Invasion History

First Non-native North American Tidal Record: 1957
First Non-native West Coast Tidal Record: 1957
First Non-native East/Gulf Coast Tidal Record: 2001

General Invasion History:

Palaemon macrodactylus is native to coastal waters and estuaries of the Northwest Pacific, from the Vladivostok area of Russia, through Japan (from Hokkaido to Kyushu), South Korea (Pusan) and Taiwan (Newman 1963; Golikov et al. 1976, but see Ashelby et al. 2013). It has been a highly successful invader, appearing in San Francisco Bay in 1957 (Newman 1963), northern Europe in 1992 (Ashelby et al. 2004; Gonsalez-Ortegon et al. 2007), Atlantic Spain in 1997 (Cuesta et al. 2004), and Argentina in 2000 (Mar del Plata, Spivak et al. 2006). This shrimp is sold as food in Japan (Holthuis 1989).

North American Invasion History:

Invasion History on the West Coast:

Palaemon macrodactylus was first observed in the Northeast Pacific in north San Francisco Bay in 1957, and by 1961, was present in the South, Central, and San Pablo Bays, and occurring upstream to Antioch at the western edge of the Delta (Newman 1963). In 1962, it was collected in Los Angeles Harbor (Carlton 1979), and subsequently has become established in Mission Bay (in 2001, Fairey et al. 2002), Los Penasquitos and San Dieguito Lagoons (in 2000, http://www.perl.sdsu.edu/Reports/LPL01-02.pdf; Cohen et al. 2002), Santa Monica Bay (in 1977, Carlton 1979), Malibu Lagoon (in 1984, USNM 205237, US National Museum if Natural History 2007), Morro Bay (in 2001, Fairey et al. 2002), and Elkhorn Slough (in 1981, Standing 1981, cited by Wasson et al. 2001). In 1987, it was found in Coos Bay, Oregon (Cohen and Carlton 1995). It has also been reported from Humboldt Bay, northern California (in 1995, Cohen and Carlton 1995; Wonham and Carlton 2005, but was not mentioned in Boyd et al. 2002); Willapa Bay, Washington (Jensen, G.C., 1995, Pacific Coast Crabs and Shrimps, cited by USGS Nonindigenous Aquatic Species Program 2009); and Boundary Bay, British Columbia (Lamb and Hanby 2005, cited by Ashelby et al. 2013).

Invasion History on the East Coast:

Palaemon macrodactylus was collected in three locations in the Bronx, New York City, on the East River, connecting the Hudson River and Long Island Sound, in 2001-2002 and 2006. About 120 specimens were collected, indicating an established population (Warkentine and Rachlin 2010). In 2009, specimens of P. macrodactylus were collected in eastern Long Island Sound, in the Mystic River, Connecticut (USGS Nonindigenous Aquatic Species Program 2010), and in 2010, this shrimp was collected in the Providence River, Rhode Island, at the head of Narragansett Bay (Cremins 2010). In 2009, a specimen of P. macrodactylus was collected in the James River estuary, Virginia, and a second was identified from the York River (Robert Llanso, personal communication). A third specimen, caught in the summer of 2007, was identified among shrimp caught from the Smithsonian Environmental Research Center dock on the Rhode River, Edgewater, Maryland (Ruiz et al., unpublished data). The extent of the P. macrodactylus invasion in East Coast waters is unknown, and may be difficult to detect, given the similarity of this shrimp to the three native Palaemonetes species. Surveys are being planned by Chesapeake Bay researchers in order to determine the abundance and distribution of this shrimp in Chesapeake Bay (Eric Bah, Greg Ruiz, personal communications). The likely sources of the East Coast P. macrodactylus are the newly established populations on the coasts of Europe, from Spain to Germany (Warkentine and Rachlin 2010).

