General Enquiries on the form should be made to: Defra, Procurements and Commercial Function (Evidence Procurement Team) E-mail: [email protected]

Evidence Project Final Report

 Note In line with the Freedom of Information Project identification Act 2000, Defra aims to place the results of its completed research projects in the public domain wherever possible. 1. Defra Project code M1102 The Evidence Project Final Report is designed to capture the information on 2. Project title the results and outputs of Defra-funded MEMFISH - Macro-ecology of marine finfish in UK waters research in a format that is easily publishable through the Defra website An Evidence Project Final Report must be completed for all projects. 3. Contractor Cefas This form is in Word format and the organisation(s) boxes may be expanded, as appropriate. Pakefield Road Lowestoft  ACCESS TO INFORMATION Suffolk The information collected on this form will NR33 0HT be stored electronically and may be sent to any part of Defra, or to individual

researchers or organisations outside 54. Total Defra project costs £ 1,104,490 Defra for the purposes of reviewing the (agreed fixed price) project. Defra may also disclose the

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EVID4 Evidence Project Final Report (Rev. 06/11) Page 1 of 28 6. It is Defra‘s intention to publish this form. Please confirm your agreement to do so...... YES NO (a) When preparing Evidence Project Final Reports contractors should bear in mind that Defra intends that they be made public. They should be written in a clear and concise manner and represent a full account of the research project which someone not closely associated with the project can follow. Defra recognises that in a small minority of cases there may be information, such as intellectual property or commercially confidential data, used in or generated by the research project, which should not be disclosed. In these cases, such information should be detailed in a separate annex (not to be published) so that the Evidence Project Final Report can be placed in the public domain. Where it is impossible to complete the Final Report without including references to any sensitive or confidential data, the information should be included and section (b) completed. NB: only in exceptional circumstances will Defra expect contractors to give a "No" answer. In all cases, reasons for withholding information must be fully in line with exemptions under the Environmental Information Regulations or the Freedom of Information Act 2000. (b) If you have answered NO, please explain why the Final report should not be released into public domain

Executive Summary 7. The executive summary must not exceed 2 sides in total of A4 and should be understandable to the intelligent non-scientist. It should cover the main objectives, methods and findings of the research, together with any other significant events and options for new work.

EVID4 Evidence Project Final Report (Rev. 06/11) Page 2 of 28 MEMFISH - Macro-ecology of marine finfish in UK waters

In contrast to previous projects, MEMFISH aimed to take a whole life-cycle approach to understand the biology and ecology of marine fish in UK waters. Specifically, MEMFISH focussed on: the relationships between fish population sub-units in spawning areas, on nursery grounds and on feeding grounds; how these vary year-on-year; and the contribution of the environment to such variation. To achieve this, MEMFISH work proceeded in three distinct phases: (1) collation and evaluation; (2) implementation; and (3) integration. This provided a unique opportunity to collate, assess and evaluate existing datasets, and identify how these could best then be deployed, both in terms of re-analysis and with supplementary research, to answer policy-relevant questions and ultimately, to reduce uncertainty in management advice relevant to future management needs.

“Macro-ecology of marine fish in UK waters” (MEMFISH)

Project Plan: Scientific Outputs: Case Studies:

Phase 1: COLLATION & Apostolaki et al. 2008 EVALUATION Metcalfe et al. 2008a Metcalfe et al. 2008b Sims et al. 2008 Sturrock et al. 2012a Year 1 Flatfish Connectivity Meeting Use existing frameworks/data Cod Mini-Symposium

Field Field Survey

Individual behaviour to population dynamics to population Individual behaviour Hunter et al. 2009a; 2012a Albaina et al. 2010 Phase 2: IMPLEMENTATION Pliru et al. 2012 Fox et al. 2012 Sturrock et al 2012b,c,d ICES Benchmark workshop on flatfish, 2009 Egg WKPESTO 2012

Natural Tags Righton et al. 2008; 2009;2010;2012 Kjesbu et al. 2010 Bendall et al. 2012a Heath et al. 2012 Larva Neuenfeldt et al. 2012 Righton & Metcalfe 2012

West et al. 2012 cycle approach cycle

- NERC-SMB Project

ICES Benchmark workshop on roundfish, 2009 Artificial Tags

ICES WKROUND, 2012 Hatchingcatching to

Juvenile Hunter & Kupschus 2007 Hunter et al. 2012b

Wholelife IFREMER Visiting Scientist 2009/10 Years 2-4 ICES WKFLAT 2012 Develop new case studies Adult Quayle et al. 2009

Hunter & de Pontual 2011 Collaboration ICES WKROUND 2012 Phase 3: INTEGRATION

Hunter et al. 2009b Bendall et al. 2009 Bendall et al. 2012b Wiegand et al. 2012 Year 5 ICES benchmarking of North Sea plaice and cod stocks, and Irish Sea Biology into FM Cod and bass stocks

Figure 1 illustrates how the scientific objectives originally proposed relate to the 3 research phases, including the four new WPs. As far as was possible, a whole life-cycle approach was applied, using both standard and novel techniques. The phased and modular structure of the MEMFISH project proved a highly effective means of making best use of, and publishing existing data, and in generating new work in terms of new collaborations and proposals (with 16 peer- reviewed publications, 3 book chapters, 5 ICES papers and 5 more manuscripts in preparation by close of project).

MEMFISH work has contributed directly to ICES benchmarking exercises and working groups, and has been disseminated widely. All of the work carried out under MEMFISH has been consistent with Defra‘s evidence investment strategy, and the Marine Strategy Framework Directive (MSFD), specifically action 3, that ―Populations of all commercially exploited fish and shellfish are within safe biological limits, exhibiting a population age and size distribution that is indicative of a healthy stock‖.

Phase 1 – Collation & Evaluation

Through a process of consultation with Defra, stakeholder groups and Cefas‘ own fisheries management advisors, four new research work-packages (WPs) were agreed.

Phase 2- Implementation

The projects proposed as a result of phase 1 were put forward to develop analytical and modelling processes that would add value to the project by combining MEMFISH data with data gathered over many

EVID4 Evidence Project Final Report (Rev. 06/11) Page 3 of 28 years, and from additional satellite projects. The focus of new work was on the relationships between fish population subunits in spawning areas, on nursery grounds, and on feeding areas, how these vary year- on-year, and the contribution of the environment to such variation:

WP1 Plaice population dynamics and extension of the North Sea Population Movement Simulator (NSPMS).

Otolith microchemistry Population Dynamics - Identification of plaice egg and larval predators using molecular probes - Impact of sprat predation on plaice egg mortality - Large-scale variation in seasonal swimming patterns Implementation of the NSPMS model.

WP2 Population structure and connectivity of cod in ICES area VII.

Field Survey of biological attributes of cod spawning aggregations in the Irish Sea 2009/11 - Overall catch composition in the western and eastern Irish Sea - Distribution and abundance of cod - Size composition of cod on spawning grounds - Sex ratio of cod - Maturity and reproductive characteristics of cod - Length–weight relationships Stock identification - Spatial dynamics of cod in the North Sea - Spatial dynamics of cod in the Irish & Celtic Seas

WP3 Population structure and connectivity of sole in UK Waters.

Exploration of best-practise for attachment of electronic data storage tags (DSTs) Potential for water chemistry to predict otolith composition in juvenile sole. Co-development of “Understanding and predicting connectivity in marine fish populations”

WP4 Distribution and abundance of sea-bass in UK waters in relation to climate change.

Bass Distribution Bass tagging (English Channel/Western Waters)

Data from the new WPs was interpreted in relation to the marine environment in the broadest sense (i.e. including human activities, local and global changes in habitats). Regional case studies have targeted issues of match or mismatch between metapopulation dynamics and management scales, conservation of essential fish habitats and the evaluation of climate change on habitat and biology. The strength of the multidisciplinary approach applied by MEMFISH is that the combined data streams provide a more comprehensive understanding of life-history characteristics than any one technique in isolation might have achieved.

Phase 3 – Integration

MEMFISH generated data and case studies were linked to the development and evaluation of management and recovery plans, and simulation modelling frameworks were used to examine the implications of the new knowledge collected during the project both to evaluate the importance of incorporating biological knowledge and how to use it to provide advice on the `safe biological limits to exploitation‘, as follows: 1. Applying individual-based telemetry data in fisheries management: A biologically-based movement model for plaice in the North Sea. 2. Using DST data to assess changes in the accessibility of Atlantic cod to trawl gears 3. Spatio-temporal dynamics of Atlantic cod in western Waters 4. Are spatial closures better than size limits for halting the decline of the North Sea thornback ray?

EVID4 Evidence Project Final Report (Rev. 06/11) Page 4 of 28

Project Report to Defra 8. As a guide this report should be no longer than 20 sides of A4. This report is to provide Defra with details of the outputs of the research project for internal purposes; to meet the terms of the contract; and to allow Defra to publish details of the outputs to meet Environmental Information Regulation or Freedom of Information obligations. This short report to Defra does not preclude contractors from also seeking to publish a full, formal scientific report/paper in an appropriate scientific or other journal/publication. Indeed, Defra actively encourages such publications as part of the contract terms. The report to Defra should include:  the objectives as set out in the contract;  the extent to which the objectives set out in the contract have been met;  details of methods used and the results obtained, including statistical analysis (if appropriate);  a discussion of the results and their reliability;  the main implications of the findings;  possible future work; and  any action resulting from the research (e.g. IP, Knowledge Exchange).

EVID4 Evidence Project Final Report (Rev. 06/11) Page 5 of 28 Contents:

1. General Introduction. 2. Summary of research results in relation to Scientific Objectives. 3. Collation and evaluation of existing data. 4. Plaice population dynamics and extension of the North Sea Population Movement Simulator (NSPMS). 5. Population structure and connectivity of cod in ICES area VII. 6. Population structure and connectivity of sole in UK Waters. 7. Distribution and abundance of sea-bass in UK waters in relation to climate change. 8. Integration. 9. Future Work

1. GENERAL INTRODUCTION

The waters around the UK harbour a diverse range of marine habitats. For fish inhabiting this environment, physical characteristics including depth, water currents, topography, sediment type, temperature, salinity, oxygen, illumination vary on a spatial and seasonal basis, as do levels of food availability and predator abundance. Consequently, fish within their broad geographic range are rarely distributed evenly or randomly. Instead, fish populations typically exhibit patchy distributions, with higher abundance in preferred habitats and lower abundances (or even total absence) elsewhere. Variability in lifetime environmental experience is further complicated for those species that show developmental (e.g. between nursery areas and adult populations) and/or seasonal (e.g. between spawning and feeding) shifts in habitat occupancy. Spatial and temporal variation in distribution and population fragmentation can result in varying responses to exploitation and climate change. In order to predict the likely responses of fish populations or sub-populations to changes in patterns of exploitation, or different management or conservation actions, it is therefore essential to have a fundamental understanding of fish population structures, and the biological and environmental processes that give rise to them.

However, knowledge of the whole life-cycle of most marine fin-fish species is at best fragmentary, focussing on specific phases of the life-cycle at different times of year and in different geographical areas. Even for the more intensively studied species, there are considerable gaps in existing knowledge. For example, the locations and long-term persistence of cod spawning grounds in the North Sea (Blanchard et al., 2005) or the connectivity between populations of sole in the western English Channel (ICES 2005) are still incompletely understood. Such gaps in knowledge are an impediment to the development of robust, multi-annual technical measures to conserve and rebuild stocks. In consequence, the population dynamics used in assessment (i.e. VPA) are estimated from commercial catch data (Fromentin et al., 2000). To reduce the uncertainty associated with stock assessment there is need for novel, fishery-independent, observations. Furthermore, without a full understanding of the ontogenetic and seasonal patterns in the distribution, movement and behaviour of individuals, together with knowledge of how individuals of the same species interact across their geographic range, the significance of knowledge deficits is not always immediately apparent. This in turn constrains the ability of fisheries scientists to identify and quantify the likely impact of possible conservation or management actions (Hunter et al., 2006; Kell et al., 2004).

