Watersipora subtorquata

Invasion History

First Non-native North American Tidal Record: 1937
First Non-native West Coast Tidal Record: 1937
First Non-native East/Gulf Coast Tidal Record:

---

General Invasion History:

Watersipora subtorquata is an encrusting bryozoan widely distributed around the globe. Its native range is poorly understood because of taxonomic confusion with related species, particularly W. subovoidea, which it has been lumped with in older literature as 'W. cucullata' (Gordon 1989; Gordon and Mawatari 1992), or by treating W. subovoidea as a synonym of W. subtorquata (Seo 1999). This species was described from Rio de Janeiro, Brazil (Gordon 1989). We regard it as cryptogenic in the western Atlantic, where early records (e.g., Osburn 1914, Dry Tortugas; Osburn 1940, Puerto Rico) refer to W. cucullata, and where it has also been recorded as W. subovoidea (Winston 1982). In recent collections, it has been identified in Jamaica (Creary 2003), Puerto Rico (in 2007, Ruiz et al. unpublished data), and in Florida: Biscayne Bay (in 2004, Ruiz et al. unpublished data), Indian River Lagoon (in 2007, Ruiz et al. unpublished data), and Jacksonville (in 2002, Ruiz et al. unpublished data). This bryozoan also occurs in the Southwest Atlantic, on the west coast of South Africa (Florence et al. 2007, cited by Mead et al. 2011b). We also consider it cryptogenic in the Northwest Pacific, where it has been reported from the Sea of Japan and East China Sea coasts of South Korea, the Yellow Sea coast of Korea and China, and the Pacific coast of Japan (Tokyo Bay and southward) south to the Paracel Islands in the South China Sea (Seo 1999; Huang 2001).

In California, molecular surveys identified two clades of Watersipora subtorquata (Clades A and B), and an additional species (Watersipora n. sp.). Clade A is widely distributed globally, including Australia, New Zealand, and South Korea. Clade B is known from China and California (Mackie et al. 2012). In California, Clade A occurred from San Diego to Humboldt Bay, but was most abundant in southern and central-region harbors. Clade B occurred over the same range, but was usually less dominant, and was very spotty in the northern part of the range (Mackie et al. 2012).

Watersipora subtorquata has been introduced to the Northeast Pacific (1st record 1888, Gulf of California, and Cabo San Lucas to Puget Sound, Cohen and Carlton 1995; Cohen 2005; Ruiz et al. unpublished data), much of the coast of Australia (1st record 1950, Sydney Harbor, Winston 1977), New Zealand (1st Record 1983, Gordon and Mawatari 1992), and the Atlantic coast of France (1st Record 1973, d'Hondt 1984, cited by Ryland et al. 2009). This organism has a short planktonic stage (Gordon and Mawatari 1992; Cohen and Carlton 1995) suggesting that ship fouling is its likeliest mode of transport to most locations. However, its appearances in France appear related to culture of the Pacific Oyster (Crassostrea gigas) (Ryland et al. 2009).

North American Invasion History:

Invasion History on the West Coast:

Bryozoans of the Watersipora subtorquata complex have been introduced to the northeast Pacific (1st record 1888, Gulf of California, and Cabo San Lucas to Puget Sound, (Cohen and Carlton 1995; Cohen 2005; Ruiz et al. unpublished data). The invasion history of the Watersipora spp. on the west coast is murky, with conflicting molecular and morphological surveys. In California, molecular surveys identified two clades of Watersipora subtorquata (Clades A and B), and an additional species (Watersipora n. sp.). Clade A is widely distributed globally, including Australia, New Zealand, and South Korea. Clade B is known from China and California (Mackie et al. 2012). In California, Clade A occurred from San Diego to Humboldt Bay, but was most abundant in southern and central-region harbors. Clade B occurred over the same range, but was usually less dominant, and was very spotty in northern part of the range They found a third form, Watersipora n. sp., ranging from southern California to Puget Sound. This species has been named formally or described morphologically (Mackie et al. 2012). The molecularly defined species Clades A and B have not been studied morphologically and are unnamed as present. 
 
In a global survey of the genus Watersipora, Vieira et al. (2014) mapped three species on the West Coast, the well-defined W. arcuata, W. subatra, and the little-known W. atrofusca. Vieira et al. (2014) identified the well-defined W. arcuata and W. subatra, and W. atrofusca (known from Mexico and possibly southern California, little information available). Watersipora subatra ranges from Mexico to Puget Sound (Vieira et al. 2014; Ruiz et al. unpublished data). The 'true' W. suborquata was not shown on the West Coast of North America on Vieira et al.’s (2014) map. However, it has been found locally in Long Beach and other sites in southern California, but is less widespread (Linda McCann, personal communication).

Invasion History in Hawaii:

The earliest record for Watersipora subtorquata in Hawaii are specimens collected in 1966 in Pearl Harbor, and the Ala Wai Marina, near Honolulu, Oahu, in 1966 (Carlton and Eldredge, 2009). This bryozoan was collected on fouling plates in 2007 in Kaneohe Bay, Barbers Point Harbor, and several locations near Honolulu (Ruiz et al., unpublished data).

Invasion History Elsewhere in the World:

Since Watersipora subtorquata is a member of a complex of cryptic species, its native and introduced ranges are uncertain. Possible native regions include the tropical southwest Atlantic (type locality in Brazil), and the Indo-West Pacific (Vieira et al. 2014). It is found in many parts of the of the Mediterranean, from France to Egypt but the dates are unknown due to confusion with the native W. cucullata (Harmelin 2014; Vieira et al. 2014). In the eastern Atlantic, W. subtorquata was found in the Azores in 1888 (Chainho et al. 201), Madeira in 2006 (Canning-Clode et al. 2016), and South Africa (Florence et al. 2007; Mead et al. 2011). So far, specimens of Watersipora on the Atlantic coast of Europe (d'Hondt 1984; Ryland et al. 2009) have been identified or re-identified as W. subatra (Vieira et al. 2014). 

