Diplosoma listerianum

Invasion History

First Non-native North American Tidal Record: 1899
First Non-native West Coast Tidal Record: 1899
First Non-native East/Gulf Coast Tidal Record: 1975

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General Invasion History:

Diplosoma listerianum was first described from the English Channel by Milne-Edwards in 1841. Initially, this species was described, under different names, from many different places around the world, such as Australia (D. rayneri Macdonald 1859) Brazil (D. macdonaldi Herdman 1886), and Japan (D. mitsukurii Oka 1892). These and many other names were synonymized by Kott (1990; 2001). This tunicate is now said to have a cosmopolitan distribution but represents a species complex (Perez-Portela et al. 2013). In most of these regions, D. listerianum was already established when researchers began surveying tunicates and, as a result, many populations have not been recognized as true introductions. One cryptic species (clade A) occurred in most of the sites sampled worldwide, while three other clades had more local distributions (Perez-Portela et al. 2013). We consider this species complex to be cryptogenic (of unknown origin) through most of its range. However, it appears to be a definite introduction in several locations, including the Northeast Pacific (California to British Columbia), the Northeast Atlantic (north of Cape Hatteras, North Carolina), Hawaii, New Zealand, and South Africa. In European waters, near the site of first description, range expansions have been noted in the North Sea (UK, Vance et al. 2008) and in the Netherlands (1st Record 1977, Gittenberger et al. 2007).

North American Invasion History:

Invasion History on the West Coast:

Diplosoma listerianum was collected in San Diego Bay, California in 1917. It was described by Ritter and Forsyth as D. pizoni from a single colony (Van Name 1945). Five additional colonies collected in 1899 were also identified as D. pizoni by Ritter (Eldredge 1966). Diplosoma listerianum apparently expanded its range northward, reaching Monterey Bay by 1939 (Van Name 1945; Eldredge 1966). In 1948, D. listerianum was found in San Francisco Bay, California (Eldredge 1966). In 1960, it was collected on Vancouver Island, British Columbia and by 1966 it was common in Puget Sound, Washington (Lambert 1966). 
 
Since D. listerianum was treated as a native species, we have not been able to find first dates of record for many West Coast water bodies. By 1980, it was considered common from San Diego, California to British Columbia, Canada (Abbott and Newberry 1980). In Southern California, Lambert and Lambert (1998) described it as occurring in 'all harbors, at all times of year' and subsequent surveys have confirmed this (Fairey et al. 2001; de Rivera et al. 2006; Ruiz et al., unpublished data). Further north, it occurs in Humboldt Bay, CA (Fairey et al. 2003; Ruiz et al. unpublished data) and Coos Bay, Oregon (1988, Hewitt 1993, 2001, Ruiz et al., unpublished data). It is widespread throughout Puget Sound (Ruiz et al., unpublished data) and has been collected at as far north as Bamfield, Vancouver Island, British Columbia (Mackie and Singla 1987). 

Invasion History on the East Coast:

Diplosoma listerianum was first collected in 1880 in the Atlantic Ocean off South Carolina, and subsequently found from Cape Fear, NC to Biscayne Bay, FL (U.S. National Museum of Natural History 2002; Van Name 1921; Ruiz et al. unpublished data). We consider it cryptogenic in these southern US waters. The first reported occurrence of D. listerianum north of Cape Hatteras, NC was in Long Island Sound, CT in the 1970s (R. Whitlatch to James T. Carlton, personal communication, 2002). In 1981, it was found at the east end of the Cape Cod Canal, in the Gulf of Maine (R. Whittaker to James T. Carlton, personal communication, 1984). In 1997, it was found in Great Bay, near Portsmouth, New Hampshire (Blezard 1999). In 1998, it was found in Narragansett Bay, Rhode Island and Vineyard Sound, Massachusetts (Whitlatch and Osman 2000). In 1999, it was found near Cape Neddick, Maine (ME) (Harris and Tyrell 2001), and in 2003 it was found in Casco Bay, ME (MIT Sea Grant 2003). It was found in the Gulf of St. Lawrence, in the Magdalene Islands, Quebec in 2008 and in Lunenburg Harbour, Nova Scotia in 2012; however establishment in both locations is uncertain (Willis et al. 2011; Moore et al. 2014).  The northward range expansion fluctuates with the variability of winter weather. and the presence of thermal refugia. It is predicted to colonize most of the Atlantic coast of Nova Scotia by 2075 (Lowen and DiBacco 2023).  There are no published records of this tunicate south of Long Island Sound, CT. However, in 2001 and 2002, specimens were found on fouling plates in lower Chesapeake Bay near Cape Charles, VA. Since there were concerns about identifications, specimens were confirmed as D. listerianum by Gretchen Lambert. Colonies in Cape Charles, Virginia (VA) appeared to be abundant (Ruiz et al. unpublished data).

Invasion History on the Gulf Coast:

Diplosoma listerianum was first found at Cedar Key, FL in 1885 (U.S. National Museum of Natural History 2002; Van Name 1921). It has been collected from Cape Sable, FL to Laguna Madre, Texas (U.S. National Museum of Natural History 2002; Van Name 1921, Lambert et al. 2005; Ruiz et al. unpublished data). We regard it as cryptogenic in the Atlantic Ocean south of Cape Hatteras, NC.

Invasion History in Hawaii:

Diplosoma listerianum was collected in Honolulu, Oahu as early as 1900. It has also been found on Maui and on remote islands, such as Midway Island and Johnston Atoll (Eldredge 1967, Coles et al. 1999; Coles et al. 2001; Carlton and Eldredge 2009).

Invasion History Elsewhere in the World:

Diplosoma listerianum has been reported worldwide from tropical to cold-temperate regions. Among locations where it is considered a recent invader are Guam (1998, Lambert 2002), New Zealand (1946, Cranfield et al. 1998), and South Africa (1st Record 1949, now found from Saldanha Bay to Durban, Monniot et al. 2001; Mead et al. 2011b). In 2018, one specimen was found in a harbor on the southwest coast of Iceland (Ramos-Espla et al. 2020). Genetic studies may result in changes to cryptogenic/introduced status of D. listerianum populations.

We consider Diplosoma listerianum to be broadly cryptogenic in the East Atlantic and tropical continental waters around the world, but introduced to more isolated regions and islands. In the Eastern Pacific, we regard D. listerianum as introduced to the Galapagos Islands (Lambert 2019) and on Cocos Island, Costa Rica, where it was found on fouling plates in 2018 (Ruiz et al., unpublished data).

