NORTH-WESTERN JOURNAL OF ZOOLOGY 8 (1): 198-201 ©NwjZ, Oradea, Romania, 2012 Article No.: 121201 http://biozoojournals.3x.ro/nwjz/index.html

Gammarus varsoviensis Jazdzewski, 1975 (Amphipoda, Gammaridae): a long overlooked species in Ukrainian rivers

Michał GRABOWSKI1*, Tomasz MAMOS1, Tomasz REWICZ1, Karolina BACELA-SPYCHALSKA1 and Mykola OVCHARENKO2

1. Department of Invertebrate Zoology & Hydrobiology, University of Lodz, Banacha 12/16, 90-237 Lodz, Poland 2. Institute of Parasitology, Polish Academy of Sciences, Twarda 51/55, 00-818 Warszawa, Poland *Corresponding author, M. Grabowski, E-mail: [email protected]

Received: 30. August 2011 / Accepted: 13. December 2011 / Available online: 08. January 2012 / Printed: June 2012

Abstract. The first findings of Gammarus varsoviensis in are reported based on samples collected in 2009 and 2011. Until now, the species was known predominantly from the Baltic drainage area, with only one locality in the upper River in Belarus (Black Sea basin). Our findings extend the known geographic range of this species ca. 830 kilometers south-east, revealing its occurrence in typically lowland streams and rivers of the system, in the forest and Pontic steppe zone of the Black Sea basin. Based on the literature data we conclude that presence of this species in Ukraine was formerly overlooked through misidentification with morphologically close Gammarus lacustris. Based on such distribution pattern we can surmise that the species is native to the Black Sea drainage area. It could possibly have migrated to the Baltic basin through the Pripyat-Bug Canal, already in the XIX century.

Key words: Dnieper River, biogeography, Pontic steppe, freshwater, gammarids, Ukraine.

Amphipods are among the most diverse crusta- tive freshwater Gammarus in Ukrainian inland wa- cean groups inhabiting inland waters with 293 ters, Martynov (1931) described several new spe- genera and nearly 1900 species known so far to oc- cies from the Crimean Peninsula: Gammarus tauri- cur in such environment (Balian et al. 2008). Most cus, G. nudus, G. spelaeus and G. kesslerianus. Kara- of this diversity concentrates in the Palearctic man & Pinkster (1977) listed several localities for realm, predominantly in middle latitudes, with G. kischinefensis in the Dniester River system. such hotspots in Europe as Iberian, Apennine and Dedju (1980) reported presence of Gammarus pulex Balkan peninsulas and also Ponto-Caspian region in Crimean Peninsula, G. lacustris in the entire (Vainola et al. 2008, Hou et al. 2011). Ukraine lies Dnieper River system, G. balcanicus in the Carpa- almost entirely within the Black Sea drainage ba- thian Mountains, Dniester system and tributaries sin, being a very important part of that region. De- of the upper Southern Bug River, and G. kischinef- spite its northern location, the area was free of ice fensis from the Dniester system. Jazdzewski & Ko- during the Pleistocene – the Dnieper River system nopacka (1988, 1989) discussed the distribution of originated around late Miocene, ca. 13.6 million G. balcanicus, G. kischinefensis and newly erected G. years ago (Matoshko et al. 2002), potentially offer- leopoliensis in the upper Dniester basin. ing a good opportunity for long term radiation of Gammarus varsoviensis Jazdzewski, 1975 is a freshwater fauna. species described originally from the vicinity of Surprisingly, amphipod freshwater fauna of Warsaw, in Central Poland. Until now the species Ukrainian inland waters has been rather poorly has been recorded from Central European low- studied. Most research activity has focused upon lands (Fig. 1), ranging from the right tributaries of large rivers such as Dniester or Dnieper and their the Elbe River (North Sea basin) in Germany on estuaries (limans). Particularly, stress was put on the west, through Oder and Vistula river systems the Ponto-Caspian oligohaline species (mainly of in Poland, Nemunas River in Lithuania to Dau- genera Pontogammarus, Obesogammarus, Dikero- gava River (Baltic basin) in Latvia on the east; only gammarus and Chaetogammarus) diversified in the few localities were recorded so far from the upper Black and Caspian Sea basins during subsequent Pripyat River (Black Sea basin) in Belarus (Jaz- transgressions and regressions and introduced dzewski 1975, Jazdzewski 1980, Jazdzewski & Ko- during Soviet times into the contemporarily built nopacka 1995, Arbaciauskas 2008, own unpub- artificial dam reservoirs (Zhuravel 1965, Pligin & lished data). Jazdzewski (1980) as well as Vainio et Yemelyanova 1989, Grigorovich et al. 2002). Now, al. (1995) noticed that G. varsoviensis distribution these studies are rather outdated. Concerning na- range does not conform to any general zo- First records of Gammarus varsoviensis in Ukraine 199 ogeographical pattern among native European 2011 did not reveal any further records of the species in other studied rivers. inland water animals (e.g. Illies, 1978). The above authors hypothesized that it might be a recent All new records of G. varsoviensis come from small immigrant from sources outside central Europe, and medium size lowland rivers, flowing through expanding its range through the canal network forest and grassy areas, with slow current, sand- connecting various river systems. Unfortunately, muddy bottom and dense vegetation (Fig. 2). Such due to lack of data outside the area, no further conditions are identical to those in which it was conclusions could be derived. found in Poland, Germany, Lithuania and Belarus

