Bugula neritina (brown bryozoan)
- Summary of Invasiveness
- Taxonomic Tree
- Distribution Table
- Habitat List
- Biology and Ecology
- Means of Movement and Dispersal
- Pathway Causes
- Pathway Vectors
- Impact Summary
- Risk and Impact Factors
- Uses List
- Similarities to Other Species/Conditions
- Prevention and Control
- Principal Source
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Bugula neritina (Linnaeus, 1758)
Preferred Common Name
- brown bryozoan
Other Scientific Names
- Anamarchis neritina
- Sertularia neritina
International Common Names
- English: common bugula
Summary of InvasivenessTop of page
Bugula neritina forms flexible bushy colonies, branching biserial, to about 10 cm high and is purplish-brown in colour. Zooids white and globular, with the outer corner pointed (Bishop Museum 2002, in Gordon and Mawatari, 1992). Zooids are large and measure an average of 0.97 x 0.28 mm. B. neritina differs from other species in this genus in that it possesses no avicularia and no spines. The lophophore measures an average of 0.764 mm in diameter and bears 23 tentacles (SMSFP 2001). Embryos brooded in ovicells are dark brown in colour and measure approximately 0.25 mm in diameter (SMSFP 2001 in Winston 1982).
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Bryozoa
- Class: Gymnolaemata
- Order: Cheilostomatida
- Family: Bugulidae
- Genus: Bugula
- Species: Bugula neritina
DescriptionTop of page
Bugula neritina forms flexible bushy colonies, branching biserial, to about 10cm high and is purplish-brown in colour. Zooids white and globular, with the outer corner pointed (Bishop Museum 2002, in Gordon and Mawatari, 1992). Zooids are large and measure an average of 0.97 X 0.28mm. B. neritina differs from other species in this genus in that it possesses no avicularia and no spines. The lophophore measures an average of 0.764mm in diameter and bears 23 tentacles (SMSFP 2001). Embryos brooded in ovicells are dark brown in colour and measure approximately 0.25mm in diameter (SMSFP 2001 in Winston 1982).
DistributionTop of page
Bugula neritina a common fouling organism worldwide, is reported from all seas except sub Arctic and sub Antarctic regions (Bishop Museum 2002). The cosmopolitan distribution of the species appears to be due to shipping introductions (Mackie et al. 2006). Furthermore, genetically divergent but morphologically unrecognised (=’cryptic’) species of B. neritina have been identified in the United States.
Distribution TableTop of page
The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Atlantic, Western Central||Present||Introduced||Invasive||ISSG, 2011|
|Korea, DPR||Present||Introduced||Invasive||ISSG, 2011|
|Korea, Republic of||Present||Introduced||Invasive||ISSG, 2011|
|USA||Present||Present based on regional distribution.|
|-South Carolina||Present||Introduced||Invasive||ISSG, 2011|
Central America and Caribbean
|Puerto Rico||Present||Introduced||1940||Invasive||ISSG, 2011|
|Ecuador||Present||Present based on regional distribution.|
|-Galapagos Islands||Present||Introduced||1930||Invasive||ISSG, 2011|
|UK||Present||Present based on regional distribution.|
|-England and Wales||Present||Introduced||1912||Invasive||ISSG, 2011|
|-New South Wales||Present||Introduced||1993||Invasive||ISSG, 2011|
|New Zealand||Present||Introduced||1949||Invasive||ISSG, 2011|
HabitatTop of page
B. neritina colonies are typically found in harbours and embayments, intertidal to 5m, attached to any available hard substrate (Bishop Museum, 2002). Larvae colonise a variety of artificial substrata including hulls (Mackie et al., 2006). Studies have shown B. neritina larvae prefer to attach to rougher surfaces and prefer to attach to organic material. For example, in nature they frequently affix themselves to algae and to established bryozoan colonies (Lynch, 1947). B. neritina is found in euhaline and polyhaline regions (water salinity around 30-18‰) (Winston, 1977).
In North America B. neritina occurs on rocky reefs and seagrass leaves (Hayes et al., 2005).
Habitat ListTop of page
|Coastal areas||Present, no further details|
|Marine||Present, no further details||Harmful (pest or invasive)|
Biology and EcologyTop of page
The bryozoan is a suspension feeder. It has a retractable U-shaped crown of tentacles (lophophore) which bear cilia that create a current, bringing microscopic plankton and organic particles toward the animal. Particles are then guided into the mouth by action of the tentacles and cilia (Bishop Museum 2002).