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Description

Diplosoma listerianum is a colonial ascidian species forming extensive thin, delicate encrusting sheets, rarely more than 2 mm thick and up to 50 mm wide. It can overgrow seagrasses, pilings, crab shells, and other sessile organisms such as corals, gorgonians, and other tunicates. Its tunic is transparent and may have white or yellowish granular bodies suspended in the tunic material. Zooids (individual animals) underneath the tunic may be white, brown, green, or black due to pigment on the abdomen, and sometimes the thoracic epithelium. Other tissues are usually light-colored ranging from white to yellowish or rusty (Van Name 1945; Kott 2001). The colonies have a variable but extensive common cloacal cavity that is subdivided by thin connective tissue in which the zooids are embedded. Each zooid is enclosed by a branch or strip of tunic material (Van Name 1945; Kott 2001). Zooids are small, about 1 mm long. The oral siphon is conspicuous and divided into six deeply divided lobes around the opening. The atrial opening is large and oval, with no atrial tube or languet. This opening has plain edges and exposes most of the branchial sac to the cloacal cavity. There are four rows of stigmata, 10 in the anterior row, and eight in the posterior row. The stomach is rounded and smooth-walled, leading to an intestine of large diameter. Conspicuous stolonic vessels extend from the ventral side of the abdomen where the post-pyloric part of the gut is bent ventrally at right angles to the vertical axis of the zooid. Two oval testis follicles lie against the dorsal side of the gut loop. The vas deferens is hooked around between them, extending anteriorly around the ascending limb of the gut loop. The ovaries and eggs are found behind the intestinal loop (Van Name 1945; Kott 2001). The larvae have a trunk 0.4–6 mm long, with three fixatory papillae (Hayward and Ryland 1990).

A recent study has shown that 'Diplosoma listerianum' consists of at least 4 cryptic species. One of these, clade A, is extremely widespread, occurring in the northeast Atlantic (Plymouth, England; Atlantic and Mediterranean Spain); western Atlantic (Panama); southeast Atlantic-Indian Ocean (South Africa); western Pacific (Japan; Melbourne Australia), and eastern Pacific (Washington, California; Chile) (Perez-Portela et al. 2013). Other clades (B, C, and D) have a more limited distribution, but overlap with clade A. The genetic structure of local populations of clade A is suggestive of anthropogenic dispersal along historic shipping routes but does not indicate a region of origin. The distribution of the clades does not easily correspond with the many historic synonyms (eg., 'D. macdonaldi, D. mitsakurii, etc.). Additional genetic surveys will be needed to determine the identity of populations in many regions of the world.

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Taxonomy

Ecology

General:

Life History- A colonial tunicate consists of many zooids, bearing most or all the organs of a solitary tunicate, but modified to varying degrees for colonial life. Colonial tunicates of the family Didemnidae have small zooids, completely embedded in an encrusting and thin tunic. Each zooid has an oral siphon and an atrial aperture which opens to a shared local chamber. Water is pumped into the oral siphon, through finely meshed ciliated gills on the pharynx, where phytoplankton and detritus are filtered and passed on mucus strings to the stomach and intestines. Excess waste is expelled in the outgoing atrial water (Van Name 1945; Barnes 1983). 
 
Colonial tunicates reproduce both asexually by budding and sexually from fertilized eggs that develop into larvae. Buds can form from the body wall of the zooids. Colonies vary in size ranging from small clusters of zooids to huge spreading masses. The zooids are hermaphroditic, which means both eggs and sperm are released into the atrial chamber. Eggs may be self-fertilized or fertilized by sperm from nearby animals, but some species have a partial block to self-fertilization. Fertilized eggs are brooded within the tunic until they hatch into lecithotrophic (non-feeding, yolk-dependent) tadpole larvae. The larva has a muscular tail and a notochord, eyespots, and a set of adhesive papillae. The larvae are expelled upon hatching and swim briefly before settlement. Swimming periods are usually less than a day, but some larvae settle immediately after release or swim for longer periods if the water temperature is low. On settlement the tail is absorbed, the gill basket expands, and the tunicate begins to feed by filtering (Van Name 1945; Barnes 1983). Populations of D. listerianum in Millport, Scotland had 1–2 generations per year, while those in Plymouth, England had up to 3 generations per year, and could produce larvae in 2–4 weeks after settlement (Millar 1954). 

Food:

Phytoplankton

Consumers:

Competitors:

Trophic Status:

Suspension Feeder

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Habitats

General Habitat	Grass Bed	None
General Habitat	Coarse Woody Debris	None
General Habitat	Oyster Reef	None
General Habitat	Marinas & Docks	None
General Habitat	Rocky	None
General Habitat	Mangroves	None
General Habitat	Vessel Hull	None
General Habitat	Unstructured Bottom	None
General Habitat	Coral reef	None
Salinity Range	Polyhaline	18-30 PSU
Salinity Range	Euhaline	30-40 PSU
Tidal Range	Subtidal	None
Vertical Habitat	Epibenthic	None

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Life History

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	2.2	Field, based on coldest site in geographical range, Boston MA (Zerebecki and Sorte 2011
Maximum Temperature (ºC)	30	Field, based on warmest site in geographical range, Miami FL (Zerebecki and Sorte 2011) Temperature tolerances vary with acclimation and geographical location. For D. listerianum, from Lynn Harbor MA, acclimated at 17 C, the median lethal 24 h temperature (LT50) was 29.41C, but signficantly lower (27.9) for this species from Bodega Bay CA (Sorte et al. 2013).
Minimum Salinity (‰)	18	Typical salinity, Black Sea. In Great Bay NH, D. listerianum occurred at a station where salinity dropped to 24 ppt in spring, but not further upstream (Blezard 1999). A population from Holyhead Harbour, Wales (Irish Sea) showed dramatic mortality when transferred from 34 to 27 and 20 PSU (Gröner et al. 2011).
Maximum Salinity (‰)	40	Field salinity (Shark Bay, Australia, Wyatt et al. 2005)
Minimum Reproductive Temperature	15	Field, larval release, Mediterranean (Brunetti et al. 1988)
Minimum Duration	0	Brunetti et al. 1988 (80% settlement within 24 hours in the light)
Maximum Duration	3	Brunetti et al. 1988 (only 40% settlement with 3 days in the dark)
Broad Temperature Range	None	Cold temperate-Tropical
Broad Salinity Range	None	Polyhaline-Euhaline

General Impacts

Economic Impacts

Fisheries: Diplosoma listerianum has been reported to foul cultured shellfish in the United Kingdom (Isle of Man), Croatia, Japan, and Hong Kong (Ross et al. 2004, Igic 1972, Arakawa 1990, Huang 2003, cited by da Rocha et al. 2009). Da Rocha et al. (2009) suggest that because of the thinness of D. listerianum colonies, its impacts on mussel growth might be small, but this hypothesis still needs to be tested.

Shipping and Industry: In many parts of the world, the colonial tunicate Diplosoma listerianum is an abundant fouling organism (Lambert 2002), known from docks, buoys, floats, and ship hulls (Woods hole Oceanographic Institution 1951).

