Alien Crustacea in Polish Waters – Amphipoda

Home , Amphipoda, Chelicorophium, Corophiidae, Gammaridae, Gammaroidea, Gammarus pulex

Research Article

Alien Crustacea in Polish waters – Amphipoda

Michał Grabowski*, Krzysztof Jażdżewski and Alicja Konopacka Department of Invertebrate Zoology and Hydrobiology, University of Łódź, Banacha 12/16, 90-237 Łódź, Poland E-mail: [email protected] *Corresponding author

Received: 1 November 2006 / Accepted: 9 February 2007

Abstract

Among ca. 750 species of Crustacea recorded from Poland, 18 representatives of 5 orders of macro-crustaceans have been identified as alien species that have invaded or have been introduced to Polish waters. Out of 44 freshwater, brackishwater and semiterrestrial species of Amphipoda occurring in Poland (not counting several stygobiotic species), 8 species from three families may be included in this group. They are Corophiidae: Chelicorophium curvispinum (=Corophium curvispinum), Gammaridae: Gammarus roeselii, G. tigrinus, Chaetogammarus ischnus (=Echinogammarus ischnus), Pontogammaridae: Dikerogammarus haemobaphes, D. villosus, Pontogammarus robustoides and Obesogammarus crassus. It is noticeable that most of them (C. curvispinum, C. ischnus, D. haemobaphes, D. villosus, P. robustoides, O. crassus) are of Ponto-Caspian origin, one species was introduced from North America (G. tigrinus), and one from south-eastern Europe (G. roeselii). All the species listed above have spread widely in Poland, in large rivers and artifical reservoirs (Chelicorophium curvispinum, Dikerogammarus villosus, D. haemobaphes, Pontogammarus robustoides, Gammarus tigrinus) or in medium sized rivers (Gammarus roeselii), in brackish coastal waters (Obesogammarus crassus) or both in fresh and brackish waters (Gammarus tigrinus, Pontogammarus robustoides). In most places they successfully outnumber or even completely replace native amphipod species. This paper presents data on biogeography, history, biology and ecology of alien species, as well as the consequences of their invasion in Poland with an extended bibliography and references to other European countries. Key words: biological invasions, alien Amphipoda, neozoa, Vistula River, Oder River, Central Europe

Introduction Chelicorophium curvispinum, Gammaridae: Gammarus roeselii, G. tigrinus, Chaeto- To date, ca. 750 species of Crustacea have been gammarus ischnus, and Pontogammaridae: recorded in Poland (Razowski 1997; Jażdżewski Dikerogammarus haemobaphes, D. villosus, et al. 2002). From this number 18 species, Pontogammarus robustoides, Obesogammarus belonging to 5 orders of macrocrustaceans, have crassus are recognised as alien (Figure 1). It is been identified as alien species that have entered noticeable that most of them are of Ponto- Polish waters in historical times. This paper, Caspian origin, one species was introduced from dedicated to alien amphipod crustaceans, is a North America and one from south-eastern continuation of work by Grabowski et al. (2005), Europe, namely the Balkan Peninsula (Annex). which described the geographical sources and The problem of alien amphipod species migration routes for alien Crustacea entering invading European waters was discussed first by Polish waters, and summarised the invasion Jażdżewski (1980) and then, more recently by history and progress of alien Decapoda in Bij de Vaate et al. (2002). Other detailed studies Poland. on particular species or areas in Poland have Amphipoda in Polish surface waters are been performed by Jażdżewski (2003), Jażdżew- represented by 44 species including freshwater, ski and Konopacka (2002), Jażdżewski et al. brackishwater and semiterrestrial taxa. Among (2002, 2004, 2005), Konopacka (2004) and by them, 8 species from three families, Corophiidae: Grabowski et al. (2006). Morphological features

25 M. Grabowski et al.

Central Europe and in Poland was given in the papers by Eggers and Martens (2001, 2004) and by Konopacka and Jażdżewski (2002). The aim of this work is to summarize the historical changes in the distribution of alien Amphipoda in Polish waters, including a bibliography and extensive references to neighbouring countries. In addition, to give an update on the very recent invasion progress of these species, to describe their biology and to discuss the ecological consequences of these invasion events.

Figure 1. Contribution of alien species to amphipod fauna in Species of Ponto-Caspian origin Poland. Chelicorophium curvispinum (G.O. Sars, 1895) (Figure 2) This is the oldest amphipod invader of Ponto- Caspian origin noted in Polish waters. Natively it occurs in large rivers (e.g. Volga, Dniester, Danube) of the Black and the Caspian seas. The earliest record of C. curvispinum outside its native range comes from Wundsch (1912). He found it in the Spree-Havel system near Berlin in Germany. Apparently, the species had arrived into the Baltic Sea and the North Sea drainage systems through the river connections defined as the central corridor (Bij de Vaate et al. 2002). The species evidently had crossed the Polish territory and perhaps had been present in the Middle Vistula and the Noteć rivers well before it was discovered in Poland in 1920s (Kulmatycki 1930; Wolski 1930). By now it has dispersed as far as the Great Britain, where it was recorded in the 1930s (Crawford 1935). Quite recently, the species has begun a rapid expansion in the Lower Rhine system (Den Hartog et al. 1992; Van den Brink et al. 1989; Van den Brink et al. 1993 and Van der Velde et al. 2000). Now in Poland, the species is very common and abundant in numerous localities along Vistula, Oder, Bug, Narew and Noteć Figure 2. Chelicorophium curvispinum. Photo by Michał (Jażdżewski 1980; Jażdżewski and Konopacka Grabowski. 1990, 2000, unpublished data). C. curvispinum is associated with clumps of another Ponto-Caspian invader, Dreissena polymorpha (Pallas, 1771) – in the lower Vistula dredged colonies of the of particular species were presented in several zebra mussel were always accompanied by C. papers (e.g. Jarocki and Demianowicz 1931; curvispinum (Jażdżewski and Konopacka 2002). Micherdziński 1959; Jażdżewski 1975; Jażdżew- This phenomenon can be also observed in other ski and Konopacka 1990; Konopacka 1998; invaded areas in Western Europe (Devin et al. Gruszka 1999; Konopacka 2004). A key for the 2003), as well as in the species’ native occur- identification of alien amphipods occur-ring in rence, i.e. in the Dnieper River. Apparently the

26 Alien Crustacea in Polish waters – Amphipoda

probable migration routes of this species in Poland are presented in Figure 3.

