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Issue n°29 - December 2015

Issue n°29 - December 2015
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In this issue

Lisa Maddison


It is hard to believe that we have reached the end of yet another year! It has been a busy one for IMBER. Two of our regional programmes, ESSAS and CLIOTOP, held conferences during the year, and in October IMBER convened IMBIZO IV which continued the tradition of delivering good science, great discussions and a lot of fun. The next one will be held in 2017, and if you would like to host it, or know of a great venue for IMBIZO V, please contact me.

This issue of the IMBER Update is dedicated mostly to articles pertaining to the CLImate Impacts On Top Oceanic Predators (CLIOTOP) Regional Programme. They held their 3rd Symposium, titled "Future of oceanic animals in a changing ocean", in Spain in September. In addition to highlighting some of the excellent work that CLIOTOP working groups and individual scientists have done in the years since the last symposium in New Caledonia, this gathering also provided the opportunity to get community input about the direction that the programme should go in its next phase. Sadly, Alistair Hobday (CSIRO, Australia) stepped down as the CLIOTOP co-Chair after the symposium. However, he has been replaced by Karen Evans (also from CSIRO), who is very well placed to take the programme forward with co-Chair Kevin Weng.

IMBER is heading into a period of change. One of our co-sponsors, the International Geosphere-Biosphere Programme (IGBP) is ending on 15 December. IMBER will move into its second 10-year phase with the continued support of the Scientific Committee on Oceanic Research (SCOR), and we have submitted a transition document to become a core project of Future Earth. Internally, there will be several changes to the IMBER Scientific Steering Committee next year - read on to see who is coming and who is going...

Wishing everyone in the IMBER community happy holidays and a peaceful and prosperous new year.

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The 3rd CLIOTOP Symposium: Future of oceanic animals in a changing ocean

CLIOTOP Scientific Steering Committee and the 3rd CLIOTOP Symposium organising committee


CLIOTOP is an international research network open to researchers, managers, and policy makers involved in research related to large marine species. Network participants organise large-scale comparative efforts to elucidate key processes involved in the interaction between climate variability and change and human uses of the ocean on the structure of pelagic ecosystems and large marine species. CLIOTOP seeks to develop predictive capability for these socio-ecological systems and evaluate adaptation options  to ensure future sustainability.

CLIOTOP, one of the four regional science programmes of IMBER, held its third open science symposium in Donostia–San Sebastián in the Basque Country, Spain during 14 – 18 September 2015. The venue for the symposium, the Palacio Miramar, once the summer holiday home for the royal family, provided a regal and historic venue. While science was foremost at the meetings, the lively bar and pintxos scene in the old town provided for congenial evening gatherings of symposium attendees.

The symposium was organised by an international committee and hosted by AZTI Tecnalia. Approximately 120 delegates attended from 24 countries (compared with nearly 200 participants from 25 countries at the first and approximately 70 participants from

18 countries at the second symposium), with research interests ranging from conservation biology, fisheries science, socio-economics, oceanography, conservation and fisheries management, meteorology and climate data management.

3rd CLIOTOP - Group photo

Figure 1. Participants of the 3rd CLIOTOP symposium in the grounds of the Palacio Miramar, Donostia


Symposium Sponsors

The financial support provided by sponsors was key to the success of the symposium in attracting a diverse international group to Donostia-San Sebastián and in also enabling the hosting of several social functions. In particular, the symposium organising committee appreciate the support provided by AZTI Tecnalia, Basquetour, CSIRO, Collecte Localisation Satellites (CLS), IMBER, Intergovernmental Oceanographic Commission (IOC), Institut de recherche pour le développement (IRD), NOAA and PICES. 


Symposium objectives

The overall objectives of the 3rd CLIOTOP symposium were to:

  1. Evaluate impacts of climate variability and change over seasonal to decadal time scales on pelagic species and dependent socio-economic and management systems
  2. Identify risk assessment and evaluation tools that incorporate climate variability in order to improve sustainable resource management (conservation, fisheries, spatial planning, etc.)
  3. Identify sustainable pathways for coupled socio-ecological oceanic systems
  4. Position CLIOTOP science for the next 10-year phase as part of Future Earth, and build a collaborating community of scientists, managers, and policy-makers.

The 3rd symposium detailed impacts on species from individual climate stressors, interactions of multiple stressors and the impacts on populations via changes in these. It also focused on understanding the responses of the mid-trophic community to change, and interactions with higher predator groups. Various management approaches and socio-economic consequences were explored, and adaptation options to reduce vulnerability to climate change were evaluated and assessed.

3rd CLIOTOP - Sympoiusm dinner

Figure 2. Symposium delegates enjoying the ambience of the cellar room at the Petritegi Cider House during the symposium dinner


Symposium themes and presentations

At the two previous symposia, presentations were orientated specifically around the CLIOTOP working groups and a small number of cross-cutting sessions. At this 3rd Symposium, however, the presentations were organized into nine themes, some of which focused on aspects of the research of the working groups. The themes highlighted a number of research initiatives that span across working groups, for example: integrating movement and behaviour with trophic ecology (WG2 and 3), exploring early life history and physiological constraints on habitat utilisation (WG 1 and 2) and incorporating socio- economics and management strategies into synthesis frameworks (WG 4 and 5).

1. Early life history of pelagic species – winners and losers in the future ocean (moderator: Maria Gasalla)

The early life history dynamics of oceanic top predators are likely driven by a combination of density-dependent and -independent processes, each effecting survival, and ultimately year-class strength. These early life history dynamics are closely linked to environmental processes, many of which are demonstrably influenced by climate variability. For example, changes in climate can impact ocean temperature distribution, timing and depth of stratification, the formation of mesoscale structures such as fronts and gyres, upwelling and consequently, production. Changes in production can directly influence rates of growth and mortality of early life stages of top predators, impacting their survival. Presentations in this theme detailed linkages between larval/early life stages and thermal habitats as well as oceanic pathways, the trophodynamics of larval/early life stages and differing vulnerabilities of life stages to fishing were also explored.

2. Implications of potential distribution and abundance shifts in oceanic organisms for food security and species conservation (moderator: Dan Costa)

Observed or projected changes in the distribution and abundance of pelagic species will have flow-on effects for ecosystem management and the societies that depend on natural resources. Altered distributions of harvested species can lead to changes in access to resources, disadvantaging some communities and therefore impacting their food security while providing new access benefits for others. Changes in distribution of bycatch or protected species can lead to conflict with stressors such as shipping, oil and gas exploration, or fishing, and may require new management interventions. Presentations within this theme explored the utility of various models in exploring and projecting changes in the habitats of species in response to a changing climate and the flow-on effect on fisheries indices and management structures/schemes.

3. Trophic pathways in open ocean ecosystems - changes in mid-trophic level community composition as a result of changes to physical, chemical and biological components of the marine environment (moderator: Shiroh Yonezaki)

There is growing evidence that climate variability affects primary producers in marine systems, imparting bottom-up forces on food webs via trophic pathways. Simultaneously, fisheries removals of upper-level predators can have cascading effects on the underlying trophic levels. The largest gap in our knowledge of trophic pathways in pelagic ecosystems remains the intermediate trophic levels. They include small fishes, cephalopods and crustaceans  and comprise the forage base for upper-level predators. Any changes in the abundance or community composition of intermediate trophic levels will, therefore, have implications for upper-level predators. Delineating the key trophic pathways linking primary production to the upper trophic levels through the forage groups, needs better understanding of how intermediate trophic groups and associated trophic pathways vary among productivity regimes, ecosystems and as a result underlying changes in these. Presentations within this theme were largely focused on classical approaches to ascertain trophic linkages (e.g. stomach content analysis) and the use of long-term trophic linkage datasets to determine change over time. The importance of such long-term datasets as inputs into ecosystem models and in linking primary production and higher trophic level components was also highlighted.

4. Integrated modelling to project and explore future patterns - evaluation of model complexity vs. generality, evidence of important processes to include in models, and evaluation of model results (moderator Sophie Bestley)

Models are increasingly available to evaluate future patterns in pelagic ecosystems. For example, quantitative indicators that characterize ecosystem status and the ongoing performance of oceanic management systems can forge links between resource managers, stakeholders, and scientists. Such indicators provide a mechanism for discussing the conservation and exploitation of living marine resources in an ecosystem context. No single modelling approach is likely to have predictive power at all spatial and temporal scales of interest, and management decisions must be made on both strategic and tactical scales, utilising a variety of approaches. In general, models with a larger ecological scale (e.g. trophic or spatially explicit ecosystem models) are expected to describe decadal scale forcing related to changes in whole-system productivity, predict the ecosystem consequences of changes in system productivity and fishing mortality, and provide strategic management advice (e.g. advice on allocating fishing mortality under alternative productivity regimes). In contrast, models with smaller ecological scales (e.g. single species assessment models that include environmental forcing) will typically describe sub-decadal scale forcing on population-dynamics processes like recruitment, predict the consequences of environmentally-forced variability in these processes under alternative patterns of fishing mortality, and provide tactical management advice (e.g. annual harvest quotas). Presentations within this theme detailed modelling frameworks ranging from complex simulations of important processes, either within species or particular regions through to basin/global scale integration of assimilated data to explore fishery/human processes. Presentations highlighted the importance of data quality control processes and evaluation and validation of modelling frameworks in building confidence in end users.