In August 2009, during collecting trip to Ukraine, we un- (Jazdzewski 1975, Arbaciauskas 2008, own unpub- expectedly found several populations of Gammarus show- lished data). ing diagnostic features of G. varsoviensis and inhabiting A question arises, why the species was not re- rivers of the middle and the lower Dnieper River drain- corded earlier from this vast area, extending the age system, within the Black Sea basin (Table 1, Fig. 1). Qualitative samples of amphipods were collected with known geographic range of this species ca. 830 benthic hand-net from all available habitats at each site. kilometers south-east? In our opinion it was sim- The materials were fixed in 96% ethanol, identified to ply overlooked, first of all for its close resemblance species level based on available literature (Jazdzewski to Gammarus lacustris, from which it was not iden- 1975, Karaman & Pinkster 1977) and stored in the collec- tion of the Department of Invertebrate Zoology & Hydro- tifiable for a long time (Jazdzewski 1975). The two biology, University of Lodz. Subsequent collecting trip in species may be separated mainly through rather

Table 1. Findings of Gammarus varsoviensis in Ukrainian rivers (* population that co-exists with G. lacustris).

Station Decimal Decimal Altitude Locality Date number Latitude longitude (m asl) 1 51.73574 25.49367 146 Stochid River near Lubeshiv town 09.08.2009 2* 51.84818 25.47230 147 Pripyat R. near Lyubyaz village 09.08.2009 3 51.77030 25.35393 148 Korostianka R. near Derevok vill. 09.08.2009 4 51.22880 26.36290 152 Horyn R. near Krychyls’k village 10.08.2009 5 50.84359 29.81191 133 R. near Blidcha vill. 11.08.2009 6 50.88941 29.98299 125 Tal R. near Rudnya-Shpylivs'ka vill. 11.08.2009 7 50.68513 31.11143 91 Krezen R. near Zavorici vill. 26.08.2009 8 47.69575 35.57920 25 R. near Dymitrovka vill. 14.08.2009

Figure 1. Distribution of Gammarus varsoviensis in Europe. Grey circles – previously known records of the species (combined literature and own data). White circles with numbers – localities of G. varsoviensis in Ukraine revealed during 2009 survey (numbers on data points correspond to the station numbers in Table 1). White triangles – other localities in Ukraine studied during 2009 and 2011 surveys, where no G. varsoviensis was found. 200 Grabowski, M. et al.

Figure 2. Habitats of Gammarus varsoviensis in Ukraine: A – Pripyat River near Lyubyaz village, B – Horyn River near Krychyls’k village, C – Tal River near Rudnya-Shpylivs'ka village, D – Konka River near Dymitrovka village (photo: Michal Grabowski).

Figure 3. Morphology of antenna II and telson in Gammarus varsoviensis (A, C) and G. lacustris (B, D). Arrow indicates shape and position of antennal gland cone (after Karaman & Pinkster 1977, modified).

