Massive consumption of gelatinous plankton by Mediterranean apex predators

Stable isotopes of carbon and nitrogen were used to test the hypothesis that stomach content analysis has systematically overlooked the consumption of gelatinous zooplankton by pelagic mesopredators and apex predators. The results strongly supported a major role of gelatinous plankton in the diet of bluefin tuna (Thunnus thynnus), little tunny (Euthynnus alletteratus), spearfish (Tetrapturus belone) and swordfish (Xiphias gladius). Loggerhead sea turtles (Caretta caretta) in the oceanic stage and ocean sunfish (Mola mola) also primarily relied on gelatinous zooplankton. In contrast, stable isotope ratios ruled out any relevant consumption of gelatinous plankton by bluefish (Pomatomus saltatrix), blue shark (Prionace glauca), leerfish (Lichia amia), bonito (Sarda sarda), striped dolphin (Stenella caerueloalba) and loggerhead sea turtles (Caretta caretta) in the neritic stage, all of which primarily relied on fish and squid. Fin whales (Balaenoptera physalus) were confirmed as crustacean consumers. The ratios of stable isotopes in albacore (Thunnus alalunga), amberjack (Seriola dumerili), blue butterfish (Stromaeus fiatola), bullet tuna (Auxis rochei), dolphinfish (Coryphaena hyppurus), horse mackerel (Trachurus trachurus), mackerel (Scomber scombrus) and pompano (Trachinotus ovatus) were consistent with mixed diets revealed by stomach content analysis, including nekton and crustaceans, but the consumption of gelatinous plankton could not be ruled out completely. In conclusion, the jellyvorous guild in the Mediterranean integrates two specialists (ocean sunfish and loggerhead sea turtles in the oceanic stage) and several opportunists (bluefin tuna, little tunny, spearfish, swordfish and, perhaps, blue butterfish), most of them with shrinking populations due to overfishing.

Trace me if you can: the use of intrinsic biogeochemical markers in marine top predators

Human activities have serious impacts on marine apex predators. Inadequate knowledge of the spatial and trophic ecology of these marine animals ultimately compromises the viability of their populations and impedes our ability to use them as environmental biomonitors. Intrinsic biogeochemical markers, such as stable isotopes, fatty acids, trace elements, and chemical pollutants, are increasingly being used to trace the spatial and trophic ecology of marine top predators. Notable advances include the emergence of the first oceanographic"isoscapes" (isotopic geographic gradients), the advent of compound-specific isotopic analyses, improvements in diet reconstruction through Bayesian statistics, and tissue analysis of tracked animals to ground-truth biogeochemical profiles. However, most researchers still focus on only a few tracers. Moreover, insufficient knowledge of the biogeochemical integration in tissues, fractionation and routing processes, and geographic and temporal variability in baseline levels continue to hamper the resolution and potential of these markers in studying the spatial and feeding ecology of top predators.

Feeding ecology and movements of the Barolo shearwater Puffinus baroli baroli in the Azores, NE Atlantic

Trophic ecology and movements are critical issues for understanding the role of marine predators in food webs and for facing the challenges of their conservation. Seabird foraging ecology has been increasingly studied, but small elusive species, such as those forming the"little shearwater" complex, remain poorly known. We present the first study on the movements and feeding ecology of the Barolo shearwater Puffinus baroli baroli in a colony from the Azores archipelago (NE Atlantic), combining global location-sensing units, stable isotope analyses of feathers (δ13C and δ15N), stomach flushings and data from maximum depth gauges. During the chick-rearing period, parents visited their nests most nights, foraged mainly south of the colony and fed at lower trophic levels than during the non-breeding period. Squid was the most diverse prey (6 families and at least 10 different taxa), but species composition varied considerably between years. Two squid families, Onychoteuthidae and Argonautidae, and the fish family Phycidae accounted for 82.3% of ingested prey by number. On average, maximum dive depths per foraging trip reached 14.8 m (range: 7.9 to 23.1 m). After the breeding period, birds dispersed offshore in all directions and up to 2500 km from the breeding colony, and fed at higher trophic levels. Overall, our results indicate that the Barolo shearwater is a non-migratory shearwater feeding at the lowest trophic level among Macaronesian seabirds, showing both diurnal and nocturnal activity and feeding deeper in the water column, principally on small schooling squid and fish. These traits contrast with those of 3 other Azorean Procellariiformes (Cory"s shearwater Calonectris diomedea, the Madeiran storm-petrel Oceanodroma castro and Monteiro"s storm-petrel O. monteiroi), indicating ecological segregation within the Azorean seabird community.

