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Potentially Misidentified Species:
Colonial Botryllus schlosseri is similar in appearance to other colonial
tunicates such as those belonging to the genus Botrylloides. The systems
(zooid clusters) of Botrylloides species tend to be elongate in shape,
often arranged in meandering double rows, but only occasionally exhibiting the
star pattern typical of B. schlosseri (Cohen 2005).
II. HABITAT AND DISTRIBUTION
Regional Occurrence:
Botryllus schlosseri was introduced from Europe and is now found on both coasts
of North America as well as Australia and the Asian Pacific (NIMPIS 2002).
On the Atlantic coast of North America, it is occurs from Maine (occasionally
north to the Bay of Fundy) south to Florida and the Gulf of Mexico (Cohen
2005). Occurrence down the Atlantic seaboard may be disjunct, possibly due to
discontinuity of hard substrates, but collection records indicate its presence
in New Jersey and the North Carolina (NIMPIS 2002, Hiscock 2007, Salem Sound
Coastwatch undated).
IRL Distribution:
Mook (1993) identified Botryllus schlosseri as established and seasonally abundant in the IRL (Carlton and Ruckelshaus 1997).
III. LIFE HISTORY AND POPULATION BIOLOGY
Age, Size, Lifespan:
Individual Botryllus schlosseri zooids are usually 2.5-5 mm in length, system
clusters around 5-10 mm in diameter and colonies are typically 10 cm or less
across (Cohen 2005).
Chadwick-Furman and Weissman (1995) reported that colonies in the field
populations they studied had lifespans ranging from an average of 82-247 days
and that colony life expectancy was seasonally dependent. This is in sharp
contrast to colonies that have been maintained in the laboratory for more than
7 years (Rinkevich and Shapira 1998).
B. schlosseri colonies grew exponentially as juveniles attaining colony
sizes of up to 1400 zooids within 69 days (Chadwick-Furman and Weissman 1995).
Abundance:
Carlton and Ruckelshaus (1997) indicate that Botryllus schlosseri is seasonally
common in Florida fouling communities, as is the co-occurring non-native
ascidian Styela plicata and the cryptogenic (native range uncertain)
B. niger (= Botrylloides nigrum).
Reproduction:
Reproduction in B. schlosseri, a sessile hermaphrodite, includes a
sexual and an asexual component.
Sexual reproduction involves the release of male gametes into the water
followed by uptake in the incurrent siphons of nearby colonies and interanal
fertilization of eggs (Phillippi et al. 2004). Asexual reproduction involves a
synchronized budding process that occurs on an approximately weekly basis to
increase the size of the clonal colony until it becomes large enough to
reproduce sexually. Closely genetically related colonies may fuse with one
another when they come into contact, resulting in larger "chimera" (made up of genetically distinct individuals) colonies
that may reach sexual maturity more rapidly than smaller unfused colonies.
Sexual maturity in field populations in Monterey, CA was attained in 49 days,
corresponding to 7 asexual replication cycles (Chadwick-Furman and Weissman
1995, Salem Sound Coastwatch undated).
Reproductive seasonality appears quite variable across the broad distribution
range of the species.
Embryology:
Internal gestation and development to a free-swimming "tadpole" larval stage
is followed by release to the water column and a short (up to 24-36 hours)
planktonic duration that is probably capable of only local (1-10 km) dispersal
(Berrill 1950, 1975, Hiscock 2007).
IV. PHYSICAL TOLERANCES
Temperature:
Broad latitudinal distribution of Botryllus schlosseri (from Florida north to
the Bay of Fundy on the east coast of North America, for example), indicates a
eurythermal condition. Sexual reproduction is likely to be seasonally
restricted to a brief summer season at higher latitudes. Brunetti et al.
(1980) report a lower reproductive temperature of 11ºC for the European
population examined by these authors.
Salinity:
Hiscock (2007) indicates Botryllus schlosseri is relatively euryhaline, tolerating salinities ranging from 18-40 ppt.
V. COMMUNITY ECOLOGY
Trophic Mode:
Botryllus schlosseri is a suspension feeder whose diet includes
suspended phytoplankton, zooplankton and suspended organic matter (Millar 1971,
NIMPIS 2002).
Associated Species:
Various invertebrate species such as flatworms, crustaceans, and gastropods
have been reported to feed on Botryllus schlosseri colonies (Cohen
2005).
VI. INVASION INFORMATION
Invasion History:
The European native range of Botryllus schlosseri is believed to encompass the
Mediterranean, and may also include the Adriatic and Black seas, Great Britain
and France, Norway, and the Faroe Islands (Cohen 2005, Salem Sound Coastwatch
undated).