Each bryozoan colony begins from a single, sexually produced, primary zooid. This zooid undergoes asexual budding to produce a group of daughter cells, which themselves form buds, and so on. Most bryozoans are hermaphroditic, each zooid capable of producing sperm and eggs. Sperm is released into the coelom and the fertilised eggs are retained and brooded for a time before being released (Bishop Museum 2002).
Bryozoans have swimming, lecithotrophic larvae that attach and metamorphose within 1 or 2 days following release from the colony. Larvae are initially photopositive but soon become photonegative/Geopositive, settling usually within a few hours of release (Lynch, 1947). Larvae may have gregarious settlement (Keough, 1984). Bugula larvae generally settle throughout the year except during midwinter (Sutherland and Karlson 1977). Field studies in Australia and North America show considerable variation in life history in B. neritina from different habitats, apparently due to genetic or early environmental effects (Keough, 1989; NEMESIS 2006). B. neritina's life history may include an annual period of dormancy, in which colonies recede to a regenerative holdfast (Dyrynda and Ryland 1982). This senescence occurs at differing times of year and appears dependent upon water temperature, with populations in cool-temperate areas receding during winter and populations in warm areas receding over summer months (Keough and Chernoff, 1987).
Means of Movement and DispersalTop of page
Introduction pathways to new locations
Aquaculture:Bugula neritina attaches to oyster shells and be transferred along with oyster shippings (Cohen 2005).
Ship ballast water:Bugula neritina can be transported via tiny colonies attached to the sides of ballast tanks or on floating material inside the ballast tanks (Cohen 2005).
Ship/boat hull fouling: Ship/boat hull fouling is a common means of movement of Bugula neritina colonies and a likely source of ongoing introductions.
Local dispersal methods
Agriculture (local):Bugula neritina attaches to oyster shells and be transferred along with oyster shippings (Cohen 2005).
Pathway CausesTop of page
Pathway VectorsTop of page
Impact SummaryTop of page
|Fisheries / aquaculture||Negative|
ImpactTop of page
Risk and Impact FactorsTop of page Invasiveness
- Proved invasive outside its native range
- Highly adaptable to different environments
- Infrastructure damage
- Negatively impacts aquaculture/fisheries
- Transportation disruption
- Highly likely to be transported internationally accidentally
- Difficult/costly to control
UsesTop of page
Bugula neritina colonies are the source of a novel chemical compound (bryostatin) which has been shown to be effective against leukaemia and a number of other kinds of cancer. A newly described species of bacterium, which is symbiotic to B. neritina cryptic species 'type D', appears to be the source of bryostatins (Davidson and Haygood, 1999; Davidson et al. 2001).
Uses ListTop of page
- Source of medicine/pharmaceutical
Similarities to Other Species/ConditionsTop of page
Three morphologically cryptic species have been identified based on mitochondrial genetic divergence and bacterial symbionts (Davidson and Haygood, 1999; McGovern and Hellberg, 2003): B. neritina, B. stolonifera and B. turrita.
Bugula turrita can be distinguished from B. turrita by the presence of avicularia. Bugula neritina does not range farther north on the Atlantic Coast than Beaufort, North Carolina, it is replaced along the northern half of the East Coast of USA by Bugula turrita (Lynch 1947).
Prevention and ControlTop of page
Compiled by IUCN SSC Invasive Species Specialist Group (ISSG)
For more details, please see Hayes et al. 2005.
The rankings determined in Hayes et al. 2005 will be used by the National Introduced Marine Pest Coordinating Group in Australia to assist in the development of national control plans which could include options for control, eradication and/or long term management.
It has been suggested that ballast water control measures be implemented to control the spread of B. nertinavia the oyster aquaculture industry (PWSRCAC 2004).
Chemical: Copper-based treatments have been used to control many pest species. The attachment of B. neritina larvae to copper, mercury and control paint was investigated by Wisely (1962) who found that the numbers attaching to the control paint strips was seven times greater than the numbers attaching to copper, and twenty times greater than the numbers attaching to mercury (NIMPIS 2001). Introduction of B. neritina by copper-painted vessels may be aided by a potential tolerance to toxicants (Piola and Johnston 2006).