Longer and less overlapping food webs in anthropogenically disturbed marine ecosystems: confirmations from the past

The human exploitation of marine resources is characterised by the preferential removal of the largest species. Although this is expected to modify the structure of food webs, we have a relatively poor understanding of the potential consequences of such alteration. Here, we take advantage of a collection of ancient consumer tissues, using stable isotope analysis and SIBER to assess changes in the structure of coastal marine food webs in the South-western Atlantic through the second half of the Holocene as a result of the sequential exploitation of marine resources by hunter-gatherers, western sealers and modern fishermen. Samples were collected from shell middens and museums. Shells of both modern and archaeological intertidal herbivorous molluscs were used to reconstruct changes in the stable isotopic baseline, while modern and archaeological bones of the South American sea lion Otaria flavescens, South American fur seal Arctocephalus australis and Magellanic penguin Spheniscus magellanicus were used to analyse changes in the structure of the community of top predators. We found that ancient food webs were shorter, more redundant and more overlapping than current ones, both in northern-central Patagonia and southern Patagonia. These surprising results may be best explained by the huge impact of western sealing on pinnipeds during the fur trade period, rather than the impact of fishing on fish populations. As a consequence, the populations of pinnipeds at the end of the sealing period were likely well below the ecosystem's carrying capacity, which resulted in a release of intraspecific competition and a shift towards larger and higher trophic level prey. This in turn led to longer and less overlapping food webs.

Small but powerful: Top predator local extinction affects ecosystem structure and function in an intermittent stream.

Top predator loss is a major global problem, with a current trend in biodiversity loss towards high trophic levels that modifies most ecosystems worldwide. Most research in this area is focused on large-bodied predators, despite the high extinction risk of small-bodied freshwater fish that often act as apex consumers. Consequently, it remains unknown if intermittent streams are affected by the consequences of top-predators' extirpations. The aim of our research was to determine how this global problem affects intermittent streams and, in particular, if the loss of a small-bodied top predator (1) leads to a 'mesopredator release', affects primary consumers and changes whole community structures, and (2) triggers a cascade effect modifying the ecosystem function. To address these questions, we studied the topdown effects of a small endangered fish species, Barbus meridionalis (the Mediterranean barbel), conducting an enclosure/exclosure mesocosm experiment in an intermittent stream where B. meridionalis became locally extinct following a wildfire.We found that top predator absence led to 'mesopredator release', and also to 'prey release' despite intraguild predation, which contrasts with traditional food web theory. In addition, B. meridionalis extirpation changed whole macroinvertebrate community composition and increased total macroinvertebrate density. Regarding ecosystem function, periphyton primary production decreased in apex consumer absence. In this study, the apex consumer was functionally irreplaceable; its local extinction led to the loss of an important functional role that resulted in major changes to the ecosystem's structure and function. This study evidences that intermittent streams can be affected by the consequences of apex consumers' extinctions, and that the loss of small-bodied top predators can lead to large ecosystem changes. We recommend the reintroduction of small-bodied apex consumers to systems where they have been extirpated, to restore ecosystem structure and function.

Consequences of warming and resource quality on the stoichiometry and nutrient cycling of a stream shredder

As a result of climate change, streams are warming and their runoff has been decreasing in most temperate areas. These changes can affect consumers directly by increasing their metabolic rates and modifying their physiology and indirectly by changing the quality of the resources on which organisms depend. In this study, a common stream detritivore (Echinogammarus berilloni Catta) was reared at two temperatures (15 and 20°C) and fed Populus nigra L. leaves that had been conditioned either in an intermittent or permanent reach to evaluate the effects of resource quality and increased temperatures on detritivore performance, stoichiometry and nutrient cycling. The lower quality (i.e., lower protein, soluble carbohydrates and higher C:P and N:P ratios) of leaves conditioned in pools resulted in compensatory feeding and lower nutrient retention capacity by E. berilloni. This effect was especially marked for phosphorus, which was unexpected based on predictions of ecological stoichiometry. When individuals were fed pool-conditioned leaves at warmer temperatures, their growth rates were higher, but consumers exhibited less efficient assimilation and higher mortality. Furthermore, the shifts to lower C:P ratios and higher lipid concentrations in shredder body tissues suggest that structural molecules such as phospholipids are preserved over other energetic C-rich macromolecules such as carbohydrates. These effects on consumer physiology and metabolism were further translated into feces and excreta nutrient ratios. Overall, our results show that the effects of reduced leaf quality on detritivore nutrient retention were more severe at higher temperatures because the shredders were not able to offset their increased metabolism with increased consumption or more efficient digestion when fed pool-conditioned leaves. Consequently, the synergistic effects of impaired food quality and increased temperatures might not only affect the physiology and survival of detritivores but also extend to other trophic compartments through detritivore-mediated nutrient cycling.

