Streblospio benedicti

Invasion History

First Non-native North American Tidal Record: 1932
First Non-native West Coast Tidal Record: 1932
First Non-native East/Gulf Coast Tidal Record:

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General Invasion History:

The spionid polychaete Streblospio benedicti is native to the Western Atlantic, from the Gulf of St. Lawrence to Venezuela (Carlton 1979; Balza et al. 2013). It occurs in soft sediments, marshes, and in fouling communities in estuaries. It is tolerant of pollution and common in disturbed areas receiving heavy nutrient inputs (Levin 1984a; Levin 1984b). Streblospio benedicti has been introduced to the West Coast, from Mexico to British Columbia (Hobson 1976; Carlton 1979; Ferrando and Mendez 2010), and Europe from Poland to Spain (da Fonseca-Genevois and Cazaux 1987; DAISIE 2009; Kocheshkova and Matviy 2009).

North American Invasion History:

Invasion History on the West Coast:

Streblospio benedicti was first collected from the West Coast in 1932 in Berkeley, California in San Francisco Bay (1932, Hartman 1936, cited by Carlton 1979). Within the Bay, it is abundant and ranges through the Central and South Basins and inland through San Pablo Bay to Antioch on Suisun Bay (Light 1977; Carlton 1979; Nichols and Thompson 1985; Cohen and Carlton 1995; Cohen et al. 2005). It is one of the most abundant organisms in shallow soft sediments of the Bay, reaching densities of 10,000-50,000 individuals per m² (Cohen and Carlton 1995). South of San Francisco Bay, this worm was reported from Elkhorn Slough in 1975 (Blake and Woodwick 1975) and Morro Bay in 1966 (Blake 1966, cited by Carlton 1979). In the Los Angeles area, it was first collected from Alamitos Bay in 1952 (Reish and Winter 1954, cited by Carlton 1979) and subsequently from Venice, on Santa Monica Bay (Woodwick 1953, cited by Carlton 1979), Anaheim Bay-Huntington Harbor (Hill and Reish 1954, cited by Carlton 1979), Newport Bay (Reish 1954, cited by Carlton 1979), and Los Angeles-Long Beach Harbors in 1973 (Hill and Reish 1975, cited by Carlton 1979). Streblospio benedicti was present in Mission Bay and the Tijuana River estuary, San Diego by 1982 (Levin 1984a; Levin 1984b) and subsequently found in San Diego Bay in 1998 (Ranasinghe et al. 2005) and Los Penasquitos Lagoon (Desmond et al. 2002). This worm is also found in Mexico, in the Topolobampo Lagoon, on the Gulf of California (Mendez-Ubach 1997), and the Urias estuary near Mazatlan (Ferrando and Mendez 2010).

North of San Francisco Bay, S. benedicti was present in Tomales Bay and Bodega Harbor by 1936 (Hartman 1936, cited by Carlton 1979), and found in Bolinas Lagoon by 1968 (Light 1969, cited by Carlton 1979). Further north, the temporal pattern of spread is spotty. In 1974, the worm was collected in Yaquina Bay, Oregon (Walker 1974, cited by Carlton 1979) and Grays Harbor, Washington (Hobson 1976, cited by Carlton 1979). Dates for intervening estuaries are later: Humboldt Bay, California in 1992 (Boyd et al. 2002); Coos Bay, Oregon in 1997 (Jeffries 1977, cited by Carlton 1979); Umpqua River Estuary, Reedsport, Oregon in 1983 (Carlton 1989); Tillamook Bay, Oregon in 1996 (Golden et al. 1998, cited by Cohen 2004); Columbia River Estuary in 1990 (Sytsma et al. 2004); and Willapa Bay in 2000 (Cohen et al. 2001). By the late 1990s, it was known from Puget Sound (Cohen et al. 1998), the San Juan Islands (Klinger et al. 1998), and the Fraser River Delta, British Columbia (Environment Canada 1994).

