Mosadýr (Bryozoa)

Picture: Guðmundur Guðmundsson

Location of Bioice samples with information on bryozoan species composition (red dots); Bioice samples with unknown Bryozoan species composition (open circles).    

Picture: Guðmundur Guðmundsson

Ocurrence of bryozoan specis in the Bioice samples. 


Bryozoans are ubiquitous in Icelandic waters and form a prominent part of the benthic comminities. They occurred in 820 samples that were collected during the BIOICE programme. Species composition has been analyzed in 445 of those 820 samples by Peter Hayward. The preliminary list of known species in Icelandic waters, that is presented here, comprises 171 species and additional 21 putative species are identified to genus. 

Bryozoans are a phylum of aquatic invertebrates, mostly marine but with some species inhabiting also fresh or brackish waters. They are the only phylum in which all species form colonies. Each colony comprises tens to many thousands of individuals called zooids. At least some of these zooids possess a lophophore, a feeding apparatus comprising an inverted cone-shaped crown of tentacles. Rhythmic beating of tiny cilia on the tentacles drives a current of water containing suspended food particles towards the mouth. Bryozoans feed on phytoplankton, like many other organisms living on the seabed. In a few instances, colonies are mobile, but most species are sessile and rely on ambient water currents to bring planktonic food into their vicinity.

Although all the zooids in a bryozoan colony are genetically identical (clonal), they may differ in morphology, a phenomenon called polymorphism. Differences in morphology reflects differences in function. For example, avicularia, named for their resemblance to a bird’s head, are defensive zooids with beak-like mandibles used to capture would-be predators. Specialized zooids gonozooids are also present in some bryozoans, their purpose being to brood the embryos before they are released as larvae. A consequence of polymorphism is that the feeding zooids must provide nutrition for the non-feeding zooids. This is accomplished using the funicular system, blood vessel-like strands that pass from zooid to zooid through pores in the walls.

Bryozoan colonies can propagate in two ways. All species produce larvae which are the products of sexual reproduction when an egg from one zooid is fertilized by a sperm usually from a separate colony. Most bryozoan larvae are short-lived, settling on a solid surface, such as a rock, shell or seaweed, within a matter of hours and undergoing metamorphosis to give the founding zooid (ancestrula) of a new colony. In a few bryozoans, the larva is of longer duration and can feed while swimming in the plankton. Larvae are the main means of dispersal in bryozoans (rafting on floating objects is another), and species having long-lived feeding larvae can potentially disperse further and more quickly than those with short-lived larvae. The second mode of colony propagation is by breakage. New colonies resulting from breakage, be it accidental, physical or biological in cause, have the same genotype as the parent colony.

Most bryozoans secrete calcareous skeletons. Features of the skeleton are of great importance in taxonomy, which means that it is usually possible to identify species using dried specimens. Indeed, it is common practice to bleach bryozoans to remove all of the soft organic parts and expose the bare skeleton. The resistant skeletons of bryozoans fossilize very well: bryozoan fossils are common in sedimentary rocks deposited as long ago as the Early Ordovician, some 450 million years ago. The calcareous skeletons of fossil bryozoans can be so abundant that they form bryozoan limestones.

Despite their high diversity there are probably more than 10,000 living bryozoan species and great abundance, bryozoans are unknown to many people. This is reflected in that fact that none of the three English vernacular names for bryozoans - moss animals, sea mats and lace corals are particularly familiar. Historically, bryozoans have had no economic value, although a marine species called Bugula neritina is the host for a bacterium that manufactures bryostatin, a natural product with tumour reducing properties which is currently in clinical trials.

