Different currencies for calculating resource phenology result in opposite inferences about trophic mismatches.

Shifts in phenology are among the key responses of organisms to climate change. When rates of phenological change differ between interacting species they may result in phenological asynchrony. Studies have found conflicting patterns concerning the direction and magnitude of changes in synchrony, which have been attributed to biological factors. A hitherto overlooked additional explanation are differences in the currency used to quantify resource phenology, such as abundance and biomass. Studying an insectivorous bird (the sanderling) and its prey, we show that the median date of cumulative arthropod biomass occurred, on average, 6.9 days after the median date of cumulative arthropod abundance. In some years this difference could be as large as 21 days. For 23 years, hatch dates of sanderlings became less synchronized with the median date of arthropod abundance, but more synchronized with the median date of arthropod biomass. The currency-specific trends can be explained by our finding that mean biomass per arthropod specimen increased with date. Using a conceptual simulation, we show that estimated rates of phenological change for abundance and biomass can differ depending on temporal shifts in the size distribution of resources. We conclude that studies of trophic mismatch based on different currencies for resource phenology can be incompatible with each other.

Global biogeography of chemosynthetic symbionts reveals both localized and globally distributed symbiont groups.

In the ocean, most hosts acquire their symbionts from the environment. Due to the immense spatial scales involved, our understanding of the biogeography of hosts and symbionts in marine systems is patchy, although this knowledge is essential for understanding fundamental aspects of symbiosis such as host-symbiont specificity and evolution. Lucinidae is the most species-rich and widely distributed family of marine bivalves hosting autotrophic bacterial endosymbionts. Previous molecular surveys identified location-specific symbiont types that "promiscuously" form associations with multiple divergent cooccurring host species. This flexibility of host-microbe pairings is thought to underpin their global success, as it allows hosts to form associations with locally adapted symbionts. We used metagenomics to investigate the biodiversity, functional variability, and genetic exchange among the endosymbionts of 12 lucinid host species from across the globe. We report a cosmopolitan symbiont species, Candidatus Thiodiazotropha taylori, associated with multiple lucinid host species. Ca. T. taylori has achieved more success at dispersal and establishing symbioses with lucinids than any other symbiont described thus far. This discovery challenges our understanding of symbiont dispersal and location-specific colonization and suggests both symbiont and host flexibility underpin the ecological and evolutionary success of the lucinid symbiosis.

Arriving late and lean at a stopover site is selected against in a declining migratory bird population.

Loss and/or deterioration of refuelling habitats have caused population declines in many migratory bird species but whether this results from unequal mortality among individuals varying in migration traits remains to be shown. Based on 13 years of body mass and size data of great knots (Calidris tenuirostris) at a stopover site of the Yellow Sea, combined with resightings of individuals marked at this stopover site along the East Asian-Australasian Flyway, we assessed year to year changes in annual apparent survival rates, and how apparent survival differed between migration phenotypes (i.e. migration timing and fuel stores). The measurements occurred over a period of habitat loss and/or deterioration in this flyway. We found that the annual apparent survival rates of great knots rapidly declined from 2006 to 2018, late-arriving individuals with small fuel stores exhibiting the lowest apparent survival rate. There was an advancement in mean arrival date and an increase in the mean fuel load of stopping birds over the study period. Our results suggest that late-arriving individuals with small fuel loads were selected against. Thus, habitat loss and/or deterioration at staging sites may cause changes in the composition of migratory phenotypes at the population-level.

Mismatch-induced growth reductions in a clade of Arctic-breeding shorebirds are rarely mitigated by increasing temperatures.

