The international Wadden Sea is an important flatfish nursery. Information from the Dutch Wadden Sea indicates that the flatfish nursery function of the area has been affected during the last decades. Increased seawater temperature has affected settling, habitat suitability for and growth performance of the various flatfish species. Settling of plaice, flounder and to a lesser extent sole larvae occurs earlier nowadays. In the 1960s, 0-, I-, II- and III-group plaice were present, but since 2000, II-group has disappeared and densities of I-group have decreased. For juvenile flounder, II-group almost disappeared, and for dab, a decline in densities of all age groups was observed from the 1990s onwards. Summer temperatures exceed the optimum for the cold-water species (plaice, flounder and dab) with increasing frequency, level and duration. Only for 0-group sole, the period with optimal growth conditions has become longer and has resulted in increased growth. Mortality rates in 0-group plaice have increased, coinciding with an increase in water temperatures and an increase in the abundance of predators. The decrease in density of juvenile plaice and dab in the Wadden Sea has not affected recruitment to North Sea stocks, suggesting that other areas have taken over part of the nursery function. The predicted increase in seawater temperature in the next decades will continue to improve the conditions for sole. The temperature tolerance of plaice and dab and to a lesser extent flounder will further reduce their scope for growth and may ultimately result in their disappearance from the Wadden Sea.
Coastal areas in north-western Europe have been influenced by elevated nutrient levels starting in the 1960s. Due to efficient measures, both nitrate and phosphate levels decreased since the mid-1980s. The co-occurring declines in nutrient loadings and fish productivity are often presumed to be causally linked. We investigated whether four resident fish species (twaite shad, bull-rout, thick-lipped grey mullet and eelpout), that spend the majority of their life in the vicinity of the coast, differed in growth between the historic eutrophication period compared to the recent lower nutrient-level period. Based on Von Bertalanffy growth models of length at age, and the analysis of annual otolith increments, we investigated the difference in sex-specific growth patterns and related these to temperature, eutrophication level (Chlorophyll a), growth window and fish density. In all four species, annual otolith growth rates during the early life stages differed between the two periods, mostly resulting in larger lengths at age in the recent period. All species showed significant correlations between increment size and temperature, explaining the observed period differences. The lack of an effect of total fish biomass provided no evidence for density dependent growth. A correlation with chlorophyll was found in bull-rout, but the relationship was negative, thus not supporting the idea of growth enhanced by high nutrient levels. In conclusion, we found no evidence for reduced growth related to de-eutrophication. Our results indicate that temperature rise due to climate change had a greater impact on growth than reduced food availability due to de-eutrophication. We discuss potential consequences of growth changes for length-based indicators used in management.
The goals of the 2019 dab exchange were (1) an estimation of ageing uncertainty for the dab.27.3a4 stock, (2) compare different preparation methods, and (3) examine seasonality in deposition of opaque/translucent material on the edge of the otolith across a latitudinal gradient. The second and third goals were follow-up actions proposed by the Workshop on Age Reading of Dab in 2015 (WKARDAB2). Sixteen age readers from seven countries (Belgium, France, Iceland, Netherlands, Germany, Denmark, England) participated in the exchange. Two sets of otoliths were examined. The first set (SmartDots event 244) consisted of 64 otoliths from ICES Subarea 4 (North Sea) and Division 3a (Skagerrak and Kattegat). These were prepared in three different manners (whole, sectioned, and sectioned-and-stained), resulting in 192 images. The second set (SmartDots event 245) consisted of 68 otoliths from ICES Subarea 4 (North Sea) and Division 5a (Icelandic waters). These were prepared in two different manners (whole and sectioned-and-stained), resulting in 136 images. Age readers were also asked to determine the edge type for each otolith in the second set. First, the data were analysed for each event and method separately, including all age readers. Then, the combined set of otoliths from Subarea 4 and Division 3a were analysed for each method separately, including only the advanced age readers who supply age data for the assessment of the dab.27.3a4 stock. Additional analyses were carried out to compare the different preparation methods, and to examine seasonality by area in the deposition of opaque/translucent material. Age reading uncertainty was high in dab compared to most other North Sea flatfish species. The percentage agreement was 69-73% and the coefficient of variation was 13-14% for dab.27.3a4, based on the age determinations of readers who supply age data for the assessment. As observed previously, uncertainty was highest in the third quarter. Comparison of methods showed better calibration results for sectioned-and-stained otoliths than for whole otoliths or sectioned otoliths. However, the ages of young fish (1-6) were on average estimated to be higher based on sectioned-and-stained otoliths than based on whole otoliths. Age reading certainty may improve if sectioned-and-stained otoliths are used by all readers, preferably after validation that this is the best method to estimate true age. The edge analysis was hampered by low number of otoliths per month and area, and inconsistency between readers. Nevertheless, the analysis did show high variability in the timing of deposition of opaque/translucent material. Consequently, a fixed rule on how to interpret the edge of the otolith in the third quarter is not applicable for dab.
