Invasion History

First Non-native North American Tidal Record: 1872
First Non-native West Coast Tidal Record:
First Non-native East/Gulf Coast Tidal Record: 1872

General Invasion History:

Bugulina simplex was described from the Adriatic Sea by Hincks in 1886, and is probably native to that region, but even in the Mediterranean its records seem to be spotty. It is known from Barcelona Harbor, Spain (Pulpeiro and Gil 1995), but is missing from several local surveys in the Mediterranean. In Turkish waters, it is reported only from the Sea of Marmara and not from the Aegean or eastern Mediterranean coasts (Koçak, and Ayden Önen 2014). On the Atlantic coast of Europe, it was first found in the British Isles in 1958, and is now known from scattered populations from northern Spain to Scotland (Pulpeiro and Gil 1995; Ryland et al. 2011; Nall et al. 2014). This bryozoan may have been established in Woods Hole, Massachusetts as early as 1872, but was known chiefly from Eel Pond, near the Marine Biological Laboratory there, where it was long misidentified as B. flabellata (Verrill and Smith 1873; Rogick, in Smith 1964; Winston and Hayward 2012). By 2010, it had expanded its range from Long Island Sound to Casco Bay, Maine (McIntyre et al. 2011; Wells et al. 2014). It has also been introduced to the Azores, Australia, and New Zealand (Brock 1983; Gordon and Mawatri 1992; Cardigos et al. 2006). At most locations where it has been found, it is associated with artificial structures, including pilings and dock floats (Rogick, in Smith 1964; Ryland et al. 2011; Winston and Hayward 2012).

North American Invasion History:

Invasion History on the East Coast:

Early surveys from 1872, of fauna in the Woods Hole, Massachusetts region reported a common bryozoan, found mostly on pilings, as Bugulina flabellata (Verrill and Smith 1873; Sumner et al. 1913). Mary Rogick re-identified this form as B. simplex (Rogick and Croasdale 1949; Rogick, in Smith 1964). Its larval biology and settlement was extensively studied in Eel Pond by researchers at the Marine Biological Laboratory (see Wendt 2000), where it was abundant. It was collected in 1979-1982 in Great Bay, New Hampshire (Dijkstra and Harris 2009); in 2003 in Long Island Sound, Connecticut (Pederson et al. 2005); and in 2010 and 2013 in Casco Bay, Freeport, Maine (MacIntyre et al. 2011; Wells et al. 2014).

Invasion History Elsewhere in the World:

Bugulina simplex is believed to be native to the Mediterranean Sea. In the Atlantic waters of Europe, it was first found in 1957 in Holyhead, Wales, on the Irish Sea (Ryland 1958). Later records from the Atlantic coast are scattered, and mostly from artificial structures in harbors, such as dock floats, pilings, and sluice gates. These include occurrences from Riá de Ribadeo, Bay of Biscay, Spain in 1979 (Pulpeiro et al. 1983); Roscoff, Britany, France in 2002 (Ryland et al. 2011); several locations in Belgium and the Netherlands in 2000 (De Blauwe et al. 2001); and a number of marinas in England and Northern Ireland (Ryland et al. 2011; Nunn and Minchin 2013). The northernmost European records, so far, are from Kirkwall Marina, on Mainland Orkney, Scotland (59°N, Nall et al. 2014), and Lerwick, Shetlands in 2012 (Collin et al. 2015). Bugulina simplex is established in the Azores, where it is probably introduced (Morton and Britton 2000; Cardigos et al. 2006).

Aside from the coasts of Europe and northeastern North America, B. simplex is known from isolated locations in South America, Australia, and New Zealand. It is established in the Puerto Belgrano Naval Base, Argentina (Lichtschein de Bastida and Bastida1980, cited by Orensanz et al. 2002); and in Australia, from Adelaide Harbour (Brock 1985) and Port Philip Bay (Holmes 1982, cited by Hewitt et al. 2004). We know of only two records from New Zealand, a single colony from Lyttleton, South Island in 1988 (Gordon and Mawatari 1992, only encountered once) and an occurrence at Whangarei, North Island in 2003 (Ryland et al. 2011).


Colonies of Bugulina simplex form broad, fan-shaped, bushy tufts, about 300 mm tall. The branches are composed of 2-6 series of zooids which are widest at their origin. The zooids are large at 560-940 µm. The zooids usually bear one spine each on the inner and outer distal corners. Some specimens have two on the outer distal corner, while in specimens from New Zealand the corner spines are reduced or absent, but otherwise the colonies conform to the morphology of the species. Avicularia are attached to the marginal zooids only. Their length equals or exceeds the width of the zooid and their beaks curve downward. The ovicells are roughly hemispherical. The apertures are wide and open, but are occluded by a membrane during brooding. The colonies are yellow to yellow-orange. Description from: Rogick, in Smith 1964; Hayward and Ryland 1990; Ryland and Hayward 1991; Gordon and Mawatari 1992; and Winston and Hayward 2012.


