Bryozoa

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Bryozoans, or "moss animals," are aquatic, mostly marine, colonial organisms, superficially rather like coral. A few to many millions of these individuals may form one colony. The colonies range from millimeters to meters in size, but the individuals that make up the colonies are tiny; usually less than a millimeter long. Usually they encrust rocky surfaces, shells, or algae. The colonies may be soft-bodied or calcareous like corals. It is the latter that are most often fossilized. There are about 5000 living species, with several times that number of fossil forms known.

After approximately 250 years of scientific study, the sum total of accumulated human wisdom on the subject of bryozoan phylogeny is Nil. Zero. Nada. It would be fair to state that we know as much about the phylogeny of entirely hypothetical Martian cryptobionts. Those, at least, have been the subject of intense speculation by learnèd folks who can get away with such speculations in the semi-popular press. Bryozoologists just throw their hands up -- like so many zoöids waving their happy lophophores about. Maybe they'll catch onto something that way -- but we are not optimistic.

For taxonomic purposes, the Bryozoa are classified as follows:

PHYLUM BRYOZOA

Class Stenolaemata
Order Cyclostomata
Order Hederellida
Order Trepostomatida
Order Cystoporida
Order Cryptostomida
Order Fenestrida
Class Gymnolaemata
Order Ctenostomata
Order Cheilostomata
Class Phylactolaemata

MAK990511 ATW050910.

Contents

Class Stenolaemata

These are marine bryozoans with tubular zooids with strongly calcified walls. The lophophore is protruded by action of annular muscles. Most forms lack an operculum. This was the predominant bryozoan group during the Paleozoic. Some grew as lacy or fan-like colonies that became important reef builders and in some regions form an abundant component of limestones. Their numbers were greatly reduced by the terminal Permian extinction event.

Order Cyclostomata (= Tubuliporata)

Bryozoa with encrusting or erect colonies, many with communication pores. The skeletal structure is typically laminated. The aperture for each zooid (little animal) is either circular or polygonal. Late Ordovician to Recent - about 250 genera

Systematic Classification

SUBORDER PALEOTUBULIPORINA
SUBORDER TUBULIPORINA
SUBORDER FASCICULINA
SUBORDER ARTICULINA
SUBORDER CANCELLATA
SUBORDER CERIOPORINA
SUBORDER RECTANGULATA

Order(?) Hederellida

A group of bryozoans similar to cyclostomatids; of uncertain relationships. Late Ordovician to Carboniferous.

Order Trepostomatida

Colonies generally robust; dendroid, encrusting, or massive. Ordovician to Triassic - 200 genera.

Order Cystoporida

Colonies encrusting or erect, robust or delicate. Crescent-shaped thickened strip or projections (lunarium) around each aperture. Early Ordovician to Late Permian - 100 genera.

Systematic Classification

SUBORDER CERAMOPORINA
SUBORDER FISTULIPORINA

Order Cryptostomida

Colonies erect, typically delicate; foliate, dendroid(tree-like), or bilaminar sheets. Skeletons without communication pores. Early Ordovician to Late Permian - 90 genera.

Systematic Classification

SUBORDER PTILODICTYINA
SUBORDER RHABDOMESINA
SUBORDER TIMANODICTYINA
SUBORDER GOLDFUSSITRYPINA
SUBORDER STREBLOTRYPINA

Order Fenestrida (=Fenestrata)

Includes some of the most amazing Bryozoa. The colonies are erect, typically delicate; reticulate (net-like) or pinnate (fern-like). This order was previously included under the Cryptostomata. Early Ordovician to Late Permian - 100 genera.

Family FENESTELLIDAE

Class Gymnolaemata

Mostly marine bryozoans with cylindrical or flattened zooids. Lophophore is protruded by action of muscles pulling on frontal wall. Includes the majority of living bryozoan species.

Order Ctenostomata

Uncalcified forms, including several that bore into calcareous substrates. Skeleton membranous or gelatinous. Opercula rare. Eggs brooded in body cavity. Almost all fossils are traces of boring forms. Ordovician to Recent - 50 named genera.

Systematic Classification

Suborder STOLONIFERINA
Superfamily TEREBRIPOROIDEA
family VINELLIDAE
Suborder VESICULARINA
Superfamily VESICULARIOIDEA
family VESICULARIIDAE
SUBORDER ALCYONIDIINA
SUPERFAMILY ALCYONIDIOIDEA
SUBORDER FLUSTRELLIDRINA
SUPERFAMILY HAYWARDOZOONOIDEA
SUPERFAMILY FLUSTRELLIDROIDEA
SUBORDER VICTORELLINA
SUPERFAMILY VICTORELLOIDEA
SUBORDER PALUDICELLINA
SUPERFAMILY PALUDICELLOIDEA
SUBORDER VESICULARINA
SUPERFAMILY VESICULARIOIDEA
SUBORDER STOLONIFERINA
SUPERFAMILY TRITICELLOIDEA
SUPERFAMILY AEVERRILLIOIDEA
SUPERFAMILY VALKERIOIDEA
SUPERFAMILY ARACHNIDIOIDEA
SUPERFAMILY TEREBRIPOROIDEA
SUPERFAMILY HISLOPIOIDEA
SUPERFAMILY PENETRANTIINA