Ecological Impacts

Competition: In experiments in Long Island Sound, Diplosoma listerianum significantly reduced recruitment of native Spirorbis spp, Bugula spp, and Balanus spp, mostly through overgrowth of newly settled individuals (Osman and Whitlatch 1995). Diplosoma listerianum also significantly reduced recruitment of Botryllus schlosseri and non-indigenous Botrylloides violaceus, mostly through overgrowth of newly settled individuals (Osman and Whitlatch 1995). Disturbance enhanced the spread of D. listerianum (Altman and Whitlatch 2007). Diplosoma listerianum was a strong competitor in utilizing empty space on fouling plates (Stachowicz et al. 2002). In Eel Pond, Woods Hole, Massachusetts, Diplosoma listerianum outgrew Botrylloides violaceus and other organisms on fouling plates, covering ~72% of the plates at the end of the experiment. In cooler water, north of Cape Cod, in Lynn Harbor, Massachusetts Bay, D. listerianum achieved a stand-off with B. violaceus, becoming co-dominant, covering ~37% of the plates at the peak of its growth (Agius 2007). While working in the English Channel, Schmidt and Warner (1986) found that D. listerianum outcompeted Trididemnum tenerum, Botryllus schlosseri and Botrylloides leachii in 39% of trials on fouling plates and reached stand-offs in 61%.  Vance et al. (2008) observed that D. listerianum rapidly overgrew other fouling organisms on plates in newly colonized regions on the North Sea coast of England. Diplosoma listerianum was one of several invasive fouling species which showed increased growth (% coverage) at temperatures 3.5 and 4.5°C above the ambient temperature in Bodega Harbor (13.5°C), while the native Distaplia occidentalis showed reduced survival (Sorte et al. 2010). 

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Regional Impacts

NA-ET3	Cape Cod to Cape Hatteras		Ecological Impact	Competition
In experiments in Long Island Sound, Diplosoma listerianum significantly reduced recruitment of native Spirorbis spp., Bugula spp., and Balanus spp., mostly through overgrowth of newly settled individuals (Osman and Whitlach 1995). Diplosoma listerianum also significantly reduced recruitment of non-indigenous Botryllus schlosseri and Botrylloides violaceus, mostly through overgrowth of newly settled individuals (Osman and Whitlach 1995). Disturbance enhanced the spread of D. listerianum (Altman and Whitlach 2007). Diplosoma listerianum was a strong competitor in filling up empty space on fouling plates (Stachowicz et al. 2002). In experiments in Eel Pond, Woods Hole, Diplosoma listerianum outgrew Botrylloides violaceus and other organisms on fouling plates, covering ~72% of the plates at the peak of its growth (Agius 2007). In years with abnormally warm winters, D. listerianum creates a community which it dominates, having lower biodiversity than native-dominated communities, especially in homogenously developed areas with extensive hard substrate (Munguia et al. 2011).
M040	Long Island Sound		Ecological Impact	Competition
In experiments in Long Island Sound, Diplosoma listerianum significantly reduced recruitment of native Spirorbis spp., Bugula spp., and Balanus spp., mostly through overgrowth of newly settled individuals (Osman and Whitlatch 1995). Diplosoma listerianum also significantly reduced recruitment of non-indigenous Botryllus schlosseri and Botrylloides violaceus, mostly through overgrowth of newly settled individuals (Osman and Whitlatch 1995). Diplosoma listerianum was a strong competitior in filling up empty space on foulng plates (Stachowicz et al. 2002). In years with abnormally warm winters, D. listerianum creates a community which it dominates, having lower biodiversity than native-dominated communities (Munguia et al. 2011).
NEA-II	None		Ecological Impact	Competition
Diplosoma listerianum has been reported to foul cultured shellfish in the United Kingdom (Isle of Man) (da Rocha et al. 2009).  Da Rocha et al. (2009) suggest that because of the thinness of D. listerianum colonies, its impact on mussel growth might be small, but this hypothesis still needs to be tested.  Vance et al. (2008) observed that D. listerianum rapidly overgrew other fouling organisms on plates in newly colonized regions on the North sea coast of England. Schmidt and Warner (1986) found that D. listerianum outcompeted Trididemnurn tenerurn, Botryllus schlosseri and Botrylloides leachii in 39% of trials on fouling plates and reached stand-offs in 61%.
NEA-II	None		Economic Impact	Fisheries
Diplosoma listerianum has been reported to foul cultured shellfish in the United Kingdom (Isle of Man) (da Rocha et al. 2009).  Da Rocha et al. (2009) suggest that because of the thinness of D. listerianum colonies, its impact on mussel growth might be small, but this hypothesis still needs to be tested.
MED-VII	None		Economic Impact	Fisheries
Diplosoma listerianum has been reported to foul cultured shellfish in Croatia (cited by da Rocha et al. 2009).  Da Rocha et al. (2009) suggest that because of the thinness of D. listerianum colonies, its impact on mussel growth might be small, but this hypothesis still needs to be tested.
NWP-3b	None		Economic Impact	Fisheries
Diplosoma listerianum has been reported to foul cultured shellfish in Japan (Arakawa 1990, cited by da Rocha et al. 2009).  Da Rocha et al. (2009) suggest that because of the thinness of D. listerianum colonies, its impact on mussel growth might be small, but this hypothesis still needs to be tested.
NWP-3b	None		Ecological Impact	Competition
Diplosoma listerianum has been reported to foul cultured shellfish in Japan (Arakawa 1990, cited by da Rocha et al. 2009). Da Rocha et al. (2009) suggest that because of the thinness of D. listerianum colonies, its impact on mussel growth might be small, but this hypothesis still needs to be tested.
NWP-2	None		Ecological Impact	Competition
Diplosoma listerianum has been reported to foul cultured shellfish in Hong Kong (Huang 2003, cited by da Rocha et al. 2009).  Da Rocha et al. (2009) suggest that because of the thinness of D. listerianum colonies, its impact on mussel growth might be small, but this hypothesis still needs to be tested.
NWP-2	None		Economic Impact	Fisheries
Diplosoma listerianum has been reported to foul cultured shellfish in Hong Kong (Huang 2003, cited by da Rocha et al. 2009).  Da Rocha et al. (2009) suggest that because of the thinness of D. listerianum colonies, its impact on mussel growth might be small, but this hypothesis still needs to be tested.
M010	Buzzards Bay		Ecological Impact	Competition
In experiments in Eel Pond, Woods Hole, Diplosoma listerianum outgrew Botrylloides violaceus and other organisms on fouling plates, covering ~72% of the plates at the peak of its growth (Agius 2007).
NA-ET2	Bay of Fundy to Cape Cod		Ecological Impact	Competition
In experiments in Lynn Harbor, Massachusetts Bay, Diplosoma listerianum achieved a stand-off with Botrylloides violaceus, becoming co-dominant, covering ~37% of the plates at the peak of its growth (Agius 2007).
NEP-V	Northern California to Mid Channel Islands		Ecological Impact	Competition
Diplosoma listerianum was one of several invasive fouling species which showed increased growth (% coverage) at temperatures 3.5 and 4.5°C above the ambient temperature in Bodega Harbor (13.5°C), while the native Distaplia occidentalis showed reduced survival (Sorte et al. 2010).
P112	_CDA_P112 (Bodega Bay)		Ecological Impact	Competition
Diplosoma listerianum was one of several invasive fouling species which showed increased growth (% coverage) at temperatures 3.5 and 4.5°C above the ambient temperature in Bodega Harbor (13.5°C), while the native Distaplia occidentalis showed reduced survival (Sorte et al. 2010).
N170	Massachusetts Bay		Ecological Impact	Competition
In experiments in Lynn Harbor, Massachusetts Bay, Diplosoma listerianum achieved a stand-off with Botrylloides violaceus, becoming co-dominant, covering ~37% of the plates at the peak of its growth (Agius 2007).
N135	_CDA_N135 (Piscataqua-Salmon Falls)		Ecological Impact	Food/Prey
Increasing abundance of the introduced colonial tunicates Didemnum vexillum and Diplosoma listerianum has resulted in population growth of the native Bloodstar starfish Henricia sanguinolenta (Dijkstra et al. 2012).
NA-ET2	Bay of Fundy to Cape Cod		Ecological Impact	Food/Prey
Increasing abundance of the introduced colonial tunicates Didemnum vexillum and Diplosoma listerianum has resulted in population growth of the native Bloodstar starfish Henricia sanguinolenta (Dijkstra et al. 2012).
N135	_CDA_N135 (Piscataqua-Salmon Falls)		Ecological Impact	Trophic Cascade
Increased abundance of the Bloodstar starfish (Henricia sanguinolenta), supported by growing populations of Didemnum vexillum and Diplosoma listerianum, has resulted in increased predation and near-disapearance of the cryptogenic sponge Halichondria panicea (Dijkstra et al. 2012).
NA-ET2	Bay of Fundy to Cape Cod		Ecological Impact	Trophic Cascade
Increased abundance of the Bloodstar starfish (Henricia sanguinolenta), supported by growing populations of Didemnum vexillum and Diplosoma listerianum, has resulted in increased predation and near-disappearance of the cryptogenic sponge Halichondria panicea (Dijkstra et al. 2012).
CA	California		Ecological Impact	Competition
Diplosoma listerianum was one of several invasive fouling species which showed increased growth (% coverage) at temperatures 3.5 and 4.5°C above the ambient temperature in Bodega Harbor (13.5°C), while the native Distaplia occidentalis showed reduced survival (Sorte et al. 2010)., Diplosoma listerianum was one of several invasive fouling species which showed increased growth (% coverage) at temperatures 3.5 and 4.5°C above the ambient temperature in Bodega Harbor (13.5°C), while the native Distaplia occidentalis showed reduced survival (Sorte et al. 2010).
NH	New Hampshire		Ecological Impact	Food/Prey
Increasing abundance of the introduced colonial tunicates Didemnum vexillum and Diplosoma listerianum has resulted in population growth of the native Bloodstar starfish Henricia sanguinolenta (Dijkstra et al. 2012).
NH	New Hampshire		Ecological Impact	Trophic Cascade
Increased abundance of the Bloodstar starfish (Henricia sanguinolenta), supported by growing populations of Didemnum vexillum and Diplosoma listerianum, has resulted in increased predation and near-disapearance of the cryptogenic sponge Halichondria panicea (Dijkstra et al. 2012).
MA	Massachusetts		Ecological Impact	Competition
In experiments in Eel Pond, Woods Hole, Diplosoma listerianum outgrew Botrylloides violaceus and other organisms on fouling plates, covering ~72% of the plates at the peak of its growth (Agius 2007)., In experiments in Lynn Harbor, Massachusetts Bay, Diplosoma listerianum achieved a stand-off with Botrylloides violaceus, becoming co-dominant, covering ~37% of the plates at the peak of its growth (Agius 2007).

