Herbarium specimen of Gastridium articulatum, from University of California, Berkeley.

Belongs within: Eukaryota.
Contains: Vendophyceae, Hacrobia, Harosa, Glaucophyta, Rhodophyceae, Chlorophyta, Streptophyta.

The Corticata is a clade of eukaryotes supported by molecular phylogenies that might be described as comprising the major algal lineages and related taxa. Cavalier-Smith et al. (2015) defined this clade as including all descendents of the common ancestor of Arabidopsis and Paramecium. Many members of this clade are photosynthetic; members of the subclade Plantae possess chloroplasts with chlorophyll a as the primary pigment that is believed to have arisen through a single primary endosymbiosis with a cyanobacterium. Photosynthetic members of the sister subclade, labelled Chromista by Cavalier-Smith (2010), usually have a chlorophyll c-containing chloroplasts that arose through secondary endosymbiosis with another photosynthetic eukaryote; debate continues, however, as to whether there was a single such event near the base of the chromist clade (followed by multiple chloroplast losses) or whether the various photosynthetic chromist lineages acquired their chloroplasts independently.

Fossil macroscopic algae have also been assumed herein to belong within the Corticata, though the true relations of groups such as Vendophyceae and Chuariaphyceae to modern multicellular algae remain uncertain.

Characters (from Cavalier-Smith et al. 2015): Eukaryotes ancestrally with ventral feeding groove and cytoskeleton comprising single dorsal centriolar microtubular root and dorsal fan, posterior split right root with I fibre and left root with C fibre, and intermediate single root; cortical alveoli often present; ciliary hairs often not associated with latticed paraxonemal rod.

<==Corticata [Biliphyta, Chromobiota, Chromophycophyta, Chromophyta, Diaphoretickes, Lecaniellaceae, Metaphyta] C-SCL15
    |--Chromista [Chromalveolata] C-S10
    |    |--Hacrobia C-S10
    |    `--Harosa C-S10
    `--Plantae [Archaeplastida] C-SCL15
         |--Glaucophyta C-S10
         `--+--Rhodophyceae C-S10
            `--Chloroplastida (see below for synonymy) AS12
                 |  i. s.: Haematococcus pluvialis OV01
                 |         Scotiellopsis oocystiformis OV01
                 |         Pterosphaera Jörgensen 1900 FT93
                 |--Chlorophyta L-BC03
                 `--Streptophyta L-BC03

Corticata incertae sedis:
  Thallophyca Zhang 1989 G03
    |--T. corrugata Zhang & Yuan 1992 X04
    `--T. ramosa Zhang 1989 X04
  Vendophyceae G03
  Bitelaria Istchenko & Istchenko 1979 G03
  Himanthaliopsis Zalessky 1915 G03
  Sarcinophycus Xiao & Knoll 2000 X04
    |--*S. radiatus Xiao & Knoll 2000 X04
    `--S. papilloformis Xiao 2004 X04
  Gastridium RV01
    |--G. articulatum RV01
    |--G. obtusum RV01
    |--G. pinnatifidum RV01
    |--G. repens RV01
    `--G. tenuissimum RV01
  ‘Archidesmus’ loganensis Peach 1899 WA04
  Scytothalia dorycarpa G05
  Caulocystis uvifera G05
  Ptilocladia pulchra G05
  Kyrtuthrix maculans K98
  Penicillus capitatus HS98
  Ahnfeltiopsis flabelliformis KY97
  Buthotrephis Hall 1847 S00, H75 [=Bythotrephis (l. c.) H75]
    `--B. rebskei S00
  Koninckopora Lee 1912 [incl. Coeloceratoides Derville 1931] T64
    `--*Coeloceratoides’ fragilis Derville 1931 T64
  Yuknessia simplex MSF05
  Eyrea Cookson & Eisenack 1971 E77
    `--E. nebula E77
  Landsburgia quercifolia L49
  Digenea simplex P75
  Delesserites Sternberg 1833 [incl. Delesserella Ruedemann 1926, Delessertites Bronn 1853] H75
    |--*D. lamourouxii (Brongniart 1823) [=Fucoides lamourouxii] H75
    `--D. salicifolia Ruedemann 1925 H75
  ‘Zonarites’ digitatus Sternberg 1833 H75
  Aataenia Gnilovskaya 1976 G79
    `--*A. reticularis Gnilovskaya 1976 G79
  Lanceoforma Walter, Oehler & Oehler 1976 G79
    `--*L. striata Walter, Oehler & Oehler 1976 G79
  Proterotainia Walter, Oehler & Oehler 1976 G79
    `--*P. montana Walter, Oehler & Oehler 1976 G79
  Prototaxites S00
    |--P. hefteri (Schaarschmidt) Schweitzer 1983 [=Mosellophyton hefteri Schaarschmidt 1974] S00
    `--P. psygmophylloides Kräusel & Weyland 1930 S00
  Archaelagena Howchin 1888 [=Archealagena (l. c.)] LT64
    `--*A. howchiniana (Brady 1876) [=Lagena howchiniana] LT64
  Cellulina Zborzewski 1834 LT64
  Coelotrochium Schlüter 1879 LT64
  Testamoebiformia LT64
    |--Cysteodictyina Carter 1880 LT64
    |    `--*C. compressa Carter 1880 LT64
    `--Holocladina Carter 1880 LT64
         `--*H. pustulifera Carter 1880 LT64
  Goniolina d’Orbigny 1850 LT64
    `--*G. hexagona d’Orbigny 1850 LT64
  Pseudogypsina Trauth 1918 LT64
    `--*P. multiformis Trauth 1918 LT64
  Siphonema Bornemann 1886 LT64
  Chuariaphyceae HR94
    |--Beltina Walcott 1899 [Beltinaceae] HR94
    |    `--*B. danai Walcott 1899 HR94
    `--Chuaria Walcott 1899 H98 (see below for synonymy)
         |--*C. circularis Walcott 1899 H98
         |--*Krishnania’ acuminata Sahni & Shrivastava 1954 H62
         |--‘Protobolella’ jonesi Chapman 1935 M65
         |--*Fermoria’ minima Chapman 1935 [=*Protobolella minima] H75
         |--C. pendjariensis H98
         `--C. wimani H75

