Monocorophium acherusicum

Invasion History

First Non-native North American Tidal Record: 1905
First Non-native West Coast Tidal Record: 1905
First Non-native East/Gulf Coast Tidal Record:

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General Invasion History:

Monocorophium acherusicum was originally described from Italy in 1857, but has been so widely dispersed by shipping that its native range is unknown. Some authors consider Europe its probable source region (Bousfield and Hoover 1997), while others consider it native to Eastern North America and introduced in European locations (Chapman 2000). It was collected in Martha’s Vineyard, Massachusetts as early as 1872 (Verrill and Smith 1873, as Corophium cylindricum Say 1818, re-identified as M. acherusicum; Yale Peabody Museum 2014). By 1937, it had also been collected from the Atlantic coasts of France, Holland, the Suez Canal, Senegal, South Africa, Hong Kong, New Zealand, and the West Coast of North America (Shoemaker 1934; Crawford 1937). It is also known from Texas to central Maine (Bousfield 1973; Lecroy 2004), Cuba (Ortiz et al. 2007), Venezuela (Ortiz et al. 2007), Brazil (Bahia to Sao Paulo 8-25°S; Valério-Berardo and Miyagi 2000), Australia (Fearn-Wannan 1968), Hawaii (Shoemaker 1947), Japan (Onbe 1966), the Black Sea and the Mediterranean (Bellan-Santini et al. 1982; Sezgün et al. 2001), the Azores (Chevreux 1900, cited by Lopes et al. 1992), and Spain to the Hebrides in Scotland (Lincoln 1979; Cacabelos et al. 2010). Recent new records from the Gulf of St Lawrence, Quebec (Adebayo et al. 2013); Dublin, Ireland (Daniels et al. 2015); and Helgoland, Germany, in the North Sea (Beerman and Franke 2011) suggest that this amphipod is still extending its range, possibly reflecting climate change and ship transport.

North American Invasion History:

Invasion History on the West Coast:

Monocorphium acherusicum was first collected in the Northeast Pacific as early as 1905 in Yaquina Bay, Oregon; 1915 in Puget Sound, Washington; and 1912 in San Francisco Bay, California. It now ranges from Vancouver Island, British Columbia, Canada to Baja California, Mexico (Carlton 1979; Cohen and Carlton 1995). Early vectors for introduction included transfer with Eastern Oyster transplants and hull fouling (Carlton 1979). This amphipod was first reported from southern California in 1946 (Mohr and Leveque 1948, cited by Carlton 1979), in Los Angeles Long Beach Harbors, where it was very abundant by 1950 (Barnard 1958). It was collected in Newport Bay before 1947 (Shoemaker 1947), San Diego Bay in 1950 (Barnard 1958), and as far south as Bahia de San Quintin, Mexico in 1960 (Barnard 1964, cited by Carlton 1979). It is now known from most of the harbors and estuaries from San Diego to Port Hueneme (Carlton 1979; Cohen et al. 2002). Ballast water and fouling were/are probably important vectors in this region.

Monocorophium acherusicum was first collected in San Francisco Bay in 1912, and now ranges throughout the estuary, into the brackish waters of Suisun Bay and the Delta, where it has been collected as far upstream as Collinsville (Cohen and Carlton 1995; Light et al. 2005). On the central California Coast, it is also known from Morro Bay (Reish and Barnard 1960, cited by Carlton 1979), Elkhorn Slough (Shoemaker 1947; Carlton 1979; Wasson et al. 2001), Bolinas Lagoon (Light 1969, cited by Carlton 1979), Tomales Bay (Johnson and Juskevice 1965, cited by Carlton 1979; 2001, Fairey et al. 2002), and Bodega Harbor (Standing 1975, cited by Carlton 1979; Fairey et al. 2002).