In contrast to previous projects, therefore, MEMFISH aimed to take a whole life-cycle approach to understand the biology and ecology of marine fish (figure 1). Specifically, MEMFISH focussed on the relationships between fish population sub-units in spawning areas, on nursery grounds and on feeding grounds, how these vary year- on-year, and the contribution of the environment to such variation. A holistic view of life-history biology requires fundamental understanding of the dispersal, growth and survival of the early life stages, and of the movements, growth, survival, maturation and reproduction of the juvenile and adult stages at both local and regional scales. To achieve this, MEMFISH work proceeded in three distinct phases: (1) collation and evaluation; (2) implementation; and (3) integration (figure 1). This provided a unique opportunity to collate, assess and evaluate existing datasets, and identify how these could best then be deployed, both in terms of re-analysis and with supplementary research, to answer policy-relevant questions and ultimately, to reduce uncertainty in management advice relevant to future management needs.

Through a process of consultation with Defra, stakeholder groups and Cefas‘ own fisheries management advisors, four new research work-packages (WPs) were agreed. These focussed on plaice population dynamics in the North Sea, population structure and connectivity of cod in ICES Area VII, population structure and connectivity in sole in UK waters, and the distribution and abundance of sea bass in UK waters in relation to climate change.

EVID4 Evidence Project Final Report (Rev. 06/11) Page 6 of 28 “Macro-ecology of marine fish in UK waters” (MEMFISH)

Project Plan: Scientific Outputs: Case Studies:

Phase 1: COLLATION & Apostolaki et al. 2008 EVALUATION Metcalfe et al. 2008a Metcalfe et al. 2008b Sims et al. 2008 Sturrock et al. 2012a Year 1 Flatfish Connectivity Meeting Use existing frameworks/data Cod Mini-Symposium

Field Field Survey

Individual behaviour to population dynamics to population Individual behaviour Hunter et al. 2009a; 2012a Albaina et al. 2010 Phase 2: IMPLEMENTATION Pliru et al. 2012 Fox et al. 2012 Sturrock et al 2012b,c,d ICES Benchmark workshop on flatfish, 2009 Egg WKPESTO 2012

Natural Tags Righton et al. 2008; 2009;2010;2012 Kjesbu et al. 2010 Bendall et al. 2012a Heath et al. 2012 Larva Neuenfeldt et al. 2012 Righton & Metcalfe 2012

West et al. 2012 cycle approach cycle

- NERC-SMB Project

ICES Benchmark workshop on roundfish, 2009 Artificial Tags

ICES WKROUND, 2012 Hatchingcatching to

Juvenile Hunter & Kupschus 2007 Hunter et al. 2012b

Wholelife IFREMER Visiting Scientist 2009/10 Years 2-4 ICES WKFLAT 2012 Develop new case studies Adult Quayle et al. 2009

Hunter & de Pontual 2011 Collaboration ICES WKROUND 2012 Phase 3: INTEGRATION

Hunter et al. 2009b Bendall et al. 2009 Bendall et al. 2012b Wiegand et al. 2012 Year 5 ICES benchmarking of North Sea plaice and cod stocks, and Irish Sea Biology into FM Cod and bass stocks

Figure 1 illustrates how the scientific objectives originally proposed relate to the 3 research phases, including the four new WPs. As far as was possible, a whole life-cycle approach was applied, using both standard (e.g. field survey) and innovative techniques (e.g. gene probes, innovative atgging techniques).The phased and modular structure of the MEMFISH project proved a highly effective means of making best use of, and publishing existing data, and in generating new work in terms of new collaborations and proposals (with 16 peer-reviewed publications, 3 book chapters, 5 ICES papers and 5 more manuscripts in preparation by close of project), which, where successful, often complemented MEMFISH work at no additional cost to the project.

The MEMFISH project has aimed to provide data that would reduce uncertainty in management advice relevant to future management needs. MEMFISH work has contributed directly to ICES benchmarking excercises and working groups, and has been publicised more widely, with 24 presentations at 13 international symposia. All of the work carried out under MEMFISH has been consistent with Defra‘s evidence investment strategy, and the Marine Strategy Framework Directive (MSFD), specifically action 3, that ―Populations of all commercially exploited fish and shellfish are within safe biological limits, exhibiting a population age and size distribution that is indicative of a healthy stock‖.

Because of the numerous MEMFISH outputs, the body of the final report provides summary details of key findings, with key outputs (including proposals generated), listed at the end of each relevant section. Note that in terms of overall spend and time allocated, as agreed, WP3 and WP4 were relatively minor and more focussed components of MEMFISH. All published material resulting from MEMFISH has been included as a supplementary annex to the final report (Annex 1).

2. SCIENTIFIC OBJECTIVES & SUMMARISED RELATED RESEARCH RESULTS

Phase 1- Evaluation

1. To collate and evaluate existing knowledge on the biological processes affecting the abundance and distribution of fish species of interest to the UK (e.g. cod, plaice, sole, bass, rays etc.) and identify important gaps in understanding of population biology relevant to fish stock assessment, management and conservation. (Complete, see Section 3, Apostolaki et al. 2008; Metcalfe et al. 2008a,b; Sims et al. 2008; Sturrock et al. 2012a)

EVID4 Evidence Project Final Report (Rev. 06/11) Page 7 of 28 2. To use existing modelling frameworks to examine the value of increasing the precision of the life-history parameters identified in Objective 1. (Risk-assessment modelling was omitted from MEMFISH to be pursued in M1201 following agreement with the customer)

3. To use the output from Objectives 1 to 2 to develop and define a series of options for research to be implemented in phase 2. (Complete, see Sections 5 - 8)

Phase 2- Implementation

4. To describe and quantify the linkages between marine fin-fish spawning grounds and nursery areas. (Complete)

4.1. Conduct acoustic and fishing surveys of fish on spawning grounds to determine spawning behaviour, fecundity and egg production (Complete, see Section 6, Pliru et al. in press, Annex)

4.2. Conduct electronic tagging studies of adult fish to the locations and frequency of spawning behaviour. (Complete, see Section 6, )

4.3. Test, improve and apply existing bio-physical models of the dispersal, growth and survival of the early life stages of fish species. (Complete, see Section 6)

4.4. Evaluate and model the vulnerability of mating systems and reproductive strategies to disruption by fishing and environment to assess the need to protect fish on spawning grounds (NERC-CASE studentship). (Complete, see Section 6)

5. To describe and quantify the natal origins, movements, growth, survival, of the juveniles on nursery grounds. (Complete)

5.1. Conduct surveys of coastal sites to determine the relative abundance of juveniles on nursery grounds and collect samples for Objective 5.2. (Complete, see Section 6)

5.2. Identify the natal origins of juveniles on nursery grounds by undertaking otolith/scale micro-increment and microchemistry studies. (Complete, see Section 5)

6. To describe and quantify the linkages between nursery grounds and adult stocks. (Complete)

6.1. Use cohort analysis with historic indices of fish abundance to determine the recruitment of juveniles to adult stocks. (Not applicable in the MEMFISH WPs selected)

6.2. Re-analyse historic tagging data and conduct new conventional tagging experiments on juvenile fish to determine the scale of their movements and their recruitment to the adult stocks. (Complete, see Section 6)

6.3. Tag 200 juvenile fish (>25cm) with temperature and depth sensing electronic tags to quantify their movements and habitat choice (thermal/ depth/geographic location) of adolescent fish, and their recruitment to adult stocks. (Complete, see Section 6)

7. To describe and quantify the origins, movements, growth, survival, maturation and reproduction of adults at local and regional scales (Complete, see sections 5 & 6)

7.1. Re-analyse, if necessary, existing data from historic tagging experiments and fishing surveys to define hypotheses of stock structure in key species of commercial fish. (Complete, see Section 6)

7.2. Use existing individual based models to simulate stock mixing and segregation and assist the design and interpretation of new tagging experiments. (Complete, see Section 6)

7.3. Conduct electronic and conventional tagging programmes to quantify the movements, growth, and reproductive behaviour of adults. (Complete, see Section 6)

7.4. Use otolith and scale microchemistry and microstructure techniques to quantify the origins, growth, maturation and reproduction of adults sampled on spawning grounds (Complete, see Section 5)

Phase 3- Integration

EVID4 Evidence Project Final Report (Rev. 06/11) Page 8 of 28

8. Integrate datasets and evaluate the consequence of incorporating improved biological knowledge into fisheries management. (Complete, see section 8 )

8.1 To link case studies to the development and evaluation of management and recovery plans by delivering relevant knowledge of biological processes to other Defra-funded research programmes that focus on evaluating possible management options (e.g. those within the MF12 programme) within the Comprehensive Risk Analysis Framework. (Complete, see Section 9, Hunter et al. 2009b; Bendall et al. 2009, 2012, Wiegand et al. 2012)

8.2 To use simulation modelling frameworks to examine the implications of the new knowledge collected during the project both to evaluate the importance of incorporating biological knowledge and how to use it to provide advice on the `safe biological limits to exploitation. (Complete, see Section 9, Hunter et al. 2009b; Bendall et al. 2009, 2012, Wiegand et al. 2012)

3. COLLATION AND EVALUATION OF EXISTING DATA

3.1 Collation & Evaluation Under MEMFISH, material from the full breadth of our fish ecology R&D work, both at the UK (e.g. MF0147, MF0148, MF0152, MF0154, MF0159, MF0432 etc…) and European level (e.g. EU FAIR PL96-2079, Codyssey, PlaiceLifeLine, etc…) was re-examined, along with data on UK finfish species characteristics listed at sites including Fishbase.com, ICES.com, Seafish.com (and see also Dulvy et al. 2005, Appendix 1, MF0729) were drawn upon to compile species life-history data. These data provided an overview of the current state of knowledge with regards important ecological processes operating at the level of individuals and populations for a range of UK finfish of commercial, conservation or strategic interest. Some of these data have been incorporated into peer-reviewed publications dealing with new technologies for observing fish behaviour and how these can be applied to enhance fisheries management (Apostolaki et al. 2008, Metcalfe et al. 2008b), how fish behave in space and time (Metcalfe et al. 2008a, Sims et al. 2008) and a review of the potential application of otoliths as natural tags in the marine environment (Sturrock et al. 2012a).

By necessity, new MEMFISH R&D work had to focus on species we were realistically able to work with, or where value (in terms of specific R&D questions or policy relevance) had been identified. However, following initial consultations with Cefas fisheries scientists, it became clear that management issues dictated that area or theme-based approaches were more relevant during the collation and evaluation stage than a species-based approach, as was initially proposed. Consequently, ―species‖ workshops, as put forward in the original proposal, were limited to two only:

“Understanding and predicting connectivity in marine fish populations using plaice and sole as models for research” (Co-convened by Cefas, hosted by Ifremer, Paris, April 2007). Researchers from the UK, France, Belgium and Portugal met to compile and evaluate current knowledge, with a view to generating new research opportunities. Proposals generated as a result of this meeting (both directly and indirectly) are listed below as follows (*denotes threshold criteria met, $ denotes successful funding): ―Understanding and predicting connectivity in marine fish populations: a methodological approach‖ (FISHCONNECT) FP7-PEOPLE-2007-1-1-ITN* ―Understanding and predicting connectivity in marine fish populations: a methodological approach‖ (FISHCONNECT) FP7-PEOPLE-2009-1-1-ITN* ―Population structure and connectivity of sole in the North-East Atlantic‖ (CONSOLE) FP7-PEOPLE- 2007-2-1-IEF* ―Validation and development of otolith microchemistry in free-ranging marine fish‖ FSBI research studentship$ ―Advancing understanding of population structure and connectivity of NE Atlantic sole: Bay of Biscay‖ IFREMER/ Region Poitou Charentes ―Visiting Scientist‖ award (to E. Hunter)$

Atlantic-cod „mini-symposium‟ (co-convened by Cefas, hosted by AFBI, Belfast, May 2007). The symposium was an important contribution to both MEMFISH and MF0154, and addressed the status of existing knowledge of cod stock spatial structure, how stock structure is dealt with in a management context, and ultimately, to identify new research to improve current knowledge. Proposals generated as a result of this meeting (both directly and indirectly) are listed below as follows (*denotes threshold criteria met, $ denotes successful funding): ―Population structuring of cod around the UK‖ NERC-SMB (2008-2011)$ ―Modelling migration and aggregation: Applications in fisheries, environmental change, and evolution‖ NERC-CASE studentship$ ―Atlantic cod tagging study: North Thames Estuary‖ – Defra FSP, 2009-2010$

EVID4 Evidence Project Final Report (Rev. 06/11) Page 9 of 28 Both meetings were important in generating new opportunities linked with the MEMFISH R&D programme. Although the FP7 proposals generated by the flatfish meeting were not ultimately funded, additional consequent benefits included collaboration opportunities with IFREMER to work on sole (WP3 – see section 7) and bass (WP4 – see section 8) tagging. The cod meeting gave rise to the successfully funded NERC-SMB proposal, which has brought together the work of UK fisheries laboratories with expertise in fisheries genetics to explore the genetic links between cod sub-stocks. Building on the genetics research already completed under project MF0159, the NERC-SMB collaboration has also made full use of the tissue samples collected during MF0154 and MEMFISH field-work. The collaborations made possible as a result of MEMFISH have contributed supplementary information on connectivity in fish stocks through access to techniques not currently available at Cefas.