Trace fossils: Delesserites foliosus Ludwig 1869 H75
               Delesserites gracilis H75
               Delesserites sinuosus H75

Chloroplastida [Chlorobionta, Chlorobiota, Chlorophycophyta, Chlorophytina, Euchlorophyceae, Euchlorophyta, Prasinomonadida, Prasinophyceae, Prasinophyta, Viridiplantae] AS12

Chuaria Walcott 1899 H98 [incl. Fermoria Chapman 1935 G79, Krishnania Sahni & Shrivastava 1954 G79, Protobolella Chapman 1935 G79, Vindhyanella Sahni 1936 G79; Chuariaceae, Chuariales H94]

*Type species of generic name indicated


[AS12] Adl, S. M., A. G. B. Simpson, C. E. Lane, J. Lukeš, D. Bass, S. S. Bowser, M. W. Brown, F. Burki, M. Dunthorn, V. Hampl, A. Heiss, M. Hoppenrath, E. Lara, E. Le Gall, D. H. Lynn, H. McManus, E. A. D. Mitchell, S. E. Mozley-Stanridge, L. W. Parfrey, J. Pawlowski, S. Rueckert, L. Shadwick, C. L. Schoch, A. Smirnov & F. W. Spiegel. 2012. The revised classification of eukaryotes. Journal of Eukaryotic Microbiology 59 (5): 429–493.

[C-S10] Cavalier-Smith, T. 2010. Kingdoms Protozoa and Chromista and the eozoan root of the eukaryotic tree. Biology Letters 6 (3): 342–345.

[C-SCL15] Cavalier-Smith, T., E. E. Chao & R. Lewis. 2015. Multiple origins of Heliozoa from flagellate ancestors: new cryptist subphylum Corbihelia, superclass Corbistoma, and monophyly of Haptista, Cryptista, Hacrobia and Chromista. Molecular Phylogenetics and Evolution 93: 331–362.

[E77] Elsik, W. C. 1977. Paralecaniella indentata (Defl. & Cooks. 1955) Cookson & Eisenack 1970 and allied dinocysts. Palynology 1: 95–102.

[FT93] Fensome, R. A., F. J. R. Taylor, G. Norris, W. A. S. Sarjeant, D. I. Wharton & G. L. Williams. 1993. A classification of living and fossil dinoflagellates. Micropaleontology Special Publication 7: i–viii, 1–351.

[G79] Glaessner, M. F. 1979. Precambrian. In: Robison, R. A., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt A. Introduction. Fossilisation (Taphonomy), Biogeography and Biostratigraphy pp. A79–A118. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[G03] Gnilovskaya, M. B. 2003. The oldest tissue differentiation in Precambrian (Vendian) algae. Paleontologicheskii Zhurnal 2003 (2): 92–98 (transl. Paleontological Journal 37 (2): 196–204).

