Larval Fish Dynamics in Coastal and Oceanic
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LARVAL FISH DYNAMICS IN COASTAL AND OCEANIC HABITATS IN THE CANARY CURRENT LARGE MARINE ECOSYSTEM (12 – 23°N) Dissertation with the aim of achieving a doctoral degree at the Faculty of Mathematics, Informatics and Natural Sciences Department of Biology of University Hamburg submitted by Maik Tiedemann M.Sc. Marine Biology B.Sc. Biology Hamburg, 2017 The present cumulative thesis is based on the scope on the bylaws of the Department of Biology's Doctoral Committee supplementing the Faculty of Mathematics, Informatics and Natural Sciences Doctoral Degree Regulations dated 1 December 2010 including adopted bylaws dated 23 February 2016. The content of chapter III may have changed in the process of publication and the submission of the present thesis. Please contact the principal author for citation purposes. Day of submission: 17. August 2017 Day of oral examination: 01. December 2017 The following evaluators recommend the admission of the dissertation: 1. Evaluator: Prof. Dr. Christian Möllmann Institute for Hydrobiology and Fisheries Science, Center for Earth System Research and Sustainability (CEN), Klima Campus, University of Hamburg, Grosse Elbstrasse 133, D-22767 Hamburg, Germany 2. Evaluator: Dr. Heino O. Fock Thünen-Institute (TI), Institute of Sea Fisheries, Federal Research Institute for Rural Areas, Forestry and Fisheries, Palmaille 9, 22767 Hamburg, Germany PREFACE The present dissertation cumulates the results of my doctoral project conducted from April 2013 to July 2017. My project was part of the tripartite German – French – African project “Ecosystem approach to the management of fisheries and the marine environment in West African waters” (AWA) funded by the German Federal Ministry of Education and Research (BMBF) under the grant number 01DG12073A. Additional funding was provided by the German Federal Office for Agriculture and Food (BLE), the French National Institute for Research for Sustainable Development (IRD), the French Research Institute for Exploitation of the Sea (IFREMER), and (especially for Chapter III) the UMR LEMAR 2012-2015 four-year plan (Team 3 on “Integrated study of ecosystem functioning and Western Africa cross-cutting orientation”). The project was designed to contribute to the understanding of oceanic processes that regulate larval fish distributions and assemblage patterns along the southern part of the Canary Current Upwelling Ecosystem. In the long-term, results are expected to support the development of an ecosystem approach to fisheries and the marine environment in West African waters. The results may further help to implement management measures (marine conservation and fisheries management) and may be used in biophysical models to increase the precision of hindcast and forecast scenarios of marine resources. My work provides insight into oceanic processes that are commonly observed in all world oceans and affect the life history of fish larvae. This dissertation comprises a general introduction to the topic, three peer-reviewed publications (Chapter I – III) and a synoptic discussion, including conclusions and an outlook. The overall concept of the doctoral work was developed by me in close collaboration with my supervisors, Dr. Heino Fock (Thünen Institute for Sea Fisheries) and Prof. Dr. Christian Möllmann (University of Hamburg). General introduction The concept was developed by me (MT). The general introduction was written by MT and includes a modified part of the proceedings: MAIK TIEDEMANN & FRANZISKA BILS (2016) Egg and larval fish studies from a bottom-up versus top down perspective. In: Bode M, Golz V, Jessen C & Lange J (2016) People and the 7 Seas – Interaction and Inspiration, YOUMARES proceedings Vol. 7. The chapter of the proceedings was written by MT and FB. Chapter I MAIK TIEDEMANN*, HEINO O. FOCK, PATRICE BREHMER, JULIAN DÖRING, CHRISTIAN MÖLLMANN Does upwelling intensity determine larval fish habitats in upwelling ecosystems? The case of Senegal and Mauritania The concept was developed by MT and HOF. The samples were collected by MT, HOF, PB, and JD during three expeditions at sea on RV Thalassa 2014 and FRV Walther Herwig III 2014/15. The sorting and identification of fish larvae were performed by MT. Data were analysed by MT with contributions from HOF and CM. The manuscript was written by MT with contributions from JD, PB, HOF, and CM. The paper is published in Fisheries Oceanography (2017) DOI: 10.1111/fog.12224. Chapter II MAIK TIEDEMANN, PATRICE BREHMER Larval fish assemblages across an upwelling front: Indication for active and passive retention The concept was developed by MT and PB. The samples were collected by BP during an expedition at sea on RV Antea 2013. The sorting and identification