3.2 Use of existing model frameworks to examine added-value In order to ensure that MEMFISH was consistent with an ecosystem-based approach to fisheries management (EBFM), objective 1 was initially run in association with MF1201 (A Risk Analysis Framework For Fisheries Management). Although initial workshops were used to gather information to be applied in a risk assessment framework, key changes in project personnel meant that it was no longer practical to pursue this technique within MEMFISH. Following agreement with the customer, this element was dropped from the MEMFISH portfolio, and risk classification work has been pursued in MF1201.

3.3 R&D options The projects proposed as a result of phase 1 (see sections 5-8 below) were put forward to develop analytical and modelling processes that would add value to the project by combining MEMFISH data with data gathered over many years, and from additional satellite projects. The focus of new work was on the relationships between fish population subunits in spawning areas, on nursery grounds, and on feeding areas, how these vary year-on- year, and the contribution of the environment to such variation:

• WP1: Plaice population dynamics and extension of the North Sea Population Movement Simulator (NSPMS) • WP2: Population structure and connectivity of cod in ICES area VII • WP3: Population structure and connectivity of sole in UK Waters • WP4: Distribution and abundance of sea-bass in UK waters in relation to climate change

Data from the new WPs have been interpreted in relation to the marine environment in the broadest sense (i.e. including human activities, local and global changes in habitats). Regional case studies have targeted issues of match or mismatch between metapopulation dynamics and management scales, conservation of essential fish habitats and the evaluation of climate change on habitat and biology. The strength of the multidisciplinary approach applied by MEMFISH is that the combined data streams have provided a more comprehensive understanding of life-history characteristics than any one technique in isolation might have achieved.

SUMMARY OF “COLLATION AND EVALUATION” SCIENTIFIC OUTPUTS & CONTRIBUTIONS Peer-reviewed papers: Apostolaki et al. 2008; Metcalfe et al. 2008a,b; Sims et al. 2008; Sturrock et al. 2012a Conferences: ―Third International Bio-logging Science Symposium‖, Pacific Grove, California, USA, August 2008. ―Advances in fish tagging and marking technology‖, Auckland, New Zealand, February 2008. ―Seventh Conference on Fish Telemetry held in Europe‖, Silkeborg, Denmark, June 2007. ―Tagging and tracking marine fish with Electronic Devices‖, San Sebastian, Spain, September 2007.

4. PLAICE POPULATION DYNAMICS AND EXTENSION OF THE NORTH SEA POPULATION MOVEMENT SIMULATOR (WP1)

4.1 Introduction The life-history biology and ontogenetic movements of North Sea plaice are relatively well understood, however the work conducted in WP1 was aimed at obtaining a fuller understanding of the relationship between spawning areas, nursery grounds and feeding areas (figure 2). Phase 1 activity generated several proposals related to WP1, and although funding for a collaborative EU-funded research project was not ultimately successful, we were successful in obtaining several smaller, but strategically important grants, and external collaboration played a significant role in completing WP1. In the research phase (phase 2), the dynamics of early life-history have been considered through examination of data collected in the Irish Sea from both MF0432 (PREDATE) and M1102 surveys (in collaboration with SAMS, Dunstaffnage, and The University of Bangor). As well as developing and enhancing novel methodologies, results from these studies have significant implications with regards the recruitment of plaice and other commercially exploited species. The movements of adult fish were re-examined (in collaboration with Simon Fraser University, Canada) based on re-analysis of electronic data storage tag (DST)data collected under projects EU-FAIR PL96 2079, MF0147 and MF0152. These data have implications with regards the spatial and temporal catchability of plaice, and the visibility of plaice to survey across the North Sea. Aspects of the whole life-cycle can in theory be interrogated through examination of the

EVID4 Evidence Project Final Report (Rev. 06/11) Page 10 of 28 chemistry of fish ear-stones or ―otoliths‖. For the first time, we have been able to experimentally examine the detailed seasonal deposition of elements into male and female plaice (in collaboration with NOCS, Southampton, and the University of Montpellier 2). This work significantly advances our understanding of fish blood chemistry, and the physiological and environmental factors that influence elemental deposition in the otolith. These observations have been coupled with new techniques originally developed under MF0152 to re- examine the chemical signatures laid down in the otoliths of fish tagged with electronic data storage tags (DSTs). This represents the first full assessment of the level of geographical information that can be extracted from the chemistry of otoliths from fully marine fish. Data from these studies have been used to better parameterise the biologically-realistic NSPMS model over the life history of the fish.

Phase 2: Phase 1: COLLATION & Impacts of predation on Phase 3: INTEGRATION EVALUATION plaice eggs & larvae, and implications for recruitment Albaina et al. 2012; Hunter et al. 2012a; Fox et al. 2012; Pliru et al. 2012 • Full age-structure and new information on plaice • Flatfish Connectivity Meeting population structure Otolith elemental chemistry – inorporated into North Sea • Dialogue with fisheries Disentangling environmental & plaice movement simulator scientists, Defra, Physiological influences to assess

stakeholders, etc… “natural tag” capacity of otoliths in • Biological data provided to fully marine fish ICES benchmarking of • Contributions to review Sturrock et al. 2012b, c, d North Sea plaice stocks + papers (Apostolaki et al. ICES WKPESTO 2008, Metcalfe et al. 2008a,b Spatial and temporal migration • Hunter et al. 2009b Sturrock et al. 2012b) and behaviour patterns of adult plaice, and implications for seasonal catchability & detection by survey Hunter et al. 2009a, b

Figure 2 A schematic illustrating the overall contribution of WP1 to MEMFISH in the context of all 3 research phases. Material reviewed during phase 1 contributed to 4 review papers, and fed into the main research phase (phase 2). New research in phase 2 followed 3 main themes ( impact areas of biological evidence in italics, scientific publications generated in bold), which were then integrated in phase 3.

4.2 Otolith microchemistry

MEMFISH studies of otolith microchemistry were undertaken mainly through collaboration around an FSBI funded studentship, ―Environmental and physiological influences on otolith chemistry in a marine fish‖ (2008- 2012), co-supervised with the National Oceanographic Centre (NOCS), University of Southampton, and an ongoing collaboration with the University of Montpellier II. The results from work originating from projects M0152 and EU MC-IEF PlaiceLifeLine were re-analysed and extended, and new research to develop and validate otolith microchemistry techniques was applied to otoliths from plaice tagged with electronic data storage tags (DSTs), whose movements have been reconstructed using DST data.

For stock discrimination and inferring connectivity patterns in fish stocks, an understanding of the mechanisms underpinning elemental incorporation into otoliths is required to liberate the full potential of otoliths as natural tags. In MEMFISH we reviewed the current understanding of the biochemical behaviours exhibited by the otolith elements most frequently used in population studies. In the same review (Sturrock et al. 2012a), individual case studies in which otolith microchemistry has been applied to describe the movements of fully marine fish species were examined. These have then been compared with new data for North sea plaice. MEMFISH results have suggested that otolith Ba/Ca and Li/Ca appeared to be under environmental control, while influences on Sr/Ca and Mg/Ca ratios were less clear, and may be subject to physiological influence. These data are currently being prepared for publication (Fig.3, Sturrock et al. 2012d).

An aquarium study was designed to disentangle the environmental vs. physiological influences on otolith chemistry. This is the first study to measure a suite of trace elements in any blood fraction from a marine fish, and the first to analyse fish plasma by any form of inductively coupled plasma-mass spectrometry (ICP-MS). Consequently, a manuscript has been prepared containing reference ranges for 12 minor and trace element concentrations in the plasma of plaice (Sturrock et al. 2012b). These represent the first measurements of Mn, Ba and Pb in any blood fraction for a marine species, and the first measurements of Rb, Ba and Pb in plasma from any fish species. These reference ranges have applications in fisheries, physiology, aquaculture, nutrition, biochemistry, ecotoxicology and animal health.

EVID4 Evidence Project Final Report (Rev. 06/11) Page 11 of 28

Figure 3 a) Map showing the migrations of male (blue) and female (red) plaice from the west and central North Sea (WNS, CNS) feeding stocks. The orange arrows indicate approximate pre-spawning migration routes, while blue arrows represent post-spawning migrations. The colour of the daily positions intensifies with time after release and represent the “most probable tracks” (estimated using Pedersen et al. (2008) HMM geolocation technique) from returned electronic data storage tag records of between 366 and 560 days. Release sites are shown by a plus (+) sign, recapture sites by a cross (x). The WNS female recapture site was within 10 kilometres of the release site and is not displayed. Note that the WNS male performed 2 pre-spawning migrations and was recaptured midway during the second post-spawning migration. b) Average otolith Mg/Ca, Li/Ca, Sr/Ca and Ba/Ca ratios (± S.E.M.) for the first 12 months of DST records (October 2004-05) for male (M) and female (F) plaice from the west and central North Sea (WNS, CNS) feeding stocks.

Figure 4 Physiology (temperature, salinity, Fulton‟s condition factor and total blood protein), blood chemistry (Ca, Cu, Sr, Se, Zn & Mg), and otolith chemistry (Sr/Ca and predicted δ18O profiles) of male and female plaice held in ambient conditions at the Cefas Laboratory Aquarium, Lowestoft from June 10 – June 11.

A summary of MEMFISH findings with regards environmental and physiological influences on otolith chemistry are presented in Fig. 4. Aquarium reared plaice showed clear seasonal changes in physiology, in keeping with observations from wild fish. Blood Ca and Sr were higher in female plaice, and peaked during the spawning period (pink bar, Fig. 3, Annex 1). However Ca peaked at the beginning of the spawning period, following vitellogenesis, while Sr peaked with GSI (gonadosomatic index). Blood Zn exhibited a clear negative relationship

EVID4 Evidence Project Final Report (Rev. 06/11) Page 12 of 28 with GSI in females. Blood Cu, Se (and possibly Mg) appeared correlated with condition and total protein concentrations, peaking just before the spawning period, although generally remaining higher in the males. A poster presentation detailing this work took the ―best poster‖ prize at the 8th International Flatfish Symposium, and the work is currently being prepared for publication (Sturrock et al. 2012c).

In addition, distinct patterns in otolith δ18O from the OTC marked aquarium fish provide an intra-annual timeline within each otolith to precisely match otolith material with the variables listed above. A method has been developed in R to allow automated curve matching. By providing new insight into how otolith chemistry records ambient conditions, these techniques should enhance our ability to geolocate individual fish in space and time. For those elements controlled primarily by fish physiology, their value as markers for fish migration is reduced, however they may prove equally useful for estimating age-at-maturity and spawning frequency, also key parameters in stock assessment and fisheries management.

