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Tricellaria inopinata

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Tricellaria inopinata

Summary

  • Last modified
  • 08 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Preferred Scientific Name
  • Tricellaria inopinata
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Bryozoa
  •       Class: Gymnolaemata
  •         Order: Cheilostomatida
  • Summary of Invasiveness
  • T. inopinata forms erect, branched colonies, anchored to the substratum by rhizoids. The overall aspect of the colony is a whitish bush growing on hard substrata, often on mussel shells. Branches are dichotomousl...

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Pictures

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PictureTitleCaptionCopyright
T. inopinata colony growing on Mytilus galloprovincialis in the lagoon of Venice.
TitleColony
CaptionT. inopinata colony growing on Mytilus galloprovincialis in the lagoon of Venice.
CopyrightAnna Occhipinti Ambrogi
T. inopinata colony growing on Mytilus galloprovincialis in the lagoon of Venice.
ColonyT. inopinata colony growing on Mytilus galloprovincialis in the lagoon of Venice.Anna Occhipinti Ambrogi
Close-up of T. inopinata colony growing on Mytilus galloprovincialis in the lagoon of Venice
TitleColony
CaptionClose-up of T. inopinata colony growing on Mytilus galloprovincialis in the lagoon of Venice
CopyrightAnna Occhipinti Ambrogi
Close-up of T. inopinata colony growing on Mytilus galloprovincialis in the lagoon of Venice
ColonyClose-up of T. inopinata colony growing on Mytilus galloprovincialis in the lagoon of VeniceAnna Occhipinti Ambrogi
Scanning electron micrograph details of a Tricellaria inopinata colony from Ria de Aveiro.  (Note scale bar)  Picture details: Marchini A, Cunha MR, Occhipinti-Ambrogi A, 2007. First observations on bryozoans and enctoprocts in the Ria de Aveiro (NW Portugal) including the first record of the Pacific invasive cheilostome Tricellaria inopinata. Marine Ecology, 28(1): 154-160.
TitleSEM
CaptionScanning electron micrograph details of a Tricellaria inopinata colony from Ria de Aveiro. (Note scale bar) Picture details: Marchini A, Cunha MR, Occhipinti-Ambrogi A, 2007. First observations on bryozoans and enctoprocts in the Ria de Aveiro (NW Portugal) including the first record of the Pacific invasive cheilostome Tricellaria inopinata. Marine Ecology, 28(1): 154-160.
CopyrightAnna Occhipinti Ambrogi
Scanning electron micrograph details of a Tricellaria inopinata colony from Ria de Aveiro.  (Note scale bar)  Picture details: Marchini A, Cunha MR, Occhipinti-Ambrogi A, 2007. First observations on bryozoans and enctoprocts in the Ria de Aveiro (NW Portugal) including the first record of the Pacific invasive cheilostome Tricellaria inopinata. Marine Ecology, 28(1): 154-160.
SEMScanning electron micrograph details of a Tricellaria inopinata colony from Ria de Aveiro. (Note scale bar) Picture details: Marchini A, Cunha MR, Occhipinti-Ambrogi A, 2007. First observations on bryozoans and enctoprocts in the Ria de Aveiro (NW Portugal) including the first record of the Pacific invasive cheilostome Tricellaria inopinata. Marine Ecology, 28(1): 154-160.Anna Occhipinti Ambrogi

Identity

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Preferred Scientific Name

  • Tricellaria inopinata d'Hondt & Occhipinti Ambrogi, 1985

Summary of Invasiveness

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T. inopinata forms erect, branched colonies, anchored to the substratum by rhizoids. The overall aspect of the colony is a whitish bush growing on hard substrata, often on mussel shells. Branches are dichotomously divided at internodes (section between branch dichotomies) and are formed by series of alternating autozooids (single feeding bryozoan individuals). Each internode is formed by a variable number of autozooids (5-10). Autozooids vary in shape and size depending on their position within the internode, but all are more elongate and taper in their proximal part, progressively enlarging in the distal part. Lateral avicularia (specialized zooids) are large and prominent and bear a triangular mandible. Generally 3-4 spines are present on the outer margin of the zooid, and 2-3 on the inner one. The scutum (shield-like flattened, or branched spine partially covering the opening of some bryozoans) is highly variable in form and size within an individual colony. Globular ovicelles (brooding chambers) with pores are usually present in some autozooids within a colony.

