Arcuatula senhousia

Invasion History

First Non-native North American Tidal Record: 1941
First Non-native West Coast Tidal Record: 1941
First Non-native East/Gulf Coast Tidal Record:

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General Invasion History:

Musculista senhousia is native from the southern Kurile Islands, Sakhalin, Russia, and the north coast of Hokkaido, Japan (Golikov et al. 1976) to Singapore (Willan 1987). It is also established in the Indian Ocean from Thailand to the Red Sea, and the coast of east Africa. The extent to which it is native in the Indian Ocean is unknown (Hewitt 2002; Academy of Natural Sciences of Philadelphia; Bachelet et al. 2009). This mussel has been widely introduced with Japanese oysters, ship fouling, and ballast water, and has invaded the northwest Pacific (from Mexico to British Columbia), the southwest Pacific (Australia, New Zealand), the northeast Atlantic (Mediterranean and Atlantic France), and the tropical west Atlantic (Venezuela) (Carlton 1979; Willan 1987; Zenentos et al. 2003; Bachelet et al. 2009). Asif and Krug (2012) found that different lineages were involved in invasions in different parts of the world. A cool-water lineage, found in northern South Korea predominated in Puget Sound, Tomales Bay, San Diego and the Mediterranean, while in other parts of Southern California and New Zealand, a warm-water lineage, found in Tokyo, was the predominant form (Asif and Krug 2012).

North American Invasion History:

Invasion History on the West Coast:

Musculista senhousia was first collected in northeast Pacific waters in Samish Bay, Washington (WA) in 1924 'on planted beds of oysters', but was not found later at this site (Kincaid 1947, cited by Carlton 1979). Established populations were found at Olympia, WA, in Puget Sound in 1959 (Rice 1971, cited by Carlton 1979) and a number of subsequent populations have been reported throughout the Sound (Carlton 1979; USGS Nonindigenous Species Program 2009). In the 1990s, M. senhousia was collected in Georgia Straits, British Columbia, and now ranges north to Barkley and Desolation Sounds on Vancouver Island (2006, Gillespie et al. 2007). It was introduced to Willapa Bay, WA with planted clams (Venerupis phillipinarum) but did not persist (Cohen et al. 2001). Populations are not yet known from bays or estuaries in Oregon.

In Northern California, M. senhousia was reported as 'rare' by one survey (in 2001, Fairey et al. 2002) in Humboldt Bay, but this identification has not been confirmed and this species was not mentioned by another study of exotic species (Boyd et al. 2002). In central California, M. senhousia was first collected in Tomales Bay in 1941 (Pitelka and Paulson 1942, cited by Carlton 1979). In 1946, it was collected in Bolinas Lagoon, where it did not become established, and in San Francisco Bay, where it became abundant and widespread (Smith 1947, cited by Carlton 1979). Within San Francisco Bay, M. senhousia is found in South, Central, and San Pablo bays, and upstream to Honker Bay (Carlton 1979; Nichols and Thompson 1985; Cohen and Carlton 1995; Cohen et al. 2005). In 1971, it was collected in Bodega Harbor, and became established (Carlton 1979). However, in Elkhorn Slough, where it was reported in 1965, it did not persist (McDonald 1969; Carlton 1979; Wasson et al. 2001). In southern California, M. senhousia was first collected in 1965 in Mission Bay (McDonald 1969; Carlton 1979), in 1976 in San Diego Bay and in Newport Bay in 1997 (Carlton 1979). In surveys in 1998-2001, it was found in other estuaries, including the Tijuana River estuary, Oceanside Harbor, Huntington Harbor, Alamitos Bay, Los Angeles-Long Beach Harbors, Rainbow Lagoon, Marina del Rey, and Port Hueneme (Cohen et al. 2002; Fairey et al. 2002; deRivera et al. 2005; Ranasinghe et al. 2005). Currently, this mussel occurs southward to the Estero de Punta Banda, Baja California, Mexico (Cohen 2005).

Invasion History on the East Coast:

This mussel was found in fouling on ships arriving in Halifax Harbor, Nova Scotia, Canada in 2007-2009 (Sylvester et al. 2011).