In more northern waters, M. acherusicum was collected in 1905 in Yaquina Bay, Oregon (OR) and occurs in Humboldt Bay, California (1st record 1971; Stout 1971, cited by Carlton 1979; Boyd et al. 2002); Coos Bay, OR (1st record 1942, Barnard 1954, cited by Carlton 1979; Carlton 1989); Tillamook Bay, OR (1st record 1996, Golden et al. 1988), and Willapa Bay, Washington (WA) (1st record 2000, Cohen et al. 2001). Monocorophium acherusicum was found very early in Puget Sound, WA (1st record 1915, USNM specimen, Carlton 1979) and in the Straits of Georgia, British Columbia (1st record 1939, Bousfield and Hoover 1997).

Invasion History in Hawaii:

Monocorophium acherusicum was first reported from Waikiki Bay, Oahu in 1943, and from Pearl Harbor in 1973. It was also collected in three small harbors in Honolulu, Keehi Lagoon, Ala Wai Harbor, and Kewalo Basin in 1997-98 (Coles et al. 1999a; Coles et al. 1999b; Carlton and Eldredge 2009).

Invasion History Elsewhere in the World:

As noted above, M. acherusicum was collected on both sides of the Atlantic in the mid-to-late 19th century, although most American specimens were identified as 'Corophium cylindricum Say 1818'. Some of the equatorial and southern records appear to be spotty, and reflective either of ship transport or limited collecting. However, populations in southern South America and South Africa are considered introduced. It was collected from the Rio de la Plata, Argentina in 1969 (U.S. National Museum of Natural History 2015), and has been collected as far south as Rio Gallegos, Argentina (51.5°S, Schwindt et al. 2014). This amphipod was first reported in South Africa in Durban, on the Indian Ocean, in 1915, but was widespread in most major ports by 1976 (Shoemaker 1947; Griffiths et al. 2009; Mead et al. 2011a; Mead et al. 2011b). By 1947, M. acherusicum was found in Dar es Salaam, Tanzania (Shoremaker 1947). It also occurs in Mauritius (Appado and Myers 2004) and India (Madras, Nayar 1959; Mumbai, Sivaprakasam 1970). It was collected in Hong Kong in 1937 (Crawford 1937; Shoemaker 1947), and by 1955, it was found in Japan at Fukuyama Harbor, in the Seto Inland Sea (1955, Onbe 1966; Doi et al. 2011). This amphipod is now widespread in the Northwest Pacific, in waters of China (Huang 2001), Korea (Hong 1993), and Vladivostok, Russia (1st record 1975, Zvyagintsev 2003). Its current northern limit is the eastern shelf of Sakhalin Island in the Sea of Japan (2002, Zvyagintsev 2003).

In the Southwest Pacific, M. acherusicum was first collected in Lyttelton Harbour, Christchurch, New Zealand in 1881 (Thomson 1881, cited by Hurley 1954). In more recent baseline port surveys, it was found in Tauranga, Gisborne, and Whangarei on the North Island and Timaru, Lyttelton, and Dunedin on the South Island, New Zealand (Inglis et al. 2005). In Australia, M. acherusicum was first discovered at Port Jackson, Sydney, New South Wales (Chilton 1921, cited by Poore and Storey 1999), and at Port Phillip Bay, in Victoria in 1963 (Fearn-Warren 1968; Poore and Storey 1999). It is also present in Western Australia in the Swan River estuary at Bunbury (Poore and Storey 1999) and at Esperance on the southern coast (Hewitt et al. unpublished).

