Baseline δ15N Research Articles

Estimating ethanol correction factors for δ13C and δ15N isotopic signatures of freshwater zooplankton from multiple lakes

AbstractIn freshwater systems, δ13C and δ15N stable isotopes can be used to differentiate between pelagic and littoral energy sources and to quantify trophic position. In these ecosystems, crustacean zooplankton are frequently used to characterize the pelagic baseline. Zooplankton samples are often preserved prior to processing and analysis, which can affect isotopic signatures. Variability in preservation effects across studies make it difficult to determine if and how to correct for preservation effects. Here, we develop a correction factor for ethanol preservation and present a flexible statistical method that can be updated with additional data to increase its applicability. We collected zooplankton from five lakes in Minnesota, USA encompassing wide isotopic ranges (δ13C from −37.23‰ to −23.96‰; δ15N from 3.07‰ to 14.44‰). Changes in zooplankton δ13C and δ15N signatures were quantified using a Bayesian hierarchical model predicting fresh values from ethanol‐preserved values. Ethanol preservation increased δ13C by a factor of 1.158 (95% CI 0.866–1.441) and had a negligible effect on δ15N (slope = 1.077; 95% CI 0.833–1.359). Lake‐specific values did not differ from the overall relationship. K‐fold and leave‐one‐out cross validation tests verified that both models were accurate; RMSE of predicted δ13C = 0.701 and RMSE of predicted δ15N = 0.590. Our correction factors could be applied to other systems in which baseline δ13C and δ15N values fall within the range of our study, and this approach also enables the inclusion of data from additional lakes to estimate new corrections.

Tropical decadal variability in nutrient supply and phytoplankton community in the Central Equatorial Pacific during the late Holocene

We have reconstructed baseline δ15N and δ13C of export production at Kingman Reef in the Central Equatorial Pacific (CEP) at sub-decadal resolution, nearly continuously over the last 2000 years. The changes in δ15N reflects the strength of the North Equatorial Counter Current (NECC) relative to the South Equatorial Current (SEC), and to a lesser extent, the North Equatorial Current (NEC). Seasonal to multi-decadal variation in the strength of these currents, through the redistribution of heat, have global climate impacts and influence marine and terrestrial ecosystems. We use modern El Niño-La Nina dynamics and the Tropical Pacific Decadal Variability (TPDV) pattern, which is defined in the CEP, as a framework for analyzing the isotopic data. The CEP δ15N and δ13C records exhibit multi-decadal (50–60 year) variability consistent with TPDV. A large multi-centennial feature in the CEP δ15N data, within age-model uncertainties, is consistent with one of the prolonged dry-pluvial sequences in the American west at the end of the Medieval Climate Anomaly, where low TPDV is correlated with drier conditions. This unique record shows that the strength of the NECC, as reflected in baseline δ15N and δ13C, has at quasi-predictable intervals throughout the late Holocene, toggled the phytoplankton community between prokaryotes and picoplankton versus eukaryotes.

Amino acid‐specific nitrogen stable isotope analysis reveals the trophic behavior of Icelandic fin whales in winter and suggests variable feeding strategies

AbstractFin whales (Balaenoptera physalus) aggregate west of Iceland during summer, where they feed mostly on krill, but their winter distribution is poorly known. This study investigates the winter behavior of fin whales that summer off Iceland by analyzing amino acid‐specific nitrogen isotope ratios in the baleen plates. This technique is increasingly used in large marine vertebrates to discriminate between changes in trophic position and changes in isotopic baseline levels, while ruling out the effect of fasting on these patterns. Analyses were conducted on samples extracted from two baleen plate segments: one corresponding to winter, another to summer. The trophic position was higher and slightly more variable in winter (3.6 ± 0.7) than in summer (3.0 ± 0.4), which indicates that during winter fin whales tend to feed on fish and are more generalist in feeding habits. Interestingly, the δ15NPhe values were not lower in winter than in summer (winter: 5.2‰ ± 2.4‰; summer: 4.0‰ ± 2.1‰). This was unexpected because baseline δ15N values (e.g., zooplankton) tend to be lower at low latitudes, where fin whales are believed to spend the winter. Fin whales may remain in relatively high latitudes during winter or visit low‐latitude areas with locally higher than average δ15N values, such as upwelling areas.

