Microchemical Analyses of Otoliths in Baltic Sea Fish

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Microchemical Analyses of Otoliths in Baltic Sea Fish Microchemical analyses of otoliths in Baltic Sea fish -Possibilities and limitations of otolith elemental analysis to describe individual life history and stock characteristics of fish in the Baltic Sea- Dissertation zur Erlangung des Doktorgrades der Mathematisch-Naturwissenschaftlichen Fakultät der Christian-Albrechts-Universität zu Kiel vorgelegt von Lasse Marohn Kiel, Juli 2011 Referent: PD Dr. Reinhold Hanel Korreferent: Prof. Dr. Carsten Schulz Tag der mündlichen Prüfung: 04.10.2011 Zum Druck genehmigt: Kiel, Der Dekan SUMMARY In this thesis otolith microchemistry analyses were used to gain insights into the individual life history and stock characteristics of three fish species from the Baltic Sea - the European eel Anguilla anguilla, the Atlantic cod Gadus morhua and the thicklip grey mullet Chelon labrosus. The special hydrographic environment of the world’s largest brackish water system provide promising conditions for the use of otolith elemental analysis to investigate individual migration patterns and stock structures of fish. Here, it was used to gain information with relevance for stock management of fish species that differ widely in their biology, ecology and stock structure. In chapter I the influence of continental migratory behaviour on health and spawner quality of the European eel was analysed. Otolith strontium (Sr) composition was used to identify characteristic migration patterns. Results show that the muscle fat contents of silver eels with strictly catadromous life cycles are significantly reduced compared to silver eels that never entered freshwaters. Furthermore, prevalence and infection intensities of the swimbladder nematode Anguillicoloides crassus are highly increased in catadromous silver eels. Both, a reduced accumulation of fat reserves and intense A. crassus infestations are assumed to impair the successful reproduction of A. anguilla. These results highlight the importance of brackish water habitats for the production of high quality spawners and question the benefit of restocking measures into inland waters. In order to investigate the influence of water temperature and feeding behaviour on the element incorporation into A. anguilla otoliths, two experimental studies were conducted (Chapter II & III). It was investigated whether experienced temperature history and individual food preferences can be reconstructed by otolith microchemistry analysis. It was further tested if temperature or diet affect the incorporation of Sr into otoliths to such a degree that the validity of otolith Sr based migration studies is impaired. Therefore, juvenile eels were reared for 15 weeks at three different water temperatures (14°C, 19°C, 24°C) and the elemental composition of the newly grown aragonite was subsequently analysed. Although the otolith composition differed significantly among treatments, detected differences were low and an in situ reconstruction of temperature history seems challenging. Similar results were obtained in the feeding experiment. Feeding of eight different diets during eight weeks caused no detectable differences in otolith elemental composition among treatments. Hence, the use of otolith microchemistry to determine individual food preferences of European eels seems to be unfeasible. However, the results presented indicate that neither temperature changes nor individual dietary behaviour impair the use of otolith Sr concentrations as a tracer of diadromy. Thus the reliability of such migration studies is fundamentally increased. i SUMMARY In chapter IV it was investigated whether multi-element otolith analysis can be used to discriminate Atlantic cod individuals according to their origin. Therefore, the concentrations of 16 elements were determined along the growth axis of otoliths from adult individuals caught at spawning grounds in the North Sea, the western Baltic Sea and the eastern Baltic Sea. Furthermore, the multi-element composition of the core region of juvenile cod otoliths from western and eastern Baltic areas was analysed. Multivariate statistical analyses successfully discriminated between adults from different stocks as well as between western and eastern Baltic Sea juveniles. Significant differences between the eastern Baltic spawning grounds, however, were not detected. These results demonstrate the potential of otolith microchemistry analysis to investigate the structure and connectivity of G. morhua stocks in the Baltic Sea. In chapter V the migratory behaviour of thicklip grey mullet was examined for the first time on individuals caught in the Baltic Sea. By detecting C. labrosus otolith Sr concentrations, this study aimed to gain first insights into preferences and whereabouts of individuals in