Stefan Neuenfeldt

Some Atlantic cod Gadus morhua in the Baltic Sea visit hypoxic water briefly but often

Individual behaviour of Atlantic cod Gadus morhua in the presence of hypoxic water was measured in situ in the vertically stratified Bornholm Basin of the Baltic Sea. Considering all recaptured individuals, the use of hypoxic habitat was comparable to data derived by traditional survey data, but some G. morhua had migrated towards the centre of the c.100 m deep basin and spent about a third of their time at oxygen saturation <50%, possibly to forage on zoobenthos. Maximal residence time per visit in such hypoxic water was limited to a few hours, allowing for the digestion of consumed prey items in waters with sufficient dissolved oxygen.

Analysing migrations of Atlantic cod Gadus morhua in the north-east Atlantic Ocean: then, now and the future

The application of data storage tags bears the potential for a quantum leap in the research on fish migrations, because not only first-capture and recapture positions are known, but at least theoretically, the migration path during the period at large can be reconstructed. Position, however, cannot be measured directly but has to be estimated using the available data on light, temperature, pressure and salinity. The reconstructed locations based on advanced estimation techniques have been termed geolocations. Examples are discussed which illustrate the applicability of geolocations in individual path descriptions, separation of reproductively isolated populations, timing and areas of spawning, tidal transport and use of protected areas. The examples are based on archival tag data from the North Sea, the Baltic Sea, the Barents Sea and Faroese and Icelandic Waters. Besides presenting the state-of-the-art geolocations for cod Gadus morhua in the north-east Atlantic Ocean, the major aim of this review is to raise awareness of gaps in knowledge and to identify ideas for new research.

Hypoxic areas, density-dependence and food limitation drive the body condition of a heavily exploited marine fish predator

Investigating the factors regulating fish condition is crucial in ecology and the management of exploited fish populations. The body condition of cod (Gadus morhua) in the Baltic Sea has dramatically decreased during the past two decades, with large implications for the fishery relying on this resource. Here, we statistically investigated the potential drivers of the Baltic cod condition during the past 40 years using newly compiled fishery-independent biological data and hydrological observations. We evidenced a combination of different factors operating before and after the ecological regime shift that occurred in the Baltic Sea in the early 1990s. The changes in cod condition related to feeding opportunities, driven either by density-dependence or food limitation, along the whole period investigated and to the fivefold increase in the extent of hypoxic areas in the most recent 20 years. Hypoxic areas can act on cod condition through different mechanisms related directly to species physiology, or indirectly to behaviour and trophic interactions. Our analyses found statistical evidence for an effect of the hypoxia-induced habitat compression on cod condition possibly operating via crowding and density-dependent processes. These results furnish novel insights into the population dynamics of Baltic Sea cod that can aid the management of this currently threatened population.

Comparative analysis of marine ecosystems: workshop on predator–prey interactions

Climate and human influences on marine ecosystems are largely manifested by changes in predator–prey interactions. It follows that ecosystem-based management of the worlds oceans requires a better understanding of food web relationships. An international workshop on predator–prey interactions in marine ecosystems was held at the Oregon State University, Corvallis, OR, USA on 16–18 March 2010. The meeting brought together scientists from diverse fields of expertise including theoretical ecology, animal behaviour, fish and seabird ecology, statistics, fisheries science and ecosystem modelling. The goals of the workshop were to critically examine the methods of scaling-up predator–prey interactions from local observations to systems, the role of shifting ecological processes with scale changes, and the complexity and organizational structure in trophic interactions.

Effects of Hypoxic Exposure during Feeding on SDA and Postprandial Cardiovascular Physiology in the Atlantic Cod, Gadus morhua

Some Atlantic cod in the Bornholm Basin undertake vertical foraging migrations into severely hypoxic bottom water. Hypoxic conditions can reduce the postprandial increase in gastrointestinal blood flow (GBF). This could subsequently postpone or reduce the postprandial increase in oxygen consumption (MO2), i.e. the SDA, leading to a disturbed digestion. Additionally, a restricted oxygen uptake could result in an oxygen debt that needs to be compensated for upon return to normoxic waters and this may also affect the ability to process the food. Long-term cardio-respiratory measurements were made on fed G. morhua in order to understand how the cardio-respiratory system of feeding fish respond to a period of hypoxia and a subsequent return to normoxia. These were exposed to 35% water oxygen saturation for 90 minutes, equivalent to the time and oxygen level cod voluntarily endure when searching for food in the Bornholm Basin. We found that i) gastric and intestinal blood flows, cardiac output and MO2 increased after feeding, ii) gastric and intestinal blood flows were spared in hypoxia, and iii) there were no indications of an oxygen debt at the end of the hypoxic period. The magnitude and time course of the measured variables are similar to values obtained from fish not exposed to the hypoxic period. In conclusion, when cod in the field search for and ingest prey under moderate hypoxic conditions they appear to stay within safe limits of oxygen availability as we saw no indications of an oxygen debt, or negative influence on digestive capacity, when simulating field observations.