In seasonal environments subject to climate change, organisms typically show phenological changes. As these changes are usually stronger in organisms at lower trophic levels than those at higher trophic levels, mismatches between consumers and their prey may occur during the consumers' reproduction period. While in some species a trophic mismatch induces reductions in offspring growth, this is not always the case. This variation may be caused by the relative strength of the mismatch, or by mitigating factors like increased temperature-reducing energetic costs. We investigated the response of chick growth rate to arthropod abundance and temperature for six populations of ecologically similar shorebirds breeding in the Arctic and sub-Arctic (four subspecies of Red Knot Calidris canutus, Great Knot C. tenuirostris and Surfbird C. virgata). In general, chicks experienced growth benefits (measured as a condition index) when hatching before the seasonal peak in arthropod abundance, and growth reductions when hatching after the peak. The moment in the season at which growth reductions occurred varied between populations, likely depending on whether food was limiting growth before or after the peak. Higher temperatures led to faster growth on average, but could only compensate for increasing trophic mismatch for the population experiencing the coldest conditions. We did not find changes in the timing of peaks in arthropod availability across the study years, possibly because our series of observations was relatively short; timing of hatching displayed no change over the years either. Our results suggest that a trend in trophic mismatches may not yet be evident; however, we show Arctic-breeding shorebirds to be vulnerable to this phenomenon and vulnerability to depend on seasonal prey dynamics.

Sulfur in lucinid bivalves inhibits intake rates of a molluscivore shorebird.

A forager's energy intake rate is usually constrained by a combination of handling time, encounter rate and digestion rate. On top of that, food intake may be constrained when a forager can only process a maximum amount of certain toxic compounds. The latter constraint is well described for herbivores with a limited tolerance to plant secondary metabolites. In sulfidic marine ecosystems, many animals host chemoautotrophic endosymbionts, which store sulfur compounds as an energy resource, potentially making their hosts toxic to predators. The red knot Calidris canutus canutus is a molluscivore shorebird that winters on the mudflats of Banc d'Arguin, where the most abundant bivalve prey Loripes orbiculatus hosts sulfide-oxidizing bacteria. In this system, we studied the potential effect of sulfur on the red knots' intake rates, by offering Loripes with various sulfur content to captive birds. To manipulate toxicity, we starved Loripes for 10 days by removing them from their symbiont's energy source sulfide. As predicted, we found lower sulfur concentrations in starved Loripes. We also included natural variation in sulfur concentrations by offering Loripes collected at two different locations. In both cases lower sulfur levels in Loripes resulted in higher consumption rates in red knots. Over time the red knots increased their intake rates on Loripes, showing their ability to adjust to a higher intake of sulfur.

Individual-Level Memory Is Sufficient to Create Spatial Segregation among Neighboring Colonies of Central Place Foragers.

AbstractCentral place foragers often segregate in space, even without signs of direct agonistic interactions. Using parsimonious individual-based simulations, we show that for species with spatial cognitive abilities, individual-level memory of resource availability can be sufficient to cause spatial segregation in the foraging ranges of colonial animals. The shapes of the foraging distributions are governed by commuting costs, the emerging distribution of depleted resources, and the fidelity of foragers to their colonies. When colony fidelity is weak and foragers can easily switch to colonies located closer to favorable foraging grounds, this leads to space partitioning with equidistant borders between neighboring colonies. In contrast, when colony fidelity is strong-for example, because larger colonies provide safety in numbers or individuals are unable to leave-it can create a regional imbalance between resource requirements and resource availability. This leads to nontrivial space-use patterns that propagate through the landscape. Interestingly, while better spatial memory creates more defined boundaries between neighboring colonies, it can lower the average intake rate of the population, suggesting a potential trade-off between an individual's attempt for increased intake and population growth rates.

Understanding spatial distributions: negative density-dependence in prey causes predators to trade-off prey quantity with quality.

Negative density-dependence is generally studied within a single trophic level, thereby neglecting its effect on higher trophic levels. The 'functional response' couples a predator's intake rate to prey density. Most widespread is a type II functional response, where intake rate increases asymptotically with prey density; this predicts the highest predator densities at the highest prey densities. In one of the most stringent tests of this generality to date, we measured density and quality of bivalve prey (edible cockles Cerastoderma edule) across 50 km² of mudflat, and simultaneously, with a novel time-of-arrival methodology, tracked their avian predators (red knots Calidris canutus). Because of negative density-dependence in the individual quality of cockles, the predicted energy intake rates of red knots declined at high prey densities (a type IV, rather than a type II functional response). Resource-selection modelling revealed that red knots indeed selected areas of intermediate cockle densities where energy intake rates were maximized given their phenotype-specific digestive constraints (as indicated by gizzard mass). Because negative density-dependence is common, we question the current consensus and suggest that predators commonly maximize their energy intake rates at intermediate prey densities. Prey density alone may thus poorly predict intake rates, carrying capacity and spatial distributions of predators.