Functional diversity of macrophyte communities within and between Pyrenean lakes

Submersed vegetation is a common feature in about 70% Pyrene an high mountain (>1500 m a.s.l.) lakes. Isoetids and soft-water elodeids are common elements of this underw ater flora and can form distinct vegetation units (i.e. patches of vegetation dominated by different species) within complex mosaics of vegetation in shallow waters (<7 m). Since is oetids exert a strong influence on sediment biogeochemistry due to high radial oxygen loss, we examined the small scale characteristics of the lake environment (water and sediment) associated to vegetation patches in order to ascertain potential functional differences among them. To do so, we characterised the species composition and biomass of the main vegetation units from 11 lakes, defined plant communities based on biomass data, and then related each community with sediment properties (redox and dissolved nutrient concentration in the pore water) and water nutrient concentration within plant canopy. We also characterised lake water and sediment in areas without vegetation as a reference. A total of twenty-one vegetation units were identified, ranging from one to five per lake. A cluster analysis on biomass species composition suggested seven different macrophyte communities that were named after the most dominant species:Nitella sp.,Potamogeton praelongus, Myriophyllum alterniflorum, Sparganium angustifolium , Isoetes echinospora,Isoetes lacustris and Carex rostrata . Coupling between macrophyte communities and their immediate environment (overlying water and sediment) was manifested mainly as variation in sediment redox conditions and the dominant form of inorganic nitrogen in pore-water. These effects depended on the specific compositi on of the community, and on the allocation between above- and belowground biomass, and could be predicted with a model relating the average and standard deviation of sediment redox potential from 0 down to -20 cm, across macrophyte communities. Differences in pore-water total dissolved phosphorus were related to the trophic state of the lakes. There was no correlation between sediment and water column dissolved nutrients. However, nitrate concentrations tended to be lower in the water overlaying isoetid communities, in apparent contradiction to the patterns of dissolved nitrates in the pore-water. These tendencies were robust even when comparing the water over laying communities within the same lake, thus pointing towards a potential effect of isoetids in reducing dissolved nitrogen in the lakes.

Sexual foraging segregation in South American sea lions increases during the pre-breeding period in the La Plata River plume

Stable carbon and nitrogen isotopes in skin and bone of South American sea lions from Brazil and Uruguay were analysed to test the hypothesis that trophic overlap between the sexes is lower during the pre-breeding season than throughout the rest of the year. The isotopic values of skin and bone were used to infer the trophic relationships between the sexes during the pre-breeding period and year round, respectively. Prey species were also analysed to establish a baseline necessary for interpreting the stable isotope ratios of skin and bone. Standard ellipse areas, estimated using Bayesian inference in the SIBER routine of the SIAR package in R, suggested that males and females used a wide diversity of foraging strategies throughout the year and that no differences existed between the sexes. However, the diversity of foraging strategies was largely reduced during the pre-breeding period, with all the individuals of each sex adopting similar strategies, but with the two sexes differing considerably in stable isotope values and the ellipse areas of males and females not overlapping at all. Nevertheless, the results revealed a general increase in the consumption of pelagic prey by both sexes during the pre-breeding period. The progressive crowding of individuals in the areas surrounding the breeding rookeries during the pre-breeding period could lead to an increase in the local population density, which could explain the above reported changes.

Effects of Temperature, Salinity and Fish in Structuring the Macroinvertebrate Community in Shallow Lakes: Implications for Effects of Climate Change

Climate warming may lead to changes in the trophic structure and diversity of shallow lakes as a combined effect of increased temperature and salinity and likely increased strength of trophic interactions. We investigated the potential effects of temperature, salinity and fish on the plant-associated macroinvertebrate community by introducing artificial plants in eight comparable shallow brackish lakes located in two climatic regions of contrasting temperature: cold-temperate and Mediterranean. In both regions, lakes covered a salinity gradient from freshwater to oligohaline waters. We undertook day and night-time sampling of macroinvertebrates associated with the artificial plants and fish and free-swimming macroinvertebrate predators within artificial plants and in pelagic areas. Our results showed marked differences in the trophic structure between cold and warm shallow lakes. Plant-associated macroinvertebrates and free-swimming macroinvertebrate predators were more abundant and the communities richer in species in the cold compared to the warm climate, most probably as a result of differences in fish predation pressure. Submerged plants in warm brackish lakes did not seem to counteract the effect of fish predation on macroinvertebrates to the same extent as in temperate freshwater lakes, since small fish were abundant and tended to aggregate within the macrophytes. The richness and abundance of most plant-associated macroinvertebrate taxa decreased with salinity. Despite the lower densities of plant-associated macroinvertebrates in the Mediterranean lakes, periphyton biomass was lower than in cold temperate systems, a fact that was mainly attributed to grazing and disturbance by fish. Our results suggest that, if the current process of warming entails higher chances of shallow lakes becoming warmer and more saline, climatic change may result in a decrease in macroinvertebrate species richness and abundance in shallow lakes

