Tricellaria inopinata

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

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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).

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Description

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.

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Taxonomy

Ecology

General:

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).

Food:

Phytoplankton

Trophic Status:

Suspension Feeder

SusFed

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Habitats

General Habitat	Marinas & Docks	None
General Habitat	Rocky	None
General Habitat	Vessel Hull	None
General Habitat	Oyster Reef	None
Salinity Range	Polyhaline	18-30 PSU
Salinity Range	Euhaline	30-40 PSU
Vertical Habitat	Epibenthic	None

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Life History

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	2	Field (Galil and Occhipinti-Ambrogi 2006; Johnson et al. 2012)
Maximum Temperature (ºC)	34.5	Field (Galil and Occhipinti-Ambrogi 2006)
Minimum Salinity (‰)	20	Field (Galil and Occhipinti-Ambrogi 2006)
Maximum Salinity (‰)	35	Field (Galil and Occhipinti-Ambrogi 2006)
Maximum Duration	0.2	Settles after a few hours (Occhipinti Ambrogi and d'Hondt 1994).
Maximum Height (mm)	40	Colony 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).

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Regional Impacts

NA-ET3	Cape Cod to Cape Hatteras		Ecological Impact	Competition
Preliminary observations in Eel Pond (Woods Hole, MA) indicate that T. inopinata has negatively impacted other Eel Pond arborescent bryozoans (ICES 2012, in prep.).
M010	Buzzards Bay		Ecological Impact	Competition
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-VII	None		Ecological Impact	Competition
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-II	None		Economic Impact	Shipping/Boating
Fouling impacts have been listed for the British Isles (Minchin et al. 2013)
NEA-V	None		Ecological Impact	Habitat 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)
MA	Massachusetts		Ecological Impact	Competition
Preliminary observations in Eel Pond (Woods Hole, MA) indicate that T. inopinata has negatively impacted other Eel Pond arborescent bryozoans (ICES 2012, in prep.).

References

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>



Leclerc, Jean-Charles; Viard, Fredérique (2017) Habitat formation prevails over predation in influencing fouling communities, Ecology and Evolution 7: 477-492

Abbe, George R. (1987) Epifauna, In: Heck, Kenneth L.(Eds.) Ecological studies in the middle reach of Chesapeake Bay- Calvert Cliffs. , Berlin. Pp. 82-91

Amor, Kounofi-Ben; Rifi, M.; Ghanem, R.; Draief, I.; Zouali, J.; Souissi, J. Ben (2016) Update of alien fauna and new records of Tunisian marine fauna, Mediterranean Marine Science 17(1): 124-143

Arenas, F. and 13 authors. (2006) Alien species and other notable records from a rapid assessment survey of marinas on the south coast of England., Journal of the Marine Biological Association of the United Kingdom 86: 329-1337

Arenas, Francisco; Sanchez, Inigo; Hawkins, Stephen J.; Jenkins, Stuart R. (2006) The invasibility of marine algal assemblages: role of functional diversity and identity., Ecology 87(11): 2851-2861

Armstrong, John D.; Bean, Colin W.; Wells, Alan (2018) The Scottish invasion of pink salmon in 2017, Journal of Fish Biology 93: 8-11
DOI: 10.1111/jfb.13680

Barnes, Robert D. (1983) Invertebrate Zoology, Saunders, Philadelphia. Pp. 883

Bishop, John D. D.; Wood, Christine A.; Lévêque, Laurent; Yunnie, Anna L. E.; Viard, Frédérique (2015b) 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: 699-706

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

Breton, Gérard; D'Hondt, Jean-Loup (2006) Tricellaria inopinata d'Hondt et Occhipinti Ambrogi, 1985 (Bryozoa: Cheilostomatida) dans le Port du Havre (manche orientale)., Bulletin Trimestrial de la Societe geologique de Normandie 9`(2): 67-72

Buschbaum, Christian; Lackschewitz, Dagmar; Reise, Karsten (2012) Nonnative macrobenthos in the Wadden Sea ecosystem, Journal of Ocean Management 68: 89-101

Buschbaum, Christian; Lackschewitz, Dagmar; Reise, Karsten (2012) Nonnative macrobenthos in the Wadden Sea ecosystem, Journal of Ocean Management 68: <missing location>

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
https://doi.org/10.1371/journal.pone.0218801

Chae , Hyun Sook Yang, Ho Jin; Min , Bum Sik; Noh , Geon Woo; Lee; Hee, Dong Lee; Dong; Seo, Jeun (2022) Three newly recorded species of Korean fouling bryozoans, Journal of Species Research 11: 180-187

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>

Cook, Elizabeth J. and 5 authors (2013) Distribution of the invasive bryozoan Tricellaria inopinata in Scotland and a review of its European expansion, Aquatic Invasions 8: In press