The invasion by the exotic species T. inopinata has been a major event affecting the bryozoan assemblage of the Lagoon of Venice during the last twenty years (d’Hondt and Occhipinti-Ambrogi, 1985). The most likely vector of introduction into the Lagoon of Venice has been the involuntary transport as non target species, such as oysters shipped from the Pacific, or alternatively by naval traffic as a fouling organism. Found in 1982, in the Giudecca canal, in the centre of Venice, the species was firstly confined to the intertidal level, often attached to the byssus threads of Mytilus galloprovincialis.
 
Being tolerant to eutrophication, aerial exposure and swift currents due to tides and salinity variations, T. inopinata thrived best inside the Lagoon, where it became the most abundant species of the hard-bottom community at the intertidal and shallow subtidal level (Occhipinti-Ambrogi, 2000a). As a consequence, most native bryozoan species have declined in abundance or almost disappeared, whereas some previously rare species have gained relative importance. Soon after its discover in the lagoon of Venice it was found in the nearby Lagoon of Grado, whose environmental characteristics are very similar to those of Venice, and some years later in another brackish environment of the Atlantic coast (Poole Harbour, southern England) (Dyrynda et al., 2000), where it showed the same invasive behaviour and ecological requirements.
 
The specieshigh reproductive potential, its ability to grow on different substrates, coupled with its tolerance to variable environmental conditions (Occhipinti-Ambrogi, 1991), determined its successful colonisation of many other locations along the European Atlantic coasts. It is seldom found in exposed sites on the open coast.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Bryozoa
  •             Class: Gymnolaemata
  •                 Order: Cheilostomatida
  •                     Family: Candidae
  •                         Genus: Tricellaria
  •                             Species: Tricellaria inopinata

Notes on Taxonomy and Nomenclature

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Tricellaria inopinata was described after samples were discovered in 1982 for the first time in the Lagoon of Venice (d’Hondt and Occhipinti-Ambrogi, 1985). The species’ name derives from the Latin term inopinatus, which means unexpected, because this taxonwas unexpectedly found in the Giudecca Canal, in the very centre of the city of Venice, one of the most studied locations in the Adriatic Sea.

The genus is known only from the Pacific Ocean. Other species belonging to the genus Tricellaria and morphologically similar to T. inopinata have been previously found in the Pacific Ocean and described as Tricellaria occidentalis (originally known as Menipea occidentalisTrask, 1853) and T. occidentalis var. catalinensis (Robertson, 1905) from the Pacific coasts of the USA, and Tricellaria porteri (originally known as Menipea porteriMcGillivray, 1889) from Australia.

Description

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T. inopinata colonies are whitish to cream in colour and arborescent in form, up to 6-8 cm in length. Adult colonies are formed by branches, dividing dichotomously and composed of alternating series of autozooids (5-10 per internode) of 0.40 - 0.65 mm in length, usually tapering at the proximal part and becoming wider at the distal part (0.16-0.20 mm in width).

Lateral avicularia are large and prominent, 0.15-0.20 mm long, sometimes exceeding 0.30 mm, bearing a triangular mandible. Generally 3-4 spines on the distal margin of the zooid, 2-3 internal spines. The scutum is divided into two, three or more parts (cuneiform margin), usually small (0.09-0.14 mm in length, 0.14-0.19 mm in width). The globular ovicell, 0.15-0.16 mm in diameter, bearing pores (d’Hondt and Occhipinti-Ambrogi, 1985).
 
From eggs brooded in the ovicells originate short lived non-feeding (lecithotrophic) larvae (uniformly covered by cilia, cylindrical in shape, 0.11-0.12 mm wide). Larvae swim for a few hours and metamorphose immediately after the settlement on the substrate, giving origin to an ancestrula, which is the first zooid of a colony (0.28 mm long and 0.1 mm wide). Ancestrula rapidly fixes to substrate by two short rhizomes.
 
