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  1. Nitrogen is a major limiting element for biological productivity, and thus understanding past variations in nitrogen cycling is central to understanding past and future ocean biogeochemical cycling, global climate cycles, and biodiversity. Organic nitrogen encapsulated in fossil biominerals is generally protected from alteration, making it an important archive of the marine nitrogen cycle on seasonal to million-year timescales. The isotopic composition of fossil-bound nitrogen reflects variations in the large-scale nitrogen inventory, local sources and processing, and ecological and physiological traits of organisms. The ability to measure trace amounts of fossil-bound nitrogen has expanded with recent method developments. In this article, we review the foundations and ground truthing for three important fossil-bound proxy types: diatoms, foraminifera, and corals. We highlight their utility with examples of high-resolution evidence for anthropogenic inputs of nitrogen to the oceans, glacial–interglacial-scale assessments of nitrogen inventory change, and evidence for enhanced CO 2 drawdown in the high-latitude ocean. Future directions include expanded method development, characterization of ecological and physiological variation, and exploration of extended timescales to push reconstructions further back in Earth's history. 
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  2. Abstract

    Eukaryotic microalgae play critical roles in the structure and function of marine food webs. The contribution of microalgae to food webs can be tracked using compound‐specific isotope analysis of amino acids (CSIA‐AA). Previous CSIA‐AA studies have defined eukaryotic microalgae as a single functional group in food web mixing models, despite their vast taxonomic and ecological diversity. Using controlled cultures, this work characterizes the amino acidδ13C (δ13CAA) fingerprints—a multivariate metric of amino acid carbon isotope values—of four major groups of eukaryotic microalgae: diatoms, dinoflagellates, raphidophytes, and prasinophytes. We found excellent separation of essential amino acidδ13C (δ13CEAA) fingerprints among four microalgal groups (mean posterior probability reclassification of 99.2 ± 2.9%). We also quantified temperature effects, a primary driver of microalgal bulk carbon isotope variability, on the fidelity ofδ13CAAfingerprints. A 10°C range in temperature conditions did not have significant impacts on variance inδ13CAAvalues or the diagnostic microalgalδ13CEAAfingerprints. Theseδ13CEAAfingerprints were used to identify primary producers at the base of food webs supporting consumers in two contrasting systems: (1) penguins feeding in a diatom‐based food web and (2) mixotrophic corals receiving amino acids directly from autotrophic endosymbiotic dinoflagellates and indirectly from water column diatoms, prasinophytes, and cyanobacteria, likely via heterotrophic feeding on zooplankton. The increased taxonomic specificity of CSIA‐AA fingerprints developed here will greatly improve future efforts to reconstruct the contribution of diverse eukaryotic microalgae to the sources and cycling of organic matter in food web dynamics and biogeochemical cycling studies.

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  3. Abstract

    Studies of Antarctic paleo‐archives have produced conflicting hypotheses on the relative impact of long‐term climate change and historic exploitation of marine mammals on Southern Ocean krill predator foraging ecology. We disentangle these hypotheses using amino acid stable isotope analysis on a 7000‐yr Holocene archive of Adélie penguin (Pygoscelis adeliae) eggshells to differentiate variation in diet and trophic dynamics from baseline biogeochemical cycling as drivers of the rapid decline in krill predator bulk tissue δ15N values in recent centuries. Contrary to previous hypotheses suggesting solely trophic dynamic mechanisms as drivers of this decline, we identified an abrupt decline in source amino acid δ15N values, indicative of major changes in biogeochemical cycling at the base of the Southern Ocean food web that mirrored the decline in penguin bulk tissue δ15N values. These abrupt shifts in penguin δ15N values and associated biogeochemical cycling aligned with climatic events during the Little Ice Age that decreased surface δ15NNO3−, likely connected to a proposed increase in Ekman upwelling via a southward migration of the Westerlies. This baseline shift was in addition to a long‐term, gradual decline in penguin trophic position over the Holocene that began prior to both recent anthropogenic climate change and a proposed “krill‐surplus” following historic marine mammal exploitation in the 19thand 20thcenturies. In resolving these outstanding hypotheses about drivers of Southern Ocean food web dynamics, this study emphasizes the fundamental importance of climate‐induced variability in biogeochemical cycling on ecological processes and improves the ability of paleo‐archives to inform the ecological consequences of future environmental change in the Southern Ocean.