subtle, yet clear morphological differences in through which the river passes. Generally, G. shape of the gland cone on peduncle of second an- lacustris is a species with a vast geographical range tenna, curved upwards in G. varsoviensis and encompassing Eurasia and boreal part of North straight in G. lacustris, as well as armature of uro- America (Karaman & Pinkster 1977). In northern some and telson – with long setae in G. varsovien- parts of its range (e.g. Central Europe, Siberia), it sis, and short ones in G. lacustris (Fig. 3). That, occurs predominantly in lakes and lowland rivers, combined with an apparent lack of access to con- while more south it thrives only in alpine lakes. temporary literature, was apparently the reason Thus we can conclude that in reality, G. lacustris for which Dedju (1980) reported only G. lacustris probably occurs in Ukraine only in the northern from the entire Dnieper drainage area. In fact, dur- part of the country, while all other old records can ing the recent sampling we have found this spe- be attributed to G. varsoviensis. Such distribution cies only in the upper Pripyat River, where it co- implies a possibility that geologically old Dnieper occurred with G. varsoviensis (Table 1, Fig. 1) in system may be in fact a home range for the species almost stagnant water just below Lyubyaz Lake, from which it subsequently colonized northern First records of Gammarus varsoviensis in Ukraine 201 and western parts of its present range, covered Grigorovich, I.A., MacIsaac, H.J., Shadrin, N.V., Mills, E.L. (2002): Patterns and mechanisms of aquatic invertebrate introductions with ice sheet during the Pleistocene. In fact, all in the Ponto-Caspian region. Canadian Journal of Fishery and rivers of the Black Sea drainage area in which the Aquatic Sciences 59: 1189-1208. species was found have been connected to the Hou, Z., Sket, B., Fišer, C., Lia, S. (2011): Eocene habitat shift from saline to freshwater promoted Tethyan amphipod Dnieper system through artificial navigable canals diversification. Proceedings of the National Academy of since the XVIII/XIX century (Bij de Vaate et al. Sciences of the United States 108(35): 14533-14538. 2002). If such, then the species would be yet an- Illies, J. (1978): Limnofauna Europaea. 2. Auflage, Stuttgart, G. Fischer Verlag. other alien aquatic species recently colonizing Jazdzewski, K. (1975): Remarks on Gammarus lacustris G. O. Sars, Central Europe from the Black Sea basin (Borza et 1863, with description of Gammarus varsoviensis n. sp. al. 2011, Semenchenko et al. 2011). (Crustacea, Amphipoda). Bijdragen tot de Dierkunde 45: 71-86. Jazdzewski, K. (1980): Range extensions of some gammaridean species in European inland waters caused by human activity.

Crustaceana, Supplement 6: 84-107. Jazdzewski, K., Konopacka, A. (1988): Notes on the gammaridean Acknowledgements: Thanks are due to Natasha Amphipoda of the Dniester River basin and Eastern Rubtsova for her help in collecting samples during the Carpathians. Crustaceana, Supp1ement 13: 72-89. 2009 survey, performed and founded within the Polish Jazdzewski, K., Konopacka, A. (1989): Gammarus leopoliensis nov. Ministry for Science & Higher Education, grant no. N sp. (Crustacea, Amphipoda) from Eastern Carpathians. Bulletin of the Zoological Museum of the University of Amsterdam 11: N304 081 535. The fieldwork in 2011 was partially funded 185-196. by Polish Ministry for Science & Higher Education, grant Jazdzewski, K., Konopacka, A. (1995): Pancerzowce (Malacostraca) no. N N304 350 139 and partially by the private funds of prócz równonogów lądowych. Katalog Fauny Polski, cz. XIII, t. Radomir Jaskula within the 6th TBQuest expedition that 1, Warszawa: 165 pp. he organized and led. Thanks are due to him and Marek Karaman, G., Pinkster, S. (1977): Freshwater Gammarus species from Michalski for help in gathering samples during that trip. Europe, North Africa and adjacent regions of Asia (Crustacea- Amphipoda). Part I Gammarus pulex-group and related species. Wanda and Manolis Plaitis are wholeheartedly thanked Bijdragen tot de Dierkunde 47 (1): 1-97. and credited for hospitality in their house in Crete, Martynov, A.V. (1931): Zur Kenntnis der Amphipoden der Krim. Greece, allowing us to analyse the results and prepare the Zoologische Jahrbücher, Abteilung für Systematik, Ökologie manuscript on their terrace in summer 2011. Finally, Und Geographie der Tiere 60: (5-6): 573-606. thanks are due to Dr. Manfred Pöckl and Dr. Kresimir Matoshko, A.V., Gozhik, P.F., Ivchenko, A.S. (2002): The fluvial Zganec for reviewing the manuscript. archive of the Middle and Lower Dnieper (a review). Netherlands Journal of Geosciences / Geologie en Mijnbouw

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