Compound-specific stable nitrogen isotope analysis of amino acids shows that bulk methods provide higher estimates of mercury biomagnification in the Gulf of St. Lawrence

Compound-specific stable isotope analysis of amino acids (CSIA-AA) provides a method to estimate baseline δ15N values of food chains, allowing less biased estimates of trophic positions for organisms. Greater accuracy in trophic positions can improve estimates of contaminant biomagnification. We calculated trophic positions with various CSIA-AA equations for four species of fish and northern gannets (Morus bassanus) from the Gulf of St. Lawrence. We examined the effect of CSIA-AA-derived trophic positions on mercury biomagnification metrics (trophic magnification factors (TMF) and biomagnification factors) and compared these with trophic position estimates and metrics obtained from traditional bulk stable isotope analysis. The TMFs for the CSIA-AA equations ranged from 10 to 19, and bulk stable isotope analysis produced TMFs of 43, one of the highest TMFs recorded yet in the literature. Biomagnification factors between prey and northern gannets ranged from 20 to 42 using dietary observations and stable isotope mixing models. Our study demonstrates that discrepancies in biomagnification assessed using different approaches may go undetected when using a single approach.

Nitrogen palaeo-isoscapes: Changing spatial gradients of faunal δ15N in late Pleistocene and early Holocene Europe.

Nitrogen isotope ratio analysis (δ15N) of animal tissue is widely used in archaeology and palaeoecology to investigate diet and ecological niche. Data interpretations require an understanding of nitrogen isotope compositions at the base of the food web (baseline δ15N). Significant variation in animal δ15N has been recognised at various spatiotemporal scales and related to changes both in baseline δ15N, linked to environmental and climatic influence on the terrestrial nitrogen cycle, and animal ecology. Isoscapes (models of isotope spatial variation) have proved a useful tool for investigating spatial variability in biogeochemical cycles in present-day marine and terrestrial ecosystems, but so far, their application to palaeo-data has been more limited. Here, we present time-sliced nitrogen isoscapes for late Pleistocene and early Holocene Europe (c. 50,000 to 10,000 years BP) using herbivore collagen δ15N data. This period covers the Last Glacial-Interglacial Transition, during which significant variation in the terrestrial nitrogen cycle occurred. We use generalized linear mixed modelling approaches for interpolation and test models which both include and exclude climate covariate data. Our results show clear changes in spatial gradients of δ15N through time. Prediction of the lowest faunal δ15N values in northern latitudes after, rather than during, the Last Glacial Maximum is consistent with the Late Glacial Nitrogen Excursion (LGNE). We find that including climatic covariate data does not significantly improve model performance. These findings have implications for investigating the drivers of the LGNE, which has been linked to increased landscape moisture and permafrost thaw, and for understanding changing isotopic baselines, which are fundamental for studies investigating diets, niche partitioning, and migration of higher trophic level animals.

Calibrating bulk and amino acid δ13C and δ15N isotope ratios between bivalve soft tissue and shell for paleoecological reconstructions

Ecological isotope proxies measured in ancient bivalve shell matrix protein have great promise for paleoecological reconstruction. Compound-specific isotopes of amino acids (CSI-AA) may be an ideal tool for developing paleoecological proxies, as initial work has indicated that CSI-AA is less subject to alteration under geologic conditions relative to bulk isotope values. While CSI-AA proxies have been applied in modern bivalve soft tissues and a few shell studies, they have yet to be systematically investigated in shell matrix protein. Here, we measured stable isotope values of carbon (δ13C) and nitrogen (δ15N) in both bulk and individual amino acids (AA), comparing soft tissue and shell organic fractions to test the fidelity of a suite of ecological isotope proxies in shell. We sampled three ubiquitous bivalve species in four seasons for one year from two common coastal environments: littoral rocky intertidal and estuarine delta ecosystems. Particulate organic matter (POM) was simultaneously collected to investigate relationships between tissue types and local POM isotope signatures. The ecological proxies tested include trophic niche breadth from bulk isotopes, and baseline δ13C and δ15N values, resource contribution, and trophic level from CSI-AA data. We found niche breadth corresponded well between tissue types, but that bulk isotope values were significantly higher in shell compared to soft tissue. In the CSI-AA record, there were no differences in baseline δ13C and δ15N proxies between tissue types. While no consistent seasonal trends were observed in the δ13C record, summer bulk and baseline δ15N values were lowest in both ecosystems in both tissues. Food resource contribution estimates from shell also closely matched soft tissue, however trophic level estimates were consistently higher in shell, attributed to a systematic offset in Glutamic acid δ15N values. We therefore propose here a new mollusk-specific trophic discrimination factor, and corresponding shell isotope enrichment factor to correct for isotopic routing, both required for accurate paleoecological reconstructions.

