Description
Taxonomy- Tunicates of the genus Botryllus are colonial tunicates, in which groups of 5-20 zooids are organized around shared cloacal (excurrent) openings in a star-like pattern (Van Name 1945). Botryllus 'schlosseri' is a circumglobal form in the temperate zone, highly variable in colony shape and color (Van Name 1945), but whether it constitutes a single species is uncertain. Boyd et al. (1990) found that at Woods Hole MA and Monterey CA, Botryllus were morphologically similar and interfertile, but colonies from the two locations displayed 'allorejection' and did not fuse in the laboratory. They concluded that the Woods Hole and Monterey populations were the same species with slight genetic differences, but cited a number of morphological differences, and a difference in chromosome numbers, between these populations and those in Naples. Boyd et al. have suggested the need for worldwide studies of the species status of Botryllus populations because of their widespread use in embryological and immunological research. Rinkevich et al. (1992) found colony fusion in 4% of Israel vs. Monterey assays, and 12% of Monterey vs. Japan tests, suggesting a high degree of genetic similarity among populations. However, in a later study (Rinkevich et al. 1995), they found no fusion in either Israel vs. Japan assays or in comparisons between different populations along the Israel vs. Japan assays or in comparisons between different populations along the Israeli coast, suggesting a great local polymorphism of allorecognition alleles. The relation of immunological responses to differentiation of other genes and its relation to status of B. schlosseri as a single species (or a species complex) is unclear at this time (Boyd et al. 1990; Rinkevich et al. 1995). A genetic comparison of East Coast (Maine and Woods Hole) vs. West Coast (Bodega Bay to Los Angeles) finds significant differences between populations, suggesting a possible difference in origin for East and West Coast populations (Europe vs. Asia) (Stoner et al. 2002). In another analysis, genotypes of B. schlosseri from Woods Hole and Maine grouped with those from European harbors (Lopez-Legentil 2006). (See comments under 'Taxonomy'). Potentially Misidentified Species - Botrylloides violaceus is a Pacific species, introduced to New England waters in the 1970s (Whitlach et al. 1995), and collected in Chesapeake Bay since 2000 (Ruiz et al. unpublished data).
Taxonomy
Kingdom | Phylum | Class | Order | Family | Genus |
---|---|---|---|---|---|
Animalia | Chordata | Ascidiacea | Phlebobranchia | Botryllidae | Botryllus |
Synonyms
Invasion History
Chesapeake Bay Status
First Record | Population | Range | Introduction | Residency | Source Region | Native Region | Vectors |
---|---|---|---|---|---|---|---|
1962 | Established | Stable | Cryptogenic | Regular Resident | Unknown-Marine | Unknown-Marine | Shipping(Fouling Community) |
History of Spread
Botryllus schlosseri (Golden Star Tunicate) now has a worldwide distribution, including harbors and ports on both sides of the North and South Atlantic (Argentina, South Africa), and both sides of the North and South Pacific (China to Russia, Mexico to Alaska; Australia, New Zealand, Chile)The region of origin of B. schlosseri is unknown, but James T. Carlton (2000, personal communication) suspects the Indo-Pacific as a possible native region. Recent genetic work supports a Mediterranean origin (e.g., Rinkevich et al. 1995; Ben-Shlomo et al. 2001; Ben Shlomo et al. 2006).
Botryllus schlosseri was first reported from North America near Boston MA by Couthouy (1838). It was later recorded from (Brooklyn, Long Island NY; Boston, Salem MA) as B. gouldi by Verrill (1871). Later accounts gave wider ranges: 'Portland, Maine, southward, and is locally abundant from the vicinity of Boston to New Jersey, inclusive. It is also abundant on the west coast of Florida, including the Tortugas, but the only record from intermediate points that has thus far come to notice is the probable one of Pearse, Humm and Wharton....who report a Botryllus sp. from Beaufort, North Carolina (Van Name 1945). 'Boston (or less regularly Bay of Fundy) south at least to Chesapeake Bay....Sporadically south to Gulf of Mexico' (Gosner 1978). The range of B. schlosseri has been extended northwards in recent years, to the Atlantic coast of Nova Scotia and the Gulf of St. Lawrence (Prince Edward Island, reached in 2001, Locke et al. 2007). Van Name (1945) wrote: 'It is, however, very probably an introduced species brought here on the bottom of ships'.
