Ratios δ15N Research Articles

Effects of current regimes and oxygenation on particulate matter preservation on the Namibian shelf: Insights from amino acid biogeochemistry

Many reconstructions of past biogeochemical states rely on proxies such as δ15N that in turn are affected by the preservation state of organic matter. N-turnover processes in the upwelling system on the Namibian shelf are difficult to reconstruct because lateral particle advection and differential degradation under oxic to anoxic conditions complicate particle tracing in suspended matter and surface sediments as well as the interpretation of sedimentary δ15N records. We analysed the amino acid composition of suspended matter, sinking particles, phytoplankton biomass and surface sediments in order to investigate the degradation pathway of organic matter from the sea surface to the surface sediment. A principal component analysis carried out with the amino acid data set results in a triangular-shaped model. Two branches evolve from phytoplankton biomass on the apex: The suspended matter branch is characterized by relatively enhanced contents of Glu, Ser and Leu, whereas the non-protein amino acids and Lys mark the sinking particle and sediment branch. The results suggest that both pools have taken different diagenetic pathways and that there is only limited exchange between them. The preservation state of organic matter in surface sediments within the oxygen minimum zone is generally better than on the oxic outer shelf, where organic matter preservation depends on water depth.A vertical increase in δ15N ratios from phytoplankton biomass to the surface sediment by ~2.5‰ is attributed to early diagenetic degradation in the water column and at the water/sediment interface. The δ15N ratios of sediments are mainly controlled by the N-isotope signature of nitrate on the outer shelf and upper continental slope. Nutrient-enriched deep water ascends at the shelf breaks due to the influence of internal waves and tides or curl-driven shelf break upwelling. The δ15N of upwelling nitrate at the shelf breaks is ~3‰ lower than that of nitrate from the inner shelf. This difference in nitrate sources is expressed in a band of decreasing δ15N ratios of surface sediments at the shelf breaks. The hydrodynamic upwelling regime at the shelf breaks flushes significant amounts of suspended matter from the mesopelagic ocean across the shelf break as indicated by amino acid monomer patterns of suspended matter.

Pup Vibrissae Stable Isotopes Reveal Geographic Differences in Adult Female Southern Sea Lion Habitat Use during Gestation.

Individuals within populations often differ substantially in habitat use, the ecological consequences of which can be far reaching. Stable isotope analysis provides a convenient and often cost effective means of indirectly assessing the habitat use of individuals that can yield valuable insights into the spatiotemporal distribution of foraging specialisations within a population. Here we use the stable isotope ratios of southern sea lion (Otaria flavescens) pup vibrissae at the Falkland Islands, in the South Atlantic, as a proxy for adult female habitat use during gestation. A previous study found that adult females from one breeding colony (Big Shag Island) foraged in two discrete habitats, inshore (coastal) or offshore (outer Patagonian Shelf). However, as this species breeds at over 70 sites around the Falkland Islands, it is unclear if this pattern is representative of the Falkland Islands as a whole. In order to characterize habitat use, we therefore assayed carbon (δ13C) and nitrogen (δ15N) ratios from 65 southern sea lion pup vibrissae, sampled across 19 breeding colonies at the Falkland Islands. Model-based clustering of pup isotope ratios identified three distinct clusters, representing adult females that foraged inshore, offshore, and a cluster best described as intermediate. A significant difference was found in the use of inshore and offshore habitats between West and East Falkland and between the two colonies with the largest sample sizes, both of which are located in East Falkland. However, habitat use was unrelated to the proximity of breeding colonies to the Patagonian Shelf, a region associated with enhanced biological productivity. Our study thus points towards other factors, such as local oceanography and its influence on resource distribution, playing a prominent role in inshore and offshore habitat use.