T. inopinata holotype is hosted in the Natural History Museum of Paris (MNHN-BRY-11596) and paratypes are hosted in the Natural History Museum of Venice (number: 10481), in the Natural History Museum of Milano (Number: r119) and in the Natural History Museum of Paris (MNHN-BRY-11597 to 11601).
 

Distribution

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The native distribution of T. inopinata is difficult to ascertain, as the debate on its taxonomic status has not reached a definite consensus on the nomenclature.

As mentioned in Notes on Taxonomy and Nomenclature, three other species of Tricellaria (T. occidentalis,T. occidentalis var. catalinensis and T. porteri) have been found along the Pacific coasts of the United States (Robertson, 1905; Osburn, 1950), in Japan (Yanagi and Okada, 1918), Australia (McGillivray, 1889; Bock, 1982; Brock, 1985) and New Zealand (Gordon and Mawatari, 1992) and are similar to T. inopinata.
 
The distinction between T. occidentalis and T. occidentalis var. catalinensis (Robertson 1905) was sustained by several authors (Yanagi and Okada, 1918; Okada, 1929; O’Donoghue and O’Donoghue, 1923; Osburn, 1950) who noted the morphological variability of the scutum within the individual colony. Silén (1941) concluded that since the two morphologies apparently integrated within a colony, T. occidentalis and T. occidentalis var. catalinensis were one and the same, the position was also retained by Soule et al. (1995).
 
The main differences between Tricellaria species are based on some morphological characteristics (number of autozooids between branch bifurcations, presence of marginal spines, scutum morphology, presence of lateral avicularia), which in some cases may present a certain degree of variability.
 
For that reason, in 1992, Gordon and Mawatari rejected T. inopinata as a new species and considered it as a junior synonym of T. occidentalis, stating that some of the morphological characteristics of the Mediterranean species fall within the range of variability of the Pacific one.
 
On the contrary, Breton and d’Hondt (2005) analysing the morphological shape and size of the scutum, excluded any possible source of confusion among the Mediterranean T. inopinata, the Pacific T. occidentalis and the Australian T. porteri.
 
The attempt to clarify the problem by Dyrynda et al. (2000) of comparing the type material and original description of the Pacific species did not produce any results as both the T. occidentalis andT. occidentalis var. catalinensis type material has been lost, and also the samples available in the different collections are not sufficient to allow complete comparisons. Only the morphological examination of large quantities of material from different locations both from European and Pacific coasts, coupled with molecular analysis will allow to clarify in a definitive way the identity and origin of the afore-mentioned Tricellaria species and confirm whether T. inopinata represents a portion of the genetic variability of one of the Pacific species, involuntarily introduced in the Lagoon of Venice.
 
From the nomenclatural point of view, the only detailed description, completed with type material, is that of the T. inopinata population by d’Hondt and Occhipinti-Ambrogi (1985) from the Lagoon of Venice.
 
T. inopinata n. sp. was described from the Lagoon of Venice: the possible route of invasion has been oysters transfer or, alternatively, the shipping traffic from the Pacific. In Italy, it is present also in the nearby Lagoon of Grado.
 
It was found along the Atlantic coast of the Iberian Peninsula, in the Ria de Ribadeo, in the Ria de Vigo, Spain, and in the Ria de Aveiro, Portugal. It is present in harbours and marinas in Belgium, the Netherlands and France, as well as in central and southern England from east Sussex to Cornwall.

Distribution Table

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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/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Sea Areas

Atlantic, NortheastWidespreadIntroduced1990s Invasive Arenas et al., 2006Growing on hard substrates, algae and other organisms in harbours and marinas
Mediterranean and Black SeaLocalisedIntroduced1982 Invasive d'Hondt and Occhipinti-Ambrogi, 1985Growing on hard substrates, algae and other organisms everywhere inside the lagoon