of fish larvae were performed by MT. Data were analysed by MT with contributions from PB. The manuscript was written by MT with contributions from PB. The paper is published in Estuarine, Coastal and Shelf Science (2017) DOI: 10.1016/j.ecss.2016.12.015. Chapter III MAIK TIEDEMANN, HEINO O. FOCK, JULIAN DÖRING, LUC BONAVENTURE BADJI, CHRISTIAN MÖLLMANN Oceanic dipole eddies drive cross-shelf dispersal of fish larvae The concept was developed by MT and HOF. The samples were collected by MT, HOF, and LBB during an expedition at sea on FRV Walther Herwig III 2015. The sorting and identification of fish larvae were performed by MT and LBB. Data were analysed by MT. The manuscript was written by MT with contributions from JD, HOF, LBB, and CM. The manuscript is under review. Synoptic discussion The concept was developed by MT. The synoptic discussion was written by MT. SUMMARY Understanding processes that regulate dispersal patterns and survival of larval fish in relation to habitat structures as bottom-up drivers in upwelling ecosystems is the objective of this dissertation. To what extent do larval fish assemblages reflect species- specific adaptations to the physical environment in marine ecosystems? That’s only one of several remaining questions that have received much attention in the past, yet remain unanswered for all globally important marine ecosystems. The abundant small pelagic fish species in the southern Canary Current Upwelling Ecosystem (CCUE) provide livelihoods for millions of people, but the physical processes that drive abundance and distribution of these species are not well known. In order to understand vital processes in the CCUE that control the dispersal of larval fish assemblages, four surveys at sea were conducted between 2014 and 2016. The data framework, which was used to answer central research questions, was based on a combination of field work (expeditions with larval fish sampling), in-situ measurements of environmental parameters (salinity, fluorescence, conductivity, temperature, depth – CTD), community analysis (e.g. Cluster Analyses, Discriminant Analyses, Similarity Percentage Analysis (SIMPER), Multivariate Analysis of Variance (MANOVA), Random Forest Models, and Generalized Additive Models (GAMs)), and satellite based remote sensing data (sea surface temperature, sea surface chlorophyll-a, wind regime, absolute dynamic topography and geostrophic current vectors). Results from data collection served as the basis for preparation of my dissertation and are presented in the form of three scientific publications (Chapter I – III). In Chapter I, a spatiotemporal niche-partitioning of larval habitats was detected for the two most dominant small pelagic species, round sardinella (Sardinella aurita) and European sardine (Sardina pilchardus), constrained by species- specific upwelling regimes. In Chapter II, active larval vertical migration behaviour was revealed in an offshore upwelling cell. Species that occurred in the inner upwelling cell took advantage of a passive retention to circumvent a loss of larvae into unproductive oceanic regions during an upwelling event. In Chapter III, two larval fish assemblages were observed, each inhabiting one of two water masses that were trapped by two mesoscale eddies along a front. Chapter I provides insight into large scale processes that mainly influence the southern CCUE during summer and winter, and summarizes general conditions of the physical environment. It contributes a baseline understanding of ecosystem function as a response to broad meteorological and oceanographic processes that affect the abundance of larvae of small pelagic fish species. Chapters II and III investigated on a mesoscale upwelling fronts, converging water masses, a dipole eddy and cross-shelf variations that regulate the dispersal and composition of larval fish assemblages at the two main nursery areas (Senegalese sub-region and Banc d’Arguin) of the southern CCUE. Within the scientific scope of Chapter I, the results identified larval fish habitats constrained by upwelling intensities in the southern CCUE. While the occurrence of S. pilchardus larvae peaked during strong upwelling in winter/spring, S. aurita larvae were most abundant during the decreased upwelling phase in summer. The shift of peak occurrences of both dominant small pelagic species indicated a species-specific adaptation to the main physical process (upwelling) in Eastern Boundary Upwelling Ecosystems. Distribution patterns of both species exhibited a spatiotemporal niche partitioning. Under future scenarios of climate warming, where upwelling intensity is expected to change, this result may explain shifting dominance structures within fish assemblages that occur in the southern CCUE. The investigation into the impacts of upwelling on larval fish