Invasion History Elsewhere in the World:

In European waters, Palaemon macrodactylus was first collected at West Thurrock Power Station, on the Thames estuary, England in 1992 (Worsfold and Ashelby 2006). It is established at several sites in the Thames estuary and in the Stour, Orwell and Medway estuaries, (in 2000, Ashelby et al. 2004). This shrimp has spread to several ports in northern Europe including Walsoorden, in the Westerschelde estuary, Netherlands (in 1999, d'Udekem d'Acoz et al. 2004, cited by González-Ortegón et al. 2007); Zeebruge, Belgium (in 2006, Kerckof et al. 2007); and Bremen, Germany (in 2005, González-Ortegón et al. 2007). To the south, P. macrodactylus is established in the Gironde Estuary, France, on the Bay of Biscay (in 2006, Beguer et al. 2007) and the Guadualquivir, Guadalete, and San Pedro River estuaries in Spain (in 1999, Cuesta et al. 2004). In 2005 and 2010, larvae of P. macrodactylus were collected in plankton in the Mediterranean, off the Balearic Islands, Spain, but adult populations have not yet been found (Torres et al. 2012; Ashelby et all. 2013). In 2009, it was collected in Constanta, Romania, on the Black Sea, where it is apparently established (Micu and Nita 2009).

Palaemon macrodactylus was collected in Australia in 1979 near a power plant in Lake Macquarie, New South Wales, but has not been collected since (Pollard and Hutchings 1990). However, it has become established in Mar del Plata, Argentina (in 2000, Spivak et al. 2006).


Description

Palaemon macrodactylus is a caridean shrimp with a distinct, well-developed rostrum.  It has at least eight, but usually 10-12 dorsal teeth with three (rarely two, J.T. Carlton) teeth behind the orbit (Gonzales-Ortegon and Cuesta 2006). The rostrum lacks a strong ventral expansion and has a double row of setae along the ventral margin. Supraorbital, suborbital and hepatic spines are absent. Both legs of the first pair are chelate (clawed). The carpus (second segment from the tip) of the second walking leg is not annulated (ringed). The length of the dactylus (toe) of the second leg is short, less than half the length of the propodus (terminal segment). The telson terminates in a short strong median spine and a pair of long, strong mediolateral spines, with a median pair of short plumose setae. The body is transparent or nearly transparent, with a reddish hue in the tail fan and antennary area. On rare occasions, it is dark green or olive-drab (Newman 1963; Kuris et al., in Carlton 2007). There are a few oblique transverse stripes on the carapace. In males, the abdomen is frequently translucent, but in females it is quite pigmented, with reddish spots covering all of the body surface and a whitish longitudinal stripe running along the back (Newman 1963; d'Udekem d'Acoz et al. 2005).


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Arthropoda
Subphylum:   Crustacea
Class:   Malacostraca
Subclass:   Eumalacostraca
Superorder:   Eucarida
Order:   Decapoda
Suborder:   Pleocyemata
Infraorder:   Caridea
Family:   Palaemonidae
Genus:   Palaemon
Species:   macrodactylus

Synonyms

Potentially Misidentified Species

Palaemon adspersus
Baltic Shrimp, native to East Atlantic, introduced to Gulf od St. Lawrence

Palaemon elegans
Rockpool Shrimp, native to East Atlantic, introduced to East Coast

Palaemon kadiakenisis
Mississippi Grass Shrimp =Palaemonetes kadiakiensis, native to Interior Basin of North America, introduced to the Sacramento-San Joaquin Delta

Palaemon modestus
Siberian Prawn, freshwater shrimp (=Exopalaemon modestus), introduced to West Coast rivers

Palaemon mundusnovus
Brackish Grass Shrimp, = Palaemonetes intermedius, NW Atlantic native, (de Grave and Ashelby 2013; González-Ortegón 2015)

Palaemon pugio
Daggerblade Grass Shrimp, = Palaemonetes pugio, NW Atlantic native (de Grave and Ashelby 2013; González-Ortegón 2015)

