Size-dependent foraging niches of European Perch <Emphasis Type="Italic">Perca fluviatilis</Emphasis> (Linnaeus, 1758) and North American Yellow Perch <Emphasis Type="Italic">Perca flavescens</Emphasis> (Mitchill, 1814)

Body size of consumer species is a fundamental trait that influences the trophic ecology of individuals and their contribution to the functioning of freshwater ecosystems. However, the relationship between body size and trophic ecology can be highly variable both within and between closely-related and similarly-sized species. In this study we compared the intra- and interspecific relationship between body size and trophic position for North American Yellow Perch Perca flavescens and European Perch Perca fluviatilis, which share similarities in morphology, life history traits and trophic requirements. We used stable isotope ratios (δ¹⁵N and δ¹³C) to characterize differences in size-dependency of trophic position and to trace consumer foraging history of Yellow Perch in lakes in the Northwestern United States and European Perch in lakes in Germany. The trophic position and stable isotope ratios of Yellow Perch and European Perch steadily increased with total body length, but European Perch were consistently feeding at higher trophic positions than Yellow Perch at a given length. European Perch occupied considerably higher trophic positions (mean trophic position = 3.9) than Yellow Perch (mean trophic position = 2.8). Large European Perch were increasingly piscivorous, whereas large Yellow Perch were more opportunistic and omnivorous predators of invertebrate prey. Overall, the trophic position among individual Yellow Perch varied more strongly than in European Perch. We conclude that both species similarly increase in trophic position with size, but the specific size-dependency of both trophic position and resource use varies with taxonomy and local ecological conditions. Thus, body size as a sole measure of trophic position should be considered cautiously when generalizing across populations and species.     

Evidence of two subaggregations of humpback whales on the Kodiak,Alaska, feeding ground revealed from stable isotope analysis

Knowledge of humpback whale (Megaptera novaeangliae) foraging on feeding grounds is becoming increasingly important as the growing North Pacific population recovers from commercial whaling and consumes more prey, including economically important fishes. We explored spatial and temporal (interannual, within‐season) variability in summer foraging by humpback whales along the eastern side of the Kodiak Archipelago as described by stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope ratios of humpback whale skin (n = 118; 2004–2013). The trophic level (TL) of individual whales was calculated using basal food web δ¹⁵N values collected within the study area. We found evidence for the existence of two subaggregations of humpback whales (“North,” “South”) on the feeding ground that fed at different TLs throughout the study period. Linear mixed models suggest that within an average year, Kodiak humpback whales forage at a consistent TL during the feeding season. TL estimates support mixed consumption of fish and zooplankton species in the “North” (mean ± SE; 3.3 ± 0.1) and predominant foraging on zooplankton in the “South” (3.0 ± 0.1). This trend appears to reflect spatial differences in prey availability, and thus, our results suggest North Pacific humpback whales may segregate on feeding aggregations and target discrete prey species.     

Consumer‐resource stoichiometry as a predictor of trophic discrimination (Δ13C, Δ15N) in aquatic invertebrates

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The trophic structure of bark-living oribatid mite communities analysed with stable isotopes (15N, 13C) indicates strong niche differentiation

The aim of the present study was to identify food sources of bark-living oribatid mites to investigate if trophic niche differentiation contributes to the diversity of bark living Oribatida. We measured the natural variation in stable isotope ratios (¹⁵N/¹⁴N, ¹³C/¹²C) in oribatid mites from the bark of oak (Quercus robur), beech (Fagus sylvatica), spruce (Picea abies) and pine (Pinus sylvestris) trees and their potential food sources, i.e., the covering vegetation of the bark (bryophytes, lichens, algae, fungi). As a baseline for calibration the stable isotope signatures of the bark of the four tree species were measured and set to zero. Oribatid mite stable isotope ratios spanned over a range of about 13 δ units for ¹⁵N and about 7 δ units for ¹³C suggesting that they span over about three trophic levels. Different stable isotope signatures indicate that bark living oribatid mites feed on different food sources, i.e., occupy distinct trophic niches. After calibration stable isotope signatures of respective oribatid mite species of the four tree species were similar indicating close association of oribatid mites with the corticolous cover as food source. Overall, the results support the hypothesis that trophic niche differentiation of bark living oribatid mites contributes to the high diversity of the group.     