5. Socio-economic aspects and management strategies – what are the key needs and resulting decisions and actions that should guide oceanic resource management under climate change (moderator: Olivier Maury)

Understanding the relationships between ecological, economic and social objectives, and the trade-offs between each is important in designing policies to manage, conserve or restore marine ecosystems. Further demands on ocean resources are expected with increasing populations, associated food security and utilisation of marine ecosystems and emerging industries such as renewable energies and offshore marine production systems. A changing climate is expected to place further pressures on marine environments and will interact with demands in varying ways. Managing these multiple uses, some with often conflicting objectives, to ensure sustainable ecosystems, industries and communities is a major challenge globally. Presentations within this theme highlighted the importance of modelling and scenario development for understanding the feedbacks between social/ecological systems in response to changes in both systems. The importance of considering a changing climate in developing and evaluating management approaches/strategies was also discussed.

6. Influence and role of biophysical processes and feedbacks on top predators (moderator: Patrick Lehodey)

Models developed for marine species and ecosystems now include many physical forcing factors, the effects of which propagate up the food web. Given the mobility of many marine species, and the lag in response of higher order animals to changes in the physical environment, there is a need to identify appropriate methods to understand how biophysical processes influence marine species, and relevant environmental or response variables for methodological approaches. Focus has been on primary environmental variables, such as temperature, and the physiology and behaviour of top predators. But what about changes in upwelling, currents, or mixed layer depth? How do these affect foraging efficiency, predation, migration and survival? Presentations within this theme highlighted the increasing use of very detailed/accurate data on animal movement and physiology and the use of statistical models to explore habitat features that may be of importance to animal movement and physiology.

7. Biodiversity, conservation and adaptive management – future strategies for incorporating long term change (moderator: Rebecca Lent)

A range of marine species are either recovering from past exploitation, or continue to be subjected to ongoing direct or indirect population threats. In some instances, conservation policy and management of pelagic species has had some success in terms of population recovery, but not so in other cases. Models can now generate scenarios of future trajectories for populations under climate change. However, thus far, they are only used to generate and explore hypotheses and as a result management strategies currently lack the ability to account for the responses of populations to long-term change. A key focus is to identify what is needed to incorporate scenarios of marine populations under climate change and what may be required to build adaptive management mechanisms to inform conservation and resource management in the future. Presentations within this theme highlighted the need for clear information about threats, early action in adaptation and mitigation and active engagement with stakeholders to inform policy on appropriate management measures. Regions such as the Arctic, where human activity is increasing and having a major transformative impact, were identified as key areas for research, as was research into the design of spatial protection areas and implementation of monitoring programs.

8. Scenarios of large marine organisms and their fisheries in a changing world (moderator: Inna Senina)

A range of modelling frameworks can be used to generate and investigate scenarios of future distributions and abundances of pelagic species and flow-on implications for industries that rely on them. These range from simple response variable models, to models of intermediate complexity and complex ecosystem models coupled to earth system models forced by emission scenarios developed by the Intergovernmental Panel on Climate Change (IPCC). Model projections are not only important for investigating changes in populations, but also for the evaluation of the effectiveness of management strategies into the future and for informing adaptations that might be required to achieve socio-economic or conservation objectives, efficiencies or trade-offs within a changing world. Presentations within this theme identified the need for (i) expanded synergies between observations and predictive models to improve understanding of the impacts of a changing environment on pelagic species and (ii) faster reactions to changes by developing effective management strategies and providing scientific advice to policy-makers. This can be facilitated through the use of data from currently under-utilised sources (e.g. echo-sounder equipped FADs), using the relationship between low-frequency variability in the environment and species population dynamics as predictive tools in population assessments, and by including human components, such as fleet dynamics, when exploring population projections under climate scenarios.

9. Data, analyses and tool development associated with understanding the impacts of climate variability on fisheries (moderator: Kylie Scales)

Variability in the marine environment is largely driven by climatic processes that occur on time scales ranging from days to decades and at spatial scales of less than a kilometre to whole ocean basins. This variability has a strong influence on marine species, through direct and indirect interactions. There is still much to learn about the scales at which these responses occur, how observations collected at the scale of the individual can be used to understand population responses, and how observations can explain responses to variability in the marine environment. Further, identification of appropriate methods for data collection and analysis, and the management of ever-increasing observation datasets collected at finer scales and for longer periods are required. Presentations in this theme detailed various modelling approaches used to identify important habitats for breeding and feeding. They also detailed biochemical and genomic tracers and species composition metrics used to identify the horizontal and vertical utilisation of habitats, and oceanographic and climate datasets of importance for forecasting habitats of importance for species.


Keynote addresses

In addition to contributed oral and poster presentations, six keynote speakers addressed topics central to the objectives of the symposium.

Dr Lisa Balance, NOAA and Scripps Institution of Oceanography, reviewed methods to assess the health and condition of marine mammals (cetaceans) and the growing awareness of the multiple ways marine mammals (top predators) may be structuring oceanic environments (e.g. via predation, nutrient fluxes).

Prof. Emanuele Di Lorenzo, Georgia Institute of Technology and PICES, provided an overview of climate-driven processes in the North Pacific Ocean and examined the mechanisms and linkages associated with the varying forms of El Niño, its relation to the Pacific Decadal Oscillation (PDO), and how these might change in the future.

Professor Molly Lutcavage, University of Massachusetts, Boston, reviewed progress in establishing the movements and habitats of large pelagic predators in the North Atlantic and their relevance to the current management frameworks for these species.

Dr Dale Squires, NOAA and University of California San Diego, reviewed management and policy approaches to bycatch reduction and highlighted the importance of careful decision making in choosing policy instruments and research strategies to support these.

Dr Einar Svendsen, IMBER and Institute of Marine Research reviewed the future needs for an ecosystem approach in research and management within the framework of IMBER, including the importance of developing marine observation networks integrated on global scales to ensure the conservation of the ocean for sustainable use.

Dr Jock Young, CSIRO Oceans and Atmosphere, provided an overview of the research activities of the CLIOTOP Working Group 3 since its inception in 2003. He highlighted the importance of bringing together researchers with regional datasets by providing a global approach for the assessment of trophic ecology and to identify how food webs may change in the future.

3rd CLIOTOP - Keynote speakers

Figure 3. CLIOTOP symposium keynote speakers (L-R): Dale Squires, Lisa Balance, Einar Svendsen, Molly Lutcavage, Jock Young (Emanuele Di Lorenzo absent).


Progress since the 1st and 2nd CLIOTOP symposia

Once again, the CLIOTOP symposium was dominated by presentations on fish and the Pacific (see Table 1). Although at this symposium, there was still a focus on single response variables, many used multi-species, multi-region, global and longer-term datasets in their comparisons. There was also a greater focus on forecasting shorter time scale responses rather than longer term projections, thereby expanding the time scales at which responses to change in the environment are being investigated.

With regard to the overall objectives of the symposium, some topics are being addressed, but those listed below require more attention and could perhaps be the focus of targeted task teams within the framework of CLIOTOP Phase 3.

1. Evaluate impacts of climate variability and change on pelagic species and dependent socio-economic and management systems over seasonal to decadal time scales.

Increased understanding of the habitat requirements of species now permit the evaluation of their responses to variability and change in the environment. Presentations showed a general increase in the assessment of these responses under formal scenario conditions, using earth system model outputs. Additional research is needed to assess responses, using a wider range of future climate scenarios and multiple earth system model outputs (ensemble approaches). It is therefore expected that research in this area will increase in the coming years, particularly with the incorporation of output from initiatives such as Fish-MIP into the IPCC process. Presentations explored projections of the impacts of climate change on socio-economic systems associated with fisheries, the applicability of adaptive management  and modelling frameworks to incorporate the responses of fleets through changes in effort into projections and assessment. As understanding of the responses of species to climate variability and change increases, it is expected that there will be greater focus on exploring the links to associated human systems.

2. Identify risk assessment and evaluation tools that incorporate climate variability in order to improve sustainable resource management (conservation, fisheries, spatial planning, etc.).

Risk assessment, adaptation and evaluation tools were not strong themes at the symposium, highlighting the need for the development of tools that incorporate climate variability for improved resource management. The need for conservation managers to move beyond ‘preserve and protect’ systems to adaptation engineering systems, and that it is critical that stakeholders are involved in scenario science to improve fisheries management policies were emphasised.