Hydrodynamic Ocean Circulation Models and Lagrangian particle tracking models are valuable tools e.g. in coastal ecology to identify the connectivity between offshore spawning and coastal nursery areas of commercially important fish, for risk assessment and more for defining or evaluating marine protected areas. Most studies are based on only one model and do not provide levels of uncertainty. Here this uncertainty was addressed by applying a suite of 11 North Sea models to test what variability can be expected concerning connectivity. Different notional test cases were calculated related to three important and well-studied North Sea fish species: herring (Clupea harengus), and the flatfishes sole (Solea solea) and plaice (Pleuronectes platessa). For sole and plaice we determined which fraction of particles released in the respective spawning areas would reach a coastal marine protected area. For herring we determined the fraction located in a wind park after a predefined time span. As temperature is more and more a focus especially in biological and global change studies, furthermore inter-model variability in temperatures experienced by the virtual particles was determined. The main focus was on the transport variability originating from the physical models and thus biological behavior was not included. Depending on the scenario, median experienced temperatures differed by 3°C between years. The range between the different models in one year was comparable to this temperature range observed between modelled years. Connectivity between flatfish spawning areas and the coastal protected area was highly dependent on the release location and spawning time. No particles released in the English Channel in the sole scenario reached the protected area while up to 20% of the particles released in the plaice scenario did. Interannual trends in transport directions and connectivity rates were comparable between models but absolute values displayed high variations. Most models showed systematic biases during all years in comparison to the ensemble median, indicating that in general interannual variation was represented but absolute values varied. In conclusion: variability between models is generally high and management decisions or scientific analysis using absolute values from only one single model might be biased and results or conclusions drawn from such studies need to be treated with caution. We further concluded that more true validation data for particle modelling are required.
The shallow coastal zone is an essential nursery habitat for juvenile flatfish species such as sole (Solea solea L.). The increased frequency of shoreface nourishments along the coast is likely to affect this nursery function by altering important habitat conditions, including sediment grain size. Sediment preference of juvenile sole (41–91mm) was studied in a circular preference chamber in order to understand the relationship between grain size and sole distribution. The preference tests were carried out at 11°C and 20°C to reflect seasonal influences. The juveniles showed a significant preference for finer sediments. This preference was not length dependent (within the length range tested) nor affected by either temperatures. Juvenile sole have a small home range and are not expected to move in response to unfavourable conditions. As a result, habitat alterations may have consequences for juvenile survival and subsequently for recruitment to adult populations. It is therefore important to carefully consider nourishment grain size characteristics to safeguard suitable nursery habitats for juvenile sole.
Concern exists about the potential effects of pile-driving sounds on fish, but evidence is limited, especially for fish larvae. A device was developed to expose larvae to accurately reproduced pile-driving sounds. Controlled exposure experiments were carried out to examine the lethal effects in common sole larvae. No significant effects were observed at zero-to-peak pressure levels up to 210 dB re 1 μPa2 and cumulative sound exposure levels up to 206 dB re 1 μPa2·s, which is well above the US interim criteria for nonauditory tissue damage in fish. Experiments are presently being carried out for European sea bass and herring larvae.
Underwater sound from human activities may affect fish behaviour negatively and threaten the stability of fish stocks. However, some fundamental understanding is still lacking for adequate impact assessments and potential mitigation strategies. For example, little is known about the potential contribution of the temporal features of sound, the efficacy of ramp-up procedures, and the generalisability of results from indoor studies to the outdoors. Using a semi-natural set-up, we exposed European seabass in an outdoor pen to four treatments: 1) continuous sound, 2) intermittent sound with a regular repetition interval, 3) irregular repetition intervals and 4) a regular repetition interval with amplitude 'ramp-up'. Upon sound exposure, the fish increased swimming speed and depth, and swam away from the sound source. The behavioural readouts were generally consistent with earlier indoor experiments, but the changes and recovery were more variable and were not significantly influenced by sound intermittency and interval regularity. In addition, the 'ramp-up' procedure elicited immediate diving response, similar to the onset of treatment without a 'ramp-up', but the fish did not swim away from the sound source as expected. Our findings suggest that while sound impact studies outdoors increase ecological and behavioural validity, the inherently higher variability also reduces resolution that may be counteracted by increasing sample size or looking into different individual coping styles. Our results also question the efficacy of 'ramp-up' in deterring marine animals, which warrants more investigation.