Genetic studies indicate that San Francisco Bay, Tijuana River, and Padilla Bay, Washington populations are closely related to East Coast populations (Galveston Bay, Texas; Bogue Sound, North Carolina; and Georgetown, South Carolina) which also have a lecithotrophic mode of development (Schulze et al. 2000; Pernet and McArthur 2006). In another genetic comparison of three West Coast and 11 East Coast populations, the West Coast populations were most similar to lecithotrohpic animals from Delaware (Zakas and Wares 2012). These studies suggest that oysters were probably not the vector of introduction, since most Eastern Oyster transplants came from Long Island Sound (Carlton 1979; Miller 2000; Miller et al. 2007). The late discovery of S. benedicti from Humboldt Bay to British Columbia, many decades after Eastern Oyster transplants, also suggests that other vectors were involved, probably transport of Streblospio by shipping from central or southern California (Schulze et al. 2000; Pernet and McArthur 2006; Zakas and Wares 2012).

Invasion History in Hawaii:

Streblospio benedicti was first found on Oahu in Halawa Stream, Pearl Harbor in 1977 (Ward 1981, cited by Carlton and Eldredge 2009) and has been found in several streams in Oahu (Englund et al. 2000).

Invasion History Elsewhere in the World:

In the northeast Atlantic, S. benedicti was first reported from the Shannon Estuary, Ireland in 1979 (O'Connor and O'Sullivan 1981) and the Loire Estuary, near Nantes, France in 1982 (da Fonseca-Genevois and Cazaux 1987). It has also been found in the Marennes-Oleron Bay in Brittany (de Montaudouin and Sauriau 2000), and the estuaries of La Arena, Plentzia and Gernika (Bay of Biscay) in the Basque country, Spain (in 1991, Garcia-Arberas and Rallo 2004). By 1999, it was found in German North Sea waters in the Ems Estuary (Nehring and Leuchs 1999) and the Wadden Sea (Sebesvari et al. 2006). In the mid-1990s, S. benedicti was found in the Baltic Sea, in a Russian portion of the Vistula Lagoon (Kocheshkova and Matviy 2009). The introduced S. benedicti in the German Wadden Sea have planktotrophic larval development, indicating origination from a different source than that of the introduced populations on the West Coast of North America (Sebesvari et al. 2006).

Reports of introduced Streblospio from the Mediterranean, Black and Caspian Seas refer to S. gynobranchiata (Cinar et al. 2005; Boltacheva 2008; Taheri et al. 2008), native to the Gulf of Mexico. Populations of Streblospio sp. resembling S. benedicti have been found in Japan (S. b. japonica; Imajima 1990) and in Brazil (S. cf. benedicti; Mendes and Soares-Gomes 2013). The Japanese population showed some morphological similarities both to S. benedicti and S. shrubsolii. Its taxonomic position is uncertain (Rice and Levin 1998).

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Description

Streblospio benedicti is a small, inconspicuous, tube-dwelling spionid polychaete. Its prostomium has four dark eyespots. The anterior end bears a pair of long, translucent, colorless palps, with a single pair of branchiae (gills) adjacent to the palps. The palps are heavily ciliated, with crenulated (scalloped) margins. The 2nd segment bears a broad collar. Chaetiger 1 has a very short pair of parapodia, whose notopodia have 1 or 2 longer chaetae, and neuropodia have five shorter setae, while chaetiger 2 has a raised dorsal collar. The posterior segments have longer, more numerous chaetae. Hooks are present from segment 7 back and alternate with pointed setae. The hooks have 4-5 teeth at the distal end. The adult worms are 5-10 mm long. The body is translucent reddish brown. The branchiae are crossed by transverse bars of dark green pigment. They inhabit soft, grey, mucoid tubes attached to surfaces or embedded in sediment (Hartman 1945; Light 1978; Levin and Creed 1986; Lippson and Lippson 1997; Blake and Ruff, in Carlton 2007). Eggs and larvae are brooded in dorsal pouches (Levin 1984; Schulze et al. 2000).

Different populations have different modes of larval development. Some populations (Massachusetts, Rhode Island, and Florida) have planktotrophic larval development (Levin 1984b ) which is illustrated by Dean (1965). Others have lecithotrophic larvae (California, Texas), while populations in North and South Carolina have both. The two groups of larvae differ in size, number of chaetae, and duration in plankton (Levin 1984a; Levin 1984b; Schulze et al. 2000). Reproductive mode appears to be genetically determined, but involves minimal genetic differences, and so these two forms are considered conspecific (Schulze et al. 2000; Zakas and Wares 2012). A form of Streblospio from the Gulf of Mexico, having a third mode of development, brooding planktotrophic larvae in the gills (branchiae) is now considered a separate species (S. gynobranchiata) (Schulze et al. 2000).