Palaemonetes vulgaris
Common Grass Shrimp,= Palaemonetes vulgaris, NW Atlantic native (de Grave and Ashelby 2013; González-Ortegón 2015)

Ecology

General:

Life History- During reproduction in caridean shrimps, the copulating pair is usually oriented at right angles to one another, with the genital regions opposing each other. The modified first and second pairs of pleopods are used to transfer a spermatophore to a receptacle between the thoracic legs of the female (Barnes 1983). After mating, female palaemonid shrimps carry broods of fertilized eggs on their abdomen. Females carry 150-2000 eggs, increasing with body size (Seigfried 1980; Beguer et al. 2011). These hatch into planktonic larvae called zoeae, which have feathery appendages. Shrimp zoeae lack the prominent spines seen in brachyuran crabs, and look quite shrimplike (Johnson and Allen 2005). They go through several molts and metamorphose into postlarvae, which have well-developed walking legs and pleopods (swimmerets). After a subsequent molt, the body takes on the adult shape. Adults mature at sizes as small as 16-17 mm length. Females tend to be larger than males, reaching maximum sizes of 45-70 mm, compared to 31.5-45 mm for males (Vazquez et al. 2012).

Ecology- Palaemon macrodactylus favors pilings, walls, debris, and other forms of shelter (Crooks et al. 2016). It is often most abundant in low-salinity waters, possibly due to reduced competition with native species (Newman 1963; Siegfried 1980; Gonzalez-Ortegon et al. 2010). It reproduces successfully at 3-34 PSU, and larvae born at higher salinities, survive and develop at 1 PSU. However, rates of survival increase at higher salinity (Vazquez et al. 2016).

Food:

Plants, Mysids, Amphipods, Crabs

Trophic Status:

Omnivore

Omni

Habitats

General HabitatGrass BedNone
General HabitatSalt-brackish marshNone
General HabitatUnstructured BottomNone
General HabitatMarinas & DocksNone
Salinity RangeMesohaline5-18 PSU
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Salinity RangeOligohaline0.5-5 PSU
Tidal RangeSubtidalNone
Vertical HabitatEpibenthicNone
Vertical HabitatNektonicNone


Tolerances and Life History Parameters

Minimum Salinity (‰)0.7Cohen and Carlton 1995; Born 1968
Maximum Salinity (‰)51Born 1968
Minimum Reproductive Salinity3Experimental, no hatching at 1 PSU, but older larvae survived and developed at this salinity(Vazquez et al. 2016)
Maximum Reproductive Salinity34Experimental, highest tested (Vazquez et al. 2016)
Minimum Duration11Laboratory rearing, animals from San Francisco Bay (Little 1969)
Maximum Duration21Laboratory rearing, animals from San Francisco Bay (Little 1969)
Minimum Length (mm)16.8Ovigerous females, Mar de la Plata estuary, Argentina (Vazquez et al. 2012); 17 mm for males (Siegfried 1980)
Maximum Length (mm)70Ovigerous females (Ashelby et al. 2004, Orwell estuary, England); 40 mm for males (San Francisco Bay estuary, Seigfried 1980).
Broad Temperature RangeNoneCold temperate-Warm temperate
Broad Salinity RangeNoneOligohaline-Hyperhaline

General Impacts

Economic Impacts

Palaemon macrodactylus is sold as food in Japan, but apparently only has minor value as a food item (Holthuis 1989). It is a potential food source for commercial and sport fish, but may affect the abundance of other fish-food species through predation.

Ecological Impacts

Palaemon macrodactylus is a potential competitor with other caridean shrimps, and though omnivorous, predominantly feeds on invertebrates.

