Tricellaria inopinata

Overview

Scientific Name: Tricellaria inopinata

Phylum: Bryozoa

Class: Gymnolaemata

Order: Cheilostomatida

Family: Candidae

Genus: Tricellaria

Species:

inopinata [Describe here as A. iricolor]

Native Distribution

Origin Realm:

Temperate Northern Pacific, Central Indo-Pacific

Native Region:

Origin Location:

CONFLICT: Northwest and northeast Pacific Temperate Northern Pacific [Japan] (NEMESIS 2016; Occhipinti-Ambrogi and Savini 2003; Calder et al 2014) STATUS STATED Native range: temperate Pacific from North America, Japan to Australia - undetermined Pacific origin of the species (belonging to the species complex T. porteri-occidentalis-inopinata) (Galil and Occhipinti-Ambrogi 2009) STATUS STATED It probably originates from the Pacific coast of North-America (Occhipinti Ambrogi and d'Hondt 1994, cited in De Blauwe and Faasse 2001; Rilov and Galil 2009) STATUS STATED Uncertain realm Native range: temperate Pacific from North America, Japan to Australia - undetermined Pacific origin of the species (belonging to the species complex T. porteri-occidentalis-inopinata) (Galil and Occhipinti-Ambrogi 2009) STATED Indo-Pacific origin (Marchini et al 2015; Occhipinti-Ambrogi 2000) STATUS NOT STATED Uncertain, assumed to be introduced from the north Pacific (NEMESIS 2016; CABI 2006; Porter et al 2015; Bishop et al 2015a; Cook et al 2013a; Lejeusne et al 2014) RELATED: Temperate Northern Pacific [T. occidentalis] Cape Flattery, Washington to Santa Barbara. (Trask 1857, cited in Osburn 1950) STATUS NOT STATED [T. occidentalis] From British Columbia (Hincks 1882, 1884, cited in Grischenko et al. 2007; O'Donoghue & O'Donoghue 1923, cited in Grischenko et al. 2007) to Baja California, Mexico (Robertson 1905, cited in Grischenko et al. 2007; Soule et al. 1995, cited in Grischenko et al. 2007) STATUS NOT STATED [T. occidentalis] With Reference to Robertson and O'Donoghue, Mawatari (1951) stated that T. o was recorded as far south as San Diego and from Queen Charflotte Islands. STATUS NOT STATED

Geographic Range:

[Western Pacific] Japan; Australia and New Zealand (NEMESIS 2016; Occhipinti-Ambrogi and Savini 2003; Connell 2001; Porter et al 2015; Bishop et al 2015b) [Eastern Pacific] Canada and US (NEMESIS 2016) [Western Atlantic] Canada and US; Narragansett Bay, Rhode Island to Hampton River, New Hampshire (NEMESIS 2016) [Eastern Atlantic] Norway; Scotland and Ireland, and south to Cadiz, Spain (Buschbaum 2012 and Cook et al 2013a, cited in NEMESIS 2016) [Mediterranean Sea] (Galil and Occhipinti-Ambrogi 2009) -2 43.2999992370605,12.4000005722046 51.8000030517578 (OBIS 2016)

General Diversity:

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 (CABI 2016) Genus only known 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 and T. occidentalis var. catalinensis from the Pacific coasts of the USA and Tricellaria porteri from Australia (CABI 2016; Occhipinti-Ambrogi 2000)) Member of the T. occidentalis complex, which also includes T. occidentalis and T. porteri (Cook et al 2013a; Gordon and Mawatari 1992, cited in Dyrynda et al 2000) Undetermined Pacific origin of the species (belonging to the species complex T. porteri-occidentalis-inopinata) (Galil and Occhipinti-Ambrogi 2009)

Non-native Distribution

Invasion History:

Yes, see inv_propens

Non-native Region:

Northwest Atlantic, Northeast Atlantic, Mediterranean Sea, Northwest Pacific, Northeast Pacific, Australia and New Zealand

Invasion Propens:

CONFLICT: Northwest and northeast Pacific Temperate Northern Pacific [Canada] East coast of Canada (NEMESIS 2016; Galil and Occhipinti-Ambrogi 2009; Johnson et al 2012, cited in Bishop et al 2015b) *Introduced Invaded the north of Japan to Taiwan (Occhipinti Ambrogi and d'Hondt 1994, cited in De Blauwe and Faasse 2001; Rilov and Galil 2009) *Introduced Introduced into Japan, Taiwan and the West Pacific (Occhipinti Ambrogi and d'Hondt 1994, cited in Porter et al 2015) *Introduced Cryptogenic along west coast of Canada/US, Japan (NEMESIS 2016) *Cryptogenic Temperate Northern Atlantic [Mediterranean] (Streftaris & Zenetos 2006) *Noted as one of the 100 worst invasive species [Atlantic Europe] 2000s: found on the Atlantic coast of Spain, Portugal and coasts of Belgium, Netherlands, Germany, France, and Norway (Pulpeiro et al 2002, de Blauwe and Faase 2001, Buschbaum et al 2012; NEMESIS 2016; Bishop et al 2015a; Cook et al 2013a; Marchini et al 2007, Breton and d'Hondt 2005 and Markert et al 2015, cited in Porter et al 2015; De Blauwe and Faasse 2001; De Blauwe 2005; Johnson and Woollacott 2015; Kerckhof et al 2007; Ramalhosa and Canning-Clode 2015; Ros and Guerra-Garcia 2012; Ros et al 2014) *Introduced [UK] 1998-2006: Present in south-central England; Observed in Dublin Bay, Ireland ; British Isles from Cornwall to Scotland; Second North Atlantic record at Poole Harbour and has spread out since (Dyrynda et al 2000; Kelso and Wyse Jackson 2012; Porter et al 2015; Dyrynda 2005; Bishop et al 2015a; NEMESIS 2016; CABI 2016; Cook et al 2013a; Nall et al 2015) *Introduced [Scotland] 2011-2012: observed in Scotland marinas and Dublin Bay, Ireland (Kelso and Wyse Jackson 2012, cited in Porter et al 2015; Cook et al 2013a) *Introduced and established [Norway] 2014: Norwegian waters at the ports of Florø and Kristiansund (Porter et al 2015) *Introduced [Germany] 2009: new to the northern and southern German Wadden Sea (Buschbaum et al 2012) *Introduced [Italy] 1982: First described in 1985 from specimens collected in May 1982 from waterways connected to the central region of the Lagoon of Venice; Invasion likely via oysters import from the Pacific (d'Hondt and Occhipinti Ambrogi 1985, cited in Porter et al 2015; NEMESIS 2016; Cook et al 2013a; Bandelj et al 2009; Sconfietti and Marino 1989; Sconfietti et al 2003) *Introduced [Spain] 1996: Observed for the first time in Galicia, northwest of Spain (De Blauwe and Faasse 2001) *Introduced [Portugal] 2004: Sampled Ria de Aveiro and found first observations, introduced from Venice, Italy (Marchini et al 2007) *Introduced [Tunisia] 2005: Gulf of Tunis collected (Ben Souissi et al 2006, cited in Lodola et al 2012 (possibly Cook et al. 2013a?)) *Invasive [Mediterranean Sea] Reported in the central region of the Mediterranean, on the northern coast of Tunisia, north-west Italy and the east coast of Sardinia. Along the eastern coast of the Atlantic, T. inopinata has been reported from Cadiz, in southern Spain to the northeast coast of Scotland (Ben Souissi et al. 2006, cited in Cook et al 2013a; Lodola et al 2012; Rosso 2003) *Invasive Introduced to the Mediterranean Sea and the European Atlantic coast (Galil and Occhipinti-Ambrogi 2009) *Introduced Invaded the Mediterranean (Occhipinti Ambrogi and d'Hondt 1994, cited in De Blauwe and Faasse 2001; Rilov and Galil 2009) *Introduced [US] 2010, reported in Woods Hole, Massachusetts, USA; Spread to New England locales (Johnson et al 2012; Porter et al 2015; NEMESIS 2016; Johnson and Woollacott 2015; Galil and Occhipinti-Ambrogi 2009; Bishop et al 2015b) *Introduced Temperate Australasia New Zealand (Occhipinti Ambrogi and d'Hondt 1994, cited in Porter et al 2015; Bishop et al 2015b; Connell 2001; Dyrynda et al 2000; Piola and Johnston 2009) *Introduced Invaded New Zealand (Occhipinti Ambrogi and d'Hondt 1994, cited in De Blauwe and Faasse 2001; Rilov and Galil 2009) *Introduced New Zealand (NEMESIS 2016) *Cryptogenic Uncertain realm Australia (Occhipinti Ambrogi and d'Hondt 1994, cited in Porter et al 2015; Bishop et al 2015b; Connell 2001; Dyrynda et al 2000; Piola and Johnston 2009) *Introduced Invaded Australia (Occhipinti Ambrogi and d'Hondt 1994, cited in De Blauwe and Faasse 2001; Rilov and Galil 2009) *Introduced Australia (NEMESIS 2016) *Cryptogenic

Status Date Non-native:

[US] First observations in 2010 at Woods Hole, Massachusetts (Johnson et al 2012; Porter et al 2015; NEMESIS 2016; Johnson and Woollacott 2015) [Norway] Found at ports of Floro and Kristiansund in 2014 (Porter et al 2015) [UK] 1998-2006 (Dyrynda et al 2000; Kelso and Wyse Jackson 2012; Porter et al 2015; Dyrynda 2005; Bishop et al 2015a; NEMESIS 2016; CABI 2016; Cook et al 2013a; Nall et al 2015) [Germany] New observations in Wadden Sea in 2009 (Buschbaum et al 2012) [Italy] New to science when discovered in Venice; Collected in May 1982 (Marchini et al 2015; d'Hondt and Occhipinti-Ambrogi 1985, cited in Porter et al 2015) [Spain] 1996: Observed for the first time in Galicia, northwest of Spain (De Blauwe and Faasse 2001) [Portugal] Sampled in 2004 (Marchini et al 2007) [Tunisia] Collection in Gulf of Tunis in 2005 (Ben Souissi et al 2006, cited in Lodola et al 2012) [Japan] 2006 in Akkeshi Bay. (Grischenko et al. 2007) [Japan] 2006-2010 in Osaka Bay. (Association for the Research of Littoral Organisms in Osaka Bay 2012) [Italy] 2010 at La Spezia Harbor. (Lodola et al. 2012, cited in Cook et al. 2013b) [United Kingdom] 2009 at Brixham, Burry Port and othe regions in England. (Oakley & Griffith pers. com., cited in Cook et al. 2013b) [United Kingdom] 2011 at Holyhead Marina, Milford Haven Area and Nayland Marine in Wales. (Holt pers. com., cited in Cook et al. 2013b) [Belgium] 2001 at Bredene, De Haan, and Zeebrugge. (De Blauwe 2002, cited in Cook et al. 2013b) [France] Pre-2012 at Arcachon Bay. (Goulletquer pers. com., cited in Cook et al. 2013b) [The Netherland] 2011 at Oesterdam. (De Blauwe pers. com., cited in Cook et al. 2013b) [Spain] Pre-2012 at Costa de Lugo and Xove. (Reverter-Gil pers. com., cited in Cook et al. 2013b) [Portugal] 2004 at Ria de Aveiro. (Marchini et al. 2007, cited in Cook et al. 2013b) [Tunisia] 2005 at Gulf of Tunis, Tunis lagoon and Harbor of Radès. (Zenetos pers. com., cited in Cooks et al. 2013b) [Germany] 2009 at Hörnum, List and Wilhelmshaven. (Buschbaum et al. 2011, cited in Cook et al. 2013b) [Madeira] 2011 at Bay of Funchal. (Clode & Jones pers. com., cited in Cook et al. 2013b)

Vectors and Spread

Initial Vector:

Aquaculture and Fisheries, Hull fouling (commercial)

Second Vector:

Hull fouling (recreational, commercial), Natural dispersal

Vector Details:

[US] Shipping traffic most likely vector that introduced it to the US (Johnson et al 2012; Johnson and Woollacott 2015) 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 (CABI 2016; Galil and Occhipinti-Ambrogi 2009; Marchini et al 2007; Savini et al 2006b) Found almost exclusively in marinas, harbours, or enclosed bays associated with aquaculture activities (Cook et al 2013a) In Europe, often found near sites of oyster culture, though secondary transport by recreational and fishing boats is likely (Galil and Occhipinti Ambrogi 2006, cited in NEMESIS 2016) Introduction from Venice to the Atlantic coasts of Spain was likely the movement of clams between the two locations (Fernandez-Pulpeiro et al 2001, cited in CABI 2016) 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 probable ways of expansion, accounting for the rapid colonization of the European Atlantic coasts from Portugal to the Netherlands (CABI 2016; Ashton et al 2006; Marchini et al 2007) The introduction of T. i. to italy in 1982 (Occhipinti-Ambrogi 2000) and France (Goulletquer pers. com., cited in Cook et al. 2013b) has been linked with the importation of the Pacific oyster, Crassostrea gigas for culture purposes. (Cook et al. 2013b) T. i. has been transported to the lagoon of Venice together with some fisheries products (e. g. oysters or clam spat) whose cultivation is widespread there. (Occhipinti -Ambrogi & D'Hondt 1994) Cook et al. (2013b) stated that De blauwe and Faasse (2001) suggested that the spread of T. i. along the coast of The Netherlands and Belguim was from fouled hulls of smaller boats (fishing boats, sport yachts or smaller trade vessels).

Spread Rate:

[US] On the east coast, population at Woods Hole Harbor, MA from 2010 had expanded to Boston Harbor by 2011. 2013 survey found its range extended southward to mouth of Narragansett Bay, Rhode Island and northward to Hampton River, New Hampshire (NEMESIS 2016) [UK] First record in 1998 from Poole Harbour, Dorset in 1998. In 1999, the species was found to extend only from Swanage to Chichester Harbour (Dyrynda et al. 2000), a distance of c. 75 km, but by 2004 it had spread westwards on the south coast as far as Falmouth (Arenas et al. 2006). Bishop et al (2015) discusses T. inopinata around the coast of England from Fleetwood in the North West to Scarborough on the east coast, but these do not represent the species’ northern limits in UK, as by 2012 it occurred on both the west and east coasts of Scotland following an initial Scottish sighting in 2006 (Cook et al. 2013; Bishop et al 2015b) [Norway] Significant northerly extension to the range in North-west Europe based on records in the Orkney Isles, Scotland (Cook et al 2013a and Nall et al 2015, cited in Porter et al 2015) [Italy] Estimated spread rate through the Lagoon of Venice of at least 6km per year (Occhipinti-Ambrogi 1990, cited in Savini et al 2006a) [Europe] First discovered in 1982 in lagoon of Venice. In 1998, discovered in the Marano Lagoon, 70km to the east (Dyrynda et al 2000, cited in NEMESIS 2016) [Europe] Dispersal throughout Europe has been rapid and represents a range expansion of ~ 5,500 km (by sea) north to the North-east coast of Scotland from where it was first reported in north-east Italy in 1982. This equates to a dispersal rate of approximately 190 km/yr (Bishop et al 2015b; Cook et al 2013a) [Europe] Rapid expansion along European Atlantic coasts (CABI 2016) Approximately 190km/year in Europe. (Cook et al. 2013b) RELATED: Spread northwards from more southerly locations in mainland Europe (De Blauwe and Faasse 2001) or by hopping across the North Sea from Scotland (Cook et al 2013a) - not mutually exclusive hypotheses (Porter et al 2015)

Date First Observed in Japan:

Mutsu Bay: 1926-1927. (Okada 1928, 1929) RELATED: [Menipea occidentalis] Yanagi & Okada (1918) stated the occurrence of Menipea occidentalis first from Japanese waters, but they don't state the date when the collection was made.

Date First Observed on West coast North America:

In 2010, species was found in Woods Hole Harbor, MA (Johnson et al 2012, cited in NEMESIS 2016)

Impacts

Impact in Japan:

NF

Global Impact:

[US] Reaches reproductive maturity earlier than other bryozoan species in Atlantic US (Johnson and Woollacott 2015) [US] On the east coast, displacing native arborescent bryozoans (NEMESIS 2016) [Italy] Invasion is a major event affecting bryozoan assemblage of the Lagoon of Venice during last twenty years; Has replaced the native Bugula spp. as the dominant bryozoan in Venice Lagoon (d'Hondt and Occhipinti-Ambrogi 1985, cited in CABI 2016; Corriero et al 2007; Occhipinti-Ambrogi 2000; Occhipinti-Ambrogi and Savini 2003; Streftaris and Zenetos 2006) [Europe] The lack of die back in the winter months and almost year-round reproduction in certain regions gives this species a distinct advantage over the native bryozoans, such as Bugula spp., which typically die back in winter and then re-establish themselves in a yearly cycle; loss of biodiversity by replacing native bryozoan species (Reverter-Gil and Fernandez-Pulperio 2001, Marchini et al. 2007, Glass 2009 and Ryland 1960, cited in Cook et al 2013a; Cook et al 2013b; NEMESIS 2016) Ability to grow on other bryozoan species while other species may not be able to settle on top of it (Johnson et al 2012) Probably contributes to fouling of recreational boats, docks, ropes, etc. But its economic impacts cannot be easily separated from those of the rest of the fouling community (NEMESIS 2016) Fouling impacts in the community (Kerckhof et al 2007) The loss of biodiversity, as a consequence of T. i., could be quantified. (Occhipinti-Ambrogi & Savini 2003, cited in Cook et al. 2013b)

Tolerences

Native Temperature Regime:

Cool temperate, Mild temperate, Warm temperate, Subtropical

Native Temperature Range:

San Francisco: max 20.0ºC in summer and min 11.0ºC in winter. (Clark et al. 2003) San Diego: max 22.0ºC in summer and min 13.0ºC in winter. (Clark et al. 2003) Cool temperate, Mild temperate, Warm temperate, Subtropical (M. Otani, pers. comm.)

Non-native Temperature Regime:

Cool temperate, Mild temperate, Warm temperate, Subtropical

Non-native Temperature Range:

VARIABLITY Venezia: max 27.0ºC in summer and min 9.0ºC in winter. (Clark et al. 2003) Minimum water temperatures in the lagoon of Venice are 2-3ºC. (Occhipinti-Ambrogi 1991, cited in DeBlauwe & Faasse 2001) 8.6˚C - 10.4 ˚C (Markert et al 2015) Temperate, optimum 20-25°C; 3-34°C tolerated (CABI 2016) Temperate and subtropical latitudes (Cook et al 2013a) Cool temperate, Mild temperate, Warm temperate (M. Otani, pers. comm.)