[G05] Goldberg, N. A. 2005. Temporal variation in subtidal macroalgal assemblages at Black Island, Recherche Archipelago. Journal of the Royal Society of Western Australia 88 (2): 65–71.

[H98] Haines, P. W. 1998. Chuaria Walcott, 1899 in the lower Wessel Group, Arafura Basin, northern Australia. Alcheringa 22 (1): 1–8.

[H62] Häntzschel, W. 1962. Trace fossils and problematica. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt W. Miscellanea: Conodonts, Conoidal Shells of Uncertain Affinities, Worms, Trace Fossils and Problematica pp. W177–W245. Geological Society of America, and University of Kansas Press.

[H75] Häntzschel, W. 1975. Treatise on Invertebrate Paleontology pt W. Miscellanea Suppl. 1. Trace Fossils and Problematica 2nd ed. The Geological Society of America: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[HS98] Hara, I., & T. Sakurai. 1998. Isolation and characterization of vanadium bromoperoxidase from a marine macroalga, Ecklonia stolonifera. Journal of Inorganic Biochemistry 72: 23–28.

[HR94] Hofmann, H. J., & R. H. Rainbird. 1994. Carbonaceous megafossils from the Neoproterozoic Shaler Supergroup of Arctic Canada. Palaeontology 37 (4): 721–731.

[K98] Kaehler, S. 1998. The non-coralline epilithic encrusting algae of Hong Kong II: additions and identification. Asian Marine Biology 15: 1–17.

[KY97] Kogame, K., & Y. Yamagishi. 1997. The life history and phenology of Colpomenia peregrina (Scytosiphonales, Phaeophyceae) from Japan. Phycologia 36 (5): 337–344.

[L49] Lindauer, V. W. 1949. Additions to the marine algae of New Zealand. Transactions and Proceedings of the Royal Society of New Zealand 77 (3): 390–393.

[LT64] Loeblich, A. R., Jr & H. Tappan. 1964. Sarcodina: chiefly “thecamoebians” and Foraminiferida. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt C. Protista 2 vol. 2. The Geological Society of America and The University of Kansas Press.

[L-BC03] López-Bautista, J. M., & R. L. Chapman. 2003. Phylogenetic affinities of the Trentepohliales inferred from small-subunit rDNA. International Journal of Systematic and Evolutionary Microbiology 53: 2099–2106.

[MSF05] Maletz, J., M. Steiner & O. Fatka. 2005. Middle Cambrian pterobranchs and the Question: what is a graptolite? Lethaia 38: 73–85.

[M65] Moore, R. C. (ed.) 1965. Treatise on Invertebrate Paleontology pt H. Brachiopoda vol. 2. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas Press: Lawrence (Kansas).

[OV01] Orosa, M., J. F. Valero, C. Herrero & J. Abalde. 2001. Comparison of the accumulation of astaxanthin in Haematococcus pluvialis and other green microalgae under N-starvation and high light conditions. Biotechnology Letters 23: 1079–1085.

[P75] Por, F. D. 1975. Pleistocene pulsation and preadaptation of biotas in Mediterranean seas: consequences for Lessepsian migration. Systematic Zoology 24 (1): 72–78.

[RV01] Redhead, S. A., R. Vilgalys, J.-M. Moncalvo, J. Johnson & J. S Hopple Jr. 2001. Coprinus Pers. and the disposition of Coprinus species sensu lato. Taxon 50: 203–241.

[S00] Schweitzer, H. J. 2000. Neue Pflanzenfunde im Unterdevon der Eifel (Deutschland). Senckenbergiana Lethaea 80 (2): 371–395.

[T64] Teichert, C. 1964. Doubtful taxa. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt K. Mollusca 3. Cephalopoda—General Features—Endoceratoidea—Actinoceratoidea—Nautiloidea—Bactritoidea pp. K484–K490. The Geological Society of America and the University of Kansas Press.

[WA04] Wilson, H. M., & L. I. Anderson. 2004. Morphology and taxonomy of Paleozoic millipedes (Diplopoda: Chilognatha: Archipolypoda) from Scotland. Journal of Paleontology 78 (1): 169–184.

[X04] Xiao, S. 2004. New multicellular algal fossils and acritarchs in Doushantuo chert nodules (Neoproterozoic; Yangtze Gorges, south China). Journal of Paleontology 78 (2): 393–401.

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