The Wadden Sea bordering the Dutch, German and Danish coast, is traditionally a region with important functions for many fish species: as a nursery area for juveniles (marine juveniles), as a feeding area, as a transit to and from fresh water, and resident species complete their whole life cycle there. Because of indications that the importance of the Dutch Wadden Sea has changed drastically for many species during the past decades, we analysed and classified trends of 24 common fish species in the last 45 years, which were assigned to 5 different ecological guilds. Trends were examined for three Wadden Sea regions and compared to trends in the adjacent two North Sea coastal regions. For these analyses we made a combined use of two longterm time series: an annual beamtrawl survey, the Demeral Fish Survey (DFS) with a high spatial but poor seasonal resolution and a Tyke series with a high seasonal but poor spatial resolution. We investigated for which species the DFS survey was appropriate for trend analysis, and we evaluated whether a change in timing may contribute to patterns in DFS time trends. Total fish biomass showed a similar pattern in all tidal basins with an increase from 1970 to 1980, a peak in the mid-1980s and a strong decline from 1980 to 2000, with a subsequent stable trend. The pattern in the coastal region deviated especially in the past 10 years, with a further decline along the Dutch Wadden coast and an increase along the mainland coast. Most dramatic declines throughout the Wadden Sea occurred in species belonging to the marine juvenile guild, notably plaice, sole and dab. A declining trend in marine juveniles is on-going in the western part, while it recently stabilised or even increased in the central and eastern part and in the coastal regions. Resident species showed more variable trends in the Wadden Sea with less pronounced directions: both increases and decreases occurred. In the coastal regions, several resident species have increased considerably in the last 15 years, a pattern not observed in the Wadden Sea. Also the size structure of the fish community changed in all regions, with generally the strongest declines in the largest size classes. The combined use of the two surveys showed that for some species the DFS was not timed in the period of peak occurrence. Although the phenology of several species has changed, the DFS survey period still encompasses the peak period of most species. (C) 2016 Elsevier B.V. All rights reserved.
Connectivity throughout the life cycle of flatfish remains an open question, especially during the early life stages. Their effective management requires understanding of how spawning grounds and nurseries are connected and what processes influence larval retention and dispersal. The case of sole (Solea solea L.) is of particular interest because it is one of the most valuable commercial species in the North Sea, although stocks are chronically overexploited and variability in interannual recruitment is high. The transport of sole larvae from the spawning grounds to the nurseries is driven by hydrodynamic processes, but the final dispersal pattern and larval survival/abundance might be influenced by both behavioral and environmental factors. Therefore it is important to understand the relative impact of hydrodynamics, environment, behavior and ecophysiology on sole larval dispersal. Here we use a particle-tracking transport model coupled to a 3D hydro-dynamic model of the North Sea to investigate interannual variability of the transport of sole larvae over a 12-year period (1995–2006). A sensitivity analysis is performed to assess the relative impact of hydrodynamics, temperature and behavior on the recruitment dynamics to the nurseries. Four scenarios have been tested: (i) constant forcing of sea surface temperature during all years but varying meteorological forcing and river runoff, (ii) constant meteorological forcing during the whole period but varying sea surface temperature and river runoff, (iii) no vertical migration and (iv) an extended drift period (max. 30days) before settlement if the larvae are not close to a suitable sediment type. Results suggest that year-to-year variability of larval supply to the nurseries is high, both in terms of abundance and larval source (balance between retention and dispersal). Sensitivity analysis shows that larval abundance at the end of the larval stage increases considerably if a settling delay is included. The impact of vertical migration on larval transport and the variations in larval retention at the nurseries due to varying meteorological conditions and sea surface temperature forcing are not spatially consistent.
