Invasion History

First Non-native North American Tidal Record: 2010
First Non-native West Coast Tidal Record:
First Non-native East/Gulf Coast Tidal Record: 2010

General Invasion History:

Tricellaria inopinata, an introduced species in the Mediterranean, was first described from the Venice Lagoon, Italy (d'Hondt and Occhipinti Ambrogi 1985). It is doubtless native to the Pacific Ocean, where one or more very closely related (or the same) species occurs, under additional names, such as T. porteri, T. occidentalis, and T. occidentalis catalinensis (Dyrynda et al. 2000). In 1982, T. inopinata was found in the Venice Lagoon (d'Hondt and Occhipinti Ambrogi 1985), in 1998 it was found in Poole Harbor, on the English Channel (Dyrynda et al. 2000), and in 2000, it was found on the Atlantic coast of Spain (Pulpeiro et al. 2002), and the coasts of Belgium, the Netherlands, Germany and Norway (de Blauwe and Faase 2001; Buschbaum et al. 2011; Cook et al. 2013; Porter et al. 2015). In Europe, it currently ranges north to Norway, west to the Atlantic coasts of Scotland and Ireland, and south to Cadiz, Spain (Buschbaum 2012; Cook et al. 2013). In 2010, this bryozoan was found on the Atlantic coast of the US, in Eel Pond, Woods Hole, Massachusetts where it is now established (Johnson et al. 2012), and since then it has been found north to the Boston region (James T. Carlton, personal communication 2013).  Tricellaria spp. washed ashore in central Oregon in 2012 on a floating dock which was swept away from the Japanese coast during the 2011 tsunami, and was identified morphologically as T. inopinata, suggesting that Japan is the native region (McCuller and Carlton 2018).

North American Invasion History:

Invasion History on the East Coast:

In 2010, Tricellaria inopinata was discovered in Eel Pond, a small lagoon adjacent to Woods Hole Harbor, Massachusetts (MA) (Johnson et al. 2012). The population expanded in 2011, displacing native arborescent bryozoans (Johnson et al. 2012). It was subsequently found in the Boston region, in Boston Harbor, Marblehead and Gloucester, where it is also now established (collections by Adrienne Pappal; James T. Carlton, personal communication; Johnson and Woollacott 2015). Genetic analysis indicates that the population in Boston Harbor was probably introduced from the Eel Pond population, but the Marblehead, and Gloucester populations were introduced independently (Johnson and Woollacott 2015). In a 2013 survey, it had extended its range southward to the mouth of Narragansett Bay, Rhode Island and northward to Hampton River, New Hampshire (Wells et al. 2014).

Invasion History Elsewhere in the World:

Tricellaria inopinata was first discovered in the lagoon of Venice, at the head of the Adriatic Sea, a region with a well-studied fauna, in 1982. This bryozoan was found associated with wooden pilings and other man-made structures, and rapidly spread within the lagoon (d'Hondt and Occhipinti Ambrogi 1985; Occhipinti Ambrogi and d'Hondt 1994). In 1998, it was discovered in the Marano Lagoon, 70 km to the east (Dyrynda et al. 2000).

In 1998, T. inopinata was found in Poole Harbor, England, on the English Channel. It is currently known in the British Isles from Cornwall to Scotland, and Ireland, in harbors of the Atlantic, Irish Sea, English Channel, and North Sea coasts (Dyrynda et al. 2000; Arenas et al. 2006; Cook et al. 2013). Tricellaria inopinata was found at a number of locations on the coast of the Netherlands and Belgium in 2000 (de Blauwe and Faase 2001), at Le Havre, France in 2003 (Breton and d'Hondt 2006) and as far east as the German Wadden Sea coast (2009, Buschbaum 2012; Cook et al. 2013). In 2014, it was found in the harbors of Floro and Kristiansund, Norway (61-63 N, Porter et al. 2014). Further south it was collected in Brittany, at Hendaye, France, near the Spanish border (in 2000, de Blauwe and Faase 2001, in 2010, De Blauwe 2010), in Galicia, Spain (Pulpeiro et al. 2001), in the Ria de Aveiro, Portugal (in 2004, Marchini et al. 2007). Its current southern limit on the Atlantic coast is Cadiz, Spain (Cook et al. 2013; Ros et al. 2013). In Europe, the sites where T. inopinata has been found are often near sites of oyster culture (Pacific Oyster, Crassostrea gigas), though secondary transport by recreational and fishing boats is likely (Galil and Occhipinti Ambrogi 2006).


