Phenological trends and trophic mismatch across multiple levels of a North Sea pelagic food web

Differential phenological responses to climate among species are predicted to disrupt trophic interactions, but datasets to evaluate this are scarce. We compared phenological trends for species from 4 levels of a North Sea food web over 24 yr when sea surface temperature (SST) increased significantly. We found little consistency in phenological trends between adjacent trophic levels, no significant relationships with SST, and no significant pairwise correlations between predator and prey phenologies, suggesting that trophic mismatching is occurring. Finer resolution data on timing of peak energy demand (mid-chick-rearing) for 5 seabird species at a major North Sea colony were compared to modelled daily changes in length of 0-group (young of the year) lesser sandeels Ammodytes marinus. The date at which sandeels reached a given threshold length became significantly later during the study. Although the phenology of all the species except shags also became later, these changes were insufficient to keep pace with sandeel length, and thus mean length (and energy value) of 0-group sandeels at mid-chick-rearing showed net declines. The magnitude of declines in energy value varied among the seabirds, being more marked in species showing no phenological response (shag, 4.80 kJ) and in later breeding species feeding on larger sandeels (kittiwake, 2.46 kJ) where, due to the relationship between sandeel length and energy value being non-linear, small reductions in length result in relatively large reductions in energy. However, despite the decline in energy value of 0-group sandeels during chick-rearing, there was no evidence of any adverse effect on breeding success for any of the seabird species. Trophic mismatch appears to be prevalent within the North Sea pelagic food web, suggesting that ecosystem functioning may be disrupted.

Wasp-Waist Interactions in the North Sea Ecosystem

In a “wasp-waist” ecosystem, an intermediate trophic level is expected to control the abundance of predators through a bottom-up interaction and the abundance of prey through a top-down interaction. Previous studies suggest that the North Sea is mainly governed by bottom-up interactions driven by climate perturbations. However, few studies have investigated the importance of the intermediate trophic level occupied by small pelagic fishes.

Scale-dependent response diversity of seabirds to prey in the North Sea

Functional response diversity is defined as the diversity of responses to environmental change among species that contribute to the same ecosystem function. Because different ecological processes dominate on different spatial and temporal scales, response diversity is likely to be scale dependent. Using three extensive data sets on seabirds, pelagic fish, and zooplankton, we investigate the strength and diversity in the response of seabirds to prey in the North Sea over three scales of ecological organization. Two-stage analyses were used to partition the variance in the abundance of predators and prey among the different scales of investigation: variation from year to year, variation among habitats, and variation on the local patch scale. On the year-to-year scale, we found a strong and synchronous response of seabirds to the abundance of prey, resulting in low response diversity. Conversely, as different seabird species were found in habitats dominated by different prey species, we found a high diversity in the response of seabirds to prey on the habitat scale. Finally, on the local patch scale, seabirds were organized in multispecies patches. These patches were weakly associated with patches of prey, resulting in a weak response strength and a low response diversity. We suggest that ecological similarities among seabird species resulted in low response diversity on the year-to-year scale. On the habitat scale, we suggest that high response diversity was due to interspecific competition and niche segregation among seabird species. On the local patch scale, we suggest that facilitation with respect to the detection and accessibility of prey patches resulted in overlapping distribution of seabirds but weak associations with prey. The observed scale dependencies in response strength and diversity have implications for how the seabird community will respond to different environmental disturbances.

Temporal changes in plankton of the North Sea: community shifts and environmental drivers

This paper analyses long-term and seasonal changes in the North Sea plankton community during the period 1970 to 2008. Based on Continuous Plankton Recorder (CPR) data covering 38 yr, major changes in both phytoplankton and zooplankton abundance and community structure were identified. Regime changes were detected around 1978, 1989 and 1998. The first 2 changes have been discussed in the literature and are defined as a cold episodic event (1978) and a regime shift towards a warm dynamic regime (1989). The effect of these 2 regime changes on plankton indicators was assessed and checked against previous studies. The 1998 change represents a shift in the abundance and seasonal patterns of dinoflagellates and the dominant zooplankton group, the neritic copepods. Furthermore, environmental factors such as air temperature, wind speed and the North Atlantic water inflow were identified as potential drivers of change in seasonal patterns, and the most-likely environmental causes for detected changes were assessed. We suggest that a change in the balance of dissolved nutrients driven by these environmental factors was the cause of the latest change in plankton community structure, which in turn could have affected the North Sea fish community.

