Survey of River and Sea Lampreys in German Waters of the Baltic Sea – Basis of Successful Rebuilding Programmes. ICES CM 2005

Home , Binz, Hiddensee, Lauterbach (Rügen), Peenestrom, Ralswiek, Recknitz

Not to be cited without prior reference to the authors

ICES Annual Science Conference 2005 CM 2005/W:06 Rebuilding Programmes for Threatened Fish Populations

Survey of river and sea lampreys in German waters of the Baltic Sea – basis of successful rebuilding programmes

R. Thiel, H. M. Winkler, P. Riel and R. Neumann

ABSTRACT In order to prepeare population rebuilding programmes, the status of river lamprey and sea lamprey was investigated within the German waters of the Baltic Sea. From August 2003 until August 2005 the following sources of data were analysed: (i) research fisheries with otter and shrimp trawls, (ii) records from commercial and recreational fisheries, (iii) records from ichthyological museum collections, (iv) commercial catch statistics and relevant publications. 317 records of river lampreys with 19,977,622 individuals and 89 records of sea lampreys with 129 individuals were obtained for the period from 1649 - 2005. The number of recorded individuals of river lampreys per year was two times higher from 1649 - 1939 than in the following period 1940 - 1989. Only 1 % of the yearly records of river lampreys from this time period was estimated from 1990 - 2005. No distinct trend of the temporal population development of sea lamprey was found. From 1829 - 1989 the highest number of records of sea lamprey (70 %) was estimated for subdivisions 22 and 24. From 1990 - 2005 all records of sea lamprey in German Baltic waters were distributed within the coastal waters and estuaries. Areas with distinct higher densities of sea lamprey records were not estimated. Most of the records of river lamprey (88 %) originated from subdivisions 24 and 26 from 1649-1989. The species was mainly distributed within the Szczecin Lagoon and adjacent waters, the lower Vistula River, the Vistula Lagoon, the Bay of Gdańsk and the Curonian Lagoon. From 1990 - 2005 most of the records (83 %) and recorded individuals (98 %) of river lamprey from German Baltic waters originated from the Szczecin Lagoon and its adjacent waters. 9 Spawning sites of river lamprey were found in the river systems of Peene, Warnow and Stepenitz. Regular annual spawning of river lamprey occurs not at all spawning sites. Actual reproduction of sea lamprey is not known from rivers of the German Baltic Sea area. From 1887-1999 about 14,377 kilogram of river lampreys were caught annually in the southern Baltic Sea. 82 % of the total lamprey catches of the southern Baltic Sea originated from subdivision 26. Highest lamprey catches were obtained during the decades from 1890 - 1919. The contribution of the lower Vistula River, Vistula Lagoon and Bay of Gdańsk increased remarkable from 1887-1939, whereas the importance of the Szczecin Lagoon and adjacent waters for the lamprey fishery decreased during the same period of time. A remarkable proportion of lamprey catches is contributed from the region of the Curonian Lagoon since 1950. However, the critical situation for the populations of river lamprey in both marine and freshwater habitats of the German Baltic Sea area requires a rebuilding programme for the still existing but threatened populations.

Keywords: river lamprey, sea lamprey, Baltic Sea, past distribution, commercial catch, recent records, rebuilding programme

Ralf Thiel, Philip Riel, Renate Neumann: German Oceanographic Museum, Katharinenberg 14-20, D- 18439 Stralsund, Germany [tel: +49 3831 2650 300, fax: +49 3831 2650 209, e-mail: [email protected]]. Helmut M. Winkler: University of Rostock, Department Biological Sciences, Universitätsplatz 2, D-18055 Rostock, Germany [tel: +49 381 498 6272, fax: +49 381 498 6262, e-mail: [email protected]]

1 INTRODUCTION

Lampreys belong to the Agnatha – the most primitive and oldest group of all living vertebrates. Their fossils are available from the late Silurian and Devonian periods, 450 million years ago (Maitland 2003). Most recent lamprey species have an antitropical distribution and occur in the temperate and arctic regions of both the northern and southern hemispheres (Kelly & King 2001). Exceptionally, the genus Tetrapleurodon occurs in subtropical Mexico, but lives here at high altitude (Salewski 2003). In Eurasia and North America lampreys are represented by the genus Lampetra, Petromyzon, Ichthyomyzon, Lethenteron, Eudontomyzon, Entosphenus and Caspiomyzon. There are 39 recognised lamprey species worldwide (Potter 1980), of which there are 34 present in the northern hemisphere (Krappe 2004). Four species represent the Petromyzontiformes in Germany (Freyhof 2002). These are sea lamprey (Petromyzon marinus Linnaeus, 1758), river lamprey (Lampetra fluviatilis (Linnaeus, 1758)), brook lamprey (Lampetra planeri (Bloch, 1784)) and Vladykov´s lamprey (Eudontomyzon vladykovi Oliva & Zanandrea, 1959). River lamprey and sea lamprey are the only two anadromous lamprey species living in German waters of the Baltic Sea, its estuaries and rivers (Spratte & Hartmann 1998, Winkler et al. 2002). However, the river lamprey is found from southern Norway to the western Mediterranean. The species inhabits the coastal waters of the whole Baltic Sea. It has also been reported from Turkey (Erguven 1989). Anadromous sea lampreys occur over much of the Atlantic coastal area of western and northern Europe, from northern Norway to the western Mediterranean, and eastern North America. In Europe it is distributed as far north as Varanger Fjord in Norway (70° N) and as far south as the western Mediterranean (Beamish 1980, Hubbs & Potter 1971). It has recently been documented from the Aegean Sea by Economidis et al. (1999) and from the Kola Bay (Reshetnikov 2003). According to Kelly & King (2001) there exist also several landlocked populations of sea lamprey, but not in the Baltic Sea region, where the species occurs in very low densities. It is known that populations of anadromous lampreys have declined dramatically in European inland waters, including Germany, since the mid of the 20th century (e.g. Kelly & King 2001). Due to their decrease in stocks and numbers in inland waters, both lamprey species are listed in Annex II of the European Union Habitats Directive as species of community interest, whose conservation requires the designation of Special Areas of Conservation; in Appendix III of the Bern Convention, which permits some exploitation of their populations; and in many Red Data Books throughout Europe, which list these species often as threatened. However, there is a great lack of information with regard to the status of anadromous lamprey species in marine waters, e.g. in the Baltic Sea. In order to check, if rebuilding programmes for anadromous lampreys are necessary and how they should be performed in that case, we analysed the status of diadromous lamprey species within the German waters of the Baltic Sea area. In addition, this paper provides useful information for biodiversity considerations with regard to river and sea lamprey and

2 outlines requirements for their conservation in Germany. Furthermore, to provide such information is particularly relevant to fulfill the requirements of the European Habitats Directive, the EU Water Framework Directive, and of coastal zone management and conservation. In detail, this study was performed to describe, analyse and discuss: (1) the past distribution of lampreys in subdivisions 20-26 of the Baltic Sea, (2) the spatial and temporal characteristics of the commercial catch of lampreys in the southern Baltic Sea, (3) the recent records of lampreys in German Baltic waters, and (4) the location of spawning sites of lampreys in German tributaries of the Baltic Sea region.

MATERIAL AND METHODS

During a 25 months period from August 2003 until August 2005 the status of anadromous lampreys in German waters of the Baltic Sea area was investigated based on the analysis of the following sources of data: (i) research fisheries with otter and shrimp trawls, (ii) records from commercial and recreational fisheries, (iii) records from ichthyological museum collections, (iv) commercial catch statistics and relevant publications.

Evaluation of hauls from research fishery The research fisheries were carried out with different bottom trawls (otter and shrimp trawls) in the Exclusive Economic Zone and in the coastal waters of Germany in the Baltic Sea region between the Islands of Ruegen, Usedom and Bornholm during autumn 2003, in spring, summer and autumn 2004, and in spring and summer 2005. Fish sampling was performed at 22 stations. Overall, 165 hauls were carried out. Trawls were towed during daylight and lasted for 30-60 min. After collection, adult and larger juvenile individuals were identified to species, counted, weighted, measured and released back into the water immediately. Fish larvae and small juveniles were investigated in the laboratory. More details of the construction of the trawls, the sampling procedures and catch efficiencies of the gears are given in Thiel et al. (2004, 2005). Additionally, the Federal Research Centre for Fisheries contributed catch records of lampreys from the German Baltic waters since 1991 based on data resulting from more than 3,000 hauls, taken with otter trawls. Some additional data originating from studies in the Pomeranian Bay and from hydro-acoustic surveys were available from 1978. More details regarding the sampling regime used by the Federal Research Centre for Fisheries are presented in the paper by Kloppmann et al. (2003).

Compilation of recent catch records from commercial and recreational fishery In order to ensure that commercial and recreational fishermen communicate actual catch records of lampreys, an information sheet containing drawings with the most important determination characteristics of the species was developed. This information sheet was

3 distributed along the Baltic coast of Germany. Furthermore, a catch award was announced to those fishermen communicating verified and accurate catches of lampreys (Fig. 1).