Watersipora subtorquata's occurrence in Europe was obscured by its confusion with W. subovoidea, but it was collected in the Bay of Arcachon, France on the Bay of Biscay in 1973 (d'Hondt 1984, cited by Ryland et al. 2009). It was collected again in the Bay of Arcachon in 2003, and in 1999–2007, found in many sites in Brittany and the Channel Islands (Ryland et al. 2009). Most of the collection sites were located near oyster-culture operations, and this species was most likely introduced to Europe with the Pacific Oyster (Crassostrea gigas) (Ryland et al. 2009). This bryozoan is also introduced on the west coast of South Africa (Florence et al. 2007, cited by Mead et al. 2011b).

---

Description

Watersipora subtorquata is an encrusting bryozoan, growing in single layers of flat substrates, but becomes multilamellar on rough substrates, and sometimes erect and leaf-like (foliaceous). Zooids are roughly elongate-rectangular, about twice as long as wide, being740–1,500 µm X 290–680 µm in size. The frontal shield is flat to slightly convex, perforated by numerous round pseudopores. Lateral-oral septulae (connecting the zooids) are absent. The orifice is slightly wider than the long, proximal edge, with a V-shaped or rounded sinus and approximately 130 x 260 (mean ~230) µm in size. It occupies less than 10% of the total zooid area with a well-defined proximal sinus demarcated by triangular condyles, ~55 × 115 µm (Ryland et al. 2009). The rim of the orifice has projecting proximo-lateral, and triangular condyles. The operculum has a dark central band and two lucida (transparent spots) adjacent to the condyles (Vieira et al. 2014). There are no oral spines or avicularia, and no ovicells. However, the zooids brood bright orange-red embryos internally. Colonies are orange to brownish-purple or black with dried or dead colonies becoming dark-orange or gray. (Description from Gordon and Mawatari 1992; Seo 1999; Ryland et al. 2009; Vieira et al. 2014). Colonies can become erect and leaf-like, with extensively overlapping calcareous crusts and curled edges. The crusts are often grayish black or dull orange, with bright orange (New Zealand) or purplish red (Korea) growing edges. The operculum is strongly pigmented with a dark, broad, biconcave band proximally, gradually spreading around paired clear areas. The polypides have orange lophophores 740 x 850 µm long, and 24 tentacles (Gordon 1989).  
 
NOTE: A recent revision of Watersipora taxonomy presented a drastic change in the nomenclature and worldwide biogeography of the genus. Vieira et al.'s (2014), map shows W. subtorquata as absent from the West Coast of North America and shows three species on the West Coast: W. subatra, W. atrofusca, and W. arcuata, with W. subatra identified only from California and Pacific Mexico. Watersipora subtorquata is widely distributed in the tropical-subtropical Atlantic and Mediterranean, Red Sea, and Indo-West Pacific. Another previously identified species, W. subovoidea, has been abandoned and partially synonymized with an earlier name W. cucullata, a name until recently regarded as obsolete (Vieira et al. 2015). This worldwide revision of the genus is at odds with current molecular studies (Mackie et al. 2006; Mackie et al. 2012). Vieira et al.'s (2014) map includes only a small number of West Coast samples. A larger regional sample will be required to reconcile morphological and molecular taxonomy of Watersipora on the West Coast of North America. Until this is available, we will use the names W. subtorquata and Watersipora n. sp. in NEMESIS. 

---

Taxonomy

Ecology

General:

Watersipora subtorquata is an encrusting, 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 (Barnes 1983). Watersipora subtorquata is known from pilings, rocks, shells, floats, oil platforms, ships' hulls, and fouling plates (Mackie et al. 2006; Page et al. 2006; Cohen and Zabin 2009; Ryland et al. 2009). 

 

Food:

Phytoplankton

Trophic Status:

Suspension Feeder

SusFed

---

Habitats

General Habitat	Coarse Woody Debris	None
General Habitat	Marinas & Docks	None
General Habitat	Rocky	None
General Habitat	Vessel Hull	None
General Habitat	Coral reef	None
Salinity Range	Polyhaline	18-30 PSU
Salinity Range	Euhaline	30-40 PSU
Tidal Range	Subtidal	None
Tidal Range	Low Intertidal	None
Vertical Habitat	Epibenthic	None

---

Life History

Ecology- Watersipora arcuata is known from pilings, rocks, floats and ships' hulls (Banta 1969; Gordon and Mawatari 1992).

---

Tolerances and Life History Parameters

Minimum Temperature (ºC)	6.7	Field, based on coldest site in geographical range, Port Townsend WA (Zerebecki and Sorte 2011)
Maximum Temperature (ºC)	30.6	Field, based on warmest site in geographical range, Red Sea (Zerebecki and Sorte 2011)
Minimum Salinity (‰)	25	Field salinity (California) (Cohen 2005)
Maximum Salinity (‰)	40	Field salinity (Shark Bay, Western Australia) (Wyatt et al. 2005)
Minimum Duration	0	Larva- Cohen and Carlton 1995
Maximum Duration	1	Larva- Cohen and Carlton 1995
Broad Temperature Range	None	Warm-Temperate-Tropical
Broad Salinity Range	None	Polyhaline-Euhaline

General Impacts

Watersipora subtorquata is an encrusting bryozoan widely distributed around the globe. Its colonies can be erect and leaf-like, with extensive overlapping calcareous crusts and curled edges, which create secondary habitat for the settlement of other marine invertebrates. Its native range is poorly understood because of taxonomic confusion with related species, particularly W. subatra. However, introduced populations have been recorded on the West coast of the United States, Hawaii, Australia, New Zealand, Europe, and South Africa. This species is known from rocks, oyster shells, pilings, floats, oil platforms, ships' hulls, and fouling plates. It is tolerant of copper and mercury antifouling paints and has outcompeted congeneric species in some areas of its introduced range.  

