Chelicorophium curvispinum (G.O. Sars, 1895)

Common Name: Caspian mud shrimp

Synonyms and Other Names:

Corophium curvispinum Sars, 1895; Corophium sowinskyi, Corophium devium Wundsch 1912




Silvia Waajen, onderwaterwereld.orgCopyright Info


Lodewijk RoelenCopyright Info

Identification: Chelicorophium curvispinum has a cylindrical, dorso-ventrally compressed, curled, grey-yellow body with unfused urosomal segments (with spines on uropods 1 and 2) and a small triangular rostrum. It has two pairs of antennae, the first pair slender and the second pair large and thick. The first segment of the first antennae has 3 or 4 spines along the ventral margin and 2 or 3 spines on the inner margin; antennae 1 in females is sparsely setose (hairy). The second antennae are moderately setose, with long hairs; distinguishing features of this species are that the fourth segment of these antennae bears a large curved spur and one or two smaller spurs on its ventrolateral tip, and the fifth segment bears a proximal triangular process on its ventral surface. Females and males differ in the presence of spines and spurs on the inner surface of the second antennae. This species has an evenly convex palm on its most anterior appendage (gnathopod 1), while teeth are present along the inner margin of the last segment (dactylus) of the second gnathopod (de Kluijver and Ingalsuo 1999, Lincoln 1979).  Largest adults up to 8 mm long; overwintering population 2.4-4.5 mm; juveniles < 1.8 mm (van den Brink et al. 1993, Rajagopal et al. 1998b)


Size: up to 8 mm long


Native Range: Chelicorophium curvispinum originated in the Ponto-Caspian basin, and may have colonized the adjacent Black-Azov basin when they were connected in prehistory (Mordukhai-Boltovskoi 1964). It is native to large river systems discharging into the Black and Caspian Seas, including the Volga, Dnieper, Dniester, and Danube (de Kluijver and Ingalsuo 1999, bij de Vaate et al. 2002).


Ecology: Chelicorophium curvispinum is found in salt, brackish, and fresh water (de Kluijver and Ingalsuo 1999). It is originally a brackish water species occurring in salinities of less than 6 ppt (Romanova 1975), with the ability to tolerate very low salinities (Bayliss and Harris 1988; van den Brink et al. 1993; Harris and Bayliss 1990; Taylor and Harris 1986a, b). In Black Sea lagoons and estuaries, its distribution follows the 1.5 ppt isohaline (Bortkevitch 1988). This species is most successful in waters with relatively high ionic content and requires a minimum sodium ion (Na+) concentration of 0.5 mM (Harris and Aladin 1997). The lethal minimum oxygen concentration for C. curvispinum is 0.300 mg O2/L (Dedyu 1980). Chelicorophium curvispinum is able to tolerate temperatures from 7.0-31.8°C (Jazdzewski and Konopacka 1990). Populations of this amphipod remain unchanged or gain biomass under conditions of moderate eutrophication (Kotta et al. 2012). It is intolerant of heavy organic pollution levels (Harris and Musko 1999, Jazdzewski 1980).

Chelicorophium curvispinum may aggregate on stones, wooden structures, dead and living shells, sandy sediments, clay sediment, and submerged aquatic vegetation (van den Brink et al. 1993, den Hartog et al. 1992). It can also be found in association with the green alga Cladophora (Ojaveer and Kotta 2006). In European waters, C. curvispinum density has been extremely variable, with some studies observing 100,000-750,000 individuals/m2 (highest at 2-3 m depths) (van den Brink et al. 1993, den Hartog et al. 1992), and others observing much lower densities (3,000-50,000/m2) (Bortkewitch 1987, Harris and Bayliss 1990, Musko 1989, Schöll 1990). Average densities in more recently invaded territory (Gulf of Finland) were reported to be between 125-1425 individuals/m2 (Ojaveer and Kotta 2006).