the Baltic Sea. Results confirmed a preference of brackish habitats for all analysed specimens and suggest a high plasticity of C. labrosus migratory behaviour. Some individuals seem to undertake annual migrations to marine waters, while others only sporadically left brackish waters. Although the behaviour of Baltic Sea C. labrosus could not be conclusively clarified, the results support the assumption that a considerable fraction of individuals entering the western Baltic Sea regularly migrate to the North Sea. The results presented in this thesis confirm the feasibility of otolith microchemistry analyses for the investigation of a broad range of questions on Baltic Sea fish. Knowledge about investigated species was expanded and opportunities for future studies were indicated. ii ZUSAMMENFASSUNG In dieser Arbeit wurden anhand mikrochemischer Otolithenanalysen Erkenntnisse über die individuelle Lebensweise und die Bestandsstruktur von drei Fischarten aus der Ostsee gewonnen – dem Europäischen Aal Anguilla anguilla, dem Atlantischen Kabeljau (Dorsch) Gadus morhua und der Dicklippigen Meeräsche Chelon labrosus. Die hydrographischen Besonderheiten der Ostsee als weltgrößtes Brackwassersystem bieten vielversprechende Voraussetzungen für die Anwendung von Otolithen-Elementanalysen zur Untersuchung des individuellen Wanderverhaltens von Fischen und deren Bestandsstrukturen. Hier wurden sie genutzt, um managementrelevante Fragestellungen von Fischarten zu untersuchen, die sich hinsichtlich ihrer Biologie, Ökologie und Bestandsstruktur sehr unterscheiden. In Kapitel I wurde der Einfluss des kontinentalen Wanderverhaltens auf die Gesundheit und die Laicherqualität des Europäischen Aals untersucht. Dazu wurden anhand der Otolithen- Strontiumverteilung charakteristische Wandermuster ermittelt. Die Ergebnisse zeigen, dass der Muskelfettgehalt von Blankaalen mit einem strikt katadromen Wanderverhalten signifikant geringer ist als der von Blankaalen, die niemals ins Süßwasser eingewandert sind. Darüber hinaus sind Prävalenz und Befallsintensität mit dem Schwimmblasen-Nematoden Anguillicoloides crassus bei katadromen Blankaalen stark erhöht. Sowohl eine verringerte Einlagerung von Fettreserven als auch der intensive Befall mit A. crassus stehen im Verdacht, den Reproduktionserfolg von A. anguilla zu vermindern. Die vorliegenden Ergebnisse heben die Bedeutung von Brackwasserhabitaten für die Produktion von gesunden Laichtieren hervor und stellen den Nutzen von Besatzmaßnahmen in Binnengewässer in Frage. Um den Einfluss von Wassertemperatur und Futterverhalten auf die Elementeinlagerung in A. anguilla-Otolithen zu prüfen, wurden zwei experimentelle Studien durchgeführt (Kapitel II & III). Es wurde untersucht, ob erlebte Temperaturveränderungen und individuelle Futterpräferenzen anhand der im Otolithen gespeicherten Elemente rekonstruiert werden können. Ferner wurde geprüft, ob Temperatur oder Diät die Einlagerung von Strontium (Sr) in einem Maße beeinflussen, das die Aussagekraft von Otolithen-Sr basierten Migrationsstudien einschränken könnte. Dazu wurden juvenile Aale für 15 Wochen bei drei unterschiedlichen Temperaturen gehalten (14°C, 19°C, 24°C). Anschließend wurde die Elementzusammensetzung des hinzu gewachsenen Aragonits analysiert. Zwar wurden statistisch signifikante Unterschiede in der Elementzusammensetzung verschiedener Temperaturbehandlungen festgestellt, diese waren jedoch so gering, dass eine Rekonstruktion erlebter Temperaturänderungen in situ schwierig erscheint. Zu einem ähnlichen Ergebnis kam auch der Futterversuch. Die Fütterung von acht verschiedenen Diäten über einen Zeitraum von acht Wochen führte zu keinen statistisch iii ZUSAMMENFASSUNG signifikanten Unterschieden in der Elementeinlagerung. Futterpräferenzen lassen sich demnach offenbar nicht anhand der Otolithen-Elementzusammensetzung nachverfolgen. Allerdings zeigen die vorliegenden Ergebnisse, dass beim Europäischen Aal weder Temperaturschwankungen noch unterschiedliches Futterverhalten die Verwendung von Sr zur Rekonstruktion diadromen Verhaltens einschränken. Damit wird die Aussagekraft solcher Migrationsstudien deutlich erhöht. In Kapitel IV wurde untersucht, ob Otolithen-Multielementanalysen dazu geeignet sind, G. morhua-Individuen ihrer Herkunft zuzuordnen. Dazu wurden die Konzentrationen von 16 Elementen entlang der Wachstumsachse von Otolithen adulter Dorsch von Laichplätzen aus der Nordsee, der westlichen und der östlichen Ostsee untersucht. Des Weiteren wurde die Multielementzusammensetzung der Kernbereich von Otolithen juveniler Dorsche aus der westlichen und der östlichen Ostsee analysiert. Multivariate statistische Analysen konnten sowohl die adulten Bestände erfolgreich trennen, als auch die Juvenilen aus der westlichen und der östlichen Ostsee unterscheiden. Signifikante Unterschiede zwischen
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