3. Identify sustainable pathways for coupled socio-ecological oceanic systems

Exploration of sustainable pathways for fisheries or conservation management was only touched on during the symposia, highlighting an area of research that is currently under-explored. There is clearly a need to develop opportunities for expanding effort in these areas of fisheries and conservation management.


Table: Comparison of the focus of presentations (% of total presentations) and the extent of author and country collaborations at the three CLIOTOP symposia held to date

3rd CLIOTOP - Table

4. Position CLIOTOP science for the next 10-year phase as part of Future Earth, and build a collaborating community of scientists, managers and policy-makers.

Presentations provided an overview of where understanding has increased, where it is still needed and where potential task teams developed for Phase 3 of CLIOTOP might have the greatest impact. The final part of the symposium focused on discussions around task teams and potential research areas. These discussions within the community will continue as the process for establishing task teams is finalised and their work is initiated.

Awards, publications and future meetings

The best oral presentation by a student was awarded to Benjamin Arthur for his presentation Predicting the availability of foraging habitats under changing ocean conditions: what can seven years of tracking a wide-ranging predator tell us? Yoshinori Aoki was given an honourable mention for his presentation Change in energy acquisition of skipjack tuna (Katsuwonus pelamis) with northward migration in the western Pacific Ocean.

The best poster presentation by a student was awarded to Stephanie Czudaj for her presentation Food-web structure of mesopelagic communities in high and low oxygen environments in the eastern tropical north Atlantic as identified by stable isotope analysis. Honourable mention went to Alice Della Penna for her presentation Quasi-planktonic movements of top marine predators: how does horizontal advection affect animal movement?

Einar Svendsen presented Alistair Hobday with a farewell and thank you gift from IMBER.  Alistair has served tirelessly as co-Chair of CLIOTOP for the past six years, and we are delighted that he will continue to contribute as a member of the CLIOTOP Scientific Steering Committee.


Figure 4. The CLIOTOP Scientific Steering Committee helping Alistair Hobday to put his feet up after stepping down from the co-chair position.

A special issue of Deep Sea Research will publish papers from the symposium, with submissions being prepared from many attendees. It is particularly pleasing to see that several papers are also being prepared as a result of discussions arising from the symposium discussion sessions.

The next CLIOTOP open science symposium is scheduled for 2018, with the venue and theme yet to be finalised. It is hoped that in the meantime, there will be CLIOTOP involvement at the IMBER IMBIZO and summer school series, as well as interactions between CLIOTOP community members through operational task teams.

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Science Highlights from the 3rd CLIOTOP Symposium

Finding winning strategies in a changing ocean

Mary-Anne Lea1,2, Jeremy T. Sterling1, Noel Pelland1, Sharon Melin1, Rolf Ream1 and Tom Gelatt1

1. National Marine Mammal Laboratory, NOAA Alaska Fisheries Science Center, 7600 Sand Point Way, Seattle 98115, WA USA.

2. Institute for Marine and Antarctic Studies, University of Tasmania, 20 Castray Esplanade, Battery Point, TAS 7004, Australia.


Understanding how young animals respond to changing oceanic conditions is critical for predicting population responses to anticipated change in marine ecosystems. For Northern fur seals (Callorhinus ursinus) that have experienced rapid population declines in the Bering Sea, and are one of only two fur seal species to conduct large-scale ocean basin-wide migrations (Fig. 5), this need is pressing.

Lea-Fig 3

The Alaska Ecosystems Program at the NOAA National Marine Laboratory has used evolving biotelemetry techniques over the last two decades to characterise the habitat requirements and foraging strategies of breeding northern fur seals (Baker 2007, Sterling 2014). More recently this approach has extended to include newly-weaned pups (n=166) tracked from the four North American populations in Alaska and California to compare and contrast regions of higher use by naïve animals with those regions favoured by experienced adult females.

Pups born in Alaskan waters, from both declining and increasing populations, travel thousands of kilometres to waters associated with the Transitional Zone Chlorophyll Front (TZCF) and upwelling features in the California Current (CC) and North American continental margin (Fig. 5). Departing during winter, pups must contend with a suite of oceanic conditions including extreme weather events and associated wave heights (see Lea et al., 2009). High wind speeds and wind direction influence pup movement behaviour by causing them to travel and move away from potentially good feeding grounds (Fig. 6c). Severe storms may also lead pups to exhaust all of their resources and starve (Scheffer 1950). While some pups travel northwards to the Bering Sea ice edge, the majority head southwards. Pups that reach west coast Canada and US waters may encounter productive upwelling ecosystems (Scheffer 1950) as well as significant domoic acid algal bloom and predation risks (García‐Reyes and Largier 2012). Recent, numerous strandings of emaciated California sea lion pups in the California Current region also highlight the impact changing/warming ocean conditions may exert on prey availability at a critical time for newly-weaned migrants and local suckling species.

The transition from early fall to winter in the Bering Sea includes rougher seas, shorter days, dropping temperatures, and upper ocean mixing due to storms – all factors known to change mid-trophic species distribution and abundance and thus influencing the northern fur seal population to migrate to more productive and energetically profitable foraging grounds. In contrast to adults, pup migration patterns were less consistent, seemingly random, and showed year-to-year variability in dispersal patterns corresponding to markedly different at-sea conditions. This suggests that young animals undergo a period of learning at sea while contending with rapidly changing environmental conditions (Fig. 5). The transition from traveling to foraging state varies with size and may reflect interactions between physiological capabilities, experience and environmental conditions. 

Adult male fur seals capable of diving far deeper than adult females and pups transition from a transit to foraging state more quickly than females and pups (Fig. 6a) and are less influenced by changes in wind strength. Adult female foraging state is susceptible to influences in wind speed during the early migration phase (Fig. 6b), while more shallow diving pups may take up to six months to transition from transit to proportionally greater foraging states (Fig. 6c).

Examination of individual tracks reveals that storms, ocean eddies, and other oceanographic phenomena elicit strong behavioural reactions in experienced, mature adults (Sterling et al. 2014). Female habitat utilization around coastal capes, eddies and regions of strong oceanographic variability suggests that the predictability of productive regions along the Californian coast is likely central to the success of the migratory life cycle adopted by Alaskan northern fur seals. Further study is needed to better understand the effects of storms and ocean climate on pup condition, survivorship, and northern fur seal demography, given observations (Ruggiero et al. 2010) and climate projections (Wang et al. 2014) of alterations in the strength and annual cycle of wave heights in the North Pacific Ocean. These trends are in addition to strong interannual and decadal climate variability that plays a currently unknown role in Bering Sea northern fur seal demography, but is known to play a significant role in adult and pup survivorship for northern fur seals breeding on San Miguel Island.  A thorough understanding of the spectrum of foraging strategies exhibited by naïve and experienced fur seals provides valuable insight into a future where productive regions may become less predictable.

Lea-Fig 1

Figure 5. Seasonal habitat use of adult male, female and newly-weaned pups tracked using satellite tags between 1991 and 2010. Tracks are presented for those animals transmitting for >60d.Coloured regions encompassing the 75% habitat utilization contour are coloured for each age class [adult males – black, adult female –red, pups – blue]. TZCF – Transition Zone Chlorophyll Front.

Lea-Fig 2

Figure 6. Behavioural state response for a. adult male, b. adult female and c. pup Northern fur seals in response to changing wind strength. Mean dive depth and wind speeds denoted in red and black respectively. Dashed blue lines indicate the number of days past October 1 when seals from each age class transition to a more resident behavioural state.