Carbon and nitrogen isotope fractionation of amino acids in an avian marine predator,the gentoo penguin (Pygoscelis papua)

Compound‐specific stable isotope analysis (CSIA) of amino acids (AA) has rapidly become a powerful tool in studies of food web architecture, resource use, and biogeochemical cycling. However, applications to avian ecology have been limited because no controlled studies have examined the patterns in AA isotope fractionation in birds. We conducted a controlled CSIA feeding experiment on an avian species, the gentoo penguin (Pygoscelis papua), to examine patterns in individual AA carbon and nitrogen stable isotope fractionation between diet (D) and consumer (C) (Δ¹³C_C‐D and Δ¹⁵N_C‐D, respectively). We found that essential AA δ¹³C values and source AA δ¹⁵N values in feathers showed minimal trophic fractionation between diet and consumer, providing independent but complimentary archival proxies for primary producers and nitrogen sources respectively, at the base of food webs supporting penguins. Variations in nonessential AA Δ¹³C_C‐D values reflected differences in macromolecule sources used for biosynthesis (e.g., protein vs. lipids) and provided a metric to assess resource utilization. The avian‐specific nitrogen trophic discrimination factor (TDF_Glu‐Phe = 3.5 ± 0.4‰) that we calculated from the difference in trophic fractionation (Δ¹⁵N_C‐D) of glutamic acid and phenylalanine was significantly lower than the conventional literature value of 7.6‰. Trophic positions of five species of wild penguins calculated using a multi‐TDF_Glu‐Phe equation with the avian‐specific TDF_Glu‐Phe value from our experiment provided estimates that were more ecologically realistic than estimates using a single TDF_Glu‐Phe of 7.6‰ from the previous literature. Our results provide a quantitative, mechanistic framework for the use of CSIA in nonlethal, archival feathers to study the movement and foraging ecology of avian consumers.     

Evolutionary convergence in Neotropical cichlids and Nearctic centrarchids: evidence from morphology,diet, and stable isotope analysis

Despite divergent evolutionary histories, Neotropical cichlids (Cichlidae) and Nearctic sunfishes (Centrarchidae) appear to have similar functional morphotypes and occupy similar ecological niches. We applied an integrative approach analyzing morphological traits, stomach contents, and stable isotope ratios (δ¹³C, δ¹⁵N) to investigate whether local assemblages of cichlids (Venezuela, Peru) and centrarchids (Texas) reveal one‐to‐one patterns of morphological and ecological convergence. Multivariate ordinations performed on diet and morphology datasets identified a broad overlap between cichlid and centrarchid assemblages. The functional morphology of the two groups has diversified in a convergent manner within the confines of ram‐suction modes of prey ingestion. Both groups had the same set of ecomorph types that corresponded to the same trophic niches, including substrate‐sifting invertivores, epibenthic invertebrate gleaners, and piscivores; the one exception was a molluscivorous sunfish, comprising a niche that was not represented in the two cichlid assemblages. Estimates of trophic positions based on stable isotope analysis revealed convergent vertical trophic structure; with few exceptions, fishes with similar morphologies had similar trophic positions. Large‐bodied piscivores had highest trophic positions, whereas small and medium‐bodied generalists and invertivores had low to intermediate trophic positions. Consistent patterns of ecomorphological convergence in these two perciform groups provide strong evidence for adaptation involving constrains in functional morphology associated with feeding. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 146–164.     

