Ecological effects of the North Atlantic oscillation

Climatic oscillations as reflected in atmospheric modes such as the North Atlantic Oscillation (NAO) may be seen as a proxy for regulating forces in aquatic and terrestrial ecosystems. Our review highlights the variety of climate processes related to the NAO and the diversity in the type of ecological responses that different biological groups can display. Available evidence suggests that the NAO influences ecological dynamics in both marine and terrestrial systems, and its effects may be seen in variation at the individual, population and community levels. The ecological responses to the NAO encompass changes in timing of reproduction, population dynamics, abundance, spatial distribution and interspecific relationships such as competition and predator-prey relationships. This indicates that local responses to large-scale changes may be more subtle than previously suggested. We propose that the NAO effects may be classified as three types: direct, indirect and integrated. Such a classification will help the design and interpretation of analyses attempting to relate ecological changes to the NAO and, possibly, to climate in general.

Strong preference for decapod prey by the western rock lobster Panulirus cygnus

Western rock lobsters, Panulirus cygnus are an abundant benthic consumer distributed along the temperate west coast of Australia and constitute the largest single species fishery in Australia. As a dominant consumer, it is important to understand their predator-prey interactions as they can potentially exert strong trophic effects, and may influence ecosystem function as seen in other spiny lobster species. While previous field studies have focused on the diet composition of P. cygnus, this study investigated their preference for various benthic invertebrate prey to better understand the likely predator-prey interactions of P. cygnus. Prey preferences of small sub-legal juvenile lobsters, as well as medium and large legal-sized mature lobsters were investigated using laboratory feeding trials to identify size-associated differences in lobster prey preference. Handling time and diet quality were investigated to estimate energetic cost and gain from consuming different prey which may explain prey choice by lobsters. It was found that large lobsters preferred crabs and mussels while medium and small lobsters preferred crabs over mussels, gastropods, and sea urchins. This suggests that strong predator-prey interactions between P. cygnus and crabs may occur in the wild.

DISCOVERY 2010: Spatial and temporal variability in a dynamic polar ecosystem

The Scotia Sea has been a focus of biological- and physical oceanographic study since the Discovery expeditions in the early 1900s. It is a physically energetic region with some of the highest levels of productivity in the Southern Ocean. It is also a region within which there have been greater than average levels of change in upper water column temperature. We describe the results of three cruises transecting the central Scotia Sea from south to north in consecutive years and covering spring, summer and autumn periods. We also report on some community level syntheses using both current-day and historical data from this region. A wide range of parameters were measured during the field campaigns, covering the physical oceanography of the region, air–sea CO2 fluxes, macro- and micronutrient concentrations, the composition and biomass of the nano-, micro- and mesoplankton communities, and the distribution and biomass of Antarctic krill and mesopelagic fish. Process studies examined the effect of iron-stress on the physiology of primary producers, reproduction and egestion in Antarctic krill and the transfer of stable isotopes between trophic layers, from primary consumers up to birds and seals. Community level syntheses included an examination of the biomass-spectra, food-web modelling, spatial analysis of multiple trophic layers and historical species distributions. The spatial analyses in particular identified two distinct community types: a northern warmer water community and a southern cold community, their boundary being broadly consistent with the position of the Southern Antarctic Circumpolar Current Front (SACCF). Temperature and ice cover appeared to be the dominant, over-riding factors in driving this pattern. Extensive phytoplankton blooms were a major feature of the surveys, and were persistent in areas such as South Georgia. In situ and bioassay measurements emphasised the important role of iron inputs as facilitators of these blooms. Based on seasonal DIC deficits, the South Georgia bloom was found to contain the strongest seasonal carbon uptake in the ice-free zone of the Southern Ocean. The surveys also encountered low-production, iron-limited regions, a situation more typical of the wider Southern Ocean. The response of primary and secondary consumers to spatial and temporal heterogeneity in production was complex. Many of the life-cycles of small pelagic organisms showed a close coupling to the seasonal cycle of food availability. For instance, Antarctic krill showed a dependence on early, non-ice-associated blooms to facilitate early reproduction. Strategies to buffer against environmental variability were also examined, such as the prevalence of multiyear life-cycles and variability in energy storage levels. Such traits were seen to influence the way in which Scotia Sea communities were structured, with biomass levels in the larger size classes being higher than in other ocean regions. Seasonal development also altered trophic function, with the trophic level of higher predators increasing through the course of the year as additional predator-prey interactions emerged in the lower trophic levels. Finally, our studies re-emphasised the role that the simple phytoplankton-krill-higher predator food chain plays in this Southern Ocean region, particularly south of the SACCF. To the north, alternative food chains, such as those involving copepods, macrozooplankton and mesopelagic fish, were increasingly important. Continued ocean warming in this region is likely to increase the prevalence of such alternative such food chains with Antarctic krill predicted to move southwards.

