Dikerogammarus haemobaphes (Eichwald, 1841)

Common Name: Scud

Synonyms and Other Names:

Small-humped scud, Dikerogammarus villosus balatonicus Ponyi, Gammarus haemobaphes

Dr. Michal GrabowskiCopyright Info

Identification: Dikerogammarus haemobaphes has a laterally compressed, curled, and semi-transparent whitish body consisting of a head (cephalon), thorax (pereon), and abdomen. Its head contains one pair of eyes, mouthparts (gnathopods), and two pairs of antennae, each with peduncle (broad stalk, connected to head) and flagellum (narrow, outer tip). Its pereon consists of seven segments, each with a pair of walking legs (pereopods). Its abdomen consists of six segments divided into two three-segment parts: pleosome (anterior) with brush-like limbs known as pleopods, and urosome (posterior) with shorter, immobile rod-like limbs called uropods.

This species can be distinguished from other Dikerogammarus species by morphological features of the second (lower) pair of antennae and of the first and second urosomal segments. In D. haemobaphes, the peduncle and flagellum of antenna 2, as well as the gnathopods, are characterized by short bristles rather than dense, long bristles. Urosomes 1 and 2 of D. haemobaphes have a shallow dorsal protuberance tipped with two spines instead of the pointed dorsal protuberance seen in other species of Dikerogammarus. Additionally, in contrast to the morphologically similar D. villosus, no color variation is exhibited by this species (Müller et al. 2002). However, these distinguishing features apply mainly to adult males and are less diagnostic in females and juveniles.

Adult male: average 16 mm (range 9-21 mm); adult female: average 11 mm (range: 7-15 mm); stage 2 egg: average 0.43 mm (Bacela et al. 2009, Kinzler et al. 2009).

Size: to 21mm

Native Range: Ponto-Caspian basin

This species is not currently in the Great Lakes region but may be elsewhere in the US. See the point map for details.

Ecology: Dikerogammarus haemobaphes is euryoecious (adapted to varied ecological conditions), preferring to inhabit solid substrates, macrophytes, and filamentous algae in large rivers and lakes (Kiticyna 1980, Muskó 1994). It is able to tolerate a relatively wide temperatures range (6-30°C) (Kiticyna 1980). Although Grigorovich et al. (2003) listed D. haemobaphes as occurring naturally in waters of 17 ppt salinity, this species is most likely restricted to 0-8 ppt (Ponomareva 1976). It is most commonly found near mouths of freshwater rivers (typically 1-1.5 PSU, practical salinity units) than in more saline waters (2-8 PSU) (Grabowski et al. 2006; Jazdzewski et al. 2002, 2004, 2005). The lethal minimum oxygen concentration for D. haemobaphes is 0.345 mg O2/L; while such conditions are considered hypoxic, several other Ponto-Caspian amphipods invaders in the Baltic Sea can tolerate even lower oxygen concentrations (Dedyu 1980).

Dikerogammarus haemobaphes is a dietary generalist (bij de Vaate et al. 2002), feeding on detritus, sediments, unicellular and filamentous algae, and other small crustaceans. Its predation intensity on animal food sources such as chironomids, oligochaetes, crustaceans, and mayflies increases during the summer months when water temperatures are higher (van der Velde et al. 2009). Cannibalism within this species is significantly higher (~50%) than that of other European gammarid species (Kinzler et al. 2009).

Dikerogammarus haemobaphes is frequently found in association with another Ponto-Caspian mass invader, Dreissena polymorpha, preferring to settle on living zebra mussel shells over other substrate types (Kobak et al. 2009, Musko´ 1993, bij de Vaate et al. 2002, Wawrzyniak- Wydrowska and Gruszka 2005). Dreissenid shell surface properties are thought to attract D. haemobaphes, with a distinct preference shown for living mussels over empty shells as well as clean shells over varnished shells (Kobak et al. 2009). Additionally, the living mussel shells serve as a better habitat for prey items, including chironomids (Botts et al. 1996, Mörtl and Otto-Rothhaupt 2003, Ricciardi et al. 1997, Stewart et al. 1998).

