Contributions to Zoology, 71 (4) 115-122 (2002)

SPB Academic Publishing bv, The Hague

Four Ponto-Caspian and one American gammarid species (Crustacea,

Amphipoda) recently invading Polish waters

Jazdzewski & Michal Grabowski Krzysztof ,Alicja Konopacka Department ofInvertebrate Zoology and Hydrobiology, University ofLodz, 12/16Banacha str., 90 237 Lodz,

Poland

Keywords: Biogeography, gammarids, European waters, Poland

in the Abstract (Di Castri 1989) and, case of freshwater fauna,

by the construction of artificial waterways joining

in the The paper discusses recent drastic changes composition formerly separate river systems (Jazdzewski 1980).

of Polish gammarid fauna, that occurred at the end of 20th Central Europe is drained by the rivers discharging invasion offive alien century. This change was caused by the into the North Sea, Baltic Sea and Black Sea. The species - four ofPonto-Caspian origin(Dikerogammarus haemo- constructions of man-made canals joining these baphes, D. villosus, Obesogammarus crassus and Pontogam- different basins started in XVIIIth and marus robustoides) and one of American origin (Gammarus century during

tigrinus). Probable invasion routes are presented. two next centuries these three basins were inter-

connected in canals several ways. Major important

to be mentioned here are those connecting the Elbe

Contents and Oder (opened in 1746), the Oder and Vistula

(1774) and the Vistula and Dnieper systems (1784,

115 Abstract Bug - Pripet’ canal) (Jazdzewski 1980). Introduction 115 The area of Poland belongs nearly entirely to Review of recent immigrants 116 the Baltic Sea basin, and some 90% of this terri- The present state of Polish gammarid fauna 117 to the Oder and Vistula Discussion 119 tory belongs drainage sys-

tems. the minor Acknowledgements 121 Only part of northern Poland is

121 References drained by smaller rivers emptying directly into the

Baltic Sea.

Until recently gammarid species were usually Introduction recognised as members of the extra-large family

Gammaridae. Amphipod genera ascribed to this

of and Geographical ranges plants change family sensu Stebbing (1906) appeared to be really

permanently in time, however these changes, when not related to each other, however this old familial

natural, are slow from the human point of view. arrangement, especially of freshwater, Holarctic

Even the formation of the Stock geologically young present , persisted very long. (1968, 1974), the last North-European fauna, that has begun after Karaman (1977), Bousficld (1977, 2001) and Bar-

Wiirm and continues till now for some nard & Barnard made the glaciation (1983) attempts to group 15 thousands is of years, from our perspective a phyletically related „gammarid” taxa using much rather The of civi- long process. rapid development more detailed morphological analysis.

lisation, and of human intercontinental from the fundamental especially Starting paper by Sars

some 5 centuries have migrations commencing ago, (1894-1895) the rich, originally Ponto-Caspian am- accelerated seriously biogeographical changes by was phipod fauna usually considered to form a part intentional acclimatisations, accidental introductions of the family sensu lato (i.a. Carausu

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and et at. 1955, Birshtejn & Romanova 1968, Mor- list of Polish gainmarids with 5 alien species,

dukhaj-Boltovskoj et al. 1969), although already to recognise their most probable invasion routes.

Martynov (1924) commented some special morpho-

of the logical features of many “gammarids” so

called Ponto-Caspian complex - the features that Review of recent immigrants

he has named “pontogammarisation”. Stock (1974)

named “ has distinguished a group Dikerogamma- Dikerogammarus haemobaphes (Eichwald, 1841) such rus-Pontogammarus complex” encompassing For the first time in the Baltic Sea basin, the spe-

Steno- cies recorded in Poland in 1997 genera as Dikerogammarus, Pontogammarus, was (Konopacka

and several others as well The of this gammarus,Niphargoides 1998). range expansion Ponto-Caspian

This as a newly erected Obesogammarus. group invader in European waters and in Poland was al-

& was formally named“family ” by ready presented by Jazdzewski Konopacka (2000).