Analysis of ichthyological collections The ichthyological collections of museums and universities which are known to contain individuals of lampreys from the Baltic Sea in their collections were analysed. In unclear cases of scientific determination, the specimens were re-determined. Ichthyological collections and/or databanks of the following 10 institutions were analysed for records of lampreys from subdivisions 20 - 26 of the Baltic Sea:

• Museum of Natural History of the Humboldt University in Berlin, Germany (NHMB), • Zoological Institute and Museum of the University of Hamburg, Germany (ZMUH), • Museum of Natural History Stuttgart, Germany (NHMS), • Zoological Museum of the Christian-Albrechts-University Kiel, Germany (ZMUK), • Department Biology of the University of Rostock, Germany (DBUR), • German Oceanographic Museum, Stralsund, Germany (GOMS; Fig. 2), • Zoological Institute and Museum of the Ernst-Moritz-Arndt-University Greifswald, Germany (ZMUG), • Zoological Museum of the University of Copenhagen, Denmark (ZMUC), • Swedish Museum of Natural History, Stockholm, Sweden (NRMS), • Smithsonian National Museum of Natural History, Washington, USA (USNM),

Synopsis of data from commercial catch statistics and publications Data from papers containing original information about catch statistics of lampreys in the southwest Baltic Sea and of information about the actual location of spawning sites in German tributaries of the Baltic Sea were assessed, selected and than compiled. Such data were used from e.g. Anonymus (1887-1920), Eichelbaum (1926-1940), Duncker (1960), Leipe & Königstedt (1988), Bartel (1993), Schröder (1995), Witkowski (1996), Raczyński & Filipiak (1999), Gaigalas (2001), Winkler et al. (1999), Waterstraat & Krappe (2000), Winkler et al. (2000), Waterstraat et al. (2001), Winkler et al. (2002), Krappe (2004), Schaarschmidt & Lemcke (2004), Spratte (2004), Thiel et al. (2004, 2005). The commercial catch data were compiled in units of kilogram, metric tons and number of individuals. To allow a clear presentation in figures 3 - 6 and 10 - 11, records of lampreys from the same localities and time periods were summarized.

4 RESULTS

Origin of available records of lampreys 317 records of river lampreys with 19,977,622 individuals and 89 records of sea lampreys with 129 individuals were obtained based on the analysis of all available data sources from 1649 - 2005 (Tables 1, 2, 4, 5). Records originated only from commercial catch statistics, publications and ichthyological collections from 1649 - 1989. During that period of time, 83.27 % of all records of river lamprey and 32.89 % of the records of sea lamprey were obtained from commercial catch statistics and publications, whereas 16.73 % of the river lamprey records and 67.11 % of the records of sea lamprey originated from the analysis of ichthyological collections. From 1990 - 2005, most records of river lampreys (78.79 %) were obtained from commercial and recreational fishery. 18.18 % of the records originated from commercial catch statistics, publications and ichthyological collections, whereas research fishery provided 3.03 % of the records of river lampreys during that time period. In contrast, most records of sea lampreys (53.85 %) resulted from the analysis of commercial catch statistics, publications and ichthyological collections between 1990 and 2005. 38.46 % of the records originated from commercial and recreational fishery, whereas research fishery provided 7.69 % of the records of sea lamprey from 1990 - 2005.

Past distribution of lampreys in subdivisions 20-26 of the Baltic Sea The analysis of relevant records in ichthyological collections and of data from commercial catch statistics and publications resulted in 36 different records of sea lampreys and 185 records of river lampreys in the southwest Baltic Sea for the time period 1649 -1939 (Table 1 and 2, Fig. 3 and 5). From 1940 - 1989, sea lamprey was recorded 40 times (Table 1, Fig. 4), whereas 66 records of river lamprey were obtained (Table 2, Fig. 5). However, the number of recorded individuals of river lampreys until 1939 was with 62,305 individuals per year (18,130,740 individuals in total) approximately two times higher than in the following time period 1940 - 1989, when 36,802 individuals per year (1,840,087 individuals in total) were recorded (Table 2). From 1940 - 1989, in average 1.5 individuals per year (75 individuals in total) of sea lampreys were recorded, whereas with 0.4 individuals per year (42 individuals in total) a lower number was obtained between 1829 and 1939 (Table 1). However, these changes were observed on a very low density level of sea lamprey. Therefore, the values are not appropriate to describe a distinct trend of the temporal population development of this species. The oldest record of river lamprey dates back to the year 1649, when this species was recorded in the Szczecin Lagoon near the city of Ueckermuende (Table 2). The oldest record of a sea lamprey originated from Skagen in Denmark from 1829 (Table 1). From 1829 - 1989, the highest number of records of sea lamprey (70.15 %) was estimated for subdivisions 22 and 24 (Table 1, Fig. 3 and 4). However, offshore records of sea lampreys were rare.

5 No record of sea lamprey was available in the eastern subdivisions 25 and 26 until 1939 (Fig. 3). The species was occasionally found in these subdivisions from 1940 - 1989 (Fig. 4). Compared to the period 1829 - 1939, a higher density of records of sea lampreys was obtained from the Baltic coast of Germany between 1940 and 1989 (Fig. 3 and 4). Most of the records of river lamprey (87.65 %) originated from subdivisions 24 and 26 from 1649 - 1989 (Table 2, Fig. 5 and 6). During that time period the species was mainly distributed within the three following areas of the southern Baltic Sea (Table 2, Fig. 5 and 6):

• Szczecin Lagoon and adjacent waters (subdivision 24), • Lower Vistula River, Vistula Lagoon and Bay of Gdańsk (subdivision 26), • Curonian Lagoon (subdivision 26).

Remarkable numbers of records of river lampreys were also obtained from the Kiel Bight, Mecklenburg Bight, mouth of the Warnow River and from the waters around the Ruegen Island from 1649 - 1939 (Table 2, Fig. 5). From 1940 - 1989, river lamprey was not recorded from the Kiel Bight and the number of records in the southwest Mecklenburg Bight was much lower than in the previous time period (Table 2, Fig. 5 and 6).

Spatial and temporal characteristics of the commercial catch of lampreys in the southern Baltic Sea Sea lamprey has not been used commercially in the southern Baltic Sea. There is evidence that river lamprey has been a commercial species in the Baltic since the end of the 19th century. However, this species has probably been exploited earlier. From 1887 - 1999 about 14,377 kilogram (kg) of river lampreys were caught annually in the southern Baltic Sea. However, remarkable higher catches were obtained during the decades from 1890 - 1919, when 44,430, 38,250 and 32,794 kg of lampreys were caught annually (Table 3). Later on, annual lamprey catches decreased significantly (Fig. 7). Mean annual catch of river lamprey ranged between 400 and 14,814 kg within the decades between 1950 and 1999 (Table 3). Within that period, the highest mean annual lamprey yield (14,814 kg) was obtained during the decade from 1970 - 1979. 81.61 % of the total lamprey catch of the southwest Baltic Sea was achieved in from subdivision 26 from 1887 - 1999 (Table 3). About 65.26 % of the mean annual yield of lampreys were contributed by the lower Vistula River, Vistula Lagoon and Bay of Gdańsk, followed by the Szczecin Lagoon and adjacent waters (18.39 %) and the Curonian Lagoon and Nemunas Delta (16.35 %, Table 3). The contribution of the lower Vistula River, Vistula Lagoon and Bay of Gdańsk increased remarkable from 1887 - 1939, whereas the importance of the Szczecin Lagoon and adjacent waters for the lamprey fishery decreased during the same period of time (Fig. 8). Lamprey catches originated nearly exclusively from the lower Vistula River, Vistula Lagoon and Bay of Gdańsk from 1920 - 1939 (Fig. 8). A remarkable proportion of lamprey catches is contributed from the Curonian Lagoon and Nemunas Delta since 1950 (Table 3, Fig. 8). Seasonal catches of river lamprey showed maximum values from August to December and

6 peaked in October and November for each of the three areas compared (Fig. 9). A small spring component (March) of seasonal catches was only estimated for the Vistula Lagoon.

Recent records of lampreys in German Baltic waters and location of their spawning sites in German tributaries of the Baltic Sea region 13 records of sea lamprey with 12 individuals (0.75 individuals per year) with total lengths between 43 and 85 cm were obtained in German Baltic waters from 1990 - 2005 (Table 4). All records were distributed within the coastal German waters and estuaries (Fig. 10). Areas with distinct higher densities of sea lamprey records were not found A total of 66 records of river lampreys with 6,795 individuals (425 individuals per year) with total lengths between 30 and 42 cm originated from German Baltic waters from 1990 - 2005 (Table 5). River lampreys were recorded from the Salzhaff, the Warnow River, the Recknitz River, from around the Ruegen Island, the Pommeranian Bay, from the Szczecin Lagoon with adjacent waters and from 9 spawning sites in German tributaries of the Baltic Sea (Fig. 11). Most of the records (83.33 %) and recorded individuals (98.32 %) of river lamprey originated from the Szczecin Lagoon and its adjacent waters (Table 5), demonstrating the importance of these water bodies for the spawning migrations of this species. Spawning sites for river lamprey were found in the river systems of Peene, Warnow and Stepenitz (Table 6, Fig. 11). The River Peene system must be regarded as the most important spawning site of river lampreys in the German Baltic Sea region. Here, 5 spawning sites were estimated, namely in the tributaries Scheidegraben, Augraben, Galgenbach, Klenzer Muehlbach and Libnower Muehlbach (Table 6). The River Warnow system provides three spawning places in the tributaries Beke and Koesterbeck and in the Muehlendamm canal near the city of Rostock (Fig.11). Only one spawning site was found in the Radegast, a tributary of the River Stepenitz (Table 6). Spawning sites of sea lamprey were not detected in German tributaries to the Baltic Sea.

