Stable isotope analysis provides a powerful tool to identify the energy sources which fuel consumers, to understand trophic interactions and to infer consumer trophic position (TP), an important ...concept that describes the ecological role of consumers in food webs. However, current methods for estimating TP using stable isotopes are limited and do not fulfil the complete potential of the isotopic approach. For instance, researchers typically use point estimates for key parameters including trophic discrimination factors and isotopic baselines, and do not explicitly include variance associated with these parameters when calculating TP.
We present “tRophicPosition,” an r package incorporating a Bayesian model for the calculation of consumer TP at the population level using stable isotopes, with one or two baselines. It combines Markov Chain Monte Carlo simulations through JAGS and statistical and graphical analyses using R. We model consumer and baseline observations using relevant statistical distributions, allowing them to be treated as random variables. The calculation of TP—a random parameter—for one baseline follows standard equations linking 15N enrichment per trophic level and the trophic position of the baseline (e.g. a primary producer or primary consumer). In the case of two baselines, a simple mixing model incorporating δ13C allows for the differentiation between two distinct sources of nitrogen, thus including heterogeneity derived from alternatives sources of δ15N.
Methods currently implemented in “tRophicPosition” include loading, plotting and summarizing stable isotope data either from multiple sites and/or communities or a local assemblage; loading trophic discrimination factors from an internal database or generating them; defining and initializing a Bayesian model of TP; sampling posterior parameters; analysing, comparing and plotting posterior estimates of TP and other parameters; and calculating a parametric (non‐Bayesian) TP estimate. Additionally, full documentation including examples, multiple vignettes and code are available for download.
Foreign Language Resumen
El análisis de isótopos estables es una poderosa herramienta para identificar qué vías energéticas alimentan a los consumidores, para entender las interacciones tróficas, y también para inferir la posición trófica (PT) de los consumidores, un concepto importante que describe el rol ecológico de los consumidores en las tramas tróficas. Sin embargo, los métodos actuales para estimar la PT utilizando isótopos estables están limitados y no permiten alcanzar el potencial de la aproximación isotópica. Por ejemplo, los investigadores típicamente utilizan estimaciones puntuales de parámetros claves incluyendo factores de discriminación trófica y líneas base isotópicas, y no incorporan explícitamente la varianza asociada con estos parámetros cuando calculan la PT.
Presentamos “tRophicPosition,” un paquete de R para isótopos estables que incorpora un modelo Bayesiano para el cálculo de la PT en consumidores al nivel ecológico de población, con una o dos líneas base. Este paquete combina simulaciones de cadenas de Markov Monte Carlo a través de JAGS y utilizando los análisis estadísticos y gráficos de R. Modelamos las observaciones de consumidores y líneas base utilizando distribuciones estadísticas relevantes, por lo que son tratadas como variables aleatorias. El cálculo de la PT—un parámetro aleatorio—para una línea base considera ecuaciones estándar que vinculan el enriquecimiento en 15N por nivel trófico y el nivel trófico de la línea base (p.e. un productor primario o un consumidor primario). En el caso de dos líneas base, un modelo de mezcla simple incorporando δ13C permite la diferenciación entre dos fuentes distintas de nitrógeno, incluyendo de esta forma la heterogeneidad derivada de fuentes alternativas de δ15N.
Los métodos actualmente implementados en “tRophicPosition” incluyen cargar, graficar y resumir datos de isótopos estables desde múltiples sitios y/o comunidades o ensambles locales; cargar factores de discriminación trófica desde una base de datos interna o generarlos; definir e inicializar un modelo Bayesiano para la PT; muestrear parámetros a posteriori; analizar, comparar y graficar estimaciones a posteriori de la TP y otros parámetros; y calcular una estimación frecuentista (no Bayesiana) para la PT. Adicionalmente, toda la documentación incluyendo ejemplos, distintas viñetas y el código están disponibles para descarga.