Combining isotopic analysis of bulk-skin and individual amino acids to investigate the trophic position and foraging areas of multiple cetacean species in the western South Atlantic

We investigated the trophic structure and habitat use of ten cetacean species occurring in the oceanic waters of the western South Atlantic using naturally-occurring stable isotopes. We analysed δ15N in individual amino acids (AA) to estimate cetacean trophic position (TP) and to evaluate the spatial differences in baseline δ15N (source AAs). We adjusted cetacean bulk-skin δ13C and δ15N for the effect of trophic level using their estimated TPs, obtaining δ13CAdjusted and δ15NAdjusted, respectively. These values were applied to estimate the overlap in the niche areas of cetacean baseline sources. Our analyses showed spatial segregation between Steno bredanensis and the remaining odontocetes, and the high δ15N in this species reflects its occurrence in neritic waters of the southern region. The highest TPs were observed in Physeter macrocephalus, Stenella attenuata and Globicephala melas, while the lowest TPs were reported for S. longirostris, S. clymene and Orcinus orca. Overall, source AA-δ15N showed similar patterns as those of baseline-δ15N (zooplankton) and were higher in species sampled in the southernmost region of the study area (e.g., Delphinus delphis). Isotopic niche areas estimated using δ13CAdjusted and δ15NAdjusted suggested high overlap in foraging area between S. frontalis and Tursiops truncatus, with the latter occupying a higher TP. Our analyses of δ15N in AAs provide a unique insight into the trophic ecology, forage areas and spatial segregation in resource use among these cetacean populations. Additionally, our work provides AA-δ15N baseline for future studies on the trophic ecology and habitat use of marine organisms in the western South Atlantic.

Shifting baselines: Integrating ecological and isotopic time lags improves trophic position estimates in aquatic consumers

Nitrogen stable isotope ratios (δ15N) are routinely used to estimate consumer trophic positions (TPs). This method is very sensitive to the chosen isotopic baseline, which should incorporate all temporal isotopic variability occurring in the base of the food web sustaining the consumers. However, there is still no consensus on the different methods to represent time intervals for isotopic assimilation. It is uncertain if researchers should compute a single baseline obtained by averaging δ15N source values across all time periods, multiple baselines based on δ15N source values collected concomitantly with consumers, or consider a time lag between sources and consumers. Aiming to provide empirical evidence to help select the best option, we designed 4 methods using different temporal lags between consumers and producers based on a 9 yr time series of carbon and nitrogen stable isotopes in an estuary. The 4 tested baseline methods considered the δ15N average of producers sampled in all years (Global); the same season as consumers (Concomitantly); the season before the collection of consumers (Delayed); and the previous season as well as the same season of consumer sampling (Interpolation). Comparison of results with an expected model (based on stomach content data) showed that the Delayed method is the most appropriate. We conclude that time lags in the incorporation of stable isotope values between sources and consumers must be considered in baseline δ15N calculations to improve TP estimates and provide more reliable modeling results. Additionally, we provide other recommendations to improve sampling designs when using stable isotopes in TP estimations.

