Chlorophyta

Chlorophyta or Prasinophyta is a taxon of green algae informally called chlorophytes.[6] The name is used in two very different senses, so care is needed to determine the use by a particular author. In older classification systems, it refers to a highly paraphyletic group of all the green algae within the green plants (Viridiplantae) and thus includes about 7,000 species[7][8] of mostly aquatic photosynthetic eukaryotic organisms. In newer classifications, it refers to the sister of the streptophytes/charophytes. The clade Streptophyta consists of the Charophyta in which the Embryophyta emerged.[9][10] In this sense the Chlorophyta includes only about 4,300 species.[4] About 90% of all known species live in freshwater.[11] Like the land plants (bryophytes and tracheophytes), green algae contain chlorophyll a and chlorophyll b and store food as starch[7] in their plastids.

This article is about a group of green algae. For the genus with the common name 'spider plant', see Chlorophytum.

Chlorophyta
Chlorophytes (A–F, H–L and O)
Scientific classification
Domain: Eukaryota
(unranked): Diaphoretickes
(unranked): Archaeplastida
Clade: Chloroplastida
Phylum: Chlorophyta
Reichenbach, 1828, emend. Pascher, 1914, emend. Lewis & McCourt, 2004[1][2][3]
Classes[4]
Synonyms[5]
  • Chlorophycophyta Papenfuss 1946
  • Chlorophycota
  • Chlorophytina
  • Chlorophyllophyceae
  • Isokontae
  • Stephanokontae
Green algae on coastal rocks at Shihtiping in Taiwan

With the exception of Palmophyllophyceae, Trebouxiophyceae, Ulvophyceae and Chlorophyceae, which show various degrees of multicellularity, all the Chlorophyta lineages are unicellular.[12] Some members of the group form symbiotic relationships with protozoa, sponges, and cnidarians. Others form symbiotic relationships with fungi to form lichens, but the majority of species are free-living. Some conduct sexual reproduction, which is oogamous or isogamous. All members of the clade have motile flagellated swimming cells.[13] While most species live in freshwater habitats and a large number in marine habitats, other species are adapted to a wide range of land environments. For example, Chlamydomonas nivalis, which causes Watermelon snow, lives on summer alpine snowfields. Others, such as Trentepohlia species, live attached to rocks or woody parts of trees. Monostroma kuroshiense, an edible green alga cultivated worldwide and most expensive among green algae, belongs to this group.

Ecology

Species of Chlorophyta (treated as what is now considered one of the two main clades of Viridiplantae) are common inhabitants of marine, freshwater and terrestrial environments.[14][15] Several species have adapted to specialised and extreme environments, such as deserts, arctic environments, hypersaline habitats, marine deep waters, deep-sea hydrothermal vents and habitats that experiences extreme changes in temperature, light and salinity.[16][17][18][19] Some groups, such as the Trentepohliales are exclusively found on land.[20] Several species of Chlorophyta live in symbiosis with a diverse range of eukaryotes, including fungi (to form lichens), ciliates, forams, cnidarians and molluscs. [15] Some species of Chlorophyta are heterotrophic, either free-living or parasitic.[21][22] Two common species of the heterotrophic green alga Prototheca are pathogenic and can cause the disease protothecosis in humans and animals.[23]

Classifications
"Siphoneae" from Ernst Haeckel's Kunstformen der Natur, 1904
Further information: Wikispecies:Chlorophyta

Characteristics used for the classification of Chlorophyta are: type of zoid, mitosis (karyokynesis), cytokinesis, organization level, life cycle, type of gametes, cell wall polysaccharides[24] and more recently genetic data.

Phylogeny

A newer proposed classification follows Leliaert et al. 2011[18] and modified with Silar 2016,[25] Leliaert 2016[26] and Lopes dos Santos et al. 2017[27] for the green algae clades and Novíkov & Barabaš-Krasni 2015[28] for the land plants clade. Sánchez-Baracaldo et al. is followed for the basal clades.[29][30][31]

Viridiplantae/
 
 

Mesostigmatophyceae

 
 
 

Spirotaenia

 
 

Chlorokybophyceae

 
 
 
 
Chlorophyta/
Palmophyllophyceae
 

Prasinococcales ("prasinophyte clade VI")

 
 

Palmophyllales

 
 
 
 
 

Pyramimonadophyceae ("prasinophyte clade I")

 
 

Mamiellophyceae ("prasinophyte clade II")

 
 
 
 

Nephroselmidophyceae ("prasinophyte clade III")

 
 
 

?Pycnococcaceae ("prasinophyte clade V")