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FZAB, GIS, IJS, KILJ, NLZOH, NUK, OILJ, SAZU, SBCE, SBMB, UL, UM, UPUK
Resource polymorphism-whereby ancestral generalist populations give rise to several specialised morphs along a resource gradient-is common where species colonise newly formed ecosystems. This ...phenomenon is particularly well documented in freshwater fish populations inhabiting postglacial lakes formed at the end of the last ice age. However, knowledge on how such differential exploitation of resources across contrasting habitats might be reflected in the biochemical compositions of diverging populations is still limited, though such patterns might be expected. Here, we aimed to assess how fatty acids (FA)-an important biochemical component of animal tissues-diverged across a polymorphic complex of European whitefish (Coregonus lavaretus) and their closely related monomorphic specialist congener vendace (Coregonus albula) inhabiting a series of six subarctic lakes in northern Fennoscandia. We also explored patterns of FA composition in whitefish's predators and invertebrate prey to assess how divergence in trophic ecology between whitefish morphs would relate to biochemical profiles of their key food web associates. Lastly, we assessed how information on trophic divergence provided by differential FA composition compared to evidence of resource polymorphism retrieved from more classical stomach content and stable isotopic (δ13C, δ15N) information. Examination of stomach contents provided high-resolution information on recently consumed prey, whereas stable isotopes indicated broad-scale patterns of benthic-pelagic resource use differentiation at different trophic levels. Linear discriminant analysis based on FA composition was substantially more successful in identifying whitefish morphs and their congener vendace as distinct groupings when compared to the other two methods. Three major FA (myristic acid, stearic acid, and eicosadienoic acid) proved particularly informative, both in delineating coregonid groups, and identifying patterns of pelagic-benthic feeding throughout the wider food web. Myristic acid (14:0) content and δ13C ratios in muscle tissue were positively correlated across fish taxa, and together provided the clearest segregation of fishes exploiting contrasting pelagic and benthic niches. In general, our findings highlight the potential of FA analysis for identifying resource polymorphism in animal populations where this phenomenon occurs, and suggest that this technique may provide greater resolution than more traditional methods typically used for this purpose.
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DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
Dramatic environmental shifts are occuring throughout the Arctic from climate change, with consequences for the cycling of mercury (Hg). This review summarizes the latest science on how climate ...change is influencing Hg transport and biogeochemical cycling in Arctic terrestrial, freshwater and marine ecosystems. As environmental changes in the Arctic continue to accelerate, a clearer picture is emerging of the profound shifts in the climate and cryosphere, and their connections to Hg cycling. Modeling results suggest climate influences seasonal and interannual variability of atmospheric Hg deposition. The clearest evidence of current climate change effects is for Hg transport from terrestrial catchments, where widespread permafrost thaw, glacier melt and coastal erosion are increasing the export of Hg to downstream environments. Recent estimates suggest Arctic permafrost is a large global reservoir of Hg, which is vulnerable to degradation with climate warming, although the fate of permafrost soil Hg is unclear. The increasing development of thermokarst features, the formation and expansion of thaw lakes, and increased soil erosion in terrestrial landscapes are increasing river transport of particulate-bound Hg and altering conditions for aquatic Hg transformations. Greater organic matter transport may also be influencing the downstream transport and fate of Hg. More severe and frequent wildfires within the Arctic and across boreal regions may be contributing to the atmospheric pool of Hg. Climate change influences on Hg biogeochemical cycling remain poorly understood. Seasonal evasion and retention of inorganic Hg may be altered by reduced sea-ice cover and higher chloride content in snow. Experimental evidence indicates warmer temperatures enhance methylmercury production in ocean and lake sediments as well as in tundra soils. Improved geographic coverage of measurements and modeling approaches are needed to better evaluate net effects of climate change and long-term implications for Hg contamination in the Arctic.
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•Current evidence indicates climate change is influencing Hg cycling in the Arctic.•Permafrost thaw, glacier melt, and coastal erosion are increasing Hg export.•Arctic permafrost is a global Hg reservoir, vulnerable to degradation and release.•Experiments show warmer temperatures increase MeHg in Arctic sediments and soils.•Net effects of climate change on Arctic Hg contamination remain poorly resolved.
