1. Individual based modeling of growth and survival of Atlantic Cod (Gadus morhua) and Lesser Sandeel (Ammodytes marinus) larval stages Zeren Gürkan, Asbjørn Christensen, Henrik Mosegaard DTU Aqua, Charlottenlund Castle, Denmark 7/13/2015Summer course 2009 1

2. My background BSc in Biology, METU, Turkey MScE in Env. Engr., DTU, Denmark Grad. study in Env. Engr., METU & PhD for 1 year in Bio.&Agr. Engr., UGA, USA 7/13/2015Summer course 2009 2

3. Contents of presentation SUNFISH project summary PhD research contents Results from literature review Prey field for fish models – Feeding conditions Existing bioenergetic models Initial bioenergetic Individual-Based Model (IBM) formulations 7/13/2015Summer course 2009 3

4. My PhD research objectives They are embedded in and address several tasks in SUNFISH Main objectives are identifying and modeling key biological processes that affect growth and survival of larval and juvenile cod and sandeel by giving emphasis to responses to local physical and feeding conditions Existing bioenergetic individual-based models (IBMs) will be developed and coupled to biogeochemical models of the North Sea together with other SUNFISH partners The coupled models will be used to simulate responses of biological processes of drift, growth and survival to climate change effects as conditions of physical and biogeochemical spatio-temporal fields, and formulate future sustainable ecosystem management strategies 7/13/2015Summer course 2009 4

5. SUNFISH - “Sustainable fisheries, climate change and the North Sea ecosystem” Global climate change will challenge fisheries and ecosystem management in the seas Primary and secondary production and distribution, feeding, growth, and survival of fish will be affected by changes in temperature, wind conditions, river runoff, currents It will be more difficult to separate effects of fishing from those of environmental changes, to identify biological reference points, and to develop management strategies for sustainable fisheries Well-founded assessment of the population dynamics of the North Sea fish stocks is necessary 7/13/2015Summer course 2009 5

6. SUNFISH is a collaborative project of several Danish and international institutes It aims to improve the scientific basis and predictions of effects of climate change on the North Sea ecosystem and the sustainable fisheries management in the North Sea It will provide an integrated modeling framework for developing optimal and sustainable fisheries management strategies, based on description of mechanisms of processes instead of extrapolations of observed trends, to predict the outcomes of climate change on the North Sea ecosystem It focuses on cod and sandeel, which are key species in the North Sea ecosystem and economically important to the Danish fishery 7/13/2015Summer course 2009 6

7. SUNFISH tasks from work packages I contribute to : Formulation of a model describing food intake of larvae based on literature & growth of cod larvae IBM model of larval cod growth based on biological process models of growth IBM model of larval cod survival  Predictions of the spatio-temporal distribution of larvae based on IBM models  Comparison of predicted larval distribution and size with survey observations  Identification of processes limiting survival of early life stages  Comparison of predicted distribution of settled cod with distribution observed in surveys  Spatio-temporal analysis of the predicted probability of survival to settling of eggs spawned in different areas  Estimation of sandeel growth in relation to biophysical conditions 7/13/2015Summer course 2009 7

8. Methods include the use of predictions of conditions in the North Sea in past and future accounting for climate change scenarios by BSHcmod and HYCOM hydrodynamic models, which are coupled to 3-D biogeochemical model, ERGOM, as basis of analyses of spatio-temporal variation in growth and survival of the fish early life stages Survival will be modeled using Letcher type IBM and hydrographic approach. Estimates of energy intake and expenditure will produce model of growth. Effects of local physical conditions and behavioral aspects will be included 7/13/2015Summer course 2009 8

9. 7/13/2015Summer course 2009 9 Marine Ecological Modeling Center workshop

10. 7/13/2015Summer course 2009 10

11. Initial bioenergetic IBM formulations Sandeel Population Analysis Model (SPAM) assumes von Bertalanffy temperature modulated growth model (Christensen) I develop this to a detailed model based on Letcher et al. (1996) formulation Energy intake and expenditure are combined Function of temperature, food abundance and quality Includes explicit descriptions of physiological processes and behavioral aspects 7/13/2015Summer course 2009 11

12. SPAM (SLAM - Christensen et al., 2008) 7/13/2015 Summer course 2009 12 Growth

13. 7/13/2015 Summer course 2009 13 SPAM (SLAM - Christensen et al., 2008)