DISCUSSION

The results of the present paper clearly indicate that subdivisions 22 and 24 are the main past and present distribution areas of sea lamprey in the Baltic Sea. Most records of sea lamprey originate from coastal waters, although a few individuals were caught offshore, especially in the Kattegat and Skagerrak. Lelek (1973) pointed out that sea lampreys can be found in the open sea up to several hundred kilometres away from the coast. According to Kelly & King (2001) sea lampreys occur in highest densities at latitudes between 35° and 45°. However, the Baltic Sea is located at higher latitudes than the main distribution area of sea lamprey. Although the sea lamprey was reported with low densities during the present study and is regarded as a rarity in the Baltic Sea region (Winkler et al. 2000), this species has been occasionally reported from Denmark, Sweden, Finland, Russia, Latvia, Lithuania, Estonia, Poland and Germany (Froese & Pauly 2005, Kull 1999, Winkler et al. 2000).

7 It is still a question, from which spawning population those sea lampreys being recorded in German waters of the Baltic Sea during this study, originate. Historically, the sea lamprey may have spawned in the River Trave system (Duncker 1960). However, there exists no observation from the last years showing that this spawning site is used by sea lampreys (Spratte 2004). Nowadays, no actual reproduction of sea lamprey in the whole German Baltic Sea area is known (Spratte & Hartmann 1998, Winkler et al. 2002). Furthermore, the results of the present study have shown that there is no clear trend of an increase or decrease of sea lamprey densities in German waters of the Baltic Sea during the last centuries. Our results demonstrate that in contrast to the rare sea lamprey, the river lamprey is still a common and widely distributed member of the ichthyofauna of the Baltic Sea, its estuaries and tributaries. In the Baltic Sea region, populations of river lamprey occur in Denmark, Sweden, Finland, Russia, Latvia, Lithuania, Estonia, Poland and Germany (Anonymus 1998, Bartel et al. 1993, Winkler et al. 2000). Although the species is mainly anadromous in the Baltic Sea area, there are a few landlocked, non-migratory populations isolated from the sea in Finland (Tuunainen et al. 1980, Valtonen 1980) and eventually in Russia (e.g. Abakumov 1956, Reshetnikov 2003). According to Hardisty (1986) river lampreys shall live predominantly in coastal areas, especially near estuaries. However, our study summarized a few records of river lampreys outside coastal waters of the Baltic Sea. The present study has shown that the German, Polish and Lithuanian waters, especially within subdivisions 24 and 26, were the main distribution and as well fishery areas of river lamprey in the past. About hundred years ago, the river lamprey was an important commercial species for coastal fisheries in the southern Baltic waters. In Germany, no lamprey fishery occurs anymore, whereas, according to our results, the river lamprey was used commercially in Poland and Lithuania at lower levels at least until 1999. In rivers of Finland, Russia, Latvia and Estonia the commercial fishery of lamprey is still active (Anonymus 1998). The estimated seasonal pattern of catch indicates that lampreys in the southern Baltic were mainly caught during their spawning migrations. This pattern has also been observed by Sterner (1918) and Imam et al. (1958) within the framework of investigations of lamprey catches in rivers flowing into the North Sea. During our study only 9 used spawning sites of river lampreys were found in German tributaries of the Baltic Sea. 8 of them were located in only two river systems (Peene and Warnow). Furthermore, Winkler et al. (1999) pointed out that nowadays regular annual spawning of river lamprey occurs not at all spawning sites. Additionally, the spawning populations of river lamprey are very small, comprising only 20 to 100 individuals (Winkler et al. 2002). According to Imam et al. (1958), Wilkens & Köhler (1977), Möller (1984), Thiel & Salewski (2003), Krappe (2004) and Spratte (2004) a considerable decrease of anadromous lamprey populations during the mid 1950s has been observed in inland waters of Germany. Lelek (1987) assumed that pollution of estuaries and the building of weirs and dams prevented lampreys from reaching their original spawning grounds. Similarly, Kelly & King (2001) concluded that populations of anadromous lampreys have declined in European inland

8 waters mainly due to pollution, to the construction of dams associated with hydroelectric power stations, weirs and other manmade barriers in rivers and channelisation and land- management practices that lead to increased siltation on spawning gravels. Recently Spratte (2004) concluded that the increased number of records of river lampreys in inland waters of Schleswig-Holstein in northwest Germany during the last years could be attributed to improved water quality in the rivers. In almost all rivers and streams in the catchment of the River Rhine, water quality has improved considerably, and the problems of low oxygen concentrations became unimportant since the early 1980s (Freyhof 2002). Therefore, populations of river and sea lamprey recovered there (Freyhof & Weibel 1999, Korte 1999). However, the situation of the river lamprey populations in Mecklenburg- Vorpommern in northeast Germany is still critical (Winkler et al. 2002). The present study summarizes records of river lampreys from their parasitic feeding phase in the marine environment and also from their spawning sites in freshwater. The results confirm the critical situation for the populations of river lamprey in both marine and freshwater habitats of the German Baltic Sea area. A rebuilding programme for the still existing but threatened populations of river lampreys is necessary. In this regard it is noteworthy that the European Union Habitats Directive requires the establishment of a European network of important, high quality conservation sites that will contribute significantly to the conservation of distinct habitat types and species. The river lamprey and sea lamprey are listed in Annex II of the Directive, which lists animal and plant species of interest to the European Community whose conservation requires the designation of special areas of conservation (SACs). The river lamprey is also listed in Annex V of the Directive, which lists those animal and plant species of European Community interest whose exploitation and taking in the wild may be subject to management measures. However, it is unlikely that a commercial fishery for river lamprey will ever start again in Germany, but if it were ever proposed it must be managed under strict guidelines to ensure sustainability. Fricke (2000) proposed marine and estuarine NATURA 2000 protected areas for Germany. This can be considered as a first step for conservation of river lampreys in those waters. In 2004 Germany has nominated a number of SACs in German Baltic waters to the EU Commission. They also cover parts of the Szczecin Lagoon with adjacent waters, where the main migration route of river lampreys to their most important spawning sites has been detected during this study. However, the rebuilding programme should also contain measures to restore lost spawning sites and the connectivity within the river systems themselves that the lampreys can reach their spawning sites during their spawning migrations. The habitat connectivity for the river lamprey can be increased with the removal of barriers and weirs or provision of fish passes in rivers, where their spawning and nursery habitats are situated. However, more detailed measures for rebuilding programmes for river lamprey in European countries are provided by, e.g., Kelly & King (2001) for rivers in Ireland and Maitland (2000) for UK rivers. Leipe & Königstedt (1988), Waterstraat & Krappe (2000), Petersen et al. (2004) and Spratte (2004) describe measures in the framework of rebuilding programmes for river lamprey in German rivers.

9 In conclusion from the results of the present paper, special rebuilding programmes for the sea lamprey are not recommended for the German waters of the Baltic Sea region. However, it should be observed, if the meanwhile improved connectivity within the River Trave system (Hartmann, pers. comm.) will lead to a successful spawning migration and spawning of sea lampreys. However, sea lamprey could also benefit from measures performed in the framework of rebuilding programmes for river lamprey.

Acknowledgements We thank Uwe Böttcher, Bernd Bruns, Tomas Gröhsler, Jens Heischkel, Volker Huckstorf, Dietmar Lill, Nardine Löser and Ronny Weigel for their helpful contributions and comments to this study. The study was assigned by the German Federal Agency for Nature Conservation and sponsored by the Federal Ministry for the Environment, Nature Conservation and Nuclear Safety under grant number 80385220.

