Résumé: Understanding the life cycle strategies and predatory impact of alien jellyfish species is critical to mitigate the impact that these organisms may have on local populations, biodiversity, and ultimately on the functioning of food webs. In the Mediterranean Sea, little is known about the dynamics of alien jellyfish, despite this biodiversity hotspot being one of the most threatened areas by increasing numbers of alien jellyfish. Here, we investigated the population dynamics and predatory impact of a non-indigenous scyphomedusa, Aurelia solida Browne 1905, in the Bizerte Lagoon, Tunisia. The study was based on bimonthly surveys performed over two consecutive years, from November 2012 to August 2014. Field observations showed that the planktonic phase of A. solida occurs from winter to early summer. Prey composition was investigated by means of gut content and field zooplankton analyses. Calanoid copepods, mollusc larvae, and larvaceans represented the main food items of A. solida. To determine the jellyfish feeding rate and their predatory impact on zooplankton populations, the digestion time for zooplankton prey was assessed at three different temperatures: 13, 18, and 23 degrees C in laboratory conditions, corresponding to the average range of temperatures encountered by A. solida in the Bizerte Lagoon. We found that A. solida consumed 0.5-22.5% and 0.02-37.3% of the daily zooplankton standing stock in 2013 and 2014, respectively. These results indicate a non-negligible but restricted seasonal grazing impact on some mesozooplankton groups, explained by the relatively short lifespan of the medusa stage (5-6 months).

Résumé: Until recently, jellyfish have been ignored as an important source of food, due to their low nutritional value. Here, quantitative PCR was used to detect and quantify the DNA of the jellyfish Aurelia coerulea in the gut contents of commercially important fish species from the Thau Lagoon. Individuals from five fish species were collected during two different periods: the bloom period, when the pelagic stages of A. coerulea are abundant, and the post-bloom period, when only the benthic stage – polyps – is present in the lagoon. The DNA of A. coerulea was detected in the guts of 41.9% of the fish analysed, belonging to four different species. The eel Anguilla anguilla and the seabream Sparus aurata were important jellyfish consumers during the bloom and post-bloom periods, respectively. These results provide new insights on the potential control of jellyfish populations and on jellyfish importance as a food source for exploited fishes.

Résumé: Aurelia aurita (Linneaus, 1758) is a cosmopolitan scyphozoan, probably the most investigated jellyfish in temperate and highly productive coastal ecosystems. Despite a prominent top-down control in plankton food webs, a mechanistic understanding of A. aurita population dynamics and trophic interactions has been barely addressed. Here we develop a food web dynamic model to assess A. aurita role in the seasonal plankton dynamics of the Kiel Fjord, southwestern Baltic Sea. The model couples low trophic level dynamics, based on a classical Nutrient Phytoplankton Zooplankton Detritus (NPZD) model, to a stage-resolved copepod model (referencing Pseudocalanus sp.) and a jellyfish model (A. aurita ephyra and medusa) as consumers and predators, respectively. Simulations showed the relevance of high abundances of A. aurita, which appear related with warm winter temperatures, promoting a shift from a copepod-dominated food web to a ciliate and medusa dominated one. The model captured the intraspecific competition triggered by the medusae abundance and characterized by a negative relationship between population density and individual size/weight. Our results provide a mechanistic understanding of an emergent trait such as size shaping the food web functioning, driving predation rates and population dynamics of A. aurita, driving its sexual reproductive strategy at the end of the pelagic phase.