A recent genetic analysis indicates that 'B. schlosseri' is a complex of at least 5 cryptic species (A-E), but only one, clade A, had a widespread distribution in the Northeast and Northwest Atlantic, and Northeast Pacific. The other clades had restricted distributions in Europe (Bock et al. 2012). However, Yund et al. (2015) have identified at least one subclade which appears to be native to the Northwest Atlantic. This subclade is most genetically diverse in that region, and 9 of its 12 haplotypes are unique to that region. The divergence between the prevailing Northwest Atlantic haplotype those from European waters is too great to be accounted for by evolution (Yund et al. 2015). We cannot exclude the occurrence of European genotypes or crytpic species in East Coast water, and so we will treat B. schlosseri as a single cryptogenic species on the East Coast.
Botryllus schlosseri was first collected on the West Coast in 1947, at Mare Island, San Francisco Bay (Carlton 1979). It now occurs in harbors from Ensenada, Mexico (Lambert and Lambert 2003) to Sitka, Alaska (2001, Ruiz et al. unpublished data).
In Chesapeake Bay, B. schlosseri was not reported in the course of an extensive survey of Chesapeake benthos in 1915-1922 (Cowles 1930) or in shoreline surveys near Norfolk by Ferguson et al. (1949). However, it was reported in fouling on the dredge Chinook in Hampton Roads in 1923 (Visscher 1928). It was listed as a rare species in deeper waters of the lower Bay by Wass (1965). A colony was collected at Virginia Institute of Marine Science, Gloucester Point in 1962 (Calder 1972), and it was common to abundant on piers at Norfolk in 1964-1965 (Calder and Brehmer 1967). 'During the drought years of the mid-sixties, B. schlosseri suddenly appeared at Gloucester Point on tray oysters and eventually erupted to cover nearly all tufts of eelgrass (Zostera marina) and (Ruppia (R. maritima , Widgeon Grass) in shallow water in the lower York River. It is a fast-growing pernicious pest on trays in the cool months of spring and fall but barely survives hot summers in Virginia....B. schlosseri was not vigorous in the wet year of 1971, but it was still present on trays of oysters in the spring of 1972. After Agnes it disappeared and no trace has been found at any fouling stations. It may not recover its distribution of the 1960's until another series of droughts occurs' (Andrews 1973). B. schlosseri was abundant on fouling panels in Lynnhaven Bay in 1977 (Otsuka and Dauer 1980) and common on' settling plates (1994-95) in all of the major lower Bay regions sampled except Norfolk (15-18 ppt), and not at all sites. B. schlosseri is much more common on spring-early summer plates than summer-fall plates (Ruiz et al. unpublished data).
The spread of B. schlosseri in the lower Bay in the 1960s is suggestive of a recent introduction, either from Europe or from further north along the East Coast, but this pattern could also represent population fluctuations in response to long-term salinity or temperature changes.
History References - Andrews 1973; Ben-Shlomo et al. 2001; Boyd et al. 1990; Calder 1972; Calder and Brehmer 1967; Carlton 1979; Cowles 1930; Couthouy 1838; Ferguson et al. 1949; Gosner 1978; Gould 1841; Otsuka and Dauer 1980; Rinkevich et al. 1995; 2001; Ruiz et al. unpublished data; Stoner et al. 2001; Verrill 1871; Visscher 1928; Van Name 1945; Wass 1963
Invasion Comments
Ecology
Environmental Tolerances
For Survival | For Reproduction | |||
---|---|---|---|---|
Minimum | Maximum | Minimum | Maximum | |
Temperature (ºC) | -1.0 | 30.0 | 13.0 | 25.0 |
Salinity (‰) | 16.0 | 44.0 | 25.0 | 40.0 |
Oxygen | hypoxic | |||
pH | ||||
Salinity Range | poly-eu |
Age and Growth
Male | Female | |
---|---|---|
Minimum Adult Size (mm) | 75.0 | 75.0 |
Typical Adult Size (mm) | 90.0 | 90.0 |
Maximum Adult Size (mm) | 300.0 | 300.0 |
Maximum Longevity (yrs) | 0.7 | 0.7 |
Typical Longevity (yrs | 0.2 | 0.2 |
Reproduction
Start | Peak | End | |
---|---|---|---|
Reproductive Season | |||
Typical Number of Young Per Reproductive Event |
|||
Sexuality Mode(s) | |||
Mode(s) of Asexual Reproduction |
|||
Fertilization Type(s) | |||
More than One Reproduction Event per Year |
|||
Reproductive Startegy | |||
Egg/Seed Form |
Impacts
Economic Impacts in Chesapeake Bay
Botryllus schlosseri (Golden Star Tunicate) is a common component of fouling communities in the lower Chespeake bay.