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10 Freyhof, J. (2002). Chapter 1. Freshwater fish diversity in Germany, threats and species extinction. In: M.J. Collares-Pereira, I.G. Cowx & M.M. Coelho (eds.). Conservation of Freshwater Fishes: Options for the Future. Fishing News Books, Blackwell Science Ltd, Oxford, London: 3-22. Freyhof, J. & Weibel, U. (1999). Vorschlag einer Neufassung der Roten Liste der bestandsgefährdeten Neunaugen und Fische (Petromyzontiformes; Teleostei) in Rheinland-Pfalz sowie ein Verzeichnis aller vorkommenden Arten. Fauna und Flora in Rheinland-Pfalz 9: 195-206. Fricke, R. (2000). Auswahl und Management mariner NATURA 2000 Gebiete für Fischarten im Anhang II der FFH-Richtlinie. Schriftenreihe Landespflege und Naturschutz 68: 113- 133. Gaigalas, K. (2001). Kuršių marių baseino žuvys ir žvejyba. Eglė, Klaipeda: 368 pp. Hardisty, M.W. (1986). A general introduction to lampreys. In: Holcik, J. (ed.) The Freshwater Fishes of Europe. Band 1/1 Petromyzontiformes. Aula Verlag, Wiesbaden. Hubbs, C.L. & Potter, I.C. (1971). Distribution, phylogeny and taxonomy. In Hardisty, M. W. & Potter, I. C. (eds.) The Biology of Lampreys, Vol. 1. London. Academic Press. Imam, A. K. E. S., Lühmann, M. & Mann, H. (1958). Über Neunaugen und Neunaugenfischerei in der Elbe. Fischwirt 9: 249-261. Kelly, F. L. & King, J. J. (2001). A review of the ecology and distribution of three lamprey species, Lampetra fluviatilis (L.), Lampetra planeri (Bloch) and Petromyzon marinus (L.): a context for conservation and biodiversity considerations in Ireland. Biology and Environment: Proceedings of the Royal Irish Academy 101B (3): 165-185. Kloppmann, M.H.F, Böttcher, U., Damm, U., Ehrich, S., Mieske, B., Schultz, N. & Zumholz, K. (2003). Erfassung von FFH-Anhang II-Fischarten in der deutschen AWZ von Nord- und Ostsee. Final report for the German Federal Agency of Nature Conservation: 82 pp. Korte, E. (1999). Bestandsentwicklung der Fischarten der hessischen Rheinaue 1994-1997. Reproduktionsstrategien, Jungfischaufkommen, Gefährdung, Entwicklungstendenzen. Umweltplanung, Arbeits- und Umweltschutz 268: 1-186. Krappe, M. (2004). Quantitative Analysen populationsbiologischer Phänomene im Lebenszyklus des Bachneunauges Lampetra planeri (Bloch, 1784). PhD thesis, University of Rostock: 241 pp. Kull, T. (1999). Estonian Biodiversity Strategy and Action Plan. EPMÜ Keskkonnakaitse Instituut: 165 pp. Leipe, T. & Königstedt, D. (1988). Zum Vorkommen und Schutz der Neunaugen (Cyclostomata, Petromyzontiformes) in Mecklenburg. Naturschutzarbeit in Mecklenburg-Vorpommern 31: 12-21. Lelek, A. (1973). Occurrence of the Sea Lamprey in midwater of Europe. Copeia 1: 136-137. Lelek, A (1987). Threatened Fishes of Europe. In: Holcik, J. (ed.) The Freshwater Fishes of Europe. Volume 9, Aula Verlag, Wiesbaden: 343 pp.

11 Maitland, P. S. (2003). Ecology of the River, Brook and Sea Lamprey - Lampetra fluviatilis, Lampetra planeri and Petromyzon marinus. Conserving Natura 2000 Rivers. Ecology Series No. 5: 1-52. Möller, H. (1984): Daten zur Biologie der Elbfische. Verlag Heino Möller, Kiel. Petersen, B., Ellwanger, G., Bless, R., Boye, P., Schröder, E. & Ssymank, A. (2004). Das europäische Schutzgebietssystem Natura 2000. Ökologie und Verbreitung von Arten der FFH-Richtlinie in Deutschland. Schriftenreihe für Landschaftspflege und Naturschutz 69 (2): 693 pp. Potter, I. C. (1980). The Petromyzontiformes with particular reference to paired species. Canadian Journal of Fisheries and Aquatic Sciences 37: 1595-1615. Raczyński, M. & Filipiak, J. (1999). A preliminary biological and morphometric characteristics of river lamprey (Lampetra fluviatilis (L.)) from Lake Dabie. Electronic Journal of Polish Agricultural Universities, Fisheries 2 (2): 12 pp. Reshetnikov, Y. S. (2003). Atlas of Russian Freshwater Fishes. Vol.1. Nauka, Moscow: 378 pp. Salewski, V. (2003). Satellite species in lampreys: a worldwide trend for ecological speciation in sympatry? Journal of Fish Biology 63: 267-279. Schaarschmidt, T. & Lemcke, R. (2004). Quellendarstellungen zur historischen Verbreitung von Fischen und Rundmäulern in Binnengewässern des heutigen Mecklenburg- Vorpommerns. Mitteilungen der Landesforschungsanstalt für Landwirtschaft und Fischerei Mecklenburg-Vorpommern 32: 1-261. Schröder, H. (1995). Meerneunaugen in den Küstengewässern Mecklenburg-Vorpommerns. Meer und Museum 11: 31-40. Spratte, S. (2004). Projektierungsstudie zur Verbesserung der Bestandsentwicklung von Neunaugen in Schleswig-Holstein. Landessportfischerverband Schleswig-Holstein e.V.: 36 pp. Spratte, S., Hartmann, U. (1998): Süßwasserfische und Neunaugen in Schleswig-Holstein. Ministerium für ländliche Räume, Landwirtschaft, Ernährung und Tourismus des Landes Schleswig-Holstein, Kiel. Sterner, E. (1918). Die niederelbische Küstenfischerei, II. Die Fischereiarten. 10. Die übrigen Fischereiarten im oberen Teil. Fischerbote 10: 245-253. Thiel, R. & Salewski, V. (2003): Verteilung und Wanderung von Neunaugen im Elbeästuar (Deutschland). Limnologica 33:214-226 Thiel, R., Winkler, H.M., Neumann, R. (2004): Erfassung von FFH-Anhang II Fischarten in der deutschen AWZ von Nord- und Ostsee. Study for the German Federal Agency of Nature Conservation. German Oceanographic Museum, Stralsund. 1. Report 02/2004: 67 pp. Thiel, R., Winkler, H.M., Neumann, R. (2005): Erfassung von FFH-Anhang II Fischarten in der deutschen AWZ von Nord- und Ostsee. Study for the German Federal Agency of Nature Conservation. German Oceanographic Museum, Stralsund. 2. Report 03/2005: 106 pp.

12 Tuunainen, P., Ikonen, E. & Auvinen, H. (1980). Lampreys and lamprey fisheries in Finland. Canadian Journal of Fisheries and Aquatic Sciences 37: 1953-1959. Valtonen, T. (1980). European river lamprey (Lampetra fluviatilis). Fishing and lamprey populations in some rivers running into Bothnian Bay, Finland. Can. J. Fish. Aquat. Sci. 38: 1967-1973. Waterstraat, A. & Krappe, M. (2000). Beiträge zur Ökologie und Verbreitung von FFH- Fischarten und Rundmäulern in Mecklenburg-Vorpommern: 1. Das Flussneunauge (Lampetra fluviatilis L.) im Peenesystem. Natur und Naturschutz in Mecklenburg- Vorpommern 35: 64-79. Waterstraat, A., Krappe, M., Spieß, H.-J. & Winkler H. M. (2001). Monitoring von Ichthyozönosen kleiner Fließgewässer - Bestandteil des ichthyologischen Artenmonitorings in Mecklenburg-Vorpommern. Naturschutzarbeit in Mecklenburg- Vorpommern 44 (1): 9-17. Wilkens, H. & Köhler, A. (1977). Die Fischfauna der mittleren und unteren Elbe: die genutzten Arten. Abh. Naturwiss. Ver. Hamburg 20: 185-222. Winkler, H. M., Skóra, K., Repečka, R., Ploks, M., Neelov, A., Urho, L., Gushin, A. & Jespersen, H. (2000). Checklist and status of fish species in the Baltic Sea. ICES CM 2000/Mini 11: 15 pp. Winkler, H.M., Spieß, H.-J., Waterstraat, A., Krappe, M. & Lemcke, R. (1999). Monitoring von FFH-Arten von Rundmäulern und Fischen in Referenzgebieten. Naturschutzarbeit in Mecklenburg-Vorpommern 42 (1): 24-40. Winkler, H. M., Waterstraat, A. & Hamann, N. (2002). Rote Liste der Rundmäuler, Süßwasser- und Wanderfische Mecklenburg-Vorpommerns, kommentiert, Stand 2002. Umweltministerium Mecklenburg-Vorpommern: 52 pp. Witkowski, A. (1996). Distribution of the river lamprey, Lampetra fluviatilis (Linnaeus, 1758), in inland waters of Poland and reasons of the species decline. Bull. Lampetra 2: 77-82.

13 Table 1. Historical records of sea lamprey in subdivisions 20 - 26 of the Baltic Sea in the time period 1829 - 1989. For abbreviations see chapter "Material and Methods".