Intra‐lake stable isotope ratio variation in selected fish species and their possible carbon sources in Lake Kyoga (Uganda): implications for aquatic food web studies

The stable isotopes of nitrogen (δ¹⁵N) and carbon (δ¹³C) provide powerful tools for quantifying trophic relationships and carbon flow to consumers in food webs; however, the isotopic signatures of organisms vary within a lake. Assessment of carbon and nitrogen isotopic signatures in a suite of plants, invertebrates, and fishes in Lake Kyoga, indicated significant variation between two sites for δ¹³C (paired t = 6.305; df = 14, P < 0.001 and δ¹⁵N paired t = 1.292; df = 14; P < 0.05). The fish fauna in Bukungu was generally more ¹³C enriched (mean δ¹³C = –16.37 ± 1.64‰) than in Iyingo (mean δ¹³C = –20.80 ± 2.41‰) but more δ¹⁵N depleted (mean δ¹⁵N = 5.57 ± 0.71‰) than in Iyingo (mean δ¹⁵N = 6.92 ± 0.83‰). The simultaneous shifts in phytoplankton and consumer signatures confirmed phytoplankton as the major source of carbon for the food chain leading to fish. Limited sampling coverage within lakes may affect lake wide stable isotope signatures, and the same error is transferred into trophic position estimation. Consideration of potential intra‐lake spatial variability in isotope ratios and size is essential in evaluating the spatial and trophic structure of fish assemblages.     

Fine‐scale spatial differences in humpback whale diet composition near Kodiak,Alaska

On the North Pacific feeding grounds, humpback whales (Megaptera novaeangliae) are recovering from commercial whaling at a rapid rate (6.8%). The potential effect that this recovery will have on trophic dynamics involving these predators is currently unknown. To better elucidate complex trophic dynamics, variability in diet composition of apex predators on their respective feeding grounds needs to be understood. Thus, we explored the diet composition of two defined subaggregations of humpback whales of the Kodiak Archipelago population (“North,” “South”) using stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope ratios of humpback whale skin and regional prey samples in Bayesian dietary mixing models. Humpback whales in the “North” region consumed proportionally more fish, dominated by capelin (Mallotus villosus), whereas, whales in the “South” region consumed predominantly krill. The difference in diet composition appears to reflect regional differences in prey availability. Thus, regional variability in diet composition by humpback whales may have disproportionate impacts on prey resources of sympatric predators depending on available prey biomass. As a result, we suggest fine‐scale studies of apex predator diets are needed to better model trophic dynamics with accuracy.     

Ecological implications of unprecedented warm water anomalies on interannual prey preferences and foraging areas of Guadalupe fur seals

The Guadalupe fur seal (GFS) currently is recovering from near extinction and prey availability is an important factor. Scat and stable isotope analyses (SIA) were used to assess GFS foraging on Guadalupe Island during 2013–2016. This period was characterized by normal (2013) ocean temperatures followed by warm conditions (2014–2016). Scat samples (~40/year) were collected for prey identification. Additionally, lanugo samples from 1-month-old pups (50/year) were processed for SIA (δ¹³C/δ¹⁵N). Interannual prey preferences were identified (ANOSIM, p < .05), including significant differences between 2013 and the anomalous years. The most important prey species among these years was the jumbo squid; followed by the neon flying squid (2014–2016). The GFS diet was more specialized in 2013–2015 and more generalized in 2016. Prey from higher trophic levels (determined by scat analysis) were consumed during 2014–2016, but with significantly lower δ¹³C/δ¹⁵N values than in 2013 (ANOVA, p < .05 for both isotope ratios). This pattern may indicate more northerly or offshore foraging areas, with the presence of oceanic prey (neon flying squid). The widest isotopic niche was observed in 2015 (2.2‰²), reflecting a broader foraging area. Our findings are an important step toward better understanding the impacts of climate change on the recovery of GFS.     