Tricellaria inopinata forms erect, bushy, branched colonies which are cream-to-buff in color and attached to hard substrates by rhizoids. The stem is composed of two rows of autozooids, with the joints between zooids in one row, roughly corresponding to the midpoint of the zooids in the other row. Internodes (branching points) occur every 3-19 zooids (odd numbers only, most commonly 3, 5 and 7). Each zooid is about 450-650 µm, with an oval opening (opesia) about 200-280 µm long, and 150-200 µm wide. A third zooid occurs in each internode, between the two branches. The number of zooids between internodes increases towards the tips of the branches. The autozooids are tapered at their proximal end, but enlarged at the distal end. Avicularia (defensive zooids) project from the sides of the branches, extending 150-200 µm on each side, and each bearing a triangular mandible. Three to four spines are present on the outer margins of the zooid and 2-3 spines on the inner margin. The most proximal of the external spines often has double tips. The middle zooid in the internodes bears a large median spine, and has at its base a small opening. The scutum is shield-like, with a flattened spine which partially covers the opening (opesium). It varies greatly in shape and size within the colony, sometimes with two or three lobes. Some autzooids in their colony usually have globular ovicells (brooding chambers), located distal to the opesium, and 150-160 µm in diameter (description from: d'Hondt and Occhipinti-Ambrogi 1985; Dyrynda et al. 2000; Johnson et al. 2012).

Among the features which distinguish T. inopinata from T. occidentalis and T. porteri are the variability of the scutum, the number of zooids between internodes, and the number of spines in the angles between the joints. Tricellaria inopinata has 3-19 zooids between internodes, T. occidentalis has 3-5, and T. porteri has 3-13. In T. inopinata, the proximal external spine of the zooids is sometimes bifid (two-tipped), but is never bifid in the other two species. A single median spine is present in the angle between branches in T. inopinata, while the other two species have two spines, with a space between. In T. inopinata, the scutum is highly variable, from a slender projection, to an antler-like structure, while in T. occidentalis, it is spatula-like, and in T. porteri it is kidney-shaped (description from: d'Hondt and Occhipinti-Ambrogi 1985; Dyrynda et al. 2000; Johnson et al. 2012).

A bryozoan from the West Coast of the US, described as T. occidentalis var. catalinensis (Robertson 1905) is similar to T. inopinatas (Dyrynda et al. 2000), but Winston and Ryland (2012) note differences in the description relative to ovicell shape, pore pattern, and spine number and position.


Taxonomic Tree

Kingdom:   Animalia
Phylum:   Bryozoa
Class:   Gymnolaemata
Order:   Cheilostomata
Suborder:   Anasca
Family:   Scrupocellariidae
Genus:   Tricellaria
Species:   inopinata


Potentially Misidentified Species

Tricellaria occidentalis
Northeast Pacific species (Dyrynda et al. 1993)

Tricellaria porteri
Southwest Pacific-Indian Ocean (Australia) (Dyrynda et al. 1993)



Life History- Tricellaria inopinata 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, and 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, which are planktonic for short periods (a few hours). Larvae settle on a substrate and metamorphose into the first zooid of a colony, an ancestrula (Barnes 1983; Occhipinti Ambrogi and d'Hondt 1994).

Ecology- Tricellaria inopinata grows attached to wood pilings, floats, rocks, seaweeds, seagrasses, oysters, mussels and other hard substrates It has been found attached to other exotic fouling species, including the seaweeds Sargassum muticum and Undaria pinnatifida, and the tunicate Styela clava (d'Hondt & Occhipinti Ambrogi 1985; Dyrynda et al. 2000). It has a strong preference for sheltered locations, and tolerates some variation in salinity (Occhipinti Ambrogi and d'Hondt 1994; Dyrynda et al. 2000).