D. Bousfield (1977) who has distinguished tentatively Recently, we discovered large population of

10 family-groups or families (i.a. Gammaridae, haemobaphes in the Vistula Lagoon, Gruszka (2000)

Pontogammaridae, Anisogammaridae etc.) form- and Muller et al. (2001) found this species in the

ing together the superfamily . Barnard lower Oder river.

& Barnard (1983) have put Gammarus-like genera

in the “Gammaroid-group” (that could be possibly Dikerogammarus villosus (Sowinsky, 1894)

in this Oder situated at the supra-familial level); group The species was recently recorded in the river,

these authors formally distinguished family Gamma- in 1999 (Gruszka 2001, Muller et al. 2001, Jazd-

ridae with genus Gammarus and several Caspian zewski and Konopacka 2002) downstream of the

of and Baikalian genera, and 13 groups, possibly canal connecting the Oder river with the Elbe ba-

such Echino- into the Oder familial level with master genera as sin. The penetration of D. villosus

and since it has used first gammarus,Dikerogammarus Pontogammams, basin is especially interesting

others. Until solid cladistic of so-called southern i.e. Danube among any analysis the corridor, river,

the all “gammarid” generais done, we follow Bous- for westwardrange expansion (Bij de Vaate et al.,

classification of Ponto-Cas- reach Danuberiver villosus field’s (1977, 2001) 2002). In the upper of D.

in in pian gammarids the family Pontogammaridae, was first recorded by Tittizer et al. (1994) 1992,

the and it has into the Rhine river via leaving genera Gammarus, Echinogammarus soon penetrated

and Chaetogammarus in the family Gammaridae the Main - Danube canal (Bij de Vaate & Klink

s.str., and retaining the widely used vernacular name 1995). From the Rhine river D. villosus continued

for both the Gammaroidea eastward the Mittelland- “gammarid” superfamily range expansion by using

the sensu Bousfield (1977) and a “Gammaroid-group” canal joining Rhine, Weser, Elbe and Oderbasins

& sensu Barnard Barnard (1983). (Grabow et al. 1998, Zettler 1998, Rudolph 2000).

Our department has studied for several decades In the lower Oder river, the species co-occurrs with

the gammarid fauna of Poland. We have collected other alien gammarids, like D. haemobaphes, P.

and determined altogether some 60,000 individu- robustoides and G. tigrinus (Miiller et al. 2001,

in 1200 taken in the whole als about samples country. own unpubl. data).

1990 The data obtained till were summarised in

two monographs by Jazdzewski (1975) and Jazd- Pontogammarus robustoides (G.O. Sars, 1894)

zewski & Konopacka (1995). At that time Polish First records of P. robustoides come from north-

15 this gammarid fauna encompassed gammarid spe- western Poland. Gruszka (1999) found species

cies. in the Szczecin Lagoon and the lower Oder river in

Results of our recent studies (1997-2001), fo- 1988. The species was also reported from the lower

than 100 col- from cusing on large rivers (more samples Vistula river (Konopacka 1998) and the Vistula

lected, altogether around 3000 specimens), together Lagoon (Jazdzewski & Konopacka 2000). Ponto-

Gruszka Vistula and Oder with the results obtained by (1995, 1999, gammamsrobustoides reached the

2001) in the Oder estuary, allowed to enrich the deltaic systems possibly with ballast waters via

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Baltic Sea from the Neman river and Curo- localities of the in system species the Vistula Lagoon are

nian Lagoon, where it was introduced in the 1960s by now the easternmost ones in Europe. (Gasjunas 1972, Arbaciauskas, 2002). However, the

species could also have entered the Vistula Lagoon

through the river the Pregola system connecting The present state of Polish gammarid fauna

Vistula river delta with the Curonian Lagoon. More

details on the origins and distribution routes of the The gammarid fauna of Poland is comparatively mentioned species in Europe can be found in well known (Jazdzewski & Konopacka 1995, Grus- Jazdzewski & Konopacka (2000). zka 1995, 1999, Konopacka 1998, Konopacka &

Jazdzewski 2002). Native freshwater taxa are

Obesogammarus crassus (G.O. Sars, 1894) Gammarus pulex, G. fossarum, G. lacustris, G. This species is most recently discovered in Polish varsoviensis, G. leopoliensis and G. balcanicus, waters, namely in the Vistula Lagoon and in the whereas the Baltic autochthonous species are G. Dead Vistula in 1998 & (Konopacka Jazdzewski, zaddachi, G. salinus, G. duebeni, G. locusta, G.