Beyond bulk δ15N: Combining a suite of stable isotopic measures improves the resolution of the food webs mediating contaminant signals across space, time and communities

Top predators are used as indicators of contaminant trends across space and time. However, signals are integrated over complex food webs, and variation in diet may confound such signals. Trophic position, assessed by bulk δ15N, is widely used to infer the variation in diet relevant to contamination, yet a single variable cannot completely describe complex food webs. Thus, we examined relationships across three aquatic systems varying from a single species to a small food web using bulk values from four isotopes and 21 amino acid-specific values. Because variation in baseline ('source') δ15N can confound estimates of trophic position , we calculated trophic position from the difference between δ15Ntrophic (δ15N for amino acids that change with trophic position) and δ15Nsource (δ15N for amino acids that do not change with trophic position). Across all three systems, variation in δ15Nsource explained over half of the variation in bulk δ15N, and stable isotope values that reflected the base of the food web (δ13C, δ18O, δ34S) predicted contaminants as well or better than δ15N—which was supported by a meta-analysis of other studies. In ospreys feeding in lakes, variation in δ15Nsource across space created a spurious relationship between ΣDDT and apparent trophic position, and masked a relationship between ΣPCB and trophic position. In a seabird guild, changes in diet over time obscured temporal variation in contaminants over five decades. In Arctic fish and invertebrates, more accurate trophic magnification factors were calculated using δ15Ntrophic-source. Thus, (1) using δ15Ntrophic-source, instead of bulk δ15N, avoided incorrect conclusions and improved accuracy of trophic magnification factors necessary to assess risk to top predators; and (2) diet assessed with multiple spatial isotopes, rather than δ15N alone, was essential to understand patterns in contaminants across space, time and biological communities. Trophic position was most important for lipophilic ‘legacy’ contaminants (ΣDDT, ΣPCB) and habitat was most important for other contaminants (ΣPBDE, ΣPFAS, mercury). We argue that the use of amino acid-specific analysis of δ15N alongside ‘non-trophic’ isotopes should be a core feature of any study that examines the influence of trophic position on chemical pollution, as required for a chemical to be added to international conventions such as the Stockholm Convention.

Body size and stable isotope composition of zooplankton in the western tropical Atlantic

Abstract Size-based approaches are paramount tools for the study of marine food webs. Here, we investigated the relationship between zooplankton body size, stable isotope composition and trophic level (TL) along a large-scale onshore-offshore gradient in the western tropical Atlantic. Samples were obtained on the Brazilian continental shelf, slope and in oceanic waters (off Fernando de Noronha archipelago and Rocas Atoll) in September and October 2015. Zooplankton was sieved into five size fractions. Zooplankton was dominated by copepods, except for the largest (>2000 μm) size fraction, that showed a high biovolume of chaetognaths, decapods, and fish larvae. Maximum zooplankton abundance and biovolume was found at the continental slope. POM showed consistently lower δ13C than zooplankton, indicating a selective use of 13C-rich primary food sources by zooplankton. Particulate organic matter (POM) was more 13C-enriched in shelf areas (average: −22.8, −23.6 and − 24.3‰ at the shelf, slope and oceanic islands, respectively), probably due to the higher abundance of diatoms nearshore. POM had δ15N values between 2.5 and 6.9‰ (average: 4.0, 4.9 and 4.2‰ at the shelf, slope and oceanic islands, respectively). Zooplankton δ15N and TL increased with body size. The δ15N of the 200–500 μm size fraction was used as baseline for TL estimation. Oceanic areas (average baseline δ15N = 5.8‰ ± 0.52, n = 14) showed a higher baseline δ15N than the shelf (average = 3.9‰ ± 0.69, n = 9) and the slope areas (average = 3.1‰ ± 0.93, n = 9). In spite of differing baselines, the δ15N data produced a consistent pattern of log-linear increase in TL with increasing size, in all areas. The choice of input trophic enrichment factor (TEF) values only slightly changed the log10 (body size) vs TL slopes, but this choice had a considerable effect on the estimates of predator/prey size ratio (PPSR) and predator/prey mass ratio (PPMR). Using a TEF above 2.3 leads to unrealistic PPSR and PPMR estimates. Overall average slope was 0.59 ± 0.08 TL μm−1 with TEF = 2.3 and 0.42 ± 0.07 TL μm−1 with TEF = 3.2.