Europe

BelgiumWidespreadIntroduced2000 Invasive Blauwe and Faasse, 2001Harbours and marinas
FranceWidespreadIntroduced2000 Invasive Blauwe and Faasse, 2001Harbours and marinas
ItalyWidespreadIntroduced1982 Invasive d'Hondt and Occhipinti-Ambrogi, 1985On wooden piles and all other hard substrates in the main and secondary canals of the Lagoon of Venice
NetherlandsWidespreadIntroduced2000 Invasive Blauwe and Faasse, 2001Harbours and marinas
PortugalWidespreadIntroduced2004 Invasive Marchini et al., 2007In harbours and touristic marinas on pontoons, wooden piles and hard substrates
SpainWidespreadIntroduced1996 Invasive Fernández-Pulpeiro et al., 2001On docks and wharves in recreational and fishery ports at the intertidal level, attached on various organisms and on floating detached algae
UKWidespreadIntroduced1998 Invasive Arenas et al., 2006Ports, harbours and marinas

History of Introduction and Spread

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T. inopinata was firstly found near the centre of Venice in the Giudecca Canal, but after a few years it colonized all of the lagoon, except the areas with lower salinity (Occhipinti-Ambrogi, 2000b).

T. inopinata is a non-indigenous species introduced in Europe probably with oyster transfer from the Pacific. Belonging to a species-complex native to the United States Pacific coast (Robertson, 1905; Osburn, 1950), Japan (Yanagi and Okada, 1918; Mawatari, 1951), Australia (McGillivray, 1889; Bock, 1982; Brock, 1985), T. inopinata made its first appearence in the Lagoon of Venice in 1982 as the first non-Pacific location of a species belonging to genus Tricellaria (Occhipinti-Ambrogi 2000a). The first appearance of the species along the Atlantic coast was in 1996, in Spain, in the Ria de Ribadeo and in Galicia in the Ria de Vigo (Reverter-Gil and Fernández-Pulpeiro, 2001; Fernández-Pulpeiro et al., 2001). The species was then collected in central southern England for the first time in 1998 (Dyrynda et al., 2000) and in Belgium, the Netherlands and France, in 2000, mainly in harbours and marinas (De Blauwe and Faasse, 2001).
 
In 2004 a further extension of the species distribution was reported by Arenas et al. (2006) from east Sussex to Cornwall (UK) and by Marchini et al. (2007) in the Ria de Aveiro (Portugal).

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
Belgium 2000 Hitchhiker (pathway cause) Yes Blauwe and Faasse (2001) Possibly from UK
France 2000 Hitchhiker (pathway cause) Yes Blauwe and Faasse (2001) Possibly from UK
Italy 1982 Aquaculture (pathway cause) Yes d'Hondt and Occhipinti-Ambrogi (1985) The genus is known only in the Pacific, but the native location remains unknown due to systematic problems
Netherlands 2000 Hitchhiker (pathway cause) Yes Blauwe and Faasse (2001) Possibly from UK
Portugal 2004 Hitchhiker (pathway cause) Yes Marchini et al. (2007) Possibly from Spain
Spain Italy 1996 Aquaculture (pathway cause) Yes Fernández-Pulpeiro et al. (2001)
UK 1998 Aquaculture (pathway cause) ,
Hitchhiker (pathway cause)
Yes Dyrynda et al. (2000) Possibly from Italy or Spain

Risk of Introduction

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Ship traffic can be considered the principal vector facilitating long-distance and regional anthropogenic dispersal of many alien species, but in the case of T. inopinata the most likely form of secondary introduction from Venice to the Atlantic coasts of Spain was the movement of clams (Ruditapes philippinarum) between the two locations, as reported by Fernández-Pulpeiro et al. (2001). From Spain to the other Atlantic localities, both commercial and recreational vessels, as well as natural dispersion due to transport on living organisms (e.g. floating algae), are the most probably ways of expansion, accounting for the rapid colonization of the European Atlantic coasts from Portugal to the Netherlands (Dyrynda et al., 2000; De Blauwe and Faasse, 2001; Marchini et al., 2007).

Habitat

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The species is typical of the intertidal and shallow subtidal zone. It has been described from enclosed or sheltered habitats, like lagoons, estuaries, harbours and marinas. It has been seldom if ever found on the open coast.

