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Evidence Project Final Report

NoteIn line with the Freedom of Information Act 2000, Defra aims to place the results of its completed research projects in the public domain wherever possible. The Evidence Project Final Report is designed to capture the information on the results and outputs of Defra-funded research in a format that is easily publishable through the Defra websiteAn Evidence Project Final Report must be completed for all projects.

This form is in Word format and the boxes may be expanded, as appropriate.

ACCESS TO INFORMATIONThe information collected on this form will be stored electronically and may be sent to any part of Defra, or to individual researchers or organisations outside Defra for the purposes of reviewing the project. Defra may also disclose the information to any outside organisation acting as an agent authorised by Defra to process final research reports on its behalf. Defra intends to publish this form on its website, unless there are strong reasons not to, which fully comply with exemptions under the Environmental Information Regulations or the Freedom of Information Act 2000.Defra may be required to release information, including personal data and commercial information, on request under the Environmental Information Regulations or the Freedom of Information Act 2000. However, Defra will not permit any unwarranted breach of confidentiality or act in contravention of its obligations under the Data Protection Act 1998. Defra or its appointed agents may use the name, address or other details on your form to contact you in connection with occasional customer research aimed at improving the processes through which Defra works with its contractors.

Project identification

1. Defra Project code   MF1105   

2. Project title

The Continuous Plankton Recorder Survey: Fisheries Investigations

3. Contractororganisation(s)

Dr Priscilla LicandroSir Alistair Hardy Foundation for Ocean Science     The Laboratory - Citadel Hill     PL1 2PB, Plymouth

54. Total Defra project costs £ 978,065(agreed fixed price)

5. Project: start date................. 01 April 2007

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end date.................. 30 March 2012

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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 Summary7. 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.

     The Continuous Plankton Recorder (CPR) survey, run by the Sir Alister Hardy Foundation for Ocean Science (SAHFOS), is the longest and geographically the most extensive marine biological survey in the world.

During 2007-2012, the Defra contract MF1105 has contributed to the maintenance and expansion of the core-monitoring programme of the CPR survey and to the associated work that has a fisheries focus. The operational management of the survey is quite complex, including logistics (the survey is globally operated by commercial vessels), workshop maintenance, plankton analysis and archiving of data and samples, possible application of the resulting data to fisheries research and the potential impact of climate change on fisheries. The Survey relies on a team of scientists, technicians and on other personnel that have many decades of experience and knowledge, including a taxonomic skill base on phytoplankton, zooplankton and fish larvae. New CPR routes have been established since 2010 in the northern North Atlantic in the proximity of the Arctic ocean, while since 2011 a CPR monitoring has been started across the Benguela current, from Angola to South Africa, by colleagues from the University of Cape Town with the support of SAHFOS. The information on plankton biomass and diversity provided by the CPR in those new regions will improve our understanding of the impact of environmental variability on the productivity of fish stocks and will help to Recent investigations on long-term changes of plankton functional groups based on CPR records have shown that some areas around the British Isles (e.g. the Flamborough Head region and the Skagerrak) are particularly sensitive to climate and to other environmental changes. In some of those regions for instance, marine plankton predators such as cnidarian jellyfish, which are competing with fish for prey, have increased in abundance and are now more persistent than in the past. It is therefore important that in those marine habitats potentially at risk, fish stocks are exploited more lightly than elsewhere. Within the MF1105 SAHFOS has generated a new index of zooplankton standing stock that is currently used by different modelling groups to validate end-to-end regional models for the North Sea and the north-western European shelf. SAHFOS is directly involved in this validation process, which is a complex but necessary task to select the most suitable regional models to be used as tools for the management of the marine resources. In the future, a yearly update of the zooplankton index could be used to assess the performance of those ecosystem models used for management purpose.

Another option for new work following up the MF1105 has arisen from the successful application of a

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rapid genetic technique for the identification of species of clupeid larvae collected by the CPR. This new application, developed by Dr Andrew Griffiths in collaboration with SAHFOS, open up to the possibility of obtaining CPR records of early stages of herring, sardine and sprat that could be utilised to improve the assessment and the management of those fish stocks.