Detecting alternate foraging ecotypes in Australian sea lion (Neophoca cinerea) colonies using stable isotope analysis

Stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotopes are used frequently to describe the trophic ecology of top marine mammal predators. Australian sea lions (Neophoca cinerea) are one of the world's rarest otariid seals and exhibit the highest levels of natal site philopatry of any seal. We report the development of a screening technique to identify different foraging ecotypes and assess their relative frequencies in Australian sea lion breeding colonies using stable isotope ratios in pups. Geospatial and dive data from 15 adult females at three breeding colonies revealed alternate foraging strategies (inshore and offshore foraging) that were reflected in significant changes in δ¹³C and δ¹⁵N. Isotope fractionation from mother to pup was validated using paired whisker and blood serum samples with no significant difference between δ¹³C and δ¹⁵N enrichment of +1.27‰ (whiskers) and +1.92‰ (blood serum) from mothers to pups. Isotope ratios from whisker samples representing over 50% of pups born at three colonies revealed significant intercolony differences in maternal foraging ecotype frequencies. These results are unique in that ecological partitioning over such a small spatial scale has not been described in any other otariid species.     

Stable isotope analysis indicates trophic differences among forest floor carabids in Japan

Differences in trophic niches among carabid beetles (Coleoptera: Carabidae) co‐occurring on the forest floors of warm temperate forests in central Japan were studied using carbon (δ¹³C) and nitrogen (δ¹⁵N) stable isotope analyses. Different carabid species showed similar δ¹⁵N values, which were higher than those of their possible invertebrate prey (herbivores and detritivores) collected from the litter layer, indicating that these species were consumers in the same trophic level. In contrast, δ¹³C values differed among carabid species, indicating interspecific differences in prey animals. The variation in the δ¹³C value was larger in summer than in autumn. In summer, δ¹³C values indicated that some carabids depended highly on either grazing (low δ¹³C values) or detrital sources (high δ¹³C values) within the food chain [Chlaenius posticalis Motschulsky and Haplochlaenius costiger (Chaudoir), respectively], although other species with intermediate δ¹³C values likely depended on both. The latter group of species comprised mostly two dominant genera (Carabus and Synuchus). Although congeners might have similar feeding habits, the stable isotope ratios indicated trophic niche differences between adults of different species and between adults and larvae of the same genus.     

Terrestrial contributions to Afrotropical aquatic food webs: The Congo River case

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Insights into the spatiotemporal component of feeding ecology: an isotopic approach for conservation management sciences

Aim Bringing new approaches to trace spatiotemporal variations in animals’ feeding ecology, which is fundamental for wildlife management and conservation since the accessibility of animals to trophic resources plays a key role in the dynamics of populations and metapopulations. Location Western Mediterranean coast. Methods The analysis of naturally occurring stable isotopes constitutes an exceptional approach to assess variations in the trophic ecology of species within the spatiotemporal dimensions. Here, we examined the spatiotemporal heterogeneity in resource exploitation of a nuisance and overpopulated gull species with a great feeding plasticity, the Yellow‐legged gull Larus michahellis, by measuring the stable isotope ratios of carbon (¹³C/¹²C, δ¹³C), nitrogen (¹⁵N/¹⁴N, δ¹⁵N) and sulphur (³⁴S/³³S, δ³⁴S) in different wing feathers. Results Tracing isotopic changes among feathers in relation to moulting sequence, we showed that isotopic analyses on first primary feathers were good indicators of breeding trophic ecology, while others, such as eighth secondary feathers, reflected the feeding behaviour during the non‐breeding period. This knowledge was then used to report on estimations of seasonal feeding patterns throughout the year for seven gull populations along the Western Mediterranean coast. Main conclusions The high diversity in the exploitation patterns of the foraging habitats found in the study area both at spatial and at temporal scale indicated the opportunistic feeding behaviour of Yellow‐legged gulls. Proposed isotopic approach has revealed as a useful tool to evaluate spatiotemporal variations in the feeding ecology of populations which may become clue for dealing with the management of wild species, not only nuisance populations but also endangered species. We finally pointed out the relevance of the isotopic variability among individuals to infer diet diversity and food availability of a given population, thereby allowing demographic forecasts when trophic resources vary in abundance.     

