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Class Anthozoa-Corals and Sea Anemones, Sea Fans, Sea Pansies Diploria labyrinthoformis Paragorgia

Heteractis magnifica

Largest Class (> 6000 spp.), 3 Subclasses:

1) Hexacorallia Anemones and true corals 2) Octocorallia Soft corals and sea fans 3) Ceriantipatharia Black Corals and Tube Anemones

Diploria spp.

Exclusively marine, solitary or colonial polypoids with no medusa stage Polyps have internal cavity divided by septa Pharynx penetrates well into G-V cavity and bears 1 or more ciliated grooves (siphonoglyphs) Cnidae epidermal and gastrodermal Margins of mesenteries or septa have cnidocytes Gonads gastrodermal 8 tentacles usually, or multiples of 6

Plexaura homomallia

Gorgonia ventalina

tube anemone

Cirrhipathes anguina

Sea Anemones
Large, solitary polyps, brightly colored Oral disk may bear many tentacles Slit shaped mouth with ciliated groove (siphonoglyph) at one end Allows water to enter G-V cavity to maintain hydrostatic pressure Polyp can contract and withdraw under muscular collar when threatened or inactive Septa increase internal surface area for digestion, gas exchange Mesoglea is much thicker than in Hydrozoans, partly cellular

Spirocysts in gastrodermis-sticky cnidocytes Cnidocytes may be used aggressively against other anemones

Metridium giganteum

Asexual fission of various forms Hermaphroditic but production of male and female gametes asynchronous so little self fertilization Fertilization often inside with variable development before release of larva Usually planktonic planula which settles and grows tentacles

Longitutidal (A) and cross section (B) of a sea anemone

Anthozoan Reproduction

Habitat utilization of reef-dwelling sea anemones

Stony Corals (Madreporarian) Grow in calcareous cups with sclerosepta-skeleton completely outside polyp (for some species) Colonial and usually interconnected Polyps are usually quite small Foundations of coral reefs Most have zooxanthellae which provide nutrients Polyps are similar to anemones except lack siphonoglyphs

Carbonate cups have sclerosepta that help to anchor polyps Carbonate is laid down in bands because of variation in rate of deposition Allows for aging of the skeleton Coral polyps feed in very similar way to anemones except that continuous bed of polyps provides a wide mesh for prey capture Coral polyps of different genotypes compete with each other for limited attachment substrate Secrete allelochemicals to kill neighbors

Octocorals - Sea Pens, sea Pansies, Sea Fans, Pipe Corals Colonial polyps Always with 8 tentacles, each of which is pinnate or with side branches Polyps joined by connective tissue called coenenchyme with gastrodermal tubes Skeleton is internal-individual spicules or fused mass- may be rigid or flexible Sea Fans oriented perpendicular to current-can bend Few cnidocytes so probably feed on small particles - not zooplankton Reproduction is similar to other anthozoans

Cnidarian fossils from the Ediacaran fauna

Fossils are both medusoid and polypoid, and probably represent all cnidarian classes (Scrutton 1979).

From Brusca and Brusca (2002)

Note: The extinct class Conulata is probably related to Scyphozoa

Competing hypotheses (Figs 1A,B) of cnidarian life-cycle evolution and systematic relationships (Bridge et al. 1995.)

H1: Anthozoans are the most basal (Polyp is the ancestral and medusa is the derived condition)

H2: Anthozoans are the most derived (medusa is the ancestral condition but has been lost in Anthozoa. Polyps evolved secondarily)

Morphological, mtDNA, and 18S rDNA data separately and combined support the basal position of Anthozoa but do not resolve the relationships among Scyphozoa, Cubozoa and Hydrozoa (Bridge et al. 1995).

Phylogeny of Cnidaria according to the medusa theory

Gordon Medusa
1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. Polypoid body form and alteration of generations Gonads relocated in gastrodermis Partitions (mesenteries) appear, subdividing the coelenteron Polyp stage secondarily lost or reduced Evolution of the rhopalium Evolution of strobilation Evolution of boxlike medusa body Complex lensed rhopalial eyes Velarium Suppression of the medusoid stage Hexaradial and octaradial symmetry Actinopharynx Siphonoglyph Coelenteron acquires mesenterial filaments Loss of operculum Loss of cnidocil Evolution of tripartite series of flaps on cnidae Evolution of special ciliary cones associated with cnidae

Cnidarian synapomorphies: radial, medusoid body; cnidae; planula larva; cnidarian coelenteron.

Phylogeny of Cnidaria according to the polyp theory (Brusca and Brusca 2002)

1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13.

Cnidocil + loss of ancestral anthozoan traits Polyp stage lost or reduced Acraspedote medusa Evolution of the rhopalium Evolution of strobilation Evolution of boxlike medusa body Complex lensed rhopalial eyes Velarium Relocation of gonad-forming tissue to epidermis Loss of gut mesenteries Simplification of the middle layer to an acellular mesoglea Evolution of the craspedote medusa form Loss of gastrodermal nematocysts