Modern bryozoans are classified into four major groups: class Phylactolaemata and orders Ctenostomata, Cheilostomata and Cyclostomata. Phylactolaemates, which are exclusively freshwater, will not be considered further. Ctenostomes are soft-bodied marine and freshwater bryozoans that form stoloniferous or gelatinous colonies, some species of the former type boring into calcareous substrates such as shells. Their feeding zooids tend to be tubular in shape, unlike those of the related cheilostomes which typically have box-shaped zooids and differ in secreting a calcareous skeleton. The morphology of both cheilostome zooids and colonies is remarkably varied, as reflected by the enormous taxonomic diversity of this marine order. Polymorphism is highly developed in cheilostomes, with avicularia, spinozooids and rhizozooids being present in many species. The majority of cheilostomes brood their embryos, one at a time, in distinctive hood-like structures called ovicells. In contrast, the final bryozoan group, the cyclostomes, have distinct polymorphic zooids called gonozooids, each of which broods a clutch of clonal embryos produced by polyembyonic fission. Cyclostomes are less diverse than cheilostomes and tend to have simple, tubular skeletons.

Cite bryozoa pages as: Peter Hayward, Piotr Kuklinski, and Gudmundur Gudmundsson (2020). Bryozoa (mosadýr) in Icelandic waters (BIOICE).


In general:

Hayward, P. J. and Ryland, J. S., 1998. Cheilostomatous Bryozoa Part I. Aeteoidea – Cribrilinoidea, in: Barnes, R. S. K. and Crothers, J.H., (eds) Synopses of the British Fauna (New Series). No.10 (Second Edition): 1-366.

Hayward, P. J. and Ryland, J.S., 1999. Cheilostomatous Bryozoa Part II. Hippothooidea – Celleporoidea, In: Barnes RSK, Crothers JH, (eds) Synopses of the British Fauna (New Series). No.14 (Second Edition): 1-416.

McKinney, F. K. and Jackson, J. B. C., 1991. Bryozoan Evolution. Chicago and London: The University of Chicago Press, 238 pp.

Ryland, J.S., 1970. Bryozoans. Hutchinson, London.


Arctic Bryozoa:

Gostilovskaya, M.G., 1978. Bryozoa of the White Sea. (in Russian). Leningrad: “Nauka”.

Kluge, G.A., 1975. Bryozoa of the northern seas of the USSR. New Delhi: Amerind Publishing Co.Pvt. pp. 711.

Ryland, J.S., 1963. Systematic and biological studies on Polyzoa (Bryozoa) from Western Norway. Sarsia, 14: 1-61.

Smitt, F.A., 1868. Bryozoa marina in regionibus arcticis et borealibus vivienta recensuit – Ofversigt af  Vetenskap-Akademiens Forhandlingar, 1867. 6:443-487.

Smitt, F.A, 1871. Kritisk Forteckning ofver Skandinaviens Hafs-Bryozoer. Ofversigt af Kongl. Vetenskaps - Akademiens Forhandlingar, 27: 1115-1134.

Web based Atlas of Arctic Bryozoa:

Bryozoan species in Icelandic waters, according to accepted names in WoRMS,

processed with Taxon Match Tool, (accessed May 21st, 2020) 