How habitat-modifying organisms structure the food web of two coastal ecosystems.

The diversity and structure of ecosystems has been found to depend both on trophic interactions in food webs and on other species interactions such as habitat modification and mutualism that form non-trophic interaction networks. However, quantification of the dependencies between these two main interaction networks has remained elusive. In this study, we assessed how habitat-modifying organisms affect basic food web properties by conducting in-depth empirical investigations of two ecosystems: North American temperate fringing marshes and West African tropical seagrass meadows. Results reveal that habitat-modifying species, through non-trophic facilitation rather than their trophic role, enhance species richness across multiple trophic levels, increase the number of interactions per species (link density), but decrease the realized fraction of all possible links within the food web (connectance). Compared to the trophic role of the most highly connected species, we found this non-trophic effects to be more important for species richness and of more or similar importance for link density and connectance. Our findings demonstrate that food webs can be fundamentally shaped by interactions outside the trophic network, yet intrinsic to the species participating in it. Better integration of non-trophic interactions in food web analyses may therefore strongly contribute to their explanatory and predictive capacity.

Diet preferences as the cause of individual differences rather than the consequence.

Behavioural variation within a species is usually explained as the consequence of individual variation in physiology. However, new evidence suggests that the arrow of causality may well be in the reverse direction: behaviours such as diet preferences cause the differences in physiological and morphological traits. Recently, diet preferences were proposed to underlie consistent differences in digestive organ mass and movement patterns (patch residence times) in red knots (Calidris canutus islandica). Red knots are molluscivorous and migrant shorebirds for which the size of the muscular stomach (gizzard) is critical for the food processing rate. In this study, red knots (C. c. canutus, n = 46) were caught at Banc d'Arguin, an intertidal flat ecosystem in Mauritania, and released with radio-tags after the measurement of gizzard mass. Using a novel tracking system (time-of-arrival), patch residence times were measured over a period of three weeks. Whether or not gizzard mass determined patch residence times was tested experimentally by offering 12 of the 46 tagged red knots soft diets prior to release; this reduced an individual's gizzard mass by 20-60%. To validate whether the observed range of patch residence times would be expected from individual diet preferences, we simulated patch residence times as a function of diet preferences via a simple departure rule. Consistent with previous empirical studies, patch residence times in the field were positively correlated with gizzard mass. The slope of this correlation, as well as the observed range of patch residence times, was in accordance with the simulated values. The 12 birds with reduced gizzard masses did not decrease patch residence times in response to the reduction in gizzard mass. These findings suggest that diet preferences can indeed cause the observed among-individual variation in gizzard mass and patch residence times. We discuss how early diet experiences can have cascading effects on the individual expression of both behavioural and physiomorphic traits. This emphasizes that to understand the ecological consequences of individual differences, knowledge of the environment during development is required.

Natural selection by pulsed predation: survival of the thickest.

Selective predation can lead to natural selection in prey populations and may alleviate competition among surviving individuals. The processes of selection and competition can have substantial effects on prey population dynamics, but are rarely studied simultaneously. Moreover, field studies of predator-induced short-term selection pressures on prey populations are scarce. Here we report measurements of density dependence in body composition in a bivalve prey (edible cockle, Cerastoderma edule) during bouts of intense predation by an avian predator (Red Knot, Calidris canutus). We measured densities, patchiness, morphology, and body composition (shell and flesh mass) of cockles in a quasi-experimental setting, i.e., before and after predation in three similar plots of 1 ha each, two of which experienced predation, and one of which remained unvisited in the course of the short study period and served as a reference. An individual's shell and flesh mass declined with cockle density (negative density dependence). Before predation, cockles were patchily distributed. After predation, during which densities were reduced by 78% (from 232 to 50 cockles/m2), the patchiness was substantially reduced, i.e., the spatial distribution was homogenized. Red Knots selected juvenile cockles with an average length of 6.9 ± 1.0 mm (mean ± SD). Cockles surviving predation had heavier shells than before predation (an increase of 21.5 percentage points), but similar flesh masses. By contrast, in the reference plot shell mass did not differ statistically between initial and final sampling occasions, while flesh mass was larger (an increase of 13.2 percentage points). In this field study, we show that Red Knots imposed a strong selection pressure on cockles to grow fast with thick shells and little flesh mass, with selection gradients among the highest reported in the literature.