In the Adriatic and northeast Atlantic species colonizes a variety of artificial hard substrata: floating pontoons, wooden piles, port highlights, vessel hulls, buoys and ropes. It also grows on other organisms.

Habitat List

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CategoryHabitatPresenceStatus
Brackish
Estuaries Principal habitat
Lagoons Principal habitat Harmful (pest or invasive)
Marine
Benthic zone Principal habitat Harmful (pest or invasive)
Inshore marine Principal habitat Harmful (pest or invasive)

Biology and Ecology

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Reproductive Biology

Species reproduction is both asexual, through zooidal budding (i.e. a new zooid buds from a already existing complete zooid), or sexual, with sperm and eggs developing in each zooid and originating, after fertilization, as embryos which are brooded in the ovicells.
 
Mature colonies with ovicells and embryos are found over much of the year, allowing a continuous recruiting population (Occhipinti-Ambrogi and d’Hondt, 1994) and a rapid colonization of the invaded habitat.
 
From the embryos originate the lecitotrophic larvae, whose low dispersal potential is thus not very effective in long-range transportation of the species, indicating alternative means of dispersal (mainly anthropogenic).
 
Nutrition
 
As with all the bryozoan species T. inopinata is a filter-feeding organism thriving on phytoplankton and other organic material such as bacteria and detritus, although no specific information on the quantity and quality of the food ingested is available.
 
Associations
 
The species has been found in association with macrophytes (Sargassum muticum, Undaria pinnatifida and Codium fragile), mussels and oysters (Mytilus galloprovincialis, Mytilus edulis, Crassostrea gigas), sponges, ascidians (Styela clava, Ascidiella aspersa, Styela plicata), polychaete worms (Ficopomatus enigmaticus, Hydroides dianthus), other bryozoans (Bugula neritina) and many other organisms constituting hard bottom communities in lagoons and harbours.

Climate

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ClimateStatusDescriptionRemark
C - Temperate/Mesothermal climate Preferred Average temp. of coldest month > 0°C and < 18°C, mean warmest month > 10°C

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
54 45

Air Temperature

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Parameter Lower limit Upper limit
Absolute minimum temperature (ºC) 3 34

Water Tolerances

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ParameterMinimum ValueMaximum ValueTypical ValueStatusLife StageNotes
Depth (m b.s.l.) Optimum Intertidal
Salinity (part per thousand) 28 30 Optimum 20-35 tolerated
Water temperature (ºC temperature) 20 25 Optimum 3-34 tolerated

Means of Movement and Dispersal

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The undetermined Pacific origin of the species (belonging to the species complex T. porteri-occidentalis-inopinata) suggests that the first Mediterranean introduction was due to accidental introduction (oyster import). Secondary introductions are likely to have been caused by accidental introductions associated with small vessels (small commercial vessels, yachts, fishing boats in coastal routes), natural dispersion (currents) or vector transport (on floating fragments of Sargassum).

Pathway Causes

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CauseNotesLong DistanceLocalReferences
Aquaculture Yes Occhipinti-Ambrogi, 2000a
Flooding and other natural disastersE.g. debris from the 2011 Tohoku tsunami carried individuals from the Japanese coast to Oregon Yes

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
AircraftAdult colonies Yes Yes
Aquaculture stock Yes Yes Occhipinti-Ambrogi, 2000a
Floating vegetation and debrisE.g. debris from the 2011 Tohoku tsunami carried individuals from the Japanese coast to Oregon Yes Yes Blauwe and Faasse, 2001
Live seafood Yes Yes Occhipinti-Ambrogi, 2000a
Ship hull fouling Yes Dyrynda et al., 2000

Impact Summary

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CategoryImpact
Environment (generally) Negative

Economic Impact

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It is a rapidly growing fouling organism, settling on vessels, pontoons, buoys and ropes and all other kinds of submerged organisms at the intertidal and upper subtidal levels.

Environmental Impact

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Impacts on Biodiversity

The arrival and spread of T. inopinata in the Venice Lagoon caused a decline in the abundance of some bryozoan species with the same colony form, such as Bugula stolonifera and Bugula neritina. On the other hand, other species as Buskia socialis and Bowerbankia gracillima, previously considered rare species within the Lagoon, gained relative importance (Occhipinti-Ambrogi, 2000a,b).