Comparative seasonal dynamics of Centropages typicus at seven coastal monitoring stations in the North Sea, English Channel and Bay of Biscay

We review current knowledge and understanding of the biology and ecology of Centropages typicus in the European shelf-seas (e.g. North Sea, English Channel and Bay of Biscay). Our study is based on observations at seven coastal time-series stations as well as on the Continuous Plankton Recorder dataset. This paper focuses on the influence of the environmental parameters (e.g. temperature and Chla) on the life cycle and distribution of C typicus and provides a comparison with its congeneric species C. hamatus and C. chierchiae in the study area. Data on abundance, seasonality and egg production have been used to define the temperature and chlorophyll optima for occurrence and reproduction of Centropages spp. within this region of the European shelf-seas. (C) 2007 Elsevier Ltd. All rights reserved.

Predator–prey reversal: A possible mechanism for ecosystem hysteresis in the North Sea?

Removal of large predatory fishes from marine ecosystems has resulted in persistent ecosystem shifts, with collapsed predator populations and super-abundant prey populations. One explanation for these shifts is reversals of predator–prey roles that generate internal feedbacks in the ecosystems. Pelagic forage fish are often predators and competitors to the young life stages of their larger fish predators. I show that cod recruitment in the North Sea has been negatively related to the spawning-stock biomass of herring for the last 44 years. Herring, together with the abundance of Calanus finmarchicus, the major food for cod larvae, were the main predictors of cod recruitment. These predictors were of equivalent importance, worked additively, and explained different parts of the dynamics in cod recruitment. I suggest that intensive harvesting of cod has released herring from predator control, and that a large population of herring suppresses cod recruitment through predation on eggs and larvae. This feedback mechanism can promote alternative stable states and therefore cause hysteresis to occur under changing conditions; however, harvesting of herring might at present prevent a shift in the ecosystem to a herring-dominated state.

Mechanisms of long-term decline in size of lesser sandeels in the North Sea explored using a growth and phenology model

The lesser sandeel Ammodytes marinus is a key species in the North Sea ecosystem, transferring energy from planktonic producers to top predators. Previous studies have shown a long-term decline in the size of 0-group sandeels in the western North Sea, but they were unable to pinpoint the mechanism (later hatching, slower growth or changes in size-dependent mortality) or cause. To investigate the first 2 possibilities we combined 2 independent time series of sandeel size, namely data from chick-feeding Atlantic puffins Fratercula arctica and from the Continuous Plankton Recorder (CPR), in a novel statistical model implemented using Markov Chain Monte Carlo (MCMC). The model estimated annual mean length on 1 July, as well as hatching date and growth rate for sandeels from 1973 to 2006. Mean length-at-date declined by 22% over this period, corresponding to a 60% decrease in energy content, with a sharper decline since 2002. Up to the mid-1990s, the decline was associated with a trend towards later hatching. Subsequently, hatching became earlier again, and the continued trend towards smaller size appears to have been driven by lower growth rates, particularly in the most recent years, although we could not rule out changes in size-dependent mortality. Our findings point to major changes in key aspects of sandeel life history, which we consider are most likely due to direct and indirect temperature-related changes over a range of biotic factors, including the seasonal distribution of copepods and intra- and inter-specific competition with planktivorous fish. The results have implications both for the many predators of sandeels and for age and size of maturation in this aggregation of North Sea sandeels.

Synchronous response of marine plankton ecosystems to climate in the Northeast Atlantic and the North Sea

Over the last few decades, global warming has accelerated both the rate and magnitude of changes observed in many functional units of the Earth System. In this context, plankton are sentinel organisms because they are sensitive to subtle levels of changes in temperature and might help in identifying the current effects of climate change on pelagic ecosystems. In this paper, we performed a comparative approach in two regions of the North Atlantic (i.e. the Northeast Atlantic and the North Sea) to explore the relationships between changes in marine plankton, the regional physico-chemical environment and large-scale hydro-climatic forcing using four key indices: the North Atlantic Oscillation (NAO), the Atlantic Multidecadal Oscillation (AMO), the East Atlantic (EA) pattern and Northern Hemisphere Temperature (NHT) anomalies. Our analyses suggest that long-term changes in the states of the two ecosystems were synchronous and correlated to the same large-scale hydro-climatic variables: NHT anomalies, the AMO and to a lesser extent the EA pattern. No significant correlation was found between long-term ecosystem modifications and the state of the NAO. Our results suggest that the effect of climate on these ecosystems has mainly occurred in both regions through the modulation of the thermal regime.

