Long-term stability in the circumpolar foraging range of a Southern Ocean predator between the eras of whaling and rapid climate change.

Assessing environmental changes in Southern Ocean ecosystems is difficult due to its remoteness and data sparsity. Monitoring marine predators that respond rapidly to environmental variation may enable us to track anthropogenic effects on ecosystems. Yet, many long-term datasets of marine predators are incomplete because they are spatially constrained and/or track ecosystems already modified by industrial fishing and whaling in the latter half of the 20th century. Here, we assess the contemporary offshore distribution of a wide-ranging marine predator, the southern right whale (SRW, Eubalaena australis), that forages on copepods and krill from ~30°S to the Antarctic ice edge (>60°S). We analyzed carbon and nitrogen isotope values of 1,002 skin samples from six genetically distinct SRW populations using a customized assignment approach that accounts for temporal and spatial variation in the Southern Ocean phytoplankton isoscape. Over the past three decades, SRWs increased their use of mid-latitude foraging grounds in the south Atlantic and southwest (SW) Indian oceans in the late austral summer and autumn and slightly increased their use of high-latitude (>60°S) foraging grounds in the SW Pacific, coincident with observed changes in prey distribution and abundance on a circumpolar scale. Comparing foraging assignments with whaling records since the 18th century showed remarkable stability in use of mid-latitude foraging areas. We attribute this consistency across four centuries to the physical stability of ocean fronts and resulting productivity in mid-latitude ecosystems of the Southern Ocean compared with polar regions that may be more influenced by recent climate change.

Inter- and intrapopulation resource use variation of marine subsidized western fence lizards.

Marine resource subsidies alter consumer dynamics of recipient populations in coastal systems. The response to these subsidies by generalist consumers is often not uniform, creating inter- and intrapopulation diet variation and niche diversification that may be intensified across heterogeneous landscapes. We sampled western fence lizards, Sceloporus occidentalis, from Puget Sound beaches and coastal and inland forest habitats, in addition to the lizards' marine and terrestrial prey items to quantify marine and terrestrial resource use with stable isotope analysis and mixing models. Beach lizards had higher average δ13C and δ15N values compared to coastal and inland forest lizards, exhibiting a strong mixing line between marine and terrestrial prey items. Across five beach sites, lizard populations received 20-51% of their diet from marine resources, on average, with individual lizards ranging between 7 and 86% marine diet. The hillslope of the transition zone between marine and terrestrial environments at beach sites was positively associated with marine-based diets, as the steepest sloped beach sites had the highest percent marine diets. Within-beach variation in transition zone slope was positively correlated with the isotopic niche space of beach lizard populations. These results demonstrate that physiography of transitional landscapes can mediate resource flow between environments, and variable habitat topography promotes niche diversification within lizard populations. Marine resource subsidization of Puget Sound beach S. occidentalis populations may facilitate occupation of the northwesternmost edge of the species range. Shoreline restoration and driftwood beach habitat conservation are important to support the unique ecology of Puget Sound S. occidentalis.

Pliocene hominins from East Turkana were associated with mesic environments in a semiarid basin.

During the middle Pliocene (∼3.8-3.2 Ma), both Australopithecus afarensis and Kenyanthropus platyops are known from the Turkana Basin, but between 3.60 and 3.44 Ma, most hominin fossils are found on the west side of Lake Turkana. Here, we describe a new hominin locality (ET03-166/168, Area 129) from the east side of the lake, in the Lokochot Member of the Koobi Fora Formation (3.60-3.44 Ma). To reconstruct the paleoecology of the locality and its surroundings, we combine information from sedimentology, the relative abundance of associated mammalian fauna, phytoliths, and stable isotopes from plant wax biomarkers, pedogenic carbonates, and fossil tooth enamel. The combined evidence provides a detailed view of the local paleoenvironment occupied by these Pliocene hominins, where a biodiverse community of primates, including hominins, and other mammals inhabited humid, grassy woodlands in a fluvial floodplain setting. Between <3.596 and 3.44 Ma, increases in woody vegetation were, at times, associated with increases in arid-adapted grasses. This suggests that Pliocene vegetation included woody species that were resilient to periods of prolonged aridity, resembling vegetation structure in the Turkana Basin today, where arid-adapted woody plants are a significant component of the ecosystem. Pedogenic carbonates indicate more woody vegetation than other vegetation proxies, possibly due to differences in temporospatial scale and ecological biases in preservation that should be accounted for in future studies. These new hominin fossils and associated multiproxy paleoenvironmental indicators from a single locale through time suggest that early hominin species occupied a wide range of habitats, possibly including wetlands within semiarid landscapes. Local-scale paleoecological evidence from East Turkana supports regional evidence that middle Pliocene eastern Africa may have experienced large-scale, climate-driven periods of aridity. This information extends our understanding of hominin environments beyond the limits of simple wooded, grassy, or mosaic environmental descriptions.