Specialized bryozoan species, occupying well-defined sectors of the Lagoon with peculiar environmental characteristics, declined in abundance, whereas T. inopinata, a generalist inside the lagoon, was found almost everywhere. When the abundance of T. inopinata peaked, bryozoan assemblages were uniform throughout the Lagoon, with the exception of few localities.

Threatened Species

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Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
Bugula neritina (brown bryozoan)No DetailsItalyCompetitionOcchipinti-Ambrogi, 2000b
Bugula stoloniferaNo details No detailsItalyCompetitionOcchipinti-Ambrogi, 2000b

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Highly adaptable to different environments
  • Is a habitat generalist
  • Tolerant of shade
  • Highly mobile locally
  • Benefits from human association (i.e. it is a human commensal)
  • Fast growing
  • Has high reproductive potential
  • Gregarious
  • Has propagules that can remain viable for more than one year
  • Reproduces asexually
Impact outcomes
  • Ecosystem change/ habitat alteration
  • Infrastructure damage
  • Modification of natural benthic communities
  • Modification of successional patterns
  • Reduced native biodiversity
  • Threat to/ loss of native species
Impact mechanisms
  • Competition - monopolizing resources
  • Competition
  • Fouling
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally
  • Difficult to identify/detect as a commodity contaminant
  • Difficult to identify/detect in the field

Similarities to Other Species/Conditions

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A possible misunderstanding occurred between genus Tricellaria and Bugulopsis due to general colony appearance and branch disposition (d’Hondt and Occhipinti-Ambrogi, 1985). Morphological features such as the scutum, avicularia and vibracularia present in Tricellaria made it distinguishable, under the microscope, from Bugulopsis.

T. inopinata from the Adriatic coast and northeast Atlantic is morphologically similar to T. occidentalis from Japan and New Zealand and to T. porteri from Australia (see Distribution).

Prevention and Control

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Aquaculture imports should be strictly quarantined, and only second generation spat should be released; special care should be taken when aquaculture facilities are located in the vicinity of ports and marinas.

Gaps in Knowledge/Research Needs

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  • Genetic information from populations of Tricellaria in different locations could highlight the actual areas of origin and clarify the taxonomic questions.
  • Physiological investigations could explain the habitat preferences for sheltered habitats.
  • Pathways for introduction could be clarified by examination of likely vectors such as shipment of molluscs or hull fouling.

References

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Arenas F; Bishop JDD; Carlton JT; Dyrynda PJ; Farnham WF; Gonzalez DJ; Jacobs MW; Lambert C; Lambert G; Nielsen SE; Pederson JA; Porter JS; Ward S; Wood CA, 2006. Alien species and other notable records from a rapid assessment survey of marinas on the south coast of England. Journal of Marine Biological Association, 86:1329-1337.

Blauwe HDe; Faasse M, 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.

Bock PE, 1982. Bryozoans (Phylum Bryozoa). In: Marine Invertebrates of Southern Australia, part 1 [ed. by Shepherd, S. A. \Thomas, I. M.]. South Australia: Government Printer, 319-394.

Breton G; d'Hondt J-L, 2005. [English title not available]. (Tricellaria inopinata d'Hondt et Occhipinti-Ambrogi, 1985 (Bryozoa: Cheilostomatida) dans le Port du Havre (Manche Orientale).) Bulletin de la Société géologique de Normandie Amis Musée du Havre, 91(2):67-72.

Brock BJ, 1985. South Australian fouling bryozoans. In: Bryozoa: Ordovician to recent [ed. by Nielsen, C. \Larwood, G. P.]. Fredensborg: Olsen and Olsen, 45-49.

DAISIE, 2006. Delivering Alien Invasive Species Inventories for Europe. http://www.europe-aliens.org/

d'Hondt JL; Occhipinti-Ambrogi A, 1985. [English title not available]. (Tricellaria inopinata, n., un nouveau Bryozoaire Chilostome de la faune méditerranéenne.) Marine Ecology, 6(1):35-46.

