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History Characteristics of Ide Leuciscus Idus in the Eastern Baltic Sea M

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History Characteristics of Ide Leuciscus Idus in the Eastern Baltic Sea M Fisheries Management and Ecology Fisheries Management and Ecology, 2015 Life-history characteristics of ide Leuciscus idus in the Eastern Baltic Sea M. ROHTLA, R. SVIRGSDEN, I. TAAL, L. SAKS, R. ESCHBAUM & M. VETEMAA Estonian Marine Institute, University of Tartu, Tartu, Estonia Abstract Otolith Sr:Ca and Ba:Ca profiles were used in parallel with age data to investigate the life-history characteristics of ide, Leuciscus idus (L.) (n = 111), in the V€ainameri Sea, West Estonia. Sr:Ca profiles were more variable and useful than Ba:Ca profiles. Flexible life-history patterns were observed within and among the three study sites. Most of the individuals (72%) hatched in semi-enclosed bays that are fresh water during spring spawning but are often flooded with brackish water during other seasons. The importance of lotic spawning varied among sites and was the highest (88%) in Matsalu Bay, moderate in Saunja Bay (33%) and lowest (0%) in K€aina Bay. Young of the year emigrated from natal sites and entered the sea within the first summer; 95% did so during the first month post- hatch. Juvenile ide undertook non-spawning, freshwater migrations in the following spring; however, the reasons behind this phenomenon remain unknown. As the importance of lotic spawning has significantly decreased and multiple historically important ide spawning rivers lack anadromous runs altogether, it is suggested that actions should be taken to aid the recovery of those imperilled spawning stocks. KEYWORDS: age, brackish water, cyprinid, Estonia, migration, Sr:Ca and Ba:Ca profiles. cyprinid, ide Leuciscus idus (L.) (Adjers et al. 2006), Introduction which are at the historical lows in multiple areas The Baltic Sea is one of the largest brackish water bodies (Vetemaa et al. 2006; Eschbaum et al. 2014). in the world. This ecosystem consists of life with marine Ide is generally considered to be a rheophilic, fresh- and freshwater origin and thrives on high productivity water species that inhabits large lowland rivers and lakes of these waters. Unfortunately, this high production (J€arvalt et al. 2003; Winter & Fredrich 2003; Kuliskova comes at a price as most of the regions are affected by et al. 2009). However, similar to many other freshwater eutrophication caused by nutrient pollution from the species inhabiting the brackish Baltic Sea, ide has catchment areas (HELCOM 2010). Eutrophication, adapted to life in the sea and to undertake anadromous together with overfishing and climate change, facilitate spawning migrations to fresh waters (Muller€ & Berg alterations in ecosystem function (Mollmann€ et al. 2008; 1982; J€arvalt et al. 2003). Some populations of such Eriksson et al. 2009). For example, the numbers of species (e.g. northern pike Esox lucius L.) even have piscivorous fishes have gone down in many regions totally shifted to spawning in the brackish environment (Nilsson et al. 2004; Lehtonen et al. 2009; Ljunggren and have lost contact with fresh waters altogether (Eng- et al. 2010), whereas the opposite trend is evident for stedt et al. 2010). Similarly, some disputable evidence prey species such as roach, Rutilus rutilus (L.), silver indicates that ide may have reproduced in the brackish bream, Blicca bjoerkna (L.), gibel carp, Carassius coastal waters of Estonia 30–40 years ago (Mikelsaar gibelio (Bloch), and threespine stickleback, Gasterosteus 1984). General knowledge on the migratory biology of aculeatus L. (Nilsson et al. 2004; Vetemaa et al. 2005, ide is relatively scarce for both freshwater resident (Win- 2006). This has produced a regime shift in the food ter & Fredrich 2003; Kuliskova et al. 2009) and anadro- chain where top-down control has weakened and mous ide (Cala 1975; Eriksson & Muller€ 1982; Johnson bottom-up control prevails (Mollmann€ et al. 2008; 1982). In the most extensive study on anadromous ide, Ljunggren et al. 2010). Interestingly, the current state Cala (1975) studied a population living near the dynamic does not include the population growth of another Øresund Strait and stated that the anadromous runs may Correspondence: M. Rohtla, Estonian Marine Institute, University of Tartu, Vanemuise 46A, 51014 Tartu, Estonia (e-mail: [email protected]) © 2015 John Wiley & Sons Ltd doi: 10.1111/fme.12120 1 2 M. ROHTLA ET AL. cover ~50 km in the River K€avlingean. Potamodromous the otolith in proportion to the ambient concentrations spawning migrations >100 km are reported from large and differences between study systems exist (Elsdon lowland rivers (Winter & Fredrich 2003; Kuliskova et al. 2008; Walther & Limburg 2012). The strontium- et al. 2009). Furthermore, the few sea recaptures of ide to-calcium ratio (Sr:Ca) is the most frequently used that had been tagged in a small river near Umea (Swe- chemical marker because the Sr:Ca ratio in fresh waters den) were