Economic Impacts Shipping and Boating- Watersipora spp. have long been known to be tolerant of copper and mercury in antifouling paint (Allen 1953; Ryland 1971; Piola and Johnston 2006). Their hard encrusting colonies are tolerant of moving water, and their colonies also provide non-toxic points of attachment for other organisms, allowing a diverse fouling community to develop (Floerl et al. 2004), which can adversely affect the speed and efficiency of ships. McKenzie et al. (2011) found that colonies from different sites varied in copper tolerance, and that tolerance was heritable in cultures. Colonies that produced large larvae tended to be more copper-tolerant (McKenzie et al. 2011).  

Ecological Impacts Competition- In New Zealand, W. subtorquata (arriving in 1983) quickly replaced W. arcuata at most locations (Gordon and Mawatari 1992). In Port Phillip Bay, Victoria, Australia W. subtorquata (arriving in 1976) also became the dominant Watersipora species (Keough and Ross 1999). In southern California, W. subtorquata was as of 2000–2003 the dominant or only form at several sites (Oceanside Harbor, Alamitos Bay, King Harbor; Geller et al. 2008 where W. arcuata was previously predominant (Banta 1969). Geller et al. (2008) suggest that W. subtorquata may be more competitive than W. arcuata in warm-temperate climates and may be expected to extend its range northward as the climate warms. Watersipora subtorquata was one of a group of seven non-native species in Bodega Harbor, most of which were rare or absent in 1970–1971 but were among the eight most abundant species in 2006. Spawning periods and abundance of species in this group appeared to be favored by a 1°C increase in average temperatures at this site over a 30-year period (Sorte and Stachowicz 2011).  

Habitat Change- Watersipora subtorquata colonies provide habitat for other organisms. Its colonies often develop elevated leaf-like folds rising above the substrate, creating additional space for colonization by other organisms. In Bodega Harbor, Geller et al. 2008 found where morphologically identical species (either W. subtorquata or Watersipora n. sp.) had more than 10% cover on fouling plates, native diversity was correlated with exotic diversity, as both groups of organisms occupied the expanded area (Sellheim et al. 2010). On the hulls of ships, and other surfaces treated with antifouling paints, W. subtorquata is often the only species able to settle, and its colonies provide surfaces on which more sensitive organisms can settle (Floerl et al. 2004). 

---

Regional Impacts

AUS-X	None		Economic Impact	Shipping/Boating
Shipping and Boating- Watersipora spp. have long been known to be tolerant of copper and mercury in antifouling paint (Piola and Johnston 2006). Their hard encrusting colonies are tolerant of moving water, and their colonies also provide non-toxic points of attachment for other organisms, allowing a diverse fouling community to develop (Floerl et al. 2007). Experimental studies on W. subtorquata's response to antifouling paint were done in Botany Bay and Port Jackson, near Sydney (Piola and Johnston 2006).
AUS-XII	None		Economic Impact	Shipping/Boating
Watersipora spp. have long been known to be tolerant of copper and mercury in antifouling paint (Allen 1950; Ryland 1971; Piola and Johnston 2006). Their hard encrusting colonies are tolerant of moving water, and their colonies also provide non-toxic points of attachment for other organisms, allowing a diverse fouling community to develop (Floerl et al. 2004). Experimental studies on W. subtorquata's response to antifouling paint were done in Townsville and Cairns, Queensland (Floerl et al. 2007).
NZ-IV	None		Ecological Impact	Competition
In New Zealand, W. subtorquata, arriving in 1983, quickly replaced W. arcuata at most locations (Gordon and Mawatari 1992).
NZ-VI	None		Ecological Impact	Competition
In New Zealand, W. subtorquata, arriving in 1983, quickly replaced W. arcuata at most locations (Gordon and Mawatari 1992).
AUS-VIII	None		Ecological Impact	Competition
In Port Phillip bay, W. subtorquata, arriving in 1976, quickly replaced W. arcuata at most locations (Keough and Ross 1999). Heavy recruitment of W. subtorquata can affect subsequent community development. However, impacts varied by site and season (Sams and Keough 2012).
NEP-VI	Pt. Conception to Southern Baja California		Ecological Impact	Competition
In southern California, W. subtorquata is now (2000-2003) the dominant or only form at several sites (Oceanside Harbor, Alamitos Bay, King Harbor; Geller et al. 2008) where W. arcuata was previously dominant (Banta 1969). Geller et al. (2008) suggest that W. subtorquata may be more competitive than W. arcuata in warm-temperate climates, and may be expected to extend its range northward as the climate warms. Experimental clearing and routine cleaning of oil platforms favored increased abundance of Watersipora subatra, recruitng in the cleared area (Viola et al. 2018). Viola et al. (2018) suggest the retention of oil platforms as artificial reefs could favor W. subatra. Removal of the upper portions of the platforms might reduce their potential as a source for coastal populations of the bryozoan.
P023	_CDA_P023 (San Louis Rey-Escondido)		Ecological Impact	Competition
In southern California, W. subtorquata is now (2000-2003) the dominant or only form at several sites (Oceanside Harbor, Alamitos Bay, King Harbor; Geller et al. 2008) where W. arcuata was previously dominant (Banta 1969). Geller et al. (2008) suggest that W. subtorquata may be more competitive than W. arcuata in warm-temperate climates, and may be expected to extend its range northward as the climate warms.
P050	San Pedro Bay		Ecological Impact	Competition
In southern California, W. subtorquata is now (2000-2003) the dominant or only form at several sites (Oceanside Harbor, Alamitos Bay, King Harbor; Geller et al. 2008) where W. arcuata was previously dominant (Banta 1969). Geller et al. (2008) suggest that W. subtorquata may be more competitive than W. arcuata in warm-temperate climates, and may be expected to extend its range northward as the climate warms.
P060	Santa Monica Bay		Ecological Impact	Competition
In southern California, W. subtorquata is now (2000-2003) the dominant or only form at several sites (Oceanside Harbor, Alamitos Bay, King Harbor; Geller et al. 2008) where W. arcuata was previously dominant (Banta 1969). Geller et al. (2008) suggest that W. subtorquata may be more competitive than W. arcuata in warm-temperate climates, and may be expected to extend its range northward as the climate warms.
AUS-XII	None		Ecological Impact	Habitat Change
On the hulls of ships and other surfaces, treated with antifouling paints, W. subtorquata is often the only species able to settle, and its colonies provide surfaces on which more sensitive organisms can settle. Experimental studies were performed in Cairns and Townsville, Queensland (Floerl et al. 2004).
AUS-X	None		Ecological Impact	Competition
Exposure to copper anti-fouling paint resulted in enhanced recruitment of W. subtorquata, despite post-settlement mortality. Surviving colonies had shorter ancestrulae and were smaller (MacKenzie et al. 2012). The ability to tolerate and even be favored by copper pollution gives W. subtorquata a competitive advantage in polluted habitats (MacKenzie et al. 2011; MacKenzie et al. 2012a)
P065	_CDA_P065 (Santa Barbara Channel)		Ecological Impact	Competition
Experimental clearing and routine cleaning of oil platforms favored increased abundance of Watersipora subatra, recruitng in the cleared area (Viola et al. 2018). Viola et al. (2018) suggest the retention of oil platforms as artificial reefs could favor W. subatra. Removal of the upper portions of the platforms might reduce their potential as a source for coastal populations of the bryozoan.
CA	California		Ecological Impact	Competition
In southern California, W. subtorquata is now (2000-2003) the dominant or only form at several sites (Oceanside Harbor, Alamitos Bay, King Harbor; Geller et al. 2008) where W. arcuata was previously dominant (Banta 1969). Geller et al. (2008) suggest that W. subtorquata may be more competitive than W. arcuata in warm-temperate climates, and may be expected to extend its range northward as the climate warms., In southern California, W. subtorquata is now (2000-2003) the dominant or only form at several sites (Oceanside Harbor, Alamitos Bay, King Harbor; Geller et al. 2008) where W. arcuata was previously dominant (Banta 1969). Geller et al. (2008) suggest that W. subtorquata may be more competitive than W. arcuata in warm-temperate climates, and may be expected to extend its range northward as the climate warms., In southern California, W. subtorquata is now (2000-2003) the dominant or only form at several sites (Oceanside Harbor, Alamitos Bay, King Harbor; Geller et al. 2008) where W. arcuata was previously dominant (Banta 1969). Geller et al. (2008) suggest that W. subtorquata may be more competitive than W. arcuata in warm-temperate climates, and may be expected to extend its range northward as the climate warms., Experimental clearing and routine cleaning of oil platforms favored increased abundance of Watersipora subatra, recruitng in the cleared area (Viola et al. 2018). Viola et al. (2018) suggest the retention of oil platforms as artificial reefs could favor W. subatra. Removal of the upper portions of the platforms might reduce their potential as a source for coastal populations of the bryozoan.

