The forgotten feeding ground: patterns in seasonal and depth-specific food intake of adult cod Gadus morhua in the western Baltic Sea.

This study presents the diet composition of western Baltic cod Gadus morhua based on 3150 stomachs sampled year-round between 2016 and 2017 using angling, gillnetting and bottom trawling, which enhanced the spatio-temporal coverage of cod habitats. Cod diet composition in shallow areas (<20 m depth) was dominated by benthic invertebrate species, mainly the common shore crab Carcinus maneas. Compared to historic diet data from the 1960s and 1980s (limited to depth >20 m), the contribution of herring Clupea harengus decreased and round goby Neogobius melanostomus occurred as a new prey species. Statistical modelling revealed significant relationships between diet composition, catch depth, fish length and season. Generalized additive modelling identified a negative relationship between catch depth and stomach content weight, suggesting reduced food intake in winter when cod use deeper areas for spawning and during peak summer when cod tend to avoid high water temperatures. The results of this study highlight the importance of shallow coastal areas as major feeding habitats of adult cod in the western Baltic Sea, which were previously unknown because samples were restricted to deeper trawlable areas. The results strongly suggest that historic stomach analyses overestimated the role of forage fish and underestimated the role of invertebrate prey. Eventually, this study shows the importance of a comprehensive habitat coverage for unbiased stomach sampling programmes to provide a more reliable estimation of top predator diet, a key information for food web analyses and multispecies models.

Hydrostatic pressure affects selective tidal stream transport in the North Sea brown shrimp (Crangon crangon).

The brown shrimp (Crangon crangon) is a highly abundant invertebrate in the North Sea, with its life cycle stages ranging from deep offshore spawning to shallow onshore nursery areas. To overcome the long distances between these two habitats, brown shrimp are suspected to use selective tidal stream transport (STST), moving with the cyclic tide currents towards their preferred water depths. However, it is not known which stimulus actually triggers STST behavior in brown shrimp. In this work, we determined the influence of different hyperbaric pressures on STST behavior of juvenile brown shrimp. Brown shrimp activity was recorded in a hyperbaric pressure chamber that supplied constant and dynamic pressure conditions simulating different depths, with and without a tidal cycle. Subsequent wavelet and Fourier analysis were performed to determine the periodicity in the activity data. The results of the experiments show that STST behavior in brown shrimp varies with pressure and therefore with depth. We further show that STST behavior can be initiated by cyclic pressure changes. However, an interaction with one or more other environmental triggers remains possible. Furthermore, a security ebb-tide activity was identified that may serve to avoid potential stranding in shallow waters and is 'remembered' by shrimp for about 1.5 days without contact with tidal triggers.

Tracing growth patterns in cod (Gadus morhua L.) using bioenergetic modelling.

Understanding individual growth in commercially exploited fish populations is key to successful stock assessment and informed ecosystem-based fisheries management. Traditionally, growth rates in marine fish are estimated using otolith age-readings in combination with age-length relationships from field samples, or tag-recapture field experiments. However, for some species, otolith-based approaches have been proven unreliable and tag-recapture experiments suffer from high working effort and costs as well as low recapture rates. An important alternative approach for estimating fish growth is represented by bioenergetic modelling which in addition to pure growth estimation can provide valuable insights into the processes leading to temporal growth changes resulting from environmental and related behavioural changes. We here developed an individual-based bioenergetic model for Western Baltic cod (Gadus morhua), traditionally a commercially important fish species that however collapsed recently and likely suffers from climate change effects. Western Baltic cod is an ideal case study for bioenergetic modelling because of recently gained in-situ process knowledge on spatial distribution and feeding behaviour based on highly resolved data on stomachs and fish distribution. Additionally, physiological processes such as gastric evacuation, consumption, net-conversion efficiency and metabolic rates have been well studied for cod in laboratory experiments. Our model reliably reproduced seasonal growth patterns observed in the field. Importantly, our bioenergetic modelling approach implementing depth-use patterns and food intake allowed us to explain the potentially detrimental effect summer heat periods have on the growth of Western Baltic cod that likely will increasingly occur in the future. Hence, our model simulations highlighted a potential mechanism on how warming due to climate change affects the growth of a key species that may apply for similar environments elsewhere.