Seasonal and ontogenetic variation of whiting diet in the Eastern English Channel and the Southern North Sea.

An accurate description of trophic interactions is crucial to understand ecosystem functioning and sustainably manage marine ecosystems exploitation. Carbon and nitrogen stable isotopes were coupled with stomach content analyses to investigate whiting (Merlangius merlangus, Linnaeus, 1758) feeding behavior in the Eastern English Channel and Southern North Sea. Whiting juveniles and adults were sampled in autumn and winter to investigate both ontogenetic and seasonal changes. In addition, queen scallops (Aequipecten opercularis) samples were collected along with fish to be used as isotopic benthic baseline. Results indicated an ontogenetic diet change from crustaceans to fish and cephalopods. In autumn, δ15N values generally increased with fish size while in winter, a decrease of δ15N values with fish size was observed, as a potential result of spatial variation in baseline δ15N values. In winter, a nutrient-poor period, an increase in feeding intensity was observed, especially on the copepod Temora longicornis. This study provides further insights into whiting trophic ecology in relation to ontogenetic and seasonal variations, and it confirms the importance of combining several trophic analysis methods to understand ecosystem functioning.

Zooplankton-based δ13C and δ15N isoscapes from the outer continental shelf and slope in the subtropical western South Atlantic

Characterizing the patterns of stable carbon (δ13C) and nitrogen (δ15N) isotopes at the base of the food webs (baseline) is essential in ecological studies for assessing the feeding habits and migration patterns of marine predators. We analysed δ13C and δ15N in zooplankton samples collected in spring and autumn (2012–2015), along the south (SCM) and southeast (NCSM) Brazilian oceanic waters. An increase in δ13C from the shelf break towards the offshore was associated with the influence of continental water input and upwellings along the shelf break, which introduce 13C-depleted inorganic carbon to the surface, where it will be incorporated into the biological system through phytoplankton growth. An opposite trend was observed in δ15N, with higher values along the shelf break related to the intrusion of upwelled deep-water nutrients, contrasting with the oligotrophic offshore waters where N2 fixation takes place, resulting in primary and secondary production that is relatively more depleted in 15N. A latitudinal (north-south) decrease in δ13C along the offshore area coincided with known isotopic patterns in inorganic carbon, which were more 13C-depleted towards higher latitudes. On the other hand, an increase in δ15N was observed towards the south, reflecting the contrast between the N2 fixation in the more oligotrophic waters in the NCSM and the higher nutrient availability in the SCM. Additionally, δ13C was significantly higher in autumn and δ15N was higher in spring. Although inter-annual differences in δ15N were non-significant, δ13C was significantly lower in 2012 than in the later years. This study provides novel information regarding the patterns of baseline δ13C and δ15N in the southwestern Atlantic Ocean, which will be useful for future investigation of the ecology of higher trophic-level organisms occurring in this area.

Food web reconstruction through isotopic compositions of fossil faeces: Insights into the ecology of a late Barremian freshwater ecosystem (Las Hoyas, Cuenca, Spain)

A total of 54 coprolites with different morphologies from the Konservat-Lagerstätte of Las Hoyas (upper Barremian, Cuenca, Spain) were analysed for stable isotopes (δ15N and δ13Corg) to examine the food-web structure of this ancient wetland. The Las Hoyas wetland exhibited wetter and drier periods that indicated water table oscillations. Differences in the δ15N values of coprolites collected from wetter and drier facies were probably due to different baseline δ15N values during the dry periods (higher) compared to those of the wetter periods. The δ13Corg values indicated that aquatic food resources were the most commonly used by the animals from Las Hoyas. Assuming a Trophic Enrichment Factor (TEF) of + 2.5 ‰, three trophic levels were established for the coprolite association. All the specimens of circular, ellipsoidal, elongated, and rosary morphotypes fell into the same trophic level, whereas other morphotypes (cylinder, irregular, and thin lace) exhibited a wide range of δ15N values and the highest % N concentrations. The covariation between δ15N and δ13Corg values describe a wide spectrum of values across the middle and upper part of the ecosystem's food web structure, suggesting that the Las Hoyas ecosystem hosted specialist feeders and omnivorous and/or opportunistic feeders, the latter being the majority. The variation of δ15N values within the specimens of some morphotypes showed that the larger coprolites do not exhibit any correspondence between the faecal size and δ15N values.