4.3 Population Dynamics

4.3.1 Identification of plaice egg predators using molecular probes Mortality during the egg and larval stages is thought to play a major role in determining year-class strength in many marine fish and predation is normally considered the main cause, but the full suite of predators responsible has rarely been assessed. In an ongoing collaboration with SAMS and the University of Bangor, potential predators on a patch of plaice eggs located in the eastern Irish Sea (data originally collected and compiled under M0432) were mapped using acoustics and sampled by trawl and a plankton multi-net (Fox et al. 2012). Gut contents of 3,373 fish, crustacea and cephalopods sampled in the area were screened using a plaice-specific TaqMan DNA probe (see Albaina et al. 2010, Hunter et al. 2012 for MEMFISH work validating gene probe methodology). Herring (Clupea harengus) and sprat (Sprattus sprattus) dominated trawl catches and showed high positive TaqMan responses (77% and 75% of individuals tested respectively). Locations of clupeid schools also broadly corresponded with the distribution of fish eggs in the plankton. Whiting (Merlangius merlangus) were also reasonably abundant in trawl hauls and 86% of their stomachs tested positive for plaice DNA. Species showing lower levels of positive TaqMan response included mackerel (Scomber scombrus), poor cod (Trisopterus minutus), squid (Loligo spp.), dogfish (Scyliorhinus canicula) and weever (Trachinus vipera). Non-reactivity of all negative controls precluded the occurrence of cross-contamination and positive reactions from demersal species, such as dogfish and weever, may have resulted from secondary predation. No benthic macro-crustaceans tested positive. Samples of planktonic organisms yielded 13% positive TaqMan reactions, mainly from clupeoid or sandeel (Ammodytidae) larvae, but also included some malacostraca and amphipoda. Use of the molecular approach allowed rapid screening of a large number of potential predators of plaice eggs and the results provide a more holistic description of the predator community than has previously been achieved.

4.3.2 Impact of sprat predation on plaice egg mortality Although the causes of fish egg and larval mortality are poorly understood, predation is thought to be the main contributing factor. To investigate the impact of predation on the young life-history stages of plaice, gut content analysis of sprat samples originally collected under M0432 was undertaken, in collaboration with EU project FACTS (Pliru et al 2012). These new results were combined with acoustic data recorded on the M0432 survey during February 2009. Acoustic observations and analysis of stomach contents revealed diurnal behaviour with dense feeding schools forming during daylight, which dispersed into thinly spread aggregations at dusk. Of 338 stomachs analysed, 95% contained identifiable prey items. The maximum number of ingested prey items peaked between 10:00 and 18:00 for all food groups. Numerically gadoid eggs were the most frequently consumed prey (64%), followed by copepods (25%) and plaice eggs (7%), but plaice eggs were present in 91% of the stomachs analysed (see Annex 1). Converting stomach content data to daily consumption suggested that sprat may consume 73% of all plaice eggs spawned in the area. Fish eggs may be an important energy source for sprat in the Irish Sea during spring time, when alternative prey is scarce. Sprats seem to be the dominant source of plaice egg mortality so the abundance and distribution of this clupeid may have important consequences for the recruitment dynamics of other fish species (Pliru et al., 2012)

4.3.3 Large-scale variation in seasonal swimming patterns The vertical swimming activity of mature female plaice. tagged with electronic data storage tags (data collected under EU-FAIR PL96-2079, M0127 and M0152) was examined to test whether swimming at different times of the year differed among areas of the North Sea with average tidal current velocities ranging from fast (West), to intermediate (East), or slow (North). Longer duration and tidal swimming activity were predicted for the western group, where fast-flowing tidal currents allow efficient selective tidal stream transport (Hunter et al. 2009a). Individual depth data were converted into binary records representing swim versus rest time, and repeated patterns of swimming were analysed according to cycle-length frequencies. Most swimming occurred during expected times of migration and spawning (October – March). Plaice infrequently spent > 5 h in mid-water, and rarely left the sea-bed during summer. As predicted, DST1 tagged plaice (West only), spent the longest times

EVID4 Evidence Project Final Report (Rev. 06/11) Page 13 of 28 swimming (P > 0.001), but there was no significant effect of Area for DST3s (all areas), suggesting that swimming plays an important behavioural role in migration in addition to transport between feeding and spawning areas. Tidal patterns of activity occurred in all three sub-stocks, predominantly during the migratory period, albeit at a significantly lower frequency in the North. These data provide one of few examples where a fish stock‘s annual behaviour patterns have been recorded across a large part of its geographical range. The results have important implications for understanding the energetics of fish migration, and the availability of demersal stocks to capture by commercial and survey vessels.

4.4 Implementation of the NSPMS model. The reconstructed movements of male and maturing female plaice tagged with DSTs were recalculated using ―guided reconstruction‖ performed using the HMM geolocation technique (Pedersen et al. 2008). Juvenile plaice mark-recapture data were summarised spatially (ICES rectangle) and temporally (decadal), including a dislocation and dispersion analysis. The model was further enhanced by the addition of age-structure, and estimations of mean weight for age, sex and time of yearm making it fully compatible with ICES assessment methodology. The effects of a range of area and seasonal closures were tested in terms of reduction in overall Addition to NSPMS of age-structure, and estimations of mean weight for age, sex and time of year. fishing mortality (Hunter et al. 2009).

4.5 SUMMARY OF WP1 SCIENTIFIC OUTPUTS & CONTRIBUTIONS Peer-reviewed papers: Albaina et al. 2010; Fox et al. 2012; Hunter et al. 2009a,2012a; Plirù et al. 2012; Sturrock et al. 2012b,c,d ICES Contributions: Benchmark Workshop on Flatfish ICES HQ, Copenhagen, Denmark, 6–13 February 2009 ICES WKPESTO, February 2012 Conferences: ―World Fisheries Congress‖, Edinburgh, May 2012. ―Eighth International Flatfish Symposium‖, IJmuiden, Netherlands, November 2011. ―Fish Diversity & Conservation‖, Bournemouth, UK, July 2011 ―ICES ASC‖, Gdansk, Poland, September 2011 ―Fourth International Science Symposium on Bio-logging‖, Hobart, Tasmania, March 2011 ―American Fisheries Society Symposium‖, Pittsburgh, Pensylvania, USA, September 2010 ―Fish & Climate Change‖, Belfast, July 2010 ―ICES ASC‖, Berlin, Germany, September 2009 ―Fourth International Otolith Symposium‖, Monterey, USA, August 2009 ―Society for Experimental Biology‖ Satellite meeting, Sete, France 2008 ―Seventh International Flatfish Symposium‖, Sesimbra, Portugal, November 2008. ―Advances in fish tagging and marking technology‖, Auckland, New Zealand, February 2008. ―Seventh Conference on Fish Telemetry held in Europe‖, Silkeborg, Denmark, June 2007. ―Tagging and tracking marine fish with Electronic Devices‖, San Sebastian, Spain, September 2007.

Supplementary awards: ―Validation and development of otolith microchemistry in free-ranging marine fish‖ FSBI research Studentship (to A. Sturrock, sponsored by NOCS & Cefas) “Disentangling environmental and physiological influences on otolith growth: unlocking the potential use of otoliths as natural tags” (to Trueman, Sturrock & Hunter, access to NERC ion- microprobe facility, School of Geosciences, Edinburgh University, UK)

5. POPULATION STRUCTURE AND CONNECTIVITY OF COD IN ICES AREA VII (WP2) Phase 2:

Phase 1: COLLATION & Field survey of Irish Sea spawning Phase 3: INTEGRATION EVALUATION grounds, assessment of fecundity in relation to environmental experience • Atlantic cod mini- • New information on cod Kjesbu et al., 2010 population and genetic symposium structure (Heath et al., in prep., Righton et al., in • Dialogue with fisheries Assessments of environmental prep) scientists, Defra, experience of cod in the North and stakeholders, etc. Irish Seas: growth, behaviour and migration. Righton et al., 2009, • Biological data provided to ICES benchmarking of • Evaluation of existing Righton et al. 2010. North Sea and Irish Sea data and contributions to cod assessments in 2009 review papers (Metcalfe Spatial dynamics of cod in UK and 2012 et al. 2008a,b, Righton et waters • al., 2008, Neuenfeldt et al., 2012) Righton et al., in prep.

Figure 5 A schematic illustrating the overall contribution of WP2 to MEMFISH in the context of all 3 research phases. Material reviewed during phase 1 contributed to 4 review papers, and fed into the main research phase (phase 2). New research in phase 2 followed 3 main themes ( impact areas of biological evidence in italics, scientific publications generated in bold), which were then integrated in phase 3.

5.1 Introduction The stock structure of cod populations, and the way that they interact, in ICES area VII remains uncertain, with little existing information on the location of nursery grounds, the connectivity between nursery grounds and adult

EVID4 Evidence Project Final Report (Rev. 06/11) Page 14 of 28 habitats or the detail of migrations of adults between feeding and spawning grounds (conventional tagging only provides information on release and recapture positions, and nothing in between). Furthermore, the underlying aspects of recruitment dynamics are still uncertain, and the links between spawning, climate and population structure are still poorly understood. For example, comparison of egg-based assessments for cod in the Irish Sea with results of conventional assessments has in the past proved difficult due to uncertainty about movements of the fish between areas.

The aim of WP2 was to evaluate the population structure and connectivity of cod in ICES area VII using a multidisciplinary and collaborative (with the Marine Institute, Galway and AFBI, Belfast, plus involvement with the NERC SMB cod genetics programme) approach that combined data from survey-based assessments of adult and egg distributions, electronic tag-based assessments of adult movements and behaviour, and field-based assessments of population structure and fecundity (figure 5).

5.2 Field Survey of biological attributes of cod spawning aggregations in the Irish Sea 2009-2011 The MEMFISH field surveys 2009-2011 focussed on the distribution of cod at known spawning areas in the Irish Sea, and assessed the biological characteristics of individual cod on the spawning grounds. Information on the size, weight and body condition of individual cod (including hepatosomatic index), and reproductive (sex, age, maturity, gonad weight) and feeding (stomach contents) status were collected. In addition, genetic samples were taken from each fish, which were supplied to the NERC-SMB project ‗Population structure of cod around the UK: scale, mechanisms and dynamics‘. Note that the full cruise reports for all 4 MEMFISH field surveys, including details of the methods applied and the results obtained, have been appended to this report (annex 1).

Box 1 755 768 54°30' 742

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53°30' Box 4 53°10' 5°50' 5°40' 5°30' 5°20' 5°10' 5°0' 4°50' 4°30' 4°20' 4°10' 4°0' 3°50' 3°40' 3°30' 3°20' 3°10' 3°0' Longitude Longitude Figure 6 Stations in the a) western and b) eastern Irish Sea sampled during the 2011 survey, 18 February–4 March 2011. Station numbers are shown where the trawl was shot, and the lines indicate the trawl haul transects. Grey circles indicate commercial trawlers cod cpue (February 2007–2009, averaged over fishing trips, from VMS). „Survey boxes‟ are indicated by red lines, and two underwater gas pipelines running from Scotland to Northern Ireland are indicated by brown dotted lines.

5.2.1 Overall catch composition in the western and eastern Irish Sea Important differences in general catch composition were observed between the western and eastern Irish Sea. Herring and sprat were the dominant pelagic species in both areas: but whereas herring outnumbered sprat in the west, the reverse was true in the east where sprat was the most numerically abundant species overall. Norway lobster was far more abundant in the west than in the east. Various gadoid species were much more commonly found in the west, most notably haddock, Norway pout and blue whiting, although whiting was ubiquitous. In the east, a range of flatfish species were very common, especially dab, plaice, flounder and solenette.

5.2.2 Distribution and abundance of cod In total, 221 Irish Sea cod were caught during 2011. These were distributed fairly evenly over the spawning ground study areas in the western and eastern Irish Sea (Fig. 6). In the western Irish Sea, 168 cod were caught

EVID4 Evidence Project Final Report (Rev. 06/11) Page 15 of 28 in 73 hauls of 1 hour duration each, i.e. at an average catch rate of 2.30 per hour. This was lower than our mean catch rates in the western Irish Sea 2009 and 2010 surveys (3.29 and 2.50 cod per hour respectively). However, it should be noted that some of the northernmost stations on rocky grounds, where many small juvenile cod occur, were not sampled during 2011 (after extensive gear damage experienced at these sites in 2010 and 2009).

Table 1 A comparison of the most abundant fish and shellfish species sampled in the western and eastern Irish Sea (2011 survey data).