Project Report to Defra8. 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).

1. To monitor and describe, in the long-term, pelagic variability and diversity in the core areas of the CPR survey over the Northwest European shelf and in the eastern and western Atlantic with a focus on applications to fisheries research and the interpretation of climate change effects on fisheries.

LogisticSAHFOS has successfully maintained and expanded the core-monitoring programme of the CPR survey and associated work that has fisheries focus. CPRs have sampled along 23 different routes on average circa ≥ 120, 000 nautical miles (nm), providing more than 100,000 analysable samples per year in the core North Atlantic area. Since 2009 the CPR monitoring has been expanded in the northernmost region of the North East Atlantic, due to the re-establishment of the ST route between Svalbard and the North Cape of Norway and thanks to the addition of a new route (NI route) between Mosjoen (Norway) and Reydarfjordur (Iceland). The plankton collected along those routes provides unique information to monitor the changes occurring in the pelagic ecosystems of the Arctic and Subarctic, two regions that have undergone significant environmental changes due to global warming. Since June 2010 CPRs are towed also in the Atlantic sector of the Antarctic Ocean between the Falkland Islands and South Georgia. Since July 2011 a regular CPR monitoring (CT route) has been implemented with the support of SAHFOS in the Benguela region, from Luanda (Angola) to Durban (South Africa).

Highlights SAHFOS has set up the framework for fostering, coordinating and harmonizing the implementation of a worldwide CPR monitoring network, i.e. the Global Alliance of Continuous Plankton Recorder Surveys (GACS, http://www.globalcpr.org/). A global CPR monitoring may significantly enhance our understanding of the ongoing changes in the marine environment as pelagic variability and related changes in the fish stocks need to be addressed not only at local, but also at regional and global scale.

ResearchDuring the 4-years of the MF1105 SAHFOS has published 37 publications relevant to the MF1105 in peer-reviewed journals. There is increasing evidence that in the last few decades the functioning of the North Atlantic has undergone significant changes, which may have had a knock on effect on the North Sea and NW European shelf fisheries. Some of the most relevant findings achieved by SAHFOS within the MF1105 have shown that: (1) increasing abundance and species shift in dominant meroplankton (i.e. plankton larvae of benthic organisms) induced by climate changes may have altered the trophodynamics of the North Sea ecosystem (Kirby et al., 2007; Lindley et al., 2010); (2) Significant ecosystem changes across multiple trophic levels (i.e. the so called ’regime shifts’) have been triggered in the North Atlantic by temperature increases above a critical thermal boundary of 9-10 °C (Beaugrand et al., 2008); (3) alteration in the

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plankton community could be related to the recent poor recruitment of North Sea autumn-spawning herring (Payne et al., 2009); (4) marine plankton predators such as jellyfish, competing with fish for prey, have recently increased or showed alterations in their geographical distribution in the north-eastern Atlantic and adjacent seas, mainly due to hydro-climatic variability (Licandro et al., 2010; Licandro et al., 2011); (5) changes in the trophic regulation of the North Sea plankton food-web have occurred after the early 1970s (Llope et al., 2011).The main findings summarised above have been achieved applying different statistical methodologies accounting for the spatio-temporal autocorrelation that characterises CPR data and other large data sets.

Highlights Some statistical methodologies developed within the MG1105, i.e. threshold Generalized Additive Models (GAMs), have shown to be useful tools to identify critical changes in the spatio-temporal variability of different plankton functional groups (see Fig. 1 below as an example). This methodology could be applied to other biotic (e.g. ICES fish data) and abiotic (e.g. sea surface temperature) data sets available in the North Sea and adjacent seas, to verify whether the variability observed in some fish stocks could be related to a shift in the regulation of food-web dynamics.

Figure 1 (from Llope, Licandro et al., 2012). Phytoplankton biomass model. Zoo/phytoplankton spatio-temporal interaction. (A) Contour map showing the characteristic distribution of zooplankton filter-feeders biomass (HER) observed in the North Sea after 1972. Since 1972 spatial effects of diatoms (B) and dinoflagellates (C) on filter-feeders differ in the western and eastern North Sea regions. (D) Threshold selection.