References

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

Marasinghe, M. M. K.I ; Ranatunga, R. R. M .K. P. Anil, Arga C. (2018) Settlement of non-native Watersipora subtorquata (d'Orbigny, 1852) in artificial collectors deployed in Colombo Port, Sri Lanka, BioInvasions Records 7: 7-14

Abdel-Salam, Kh. M.; Ramadan, Sh. E. (2008) Fouling Bryozoa from some Alexandria harbours, Egypt. (II) Encrusting species, Mediterranean Marine Science 9(2): 5-20

Abdelsalam, Khaled Mahmood (2018) First record of the exotic lysmatid shrimp Lysmata vittata (Stimpson, 1860) (Decapoda: Caridea: Lysmatidae) from the Egyptian Mediterranean coast, Mediterranean Marine Science 19(1): 124-131

Allen, F. E. (1953) Distribution of marine invertebrates by ships, Australian Journal of Marine and Freshwater Research 4(2): 307-316

Ashton, Gail; Davidson, Ian; Ruiz, Gregory (2014) Transient small boats as a long-distance coastal vector for dispersal of biofouling organisms, Estuaries and Coasts 37: 1572-1581

Australian Museum Business Services (2002) <missing title>, Australian Museum Business Services, for Sydney Ports Corporation, Sydney. Pp. <missing location>

Baker, H. R. (1984) Diversity and zoogeography of marine Tubificidae (Annelida, Oligochaeta), with notes on variation in widespread species, Hydrobiologia 115: 191-196

Banta, W. C. (1969) The recent introduction of Watersipora arcuata Banta (Bryozoa, Cheilostomata) as a fouling pest in Southern California, Bulletin of the Southern California Academy of Sciences 68: 248-251

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

Birdsey, Emma M.; Johnston, Emma L.; Poore, Alistair G. B. (2012) Diversity and cover of a sessile animal assemblage does not predict its associated mobile fauna, Marine Biology 15: <missing location>

Boyd, Milton J.; Mulligan, Tim J; Shaughnessy, Frank J. (2002) <missing title>, California Department of Fish and Game, Sacramento. Pp. 1-118

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

Campbell, Marnie L] ; Hewitt, Chad L.[ Miles, Joel (2016) Marine pests in paradise: capacity building, awareness raising and preliminary introduced species port survey results in the Republic of Palau, Biological Invasions 7(4): 351-363

Canning-Clode, João; Fofonoff, Paul; McCann, Linda; Carlton, James T.; Ruiz, Gregory (2013) Marine invasions on a subtropical island: fouling studies and new records in a recent marina on Madeira Island (Eastern Atlantic Ocean), Aquatic Invasions 8(3): 261-270

Carlton, James T. (1979) History, biogeography, and ecology of the introduced marine and estuarine invertebrates of the Pacific Coast of North America., Ph.D. dissertation, University of California, Davis. Pp. 1-904