Western Irish Sea (74 stations) Eastern Irish Sea (21 stations) 187273 Herring Clupea harengus 103007 Sprat Sprattus sprattus 49254 Whiting Merlangius merlangus 84543 Herring Clupea harengus 42748 Sprat Sprattus sprattus 20090 Dab Limanda limanda 16145 Norway lobster Nephrops norvegicus 18539 Whiting Merlangius merlangus 9322 Dab Limanda limanda 2866 Plaice Pleuronectes platessa 7550 Haddock Melanogrammus 595 Norway lobster Nephrops norvegicus aeglefinus 5085 Norway pout Trisopterus esmarki 282 Lesser sp. Scyliorhinus canicula dogfish 4096 Lesser sp. Scyliorhinus canicula 210 Flounder Platichthys flesus dogfish 3497 Grey gurnard Eutrigla gurnardus 208 Lesser weever Echiichthys vipera 1861 Red gurnard Aspitrigla cuculus 183 Solenette Buglossidium luteum

In the eastern Irish Sea (not visited in 2010), cod catch rates were far higher than in 2009 when only 17 individuals were caught in a total of 26 hauls (mean 0.65 per hour); this year 52 cod were caught in 21 hauls (mean 2.48 per hour). Whereas in 2009 cod were much less common in the eastern than western Irish Sea, in

2011 there appeared to be no appreciable difference in cod abundance between east and west (Fig. 7). 54°30'

+ + ++

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+ Latitude + + + +

+ + +

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+ 53°30' Cod per hour 0 1 5 10 +

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Figure 7 Distribution of cod catches in the Irish Sea. Symbol sizes indicate catch rates as number of cod per 1-hour haul (+ indicates a station where no cod were caught).

5.2.3 Size composition of cod on spawning grounds In the western Irish Sea, a broad size distribution of cod was represented, including juveniles, smaller and larger mature fish. There were more large individuals present in 2011 than in 2010 (Fig. 8). A few very large cod were caught; the largest two individuals (both females) measured 105.5 and 99.0 cm and weighed 13.56 and 11.66 kg, respectively. Adult cod showed a peak in the length distribution around 60–75 cm, progressed from a peak around 40–60 cm in 2010.

In 2009, few medium to large adult cod and no juveniles were encountered in the eastern Irish Sea. In 2011, a wide range of sizes was observed, including juveniles and medium- to moderately large adult cod, although no cod >80 cm were found in this survey. This tentatively suggests a rejuvenated, growing young population of cod in the eastern Irish Sea.

EVID4 Evidence Project Final Report (Rev. 06/11) Page 16 of 28 (a) 2009 West (b) 2009 East 0.4 0.4 Females 0.3 0.3 Males

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Number per 10 h fishing10 per Number h fishing10 per Number 10 20 30 40 50 60 70 80 90 10 20 30 40 50 60 70 80 90

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Number per 10 h fishing10 per Number h fishing10 per Number 10 20 30 40 50 60 70 80 90 10 20 30 40 50 60 70 80 90 Length Length Figure 8 Length distributions of female (black bars) and male cod (grey bars) observed in the western (left) and eastern (right) Irish Sea during the Memfish surveys 2009 (a, b), 2010 (c, d), and 2011 (e, f).

Table 2. Numbers of female and male cod (and some unsexed individuals) smaller and larger than 30 cm, caught in the 2009–2011 surveys. Sex ratios expressed as % females and % males of the total numbers of known-sex cod. Figures shown separately for the western and eastern Irish Sea.

Area Size Year Females Males Unsexed % % Males Females West <30 cm 2009 28 21 0 57% 43% 2010 29 17 1 63% 37% 2011 22 11 3 67% 33%

>30 cm 2009 71 28 0 72% 28% 2010 93 128 1 42% 58% 2011 80 52 0 61% 39%

East <30 cm 2009 0 0 2 — — 2011 8 4 0 67% 33%

>30 cm 2009 8 7 0 53% 47% 2011 8 32 0 20% 80%

5.2.3 Sex ratio of cod There were local differences in the sex ratio of cod for (near-) adult individuals (here defined as >30cm long). In the east, far more male than female (near-) adult cod were found (80% male, 20% female) – notably, several catches contained groups of 4–7 males only. By contrast, in the west, more female than male (near-) adult cod were caught (61% female, 39% male). The sex ratio in the west was similar to that in 2009 when females also clearly predominated here (72%), but quite different from the intervening year when females were less often caught than males (42% only). Amongst juveniles (<30 cm), females also predominated in both the western and eastern Irish Sea (67% in both regions).

5.2.4 Maturity and reproductive characteristics of cod

EVID4 Evidence Project Final Report (Rev. 06/11) Page 17 of 28 (a) Females 2009 (b) Males 2009 50 50 I I 40 M 40 M Number H Number R 30 R 30 S 20 20

10 10

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20 20

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0 0 10 20 30 40 50 60 70 80 90 100 10 20 30 40 50 60 70 80 90 100 Length Length

Figure 9 Maturity stages by 10-cm length groups of cod, for females (left) and males (right) caught during the surveys of 2009 (a, b), 2010 (c, d) and 2011 (e, f). I immature; M maturing; H hyaline (females only); R running; S spent; A atretic.

Maturity stages determined in all 3 years showed that all cod <30 cm were immature, and all cod >60 cm were mature (Fig. 9). Maturity in females during 2011 tended to be at a further advanced stage than in both earlier surveys, in spite of almost the same timing of the survey within the year and a very cold winter preceding. The proportion of females in the more advanced maturity stages (i.e. hyaline, running and/or spent) was 27%; this was 10% in 2009 and 14% in 2010. Conversely, maturity in males in 2011 was less advanced than in 2010 but more than in 2009; the proportion of males in the advanced maturity stages (running and spent) was 18% in 2009, 33% in 2010 and 21% in 2011. Only 2 spent males were observed, both during 2011.

Length–weight and length–gonad relationships for the 2011 Memfish survey are shown in Fig. 10, for female and male cod. These figures show a steady length–weight relationship, and also illustrate that gonadal investment remains quite limited for cod up to 50 cm length. Above that length, however, cod invest heavily in gonadal development. The highest ovary weights recorded were 2804 g and 2602 g, above the highest ovary weights recorded in 2009 (2529 g) and 2010 (2084 g).

5.2.4 Length–weight relationships The length–weight relationships for cod sampled during 2009, 2010 and 2011 are shown in Fig. 11. The relationships were highly similar during the initial two years, but during the current year tended to be slightly lower, indicating a slight decrease in the average body condition of cod at a given length.

5.3 Stock identification

5.3.1 Spatial dynamics of cod in the North Sea The results of a large-scale electronic tagging programme of Atlantic cod in the North Sea, spanning the years 1999 to 2008 was reported at ICES ASC 2008 (Righton et al. 2008). During this time, 1126 tags were deployed on cod between 37cm and 110cm, and 263 tags were returned yielding over 31,000 days of data. A tidal geolocation model was used to determine the daily locations of each cod, and to ascertain the scale and rate of their movements. This dataset was used to describe fundamental features of cod ecology, such as the balance

EVID4 Evidence Project Final Report (Rev. 06/11) Page 18 of 28 between homing and resident behaviour, the location of feeding and spawning grounds (and the migratory pathways between them), and the seasonality of migration and its impact upon sub-stock structure. The results provide a step forward in our understanding of how Atlantic cod exploit and utilise their environment and, as a result, have implications for our expectations of how cod stocks may recover following depletion (Righton et al. 2008). (a) Females 14000

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Figure 10 Length–weight relationships (left panels) and length–gonad weight relationships (right panels) of (a) female and (b) male cod caught in the 2011 survey. For each individual fish, the colour of the symbol indicates its maturity stage (see legend: I immature; M maturing; H hyaline [females only]; R running; S spent). (a) Females (b) Males 14000 14000

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Figure 11 Length–weight relationships of (a) female and (b) male cod during the 2009, 2010 and 2011 surveys, shown for the western Irish Sea only.

5.3.2 Spatial dynamics of cod in the Irish & Celtic Seas

Work to describe cod movements between two adjacent cod management areas, one in the Irish Sea (ICES area VIIa) and the other in the western English Channel and Celtic Sea (ICES areas VIIe–k), commenced in early 2008 with a joint cruise (collaboration with AFBI), and a joint cod tagging programme (with AFBI and MI). The cod tagging fieldwork programme was completed in March 2010. A summary of all cod tagged and recaptured (to date) as part of this work is provided in table 3 (below).

Table 3 Details of DST-tagged cod released and recaptured by month and year of release in each ICES management area. Mean fish lengths in cm ±SD. Tracks for all cod at liberty > 10 days were reconstructed using the Hidden Markov Model (HMM).

EVID4 Evidence Project Final Report (Rev. 06/11) Page 19 of 28

Area Year Month ICES DSTs Length Method of DSTs Cod tracks Rectangle capture returned reconstructed (using HMM) VIIa 1999 March 35 & 36 E4 19 64±9 Long line 4 4 2008 February 35 & 36 E4 17 67±6 Midwater trawl 3 3 2009 Feb- March 35 & 36 E4 80 72±10 Midwater trawl 18 17 2010 March 36 & 37 E6 44 70±13 Midwater trawl 9 8 Total:- 160 33 32

VIIe 2007 Jan – Feb & 29 E5 & 28 E7 169 72±7 Fixed gill net & 27 17 May hook and line Total:- 169 27 17

VIIf 2004 March 31 E5 & 31 E6 13 68±10 Otter trawl 1 1 2008 May 31 E5 & 31 E6 17 67±9 Otter trawl 1 1 2010 March 30 E4 & 30 E5 60 75±8 Fixed gill net 6 6 Total:- 90 8 8

VIIg 2008 March 31 E2 & 32 E2-3 100 77±9 Seine net 9 9 2009 March 31 E2 & 32 E2-3 141 71±9 Seine net 18 18 2010 March 31 E2 & 32 E2-3 50 Seine net 10 6 Total: 291 38 33

Figure 12 Spatial distributions of 81 cod (11,170 days of data) released in the following ICES areas: eastern Irish Sea VIIa (yellow; 8 cod = 1457 days of data), western Irish Sea VIIa (red; 24 cod = 4574 days of data), eastern Celtic Sea VIIf (white; 7 cod = 1135 days of data), western Celtic Sea VIIg (dark blue; 24 cod = 2533 days of data), western English Channel VIIe (green; 17 cod = 966 days of data). Coloured circles show individual daily geolocated positions.

Out of 106 DSTs returned to date (29 February, 2012), 81 cod tracks (>10 days at liberty) have been reconstructed using HMM geolocation (Pedersen et al. 2008) to estimate the most probable daily locations for each individual cod throughout their time at liberty (Results shown in Fig. 12 for cod tagged in adjacent ICES areas VIIa & VIIe-VIIg). These results suggest movement between different ICES areas, particularly within the English Channel (VIId-e) and adjacent areas of the Celtic Sea (VIIf-h), where adult cod seem to mix in deep offshore feeding grounds (―Hurd Deep‖, the deep water mid-Channel trench within ICES areas VIIe & VIIh) during spring and summer. Eastern and western Irish Sea (VIIa) cod appear to use the deep water of the North Channel and St George‘s Channel as migratory ‗highways‘ during the spring and summer, at which time they head either north into VIIa, or south into the Celtic Sea (VIIg) to feed. By contrast, Celtic Sea cod tagged in VIIg tended to remain within VIIg throughout the year, with very little movement into neighbouring sub-divisions VIIj and VIIh.

Analyses of historic mark recapture data further suggests that cod tagged on spawning grounds in VIIa, VIIf and

EVID4 Evidence Project Final Report (Rev. 06/11) Page 20 of 28 VIIg tend to return to the same spawning grounds in successive years. Irish Sea and Celtic Sea cod generally appear to remain within these respective management areas during the autumn and winter, then disperse more widely during the spring and summer, resulting in some crossover into neighbouring management areas. Cod tagged on spawning grounds in VIIe, however, appear to migrate to other adjacent spawning grounds in neighbouring areas VIId, VIIf & VIIg during the winter within the same spawning year.

