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Bio 370 Lophophorates

BiologyMetazoan phylogenies. 370 (A) The traditional phylogeny based on and , adapted 1 from Hyman (11). (B) The new molecule-based phylogeny. A conservative approach was taken in B: i.e., some datasets provide resolution within some of the unresolved multifurcations displayed, but we have limited the extent of resolution displayed to that solidly provided by rRNA only. http://www.pnas.org/content/97/9/4453.full

Lophophorates • • 3 suspension-feeding Phyla possessing a : a ring of ciliated , used for suspension feeding. • The tentacles contain extensions of the () • The mouth is inside of the ring, the is outside.

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Lophophore forms

Mouth Anus Tentacles

Lophophorate phyla

• Phoronida (tube-dwelling marine worms) • Brachiopoda (marine bivalves, mainly but some modern) • (=Ectoprocta) marine and freshwater, colonial

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Phylum Phoronida Small group of sessile, tubicolous, marine lophophorates

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Phoronids, cont.

• Worldwide distribution, marine . • Secreted tube of lines the burrow. • Morphologically convergent with tubicolous

Phoronopsis up to 12 inches (most are much smaller)

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Phylum Brachiopoda

• Only ~380 living , but >30,000 Paleozoic species have been described • Sessile, bivalve suspension feeders, superficially similar to bivalve molluscs • Abundant and diverse - important stratigraphic indicators

Brachiopods, continued

• Shells are divided dorso-ventral, with posterior hinge • Attachment to substrate via pedicle (stalk)

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Don’t confuse with clams- shells are dorsal and ventral rather than left and right.

Brachiopoda () Dorsal valve Ventral valve Left valve Right valve

Attachment to substrate via usually burrowing pedicle through ventral valve

Brachiopods, continued Inarticulata • Appeared and radiated early, includes modern and Glottidia

• Shells usually of (CaPO4), not hinged, no brachidium Class • Most extinct and living Brachiopods are in this class, eg. • Hinged shell of , opened by diductor muscles, closed by adductor muscles • Lophophore support (brachidium) on dorsal valve • Lack anus

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Inarticulata Lingula– a living

Living articulate brachiopods

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An articulate brachiopod Dorsal valve with brachidia

Ventral valve with pedicle opening

Articulate brachiopod fossils from Kansas: Phricodothyris, Derbyia, , Hustedia, and

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More Brachiopod lore • Shell includes chitin • Unlike the other lophophorates, brachiopods have no • The inarticulate brachiopod Lingula is considered to be a - little changed in over 400 million years. • Lingula is common enough to be used as food in Thailand (“worms and dirty fingernails”)

Lophophorate phyla

• Phoronida (tube-dwelling marine worms) • Brachiopoda (marine bivalves, mainly Paleozoic but some modern) • Bryozoa (= Ectoprocta) marine and freshwater, colonial

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Bryozoa (Ectoprocta) • ~5,000 living species, many more extinct • Mainly clonal colonies of interconnected individuals (). • colonies range from millimeters to meters in size, but the zooids seldom exceed 1 mm. • Reproduction – , statoblasts, sexual • bryozoans produce bryostatin, currently under testing as an anti-cancer and memory- enhancing drug. link

Bryozoan body plan Reproduction by budding yields of connected individuals (zooids) Anus outside the lophophore (“ectoproct”) Calcified in many taxa Each has its own compartment & operculum

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Bryozoan taxa • (“guarded throat”): Freshwater, no zooid polymorphism, no ; form statoblasts (resting buds). • (“narrow throat”): Marine bryozoans with tubular zooids with calcified walls. • (“naked throat”): Mostly marine with cylindrical or flattened zooids in calcified . Includes most living bryozoans.

Gymnolaemata

operculum

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Some Bryozoa have polymorphic zooids Note the defensive avicularium

Phylactolaemata Freshwater bryozoan

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Phylactolaemata magnifica

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Zooids of Pectinatella

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Statoblasts- resting buds produced for dispersal and overwintering

Hooks may facilitate dispersal on wading birds, mammals or or insects

Archimedes An extinct bryozoan that formed spiral colonies (Mississippian period)

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Fossil bryozoans in chert from the Burlington formation (Mississippian period) in Greene County

Everlasting air … the wonder • Dried, painted marine bryozoan sold as a plant • NO care needed- no soil, no water!

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