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Description

Monocorophium acherusicum has a slender, depressed body, with small, separated coxal plates. Its urosome segments are fused and it lacks a dorso-lateral ridge. The antennae and the anterior edge of the head are sexually dimorphic in this species. In males, the rostrum is very small and recessed behind the eyes, but much more prominent in females, and projecting ahead of the eyes. In males, the ocular lobes extend forward, and Antenna 2 is much larger and heavier than Antenna 1. While in females, Antenna 2 is only slightly longer, and somewhat more robust than Antenna 1. In males, the first segment (peduncle) of antenna 1 is enlarged at the proximal end, forming a dorsal bump. The inner margin of segment 1 has one small distal spine. The flagellum has 6-8 segments. In females, segment 1 of Antenna 1 is not enlarged, and the inner margin bears 3-4 stout spines, with 4-5 more spines on the ventral margin. The male Antenna 2 has segment 4 enlarged, with its distal end prolonged into a large, curved tooth. Segment 5 has a tooth near the proximal end which nearly opposes the tooth on segment 4. Segments 4 and 5 of the male's Antenna 2 have relatively few setae. In females, segment 4 of antenna 2 bears 3 pairs of stout posterior spines, and a single distal spine. Segments 4 and 5 of the female's Antenna 2 also have more setae than the males. 

The gnathopods are not especially large or conspicuous in this genus. Segment 5 of Gnathopod 1 is longer than segment 6, and the dactyl (segment 7) is longer than the palm of segment 6. On Gnathopod 2, segment 5 is longer than segment 2, and the dactyl bears 2 prominent teeth. Pereiopods 3 and 4 have segment 2 lacking setae, and have long, backward-curving dactyls. Pereiopod 7 is nearly twice as long as Pereiopod 6, and has a long, forward-curving dactyl. As noted above, the urosome segments are fused, without lateral ridges. The peduncle of Uropod 1 has 3 stout spines on the inner margin. The outer ramus of Uropod 2 is shorter than the inner one. Uropod 3 is uniramous, with the ramus shorter than the peduncle. Adults are about 3-6 mm long (Shoemaker 1934a; Bousfield 1973; Lincoln 1979; Bousfield and Hoover 1997; Lecroy 2004; Chapman 2007). The centers of the pleonites and the segments of Antenna 2 are dark brown, mottled with white. There is a patch of pigment between the eyes, but it does not extend to the back of the head (Bousfield 1973). The borders of the pleonites and most of the appendages are white (photo in Barnett 2013). Description based on: Shoemaker 1934a, Bousfield 1973, Lincoln 1979, Bousfield and Hoover 1997, Lecroy 2004, Chapman 2007 and Barnett 2013. 

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Taxonomy

Ecology

General:

Moncorophium acherusicum is a sedentary tube dwelling amphipod, which inhabits soft and hard substrates, as epifauna and infauna. Gammarid amphipods have separate sexes, brooded embryos, and direct development (Bousfield 1973). Females tend to be larger than males, reaching 6 mm, versus 4.5 mm for males (Onbe 1966; Bousfield 1973).  In temperate climates, breeding is seasonal, e.g. May-September in New England (Bousfield 1973), or year-round, but more frequent, and starting at smaller female body-sizes in summer (2.2 mm) and winter (3.0 mm) in Fukuyama Harbor, Japan (Onbe 1966). The brood size of females is related to body size, with 2-4 eggs for the smallest females, and 60-70 eggs for an ~6 mm female (Onbe 1966).

Moncorophium acherusicum ranges from cold-temperate to tropical climates, and tolerates ice-covered winter conditions and temperatures as high as 30C (Lee et al. 2005). In experiments, it tolerated salinities as low as 6 PSU (Onbe 1966). In San Francisco Bay, it was found in the Delta and Suisun Bay during drought periods at 5-15 PSU, but was most abundant in San Pablo Bay, and the South and Central Bays at 25-30 PSU (Cohen and Carlton 1995; Lee et al. 2003; Peterson and Vaysierres 2010). Monocorophium acherusicum secretes threads of 'amphipod silk', to which the sediment detritus is attached, to form its tubes. When the amphipods are abundant, the tubes form a mass in which the openings point outward or upward (Barnard et al. 1988). The tubes can be formed on the sediment surface, or attached to fouling on vertical surfaces such as rocks or pilings, or other fouling organisms (Barnard 1958). Monocorophium acherusicum is a sedentary tube dweller much of the time, but does swim and occurs in the zooplankton, especially at night, or following disturbance by storms and river runoff (Grabe 1996). 'Unlike other tubicolous animals, the amphipods are not obligatorily sessile, but move in and out of the tubes in search of food and to mate. Migration rates are high among the tubicolous amphipods, as evidence by their early appearance on fresh blocks (experimental substrates)' (Barnard 1958). Reported substrates of M. acherusicum include: rocks, pilings, pontoons, floats, buoys, ropes of set-nets, ships' hulls, oyster shells, Eelgrass (Zostera marina, Z. japonica), hydroids, macroalgae, and sponges (Crawford 1937; Barnard 1958; Onbe 1966; Fearn-Wannan 1968; Long 1968; Marsh 1972; Bellan-Santini et al. 1982; Berkenbusch and Rowden 2007).