Invasion History Elsewhere in the World:

In 1964, M. senhousia was discovered in the eastern Mediterranean Sea, at Tel Aviv, Israel and later the Bardawil Lagoon, Egypt. In 1978, this mussel appeared in lagoons, such as the Etang de Thau, on the French Mediterranean coast, where Pacific Oysters were cultured (Zenetos et al. 2003). Subsequently, M. senhousia has appeared in other parts of the Mediterranean, including the Adriatic (Ravenna Lagoon in 1986, Zenetos et al. 2003); Po River Delta (in 1994, Mistri et al. 2002); Lagoon of Venice (in 2002, Museo di Storia Naturale di Venezia 2013), and the Gulf of Taranto (in 2001, Mastrotaro and Dappiano 2005). In 2002, it was discovered in Arcachon Bay, on the Biscay coast of France, where it now occurs in several lagoons on the France/Spain border (Bachelet et al. 2009). The introductions to the Eastern Mediterranean may have occurred through the Suez Canal, but western Mediterranean and Biscay introductions are closely associated with Japanese oyster culture (Zenetos et al. 2003; Bachelet et al. 2009).

In the Southwest Pacific, M. senhousia was first collected in Waitemata Harbour, North Island, New Zealand, in 1980, and subsequently found in several other harbors on the North Island, sometimes in very high abundances (Willan 1987). In 1982, the mussel colonized the Swan River Estuary, near Perth, Western Australia (Willan 1987). Although this population reached very high densities, it disappeared completely in 2000, after a very heavy rain, forming an extensive freshwater lens, accompanied by a toxic algae bloom (McDonald and Wells 2010). In the late 1980s, M. senhousia appeared in Port Phillip and Westernport Bays, Victoria (Boyd 1999), in St. Vincent Gulf, South Australia, and in the Tamar estuary, Tasmania (1995, Hewitt 2002).

In 2002, M. senhousia was discovered for the first time in the Western Atlantic, in the Caribbean Sea of Delta Amacuro State, Venezuela. Specimens were growing on mangroves in Boca Pedernales, Gulf of Paria and the Orinoco Delta (2002, Martinez-Escarbassiro et al. 2004; Perez et al. 2007).

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Description

Arcuatula senhousia (Senhouse Mussel) is rather fragile, smooth, and thin-shelled, 2ith a rounded triangular shape. The two valves are equal. The beak is near the anterior end, but not terminal. There are concentric lines in the middle of the shell, while the posterior and anterior ends have fine radiating striae. The lunule (depressed area near the beak) has a crenulated margin. There are also crenulations along and behind the ligament. The exterior of the shell is green to bluish green, and the periostracum is shiny. Adult mussels are ~10-30 mm in size. When settling on soft surfaces, this mussel creates "bags" from byssal threads which enclose the mussel and whole colonies, creating mats of mussels and byssus. (Description from: Abbott 1974; Willan 1987; Coan et al. 2000; Zenetos et al. 2003; Coan and Valentich-Scott, in Carlton 2007)

Larvae are described by Semenikhina et al. (2008). They settled at a size of about 280 µm after 18 days in the plankton at 25-30⁰C. At 15⁰C, they remained in the plankton for 60 days but did not settle (Kimura and Sekiguchi 1996).

The name Musculista senhousia is still widely used, but Arcuatula senhousia is recommended by WoRMS (World Registry of Marine Species, Appeltans et al. 2020).

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Taxonomy

Ecology

General:

Arcuatula senhousia has separate sexes, as other mussels do (Mistri 2002). Animals mature in about one year, and die in their second year (Creese et al. 1997; Mistri 2002). Eggs are brooded, but sperm are released into the water column. A single female may release as many as 137,000 eggs (Sgro et al. 2002). Fertilized eggs develop into a planktonic trochophore larva, then into a shelled veliger. The larvae settle at 280 µm (Semenikhina et al. 2008) after about 2-6 weeks in the plankton (Kimura and Sekiguchi 1996). In San Diego County, California larvae dispersed over a mean distance of 15 km, with much genetic exchange between Mission and San Diego Bays, but also much local recruitment within bays (Lopez-Duarte et al. 2012), as determined by genetic and trace-metal analysis.

When settling on soft surfaces, this mussel creates bags of byssal threads which enclose the mussel and whole colonies, creating mats, which can provide a major element of structure in mudflats (Willan 1987; Crooks 1998; Buschbaum et al. 2010). When settling on hard surfaces, A. senhousia does not create mats, and occurs in lower densities (Willan 1987). Like other mussels, A. senhousia is a suspension feeder, and filters phytoplankton and other suspended particles out of the water column. Arcuatula senhousia tolerates a wide range of temperatures and can tolerate salinities over the whole polyhaline-euhaline range (17-27 PSU) (Mistri 2002; Cohen 2005; Galil 2006). It is intolerant of prolonged hypoxia (Mistri 2002).