Sub- Number of No. Year Country Locality Data source division individuals 1 1829 20 Denmark Skagen 1 ZMUC 2 1855 24 Germany Greifswald Bodden unknown Schaarschmidt & Lemcke (2004) 3 1856 24 Germany Strelasund 1 ZMUG 4 1862 20 Sweden Stroemstad 1 NRMS 5 1868 22 Denmark Korsor 1 ZMUC 6 1868 24 Germany Hiddensee Island 1 NHMB 7 1872 23 Denmark Hellebaek 1 ZMUC 8 1873 20 Denmark Skagen 1 ZMUC 9 1873 22 Germany Kiel Bight 1 ZMUK 10 1873 22 Germany Kiel Bight 1 ZMUKl 11 1874 21 Denmark Kattegat 1 ZMUC 12 1875 20 Denmark Skagen 1 ZMUC 13 1883 22 Germany Kiel Bight 1 NHMS 14 1884 24 Denmark Maribo unknown ZMUC 15 1885 21 Denmark Isefjord 1 ZMUC 16 1888 20 Sweden Stroemstad 1 NRMS 17 1891 20 Sweden Stroemstad 1 NRMS 18 1892 20 Sweden Stroemstad 1 NRMS 19 1894 20 Sweden Stroemstad 1 NRMS 20 1898 22 Germany Neustadt Bight 1 ZMUH 21 1899 21 Sweden Bohuslaen 1 NRMS 22 1904 22 Denmark Langeland Belt 1 ZMUC 23 1908 20 Sweden Stroemstad 2 NRMS 24 1910 22 Denmark Langeland Belt 1 ZMUC 25 1928 20 Sweden Gullmarn Fjord 1 NRMS 26 1928 20 Sweden Gullmarn Fjord 1 NRMS 27 1929 22 Denmark Great Belt 1 ZMUC 28 1929 24 Germany Lower Warnow River unknown Schaarschmidt & Lemcke (2004) 29 1930 20 Sweden Gullmarn Fjord 1 NRMS 30 1932 24 Germany Warnemuende unknown Schaarschmidt & Lemcke (2004) 31 1934 24 Germany Saal Bodden unknown Schaarschmidt & Lemcke (2004) 32 1935 21 Sweden Roeroe Island 2 NRMS 33 1936 21 Sweden Roeroe Island 1 NRMS 34 1937 20 Sweden Gullmarn Fjord 1 NRMS 35 1938 21 Denmark Gilleleje 1 ZMUC 36 1939 22 Germany Trave River 10 Duncker (1960) 37 1945 21 Sweden Bohuslaen 1 NRMS 38 1948 23 Denmark Humlebaek 1 ZMUC 39 1951 21 Denmark Aalborg 1 ZMUC 40 1951 22 Denmark Juelsminde 1 ZMUC 41 1951 22 Denmark Sejerobugt 1 ZMUC 42 1952 24 Germany Darss and Zingst 1 Schröder (1995) 43 1952 24 Germany Stralsund 1 Schröder (1995) 44 1953 22 Denmark Great Belt 1 ZMUC 45 1953 24 Denmark Mon Island 1 ZMUC 46 1956 22 Germany Boltenhagen 1 GOMS 47 1957 21 Sweden Laerjean River unknown NRMS 48 1957 26 Lithuania Curonian Lagoon 1 Gaigalas (2001)

14 Sub- Number of No. Year Country Locality Data source division individuals 49 1960 20 Denmark Skagen 1 NRMS 50 1961 24 Germany Lower Warnow River 1 DBUR 51 1961 26 Lithuania Curonian Lagoon 24 Gaigalas (2001) 52 1964 25 Sweden Torhamn 2 NRMS 53 1967 21 Sweden Aetran River 13 NRMS 54 1972 24 Germany Ruegen Island near Lauterbach 1 GOMS 55 1977 24 Germany Hiddensee Island 1 GOMS 56 1980 24 Germany Arkona 1 Schröder (1995) 57 1980 24 Germany Warnemuende 1 GOMS 58 1981 24 Germany Baltic Sea near Wustrow 1 Schröder (1995) 59 1981 24 Germany Stubbenkammer 1 Schröder (1995) 60 1982 22 Germany Wohlenberger Wiek 1 Schröder (1995) 61 1982 24 Germany Usedom Island near Bansin 1 GOMS 62 1982 24 Germany Usedom Island near Bansin 1 GOMS 63 1983 24 Germany Greifswald Bodden 1 Schröder (1995) 64 1983 24 Germany Baltic Sea near Binz 1 Schröder (1995) 65 1983 24 Germany Prohner Wiek 1 Schröder (1995) 66 1985 24 Germany Markgrafenheide 1 GOMS 67 1985 24 Germany Lower Warnow River 1 Schröder (1995) 68 1987 24 Germany Hiddensee Island 1 Schröder (1995) 69 1987 24 Germany Mouth of River Peene 1 Schröder (1995) 70 1988 24 Germany Prohner Wiek 1 Schröder (1995) 71 1989 22 Germany Kuehlungsborn 1 Schröder (1995) 72 1989 22 Germany Salzhaff 1 Schröder (1995) 73 1989 24 Germany Goehren 1 Schröder (1995) 74 1989 24 Germany Greifswald Bodden 1 GOMS 75 1989 24 Germany Mukran 1 Schröder (1995) 76 1989 24 Germany East of Ruegen Island 1 Schröder (1995)

15 Table 2. Historical records of river lamprey in subdivisions 20 - 26 of the Baltic Sea in the time period 1649 - 1989. For abbreviations see chapter "Material and Methods".

Sub- Number of No. Year Country Locality Data source division individuals Szczecin Lagoon near 1 1649 24 Germany unknown Schaarschmidt & Lemcke (2004) Ueckermuende 2 1723 26 Poland Vistula Lagoon unknown Schaarschmidt & Lemcke (2004) 3 1794 24 Germany Lower Warnow River unknown Schaarschmidt & Lemcke (2004) 4 1831 26 Russland Kaliningrad 1 ZMUC 5 1837 24 Germany Greifswald Bodden 1 ZMUG 6 1837 24 Germany Peenestrom near Lassan 1 ZMUG 7 1842 22 Germany Holstein 1 ZMUC 8 1844 24 Germany Greifswald Bodden 1 ZMUG 9 1854 24 Germany Greifswald Bodden 2 ZMUG 10 1855 24 Germany Greifswald Bodden unknown Schaarschmidt & Lemcke (2004) 11 1855 24 Germany Upper Warnow River unknown Schaarschmidt & Lemcke (2004) 12 1858 24 Germany Greifswald Bodden 1 ZMUG 13 1867 24 Germany Greifswald Bodden 1 ZMUG 14 1868 23 Denmark Oeresund 1 ZMUC 15 1868 23 Denmark Oeresund 1 ZMUC 16 1868 24 Germany Lower Warnow River unknown Schaarschmidt & Lemcke (2004) 17 1868 24 Poland Odra River 3 NHMB 18 1875 22 Germany Kiel 1 ZMUK 19 1875 24 Germany Lower Warnow River unknown Schaarschmidt & Lemcke (2004) 20 1878 22 Germany Kiel 1 ZMUK 21 1880 24 Germany Lower Warnow River unknown Schaarschmidt & Lemcke (2004) 22 1881 22 Germany Kiel 1 ZMUK 23 1881 22 Germany Kiel 1 ZMUK 24 1881 22 Germany Kiel 1 ZMUK 25 1881 24 Germany Lower Warnow River unknown Schaarschmidt & Lemcke (2004) 26 1882 22 Germany Dassower See unknown Schaarschmidt & Lemcke (2004) 27 1882 24 Germany Spring near Sagard unknown Schaarschmidt & Lemcke (2004) 28 1882 24 Germany Stralsund unknown Schaarschmidt & Lemcke (2004) 29 1883 24 Germany Greifswald Bodden 1 ZMUG 30 1885 24 Germany Upper Warnow River 1 DBUR 31 1887 24 Germany Szczecin Lagoon 39771 Anonymous (1887-1920) 32 1887 24 Poland Lower Odra River 48230 Anonymous (1887-1920) 33 1887 26 Lithuania Curonian Lagoon 252473 Anonymous (1887-1920) 34 1887 26 Poland Vistula Lagoon 153594 Anonymous (1887-1920) 35 1888 24 Germany Szczecin Lagoon 13356 Anonymous (1887-1920) 36 1888 26 Lithuania Curonian Lagoon 58796 Anonymous (1887-1920) 37 1888 26 Poland Vistula Lagoon 66131 Anonymous (1887-1920) 38 1889 24 Germany Szczecin Lagoon 11130 Anonymous (1887-1920) 39 1889 24 Poland Odra River 1 NHMB 40 1889 26 Lithuania Curonian Lagoon 126000 Anonymous (1887-1920) 41 1889 26 Poland Vistula Lagoon 56997 Anonymous (1887-1920) 42 1890 24 Germany Szczecin Lagoon 14098 Anonymous (1887-1920) 43 1890 24 Poland Lower Odra River 97202 Anonymous (1887-1920) 44 1890 26 Lithuania Curonian Lagoon 92668 Anonymous (1887-1920) 45 1890 26 Poland Vistula Lagoon 69418 Anonymous (1887-1920) 46 1892 24 Germany Szczecin Lagoon 17808 Anonymous (1887-1920) 47 1892 24 Poland Lower Odra River 41552 Anonymous (1887-1920)