Effects of Nile perch, Lates niloticus, on functional and specific fish diversity in Uganda's Lake Kyoga system

The introduction of Nile perch, Lates niloticus, to Lake Victoria, East Africa, interacted with eutrophication to cause a reorganization of the lake's food web and the extirpation of many endemic fishes. The Lake Kyoga satellite system lies downstream from Lake Victoria. It encompasses species‐rich lakes where Nile perch are absent or very rare, and low diversity lakes where L. niloticus is abundant. In 1999 we surveyed seven lakes in the Kyoga system using experimental monofilament gill nets (1/4–1 inches variable mesh). At Boston University we assessed δ¹⁵N signatures of epaxial muscle from subsamples of the catch (n = 361). These signatures are often highly correlated with the near‐term mean realized trophic position of an individual organism. A neural network analysis of fish length, species name, trophic level, and lake of origin fish explained 94% of the sample variance in δ¹⁵N. We analysed statistical patterns in these signatures at a number of spatial scales. The relationship between trophic level and δ¹⁵N varied greatly among lakes. Higher diversity perch‐free lakes had greater variance in δ¹⁵N values and fish lengths than lower diversity Nile perch lakes, suggesting an important relationship between species diversity and functional diversity. Against expectations, lake size was negatively correlated with δ¹⁵N. Patterns in stable isotope signatures indicated that Nile perch lakes have shorter food chains than perch‐free lakes. The results throw up two management problems for the Kyoga system. Impacted lakes need to be studied to understand and ameliorate the community‐level effects of Nile perch introduction, whereas the species‐rich nonperch lakes, which harbour a large proportion of the remaining diversity of regionally endemic taxa, are in need of conservation planning.     

Seasonal changes in the stable isotope values of lake-dwelling chironomid larvae in relation to feeding and life cycle variability

1. We studied seasonal changes in the carbon and nitrogen stable isotope ratios of larval Chironomus anthracinus and C. plumosus from the profundal sediments of four contrasting lakes. 2. Pronounced seasonal changes in both δ¹³C and δ¹⁵N values were evident in chironomid larvae of both species from two summer‐stratified, eutrophic lakes: Esthwaite Water and Wyresdale Park. Changes were most marked in the larvae of C. plumosus and in larvae from greater depths. In contrast, neither C. anthracinus in summer‐stratified but mesotrophic Schöhsee, nor C. plumosus in polymictic Großer Binnensee, showed marked seasonality in larval stable isotope ratios. 3. The particularly strong ¹³C‐depletion of larvae from the stratified, eutrophic lakes is attributed to a significant contribution of methane‐derived carbon to their diets. Feeding by larvae on isotopically light methanotrophic bacteria appears to occur mainly when autumn overturn of the water column restores oxygenated conditions to the sediment surface. At this time both δ¹³C and δ¹⁵N values of larvae decreased sharply. 4. Changes in the mean stable isotope ratio of the larval populations can also occur when larger, more isotopically light, larvae pupate and emigrate from the population to hatch as imagos. This effect can induce seasonal changes in larval isotope values even in lakes in which there is no evidence of a significant involvement of methane‐derived carbon in their diets. Variations in emergence patterns between species and between lakes may generate differences in the seasonal pattern of change in stable isotope ratios in larval populations. 5. Our results emphasise the importance of adequate seasonal sampling if stable isotope ratios are to be used as biomarkers to study the role of key groups, such as chironomid larvae, in the trophic structure of lakes.     

Fish determine macroinvertebrate food webs and assemblage structure in Greenland subarctic streams

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Structure of tropical river food webs revealed by stable isotope ratios