Chelicorophium curvispinum is a tube-dwelling amphipod; it collects minerals and organic particles from the water column and secretes a 1-4 cm thick layer of muddy tubes on colonization surfaces (Paffen et al. 1994). Mud used to build these tubes has ranged from 61 to 609 grams dry weight/m2 stone surface, smothering previously established organisms (van der Velde et al. 1994). Oxygen consumption, and therefore metabolism, of individuals within tubes (39 µmol/g/h) is approximately twice as high as that by free swimming individuals (22 µmol/g/h) outside of tubes (Harris and Musko 1999; Musko et al. 1995, 1998).

This amphipod is a suspension feeder, filtering phytoplankton, especially diatoms, and other suspended matter from the water (van den Brink et al. 1993). This feeding strategy results in population densities being highest where current velocities are strongest (e.g., 1-1.23 m/s), allowing C. curvispinum to most efficiently filter the largest quantity of food and obtain the greatest amount of oxygen for metabolism (van den Brink et al. 1993, van der Velde et al. 2000). This species is an important food source for a variety of fish species, including sculpin, eels, perch, ruffe, and pike perch (van den Brink et al. 1993). Other predators include birds, crayfish, and other predatory macroinvertebrates (Biro 1974; Kelleher et al. 1998, 1999; Marguillier et al. 1998).

Reproduction in C. curvispinum occurs from May to October in the Black Sea (Bortkevitch 1988) and from April to September in the Baltic (van den Brink et al. 1993). These are the warmest periods of the year (water temperature 12-20°C). Sex ratios exhibit a female bias—females outnumber males at all times of year and at many times more than double the male population (van den Brink et al. 1993). Large females (5.00-6.30 mm) become ovigerous a few weeks before smaller individuals (3.80-4.75 mm). Three generations of offspring are produced each year, following an overwintering period—the first in April to May, the second in June to July, and the third in September to October (den Hartog et al. 1992). The progeny of the first generation of summer animals (generation 2), along with the late autumn brood (generation 3), make up the next overwintering generation (Rajagopal et al. 1998b).

Brooded egg sizes range from 360 x 280 µm (Stage I) to 520 x 440 µm (Stage IV). The number of eggs carried by females and total female body length are correlated, ranging in the Rhine from 3 to 34 eggs (mean = 12) (van den Brink et al. 1993) and in Lake Balaton from 1 to 25 (mean = 6) (Musko 1989, 1990). These differences in clutch size are thought to be due to differences in food availability. Both average clutch size (Rajagopal 1998b) and growth rate (Rajagopal 1998a) have been positively correlated with the availability of chlorophyll a, which leads to increased planktonic development and greater food availability.

Embryonic development lasts about two weeks and larval development takes approximately four weeks. Most rapid growth rates occur from May to August, when water temperatures range from 15-20°C (van den Brink et al. 1993). The life span of C. curvispinum lasts no longer than 8 months (van der Velde et al. 2000).


Means of Introduction: Chelicorophium curvispinum has a moderate probability of introduction to the Great Lakes (Confidence level: High).

Potential pathway(s) of introduction: Trans-oceanic shipping

Chelicorophium curvispinum is indigenous and widespread throughout the Ponto-Caspian region and recently has invaded the Baltic Sea and many large European river systems (bij de Vaate et al. 2002). In Europe, shipping—including transport by ballast water and hull fouling—has been the primary vector of dispersal of this species outside of its native range (Harris 1991, den Hartog et al. 1992, Leppäkoski and Olenin 2001, van der Velde et al. 2000). Introduction to Great Britain, for instance, likely occurred via ballast water released from ships trading with ports on the Elbe River in northern Germany (Harris 1991). The discovery of C. curvispinum west of the Rhine River system before it was discovered in the Rhine lends further support to shipping mediated spread (den Hartog et al. 1992), as this sort of “jump” dispersal is a common indicator of human-mediated transport (MacIsaac et al. 2001).