BibliographyTop of page
Bishop Museum. 2002. Bugula neritina (Waters, 1878), Guidebook of introduced marine species of Hawaii. Hawaii Biological Survey, Bishop Museum.
Centre for Environment, Fisheries & Aquaculture Science (CEFAS)., 2008. Decision support tools-Identifying potentially invasive non-native marine and freshwater species: fish, invertebrates, amphibians. http://www.cefas.co.uk/projects/risks-and-impacts-of-non-native-species/decision-support-tools.aspx
Cohen, A.N. 2005. Guide to the Exotic Species of San Francisco Bay. San Francisco Estuary Institute: Oakland, USA. http://www.exoticsguide.org/species_pages/b_neritina.html
CONABIO. 2008. Sistema de información sobre especies invasoras en México. Especies invasoras - Otros invertebrados. Comisión Nacional para el Conocimiento y Uso de la Biodiversidad. Fecha de acceso. http://www.conabio.gob.mx/invasoras/index.php/Especies_invasoras_-_Otros_invertebrados
Davidson, S. K. and Haygood, M. G. 1999. Identification of sibling species of the bryozoan Bugula neritina that produce different anticancer bryostatins and harbour distinct strains of the bacterial symbiont "Candidatus Endobugula sertula". Biological Bulletin 196: 273-280.
Davidson, S. K., Allen, S. W., Lim, G. E., Anderson, C. M. and Haygood, M. G. 2001. Evidence for the biosynthesis of bryostatins by the bacterial symbiont "Candidatus Endobugula sertula" of the bryozoan Bugula neritina. Applied Environmental Microbiology 67(10): 4531-4537.
Ghobashy, A.F.A. and El Komy, M.M. 1980. Fouling in the southern region of the Suez Canal, Aquatic Ecology 14(3): 179-185.
Hayes, K., Sliwa, C., Migus, S., McEnnulty, F., Dunstan, P. 2005. National priority pests: Part II Ranking of Australian marine pests. An independent report undertaken for the Department of Environment and Heritage by CSIRO Marine Research. http://www.marine.csiro.au/crimp/reports/PriorityPestsFinalreport.pdf
Hill, K., 2001. Bugula neritina. Smithsonian Marine Station at Fort Pierce. http://www.sms.si.edu/IRLSpec/Bugula_neriti.htm
Keough, M. J. and Chernoff, H. 1987. Dispersal and population variation in the bryozoan Bugula neritina Ecology 68(1): 199-210.
Keough, M. J. and Ross, J. 1999. Introduced fouling species in Port Phillip Bay. In Marine Biological Invasions of Port Phillip Bay Victoria: 9-11. L., H. C., Campbell, M. L., Thresher, R. E. and Martin, R. B. (Eds.). Hobart: CSIRO Marine Research.
Lynch, W. F. 1947. The behavior and metamorphosis of the larva of Bugula neritina (Linnaeus): experimental modification of the length of the free-swimming period and the responses of the larvae to light and gravity. Biological Bulletin 92: 115-150.
Lynch, W.L. 1947. The behavior and metamorphosis of the larva of Bugula neritina (Linnaeus): experimental modification of the length of the free-swimming period and the responses of the larvae to light and gravity, Biological Bulletin 92: 115-150 http://www.biolbull.org/cgi/reprint/92/2/115
Mackie, J. A., Keough, M. J. and Christidis, L. 2006. Invasion patterns inferred from cytochrome oxidase I sequences in three bryozoans, Bugula neritina, Watersipora subtorquata, and Watersipora arcuata. Marine Biology 149: 285-295.
Mackie, J.A., Keough, M.J. and Christidis, L. 2006. Invasion patterns inferred from cytochrome oxidase I sequences in three bryozoans, Bugula neritina, Watersipora subtorquata, and Watersipora arcuata, Marine Biology 149: 285–295
McEnnulty, F.R., Jones, T.E. and Bax, N.J. 2001. The Web-Based Rapid Response Toolbox. Retrieved 7 December 2006, from NIMPIS database. http://crimp.marine.csiro.au/NIMPIS/controls.htm
McGovern, T. and Hellberg, M. E. 2003. Cryptic species, cryptic endosymbionts, and geographic variation in chemical defenses in the bryozoan Bugula neritina. Molecular Ecology 12: 1207-1215.