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Taxonomy

Ecology

General:

Streblospio benedicti is a small, opportunistic, tube-building worm found in the upper 4 cm of mudflats and soft-sediments in estuaries and coastal waters. Sexes are separate and maturity is rapid at around 9-14 weeks (Levin 1986; Levin and Bridges 1994). Populations and individuals vary in developmental mode (lecithotrophic and planktotrophic populations) (Levin 1984a; Levin 1984b). Some populations have exclusively planktotrophic development, with smaller eggs (60-70 µm), less developed larvae (3-7 chaetigers), and a longer planktonic period (2-7 weeks), compared to those with lecithotrophic larvae (100-200 µm eggs, 8-10 chaetiger larvae, <8 days in the plankton) (Levin and Bridges 1994, reared at 20°C). These developmental differences, called poecilogony, do not seem to be influenced by the maternal condition, the environment and genetic isolation (Zakas and Rockman 2015). While the lecithotrophic larvae derive most of their nutrition from yolk, they are capable of feeding on and developing more rapidly in the presence of phytoplankton (Pernet and McArthur 2005).

Most Atlantic Coast populations are planktotrophic, while all West Coast populations are lecithotrophic (Levin 1984a; Levin 1984b; Pernet and McArthur 2005; Zakas and Wares 2012). Some Atlantic Coast populations contain both lecithotrophic and planktotrophic individuals. These individuals are identical morphologically and in DNA sequences at mitochondrial loci. The mode of development is genetically determined, with a polymorphism maintained in these populations by a balance of trade-offs (Levin 1991; Zakas and Wares 2012). West Coast populations were most similar to a lecithotrophic population from Delaware (Zakas and Wares 2012). Planktotrophic larvae settle at 8-12 chaetigers, after 5 to 40 days in the plankton (Dean 1965; Sebesvari et al. 2006). Lecithotrophic larvae settle at a similar size, 9-12 chaetigers, but they reach that size in the brood pouch and can settle immediately after release, but usually within seven days of release (Levin 1984a; Levin 1984b). Settlement is stimulated by the presence of sediment containing organic material (Sebesvari et al. 2006). Settled worms build vertical mucoid tubes in soft sediment, and can be found in oyster beds, or in fouling communities, where sediment can accumulate among shells or in crevices (Wells 1961; Llanso et al. 2011).

The broad range of S. benedicti indicates a wide temperature tolerance (Levin and Creed 1986; Schulze 2000; Zakas and Wares 2012). This polychaete also tolerates a wide range of salinity and has been found in salinities as low as 4 PSU in the Baltic Sea (Kocheshkova and Matviy 2009) and as high as ~45 PSU in the Salton Sea, California (Detwiler et al. 2002). Streblospio benedicti is tolerant of hypoxia and survived for two weeks at 7% of O2 saturation (0.5 mg/l) (Llanso 1991). They tolerate high organic contents and settle in new habitats as a pioneer organism (Garcia-Arberas and Rallo 2004).

Worms of the genus Streblospio use their long, ciliated palps to remove detritus and algae from the sediment surface and transport it to their mouths (Dauer et al. 2003). They are also capable of using their long, ciliated palps to remove particles either from the sediment surface or the water column, and so can function both as a suspension and a deposit feeder (Nichols and Thompson 1985). They play an important part in estuarine food webs, despite their small size, because they can reach very high population densities (Nicols and Thompson 1985; Kocheshkova and Matviy 2009). Predators include shrimps, crabs, and small fishes (Posey and Hines 1991; Lippson and Lippson 1997).