Carlton, James T.; Eldredge, Lucius (2009) Marine bioinvasions of Hawaii: The introduced and cryptogenic marine and estuarine animals and plants of the Hawaiian archipelago., Bishop Museum Bulletin in Cultural and Environmental Studies 4: 1-202

Cesar-Aldariz, J.; Fernandez, E.; Reverter-Gil, O. (1997) Briozoos de la costa Lugo (N. O. Espana), Nova Acta Cientifica Compostelana (Bioloxia) 7: 207-220

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>

Clark, Graeme F.; Johnston, Emma L (2009) Propagule pressure and disturbance interact to overcome biotic resistance of marine invertebrate communities, Oikos 118: 1679-1686

Clark, Graeme F.; Johnston, Emma L. (2010) Temporal change in the diversity-invasibility relationship in the presence of a disturbance regime, Ecology Letters 14: 52-57

Cockrell, Marcy L.; Sorte, Cascade J.B. (2013) Predicting climate-induced changes in population dynamics of invasive species in a marine epibenthic community, Journal of Experimental Marine Biology and Ecology 440: 42-48

Cohen, Andrew N. 2005-2024 Exotics Guide- Non-native species of the North American Pacific Coat. https://www.exoticsguide.org/



Cohen, Andrew N. and 10 authors (2005) <missing title>, San Francisco Estuary Institute, Oakland CA. Pp. <missing location>

Cohen, Andrew N. and 12 authors (2002) Project report for the Southern California exotics expedition 2000: a rapid assessment survey of exotic species in sheltered coastal waters., In: (Eds.) . , Sacramento CA. Pp. 1-23

Cohen, Andrew N.; Carlton, James T. (1995) Nonindigenous aquatic species in a United States estuary: a case study of the biological invasions of the San Francisco Bay and Delta, U.S. Fish and Wildlife Service and National Sea Grant College Program (Connecticut Sea Grant), Washington DC, Silver Spring MD.. Pp. <missing location>

Cohen, Andrew N.; Zabin, Chela J. (2009) Oyster shells as vectors for exotic organisms, Journal of Shellfish Research 28(1): 163-167

Coles, S. L.; Reath, P. R.; Skelton, P. A.; Bonito, V; DeFelice; Basch, L. (2003) Introduced marine species in Pago Pago Harbor, Fagatele Bay and the National Park Coast, American Samoa., Bishop Museum Technical Report 26: 1-24

Coutts, Ashley D. M.; Moore, Kirrily M.; Hewitt, Chad L. (2003) Ships' sea-chests: an overlooked transfer mechanism for non-indigenous marine species., Marine Pollution Bulletin 46: 1504-1515

Cranfield, H.J.; Gordon, D.P.; Willan, R.C.; Marshall, B.A; Battershill, C.N.; Francis, M.P.; Nelson, W.A.; Glasby, C.J.; Read, G.B. (1998) <missing title>, The National Institute of Water and Atmospheric Research, New Zealand. Pp. <missing location>

Creary, Marcia M. (2003) A simplified field guide to the bryozoan species found on the roots of the red mangrove (Rhizophora mangle) in and around Kingston Harbour, Jamaica, W.I., Bulletin of Marine Science 73(2): 521-526

Creary, Marcia; Webber, Mona (2009) <missing title>, Environmental Foundation of Jamaica, Kingston, Jamaica. Pp. 1-18

Crivellaro, Marcelo Schuler; Candido, Davi Volney; ilveira, Thiago Cesar Lima; Fonseca, Adriana Carvalhal; Segal, Barbara ´ (2022) A tool for a race against time: Dispersal simulations to support ongoing monitoring program of the invasive coral Tubastraea coccinea, Marine Pollution Bulletin 185(114354): Published online
https://doi.org/10.1016/j.marpolbul.2022.114354

Currie, D. R.; McArthur, M. A.; Cohen, B. F. (1999) Exotic Marine Pests in the Port of Geelong, Victoria, In: Hewitt, Campbell, Thresher & Martin(Eds.) Marine Biological Invasions of Port Phillip Bay, Victoria. , Hobart, Tasmania. Pp. 227-246

David, Andrew A.; Cote, Sara C. (2019) Genetic evidence confirms the presence of the Japanese mystery snail, Cipangopaludina japonica (von Martens, 1861) (Caenogastropoda: Viviparidae) in northern New York, BioInvasions Records 8: 793-803

de Rivera, Catherine, and 27 authors (2005) Broad-scale non-indigenous species monitoring along the West Coast in National Marine Sanctuaries and National Estuarine Research Reserves report to National Fish and Wildlife Foundation, National Fish and Wildlife Foundation, Washington, D.C.. Pp. <missing location>

Duncan, Meredyth; Chow, Benson; Myron, Kevin; Stone, Jaden; Hubbell, Mark; Schriock, Elizabeth; Hunt, Carol; Khtikian, W. Kent ; Cohen, C. Sarah (2022) First report of genetic data from two invasive Watersipora (Bryozoa) species in the central California coast rocky intertidal, Aquatic Invasions 17: Published online

Eitan, G. (1972) Additions to the bryozoan fauna of the Suez canal., Israel Journal of Zoology 21: 377-384

Fairey, Russell; Dunn, Roslyn; Sigala, Marco; Oliver, John (2002) Introduced aquatic species in California's coastal waters: Final Report, California Department of Fish and Game, Sacramento. Pp. <missing location>

Ferreira-Rodriguez, Noe; Pavel, Ana Bianca; Cogalniceanu, Dan (2021) Integrating expert opinion and traditional ecological knowledge in invasive alien species management: Corbicula in Eastern Europe as a model, Biological Invasions 23: 1087-1099

Floerl, Oliver; Pool, Thomas K.; Inglin, Graeme J. (2004) Positive interactions between nonindigenous species facilitate transport by human vectors., Ecological Applications 14(6): 1724-1736

Florence, Wayne K.; Hayward. Peter J.; Gibbons, Mark J. (2007) Taxonomy of shallow-water Bryozoa from the west coast of South Africa, African Natural History 3: 1-58