2. To provide an oversight of monitoring results from the independently funded North Pacific CPR survey and extend the coverage of the survey outside the core area as and when additional funding is available.

More than 10,000 nm have been regularly sampled every year in the North Pacific along the AT (Anchorage-Tacoma) and VJ (Vancouver-Japan) routes. During the four years of the MF1105 project, 5 papers based on the North Pacific CPR monitoring have been published in peer-reviewed journals.

The CPR monitoring in the North-eastern Pacific has shown that different hydroclimatic conditions are usually associated with different plankton communities. Phase changes in the dominant climatic signal, i.e. the Pacific Decadal Oscillation, are reflected by the alternation of relatively cool (2000-2002 and from 2007 onwards) and very warm (2004-2006) years, characterised in turn by a zooplankton community usually dominated by relatively large copepods in cool years and by small, typically more southern

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copepod species, in warmer years. Since 2010 the analysis of CPR samples collected along the VJ route in the North-western Pacific (sector <170°E) has been undertaken by JAMSTEC (Japan), under the coordination and responsibility of Sanae Chiba.

3. To collaborate with UK fishery agencies (Cefas, FRS, DARD), NERC laboratories and associated organisations and especially modelling groups outside SAHFOS to increase understanding of links between CPR data and long-term changes in fish stocks with the aim of developing a predictive capability.

SAHFOS has been proactive in developing collaborations with fishery agencies inside and outside the UK, including CEFAS, IFREMER (France), Tecnalia-AZTI and CSIC-ICM (Spain), DTU-AQUA (Denmark), vTI-SF (Germany), IMARES (the Netherlands), MIR (Poland), IMR (Norway), MRI-HAFRO (Iceland). In collaboration with those agencies within the EuFP7 projects BASIN and VECTORS (both projects started in 2011), SAHFOS is using CPR data to increase our understanding of the effects of environmental variability on long-term changes in North Sea fish stocks. As a contribution to develop a predictive capability and a better management of the marine environment, SAHFOS has produced a new index of zooplankton standing stock that account not only for the size by also for the composition of the zooplankton community. This is not trivial as zooplankton organisms of the same size may be characterized by different stochiometries and then account for a very different biomass (Fig. 2).

Figure 2 (from SAHFOS Report 2012). Mass (dry weight) per prosome unit length (mm) for different genera of dominant copepods.

HighlightsThe index of zooplankton standing stock produced by SAHFOS is currently utilised to validate end-to-end regional models developed within the Eu-FP7 MEECE (more information are available at http://www.sahfos.ac.uk/data-archive/cpr-data-for-model-validation.aspx). Those models (i.e. POLCOSM/ERSEM, ECOSMO, ECOSIM-ECOPATH) are intended to be used in the future to improve the predictive capability of fish stocks fluctuations in the North Sea and on the NW European shelf. SAHFOS zooplankton biomass index will also be included as a new environmental descriptor for the Skagerrak ecosystem assessment and used for validation of the coupled physical-biological shelf seas ecosystem model run by the National Centre for Ocean Forecasting at the UK MetOffice (ERSEM/NEMO model).

4. To increase the use of the CPR database through involvement with external researchers/organisations outside the UK and especially within ICES working groups and in collaboration with OSPAR.From March 2007 to March 2012 SAHFOS has received more than 340 data requests from all over the world. SAHFOS has participated and lead in different ICES documents, including 3 ICES Cooperative Research Reports, the annual reports of the ICES WG on Zooplankton Ecology (WGZE), the report of the ICES Study Group on Recruitment Variability in North Sea Planktivorous Fish (SGRECVAP) and the report of the Study Group on Working Hypotheses Regarding effects of Climate Change (SGWRECC). SAHFOS was also leading author of the chapter on trends in plankton communities included in the ICES

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Position Paper on climate change (Licandro et al., 2011).CPR data are used every year for the implementation of an integrated assessment of the North Sea ecosystem, carried out by the ICES Working Group WGINOSE.

