Multi-decadal trends in biomarkers in harp seal teeth from the North Atlantic reveal the influence of prey availability on seal trophic position

Arctic food webs are being impacted by borealisation and environmental change. To quantify the impact of these multiple forcings, it is crucial to accurately determine the temporal change in key ecosystem metrics, such as trophic position of top predators. Here, we measured stable nitrogen isotopes (δ¹⁵N) in amino acids in harp seal teeth from across the North Atlantic spanning a period of 60 years to robustly assess multi-decadal trends in harp seal trophic position, accounting for changes in δ¹⁵N at the base of the food web. We reveal long-term variations in trophic position of harp seals which are likely to reflect fluctuations in prey availability, specifically fish- or invertebrate-dominated diets. We show that the temporal trends in harp seal trophic position differ between the Northwest Atlantic, Greenland Sea and Barents Sea, suggesting divergent changes in each local ecosystem. Our results provide invaluable data for population dynamic and ecotoxicology studies.     

Mixed interactions among life history stages of two harvested related species

Climate change and harvesting can affect the ecosystems'' functioning by altering the population dynamics and interactions among species. Knowing how species interact is essential for better understanding potentially unintended consequences of harvest on multiple species in ecosystems. I analyzed how stage‐specific interactions between two harvested competitors, the haddock (Melanogrammus aeglefinus) and Atlantic cod (Gadus morhua), living in the Barents Sea affect the outcome of changes in the harvest of the two species. Using state‐space models that account for observation errors and stochasticity in the population dynamics, I run different harvesting scenarios and track population‐level responses of both species. The increasing temperature elevated the number of larvae of haddock but did not significantly influence the older age‐classes. The nature of the interactions between both species shifted from predator‐prey to competition around age‐2 to ‐3. Increased cod fishing mortality, which led to decreasing abundance of cod, was associated with an increasing overall abundance of haddock, which suggests compensatory dynamics of both species. From a stage‐specific approach, I show that a change in the abundance in one species may propagate to other species, threatening the exploited species'' recovery. Thus, this study demonstrates that considering interactions among life history stages of harvested species is essential to enhance species'' co‐existence in harvested ecosystems. The approach developed in this study steps forward the analyses of effects of harvest and climate in multi‐species systems by considering the comprehension of complex ecological processes to facilitate the sustainable use of natural resources.     

Ancient DNA reveals a southern presence of the Northeast Arctic cod during the Holocene

Climate change has been implicated in an increased number of distributional shifts of marine species during the last century. Nonetheless, it is unclear whether earlier climatic fluctuations had similar impacts. We use ancient DNA to investigate the long-term spawning distribution of the Northeast Arctic cod (skrei) which performs yearly migrations from the Barents Sea towards spawning grounds along the Norwegian coast. The distribution of these spawning grounds has shifted northwards during the last century, which is thought to be associated with food availability and warming temperatures. We genetically identify skrei specimens from Ruskeneset in west Norway, an archaeological site located south of their current spawning range. Remarkably, ¹⁴C analyses date these specimens to the late Holocene, when temperatures were warmer than present-day conditions. Our results either suggest that temperature is not the only driver influencing the spawning distribution of Atlantic cod, or could be indicative of uncertainty in palaeoclimate reconstructions in this region. Regardless, our findings highlight the utility of aDNA to reconstruct the historical distribution of economically important fish populations and reveal the complexity of long-term ecological interactions in the marine environment.     

Integrating omics to characterize eco‐physiological adaptations: How moose diet and metabolism differ across biogeographic zones