all widely distributed along the coast, meaning is usually several times lower than sea water (Kraus & that ide may cover long distance in the sea as well Secor 2004; Rohtla et al. 2014). This allows the move- (Johnson 1982). ments of diadromous species to be tracked because the Ide is classified as least concern in the IUCN Red List ambient Sr:Ca levels are recorded in the otoliths (Shiao (2014), whereas in Estonia, it is classified as data defi- et al. 2006; Magath et al. 2013; Rohtla et al. 2014). The cient (http://elurikkus.ut.ee). In Estonia, catches of fresh- barium-to-calcium ratio (Ba:Ca) is often used as an addi- water resident and coastal ide peaked in 1920–1930s and tional marker for interpreting Sr:Ca profiles, but with 1980s at 54 and 177 t yrÀ1, respectively (http://www. mixed results (Crook et al. 2006; Hamer et al. 2006; agri.ee). Abundances and catches have constantly Rohtla et al. 2014; Smith & Kwak 2014). By contrast to decreased since the 1980s in the coastal sea (Vetemaa Sr:Ca, the Ba:Ca ratio is generally enriched in fresh et al. 2006, 2010; Eschbaum et al. 2014). However, an waters and decreased in sea waters. increase in juvenile ide has been recorded in some In this study, life-history migrations of ide from the coastal areas in recent years (Eschbaum et al. 2014). It V€ainameri Sea were reconstructed using otolith Sr:Ca is likely that overfishing lead to the collapse of most ide and Ba:Ca profiles. Specifically, the aims were to inves- spawning stocks in Estonia (Vetemaa et al. 2001). It tigate migrations on a temporal and spatial scale using remains unclear, however, why most of those spawning combined age and chemical data, and assess potential stocks have not recovered despite a virtual cessation of spawning biome preferences, with fresh waters assumed commercial fishing for ide. Therefore, unfavourable to be the main spawning biome for ide. environmental conditions may be inhibiting recovery. For example, Eriksson and Muller€ (1982) observed large Methods numbers of adult ide migrating up a small Swedish river for spawning, but no descending juveniles were subse- Ide were sampled from three sites in the V€ainameri Sea quently recorded. Although they caught large numbers (Fig. 1). K€aina Bay (KB) is a semi-enclosed lagoon in of dead or developing eggs and newly emerged larvae, which salinity varies from 0 to 4 psu. Water level is reg- they concluded that reproduction had failed, possibly ulated by locks situated on the two outlets. KB becomes due to extreme variations in river water pH. They also a freshwater lake during spring snowmelt and is there- noted that the individuals in that spawning stock were fore used for spawning by different freshwater species. large (old) and of uniform size. It was also recently dem- During summer, water levels may drop to <50 cm, and onstrated that some low-density spawning stocks of ide in winter, the bay often freezes to the bottom. Fish in Estonia are predominantly composed of old individu- assemblage structure is shaped by predation pressure als (Rohtla et al. 2015), and this in turn may hinder suc- from common cormorant Phalacrocorax carbo sinensis cessful reproduction in those spawning stocks due to (L.) (Vetemaa et al. 2010). Nevertheless, the ide spawn- reduced egg quality (Targonska et al. 2012). Thus, it is ing stock in KB is probably the largest and most viable likely that the current state of ide is a result of complex in the entire V€ainameri Sea. There are no recent records interplay between fishing pressure, environmental fac- of ide spawning in nearby rivers, such as occurred his- tors, species-specific traits and other factors such as torically. Matsalu Bay (MB) is a relatively dynamic sys- interspecies competition and predator–prey relationships. tem with an east-to-west salinity gradient (0–6 psu) that Further studies on migration patterns, natal origins and is 10 km long, and a delta estuary formed by multiple environmental factors that are needed for successful rivers. Ide penetrates these rivers for spawning, but no spawning would help to gain more information for con- reproduction in the bay has been recorded. Saunja Bay serving, restoring and managing the anadromous stocks (SB) is a shallow and semi-enclosed, mostly fresh water, of this large bodied cyprinid. bay with three small inflowing rivers. Winter hypoxia Otolith microchemistry has recently emerged as a events occur about once every 6 years and may incite powerful tool for studying fish migration patterns (Secor major fish kills. According to local fishers, ide spawns 2010). Besides recording the age of the fish, otoliths also in SB only, despite it penetrating the rivers some dec- incorporate and retain different minor and trace elements ades ago. Together with the inflowing rivers, MB and from the water. These chemical data can be then used to SB act as important spawning and nursery grounds for infer migration patterns if elements are incorporated to many freshwater fishes in the V€ainameri Sea area.
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