Native Salinity Regime:

Polyhaline, Euhaline

Native Salinity Range:

San Francisco: max 32.0psu in dry period and min 10.0psu in wet period. (Clark et al. 2003) San Diego: max 37.0psu in dry period and min 33.5psu in wet period. (Clark et al. 2003)

Non-native Salinity Regime:

Polyhaline, Euhaline

Temperature Regime Survival:

Cool temperate, Mild temperate, Warm temperate, Subtropical, See details

Temperature Range Survival:

2-34.5°C (Galil and Occhipinti-Ambrogi 2006 and Johnson et al 2012, cited in NEMESIS 2016; Galil and Occhipinti-Ambrogi 2009) Freezing to near-freezing to 28 °C (Johnson et al 2012 and Marchini et al 2007, cited in Cook et al 2013a) 11.597 ºC (OBIS 2016) T. i. will tolerate from freezin or near-freezing to 28ºC. (Johnson et al. 2012 and Marchini et al. 2007, cited in Cook et al. 2013b) Cool temperate, Mild temperate, Warm temperate, Subtropical (M. Otani, pers. comm.) RELATED: [Tricellaria spp.] -0.262 - 16.802 ºC (OBIS 2016)

Temperature Regime Reproduction:

Cool temperate, Mild temperate, Warm temperate

Temperature Range Reproduction:

Able to brood embryos over a wide range of temperatures from 8.6–16.7°C although the optimal temperature range was 12.6–15.5°C for colonies sampled in Plymouth, south coast of the UK (Glass 2009, cited in Cook et al 2013a). T. i. is able to brood embryos over a wide range of temperature from 8.6-16.7ºC for colonies sampled in Plymouth. (Grass 2009, cited in Cook et al. 2013b) Cool temperate, Mild temperate, Warm temperate (M. Otani, pers. comm.)

Salinity Regime Survival:

Polyhaline, Euhaline

Salinity Range Survival:

20-38 PSU (Galil and Occhipinti-Ambrogi 2006, cited in NEMESIS 2016; Galil and Occhipinti-Ambrogi 2009; Cook et al 2013a) 34.849 PPS (OBIS 2016) T. i. will tolerate sakinities from 20-38psu with the optimal range fo 20-35psu. (Breton & D'Hondt 2005; Corriero et al. 2012; Marchini et al. 2007, cited in Cook 2013, Dyrynda et al. 2000) T. i. did not occur in the innermost section of the canals where salinity drops frequently <20ppt, nor outside the lagoon in the Adriatic Sea. (Occhipinti-Ambrogi 2000) RELATED: [Tricellaria spp.] 27.473 - 35.498 PPS (OBIS 2016)

Salintiy Regime Reproduction:

Polyhaline, Euhaline

Salinity Range Reproduction:

Polyhaline, Euhaline (M. Otani, pers. comm.)

Depth Regime:

Upper intertidal, Mid intertidal, Lower intertidal, Shallow subtidal

Depth Range:

5 - 13m depth (Markert et al 2015) Species first confined to the intertidal level, attached to the byssus threads of Mytilus galloprovincialis (CABI 2016) Became most abundant species of the hard-bottom community at the intertidal and shallow subtidal level (Occhipinti-Ambrogi 2000, cited in CABI 2016) Infra-littoral to the shallow sub-tidal zone (Dyrynda et al 2000, cited in Cook et al 2013a) T. i. has been found within the infralittoral fringe (lowest shore/shollow subtidal zone). (Dyrynda et al. 2000) RELATED: [T. occidentalis] Found commonly between tide marks. (Mawatari 1951) [T. occidentalis] Found at 20-30m deep. (Inaba 1988)

Non-native Salinity Range:

Native Abundance:

See details

Reproduction

Fertilization Mode:

Internal

Reproduction Mode:

Hermaphrodite/monoecious

Spawning Type:

NA

Development Mode:

Lecithotrophic planktonic larva (non-feeding)

Asexual Reproduction:

Budding/fragmentation (Splitting into unequal parts. Buds may form on the body of the “parent”)

Reproduction Details:

Zooids are hermaphroditic and produce large yolky eggs, which hatch into lecithotrophic larvae; 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, cited in NEMESIS 2016) Hermaphroditic; broods its lecithotrophic eggs in a special external chamber (ovicell). Larvae are planktonic, non-feeding and last only a few hours. Metamorphosis occurs within minutes of settlement to substrate. Mature colonies with ovicells and embryos are found over much of the year, allowing for a continuous recruiting (Galil and Occhipinti-Ambrogi 2009) Species reproduction is both asexual, through zooidal budding, or sexual, with sperm and eggs developing in each zooid and originating, after fertilization, as embryos which are brooded in the ovicells (CABI 2016; Cook et al 2013a) Short free swimming larval stage, which settles within two hours (Johnson et al. 2012, cited in Cook et al 2013a; Occhipinti-Ambrogi and d'Hondt 1994) RELATED: [Order: Cheilostomata] Free spawning species produce the characteristic triangular cyphonautes larva. These larvae are long-lived and planktotrophic. The larval body is enclosed in a membranous shell; the size can be up to little over 1 mm. Cyphonautes larvae are not keyed out - if possible at all. (van Couwelaar 2003) [Gymnolaemates] Internal fertilization, whether intracoelomic or intraovarian, is obligatory (Temkin 1994 and 1996, cited in Ostrovsky 2013) [Gymnolaemates] Differ from most organisms in that sperm-egg fusion does not stimulate egg activation. Egg activation may not occur until "spawned" outside of maternal zooid (Temkin 1991) [Bryozoans] While sperm is spawned through pores in lophophore tentacles, eggs are usually harbored inside the body wall, and are internally fertilized by sperm, coming in on lophophore feeding currents (Brusca and Brusca 2003, cited in Rouse 2011; Kozloff 1990, cited in Rouse 2011) [Bryozoans] Colonial hermaphrodites, with testes (spermatogenic tissue) and ovaries developing either within the same zooid (zooidal hermaphroditism) or in different zooids within the same colony (zooidal gonochorism) (Ostrovsky 2013) Members of the phylum Bryozoa are hermaphroditic. Both fertilization and egg brooding may either be internal or external (Ruppert et al. 2004) [Bryozoa] All bryozoan colonies are hermaphroditic. Autozooids may be dioecious; or monoecious, and protandrous or protogynous. (Hayward & Ryland 1998) [Bryozoa] Reproduces asexually by budding. (Mawatari 1976)