The reproductive period of D. haemobaphes lasts from April to October, with the production of spring, summer, and autumn (overwintering) generations, each with 3-5 cohorts (Bacela et al. 2009, Mordukhai- Boltovskoi 1949). The autumn (overwintering) generation begins to reproduce in April, and by mid May its progeny (spring generation) appear. At this point, population size structure is strongly bimodal due to the presence of the parental generation. In June, the spring generation begins to reproduce and the autumn generation dies off. In early July, the summer generation is released and rapidly matures, giving rise to the next autumn generation. As in many other gammarid species, mean size of mature individuals, mean size of breeding females, and fecundity declines over subsequent generations (autumn to spring to summer) (Bacela et al. 2009, Kuradina 1975, Musko 1993).

Male to female sex ratios are generally close to 1:1, with intersex individuals (able to both lay and fertilize eggs in a brood pouch) representing roughly 1% of the population (Bacela et al. 2009). The maturity index (minimal size/mean size of gravid females) of D. haemobaphes is low (~0.67), indicating this species reaches maturity at a relatively young age. Clutch size varies from 5 to 128 eggs, with mean estimates ranging from 20-52 eggs (Bacela et al. 2009, Briskina 1950, Gudkova and Melnikova 1969, Kiticyna 1980, Kurandina 1975, Musko´ 1993). Clutch size has been observed to increase with female body length according to a log-linear relationship, with both clutch and body size varying in different habitats (e.g., mean brood sizes of 35 and 52 in river and reservoir populations, respectively) (Bacela et al. 2009). The partial fecundity index (mean clutch size/female size) for D. haemobaphes reported in this same study is relatively high, with values of 3.3 for river females and 4.3 for reservoir females. These variations in size and fecundity based sampling location are possibly due to differences in predation pressure and/or hydrological conditions such as water velocity (Adams et al. 1989).

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

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

Dikerogammarus haemobaphes has been proposed to be able to survive partial to complete ballast water exchange due to one reported natural occurrence in 17 ppt salinity waters (Grigorovich et al. 2003), but other estimates of this species’ salinity tolerance are more restrictive (0-8 ppt) (Ponomareva 1976). Based on the former, Grigorovich et al. (2003) assessed D. haemobaphes as having a high Great Lakes invasion risk mediated by both BOB and NOBOB vessels. However, the closely related European invader D. villosus exhibited 0% survival after 24 hours in 34 PSU empty-refill and flow-through salinity treatments (Santagata et al. 2008), suggesting that D. haemobaphes may be similarly impacted by current ballast water regulations.





Status: Not established in North America, including the Great Lakes
Dikerogammarus haemobaphes has a high probability of establishment if introduced to the Great Lakes (Confidence level: Moderate).

Ponto-Caspian amphipod identified as having high probability of invasion if introduced to the Great Lakes (Grigorovich et al. 2003, Ricciardi and Rasmussen 1998, U.S. EPA 2008); listed as invasive in Baltic Sea (Baltic Sea Alien Species Database 2007).The climatic conditions of the native (Ponto-Caspian) and introduced (Baltic) ranges of D. haemobaphes are very similar to those of the Great Lakes. While D. haemobaphes is able survive in moderately wide temperature and salinity ranges, as well as hypoxic conditions, it is primarily a freshwater, riverine species (see Ecology). Its westward spread through all major European river systems connecting the Ponto-Caspian with the Baltic may have been limited by salinity levels to inland water courses (see bij de Vaate et al. 2002). Nevertheless, its physiological tolerances are well within conditions occurring in the Great Lakes. Increased salinization as a potential effect of climate change (Rahel and Olden 2008) may be inconsequential to this species’ establishment in the Great Lakes, as it tolerates salinities up to 8 ppt (Ponomareva 1976). Arbaciauskas (2002) hypothesized that oxygen concentration is the principal limiting factor in determining the survival and sustainability of populations of Ponto-Caspian amphipods. Therefore, anoxic conditions, as present in the central basin of Lake Erie (Summers 2001), may prevent D. haemobaphes from establishing in some regions of the Great Lakes.

