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Population Dynamics and Trophic Interactions of Atlantic Cod Along the Greenland Shelf

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Population Dynamics and Trophic Interactions of Atlantic Cod Along the Greenland Shelf Population dynamics and trophic interactions of Atlantic cod along the Greenland shelf Dissertation with the aim of achieving a doctoral degree at: Faculty of Mathematics, Informatics and Natural Sciences Department of Biology, University of Hamburg Institute for Marine Ecosystems and Fisheries Science submitted by Karl-Michael Werner M.Sc. Fisheries biology and management B.Sc. Environmental Science Bremerhaven, 2020 Commission: Prof. Christian Möllmann (supervisor and evaluator), University of Hamburg Dr. Heino O. Fock (supervisor and evaluator), Thünen Institute of Sea Fisheries Bremerhaven Prof. Myron Peck, University of Hamburg Prof. Kathrin Dausmann, University of Hamburg Die mündliche Prüfung fand am 10. Juni 2020 statt. The oral examination took place on 10th of June 2020. Preface The results of this cumulative PhD dissertation summarize the findings of the research I have conducted as part of my PhD project from Mai 2016 to December 2019. This work was funded by the CLIMA project, reference RER 15/0008, Ministry of Foreign Affairs Norway. Biological samples were collected on nine research cruises with FRV Walther Herwig 3 between 2006 and 2017. I participated in the survey in 2016 as staff scientist and in 2017 as cruise leader. The work was focussed on investigating the impacts of natural variability and anthropogenic pressures on stock productivity and body condition of Atlantic cod (Gadus morhua) in Greenland waters. Because climate change and fishing have already affected all cod stocks in the North Atlantic and changes in abundance of cod affect ecosystems and socio-economic stability of coastal communities, it is important to disentangle the role of natural variability and fishing on population replenishment. This thesis consists of an introduction (Chapter 1), six publications presented below (Chapter 2-7) and a synoptic discussion (Chapter 8). The concept of this PhD dissertation was designed by myself with support of my advisors Heino O. Fock and Christian Möllmann. Chapter 2 Karl Michael Werner, Hans-Joachim Rätz, Søren Anker Pedersen, Christian Möllmann Population and abundance dynamics in the offshore stock of Atlantic cod in Greenland waters The concept was mainly developed by myself with support from Hans-Joachim Rätz and Christian Möllmann. Data were taken from the Thünen Institute data base. The literature review was conducted by mself. Hans-Joachim Rätz and myself extracted and prepared the data from our database. Hans-Joachim Rätz ran the virtual population analysis. The first draft of the paper was written by myself and all co-authors contributed with comments and advice. The manuscript is currently in preparation for submission. i Chapter 3 Camilla Sguotti, Saskia A. Otto, Xochitl Cormon, Karl Michael Werner, Ethan Deyle, George Sugihara, Christian Möllmann Non-linearity in stock-recruitment relationships of Atlantic cod: insights from a multi- model approach The concept was mainly developed by Camilla Sguotti and Christian Möllmann. I contributed data from the stock assessment in chapter 2. The first draft was written by Camilla Sguotti and all authors contributed with comments and advice afterwards. The manuscript was published in the ICES Journal of Marine Science (2019), doi:10.1093/icesjms/fsz113. Chapter 4 Fabian Zimmermann* and Karl Michael Werner* *both authors contributed equally to this study Improved management is the main driver behind recovery of Northeast Atlantic fish stocks Both authors developed the concept. Data analysis was mainly conducted by Fabian Zimmermann, writing of the first draft mainly by myself. The paper is published in Frontiers in Ecology and the Environment (2019), 17(2): 93-99 Chapter 5 Karl Michael Werner, Marc. H. Taylor, Rabea Diekmann, Josep Lloret, Christian Möllmann, Raul Primicerio, Heino O. Fock Evidence for limited adaptive responsiveness to large-scale spatial variation of habitat quality The concept was developed by myself and Heino O. Fock. And data were collected by myself and Heino O. Fock. Samples were taken and stomachs dissected by myself, Heino O. Fock and five master students. Samples were collected on Thünen surveys in Greenland between 2006 and 2016. Data were analysed by myself, Marc H. Taylor and Rabea Diekmann. I wrote the first draft of the manuscript and all co-authors contributed with comments and advice. The paper is published in Marine Ecology Progress Series (2019), 629: 179-191. ii Chapter 6 Ina Stoltenberg, Karl-Michael Werner, Stefanie M.H. Ismar-Rebitz, Heino Fock Trophic niche variability influences body condition on organosomatic and biochemical level of mature female Atlantic cod (Gadus morhua) in