Adult Mobility:

Sessile

Adult Mobility Details:

Encrusting, bush-like, calcified bryozoan (NEMESIS 2016) RELATED: [Bryozoa] The abundance and taxonomic diversity of benthic bryozoan faunas are directly related to substratum. (Hayward & Ryland 1998) [Bryozoa] Colonies are sessile (Hayami 1975) [Bryozoa] Phylum of sessile, colonial suspension feeders found throughout the world in both marine and freshwater environments. (Tilbrook 2012)

Maturity Size:

Zooid: 450-650μm (NEMESIS 2016) Colony: up to 6-8cm in length (CABI 2016) Colony maximum height: 40mm (Johnson et al 2012, cited in NEMESIS 2016; Cook et al 2013a) Large number of colonies come to sexial activity within six weeks in spring and somewhat earlier in summer with the average length of 10mm at spring generation. (Mawatari 1951)

Maturity Age:

Large number of colonies come to sexial activity within six weeks in spring and somewhat earlier in summer. (Mawatari 1951)

Reproduction Lifespan:

VARIABILITY Embryos were found year round. (Occhipinti-Ambrogi 2000) Reproductively active colonies have been found all year round in parts of Europe. (Reverter-Gil & Fernandez-Pulperio 2001, cited in Cook et al. 2013) RELATED: [Tricellaria occidentalis] According to the observations of the test plates monthly submerged at Yokohama harbour, the first appearance of young colonies seemed to be early in May, and the last settlement of new colonies late in November. (Mawatari 1951) [Tricellaria occidentalis] Breeding season at Asamushi, Mutsu Bay is from January to December. (Tsuchiya & Osanai 1978)

Longevity:

RELATED: Annual cycle of colony die-back and re-growth. During this cycle, the arborescent portion of the colony will die off, most likely due to deterioration in environmental conditions. When conditions improve however, colonies will grow back, presumably stemming from the root-like projections that remained attached to the substrate (Johnson et al 2012) [T. occidentalis] The falling off of the species that occurs only in winter may imply the longevity of the species is 6-7 months. (Mawatari 1976)

Broods per Year:

NF

Reproduction Cues:

RELATED: [Bryozoans] Experiments often used light as a cue to collect embryos/larvae (Woollacott and Zimmer 1977) [Tricellaria occidentalis] Accelerating effect of light stimulus to the liberation of larvae is obvously noticed in T. o. as wa seen in Bugula neritina. (Mawatari 1951) [Bryozoa] In coastal species light is an important stimulus to larval release, and many cheilostomates shed larvae during the first few hours of daylight. (Hayward & Ryland 1998) [Bryozoa] In various degrees of intensity according to the species temperature also stimulates sexual reproduction. (Winston 1977)

Reproduction Time:

VARIABILITY Embryos were found year round. (Occhipinti-Ambrogi 2000) Reproductively active colonies have been found all year round in parts of Europe. (Reverter-Gil & Fernandez-Pulperio 2001, cited in Cook et al. 2013b) Mature colonies with ovicells and embryos are found over much of the year, allowing a continuous recruiting population (Occhipinti-Ambrogi and d'Hondt 1994, cited in CABI 2016; Cook et al 2013a) [US] Reaches reproductive maturity by late-May or early-June; Began to decrease in mid-December (Johnson and Woollacott 2015; Johnson et al 2012) RELATED: [Tricellaria occidentalis] According to the observations of the test plates monthly submerged at Yokohama harbour, the first appearance of young colonies seemed to be early in May, and the last settlement of new colonies late in November. (Mawatari 1951) [Tricellaria occidentalis] Breeding season at Asamushi, Mutsu Bay is from January to December. (Tsuchiya & Osanai 1978)

