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Diel Vertical Migration Patterns of Baltic Sea Copepods Analysed With Diel Vertical Migration Patterns of Baltic Sea Copepods Analysed with a Video Plankton Recorder Dissertation zur Erlangung des Doktorgrades an der Fakultät für Mathematik, Informatik und Naturwissenschaften Fachbereich Biologie der Universität Hamburg vorgelegt von Kristin Hänselmann aus Hamburg Hamburg, 2019 Gutachter der Dissertation: Prof. Dr. Axel Temming Jun.-Prof. Dr. Elisa Schaum Tag der Disputation: 20.09.2019 Mitglieder der Prüfungskommission: Prof. Dr. Kathrin Dausmann - Vorsitz Prof. Dr. Inga Hense Prof. Dr. Axel Temming Jun.-Prof. Dr. Elisa Schaum CONTENTS SUMMARY _________________________________________________________________ 1 ZUSAMMENFASSUNG ________________________________________________________ 3 GENERAL INTRODUCTION ____________________________________________________ 5 THE BALTIC SEA ______________________________________________________________________ 5 COPEPODS IN THE BALTIC SEA __________________________________________________________ 7 COPEPOD DIEL VERTICAL MIGRATION ____________________________________________________ 9 PLANKTIVOROUS FISH IN THE BALTIC SEA ________________________________________________ 10 PLANKTON PATCHINESS ______________________________________________________________ 11 VIDEO PLANKTON RECORDER __________________________________________________________ 11 AIM OF THE THESIS __________________________________________________________________ 12 REFERENCES __________________________________________________________________________ 15 Chapter I _________________________________________________________________ 25 Predator density triggered vertical migration of Pseudocalanus acuspes in the Baltic Sea ____ 25 ABSTRACT ____________________________________________________________________________ 25 INTRODUCTION _______________________________________________________________________ 26 METHODS ____________________________________________________________________________ 29 SAMPLING AREA & STRATEGY _________________________________________________________ 29 VIDEO PLANKTON RECORDER __________________________________________________________ 29 PSEUDOCALANUS ACUSPES IMAGE DATA ________________________________________________ 31 CLUPEID FISH DATA __________________________________________________________________ 32 DATA ANALYSIS _____________________________________________________________________ 35 RESULTS _____________________________________________________________________________ 37 STAGNATION PERIODS _______________________________________________________________ 37 POST INFLOW SITUATION _____________________________________________________________ 41 DISCUSSION __________________________________________________________________________ 45 METHODS _________________________________________________________________________ 45 RESULTS ___________________________________________________________________________ 45 CLUPEID FISH DATA _______________________________________________________________ 46 DOWNWARD COPEPOD MIGRATION _________________________________________________ 47 UPWARD COPEPOD MIGRATION _____________________________________________________ 49 TOP-DOWN / BOTTOM-UP CONTROL _________________________________________________ 50 CONCLUSION _________________________________________________________________________ 51 REFERENCES __________________________________________________________________________ 52 i Chapter II _________________________________________________________________ 61 DVM patterns of thermocline associated copepods in different basins of the Baltic Sea _____ 61 ABSTRACT ____________________________________________________________________________ 61 INTRODUCTION _______________________________________________________________________ 62 METHODS ____________________________________________________________________________ 65 SAMPLING AREA & STRATEGY _________________________________________________________ 65 VIDEO PLANKTON RECORDER __________________________________________________________ 65 COPEPOD DATA _____________________________________________________________________ 67 MULTINET COPEPOD DATA ___________________________________________________________ 67 CLUPEID FISH DATA __________________________________________________________________ 67 DATA ANALYSIS _____________________________________________________________________ 68 RESULTS _____________________________________________________________________________ 70 MULTINET COPEPOD DATA ___________________________________________________________ 70 COPEPOD DVM PATTERNS ____________________________________________________________ 70 COPEPOD DVM PATTERNS—INDIVIDUAL SPECIES _________________________________________ 73 CLUPEID DVM PATTERNS _____________________________________________________________ 