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Small and Meso-Scale Distribution Patterns of Key Copepod Species in the Central Baltic Sea and Their Relevance for Larval Fish

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Small and Meso-Scale Distribution Patterns of Key Copepod Species in the Central Baltic Sea and Their Relevance for Larval Fish Small and meso-scale distribution patterns of key copepod species in the Central Baltic Sea and their relevance for larval fish survival Dissertation zur Erlangung des Doktorgrades der Mathematisch-Naturwissenschaftlichen Fakultät der Christian-Albrechts-Universität zu Kiel vorgelegt von Jörn Oliver Schmidt Kiel, März 2006 Referent: Prof. Dr. D. Schnack Ko-Referent: Prof. Dr. M. St. John Tag der mündlichen Prüfung: 27.04.2006 Zum Druck genehmigt: Kiel, Der Dekan Wenn [...] haline Schichtung in der Natur auf kurzfristige Vertikalbewegungen hemmend einwirkt, dürfte es sich nicht um ein physiologisches, sondern eher um ein psychologisches Problem handeln. Banse 1959 Nothing in the world is purely spatial or purely temporal; everything is process Blaut 1961 Summary This thesis shows different aspects of the distri- The distribution of reproducing female P. bution of zooplankton and the resulting influ- acuspes was investigated with the Video ence on larval fish survival. Plankton Recorder, an in situ imaging device. The vertical distribution of the key calanoid The distribution was closely related to high copepod species in the Bornholm Basin (Baltic salinity although low oxygen values were Sea), Pseudocalanus acuspes (Giesbrecht 1881), avoided. Based on images of female P. acuspes Temora longicornis (Müller 1792), Acartia carrying egg sacs the number of eggs per clutch bifilosa (Giesbrecht 1882), Acartia longiremis was calculated and using the temperature (Lilljeborg 1853) and the only cyclopoid dependent development time an in situ egg copepod Oithona similis (Claus 1863), were production rate was estimated. Albeit the investigated. Sampling was carried out with production was the same between years, the multiple opening/closing nets. The results naupli to female ratio was higher in 2003 showed distinct seasonal differences in the leading to the conclusion that the naupli vertical distribution for all species. For P. mortality might have been reduced. acuspes an ontogenetic migration was A simple individual based model of P. acuspes detected, with the younger stage dwelling near connected to a circulation model of the Baltic the surface and the older stages moving Sea was used to reveal possible influence of successively deeper. The adults inhabited the drift to biomass changes. The results showed water layers around the halocline. In November that in contrast to the 1980’s during the 1990’s the vertical distribution disintegrated and all the prevailing barotrophic circulation forced the stages were distributed over the whole water majority of P. acuspes out of the Bornholm column. However in this period the older Basin into easterly Basins and shallow coastal stages (C4, C5) dominated the community. A regions. However, the transport of organisms diel vertical migration was observed for T. produced in the eastern Basins (Gotland Basin longicornis in the summer month (June 2001, and Gdansk Deep) back to the Bornholm Basin July and August 2002). The migration was most was limited. Lower salinity and higher tempera- distinc for the older stages, moving from 20 ture in the near shore water might influence the metre at night to almost 60 metre at day. No survival and reproduction. diel migration was observed in April and The same circulation model was used to inves- November. For Acartia species an extended tigate the drift of larval cod and possible conse- daily migration was not observed, even though quences on the overlap with food organisms. the weighted mean depth of Acartia longiremis Again, a shift in the baroclinic circulation in July shifted from 10 metre at night to 20 between the 1980’s to the 1990’s led to an metre at day. Both Acartia species inhabited the increased dispersion of larvae in the latter pe- water column above the thermocline in spring riod. It was shown that this decreased the and summer. However, in November the overlap with larval prey and thus decreased the distribution was broader for all stages in both survival probability of larval cod. species. All stages of O. similis inhabited the Furthermore, the influence of variability in prey water layers around the halocline irrespective abundance and diversity on the results of an to the season, showing no vertical migration. individual based model on larval cod growth v Summary and survival was investigated. It was shown, factor was the size of prey organisms. Again, that the use of idealized mean prey fields led to the use of mean value led to an overestimation an overestimation of larval survival. Using field of survival. Using measurements from samples data derived from net sampling the survival was reduced the survival up to 75%. partly reduced