Chlorophyta
Chlorophyta | |
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Chlorophytes (A–F, H–L and O) | |
Scientific classification ![]() | |
Domain: | Eukaryota |
Clade: | Diaphoretickes |
(unranked): | Archaeplastida |
(unranked): | Viridiplantae |
Division: | Chlorophyta Reichenbach, 1828, emend. Pascher, 1914, emend. Lewis & McCourt, 2004[1][2][3] |
Classes[4] | |
Diversity | |
7,934 species (6,851 extant, 1,083 fossil)[4] | |
Synonyms | |
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![](http://upload.wikimedia.org/wikipedia/commons/thumb/a/af/Taiwan_2009_East_Coast_ShihTiPing_Giant_Stone_Steps_Algae_FRD_6581.jpg/220px-Taiwan_2009_East_Coast_ShihTiPing_Giant_Stone_Steps_Algae_FRD_6581.jpg)
Chlorophyta is a taxon of green algae informally called chlorophytes.[6]
Systematics
[edit]Taxonomy
[edit]The name Chlorophyta was historically attributed to a paraphyletic group of all green algae, i.e., all organisms belonging to the Viridiplantae with the exception of land plants (Embryophyta).[7] In modern classifications, Chlorophyta is only one of the many clades of green algae, sister to the Streptophyta clade from which land plants evolved.[8][9] In this latter sense, the Chlorophyta includes only about 7,900 species.[4] About 90% of all known species live in freshwater.[10] Like other green algae and land plants, chlorophytes contain chlorophylls a and b, and store food as starch in their plastids.[7]
With the exception of the three classes Ulvophyceae, Trebouxiophyceae and Chlorophyceae in the UTC clade, which show various degrees of multicellularity, all the Chlorophyta lineages are unicellular.[11] 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.[12] Monostroma kuroshiense, an edible green alga cultivated worldwide and most expensive among green algae, belongs to this group.
Ecology
[edit]Most species of Chlorophyta are aquatic, prevalent in both marine and freshwater environments. However, some species have adapted to a wide range of terrestrial environments. For example, Chlamydomonas nivalis lives on summer alpine snowfields, and Trentepohlia species, live attached to rocks or woody parts of trees.[13][14] 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.[15][16][17][18] Some groups, such as the Trentepohliales are exclusively found on land.[19] Several species of Chlorophyta live in symbiosis with a diverse range of eukaryotes, including fungi (to form lichens), ciliates, forams, cnidarians and molluscs.[14] Some species of Chlorophyta are heterotrophic, either free-living or parasitic.[20][21] Others are mixotrophic bacterivores through phagocytosis.[22] Two common species of the heterotrophic green alga Prototheca are pathogenic and can cause the disease protothecosis in humans and animals.[23]
Classifications
[edit]![](http://upload.wikimedia.org/wikipedia/commons/thumb/7/7d/Haeckel_Siphoneae.jpg/220px-Haeckel_Siphoneae.jpg)
Characteristics used for the classification of Chlorophyta are: type of zoid, mitosis (karyokinesis), cytokinesis, organization level, life cycle, type of gametes, cell wall polysaccharides[24] and more recently genetic data.
Phylogeny
[edit]Leliaert et al. 2012 proposed the following phylogeny. He marked the "prasinophytes" as paraphyletic, with the remaining Chlorophyta groups as "core chlorophytes". He described all Streptophyta except the land plants as paraphyletic "charophytes".[14]
"Hypothetical ancestral |
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green flagellate" |
A 2020 paper places the "Prasinodermophyta" (i.e. Prasinodermophyceae + Palmophyllophyceae) as the basal Viridiplantae clade.[25]
Viridiplantae/ |
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Green Algae/ Prasinophyta s.l. |
Leliaert et al. 2012
[edit]Simplified phylogeny of the Chlorophyta, according to Leliaert et al. 2012.[14] Note that many algae previously classified in Chlorophyta are placed here in Streptophyta.