75 OVERLAP COPEPOD & CLUPEID DVM ____________________________________________________ 76 COPEPOD & AURELIA AURITA DISTRIBUTION _____________________________________________ 81 DISCUSSION __________________________________________________________________________ 83 MULTINET COPEPOD DATA ___________________________________________________________ 83 COPEPOD DVM PATTERNS ____________________________________________________________ 83 OVERLAP COPEPOD & CLUPEID DVM ____________________________________________________ 84 COPEPOD & AURELIA AURITA DISTRIBUTION _____________________________________________ 85 OTHER CAUSES FOR DVM _____________________________________________________________ 86 CONCLUSION _________________________________________________________________________ 87 REFERENCES __________________________________________________________________________ 88 Chapter III ________________________________________________________________ 95 Investigating copepod patches and their implications on sprat (Sprattus sprattus) feeding in the Baltic Sea _____________________________________________________________________ 95 ABSTRACT ____________________________________________________________________________ 95 INTRODUCTION _______________________________________________________________________ 96 METHODS ____________________________________________________________________________ 99 SAMPLING AREA & STRATEGY _________________________________________________________ 99 VIDEO PLANKTON RECORDER __________________________________________________________ 99 VPR DATA ________________________________________________________________________ 101 COPEPOD PATCH DEFINITION _________________________________________________________ 102 VERTICAL INTERPOLATION OF VPR DATA _______________________________________________ 102 ii PREDICTED STOMACH CONTENT WEIGHTS ______________________________________________ 103 SPRAT STOMACH FIELD SAMPLES _____________________________________________________ 108 MULTINET COPEPOD DATA __________________________________________________________ 108 DATA ANALYSIS ____________________________________________________________________ 109 RESULTS ____________________________________________________________________________ 110 VALIDATING VERTICAL INTERPOLATION ________________________________________________ 110 MEASURING COPEPOD PATCHES ______________________________________________________ 110 PREDICTING STOMACH CONTENT WEIGHTS _____________________________________________ 116 DISCUSSION _________________________________________________________________________ 121 METHODS ________________________________________________________________________ 121 RESULTS __________________________________________________________________________ 121 MEASURING COPEPOD PATCHES ___________________________________________________ 121 PREDICTING STOMACH CONTENT WEIGHTS ___________________________________________ 122 CONCLUSION ________________________________________________________________________ 125 REFERENCES _________________________________________________________________________ 126 GENERAL DISCUSSION _____________________________________________________ 131 EFFICIENCY OF PLANKTON SAMPLING WITH THE VPR SYSTEM ______________________________ 131 PREDATOR AVOIDANCE IN PSEUDOCALANUS ACUSPES DVM _______________________________ 133 SPECIES-SPECIFIC DVM DRIVERS ______________________________________________________ 133 COPEPOD DVM, PATCHINESS & IMPLICATIONS ON SPRAT FEEDING __________________________ 136 GENERAL CONCLUSION & OUTLOOK ___________________________________________________ 138 REFERENCES _________________________________________________________________________ 140 Individual contributions to chapters with multiple authors ________________________ 145 Danksagung _____________________________________________________________ 147 Eidesstattliche Versicherung ________________________________________________ 149 Certification of Written English Quality ________________________________________ 151 iii SUMMARY In this thesis, interspecies differences of Baltic Sea copepod diel vertical migration (DVM) are investigated with a special emphasis on drivers of these migration behaviours. Additionally, for the first time information on the spatial dimensions of Baltic Sea copepod patches are provided. Furthermore, implications of a patchy copepod distribution on feeding of clupeid fish like sprat (Sprattus sprattus) are elucidated. The investigations on copepod DVM and patch dimensions are based on analyses of Video Plankton Recorder (VPR) data. This unobtrusive underwater microscope system provides small- as well as large-scale information on plankton composition and hydrographic conditions. The unobtrusive nature of sampling makes the VPR the
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