16 Sub- Number of No. Year Country Locality Data source division individuals

48 1892 26 Lithuania Curonian Lagoon 142343 Anonymous (1887-1920) 49 1892 26 Poland Vistula Lagoon 40035 Anonymous (1887-1920) 50 1893 25 Sweden North east of Kristianstad 3 NRMS 51 1894 24 Germany Szczecin Lagoon 29680 Anonymous (1887-1920) 52 1894 24 Poland Lower Odra River 267120 Anonymous (1887-1920) 53 1894 26 Lithuania Curonian Lagoon 169649 Anonymous (1887-1920) 54 1894 26 Poland Vistula Lagoon 34319 Anonymous (1887-1920) 55 1895 24 Poland Lower Odra River 242040 Anonymous (1887-1920) 56 1895 26 Lithuania Curonian Lagoon 120237 Anonymous (1887-1920) 57 1895 26 Poland Vistula Lagoon 16082 Anonymous (1887-1920) 58 1895 26 Poland Mouth of Vistula River 383641 Anonymous (1887-1920) 59 1896 24 Germany Lower Warnow River unknown Schaarschmidt & Lemcke (2004) 60 1896 24 Poland Lower Odra River 409287 Anonymous (1887-1920) 61 1896 26 Lithuania Curonian Lagoon 156622 Anonymous (1887-1920) 62 1896 26 Poland Vistula Lagoon 28581 Anonymous (1887-1920) 63 1896 26 Poland Mouth of Vistula River 193470 Anonymous (1887-1920) 64 1897 21 Denmark Kattegat 1 ZMUC 65 1897 24 Germany Lower Warnow River unknown Schaarschmidt & Lemcke (2004) 66 1897 24 Poland Lower Odra River 423534 Anonymous (1887-1920) 67 1897 26 Lithuania Curonian Lagoon 131471 Anonymous (1887-1920) 68 1897 26 Poland Vistula Lagoon 176981 Anonymous (1887-1920) 69 1897 26 Poland Mouth of Vistula River 446079 Anonymous (1887-1920) 70 1898 21 Denmark Gilleleje 1 ZMUC 71 1898 22 Germany Neustadt Bight 2 ZMUH 72 1898 24 Poland Lower Odra River 248125 Anonymous (1887-1920) 73 1898 26 Lithuania Curonian Lagoon 135517 Anonymous (1887-1920) 74 1898 26 Poland Vistula Lagoon 76508 Anonymous (1887-1920) 75 1898 26 Poland Mouth of Vistula River 443661 Anonymous (1887-1920) 76 1899 22 Denmark Langeland Belt 1 ZMUC 77 1899 24 Germany Fischerbruch near Rostock 1 GOMS 78 1899 24 Poland Lower Odra River 603098 Anonymous (1887-1920) 79 1899 26 Lithuania Curonian Lagoon 157997 Anonymous (1887-1920) 80 1899 26 Poland Vistula Lagoon 129273 Anonymous (1887-1920) 81 1899 26 Poland Mouth of Vistula River 1055509 Anonymous (1887-1920) 82 1900 24 Poland Lower Odra River 298121 Anonymous (1887-1920) 83 1900 26 Lithuania Curonian Lagoon 14895 Anonymous (1887-1920) 84 1900 26 Poland Vistula Lagoon 205561 Anonymous (1887-1920) 85 1900 26 Poland Mouth of Vistula River 333625 Anonymous (1887-1920) 86 1901 24 Poland Lower Odra River 189180 Anonymous (1887-1920) 87 1901 26 Lithuania Curonian Lagoon 25701 Anonymous (1887-1920) 88 1901 26 Poland Vistula Lagoon 282510 Anonymous (1887-1920) 89 1901 26 Poland Mouth of Vistula River 367977 Anonymous (1887-1920) 90 1902 24 Poland Lower Odra River 119269 Anonymous (1887-1920) 91 1902 26 Lithuania Curonian Lagoon 49687 Anonymous (1887-1920) 92 1902 26 Poland Vistula Lagoon 40453 Anonymous (1887-1920) 93 1902 26 Poland Mouth of Vistula River 177311 Anonymous (1887-1920) Waters around Ruegen 94 1903 24 Germany 1 ZMUH Island 95 1903 24 Poland Lower Odra River 83772 Anonymous (1887-1920) 96 1903 26 Lithuania Curonian Lagoon 85083 Anonymous (1887-1920) 97 1903 26 Poland Vistula Lagoon 81565 Anonymous (1887-1920)

17 Sub- Number of No. Year Country Locality Data source division individuals

98 1903 26 Poland Mouth of Vistula River 255413 Anonymous (1887-1920) 99 1904 24 Poland Lower Odra River 85018 Anonymous (1887-1920) 100 1904 26 Lithuania Curonian Lagoon 27701 Anonymous (1887-1920) 101 1904 26 Poland Vistula Lagoon 73870 Anonymous (1887-1920) 102 1904 26 Poland Mouth of Vistula River 301087 Anonymous (1887-1920) 103 1905 24 Poland Lower Odra River 221442 Anonymous (1887-1920) 104 1905 26 Lithuania Curonian Lagoon 27591 Anonymous (1887-1920) 105 1905 26 Poland Vistula Lagoon 104100 Anonymous (1887-1920) 106 1905 26 Poland Mouth of Vistula River 276684 Anonymous (1887-1920) 107 1906 24 Poland Lower Odra River 74304 Anonymous (1887-1920) 108 1906 26 Lithuania Curonian Lagoon 40343 Anonymous (1887-1920) 109 1906 26 Poland Vistula Lagoon 43311 Anonymous (1887-1920) 110 1906 26 Poland Mouth of Vistula River 218917 Anonymous (1887-1920) 111 1907 24 Poland Lower Odra River 42294 Anonymous (1887-1920) 112 1907 26 Lithuania Curonian Lagoon 36825 Anonymous (1887-1920) 113 1907 26 Poland Vistula Lagoon 24623 Anonymous (1887-1920) 114 1907 26 Poland Mouth of Vistula River 452785 Anonymous (1887-1920) 115 1908 24 Poland Lower Odra River 39296 Anonymous (1887-1920) 116 1908 26 Lithuania Curonian Lagoon 71012 Anonymous (1887-1920) 117 1908 26 Poland Vistula Lagoon 17698 Anonymous (1887-1920) 118 1908 26 Poland Mouth of Vistula River 436681 Anonymous (1887-1920) 119 1909 24 Germany Lower Warnow River unknown Schaarschmidt & Lemcke (2004) 120 1909 24 Poland Lower Odra River 28196 Anonymous (1887-1920) 121 1909 26 Lithuania Curonian Lagoon 34956 Anonymous (1887-1920) 122 1909 26 Poland Vistula Lagoon 26712 Anonymous (1887-1920) 123 1909 26 Poland Mouth of Vistula River 360722 Anonymous (1887-1920) 124 1910 24 Poland Odra River 1 NHMB 125 1910 24 Poland Lower Odra River 47948 Anonymous (1887-1920) 126 1910 26 Lithuania Curonian Lagoon 53424 Anonymous (1887-1920) 127 1910 26 Poland Vistula Lagoon 54908 Anonymous (1887-1920) 128 1910 26 Poland Mouth of Vistula River 304693 Anonymous (1887-1920) 129 1911 24 Poland Lower Odra River 60903 Anonymous (1887-1920) 130 1911 26 Lithuania Curonian Lagoon 26547 Anonymous (1887-1920) 131 1911 26 Poland Vistula Lagoon 73925 Anonymous (1887-1920) 132 1911 26 Poland Mouth of Vistula River 390237 Anonymous (1887-1920) 133 1912 20 Denmark Skagen 1 ZMUC 134 1912 24 Poland Lower Odra River 132566 Anonymous (1887-1920) 135 1912 26 Lithuania Curonian Lagoon 17698 Anonymous (1887-1920) 136 1912 26 Poland Vistula Lagoon 101616 Anonymous (1887-1920) 137 1912 26 Poland Mouth of Vistula River 370725 Anonymous (1887-1920) 138 1913 24 Poland Lower Odra River 60518 Anonymous (1887-1920) 139 1913 26 Lithuania Curonian Lagoon 35066 Anonymous (1887-1920) 140 1913 26 Poland Vistula Lagoon 21447 Anonymous (1887-1920) 141 1913 26 Poland Mouth of Vistula River 467405 Anonymous (1887-1920) Szczecin Lagoon near 142 1914 24 Germany unknown Schaarschmidt & Lemcke (2004) Ueckermuende 143 1914 24 Poland Lower Odra River 14647 Anonymous (1887-1920) 144 1914 26 Lithuania Curonian Lagoon 8904 Anonymous (1887-1920) 145 1914 26 Poland Vistula Lagoon 21985 Anonymous (1887-1920) 146 1914 26 Poland Mouth of Vistula River 614486 Anonymous (1887-1920) 147 1915 24 Poland Lower Odra River 12822 Anonymous (1887-1920)