1. With accelerated land conversion and global heating at northern latitudes, it becomes crucial to understand, how life histories of animals in extreme environments adapt to these changes. Animals may either adapt by adjusting foraging behavior or through physiological responses, including adjusting their energy metabolism or both. Until now, it has been difficult to study such adaptations in free‐ranging animals due to methodological constraints that prevent extensive spatiotemporal coverage of ecological and physiological data.
2. Through a novel approach of combining DNA‐metabarcoding and nuclear magnetic resonance (NMR)‐based metabolomics, we aim to elucidate the links between diets and metabolism in Scandinavian moose Alces alces over three biogeographic zones using a unique dataset of 265 marked individuals.
3. Based on 17 diet items, we identified four different classes of diet types that match browse species availability in respective ecoregions in northern Sweden. Individuals in the boreal zone consumed predominantly pine and had the least diverse diets, while individuals with highest diet diversity occurred in the coastal areas. Males exhibited lower average diet diversity than females.
4. We identified several molecular markers indicating metabolic constraints linked to diet constraints in terms of food availability during winter. While animals consuming pine had higher lipid, phospocholine, and glycerophosphocholine concentrations in their serum than other diet types, birch‐ and willow/aspen‐rich diets exhibit elevated concentrations of several amino acids. The individuals with highest diet diversity had increased levels of ketone bodies, indicating extensive periods of starvation for these individuals.
5. Our results show how the adaptive capacity of moose at the eco‐physiological level varies over a large eco‐geographic scale and how it responds to land use pressures. In light of extensive ongoing climate and land use changes, these findings pave the way for future scenario building for animal adaptive capacity.

     

Stock collapse and its effect on species interactions: Cod and herring in the Norwegian‐Barents Seas system as an example

Both the Norwegian Spring Spawning herring (Clupea harengus) and the Northeast Arctic (NEA) cod (Gadus morhua) are examples of strong stock reduction and decline of the associated fisheries due to overfishing followed by a recovery. Cod and herring are both part of the Barents Sea ecosystem, which has experienced major warming events in the early (1920–1940) and late 20th century. While the collapse or near collapse of these stocks seems to be linked to an instability created by overfishing and climate, the difference of population dynamics before and after is not fully understood. In particular, it is unclear how the changes in population dynamics before and after the collapses are associated with biotic interactions. The combination of the availability of unique long‐term time series for herring and cod makes it a well‐suited study system to investigate the effects of collapse. We examine how species interactions may differently affect the herring and cod population dynamic before and after a collapse. Particularly we explore, using a GAM modeling approach, how herring could affect cod and vice versa. We found that the effect of cod biomass on herring that was generally positive (i.e., covariation) but the effect became negative after the collapse (i.e., predation or competition). Likewise a change occurred for the cod, the juvenile herring biomass that had no effect before the collapse had a negative effect after. Our results indicate that the population collapses may alter the inter‐specific interactions and response to abiotic environmental changes. While the stocks are at similar abundance levels before and after the collapses, the system is potentially different in its functioning and may require different management action.     

Mitochondrial DNA polymorphism of atlantic cod of the Barents and White seas

High level of intraspecific polymorphism of the mtDNA cytochrome b gene in the Atlantic cod Gadus morhua L. from the northeastern part of the species range (Barents and White seas) is detected. The absence of genetic differentiation between the samples from different areas of the Norwegian and Barents seas is shown, and the fact that these samples belong to the same population of Northeast Arctic cod is confirmed. Significant differences in the frequences of mitochondrial DNA haplotypes between the cod from the Barents and White seas, as well as from the northwestern part of the range, are found. The cod Gadus morhua kildinensis Derjugin, 1920 from Mogilnoye Lake and Gadus morhua marisalbi Derjugin, 1920 from the White Sea were revealed to have common haplotypes with the cod from the Barents Sea, confirming the recent origin of these forms from the Atlantic cod G. morhua.     

Temporal patterns in the social network of core units in Rwenzori Angolan colobus monkeys: Effects of food availability and interunit dispersal