Cornelio, Michele; Manzoni, Alberto (1999) Caratterizzazione stagionale degli insediamenti di organismi macrobentonici su substrati sperimentali nel bacino centrale della laguna di Venezia., Bollettino del Museo Civico di Storia Naturale di Venezia 49: 135-144

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

D'Hondt, Jean-Loup; Occhipinti Ambrogi, Anna (1985) Tricellaria inopinata n. sp., un nouveau bryozoiare Cheilostome de la faune méditerranée, Marine Ecology 6(1): 35-36

DAISIE (Delivering Alien Invasive Species Inventories to Europe) (2009) Handbook of alien species in Europe, Springer, Dordrecht, Netherlands. Pp. 269-374

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

Dyrynda, P. E. J.; Fairall, V. R.; A. Occhipinti Ambrogi; D’Hondt, J.-L. (2000) The distribution, origins 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

Galil, B. S.; Occhipinti-Ambrogi, A. 2006 <i>Tricellaria inopinata</i>- DAISIE Factsheet. <missing URL>



Gueron, Rodrigo; Almeida, Alexandre Oliveira; Aguilar, Robert; Ogburn, Matthew B.; Prakash, Sanjeevi ; Baeza, J. Antonio (2022) Delimiting species within the Lysmata vittata (Stimpson, 1860) (Decapoda: Lysmatidae) species complex in a world full of invaders, Zootaxa 5150(2): 189–216
https://doi.org/10.11646/zootaxa.5150.2.2

ICES Advisory Committee on the Marine Environment (2011) <missing title>, International Council for the Exploration of the Seas, Copenhagen. Pp. 1-180

ICES Advisory Committee on the Marine Environment (2012) <missing title>, International Council for the Exploration of the Seas, Copenhagen. Pp. <missing location>

Inglis, Graeme and 6 authors (2006a) Port of Lyttelton: Baseline survey for non-indigenous marine species, Biosecurity New Zealand Technical Paper 2005/01: 1-64

Inglis, Graeme and 5 authors (2006b) Port of Taranaki: Baseline survey for non-indigenous marine species, Biosecurity New Zealand Technical Paper 2005/04: 1-47

Inglis, Graeme and 6 authors (2006e) Whangarei Harbour (Whangarei Port and Marsden Point: Baseline survey for non-indigenous species, Biosecurity New Zealand Technical Paper 2005(16): 1-52

Johnson, Collin H.; Winston, Judith E.; Woollacott, Robert M. (2012) Western Atlantic introduction and persistence of the marine bryozoan Tricellaria inopinata, Aquatic Invasions 7(3): 295–-03

Johnson, Collin H.; Woollacott, Robert M. (2015) Analyses with newly developed microsatellite markers elucidate the spread dynamics of Tricellaria inopinata d’Hondt and Occhipinti-Ambrogi, 1985 - a recently established bryozoan along the New England seashore, Aquatic Invasions 10(2): 135-145

Kelso, Antoinette; Wyse Jackson, Patrick N. (2012) Invasive bryozoans in Ireland: first record of Watersipora subtorquata (d'Orbigny, 1852) and an extension of the range of Tricellaria inopinata d’Hondt and Occhipinti Ambrogi, 1985, Bioinvasion Records 1(3): 209-214

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

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

Lodola, Alice; Savini, Dario; Anna Occhipinti-Ambrogi (2012) First record of Tricellaria inopinata (Bryozoa: Candidae) in the harbours of La Spezia and Olbia, Western Mediterranean Sea (Italy), Marine Biodiversity Records 5: e41

Lord, Joshua P.; Calini, Jeremy M.; Whitlatch, Robert B. (2015) Influence of seawater temperature and shipping on the spread and establishment of marine fouling species, Marine Biology 162: 2481-2492

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

McCuller, Megan I.; Carlton, James (2018) Transoceanic rafting of Bryozoa (Cyclostomata, Cheilostomata, and Ctenostomata) across the North Pacific Ocean on Japanese tsunami marine debris, Aquatic Invasions 13(1): 137-162
DOI: https://doi.org/10.3391/ai.2018.13.1.11

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

Minchin, Dan; Cook, Elizabeth J.; Clark, Paul F. (2013) Alien species in British brackish and marine waters, Aquatic Invasions 8: in press

Morri, Carla; , Puce. Stefania; Bianchi. Carlo Rike Bitar, Ghazi , Zibrowius, Helmut; Bavestrello, Giorgio (2009) Hydroids (Cnidaria: Hydrozoa) from the Levant Sea (mainly Lebanon), with emphasis on alien species, Journal of the Marine Biological Association of the United Kingdom 89: 49-62
https://doi.org/10.1017/S0025315408002749

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
http://dx.doi.org/10.3391/ai.2015.10.1.11