Order Family Species Author
Cheilostomatida Aeteidae Aetea sica (Couch, 1844)
Cheilostomatida Aeteidae Aetea truncata (Landsborough, 1852)
Cheilostomatida Beaniidae Beania thula Hayward, 1994
Cheilostomatida Bitectiporidae Hippomonavella borealis (Smitt, 1872)
Cheilostomatida Bitectiporidae Hippoporina harmsworthi (Waters, 1900)
Cheilostomatida Bitectiporidae Hippoporina reticulatopunctata (Waters, 1900)
Cheilostomatida Bitectiporidae Schizomavella (Schizomavella) linearis Norman, 1893
Cheilostomatida Bitectiporidae Smittoidea propinqua (M.Sars, 1863)
Cheilostomatida Bryocryptellidae Cystisella saccata (Busk, 1856)
Cheilostomatida Bryocryptellidae Marguetta lorea (Alder, 1864)
Cheilostomatida Bryocryptellidae Palmiskenea aquilonia Hayward, 1994
Cheilostomatida Bryocryptellidae Palmiskenea faroensis Hayward, 1994
Cheilostomatida Bryocryptellidae Palmiskenea plana (Hincks, 1888)
Cheilostomatida Bryocryptellidae Palmiskenea skenei (Ellis & Solander, 1786)
Cheilostomatida Bryocryptellidae Palmiskenea sp. (Norman, 1876)
Cheilostomatida Bryocryptellidae Porella aperta (Dawson, 1859)
Cheilostomatida Bryocryptellidae Porella belli Hayward, 1994
Cheilostomatida Bryocryptellidae Porella compressa (Fleming, 1828)
Cheilostomatida Bryocryptellidae Porella cymosa (M.Sars, 1851)
Cheilostomatida Bryocryptellidae Porella laevis (Nordgaard, 1905)
Cheilostomatida Bryocryptellidae Porella patula (Norman, 1868)
Cheilostomatida Bryocryptellidae Porella peristomata (Norman, 1868)
Cheilostomatida Bryocryptellidae Porella struma (Smitt, 1868)
Cheilostomatida Bugulidae Bicellariella ciliata (Linnaeus, 1758)
Cheilostomatida Bugulidae Bicellarina alderi (Busk, 1859)
Cheilostomatida Bugulidae Bugulella fragilis Verrill, 1879
Cheilostomatida Bugulidae Bugulella gracilis (Nichols, 1911)
Cheilostomatida Bugulidae Corynoporella tenuis Hincks, 1888
Cheilostomatida Bugulidae Crisularia harmsworthi (Waters, 1900)
Cheilostomatida Bugulidae Crisularia purpurotincta (Norman, 1868)
Cheilostomatida Bugulidae Crisularia turrita (Desor, 1848)
Cheilostomatida Bugulidae Dendrobeania decorata (Verrill, 1879)
Cheilostomatida Bugulidae Dendrobeania fessa Kluge, 1955
Cheilostomatida Bugulidae Dendrobeania fruticosa (Packard, 1863)
Cheilostomatida Bugulidae Dendrobeania murrayana (Bean in Johnston, 1847)
Cheilostomatida Bugulidae Dendrobeania pseudomurrayana Kluge, 1955
Cheilostomatida Bugulidae Kinetoskias arborescens Danielssen, 1868
Cheilostomatida Bugulidae Kinetoskias smitti Danielssen, 1868
Cheilostomatida Bugulidae Nordgaardia pusilla (Nordgaard, 1907)
Cheilostomatida Bugulidae Uschakovia gorbunovi Kluge, 1946
Cheilostomatida Calloporidae Amphiblestrum flemingii (Busk, 1854)
Cheilostomatida Calloporidae Amphiblestrum solidum (Packard, 1863)
Cheilostomatida Calloporidae Amphiblestrum sp. Gray, 1848
Cheilostomatida Calloporidae Callopora craticula (Alder, 1856)
Cheilostomatida Calloporidae Cauloramphus cymbaeformis (Hincks, 1877)
Cheilostomatida Calloporidae Crassimarginatella tensa (Norman, 1903)
Cheilostomatida Calloporidae Flustrellaria whiteavesi (Norman, 1903)
Cheilostomatida Calloporidae Megapora ringens (Busk, 1856)
Cheilostomatida Calloporidae Ramphonotus minax (Nordgaard, 1907)
Cheilostomatida Calloporidae Ramphonotus septentrionalis (King, 1846)
Cheilostomatida Calloporidae Tegella arctica Kluge, 1955
Cheilostomatida Calloporidae Tegella kildinensis Levinsen, 1909
Cheilostomatida Calloporidae Tegella sp. (Bidenkap, 1897)
Cheilostomatida Calloporidae Tegella spitzbergensis (Norman, 1864)
Cheilostomatida Candidae Aquiloniella paenulata (Norman, 1903)
Cheilostomatida Candidae Aquiloniella scabra (van Beneden, 1848)
