Individual diet variability shapes the architecture of Antarctic benthic food webs.

Antarctic biodiversity is affected by seasonal sea-ice dynamics driving basal resource availability. To (1) determine the role of intraspecific dietary variability in structuring benthic food webs sustaining Antarctic biodiversity, and (2) understand how food webs and the position of topologically central species vary with sea-ice cover, single benthic individuals' diets were studied by isotopic analysis before sea-ice breakup and afterwards. Isotopic trophospecies (or Isotopic Trophic Units) were investigated and food webs reconstructed using Bayesian Mixing Models. As nodes, these webs used either ITUs regardless of their taxonomic membership (ITU-webs) or ITUs assigned to species (population-webs). Both were compared to taxonomic-webs based on taxa and their mean isotopic values. Higher resource availability after sea-ice breakup led to simpler community structure, with lower connectance and linkage density. Intra-population diet variability and compartmentalisation were crucial in determining community structure, showing population-webs to be more complex, stable and robust to biodiversity loss than taxonomic-webs. The core web, representing the minimal community 'skeleton' that expands opportunistically while maintaining web stability with changing resource availability, was also identified. Central nodes included the sea-urchin Sterechinus neumayeri and the bivalve Adamussium colbecki, whose diet is described in unprecedented detail. The core web, compartmentalisation and topologically central nodes represent crucial factors underlying Antarctica's rich benthic food web persistence.

Effects of Sea-Ice Persistence on the Diet of Adélie Penguin (Pygoscelis adeliae) Chicks and the Trophic Differences between Chicks and Adults in the Ross Sea, Antarctica.

In Antarctica, prey availability for the mesopredator Adélie penguin, Pygoscelis adeliae, depends on sea-ice dynamics. By affecting cycles of sea-ice formation and melt, climate change could thus affect penguin diet and recruitment. In the light of climate change, this raises concerns about the fate of this dominant endemic species, which plays a key role in the Antarctic food web. However, few quantitative studies measuring the effects of sea-ice persistence on the diet of penguin chicks have yet been conducted. The purpose of this study was to fill this gap by comparing penguin diets across four penguin colonies in the Ross Sea and evaluating latitudinal and interannual variation linked to different sea-ice persistence. Diet was evaluated by analysing the δ13C and δ15N values of penguin guano, and sea-ice persistence by means of satellite images. Isotopic values indicate that penguins consumed more krill in colonies with longer sea-ice persistence. In these colonies, the δ13C values of chicks were lower and closer to the pelagic chain than those of adults, suggesting that the latter apparently catch prey inshore for self-feeding and offshore for their chicks. The results indicate that sea-ice persistence is among the principal factors that influence the spatiotemporal variability of the penguins' diet.

The Feeding Behaviour of Gall Midge Larvae and Its Implications for Biocontrol of the Giant Reed: Insights from Stable Isotope Analysis.

The gall midge Lasioptera donacis, whose larval stage interferes with the reed's leaf development, is a potential candidate agent for the biological control of Arundo donax. Reed infestation is always associated with the presence of a saprophytic fungus, Arthrinium arundinis, which is believed to provide food for the larvae. Larvae also interact with a parasitic nematode, Tripius gyraloura, which can be considered its natural enemy. To deepen our knowledge of the plant-fungus-insect trophic interactions and to understand the effects of the nematode on midge larval feeding behaviour, we applied stable isotope analysis, one of the most effective methods for investigating animal feeding preferences in various contexts. The results showed that on average the fungus accounted for 65% of the diet of the midge larvae, which however consumed the reed and the fungus in variable proportions depending on reed quality (expressed as the C:N ratio). No differences in feeding behaviour were observed between parasitised and non-parasitised midge larvae, indicating that nematodes have no effect in this regard. Due to its trophic habits, L. donacis could be an effective control agent of A. donax and these results need to be considered when implementing biological control measures.

Climate-related drivers of nutrient inputs and food web structure in shallow Arctic lake ecosystems.