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>



O'Shaughnessy, Kathryn A.; Lyons, David; Ashelby,Christopher W; R Counihan, Randall; Pears, Eliot; Taylor; Davies, Rebecca; PStebbing, aul D. (2-023) Rapid assessment of marine non-native species in Irish marinas, Management of Biological Invasions 14: 245–267
, https://doi.org/10. 3391/mbi.2023.14.2.05 Received: 4 August 2022

Occhipinti Ambrogi, A. (2001) Transfer of marine organisms: a challenge to the conservation of coastal biocoenoses, Aquatic Conservation: Marine and Freshwater Ecosystems 11: 243-251

Occhipinti Ambrogi, Anna (2000) Biotic invasions in a Mediterranean lagoon., Biological Invasions 2: 165-176

Occhipinti Ambrogi, Anna; D'Hondt, J.-L. (1994) Biology and Palaeobiology of Bryozoans, Olsen & Olsen, Fredensborg, Denmark. Pp. 139-144

Occhipinti Ambrogi, Anna; D'Hondt, Jean-Loup (1981) Distribution of bryozoans in brackish waters of Italy., In: Larwood, Gilbert P. and Nielsen, Claus(Eds.) Recent and Fossil Bryozoa.. , Fredensborg. Pp. <missing location>

Occhipinti-Ambrogi, A.; Savini, D. (2003) Biological invasions as a component of global change in stressed marine ecosystems., Marine Pollution Bulletin 46: 542-551

Occhipinti-Ambrogi, Anna (2007) Global change and marine communities: Alien species and climate change., Marine Pollution Bulletin 55: 342-352

Pederson, Judith, and 13 authors (2021) 2019 Rapid Assessment Survey of marine bioinvasions of southern New England and New York, USA, with an overview of new records and range expansions, Bioinvasions Records 10(2): 22-–237

Piola, Richard F.; Johnston, Emma L. (2008) Pollution reduces native diversity and increases invader dominance in marine hard-substrate communities., Diversity and Distributions 14: 329-342

Pongparadon, Supattra; Zuccarello, Giuseppe C.; Prathep, Anchana (2017) High morpho-anatomical variability in Halimeda macroloba (Bryopsidales, Chlorophyta) in Thai waters, Phycological Research 65: 136-145
doi: 10.1111/pre.12172

Porter, Joanne S.; Spencer Jones, Mary E.; Kuklinski, Piotr; Rouse, Sally (2015) First records of marine invasive non-native Bryozoa in Norwegian coastal waters from Bergen to Trondheim, BioInvasions Records 4: In press

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

Reverter-Gil. Oscar; Souto, Javier (2020) Watersiporidae (Bryozoa) in Iberian waters: an update on alien and native species, Marine Biodiversity 49: 2735-2752

Ribeiro, Romeu S.; Mata, Ana M. T. ; Salgado, Ricardo; Gandra, Vasco; Afonso, Inês; Galhanas, Dina; Dionísio, Maria Ana; Chainho, Paula (2023) Undetected non-indigenous species in the Sado estuary (Portugal), a coastal system under the pressure of multiple vectors of introduction, Journal of Coastal Conservation 27(53): Published online
https://doi.org/10.1007/s11852-023-00979-3

Rifi, Mouna; Basti, Leila; Rizzo, Lucia; Tanduo, Valentina; Radulovici, Adriana; Jaziri, Sabri; Uysal, Irfan; Souissi, Nihel; Mekki, Zeineb; Crocetta. Fabio (2023) Tackling bioinvasions in commercially exploitable species through interdisciplinary approaches: A case study on blue crabs in Africa’s Mediterranean coast (Bizerte Lagoon, Tunisia, Estuarine, Coastal and Shelf Science 291(108419): Published online
https://doi.org/10.1016/j.ecss.2023.108419

Ros, Macarena; Guerra-García, José M.; González-Macías, Manuel; Saavedra, Ángela; López-Fe, Carlos M. L (2013) Influence of fouling communities on the establishment success of alien caprellids (Crustacea: Amphipoda) in Southern Spain, Marine Biology Research 9(3): 261-273

Schmitt, Joseph D. Emmel, Jason A. Bunch, Aaron J. Hilling, Corbin D. Orth, Donald J (2019) Feeding ecology and distribution of an invasive apex predator: Flathead Catfish in subestuaries of the Chesapeake Bay, Virginia, North American Journal of Fisheries Management 39: 390-402
https://doi.org/10.1002/nafm.1027

Ulman, Aylin and 17 authors (2017) A massive update of non-indigenous species records in Mediterranean marinas, PeerJ 5( e3954): <missing location>

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

Zambrano, René; Ramos, John (2021) Alien crustacean species recorded in Ecuador, Nauplius 29: e2021043

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