Comparative isotope ecology of western Amazonian rainforest mammals.

Closed-canopy rainforests are important for climate (influencing atmospheric circulation, albedo, carbon storage, etc.) and ecology (harboring the highest biodiversity of continental regions). Of all rainforests, Amazonia is the world's most diverse, including the highest mammalian species richness. However, little is known about niche structure, ecological roles, and food resource partitioning of Amazonian mammalian communities over time. Through analyses of δ13Cbioapatite, δ13Chair, and δ15Nhair, we isotopically characterized aspects of feeding ecology in a modern western Amazonian mammalian community in Peru, serving as a baseline for understanding the evolution of Neotropical rainforest ecosystems. By comparing these results with data from equatorial Africa, we evaluated the potential influences of distinct phylogenetic and biogeographic histories on the isotopic niches occupied by mammals in analogous tropical ecosystems. Our results indicate that, despite their geographical and taxonomic differences, median δ13Cdiet values from closed-canopy rainforests in Amazonia (-27.4‰) and equatorial Africa (-26.9‰) are not significantly different, and that the median δ13Cdiet expected for mammalian herbivores in any closed-canopy rainforest is -27.2‰. Amazonian mammals seem to exploit a narrower spectrum of dietary resources than equatorial African mammals, however, as depicted by the absence of highly negative δ13Cdiet values previously proposed as indicative of rainforests (<-31‰). Finally, results of keratin and bioapatite δ13C indicate that the predictive power of trophic relationships, and traditional dietary ecological classifications in bioapatite-protein isotopic offset expectations, must be reconsidered.

A new era of isotope ecology: Nitrogen isotope ratio of amino acids as an approach for unraveling modern and ancient food web.

Food web research is rapidly expanding through study of natural fractional abundance of 15N in individual amino acids. This paper overviews the principles of this isotope approach, and from my perspective, reanalyzes applications, and further extends the discussion. It applies kinetic isotope effects that enriches 15N in certain amino acids associated with the metabolic processes, which was clearly demonstrated by observations of both natural ecosystem and laboratory experiments. In trophic processes 'trophic amino acids' such as glutamic acid that significantly enrich 15N, whereas 'source amino acids' such as phenylalanine and methionine show little 15N enrichment. Through various applications conducted over the years, the principles of the method have shown to operate well and disentangle complex food webs and relevant problems. Applications include food chain length estimate, nitrogen resource assessment, tracking fish migration, and reconstruction of paleodiet. With this approach, considerations of a wide range of classical issues have been reinvigorated, while in the same time, new challenging frontiers are emerging.

Dietary isotopes of Madagascar's extinct megafauna reveal holocene browsing and grazing guilds.