to 50%. Another investigated vi Zusammenfassung In dieser Arbeit werden verschiedene Aspekte ter über die Wassersäule verteilten. Alle Stadien der Verteilung von Zooplankton und der daraus von O. similis hielten sich über das ganze Jahr resultierende Einfluss auf das Überleben von hinweg im Bereich der Salzgehaltssprungschicht Fischlarven gezeigt. auf, ohne Wanderungsbewegungen zu zeigen. Die Vertikalverteilung der wichtigsten calanoi- Die Untersuchung der Verteilung von sich re- den Copepoden im Bornholm Becken, Pseudo- produzierenden P. acuspes Weibchen mit Hilfe calanus acuspes, Temora longicornis, Acartia des Videoplankton Rekorders in zwei hydro- bifilosa, Acartia longiremis und dem einzigen graphisch unterschiedlichen Jahren zeigte eine cyclopoiden Copepoden Oithona similis, wurde enge Bindung an hohe Salzgehalte (13.25) und mit Hilfe von Multischließnetzen untersucht. die Vermeidung zu niedriger Sauerstoffgehalte Die Ergebnisse zeigen deutliche saisonale Un- (0.88 ml/l). Zusätzlich zur Darstellung der terschiede in der Vertikalverteilung aller Arten. Vertikalverteilung konnte mit Hilfe der Dabei konnte für P. acuspes eine ontoge- Aufnahmen eisacktragender Weibchen die netische Vertikalwanderung gezeigt werden, Anzahl der Eier pro Eisack bestimmt werden bei der sich die jungen Stadien in den oberen und über die temperaturabhängige Entwicklung Wasserschichten aufhalten und die Älteren die Eiproduktion. Obwohl die Produktion in sukzessive in tiefere Wasserschichten abwan- beiden Jahren gleich war, lag das Verhältnis von dern. Dabei halten sich die Adulten im Gebiet Nauplien zu Weibchen im Einstromjahr höher, der Salzgehaltssprungschicht auf. Gegen Winter was auf eine erhöhte Sterblichkeit in dem (November) löst sich diese Vertikalstruktur auf Stagnationsjahr schließen lässt. und alle Stadien verteilen sich über den gesam- Ein vereinfachtes individuen-basiertes Modell ten Wasserkörper. Allerdings dominieren zu für P. acuspes, gekoppelt mit einem Zirkulati- diesem Zeitpunkt ältere Copepodidstadien (C4, onsmodell der Ostsee, wurde benutzt, um den C5) die Stadienzusammensetzung. Für T. Einfluss zu untersuchen, den die Drift von die- longicornis konnte eine tägliche Vertikalwan- sen Copepoden auf eine Abnahme ihrer Bio- derung in den Sommermonaten (Juni 2001, Juli masse haben kann. Die Ergebnisse zeigen, dass und August 2002) gezeigt werden. Am ausge- in den 1990er Jahren, im Gegensatz zu den prägtesten war die Wanderung für die älteren 1980er Jahren, die vorherrschenden ba- Stadien, die von 20 Meter Wassertiefe in der rotrophen Strömungen zu einer starken Verdrif- Nacht bis auf 60 Meter am Tag wandern. Für tung aus dem Bornholmbecken in die östlichen April und November konnte keine Wanderung Becken und in die Küstenbereiche führten. Der gezeigt werden. Die Acartia Arten zeigten keine Rückstrom aus den östlichen Becken zurück in solche ausgedehnte Wanderungsbewegung, die Bornholmsee war hingegen zu keiner Zeit auch wenn sich der Verteilungsschwerpunkt bei signifikant. Die niedrigeren Salzgehalte und die A. longiremis von 10 Meter am Tag auf 20 höheren Temperaturen im küstennahen Wasser Meter in der Nacht verlagerte. Beide Arten könnten einen negativen Einfluss auf das halten sich im Frühjahr und den Som- Überleben oder die Reproduktion gehabt mermonaten vorwiegend über der Temperatur- haben. sprungschicht auf. Diese Bindung verschwand Dasselbe Zirkulationsmodell wurde benutzt, um im Winter (November), wo sich alle Arten brei- die Drift von Dorschlarven und die Konsequenz vii Zusammenfassung in Hinsicht auf Überlappung mit ihrer Nahrung Wachstum und Überleben von Dorschlarven. zu untersuchen. Auch hier konnte gezeigt wer- Es konnte gezeigt werden, dass die den, dass eine Veränderung der baroklinen Verwendung von idealisierten mittleren Nah- Strömungen in den 1990er Jahren im Gegen- rungsfeldern zu einer Überschätzung des Über- satz zu den 1980er Jahren, zu einer erhöhten lebens führen kann. Werden ‚echte’ Nahrungs- Dispersion der Larven führten. Es konnte ge- felder in dem Modell benutzt, lag das Überle- zeigt werden, dass dies zu einer Verringerung ben zum Teil um 50% niedriger. Als weiterer der Überlappung mit der Nahrung führte, wel- Faktor wurde die Größe der Nah- ches ein niedrigeres Überleben zur Folge hatte. rungsorganismen identifiziert. Auch hier zeigte Untersucht wurde weiterhin der Einfluss der sich, dass die Annahme einer mittleren Größe Variabilität in der Abundanz und Diversität der zu einer Überschätzung führt, und die Zooplankton-Nahrungsfelder auf die Ergebnisse Benutzung realer Größen das Überleben um eines individuen-basierten Modells zum bis zu 75% verringert. viii Content Summary ..............................................................................................................................................................v Zusammenfassung ...............................................................................................................................................vii
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