In order to predict the effects of climate change on polar ecosystems, disentangling mechanisms of nutrient transfer in food webs is crucial. We investigated sources of nutrients in tundra lakes, tracing their transfer through the food web and relating the observed patterns to runoff, snow coverage, and the presence of migratory geese in lake catchments. C and N content (elemental and isotopic) of several food web components including Lepidurus arcticus (Notostraca, at the top of the lake food webs) in 18 shallow Arctic lakes was compared. Terrestrial productivity and geese abundance were key biotic factors that interacted with abiotic variables (snow coverage, lake and catchment size) in determining the amount and origin of nutrient inputs, affecting the trophic interactions among aquatic species, food chain length and nutrient flow in Arctic lake food webs. Decreasing snow coverage, increasing abundance and expansion of the geese's range are expected across the Arctic due to climate warming. By relating nutrient inputs and food web structure to snow coverage, vegetation and geese, this study contributes to our mechanistic understanding of the cascade effects of climate change in tundra ecosystems, and may help predict the response of lakes to changes in nutrient inputs at lower latitudes.

Increasing nutrient inputs over the last 500 years in an Italian low-impacted seagrass meadow.

Posidonia oceanica is a seagrass endemic to the Mediterranean and it has been widely used as a bioindicator. We studied the layers of a 500-year-old matte using a multiproxy approach (δ13C, δ15N, 14C and C and N concentrations in seagrass debris) in order to evaluate the potential of P. oceanica as a long-term environmental indicator of N pollution and CO2 emissions. From 1581 to 1800, accumulation rate was ca. 0.35 cm year-1, while in the last 100 years it has amounted to ca. 0.51 cm year-1. We observed increasing δ15N values with height in the vertical matte profile, indicating an increase in anthropogenic organic N inputs over time. In contrast, no clear trend in the δ13C values was observed. This study reconstructs the long-term impact of human activities on a seagrass meadow located off the Italian coast, yielding long-term background information that can help managers to implement efficient plans.

Evaluation of nitrogen and heavy metal pollution in southern Caspian Sea: Risk assessment and modeling approach.

Industrial, agricultural, and recreational activities dump several pollutants into the Caspian Sea, which is one of the main water bodies of Iran. Therefore, performing risk assessments would be required as part of the monitoring programs. Herein, non-cancer health risk assessment of the consumption of macroalgae was performed and the ecological risk assessment of metal pollution in sediments of the southern Caspian Sea was presented using sediment quality guidelines (SQGs), enrichment factors (EF), contamination factors (CF), contamination degrees (CD), pollution load indices (PLI), geo accumulation indices (Igeo), and potential ecological risk indices (RI). Next, machine-learning approaches were used to predict the stable isotope value of nitrogen in macroalgae, in which physicochemical information of water and heavy metal levels in macroalgae were used separately and together as predictor variables. Results indicated that simultaneous use of physicochemical properties of water and heavy metal levels resulted in the best prediction of isotopic nitrogen content.

Gaining insight into the assimilated diet of small bear populations by stable isotope analysis.

Apennine brown bears (Ursus arctos marsicanus) survive in an isolated and critically endangered population, and their food habits have been studied using traditional scat analysis. To complement current dietary knowledge, we applied Stable Isotope Analysis (SIA) to non-invasively collected bear hairs that had been individually recognized through multilocus genotyping. We analysed carbon (δ13C) and nitrogen (δ15N) stable isotopes of hair sections and bear key foods in a Bayesian mixing models framework to reconstruct the assimilated diet on a seasonal basis and to assess gender and management status effects. In total, we analysed 34 different seasonal bear key foods and 35 hair samples belonging to 27 different bears (16 females and 11 males) collected during a population survey in 2014. Most bears showed wide δ15N and δ13C ranges and individual differences in seasonal isotopic patterns. Vegetable matter (herbs, fleshy fruits and hard mast) represented the major component of the assimilated diet across the dietary seasons, whereas vegetable crops were rarely and C4 plants (i.e., corn) never consumed. We confirmed an overall low consumption of large mammals by Apennine bears consistently between sexes, with highest values in spring followed by early summer but null in the other seasons. We also confirmed that consumption of fleshy fruits peaked in late summer, when wild predominated over cultivated fleshy fruits, even though the latter tended to be consumed in higher proportion in autumn. Male bears had higher δ 15N values than females in spring and autumn. Our findings also hint at additional differences in the assimilated diet between sexes, with females likely consuming more herbs during spring, ants during early summer, and hard mast during fall compared to males. In addition, although effect sizes were small and credibility intervals overlapped considerably, management bears on average were 0.9‰ lower in δ 13C and 2.9‰ higher in δ 15N compared to non-management bears, with differences in isotopic values between the two bear categories peaking in autumn. While non-management bears consumed more herbs, wild fleshy fruits, and hard mast, management bears tended to consume higher proportions of cultivated fruits, ants, and large mammals, possibly including livestock. Although multi-year sampling and larger sample sizes are needed to support our findings, our application confirms that SIA can effectively integrate previous knowledge and be efficiently conducted using samples non-invasively collected during population surveys.