Megafauna play a disproportionate role in developing and maintaining their biomes, by regulating plant dispersal, community structure and nutrient cycling. Understanding the ecological roles of extinct megafaunal communities, for example through dietary reconstruction using isotope analysis, is necessary to determine pre-human states and set evidence-based restoration goals. We use δ13C and δ15N isotopic analyses to reconstruct Holocene feeding guilds in Madagascar's extinct megaherbivores, which included elephant birds, hippopotami and giant tortoises that occurred across multiple habitats and elevations. We compare isotopic data from seven taxa and two elephant bird eggshell morphotypes against contemporary regional floral baselines to infer dietary subsistence strategies. Most taxa show high consumption of C3 and/or CAM plants, providing evidence of widespread browsing ecology. However, Aepyornis hildebrandti, an elephant bird restricted to the central highlands region, has isotope values with much higher δ13C values than other taxa. This species is interpreted as having obtained up to 48% of its diet from C4 grasses. These findings provide new evidence for distinct browsing and grazing guilds in Madagascar's Holocene megaherbivore fauna, with implications for past regional distribution of ecosystems dominated by endemic C4 grasses.

Isotopes in teeth and a cryptic population of coastal freshwater seals.

Human activities threaten the biodiversity of aquatic mammals across the globe. Conservation of these species hinges on the ability to delineate movements and foraging behaviors of animals, but gaining such insights is hampered by difficulties in tracing individuals over their lives. We determined isotope ratios in teeth (87 Sr/86 Sr, 13 C/12 C, and 18 O/16 O) to examine lifelong movement and resource-use patterns of a unique freshwater population of a wide-ranging pinniped species (harbor seal [Phoca vitulina]) that resides in Iliamna Lake, Alaska (U.S.A.). This population's potentially unique migratory behavior and use of different trophic resources are unknown. The isotope ratios we measured in teeth showed that seals were born in the lake, remained lifelong residents, and relied principally on resources produced from in the lake, even when seasonally abundant and nutrient-dense spawning anadromous fish (i.e., sockeye salmon [Oncorhynchus nerka]) were available in the lake. Our results illustrate how serial isotope records in teeth, particularly 87 Sr/86 Sr ratios, can be used to quantify how coastal mammal populations exploit both freshwater and marine ecosystems. Understanding lifelong patterns of habitat and resource use is essential information when designing effective conservation plans for threatened coastal mammals. We present the Iliamna Lake harbor seals as a unique case study into how isotope records within teeth can help reveal the cryptic ecology of such a population residing in an intact ecosystem. The results also provide critical baseline information for the Kvichak River system, which is facing an uncertain future due to proposed large-scale industrial development and a rapidly changing climate.

Isotopos Dentales y una Población Críptica de Focas Costeras de Agua Dulce Resumen Las actividades humanas amenazan a la diversidad de mamíferos acuáticos en todo el mundo. La conservación de estas especies depende de la habilidad para delinear los movimientos y los comportamientos de búsqueda de alimento de los animales, pero la obtención de dicha información está obstaculizada por las dificultades en el rastreo de individuos a lo largo del transcurso de sus vidas. Determinamos la proporción de isotopos dentales (87 Sr/86 Sr, 13 C/12 C y 18 O/16 O) para examinar el movimiento a lo largo de la vida y los patrones de uso de recursos de una población única de una especie de pinnípedos de agua dulce con una distribución amplia (foca común [Phoca vitulina]), la cual reside en el lago Iliamna, Alaska (E.U.A.). Se desconocen el comportamiento migratorio potencialmente único de esta población y el uso que le dan a los diferentes recursos tróficos. La proporción de isotopos que medimos en los dientes mostró que las focas nacieron en el lago, permanecieron como residentes de toda la vida y dependieron principalmente de los recursos producidos en el lago, incluso cuando estaban disponibles en aquel lugar por razones reproductivas los peces anádromos abundantes estacionalmente y con densidad de nutrientes (es decir, el salmón rojo [Oncorhynchus nerka]). Nuestros resultados ilustran cómo los registros seriales de isotopos dentales, particularmente la proporción 87 Sr/86 Sr, pueden usarse para cuantificar cómo las poblaciones de mamíferos costeros explotan tanto los ecosistemas marinos como los de agua dulce. El entendimiento de los patrones ontogénicos del uso de recursos y de hábitat es esencial cuando se diseñan planes efectivos de conservación para los mamíferos costeros en peligro. Presentamos a las focas comunes del lago Iliamna como un estudio de caso único sobre cómo los registros de isotopos dentales pueden ayudar a revelar la ecología críptica de dicha población que reside en un ecosistema intacto. Los resultados también proporcionan información importante de línea base para el sistema el río Kvichak, el cual está enfrentando un futuro incierto debido a la propuesta de un desarrollo industrial de gran escala y al rápido clima cambiante.