Order Cheilostomata

These are calcified forms, usually with an operculum. Short-box-like zooeica (chambers for the little animal). The eggs are usually brooded in ovicells (swollen spherical chambers in which the fertilized egg develops into a larva). The suborders are distinguished according to frontal calcification and mechanism of lophophore protrusion. This order includes the majority of recent Bryozoa. Late Jurassic - Recent. 1000 genera.

Systematic Classification

Infraorder FLUSTRINA
Superfamily CALLOPOROIDEA
family Flustridae
Suborder NEOCHEILOSTOMINA
Superfamily CALLOPOROIDEA
family CUPULADRIIDAE
Suborder ASCOPHORINA
Infraorder LEPRALIOMORPHA Gordon, 1989
Superfamily SMITTINOIDEA
Family BITECTIPORIDAE
Infraorder LEPRALIOMORPHA
Superfamily SCHIZOPORELLOIDEA
Family SCHIZOPORELLIDAE
SUBORDER PROTOCHEILOSTOMATINA
SUPERFAMILY LABIOSTOMELLOIDEA
SUBORDER INOVICELLINA
SUPERFAMILY AETEOIDEA
SUBORDER SCRUPARIINA
SUPERFAMILY SCRUPARIOIDEA
SUBORDER MALACOSTEGINA
SUPERFAMILY ELECTROIDEA
SUBORDER NEOCHEILOSTOMINA
INFRAORDER FLUSTRINA
SUPERFAMILY CALLOPOROIDEA
INFRAORDER CELLULARIOMORPHA
SUPERFAMILY BUGULOIDEA
SUPERFAMILY MICROPOROIDEA
SUPERFAMILY CELLARIOIDEA
SUBORDER ASCOPHORINA
INFRAORDER ACANTHOSTEGOMORPHA
SUPERFAMILY CRIBRILINOIDEA
SUPERFAMILY BIFAXARIOIDEA
SUPERFAMILY NEPHROPOROIDEA
SUPERFAMILY CATENICELLOIDEA
INFRAORDER HIPPOTHOOMORPHA
SUPERFAMILY HIPPOTHOOIDEA
INFRAORDER UMBONULOMORPHA
SUPERFAMILY ARACHNOPUSIOIDEA
SUPERFAMILY ADEONOIDEA
SUPERFAMILY PSEUDOLEPRALIOIDEA
SUPERFAMILY LEPRALIELLOIDEA (UMBONULOIDEA)
SUPERFAMILY CHLIDONIOPSOIDEA
INFRAORDER LEPRALIOMORPHA
SUPERFAMILY SMITTINOIDEA
SUPERFAMILY SCHIZOPORELLOIDEA
SUPERFAMILY URCEOLIPOROIDEA
SUPERFAMILY DIDYMOSELLOIDEA
SUPERFAMILY EUTHYRISELLOIDEA
SUPERFAMILY SIPHONICYTAROIDEA
SUPERFAMILY MAMILLOPOROIDEA
SUPERFAMILY CELLEPOROIDEA
SUPERFAMILY CONESCHARELLINOIDEA

Class Phylactolaemata

Non-calcareous freshwater bryozoans with no zooid polymorphism. Horse-shoe shaped lophophore. No fossil record except for a few statoblasts (resistant wintering structures) from Quaternary sediments.

Systematic Classification

ORDER PLUMATELLIDA

Phylogeny

<==Bryozoa [Ectoprocta, Gymnolaemata, Halcyonellea]
   |  i. s.: Hyalinella punctata
   |         Stolella
   |         Lophopodella carteri
   |         Metrarabdotos
   |           |--M. micropora
   |           `--+--M. chipolanum
   |              `--+--+--M. colligatum
   |                 |  `--+--M. auriculatum
   |                 |     `--M. tenue
   |                 `--+--M. kugleri
   |                    `--+--M. lacrymosum
   |                       `--M. unguiculatum
   |         Rectonychocella dimorphocella Canu & Bassler 1935 (n. d.)
   |         Amphiblestrum ovatum Maplestone 1901
   |         Semicoscinium tenuiceps
   |         Constellaria florida
   |         Rhombotrypa
   |         Parvohallopora
   |         Monticulipora
   |         Hippoporina perforata
   |         Celleporella hyalina
   |         Cryptosula
   |           |--C. pallasiana
   |           `--C. zavjalovensis
   |         Fenestrulina malusii
   |         Hippopodinella adpressa
   |         Adeonella
   |         Uscia mexicana
   |         Tendra
   |         Flustrellidra hispida
   |         Pentapora foliacea [=Lepralia foliacea]
   |         Camptoplites atlanticus
   |         Terebripora comma
   |         Eurystomella foraminigera
   |         Carbasea indivisa
   |         Kinetoskias
   |         Escharella immisca
   |         Forella repens
   |         Flustrella Ehrenberg 1839 (n. d.)
   |           `--*F. concentrica Ehrenberg 1839 (n. d.)
   |         Pherusa tubulosa
   |         Gemellaria loricata
   |         Schizobrachiella sanguinea
   |         Schizomavella linearis
   |         Casteropora vetusta Oehlert 1888
   |         Kazakhstanella Nekhoroshev 1956
   |         Caberea boryi
   |--Ctenostomata
   `--+--Cheilostomata
      `--+--Stenolaemata
         `--Phylactolaemata [Lophopoda]
              |--Cristatella mucedo
              |--Lophopus cristallinus
              `--Plumatella [Plumatellida]
                   |--P. fungosa [=Alcyonella fungosa, A. stagnorum]
                   `--P. repens