Chaetogammarus ischnus (Stebbing, 1899) This is also an older amphipod immigrant to Polish waters. The native range of this species consists of the Black and Caspian sea drainages, including large Ponto-Caspian rivers, lagoons (limans), and dam reservoirs where it was intentionally introduced (Jażdżewski 1980). Its first record outside the Ponto-Caspian region was from the Vistula River between Warsaw and Tczew (Jarocki and Demianowicz 1931). Jażdżewski (1975) found this species in the Vistula mouth. A very interesting locality is also Figure 3. Distribution and possible migration routes in Poland of the Licheńskie Lake, an artificially heated water Chelicorophium curvispinum. Legend: (1) migration route and body belonging to a power plant cooling system. approximate dates, (2) river basin boundaries, (3) country borders, (4) artificial canals, C.L. – Curonian Lagoon, V.L. – Vistula The species was found there by Jażdżewski and Lagoon, S.L. – Szczecin Lagoon. Konopacka (1990), Konopacka and Jesionowska (1995) and it still occurs in the lake (unpublished data). Samples collected in 1998 by Jażdżewski and Konopacka (2000) confirmed the occurrence of this species in the middle and lower Vistula and in its largest tributary – Bug. However, although it still occurred there, C. ischnus was not common and was evidently outnumbered by new Ponto-Caspian gammarid invaders. This is possibly an example of an invader, whose population had stabilised after an initial phase of rapid population growth. Interestingly, the species has been recently observed more frequently in both previous and new localities. Jażdżewski (2003) found an abundant population inhabiting the Narew River near Nowogród. The species co-occurred there with another alien D. haemobaphes and two native species: Gammarus fossarum Koch, in Panzer, 1836, and G. varsoviensis Jażdżewski, 1975. In 2001, C. Figure 4. Distribution and possible migration routes in Poland of Chaetogammarus ischnus. Legend as in Figure 1. ischnus was also found in the Noteć River (unpublished data). The river is an important element of the so called central migration corridor (Bij de Vaate et al. 2002). The river shells offer a good solid base for building silt- empties to the Warta River, which is a tributary tubes, which are inhabited by this amphipod. of the Oder River, but at the same time Noteć is Colonisation success of this species in the Rhine connected to Vistula through the Bydgoski canal was discussed by den Hartog et al. (1992), and (Bij de Vaate et al. 2002, Grabowski et al. 2005). explained by its opportunistic characters, rapid However, our very recent findings (unpublished growth, early maturity, ability to produce several data) have not indicated the presence of C. generations per year, and high fecundity. As ischnus in the Oder, but this species has been shown recently, the species is now one of the previously found further west in the 1970s, in most important primary consumers, and is an canals joining the Elbe, Weser and Ems (Herhaus important food source for fish in the above river, 1978, Herbst 1982), and later in some Mecklen- determining its functional diversity and food web burgian lakes (Waterstraat and Köhn 1989). structure (Van Riel at al. 2006). Distribution and Quite recently, the species has also become an

27 M. Grabowski et al.

species invaded the Vistula and Oder deltaic systems from Lithuanian waters, through the Pregel (Pregoła) river system connecting the Kuronskij and the Vistula Lagoon (Jażdżewski et al. 2002; Grabowski et al. 2003). In addition, it could have migrated through the mesohaline coastal Baltic waters. Our findings (unpublished data) indicate also that the species prefers lentic parts of large rivers of very slow current or stagnant water bodies rich in nutrients (e.g. Zegrzyński Reservoir, Włocławski Reservoir, Szczecin Lagoon, Vistula Lagoon), where it is