Carbon and nitrogen isotopic variability in bone collagen during the Neolithic period: Influence of environmental factors and diet

Studies on Holocene periods in France and Liguria over the past 15 years provide an important isotopic database (δ13C and δ15N) on human and animal bone collagen during the Neolithic period (ca. 5500–3100 BC cal.; n = 573). The distribution of archaeological sites (n = 30) along a latitudinal transect from the Mediterranean to the Channel offers a broad data base reflecting a variety of environments and potential cultural practices. We propose a new insight into δ13C and δ15N data to understand the variability in both environment and human diet at the onset of farming. Statistical comparisons highlight significant geographical variation in both δ13C and δ15N ratios in most of the examined species and particularly in wild herbivores (Δ13C = 3.8‰; Δ15N = 8.5‰). Higher δ15N and lower δ13C ratios are found in samples from northern France. Conversely, lower nitrogen and higher carbon isotopic ratios are present in samples from the Mediterranean area. Results indicate the probable strong influence of natural factors impacting soil and plant isotopic ratios and passing this variation further on into the whole food chain. Our data indicate that the isotopic baseline depends on the local environmental particularities which must be taken into account in reconstructing human palaeodiets.

A specialised cannibalistic Arctic charr morph in the piscivore guild of a subarctic lake

Arctic charr (Salvelinus alpinus) is generally considered an inferior piscivore compared to brown trout (Salmo trutta). However, we demonstrate that a recently described profundal spawning piscivore PP-morph of Arctic charr in a subarctic lake has evolved typical piscivore traits (i.e. large jaws, robust skulls), similar to the co-occurring brown trout but different from the sympatric littoral spawning omnivore LO-morph of Arctic charr. A few large-sized LO-morph, most large-sized PP-morph and trout constitute the piscivore guild, but they differed in prey fish selection. Of the fish-eating deep-water PP-morph, 52% had consumed charr, with a peak during the polar night period (68–80% in November–January). In contrast, the LO-morph and trout were mainly (~90%) inter-specific piscivores eating stickleback. Consequently, the PP-morph was the main charr predator and started to feed on fish at a smaller size and took larger prey relative to their body length than the trout. Stable isotope (δ13C, δ15N) ratios reflect the dietary specialisations in the upper-water (LO-morph, trout) and the deep-water (PP-morph) environments, besides the piscivorous behaviour of the PP-morph and trout. The existence of a shallow-water piscivore (trout) may explain the origin of this PP-morph, taking benefit of underutilised resources of small-sized Arctic charr in the deep-water habitat.

Stable carbon and nitrogen isotopic compositions of ambient aerosols collected from Okinawa Island in the western North Pacific Rim, an outflow region of Asian dusts and pollutants

Stable carbon (δ13C) and nitrogen (δ15N) isotope ratios were measured for total carbon (TC) and nitrogen (TN), respectively, in aerosol (TSP) samples collected at Cape Hedo, Okinawa, an outflow region of Asian pollutants, during 2009–2010. The averaged δ13C and δ15N ratios are −22.2‰ and +12.5‰, respectively. The δ13C values are similar in both spring (−22.5‰) and winter (−22.5‰), suggesting the similar sources and/or source regions. We found that δ13C from Okinawa aerosols are ca. 2‰ higher than those reported from Chinese megacities probably due to photochemical aging of organic aerosols. A strong correlation (r = 0.81) was found between nss-Ca and TSP, suggesting that springtime aerosols are influenced from Asian dusts. However, carbonates in the Asian dusts were titrated with acidic species such as sulfuric acid and oxalic acid during atmospheric transport although two samples suggested the presence of remaining carbonate. No correlations were found between δ13C and tracer compounds (levoglucosan, elemental carbon, oxalic acid, and Na+). During winter and spring, coal burning is significant source in China. Based on isotopic mass balance, contribution of coal burning origin particles to total aerosol carbon was estimated as ca. 97% in winter, which is probably associated with the high emissions in China. Contribution of NO3− to TN was on average 45% whereas that of NH4+ was 18%. These results suggest that vehicular exhaust is an important source of TN in Okinawa aerosols. Concentration of water-soluble organic nitrogen (WSON) is higher in summer, suggesting that WSON is more emitted from the ocean in warmer season whereas inorganic nitrogen is more emitted in winter and spring from pollution sources in the Asian continent.