14. 7/13/2015 Summer course 2009 14 SPAM (SLAM - Christensen et al., 2008) Suitable sandeel banks Interbank exchanges of sandeel larvae Av. Spatial SD (bank variability) Sandeel Year

15. Flow diagram - All functions are dependent on larval size. 5 submodels determine prey encounter, foraging, bioenergetics-growth, starvation, and predation rates. Major processes affecting each submodel are in the pointed boxes. Characteristics of the larvae’s environment (food and predators) are in the boxes with horizontal arrows. The model calculates growth rates for each larva every time step. Each simulation may start with a certain number of larvae all with a specified length. 7/13/2015Summer course 2009 15 Generic larval IBM - Letcher et al. (1996)

16. 7/13/2015Summer course 2009 16 (Assimilation efficiency) (Total costs) Specific dynamic action, SDA & egestion, E Growth submodel (Routine metabolism) Weight, W Ingestion, I

17. 7/13/2015Summer course 2009 17 Growth submodel to examine effects of food patchiness and temporal variation Constant ingestion, I

18. 7/13/2015Summer course 2009 18 Maximum consumption, total mass eaten, and masses of four prey types eaten under nominal conditions and optimal foraging model Foraging submodel : Applies optimal foraging model (Charnov, 1976)

19. 7/13/2015Summer course 2009 19 Foraging submodel varying ingestion 2 preys

20. Hypothesis - Bottom-up cascade. Recent statement by Beaugrand et al. (2003) says recruitment failure of many species in the North Sea is linked to the change in distribution and composition of warm (Calanus helgolandicus) and cold (C. finmarchicus) water zooplankton, which is potentially driven by regional warming. It will be analyzed further with the bioenergetic IBMs developed using zooplankton model outputs from SUNFISH partners and ‘Continuous Plankton Recorder’ (CPR) time series data. 7/13/2015Summer course 2009 20

21. 7/13/2015Summer course 2009 21 Sir Alister Hardy Foundation for Ocean Science (SAHFOS) provided the CPR data for NS area of several taxa of plankton included in diets of cod and sandeel One use of this database will be to realize the prey conditions as input to the model Calibration & validation of model results

22. Thank you. Questions & feedback? 7/13/2015Summer course 2009 22

23. PhD project contents - Hypotheses Climate change effects. Recent progress by SUNFISH partners allows improved description of water turbidity, turbulence, and light penetration in relation to meteorological forcing factors. These three factors strongly affect feeding success and efficiency as well as predation risk. Future climate scenarios anticipate stronger winds on average and therefore, it is highly relevant to explore the biological impacts on early life stages using the developed bioenergetic IBMs. 7/13/2015PhD Seminar 2009 23

24. PhD project contents - Hypotheses Optimal life history analysis. The ecological fitness of a species depends on the traits characterizing its entire life history. Traits in early life stages and of individuals including behavioral aspects may matter in determining the ecological fitness of a species. Evolutionary mechanisms have optimized these traits for each species in its ecosystem niche. The fitness of alternative life histories may shift relative to each other due to direct and indirect climate change effects, which may mean a regime shift in the ecosystem. The project can explore these aspects for cod and sandeel early life stages. Optimal life history analysis also constitutes a useful consistency check of the developed bioenergetic IBMs. 7/13/2015PhD Seminar 2009 24

25. PhD project contents - Hypotheses The physical and biogeochemical fields with sufficient spatial resolution will be available within the SUNFISH project to explore biological consequences. The processes on which climatic effects are quantified include drift, growth and survival of zooplankton and fish. The bioenergetic IBMs developed will be coupled to operational biogeochemical models for North Sea with other members of the SUNFISH project simulating climatic effects on total biomass and on spatio-temporal distribution, composition and nutritional value of zooplankton. 7/13/2015PhD Seminar 2009 25

26. PhD project contents - Hypotheses The distribution and composition of warm (C. helgolandicus) and cold (C. finmarchicus) water zooplankton has a distinct spatial and seasonal pattern. How the spatial and temporal heterogeneity and composition of the zooplankton community, which may include species with different nutritional values, affect recruitment predictions of cod and sandeel will be examined. Operational biogeochemical models most often assume a constant or otherwise simplified grazing rate from planktivorous biomass while elementary estimates suggest the grazing rate is highly variable over the seasonal cycle. Possibility of improving the performance of current operational biogeochemical models by implementing a feedback loop of fish grazing on zooplankton to demonstrate the importance of grazing dominated either by cod or sandeel larvae at a locality will be examined. 7/13/2015PhD Seminar 2009 26