Gauff, Robin P. M.; Bouchoucha, Marc; Curd, Amelia; Droual, Gabin; Evrard, Justine; Gayet, Nicolas; Nunes. Flavia (2023) First joint morphological and molecular detection of Watersipora subatra in the Mediterranean Sea presented in an updated genus phylogeny to resolve taxonomic confusion, Aquatic Invasions 18(3): 295-312

Glasby, Tim M.; Connell, Sean D.; Holloway, Michael G.; Chad L. Hewitt (2007) Nonindigenous biota on artificial structures: could habitat creation facilitate biological invasions?, Marine Biology 151: 887-895

Goddard, Jeffrey H. R.; Love, Milton S. (2010) Megabenthic invertebrates on shell mounds associated with oil and gas platforms off California, Bulletin of Marine Science 86: 533-554

Gordon, D. P. (1989) The marine fauna of New Zealand: Bryozoa: Gymnolaemata (Cheilostomida Ascophorina) from the Western South Island continental shelf and slope., New Zealand Oceanographic Institute Memoir 97: 5-95

Gordon, Dennis P. (2016) Bryozoa of the South China Sea: an overview, Raffles Bulletin of Zoology 34: 604-618

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

Green, Stephanie J. and 7 authors (2021) Broad-scale acoustic telemetry reveals long-distance movements and large home ranges for invasive lionfish on Atlantic coral reefs, Marine Ecology Progress Series 673: 117-134

Grey, Erin K. (2009) Scale-dependent relationships between native richness, resource stability and exotic cover in dock fouling communities of Washington, USA, Diversity and Distributions 15: 1073-1080

Harmelin, Jean-Georges (2014) Alien bryozoans in the eastern Mediterranean Sea- new records from the coast of Lebanon, Zootaxa 3893: 301-338

Hart, Simon P.; Keough, Michael J. (2009) Does size predict demographic fate? Modular demography and constraints on growth determine response to decreases in size, Ecology 90(6): 1670-1678

Hart, Simon P.; Burgin, Jacqueline R.; Marshall, Dustin J. (2012) Revisiting competition in a classic model system using formal links between theory and data, Ecology 93(12): 2015-2022

Hayward, Peter J.; McKinney, Frank K. (2002) Northern Adriatic Bryozoa from the vicinity of Rovinj, Croatia., Bulletin of the American Museum of Natural History 270: 1-139

Hedge, Luke H.; Johnston, Emma L. (2012) Propagule pressure determines recruitment from a commercial shipping pier, Biofouling 28(1): 73-85

Huang, Zongguo (Ed.), Junda Lin (Translator) (2001) Marine Species and Their Distributions in China's Seas, Krieger, Malabar, FL. Pp. <missing location>

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

Invasive Species Specialist Group 2001-2016 100 Of The World's Worst Invasive Species. <missing URL>



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

Keough, M. J. ; Ross, J. (1999) Introduced fouling species in Port Phillip Bay., In: Hewitt, C. L.; Campbell; M.;Thresher, R.; Martin,(Eds.) Marine Biological Invasions of Port Phillip Bay, Victoria. , Hobart, Tasmania. Pp. 193-225

Kim, Daemin; Taylor, Andrew T.; Near, Thomas J. (2022) Phylogenomics and species delimitation of the economically important Black Basses (Micropterus), Scientific Reports 12(9113): Published online
https://doi.org/10.1038/s41598-022-11743-2

Koçak, Ferah (2007) Bryozoan assemblages at some marinas in the Aegean Sea, JMBA2- Biodiversity Records (online) <missing volume>: <missing location>

Kocak, Ferah (2023) Alien bryozoan species along the Aegean coast of Turkey, Cahiers de Biologie Marine 64: 137-151
DOI: 10.21411/CBM.A.B3634AD9

Lange, Rolanda; Marshall, Dustin J. (2016) Relative contributions of offspring quality and environmental quality to adult field performance, Oikos 125: 210-217

Laruson, Aki J.; Craig, Sean F.; Messer, Kirk J.; Mackie, Joshua A. (2012) Rapid and reliable inference of mitochondrial phylogroups among Watersipora species, an invasive group of ship-fouling species (Bryozoa, Cheilostomata), Conservation Genetic Resources 4: 617-619

Leca, L.; d'Hondt, J.-L. (1993) [Hippopetraliella tahitiensis n. sp. n. sp., a new Cheilostome bryozoan (Petraliellidae) from French Polynesia], Cahiers de Biologie Marine 34: 401-209

Leonard, Kaeden; Hewitt , Chad L. Campbell, Marnie L.; Carmen Primo; Miller, Steven D. (2017) Epibiotic pressure contributes tobiofouling invader success, Scientific Reports <missing volume>(173): <missing location>

Li, C. Y. (1989) Collections of Papers on Marine Ecology in the Daya Bay, Third Institute of Oceanography (SOA) China Ocean Press, China. Pp. 106-111

Lins, Daniel M. ; Rocha, Rosana M. (2023) Marine aquaculture as a source of propagules of invasive fouling species , Polar Biology 11(e5456): Published online

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

Long, Edward R.; Rucker, James B. (1969) A comparative study of cheilostome bryozoa at Yokosuka, Maizuru, and Sasebo, Japan, Pacific Science 23: 56-69

Looby, Audrey; Ginsburg, David W. (2021) Nearshore species biodiversity of a marine protected area off Santa Catalina Island, California, Western North American Naturalist 81(1): 113-130

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

Mackie, Joshua A.; Darling, John A.; Geller, Jonathan B. (2012) Ecology of cryptic invasions: latitudinal segregation among Watersipora (Bryozoa) species, Scientific Reports 2(871): 1-10

Mackie, Joshua A.; Keough, Michael J.; Christidis, Les (2006) Invasion patterns inferred from cytochrome oxidase I sequences in three bryozoans, Bugula neritina, Watersipora subtorquata; and Watersipora arcuata., Marine Biology 149: 285-295