Stable isotope ecology of black rhinos (Diceros bicornis) in Kenya.

Stable isotope and elemental ratios in hair are influenced by the environment, including both climate and geology. Stable carbon isotopes can be used to give estimates of the C4/CAM fraction of diets of herbivorous mammals; stable nitrogen isotopes are related to the local water deficit; strontium isotopes are determined by the local geology. We studied hair from rhinos in Kenya to determine spatial patterns in δ13C, δ15N, and 87Sr/86Sr ratios. The samples of rhino hair were collected during Kenya Wildlife Service translocation or veterinary activities. δ13C values showed diets dominated by C3 foods, but in some regions the diet, at least seasonally, contained significant quantities (i.e., > ca. 20%) of C4/CAM foods. δ15N values were related to water deficit, with higher δ15N values in regions with high water deficit. 87Sr/86Sr isotope ratios were found to be related to the local geological substrate suggesting that 87Sr/86Sr isotope ratios are provisionally useful for determining the origins of illegal wildlife materials in Kenya and elsewhere in Africa.

Individual isotopic specializations predict subsequent inter-individual variation in movement in a freshwater fish.

Despite many similarities and intuitive links between individual dietary specialization and behavioral inter-individual variation, these phenomena have been studied in isolation, and empirical data confirming relationships between these intraspecific variance sources are lacking. Here we use stable isotope analysis and acoustic telemetry to test the hypothesis that individual specialization in trophic (δ15 N) and littoral/pelagic prey reliance (δ13 C) covary with inter-individual variation in movement in a group of 34 free-swimming burbot (Lota lota). By performing stable isotope analysis on tissues with differing isotopic turnover rates (anal fin and dorsal muscle), in 24 lethally sampled burbot, we demonstrate that individual specialization in trophic niche (δ15 N) and littoral/pelagic prey reliance (δ13 C) occurred within the population. By performing stable isotope analysis on anal fins of a group of telemetry tagged burbot, we were able to show that interactions between trophic niche and littoral/pelagic prey reliance, explained a significant proportion of the subsequent between-individual variance in mean movement rates. These findings demonstrate an empirical connection between behavioral inter-individual variation and dietary specialization, thus providing a substantial expansion of our understanding of the wider ecological consequences of these interesting phenomena.

Stable carbon, oxygen, and nitrogen, isotope analysis of plants from a South Asian tropical forest: Implications for primatology.

Stable isotope analysis of primate tissues in tropical forest contexts is an increasingly popular means of obtaining information about niche distinctions among sympatric species, including preferences in feeding height, forest canopy density, plant parts, and trophism. However, issues of equifinality mean that feeding height, canopy density, as well as the plant parts and plant species consumed, may produce similar or confounding effects. With a few exceptions, researchers have so far relied largely on general principles and/or limited plant data from the study area as references for deducing the predominant drivers of primate isotope variation. Here, we explore variation in the stable carbon (δ13 C), nitrogen (δ15 N), and oxygen (δ18 O) isotope ratios of 288 plant samples identified as important to the three primate species from the Polonnaruwa Nature Sanctuary, Sri Lanka, relative to plant part, season, and canopy height. Our results show that plant part and height have the greatest effect on the δ13 C and δ18 O measurements of plants of immediate relevance to the primates, Macaca sinica, Semnopithecus priam thersites, and Trachypithecus vetulus, living in this monsoonal tropical forest. We find no influence of plant part, height or season on the δ15 N of measured plants. While the plant part effect is particularly pronounced in δ13 C between fruits and leaves, differential feeding height, and plant taxonomy influence plant δ13 C and δ18 O differences in addition to plant organ. Given that species composition in different regions and forest types will differ, the results urge caution in extrapolating general isotopic trends without substantial local baselines studies.