Food:

Phytoplankton; Detritus

Consumers:

crabs, fishes, spiny lobsters, birds, snails

Trophic Status:

Suspension Feeder

SusFed

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Habitats

General Habitat	Unstructured Bottom	None
General Habitat	Salt-brackish marsh	None
General Habitat	Marinas & Docks	None
General Habitat	Rocky	None
General Habitat	Oyster Reef	None
General Habitat	Vessel Hull	None
Salinity Range	Polyhaline	18-30 PSU
Salinity Range	Euhaline	30-40 PSU
Tidal Range	Subtidal	None
Tidal Range	Low Intertidal	None
Tidal Range	Mid Intertidal	None
Vertical Habitat	Epibenthic	None

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	5	(Mistri 2002, Italy)
Maximum Temperature (ºC)	31	(Mistri 2002, Italy)
Minimum Salinity (‰)	17	Field data, San Francisco Bay (Cohen 2005)
Maximum Salinity (‰)	37	Field data, San Francisco Bay (Cohen 2005)
Minimum Reproductive Temperature	25	Development incomplete at 15 C, no intermediate temperatures tested (Kimura and Sekiguchi 1996)
Maximum Reproductive Temperature	30	Highest tested (Kimura and Sekiguchi 1996)
Minimum Duration	18	25 & 30 C- Kimura and Sekiguchi 1996
Maximum Duration	42	Uchida 1965, cited by Mistri 2002
Minimum Length (mm)	14	Mistri 2002, age at maturity, Italy
Maximum Length (mm)	32	Mistri 2002, age at maturity, Italy
Broad Temperature Range	None	Cold temperate-Tropical
Broad Salinity Range	None	Polyhaline-Euhaline

General Impacts

Ecological Impacts

Competition- The invasion of Musculista senhousia in Mission Bay, California; the Po River Delta, Italy; and New Zealand estuaries has resulted in the decline, and sometimes disappearance of some suspension feeding bivalves and serpulid polychaetes. This species reaches very high densities and biomasses on mudflats, and presumably competes with other suspension-feeders (Willan 1987; Creese et al. 1997; Crooks 2001; Mistri 2003; Hayward et al. 2008). The mats provide structure for some organisms, providing protection from predation and sites of attachment.

Habitat Change- In its native and invaded ranges, M. senhousia creates large mats of mussels surrounded by byssus threads and embedded in the sediment, increasing shear strength and stability of the sediment, and creating a new habitat which can be colonized by some organisms, but can have negative impacts on others. The mats provide protection from predation, and also concentrate organic matter favoring some deposit feeders (Buschbaum et al. 2010; Crooks 1998; Crooks and Khim 1999; Mistri 2003; Munari et al. 2008). In brackish water, the shells of Musculista favored foraminifera communities by stabilizing pH (Hayward et al. 2008). Negative impacts may include the alteration of sediment structure and texture, interfering with burrowing, or chemical effects on sediment due to deposition of organic compounds (Munari et al. 2008). Impacts on eelgrass (Zostera marina) were mixed, with decreased rhizome growth, but increased leaf growth due to increased sediment ammonium (Dexter and Crooks 2000).

Food/Prey- The development of large biomasses of mussels in intertidal mudflats provide a major food resource for predators, including fishes, invertebrates, and birds. In Mission Bay, California, abundances of M. senhousia decline in a seaward direction, due largely to predation by spiny lobsters and murex snails (Cheng and Hovel 2011). However, effects on predator populations are not known.