offshore Greenland waters The concept was developed by Ina Stoltenberg and myself as part of supervising Ina Stoltenberg’s Master thesis. Heino O. Fock, Stefanie Ismar-Rebitz and myself supervised Ina. Data were collected on the Thünen Greenland survey 2017, where I was cruise leader, by Ina Stoltenberg and myself. Ina Stoltenberg and myself wrote the first draft of the manuscript and Heino O. Fock and Stefanie M. H. Ismar-Rebitz contributed with comments and advice. The paper is in preparation for publication. Chapter 7 Robert Boenish, Jacob P. Kritzer, Kristin Kleisner, Robert Steneck, Karl Michael Werner, Wenbin Zhu, Frederick Schram, Douglas Rader, William Cheung, Jose Ingles, Yongjun Tian, John Mimikakis The global rise of crustacean fisheries Robert Boenish compiled the data and conducted the analysis. Robert Boenish and myself developed the current draft of the manuscript and all co-authors contributed with comments and advice. The paper is currently under revision in Frontiers in Ecology in the Environment (major revisions, 22.01.2020, FEE19-0294). iii Table of contents Preface i Thesis summary v 1 Introduction 1 2 Publication 1 17 Werner et al. (in prep) Population and abundance dynamics in the offshore stock of Atlantic cod in Greenland waters 3 Publication 2 37 Sguotti et al. (2019) Non-linearity in stock-recruitment relationships of Atlantic cod: insights from a multi-model approach 4 Publication 3 72 Zimmermann & Werner (2019) Improved management is the main driver behind recovery of Northeast Atlantic fish stocks 5 Publication 4 103 Werner et al. (2019) Evidence for limited adaptive responsiveness to large-scale spatial variation of habitat quality 6 Publication 5 133 Stoltenberg et al. (in prep.) Trophic niche variability influences body condition on organosomatic and biochemical level of mature female Atlantic cod (Gadus morhua) in offshore Greenland waters 7 Publication 6 162 Boenish et al. (in rev.) The global rise of crustacean fisheries 8 Synoptic discussion and conclusions 183 Acknowledgements 205 iv Thesis Summary Atlantic cod (Gadus morhua) is the most abundant high-trophic-level fish predator in continental shelf habitats in the North Atlantic and has been a vital economic and nutritious resource for many nations. Climate change and overexploitation have already affected all stocks of Atlantic cod and changes in their abundance influence entire ecosystems and the socioeconomic stability of fishing communities. The short, medium and long-term future of marine ecosystem compositions and global fishery patterns will depend on global warming and how marine resources are managed. Investigating how wild fish stocks respond to environmental change and anthropogenic pressures and how they can be sustainably managed in a changing environment is one of the most important questions to be investigated in marine ecology and fishery science. Although Atlantic cod is one of the most important fishery resources in Greenland, scientific knowledge on the stock is patchy ever since the fishery collapsed. Here, the cod stock belonged to the once largest cod stocks worldwide with a total stock biomass of ~ 4 million tons. The stock supported a fishery with annual landings up to 460.000 tons until it collapsed in the 1970s. It is the objective of this thesis to investigate natural and anthropogenic drivers impacting the life history and abundance of cod in Greenland waters. In particular, I focus on fishing as anthropogenic driver and on the influence of habitat heterogeneity as reflected through diet composition as natural driver. The thesis consists of six scientific publications, which deal with 1) fishing and environmental change as drivers of abundance of the cod stock in Greenland waters since 1955; 2) non-linear stock-recruitment dynamics of North Atlantic cod stocks and their implications for recovery potential; 3) the impacts of natural variation and fishing as drivers of Northeast Atlantic fish stocks; 4) the influence of diet composition on energy reserves and the ability of Greenland cod to adapt to spatial and temporal variability of habitat quality and how this influences their body condition; 5) if different trophic niches impact the reproductive capacities and body condition of the cod population in East Greenland and 6) how anthropogenically altered v marine ecosystems including an example from West Greenland influence the composition of fishery harvests and if these changes come with ecological and socioeconomic trade-offs. Results of our review in publication 1 show that while the collapse of the Greenland cod stock in the 1970s was mainly driven by recruitment failure associated with dropping temperatures, overfishing in the late 1980s and early 1990s induced hysteresis effects, which have ever since the collapse in the 1970s prevented a recovery. The stock can benefit from irregular influx events of eggs
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