Conclusions & Perspectives Connectivity of flatfish remains an open question, especially at early life stages. The impact of anthropogenic factors, such as climate change on larval dispersal remains unknown. The case of sole (Solea solea) is of particular interest because sole is one of the most valuable commercial species in the North Sea. It is important to understand how the retention/dispersal of larvae would be affected by climate change in order to propose appropriate measures for the management of the North Sea stock. Introduction
In view of the rapid extension of offshore wind farms, there is an urgent need to improve our knowledge on possible adverse effects of underwater sound generated by pile-driving. Mortality and injuries have been observed in fish exposed to loud impulse sounds, but knowledge on the sound levels at which (sub-)lethal effects occur is limited for juvenile and adult fish, and virtually non-existent for fish eggs and larvae. A device was developed in which fish larvae can be exposed to underwater sound. It consists of a rigid-walled cylindrical chamber driven by an electro-dynamical sound projector. Samples of up to 100 larvae can be exposed simultaneously to a homogeneously distributed sound pressure and particle velocity field. Recorded pile-driving sounds could be reproduced accurately in the frequency range between 50 and 1000 Hz, at zero to peak pressure levels up to 210 dB re 1µPa(2) (zero to peak pressures up to 32 kPa) and single pulse sound exposure levels up to 186 dB re 1µPa(2)s. The device was used to examine lethal effects of sound exposure in common sole (Solea solea) larvae. Different developmental stages were exposed to various levels and durations of pile-driving sound. The highest cumulative sound exposure level applied was 206 dB re 1µPa(2)s, which corresponds to 100 strikes at a distance of 100 m from a typical North Sea pile-driving site. The results showed no statistically significant differences in mortality between exposure and control groups at sound exposure levels which were well above the US interim criteria for non-auditory tissue damage in fish. Although our findings cannot be extrapolated to fish larvae in general, as interspecific differences in vulnerability to sound exposure may occur, they do indicate that previous assumptions and criteria may need to be revised.
We investigated long-term trends in abundance of the NE Atlantic population of brown shrimp Crangon crangon based on data collected in annual autumn surveys carried out along the coasts of the North Sea in The Netherlands, Germany and Denmark. Surveys covered some estuaries and intertidal areas, as well as shrimp fishery grounds. The 40 yr period showed distinct regional trend differences, but only in estuarine areas: the Western Dutch Wadden Sea (increase), the Eastern Dutch Wadden Sea (decrease) and the Oosterschelde (decrease). The decrease in the Oosterschelde coincided with the closure of this sea arm from fresh water inflow. Increases in the Western and Eastern Dutch Wadden Sea were driven primarily by increased abundance of small shrimp, suggesting that the causative mechanism likely affected younger shrimp. Trends in potential causes were examined in the 2 regions. Of all abiotic factors investigated, only water clarity decreased significantly in the west but not in the east. Mean abundance of potential predators (cod, whiting, shore crab and swimming crab) was used as an indicator of predation pressure. Shore crab showed similar increases in both areas. In the other 3 species, predator abundance in the east was generally initially higher than in the west, but declined towards similar densities in recent years. Shrimp fishing pressure in NW Europe has increased strongly since the 1990s. Preliminary, spatially resolved logbook data show that shrimp landings from the Western Wadden Sea followed this increase, but those from the Eastern Wadden Sea remained stable. Apart from mechanisms related to fishing pressure and water clarity, a candidate cause of differences in abundance may be differences in food condition or competition for food. Generally biomass of benthos, a primary food for shrimp, increased in the west and remained stable in the east. However, because brown shrimp are omnivorous, quantification of changing fractions of food available to them is difficult.
Driven by the concern that impulsive noise produced by offshore pile driving may lead to mortality of fish larvae, a device was developed for testing the sensitivity of small fish and fish larvae to sound exposure. The device consists of a rigid-walled cylindrical chamber (110-mm diameter, 160-mm height), driven by an electrodynamical sound projector. Samples of up to 100 larvae can be exposed simultaneously to a homogeneously distributed sound pressure and particle velocity field, at a controllable static pressure up to 3 bars. Two configurations are available with either a dominant sound pressure or a dominant particle velocity exposure. Recorded piling noise can be reproduced in a controlled way, in the frequency range between 50 Hz and 1 kHz, at zero to peak pressure up to 40 kPa and single pulse sound exposure levels up to 187 dB re 1 μPa2 s, or peak particle velocity up to 2.2 cm/s and integrated square particle velocity level 124 dB re 1 (nm/s)2 s. Tests are carried out in which sole (Solea solea) larvae at different stages of development were exposed to various levels and durations of piling noise.