Hydrology, light and the use of organic and inorganic materials as structuring factors of biological communities in Mediterranean streams

Hydrological disturbances, light availability and nutrients are the most relevant factors determining the structure of the biological communities in Mediterranean rivers. While some hydrological disturbances are able to induce catastrophic effects, which may cause a complete reset in physical and biological conditions, continued enrichment or changes in light availability are factors leading to the progressive shift in the communities of autotrophs and heterotrophs in the systems. Primary production in Mediterranean streams shows relevant seasonal changes which mainly follows the variations in light availability. In most forested streams, the algal community is shade-adapted. Nutrient enrichment (especially phosphorus) leads to marked increases in primary production, but this increase is not lineal and there is a saturation of algal biomass even in the most enriched systems. The heterotrophs (bacteria, fungi) are related to the pattern of DOC availability (which most depends on the seasonal discharge and leaf fall dynamics) and to the available substrata in the stream. It has been repeatedly observed that shorttime increases of extracellular enzyme activities are related to the accumulation of autochthonous (algal) and/or allochthonous (leaves) organic matter on the streambed during spring and summer, this being more remarkable in dry than in wetter years. Flow reduction favours detritus concentration in pools, and the subsequent increase in the density and biomass of the macroinvertebrate community. In Mediterranean streams collectors are accounting for the highest density and biomass, this being more remarkable in the least permanent systems, in accordance with the effect of floods on the organic matter availability. Nutrients, through the effect on the primary producers, also affect the trophic food web in the streams by favouring the predominance of grazers

Factors influencing zooplankton size structure at contrasting temperatures in coastal shallow lakes: Implications for effects of climate change

We assessed the importance of temperature, salinity, and predation for the size structure of zooplankton and provided insight into the future ecological structure and function of shallow lakes in a warmer climate. Artificial plants were introduced in eight comparable coastal shallow brackish lakes located at two contrasting temperatures: cold-temperate and Mediterranean climate region. Zooplankton, fish, and macroinvertebrates were sampled within the plants and at open-water habitats. The fish communities of these brackish lakes were characterized by small-sized individuals, highly associated with submerged plants. Overall, higher densities of small planktivorous fish were recorded in the Mediterranean compared to the cold-temperate region, likely reflecting temperature-related differences as have been observed in freshwater lakes. Our results suggest that fish predation is the major control of zooplankton size structure in brackish lakes, since fish density was related to a decrease in mean body size and density of zooplankton and this was reflected in a unimodal shaped biomass-sizespectrum with dominance of small sizes and low size diversity. Salinity might play a more indirect role by shaping zooplankton communities toward more salt-tolerant species. In a global-warming perspective, these results suggest that changes in the trophic structure of shallow lakes in temperate regions might be expected as a result of the warmer temperatures and the potentially associated increases in salinity. The decrease in the density of largebodied zooplankton might reduce the grazing on phytoplankton and thus the chances of maintaining the clear water state in these ecosystems

Environmental determinants of foraging strategies in Cory"s shearwaters Calonectris diomedea

Despite the advent of devices to track seabird movements, the extent to which productive areas and oceanic winds influence foraging strategies is still not fully understood. We investigated the main environmental determinants of foraging strategies in Cory"s shearwaters Calonectris diomedea by combining satellite-tracking information from 14 birds breeding on the Canary Islands with concurrent data on chlorophyll a (chl a) concentrations and oceanic winds. Additionally, we took blood samples at the end of each foraging trip and analysed carbon (C) and nitrogen (N) stable isotopes to examine the use of trophic resources. All birds showed commuting trips, concentrating foraging activity exclusively on the African continental shelf. Foraging locations showed a strong association with chl a concentrations, suggesting birds select foraging areas according to prey availability. In contrast with other breeding colonies where Cory"s shearwaters use a dual-foraging method, birds showed a unimodal strategy and did not show differences in C and N isotope signatures in plasma, confirming that close proximity to highly productive areas strongly influences foraging strategies. In addition, birds tracked during 2 consecutive trips foraged in the same area, suggesting that high resource availability promotes fidelity to feeding grounds also at coarse scales. Persistent northeast trade winds blew during the study period, and commuting trips followed a consistent clockwise movement with a southwest heading while the birds foraged along the continental shelf, suggesting that birds used tail winds to reduce their flying costs. Our results corroborate that oceanographic conditions in the vicinity of the breeding colony have a strong effect on foraging strategies of pelagic seabirds.