olivier (2012). "Phylogeny and Molecular Evolution of the Green Algae" (PDF). Critical Reviews in Plant Sciences. 31: 1–46. doi:10.1080/07352689.2011.615705.
  14. ^ Marin, Birger (2012). "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". Protist. 163 (5): 778–805. doi:10.1016/j.protis.2011.11.004. PMID 22192529.
  15. ^ Laurin-Lemay, Simon; Brinkmann, Henner; Philippe, Hervé (2012). "Origin of land plants revisited in the light of sequence contamination and missing data". Current Biology. 22 (15): R593–R594. Bibcode:1996CBio....6.1213A. doi:10.1016/j.cub.2012.06.013. PMID 22877776.
  16. ^ Leliaert, Frederik; Tronholm, Ana; Lemieux, Claude; Turmel, Monique; DePriest, Michael S.; Bhattacharya, Debashish; Karol, Kenneth G.; Fredericq, Suzanne; Zechman, Frederick W. (2016-05-09). "Chloroplast phylogenomic analyses reveal the deepest-branching lineage of the Chlorophyta, Palmophyllophyceae class. nov". Scientific Reports. 6: 25367. Bibcode:2016NatSR...625367L. doi:10.1038/srep25367. ISSN 2045-2322. PMC 4860620. PMID 27157793.
  17. ^ Cook, Martha E.; Graham, Linda E. (2017). Archibald, John M.; Simpson, Alastair G. B.; Slamovits, Claudio H. (eds.). Handbook of the Protists. Springer International Publishing. pp. 185–204. doi:10.1007/978-3-319-28149-0_36. ISBN 9783319281476.
  18. ^ Lewis, Louise A.; Richard M. McCourt (2004). "Green algae and the origin of land plants" (abstract). American Journal of Botany. 91 (10): 1535–1556. doi:10.3732/ajb.91.10.1535. PMID 21652308.
  19. ^ Ruhfel, Brad R.; Gitzendanner, Matthew A.; Soltis, Pamela S.; Soltis, Douglas E.; Burleigh, J. Gordon (2014-02-17). "From algae to angiosperms–inferring the phylogeny of green plants (Viridiplantae) from 360 plastid genomes". BMC Evolutionary Biology. 14: 23. doi:10.1186/1471-2148-14-23. ISSN 1471-2148. PMC 3933183. PMID 24533922.
  20. ^ Adl, Sina M.; Simpson, Alastair G. B.; Lane, Christopher E.; Lukeš, Julius; Bass, David; Bowser, Samuel S.; Brown, Matthew W.; Burki, Fabien; Dunthorn, Micah (2012-09-01). "The Revised Classification of Eukaryotes". Journal of Eukaryotic Microbiology. 59 (5): 429–514. doi:10.1111/j.1550-7408.2012.00644.x. ISSN 1550-7408. PMC 3483872. PMID 23020233.
  21. ^ Sánchez-Baracaldo, Patricia; Raven, John A.; Pisani, Davide; Knoll, Andrew H. (2017-09-12). "Early photosynthetic eukaryotes inhabited low-salinity habitats". Proceedings of the National Academy of Sciences. 114 (37): E7737–E7745. doi:10.1073/pnas.1620089114. PMC 5603991. PMID 28808007.
  22. ^ Gitzendanner, Matthew A.; Soltis, Pamela S.; Wong, Gane K.-S.; Ruhfel, Brad R.; Soltis, Douglas E. (2018). "Plastid phylogenomic analysis of green plants: A billion years of evolutionary history". American Journal of Botany. 105 (3): 291–301. doi:10.1002/ajb2.1048. ISSN 0002-9122. PMID 29603143.