A metacommunity perspective on source–sink dynamics and management: the Baltic Sea as a case study

The degree to which metapopulation processes influence fish stock dynamics is a largely unresolved issue in marine science and management, especially for highly mobile species such as Atlantic cod (Gadus morhua) and herring (Clupea harengus). The Baltic Sea comprises a heterogeneous oceanographic environment that structures the spatial and temporal distribution of the dominant species cod, herring, and sprat (Sprattus sprattus). Despite local differences, the stocks are traditionally managed as homogeneous units. Here, we present a metacommunity-perspective on source–sink dynamics of Baltic Sea fish stocks by using a spatially disaggregated statistical food web model. The model is fitted to area-specific time series of multiple abiotic and biotic variables using state-space methods. Our analysis reveals pronounced net fluxes between areas, indicative of source–sink dynamics, as well as area-specific differences in species interactions (i.e., density dependence, competition, and predator–prey) and the degree of fishing and climate impact on survival and recruitment. Furthermore, model simulations show that decreasing exploitation pressure in the source area for cod (without reallocating fishing effort) produces an increase in neighboring sink habitats, but a decline of prey species in response to increased predation. Our approach provides valuable insight concerning metacommunity-structuring of marine fish and may serve as an important tool for implementing sustainable management strategies under the ecosystem approach to marine and fisheries management.

Recovery of gastric evacuation rate in Atlantic cod Gadus morhua L surgically implanted with a dummy telemetry device

The current study investigated how the gastric evacuation rate (GER) was affected after surgically introducing dummies of a blood flow biotelemetry system into the abdominal cavity of Atlantic cod, Gadus morhua. Gastric evacuation experiments were performed two and 10 days postsurgery on surgically implanted and control G. morhua force-fed sandeel, Ammodytes tobianus. The results were compared with previously obtained estimates from unstressed conspecifics voluntarily feeding on a similar diet. After two days, GER was significantly lower in the group of fish with the dummy implants compared with the control group, but following 10 days of recovery no significant difference was seen between the two groups. The difference between implanted and control fish observed two days postsurgery may have resulted either from surgery, postsurgical stress and/or the presence of the implant. The conclusion is that 10 days of postsurgical recovery will stabilize GER in G. morhua, thus indicating that at this point the implant per se did not affect GER. Both the fish with surgical implants and controls in this study evacuated their stomachs much slower and with much higher interindividual variation compared with G. morhua feeding voluntarily on similar prey items. The lower GER and higher interindividual variation for force-fed fish indicate that handling, anaesthetization and force-feeding impair GER and that individual fish respond differently to the suppressing effects.

Marine ecosystem connectivity mediated by migrant–resident interactions and the concomitant cross‐system flux of lipids

Abstract Accumulating research argues that migrants influence the functioning and productivity of local habitats and ecosystems along migration routes and potentially drive cross‐system energy fluxes of considerable magnitude, yet empirical documentation of local ecological effects and descriptions of the underlying mechanisms are surprisingly rare. In this study, we discovered migrant–resident interactions and substantial cross‐system lipid transportation in the transition zone between the Baltic Sea and the North Sea where a resident cod population (predators) was found to interact with a herring population (prey) on a seasonal basis. We traced the lipids, using fatty acid trophic markers (FATM), from the herring feeding grounds in the North Sea to the cod livers in the Western Baltic Sea. Time series analysis of population dynamics indicated that population‐level production of cod is positively affected by the herring subsidies. However, the underlying mechanisms were more complicated than anticipated. During the herring season, large cod received most of its dietary lipids from the herring, whereas smaller cod were prevented from accessing the lipid pool due to a mismatch in predator–prey size ratio. Furthermore, while the herring were extremely rich in bulk energy, they were surprisingly poor in a specific functional fatty acid. Hence, our study was the first to illustrate how the magnitude cross‐system fluxes of subsidies in migrant–resident systems are potentially constrained by the size structure of the resident predator population and the nutritional quality of the migrants.

Sustainable use of Baltic Sea resources

The BONUS symposium “Science delivery for sustainable use of the Baltic Sea living resources” held in Tallinn, Estonia, in October 2017 was an opportunity for the presentation and discussion of 107 papers that examined the state and dynamics of living resources of the Baltic Sea, and associated management challenges. The symposium included a half-day stakeholder panel discussion that addressed the main challenges related to sustainable management and matching research and policy/management needs. Based on the five symposium papers published in this Special Issue as well as the stakeholder panel discussion, it can be concluded that (i) new observations about the feeding ecology of clupeids supports a more complete understanding of trophic interactions in the pelagic realm and improved calibration of ecosystem models, (ii) to safequard sustainable and diverse fisheries resources, one should take into account the specific local characteristics of the fish community, (iii) to safeguard sustainable use of marine resources and mitigate cross-sectoral and transboundary conflicts, a risk-based approach should be adopted, and (iv) incorporation of scientific advice into management faces several obstacles including the reality that not all readily available knowledge is currently being incorporated into the decision-making process.

Supplementary material from "Hypoxic areas, density-dependence and food limitation drive the body condition of a heavily exploited marine fish predator"

Investigating the factors regulating fish condition is crucial in ecology and the management of exploited fish populations. The body condition of cod (Gadus morhua) in the Baltic Sea has dramatically decreased during the past two decades, with large implications for the fishery relying on this resource. Here, we statistically investigated the potential drivers of the Baltic cod condition during the past 40 years using newly compiled fishery-independent biological data and hydrological observations. We evidenced a combination of different factors operating before and after the ecological regime shift that occurred in the Baltic Sea in the early 1990s. The changes in cod condition related to feeding opportunities, driven either by density-dependence or food limitation, along the whole period investigated and to the fivefold increase in the extent of hypoxic areas in the most recent 20 years. Hypoxic areas can act on cod condition through different mechanisms related directly to species physiology, or indirectly to behaviour and trophic interactions. Our analyses found statistical evidence for an effect of the hypoxia-induced habitat compression on cod condition possibly operating via crowding and density-dependent processes. These results furnish novel insights into the population dynamics of Baltic Sea cod that can aid the management of this currently threatened population.