Marine ecosystems evolve under many interconnected and area-specific pressures. To fulfil society's intensifying and diversifying needs while ensuring ecologically sustainable development, more effective marine spatial planning and broader-scope management of marine resources is necessary. Integrated ecological-economic fisheries models (IEEFMs) of marine systems are needed to evaluate impacts and sustainability of potential management actions and understand, and anticipate ecological, economic and social dynamics at a range of scales from local to national and regional. To make these models most effective, it is important to determine how model characteristics and methods of communicating results influence the model implementation, the nature of the advice that can be provided and the impact on decisions taken by managers. This article presents a global review and comparative evaluation of 35 IEEFMs applied to marine fisheries and marine ecosystem resources to identify the characteristics that determine their usefulness, effectiveness and implementation. The focus is on fully integrated models that allow for feedbacks between ecological and human processes although not all the models reviewed achieve that. Modellers must invest more time to make models user friendly and to participate in management fora where models and model results can be explained and discussed. Such involvement is beneficial to all parties, leading to improvement of models and more effective implementation of advice, but demands substantial resources which must be built into the governance process. It takes time to develop effective processes for using IEEFMs requiring a long-term commitment to integrating multidisciplinary modelling advice into management decision-making. K E Y W O R D Sbio-economic models, comparative model evaluation, fisheries management advice, integrated ecological-economic fisheries models, marine spatial planning and cross-sector management, performance criteria and scales and risks, use and acceptance and implementation and communication and flexibility and complexity | INTRODUCTIONThere is a growing need for tools to evaluate policies and assess tradeoffs in management of marine resources and provision of ecosystem services such as fishing, aquaculture, renewable energy, shipping, conservation and recreation (Cormier, Kannen, Elliott, & Hall, 2015;Degnbol & Wilson, 2008;EU 2014;Langlois, Fréon, Steyer, Delgenés, & Hélias, 2014;White et al., 2012). It is necessary to elaborate and apply common principles and broader, interdisciplinary management evaluation in the use of marine space involving several types of activities and sectors Soma et al., 2013;Stelzenmüller et al., 2013;Sundblad et al., 2014). Policymakers need to know the costs and benefits of conserving ecosystem goods and services to manage them sustainably. Moreover, according to an ecosystembased approach to management, specific pressures, associated uncertainties and risks need to be taken into account (Douvere, 2008;Ehler & Douvere, 2009;Gi...

Abstract. Eastern boundary upwelling systems (EBUS) are among the most productive marine ecosystems on Earth. The production of organic material is fueled by upwelling of nutrient-rich deep waters and high incident light at the sea surface. However, biotic and abiotic factors can modify surface production and related biogeochemical processes. Determining these factors is important because EBUS are considered hotspots of climate change, and reliable predictions of their future functioning requires understanding of the mechanisms driving the biogeochemical cycles therein. In this field experiment, we used in situ mesocosms as tools to improve our mechanistic understanding of processes controlling organic matter cycling in the coastal Peruvian upwelling system. Eight mesocosms, each with a volume of ∼55 m3, were deployed for 50 d ∼6 km off Callao (12∘ S) during austral summer 2017, coinciding with a coastal El Niño phase. After mesocosm deployment, we collected subsurface waters at two different locations in the regional oxygen minimum zone (OMZ) and injected these into four mesocosms (mixing ratio ≈1.5 : 1 mesocosm: OMZ water). The focus of this paper is on temporal developments of organic matter production, export, and stoichiometry in the individual mesocosms. The mesocosm phytoplankton communities were initially dominated by diatoms but shifted towards a pronounced dominance of the mixotrophic dinoflagellate (Akashiwo sanguinea) when inorganic nitrogen was exhausted in surface layers. The community shift coincided with a short-term increase in production during the A. sanguinea bloom, which left a pronounced imprint on organic matter C : N : P stoichiometry. However, C, N, and P export fluxes did not increase because A. sanguinea persisted in the water column and did not sink out during the experiment. Accordingly, export fluxes during the study were decoupled from surface production and sustained by the remaining plankton community. Overall, biogeochemical pools and fluxes were surprisingly constant for most of the experiment. We explain this constancy by light limitation through self-shading by phytoplankton and by inorganic nitrogen limitation which constrained phytoplankton growth. Thus, gain and loss processes remained balanced and there were few opportunities for blooms, which represents an event where the system becomes unbalanced. Overall, our mesocosm study revealed some key links between ecological and biogeochemical processes for one of the most economically important regions in the oceans.