Inherent variation in carbon and nitrogen isotopic assimilation in the freshwater macro-invertebrate Cherax destructor

Individual variability in diet source selection has often been cited as the main factor for intra-specific variation of isotopic signatures among food-web consumers. We conducted a laboratory study to test how well the individual variability of the δ13C and δ15N ratios in the muscle of an omnivore consumer (yabby: Cherax destructor) corresponded to the variability of various diet types and diet combinations. We found that C. destructor muscle isotope signatures varied in concert with the composition of single-source diets, and that this variability was low. However, when fed the same proportional mixture of multiple diet sources, comparatively high isotopic variability was observed among specimens. Results suggest that a substantial component of isotopic variability in wild populations may be owing to inherent differences in uptake, absorption, and sequestration among individuals, which is distinct from behaviourally driven individualised diet selection. Considering the potential of such individual variability in assimilation to be present in many different consumer populations, we suggest further testing for a range of species and inclusion of this source of variation, for interpretation of isotopic data for trophic ecology.

Zoobenthos are minor dietary components of small omnivorous fishes in a shallow eutrophic lake

We examined whether small omnivorous fishes (smaller than ~100mm long) integrate littoral, pelagic and benthic pathways in a shallow, eutrophic lake (Lake Izunuma, Japan). The surface of the lake was covered by a dense vegetation of floating-leaved macrophytes, and small species dominated the icthyofauna. We determined the δ13C and δ15N ratios of five dominant species of small omnivorous fishes. Using a stable isotope analysis in the R mixing model, we determined the possible contribution of three potential food sources (epiphytic algae, zooplankton and zoobenthos (larval chironomids)) to omnivorous fish tissue compositions. Four omnivorous fishes (Gnathopogon elongatus elongatus, Pseudorasbora parva, Biwia zezera and Tridentiger obscurus) subsisted largely on epiphytic algae and zooplankton, whereas zoobenthos contributed little to their diets. Acheilognathus rhombeus subsisted mostly on epiphytic algae. Thus, in this shallow, eutrophic lake, omnivorous fishes incorporated both littoral and pelagic production into the food web, but rarely benthic production. The dominant benthic chironomid larvae often burrow several centimetres into the sediment, and the low dietary contribution of zoobenthos to small fishes may be due to inefficiency at foraging on buried benthos associated with fish body size.

Morphologically similar, coexisting hard corals (Porites lobata and P. solida) display similar trophic isotopic ratios across reefs and depths

Recent studies using stable isotope analysis in scleractinian corals have highlighted strong inter- and intra-specific variability in isotopic ratios, but few have excluded the effects of morphology, which affects resource acquisition, potentially confounding this with metabolic differences among species. Differences in the stable isotopic (δ13C and δ15N) ratios of the coral host tissue and photosymbionts of two co-existing, morphologically similar Porites corals (P. lobata and P. solida) were examined across nested spatial scales (inter-reefs and intra-reef) and across depths in Zanzibar, Tanzania. There were few differences between species in either coral host or photosymbiont isotopic ratios, but the two tissues showed different spatial patterns. Photosymbionts showed variation only in their δ13C ratios, which differed among reefs, but not by depth. In contrast, the coral hosts differed in δ13C and δ15N values among reefs and also by depth. Within-reef differences among sites occurred only for photosymbionts at one reef. The absence of differences in isotopic ratios between the two Porites species across reefs and depths, confirms that highly related and morphologically similar scleractinian corals may occupy similar ecosystem niches, metabolising resources in a similar fashion. This suggests that resource partitioning among corals, and subsequent isotopic variability, is most likely driven by resource acquisition, rather than being inherently species-specific.

