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Haptophyte

The haptophytes, classified either as the Haptophytina, Haptophyta or Prymnesiophyta (named for Prymnesium), are a clade of algae that can produce minerals.

The names Haptophyceae or Prymnesiophyceae are sometimes used instead. This ending implies classification at the class rank rather than as a division. Although the phylogenetics of this group has become much better understood in recent years, there remains some dispute over which rank is most appropriate.

Characteristics

The chloroplasts are pigmented similarly to those of the heterokonts, but the structure of the rest of the cell is different, so it may be that they are a separate line whose chloroplasts are derived from similar red algal endosymbionts. Haptophyte chloroplasts contain chlorophylls a, c<sub>1</sub>, and c<sub>2</sub> but lack chlorophyll b. For carotenoids, they have beta-, alpha-, and gamma- carotenes. Like diatoms and brown algae, they have also fucoxanthin, an oxidized isoprenoid derivative that is likely the most important driver of their brownish-yellow color.

The cells typically have two slightly unequal flagella, both of which are smooth, and a unique organelle called a haptonema, which is superficially similar to a flagellum but differs in the arrangement of microtubules and in its use. The name comes from the Greek hapsis, touch, and nema, round thread. The mitochondria have tubular cristae.

Most haptophytes reportedly produce chrysolaminarin rather than starch as their major storage polysaccharide, but some Pavlovaceae produce paramylon. The chain length of the chrysolaminarin is reportedly short (polymers of 20–50 glycosides, unlike the 300+ of comparable amylose), and it is located in cytoplasmic membrane-bound vacuoles.

Significance

The best-known haptophytes are coccolithophores, which make up 673 of the 762 described haptophyte species, and have an exoskeleton of calcareous plates called coccoliths. Coccolithophores are some of the most abundant marine phytoplankton, especially in the open ocean, and are extremely abundant as microfossils, forming chalk deposits. Other planktonic haptophytes of note include Chrysochromulina and Prymnesium, which periodically form toxic marine algal blooms, and Phaeocystis, blooms of which can produce unpleasant foam which often accumulates on beaches.

Haptophytes are economically important, as species such as Pavlova lutheri and Isochrysis sp. are widely used in the aquaculture industry to feed oyster and shrimp larvae. They contain a large amount of polyunsaturated fatty acids such as docosahexaenoic acid (DHA), stearidonic acid and alpha-linolenic acid. Tisochrysis lutea contains betain lipids and phospholipids.

Classification

The haptophytes were first placed in the class Chrysophyceae (golden algae), but ultrastructural data have provided evidence to classify them separately. Both molecular and morphological evidence supports their division into five orders; coccolithophores make up the Isochrysidales and Coccolithales. Very small (2–3μm) uncultured pico-prymnesiophytes are ecologically important.

Haptophytes was discussed to be closely related to cryptomonads.

Haptophytes are closely related to the SAR clade.