usually found in higher densities than in running Figure 5. Pontogammarus robustoides. Photo by Michał water sections. A very recent and surprising Grabowski. finding of this species comes from the Lucieńskie Lake located in the Vistula valley (Grabowski and Bącela 2005). The lake is a invader in the Great Lakes in North America channel valley lake of mesotrophic character – (Witt et al. 1997). Life history of the species in until now P. robustoides was usually found in Central and Western Europe was studied by large, warm, eutrophic rivers, lagoons or dam Konopacka and Jesionowska (1995) as well as by reservoirs. In the lake, this species co-occurs Kley and Maier (2005). Distribution and with an alien D. haemobaphes and native probable migration routes of this species in amphipod Gammarus lacustris G.O. Sars, 1863. Poland are presented in Figure 4. Most probably the species was accidentally introduced to that lake with boat traffic from the Pontogammarus robustoides (G.O. Sars, 1894) Vistula River. In laboratory conditions, the (Figure 5) species appears to show some predatory behaviour, hunting for fly larvae (unpublished This species occurred originally in the lower data). Bącela and Konopacka (2005) studied the courses of large Ponto-Caspian rivers (e.g. life cycle of P. robustoides in Poland, and Volga, Don, Dnieper, Dniester and Danube), as showed that this species is one of the most well as in some freshwater and brackish coastal fecund amphipods (particularly if compared to lakes and Black Sea lagoons (Jażdżewski 1980). native species) inhabiting Polish waters. That, In the 1960s, it was successfully naturalised in combined with early sexual maturity, body size, many Ukrainian, Lithuanian and Caucasian lakes euryoeciousness and eurytopicity, may be one of and artificial reservoirs. In Lithuania it was also features enhancing the invasive potential of this introduced to the Neman drainage system, species. including the Curonian Lagoon. Thus, the Very recently Ovcharenko et al. (2006) found species reached the Baltic Sea basin (Gasjunas one species of parasitic microsporidia (Nosema 1968; Jażdżewski 1980; Arbaciauskas 2002). pontogammari Ovcharenko & Kurandina, 1987) Some years ago P. robustoides was also reported and two gregarines (Uradiophora ramosa from Mecklenburg (Rudolph 1997; Zettler 1998). Balcescu-Codreanu, 1974, Cephaloidophora Very recent records of this species come from mucronata Codreanu-Balcescu, 1995) of Ponto- Belarus, where it is present in Dnieper, wher it Caspian origin, infecting P. robustoides in Polish could migrate from Lithuania (Mastitsky and waters. Apparently the amphipod served as a Makarevich 2007). In Poland the species was vector for the above microparasites. At the found for the first time in 1988 in the Oder moment, there are no reports on possible transfer estuary – Szczecin Lagoon, by Gruszka (1999). of these pathogens to native species. Our most recent sampling (unpublished data) has Distribution and probable migration routes of also indicated the occurrence of P. robustoides this species in Poland are presented in Figure 6. in the lower Oder River. Further discoveries come from Konopacka (1998), Jażdżewski and Obesogammarus crassus (G.O. Sars, 1894) Konopacka (2000) and Jażdżewski et al. (2004), who found P. robustoides in the lower Vistula Original distribution of this species includes and in the Vistula Lagoon. Most probably this coastal Caspian Sea and lower courses of its

28 Alien Crustacea in Polish waters – Amphipoda

Figure 7. Distribution and possible migration routes in Poland of Figure 6. Distribution and possible migration routes in Poland of Obesogammarus crassus. Legend as in Figure 1 Pontogammarus robustoides. Legend as in Figure 1.

rivers. In the Black Sea it lives in lagoons, entering the lower sections of the Kuban, Don, Dnieper, Ingulec, Boh, Dniester and the Danube as far as Serbia (Dudich 1947; Jażdżewski 1980). As for P. robustoides, it was intentionally introduced and naturalised in the 1960s to the Neman river system in Lithuania, namely to the Kaunas Reservoir and from there transferred, both naturally and through human activity, to Curonian Lagoon (Gasjunas 1972; Arbaciauskas 2002). In 1999 and 2000, Konopacka and Jażdżewski (2002) found O. crassus in the littoral zone of the Vistula Lagoon and of the Dead Vistula (Martwa Wisła), both brackish, with salinities between 1 and 6 PSU. Our recent Figure 8. Dikerogammarus haemobaphes. Photo by Michał Grabowski. studies (Jażdżewski et al. 2004, 2005) prove occurrence of the species only in such oligohaline waterbodies in Poland. Evidently, the Dikerogammarus haemobaphes (Eichwald, 1841) species invaded there from Lithuanian waters, (Figure 8) most probably through the Pregel (Pregoła) river system connecting the Curonian and the Vistula This species occurs naturally in the lower and Lagoons. In addition, it could have migrated middle courses of the Black and Caspian Sea through the mesohaline coastal Baltic waters. basin rivers and brackish lagoons, down to the The latter is highly probable as in 2003 this Sea of Marmara. It is a species with a very wide species was found in the Szczecin Lagoon ecological tolerance, e.g. it occurs in salinities (Konopacka 2003), which is a brackish part of from freshwater up to 8 PSU (Ponomareva 1975) the Oder River delta. Soon after, Eggers and and in temperatures ranging from 6 to 30oC Anlauf (2005) recorded O. crassus in the Elbe (Kititsyna 1980). Accounting for the above River in Germany, which is now the westernmost features, Mordukhai-Boltovskoi (1964) predicted limit of its distribution in Europe. Distribution that it would soon invade sea basins beyond the and probable migration routes of this species in Ponto-Caspian area. Indeed, it invaded through Poland are presented in Figure 7. the southern corridor namely upstream through

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2007). Distribution and probable migration routes of this species in Poland are presented in Figure 9.