As Great Lakes shipping traffic commonly originates in areas where this species has become established, Ricciardi and Rasmussen (1998) identified C. curvispinum as a species likely to be transported to the Great Lakes via ballast water. The discovery (but not subsequent establishment) of a related Ponto-Caspian amphipod, Corophium mucronatum, in a benthic sample collected from Lake St. Clair in 1997, suggests that C. curvispinum may at least have the potential for introduction to the Great Lakes (Grigorovich and MacIsaac 1999, Ricciardi and MacIsaac 2000). Corophium spp. appear to be unaffected by flow-through salinity shock experiments (increase of roughly 5 ppt from ambient salinity), but undergo ~35% mortality in empty-refill treatments after 48 hours of exposure (Johengen et al. 2005). However, given this species’ physiological salinity constraints, under current mandatory ballast water regulations (saltwater flushing of at least 30 ppt), the risk of its entry into the Great Lakes has been modeled as having low likelihood (Grigorovich et al. 2003).

 

 


Status: Not established in the Great Lakes
 

Chelicorophium curvispinum has a moderate probability of establishment if introduced to the Great Lakes (Confidence level: Moderate).

This species has been one of the most successful macroinvertebrate invaders in Europe, establishing populations much larger than those of any native invertebrate species within a few years of colonization (van den Brink et al. 1993, den Hartog et al. 1992, bij de Vaate et al. 2002). Densities have reached 750,000 individuals/m2 in some areas of the Rhine (van den Brink et al. 1993). With a high fecundity (see Ecology), reproducing populations are now established throughout all major European river systems and as far west as Great Britain (bij de Vaate et al. 2002). This species is able to readily disperse through ballast water transport, ship hull fouling, passive drift, and active migration (van Riel et al. 2006, van der Velde et al. 2000), with secondary spread across Europe occurring in a pattern similar to, though at a much slower rate than, that of the zebra mussel (Tittizer et al. 1994).

Chelicorophium curvispinum is a non-specific feeder (bij de Vaate et al. 2002), filtering diatoms, organic particles, and small minerals from the water column. Its superior competitive abilities—including spatial adaptation, gregarious behavior, and relatively short lifespan and generation time—have contributed to this species’ invasion success (bij de Vaate et al. 2002, van den Brink et al. 1993). Competition with other macroinvertebrate species has been well documented, most notably with the highly successful Great Lakes invader, the zebra mussel (Dreissena polymorpha).

The water temperature (up to 31.8°C) and salinity (<6 ppt) ranges tolerated by C. curvispinum are well within those of the Great Lakes and have allowed this species to be extremely successful in invasions of European rivers. Furthermore, C. curvispinum produces overwintering populations of smaller individuals (van den Brink et al. 1993) in waters of the Ponto-Caspian basin with very similar climatic conditions to those of the Great Lakes. However, its physiological tolerance (see Ecology) is restricted by other factors, such as ion concentrations, oxygen availability, chlorophyll a concentrations, flow rate, and organic pollution levels (van der Velde et al. 2000). Individuals’ ability to retain and replace Na+ and Cl- varies among populations in different locations, and some populations have adapted to freshwater by means of lower ion permeability (Harris 1991, van der Velde et al. 2000). The changing conditions of the Rhine River throughout the 20th century, specifically increases in temperature and salinity, have created more suitable conditions for the invasion of foreign species originating in brackish waters, including C. curvispinum (van den Brink et al. 1993, den Hartog et al. 1992). These conditions are consistent with the physical changes forecast for the Great Lakes as a result of climate change (Rahel and Olden 2008), suggesting that this species may benefit from the resulting habitat shifts if introduced.