 
 

Picocystophyceae ("prasinophyte clade VII C")

 
 
 

Chloropicophyceae ("prasinophyte clade VII A/B")

 
Tetraphytina
 

Pedinophyceae

 
Chlorophytina
 

Chlorodendrophyceae ("prasinophyte clade IV")

 
 
 

Trebouxiophyceae

 
 
 

Ulvophyceae

 
 

Chlorophyceae

 
 
 
 
 
 
 
 
 
Prasinophyta
Streptophyta/
 

Klebsormidiophyta

 
Phragmoplastophyta
 

Charophyta (stoneworts)

 
 
 

?Chaetosphaeridiales

 
 

Coleochaetophyta

 
 
 
 

Zygnematophyta

 
 

Mesotaeniaceae

 
 
 

Embryophyte (Land plants)

 
 
 
 
Charophyta
 
Green Algae

A 2020 paper places the "Prasinodermophyta" (i.e. Prasinodermophyceae + Palmophyllophyceae) as the basal Viridiplantae clade.[32]

Viridiplantae/
Prasinodermophyta
 

Prasinodermophyceae

 
Palmophyllophyceae
 

Prasinococcales ("prasinophyte clade VI")

 
 

Palmophyllales

 
 
 
 
Chlorophyta
 
 

Pyramimonadophyceae ("prasinophyte clade I")

 
 

Mamiellophyceae ("prasinophyte clade II")

 
 
 
 

Nephroselmidophyceae ("prasinophyte clade III")

 
 
 

Pycnococcaceae ("prasinophyte clade V")

 
 
 

Chloropicophyceae ("prasinophyte clade VII A/B/C")

 
Tetraphytina
 

Pedinophyceae

 
 
 

Trebouxiophyceae

 
 
 

Chlorodendrophyceae ("prasinophyte clade IV")

 
 
 

Ulvophyceae

 
 

Chlorophyceae

 
 
 
 
 
 
 
 
 
Streptophyta/
Mesostigmatophyceae
 

Mesostigma viride

 
 
 

Spirotaenia

 
 

Chlorokybophyceae

 
 
 
 
 

Klebsormidiophyta

 
Phragmoplastophyta
 

Charophyceae (stoneworts)

 
 
Coleochaetophyceae
 

Chaetosphaeridiales

 
 

Coleochaetophyta

 
 
 
Zygnematophyceae
 

Zygnematophyta

 
 

Mesotaeniaceae

 
 
 

Embryophyte (Land plants)

 
 
 
 
 
Charophyta
 
Green Algae/Prasinophyta s.l.

Leliaert et al. 2012

Simplified phylogeny of the Chlorophyta, according to Leliaert et al. 2012.[15] Note that many algae previously classified in Chlorophyta are placed here in Streptophyta.

Pombert et al. 2005

A possible classification when Chlorophyta refers to one of the two clades of the Viridiplantae is shown below.[33]

Lewis & McCourt 2004

Hoek, Mann and Jahns 1995

Classification of the Chlorophyta, treated as all green algae, according to Hoek, Mann and Jahns 1995.[7]

In a note added in proof, an alternative classification is presented for the algae of the class Chlorophyceae:

Bold and Wynne 1985

Classification of the Chlorophyta and Charophyta according to Bold and Wynne 1985.[35]

Mattox & Stewart 1984

Classification of the Chlorophyta according to Mattox & Stewart 1984:[34]

Fott 1971

Classification of the Chlorophyta according to Fott 1971.[7]:483

  • Suborder Chlorosarcineae
  • Suborder Ulotrichineae
  • Suborder Oedogoniineae
  • Suborder Chaetophorineae

Round 1971

Classification of the Chlorophyta and related algae according to Round 1971.[36]

Smith 1938

Classification of the Chlorophyta according to Smith 1938:

  • Order 5. Schizogoniales
  • Family Schizogoniaceae
  • Order 10. Siphonales
  • Order 11. Siphonocladiales

Research and Discoveries

In February 2020, the fossilized remains of green algae, named Proterocladus antiquus were discovered in the northern province of Liaoning, China.[37] At around a billion years old, it is believed to be one of the oldest examples of a multicellular chlorophyte.[38]