Feeding plasticity, high reproductive capacity, relatively small eggs, short egg development time, fast sexual maturation, brooding, and production of multiple generations per year are factors thought to contribute to the invasion success of this species (Bacela et al. 2009, bij de Vaate et al. 2002). Dikerogammarus haemobaphes fecundity is moderate compared to that of the invasive gammarids D. villosus and P. robustoides (Bacela et al. 2009), but it is high in relation to invasive European gammarids as a group (Grabowski et al. 2007a). The autumn generation typically overwinters, but in thermally polluted waters (e.g., hydroelectric cooling water discharge), this species may reproduce year round (Kiticyna 1980); therefore, warming waters as a result of climate change could be beneficial to its invasion success.

D. haemobaphes has outcompeted and displaced native European gammarids, but it has also experienced declines in European waterways following expansion of the related invasive amphipod Dikerogammarus villosus (Jazdzewski et al. 2004, 2005; Kinzler et al. 2009). Dikerogammarus haemobaphes constitutes a food base for multiple fish genera (Grabowska and Grabowski 2005; Kelleher et al. 1998, 2000; Kostrzewa and Grabowski 2003), though the extent to which this predation will have an effect on potential populations in the Great Lakes is unknown.

Invasive dreissenid mussels are likely to facilitate the establishment of D. haemobaphes in the Great Lakes, as this amphipod preferentially colonizes living zebra mussel shells over other substrate types, including empty shells and stones, in the Ponto-Caspian and other newly invaded regions (see Ecology).

The dispersal rate of D. haemobaphes across Europe is similar to that of many other Ponto-Caspian invasive amphipods (e.g., D. villosus), spreading across the entire European continent in roughly 50 years (bij de Vaate et al. 2002).

Great Lakes Impacts: Current research on the potential for environmental impact of Dikerogammarus haemobaphes if introduced to the Great Lakes is inadequate to support proper assessment.

Upon introduction to new waterways, D. haemobaphes has outcompeted native European gammarids, including displacing Chaetogammarus ischnus in the lower Vistula Lagoon (Jazdzewski et al. 2004, 2005). Additionally, D. haemobaphes is a vector of gregarines, a group of unicellular parasites that infect the intestines of invertebrates (Codreanu-Balcescu 1995). However, the transfer of these parasites to native species is unknown (Grabowski et al. 2007b).

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

There is little or no evidence to support that Dikerogammarus haemobaphes has the potential for significant beneficial effects if introduced to the Great Lakes.

Dikerogammarus haemobaphes constitutes a food base for perches, gobies, and eels (Grabowska and Grabowski 2005; Kelleher et al. 1998, 2000; Kostrzewa and Grabowski 2003). It was intentionally stocked in large European rivers prior to the 1970s to enhance the fish fauna (Karpevich 1975, Jazdzewski 1980, bij de Vaate et al. 2002).

Management: Regulations (pertaining to the Great Lakes)

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.


There are no known biological control methods for this species.

There are no known physical control methods for this species

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: Dikerogammarus haemobaphes was first observed in a non-native location in 1976, when it migrated up the Danube River through the southern corridor and arrived in the German section of the upper Danube (Tittizer 1996). It was subsequently observed in the Main-Danube canal in 1993 (Schleuter et al. 1994), through which it migrated to the North Sea basin via the Rhine River (Schöll et al. 1995). In 1997, this species was first discovered in Poland in the Vistula River (Konopacka 1998). Around this time, it was also discovered in the Notec and Bug rivers, tributaries of the Oder and Vistula rivers, respectively (Jazdzewski and Konopacka 2000, 2002; Jazdzewski et al. 2002). Dikerogammarus haemobaphes was observed in the central and southern corridors of the Volga River, as well as the upper Volga basin, for the first time around the year 2000 and quickly became abundant (L’vova et al. 1996, Jazdzewski et al. 2004). It is now also present in the Great Masurian Lakes (Jazdzewski 2003) and in a small mesotrophic lake in the Vistula valley (Grabowski and Bacela 2005).


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

Contributing Agencies:

Revision Date: 10/14/2021

Citation for this information:
Baker, E., L. Dettloff and A. Fusaro, 2021, Dikerogammarus haemobaphes (Eichwald, 1841): 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?Species_ID=3613&Potential=Y&Type=2&HUCNumber=, Revision Date: 10/14/2021, Access Date: 12/7/2021

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.