Dikerogammarus villosus (Sowinsky, 1894) This is a very recent gammarid invader to Polish waters. Similar to other Ponto-Caspian invaders it is naturally distributed in the lower courses of large rivers in the Black and Caspian Sea basins (Mordukhai-Boltovskoi 1969). The original dispersal route of D. villosus was the southern migration corridor (Bij de Vaate et al. 2002). Outside its natural range, it was first recorded from the upper Danube in 1992, outcompeting D. haemobaphes – the earlier invader (Nesemann et Figure 9. Distribution and possible migration routes in Poland of al. 1995). Soon after, it was recorded in high Dikerogammarus haemobaphes. Legend as in Figure 1. abundance in the lower Rhine River (Bij de Vaate and Klink 1995, Bij de Vaate et al. 2002). From there it moved eastwards through the Mittelland canal to the Elbe (Grabow et al. 1998, the Danube River (Nesemann et al. 1995). In the Zettler 1999, Rudolph 2000). From the Elbe, it 1950s, it was recorded as far as 1700km most probably dispersed through the above upstream from its mouth (Straškraba 1962). In mentioned Havel-Spree system, to the Oder 1993, Schleuter et al. (1994) observed the River where it was recorded for the first time species in the Main-Danube Canal, followed by downstream of the canal which connects this its entrance to the North Sea basin through the river to Havel-Spree system (Müller et al. 2001, Rhine River (Schöll et al. 1995). In Poland, D. Jażdżewski et al. 2002). At the moment this haemobaphes was found in 1997 by Konopacka species has spread almost along the entire stretch (1998) in the middle Vistula River. Further of the Oder within Polish borders (unpublished research made by our team in the middle and data), including the brackish Szczecin Lagoon in lower sections of the river showed that this the river’s delta (Jażdżewski et al. 2005). species was definitely a dominant gammarid, Surprisingly, in 2003 this species was also followed by two other Ponto-Caspian invaders: discovered in the Bug River (Konopacka 2004), abundant P. robustoides and scarce C. ischnus. where it migrated along the central migration In the Vistula River, upstream of Warsaw as far corridor, through the Pripet-Bug connection. as Sandomierz, it was the only gammarid present This is supported by the recent records of D. (Jażdżewski et al. 2002). We also found this villosus species in the Dnieper in Belarus species in the Vistula’s main tributary – Bug as (Mastitsky and Makarevich 2007). In the Bug well as in Noteć, Warta and Oder (Odra) River, this species occurs from this connection (Jażdżewski and Konopacka 2000, unpublished down to the Zegrzyński Reservoir, although it data). Furthermore, we have found the gammarid has not yet reached the Vistula River (unpub- to be abundant along the coasts of the Vistula lished data). In this case, we are seeing two Lagoon (Jażdżewski et al. 2004). At the same independent invasion events of this species into time Gruszka (2000) and Müller et al. (2001) Polish waters. reported it from the lowest sections of the Oder This species appears to be a very versatile River. Recently, Jażdżewski (2003) found it feeder, either acting as a filter-feeder exploiting along the Narew River invading the Masurian micro-algae (Platvoet et al. 2006) or as a very lakes in north-eastern Poland. The distribution of effective predator preying upon other D. haemobaphes in Polish waters clearly macroinvertebrates (MacNeil and Platvoet 2005; indicates that the species also uses the central Krisp and Maier 2005; Kley and Maier 2003; corridor to invade the Baltic and North Sea Devin et al. 2003; Dick and Platvoet 2000, Van basins (Bij de Vaate et al. 2002). The hypothesis Riel et al. 2006) and hunting for fish eggs or is also well supported by findings of this species juveniles (Casellato et al. 2005). Dick (1996) in Dnieper in Belarus (Mastitsky and Makarevich reports that this can be attributed to the species

30 Alien Crustacea in Polish waters – Amphipoda

Species of North American origin

Gammarus tigrinus Sexton, 1939 The invasion history of this species in Europe is one of the best documented so far. This gammarid occurs natively in brackish waters, along the Atlantic shores of North America from southern Labrador to Florida. It was reported to tolerate salinities between 1 and 25 PSU (Bousfield 1973). It is unknown when G. tigrinus was first introduced to Great Britain (Gledhill et al. 1976), but it was transplanted to the rivers Werra and Weser in Germany in 1957 (Schmitz 1960, Fries and Tesch 1965). Then, Bulnheim (1976) noted the occurrence of G. tigrinus in the Figure 10. Distribution and possible migration routes in Poland of Dikerogammarus villosus. Legend as in Figure 1. Baltic Sea drainage system (Schlei estuary). Simultaneously, this species dispersed into the Netherlands, probably from Iljsselmeer and soon occupied the brackish and fresh waters of this relatively large body size. Actually D. villosus region (Nijssen and Stock 1966, Dieleman and can have a large impact on local amphipod Pinkster 1977, Pinkster et al. 1977, 1980, 1992). populations. It was observed to exterminate Further eastward steps of G. tigrinus along the communities of the alien G. tigrinus in the Baltic Sea shores and inland localities in NE Rhine, and native Gammarus duebeni Liljeborg, Germany were documented by Bulnheim (1985), 1852 in Lake Ijsselmeer (Kelleher et al. 1999, Rudolph (1994 a, b, 2000) and Zettler (1995, Dick and Platvoet 2000). Similarly, in Germany 1998). In Poland, this species was first recorded the former invader D. haemobaphes was almost by Gruszka (1995, 1999) in the Szczecin Lagoon eliminated after colonisation by D. villosus (Kley in 1988. Our recent survey of the entire Oder and Maier 2003). Laboratory experiments by River flow (unpublished data) proved that G. MacNeil and Platvoet (2005) also showed a tigrinus entered this river upstream as high as the severe predatory impact of this species upon city of Kędzierzyn-Koźle, where it occurs in native G. pulex (Linnaeus, 1758). The same freshwater conditions. Thus this species has pattern was observed very recently in the Odra considerably extended its European range south- river, where the former coloniser, G. tigrinus, is eastwards. In the years 1998-2000, we found this now being replaced by D. villosus (unpublished species dominating some sections of the Vistula data). In the Rhine River it has become one of River mouth, including the Dead Vistula and the the key secondary consumers occupying high Vistula Lagoon (Jażdżewski and Konopacka trophic levels comparable to fish (Van Riel et al. 2000, Jażdżewski et al. 2002, 2004, Grabowski 2006). For the above reasons, it is of key et al. 2006). Interestingly, in the Vistula deltaic importance to monitor further expansion and system this species distribution was limited naturalisation processes of this species in Polish exclusively to its brackishwater parts. Also our waters. Life history of D. villosus in Western survey of coastal Malacostraca performed in Europe was previously studied by Kley and 2004 along the entire stretch of the Polish Baltic Maier (2003), Piscart et al. (2003), Devin et al. shoreline, revealed that G. tigrinus is present (2004) and by Pöckl (2007). Generally, this along the entire studied area, occurring both in particular invader may have an extremely coastal Baltic waters and in the estuaries of small deteriorative effect on local macroinvertebrate rivers, however, it did not disperse further and fish populations and due to its high fertility upstream (Jażdżewski et al. 2005). Most and fast growth rate it may become a recently, G. tigrinus was found further east, cosmopolitan species that creates problems in along the Finnish coast to the Gulf of Finland many parts of the world (Devin et al. 2003). (Pienimäki et al. 2004), in the brackish Curonian Distribution and probable migration routes of Lagoon in Lithuania (Daunys and Zettler 2006) this species in Poland are presented in Figure 10. and in the Gulf of Riga (Kotta 2005).