Testosterone, migration distance, and migratory timing in song sparrows Melospiza melodia.

In seasonally migratory animals, migration distance often varies substantially within populations such that individuals breeding at the same site may overwinter different distances from the breeding grounds. Shorter migration may allow earlier return to the breeding grounds, which may be particularly advantageous to males competing to acquire a breeding territory. However, little is known about potential mechanisms that may mediate migration distance. We investigated naturally-occurring variation in androgen levels at the time of arrival to the breeding site and its relationship to overwintering latitude in male and female song sparrows (Melospiza melodia). We used stable isotope analysis of hydrogen (δ(2)H) in winter-grown claw tissue to infer relative overwintering latitude (migration distance), combined with 14years of capture records from a long-term study population to infer the arrival timing of males versus females. Relative to females, males had higher circulating androgen levels, migrated shorter distances, and were more likely to be caught early in the breeding season. Males that migrate short distances may benefit from early arrival at the breeding grounds, allowing them to establish a breeding territory. Even after controlling for sex and date, androgen levels were highest in individuals that migrated shorter distances. Our findings indicate that androgens and migration distance are correlated traits within and between sexes that may reflect individual variation within an integrated phenotype in which testosterone has correlated effects on behavioral traits such as migration.

Bioenergetics, Trophic Ecology, and Niche Separation of Tunas.

Tunas are highly specialized predators that have evolved numerous adaptations for a lifestyle that requires large amounts of energy consumption. Here we review our understanding of the bioenergetics and feeding dynamics of tunas on a global scale, with an emphasis on yellowfin, bigeye, skipjack, albacore, and Atlantic bluefin tunas. Food consumption balances bioenergetics expenditures for respiration, growth (including gonad production), specific dynamic action, egestion, and excretion. Tunas feed across the micronekton and some large zooplankton. Some tunas appear to time their life history to take advantage of ephemeral aggregations of crustacean, fish, and molluscan prey. Ontogenetic and spatial diet differences are substantial, and significant interdecadal changes in prey composition have been observed. Diet shifts from larger to smaller prey taxa highlight ecosystem-wide changes in prey availability and diversity and provide implications for changing bioenergetics requirements into the future. Where tunas overlap, we show evidence of niche separation between them; resources are divided largely by differences in diet percentages and size ranges of prey taxa. The lack of long-term data limits the ability to predict impacts of climate change on tuna feeding behaviour. We note the need for systematic collection of feeding data as part of routine monitoring of these species, and we highlight the advantages of using biochemical techniques for broad-scale analyses of trophic relations. We support the continued development of ecosystem models, which all too often lack the regional-specific trophic data needed to adequately investigate climate and fishing impacts.

Oxygen isotope ratios in primate bone carbonate reflect amount of leaves and vertical stratification in the diet.

The stable isotopic biogeochemistry of free-ranging primates is a unique tool to assess dietary and ecological adaptions among sympatric populations. The present study tested the hypothesis that oxygen isotopes in the bone carbonate of five primate and four ungulate species that live in Kibale National Park, Uganda, would show minimal variability since the species obtain water from a single water source. Bones were analyzed for stable carbon (δ(13) C) and oxygen (δ(18) O) isotope ratios. Results for apatite δ(13) C are consistent with all species feeding in a closed forest habitat and do not exhibit niche partitioning. δ(18) O values, in contrast, cluster by species and correlate positively with the relative contribution of leaves to the whole diet are likely also modified by vertical niche partitioning between taxa within the forest canopy. These results show that biochemical markers from naturally deceased primate remains can aid our understanding of how living animals exploit available resources. Am. J. Primatol. 78:1086-1097, 2016 © 2015 Wiley Periodicals, Inc.

Variation in carbon isotope values among chimpanzee foods at Ngogo, Kibale National Park and Bwindi Impenetrable National Park, Uganda.

