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Biology 222 Invertebrate Zoology II

Fall 2004Guest Lecturer: Helen DyerNorthern Arizona University

Lectures 6-7

PHYLUM ANTHOZOA

PHYLUM MOLLUSCA

PHYLUM ANNELIDA

SUPER PHYLUM LOPHOPHORATA

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Phylogeny• Page 32 Figure 2.3 (A)• Originally placed with

Cnidarians• Originally thought to be

Deuterostomes• Now placed with

Protostomes• Still lots of controversy• Final Answer?

Lophophorate Phylogeny

Page 799

Lophophorate Phyla

• General characteristics

• Common character: Lophophore– U-shaped food

gathering structure– 1-2 rows, hollow ,

ciliated tentacles

Phylum Phoronida

Phylum BryozoaPhylum Brachiopoda

Micrograph of Lophophore

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The Lophophore

Physical Characteristics of Lophophorates

• Trimeric– Prosome – Mesosome– Metasome

• Body cavities– Protocoel– Mesocoel– Metacoel

• Protostomes– Schizocoely

• Peritoneum– Distinct from

enterocoely -Deuterostomes

• Sessile suspension feeders– Reduced cephalization– U-Shaped gut– Nervous system

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• Previously classified as Deuterostomes– Radial determinate

cleavage– Enterocoeluous

coelom formation• Molecular Phylogeny

– Doesn’t fit• Haeckel’s Maxim

– 3 major phyla

Phylum Phoronida

• The Phoronids• Worm-like tube dwellers• Two Genera

– Phoronis– Phoronopsis

• 12 species• Intertidally to 400+m

– Common at 70m

• 0.5 – 50cm long• Fossil Record Absent

General Characteristics

Page 773

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Physiology

Page 774 Figure 21.1

Reproduction• Asexual

– Transverse fission

• Gonochoristic or Hermaphroditic

• Sexual structures– Form in lophophoral organs

• Near lophophore– Form nidamental glands in

females– Spermatophoral glands in

males

• Internal fertilization in P. harmeri– Spermatophores

released• Captured on female

lophophores

– Amoeboid sperm• To lophophore coelom

– Travel through meso and metacoel to eggs

• Fertilization occursPage 777 Figure 21.3

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Early Development

Page 778 Figure 21.4

Actinotroch larvae

Phylogeny and Systematics• Widespread• Earliest possible fossil

from the Jurassic/ Upper Cretaceous– 202mya/145mya– Suspect– Trace Fossil

• Borings– Talpina ramosa– Conchotrema

Composita subtilita

Phoronids a la Carteor

Predators of Phoronids

• Some species of Fish• Gastropods• Nematodes

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Research

• http://www.com.univ- mrs.fr/DIMAR/Phoro/

Phylum Brachiopoda

• The Lamp shells• Resemble bivalves• Highly modified

lophophore• Excellent fossil record

General Characteristics

Page 793Page 792 Figure 21.18

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Physiology

Page 794 Figure 21.19

Water and Nutrient Flow

Musculature

Page 796

Bivalve

Brachiopod

Adductor

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Reproduction

• No Asexual • Most species

Gonochoristic• Fertilization external

– Eggs and sperm shed– Some species brood

embryos• Requires Internal

fertilization

Early Development

• Strong resemblance to deuterostomes

• Larvae metamorphose in Articulates– Not in Inarticulates

• Mantle lobes fold up to enclose visceral mass

Page 797 Figure 21.22

Lobate Larvae

Phylogeny and Systematics

• Widespread and speciose

• Arose in Devonian– 400 mya

• Crash in Permian-Triassic extinction– 250 mya

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Class Inarticulata• Without calcified valves• Order Lingulida – the

Lamp Shells– Lingula– Oldest genus on planet– Virtually identical to

specimens 400 myr

• Order Acrotretida– Crania– Limpet like in form

Class Articulata

• With calcified valves• Order Rhynchonellida

– permanently attached to solid substrates

– often with complete gut

Class Articulata

• Order Terebratulida– often found in fossil

assemblages– Composita,

Anthracospirifer, Cleiothyridina

– Occasionally with movable peduncle, often large

– TerebratellaCompositaComposita

AnthracospiriferAnthracospirifer

CleiothyridinaCleiothyridina

TerebratellaTerebratella

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Brachiopods on the half shellor