  • Baker J.D. 2007. Post-weaning migration of northern fur seal Callorhinus ursinus pups from the Pribilof Islands, Alaska. Marine Ecology-Progress Series 341: 243-255.
  • Loughlin T.R., Baba N., Robson B.W. in Dynamics of the Bering Sea, T. R. Loughlin, K. Ohtani, Eds. 1999. University of Alaska Sea Grant, Fairbanks, AK,  chap. 27, pp. 615-630.
  • Pelland N.A., Sterling J.T., Lea, M-A., Bond, N.A., Ream, R.R., Lee, C.M., Eriksen, C.C. 2014. Fortuitous Encounters between Seagliders and Adult Female Northern Fur Seals (Callorhinus ursinus) off the Washington (USA) Coast: Upper Ocean Variability and Links to Top Predator Behavior. e101268.
  • Ream R.R., Sterling, J.T., Loughlin, T.R. 2005)Oceanographic influences on northern fur seal migratory movements. Deep Sea Res. Part II, 52, 823-843.
  • Sterling J.T., Springer, A.M., Iverson, S.J., Johnson, S.P., Pelland, N.A., Johnson, D.S., Lea, M-A., Bond, N.A. 2014. The sun, moon, wind, and biological imperative–shaping contrasting wintertime migration and foraging strategies of adult male and female northern fur seals (Callorhinus ursinus). PloS ONE 9(4).
  • Lea M.-A., Johnson, D., Ream, R., Sterling, J., Melin, S., Gelatt, T. 2009. Extreme weather events influence dispersal of naive northern fur seals. Biology Letters 5: 252-257.
  • Scheffer V.B. 1950. Winter injury to young fur seals on the northwest coast. Calif. Fish Game 36, 378-379.
  • García‐Reyes M., Largier J.L. 2012. Seasonality of coastal upwelling off central and northern California: New insights, including temporal and spatial variability. Journal of Geophysical Research: Oceans (1978–2012) 117(C3).
  • Rust L., Gulland F., Frame E., Lefebvre K. 2014. Domoic acid in milk of free living California marine mammals indicates lactational exposure occurs. Marine Mammal Science 30: 1272-1278.
  • Ruggiero P., Komar P.D., Allan J.C. 2010. Increasing wave heights and extreme value projections: The wave climate of the US Pacific Northwest. Coastal Engineering 57(5): 539-552.
  • Wang X.L.,  Feng Y.,  Swail V. R. 2014. Changes in global ocean wave heights as projected using multimodel CMIP5 simulations, Geophys. Res. Lett., 41, 1026–1034, doi:10.1002/2013GL058650
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Changing climate and the winter foraging ecology of Antarctic fur seal populations

Ben Arthur

Institute for Marine and Antarctic Studies (IMAS), University of Tasmania, Hobart, Tasmania, Australia.

Email: Benjamin.Arthur@utas.edu.au

Websites: www.imas.utas.edu.au, www.researchgate.net/profile/Benjamin_Arthur


The Southern Ocean is a rapidly changing environment and there is a pressing need to increase our understanding of this complex ecosystem, the impacts of large-scale climate changes, and the effectiveness of conservation and management measures. Despite many studies into the effects of large-scale climatic changes on the physical structure of the Southern Ocean, how such changes will impact ecosystem structure and function are poorly understood. Any changes will, however, be reflected in the responses of high-trophic level species, such as the Antarctic fur seal (Arctocephalus gazella), making them ideal indicators of wider ecosystem change.

Arthur-Fig 1

Figure 7. Antarctic fur seals on Marion Island (Southern Indian Ocean). Credit Ben Arthur

In the Southern Ocean, variation in prey availability to higher trophic levels occurs as part of normal oceanographic and atmospheric phenomena. However, there is increasing evidence of anthropogenic changes in the distribution and abundance of prey, particularly through large-scale climate changes. Several studies have documented demographic changes in predator populations associated with the predicted effects of climate change and there are growing signs that responses such as these are indicative of large-scale ecosystem shifts. However, despite these studies, the links between the physical effects of climate change, biological productivity and the response of top predators in the Southern Ocean remain poorly understood.

During the winter, Antarctic marine predators face a substantially different environment, both physically and biologically, from the summer season. As a result, winter foraging patterns can differ markedly from the more constrained summer foraging behaviour that are well documented for many species. Quantitative studies into the winter habitat use and foraging ecology of top predators are therefore critical to better understand the influence of environmental variability and fluctuations in prey resources on higher trophic level species. Until recently, quantifying the habitat use of Antarctic marine predators during the pre-breeding winter period was difficult, but advances in bio-logging technology have made tracking the at-sea movements of animals easier and more affordable.

The winter foraging behaviour of the Antarctic fur seal is the focus of an international collaborative study led by the Institute for Marine and Antarctic Studies (IMAS) at the University of Tasmania, Australia. The Antarctic fur seal is an abundant and key Southern Ocean predator. The global population has increased rapidly since sealing operations ceased in the early 20th century after the species was driven to economic extinction.

Since 2008, the winter foraging movements of over 200 female Antarctic fur seals has been studied at three, circumpolar breeding sites (Marion Island, Bird Island and Cape Shirreff) using miniaturized geo-locating (GLS) tags. The project is a collaboration between the Antarctic research programs of Australia, South Africa, Britain, the United States and, soon, France. Such a coordinated study is greatly improving our understanding of the habitat and dietary preferences of this top predator, information that is needed to inform ecosystem models and management measures in the Southern Ocean. Such information is directly applicable to a number of international objectives, including the management imperatives of The Convention for the Conservation of Antarctic Marine Living Resources (CCAMLR) and the Southern Ocean Observing System (SOOS).

Arthur-Fig 4

Figure 8. Female Antarctic fur seal carrying a geolocation tag and her pup. Credit Chris Oosthuizen.

Arthur-Fig 2

Figure 9. Tracks of the winter foraging trips of ~200 female Antarctic fur seals between 2008-14. Locations in red indicate foraging behaviour while location in grey represent travelling behaviour.

Arthur-Fig 3

Figure 10. Core winter foraging habitats of female Antarctic fur seals (green) in relation to CCAMLR (Commission for the Conservation of Antarctic Marine Living Resources) management areas.

The unique behavioural data set resulting from this program has provided an opportunity to understand the effects of climate shifts on a higher-level predator. To date, this project has revealed strong fidelity to oceanic foraging grounds by individuals over multiple years (Arthur et al. 2015), which raises questions about the ability of long-lived animals to respond to future ocean change. We have also provided the first insights into the diving behaviour and vertical habitat use of Antarctic fur seals during the winter period, showing that animals from sub-Antarctic Marion Island employ two distinct foraging strategies, having to balance prey type and accessibility with the energetic costs associated with traveling to remote Southern Ocean habitats during winter (Arthur et al. In review).

Current work has, for the first time, identified important winter foraging habitats for female Antarctic fur seals from several Southern Ocean colonies, and the oceanographic factors influencing the species's use of these (Arthur et al. In prep). The habitat modelling approach is being extended to incorporate long-term anomalies of key environmental predictors, allowing past estimates of fur seal foraging habitat in the Southern Ocean over recent decades to be compared with current observations. Such information will ultimately enable predictions of foraging habitat under various climate change scenarios to reveal how the winter foraging habitats of this important top consumer in the Antarctic marine ecosystem may shift in the future.

Thank you to the Australian Wildlife Society and the Antarctic Climate & Ecosystems Cooperative Research Centre for their generous support which allowed me to attend the 3rd CLIOTOP Symposium. 



  • Arthur B., Hindell M., Bester M., Trathan P., Jonsen I., Staniland I., Oosthuizen W.C., Wege, M., Lea, M-A. 2015. Return customers: Foraging site fidelity and the effect of environmental variability in wide-ranging Antarctic fur seals. PLos ONE 10(3): e0120888.
  • Arthur B., Hindell M., Bester M.N., Oosthuizen W.C., Wege, M., Lea M.-A. South for the winter? Within-dive foraging effort reveals the trade-offs between divergent foraging strategies in a free-ranging predator. (In review).
  • Arthur B., Hindell M., Bester M.N., Trathan P., Goebel,M., Lea M.-A. Winter foraging habitats of a key Southern Ocean predator, the Antarctic fur seal (Arctocehpalus gazella). (In prep.).
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Modelling the oceanic habitats of pelagic fish using recreational fisheries data

Stephanie Brodie1, 2,*, Hobday A.J.3, Smith J.A.1,2, Hartog J.R.3, Spillman C. M.4, Everett J.D.1, 2, Taylor M.D.1,5, Gray C.A.1,6 & Suthers I.M.1, 2

1. School of Biological, Earth and Environmental Sciences, University of New South Wales, Sydney, Australia

2. Sydney Institute of Marine Science, Sydney, Australia

3. Commonwealth Scientific and Industrial Research Organisation, Hobart, Australia

4. Bureau of Meteorology, Melbourne, Australia

5. New South Wales Department of Primary Industries, Port Stephens Fisheries Institute, Nelson Bay, Australia

6. WildFish Research, Grays Point, Australia

Contact email: stephanie.brodie@unsw.edu.au



Understanding the ecological processes that drive the distributions and movements of organisms is a fundamental goal of ecology. Species distribution models are useful tools for describing the environmental requirements of species, and can also support dynamic marine management strategies (Maxwell et al., 2015).  However, such ecological models require substantial amounts of data sampled over long periods of time. We provide an example of how alternate data resources from a citizen science program can be applied to distribution modelling of pelagic fish (Brodie et al., 2015).

Species distribution models of two pelagic fish - dolphinfish (Coryphaena hippurus) and yellowtail kingfish (Seriola lalandi) were developed using 19 years of presence-only data from a recreational angler-based catch-and-release fishing program. The application of the catch-and-release data had issues with fisheries-dependence and a lack of recorded fishing effort, but these issues were solved through the use of modelling techniques and data thinning (Brodie et al., 2015). A Poisson point process model within a generalised additive modelling (GAMM) framework was used to determine the species distributions off the east coast of Australia as a function of several oceanographic covariates. This modelling approach converted the probability of fish occurrence to a metric of fish abundance (fish km-2; Renner et al., 2015).