Ways to be different: foraging adaptations that facilitate higher intake rates in a northerly wintering shorebird compared with a low-latitude conspecific.

At what phenotypic level do closely related subspecies that live in different environments differ with respect to food detection, ingestion and processing? This question motivated an experimental study on rock sandpipers (Calidris ptilocnemis). The species' nonbreeding range spans 20 deg of latitude, the extremes of which are inhabited by two subspecies: C. p. ptilocnemis that winters primarily in upper Cook Inlet, Alaska (61°N) and C. p. tschuktschorum that overlaps slightly with C. p. ptilocnemis but whose range extends much farther south (∼40°N). In view of the strongly contrasting energetic demands of their distinct nonbreeding distributions, we conducted experiments to assess the behavioral, physiological and sensory aspects of foraging and we used the bivalve Macoma balthica for all trials. C. p. ptilocnemis consumed a wider range of prey sizes, had higher maximum rates of energy intake, processed shell waste at higher maximum rates and handled prey more quickly. Notably, however, the two subspecies did not differ in their abilities to find buried prey. The subspecies were similar in size and had equally sized gizzards, but the more northern ptilocnemis individuals were 10-14% heavier than their same-sex tschuktschorum counterparts. The higher body mass in ptilocnemis probably resulted from hypertrophy of digestive organs (e.g. intestine, liver) related to digestion and nutrient assimilation. Given the previously established equality of the metabolic capacities of the two subspecies, we propose that the high-latitude nonbreeding range of ptilocnemis rock sandpipers is primarily facilitated by digestive (i.e. physiological) aspects of their foraging ecology rather than behavioral or sensory aspects.

Field measurements give biased estimates of functional response parameters, but help explain foraging distributions.

Mechanistic insights and predictive understanding of the spatial distributions of foragers are typically derived by fitting either field measurements on intake rates and food abundance, or observations from controlled experiments, to functional response models. It has remained unclear, however, whether and why one approach should be favoured above the other, as direct comparative studies are rare. The field measurements required to parameterize either single or multi-species functional response models are relatively easy to obtain, except at sites with low food densities and at places with high food densities, as the former will be avoided and the second will be rare. Also, in foragers facing a digestive bottleneck, intake rates (calculated over total time) will be constant over a wide range of food densities. In addition, interference effects may depress intake rates further. All of this hinders the appropriate estimation of parameters such as the 'instantaneous area of discovery' and the handling time, using a type II functional response model also known as 'Holling's disc equation'. Here we compare field- and controlled experimental measurements of intake rate as a function of food abundance in female bar-tailed godwits Limosa lapponica feeding on lugworms Arenicola marina. We show that a fit of the type II functional response model to field measurements predicts lower intake rates (about 2.5 times), longer handling times (about 4 times) and lower 'instantaneous areas of discovery' (about 30-70 times), compared with measurements from controlled experimental conditions. In agreement with the assumptions of Holling's disc equation, under controlled experimental settings both the instantaneous area of discovery and the handling time remained constant with an increase in food density. The field data, however, would lead us to conclude that although handling time remains constant, the instantaneous area of discovery decreased with increasing prey densities. This will result into highly underestimated sensory capacities when using field data. Our results demonstrate that the elucidation of the fundamental mechanisms behind prey detection and prey processing capacities of a species necessitates measurements of functional response functions under the whole range of prey densities on solitary feeding individuals, which is only possible under controlled conditions. Field measurements yield 'consistency tests' of the distributional patterns in a specific ecological context.

Moving on with foraging theory: incorporating movement decisions into the functional response of a gregarious shorebird.

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.

Digestive capacity and toxicity cause mixed diets in red knots that maximize energy intake rate.