Résumé: The costs and benefits of group living often depend on the spatial position of individuals within groups and the ability of individuals to occupy preferred positions. For example, models of predation events for moving prey groups predict higher mortality risk for individuals at the periphery and front of groups. We investigated these predictions in sardine (Sardinella aurita) schools under attack from group hunting sailfish (Istiophorus platypterus) in the open ocean. Sailfish approached sardine schools about equally often from the front and rear, but prior to attack there was a chasing period in which sardines attempted to swim away from the predator. Consequently, all sailfish attacks were directed at the rear and peripheral positions of the school, resulting in higher predation risk for individuals at these positions. During attacks, sailfish slash at sardines with their bill causing prey injury including scale removal and tissue damage. Sardines injured in previous attacks were more often found in the rear half of the school than in the front half. Moreover, injured fish had lower tail-beat frequencies and lagged behind uninjured fish. Injuries inflicted by sailfish bills may, therefore, hinder prey swimming speed and drive spatial sorting in prey schools through passive self-assortment. We found only partial support for the theoretical predictions from current predator-prey models, highlighting the importance of incorporating more realistic predator-prey dynamics into these models. This article is part of the themed issue 'Physiological determinants of social behaviour in animals'.

Résumé: The invasive ctenophore Mnemiopsis leidyi has been reported in various coastal locations in the southern North Sea in the past years. Since 2009, International Bottom Trawl Surveys have recorded this species each winter in open waters. As this species, well-known for its dramatic disturbance of ecosystems, was expected not to be able to overwinter offshore it is crucial to understand its distribution dynamics. Two modelling methods, a quantile regression and a particle tracking model, were used (1) to identify habitats where the invasive ctenophore M. leidyi could survive the North Sea cold winters and (2) to investigate the dispersal of individuals between these different habitats, emphasizing favorable areas where sustainable populations could have been established. Temperature was found to be the crucial factor controlling the winter distribution of M. leidyi in the North Sea. High abundance predictions in winter were associated with low values of temperature, which characterise south-eastern coastal areas and estuaries influenced by riverine runoff. A retention-based M. leidyi population was indicated along the northern Dutch coast and German Bight and a transport-based population offshore from the western Danish coast. Individuals found in the open waters were transported from southern coasts of the North Sea, thus the open water population densities depend on the flux of offspring from these areas. This study provides the first estimates of the overwinter areas of this invasive species over the cold winters in the North Sea. Based on the agreement of habitat and dispersal model results, we conclude that M. leidyi has become established along south-eastern coasts of the North Sea where the environment conditions allows overwintering and it can be retained for later blooms.

Résumé: 1. Models relating intake rate to food abundance and competitor density (generalized functional response models) can predict forager distributions and movements between patches, but we lack understanding of how distributions and small-scale movements by the foragers themselves affect intake rates. Using a state-of-the-art approach based on continuous-time Markov chain dynamics, we add realism to classic functional response models by acknowledging that the chances to encounter food and competitors are influenced by movement decisions, and, vice versa, that movement decisions are influenced by these encounters. We used a multi-state modelling framework to construct a stochastic functional response model in which foragers alternate between three behavioural states: searching, handling and moving. Using behavioural observations on a molluscivore migrant shorebird (red knot, Calidris canutus canutus), at its main wintering area (Banc d'Arguin, Mauritania), we estimated transition rates between foraging states as a function of conspecific densities and densities of the two main bivalve prey. Intake rate decreased with conspecific density. This interference effect was not due to decreased searching efficiency, but resulted from time lost to avoidance movements. Red knots showed a strong functional response to one prey (Dosinia isocardia), but a weak response to the other prey (Loripes lucinalis). This corroborates predictions from a recently developed optimal diet model that accounts for the mildly toxic effects due to consuming Loripes. Using model averaging across the most plausible multi-state models, the fully parameterized functional response model was then used to predict intake rate for an independent data set on habitat choice by red knot. Comparison of the sites selected by red knots with random sampling sites showed that the birds fed at sites with higher than average Loripes and Dosinia densities, that is sites for which we predicted higher than average intake rates. We discuss the limitations of Holling's classic functional response model which ignores movement and the limitations of contemporary movement ecological theory that ignores consumer-resource interactions. With the rapid advancement of technologies to track movements of individual foragers at fine spatial scales, the time is ripe to integrate descriptive tracking studies with stochastic movement-based functional response models.