HighlightsIn May 2007 SAHFOS has hosted the 2nd meeting of the ICES Study Group on Recruitment Variability in North Sea Planktivorous Fish (SGRECVAP). A close collaboration between SAHFOS and the ICES fishery community has been developed throughout the years. As a result, Dr. Priscilla Licandro from SAHFOS is now co-chair of the newly formed ICES Working Group on Small Pelagic Fishes, their Ecosystems and Climate Impact (WGSPEC).

5. To secure the long-term maintenance of the sample archive and maximise its use by making it available for molecular, biogeochemical and other studies, including work on fish larvae.

SAHFOS has regularly revisited and ensured a good quality of the state of preservation of the samples in the CPR archive, which contains more than 2500 sealed storage boxes holding an estimated 176,000 samples in labelled plastic envelopes on a bed of formaldehyde impregnated cotton wool. Molecular biology has been used to study different plankton organisms collected from CPR samples such as non-indigenous (i.e. the diatom Neodenticula seminae) and HaBs (i.e. the dinoflagellate Karenia spp.) phytoplankton species, in collaboration with Dr. Declan Schroeder at the MBA. Zooplankton taxa not routinely identified by morphological recognition such as cnidarian jellyfish and decapod larvae were also studied by applying molecular techniques to CPR samples, in collaboration with Dr. Richard Kirby of the University of Plymouth.

HighlightsDr. Andrew Griffiths at the MBA has developed a rapid genetic technique, involving real-time PCR and high-resolution melt (HMR) analysis, which has successfully identified species of clupeid larvae sampled by the CPR. This methodology could be utilised to retrospectively identify and count the larvae of herring, sardine and sprat collected by the CPR in the seas around UK.

6. To add value to the CPR survey by instrumenting routes when possible and affordable and by implementing relevant new methodologies, in partnership with other organisations.

SAHFOS has regularly measured water temperature and salinity using different sensors (i.e. Brancker XR620 CTD+F unit, Minipack CTD+F and Star Oddi, CTD units) in 2 trans-Atlantic routes (BB/BA and DA/D), 2 routes across the Bay of Biscay (IB and SA) and on the Plymouth to Roscoff route (PR). Temperature and salinity were also recorded on the CPR tow from Immingham to Iceland (LR/V-) and along 2 routes in the Norwegian Sea from Norway to Iceland (NI) and from Norway to Svalbard (ST).In addition temperature and salinity were measured on the 3 trans-Pacific CPR routes (VJ) and temperature on the Canadian west-coast tow (AT).A ‘Water and Microplankton Sampler (WaMS)’, set up to collect 10 water samples, was designed and fitted to a standard Continuous Plankton Recorder body to allow the collection in continuous of surface sea-water samples. WaMS were towed in the southern North Sea along the R route and in the Western English Channel along the PR route. Water samples collected by the WaMS, approximately 100ml of sea-water preserved in acidic Lugol’s iodine, were used to count the smallest component of the plankton (i.e. micro, nano- and picoplankton) through counting by inverted microscopy, flow cytometry and molecular biology. HighlightsMonitoring by the WaMS could provide unique information at regional scale on small eukaryotes cells including Harmful Algal Bloom (HABs) taxa, bacteria, viruses and potential plankton pathogens.

7. To contribute to the Initial Observing System of GOOS and the developing plans of GOOS COOP.

The CPR programme has been source of advice for programmes under GOOS, operating an open data policy, which is fully compliant with GOOS. SAHFOS contributed to GOOS posting annually on the SAHFOS website CPR results for Calanus finmarchicus and Phytoplankton Colour Index (PCI), which is a proxy for phytoplankton standing stock, averaged for CPR standard areas in the North Atlantic. Temperature data measured on selected CPR routes is also available on the website. The CPR has been recognised as a fundamental tool for an integrated and sustained marine observing system (General Joint Research Centre of the European Community, EU JRC Report 2006).

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8. To produce reports, publications and web based products on the results of the monitoring programme, including annual contributions to the Report Cards of the MCCIP as part of the SAHFOS Knowledge Transfer and Education programme.