Encounter success of free-ranging marine predator movements across a dynamic prey landscape

Movements of wide-ranging top predators can now be studied effectively using satellite and archival telemetry. However, the motivations underlying movements remain difficult to determine because trajectories are seldom related to key biological gradients, such as changing prey distributions. Here, we use a dynamic prey landscape of zooplankton biomass in the north-east Atlantic Ocean to examine active habitat selection in the plankton-feeding basking shark Cetorhinus maximus. The relative success of shark searches across this landscape was examined by comparing prey biomass encountered by sharks with encounters by random-walk simulations of ‘model’ sharks. Movements of transmitter-tagged sharks monitored for 964 days (16754km estimated minimum distance) were concentrated on the European continental shelf in areas characterized by high seasonal productivity and complex prey distributions. We show movements by adult and sub-adult sharks yielded consistently higher prey encounter rates than 90% of random-walk simulations. Behavioural patterns were consistent with basking sharks using search tactics structured across multiple scales to exploit the richest prey areas available in preferred habitats. Simple behavioural rules based on learned responses to previously encountered prey distributions may explain the high performances. This study highlights how dynamic prey landscapes enable active habitat selection in large predators to be investigated from a trophic perspective, an approach that may inform conservation by identifying critical habitat of vulnerable species.

Ranking of length-class seasonal and regional effects on dietary compositions of the co-occurring Pagrus auratus (Sparidae) and Pseudocaranx georgianus (Carangidae)

Using an effective combination of multivariate testing and ordination analyses, this study compares the extents to which the diets of two co-occurring fish species (Pagrus auratus and Pseudocaranx georgianus) are related to body size (length class), season and region and the rank order importance of those effects. Thus, volumetric dietary compositions were determined for these species on the lower west coast of Australia, where both are abundant, and for P. auratus from the mid west coast and P. georgianus from the south coast. The diet of P. auratus on the lower west coast was strongly related to body size and slightly less to season. With increasing body size, its diet shifted from predominantly ophiuroids to larger prey, such as brachyuran crabs, teleosts, echinoids and ultimately asteroids, probably reflecting a shift from foraging over soft sediments to areas over and around reefs. Seasonal changes on the lower west coast were restricted mainly to small P. auratus, while larger fish underwent seasonal changes further north. Analyses using a common size range of medium to larger P. auratus demonstrated that dietary composition differed more between regions than seasons. The relationships between diet and length class of P. georgianus on both the lower west and south coasts were less pronounced than for P. auratus and seasonal changes were restricted to the south coast, where amphipod consumption increased markedly in summer. The diet of P. georgianus was related far more to region than length class and season, with more small teleosts, small crabs, carideans and littorinids and less amphipods, isopods and small bivalves being ingested on the lower west than south coasts. Although crabs and teleosts were important typifying prey of P. auratus and P. georgianus, when co-occurring, the former predator tended to ingest greater volumes of larger and often less mobile prey. This reflects differences in dentition, jaw morphology and feeding behaviour and reduces the potential for competition for food resources. The results imply that P. auratus and P. georgianus are opportunistic feeders and that the effects of length class, season and region on dietary composition and their rank orders can vary markedly between species and for length class and season between regions for the same species.

An innovative statistical approach to constructing a readily comprehensible food web for a demersal fish community

Many food webs are so complex that it is difficult to distinguish the relationships between predators and their prey. We have therefore developed an approach that produces a food web which clearly demonstrates the strengths of the relationships between the predator guilds of demersal fish and their prey guilds in a coastal ecosystem. Subjecting volumetric dietary data for 35 abundant predators along the lower western Australia coast to cluster analysis and the SIMPROF routine separated the various species x length class combinations into 14 discrete predator guilds. Following nMDS ordination, the sequence of points for these predator guilds represented a 'trophic' hierarchy. This demonstrated that, with increasing body size, several species progressed upwards through this hierarchy, reflecting a marked change in diet, whereas others remained within the same guild. A novel use of cluster analysis and SIMPROF then identified each group of prey that was ingested in a common pattern across the full suite of predator guilds. This produced 12 discrete groups of taxa (prey guilds) that each typically comprised similar ecological/functional prey, which were then also aligned in a hierarchy. The hierarchical arrangements of the predator and prey guilds were plotted against each other to show the percentage contribution of each prey guild to the diet of each predator guild. The resultant shade plot demonstrates quantitatively how food resources are spread among the fish species and revealed that two prey guilds, one containing cephalopods and teleosts and the other small benthic/epibenthic crustaceans and polychaetes, were consumed by all predator guilds.

