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
Dense kelp forest with understorey at Partridge Point near Dave's Caves, Cape Peninsula
Dense kelp forest with understory at Partridge Point near Dave's Caves, Cape Peninsula
Scientific classification Edit this classification
Domain: Eukaryota
Clade: Diaphoretickes
Clade: SAR
Clade: Stramenopiles
Phylum: Gyrista
Subphylum: Ochrophytina
Cavalier-Smith, 1995[1]
Classes
Synonyms
  • Heterokontophyta van den Hoek et al., 1978
  • Ochrista Cavalier-Smith, 1986[2][3]
  • Stramenochromes Leipe et al., 1994[4]

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
Chrysista
"SI clade"
Fucistia

Xanthophyceaeyellow-green algae
Allorge ex Fritsch 1935

Chrysoparadoxophyceae
Wetherbee et al. 2019

Phaeosacciophyceae
R.A.Andersen, L.Graf & H.S.Yoon 2020

Schizocladiophyceae
Henry, Okuda & Kawai 2003

Phaeophyceaebrown algae
Kjellman 1891

Aurophyceae
Cavalier-Smith 2013

"PX clade"
Raphidoistia

Raphidomonadea
Chadefaud ex Silva 1980

"SII clade"

Olisthodiscophyceae
Barcytė, Eikrem & M. Eliáš 2021

Limnistia

Chrysophyceaegolden algae
Pascher 1914

Synchromophyceae
Horn et al. 2007

Eustigmatophyceae
Hibberd & Leedale 1971

Diatomista
Hypogyrista

Dictyochophyceae
Silva 1980

Pinguiophyceae
Kawachi et al. 2002

Khakista

Bolidophyceae
Guillou & Chretiennot-Dinet 1999

Bacillariophyceaediatoms
Dangeard 1933

"SIII clade"

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
    • Superclass Fucistia Cavalier-Smith 1995
      • Class Aurophyceae Cavalier-Smith 2013
      • 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")
      • Class Xanthophyceae Allorge ex Fritsch 1935 [=Tribophyceae Hibberd 1981;[21] Heterokontae Luther 1899] ("yellow-green algae")
  • 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
    • 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]
  • Ochrophytina incertae sedis

Notes

  1. 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

  1. Cavalier-Smith T (1995). "Zooflagellate phylogeny and classification". Cytology. 37 (11): 1010–1029. PMID 8868448.
  2. 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.
  3. Reviers, B. de. (2006). Biologia e Filogenia das Algas. Editora Artmed, Porto Alegre, p. 157.
  4. Reviers, B. de. (2006). Biologia e Filogenia das Algas. Editora Artmed, Porto Alegre, pp. 15-16.
  5. 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.
  6. 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.
  7. 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.
  8. 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.
  9. "Heterokontophyta". SHIGEN. National Institute of Genetics. Retrieved June 18, 2009.
  10. 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.
  11. 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.
  12. 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.
  13. 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.
  14. Mikrjukov, Kirill A.; Patterson, David J. (2001). "Taxonomy and phylogeny of Heliozoa. III. Actinophryids" (PDF). Acta Protozoologica. 40: 3–25.
  15. 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.
  16. 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.
  17. 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.
  18. 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.
  19. 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.
  20. 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.
  21. 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.
  22. 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.


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