Fecundity:

High reproductive rate with high numbers in ovicells by each brooding zooid (Occhipinti-Ambrogi and d'Hondt 1994)

Egg Size:

RELATED: Ovicell: 150-160μm in diameter (NEMESIS 2016) [Gymnolaemata] About 200µm (Woollacott and Zimmer 1977)

Egg Duration:

NF

Early Life Growth Rate:

Planktonic for short periods (a few hours); larvae settle on substrate and metamorphose into the first zooid of a colony (NEMESIS 2016) Free swimming larval stage, which settles within two hours (Johnson et al. 2012, cited in Cook et al 2013a) Relatively rapid metamorphosis (<32 hours) (Occhipinti Ambrogi and d'Hondt 1994 and Johnson et al. 2012, cited in Cook et al 2013a) 90% of sampled individuals initiated metamorphosis within 2hr and 90% had completed metamorphosis within 32hr (Johnson et al 2012) RELATED: [Gymnolaemata] Two phases of larvae metamorphosis: first stage about 20mins; second stage 1-6 days (Woollacott and Zimmer 1977)

Adult Growth Rate:

Relatively rapid metamorphosis (<32 hours) (Occhipinti Ambrogi and d'Hondt 1994 and Johnson et al. 2012, cited in Cook et al 2013a) According to Mawatari (1951), the growth of the colony is as follows. Within the first three weeks: colony reaches 5 or 6 mm. At the end of the sixth week: average length of colonies of spring generation is 10 mm. In two months: the largest colony of summer generation reaches a height of 20 mm. At the end of the third month: the colonies may reach 25 mm or more in height , and 30 mm or more in diameter.

Population Growth Rate:

NF

Population Variablity:

After the first record of T. i. at Venice lagoon in 1982 (Occhipinti-Ambrogi 2000), its population reached its maximum abundance and geographical spread c. 1988-1989 (Occhipinti-Ambrogi 1991, cited in Dyrynda et al. 2000), but declined in both frequency and abundance from 1993 onwards (Occhipinti-Ambrogi 2000), and population in the Wadden Sea, northern Germany, have been absent since 2009-2010. (D. Lackschewitz pers. comm., cited in Cook et al. 2013b)

Habitat

Ecosystem:

SAV, Rocky intertidal, Rocky subtidal, Mussel reef, Oyster reef, Worm reef, Macroalgal beds, Flotsam, Floating plants or macroalgae, Fouling,

Habitat Type:

Epibenthic, Epiphytic; Epizoic

Substrate:

Rock, Gravel, Biogenic, Artificial Substrate

Exposure:

Semi-exposed, Protected, Very protected

Habitat Expansion:

NF

Habitat Details:

Abundant on natural surfaces; artificial surfaces such as ropes and buoys; and on other sessile fauna (De Blauwe and Faasse 2001, cited in Porter et al 2015; Arenas et al 2006) Grows attached to wood pilings, floats, rocks, boulders, seaweeds, seagrasses, oysters, mussels, and other hard substrates. Also been found attached to other exotic fouling species; strong preference for sheltered locations (i.e. lagoons, estuaries, harbours and marinas), and tolerates some variation in salinity (NEMESIS 2016; Galil and Occhipinti-Ambrogi 2009; Dyrynda et al 2000) Seldom, if ever, found in exposed sites on the open coast (CABI 2016) Settles on a wide range of anthropogenic and natural substrata. Dispersal with natural rafting such as current-dispersed algal fragments or pumice; or as fouling organisms on vessels and marine flotsam (Galil and Occhipinti-Ambrogi 2009) Littoral rock and other hard substrata, marine infralittoral; Marine benthic intertidal and subtidal, preferably in marinas, harbours and brackish water bodies (Galil and Occhipinti-Ambrogi 2009) Will attach to a wide range of biotic and abiotic substrates, including artificial and natural structures such as vessel hulls, floating pontoons, navigation buoys, and infra-littoral boulders. It readily grows on macroalgae, such as Chondrus crispus and the non-native Sargassum muticum and on other invertebrates such as the non-native Styela clava, Mytilus galloprovincialis, sponges, hydroids and even other bryozoans (Occhipinti Ambrogi and d'Hondt 1994, De Blauwe and Faasse 2001, Fernandez-Pulperio et al. 2001 and Dyrynda et al. 2000, cited in Cook et al 2013a; Savini et al 2006a) Species has been found in association with macrophytes (Sargassum muticum, Undaria pinnatifida and Codium fragile), mussels and oysters (Mytilus galloprovincialis, Mytilus edulis, and Crassostrea gigas), sponges, ascidians (Styela clava, Ascidiella aspersa, and Styela plicata), polychaete worms (Ficopomatus enigmaticus and Hydroides dianthus), other bryozoans (Bugula neritina) and many other organisms constituting hard bottom communities in lagoon and harbours (CABI 2016; Mineur et al 2014) It readily grows on macroalgae, such as Chondrus crispus and the non-native Sargassum muticum and on other invertebrates such as the non-native Styela clava and Mytilus galloprovincialis (Cook et al 2013a) T. i. was found growing on ships' hulls, floats, buoys, ropes, stones, the algae Sargassum muticum, Codium fragile and Ulva sp., on an unidentified campanulariid hydrozoan, on mussels, Mytilus edulis and the ascidians Styela clava and Ascidiella aspersa at North Sea coast. In Hendaye it was found on the bryozoan Bugula neritina and the polychaete worm Ficopomatus enigmaticus. Ancestrulas settle on colonies of their own species as well. (De Blauwe & Faasse 2001) T. i. colonized primary (abiotic) substrata, most notably floats associated with berthing pontoons, but also infralittoral boulders. Secondary (biotic) substrata colonized within the port area, marinas and natural shores include macro-algae such as Chondrus crispus and Sargassum muticum and sessile invertebrate such as Styela clava. (Dyrynda et al. 2000) [As Tricellaria occidentalis (Synonymized taxon)] Known from various sorts of seaweeds and as one of the most common fouling organisms on setting nets and ships' bottoms in Japan. (Mawatari 1951) Known from gravel, rock, ship, and etc. (Inaba 1988) Semi-exposed (M. Otani, pers. comm.) [As Menipea occidentalis (Synonymized taxon)] Attached to sea-weeds. (Okada 1929)

