Ochrophyte
The ochrophytes are is a group of mostly photosynthetic stramenopiles (heterokonts), placed either in phylum Ochrophyta or subphylum Ochrophytina.[5][6][7] Their plastid is of red algal origin.[8][7]
Ochrophyte | |
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Dense kelp forest with understory at Partridge Point near Dave's Caves, Cape Peninsula | |
Scientific classification ![]() | |
Domain: | Eukaryota |
Clade: | Diaphoretickes |
Clade: | SAR |
Clade: | Stramenopiles |
Phylum: | Gyrista |
Subphylum: | Ochrophytina Cavalier-Smith, 1995[1] |
Classes | |
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Synonyms | |
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The classification of the group is still being worked out. Originally, the ochrophytes were regarded as a phylum denominated Ochrophyta. Some authors (e.g., Cavalier-Smith) divided it into two subphyla, Phaeista Cavalier-Smith 1995 (comprising Hypogyristea and Chrysista in some classifications, or Limnista and Marista in others) and Khakista Cavalier-Smith, 2000 (comprising Bolidophyceae and diatoms).[9] Others prefer not to use the subphyla, listing only lower taxa (e.g., Reviers, 2002, Guiry & Guiry, 2014).
However, it is currently regarded as a subphylum inside of the phylum Gyrista, along with Pseudofungi and Bigyromonada. It contains two infraphyla: Diatomista (containing diatoms and related groups) and Chrysista (containing brown and golden algae and related groups).[7]
Phylogeny
The cladogram below shows the evolutionary relationships between all ochrophyte classes. It is based on the latest phylogenetic analyses:[10][7][11][12]
Ochrophytina |
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The probable sister clade to Ochrophyta/Ochrophytina is Actinophryidae,[13] originally classified as heliozoan protists.[14]
Classification
Below is the present classification of ochrophytes,[15][7] with the inclusion of three new classes of algae described in 2019,[16] 2020[11] and 2021.[12]
- Infraphylum Chrysista Cavalier-Smith 1991
- Superclass Limnistia Cavalier-Smith 1996 emend. 2006
- Class Eustigmatophyceae Hibberd & Leedale 1971
- Class Chrysomonadea Saville-Kent 1881 stat. nov. Pascher 1910 emend. Cavalier-Smith 2017 [=Chrysophyceae Pascher 1914]
- Class Picophagea Cavalier-Smith 2006 emend. 2017 [=Synchromophyceae Horn et al. 2007]
- Superclass Raphidoistia Cavalier-Smith 1986 orth. mut. 2006
- Class Raphidomonadea Chadefaud ex Silva 1980
- Subclass Raphidophycidae Cavalier-Smith 2013
- Subclass Raphopoda Cavalier-Smith 2013
- Class Raphidomonadea Chadefaud ex Silva 1980
- Superclass Fucistia Cavalier-Smith 1995
- Class Aurophyceae Cavalier-Smith 2013
- Subclass Aurearenophycidae Kai, Yoshii, Nakayama & Inouye 2008 (as Aurearenophyceae)[17] stat. nov. Cavalier-Smith 2013
- Subclass Phaeothamniophycidae Andersen & Bailey 1998 (as Phaeothamniophyceae)[18] stat. nov. Cavalier-Smith 2006[19]
- Class Chrysomerophyceae Cavalier-Smith 1995
- Class Chrysoparadoxophyceae Wetherbee et al. 2019[16]
- Class Phaeosacciophyceae R.A.Andersen, L.Graf & H.S.Yoon 2020[11]
- Class Schizocladiophyceae Henry, Okuda & Kawai 2003[20]
- Class Phaeophyceae Kjellman 1891 [=Fucophyceae Warming 1884; Melanophyceae Rabenhorst 1863] ("brown algae")
- Subclass Discosporangiophycidae
- Subclass Ishigeophycidae Silberfeld, F. Rousseau & Reviers 2014
- Subclass Dictyotophycidae Silberfeld, F. Rousseau & Reviers 2014
- Subclass Fucophycidae Cavalier-Smith 1986
- Class Xanthophyceae Allorge ex Fritsch 1935 [=Tribophyceae Hibberd 1981;[21] Heterokontae Luther 1899] ("yellow-green algae")
- Class Aurophyceae Cavalier-Smith 2013