2002). Original distribution areas of O. crassus en- inaequicauda, G. oceanians and Chaetogammarus

compassed offshore Caspian Sea waters and lower stoerensis. Gammarus roeselii was recognized by of courses rivers emptying to this water body; in & Jazdzewski Roux (1988) as a species of Balkan the river the Volga species penetrated as upstream origin, possibly recently (in XIX century?) enter- far as to Volgograd (Mordukhaj-Boltovskoj 1979). ing western and northern Europe via the Danube In the Black Sea O. occurred system crassus origi- system. It has possibly used the Danube-Rheincanal

nally in brackish lagoons and in the lower courses and Mittelland-canal in its westward and then east- of large rivers (Dedju 1980, Jazdzewski 1980). In ward range expansion. Until recently the only evi-

the Danube river it was noted as far as upstream, dently alien gammarid species was Chaetogammarus in its sector (Dudich 1967). Like P. Yugoslavian ischnus, discovered in the Vistula river in 1928

robustoides, O. crassus was transplanted in early & (Jarocki Demianowicz 1931). This species has 1960s into the Kaunas artificial reservoir on the used the canal surely Bug-Prypet’ for range exten- Neman river in Lithuania and from there, after sion from the Dnieper system westward. In the last acclimatisation, it entered the Curonian Lagoon decades we face an increasing number of invasions

(Gasjunas 1972, Arbaciauskas, 2002). Subsequently new of alien species in Polish waters (Fig. 1). In it entered the Vistula Lagoon, most via probably quite a short time Polish gammarid fauna has been the Pregel river system. However, due to its com- enriched by 5 species: four of Ponto-Caspian ori- paratively high euryhalinity, O. crassus could have gin (Dikerogammarus haemobaphes, D. villosus, also dispersed south-westwards along the Baltic Sea Pontogammarus robustoides and Obesogammarus shores, with the salinity of 7 PSU in this average crassus) and one from North America (Gammarus region. tigrinus). Two of these immigrants, G. tigrinus and

D. haemobaphes dominate now the , gammarid fauna Gammarus tigrinus Sexton, 1939 of lotic environments in the Oder and Vistula rivers, This North American euryhaline species was ob- respectively. In the Oder river G. tigrinus is the served in waters with salinities ranging from 1 to most common and widespread species, entering 25 PSU (Bousfield 1973). Information on its intro- upstream as far as nearly to the city of Opole, be- in duction and distribution routes Europe have been ing the only gammarid species there. In the lower summarised Jazdzewski & by Konopacka (2000). Oder its river, especially downstream of connec- First of observations G. tigrinus in Polish waters tion with - Spree Havel system and of the Warta 1988 in were done in the Szczecin Lagoon (Gruszka river mouth, Dikerogammarus haemobaphes joins 1995, 1999, Wawrzyniak-Wydrowska & Gruszka G. tigrinus, and, according to Muller et al. (2001) Recent of 2001). survey the entire Oder river could (un- be even a dominant in mixed gammarid po- published data) proved that G. tigrinus entered this pulations. On the other hand D. haemobaphes has river far to upstream as as the city of Opole. The the entire Vistula conquered nearly river, occurring

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Fig. I. Changes in gammarid fauna ofPolish waters. Place of first record indicated in brackets. PC - Ponto-Caspian species. NA -

North American species.