Dietary ontogeny of the blue shark, Prionace glauca, based on the analysis of δ13C and δ15N in vertebrae

Ontogenetic changes in habitat and food preferences are common in nature; they reflect changes in the needs of organisms during their lifetime. Studying the dietary ontogeny of pelagic species is a difficult task, because of migratory processes and the inaccessibility of their habitats. As a result, their life history remains poorly understood, or even unknown. Here, we studied the dietary ontogeny of 18 blue sharks, Prionace glauca, using isotopic analysis in vertebrae. A total of 132 samples of vertebral collagen were taken (64 from males and 68 from females). The wide range of δ13C values (− 16.8 to − 13.1‰) suggests that these sharks use both coastal and oceanic areas for feeding. Small juveniles and adults preferred coastal areas, while medium-sized and large juveniles preferred oceanic areas. The estimated δ15N values (9.5–19.0‰) suggest that P. glauca is a top predator that occupies various trophic levels and/or it feeds across areas with different baseline δ15N (trophic position 3.9–8.4). Isotopic enrichment and differences in δ13C and δ15N thus suggest ontogenetic changes in habitat use and prey consumption between maturity stages. The use of hard anatomical structures (vertebrae) is highly relevant because they integrate information on the dietary ontogeny of this shark species.

Paired bulk organic and individual amino acid δ15N analyses of bivalve shell periostracum: A paleoceanographic proxy for water source variability and nitrogen cycling processes

Abstract Developing high-resolution, well-dated, marine proxies of environmental, climatic, and oceanographic conditions is critical in order to advance our understanding of the ocean’s role in the global climate system. While some work has investigated bulk and compound specific stable nitrogen isotope (δ15N) values in bivalve shells as proxies for environmental variability, the small concentrations of nitrogen found in the organic matrix of the shell calcium carbonate (hereafter carbonate) makes developing high resolution records challenging. This study investigates the potential of using the bulk and amino acid δ15N values of bivalve periostracum, the protein layer on the outside of the shell, as a proxy archive of nitrogen cycling processes and water source variability. Bulk δ15N values were measured on the periostracum, carbonate, and adductor muscle of Arctica islandica shells collected in the Gulf of Maine. Increased variability of isotopic values across growth lines compared to along growth lines support mechanistic reasoning based on growth processes that periostracum is recording changes in δ15N values over the course of the clam’s lifetime (up to 500 years). In addition, the statistically significant relationship between periostracum δ15N values and contemporaneous carbonate δ15N values of the same shell (r = 0.82, p Compound specific δ15N analyses of the periostracum of A. islandica shells were used to determine that the calculated trophic position of the clams in this study (1.4 ± 0.4) did not change significantly between 1783 and 1997. Phenylalanine δ15N values over the last 70 years show similar trends to that of the bulk record, suggesting that changes in bulk δ15N values over that time period are related to changes in baseline δ15N values. Periostracum δ15N values from shells collected in the western Gulf of Maine have decreased by ∼1‰ since the mid-1920s. This trend (−0.008‰/year) is not statistically different from the trend of previously published δ15N values of deep-sea corals from the entrance to the Gulf of Maine over the same time period. This coral record has been shown to indicate a shift in water source in the region and therefore the similarity between the two records suggest that changes in periostracum δ15N values are reflecting broader North Atlantic hydrographic changes. Our study introduces a new, high-resolution, and absolutely dated paleoceanographic proxy of baseline δ15N values, presenting the opportunity for future reconstructions of aspects of nitrogen cycling and water source changes in the global oceans.