1. Multi‐level societies are complex, nested social systems where basic social groups (i.e., core units) associate in a hierarchical manner, allowing animals to adjust their group sizes in response to variables such as food availability, predation, or conspecific threat. These pressures fluctuate over time and examining the extent to which this variation affects the clustering of core units into different tiers may be instrumental in understanding the evolution of multi‐level societies.
2. The goal of our study was to determine the degree of temporal variability in interunit associations in a multi‐level society of Rwenzori Angolan colobus monkey (Colobus angolensis ruwenzorii), and to determine the social and ecological factors that underlie association patterns. The C. a. ruwenzorii multi‐level society consists of at least three tiers, with core units clustering into clans that share a home range in a band tier.
3. We performed social network analyses on 21 months of association data from 13 core units (totaling 139 identifiable individuals) at Lake Nabugabo, Uganda. We described the patterns of variation in core‐unit associations over time and investigated how changes in rainfall, food availability, and interunit dispersals were correlated with these associations over the short‐term (month to month) and long‐term (year to year).
4. Although clans were relatively stable, larger‐scale changes in association patterns included the formation of an all‐male unit and the transfer of one core unit between clans (within the band tier). Seasonally, core units associated significantly more when fruit, their preferred food source, was abundant (i.e., social networks were denser and more clustered) and there was no direct effect of rainfall seasonality or young leaf availability. Male dispersals also occurred more during periods of high fruit availability, suggesting that greater band cohesion allowed males to prospect and transfer between core units. Once males transferred, their previous and new units associated significantly more with one another than with other core units for 1–2 months postdispersal. The dispersal of five males from one core unit to another in a different clan co‐occurred with this core unit switching its clan affiliation.
5. By examining temporal shifts in social network structure among core units, this study shows the interconnected roles that food availability and dispersal have in shaping the C. a. ruwenzorii multi‐level social system. Our findings highlight how ecological conditions can drive association patterns, impact interunit relationships, and influence social organization.

     

Diets and body condition of polar cod (<Emphasis Type="Italic">Boreogadus saida</Emphasis>) in the northern Bering Sea and Chukchi Sea

To understand trophic responses of polar cod Boreogadus saida (a key species in Arctic food webs) to changes in zooplankton and benthic invertebrate communities (prey), we compared its stomach contents and body condition between three regions with different environments: the northern Bering Sea (NB), southern Chukchi Sea (SC), and central Chukchi Sea (CC). Polar cod were sampled using a bottom trawl, and their potential prey species in the environment were sampled using a plankton net and a surface sediment sampler. Polar cod fed mainly on appendicularians in the NB and SC where copepods were the most abundant in the environment, while they fed on copepods, euphausiids, and gammarids in the CC where barnacle larvae were the most abundant species in plankton samples on average. The stomach fullness index of polar cod was higher in the NB and SC than CC, while their body condition index did not differ between these regions. The lower lipid content of appendicularians compared to other prey species is the most plausible explanation for this inconsistency.     

Forecasting future recruitment success for Atlantic cod in the warming and acidifying Barents Sea

Productivity of marine fish stocks is known to be affected by environmental and ecological drivers, and global climate change is anticipated to alter recruitment success of many stocks. While the direct effects of environmental drivers on fish early life stage survival can be quantified experimentally, indirect effects in marine ecosystems and the role of adaptation are still highly uncertain. We developed an integrative model for the effects of ocean warming and acidification on the early life stages of Atlantic cod in the Barents Sea, termed SCREI (Simulator of Cod Recruitment under Environmental Influences). Experimental results on temperature and CO₂ effects on egg fertilization, egg and larval survival and development times are incorporated. Calibration using empirical time series of egg production, temperature, food and predator abundance reproduces age‐0 recruitment over three decades. We project trajectories of recruitment success under different scenarios and quantify confidence limits based on variation in experiments. A publicly accessible web version of the SCREI model can be run under www.oceanchange.uni-bremen.de/ ;SCREI. Severe reductions in average age‐0 recruitment success of Barents Sea cod are projected under uncompensated warming and acidification toward the middle to end of this century. Although high population stochasticity was found, considerable rates of evolutionary adaptation to acidification and shifts in organismal thermal windows would be needed to buffer impacts on recruitment. While increases in food availability may mitigate short‐term impacts, an increase in egg production achieved by stock management could provide more long‐term safety for cod recruitment success. The SCREI model provides a novel integration of multiple driver effects in different life stages and enables an estimation of uncertainty associated with interindividual and ecological variation. The model thus helps to advance toward an improved empirical foundation for quantifying climate change impacts on marine fish recruitment, relevant for ecosystem‐based assessments of marine systems under climate change.     

