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Rugosa

The Rugosa are an extinct class of solitary or colonial corals that were abundant in Middle Ordovician to Late Permian seas.

Morphology
Each polyp of a rugose coral is hosted on a corallite, the fundamental skeletal structure of the coral. Unlike most living corals, many rugosan species live a solitary life, with a relatively large polyp (and corresponding corallite) surviving on its own. These horn-shaped solitary species develop from a pointed tip (apex) up towards the polyp-bearing cup (calice). Other rugosan species have polyps which grow together as a colony, with the entire colony's skeleton known as a corallum. Axial structures Rugose corals often have a columella, a dense rod running up the center (axis) of the corallite. It is common in rugose corals because they were mainly solitary, and so required the extra support. Tabulate corals have no columella because they were always colonial and relied on the support of neighboring corallites. Alternatively, the center of the corallite may be supported by the aulos, a tube filled with tabulae. Colony forms Colonial rugose corals can show many different growth forms in their corallum (entire colony structure): • Fasciculate corals have their corallites joined at the base, but the edge of each corallite is not connected, leaving a gap of open water between each. • Dendroid corals have a branching bush-like system of corallites. • Phaceloid corals have corallites running in parallel from a common base. Some phaceloid corals have outgrowths bridging between the gaps. • Massive corals have no gaps between their corallites. • Cerioid corals retain walls between adjacent corallites. Petoskey stones (Hexagonaria) are an example of cerioid corals. • Astreoid corals have no walls between adjacent corallites, but each corallite retains its own set of septa which contact the septa of adjacent corallites. • Thamnasteroid corals have no walls, and septa are shared equally between adjacent corallites. • Aphroid corals have no walls, and septa are completely isolated from each other within a colony-wide mesh of dissepiments. == Taxonomy ==
Taxonomy
Taxonomy to the suborder level, mostly based on Treatise on Invertebrate Paleontology (Part F, 1981) (Early Permian – Middle Permian) • Suborder Ketophyllina Zhavoronkova, 1972 (Late Ordovician – Late Devonian) • Suborder Lithostrotionina Spasskiy & Kachanov, 1971 (Mississippian – Late Permian) • Suborder Lonsdaleiina Spasskiy, 1974 (Mississippian – Late Permian) • Suborder Lycophyllina Zhavoronkova, 1972 (Late Ordovician – Middle Devonian) • Suborder Metriophyllina Spasskiy, 1965 (Middle Ordovician – Late Permian) • Suborder Plerophyllina Sokolov, 1960 (Late Silurian – Late Permian) • Suborder Ptenophyllina Wedekind, 1927 (Silurian – Late Devonian) • Suborder Stauriina Verrill, 1865 (Middle Ordovician – Mississippian) • Suborder Stereolasmatina Hill, 1981 (Early Devonian – Late Permian) • Suborder Streptelasmatina Wedekind, 1927 (Middle Ordovician – Late Devonian) • Suborder Tachylasmatina Fedorowski, 1973 (Early Devonian – Late Permian) == Evolutionary history ==
Evolutionary history
Ordovician Rugosans likely originated from a non-skeletal anthozoan similar to modern sea anemones. The oldest rugosan fossils appear near the end of the Middle Ordovician, during the transition from the Darriwilian stage to the Sandbian stage at the start of the Late Ordovician. Lambelasma, a calostyline from Darriwilian-age Iran, is the earliest confirmed example. By the end of the Sandbian stage, rugose corals were common in the shallow seas of North America and Baltoscandia. Five rugosan suborders filled out the reef-building faunas of the Late Ordovician: Cystiphyllina, Calostylina, Stauriina, Streptelasmatina, and Metriophyllina. Cystiphyllines, streptelasmatines, and metriophyllines were usually solitary, stauriines were colonial, and calostylines included many species of both solitary and colonial corals. Some early rugosans were very simple, such as Primitophyllum (a cystiphylline) and Lambeophyllum (a calostyline). These simple forms resembled tiny cones with rudimentary septa and no tabulae, and they may be a good estimate for an ancestral rugosan. Others were more typical, such as the solitary Streptelasma (a streptelasmatine) and colonial Favistina (a stauriine). Silurian The Ordovician suborders all persisted into the Silurian, though newer groups achieved even greater abundance. The Arachnophyllina, Ketophyllina, and Lycophyllina diversify quickly, while others (the Columnariina, Cyathophyllina, and Ptenophyllina) slowly take hold. Calostylines diminish and go extinct near the Silurian-Devonian boundary. Lycophyllines were usually solitary, while arachnophyllines, columnariines, and ptenophyllines were usually colonial. Cyathophyllines and ketophyllines included many solitary and colonial species. Devonian The Early Devonian was the apex of diversity for rugosans and many other marine animals. The Middle to Late Devonian, however, saw a series of extinction events which profoundly reshaped rugosan faunas. Arachnophyllines, cystiphyllines, lycophyllines, and streptelasmatines all die out near the start of the Late Devonian. Others (ketophyllines, columnariines, and ptenophyllines) last longer, though they too would vanish by the end of the period. However, the Devonian was also the origin of a few more rugosan suborders: Plerophyllina and Stereolasmatina, both of which were small and solitary. Carboniferous-Permian Rugosans recovered from extinction quickly in the Mississippian (early Carboniferous). Plerophyllines and stereolasmatines lead the charge, alongside a substantial diversification of four new suborders: Aulophyllina, Caniniina, Lithostrotionina, and Lonsdaleiina. Aulophyllines and caniniines were usually large and solitary, while lithostrotionines and lonsdaleiines were usually colonial. An extinction at the end of the Mississippian eliminated the last stauriines and cyathophyllines, while the other suborders survived at a slightly reduced capacity through the Pennsylvanian (late Carboniferous). Rugosans experienced a few small extinctions within the Permian period, but on the whole they were still abundant reef builders until the entire class died out during the Permian-Triassic mass extinction. == Gallery ==
Gallery
File:RugosaOrdovician.jpg|Three views of Grewingkia canadensis, a solitary streptelasmatine from Ordovician Indiana File:Cyathophylloides sp. (colonial coral).jpg|Cyathophylloides sp., a colonial stauriine from the Late Ordovician, Georgian Bay Formation, Ontario, Canada File:Streptelasma divaricans (Nicholson, 1875).jpg|Streptelasma divaricans (Nicholson, 1875), a solitary streptelasmatine from the Liberty Formation (Late Ordovician) of southern Ohio File:Acervularia-ananas gotland.jpg|Acervularia, a colonial columnariine from Silurian Sweden File:The Childrens Museum of Indianapolis - Polished Petoskey stone.jpg|A polished Petoskey stone (Hexagonaria, a colonial columnariine from Devonian Michigan) File:Zaphrentis phrygia fossil rugose coral (Jeffersonville Limestone, Middle Devonian; Falls of the Ohio, southern Indiana, USA) 4 (14968944624).jpg|Zaphrentis, a solitary cyathophylline from Middle Devonian Indiana, looking down onto the calyx. File:Stereolasma cross section.jpg|Cross-section of Stereolasma rectum, a solitary stereolasmatine from the Middle Devonian of Erie County, New York File:Fossilised Coral near Ogmore-by-Sea in Wales 2.jpg|Solenodendron, a colonial lithostrotionine from Mississippian Wales, seen from below == References ==
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