Bryozoa

Trembling sea mat Victorella pavida, copyright Marine Biological Association of the UK.

Belongs within: Lophotrochozoa.
Contains: Cheilostomata, Phylactolaemata, Stenolaemata, Stoloniferina, Vesiculariidae, Alcyonidiidae, Arachnidiidae.

The Bryozoa, sometimes known as moss animals or lace animals, are aquatic (mostly marine) minute colonial animals with the mouth surrounded by a ring of tentacles (the lophophore) and a U-shaped gut with the anus situated outside the lophophore. The lophophore can be retrated within the tentacle sheath. A single nerve ganglion is present within the coelom, but no vascular, respiratory or excretory system (Prothero 1998). Bryozoan colonies are calcified in the Cheilostomata and Stenolaemata, giving these groups an extensive fossil record. While individual zooecia (zooid body walls) may grow continually in Stenolaemata, the Ctenostomata and Cheilostomata have zooecia that grow to a fixed size and colony growth is by the addition of zooecia (Prothero 1998).

<==Bryozoa [Ectoprocta, Gymnolaemata, Halcyonellea]
    |--+--Cheilostomata P98
    |  `--+--Phylactolaemata P98
    |     `--Stenolaemata P98
    `--Ctenostomata [Ctenostomida] P98
         |  i. s.: Metalcyonidium gautieri R79
         |         Pseudalcyonidium bobinae R79
         |         Monobryozoon Remane 1936 [Monobryozoontidae] H86
         |           |--M. ambulans Remane 1936 H86
         |           |--M. bulbosum Ott 1972 H86
         |           `--M. limicola R79
         |         Orbignyopora devonica (Richards 1974) BM05
         |         Flustrellidra [Flustrellidridae] WT06
         |           `--F. hispida WT06
         |--Stoloniferina H15
         |--Vesicularina H15
         |    |  i. s.: Avenella H15
         |    |         Cryptopolyzoon H15
         |    `--Vesiculariidae H15
         |--Carnosa [Alcyonellea, Carnosina] H15
         |    |--Alcyonidiidae H15
         |    `--Flustrellidae H15
         |         |--Echinella placoides Korotneff 1901 H15
         |         |--Verrucularia v. Suhr 1834 H15
         |         |--Elzerina Lamouroux 1816 H15
         |         |    `--E. blainvillii Lamouroux 1816 (see below for synonymy) H15
         |         `--‘Flustrella’ Gray 1848 non Ehrenberg 1839 H15
         |              |--F. binderi [=Farciminaria binderi] H15
         |              |--F. flabellaria H15
         |              `--F. hispida [incl. F. h. var. cylindrica] H15
         `--Paludicellea [Paludicellina] H15
              |--Arachnidiidae H15
              |--Paludicella [Paludicellidae] H15
              |    `--P. articulata [=Alcyonella articulata] H15
              `--Victorella Saville Kent 1870 [Homodiaetidae, Victorellidae, Victorellides] H15
                   |--*V. pavida Saville Kent 1870 H15
                   |--V. bengalensis Annandale 1911 H15
                   |--V. continentalis Braem 1911 H15
                   |--‘Paludicella’ muelleri Kraepelin 1887 H15
                   |--V. sibogae Harmer 1915 H15
                   `--V. symbiotica Rousselet 1907 H15