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GEOZS, IJS, IMTLJ, KILJ, KISLJ, NLZOH, NUK, OILJ, PNG, SAZU, SBCE, SBJE, UILJ, UL, UM, UPUK, ZAGLJ, ZRSKP
Subarctic lakes are characterised by extreme seasonal variation in light and temperature which influences growth, maturation, condition and resource use of fishes. However, our understanding of how ...seasonal changes affect mercury concentrations of fishes is limited. We conducted a year-round study (3 ice-covered months, 3 open-water months) with open-water inter-annual aspect (3 years: samples from August/September), focusing on total mercury (THg) concentrations and ecological characteristics of a common freshwater fish, European whitefish (Coregonus lavaretus (L.)) from a subarctic lake. We measured THg concentrations from tissues with fast (liver, n = 164) and moderate (muscle, n = 225) turnover rates, providing information on THg dynamics over different temporal scales. In both tissues, lipid-corrected THg concentrations were highest in winter (liver: 1.70 ± 0.88 μg/g, muscle: 0.24 ± 0.05 μg/g) and lowest in summer (liver: 0.87 ± 0.72 μg/g, muscle: 0.19 ± 0.04 μg/g). THg concentrations increased in winter following the summer-autumn dietary shift to pelagic zooplankton and starvation after spawning. Whitefish THg concentrations decreased towards summer, and were associated with consumption of benthic macroinvertebrates and subsequent growth dilution. Mercury bioaccumulated in both tissues with age, both showing the strongest regression slopes in winter and lowest in summer. THg concentrations in liver and muscle tissue were correlated throughout the year, however the correlation was lowest in summer, indicating high metabolism during somatic growing season in summer and growth dilution. Multiple linear regression models explained 50% and 55% of the THg variation in liver and muscle both models dominated by seasonally-variable factors i.e. sexual maturity, δ13C, and condition factor. Seasonally varying bioaccumulation slopes and the higher level of intra-annual variation (21%) in whitefish THg concentration in muscle than the inter-annual accumulation (8%) highlight the importance of including seasonal factors in future THg studies.
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•Year-round variation of THg in fish tissues is poorly understood in subarctic lakes.•THg concentrations of liver and muscle were highest in winter and lowest in summer.•Starvation and planktivory increased THg, while growth dilution decreased THg.•Intra-annual variation of THg in tissues was higher than inter-annual accumulation.•Bioaccumulation of THg was highest in winter and lowest in summer for both tissues.
Strong seasonal variation was observed in THg concentrations and bioaccumulation slopes in muscle and liver tissues, suggesting that the temporal component of sampling should be considered in future THg monitoring and risk assessment programmes.
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GEOZS, IJS, IMTLJ, KILJ, KISLJ, NUK, OILJ, PNG, SAZU, SBCE, SBJE, UL, UM, UPUK, ZRSKP
1. The adaptive radiation of fishes into benthic (littoral) and pelagic (lentic) morphs in post-glacial lakes has become an important model system for speciation. Although these systems are well ...studied, there is little evidence of the existence of morphs that have diverged to utilize resources in the remaining principal lake habitat, the profundal zone. 2. Here, we tested phenotype-environment correlations of three whitefish (Coregonus lavaretus) morphs that have radiated into littoral, pelagic and profundal niches in northern Scandinavian lakes. We hypothesized that morphs in such trimorphic systems would have a morphology adapted to one of the principal lake habitats (littoral, pelagic or profundal niches). Most whitefish populations in the study area are formed by a single (monomorphic) whitefish morph, and we further hypothesized that these populations should display intermediate morphotypes and niche utilization. We used a combination of traditional (stomach content, habitat use, gill raker counts) and more recently developed (stable isotopes, geometric morphometrics) techniques to evaluate phenotype-environment correlations in two lakes with trimorphic and two lakes with monomorphic whitefish. 3. Distinct phenotype-environment correlations were evident for each principal niche in whitefish morphs inhabiting trimorphic lakes. Monomorphic whitefish exploited multiple habitats, had intermediate morphology, displayed increased variance in gillraker-counts, and relied significantly on zooplankton, most likely due to relaxed resource competition. 4. We suggest that the ecological processes acting in the trimorphic lakes are similar to each other, and are driving the adaptive evolution of whitefish morphs, possibly leading to the formation of new species.
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BFBNIB, FZAB, GIS, IJS, INZLJ, KILJ, NLZOH, NMLJ, NUK, OILJ, PNG, SAZU, SBCE, SBMB, UL, UM, UPUK, ZRSKP
European whitefish is a model species for adaptive radiation of fishes in temperate and subarctic lakes. In northern Europe the most commonly observed morphotypes are a generalist (LSR) morph and a ...pelagic specialist (DR) morph. The evolution of a pelagic specialist morph is something of an enigma, however, as this region is characterized by long, dark winters with pelagic primary production limited to a brief window in late summer. We conducted the first winter‐based study of polymorphic whitefish populations to determine the winter ecology of both morphs, and we combined seasonal diet and stable isotope analysis with several proxies of condition in three polymorphic whitefish populations. The generalist LSR morph fed on benthic and pelagic prey in summer but was solely reliant on benthic prey in winter. This was associated with a noticeable but moderate reduction in condition, lipid content and stomach fullness in winter relative to summer. In contrast, the DR whitefish occupied a strict pelagic niche in both seasons. A significant reduction in pelagic prey during winter resulted in severe decrease in condition, lipid content and stomach fullness in DR whitefish in winter relative to summer, with the pelagic morph apparently approaching starvation in winter. We suggest that this divergent approach to seasonal foraging is associated with the divergent life‐history traits of both morphs.