Trace elements and stable isotopes in penguin chicks and eggs: A baseline for monitoring the Ross Sea MPA and trophic transfer studies.

Multi-tissue trace elements (TEs), C, N concentrations and stable isotopes (δ13C, δ15N) of chick carcasses and eggs of Adélie and Emperor penguins were studied to i) provide reference data before the recent institution of the Ross Sea Marine Protected Area (Antarctica), and ii) provide conversion factors that allow estimating C, N, δ13C and δ15N in edible tissues from non-edible ones, thus improving the use of stable isotopes in contamination and trophic transfer studies. Higher concentrations of As, Cd, Cr, Cu, Hg, Mn and Pb were found in chick carcasses than in eggs, suggesting increasing contamination in recent decades and high toxicity risks for penguin consumers. Isotopic conversion factors highlighted small differences among body tissues and conspecifics. These values suggest that chick carcasses are reliable indicators of the energy pathways underlying the two penguin species, their trophic position in the food web and their exposure to TEs.

Stable isotope ratios (δ13C and δ15N) and heavy metal levels in macroalgae, sediment, and benthos from the northern parts of Persian Gulf and the Gulf of Oman.

In this investigation, δ13C and δ15N isotope ratios and heavy metal levels were assessed in macroalgae, sediment samples, and benthic species from northern parts of the Persian Gulf and the Gulf of Oman. The highest δ15N values in algal samples (11±0.42), indicative of anthropogenic organic N inputs, were detected in the Sadaf region, whereas the lowest values (3.17 ±â€¯0.12), indicative of anthropogenic inorganic inputs, were detected in the Parvaz region. In addition to sediment quality guidelines (SQGs), contamination factors (CF), enrichment factors (EF), contamination degree (CD), pollution load index (PLI), geo accumulation index (Igeo), and potential ecological risk index (RI) were employed to assess the anthropogenic influence on sediment quality and to describe the sensitivity of the biota to toxic heavy metals. The obtained results demonstrate that the analyzed elements (Mn, Cr, Ni, Pb, Zn, Cu, Co, and V) had no ecological risk in the sampling area.

Nitrogen and metal pollution in the southern Caspian Sea: a multiple approach to bioassessment.

The Caspian Sea hosts areas of high ecological value as well as industrial, leisure, and agricultural activities that dump into the water body different kinds of pollutants. In this complex context, a proper description of the origin and potential sources of pollution is necessary to address management and mitigation actions aimed at preserving the quality of the water resource and the integrity of the ecosystems. Here, we aimed at detecting sources of both nitrogen inputs, by N stable isotope analysis of macroalgae, and metals in macroalgae and sediments in two highly anthropized coastal stretches at the Iranian side of the Caspian Sea. Sampling was done near the mouth of rivers and canals draining agricultural and urbanized areas. In the westernmost waters, facing a port city, low macroalgal δ15N signatures indicated industrial fertilizers as the principal source of pollution. By contrast, in the central coastal waters, facing touristic areas, the high macroalgal δ15N indicated N inputs from wastewaters. Here the lowest dissolved oxygen concentrations in waters were associated with excess dissolved inorganic nitrogen. Metal concentrations varied largely in the study areas and were lower in macroalgae than in sediments. Localized peaks of Pb and Zn in sediments were observed in the central coastal sites as probable byproducts of mining activity transported downstream. By contrast, Cr and Ni concentrations were high in all sampling sites, thus potentially representing hazardous elements for marine biota. Overall, macroalgal δ15N coupled with metal analysis in macroalgae and sediments was useful for identifying the main sources of pollution in these highly anthropized coastal areas. This double approach in comprehensive monitoring programs could thus effectively inform stakeholders on major environmental threats, allowing targeted management measures.

Species richness and vulnerability to disturbance propagation in real food webs.