SUMMARY OF WP2 SCIENTIFIC OUTPUTS & CONTRIBUTIONS Peer-reviewed papers: Righton et al. 2008;2010;2012; Kjesbu et al. 2010; Bendall et al. 2012, Heath et al. 2012, Neuenfeldt et al. 2012, Righton & Metcalfe 2012, West et al. 2012. ICES Contributions: Benchmark Workshop on Roundfish ICES HQ, Copenhagen, Denmark, 16–23 January 2009 ICES WKROUND Aberdeen 22–29 February 2012 Conferences ―ICES ASC‖, Berlin, Germany, September 2009 ―Third International Science Symposium on Bio-logging‖, Pacific Grove, California, USA, August 2008. ―Advances in fish tagging and marking technology‖, Auckland, New Zealand, February 2008.

Supplementary awards: ―Population structuring of cod around the UK‖ NERC-SMB, 2008-2011 West, Christopher (2010) Modelling migration and aggregation: Applications in fisheries, environmental change, and evolution. PhD thesis, University of York (co-supervised by Cefas). “Atlantic cod tagging study: North Thames Estuary” – Fisheries-Science Partnership project, 2009-2010.

6. POPULATION STRUCTURE AND CONNECTIVITY OF SOLE IN UK WATERS (WP3)

6.1 Introduction In UK waters the Channel sole fisheries represent a commercially valuable asset, both at the regional and national level. Western Channel sole support a sizeable beam-trawl fleet on the southern English coast. ICES, however, has indicated some concerns about the sustainable future of the stock, and since 2003, has been advising a recovery plan. The EC (Council Regulation No 509/2007 of 7 May 2007) has established a multi- annual plan for the sustainable exploitation of the Western Channel sole, developed in association with industry and scientists. However, some of the basic questions about this stock‘s migration patterns and recruitment dynamics need to be answered in order to support stock assessments and the provision of appropriate technical measures. Connectivity between sole populations has never been quantified, and there is little existing information on the movements of juveniles within and between nursery areas, the connectivity between juvenile and adult habitats, or the migrations of adults between feeding and spawning grounds. The relative contribution of juvenile habitats to the recruitment in any specific stock and the ecological limits of each stock, remain key questions to be answered in order to achieve sustainable management. The overall aim of WP3 of MEMFISH was to initiate an evaluation of the population structure and connectivity of sole in UK waters using a multidisciplinary approach that combines natural (otolith microchemistry and parasites) and artificial tags (mark- recapture and DSTs). It was never our intention to achieve this directly, but rather, through seeking additional funding for a focussed large-scale project.

In addition, based on the proposal ―Advancing understanding of population structure and connectivity of NE Atlantic sole: Bay of Biscay‖ (submitted to IFREMER/Region Poitou Charentes), E. Hunter was awarded the position of ―IFREMER visiting scientist‖ from September 2009 until June 2010, having successfully obtained funding from both IFREMER (3 months) and region Poitou-Charentes (7 months). The aims of the work undertaken as IFREMER visiting scientist were to strengthen links between Cefas and IFREMER, and to extend the scope of MEMFISH (at no additional cost to the project). The specific aims and objectives of the collaboration were to 1) Participate in ongoing research into population connectivity of sole populations with a focus on methodological development (otolith microchemistry and tagging), and 2) Co-development of EU FP7 proposal, ―Understanding and predicting connectivity in marine fish populations‖

6.2 Exploration of best-practice for attachment of electronic data storage tags (DSTs)

Figure 14 Internal tagging of the sole, Solea solea. From left to right, insertion of dummy G5 tag into the body cavity, closure of the wound using sutures, sole immediately post-operation.

Experiments were conducted in the laboratory aquarium at IFREMER, L‘ Houmeau, between October 2009 and June 2010. Sole of >19 cm TL were collected by 2 m beam trawl from the Pertuis Charente during October 2009

EVID4 Evidence Project Final Report (Rev. 06/11) Page 21 of 28 (n = 25). Each individual was measured (TL, SL), weighed, and tagged with an identifying pit tag, and were maintained in the laboratory aquarium under ambient conditions. The experimental fish were divided into 3 groups, each of 5 fish: Group 1 was tagged internally with regular Cefas G5 dummy tags (figure 14). Group 2 was tagged externally with Cefas G5 dummy tags. Group 3 fish were not provided with an additional tag so as to act as the control group. Following tagging (under anaesthesia), fish were placed in a 90 x 90 cm arena in 50 l of seawater, and their behaviour was filmed over 23 h. This process was repeated after 6 weeks, then again at 12 weeks. Noldus Ethovision® XT 6.0 is being used to analyze the tagged-fish behaviour. In addition, a second group of larger sole obtained from Aquaculture (n = 57, average wt 250 g approx) was tagged (as above) and monitored for growth and condition. The experiment was completed in June 2010. Results from our study on the impacts of tagging on the growth and behavior of sole were presented at 8IFFS, and are currently being prepared for publication (Hunter et al. 2012b).

6.3 Potential for water chemistry to predict otolith composition in juvenile sole. The deployment of DGT samplers was ultimately not possible as part of this study. However, a 2nd cohort of juvenile sole samples from sites previously sampled by IFREMER were obtained, and their otoliths analysed to obtain elemental fingerprints. This work will ultimately result in a collaborative research paper entitled ―Discrimination of Solea solea nurseries along the French Atlantic coast using otolith elemental signatures‖. The results from this study are relevant to future studies to discriminate sole nurseries in the English Channel and surrounding areas. 6.4 Co-development of “Understanding and predicting connectivity in marine fish populations” Originally co-drafted by IFREMER and Cefas (under MEMFISH), with input from associated European laboratory colleagues (IMARES, KULeuven, FFCUL, NIOZ, UNIABDN, WU & CNRS) the EU Initial Training Network ―Understanding and predicting connectivity in marine fish populations‖ was originally submitted in the 2007 FP7 call, then a much revised version was submitted in 2009. On each occasion the proposal scored well, passing all threshold criteria, but ultimately did not win funding. Similarly, an EU IEF fellowship proposal ―Population structure and connectivity of sole in the North-East Atlantic‖ passed the threshold criteria but was not ultimately funded. Both proposals may, however, be used as the basis of future proposals as and when appropriate opportunities arise.

SUMMARY OF WP3 SCIENTIFIC OUTPUTS & CONTRIBUTIONS Peer-reviewed papers: Hunter et al. 2012b ICES Contributions: ICES WKFLAT 2012 Conferences: ―Eighth International Flatfish Symposium‖, IJmuiden, Netherlands, November 2011. Proposals: ―Population structure and connectivity of sole in the North-East Atlantic‖ (CONSOLE) FP7- PEOPLE-2007-2-1-IEF (not funded) ―Understanding and predicting connectivity in marine fish populations: a methodological approach‖ (FISHCONNECT) FP7- PEOPLE-2009-1-1-ITN (not funded) Supplementary awards: ―Advancing understanding of population structure and connectivity of NE Atlantic sole: Bay of Biscay‖ IFREMER/ Region Poitou Charentes ―Visiting Scientist‖ award (to E. Hunter, September 2009 – June 2010) Other: Hunter & Kupschus 2007, Fishing News

7. DISTRIBUTION AND ABUNDANCE OF SEA BASS IN UK WATERS IN RELATION TO CLIMATE CHANGE (WP4) 7.1 Introduction Bass tolerate a wide range of temperatures (2° to 32°C) and, in British waters, are typically found from 7° to 27°C where they are towards the northern limit of their geographical distribution. Adult bass have been shown to undertake substantial autumn migrations; from the southern North Sea to the western English Channel, from the Irish Sea to Cornwall, and from the English south coast well into the Bay of Biscay. In recent years, the distribution of nursery grounds for bass has shifted north, and juveniles are now able to overwinter in some parts of the North Sea. Under MEMFISH, a desk study was proposed to assess the changing distribution of adult and juvenile bass over the last 30 years, using data from tagging studies, ground-fish surveys, and dedicated bass monitoring studies.

7.2 Bass Distribution Bass landings data (by ICES rectangle) from 1985 - 2010 were extracted from the Fisheries Activity Database (FAD) (Fig.16). These data show an expansion of the bass fishery since 1985, but the centre of distribution of bass based on these data shows no change over the 25 year period. These data don‘t necessarily contradict the current perception of a northward expansion of bass stocks in response to warming sea-temperatures. Rather, they could suggest that the expansion of the fishery is related to the ability of bass to overwinter and breed in areas that have previously been too cold. These data will be presented at the forthcoming ICES WGNEW meeting in March 2012 which will include a data compilation and evaluation exercise for bass, in preparation for a full benchmark stock assessment later in the year.

EVID4 Evidence Project Final Report (Rev. 06/11) Page 22 of 28

Figure 16 Landings of bass, Dicentrarchus labrax, extracted from the Fisheries Activity Database for the years 1985 to 2010. Rectangles indicate ICES rectangles from which bass have been recorded during this period. Black dots represent the annual centres of density, based on annual FAD landings, weighted by catch weight in each rectangle.

7.3 Bass Tagging 7.3.1 English Channel The distribution of European sea bass changes seasonally as they migrate between feeding and spawning grounds, yet their behaviour during these migrations remains largely unexplored. We used DSTs that record and store depth and temperature (originally released under M0154) to identify characteristic behaviours at a fine temporal scale during migrations from and to spawning grounds (Quayle et al. 2009). Eighty-nine sea bass in the North Sea and English Channel were tagged during 2005–2006, 11 of which were recaptured. Five bass showed long distance (>100 km) migration, while the remaining six were recaptured relatively quickly, close to their release locations. Recovered tags yielded 422 days of depth data, from which three characteristic behaviours were identified. The most commonly observed behaviour (― inshore behaviour‖) was characterised by maintenance of position in mid-water with frequent ascents and descents of between 2 and 10 metres. This latter behaviour was often succeeded by ―diving‖, when individuals made deep descents (<120 m) away from surface waters. Bass that remained close to their point of release exhibited ―inshore‖ behaviour for most of their time at liberty, and rarely broke the sequence of inshore behaviour, whereas bass that migrated long distances showed deep-diving and mid-water behaviour for extended periods, with relatively little inshore behaviour. ―Shoaling‖ was also evident in two bass that mirrored their diurnal variability (shallower at night than during the day) for two weeks before separating (Quayle et al. 2009).

7.3.2 Bay of Biscay/English Channel/Celtic Sea WP4 benefited as a result of the IFREMER visiting scientist award to E. Hunter (see WP3, section 6 above). With reference to Cefas‘ previous experience in tagging sea-bass with DSTs (see Quayle et al. 2009), we were asked to collaborate on an IFREMER-led bass-tagging study. To date, this project has tagged and released 145 adult sea-bass with Cefas G5 floated electronic tags to provide data on annual migration routes. The fish were released in the Mer d‘Iroise Marine Park off north-west Brittany. Ten of these tags have already been recovered from the Bay of Biscay, Celtic Sea and English Channel, the most recent of which appears to show indication of an attack by a predator. A further 15 Vemco acoustic tags were used to make detailed observations of individual behaviour of mature fish on the spawning grounds located in the Marine Park. The high levels of mobility of these fish in both English, French and international waters makes this project of relevance to Defra. This project represents a significant added-value to the MEMFISH portfolio, at no significant additional cost.

SUMMARY OF WP4 SCIENTIFIC OUTPUTS & CONTRIBUTIONS Proposals: ―Identifying key population metrics of UK sea bass stocks in relation to climate change‖, Defra 2011 (not funded) ICES contributions: ICES WKNEW, March 2012

EVID4 Evidence Project Final Report (Rev. 06/11) Page 23 of 28 Peer-reviewed papers: Quayle et al. 2009 Other: Hunter & de Pontual 2011, Fishing News, IFREMER collaboration (see above)

8. INTEGRATION

8.1 Applying individual-based telemetry data in fisheries management: A biologically-based movement model for plaice in the North Sea. Recent field studies of commercially exploited marine fish stocks have demonstrated compelling evidence of complex population structure and varying rates of exchange between different management areas. In spite of this, the stock assessment methods currently applied by ICES to advise on total allowable catches (TACs) and technical conservation measures still make only limited concessions to seasonal migrations and movements. As an example, in the North Sea, plaice is currently managed as a single stock. However results from a century of mark-recapture experiments, and more recently, from the release of hundreds of individuals tagged with electronic data storage tags, have allowed the identification of spatial population substructure and the characterisation of annual migration routes and spawning areas. In addition, recent work using otolith microchemistry has been applied to determine the contribution of larvae from different spawning areas to different nursery grounds, and the input from the nursery grounds to the adult population subunits. Hunter et al. (2009) present a biologically-based population movement simulation model (originally developed under M0152) which can utilize data from all of these sources. The reconstructed movements of male and maturing female plaice tagged with DSTs were recalculated (Fig. 5) using ―guided reconstruction‖ performed using the HMM geolocation technique (Pedersen et al. 2008). Juvenile plaice mark-recapture data were summarised spatially (ICES rectangle) and temporally (decadal), including a dislocation and dispersion analysis. The model was further enhanced by the addition of age-structure, and estimations of mean weight for age, sex and time of year making it fully compatible with ICES assessment methodology. The effects of a range of area and seasonal closures were tested in terms of addition to NSPMS of age-structure, and estimations of mean weight for age, sex and time of year. reduction in overall fishing mortality.

