Haptophyte The haptophytes , classified either as the Haptophyta , Haptophytina or Prymnesiophyta , are a clade of algae . 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 . 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 , thread. The mitochondria have tubular cristae .Significance The best-known haptophytes are coccolithophores, which make up 673 of the 762 described haptophyte species, 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 , stearidonic acid and alpha-linolenic acid . Tisochrysis lutea contains betain lipids and phospholipids.Classification The haptophytes were first placed in the class Chrysophyceae , 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 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 Cavalier-Smith 2015 Clade Rappemonada Kim et al . 2011 * Class Rappephyceae Cavalier-Smith 2015 ** Order Rappemonadales *** Family Rappemonadaceae Clade Haptomonada * Class Pavlovophyceae Cavalier-Smith 1986 ** Order Pavlovales Green 1976 *** Family Pavlovaceae Green 1976 * Class Prymnesiophyceae Christensen 1962 emend. Cavalier-Smith 1996 ** Family †Eoconusphaeraceae Kristan-Tollmann 1988 ** Family †Goniolithaceae Deflandre 1957 ** Family †Lapideacassaceae Black, 1971 ** Family †Microrhabdulaceae Deflandre 1963 ** Family †Nannoconaceae Deflandre 1959 ** Family †Polycyclolithaceae Forchheimer 1972 emend Varol, 1992 ** Family †Lithostromationaceae Deflandre 1959 ** Family †Rhomboasteraceae Bown, 2005 ** Family Braarudosphaeraceae Deflandre 1947 ** Family Ceratolithaceae Norris 1965 emend Young & Bown 2014 ** Family Alisphaeraceae Young et al., 2003 ** Family Papposphaeraceae Jordan & Young 1990 emend Andruleit & Young 2010 ** Family Umbellosphaeraceae Young et al., 2003 ** Order †Discoasterales Hay 1977 *** Family †Discoasteraceae Tan 1927 *** Family †Heliolithaceae Hay & Mohler 1967 *** Family †Sphenolithaceae Deflandre 1952 *** Family †Fasciculithaceae Hay & Mohler 1967 ** Order Phaeocystales Medlin 2000 *** Family Phaeocystaceae Lagerheim 1896 ** Order Prymnesiales Papenfuss 1955 emend. Edvardsen and Eikrem 2000 *** Family Chrysochromulinaceae Edvardsen, Eikrem & Medlin 2011 *** Family Prymnesiaceae Conrad 1926 ex Schmidt 1931 ** Subclass Calcihaptophycidae *** Order Isochrysidales Pascher 1910 **** Family †Prinsiaceae Hay & Mohler 1967 emend. Young & Bown, 1997 **** Family Isochrysidaceae Parke 1949 **** Family Noëlaerhabdaceae Jerkovic 1970 emend. Young & Bown, 1997 *** Order †Eiffellithales Rood, Hay & Barnard 1971 **** Family †Chiastozygaceae Rood, Hay & Barnard 1973 **** Family †Eiffellithaceae Reinhardt 1965 **** Family †Rhagodiscaceae Hay 1977 *** Order Stephanolithiales Bown & Young 1997 **** Family Parhabdolithaceae Bown 1987 **** Family †Stephanolithiaceae Black 1968 emend. Black 1973 *** Order Zygodiscales Young & Bown 1997 **** Family Helicosphaeraceae Black 1971 **** Family Pontosphaeraceae Lemmermann 1908 **** Family †Zygodiscaceae Hay & Mohler 1967 *** Order Syracosphaerales Ostenfeld 1899 emend. Young et al., 2003 **** Family Calciosoleniaceae Kamptner 1927 **** Family Syracosphaeraceae Lohmann, 1902 **** Family Rhabdosphaeraceae Haeckel, 1894 *** Order †Watznaueriales Bown 1987 **** Family †Watznaueriaceae Rood, Hay & Barnard 1971 *** Order †Arkhangelskiales Bown & Hampton 1997 **** Family †Arkhangelskiellaceae Bukry 1969 **** Family †Kamptneriaceae Bown & Hampton 1997 *** Order †Podorhabdales Rood 1971 **** Family †Axopodorhabdaceae Wind & Wise 1977 **** Family †Biscutaceae Black, 1971 **** Family †Calyculaceae Noel 1973 **** Family †Cretarhabdaceae Thierstein 1973 **** Family †Mazaganellaceae Bown 1987 **** Family †Prediscosphaeraceae Rood et al., 1971 **** Family †Tubodiscaceae Bown & Rutledge 1997 *** Order Coccolithales Schwartz 1932 **** Family Reticulosphaeraceae Cavalier-Smith 1996 **** Family Calcidiscaceae Young & Bown 1997 **** Family Coccolithaceae Poche 1913 emend. Young & Bown, 1997 **** Family Pleurochrysidaceae Fresnel & Billard 1991 **** Family Hymenomonadaceae Senn 1900
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