The distribution models revealed the oceanic habitats of both dolphinfish and kingfish, and the importance of frontal features for these two species. Sea surface temperature, sea level anomaly, sea surface temperature frontal index, and eddy kinetic energy were significant environmental predictors for both dolphinfish and kingfish distributions. Predicted habitats for both species indicate greater fish intensity off the east Australian coast during summer and autumn in response to the regional oceanography, namely shelf incursions by the East Australian Current (Figs. 11 & 12). This study provides a framework for using presence-only recreational fisheries data to create species distribution models that can contribute to future dynamic spatial management of pelagic fisheries.

Brodie-Fig 1

Figure 11. Spatial prediction of dolphinfish intensity off east Australia across four seasons: A) spring B) summer C) autumn and D) winter. Colours represent the intensity of dolphinfish in number of fish km-2, and the grey line represents the 500 m isobath. 

Brodie-Fig 2

Figure 12. Spatial prediction of kingfish intensity off east Australia across four seasons: A) spring B) summer C) autumn and D) winter. Colours represent the intensity of kingfish in number of fish km-2, and the black line represents the 500 m isobath. Predictions are limited to 500m isobath due to kingfish association with coastal reefs.

Using this model framework, we created a seasonal forecast of fish habitats that predicts the distribution of dolphinfish abundance along the east coast of Australia. Seasonal forecasting of marine species distributions has been used as a decision support tool to help reduce uncertainty in commercial and aquaculture fisheries in Australia (Eveson et al., 2015; Hobday et al., 2015). However, there has not been any application of these tools by the recreational fisheries sector despite their increasing use of marine resources, and important economic and social contributions to coastal communities. We have developed a seasonal forecast of dolphinfish abundance off the east coast of Australia as a decision support tool for the recreational fisheries sector. The forecasting model showed good predicative skill up to six months into the future, and could effectively communicate fish abundance along the east coast of New South Wales, Australia (Fig. 13).

Brodie-Fig 3

Figure 13. Visualisation of the forecast for an example month (January 2012): A) the spatial map of observed dolphinfish intensity (fish km-2); B) spatial map of forecast dolphinfish intensity (fish km-2) for January 2012, lead time 1.


Continued development and application of seasonal forecasts will likely improve resilience and reduce uncertainty in fishery industries and marine resource management. The use of data from citizen science programs, such as catch-and-release fishing programs, can contribute to future dynamic spatial management of pelagic fisheries. Furthermore, combining global, or broad-scale, fisheries records may provide large enough datasets to comparatively analyse species distributions across, and between, ocean basins (e.g. Arrizabalaga et al., 2015). A wider application of such datasets may provide greater understanding of the key environmental and biological drivers of key trophic groups in pelagic ecosystems.



  • Arrizabalaga H., Dufour F., Kell L., Merino G., Ibaibarriaga L., Chust G., Irigoien X., Santiago J., Murua H., Fraile I., Chifflet M., Goikoetxea N., Sagarminaga Y., Aumont O., Bopp L., Herrera M., Marc Fromentin J., Bonhomeau S. 2015. Global habitat preferences of commercially valuable tuna. Deep-Sea Research Part II: Topical Studies in Oceanography 113: 102-112.
  • Brodie S., Hobday A.J., Smith J.A., Everett J.D., Taylor M.D., Gray C.A., Suthers I.M. 2015. Modelling the oceanic habitats of two pelagic species using recreational fisheries data. Fisheries Oceanography 24: 463-477.
  • Eveson J.P., Hobday A.J., Hartog J.R., Spillman C.M., Rough K.M. 2015. Seasonal forecasting of tuna habitat in the Great Australian Bight. Fisheries Research 170: 39-49.
  • Hobday A.J., Spillman C.M., Hartog J.R., Eveson J.P. 2015. Seasonal forecasting for decision support in marine fisheries and aquaculture. Fisheries Oceanography doi:10.1111/fog.12083. 
  • Maxwell S.M., Hazen E.L., Lewison R.L., Dunn D.C., Bailey H., Bograd S.J., Briscoe D.K., Fossette S., Hobday A.J., Bennett M., Benson S., Caldwell M.R., Costa D.P., Dewar H., Eguchi T., Hazen L., Kohin S., Sippel T., Crowder L.B. 2015. Dynamic ocean management: Defining and conceptualizing real-time management of the ocean. Marine Policy 58: 42-50.
  • Renner I.W., Elith J., Baddele, A., Fithian W., Hastie T., Phillips S.J., Popovic G., Warton D.I. 2015. Point process models for presence‐only analysis. Methods in Ecology and Evolution 6: 366-379.
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Towards comprehensive open ocean food web understanding

Anela Choy

Monterey Bay Aquarium Research Institute (MBARI)

Contact email: anela@mbari.org


Waters seaward of Earth’s continental shelves comprise the vast majority of living space on the planet. From the very productive, sunlit surface skin of the ocean to the cold and darkest abyssal depths, diverse communities of marine animals rise and fall within fragile and mysterious food web interactions. Human-induced changes to open ocean ecosystems have been broadly documented, raising concern about the sustainable supply of marine protein for global societies. Thus, healthy open ocean ecosystems are likely to benefit from ecosystem-based fisheries management, the main premise of which is to explicitly address the potentially deleterious implications of fishing activities on all components of underlying ecosystems, while taking into account bottom-up variability forced by the environment (e.g., Marasco et al. 2007).

Ecosystem-based fisheries management requires accurate ecological information for the principal ecosystem elements across broad temporal and spatial scales, and ultimately, the ability to model the system’s potential evolution in response to anthropogenic pressures. The size and complexity of large open ocean ecosystems unfortunately makes such a task difficult, if not impossible. The diversity of pelagic organisms is immense, and often there are multiple, shifting interactions between these organisms. Combining multiple approaches is thus necessary to monitor and understand marine food web structure and overall energy flow. Here, I highlight a subset of these methods being used in the North Pacific Subtropical Gyre and the Northern California Current ecosystems (Fig. 14).

Choy-Fig 1

Figure 14. General spatial coverage of the Hawaii Longline Fishery for the years 1990 to 2008. Qualitative fishing effort is shown by non-confidential 5° × 5° locations from the NOAA Hawaii Longline Observer Program, where darker circles indicate greater effort.

Predator-prey interactions have traditionally been derived from stomach content analysis (SCA), and are commonly used in ecosystem models to infer food web connectivity. Analysis of commercial fishery data suggests large-scale changes in catch composition, with generally smaller, lower trophic level species replacing large, apex predator species in relative abundance (Polovina et al. 2008). We used targeted SCA to investigate the diets and trophic roles of a subset of these key commercial and non-target predatory fish species sampled from the Hawaii Longline Fishery. First diet descriptions of moonfish, lancetfish and snake mackerel demonstrated unique exploitation of midwater micronekton communities, different from adult tunas and billfishes occupying a shared vertical habitat (Choy et al. 2013). Another key finding of this work was that some of these species (Alepisaurus ferox and Lampris spp.) consume surprisingly large amounts of plastic and other anthropogenic rubbish (Choy and Drazen 2013). Substantial amounts of rubbish were found in the stomachs of these presumably deeper-dwelling species, suggesting that anthropogenic litter extends into the deep sea (Fig. 15).

Choy-Fig 2

Figure 15. Examples of anthropogenic marine debris found in Lampris spp. (all but top left image) and Alepisaurus ferox (top left image) stomachs. Each of the five images represents debris found within one individual stomach. Scale bars are 1 cm. From Choy and Drazen 2013.

Temporal and taxonomic resolution derived from SCA is not easily replicated with more modern approaches, but there are key methodological drawbacks. SCA is highly labour intensive, with the stomach content representing only the most recently eaten meal, where long-term feeding habits cannot be integrated without extensive sample sizes. Furthermore, varying rates of digestion can inhibit genus or species-level identification of prey, biasing results towards groups that are more difficult to digest or easier to identify from remains. Alternatively, measuring the natural abundances of elements relating to trophic interactions have proved to greatly augment the utility of SCA, integrating underlying spatial and temporal variability expected from complex open ocean food web pathways. The second food web approach we utilized consists of a suite of biochemical tracers, including bulk and compound-specific stable isotopes, trace metals, and fatty acids.