References

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

Abbott, Donald P.; Lambert, Charles C.; Lambert, Gretchen; Newberry, A. Todd (2007) The Light and Smith Manual: Intertidal Invertebrates from Central California to Oregon (4th Edtion), University of California Press, Berkeley, CA. Pp. 949-964

Abbott, Donald P.; Newberry, Andrew Todd (1980) Intertidal invertebrates of California, Stanford University Press, Stanford, California. Pp. 177-227

Agius, Brad P. (2007) Spatial and temporal effects of pre-seeding plates with invasive ascidians: Growth, recruitment and community composition., Journal of Experimental Marine Biology and Ecology 342: 30-39

Airoldi, Laura; Turon, Xavier; Perkol-Finkel, Shimrit; Rius, Marc (2015) Corridors for aliens but not for natives: effects of marine urban sprawl at a regional scale, Diversity and Distributions 21: 755-768

Altman, Safra; Whitlatch, Robert B. (2007) Effects of small-scale disturbance on invasion success in marine communities., Journal of Experimental Marine Biology and Ecology 342: 15-29

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

Baldwin, Andy; Leason, Diane (2016) Potential Ecological impacts of Emerald Ash Borer on Maryland's Eastern Shore, In: None(Eds.) None. , <missing place>. Pp. <missing location>

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

Berrill, N. J. (1932) Ascidians of the Bermudas, Biological Bulletin 62(1): 77-88

Berrill, N. J. (1950) <missing title>, Ray Society, London. Pp. <missing location>

Beshai, Ryan A.; Truong, Danny A.; Henry, Amy K. Sorte, Cascade J. B. (2022) Biotic resistance or invasional meltdown? Diversity reduces invasibility but not exotic dominance in southern California epibenthic communities, Biological Invasions 25(2): 533-549
https://doi.org/10.1007/s10530-022-02932-1

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>

Blezard, David J. (1999) <missing title>, M.S. Thesis, University of New Hampshire, Durham, New Hampshire. Pp. <missing location>

Blum, Julia C.; Chang, Andrew L.; Liljesthröm, Marcela; Schenk, Michelle E. Steinberg, Mia K.; Ruiz, Gregory M. (2007) The non-native solitary ascidian Ciona intestinalis depresses species richness., Journal of Experimental Marine Biology and Ecology 342: 5-14

Bram, Jason; Page, Henry M.; Dugan, Jenifer E. (2005) Spatial and temporal variability in early succession patterns of an invertebrate assemblage at an offshore oil platform., Journal of Experimental Marine Biology and Ecology 317: 223-237

Brewin, Beryl L. (1946) Ascidians in the vicinity of the Portobello Marine Station, Transactions of the Royal Society of New Zealand 76(2): 87-81

Brewin, Beryl L. (1950) Ascidians of New Zealand Part IV. Ascidians in the vicinity of Christchurch, Transactions of the Royal Society of New Zealand 78(2): 344-353

Brunetti, R.; Bressan, M.; Marin, M.; Libralato, M. (1988) On the ecology and biology of Diplosoma listerianum (Ascidiacea, Didemnidae), Vie et Milieu 38(2): 123-131

Bullard, Stephan G.; Whitlatch, Robert B.; Osman, Richard W. (2004) Checking the landing zone: do invertebrate larvae avoid settling near superior spatial competitors?, Marine Ecological Progress Series 280: 239-247

California Department of Fish and Wildlife (2014) Introduced Aquatic Species in California Bays and Harbors, 2011 Survey, California Department of Fish and Wildlife, Sacramento CA. Pp. 1-36

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

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

Carman, M. R.; Morris, J. A.; Karney, R. C.; Grunden, D. W. (2010) An initial assessment of native and invasive tunicates in shellfish aquaculture of the North American east coast, Journal of Applied Ichthyology 26(Suppl. 2): 8-11

