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Chordates Echinoderms Molluscs Segmented worms Bryozoans Arthropods Cnidarians Sponges Brachiopods Deuterostomes (anus forms before mouth) Coelomates (true body cavity) Animals with nerve and muscle cells Fossils &animals Evolution—Cnidaria Multicellular 1 Cnidarians—Stuff to know • All bold font morphologic terms in text • Classification and stratigraphic ranges of paleontologically important groups • Skeletal mineralogy • Septal insertion patterns in rugosans and scleractinians • Hermatypic vs. ahermatypic ecology Fossils & Evolution—Cnidaria 2 Cnidarians—Be able to identify: • Order Scleractinia – Genus Diploria; Genus Montastrea; Genus Dichocoenia • Order Rugosa – Genus Hexagonaria; Genus Pachyphyllum • Order Tabulata – Genus Favosites; Genus Halysites; Genus Aulopora Fossils & Evolution—Cnidaria 3 Cnidaria—Phylum overview • Colonial and solitary invertebrates • Examples include hydroids, jellyfish, sea anemones, corals • Two body layers (ectoderm and endoderm) separated by middle, non-cellular (“jelly”) layer (mesogleoa) • No coelom (no true body cavity) • No organs • Primary radial symmetry • Possess specialized stinging structures (nematocysts) Fossils & Evolution—Cnidaria 4 Cnidaria—Phylum overview (cont.) • Body is a polyp (mouth up) or medusa (mouth down) • Digestive system is a central mouth that leads to a digestive cavity (enteron) • Mouth may be surrounded by tentacles • Muscle cells and nerve cells facilitate simple movements Fossils & Evolution—Cnidaria 5 Basic body forms Fossils & Evolution—Cnidaria 6 Cnidaria—Phylum overview (cont.) • Skeleton may be absent, internal, or external – If present, organic or calcareous • • • • Aquatic (fresh and marine) Suspension feeders Sessile, planktonic, or nektonic Stratigraphic range is Late Proterozoic (Ediacaran) to Recent Fossils & Evolution—Cnidaria 7 Classification • Class Hydrozoa (“hydroids,” unimportant as fossils) • Class Scyphozoa (jellyfish, unimportant as fossils) • Class Anthozoa (true corals and others) – Exclusively marine – Polyp stage only; no medusa – Free-swimming larvae Fossils & Evolution—Cnidaria 8 Cnidaria classification Phylum Cnidaria Class Hydrozoa (unimportant) Class Scyphozoa (unimportant) Class Anthozoa (corals et al.) Subclass Octocorallia (relatively unimportant) Subclass Zoantharia Order Tabulata (tabulates) Early Ordovician-Permian Order Rugosa (rugose) Middle Ordovician-Permian Order Scleractinia (modern corals) Middle Triassic-Recent Note: Permian scleractinian-like forms are now known Fossils & Evolution—Cnidaria 9 Skeletal morphology • Coral skeletons are external and calcareous – Aragonite or calcite • Skeleton is secreted by the epidermis at the base of the polyp • Skeleton consists of basal plate, radial septa, and outer wall (theca) • As skeleton grows upward, new basal plates may be added – Tabulae (transverse plates) – Dissepiments (smaller, curved plates) Fossils & Evolution—Cnidaria 10 Polyp and skeleton Fossils & Evolution—Cnidaria 11 Skeletal morphology (cont.) • Polyp occupies the calice, the part of the skeleton above the last-formed tabula or dissepiments • Skeleton of one coral (solitary or colonial) is a corallum • Skeleton of one polyp in a colony is a corallite • Skeletal tissue between corallites in a colony is coenosteum Fossils & Evolution—Cnidaria 12 Skeletal morphology (cont.) Fossils & Evolution—Cnidaria 13 Skeletal morphology (cont.) Fossils & Evolution—Cnidaria 14 Skeletal morphology (cont.) Fossils & Evolution—Cnidaria 15 Septal insertion • Tabulates lack septa or possess only minor septa • Order of septal insertion is the most important aspect of classification in the rugose corals and scleractinians Fossils & Evolution—Cnidaria 16 Septal insertion in Rugose corals • First six septa are “protosepta” – Stage 1: cardinal and counter septa – Stage 2: alar septa (on either side of cardinal septum) – Stage 3: counterlateral septa (on either side of counter septum) • All subsequent septa (metasepta) are added on either side of cardinal septum and on counter side of alar septa • Septa cluster into four quadrants hence, “Tetracorals” Fossils & Evolution—Cnidaria 17 Septal insertion in Rugose corals Six protosepta x = cardinal sector y = alar sector Fossulae = gaps between sectors 18 Fossils & Evolution—Cnidaria Septal insertion in scleractinians • Stage 1: six protosepta • Stages 2 and higher: metasepta added in the center of spaces between existing septa – Metasepta added in groups of 6, 12, 24, 48, etc. Fossils & Evolution—Cnidaria 19 Septal insertion in scleractinians Fossils & Evolution—Cnidaria 20 Coral evolution • Among the common corals, tabulates (Early Ordovician-Permian) were first to originate • Rugose corals (Middle Ordovician-Permian) might have evolved from tabulates or they might have a separate ancestor • Scleractinians might have evolved from rugosans (?), or from a naked sea anemone – – – – Permian “scleractinian-like” forms are known Late Paleozoic aragonitic rugosans are known No Early Triassic corals are known Pattern of septal insertion is quite different in rugosans and scleractinians Fossils & Evolution—Cnidaria 21 Coral evolution Fossils & Evolution—Cnidaria 22 Coral ecology and reefs • Hermatypic = reef corals that possess zooxanthellae (symbiotic algae) – Shallow, tropical water (25–29°C; < 90 m depth) – Rapid skeletal growth – Oligotrophic (low nutrient) environments • Ahermatypic = non-reef corals without zooxanthellae – Wide environmental range (all latitudes) – Up to 6000 m depth; down to 1°C temp Fossils & Evolution—Cnidaria 23