Thyme travels: 15N isoscapes of Thymus vulgaris L. invasion in lightly grazed pastoral communities

AbstractAlterations to ecosystem nitrogen (N) cycling by introduced plant species may increase the invasibility of habitat providing a positive feedback for the introduced species to become invasive. Spatial patterns of foliar and soil δ15N ratios reflect variation in rates and process of N‐cycling across invaded landscapes and provide insight into N‐source uptake and utilization strategies of invasive plant species. To evaluate invasion‐associated changes in soil and foliar δ15N at different scales: regional (among different sites), local (between north‐ and south‐facing aspect at the same site), and microsite (within populations in the same community), we measured foliar and soil δ15 N, animal faeces cover (as a proxy for grazing intensity) and N2‐fixing species cover from inside to outside Thymus vulgaris L. (thyme)‐invaded lightly grazed pastoral communities in Central Otago, southern South Island, New Zealand. Mean thyme foliar δ15N were near‐zero across the invaded landscape, and did not change across the advancing edge of invasion or with aspect. There was no evidence that associations with N2‐fixing species provide a potential N source. Soil δ15N was lower inside of thyme compared to at the edge or outside of thyme and was varied between aspects at some sites. Animal faeces cover as a proxy for grazing intensity explained only 23% of this observed variation of soil δ15N. Thyme invasion may result in lowered soil δ15N reflecting alterations to N dynamics. Associated invasion‐related impacts of animal grazing may also impact soil δ15N. Further studies are required to distinguish the underlying mechanism responsible for the observed patterns of foliar and soil δ15N values across thyme‐invaded Central Otago landscapes.

Isotopic Evidence for Diet Among Historic Bedouin of Khirbat al‐Mudayna, Jordan

AbstractInterpreting the behaviours of nomadic groups from the archaeological record is particularly challenging owing to the ephemeral remains produced by their lifestyle. As such, human burials – even from isolated contexts – provide a wealth of information about these transient groups. Stable carbon and nitrogen isotope analysis of a series of human skeletal remains (n = 22) from isolated burials were utilized to reconstruct the diet of historic (ca. 13th–19th century CE) Bedouin from the Khirbat al‐Mudayna vicinity in modern‐day Jordan. Carbon isotope values ranging from −17.7 to −12.5‰ point to variable dietary contributions from both C3‐ and C4‐based foods, while corresponding faunal δ13C ratios ( = −19.0 ± 1.7‰, 1σ) exhibit depletion in 13C characteristic of a predominantly C3 diet. While mean human δ15N values ( = 12.7 ± 1.9‰, 1σ) may suggest that the majority of protein consumed by these nomads came from animal and not plant sources, elevated δ15N ratios in some fauna may be indicative of 15N‐enriched soils as reported in some arid desert environments. Considerable variability in both δ13C and δ15N values also points to differences in individual life histories and subsistence economies over time. Despite a lack of temporal cohesion, these data illustrate the information that can be gleaned from the isolated interments of nomadic or semi‐nomadic individuals. Copyright © 2015 John Wiley & Sons, Ltd.

Nitrogen and argon in Sung Valley and Ambadongar carbonatite complexes: Evidence of incomplete homogenization of mantle and recycled components

Nitrogen and argon concentrations and isotopic ratios have been measured in mineral separates (carbonates, apatites and magnetites) of two temporally and spatially different Indian carbonatites from Sung Valley (107Ma) and Ambadongar (65Ma). Gases were extracted by vacuum crushing (carbonates, apatites and magnetites) and stepwise pyrolysis (only apatite) to look for the trapped gases and resolve the different components. δ15N varies from +1.4‰ to +13.6‰ (with accompanying 40Ar/36Ar varying from 900 to 4956) in Sung Valley, and from −3.4 to +14.2 (with accompanying 40Ar/36Ar varying from 353 to 5301) in Ambadongar. This clearly shows the presence of more than one component in these carbonatites. We identify primary and recycled nitrogen and argon in these carbonatites. Additionally, N may be present in other speciation, like solid inorganic form with elevated δ15N ratios up to 16.9‰. Unlike the well-studied carbonatites from Kola, these Indian carbonatites do not show complete homogenization of the different components that can be hosted in different populations of inclusions and, as a result, can be identified. Preservation of such heterogeneity indicates that carbonatite magmatism originated from a small-scale heterogeneous subcontinental mantle initiated during lithospheric rifting.

High pH microbial ecosystems in a newly discovered, ephemeral, serpentinizing fluid seep at Yanartaş (Chimera), Turkey.