Trophic Status:

Suspension Feeder



General HabitatMarinas & DocksNone
General HabitatRockyNone
General HabitatVessel HullNone
General HabitatOyster ReefNone
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Vertical HabitatEpibenthicNone

Life History

Tolerances and Life History Parameters

Minimum Temperature (ºC)2Field (Galil and Occhipinti-Ambrogi 2006; Johnson et al. 2012)
Maximum Temperature (ºC)34.5Field (Galil and Occhipinti-Ambrogi 2006)
Minimum Salinity (‰)20Field (Galil and Occhipinti-Ambrogi 2006)
Maximum Salinity (‰)35Field (Galil and Occhipinti-Ambrogi 2006)
Maximum Duration0.2Settles after a few hours (Occhipinti Ambrogi and d'Hondt 1994).
Maximum Height (mm)40Colony height, Johnson et al. 2012

General Impacts

Economic Impacts

Shipping/boating- Tricellaria inopinata probably contributes to fouling of recreational boats, docks, ropes, etc. (Dyrynda et al. 2000; Galil and Occhipinti-Ambrogi 2006), but its economic impacts cannot be easily separated from those of the rest of the fouling community.

Ecological Impacts

Competition- Since its invasion in the Lagoon of Venice, Tricellaria inopinata has largely replaced other species of arborescent bryozoans, reducing diversity within this community (Occhipinti-Ambrogi 2000; Occhipinti-Ambrogi and Savini 2003; Galil and Occhipinti-Ambrogi 2006). Preliminary observations in Eel Pond, Woods Hole, Massachusetts, indicate that this bryozoan is competing with native arborescent bryozoans (ICES 2012, in prep).

Regional Impacts

NA-ET3Cape Cod to Cape HatterasEcological ImpactCompetition
Preliminary observations in Eel Pond (Woods Hole, MA) indicate that T. inopinata has negatively impacted other Eel Pond arborescent bryozoans (ICES 2012, in prep.).
M010Buzzards BayEcological ImpactCompetition
Preliminary observations in Eel Pond (Woods Hole, MA) indicate that T. inopinata has negatively impacted other Eel Pond arborescent bryozoans (ICES 2012, in prep.).
MED-VIINoneEcological ImpactCompetition
Tricellaria inopinata has replaced most of the native arborescent bryozoans in the Venice Lagoon, resulting in a reduction of biodiversity (Occhipinti Ambrogi 2000; Occhipinti Ambrogi and Savini 2003).
NEA-IINoneEconomic ImpactShipping/Boating
Fouling impacts have been listed for the British Isles (Minchin et al. 2013)
NEA-VNoneEcological ImpactHabitat Change
Tricellaria inopinata provided habitat for at least 19 native and introduced species in Puerto América, Cádiz, Spain. The introduced species Caprella scaura, Monocorophium sextonae (Amphipoda) and Paracerceis sculpta (Isopoda). However, its effects may not differ from other temperate bryozoans, such as Bugula neritina (Gavira-O’Neill et al. 2019)
MAMassachusettsEcological ImpactCompetition
Preliminary observations in Eel Pond (Woods Hole, MA) indicate that T. inopinata has negatively impacted other Eel Pond arborescent bryozoans (ICES 2012, in prep.).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
MED-VII None 1982 Def Estab
NEA-II None 1998 Def Estab
NA-ET3 Cape Cod to Cape Hatteras 2010 Def Estab
M010 Buzzards Bay 2010 Def Estab
NEA-III None 2004 Def Estab
NEP-VI Pt. Conception to Southern Baja California 1905 Crypto Estab
NEP-III Alaskan panhandle to N. of Puget Sound 0 Crypto Estab
NZ-IV None 0 Crypto Estab
AUS-VII None 0 Crypto Estab
NEA-V None 2000 Def Estab
NEA-IV None 2010 Def Estab
NA-ET2 Bay of Fundy to Cape Cod 2013 Def Estab
N170 Massachusetts Bay 2013 Def Estab
N140 Hampton Harbor 2013 Def Estab
N180 Cape Cod Bay 2013 Def Estab
M020 Narragansett Bay 2013 Def Estab
AR-V None 2014 Def Estab
NEA-VI None 2015 Def Estab
MED-VI None 2015 Def Estab
MED-II None 2015 Def Estab
MED-III None 2018 Def Estab
WA-I None 2014 Def Estab
M023 _CDA_M023 (Narragansett) 2019 Def Estab
M040 Long Island Sound 2019 Def Estab
M060 Hudson River/Raritan Bay 2019 Def Estab
N130 Great Bay 2018 Def Estab
N100 Casco Bay 2018 Def Estab
M026 _CDA_M026 (Pawcatuck-Wood) 2022 Def Estab
NWP-3b None 0 Native Estab
NWP-4b None 0 Native Estab
MED-VIII None 2015 Def Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude


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