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Regional Impacts

NZ-IV	None		Ecological Impact	Competition
Arcuatula senhousia replaced most of the native bivalve fauna in the Tamaki Estuary, North Island, New Zealand (Hayward et al. 2008). Its invasion resulted in the reduction of infauna diversity, particularly bivalves, under mussel mats (Creese et al. 1997). Arcuatula senhousia replaced a subtidal scallop bed. It appears to be competing with, and reducing the abundance of the native mussel Xenostrobus pulex (Willan 1987).
NZ-IV	None		Ecological Impact	Habitat Change
At lower salinities, shells buffer pH changes resulting from land clearance, causing persistence of some foraminifera in mussel sites. Sites invaded by mussels showed changes in ostracod species composition (Hayward et al. 2008).
NEP-VI	Pt. Conception to Southern Baja California		Ecological Impact	Competition
The invasion of Arcutatuls senhousia was accompanied by the disappearance of the native bivalve Chione fluctifraga. In experiments, survival and growth of C. fluctifraga and C. undatella was reduced in the presence of A. senhousia's mats. Competition for food with M. senhousia may be a factor in the decline of these suspension-feeding clams (Crooks 2001). In experiments in Mission and San Diego Bays, settling young of M. senhousia out-compete native bivalves (Chione undatella and Laevicardium substriatum) for space, by overgrowing them (Castorani and Hovel 2015).
NEP-VI	Pt. Conception to Southern Baja California		Ecological Impact	Habitat Change
Arcuatula senhousia's mats cover large areas of subtidal flats in Mission Bay, San Diego, California. These mats increase the shear strength and stability of the sediment, favoring increased densities of other benthic organisms in the mats, including the tanaid, Leptochelia dubia, and the gastropod Barleia subtenus. The mats provide structure for tube-building organisms such as L. dubia. The mats also concentrate organic matter, favoring deposit-feeding organisms. (Crooks 1998; Crooks and Khim 1999). Negative impacts of A. senhousia are not obvious or numerous, but negative correlations were seen with the deposit-feeding bivalve Solen rostriformis, and the polychaetes Euchone limnicola and Diplocirrus sp. (Dexter and Crooks 2000). Alteration of sediments by A. senhousia may be a factor in the decline of Chione fluctifraga and C. undatella (Crooks 2001). In San Diego Bay, A. senhousia had both negative and positive effects on the eelgrass Zostera marina, in experiments, reducing rhizome growth, but increasing sediment ammonium and enhancing leaf growth (Dexter and Williams 1998).
P030	Mission Bay		Ecological Impact	Competition
The invasion of Arcuatula senhousia was accompanied by the disappearance of the native bivalve Chione fluctifraga. In experiments, survival and growth of C. fluctifraga and C. undatella was reduced in the presence of A. senhousia's mats. Competition for food with M. senhousia may be a factor in the decline of these suspension-feeding clams (Crooks 2001). In experiments in Mission and San Diego Bays, settling young of A. senhousia out-compete native bivalves (Chione undatella and Laevicardium substriatum) for space, by overgrowing them (Castorani and Hovel 2015).
P030	Mission Bay		Ecological Impact	Habitat Change
Arcuatula senhousia's mats cover large areas of subtidal flats in Mission Bay, San Diego, California. These mats increase the shear strength and stability of the sediment, favoring increased densities of other benthic organisms in the mats, including the tanaid, Leptochelia dubia, and the gastropod Barleia subtenus. The mats provide structure for tube-building organisms such as L. dubia. The mats also concentrate organic matter, favoring deposit-feeding organisms (Crooks 1998; Crooks and Khim 1999). Negative impacts of A. senhousia are not obvious or numerous, but negative correlations were seen with the deposit-feeding bivalve Solen rostriformis, and the polychaetes Euchone limnicola and Diplocirrus sp. (Dexter and Crooks 2000). Alteration of sediments by A. senhousia may be a factor in the decline of Chione fluctifraga and C. undatella (Crooks 2001).
P020	San Diego Bay		Ecological Impact	Habitat Change
In San Diego Bay, A. senhousia had both negative and positive effects on the eelgrass Zostera marina in experiments, reducing rhizome growth, but increasing sediment ammonium and enhancing leaf growth (Dexter and Williams 1998).
MED-VII	None		Ecological Impact	Competition
In the Sacco di Goro, in the Po River delta, the invasion of A. senhousia was accompanied by the decline of the exotic suspension-feeders Ficopomatus enigmaticus, Hydroides dianthus, and the native Mytilaster minimus, presumably due to competition for suspended food particles (Mistri 2003).
MED-VII	None		Ecological Impact	Habitat Change
In the Sacco di Goro, in the Po River delta, the invasion of M\A. senhousia was accompanied by the increase of deposit-feeding taxa (Neanthes succinea, Streblospio shrubsolei, and Microdeutopus gryllotalpa), probably favored by changes in the sediment (Mistri 2003). Patches of mussels of moderate density, had higher abundances, biomasses and species richness than areas without mussels, but at extremely high mussel densities, benthic diversity decreased (Munari et al. 2008).