Gas seeps emanating from Yanartaş (Chimera), Turkey, have been documented for thousands of years. Active serpentinization produces hydrogen and a range of carbon gases that may provide fuel for life. Here we report a newly discovered, ephemeral fluid seep emanating from a small gas vent at Yanartaş. Fluids and biofilms were sampled at the source and points downstream. We describe site conditions, and provide microbiological data in the form of enrichment cultures, Scanning electron microscopy (SEM), carbon and nitrogen isotopic composition of solids, and PCR screens of nitrogen cycle genes. Source fluids are pH 11.95, with a Ca:Mg of ~200, and sediments under the ignited gas seep measure 60°C. Collectively, these data suggest the fluid is the product of active serpentinization at depth. Source sediments are primarily calcite and alteration products (chlorite and montmorillonite). Downstream, biofilms are mixed with montmorillonite. SEM shows biofilms distributed homogeneously with carbonates. Organic carbon accounts for 60% of the total carbon at the source, decreasing downstream to <15% as inorganic carbon precipitates. δ13C ratios of the organic carbon fraction of solids are depleted (−25 to −28‰) relative to the carbonates (−11 to −20‰). We conclude that heterotrophic processes are dominant throughout the surface ecosystem, and carbon fixation may be key down channel. δ15N ratios ~3‰, and absence of nifH in extracted DNA suggest that nitrogen fixation is not occurring in sediments. However, the presence of narG and nirS at most locations and in enrichments indicates genomic potential for nitrate and nitrite reduction. This small seep with shallow run-off is likely ephemeral, but abundant preserved microterracettes in the outflow and the surrounding area suggest it has been present for some time. This site and others like it present an opportunity for investigations of preserved deep biosphere signatures, and subsurface-surface interactions.

Isotopic variability in a stream longitudinal gradient: implications for trophic ecology

Stable isotope analysis (SIA) constitutes a powerful tool for evaluating trophic ecology. However, natural spatio-temporal isotopic variability may complicate the application of this approach in rivers. In this study, we analysed different sources of variability and potential limitations in the use of SIA along the Tea Stream (NW Iberian Peninsula). Gut content analyses (GCA), δ13C and δ15N ratios for major food web components were evaluated in replicate samples collected at six sites along the mid-downstream gradient of the stream in 2010, 2006 and 2007. In this study, we tested (1) isotopic changes on each basal resource and consumer across sites (spatial variability), (2) specific isotopic patterns on the different food web components within each studied site (within-site variability), (3) isotopic consistency through years (temporal variability), and (4) isotopic differences due to fish size (ontogenetic variability). We found higher variability for δ13C than for δ15N ratios in major food web components along the studied gradient. Additionally, autochthonous basal sources exhibited higher variability than allochthonous ones. A clear decreasing pattern in the δ13C ratios of biofilm was observed towards the stream mouth, which was also reflected in both primary and secondary consumers. The high parallelism between biofilm isotopic patterns and those of higher trophic levels stress the importance of exploring the relevance of this resource fuelling food webs along the Tea Stream. Moreover, the importance of combining GCA and SIA was confirmed, allowing a better interpretation of fish trophic interactions in the stream. Our results proved that a site-specific study perspective (considering longitudinal variability) would be more appropriate when evaluating food web issues in greater detail for the Tea Stream.

Individual dietary patterns during childhood: an archaeological application of a stable isotope microsampling method for tooth dentin

Diet from the late medieval Fishergate House cemetery site (York, UK) is reconstructed using nitrogen and carbon stable isotope ratio analysis from tooth dentin. Deciduous teeth from 42 subadult individuals (fetal to 5–6 years) were used to reconstruct weaning practices at a population and an individual level. This is the first archaeological use of this microsampling method (dentin ≥.3 mg). This method allows an individual's changing diet to be reconstructed from the fetal period through weaning. The fetal signals show a complicated relationship with adult female ratios, having higher δ15N values than expected. At this site, there is an unusual decoupling between peak mortality (4–6 years) and weaning (2 years). The mean δ15N ratios for weaned children were enriched when compared to the adult females (12.4‰ ± 1.29 and 11.4‰ ± 1.1; statistically significant to p < .05). Early childhood diet is surprisingly high in marine fish and/or pork given the low socioeconomic class of the sample. This is a departure in weaned diet from contemporary communities and may be responsible for the unusual disconnect between peak mortality and weaning. When the individual dietary reconstructions were combined with each individual's rib reconstruction the presence of a true child specific diet was clear starting at approximately 2 years of age. Some individuals diverge from the population norm and have an extended breastfeeding period linked to poor health. The increased resolution of microsampling allows bioarchaeologists to test detailed time depended questions about early childhood diet and health.