Fisheries - Botryllus schlosseri is a 'a fast-growing pernicious pest' on oyster trays, but is absent or rare on natural oyster beds, presumably because of siltation (Andrews 1973).
Industries, Boating - In lower Chesapeake Bay, B. schlosseri varies greatly in abundance from year to year in abundance with salinity and temperature, and is not as important in fouling as Molgula manhattensis (Calder 1966; Andrews 1973).
Economic Impacts Outside of Chesapeake Bay
Botryllus schlosseri (Golden Star Tunicate) is a common fouling organism in temperate harbors and on ships (Milkman 1967; Visscher 1928). It is also important as an experimental organism in embryology, genetics, and immunology (Milkman 1967; Rinkevich et al. 1995).
References- Milkman 1967; Rinkevich et al. 1995; Visscher 1928;
Ecological Impacts on Chesapeake Native Species
Botryllus schlosseri (Golden Star Tunicate) is a common component of fouling communities in the lower Chespeake bay.
Habitat Change - B. schlosseri colonies provide habitat for a wide variety of small motile animals (copepods; amphipods; polychaetes; etc.) (Milkman 1967; Ruiz et al. unpublished data).
Competition - Bancroft (1904) listed hydroids, bryozoans, and 'worms' as competitors for space with B. schlosseri at Woods Hole and Naples. B. schlosseri are abundant and occasionally dominant in spring-early summer fouling communities in the lower Bay (Andrews 1973; Ruiz et al. unpublished data). When fouling plates in Lynnhaven Bay were covered with wire mesh to exclude larger predators, B. schlosseri became dominant during December-March (Otsuka and Dauer 1980). Recruitment of other fouling organisms including native Spirorbis spp. was reduced in the vicinity of B. schlosseri colonies in Long Island Sound (Osman and Whitlach 1995).
Food/Prey- B. schlosseri is eaten by snails and crabs, but Bancroft (1904) considered predation to be much more intense on Naples than Woods Hole populations. Mitrella lunata (Lunate Dove Shell ), a common East Coast fouling community snail was observed to eat B. schlosseri in laboratory cultures (Milkman 1967).
Toxicity - Pellets incorporating ground dried fish were not eaten by green crabs (Carcinus maenas) when treated with B. schlosseri extract, suggesting some anti-feeding properties (Teo and Ryland 1994). However, effects on native crab species are not known, and observations of Bancroft (1904) and others indicate that crabs will eat B. schlosseri on occasion. It has long been believed that some of the unusual chemistry associated with tunicates served, at least in part, to discourage predation, but these effects appear to be weak and variable, at least with the species tested (Teo and Ryland 1994).
References- Andrews 1973; Bancroft 1904; Milkman 1967; Osman and Whitlach 1995; Otsuka and Dauer 1980; Ruiz et al. unpublished data; Teo and Ryland 1994
Ecological Impacts on Other Chesapeake Non-Native Species
Botryllus schlosseri (Golden Star Tunicate) is a common component of fouling communities in the lower Chespeake bay.
Competition - A number of introduced species are competitors for space with B. schlosseri, including Garveia franciscana, Diadumene lineata, and Ficopomatus enigmaticus, but interactions between these species and B. schlosseri have not been documented (Ruiz et al., unpublished data). Recruitment of other introduced fouling organisms (Diplosoma listerianum; Botrylloides spp.) was reduced in the vicinity of B. schlosseri colonies in Long Island Sound (Osman and Whitlach 1995).
Toxicity - B. schlosseri has possible anti-feeding effects on Carcinus maenas (green crabs) (Teo and Ryland 1994).
References- Teo and Ryland 1994
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