We evaluated annual and regional variation in the dietary niche of Pygoscelis penguins including the sea ice-obligate Adélie penguin ( Pygoscelis adeliae ), and sea ice-intolerant chinstrap ( Pygoscelis antarcticus ) and gentoo ( Pygoscelis papua ) penguins, three species that nest throughout the western Antarctic Peninsula (AP) to test the sea ice trophic interaction hypothesis , which posits that penguin breeding populations with divergent trends, i.e., declining or increasing, are reliant on differing food webs. Our study relies on values of naturally occurring carbon ( 13 C/ 12 C, δ 13 C) and nitrogen ( 15 N/ 14 N, δ 15 N) stable isotopes as integrated proxies of penguin food webs measured over three years at three different breeding colonies. At Anvers Island in the north, where reductions in sea ice and changes in breeding population trends among sympatric sea ice-obligate (Adélie) and sea ice-intolerant (chinstrap and gentoo) penguins have been most notable, our analyses show that all three species of Pygoscelis penguins became more similar isotopically over the reproductive period. By late chick-rearing at Anvers Island, crèched chicks at 5-weeks-old for all species occupied similar trophic positions. Isotopic mixing models indicated that the proportions of prey provisioned by adult penguins to 5-week-old chicks at Anvers Island were generally similar across species within years, consisting primarily of Antarctic krill ( Euphausia superba ). Crèched Adélie chicks had higher δ 13 C and δ 15 N values at Avian and Charcot Islands, southern breeding colonies where sea ice is more prominent and populations of Adélie penguins have increased or remain stable. Trophic position increased with latitude, while the proportions of prey provisioned by Adélie penguin adults to chicks at southern breeding colonies included species typical of high Antarctic marine food webs, especially crystal krill ( Euphausia crystallorophias ). A Bayesian metric for dietary niche width, standard ellipse area (SEA-B), indicated that Pygoscelis penguins with greater population changes in the north had more variability in dietary niche width than stable populations further south. Our results lend insight on marine food web drivers of Pygoscelis penguin reproduction at the regional scale and question the long-standing paradigm that Antarctic krill are the only food web component critical to penguin reproductive survival in this region of the Southern Ocean.
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Isoscape Models of the Southern Ocean: Predicting Spatial and Temporal Variability in Carbon and Nitrogen Isotope Compositions of Particulate Organic Matter
Polar marine ecosystems are particularly vulnerable to the effects of climate change. Warming temperatures, freshening seawater, and disruption to sea‐ice formation potentially all have cascading effects on food webs. New approaches are needed to better understand spatiotemporal interactions among biogeochemical processes at the base of Southern Ocean food webs. In marine systems, isoscapes (models of the spatial variation in the stable isotopic composition) of carbon and nitrogen have proven useful in identifying spatial variation in a range of biogeochemical processes, such as nutrient utilization by phytoplankton. Isoscapes provide a baseline for interpreting stable isotope compositions of higher trophic level animals in movement, migration, and diet research. Here, we produce carbon and nitrogen isoscapes across the entire Southern Ocean (>40°S) using surface particulate organic matter isotope data, collected over the past 50 years. We use Integrated Nested Laplace Approximation‐based approaches to predict mean annual isoscapes and four seasonal isoscapes using a suite of environmental data as predictor variables. Clear spatial gradients in δ13C and δ15N values were predicted across the Southern Ocean, consistent with previous statistical and mechanistic views of isotopic variability in this region. We identify strong seasonal variability in both carbon and nitrogen isoscapes, with key implications for the use of static or annual average isoscape baselines in animal studies attempting to document seasonal migratory or foraging behaviors.
more » « less- Award ID(s):
- 1643845
- NSF-PAR ID:
- 10360177
- Publisher / Repository:
- DOI PREFIX: 10.1029
- Date Published:
- Journal Name:
- Global Biogeochemical Cycles
- Volume:
- 35
- Issue:
- 9
- ISSN:
- 0886-6236
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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BACKGROUND The availability of nitrogen (N) to plants and microbes has a major influence on the structure and function of ecosystems. Because N is an essential component of plant proteins, low N availability constrains the growth of plants and herbivores. To increase N availability, humans apply large amounts of fertilizer to agricultural systems. Losses from these systems, combined with atmospheric deposition of fossil fuel combustion products, introduce copious quantities of reactive N into ecosystems. The negative consequences of these anthropogenic N inputs—such as ecosystem eutrophication and reductions in terrestrial and aquatic biodiversity—are well documented. Yet although N availability is increasing in many locations, reactive N inputs are not evenly distributed globally. Furthermore, experiments and theory also suggest that global change factors such as elevated atmospheric CO 2 , rising temperatures, and altered precipitation and disturbance regimes can reduce the availability of N to plants and microbes in many terrestrial ecosystems. This can occur through increases in biotic demand for N or reductions in its supply to organisms. Reductions in N availability can be observed via several metrics, including lowered nitrogen concentrations ([N]) and isotope ratios (δ 15 N) in plant tissue, reduced rates of N mineralization, and reduced terrestrial N export to aquatic systems. However, a comprehensive synthesis of N availability metrics, outside of experimental settings and capable of revealing large-scale trends, has not yet been carried out. ADVANCES A growing body of observations confirms that N availability is declining in many nonagricultural ecosystems worldwide. Studies have demonstrated declining wood δ 15 N in forests across the continental US, declining foliar [N] in European forests, declining foliar [N] and δ 15 N in North American grasslands, and declining [N] in pollen from the US and southern Canada. This evidence is consistent with observed global-scale declines in foliar δ 15 N and [N] since 1980. Long-term monitoring of soil-based N availability indicators in unmanipulated systems is rare. However, forest studies in the northeast US have demonstrated decades-long decreases in soil N