MED-III	None		Ecological Impact	Habitat Change
In the Rio di Padrongiano Delta, Tyrhennian Sea (Munari et al. 2008) patches of mussels of moderate density, had higher abundances, biomasses and species richness than areas without mussels, but at extremely high mussel densities, benthic diversity decreased (Munari et al. 2008).
P030	Mission Bay		Ecological Impact	Food/Prey
Arcuatula senhousia populations in Mission Bay decline in a seaward direction, as a result of predation by Spiny Lobsters (Panulirus interruptus) and murex snails (Pteropurpura festiva) (Cheng and Hovel 2011). However, effects on predator populations are not known.
NEP-VI	Pt. Conception to Southern Baja California		Ecological Impact	Food/Prey
Arcuatula senhousia populations in Mission Bay decline in a seaward direction, as a result of predation by Spiny Lobsters (Panulirus interruptus) and murex snails (Pteropurpura festiva) (Cheng and Hovel 2011). However, effects on predator populations are not known.
NEP-VI	Pt. Conception to Southern Baja California		Ecological Impact	Trophic Cascade
One factor in the replacement of native bivalves by Arcuatula senhousia is that the heavy settlement of the mussel attract native predators (Castorani and Hovel 2015)
P030	Mission Bay		Ecological Impact	Trophic Cascade
One factor in the replacement of native bivalves by Arcuatula senhousia is that the heavy settlement of the mussel attract native predators(Castorani and Hovel 2015)
P020	San Diego Bay		Ecological Impact	Competition
In experiments in Mission and San Diego Bays, settling young of A. senhousia out-compete native bivalves (Chione undatella and Laevicardium substriatum) for space, by overgrowing them (Castorani and Hovel 2015).
P020	San Diego Bay		Ecological Impact	Trophic Cascade
ne factor in the replacement of native bivalves by Arcuatula senhousia is that the heavy settlement of the mussel attract native perdators(Castorani and Hovel 2015)
NZ-IV	None		Ecological Impact	Toxic
In 2009, dogs in New Zealand died after consuming a native sea-slug, Pleurobranchaea maculata. The slugs contained high levels of tetradotoxin (TTX), accumulated from predation on Arculatula senhousia, which had been feeding on toxic phytoplankton (Taylor et al. 2015).
CA	California		Ecological Impact	Competition
In experiments in Mission and San Diego Bays, settling young of A. senhousia out-compete native bivalves (Chione undatella and Laevicardium substriatum) for space, by overgrowing them (Castorani and Hovel 2015)., The invasion of Arcuatula senhousia was accompanied by the disappearance of the native bivalve Chione fluctifraga. In experiments, survival and growth of C. fluctifraga and C. undatella was reduced in the presence of A. senhousia's mats. Competition for food with M. senhousia may be a factor in the decline of these suspension-feeding clams (Crooks 2001). In experiments in Mission and San Diego Bays, settling young of A. senhousia out-compete native bivalves (Chione undatella and Laevicardium substriatum) for space, by overgrowing them (Castorani and Hovel 2015).
CA	California		Ecological Impact	Food/Prey
Arcuatula senhousia populations in Mission Bay decline in a seaward direction, as a result of predation by Spiny Lobsters (Panulirus interruptus) and murex snails (Pteropurpura festiva) (Cheng and Hovel 2011). However, effects on predator populations are not known.
CA	California		Ecological Impact	Habitat Change
In San Diego Bay, A. senhousia had both negative and positive effects on the eelgrass Zostera marina in experiments, reducing rhizome growth, but increasing sediment ammonium and enhancing leaf growth (Dexter and Williams 1998)., Arcuatula senhousia's mats cover large areas of subtidal flats in Mission Bay, San Diego, California. These mats increase the shear strength and stability of the sediment, favoring increased densities of other benthic organisms in the mats, including the tanaid, Leptochelia dubia, and the gastropod Barleia subtenus. The mats provide structure for tube-building organisms such as L. dubia. The mats also concentrate organic matter, favoring deposit-feeding organisms (Crooks 1998; Crooks and Khim 1999). Negative impacts of A. senhousia are not obvious or numerous, but negative correlations were seen with the deposit-feeding bivalve Solen rostriformis, and the polychaetes Euchone limnicola and Diplocirrus sp. (Dexter and Crooks 2000). Alteration of sediments by A. senhousia may be a factor in the decline of Chione fluctifraga and C. undatella (Crooks 2001).
CA	California		Ecological Impact	Trophic Cascade
Tne factor in the replacement of native bivalves by Arcuatula senhousia is that the heavy settlement of the mussel attract native perdators(Castorani and Hovel 2015), One factor in the replacement of native bivalves by Arcuatula senhousia is that the heavy settlement of the mussel attract native predators(Castorani and Hovel 2015)

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