- Superclass Limnistia Cavalier-Smith 1996 emend. 2006
- Infraphylum Diatomista Derelle et al. ex Cavalier-Smith 2017
- Superclass Hypogyrista Cavalier-Smith 1995 stat. nov. 2006
- Class Dictyochophyceae Silva 1980 [=Dictyochia Haeckel 1894; Alophycidae Cavalier-Smith 2006]
- Subclass Pedinellia Cavalier-Smith 1986 stat. nov. 2017 [=Actinochrysophyceae Cavalier-Smith 1995] ("axodines")
- Subclass Pelagophycidae Andersen & Saunders 1993 ex Cavalier-Smith 2006 [=Pelagophyceae Andersen & Saunders 1993]
- ?Subclass Sulcophycidae Cavalier-Smith 2013[lower-alpha 1]
- Class Pinguiophyceae Kawachi et al. 2002
- Class Dictyochophyceae Silva 1980 [=Dictyochia Haeckel 1894; Alophycidae Cavalier-Smith 2006]
- Superclass Khakista Cavalier-Smith 2000 stat. nov. 2017[7]
- Class Bolidophyceae Guillou & Chretiennot-Dinet 1999
- Class Diatomeae Dumortier 1821 [=Bacillariae Haeckel 1878; Bacillariophyceae Dangeard 1933]
- Subclass Corethrophycidae Round & Crawford 1990
- Subclass Rhizosoleniophycidae Round & Crawford 1990
- Subclass Eucentricophycidae Cavalier-Smith 2000
- Subclass Bacillariophycidae [=Pennatia Pennatia Schütt 1896; =Pennatophycidae]
- Superclass Hypogyrista Cavalier-Smith 1995 stat. nov. 2006
- Ochrophytina incertae sedis
- Class Olisthodiscophyceae Barcytė, Eikrem & M. Eliáš 2021[12]
Notes
- Cavalier-Smith suggested that Sulcochrysis is possibly related to Olisthodiscus, and grouped both genera within this subclass.[22][7] However, a study in 2021 reveals that Olisthodiscales is part of a new class Olisthodiscophyceae, which includes only the genus Olisthodiscus. The position of the genus Sulcochrysis remains uncertain due to a lack of DNA sequences.[12]
References
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- Cavalier-Smith T (1995). "Zooflagellate phylogeny and classification". Cytology. 37 (11): 1010–1029. PMID 8868448.
- Cavalier-Smith, T. (1986). The kingdom Chromista, origin and systematics. In: Round, F.E. and Chapman, D.J. (eds.). Progress in Phycological Research. 4: 309–347.
- Reviers, B. de. (2006). Biologia e Filogenia das Algas. Editora Artmed, Porto Alegre, p. 157.
- Reviers, B. de. (2006). Biologia e Filogenia das Algas. Editora Artmed, Porto Alegre, pp. 15-16.
- Thomas Cavalier-Smith & Ema E.-Y. Chao (2006). "Phylogeny and megasystematics of phagotrophic heterokonts (kingdom Chromista)". Journal of Molecular Evolution. 62 (4): 388–420. Bibcode:2006JMolE..62..388C. doi:10.1007/s00239-004-0353-8. PMID 16557340. S2CID 29567514.
- Ingvild Riisberga; Russell J. S. Orr; Ragnhild Kluge; Kamran Shalchian-Tabrizi; Holly A. Bowers; Vishwanath Patil; Bente Edvardsen; Kjetill S. Jakobsen (2009). "Seven gene phylogeny of heterokonts". Protist. 160 (2): 191–204. doi:10.1016/j.protis.2008.11.004. PMID 19213601.
- Cavalier-Smith, Thomas (2018). "Kingdom Chromista and its eight phyla: a new synthesis emphasising periplastid protein targeting, cytoskeletal and periplastid evolution, and ancient divergences". Protoplasma. 255 (1): 297–357. doi:10.1007/s00709-017-1147-3. PMC 5756292. PMID 28875267.
- Ševčíková T, Horák A, Klimeš V, et al. (2015). "Updating algal evolutionary relationships through plastid genome sequencing: did alveolate plastids emerge through endosymbiosis of an ochrophyte?". Sci Rep. 5: 10134. Bibcode:2015NatSR...510134S. doi:10.1038/srep10134. PMC 4603697. PMID 26017773.
- "Heterokontophyta". SHIGEN. National Institute of Genetics. Retrieved June 18, 2009.
- Derelle R, López-García P, Timpano H, Moreira D (November 2016). "A Phylogenomic Framework to Study the Diversity and Evolution of Stramenopiles (=Heterokonts)". Mol Biol Evol. 33 (11): 2890–2898. doi:10.1093/molbev/msw168. PMC 5482393. PMID 27512113.