A central issue in ecology is understanding how complex and biodiverse food webs persist in the face of disturbance, and which structural properties affect disturbance propagation among species. However, our comprehension of assemblage mechanisms and disturbance propagation in food webs is limited by the multitude of stressors affecting ecosystems, impairing ecosystem management. By analysing directional food web components connecting species along food chains, we show that increasing species richness and constant feeding linkage density promote the establishment of predictable food web structures, in which the proportion of species co-present in one or more food chains is lower than what would be expected by chance. This reduces the intrinsic vulnerability of real food webs to disturbance propagation in comparison to random webs, and suggests that biodiversity conservation efforts should also increase the potential of ecological communities to buffer top-down and bottom-up disturbance in ecosystems. The food web patterns observed here have not been noticed before, and could also be explored in non-natural networks.

Antarctic food web architecture under varying dynamics of sea ice cover.

In the Ross Sea, biodiversity organisation is strongly influenced by sea-ice cover, which is characterised by marked spatio-temporal variations. Expected changes in seasonal sea-ice dynamics will be reflected in food web architecture, providing a unique opportunity to study effects of climate change. Based on individual stable isotope analyses and the high taxonomic resolution of sampled specimens, we described benthic food webs in contrasting conditions of seasonal sea-ice persistence (early vs. late sea-ice break up) in medium-depth waters in Terra Nova Bay (Ross Sea). The architecture of biodiversity was reshaped by the pulsed input of sympagic food sources following sea-ice break up, with food web simplification, decreased intraguild predation, potential disturbance propagation and increased vulnerability to biodiversity loss. Following our approach, it was possible to describe in unprecedented detail the complex structure of biodiverse communities, emphasising the role of sympagic inputs, regulated by sea-ice dynamics, in structuring Antarctic medium-depth benthic food webs.

Horizontal and vertical food web structure drives trace element trophic transfer in Terra Nova Bay, Antarctica.

Despite a vast amount of literature has focused on trace element (TE) contamination in Antarctica during the last decades, the assessment of the main pathways driving TE transfer to the biota is still an overlooked issue. This limits the ability to predict how variations in sea-ice dynamics and productivity due to climate change will affect TE allocation in the food web. Here, food web structure of Tethys Bay (Terra Nova Bay, Ross Sea, Antarctica) was first characterised by analysing carbon and nitrogen stable isotopes (δ13C, δ15N) in organic matter sources (sediment and planktonic, benthic and sympagic primary producers) and consumers (zooplankton, benthic invertebrates, fish and birds). Diet and trophic position were also characterised using Bayesian mixing models. Then, relationships between stable isotopes, diet and TEs (Cd, Cr, Co, Cu, Hg, Ni, Pb and V) were assessed in order to evaluate if and how horizontal (organic matter pathways) and vertical (trophic position) food web features influence TE transfer to the biota. Regressions between log[TE] and δ13C revealed that the sympagic pathway drives accumulation of V in primary consumers and Cd and Hg in secondary consumers, and that a coupled benthic/pelagic pathway drives Pb transfer to all consumers. Regressions between log[TE] and δ15N showed that only Hg biomagnifies across trophic levels, while all the others TEs showed a biodilution pattern, consistent with patterns observed in temperate food webs. Although the Cd behavior needs further investigations, the present findings provide new insights about the role of basal sources in the transfer of TEs in polar systems. This is especially important nowadays in light of the forecasted trophic changes potentially resulting from climate change-induced modification of sea-ice dynamics.

Time- and depth-wise trophic niche shifts in Antarctic benthos.