High Individual Variability in Beak Stable Isotopes of Jumbo Squid off Peruvian Exclusive Economic Zone (EEZ) Waters in the Analysis of Migratory and Foraging Ecology

Stable isotopes (δ13C and δ15N) have been widely used to track cephalopod habitat, migration and trophic structure. In this study, we analyzed the δ13C and δ15N values in 245 Dosidicus gigas beaks collected during 2009–2010 and in 2013 off Peruvian Exclusive Economic Zone (EEZ) waters. High individual variation in beak stable isotopes was shown with values ranging from −19.4‰ to −15.8‰ for δ13C and from 5.0‰ to 15.1‰ for δ15N. A generalized additive model (GAM) showed that latitude, mantle length and distance to shelf break significantly described the isotope variation with deviance ranging from 16.6% in δ13C to 36.3% in δ15N. Large variability in beak δ13C values for a given size and sampling station indicated that D. gigas off the Peruvian EEZ waters migrate in different ways when they occupy a large range of habitats for their ontogeny. Low baseline δ15N values suggested that spatial changes in diet trophic level may be the main determinative factor in beak δ15N variation. We conclude that high variability in beak δ15N values for a given set of explanatory variables indicated that D. gigas is an opportunistic predator with highly diverse dietary habitats. This study further develops our knowledge of the life history of D. gigas in such a highly dynamic region.

A global meta‐analysis of marine predator nitrogen stable isotopes: Relationships between trophic structure and environmental conditions

AbstractAimWe examined potential environmental drivers of broad‐scale spatial patterns in the trophic structure of marine ecosystems as represented by nitrogen stable isotopes in globally distributed marine predators. Additionally, we assessed the effects of spatial scale on the predictive capabilities of environmental variables.LocationGlobal oceans.Time period2000 to 2015.Major taxa studiedTunas: Thunnus albacares, Thunnus obesus, Thunnus alalunga.MethodsWe undertook a global compilation and meta‐analysis of the bulk nitrogen stable isotope ratios (δ15N values) of three tuna species (n = 4,281). After adjusting for regional variations in baseline δ15N values using a global ocean biogeochemistry model, generalized additive mixed models were employed to infer global‐scale oceanographic controls of trophic structure, using cosmopolitan tuna species as a model.ResultsFor the three tuna species, variation in trophic position estimated using bulk δ15N values was largely explained by geographical location and the corresponding oxygen minimum layer depth. Tuna trophic positions declined in areas with reduced oxygen at depth. Food‐chain length, as captured by maximum trophic position, was longer in areas of the western Pacific Ocean and shorter in the northern Atlantic and eastern Pacific Oceans. Trophic adaptability of the tuna predators, as indicated by intraspecific variability, was highest in the western and central Pacific Ocean and lowest in the northern Atlantic Ocean. Our analysis demonstrated that while tunas share similar functional trophic roles, deeper‐foraging tuna species had higher trophic positions globally. The predictive capacity of environmental variables decreased at finer (regional) spatial scales.Main conclusionsOur work suggests that habitat compression resulting from the predicted global expansion of oxygen minimum zones with ocean warming will impact the trophic structure of marine food webs and the corresponding foraging habits of marine predators. Spatial scale analyses highlighted the importance of representing differences in regional ecological dynamics in global‐scale trophic and ecosystem models.

Nitrogen isotopic analysis of carbonate-bound organic matter in modern and fossil fish otoliths