- Graf L, Yang EC, Han KY, Küpper FC, Benes KM, Oyadomari JK, Herbert RJH, Verbruggen H, Wetherbee R, Andersen RA, Yoon HS (December 2020). "Multigene Phylogeny, Morphological Observation and Re-examination of the Literature Lead to the Description of the Phaeosacciophyceae Classis Nova and Four New Species of the Heterokontophyta SI Clade". Protist. 171 (6): 125781. doi:10.1016/j.protis.2020.125781. PMID 33278705. S2CID 227315556.
- Dovilė Barcytė; Wenche Eikrem; Anette Engesmo; Sergio Seoane; Jens Wohlmann; Aleš Horák; Tatiana Yurchenko; Marek Eliáš (2 March 2021). "Olisthodiscus represents a new class of Ochrophyta". Journal of Phycology. 57 (4): 1094–1118. doi:10.1111/jpy.13155. hdl:10852/86515. PMID 33655496.
- Azuma, Tomonori; Pánek, Tomáš; Tice, Alexander K.; Kayama, Motoki; Kobayashi, Mayumi; Miyashita, Hideaki; Suzaki, Toshinobu; Yabuki, Akinori; Brown, Matthew W.; Kamikawa, Ryoma (10 April 2022). "An Enigmatic Stramenopile Sheds Light on Early Evolution in Ochrophyta Plastid Organellogenesis". Molecular Biology and Evolution. 39 (4). doi:10.1093/molbev/msac065. PMC 9004409. PMID 35348760.
- Mikrjukov, Kirill A.; Patterson, David J. (2001). "Taxonomy and phylogeny of Heliozoa. III. Actinophryids" (PDF). Acta Protozoologica. 40: 3–25.
- Silberfeld T, Rousseau F, de Reviers B (2014). "An Updated Classification of Brown Algae (Ochrophyta, Phaeophyceae)". Cryptogamie, Algologie. 35 (2): 117–156. doi:10.7872/crya.v35.iss2.2014.117. S2CID 86227768.
- Wetherbee R, Jackson CJ, Repetti SI, Clementson LA, Costa JF, van de Meene A, Crawford S, Verbruggen H (April 2019). "The golden paradox - a new heterokont lineage with chloroplasts surrounded by two membranes". J Phycol. 55 (2): 257–278. doi:10.1111/jpy.12822. hdl:11343/233613. PMID 30536815. S2CID 54477112.
- Kai A, Yoshii Y, Nakayama T, Inouye I (2008). "Aurearenophyceae classis nova, a New Class of Heterokontophyta Based on a New Marine Unicellular Alga Aurearena cruciata gen. et sp. nov. Inhabiting Sandy Beaches". Protist. 159 (3): 435–457. doi:10.1016/j.protis.2007.12.003. ISSN 1434-4610. PMID 18358776.
- Craig Bailey J, Bidigare RR, Christensen SJ, Andersen RA (September 1998). "Phaeothamniophyceae Classis Nova: A New Lineage of Chromophytes Based upon Photosynthetic Pigments, rbcL Sequence Analysis and Ultrastructure". Protist. 149 (3): 245–63. doi:10.1016/S1434-4610(98)70032-X. PMID 23194637.
- Cavalier-Smith T, Chao EE (22 March 2006). "Phylogeny and Megasystematics of Phagotrophic Heterokonts (Kingdom Chromista)". J Mol Evol. 62 (4): 388–420. Bibcode:2006JMolE..62..388C. doi:10.1007/s00239-004-0353-8. PMID 16557340. S2CID 29567514.
- Kawai, Hiroshi; Maeba, Shunsuke; Sasaki, Hideaki; Okuda, Kazuo; Henry, Eric C. (2003). "Schizocladia ischiensis: A New Filamentous Marine Chromophyte Belonging to a New Class, Schizocladiophyceae". Protist. 154 (2): 211–228. doi:10.1078/143446103322166518. ISSN 1434-4610. PMID 13677449.
- Hibberd DJ (February 1981). "Notes on the taxonomy and nomenclature of the algal classes Eustigmatophyceae and Tribophyceae (synonym Xanthophyceae)". Botanical Journal of the Linnean Society. 82 (2): 93–119. doi:10.1111/j.1095-8339.1981.tb00954.x.
- Cavalier-Smith, Thomas; Scoble, Josephine Margaret (2012). "Phylogeny of Heterokonta: Incisomonas marina, a uniciliate gliding opalozoan related to Solenicola (Nanomonadea), and evidence that Actinophryida evolved from raphidophytes". European Journal of Protistology. 49 (3): 328–353. doi:10.1016/j.ejop.2012.09.002. PMID 23219323.