The isotopic composition of a consumer reflects that of its prey and/or primary nutrient source in a predictable manner. At each trophic level, an increase of ~3‰ has been observed in the bulk tissue δ15N values of many consumers, while δ13C values change little within marine food webs, ~1‰ per trophic level (e.g., Fig. 16). These bulk stable isotope measurements augment snapshot trophic interactions inferred from SCA by integrating a longer feeding period reflecting the turnover rates of the tissue(s) analyzed. We utilized both bulk and compound-specific stable isotope approaches to characterize the trophic structure of epipelagic, mesopelagic and upper bathypelagic fish consumers, as well as to partition the importance of particulate organic matter and zooplankton basal food resources (Hannides et al. 2013; Choy et al. 2015). With this work we provide new evidence that large open ocean predatory fishes and micronekton species access a food web fueled by particles formed in surface waters but that are highly modified by microbes as they slowly settle into cold, dark deep waters.

Choy-Fig 3

Figure 16. Biplot of mean bulk tissue δ15N and δ13C values for 23 species of large pelagic (empty circles) and micronekton fishes (filled squares) from the North Pacific Subtropical Gyre ecosystem. Error bars are standard error. From Choy et al. 2015.

In summary, by combining multiple approaches tremendous insights have been gained into how open ocean ecosystems function and how marine communities are connected across vast vertical areas. However, open ocean ecosystems remain vastly understudied, and growing scientific evidence suggests that human-induced ecosystem changes are more widespread than previously acknowledged. To assure proper delivery of highly valuable open ocean ecosystem services into the future, continued and comprehensive monitoring and synthesis of these ecosystems is critical.


  • Choy C.A., Popp B.N., Hannides C.C.S., Drazen J.C. 2015. Trophic structure and food resources of epipelagic and mesopelagic fishes in the North Pacific Subtropical Gyre ecosystem inferred from nitrogen isotopic compositions. Limnology and Oceanography 60: 1156-1171.
  • Choy C.A., Portner E., Iwane M., Drazen J.C.. 2013. Diets of five important predatory mesopelagic fishes of the central North Pacific, Marine Ecology Progress Series 492: 169-184.
  • Choy C.A. and Drazen J.C. 2013. Plastic for dinner? Observations of frequent debris ingestion by pelagic predatory fishes from the central North Pacific. Marine Ecology Progress Series 485: 155-163.
  • Hannides C.C.S., Popp B.N., Choy C.A., Drazen J.C. 2013. Midwater zooplankton and suspended particle dynamics in the North Pacific Subtropical Gyre: a stable isotope perspective. Limnology and Oceanography 58: 1931-1946.
  • Marasco R.J., Goodman D., Grimes C.B., Lawson P.W., Punt A.E., Quinn Ii T.J. 2007. Ecosystem-based fisheries management: some practical suggestions. Can. J. Fish. Aquat. Sci. 64: 928-939.
  • Polovina J.J., Abecassis M., Howell E.A., Woodworth P. 2009. Increases in the relative abundance of mid-trophic level fishes concurrent with declines in apex predators in the subtropical North Pacific, 1996-2006. Fish. Bull. 107: 523-531.
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A curious case of dissolved oxygen and striped marlin

Chi Hin (Tim) Lam

Large Pelagics Research Center (www.tunalab.org), School for the Environment, University of Massachusetts, Boston, MA, USA

 For all things striped marlin, billfishes and tuna, contact Tim at: tim.lam@umb.edu


Striped marlin is prized all over the world as an acrobatic and feisty opponent to any sport fisher, and represents a commercially important catch in some countries and a delicacy for the Asian sashimi market. Found throughout the Pacific and Indian Oceans, striped marlin stays mostly within the mixed layer and near the sea surface, where sun-lit waters provide the best conditions for this visual predator to chase down sardines, anchovies and mackerels. The preference of striped marlin and their billfish cousins for the epipelagic zone sometimes makes them seem too simple a research subject for many fish biologists. It is not uncommon to hear seasoned scientists remarking about how “boring” billfish are as they just hang out near the surface. Our latest research findings in Fisheries Research (166:80-91) begs to differ and invites a rethink of the physiological capabilities of billfishes.

Our aim was to understand the population connectivity of striped marlin in various parts of the Pacific Ocean. Striped marlin occupy the widest latitudinal range of all Pacific billfishes and genetics has identified three distinct stocks: North Pacific, Southwest Pacific and Southeast Pacific (Purcell and Edmands 2011). With generous support from the late Paxson “Packy” Offield, an International Game Fish Association Fishing Hall of Fame Inductee, we deployed 248 popup satellite archival tags (PSATs) at eight locations from Australia to Mexico over a five-year period (Fig. 17). PSAT technology enables satellite transmission of collected depth, temperature and light records, without needing to recover the tags, which can be problematic in the vast Pacific Ocean. This is the largest billfish PSAT study ever conducted; only bluefin tuna has the commercial and conservation clout to attract enough funding to deploy more PSATs.

A striped marlin carrying a popup satellite archival tag (PSAT).

Figure 17.  A striped marlin carrying a popup satellite archival tag (PSAT). 

Although the results did not reveal cross-basin migrations, they showed extensive regional mixing. Moreover, they revealed a west-east deepening in maximum swimming depth across the Pacific (Fig. 18), which can be best explained by Australia and New Zealand having a deeper mixed layer, warmer temperatures and higher dissolved oxygen concentrations in the water column. Dissolved oxygen (DO), in particular, is known as a constraint to vertical habitat range for pelagic fishes (Prince et al. 2010). DO concentrations of below 3.5 ml L-1 are widely regarded as the physiological limit to billfishes, as they are thought to avoid any “suffocating” conditions. Concerns over increasing deoxygenation of the upper oceans under climate change have heightened the importance of DO for pelagic fishes (Stramma et al. 2012). This is where our striped marlin story gets interesting, thanks to tag data recovered off Mexican waters.

Lam-Fig 2

Figure 18. Maximum depth of striped marlin. Tag observations are binned to generate average values in a 1°x 1° grid and plotted in false colour. Contours are generated from a generalized additive mixed model. The maximum depth deepened from west to east across the longitudinal range of tagged individuals.

Off Baja California in Mexico, striped marlin inside the Gulf of California often dived about 20m deeper than when they were outside the Gulf (Fig. 19). They spent most of the day in the well-lit and oxygenated mixed layer, but about 5-10% of their day was spent at depths where dissolved oxygen concentrations were less than 3.5 ml L-1, and sometimes below 1 ml L-1. This is contrary to the established understanding of a strict oxygen constraint for billfish. Interestingly, striped marlin performed many “bounce” dives over the course of a day, for several days at a time (Fig. 20). Such behaviour has only previously been observed in tunas (Schaefer et al. 2009) and sharks (Nasby-Lucas et al. 2009), and is interpreted as the animal scanning up and down the water column for potential prey. However, what has not been recorded in the literature is that the depths of the bounce dives coincide with low DO concentrations. Striped marlin appeared to be attracted by the boundary conditions caused by low DO, much like the conditions present at the bottom of the mixed layer (Fig. 20). Our results highlight the diversity that exists in billfish diving behaviours, which has thus far been overlooked. Short-term tolerance for low dissolved oxygen concentrations, often occurring in tandem with cold temperatures, means that striped marlin are able to exploit enhanced foraging opportunities at the vertical or horizontal boundaries in the ocean. Furthermore, their preference for the epipelagic zone suggests that they are able to maintain a routine oxygen surplus, thus giving them an advantage over prey species whose vertical range might be “squeezed” by the expanding oxygen minimum zones.

Lam-Fig 3

Figure 19. Water column utilization by striped marlin in the Gulf of California.  Striped marlin spent between 5-10 % of their day in waters with dissolved oxygen (DO) of <3.5 ml L-1. A sharp decline in DO occurred at depths below the second chlorophyll maximum, at around 50 m.

Lam-Fig 4

Figure 20. Water column properties and depth-temperature time series from a striped marlin exhibiting bounce dive behaviour in the Gulf of California. One day within the week is magnified to show the constant swimming up and down the water column. This behaviour can repeat over the course of a few days, punctuated by shallow nighttime swimming. 

It is not only fish that school into near-surface bait balls that are prey for striped marlin. Gut content analyses showed squids are also a dominant prey group, especially when striped marlin are off the continental shelf (Shimose et al. 2010). Squid could hold the key to understanding the motivation behind their bounce diving behaviour, even though PSAT data from striped marlin and squid only show a small overlap around 150 m (Bazzino et al. 2010). To investigate this further, we are teaming up with Drs. Anela Choy and Bruce Robison of Monterey Bay Aquarium Research Institute. Using their extensive cruise data from the Gulf of California, we will investigate vertical overlap and occurrence of prey species with top predators, from striped marlin to sharks and whales. This work will offer unique insights into the ecosystem dynamics of co-occurring pelagic predators and prey, highlighting vertical habitat use of predators relative to environmental factors. Understanding the vertical habitat characteristics of striped marlin is necessary to evaluate their vulnerability to fishing gear, to identify realistic bycatch reduction measures, and to provide spatially explicit information for stock assessments to ensure a sustainable future for this iconic billfish.