Competition- The extent of competition between P. macrodactylus and native shrimp species has been difficult to assess, due to differing habitat preferences among the species. In San Francisco Bay, the native shrimps Crangon spp. prefers soft, bare sediment, while P. macrodactylus prefers rubble, man-made structures, and vegetation (Newman 1963). However, both species feed on the mysid Neomysis mercedis, and could be competing for food, particularly in the fall, when N. mercedis numbers are low (Sitts and Knight 1979). In the Guadalquivir Estuary, Spain, competition between P. macrodactylus and the native P. longirostris is considered possible, since diets strongly overlap, but may be limited, since P. macrodactylus primarily utilizes the low-salinity regions of the estuary, where P. longirostris is rare (Gonzalez-Ortegon et al. 2010). One competitive advantage of P. macrodactylus over P. longirostris, is its greater ability to use areas of low oxygen concentration (Gonzalez-Ortegon et al. 2010). In the Gironde estuary, France, P. macrodactylus initially colonized areas little-used by P. longirostris, but has begun to displace it due to higher reproductive rates (Beguer et al. 2011). In the Guadalquivir estuary, in Spain, P. macrodactylus showed wider temperature and salinity tolerances than the native P. longirostris and P. varian (Lejeusne et al. 2014).

Predation- In the San Francisco Bay Delta in the 1970s, the major prey of P. macrodactylus was the native mysid Neomysis mercedis (Sitts and Knight 1979). Mysids are also important prey for juvenile fish. Since then, N. mercedis has been replaced by smaller species of Asian mysids. We have no recent data from San Francisco Bay or elsewhere on the role of P. macrodactylus as a predator.