  • Subphylum Haptophytina <small>Cavalier-Smith 2015</small> [Haptophyta <small>Hibberd 1976 sensu Ruggerio et al. 2015</small>]
  • Class Rappephyceae <small>Cavalier-Smith 2015</small>
  • Order Rappemonadales
  • Family Rappemonadaceae
  • Class Pavlovophyceae <small>Cavalier-Smith 1986</small> [Pavlovophycidae <small>Cavalier-Smith 1986</small>]
  • Order Pavlovales <small>Green 1976</small>
  • Family Pavlovaceae <small>Green 1976</small>
  • Class Prymnesiophyceae <small>Christensen 1962 emend. Cavalier-Smith 1996</small> [Haptophyceae s.s.; Prymnesiophycidae <small>Cavalier-Smith 1986</small>; Coccolithophyceae <small>Casper 1972 ex Rothmaler 1951</small>]
  • Family †Eoconusphaeraceae <small>Kristan-Tollmann 1988</small> [Conusphaeraceae]
  • Family †Goniolithaceae <small>Deflandre 1957</small>
  • Family †Lapideacassaceae <small>Black, 1971</small>
  • Family †Microrhabdulaceae <small>Deflandre 1963</small>
  • Family †Nannoconaceae <small>Deflandre 1959</small>
  • Family †Polycyclolithaceae <small>Forchheimer 1972 emend Varol, 1992</small>
  • Family †Lithostromationaceae <small>Deflandre 1959</small>
  • Family †Rhomboasteraceae <small>Bown, 2005</small>
  • Family Braarudosphaeraceae <small>Deflandre 1947</small>
  • Family Ceratolithaceae <small>Norris 1965 emend Young & Bown 2014</small> [Triquetrorhabdulaceae <small>Lipps 1969 - cf Young & Bown 2014</small>]
  • Family Alisphaeraceae <small>Young et al., 2003</small>
  • Family Papposphaeraceae <small>Jordan & Young 1990 emend Andruleit & Young 2010</small>
  • Family Umbellosphaeraceae <small>Young et al., 2003</small> [Umbellosphaeroideae]
  • Order †Discoasterales <small>Hay 1977</small>
  • Family †Discoasteraceae <small>Tan 1927</small>
  • Family †Heliolithaceae <small>Hay & Mohler 1967</small>
  • Family †Sphenolithaceae <small>Deflandre 1952</small>
  • Family †Fasciculithaceae <small>Hay & Mohler 1967</small>
  • Order Phaeocystales <small>Medlin 2000</small>
  • Family Phaeocystaceae <small>Lagerheim 1896</small>
  • Order Prymnesiales <small>Papenfuss 1955 emend. Edvardsen and Eikrem 2000</small>
  • Family Chrysochromulinaceae <small>Edvardsen, Eikrem & Medlin 2011</small>
  • Family Prymnesiaceae <small>Conrad 1926 ex Schmidt 1931</small>
  • Subclass Calcihaptophycidae
  • Order Isochrysidales <small>Pascher 1910</small> [Prinsiales <small>Young & Bown 1997</small>]
  • Family †Prinsiaceae <small>Hay & Mohler 1967 emend. Young & Bown, 1997</small>
  • Family Isochrysidaceae <small>Parke 1949</small> [Chrysotilaceae; Marthasteraceae <small>Hay 1977</small>]
  • Family Noelaerhabdaceae <small>Jerkovic 1970 emend. Young & Bown, 1997</small> [Gephyrocapsaceae <small>Black 1971</small>]
  • Order †Eiffellithales <small>Rood, Hay & Barnard 1971</small> (loxolith; imbricating murolith)
  • Family †Chiastozygaceae <small>Rood, Hay & Barnard 1973</small> [Ahmuellerellaceae <small>Reinhardt, 1965</small>]
  • Family †Eiffellithaceae <small>Reinhardt 1965</small>
  • Family †Rhagodiscaceae <small>Hay 1977</small>
  • Order Stephanolithiales <small>Bown & Young 1997</small> (protolith; non-imbrication murolith)
  • Family Parhabdolithaceae <small>Bown 1987</small>
  • Family †Stephanolithiaceae <small>Black 1968 emend. Black 1973</small>
  • Order Zygodiscales <small>Young & Bown 1997</small> [Crepidolithales]
  • Family Helicosphaeraceae <small>Black 1971</small>
  • Family Pontosphaeraceae <small>Lemmermann 1908</small>
  • Family †Zygodiscaceae <small>Hay & Mohler 1967</small>
  • Order Syracosphaerales <small>Ostenfeld 1899 emend. Young et al., 2003</small> [Rhabdosphaerales <small>Ostenfeld 1899</small>]
  • Family Calciosoleniaceae <small>Kamptner 1927</small>
  • Family Syracosphaeraceae <small>Lohmann, 1902</small> [Halopappiaceae <small>Kamptner 1928</small>] (caneolith & cyrtolith; murolith)
  • Family Rhabdosphaeraceae <small>Haeckel, 1894</small> (planolith)
  • Order †Watznaueriales <small>Bown 1987</small> (imbricating placolith)
  • Family †Watznaueriaceae <small>Rood, Hay & Barnard 1971</small>
  • Order †Arkhangelskiales <small>Bown & Hampton 1997</small>
  • Family †Arkhangelskiellaceae <small>Bukry 1969</small>
  • Family †Kamptneriaceae <small>Bown & Hampton 1997</small>
  • Order †Podorhabdales <small>Rood 1971</small> [Biscutales <small>Aubry 2009</small>; Prediscosphaerales <small>Aubry 2009</small>] (non-imbricating or radial placolith)
  • Family †Axopodorhabdaceae <small>Wind & Wise 1977</small> [Podorhabdaceae <small>Noel 1965</small>]
  • Family †Biscutaceae <small>Black, 1971</small>
  • Family †Calyculaceae <small>Noel 1973</small>
  • Family †Cretarhabdaceae <small>Thierstein 1973</small>
  • Family †Mazaganellaceae <small>Bown 1987</small>
  • Family †Prediscosphaeraceae <small>Rood et al., 1971</small> [Deflandriaceae <small>Black 1968</small>]
  • Family †Tubodiscaceae <small>Bown & Rutledge 1997</small>
  • Order Coccolithales <small>Schwartz 1932</small> [Coccolithophorales]
  • Family Reticulosphaeraceae <small>Cavalier-Smith 1996</small> [Reticulosphaeridae]
  • Family Calcidiscaceae <small>Young & Bown 1997</small>
  • Family Coccolithaceae <small>Poche 1913 emend. Young & Bown, 1997</small> [Coccolithophoraceae]
  • Family Pleurochrysidaceae <small>Fresnel & Billard 1991</small>
  • Family Hymenomonadaceae <small>Senn 1900</small> [Ochrosphaeraceae <small>Schussnig 1930</small>]

References

External links