Climate change is expected to affect resource-consumer interactions underlying stability in polar food webs. Polar benthic organisms have adapted to the marked seasonality characterising their habitats by concentrating foraging and reproductive activity in summer months, when inputs from sympagic and pelagic producers increase. While this enables the persistence of biodiverse food webs, the mechanisms underlying changes in resource use and nutrient transfer are poorly understood. Thus, our understanding of how temporal and spatial variations in the supply of resources may affect food web structure and functioning is limited. By means of C and N isotopic analyses of two key Antarctic benthic consumers (Adamussium colbecki, Bivalvia, and Sterechinus neumayeri, Echinoidea) and Bayesian mixing models, we describe changes in trophic niche and nutrient transfer across trophic levels associated with the long- and short-term diet and body size of specimens sampled in midsummer in both shallow and deep waters. Samplings occurred soon after the sea-ice broke up at Tethys Bay, an area characterised by extreme seasonality in sea-ice coverage and productivity in the Ross Sea. In the long term, the trophic niche was broader and variation between specimens was greater, with intermediate-size specimens generally consuming a higher number of resources than small and large specimens. The coupling of energy channels in the food web was consequently more direct than in the short term. Sediment and benthic algae were more frequently consumed in the long term, before the sea-ice broke up, while consumers specialised on sympagic algae and plankton in the short term. Regardless of the time scale, sympagic algae were more frequently consumed in shallow waters, while plankton was more frequently consumed in deep waters. Our results suggest a strong temporal relationship between resource availability and the trophic niche of benthic consumers in Antarctica. Potential climate-driven changes in the timing and quality of nutrient inputs may have profound implications for the structure of polar food webs and the persistence of their constituent species, which have adapted their trophic niches to a highly predictable schedule of resource inputs.

Space-time monitoring of coastal pollution in the Gulf of Gaeta, Italy, using δ15N values of Ulva lactuca, landscape hydromorphology, and Bayesian Kriging modelling.

We investigated the space-time dynamics of N pollution in a Mediterranean gulf (Gulf of Gaeta) by means of δ15N variation in seaweed fronds (Ulva lactuca) previously collected from an unpolluted habitat. We used a comprehensive deployment grid that enabled the generation of isotopic seascapes (isoseascapes) describing the topography of N pollution in coastal waters and identifying N input hotspots and their pathways of dispersion at sea. The δ15N values of U. lactuca increased during 48h of exposure to the gulf waters, indicating anthropogenic N inputs from wastewater-derived sources. Comparison of the isoseascapes between two years differing in terms of rainfall identified coastal and offshore areas that were vulnerable to freshwater-transported nutrients, consistent with terrestrial hydromorphology and sea surface-water circulation. Isoseacapes were robust enough to reduce deployment effort, representing a powerful tool for monitoring and management strategies and useful for Environmental Protection Agencies, the main target audience of applied ecological research.

Effect of habitat degradation on competition, carrying capacity, and species assemblage stability.

Changes in species' trophic niches due to habitat degradation can affect intra- and interspecific competition, with implications for biodiversity persistence. Difficulties of measuring species' interactions in the field limit our comprehension of competition outcomes along disturbance gradients. Thus, information on how habitat degradation can destabilize food webs is scarce, hindering predictions regarding responses of multispecies systems to environmental changes. Seagrass ecosystems are undergoing degradation. We address effects of Posidonia oceanica coverage reduction on the trophic organization of a macroinvertebrate community in the Tyrrhenian Sea (Italy), hypothesizing increased trophic generalism, niche overlap among species and thus competition and decreased community stability due to degraded conditions. Census data, isotopic analysis, and Bayesian mixing models were used to quantify the trophic niches of three abundant invertebrate species, and intra- and interspecific isotopic and resource-use similarity across locations differing in seagrass coverage. This allowed the computation of (1) competition strength, with respect to each other and remaining less abundant species and (2) habitat carrying capacity. To explore effects of the spatial scale on the interactions, we considered both individual locations and the entire study area ("'meadow scale"). We observed that community stability and habitat carrying capacity decreased as P. oceanica coverage declined, whereas niche width, similarity of resource use and interspecific competition strength between species increased. Competition was stronger, and stability lower, at the meadow scale than at the location scale. Indirect effects of competition and the spatial compartmentalization of species interactions increased stability. Results emphasized the importance of trophic niche modifications for understanding effects of habitat loss on biodiversity persistence. Calculation of competition coefficients based on isotopic distances is a promising tool for describing competitive interactions in real communities, potentially extendible to any subset of ecological niche axes for which specimens' positions and pairwise distances can be obtained.

The effects of nitrogen pollutants on the isotopic signal (δ15N) of Ulva lactuca: Microcosm experiments.