Bryozoa incertae sedis:
  Stolella OC02
  Lophopodella carteri R79
  Metrarabdotos CB01
    |--M. micropora GE93
    `--+--M. chipolanum GE93
       `--+--+--M. colligatum GE93
          |  `--+--M. auriculatum GE93
          |     `--M. tenue GE93
          `--+--M. kugleri GE93
             `--+--M. lacrymosum GE93
                `--M. unguiculatum GE93
  Rectonychocella dimorphocella Canu & Bassler 1935 (n. d.) CB01
  Amphiblestrum ovatum Maplestone 1901 CB01
  Semicoscinium tenuiceps EB01
  Constellaria florida EB03
  Rhombotrypa EB03
  Parvohallopora EB03
  Hippoporina perforata KBC03
  Celleporella hyalina KBC03
  Fenestrulina malusii KBC03
  Hippopodinella adpressa R79
  Adeonella C79
  Uscia mexicana C79
  Tendra C79
  Pentapora foliacea R79 [=Lepralia foliacea K92]
  Camptoplites atlanticus R79
  Terebripora d’Orbigny 1842 H75
    `--T. comma R79
  Eurystomella foraminigera R79
  Carbasea indivisa R79
  Kinetoskias R79
  Escharella immisca Bo79
  Forella repens Bu79
  Pherusa tubulosa H15
  Gemellaria loricata BK77
  Schizobrachiella sanguinea BH88
  Schizomavella linearis BH88
  Casteropora vetusta Oehlert 1888 BM05
  Kazakhstanella Nekhoroshev 1956 PP05
  Caberea boryi GD00
  Calpensia nobilis (Esper 1796) (see below for synonymy) H30
  Eschara Linnaeus 1758 H30, L58
    |--E. divaricata Linnaeus 1758 L58
    |--E. fistulosa Linnaeus 1758 L58 [=Cellaria fistulosa H30]
    |--E. foliacea Linnaeus 1758 L58 [=Flustra foliacea H30]
    |--E. fragilis Linnaeus 1758 L58
    `--E. verticillata Linnaeus 1758 L58
  ‘Sertularia’ ciliata Linnaeus 1758 H30, L58
  ‘Sertularia’ eburnea Linnaeus 1758 H30, L58
  ‘Sertularia’ lendigera Linnaeus 1758 H30, L58
  ‘Sertularia’ reptans Linnaeus 1758 H30, L58
  Fredericella sultana KA01
  ‘Lagena’ acuta F71
  Batostomella F71
    |--B. cylindrica Crockford 1941 F71
    `--B. lineata Crockford 1947 F71
  Calopora hattonensis Ross 1961 F71
  Hexagonella F71
    |--H. australe (Cretnall 1926) [=Coscinium australe] F71
    `--H. lineata Crockford 1944 F71
  Dichotrypa fragilis Crockford 1947 F71
  Helicopora australis Etheridge 1921 F71
  Leioclema porosa Crockford 1947 F71
  Minilya duplaris F71
  Protoretepora ampla F71
  Pseudohornera retiformis Crockford 1941 F71
  Ptilopora carinata Crockford 1941 F71
  Saffordotaxis multigranulata (Bretnall 1926) [=Rhombopora multigranulata] F71
  Streblotrypa F71
    |--S. etheridgei Bretnall 1926 F71
    `--S. parallela Crockford 1947 F71
  Haplopoma bimucronatum BBB-S95
  Margaretta ceroeides BBB-S95
  Vinella Ulrich 1890 H75
  Spathipora Fischer 1866 H75
    `--S. prima Voigt 1972 H75
  Penetrantia Silén 1946 H75
  Harmeriella Borg 1940 H75
  Immergentia Silén 1946 H75
  Stolonicella schindewolfi M79
  Taeniocellaria setifera M79
  Reteporidea cancellata M79
  Corynotrypa Bassler 1911 [incl. Cochleatina Bykova 1956] LT64
    `--‘*Cochleatina’ plavinensis Bykova 1956 LT64
  ‘Polytrema’ Risso 1826 nec Rafinesque 1819 nec Férussac 1822 LT64
  Myriaporidae W77
    |--Synocladia King 1849 W77
    |    `--S. biserialis Swallow & Hawn 1858 [incl. S. virgulacea Geinitz 1866] W77
    `--Glauconome Goldfuss 1826 W77
         |--G. nereidis White 1874 W77
         `--G. triliniata W77

Borings: Harmeriella cretacea Voigt 1957 H75
         Immergentia lissajousi Walter 1965 H75
         Penetrantia gosaviensis Voigt & Soule 1973 H75

Calpensia nobilis (Esper 1796) [=Cellepora nobilis; incl. Membranipora bifoveolata, Eschara impressa Moll 1803, Micropora impressa] H30

Elzerina blainvillii Lamouroux 1816 [=Farciminaria blainvillii; incl. Verrucularia dichotoma v. Suhr 1834, Farciminaria dichotoma, Flustrella dichotoma] H15

*Type species of generic name indicated

REFERENCES

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[Bo79] Bosence, D. W. J. 1979. The factors leading to aggregation and reef formation in Serpula vermicularis L. In: Larwood, G., & B. R. Rosen (eds) Biology and Systematics of Colonial Organisms pp. 299–318. Academic Press: London.