References
  1. Reichenbach HG (1828). Conspectus Regni Vegetabilis. p. 23.
  2. Pascher A (1914). "Über Flagellaten und Algen". Berichte der Deutschen Botanischen Gesellschaft. 32: 136–160.
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  4. Guiry MD, Guiry GM (2011). "AlgaeBase : Chlorophyta". World-wide electronic publication, National University of Ireland, Galway. Retrieved 2011-07-26. Cite journal requires |journal= (help)
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  11. Chlorophyta (Chapter 5) - Phycology - Cambridge University Press
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  14. Graham LE, Graham JM, Wilcox LW (2009) Algae. 2nd Edition. Benjamin Cummings (Pearson), San Francisco, CA
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  22. Nedelcu AM (December 2001). "Complex patterns of plastid 16S rRNA gene evolution in nonphotosynthetic green algae". Journal of Molecular Evolution. 53 (6): 670–9. Bibcode:2001JMolE..53..670N. doi:10.1007/s002390010254. PMID 11677627. S2CID 21151223.
  23. Tartar A, Boucias DG, Adams BJ, Becnel JJ (January 2002). "Phylogenetic analysis identifies the invertebrate pathogen Helicosporidium sp. as a green alga (Chlorophyta)". International Journal of Systematic and Evolutionary Microbiology. 52 (Pt 1): 273–9. doi:10.1099/00207713-52-1-273. PMID 11837312.
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  27. Lopes Dos Santos A, Pollina T, Gourvil P, Corre E, Marie D, Garrido JL, Rodríguez F, Noël MH, Vaulot D, Eikrem W (October 2017). "Chloropicophyceae, a new class of picophytoplanktonic prasinophytes". Scientific Reports. 7 (1): 14019. Bibcode:2017NatSR...714019L. doi:10.1038/s41598-017-12412-5. PMC 5656628. PMID 29070840.
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  29. Sánchez-Baracaldo P, Raven JA, Pisani D, Knoll AH (September 2017). "Early photosynthetic eukaryotes inhabited low-salinity habitats". Proceedings of the National Academy of Sciences of the United States of America. 114 (37): E7737–E7745. doi:10.1073/pnas.1620089114. PMC 5603991. PMID 28808007.
  30. Gitzendanner MA, Soltis PS, Wong GK, Ruhfel BR, Soltis DE (March 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. PMID 29603143.
  31. Rockwell NC, Martin SS, Li FW, Mathews S, Lagarias JC (May 2017). "The phycocyanobilin chromophore of streptophyte algal phytochromes is synthesized by HY2". The New Phytologist. 214 (3): 1145–1157. doi:10.1111/nph.14422. PMC 5388591. PMID 28106912.
  32. Li L, Wang S, Wang H, Sahu SK, Marin B, Li H, et al. (September 2020). "The genome of Prasinoderma coloniale unveils the existence of a third phylum within green plants". Nature Ecology & Evolution. 4 (9): 1220–1231. doi:10.1038/s41559-020-1221-7. PMC 7455551. PMID 32572216.
  33. Pombert JF, Otis C, Lemieux C, Turmel M (September 2005). "The chloroplast genome sequence of the green alga Pseudendoclonium akinetum (Ulvophyceae) reveals unusual structural features and new insights into the branching order of chlorophyte lineages". Molecular Biology and Evolution. 22 (9): 1903–18. doi:10.1093/molbev/msi182. PMID 15930151.
  34. Mattox KR, Stewart KD, et al. (The Systematics Association) (1984). "Classification of the green algae: a concept based on comparative cytology.". In Irvine DE, John DM (eds.). The systematics of Green Algae. 27. London: Academic Press. pp. 29–72.
  35. Bold HC, Wynne MJ (1985). Introduction to the algae : structure and reproduction (2nd ed.). Englewood Cliffs, N.J.: Prentice-Hall. ISBN 978-0-13-477746-7.
  36. Round FE (1971). "The taxonomy of the Chlorophyta, 2". Br. Phycol. J. 6 (2): 235–264. doi:10.1080/00071617100650261.
  37. McCall R (2020-02-24). "Billion-year-old green algae found in China is the oldest ever discovered". Newsweek. Retrieved 2020-02-25.
  38. Sandhya Ramesh (2020-02-25). "Scientists discover world's oldest green algae fossil in China". ThePrint. Retrieved 2020-02-25.

Further reading
  • Burrows EM (1991). Seaweeds of the British Isles. 2 (Chlorophyta). London: Natural History Museum. ISBN 978-0-565-00981-6.
  • Lewis LA, McCourt RM (October 2004). "Green algae and the origin of land plants". American Journal of Botany. 91 (10): 1535–56. doi:10.3732/ajb.91.10.1535. PMID 21652308.
  • Pickett-Heaps JD (1975). Green Algae. Structure, Reproduction and Evolution in Selected Genera. Stamford, CT: Sinauer Assoc. p. 606.
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