Taxonomic Tree

Kingdom:   Animalia
Phylum:   Bryozoa
Class:   Gymnolaemata
Order:   Cheilostomata
Suborder:   Anasca
Family:   Bugulidae
Genus:   Bugulina
Species:   simplex


Potentially Misidentified Species

Bugulina flabellata
In early (pre-1960s) literature dealing with the Woods Hole area, Bugulina simplex was identified as 'B. flabellata' (Ryland and Hayward 1991).



Life History- Bugulina simplex is a bush-like, calcified bryozoan, composed of many individual zooids. The zooids feed by extending the ciliated tentacles of the lophophore as a funnel, creating a current, driving food particles into their mouths. The food is guided along the tentacles and through the pharynx by the cilia. Larger food particles can be moved or captured by flicking or contracting the tentacles. The zooids are hermaphroditic and produce large yolky eggs, which hatch into lecithotrophic larvae that are planktonic for short periods (less than 1 day) (Wendt 2000). Larvae settle on a substrate and metamorphose into the first zooid of a colony, an ancestrula (Barnes 1983).

Ecology- Bugulina simplex has been found on rocky and shelly bottoms in Vineyard Sound, Massachusetts but is much more frequently found on pilings and floats in harbors and marinas (Verrill and Smith 1873; Sumner et al. 1913; Gordon and Mawatari 1992; De Blawuwe and Faase 2001; Ryland et al. 2011).


Phytoplankton, detritus

Trophic Status:

Suspension Feeder



General HabitatMarinas & DocksNone
General HabitatRockyNone
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Tidal RangeSubtidalNone
Vertical HabitatEpibenthicNone

Tolerances and Life History Parameters

Maximum Duration1Maximum larval swimming period before metamorphosis (Wendt 2000)
Maximum Height (mm)30Winston and Hayward 2012
Broad Temperature RangeNoneCold temperate-Warm Temperate
Broad Salinity RangeNonePolyhaline-Euhaline

General Impacts

No impacts have been reported for Bugulina simplex in its presumed native and introduced ranges.

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
NA-ET3 Cape Cod to Cape Hatteras 1872 Def Estab
M010 Buzzards Bay 1872 Def Estab
NA-ET2 Bay of Fundy to Cape Cod 1982 Def Estab
N100 Casco Bay 2010 Def Estab
N170 Massachusetts Bay 2003 Def Estab
N130 Great Bay 1982 Def Estab
N180 Cape Cod Bay 2010 Def Estab
M040 Long Island Sound 2003 Def Estab
M020 Narragansett Bay 2010 Def Estab
M026 _CDA_M026 (Pawcatuck-Wood) 2010 Def Estab
N140 Hampton Harbor 2013 Def Estab
NEA-II None 1957 Def Estab
NEA-V None 1994 Def Estab
MED-II None 0 Crypto Estab
MED-III None 0 Crypto Estab
AUS-VII None 1983 Def Estab
SA-I None 1980 Def Estab
AUS-VIII None 1982 Def Estab
NEA-IV None 2002 Def Estab
NEA-III None 2004 Def Estab
MED-VII None 1886 Crypto Estab
NZ-IV None 8 Def Unk
MED-VIII None 0 Crypto Estab
MED-IV None 0 Crypto Estab
NEA-VI None 1998 Def Estab
N195 _CDA_N195 (Cape Cod) 1872 Def Estab
N185 _CDA_N185 (Cape Cod) 1913 Def Estab
AR-V None 2014 Def Estab
WA-I None 2014 Def Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude


Moisander; Pia ; Meghan Dailey; Daley, Meaghan D, M.S., Kelsey Garlick, M.S. and Katyanne M Shoemaker 11/8 2017 • Influence of nutrients on the benthic bloom-forming cyanobacterium <em>Hydrocoleum</em> sp. in the Nantucket Island. <missing URL>

Bishop, John D.D.; Wood, Christine A.; Yunnie, Anna L. E.; Griffiths, Carly A. (2015a) Unheralded arrivals: non-native sessile invertebrates in marinas on the English coast, Aquatic Invasions 10: 249-264

Brock, Brian J. (1985) Bryozoa: Ordovician to Recent, Olsen & Olsen, Fredensborg. Pp. 45-49