Résumé: Effects of predation may cascade down the food web. By alleviating interspecific competition among prey, predators may promote biodiversity, but the precise mechanisms of how predators alter competition have remained elusive. Here we report on a predator-exclosure experiment carried out in a tropical intertidal ecosystem, providing evidence for a three-level trophic cascade induced by predation by molluscivore Red Knots (Calidris canutus) that affects pore water biogeochemistry. In the exclosures the knots' favorite prey (Dosinia isocardia) became dominant and reduced the individual growth rate in an alternative prey (Loripes lucinalis). Dosinia, a suspension feeder, consumes suspended particulate organic matter (POM), whereas Loripes is a facultative mixotroph, partly living on metabolites produced by sulfur-oxidizing chemoautotrophic bacteria, but also consuming suspended POM. Reduced sulfide concentrations in the exclosures suggest that, without predation on Dosinia, stronger competition for suspended POM forces Loripes to rely on energy produced by endosymbiotic bacteria, thus leading to an enhanced uptake of sulfide from the surrounding pore water. As sulfide is toxic to most organisms, this competition-induced diet shift by Loripes may detoxify the environment, which in turn may facilitate other species. The inference that predators affect the toxicity of their environment via a multi-level trophic cascade is novel, but we believe it may be a general phenomenon in detritus-based ecosystems.

Résumé: A statistical catch-at-age model was developed to assess the effects of predation by the northwest Atlantic harp seal population on northern Gulf of St. Lawrence cod by estimating the relative importance of different sources of mortality that affected the stock during a period of collapse and non-recovery. Cod recruitment at age 1 is modeled via a non-linear stock-recruitment relationship based on total egg production and accounts for changes in female length-at-maturity and cod condition. Natural mortality other than seal predation also depends on cod condition used as an integrative index of changes in environmental conditions. The linkage between seals and cod is modeled through a multi-age functional response that was derived from the reconstruction of the seal diet using morphometric relationships and stomach contents of more than 200 seals collected between 1998 and 2001. The model was fitted following a maximum likelihood estimation approach to a scientific survey abundance index (1984 to 2006). Model results show that the collapse of the northern Gulf of St. Lawrence cod stock was mainly due to the combination of high fishing mortality rates and poor environmental conditions in the early to mid-1990s contributing to the current state of recruitment overfishing. The increase in harp seal abundance during 1984 to 2006 was reflected by an increase in predation mortality for the young cod age-groups targeted by seals. Although current levels of predation mortality affect cod spawning biomass, the lack of recovery of the NGSL cod stock seems mainly due to the very poor recruitment.

Résumé: The trophic level (TL) mean and variance, and the degree of omnivory for five Celtic Sea fish predators were estimated using a database of stomach content records characterized by a high level of taxonomic resolution. The predators occupied a high position in the food web, i.e. 4·75 for Atlantic cod Gadus morhua, 4·44 for haddock Melanogrammus aeglefinus, 4·88 for European hake Merluccius merluccius, 5·00 for megrim Lepidorhombus whiffiagonis and 5·27 for whiting Merlangius merlangus. The level of taxonomic resolution of the prey did not greatly affect mean TL predator values; an effect on variance was evident, low resolution masking intra-population variability in TL. Generalized additive models (GAM) were used to explain the variability of predator TL caused by environmental variables (International Council for the Exploration of the Sea, ICES, division and season) and predator characteristics (total length, LT). Significant year, location season and interaction effects were found for some species and with LT at the scale of ICES subdivision. The species-specific variability of TL could be due to spatio-temporal variations in prey availability and in predator selectivity following ontogenetic changes. Omnivorous fish TL was less affected by spatio-temporal variations. In addition, results showed that the omnivory index and TL variability provide dissimilar information on predator feeding strategy. Combining information on TL variability and omnivory allowed between within-individual and between-individual components contributing to trophic niche width to be separated and the type of generalization of fish predators to be identified.