Stable isotope values in primate tissues can be used to reconstruct diet in the absence of direct observation. However, in order to make dietary inferences, one must first establish isotopic variability for potential food sources. In this study we examine stable carbon isotope (δ(13) C) values for chimpanzee (Pan troglodytes) food resources from two Ugandan forests: Ngogo (Kibale National Park), and Bwindi Impenetrable National Park. Mean δ(13) C values for plant samples are equivalent at both sites. Plant δ(13) C values are best explained by a multivariate linear model including plant part (leaves, pith, flowers, and fruit), vertical position within the canopy (canopy vs. ground), and taxon (R(2) = 0.6992). At both sites, leaves had the lowest δ(13) C values followed by pith and fruit. Canopy resources have comparable δ(13) C values at the two sites but ground resources have lower δ(13) C values at Ngogo than Bwindi (-30.7 vs. -28.6‰). Consequently, isotopic differences between ground and canopy resources (4.2 vs. 2.2‰), and among plant parts are more pronounced at Ngogo. These results demonstrate that underlying environmental differences between sites can produce variable δ(13) C signatures among primate food resources. In the absence of observation data or isotope values for local vegetation, caution must be taken when interpreting isotopic differences among geographically or temporally separated populations or species. Am. J. Primatol. 78:1031-1040, 2016. © 2016 Wiley Periodicals, Inc.

Interspecific, ontogenetic and temporal variations in stable isotopes of small tuna species in the northeast Atlantic Ocean.

In order to study the trophic level of small tuna species and their contribution to the carbon flow in pelagic food webs, an analysis of carbon and nitrogen stable isotopes was carried out. The investigation was focused on four small tuna species (Auxis rochei, Auxis thazard, Euthynnus alletteratus and Sarda sarda) commonly harvested in the northeast Atlantic Ocean. The isotope analysis showed how the results for S. sarda are different from the rest of the species analysed, with a higher trophic level, similar to other major tuna species. The greatest niche overlap in δ13C and δ15N occurs among A. rochei, A. thazard and E. alletteratus. Auxis rochei and E. alletteratus showed a size-dependent variability in δ15N, and in δ13C for S. sarda. The small tuna S. sarda exhibits the highest migration rates among various geographical areas in comparison to other small pelagic tunas, and the seasonal variability of isotope values in the area studied can be attributed to the incorporation of larger individuals with a higher lipid content. The results of this work provide new information on the ecological role played by small tuna in food webs, which is more complex and varied than currently thought. This knowledge is essential for a more effective management of fisheries.

Effects of lipid extraction on stable isotope ratios of carbon and nitrogen in muscles of freshwater fish.

The extraction of lipids by the Folch method from the muscles of all the fish studied led to statistically significant differences in the values of δ15N. At the same time, lipid extraction led to a statistically significant increase in δ13C in pike and roach, and to a statistically insignificant decrease in δ13C in perch and bream. Thus, lipid extraction cannot serve as a universal method of sample preparation for the analysis of the isotopic composition of carbon (13C/12C) and nitrogen (15N/14N) in fish muscles. The differences between the δ13C values in the samples before and after lipid extraction were statistically investigated by different models. It is shown that mathematical correction method models can be used, but the results are depending on the fish types.

The stable isotope hydrology of Sable Island, Nova Scotia, Canada with implications for evaluating the water budget of wild horses.

We investigated the stable isotope hydrology of Sable Island, Nova Scotia, Canada over a five year period from September, 2017 to August, 2022. The δ2H and δ18O values of integrated monthly precipitation were weakly seasonal and ranged from -66 to -15 ‰ and from -9.7 to -1.9 ‰, respectively. Fitting these monthly precipitation data resulted in a local meteoric water line (LMWL) defined by: δ2H = 7.22 ± 0.21 · δ18O + 7.50 ± 1.22 ‰. Amount-weighted annual precipitation had δ2H and δ18O values of -36 ± 11 ‰ and -6.1 ± 1.4 ‰, respectively. Deep groundwater had more negative δ2H and δ18O values than mean annual precipitation, suggesting recharge occurs mainly in the winter, while shallow groundwater had δ2H and δ18O values more consistent with mean annual precipitation or mixing of freshwater with local seawater. Surface waters had more positive values and showed evidence of isolation from the groundwater system. The stable isotopic compositions of plant (leaf) water, on the other hand, indicate plants use groundwater as their source. Fog had δ2H and δ18O values that were significantly more positive than those of local precipitation, yet had similar 17O-excess values. δ2H values of horsehair from 4 individuals lacked seasonality, but had variations typical to those of precipitation on the island. Differences in mean δ2H values of horsehair were statistically significant and suggest variations in water use may exist between spatially disparate horse communities. Our results establish an important initial framework for ongoing isotope studies of feral horses and other wildlife on Sable Island.