Climate-Driven Ichthyoplankton Drift Model Predicts Growth of Top Predator Young

Climate variability influences seabird population dynamics in several ways including access to prey near colonies during the critical chick-rearing period. This study addresses breeding success in a Barents Sea colony of common guillemots Uria aalge where trophic conditions vary according to changes in the northward transport of warm Atlantic Water. A drift model was used to simulate interannual variations in transport of cod Gadus morhua larvae along the Norwegian coast towards their nursery grounds in the Barents Sea. The results showed that the arrival of cod larvae from southern spawning grounds had a major effect on the size of common guillemot chicks at fledging. Furthermore, the fraction of larvae from the south was positively correlated to the inflow of Atlantic Water into the Barents Sea thus clearly demonstrating the mechanisms by which climate-driven bottom-up processes influence interannual variations in reproductive success in a marine top predator.     

Feeding habits of harp seals (Phoca groenlandica) during early summer and autumn in the northern Barents Sea

The feeding habits of harp seals (Phoca groenlandica) in the Barents Sea were examined in studies conducted during June 1991, September 1990 and 1991, and October 1992. Analyses of stomach and intestinal contents were carried out and concurrent estimates of prey abundance were made using trawl gear. Harp seals appeared to feed at low intensity in the pack ice belt during the first half of June. There was little potential prey in the water column, but prawns (Pandalus borealis), capelin (Mallotus villosus) and polar cod (Boreogadus saida) were abundant close to the bottom. In September, the seals sampled in the northern pack ice areas of the Barents Sea fed on the pelagic amphipod Parathemisto libellula, krill (Thysanoessa spp.), prawns and, to a lesser extent, on fish species such as polar cod, sculpins (Cottidae) and snailfish (Liparidae). Trawling revealed that large quantities of Parathemisto libellala were present in the upper layers of the water column. Fish, mainly capelin and polar cod, were less abundant and occurred in deeper waters. In mid-October, the diet of seals in the northern Barents Sea consisted mainly of amphipods (Parathemisto sp.). Later in October, when increasing pack ice cover forced the harp seals to move south, the diet seemed to change from amphipods to fish prey, predominantly capelin and polar cod.     

Distribution and diet of 0-group cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) and haddock (<Emphasis Type="Italic">Melanogrammus aeglefinus</Emphasis>) in the Barents Sea in relation to food availability and temperature

Distribution of 0-group cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) in August–September 2005 and 2006 was mainly restricted to the Atlantic waters of the western and central areas of the Barents Sea. The main distribution of 0-group fish overlapped largely with areas of high biomass (>7 gm⁻² dry weight) of zooplankton. The copepod Calanus finmarchicus and krill Thysanoessa inermis, which are dominant zooplankton species in both Atlantic and boreal waters of the Barents Sea, were the main prey of 0-group cod and haddock. The main distribution, feeding areas and prey of 0-group cod and haddock overlapped, implying that competition for food may occur between the two species. However, though their diet coincided to a certain degree, haddock seems to prefer smaller and less mobile prey, such as Limacina and appendicularians. As 0-group fish increased in size, there seems to be a shift in diet, from small copepods and towards larger prey such as krill and fish. Overall, a largely pelagic feeding behaviour of 0-group cod and haddock was evident from this study.     

Thermal growth potential of Atlantic cod by the end of the 21st century

Ocean warming may lead to smaller body sizes of marine ectotherms, because metabolic rates increase exponentially with temperature while the capacity of the cardiorespiratory system to match enhanced oxygen demands is limited. Here, we explore the impact of rising sea water temperatures on Atlantic cod (Gadus morhua), an economically important fish species. We focus on changes in the temperature‐dependent growth potential by a transfer function model combining growth observations with climate model ensemble temperatures. Growth potential is expressed in terms of asymptotic body weight and depends on water temperature. We consider changes between the periods 1985–2004 and 2081–2100, assuming that future sea water temperatures will evolve according to climate projections for IPCC AR5 scenario RCP8.5. Our model projects a response of Atlantic cod to future warming, differentiated according to ocean regions, leading to increases of asymptotic weight in the Barents Sea, while weights are projected to decline at the southern margin of the biogeographic range. Southern spawning areas will disappear due to thermal limitation of spawning stages. These projections match the currently observed biogeographic shifts and the temperature‐ and oxygen‐dependent decline in routine aerobic scope at southern distribution limits.     