Chlorophyll a concentrations required by this species are currently present only in Lake Erie’s central basin, with less than 3 µg/L typically occurring in the other lake basins (USEPA 2012). This is consistent with the predicted distribution of C. curvispinum in the Great Lakes according to the Genetic Algorithm for Rule-Set Production (GARP) model, which incorporates variable chlorophyll a levels (USEPA 2008). However, anoxic conditions have recently been present in the central basin of Lake Erie, dropping below 0.5 mg/L at certain times of year (USEPA 2012). As a result, C. curvispinum distribution is likely to be restricted to areas with sufficient flow rates, high dissolved oxygen levels, and high phytoplankton productivity.


Great Lakes Impacts: Chelicorophium curvispinum has the potential for moderate environmental impact if introduced to the Great Lakes.

In invasions across Europe, high densities of C. curvispinum have been associated with reductions in macroinvertebrate species richness (van den Brink et al. 1991, van der Velde et al. 1998). This species has outcompeted several native macroinvertebrates, including the Great Lakes invader Dreissena polymorpha (zebra mussel) (van der Velde et al. 1994). Additionally, this species has outcompeted the freshwater isopod Asellus aquaticus and several species of chironomid larvae within their native ranges in the Rhine (Kinzelbach 1997). Competition for food with other filter feeding species is expected if this species were to reach the Great Lakes (van den Brink et al. 1993), though competition for space (due to the smothering of settled individuals with their tube-building material and creating future unsuitable settling surface) has been the primary factor in reducing macroinvertebrate abundance in invaded areas of the Rhine (van der Velde et al. 1994). This species may have prevented the Great Lakes non-native Echinogammarus ischnus from forming dense populations in the Rhine (where it is native) due to the smothering of potential hard substrate settlement areas (van der Velde et al. 2000), and the production of these mud tubes has additionally led to the displacement of native filter-feeding caddisflies (Hydropsyche sp.) (van der Velde et al. 1994).

This species plays an important role in the food web of the Rhine as well as other invaded areas, driving prey switching among some fish predators (van der Velde et al. 2000). Due to the reduction in invertebrate diversity in areas where C. curvispinum has invaded, a decline in the number of predators, such as leeches, has been observed (van der Velde et al. 2000). It has been hypothesized that the prevalence of C. curvispinum in invaded rivers may lead to alterations of entire food webs (Ricciardi and Rasmussen 1998), although its role as a food source for predatory macroinvertebrates (e.g., as crayfish) requires further study (Rajagopal et al. 1998a).
The population explosion of C. curvispinum in the Rhine from 1989-1991 coincided with a decrease in the levels of total organic carbon and total suspended matter (van den Brink et al. 1993). An increase in water clarity due to particle filtration by large populations of C. curvispinum is thought to have expanded the euphotic zone, increasing the transfer of energy and nutrients to the benthos and leading to greater levels of benthic production (Rajagopal et al. 1998a).

There is little or no evidence to support that Chelicorophium curvispinum has the potential for significant socio-economic impacts if introduced to the Great Lakes.

The resulting reduction in macroinvertebrate abundance associated with C. curvispinum invasions may have a negative impact on the diet of native fish species; however, predation pressure on exotic amphipods changes rapidly depending upon prey availability. Hence, further biological monitoring is necessary in order to determine the full extent of the impact of C. curvispinum on fish diet and subsequent effect on commercial fisheries (van der Velde et al. 2000).

Chelicorophium curvispinum has the potential for moderate beneficial effect if introduced to the Great Lakes.

In areas where these species have colonized together, C. curvispinum has either greatly reduced or eliminated D. polymorpha populations by smothering settled individuals and larvae with a thick layer of dense, muddy material used for construction of tubes (van der Velde et al. 1994). After introduction of C. curvispinum to the Rhine, zebra mussel populations were seen to decrease from 1000s of individuals/m2 to 100s of individuals/m2 within four years (Paffen et al. 1994, Rajagopal et al. 1998a, van der Velde et al. 1994, 1998). This species has also potentially prevented populations of another Great Lakes invader, Echinogammarus ischnus, from forming dense populations in the Rhine (van der Velde et al. 2000). While populations of previously established invaders in the Great Lakes may be reduced by the introduction of this species, its effectiveness as a control agent is likely to be limited due to independent negative ecological consequences.
Chelicorophium curvispinum is fed upon by many fish genera, including those with species represented in the Great Lakes (van den Brink et al. 1993). In the past, this species was intentionally introduced to large rivers in the Ponto-Caspian region to increase faunal diversity and to feed fish (Jazdzewski 1980, Karpevich 1975, Mordukhai-Boltovskoi 1979). However, its potential beneficial impact as a food source for commercial fisheries in the Great Lakes is unknown.