18 Sub- Number of No. Year Country Locality Data source division individuals

148 1915 26 Lithuania Curonian Lagoon 10642 Anonymous (1887-1920) 149 1915 26 Poland Mouth of Vistula River 578700 Anonymous (1887-1920) 150 1916 24 Poland Lower Odra River 12807 Anonymous (1887-1920) 151 1916 26 Lithuania Curonian Lagoon 29680 Anonymous (1887-1920) 152 1916 26 Poland Vistula Lagoon 32871 Anonymous (1887-1920) 153 1916 26 Poland Mouth of Vistula River 217109 Anonymous (1887-1920) 154 1917 24 Poland Lower Odra River 5491 Anonymous (1887-1920) 155 1917 26 Lithuania Curonian Lagoon 36506 Anonymous (1887-1920) 156 1917 26 Poland Vistula Lagoon 13059 Anonymous (1887-1920) 157 1917 26 Poland Mouth of Vistula River 441342 Anonymous (1887-1920) 158 1918 26 Lithuania Curonian Lagoon 129702 Anonymous (1887-1920) 159 1918 26 Poland Mouth of Vistula River 352109 Anonymous (1887-1920) 160 1919 26 Lithuania Curonian Lagoon 99428 Anonymous (1887-1920) 161 1919 26 Poland Vistula Lagoon 13133 Anonymous (1887-1920) 162 1920 26 Poland Vistula Lagoon 7806 Anonymous (1887-1920) 163 1923 25 Poland Kolberg 2 NHMB 164 1929 22 Denmark Vejle Fjord 1 ZMUH 165 1930 22 Denmark Great Belt 1 ZMUC 166 1930 24 Germany Upper Warnow River unknown Schaarschmidt & Lemcke (2004) 167 1932 23 Denmark Koge Bight 1 ZMUC 168 1932 26 Poland Vistula Lagoon 10360 Eichelbaum (1926-1940) 169 1933 26 Poland Vistula Lagoon 23680 Eichelbaum (1926-1940) 170 1934 24 Germany Saal Bodden unknown Schaarschmidt & Lemcke (2004) 171 1934 26 Poland Vistula Lagoon 37000 Eichelbaum (1926-1940) 172 1935 26 Poland Vistula Lagoon 39960 Eichelbaum (1926-1940) 173 1936 26 Poland Vistula Lagoon 7400 Eichelbaum (1926-1940) 174 1937 26 Poland Vistula Lagoon 11840 Eichelbaum (1926-1940) 175 1937 26 Poland Vistula River 1 ZMUH 176 1938 26 Lithuania Curonian Lagoon 13320 Eichelbaum (1926-1940) 177 1938 26 Poland Vistula Lagoon 13320 Eichelbaum (1926-1940) 178 1939 22 Denmark Little Belt 1 ZMUC 179 1939 22 Germany Luebeck 10 Duncker (1960) 180 1939 22 Germany Schlei 10 Duncker (1960) 181 1939 22 Germany Schwartau 10 Duncker (1960) 182 1939 22 Germany Stecknitz 10 Duncker (1960) 183 1939 22 Germany Trave River 100 Duncker (1960) 184 1939 22 Germany Wakenitz 10 Duncker (1960) 185 1939 26 Lithuania Curonian Lagoon 4440 Eichelbaum (1926-1940) 186 1941 24 Germany Szczecin Lagoon unknown Schaarschmidt & Lemcke (2004) 187 1955 24 Germany Greifswald Bodden unknown Schaarschmidt & Lemcke (2004) 188 1955 26 Lithuania Curonian Lagoon 8000 Gaigalas (2001) 189 1956 26 Lithuania Curonian Lagoon 8000 Gaigalas (2001) 190 1957 22 Germany Trave River 1 ZMUH 191 1957 26 Lithuania Curonian Lagoon 8000 Gaigalas (2001) 192 1958 26 Lithuania Curonian Lagoon 8000 Gaigalas (2001) 193 1959 26 Lithuania Curonian Lagoon 8000 Gaigalas (2001) 194 1960 24 Germany Ralswiek unknown Schaarschmidt & Lemcke (2004) 195 1960 26 Lithuania Curonian Lagoon 6800 Gaigalas (2001) 196 1961 26 Lithuania Curonian Lagoon 6800 Gaigalas (2001) 197 1962 26 Lithuania Curonian Lagoon 6800 Gaigalas (2001)

19 Sub- Number of No. Year Country Locality Data source division individuals

198 1963 26 Lithuania Curonian Lagoon 6800 Gaigalas (2001) 199 1964 26 Lithuania Curonian Lagoon 6800 Gaigalas (2001) 200 1965 26 Lithuania Curonian Lagoon 53200 Gaigalas (2001) 201 1966 26 Lithuania Curonian Lagoon 53200 Gaigalas (2001) 202 1967 26 Lithuania Curonian Lagoon 53200 Gaigalas (2001) 203 1968 26 Lithuania Curonian Lagoon 53200 Gaigalas (2001) 204 1969 26 Lithuania Curonian Lagoon 53200 Gaigalas (2001) 205 1970 24 Germany Barth Bodden unknown Schaarschmidt & Lemcke (2004) 206 1970 26 Lithuania Curonian Lagoon 69600 Gaigalas (2001) 207 1970 26 Poland Vistula River 327870 Bartel (1993) 208 1971 26 Lithuania Curonian Lagoon 69600 Gaigalas (2001) 209 1971 26 Poland Vistula River 299110 Bartel (1993) 210 1972 26 Lithuania Curonian Lagoon 69600 Gaigalas (2001) 211 1972 26 Poland Vistula River 241730 Bartel (1993) Szczecin Lagoon near 212 1973 24 Germany 20 Schaarschmidt & Lemcke (2004) Altwarp 213 1973 24 Germany Strelasund 1 GOMS 214 1973 26 Lithuania Curonian Lagoon 69600 Gaigalas (2001) 215 1973 26 Poland Vistula River 115380 Bartel (1993) 216 1974 24 Germany Prohner Wiek 1 GOMS 217 1974 26 Lithuania Curonian Lagoon 69600 Gaigalas (2001) 218 1974 26 Poland Vistula River 7730 Bartel (1993) 219 1975 26 Lithuania Curonian Lagoon 2600 Gaigalas (2001) 220 1975 26 Poland Vistula River 840 Bartel (1993) 221 1976 26 Lithuania Curonian Lagoon 2600 Gaigalas (2001) 222 1976 26 Poland Vistula River 32550 Bartel (1993) 223 1977 26 Lithuania Curonian Lagoon 2600 Gaigalas (2001) 224 1977 26 Poland Vistula River 95150 Bartel (1993) 225 1978 26 Lithuania Curonian Lagoon 2600 Gaigalas (2001) 226 1979 26 Lithuania Curonian Lagoon 2600 Gaigalas (2001) 227 1980 24 Germany Greifswald Bodden unknown Schaarschmidt & Lemcke (2004) 228 1981 22 Germany Wallensteingraben unknown Schaarschmidt & Lemcke (2004) 229 1981 24 Germany Schmachter See unknown Schaarschmidt & Lemcke (2004) 230 1981 24 Germany Szczecin Lagoon unknown Schaarschmidt & Lemcke (2004) 231 1982 24 Germany Zicker 1 DBUR 232 1984 24 Germany Wehrland 1 ZMUG 233 1985 26 Lithuania Curonian Lagoon 800 Gaigalas (2001) 234 1986 24 Germany River Peene 1 NHMB 235 1986 24 Germany River Warnow unknown Schaarschmidt & Lemcke (2004) 236 1986 24 Germany Wehrland 1 ZMUG 237 1986 24 Germany Wehrland 30 Schaarschmidt & Lemcke (2004) 238 1986 24 Poland Odra River 1 NHMB 239 1986 26 Lithuania Curonian Lagoon 800 Gaigalas (2001) 240 1987 24 Germany Wehrland 60 Schaarschmidt & Lemcke (2004) 241 1987 26 Lithuania Curonian Lagoon 800 Gaigalas (2001) North east of 242 1988 24 Germany 9 Schröder (1995) Stubbenkammer 243 1988 26 Lithuania Curonian Lagoon 800 Gaigalas (2001) 244 1988 26 Poland Vistula River 7500 Bartel (1993) Szczecin Lagoon near 245 1989 24 Germany 100 Waterstraat & Krappe (2000) Ueckermuende 246 1989 25 Poland Grabowska River unknown Witkowski (1996)

20 Sub- Number of No. Year Country Locality Data source division individuals

247 1989 25 Poland Parseta River unknown Witkowski (1996) 248 1989 25 Poland Rega River unknown Witkowski (1996) 249 1989 26 Lithuania Curonian Lagoon 800 Gaigalas (2001) 250 1989 26 Poland Pasleka River unknown Witkowski (1996) 251 1989 26 Poland Vistula River 7000 Bartel (1993)

Table 3. Mean annual catch (kg) of river lamprey in the entire southern Baltic Sea and in selected areas in different time periods from 1887 - 1999.

Subdivision 24 Subdivision 26 Lower Vistula River, Total catch Szczecin Lagoon Curonian Lagoon Vistula Lagoon and Southern Baltic Sea and adjacent waters and Nemunas Delta Bay of Gdańsk

1887-1889 18,564.33 2,526.67 6,215.67 9,822.00 1890-1899 44,430.40 16,129.00 20,845.20 7,456.20 1900-1909 38,249.90 7,957.50 27,504.00 2,788.40 1910-1919 32,793.60 2,343.00 27,434.40 3,016.20 1920-1929 52.60 0.00 52.60 0.00 1930-1939 1,090.00 0.00 970.00 120.00 1950-1959 400.00 0.00 0.00 400.00 1960-1969 9,000.00 0.00 6,000.00 3,000.00 1970-1979 14,813.60 0.00 11,203.60 3,610.00 1980-1989 595.60 0.00 555.60 40.00 1990-1999 1,083.60 45.00 198.60 840.00 1887-1999 14,376.56 2,643.93 9,381.43 2,351.20 100% 18.39% 65.26% 16.35%

21 Table 4. Recent records of sea lamprey in German waters of the southern Baltic Sea from 1990 - 2005. For abbreviations see chapter “Material and Methods”.

Total Sub- Number of No. Year Locality length Data source division Individuals (cm) 1 1991 24 Strelasund near Stralsund 1 49 Schröder (1995) 2 1992 24 Strelasund 1 85 Schröder (1995) 3 1998 24 Lower Warnow River 1 DBUR 4 1999 22 Kiel Bight 1 ZMUK 5 2000 22 Neustadt Bight 1 Spratte (2004) 6 2001 24 Coastal waters of Usedom Island 1 43 cm Thiel et al. (2005) 7 2002 22 Flensburg Bight 1 50 Spratte (2004) 8 2002 22 Schlei unknown Spratte (2004) 9 2003 22 Howacht Bight 1 64.5 Thiel et al. (2004) 10 2003 24 Warnow River 1 -- Thiel et al. (2005) 11 2004 22 Dassower See 1 -- Thiel et al. (2005) 12 2005 22 Poel Island near Gollwitz 1 60 Thiel et al. (2005) 13 2005 24 Peenestrom 1 80 Thiel et al. (2005)

Table 5. Recent records of river lamprey in German and adjacent Polish waters of the southern Baltic Sea. For abbreviations see chapter “Material and Methods”.