Climate effects on size‐at‐age: growth in warming waters compensates for earlier maturity in an exploited marine fish

Over the past 3 decades, North Sea Atlantic cod (Gadus morhua) have exhibited variable length‐at‐age along with declines in spawning stock biomass and timing of maturity. Multiple factors affecting growth and development in fish acted on this economically important stock over the same period including warming waters and an intensive fishery. Here, we employ North Sea cod as a model population, exploring how a physiologically relevant temperature metric (the growing degree‐day, GDD; °C day) can be used to compare year‐classes on a physiologically relevant time‐scale, disentangling influences of climate (thermal history) on observed length‐at‐age trends. We conclude that the trends in North Sea cod length‐at‐age observed during the last three decades can be explained by a combination of temperature‐dependent growth increases and a trend toward earlier maturation, the latter likely induced by the intensive fishing pressure, and possibly evidence of fisheries‐induced evolution.     

New data on morphometry, feeding, and parasitic fauna of Atlantic spiny lumpsucker Eumicrotremus spinosus (Cyclopteridae) from the Barents Sea

Data on morphometric analysis, feeding, and parasitic fauna of the Atlantic spiny lumpsucker Eumicrotremus spinosus from the Northwestern Barents Sea are given. The amount of food in the whole digestive tract of fish in the evening (18.00–02.00) is high; the weight of food bolus averaged 1/9 of the weight of the fish without guts; the average index of fullness of the stomach was 1018?. The small hyperiids Themisto abyssorum constituted the basis of food; Th. libellula, gammarides of the family Lysianassidae, and Gammarus wilkitzkii were consumed to a lesser degree. The pink shrimp Pandalus borealis and euphausiid Thysanoessa inermis are noted singly; the polychaetes of genus Nereis also serve as occasional food. Parasitic fauna of the Atlantic spiny lumpsucker is represented by the larvae of Anisakis simplex that are located on the liver parenchyma and pyloric appendages. The parasitic copepod Lernaeocera branchialis was found on the gills of one of the females.     

Do Norwegian Atlantic salmon feed in the northern Barents Sea? Tag recoveries from 70 to 78° N

Three tagged Atlantic salmon Salmo salar were recaptured as subadults or adults (1·4–3 kg) between 70·5 and 78° N in the western Barents Sea, two of which originated from the Alta Fjord region in northern Norway and one from the Drammen River, south-eastern Norway. An additional tag was recovered from the stomach of a Greenland halibut Reinhardtius hippoglossoides captured south-west of Bear Island at >600 m depth; this tag was from a smolt released in the River Alta 1 month earlier. These are the northernmost tag recoveries reported for Atlantic salmon, and indicate that Norwegian Atlantic salmon, especially the fish from northern populations, may use the northern Barents Sea as a feeding area during part of their life cycle.     

Disparate movement behavior and feeding ecology in sympatric ecotypes of Atlantic cod

Coexistence of ecotypes, genetically divergent population units, is a widespread phenomenon, potentially affecting ecosystem functioning and local food web stability. In coastal Skagerrak, Atlantic cod (Gadus morhua) occur as two such coexisting ecotypes. We applied a combination of acoustic telemetry, genotyping, and stable isotope analysis to 72 individuals to investigate movement ecology and food niche of putative local “Fjord” and putative oceanic “North Sea” ecotypes—thus named based on previous molecular studies. Genotyping and individual origin assignment suggested 41 individuals were Fjord and 31 were North Sea ecotypes. Both ecotypes were found throughout the fjord. Seven percent of Fjord ecotype individuals left the study system during the study while 42% of North Sea individuals left, potentially homing to natal spawning grounds. Home range sizes were similar for the two ecotypes but highly variable among individuals. Fjord ecotype cod had significantly higher δ¹³C and δ¹⁵N stable isotope values than North Sea ecotype cod, suggesting they exploited different food niches. The results suggest coexisting ecotypes may possess innate differences in feeding and movement ecologies and may thus fill different functional roles in marine ecosystems. This highlights the importance of conserving interconnected populations to ensure stable ecosystem functioning and food web structures.     