Carman, Mary R. and 13 authors (2016) Distribution and diversity of tunicates utilizing eelgrass as substrate in the western North Atlantic between 39° and 47° north latitude (New Jersey to Newfoundland), Management of Biological Invasions Published online: <missing location>

Carman, Mary R. and 7 authors 2010 Ascidians at the Pacific and Atlantic entrances to the Panama Canal. <missing URL>



Carman, Mary R.; Bullard, S.G.; Donnelly, J.P. (2007) Water quality, nitrogen pollution, and ascidian diversity in coastal waters of southern Massachusetts, USA., Journal of Experimental Marine Biology and Ecology 342: 175-178

Carman, Mary R.; Grunden, David W. (2010) First occurrence of the invasive tunicate Didemnum vexillum in eelgrass habitat, Aquatic Invasions 5(1): <missing location>

Carman, Mary, and 8 authors (2011) Ascidians at the Pacific and Atlantic entrances to the Panama Canal, Aquatic Invasions 6(4): 371-380

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

Colarusso, Phil; Nelson, Eric; Ayvazian, Suzanne; Carman, Mary R.; Chintala, Marty; Grabbert, Sinead; Grunden, David (2016) Quantifying the ecological impact of invasive tunicates to shallow coastal water systems, Management of Biological Invasions 7: 33-42

Coles S. L., DeFelice R. C., Eldredge, L. G. (1999a) Nonindigenous marine species introductions in the harbors of the south and west shores of Oahu, Hawaii., Bishop Museum Technical Report 15: 1-212

Coles, S. L.; DeFelice, R. C. : Eldredge, L. G. (2002a) Nonindigenous marine species in Kaneohe Bay, Oahu, Hawai`i, Bishop Museum Technical Report 24: 1-364

Coles, S. L.; DeFelice, R. C.; Eldredge, L. G.; Carlton, J. T. (1999b) Historical and recent introductions of non-indigenous marine species into Pearl Harbor, Oahu, Hawaiian Islands., Marine Biology 135(1): 147-158

Coles, S. L.; DeFelice, R. C.; Minton, D. (2001) Marine species survey of Johnston Atoll, Central Pacific Ocean, June 2000, Bishop Museum Technical Report 19: 1-59

Coles, S. L.; Reath, P. R.; Longenecker, K.; Bolick, Holly; Eldredge, L. G. (2004) <missing title>, Hawai‘i Community Foundation and the U. S. Fish and Wildlife Service, Honolulu. Pp. 1-187

Crane, Laura C.; Goldstein, Jason S.; Thomas, Devin W.; Rexroth, Kayla S.; Watts, Alison W. (2021) Effects of life stage on eDNA detection of the invasive European green crab (Carcinus maenas) in estuarine systems, Ecological Indicators 124(107412): Published online

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>

da Rocha, R. M. (1991) Replacement of the ascidian species in a southeastern Brazilian fouling community., Boletim de Institutode Oceanographia 39(2): 141-153

da Rocha, Rosana M.; Faria, Suzana B.; Moreno, Tatiane R. (2005) Ascidians from Bocas del Toro, Panama, Caribbean Journal of Science 41(3): 600-612

da Rocha, Rosana M.; Kremer, Laura P.; Baptista, Mariah S.; Metri, Rafael (2009) Bivalve cultures provide habitat for exotic tunicates in southern Brazil., Aquatic Invasions 4(1): 195-205

da Rocha, Rosana M.; Kremer, Laura P. (2005) Introduced ascidians in Paranagua Bay, Parana, southern Brazil., Revista Brasileira da Zoologia 22(4): 1170-1184

da Rocha, Rosana M.; Moreno, Tatiane M.; Metri, Rafael (2005) [Ascidians of the Marine Biological Reserve of Avoredro, Santa Catarina, Brazil)., Revista Brasileira de Zoologia 22(2): 461-476

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>

Dias, G. M.; Rocha, R. M.; Lotufo, T. M. C.; Kremer, L. P. (2013) Fifty years of ascidian biodiversity research in Sao Sebastiao, Brazil, Journal of the Marine Biological Association of the United Kingdom 93(1): 273-282

Dijkstra, Jennifer; Harris, Larry G.; Westerman, Erica (2007) Distribution and long-term temporal patterns of four invasive colonial ascidians in the Gulf of Maine, Journal of Experimental Marine Biology and Ecology 342: 61-68

Dijkstra, Jennifer; Sherman,Hillary; Harris, Larry G. (2007) The role of colonial ascidians in altering biodiversity in marine fouling communities., Journal of Experimental Marine Biology and Ecology 342: 169171

Eldredge, L. G. (1966) Taxonomic review of Indo-Pacific didemnid ascidians and descriptions of twenty-three central Pacific species., Micronesica 2: 161-261

Espla, A. A. Ramos; Buencuerpo, V.; Vazquez, E.; Lafargue, F. (1992) Some biogeographical remarks about the ascidian littoral fauna of the straits of Gibraltar (Iberian sector), Bulletin de l'Institut Océanographique 9 (special): 125-131

Faasse, Marco (2012) The exotic isopod Synidotea in the Netherlands and Europe, A Japanese or American invasion (Pancrustacea: Isopoda)?, Nederlandse Faunistiche Mededelingen 108: 103-106

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>

Fisheries and Oceans Canada 2018b Haplosporidium costale (SSO) of Oysters. https://www.dfo-mpo.gc.ca/science/aah-saa/diseases-maladies/hcoy-eng.html



Freeman, Aaren S.; Frischeisen, Alejandro; Blakeslee, April M. H. (2016) Estuarine fouling communities are dominated by nonindigenous species in the presence of an invasive crab, Biological Invasions Published online: <missing location>

Freestone, Amy L.; Ruiz, Gregory M.; Torchin, Mark E. (2013) Stronger biotic resistance in tropics relative to temperate zone: Effects of predation on marine invasion dynamics, Ecology 94(6): 1370-1377

Gaonkar, Chetan A.; Sawant, Subhash S.; Anil, Arga Chandrashekar; Krishnamurthy, Venkat; Harkantra, Sandanand (2010) Changes in the occurrence of hard substratum fauna: A case study from Mumbai Harbour, India, Indian Journal of Marine Science 39(1): 74-84

Gittenberger, Adriaan (2007) Recent population expansions of non-native ascidians in the Netherlands., Journal of Experimental Marine Biology and Ecology 342: 122-126

Gittenberger, Adriaan (2009) Invasive tunicates on Zeeland and Prince Edward Island mussels, and management practices in The Netherlands., Aquatic Invasions 4(1): 279-281

Glasby, T. M. (2001) Development of sessile marine assemblages on fixed versus moving substrata., Marine Ecology Progress Series 215: 37-47

Goodbody, Ivan (2000) Diversity and distribution of ascidians (Tunicata) in the Pelican Cays, Belize., Atoll Research Bulletin 480: 1-33

Goodbody, Ivan (2004) Diversity and distribution of ascidians (Tunicata) at Twin Cays, Belize, Atoll Research Bulletin 524: 1-22