The utilization of stable isotope analysis for the estimation of the geographic origins of unidentified cadavers

The number of unidentified cadavers is increasing worldwide and the effective methods which reveal their geographic origin are not well known. This study reports on the utilization of δ18O, δ13C, δ2H and δ15N ratios gained through stable isotope analysis of urine samples collected from eight locations: Chiba, Japan; Fuzhou, China; and Denpasar, Indonesia in our pilot study with data from healthy volunteers from five further locations from healthy volunteers: Melbourne and Perth, Australia; Qingdao, China; Turku, Finland and Oklahoma, USA. This study posits that the utilization of δ18O and δ2H is more feasible than δ13C and δ15N stable isotope ratios in differentiating or estimating the origin of human samples. Secondly, this study demonstrated that the δ18O and δ2H stable isotope ratios of urine samples from eight locations differed significantly.

Applicability of stable C and N isotope analysis in inferring the geographical origin and authentication of commercial fish (Mackerel, Yellow Croaker and Pollock)

Globalisation of seafood and aquaculture products and their convenient marketing worldwide, increases the possibility for the distribution of mislabelled products; thereby, underlining the need to identify their origin. Stable isotope analysis is a promising approach to identify the authenticity and traceability of seafood and aquaculture products. In this investigation, we measured carbon and nitrogen stable isotope ratios (δ13C and δ15N) of three commercial fish, viz. Mackerel, Yellow Croaker and Pollock, originating from various countries. Apart from the species-dependent variation in the isotopic values, marked differences in the δ13C and δ15N ratios were also observed with respect to the country of origin. This suggests that C and N isotopic signatures could be reliable tools to identify and trace the origin of commercial fish.

Effect of habitat use and diet on the gastrointestinal parasite community of an avian omnivore from an urbanized environment

With urbanization, anthropogenic foods have grown in importance for several species. A shift away from natural foods is likely to affect trophically transmitted parasites, although this has received limited attention in urban-adapted avian omnivores. We examined the effect of habitat use and diet on gastrointestinal parasite assemblages of Ring-billed Gulls (Larus delawarensis Ord, 1815) breeding in an urbanized region (Montréal, Quebec, Canada) using three approaches depicting different temporal scales. Birds were fitted with global positioning system data loggers to determine habitat-use patterns during a 2–3 day period. Stomach contents were examined, providing dietary information during the preceding hours. Liver carbon (δ13C) and nitrogen (δ15N) stable isotope signatures served as proxy of nutrient assimilation during the preceding 10 days. Endoparasite community variations were best explained by δ13C and δ15N ratios. Individuals with lower δ15N had lower parasite abundance, while δ13C was enriched in birds with less diverse parasites (fewer helminth genera). Tissue depletion in15N in Ring-billed Gulls from this colony has previously been associated with predominant consumption of human-related foods. The present study supports the hypothesis that Ring-billed Gulls exhibiting preference for anthropogenic food sources encounter fewer trophically transmitted parasites, which may have contributed, in part, to their population expansion during the last century.