A protocol for distilling animal body water from biological samples and measuring oxygen and hydrogen stable isotopes via cavity ring-down spectroscopy.

The application of stable isotope analysis (SIA) to the fields of ecology and animal biology has rapidly expanded over the past three decades, particularly with regards to water analysis. SIA now provides the opportunity to monitor migration patterns, examine food webs, and assess habitat changes in current and past study systems. While carbon and nitrogen SIA of biological samples have become common, analyses of oxygen or hydrogen are used more sparingly despite their promising utility for tracing water sources and animal metabolism. Common ecological applications of oxygen or hydrogen SIA require injecting enriched isotope tracers. As such, methods for processing and analyzing biological samples are tailored for enriched tracer techniques, which require lower precision than other techniques given the large signal-to-noise ratio of the data. However, instrumentation advancements are creating new opportunities to expand the applications of high-throughput oxygen and hydrogen SIA. To support these applications, we update methods to distill and measure water derived from biological samples with consistent precision equal to, or better than, ± 0.1 ‰ for δ17O, ± 0.3 ‰ for δ18O, ± 1 ‰ for δ2H, ± 2 ‰ for d-excess, and ± 15 per meg for Δ17O.

Stable isotope analysis in food web research: Systematic review and a vision for the future for the Baltic Sea macro-region.

Food web research provides essential insights into ecosystem functioning, but practical applications in ecosystem-based management are hampered by a current lack of knowledge synthesis. To address this gap, we provide the first systematic review of ecological studies applying stable isotope analysis, a pivotal method in food web research, in the heavily anthropogenically impacted Baltic Sea macro-region. We identified a thriving research field, with 164 publications advancing a broad range of fundamental and applied research topics, but also found structural shortcomings limiting ecosystem-level understanding. We argue that enhanced collaboration and integration, including the systematic submission of Baltic Sea primary datasets to stable isotope databases, would help to overcome many of the current shortcomings, unify the scattered knowledge base, and promote future food web research and science-based resource management. The effort undertaken here demonstrates the value of macro-regional synthesis, in enhancing access to existing data and supporting strategic planning of research agendas.

High δ13C values in red squirrels Sciurus vulgaris explained by a reliance on conifer seeds.

Eurasian red squirrels Sciurus vulgaris have unusually high δ13C values compared to other forest rodents, which is seemingly related to the consumption of 13C-enriched conifer seeds. To test this hypothesis, we analyzed the hair of wild and zoo-kept red squirrels, seeds of pine and spruce, and feathers of specialized and opportunistic consumers of conifer seeds, crossbills Loxia spp., and woodpeckers Dendrocopos major. Data on the isotopic composition (δ13C and δ15N values) of hair or feathers of other species of forest rodents and birds were obtained from published studies. The range of mean δ13C values of hair of wild forest rodents (19 species) exceeded 10 ‰. All squirrel species had high 13C content, S. vulgaris having maximum δ13C values. In contrast, S. vulgaris kept in captivity had an isotopic composition similar to other captive rodents. The feathers of crossbills were enriched in 13C compared to other forest birds (15 species), while seeds of coniferous trees often had higher δ13C values compared to seeds of other woody plants. Distinctiveness of the isotopic composition of mammals and birds feeding on the seeds of coniferous trees suggests that this resource can be traced through the entire forest food web.