Goodbody, Ivan; Webber, Mona (2003) The ascidian fauna of Port Royal, Jamaica I. Harbor and mangrove-dwelling species., Bulletin of Marine Science 73(2): 457-476

Goodbody, Ivan; Webber, Mona (2003) <missing title>, 3 Environmental Foundation of Jamaica, Kingston. Pp. <missing location>

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

Greene, Charles H.; Schoener, Amy; Corets, Elaine (1983) Succession on marine hard substrata: the adaptive significance of solitary and colonial strategies in temperate fouling communities*, Marine Ecology Progress Series 13: 121-129,

Griffiths, Charles L.; Robinson, Tamara B.; Mead, Angela (2009) Biological Invasions in Marine Ecosystems., Springer-Verlag, Berlin Heidelberg. Pp. <missing location>

Gröner, Frederike; Lenz, Mark; Wahl, Martin; Jenkins, Stuart R. (2011) Stress resistance in two colonial ascidians from the Irish Sea: The recent invader Didemnum vexillum is more tolerant to low salinity than the cosmopolitan Diplosoma listerianum, Journal of Experimental Marine Biology and Ecology 409: 48-52

Harris, Larry G.; Dijkstra, Jennifer A. (2007) <missing title>, New Hampshire Estuaries Project, <missing place>. Pp. <missing location>

Harris, Larry; Tyrrell, Megan (2001) Changing community states in the Gulf of Maine., Biological Invasions 3: 9-21

Haupt, T. M.; Griffiths, C. L.; Robinson, T. B.;Tonin, A. F. G. (2010) Oysters as vectors of marine aliens, with notes on four introduced species associated with oyster farming in South Africa, African Zoology 45: 52-62

Hewitt, Chad L. (1993) <missing title>, Ph.D. Dissertation, University of Oregon, Eugene. Pp. <missing location>

Huang, Xuguang, Bingyu;; Guo, Donghu; Zhong;, Yanping; Li, Shunxing; Liu, Xin;; Laws, Edward A.; Huang, Bangqin (2021) Blackfordia virginica blooms shift the trophic structure to smaller size plankton in subtropical shallow waters, Marine Pollution Bulletin 182(111990): Published online

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

Hutchings, Brenna; Emma Stiles; Erwin, Patrick M () Hurricane events facilitate the dominance of introduced invertebrate species in harbors, Biological Invasions <missing volume>: Published online
https://doi.org/10.1007/s10530-023-03056-w

Ignacio, Barbara L.; Julio, Luciana M.; Junqueira, Andrea O. R; Ferreira-Silva, Maria A. G. (2010) Bioinvasion in a Brazilian Bay: filling gaps in the knowledge of southwestern Atlantic biota, PLOS ONE 5(9): <missing location>

Karlson, Ronald H.; Osman, Richard W (2012) Species composition and geographic distribution of invertebrates in fouling communities along the east coast of the USA: a regional perspective, Marine Ecology Progress Series 458: 255–268

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

Kott, P. (1998) Tunicata, Zoological Catalogue of Australia 34: 51-252

Kott, P. (2005) Catalogue of Tunicata in Australian waters, Queensland Museum, Brisbane. Pp. 1-301

Kott, Patricia (1990) The Australian Ascidiacea, part 2, Aplousobranchia (1), Memoirs of the Queensland Museum 29(1): 1-266

Kott, Patricia (2001) The Australian Ascidiacea, part 4, Aplousobranchia (3), Didemnidae., Memoirs of the Queensland Museum 47(1): 1-407

Koukouras, Athanasios; Voultisiado-Koukoura, Eleni; Kevrekidis, Theodoros; Vafidis, Dimitri (1995) Ascidian fauna of the Aegean Sea with a checklist of the Mediterranean and Black Sea species, Annales de l Institut Oceanographique, Paris 71(1): 19-34

LaFargue, F.; Tursi, A. (1975) Contribution a la connaissance des didemnidae (ascides composees) des cotes des pouilles (mer ionienne et mer adriatique), Vie et Milieu 25(1B): 33-48

Lambert, C. C.; Lambert, G. (1998) Non-indigenous ascidians in southern California harbors and marinas., Marine Biology 130: 675-688

Lambert, Gretchen (1966) The general ecology and growth of a solitary sscidian, Corella willmeriana., Biological Bulletin 135(2): 296-307

Lambert, Gretchen (2002) Nonindigenous ascidians in tropical waters., Pacific Science 56(3): 191-298

Lambert, Gretchen (2003) Marine biodiversity of Guam: the Ascidiacea., Micronesica 35-36: 584-593

Lambert, Gretchen; Faulkes, Zen; Lambert, Charles C.; Scofield, Virginia L. (2005) Ascidians of South Padre Island, Texas, with a key to species., Texas Journal of Science 57(3): 251-262

Lenz, Mark and 11 authors (2011) Non-native marine invertebrates are more tolerant towards environmental stress than taxonomically related native species: Results from a globally replicated study, Environmental Research 111: 943-952

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>

Locke, Andrea (2009) A screening procedure for potential tunicate invaders of Atlantic Canada., Aquatic Invasions 4(1): 71-79

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

Lopez-Legentil, Susanna; Legentil, Miquel L.; Erwin, Patrick M.; Turon, Xavier (2015) Harbor networks as introduction gateways: contrasting distribution patterns of native and introduced ascidians, Biological Invasions 17: 1623-1638
DOI 10.1007/s10530-014-0821-z

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

Lowen, J. B.; DiBacco, C. (2023) Range expansion and establishment of a non-indigenous tunicate (Diplosoma listerianum) in thermal refugia is mediated by environmental variability in changing coastal environments, Canadian Journal of Fisheries and Aquatic Science 80: 330–345
dx.doi.org/10.1139/cjfas-2022-0082

MacIntyre, Chris; Adrienne Pappal; Pederson, Judy; Smith, Jan P. (2011) Marine Invaders in the Northeast Rapid Assessment Survey of non-native and native marine species of floating dock communities, Massachusetts Coastal Zone Management, Boston MA. Pp. <missing location>

Mackie, G. O.; Singla, C. L. (1987) Impulse propagation and contraction in the tunic of a compound ascidian, Biological Bulletin 173: 188-204

Marins, Flavia O.; Novaes, Roberto L. M.; Rocha, Rosana M.; Junquiera, Andrea O. R. (2010) Non indigenous ascidians in port and natural environments in a tropical Brazilian bay, Zoologia 27(2): 213-222

MarLin- Marine Life Information Network 2006-2024 MarLin- Marine Life Information Network. <missing URL>



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

Marshall, J. I.; Rowe, F. W. E.; Fisher; Smith, D. F. (1980) Alterations to the relative species-abundance of ascidians and barnacles in a fouling community due to screens, Australian Journal of Marine and Freshwater Research 31: <missing location>

Massachusetts Office of Coastal Zone Management (2013) Rapid assessment survey of marine species at New England floating docks and rocky shores, Massachusetts Office of Coastal Zone Management, Boston MA. Pp. <missing location>