SAHFOS has regularly updated and spread the results of the CPR monitoring, through the yearly publication of an Annual Report and Ecological Status Report summarising monitoring and research outputs, participation in ICES Working Groups and SAHFOS Education/KT initiatives. SAHFOS was involved and often led on several Knowledge Transfer activities, producing high quality policy documents. Those includes: 2 reports and 1 ICES Cooperative Research Report replying to requests from OSPAR and the ICES (Licandro et al., 2007; Anken-Nissen et al., 2008; ICES 2008); the Report Cards of the MCCIP (Edwards et al., 2007; Elliot et al., 2007; Raine et al., 2007; Heath et al., 2009; Reid et al, 2009; Edwards et al., 2010); Charting in Progress 2: An Integrated Assessment of the State of UK Seas (Reid et al., 2010). Other Knowledge Exchange Publications include “Climate Change and European Marine System Research (CLAMER)” and the “Atlas of Calcareous Plankton” that can be downloaded at the page http://www.sahfos.ac.uk/research/publications/knowledge-exchange-publications.aspx

9: Additional Milestones 10/11

Molecular analysis of jellyfishAdditional funding from DEFRA was allocated for the taxonomic identification of swarming jellyfish collected around UK by the CPR during 2010 and 2011. The identification is possible by using molecular methods on the gelatinous tissue collected from samples fully covered in jellyfish material. In the period of study jellyfish swarms were mainly detected on the western European shelf and in the English Channel, while only 2 samples collected from the CPR in September 2010 in the central North Sea were extremely reach in jellyfish material. Records from the time series of Stonehaven seem to confirm a relatively low abundance of hydrozoan jellyfish in 2010-2011 in the north-western North Sea.While the molecular analysis of jellyfish collected by the CPR is still ongoing, part of the funding has been spent to build up/expand a reference database of jellyfish DNA sequences that can be used to improve the taxonomic identification of jellyfish material extracted from CPR samples. Historical archive/formalin storeThe books reporting CPR zooplankton records of the period previous to the second world war have been archived in the database. A searchable database has been designed and populated with information on all stored CPR samples (e.g. location of the sample in the CPR archive, state of the sample), including records of the samples that have been given away for further analysis (e.g. molecular analysis).

Digitisation of historic paper recordsThe most relevant pre-war CPR analysis books have been chosen to be digitized. Zooplankton data collected along c. 200 North Sea routes in the middle-late 1930s have been included in the database, while the digitalisation of the Phytoplankton Colour Index (PCI) data collected during the same period is ongoing and will be soon completed.

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References to published material9. 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.

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Papers published in peer-reviewed journals generated within this project

1. Attrill, M.J., Wright, J. & Edwards, M. 2007. Climate-related increases in jellyfish frequency suggest a more gelatinous future for the North Sea. Limnology and Oceanography, 52: 480-485.

2. Batten, S.D. & Freeland, H.J. 2007. Plankton populations at the bifurcation of the North Pacific Current. Fisheries Oceanography, 16: 536-546.

3. Batten, S.D. & Burkill, P.H., 2010. The Continuous Plankton Recorder: towards a global perspective. Journal of Plankton Research, 32: 1619-1621.

4. Batten, S.D. & Walne, A.W., 2011. Variability in northwards extension of warm water copepods in the NE Pacific. Journal of Plankton Research, 33: 1643– 1653.

5. Baxter, E.J., Walne, A.W., Purcell, J., McAllen, R. & Doyle, T.K., 2010. Identification of jellyfish from Continuous Plankton Recorder samples. Hydrobiologia, 645: 193-201.

6. Beaugrand, G., Edwards, M., Brander, K., Luczak, C. & Ibanez, F., 2008. Causes and prediction of abrupt climate-driven ecosystem shifts in the North Atlantic. Ecology Letters, 11: 1157–1168.

7. Castellani, C., Lindley, J.A., Wootton, M. & Kirby, R.R., 2011. Morphological and genetic variation in the north Atlantic copepod, Centropages typicus. Journal of the Marine Biological Association of the UK, 92.1.

8. Edwards, M., Beaugrand, G., Hays, G.C., Koslow, J.A. & Richardson, A.J., 2010. Multi-decadal oceanic ecological datasets and their application in marine policy and management. Trends in Ecology and Evolution, 25: 602-610.