Combining histology, stable isotope analysis and ZooMS collagen fingerprinting to investigate the taphonomic history and dietary behaviour of extinct giant tortoises from the Mare aux Songes deposit on Mauritius

Taphonomic research of bones can provide additional insight into a site's formation and development, the burial environment and ongoing post-mortem processes. A total of 30 tortoise (Cylindraspis) femur bone samples from the Mare aux Songes site (Mauritius) were studied histologically, assessing parameters such as presence and type of microbial alteration, inclusions, staining/infiltrations, the degree of microcracking and birefringence. The absence of microbial attack in the 4200year old Mare aux Songes bones suggests the animals rapidly entered the soil whole-bodied and were sealed anoxically, although they suffered from biological and chemical degradation (i.e. pyrite formation/oxidation, mineral dissolution and staining) related to changes in the site's hydrology. Additionally, carbon and nitrogen stable isotopes were analysed to obtain information on the animals' feeding behaviour. The results show narrowly distributed δ13C ratios, indicating a terrestrial C3 plant-based diet, combined with a wide range in δ15N ratios. This is most likely related to the tortoises' drought-adaptive ability to change their metabolic processes, which can affect the δ15N ratios. Furthermore, ZooMS collagen fingerprinting analysis successfully identified two tortoise species (C. triserrata and C. inepta) in the bone assemblage, which, when combined with stable isotope data, revealed significantly different δ15N ratios between the two tortoise species. As climatic changes around this period resulted in increased aridity in the Mascarene Islands, this could explain the extremely elevated δ15N ratio in our dataset. The endemic fauna was able to endure the climatic changes 4200years ago, although human arrival in the 17th century changed the original habitat to such an extent that it resulted in the extinction of several species. Fortunately we are still able to study these extinct tortoises due to the beneficial conditions of their burial environment, resulting in excellent bone preservation.

Nearshore energy subsidies support Lake Michigan fishes and invertebrates following major changes in food web structure.

Aquatic food webs that incorporate multiple energy channels (e.g., nearshore benthic and pelagic) with varying productivity and turnover rates convey stability to biological communities by providing independent energy sources. Within the Lake Michigan food web, invasive dreissenid mussels have caused rapid changes to food web structure and potentially altered the channels through which consumers acquire energy. We used stable C and N isotopes to determine how Lake Michigan food web structure has changed in the past decade, coincident with the expansion of dreissenid mussels, decreased pelagic phytoplankton production, and increased nearshore benthic algal production. Fish and invertebrate samples collected from sites around Lake Michigan were analyzed to determine taxa-specific 13C:12C (delta13C) and 15N:14N (delta15N) ratios. Sampling took place during two distinct periods, 2002-2003 and 2010-2012, that spanned the period of dreissenid expansion, and included nearshore, pelagic and profundal fish and invertebrate taxa. The magnitude and direction of the delta13C shift indicated significantly greater reliance upon nearshore benthic energy sources among nearly all fish taxa as well as profundal invertebrates following dreissenid expansion. Although the mechanisms underlying this delta13C shift likely varied among species, possible causes include the transport of benthic algal production to offshore waters and increased feeding on nearshore prey items by pelagic and profundal species. delta15N shifts were more variable and of smaller magnitude across taxa, although declines in delta15N among some pelagic fishes suggest a shift to alternative prey resources. Lake Michigan fishes and invertebrates appear to have responded to dreissenid-induced changes in nutrient and energy pathways by switching from pelagic to alternative nearshore energy subsidies. Although large shifts in energy allocation (i.e., pelagic to nearshore benthic) resulting from invasive species appear to affect total production at upper trophic levels, changes in trophic structure and utilization of novel energy pathways may help to stabilize food webs following species invasions.

Paleodietary changes by penguins and seals in association with Antarctic climate and sea ice extent

Positioned near the top of the food web, the dietary composition of Antarctic penguins and seals can be an excellent indicator of the regional food web and thus the status of the marine ecosystem. The dietary composition of modern penguins and seals has been well investigated; a long-term time series of data on penguin and seal diets, however, are rare. Such data, especially any predating the initiation of human harvesting of fish, whales and seals in Antarctica, are crucial for understanding and predicting responses of regional marine food webs to natural climate changes. Here we review recent progress on research of paleodietary change in Antarctic penguins and seals, specifically the Adelie penguin (Pygoscelis adeliae) and Antarctic fur seal (Arctocephalus gazella). These studies indicate that the dietary changes of penguins correspond quite well with fluctuations in climate and sea ice extent during the Holocene. The depleted δ 15N ratios found in modern Adelie penguins support the “krill surplus hypothesis” in relation to historic human depletion of krill-eating fish, seals and whales.