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:

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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 unpublished data), and in Florida: Biscayne Bay (in 2004, Ruiz et al. unpublished data), Indian River Lagoon (in 2007, Ruiz et al. unpublished data), and Jacksonville (in 2002, Ruiz et al. unpublished data).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 unpublished data), much of the coast of Australia (1st record 1950, Sydney Harbor, Winston 1977), New Zealand (1st Record 1983, Gordon and Mawatari 1992), and the Atlantic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).

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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. 

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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

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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

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Life History

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

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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). 

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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.

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