National Introduced Marine Pest Information System (NIMPIS), 2002. Bugula neritina species summary. In: Hewitt, C.L., Martin, R.B., Sliwa, C., McEnnulty, F.R., Murphy, N.E., Jones, T. and Cooper, S. (eds). NIMPIS. Retrieved 7 December 2006, from NIMPIS database. http://www.marine.csiro.au/crimp/nimpis/spSummary.asp?txa=6929
NEMESIS (National Exotic Marine and Estuarine Species Information System). 2006. Bugula neritina. The Smithsonian Environmental Research Center. Retrieved 7 December 2006, from Chesapeake Bay Introduced Species Database http://invasions.si.edu/nemesis/CH-TAX.jsp?Species_name=Bugula%20neritina
NEMESIS (National Exotic Marine and Estuarine Species Information System). 2005. Bugula neritina -Invasion History. The Smithsonian Environmental Research Center. Retrieved 7 December 2006, from Chesapeake Bay Introduced Species Database. http://invasions.si.edu/nemesis/CH-INV.jsp?Species_name=Bugula+neritina
NEMESIS (National Exotic Marine and Estuarine Species Information System). 2004. Bugula neritina - Ecology. The Smithsonian Environmental Research Center. Retrieved 7 December 2006, from Chesapeake Bay Introduced Species Database. http://invasions.si.edu/nemesis/CH-ECO.jsp?Species_name=Bugula+neritina
NOBANIS (North European and Baltic Network on Invasive Alien Species). 2005. Bugula neritina. Retrieved 19 December 2006, from NOBANIS database. http://www.nobanis.org/NationalInfo.asp?countryID=DE&taxaID=5818
Perkol-Finkel, S. and Benayahu, Y. 2004. Recruitment of benthic organisms onto a planned artificial reef: shifts in community structure one decade post-deployment. Article in press. http://www.tau.ac.il/lifesci/departments/zoology/members/benayahu/documents/3aip.pdf
Piola, R.F. and Johnston, E.L. 2006. Differential resistance to extended copper exposure in four introduced bryozoans, Marine Ecology Progress Series 311: 103-114. http://www.int-res.com/articles/meps2006/311/m311p103.pdf
Prince William Sound Regional Citizens' Advisory Council. 2004. Non-indigenous Aquatic Species of Concern for Alaska. Fact Sheet 9. Single Horn Bryozoan. http://www.pwsrcac.org/docs/d0015800.pdf
Sutherland, J.P. and Karlson, R.H. 1977. Development and Stability of the Fouling Community at Beaufort, North Carolina, Ecological Monographs 47(4): 425-446.
USGS (United States Geological Survey). 2005. Bugula neritina. http://nas.er.usgs.gov/queries/collectioninfo.asp?NoCache=12%2F5%2F2006+4%3A59%3A44+PM&SpeciesID=266&State=&HUCNumber=
Virginia Museum of Natural History (VMNH), 2005. More Bryozoan Information. VMNH Virginia, USA.
Walters, L.J. 1992. Field Settlement Locations on Subtidal Marine Hard Substrata: Is Active Larval Exploration Involved?, Limnology and Oceanography 37(5): 1101-1107. http://aslo.org/lo/toc/vol_37/issue_5/1101.pdf
Winston, J.E. 1977. Distribution and Ecology of Estuarine Ectoprocts: A Critical Review, Chesapeake Science18(1): 34-57.
Wyatt, A.S.J., Hewitt, C.L., Walker, D.I. and Ward, T.J. 2005. Marine introductions in the Shark Bay World Heritage Property, Western Australia: a preliminary assessment, Diversity and Distributions 11(1): 33–44
ReferencesTop of page
Occhipinti-Ambrogi A, 2000. Recent developments in the history of the bryozoans of the lagoon of Venice: biodiversity and environmental stress. In: Proceedings of the 11th International Bryozoology Association Conference. Allen Press, 305-315.
ContributorsTop of page
Reviewed by: Dr. Josh Mackie, Invertebrate Zoology and Molecular Ecology Lab. Moss Landing Marine Laboratories. California USA
Principal sources: Bishop Museum 2002. Bugula neritina (Linnaeus, 1758). Guidebook of introduced marine species of Hawaii.
SMSFP 2001. Bugula neritina (Linnaeus, 1758). Smithsonian Marine Station at Fort Pierce.
Distribution MapsTop of page
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