- Viridiplantae
- Chlorophyta
- core chlorophytes
- prasinophytes (paraphyletic)
- Streptophyta
- Chlorophyta
Pombert et al. 2005
[edit]A possible classification when Chlorophyta refers to one of the two clades of the Viridiplantae is shown below.[26]
- Class Prasinophyceae T. A. Chr. ex Ø. Moestrup & J. Throndsen
- Class Chlorophyceae Wille
- Class Trebouxiophyceae T. Friedl
- Class Ulvophyceae
Lewis & McCourt 2004
[edit]- Division Chlorophyta (green algae sensu stricto)
- Subdivision Chlorophytina
- Class Chlorophyceae (chlorophytes)
- Order Chlamydomonadales (+ some Chlorococcales + some Tetrasporales + some Chlorosarcinales)
- Order Sphaeropleales (sensu Deason, plus Bracteacoccus, Schroederia, Scenedesmaceae, Selanastraceae)
- Order Oedogoniales
- Order Chaetopeltidales
- Order Chaetophorales
- Incertae Sedis (Cylindrocapsa clade, Mychonastes clade)
- Class Ulvophyceae (ulvophytes)
- Order Ulotrichales
- Order Ulvales
- Order Siphoncladales/Cladophorales
- Order Caulerpales
- Order Dasycladales
- Class Trebouxiophyceae (trebouxiophytes)
- Order Trebouxiales
- Order Microthamniales
- Order Prasiolales
- Order Chlorellales
- Class Prasinophyceae (prasinophytes)
- Order Pyramimonadales
- Order Mamiellales
- Order Pseudoscourfieldiales
- Order Chlorodendrales
- Incertae sedis (Unnamed clade of coccoid taxa)
- Class Chlorophyceae (chlorophytes)
- Subdivision Chlorophytina
- Division Charophyta (charophyte algae and embryophytes)
- Class Mesostigmatophyceae (mesostigmatophytes)
- Class Chlorokybophyceae (chlorokybophytes)
- Class Klebsormidiophyceae (klebsormidiophytes)
- Class Zygnemophyceae (conjugates)
- Order Zygnematales (filamentous conjugates and saccoderm desmids)
- Order Desmidiales (placoderm desmids)
- Class Coleochaetophyceae (coleochaetophytes)
- Order Coleochaetales
- Subdivision Streptophytina
- Class Charophyceae (reverts to use of GM Smith)
- Order Charales (charophytes sensu stricto)
- Class Embryophyceae (embryophytes)
- Class Charophyceae (reverts to use of GM Smith)
Hoek, Mann and Jahns 1995
[edit]Classification of the Chlorophyta, treated as all green algae, according to Hoek, Mann and Jahns 1995.[7]
- Class Prasinophyceae (orders Mamiellales, Pseudocourfeldiales, Pyramimonadales, Chlorodendrales)
- Class Chlorophyceae (orders Volvocales [including the Tetrasporales], Chlorococcales, Chaetophorales, Oedogoniales)
- Class Ulvophyceae (orders Codiolales, Ulvales)
- Class Cladophorophyceae (order Cladophorales)
- Class Bryopsidophyceae (orders Bryopsidales, Halimedales)
- Class Dasycladophyceae (order Dasycladales)
- Class Trentepohliophyceae (order Trentepohliales)
- Class Pleurastrophyceae (order Pleurastrales)
- Incertae sedis (order Prasiolales)
- Class Klebsormidiophyceae (orders Klebsormidiales, Coleochaetales)
- Class Zygnematophyceae (order Zygnematales, Desmidiales)
- Class Charophyceae (order Charales)
In a note added in proof, an alternative classification is presented for the algae of the class Chlorophyceae:
- Class Chlamydophyceae (orders Volvocales, Chlorococcales, Chaetophorales)
- Class Oedogoniophyceae (order Oedogoniales)
- Class Chlorophyceae (order Chlorellales)
Bold and Wynne 1985
[edit]Classification of the Chlorophyta and Charophyta according to Bold and Wynne 1985.[27]