Lastly, while water clarity is increased by the presence of large populations of this species (Rajagopal 1998a), the positive impact of this effect for humans and/or native species is likely limited.


Management: Regulations
There are no known regulations for this species.*

*Ballast water regulations applicable to this species are currently in place to prevent the introduction of nonindigenous species to the Great Lakes via shipping. See Title 33: Code of Federal Regulations, Part 151, Subparts C and D (33 CFR 151 C) for the most recent federal ballast water regulations applying to the Great Lakes and Hudson River.

Note: Check federal, state/provincial, and local regulations for the most up-to-date information.

Control
Biological
There are no known biological control methods for this species.

Physical
There are no known physical control methods for this species.

Chemical
There are no known chemical control methods for this species.

Note: Check state/provincial and local regulations for the most up-to-date information regarding permits for control methods. Follow all label instructions.


Remarks: Chelicorophium curvispinum often occurs in conjunction with C. sowinskyi (Mordukhai-Boltovskoi 1964, Ricciardi and Rasmussen 1998), which has a similar appearance. This can potentially make it difficult to distinguish between the two (Jazdzewski 1980, Jazdzewski and Konopacka 1988, Pirogov et al 1991).

The nonindigenous distribution of C. curvispinum includes: Baltic and North Seas; rivers adjacent to Ponto-Caspian (circa 1900) (Jazdzewski 1980), Spree-Havel system near Berlin (1912) (Jazdzewski and Konopacka 1996), Oder system (1913-20), Vistula and Notec (1920-33) (Leppäkoski 1984), Elbe (1923), Neman (1924), Russian/Ukrainian canals and reservoirs (1930-75), Ural River (early to mid-20th century), Lake Balaton (1934), Great Britain (1918, 1935, 1962, 1969-70) (Crawford 1935, Harris 1991, Moon 1970), Mittelland Canal (1956) (van den Brink et al. 1989), Upper Danube (1959) (van der Velde et al. 2000), lakes and reservoirs in western Asia and European USSR (1965-75), Volga through canal from Don, Dortmund-Ems Canal (1977) (van den Brink et al. 1989), Rhine River (1987) (van den Brink et al. 1989, Scholl 1990), Main River (1988) (van der Velde et al. 2000), Main-Danube Canal (1993) (Tittizer 1996), Eastern Gulf of Finland (2005) (Kotta et al. 2006).


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Author: Baker, E., L. Dettloff and A. Fusaro


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Revision Date: 1/26/2015


Citation for this information:
Baker, E., L. Dettloff and A. Fusaro, 2018, Chelicorophium curvispinum (G.O. Sars, 1895): U.S. Geological Survey, Nonindigenous Aquatic Species Database, Gainesville, FL, and NOAA Great Lakes Aquatic Nonindigenous Species Information System, Ann Arbor, MI, https://nas.er.usgs.gov/queries/greatlakes/FactSheet.aspx?SpeciesID=18&Potential=Y&Type=2&HUCNumber=, Revision Date: 1/26/2015, Access Date: 12/13/2018

This information is preliminary or provisional and is subject to revision. It is being provided to meet the need for timely best science. The information has not received final approval by the U.S. Geological Survey (USGS) and is provided on the condition that neither the USGS nor the U.S. Government shall be held liable for any damages resulting from the authorized or unauthorized use of the information.