Mackie, Joshua A.; Wostenberg, Darren; Doan, Michael; Craig, Sean F. ; Darling, John A. (2014) High-throughput Illumina sequencing and microsatellite design in Watersipora (Bryozoa), a complex of invasive species, Conservation Genetic Resources 6: 1053-1055

Marraffini, M. L.; Geller, J. B. (2015) Species richness and interacting factors control invasibility of a marine community, Proceedings of the Royal Society of London B 282: Published online

McKenzie, Louise A.; Brooks, Robert C.; Johnston, Emma L. (2012b) A widespread contaminant enhances invasion success of a marine invader, Journal of Applied Ecology 49(4): 767-773

McKenzie, Louise A.; Brooks, Rob; Johnston, Emma L. (2011) Heritable pollution tolerance in a marine invader, Environmental Research 111: 926-932

McKenzie, Louise A.; Johnston, Emma L.; Brooks, Robert (2012a) Using clones and copper to resolve the genetic architecture of metal tolerance in a marine invader, Ecology and Evolution 2(6): 1319-1329

Mead, A.; Carlton, J. T.; Griffiths, C. L.; Rius, M. (2011a) Revealing the scale of marine bioinvasions in developing regions: a South African re-assessment, Biological Invasions 13(9): 1991-2008

Mead, A.; Carlton, J. T.; Griffiths, C. L. Rius, M. (2011b) Introduced and cryptogenic marine and estuarine species of South Africa, Journal of Natural History 39-40: 2463-2524

Mook, David (1983) Indian River fouling organisms, a review, Florida Scientist 26(3/4): 162-167

Moran, Z.; Orth, D. J.; Schmitt, J. D.; Hallerman. E.; Aguilar, R. (2016) Effectiveness of DNA barcoding for identifying piscine prey items in stomach contents of piscivorous catfishes , Environmental Biology of Fishes 19: 161–167
DOI 10.1007/s10641-015-0448-

Needles, Lisa A. (2007) <missing title>, M.S. Thesis, California Polytechnic State University, San Luis Obispo. Pp. <missing location>

Needles, Lisa A.; Gosnell, J. Stephen; Waltz, Grant T.; Wendt, Dean E.; Gaines, Steven D. (2015) Trophic cascades in an invaded ecosystem: native keystone predators facilitate a dominant invader in an estuarine community, Oikos 124: 1282-1292

Needles, Lisa A.; Wendt, Dean E. (2013) Big changes to a small bay: Introduced species and long-term compositional shifts to the fouling community of Morro Bay (CA), Biological Invasions 15(6): 1231-1251

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



Nyberg, Cecilia D.; Thomsen, Mads S.; Wallentinus, Inger (2009) Flora and fauna associated with the introduced red alga Gracilaria vermiculophylla, European Journal of Phycology 44(3): 395-403

Osburn, R. C. (1914) The Bryozoa of the Tortugas Islands, Florida, Papers from the Tortugas Islands Laboratory of the Carnegie Institution 5: 181-222

Osburn, Raymond C. (1940) Bryozoa of Porto Rico, N. Y. Academy of Sciences - Scientific Survey of Puerto Rico and the Virgin Islands 16(3): 321-486

Page, Henry M.; Dugan, Jenifer E.; Culver, Carolynn S.; Hoesterey, Justin C. (2006) Exotic invertebrate species on offshore oil platforms., Marine Ecology Progress Series 325: 101-107

Panicz, R., Eljasik, P.; Wrzecionkowski, K.; Smietana, N. Biernaczyk. M. (2022) First report and molecular analysis of population stability of the invasive Gulf wedge clam, Rangia cuneata (G.B. Sowerby I, 1832) in the Pomerian Bay (Southern Baltic Sea), European Journal of Zoology 89(1): 568-578
https://doi.org/10.1080/24750263.2022.2061612

Pestana, L. B.; Dias, G. M.; Marques, A. C . (2021) Spatial and temporal diversity of non-native biofouling species associated with marinas in two Angolan bays, African Journal of Marine Science 42(4): 413-422

Piola, Richard F.; Johnston, Emma L. (2006a) Differential resistance to extended copper exposure in four introduced bryozoans., Marine Ecology Progress Series 311: 103-114

Pister, Benjamin (2009) Urban marine ecology in southern California: the ability of riprap structures to serve as rocky intertidal habitat, Marine Biology 156: 861-873

Pollard, D.A.; Pethebridge, R.L. (2002) Report on Port of Botany Bay: Introduced marine pest species survey, NSW Fisheries Final Report Series 40: 1-69

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

Quintanilla, Elena; Thomas Wilke; Ramırez-Portilla, Catalina; Sarmiento, Adriana; Sanchez, Juan A. () , None <missing volume>: <missing location>

Revanales, Triana; Guerra-García, José M.; Ros, Macarena (2022) Colonization dynamics of potential stowaways inhabiting marinas: Lessons from caprellid crustaceans, Water 14(2659.): Published online
doi.org/10.3390/w14172659

Rodriguez, Laura F.; Ibarra-Obando, Silvia E. (2008) Cover and colonization of commercial oyster (Crassostrea gigas) shells by fouling organisms in San Quintin Bay, Mexico, Journal of Shellfish Research 27(2): 337-343

Rogers, Tanya L.; Byrnes, Jarrett E.; Stachowicz, John J. (2016) Native predators limit invasion of benthic invertebrate communities in Bodega Harbor, California, USA, Marine Ecology Progress Series 545: 161-173

Ruiz, Gregory M.; Geller, Jonathan (2018) Spatial and temporal analysis of marine invasions in California, Part II: Humboldt Bay, Marina del Re, Port Hueneme, and San Francisco Bay, Smithsonian Environmental Research Center & Moss Landing Laboratories, Edgewater MD, Moss Landing CA. Pp. <missing location>