- Chlorophyta, Chlorophyceae (16 orders)
- Charophyta, Charophyceae (1 order)
Mattox & Stewart 1984
[edit]Classification of the Chlorophyta according to Mattox & Stewart 1984:[28]
- Micromonadophyceae (similar to Prasinophyceae; Tetraselmidiales transferred to Pleurastrophyceae)
- Charophyceae Rabenhorst
- Ulvophyceae
- Pleurastrophyceae
- Chlorophyceae
Fott 1971
[edit]Classification of the Chlorophyta according to Fott 1971.[29]
- Class Chlorophyceae
- Order Volvocales
- Order Tetrasporales
- Order Chlorococcales
- Order Ulotrichales
- Suborder Chlorosarcineae
- Suborder Ulotrichineae
- Suborder Oedogoniineae
- Suborder Chaetophorineae
- Order Siphonocladales
- Order Bryopsidales
- Class Conjugatophyceae
- Class Charophyceae
Round 1971
[edit]Classification of the Chlorophyta and related algae according to Round 1971.[30]
- "green algae"
- Euglenophyta
- Prasinophyta
- Charophyta
- Chlorophyta
Smith 1938
[edit]Classification of the Chlorophyta according to Smith 1938:
- Class 1. Chlorophyceae
- Order 1. Volvocales
- Family 1. Chlamydomonadaceae
- Family 2. Volvocaceae
- Order 2. Tetrasporales
- Order 3. Ulotrichales
- Family 1. Ulotrichaceae
- Family 2. Microsporaceae
- Family 3. Cylindrocapsaceae
- Family 4. Chaetophoraceae
- Family 5. Protococcaceae
- Family 6. Coleochaetaceae
- Family 7. Trentepohliaceae
- Order 4. Ulvales
- Family 1. Ulvaceae
- Family 2 Schizomeridaceae
- Order 5. Schizogoniales
- Family Schizogoniaceae
- Order 6. Cladophorales
- Family 1. Cladophoraceae
- Family 2. Sphaeropleaceae
- Order 7. Oedogoniales
- Family Oedogoniaceae
- Order 8. Zygnematales
- Family 1. Zygnemataceae
- Family 2. Mesotaeniaceae
- Family 3. Desmidiaceae
- Order 9. Chlorococcales
- Family 1. Chlorococcaceae
- Family 2. Endosphaeraceae
- Family 3. Characiaceae
- Family 4. Protosiphonaceae
- Family 5. Hydrodictyaceae
- Family 6. Oöcystaceae
- Family 7. Scenedesmaceae
- Order 10. Siphonales
- Family 1. Bryopsidaceae
- Family 2. Caulerpaceae
- Family 3. Halicystaceae
- Family 4. Codiaceae
- Family 5. Derbesiaceae
- Family 6. Vaucheriaceae
- Family 7. Phyllosiphonaceae
- Order 11. Siphonocladiales
- Family 1. Valoniaceae
- Family 2. Dasycladaceae
- Class 2. Charophyceae
Fossil record
[edit]In February 2020, the fossilized remains of green algae, named Proterocladus antiquus were discovered in the northern province of Liaoning, China. At around a billion years old, it is believed to be one of the oldest examples of a multicellular chlorophyte.[31]
References
[edit]Citations
[edit]- ^ Reichenbach 1828, p. 23.
- ^ Pascher 1914.
- ^ Adl et al. 2019, p. 36.
- ^ a b c Guiry 2024.
- ^ Papenfuss 1955.
- ^ Rockwell et al. 2017.
- ^ a b c van den Hoek, Mann & Jahns 1995.
- ^ Lewis & McCourt 2004.
- ^ Becker & Marin 2009.
- ^ Lee 2018.
- ^ Umen 2014.
- ^ Kapraun 2007.
- ^ Graham et al. 2022.
- ^ a b c d Leliaert et al. 2012.
- ^ Lewis & Lewis 2005.
- ^ De Wever et al. 2009.
- ^ Leliaert, Verbruggen & Zechman 2011.
- ^ Foflonker et al. 2016.
- ^ López-Bautista, Rindi & Guiry 2006.
- ^ Joubert & Rijkenberg 1971.
- ^ Nedelcu 2001.
- ^ Anderson, Charvet & Hansen 2018.
- ^ Tartar et al. 2002.
- ^ Lobban & Wynne 1981, p. 88.
- ^ Li et al. 2020.
- ^ Pombert et al. 2005.
- ^ Bold & Wynne 1985.
- ^ Mattox & Stewart 1984.
- ^ van den Hoek, Mann & Jahns 1995, p. 483.
- ^ Round 1971.