9. Edwards, M., 2011. Biodiversity and ecosystems: Change at the community level. Nature Climate Change, 1: 398–399.

10. Fauchald, P., Skov, H., Skern-Mauritzen, M., Johns, D.G & Tveraa, T., 2011. Wasp-Waist Interactions in the North Sea Ecosystem. PLOS-one, 6: e22729.

11. Fauchald, P., Skov, H., Skern-Mauritzen, M., Johns, D.G. & Tveraa, T., 2011. Scale-dependent response diversity of seabirds to prey in the North Sea. Ecology, 92: 228–239.

12. Fredericksen, M., Elston, D.A., Edwards, M., Mann, A.D. & Wanless, S., 2011. Mechanisms of long-term decline in size of lesser sandeels in the North Sea explored using a growth and phenology model Marine Ecology Progress Series, 432: 137-147.

13. Helaouët, P. & Beaugrand, G. 2007. Macroecology of Calanus finmarchicus and C. helgolandicus in the North Atlantic Ocean and adjacent seas. Marine Ecology Progress Series, 345: 147-165.

14. Helaouët, P., Beaugrand, G. & Reid, P.C., 2011. Macrophysiology of Calanus finmarchicus in the North Atlantic Ocean. Progress in Oceanography, 91: 217–228.

15. Kirby, R.R., Beaugrand, G., Lindley, J.A., Richardson, A.J., Edwards, M. & Reid, P.C. 2007. Climate effects and benthic-pelagic coupling in the North Sea. Marine Ecology Progress Series, 330: 31-38.

16. Kirby, R.R., Lindley, J.A. & Batten, S.D. 2007. Spatial heterogeneity and genetic variation in the copepod Neocalanus cristatus along two transects in the north Pacific sampled by the Continuous Plankton Recorder. Journal of Plankton Research, 29: 97-106.

17. Kirby, R.R., Beaugrand, G. & Lindley, J.A., 2008. Climate-induced effects on the meroplankton and the benthic-pelagic ecology of the North Sea. Limnology and Oceanography, 53: 1805-1815.

18. Lampitt, R.S., Salter, I. & Johns, D., 2009. Radiolaria: major exporters of organic carbon to the deep ocean. Global Biogeochemical Cycles, doi:10.1029/2008GB003221.

19. Leterme, S.C., Pingree, R.D., Skogen, M.D., Seuront, L., Reid, P.C. & Attrill, M.J., 2008. Decadal fluctuations in North Atlantic water inflow in the North Sea between 1958-2003: impacts on temperature and phytoplankton populations. Oceanologia, 50: 59-72.

20. Licandro, P. & Icardi, P., 2009. Basin scale distribution of zooplankton in the Ligurian Sea (north-western Mediterranean) in late autumn. Hydrobiologia, 617: 17-40. doi 10.1007/s10750-008-9523-9.

21. Licandro, P., Conway, D.V.P., Yahia, M.N.D., Puelles, M.L.F.d., Gasparini, S., Hecq, J.H., Tranter, P. & Kirby, R.R., 2010. A blooming jellyfish in the northeast Atlantic and Mediterranean. Biology Letters, 6: 688-691. (http://rsbl.royalsocietypublishing.org/content/6/5/688.full.pdf+html).

22. Licandro, P., Souissi, S., Ibanez, F. & Carré. C. 2011. Long-term variability and environmental preferences of calycophoran siphonophores in the Bay of Villefranche (North-Western Mediterranean). Progress in Oceanography (http://dx.doi.org/10.1016/j.pocean.2011.11.004).

23. Lindley, J.A. & Batten, S.D., 2008. Distribution and Seasonal Cycles of decapod crustacean larvae in Continuous Plankton Records from the North Pacific Ocean. Journal of the Marine Biological Association of the United Kingdom, 88: 443-451.

24. Lindley, J.A., Beaugrand, G., Luczak, C., Dewrumez, J.-M. & Kirby, R.R., 2010. Warm-water decapods and the trophic amplification of climate in the North Sea. Biology Letters, 6: 773-776 .

25. Lindley, J.A. & Kirby, R.R., 2010. Climate-induced changes in the North Sea Decapoda over the last 60 years. Climate research, 42: 257-264.

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