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BFBNIB, FZAB, GIS, IJS, KILJ, NLZOH, NUK, OILJ, SAZU, SBCE, SBMB, UL, UM, UPUK
A mechanistic understanding of how environmental change affects trophic ecology of fish at the individual and population level remains elusive. To address this, we conducted a space-for-time approach ...incorporating environmental gradients (temperature, precipitation and nutrients), lake morphometry (visibility, depth and area), fish communities (richness, competition and predation), prey availability (richness and density) and feeding (population niche breadth and individual trophic specialisation) for 15 native fish taxa belonging to different thermal guilds from 35 subarctic lakes along a marked climate-productivity gradient corresponding to future climate change predictions. We revealed significant and contrasting responses from two generalist species that are abundant and widely distributed in the region. The cold-water adapted European whitefish (
Coregonus lavaretus
) reduced individual specialisation in warmer and more productive lakes. Conversely, the cool-water adapted Eurasian perch (
Perca fluviatilis
) showed increased levels of individual specialism along climate-productivity gradient. Although whitefish and perch differed in the way they consumed prey along the climate-productivity gradient, they both switched from consumption of zooplankton in cooler, less productive lakes, to macrozoobenthos in warmer, more productive lakes. Species with specialist benthic or pelagic feeding did not show significant changes in trophic ecology along the gradient. We conclude that generalist consumers, such as warmer adapted perch, have clear advantages over colder and clear-water specialised species or morphs through their capacity to undergo reciprocal benthic–pelagic switches in feeding associated with environmental change. The capacity to show trophic flexibility in warmer and more productive lakes is likely a key trait for species dominance in future communities of high latitudes under climate change.
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CEKLJ, EMUNI, FIS, FZAB, GEOZS, GIS, IJS, IMTLJ, KILJ, KISLJ, MFDPS, NLZOH, NUK, OILJ, PNG, SAZU, SBCE, SBJE, SBMB, SBNM, UKNU, UL, UM, UPUK, VKSCE, ZAGLJ
Ambient light and temperature show extreme seasonal variation in subarctic lakes due to the midnight sun period in summer and cold polar night period in winter. These changes have clear impacts on ...fish feeding and reproduction cycles, potentially affecting the fatty acid (FA) composition of muscle. Despite extensive research into fish FA over recent decades, we know little about intra‐annual changes of fish FA profile and content.
We studied intra‐annual changes in the FA profile (mol%) and content (mg g‐1 dry weight) of sexually mature European whitefish (Coregonus lavaretus) muscle in a large and deep subarctic lake located in northern Fennoscandia. We collected fish, zooplankton, and benthic macroinvertebrate samples during 3 ice‐covered months, including December (during whitefish spawning), and 3 open‐water months. Fish size, age, sex, stomach content and fullness, and gonadosomatic index were also assessed as co‐variates.
Whitefish changed diet from benthic macroinvertebrates in winter to zooplankton in summer. Generally, whitefish somatic growth was slow and most energy was used for gonad growth. Zooplankton had higher total content and different profile of FA compared to benthic macroinvertebrates. Increased zooplanktivory in summer was detected with higher α‐linolenic acid (ALA, 18:3n‐3) and stearidonic acid (SDA, 18:4n‐3) percentage and content as well as increased the ratio of polyunsaturated FAs (PUFAs) of n‐3 and n‐6 family (n‐3/n‐6 ratio) in fish muscle.
Whitefish gonadal growth and development occur during the summer growing season and continue until the initiation of spawning in early winter. We found that the content of physiologically crucial PUFA, eicosapentaenoic acid (EPA, 20:5n‐3), docosahexaenoic acid (DHA, 22:6n‐3), and arachidonic acid (ARA, 20:4n‐6) decreased by c. 60% between late summer and the spawning period in early winter. After spawning, total FA content of whitefish muscle increased rapidly, reaching the maximum recorded level in mid‐summer.