Food:

Phytoplankton; detritus

Consumers:

Shrimp, Crabs, Fishes

Trophic Status:

Deposit Suspension Feeder

DepSusFed

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Habitats

General Habitat	Unstructured Bottom	None
General Habitat	Oyster Reef	None
General Habitat	Marinas & Docks	None
General Habitat	Mangroves	None
General Habitat	Grass Bed	None
Salinity Range	Mesohaline	5-18 PSU
Salinity Range	Polyhaline	18-30 PSU
Salinity Range	Euhaline	30-40 PSU
Tidal Range	Subtidal	None
Tidal Range	Low Intertidal	None
Vertical Habitat	Endobenthic	None
Vertical Habitat	Epibenthic	None

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	-2	Based on geographical distribution
Maximum Temperature (ºC)	34.5	Field, Terminos Lagoon, Mexico (Cruz-Abrego et al. 1994)
Minimum Salinity (‰)	4	Field, 4-7 PSU, Kocheshkova and Matviy 2010, Vistula Lagoon, Baltic Sea.
Maximum Salinity (‰)	45	Salton Sea (g/L). Note that the composition of this lake's salts differs from seawater (Detwiler et al. 2002).
Minimum Dissolved Oxygen (mg/l)	0.5	Experimental at 26 C (Llanso 1991)
Minimum Reproductive Temperature	7.5	Experimental, animals from Bogue Sound NC. Reproduction reduced below 12.5 C (Levin and Creed 1986)
Maximum Reproductive Temperature	30	Reproduction reduced above 25 C C (Levin and Creed 1986)
Minimum Duration	0	Lecithotrophic larvae are competent to settle upon release (Levin 1984a; Levin 1984b).
Maximum Duration	40	Planktotrophic larvae, in glass tubes, without substrate, 20 C (Levin 1984a; Levin 1984b).
Minimum Length (mm)	5	Hartman 1945
Maximum Length (mm)	9.7	Levin and Creed 1986
Broad Temperature Range	None	Cold temperate-Tropical
Broad Salinity Range	None	Mesohaline-Euhaline

General Impacts

Ecological Impacts

The spionid polychaete Streblospio benedicti can attain very high abundances, especially in the shallow, fine sediments of estuaries. It is probably a significant grazer of phytoplankton in native and invaded estuaries, and may have effects on sediment properties. Competition with other suspension and deposit feeding invertebrates (polychaetes, amphipods, mollusks) is likely. However, these impacts have not been studied experimentally in invaded estuaries. In European waters, it is also a potential competitor with the native S. shrubsolii (Fonseca-Genevois and Cazaux 1987).

Herbivory- Nichols and Thompson (1985) suggested that an upstream movement of abundant suspension-feeding benthos, including S. benedicti, was responsible for a decline in phytoplankton biomass in Suisun Bay, California during a period of low river flow and high salinity in 1976-1977. Da Fonseca-Genevois and Cazaux (1987) found that phytoplankton comprised a significant portion of the stomach contents of S. benedicti in the Loire Estuary, France.

Competition- In the Loire estuary, France, S. benedicti largely replaced the native S. shrubsolii, after the dredging of a shipping channel. However, the precise mechanism of competition was not studied (da Fonseca-Genevois and Cazaux 1987).

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Regional Impacts

NEP-V	Northern California to Mid Channel Islands		Ecological Impact	Herbivory
Nichols and Thompson (1985) suggested that an upstream movement of abundant suspension-feeding benthos, including S. benedicti, was responsible for a decline in phytoplankton biomass in Suisun Bay during a period of low river flow and high salinity 1976-1977.
P090	San Francisco Bay		Ecological Impact	Herbivory
Nichols and Thompson (1985) suggested that an upstream movement of abundant suspension-feeding benthos, including S. benedicti, was responsible for a decline in phytoplankton biomass in Suisun Bay during a period of low river flow and high salinity 1976-1977.
NEA-IV	None		Ecological Impact	Competition
In the Loire estuary, France, the native Streblospio shrubsolii was largely replaced by S. benedicti. The species replacement was attributed to S. benedicti's greater tolerance to the stress, disturbance, and sediment changes caused by the dredging of shipping channels (Fonseca-Genevois and Cazaux 1987).
CA	California		Ecological Impact	Herbivory
Nichols and Thompson (1985) suggested that an upstream movement of abundant suspension-feeding benthos, including S. benedicti, was responsible for a decline in phytoplankton biomass in Suisun Bay during a period of low river flow and high salinity 1976-1977., Nichols and Thompson (1985) suggested that an upstream movement of abundant suspension-feeding benthos, including S. benedicti, was responsible for a decline in phytoplankton biomass in Suisun Bay during a period of low river flow and high salinity 1976-1977.

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