Ryland, J. S. (1971) Bryozoa (Polyzoa) and marine fouling., In: Gareth Jones E.B; Eltringham, S. K.(Eds.) Marine Borers, fungi and fouling organisms of wood. , Paris. Pp. 137-154

Ryland, John S.; De Blauwe, Hans; Lord, Richard; Mackie, Joshua A. (2009) Recent discoveries of alien Watersipora (Bryozoa) in Western Europe, with redescriptions of species, Zootaxa 2093: 43-59

Sams, Michael A.; Keough, Michael J.1 AND MICHAEL J. KEOUGH (2012) Contrasting effects of variable species recruitment on marine sessile communities, Ecology 93: 1134-1146

Sellheim, Kirsten; Stachowicz, John J.; Coates, R. Cameron (2010) Effects of a nonnative habitat-forming species on mobile and sessile epifaunal communities, Marine Ecology Progress Series 398: 69-80

Seo, Ji Eun (1999) Taxonomic review of Korean Watersipora (Bryozoa, Gymnolaemata, Cheilostomata)., Korean Journal of Systematic Zoology 15(2): 221-229

Smale, Dan A.; Wernberg, Thomas (2012) Short-term in situ warming influences early development of sessile assemblages, Marine Ecology Progress Series 453: 129-136

Soors, Jan; Faasse, Marco; Stevens, Maarten; Verbessem, Ingrid; De Regge, Nico;Van den Bergh, Ericia (2010) New crustacean invaders in the Schelde estuary (Belgium), Belgian Journal of Zoology 140: 3-10

Sorte, Cascade J. B.; Stachowicz, John J. (2011) Patterns and processes of compositional change in a California epibenthic community, Marine Ecology Progress Series 435: 63-74

Sorte, Cascade, J. B.; Williams, Susan L.; Zerebecki, Robyn A. (2010) Ocean warming increases threat of invasive species in a marine fouling community, Ecology 91(8): 2198-2204

Soule, Dorothy F.; Soule, John D.; Morris, Penny A.; Chaney, Henry A (2007) The Light and Smith Manual: Intertidal Invertebrates from Central California to Oregon (4th edition), University of California Press, Berkeley CA. Pp. 866-904

Stachowicz, John J.; Byrnes, Jarrett E. (2006) Species diversity, invasion success, and ecosystem functioning: disentangling the influence of resource competition, facilitation, and extrinsic factors., Marine Ecology Progress Series 311: 251-262

Stafford, Nathaniel B.; Bell, Susan S. (2006) Space competition between seagrass and Caulerpa prolifera (Forsskal) Lamouroux following simulated disturbances in Lassing Park,FL, Journal of Experimental Marine Biology and Ecology 333: 49-57

Svanfeldt, Karin ; Monro, Keyne; Marshall, Dustin J. (2017) Dispersal duration mediates selection on offspring size, Oikos 126: 480-487,

U.S. National Museum of Natural History 2002-2021 Invertebrate Zoology Collections Database. http://collections.nmnh.si.edu/search/iz/



Vail, Lyle L.; Wass, Robin E. (1981) Experimental studies on the settlement and growth of Bryozoa in the natural environment., Australian Journal of Marine and Freshwater Research 32: 639-656

van Soest, R. W. M. (1976) First European record of Haliclona loosanoffi Hartman, 1958 (Porifera, Haplosclerida), a species hitherto known only from the New England coast (U.S.A.)., Beaufortia 24(316): 177-187

Vieira, Leandro M.; Jones, Mary Spencer; Taylor, Paul D. (2014) The identity of the invasive fouling bryozoan Watersipora subtorquata (d’Orbigny) and some other congeneric species, Zootaxa 3857: 151-182

Vieira, Leandro M.; Migotto, Alvaro E.; Winston, Judith E. (2008) Synopsis and annotated checklist of Recent marine Bryozoa from Brazil, Zootaxa 1810: 1-39

Ward, David L.; Finch, Colton; Blasius, Heidi (2015) Could high salinity be used to control bullfrogs in small ponds? , Journal of the Arizona-Nevada Academy of Science 46(2): 50-52

Wasson, Kerstin; Zabin, C. J.; Bedinger, L.; Diaz, M. C.; Pearse J. S. (2001) Biological invasions of estuaries without international shipping: the importance of intraregional transport, Biological Conservation 102: 143-153

Wiltshire, K.; Rowling, K.; Deveney, M. (2010) <missing title>, South Australian Research and Development Institute, Adelaide. Pp. 1-232

Winston, Judith E. (1977) Distribution and ecology of estuarine ectoprocts: a critical review., Chesapeake Science 18(1): 34-57

Winston, Judith E. (1982) Marine bryozoans (Ectoprocta) of the Indian River area (Florida)., Bulletin of the American Museum of Natural History 173: 99-176

Winston, Judith E. (1986) An annotated checklist of coral-associated bryozoans, American Museum Novitates 2859: 1-39

Winston, Judith E. (1995) Ectoproct diversity of the Indian River coastal lagoon, Bulletin of Marine Science 57(1): 84-93

Winston, Judith E. (2010) Stability and change in the Indian River area bryozoan fauna over a twenty-four year period, Smithsonian Contributions to the Marine Sciences 38: 229-239

Wyatt, Alex S. J.; Hewitt, Chad L.; Walker, Di I.; Ward, Trevor J. (2005) Marine introductions in the Shark Bay world heritage property, Western Australia: a preliminary assessment., Diversity and Distributions 11: 33-44

Zabin, Chela J.; Marraffini, Michelle; Lonhart, Steve I.; McCann, Linda · Ceballos, Lina; King, Chad (2018) Non-native species colonization of highly diverse, wave swept outer coast habitats in central california, Marine Biology 165(31): Published online

Zerebecki, Robyn A.; Sorte, Cascade J. B. (2011) Temperature tolerance and stress proteins as mechanisms of invasive species success, PLOS ONE 6(4): e14806, online

---