Predators of Brachiopods

• Echinoderms• Crustaceans• Gastropods• Cephalopods • Some species of Fish

Research

• International Brachiopod Congress– http://paleopolis.rediris.es/BrachNet/index.htm

Phylum Bryozoa

• Moss animals, Polyzoa, Ectoprocts

• Mostly sessile, modular– Zooids – No larger than 1mm– Colony – mm to m in size

• Retractable tentacles• Excellent fossil record

– Example of punctuated evolution

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• 5000 known extant species

• 16000 known extinct species

• Class Gymnolaemata• Class Stenolaemata• Class

Phylactolaemata

Sessile?

Page 780 Figure 21.5

Cristatella spp.

Cupuladriids spp.

General Characteristics

Page 779

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Physiology

Page 784 Figure 21.9

Autozooids

Heterozoids

AviculariaVibracula and Spines

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Ovicells

Autozooid

OvicellsZooecium

Reproduction

• Hermaphroditic or Gonochoristic

• Asexual – Budding

• Sexual– Eggs and Sperm

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2.1. Page 786 Figure 21.10 2. Page 780 Figure 21.5

Early Development

• Free swimming larvae• Ancestrula• Colony Development

Egg and Larvae

Ancestrula

Colony

Cyphonautes larvae

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Phylogeny and Systematics

• Widespread • First arose Ordovician

– 500mya• Cambrian?

– 570mya• Permian/Triassic

extinction– 250mya

• Identification Problematic

Class Gymnolaemata• Marine• Circular Lophophore• Colonies leathery

– Some Calcification• Box like or cylindrical

zooids• 650 genera• Early Ordovician

Alcionidium gelatinosum

Reteporella spp.

Bugula californica,Bugula spp.

• Order Ctenostomata– Zooids cyndrical to flat– No Ooecia– No Avicularia– Paleozoic to Recent

• Order Cheilostomata– Zooids flat box– Ooecia (brood

chambers)– Avicularia– Mesozoic to Recent

Membranipora membranacea

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Class Stenolaemata

• Marine• Tube shaped zooids• Circular lophophore• Some calcification• Mostly Aborescent

and Fenestrate• 550 Genera

Crisia occidentalis

Tubulipora sp.

Septopora spp.

• 5 Orders– 4 Extinct

• E. Ordovician

• Order Cyclostomata– Zooids cylindrical– Special reproductive

zooids– No Epistome– Paleozoic - Present

Class Phylactolaemata• Fresh Water• Zooids cylindrical• Epistome present• Lophophore

– Horseshoe shape– Large

• Non-Calcareous– chitinous or gelatenous

• Statoblasts• 12 Genera• Mesozoic - Recent

Pectinatella magnifica

Cristatella Cristatella mucedo

Plumatella spp. statoblast

Plumatella spp.

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Bryozoa Under Glassor

Predators of Bryozoans

• Nudibranchs• Gastropods, Chitons• Sea Spiders• Sea Urchins• Flounder, Angel Fish and Puffers

Research

• Bryostatin 1• Myxozoan Parasites – PKD in Salmon• International Bryozoology Association

– http://www.nhm.ac.uk/hosted_sites/iba/

The Lophophorates

• Circular or U-shaped Lophophore– Suspension or Filter feeders

• Reproduction strategies• Don’t need Sunlight to Survive

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Life Styles

Marine Environment

References and Photo Credits• Slide 1• http://eebweb.arizona.edu/collections/Fishes/Photographs.html• http://www.coralreefnetwork.com/stender/diving/california/invert/invert.html• http://www.personal.psu.edu/faculty/s/b/sbj4/aquarium/reef_pictures/• http://www.winterset.net/photos/reef/reef.html• http://www.marinbi.com/bryozoa/• http://www.pznow.co.uk/marine/bivalves.html• Slide 2• http://www.amnh.org/exhibitions/permanent/ocean/03_oceanlife/g3_permia

n.php• http://www.coralreefnetwork.com/stender/diving/california/invert/invert.html• http://www.marinbi.com/bryozoa/• Slide 3• http://www.sportesport.it/plancton03.html• http://www.imagequest3d.com/catalogue/larvalforms/pages/d002_jpg.html• http://www.microscopy-uk.org.uk/micropolitan/index.html