Mastrototaro, F.; D’Onghia, G.; Tursi, A. (2008) Spatial and seasonal distribution of ascidians in a semi-enclosed basin of the Mediterranean Sea., Journal of the Marine Biological Association of the United Kingdom 88(5): 1053-1061

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

Millar, R. H. (1954) The annual growth and reproductive cycle of four ascidians, Journal of the Marine Biological Association of the United Kingdom 31(1): 41-61

Millar, R.H. (1978) Ascidians from the Guyana shelf., Journal of Sea Research 32(1): 99-106

MIT Sea Grant 2003-2008 Introduced and cryptogenic species of the North Atlantic. <missing URL>



Monniot, C.; Monniot, F. (1997) Records of ascidians from Bahrain, Arabian Gulf with three new species., Journal of Natural History 31: 1623-1643

Monniot, Claude; Monniot, Francoise (1994) Additions to the inventory of Eastern tropical Atlantic Ascidians: arrival of cosmopolitan species., Bulletin of Marine Science 54(1): 71-93

Monniot, Claude; Monniot, Francoise; Griffiths, Charles; Schleyer, Michael (2001) South African Ascidians., Annals of the South African Museum 108(1): 1-141

Monniot, Claude; Monniot, Francoise; Laboutte, Pierre (1985) [Ascidians of the port of Papeete (French Polynesia); Relation to the environment and to intercontinental transport by navigation] (French), Bulletin du Museum National d'Histoire Naturelle. 4e Serie. Section A. Zoologie, Biologie et Ecologie Animales 7(3): 481-495

Monniot, Francoise (2018) Ascidians collected during the Madibenthos expedition in Martinique: 1—Phlebobranchia, Zootaxa 4387(3): 451-472
https://doi.org/10.11646/zootaxa.4387.3.3

Monniot, Francoise; Monniot, Claude (1997) Ascidians collected in Tanzania, Journal of East African Natural History 86: 1-35

Monniot, Francoise; Monniot, Claude (2001) Ascidians from the tropical western Pacific., Zoosystema 23(2): 201-383

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

Moore, Andrea M.; Vercaemer, Bénédikte; DiBacco, Claudio; Sephton, Dawn; Ma, Kevin C. K. (2014) Invading Nova Scotia: first records of Didemnum vexillum Kott, 2002 and four more non-indigenous invertebrates in 2012 and 2013, BioInvasions Records 3(4): 225-234

Moreno-Davila, Betzabe Berenice (2010) <missing title>, Universidad del Mar, campus Puerto Angel, Oaxaca, Puerto Angel, Oaxaca, Mexico. Pp. 116

Munguia, Pablo; Osman, Richard W.; Hamilton, John; Whitlatch, Robert; Zajac, Roman (2011) Changes in habitat heterogeneity alter marine sessile benthic communities 4, Ecological Applications 21(3): 925-935

Murray, Cathryn Clarke, and 5 authors (2014) Spatial distribution of marine invasive species: environmental, demographic and vector drivers, Diversity and Distributions 20: 824-836

Murray, Cathryn Clarke; Pakhomov, Evgeny A.; Therriault, Thomas W. (2011) Recreational boating: a large unregulated vector transporting marine invasive species, Diversity and Distributions 17: 1161-1172

Naranjo, S. A.; Carballo, J. C.; Garcia-Gomez, J. C. (1996) Effects of environmental stress on ascidian populations in Algeciras Bay (southern Spain)., Marine Ecology Progress Series 144: 119-131

Nichols, Claire L.; Lambert, Gretchen; Nydam, Marie L. (2023) Continued persistence of non-native ascidians in Southern California harbors and marinas, Aquatic Invasions 18(1): 1-22
https://doi.org/10.3391/ ai.2023.18.1.101962

Olden, Julian 2013 Rhinogobius brunneus; Amur Goby; Yoshinobori. <missing URL>



Oliveira Filho, R. R.; Lotufo, T. M. C.; 2010 New records of introduced ascidians at Ceara State harbors, Northern Brazil. <missing URL>



Osman, R. W., Whitlatch, R. B. (2000) Ecological interactions of invading ascidians within epifaunal communities of southern New England., In: Pederson, Judith(Eds.) First National Conference on Marine Bioinvasions. , Cambridge MA. Pp. 164-174

Osman, Richard W.; Whitlatch, Robert B. (1995) The influence of resident adults on recruitment: a comparison to settlement., Journal of Experimental Marine Biology and Ecology 190: 169-198

Paulay, Gustav; Kirkendale, Lisa; Lambert, Gretchen; Meyer, Chris (2002) Anthropogenic biotic interchange in a coral reef ecosystem: a case study from Guam, Pacific Science 56(2): 403-422

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

Perez-Portela, R.; Arranz, V.; Rius, M.; Turon, X. (2013) Cryptic speciation or global spread? The case of a cosmopolitan marine invertebrate with limited dispersal capabilities, Scientific Reports 3(1397): 1-10

Pestana, Lueji Barros; Dias, Gustavo Muniz; Marquesa, Antonio Carlos (2017) A century of introductions by coastal sessile marine invertebrates in Angola, South East Atlantic Ocean, Marine Pollution Bulletin 125: 426-a432

Pezy, Jean-Philippe; Pezy, Ambre; Raoux, Aurore (2022) The invasive species Rangia cuneata: A new food source for herring gull (Larus argentatus)?, Ecosphere <missing volume>: ecs2.4058
https://doi.org/10.1002/ecs2.4058

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

Rabalais, Nancy N. (1977) South Texas fauna- a symposium honoring Dr. Allan H. Chaney., Caesar Kleger Wildlife Research Institute, Kingsville Texas. Pp. 65-74

Rajagopal, S.;Nair, K.V.K.; Van Der Velde, G.; Jenner, H.A. (1997) Seasonal settlement and succession of fouling communities in Kalpakkam, east coast of India., Netherlands Journal of Aquatic Ecology 30(4): 309-325

Ramadan, Sh. E.; Kheirallah, A. M.; Abdel-salam, Kh. M. (2006) Marine fouling community in the Eastern harbour of Alexandria, Egypt compared with four decades of previous studies, Mediterranean Marine Science 7/2: 19-29

Rho, Boon Jo; Lee, Ji-Eun (1991) A systematic study of the Ascidians in Korea, Korean Journal of Systematic Zoology 7(2): 195-220

Rho, Boon Jo; Park, Kyung-Sook (1998) Taxonomy of ascidians from Geojedo Island in Korea, Korean Journal of Systematic Zoology 14(3): 173-192

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

Rius, M.; Griffths, C. W. (2011) Alien & Invasive Animals: A South African Perspective, Random House Struik, Johannesburg, South Africa. Pp. 71-75

Robinson, T. B.; Griffiths, C. L.; McQuaid, C. D.; Rius, M. (2005) Marine alien species of South Africa-- status and impacts, African Journal of Marine Science 27(1): 297-306

Rodrigues, S. A.; da Rocha, R. M. (1993) Littoral compound Ascidians (Tunicata) from Sao Sebastiao, Estado de Sao Paulo, Brazil., Proceedings of the Biological Society of Washington 106: 728-739

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>

Ruiz, Gregory; Geller, Jonathan (2021) Spatial and temporal analysis of marine invasions: supplemental studies to evaluate detection through quantitative and molecular methodologies, Marine Invasive Species Program, California Department of Fish and Wildlife, Sacramento CA. Pp. 153 ppl.