Alternative reproductive tactics are associated with sperm performance in invasive round goby from two different salinity environments.

During male-male competition, evolution can favor alternative reproductive tactics. This often results in a dominant morph that holds a resource, such as a nest for egg laying, which competes with a smaller sneaker morph that reproduces by stealing fertilizations. The salinity environment can influence male growth rates, for example, via osmoregulatory costs, which in turn may influence the use of sneaker tactics for small males competing for mating opportunities. Salinity can also affect sperm directly; however, little is known of how salinity influences sneaker tactics through sperm performance. We sampled males of the invasive round goby (Neogobius melanostomus) from two environments, a freshwater river and a brackish estuary. This fish has two male morphs: nest-holding dark males and non-nest-holding light males. We examined the role of water salinity of 0, 8, and 16 on sperm performance and found that for estuarine males, a salinity of 0 reduced sperm velocity compared to a salinity of 8 and 16. Riverine males had low velocity in all salinities. Sperm viability also decreased by over 30% in 0 salinity, compared to 8 and 16, for fish from both environments. Gobies produce ejaculate contents in specialized glands that could in theory shield sperm in an adverse environment. However, gland contents did not improve sperm performance in our tests. Body mass and age estimates indicate that riverine males invested more in somatic growth compared to estuarine males. Estuarine light morph males had a high enough gonadosomatic index to indicate sneaker tactics. We propose that when sperm performance is low, such as for the riverine males, sneaker tactics are ineffective and will be selected against or phenotypically suppressed. Instead, we interpret the increased investment in somatic growth found in riverine males as a life-history decision that is advantageous when defending a nest in the next reproductive season.

Predation risk triggers copepod small-scale behavior in the Baltic Sea.

Predators not only have direct impact on biomass but also indirect, non-consumptive effects on the behavior their prey organisms. A characteristic response of zooplankton in aquatic ecosystems is predator avoidance by diel vertical migration (DVM), a behavior which is well studied on the population level. A wide range of behavioral diversity and plasticity has been observed both between- as well as within-species and, hence, investigating predator-prey interactions at the individual level seems therefore essential for a better understanding of zooplankton dynamics. Here we applied an underwater imaging instrument, the video plankton recorder (VPR), which allows the non-invasive investigation of individual, diel adaptive behavior of zooplankton in response to predators in the natural oceanic environment, providing a finely resolved and continuous documentation of the organisms' vertical distribution. Combing observations of copepod individuals observed with the VPR and hydroacoustic estimates of predatory fish biomass, we here show (i) a small-scale DVM of ovigerous Pseudocalanus acuspes females in response to its main predators, (ii) in-situ observations of a direct short-term reaction of the prey to the arrival of the predator and (iii) in-situ evidence of pronounced individual variation in this adaptive behavior with potentially strong effects on individual performance and ecosystem functioning.

Metabolic costs of spontaneous swimming in Sprattus sprattus L., at different water temperatures.

Oxygen uptake (MO2; mgO2 fish-1h-1) of fish groups was measured at temperatures between 10-19°C in an intermittent-flow respirometer to quantify the metabolic costs of spontaneous swimming patterns in the small clupeid Sprattus sprattus. Movements of individual fish within the school were tracked automatically during respirometry. Oxygen uptake was then related to mean swimming speeds and the number of sharp turns (>90°), which are common behavioural elements of spontaneous swimming in clupeid fish. Different possible model formulations for describing the relationship between respiration and swimming patterns were compared via the AIC. The final model revealed that costs for sharp turns at a frequency of 1 s-1 doubled the metabolic costs compared to those with zero turns but with likewise a moderate swimming speed of 0.28 body length -1. The cost for swimming doubled if the swimming speed was doubled from 0.28 to 0.56 BLs-1 but increased by a factor of 4.5 if tripled to 0.84 BLs-1. Costs for transport were minimal at a speed of 0.4 body lengths s-1 at all temperatures. New basic input parameters to estimate energy losses during spontaneous movements, which occur typically during foraging in this small pelagic fish, are provided.