Cheilostomatida Candidae Bugulopsis peachii (Busk, 1851)
Cheilostomatida Candidae Caberea ellisii (Fleming, 1814)
Cheilostomatida Candidae Notoplites evocatus (Jullien, 1882)
Cheilostomatida Candidae Notoplites harmeri Ryland, 1963
Cheilostomatida Candidae Notoplites jeffreysii (Norman, 1868)
Cheilostomatida Candidae Notoplites sibiricus (Kluge, 1929)
Cheilostomatida Candidae Notoplites smittii (Norman, 1868)
Cheilostomatida Candidae Notoplites sp. Harmer, 1923
Cheilostomatida Candidae Scrupocellaria intermedia (Linnaeus, 1758)
Cheilostomatida Candidae Scrupocellaria scruposa van Benden, 1845
Cheilostomatida Candidae Scrupocellaria sp. (Pallas, 1766)
Cheilostomatida Candidae Tricellaria arctica (Smitt, 1868)
Cheilostomatida Candidae Tricellaria elongata (Van Beneden, 1848)
Cheilostomatida Candidae Tricellaria gracilis Fleming, 1828
Cheilostomatida Candidae Tricellaria sp. (Ellis & Solander, 1786)
Cheilostomatida Candidae Tricellaria ternata Dalyell, 1848
Cheilostomatida Cellariidae Cellaria fistulosa (Linnaeus, 1758)
Cheilostomatida Cellariidae Cellaria sinuosa (Hassall, 1840)
Cheilostomatida Celleporidae Celleporina caliciformis (Lamouroux, 1816)
Cheilostomatida Celleporidae Celleporina pygmaea (Norman, 1868)
Cheilostomatida Celleporidae Celleporina sp. Gray, 1848
Cheilostomatida Celleporidae Celleporina ventricosa (Lorenz, 1886)
Cheilostomatida Celleporidae Lagenipora lepralioides (Norman, 1868)
Cheilostomatida Celleporidae Turbicellepora avicularis Hayward & Hansen, 1999
Cheilostomatida Celleporidae Turbicellepora boreale Hayward & Hansen, 1999
Cheilostomatida Celleporidae Turbicellepora canaliculata (Busk, 1881)
Cheilostomatida Celleporidae Turbicellepora magnicostata (Barroso, 1919)
Cheilostomatida Celleporidae Turbicellepora nodulosa (Lorenz, 1886)
Cheilostomatida Celleporidae Turbicellepora smitti (Kluge, 1962)
Cheilostomatida Celleporidae Turbicellepora sp. Ryland, 1963
Cheilostomatida Chaperiidae Larnacicus corniger (Busk, 1859)
Cheilostomatida Cheiloporinidae Cheilopora sincera (Smitt, 1867)
Cheilostomatida Cribrilinidae Cribrilina (Cribrilina) cryptooecium Norman, 1903
Cheilostomatida Cribrilinidae Cribrilina watersi Andersson, 1902
Cheilostomatida Cribrilinidae Gephyrotes nitidopunctata (Smitt, 1868)
Cheilostomatida Cribrilinidae Membraniporella nitida (Johnston, 1838)
Cheilostomatida Cryptosulidae Harmeria scutulata Levinsen, 1909
Cheilostomatida Doryporellidae Doryporellina reticulata (Linnaeus, 1767)
Cheilostomatida Electridae Electra pilosa (Busk, 1859)
Cheilostomatida Electridae Pyripora catenularia (Busk, 1860)
Cheilostomatida Escharellidae Escharella abyssicola (Ridley, 1881)
Cheilostomatida Escharellidae Escharella connectens (Fleming, 1828)
Cheilostomatida Escharellidae Escharella immersa Hayward, 1979
Cheilostomatida Escharellidae Escharella klugei (Boeck in Smitt, 1868)
Cheilostomatida Escharellidae Escharella labiata (Norman, 1864)
Cheilostomatida Escharellidae Escharella laqueata Hayward, 1994
Cheilostomatida Escharellidae Escharella levinseni (Hincks, 1880)
Cheilostomatida Escharellidae Escharella octodentata Gray, 1848
Cheilostomatida Escharellidae Escharella sp. (Johnston, 1838)
Cheilostomatida Escharellidae Escharella variolosa (Hassall, 1842)
Cheilostomatida Escharellidae Escharella ventricosa (Busk, 1856)
Cheilostomatida Escharellidae Hemicyclopora labrata (Busk, 1855)
Cheilostomatida Escharellidae Hemicyclopora polita Hayward, 1994
Cheilostomatida Escharellidae Neolagenipora rugosa Hayward, 1994
Cheilostomatida Escharellidae Temachia microstoma (Busk, 1860)
Cheilostomatida Escharinidae Escharina alderi (Busk, 1856)