A modified niche model for generating food webs with stage‐structured consumers: The stabilizing effects of life‐history stages on complex food webs

1. Almost all organisms grow in size during their lifetime and switch diets, trophic positions, and interacting partners as they grow. Such ontogenetic development introduces life‐history stages and flows of biomass between the stages through growth and reproduction. However, current research on complex food webs rarely considers life‐history stages. The few previously proposed methods do not take full advantage of the existing food web structural models that can produce realistic food web topologies.
2. We extended the niche model developed by Williams and Martinez (Nature, 2000, 404, 180–183) to generate food webs that included trophic species with a life‐history stage structure. Our method aggregated trophic species based on niche overlap to form a life‐history structured population; therefore, it largely preserved the topological structure of food webs generated by the niche model. We applied the theory of allometric predator–prey body mass ratio and parameterized an allometric bioenergetic model augmented with biomass flow between stages via growth and reproduction to study the effects of a stage structure on the stability of food webs.
3. When life‐history stages were linked via growth and reproduction, more food webs persisted, and persisting food webs tended to retain more trophic species. Topological differences between persisting linked and unlinked food webs were small to modest. The slopes of biomass spectra were lower, and weak interaction links were more prevalent in the linked food webs than the unlinked ones, suggesting that a life‐history stage structure promotes characteristics that can enhance stability of complex food webs.
4. Our results suggest a positive relationship between the complexity and stability of complex food webs. A life‐history stage structure in food webs may play important roles in dynamics of and diversity in food webs.

     

Thermal ecology and baseline energetic requirements of a large‐bodied ectotherm suggest resilience to climate change

1. Most studies on how rising temperatures will impact terrestrial ectotherms have focused on single populations or multiple sympatric species. Addressing the thermal and energetic implications of climatic variation on multiple allopatric populations of a species will help us better understand how a species may be impacted by altered climates.
2. We used eight years of thermal and behavioral data collected from four populations of Pacific rattlesnakes (Crotalus oreganus) living in climatically distinct habitat types (inland and coastal) to determine the field‐active and laboratory‐preferred body temperatures, thermoregulatory metrics, and maintenance energetic requirements of snakes from each population.
3. Physical models showed that thermal quality was best at coastal sites, but inland snakes thermoregulated more accurately despite being in more thermally constrained environments. Projected increases of 1 and 2°C in ambient temperature result in an increase in overall thermal quality at both coastal and inland sites.
4. Population differences in modeled standard metabolic rate estimates were driven by body size and not field‐active body temperature, with inland snakes requiring 1.6× more food annually than coastal snakes.
5. All snakes thermoregulated with high accuracy, suggesting that small increases in ambient temperature are unlikely to impact the maintenance energetic requirements of individual snakes and that some species of large‐bodied reptiles may be robust to modest thermal perturbations under conservative climate change predictions.

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Tracking nutrients in space and time: Interactions between grazing lawns and drought drive abundances of tallgrass prairie grasshoppers

1. We contrast the response of arthropod abundance and composition to bison grazing lawns during a drought and non‐drought year, with an emphasis on acridid grasshoppers, an important grassland herbivore.
2. Grazing lawns are grassland areas where regular grazing by mammalian herbivores creates patches of short‐statured, high nutrient vegetation. Grazing lawns are predictable microsites that modify microclimate, plant structure, community composition, and nutrient availability, with likely repercussions for arthropod communities.
3. One year of our study occurred during an extreme drought. Drought mimics some of the effects of mammalian grazers: decreasing above‐ground plant biomass while increasing plant foliar percentage nitrogen.
4. We sampled arthropods and nutrient availability on and nearby (“off”) 10 bison‐grazed grazing lawns in a tallgrass prairie in NE Kansas. Total grasshopper abundance was higher on grazing lawns and the magnitude of this difference increased in the wetter year of 2019 compared to 2018, when drought led to high grass foliar nitrogen concentrations on and off grazing lawns. Mixed‐feeding grasshopper abundances were consistently higher on grazing lawns while grass‐feeder and forb‐feeder abundances were higher on lawns only in 2019, the wetter year. In contrast, the abundance of other arthropods (e.g., Hemiptera, Hymenoptera, and Araneae) did not differ on and off lawns, but increased overall in 2019, relative to the drought of 2018.
5. Understanding these local scale patterns of abundances and community composition improves predictability of arthropod responses to ongoing habitat change.