- ^ Tang et al. 2020.
Cited literature
[edit]- Adl, Sina M.; Bass, David; Lane, Christopher E.; Lukeš, Julius; Schoch, Conrad L.; Smirnov, Alexey; Agatha, Sabine; Berney, Cedric; Brown, Matthew W.; Burki, Fabien; et al. (2019). "Revisions to the Classification, Nomenclature, and Diversity of Eukaryotes". Journal of Eukaryotic Microbiology. 66 (1): 4–119. doi:10.1111/JEU.12691. PMC 6492006. PMID 30257078.
- Anderson, R.; Charvet, S.; Hansen, P. J. (2018). "Mixotrophy in Chlorophytes and Haptophytes—Effect of Irradiance, Macronutrient, Micronutrient and Vitamin Limitation". Frontiers in Microbiology. 9: 1704. doi:10.3389/fmicb.2018.01704. PMC 6080504. PMID 30108563.
- Becker, Burkhard; Marin, Birger (2009). "Streptophyte algae and the origin of embryophytes". Annals of Botany. 103 (7): 999–1004. doi:10.1093/aob/mcp044. PMC 2707909. PMID 19273476.
- Bold, Harold Charles; Wynne, Michael James (1985). Introduction to the algae : structure and reproduction (2nd ed.). Englewood Cliffs, N.J.: Prentice-Hall. ISBN 978-0-13-477746-7.
- De Wever, Aaike; Leliaert, Frederik; Verleyen, Elie; Vanormelingen, Pieter; Van der Gucht, Katleen; Hodgson, Dominic A.; Sabbe, Koen; Vyverman, Wim (October 2009). "Hidden levels of phylodiversity in Antarctic green algae: further evidence for the existence of glacial refugia". Proceedings of the Royal Society B: Biological Sciences. 276 (1673): 3591–3599. doi:10.1098/rspb.2009.0994. PMC 2817313. PMID 19625320.
- Foflonker, Fatima; Ananyev, Gennady; Qiu, Huan; Morrison, Andrenette; Palenik, Brian; Dismukes, G. Charles; Bhattacharya, Debashish (June 2016). "The unexpected extremophile: Tolerance to fluctuating salinity in the green alga Picochlorum". Algal Research. 16: 465–472. doi:10.1016/j.algal.2016.04.003.
- Graham, Linda E.; Graham, James M.; Wilcox, Lee W.; Cook, Martha E. (2022). Algae (4th ed.). LJLM Press. ISBN 978-0-9863935-4-9.
- Guiry, Michael D. (2024). "How many species of algae are there? A reprise. Four kingdoms, 14 phyla, 63 classes and still growing". Journal of Phycology. 60 (2): 214–228. doi:10.1111/jpy.13431.
- Joubert, J. J.; Rijkenberg, F. H. (1971). "Parasitic green algae". Annual Review of Phytopathology. 9: 45–64. doi:10.1146/annurev.py.09.090171.000401.
- Kapraun, Donald F. (April 2007). "Nuclear DNA content estimates in green algal lineages: chlorophyta and streptophyta". Annals of Botany. 99 (4): 677–701. doi:10.1093/aob/mcl294. PMC 2802934. PMID 17272304.
- Lee, Robert Edward (2018). "Chlorophyta". Phycology (5th ed.). Cambridge University Press. pp. 133–230. doi:10.1017/9781316407219. ISBN 9781316407219.
- Leliaert, Frederik; Verbruggen, Heroen; Zechman, Frederick W. (September 2011). "Into the deep: new discoveries at the base of the green plant phylogeny". BioEssays. 33 (9): 683–692. doi:10.1002/bies.201100035. PMID 21744372. S2CID 40459076.
- 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. S2CID 17603352. Archived (PDF) from the original on 2015-06-26.
- Lewis, Louise A.; McCourt, Richard M. (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.
- Lewis, Louise A.; Lewis, Paul O. (December 2005). "Unearthing the molecular phylodiversity of desert soil green algae (Chlorophyta)". Systematic Biology. 54 (6): 936–47. doi:10.1080/10635150500354852. PMID 16338765.