Trophic Level:

Suspension feeder

Trophic Details:

Feeds on phytoplankton, suspension feeder (NEMESIS 2016) RELATED: [Bryozoans] Suspension feeder...filter phytoplankton less than 0.045mm in size from the water column. (Hill 2001) [Bryozoa] Many phytoplankton species are cleary unsuitable as food for bryozoans. (Hayward & Ryland 1998) [Cheilostomata] Main food is diatom, protozoans and etc. and unappropriate sized particles are ejected (Mawatari 1976)

Forage Mode:

Generalist

Forage Details:

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 (NEMESIS 2016) RELATED: [Bryozoans] Suspension feeder...filter phytoplankton less than 0.045mm in size from the water column. (Hill 2001) [Bryozoa] Many phytoplankton species are cleary unsuitable as food for bryozoans. (Hayward & Ryland 1998) [Cheilostomata] Main food is diatom, protozoans and etc. and unappropriate sized particles are ejected (Mawatari 1976)

Natural Control:

COMPETITION: Other sessile fauna although it generally outcompetes local species (CABI 2016) RELATED: PREDATION [Predation] [Bryozoa] Browsers and grazers, including sea urchins, fish, crabs and some prosobranchs, are known to include bryozoans in their diet. (Hayward & Ryland 1998) [Predation] [Bryozoa] Bryozoans are also the prey of very many small, selective predators, some of which may be adapted to a very narrow spectrum of prey species. Among them opisthobranch predators of bryozoans are well known. (Hayward & Ryland 1998) [Predation] [Bryozoa] Other than opisthobranchs as a predator, amphipods, isopods, mites and pycnogonids have all been recorded preying on bryozoan colonies. (Hayward & Ryland 1998) EPIBIONTS [Epibionts] [Cheilostomata] It is frequently observed in Japan that several species of hydroids flourish on Cheilostomata cause damages to them. (Mawatari 1976)

Associated Species:

Ancestrulas of T. i. settle on colonies of their own species as well. (De Blauwe & Faasse 2001)

References and Notes

References:

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Modelling spatial distribution of hard bottom benthic communities and their functional response to environmental parameters. Ecological Modelling, 220(21), 2838-2850. Doi: 10.1016/j.ecolmodel.2009.04.024 Bishop, J. D. D., Wood, C. A., Lévêque, L., Yunnie, A. L. E., & Viard, F. (2015a). Repeated rapid assessment surveys reveal contrasting trends in occupancy of marinas by non-indigenous species on opposite sides of the western English Channel. Marine Pollution Bulletin, 95(2), 699-706. Doi: 10.1016/j.marpolbul.2014.11.043 Bishop, J. D. D., Wood, C. A., Yunnie, A. L. E., & Griffiths, C. A. (2015b). Unheralded arrivals: non-native sessile invertebrates in marinas on the English coast. Aquatic Invasions, 10(3), 249-264. Doi: 10.3391/ai.2015.10.3.01. Buschbaum, C., Lackschewitz, D., & Reise, K. (2012). Nonnative macrobenthos in the Wadden Sea ecosystem. Ocean & Coastal Management, 68, 89-101. Doi: 10.1016/j.ocecoaman.2011.12.011 CABI. (2016). Invasive Species Compendium. 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Literature:

Extensive scientific information; peer-reviewed information; data specific to the location; supported by long-term datasets (10 years or more)

Notes:

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 (CABI 2016) May be a complex with T. occidentaris (M. Otani, pers. comm.)