Cheilostomatida Escharinidae Escharina boreale Hayward, 1994
Cheilostomatida Escharinidae Escharina dutertrei haywardi Zabala, Maluquer & Harmelin, 1993
Cheilostomatida Escharinidae Escharina sp. Milne Edwards, 1836
Cheilostomatida Eucrateidae Eucratea loricata (Linnaeus, 1758)
Cheilostomatida Exochellidae Escharoides jacksonii (Waters, 1900)
Cheilostomatida Flustridae Carbasea carbasea (Ellis & Solander, 1786)
Cheilostomatida Flustridae Sarsiflustra abyssicola (Hassall, 1841)
Cheilostomatida Flustridae Securiflustra securifrons (Busk, 1860)
Cheilostomatida Flustridae Terminoflustra barleei (Smitt, 1868)
Cheilostomatida Flustridae Terminoflustra membranaceotruncata (Busk, 1860)
Cheilostomatida Haplopomidae Haplopoma planum Ryland, 1963
Cheilostomatida Haplopomidae Haplopoma sp. Ryland, 1963
Cheilostomatida Hincksiporidae Hincksipora stenostoma (Norman, 1864)
Cheilostomatida Hippoporidridae Hippoporella obesa (Waters, 1900)
Cheilostomatida Hippothoidae Celleporella hyalina (Linnaeus, 1767)
Cheilostomatida Lepraliellidae Lepraliella contigua (Smitt, 1868)
Cheilostomatida Microporellidae Calloporina Neviani, 1895
Cheilostomatida Microporellidae Microporella ciliata (Pallas, 1766)
Cheilostomatida Microporellidae Microporella sp. Hincks, 1877
Cheilostomatida Myriaporidae Leieschara coarctata M.Sars, 1863
Cheilostomatida Myriaporidae Leieschara subgracilis (d'Orbigny, 1853)
Cheilostomatida Onychocellidae Rectonychocella solida (Jullien, 1903)
Cheilostomatida Phidoloporidae Phidolopora elongata (Lamarck, 1816)
Cheilostomatida Phidoloporidae Reteporella beaniana (Jullien, 1903)
Cheilostomatida Phidoloporidae Reteporella grimaldii Busk, 1884
Cheilostomatida Phidoloporidae Reteporella sp. (Nordgaard, 1907)
Cheilostomatida Phidoloporidae Reteporella watersi Hayward, 1994
Cheilostomatida Phidoloporidae Reteporellina hyperborea Lorenz, 1886
Cheilostomatida Romancheinidae Arctonula arctica (M. Sars, 1851)
Cheilostomatida Romancheinidae Ragionula rosacea (Kluge, 1906)
Cheilostomatida Setosellidae Setosella vulnerata (Busk, 1860)
Cheilostomatida Smittinidae Parasmittina jeffreysi Osburn, 1952
Cheilostomatida Smittinidae Parasmittina sp. (Johnston, 1838)
Cheilostomatida Smittinidae Parasmittina trispinosa (Linnaeus, 1758)
Cheilostomatida Smittinidae Pseudoflustra hincksi (Andersson, 1902)
Cheilostomatida Smittinidae Pseudoflustra minima (Stimpson, 1854)
Cheilostomatida Smittinidae Pseudoflustra sinuosa Bidenkap, 1897
Cheilostomatida Smittinidae Pseudoflustra solida Hayward, 1994
Cheilostomatida Smittinidae Pseudoflustra sp. (Fleming, 1828)
Cheilostomatida Smittinidae Pseudoflustra virgula (Busk, 1860)
Cheilostomatida Smittinidae Smittina bella Lorenz, 1886
Cheilostomatida Smittinidae Smittina rigida Hayward, 1994
Cheilostomatida Smittinidae Smittoidea exilis (Smitt, 1868)
Cheilostomatida Stomachetosellidae Stomachetosella normani (Busk, 1854)
Cheilostomatida Stomachetosellidae Stomacrustula cruenta (Busk, 1860)
Cheilostomatida Stomachetosellidae Stomacrustula sinuosa (d'Orbigny, 1853)
Cheilostomatida Tessaradomidae Tessaradoma boreale Busk, 1855
Cheilostomatida Umbonulidae Posterula sarsii Hayward, 1994
Cheilostomatida Umbonulidae Rhamphostomella costata (Smitt, 1868)
Cheilostomatida Umbonulidae Rhamphostomella ovata (Smitt, 1868)
Cheilostomatida Umbonulidae Rhamphostomella plicata (O. Fabricius, 1780)
Cheilostomatida Umbonulidae Rhamphostomella scabra (Sars G.O., 1872)
Cheilostomatida Cheilostomata sp.
Ctenostomatida Alcyonidiidae Alcyonidium candidum Ryland, 1963
Ctenostomatida Triticellidae Triticella flava (Pallas, 1766)
Ctenostomatida Vesiculariidae Amathia imbricata (Adams, 1800)