* Type species of genus indicated

References

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Bosence, D. W. J. 1979. The factors leading to aggregation and reef formation in Serpula vermicularis L. In Biology and Systematics of Colonial Organisms (G. Larwood & B. R. Rosen, eds.) pp. 299-318. Academic Press: London.

Botquelen, A., & E. Mayoral. 2005. Early Devonian bioerosion in the Rade de Brest, Armorican Massif, France. Palaeontology 48 (5): 1057-1064.

Boxshall, G. A., & A. G. Humes. 1988. A new genus of Lichomolgidae (Copepoda: Poecilostomatoida) associated with a phoronid in Hong Kong. Bulletin of the British Museum (Natural History), Zoology Series 54 (6): 301-307.

Buss, L. W. 1979. Habitat selection, directional growth and spatial refuges: Why colonial animals have more hiding places. In Biology and Systematics of Colonial Organisms (G. Larwood & B. R. Rosen, eds.) pp. 459-497. Academic Press: London.

Cook, P. L. 1979. Some problems in interpretation of heteromorphy and colony integration in Bryozoa. In Biology and Systematics of Colonial Organisms (G. Larwood & B. R. Rosen, eds.) pp. 193-210. Academic Press: London.

Cook, P. L., & P. E. Bock. 2001. Calescharidae, a new family for the Tertiary to Recent genera Caleschara MacGillivray and Tretosina Canu & Bassler (Bryozoa: Cheilostomata). Invertebrate Taxonomy 15: 527-550.

Eckert, J. D., & C. E. Brett. 2001. Early Silurian (Llandovery) crinoids from the Lower Clinton Group, western New York State. Bulletins of American Paleontology 360: 1-88.

Erickson, J. M., & T. D. Bouchard. 2003. Description and interpretation of Sanctum laurentiensis, new ichnogenus and ichnospecies, a domichnium mined into Late Ordovician (Cincinnatian) ramose bryozoan colonies. Journal of Paleontology 77 (5): 1002-1010.

Giribet, G., D. L. Distel, M. Polz, W. Sterrer & W. C. Wheeler. 2000. Triploblastic relationships with emphasis on the acoelomates and the position of Gnathostomulida, Cycliophora, Plathelminthes, and Chaetognatha: A combined approach of 18S rDNA sequences and morphology. Systematic Biology 49: 539-562.

Gould, S. J., & N. Eldredge. 1993. Punctuated equilibrium comes of age. Nature 366: 223-227.

Harmer, S. F. 1915. The Polyzoa of the Siboga Expedition – Part I. Entoprocta, Ctenostomata and Cyclostomata. E. J. Brill: Leyden.

Kashin, I. A., E. V. Bagaveeva & S. F. Chaplygina. 2003. Fouling communities of hydrotechnical constructions in Nakhodka Bay (Sea of Japan). Russian Journal of Marine Biology 29: 267-283.

Kozloff, E. N. 1992. The genera of the phylum Orthonectida. Cahiers de Biologie Marine 33: 377-406.

Okamura, B., A. Curry, T. S. Wood & E. U. Canning. 2002. Ultrastructure of Buddenbrockia identifies it as a myxozoan and verifies the bilaterian origin of the Myxozoa. Parasitology 124: 215-223.

Prothero, D. R. 1998. Bringing Fossils to Life: An introduction to paleobiology. WCB McGraw-Hill: Boston.

Pushkin, V. I., & L. E. Popov. 2005. Two enigmatic bryozoans from the Middle Orodovician of the East Baltic. Palaeontology 48 (5): 1065-1074.

Ryland, J. S. 1979. Structural and physiological aspects of coloniality in bryozoans. In Biology and Systematics of Colonial Organisms (G. Larwood & B. R. Rosen, eds.) pp. 211-242. Academic Press: London.

Credits

MAK?; phylogeny and references CKT071129

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