Among energy-maximizing animals, preferences for different prey can be explained by ranking the prey according to their energetic content. However, diet choice also depends on characteristics of the predator, such as the need to ingest necessary nutrients and the constraints imposed by digestion and toxins in food. In combination, these factors can lead to mixed diets in which the energetically most profitable food is not eaten exclusively even when it is abundant. We studied diet choice in red knots (Calidris canutus canutus) feeding on mollusks at a West African wintering site. At this site, the birds fed primarily on two species of bivalves, a thick-shelled one (Dosinia isocardia) that imposed a digestive constraint and a thin-shelled one (Loripes lucinalis) that imposed a toxin constraint. The latter species is toxic due to its symbiotic association with sulfide-oxidizing bacteria. We estimated experimentally the parameters of a linear programming model that includes both digestive and toxin constraints, leading to the prediction that red knots should eat a mixture of both mollusk species to maximize energy intake. The model correctly predicted the preferences of the captive birds, which depended on the digestive quality and toxicity of their previous diet. At our study site, energy-maximizing red knots appear to select a mixed diet as a result of the simultaneous effects of digestive and toxin constraints.

Personality drives physiological adjustments and is not related to survival.

The evolutionary function and maintenance of variation in animal personality is still under debate. Variation in the size of metabolic organs has recently been suggested to cause and maintain variation in personality. Here, we examine two main underlying notions: (i) that organ sizes vary consistently between individuals and cause consistent behavioural patterns, and (ii) that a more exploratory personality is associated with reduced survival. Exploratory behaviour of captive red knots (Calidris canutus, a migrant shorebird) was negatively rather than positively correlated with digestive organ (gizzard) mass, as well as with body mass. In an experiment, we reciprocally reduced and increased individual gizzard masses and found that exploration scores were unaffected. Whether or not these birds were resighted locally over the 19 months after release was negatively correlated with their exploration scores. Moreover, a long-term mark-recapture effort on free-living red knots with known gizzard masses at capture confirmed that local resighting probability (an inverse measure of exploratory behaviour) was correlated with gizzard mass without detrimental effects on survival. We conclude that personality drives physiological adjustments, rather than the other way around, and suggest that physiological adjustments mitigate the survival costs of exploratory behaviour. Our results show that we need to reconsider hypotheses explaining personality variation based on organ sizes and differential survival.

Toxin constraint explains diet choice, survival and population dynamics in a molluscivore shorebird.

Recent insights suggest that predators should include (mildly) toxic prey when non-toxic food is scarce. However, the assumption that toxic prey is energetically as profitable as non-toxic prey misses the possibility that non-toxic prey have other ways to avoid being eaten, such as the formation of an indigestible armature. In that case, predators face a trade-off between avoiding toxins and minimizing indigestible ballast intake. Here, we report on the trophic interactions between a shorebird (red knot, Calidris canutus canutus) and its two main bivalve prey, one being mildly toxic but easily digestible, and the other being non-toxic but harder to digest. A novel toxin-based optimal diet model is developed and tested against an existing one that ignores toxin constraints on the basis of data on prey abundance, diet choice, local survival and numbers of red knots at Banc d'Arguin (Mauritania) over 8 years. Observed diet and annual survival rates closely fit the predictions of the toxin-based model, with survival and population size being highest in years when the non-toxic prey is abundant. In the 6 of 8 years when the non-toxic prey is not abundant enough to satisfy the energy requirements, red knots must rely on the toxic alternative.

Economic design in a long-distance migrating molluscivore: how fast-fuelling red knots in Bohai Bay, China, get away with small gizzards.