Effects of two chemical forms of Nitrogen (NH4+ and NO3-) on δ15N in Ulva lactuca were analysed separately and in mixture at two concentrations. We assessed whether the δ15N values of U. lactuca discriminate between Nitrogen from synthetic fertilisers (inorganic) and from fresh cow manure (organic), and the isotopic ability of the macroalga to reflect Nitrogen concentrations. Isotopic signature and N content of the macroalga reflected different nitrogenous sources and their concentrations after 48h. The inorganic Nitrogen source (NH4NO3) altered the isotopic values of the macroalgae more than Nitrogen from fresh cow manure (NO3-). δ15N values observed in the mixed solution did not differ from those displayed in NH4NO3 treatment alone. We conclude that stable isotope analysis of U. lactuca collected in an unpolluted site and experimentally submerged in sites suspected of being affected by disturbance is a useful tool for rapid monitoring of anthropogenic discharges of Nitrogen pollutants.

Stable isotope variations in benthic primary producers along the Bosphorus (Turkey): A preliminary study.

The Bosphorus Strait is a dynamic and complex system. Recent evidences showed nitrogen and heavy metal concentrations to follow opposite patterns across the Strait, suggesting a complex spatial organisation of the anthropogenic disturbance in this system. Here, we provide isotopic information on the origin and transportation of dissolved nitrogen along the Bosphorus. C and N isotopic and elemental analyses were performed on specimens of Ulva lactuca and associated epiphytes sampled in five locations across the Strait. Variations in C and N isotopic signatures were observed in U. lactuca, pointing to a decrease in the availability of anthropogenic organic dissolved nitrogen along a north-south direction. Conversely, epiphytes did not show isotopic or elemental patterns across the Strait. These results suggest that preliminary stable isotope surveys in extended costal systems basing on U. lactuca can represent a valuable tool to focus meaningful targets and hypotheses for pollution studies in the Mediterranean region.

δ(15)N variation in Ulva lactuca as a proxy for anthropogenic nitrogen inputs in coastal areas of Gulf of Gaeta (Mediterranean Sea).

We tested the capacity of Ulva lactuca to mark N sources across large marine areas by measuring variation in its δ(15)N at several sites in the Gulf of Gaeta. Comparisons were made with the macroalga Cystoseira amentacea. Variation of δ(15)N values was assessed also in the coastal waters off the Circeo Natural Park, where U. lactuca and C. amentacea were harvested, as these waters are barely influenced by human activities and were used as reference site. A small fragment from each frond was preserved before deployment in order to characterize the initial isotopic values. After 48 h of submersion, U. lactuca was more responsive than C. amentacea to environmental variation and δ(15)N enrichment in the Gulf of Gaeta was observed. The spatial distribution of δ(15)N enrichment indicated that different macro-areas in the Gulf were affected by N inputs from different origins. Comparison of the δ(15)N values of fragments taken from the same transplanted frond avoided bias arising from natural isotopic variability.

Predators and resources influence phosphorus transfer along an invertebrate food web through changes in prey behaviour.

Predators play a fundamental role in prey trophic behaviour, with indirect consequences for species coexistence and ecosystem functioning. Resource quality and availability also influence prey trophic behaviour, with potential effects on predator-prey dynamics. Although many studies have addressed these topics, little attention has been paid to the combined effects of predators and resources on prey species coexistence and nutrient transfer along food chains, especially in detritus-based systems. To determine the influence of predators and resource quality on the movement and P uptake of detritivores, we carried out a field experiment on the River Kelvin (Scotland) using (32)P to test the hypothesis of reduced prey vagility among resource patches as a strategy to avoid predation. Thirty leaf sacks containing alder leaves and two detritivore prey populations (Asellus aquaticus and Lymnaea peregra) were placed in cages, half of them with two predator species (Dendrocoelum lacteum and Erpobdella octoculata) and the other half without predators. Five alder leaf bags, each individually inoculated with a different fungus strain to simulate a patchy habitat, were placed inside each leaf sack. One bag in each sack was labelled with (32)P, in order to assess the proportion of detritivores using it as food and thus their movement among the five resource patches. Three replicates for each labelled fungus and each predation treatment (i.e. with and without predators) were left on the riverbed for 7 days. The presence of predators had negligible effects on the number of detritivores in the leaf bags, but it did reduce the proportion of (32)P-labelled detritivores and their P uptake. The most strongly affected species was A. aquaticus, whose vagility, trophic overlap with L. peregra and P uptake were all reduced. The results confirm the importance of bottom-up and top-down forces acting simultaneously to regulate nutrient transfer along food chains in patchy habitats.