Sanamyan, Karen (1999) Ascidians from the North-Western Pacific region: 6. Didemnidae, Ophelia 51(2): 143-161

Sanamyan, Karen; Schories, Dirk (2005) Ascidians from Peru, Spixiana 27(3): 193-197

Schmidt, Gunter H.; Warner, George F. (1986) Spatial competition between colonial ascidians: the importance of stand-off., Marine Ecology Progress Series 31: 101-104

Schwindt, Evangelina and 15 authors (2014) Marine fouling invasions in ports of Patagonia (Argentina) with implications for legislation and monitoring programs, Marine Environmental Research 99: 60-68

Shenkar, N.; Loya, Y. (2008) Aqaba-Eilat- the improbable gulf. Envrionment, biodiversity, and perservation., Magnes Press, Jerusalem. Pp. 197-237

Simkanin, Christina; Fofonoff, Paul W.; Larson, Kriste; Lambert, Gretchen; Dijkstra, Jennifer A.; Ruiz, Gregory M. (2016) Spatial and temporal dynamics of ascidian invasions in the continental United States and Alaska, Marine Biology 163: Published online

Skinner, Luís F.; Barboza, Danielle F.; Rocha, Rosana M. (2016) Rapid Assessment Survey of introduced ascidians in a region with many marinas in the southwest Atlantic Ocean, Brazil, Management of Biological Invasions 7(1): 13-20

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.; Jones, Sierra J.; Miller, Luke P. (2013) Geographic variation in temperature tolerance as an indicator of potential population responses to climate change, Journal of Experimental Marine Biology and Ecology 400: 209-217

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

Stachowicz, John J.; Fried, Heather; Osman, Richard W.; Whitlatch, Robert B. (2002a) Biodiversity, invasion resistence, and marine ecosystem function: reconciling pattern and process., Ecology 83(9): 2575-2590

Stachowicz, John J.; Terwin, Jeffrey R.; Whitlatch, Robert B.; Osman, Richard W. (2002b) Linking climate change and biological invasions: ocean warming facilitates nonindigenous species invasions., Proceedings of the National Academy of Sciences of the U.S.A. 99(24): 15497-15500

Streit, Olivia T; Lambert, Gretchen; Erwin, Patrick M.; Lopez-Legentil, Susanna (2021) Diversity and abundance of native and non-native ascidians in Puerto Rican harbors and marinas, Marine Pollution Bulletin 167(112262): Published online

Swami, B. S.; Chapgar, B. F. (2002) Settlement pattern of ascidians in harbor waters of Mumbai, West Coast of India., Indian Journal of Marine Science 31(3): 207-212

Tamburini, M. ; Ferrario, J.;; Marchini, A.;; Piazza, A.; Lo Vullo; M.; Occhipinti-Ambrogi, A. (2023) Are Fouling Non-indignousspecies invading Porto Venere Bay? An Assessment through image analysis , Biologia Marina Mediterranea <missing volume>(27): 213-216

Tovar-Hernandez, Maria Ana; Dean, Harlan (2014) A new gregarious sabellid worm from the Gulf of California reproducing by spontaneous fission (Polychaeta, Sabellidae), Journal of the Marine Biological Association of the United Kingdom 94(5): 935–946.

Tracy, Brianna M.; Reyns, Nathalie B. (2014) Spatial and temporal patterns of native and invasive ascidian assemblages in a Southern California embayment, Aquatic Invasions 9: In press

Turon, Xavier; Cañete, Juan I.; Sellanes, Javier; Rocha, Rosana M.; López-Legentil, Susanna (2016) Too cold for invasions? Contrasting patterns of native and introduced ascidians in subantarctic and temperate Chile, Management of Biological Invasions 7: 77-86

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



Van Name, Willard G. (1921) Ascidians of the West Indian region and southeastern United States., Bulletin of the American Museum of Natural History 44: 283-494

Van Name, Willard G. (1945) The North and South American ascidians, Bulletin of the American Museum of Natural History 84: 1-462

Vance, Thomas; Lauterbach, Lars; Lenz, Mark;Wahl, Martin; Sanderson, Roy A.; Thomason, Jeremy C. (2008) Rapid invasion and ecological interactions of Diplosoma listerianum in the North Sea, UK., Occasional Papers of the Bishop Museum <missing volume>: 1-5

Vaz-Pinto, F. and 5 authors (2014) Invasion success and development of benthic assemblages: Effect of timing, duration of submersion and substrate type, Marine Environmental Research 94: 72-79

Vazquez, E.; Urgorri, V. (1992) [Fouling ascidians in the inlet of A Grana, Ria de Ferrol, (Galicia, Spain)], Nova Acta Cientifica Compostelana (Bioloxia) 3: 161-167

Vazquez, Lafargue, Elsa; Francoise ; Urgorri, Vicotia (1994) Nuevos datos sobre distribucion de tres especies de la familia Didmenidae (Tunicata, Ascidiacea) de la costas ibericas., Boletin de la Real Sociedad Espaniola de Historia Natural 91(1-4): 219-224.

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

Whitlatch, Robert B.; Osman, Richard (2000) Geographical distributions and organism-habitat associations of shallow water introduced marine fauna in New England., In: Pederson, Judith(Eds.) Marine Bioinvasions. , Cambridge MA. Pp. 61-65

Whitlatch, Robert B.; Osman, Richard W. (2009) Post-settlement predation on ascidian recruits: predator responses to changing prey density., Aquatic Invasions 4(1): 121-131

Whitlatch, Robert B.; Osman, Richard W.; Frese, Annette, Malatesta, Richard, Mitchell, Patricia, Sedgwick, Lynn (1995) The ecology of two introduced marine ascidians and their effects on epifaunal organisms in Long Island Sound, In: Balcom, N. C.(Eds.) Proceedings of the Northeast Conference on Non-Indigenous Aquatic Nuisance Species. , Groton, CT. Pp. 28-29

Willis, Jessica E.; Stewart-Clark, Sarah; Greenwood, Spencer J.; Davidson, Jeff; Quijon, Pedro (2011) A PCR-based assay to facilitate early detection of Diplosoma listerianum in Atlantic Canada, Aquatic Invasions 6: corrected proof

Witman, Jon D.; Smith, Franz (2003) Rapid community change at a tropical upwelling site in the Galapagos Marine Reserve, Biodiversity and Conservation 12: 25-45

Woods Hole Oceanographic Institution, United States Navy Dept. Bureau of Ships (1952) Marine fouling and its prevention., United States Naval Institute., Washington, D.C.. Pp. 165-206

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

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

Zvyagintsev, A. Yu;. Korn, O. M.; Kulikova, V. A. (2004) Seasonal dynamics of pelagic larvae and settling of the fouling organisms in conditions of thermal pollution., Russian Journal of Marine Biology 30(4): 266-277

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