Dyrynda PEJ; Fairall VR; Occhipinti-Ambrogi A; d'Hondt JL, 2000. The distribution, origin and taxonomy of Tricellaria inopinata d'Hondt and Occhipinti-Ambrogi, 1985, an invasive bryozoan new to the Atlantic. Journal of Natural History, 34:1993-2006.

Fernández-Pulpeiro E; César-Aldariz J; Reverter-Gil O, 2001. [English title not available]. (Sobre la presencia de Tricellaria inopinata d'Hondt & Occhipinti-Ambrogi, 1985 (Bryozoa, Cheilostomatida) en el litoral gallego (N. España).) Nova Acta Científica Compostelana (Bioloxía), 11:207-213.

Gordon PD; Mawatari SF, 1992. Atlas of marine-fouling Bryozoa of New Zealand ports and harbours. Miscellaneous Publications of New Zealand Oceanographic Institute, 107:1-52.

IBA, 2009. International Bryozoology Association. http://www.bryozoa.net/

Marchini A; Cunha MR; Occhipinti-Ambrogi A, 2007. First observations on bryozoans and enctoprocts in the Ria de Aveiro (NW Portugal) including the first record of the Pacific invasive cheilostome Tricellaria inopinata. Marine Ecology, 28(1):154-160.

Mawatari SF, 1951. On Tricellaria occidentalis, (Trask), one of the fouling bryozoans in Japan. Miscellaneous Report of the Research Institute for Natural Resources, 22:9-16.

McGillivray PH, 1889. On some Australian Polyzoa. Transactions and Proceedings and Report of the Royal Society of South Australia, 12:24-30.

Occhipinti-Ambrogi A, 1991. [English title not available]. (The spread of Tricellaria inopinata into the lagoon of Venice: an ecological hypothesis. Bulletin de la Société de Science Naturelles de l'Ouest France Mémoires Hors Série.) In: Bryozoaires actuels et fossiles: Bryozoa living and fossil, 1 [ed. by Bigey, F. P.]. 299-308.

Occhipinti-Ambrogi A, 2000. Biotic invasions in a Mediterranean Lagoon. Biological Invasions, 2:167-176.

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.

Occhipinti-Ambrogi A; d'Hondt JL, 1994. The invasion ecology of Tricellaria inopinata into the lagoon of Venice: morphological notes on larva and the ancestrula. In: Biology and Palaeobiology of Bryozoans [ed. by Hayward, P. J. \Ryland, J. S. \Taylor, P. D.]. Fredensborg, Danmark: Olsen & Olsen, 139-144.

Okada Y, 1929. Report of the Biological Survey of Muts Bay, 12, Cheilostomatous Bryozoa of Muts Bay. Sciences Report of the Tohoku imperial University, 4(4):11-35.

Olsen; Olsen; FredensboO'Donoghue CH; O'Donoghue E, 1923. A preliminary list of Polyzoa (Bryozoa) from the Vancouver Island region. Contribution to Canadian Biology, 1:145-201.

Osburn RC, 1950. Bryozoa of the Pacific Coast of America. Part 1, Cheilostomata-Anasca. Allan Hancock Pacific Expedition, 14:1-269.

Reverter-Gil O; Fernández-Pulpeiro E, 2001. [English title not available]. (Inventario y cartografia de los Briozoos marinos de Galicia (N. de España).) Monografías de Nova Acta Científica Compostelana,Serie Bioloxía, 1. 1-243.

Robertson A, 1905. Non-incrusting Cheilostomatous Bryozoa of the West Coast of North America. University of California Publication, Zoology, 2(5):235-322.

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Links to Websites

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WebsiteURLComment
DAISIE Delivering Alien Invasive Species Inventories for Europehttp://www.europe-aliens.org/index.jsp
International Bryozoology Association (IBA)http://www.bryozoa.net/

Contributors

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18/10/09 Original text by:

Anna Occhipinti Ambrogi, Director Department "Ecologia del Territorio", Via S.Epifanio4, I-27100 Pavia, Italy

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