Central-place foraging in large seabird colonies leads to high levels of intra-specific competition for food resources, often resulting in between-colony spatial segregation. However, little is known about within-colony variation in foraging behaviour that may arise from breeding locations. Using little penguins Eudyptula minor from a large colony (ca. 32 000 individuals) on Phillip Island (Australia), we present a novel approach combining GPS, diving, acceleration and bathymetry data. We investigated within-colony variation in 3-dimensional distribution of prey encounters and its consequences for foraging behaviour and breeding success. Over 1 breeding season, we simultaneously tracked 63 little penguins from 2 breeding sites located ~2 km apart and monitored their breeding success. We recorded 58 452 dives, of which 11 992 had prey encounter events associated. Results revealed strong spatial foraging segregation between sites throughout the breeding season and differences between sites in prey encounter depth during chick-rearing (mean ± SE, 11.8 ± 0.2 m vs. 17.3 ± 0.3 m). Birds from one site foraged in deeper waters and apparently experienced higher levels of competition, resulting in lower prey encounter rates and lower foraging efficiency (i.e. body mass gain after foraging trips), hence these birds seemed less successful. However, breeding success was high (>1.5 chicks fledged per pair) at both sites, indicating that food was not a limiting factor that year. Nonetheless, breeding success records over the last 12 yr showed that the site where birds foraged at deeper depths produced fewer chicks. Our findings highlight the importance of understanding small-scale spatial segregation to capture foraging behaviour variation within large seabird colonies.

The epoch of the Anthropocene, a period during which human activity has been the dominant influence on climate and the environment, has witnessed a decline in oxygen concentrations and an expansion of oxygendepleted environments in both coastal and open ocean systems since the middle of the 20th century. This paper provides a review of system-specific drivers of low oxygen in a range of case studies representing marine systems in the open ocean, on continental shelves, in enclosed seas and in the coastal environment. Identification of similar and contrasting responses within and across system types and corresponding oxygen regimes is shown to be informative both in understanding and isolating key controlling processes and provides a sound basis for predicting change under anticipated future conditions. Case studies were selected to achieve a balance in system diversity and global coverage. Each case study describes system attributes, including the present-day oxygen environment and known trends in oxygen concentrations over time. Central to each case study is the identification of the physical and biogeochemical processes that determine oxygen concentrations through the tradeoff between ventilation and respiration. Spatial distributions of oxygen and time series of oxygen data provide the opportunity to identify trends in oxygen availability and have allowed various drivers of low oxygen to be distinguished through correlative and causative relationships. Deoxygenation results from a complex interplay of hydrographic and biogeochemical processes and the superposition of these processes, some additive and others subtractive, makes attribution to any particular driver challenging. System-specific models are therefore required to achieve a quantitative understanding of these processes and of the feedbacks between processes at varying scales.

Landscape conversion by humans may have detrimental effects on animal populations inhabiting managed ecosystems, but human-altered areas may also provide suitable environments for tolerant species. We investigated the spatial ecology of a highly mobile nocturnal avian species–the red-necked nightjar (Caprimulgus ruficollis)–in two contrastingly managed areas in Southwestern Spain to provide management recommendations for species having multiple habitat requirements. Based on habitat use by radiotagged nightjars, we created maps of functional heterogeneity in both areas so that the movements of breeding individuals could be modeled using least-cost path analyses. In both the natural and the managed area, nightjars used remnants of native shrublands as nesting sites, while pinewood patches (either newly planted or natural mature) and roads were selected as roosting and foraging habitats, respectively. Although the fraction of functional habitat was held relatively constant (60.9% vs. 74.1% in the natural and the managed area, respectively), landscape configuration changed noticeably. As a result, least-cost routes (summed linear distances) from nest locations to the nearest roost and foraging sites were three times larger in the natural than in the managed area (mean ± SE: 1356±76 m vs. 439±32 m). It seems likely that the increased proximity of functional habitats in the managed area relative to the natural one is underlying the significantly higher abundances of nightjars observed therein, where breeders should travel shorter distances to link together essential resources, thus likely reducing their energy expenditure and mortality risks. Our results suggest that landscape configuration, but not habitat availability, is responsible for the observed differences between the natural and the managed area in the abundance and movements of breeding nightjars, although no effect on body condition was detected. Agricultural landscapes could be moderately managed to preserve small native remnants and to favor the juxtaposition of functional habitats to benefit those farm species relying on patchy resources.