Fish assemblages in tropical river food webs are characterized by high taxonomic diversity, diverse foraging modes, omnivory, and an abundance of detritivores. Feeding links are complex and modified by hydrologic seasonality and system productivity. These properties make it difficult to generalize about feeding relationships and to identify dominant linkages of energy flow. We analyzed the stable carbon and nitrogen isotope ratios of 276 fishes and other food web components living in four Venezuelan rivers that differed in basal food resources to determine 1) whether fish trophic guilds integrated food resources in a predictable fashion, thereby providing similar trophic resolution as individual species, 2) whether food chain length differed with system productivity, and 3) how omnivory and detritivory influenced trophic structure within these food webs. Fishes were grouped into four trophic guilds (herbivores, detritivores/algivores, omnivores, piscivores) based on literature reports and external morphological characteristics. Results of discriminant function analyses showed that isotope data were effective at reclassifying individual fish into their pre-identified trophic category. Nutrient-poor, black-water rivers showed greater compartmentalization in isotope values than more productive rivers, leading to greater reclassification success. In three out of four food webs, omnivores were more often misclassified than other trophic groups, reflecting the diverse food sources they assimilated. When fish δ¹⁵N values were used to estimate species position in the trophic hierarchy, top piscivores in nutrient-poor rivers had higher trophic positions than those in more productive rivers. This was in contrast to our expectation that productive systems would promote longer food chains. Although isotope ratios could not resolve species-level feeding pathways, they did reveal how top consumers integrate isotopic variability occurring lower in the food web. Top piscivores, regardless of species, had carbon and nitrogen profiles less variable than other trophic groups.     

δ15N of zooplankton species in subarctic lakes in northern Sweden: effects of diet and trophic fractionation

1. To assess the use of stable nitrogen isotopes (δ¹⁵N) for reconstructing trophic relationships in planktonic food webs, crustacean zooplankton species and pelagic dissolved and particulate matter were analysed in 14 subarctic lakes in northern Sweden. The lakes are situated along an altitudinal gradient and show a substantial variation in nutrient content and energy mobilization by bacterioplankton and phytoplankton. 2. The δ¹⁵N of dissolved and particulate matter was comparatively low, suggesting efficient N recycling and low losses of depleted N from the pelagic zone of these unproductive lakes. 3. Copepods had a systematically higher δ¹⁵N than cladocerans, with an average difference of 3.1–4.9‰ within lakes, implying different trophic positions of the two groups. Comparisons of nitrogen pools and energy fluxes suggest that the low cladoceran δ¹⁵N was a result of feeding on bacteria. 4. The difference in δ¹⁵N between copepods and cladocerans declined with decreasing bacterioplankton production among lakes, due either to increasing trophic isotope fractionation or decreasing relative importance of bacteria in the diet of cladocerans.     

Aquatic food‐web structure along a salinized dryland river

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Are you what you eat? Effects of trophic discrimination factors on estimates of food assimilation and trophic position with a new estimation method

A key factor for estimates of assimilation of resources and trophic position based on stable isotope data is the trophic discrimination factor (TDF). TDFs are assumed based on literature reviews, but may vary depending on a variety of factors, including the type of diet. We analyzed effects of alternative TDFs on estimates of assimilated resources and trophic positions for an omnivorous fish, Jenynsia multidentata, that reveals dietary variation among locations across a salinity gradient of a coastal lagoon in southern Brazil. We also compared estimates of foods ingested vs. foods assimilated. Food assimilation was estimated using carbon (δ¹³C) and nitrogen (δ¹⁵N) stable isotope ratios of food sources and consumer muscle tissue and an isotopic mixing model (SIAR); consumer trophic position (TP) was estimated from consumer and production source δ¹⁵N values. Diet was estimated using an index of relative importance based on frequency of occurrence and volumetric and numeric proportions of food items from stomach contents. The effect of variation in TDF on food assimilation and TP was tested using three alternative TDFs reported in review papers. We then created a new method that used food source-specific TDFs (reported separately for herbivores and carnivores) weighted in proportion to estimated assimilation of resources according to mixing model estimates to estimate TP (hereafter TP_WAR). We found that plant material was not assimilated in a proportion similar to its importance in the diet of fish at a freshwater site, and the new method yielded best assimilation estimates. Animal material made greatest contributions to fish biomass irrespective of TDFs used in the mixing model. The new method produced TP estimates consistent with differences in estimated food assimilation along the salinity gradient. Our findings support the idea that food source-specific TDFs should be used in trophic studies of omnivores, since the method improved our ability to estimate trophic position and resource assimilation, two important ecological indicators.