31 M. Grabowski et al.

zaddachi, may breed only in low water temperatures having, in our latitudes, a reproductive break during the summer. In contrary, G. tigrinus, is able to reproduce throughout the year with the exception of the coldest months when water temperatures are close to 0ºC. Also, the natives cease reproduction in lowered salinities, which is not the case for G. tigrinus. All that, combined with higher fecundity, more generations per year and the ability to thrive in heavily polluted waters makes this species one of the most successful amphipod invaders in Europe (Pinkster et al. 1977). Distribution and probable migration routes of this species in Poland are presented in Figure 11. Figure 11. Distribution and possible migration routes in Poland of Gammarus tigrinus. Legend as in Figure 1. Species of Balkan origin

Gammarus roeselii Gervais, 1835 Gammarus tigrinus is well known for its high invasive potential. In most colonised habitats, This species is often treated as a native to our the species has replaced native invertebrates, fauna, but has been recognised as a species of including other gammarid species. In Watch Balkan origin (Karaman and Pinkster 1977, Lane Flash (UK) it has eliminated the native G. Jażdżewski 1980, Jażdżewski and Roux 1988). duebeni (Savage 1982). In Northern Ireland, this Occurring naturally in the freshwaters of the species was shown to predate upon native Balkan, Peloponesian and Anatolian peninsulas, possum shrimp, Mysis relicta Loven, 1862, it entered (possibly starting from XIX century) however juvenile G. tigrinus were also under western and northern Europe through the predation pressure by the mysid (Bailey et al. southern invasion corridor, namely the Danube 2006). In Germany this species has eliminated system. The Rhine and Danube systems were many other macroinvertebrates (Fries and Tesch first connected by the Ludwigskanal in 1845, 1965). In the Netherlands, it was recorded that however G. roeselii was already recorded and besides replacing native G. duebeni and G. described from the vicinity of Paris in 1835 zaddachi Sexton, 1912, it was even injuring (Karaman and Pinkster 1977). Therefore, trapped fish (Pinkster et al. 1977). In Poland, Jażdżewski (1980) suggests that this species G. tigrinus successfully colonized oligohaline could have crossed the watershed with Baltic lagoons, eventually replacing not only unintentional human help, e.g. with aquatic native brackishwater gammarids, G. duebeni and plants. Currently it is distributed throughout the G. zaddachi, but also dominating other aliens: whole southern, middle and western continental P. robustoides, O. crassus and D. haemobaphes Europe, excluding the Apennine and Iberian (Grabowski et al. 2006). In Puck Bay, the peninsulas and Atlantic coasts. G. roeselii could species is abundant and dominates the coastal have reached the territory of Poland in two zone, while in the deeper parts native species are different ways. Firstly, from the west using the still the main component of the amphipod Oder-Spree and Oder-Havel canal systems, community. Similarly, along the Baltic coast, secondly – from the south, crossing a rather low G. tigrinus dominates in estuaries and coastal watershed between Oder and Morava (the lakes, while in more saline Baltic waters it is Danube tributary) in the Czech Republic present in lower densities (Jażdżewski et al. (Straškraba 1958). The distribution of this 2005). As discussed by Savage (1982) and by species in Polish waters is rather well known. It Pinkster et al. (1977) the potential replacement occurs in freshwaters – small and bigger rivers mechanism in the case of G. tigrinus and the of slow current, in lakes and in artificial canals native species may be based on a higher mainly in western Poland (Jażdżewski and reproductive capacity of the alien with relation Konopacka 1995). Its localities in the lower to temperature. Both natives, G. duebeni and G. Vistula system (affluents of Elbląg Canal) are at

