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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

2

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

3

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

4

• 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

5

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

6

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

7

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

8

Physiology

Page 794 Figure 21.19

Water and Nutrient Flow

Musculature

Page 796

Bivalve

Brachiopod

Adductor

9

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

12

• 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

13

Physiology

Page 784 Figure 21.9

Autozooids

Heterozoids

AviculariaVibracula and Spines

14

Ovicells

Autozooid

OvicellsZooecium

Reproduction

• Hermaphroditic or Gonochoristic

• Asexual – Budding

• Sexual– Eggs and Sperm

1.

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

15

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

16

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.

17

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

20

• 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.

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