Sensitivity of multispecies maximum sustainable yields to trends in the top (marine mammals) and bottom (primary production) compartments of the southern North Sea food-web.

In marine ecosystems, maximum sustainable yield considerations are affected by any substantial changes that occur in the top and bottom compartments of the food-web. This study explores how the southern North Sea's fisheries may need to adjust their fishing efforts to maintain optimum yields of sole, plaice, cod and brown shrimps under increased marine mammal populations and a reduced primary productivity. We constructed plausible scenarios of ongoing food-web changes using the results of Bayesian age-structured population models to estimate carrying capacities of harbour porpoises (Phocoena phocoena) and grey seals (Halichoerus grypus). Losses in primary productivity were predicted by lower trophic level ecosystem models. These scenarios were implemented in a food-web model of the southern North Sea. For each scenario, we sought mixed-fleet fishing efforts that would deliver maximum yields of sole, plaice, cod and brown shrimp combined. We also did so for a baseline run with unaltered mammal and primary production, and compared the differences in optimal fishing strategies, predicted yields, and states of the stocks between the scenarios. We found stocks and yields to be far more sensitive to changes in primary productivity than to increased marine mammal predation. The latter predominantly impacted cod, and even benefitted brown shrimps compared to the baseline run. Under 30% reduced primary productivity, fishing efforts had to be reduced by 50% to still provide maximum yields, whereas the marine mammal scenario induced no need to adjust the fishing regime. This draws attention to the potential gains of incorporating bottom-up processes into long-term management considerations, while marine mammal predation may be less of a concern, in particular for flatfish fisheries in the North Sea, and may even benefit shrimp trawlers because of reduced predation on shrimp from fish predators.

Predictive framework for codend size selection of brown shrimp (Crangon crangon) in the North Sea beam-trawl fishery.

The brown shrimp (Crangon crangon) fishery is of great socio-economic importance to coastal communities on the North Sea. The fishery is exploited by beam trawlers often using codends with very small mesh sizes, leading to concerns about catch rates of undersized shrimp. However, little information is available on codend size selection, making it difficult to provide scientifically based advice on alternative codend designs. Therefore, this study establishes a predictive framework for codend size selection of brown shrimp, based on a large selectivity dataset from 33 different codend designs tested during four experimental fishing cruises, during which more than 350,000 brown shrimp were length measured. Predictions by the framework confirm concerns about the exploitation pattern in the fishery, because the retention probability of undersized shrimp reaches 95% with the currently applied designs. The framework predictions allow the exploration of obtainable exploitation patterns depending on codend design. For example, increasing codend mesh size to 25-29 mm would reduce the retention rate of undersized shrimp to a maximum of 50%, depending on codend mesh type.

A spatially explicit risk approach to support marine spatial planning in the German EEZ.

An ecosystem approach to marine spatial planning (MSP) promotes sustainable development by organizing human activities in a geo-spatial and temporal context. (1) This study develops and tests a spatially explicit risk assessment to support MSP. Using the German exclusive economic zone (EEZ) of the North Sea as a case study area, current and future spatial management scenarios are assessed. (2) Different tools are linked in order to carry out a comprehensive spatial risk assessment of current and future spatial management scenarios for ecologic and economic ecosystem components, i.e. Pleuronectes platessa nursery grounds. With the identification of key inputs and outputs the suitability of each tool is tested. (3) Here, the procedure as well as the main findings of the spatially explicit risk approach are summarised to demonstrate the applicability of the framework and the need for an ecosystem approach to risk management techniques using geo-spatial tools.

Comment on "Impacts of biodiversity loss on ocean ecosystem services".

Worm et al. (Research Articles, 3 November 2006, p. 787) investigated the importance of biodiversity to marine ecosystem services across temporal and spatial scales. In projecting the extent of future fisheries collapse, we argue that the authors inappropriately extrapolated beyond their available observations and used data on marine reserves and fishery closures that are not representative of global fisheries.