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• Slide 4• Brusca, R.C., Brusca, G.J., 2003, Invertebrates; 2nd edition; Sinaur Assoc, Inc.,

Publishers; Mass., pg. 799• Slide 5• http://www.personal.psu.edu/users/m/x/mxm669/• http://cas.bellarmine.edu/tietjen/Diversity/ectoprocta.html• http://bioweb.uwlax.edu/zoolab/Table_of_Contents/Lab-

6a/Phylum_Ectoprocta_2/Phylum_Ectoprocta_2a/phylum_ectoprocta_2a.html• http://biology.fullerton.edu/courses/biol_404/web/organism.html• Slide 6• Sinaur Associates Inc. Copyright 2001• http://www.lander.edu/rsfox/310images/310bilatImage.html• Slide 7• http://www.lander.edu/rsfox/310images/310LophoImage.html• Slide 8• http://www.lander.edu/rsfox/310images/310bilatImage.html• Slide 9• No Credits• Slide 10• http://www.zoology.uwa.edu.au/staff/rblack/13062001.html• http://virtual.yosemite.cc.ca.us/randerson/Marine%20Invertebrates/phoronid.html

• Slide 11• Brusca, R.C., Brusca, G.J., 2003, Invertebrates; 2nd edition; Sinaur Assoc, Inc., Publishers;

Mass., pg. 773.• Unknown• Slide 12• Brusca, R.C., Brusca, G.J., 2003, Invertebrates; 2nd edition; Sinaur Assoc, Inc., Publishers;

Mass., pg. 774.• Slide 13• http://www.sportesport.it/plancton03.html• Slide 14• Brusca, R.C., Brusca, G.J., 2003, Invertebrates; 2nd edition; Sinaur Assoc, Inc., Publishers;

Mass., pg. 774. • Slide 15• http://scaa.usask.ca/gallery/lacalli/phoronid2.shtml• Slide 16• http://www.com.univ- mrs.fr/DIMAR/Phoro/SYST/PHORONIDA/Map_E.html• Compista subtilita with phoronid borings. Photograph taken by Helen C. Dyer 2002. May only be

used for educational purposes. • Slide 17• No Credits• Slide 18• http://www.com.univ- mrs.fr/DIMAR/Phoro• Slide 19• http://paleo.cortland.edu/tutorial/Brachiopods/brachmorph.html• http://earthsystems.uta.edu/historical_labs/7phylum_brachiopoda.html• Slide 20• Brusca, R.C., Brusca, G.J., 2003, Invertebrates; 2nd edition; Sinaur Assoc, Inc., Publishers;

Mass., pg. 792- 793.

• Slide 21• Brusca, R.C., Brusca, G.J., 2003, Invertebrates; 2nd edition; Sinaur Assoc, Inc., Publishers; Mass., pg. 794• Slide 22• http://www.biology.ualberta.ca/courses.hp/zool250/Labs/Lab12/Lab12.html• Slide 23• http://www.brookdale.cc.nj.us/staff/sandyhook/taxonomy/shells/bivalves.html• Brusca, R.C., Brusca, G.J., 2003, Invertebrates; 2nd edition; Sinaur Assoc, Inc., Publishers; Mass., pg. 796• Slide 24• http://www.imagequest3d.com/catalogue/larvalforms/pages/d002_jpg.html• Slide 25• http://www.imagequest3d.com/catalogue/larvalforms/pages/d002_jpg.html• Brusca, R.C., Brusca, G.J., 2003, Invertebrates; 2nd edition; Sinaur Assoc, Inc., Publishers; Mass., pg. 797.• Slide 26• http://www-personal.umich.edu/~damstra/fossils/brachioreef.html• Rudwick, M.J.S., 1970, Living and Fossil Brachiopods;Hutchinson University Library, London.• Slide 27• http://paleo.cortland.edu/tutorial/Brachiopods/brachmorph.html• http://www.uky.edu/OtherOrgs/KPS/pages/fossilphoto.html• http://users.unimi.it/vertpal/collezione%20paleontologica/brachiopodi/inarticulata/18est.html• http://privat.egersund.com/erling/Armfottinger/default.html• Slide 28• http://eebweb.arizona.edu/collections/Fishes/Photographs.html• http://scilib.ucsd.edu/sio/nsf/fguide/brachiopoda1.html• http://biology.fullerton.edu/courses/biol_404/web/organism.html• Slide 29• http://www.itano.net/fossils/chaffee/brach05.html• http://www.colossal-fossil-site.com/522-list.html• Unknown• Slide 30• No Credits