Learn more

Lam C. H., Kiefer D., Domeier M.L. 2015. Habitat utilization of striped marlin in the Pacific Ocean. Fisheries Research 166: 80-91. http://dx.doi.org/10.1016/j.fishres.2015.01.010


  • Bazzino G., Gilly W.F., Markaida U., Salinas-Zavala C.A., Ramos-Castillejos J. 2010. Horizontal movements, vertical-habitat utilization and diet of the jumbo squid (Dosidicus gigas) in the Pacific Ocean off Baja California Sur, Mexico. Prog. Oceanogr. 86(1–2): 59-71.
  • Nasby-Lucas N., Dewar H., Lam C.H., Goldman K.J., Domeier M.L. 2009. White Shark Offshore Habitat: A Behavioral and Environmental Characterization of the Eastern Pacific Shared Offshore Foraging Area. PloS ONE 4(12): e8163:8161-8114.
  • Prince E.D., Luo J.G., Goodyear C.P., Hoolihan J.P., Snodgrass D., Orbesen E.S., Serafy J.E., Ortiz M., Schirripa M.J. 2010. Ocean scale hypoxia-based habitat compression of Atlantic istiophorid billfishes. Fish. Oceanogr. 19(6): 448-462.
  • Purcell C.M.,Edmands S. 2011. Resolving the genetic structure of striped marlin, Kajikia audax, in the Pacific Ocean through spatial and temporal sampling of adult and immature fish. Can. J. Fish. Aquat. Sci.  68(11): 1861-1875.
  • Schaefer K.M., Fuller D.W., Block B.A. 2009. Vertical movements and habitat utilization of skipjack (Katsuwonus pelamis), yellowfin (Thunnus albacares), and bigeye (Thunnus obesus) tunas in the equatorial Eastern Pacific Ocean, ascertained through archival tag data. In: Nielsen J.L., Arrizabalaga H., Fragoso N., Hobday A., Lutcavage M., Sibert J., Eds. Tagging and Tracking of Marine Animals with Electronic Devices, Reviews: Methods and Technologies in Fish Biology and Fisheries 9: Springer. p.121-144.
  • Shimose T., Yokawa K., Saito H. 2010. Habitat and food partitioning of billfishes (Xiphioidei). J. Fish. Biol. 76(10): 2418-2433.
  • Stramma L., Prince E.D., Schmidtko, S., Luo J., Hoolihan J.P., Visbeck M., Wallace D.W.R., Brandt P., Kortzinger A. 2012. Expansion of oxygen minimum zones may reduce available habitat for tropical pelagic fishes. Nature Climate Change 2(1): 33-37.
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Resolving the role of mesopelagics in Southern Ocean food webs

Rowan Trebilco1, Andrea Walters2, Jess Melbourne-Thomas1,3, Andrew Constable1,3

1. Antarctic Climate and Ecosystems Cooperative Research Centre, University of Tasmania

2. Antarctic Gateway Partnership, University of Tasmania

3. Australian Antarctic Division, Kingston, Tasmania


The Southern Ocean is rapidly changing, with future environmental changes likely to have wide-reaching consequences for the structure and function of marine ecosystems (Constable et al. 2014). In addition, the coming decades are likely to see expansion of krill fisheries, and potentially the development of other new fisheries (Kock et al. 2007). Along with well-designed observation programs, ecosystem models are the central means by which we can gain insight into what these changes will mean for the structure and function of Southern Ocean ecosystems, providing the necessary information to inform management decisions into the future.

Mesopelagic fishes and squid, together with krill, dominate mid-trophic levels in Southern Ocean ecosystems, comprising the pathways by which energy from primary producers is made accessible to higher-order predators including whales, seals, penguins, flying seabirds, and large (often commercially valuable) fish (Fig. 21; Kozlov 2006, Connan et al. 2007).  The short, krill-dominated food chains are well studied and relatively well represented in ecosystem models (Hill et al. 2006, Murphy et al. 2012). However, mesopelagic fish and squid are far less well studied and represent a key area of uncertainty in current ecosystem modelling efforts (Hill et al. 2006, Murphy et al. 2012, Melbourne-Thomas et al. in prep). Knowledge of mesopelagic fish and squid has also been identified as a key uncertainty for understanding top predator trophodynamics globally (Young et al. 2015).

While food chains based on krill (“krill dominated”) and mesopelagic fishes and squids (“fish dominated”) may occur together at the same location and at the same time, distinct krill vs. fish dominated ecosystem states are identifiable (Duhamel et al. 2014, Hosie et al. 2014). In some regions, such as the Scotia Sea, changes between krill and fish dominance occur through time, with some years exhibiting a krill regime and others exhibiting a fish regime (Murphy et al. 2007a, 2007b). Elsewhere, the transition occurs over space, like on the Kerguelen Plateau where there is a south-to north shift from krill to fish dominance (Hulley et al. 2011, Nicol and Raymond 2012).  It is not clear what determines whether the krill or fish pathway dominates, nor whether the relative importance of these energy pathways will change in the future (Constable et al. 2014).

Trebilco-Fig 1

Figure 21. A generic network model representation of Southern Ocean food webs showing the two major pathways, via krill (orange) and fish (blue) by which energy makes its way from primary producers to higher trophic levels (from Melbourne-Thomas et al. in prep).

Uncertainties regarding mid- trophic levels are a major impediment to the development of conservation and management strategies (Constable et al. 2014, IMBER 2015). Within the Antarctic Climate and Ecosystems Cooperative Research Centre (ACE CRC) and through the Antarctic Gateway Partnership, together with an international network of colleagues and collaborators, we have initiated a research program that seeks to address these knowledge gaps through an integrated program of modelling and targeted field observation. This work is initially focused on the Kerguelen Plateau and the nearby continent (herein collectively referred to as the Kerguelen Axis; Fig. 22). The Kerguelen Axis represents the most important area for primary production in the southern Indian Ocean (Quéguiner et al. 2015), and is home to high-value toothfish and icefish fisheries (Constable and Welsford 2011), historical and potential future krill fisheries in the south (Nicol and Raymond 2012), and important foraging habitat for many predators (Raymond et al. 2014).

Improved model representation for mesopelagics

Given the potential importance of mesopelagic fish and squid in Southern Ocean food webs, there is a strong imperative to develop robust and informative model representations to help reduce uncertainty regarding the role played by mesopelagic taxa. Over the past decade, two modelling approaches have been developed that offer great promise in this regard. These are the Spatial Ecosystem and Population Dynamics Model (SEAPODYM) framework (Lehodey et al. 2008, 2010), and size-based ecosystem models (Law et al. 2009, Scott et al. 2014). Neither approach has yet been implemented fully in the Southern Ocean. We plan to develop implementations of both approaches for the Kerguelen Axis. 

Trebilco-Fig 2

Figure 22. The Kerguelen Plateau, with the planned track of the 2016 Kerguelen Axis voyage (red). Chlorophyll A and SST are 10-year summer climatologies (2005-2014; October-January for Chl; January for SST). White lines in top panel show the 10-year average of the maximum extent of ice (15% concentration) for the months of October (winter) and January (summer). Pink contours in bottom panel show areas of 80% overlap for important predator habitat from Raymond et al. 2014.

In addition to SEAPODYM and size-based models, implementations of Atlantis (Fulton et al. 2004), Ecopath with Ecosim (EwE; Pruvost et al. 2005), and in-house modelling approaches (Constable 2005) are also being developed with the goal of establishing an ensemble of consistent and comparable models for the region, which have model behaviours that can replicate available mesopelagic data. Once tested, these models are expected to be able to be used for the following purposes: (i) assess the foodweb effects of finfish fisheries and climate change in the region; (ii) identify and evaluate potential indicators of change, and;

(iii) understand the implications of change for higher trophic levels (marine mammals, seabirds and large fishes). Having an ensemble of models available for these tasks will enable more robust assessment of whether the outcomes are sensitive to model assumptions (Fulton and Link 2014). The different models will also enable a greater breadth of ecosystem scenarios to be evaluated (Fig. 23).

Targeted observations to understand the drivers of the krill to fish-transition

Along the Kerguelen Axis, there is a transition from krill-dominated food webs in the south to fish-dominated food webs in the north (Hulley et al. 2011, Nicol and Raymond 2012). However, there has not yet been a systematic survey along this gradient to elucidate what drives this transition. Understanding the dichotomy in food web structure along the Kerguelen Axis will enable us to make informed decisions about how environmental change and harvesting might influence Southern Ocean ecosystems into the future, and will underpin the development of evidence-based management decisions.