8.2 Using DST data to assess changes in the accessibility of Atlantic cod to trawl gears The accessibility of fish to fishing gear is an important component of their ‖catchability‖. We assessed the seasonal and inter-individual variation in accessibility to trawl gear of Atlantic cod (Gadus morhua) in the North Sea by using depth data retrieved from 45 archival tags (~9000 days‘data). To do this, we calculated the proportion of time that cod would have been within 5 m of the seabed (i.e. vulnerable to otter or beam trawls) each day, and used this as an index of catchability. Overall, cod were vulnerable to trawl gear for 74% of the time. Cod were less accessible to trawl when they exhibited migratory behaviour, which generally occurred between October and March. During the feeding season (April to October), cod spent approximately 90% of their time within the reach of trawl gear. Landings (and cpue) data from cod fisheries in the UK between the 1970s and the present day demonstrate that cod landings have typically been greater in summer, although the strength of this pattern has weakened each decade. Our analysis suggests that data from archival tags can be used to assess the ―catchability‖ of cod, and that seasonal changes in catchability are attributable to the interaction between fishing fleet and migratory activity. These data were originally presented at ICES ASC 2009 (Righton et al. 2009).

8.3 Spatio-temporal dynamics of Atlantic cod in western Waters Using data from mark recovery and electronic tag studies, the exchange of cod between three adjacent cod management areas (the Irish Sea (ICES Division VIIa), Scottish waters (ICES Division IVa) and the western English Channel and Celtic Sea, ICES Divisions VIIe-k) was examined. We evaluate 2209 archived and current mark recovery data (1960‘s – to 2010) to assess the seasonal changes in distribution of cod within ICES Divisions VIa, VIIa and VIIe-g. In addition, we report on the results of a collaborative electronic data storage tagging (DST) programme of cod in the Irish and Celtic Sea, spanning the years 1999 to 2010. During this time, 710 DSTs have been deployed on cod between 43cm and 110cm, and 107 of the tags have been returned yielding over 11000 days of data. We use a tidal geolocation model to determine the daily locations of each cod to reconstruct in detail the migrations of cod tagged with DSTs. The results describe fundamental features of cod spatial ecology in the Irish and Celtic Sea, such as the seasonality of migration and habitat occupation and the potential impact upon sub-stock structure. This work has been put forward for ICES benchmarking of cod stocks in the Irish Sea, and is currently being prepared for publication (Bendall et al. 2012).

8.4 Are spatial closures better than size limits for halting the decline of the North Sea thornback ray? A key challenge of the ecosystem approach to fisheries management is to sustain viable populations of large bodied less-productive vulnerable elasmobranchs that are the by-catch of fisheries that target more productive species. The North Sea population of the thornback ray (Raja clavata) is now mainly confined to the Thames Estuary and surrounding SW North Sea and is exploited by a flatfish trawl fishery. Using data originally collected under M0148, we explored the relative effectiveness of seasonal closures versus size-based landing restrictions

EVID4 Evidence Project Final Report (Rev. 06/11) Page 24 of 28 using a four-season age-structured model (Wiegand et al. 2011). More than a third of adult thornback rays are currently removed by fishing each year, and without effective management, a further 90% decline within 30 years is likely. Model results indicate that a three-season closure of the Thames Estuary is the shortest closure that will ensure thornback ray recovery and minimal loss of fishery yield. Minimum and maximum landing size restrictions are nearly as effective at recovering thornback rays but less so at improving yield. While long seasonal closures and full marine protected areas are more effective at ensuring the recovery of thornback rays, length restrictions may be simpler to implement under the current institutional framework and may have less impact on the multispecies trawl fisheries operating in the area.

9. FUTURE WORK

Applying biomineralised materials in climate change research Cefas holds an internationally recognised repository of otoliths and other biomineralised materials (scales, vertebrae, etc…) dating back at least 6 decades. Work is currently underway to comprehensively catalogue this material, and to scope how these materials might best be deployed in research to address ―big science‖ questions, for example, in identifiying the impact of inter-decadal climate change and consequent impacts on fish populations. Methods developed under MEMFISH WP1 will play an important role in taking this work forward.

Migration of European seabass As a result of the IFREMER visiting scientist award to E. Hunter (see WP3, above), and due to Cefas‘ previous experience in tagging sea-bass with DSTs (Quayle et al. 2009), we were asked to collaborate on an IFREMER- led sea-bass-tagging study. To date, this project has tagged and released 145 adult sea-bass (paid for by IFREMER) with Cefas G5 floated electronic tags to provide data on annual migration routes. The high levels of mobility of these fish in both English, French and international territorial waters makes this project of relevance to Defra. We are currently seeking external funding to allow analysis of data already gathered, and to extend the release of tagged bass at sites in the Irish Sea and English Channel.

Energetics of Atlantic cod Fieldwork undertaken under MEMFISH using research vessels and electronic tagging studies have yielded some new avenues for research. Firstly, MEMFISH results have suggested that the reproductive behaviour and reproductive potential of cod appear to be closely linked to environmental temperature. Existing collaborations with IMR Norway will be developed to exploit the potential for further studies on these relationships. Secondly, a greater understanding of the spatial dynamics of cod, and how this relates to overall energetics and energy capture, is beginning to emerge. These give rise to the potential for innovative studies of migration energetics and food acquisition, which we anticipate being taken forward in collaboration with potential collaborators in the UK, Ireland and Denmark.

Significance of predation on early life-history stages in stock-recruitment In the Irish Sea, a steady increase in spawning stock biomass of plaice since the late 1990s, due to reduced fishing pressure, has yet to be matched with a clear improvement in recruitment. Analysis of stomach content data from predators sampled during PREDATE (MF0432) and MEMFISH cruises (Fox et al. 2012) suggest that suppressed plaice recruitment may be at least partially due to high rates of predation on the eggs. In Liverpool Bay, based on the analysis of more than 400 sprat stomachs combined with simultaneous observations of plaice egg density in the water column, we estimate that sprat may consume between 20% and 76% of all spawned plaice eggs (Pliru et al. 2012). These data suggest a possible ‗top-down‘ effect on the population dynamics of the bigger fish species based on predation on fish eggs by the smaller forage fish. As evidence suggests that sprat abundance is increasing in the Irish Sea, egg predation may become an increasingly constraining factor on stock dynamics in the future.

Field work carried out under PREDATE (and followed up during MEMFISH) was limited to just two 10 day surveys, one of which was negatively impacted by adverse weather conditions. The work undertaken therefore represents ―proof of concept‖, where we have demonstrated our ability to model egg dispersion in real-time, to sample and map predator communities, and to describe predator-prey dynamics. Given the failure of the western Irish Sea cod stock to recover following 10 years of protection and a recovery plan, a next logical step would be to investigate whether the same species predate the early-life history stages of Irish Sea (and North Sea) cod, so as to determine whether this is a significant factor in cod stock recovery. Since early stage cod eggs cannot be identified visually, the use of the molecular approach pioneered in the former study will be essential.

EVID4 Evidence Project Final Report (Rev. 06/11) Page 25 of 28

References to published material 9. This section should be used to record links (hypertext links where possible) or references to other published material generated by, or relating to this project. 10. LIST OF PUBLICATIONS

Albaina, A, Fox, C.J., Taylor, N., Hunter, E., Maillard, M. & Taylor, M.I. (2010). A TaqMan real-time PCR based assay to detect predation of plaice (Pleuronectes platessa L.) DNA by the brown shrimp (Crangon crangon L.) and the shore crab (Carcinus maenas L.) – Assay development and validation. Journal of Experimental Marine Biology and Ecology 391: 178-189.

Apostolaki, P., Pilling, G.M., Armstrong, M.J., Metcalfe, J.D. & Forster, R. (2008). Accumulation of new knowledge and advances in fishery management; two complementary processes? In: ―Advances in Fisheries Science: 50 years on from Beverton and Holt‖. pp 229-254 Eds. A. Payne, J. Cotter & E. Potter, Blackwell Publishing.

Bendall, V., Macdara, O‘Cuaig, Schön, P.J., Hetherington, S., Armstrong, M., Graham, N. & Righton, D. (2009). Spatio-temporal dynamics of Atlantic cod (Gadus morhua) in the Irish and Celtic Sea: results from a collaborative tagging programme. ICES CM2009/ J0:4, 35 pp.

Bendall, V., Ơ Cuaig, M., Schön, P.J Hetherington, S., Armstrong, M., Graham, N. & Righton, D. (2012a). Spatio-temporal dynamics of Atlantic cod (Gadus morhua) in the Irish and Celtic Sea: Latest results from a collaborative tagging programme (as of January 2012). Paper for the ICES Benchmark Workshop on Western Waters Roundfish in Aberdeen.

Bendall, V., Ơ Cuaig, M., Schön, P.J Hetherington, S., Armstrong, M., Graham, N. & Righton, D. (2012b). Spatio-temporal dynamics of Atlantic cod (Gadus morhua) in Western Waters. Marine Ecology Progress Series In prep.

Fox, C.J., Taylor, M., van der Kooij, J., N. Taylor, Pliuru, A., Milligan, S.M., Sonia Pascoal, S., Lallias, D. Maillard, M. & Hunter, E. (2012). Predators of plaice eggs in the eastern Irish Sea identified using molecular probes. Marine Ecology Progress Series In press.

Heath, M.R., Preedy, K.F., Culling, M.A., Crozier, W.W., Fox, C.J., Gurney, W.S.C., Hutchinson, W.F., Nielsen, E.E., O‘Sullivan, M., Righton, D.A., Speirs, D.C., Taylor, M.I., Wright, P.J. & and Carvalho G.R. (2012). Genetic diversity in fish populations is threatened by harvesting patterns. Proceedings of the National Academy of Sciences Submitted.

Hunter, E., Buckley, A., Darnaude, A & Bell, M. (2009). Applying individual-based telemetry data in fisheries management: A biologically-based movement model for plaice in the North Sea. ICESCM 2009/J:04, 26 pp.

Hunter, E., Cotton, R.J., Metcalfe, J.D. & Reynolds, J.D. (2009). Spatial and temporal variation in swimming activity and swimming patterns by plaice, Pleuronectes platessa L., in the North Sea. Marine Ecology Progress Series 392: 167-178.

Hunter, E. and de Pontual, H. (2011). ―Sea bass on the move: Help needed from Fishermen‖ Fishing News, 28 Jan 2011

Hunter, E. & Kupschus, S. (2007). Channel sole targeted with electronic tags. Fishing News

Hunter, E., Péan, S., & Bégout, M.L. (2012b). Tagging effects on growth, condition and behaviour of the common sole, Solea solea L. Journal of Sea Research In prep.

Hunter, E., Taylor, N., Fox, C.J., Maillard, M. & Taylor, M.I. (2012a). Effectiveness of TaqMan probes for detection of fish eggs and larvae in the stomach contents of a teleost predator. Journal of Fish Biology. In press.

Kjesbu, O.S., Righton, D., Krüger-Johnsen, M., Thorsen, A., Michalsen, K., Fonn, M. & Witthames, P.R. (2010). Thermal dynamics of ovarian maturation in Atlantic cod (Gadus morhua). Canadian Journal of Fisheries and Aquaculture Science. 67: 605-625.