We carried out an observational and experimental study to decipher how resource characteristics, in interaction with the predator's phenotype, constrain a fitness-determining performance measure, i.e. refuelling in a migrant bird. Two subspecies of red knot (Calidris canutus rogersi and C. c. piersmai) use northern Bohai Bay, Yellow Sea, China, for the final prebreeding stopover, during their 10,000-15,000 km long migrations between wintering and breeding areas. Here, they feed on small bivalves, especially 2-7 mm long Potamocorbula laevis. With an average stay of 29 days, and the need to store 80 g of fat for the onward flights to high-Arctic breeding grounds, red knots need to refuel fast. Using existing knowledge, we expected them to achieve this on the basis of (1) prey with high flesh to shell mass ratios, (2) large gizzards to crush the ingested molluscs, or (3) a combination of the two. Rejecting all three predictions, we found that red knots staging in Bohai Bay had the smallest gizzards on record (4.9 ± 0.8 g, mean ± s.e.m., N = 27), and also found that prey quality of P. laevis is much lower than predicted for the measured gizzard size (i.e. 1.3 rather than the predicted 4.5 kJ g(-1) dry shell mass, DM(shell)). The estimated handling time of P. laevis (0.2 s) is much shorter than the observed time between two prey ingestions (0.7 s), indicating that prey handling time is no constraint. Based on field observations of dropping rates and on indoor digestion trails, the shell processing rate was estimated at 3.9 mg DM(shell) s(-1), i.e. three times higher the rate previously predicted for red knots eating as fast as they can with the measured gizzard size. This is explained by the small and easily crushed P. laevis enabling high processing rates. As P. laevis also occurred in high densities, the metabolizable energy intake rate of red knots with small gizzards at 5 J s(-1) was as high as at northward staging sites elsewhere in the world. Currently, therefore, food characteristics in Bohai Bay are such that red knots can refuel fast whilst economizing on the size of their gizzard. These time-stressed migrants thus provide an elegant example of symmorphosis.

Site-specific length-biomass relationships of arctic arthropod families are critical for accurate ecological inferences.

Arthropods play a crucial role in terrestrial ecosystems, for instance in mediating energy fluxes and in forming the food base for many organisms. To better understand their functional role in such ecosystem processes, monitoring of trends in arthropod biomass is essential. Obtaining direct measurements of the body mass of individual specimens is laborious. Therefore, these data are often indirectly acquired by utilizing allometric length-biomass relationships based on a correlative parameter, such as body length. Previous studies have often used such relationships with a low taxonomic resolution and/or small sample size and/or adopted regressions calibrated in different biomes. Despite the scientific interest in the ecology of arctic arthropods, no site-specific family-level length-biomass relationships have hitherto been published. Here we present 27 family-specific length-biomass relationships from two sites in the High Arctic: Zackenberg in northeast Greenland and Knipovich in north Taimyr, Russia. We show that length-biomass regressions from different sites within the same biome did not affect estimates of phenology but did result in substantially different estimates of arthropod biomass. Estimates of daily biomass at Zackenberg were on average 24% higher when calculated using regressions for Knipovich compared to using regressions for Zackenberg. In addition, calculations of daily arthropod biomass at Zackenberg based on order-level regressions from frequently cited studies in literature revealed overestimations of arthropod biomass ranging from 69.7% to 130% compared to estimates based on regressions for Zackenberg. Our results illustrate that the use of allometric relationships from different sites can significantly alter the biological interpretation of, for instance, the interaction between insectivorous birds and their arthropod prey. We conclude that length-biomass relationships should be locally established rather than being based on global relationships.

Trophic cascade induced by molluscivore predator alters pore-water biogeochemistry via competitive release of prey.

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.

Scaling up ideals to freedom: are densities of red knots across western Europe consistent with ideal free distribution?

Local studies have shown that the distribution of red knots Calidris canutus across intertidal mudflats is consistent with the predictions of an ideal distribution, but not a free distribution. Here, we scale up the study of feeding distributions to their entire wintering area in western Europe. Densities of red knots were compared among seven wintering sites in The Netherlands, UK and France, where the available mollusc food stocks were also measured and from where diets were known. We tested between three different distribution models that respectively assumed (i) a uniform distribution of red knots over all areas, (ii) a uniform distribution across all suitable habitat (based on threshold densities of harvestable mollusc prey), and (iii) an ideal and free distribution (IFD) across all suitable habitats. Red knots were not homogeneously distributed across the different European wintering areas, also not when considering suitable habitats only. Their distribution was best explained by the IFD model, suggesting that the birds are exposed to interference and have good knowledge about their resource landscape at the spatial scale of NW Europe, and that the costs of movement between estuaries, at least when averaged over a whole winter, are negligible.