Long-term changes in abundance and distribution of microzooplankton in the NE Atlantic and North Sea

Long-term changes in mesozooplankton and phytoplankton populations have been well documented in the North Atlantic region, whereas data for microzooplankton are scarce. This neglected component of the plankton is a vital link in marine food-webs, grazing on smaller flagellates and cyanobacteria and in turn providing food for the larger mesozooplankton. We use the latest tintinnid (Ciliophora, Protista) data from the Continuous Plankton Recorder (CPR) survey in the NE Atlantic and North Sea to examine the phenology, distribution and abundance of this important group of ciliates. Presence/absence data came from 167 122 CPR samples collected between 1960 and 2009 and abundance data from 49 662 samples collected between 1996 and 2009. In the North Atlantic the genus Dictyocysta spp. dominated and Parafavella gigantea showed an increase in abundance around Iceland and Greenland. In the North Sea higher densities of Tintinnopsis spp., Favella serrata and Ptychocylis spp. were found. The presence of tintinnids in CPR samples collected in the North Atlantic has increased over the last 50 years and the seasonal window of high abundance has lengthened. Conversely in the North Sea there has been an overall reduction in abundance. We discuss possible drivers for these long-term changes and point the way forward to more holistic studies that examine how ecosystems, rather than just selected taxa, are responding to climate change.

Temperature affects the timing of spawning and migration of North Sea mackerel

Climate change accentuates the need for knowing how temperature impacts the life history and productivity of economically and ecologically important species of fish. We examine the influence of temperature on the timing of the spawning and migrations of North Sea Mackerel using data from larvae CPR surveys, egg surveys and commercial landings from Danish coastal fisheries in the North Sea, Skagerrak, Kattegat and inner Danish waters. The three independent sources of data all show that there is a significant relationship between the timing of spawning and sea surface temperature. Large mackerel are shown to arrive at the feeding areas before and leave later than small mackerel and the sequential appearance of mackerel in each of the feeding areas studied supports the anecdotal evidence for an eastward post-spawning migration. Occasional commercial catches taken in winter in the Sound N, Kattegat and Skagerrak together with catches in the first quarter IBTS survey furthermore indicate some overwintering here. Significant relationships between temperature and North Sea mackerel spawning and migration have not been documented before. The results have implications for mackerel resource management and monitoring. An increase in temperature is likely to affect the timing and magnitude of the growth, recruitment and migration of North Sea mackerel with subsequent impacts on its sustainable exploitation.

Long-Term Retrospective Analysis of Mackerel Spawning in the North Sea: A New Time Series and Modeling Approach to CPR Data

We present a unique view of mackerel (Scomber scombrus) in the North Sea based on a new time series of larvae caught by the Continuous Plankton Recorder (CPR) survey from 1948-2005, covering the period both before and after the collapse of the North Sea stock. Hydrographic backtrack modelling suggested that the effect of advection is very limited between spawning and larvae capture in the CPR survey. Using a statistical technique not previously applied to CPR data, we then generated a larval index that accounts for both catchability as well as spatial and temporal autocorrelation. The resulting time series documents the significant decrease of spawning from before 1970 to recent depleted levels. Spatial distributions of the larvae, and thus the spawning area, showed a shift from early to recent decades, suggesting that the central North Sea is no longer as important as the areas further west and south. These results provide a consistent and unique perspective on the dynamics of mackerel in this region and can potentially resolve many of the unresolved questions about this stock

Climate-induced effects on the meroplankton and the benthic-pelagic ecology of the North Sea

Analyses of long-term time series of North Sea plankton and sea surface temperature (SST) data reveal that the annual planktonic larval abundance of three benthic phyla, Echinodermata, Arthropoda, and Mollusca, responds positively and immediately to SST. Long-term outcomes for the planktonic abundance of these three phyla are different, however. The planktonic larvae of echinoderms and decapod crustaceans have increased in abundance from 1958 to 2005, and especially since the mid-1980s, as North Sea SST has increased. In contrast, the abundance of bivalve mollusc larvae has declined, despite the positive year-to-year relationship between temperature and bivalve larval abundance continuing to hold. We argue that the changes in meroplankton abundance, coincident with increased phytoplankton and declining holoplankton, reflect the synchronous effect of rising SST and related changes in the pelagic community on the reproduction and recruitment of many benthic marine invertebrates. Under this scenario, the long-term decline in bivalve mollusc larvae will reflect increased predation on the settled larvae and adults by benthic decapods. These alterations in the zooplankton may therefore describe an ecosystem-wide restructuring of North Sea trophic interactions.