32 Alien Crustacea in Polish waters – Amphipoda

indigenous gammarids, b) predation pressure from the newcomers, and c) greater reproductive capacity of the alien species. Among the interactions between the alien invaders and local species, trophic relationships should be particularly studied as they may alter the functional structure of entire ecosystems. Of the above invaders, one – D. villosus – is known to be a voracious predator exterminating local fauna, including other gammarids, in newly invaded areas. We also observed predatory behaviour of P. robustoides in laboratory conditions (unpublished data). These two species, taking into account their size, can become lethal for the local benthic communities in Polish waters. Predatory behaviour was also shown in the case Figure 12. Distribution and possible migration routes in Poland of G. tigrinus. From the other side, the new- of Gammarus roeselii. Legend as in figure 1. Unknown date of comers can become an important food source for colonization. local species. Even the same species can be both: a predator and a sought after prey item, which is true for both mentioned pontogammarids. In the most northeastern limits of its distribution in many artificial reservoirs in the Ukraine and Europe (Jażdżewski 1980). Lithuania, these alien species have been intro- The species does not spread quickly and does duced as a productive food source for com- not exhibit a high invasive potential. Pöckl mercial fishes. We have frequently observed C. (1993, 2003) studied the reproduction of G. curvispinum, P. robustoides and D. haemo- roeselii and its ability to compete with baphes individuals in fish stomachs; interesting- Gammarus fossarum. Apparently the former ly these species are an important food source for prefers warmer waters, where it may mature Ponto-Caspian fish invaders: racer goby earlier and give larger broods, and hence is able (Neogobius gymnotrachelus (Kessler, 1857)) and to dominate the latter, which performs better in monkey goby (Neogobius fluviatilis (Pallas, colder temperatures. Generally, G. roeselii may 1814)) which commonly occur in the Bug and be defined as an old alien coloniser, well Vistula rivers (Kostrzewa and Grabowski 2003; established but not invasive. Grabowska and Grabowski 2005, unpublished Distribution and probable migration routes of data). This brings another possible threat for this species in Poland are presented in Figure 12. local fauna – an infection by alien plagues or parasites, e.g. P. robustoides is known to be a vector for one species of microsporidia and two Conclusions gregarines coming from the Ponto-Caspian region (Ovcharenko et al. 2006). In summary, the alien amphipods constitute a Greater reproductive capacity of alien species significant part of the benthic communities in compared to natives has been proven for almost large Polish rivers and in coastal Baltic waters. all alien amphipod species in European fresh and In lower sections of rivers such as the Bug, brackish waters. Combined with high tolerance Vistula, Oder, and also in Szczecin and Vistula to warm water temperatures, heavy pollution and lagoons, the amphipod fauna is now limited the above mentioned predatory behaviour, it exclusively to alien species (Jażdżewski 2003; makes the alien amphipods extremely successful Jażdżewski et al. 2002, 2004, 2005; Grabowski invaders. et al. 2006, unpublished data). The retreat of Another effect of invasion is the formation of native species is a serious problem in many new biotic communities in local habitats – an places colonised by alien amphipods not only in “Invasional Meltdown” phenomenon. It occurs Poland but across Europe. The possible reasons when species co-evolving in their native range behind this phenomenon include: a) pollution of facilitate the invasion through mutual interaction aquatic ecosystems which are intolerable to the of newly colonized areas. A good example may

33 M. Grabowski et al. be the Ponto-Caspian C. curvispinum using References colonies of the bivalve D. polymorpha (of the same geographic origin) as a substrate for Arbaciauskas K (2002) Ponto-Caspian amphipods and mysids in building silt-tubes (Jażdżewski and Konopacka the inland waters in Lithuania: History of introduction, current distribution and relations with native malacostracans. 