Trebilco-Fig 3

Figure 23. Existing data will be integrated with new field observations to develop and evaluate implementations of size-based and SEAPODYM ecosystem models. These models will form part of ensemble-of-models approach for exploring scenarios of change and evaluating the performance of indicators of change 

Five hypotheses have been identified that may explain the transition from krill to fish dominance on the Kerguelen Axis: (i) the Southern Boundary of the Antarctic Circumpolar Current (ACC) (Nicol et al. 2000); (ii) the Southern ACC Front (SACCF) (Nicol and Raymond 2012); (iii) the winter extent of sea ice (Nicol et al. 2008, Murphy et al. 2013); (iv) extent of tolerance for warmer temperatures (Hill et al. 2013); and (v) availability of diatoms, the primary food of krill (Constable et al. 2014). In January 2016, the ACE CRC will conduct a voyage with a sampling design that aims to distinguish the relative contributions of these factors (Figure 2). Understanding the krill-fish transition will be a key factor in resolving the role of mesopelagics in Southern Ocean food webs, and how it may change with changing environmental conditions and changes to fisheries into the future.



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  • Constable A.J., 2005. Implementing plausible ecosystem models for the Southern Ocean: an ecosystem, productivity, ocean, climate (EPOC) model. WG-EMM-05/ 33. CCAMLR Ecosystem Monitoring and Management Working Group. CCAMLR, Hobart, Australia.
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Where are they now?


Oceans, marine ecosystems, and society facing climate change. This is what brought 70 early career scientists and students from 26 countries together in Brest, France in August 2010. Apart from several international oceanography conferences, I had at that point in my career had little interaction with ocean researchers outside of the United States. Little did I know that four years later I would be immersed in European and Norwegian marine research and living in northern Europe.

The IMBER ClimECO2 summer school was held at the European Institute for Marine Studies (IUEM). It was organized as a series of lectures on climate change effects on the ocean, contributed oral and poster presentations, and an interesting evening of role-playing as representatives of governments, stakeholders, and scientists in the decision-making process about different climate change scenarios and the resultant socio-economic consequences (complete with Bretagne crepes and appetizers!). Here we were encouraged to discuss future ocean change, to form a cross-disciplinary network of scientists from diverse backgrounds, and to consider our research in the scope of policy and societal needs and actions.

Some months later, I was invited to attend the IMBIZO II conference in Crete, Greece. This meeting brought the international marine biogeochemistry and ecosystem community together to discuss, share, and strategize about research directions and goals for the coming years. Unlike typical meetings, IMBIZO II had dedicated time for open discourse and discussion pertaining to topics ranging from environmental elemental stoichiometry to marine food webs. This was the first of many more meetings and workshops for me in which the mold of formal talks followed by questions was broken.

Andrew King

Andrew King

Norwegian Institute for Water Research

Bergen, Norway

Fortuitously and fittingly, the 2014 Future Oceans IMBER Open Science Conference held in Bergen, Norway coincided with the start of my current position at the Norwegian Institute for Water Research (NIVA) in Bergen. There were many familiar faces at the meeting and it provided an excellent introduction to what have now become familiar landmarks – the historic Bryggen where the conference was held, Håkons Hall for the welcome reception, and Fløyen where the conference dinner was held. The theme of the meeting matched my research interests in present day and future variability in nutrient biogeochemistry.

From my perspective, these events have collectively served as an opportunity to link with fellow scientists and to learn about new approaches from an international perspective, and have ultimately fostered international cooperation (in my case, and very likely for others). I look forward to the next IMBER event to hear about and discuss new research developments, to meet new colleagues, and to re-connect with old friends.

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Comings and goings on the IMBER Scientific Steering Committee

The goings

As the end of the year approaches, several people will be rotating off the IMBER Scientific Steering Committee (SSC).

We would like to thank the following people for their superb contribution to the IMBER SSC for at least the past six years:

  • Alberto Piola –Executive Committee member (Argentina)
  • Claudio Campagna (Argentina)
  • Ken Drinkwater - Executive Committee member (Norway)
  • Su Meil Liu (China), and last, but definitely not least,
  • Eileen Hofmann, who has tirelessly chaired the IMBER SSC since 2010. Her guidance and wisdom have been invaluable, and on an almost daily basis she went way beyond the call of duty and her bedtime! We are delighted that she will remain on the SSC as Past-Chair for another year.
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The comings

We are very pleased to welcome the following people who will join the IMBER SSC from 1 January 2016:

Ingrid van Puten
  Ingrid van Putten is a human behaviour modeller with the Ecosystems Modelling team at the Commonwealth Scientific & Industrial Research Organisation (CSIRO) in Hobart, Australia. Her research focuses on modelling the interaction of social and economic behaviour with the biophysical environment, and understanding coupled social-ecological systems. Because the complexity of the bio-physical sphere is mirrored in social and economic systems, she focuses on tools that effectively model social and economic data, and provide the optimum level of complexity for human behaviour models.
Cisco Werner
  Cisco Werner is Director of the Southwest Fisheries Science Center at NOAA in La Jolla, CA, USA. He brings to IMBER a blend of marine ecosystem research, ranging from the structure and function of marine ecosystems, ocean circulation physics, and the development and implementation of ocean and coastal observing and forecasting systems. He also has considerable science management expertise, and a wealth of experience in international science collaboration having chaired the GLOBEC SSC. His research experience within physical-biological modelling provides him with an excellent background for the integration of the biogeochemistry and the marine ecosystems. 
Ying Wu
  Ying Wu is a professor at the State Kay Laboratory of Estuarine and Coastal Research, East China Normal University in Shanghai, China. Her research focuses on the composition and biogeochemistry of organic matter from source to sink, using isotopes and biomarkers to trace the sources, fluxes, cycles and transformations. Ying has been actively involved in Chinese GLOBEC/IMBER projects since the very beginning. She is currently leading one of the sub-projects of a large national project that aims to evaluate the role of migrating behaviour of mesopelagic fishes on marine carbon cycle.
Dan Costa
  Dan Costa is a Distinguished Professor of Ecology and Evolutionary Biology at the University of California at Santa Cruz, CA, USA. His main research focus is the ecology and physiology of marine mammals and seabirds, their adaptations to the marine environment, especially the movements, foraging ecology and energetics of pinnipeds and seabirds. His current work is aimed at recording the movement and distribution patterns of marine mammals and seabirds to understand their habitat needs. This work is helping to identify biodiversity hotspots and the factors that create them.
Carol Robinson
  Carol Robinson will take over the reins of the IMBER SSC chair from Eileen Hofmann. Carol is not a new comer to IMBER, having served two terms as an IMBER SSC and Executive Committee member. We are thrilled to welcome her back! Carol´s professional title is “Reader in Marine Biogeochemistry” at the School of Environmental Sciences at the University of East Anglia in Norwich, UK, where she leads a team which studies the role of marine bacteria, phytoplankton and zooplankton in the global cycling of carbon and oxygen, and how this varies in space and time and with changing environmental conditions, such as increasing nutrient supply, temperature and carbon dioxide and decreasing dissolved oxygen. 
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IMBER future events

7th CJK

The symposium will be held from 24-26 March 2016 in Jeju, Korea. Join us to present your work and discuss the methodological and empirical challenges involved in addressing the symposium themes. Visit our website for all the details, and submit an abstract  by 5 January 2016.

We look forward to welcoming you in Jeju!

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IMBER ClimEco5 Summer School (10-17 August 2016, Natal, Brazil)

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  • Williams N.L., Feely R.A., Sabine C.L., Dickson A.G., Swift J.H., Talley L.D. & Russell J.L. 2015. Quantifying anthropogenic carbon inventory changes in the Pacific sector of the Southern Ocean. Marine Chemistry 174: 147-160. doi:10.1016/j.marchem.2015.06.015
  • Yang Y., Hansson L. & Gattuso J.-P. 2015. Data compilation on the biological response to ocean acidification: an update. Earth Syst. Sci. Data Discuss. 8: 889-912. DOI: 10.5194/essdd-8-889-2015 
  • Zark M., Riebesell U. & Dittmar T. 2015. Effects of ocean acidification on marine dissolved organic matter are not detectable over the succession of phytoplankton blooms. Science Advances 1(9): e15005. doi: 10.1126/sciadv.1500531
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Meeting Calendar



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And last but not least, many thanks to to everyone in the IMBER community for their contribution to the project during the year. We have made good progress and look forward to kicking off the new year with a new science plan developed by all of you.

We would not be able to do anything without our sponsors - SCOR and IGBP, the Institute of Marine Research and the Research Council of Norway, and the East China Normal University - grateful thanks for the all the support.

We are very sad to see IGBP go after 10 years of guidance and funding. ....see you on the other side when we become a core project of Future Earth and SCOR.

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( Photo: Paradise Harbour, Antarctica by Liam Quinn  CC BY-SA 2.0 )