When feeding, Moncorophium acherusicum sit at the mouths of their tubes, waving their antennae, capturing phytoplankton and organic detritus (Barnard 1958). We have not found studies of the feeding of M. acherusicum, but like other corophiid amphipods, it probably is capable of feeding on detritus and benthic microalgae on the sediment surface, and grazing on filamentous epiphytic algae growing on seaweeds and seagrasses (Bousfield 1973). Fishes appear to be the major predators on M. acherusicum. In a mesocosm experiment in the Fraser River Delta, British Columbia, the exclusion of fishes led to a dramatic increase in this amphipod's abundance (Amundred et al. 2015).

Food:

Phytoplankton; Detritus

Consumers:

Fishes, crabs, shrimps

Competitors:

Other filter-feeding amphipods

Trophic Status:

Suspension Feeder

SusFed

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Habitats

General Habitat	Grass Bed	None
General Habitat	Coarse Woody Debris	None
General Habitat	Unstructured Bottom	None
General Habitat	Oyster Reef	None
General Habitat	Marinas & Docks	None
General Habitat	Vessel Hull	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	Epibenthic	None
Vertical Habitat	Endobenthic	None

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	0	Based on range (Bousfield 1973)
Maximum Temperature (ºC)	30	Experimental (Lee et al. 2005, highest tested), Korea
Minimum Salinity (‰)	6	Experimental (Onbe 1966)
Maximum Salinity (‰)	40	Experimental (Lee et al. 2005, highest tested), Korea
Minimum Length (mm)	1.3	Adult males, 2.2 for females (Onbe 1966, Japan)
Maximum Length (mm)	6	Adult females, 4.5 for males (Onbe 1966, Japan)
Broad Temperature Range	None	Cold temperate-Tropical
Broad Salinity Range	None	Mesohaline-Euhaline

General Impacts

Monocorophium acherusicum is a widespread fouling organism, building masses of tubes, constructed of sediment and detritus, on ships' hulls, docks, and aquaculture structures (Crawford 1937; Shoemaker 1947; Woods Hole Oceanographic Institution 1952; Barnard 1958; Onbe 1966). It is also abundant in soft substrates of bays and estuaries, and is probably an important grazer of phytoplankton and benthic and epiphytic microalgae, as well as an important prey item for fishes. Monocorophium acherusicum's tube-building is a form of ecological engineering. It is often an early colonizer of fresh pilings, creating habitat which is used by other organisms (Barnard 1958).

Economic Impacts

Monocorophium acherusicum can foul pilings, covering them with masses of tubes covered with sediment, which is probably unattractive in marinas. However, the fouling perhaps provides a benefit by discouraging boring organisms such as shipworms and gribbles (Barnard 1958). In Japan, fouling by M. acherusicum affected the culture of seaweeds and oysters (Onbe 1966).