Intra‐annual changes in whitefish muscle FA profiles and contents were modified both by available diet and reproductive phase; however, reproductive physiology was clearly a stronger driver of the changes in muscle FA composition. Results suggest marked changes in intra‐annual FA composition of fish muscle, an important factor that should be considered in future studies and especially in long term monitoring programs. Future studies are needed to determine whether these inter‐annual FA patterns revealed in this study can be extended to different regions and to e.g. adipose or spring spawning species.
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BFBNIB, FZAB, GIS, IJS, KILJ, NLZOH, NUK, OILJ, SAZU, SBCE, SBMB, UL, UM, UPUK
Climate change in the Arctic is outpacing the global average and land‐use is intensifying due to exploitation of previously inaccessible or unprofitable natural resources. A comprehensive ...understanding of how the joint effects of changing climate and productivity modify lake food web structure, biomass, trophic pyramid shape and abundance of physiologically essential biomolecules (omega‐3 fatty acids) in the biotic community is lacking. We conducted a space‐for‐time study in 20 subarctic lakes spanning a climatic (+3.2°C and precipitation: +30%) and chemical (dissolved organic carbon: +10 mg/L, total phosphorus: +45 µg/L and total nitrogen: +1,000 µg/L) gradient to test how temperature and productivity jointly affect the structure, biomass and community fatty acid content (eicosapentaenoic acid EPA and docosahexaenoic acid DHA) of whole food webs. Increasing temperature and productivity shifted lake communities towards dominance of warmer, murky‐water‐adapted taxa, with a general increase in the biomass of primary producers, and secondary and tertiary consumers, while primary invertebrate consumers did not show equally clear trends. This process altered various trophic pyramid structures towards an hour glass shape in the warmest and most productive lakes. Increasing temperature and productivity had negative fatty acid content trends (mg EPA + DHA/g dry weight) in primary producers and primary consumers, but not in secondary nor tertiary fish consumers. The massive biomass increment of fish led to increasing areal fatty acid content (kg EPA + DHA/ha) towards increasingly warmer, more productive lakes, but there were no significant trends in other trophic levels. Increasing temperature and productivity are shifting subarctic lake communities towards systems characterized by increasing dominance of cyanobacteria and cyprinid fish, although decreasing quality in terms of EPA + DHA content was observed only in phytoplankton, zooplankton and profundal benthos.
Joint impacts of climate and productivity on food webs biomass and quality components have remained understudied. Biomass and essential fatty acid content (eicosapentaenoic acid EPA and docosahexaenoic acid DHA) in food webs of 20 lakes along climatic and productivity gradient of subarctic Fennoscandia were collected. Increasing temperature and productivity shifted communities towards food webs characterised by increasing biomass dominance of cyanobacteria and cyprinid fish. This process altered various trophic pyramid structures into hourglass shape in the warmest and most productive lakes. Decreasing EPA + DHA content was observed at two lowest trophic levels, but not in next levels composing of fish. Collectively, our results showed that the increasing temperature and productivity promoted both biomass and fatty acid areal content in lake food webs.
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BFBNIB, FZAB, GIS, IJS, KILJ, NLZOH, NUK, OILJ, SAZU, SBCE, SBMB, UL, UM, UPUK
The European whitefish (Coregonus lavaretus) species complex is a classic example of recent adaptive radiation. Here, we examine a whitefish population introduced to northern Finnish Lake Tsahkal in ...the late 1960s, where three divergent morphs (viz. littoral, pelagic, and profundal feeders) were found 10 generations after. Using demographic modeling based on genomic data, we show that whitefish morphs evolved during a phase of strict isolation, refuting a rapid sympatric divergence scenario. The lake is now an artificial hybrid zone between morphs originated in allopatry. Despite their current syntopy, clear genetic differentiation remains between two of the three morphs. Using admixture mapping, we identify five SNPs associated with gonad weight variation, a proxy for sexual maturity and spawning time. We suggest that ecological adaptations in spawning time evolved in allopatry are currently maintaining partial reproductive isolation in the absence of other barriers to gene flow.
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BFBNIB, FZAB, GIS, IJS, KILJ, NLZOH, NUK, OILJ, SAZU, SBCE, SBMB, UL, UM, UPUK
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