Carballo-Bolaños, Rodrigo Denis, Vianney Huang, Ya-Yi Keshavmurthy, Shashank Chen, Chaolun Allen (2019) Temporal variation and photochemical efficiency of species in Symbiodinaceae associated with coral Leptoria phrygia (Scleractinia; Merulinidae) exposed to contrasting temperature regimes, PLOSOne 14: e0218801

Cardigos, F. and 5 authors (2006) Non-indigenous marine species of the Azores., Helgoland Journal of Marine Research 60: 160-169

Chainho, Paula and 20 additional authors (2015) Non-indigenous species in Portuguese coastal areas, lagoons, estuaries, and islands, Estuarine, Coastal and Shelf Science <missing volume>: <missing location>

Collin, Samuel B.; Tweddle, Jacqueline F.; Shucksmith, Rachel J. (2015) Rapid assessment of marine non-native species in the Shetland Islands, Scotland, BioInvasions Records 4: In press

Corriero, G.; Longo, C.; Mercurio, M.; Marchini, A.; Occhipinti-Ambrogi, A. (2007) Porifera and Bryozoa on artificial hard bottoms in the Venice Lagoon: Spatial distribution and temporal changes in the northern basin, Italian Journal of Zoology 74(1): 21-29

Crivellaro, Marcelo Schuler; Candido, Davi Volney; ilveira, Thiago Cesar Lima; Fonseca, Adriana Carvalhal; Segal, Barbara ´ (2022) A tool for a race against time: Dispersal simulations to support ongoing monitoring program of the invasive coral Tubastraea coccinea, Marine Pollution Bulletin 185(114354): Published online

de Blauwe, Hans; Faasse, Marco (2001) Extension of the range of the bryozoans Tricellaria inopinata and Bugula simplex in the north-east Atlantic Ocean (Bryozoa: Cheilostomatida), Nederlandse Faunistiche Mededelingen 14: 103-112

Dijkstra, Jennifer A.; Harris, Larry G. (2009) Maintenance of diversity altered by a shift in dominant species: implications for species coexistence, Marine Ecology Progress Series 387: 71-80

Fehlauer-Ale, Karin H.; Winston, Judith E.; Tilbrook, Kevin J.; Nascimento, Karine B.; Vieira, Leandro M. (2015) Identifying monophyletic groups within Bugula sensu lato (Bryozoa, Buguloidea), Zoologica Scripta <missing volume>: <missing location>

Gittenberger, Arjan; Mirimin, Luca; Boyd, John; O’Beirn, Francis; Devine, Grainne; O’Brien, Martina; Rensing, Marjolein; O’Dwyer, Katie; Gittenberger, Edmund (2023) Marine Non-Indigenous Species Dynamics in Time and Space within the Coastal Waters of the Republic of Ireland, Diversity 15(1019): Published online

Gordon, Dennis P.; Mawatari, S.F. (1992) Atlas of marine-fouling bryozoa of New Zealand ports and harbors., Miscellaneous Publications of the New Zealand Oceanographic Institute 107: 1-52

Hayward, P.J.; Ryland, J. S. (1990) <missing title>, 2 Clarendon Press, Oxford. Pp. <missing location>

Hewitt, Chad L. and 14 authors. (2004) Introduced and cryptogenic species in Port Phillip Bay, Victoria, Australia., Marine Biology 144: 183-202

Kennedy, C., Pappal, A. L.; Bastidas, C.; ; Carlton, J. T.; David, A. A.; Dijkstra, J.A.; Duffey, S; Gibson, J.; Grady, S. P.; Green-Gavrielidis, (2020) Report on the 2018 Rapid Assessment Survey of Introduced, Cryptogenic, and Native Marine Species at New England Marinas: Massachusetts to Maine, <missing publisher>, Boston MA. Pp. <missing location>

Keppel, E. ; Ruiz, G. M.; Tovar–Hernández, M. A. (2020) Re-description of Parasabella fullo (Grube, 1878) (Polychaeta: Sabellidae) and diagnostic characteristics for detection in California, European Journal of Zoology 87(1): 105-115
DOI: 10.1080/24750263.2020.1721578

Koçak, Ferah; Aydin Önen, Sinem (2014) Checklist of Bryozoa on the coasts of Turkey, Turkish Journal of Zoology 38: 880-891

Liu, Wenliang; Liang, Xiaoli ; Zhu, Xiaojing (2015) A new record and mitochondrial identification of Synidotea laticauda Benedict, 1897 (Crustacea: Isopoda: Valvifera: Idoteidae) from the Yangtze Estuary, China, Zootaxa 4294: 371-380