Ruiz et al. (2000) indicate that the first records of B. schlosseri on
the east coast of North America date to 1841 in Massachusetts, while the
earliest reported occurrence in the Gulf of Mexico appears to be 1921. On the
U.S. west coast, the earliest reports of B. schlosseri include reports
from San Francisco Bay dating to the mid 1940s, San Diego Bay and Mission Bay
dating to the early 1960s, and from a Puget Sound oyster farm in the late 1960s
or early 1970s. Broader distribution up and down the Pacific coast of North
America from British Columbia to Mexico was only documented starting in the
mid-1990s (Lambert and Lambert 1998, Cohen 2005).
The species has been broadly introduced elsewhere as well, and can now be found
in Australia (since 1905), Tasmania and New Zealand (since 1928), Japan, and
Hong Kong. Ship hull fouling is probably the most common introduction vector
for B. schlosseri, but accidental introduction in shipments of live
oysters or other wild-harvested or cultured organisms has probably also played
a role in expanding the range of this organism (Cohen 2005).
Potential to Compete With
Natives:
B. schlosseri competes with other attached benthic filter feeders for
space and possibly food as well. Fast-growing B. schlosseri colonies
may overgrow neighboring organisms and alter community dynamics (NIMPIS 2002,
Cohen 2005). Harms and Anger (1983) report barnacles and mussels as among the
most important space competitors with B. schlosseri.
Schmidt and Warner (1986) examined spatial competition among B. schlosseri and
three other encrusting colonial ascidian species and reported no significant
overgrowth among competing species.
Possible Economic Consequences of Invasion:
An abundant harbor fouling organism, Botryllus schlosseri is a nuisance species
that fouls boat hulls, marine equipment, aquaculture gear, and other submerged
structures. It can also overgrow and compete for space with cultured oysters
and mussels (NIMPIS 2002, Cohen 2005).
VII.
REFERENCES
Berrill N.J. 1950. The Tunicata with an account of the British species. The Ray
Society, London. 354 p.
Berril N.J. 1975. Chordata: Tunicata. pp. 241-282 In: (Geise A.C, and J.S.
Pearse, eds). Reproduction of Marine Invertebrates, vol. II,. Academic Press,
NY.
Brunetti R., Beghi L., Bressan M., and M.G. Marin. 1980. Combined effects of
temperature and salinity on colonies of Botryllus schlosseri and
Botrylloides leachi (Ascidiacea) from the Venetian Lagoon. Marine
Ecology Progress Series 1980:202-314.
Carlton J.T. and M.H. Ruckelshaus. 1997. Nonindigenous marine invertebrates and
algae. Pp 187-201 in: Simberloff D., Schmitz D.C., and T.C. Brown (eds).
Strangers in Paradise. Island Press, Washington, D.C. 467 p.
Chadwick-Furman N.E., and I.L. Weissman. 1995. Life history plasticity in
chimaeras of the colonial ascidian Botryllus schlosseri. Biological
Sciences 262:157-162.
Cohen A.N. 2005 Guide to the Exotic Species of San Francisco Bay. San Francisco
Estuary Institute, Oakland, CA. Available online.
Harms J., and K. Anger. 1983. Seasonal, annual, and spatial variation in the
development of hard bottom communities. Helgolaender Meeresuntersuchungen
36:137-150.
Hiscock K. 2007. Botryllus schlosseri. Star ascidian. Marine Life
Information Network: Biology and Sensitivity Key Information Sub-programme.
Plymouth: Marine Biological Association of the United Kingdom. Available online.
Lambert C.C., and G. Lambert. 1998. Non-indigenous ascidians in southern
California harbors and marinas. Marine Biology 130:675-688.
Millar R.H. 1971. The biology of ascidians. Advances in Marine Biology 9:1-100.
Mook D. 1983. Responses of common fouling organisms in the Indian River,
Florida, to various predation and disturbance intensities. Estuaries 6:372-379.
NIMPIS. 2002. Botryllus schlosseri species summary. CSIRO National
Introduced Marine Pest Information System (Hewitt C.L., Martin R.B., Sliwa C.,
McEnnulty, F.R., Murphy, N.E., Jones T. and S. Cooper Eds). Available online.
Phillippi A., Hamann E., and P.O. Yund. 2004. Fertilization in an egg-brooding
colonial ascidian does not vary with population density. Biological Bulletin
206:152-160.
Rinkevich B., and M. Shapira. 1998. An improved diet for inland broodstock and
the establishment of an inbred line from Brotryllus schlosseri, a
colonial sea squirt (Ascidiacea). Aquatic Living Resources 11:163-171.
Ruiz G.M., Fofonoff P.W., carlton J.T., Wonham M.J., and A.H. Hines. 2000.
Invasion of coastal marine communities on North America: Apparent patterns,
processes, and biases. Annual Review of Ecological Systematics 31:481-531.
Salem Sound Coastwatch. Undated. Guide to marine invaders in the Gulf of Maine:
Botryllus schlosseri fact sheet. 2p.
Schmidt G.H., and G.F. Warner. 1986. Spatial competition between colonial
ascidians: the importance of stand-off. Marine Ecology Progress Series
31:101-104.
Report by:
J. Masterson, Smithsonian Marine Station
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