cycling and N exports to air and water, even in the face of elevated atmospheric N deposition. Collectively, these studies suggest a sustained decline in N availability across a range of terrestrial ecosystems, dating at least as far back as the early 20th century. Elevated atmospheric CO 2 levels are likely a main driver of declines in N availability. Terrestrial plants are now uniformly exposed to ~50% more of this essential resource than they were just 150 years ago, and experimentally exposing plants to elevated CO 2 often reduces foliar [N] as well as plant-available soil N. In addition, globally-rising temperatures may raise soil N supply in some systems but may also increase N losses and lead to lower foliar [N]. Changes in other ecosystem drivers—such as local climate patterns, N deposition rates, and disturbance regimes—individually affect smaller areas but may have important cumulative effects on global N availability. OUTLOOK Given the importance of N to ecosystem functioning, a decline in available N is likely to have far-reaching consequences. Reduced N availability likely constrains the response of plants to elevated CO 2 and the ability of ecosystems to sequester carbon. Because herbivore growth and reproduction scale with protein intake, declining foliar [N] may be contributing to widely reported declines in insect populations and may be negatively affecting the growth of grazing livestock and herbivorous wild mammals. Spatial and temporal patterns in N availability are not yet fully understood, particularly outside of Europe and North America. Developments in remote sensing, accompanied by additional historical reconstructions of N availability from tree rings, herbarium specimens, and sediments, will show how N availability trajectories vary among ecosystems. Such assessment and monitoring efforts need to be complemented by further experimental and theoretical investigations into the causes of declining N availability, its implications for global carbon sequestration, and how its effects propagate through food webs. Responses will need to involve reducing N demand via lowering atmospheric CO 2 concentrations, and/or increasing N supply. Successfully mitigating and adapting to declining N availability will require a broader understanding that this phenomenon is occurring alongside the more widely recognized issue of anthropogenic eutrophication. Intercalibration of isotopic records from leaves, tree rings, and lake sediments suggests that N availability in many terrestrial ecosystems has steadily declined since the beginning of the industrial era. Reductions in N availability may affect many aspects of ecosystem functioning, including carbon sequestration and herbivore nutrition. Shaded areas indicate 80% prediction intervals; marker size is proportional to the number of measurements in each annual mean. Isotope data: (tree ring) K. K. McLauchlan et al. , Sci. Rep. 7 , 7856 (2017); (lake sediment) G. W. Holtgrieve et al. , Science 334 , 1545–1548 (2011); (foliar) J. M. Craine et al. , Nat. Ecol. Evol. 2 , 1735–1744 (2018)more » « less
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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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Abstract Stable isotope ratios of H (
δ 2H ), O (δ 18O), and C (δ 13C) are linked to key biogeochemical processes of the water and carbon cycles; however, the degree to which isotope-associated processes are reflected in macroscale ecosystem flux observations remains unquantified. Here through formal information assessment, new measurements ofδ 13C of net ecosystem exchange (NEE ) as well asδ 2H andδ 18O of latent heat (LH ) fluxes across the United States National Ecological Observation Network (NEON) are used to determine conditions under which isotope measurements are informative of environmental exchanges. We find all three isotopic datasets individually contain comparable amounts of information aboutNEE andLH fluxes as wind speed observations. Such information from isotope measurements, however, is largely unique. Generally,δ 13C provides more information aboutLH as aridity increases or mean annual precipitation decreases.δ 2H provides more information aboutLH as temperatures or mean annual precipitation decreases, and also provides more information aboutNEE as temperatures decrease. Overall, we show that the stable isotope datasets collected by NEON contribute non-trivial amounts of new information about bulk environmental fluxes useful for interpreting biogeochemical and ecohydrological processes at landscape scales. However, the utility of this new information varies with environmental conditions at continental scales. This study provides an approach for quantifying the value adding non-traditional sensing approaches to environmental monitoring sites and the patterns identified here are expected to aid in modeling and data interpretation efforts focused on constraining carbon and water cycles’ mechanisms. -
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Abstract Degradation of peatlands via drainage is increasing globally and destabilizing peat carbon (C) stores. The effects of drainage on the timing and magnitude of lateral C losses from degraded peatlands remains understudied. We measured spatial and temporal variability in lateral C exports from three drained peat islands in the Sacramento‐San Joaquin Delta in California across the 2017 and 2018 water years using measurements of dissolved inorganic C (DIC), dissolved organic C (DOC), and suspended particulate organic C (POC) concentration combined with discharge. These measurements were supplemented with stable isotope data (δ13C‐DIC, δ13C‐POC, δ15N‐PON, and δ2H‐H2O values) to provide insight into hydrological and biogeochemical controls on lateral C exports from drained peatlands. Drainage DOC and DIC concentrations were seasonally variable with the highest values in the winter rainy season, when discharge was also elevated. Seasonal differences in the mobilization of dissolved C appeared to result from changing water sources and water table levels. Peat island drainage C contributions to surrounding waterways were also greatest during the winter. Although temporal variability in C cycling processes and trends were generally similar across islands, baseline drainage DIC, DOC, and POC concentrations were spatially variable, likely a result of sub‐island‐scale differences in soil organic matter content and hydrology. This spatial variability complicates system‐wide assessments of C budgets. Net lateral C exports were water year dependent and comparable to previously published vertical C emission rates for this system. This work highlights the importance of including lateral C exports from drained peatlands in local and regional C budgets.