2002), when other suitable substrates are In: Leppäkoski E, Gollasch S, Olenin S (eds), Invasive lacking. The same relationship may be observed Aquatic Species of Europe – Distribution, Impacts and in both species’ native area of occurrence. Management, Kluwer Academic Publishers, Dordrecht, pp We can expect further successful invasions, 104-115 Bailey RJE, Dick JTA, Elwood RW, MacNeil C (2006) Predatory particularly from the Ponto-Caspian region. The interactions between the invasive amphipod Gammarus first reason for this is the present lack of tigrinus and the native opossum shrimp Mysis relicta. isolation between this basin and the Baltic Journal of the North American Benthological Society 25(2): drainage system. Secondly, there are numerous 393-405, http://dx.doi.org/10.1899/0887-3593(2006)25[393:PIB TIA]2.0.CO;2 naturalisations of Ponto-Caspian species in Barnard JL, Barnard CM (1983) Freshwater Amphipoda of the former Soviet Union countries, close to Polish world. Hayfield Association, Mt. Vernon, Virginia, 2 vols: 1- territory, which may facilitate the expansion 830 Bącela K, Konopacka A (2005) The life history of ofthe alien species by establishing new source Pontogammarus robustoides, an alien amphipod species in populations. Thirdly, a bit more complex, is a set Polish waters. Journal of Crustacean Biology 25(2): 190-195, of physio-ecological features shared by the http://dx.doi.org/10.1651/C-2519 species coming from the Ponto-Caspian region. Bij de Vaate A, Klink AG (1995) Dikerogammarus villosus Sowinsky (Crustacea: Gammaridae) a new immigrant in the All these species are euryoecious and rather Dutch part of the Lower Rhine. Lauterbornia 20: 51-54 euryhaline, occurring from freshwater to, in Bij de Vaate A, Jażdżewski K, Ketelaars HAM, Gollasch S, Van some cases, mesohaline conditions (Bij de Vaate der Velde G (2002) Geographical patterns in range extension et al. 2002; Konopacka 2004). Besides of Ponto-Caspian macroinvertebrate species in Europe. Canadian Journal of Fisheries and Aquatic Sciences 59(7): crustaceans, successful invasions in European 1159-1174, http://dx.doi.org/10.1139/f02-098 waters are also known for Ponto-Caspian gobiid Bousfield EL (1973) Shallow-water Gammaridean Amphipoda of fishes, molluscs, triclads, hydrozoans, leeches New England. Cornell University Press, Ithaca, pp 1-312 and water-mites (Bij de Vaate et al. 2002; Bousfield EL (1977) A new look at the systematics of Gammaridean amphipods of the world. Crustaceana Jażdżewski and Konopacka 2002). It is presumed Supplement 4: 282-316 that the next Ponto-Caspian amphipod to enter Bousfield EL (2001) An updated commentary on phyletic Polish waters may be Chaetogammarus classification of the amphipod Crustacea and its applicability warpachowskii (G.O. Sars, 1894), which occurs to the North American fauna. Amphipacifica 3(1): 49-113 Bousfield EL, Hoover PM (1997) The amphipod superfamily in the Curonian Lagoon (Konopacka 2004). It is Corophioidae on the Pacific Coast of North America. Part V. likely that it may follow the route already taken Family Corophiidae, new subfamily. Systematics and by P. robustoides and O. crassus. distributional ecology. Amphipacifica 2(3): 67-139 To conclude, the amphipod fauna of Poland Bulnheim HP (1976) Gammarus tigrinus, ein neues Faunenelement der Ostseeförde Schlei. Schriften des has undergone a very dynamic change. However, Naturwissenschaftlichen Vereins für Schleswig-Holstein: 46- the exact nature of the processes, particularly the 84 biological features of each species, physiological Bulnheim HP (1985) Genetic differentiation between natural tolerance to environmental stress and trophic populations of Gammarus tigrinus (Crustacea, Amphipoda) with reference to its range extension in European continental relationships with local species remain largely waters. Archiv für Hydrobiologie 102: 273-290 unknown. It will, therefore, be a great challenge Casellato S, La Piana G, Latella L, Ruffo S (2005) for scholars in the near future. Dikerogammarus villosus (Sowinsky, 1894), a new invasive species in the Garda Lake (Northern Italy). Abstracts for Biological Invasions in Inland Waters (INWAT) Workshop, Acknowledgements May 5–7, 2005, p 26 Crawford GI (1935) Corophium curvispinum G.O. Sars var. devium Wundsch, in England. Nature 136: 685, http://dx.doi. The data presented in this paper come from activity supported org/10.1038/136685c0 financially by the Polish State Committee of Scientific Research Daunys D, Zettler ML (2006) Invasion of the North-American (KBN), grant No. 2 P04G 076 026 p01, as well as from the amphipod (Gammarus tigirnus Sexton, 1939) into the internal grants and funds from the University of Łódź. Authors Curonian Lagoon, south-eastern Baltic Sea. Acta Zoologica are indebted to Ewa Janowska and Karolina Bącela for their Lituanica 16(1): 20-26 extensive help in the fieldwork. Many thanks are due also to Dr Den Hartog C, Van den Brink FWB, Van der Velde G (1992) Elizabeth Cook, Dunstaffnage Marine Labora-tory, Oban, UK for Why was the invasion of the river Rhine by Corophium improving the language of this paper. curvispinum and Corbicula species so successful? Journal of Natural History 26: 1121-1129, http://dx.doi.org/10.1080/002 22939200770651