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.

Diet of auklet chicks in the Aleutian Islands, Alaska: similarity among islands, interspecies overlap, and relationships to ocean climate

Seabirds are effective samplers of the marine environment, and can be used to measure resource partitioning among species and sites via food loads destined for chicks. We examined the composition, overlap, and relationships to changing climate and oceanography of 3,216 food loads from Least, Crested, and Whiskered Auklets (Aethia pusilla, A. cristatella, A. pygmaea) breeding in Alaska during 1994–2006. Meals comprised calanoid copepods (Neocalanus spp.) and euphausiids (Thysanoessa spp.) that reflect secondary marine productivity, with no difference among Buldir, Kiska, and Kasatochi islands across 585 km of the Aleutian Islands. Meals were very similar among species (mean Least–Crested Auklet overlap C = 0.68; Least–Whiskered Auklet overlap C = 0.96) and among sites, indicating limited partitioning of prey resources for auklets feeding chicks. The biomass of copepods and euphausiids in Least and Crested Auklet food loads was related negatively to the summer (June–July–August) North Pacific Gyre Oscillation, while in Whiskered Auklet food loads, this was negatively related to the winter (December–January–February) Pacific Decadal Oscillation, both of which track basin-wide sea-surface temperature (SST) anomalies. We found a significant quadratic relationship between the biomass of calanoid copepods in Least Auklet food loads at all three study sites and summer (June–July) SST, with maximal copepod biomass between 3–6°C (r 2 = 0.71). Outside this temperature range, zooplankton becomes less available to auklets through delayed development. Overall, our results suggest that auklets are able to buffer climate-mediated bottom-up forcing of demographic parameters like productivity, as the composition of chick meals has remained constant over the course of our study.

Scale-dependent foraging ecology of a marine top predator modelled using passive acoustic data

1.Understanding which environmental factors drive foraging preferences is critical for the development of effective management measures, but resource use patterns may emerge from processes that occur at different spatial and temporal scales. Direct observations of foraging are also especially challenging in marine predators, but passive acoustic techniques provide opportunities to study the behaviour of echolocating species over a range of scales. 2.We used an extensive passive acoustic data set to investigate the distribution and temporal dynamics of foraging in bottlenose dolphins using the Moray Firth (Scotland, UK). Echolocation buzzes were identified with a mixture model of detected echolocation inter-click intervals and used as a proxy of foraging activity. A robust modelling approach accounting for autocorrelation in the data was then used to evaluate which environmental factors were associated with the observed dynamics at two different spatial and temporal scales. 3.At a broad scale, foraging varied seasonally and was also affected by seabed slope and shelf-sea fronts. At a finer scale, we identified variation in seasonal use and local interactions with tidal processes. Foraging was best predicted at a daily scale, accounting for site specificity in the shape of the estimated relationships. 4.This study demonstrates how passive acoustic data can be used to understand foraging ecology in echolocating species and provides a robust analytical procedure for describing spatio-temporal patterns. Associations between foraging and environmental characteristics varied according to spatial and temporal scale, highlighting the need for a multi-scale approach. Our results indicate that dolphins respond to coarser scale temporal dynamics, but have a detailed understanding of finer-scale spatial distribution of resources.

Scaling laws of ambush predator 'waiting' behaviour are tuned to a common ecology

The decisions animals make about how long to wait between activities can determine the success of diverse behaviours such as foraging, group formation or risk avoidance. Remarkably, for diverse animal species, including humans, spontaneous patterns of waiting times show random ‘burstiness’ that appears scale-invariant across a broad set of scales. However, a general theory linking this phenomenon across the animal kingdom currently lacks an ecological basis. Here, we demonstrate from tracking the activities of 15 sympatric predator species (cephalopods, sharks, skates and teleosts) under natural and controlled conditions that bursty waiting times are an intrinsic spontaneous behaviour well approximated by heavy-tailed (power-law) models over data ranges up to four orders of magnitude. Scaling exponents quantifying ratios of frequent short to rare very long waits are species-specific, being determined by traits such as foraging mode (active versus ambush predation), body size and prey preference. A stochastic–deterministic decision model reproduced the empirical waiting time scaling and species-specific exponents, indicating that apparently complex scaling can emerge from simple decisions. Results indicate temporal power-law scaling is a behavioural ‘rule of thumb’ that is tuned to species’ ecological traits, implying a common pattern may have naturally evolved that optimizes move–wait decisions in less predictable natural environments.