Ecological impacts

Monocorophium acherusicum's tubes on man-made and natural surfaces provide a major habitat alteration, probably facilitating settlement of some organisms, but discouraging others. Barnard (1958) described a pattern of succession in Los Angeles Harbor, in which corophiid amphipods, dominated by M. acherusicum, settled first on new pilings, followed by polychaetes and tunicates. However, Onbe (1966) found that few other organisms settled on surfaces heavily covered with M. acherusicum's tubes. As an opportunistic suspension/deposit feeder, M. acherusicum could affect the abundance of phytplankton in estuaries (Nichols and Thompson 1985a). High densities of corophiid amphipods could also affect the stability of sediments and the degree of erosion in estuaries (Talman et al. 1999).

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Regional Impacts

NWP-3b	None		Economic Impact	Fisheries
The growth of cultured seaweed (Undaria pinnatifida was reduced by being fouled by the tubes of M. acherusicum and Erichthonius brasiliensis. The tubes were also found on the shells of cultivated edible and pearl oysters (Onbe 1966).
NWP-3b	None		Ecological Impact	Competition
Onbe (1966) observed that no significant fouling by other organisms occurred on buoys which were completely covered by M. acherusicum's tubes.
NWP-3b	None		Ecological Impact	Habitat Change
Onbe (1966) observed that no significant fouling by other organisms occurred on buoys which were completely covered by M. acherusicum's tubes.
NEP-VI	Pt. Conception to Southern Baja California		Economic Impact	Shipping/Boating
Monocorophium acherusicum fouls pilings, covering them with masses of tubes covered with sediment, but perhaps a benefit by discouraging boring organisms (Barnard 1958).
NEP-VI	Pt. Conception to Southern Baja California		Ecological Impact	Habitat Change
Dense masses of corophiid tubes may discourage settlement by boring organisms and by other foulers, such as the tunicate Ciona intestinalis (Barnard 1958). They also may provide habitat for other organisms, such as predatory polychaetes (Barnard 1958).
AUS-VIII	None		Ecological Impact	Habitat Change
Talman et al. (1999, suggested that the very high densities of corophiids (presumably introduced) observed in Port Phillip Bay could decrease the stability of sediment, resulting in increased erosion.
P050	San Pedro Bay		Economic Impact	Shipping/Boating
Fouling pilings, covering them with masses of tubes covered with sediment, but perhaps a benefit by discourage boring organisms (Barnard 1958).
P050	San Pedro Bay		Ecological Impact	Habitat Change
Dense masses of corophiid tubes may discourage settlement by boring organisms and by other foulers, such as the tunicate Ciona intestinalis (Barnard 1958). They also may provide habitat for other organisms, such as predatory polychaetes (Barnard 1958).
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 M. acherusicum, was responsible for a decline in phytoplankton biomass in Suisun Bay during 1976-1977, a dry period of high salinity (Nichols and Thompson 1985).
P090	San Francisco Bay		Ecological Impact	Herbivory
Nichols and Thompson (1985) suggested that an upstream movement of abundant suspension-feeding benthos, including M. acherusicum, was responsible for a decline in phytoplankton biomass in Suisun Bay during 1976-1977, a dry period of high salinity (Nichols and Thompson 1985).
CA	California		Ecological Impact	Habitat Change
Dense masses of corophiid tubes may discourage settlement by boring organisms and by other foulers, such as the tunicate Ciona intestinalis (Barnard 1958). They also may provide habitat for other organisms, such as predatory polychaetes (Barnard 1958).
CA	California		Ecological Impact	Herbivory
Nichols and Thompson (1985) suggested that an upstream movement of abundant suspension-feeding benthos, including M. acherusicum, was responsible for a decline in phytoplankton biomass in Suisun Bay during 1976-1977, a dry period of high salinity (Nichols and Thompson 1985)., Nichols and Thompson (1985) suggested that an upstream movement of abundant suspension-feeding benthos, including M. acherusicum, was responsible for a decline in phytoplankton biomass in Suisun Bay during 1976-1977, a dry period of high salinity (Nichols and Thompson 1985).
CA	California		Economic Impact	Shipping/Boating
Fouling pilings, covering them with masses of tubes covered with sediment, but perhaps a benefit by discourage boring organisms (Barnard 1958).

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