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• Slide 31• http://paleopolis.refiris.es/BrachNet/index.html• Slide 32• http://people.freenet.de/haeussermann/• Fossil Pennsylvanian Fenestrate Bryozoan. Photograph by Helen C. Dyer 2004. May be used for

educational purposes only.• Slide 33• http://cas.bellarmine.edu/tietjen/Diversity/ectoprocta.html• Unknown• Slide 34• Brusca, R.C., Brusca, G.J., 2003, Invertebrates; 2nd edition; Sinaur Assoc, Inc., Publishers;

Mass., pg. 780• Unknown• Slide 35• http://www.microscopy- uk.org.uk/micropolitan/index.html• Brusca, R.C., Brusca, G.J., 2003, Invertebrates; 2nd edition; Sinaur Assoc, Inc., Publishers;

Mass., pg. 779.• Slide 36• Brusca, R.C., Brusca, G.J., 2003, Invertebrates; 2nd edition; Sinaur Assoc, Inc., Publishers;

Mass., pg. 784.• Slide 37• http://www.microscopy- uk.org.uk/mag/indexmag.html• Slide 38• http://www.microscopy- uk.org.uk/mag/indexmag.html• Slide 39• http://www.microscopy- uk.org.uk/micropolitan/index.html

• Slide 40• Brusca, R.C., Brusca, G.J., 2003, Invertebrates; 2nd edition; Sinaur Assoc, Inc., Publishers; Mass., pg. 780, 786.• Slide 41• http://www.imagequest3d.com/pages/current/pictureoftheweek/bryozoa/bryozoa.html• http://www.gpi.uni-kiel.de/~ae/• Unknown• Slide 42• McKinney, F. K., and Jackson, J.B.C., 1989, Bryozoan Evolution, University of Chicago Press, Chicago. Pg. 15.• Slide 43• http://www3.bc.sympatico.ca/kerryw/creature/spiral.html• http://www.gpi.uni-kiel.de/~ae/index_e.html• http://www.seawater.no/fauna/Mosdyr/nettmosdyr.html• Slide 44• http://www3.bc.sympatico.ca/kerryw/creature/kbryo.html• Unknown• Slide 45• http://www.marinbi.com/bryozoa/• Unknown• Fossil Fenestrate Bryozoan. Photographs by Helen C. Dyer and Ernest H. Gilmour. 2004. May be used for educational purposes only.• Slide 46• Unknown• Slide 47• http://www.civgeo.rmit.edu.au/bryozoa/phylactolaemata/pectinatellidae/pectmag.html• http://www.civgeo.rmit.edu.au/bryozoa/phylactolaemata/cristatellidae/crismuc.html• http://tolweb.org/tree?group=Bryozoa&contgroup=Bilateria• http://www.ct.sakura.ne.jp/~gen-yu/2000/fwbryo.html• Slide 48• Slide 49• http://www.rusbiotech.ru/old/arhiv_s_11.html• http://www.nhm.ac.uk/hosted_sites/iba• Slide 50• Slide 51• http://www.amnh.org/exhibitions/permanent/ocean/03_oceanlife/g3_permian.php• http://mangrove.nus.edu.sg/guidebooks/text/2005.html• http://www.zoology.uwa.edu.au/staff/rblack/13062001.html• http://www.marinbi.com/bryozoa/• http://www.microscopy-uk.org.uk/micropolitan/index.html• Unknown• Slide 52• Carbonate Depositional Environments, 1983, AAPG.