Metcalfe, J.D., Fox, C.F., Righton, D.A., Wright, P.J. & Casey, J. (2009). A review of the biological evidence for cod stock sub- structure in the North Sea: a report for the ICES roundfish benchmarking meeting in Copenhagen.

Metcalfe, J.D., Righton, D.A., Hunter, E. & Eastwood, P. (2008). Migration and Habitat Choice in Marine Fish. In: ―Fish Behaviour‖ Eds. C. Magnhagen, V.A. Braithwaite, E. Forsgren & B.G. Kapoor. Science Publishers Inc., Enfield, USA. Pp 187-233.

Metcalfe, J.D., Righton, D.A., Hunter, E., Neville, S. & Mills, D.K. (2008). New technologies for the advancement of fisheries science. In: ―Advances in Fisheries Science: 50 years on from Beverton and Holt‖. pp 255-279. Eds. A. Payne, J. Cotter & E. Potter, Blackwell Publishing.

Neuenfeldt, S., Righton, D., Neat, F., Wright, P.J., Svedäng, H., Michalsen, K., Subbey, S., Steingrund, P., Thorsteinssson, V., Pampoulie, C., Andersen, K.H., Pedersen, M.W., & Metcalfe, J.D. (2012). Migrations of Atlantic cod in the NE Atlantic: then, now and the future. Journal of Fish Biology In press

Pliuru, A., van der Kooij, J., Fox, C.J., Milligan, S.M., & Hunter, E. (2011). Feeding behaviour, selective predation and daily egg consumption rates by sprat. ICES Journal of Marine Science In press.

Quayle, V., Righton, D., Hetherington, S. & Pickett, G. (2009). Observations of the Behaviour of European Sea Bass (Dicentrarchus labrax) in the North Sea. in, J.L. Nielsen et al. (eds.), Tagging and Tracking of Marine Animals with Electronic Devices, Reviews: Methods and Technologies in Fish Biology and Fisheries 9, 103-119.

Righton, D.A., Andersen, K.H., Neat, F., Thorsteinsson, V., Steingrund, P., Svedäng, H., Michalsen, K., Hinrichsen, H.H., Bendall, V., Neuenfeldt, S., Wright, P., Jonsson, P., Huse, G., van der Kooij, J., Mosegaard, H., Hüssy, K. & Metcalfe, J. (2010). Thermal niche of Atlantic cod Gadus morhua: limits, tolerance and optima. Marine Ecology Progress Series. 420: 1-13.

Righton, D., & Metcalfe, J.D. (2012). Through the looking glass:how archival; tagging programmes can provide fundamental biological information and be useful for fisheries management. Proceedings of the Second International Conference on Fish Tagging

EVID4 Evidence Project Final Report (Rev. 06/11) Page 26 of 28 and Tracking. In press.

Righton, D., Neat, F., Bendall, V., Berx, B., Wright, P & Metcalfe, J.D. (2012). Spatial dynamics of cod in UK waters. PLoS One, in prep.

Righton, D., Quayle, V., Neat, F., Pedersen, M., Wright, P., Armstrong, M., Svedäng, H., Hobson, V. & Metcalfe, J. (2008). Spatial dynamics of Atlantic cod (Gadus morhua) in the North Sea: results from a large-scale electronic tagging programme. ICES CM 2008/P:04.

Righton, D., Townhill, B. & van der Kooij, J. (2009). Catch me if you can: archival tagging studies can help assess changes in the accessibility of Atlantic cod (Gadus morhua) to trawl gears. ICES CM2009/ J0:8

Sims, D.W., Southall, E.J., Humphries, N.J., Hays, G.C., Bradshaw, C.J.A., Pitchford, J.W., Mohammed, A.J., Ahmed, Z., Brierley, A.S., Hindell, M.A., Morritt, D., Musyl, M.K., Righton, D., Shepard, E. L.C., Wearmouth, V.J., Wilson, R.P., Witt, M.J. & Metcalfe, J.D. (2008). General scaling laws of marine predator search behaviour. Nature. 451: 1098-1101.

Sturrock, A.M., Trueman, C.T., Darnaude, A.M. & Hunter, E. (2012). Can otolith microchemistry track individual movements of fully marine fish? Journal of Fish Biology. In press.

Sturrock, A.M., Hunter, E., Milton, A. &Trueman, C.T. (2012) Optimisation of a simple dilution method for measuring trace metals in fish blood plasma using HR-ICPMS. Journal of Applied Spectrometry Submitted.

Sturrock, A.M., Hunter, E., & Trueman, C.T. (2012). Intrinsic and extrinsic influences on multi-element chemistry in the blood of a marine flatfish Canadian Journal of Fisheries & Aquatic Science In prep.

Sturrock, A.M., Hunter, E., & Trueman, C.T. (2012). Disentangling intrinsic and extrinsic influences on otolith microchemistry in a fully marine, adult flatfish: an experimental approach – Marine Ecology Progress Series In prep.

West, C., Pitchford, J., Righton, D., Dytham, C. (2012). Effects of climate change on reproductive aggregations: a model and application to cod in the North Sea. Journal of Animal Ecology Submitted

Wiegand, J., Hunter, E. & Dulvy, N. (2011). Evaluating management strategies for the thornback ray Raja clavata: Are marine protected areas better than traditional fisheries management? Marine and Freshwater Research 62: 722-733.

11. ADDITIONAL REFERENCES CITED IN THE REPORT

Blanchard, J. L., Heffernan, O. A. & Fox, C. J. North Sea cod, International Council for the Exploration of the Seas, ICES Cooperative Research Report, 274, 76-88 pp., (2005).

Fromentin, J.-M., Ernande, B., Fablet, R. & de Pontual, H. (2009). Importance and future of individual markers for the ecosystem approach to fisheries. Aquatic Living Resources 22, 395-408.

Hunter, E., Berry, F., Buckley, A.A., Stewart, C. & Metcalfe, J.D. (2006). Seasonal migration of thornback rays and implications for closure management. Journal of Applied Ecology 43: 710-720.

Kell, L.T., Scott, R. & Hunter, E. 2004. Implications for current management advice for North Sea plaice: Part I. Migration between the North Sea and English Channel. Journal of Sea Research 52: 287-299.

Pedersen, M.W., Righton, D., Thygesen, U.H., Andersen, K.H. & Madsen, H., 2008. Geolocation of North Sea cod (Gadus morhua) using hidden Markov models and behavioural switching. Canadian Journal of Fisheries and Aquatic Science 65: 2367–2377.

12. LIST OF PRESENTATIONS

Sturrock, A., Trueman, C. & Hunter, E. (2012) Validation of otolith chemistry as an indicator of movement, connectivity and stock structure in North Sea plaice. World Fisheries Congress, Edinburgh, UK, May 2012.

Hunter, E., Stewart, C., Riley, A. & Metcalfe, J.D. (2011) Growing with the flow: new observations on the behaviour of male and maturing female plaice in the North Sea. Eighth International Flatfish Symposium 2011, Ijmuiden, The Netherlands.

Hunter, E., Péan, S. & Bégout, M.L. (2011) Impacts of tagging on the common sole, Solea solea. Eighth International Flatfish Symposium 2011, Ijmuiden, The Netherlands.

Sturrock, A.M., Trueman, C., Darnaude, A.M. & Hunter, E. (2011) There‘s no plaice like home: Using data storage tags to validate otolith chemistry in North Sea plaice. Eighth International Flatfish Symposium 2011, Ijmuiden, The Netherlands.

Sturrock, A.M., Trueman, C. & Hunter, E. (2011) Disentangling environmental and physiological influences on otolith chemistry using a flatfish model. Eighth International Flatfish Symposium 2011, Ijmuiden, The Netherlands.

Plirú, A., van der Kooij, J., Engelhard, G.H., Fox, C.J., Milligan, S.P. & Hunter, E. (2011) Egg predation by sprat may impose a ―top- down‖ effect on plaice recruitment in the Irish Sea. Eighth International Flatfish Symposium 2011, Ijmuiden, The Netherlands.

Plirú, A., van der Kooij, J., Engelhard, G.H., Fox, C.J., Milligan, S.P. & Hunter, E. (2011) Is recruitment of plaice in the Irish Sea constrained by predation of eggs by sprat? ICES Annual Science Conference 2011, Theme Session H

Lewis, A.M., Trueman, C.N., Darnaude, A. & Hunter, E. (2011) Unlocking the bio-logging potential of otoliths as natural tags: Disentangling environmental and physiological influences on otolith chemistry. ―Fourth International Science Symposium on Bio- logging‖, Hobart, Australia.

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Lewis, A.M., Trueman, C.N. & Hunter, E. (2010) Disentangling environmental and physiological influences on otolith chemistry: unlocking the potential use of otoliths as natural tags. ―American Fisheries Society Symposium‖, Pittsburgh, Pennsylvania USA.

Fox, C.J., Taylor, M., Taylor, N., van der Kooij, J., Milligan S. and Hunter, E. (2010) Describing the dynamics of predation in a changing predator landscape. FSBI Annual Symposium ―Fish and Climate Change‖, Belfast, UK.

Lewis, A.M., Trueman, C.N. & Hunter, E. (2010) Can otolith chemistry interpret spatial dynamics in a changing environment? An experimental approach. FSBI Annual Symposium ―Fish and Climate Change‖, Belfast, UK.

Bendall, V., Macdara, O‘Cuaig, Schön, P.J., Hetherington, S., Armstrong, M., Graham, N. & Righton, D. (2009) Spatio-temporal dynamics of Atlantic cod (Gadus morhua) in the Irish and Celtic Sea: results from a collaborative tagging programme. ―ICES Annual Science Conference‖, Berlin.

Hunter, E., Buckley, A., Darnaude, A & Bell, M. (2009) Applying individual-based telemetry data in fisheries management: A biologically-based movement model for plaice in the North Sea. ―ICES Annual Science Conference”, Berlin.

Hunter, E. (2009) Finding one‘s plaice: Individual behaviour and it‘s application in fisheries management. IFREMER visiting scientist lecture (L‘ Houmeau, Brest, Nantes).

Lewis, A., Darnaude, A., Hunter, E. & Trueman, C. (2009) Development and validation of otolith microchemistry techniques in support of the retrospective geolocation of fully marine fishes. ―4th International Otolith Symposium”, Monterey, California, USA.

Lewis, A., Hunter, E. & Trueman, C. Investigation into the influence of spawning on the chemistry of plaice (Pleuronectes platessa L.) otoliths. ―4th International Otolith Symposium”, Monterey, California, USA (poster)

Darnaude, A. & Hunter, E. (2008) Determination of Plaice lifetime movements in the North Sea by linking natural and electronic data records. ―Society for Experimental Biology”, Satellite Meeting, Sete, France.

Metcalfe, J. & Righton, D. (2008) ―Reaching for the stars: a perspective from 25 years of fish tagging‖ (invited key-note paper). ―Third International Bio-logging Science Symposium‖, Pacific grove, California, USA.

Darnaude, A. & Hunter, E. (2008) Coupled analysis of natural and electronic data records to determine the lifetime movements of individual fish. ―Advances in fish tagging and marking technology‖, Auckland, New Zealand.

Hunter, E., Buckley, A., Neville, S., Darnaude, A. & Bell, M. (2007) Applying telemetry data in fisheries management: A biologically- based movement model for plaice in the North Sea. ―Seventh Conference on Fish Telemetry held in Europe‖, Silkeborg, Denmark.

Darnaude, A. & Hunter, E. (2007) Otolith δ18O and data storage tags: key tools in the assessment of marine fish stock structure and lifetime migration patterns? ―Seventh Conference on Fish Telemetry held in Europe‖, Silkeborg, Denmark.

Quayle, V.A & Righton, D. (2007) Preliminary observations of the behaviour of sea bass (Dicentrarchus labrax) during migration. ―Seventh Conference on Fish Telemetry held in Europe‖, Silkeborg, Denmark.

Darnaude, A. & Hunter, E. (2007) Migration history and stock discrimination in North Sea plaice through coupled analysis of otolith δ18O and archival tag data. ―Tagging and tracking marine fish with Electronic Devices‖, San Sebastian, Spain.

Hunter, E. (2007) Cefas research into thornback ray migration. Presentation to Kent & Essex Sea Fisheries Committee.

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