Cyclostomata sp.

Cyclostomatida Crisiidae Bicrisia abyssicola Kluge, 1962
Cyclostomatida Crisiidae Crisia constans Kluge, 1946
Cyclostomatida Crisiidae Crisia eburnea (Linnaeus, 1758)
Cyclostomatida Crisiidae Crisia klugei Ryland, 1967
Cyclostomatida Crisiidae Crisia sp. Lamouroux, 1812
Cyclostomatida Crisiidae Filicrisia smitti (Kluge, 1946)
Cyclostomatida Frondiporidae Infundibulipora lucernaria (Sars, 1851)
Cyclostomatida Horneridae Hornera lichenoides (Hincks, 1877)
Cyclostomatida Horneridae Hornera sp. Lamouroux, 1821
Cyclostomatida Lichenoporidae Disporella hispida (Fleming, 1828)
Cyclostomatida Lichenoporidae Patinella verrucaria (Fabricius, 1780)
Cyclostomatida Oncousoeciidae Oncousoecia diastoporides (Norman, 1869)
Cyclostomatida Oncousoeciidae Oncousoecia dilatans (Johnston, 1847)
Cyclostomatida Plagioeciidae Diplosolen intricarium (Smitt, 1872)
Cyclostomatida Plagioeciidae Diplosolen obelium (Johnston, 1838)
Cyclostomatida Plagioeciidae Entalophoroecia clavata (Couch, 1842)
Cyclostomatida Plagioeciidae Entalophoroecia deflexa (Norman, 1869)
Cyclostomatida Plagioeciidae Plagioecia patina Boeck, 1862
Cyclostomatida Stigmatoechidae Stigmatoechos violacea (M.Sars, 1863)
Cyclostomatida Tubuliporidae Exidmonea atlantica (Forbes in Johnston, 1847)
Cyclostomatida Tubuliporidae Pencilletta penicillata (Smitt, 1866)
Cyclostomatida Tubuliporidae Tubulipora liliacea (Fabricius, 1780)
Cyclostomatida Tubuliporidae Tubulipora sp. (Hincks, 1860)

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