usually as an only gammarid species as far upstream the gammarid fauna was dominated by either Obe-

km below Cracow. In the lentic and G. R as about 100 condi- sogammarus crassus tigrinus or robust-

in tions of the artificial Wloclawek reservoir the oides and O. crassus. In both cases a small admixture middle/lower Vistula section, D. haemobaphes of Gammarus duebeni occurred. D. haemobaphes was outcompeted by Pontogammarus robustoides. also occurred in numbers in the less saline (P-

Downstream of this reservoir the more rheophilous oligohaline) part of the Lagoon influenced by the

D. haemobaphes regained its dominance. Nogat arm of the Vistula. The gammarids occurring

- northern the Populations ofnative gammarid species Gam- along the shores of Vistula Lagoon were

G. G. marus pulex, G. fossarum, G. varsoviensis, and the mostly dominated by duebeni or by tigrinus, earlier immigrant G. roeselii - were only recorded other species found were P. robustoides, Gammarus in some tributaries. zaddachi and O. crassus. In brackish water of the

In the P-oligohaline wafers of the Szczecin La- former Vistula section, called the Dead Vistula

the 2-7 the and goon two alien species dominated gammarid (salinity PSU) most common usually fauna- G. tigrinus and P. robustoides (Wawrzyniak- dominantgammarid is G. tigrinus, most often accom-

Wydrowska & Gruszka 2001). In the mostly a- panied by G. zaddachi, sometimes by D. haemoba-

and oligohaline Vistula Lagoon, in its southern part, phes rarely by G. duebeni, which was, however,

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the dominant the station of freshwater species at least saline near sions or oligohaline species (e.g. the the dam Dead the separating Vistula from Vistula case of Chaetogammarus ischnus, Witt el al. 1997)

It these river. is worth while to note, that same such transport seems to be the major possibility. brackishwater bodies «- Vistula Lagoon and Dead However, in European waters, after breaking

- Vistula at least till the 1970s, were mainly inhab- physical barriers, migrations through canals and ited G. zaddachi and G. duebeni with Sea by varying along the brackish Baltic littoral waters were dominanceof another and a the one or species very rare most important way of range extensions. This

of Gammarus salinus and G. oceanicus presence at semi-natural penetration can occur within the same

the Sea the entrance ofDead Vistula to Baltic (Zmud- or adjacent biogeographical province of identical

Arndt and zinski 1957, 1965, Jazdzewski 1975, or similar climatic conditions.

Recent invasion of alien unpubl. observations). routes gammarid spe-

cies in Polish waters are illustrated by Fig. 2. This

scheme is based upon the distribution of new and

Discussion old records of alien gammarids along the Polish

in river courses given detail by Jazdzewski & Kono-

There several for are possibilities gammarid spe- packa (2002).

The invasion cies to extend their original distribution areas. Quite of many Ponto-Caspian species in

via natural way is their upstream migration, especially European freshwaters, and, freshwaters, into in large rivers. Segerstrale (1954) suggested that, brackish coastal waters of the Baltic and North Seas,

in is at least the case of Gammarus lacustris, the trans- related to their typically oligohaline preferences

birds could be for the wide and Most of these port by responsible relatively high euryhalinity. spe- distribution of this species in Holarctic. However, cies originally live in estuaries and lagoons of the

do Azov in most of the cases discussed we have to with Black and Seas of a low salinity (0.1-7 PSU

but various kinds of human impact. The construction mostly 0,5-5 PSU (Mordukhaj-Boltovskoj et

al. of canals connecting different drainage areas is one 1969, Dedju 1980)) as a relict fauna of the Sar- of of matian or Pontian fundamental reasons of the penetration par- Age. ticular species into sometimes distant regions. An- The endemic Ponto-Caspian fauna is

of other factor, often connected with the former one, probably a freshwater origin; at present various exhibit are intentional introductions of species aimed at species various grades of euryhalinity. In

the Sea the enrichment of fish food resources (Karpevich Caspian itself over 70 endemic malaco-