34 Alien Crustacea in Polish waters – Amphipoda

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Pöckl M (2007) Strategies of a successful new invader in Van den Brink FWB, Van der Velde G, Bij de Vaate A (1993) European fresh waters: fecundity and reproductive Ecological aspects, explosive range extension and impact potential of the Ponto-Caspian amphipod Dikerogamma- of a mass invader, Corophium curvispinum Sars, 1895 rus villosus in the Austrian Danube, compared with the (Crustacea: Amphipoda), in the Lower Rhine (The indigenous Gammarus fossarum and G. roeseli. Netherlands). Oecologia 93: 224-232, http://dx.doi.org/10. Freshwater Biology 52: 50-63, http://dx.doi.org/10.1111/ 1007/BF00317675 j.1365-2427.2006.01671.x Van der Velde G, Rajagopal S, Musko I, Bij de Vaate A Razowski J, ed (1997) Checklist of Animals of Poland. (2000) Ecological impact of crustacean invaders. General Volume IV. Wydawnictwa Instytutu Systematyki i considerations and examples from the Rhine River. In: Ewolucji Zwierząt PAN, Kraków: 1-303 von Vaupel Klein JC, Schram FR (eds), The biodiversity Rudolph K (1994a) Erstnachweis des Amphipoden Gammarus crisis and Crustacea, Proceedings of 4th International tigrinus Sexton, 1939 (Crustacea: Gammaridea) im Crustacean Congress, Brill, Leiden, Crustacean Issues 12, Peenestrom und Achterwasser (südliche Ostseeküste). pp 3-33 Naturschutzarbeit im Macklenburg-Vorpommern 37: 23- Van Riel MC, Van der Velde G, Rajagopal S, Marguillier S, 29 Dehairs F, A bij de Vaate (2006) Trophic relationships in Rudolph K (1994b) Funde des Amphipoden Gammarus the Rhine food web during invasion and after tigrinus Sexton, 1939 in zwei Havelseen der Region establishment of the Ponto-Caspian invader Berlin/Brandenburg (Crustacea: Amphipoda: Gam- Dikerogammarus villosus. Hydrobiologia 565: 39-58, maridae). Faunistische Abhandlungen Staatliches http://dx.doi.org/10.1007/s10750-005-1904-8 Museum für Tierkunde, Dresden 19: 129-133 Waterstraat A, Köhn J (1989) Ein Beitrag zur Fauna des Rudolph K (1997) Zum Vorkommen des Amphipoden Kummerower Sees, Erstnachweis des Amphipoden Pontogammarus robustoides Sars, 1894 im Peenemün- Echinogammarus ischnus Stebbing, 1899, in der DDR. dungsgebiet. Natur und Museum 127: 306-312 Archiv der Freunde der Naturgeschichte in Mecklenburg Rudolph K (2000) Gebietsfremde malakostrake Krebse in 29: 93-106 mittleren Teil Brandenburgs. Aktueller Stand der Witt JDS, Hebert PDN, Morton WB (1997) Echinogammarus Verbreitung. Neozoen 3: 10-11 ischnus: another crustacean invader in the Laurentian Savage AA (1982) The survival and growth of Gammarus Great Lakes basin. Canadian Journal of Fishersies and tigrinus Sexton (Crustacea: Amphipoda) in relation to Aquatic Sciences 54:264-268, http://dx.doi. org/10.1139/f96- salinity and temperature. Hydrobiologia 94: 201-212, 292 http://dx.doi.org/10.1007/BF00016400 Wolski T (1930) Corophium curvispinum G.O. Sars in der Schleuter M, Schleuter A, Potel S, Banning M (1994) Prypeć und in der Warschauer Wasserleitungsanlagen. Dikerogammarus haemobaphes (Eichwald 1841) Fragmenta Faunistica Musei Zoologici Poloniae 1: 152- (Gammaridae) aus der Donau erreicht über den Main- 159 Donau-Kanal den Main. Lauterbornia 19: 155-159 Wundsch HH (1912) Eine neue Species des Genus Corophium Schmitz W (1960) Die Einbürgerung von Gammarus tigrinus Latr. aus dem Müggelsee bei Berlin. Zoologische Sexton auf dem europäischen Kontinent. Archiv für Anzeiger 39: 729-738 Hydrobiologie 57: 223-225 Zettler ML (1995) Erstmachweis von Gammarus tigrinus Schöll F, Becker C, Tittizer T (1995) Das Makrozoo-benthos Sexton, 1939 (Crustacea: Amphipoda) in der Daß- des schiffbaren Rheins von Basel bis Emmerich 1986- Zingster Boddenkette und seine derzeitige Verbreitung an 1995. Lauterbornia 21: 115-137 der deutschen Ostseeküste. Archiv der Freunde der Stock JH (1974) The systematics of certain Ponto-Caspian Naturgeschichte in Mecklenburg 34: 123-140 Gammaridae (Crustacea, Amphipoda). Mitteilungen aus Zettler ML (1998) Zur Verbreitung der Malacostraca dem Hamburgischen Zoologischen Museum und Institut (Crustacea) in den Binnen- und Küstengewässer von 70: 75-95 Mecklenburg-Vorpommern. Lauterbornia 32: 49 Straškraba M (1958) Beitrag zur Kenntnis der Verbreitung der Zettler ML (1999) Erstnachweis von Dikerogammarus Amphipoden in der Tschechoslowakei aus dem zoogeo- villosus (Sovinski, 1894) und Wiederfund von Gammarus graphischen Gesichtpunkt. Acta Universitatis Carolinae, varsoviensis Jażdżewski, 1975 in Mecklenburg- Biologica 2: 197-208 Vorpommern (Crustacea: Amphipoda). Archiv der Straškraba M (1962) Amphipoden der Tschechoslovakei nach Freunde der Naturgeschichte in Mecklenburg 38: 231-233 den Sammlungen von Prof. Hrabe I. Vestnik Cesko- slovenske Zoologicke Spolecnosti 26(2): 117-145 Van den Brink FWB, Van der Velde G and Bij de Vaate A (1989) A note on the immigration of Corophium curvispinum Sars, 1895 (Crustacea: Amphipoda) into the Netherlands via the River Rhine. Bulletin of the Zoological Museum of the University of Amsterdam 11: 211-213

37 M. Grabowski et al.

Annex . Systematics, origins and ecological preferences of alien Amphipoda occurring in Polish waters

Habitats used First record Systematic position Geographic origins in Poland in Poland

Subphylum: Crustacea Pennant, 1777 Class: Malacostraca Latreille, 1806 Order: Amphipoda Latreille, 1816 Family: Corophiidae Dana, 1849 Chelicorophium curvispinum (G.O. Sars, 1895) 1 PC F 1920’s Family: Gammaridae Leach, 1813 Gammarus roeselii Gervais, 1835 BL F XIX c. ? Gammarus tigrinus Sexton, 1939 NA F, B 1988 Chaetogammarus ischnus (Stebbing 1899) 2 PC F 1931 Family: Pontogammaridae Bousfield, 1977 3 Dikerogammarus haemobaphes (Eichwald, 1841) 4 PC F, B 1997 Dikerogammarus villosus (Sowinsky, 1894) PC F, B 2001 Pontogammarus robustoides G.O. Sars, 1894 PC F, B 1988 Obesogammarus crassus (G.O. Sars, 1894) 5 PC B 1999 Symbols used: Geographic origins: NA – North American, PC – Ponto-Caspian, BL – Balkan Habitats used in Poland: F – freshwater, B – brackishwater First record in Poland: ? – see explanation in text

Annotations by species name: 1 After the recent revision by Bousfield and Hoover (1997), the new generic name Chelicorophium for former Corophium curvispinum has been accepted 2 Formerly known as Echinogammarus ischnus, Chaetogammarus tenellus, C. tenellus behningi and C. ischnus sowinskyi. See Jażdżewski and Konopacka (2002) for detailed taxonomic discussion on this matter 3 A discussion on the taxonomy of Ponto-Caspian gammarid species was given by Stock (1974) and Barnard and Barnard (1983). The concept of the family Pontogammaridae, established by Bousfield (1977, 2001) is adopted here 4 The freshwater populations of this species in older literature are often referred to as D. haemobaphes fluviatilis or even as D. fluviatilis. However, after some morphological studies, Konopacka (1998) retained the name D. haemobaphes for specimens found in Polish waters. For details see Jażdżewski (1980) and Jażdżewski and Konopacka (1988) 5 Known in older literature as Pontogammarus crassus, this species was moved by Stock (1974), along with some other related Ponto- Caspian species, to the newly created genus Obesogammarus. For details see Konopacka and Jażdżewski (2002)