Lopez Gappa, Juan (2000) Species richness of marine Bryozoa in the continental shelf and slope off Argentina (south-west Atlantic), Diversity and Distributions 6: 15-27

MacIntyre, Chris; Adrienne Pappal; Pederson, Judy; Smith, Jan P. (2011) Marine Invaders in the Northeast Rapid Assessment Survey of non-native and native marine species of floating dock communities, Massachusetts Coastal Zone Management, Boston MA. Pp. <missing location>

Micael, Joana; Tempera, Fernando; Berning, Björn; López-Fé, Carlos M.; Occhipinti-Ambrogi, Anna (2019) Shallow-water bryozoans from the Azores (central North Atlantic): native vs. non-indigenous species, and a method to evaluate taxonomic uncertainty, Marine Biodiversity 49: 469–480

Morton, B.; Britton, J.C. (2000) Origins of the Azorean intertidal biota: the significance of introduced species, survivors of chance events., Arquipelago. Life and Marine Sciences supplement 2 (part A): 29-51

Nall, Christopher R.; Guerin, Andrew J.; Cook, Elizabeth J. (2015) Rapid assessment of marine non-native species in northern Scotland and a synthesis of existing Scottish records, Aquatic Invasions 10(1): 107–121

New Zealand Ministry of Primary Industries 2023 Exotic Caulerpa seaweeds. <missing URL>

Nunn, Julia; Minchin, Dan 2013 Marine non-native invasive species in Northern Ireland. <missing URL>

Orensanz, Jose Maria and 14 other authors (2002) No longer the pristine confines of the world ocean: a survey of exotic marine species in the southwestern Atlantic, Biological Invasions 4(1-2): 115-143

Osburn, Raymond C. (1912) Bryozoa of the Woods Hole region, Fisheries Bulletin 30: 105-266

Pederson, Judith and 15 authors (2003) <missing title>, MIT Sea Grant College Program, Cambridge. Pp. <missing location>

Pulpeiro, E. Fernandez; Gil, O. Reverter (1995) Les Bryozoaires ctenostomes de la ria de Ferrol (Espagne nord-occidentale), Cahiers de Biologie Marine 36: 47-55

Ramalhosa, Patrício; Souto, Javier; Canning-Clode, João (2016) Diversity of Bugulidae (Bryozoa, Cheilostomata) colonizing artificial substratesin the Madeira Archipelago (NE Atlantic Ocean), Helgoland Marine Research <missing volume>: <missing location>

Rogick, Mary D.; Croasdale, Hannah (1949) Studies on marine Bryozoa, III. Woods Hole Region Bryozoa associated with algae, Biological Bulletin 96(1): 32-69

Rosso, Antonietta; Chimenz, Cararla; Balduzzi, Andrea (2010) Bryozoa, Biologia Marina Mediterranea 17(Suppl, 1): 589-615

Ryland, J. S. (1958) Bugula simplex hincks, a newly recognized polyzoan from British waters, Nature 181: 1146-1147

Ryland, John S. and 6 authors (2011) Alien species of Bugula (Bryozoa) along the Atlantic coasts of Europe, Aquatic Invasions 6(1): 17-31

Ryland, John S.; Hayward, Peter J. (1991) Marine flora and fauna of the Northeast United States: erect Bryozoa, NOAA Technical Report NMFS 99: 1-48

Smith, Ralph I. (1964) <missing title>, Systematics-Ecology Program, Marine Biological Laboratory, Woods Hole, Massachusetts. Pp. <missing location>

Sumner, Francis B.; Osburn, Raymond C.; Cole, Leon J.; Davis, Bradley M. (1913b) A biological survey of the waters of Woods Hole and vicinity Part II. Section III. A catalogue of the marine fauna Part II. Section IV. A catalogue of the marine flora, Bulletin of the Bureau of Fisheries 31: 539-860

Verrill, A.E.; Smith, S.I. (1873) <missing title>, 1 Report of the United States Commission of Fish and Fisheries, <missing place>. Pp. 1-757

Wells, Christopher D. and 23 authors (2014) Report on the 2013 rapid assessment survey of marine species at New England bays and harbors, Massachusetts Office of Coastal Zone Management, Boston MA. Pp. 32

Wendt, Dean E. (2000) Energies of larval swimming and metamorphosis in four species of Bugula (Bryozoa)., Biological Bulletin 198: 346-356

Winston, Judith E.; Hayward, Peter J. (2012) The marine bryozoans of the northeast coast of the United States: Maine to Virginia, Virginia Museum of Natural History Memoir 11: 1-180

Yale Peabody Museum of Natural History 2008-2016 YPM Invertebrate Zoology - Online Catalog. <missing URL>