1975, Arbaciauskas 2002). In Europe the impact stracan species were recorded, of them some 15 of these both factors the extensions of in differ- upon range species (mostly amphipods) penetrated

ent distances various amphipod species were amply discussed upstream the Volga river - those being

the most by Jazdzewski (1980) and, more recently, various euryhaline taxa (Mordukhaj-Boltovskoj alien freshwater invertebrates penetration in west- & Dzjuban 1976). ern Europe as well as the ecological impact of these Gammarid species of the so-called Ponto-Cas- invaders were summarised, i.a., by Kinzelbach pian complex (see Mordukhaj-Boltovskoj 1964) in

do (1995), Tittizer (1996), Jazdzewski & Konopacka general not occur in the open (central) Baltic

(2000), Van der Velde et id. (2000), Tittizer et id. Sea of the surface salinity 7-8 PSU, or at least do

(2000) and Bij dc Vaate et id. (2002). not compete with native fauna. On the other hand

One such like should consider of course, also the pos- species, Chaetogammarus ischnus, Ponto-

sibility of introductions of alien gammarids, for gammarus robustoides, Dikerogammarus haemo-

D. villosus instance by the transfer ofaquatic plants; such possi- baphes, and Obesogammarus crassus bility was suggested for Gammarus roeselii by are found only in freshwaters or in oligohaline la-

Roux like Vistula Jazdzewski & (1988). goons Lagoon (salinity 2-5 PSU) or

The ballast water transport also cannot be ex- Szczecin Lagoon (0,5-1,5 PSU) and only there they cluded factor with with native as a accelerating gammarid range really may compete success fauna;

for extensions and, in the case of transatlantic inva- the same is true Curonian Lagoon, possibly for

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Fig. 2. Immigration routes of invasive gammarid species in Polish waters. I - Gammarus tigrinus, 2 - Pontogammarus robustoides,

3 - Dikerogammarus haemobaphes (and Chaetogammarus ischnus, except upstream of Warsaw), 4 - Obesogammarus crassus, - - - - - 5 D. villosus, 6 - basin boundaries, 7 country borders, 8 artificial waterways, V.L. Vistula Lagoon, S.L. Szczecin Lagoon,

C.L. - Curonian Lagoon.

Gulf of Riga and Gulf of Finland. Europe? (Jazdzewski & Konopacka 2000, Tittizer

Similar oligohaline preferences of Gammarus et al 2000, Bij de Vaate et al. 2002, and unpubl. tigrinus have been mentioned several times in the 2000/2001 observations). In Poland one of the rea-

be the ionic literature (Bulnheim 1976, Pinkster et al. 1992). sons can increasing content of large

the of the rivers in last caused the industrial Although first record invading spe- decades, by pol- cies be somewhat and the first lution & may delayed really (Dojlido Woyciechowska 1985, Szymanska

& in occurrence of a taxon in new place surely precedes 1990, Ficek Ficek 1994). This rise the salin- this first record, one can assume that this delay is ity” of such rivers like Vistula and Oder would

3-5 reach the several not longer than, say, years, taking into account finally „critical point” allowing still more and more detailed monitoring of Euro- species of oligohaline preferences to start their rather pean rivers. quick conquest of new basins. Obviously the in-

but An - interesting question arises why we do ob- creasing transport is also responsible, when serve this rather recent massive invasions of vari- looking for a „trigger” of these invasions and at- ous Ponto-Caspian species in central and western taining rich populations in the whole river flows in

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comparatively shortperiod the present authors would Bulnheim H-P. 1976. Gammarus tigrinus, ein neues Faunen-

element der Ostseetorde Schlei. Schr. Naturw. Ver. Schlesw.- rather favor the above mentioned hypothesis. Holst. 46: 79-84. Serious studies on the ecological impact of alien Carausu S, Dobreanu E, Manolache C. 1955. Amphipoda the native fauna in the Vistula and species upon forme si de dulce. salmastre apa Fauna Rep. popul. Romine, Oder still undertaken. systems are not Quantita- Crustacea 4: 1-409.

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Drastic changes in the amphipod fauna (Crustacea) of Dutch Received: 12 February 2002

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