The nitrogen isotopic composition (δ15N) of otolith-bound organic matter (OM) is a potential source of information on dietary history of bony fishes. In contrast to the δ15N of white muscle tissue, the most commonly used tissue for ecological studies, the δ15N of otolith-bound OM (δ15Noto) provides a record of whole life history. More importantly, δ15Noto can be measured in contexts where tissue is not available, for example, in otolith archives and sedimentary deposits. The utility and robustness of otolith δ15N analysis was heretofore limited by the low N content of otoliths, which precluded the routine measurement of individual otoliths as well as the thorough cleaning of otolith material prior to analysis. Here, we introduce a new method based on oxidation to nitrate followed by bacterial conversion to N2O. The method requires 200-fold less N compared to traditional combustion approaches, allowing for thorough pre-cleaning and replicated analysis of individual otoliths of nearly any size. Long term precision of δ15Noto is 0.3‰. Using an internal standard of Atlantic cod (Gadus morhua) otoliths, we examine the parameters of the oxidative cleaning step with regard to oxidant (potassium persulfate and sodium hypochlorite), temperature, and time. We also report initial results that verify the usefulness of δ15Noto for ecological studies. For three salmonid species, left and right otoliths from the same fish are indistinguishable. We find that the δ15Noto of pink salmon (Oncorhynchus gorbuscha) is related to the size of the fish for this species. We find that intra-cohort δ15Noto standard deviation for wild pink salmon, farmed brown trout (Salmo trutta), and farmed rainbow trout (Oncorhynchus mykiss) are all 0.4‰ or less, suggesting that δ15Noto will be valuable for population-level studies. Lastly, our protocol yields reproducible data for both δ15Noto and otolith N content in 17th century Atlantic cod otoliths. We find that 17th century cod are approximately 2 ‰ higher than modern cod, arguably consistent with either the larger size of the otoliths (and thus inferred for the fish) or with changes in baseline (primary producer) δ15N in the modern coastal ocean compared to the past. All told, the results of this study bode well for the utility of otolith-bound δ15N for investigating the environment and ecology of modern and past fish.

Trophic position increases with thermocline depth in yellowfin and bigeye tuna across the Western and Central Pacific Ocean

Estimates of trophic position are used to validate ecosystem models and understand food web structure. A consumer’s trophic position can be estimated by the stable nitrogen isotope values (δ15N) of its tissue, once the baseline isotopic variability has been accounted for. Our study established the first data-driven baseline δ15N isoscape for the Western and Central Pacific Ocean using particulate organic matter. Bulk δ15N analysis on 1039 muscle tissue of bigeye and yellowfin tuna were conducted together with amino acid compound-specific δ15N analysis (AA-CSIA) on a subset of 21 samples. Both particulate organic matter and tuna bulk δ15N values varied by more than 10‰ across the study area. Fine-scaled trophic position maps were constructed and revealed higher tuna trophic position (by ∼1) in the southern latitudes compared to the equator. AA-CSIA confirmed these spatial patterns for bigeye and, to a lesser extent, yellowfin tuna. Using generalized additive models, spatial variations of tuna trophic positions were mainly related to the depth of the 20°C isotherm, a proxy for the thermocline behavior, with higher tuna trophic position estimates at greater thermocline depths. We hypothesized that a deeper thermocline would increase tuna vertical habitat and access to mesopelagic prey of higher trophic position. Archival tagging data further suggested that the vertical habitat of bigeye tuna was deeper in the southern latitudes than at the equator. These results suggest the importance of thermocline depth in influencing tropical tuna diet, which affects their vulnerability to fisheries, and may be altered by climate change.

Amino Acid Specific Stable Nitrogen Isotope Values in Avian Tissues: Insights from Captive American Kestrels and Wild Herring Gulls.

Through laboratory and field studies, the utility of amino acid compound-specific nitrogen isotope analysis (AA-CSIA) in avian studies is investigated. Captive American kestrels (Falco sparverius) were fed an isotopically characterized diet and patterns in δ15N values of amino acids (AAs) were compared to those in their tissues (muscle and red blood cells) and food. Based upon nitrogen isotope discrimination between diet and kestrel tissues, AAs could mostly be categorized as source AAs (retaining baseline δ15N values) and trophic AAs (showing 15N enrichment). Trophic discrimination factors based upon the source (phenylalanine, Phe) and trophic (glutamic acid, Glu) AAs were 4.1 (muscle) and 5.4 (red blood cells), lower than those reported for metazoan invertebrates. In a field study involving omnivorous herring gulls (Larus argentatus smithsonianus), egg AA isotopic patterns largely retained those observed in the laying female's tissues (muscle, red blood cells, and liver). Realistic estimates of gull trophic position were obtained using bird Glu and Phe δ15N values combined with β values (difference in Glu and Phe δ15N in primary producers) for aquatic and terrestrial food webs. Egg fatty acids were used to weight β values for proportions of aquatic and terrestrial food in gull diets. This novel approach can be applied to generalist species that feed across ecosystem boundaries.