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  4. null (Ed.)
    Characterizing energy flow and trophic linkages is fundamental to understanding the functioning and resilience of Arctic ecosystems under increasing pressure from climate change and anthropogenic exploitation. We used carbon and nitrogen stable isotopes to examine trophic dynamics and the relative contribution of terrestrial organic matter, water column phytoplankton, and phytobenthos (benthic micro- and macro-autotrophs as well as sea ice algae) to the food webs supporting 45 macroconsumers in three Arctic coastal lagoon ecosystems (Krusenstern, Sisualik, Akulaaq) and the adjacent Kotzebue Sound with varying degrees of connectivity in Cape Krusenstern National Monument, Alaska. A two-source (water column particulate organic matter and benthic sediment organic matter), two-isotope trophic dynamics model informed by a Bayesian isotope mixing model revealed that the Lagoon-Kotzebue Sound coastal ecosystem supported consumers along a trophic position continuum from primary consumers, including amphipods, copepods, and clams to trophic level five predators, such as seastars, piscivorous fishes, seals, and seabirds. The relative contribution of the three primary producer end members, terrestrial organic matter (41 ± 21%), phytoplankton (25 ± 21%), and phytobenthos (34 ± 23%) varied as a function of: 1) consumer foraging ecology and 2) consumer location. Suspension feeders received most of their carbon from food webs based on phytoplankton (49 ± 11%) and terrestrial organic matter (23 ± 5%), whereas herbivores and detritivores received the majority of their carbon from phytobenthos-based food webs, 58 ± 10% and 60 ± 8%, respectively. Omnivores and predators showed more even distributions of resource reliance and greater overall variance among species. Within the invertebrates, the importance of terrestrial organic matter decreased and phytobenthos increased with increasing trophic position. The importance of terrestrial organic matter contribution increased with lagoon proximity to major rivers inputs and isolation from Kotzebue Sound. Several taxa with cultural and subsistence food importance to local communities showed significant reliance (30–90% of baseline carbon) on food chains linked to fresh terrestrial organic matter. Our study indicates that terrestrial-marine linkages are important to the function of Arctic coastal lagoon ecosystems and artisanal fisheries. These linkages are likely to strengthen in the future with regional changes in erosion and runoff associated with climate change and anthropogenic disturbance. 
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  5. Abstract

    Compound‐specific stable isotope analysis of individual amino acids (CSIA‐AA) has emerged as a transformative approach to estimate consumer trophic positions (TPCSIA) that are internally indexed to primary producer nitrogen isotope baselines. Central to accurate TPCSIAestimation is an understanding of beta (β) values—the differences between trophic and source AA δ15N values in the primary producers at the base of a consumers’ food web. Growing evidence suggests higher taxonomic and tissue‐specificβvalue variability than typically appreciated.

    This meta‐analysis fulfils a pressing need to comprehensively evaluate relevant sources ofβvalue variability and its contribution to TPCSIAuncertainty. We first synthesized all published primary producer AA δ15N data to investigate ecologically relevant sources of variability (e.g. taxonomy, tissue type, habitat type, mode of photosynthesis). We then reviewed the biogeochemical mechanisms underpinning AA δ15N andβvalue variability. Lastly, we evaluated the sensitivity of TPCSIAestimates to uncertainty in meanβGlx‐Phevalues and Glx‐Phe trophic discrimination factors (TDFGlx‐Phe).

    We show that variation inβGlx‐Phevalues is two times greater than previously considered, with degree of vascularization, not habitat type (terrestrial vs. aquatic), providing the greatest source of variability (vascular autotroph = −6.6 ± 3.4‰; non‐vascular autotroph = +3.3 ± 1.8‰). Within vascular plants, tissue type secondarily contributed toβGlx‐Phevalue variability, but we found no clear distinction among C3, C4and CAM plantβGlx‐Phevalues. Notably, we found that vascular plantβGlx‐Lysvalues (+2.5 ± 1.6‰) are considerably less variable thanβGlx‐Phevalues, making Lys a useful AA tracer of primary production sources in terrestrial systems. Our multi‐trophic level sensitivity analyses demonstrate that TPCSIAestimates are highly sensitive to changes in bothβGlx‐Pheand TDFGlx‐Phevalues but that the relative influence ofβvalues dissipates at higher trophic levels.

    Our results highlight that primary producerβvalues are integral to accurate trophic position estimation. We outline four key recommendations for identifying, constraining and accounting forβvalue variability to improve TPCSIAestimation accuracy and precision moving forward. We must ultimately expand libraries of primary producer AA δ15N values to better understand the mechanistic drivers ofβvalue variation.

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  6. Abstract

    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 inAAisotope fractionation in birds. We conducted a controlledCSIAfeeding experiment on an avian species, the gentoo penguin (Pygoscelis papua), to examine patterns in individualAAcarbon and nitrogen stable isotope fractionation between diet (D) and consumer (C) (Δ13CC‐Dand Δ15NC‐D, respectively). We found that essentialAAδ13C values and sourceAAδ15N 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 nonessentialAAΔ13CC‐Dvalues 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 (TDFGlu‐Phe= 3.5 ± 0.4‰) that we calculated from the difference in trophic fractionation (Δ15NC‐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‐TDFGlu‐Pheequation with the avian‐specificTDFGlu‐Phevalue from our experiment provided estimates that were more ecologically realistic than estimates using a singleTDFGlu‐Pheof 7.6‰ from the previous literature. Our results provide a quantitative, mechanistic framework for the use ofCSIAin nonlethal, archival feathers to study the movement and foraging ecology of avian consumers.

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