[BM05] Botquelen, A., & E. Mayoral. 2005. Early Devonian bioerosion in the Rade de Brest, Armorican Massif, France. Palaeontology 48 (5): 1057–1064.

[BBB-S95] Boubezari, K., G. Bitar & D. Bellan-Santini. 1995. Structure et organisation de trois moulières (Mytilus galloprovincialis et Perna perna) de la région d’Alger. Mésogée 54: 63–72.

[BH88] 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.

[Bu79] Buss, L. W. 1979. Habitat selection, directional growth and spatial refuges: why colonial animals have more hiding places. In: Larwood, G., & B. R. Rosen (eds) Biology and Systematics of Colonial Organisms pp. 459–497. Academic Press: London.

[C79] Cook, P. L. 1979. Some problems in interpretation of heteromorphy and colony integration in Bryozoa. In: (G. Larwood & B. R. Rosen, eds) Biology and Systematics of Colonial Organisms pp. 193–210. Academic Press: London.

[CB01] 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.

[EB01] 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.

[EB03] 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.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.

[GD00] 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.

[GE93] Gould, S. J., & N. Eldredge. 1993. Punctuated equilibrium comes of age. Nature 366: 223–227.

[H75] Häntzschel, W. 1975. Treatise on Invertebrate Paleontology pt W. Miscellanea Suppl. 1. Trace Fossils and Problematica 2^nd ed. The Geological Society of America: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[H15] Harmer, S. F. 1915. The Polyzoa of the Siboga Expedition—Part I. Entoprocta, Ctenostomata and Cyclostomata. E. J. Brill: Leyden.

[H30] Harmer, S. F. 1930. Presidential address. Proceedings of the Linnean Society of London 141: 68–118.

[H86] d’Hondt, J.-L. 1986. Bryozoa. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 145–146. E. J. Brill/Dr W. Backhuys: Leiden.

[KBC03] 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.

[KA01] Kent, M. L., K. B. Andree, J. L. Bartholomew, M. El-Matbouli, S. S. Desser, R. H. Devlin, S. W. Feist, R. P. Hedrick, R. W. Hoffmann, J. Khattra, S. L. Hallett, R. J. G. Lester, M. Longshaw, O. Palenzuela, M. E. Siddall & C. Xiao. 2001. Recent advances in our knowledge of the Myxozoa. Journal of Eukaryotic Microbiology 48: 395–413.

[K92] Kozloff, E. N. 1992. The genera of the phylum Orthonectida. Cahiers de Biologie Marine 33: 377–406.

[L58] Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii: Holmiae.

[LT64] Loeblich, A. R., Jr & H. Tappan. 1964. Sarcodina: chiefly “thecamoebians” and Foraminiferida. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt C. Protista 2 vol. 2. The Geological Society of America and The University of Kansas Press.

[M79] Müller, A. H. 1979. Fossilization (taphonomy). In: Robison, R. A., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt A. Introduction. Fossilisation (Taphonomy), Biogeography and Biostratigraphy pp. A2–A78. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[OC02] 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.

[P98] Prothero, D. R. 1998. Bringing Fossils to Life: An introduction to paleobiology. WCB McGraw-Hill: Boston.

[PP05] Pushkin, V. I., & L. E. Popov. 2005. Two enigmatic bryozoans from the Middle Ordovician of the East Baltic. Palaeontology 48 (5): 1065–1074.

[R79] Ryland, J. S. 1979. Structural and physiological aspects of coloniality in bryozoans. In: Larwood, G., & B. R. Rosen (eds) Biology and Systematics of Colonial Organisms pp. 211–242. Academic Press: London.

[WT06] Waeschenbach, A., M. J. Telford, J. S. Porter & D. T. J. Littlewood. 2006. The complete mitochondrial genome of Flustrellidra hispida and the phylogenetic position of Bryozoa among the Metazoa. Molecular Phylogenetics and Evolution 40 (1): 195–207.

[W77] White, C. A. 1877. Report upon the invertebrate fossils collected in portions of Nevada, Utah, Colorado, New Mexico, and Arizona, by parties of the expeditions of 1871, 1872, 1873, and 1874. U.S. Geographical Surveys West of the One Hundredth Meridian 4 (1): 1–219, pls 1–21.