- Li, Linzhou; Wang, Sibo; Wang, Hongli; Sahu, Sunil Kumar; Marin, Birger; Li, Haoyuan; Xu, Yan; Liang, Hongping; Li, Zhen; Cheng, Shifeng; 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.
- Lobban CS, Wynne MJ (1981). The Biology of Seaweeds. Botanical Monograph Series 17. University of California Press. ISBN 9780520045859.
- López-Bautista, Juan M.; Rindi, Fabio; Guiry, Michael D. (July 2006). "Molecular systematics of the subaerial green algal order Trentepohliales: an assessment based on morphological and molecular data". International Journal of Systematic and Evolutionary Microbiology. 56 (7): 1709–1715. doi:10.1099/ijs.0.63990-0. hdl:10379/9448. PMID 16825655.
- Mattox, K. R.; Stewart, K. D. (1984). "Classification of the Green Algae: A Concept Based on Comparative Cytology". In Irvine, D. E. G.; John, D. M. (eds.). Systematics of the Green Algae. Systematics Association Special. Vol. 27. Academic Press. pp. 29–72. ISBN 0-12-374040-1.
- Nedelcu, Aurora M. (December 2001). "Complex patterns of plastid 16S rRNA gene evolution in nonphotosynthetic green algae". Journal of Molecular Evolution. 53 (6): 670–679. Bibcode:2001JMolE..53..670N. doi:10.1007/s002390010254. PMID 11677627. S2CID 21151223.
- Papenfuss, George F. (1955). "The Classification of the Algae". A century of progress in the natural sciences, 1853-1953. California Academy of Sciences. pp. 115–224.
- Pascher, A. (1914). "Über Flagellaten und Algen". Berichte der Deutschen Botanischen Gesellschaft. 32: 136–160. doi:10.1111/j.1438-8677.1914.tb07573.x. S2CID 257830577.
- Pombert, Jean-François; Otis, Christian; Lemieux, Claude; Turmel, Monique (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): 1903–1918. doi:10.1093/molbev/msi182. PMID 15930151.
- Reichenbach, H. TH. L. (1828). Conspectus Regni Vegetabilis per Gradus Naturales Evoluti. Lipsiae (Leipzig): apud Carolum Cnobloch. doi:10.5962/bhl.title.127418. OCLC 5359726.
- Rockwell, Nathan C.; Martin, Shelley S.; Li, Fay-Wei; Mathews, Sarah; Lagarias, J. Clark (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.
- Round, F. E. (1971). "The taxonomy of the Chlorophyta. II". British Phycological Journal. 6 (2): 235–264. doi:10.1080/00071617100650261.
- Tang, Qing; Pang, Ke; Yuan, Xunlai; Xiao, Shuhai (February 2020). "A one-billion-year-old multicellular chlorophyte". Nature Ecology & Evolution. 4: 543–549. doi:10.1038/s41559-020-1122-9.
- Tartar, Aurélien; Boucias, Drion G.; Adams, Byron J.; Becnel, James J. (January 2002). "Phylogenetic analysis identifies the invertebrate pathogen Helicosporidium sp. as a green alga (Chlorophyta)". International Journal of Systematic and Evolutionary Microbiology. 52 (1): 273–279. doi:10.1099/00207713-52-1-273. PMID 11837312.
- Umen, James G. (October 2014). "Green algae and the origins of multicellularity in the plant kingdom". Cold Spring Harbor Perspectives in Biology. 6 (11): a016170. doi:10.1101/cshperspect.a016170. PMC 4413236. PMID 25324214.
- van den Hoek, Christiaan; Mann, D. G.; Jahns, H. M. (1995). Algae: An Introduction to Phycology. Cambridge University Press. ISBN 978-0-521-30419-1.
Further reading
[edit]![](http://upload.wikimedia.org/wikipedia/commons/thumb/d/df/Wikispecies-logo.svg/34px-Wikispecies-logo.svg.png)
![](http://upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/30px-Commons-logo.svg.png)
- Burrows EM (1991). Seaweeds of the British Isles. Vol. 2 (Chlorophyta). London: Natural History Museum. ISBN 978-0-565-00981-6.
- Pickett-Heaps JD (1975). Green Algae. Structure, Reproduction and Evolution in Selected Genera. Stamford, CT: Sinauer Assoc. p. 606.