Total Sub- Number of No. Year Country Locality length Data source division individuals (cm) 1 1990 24 Poland Odra River 1 NHMB 2 1990 24 Poland Szczecin Lagoon 4500 Raczyinski & Filipiak (1999) 3 1994 24 Germany Ruden Island 50 max. 35 Schröder (1995) 4 1994 24 Germany Usedom Island 50 max. 35 Schröder (1995) 5 1995 24 Germany North of Ruegen Island 1 GOMS 6 1996 24 Germany North east of Ruegen Island 1 GOMS 7 1997 22 Germany Salzhaff 1 GOMS 8 1997 24 Germany East of Ruegen Island 1 GOMS 9 1997 24 Poland Lake Dabie 35 Raczyinski & Filipiak (1999) 10 1998 24 Poland Lake Dabie 53 Raczyinski & Filipiak (1999) 11 1999 24 Germany Libnower Muehlbach 2000 Waterstraat & Krappe (2000) 12 1999 24 Germany North of Ruegen Island 1 Kloppmann et al. (2003) 13 2003 24 Germany Coastal Waters of Usedom Island 1 appr.35 Thiel et al. (2004) 14 2003 24 Germany Szczecin Lagoon near Altwarp 1 adult Thiel et al. (2005) 15 2003 24 Germany Warnow River 6 adult Thiel et al. (2005) 16 2004 24 Germany Krumminer Wiek 1 36 Thiel et al. (2005) 17 2004 24 Germany Krumminer Wiek 3 30-35 Thiel et al. (2005) 18 2004 24 Germany Krumminer Wiek 7 30-40 Thiel et al. (2005) 19 2004 24 Germany Krumminer Wiek 1 38 Thiel et al. (2005) 20 2004 24 Germany Krumminer Wiek 1 35 Thiel et al. (2005) 21 2004 24 Germany Krumminer Wiek 1 40 Thiel et al. (2005) 22 2004 24 Germany Krumminer Wiek 1 34 Thiel et al. (2005) 23 2004 24 Germany Krumminer Wiek 1 36 Thiel et al. (2005) 24 2004 24 Germany Krumminer Wiek 1 32 Thiel et al. (2005) 25 2004 24 Germany Krumminer Wiek 1 38 Thiel et al. (2005) 26 2004 24 Germany Krumminer Wiek 3 30-34 Thiel et al. (2005) 27 2004 24 Germany Krumminer Wiek 2 33; 40 Thiel et al. (2005)

22 Total Sub- Number of No. Year Country Locality length Data source division individuals (cm) 28 2004 24 Germany Krumminer Wiek 1 40 Thiel et al. (2005) 29 2004 24 Germany Krumminer Wiek 2 38; 40 Thiel et al. (2005) 30 2004 24 Germany Krumminer Wiek 1 33 Thiel et al. (2005) 31 2004 24 Germany Krumminer Wiek 1 34 Thiel et al. (2005) 32 2004 24 Germany Krumminer Wiek 1 32 Thiel et al. (2005) 33 2004 24 Germany Krumminer Wiek 1 37 Thiel et al. (2005) 34 2004 24 Germany Krumminer Wiek 3 31-35 Thiel et al. (2005) 35 2004 24 Germany Krumminer Wiek 2 40 Thiel et al. (2005) 36 2004 24 Germany Krumminer Wiek 1 31 Thiel et al. (2005) 37 2004 24 Germany Krumminer Wiek 1 37 Thiel et al. (2005) 38 2004 24 Germany Krumminer Wiek 3 30-35 Thiel et al. (2005) 39 2004 24 Germany Krumminer Wiek 2 37; 40 Thiel et al. (2005) 40 2004 24 Germany North west of Adlerground 1 43 Thiel et al. (2004 41 2004 24 Germany Recknitz River 1 -- Thiel et al. (2005) 42 2004 24 Germany Szczecin Lagoon near Ueckermuende 1 39 Thiel et al. (2005) 43 2004 24 Germany Szczecin Lagoon near Ueckermuende 1 36 Thiel et al. (2005) 44 2004 24 Germany Szczecin Lagoon near Ueckermuende 1 38 Thiel et al. (2005) 45 2004 24 Germany Szczecin Lagoon near Ueckermuende 1 36 Thiel et al. (2005) 46 2004 24 Germany Szczecin Lagoon near Ueckermuende 1 36.5 Thiel et al. (2005) 47 2004 24 Germany Szczecin Lagoon near Ueckermuende 1 37.5 Thiel et al. (2005) 48 2004 24 Germany Szczecin Lagoon near Ueckermuende 1 38 Thiel et al. (2005) 49 2004 24 Germany Szczecin Lagoon near Ueckermuende 1 32 Thiel et al. (2005) 50 2004 24 Germany Szczecin Lagoon near Ueckermuende 1 35 Thiel et al. (2005) 51 2004 24 Germany Szczecin Lagoon near Ueckermuende 1 35 Thiel et al. (2005) 52 2004 24 Germany Szczecin Lagoon near Ueckermuende 23 32 - 42 Thiel et al. (2005) 53 2004 24 Germany Szczecin Lagoon near Ueckermuende 1 42 Thiel et al. (2005) 54 2004 24 Germany Szczecin Lagoon near Ueckermuende 1 36.5 Thiel et al. (2005) 55 2004 24 Germany Szczecin Lagoon near Ueckermuende 3 -- Thiel et al. (2005) 56 2004 24 Germany Szczecin Lagoon near Ueckermuende 1 35 Thiel et al. (2005) 57 2004 24 Germany Szczecin Lagoon near Ueckermuende 1 34 Thiel et al. (2005) 58 2004 24 Germany Szczecin Lagoon near Ueckermuende 1 40 Thiel et al. (2005) 59 2004 24 Germany Szczecin Lagoon near Ueckermuende 1 38.5 Thiel et al. (2005) 60 2004 24 Germany Szczecin Lagoon near Ueckermuende 1 40.5 Thiel et al. (2005) 61 2004 24 Germany Szczecin Lagoon near Ueckermuende 1 37 Thiel et al. (2005) 62 2004 24 Germany Szczecin Lagoon near Ueckermuende 1 41 Thiel et al. (2005) 63 2004 24 Germany Szczecin Lagoon near Ueckermuende 1 35 Thiel et al. (2005) 64 2004 24 Germany Szczecin Lagoon near Ueckermuende 1 38.5 Thiel et al. (2005) 65 2004 24 Germany Szczecin Lagoon near Ueckermuende 1 38 Thiel et al. (2005) 66 2004 24 Germany Szczecin Lagoon near Ueckermuende 1 40 Thiel et al. (2005)

23 Table 6. Location of detected spawning sites of river lamprey in German tributaries of the southern Baltic Sea.

No. River System Location Data source

1 River Peene tributaries Scheidegraben Winkler et al. (1999) 2 River Peene tributaries Augraben Waterstraat & Krappe (2000) 3 River Peene tributaries Galgenbach Waterstraat & Krappe (2000) 4 River Peene tributaries Klenzer Muehlbach Waterstraat & Krappe (2000) 5 River Peene tributaries Libnower Muehlbach Waterstraat & Krappe (2000) 6 River Warnow tributaries Beke Winkler et al. (1999) 7 River Warnow tributaries Koesterbeck Winkler et al. (1999) 8 River Warnow tributaries Muehlendamm canal near Rostock City Winkler pers. comm. (2005) 9 River Stepenitz tributaries Radegast River Winkler et al. (1999)

Fig. 1. Adult river lamprey recorded from commercial fishery in the Peenestrom near the Szczecin Lagoon from August 2004.

Fig. 2. Largest individuals of sea lamprey recorded from the ichthyological collection of the German Oceanographic Museum in the city of Stralsund (GOMS). Above: Record from the Greifswald Bodden from April 1972; Below: Record from the Greifswald Bodden from February 1982.

Fig. 3. Distribution of historical records of sea lamprey in subdivisions 20 - 26 of the Baltic Sea in the time period 1829 - 1939.

Fig. 4. Distribution of historical records of sea lamprey in subdivisions 20 – 26 of the Baltic Sea in the time period 1940 – 1989.

Fig. 5. Distribution of historical records of river lamprey in subdivisions 20 – 26 of the Baltic Sea in the time period 1649 – 1939.

Fig. 6. Distribution of historical records of river lamprey in subdivisions 20 – 26 of the Baltic Sea in the time period 1940 – 1989.

Fig. 7. Total landings of lampreys in the southern Baltic Sea from 1887 – 1999

Fig. 8. Contribution (%) of the main fishery areas in the southern Baltic Sea to the total annual yield of lampreys for different time periods from 1887 - 1999.

Fig. 8. Contribution (%) of the main fishery areas in the southern Baltic Sea to the total annual yield of lampreys for different time periods from 1887 – 1999.

Fig. 9. Seasonal variation of total catches of lamprey in three different fishery areas of the southern Baltic Sea from 1887 - 1920.

Fig. 10. Distribution of recent records of sea lamprey in German Baltic waters from 1990 - 2005.

Fig. 11. Distribution of recent records of river lamprey in German and adjacent Polish Baltic waters (circles) and location of detected spawning sites (triangles) of river lamprey in German tributaries of the southern Baltic Sea.