Regional Impacts

P090San Francisco BayEcological ImpactCompetition
In San Francisco Bay, differences in habitat preference between the native shrimp Crangon spp. and P. macrodactylus may minimize competition (Newman 1963). However, both species feed on the mysid Neomysis mercedis, and could be competing for food, particularly in the fall, when N. mercedis numbers are low (Sitts and Knight 1979).
NEP-VNorthern California to Mid Channel IslandsEcological ImpactCompetition
In San Francisco Bay, differences in habitat preference between the native shrimps Crangon spp. and P. macrodactylus may minimize competition (Newman 1963). However, both species feed on the mysid Neomysis mercedis, and could compete for food, particularly in the fall, when N. mercedis numbers are low (Sitts and Knight 1979). Neomysis mercedis is now rare in the San Fracisco estuary, largely replaced by a smaller biomass of Asian mysids. We have no information on how these changes have affected interactions between P. macrodactylus and native Crangon spp.
NEP-VNorthern California to Mid Channel IslandsEcological ImpactPredation
The most frequent food item for both P. macrodactylus and Crangon franciscorum in San Francisco Bay was the mysid Neomysis mercedis. In 1976, C. franciscorum consumed up to 6.2% of the mysid biomass, while P. macrodactylus consumed up to 4.8% (Sitts and Knight 1979). Predation on N. mercedis is significant, because this mysid is also an important food source for planktivorous adult and juvenile fish. However, this mysid is now scarce in the San Francisco estuary, largely replaced by smaller species of Asian mysids, as part of a reorganization of the food web caused by the invasion of the Asian Brackish-Water Clam (Corbula amurensis). These changes have probably altered the diet of P. macrodactylus, but we have found no recent diet studies on this shrimp.
P090San Francisco BayEcological ImpactPredation
The most frequent food item for both P. macrodactylus and Crangon franciscorum in San Francisco Bay was the mysid Neomysis mercedis. In 1976, C. franciscorum consumed up to 6.2% of the mysid biomass, while P. macrodactylus consumed up to 4.8% (Sitts and Knight 1979). Predation on N. mercedis is significant, because this mysid is also an important food source for planktivorous adult and juvenile fish. However, this mysid is now scarce in the San Francisco estuary, largely replaced by smaller species of Asian mysids, as part of a reorganization of the food web caused by the invasion of the Asian Brackish-Water Clam (Corbula amurensis). These changes have probably altered the diet of P. macrodactylus, but we have found no recent diet studies on this shrimp.
NEA-VNoneEcological ImpactCompetition
In the Guadalquivir Estuary, Spain, competition between P. macrodactylus and the native Palaemon longirostris is considered possible, since diets strongly overlap (Gonzalez-Ortegon et al. 2010). Competition may be limited, however, since P. macrodactylus primarily utilizes the low-salinity regions of the estuary, where P. longirostris is rare (Gonzalez-Ortegon et al. 2010). One competitive advantage of P. macrodactylus over P. longirostris, and other species, is its greater ability to use areas of low oxygen concentration (Gonzalez-Ortegon et al. 2010; Gonzalez-Ortegon et al. 2013). In the Gironde estuary, France, P. macrodactylus initially colonized areas little-used by P. longirostris, but has begun to displace it due to higher reproductive rates (Beguer et al. 2011).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
NWP-4a None 0 Native Estab
NEP-V Northern California to Mid Channel Islands 1957 Def Estab
NEP-IV Puget Sound to Northern California 1987 Def Estab
NEP-VI Pt. Conception to Southern Baja California 1962 Def Estab
AUS-X None 1979 Def Unk
NWP-3b None 0 Native Estab
NWP-4b None 0 Native Estab
NEA-V None 1997 Def Estab
NEA-II None 1992 Def Estab
P050 San Pedro Bay 1962 Def Estab
P170 Coos Bay 1987 Def Estab
SA-I None 2000 Def Estab
P022 _CDA_P022 (San Diego) 1994 Def Estab
P030 Mission Bay 2001 Def Estab
P060 Santa Monica Bay 1977 Def Estab
P070 Morro Bay 2000 Def Estab
NWP-3a None 0 Native Estab
P080 Monterey Bay 1981 Def Estab
P090 San Francisco Bay 1957 Def Estab
P130 Humboldt Bay 1995 Def Estab
P270 Willapa Bay 1995 Def Unk
P010 Tijuana Estuary 1995 Def Estab
P093 _CDA_P093 (San Pablo Bay) 1959 Def Estab
NA-ET3 Cape Cod to Cape Hatteras 2001 Def Estab
M130 Chesapeake Bay 2007 Def Estab
MED-IX None 2009 Def Estab
M060 Hudson River/Raritan Bay 2001 Def Estab
M040 Long Island Sound 2001 Def Estab
M020 Narragansett Bay 2010 Def Estab
NEA-IV None 2007 Def Estab
P061 _CDA_P061 (Los Angeles) 1984 Def Estab
NEP-III Alaskan panhandle to N. of Puget Sound 2005 Def Unk
P297 _CDA_P297 (Strait of Georgia) 2005 Def Unk
MED-II None 2005 Def Unk
SA-II None 2007 Def Estab
MED-VII None 2012 Def Unk
P020 San Diego Bay 2011 Def Estab
NA-ET2 Bay of Fundy to Cape Cod 2012 Def Estab
N180 Cape Cod Bay 2014 Def Estab
N170 Massachusetts Bay 2012 Def Estab
N130 Great Bay 2014 Def Estab
M010 Buzzards Bay 2014 Def Estab
B-VII None 2014 Def Estab
M023 _CDA_M023 (Narragansett) 2019 Def Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
30029 Cohen et al. 2005 (SF Bay Area RAS) 2004 2004-05-28 Moores Landing, San Francisco Bay Def 38.2261 -122.3076
30390 Wasson et al, 2001 (Elkhorn Slough Survey) 1998 1998-03-01 Elkhorn Slough General Location Def 36.8086 -121.7856
768149 Ruiz et al., 2015 2012 2012-09-06 Loch Lomond Marina, San Francisco Bay, CA, California, USA Def 37.9736 -122.4802
768216 Ruiz et al., 2015 2012 2012-09-10 Pittsburg Marina, San Francisco Bay, CA, California, USA Def 38.0346 -121.8829
768331 Ruiz et al., 2015 2013 2013-08-23 Loch Lomond Marina, San Francisco Bay, CA, California, USA Def 37.9723 -122.4829

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